Launois 1899015133J TH

UNIVERSITE AIX-MARSEILLE I - Université de Provence
ECOLE DOCTORALE DES SCIENCES DE L’ENVIRONNEMENT (251) – CEREGE Aix-

en-Provence
THESE
présentée à
L’UNIVERSITE DE PROVENCE
AIX-MARSEILLE I
Ecole Doctorale
SCIENCES DE L’ENVIRONNEMENT
Pour obtenir le grade de DOCTEUR

Spécialité : ECOLOGIE
par
Lionel LAUNOIS
Impact des facteurs anthropiques sur les

communautés piscicoles lentiques : vers
l’élaboration d’un indice poisson
Soutenue publiquement le 24 juin 2011
Composition du Jury :
Mme. Christine ARGILLIER (Directeur de Recherche, Cemagref) Directeur de thèse
M. Rémi CHAPPAZ (Professeur, Université de Provence)
Mme. Lourdes ENCINA (Professeur, Université de Séville) Rapporteur
M. Didier PONT (Directeur de Recherche, Cemagref)
M. Yorick REYJOL (Chargé de mission, ONEMA – DAST)
M. Frédéric SANTOUL (Maître de conférence, Université de Toulouse) Rapporteur
Travaux réalisés dans l’Equipe Ecosystèmes lacustres

Cemagref d’Aix-en-Provence – Unité de Recherche Hydrobiologie, HYAX
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Remerciements
Cette partie qui me tient particulièrement à cœur me permet de témoigner ma gratitude

aux nombreuses personnes qui, de près ou de loin, ont participé à ce travail et sans le concours
desquelles je n’aurais pu mener à bien ce travail.
Je tiens tout d’abord à remercier Christine Argillier d’avoir accepté la direction de ce

travail et de m’avoir témoigné sa confiance et prodigué ses conseils et ses encouragements
tout au long de cette thèse. Je voudrais également remercier Christine Argillier, Pascal Irz et
Maxime Logez, de leur relecture minutieuse et critique du présent manuscrit, et de leurs
judicieux commentaires.
Je tiens à remercier tout particulièrement, Lourdes Encina (Université de Séville) et
Frédéric Santoul (Université de Toulouse), de m’avoir fait l’honneur d’être les rapporteurs de
ce manuscrit. Merci aussi à Rémi Chappaz (Université d’Aix-Marseille), Yorick Reyjol
(ONEMA – DAST) et Didier Pont (Cemagref d’Antony) d’avoir accepté d’évaluer ce travail
en étant membre de mon jury.
Je tiens également à remercier, Jacques Veslot, Pierre Bady et Muriel Gevrey (Cemagref
d’Aix-en-Provence), pour leur aide précieuse, notamment dans les corrections apportées aux
articles. Un merci tout particulier pour leur gentillesse, leur soutien et leur amitié. Ils m’ont
permis d’avoir des discussions stimulantes, motivantes, parfois houleuses, mais toujours
constructives et pertinentes.
Je remercie également Pascal Kosuth (Cemagref de Montpellier) d’avoir accepté de co-
encadrer ce travail, Philippe Duchène (Cemagref d’Antony) pour m’avoir conseillé dans ma
démarche académique et Yann Le Coarer (Cemagref d’Aix-en-Provence) pour m’avoir
accueilli au sein de l’unité Hydrobiologie du Cemagref d’Aix-en-Provence.
Un grand merci à Anik Brind’Amour (IFREMER), Romuald Berrebi (ONEMA – DAST),
Martin Daufresne (Cemagref d’Aix-en-Provence), Corinne Grac, Caroline Grégoire et Thierry
Leviandier [Ecole Nationale de l’Eau et de l’Environnement de Strasbourg (ENGEES)], qui
m’ont prodigué, tout au long de ce travail, la pertinence de leurs conseils et de leurs
suggestions, ainsi que leur contribution scientifique déterminante dans l’encadrement de ma
thèse.
Je voudrais aussi remercier Carole Heyd, Virgine Raymond et Samuel Alleaume pour leur
aide en matière de télédétection, de SIG et d’analyses spatiales.
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Merci à l’équipe Ecosystèmes lacustres pour les expériences de terrain mémorables. J’ai
apprécié de travailler avec eux en équipe sur le terrain. Merci pour leur aide, leur
collaboration et leur bonne humeur.
Merci enfin à l’ensemble des personnes de l’unité Hydrobiologie du Cemagref d’Aix-en-
Provence pour leur grande disponibilité à mon égard. La qualité des discussions a fortement
contribué à enrichir le contenu de ce mémoire. Un merci tout particulier à Maxime, Renaud,
Jacques, Olivier, Pierre, Jonathan, Marlène, Caroline, Muriel, Carole, Didier, Bernard, Yann,
Georges, Virginie et Jean-Pierre, mes compagnons des pauses café et autres excursions
gastronomiques dans le massif de la Sainte-Victoire, pour les très bons moments passés
ensemble, ainsi que pour les très nombreux fous rires. Un merci également à Isabelle Majka,
secrétaire de compétition de l’Unité Hydrobiologie, que je remercie pour sa bonne humeur et
sa contribution pour tous les à-côtés liés à cette thèse.
Merci à mes parents et à mon frère de m’avoir toujours motivé et fait confiance, à Sophie,
ma compagne, de m’avoir soutenu au quotidien pendant toute cette période, et à mes amis de
m’avoir accompagné tout au long de cette expérience. J’ai particulièrement apprécié la
compréhension, l’optimisme et les encouragements que chacun m’a témoignés.
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Table des matières
Table des matières ..................................................................................................................- 5 -

Résumé ...................................................................................................................................- 9 -
Abstract ................................................................................................................................- 11 -
Liste des tableaux .................................................................................................................- 13 -
Liste des figures ...................................................................................................................- 15 -
Abréviations utilisées ...........................................................................................................- 17 -
CHAPITRE 1 .......................................................................................................................- 19 -
Introduction ..........................................................................................................................- 19 -
1.1. Les milieux lentiques, fonctions et enjeux ...........................................................- 20 -
1.2. Les poissons en bioindication lacustre .................................................................- 21 -
1.3. Objectifs de l’étude et organisation générale du mémoire ...................................- 23 -
CHAPITRE 2 .......................................................................................................................- 27 -
Données disponibles à l’échelle de la France.......................................................................- 27 -
2.1. L’ichtyofaune .......................................................................................................- 28 -
2.1.1. Les inventaires piscicoles.............................................................................- 28 -
2.1.2. Les caractéristiques biologiques des espèces ...............................................- 29 -
2.2. Les caractéristiques environnementales naturelles des lacs et de leurs bassins
versants.............................................................................................................................- 30 -
2.3. Les facteurs anthropiques.....................................................................................- 31 -
2.3.1. Les pressions anthropiques à l’échelle du bassin versant ............................- 32 -
2.3.2. Les pressions anthropiques locales ..............................................................- 32 -
CHAPITRE 3 .......................................................................................................................- 35 -
Sélection de métriques piscicoles par une approche intuitive : une première démarche basée
sur une typologie physique des plans d’eau .........................................................................- 35 -
3.1. Introduction ..........................................................................................................- 38 -
3.2. Matériel et méthodes ............................................................................................- 39 -
3.2.1. Réalisation de la typologie lacustre..............................................................- 39 -
3.2.2. Analyse de la réponse de l’ichtyofaune aux pressions dans chaque type ....- 41 -
3.2.2.1. Les données poissons ...............................................................................- 41 -
3.2.2.2. Les données de pressions .........................................................................- 45 -
3.2.2.3. Réponse des métriques aux pressions ......................................................- 46 -
3.3. Résultats ...............................................................................................................- 47 -
3.3.1. Typologie physique des plans d’eau français...............................................- 47 -
3.3.2. Intensité des pressions ..................................................................................- 50 -
3.3.2.1. Analyse globale ........................................................................................- 50 -
3.3.2.2. Analyse par types .....................................................................................- 52 -
3.3.3. Calcul des métriques piscicoles et réponse aux pressions............................- 53 -
3.4. Discussion et conclusions.....................................................................................- 57 -
CHAPITRE 4 .......................................................................................................................- 63 -
Sélection de métriques piscicoles et variabilité environnementale : comparaison entre lacs
naturels et réservoirs artificiels.............................................................................................- 63 -
4.1. Introduction ..........................................................................................................- 66 -
4.2. Study Sites............................................................................................................- 67 -
4.3. Material and methods ...........................................................................................- 69 -
4.4. Results ..................................................................................................................- 78 -
4.5. Discussion ............................................................................................................- 81 -
4.6. Conclusion............................................................................................................- 85 -
CHAPITRE 5 .......................................................................................................................- 87 -
Développement d’un indice poisson pour évaluer l’état écologique des plans d’eau en France :
une approche par hindcasting...............................................................................................- 87 -
5.1. Introduction ..........................................................................................................- 91 -
5.2. Methods ................................................................................................................- 92 -
5.2.1. Study sites ....................................................................................................- 92 -
5.2.2. Abiotic environmental characteristics of lakes ............................................- 94 -
5.2.3. Fish sampling procedures.............................................................................- 95 -
5.2.4. Candidate metrics .........................................................................................- 96 -
5.2.5. Catchment-scale anthropogenic pressures ...................................................- 97 -
5.2.6. Local-scale anthropogenic pressures............................................................- 98 -
5.2.7. Statistical approach ......................................................................................- 98 -
5.2.7.1. Metrics selection ......................................................................................- 99 -
5.2.7.2. Hindcasting approach ...............................................................................- 99 -
5.2.7.3. Aggregation of metrics into total scores (FBIs) .....................................- 102 -
5.2.7.4. FBI evaluation ........................................................................................- 102 -
5.3. Results ................................................................................................................- 102 -
5.3.1. Study sites, environmental variables and anthropogenic pressures ...........- 102 -
5.3.2. Fish assemblages ........................................................................................- 103 -
5.3.3. Metrics selection ........................................................................................- 103 -
5.3.4. Hindcast modeling and normalized deviation transformation ...................- 105 -
5.3.5. FBI evaluation ............................................................................................- 106 -
5.4. Discussion ..........................................................................................................- 108 -
5.4.1. Metrics selected for the FBI .......................................................................- 108 -
5.4.2. Evaluation of the structure of the FBI ........................................................- 110 -
5.4.3. Analytical procedure issues and limitations of hindcast models................- 111 -
5.5. Conclusion..........................................................................................................- 113 -
CHAPITRE 6 .....................................................................................................................- 115 -
Conclusions et perspectives ...............................................................................................- 115 -
Références bibliographiques ..............................................................................................- 123 -
ANNEXES .........................................................................................................................- 155 -
Annexe 1 : Caractérisation des pressions anthropiques s’exerçant à l’échelle régionale - 156
-
1. Introduction ........................................................................................................- 156 -
2. Impacts of chemical pressures on lake biotic communities (Tableau 6. 1)........- 157 -
3. Impacts of morphological pressures on lake biotic communities (Tableau 6. 2).- 167
-
4. Impacts of hydrological pressures on lake biotic communities (Figure 6. 3, Tableau
6. 3) - 174 -
5. Impacts of species manipulation on lake biotic communities (Figure 6. 4, Tableau 6.
4) - 179 -
6. Conclusion and perspectives for biological monitoring.....................................- 186 -
Annexe 2 : Liste des 138 plans d’eau français pour lesquels des données environnementales
naturelles et des pressions anthropiques sont disponibles..............................................- 191 -
Annexe 3 : Classification des 42 espèces pisicoles en guildes de reproduction, trophiques et
de tolérance. ...................................................................................................................- 204 -
Annexe 4 : Classification des 41 espèces piscicoles en guildes de reproduction, trophiques
et de tolérance.................................................................................................................- 206 -
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Annexe 5 : Caractéristiques des 41 espèces de poissons capturées lors d’une campagne
d’échantillonnage par plan d’eau, effectuée selon un protocole d’échantillonnage
standardisé (C.E.N. 2005). .............................................................................................- 208 -
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Résumé
L’ichtyofaune est encore peu utilisée en bioindication sur les plans d’eau, alors que la
demande des gestionnaires est forte, notamment dans le contexte de la mise en œuvre de la
Directive Cadre Européenne sur l’Eau. L’objectif de cette thèse est d’analyser la réponse de
l’ichtyofaune à diverses pressions anthropiques, afin de définir des variables biologiques de
bioindication et construire un indice rendant compte de l’état écologique des lacs naturels et
des retenues français. Plusieurs questions sont successivement abordées.
En première analyse, les possibilités de sélection de métriques répondant à des pressions
s’exerçant sur le bassin versant sont explorées par une approche typologique. L’analyse
consiste à effectuer des classifications des plans d’eau, puis à établir, au sein de chaque
groupe homogène d’un point de vue environnemental, des régressions linéaires entre les
métriques candidates et les pressions. Les résultats montrent la pertinence de l’ichtyofaune en
bioindication sur ces milieux, mais mettent en exergue des limites à cette approche
typologique.
Ainsi, afin de tester la possibilité de construction d’un bioindicateur commun aux lacs
naturels et aux retenues, une méthode alternative dite « site spécifique » a ensuite été
développée. Une modélisation statistique a été appliquée pour contrôler l’effet de
l’environnement naturel sur les communautés piscicoles lacustres. Cette analyse montre que
les métriques piscicoles répondant aux pressions sont différentes entre les deux types de
milieux.
La dernière étape, explique le développement d’un indicateur, construit sur le principe de
la mesure d’un écart à la référence, en se basant sur une approche de type statistique par
hindcasting. Cette méthode encore peu utilisée permet de s’affranchir de la sélection de sites
de référence en France en modélisant les conditions biologiques de référence, i.e., telles
qu’elles seraient en l’absence de pression anthropique. Les caractéristiques des communautés
piscicoles des sites non perturbés sont ensuite comparées à celles des sites plus ou moins
impactés par des pressions anthropiques. Ont été considérées ici les pressions s’exerçant à
l’échelle du bassin versant et localement sur les plans d’eau. Suivant ce protocole, un indice
poisson lacustre issu de la combinaison de métriques répondant conjointement à des pressions
anthropiques globales et locales est proposé pour les lacs naturels et pour les retenues.
Les avancées de ce travail et les perspectives sont ensuite discutées au regard du contexte
européen de la gestion des milieux aquatiques.
Mots-clés : bioindication ▪ communauté piscicole ▪ lac naturel ▪ retenue artificielle ▪ facteurs
environnementaux ▪ pressions anthropiques ▪ échelles spatiales ▪ traits d’histoire de vie ▪
Directive Cadre Européenne sur l’eau.
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Abstract
Fish communities remain underused in the domain of lentic ecosystem bioindication, even
though water managers need such tools in the current context of the Water Directive
Framework. The objective of this thesis was to analyse the fish communities’ response to
various anthropogenic pressures, in order to define the candidate biological variables in
bioindication and to develop an index to evaluate the ecological status of French natural lakes
and reservoirs. Several questions are raised.
First of all, the means available to select fish-based metrics responding to catchment-scale
anthropogenic pressures are analysed using a typological approach. This analysis consists in
classifying lentic ecosystems using classifications, and then, within each homogenous group
of ecosystems identified based on environmental criteria, performing linear regressions
between candidate fish-based metrics and anthropogenic pressures. The results clearly show
that fish communities could be used as good bioindicators for the lentic ecosystems, but they
also underline the limits of typological approaches.
To test how a bioindicator shared by natural lakes and reservoirs could be developed, an
alternative site-specific method was subsequently developed. Statistical modeling was applied
to control the natural environmental effects on lentic fish communities. This analysis shows
that fish-based metrics responding to catchment-scale anthropogenic pressures differ in the
two types of lentic ecosystems.
The last part reports the development of an index, based on the reference condition
approach, using a hindcasting statistical approach. This method, still infrequently used,
enables one to sidestep the problem related to the selection of reference sites in France.
Furthermore, this method allows modeling biological reference conditions, i.e. conditions
without anthropogenic pressures. Fish communities’ characteristics are then compared for
both undisturbed sites and sites that are more or less impacted by anthropogenic pressures.
Both catchment-scale and local anthropogenic pressures were considered in this analysis.
Following this protocol, a lake fish-based index resulting from the combination of metrics that
respond jointly to global and local anthropogenic pressures was developed for natural lakes
and reservoirs.
Finally, the advances made as a result of this research and its perspectives are discussed
with regards to the European context of aquatic ecosystem management.
- 11 -
Key-words: bioindication ▪ fish community ▪ natural lake ▪ reservoir ▪ environmental factors ▪
anthropogenic pressures ▪ spatial scales ▪ life history traits ▪ Water Framework Directive.
- 12 -
Liste des tableaux
Tableau 2. 1 : Critères de classification des espèces piscicoles en guildes fonctionnelles. .- 30 -
Tableau 3. 1: Valeur moyenne et écart type des principales caractéristiques environnementales

des plans d’eau appartenant aux cinq types définis par l’analyse (pour les types, voir
« 1.3.1 Typologie physique des plans d’eau français »). .............................................- 41 -
Tableau 3. 2: Caractéristiques des 29 espèces de poissons capturées dans 16 plans d’eau du
groupe LN1 et 10 du groupe RA3, lors d’une campagne d’échantillonnage par plan d’eau
effectuée entre 2005 et 2009, selon un protocole d’échantillonnage standardisé (C.E.N
2005).............................................................................................................................- 43 -
Tableau 3. 3: Liste des 14 métriques testées pour l’analyse des réponses de l’ichtyofaune de 2
types de plans d’eau aux pressions...............................................................................- 45 -
Tableau 3. 4: Corrélation de Spearman entre les métriques et les pressions au sein de chaque
type de plans d’eau : * significatif à p<0,05. ...............................................................- 54 -
Tableau 4. 1: Environmental parameter mean values and extreme values of the 89 study sites.-
68 -
Tableau 4. 2: Description of the anthropogenic variables included in the analysis; and
principal component analysis (PCA) carried out on the watershed pressure variables..- 71
-
Tableau 4. 3: Occurrence rates of the 42 fish species in natural lakes and reservoirs. ........- 73 -
Tableau 4. 4: Proposed guild classification for French lake fishes. .....................................- 74 -
Tableau 4. 5: Mean values (standard deviation) of the fish-based metrics and p-values of
Student t-test (with Welch correction if needed) obtained from the functional traits
between natural lakes and reservoirs; P ≤ 0.05*, P ≤ 0.01**, P ≤ 0.001***. ..............- 76 -
Tableau 4. 6: Regression of fish metrics versus the environmental and anthropogenic
variables. ......................................................................................................................- 80 -
Tableau 5. 1: Left: Principal component analyses (PCAs) carried out separately on the
watershed agricultural and urban pressure variables (AP and UP respectively). Table
entries are the percentage of variance (inertia) explained by each first principal
component (PC) and the correlations between the variables and the first PCs. Those
loading most heavily are in bold. Right: descriptive statistics of the anthropogenic
variables included in the analysis.................................................................................- 94 -
Tableau 5. 2: Environmental parameters, mean and extreme values for the 91 study sites.- 95 -
Tableau 5. 3: Guild classification for French lake fishes. ....................................................- 97 -
Tableau 5. 4: Example calculations for hindcasting approach using the model for log(B_OTol
+1) at Angle Guignard (ANG85, reservoir site). Adapted from Kilgour and Stanfield
(2006). ........................................................................................................................- 100 -
Tableau 5. 5: Pearson correlation coefficients between selected normalized deviations; P ≤
0.05*. P ≤ 0.01**. P ≤ 0.001***................................................................................- 101 -
variables. ....................................................................................................................- 104 -
- 13 -
- 14 -
Liste des figures
Figure 2. 1 : Localisation des sites d’étude en France et délimitation des principales zones
hydrographiques. ..........................................................................................................- 29 -
Figure 3. 1 : Localisation géographique des plans d’eau étudiés (1a) pour la réalisation de la
typologie et (1b) pour l’analyse de la réponse des métriques aux pressions................- 40 -
Figure 3. 2 : Illustration de la méthode K-means, avec le partitionnement des données en 3
groupes, ou « clusters ». ...............................................................................................- 41 -
Figure 3. 3: Identification des deux groupes de lacs naturels (1 : groupe LN1 ; 2 : groupe
LN2) par la méthode des K-means – analyse sur 34 lacs naturels. ..............................- 47 -
Figure 3. 4: Identification des trois groupes de retenues (1 : groupe RA1 ; 2 : groupe RA2 ; 3 :
groupe RA3) par la méthode des K-means – analyse sur 104 sites. ............................- 48 -
Figure 3. 5: Représentation des principales caractéristiques environnementales naturelles des
cinq types de plans d’eau identifiés..............................................................................- 49 -
Figure 3. 6: Analyse en Composantes Principales (ACP) réalisée sur les neuf variables de
pressions anthropiques. ................................................................................................- 50 -
Figure 3. 7: Analyse en Composantes Principales (ACP) réalisée sur les cinq variables de
pressions agricoles (7a), et les quatre variables de pressions urbaines (7b). ...............- 52 -
Figure 3. 8: Représentation des trois indices synthétiques de pressions anthropiques pour
chacun des cinq types de plans d’eau identifiés : indice synthétique des pressions
agricoles (8a), indice synthétique des pressions urbaines (8b), indice de pression global
(8c). ..............................................................................................................................- 53 -
Figure 3. 9: Représentations graphiques des corrélations de Spearman (Rsp : valeur du
coefficient de corrélation) entre les métriques sélectionnées dans le type RA3 des
retenues et les indices synthétiques de pressions. ........................................................- 55 -
coefficient de corrélation) entre les métriques sélectionnées dans le type LN1 des lacs
naturels et les indices synthétiques de pressions. .........................................................- 56 -
Figure 4. 1 : Partitionnement de la variance d’une métrique biologique en quatre

composantes : [a] fraction exclusivement liée aux pressions anthropiques, [b] fraction
conjointe entre pressions anthropiques et environnement, [c] fraction exclusivement liée
à l’environnement et [d] fraction inexpliquée. Adapté de Legendre & Legendre (1998)...-
65 -
Figure 4. 2: Location of the 89 study sites in the French hydrographic network.................- 68 -
Figure 5. 1: Location of the 91 study sites in France. ..........................................................- 93 -

Figure 5. 2: Boxplot of deviations (i.e., observed metric – expected metric), scaled in terms of
standard deviations for (a) reservoirs and (b) natural lakes in France. ......................- 106 -
Figure 5. 3: Relationships between: (a) FBI and agricultural pressure for reservoirs (n=67); (b)
FBI and LHMS for natural lakes (n=24); and (c) FBI and urban pressure for natural lakes
(n=24). ........................................................................................................................- 108 -
Figure 6. 1: Relationships between chemical pressures and lake biological communities..- 160
-
Figure 6. 2: Relationships between morphological pressures and lake biota.....................- 169 -
Figure 6. 3: Relationships between hydrological pressures and lake biological communities
(solid line: direct effects, dotted line: indirect effects)...............................................- 174 -
- 15 -
Figure 6. 4: Relationships between species manipulations and lake biota, especially on native
fish communities (solid line: direct effects, dotted line: indirect effects). .................- 179 -
- 16 -
Abréviations utilisées
ACP : Analyse en Composantes Principales.

BRGM : Bureau de Recherches Géologiques et Minières.
BV : Bassin Versant.
CEN : Comité Européen de Normalisation.
CNRS : Centre National de la Recherche Scientifique.
CWA : Clean Water Act.
DAST : Direction de l’Action Scientifique et Technique.
DCE : Directive Cadre Européenne sur l’eau.
EEA : European Environment Agency.
ENGEES : Ecole Nationale de l’Eau et de l’Environnement de Strasbourg.
EPCN : European Pond Conservation Network.
EU : European Union.
FAO : Food and Agriculture Organization.
FBI : Fish-based Index
IBI : Index of Biotic Integrity.
IFEN : Institut Français de l’ENvironnement.
IFREMER : Institut Français de Recherche pour l’Exploitation de la MER.
IGN : Institut Géographique National.
INERIS : Institut National de l’EnviRonnement Industriel et des riSques.
INRA : Institut National de la Recherche Agronomique.
LHMS : Lake Habitat Modification Score.
LHS : Lake Habitat Survey.
LN (ou N) : Lacs Naturels.
MNT : Modèle Numérique de Terrain.
NTotal : Azote total.
NOAA : National Oceanic and Atmospheric Administration.
ONEMA : Office National de l’Eau et des Milieux Aquatiques.
PTotal : Phosphore total.
RA (ou A) : Réservoirs.
RCA : Reference Condition Approach.
RDA : Redundancy Analysis (Analyses de redondance)
- 17 -
REFCOND : REFerence CONDition.
RFAI : Reservoir Fish Assemblage Index.
SIG : Système d’Information Géographique.
TVA : Tennessee Valley Authority.
US : United States.
USEPA : United States Environmental Protection Agency.
USGS : United States Geological Survey.
WFD : Water Framework Directive.
- 18 -
CHAPITRE 1
Introduction
1.1. Les milieux lentiques, fonctions et enjeux
Les lacs naturels et les plans d’eau d’origine artificielle – tous deux regroupés sous le
terme générique de « milieux lentiques » - représentent plus de 90%, en volume, de l’eau
douce de surface disponible sur Terre (Shiklomanov 1993). En Europe, on dénombrait plus de
500 000 lacs naturels ayant une superficie supérieure à 10 ha (Kristiansen & Hansen 1994),
ainsi que 10 000 retenues artificielles couvrant une surface totale de plus de 100 000 km2
(World Commission on Dams 2000). Ce dernier nombre est probablement bien supérieur
aujourd’hui car de nouveaux plans d’eau sont régulièrement créés pour divers usages tels que
les loisirs, le soutien d’étiage ou bien encore l’irrigation des terres agricoles.
Les milieux lentiques, en particulier les petits plans d’eau d’origine naturelle, abritent le
plus souvent une diversité faunistique et floristique exceptionnelle, fréquemment supérieure à
celles des rivières et des fleuves (Davies et al. 2008, Williams et al. 2004). Ils présentent une
grande diversité morphologique (Maitland 1995) et forment une véritable mosaïque d’habitats
hétérogènes, peuplés par un grand nombre d’espèces menacées (Baillie et al. 2004). Outre ce
rôle de réservoirs biologiques, les plans d’eau remplissent des fonctions socio-économiques
multiples, soit directement (alimentation en eau potable, activités de pêche, etc.) soit
indirectement (fonctions d’agréments et touristiques) reliées aux ressources en eau et à la
faune qu’ils hébergent. Il est donc essentiel de préserver la qualité de ces milieux aquatiques
menacés par l’urbanisation croissante et l’agriculture (Jackson et al. 2001b, Naiman & Turner
2000).
Ainsi, on assiste à un renforcement des mesures de protection et de surveillance des
milieux aquatiques (Carpenter 2002, E.P.C.N 2007). Parmi celles-ci, nous pouvons citer l’US
Clean Water Act (CWA, 1972) et la Directive Cadre Européenne sur l’Eau (DCE,
2000/60/EC, Communauté Européenne 2000), qui traduisent réglementairement les politiques
des Etats. Leurs objectifs environnementaux sont identiques (Novotny 2003), à savoir « la
restauration et le maintien de l’intégrité physique, chimique et biologique de l’ensemble des
masses d’eau continentales». Ceci conduit au concept « d’état écologique », défini comme
« l’expression de la qualité, de la structure et du fonctionnement des milieux aquatiques »
(Solimini et al. 2006). Ainsi, la DCE impose pour l’ensemble des masses d’eau de surface,
incluant les plans d’eau, l’atteinte du « bon état écologique » d’ici 2015 (Noges et al. 2009).
Enfin, ces deux textes réglementaires établissent une série de critères et de normes décrivant
le bon état écologique. Quatre compartiments biologiques (l’ichtyofaune, le phytoplancton,
- 20 -
les macrophytes et le phytobenthos, et la faune benthique invertébrée) peuvent contribuer
selon les masses d’eau considérées, à définir cet état écologique qui intègre également, en
soutien à la biologie, la physico-chimie et l’hydromorphologie.
En France, lors du dernier rapportage à la Commission Européenne relatif à l’état des eaux
de surface en date du 22 mars 2010
(http://www.rapportage.eaufrance.fr/annexes/dce/2010/FR.htm), on dénombrait 439 plans
d’eau soumis à la DCE (de plus de 50 ha pour la grande majorité, à l’exception de certains
lacs naturels). Ils se répartissaient de la manière suivante : 368 réservoirs et 71 lacs naturels.
En termes de gestion écologique, les plans d’eau français ont été peu considérés au cours
de ces 20 dernières années. Ainsi, pour définir la qualité écologique des milieux lentiques en
France, les gestionnaires (administrations nationales, régionales et locales) et les bureaux
d’études en environnement restent démunis ; en effet, les méthodes d’évaluation, en
particulier celles basées sur les bioindicateurs, demeurent rares (Appelberg et al. 2000, Drake
& Valley 2005, Hickman & McDonough 1996, Lafont et al. 1991, Mouthon 1993, Verneaux
et al. 2004). D’autre part, quand elles existent, ces méthodes concernent des types
d’environnement spécifiques – par exemple, les lacs de montagne ou les zones humides – et
sont basées sur les compositions faunistiques, ce qui les rend au final peu généralisables.
1.2. Les poissons en bioindication lacustre
Par rapport aux autres organismes à même de fournir une estimation de l’état écologique
des plans d’eau, les poissons présentent un certain nombre de caractéristiques intéressantes et
originales (Simon & Lyons 1995) :
Ils intègrent une variabilité environnementale sur de larges échelles d’espace et de
temps du fait de leur capacité de déplacement et de leur longévité ;
Ils sont à la fois sensibles à la qualité de l’eau et aux altérations de l’habitat en raison
de leurs besoins d’habitats de reproduction, de croissance, de repos et de refuge ;
Ils constituent le maillon le plus élevé des chaînes trophiques aquatiques et sont donc
sensibles aux altérations des autres éléments de la biocénose (producteurs primaires et
consommateurs secondaires essentiellement) ;
Leurs exigences écologiques sont relativement bien connues.
- 21 -
Depuis une vingtaine d’années, de nombreux travaux ont été conduits sur les relations
entre environnement et ichtyofaune lacustre. En effet, des approches macroécologiques
(Amarasinghe & Welcomme 2002, Gardezi & Gonzalez 2008, Griffiths 1997, Irz 2006,
Jackson et al. 2001a, Matthews 1998) ont mis en évidence que la structure des communautés
piscicoles résultait de processus agissant à différentes échelles de temps et d’espace (Beisner
et al. 2006, Tonn 1990, Tonn et al. 1990). Il est également acquis que le fonctionnement de
ces communautés piscicoles découle d’interactions entre les caractéristiques du biotope
(climatologie, qualité de l’eau, diversité et zonation des habitats, ressources alimentaires
disponibles), les exigences et les caractéristiques des espèces, les relations interspécifiques et
les pressions anthropiques s’exerçant sur les milieux (Jackson et al. 2001a). Un certain
nombre de lois générales (relation richesse spécifique / latitude ou / surface, par exemple)
régissant la distribution des espèces piscicoles dans les milieux lentiques ont ainsi été
identifiées (Barbour & Brown 1974, Irz 2006, Mehner et al. 2005, Pronier 2000, Rahel 1984,
Tonn & Magnuson 1982). Ces connaissances sur les variables environnementales structurant
les communautés piscicoles, représentent un préalable au développement d’outils de
bioindication. Elles permettent, en effet, d’orienter les recherches sur les liens de type
pressions / impacts à la base de la définition des bioindicateurs.
Bien que l’utilisation des organismes biologiques pour rendre compte de l’état des
systèmes soit assez ancienne, notamment en Europe (Kolkwitz & Marsson 1908), il s’agit
d’un domaine qui s’est fortement développé suite aux travaux de Karr (1981), aux Etats-Unis.
Ce dernier définit l’indice d’intégrité biotique (IBI) sur des principes largement repris dans la
DCE. Cet indice représente ainsi l’état du milieu, évalué au final par une note qui le
positionne sur une gamme de valeurs entre « l’état de référence » (ou « pristine », c’est-à-dire
non perturbé par l’Homme) et l’état dégradé (ou « mauvais état », au sens de la DCE). Une
autre caractéristique de cet indice d’intégrité biotique est de ne pas utiliser seulement la
position taxonomique des espèces, mais d’asseoir le diagnostic par la prise en compte de
certaines caractéristiques des espèces. L’utilisation des traits biologiques a permis de décrire
les peuplements par la présence, ou l’importance, de différents groupes ou guildes d’espèces
ayant les mêmes affinités biologiques et/ou caractéristiques fonctionnelles. Ce sont donc
toutes ces caractéristiques fonctionnelles et/ou taxonomiques appelées « métriques », qui sont
ainsi combinées au sein d’un indice multimétrique pour diagnostiquer l’état écologique des
systèmes.
Pour les plans d’eau, cette méthode a été développée une dizaine d’années plus tard en
Amérique du Nord dans le cadre du « Lake and Reservoir Bioassessment Program » de
- 22 -
l’USEPA (USEPA 1998). Par exemple, Hughes and Noss (1992), Lyons et al. (2000) et
Seilheimer and Chow-Fraser (2006) ont testé des métriques candidates à l’élaboration d’un
indice poisson pour les lacs naturels ; Karr et la Tennessee Valley Authority (TVA) ont mené,
quant à eux, des recherches pour appliquer les IBI sur les réservoirs (Dionne & Karr 1992,
Dycus & Meinert 1992, Jennings et al. 1995, McDonough & Hickman 1999). En Europe,
plusieurs indices poisson ont également été développés durant la dernière décennie
(Appelberg et al. 2000, Belpaire et al. 2000, Gassner et al. 2003, Tammi et al. 2001), le plus
abouti étant celui développé en Suède (Holmgren et al. 2007). Néanmoins, très récemment,
Beck & Hatch (2009) ont discuté des obstacles et des imperfections fréquemment rencontrés
dans le développement des indices poisson actuels. En particulier, les critères qui sont utilisés
pour classer l’état d’altération des plans d’eau ne prennent pas en compte les effets de
certaines perturbations anthropiques sur les biotopes aquatiques, telles que la perte de
connectivité hydrologique, la pêche sportive et récréative, ou la présence d’espèces invasives
ou introduites. En effet, actuellement, seules les pressions anthropiques qui se traduisent
directement par des altérations physiques ou chimiques des écosystèmes aquatiques sont
considérées (Beck & Hatch 2009, Drake & Pereira 2002). De plus, malgré de nombreuses
études, un point faible essentiel dans les travaux de bioindication lacustre reste la prise en
compte très partielle de la variabilité environnementale naturelle, en particulier dans la plupart
des études conduites en Amérique du Nord où les jeux de données utilisés sont très
conséquents, mais considérés comme homogènes (Whittier et al. 2002). Cette faiblesse a pour
corollaire l’impossibilité d’utiliser un même indice sur des territoires vastes où les conditions
environnementales sont très variables. Enfin, rares sont les méthodes actuelles qui sont
conformes aux exigences de la DCE ou du US CWA pour l’évaluation de la condition des
biotopes lacustres. En effet, la question de la référence biologique est, rarement, clairement
explicitée, généralement du fait de l’absence ou du manque de sites de référence.
1.3. Objectifs de l’étude et organisation générale du mémoire
L’objectif de la présente étude est d’évaluer la contribution relative des facteurs naturels et
anthropiques sur la structure des communautés piscicoles des plans d’eau français. Cette
démarche devra conduire à l’identification de variables de bioindication, ou métriques, qui
pourront entrer dans la construction d’un indice poisson national. Cet indice devra rendre
compte de la structure fonctionnelle, de la composition et de l’abondance des espèces
- 23 -
piscicoles, trois caractéristiques des communautés à considérer dans le cadre réglementaire de
la DCE.
Ainsi, ce mémoire de thèse est structuré en 4 chapitres.
Le chapitre 2 décrit brièvement l’ensemble des données piscicoles, environnementales et

de pressions anthropiques, pour 106 plans d’eau répartis sur l’ensemble de la France.
Chacun des chapitres suivant aborde une question liée au processus d’élaboration d’un
indice poisson.
Le chapitre 3 est structuré autour d’un article (Launois et Argillier, sous presse dans
Hydroécologie appliquée). Il s’agissait, ici, d’explorer la possibilité de dégager des relations
entre l’ichtyofaune et les pressions anthropiques qui s’exercent sur les plans d’eau français,
après regroupement des plans d’eau par types homogènes d’un point de vue environnemental.
Rappelons que cette démarche, adoptée dans de nombreuses études, est la seule reconnue par
la DCE et les directives nationales afférentes pour l’expression des états écologiques. Les
limites de cette approche sont discutées, et elles nous ont conduits à envisager des méthodes
alternatives.
Dans le chapitre 4, nous nous sommes interrogés sur la possibilité d’utiliser des
caractéristiques des communautés piscicoles (mêmes métriques) communes aux lacs et aux
retenues, pour rendre compte des altérations anthropiques. Dans ce travail, compte tenu des
limites de l’approche typologique mise en évidence dans les premiers travaux et de
l’amélioration de la qualité du jeu de données, une approche par modélisation, de type « site
spécifique », a été développée. L’article publié constitue le cœur de cette partie du manuscrit
(Launois et al., 2011, Ecology of Freshwater Fish).
Après avoir démontré la nécessité de développer des outils distincts pour chacun des
milieux, et la possibilité de mettre en œuvre une approche « site spécifique » pour établir des
corrélations entre les métriques piscicoles et les pressions, nous avons travaillé à la
construction de deux indices. La démarche est décrite dans le chapitre 5. Pour s’affranchir des
difficultés liées au manque de sites de référence à l’échelle nationale, une solution originale a
été appliquée. L’article correspondant (Launois et al., sous presse dans Ecological Indicators)
décrit les différentes étapes jusqu’à l’élaboration d’un indice, en se basant sur une approche
encore peu utilisée, mais prometteuse : la hindcasting model approach (Baker et al. 2005,
Kilgour & Stanfield 2006).
- 24 -
Les principaux résultats des études présentées dans ce mémoire sont repris dans une
conclusion générale (chapitre 6) qui propose des perspectives découlant de ce travail.
- 25 -
- 26 -
CHAPITRE 2
Données disponibles à l’échelle de la France

Ce chapitre a pour but de fournir des informations succinctes sur les jeux de données –
biologiques, écologiques et anthropiques – utilisés dans le cadre de ce travail de thèse pour
faciliter la compréhension de l’ensemble du manuscrit. Ces données sont décrites de façon
plus détaillée dans les différents articles.
Les termes « plans d’eau » et « lacs » seront utilisés indifféremment dans la suite du
mémoire.
2.1. L’ichtyofaune
2.1.1. Les inventaires piscicoles

Les inventaires piscicoles ont été réalisés sur 146 plans d’eau (en date du 1er janvier 2010)
répartis sur l’ensemble de la France, au moyen d’une méthode standardisée : le protocole
C.E.N. nordique (C.E.N. 2005a). Les données de pêches collectées présentent l’avantage
d’être homogènes à l’échelle nationale. Les différents inventaires peuvent ainsi être comparés
dans le temps pour un lac donné, mais également entre lacs. Une description plus détaillée
concernant la méthode de pêche standardisée est disponible au chapitre 4.
Ces pêches d’échantillonnage, dont les données ont été mises à disposition par l’ONEMA,
les Agences de l’Eau et des bureaux d’étude, concernent 106 réservoirs et 40 lacs naturels
(Figure 2. 1). D’autres pêches ont été réalisées sur des lacs situés à une altitude supérieure à
1500 m, mais ces données ont été exclues de nos analyses. En effet, comme le rapportent
Argillier et al. (2002b), ces plans d’eau présentent des richesses spécifiques très faibles, et
sont quasi exclusivement peuplés par des espèces introduites. De plus, l’ichtyofaune y a été
écartée du diagnostic pour la DCE. Les enjeux en termes de développement d’outils de
bioindication reposent donc sur d’autres compartiments biologiques. De même, parmi les
milieux échantillonnés au niveau national, certains n’ont pu être que partiellement intégrés
dans les analyses. En effet, certaines variables environnementales ou de pressions n’étaient
pas disponibles.
- 28 -
Figure 2. 1 : Localisation des sites d’étude en France et délimitation des principales zones
hydrographiques.
Les symboles ◊ et ♦ représentent, respectivement, les lacs naturels et les plans
d’eau artificiels.
Au total, 42 espèces de poissons ont été pêchées entre 2005 et 2009 (cf. Chapitres 3, 4 et
5). Pour chacune, tous les individus ont été identifiés. Les captures de chacune de ces espèces
par unité d’effort de pêche, en nombre d’individus et en biomasse sont disponibles pour
l’ensemble de ces plans d’eau.
2.1.2. Les caractéristiques biologiques des espèces

Les critères de classification des espèces selon leur tolérance, préférenda d’habitat de
reproduction et guilde trophique sont donnés dans le Tableau 2. 1. Ils reposent sur des études
publiées (Balon 1975, Bruslé & Quignard 2001, Keith & Allardi 2001), des bases de données
- 29 -
en ligne (Pont et al. 2006) et des avis d’experts (H. Persat, N. Poulet, et O. Schlumberger,
communications personnelles).
Tableau 2. 1 : Critères de classification des espèces piscicoles en guildes fonctionnelles.

