Freshwater Biology (1999) 41, 809±820

Predicting the structure and diversity of young-of-the-

year fish assemblages in large rivers
R. E. GOZLAN*y , S. MASTRORILLO y , G. H.COPP* AND S. LEK y
*Landscape & Ecology Research Group, Department of Environmental Sciences, University of Hertfordshire, College Lane,
Hatfield, Herts. AL10 9AB, U.K.
y
CESAC, UMR C5576, CNRS-University of Paul Sabatier, 118 route de Narbonne, 31062 Toulouse Cedex 04, France

SUMMARY

1. Interactions between environmental variables and 0+ fish assemblages in the upper

River Garonne (France) were quantified during late August 1995.
2. The abundance and diversity of the fish assemblages in floodplain channels were
modelled using Artificial Neural Network (ANN) analysis and nine variables: the
abundance of the six dominant species, fish specific richness, overall abundance of 0+ fish
and the Shannon index of diversity. Multiple regression analysis was also used to assess
ANN performance.
3. Using 596 samples, correlation coefficients (r adjusted) between observed and
estimated values of the nine dependent parameters were all highly significant (P < 0.01).
Expected values from the tested data were significantly related to the observed values. The
correlation coefficient between observed and estimated values (r) varied from 0.70 to 0.85.
4. The ANN provided a high quality prediction, despite the complex nature of the
relationship between microhabitat composition and fish abundance.
5. Garson's algorithm was used to provide the explanatory power needed in ecology
when using black-box models. Parameters contained in the models (i.e. weighting) were
used to determine the relative contributions of explanatory variables and thus to ascertain
the structure of fish communities.

Keywords: artificial neural network, modelling, flood plains, point abundance sampling,
River Garonne, 0+ fish

Introduction assess environmental influences on recruitment using

Predicting the abundance of young-of-the-year (0+)
fishes is essential in determining year-class strength
(Hatcher, Nester & Muth, 1991) as well as detecting
environmental influences on recruitment (Kope &
Botsford, 1988). Moreover, interactions between the
organism and its environment also influence the
taxonomic and numerical composition of assem-
blages. Modelling and simulation are powerful tools
that have been used to describe and predict ecosystem
structure. Ricker (1975) used correlation analysis to
Correspondence: R. E. Gozlan, Landscape & Ecology Research
Group, Department of Environmental Sciences, University of
Hertfordshire, College Lane, Hatfield, Herts. AL10 9AB, U.K.
E-mail: r.gozlan@herts.ac.uk
abundance data (Kope & Botsford, 1988). Multiple
regression (Binns & Iserman, 1979), as well as
canonical correspondence analysis (ter Braak &
Verdonschot, 1995), have frequently been used as
qualitative methods to elucidate the relationships
between species assemblages and their habitat pre-
ferences. The aim of our study was to predict the
abundance of six fish species in the River Garonne
using a non-linear multivariate method, namely a
neural network trained by backpropagation (Hornik,
Stinchcomb & White, 1989; Chen, Billings & Grant,
1990; Specht, 1991). Various scientific fields (physics,
geology, chemistry, biology) have already used non-
linear modelling methods successfully. Similarities
between neural networks and ecosystems have been
highlighted (Colasanti, 1991) and thus neural net-

