current status of the riverine fishes of the french mediterranean basin

Transcription

Biological Conservation 72 (1995) 137-158
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ELSEVIER
0006-3207(94)00077-8
© 1995 ElsevierScienceLimited
Printed in Great Britain. All rights reserved
0006-3207/95/$09.50+.00
C U R R E N T STATUS OF THE RIVERINE FISHES OF THE
F R E N C H M E D I T E R R A N E A N BASIN
Thomas Changeux & Didier Pont
Laboratoire d'~cologie des systOmesfluviaux, URA, CNRS 1451, 1, Rue Parmentier, Arles 13200, France
Abstract
the centre of the Gulf of Genoa (Fig. 1). This area
would appear to possess great diversity both in geographic and climatic terms. In his classification of
world ecoregions, Bailey (1989) distinguished no less
than four types within this area, all belonging to the
wet temperate division.
This physical diversity is accentuated in biological
terms by geological history. In recent periods, dominated by the quaternary glacial eradications, the fish
populations of the Rh6ne basin benefited from a previous connection in the Pliocene with the Danube system
(Thienemann, 1950), recognized as the centre of origin
for European freshwater populations (Banarescu,
1989a). In addition, the north-south orientation of the
main river valley (Sa6ne valley continued south by the
Rh6ne corridor) and the persistence of a temperate climate in the southernmost regions during the maximum
extension of the ice caps (Riss: 200,000 BP) (Monjuvent
1984a, b) allowed refuge areas to be maintained for
thermophilic species (Persat & Berrebi, 1990).
The combination of these special characteristics
of our study area explains the presence of seven species
(discussed below), which, according to the most recent
of European freshwater fish atlases (Maitland,
1981; Lelek, 1987; Terofal, 1987), all have a restricted
geographical range. None of these species has
been recorded in the Swiss part of the Rh6ne basin,
situated upstream of Lake Geneva (Pedroli et al.,
1991).
An additional three species have been recorded from
France since 1950 by single authorities, but their occurrence has not been subsequently confirmed: (i) Aphanius iberus Valenciennes, 1846 from near the Spanish
border (Arnoult, 1957); (ii) Valencia hispanica Valenciennes, 1846, another cyprinodont, from the same sites
(Arnoult, 1957); and (iii) Cottus petiti Bacescu, 1964, a
cottid from the Lez, a short coastal river flowing into
the Gulf of Lion (Bacescu & Bacescu-Mester, 1964).
These species have not been accepted, and neither has
the Corsican subspecies of the brown trout, Salmo
trutta macrostigma Dum6ril, 1858.
The current status of these endemic taxa is worthy of
re-examination because of the major man-made changes
that have occurred to the French rivers, particularly
during the second half of the 20th century (Persat,
1988).
This work aims to provide a review of the current
status of these seven species or subspecies in France
We have undertaken a study of current knowledge on the
distribution and abundance of 10 freshwater fish species
with a restricted geographical range, recorded since 1950
from French rivers flowing into the Mediterranean,
including those in Corsica. In order of decreasing abundance these are: Leuciscus (Telestes) souffia, Chondrostoma toxostoma, Barbus meridionalis, Zingel asper,
Blennius fluviatilis, Aphanius fasciatus, Alosa fallax
rhodanensis, Aphanius iberus, Valencia hispanica and
Cottus petiti. Distribution maps for the first seven taxa
have been produced on the basis of 3124 fisheries
surveys included within Fisheries Orientation Schemes
carried out in French ddpartements. These taxa are
distributed in three different zoogeographical regions:
a northern mainland region, a southern mainland region
and the island of Corsica. The distribution of these taxa
in the major rivers is shown on maps by means of lines of
varying thickness and for the minor river courses by
shading. The detailed habitat preferences of the first
three species have been determined, showing a succession
of Barbus meridionalis, Leuciscus (Telestes) souffia and
Chondrostoma toxostoma going downstream within the
southern mainland region. Following a discussion of the
requirements of each of these species and of the threats
that they face within the study area, their status is
defined according to the International Union for the
Conservation of Nature definitions.
Keywords: endemic, fish, conservation, France, riverine.
INTRODUCTION
The distinctive character of the fish populations of running waters in Mediterranean France and Corsica has
been known for a long time (Roule, 1904, 1933; Mathias, 1921; Kreitmann, 1932). This distinctiveness has,
however, rarely been integrated into studies that are
currently used in this country to analyse fishery policies
(Leger, 1937; Huet, 1954; lilies & Botosaneanu, 1963;
Verneaux, 1973).
