Apes

Hydrobiologia (2013) 704:417436
DOI 10.1007/s10750-012-1250-6
WATER BODIES IN EUROPE
Global warming and potential shift in reference conditions:

the case of functional fish-based metrics
Maxime Logez Didier Pont
Received: 29 March 2012 / Accepted: 7 July 2012 / Published online: 1 August 2012
Springer Science+Business Media B.V. 2012
Abstract The reference condition approach, advocated by the Water Framework Directive, is the basis
of most currently used multimetric indices using
functional traits of fish species. The ecological status
of streams is assessed by measuring the deviation of
the observed trait values from the theoretical values of
reference conditions in the absence of anthropogenic
disturbances. While reference conditions serve as
baselines for ecological assessment, they vary with
natural environmental conditions. Therefore, global
warming appears to be a major threat to the use of
current indices for diagnosing future stream conditions, as climate change is projected to modify
assemblage composition, suggesting that the functional structure of fish assemblages will also be
affected. The main objectives of this study are to
assess the potential effect of climate change on the
trait composition of fish assemblages and the consequences for the establishment of reference conditions.
Guest editors: C. K. Feld, A. Borja, L. Carvalho & D. Hering /

Water bodies in Europe: integrative systems to assess
ecological status and recovery
Electronic supplementary material The online version of

this article (doi:10.1007/s10750-012-1250-6) contains
supplementary material, which is available to authorized users.
M. Logez (&) D. Pont
Irstea, UR HBAN, 1 rue Pierre-Gilles de Gennes,
CS 10030, 92761 Antony, France
e-mail: maxime.logez@irstea.fr
The results highlight the relation between environmental, especially climatic, conditions and functional
traits and project the effects of climate change on
trait composition. Traits based on species intolerance
are expected to be most negatively affected by the
projected climatic shift. The consequences for the
development of multimetric indices based on fish
functional traits are discussed.
Keywords IBI Climate change Riverine fish
assemblages Functional trait Reference condition
Water Framework Directive Local species richness
Introduction
Reference conditions, as defined by the European
Water Framework Directive (WFD), serve as baselines to assess the ecological status of streams by
comparing the observed biotic assemblages with
theoretical assemblages in the absence of human
disturbance (Bailey et al., 1998; Hawkins et al.,
2010). The WFD also advocates that the ecological
assessment should integrate several biotic assemblage
descriptors, also called metrics (Hering et al., 2006),
into a synthetic index. These multimetric indices must
reflect the level of impairment of a given site (Karr &
Chu, 1999; Oberdorff et al., 2002; Pont et al., 2006,
2007). Several multimetric indices based on fish
assemblages (Oberdorff et al., 2002; Pont et al.,
2006, 2007, 2011; Argillier et al., this issue) use the
123
418
principles of the index of biotic integrity (IBI)

developed by Karr (1981), which compares observed
values of several assemblage descriptors with theoretical values that would be observed in pristine
sites, a concept very close to that of reference
conditions (Karr, 1981). Since the first IBI, several
changes have been made to enhance the assessment
of ecological conditions with multimetric indices. One
of the main improvements was to consider metric
variability (Fausch et al., 1984; Karr & Chu, 1999;
Pont et al., 2006, 2007; Roset et al., 2007; Logez &
Pont, 2011) to disentangle the effects of impairment
from natural variability (Oberdorff et al., 1995, 2002;
Hughes et al., 1998; Stoddard et al., 2008). Nevertheless, the establishment of reference conditions remains
essential to develop and use accurate bioassessment
tools.
The historical approach was used by several authors
to define the reference conditions from past evidence
of system functioning (Swetnam et al., 1999; Muxika
et al., 2007). In contrast to this palaeoecological
approach, for most ecosystem types, reference conditions are very often defined in a more pragmatic
manner (Stoddard et al., 2006; Hering et al., this
issue), and very often the minimally disturbed conditions that are observed in a region are used (Logez &
Pont, 2011). All these approaches are based on the
assumption of relatively stable biotic communities,
so that the observed deviations consistently reflect
the effects of human disturbances. Comparing the
observed biota with reference values of a differently
functioning river, however, would lead to inaccurate
assessment due to inconsistencies in reference condition baselines. Nevertheless, with the projected modifications of climatic conditions, the assumption of
stable and consistent reference conditions is less likely
to be met. Temperature and precipitation are two
major components of hydrosystem functioning (Allan
& Castillo, 2007) which are strongly affected by
climatic changes (Webb & Nobilis, 1995; Webb,
1996; Webb & Nobilis, 2007; Albelk & Albek, 2009).
The consequences of global change, and in particular of temperature modifications, for riverine fish
assemblages are mostly studied by addressing fish
species distributions rather than assemblage structure.
Projected distributions using various gas emission
scenarios (IPCC, 2007) highlight a shift in fish species
distributions following the predicted changes in
climatic conditions (Buisson et al., 2008; Buisson &
123
Grenouillet, 2009; Grenouillet et al., 2011). Following

the theory of habitat filtering, local assemblages are
composed of species selected on their traits by the
environmental conditions (Keddy, 1992). Among the
environmental drivers, temperature was shown to be
the major determinant of the functional structure of
European fish assemblages (Logez et al., 2012a).
Therefore, global warming will likely affect both the
composition and the functional structure of fish
assemblages.
Functional traits which group species having
a common biological or ecological characteristic into
a single variable, e.g. species intolerant of habitat
degradation, are now commonly used to develop
bioassessment tools based on fish assemblages (Oberdorff et al., 2002; Pont et al., 2006, 2007, 2009;
Logez & Pont, 2011). Ecological theory suggests that
these traits are directly or indirectly related to
ecosystem functioning (Lavorel & Garnier, 2002),
because of either their response to or their effect on
the ecosystem. Functional traits are assumed to react
to human pressure in a consistent way across a wide
range of streams and over a very broad spatial extent
(Pont et al., 2006, 2007). While patterns derived from
assemblage compositions are controlled by species
distribution areas and thus by biogeographical drivers
(Hoeinghaus et al., 2007), patterns derived from
functional traits are mostly related to the environmental conditions present (e.g. Lamouroux et al., 2002;
Goldstein & Meador, 2004; Statzner et al., 2004;
Bonada et al., 2007; Ibanez et al., 2009; Logez et al.,
2010). In addition to the projected shift in species
distribution, traitenvironment relationships suggest
that fish assemblage structures will change to cope
with the new climatic conditions. It is hypothesised,
however, that functional traits will respond differently
to global change compared with species composition.
Two major approaches are used to control the
environmental variability in metrics. The type-specific
approach groups sites based on their environmental
similarities, whereas with the site-specific approach
the assemblage functional structure is directly predicted from environmental variables (Roset et al.,
2007). Some recent bioassessment methods use this
latter predictive approach to estimate the theoretical
value of a metric that should have been observed in the
absence of human pressures (Pont et al., 2006, 2007;
Hawkins et al., 2010; Logez & Pont, 2011). For
instance, the European Fish Index (EFI), a multimetric
418
principles of the index of biotic integrity (IBI)