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810 R. E. Gozlan et al.
works offer great potential for ecological modelling
and biodiversity research (Edwards & Morse, 1995).
In recent years, the questions put to scientists by
river management agencies reflect an integrated
approach to river systems, which requires quantita-
tive estimators of abundance. Such estimators are
inherent in the distribution and density of 0+ fish as
functional describers to define the ecological function
of floodplain ecosystems (Copp, 1989; Gozlan et al.,
1998). Much ecological research has focused on the
strength of the relationship between 0+ fishes and
microhabitat characteristics (Lightfoot & Jones, 1979;
Souchon et al., 1989; Copp, 1992; Baras, Nimbada &
Philippart, 1995; Garner, 1996; Gozlan et al., 1998). The
end of the summer period has been identified (Copp,
1989) as a particularly suitable phase during which
the intrinsic ecological needs of 0+ fishes can be
evaluated, as this relatively `stable' interval separates
the phase of high mortality characteristic of the
preceding embryonic/larval periods and that of the
subsequent winter season.
In order to highlight the fish assemblages and their
link with habitat features (Oberdorff & Porcher, 1992),
it is essential to be able to describe, quantify and
model this relationship. The habitat variables intro-
duced into the modelling procedure need to be chosen
carefully, with a full understanding of their ecological
significance, as this will provide greater precision for
the prediction. By sampling a large number of
microhabitats at the end of the summer, an instanta-
neous picture of the structure, abundance and
distribution of 0+ fish assemblages in floodplain
ecosystems can be obtained. This abundance is the
result of their reproductive success, their mortality
since hatching (natural mortality rate), habitat avail-
ability and of trophic competition (intra and inter
specific). Because the abundance of 0+ fishes commu-
nities at the end of the summer changes over the
years, we do not try to predict the recruitment of 0+
fishes quantitatively but rather their dispersion in
terms of different microhabitat characteristics. There-
fore, variables influencing the distribution of 0+ fishes
must be determined with the aim of highlighting the
stochastic or deterministic features of those assem-
blages.
Artificial neural networks (ANN) are more power-
ful than multiple linear regressions in achieving our
objectives because of their ability to process non-linear
relationships (Hornick, Stinchcombe & Muth, 1989;
Lek et al., 1996b), which often occur in ecology (ter
Braak & Verdonschot, 1995). It has repeatedly been
noted that ANN performance surpasses that of
multiple regression analysis (Ehrman, Clair & Bou-
chard, 1996; Scardi, 1996). For example, behaviour of
real systems cannot be effectively reproduced by
multiple linear regression models when considering
very low or high variable values (Lek et al., 1996b).
Non-linear transformations (logarithmic, power,
exponential) of independent and/or dependent vari-
ables have often been used to improve results
(Cancela Da Fonseca, 1991). Despite this, multiple
regression, results have often proved unsatisfactory,
with a low percentage of variance being explained. On
the other hand, in ecology (Goh, 1995; Lek et al.,
1996b; Scardi, 1996) even the simplest ANN with only
one hidden layer can effectively model non-linear
systems. ANNs using the error back-propagation
procedure have recently been applied for ecological
purposes as a predictive tool (Lek, Dinopoulos &
Fabre, 1996c; Mastrorillo et al., 1997; GueÂgan, Lek &
Oberdorff, 1998), but often without explanation of the
identifying variables nor assessment of their influence
on the output. To address this shortcoming, we
propose here the use of Garson's algorithm to
evaluate the sensitivity of each variable (Garson,
1991). Graphic representation of the contribution of
environmental variables in predicting fish abundance
thus facilitates our understanding of the impact of
habitat modification on 0+ structures, in contrast with
the previous uses of ANN.
Materials and methods
Study site
The River Garonne's source is the Maladetta's glacier
(Spain), and the river disappears into `Tora's Hole' at
2050 m a.s.l (Fig. 1). It reappears in the `Val d'Aran' at
1872 m a.s.l. and flows for 575 km, discharging into
the Atlantic Ocean at its confluence with the River
Gironde. The average annual rainfall (based on data
from 1954 to 1990) is 612.7 mm, and the average daily
temperature is 5.4 °C in January, which is the coldest
month, and 21.9 °C in the warmest month, July. The
average yearly temperature is 13.3 °C (Bengen, 1992).
Compared with other French rivers (e.g. Seine and
RhoÃne), the Garonne is less impacted by industrial
pollution. However, its natural flow has been partially
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 Structure and diversity of young-of-the year fish assemblages 811