Guildes Définitions
Guilde de tolérance Espèces insensibles à tout impact dû aux altérations du
(Karr et al. 1986, Tolérants régime hydrologique, du flux des nutriments, de la structure
www.fame.boku.ac.at) de l’habitat et de la chimie de l’eau.
Espèces dont la perte est une réponse à la dégradation de
Intolérants l’état écologique, alors que le nombre d’espèces tolérantes
tend à augmenter avec le degré de perturbation.
Guilde de reproduction
Espèces préférant se reproduire sur des substrats
(Balon 1975, Dycus & Lithophiles
minéralisés.
Meinert 1992)
Espèces nécessitant des substrats minéralisés pour se
Stricts Lithophiles
reproduire.
Espèces nécessitant des substrats végétalisés pour se
Stricts Phytophiles
reproduire et dont les alevins ne sont pas photophobiques.
Guilde trophique
(Bruslé & Quignard 2001,
Omnivores Espèces se nourrissant à la fois d’animaux et de végétaux.
Dycus & Meinert 1992,
Goldstein & Simon 1999)
Espèces se nourrissant majoritairement
Invertivores
d’invertébrés/insectes.
Espèces se nourrissant principalement de zooplancton
Planctivores
et/ou phytoplancton.
Herbivores Espèces se nourrissant de végétaux.
Piscivores Espèces se nourrissant de poissons.
L’utilisation de ces guildes permet une caractérisation fonctionnelle des assemblages,

plutôt que taxonomique. Elle autorise donc à comparer les communautés piscicoles entre
zones géographiques qui abritent des cortèges d’espèces différents. Les caractéristiques
biologiques attribuées à chaque espèce correspondent aux caractéristiques des adultes. Le
changement de régime alimentaire par exemple entre juvéniles et adultes n’a pas été pris en
considération. De plus, l’on suppose la variabilité inter-spécifique plus importante que la
variabilité intra-spécifique (Blanck 2007).
2.2. Les caractéristiques environnementales naturelles des lacs et de leurs bassins

versants
Les caractéristiques naturelles des plans d’eau, connues pour influencer les peuplements
de poissons, ont été collectées. Ces paramètres ont, en partie, été identifiés lors de deux thèses
- 30 -
antérieures sur le sujet (Irz 2006, Pronier 2000). Des informations complémentaires, relatives
à la géologie sur le bassin versant des plans d’eau ou aux régimes climatique et hydrologique,
ont été collectées à partir d’un Système d’Information Géographique (SIG) ArcGIS® (version
9.2). Les caractéristiques environnementales naturelles décrivent donc les systèmes aux
échelles régionale et locale. Les sources de données utilisées ont été :
Carthage® pour les délimitations des bassins versants et des plans d’eau (Institut
Géographique National (IGN) – surface bassin versant, surface plan d’eau et
indice de développement des berges),
Les cartes géologiques© BRGM au 1/50000ème et au 1/1000000ème pour la géologie
sur les bassins versants (Bureau de Recherches Géologiques et Minières (BRGM)
– indice géologique),
L’Analyse Utilisant le RELief pour l’HYdrométérologie (AURELHY – résolution à
1km) afin de calculer les températures moyennes mensuelles (Météo-France© -
Températures moyennes annuelles des mois de janvier et de juillet),
Le modèle numérique de terrain (MNT) à 250 m pour la topographie et la
délimitation des bassins versants des lacs (IGN – profondeur maximale du plan
d’eau, volume de développement),
BD Topo© pour l’altitude des plans d’eau (IGN – altitude),
Les modèles hydrologiques développés par H. Pella et E. Sauquet (Pella et al.
2006), et N. Folton (logiciel LOIEAU 2) pour le calcul des débits transitant dans
les plans d’eau et le calcul des temps de résidence (temps de résidence, marnage
des plans d’eau).
2.3. Les facteurs anthropiques
En France, la biodiversité des écosystèmes aquatiques est fortement menacée,

essentiellement par l’urbanisation croissante et l’agriculture intensive (Colin et al. 2000, Even
et al. 1998). En effet, outre l’altération physique et chimique des écosystèmes aquatiques, ces
pressions anthropiques altèrent les communautés biologiques (Jennings et al. 1999a,
Scheuerell & Schindler 2004, Solimini et al. 2006). Ces pressions peuvent avoir soit une
action directe sur la composition spécifique ou la structure des communautés piscicoles, c’est
le cas des empoissonnements ou des introductions d’espèces fréquentes sur les plans d’eau
- 31 -
(Argillier et al. 2002a, Crivelli 1995, Welcomme 1988), soit une action indirecte lorsqu’elles
détériorent les habitats ou la qualité de l’eau (Crowder et al. 1981, Fischer & Eckmann 1997,
Mosisch & Arthington 1998, Wolter & Arlinghaus 2003).
Pour asseoir le choix des pressions anthropiques à prendre en considération dans cette
étude, une synthèse bibliographique, présentée en Annexe 1, a recensé les impacts des
pressions anthropiques sur l’ensemble des organismes vivants lacustres. Ce travail omet
toutefois volontairement les pressions chimiques, en particulier les rejets en nutriments et
l’acidification, car elles ont fait l’objet de nombreuses synthèses (Brönmark & Hansson 2002,
Haines 1981, Kidd et al. 2007, Ostendorp et al. 2004a, Schindler 1988).
2.3.1. Les pressions anthropiques à l’échelle du bassin versant

Neuf alias de pressions anthropiques s’exerçant à l’échelle régionale ont été sélectionnés
et inclus dans les différentes analyses menées au cours de cette thèse (cf. chapitres 3, 4 et 5).
Les sources de données utilisées pour collecter ces variables ont été :
Corine Land Cover 2002 pour l’occupation des sols sur les bassins versants
(Institut Français de l’Environnement (IFEN) – occupations du sol de types urbain
et agricole, surface agricole),
BD Route 500® pour les linéaires de voies de communication sur les bassins
versants (IGN – densité de réseaux routiers et ferrés sur le bassin versant),
GTOPO Landscan Global Population (résolution à 1km) pour les densités de
population sur les bassins versants (Oak Ridge Laboratory – densité de population
sur le bassin versant et sur une zone tampon de 1000 m autour du lac).
Le modèle NOPOLU System 2 pour les valeurs de bilans et de surplus en
nutriments (Azote total (NTot) et Phosphore total (PTot) – surplus en NTot, bilans en
NTot et PTot – Pöyry Environnement© - BETURE CEREC / JAAKKO PÖYRY
INFRA 2000).
La carte de l’aléa « d’érosion des sols » pour coupler cette valeur aux teneurs en
PTot (Institut National de la Recherche Agronomique (INRA)).
2.3.2. Les pressions anthropiques locales

Les données ont été obtenues en application d’un protocole d’échantillonnage écossais, en
voie de standardisation, appelé Lake Habitat Survey (LHS, voir Rowan et al. 2006). Le
- 32 -
principe de cette méthode est une caractérisation des habitats physiques des zones rivulaires,
depuis la berge jusqu’à la zone littorale. Un inventaire des pressions anthropiques modifiant la
structure naturelle des berges et des environs proches du plan d’eau est réalisé. Les usages
anthropiques dont le plan d’eau fait l’objet sont également recensés. Ces pressions sont
ensuite agrégées pour constituer un indice synthétique qui a été utilisé dans la dernière étude
menée au cours de cette thèse (cf. chapitre 5).
- 33 -
- 34 -
CHAPITRE 3
Sélection de métriques piscicoles par une approche intuitive : une première démarche
basée sur une typologie physique des plans d’eau
- 35 -
Classiquement, les indicateurs biologiques, quelle que soit la communauté considérée,
sont développés pour des types de milieux réputés homogènes (Moss et al. 2003, Plafkin et al.
1989, Schupp 1992). En effet, cette approche, dite « type-spécifique », repose sur des
techniques de clustering afin de grouper préalablement les sites de référence selon leur
homogénéité faunistique ou environnementale (Roset et al. 2007). Elle est utilisée dans de
nombreuses études, avant de modéliser les assemblages biologiques (Wright et al. 2000).
Ainsi, il est admis que la variabilité des métriques au sein de ces types résulte principalement
de l’effet des facteurs anthropiques, la variabilité liée à l’environnement naturel étant réduite.
Cette approche par types a été adoptée aux Etats-Unis au début des années 80 pour le
développement de la plupart des premiers indicateurs biologiques lacustres (Plafkin et al.
1989, Schupp 1992). Néanmoins, elle a été peu suivie en Europe, vraisemblablement pour
deux raisons : (i) une forte hétérogénéité environnementale des plans d’eau en regard de leur
faible densité dans la plupart des régions et (ii) une absence de standardisation des protocoles
de recueil d’informations qui conduit a une hétérogénéité des données disponibles. Bien que
ce ne soit pas nécessairement explicité, cette notion de typologie est toutefois sous-jacente
dans le développement de quelques indicateurs utilisés régionalement. En France, c’est le cas
des indices oligochètes et mollusques développés sur les lacs naturels du secteur Alpin
(AFNOR 2005, Lafont et al. 1991, Mouthon 1993) ; c’est aussi le cas, par exemple, avec des
indices poisson développés en Espagne pour les retenues de Catalogne
(Agència Catalana de l'Aigua 2003) et en Autriche pour les lacs naturels (Gassner et al. 2003,
Gassner et al. 2005).
De plus, il s’agit de la méthodologie soutenue par la DCE (European Community 2000) et
donc préconisée pour construire les indicateurs sur lesquels les Etats membres doivent établir
leur diagnostic d’état écologique. C’est aussi sur une base régionale et typologique que se fait
l’harmonisation (intercalibration) des méthodes d’évaluation de l’état des milieux.
Dans cette perspective, ce chapitre présente un premier essai d’application d’une approche
typologique pour sélectionner des métriques piscicoles candidates à l’élaboration d’un indice
poisson répondant aux exigences de la DCE. Cette typologie a été réalisée à partir de critères
environnementaux naturels dont l’influence sur la structure des communautés de poissons
lacustres a été démontrée sur un jeu de données comparables (Irz 2006). Au sein des types
identifiés, l’option a été prise de ne mesurer que la réponse des métriques à des pressions
s’exerçant à l’échelle du bassin versant.
- 36 -
Les résultats présentés sont encourageants ; en effet, certaines métriques classiquement
sélectionnées dans la construction d’indices poisson pour les milieux lacustres, répondent aux
pressions de type urbanisme, agriculture ou globale. De plus, au sein d’un groupe de lacs, les
métriques qui répondent aux pressions agricoles sont distinctes de celles qui répondent à
l’urbanisation. Ces résultats permettent d’envisager la possibilité d’utiliser les communautés
piscicoles pour évaluer l’effet des perturbations anthropiques sur les milieux lentiques
français.
Toutefois, ce travail souligne aussi les limites de l’approche typologique dans le contexte
national. En particulier, nous avons montré que les groupes définis pouvaient regrouper des
milieux encore très hétérogènes par certains critères (géographie, origine, etc.). Ainsi, une part
significative de la variabilité naturelle peut demeurer dans l’indice biologique final et le risque
de faible sensibilité de l’indice aux pressions anthropiques pourrait donc être élevé (Beck &
Hatch 2009, Borja et al. 2009, Carvalho et al. 2009, Schmutz et al. 2007). Pour réduire ces
biais, la prise en compte de variables environnementales supplémentaires conduirait
certainement à l’identification d’un nombre plus élevé de types. Toutefois, pour chacun
d’entre eux, les effectifs seraient plus faibles, ce qui ne serait plus compatible avec l’analyse
des relations pressions/ichtyofaune. De plus, on peut s’interroger sur la pertinence d’une
définition de types de milieu sur la base de critères environnementaux, dont les valeurs
mesurées sont réparties sur toute l’étendue d’une échelle continue.
Ces travaux montrent également que le nombre de sites pas ou peu perturbés pour certains
types est restreint, voire nul dans certains cas. Dans ces conditions, il paraît difficilement
envisageable de définir des objectifs environnementaux tels qu’ils sont définis par les textes
réglementaires, à savoir par mesure d’un écart à la référence (Anonymous 2003b). De plus,
entre les différents groupes, les gammes de pressions mesurées sont elles-mêmes distinctes, ce
qui hypothèque la comparabilité et la pérennité des indices éventuellement développés.
Enfin, la dernière limite à cette approche typologique repose sur un certain nombre de
paramètres qui se révèlent être rapidement affectés par le réchauffement climatique (Noges et
al. 2007). Certains milieux, en particulier ceux situés à la limite de certains types, pourraient
changer de groupe typologique.
Notons par ailleurs, que d’un point de vue opérationnel, la différence entre la typologie
établie ici et la typologie nationale (Ministère de l'Ecologie et du Développement Durable
2005) complexifie l’établissement des objectifs environnementaux assignés à chaque site. En
- 37 -
effet, la mise en œuvre de cette démarche pour le développement de tous les bioindicateurs
supposerait que, selon l’indicateur considéré, l’on se réfère à l’une ou l’autre des typologies.
Par conséquent, ce travail souligne non seulement la nécessité de collecter d’autres

données à l’échelle nationale, mais également le besoin de rechercher une solution alternative
à cette approche typologique.
Intérêts et limites d’une approche par type de milieu pour le développement d’un indice
poisson lacustre français (sous presse dans Hydroécologie appliquée).
Lionel Launois et Christine Argillier
3.1. Introduction
Les systèmes lentiques ont par nature des caractéristiques physiques, chimiques et
biologiques différentes. En effet, les processus hydrologiques, la géologie du bassin versant,
la position géographique des plans d’eau et la morphologie de la cuvette lacustre, sont autant
de paramètres environnementaux qui affectent les processus biologiques, l’assemblage des
communautés (Plafkin et al. 1989) et qui contribuent à rendre les caractéristiques physiques et
chimiques d’un plan d’eau uniques. Ce caractère unique conféré par les variables
environnementales aux lacs, rend difficile le développement d’outils d’évaluation
transposables à de larges échelles spatiales (Schmutz et al. 2007, Schupp 1992).
Historiquement, le recours à une approche typologique apparaît comme le moyen le plus
intuitif et adéquat pour s’affranchir de cette variabilité environnementale (Drake & Pereira
2002, Hawkins et al. 2000, Schupp 1992). En effet, en passant par la création de groupes de
lacs homogènes d’un point de vue environnemental, on espère que les processus écologiques
qui s’opèrent au sein de chacun de ces groupes soient similaires, tout comme les réponses des
communautés piscicoles aux pressions anthropiques. Quel que soit le type de milieux
continental et côtier, c’est ce cadre théorique qui a été retenu pour la mise en place des
indicateurs à utiliser en application de la Directive Cadre Européenne sur l’Eau (DCE ;
Communauté Européenne 2000). Ainsi, en France une typologie des plans d’eau a été définie.
Elle est basée sur l’origine naturelle ou anthropique du plan d’eau, sur l’hydroécorégion
(géologie, relief et climat), la morphologie de la cuvette et pour certains types sur le
fonctionnement hydraulique. Douze types de lacs naturels et 18 types de plans d’eau d’origine
anthropique sont décrits (Ministère de l'Ecologie et du Développement Durable 2005).
- 38 -
L’objectif de ce travail est d'examiner, à partir des données d’inventaire disponibles, les
possibilités de développement d’un indicateur poisson adapté aux plans d’eau français,
d’origine naturelle et anthropique, dans le cadre d’une approche par type de masses d’eau,
conforme aux principes généraux de la DCE. Pour cela, compte tenu de la très forte variabilité
environnementale résiduelle dans les types nationaux cités précédemment [exemple de la
profondeur décrit dans De Bortoli and Argillier (2008)], la démarche consiste à (1) établir une
nouvelle typologie physique des plans d’eau sur la base de variables environnementales,
connues pour influencer l’ichtyofaune, disponibles au niveau national, puis à (2) analyser la
réponse aux pressions de métriques candidates, dans les types de plans d’eau où les données
relatives à l’ichtyofaune sont les plus nombreuses. Ce travail vise à dégager les avantages et
les limites de cette approche typologique au niveau national, pour la mise au point d’un
bioindicateur basé sur l’ichtyofaune des plans d’eau.
3.2. Matériel et méthodes
3.2.1. Réalisation de la typologie lacustre

Le jeu de données environnementales disponible concerne 34 lacs naturels et 104
réservoirs sur un total de 71 lacs et 368 retenues sur lesquels doit s’effectuer un rapportage au
niveau européen. La localisation de ces 138 plans d’eau est présentée sur la Figure 3. 1a et
leur liste en annexe 2. Ces plans d’eau sont bien répartis sur l’ensemble du territoire, mais ils
sont tous situés à moins de 1500 m d’altitude, limite au-delà de laquelle les densités et les
richesses spécifiques des poissons sont très faibles (Argillier et al. 2002b) et donc
potentiellement limitantes pour le calcul de certaines métriques.
Ces sites ont été décrits par cinq variables environnementales « physiques »,
indépendantes entre elles et non influencées par les pressions : la surface du bassin versant,
l’altitude, la surface et la profondeur maximale du plan d’eau, ainsi que l’indice de
développement des berges [Shoreline Development Factor (SLDF)] (Eadie & Keast 1984).
Ces variables ont été log transformées. En l’absence de données homogènes des températures
de l’eau, celles-ci n’étant prises que ponctuellement lors des pêches effectuées à des dates
variables, la température moyenne de l’air des mois de janvier et de juillet ont été calculées à
partir du modèle AURHELY, qui recense ces valeurs moyennes sur la période 1961-1990
(Benichou & Le Breton 1987). Elles ont été utilisées pour créer deux nouvelles variables
relatives à la température non corrélées, comme pour l’indice poissons rivière : T janvier + T°
- 39 -
juillet et T° juillet – T° janvier (Oberdorff et al. 2001, Oberdorff et al. 2002, Pont et al. 2006,
Pont et al. 2007). Ces deux variables ont été utilisées telles quelles.
Pour la réalisation de groupes de systèmes homogènes du point de vue de l’environnement

naturel, les plans d’eau naturels et d’origine artificielle ont été traités séparément. Les groupes
de lacs qui s’apparentent le plus du point de vue environnemental ont été définis par la
réalisation d’une classification non supervisée, ascendante (méthode des K-means). Les sept
variables environnementales précédemment décrites et présentées dans le Tableau 3. 1 ont
d’abord été centrées et réduites. Ensuite, une analyse en composantes principales a été réalisée
sur ces variables environnementales. Les cinq premières composantes ont été sélectionnées,
car elles expliquaient plus de 90% de la variabilité. Finalement, ces cinq composantes ont été
analysées selon la méthode des K-means (MacQuenn 1967). Cette méthode partitionne les
données en K groupes, ou "clusters", ne se chevauchant pas. Ce résultat est obtenu en
positionnant K "prototypes" ou "centroïdes" dans les régions de l'espace les plus peuplées
(Figure 3. 2). Chaque observation est alors affectée au prototype le plus proche (règle dite "de
la Distance Minimale"). Chaque groupe contient donc les observations qui sont plus proches
d'un certain prototype que de tout autre prototype.
(1a) (1b)
Figure 3. 1 : Localisation géographique des plans d’eau étudiés (1a) pour la réalisation de la
typologie et (1b) pour l’analyse de la réponse des métriques aux pressions.
Dans ce dernier cas, les plans d’eau en grisé sont exclus.
- 40 -
Figure 3. 2 : Illustration de la méthode K-means, avec le partitionnement des données en 3
groupes, ou « clusters ».
Ce résultat est obtenu en positionnant 3 « prototypes » (c1, c2 et c3) dans les
régions de l’espace les plus peuplées. Adapté de MacQuenn (1967).
Tableau 3. 1: Valeur moyenne et écart type des principales caractéristiques

environnementales des plans d’eau appartenant aux cinq types définis par l’analyse
(pour les types, voir « 1.3.1 Typologie physique des plans d’eau français »).
Retenues Lacs naturels
Groupe 1 Groupe 2 Groupe 3 Groupe 1 Groupe 2
(n = 20) (n = 35) (n = 49) (n = 24) (n = 10)
824,2
Altitude (m) 371,4 (166,1) 452,1 (257,6) 82,9 (67,4) 14,5 (10,5)
(271,2)
Surface du bassin versant 1227,8 373,6 770,3
2 77,9 (217,1) 403,1 (255,7)
(km ) (1603,7) (1644,9) (2222,0)
203,6 2029,1
Surface du plan d’eau (ha) 613,2 (610,5) 96 (59,4) 107,5 (99,5)
(534,6) (2458,9)
Indice de dévelopement des
† 4,43 (1,72) 2,39 (0,85) 3,35 (1,60) 1,54 (0,58) 1,81 (0,56)
berges (SLDF ; sans unité)
Profondeur maximale (m) 53,2 (30,9) 22,9 (21,1) 13,0 (10,5) 29,7 (27,1) 9,7 (8,1)
T°C moy. Jan + T°C moy Jul 24,6 (3,1) 21,2 (2,7) 25 ,6 (2,0) 17,7 (3,5) 29,1 (1,6)
T°C moy. Jul - T°C moy Jan 17,2 (1,1) 17,6 (1,0) 13 ,7 (1,7) 18,4 (0,6) 14,3 (1,2)
†
Calculé ainsi : périmètre du lac / (√4π×surface du lac).
3.2.2. Analyse de la réponse de l’ichtyofaune aux pressions dans chaque type
3.2.2.1. Les données poissons

L’ichtyofaune a été échantillonnée entre 2005 et 2008, sur 44 des 138 plans d’eau français
sur lesquels les variables environnementales sont disponibles (voir liste en annexe 2). Le
protocole utilisé est le protocole standardisé aux filets maillants (C.E.N. 2005b). Il prévoit une
campagne d’échantillonnage aléatoire stratifiée par zones de différentes profondeurs, durant la
période estivale (juin à octobre, la température de l’eau en surface devant être supérieure à
15°C). L’effort de pêche est défini en fonction de la profondeur et de la surface des systèmes.
- 41 -
Les espèces présentes ont été recensées et tous les poissons ont été comptés et pesés.
A partir de ces données, les abondances exprimées en nombre d’individus et en biomasse
par unité d’effort (surface x temps de pose) ont été calculées. De plus, onze traits d’histoire de
vie associés aux espèces ont été définis sur la base de données de la littérature (Balon 1975,
Bruslé & Quignard 2001, Keith & Allardi 2001), sur les bases en ligne (Pont et al. 2006) et
sur avis d’experts en particulier pour les traits liés au régime alimentaire (H. Persat, N. Poulet,
O. Schlumberger) (Tableau 3. 2). Ces traits ont été utilisés pour le calcul de quatorze
métriques piscicoles choisies parmi celles les plus fréquemment utilisées en bioindication sur
les plans d’eau et dont le calcul repose sur plus de deux espèces dans l’ensemble du jeu de
données (Tableau 3. 3).
- 42 -
Tableau 3. 2: Caractéristiques des 29 espèces de poissons capturées dans 16 plans d’eau du groupe LN1 et 10 du groupe RA3, lors d’une
campagne d’échantillonnage par plan d’eau effectuée entre 2005 et 2009, selon un protocole d’échantillonnage standardisé (C.E.N
2005).
1 = oui et 0 = non.
Abondance Abondance
Nom Fréquence Fréquence Relative Relative Lithophiles Phytophiles Omnivores
Code Espèce Tolérants Invertivores Planctivores Piscivores
commun (%) RA3 (%) LN1 moyenne moyenne stricts stricts Généralistes
(%) RA3 (%) LN1
GAR Rutilus rutilus Gardon 100,00 93,75 29,01 37,26 1 0 0 1 1 1 0
Perche
PER Perca fluviatilis 100,00 93,75 19,63 36,71 1 0 0 0 1 1 1
franche
Brème
BRE Abramis brama 100,00 43,75 20,98 4,96 1 0 0 1 1 1 0
commune
SAN Sander lucioperca Sandre 90,00 25,00 7,44 0,72 0 0 0 0 0 0 1
Scardinius
ROT Rotengle 80,00 56,25 0,99 7,54 0 0 1 1 1 1 0
erythrophthalmus
ABL Alburnus alburnus Ablette 80,00 43,75 1,49 0,85 1 0 0 1 1 1 0
Gymnocephalus
GRE Grémille 70,00 37,50 3,11 1,83 0 0 0 1 1 0 0
cernuus
Brème
BRB Blicca bjoerkna 60,00 6,25 6,67 1,21 1 0 1 0 1 1 0
bordelière
BRO Esox lucius Brochet 60,00 62,50 0,18 0,42 0 0 1 0 0 0 1
Poisson
PCH Ameiurus melas 40,00 6,25 9,56 0,32 1 1 0 1 1 0 1
chat
Perche
PES Lepomis gibbosus 40,00 6,25 0,07 0,25 1 1 0 0 1 1 0
soleil
CCO Cyprinus carpio Carpe 40,00 25,00 0,04 0,05 1 0 0 1 1 1 0
TAN Tinca tinca Tanche 20,00 37,50 0,03 0,53 1 0 1 1 1 0 1
Carassius
CAS Carassin 20,00 12,50 0,01 0,21 1 0 1 1 1 0 0
carassius
GOU Gobio gobio Goujon 10,00 43,75 0,03 1,97 0 0 0 0 1 0 0
Oncorhynchus Truite arc-
TAC 10,00 12,50 0,08 0,76 0 1 0 0 1 0 1
mykiss en-ciel
SIL Silurus glanis Silure 10,00 6,25 0,02 0,02 0 0 1 0 1 0 1
MUP Liza ramada Mulet porc 10,00 0,00 0,50 0,00 1 1 0 0 1 0 0
BOU Rhodeus amarus Bouvière 10,00 0,00 0,16 0,00 0 0 0 0 1 0 0
CHE Squalius cephalus Chevaine 0,00 50,00 0,00 1,95 0 0 0 1 1 1 1
Coregonus Corégone
COR 0,00 37,50 0,00 0,93 0 1 0 0 1 1 0
lavaretus lavaret
Omble
OBL Salvelinus alpinus 0,00 12,50 0,00 0,74 0 1 0 0 1 0 1
chevalier
TRF Salmo trutta fario Truite fario 0,00 18,75 0,00 0,53 0 1 0 0 1 0 1
CHA Cottus gobio Chabot 0,00 12,50 0,00 0,08 0 1 0 0 1 0 0
Salvelinus
CRI Cristivomer 0,00 6,25 0,00 0,08 0 1 0 0 1 0 1
namaycush
Alburnoides
SPI Spirlin 0,00 6,25 0,00 0,04 0 1 0 0 1 1 0
bipunctatus
Leuciscus
VAN Vandoise 0,00 6,25 0,00 0,02 0 1 0 1 1 1 0
leuciscus
Blennie
BLE Salaria fluviatilis 0,00 6,25 0,00 0,02 0 1 0 0 1 0 0
fluviatile
Lote de
LOT Lota lota 0,00 6,25 0,00 0,01 0 1 0 0 1 0 1
rivière
- 44 -
Tableau 3. 3: Liste des 14 métriques testées pour l’analyse des réponses de l’ichtyofaune de 2
types de plans d’eau aux pressions.
Type RA3 (n = 10) Type LN1 (n = 16)
Métriques candidates Ecart- Ecart-
Moyenne Variations Moyenne Variations
type type
Richesse spécifique totale
9,50 1,96 7-12 7,75 2,57 5-13
(RS)
†
Richesse en espèces natives 5,90 1,29 3-8 6,25 2,52 3-13
Richesse en espèces
6,10 1,52 4-8 3,69 1,35 1-6
tolérantes
1,00 0,94 0-3 1,38 1,75 0-6
lithophiles strictes
2,50 1,18 1-4 1,81 1,11 0-3
phytophiles strictes
8,00 1,89 6-11 6,88 2,53 4-12
invertivores
6,00 1,15 4-7 4,69 1,66 2-7
planctivores
3,30 0,82 2-4 3,38 1,41 1-6
piscivores
5,50 1,65 4-8 4,13 1,54 1-6
omnivores généralistes
Indice de Shannon-Wiener 1,35 0,27 0,94-1,81 1,11 0,21 0,61-1,52
Indice de diversité de Simpson 0,65 0,09 0,48-0,79 0,56 0,11 0,27-0,68
Indice d'équitabilité (Indice de
0,15 0,03 0,07-0,18 0,16 0,06 0,08-0,25
Shannon / RS)
Captures par unité d’effort de
2407 1407 591-4729 967 1362,68 128-5811
pêche (CPUE_Nb)
Biomasse par unité d’effort de
124 54 55-206 57 35 15-134
pêche (BPUE_gr)
†
Les espèces natives sont définies comme étant des espèces de poissons natifs sur le bassin versant
du plan d’eau étudié (Keith and Allardi 2001).
3.2.2.2. Les données de pressions

Les données de pressions étaient disponibles pour 44 lacs sur lesquels l’ichtyofaune a été
échantillonnée. Les pressions s’exerçant localement étant trop partielles sur l’ensemble des
sites, on s’est intéressé uniquement aux pressions sur le bassin versant des plans d’eau (BV).
Neuf variables de pressions (ou de forces motrices à l’origine des pressions, voir IMPRESS
2002) ont été décrites afin de construire des indices synthétiques de pressions calculables pour
l’ensemble des plans d’eau. Ces variables sont :
- la surface en zones urbaines sur le BV, exprimée en % de la surface totale du BV
(source : Corine Land Cover 2000),
- la surface en zones agricoles sur le BV, également exprimée en % de la surface du
BV (source : Corine Land Cover 2000),
- la densité de population humaine sur le BV, en nombre d’individus à l’hectare,
- la densité de population humaine sur une zone tampon de 1km autour du lac, en
nombre d’individus à l’hectare,
- la densité de réseau de communication sur le BV, exprimée en km/km2,
- les surplus en azote (NTotal) et en phosphore (PTotal) sur le BV (source : NOPOLU –
PÖYRY Environment), exprimés en tonnes/km2,
- le bilan en azote sur le BV, exprimé en tonnes/km2, qui correspond à la somme des
surplus en azote et des dépôts atmosphériques en azote d’origine anthropique,
- l’aléa moyen d’érosion couplé aux surplus en PTotal du BV (source : INRA – Ifen –
GIS Sol, 2000). En effet, comme le phosphore est un élément chimique peu mobile
(Carpenter et al. 1998b), et comme le préconise Coale (2000), nous avons
multiplié le surplus en PTotal par l’aléa moyen d’érosion du BV. Ce dernier est un
coefficient adimensionnel intégrant la battance des sols, l’érodabilité, la
topographie, l’occupation du sol, les précipitations, ainsi que les zones où les
cultures sont prépondérantes
(http://erosion.orleans.inra.fr/alea_france_version2000/index.html).
Une analyse en composantes principales (ACP) a ensuite été réalisée sur ces neuf
variables. Deux nouvelles ACP ont ensuite été effectuées, l’une sur les variables liées à
l’agriculture, l’autre sur les variables liées à l’urbanisme.
Les coordonnées des lacs sur l’axe principal de ces ACP constituent les indices
synthétiques de pression. Ces deux indices sont ensuite sommés pour donner un indice global
de pression s’exerçant sur le BV, à chaque lac. Avant d’être log transformés, ces indices sont
translatés pour ne pas prendre de valeur négative. La gamme de variation des pressions a été
analysée au sein de chaque type de plans d’eau.
3.2.2.3. Réponse des métriques aux pressions

Les groupes typologiques regroupant le plus de sites avec des données poissons sont le
type RA3, i.e. 10 retenues peu profondes, à berges découpées et à très faible altitude et, le
type LN1, i.e. 16 lacs naturels plutôt profonds et de forme arrondie, situés à des altitudes
moyennes (Figure 3. 1b). La réponse des métriques aux pressions a été testée sur ces deux
types par régression linéaire entre les indices de pressions et les métriques (coefficient de
corrélation de Spearman (ρ), P < 0.05).
Toutes les analyses ont été réalisées sous « R » (Ihaka and Gentleman 1996, R
Development Core Team 2008).
- 46 -
3.3. Résultats
3.3.1. Typologie physique des plans d’eau français

La méthode des K-means permet d’identifier deux types de lacs naturels (Figure 3. 3) et
trois types de retenues (Figure 3. 4). Les caractéristiques environnementales moyennes de ces
cinq groupes de sites sont données dans le Tableau 3. 1 et représentées sur la Figure 3. 5.
Figure 3. 3: Identification des deux groupes de lacs naturels (1 : groupe LN1 ; 2 : groupe
LN2) par la méthode des K-means – analyse sur 34 lacs naturels.
Les axes horizontal et vertical correspondent, respectivement au premier et au
deuxième axe de l’ACP réalisée sur les sept variables environnementales
disponibles pour les 34 lacs naturels. Pour les numéros des plans d’eau, voir la
liste détaillée à l’annexe 2.
- 47 -
Figure 3. 4: Identification des trois groupes de retenues (1 : groupe RA1 ; 2 : groupe RA2 ;
3 : groupe RA3) par la méthode des K-means – analyse sur 104 sites.
Les axes horizontal et vertical correspondent, respectivement au premier et au
deuxième axe de l’ACP réalisée sur les sept variables environnementales
disponibles pour les 104 retenues. Pour les numéros des plans d’eau, voir la liste
détaillée à l’annexe 2.
Le premier groupe de lacs naturels (LN1) est constitué de 24 sites d’altitude, plutôt petits,
profonds et de forme plutôt arrondie (SLDF proche de 1). Le deuxième groupe de lacs
naturels (LN2) est constitué de 10 sites de très faible altitude, grands, peu profonds et peu
découpés, mais dont les superficies peuvent être très différentes. Ces deux types de lacs
naturels sont respectivement les plus froids et les plus chauds du jeu de données, les premiers
pouvant avoir des températures moyennes hivernales négatives. Ils se distinguent clairement
d’un point de vue géographique. Le premier groupe est constitué des plans d’eau des Alpes,
du Jura et du Massif Central, le deuxième des lacs des littoraux atlantique et méditerranéen.
Le premier groupe de réservoirs (RA1) est constitué de 20 plans d’eau caractérisés par des
altitudes intermédiaires, des surfaces et des profondeurs importantes, et des berges découpées
(SLDF élevés). Le deuxième groupe (RA2) est constitué de 35 plans d’eau situés également à
moyenne altitude mais de surface plutôt petite et de forme arrondie. Leur profondeur
moyenne est intermédiaire. Le troisième groupe (RA3) rassemble 49 plans d’eau de faible
altitude, de petite surface, à berges découpées, mais très peu profonds.
- 48 -
Les types RA1 et RA2 de retenues ne se distinguent pas forcément d’un point de vue
géographique, ni du point de vue des températures, par contre le type RA3 regroupe
majoritairement des retenues du bassin Loire/Bretagne, dont les températures hivernales et
estivales sont moins contrastées que dans les deux autres types de retenues.
Figure 3. 5: Représentation des principales caractéristiques environnementales naturelles des

cinq types de plans d’eau identifiés.
Surface du plan d’eau (SuperficiePla ; en ha) ; altitude du plan d’eau (Altitude ; en
m) ; profondeur maximale du plan d’eau (ProfondeurMaxPla ; en m) ; surface du
bassin versant (Surface_BV ; en km2) ; indice de développement des berges
(SLDF ; sans unité) ; somme des températures moyennes de l’air des mois de
janvier et de juillet (JanJuil ; en °C) ; différence des températures moyennes de
l’air entre le mois de juillet et le mois de janvier (Juil_Jan ; en °C).
- 49 -
3.3.2. Intensité des pressions
3.3.2.1. Analyse globale

L’ACP réalisée sur l’ensemble des variables de pression des 138 plans d’eau permet
d’identifier deux groupes de variables, les unes relatives à l’urbanisme, les autres en relation
avec l’agriculture (Figure 3. 6).
Figure 3. 6: Analyse en Composantes Principales (ACP) réalisée sur les neuf variables de
pressions anthropiques.
Surface en zones urbaines sur le BV (X.du.BV.Urbain) ; surface en zones
agricoles sur le BV (X.du.BV.Agri) ; densité de population sur le BV
(DensitePop_BV_1998) ; densité de population sur un buffer de 1km autour du lac
(DensitePop_Buff1000_1998) ; densité de réseau de communication sur le BV
(Densite_Reso) ; surplus en azote (NTotal) et en phosphore (PTotal) sur le BV
(SurplusN.T._Moy et SurplusP.T._Moy) ; bilan en azote sur le BV
(BilanN.T._Moy) ; aléa moyen d’érosion couplé aux surplus en Ptotal du BV
(SurplusP.T._Moy.1).
- 50 -
L’ACP réalisée sur les 5 variables de pressions agricoles (Figure 3. 7a) montre que les 138
plans d’eau se répartissent le long du premier axe qui explique 73 % de la variabilité. Il porte
la variable de transfert de phosphore et dans une moindre mesure les variables bilan et surplus
d’azote total. Globalement, les sites peu impactés par ces variables sont plus nombreux que
les plans d’eau avec des valeurs fortes. Néanmoins, toute la gamme des pressions
anthropiques est observée.
Les premier et deuxième axes de l’ACP, réalisée sur les 4 variables de pressions urbaines
disponibles pour les 138 plans d’eau, expliquent respectivement 70 % et 17 % de la variabilité
des sites (Figure 3. 7b). Le premier axe est fortement tiré par deux sites très impactés
[retenues de Grand-Large (n°101 sur la Figure 3. 7b) et de Torcy-Neuf (n°113 sur la Figure 3.
7b) avec, respectivement, 42 % et 55 % du BV urbanisé], qui s’opposent à l’ensemble des
autres plans d’eau. Un lac [Nantua (n°80, sur la Figure 3. 7b)] se distingue sur cet axe par son
important réseau de communication sur son BV.
(7a)
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(7b)
Figure 3. 7: Analyse en Composantes Principales (ACP) réalisée sur les cinq variables de
pressions agricoles (7a), et les quatre variables de pressions urbaines (7b).
3.3.2.2. Analyse par types

L’analyse par types, menée sur les 44 plans d’eau présentant des données poissons,
indique que l’intensité des pressions agricoles est comparable entre les groupes LN1 et LN2
des lacs naturels (respectivement, n=16 et n=3) et RA1 et RA2 des retenues (respectivement,
n=8 et n=7) (Figure 3. 8a). Au sein de ces 4 groupes, les intensités de pressions sont beaucoup
plus faibles que dans le groupe 3 des retenues (RA3 ; n=10), qui se démarque très nettement.
Dans ce dernier type, tous les sites sont plus impactés que ceux des autres types.
Les pressions urbaines sont aussi globalement assez faibles sur l’ensemble des sites, quel
que soit le groupe de plans d’eau considéré (Figure 3. 8b). On relève pourtant des valeurs très
élevées de pressions urbaines pour deux sites, à savoir Torcy-Neuf et Annecy, classés
respectivement dans le groupe 2 des retenues (RA2) et dans le groupe 1 des lacs naturels
(LN1).
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Le groupe 3 des retenues (RA3), qui se distingue par de fortes pressions agricoles,
présente également un indice de pressions globales élevé. De même, les deux sites fortement
marqués par les pressions urbaines sont très impactés lorsque l’on considère les pressions
totales (Figure 3. 8c).
Figure 3. 8: Représentation des trois indices synthétiques de pressions anthropiques pour

chacun des cinq types de plans d’eau identifiés : indice synthétique des pressions
agricoles (8a), indice synthétique des pressions urbaines (8b), indice de pression
global (8c).
3.3.3. Calcul des métriques piscicoles et réponse aux pressions

Vingt neuf espèces de poissons ont été recensées sur les seize lacs naturels du groupe LN1
et les 10 retenues du groupe RA3. Les caractéristiques des espèces piscicoles capturées sur
ces sites sont listées dans le Tableau 3. 2. La perche franche (Perca fluviatilis) et le gardon
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(Rutilus rutilus) sont les plus fréquents quel que soit le type de milieu et la plupart des espèces
sont communes aux deux types de systèmes. Inversement, dix espèces ne sont présentes que
dans les lacs naturels. Parmi ces espèces, on compte trois salmonidés (Salmo trutta fario,
Salvelinius alpinus et Salvelinius namaycush). La bouvière (Rhodeus amarus) et le mulet porc
(Liza ramada) n’ont été capturés que dans des retenues.
Dans les retenues du groupe 3, aucune des métriques calculées ne répond à l’indice de
pression urbaine. Par contre trois métriques répondent positivement aux pressions agricoles
qui s’exercent sur le bassin versant (Figure 3. 9, Tableau 3. 4): il s’agit de la richesse en
espèces phytophiles et des densités de poissons, exprimées en biomasse ou en effectifs. La
richesse en espèces phytophiles est la seule métrique répondant positivement à l’indice de
pression globale.
Tableau 3. 4: Corrélation de Spearman entre les métriques et les pressions au sein de chaque
type de plans d’eau : * significatif à p<0,05.
Indices synthétiques de pressions
Métriques
urbaines agricoles urbaines + agricoles
Groupe 3 des retenues (n = 10)