developed by Karr (1981), which compares observed
values of several assemblage descriptors with theoretical values that would be observed in pristine
sites, a concept very close to that of reference
conditions (Karr, 1981). Since the first IBI, several
changes have been made to enhance the assessment
of ecological conditions with multimetric indices. One
of the main improvements was to consider metric
variability (Fausch et al., 1984; Karr & Chu, 1999;
Pont et al., 2006, 2007; Roset et al., 2007; Logez &
Pont, 2011) to disentangle the effects of impairment
from natural variability (Oberdorff et al., 1995, 2002;
Hughes et al., 1998; Stoddard et al., 2008). Nevertheless, the establishment of reference conditions remains
essential to develop and use accurate bioassessment
tools.
The historical approach was used by several authors
to define the reference conditions from past evidence
of system functioning (Swetnam et al., 1999; Muxika
et al., 2007). In contrast to this palaeoecological
approach, for most ecosystem types, reference conditions are very often defined in a more pragmatic
manner (Stoddard et al., 2006; Hering et al., this
issue), and very often the minimally disturbed conditions that are observed in a region are used (Logez &
Pont, 2011). All these approaches are based on the
assumption of relatively stable biotic communities,
so that the observed deviations consistently reflect
the effects of human disturbances. Comparing the
observed biota with reference values of a differently
functioning river, however, would lead to inaccurate
assessment due to inconsistencies in reference condition baselines. Nevertheless, with the projected modifications of climatic conditions, the assumption of
stable and consistent reference conditions is less likely
to be met. Temperature and precipitation are two
major components of hydrosystem functioning (Allan
& Castillo, 2007) which are strongly affected by
climatic changes (Webb & Nobilis, 1995; Webb,
1996; Webb & Nobilis, 2007; Albelk & Albek, 2009).
The consequences of global change, and in particular of temperature modifications, for riverine fish
assemblages are mostly studied by addressing fish
species distributions rather than assemblage structure.
Projected distributions using various gas emission
scenarios (IPCC, 2007) highlight a shift in fish species
distributions following the predicted changes in
climatic conditions (Buisson et al., 2008; Buisson &
123
Grenouillet, 2009; Grenouillet et al., 2011). Following

the theory of habitat filtering, local assemblages are
composed of species selected on their traits by the
environmental conditions (Keddy, 1992). Among the
environmental drivers, temperature was shown to be
the major determinant of the functional structure of
European fish assemblages (Logez et al., 2012a).
Therefore, global warming will likely affect both the
composition and the functional structure of fish
assemblages.
Functional traits which group species having
a common biological or ecological characteristic into
a single variable, e.g. species intolerant of habitat
degradation, are now commonly used to develop
bioassessment tools based on fish assemblages (Oberdorff et al., 2002; Pont et al., 2006, 2007, 2009;
Logez & Pont, 2011). Ecological theory suggests that
these traits are directly or indirectly related to
ecosystem functioning (Lavorel & Garnier, 2002),
because of either their response to or their effect on
the ecosystem. Functional traits are assumed to react
to human pressure in a consistent way across a wide
range of streams and over a very broad spatial extent
(Pont et al., 2006, 2007). While patterns derived from
assemblage compositions are controlled by species
distribution areas and thus by biogeographical drivers
(Hoeinghaus et al., 2007), patterns derived from
functional traits are mostly related to the environmental conditions present (e.g. Lamouroux et al., 2002;
Goldstein & Meador, 2004; Statzner et al., 2004;
Bonada et al., 2007; Ibanez et al., 2009; Logez et al.,
2010). In addition to the projected shift in species
distribution, traitenvironment relationships suggest
that fish assemblage structures will change to cope
with the new climatic conditions. It is hypothesised,
however, that functional traits will respond differently
to global change compared with species composition.
Two major approaches are used to control the
environmental variability in metrics. The type-specific
approach groups sites based on their environmental
similarities, whereas with the site-specific approach
the assemblage functional structure is directly predicted from environmental variables (Roset et al.,
2007). Some recent bioassessment methods use this
latter predictive approach to estimate the theoretical
value of a metric that should have been observed in the
absence of human pressures (Pont et al., 2006, 2007;
Hawkins et al., 2010; Logez & Pont, 2011). For
instance, the European Fish Index (EFI), a multimetric
419
Fig. 1 Locations of the 1,548 sampling sites in the 13

ecoregions adapted from Illies (1978). Due to the low number
of sites occurring in some regions, we regrouped some
ecoregions into larger geographical areas: the Alps and Pyrenees
in Alps (ALPS); the Hungarian lowlands, Eastern plains and
Pontic province in the Eastern region (EAST); and the
Fennoscandian Shield and Borealic uplands in the Nordic region
(NOR). To take into account the specificity of the Mediterranean
areas (Gasith & Resh, 1999), we defined a Mediterranean region

(MED, coloured in dark grey) as Mediterranity level 1 of
Segurado et al.s (2008) classification. The former IberoMacaronesian (IBE) and Italy, Corsica and Malta ecoregions
(ITA) were thus divided into two distinct areas. The Baltic
province (BAL), the Carpathians (CAR), Great Britain (ENG),
Central plains (C.P), Central highlands (C.H), Western plains
(W.P) and Western highlands (W.H) remain unchanged
index designed for European streams and based on ten

fish functional traits, used statistical models to predict
the expected metric values, in the absence of human
pressures, from the environmental conditions observed
at a given site (Pont et al., 2006, 2007; Bady et al.,
2009). The impairment level of a site, i.e. its deviation
from the reference conditions, is quantified by comparing the field-observed values with the predicted
theoretical values (Bailey et al., 1998). The scores
associated with each functional trait are thus independent of the environmental conditions and comparable
among stream reaches. The predictive approach

explicitly takes into account the climatic conditions
in modelling the reference baselines for functional
traits, because climatic factors (e.g. mean air temperature in July) are integrated as predictors into the
statistical models (Pont et al., 2006, 2007; Logez &
Pont, 2011). This makes it possible to test whether
climate change will limit the representativeness of
certain functional traits in European fish assemblages,
which could have important consequences for the use
of bioassessment tools based on such traits.
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420
Table 1 Description of the eight functional traits considered