Fig. 1 Map with locations of the study sites on the upper River Garonne, between Muret and Golfech, France.

modified by the presence of several dams, thus Nilsonn, 1994). From south to north-west, topography
promoting the fragmentation of animal and plant and climate determine three principal landscape
communities within the river channel and its alluvial types: the Pyrenean mountains with a pronounced
flood plain (DeÂcamps & Naiman, 1989; Dynesius & relief, a large green hill zone of Piedmont, and the

ã 1999 Blackwell Science Ltd, Freshwater Biology, 41, 809±820

812 R. E. Gozlan et al.
valley of the River Garonne with inundation zones
and alluvial terraces. An oceanic influence predomi-
nates over the whole basin, but lessens to the south-
east, which is subjected to Mediterranean influence
with dry winds and less rainfall.
Sampling design
During August 1995, 0+ fish and seventeen environ-
mental variables were characterised at 1160 sampling
points at each of thirty-eight sites (Gozlan et al., 1998),
with one sampling point taken approximately every
10 m2 using point abundance sampling by electro-
fishing (P.A.S.E.) adapted for 0+ fishes (Copp &
Garner, 1995). The structure of the spatial variability
was investigated within a large section of the River
Garonne. As recommended by Persat, Olivier & Pont
(1994), the longitudinal and transversal aspects of this
habitat variability were taken into account by dis-
tributing our 1160 sampling points equally over three
types of channels from upstream to downstream of
Toulouse between Muret and Golfech (Fig. 1): flowing
channel (FC), partially abandoned channel (PA) and
abandoned channel (AC). Sampling points were
selected randomly in a zig-zag manner. A dinghy
was used to approach individual sampling points, the
operator plunged the activated anode of a portable
electrofishing apparatus as quickly as possible to a
depth of about 0.5 m. The anode was activated for a
period of 4±5 s, with a dip net immediately immersed
Table 1 Input and output parameters from the River Garonne (data from Gozlan et al., 1998) used in the ANN modelling. See Table 2
for fish codes
0.5 m below the anode, and both were lifted directly
out of the water. Young-of-the-year fish were pre-
served in 4% formalin. Nine semiquantitative and
seven quantitative environmental variables were
measured at each sampling point: distance from the
bank (in metres), water depth, slope of bank (depth
divided by the distance from the bank), percentage of
clay (< 0.05 mm),% silt (< 0.06 cm), % sand (0.06±
0.2 cm), % gravel (0.2±6.3 cm), % pebbles (6.4±25 cm)
and % cobbles (> 25 cm) as well as % of roots.
Macrophytes and algae were recorded as absent or
present. Water temperature was measured with a
thermometer, and water velocity was determined
semiquantitatively as in Copp (1992): absent (0 cm
s±1), weak (> 0, but < 5 cm s±1), medium (5±10 cm
s±1), strong (> 10 cm s±1).
The ANN model
From seventeen environmental variables (Table 1),
nine biological (dependant) variables were predicted
using an Artificial Neural Network (ANN): the
abundance of six dominant fish species, fish specific
richness, total number of 0+ fishes and the Shannon
index of diversity. The predictive model of fish
abundance based on habitat characteristics was con-
structed using the mathematical algorithm of back-
propagation (Rumelhart, Hinton & William, 1986;
Carpenter, 1989; Weigend, Huberman & Rumelhart,
1992) to transform the activation into a non-linear type

Input parameters Code Output parameters Code

Flowing channel (0/1) FC Abundance of Bj ABj

Partially abandoned channel (0/1) PA Abundance of Lc ALc
Abandoned channel (0/1) AC Abundance of Lg ALg
Distance from the bank (in metres) DIS Abundance of Bb ABb
Water depth (in cm) DEP Abundance of Gb AGg
Slope of bank (DEP/DIS) SLO Abundance of Rr ARr
% of clay CLA Specific richness FSR
% of silt SIL Number of 0+ fishes Ind
% of sand SAN Shannon diversity index SH
% of gravel GRA
% of pebble PEB
% of cobbles COB
Temperature (in °C) TEM
Water velocity VEL
% of roots ROO
Macrophytes (0/1) VEG
Algae (0/1) ALG