In France, the catchment area of rivers flowing into
the Mediterranean covers 134,360 km 2 (24.4% of the
area of France) and comprises mainly the catchment of
the Rh6ne (15% of the total area of France), part of
which lies in Switzerland and has been excluded from
this study, plus small coastal rivers along the entire
Mediterranean shoreline and the island of Corsica in
137
T. Changeux, D. Pont
138
with Meuse
basin
Col
with Seine
basi~,~
with Rhine
basin
with Lo
basin
Lake Geneva
Large rivers
.
selected
- - - excluded
Small tributaries
[ ] selected
[ ] excluded
connection by canal
with adjacent basin
•
I
Town
1 O0 kin
I
Nice
with G a n
basin
Marseille
Fig. 1. A, Study area defined as the entire area in France draining towards the Mediterranean. a, The Rh6ne basin; b, Corsica;
c, coastal river to the east of the Rh6ne; d, Western coastal rivers. B, Rivers and drainage units identified within study area. The
numbers given increase in order from west to east and from downstream to upstream and indicate the rivers and lakes cited in
the text (1, Aude; 2, H6rault; 3, Lez; 4, Vidourle; 5, Lower Rh6ne; 6, Upper Rh6ne; 7, Durance; 8, Ard6che; 9, Is~re; 10, Sa6ne;
11, Lac du Bourget; 12, Lac d'Annecy; 13, Doubs; 14, Argens; 15, Var).
by defining their distribution range, the habitat types
in which they occur and the accompanying fish fauna
in the areas where they are still recorded. These results
are compared with data from recent publications
(Kiener, 1985; Allardi & Keith, 1991) and older maps
published between 1924 and 1956 (references given
later in the text) in order to produce a diagnosis of the
risks of extinction and o f measures that could be taken
to guarantee the survival of the species concerned.
M A T E R I A L AND M E T H O D S
Study area
The study area is typified by its location at the western
end of the east-west chain of the Alps, which separates
the north European plain from the catchment of the
Mediterranean (BRGM, 1980). The Alpine geological
system occupies the entire area to the east of the Rh6ne,
including Corsica, and is prolonged to the south-east by
Status of French river&e Mediterranean fish
the chain of the Pyrenees, the southern limit of the
study area. The valley of the Sa6ne, which represents
the ancient river axis, communicates with the Mediterranean basin via a special geological formation (the
Rh6ne trench), through which the River Rh6ne now
flows in a north-south direction. The western boundary
of the study area is the Massif Central.
Vegetation is an indicator that is much used to
describe terrestrial habitat characteristics. From this point
of view, the study area contains 10 different phytoecological regions out of the 14 recorded for France as
a whole (Dupias & Rey, 1985). The special features of
the valleys of the Rh6ne and Sa6ne justify their being
attributed to regions of their own. The remaining area
is mainly occupied by the Mediterranean lowlands and
regions associated with the various mountain massifs
such as the Alps, Massif Central and Corsica.
The Mediterranean climate of the south of France
is typified by a negative hydrological balance (annual precipitation lower than evapotranspiration according to
the definition of Pagney (1988)). The water supply of
most rivers is therefore ensured by their sources being in
other climatic regions. The Mediterranean rainfall pattern
dominant in the south is typified by a summer drought
that can lead to complete drying out of small rivers. With
increasing altitude, the pluvial, pluvio-nival, nivo-pluvial,
nival and glacial regimes succeed one another.
The combination of these regimes makes the discharge
of the Rh6ne very high in comparison with its length:
it has the 42nd highest discharge in the world, but its
length is less than the 100th longest in the world according to Welcomme (1985).
Data sources and selection
French fresh waters, with the exception of the largest
natural lakes, have recently been subjected to systematic
investigations as part of the Departmental Fisheries
Orientation Schemes, using a standardized method
drawn up at a national level (Souchon & Trocherie,
1990). Rivers were divided into sections corresponding
to ecologically homogeneous units. Habitat and faunal
composition information was made accessible to our
analyses in the form of an Oracle relational database
(Trutna & Mone, 1990).
The field of study was purposely restricted to water
courses showing a wide range of species diversity: only
rivers containing at least three species in at least one
section of their course were retained. If this was the
case, the entire river course was selected, including the
headwaters, where there may be fewer than three
species. This preliminary selection led to the elimination of many small mountain streams (Alps and Corsica) and most of the tributaries of coastal rivers (Fig.