Trait
Categories
Richness
Local species richness (RICH)
Tolerance to oxygen
Intolerant (O2INTOL): species

requiring more than 6 mg oxygen
per litre
Tolerance to habitat
degradation
Intolerant (HINTOL): species

intolerant to hydromorphological
degradations
Affinity for flow

velocity (habitat)
Rheophilic (RH): species preferring to

live in high-flow conditions
Eurytopic (EURY): species with broad
tolerance to flow conditions
Spawning habitat
(RHPAR) species preferring to spawn

in running waters
(EUPAR) species without clear
spawning preferences
Reproduction
Lithophilic (LITH): species spawning

exclusively on gravel, rocks, stones,
rubbles or pebbles and with
photophobic hatchlings
Table 2 Description of the four gas emission scenarios

Scenario
Description
A1F1
Rapid economic growth, global population peaks in

mid-century and more efficient technologies with
intensive consumption of fossil energy
A2
Heterogeneous world, high population growth, slow

economic growth and slow technological changes
B1
Same global population as A1F1 but convergent

world with rapid changes in economic structure
toward a service and information economy
B2
Intermediate population and economic growth,

emphasising local solutions to economic, social
and environmental sustainability
The aims of this study are to assess the effect of

climate change on eight functional traits widely used
in bioindication by: (1) assessing the relative influence
of environmental drivers (including climatic drivers)
on the field variability of functional traits, (2)
estimating the changes in functional traits in assemblages that are not or only slightly impacted, (3)
testing whether the trait sensitivity to climate change
differs among ecoregions and (4) studying the
potential shift in reference condition baselines, defined
as the deviation between the observed and theoretical
functional trait values.
123
The EFI? database integrating both natural environmental and anthropogenic factors and the data from
the UK Met Office Hadley Centre for Climate
Prediction and Research allowed us to test the effects
of climate change on riverine fish assemblage for
different scenarios at the European scale.
Materials and methods

We used the same data set as Logez et al. (2012b). All
the data used, except the forecasted climate data, were
gathered during the European EFI? project (http://
efi-plus.boku.ac.at/downloads/EFI?%200044096%
20Deliverable%20D1_1-1_3.pdf). Data were gathered
from national fish surveys conducted by several laboratories and governmental environmental agencies
(19742007, 95 % after 1990). The sites were sampled
using electrofishing methods either by wading or by
boat, depending on the stream depth. A total of 88
species were sampled, and all fish caught were identified to species level. To homogenise the sampling
effort among regions, only fish collected during the
first pass were considered.
Site selection
Sites were selected as being not impacted or only
slightly impacted by anthropogenic activities (Pont
et al., 2005) based on objective criteria (Stoddard
et al., 2008), because human alterations can modify
local assemblage composition and structure. This
selection is of major importance because all outcomes
will rely on the accuracy of the estimated trait
environment relationships. The sites chosen had good
water quality, no or few modifications of the river
cross-section, river channel and water flow, no
impoundment, no or few alterations of the river banks
and bottom habitat and no major alteration to the
rivers connectivity (Logez & Pont, 2011; Logez
et al., 2012b).
To limit the spatial autocorrelation among sites,
which is a requirement for the application of generalised linear models (GLMs; McCullagh & Nelder,
1989; Faraway, 2006), a 0.2-decimal degree latitude
and longitude grid (184396 km2) was defined and
only one site per cell was randomly selected. A total of
1,548 sites were used (Fig. 1).
421
Table 3 Overview of the current environmental conditions:

upstream drainage area (UDA), mean air temperature in July
(TJUL), mean thermal amplitude between July and January
Dataset
Scenario
(TDIF) and stream power (STP) and of the deviations between

current and projected climatic conditions for scenarios A1F1,
A2, B1 and B2 and periods 20202030 and 20502060
UDA
TJUL
TDIF
STP
Current, mean (SD)
4.50 (1.80)
18.09 (2.16)
16.94 (4.30)
11.25 (1.54)
Range
0.0011.51
11.3025.10
8.4028.80
3.9517.43
1.74 (0.38)
-0.13 (0.71)
-0.15 (0.21)
A2
1.68 (0.38)
-0.16 (0.69)
-0.15 (0.21)
B1
1.71 (0.41)
-0.29 (0.65)
-0.15 (0.22)
20202030
A1F1
20502060
B2
1.73 (0.41)
-0.30 (0.66)
-0.13 (0.20)
A1F1
3.75 (0.93)
1.23 (1.14)
-0.25 (0.25)
A2
3.11 (0.75)
1.03 (0.94)
-0.22 (0.23)
B1
2.59 (0.65)
0.66 (0.71)
-0.18 (0.22)
B2
2.66 (0.63)
0.66 (0.73)
-0.16 (0.19)
associated with each environmental variable (alone or with

its quadratic term) expressed as a percentage of the total
independent effect I
Table 4 Hierarchical partitioning of the reduction (R) of