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 Structure and diversity of young-of-the year fish assemblages 813
2
Table 2 The fish codes, latin and common names, the number (n), standard length mean (SL), standard error (SE) and variance (s ) for
the standard lengths of 0+ fish captured in the River Garonne in August 1995 (from Gozlan et al., 1998)

Code Latin name Common name n Mean SL SE s2

Am Ameturus melas (Rafinesque 1820) black bullhead 1037 22.80 0.38 43.4
Bj Blicca bjoerkna (L. 1758) silver bream 572 23.87 0.25 46.8
Lc Leuciscus cephalus (L. 1758) chub 489 25.73 0.39 72.1
Gg Gobio gobio (L. 1758) gudgeon 398 23.48 0.38 52.4
Lg Lepomis gibbosus (L. 1758) pumpkinseed 253 21.24 0.44 36.4
Bb Barbus barbus (L. 1758) barbel 168 27.10 0.81 108.5
Rr Rutilus rutilus (L. 1758) roach 168 37.68 1.73 132.5
Rs Rhodeus sericeus (Bloch. 1783) bitterling 164 24.89 0.68 72.6
Aa Abramis brama (L. 1758) bleak 125 22.64 0.92 85.2
Ca Carassius auratus (L. 1758) goldfish 115 33.20 1.30 103.2
Cc Cyprinus carpio (L. 1758) carp 113 23.55 1.02 109.7
Se Scardinius erythrophthalmus (L. 1758) rudd 84 32.66 0.89 64.3
Gb Gambusia affinis (L. 1758) mosquito fish 77 16.13 0.69 34.8
Ab Alburnus alburnus (L. 1758) common bream 50 36.38 2.33 141.1
Cx Chondrostoma toxostoma (Vallot 1836) sofie 32 30.29 2.44 143.9
Ms Micropterus salmoides (L. 1758) largemouth bass 22 20.86 3.13 177.4
Ll Leuciscus leuciscus (L. 1758) dace 18 26.16 2.34 115.8
Nb Barbatula barbatula (L. 1758) stone loach 13 34.14 3.75 98.8
Pf Perca fluviatilis (L. 1758) perch 8 51.85 3.23 73.1
Tt Tinca tinca (L. 1758) tench 4 22.33 2.58 20.1
Sl Stizostedion lucioperca (L. 1758) pikeperch 1

response. In preparation for ANN analysis, null-
samples (i.e. without any fish) were eliminated. The
remaining data matrix contained 596 samples. The
analyses were conducted on an IBM compatible PC
using Matlab with the neural network toolbox. A
multilayered, feed-forward neural network with back-
propagation typically comprises three kinds of neuron
layer (Fig. 2): an input layer (I), one or several hidden
layers (H), and an output layer (O), which consists of
one or several neurons. The processing elements in
the network, called neurons or nodes, are organised in
a layered structure and are discrete in time. The user
can choose the number of neurons in the hidden layer
depending on the accuracy of the results required. In
the majority of cases, ANN with one hidden layer,
and a sufficient number of nodes, is capable of
achieving any mapping with an arbitrary degree of
accuracy (Hornik et al., 1989; Bhat & McAvoy, 1992).
However, if the network is modelled with a more
complicated structure (e.g. using a large number of
neurons), then a strong tendency to overfit the
training dataset may occur, resulting in a poor
prediction performance (Smith, 1994). In these net-
works, signals are propagated from the input layer
through the hidden layers to the output layer. All the
neurons of a given layer, except those of the last, emit
an axon to each neuron of the downstream layer. A
neuron sends signals to and from other neurons via
the network connections. During training, each signal
is given a weighting (modifiable weight) by a factor
associated with each connection between two succes-
sive neuron layers. This factor is determined by the
entry and exit datasets.
The independent variable values are entered into
the first layer of the model. Input data must be
adapted because their magnitudes differ greatly from
one variable to another. In order to standardise the
measurement scales, input variables are converted
into a reduced-centred variable using the formula:
Xc − X
Zs =
σX
Each neuron from the hidden layer and the output
layer performs a weighted summation of input data
received from every upstream layer neuron. The
summation is computed as follows:
1
a j = ∑ Xi Wji
i =1
where aj is activation of the jth current neuron, is the
output value of the ith neuron of the previous layer, is
the weight factor of the connection between the ith