1). However, no river section containing endemic
species was eliminated. In total, 20,181 km of river
course were taken into account, divided into 2431 sections of lengths between 0.2 and 149 km, with a mean
of 8.2 km (Sx = 7.5). 74.2% of sections had a length of
less than l0 km. The longest sections (>20 km: 3-6%)
belong to the lower courses of major rivers.
139
Faunal lists
Of the 2431 sections retained, 1 781 had been subject to
one or more fishery investigations (n = 3 124). These
consisted of electric or net fishing operations in 72.8%
and 0.3% of sections, respectively, or of angler questionnaires in 12% of cases. In the remaining cases
04-9%) the method of investigation was not stated in
the original documents.
As far as the seven species considered are concerned,
the frequencies at which the various investigation methods were used were comparable, except in the case of
Barbus meridionalis, for which questionnaires were
most frequent.
A total of 50 non-marine species was recorded from
all of the rivers flowing into the Mediterranean, 14 of
them non-native (Appendix 1). Among the latter, some
were intentionally or accidentally released into the natural environment and have acclimatised (Holcik, 1991).
Others, generally originating from the Danube, have
extended their distribution towards the west by means
of connections between river basins formed by navigation canals. Finally, some American species of
Salmonidae, such as the rainbow trout Oncorhynchus
mykiss, brook trout Salvelinus fontinalis and lake trout
Salvelinus namaycush are mainly represented by artificially stocked populations.
The number of sections in which each endemic species
is present provided a measure of its abundance divided
into three classes: rare (_<30 sections), common (between
30 and 140 sections) and very common (_>140 sections).
For each species and section, a list of accompanying
species was drawn up, distinguishing those accompanying species that were dominant (recorded from more
than 50% of sections). This list was compared with the
potential fish populations of Verneaux (1981), which in
a non-perturbed situation would be equivalent to the
association of species occurring in a river typed according to the classification of Illies and Botosaneanu
(1963).
Habitat variables
A maximum of nine habitat variables was recorded for
each section. Quantitative variables were grouped into
classes:
(1) the distance (km) from the source (recorded for
2 431 sections), measured with a map measurer
on a 1 : 50,000 map from the source to the centre
of the section and grouped into the following
classes: <10, 11-30, 31-100 and >100 km.
(2) mean altitude of the section, or if unknown, the
mean altitude of the fishing stations (recorded for
1040 sections): altitude classes <100, 101-250,
251-500, 501-1000 and >1000 m.
(3) the mean discharge at the head of the section
(recorded for 368 sections): discharge classes <1,
1-3, 3.1-9, 9.1-27 and >27 m 3 s i.
(4) the mean gradient of the section (recorded for
2 254 sections): gradient classes <1, 1-3, 3.1-10,
10.1-30 and >30%0.
140
(5) the mean width of the section (recorded for 2294
sections): width classes <2, 2-5, 5.1-10, 10-1-25
and >25 m.
(6) mean temperature for the hottest month of the
year (recorded for 310 sections), hereafter called
the summer temperature: summer temperature
classes <17, 17-20, 20.1-23 and >23°C.
(7) the hydrological status of the section (recorded
for 2365 sections) according to four categories:
main course (MC) secondary course or short-circuited section (SC), reservoir (R) and rivers drying out in summer (TW).
(8) the physico-chemical water quality for the section (recorded for 2011 sections) defined in terms
of the physico-chemical index currently used by
French river authorities (Anon., 1989) according
to five categories: 1A, waters considered free from
pollution; 1B, water quality slightly poorer, but
suitable for all uses; 2, water quality satisfactory,
possibly harmful for the reproduction of the
most sensitive fish species; 3, poor quality, of
possible danger for fish survival particularly
at periods of low discharge; 4, water considered
unsuitable for most uses.
(9) the influence of neighbouring bodies of standing
water according to two categories: no nearby
water bodies, or the presence of standing water
upstream or downstream of the section whether
directly connected or not.
The selection of sections occupied by a given species
provided the distribution of the categories of each variable. This was then compared with the distribution for
the entire 2431 sections by means of a chi-squared test.
This comparison could only be carried out for the three
most frequent species (Barbus meridionalis, Leuciscus
souffia and Chondrostoma toxostoma) for which the
statistical validity of the method could be ensured (Alldredge & Ratti, 1985, 1992). For these three species,
the habitat preferences were calculated for each variable using the following formula:
F s i - Fei.100
Fti
'
where Fsi is the number of sections in which the species
occurred per 100 sections with category i of the variable in question; Fei is the total number of sections
with category i in which the species occurs in the entire
number of sections sampled; and Ft~ is the total number
of sections with category i of the variable.