deviance owing to the environmental factors to assess the
relative effect of each factor on functional traits: independent
(I), joint contribution (J) and the independent effect (%Ii)
Trait
Total partition
R
RICH
O2INTOL
HINTOL
EURY
%Ii
I
|I/J|
UDA2
STP2
STP
TJUL2
TDIF
-3,096.8
-2,289.8
-807.1
2.8
62.4
12
18.6
-361.4
-320.5
-40.9
7.8
25.2
44.4
30.4
-419.9
-1,954.4
-349.9
-1,601.8
-70
-352.7
5
4.5
41.5
67.6
4.7
3.1
27
17.2
28.4
10.5
RH
-1,115.5
-882.1
-233.4
3.8
55.9
3.4
15.8
24.9
EUPAR
-1,995.8
-1,455.3
-540.5
2.7
57.9
7.8
12.9
21.4
RHPAR
-861.9
-613.1
-248.9
2.5
64.3
5.8
8.1
21.8
LITH
-673.7
-528.3
-145.4
3.6
72.7
12.8
14.6
The environmental variables: temperature in July (TJUL the sum of TJUL and its quadratic term), thermal amplitude between July
and January (TDIF), stream power (STP or STP2 the sum of STP and its quadratic term) and the upstream drainage area (UDA2 the
sum of UDA and its quadratic term)
Table 5 Average differences between the theoretical trait values predicted for the current and projected climatic conditions, for the
four scenarios and the two periods
Trait
20202030
A1F1
RICH
20502060
A2
B1
B2
A1F1
A2
B1
B2
0.0959
0.1046
0.0708
0.0721
-0.3264
-0.0912
0.0108
-0.0039
O2INTOL
-0.3844
-0.3728
-0.3889
-0.3932
-0.8069
-0.6618
-0.5513
-0.5685
HINTOL
-0.3224
-0.311
-0.3281
-0.3307
-0.75
-0.5954
-0.4844
-0.4997
0.3669
0.3435
0.3483
-0.823
-0.6019
-0.4597
-0.4794
EURY
RH
0.2798
-0.271
0.2776
0.2733
-0.2575
-0.2841
EUPAR
0.0458
0.05
RHPAR
0.0114
0.0131
LITH
0.0521
0.0539
0.0339
-0.002
0.0432
0.277
-0.2863
0.3495
0.0348
-0.1038
-0.0098
0.0296
0.0225
0.0001
-0.0987
-0.0216
0.0028
0.001
0.0461
-0.0873
-0.005
0.0275
0.0272
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422
Summary of the four scenarios, period 20202030
10
15
20
25
10
10
15
20
15
Metric theoretical values
10
15
20
25
10
15
20
10
15
Metric projected values

Fig. 2 Relationships between predicted trait values for the
current (y-axis) and projected climatic conditions for
20202030 (average of the four scenarios) (x-axis): a RICH,
b O2INTOL, c HINTOL, d EURY, e RH, f EUPAR, g RHPAR,
h LITH (Table 1). Grey lines represent the y = x curve, and

dashed lines represent a general trend (Loess regression curve,
Hastie et al., 2009)
Functional traits
project. We considered seven traits that are widely

used to assess the ecological status of streams based on
fish assemblages, together with local species richness
(Oberdorff et al., 2002; Pont et al., 2006, 2007, 2009;
Logez & Pont, 2011). These traits consider tolerance
A previous classification of biological and ecological

traits of European fish species (Noble et al., 2007) was
revised and completed during the European EFI?
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423
Scenario B1, period 20502060
10
15
20
25
10
0
10
15
20
d
15
10
15
20
25
10
15
20
10
15

current (y-axis) and projected climatic conditions for 20502060
under the B1 scenario (x-axis): a RICH, b O2INTOL, c HINTOL,
d EURY, e RH, f EUPAR, g RHPAR, h LITH (Table 1). Grey

lines represent the y = x curve, and dashed lines represent a
general trend (Loess regression curve; Hastie et al., 2009)
to dissolved oxygen concentration, tolerance to habitat

degradation, affinity to flow velocity, spawning habitat
and type of reproduction (Table 1); e.g., in the case of
affinity to flow velocity, the affinity or lack of affinity
of the 88 species was evaluated, and each species was
assigned to one of the different states (e.g. species are
either rheophilic or eurytopic) corresponding to

different species attributes (Logez et al., 2012a).
Therefore, traits did not overlap and provided specific
information on the functional structure of fish assemblages; e.g. eurytopic species do not include rheophilic
species (RH). The eight metrics were composed of the
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424
Scenario A1F1, period 20502060
10
15
20
25
10
15
20
10
0
10
15
10
15
20
d
15
10
15
20
25

current (y-axis) and projected climatic conditions for 20502060
under the A1F1 scenario (x-axis): a RICH, b O2INTOL,
c HINTOL, d EURY, e RH, f EUPAR, g RHPAR, h LITH
(Table 1). Grey lines represent the y = x curve and dashed lines
represent a general trend (Loess regression curve; Hastie et al.,
2009)
number of species at a site that share a given trait, e.g.

the number of RH.
traits: upstream drainage area (UDA, km2), stream

power (STP, watt m-1), mean air temperature in July
(TJUL, C) as a proxy for water temperature (Allan &
Castillo, 2007) and the thermal amplitude between
July and January (TDIF, C). UDA reflects habitat
diversity and is a descriptor of the stream reach
position along the river network (Matthews, 1998;
Environmental factors
Four environmental factors were used (Logez et al.,
2012b) to explain the field variability of the eight
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425
Table 6 Summary of average standardized deviation between observed and projected theoretical trait values (mean and SD) for
20202030 (see Materials and methods) under the four scenarios
Trait
A1F1
A2
B1
B2
RICH
-0.031 (1.064)
-0.033 (1.062)
-0.022 (1.068)
-0.023 (1.068)
O2INTOL
0.325 (1.034)
0.315 (1.033)
0.329 (1.035)
0.333 (1.034)
HINTOL
0.235 (1.046)
0.227 (1.043)
0.240 (1.045)
0.242 (1.045)
-0.178 (1.031)
-0.176 (1.031)
-0.174 (1.030)
-0.176 (1.031)
EURY
RH
0.140 (1.072)
0.133 (1.069)
0.147 (1.073)
0.148 (1.073)
EUPAR
-0.028 (1.059)
-0.031 (1.057)
-0.020 (1.062)
-0.021 (1.063)
RHPAR
-0.007 (1.027)
-0.008 (1.026)
0.001 (1.029)
0.000 (1.029)
LITH
-0.032 (1.021)
-0.033 (1.019)
-0.026 (1.023)
-0.028 (1.021)
Oberdorff et al., 2001). STP is the rate of potential