Richesse en phytophiles stricts -0,27 0,65* 0,65*
CPUE-Nb -0,50 0,65* 0,41
BPUE_gr -0,53 0,72* 0,52
Groupe 1 des lacs naturels (n = 16)
Richesse spécifique 0,56* 0,36 0,49
Richesse en espèces natives 0,53* 0,32 0,45
Richesse en espèces tolérantes 0,59* 0,6* 0,62*
Richesse en espèces planctivores 0.56* 0.43 0.50*
Richesse en espèces omnivores –
0,57* 0,49 0,58*
généralistes
Sur les lacs naturels du type un, cinq métriques répondent positivement aux pressions
urbaines. Il s’agit des richesses spécifiques totales, en espèces natives, en espèces tolérantes,
en espèces omnivores-généralistes et en espèces planctivores (Tableau 3. 4). Cependant, les
corrélations observées sur les richesses totales et en espèce natives sont fortement tirées par
un point (Figure 3. 10). Les richesses en espèces tolérantes, planctivores et omnivores-
généralistes sont aussi positivement corrélées à l’indice de pression totale. La richesse en
espèces tolérantes est la seule métrique qui répond aux trois indices de pression.
- 54 -
coefficient de corrélation) entre les métriques sélectionnées dans le type RA3 des
retenues et les indices synthétiques de pressions.
Avec Pr A : indice synthétique des pressions agricoles ; Pr U : indice synthétique
des pressions urbaines ; Pr all : indice de pression global ; R Phyto : nombre
d’espèces de poissons phytophiles ; CPUE-Nb : captures par unité d’effort de
pêche et BPUE-gr : biomasse par unité d’effort de pêche.
- 55 -
coefficient de corrélation) entre les métriques sélectionnées dans le type LN1 des
lacs naturels et les indices synthétiques de pressions.
Avec Pr A : indice synthétique des pressions agricoles ; Pr U : indice synthétique
des pressions urbaines ; Pr all : indice de pression global ; R Sp : richesse
spécifique ; R natives : nombre d’espèces de poissons natifs ; R tolerantes :
nombre d’espèces de poissons tolérants ; R planctivores : nombre d’espèces de
poissons planctivores et R omnivores : nombre d’espèces de poissons omnivores.
- 56 -
3.4. Discussion et conclusions
Ce travail constitue une étape exploratoire de développement d’un indicateur poisson pour
les plans d’eau français, réalisée à partir des données nationales disponibles fin 2008.
La typologie effectuée ici prend en compte des variables environnementales connues pour
expliquer la variabilité des communautés de poissons. Néanmoins, d’autres variables
importantes, vis-à-vis des propriétés physico-chimiques de l’eau, telles que la géologie et
l’hydrologie auraient dû être prises en compte (Rowan et al. 2003, Winfield & Durie 2004),
mais elles ne sont pas disponibles pour l’ensemble des sites. La typologie regroupe donc des
milieux encore hétérogènes par certains aspects environnementaux.
Sur les lacs naturels, la typologie a été effectuée sur les données de 34 plans d’eau, ce qui
représente la moitié des milieux concernés par la DCE (71 lacs, R. Lallemand, communication
personnelle) et deux types de milieux ont été identifiés. Ainsi, quelle que soit la
représentativité de nos milieux, il est évident que scinder les lacs français en groupes
homogènes va nous amener à des types avec des effectifs très faibles, et/ou à des types au sein
desquels va demeurer une forte variabilité environnementale, source de variabilité naturelle
des communautés piscicoles.
Pour les retenues, trois types de milieux ont été identifiés avec notre jeu de données qui
représente moins d’un tiers de l’ensemble des milieux artificiels français. En supposant que
notre échantillonnage soit représentatif de la diversité des sites nationaux, il est vraisemblable
que les retenues non considérées ici se répartissent au sein des types existants. Dans le cas
contraire, quelques nouveaux types pourraient être identifiés. Néanmoins, compte tenu que
cette typologie n’intègre pas de variable hydrologique, telle que le temps de renouvellement
des masses d’eau par exemple, il est probable que la variabilité intra-type demeure forte.
Il faut également noter qu’en raison de la prise en compte de variables environnementales
mieux adaptées à l’ichtyofaune, dans la méthode utilisée ici pour identifier des types plus
homogènes, ces derniers ne recoupent pas parfaitement ceux identifiés dans la typologie
nationale (Ministère de l'Ecologie et du Développement Durable 2005).
Les milieux ont ensuite été caractérisés par les pressions (ou forces motrices, voir
IMPRESS 2002) qui s’exercent sur les bassins versants et qui sont les seules données
homogènes disponibles pour l’ensemble des sites. Ainsi, les données de pressions analysées
reflètent bien l’ensemble des sources de pollution chimique, et, partiellement, les pressions
s’exerçant sur la morphologie des lacs. En effet, la densité de population et la densité du
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réseau routier sont de bons indicateurs de la fréquentation des milieux (Forman & Alexander
1998, Jones et al. 2000, Pitt et al. 1995).
Il faut noter que la variabilité des pressions agricoles et urbaines au sein de la plupart des
types est très faible. De plus, elle n’est pas comparable sur l’ensemble des types. Dans
quelques cas (types RA2 et LN1), des pressions urbaines très fortes sont mesurées sur
quelques sites isolés, sans pour autant que la gamme de pressions soit couverte de manière
homogène par l’ensemble des plans d’eau. Si elles devaient être confirmées, ces distributions
hétérogènes de sites plus ou moins impactés, au sein des types et entre les types, pourraient
poser des problèmes quant à la définition de seuils homogènes de qualité des métriques à
l’échelle nationale.
Le développement de bioindicateurs suppose que l’on dispose de mesures sur des sites pas
ou peu impactés, ces sites servant de base de référence aux métriques sélectionnées pour leur
réponse aux pressions (Noble et al. 2007). Cette approche typologique suppose donc
l’établissement de conditions de référence au sein de chaque type. Nous avons montré que,
comparativement aux autres, le type 3 des retenues ne présentait pas de site peu impacté ; par
conséquent, nous constatons que ceci constitue une limite au développement d’un indice
poisson.
Une condition supplémentaire à la sélection de métriques réside dans l’obtention d’une
gamme assez large et complète de pressions. Or, nous avons montré que ceci n’était le cas
pour aucun des types et aucune des pressions, hormis pour les pressions agricoles du type
RA3, type pour lequel se posait le problème de l’absence de référence.
De plus, la gamme de variations des pressions anthropiques doit être à peu près identique
au sein de chaque type, sans quoi le système de notation ne pourra pas être harmonisé. Même
si ces gammes de variations des pressions anthropiques se chevauchent souvent entre types,
cette condition n’est pas remplie, notamment si l’on considère les quelques milieux très
perturbés du types 1 des lacs naturels et du type 2 des retenues.
D’autres pressions s’exercent directement sur les systèmes et sont à même d’influencer
l’ichtyofaune. C’est le cas par exemple des variations de niveau d’eau (Argillier et al. 2004,
Poirel et al. 1996). L’exercice conduit ici devra donc être élargi à l’ensemble des pressions, en
particulier à la mesure de l’altération de l’hydromorphologie des systèmes et des usages
associés.
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L’analyse de la sélection des métriques repose sur les observations réalisées sur les types
de milieux les mieux représentés. Malgré cela, le nombre de sites reste très faible, en
particulier sur les retenues. Sur ces types avec un petit nombre d’échantillons, les résultats
sont très dépendants des données. Cet exercice de sélection de métriques doit donc être
considéré comme une étape exploratoire.
Nous avons montré qu’il était possible d’identifier des métriques élaborées à partir des
caractéristiques de communautés piscicoles, répondant aux pressions anthropiques.
Finalement, au sein de chacun des deux groupes de plans d’eau étudiés, la plupart des
corrélations ne sont observées qu’avec des pressions dont la gamme de variation est la plus
forte, à savoir, les pressions agricoles dans le type RA3 et les pressions urbaines dans le type
LN1. Selon la distribution des gammes de pressions au sein des types, l’agrégation des
pressions n’est donc pas forcément une étape intéressante pour l’élaboration des indices.
Dans le groupe des retenues, toutes les métriques sont positivement corrélées à
l’agriculture. La réponse des densités de poissons (en effectif ou en biomasse) aux pressions
est assez contrastée selon les études. Une diminution des densités piscicoles avec
l’augmentation des pressions a été reportée sur les retenues des Etats-Unis (Hickman &
McDonough 1996) ; inversement, sur les lacs naturels d’Europe, c’est une augmentation qui
est mesurée (Pedron et al. 2010), conformément à ce que nous observons ici. L’augmentation
du nombre d’espèces phytophiles strictes avec l’agriculture n’a pas été documentée. On aurait
pu s’attendre à une diminution du nombre d’espèces, dont la reproduction est liée à la qualité
des habitats littoraux, avec l’augmentation des pressions du bassin versant. Ce résultat met en
avant la limite du jeu de données dans ce type d’approche.
Sur les lacs naturels, le nombre de métriques potentiellement intéressantes est beaucoup
plus important que sur les retenues. Toutefois, parmi les neuf corrélations significatives,
celles mesurées entre la richesse spécifique totale ou la richesse spécifique en espèces natives
et la pression urbaine sont douteuses du fait de la distribution des valeurs mesurées.
Inversement, l’analyse graphique des sept autres corrélations renforce les résultats issus des
analyses statistiques.
L’inclusion de métriques liées à la tolérance est très fréquente dans les indices poisson
(Jennings et al. 1999b, Whittier 1999) et la relation positive observée ici avec toutes les
pressions va dans le sens attendu selon la littérature. Bien que très répandue, l’utilisation de
cette métrique est discutable ; en effet, la bonne réponse aux pressions peut être attribuée au
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mode de construction de cette métrique, la tolérance des espèces étant définie, précisément, à
partir de leur fréquence d’observation en milieux perturbés !
La réponse des espèces omnivores-généralistes aux pressions est également bien
documentée et les résultats obtenus sont en cohérence avec la plupart des observations (Drake
& Pereira 2002, Schulz et al. 1999).
Pour conclure, ces résultats mettent en avant un certain nombre de limites à l’approche
typologique afin de développer un indice harmonisé au niveau national. Le faible nombre de
lacs naturels par type limite la pertinence des corrélations potentielles entre les métriques et
les pressions. De plus, les différences inter- et intra-types de gradients de pressions rend
difficilement envisageable une approche harmonisée au niveau national. Il conviendra de
vérifier ces limites lorsque plus de données seront disponibles. Si elles devaient être
confirmées, d’autres approches, notamment par modélisation sur l’ensemble des sites,
devraient être envisagées.
Au contraire, lorsque le jeu de données est significatif, il est possible de définir certaines
métriques relatives à l’ichtyofaune répondant de manière cohérente à des pressions d’origines
agricole ou urbaine. Ces premiers résultats sont donc très encourageants quant à la possibilité
d’utiliser l’ichtyofaune pour rendre compte de perturbations anthropiques, ceci malgré l’idée
fréquemment avancée que ces communautés piscicoles des plans d’eau sont très manipulées
par l’Homme et qu’elles reflètent donc peu les conditions environnementales.
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CHAPITRE 4
Sélection de métriques piscicoles et variabilité environnementale : comparaison entre

lacs naturels et réservoirs artificiels
Dans le domaine de la bioindication en milieux lacustres, quelques travaux mettent en
exergue la spécificité des milieux d’origine artificielle (de Freitas Terra & Araujo 2011,
Gelwick & Matthews 1990, Irz et al. 2006). Par exemple, comme le notent Jennings et al.
(1995) et Karr and Dionne (1991), l’âge moyen relativement faible des réservoirs, par
comparaison à celui des milieux naturels, fait que la notion d’intégrité biotique s’adapte mal
aux réservoirs. Aux Etats-Unis, cette distinction apparaît dans la dénomination des indices.
Pour les réservoirs, il est fait référence à des indices de peuplements [Reservoir Fish
Assemblage Index (RFAI)] (Hickman & McDonough 1996, McDonough & Hickman 1999).
De plus, ces auteurs montrent que des ensembles de métriques spécifiques à chaque type de
système, lacs naturels ou réservoirs, sont à utiliser pour la bioindication.
En France, Irz et al. (2006), et de Freitas Terra and Araujo (2011) au Brésil, mettent en
évidence que les communautés piscicoles des retenues présentent des caractéristiques
intermédiaires entre celles des lacs naturels et des cours d’eau. Elles abritent à la fois des
espèces inféodées aux milieux lotiques et des espèces inféodées aux milieux lentiques. On
peut donc supposer que les indicateurs doivent tenir compte de l’origine des systèmes.
Compte tenu de ces éléments, nous avons cherché à savoir si dans le contexte national, les
mêmes métriques pouvaient être utilisées pour rendre compte de l’état écologique des lacs
naturels et des retenues artificielles.
D’un point de vue méthodologique, nous avons exposé dans le chapitre précédent un
certain nombre de limites à l’approche typologique dans le contexte national. Nous avons
donc envisagé ici une solution alternative pour exercer un contrôle de la réponse des
communautés de poissons aux gradients environnementaux naturels. Ce type d’approche a
déjà fait ses preuves dans le développement des indices poisson pour les rivières (Oberdorff et
al. 2002, Pont et al. 2009, Pont et al. 2007).
L’article suivant vise donc à montrer comment, par modélisation, il est possible
d’identifier des métriques piscicoles qui répondent à une pression telle que l’enrichissement
en nutriments et/ou les types d’occupation du sol sur le bassin versant des lacs naturels et des
retenues artificielles français, en contrôlant l’effet propre aux gradients environnementaux
naturels.
L’analyse statistique mise en oeuvre consiste à étudier l’effet relatif de l’environnement et
des pressions anthropiques sur la variabilité de la structure des peuplements :
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- en modélisant les métriques en fonction des facteurs environnementaux et des
pressions, à l’aide de régressions multiples ;
- et en utilisant la méthode de partitionnement de la variance (Borcard et al. 1992,
Legendre & Legendre 1998) afin d’évaluer la part de la variabilité des métriques
piscicoles qui est due, soit à l’environnement exclusivement, soit aux pressions
anthropiques exclusivement, ou bien à leurs effets conjoints (Figure 4. 1).
Figure 4. 1 : Partitionnement de la variance d’une métrique biologique en quatre

composantes : [a] fraction exclusivement liée aux pressions anthropiques, [b]
fraction conjointe entre pressions anthropiques et environnement, [c] fraction
exclusivement liée à l’environnement et [d] fraction inexpliquée. Adapté de
Legendre & Legendre (1998).
Cette méthode est très utile lorsque l’on souhaite quantifier globalement les parts de
variabilité liées à plusieurs ensembles de prédicteurs, afin de mettre en évidence ce qui est
purement attribuable à un ensemble et ce qui est indistinctement attribuable à plusieurs
ensembles (Borcard et al. 1992, Legendre & Legendre 1998).
Comme dans l’analyse typologique présentée au chapitre précédent, les résultats obtenus
sont impactés par la faiblesse du jeu de données, en particulier pour les lacs naturels.
Toutefois, ces résultats permettent de dégager des métriques poissons répondant à
l’anthropisation des bassins versants des plans d’eau, lorsque la variabilité naturelle est prise
en considération.
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Sans détailler les résultats publiés, il s’avère également que les métriques à même de
rendre compte des perturbations ne sont pas les mêmes dans le cas des lacs naturels et des
retenues, ce qui va dans le sens des études citées précédemment. Ceci suggère la nécessité de
développer, en France, des bioindicateurs différents pour ces deux types de milieux. De plus,
cela confirme les résultats de Irz et al. (2006), à savoir que les lacs naturels ne peuvent pas
servir a priori comme références à atteindre pour les retenues.
Selecting fish-based metrics responding to human pressures in French natural lakes and
reservoirs: towards the development of a fish-based index (FBI) for French lakes
(Ecology of Freshwater Fish 2011: 20: 120-132 - DOI: 10.1111/j.1600-0633.2010.00467.x).
Lionel Launois, Jacques Veslot, Pascal Irz and Christine Argillier
4.1. Introduction
Frey (1977) and then Karr and Dudley (1981) defined the concept of ecological integrity
of freshwater ecosystems as their ability to support and maintain “a balanced, integrated,
adaptive community of organisms having a species composition diversity and functional
organization comparable to that of the natural habitat of the region.” Since this publication,
fish assemblages have become accepted as reflecting the “overall integrity of biological
communities” and as “excellent indicators of the environmental quality of aquatic
ecosystems” (Fausch et al. 1990).
Following Karr’s studies on Midwestern U.S. streams, many other multimetric fish-based
indices of biotic integrity (IBIs) have been developed for the running waters of North and
Central America, and Europe (Hughes & Oberdorff 1999, Miller et al. 1988, Roset et al.
2007, Simon & Lyons 1995).
Conversely, the development of IBIs for lakes remains scarce (Appelberg et al. 2000,
Drake & Pereira 2002). The main difficulty hampering this development was heterogeneity in
fish sampling (Jackson & Harvey 1997). Even scarcer are the attempts to assess the quality of
reservoirs using fish assemblages (but see Hickman and McDonough 1996; Jennings et al.
1995), despite their importance for water resource managers (Jennings et al. 1995). One
reason put forward was that reservoirs are artificial systems that lack natural reference sites to
assess the deviation from unaffected natural conditions (Hickman & McDonough 1996;
Jennings et al. 1995).
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In Europe, the implementation of the European Water Framework Directive (WFD)
(European Community 2000) requires the use of fish-based tools to assess the ecological
status of all lentic water bodies, i.e., natural lakes and reservoirs, over 50 ha. The first multi-
metric index for lake fish was developed in Sweden (Appelberg et al. 2000; Holmgren et al.
2007). However, completed fish-based indices still remain rare and concern almost solely
natural lakes. Moreover, they generally focus on specific types of environments (e.g.,
mountain or wetland lakes) and they are often based on taxonomic metrics, which are poorly
transferable due to limited fish species distribution areas. In this context, the development of
methods for monitoring the biological responses of fish assemblages to natural and
anthropogenic factors is essential and remains to be improved. This is particularly true in
some countries (e.g., France, Spain, Portugal, the Czech Republic) where most lakes are
manmade.
Reservoirs are frequently considered as an intermediate between rivers and lakes (Gelwick
& Matthews 1990, Irz et al. 2006, Wetzel 1990): however, biological cross-ecosystem studies
are currently not common (Pace 1991). Nevertheless, reservoirs satisfy some of the definition
criteria of lakes (Politou et al. 1993), because most of the major processes (e.g., internal
mixing, nutrient uptake, predator–prey interactions) occur in both natural lakes and reservoirs
(Thornton 1990). Currently, the question of whether a common fish-based index could be
used for both lentic ecosystems has been raised.
Therefore, the goals of this study were (i) to test if some candidate metrics, including
functional metrics, are correlated to anthropogenic pressure gradients when the environment
is controlled for in each type of lentic ecosystem and (ii) to compare the selected metrics
between natural lakes and reservoirs.
4.2. Study Sites

Throughout this article we refer to lakes of natural origin as natural lakes and to man-
made lentic systems as reservoirs; the generic term ‘lakes’ is used for both types. Only
freshwater lakes are included in the present study.
Anthropogenic pressure indices were computed using 40 natural lakes and 106 reservoirs
located throughout France. Fish data were collected from 30 natural lakes and 59 reservoirs
among these 146 lakes (Figure 4. 2). The lake data set represented a wide range of each
environmental condition (Tableau 4. 1), but as demonstrated in previous works on French
lakes (Argillier et al. 2002b, Irz et al. 2004a, Irz et al. 2002), sites with altitudes over 1500
- 67 -
meters above sea level had to be excluded from these analyses because they shelter only
introduced fish species.
Figure 4. 2: Location of the 89 study sites in the French hydrographic network.

◊ Natural lakes ♦ Reservoirs.
Tableau 4. 1: Environmental parameter mean values and extreme values of the 89 study sites.
Reservoirs (n=59) Natural lakes (n=30)
Parameter Mean Range Mean Range
Geologic Index (GI) (% of the drainage
basin as calcareous) 0.26 0~1 0.38 0~1
Residence Time (RT) (day) 137.09 0.11~922.80 692.08 5~5095
a
Water level maximal amplitude (WL) 1.53 0~2 0.45 0~2
2
Catchment area (ADB) (km ) 662.23 8.42~10533.34 152.80 1.25~1033.34
Lake area (AL) (ha) 251.00 44~2272 781.00 13~5768
Maximum depth (Dmax) (m) 23.87 2~100 23.13 2~110
b
Shoreline development factor (SLDF) 3.34 1.01~9.20 1.62 1.01~3.80
T°C mean Jan + T°C mean Jul (SumT) 23.86 15.49~31.27 20.44 13.18~31.51
T°C mean Jul - T°C mean Jan (DiffT) 15.97 10.43~19. 42 17.19 13.48~19.91
c
Volume development (Vd) 1.21 0.37~2.96 1.42 0.30~2.66
b
3 classes: 0 (no variation), 1 (≤3m), 2(>3m)
b
Calculated as: perimeter / (√4π×surface).
c
Calculated as: 3 × Lake volume / (Maximum depth × Lake area).
- 68 -
4.3. Material and methods
Environmental parameters. – Five large-scale environmental parameters related to lake and
drainage basin morphology, and known to affect the distribution patterns of fish assemblages,
were considered (Tableau 4. 1). The lake area (AL) is a strong predictor of fish richness
(Barbour & Brown 1974, Eadie & Keast 1984); the drainage basin area (ADB) is an indicator
of habitat diversity upstream of lake and of inflows into lakes (Irz et al. 2004b); the maximum
depth (Dmax) acts both directly on fish habitat availability by determining the importance of
the lake littoral zone (Argillier et al. 2002c) and indirectly on the lake water dissolved oxygen
level (Hondzo & Stefan 1996); the lake shoreline development factor (SLDF) (Eadie & Keast
1984) accounts for an indicator of habitat diversity into lakes; and the volume development
(Vd) (Håkanson 1981) gives information on the lake basin shape and therefore on the size of
the lake littoral zone. The AURELHY climate model (Benichou & Le Breton 1987) was used
to compute January and July mean air temperatures, because air temperatures are known to
control the distribution of fish species (Daufresne & Boet 2007, Irz et al. 2007, Mason et al.
2008) by affecting the metabolism (Gillet 1991) and the growth of juveniles (Jensen 1990,
Staggs & Otis 1996) (SumT; DiffT; see Tableau 4. 1). The hydrological model developed by
Pella et al. (2006) was used to evaluate the lake yearly inflow in order to calculate residence
time (RT). A geologic index (GI) displaying the percentage of the catchment area as
calcareous, based on 1/50,000 geological maps (BRGM: http://www.brgm.fr/cartegeol.jsp),
was computed on the whole lake catchment area. This GI enables us to understand lake
catchment chemistry, which is an important factor affecting fish abundance (Shaw et al.
2004). Finally, the water level maximum amplitude (WL) was considered as an additional
hydrological parameter, due to its detrimental effect on fish feeding conditions, sheltering and
reproduction (Bragg et al. 2003, Winfield 2004).
Anthropogenic pressures. – Anthropogenic pressures were assessed for the overall French
lake data set (n = 146) using seven variables derived from features of the lakes’ catchments
(Tableau 4. 2). Land cover percentages allow evaluation of the agricultural and urban land use
(Drake & Pereira 2002). Population density relates to the watershed dynamic, especially
chemical discharges into lakes (see Pitt et al. 1995). The network density of roads and
railways reflects fragmentation (Forman & Alexander 1998, Jones et al. 2000). The soil
surface nitrogen balance and phosphorus balance were simulated using the NOPOLU System
2 model (PÖYRY) (LeGall, personal communication). These are the differences between the
total quantity of nutrients entering the soil and the quantity leaving it annually. The nitrogen
- 69 -
balance was kept as a pressure variable, but the phosphorus balance was adjusted according to
the recommendation of Coale et al. (2002). As particulate phosphorus is not very mobile
(Carpenter et al. 1998b), Coale proposed to multiply the phosphorus balance by the mean
value of the lake’s watershed soil erosion after log transformation (log(x+1)). This watershed
soil erosion was estimated with a model developed by the French National Institute for
Agricultural Research (INRA – Ifen – Gis Sol, 2000 –
http://eusoils.jrc.ec.europa.eu/ESDB_Archive/serae/grimm/erosion/inra/europe/analysis/maps
_and_listings/web_erosion/index.html). This model is based on soil slumping, erodibility,
topography, land cover, rainfalls and dominant cultivated land cover.
We assume that these seven variables derived from GIS data reflected the anthropogenic
pressures undergone by lakes at the catchment scale. The population density and agricultural
land cover were log-transformed (log(x+1)) and the percentage of the urbanised catchment
area was transformed to arcsine√(X). The other four anthropogenic variables were kept raw.
In order to reduce dimensionality, a standardised principal component analysis (PCA) was
performed on each group of anthropogenic variables (Tableau 4. 2). On the one hand, we
carried out a PCA on urban variables, and lake coordinates on the first principal component
were considered a synthetic urban pressure variable (PU). The same was done on agricultural
variables in order to create a synthetic agricultural pressure variable (PA).
- 70 -
Tableau 4. 2: Description of the anthropogenic variables included in the analysis; and principal component analysis (PCA) carried out on the
watershed pressure variables.
Table entries are the variable correlations with the first principal component on each of the two PCAs, one carried out on agricultural
variables only (PA) and another carried out on urban variables only (PU). Those loading most heavily on each PC are in bold.
All lakes (n = 146) Reservoirs (n=59) Naturals (n=30)
PA PU Mean Range Mean Range
Variable Inertia 63% 72%
% catchment agricultural 0.84 - 51.41 0 ~ 93.80 24.30 0 ~ 87.50
2
Agriculture land cover area (km ) 0.65 - 427.06 0 ~ 8783.40 54.35 0 ~ 790.60
Nitrogen balance (t) 0.79 - 122.19 -598.20 ~ 877.10 -40.44 -346.80 ~ 432.30
Phosphorus balance (t) * erodibility coefficient 0.88 - 421.71 -156.60 ~ 2523.10 -12.95 -389.10 ~ 849.20
% catchment urban - 0.90 3.95 0 ~ 55 3.40 0 ~ 15.30
2
Population density in catchment (people/km ) - 0.86 65.33 4.10 ~ 1006.50 40.02 3.70 ~ 260.50
Transportation network density - 0.78 2.17 0 ~ 10.30 2.09 0 ~ 11.30
Fish sampling procedures. – Fish data for the 89 French lakes were collected between 2005
and 2009 using a standardised method (C.E.N. 2005b). A stratified random sampling scheme
was implemented during summer (June–October, surface water temperature over 15°C). The
sampled lakes were divided into depth strata and randomly located samplings were performed
within each depth stratum. Both pelagic and benthic multi-mesh gillnets made out of
uncolored nylon were used. Benthic nets were 30 m long and 1.5 m deep and composed of 12
different mesh sizes (the panel for each mesh size was 2.5 m long) ranging between 5 mm to
55 mm following geometry series, with an approximately 1.25 ratio between mesh sizes. The
pelagic nets were 27.5 m long and 6 m high and had the same mesh sizes without the 5 mm
mesh size. The number of nets increased with lake depth and area. Nets were set around 7
p.m. for approximately 12 h. Fish were identified to species and counted. The method
provides the number of fish that were caught and the biomass for each one. We then
calculated a whole-lake estimate for species relative abundances (RA), expressed as the ratio
of the number of individuals belonging to a fish species to the total number of individuals in
the lake; catch per unit effort (CPUE), and biomass (in grams) per unit effort (BPUE)
(including both the benthic and the pelagic nets), respectively expressed as the overall number
of individuals or biomass captured divided by the unit-of-effort (i.e., nets’ surfaces multiplied
by the sampling duration). Finally, this method provided fish data that were comparable over
time within a lake and between lakes.
Candidate metrics. – Altogether, 42 species were caught in these 89 lakes (Tableau 4. 3).
Tableau 4. 3: Occurrence rates of the 42 fish species in natural lakes and reservoirs.
Rates over 75% are in bold.
Frequency Average Relative Average Relative
Frequency
Common name Species Reservoirs Abundance Abundance
Naturals (%)
(%) Reservoirs (%) Naturals (%)
Bream Abramis brama 76.67 51.61 9.25 4.87
Sterlet Acipenser ruthenus 1.67 0.00 0.00 0.00
Alburnoides
Spirlin 0.00 3.23 0.00 0.02
bipunctatus
Bleak Alburnus alburnus 53.33 29.03 2.05 2.25
Brown Bullhead Ameiurus melas 28.33 29.03 7.13 1.18
European eel Anguilla anguilla 1.67 3.23 0.01 0.00
Stone Loach Barbatula barbatula 5.00 6.45 0.22 0.10
Barbel Barbus barbus 6.67 0.00 0.04 0.00
White Bream Blicca bjoerkna 66.67 22.58 8.05 3.46
Carassius auratus
Goldfish 8.33 0.00 0.01 0.00
auratus
Crucian Carp Carassius carassius 11.67 9.68 0.03 0.10
Common Whitefish Coregonus lavaretus 3.33 32.26 0.03 1.61
Bullhead Cottus gobio 1.67 6.45 0.02 0.04
Common Carp Cyprinus carpio 38.33 19.35 0.23 0.03
Northern Pike Esox lucius 58.33 67.74 0.35 0.27
Mosquitofish Gambusia affinis 1.67 0.00 0.01 0.00
Gudgeon Gobio gobio 15.00 32.26 2.07 1.07
Gymnocephalus
Ruffe 63.33 38.71 6.08 3.17
cernuus
Hypophthalmichthys
Silver carp 1.67 0.00 0.01 0.00
molitrix
Pumpkinseed Lepomis gibbosus 30.00 32.26 0.19 0.23
Belica Leucaspius delineatus 3.33 3.23 0.00 0.25
Ide Leuciscus idus 1.67 0.00 0.00 0.00
Common Dace Leuciscus leuciscus 3.33 9.68 0.03 0.03
Golden grey mullet Liza aurata 1.67 0.00 0.00 0.00
Thinlip Mullet Liza ramada 1.67 0.00 0.12 0.00
Burbot Lota lota 0.00 3.23 0.00 0.00
Rainbow Trout Oncorhynchus mykiss 10.00 9.68 0.83 0.42
European Perch Perca fluviatilis 93.33 93.55 21.95 28.72
Eurasian Minnow Phoxinus phoxinus 5.00 3.23 0.41 1.81
Bitterling Rhodeus amarus 3.33 0.00 0.03 0.00
Roach Rutilus rutilus 91.67 96.77 31.31 38.85
Freshwater Blenny Salaria fluviatilis 0.00 3.23 0.00 0.01
Brown Trout Salmo trutta fario 15.00 16.13 1.13 0.65
Charr Salvelinus alpinus 0.00 9.68 0.00 0.78
Lake Trout Salvelinus namaycush 0.00 3.23 0.00 0.04
Pike Perch Sander lucioperca 78.33 41.94 4.41 1.83
Scardinius
Rudd 81.67 70.97 3.39 5.88
erythrophthalmus
Wels Catfish Silurus glanis 21.67 9.68 0.05 0.01
European Chub Squalius cephalus 21.67 38.71 0.44 1.06
Western vairone Telestes souffia 0.00 3.23 0.00 0.79
Grayling Thymallus thymallus 1.67 0.00 0.02 0.00
Tench Tinca tinca 25.00 35.48 0.09 0.45
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Ten biological traits (Tableau 4. 4) derived from functional guilds were used: two from
tolerance guilds (overall tolerant (OTol) and overall intolerant (OIntol)), three from
reproductive guilds (lithophilic species (Lith), strict lithophilic species (StLith), and strict
phytophilic species (StPhyto)), and five from trophic guilds (omnivorous (Omni),
invertivorous (Invert), planktivorous (Plankt), herbivorous (Herb) and piscivorous species
(Pisc)). The 42 fish species caught were assigned to these attributes (Appendix 3), based on
previous grey or published literature, online resources (Froese & Pauly 2009 –
http://www.fishbase.org/; Pont et al. 2006) or expert judgments (H. Persat, O. Schlumberger
& N. Poulet, personal communications). In addition, native species (Nat) were distinguished
using Keith & Allardi’s (2001) classification.
Tableau 4. 4: Proposed guild classification for French lake fishes.

Guilds Definitions
Overall Tolerants Species sensitive to any common impact related to altered
flow regime, nutrient regime, habitat structure and water
Tolerance guild
chemistry. Loss of intolerant species is a response to
(Karr et al. 1986) Overall Intolerants
degradation, whereas the number of tolerant species will
tend to increase with disturbance.
Can reproduce on unsilted mineral substrates whose

Lithophilic
Reproduction guild larvae are photophobic.
(Balon 1975; Dycus & Strict Lithophilic Need mineral substrates to reproduce.
Meinert 1994) Need plant substrates to reproduce and whose larvae are
Strict Phytophilic
not photophobic.
Feed on both animal and vegetal items (more than 25%

Omnivores
plant material and more than 25% animal material).
Trophic guild Feed at least partially on invertebrates/insects (more than
Invertivores
(Bruslé & Quignard 2001; 75% macro-invertebrates in the diet).
Dycus & Meinert 1994; Feed mainly on zooplankton and/or phytoplankton (more
Planktivores
Goldstein & Simon 1999; than 75% of plankton in the diet).
Pont et al. 2006) Feed on plant items (more than 75% of plant material in the
Herbivores
diet).
Feed on fish at least partially when adults (more than 75%
Piscivores
fish in the diet).
A set of candidate metrics was derived from species richness and faunal composition as in
other lake and stream IBIs (Drake & Pereira 2002; Holmgren et al. 2007; Karr et al. 1986;
Minns et al. 1994; Schulz et al. 1999). These metrics (Tableau 4. 5) were expressed as (i)
number of species sharing a trait (Sp_trait), (ii) number of species sharing a trait divided by
the total number of species in the lake (%Sp_trait), (iii) number of individuals sharing a trait
caught by unit effort (Nb_trait), (iv) number of individuals sharing a trait divided by the total
number of individuals in the lake (%Nb_trait), (v) biomass of individuals sharing a trait
- 74 -
caught by unit effort (B_trait) or (vi) biomass of individuals sharing a trait divided by the
total biomass of individuals in the lake (%B_trait). The total species richness (RS) metric was
calculated, and three diversity indices based on fish abundances in each sampled site – i.e.,
Shannon’s index (SWI) (Shannon & Weaver 1949), Simpson’s index (SI) (Simpson 1949)
and evenness index (EI) (Hill 1973) – were also considered, as well as BPUE, CPUE and the
average biomass (AB) calculated for each lake sampled as the ratio of BPUE to CPUE
(Tableau 4. 5).
- 75 -
Tableau 4. 5: Mean values (standard deviation) of the fish-based metrics and p-values of Student t-test (with Welch correction if needed)
obtained from the functional traits between natural lakes and reservoirs; P ≤ 0.05*, P ≤ 0.01**, P ≤ 0.001***.
Reservoirs (n=59) Naturals (n=30)
Proportion Proportion in Proportion Proportion in
Species Species
Functional traits CPUE in number of BPUE biomass of CPUE in number of BPUE biomass of
Richness Richness
individuals individuals individuals individuals
Overall Species
- - - - 9.50 (2.60) - - - - 8.40 (2.50)
Richness
Shannon index - - - - 1.29 (0.32)* - - - - 1.15 (0.21)*
Simpson index - - - - 0.63 (0.13) - - - - 0.58 (0.10)
Evenness index - - - - 0.59 (0.13) - - - - 0.56 (0.11)
Natives 0.08 (0.09) 0.66 (0.32) 3.30 (2.70) 0.62 (0.30) 5.70 (2.40) 0.06 (0.10) 0.78 (0.35) 2.40 (2.20) 0.74 (0.29) 5.70 (2.60)
Intolerants 0.00 (0.00) 0.01 (0.07) 0.04 (0.20)* 0.02 (0.09)** 0.30 (0.60)* 0.00 (0.01) 0.04 (0.08) 0.2 (0.30)* 0.09 (0.15)** 0.80 (1.10)*
3.90 5.40 2.40 4.20
Tolerants 0.09 (0.09) 0.80 (0.23) 0.74 (0.20)* 0.08 (0.11) 0.80 (0.21) 0.64 (0.20)*
(2.70)** (1.90)** (2.10)** (1.70)**
4.20 2.60
Lithophilic 0.09 (0.09) 0.88 (0.19) 0.83 (0.17) 6.60 (2.00) 0.08 (0.12) 0.91 (0.14) 0.79 (0.23) 6.20 (2.20)
(2.60)** (2.00)**
Strict lithophilic 0.00 (0.02) 0.10 (0.19) 0.50 (0.80) 0.11 (0.19) 1.10 (1.00) 0.00 (0.01) 0.08 (0.17) 0.30 (0.40) 0.13 (0.20) 1.70 (1.50)
Strict phytophilic 0.02 (0.04) 0.12 (0.19) 0.90 (1.10) 0.16 (0.16) 2.80 (1.30)* 0.01 (0.02) 0.08 (0.14) 0.80 (1.60) 0.21 (0.23) 2.20 (1.20)*
Omnivores 0.05 (0.05) 0.47 (0.22) 2.90 (2.10)* 0.55 (0.19) 3.90 (1.50)* 0.06 (0.10) 0.54 (0.26) 2.00 (1.70)* 0.51 (0.21) 3.20 (1.20)*
0.95 0.98
Invertivores 0.10 (0.09) 4.40 (2.70)* 0.86 (0.10) 8.10 (2.30) 0.09 (0.12) 3.10 (2.30)* 0.90 (0.11) 7.30 (2.20)
(0.05)** (0.03)**
Planktivores 0.06 (0.07) 0.55 (0.26) 3.40 (2.60)* 0.62 (0.20) 5.00 (1.90)* 0.06 (0.11) 0.57 (0.26) 2.20 (1.90)* 0.58 (0.20) 4.20 (1.60)*
Herbivores 0.03 (0.04) 0.34 (0.22) 1.30 (0.90) 0.28 (0.17) 1.60 (0.80) 0.04 (0.07) 0.41 (0.21) 0.90 (0.80) 0.27 (0.13) 1.80 (0.90)
Piscivores 0.04 (0.06) 0.36 (0.25) 1.60 (1.20) 0.35 (0.19) 3.60 (1.20) 0.03 (0.04) 0.36 (0.24) 1.30 (1.00) 0.44 (0.21) 3.60 (1.20)
All functional fish traits considered here are shared by at least three species. Moreover, as
in other studies (Irz et al. 2008, Jennings et al. 1995, Pont et al. 2007), introduced species
were not excluded from the metrics calculations because they may play an important
functional role in lake ecosystems.
This set of metrics reflects complementary aspects of assemblage richness and
functioning.
Statistical analysis. – For environmental variables, to retrieve less skewed distributions, the
residence time, catchment area, lake area and maximum depth were log-transformed. For
biological variables, classic monotonic transformations were used to meet the requirements of
the linear model (normality, linearity, homoscedasticity): count (abundance, richness) and
biomass metrics were log-transformed, proportion metrics were arcsine-square-root-
transformed, whereas diversity indices were kept untransformed. These metrics were then
scaled and the variation partitioning of each metric was performed using constrained
redundancy analysis (Legendre & Legendre 1998). Here, with a single response variable,
redundancy analysis consists in a multiple linear regression on the principal components of
predictors with orthogonal residuals. The three sets of predictors involved are: the ten
environmental predictors, the agricultural pressure index (PA) and the urban pressure index
(PU). A permutation test based on residuals after partial RDA was used to assess the
significance of the parts of variance explained solely by PA and solely by PU.
Each metric significantly explained by at least one of the two pressure indices was then
regressed using a stepwise “both sides” selection of predictors based on Bayesian information
criterion (BIC) from the complete model:
Metric ~ (AL + ADB + Dmax + SLDF + Vd + SumT + DiffT + RT + GI + WL) + PA + PU (1)
The pressure effect of a given pressure on a metric was considered only if: (i) the adjusted
R-squared of the resulting model was higher than 0.2; (ii) the coefficient of that pressure in
the resulting model was significantly different from zero; and (iii) the pressure was also
included in the best model selected on the basis of the root mean square error (RMSE) using a
leave-one-out cross-validation procedure (LOO).
All statistical analyses were performed using R statistical software (R.2.9.1) as well as
‘vegan’, ‘pls’, and ‘caret’ R packages (Ihaka & Gentleman 1996, R Development Core Team
2008).
4.4. Results
Study sites and environmental parameters. – In the present study, natural lakes are located
either in mountain areas (e.g., the Alps, the Massif Central) or next to the coast (Figure 4. 2).
On the contrary, reservoirs are roughly randomly spread throughout France. The mean values
of most environmental parameters are quite similar between natural lakes and reservoirs,
although natural lakes have larger residence time (692 days vs. 137 days) (Student t-test with
Welch correction, t = 2.16, P < 0.05) (Tableau 4. 1), lower SLDF (with Welch correction, t = -
6.83, P < 0.001), smaller catchment areas (t = -3.29, P < 0.001) and lower water level
fluctuations (t = -7.53, P < 0.001) than reservoirs.
Differences between lake types are clearer with respect to anthropogenic pressures
(Tableau 4. 2). Apart from network densities and urbanised areas, all anthropogenic pressures
displayed higher mean values for artificial lakes than for natural ones. For natural lakes, the
negative nutrient balances indicate a deficit of nutrients in the soil surface of the catchment,
whereas their highly positive values for reservoirs show a nutrient accumulation. In addition,
all agricultural variables reached higher maximum values for reservoirs than for natural lakes
– more than twice as much. For example, reservoirs have higher maximum values of
agriculture land cover areas (8,783.4 km2 vs. 790.6 km2) and nutrient balances (877.1 t(N) vs.
432.3 t(N), and 2,523.1 t(P) vs. 849.2 t(P)) than natural lakes. In summary, reservoirs undergo
much more agricultural pressure than natural lakes do.
Fish sampling and candidate metrics. – Lakes and reservoirs are very similar in terms of
common and rare species (Tableau 4. 3). Indeed, 68% of the fish species occur in both types
of ecosystems. In both natural lakes and reservoirs, Perca fluviatilis (Linnaeus, 1758),
Rutilus, rutilus (L., 1758) and Scardinius erythrophthalmus (L., 1758) were widespread (more
than two-thirds of the sampling sites) and had a high mean relative abundance (>5% in at least
one of two types). In addition, fish species with low occupancy (<10% of the sampling sites)
and poor average relative abundance (<1% in at least one of the system types) were nearly the
same as well (Tableau 4. 3). On the contrary we observed clear differences (≥ 25% in
frequency) between lake types concerning Abramis brama (L., 1758), Alburnus alburnus (L.,
1758), Blicca bjoerkna (L., 1758), Gymnocephalus cernuus (L., 1758), Sander lucioperca (L.,
1758) and Coregonus lavaretus (L., 1758) (Tableau 4. 3). All of them, except the last one
were more frequent in reservoirs than in natural lakes.
Most of the functional fish-based metrics also indicated similarities between lake types
(Student t-test, with Welch correction if needed, see Tableau 4. 5).
- 78 -
Overall, 11 out of 73 candidate fish metrics displayed a significant response to
anthropogenic pressures (Tableau 4. 6). None of the metrics based on the lithophilic, strict
lithophilic, intolerance, and invertivorous traits displayed a significant response. Moreover,
anthropogenic impacts do not appear significant on species richness or on diversity indices. In
addition, none of the candidate metrics indicated a significant anthropogenic impact on both
natural lakes and reservoirs.
- 79 -
Tableau 4. 6: Regression of fish metrics versus the environmental and anthropogenic variables.
Table entries are the regression coefficient model, R square adjusted, coefficient significance level, and variance partitioning; var env
is the percentage of the total variation attributable to pure environmental effects, var PA to pure agricultural pressure effects, var PU
to pure urban pressure effects, and unexpl variation unexplained by the model. For abbreviations see section “Candidate metrics”.
2
PA PU R -adjusted var env var PA var PU unexpl
Reservoirs BPUE 0.09* - 0.56 0.37 0.03* <0.00 0.40
n = 59 Sp_OTol 0.20** - 0.39 0.18 0.07* 0.00 0.59
B_OTol 0.20*** - 0.68 0.38 0.06** 0.01 0.33
Sp_Omni 0.20*** - 0.25 0.01 0.10* 0.02 0.79
Sp_Plankt 0.20*** - 0.34 0.10 0.10* 0.01 0.70
%Sp_Plankt 0.06*** - 0.21 <0.00 0.12** <0.00 0.85
B_Herb 0.20*** - 0.33 0.14 0.13** 0.02 0.71
%Sp_Pisc -0.08*** - 0.28 0.06 0.17** <0.00 0.73
Natural lakes %B_OTol - 0.20*** 0.43 0.27 <0.00 0.30** 0.51
n = 30 Nb_StPhyto 0.03*** - 0.66 0.23 0.20** 0.01 0.39
%B_Plankt 0.20* - 0.42 0.28 0.17* <0.00 0.65
*significant at 0.05 level; **significant at 0.01 level; ***significant at 0.001 level
Three metrics displayed a significant response to anthropogenic pressures in natural lakes
(Tableau 4. 6). %B_OTol increases with urbanisation stress (30% of the variance explained).
Nb_StPhyto and %B_Plankt seem to be good indicators of agricultural impacts (Tableau 4. 6)
(20% and 17% of variance, respectively, explained exclusively by agricultural pressure).
For the reservoirs, eight metrics were selected, including BPUE (Tableau 4. 6). Five of
them relate to assemblage trophic structure and two to tolerance guild metrics (Sp_OTol and
B_OTol). The only type of pressure significantly contributing to the model was PA (Tableau
4. 6). Apart from the percentage of piscivorous species, the other seven metrics displayed a
positive response to the agricultural pressure when the environment was controlled for. The
metric %Sp_Pisc showed the strongest response with 17% of its variance attributed to PA.
B_Herb and %Sp_Plankt came next with 13% and 12%, respectively.
All linear models included at least one significant coefficient for environmental variables
(unpublished results). This highlights the importance of accounting for the environmental
patterns of variability when studying the response of fish-based metrics to anthropogenic
pressures at broad spatial scales. For the natural lakes, this is particularly true for the
%B_Plankt and %B_OTol (Tableau 4. 6), with 28%, and 27%, respectively, of their variance
attributed to environmental variables. For reservoirs, this mostly concerns B_OTol and
BPUE, with 38%, and 37%, respectively, of their variance explained by environmental
variables.
4.5. Discussion
The data set processed here can be considered representative of the diversity of lakes at
the national scale (excluding lakes above 1500 m). Although natural lakes and reservoirs both
meet the generic definition of lakes (Politou et al. 1993), they clearly differ in terms of
morphological characteristics, functioning (e.g., water level fluctuation (Tableau 4. 1)) and
anthropogenic pressures (Cooke et al. 1993, Thornton 1990). Reservoirs undergo much more
agricultural pressures than natural lakes do certainly because the storage of water in artificial
lakes favors the development of irrigation.
Assessing lake conditions through fish assemblages