The habitat preferences (Fig. 2) are presented for
each species in the form of relative frequencies for each
of the n categories of the variable in question (Psi)
P s i - Fsi'lO0
n
~ F s i.
i=l
Distribution maps
The geographical distribution of each species was
mapped by distinguishing between major river courses
and their smaller tributaries. The area containing
the 3431 sections selected was first divided into 163
drainage units (Fig. 1). Within each unit, fish distributions in sections occurring on small water courses were
mapped by shading the areas of the units (n -- 163),
whereas sections on main river courses were shown as
lines corresponding to the number of main rivers (n -222).
Species occurrences per linear or area unit were
grouped into two classes. The entire data set was digitized with graphics software (Cohn et al., 1987-90) and
exported to a cartographic analysis program (Brossier,
1990).
RESULTS
Cyprinidae
Leuciscus (Telestes) souffia Risso, 1826
Status and geographical distribution. Leuciscus (Telestes) souffia belongs to one of the genera of Cyprinidae
(Leuciscus) best represented in French freshwater fish
populations. The polymorphisms and possibilities of
hybridization within the genus complicate its classification
and several sub-genera, traditionally shown in brackets,
are admitted.
The European distribution of L. souffia is centred on
the Alps, around which it has developed several distinct
morphological types ranging from Bavaria and Romania through to the Sa6ne and Rh6ne basins as far
as Italy (Spillmann, 1959). In France it is restricted to
the basins of the Rhine and the Mediterranean (Allardi
& Keith, 1991). Three forms with the status of subspecies can be distinguished (Spillmann, 1961). The
main dividing line is located in south-east France at
the River Var (Spillmann, 1960, 1962, 1965a,b), which
separates the Italian form Leuciscus souffia multicellus
to the east from the form inhabiting the Rh6ne basin
and rivers on the coastal plain (Leuciscus souffia agassizi). An intermediate variety Leuciscus souffia souffia,
inhabiting the River Var, has been retained as the type.
The polymorphism of this species has been interpreted
as phenotypic variations resulting from environmental
differences between (Spillmann, 1962) and within (Spillmann, 1970) basins and has been analysed by Aubenton
et al. (1970-1971) and Daget and Bauchot (1976).
This very common fish (478 sections) occurs mainly
in the Rh6ne basin and in coastal rivers to the east (Fig.
3). In agreement with Spillmann (1970) and in contrast
to the observations of Kiener (1985) and Hillenius
(1965), the H6rault basin forms the western limit of the
species' distribution. In contrast to earlier records,
which showed a discontinuous distribution (Leger,
1901, 1927; Hesse & Paris, 1924, 1927; Leger et al.,
1945; Spillmann, 1961) our study indicates that it is present in rivers around the confluence of the Rh6ne and
Sa6ne. The current distribution is more extensive not
only in terms of basins but also in terms of the length of
river occupied. There is, however, no evidence to suggest that this is due to a real extension in the distribution range, rather than faulty identification in old
Status of French riverine Mediterranean fish
Barbus meridionalis
Leuciscus soufia
141
Chondrostoma toxostoma
Slope ( m.km-I )
l >0-30..130~11111
0
l'(n<O.(X)l
192)
=
20
Width (m)
,25_~______~
10 - 2 $_111~11111111
l
4(1
0
2(1
~
40
0
40
~~-~
0
Water Quality
p <0.05
(n= 176)
3 II
0
0
,
20
20
4:1
l
<0.001
1 = 112)
2(1
40
p <0.00l
(n = 4.~0)
p <O.(X)I
(. = 1 9 2 )
~
/'(,l<0.(X)l=
444)
•
20
1
~ l l l l m
0
40
p
. . . . .
0
r I
2O
40
<O.(X)I
p <I).(X)I
(n = 112)
( n = 4111))
20
40
20
0
40
Hydrological status
T.W.
--
p <O.(X)I
S.A_. ~11
M.C.