energy expenditure over a reach or STP per unit of
stream length (Gordon et al., 2004) and reflects the
hydraulics at reach scale and integrates annual
precipitations (see Logez et al., 2012b for further
details). Temperature is a very well-known environmental driver that influences both species distribution
and functional traits (e.g. Somero, 1997; Logez et al.,
2010, 2012a, b). UDA and STP were log-transformed
due to the skewness of their distributions.
To assess the effect of global change, we used four
different climate projections developed by the UK
Met Office Hadley Centre for Climate Prediction and
Research (Mitchell et al., 2004; Mitchell & Jones,
2005). These projections were averaged for the
20202030 (referred to as 2020) and 20502060
(referred to as 2050) periods. These projections are
based on four different socio-economic scenarios
proposed by the Intergovernmental Panel on Climate
Change (Nakicenovic & Swart, 2000), used in its
fourth and latest assessment report (IPCC, 2007):
A1FI, A2, B1 and B2 (Table 2). Projections were
derived and averaged from three global circulation
models (GCM): HadCM3 (Mitchell et al., 1998;
Gordon et al., 2000), CGCM2 (Flato & Boer, 2001)
and CSIRO-Mk2 (Hirst, 1999; Hirst et al., 2000). All
climatic data were extracted from the TYN SC 1.06
data set (Mitchell et al., 2004) with 100 9 100 grid
resolution (127275 km2).
Ecoregions
To test whether the response to climate change could
differ among European regions and size of streams, we
used Illies ecoregions (Illies, 1978). Some regions

were grouped into larger geographic areas because of
their very low representations in the data sets: the
Fennoscandian Shield and Borealic uplands into the
Nordic region, the Alps and Pyrenees into Alps,
and the Eastern plains, Hungarian lowlands and Pontic
province into the Eastern region. To take into
account the specificity of the Mediterranean areas
(Gasith & Resh, 1999), we defined a Mediterranean
region as Mediterranity level 1 of Segurado et al.s
(2008) classification.
Statistical analyses
Due to the nature of the metrics (count data), log-linear
models (McCullagh & Nelder, 1989; Faraway, 2006)
were used to relate metric variations to environmental
conditions. Log-linear models use a non-normal
distribution for the model errors, and dependent
variables are linearly related to predictors through a
link function (the logarithm function for the Poisson
distribution; McCullagh & Nelder, 1989; Cameron &
Trivedi, 1998). Therefore,
the models are formulated
P
as: log(Y) = a ?
biXi, where Y is the dependent
variable (i.e. each metric), a is the intercept, and bi is
the ith parameter associated with environmental
variable Xi. The coefficients are estimated by maximising the likelihood (McCullagh & Nelder, 1989;
Faraway, 2006) rather than by ordinary least squares
as for linear models (Kutner et al., 2005; Montgomery
et al., 2006).
Environmental variables and their quadratic terms
(except for TDIF) were integrated into the models
to allow for nonlinear relationships, and the most
123
426
0.3
0.3
0.0
0.0
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0.0
0.1
0.1
0.2
0.2
0.3
0.3
0.4
0.5
0.0
0.0
0.0
0.1
0.2
0.2
0.2
0.4
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0.4
0.4
0.5
0.6
0.0
0.6
0.8
0.4
0.4
Density
0.5
0.4
0.5
Summary of the four scenarios, period 20202030
Standardized deviation between observed and theoretical metric values

Fig. 5 Densities (kernel estimation; Hastie et al., 2009) of the
functional trait scores on the current environmental conditions
(in grey) and for 20202030 (average of the four scenarios)
(dashed lines): a RICH, b O2INTOL, c HINTOL, d EURY,

e RH, f EUPAR, g RHPAR, h LITH (Table 1)
relevant variables were selected with a stepwise

procedure based on Akaikes information criterion
(AIC) (Logez & Pont, 2011). The complete models
were formulated as: log(metric) * UDA ? UDA2 ?
STP ? STP2 ? TJUL ? TJUL2 ? TDIF.
The relative effect of each environmental variable

selected on a given metric was assessed by the
hierarchical partitioning method (Chevan & Sutherland, 1991; Mac Nally, 2000). This method estimates
the independent and joint contributions of each
123
427
0.5
0.4
0.3
0.1
0.0
0.1
0.0
4
0.4
0.0
0.0
0.1
0.1
0.2
0.2
0.3
0.3
0.4
0.5
0.0
0.0
0.0
0.1
0.2
0.2
0.2
0.4
0.3
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0.4
0.6
0.5
0.2
0.3
0.2
0.3
0.2
0.1
0.0
0.6
0.8
Density
0.4
0.5
0.4
0.5
Scenario B1, period 20502060

(in grey) and for 20502060 under the B1 scenario (dashed
lines): a RICH, b O2INTOL, c HINTOL, d EURY, e RH,

f EUPAR, g RHPAR, h LITH (Table 1)
explanatory variable to the models goodness of fit

by jointly considering all the possible models and
using the theorem of hierarchies. The reduction of
deviance was used as a measure of the models
goodness of fit (Pont et al., 2005; Logez et al., 2012b).
Multicollinearity between environmental factors was

assessed by computing the general variance inflation
factor (GVIF) (Fox, 2002).
The shift in functional traits was assessed by
comparing the trait values predicted for the current
123
428
0.3
0.2
0.2
0.1
0.1
0.0
0.0
4
h
0.4
0.0
0.0
0.1
0.1
0.2
0.2
0.3
0.3
0.4
0.5
0.0
0.0
0.0
0.1
0.2
0.2
0.2
0.4
0.3
0.4
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0.2
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0.0
0.6
0.8
Density
0.5
0.5
0.4
0.5
Scenario A1F1, period 20502060

(in grey) and for 20502060 under the A1F1 scenario (dashed
lines): a RICH, b O2INTOL, c HINTOL, d EURY, e RH,

f EUPAR, g RHPAR, h LITH (Table 1)
environmental conditions and for the projected climatic conditions for the two periods and for the four
gas emission scenarios. The other physical components were assumed to be constant. If climate change
does not have any effect on the functional fish
structure, the predictions should be distributed around