The present study provides evidence that fish assemblages are good potential indicators of
urban and agricultural impacts on lakes. Eleven metrics (among the initial set of 73)
responded to the lake catchment pressures when the environment was controlled for.
Most of the selected metrics responded to agricultural pressures. This prevailing effect of
agriculture versus urban land use can be explained by the differences in intensity of these two
pressures. The mean percentage of agricultural land cover in catchments exceeds by one order
of magnitude that of urban land use (Tableau 4. 2). The buiding of treatment plants for
domestic and industrial sewage certainly contributed to mitigate the impact of urban land use
on freshwater ecosystems. Conversely, agriculture is still a major user of fertilisers and
pesticides, and it contributes to soil erosion. In this context the prominent effect of agriculture
on freshwater assemblages is not really surprising, even though the processes involved remain
unclear. Direct impacts (e.g., by toxicity) are certainly less important than those explained by
habitat alteration or eutrophication. Finally, there is also more variability in agricultural than
in urban land cover (synthesized by higher SD, see Tableau 4. 2), which may explain a clearer
response of fish-based metrics. Moreover, in the present study, the urbanisation of catchments
was very low compared to that in other studies (Drake & Pereira 2002, Whittier 1999). In
addition, the lack of significance might also be due to the fact that these relationships between
metrics and pressures are actually non-linear, which was not considered here. Metrics might
indeed be, for example, sensitive to a given pressure above a certain threshold.
Regarding natural lakes only three metrics displayed a significant response to
anthropogenic pressures (Tableau 4. 6). This is less than for reservoirs, which may be
explained by the above-mentioned differences in the agricultural pressure between natural
lakes and reservoirs. Furthermore, we have to acknowledge that the number of natural lakes
included in the present study and the variability between these lakes (i.e. coastal to mountain
lakes) may limit the possibility of identifying certain relevant metrics. Nevertheless,
considering the low number (less than 70) of French natural lakes over 50 ha, this exercise
will remain difficult at a national scale, whatever the sampling method used. At the European
level, similar tests would be more powerful.
Finally, it is obvious that fish management practices can influence fish species
composition and abundance (Argillier et al. 2002a). However, in this study, fish species were
considered self-sustaining. Indeed, all species are not stocked and when they are, according to
our knowledge of local management practices, this is infrequent. Furthermore, even if none of
the lakes studied was concerned with commercial fishing, we cannot exclude bias in our
metric calculations (i.e., metrics related to the abundance of piscivores and salmonids) due to
intensive recreational angling.
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Fish-based metric responses on both lentic ecosystems
For planktivorous species, for example, agriculture has a significant effect on %B_Plankt
for natural lakes and on %Sp_Plankt for reservoirs (Tableau 4. 6), but no metric revealed a
common response to a pressure on both natural lakes and reservoirs.
The selected fish-based metrics

Only fish-related metrics based on tolerance, reproduction, and trophic guilds were
retained in this study. The tolerance-related metrics seemed to display clear responses to
anthropogenic pressures both on reservoirs and natural lakes. Most IBIs developed on rivers
(Oberdorff et al. 2002, Pont et al. 2006), reservoirs (Karr & Dionne 1991, McDonough &
Hickman 1999) or natural lakes (Belpaire et al. 2000; Drake & Pereira 2002) use some
measure of the relative dominance of tolerant species to assess moderate to high levels of
degradation (Simon & Lyons 1995). Nevertheless, as discussed previously, the metric units of
the selected tolerance-related metrics were different between natural lakes and reservoirs.
Moreover, it is apparent that 5 of 11 intolerant species are salmonids (see Appendix 3) and
they likely occupy mainly the coldwater lakes located above 1500 m that were excluded from
our analyses. In spite of the absence of response of intolerant species-based metrics, most of
our results confirm the value of tolerant-related metrics.
Apart from the proportion of piscivorous species, all other trophic-based metrics displayed
significantly positive relationships with agricultural pressure, among both types of lakes. All
six trophic-based metrics selected in this study responded similarly to those found in other IBI
adaptation studies (Appelberg et al. 2000; Drake & Pereira 2002; Jennings et al. 1995; Karr &
Dionne 1991). In contrast to temperate stream ecosystems, it is acknowledged that lentic
ecosystems are considered more stable from an environmental point of view, thereby
promoting intra-assemblage interactions (e.g., competition, predation) and giving rise to
trophically structured assemblages (Hugueny & Paugy 1995, Ibañez et al. 2009, Oberdorff et
al. 1998). Hence, studying the response of the trophic composition metrics to anthropogenic
pressures in lentic ecosystems was useful, while nutrient loadings induced by agricultural land
use could probably modify the equilibrium between the different trophic levels (Carpenter et
al. 1985, Vitousek et al. 1997).
Among the metric based on reproductive guilds, Nb_StPhyto was the only one that
showed a significant response to agricultural pressure; this was observed only for natural
lakes (Tableau 4. 6).
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Candidate metrics displaying no response
Species richness was not correlated to catchment-scale anthropogenic pressures. This is
congruent with previous results (Irz et al. 2008). Various responses were reported for this
metric. It can increase due to eutrophication (Dodson et al. 2000, Mittelbach et al. 2001) or
species introduction (Irz et al. 2004a), but decrease due to the extirpation of habitat-sensitive
taxa (Corbacho & Sanchez 2001).
The three diversity indices were not either correlated to anthropogenic factors. These
indices have rarely been used in fish index development, probably because they are highly
correlated with each other and because the information they provide is difficult to interpret
(Holmgren et al. 2007). In addition, zoogeographic factors (e.g., post-glacial dispersal,
hydrographic regions, and pH) relevant for explaining species richness and diversity indices
have not been considered in this analysis (Griffiths 1997; Irz et al. 2004a).
Despite a strong negative relationship between the native related metrics and
anthropogenic pressures observed in other studies (e.g., Jennings et al. 1999), this association
was not detected. However, the species driving this relation in the French data set were in the
majority Rutilus rutilus (L., 1758), Perca fluviatilis (L., 1758), Blicca bjoerkna (L., 1758) and
Abramis brama (L., 1758). These species are overall tolerant as regards degraded lake
conditions. Hence, the selection of a fish species able to respond, depending on the biological
trait that is studied, in both directions to degraded lake conditions explains why the response
of those candidate metrics could be concealed in our study. The same explanation can be
advanced to account for the absence of relationships between agricultural or urban pressure
and fish-based metrics related to lithophilic, strict lithophilic, and invertivorous species.
In addition, the near absence of reproduction-related metric responses highlighted by this
study may be the result of the poor relevance or the deficiency of the lake descriptors
considered. It is commonly accepted that both local and large-scale processes influence local
fish assemblages in lakes (Irz et al. 2008, Irz et al. 2007, Petesse et al. 2007a). In this study,
only large-scale pressures were considered and more relevant descriptors of habitat and
hydromorphological alteration have to be collected; further studies will then be required to
better understand the influence of regional versus local anthropogenic pressures on fish
assemblages (Irz et al. 2008).
Finally, the relationship between intolerant species-related metrics and anthropogenic
pressures can be concealed, as reported by Irz et al. (2008), because “a species could be
considered as intolerant in some regions where it is restricted to some particular type of
environment, and tolerant in another region where it is widespread.” However, in this study
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we can note that, apart from Gobio gobio (L., 1758), all other fish species driving that relation
had either low occupancy (<3% of the sampling sites) or poor average relative abundance
(<1% in at least one of both types). Consequently, intolerant species-related metrics displayed
a very narrow range of variance, thus explaining their lack of response to anthropogenic
pressures.
Additional work is needed, and as suggested by Berrebi-dit-Thomas et al. (1998) and
Whittier et al. (2007), metrics based on biological traits that are the expression of specialized
attributes (e.g., invertivores, lithophilic) should be calculated without considering overall
tolerant fish species. Indeed, this modification may prevent the sensitivity of a specific
biological parameter from being concealed due to the heterogeneity of the fish species
contributing to the calculation of the fish-based metric.
Multiple testing
No corrections were made for multiple testing. We are aware of the increasing risk of
false positive. However, there is much true dependence between the metrics tested since they
are based on guild composed by partly the same species and for a given guild six metrics are
based on the same species but measured differently. In short, because of this dependence, the
true number of effective testing must be largely fewer than the number of metrics considered
(Cheverud 2001). Nevertheless, much attention must be paid to the levels of significance.
4.6. Conclusion
The present study indicates that fish assemblages are appropriate ecological indicators for
both natural and artificial lakes. Our results underline the necessity to control for the
environment when developing FBIs at broad geographical scales. They also prove that the
relevant metrics differ between natural lakes and for reservoirs. This challenges the attempts
to use natural systems as references to assess the ecological potential of artificial ones.
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- 86 -
CHAPITRE 5
Développement d’un indice poisson pour évaluer l’état écologique des plans d’eau en
France : une approche par hindcasting
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L’une des difficultés majeures dans le développement des indices biologiques réside dans
la définition des conditions de référence à utiliser pour étalonner le système de classification
(Stoddard et al. 2006). Pour les milieux aquatiques, la notion de conditions de référence
correspond à la meilleure situation observée dans le groupe de masses d’eau étudié (Davies &
Jackson 2006, Holmgren et al. 2007, Hughes 1995, Jackson & Davis 1994, Steedman 1994,
Stoddard et al. 2006). Ainsi, en pratique, les sites dits « de référence » sont très souvent les
sites qui présentent un degré minimal d’altération (Pont et al. 2006).
L’approche dite « des conditions de référence » (Bailey et al. 1998), bien que très utilisée,
présente des imprécisions d’un point de vue méthodologique (Bowman & Somers 2005,
Stoddard et al. 2006). En effet, les niveaux de perturbation tolérés pour chaque classe de
référence peuvent grandement varier selon que l’on considère des sites peu ou pas impactés
comme pouvant être définis en tant que sites de référence (Stoddard et al. 2006). Selon les
niveaux choisis, les interprétations biologiques qui en découlent peuvent alors être également
très différentes. De plus, cette approche introduit une part d’incertitude de par l’utilisation de
termes et/ou d’expressions polysémantiques pour évaluer les impacts anthropiques. Par
exemple, Stoddard et al. (2006) indiquent que l’expression « conditions de référence » est
utilisée de manière impropre pour qualifier de nombreux états biologiques différents, incluant
à la fois l’état d’un écosystème à un moment déterminé du passé, les meilleurs situations
observées actuellement, l’état d’un écosystème en l’absence de pressions anthropiques
significatives, ou bien encore l’état que devrait avoir les sites s’ils étaient mieux gérés. Cette
approche présente cependant aussi des avantages et implique, notamment, de définir les
caractéristiques attendues des communautés biologiques dans un site donné en l’absence
d’impact d’origine anthropique significatif (Bowman & Somers 2005, Hawkins et al. 2010,
Stoddard et al. 2006, USEPA 1998). Comme pour les autres méthodes de bioindication, la
question de l’échantillonnage est centrale et conditionne en grande partie l’efficacité des
indices (Stoddard et al. 2006). Le prélèvement doit ainsi être suffisamment important pour
permettre une bonne évaluation des occurrences et abondances ainsi que de la richesse
(Angermeier et al. 2000).
La DCE définit les conditions de référence, ce qui a pour but d’harmoniser les diverses
approches en leur fixant un cadre (European Community 2000). En effet, la guidance
REFCOND (Anonymous 2003b) précise que les conditions de référence sont définies à partir
de sites pas ou très légèrement perturbés. Il est aussi fait référence à un recours potentiel à des
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données historiques. Cette dernière option a été relativement peu utilisée en Europe par les
Etats membres1 afin de définir des sites de référence (Gassner et al. 2005).
Pour les retenues, qui sont classées comme masses d’eau fortement modifiées, la DCE
stipule que la référence est à définir sur le type de milieu naturel le plus proche. Si cette
notion peut avoir un sens pour certains systèmes naturels modifiés, elle est peut-être plus
critiquable pour d’autres. Nous avons vu en effet que les réservoirs sont souvent considérés
comme des systèmes intermédiaires entre les milieux lotiques et les lacs naturels (Gelwick &
Matthews 1990, Irz et al. 2006, Wetzel 1990).
En France, quel que soit le type de milieu, les sites de références ont été définis par
jugement d’expert par les Agences de l’eau. Les critères de pressions à considérer sont fixés
par une circulaire ministérielle (Ministère de l'Ecologie et du Développement Durable 2004),
mais leur description laisse une large part d’interprétation possible, qui est probablement
source d’hétérogénéité. Suivant cette circulaire, vingt-trois lacs naturels ont été considérés
comme référence, mais la plupart d’entre eux sont petits, situés en tête de bassin, à une
altitude supérieure à 1500 m, c’est-à-dire dans des conditions environnementales où
l’ichtyofaune n’est pas considérée pertinente pour établir des diagnostics d’états. Pour les
retenues, utilisant les mêmes critères, quelques sites ont été considérés comme référence sur le
bassin Rhône Méditerranée et Corse, mais il s’agit d’une démarche isolée et qui n’a pas été
suivie à l’échelle nationale. De plus, aucun consensus sur les critères de classification des
retenues en référence n’émerge et nous avons montré la non pertinence de considérer un lac
naturel comme une référence pour une retenue.
Parmi les alternatives possibles pour le développement d’un indice, et compatibles aux
exigences de la DCE [Annex II, 1.3 (i-vi)] (Anonymous 2003a), l’approche basée sur une
approche par hindcasting a été explorée ici (Baker et al. 2005, Kilgour & Stanfield 2006).
Cette méthode, basée sur des modèles statistiques incluant comme prédicteurs à la fois des
des variables environnementales et des facteurs anthropiques connus pour influencer les
communautés piscicoles, a été initialement développée au Canada par Reynoldson and Day
(1998) pour étudier les facteurs structurant les peuplements benthiques. Cette méthode,
comporte de nombreux avantages, notamment celui de nécessiter juste un nombre restreint de
sites de références ou peu impactés. Elle permet en premier lieu, de contourner le problème de
1
Cependant l’Autriche et les régions frontalières (nord de l’Italie, Bavière et Slovénie) utilisent cette notion sur
les lacs alpins, voir Gassner et al. (2005).
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la sélection de sites de références nécessaires à la modélisation des conditions biologiques
attendues en l’absence de pressions (Pont et al. 2006, Bailey 1998). En second lieu, elle
permet de conserver les avantages d’une approche par modélisation statistique de la réponse
des métriques à l’environnement naturel, sans recours à la classification des plans d’eau.
L’article suivant décrit l’application de l’approche hindcasting pour le développement
d’un indicateur poisson pour les lacs naturels français, et d’un indicateur pour les retenues
françaises.
Dans cette étude, nous avons considéré, en plus des pressions à large échelle utilisées dans
les analyses précédentes, celles s’exerçant localement. En effet, ces deux types de pressions
n’ont a priori pas les mêmes effets sur les communautés piscicoles lacustres. Contrairement
aux pressions qui s’exercent à l’échelle régionale et dont les effets sont énumérés en Annexe
1, les pressions qui s’exercent à l’échelle locale se manifestent, en particulier, par la nature
des aménagements rivulaires des plans d’eau et par les activités humaines qui s’y déroulent
(Rowan et al. 2006). Ainsi, les pressions locales semblent plutôt agir sur la quantité et la
qualité des habitats disponibles pour les organismes aquatiques (McGoff & Irvine 2009,
Moore 1980, Ostendorp et al. 2004a, Schmieder 2004).
Nous avons montré que l’utilisation de cette approche statistique permettait de mettre en
évidence de fortes corrélations entre certaines caractéristiques des peuplements de poissons et
les perturbations subies par les retenues et les lacs naturels français. Toutefois, des limites
sont à formuler en ce qui concerne les lacs naturels. En effet, pour ces derniers, aucune
relation entre les métriques piscicoles et les pressions agricoles s’exerçant à l’échelle du
bassin versant n’a pu être établie. De plus, contrairement aux réservoirs, aucune métrique
piscicole associée aux guildes de trophie et de reproduction n’a été incluse dans l’indice
poisson final pour les lacs naturels.
Par ailleurs, il apparaît clairement que certaines métriques répondent exclusivement à des
pressions anthropiques locales. Ces résultats confirment ceux obtenus dans plusieurs études
américaines (Jennings et al. 1999a, Jennings et al. 2003, Minns et al. 1994, Thoma 1999),
mettant en évidence des relations entre l’altération hydromorphologique et l’ichtyofaune des
plans d’eau, à large échelle spatiale. Ils sont particulièrement intéressants dans le contexte de
mise en œuvre de la DCE. Ils soulignent la pertinence de l’utilisation de l’ichtyofaune pour
l’évaluation de l’état écologique des plans d’eau, du fait de sa complémentarité avec d’autres
indicateurs, tels que ceux utilisant le phytoplancton.
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Development of a fish-based index (FBI) of biotic integrity for French lakes using the
hindcasting approach (sous presse dans Ecological Indicators – DOI:
10.1016/j.ecolind.2011.03.028)
Lionel Launois, Jacques Veslot, Pascal Irz, and Christine Argillier
5.1. Introduction
The European Water Framework Directive (WFD, 2000/60/EC) (European Community

2000) aims at “restoring and maintaining the chemical, physical and biological integrity of the
Nation’s waters.” To achieve these objectives, it promotes basin-scale planning of
management actions. Hence, there is a need to develop tools for the assessment of the
integrity of water bodies. The directive indicates that fish is one of the “quality elements” to
be considered for the ecological evaluation of surface waters. Pioneering work on the
midwestern streams of the US (Karr 1981, Karr & Dudley 1981) established the basis for the
successful development of fish-based indices of biotic integrity (IBIs) now used for European
lotic systems (Oberdorff et al. 2002, Pont et al. 2006). On streams and rivers, the early
implementation of monitoring networks allowed the collection of large and accurate data sets.
Scientists could therefore (i) assess reference conditions by modeling fish community
attributes (metrics) vs. environmental variables using a set of undisturbed sites, (ii) select
metrics displaying response to anthropogenic pressures, and (iii) aggregate individual metrics
into a synthetic index.
In contrast, with the exceptions of the Scandinavian programs related to acidification or of
EU-funded research [see Moss et al. (2003)], the monitoring of lakes in Europe has long been
considered on a site-by-site basis. Focusing on the environmental differences between these
well-defined, apparently independent entities, any unifying framework appeared doomed to
failure, overwhelmed by idiosyncrasies. However, progress in macroecological and
biogeographical research, along with the political will and its legislative translation, have
recently made lake monitoring feasible and solvent; consequently survey data are now
available.
However, problems remain to be solved when following a reference condition approach
(RCA) if there are too few lakes of any type to construct predictive models (Baker et al. 2005,
Kilgour & Barton 1999) or if too few lentic systems could clearly be classified as unimpacted
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or at least minimally impacted (Herlihy et al. 2008, Stoddard et al. 2006). In this instance,
according to the WFD (Annex II, 1.3 (i-vi)) (Anonymous 2003a), reference conditions can be
defined using other approaches, such as statistical hindcasting. This alternative approach to
RCAs, based on models including as predictors both anthropogenic factors and environmental
variables known to influence natural ecological patterns, has been developed in Canada by
Reynoldson and Day (1998) to assess benthos assemblages. Baker et al. (2005) and Kilgour
and Stanfield (2006) added that these alternative approaches to RCA models using landscape
context models could also be adapted to compute reference fish assemblages in freshwater
systems by setting anthropogenic pressure variables to zero, “so long as the variables used to
predict the expected condition do not vary with anthropogenic disturbance.” Hence, the
relationship between a metric and anthropogenic pressure variables is used to hindcast the
value of the metric in the absence of such pressures. In this approach each fish-based metric is
re-expressed as a deviation from its expected score based on explicitly modeled reference
conditions at each sampled lake. Then, after having scaled this deviation, a normalized score
is associated with each metric (Baker et al. 2005, Kilgour & Stanfield 2006).
The aim of this study was to develop a fish-based index (FBI) of biotic integrity for both
natural and artificial French lakes. The first step was to select a set of functional metrics
responding to local and catchment-scale anthropogenic pressures when environmental
variability is controlled for. Second, a subset, reflecting complementary aspects of community
functioning and chosen to avoid redundancy, was aggregated into a synthetic index.
5.2. Methods
5.2.1. Study sites

Throughout this article, we refer to lakes of natural origin as “natural lakes” and to lentic
systems created artificially by damming as “reservoirs”; “lakes” is the generic term used for
both types of lentic ecosystems.
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Figure 5. 1: Location of the 91 study sites in France.
Grey lines represent ecoregion limits. ○Natural lakes ●Reservoirs
Anthropogenic catchment-scale pressure indices were computed for 40 natural and 106
artificial lakes, while local pressure data, requiring a field survey, were collected for a subset
encompassing 24 natural and 67 artificial lakes. Fish data and environmental features were
also collected from these 91 sites. The study lakes were selected because they belonged to the
French WFD monitoring network. This network encompasses all lakes larger than 50 ha in
surface area or included as reference sites. At the national scale, the data set processed here
accounts for one-third of such natural lakes and one-fifth of such reservoirs. They were
located in 14 out of 22 French ecoregions (Wasson et al. 2001) (Figure 5. 1). We excluded
brackish littoral lakes as well as mountain lakes (over 1500 m above sea level). The latter
shelter exclusively introduced fish species (Argillier et al. 2002b). The range of anthropogenic
pressures covered by natural lakes is much narrower than that covered by artificial lakes
(Tableau 5. 1).
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Tableau 5. 1: Left: Principal component analyses (PCAs) carried out separately on the
watershed agricultural and urban pressure variables (AP and UP respectively).
Table entries are the percentage of variance (inertia) explained by each first
principal component (PC) and the correlations between the variables and the first
PCs. Those loading most heavily are in bold. Right: descriptive statistics of the
anthropogenic variables included in the analysis.
PCA Descriptive statistics
All lakes (N=146) Reservoirs (N=67) Naturals (N=24)
AP UP Mean SD Range Mean SD Range
Variable Inertia 63% 75%
% catchment agricultural 0.84 - 57.8 30.1 0.0 - 94.4 22.9 21.3 0.0 - 72.0
Agriculture land cover area
2 0.68 - 413 1235 0 - 8783 34 93 0 - 465
(km )
-598.2 to -346.8 to
Nitrogen balance (ton) 0.77 - 159.2 331.4 -72.8 97.8
1101.3 78.3
Phosphorus balance (ton) -152.0 to -389.1 to
0.86 - 507.9 651.4 -39.3 153.1
* erodability coefficient 2523.1 314.9
% catchment urban - 0.90 3.7 8.0 0.0 - 55.0 3.6 4.1 0.0 - 15.3
Population density in
2 - 0.89 64.2 130.6 5.6- 1006.5 41.0 53.8 3.7 - 260.5
catchment (pers./km )
Transportation network
- 0.79 2.1 1.7 0.0 - 10.3 2.2 2.6 0.0 - 11.3
density
Lake Habitat Modification
- - 18.9 6.8 6.0 - 36.0 17.2 7.1 6.0 - 32.0
Score (LHMS)
5.2.2. Abiotic environmental characteristics of lakes

Efficient impact assessment at broad spatial scales requires the consideration of the
environmental variables that are known to determine natural ecological patterns and that are
not modified by human activities. Hence, the suite of variables given in Tableau 5. 2 was
included in the models. Maximum depth (Dmax), shoreline development factor (SLDF) (Eadie
& Keast 1984), and volume development (Vd) (Håkanson 1981) determine the abundance of a
lake’s littoral habitat (Argillier et al. 2002c) and influence the water’s dissolved oxygen level
(Hondzo & Stefan 1996). Lake area (AL) is a strong driver of fish species richness (Barbour &
Brown 1974, Eadie et al. 1986). Drainage basin area (ADB) can be considered a surrogate for
habitat diversity upstream from the lake (Irz et al. 2004b). January and July mean monthly
maximum air temperatures (TJanuary & TJuly) were computed using the AURELHY climate
model (Benichou & Le Breton 1987) which allowed us to derive the following two
independent variables related to the temperature requirements of living organisms (Daufresne
& Boet 2007, Irz et al. 2007, Mason et al. 2008, Rathert et al. 1999):
(i) SumT = TJanuary + TJuly
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(ii) DiffT = TJuly - TJanuary
Residence time (RT) strongly influences within-lake physicochemical as well as biological
processes (Thornton 1990). It was computed based on a hydrological model of mean yearly
runoff (Sauquet 2006, Sauquet et al. 2000). The geological index (GI) represents the
percentage of lake catchment area as calcareous (derived from 1:50,000 geological maps –
BRGM – http://www.brgm.fr/cartegeol.jsp). Therefore, it relates to the water chemistry and
buffering capacity (Alpay et al. 2006, Brousseau et al. 1985). Water level maximal amplitude
(WL) was also included because of its detrimental effect on fish feeding conditions,
sheltering, and reproduction (Rowan et al. 2003, Winfield 2004).
Tableau 5. 2: Environmental parameters, mean and extreme values for the 91 study sites.
Reservoirs (N=67) Natural lakes (N=24)
Parameter Mean Range Mean Range
Geologic index (GI) (% of the drainage basin as calcareous) 0.24 0.00-1.00 0.38 0.00-1.00
Residence time (RT) (day) 128.82 0.11-922.80 771.40 5.00-5095.00
a
Water level maximum amplitude (WL) 1.58 0.00-2.00 0.42 0.00-2.00
2
Catchment area (ADB) (km ) 653 1-10533 134 1-1033
Lake area (AL) (ha) 216 38-2203 772 13-5768
Maximum depth (Dmax) (m) 23 2-100 29 2-110
b
Shoreline development factor (SLDF) 3.49 1.09-9.62 1.65 1.01-3.80
TJanuary + TJuly (SumT) 24.14 15.49-31.27 19.24 13.18-28.91
TJuly - TJanuary (DiffT) 15.81 10.43-19.42 17.46 13.63-19.91
c
Volume development (Vd) 1.18 0.37-2.96 1.39 0.30-2.66
a
3 classes: 0 (no variation), 1 (≤3 m), 2 (>3 m).
b
Calculated as: perimeter / (√4π×surface).
c
Calculated as: Lake volume / (Maximum depth × Lake area).
5.2.3. Fish sampling procedures

Fish data for 91 lakes were collected between 2005 and 2009 using a standardized method
(C.E.N. 2005b). A stratified random sampling scheme was implemented during summer (June
– October, surface water temperature over 15°C). The sampled lakes were divided into depth
strata and random samplings were performed within each depth stratum using both benthic
and pelagic multimesh gillnets. The benthic gillnets were 30 m long, 1.5 m high, and
composed of 12 different mesh sizes ranging from 5 mm to 55 mm following a geometric
series chosen to maximize the evenness of selectivity across fish sizes. Lakes deeper than 6 m
(N=50) were complementarily sampled with pelagic multimesh gillnets 27.5 m long and 6 m
high. The number of nets increased with the lakes’ depth and area. Gillnets were set around 7
p.m. for approximately 12 h. Fish individuals were identified as to species, weighed, and
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measured. A whole-lake estimate was then calculated for species relative abundance and
biomass, as well as the means of catch per unit effort (CPUE) and biomass per unit effort
(BPUE) (including both the benthic and the pelagic nets), respectively expressed as the
overall number of individuals or biomass captured divided by the unit-of-effort (i.e., the nets’
surfaces multiplied by the sampling duration). Fish data collected using this method are
comparable over time within a lake and between lakes (Appelberg 2000, Appelberg et al.
1995).
5.2.4. Candidate metrics

Candidate metrics were selected for their inclusion in other lake and stream IBIs (Drake &
Pereira 2002, Holmgren et al. 2007, Jennings et al. 1999b, Karr et al. 1986, Minns et al. 1994,
Schulz et al. 1999, Simon & Lyons 1995, Whittier 1999). Each of the 41 fish species collected
in this study (Appendix 4) was classified as to tolerance (overall tolerant, OTol and overall
intolerant, OIntol), reproduction (lithophilic, Lith; strict lithophilic, StLith; and strict
phytophilic, StPhyto), trophic group (omnivorous, Omni; invertivorous, Invert; planktivorous,
Plankt; herbivorous, Herb; and piscivorous, Pisc), and origin (native, Nat), using information
from previous grey or published literature (Keith & Allardi 2001, Kottelat & Freyhof 2007,
Pont et al. 2006), online resources (Froese & Pauly 2009 – http://www.fishbase.org/), or
experts’ estimations (H. Persat, O. Schlumberger & N. Poulet, personal communications)
(Tableau 5. 3). For each of these functional guilds, six metrics were computed: (i) number of
species sharing a trait (Sp_trait), (ii) number of species sharing a trait divided by the total
number of species (%Sp_trait), (iii) number of individuals sharing a trait caught by unit effort
(Nb_trait), (iv) number of individuals sharing a trait divided by the total number of
individuals (%Nb_trait), (v) biomass of individuals sharing a trait caught by unit effort
(B_trait), and (vi) biomass of individuals sharing a trait divided by the total biomass of
individuals (%B_trait). Community-level metrics were also considered: diversity indices
based on fish abundances [the Shannon-Wiener index (Shannon & Weaver 1949), SWI; the
Simpson index (Simpson 1949), SI; and the evenness index (Hill 1973), EI], total species
richness (RS), the BPUE, the CPUE, and the average biomass (AB) calculated for each
sampled site as the ratio of BPUE to CPUE. Altogether that comes to 73 candidate metrics
expected to adequately reflect community structure and functioning.
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Tableau 5. 3: Guild classification for French lake fishes.
Guilds Classes Definitions
Tolerance guild Overall tolerants Species insensitive to any common impact related to nutrient
(Karr et al. 1986) regime, habitat structure and water chemistry.
Overall intolerants Loss of intolerant species is a response to degradation,
whereas the number of tolerant species will tend to increase
with disturbance.
Reproduction guild Lithophilic Can reproduce on unsilted mineral substrates whose larvae
(Balon 1975, Dycus & are photophobic.
Meinert 1994) Strict lithophilic Need mineral substrates to reproduce.
Strict phytophilic Need plant substrates to reproduce and whose larvae are not
photophobic.
Origin guild (Keith & Allardi Natives Species considered as native to the drainage basin
2001) considered.
Trophic guild Omnivores Feed on both animal and vegetal items (more than 25% plant
(Bruslé & Quignard 2001; material and more than 25% animal material).
Dycus & Meinert 1994, Invertivores Feed at least partially on invertebrates/insects (more than
Goldstein & Simon 1999; 75% macro-invertebrates in the diet).
Pont et al. 2006) Planktivores Feed mainly on zooplankton and/or phytoplankton (more
than 75% of plankton in the diet).
Herbivores Feed on plant items (more than 75% of plant material in the
diet).
Piscivores Feed on fish at least partially when adults (more than 75%
fish in the diet).
We examined the distribution and species composition of each metric. The metrics based
on functional traits shared by less than three species were omitted. Non-native species were
not excluded because they also have potential indicator properties (Halliwell et al. 1999, Irz et
al. 2008, Reyjol et al. 2007).
5.2.5. Catchment-scale anthropogenic pressures

Seven catchment-scale pressure variables were collected for the overall data set (n = 146)
(Tableau 5. 1). Within each lake catchment, we estimated (i) urban and agricultural land cover
percentages, (ii) agricultural land use area (km2), (iii) population density (people/km2), and
(iv) network density of roads and railways (km.km-2). These density variables relate to the
watershed dynamics, especially chemical discharges into lakes (see Pitt et al., 1995) and soil
fragmentation (Forman & Alexander 1998, Jones et al. 2000). In addition, the soil surface
nitrogen (N) and phosphorus (P) balances were computed using the NOPOLU System 2
model (Commercial name, for further details see BETURE CEREC / JAAKKO PÖYRY
INFRA). Nutrient balance represents the annual mean difference between the total quantity of
nutrient entering the soil (e.g., inorganic fertilisers, livestock manure, nitrogen fixation) and
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the quantity leaving it (e.g., harvested crop production, grass and fodder crop production). As,
the long term, there is no infinite accumulation, this difference gives an estimate of the
nutrient load in the runoff. Furthermore, as phosphorus is not a very mobile chemical element
(Carpenter et al. 1998b) and according to the recommendation of Coale (2000), we multiplied
the phosphorus balance by the mean value of the lake’s watershed soil erosion factor. This
factor was estimated with a model including soil slumping and erodibility, topography, land
cover, rainfall, and dominant cultivated land covers (Van der Knijff et al., 2000 -
http://eusoils.jrc.ec.europa.eu/ESDB_Archive/serae/grimm/erosion/inra/europe/analysis/maps
_and_listings/web_erosion/index.html).
We assumed that these seven variables derived from GIS data reflected the anthropogenic
pressures undergone by lakes at the catchment scale.
5.2.6. Local-scale anthropogenic pressures

Local pressure variables were collected between 2005 and 2009 using the Lake Habitat
Survey (LHS, see Rowan et al., 2006) standard procedure. This technique was designed in
Scotland to meet the hydromorphological and environmental impact assessment needs of the
WFD. It requires a detailed recording, of shoreline features at a number of plots,
complemented by a meso-scale survey of the entire lake, including shoreline characteristics
and pressures as well as modifications of the hydrological regime. For modeling purposes, a
Lake Habitat Modification Score (LHMS) was computed from the raw LHS data (Tableau 5.
1). According to Rowan et al. (2006), this score “synthesizes a wide array of anthropogenic
pressures” at the lake’s local scale. LHMS values ranged from 0 to 42, the latter value
corresponding to high levels of anthropogenic pressure
5.2.7. Statistical approach

Some of the variables were transformed in order to meet the requirements of the linear
model. The environmental variables displaying skewed distributions (residence time,
catchment area, lake area and maximum depth) were log-transformed, along with abundance,
richness and biomass metrics as well as agricultural land cover area, population density, and
nutrient balances. Proportion metrics and percentages of land use in catchments were arcsine-
square-root-transformed.
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Principal component analyses (PCAs) were performed on agricultural and urban variables
separately (Tableau 5. 1) to reduce dimensionality and eliminate colinearity. For each group
of pressures, the first principal component (AP & UP) was retained as a synthetic pressure
variable.
A comparative analysis of the response of fish-based metrics to anthropogenic stressors
for natural and artificial lakes was carried out in a previous study (see Launois et al., 2011).
Consequently, the two types of lentic ecosystems were analyzed separately.
5.2.7.1.Metrics selection
The metrics were selected using an elimination process. They were scaled and submitted
to variation partitioning using constrained redundancy analysis (Legendre & Legendre 1998).
With a single response variable, redundancy analysis consisted in a multiple linear regression
(MLR) on the principal components of predictors with orthogonal residuals. The three sets of
predictors involved were: the ten environmental predictors, the regional pressure indices (AP
and UP) and the Lake Habitat Modification Score (LHMS). A permutation test based on
residuals after partial RDA was used to assess the significance of the part of variance
explained solely by AP, solely by UP, and solely by LHMS.
For each type of lake, each metric significantly explained by at least one of the three
pressure indices was then regressed using a stepwise selection of predictors based on the
Bayesian information criterion (BIC) from the complete model:
(1) Observed metric ~ (AL + ADB + Dmax + SLDF + Vd + SumT + DiffT + RT + GI + WL)
+ AP + UP + LHMS
The effect of a given pressure on a metric was considered only if: (i) the adjusted R-
squared of the resulting model was higher than 0.2, (ii) the coefficient of that pressure in the
resulting model was significantly different from zero, and (iii) the pressure was also included
in the best model selected on the basis of the root mean square error (RMSE) using a leave-
one-out cross-validation procedure (LOO). The other metrics were excluded.
5.2.7.2.Hindcasting approach
The hindcasting approach was restricted to the selected metrics. Their expected values in
the absence of anthropogenic pressures were predicted by setting the anthropogenic pressure
variables (i.e., AP, UP, and LHMS) to zero in the models obtained above. A normalized
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deviation (Baker et al. 2005, Kilgour & Stanfield 2006) was then obtained for each metric at
each site by dividing the difference between the observed (i.e., 1.67 in Tableau 5. 4) and the
expected (i.e., 1.09 in Tableau 5. 4) values of the metric by the square-root of the model’s
residual variance (i.e., 1.81 in Tableau 5. 4).
Tableau 5. 4: Example calculations for hindcasting approach using the model for
log(B_OTol +1) at Angle Guignard (ANG85, reservoir site). Adapted from
Kilgour and Stanfield (2006).
Model parameter Model coefficients Site conditions Result
Constant 0.90 1.00 0.90
Maximum depth (Dmax) -0.40 2.40 -0.96
Volume development (Vd) -0.10 1.22 -0.12
LHMS 0.40 0.00 0.00
Hindcast (expected) log(x+1) B_OTol (1) 1.09

Observed log(x+1) B_OTol (2) 1.67
Deviation score (3) (2) - (1) 0.58
Standard deviation (SD) 0.32
Normalized deviation expressed in units of