(n = 178)
0
40
80
II
/
ns
(n = 100)
(n = 386)
0
40
80
0
r
i
40
80
W a t e r f l o w ( m 3 . s-I )
ns
(n = 103)
-I
i
I
(1
~
'
30
i
(~)
0
Average
temperature
(°C)
'
ns
(n = 26)
-~7
ill
'
~ ~
0
'
31)
60
'
'
i
'
p <11.05
(n = 77)
---~---~--I
'
30
'
p <0.05
(n = 36)
~11
F --r
60
~
~
0
'
3(/
L...._ m
0
!
3()
..~
6()
p <0.01
r
6()
0
30
60
Fig. 2. Habitat preferences of the three most abundant species. The histograms represent the frequency of occurrence of each
species (the total is equal to 100%). n is the number of sections included for each variable, nc indicates that the distribution could
not be tested by chi-squared, p is the probability of the null-hypothesis of a homogeneous distribution of the species and ns
indicates that the species is uniformly distributed (p > 0.05).
studies caused by the similarity in vernacular names of
this species (souffle) and Chondrostoma toxostoma (so{fie
or sofie) in the south of France (Doffer, 1957; Spillmann, 1970). In contrast, the species is probably now
less frequent in the River Rh6ne than it was at the
beginning of the century (Agence de Bassin RMC, 1988).
Habitat and accompanying species. The habitat preferences (Fig. 2) indicate that it is under-represented in the
upper reaches of rivers (distance from source less than
10 km; chi-squared test: p < 0-001) and in rivers with a
width of less than 5 m (chi-squared test: p < 0.001). It
occurs preferentially in sections with a gradient of
between 1 and 10%o (chi-squared test: p < 0-001) and at
altitudes of 100-500 m (chi-squared test: p < 0-001). In
contrast, mean discharge has no influence on its distri-
but{on (chi-squared test: p > 0.05). The summer temperature in the sections in which it occurs varies
between 11 and 26.5°C, with a mean of 20.8°C (% =
2.8°C, n -- 77). It is located preferentially in
waters where the summer temperature is greater than
20°C (chi-squared test: p < 0.05). It is affected little by
the nearby presence of standing water (chi-squared test:
p > 0.05) or by the hydrological status. Its under-representation in waters of very high quality (category 1A)
is probably related to the fact that such waters tend to
occur in the upper reaches. On the other hand it does
not seem to be affected by a degradation of water quality (categories 3 and 4).
The only dominant accompanying species are Salmo
trutta and Leuciscus cephalus. As indicated by Verneaux
(1981), L. souffia forms part of the community that
142
,~.
k
h
'
Presence of L. soufia
in 40 to 100 % of the included sections
Absence of L. soufia
....
i il
i >,,,
......................
Presence of L. soufia
(,
....
in 35 to 100 % of
the included sections
in 1 to 35 % of
\
i?,
,
Absence of L. soufia
Non sampled areas
r"
!'~ .,-""k
•-,..~;,..-,
!~-"
,£
:
9
I
5°~1
t"
"I
../~:'~,.,'....-'" _;7"
Leuciscus soufia
Fig. 3. Distribution map of the blageon Leuciscus (Telestes) souffia. Main river courses are shown as lines. Small water courses
are pooled in surface drainage units.
occupies the zone extending from the mesorhithron to the
epipotamon. However, this species is better represented
than this author suggests in the part further upstream
(mesorhithron), where the species diversity is low and
the community restricted to the most rheophilic species.
Particular biological characteristics. Leger (1937) and
Dorier (1957) suggested that L. souffia should be considered as the indicator species replacing Thymallus thymallus
in southern rivers. In the northern part of the Rh6ne basin
the two species are frequently associated (Verneaux,
1973). In the southern part, where T. thymallus is absent,
since it belongs to a different zoogeographical assemblage,
L. souffia tends to inhabit stations with a steeper gradient
and narrower width than those occupied by T. thymallus
(according to the diagram of Huet, in Anon., 1984). L.
souffia has rarely been managed by man since it is difficult to rear in standing water (Spillmann, 1974): fish
reared under such conditions have a smaller head compared with the rest of the body (SpiUmann, 1965b).
Chondrostoma toxostoma Vallot, 1836
Status and geographical distribution. Chondrostoma toxostoma is the only native representative of the European
genus Chondrostoma occurring in the south of France
(Mathias, 1921).
This common species (129 sections) occurs in most
of the Rhrne basin and in coastal rivers to the west
(Fig. 4). Nelva (1988), Nelva-Pasqual (1985) and
Kiener (1985) indicated that it was present in coastal
rivers to the east of the Rhrne. Allardi and Keith
(1991) also recorded it from several stations in the
143
c
,.."