the y = x curve.
The potential effects of climate change on reference
condition baselines were assessed by computing the
standardised deviations between present observed
123
429
Table 7 Summary of average standardized deviation between observed and projected theoretical trait values (mean and SD) for
20502060 (see Materials and methods) under the four scenarios
Trait
A1F1
A2
B1
B2
RICH
0.112 (1.170)
0.032 (1.127)
-0.002 (1.104)
0.003 (1.108)
O2INTOL
0.683 (1.063)
0.560 (1.051)
0.466 (1.043)
0.481 (1.044)
HINTOL
0.548 (1.088)
0.435 (1.068)
0.354 (1.057)
0.365 (1.058)
-0.223 (1.075)
-0.234 (1.060)
-0.219 (1.048)
-0.222 (1.049)
EURY
RH
0.424 (1.167)
0.310 (1.131)
0.237 (1.107)
0.247 (1.111)
EUPAR
0.072 (1.165)
0.009 (1.126)
-0.017 (1.103)
-0.013 (1.108)
RHPAR
0.063 (1.064)
0.014 (1.045)
-0.002 (1.039)
-0.001 (1.039)
LITH
0.056 (1.068)
0.004 (1.046)
-0.016 (1.037)
-0.016 (1.036)
Table 8 Average deviation of trait scores (from the four scenarios) between projected climatic conditions for 20202030 and current
climatic conditions for each ecoregion (Fig. 1)
Ecoregion
RICH
O2INTOL
HINTOL
EURY
RH
EUPAR
RHPAR
LITH
ALPS
-0.220
0.204
0.066
-0.212
-0.097
-0.177
-0.152
-0.192
BAL
-0.165
0.360
0.215
-0.356
0.057
-0.216
-0.034
-0.108
C.H
-0.194
0.293
0.148
-0.200
-0.033
-0.178
-0.104
-0.147
C.P
-0.117
0.413
0.270
-0.350
0.134
-0.142
-0.036
-0.086
CAR
-0.330
0.431
0.230
-0.361
-0.053
-0.350
-0.180
-0.221
ENG
-0.137
0.059
-0.014
-0.104
-0.100
-0.096
-0.083
-0.120
0.345
0.664
0.581
-0.160
0.600
0.404
0.188
0.154
EAST
IBE
-0.027
0.266
0.210
-0.131
0.128
-0.026
-0.002
-0.009
ITA
0.275
0.462
0.438
0.032
0.430
0.216
0.181
0.205
MED
0.260
0.334
0.344
0.033
0.370
0.205
0.177
0.203
NOR
-0.410
0.029
-0.101
-0.337
-0.309
-0.456
-0.185
-0.244
W.H
-0.088
0.309
0.202
-0.139
0.070
-0.072
-0.041
-0.072
W.P
0.036
0.370
0.298
-0.170
0.228
0.028
0.041
0.014
values and theoretical (predicted) trait values in the

absence of pressure (Pont et al., 2006), also called trait
scores, for the different climate scenarios. This
computation tests how an assemblage structure that
is currently observed in minimally disturbed conditions (i.e. close to the reference condition) would be
considered in the future. For a given scenario, if the
future climatic conditions would lead to a lesser
representation of a given metric, the deviation
between observed and theoretical values increases
and thus the difference between the deviations computed for the future and current climate is positive. For
example, if in a given site the observed RICH was 2
and the theoretical value was 3, then the deviation
between the observed and theoretical values is -1, for
the current climatic conditions. If climate change

decreased the theoretical value to 0, then the deviation
between observed and theoretical values equals 2 and
the difference between the two deviations, future
minus current, is 3 (i.e. 2 minus -1). On the other
hand, if the climate change led to a better representation of the metric in fish assemblages then a negative
difference would be observed between the deviations
computed for the future and current climatic conditions. In summary, a positive difference means that the
metric is less represented in the future assemblage,
whereas a negative difference means a higher value of
the given metric in undisturbed assemblages.
We used Wilcoxon tests to compare the average
trait deviations computed for each period and each
123
430
Table 9 Average deviation of trait scores (from the four scenarios) between projected climatic conditions for 20502060 and current
climatic conditions for each ecoregion (Fig. 1)
Ecoregion
RICH
O2INTOL
ALPS
-0.290
0.385
HINTOL
EURY
RH
EUPAR
RHPAR
LITH
0.180
-0.319
-0.062
-0.245
-0.226
-0.266
BAL
-0.286
0.666
0.432
-0.678
0.169
-0.385
-0.079
-0.176
C.H
-0.232
0.462
0.264
-0.275
0.026
-0.225
-0.151
-0.185
C.P
-0.187
0.554
0.371
-0.517
0.186
-0.228
-0.085
-0.128
CAR
-0.310
0.905
0.620
-0.542
0.243
-0.351
-0.188
-0.216
ENG
-0.264
0.072
-0.039
-0.196
-0.189
-0.195
-0.176
-0.214
EAST
0.706
1.021
0.933
-0.030
1.021
0.764
0.354
0.345
IBE
0.118
0.604
0.519
-0.164
0.417
0.077
0.062
0.078
ITA
0.477
0.648
0.631
0.119
0.646
0.357
0.327
0.363
MED
0.720
0.634
0.680
0.330
0.784
0.517
0.480
0.555
NOR
-0.714
0.033
-0.174
-0.590
-0.526
-0.800
-0.359
-0.412
W.H
-0.099
0.473
0.326
-0.195
0.147
-0.088
-0.061
-0.086
W.P
0.080
0.548
0.454
-0.236
0.366
0.050
0.042
0.034
scenario with the average deviations computed for the

current climatic conditions. Due to the multiple
comparisons testing the same null hypothesis (Shaffer,
1995), as the two samples come from populations in
the same location (Sokal & Rohlf, 1995), the P-values
were adjusted using the method proposed by Benjamini and Yekutieli (2001) and Dudoit & van der Laan
(2008). This method controls the false discovery rate
and is more powerful, less conservative and better
limits the number of hypotheses inconsistently not
rejected than classical Bonferroni correction (Logez
et al., 2010).
The potential differences in reference baseline shift
among ecoregions were tested using KruskalWallis
tests, which are the nonparametric equivalent of oneway analysis of variance (Sokal & Rohlf, 1995). The
P-values were adjusted using the same methodology
as for Wilcoxon tests.
broad amplitude but lower variance than the other

environmental parameters (Table 3).
For the 20202030 period, the change in climatic
conditions was mostly visible on TJUL with an
average increase of 1.72 C. At the same time, the
thermal amplitude was expected to decrease slightly,
suggesting that the temperature increase would be
greater in January than in July for this period. In
contrast, the increase of temperature projected for the
20502060 period would be higher in July than in
January, respectively, 3.03 and 2.13 C (average of
the four scenarios). STP was on average expected to
decrease over time. The same climate change patterns
were predicted with the four scenarios (all Spearman
q [ 0.95); only the magnitudes were different, with
scenarios A1F1 and A2 predicting the greatest temperature increases over 20502060 (Table 4).
Effect of environmental factors
Results
Data sets
The 1,548 stream reaches were mostly located in small
to medium-sized streams; 90 % of sites had UDA less
than 1,000 km2. The climatic conditions were highly
contrasted in terms of both temperatures and thermal
amplitude. TJUL ranged from 11.3 to 25.1 C, and
TDIF varied between 8.4 and 28.8 C. STP had a very
123
Among the four environmental factors considered,