(3) / SD 1.81
standardized deviation (SDs)
Sites with normalized deviations near zero were considered unimpacted (Baker et al.
2005, Kilgour & Barton 1999). On the other hand, the farther the normalized deviation was
from zero, the greater the impact. Variations of ± 2 standard deviations from the expected
values were arbitrarily interpreted as describing the statistically expected range with no
anthropogenic pressure (Wiley et al. 2003).
To avoid redundancy, if two normalized deviations were highly correlated (Pearson
correlation r > 0.8), we decided to retain only one (Tableau 5. 5). As the final IBI was
expected to be sensitive to all three types of anthropogenic pressures, we gave priority to the
one metric responding to the anthropogenic pressure that had no related indicator otherwise.
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Tableau 5. 5: Pearson correlation coefficients between selected normalized deviations; P ≤ 0.05*. P ≤ 0.01**. P ≤ 0.001***.
Those over 0.8 are in bold. Normalized deviations retained for the final index are in italic and bold. See section 2.4 Candidate metrics
for metrics abbreviations.
Reservoirs
Normalized
N = 67 BPUE Sp_StLith %Sp_StLith %Sp_Pisc Nb_Herb B_OTol B_Omni B_Plankt %B_Lith
deviation
BPUE 1.00 -0.06 -0.05 -0.08 0.33** 0.88*** 0.74*** 0.82*** -0.11
Sp_StLith 0.91*** 0.03 -0.19 -0.07 -0.15 -0.25* 0.14
%Sp_StLith 0.14 -0.24 -0.18 -0.22 -0.33** 0.17
%Sp_Pisc -0.12 -0.19 -0.06 -0.18 0.03
Nb_Herb 0.39*** 0.41*** 0.45*** 0.03
B_OTol 0.75*** 0.86*** -0.11
B_Omni 0.86*** 0.16
B_Plankt -0.19
%B_Lith 1.00
Natural
lakes
Normalized
N = 24 Sp_OIntol Sp_Invert Sp_Plankt B_OIntol B_StLith %B_OTol
deviation
Sp_OIntol 1.00 0.81*** 0.65*** 0.54** 0.88*** 0.18
Sp_Invert 0.84*** 0.37 0.74*** 0.35
Sp_Plankt 0.41* 0.64*** 0.40
B_OIntol 0.64*** -0.16
B_StLith 0.22
%B_OTol 1.00
5.2.7.3.Aggregation of metrics into total scores (FBIs)
First, normalized deviations were transformed into absolute values. Second, these positive
values (ND) were scored with values between 0 and 10 (see Minns et al., 1994), using the
following equation and conditions:
(2.1) if ND ≤ ND95 , Score = [10 × ( ND95 − ND) /( ND95 − NDmin )]
(2.2) else, Score = 0

where NDmin and ND95 are the minimum observed value and 95th percentile, respectively, of
normalized deviations.
5.2.7.4.FBI evaluation
FBIs were calculated for 67 reservoirs and 24 natural lakes by multiplying the sum of
metric scores by 10 N M , N M being the number of scores retained. This procedure bounds the
FBI between 0 and 100. We used the Pearson correlation analysis to test FBIs against the
pressure variables. Finally, FBI ranges were divided into five equal-width intervals: 0≤20 =
bad, >20-40 = poor, >40-60 = moderate, >60-80 = good, and >80 = high (Karr 1981, Minns et
al. 1994).
All statistical analyses were performed using R statistical software (R.2.9.1) as well as
vegan (variation partitioning), pls and caret (statistical models’ validations) R packages (Ihaka
& Gentleman 1996, R Development Core Team 2008).
5.3. Results
5.3.1. Study sites, environmental variables and anthropogenic pressures

The reservoirs included in the present study represented a broad assortment of
characteristics (Tableau 5. 2) and were fairly evenly spread throughout France (Figure 5. 1).
In contrast, natural lakes were concentrated either in mountain areas (e.g., the Alps, the
Massif Central) or near the coast. Natural lakes had significantly longer residence time (771
days vs. 129 days, p < 0.001), lower SLDF (p < 0.001), greater surface area (p < 0.01), lower
water level fluctuations (p < 0.001), and lower SumT (p < 0.01).
Differences between natural and man-made lakes were also clear with respect to
anthropogenic pressures (Tableau 5. 1). Apart from transportation network densities,
urbanized area, and LHMS score, all anthropogenic pressures displayed higher mean values
for artificial lakes than for natural ones. In addition, all agricultural variables reached higher
maximum values – more than twice as high – for reservoirs than for natural lakes. The
negative nutrient balances for natural lakes indicated a deficit of nutrients in the soil surface
of the catchments, whereas their highly positive values for reservoirs showed a nutrient
accumulation in the upstream catchment.
5.3.2. Fish assemblages

Forty-one species were collected, of which 10 were intolerant and 15 were tolerant
(Appendix 4). Regarding trophic groups, invertivores were the most common (37 species),
followed by planktivores (18), piscivores (14), omnivores (11), and herbivores (10). We
collected 17 strictly lithophilic and 11 non-obligatory lithophilic species, as well as eight
strictly phytophilic species. In both natural lakes and reservoirs, Rutilus rutilus (Linnaeus,
1758), Perca fluviatilis (L., 1758), and Scardinius erythrophtalamus (L., 1758) were
widespread (> 70% of the sampling sites) and had a high mean average abundance (Appendix
5). On the contrary, differences were clear between lake types (Appendix 5) for Blicca
bjoerkna (L., 1758), Sander lucioperca (L., 1758), Abramis brama (L., 1758), Cyprinus
carpio (L., 1758), and Gymnocephalus cernuus (L., 1758), which were more frequent in
reservoirs than in natural lakes, and for Squalius cephalus (L., 1758) and Coregonus lavaretus
(L., 1758) which occurred more frequently in natural lakes.
5.3.3. Metrics selection

Overall, 15 out of 73 candidate fish metrics, covering all three groups of functional traits,
displayed a significant response to anthropogenic pressures (Tableau 5. 6). None of them was
derived from the strictly phytophilic trait (Tableau 5. 6). Species richness, diversity indices,
and native status-related metrics were also rejected. None of the candidate metrics indicated a
significant anthropogenic impact on both natural lakes and reservoirs, thereby confirming the
relevance of analyzing each lake type separately.
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variables.
Table entries are the regression coefficient, adjusted R squared and variance
partitioning; var env is the percentage of the total variation attributed to purely
environmental effects, var AP to purely agricultural pressure effects, var UP to
purely urban pressure effects, var LHMS to purely local pressure effects, and
unexpl variation unexplained by the model. Metrics retained for the final index are
in italic and bold. For abbreviations see section 2.4 Candidate metrics.
Adjusted var var var var
AP UP LHMS 2 unexpl
Metric value R env AP UP LHMS
Reservoirs BPUE - - 0.60** 0.57 0.43 0.02 <0 0.03* 0.43
N = 67 Sp_StLith -0.50* - - 0.27 0.14 0.05* <0 <0 0.79
%Sp_StLith -0.30** - - 0.29 0.11 0.08** <0 <0 0.76
%Sp_Pisc -0.20*** - - 0.24 0.05 0.09** <0 <0 0.79
Nb_Herb - 0.06* - 0.22 0.02 0.00 0.05* <0 0.90
B_OTol 0.40* - - 0.64 0.42 0.03* 0.00 0.01 0.38
B_Omni - - 0.70** 0.55 0.4 <0 <0 0.05** 0.51
B_Plankt 0.50** - 0.70** 0.66 0.47 0.04** <0 0.03* 0.35
%B_Lith -0.20* - - 0.23 0.15 0.05* <0 <0 0.83
Natural
Sp_OIntol - - 2.00*** 0.85 0.45 <0 <0 0.12* 0.22
lakes
N = 24 Sp_Invert - - 1.00*** 0.70 0.20 <0 <0 0.23* 0.44
Sp_Plankt - - 1.00*** 0.55 <0 <0 0.00 0.25* 0.61
B_OIntol - -0.70** 0.60*** 0.67 0.34 <0 0.13* 0.22* 0.34
B_StLith - - 0.90*** 0.52 0.03 <0 <0 0.31* 0.74
%B_OTol - 1.00** - 0.54 0.21 0.06 0.40** <0 0.63
*significant at 0.05 level; **significant at 0.01 level; ***significant at 0.001 level
Six metrics displayed a significant response to anthropogenic pressures in natural lakes

(Tableau 5. 6). Two of them, %B_OTol and B_OIntol, displayed a response to urbanization
stress (40% and 13% of their variance explained exclusively by UP). Apart from %B_OTol,
all other candidate metrics were related to LHMS (from 12% to 31% of variance explained
exclusively by local pressures) (Tableau 5. 6). No metric displayed a response to agricultural
pressure for natural lakes.
For the reservoirs, nine metrics were selected, including BPUE (Tableau 5. 6). The most
significant effects were attributable to AP. Indeed, three metrics related to mineral substrate
spawning habitats (Sp_StLith, %Sp_StLith and %B_Lith), two related to community trophic
structure (%Sp_Pisc and B_Plankt), and B_OTol, displayed a response to agricultural stress
(from 3% to 9% of the variance explained exclusively by AP). LHMS had significant effects
on B_Omni, BPUE and B_Plankt, with, respectively, 5% and 3% of their variance explained
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exclusively by local pressures. Finally, apart from Nb_Herb, no significant effect caused by
UP was detected (Tableau 5. 6).
All linear models included at least one significant coefficient for environmental variables
(not shown here), thereby confirming that environmental patterns have to be taken into
account when studying the broad-scale relationships between fish-based metrics and
anthropogenic pressures. For the natural lakes, this is particularly true for Sp_OIntol and
B_OIntol (Tableau 5. 6), with 45% and 34%, respectively, of their variance explained
exclusively by environmental variables. For the reservoirs, this mostly concerns B_Plankt,
BPUE, B_OTol, and B_Omni, with 47%, 43%, 42%, and 40%, respectively, of their variance
explained solely by environmental variables.
5.3.4. Hindcast modeling and normalized deviation transformation

The final selection included three scores for natural lakes and six for reservoirs (Tableau
5. 6).
The majority of the reservoirs were little impaired (Figure 5. 2a). Most of the normalized
deviations were distributed within ± 1 SD of the expected value (zero), and only a few were
out of the ± 2 SD range. In contrast, for natural lakes (Figure 5. 2b), most of the normalized
deviations were distributed beyond ± 2 SD from the expected values, thereby suggesting that
the majority of sites were more impaired than reservoirs were. In addition, for both types of
lake, all selected metrics had some normalized deviations that exceeded ± 2 SD in some sites,
indicating severe impacts (Figure 5. 2). Across metrics, from 17% to 83% of the natural lakes
deviated from the expected value, whereas only 4%-38% of the reservoirs exceeded this
value.
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Fig. 5.2a
Fig. 5.2b
Figure 5. 2: Boxplot of deviations (i.e., observed metric – expected metric), scaled in terms
of standard deviations for (a) reservoirs and (b) natural lakes in France.
Black lines indicate ± 2 SD thresholds. For labels see section 2.4 Candidate
metrics.
5.3.5. FBI evaluation

The final FBIs had a potential range from 0 to 100. For reservoirs, FBIs were significantly
correlated with agricultural pressures (r = 0.39, p < 0.005) (Figure 5. 3a), whereas for natural
- 106 -
lakes they were correlated with urban (r = 0.52, p < 0.01) and local (r = 0.87, p < 0.001)
pressures (Figure 5. 3b&c).
Fig 5.3a
Fig 5.3b
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Fig 5.3c
Figure 5. 3: Relationships between: (a) FBI and agricultural pressure for reservoirs (n=67);
(b) FBI and LHMS for natural lakes (n=24); and (c) FBI and urban pressure for
natural lakes (n=24).
Regression lines were drawn only when the regression was significant at the 0.05
level. Pearson’s correlation coefficients (r) and p-values (p) are indicated.
5.4. Discussion
The present study demonstrated that hindcasting modeling of fish-based metrics made it
possible to assess the current condition of lakes. A set of non-redundant metrics displaying
significant responses to anthropogenic pressure was selected and combined into an index of
biotic integrity for French lakes.
5.4.1. Metrics selected for the FBI

Karr et al. (1986) suggested six criteria that IBIs should meet to ensure their ability to
detect environmental degradation in a freshwater ecosystem. The indices must be (i)
biological, (ii) interpretable across many trophic levels, (iii) sensitive to environmental
conditions being monitored, (iv) responsive over their range to intended uses, (v) reproducible
and precise when defined with acceptable limits, and (vi) of low variability.
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The lake FBIs developed in the present study are based entirely on combining fish survey
data and species ecological features, so they meet criterion (i). The inclusion of trophic
metrics reflects the food available to fish and therefore relates to various compartments of the
food web (ii). These FBIs partially meet criterion (iii), as they respond to catchment-scale
and/or local pressures. The within-FBI-class analyses (unpublished results) show their
sensitivity in both types of lakes (iv). Carefully described and standardized field sampling
methods ensure reproducibility (v). The last criterion (vi) can be considered met here because
of appropriate sampling and a statistical control of the natural variability in the metrics across
sites.
Metrics based on tolerance, reproduction, and trophic guilds were retained. The tolerance-
related metrics displayed clear responses to both urban and local anthropogenic pressures
exclusively for natural lakes (Tableau 5. 6). B_OIntol and %B_OTol were the two biomass-
related metrics that decreased and increased, respectively, with increasing urban pressures.
These relationships are congruent with those found in US lakes (Drake & Pereira 2002,
Jennings et al. 1999b, Whittier 1999). In addition, the richness and biomass of intolerant fish
species (i.e., S_OIntol and B_OIntol, respectively) have been found to increase with local
anthropogenic pressures. Most IBIs developed on lentic systems include some measure of the
relative dominance of tolerant and/or intolerant species to assess lake status (Belpaire et al.
2000, Drake & Pereira 2002, Simon & Lyons 1995), but usually not expressed as biomass.
For reservoirs, all trophic guilds appear to be related to anthropogenic pressures (Tableau
5. 6), although some were removed from the final index to avoid redundancy. This strongly
suggests that a fish-based index is a relevant indicator of eutrophication (Appelberg et al.
2000, Drake & Pereira 2002, Tammi et al. 2001). Nutrient inputs have long been considered
major drivers of fish communities in these ecosystems because eutrophication implies oxygen
depletion, organic sediment accumulation (Carpenter et al. 1998b, Harper 1992), and a
transfer of primary productivity from macrophytes to phytoplankton (Leach et al. 1977).
Despite the complexity of these processes, the response of the fish communities analyzed here
to agricultural land use can be summarized by a decrease in piscivores and an increase in
planktivores. The absence of trophic metrics in the final index for natural lakes is puzzling,
but does not necessarily indicate their irrelevance. Indeed, in the set of lakes studied here, the
natural ones undergo far less eutrophication than the artificial ones. A limited gradient of
agricultural pressures and the small sample size may have prevented us from identifying the
expected biological response to the eutrophication of natural lakes because of the model’s
reduced statistical power.
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Reproduction-related metrics also displayed clear responses to pressures exclusively for
reservoirs (Tableau 5. 6). As expected from the literature survey, we observed negative
relationships between %Sp_StLith or %B_Lith and reservoir quality (Hickman &
McDonough 1996). For natural lakes, the absence of reproductive-based metrics in the final
FBI can be explained considering the biological traits of the species. The fish species driving
the relation between the metrics derived from lithophilic and strict lithophilic traits and
anthropogenic pressures were in the majority Rutilus rutilus (L., 1758) and Perca fluviatilis
(L., 1758). These species also happen to be overall tolerant to degraded lake conditions,
thereby creating a strong correlation between the metrics initially chosen to reflect
complementary aspects of communities. Consequently, B_StLith was removed to avoid
redundancy among metrics (Tableau 5. 5), despite its strong response to local anthropogenic
pressures.
5.4.2. Evaluation of the structure of the FBI

Whereas the FBI for reservoirs fully meets the set of criteria, recommended by Karr et al.
(1986), the FBI developed for natural lakes in France suffers from the absence of a metric
responding to the agricultural pressures that have been proved to affect trophic-related metrics
in other contexts (Appelberg 2000, Drake & Pereira 2002, Jennings et al. 1995, Karr &
Dionne 1991). This absence is likely due to the low agricultural land use in the catchments of
the sampled lakes (see section 4.1 Selected metrics for use in FBI above).
Minns et al. (1994) recommended reducing the redundancy among metrics when
developing IBIs in order to optimize the effort required to obtain the information. Despite our
strict selection process, correlations among the metrics included in the FBI for both types of
lakes brought out substantial redundancy. However, we believe this cannot be avoided for
several reasons: (i) the number of species was only 41, so the most widespread tend to
increase correlations between traits, (ii) habitat availability exerts constraints on several
aspects of species ecology (e.g., headwater communities encompass mainly cold water
species, benthic invertivores, and lithophilic species), and (iii) phylogenetic relatedness
associated with the inheritability of certain characteristics imposes evolutionary constraints on
the suite of traits present in the regional pool of species and consequently on those making up
local communities.
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5.4.3. Analytical procedure issues and limitations of hindcast models
According to the WFD, reference conditions can be modeled when insufficient numbers
of reference sites are available. In this case, the hindcasting approach may be useful (Baker et
al. 2005). To our knowledge, no previous study has attempted to use this methodological
approach to assess the ecological status of water bodies to meet the requirements of the WFD.
No correction was made for multiple testing in this study. Since the colinearity between
fish-based metrics – particularly within a given biological trait – is high, there is far less real
testing than the number of metrics considered. Hence, p-values that are of the order of
magnitude of the significance threshold should be considered very carefully.
Using hindcasting modeling, we assumed that the reference condition was adequately
simulated by setting all anthropogenic factors to zero. However, in the WFD, the reference for
a given lake corresponds to the state in “the absence of anthropogenic pressures, or nearly
undisturbed conditions” (Stoddard et al. 2006). Therefore, the method implemented here was
even more restrictive in the assessment of the reference than in the spirit of the WFD, leading
to overestimated FBI values.
Hindcasting to a pristine condition is not recommended in the absence of data on
minimally disturbed or reference sites (Kilgour & Stanfield 2006) and gives its best results
when the data set covers the whole range of the anthropogenic pressure gradients. The data set
of natural lakes processed here did not include very impacted sites and the range of
agricultural pressure gradient was restricted to low pressures in comparison to reservoirs. This
hampered our ability to assess agricultural impacts on fish communities. Furthermore, for
both types of lentic ecosystems, urban pressures contributed to the models only very slightly,
perhaps because of limited population density in most catchments. In addition, the accuracy
of the proxies used to describe urban pressures may be insufficient to show urban impacts.
Complementary to the classic catchment-scale pressures (Drake & Pereira 2002, Jennings
et al. 1999b, Lyons et al. 2000, Thoma 1999), we included an assessment of local habitat
degradation (Rowan et al. 2006). The LHMS strongly contributed to models and improved
them compared to a previous study (Launois et al. 2011). This demonstrates that restricting
pressure evaluation to GIS-derived variables is not sufficient to develop a FBI, at least in the
French context. It also stresses the impact of littoral habitat alteration on fish communities.
In Baker’s et al. (2005) study, biological metrics are not or hardly sensitive to
anthropogenic pressures. The other metrics which are considered in that study are physico-
chemical ones. For these metrics, the authors can be fairly confident in the a priori direction
of the response to stressors. Besides, for this kind of metrics, it could be relevant to consider
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conditions “better that pristine”. Karr and Dudley (1981) defined the concept of ecological
integrity of freshwater ecosystems as their ability to support and maintain “a balanced,
integrated, adaptive community of organisms having a species composition diversity and
functional organization comparable to that of the natural habitat of the region.” Therefore,
“better conditions than pristine” has no real meaning for biological metrics. Indeed, any
significant durable deviation from the expected state of communities in a given environment
must be considered as impairment whatever the direction of the deviation.
In the hindcasting approach, the explicit inclusion of stressors in models means that the
‘expected’ direction is somewhat known a posteriori, since after having fitted models to
calibration dataset, the sign of pressure effects are known. However, hindcasting scores are
not exactly pressure effects but it includes model residuals. In our opinion, deviations in the
opposite direction of the pressure effect, which is then mostly due to residuals, should not be
considered as a misspecification of the environmental part of the model, though it could
possibly be improved, but rather to other omitted pressures or to a misspecification of the
anthropogenic part of the model. The environmental part of the model is indeed what provides
the expectation of the reference state. Its accuracy can be discussed but we have to
hypothesize that the model is well specified to address the hindcasting approach to integrity
index development. This is why we have decided to use absolute values, considering
deviations in the opposite direction as impairment too, but impairment due to other unknown
pressures.
In the case of a metric responding positively to one pressure and negatively to another – in
this study, only B_OIntol – antagonist effects of both stressors can in some lakes be involved
but we have preferred retaining this metric because of its sensitivity to stressors, even if it
could be discussed. In this case, the use of the absolute value of the deviation is the only
option.
Finally, in the absence of adequate data, we did not evaluate the statistical significance of
the deviation from historical reference values. Instead, we arbitrarily set the threshold of
impairment for all metrics at ± 2 SD. Although this method is becoming popular (Kilgour &
Stanfield 2006), and could, be thought of as associated with 95% confidence intervals (Wiley
et al. 2003), the reader has to keep in mind that it includes a certain subjectivity. The
reproducibility and precision of the index will be usefully tested when new fish data are
added.
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5.5. Conclusion
The development of fish-based assessment tools for lentic systems is a necessary step in
the implementation of the WFD. The present study proves that this can be done in some lake-
poor regions when standard field methods are implemented. Many IBI studies restricted
pressure evaluation to broad-scale variables. We believe that the assessment of local pressures
is worth the additional cost of a field survey.
Methodological difficulties, as well as the amount of costly data required, have so far
hampered the development of lake IBIs. Our study provides a significant progress in this area.
Although the FBIs developed here have certain weaknesses, the ultimate objective of
monitoring and assessment tools should be kept in mind; protecting the environment. The
imperfection of tools should not be made a pretext for discarding ambitious environmental
policy. Ecological knowledge is sufficient to provide guidelines to improve the sustainability
of water and aquatic resources.
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CHAPITRE 6
Conclusions et perspectives
La prise de conscience généralisée des services rendus par les écosystèmes d’eau douce a
conduit à une politique de l’eau volontariste en Europe. D’ambitieux objectifs de qualité ont
été définis (European Community 2000). Pour autant, sur les plans d’eau, les outils pour
atteindre ces objectifs restent encore en partie à développer. Ce mémoire démontre que
l’ichtyofaune peut contribuer efficacement à l’établissement d’un diagnostic de l’état
écologique des milieux lentiques.
Analyse critique des données

Mon travail s’est concentré sur le développement d’un indice multimétrique applicable
aux plans d’eau français. Il s’appuie sur des travaux antérieurs qui ont mis en évidence un
certain nombre de relations entre les caractéristiques des assemblages piscicoles lacustres et
des facteurs environnementaux (Irz 2006, Pronier 2000). Néanmoins, une partie de mon
travail a consisté à améliorer la qualité des données nationales disponibles. En effet, les
variables environnementales naturelles telles que la géologie et l’hydrologie ont été
identifiées comme des facteurs influençant, directement ou indirectement, la structure des
peuplements piscicoles lacustres (Alpay et al. 2006, Brousseau et al. 1985, Rowan et al. 2003,
Thornton 1990, Winfield 2004). Leur effet n’avait jamais été étudié à l’échelle nationale et
pourtant, nous avons montré ici que la prise en compte de ces variables contribue à mieux
expliquer la variabilité naturelle de l’ichtyofaune. Par exemple, l’étude menée par Irz et al.
(2008) révélait que la variabilité liée à l’environnement représentait entre 8% et 19% de la
variabilité des métriques de tolérance (tolérance et intolérance globales) et du caractère
invertivore des espèces piscicoles. Les deux études présentées aux chapitres 4 et 5 de cette
thèse, avec la prise en considération de variables hydrologiques et géologiques, montraient
que la variabilité environnementale (hydrologie et géologie) expliquait entre 18% et 45% de
la variabilité de ces mêmes métriques. Il en découle que la part de la variabilité non expliquée
des métriques a diminué entre l’étude menée par Irz et al. (2008) et celle conduite ici. Notons
toutefois que les facteurs zoogéographiques pertinents (dispersion postglaciaire,
hydroécorégions, pH, etc.) pour interpréter la richesse spécifique et les indices de diversité
n’ont pas été inclus dans ces études (Griffiths 1997, Irz et al. 2004a). La prise en compte
future de ces facteurs pourrait permettre d’améliorer encore la part de variabilité des
métriques expliquée par l’environnement.
De plus, le développement d’indicateurs nécessite une bonne caractérisation des pressions
anthropiques, que nous avons dû entièrement mener. Un effort particulier a porté sur la
pondération des pressions liées aux usages du sol sur le bassin versant. Pour cela, les surplus
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en phosphore ont été pondérés par la pente, la géologie, la battance, la pédologie, l’occupation
des sols et le climat, comme préconisé par Coale (2000). Nous avons également exploité des
données standardisées de pressions locales, pressions rarement utilisées et/ou bien décrites
dans de telles approches de type pressions/impacts.
Pour ce qui est de l’ichtyofaune, cette étude a pu s’appuyer sur des données collectées en
application d’une méthode de pêche standardisée (C.E.N. 2005a). Cela permet de collecter
des données homogènes et comparables. Notre hypothèse de départ, à savoir que les données
de pêche sont comparables entre les lacs, peut être discutée car nous travaillons sur plusieurs
espèces, dans plusieurs lacs de plusieurs écorégions différentes ce qui pourrait engendrer des
différences de sélectivité. Toutefois, comme en attestent les travaux de Larranaga (2010), la
sélectivité des filets influence peu les captures et on peut considérer que la variabilité est
réduite du fait que les engins sont identiques, les périodes de pêche comparables et l’ensemble
du lac toujours prospecté par des poses aléatoires sur des strates définies.
Un problème récurent dans ce type d’approche concerne la méconnaissance de
l’importance de la gestion piscicole sur ces milieux que l’on sait très manipulés (Argillier et
al. 2002a). En effet, même si l’on considère que toutes les espèces échantillonnées sont
capables de se maintenir dans le milieu, il n’en demeure pas moins que l’abondance de
certaines de ces espèces peut être affectée par la gestion, de même que leur structure en taille.
Malgré des avancées certaines sur la qualité du jeu de données, il faut reconnaître que la
collecte de toutes les variables environnementales et de pressions anthropiques a été
particulièrement laborieuse. L’ensemble des informations utilisées ici n’est pas disponible
pour l’ensemble des lacs français. La poursuite de l’étude nécessitera donc, parallèlement à
l’acquisition des données biologiques, encore un effort important pour tendre vers la
caractérisation des pressions anthropiques. Notons également que de ce point de vue, la
situation nationale n’est pas une situation isolée. En effet, des difficultés analogues pour
décrire les caractéristiques environnementales des plans d’eau ont été rencontrées dans des
travaux menés au niveau européen (Pedron et al. 2010).
Choix de la méthode
La majorité des indices poisson qui ont jusqu’à présent été développés pour les milieux
lacustres reposent sur (i) l’hypothèse d’homogénéité des caractéristiques environnementales
entre les plans d’eau (e.g., Drake and Pereira 2002) et (ii) une approche de modélisation sur
sites de référence (RCA, voir Stoddard et al. 2006 ; e.g., Holmgren et al. 2007). Dans le
contexte français, nous avons montré qu’il serait impossible de définir des groupes de plans
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d’eau comprenant un nombre significatif de lacs présentant des conditions environnementales
comparables. Ainsi, cette approche typologique pourtant soutenue par la DCE (European
Commission 2002) et largement utilisée pour développer des outils de bioindication (Ferreol
et al. 2005, Marchant et al. 1999, Omernik 1995, Sandin 2003), s’avère peu adaptée. Par
ailleurs, l’application de cette méthode est rendue d’autant plus difficile que la variabilité des
pressions anthropiques au sein de la plupart des types de plans d’eau est très faible et non
comparable entre les types. Ces distributions hétérogènes de sites plus ou moins impactés, au
sein et entre les groupes, posent alors des problèmes quant à la comparaison des scores des
différents indices poisson à l’échelle nationale. C’est la raison pour laquelle nous avons
envisagé d’appliquer une méthode alternative dite « site-spécifique » (Roset et al. 2007). Le
contexte national marqué par la quasi-absence de lacs de basse altitude avec un degré
d’altération négligeable, n’a pas permis l’utilisation de l’approche usuelle par conditions de
référence (Bailey et al. 1998). Il se pose également la question de savoir comment définir une
« retenue de référence ». Contrairement à ce qu’il est préconisé dans la DCE, les travaux
menés dans le cadre de cette thèse, ainsi que des travaux antérieurs (Irz et al. 2006) ont
démontré qu’il n’était pas possible d’utiliser les lacs naturels comme référence pour évaluer le
potentiel écologique des retenues. Ainsi, le choix méthodologique s’est porté sur une
approche innovante, l’hindcasting, basée sur un modèle statistique, qui s’est révélée être tout
à la fois pertinente et compatible avec les exigences de la DCE [Annex, II, 1.3 (i-vi)]
(Anonymous 2003a). En effet, nous avons montré que l’utilisation de cette approche
permettait de mettre en évidence de fortes corrélations entre certaines caractéristiques des
peuplements piscicoles et les perturbations subies par les retenues et les lacs naturels français.
En outre, l’utilisation de cette approche, qui fut précédemment testée dans les milieux lotiques
sur les macroinvertébrés benthiques, l’ichtyofaune et des paramètres chimiques (Baker et al.
2005, Kilgour & Stanfield 2006), a permis de quantifier le degré de perturbation subi par les
plans d’eau français. Toutefois, comme précisé dans le chapitre 5, des limites sont également
à formuler quant à l’utilisation de cette méthode pour ce qui concerne les lacs naturels.
Notamment, les modèles développés pour ces lacs ne peuvent être utilisés pour réaliser des
extrapolations. Ils ne peuvent être utilisés que pour prédire des conditions écologiques à partir
des jeux de données de calibration. Nous pouvons donc penser que les données
prochainement acquises, notamment à partir du réseau de contrôle opérationnel concernant les
sites qui risquent de ne pas atteindre le bon état écologique en 2015
(Ministère de l'Ecologie et du Développement Durable 2006), permettront d’améliorer la
qualité des indices. Ainsi, comme nous l’avons vu, un des intérêts de la méthode prédictive
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proposée est d’adopter une approche plus précise des conditions de référence et de prendre en
compte la variabilité environnementale des métriques. Elle permet donc d’échapper au
recours à la recherche de conditions de référence « historiques » difficiles à établir, du moins
du point de vue de la description de l’abondance des espèces dans les milieux, critère de
bioindication requis par la DCE.
Construction de l’indice
A partir des données nationales, l’application de l’approche par hindcasting dont les
forces et les faiblesses ont été discutées, a permis la sélection d’un certain nombre de
métriques rendant compte des pressions qui impactent les retenues artificielles et les lacs
naturels.
Parmi les métriques candidates, nous avons inclus des indices de diversité très utilisés
dans les développements d’indices macroinvertébrés (Garcia-Criado et al. 2005, Smith et al.
1999, Trigal et al. 2006), mais peu utilisés pour l’ichtyofaune sauf dans les études
scandinaves (Appelberg et al. 2000, Holmgren et al. 2007, Tammi et al. 2001). Ces indices de
diversité se sont avérés fortement corrélés les uns aux autres, et aucune réponse aux pressions
anthropiques n’a pu être mise en évidence dans nos analyses. Si l’on considère, par ailleurs,
que leur interprétation est difficile (Holmgren et al. 2007), ce type de métrique n’est pas
forcément pertinent dans la construction des indices.
Globalement, nos études confirment les résultats de travaux antérieurs conduits dans
d’autres environnements. En particulier, elles montrent l’intérêt d’utiliser des métriques
utilisant les traits d’histoire de vie, ainsi que les réponses des métriques aux pressions
anthropiques du bassin versant (Appelberg et al. 2000, Drake & Pereira 2002, Irz et al. 2008,
Jennings et al. 1995, Karr & Dionne 1991). Cependant, elles mettent aussi en évidence
l’impact des pressions locales sur l’ichtyofaune. Il est reconnu que certaines zones lacustres,
telles que les zones littorales, peuvent être influencées par des pressions locales in situ, des
types d’occupation du sol à proximité immédiate des plans d’eau, et par des intrants
d’origines biotiques et/ou abiotiques (Minns et al. 1994). Toutefois, les réponses de
l’ichtyofaune à ce type de pressions n’a actuellement été que peu étudiée à large échelle (mais
voir Jennings et al. 1999a ; Moyle & Randall 1998 ; Thoma 1999). La mise en évidence de
ces relations est particulièrement intéressante dans la mise en œuvre des diagnostics
écologiques. En effet, la plupart des indicateurs qui évaluent l’état écologique des plans d’eau
répondent majoritairement à l’eutrophisation et à l’acidification (Appelberg et al. 2000, Beck
& Hatch 2009, Tammi et al. 2001). Cependant, un point mis clairement en évidence durant les
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analyses a été la difficulté de décrire précisément les réponses des indices aux différents types
de pression anthropiques. Il s’avère, en effet, que la plupart des plans d’eau sont soumis à des
pressions multiples, s’exerçant à plusieurs échelles spatiales. Ainsi, les sites perturbés le sont
à la fois par des altérations de la qualité de l’eau et des conditions hydromorphologiques. De
plus, les conséquences des interactions entre pressions anthropiques sur la biologie demeurent
actuellement quasiment inexplorées. Il restera donc difficile d’établir des relations pression /
impact spécifiques à chaque pression.
Perspectives
Dans cette thèse, nous avons testé la réponse de métriques classiquement utilisées en
bioindication lacustre. Cependant, dans la perspective d’une amélioration de l’indice, on peut
envisager l’inclusion de métriques complémentaires, telles que des métriques portant sur les
espèces piscicoles hybrides, sur les espèces invasives ou sur la santé des poissons (Karr &
Chu 1999). Des travaux additionnels devraient aussi être menés, comme le suggèrent Berrebi
dit Thomas et al. (1998) et Whittier et al. (2007), sur les calculs des métriques piscicoles,
notamment en soustrayant de ces métriques toutes les espèces tolérantes. Cette modification
permettrait d’éviter que la sensibilité d’un trait biologique spécifique ne soit masquée par
l’hétérogénéité des espèces piscicoles contribuant au calcul de la métrique associée. Une
dernière idée serait d’inclure des indices de diversité fonctionnelle. Même si les connaissances
dans le domaine sont encore limitées sur le poisson, la diversité fonctionnelle est désormais
largement reconnue comme étant un meilleur prédicteur de la productivité et de la
vulnérabilité des écosystèmes que la diversité spécifique (Diaz & Cabido 2001, Heemsbergen
et al. 2004, Hulot et al. 2000, Tilman et al. 1997).
Les travaux initiés dans cette thèse se poursuivent dans un cadre européen. L’applicabilité
des indices sera testée dans d’autres régions biogéographiques. Ce travail est programmé au
niveau de l’écorégion Alpes dans le cadre des travaux d’intercalibration, ainsi que sur les
retenues européennes dans le cadre du projet européen WISER
(http://www.wiser.eu/background/lakes/). Par ailleurs, des tests plus approfondis devront être
menés sur l’indicateur en termes de robustesse (ré-échantillonnage, inclusion de nouvelles
données piscicoles et/ou de pressions) et d’évaluation du risque de mauvais classement (calcul
d’incertitudes associées aux modèles) (Noges et al. 2009). Afin d’observer le comportement
de l’indicateur dans le cadre des futurs suivis à long terme, il sera également indispensable,
lorsque nous disposerons de suffisamment de résultats issus des réseaux de suivis DCE,
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d’évaluer la variabilité interannuelle et les capacités de détection des tendances, points de
rupture, etc.
Enfin, un indice qui résume la qualité de l’écosystème, dans des conditions bien
déterminées, peut se révéler être un vecteur efficace d’information auprès du grand public et
donc un outil de gestion concertée du milieu naturel (Simon 1999). Toutefois, si cet indice
permet d’évaluer l’état écologique des plans d’eau, il n’autorise pas de faire l’économie
d’études spécifiques et détaillées pour la définition des mesures de restauration à prendre sur
un site donné.
En accord avec la perspective écosystémique prônée par la DCE, une étape ultérieure
consistera aussi à comparer, au niveau national, la cohérence / complémentarité des résultats
des diagnostics basés sur les différents compartiments biologiques pour un même type de
masse d’eau (lacs, retenues, rivières, estuaires).
Pour terminer, il est important de noter qu’actuellement, les différents milieux sont
considérés de manière isolée. Or, ces écosystèmes lotiques et lentiques, comme le faisait déjà
remarquer dans ses études Irz (2006), font partie intégrante d’une mosaïque de paysages
(Wiens 2002), et sont interdépendants car échangeant des organismes vivants (Jackson et al.
2001a), de la matière et de l’énergie (Kling et al. 2000). La prise en compte des relations entre
plans d’eau et cours d’eau et des effets réciproques de la gestion d’un type de milieu sur
l’autre, représente également une perspective intéressante de recherche pour l’amélioration de
la gestion intégrée des hydrosystèmes.
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ANNEXES
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Annexe 1 : Caractérisation des pressions anthropiques s’exerçant à l’échelle régionale
Cet article en cours de révision présente les impacts des pressions anthropiques sur
l’ensemble des organismes vivants lacustres. En effet, compte tenu des interactions entre
niveaux trophiques, on s’est intéressés aux effets des pressions sur l’ensemble des
communautés biologiques lacustres, plutôt que de se restreindre aux poissons en particulier.
A review of the anthropogenic pressures undergone by lake biological communities and

of their impacts (en révision)
Lionel Launois and Christine Argillier
1. Introduction
Natural lakes and reservoirs account for more than 90% in volume of the overall surface
freshwater on Earth (Shiklomanov 1993). In Europe, there are an estimated 500,000 natural
lakes with surface areas greater than 0.01 km2 (Fritzner et al. 2001, Kristiansen & Hansen
1994) and approximately 10,000 reservoirs (EEA 1995, World Commission on Dams 2000).
Moreover, new reservoirs being regularly created for recreational activities, amenity use, or
water supply, this number may very well be higher today.
The importance of these natural lakes and reservoirs for conservation and resource
management has been widely acknowledged. Indeed, lakes have a number of ecological and
economic functions related to water resources, and the human benefits gained from these
lakes include: (i) water supply for drinking, agriculture, industry, and hydroelectricity
production; (ii) fish production, which serves as a resource for food and cash income; and (iii)
aesthetic value, which attracts tourists, supports the aquarium trade, and has other
noneconomic benefits. Nevertheless, over the last few centuries the lake environment has
increasingly been modified by anthropogenic pressures that may threaten these resources
(Mensing et al. 1998). Moreover, these pressures are expected to continue to increase in
surface waters in the future due to rising population densities, farming and industrialization
(Bragg et al. 2003, Elias & Meyer 2003, Jackson et al. 2001b, Naiman et al. 1998).
Consequently, the health of lentic ecosystems is increasingly at risk, raising society’s
awareness of the need to preserve freshwaters quality. Concomitantly, there is also a need for
monitoring the health of lakes subjected to anthropogenic pressures (Carpenter 2002, Pace
2001). Implementing such surveillance requires good knowledge of the relationships between
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anthropogenic pressures and biological communities, and a good understanding of the
processes involved.
Understanding the relationships between anthropogenic pressures and lake biological
communities presents many advantages, particularly for the development of quantitative
dose–response relationships, classification tools and models, for effective lake management
practices (Solimini et al. 2006). First, determining the impacts resulting from the various
pressures on lake environmental parameters allows aquatic scientists to select undisturbed
environmental variables in order to define reference conditions on lakes (European
Community 2000, Noges et al. 2009, Ostendorp 2004b, Solheim 2005). In addition, reviewing
these relationships helps scientists develop suitable pressure–impact relationship models for
bioindication, by explaining and quantifying the deviation between observed and reference
biological assemblages in relation to human disturbances (Van de Bund & Solimini 2006).
Finally, focusing on anthropogenic pressures and their impacts contributes to effective
conservation and management of lake ecosystems, perhaps the most valuable potential result
(Annadotter et al. 1999, Born 1979, Carpenter & Cottingham 1997, Cooke et al. 2005, Martin
et al. 2000, Shapiro 1979). Indeed, understanding both the causes and the consequences of the
impairment of biological integrity and proposing a remedy are now key components of lake
management programs (Novotny et al. 2005). With a better understanding of the relationships
between biological communities and pressures, water managers and civil engineers will be
better informed in assessing the risks of anthropogenic impacts on lake biodiversity (Novotny
et al. 2005).
The objective of this paper is to review the relationships between the different lake
biological communities and anthropogenic pressures commonly described in the literature.
This review focuses mainly on strong direct impacts of anthropogenic pressures on lake biota.
Given the wide variety of anthropogenic pressures, this review has been organized by the type
of pressures on lakes: (i) chemical pressures on lake biota, which have been demonstrated
from both experimental laboratory studies and studies in natural lake ecosystems, (ii)
morphological alterations, (iii) hydrological modifications on various biological communities
in lakes, and (iv) biological manipulations on lake biotic communities, mainly fish
manipulations in lentic ecosystems.
2. Impacts of chemical pressures on lake biotic communities (Tableau 6. 1)

The wide array of chemical pollutants entering lake ecosystems along with artificial land
use runoff can cause numerous biological effects (Tammi et al. 2003). Among these chemical
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pollutants, nutrient loading, i.e., eutrophication in lakes, appears to be the most common
pressure affecting most lakes worldwide (Carpenter et al. 1998a, Vitousek et al. 1997, Watt et
al. 2007). In North America and Scandinavia, artificial acidification has also been considered
a critical pressure for aquatic communities (Driscoll et al. 2001, Moiseenko 2005). Today
scientists are also paying attention to other chemical pressures, focusing most particularly on
the increasing quantity of heavy metal and pharmaceutical pollutants entering lakes
(Demirezen & Aksoy 2004, Giguère et al. 2005, Versteeg et al. 2005). Different chemical
constituents impact lake ecosystems and their biological communities in different ways
(Ostendorp et al. 2004a).
Effects of nutrient inputs (Figure 6. 1a)