.~
g---i
,.'...," /
j
....
,."
Presence of C.
toxostoma
....g:.---.i(
i
I
_o. o . l , .
Absence of C.
toxostoma
......................
g
Presence of C.
wxostoma
'n
..""...l'
in 40 to 1130% of
in 1 to 40 % of
Absence of C.
.....
toxostoma
Non sampled areas
---'"
- i'",
.a
x ,. . '
.."
"
"
""
~ .~t, ..... .-,
( t ~ - ',k
~.._--,
'.,,'-:,"
"~'~_ ~
~
"-.
:
,
~
--
N
~
,,
..
50 km
i ....
.-
-~ "
I
1'.
" ......
':. ,"" -'!
,,-....,,..,.,
"
~":" 2:.-"
......
'...'"
"~
(.
_.r')(""',
2.-,-
.i:J i...:]!:,
, _ _ ~
"'"', . . . . " ~
,......
,,
..:
~"" . F . . . . . . , . . . . :
Nk~
l'""-, ~
o....-;
'c---'""....... ..
.--",,
-
- ....
:
";" "~
-...
"
"::(~.!. i;
Chondrostoma toxostoma
Fig. 4. Distribution map of the south-western European nase Chondrostoma toxostoma. Main river courses are shown as lines.
Small water courses are pooled in surface drainage units.
G a r o n n e basin and in the upper reaches of the Loire
and Seine basins.
Old distribution records are not very reliable because
of the similarity of the vernacular names of C toxostoma
and L. souffia and the difficulty in distinguishing the immature stages of Chondrostoma nasus and C toxostoma.
Habitat and accompanying species. Most of the variables which are significantly different f r o m the mean
rank or mean profile of the habitats surveyed (Fig. 2)
indicate a preferred distribution for the lower reaches
of rivers, more than 100 km from the source (chisquared test: p < 0.001), with altitudes o f below 100 m
(chi-squared test: p < 0-001), gradients of less than 3%o
(chi-squared test: p < 0.001), widths of greater than 25 m
(chi-squared test: p< 0.001) and discharges of greater than
27 m 3 s-l (chi-squared test: p < 0.05). It occurs preferentially in waters where the summer temperature greatly
exceeds 20°C (chi-squared test: p < 0.01): the mean summer
temperature in all of the sections in which it is recorded is
224°C (Sx = 3.03), with values ranging from 13 to 27°C.
Its absence from waters of good physico-chemical quality
(category 1A) is related to its location in the lower reaches,
whereas its presence in waters of poor quality shows that
it has a certain resistance. Its distribution in the various
hydrological categories could not be statistically tested.
It does not seem to be affected by the presence of
nearby water bodies (chi-squared test: p > 0.05).
This fish belongs to a community of 44 fish species,
of which 12 are non-native. The dominant accompany-
144
French rivers by way of canals connecting river basins
(Nelva, 1988). It is still difficult to assess the impact
of the arrival of C. nasus in the R h f n e basin on the
distribution of C. toxostorna. It seems that at the
start of this century, C. toxostoma was really subject
to competition since a decline was recorded in the
Safne (Paris, 1932). It is possible, however, that C. toxostoma was never very common in river habitats
(Nelva, 1988) and C. nasus does not seem to have
colonised very far up tributary rivers (Spillmann, 1961).
In the Ardfche, an equilibrium seems to have been
established between C. toxostoma, which occupies the
areas upstream, and C. nasus, which is combined to the
downstream parts (Nelva-Pasqual, 1985). The separation of the spawning seasons and dietary differences
are sufficient to allow the two species to coexist in the
contact zones (Chappaz et al.. 1989). Nevertheless, in the
ing species are, in order of frequency: Leuciscus
cephalus, Gobio gobio, Barbus meridionalis, Phoxinus
phoxinus, Rutilus rutilus and Anguilla anguilla. These
data are more or less in agreement with the potential
communities of Verneaux (1981) for the metarhithron,
but extend downstream to beyond the hypopotamon.
Particular biological characteristics. As with the other
species of the genus, it is typified by its inferior mouth,
especially adapted for scraping the perilithon. Its
growth, distribution, reproduction and diet during the
filling of an artificial reservoir in the south of France
have been studied by Chappaz (1986). It was located in
the lotic habitats in the upstream part of the water body,
and a decline in numbers by about 10% was recorded.
During the 19th century, a related but larger species
(Chondrostoma nasus) spread from the Danube to most
q;
.... ..'.'.