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814 R. E. Gozlan et al.
Fig. 2 Illustration of the neural network structure, with
seventeen input neurons (I), the hidden layer (H) with fifteen
neurons, and one output layer (O). See Table 2 for variable
codes.
neuron of the previous layer and the jth neuron of the
current layer, Wj 1 is the bias on the jth current neuron
always equal to 1.
After computing the weighted sum, as shown in
eqn 1, the method of calculating the neuron output
according to its net input depends on the type of
activation function used in the ANN. A common
transfer function is the sigmoid function (Lek et al.,
1996c):
1
X j = f (a j ) =
1 + exp − aj
Finally it should be noted that parameters such as h
(learning coefficient) and a (momentum) can be used.
These serve to accelerate learning and prevent the
network from falling into local minima. Network
learning, performed on a representative data set, runs
until the Mean Square of Errors (MSE) is minimised
1 N
MSE = ∑ Yn − Yn )
2
N n =1
where Yn is the expected network output value
(`theoretical value'), Yn is the value estimated by the
network (output layer neuron) and N is the number of
observations.
Modelling was carried out in two steps:
First, the entire data matrix of 596 samples ´ 17
environmental variables was randomly divided into
two subsets of which the first consisted of 75% of the
data (447 records); this was used to determine the
connection weighting after 1000 iterations (which is
low in neural network modelling, Bhat & McAvoy,
1992) with the best compromise between bias and
variance (see Kohavi, 1995). This process of updating
the weightings to a correct set of values (so as to
minimise the error of the response) is known as
`training' or `learning' (Rumelhart et al., 1986; Gallant,
1993; Smith, 1994). The output neuron with the largest
outgoing signal indicated the network value that
estimated the case under study. The estimation of the
error signal differed according to the layers consid-
ered. Second, the remaining 149 records (25% of total)
were used to test the model.
To determine the response of the model to each of
the input variables separately, a sensitivity analysis of
the ANN was performed, using Garson's (1991)
algorithm to interpret neural network connection
weighting (Goh, 1995). This procedure partitions the
neural network connection weightings in order to
determine the relative importance of various input
variables.
Results were represented by histograms to illustrate
the contribution of each independent variable to
model each dependent variable. However, correspon-
dence analysis of the contribution of studied variables
was carried out on the contribution matrix to identify
the best simultaneous representation of two sets
taking into account environmental variables and
biological variables.
Results
A total of 4250 fish where caught in the 596 samples
(Fig. 1), including 3911 0+ fish (Table 2). The compo-
sition of the fish fauna of the River Garonne is typical
of temperate rivers, with only a few families repre-
sented and the community strongly dominated by
Cyprinidae (81% of the fishes collected). The six most
abundant species selected for the model represented
71% of the total numbers of fish (Table 2, Appendix
1).
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 Structure and diversity of young-of-the year fish assemblages 815
Table 3 Correlation coefficients between observed and esti- Table 4 Correlation coefficient between observed and estimated
mated values of multiple linear regression (MLR) models: values in the training set (n = 447 samples) and between
classical MLR, MLR with log(x + 1) transformations of quanti- observed and predicted values in the testing set of the ANN
tative variables and alternating least square method using a modelling procedure (n = 149 samples). All correlations sig-
cubic spline transformations of variables (Transreg procedure in nificant at > 99.9%. See Table 2 for the fish codes
SAS software). All models were significant (P < 0.05), except for
Bj in MLR (P = 0.401). See Table 2 for the fish codes Correlation coefficients