UDA, TJUL and TDIF were always selected by the
stepwise procedure, as were the quadratic terms of
UDA and TJUL. STP was selected for six traits and
was associated with its quadratic term for five traits
(Table 4). All GVIF values were lower than 2.34,
suggesting that multicollinearity was a limited
phenomenon.
The hierarchical partitioning supported this result:
the ratios between independent and joint contributions
(I/J) ranged between 2.5 and 7.8. UDA had the greatest
relative independent contributions (%Ii) to almost all
traits except O2INTOL, ranging from 25.2 to 72.7 %.
Temperatures (TJUL and TDIF) were the second most
important variables, and STP always had the lowest
relative independent contributions compared with the
other environmental factors (when this variable was
selected) (Table 4).
Change in functional traits
The four scenarios provided relatively similar results
for the 20202030 period; the average deviations
between the predicted values for the current and
projected climatic conditions were consistent among
scenarios. O2INTOL, HINTOL, EURY and RH
showed the largest differences among the predicted
trait values (Table 5). O2INTOL and HINTOL predictions for 20202030 were almost always under the
y = x curve (Fig. 2), suggesting that the representation of these traits would decrease. The average
deviation for RH was also negative (Table 5), but the
deviation from the current conditions seemed to
appear for high RH values (Fig. 2). In contrast to the
three former traits, EURY values for 20202030 were
on average higher than the predictions for the current
conditions (Table 5; Fig. 2).
Compared with 20202030, the results obtained
with the four scenarios were much more contrasted
over 20502060. The magnitude of responses was
greater for the A scenarios, especially for A1F1,
than for the B scenarios, but the response pattern
was similar (Table 5). The only noticeable difference
was for RICH, which was expected to decrease on
average under the A1F1 scenarios and to remain stable
under the three other scenarios. As for 20202030, the
four metrics that deviated the most from projected and
current climatic conditions were O2INTOL, HINTOL,
EURY and RH (Table 5; Figs. 3, 4). Under A1F1, the
deviations from the y = x curve seemed to occur for
lower predicted values of RH than for the 20202030
period (Fig. 2). Otherwise, the response patterns over
the two periods are relatively similar but more
scattered for 20502060.
Changes in reference condition baselines
The four scenarios provided very close trait scores
(standardised trait values, see Materials and
431
methods section) between current and climatic

conditions projected for 20202030 (Table 6), and
the patterns observed were very similar to those
observed for functional traits. O2INTOL and HINTOL
were expected to shift toward positive values on
average. The score distributions were very close for
the other traits (Fig. 5).
The patterns predicted for 20502060 seemed
amplified compared with 20202030: the magnitude
of the differences between trait scores increased.
O2INTOL and HINTOL displayed much more distinct
distributions from the current climatic conditions
(Figs. 6, 7), and the deviations among average trait
scores were more pronounced (Table 7). Not surprisingly, the metric score deviations were higher for
the A scenarios than for the B scenarios. For
instance, distributions of RH scores for the current and
20502060 climates were close under the B1 scenario
(Fig. 6), whereas these two distributions were relatively distinct under the A1F1 scenario (Fig. 7).
All KruskalWallis tests were significant (adjusted
P-values \0.001), whatever the period and the
scenario considered, suggesting that the shift in the
reference condition baselines differ among ecoregions. Because the results from all scenarios were
comparable (only the magnitude of the response
differed, see Electronic Supplementary Material), we
present only the average values from the four scenarios (Tables 8, 9). O2INTOL and HINTOL traits were
expected to decrease in almost all regions (positive
average deviation of trait scores), but the magnitudes
of the decline varied among regions. For the other
traits, the results were more contrasted among regions
(Tables 8, 9).
Discussion
The data used in this study allowed assessment of the
influence of environmental drivers on fish functional
traits and for accurate forecasting of species traits for
future decades. The projections showed that mainly
four traits will be highly affected by climate change:
species intolerant to habitat degradation (HINTOL),
species with high oxygen needs (O2INTOL), RH and
eurytopic species (EURY).
The main environmental driver of the eight traits
was the UDA, which could be explained by the type of
metrics considered in this study, as the eight traits used
123
432
in this study were expressed by species number. Local