",4::'
,.-""k.',
S;
,:
"v
"~--.~i
;
-~-"
l
,-
,.-
,_,:~ ",,..
C"......... .,' ....
:~..I . .-...'~f
,
j
Presence o f B. meridionalis
*'°
>,,
,_,
¢......,.,
.'
~
\ ',. :/,,
..'J .~*
f' ,-"
,.
;,
."
.
/
.,
,,. ,;
in 60 to 100 % o f the included sections
;',
in 1 to 60 % o f the included sections
~, ';.: ','~'~
.,..
,,i
"'.
;, -
A b s e n c e o f B. meridionalis
~".
~ ,',
),~,-.~ ,~ ~ , , '
•'
~,
,"
~-
,~
i. ~.,.,"
"':
...
c
j
,,,
;
".
,
~-
,I-'
,,.
......................
:
.,
.;"
Presence o f B. meridionalis
I
in5OtolOO%of
"•; . , , x" .
-'"
,, ,.,-- ,, . . . . " ' "
~-
............... • in 1 to 50 % of
"'",,,,,_.,~--='.
A b s e n c e of B. m e r i d i o n a l i s . ,
-..,.
Non s a m p l e d areas
";
)
"-,".4,,.
., --,.,,
~-...,"",........ .
:
:
"-.,, . _ , , - "
.r,.
~
..'-.,.
"~.;" ,,' "
,:'t'" . . . . .~L
'.
/"
"".... '
. . . . "..
,;
.
- ,IZ"
IY!: "~
Barbus meridionalis
Fig. 5, Distribution map of the Mediterranean barbel Barbus meridionalis. Main river courses are shown as lines. Small water
courses are pooled in surface drainage units.
absence of C. nasa, C. toxostoma occupies both upstream
and downstream zones by adjusting its microhabitat preferences (Grossman et al., 1987a,b). C. nasus is currently continuing its spread towards the west via navigation canals
(Kiener, 1985) although little is known of the interactions
that exist between the two species. C. toxostoma does not
seem to have been directly managed by man.
Barbus meridionalis Risso, 1826
Status and geographical distribution. Barbus meridionalis
belongs to a genus of Cyprinidae whose distribution
extends from Asia to Europe and North Africa. Taxonomic and zoogeographical problems associated with
this genus were discussed in the 'International Barbus
Workshop' (Berrebi, 1990), but no definitive solution
to the classification was found, and some details still
vary from one authority to another (AlmaCa, 1990;
Doadrio, 1990). B. meridionalis is nevertheless always
placed in the tetraploid group of European species
along with the only other representative of the genus
Barbus in France: B. barbus. B. meridionalis has been
present since the Oligocene (c. 25 million years),
whereas B. barbus probably arrived from the Danube
basin at a much more recent time (Persat & Berrebi,
1990). Despite this, B. barbus is currently the more
common and more widespread of the two species.
This very common fish (215 sections) seems to predominate in the Mediterranean region (Fig. 5) in coastal
rivers and some of the southern tributaries of the Rh6ne.
Kiener (1985) also recorded it from all the coastal
rivers, with the exception of the Aude and the Arc. It
has for a long time been recognized as a member of the
communities of rivers in the south of France (Roule,
1904; Dorier, 1957). Outside our study area, Allardi and
Keith (1991) mention it from four stations in adjoining
rivers in the basins of the Garonne and Tam.
Habitat and accompanying species. B. meridionalis is
absent from sections further than 100 km from the
source (chi-squared test: p < 0-01) (Fig. 2). It occurs
preferentially in zones with a gradient of between 3 and
30%o (chi-squared test: p < 0.001) and of widths of
between 2 and 10 m (chi-squared test: p < 0.001). The
mean discharge in the sections that it inhabits is very
low (mean 3.42 m 3 s i, Sx 3-35) and does not exceed
11.5 m 3 s i. In the light of such results, the preference
of this species for altitudes of less than 100 m (chisquared test: p < 0.01) can only be explained by its
strictly southern distribution. Indeed, the Mediterranean region is characterized by the association of a
particularly low discharge and a lower altitude than the
average for the entire study area. Its preference for the
upper reaches of rivers in this region also leads to an
absence of any significant temperature profile (chisquared test: p > 0-05).