MLR-log Transreg-Spline Output parameters Code Training Testing

MLR transformed procedure
Abundance of Bj ABj 0.933 0.803
Bj 0.168 0.370 0.330 Abundance of Lc ALc 0.909 0.857
Lc 0.301 0.307 0.462 Abundance of Lg ALg 0.951 0.857
Lg 0.321 0.302 0.436 Abundance of Bb ABb 0.867 0.819
Gg 0.258 0.390 0.500 Abundance of Gb AGg 0.932 0.841
Rr 0.209 0.310 0.430 Abundance of Rr ARr 0.928 0.799
Bb 0.314 0.406 0.450 Specific richness FSR 0.834 0.736
SR 0.303 0.280 0.430 Number of 0+ fishes Ind 0.900 0.771
Ind 0.244 0.280 0.355 Shannon diversity index SH 0.920 0.709
Sh 0.290 0.250 0.430

output variable obtained using Garson's algorithm

The correlation coefficients between observed and
estimated values of multiple regression models were
low (Table 3). The correlation coefficients of multiple
linear regression calculated with and without trans-
formation of the variables were highest for pumkin-
seed Lepomis gibbosus with 0.32 and barbel Barbus
barbus with 0.40, respectively. The correlation coeffi-
cients were stronger with the alternating least square
method using a cubic spline transformation of
variables (procedure Transreg in SAS software).
Gudgeon Gobio gobio achieved the best score with
0.50 but even this is low when compared to the results
of the ANN model. Correlation coefficients obtained
for all the observed (and estimated) values in the
training set (n = 447) were greater than 0.80 (Table 4).
Residuals and estimated values in the training
procedure were completely independent (P < 0.05).
Tests of the ANN model on the remaining 149
samples revealed a significant correlation (all > 0.7
and P < 0.001) between predicted and observed
values (Table 4), with complete independence
between the residuals and predicted values
(P > 0.05). Correlations were slightly lower for spe-
cific richness and the Shannon diversity index,
probably due to their more complex nature, but still
remain very high. The majority of records were then
aligned on the diagonal of co-ordinate 1 : 1 for all
output parameters (Fig. 3), and despite the correlation
coefficients being lower than in the training set, they
still remained clearly significant.
The contribution of environmental variables to each
(Fig. 4) showed that, when considered separately,
none of the input variables contributed more than
11% to the abundance of a fish species. However, this
is not to say that the contribution of any one variable
was not important. A species' microhabitat is char-
acterised by a complex combination of different
habitat variables, therefore rendering any conclusion
based on the importance of one particular variable
inaccurate. Note, that as complex as this combination
of habitat variables can be, a group of variables will
always contribute the same percentage to the predic-
tion of abundance of a fish species within the model.
For example (Fig. .4), the spatial habitat characteristics
(distance from the bank, depth and slope) contributed
» 18% to the abundance of any species, although
substratum contributed around 33% and aquatic
vegetation around 18%. Some variations may occur,
such as in roach Rutilus rutilus for which the spatial
habitat variables contributed 23%, or the gudgeon,
where substratum contributed 37% to its abundance.
The first two factors of the correspondence analysis
accounted for 74.7% of total variation (Fig. 5). The first
factor (43.8%) mainly accounted for the abundance of
barbel and silver bream Blicca bjoerkna (Table 5),
whereas the second factor (20.9%) accounted for the
abundance of pumpkinseed. Water velocity is an
important input variable to introduce into the model-
ling of 0+ abundance (Figs 3 and 4) because it
contributes with a fairly high percentage to the
prediction of typical lotic species (barbel, gudgeon)
or typical lentic species (silver bream).