For the last several decades, many authors have described the consequences of nutrient
loading in lakes. The first aquatic organisms directly concerned by nutrient enrichment in
lakes are primary producers, i.e., phytoplankton (Noges et al. 2009), through modifications in
their nutrient assimilation capacity and metabolism (Anderson et al. 2002, Chróst & Overbeck
1987). The most obvious consequences are a shift in species composition (Solimini et al.
2006). Green algae species (e.g., Scenedesmus), diatoms (e.g., Stephanodiscus), and many
groups of cyanobacteria (Microcystis, Aphanizomenon, Anabaena, etc.) have been shown to
be positively impacted by increasing nitrogen (N) and phosphorus (P) concentrations
(Anderson 1989, Coveney et al. 1977, Horstmann 1975, Skulberg et al. 1984). Conversely,
species of chrysophytes (e.g., Dinobryon), desmids (e.g., Cosmarium), and some diatoms
(e.g., Cyclotella comensis) are negatively impacted by increasing nutrient concentrations in
lakes (Alefs & Müller 1999, Carney & Sandgren 1983, Coesel 1982, Oesel et al. 1978).
Furthermore, nutrient enrichment is also provoking an increase in the frequency and intensity
of algae blooms during summer (Carpenter et al. 1998c, Smith et al. 2006). Harmful
cyanobacteria algal blooms are the most common and a major threat to freshwater ecosystems
(Anderson et al. 2002).The development of such large biomass blooms most often leads to
low-oxygen events that suffocate and kill fish and bottom-dwelling organisms (Bernhard et al.
2001, Oberemm et al. 1999). In addition, cyanobacteria blooms produce hepatotoxins (e.g.,
microcystins [MCs]), a substantial threat for target organisms such as mollusc gastropods
(pulmonate Lymnaea stagnalis, prosobranch Potamopyrgus antipodarum, etc.), because MCs
principally induce a decrease in growth and fecundity and a MC accumulation in gastropod
tissues (Lance 2008). Finally, high biomass blooms can also block sunlight penetration into
the water column, preventing growth of beneficial algae.
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Phytoplankton communities are the first layer in the “trophic cascade” (Carpenter et al.
1985) and their modification will necessarily impact other lake biological communities. This
was demonstrated in experiments conducted in mesocosms. It was shown for example that
nutrient additions increased densities of most phytoplankton taxa but also significantly
increased the production, growth, and densities of bacterioplankton, nanoflagelates, copepods,
benthic invertebrates, and fish throughout the food web (Breitburg et al. 1999).
Moreover, phytoplankton responses to nutrient enrichment result in an increase in lake
turbidity and oxygen deficiency in bottom waters (Noges et al. 2009), which consequently
modifies habitats for other organisms. Macrophyte cells, which are highly influenced by the
transmission of light, can deteriorate or die if lake turbidity impacts photosynthetic processes
(Hough & Fornwall 1988). Depending on a specie’s sensitivity to light and oxygen,
macrophyte composition and development will also be more or less affected (Sculthorpe
1967, Van Donk & Van de Bund 2002). Similarly, because of differences in sensitivity
between species, a decrease in oxygen conditions would also be reflected by changes in the
structure and the abundance of benthic macrofauna (Brinkhurst 1974, Lang 1999).
Eutrophication may also indirectly result in extensive fish kills due to deoxygenation of lakes
(Dybas 2005, Jeppesen et al. 1998), reducing fish recruitment due to incubating fish egg
suffocation (Wilkonska & Zuromska 1982) or altering the balance of condition-specific
competition especially for salmonids (Persson 1991, Winfield 2004). Thus, composition and
biomass of fish communities can be impacted by the indirect effects of nutrients on
phytoplankton.
Nutrient loading also directly affects macrophyte communities. Excessive growth of
aquatic macrophytes (especially during bloom periods) have often been reported (Carpenter et
al. 1998a, Carpenter et al. 1998b, King & Mace Jr 1974, Lachavanne 1985, Ostendorp et al.
2004a). This was explained by nutrient accumulation and retention in sediments (Fillos &
Swanson 1975, Serruya et al. 1974) directly available to the macrophytes through their roots
(Barko & Smart 1986, Carignan & Kalff 1980).
Finally, in a number of studies all the direct effects of nutrient loading affecting aquatic
organisms have proven to result from modifications of the C:N:P stoichiometry balance of
these organisms (Andersen 1997, Elser et al. 2000, Sterner & Hessen 1994, White 1993).
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Figure 6. 1: Relationships between chemical pressures and lake biological communities.
Examples of the effects of artificial eutrophication (1a) and artificial acidification
(1b) (solid line: direct effects, dotted line: indirect effects).
Human-induced acidification (Figure 6. 1b)

In the 20th century, the problem of acidification in lakes has been studied for the most part
in the United States, Canada, and Scandinavia, but it remains an acute problem for the 21st
century (Driscoll et al. 2001, Hultberg & Grennfelt 1986, Moiseenko 2005). The source of
this acidification has been attributed to acid precipitation related to industrial development
and anthropogenic acid-producing substances. Coniferous plantations in lake catchments also
contribute to the decrease in water pH. Two recent reviews have stressed the importance of
this chemical parameter on the entire range of ecosystems’ biological components
(Moiseenko 2005, Solimini et al. 2006).
The effects of acidification have been described as the results of physiological responses
of organisms to increased hydrogen ions (H+) (i.e., decreased pH values), but also to increased
doses of inorganic aluminium (Al3+) and harmful elements (Hg, Cd, Zn, Pb, etc), which are
primarily related to the decreased concentrations of calcium ions (Ca2+) in water. However,
Al3+ has been shown to be the most substantial toxic metal element in acidified water
(Berntssen et al. 1997, Driscoll 1985, Leivestad et al. 1987, Spry & Wiener 1991).
The first direct effect of acidification involves shell-bearing organisms such as molluscs,
mussels, or other crustaceans including crayfish, which are directly sensitive to low pH values
because H+ ions dissolve the calcium carbonates (CaCO3) making up their shells (Moiseenko
2005, Økland & Økland 1986, Singer 1984). Shell-bearing organisms therefore have a limited
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development and may be killed in acidified waters, leading to a decrease in lake community
abundance and richness (Jeffries 1997, Sutcliffe & Carrick 1973).
Others have reported the effects of acidification associated with the decrease in pH values
and the consequences on water chemistry affecting the physiology of these organisms. The
results of research in both acidified freshwaters and laboratory experiments have
demonstrated relations between acidification and (i) disturbances of ionic and osmotic
regulations by blocking cell membrane permeability (Herrmann 1990), (ii) disturbance of the
respiratory function through metal precipitations on the gills (Herrmann et al. 1993, Muniz
1991), and (iii) disturbance of gamete maturation through loss of energy induced by stress
(Davies 1989, Moiseenko 2005). These physiological responses account for species richness
and a decrease in biomass of invertebrate communities; acidification is also an explanatory
factor of the shift in invertebrate composition because it supports predatory species
(Herrmann et al. 1993, Moiseenko 2005).
The physiological effects of acidification have also been studied for fish in laboratory
experiments. Fish exposed to acidified waters increase their standard metabolism in relation
to increased cellular activity for survival only (Edwards & Hjeldnes 1977, Rosseland 1986).
Consequently, energy for fish growth might be lacking (Bonga & Dederen 1986). In addition,
laboratory studies have shown that fish oogenesis and spawning period are sensitive to low
pH conditions (Craig & Baksi 1977, Peterson et al. 1982), which induce lower Ca2+
concentrations and therefore limit production of viable eggs (Beamish 1976). Finally, it has
also been shown that the role of Al ions as a toxic factor increases after hatching by
increasing electrolyte depletion in fish fry (Herrmann et al. 1993). Consequently, changes in
fish species composition and/or abundance can be expected and in most acidified lakes,
acidification has been considered to cause population extinction. The following example is
illustrative: Hesthagen et al. (1999) found that 9630 fish populations (Brown trout, Salmo
trutta, representing 80% of the data set) have been lost in Norwegian lakes during the 50-year
period from 1940 to 1990 due to the effects of acidification.
Similar effects of acidification on lake biota composition and/or abundance have been
reported for plant communities (Driscoll 1985), phytoplankton (Driscoll 1985, Herrmann et
al. 1993, Korneva 1996), and zooplankton populations (Havens & Heath 1989, Moiseenko
2005). The main physiological processes involved are (i) the P assimilation process by aquatic
organisms have been altered by P adsorption or precipitation reactions with Al ions and (ii)
the inorganic carbon assimilation process, which is a key element in the build-up of organic
material and is disturbed by the change in its relative proportions in acidified waters (Driscoll
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1985, Havens & Heath 1989, Solimini et al. 2006). For example, Korneva’s (1996)
experiment in acidic Russian reservoirs demonstrated that with a decrease in pH the number
of phytoplankton species decreased, particularly among centric diatoms, blue-greens, and
Euglenophyta. However, some acidophilic diatom and algae (e.g., Eunotia spp.) increase with
combined low pH and increased N levels (Planas et al. 1989). Differences in sensitivity
between species can explain changes in community composition and structure in acidified
waters.
Metals
In addition to nutrient loading and acidification, many other chemical compounds
resulting from rapid industrialization and urbanization since the Industrial Revolution are also
influencing aquatic communities.
These include metals now found in lake ecosystems resulting mainly from industrial
activities (e.g., mining, landfill leachate, vehicle maintenance, steelworks, refineries)
(Flemming & Trevors 1989, Moraes et al. 2002, Steuer et al. 1997), roads (Pitt et al. 1995),
and parking areas (Davies 1986, Wilber & Hunter 1977) have direct harmful effects on
aquatic communities (Moraes et al. 2002, Phillips & Rainbow 1993). Soluble metals released
into the water column are easily available for plankton, macrophytes, molluscs, and fish
(Chapman et al. 1999, Davies 1986, Demirezen & Aksoy 2004, Lau et al. 1998, Prasad 1997,
Radwan et al. 1990). Even if many trace metals play essential roles in the metabolism and
growth of both plants and animals (Allen et al. 1970), an excessive bioaccumulation of metals
has clearly been demonstrated in several taxa (Chen et al. 2000, Giguère et al. 2005,
Goodyear & McNeill 1999). Materials from lake catchments carried into streams are
accumulated in lake sediments (Ahn et al. 2001, Kim 2005). Benthic macroinvertebrates are
the most sensitive to metal accumulation (Hare 1992, Kiffney & Clements 1993, Kim & Kim
2007), but all plant communities are able to concentrate metals. Hence, benthic macrofauna
and macrophytes have been proposed as pollution-monitoring organisms by many authors
(Greger & Kautsky 1993, Shine et al. 1998, Zhulidov 1996). Finally, even if the
bioaccumulation of these metals decreases as the trophic level increases (Chen et al. 2000),
fish can also accumulate metals in their tissues and be used as bioindicators of metal
accumulation (Mansour & Sidky 2002, Rashed 2001).
This bioaccumulation may also cause physiological changes (e.g., enzyme inhibition) or
have pathological effects (e.g., tissue lesions) (Das et al. 1997, Davies 1986, Kime 1995,
McGeer et al. 2000, Pagenkopf 1983, Weis & Weis 1976). This was documented by a large
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number of studies for many benthic macroinvertebrate species (e.g., amphipod crustaceans,
gastropod molluscs, oligochaetes) (Hodson et al. 1979). In laboratory experiments, Naimo
(1995) demonstrated that sublethal exposure to metals could alter growth, filtration efficiency,
enzyme activity, and behavior in benthic macroinvertebrates. Das et al. (1997) reviewed metal
toxicity in plants. They stated that metals have adverse effects on aquatic plants by changing
root permeability to ions and molecules (e.g., potassium in Thalassiosira) and by changing
the structure and the function of plant membrane, which decreases the concentration of
chlorophyll and oxygen production in many species. It has been shown that decreases in fish
reproduction may occur as well as symptoms of chronic toxicity (e.g., kidney injuries,
hypertension) (Luckey & Venugopal 1977, Moraes et al. 2002, Wren 1986). Laboratory-
based studies have also demonstrated that some zooplankton communities (e.g., Daphnia
magna) were highly sensitive to metals (Biesinger & Christensen 1972, Yan et al. 1996),
especially to nickel (Ni) and copper (Cu) (Ingersoll & Winner 1982, Koivisto et al. 1992),
which reduce organism longevity and impair reproduction.
As a consequence of these physiological effects and considering differential taxa and
individual tolerance, community composition, density, and diversity can be affected by metal
discharges in the water and sediments (De Vevey et al. 1993, Phillips & Rainbow 1993).
Nevertheless, numerous authors studying concentrations of heavy metals in water and bottom
sediments have reported that the real impact on lake biological communities lay in long-term
exposure to metals accumulating in benthic sediments (Garie & McIntosh 1986). These
bioaccumulation processes in the different communities (by direct or indirect effects via the
food web) have not yet been sufficiently studied (Chen et al. 2000, Goodyear & McNeill
1999), particularly in view of understanding and assessing metal pathways in the lower
trophic levels in freshwater ecosystems, because of technical limitations and incomplete
information on the structure of these communities (Chen et al. 2000).
Pharmaceutical pollutants
Among the chemical products released in water bodies, the effects of pharmaceutical
pollutants are increasingly highlighted. Indeed, these pollutants, which contain detectable
quantities of surfactants, antibiotics, and other types of antimicrobial chemicals, have
potential adverse aquatic effects. These direct and indirect effects are usually predicted from
laboratory acute and chronic ecotoxicity data in species such as algae, crustaceans, and fish
(Crane et al. 2006, Versteeg et al. 2005). These pollutants which act mainly on metabolic
processes, can affect homeostasis and physiological functions of aquatic organisms (e.g.,
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musculature atrophy, cardiovascular system pathology, and reproductive development
perturbation) (Ferrari et al. 2003). Modification of the aquatic communities’ composition and
abundance as well as changes in genetic integrity have been observed in individual organisms
(Nash et al. 2004).
Biological effects have been tested in in vivo laboratory experiments on invertebrates and
fish, in particular finding endocrine disruption (i.e., gonads disruptions) due to high sensitivity
to the sex hormones excreted by women either naturally or as a result of oral contraception
(Crane et al. 2006, Zerulla et al. 2002). These pharmaceutical effects on biota can manifest
through reduced fecundity, embryonic development disturbances, decreases in egg
fertilization and sex ratio (skewed toward females), and in demasculinization (lower male
secondary sex characteristic index) (Cheek & McLachlan 1998, Fent et al. 2006, Hutchinson
et al. 1999, Nentwig 2007).
In laboratory experiments, alterations in reproduction and anabolic processes and the
inhibition of bioluminescence in microbial assemblages (e.g., Vibrio harveyi, V. fischeri) after
exposure to antibiotics were highlighted (Al-Ahmad et al. 1999, Backhaus & Grimme 1999,
Kümmerer et al. 2000, Stanislawski 1979). In addition, direct population-level consequences
of exposure to specific pharmaceuticals have also been demonstrated in natural ecosystems.
In the United Kingdom, reduced reproductive success and localized extinctions in marine
gastropods were attributed to the antifouling agent tributyltin (TBT) (Gibbs et al. 1991,
Matthiessen & Gibbs 1998).
Finally, in in situ and laboratory experiments Wilson et al. (2003) demonstrated that
pharmaceutical and personal-care-products (PPCPs) may also induce an adaptive response of
the algal community to prolonged exposure, as is commonly found in bacteria–
pharmaceutical interactions. Indeed, even if further studies are required to precisely
understand the ecological processes, they stated that these chemical compounds on algae
communities can result in final algae biomass yield changes, and more generally can
influence both the structure and the function of algae communities in freshwater ecosystems
receiving such pharmaceutical pollutants.
Others
Urban and suburban lakes are potentially highly impacted by the organic pollutants used
to protect lawns, parks, and home gardens from pests (Overmyer et al. 2005, Phillips &
Rainbow 1993), polycyclic aromatic hydrocarbons (PAH) entering aquatic environments from
many impervious surfaces (e.g., roads, parking, trampled fields) (Stein et al. 2006, Van Metre
- 164 -
et al. 2000), and chlorides resulting primarily from road de-icer applications during the winter
months (Godwin et al. 2003, Kaushal et al. 2005). These chemical pollutants enter lakes
primarily through runoff from rain events and are commonly detected in these aquatic
freshwater ecosystems (United States Geological Survey (USGS) 1999). Moreover, many
authors have shown that several chemical pollutants have been detected at high concentrations
in freshwater and even at concentrations which may be lethal for aquatic life (United States
Geological Survey (USGS) 1999).
These compounds have the same direct impacts on aquatic communities as those induced
by heavy metals. The bioaccumulation of these toxic compounds results in physiological and
pathological problems in aquatic organisms, such as changes in levels of reproductive
hormones, hypertension, tumors, hepatic dysfunction or more widely impairment of
endocrine, immune, and behavioral functions (Beasley & Kneale 2002, Liess & Von Der Ohe
2005, Ostendorp et al. 2004a, Phillips & Rainbow 1993, Rao 1999, Sarkar et al. 2004,
Thornton & Wilhm 1974, Walsh et al. 1994).
With more or less sensitive species, these toxic pollutants may affect the taxonomic
structure (composition and abundance), density, and diversity of aquatic communities, leading
to modification of the entire ecological structure, as heavy metals do (Beasley & Kneale 2002,
Brooker & Edwards 1975, Galat & Robinson 1983, Liess & Von Der Ohe 2005, Robson &
Barrett 1977, Scorgie 1980). However, investigations of the uptake of trace organic
contaminants in freshwater organisms have been restricted largely to fish and
macroinvertebrates (Phillips & Rainbow 1993), and it has been widely recognized that aquatic
macroinvertebrates can be used as biomonitors to assess the effects of these
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Tableau 6. 1: Chemical pressures affecting lake aquatic species and their related effects (direct and indirect effects are presented).
Chemical pressures Effects on lake biological communities References
Nutrient loadings Modifications of C:N:P stoichiometry balance of aquatic organisms Sterner and Hessen 1994; Andersen 1997; Elser et al. 2000
Increase production, growth, and densities of lake biota Breitburg et al. 1999; Lang 1999
Increase frequency and intensity of algae blooms Carpenter et al, 1998a, 1998b, 1998c; Ostendorp et al. 2004a; Smith et al. 2006
Shift in species composition Jeppesen et al. 1998; Dybas 2005; Solimini et al. 2006
Human-induced acidification Dissolution of CaCO3 constitutive of shells Okland and Okland 1986; Moiseenko 2005
Disturbance of ionic and osmotic regulations Herrmann 1990
Disturbance of the respiratory function Muniz 1991; Herrmann et al. 1993
Disturbance of oogenesis and gamete maturation Craig and Baksi 1977; Peterson et al. 1982; Davies 1989; Moiseenko 2005
Decrease in biota composition and/or abundance Driscoll 1985; Korneva 1996; Hesthagen et al. 1999; Moiseenko 2005; Solimini 2006
Metals Physiological effects (e.g. enzyme inhibition, filtration deficiency) Naimo 1995; Das et al. 1997
Pathological effects (e.g. tissue lesions, hypertension) Wren 1986; Moraes et al. 2002
Reduction in organisms longevity and reproduction impairment Biesinger and Christensen 1972; Yan et al. 1996
Decrease in communities’ composition, density, and diversity De Vevey et al. 1993; Phillips and Rainbow 1993
Pharmaceuticals pollutants Alteration of homeostasis of aquatic organisms Ferrari et al. 2003

Endocrine disruption, embryonic development disturbances, decrease
in sex ratio, demasculinization Cheek and McLachlan 1998; Hutchinson et al. 1999; Fent et al. 1996; Nentwig 2007
Modification of other physiological functions (e.g. musculature atrophy) Ferrari et al. 2003
Decrease in communities’ composition and abundance Wilson et al. 2003; Nash et al. 2004
Changes in aquatic communities’ genetic integrity Nash et al. 2004
Other chemical pollutants

Physiological effects (e.g. enzyme inhibition, filtration deficiency) Thornton and Wilhm 1974; Phillips and Rainbow 1993; Walsh et al. 1994; Rao 1999;
(e.g. Organic pollutants, Pathological effects (e.g. tissue lesions, hypertension) Beasley and Kneale 2002; Sarkar et al. 2004; Ostendorp et al. 2004a; Liess and Von Der Ohe 2005
Polycyclic aromatic hydrocarbons, Decrease in communities’ composition, density, and diversity Scorgie 1980; Galat and Robinson 1983; Beasley and Kneale 2002; Liess and Von Der Ohe 2005
Chlorides)
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contaminants on freshwater ecosystems (Overmyer et al. 2005, Robson et al. 2006, Simpson
& Roger 1995).
3. Impacts of morphological pressures on lake biotic communities (Tableau

6. 2)
One of the confusing aspects of water quality management in lakes is that often only the
chemical component of water quality is considered, whereas morphological changes in
ecosystem structure, and the degradation of physical habitats, are current threats to lake
ecological integrity (Elias & Meyer 2003, Engel & Pederson 1998, Treiber 2003). In the last
few decades, urban settlement and recreational activities (e.g., motorboat activities,
navigation) in lakes or in shore areas have dramatically increased (Doyle & Hassett 2004,
Schmieder 2004) and are expected to continue to increase in the future (Brauns et al. 2007a,
Scheuerell & Schindler 2004), probably leading to increased morphological alterations in
lakes. As stated in the European Water Framework Directive (WFD), the morphological
parameters commonly considered are lake shoreline characteristics, the lake substrate, and the
shape of the lake basin (European Community 2000).
Modifications and alterations of the lake shoreline characteristics (Figure 6. 2)

Regarding the lake littoral zones, direct modifications and alterations such as
reinforcement structures (e.g., embankments, sheet pillings, concrete walls), water level
control structures, and artificial areas (e.g., recreational beaches, residential and/or
commercial areas, development of marinas) are frequent. Their impacts on aquatic biota are
broadly similar and have been frequently studied (Elias & Meyer 2003, Radomski & Goeman
2001, Schmieder 2004, Sukopp 1971).
One aquatic organism directly concerned by these direct modifications and alterations of
lake shorelines are macrophytes (Radomski & Goeman 2001, Schmieder 2004), which suffer
from their normal habitat being transformed into large areas of solid substrata (Liddle &
Scorgie 1980). These shoreline modifications break the natural continuity of the littoral
substrate and the moisture gradient (Beauchamp et al. 1994, Jennings et al. 2003), resulting in
changes in the vegetation gradient (Schmieder 2004). Moreover, as reported by Maynard and
Wilcox (1997), these changes or losses of marginal aquatic plants can be the consequences of
the combined effects of reducing littoral zones and changing the wave climate in the near-
shore area. Shore fortifications may shift wave energy to other nearby areas, which can cause
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direct damage to aquatic macrophytes by the direct uprooting actions of boat-generated waves
(Vermaat & De Bruyne 1993). In addition, aquatic macrophytes can also be indirectly
impacted by these recreational activities and power-boat-induced waves because they increase
turbidity, which can inhibit the penetration of solar radiation (Cragg et al. 1980, Murphy et al.
1995). According to Ostendorp (1989) and Strang and Dienst (2004) amphibious reed
vegetation and communities of small-sized plants on nutrient-poor gravel shores are the plant
communities most severely affected by lake shoreline modifications (Elias & Meyer 2003,
Radomski & Goeman 2001, Sukopp 1971).
Macroinvertebrate communities in the lake littoral zone are also significantly reduced
with increasing shoreline modifications (Doyle & Hassett 2004). Increased turbidity in this
modified littoral area may have deleterious effects on aquatic invertebrates (Mosisch &
Arthington 1998). Many species of aquatic invertebrate communities (e.g., Potamopyrgus
antipodarum, Gyraulus albus, Sphaerium spp.) find shelter in the large pore spaces among
cobbles and boulders in lake littoral zones (Bänziger 1995). Thus, when fine sediments fill
these pore spaces, it reduces the quality and quantity of habitat available to
macroinvertebrates (Barton & Hynes 1978a, 1978b, Moore 1980). Furthermore, Liddle (1998)
stated that fine suspensoids in water may obstruct aquatic invertebrate gills by an
accumulation of silt, which results in an increase in invertebrates drift (Murphy et al. 1995).
It has also recently been shown that shoreline development impacts the littoral food-web
by reducing resource availability and consumer species richness due to loss of habitats
(Brauns et al. 2007a, Brauns et al. 2007b) and modifies carbon budget sources in lake
ecosystems (Brauns 2008).
The biological impacts of anthropogenic shoreline development have mainly been
quantified for littoral fish communities, particularly in urban lakes, through the loss of
reproductive or other critical habitats for fish (Bryan & Scarnecchia 1992). Lake shoreline
development, such as bottom substrate modifications (Beauchamp et al. 1994, Jennings et al.
2003) and the removal of coarse woody debris (Christensen et al. 1996b, Marburg et al.
2006), has direct cumulative effects on the physical components of lake littoral habitats,
which result in the loss of fish spawning habitats (NOAA Fisheries 2004). Moreover, these
fine sediments have been shown to obstruct fish gills by an accumulation of silt, which can
clog of respiratory structures, reducing feeding rates and increasing egg kill (Mosisch &
Arthington 1998, Sutherland & Ogle 1975). Consequently, direct modifications and
alterations of lake shorelines may result in a reduction of fish growth rates and have the
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potential to alter fish spatial distributions (Scheuerell & Schindler 2004, Schindler et al.
2000).
In addition, both macroinvertebrate and fish communities are indirectly impacted by lake
shoreline development, through the destruction of aquatic vegetation in littoral areas.
Undoubtedly, the major impact of morphological alterations of the lake littoral zones is an
alteration of native macrophyte community composition and density (Bryan & Scarnecchia
1992), which implies changes to other communities (Engel 1990, Janecek 1988, Radomski &
Goeman 2001, Wetzel 2001), trophic interactions (Jeppesen et al. 1997), and more generally
the entire lake ecosystem function (Christensen et al. 1996b). Aquatic macrophytes provide
lake organisms with food, spawning or nesting habitat, foraging substrates, and cover from
sun and predators (Engel 1990). Fish growth rates and spatial aggregation decrease when
macrophytes disappear, and their spatial distributions are also modified (Scheuerell &
Schindler 2004, Schindler et al. 2000). Furthermore, in the littoral area of lakes, benthic
macroinvertebrate species diversity and abundance decrease when aquatic vegetation is
altered (Bänziger 1995, Brauns et al. 2007a).
Finally, these indirect impacts caused by anthropogenic pressures on macroinvertebrate
and fish communities, which are induced by changes and/or the loss of macrophyte
communities in lakes, occur similarly for morphological and hydrological pressures.
Figure 6. 2: Relationships between morphological pressures and lake biota.
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Example of the consequences of direct shoreline modifications (solid line: direct
effects, dotted line: indirect effects).
Lake substrate modifications
At some point of their life cycle, many aquatic organisms have their principal habitat in,
on, or near sediment. Therefore, the lake’s substrate is a morphological parameter for which
the impacts of modifications on aquatic communities have been well studied at least in the
littoral area (NOAA Fisheries 2004). Numerous authors have recently reviewed the impacts
of suspended and bedded sediments in aquatic ecosystems (Bilotta & Brazier 2008, Naiman &
Bilby 1998, Reid & Dunne 1996, Waters 1995, Wilber & Clarke 2001).
The deleterious effects of high suspended solid loads and sedimentation on lake habitats
have been demonstrated to modify the substrate surface conditions and the volume of fine
sediment within the aphotic zone (Graham 1990, Wood & Armitage 1997). In extreme cases,
fine sediments smother the entire lake substrate by filling interstitial spaces, leading to
modification of the physical, chemical, and biological properties of the water body (Bilotta &
Brazier 2008).
The impacts of these suspended solids settling on substrate have been shown to influence
phytoplankton, periphyton, and macrophytes, primarily through changing the underwater light
climate in aquatic ecosystems (Best et al. 2001, Bilotta & Brazier 2008, Schmieder 2004).
Indeed, the reduction of light penetration through the water column will restrict the rate at
which algae and macrophytes can assimilate energy through photosynthesis (Van
Nieuwenhuyse & LaPerriere 1986). Consequently, turbidity will limit the growth and the
distribution of aquatic plants, which will impact directly on primary consumers (Berry et al.
2003, Bilotta & Brazier 2008, Hecky & Guildford 1984). Moreover, fine sediment suspension
and deposition affect periphyton organisms by reducing the organic content of their cells
(Cline et al. 1982, Graham 1990). In addition, high levels of sediments can also have abrasive
and damaging effects on the photosynthetic structures of organisms (Alabaster & Lloyd 1982,
Steinman & McIntire 1990). In extreme cases, sedimentation causes burial effects on
macrophytes, resulting in changes in species composition and in overall loss of aquatic plants
(Fonseca & Fisher 1986, Terrados et al. 1998). Finally, as in the case of lake shoreline
modifications, sedimentation can also indirectly affect the abundance of aquatic plants by
acting as a vector of nutrients (Heathwaite 1994) and toxic compounds (e.g., pesticides,
herbicides) (Kronvang et al. 2003) from the land surface to the waterbody.
Benthic invertebrates (e.g., mayflies [Ephemeroptera], stoneflies [Plecoptera], caddisflies
[Tricoptera]) are also affected by substrate modifications (Bjornn et al. 1974, Bjornn et al.
- 170 -
1977, Donohue & Irvine 2003, Waters 1995), especially by sediment removal operations
(Benhke & six coauthors 1987). A number of studies have shown that this decrease in
sedimentation increased in invertebrate drift (Gammon 1970, Ryder 1989). In addition, for
filter-feeding invertebrates (e.g., mussels, copepods) (Arruda et al. 1983, Tester & Turner
1989), laboratory studies (Aldridge et al. 1987) have shown that high levels of sediments can
clog feeding and respiratory structures, reducing their feeding and respiratory efficiency,
therefore reducing growth rates, stressing, and even killing these organisms (Hynes 1970).
Another taxonomic group, fish, particularly salmonids, have also received a great deal of
attention from researchers (Alabaster & Lloyd 1982, Berry et al. 2003, Schuett-Hames &
Pleus 1996, Zimmermann & Lapointe 2005). Three major effects of substrate modifications
can affect fish: (i) direct physiological effects of suspended sediment, such as suffocation, (ii)
effects caused by sediment deposition, leading to increased embeddedness or burial of eggs
and larvae, and (iii) effects attributable to decreased in water clarity.
The suffocating effect of the fine sediments may directly affect free-living fish by
clogging fish gill structures (Cordone & Kelley 1961) and reducing the amount of dissolved
oxygen available in water (Bjornn & Reiser 1991, Winfield 2004). Thus, fish population
abundance may decrease catastrophically, and local extinction may occur owing to substrate
modifications (Marschall & Crowder 1996, Rose et al. 1999). The second impact induced by
substrate disturbances involves the deposition and settling of sediments in interstitial spaces
of cobbles and gravel, which are utilized as spawning habitats by fish (Bilotta & Brazier
2008, Scrivener & Brownlee 1989, Wood & Armitage 1997). This has been recognized as a
major cause of the reduced development and survival of salmonid eggs and larvae (NOAA
Fisheries 2004). The deposit of materials blocking the pores in the substrate structure prevents
sufficient exchange of dissolved oxygen and carbon dioxide between breathing eggs and
larvae and flowing water (Greig et al. 2005, Pauley et al. 1989, Walling et al. 2003). Finally,
the decrease in water clarity directly affects the natural movement and swimming of fish in
water (Bisson & Bilby 1982, Bruton 1985, Whitman et al. 1982). Moreover, substrate
modification impacts fish indirectly by reducing the abundance of food available to fish due
to a reduction in primary production and a reduction in habitat available for insectivore prey
items (Bruton 1985, Doeg & Koehn 1994, Gray & Ward 1982). The decrease in water clarity,
which reduces hunting efficiency, particularly in visual feeders, also affects fish populations
and/or communities (Bruton 1985, Ryan 1991).
Hence, all the impacts related to substrate modifications in lakes finally influence lake
biota diversity and abundance (Rowan & Soutar 2005).
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Lake basin shape modifications
Significant alterations in the shape of the lake basin induced by anthropogenic pressures
are currently rare. Nonetheless, in some cases artificial dams are built in order to increase the
natural stocked water volume in certain lakes. As a consequence, the lake’s littoral area is
completely modified and crucial habitats for lake biota can disappear, modifying the lake’s
biological communities in a similar way as those described for changes related to lake
shoreline modifications (see Section 2.1, “Modifications and alterations of lake shoreline
characteristics”) and water level fluctuation (see Section 3.1, “Water level control”).
However, the impacts resulting from lake basin shape modification are not well documented
at this time. One reason is that the status (biological, geomorphological, etc.) of lakes is
seldom assessed before and after dam construction. Moreover, after creating artificial lakes,
many factors covary (e.g., morphological modifications vary with hydrological parameters
such as residence time and vary positively with an increase in services), making clear
pressure–impact relationship analysis difficult.
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Tableau 6. 2: Morphological pressures affecting lake aquatic species and their related effects (direct and indirect effects are presented).
Morphological pressures Effects on lake biological communities References
Modifications and alterations of Damage to aquatic macrophytes by direct uprooting actions Vermaat and De Bruyne 1993
the lake shoreline characteristics Clogging respiratory structures of aquatic organisms Mosisch and Arthington 1998
Reduction in fish growth rates Schindler et al. 2000; Scheuerell and Schindler 2004
Reduction in the quality and quantity of habitat available to aquatic organisms Barton and Hynes 1978a, 1978b; Moore 1980; NOAA Fisheries 2004
Increase in invertebrates drift and alteration of communities’ spatial distribution Murphy et al. 1995; Schindler et al. 2000; Scheuerell and Schindler 2004
Alteration of aquatic communities’ composition and densities Bryan and Scarnecchia 1992; Radomski and Goeman 2001; Wetzel 2001
Lake substrate modifications Decrease in the organic content of aquatic organism cells Cline et al. 1982; Graham 1990
Reduction in organism feeding and respiratory efficiencies Aldridge 1987; Bjornn and Reiser 1991; Winfield 2004
Abrasive and damaging effects to the photosynthetic structures of organisms Alabaster and Lloyd 1982; Steinman and McIntire 1990
Limitation of aquatic plant growth Hecky and Guildford 1984; Berry et al. 2003; Bilotta and Brazier 2008
Reduction in the development and survival of organism eggs and larvae NOAA Fisheries 2004
Changes in fish natural movement and swimming in water Bisson and Bilby 1982; Witman et al. 1982; Bruton 1985
Changes in communities’ spatial distributions Gammon 1970; Ryder 1989; Berry et al. 2003; Bilotta and Brazier 2008
Decrease in communities’ composition, diversity and abundance Fonseca and Fisher 1986; Terrados et al. 1998; Rowan and Soutar 2005
- 173 -
4. Impacts of hydrological pressures on lake biotic communities (Figure 6. 3,
Tableau 6. 3)
During the 20th century, the global population increased fourfold and water withdrawn
from natural freshwater ecosystems increased eightfold (Gleick 1998). Human society has
used freshwater from lakes, groundwater, and wetlands for many different urban (e.g.,
consumption), agricultural (e.g., irrigation), and industrial (e.g., hydroelectric power)
activities (Baron et al. 2002, Jackson et al. 2001b). Moreover, these human water needs are
estimated to continue to increase in the future, and they must also be available throughout the
year (Postel 2000, Postel & Peterson 1996). Consequently, in instigating hydrological
changes, human society produces negative consequences or trade-offs in freshwater
ecosystems, especially in terms of supporting aquatic life and maintaining suitable water
quality (Baron et al. 2002, Naiman et al. 1995, Postel 2000, Stein et al. 2000).
According to the WFD, hydrological parameters relevant to supporting ecosystem
functioning are flow quantity and dynamics, water level, residence time, and the resultant
connection to groundwater (European Community 2000). Most of these parameters are
controlled in artificial water bodies as well as in some natural lakes through hydraulic
structures created to meet human needs (Bergkamp et al. 2000).
Figure 6. 3: Relationships between hydrological pressures and lake biological communities

(solid line: direct effects, dotted line: indirect effects).
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Water level control
Among the hydrological pressures, water level control was probably the most heavily
emphasized in the grey literature, scientific papers, and reviews (Bragg et al. 2003, James et
al. 2002, Maynard & Wilcox 1997, Van Groenendael et al. 1996). The loss of natural
submersed and emerged vegetation in littoral zones resulting from physically removing
macrophyte habitats has been shown for different environmental situations (Fuller 2002,
Schmieder 2004, Wilcox 1995). This loss of aquatic vegetation reduces lake’s littoral prey
populations including epiphyton and macroinvertebrates (Heikinheimo-Schmid 1985,
Hofmann 1998, Smith et al. 1987). It is also detrimental to fish feeding conditions, sheltering,
and reproduction. Therefore, via its impacts on macrophytes, water level regulation may
impact other lake biological communities, modifying the entire food-web structure. For
example, Brauns et al. (2008) stated that loss of eulittoral root habitats in lakes will cause a
significant alteration of the littoral invertebrate community. These modifications of
invertebrates’ biomass and diversity may impact invertivorous fish communities.
Nevertheless, fluctuations in water level may severely and directly act on reproduction of
fish, threaten egg incubation and even their deposition in suitable littoral fish spawning
habitat whatever the species (Michaletz 1997, Winfield 2004, Winfield et al. 1998).
Moreover, water level variation instigates an increase in lake turbidity, in concentrations
of the end products of organic decomposition, and a decrease in oxygen concentrations in
lakes (Hambright et al. 1997). Water fluctuation may result in the lack of mixing and sunlight
for photosynthesis. Anoxic conditions, especially in the bottom layer of lake water, are also
noted (Bergkamp et al. 2000). Thus, water level impacts the composition and the productivity
of the phytoplankton communities (Hofmann 1998).
Residence time
Ambrosetti et al. (2003) defined residence time as “what controls the concentrations and
accumulative capacity of all substances entering a lake basin, whether naturally or
artificially.” Since the classical empirical model of lake eutrophication (Vollenweider 1976)
recognized that the biogeochemical processing of phosphorus varies with residence time,
other authors have demonstrated that variable water retention and flushing influence thermal
stratification (Hamilton & Lewis Jr 1987), isotopic composition (Herczeg & Imboden 1988),
dissolved organic carbon concentration (Christensen et al. 1996a), heavy metal concentration
(Hilton et al. 1995), but also nutrient concentrations (Dettmann 2001, Hecky et al. 1993,
Kaste et al. 2003), mineralization rates of organic matter (Heyer & Kalff 1998), and finally
- 175 -
primary production (Jassby et al. 1990). High water retention time has been shown to
negatively affect plankton abundance and composition (Basu & Pick 1996, Bledzki & Ellison
2001, Gophen 2003). Moreover, due to the significant role played by plankton in nutrient
cycling in lakes, modification of the residence time may also indirectly impact other aquatic
communities. Indirect effects of residence time modifications on juveniles in fish
communities, by the loss of food resources, was described by Petesse et al. (2007b), but to our
knowledge, nothing is clearly shown on the other biological communities.
Natural hydrological cycle