species richness is known to vary along the longitudinal gradient (Horwitz, 1978; Vannote et al., 1980;
Hugueny, 1989; Oberdorff et al., 1993, 1995; Grenouillet et al., 2004; Ibanez et al., 2009; Logez et al.,
2010), which could explain the importance of UDA for
these metrics. Moreover, these results are consistent
with previous studies which used environmental
variables reflecting the longitudinal gradient (e.g.
distance from source) to model metrics based on
functional traits and expressed in the number of
species (Pont et al., 2006, 2007; Logez et al., 2010).
The second major environmental factor was temperature, expressed either in monthly temperature
(TJUL) or in thermal amplitude (TDIF). The climatic
changes, especially thermal changes, projected by the
four different gas emission scenarios (IPCC, 2007) are
expected to modify the functional structure of European
fish assemblages. This shift in functional structure can
be compared with the shift in distribution areas
projected using species distribution models (Xenopoulos et al., 2005; Buisson et al., 2008, 2010; Buisson &
Grenouillet, 2009; Grenouillet et al., 2011). The rise in
observed temperature (Webb & Nobilis, 1995, 2007;
Webb, 1996) and expected temperature caused by
global change (IPCC, 2007) will negatively affect coldand cool-water species such as salmonids (Isaak et al.,
2012) and modify the composition of the species
assemblage (Xenopoulos et al., 2005; Graham &
Harrod, 2009). Compared with the current situation,
suitable climatic conditions for coldwater species
presence would be greatly limited (Isaak et al., 2012),
leading to extinction of local populations.
Coldwater species are very often intolerant to
both habitat degradation and low oxygen concentration (Halliwell et al., 1999; Zaroban et al., 1999;
Holzer, 2008; Logez & Pont, 2011). Therefore, the
projected decline of the two corresponding metrics,
HINTOL and O2INTOL, must be directly linked to
the projected decline of coldwater species under
climate warming. The same tendency is observed for
the RH metric, but to a lesser extent due to the
wider range of thermal preference of RH (Logez
et al., 2012b), including both salmonid and cyprinid
species. At the opposite extreme, the eurytopic
metric (EURY) tended to be better represented in
the predicted future undisturbed fish assemblage,
due to their ability to adapt to a wider range of
environmental conditions.
123
The traits related to reproduction (RHPAR, EUPAR, LITH) were not expected to be greatly affected
by climate change. This suggests either that only some
species will be affected in the same manner by climate
change depending on their thermal preferences (Buisson et al., 2008, 2010; Hering et al., 2009; Grenouillet
et al., 2011; Logez et al., 2012b) or that the same
function will be represented by other species with
higher thermal preferences but with similar characteristics. Finally, local species richness (RICH) was also
expected to remain stable, suggesting replacement
of intolerant coldwater species with more eurytopic
species.
These results will have consequences for the
bioassessment of European streams based on fish
assemblages, especially when using most of the IBItype indices commonly used nowadays (Pont et al.,
2011). Most of these indices integrate several metrics
based on functional traits that will be impacted by
climate change. For the metrics that are expected to
be better (EURY) or slightly less (RH) represented
in assemblages, reference condition baselines would
have to be changed because they will become poorly
adapted to the future climate. Otherwise, under the
reference condition approach (Bailey et al., 1998), the
observed values would be compared with inconsistent
values, leading to misclassification of both impaired
and non-impaired sites. This redefinition of baseline
reference conditions could be partially obtained with
predictive fish indices (Pont et al., 2006, 2007)
including climatic conditions in the computation of
the predicted metric values in the absence of pressure.
For the intolerance-based metrics expected to be
highly negatively affected by climate change (HINTOL, O2INTOL), a change in the baselines of
reference conditions could be considered as long as
global warming does not excessively reduce the
diversity of these functional groups. For example, in
view of a huge disappearance of cold-intolerant
species (Buisson et al., 2008; Lassale et al., 2008;
Buisson & Grenouillet, 2009), these metrics must not
be considered in future bioassessment tools, because
the metrics must reflect the faunal specificity of the
regional fauna (Karr, 1981, 1991; Logez & Pont,
2011). Including a metric which has little or no
representation in the region limits the discrimination
between impaired and unimpaired sites (Harris &
Silveira, 1999), which could lead to limited use of the
metric based on species intolerance, the most sensitive
metric to human pressures (Fausch et al., 1984; Simon

& Lyons, 1995; Oberdorff et al., 2002; Joy & Death,
2004; Pont et al., 2006, 2007, 2009; Melcher et al.,
2007; Vehanen et al., 2010; Logez & Pont, 2011).
A final consequence would be reduced sensitivity of
fish-based indices to local anthropogenic alterations.
Depending on the ecoregion, different effects of
climate change were forecasted, suggesting that for
some ecoregions the current bioassessment tools will
more rapidly become inadequate, whereas for the
others these tools will remain consistent when used to
assess the ecology of streams. Climate change consequences would depend on the shift in climatic
conditions projected for these regions.
All these results clearly stress the need to maintain
a biomonitoring network on undisturbed sites considered as reflecting present reference conditions. The
functional changes of European fish assemblages
could be missed and the reliability of the currently
used indices may not be assessed if monitoring is not
pursued.
Acknowledgments We are grateful to all the members of the
European EFI? project (contract number 044096, http://efiplus.boku.ac.at/), which provided all the data. This paper is a
result of the WISER (Water bodies in Europe: Integrative Systems to assess Ecological status and Recovery) project funded
by the European Union under the 7th Framework Programme,
Theme 6 (Environment including Climate Change) (contract no.
226273), http://www.wiser.eu.
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Hydrobiologia (2013) 704:417436

WATER BODIES IN EUROPE

Global warming and potential shift in reference conditions:

Guest editors: C. K. Feld, A. Borja, L. Carvalho & D. Hering /

Electronic supplementary material The online version of

principles of the index of biotic integrity (IBI)

Hydrobiologia (2013) 704:417436

Grenouillet, 2009; Grenouillet et al., 2011). Following

principles of the index of biotic integrity (IBI)

Hydrobiologia (2013) 704:417436

Grenouillet, 2009; Grenouillet et al., 2011). Following

Hydrobiologia (2013) 704:417436

Fig. 1 Locations of the 1,548 sampling sites in the 13

areas (Gasith & Resh, 1999), we defined a Mediterranean region

index designed for European streams and based on ten

among stream reaches. The predictive approach

Hydrobiologia (2013) 704:417436

Table 1 Description of the eight functional traits considered

Local species richness (RICH)

Intolerant (O2INTOL): species

Intolerant (HINTOL): species

Affinity for flow

Rheophilic (RH): species preferring to

(RHPAR) species preferring to spawn

Lithophilic (LITH): species spawning

Table 2 Description of the four gas emission scenarios

Rapid economic growth, global population peaks in

Heterogeneous world, high population growth, slow

Same global population as A1F1 but convergent

Intermediate population and economic growth,

The aims of this study are to assess the effect of

Materials and methods

Hydrobiologia (2013) 704:417436

Table 3 Overview of the current environmental conditions:

(TDIF) and stream power (STP) and of the deviations between

Current, mean (SD)

associated with each environmental variable (alone or with

Table 4 Hierarchical partitioning of the reduction (R) of

Hydrobiologia (2013) 704:417436

Summary of the four scenarios, period 20202030

Metric theoretical values

Metric projected values

h LITH (Table 1). Grey lines represent the y = x curve, and

project. We considered seven traits that are widely

A previous classification of biological and ecological

Hydrobiologia (2013) 704:417436

Scenario B1, period 20502060

Metric projected values

d EURY, e RH, f EUPAR, g RHPAR, h LITH (Table 1). Grey

to dissolved oxygen concentration, tolerance to habitat

either rheophilic or eurytopic) corresponding to

Hydrobiologia (2013) 704:417436

Scenario A1F1, period 20502060

Metric projected values

number of species at a site that share a given trait, e.g.

traits: upstream drainage area (UDA, km2), stream

Hydrobiologia (2013) 704:417436

Oberdorff et al., 2001). STP is the rate of potential

used Illies ecoregions (Illies, 1978). Some regions

Hydrobiologia (2013) 704:417436

Summary of the four scenarios, period 20202030

Standardized deviation between observed and theoretical metric values

(dashed lines): a RICH, b O2INTOL, c HINTOL, d EURY,

relevant variables were selected with a stepwise