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816 R. E. Gozlan et al.
Fig. 3 Results of the ANN modelling. Scatter plot of observed values vs. predicted values (the solid line indicates the perfect fit line (for
which y = x).
Discussion
The ANN model succeeded in predicting the numer-
ical abundance of the six 0+ fish species as well as
specific richness. It revealed that the microhabitat
variables introduced into the model accurately char-
acterised some important abiotic needs of the six
species. However, only small variations occurred in
the contribution of the different variables to the
overall habitat choice of the six species (Fig. 4).
These tiny variations may be considered as mean-
ingless in overall percentages, but the model's
prediction has used them to extract more ecologically
relevant information than other techniques (Tables 3
and 4). In other words, on a microhabitat scale for this
size of fish, these small variations in environmental
variables quantitatively influence the dispersion of a
species and therefore its abundance, providing a wide
range of microhabitats are available. For this reason,
in the context of the River Garonne, the six species
may be considered to be well adapted to their
ecosystem and its spatial habitat variability, thus
resulting in their high abundance (Appendix 1).
Young-of the year fishes form shoals constituted of
small or large groups of individuals. Some species are
more likely to form larger aggregations (silver bream,
chub Leuciscus cephalus, gudgeon) as a result of their
reproductive success, environmental requirements
and behaviour (Fig. 3). As these sampling points
with high 0+ fish abundance (i.e. 160 0+ silver
bream in one sampling point, 38 0+ gudgeon in
another) resulted from determinist choices, the model
is able to give a reasonably accurate prediction of
these abundances, based only on habitat variables. It
is therefore possible to understand that certain
microhabitat combinations are strong attractive poles
for fish species of a particular age, providing a sort of
optimal habitat underlying the deterministic patterns
of fish assemblages composition (Oberdorff &
Porcher, 1992). This understanding is a key factor
for good river management in terms of habitat
variability. The presence of dams has completely
changed the River Garonne's hydrology and has at the
same time considerably reduced microhabitat diver-
sity. Transitory habitats on the main stretch of the
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 Structure and diversity of young-of-the year fish assemblages 817

Fig. 4 Representation of the contribution of environmental variables to each ANN model. (see Table 2 for fish codes).

river have disappeared in favour of more drastic
changes from lentic to lotic habitat types. As a result,
the progeny of typical rheophil species (e.g. barbel) as
well as typical limnophil species (e.g. silver bream)
have become numerically dominant in the Garonne
(Fig. 5). At the same time, fairly robust species with
very flexible ecological needs (e.g. chub), as well as
species preferring weak-to-moderate depths and
slope (e.g. roach, gudgeon, pumpkinseed), were not
affected by human impacts on the river (Gozlan et al.,
1998).
The ANN was successful in predicting the abun-
dance of 0+ fishes on a microhabitat scale in the River
Garonne, indicating that this technique merits more
frequent use in ecology and biodiversity studies for
which conventional models (mostly multiple linear
regression) are poorly suited (Table 3). Of course,
complex systems require more complex networks (i.e.
more units in the hidden layer or more than one

Table 5 Correlation of the variables accounting for the trace of

the ordination. See Table 2 for the fish codes

Variables Axis 1 Axis 2

ABj 0.96 0.00

ABb 0.60 0.19
ALg 0.01 0.70
AGg 0.26 0.17
ARr 0.12 0.20
Fig. 5 Correspondence analysis biplot, axes 1 and 2, for the nine ALc 0.07 0.05
biological (output) variables and the seventeen (input) Sh 0.00 0.17
environmental variables collected in the River Garonne (eigen FSR 0.07 0.17
values given in the inset). See Table 2 for codes. Ind 0.17 0.05