The modification of the natural hydrological cycle, especially the alteration of flow
regimes, influences the thermal structure of lakes (Carmack et al. 1986, Killworth & Carmack
1979) and also controls the spatiotemporal distribution of nutrients, sediments, and other
water quality parameters (Gloss et al. 1980, Pharo & Carmack 1979, Smith 1978, Soltero et
al. 1974). Several numerical models developed since the 1970s showed that hydrological
cycle variation indirectly impacts phytoplankton structure and abundance by modifying
nutrient input rates in lakes, especially during the phytoplankton growing season (Carmack &
Gray 1982, Carmack et al. 1986, Gophen 2004).
The alteration of flow regimes may also have direct impacts on fish communities,
especially by modifying natural paths for migratory species (e.g. Salmo trutta L. and
Oncorhynchus mykiss) (Lake 2003). The overall rate of throughflow of a lake may be
modified, because the flow rates of rivers which feed this lake can be controlled. However,
attempts to analyze biological responses to modifications of natural flows in lakes remain
extremely scarce (Gloss et al. 1980, Killworth & Carmack 1979, Lake 2003). Moreover, most
of these analyses have been studied in morphologically specific lakes (i.e., fairly narrow
lakes, very extended lakes, etc.) (Killworth & Carmack 1979), where dynamical synergistic
effects (i.e., rotation, flood events) may interact with flow regime alteration impacts
(Ambrosetti et al. 2003, Gloss et al. 1980, Killworth & Carmack 1979).
Groundwater exchange
Changes related to groundwater have been investigated for two decades using
groundwater flow modeling (Cheng & Anderson 1994, Hunt et al. 1998, Hunt et al. 2003b,
Hunt et al. 2003a) and geochemical/isotope tracing of water (Walker et al. 2003, Walker &
Krabbenhoft 1998). These studies have shown that the variety of water sources (terrestrial and
lake recharge) can lead to complex flow systems with a variety of provenances occurring
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where groundwater discharges into lakes (Hagerthey & Kerfoot 1998, Hurley et al. 1985,
Lodge et al. 1989, Wentz & Rose 1989).
Lowering groundwater recharge directly affects the properties of surface waters by
reducing the water level of groundwater-fed lakes, impacting stable-temperature habitats and
modifying nutrient and inorganic ion supplies (Hagerthey & Kerfoot 1998, Hayashi &
Rosenberry 2002, Hunt et al. 2003b), consequently impacting aquatic habitat structure, food
resources and biotic interactions (Finkenbine et al. 2000). This has been demonstrated most
particularly for rooted macrophyte communities. Several studies have shown that the biomass
and the diversity of littoral epibenthic algal communities is influenced by modifications in
groundwater exchanges, because alterations of water flows across the sediment–water
interface can create and maintain a physicochemical spatial heterogeneity both within and
between lakes (Hagerthey & Kerfoot 1998, Lillie & Barko 1990, Lodge et al. 1989, Loeb &
Hackley 1988).
Modifications of groundwater exchange also directly affect slope stability in lake littoral
zones and erosion processes, impacting the strength of bank materials and sediment porosity
(Hunt et al. 2006). These changes indirectly affect a lake’s resident organisms, especially
benthic invertebrate and fish abundance and taxonomic richness, by modifying their spawning
habitats (Finkenbine et al. 2000, Hunt et al. 2006, Malard et al. 1996). Furthermore, lake–
groundwater exchange modifications may also influence the spawning behavior of some fish
(Hayashi & Rosenberry 2002): it has been shown that areas of groundwater discharges have a
stable temperature regime and thus provide thermal refuge for fish both in winter and summer
(Curry & Noakes 1995).
Finally, in some cases, lake riparian vegetation can also be indirectly impacted by changes
in groundwater relations, because these modifications may reduce the bank moisture of lake
shorelines (Hunt et al. 2006).
However, even if the modification of lake–groundwater exchanges has been shown to
affect entire freshwater ecosystems (Hayashi & Rosenberry 2002, Klein 1979, Pawlow &
Scientist 1977), the relations between microhabitat quality and groundwater flow have not
been well documented for benthic organisms to date (Hunt et al. 2006). One reason is that it is
necessary to better understand the effects of topography, geology, and climate on groundwater
flow systems to understand the interactions between groundwater and surface water (Winter
1999). Moreover, future studies should also be undertaken to understand the effects of
complex biogeochemical processes within lake bottom sediments, which may affect lake
aquatic communities (Hayashi & Rosenberry 2002).
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Tableau 6. 3: Hydrological pressures affecting lake aquatic species and their related effects (direct and indirect effects are presented).
Hydrological pressures Effects on lake biological communities References
Water level fluctuation Decrease in fish reproduction and fish eggs incubation Michaletz 1997; Winfield et al. 1998; Winfield 2004
Physical removing of macrophyte habitats in lake littoral zone Wilcox 1995; Fuller 2002; Schmieder 2004
Changes in communities’ composition and productivity Hofmann 1998; Brauns et al. 2008
Residence time changes Loss of food resources Petesse et al. 2007

Decrease in plankton abundance and composition Basu and Pick 1996; Bledzki and Ellison 2001; Gophen 2003
Natural hydrological cycle modification Modification of natural paths for migratory species Lake 2003
Changes in phytoplankton structure and abundance Carmack and Gray 1982; Carmack et al. 1986; Gophen 2004
Groundwater exchange modifications Influence the spawning behavior of some fish Hayashi and Rosenberry 2002
Changes in lake riparian vegetation and habitat structure Finkenbine et al. 2000; Hunt et al. 2006
Changes in food resources and biotic interactions Finkenbine et al. 2000
Lodge et al. 1989; Lillie and Barko 1990; Malard et al. 1996; Hagerthey and Kerfoot 1998; Hunt
Changes in communities’ biomass and diversity et al. 2006
- 178 -
5. Impacts of species manipulation on lake biotic communities (Figure 6. 4,
Tableau 6. 4)
The present review will now focus on biological manipulations, including intentional and
unintentional introduction of new species into aquatic systems, which is sometimes
considered one of the greatest environmental threats facing aquatic communities (Lowe-
McConnell 1990). This has been assessed in lacustrine aquatic communities for many years
by many authors, who have documented the type and management of manipulations mainly
regarding fish (e.g., Beveridge 1984; Gerdeaux 2004; Winfield 2004; Allan et al. 2005).
These manipulations largely result from historical commercial activities, particularly in lake
fisheries exploiting fish populations in most parts of the world (Winfield 2004). Nevertheless,
direct modifications of communities are not exclusively limited to fish. Scientists and
managers are also beginning to pay attention to introduction of other organisms in today’s
growing concern for biodiversity.
Figure 6. 4: Relationships between species manipulations and lake biota, especially on native
fish communities (solid line: direct effects, dotted line: indirect effects).
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Introduction of individuals or species
The impact of fish introduction has been well documented and several impacts have been
reported (Allendorf 1991, Holcik 1991). First, the introduction of new individuals of native
species or new species sparks changes in the trophic equilibrium of the community or
ecosystem. By modifying competition and/or predation relationships, these actions can
modify ecosystem functioning and sometimes species richness and biodiversity on the lake
scale (Englbrecht et al. 2002, Ostendorp et al. 2004a). Introducing individuals and species in
lakes has been demonstrated to have direct effects on the entire food web by varying fish
populations and the resulting effects on the lower trophic levels in lakes. For example, in fish
manipulations in Tuesday Lake, Carpenter et al. (1985) and Carpenter et al. (1987) showed
that the addition of piscivores and the reduction of planktivores caused an increase in
zooplankton biomass, a compositional shift from a copepod/rotifer assemblage to a
cladoceran assemblage, a reduction in algal biomass, and a continuous reduction in primary
productivity. Numerous additional studies, such as Carpenter and Kitchell (1988), Carpenter
and Kitchell (1992), Stein et al. (1995), and Pace et al. (1999), have confirmed that fisheries
and fish management are altering trophic cascades by modifying fish community structure
and diversity, with profound consequences for food webs in lake ecosystems.
Modifications of ecosystem functioning often remain very difficult to prove because of
the lack of complete data series before and after species introduction (Englbrecht et al. 2002).
Nevertheless, local extinctions of native species were frequently related to the introduction of
new species (Crivelli 1995, Holcik 1991, Olenin et al. 2007, Welcomme 1974, Winfield
2004). For example, in the 1980s an explosive increase of the Nile perch (Lates sp.; Harrison
1991) that had been introduced in Lake Victoria caused the destruction of approximately 65%
(approximately 200 species) of the endemic haplochromine cichlids (Goldschmidt et al. 1993,
Kaufman 1992, Ogutu-Ohwayo 1993, Witte et al. 1992a, Witte et al. 1992b). Moreover, a
vast literature on the introduction of new species showed that introducing alien species in
lakes has been implicated in the decline of aquatic communities other than fish communities,
such as amphibians (Kiesecker & Blaustein 1997, Lawler et al. 1999), invertebrates (Gamradt
& Kats 1996, Lodge et al. 2000, Mackie 1991, Phelps 1994, Pinkster et al. 1977, Williamson
1996), and macrophyte communities (Beltrami et al. 2008, Les & Mehrhoff 1999, Mazzeo et
al. 2003, Simpson 1984).
Another consequence of introducing new individuals of native species and/or new species
is hybridization. The homogenization of the genetic pool is considered a threat for
biodiversity, increasing the risk of extinction in case of environmental damage (Allendorf
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1991, Englbrecht et al. 2002, Holcik 1991). For example, in Scandinavia, brown trout (Salmo
trutta L.) has been widely stocked in natural ecosystems. These massive releases of hatchery-
produced brown trout have reduced the genetic diversity among wild populations (Ryman
1981), and recent studies suggest interbreeding between hatchery and wild fish may be
common in brown trout (Berrebi et al. 2000, Garcia-Marin et al. 1999, Hansen et al. 2000).
Interbreeding may lead to domestication, and if massive numbers of hatchery fish are stocked
these may even replace ancestral wild populations (Englbrecht et al. 2002, Hansen &
Loeschcke 1994).
A frequent indirect consequence of introducing individuals or species is the concomitant
introduction of diseases (Lambert 1997, Weir & Grant 2005): bacteria, protozoa, and virus
pollution. For example, the nematode Anguillicosa crassus introduced with Anguilla japonica
in Europe was responsible for large-scale mortality of the European eel Anguilla anguilla
(Hickley & Chare 2004).
Finally, introducing fish species also modifies lake habitats. For example, European carp
species (e.g., Cyprinus carpio), known to have been taken and used at Bassenthwaite Lake
and Windermere (United Kingdom) as live bait, have significant consequences for sediment–
water nutrient flows and water turbidity due to their activities deteriorating shallow areas
(Winfield & Durie 2004).
The impact of introducing other organisms has generally not been thoroughly studied
(Horne et al. 1992, Rodríguez et al. 2003). Whenever they are discussed they are related to
macrophyte introduction (Keast 1984, Ozimek et al. 1990). These impacts concern mainly
habitat modification, particularly with exotic macrophyte proliferation. For example, Ozimek
et al. (1990) demonstrated in Lake Zwemlust (Netherlands) that the abundance of exotic
species (e.g., Clara globularis, Elodea sp.) contributed to transient N-limitation of
phytoplankton causing a persistent clear-water phase in lakes.
Microbial pollution
Microbial pollution, especially pathogenic bacteria pollution, is generated by point
sources of pollution (i.e., outfalls of municipal wastewater treatment plants, outfalls of
industrial and hospital sewage) and non-point pollution sources (i.e., broken sewer lines,
sanitary sewer overflows, agricultural runoff, and waste holding-tank discharges) (Gunnison
1999, Toroglu & Toroglu 2009). Moreover, different types of anthropogenic activity, such as
the aquarium industry, aquaculture, the bait industry, and ship hull fouling are additional
sources of microbial pollution in aquatic ecosystems (Cariton & Geller 1993, Ruiz et al. 1997,
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Vignon & Sasal 2009). This microbial pollution can potentially result in pathology and
physiological changes in aquatic communities (Barber et al. 2000, Gozlan et al. 2006). Thus,
according to the comparable nature of these impacts on lake biota, this microbial pressure is
discussed here, in the same section as the species manipulation pressures.
Microbial pollution, including bacteria, protozoans, and viruses may have both direct and
indirect effects on a large variety of aquatic species (Barber et al. 2000, Jackson & Füreder
2006, Kagami & Urabe 2002). However, although the potential importance of these
pathogens has been known for over a decade, studies of pathogens have focused nearly
exclusively on fish and shellfish populations (Brookes et al. 2004, Jackson & Füreder 2006),
probably in response to human health concerns. Even if the presence of fish and shellfish
pathogens throughout Europe is frequently reported, relatively innocuous effects are generally
registered (Gozlan et al. 2006). When serious parasites were identified, they had immediate
and pronounced negative effects on hosts (Bartholomew & Reno 2002, Gozlan et al. 2006,
Johnsen & Jensen 1991). Barber et al. (2000) reported that diseases associated with these
microbial pathogens could either act directly through mortality or indirectly (e.g., reducing
fecundity, increasing risk of predation, reducing swimming speeds, and altering host
behavior). Indeed, since parasites generally utilize host energy resources, these deleterious
effects are frequently nutritional. Furthermore, parasites can also have additional impacts – as
they invade, move around, or grow within the host – through associated pathology and
physiological imbalance. Restricting the review to the effects of parasites on fish hosts, some
physiological effects to mention are muscle atrophy, nervous system pathology, respiratory
distress, and reproductive development infections (Hoffman 1999, Read 1990, Sweeting
1977). In addition, organisms such as mussels and clams can further concentrate pathogens by
their filtering action (Stewart et al. 2008), and thus may be infected or killed.
Consequently, microbial pathogens play essential roles as interacting components of a
lake’s natural environment, in regulating host population abundance and dynamics and thus
influencing the entire lake’s community structure (Malan et al. 1997, Melanie et al. 2006,
Zöllner et al. 2003). The introduction of microbial pathogens (Myxobolus cerebralis and
salmonids, Koi herpes virus and Cyprinus carpio, etc.) alters lake ecosystem processes and is
responsible for the loss of aquatic biodiversity, thereby causing major economic losses
(Gozlan et al. 2006, Vignon & Sasal 2009).
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Overexploitation of aquatic resources
Overexploitation of aquatic resources, especially fishery resources, has been changing
lake ecosystems for decades. Several studies presented intensive commercial fishing on lakes
as the primary disturbance over time (Hartman 1973, Leach & Nepszy 1976). Nevertheless,
given that the current fishery crisis has focused nearly exclusively on marine resources, the
effects of overfishing in freshwater ecosystems have for the most part gone unnoticed within
the global analysis and therefore have been poorly documented (Hilborn et al. 2004, Kura et
al. 2004).
The first impact of these activities is “the removal of apex predators, with the potential for
substantial top-down effects on food web dynamics” (Allan et al. 2005). The impact of
fisheries on the food web were demonstrated through the successive removal of larger
elements of a multispecies fish assemblage (e.g., Lates niloticus, Heterotis niloticus, and
Distichodus in Lake Victoria) and their replacement by smaller elements (e.g., Labeo,
Clarias, Heterobranchus, Schilbe, and Synodontis), which typically are at lower trophic levels
(Pauly et al. 1998, Regier & Loftus 1972). This impact, known as “fishing down the food
web” (Allan et al. 2005), induces the loss of apex predators and results in relaxation of top-
down control of prey populations and stronger top-down control at the next trophic level
below. However, overfishing may not concern only the top predators: it has also been found
to control dominant herbivores (e.g., small cyprinids), thereby influencing primary production
and abundance of basal resources via indirect pathways in trophic cascades and bottom-up
controls (Bain 1993, Flecker 1996). Moreover, Crivelli et al. (1997) found that overfishing
has obviously affected long-lived species with delayed sexual maturity more than short-lived
species. As a result, overexploitation has led to declines in spawning runs of fish species (e.g.,
salmonids) through a decrease in catch size, size at maturation and decreasing stocks (Crivelli
1984, 1990).
Thus, overfishing reduces both the mean size of individuals (i.e., the size structure of the
population) and fish species abundance in the assemblage. In some extreme cases, overfishing
can lead to the removal of some fish species. For example, Smith (1968) discussed the fish
assemblage of the Laurentian Great Lakes. In these lakes, major changes after the late 1800s
and continuing into the first half of the 20th century reflect an intensive and selective fishery
targeting a succession of species, such as lake sturgeon (Acipenser fulvescens), cisco
(Coregonus artedi), lake whitefish (Coregonus clupeaformis), and sea lamprey (Petromyzon
marinus), and resulting in their successive collapses.
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Therefore, the reduction in fish assemblage diversity signifies the lower number of fish
species involved in the critical functions of lakes. Since overfishing is a contributing factor to
the decline of freshwater biodiversity, it may contribute, for example, to the removal of
herbivorous fish (e.g. Cyprinus spp.), which are particularly important in primary producer
regulation through direct grazing on benthic algae and macrophytes (Bain 1993, Flecker
1996) and in the changes in piscivorous fish density (Allan et al. 2005).
Finally, inland capture fisheries are declining in North America, Europe and the former
Soviet Union, and many inland commercial fisheries have been abandoned and replaced by
recreational fisheries (Allan et al. 2005, Cooke & Cowx 2004). The FAO (1999) estimates
global recreational harvest may be on the order of 2 million tons. Moreover, Cooke and Cowx
(2004) and Muoneke and Childress (1994) have stated that these recreational fisheries have
adverse effects on aquatic ecosystems, because even if fish are released recreational fishing
can result in substantial post-release mortality, reducing fish growth and fitness as well.
However, the global recreational harvest remains poorly documented today, because of
inadequate records and lack of a historical framework (Allan et al. 2005).
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Tableau 6. 4: Species manipulation pressures affecting lake aquatic species and their related effects (direct and indirect effects are presented).
Species manipulation pressures Effects on lake biological communities References
Introduction of individuals or species Modification of competition and/or predation relationships Englbrecht et al. 2002; Ostendorp et al. 2004a
Changes in communities’ structure and biodiversity, and alteration of
trophic cascades Carpenter and Kitchell 1988, 1992; Stein et al. 1995; Pace et al. 1999
Local extinctions of native species Welcomme 1974; Holcik 1991; Crivelli 1995; Winfield 2004; Olenin et al. 2007
Homogenization of the communities’ genetic pool (hybridization) Allendorf 1991; Holcik 1991; Englbrecht et al. 2002
Microbial pollution Physiological effects (e.g. enzyme inhibition, filtration deficiency)

Sweeting 1977; Read 1990; Hoffman 1999; Barber et al. 2000; Gozlan et al. 2006
Pathological effects (e.g. tissue lesions, hypertension)
Host population abundance and dynamics regulation Malan et al. 1997; Melanie et al. 2006
Loss of aquatic biodiversity Gozlan et al. 2006; Vignon and Sasal 2009
Loss of apex predators and relaxation of the top-down control of prey