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818 R. E. Gozlan et al.
hidden layer), adequate training as well as the
modelling of larger data sets. With Garson's algorithm
(1991), despite the fact that it is a type of black box
model, it is still possible to visualise the contribution
of each variable in constructing the model. The
explanatory part of the analysis, coupled with the
predictive power of ANN, should facilitate the
ecologically oriented management of aquatic ecosys-
tems, providing that the duration of the study is
extended. It would be necessary to monitor the river
over at least a decade in order to introduce temporal
variability into the model, thus giving a more accurate
prediction. Other environmental variables, such as
discharge and temperature, may require more accu-
rate and systematic measurement to predict variations
due to season (spawning) and time of day (see Garner
et al., 1998).
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Appendix 1 The relative density (ind m±2) of the six most abundant 0+ fish species at various sites in the River Garonne. The site
number and type are indicated as flowing channel (fc), partially abandoned channel (pa), abandoned channel (ac). The number of
sampling point per site as well as the total number of specimens (n) by species and site and the total relative density (RD, ind.m±2) at
each site are indicated (see Table 2 for the fish codes)

Species

No. of sampling No. of fish

Sites points Bb Gg Lc Rr Lg Bj sample per site RD

1 pa 40 1.4 0.7 5.6 4.9 0.4 56 19.9

2 fc 24 0.6 3 1.8
3 fc 37 2.3 4.3 10.6 7.4 0.4 72 28.2
4 fc 30 0.5 0.9 0.5 0.5 27 12.7
5 pa 40 0.4 34.5 6.3 19.7 3.5 12.7 458 160.3
6 fc 30 0.5 27.7 8.5 0.9 0.9 126 59.1
7 ac 40 0.4 3.2 56 20.0
8 pa 40 6.7 1.1 2.1 16.9 340 120.0
9 ac 30 1.4 2.8 4.7 48 22.5
10 ac 30 1.4 1.4 2.8 7.0 57 26.8
11 fc 15 8.5 6.6 45 42.3
12 ac 20 13.4 9.9 0.7 7.7 7.0 118 83.1
13 fc 25 0.6 0.6 1.7 0.6 0.6 1.1 15 8.6
14 pa 30 1.4 13.1 0.5 1.9 22.1 128 60.1
15 pa 20 5.6 2.8 16 11.2
16 ac 40 3.2 3.2 7.4 10.9 104 36.6
17 pa 70 8.5 10.3 12.1 2.0 0.2 0.6 290 58.4
18 pa 50 2.5 9.6 20.6 1.1 1.4 22 368 104.0
19 pa 20 1.4 2.8 3.5 7.0 110 77.5
20 fc 15 7.5 17.8 44.1 0.9 1.9 77.9 221 208.0
21 pa 25 0.6 3.9 29 16.3
22 fc 20 2.8 5.6 1.4 7.0 30 21.0
23 pa 45 0.3 1.3 0.6 12.5 26.9 152 47.6
24 ac 30 0.5 2.8 23 3.8 197 92.5
25 ac 60 0.2 0.7 10.8 1.2 93 21.8
26 ac 15 1.9 11.3 4.7 60 56.3
27 ac 60 0.2 0.2 12 2.7
28 ac 20 1.4 0.7 1.4 10 7.0
29 fc 15 1.9 3.8 1.9 3.8 21 19.8
30 pa 10 4.2 25.4 1.4 5.6 7.0 141 198.3
31 fc 75 0.4 2.3 1.9 2.1 11.5 198 37.3
32 fc 30 19.2 22.5 3.8 115 53.9
33 fc 20 3.5 6.3 10.6 32 22.5
34 fc 20 11.3 13.4 4.9 43 29.3
35 fc 15 1.9 0.9 0.9 7 6.6
36 fc 20 9.9 4.9 19 1.4 62 43.6
37 fc 25 5.1 2.8 6.2 35 19.8
38 fc 10 5.6 16.9 16 22.5
n= 1161 168 398 489 168 253 572 3911 47.5

ã 1999 Blackwell Science Ltd, Freshwater Biology, 41, 809±820