Overexploitation of aquatic resources populations Regier and Loftus 1972; Pauly et al. 1998; Allan et al. 2005
Stronger top-down control at the next trophic level below
Declines in spawning runs of fish species Crivelli et al. 1997
Decrease in catch size and size at maturation Crivelli 1984, 1990; Muoneke and Childress 1994; Cooke and Cowx 2004
Changes in fish species abundance, density, and decreasing stocks Smith 1968; Allan et al. 2005
Declines in freshwater biodiversity Bain 1993; Flecker 1996
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6. Conclusion and perspectives for biological monitoring
The worldwide published literature regarding anthropogenic pressures affecting lakes and
reservoirs and their impacts on biological communities is abundant. Nonetheless, existing
reviews on this subject are rare and partial (Allan et al. 2005, Bragg et al. 2003, Liddle &
Scorgie 1980, Solimini et al. 2006). This literature review has provided an overview of the
expected relationships between the four major types of pressures occurring on freshwater
ecosystems and the different biological communities: (i) chemical pressures, (ii)
morphological pressures, (iii) hydrological pressures, and (iv) species manipulations in an
attempt to respond to the increasing need for this type of information in discussing
environmental management of lake ecosystems.
Most of the information on pressures and impacts on lake biota relationships is scattered
over the journals, grey literature, and technical reports from many different fields; this review
therefore cannot claim to be a perfectly exhaustive census of anthropogenic pressure–impact
relationships.
This review highlights the problems assessing general patterns in the responses of biotic
communities to the different pressures considered. First, many papers are case studies,
particularly those investigating the morphological and hydrological effects on lake biota,
which focus on specific pressures (e.g., impoundment, shoreline reinforcement) and/or
specific biological communities (Hecky & Guildford 1984, Marschall & Crowder 1996).
Second, many relationships are experimentally demonstrated in laboratories without being
observed in the natural environment, such as the impacts of chemicals on biota (Backhaus &
Grimme 1999, Craig & Baksi 1977, Koivisto et al. 1992, Nash et al. 2004, Wilson et al. 2003,
Zerulla et al. 2002). Hence, it is difficult to draw reliable general conclusions regarding the
impacts of anthropogenic pressures on lake biota.
In addition, the effects related to the different types of pressure have not been uniformly
identified and documented. The knowledge of eutrophication, acidification, and the
hydrological effects on the various lake communities appears to be highly developed (Bragg
et al. 2003, Carpenter et al. 1998a, Carpenter et al. 1998b, Carpenter et al. 1998c, Driscoll et
al. 2001, James et al. 2002, Moiseenko 2005, Solimini et al. 2006, Vitousek et al. 1997).
Conversely, Schmieder (2004) found only a few papers on the effects of shore
reinforcements, structural burdens, tourism, and nature protection on lake biota. Moreover,
information on the effects of biotic introduction on lake biota is even more limited. It
currently focuses exclusively on fish communities in lakes (Lowe-McConnell 1990, Winfield
2004). Consequently, it is clear that more research is required on several anthropogenic
pressures in order to understand their impacts on lake biota. As suggested by Carpenter and
Cottingham (1997), a conceptual framework for understanding the interactions between
humans and lakes should be drawn up so as to expand the knowledge on the understudied
impacts.
The hierarchical organization of the effects on different types of pressures on the
biological features of lakes is another consideration. Because of the “trophic cascade”
relationships, any modification of phytoplankton communities, the first layer in this trophic
cascade, will necessarily impact other biological communities in lakes. Nevertheless, the
extent of these trophic relationships has not been fully assessed. Moreover, anthropogenic
pressures are generally combined (rather than occurring in isolation) so that in most studies it
is difficult to disentangle specific pressure–impact relationships. For example, many authors
have highlighted the relations between the physicochemical parameters and the
hydromorphological components (Fillos & Swanson 1975, Novotny et al. 2005).
Finally, reviewing the impacts of anthropogenic pressures on lake biota has highlighted
the heterogeneity of the observation methods used, especially for chemical pressures.
Consequently, comparing and summarizing the collected information posed a number of
problems. This point also argues in favour of Carpenter and Cottingham’s (1997) wish to
create a conceptual framework in order to homogenize the existing knowledge on the
interactions between anthropogenic pressures and lake biota. Water managers, stakeholders,
and aquatic scientists working to preserve lake biodiversity and biological integrity require
common knowledge for management decisions.
Furthermore, this literature review has enabled us to identify fields in need of research and
draw attention to a certain number of future perspectives.
Today there are no in situ studies taking into account possible synergistic or additive
effects stemming from exposure to multiple chemical pollutants (e.g., heavy metals, organic
pollutants, pharmaceuticals) when they are mixed at various concentrations in aquatic
ecosystems. Moreover, the toxicities of many of the degradation products of these chemical
pollutants are as yet unknown (Overmyer et al. 2005). In addition, as stated by the United
States Environmental Protection Agency (USEPA), toxicological synergies are a public health
and water management concern because chemicals individually considered safe might pose
unacceptable health or ecological risks when combined. This lack of information also applies
to many other types of anthropogenic pressure combination to which lake biota are exposed,
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including hydrological, morphological, and biological pressure synergies (Allan et al. 2005,
Radomski & Goeman 2001). In addition, cumulative effects, such as the same pressure
applied later in time to an already stressed system can have a much larger effect but remain
largely unstudied (Doyle & Hassett 2004). Nevertheless, the answer to these questions
requires long time series of in situ observations in combination with laboratory experiments
on various types of pressures exerted on different lake communities. We can expect that the
European observation networks under development (e.g., Eionet (see
http://www.eionet.europa.eu/about), SEIS (see
http://ec.europa.eu/environment/seis/index.htm)) will improve this kind of knowledge.
Moreover, this review suggests that an adequate approach to assess the effects of the
different types of pressure on ecosystem functioning requires a precise understanding of the
processes impacted by these pressures, to measure the responses to multiple lake taxa and to
monitor the effects of anthropogenic pressures on several spatial and temporal scales. One
illustrative example is the pathways of metal movement in the lower trophic levels in
freshwater systems, which remain particularly poorly understood, as does the dominant
source (food vs. water) of metals to biota (Chen et al. 2000). Similarly, recent research on
microbial pollution has shown that detecting particular pathogens in aquatic ecosystems was
extremely difficult, because of the lack of knowledge on the possible hosts across various
taxonomic groups (Gozlan et al. 2005). Moreover, research on several anthropogenic
pressures seems to have been restricted largely to certain lake communities, such as the
studies on the uptake of trace organic contaminants focusing mainly on fish and
macroinvertebrates (Phillips & Rainbow 1993), lake shoreline modification studies, focusing
largely on littoral fish (Brauns et al. 2007a, Scheuerell & Schindler 2004), and water level
modification studies, exploring plankton community responses (Bledzki & Ellison 2001).
The lack of quantitative measurement of the effects of pressures must also be emphasized.
Solimini et al. (2006) reported that there are no data or only limited data available for some of
the relationships between lake biota responses and certain eutrophication parameters. In
particular, data on phytobenthos, benthic invertebrates, and oxygen conditions in lakes are
very limited, as are data on macrophytes and fish abundance. In addition, Bartholomew and
Reno (2002) added that due to the lack of, and inconsistencies in, survey data over the present
distribution of Myxobolus cerebralis, relationships between lake biota responses and some
parasite impacts remain poorly understood. Fent et al. (2006) discussed the general lack of
chronic toxicity data on pharmaceuticals, which highlights the limited assessment of the
chronic effects of pharmaceuticals on fish communities. Finally, Englbrecht et al. (2002) have
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also shown that the records of artificial stocking in natural lake ecosystems were often
incomplete or uncertain.
Even when anthropogenic pressures are qualitatively and quantitatively assessed, their
potential long-term effects are not investigated. Ecotoxicological long-term effects, especially
with respect to potential disturbances in hormonal homeostasis, immunological status, or gene
activation and silencing during long-term exposure, remain unexplored (Fent et al. 2006). The
same problem occurs for anthropogenic hydromorphological pressures (e.g., lake shoreline
modifications, lake substrate modifications, water level regulation), whose direct effects are
not always visible and whose long-term consequences on lake ecosystem structure and
function have never been evaluated (Bragg et al. 2003).
The strategy for monitoring and characterizing anthropogenic pressures and their effects
on lake biota is built upon measurement strategies that have evolved greatly over the past 25
years to produce new tools and instruments (European Commission 2002). This included
measurements of new important parameters, definition of standards, new technologies,
improvements in procedures, and introduction of new regulations (Solimini et al. 2006). Thus,
data acquisition dynamics should continue to improve in the future, and our knowledge of
pressures–lake biota relationships should also continue to increase. Furthermore, encouraging
ecologists, scientists, and water managers to share their knowledge more effectively to protect
and restore water resources should also result in a more integrated approach to the
development of new management strategies.
Finally, this literature review focuses on pressures related to local and regional
anthropogenic activities; however we have also kept in mind the existing global pressures,
including climate change, probably the most worrying today. Improving the understanding of
the relationships between large-scale pressures and biotic communities, and the interactions
between global and smaller-scale pressures are also daunting challenges in the coming years.
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Annexe 2 : Liste des 138 plans d’eau français pour lesquels des données environnementales naturelles et des pressions anthropiques sont
disponibles.
Plans d’eau analysés dans l’étude présentée au chapitre 3. Ce tableau donne le nom du plan d’eau (Nom_Lac), le groupe typologique auquel le
plan d’eau appartient (Groupe Lac ; LNX : lac naturel du type X et RAY : réservoir du type Y) et le numéro du plan d’eau sur les figures
(Numéro Lac).
(a) Ce tableau donne également les caractéristiques environnementales naturelles disponibles pour les 138 plans d’eau listés [Surface BV :
superficie du bassin versant, en km2 ; Indice Géologie : indice géologique traduisant le pourcentage du bassin versant constitué par des roches
calcaires (maximum = 1), sans unité ; Superficie : superficie du plan d’eau, en km2 ; Profondeur Max : profondeur maximale du plan d’eau, en
m ; SLDF : indice de développement des berges, sans unité ; Indice Cuvette : indice traduisant la forme de la cuvette lacustre, sans unité ; Jan +
Juil : somme des températures moyennes des mois de juillet et janvier, en °C ; Juil - Jan : différence entre les températures moyennes des mois
de juillet et janvier, en °C ; Altitude : altitude du plan d’eau, en m ; Temps Séjour : temps de résidence associé au plan d’eau, en jours ; Marnage
Hauteur : hauteur du marnage associé au plan d’eau ; 3 classes : 0 (aucune variation), 1 (≤3 m), 2 (>3 m)] . Les 44 plans d’eau, pour lesquels des
données poissons sont également disponibles, apparaissent en gras dans la liste.
(b) Ce tableau donne également les variables de pressions anthropiques disponibles pour les 138 plans d’eau listés [DensitePopBV : la densité de
population sur le bassin versant, en habitants / km2 ; DensitePop Buff1000 : la densité de population dans une zone de 1000 m autour du plan
d’eau, en habitants / km2 ; Densite Reso : la densité de réseaux de communication sur le bassin versant, en m / km2 ; % du BV Urbain : le
pourcentage d’occupation du sol de type urbain à l’échelle du bassin versant ; % du BV Agricole : le pourcentage d’occupation du sol de type
agricole à l’échelle du bassin versant ; Bilan Azote BV : le bilan en azote à l’échelle du bassin versant, en tonnes ; Bilan Phosphore BV : le bilan
en phosphore à l’échelle du bassin versant, en tonnes ; Surplus Azote BV : le surplus en azote à l’échelle du bassin versant prenant ainsi en
considération les dépôts atmosphériques d’azote, en tonnes ; Bilan Phosphore * Erosion : les teneurs en phosphore entraînées par érosion sur le
bassin versant, sans unité ; LHMS score : le Lake Habitat Modification Score issus de l’application du protocole Lake Habitat Survey, sans
unité]. Les 44 plans d’eau, pour lesquels des données poissons sont également disponibles, apparaissent en gras dans la liste.
(a)
Indice Indice Temps Marnage
Groupe Numéro Surface BV Superficie Profondeur SLDF Jan+Juil Juil-Jan Altitude
Nom_Lac 2 Géologie 2 Cuvette Séjour Hauteur
Lac Lac (km ) (km ) Max (m) (SI) (en °C) (en °C) (m)
(SI) (SI) (jours) (0,1 ou 2)
Abbaye (l' ) LN1 1 8,68 1,00 0,82 19,5 1,84 0,36 22,82 18,40 879,0 345,0 0
Annecy (lac d' ) LN1 6 276,50 0,90 26,50 65,0 2,10 0,64 21,39 19,57 447,0 1168,0 1
Aydat (lac d') LN1 14 27,89 0,00 0,56 15,0 1,77 0,50 17,27 17,93 837,0 173,0 0
Barterand (lac de ) LN1 16 7,82 1,00 0,21 15,0 1,24 0,53 26,01 17,67 295,0 138,0 1
Bordes (lac des ) LN1 21 1,56 0,00 0,14 2,0 1,26 0,52 13,75 17,85 1193,0 44,0 0
Bouchet (lac du ) LN1 22 2,12 0,00 0,43 28,0 1,02 0,60 13,69 19,01 1200,0 2965,8 0
Bourdouze (lac de ) LN1 23 1,25 0,00 0,22 5,0 1,68 0,60 13,70 18,06 1168,0 232,9 0
Cassiere (lac de la ) LN1 29 1,88 0,00 0,13 5,0 1,41 0,40 17,23 17,73 861,0 138,7 0
Chaillexon (lac de ) LN1 65 1033,35 1,00 0,70 31,5 3,81 0,27 16,79 18,35 750,0 5,0 2
Chalain (lac de ) LN1 38 61,53 1,00 2,30 45,0 1,39 0,49 20,58 17,74 488,0 315,0 1
Chambon (lac ) LN1 40 39,18 0,00 0,48 11,0 1,38 0,28 16,58 18,22 875,0 15,0 0
Clairvaux (Grand lac) LN1 55 6,97 1,00 0,56 20,0 1,05 0,60 20,26 18,12 525,0 105,0 0
Etival (grand lac ) LN1 49 4,65 1,00 0,15 9,5 1,60 0,48 17,01 18,37 794,0 10,0 0
Grand maclu (lac du
LN1 68 2,16 1,00 0,21 24,0 1,55 0,48 17,15 18,25 779,0 103,0 0
)
Issarlès (lac d' ) LN1 60 2,01 0,00 0,90 110,0 1,03 0,89 13,73 19,91 1001,0 5095,0 2
Montcineyre (lac de ) LN1 77 2,54 0,00 0,40 22,0 1,23 0,35 13,53 18,07 1180,0 657,0 0
Nantua (lac de ) LN1 80 21,27 1,00 1,42 42,8 1,43 0,71 21,26 18,56 475,0 251,0 0
Paladru LN1 83 47,08 0,00 3,82 35,9 1,74 0,76 22,45 18,67 500,0 1460,0 1
Pavin (lac ) LN1 87 2,13 0,00 0,45 96,0 1,04 0,53 13,18 18,18 1196,0 3192,0 0
Petichet LN1 89 13,90 0,69 0,86 19,2 1,27 0,56 18,22 19,46 923,0 420,0 1
Pierre-châtel (lac de ) LN1 70 2,32 0,36 1,10 10,3 1,02 0,64 18,22 19,46 923,0 239,0 1
Rousses (lac des ) LN1 71 16,20 1,00 0,90 18,2 1,52 0,42 14,31 18,23 1059,0 164,0 0
Saint Point LN1 127 275,29 1,00 5,12 42,0 2,10 0,56 15,94 18,14 849,5 200,0 1
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Sylans (lac de ) LN1 129 11,89 1,00 0,50 22,0 1,55 0,60 20,07 18,51 584,0 210,0 2
Aulnes (étang des) LN2 10 6,18 0,00 0,88 5,5 1,12 0,69 32,06 16,48 11,0 300,0 1
Aureilhan (étang d' ) LN2 12 416,30 0,00 3,29 8,0 1,93 0,25 28,77 13,75 6,0 7,0 0
Biscarrosse (petit
LN2 19 363,61 0,00 0,71 2,0 1,45 0,46 28,77 13,71 21,0 6,0 0
étang de )
Blanc (étang du ) LN2 20 17,92 0,00 1,62 2,0 1,18 0,30 29,16 13,48 12,0 61,0 0
Carcans-Hourtin
LN2 47 438,31 0,00 57,68 15,0 2,47 0,24 27,84 14,16 13,0 665,0 0
(étang de )
Cazaux (étang de ) LN2 30 262,24 0,00 48,39 24,0 1,73 0,43 28,58 13,72 21,0 1587,0 1
Entressen (d' ) LN2 48 177,28 0,00 0,93 10,0 1,26 0,48 31,51 16,89 36,0 3057,0 0
Grand-lieu LN2 54 903,83 0,00 53,24 2,7 2,70 0,67 26,58 13,64 1,0 113,0 0
Parentis LN2 85 310,46 0,00 31,97 22,0 1,94 0,34 28,69 13,79 21,0 372,0 1
Soustons (étang de ) LN2 125 222,93 0,00 4,20 6,0 2,30 0,10 28,91 13,63 3,0 7,7 0
Amance (barrage-
RA1 4 1642,04 1,00 4,50 15,0 2,97 0,30 22,70 16,84 135,0 365,0 2
réservoir aube lac )
Auzon-Temple
(barrage-réservoir RA1 64 52,86 0,01 19,21 20,0 3,54 0,40 22,72 16,88 125,0 365,0 2
aube lac )
Avène (réservoir d' ) RA1 13 142,26 0,61 1,94 57,0 3,90 0,46 25,49 17,35 430,0 120,0 2
Bimont (lac du ) RA1 18 44,50 1,00 0,97 65,0 4,27 0,34 27,78 17,10 330,0 240,0 2
Caramany (retenue de
RA1 27 195,99 0,55 1,78 43,0 3,83 0,35 30,38 16,72 170,0 64,0 2
)
Coiselet (de ) RA1 43 1898,36 1,00 3,80 21,0 4,91 0,55 22,33 18,75 304,0 6,0 1
Fades Besserve
RA1 100 1293,21 0,00 3,52 61,0 6,48 0,32 21,77 16,65 502,0 40,0 2
(retenue des )
Grangent RA1 57 3771,55 0,00 4,09 47,4 6,48 0,29 23,68 17,42 422,0 20,0 2
Laouzas (lac de) RA1 63 147,21 0,00 2,71 48,0 3,27 0,33 22,18 16,70 775,0 51,0 2
Madine (étang de la ) RA1 72 46,68 1,00 10,98 14,0 3,02 0,23 21,00 17,26 227,0 922,8 2
Monteynard-
RA1 76 2019,95 0,29 6,57 135,0 5,10 0,39 22,89 19,53 490,0 55,0 2
Avignonet
Notre-Dame de
RA1 81 2046,41 0,29 1,65 40,0 2,36 0,51 24,32 19,42 363,0 6,0 2
Commiers (de )
Saint Cassien (lac de
RA1 116 161,53 0,25 4,50 55,0 5,09 0,30 31,24 15,80 147,4 90,0 2
)
Saint Etienne de RA1 118 694,51 0,00 5,47 63,0 6,27 0,38 22,63 16,59 517,0 33,0 2
- 193 -
Cantalès
Saint pardoux RA1 126 78,25 0,00 3,25 16,7 4,51 0,42 23,25 15,73 361,0 334,0 1
Sainte Croix (lac de ) RA1 117 1610,63 0,83 21,82 83,0 3,04 0,42 26,61 18,01 477,0 280,0 2
Tolla (lac de ) RA1 130 132,59 0,00 1,15 88,0 2,28 0,55 27,86 15,58 560,0 86,0 2
Villerest (retenue de ) RA1 134 6582,62 0,00 6,97 46,0 9,16 0,55 23,06 17,30 420,0 23,0 2
Vinça (retenue de ) RA1 135 913,41 0,00 1,77 46,0 2,84 0,35 29,25 16,55 244,0 32,0 2
Vouglans (lac de ) RA1 137 1082,16 1,00 16,00 100,0 5,31 0,38 21,11 18,25 429,0 184,0 2
Age (l' ) RA2 2 1126,28 0,00 0,45 20,0 1,56 0,20 24,26 15,88 249,0 2,0 1
Bairon (étang de ) RA2 15 59,28 1,00 0,94 9,0 1,87 0,53 21,49 16,37 174,0 63,3 2
Cercey (barrage de ) RA2 32 4,75 1,00 0,58 12,3 1,25 0,44 21,19 17,67 372,0 699,0 2
Chambon (lac du ) RA2 37 253,09 0,43 1,28 80,0 2,17 0,48 17,37 18,85 1044,0 69,0 2
Chamboux (retenue
RA2 33 17,18 0,00 0,72 18,0 2,10 0,43 19,61 18,13 496,0 236,0 1
de )
Chammet (retenue
RA2 34 109,17 0,00 0,75 15,0 2,72 0,56 18,32 17,16 714,0 51,0 2
du )
Champagney (bassin
RA2 96 0,10 0,00 1,02 30,6 1,75 0,42 19,67 18,37 416,0 60147,0 2
de )
Charmines-Moux (de ) RA2 95 286,95 1,00 0,77 22,0 2,62 0,29 21,91 18,67 381,1 7,0 2
Châtelot (du ) RA2 35 3,23 1,00 0,64 65,0 2,84 0,37 15,85 18,05 716,0 10,0 2
Chaumeçon (barrage
RA2 39 99,50 0,00 1,40 32,0 3,33 0,45 20,74 17,58 385,0 113,0 2
de )
Drapeau (du ) RA2 44 9760,91 0,99 0,57 7,0 1,19 NA 25,35 18,31 170,2 NA NA
Feyt (lac de ) RA2 51 36,77 0,00 0,49 9,2 3,01 0,47 23,37 15,85 487,1 24,0 2
Filleit (barrage du ) RA2 52 10,39 1,00 0,63 20,4 2,39 0,46 26,05 16,53 317,0 18,0 2
Grand-large (du ) RA2 101 12,26 0,01 1,60 3,7 1,35 NA 25,34 18,34 180,0 NA NA
Laprade basse (de ) RA2 104 12,56 0,00 0,89 27,0 1,81 0,33 20,93 16,67 770,0 238,0 2
Méouze (étang de ) RA2 74 8,84 0,00 0,52 5,0 1,86 0,39 18,23 17,19 763,0 72,0 1
Montaubry (barrage de
RA2 78 25,49 0,00 1,28 15,2 2,83 0,37 22,32 18,34 278,0 592,0 1
)
Mouche (barrage de
RA2 79 66,27 1,00 0,83 22,0 2,04 0,41 20,09 18,37 350,0 100,0 1
la )
Ospédale (l' ) RA2 82 2,50 0,00 0,45 25,0 1,52 0,34 27,22 16,39 949,0 180,0 2
Panthier (lac de ) RA2 84 34,68 1,00 1,05 14,3 1,16 0,48 21,21 17,63 373,2 480,0 2
Parroy (étang de ) RA2 86 157,92 1,00 0,56 4,6 1,99 0,47 21,23 17,79 228,0 138,9 2
- 194 -
Pont du Roi (retenue
RA2 110 45,67 0,99 0,45 28,0 2,63 0,31 20,70 18,12 422,0 69,0 2
du )
Puylaurent (retenue de
RA2 91 78,28 0,23 0,60 95,0 4,00 0,25 17,78 19,00 938,8 77,0 2
)
Roujanel RA2 109 97,58 0,00 0,43 47,0 4,19 0,34 21,89 18,59 610,0 33,0 2
Saint-agnan (barrage
RA2 24 47,30 0,04 1,24 10,5 2,15 0,33 19,00 17,84 523,0 70,0 1
de )
Saut de vezoles
RA2 112 8,42 0,00 0,60 20,0 1,63 0,28 15,49 18,67 964,5 105,0 2
(retenue du )
Settons (barrage des
RA2 119 48,45 0,00 3,29 18,0 2,01 0,37 18,72 17,98 580,0 200,0 1
)
Sidiailles (retenue
RA2 121 161,28 0,00 0,75 18,0 3,51 0,41 23,68 16,34 257,0 55,0 1
de)
Sorme (retenue de la) RA2 124 66,63 0,00 2,37 13,0 4,14 0,41 22,40 18,32 296,0 165,0 1
Torcy Neuf (retenue
RA2 113 20,14 0,00 1,51 17,0 3,00 0,36 21,97 18,37 316,0 540,0 1
de )
Torcy Vieux (retenue
RA2 114 13,79 0,00 0,60 11,0 3,22 0,36 21,88 18,34 316,0 181,0 1
de )
Vaivre Vesoul (lac du ) RA2 138 267,62 1,00 0,92 2,0 1,55 0,99 23,48 17,86 215,2 120,0 1
Val Joly (lac du ) RA2 50 73,52 0,00 1,54 5,3 3,62 0,45 20,96 15,60 205,0 6,0 1
Vallee (étang de la ) RA2 132 16,32 0,00 0,54 51,0 2,48 0,40 23,33 15,91 125,0 196,0 1
Vieilles forges
RA2 133 41,85 0,01 1,34 9,5 2,29 0,39 20,20 16,72 241,0 72,0 1
(retenue des )
Alesani (retenue de l' ) RA3 3 58,38 0,01 0,60 60,0 1,96 0,38 27,95 15,75 160,0 165,0 2
Angle guignard
RA3 5 422,94 0,27 0,44 10,0 3,98 0,41 26,25 14,69 28,0 6,0 1
(retenue de )
Apremont (retenue d'
RA3 7 291,46 0,00 1,59 9,0 5,23 0,27 26,84 13,34 12,0 15,0 2
)
Arzal (retenue d' ) RA3 8 10533,35 0,00 6,29 20,0 9,21 0,28 26,00 12,74 0,0 6,0 1
Audomarois (marais
RA3 9 1,49 0,70 0,59 2,8 1,38 0,22 22,59 13,93 3,0 2,0 1
de l' ) (Romelaere)
Aumée (étang ) RA3 11 15,01 0,00 0,52 6,0 1,54 NA 25,92 12,92 6,0 NA NA
Beauregard (étang ) RA3 17 9,01 0,00 0,62 2,5 1,69 0,40 25,40 15,70 89,0 103,0 1
Bois Joli (retenue du
) [complexe du Bois RA3 93 49,01 1,00 0,55 12,0 4,47 NA 25,90 10,82 17,0 NA NA
Joli]
- 195 -
Bultiere (retenue de la
RA3 25 159,56 0,00 0,92 21,0 3,92 0,40 25,98 14,10 51,0 36,0 2
)
Cannes-Ecluse la
maserotte (gravière de RA3 26 10816,88 0,62 0,91 10,0 2,41 NA 24,02 16,26 50,0 NA NA
)
Carcès (lac de ) RA3 28 450,35 0,98 0,91 16,0 2,67 0,54 30,87 16,35 169,0 26,0 1
Cebron (retenue du ) RA3 31 162,40 0,00 1,55 18,0 3,78 0,41 25,08 15,28 117,0 200,0 2
Champsanglard
RA3 97 1053,64 0,00 0,21 19,0 4,77 0,39 23,85 15,91 290,0 2,0 1
(retenue de )
Chapelle Erbrée
RA3 94 125,40 0,00 1,28 13,0 3,15 NA 24,33 13,55 83,0 NA NA
(retenue de la )
Chardes (retenue de )
[complexe de RA3 41 5036,04 0,02 1,05 15,0 3,35 NA 25,77 15,43 123,0 NA NA
Chardes]
Chatillon (étang de ) RA3 36 45,65 0,00 0,82 3,0 2,28 0,81 23,92 13,52 105,0 50,0 1
Cheze (retenue de la ) RA3 111 30,31 0,00 1,18 20,0 5,11 0,42 24,83 12,55 77,0 557,0 2
Codole (retenue de ) RA3 42 41,94 0,00 0,80 25,0 1,14 0,55 31,22 14,28 113,0 167,0 1
Drennec (retenue du ) RA3 45 24,01 0,00 1,13 25,0 2,09 0,31 23,50 10,44 142,0 133,0 2
Duc (étang au ) RA3 46 368,01 0,00 2,12 7,0 2,69 0,34 25,10 12,56 33,0 23,0 0
Forge (étang de la ) RA3 53 97,65 0,00 0,54 2,5 3,81 0,35 25,27 13,29 62,0 5,0 1
Gouet (retenue du ) RA3 56 197,48 0,00 0,76 37,0 4,70 0,28 24,24 11,20 88,0 47,0 2
Graon (retenue du ) RA3 58 44,26 0,00 0,68 18,0 5,17 0,30 26,55 13,93 34,0 62,0 2
Hardouinais (étang de
RA3 59 18,02 0,00 0,37 2,5 1,37 0,72 23,78 12,16 134,0 34,0 0
la )
Jaunay (retenue du) RA3 66 142,35 0,00 1,07 10,7 3,74 0,32 26,78 13,40 10,0 124,0 2
Jouarres (étang de ) RA3 61 8,59 1,00 1,04 4,0 1,10 0,66 31,27 16,73 50,0 426,0 2
Jugon (étang de )
[complexe de Ville- RA3 62 211,00 0,00 0,51 8,0 3,13 NA 25,33 11,49 29,0 NA NA
Hatte]
Kerne Uhel (retenue
RA3 103 75,65 0,00 0,61 10,0 4,50 0,40 23,01 10,93 215,0 17,0 1
de )
Louroux (étang du ) RA3 69 25,20 1,00 0,62 4,0 2,50 0,25 25,10 15,40 98,0 34,4 1
Marcille (étang de ) RA3 73 287,72 0,00 0,91 2,5 3,89 0,40 25,34 13,22 42,0 7,0 0
Marillet (lac du )
RA3 67 132,28 0,00 0,93 16,0 4,74 NA 26,55 14,17 18,0 NA NA
[complexe du Marillet]
- 196 -
Miélan (lac de ) RA3 75 22,29 0,00 0,67 10,0 1,65 0,56 26,91 15,61 217,0 225,0 2
Moulin Neuf (retenue
RA3 105 52,33 0,00 0,54 5,0 2,33 0,60 25,86 10,54 10,0 22,0 2
du )
Moulin Papon (retenue
RA3 106 94,52 0,00 0,85 9,0 3,78 0,57 26,08 13,90 56,0 54,0 2
de )
Moulin ribou (retenue
RA3 107 135,02 0,00 0,88 9,0 3,69 0,44 25,30 14,68 89,0 33,0 2
de )
Pas du houx (étang du
RA3 88 5,20 0,00 0,57 3,0 1,64 0,95 23,72 12,74 156,0 323,0 1
)
Pont-l'Evêque (plan
RA3 90 49,18 NA 0,57 10,0 1,26 NA 23,69 13,27 12,0 NA NA
d'eau de )
Rabodanges (barrage
RA3 92 1018,86 0,52 0,83 17,0 3,62 0,41 22,95 13,61 125,0 7,0 2
de )
Réaltor (bassin du ) RA3 99 19,42 1,00 0,60 10,0 1,68 0,16 29,99 16,89 158,8 127,0 2
Rivière Kersan
RA3 102 2472,87 0,00 0,71 3,0 3,31 NA 25,83 12,81 16,0 NA NA
(gravières de la )
Rochereau RA3 108 211,55 0,01 1,21 13,0 4,81 0,33 25,73 15,13 50,0 27,0 2
Saint Fraimbault de
RA3 120 1799,54 0,01 1,21 8,0 4,53 0,52 24,05 13,67 97,0 3,0 2
Prieres (retenue de )
Saint Michel RA3 123 35,38 0,00 4,48 8,0 2,26 0,37 22,79 10,43 226,0 120,0 1
Sillonniere (retenue
RA3 122 327,05 0,23 1,13 19,0 4,77 0,40 26,17 14,41 39,0 15,8 2
de la )
Sudais (étang de ) RA3 128 12,34 0,57 0,61 1,0 1,73 1,01 24,77 15,45 102,0 10,0 1
Touche Poupard
RA3 131 59,77 0,00 1,25 30,0 4,95 0,37 25,27 15,45 130,0 365,0 2
(retenue de la )
Valiere (retenue de la ) RA3 98 66,19 0,00 1,02 17,0 2,68 0,33 24,66 13,32 83,0 165,0 2
Ville-Hatte (retenue
de la ) - retenue RA3 115 389,77 0,00 1,57 14,0 6,71 0,55 25,67 11,13 19,0 61,0 2
d'Arguenon
Vioreau (étang de ) RA3 136 39,46 0,00 1,88 22,0 3,30 0,18 25,54 13,78 29,0 232,0 1
- 197 -
(b)
Bilan
DensitePop Bilan Bilan Surplus
Groupe Numéro DensitePop BV Densite Reso % du BV % du BV Phosphore LHMS
Nom_Lac Buff1000 (hab Azote BV Phosphore Azote
Lac Lac (hab / km2) (m / km2) Urbain Agric (t) * score
/ km2) (t) BV (t) BV (t)
Erosion
Abbaye (l' ) LN1 1 20,03 21,56 4,89 2,66 41,44 195,34 -67,57 -36,84 -67,57 NA
Annecy (lac d' ) LN1 6 260,46 693,88 2,18 9,39 15,28 190,95 -6,41 -91,42 -23,57 30
Aydat (lac d') LN1 14 14,61 24,87 0,90 1,97 72,04 150,26 13,90 -98,42 21,33 8
Barterand (lac de ) LN1 16 76,74 75,14 0,00 0,00 50,54 171,39 37,17 60,53 81,40 NA
Bordes (lac des ) LN1 21 8,55 8,28 0,00 0,00 67,76 200,15 -134,37 -122,92 -268,73 12
Bouchet (lac du ) LN1 22 23,62 27,58 0,00 0,00 3,80 49,37 13,77 -25,04 36,43 20
Bourdouze (lac de ) LN1 23 8,78 9,76 0,00 0,00 0,00 200,15 -134,37 -122,92 -364,01 6
Cassiere (lac de la ) LN1 29 22,02 29,30 3,85 15,31 39,13 150,78 13,63 -98,32 27,27 14
Chaillexon (lac de ) LN1 65 68,82 93,62 2,67 4,22 45,01 48,83 -20,01 -293,86 -39,14 26
Chalain (lac de ) LN1 38 19,60 26,82 0,46 2,07 33,21 253,12 -2,25 31,29 -4,39 32
Chambon (lac ) LN1 40 13,78 29,23 1,99 1,95 29,32 172,96 -22,64 -38,14 -45,25 16
Clairvaux (Grand lac) LN1 55 37,94 105,58 5,05 4,73 10,88 122,78 -18,16 -37,84 -35,16 NA
Etival (grand lac ) LN1 49 27,67 45,17 4,41 10,78 31,60 129,34 -14,52 -27,78 -14,52 14
Grand maclu (lac du ) LN1 68 11,90 58,23 0,00 0,00 0,00 176,63 -69,03 -44,42 -69,03 20
Issarlès (lac d' ) LN1 60 21,55 24,03 0,00 0,00 1,72 57,78 -22,22 -2,26 -44,43 16
Montcineyre (lac de ) LN1 77 3,70 6,42 0,00 0,00 2,03 197,24 -132,73 -121,88 -265,46 8
Nantua (lac de ) LN1 80 97,20 186,42 11,32 10,46 3,46 233,96 5,62 17,34 24,21 24
Paladru LN1 83 65,60 96,63 0,86 5,89 66,15 247,47 135,38 128,70 676,88 NA
Pavin (lac ) LN1 87 15,52 18,83 0,00 0,00 6,80 191,83 -129,69 -119,94 -389,06 6
Petichet LN1 89 16,38 52,47 1,83 0,00 35,43 228,83 6,49 17,32 22,62 NA
Pierre-châtel (lac de ) LN1 70 33,85 50,62 5,74 0,00 14,13 103,84 10,33 33,24 47,49 22
Rousses (lac des ) LN1 71 106,85 49,55 4,05 5,06 53,55 184,32 -88,41 -74,65 -88,41 14
Saint Point LN1 127 41,71 75,77 2,05 2,73 30,27 112,14 -56,95 -118,78 -141,78 NA
Sylans (lac de ) LN1 129 13,54 17,11 6,12 4,12 18,20 403,98 -5,18 20,84 -6,56 NA
Aulnes (étang des) LN2 10 11,42 9,64 0,00 0,00 49,57 1494,62 349,19 868,95 349,19 NA
Aureilhan (étang d' ) LN2 12 23,11 24,14 2,30 2,16 8,55 87,37 15,79 8,71 15,79 NA
Biscarrosse (petit étang
LN2 19 40,02 489,31 1,97 5,34 11,00 12,72 25,66 -55,31 25,66 12
de )
Blanc (étang du ) LN2 20 62,82 81,30 0,00 3,66 23,54 67,09 34,93 14,80 34,93 NA
Carcans-Hourtin (étang
LN2 47 18,59 6,19 1,33 1,76 13,66 354,83 90,31 78,30 90,31 16
de )
Cazaux (étang de ) LN2 30 22,09 40,48 0,96 4,98 11,74 1,00 102,36 -167,39 102,36 18
Entressen (d' ) LN2 48 NA 48,47 5,32 8,69 54,79 1401,89 328,12 808,99 395,78 NA
Grand-lieu LN2 54 70,93 49,14 2,87 5,01 87,47 719,93 656,94 432,30 849,22 NA
Parentis LN2 85 25,90 46,42 1,88 4,28 10,35 13,24 26,95 -53,73 26,95 22
Soustons (étang de ) LN2 125 26,75 42,10 1,82 4,02 17,21 71,09 28,01 15,48 28,10 22
Amance (barrage-
RA1 4 18,69 28,99 2,28 1,43 45,93 427,31 108,52 218,38 226,17 NA
réservoir aube lac )
Auzon-Temple
(barrage-réservoir aube RA1 64 4,11 7,11 0,30 0,00 5,25 318,53 115,89 126,02 186,97 NA
lac )
Avène (réservoir d' ) RA1 13 10,40 5,76 1,50 0,00 27,54 414,82 34,63 198,37 78,67 NA
Bimont (lac du ) RA1 18 17,16 22,91 0,00 2,36 6,39 110,93 37,75 35,14 38,91 10
Caramany (retenue de ) RA1 27 7,81 4,18 0,89 0,57 23,01 213,02 -30,32 30,03 -43,93 NA
Coiselet (de ) RA1 43 32,20 51,41 2,27 3,10 31,47 164,97 -27,46 -35,12 -47,24 NA
Fades Besserve
RA1 100 20,02 17,87 2,12 0,96 65,69 131,13 88,16 -61,01 123,48 26
(retenue des )
Grangent RA1 57 58,64 88,19 2,23 2,71 53,35 134,22 38,56 -30,67 58,45 18
Laouzas (lac de) RA1 63 12,22 7,48 0,99 0,37 59,87 127,16 88,96 59,67 174,44 26
Madine (étang de la ) RA1 72 13,91 11,86 1,40 2,29 45,04 20,08 17,05 -62,54 20,63 18
Monteynard- Avignonet RA1 76 12,80 14,56 1,42 0,52 20,07 200,77 28,60 96,03 113,90 NA
Notre-Dame de
RA1 81 12,77 9,26 1,43 0,51 20,26 198,70 28,31 94,95 112,20 26
Commiers (de )
Saint Cassien (lac de ) RA1 116 60,85 19,83 2,35 6,60 20,78 177,68 40,66 41,49 68,86 32
Saint Etienne de
RA1 118 82,97 21,64 2,89 4,71 59,86 27,05 40,37 -43,17 123,85 NA
Cantalès
Saint pardoux RA1 126 17,47 13,99 1,13 2,10 26,07 113,08 58,58 -30,24 82,15 NA
Sainte Croix (lac de ) RA1 117 7,73 19,80 1,30 0,35 7,91 134,10 -0,85 19,81 -2,81 22
Tolla (lac de ) RA1 130 6,07 5,91 0,52 0,38 0,81 62,00 -29,59 -43,43 -88,77 24
Villerest (retenue de ) RA1 134 99,53 39,32 2,47 3,93 58,79 181,36 76,58 25,78 111,75 24
- 199 -
Vinça (retenue de ) RA1 135 17,59 120,70 1,30 0,86 8,13 243,82 -37,59 23,18 -85,66 NA
Vouglans (lac de ) RA1 137 33,51 22,67 2,23 2,62 39,52 126,09 -11,11 -50,72 -20,02 10
Age (l' ) RA2 2 29,27 9,54 2,04 2,03 58,07 111,84 75,40 -85,55 123,24 NA
Bairon (étang de ) RA2 15 29,84 34,58 1,37 1,79 61,16 118,57 54,17 1,11 101,48 26
Cercey (barrage de ) RA2 32 12,67 16,14 0,00 2,06 62,62 297,06 211,96 -133,24 319,84 NA
Chambon (lac du ) RA2 37 5,42 95,15 0,92 0,00 1,30 120,09 0,63 -5,88 3,17 NA
Chamboux (retenue de
RA2 33 11,81 7,70 0,78 0,00 59,43 71,01 57,27 -147,69 57,27 8
)
Chammet (retenue du
RA2 34 15,51 10,72 0,55 0,66 36,77 79,30 -20,55 -3,19 -38,79 16
)
Champagney (bassin
RA2 96 7,83 16,48 0,00 0,00 42,56 216,04 21,82 -16,53 21,82 NA
de )
Charmines-Moux (de ) RA2 95 85,06 32,40 4,50 7,02 29,22 198,94 12,56 26,89 31,10 32
Châtelot (du ) RA2 35 8,96 23,39 0,00 0,00 56,81 26,46 27,04 -188,10 134,55 NA
Chaumeçon (barrage
RA2 39 8,74 8,48 1,79 0,49 45,37 74,13 50,01 -100,62 50,01 NA
de )
Drapeau (du ) RA2 44 103,02 552,55 2,91 5,85 40,96 228,10 52,71 29,24 173,70 NA
Feyt (lac de ) RA2 51 29,20 29,95 0,42 0,36 40,46 42,27 13,78 -11,93 17,60 NA
Filleit (barrage du ) RA2 52 5,57 6,39 0,00 0,00 22,35 83,18 -3,59 28,79 -10,41 16
Grand-large (du ) RA2 101 1176,28 1108,48 8,36 41,93 42,12 1374,95 545,68 690,82 3244,51 NA
Laprade basse (de ) RA2 104 16,82 13,25 0,00 0,00 14,22 478,23 121,17 338,55 226,05 20
Méouze (étang de ) RA2 74 11,55 11,85 1,59 0,00 26,21 72,60 3,68 -63,38 5,72 30
Montaubry (barrage de
RA2 78 51,06 24,09 5,45 6,93 63,61 -24,72 98,90 -263,37 105,15 28
)
Mouche (barrage de la
RA2 79 19,59 22,33 3,32 0,76 69,77 452,85 169,10 204,75 233,93 NA
)
Ospédale (l' ) RA2 82 16,86 14,90 6,30 0,00 11,31 149,14 -1,51 26,74 -4,53 12
Panthier (lac de ) RA2 84 14,37 15,73 0,58 2,18 75,52 369,00 172,05 37,37 200,21 36
Parroy (étang de ) RA2 86 33,06 34,94 2,05 2,73 63,55 121,97 40,65 60,15 46,09 16
Pont du Roi (retenue du
RA2 110 21,07 21,91 3,32 0,00 63,03 16,26 91,27 -217,69 91,27 10
)
Puylaurent (retenue de
RA2 91 11,89 2,87 2,87 0,00 22,66 865,74 -81,24 30,56 -156,60 NA
)
Roujanel RA2 109 6,20 3,53 2,58 0,00 1,17 884,14 -83,26 30,28 -218,76 NA
- 200 -
Saint-agnan (barrage
RA2 24 10,69 12,97 1,98 0,00 34,51 59,86 51,52 -102,87 51,52 16
de )
Saut de vezoles
RA2 112 8,82 9,23 0,00 0,00 3,57 301,64 10,67 52,30 13,08 8
(retenue du )
Settons (barrage des ) RA2 119 9,69 8,32 1,45 0,99 37,13 163,47 85,18 -158,40 117,48 NA
Sidiailles (retenue de) RA2 121 14,02 10,49 0,73 0,75 94,40 125,42 113,81 30,31 250,40 18
Sorme (retenue de la) RA2 124 21,98 38,17 1,02 0,69 79,42 -192,72 263,36 -597,79 263,36 20
Torcy Neuf (retenue de
RA2 113 1006,53 626,40 10,28 55,03 27,39 -190,25 261,79 -595,27 1448,66 24
)
Torcy Vieux (retenue de
RA2 114 403,74 420,03 1,13 32,81 47,69 -192,11 263,47 -598,24 700,94 8
)
Vaivre Vesoul (lac du ) RA2 138 82,98 263,20 3,47 7,18 51,81 148,27 78,28 -15,99 149,41 24
Val Joly (lac du ) RA2 50 63,58 23,76 0,71 3,28 43,68 1294,17 342,93 691,52 342,93 14
Vallee (étang de la ) RA2 132 26,44 31,93 0,00 1,57 20,07 115,95 49,58 32,64 49,58 16
Vieilles forges (retenue
RA2 133 34,95 20,06 2,82 2,01 13,56 72,98 6,05 -14,19 6,44 26
des )
Alesani (retenue de l' ) RA3 3 9,17 18,43 2,21 0,43 0,00 96,85 -50,67 -18,91 -152,00 20
Angle guignard (retenue
RA3 5 56,54 23,45 3,04 4,70 91,06 825,48 718,94 664,35 1834,13 18
de )
Apremont (retenue d' ) RA3 7 68,98 39,23 2,50 4,30 93,62 564,95 594,09 398,79 953,01 20
Arzal (retenue d' ) RA3 8 90,07 52,80 2,38 4,46 83,39 571,02 372,83 354,37 980,61 12
Audomarois (marais de
RA3 9 100,35 126,35 0,00 0,01 62,76 1808,92 855,63 1101,29 855,63 10
l' ) (Romelaere)
Aumée (étang ) RA3 11 36,28 50,67 4,15 0,00 80,95 343,76 184,51 224,98 456,98 12
Beauregard (étang ) RA3 17 3,45 28,80 0,00 0,00 7,91 185,64 47,56 51,44 47,56 NA
Bois Joli (retenue du )
[complexe du Bois RA3 93 90,56 50,58 2,91 6,03 84,56 557,63 327,15 300,65 937,53 NA
Joli]
Bultiere (retenue de la ) RA3 25 72,83 34,54 3,21 5,73 88,20 856,76 830,61 669,96 1454,73 22
Cannes-Ecluse la
RA3 26 35,02 632,65 2,51 2,53 64,09 738,69 220,66 413,41 376,74 NA
maserotte (gravière de )
Carcès (lac de ) RA3 28 47,36 13,09 2,60 6,56 25,25 391,67 45,82 109,60 114,57 30
Cebron (retenue du ) RA3 31 43,42 31,72 2,89 1,33 92,86 767,42 476,79 561,50 847,41 16
Champsanglard
RA3 97 30,32 8,65 2,17 2,14 57,12 116,98 73,14 -80,70 118,63 6
(retenue de )
- 201 -
Chapelle Erbrée
RA3 94 32,43 52,67 1,82 1,21 95,21 498,14 346,67 324,42 983,30 NA
(retenue de la )
Chardes (retenue de )
RA3 41 64,15 155,93 2,40 3,20 60,92 129,60 107,81 -19,36 257,86 NA
[complexe de Chardes]
Chatillon (étang de ) RA3 36 34,40 33,33 2,02 2,21 92,39 373,99 276,23 222,70 811,98 NA
Cheze (retenue de la ) RA3 111 32,38 27,01 3,13 2,46 88,07 303,00 212,08 164,55 644,39 NA
Codole (retenue de ) RA3 42 13,12 16,33 2,47 0,04 18,68 139,88 -34,13 52,06 -102,40 NA
Drennec (retenue du ) RA3 45 33,81 28,57 0,35 0,13 48,85 795,71 440,51 635,55 1650,51 NA
Duc (étang au ) RA3 46 64,77 103,52 2,59 3,35 80,53 705,55 434,44 410,91 882,59 16
Forge (étang de la ) RA3 53 55,00 60,02 2,07 3,39 82,47 706,68 551,69 435,48 1100,25 14
Gouet (retenue du ) RA3 56 58,98 105,92 3,06 3,34 84,72 563,37 383,86 357,94 1400,59 NA
Graon (retenue du ) RA3 58 44,58 56,06 1,95 3,04 91,32 168,53 181,63 95,26 328,21 16
Hardouinais (étang de
RA3 59 55,92 78,40 2,79 1,28 41,79 770,10 445,74 479,48 1501,37 NA
la )
Jaunay (retenue du) RA3 66 50,90 36,87 1,39 2,77 93,17 662,74 613,60 456,45 1456,87 18
Jouarres (étang de ) RA3 61 76,36 56,73 3,63 0,96 86,08 218,29 -6,32 30,54 -6,32 20
Jugon (étang de )
[complexe de Ville- RA3 62 59,95 74,22 2,15 3,16 84,32 1320,43 740,36 877,08 1346,42 NA
Hatte]
Kerne Uhel (retenue de
RA3 103 34,40 23,71 0,78 0,40 91,89 564,51 503,12 391,83 1131,72 20
)
Louroux (étang du ) RA3 69 17,14 38,41 1,30 2,49 89,15 531,48 196,99 363,69 209,14 NA
Marcille (étang de ) RA3 73 68,43 76,86 2,13 2,75 90,72 1080,05 789,03 730,67 1429,49 18
Marillet (lac du )
RA3 67 55,00 74,01 1,92 2,45 90,83 627,77 559,36 329,80 827,31 16
[complexe du Marillet]
Miélan (lac de ) RA3 75 42,25 54,11 4,88 1,80 78,23 159,39 78,45 124,31 381,34 24
Moulin Neuf (retenue du
RA3 105 63,80 118,46 1,00 1,22 87,39 329,16 273,81 230,11 548,46 20
)
Moulin Papon (retenue
RA3 106 121,58 304,55 1,64 6,66 89,33 510,82 609,45 267,42 774,21 20
de )
Moulin ribou (retenue
RA3 107 41,77 117,68 2,55 2,76 86,75 1014,95 821,24 617,32 1976,21 NA
de )
Pas du houx (étang du ) RA3 88 35,43 36,81 0,00 0,00 29,16 282,97 98,62 90,97 98,62 NA
Pont-l'Evêque (plan
RA3 90 46,42 177,94 5,39 0,25 93,49 184,32 27,98 105,30 33,39 NA
d'eau de )
- 202 -
Rabodanges (barrage
RA3 92 40,10 14,72 3,09 2,41 83,85 247,16 75,87 137,48 126,75 18
de )
Réaltor (bassin du ) RA3 99 269,81 116,47 5,88 21,70 22,29 109,74 28,51 17,26 28,51 12
Rivière Kersan
RA3 102 182,38 169,09 2,96 7,56 82,08 573,30 355,50 317,58 808,50 NA
(gravières de la )
Rochereau RA3 108 51,09 37,06 2,75 4,00 92,71 927,81 824,00 754,94 2267,05 20
Saint Fraimbault de
RA3 120 30,74 52,43 2,02 1,64 90,36 659,82 345,61 356,41 938,56 12
Prieres (retenue de )
Saint Michel RA3 123 53,81 36,50 2,89 0,51 20,54 548,81 344,49 319,91 643,98 20
Sillonniere (retenue
RA3 122 47,27 40,88 1,36 2,88 91,34 1103,64 1012,32 832,45 2204,68 10
de la )
Sudais (étang de ) RA3 128 15,05 13,45 1,47 0,00 72,14 340,91 115,94 190,58 192,14 NA
Touche Poupard
RA3 131 34,03 21,90 0,81 1,17 93,76 522,10 315,12 361,35 1029,39 20
(retenue de la )
Valiere (retenue de la ) RA3 98 84,68 104,81 4,25 6,51 79,71 998,83 592,50 659,28 1977,39 NA
Ville-Hatte (retenue de
la ) - retenue RA3 115 52,43 48,63 2,41 2,74 83,62 1125,40 607,64 773,39 1343,16 NA
d'Arguenon
Vioreau (étang de ) RA3 136 30,43 14,89 2,45 3,28 63,15 1371,84 877,84 804,68 2135,97 34
- 203 -
Annexe 3 : Classification des 42 espèces pisicoles en guildes de reproduction, trophiques et de tolérance.
Ces espèces piscicoles ont été échantillonnées dans les plans d’eau analysés de l’étude présentée au chapitre 4.
Overall Overall
Strict Strict Omnivores
Species Intolerant Tolerant Lithophilic Invertivores Planktivores Herbivores Piscivores
Lithophilic Phytophilic Generalist
species species
Abramis brama 0 1 1 0 0 1 1 1 0 0
Acipenser ruthenus 0 0 1 1 0 0 1 0 0 0
Alburnoides bipunctatus 1 0 1 1 0 0 1 1 0 0
Alburnus alburnus 0 1 1 0 0 1 1 1 0 0
Ameiurus melas 0 1 1 1 0 0 1 0 0 1
Anguilla anguilla 0 1 0 0 0 0 1 0 0 1
Barbatula barbatula 0 0 1 0 0 0 1 0 0 0
Barbus barbus 0 0 1 1 0 0 1 0 0 0
Blicca bjoerkna 0 1 0 0 1 0 1 1 0 0
Carassius auratus auratus 0 1 0 0 1 1 1 1 1 0
Carassius carassius 0 1 0 0 1 1 1 1 1 0
Coregonus lavaretus 1 0 1 1 0 0 1 1 0 0
Cottus gobio 1 0 1 1 0 0 1 0 0 0
Cyprinus carpio 0 1 1 0 0 1 1 1 0 0
Esox lucius 0 0 0 0 1 0 0 0 0 1
Gambusia affinis 0 1 0 0 0 0 1 1 0 0
Gobio gobio 0 0 1 0 0 0 1 0 1 0
Gymnocephalus cernuus 0 0 1 0 0 0 1 0 0 0
Hypophthalmichthys molitrix 0 1 0 0 0 0 0 1 1 0
Lepomis gibbosus 0 1 1 1 0 0 1 1 0 0
Leucaspius delineatus 1 0 0 0 1 0 1 1 0 0
Leuciscus idus 1 0 1 0 0 1 1 1 1 1
Leuciscus leuciscus 0 0 1 1 0 1 1 1 0 0
Liza aurata 0 0 0 0 0 1 0 1 1 0
Liza ramada 0 1 1 1 0 0 1 0 1 0
Lota lota 0 0 1 1 0 0 1 0 0 1
Oncorhynchus mykiss 0 0 1 1 0 0 1 0 0 1
Perca fluviatilis 0 1 1 0 0 0 1 0 0 1
Phoxinus phoxinus 0 0 1 1 0 0 1 0 0 0
- 204 -
Rhodeus amarus 1 0 0 0 0 0 1 0 1 0
Rutilus rutilus 0 1 1 0 0 1 1 1 1 0
Salaria fluviatilis 0 0 1 1 0 0 1 0 0 0
Salmo trutta fario 1 0 1 1 0 0 1 0 0 1
Salvelinus alpinus 1 0 1 1 0 0 1 0 0 1
Salvelinus namaycush 1 0 1 1 0 0 1 0 0 1
Sander lucioperca 0 0 1 0 0 0 0 0 0 1
Scardinius
erythrophthalmus 0 0 0 0 1 1 1 1 0 0
Silurus glanis 0 0 0 0 1 0 1 0 0 1
Squalius cephalus 0 0 1 0 0 1 1 1 1 1
Telestes souffia 1 0 1 1 0 0 1 0 0 0
Thymallus thymallus 1 0 1 1 0 0 1 0 0 0
Tinca tinca 0 1 0 0 1 0 1 0 0 1
- 205 -
Annexe 4 : Classification des 41 espèces piscicoles en guildes de reproduction, trophiques et de tolérance.
Overall intolerant Overall tolerant Strict Strict Omnivores
Species Lithophilic Invertivores Planktivores Herbivores Piscivores
species species lithophilic phytophilic generalist
Abramis brama 0 1 1 0 0 1 1 1 0 0
Acipenser ruthenus 0 0 1 1 0 0 1 0 0 0
Alburnoides
1 0 1 1 0 0 1 1 0 0
bipunctatus
Alburnus alburnus 0 1 1 0 0 1 1 1 0 0
Ameiurus melas 0 1 1 1 0 0 1 0 0 1
Anguilla anguilla 0 1 0 0 0 0 1 0 0 1
Barbatula barbatula 0 0 1 0 0 0 1 0 0 0
Barbus barbus 0 0 1 1 0 0 1 0 0 0
Blicca bjoerkna 0 1 0 0 1 0 1 1 0 0
Carassius auratus
0 1 0 0 1 1 1 1 1 0
auratus
Carassius carassius 0 1 0 0 1 1 1 1 1 0
Coregonus lavaretus 1 0 1 1 0 0 1 1 0 0
Cottus gobio 1 0 1 1 0 0 1 0 0 0
Cyprinus carpio 0 1 1 0 0 1 1 1 0 0
Esox lucius 0 0 0 0 1 0 0 0 0 1
Gambusia affinis 0 1 0 0 0 0 1 1 0 0
Gobio gobio 0 0 1 0 0 0 1 0 1 0
Gymnocephalus
0 0 1 0 0 0 1 0 0 0
cernuus
Hypophthalmichthys
0 1 0 0 0 0 0 1 1 0
molitrix
Lepomis gibbosus 0 1 1 1 0 0 1 1 0 0
Leucaspius delineatus 1 0 0 0 1 0 1 1 0 0
Leuciscus idus 1 0 1 0 0 1 1 1 1 1
Leuciscus leuciscus 0 0 1 1 0 1 1 1 0 0
Liza aurata 0 0 0 0 0 1 0 1 1 0
Liza ramada 0 1 1 1 0 0 1 0 1 0
- 206 -
Lota lota 0 0 1 1 0 0 1 0 0 1
Oncorhynchus mykiss 0 0 1 1 0 0 1 0 0 1
Perca fluviatilis 0 1 1 0 0 0 1 0 0 1
Phoxinus phoxinus 0 0 1 1 0 0 1 0 0 0
Rhodeus amarus 1 0 0 0 0 0 1 0 1 0
Rutilus rutilus 0 1 1 0 0 1 1 1 1 0
Salaria fluviatilis 0 0 1 1 0 0 1 0 0 0
Salmo trutta fario 1 0 1 1 0 0 1 0 0 1
Salvelinus alpinus 1 0 1 1 0 0 1 0 0 1
Salvelinus namaycush 1 0 1 1 0 0 1 0 0 1
Sander lucioperca 0 0 1 0 0 0 0 0 0 1
Scardinius
0 0 0 0 1 1 1 1 0 0
erythrophthalmus
Silurus glanis 0 0 0 0 1 0 1 0 0 1
Squalius cephalus 0 0 1 0 0 1 1 1 1 1
Thymallus thymallus 1 0 1 1 0 0 1 0 0 0
Tinca tinca 0 1 0 0 1 0 1 0 0 1
- 207 -
Annexe 5 : Caractéristiques des 41 espèces de poissons capturées lors d’une campagne d’échantillonnage par plan d’eau, effectuée selon
un protocole d’échantillonnage standardisé (C.E.N. 2005).
Fréquence des occurrences des espèces piscicoles (les valeurs supérieures à 75% sont en gras) et abondances relatives moyennes (RA) par espèce
dans les lacs naturels et les reservoirs échantillonnés. La RA de chaque espèce a été calculée comme le ratio entre le nombre total d’individus
appartenant à une espèce, et le nombre total d’individus capturés durant la campagne d’échantillonnage.
Frequency (%) Mean relative abundance (%)
Common name Species
Reservoirs Natural lakes Reservoirs Natural lakes
Bream Abramis brama 83.58 50.00 10.15 5.06
Sterlet Acipenser ruthenus 1.49 0.00 0.00 0.00
Spirlin Alburnoides bipunctatus 0.00 4.17 0.00 0.02
Bleak Alburnus alburnus 61.19 37.50 2.62 2.91
Brown bullhead Ameiurus melas 34.33 25.00 7.65 1.25
Europena eel Anguilla anguilla 1.49 0.00 0.00 0.00
Stone loach Barbatula barbatula 2.99 4.17 0.14 0.00
Barbel Barbus barbus 7.46 0.00 0.04 0.00
White bream Blicca bjoerkna 70.15 12.50 10.11 1.45
Goldfish Carassius auratus auratus 7.46 0.00 0.01 0.00
Crucian carp Carassius carassius 13.43 8.33 0.03 0.13
Common
Coregonus lavaretus 2.99 29.17 0.03 2.12
whitefish
Bullhead Cottus gobio 1.49 8.33 0.02 0.05
Common carp Cyprinus carpio 44.78 16.67 0.38 0.02
Northern pike Esox lucius 59.70 66.67 0.15 0.33
Mosquitofish Gambusia affinis 1.49 0.00 0.01 0.00
Gudgeon Gobio gobio 13.43 37.50 0.96 1.83
Ruffe Gymnocephalus cernuus 67.16 41.67 5.03 3.84
Silver carp Hypophthalmichthys molitrix 1.49 0.00 0.01 0.00
Pumpkinseed Lepomis gibbosus 32.84 25.00 0.20 0.15
Belica Leucaspius delineatus 2.99 0.00 0.00 0.00
Ide Leuciscus idus 1.49 0.00 0.00 0.00
Common dace Leuciscus leuciscus 5.97 12.50 0.03 0.04
Golden grey
Liza aurata 1.49 0.00 0.00 0.00
mullet
Thinlip mullet Liza ramada 1.49 0.00 0.10 0.00
Burbot Lota lota 0.00 4.17 0.00 0.01
Rainbow trout Oncorhynchus mykiss 8.96 12.50 0.71 0.47
European perch Perca fluviatilis 97.01 95.83 23.29 31.19
Eurasian
Phoxinus phoxinus 1.49 0.00 0.00 0.00
minnow
Bitterling Rhodeus amarus 2.99 0.00 0.03 0.00
Roach Rutilus rutilus 95.52 100.00 29.94 37.65
Freshwater
Salaria fluviatilis 0.00 4.17 0.00 0.01
blenny
Brown trout Salmo trutta fario 10.45 16.67 0.85 0.07
Charr Salvelinus alpinus 1.49 16.67 0.00 1.01
Lake trout Salvelinus namaycush 0.00 8.33 0.00 0.78
Pike perch Sander lucioperca 83.58 41.67 4.87 1.70
Rudd Scardinius erythrophthalmus 82.09 70.83 2.04 6.49
Wels catfish Silurus glanis 19.40 4.17 0.04 0.01
European chub Squalius cephalus 19.40 45.83 0.40 0.83
Grayling Thymallus thymallus 1.49 0.00 0.02 0.00
Tench Tinca tinca 22.39 41.67 0.10 0.57
- 209 -

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UNIVERSITE AIX-MARSEILLE I - Université de Provence

ECOLE DOCTORALE DES SCIENCES DE L’ENVIRONNEMENT (251) – CEREGE Aix-

Pour obtenir le grade de DOCTEUR

Impact des facteurs anthropiques sur les

Soutenue publiquement le 24 juin 2011

Travaux réalisés dans l’Equipe Ecosystèmes lacustres

Cette partie qui me tient particulièrement à cœur me permet de témoigner ma gratitude

Je tiens tout d’abord à remercier Christine Argillier d’avoir accepté la direction de ce

Table des matières ..................................................................................................................- 5 -

Tableau 2. 1 : Critères de classification des espèces piscicoles en guildes fonctionnelles. .- 30 -

Tableau 3. 1: Valeur moyenne et écart type des principales caractéristiques environnementales

Figure 4. 1 : Partitionnement de la variance d’une métrique biologique en quatre

Figure 5. 1: Location of the 91 study sites in France. ..........................................................- 93 -

ACP : Analyse en Composantes Principales.

1.2. Les poissons en bioindication lacustre

1.3. Objectifs de l’étude et organisation générale du mémoire

Ainsi, ce mémoire de thèse est structuré en 4 chapitres.

Le chapitre 2 décrit brièvement l’ensemble des données piscicoles, environnementales et

Données disponibles à l’échelle de la France

2.1.1. Les inventaires piscicoles

2.1.2. Les caractéristiques biologiques des espèces

Tableau 2. 1 : Critères de classification des espèces piscicoles en guildes fonctionnelles.

L’utilisation de ces guildes permet une caractérisation fonctionnelle des assemblages,

2.2. Les caractéristiques environnementales naturelles des lacs et de leurs bassins

2.3. Les facteurs anthropiques

En France, la biodiversité des écosystèmes aquatiques est fortement menacée,

2.3.1. Les pressions anthropiques à l’échelle du bassin versant

2.3.2. Les pressions anthropiques locales

Par conséquent, ce travail souligne non seulement la nécessité de collecter d’autres

3.2. Matériel et méthodes

3.2.1. Réalisation de la typologie lacustre

Pour la réalisation de groupes de systèmes homogènes du point de vue de l’environnement

Tableau 3. 1: Valeur moyenne et écart type des principales caractéristiques

3.2.2. Analyse de la réponse de l’ichtyofaune aux pressions dans chaque type

3.2.2.1. Les données poissons

3.2.2.2. Les données de pressions

3.2.2.3. Réponse des métriques aux pressions

3.3.1. Typologie physique des plans d’eau français

Figure 3. 5: Représentation des principales caractéristiques environnementales naturelles des

3.3.2.1. Analyse globale

3.3.2.2. Analyse par types

Figure 3. 8: Représentation des trois indices synthétiques de pressions anthropiques pour

3.3.3. Calcul des métriques piscicoles et réponse aux pressions

Groupe 3 des retenues (n = 10)

Sélection de métriques piscicoles et variabilité environnementale : comparaison entre

Figure 4. 1 : Partitionnement de la variance d’une métrique biologique en quatre

4.2. Study Sites

Figure 4. 2: Location of the 89 study sites in the French hydrographic network.

Tableau 4. 4: Proposed guild classification for French lake fishes.

Can reproduce on unsilted mineral substrates whose

Feed on both animal and vegetal items (more than 25%

Metric ~ (AL + ADB + Dmax + SLDF + Vd + SumT + DiffT + RT + GI + WL) + PA + PU (1)

Assessing lake conditions through fish assemblages

The selected fish-based metrics

hindcasting approach (sous presse dans Ecological Indicators – DOI:

Lionel Launois, Jacques Veslot, Pascal Irz, and Christine Argillier

The European Water Framework Directive (WFD, 2000/60/EC) (European Community

5.2.1. Study sites

5.2.2. Abiotic environmental characteristics of lakes

5.2.3. Fish sampling procedures

5.2.4. Candidate metrics

5.2.5. Catchment-scale anthropogenic pressures

5.2.6. Local-scale anthropogenic pressures