The hydrological status of the sections that it occupies shows that it is absent or rare in habitats that have
undergone major modifications (categories SC and R),
whereas it is very frequent in sections of river that are
subject to drying out in summer (chi-squared test: p <
TM
145
0-001). Similarly, the proximity of a standing water
body leads to its under-representation (chi-squared
test: p < 0-001). Waters of poor quality (categories 3
and 4) are also unfavourable for it (chi-squared test:
p < 0-05).
This fish forms part of the fish community that occupies the upper and middle reaches of rivers. The cumulative number of species occurring in the sections where
it is present is 38, of which eight are non-native. The
dominant accompanying species are, in decreasing
order of frequency: Salmo trutta, Leuciscus cephalus,
Phoxinus phoxinus and Gobio gobio. It does not figure
in the potential communities of Verneaux (1981).
Particular biological characteristics. There is still little
information on the ecology of B. meridionalis in our
study area. However, there are publications on the eastern subspecies concerning: its growth, reproduction,
density and mortality in Greece (Neophitou, 1987) and in
Yugoslavia (Soric & Jankovic, 1989) and its essentially
benthic, but selective, diet in Serbian rivers (Filipovic &
Jankoic, 1978) and the great temperature resistance of
its skeletal muscles (Pocrnjic & Omeragic, 1976; Omeragic et al., 1980). The closely related Spanish species (B.
haasi) only survives in the highest reaches of the River
Matarrafia in shallow areas, where it lives close to the
substrate or under shelter (Grossman et al., 1987b).
Similarly the distribution of B. meridionalis in France is
restricted to the most isolated tributaries (Leger, 1910;
Blaive, 1934; Dorier, 1957). It has been subject to management and artificial rearing (Raveret-Wattel, 1913;
Dorier, 1957; Philippart & Berrebi, 1990).
Percidae
Zingel asper L.1758
Status and geographical distribution. Zingel asper belongs
to a genus of Percidae whose distribution is restricted
to rivers zoogeographically connected with the Danube
(Colette, 1977). Its endemic status within the Rh6ne
basin is witness to an ancient connection between the
two rivers.
This rare fish (22 sections) only occurs in major river
courses within the Rh6ne basin (Fig. 6). It occurs in
four separate geographical areas: the north-east of the
Sa6ne basin, the middle Rh6ne (around the city of
Lyon), a few tributaries of the lower Rhone and the
upper reaches of the Durance basin. Kiener (1985) also
mentioned its presence in a few additional sites.
In 1900 it had a continuous distribution throughout
the major river courses (Rh6ne and main tributaries),
whereas currently it only occupies 17% of this area
(Boutitie, 1984a).
Habitat and accompanying species. The few habitat data
available in our database indicate a preference by the
species for sections typified by gradients of between 3
and 10%o, with widths of 2 to 5 m, distant from the
source (more than 30 km), but having water of good
quality (category 1A). The five values of mean dis-
146
J
1
<3
....
f"
L
', /t'~.~ ~"
•• :,.'""~-,
".:,
....-.,'"~
\°;
? ..<,,'.:,..:'
~ --:" i
i...!-..
~.
•
/"
:#
......,,.>.....
:"
I"
•
¢.-"k '
.
:-'--:z-,' :'.
.,.'..;
} t.-'r
.:
i
".,t. ~:,,"
; "',-.-.,",. "/4"
,,, ::
",'~2
",.i.,~
,°.....
Presence of Z. asper
',
~
::
....,,
i
,~... ....
-.,-,.,,.,-
'
Absence of Zasper
......................
.j~,
--.
,.J"
),._.,",
.......:
¢...:-,.
("--.. "-'"'.~ ~ .';
50 km
::
!.. /.",.,,
~(,,
";,:" ,:::'4
.,""
.__".......""
I "..
'. v
......
'"",.,.,i
I
:! )
i~~, -.:
¢
~:iii~
f-: ....
Zingel asper
Fig. 6. Distribution map of the asper Zingel asper. Main river courses are shown as lines. This species does not occur in small
water courses.
charge recorded for this species are between 7.5 and
126 m 3 s-1. Its preference for faster flowing waters in
the south of the region (Boutitie, 1984b) can be related
to its requirements for a spawning temperature (between 6 and 14°C) that is particularly low for a percid
(Perrin, 1988).
The cumulative number of species recorded in all
sections in which it is present is 34, of which eight are
non-native. The dominant associated species are, in
decreasing order of frequency: Salmo trutta, Barbus
barbus, Noemacheilus barbatulus, Leuciscus cephalus,
Leuciscus souffia, Gobio gobio, Chondrostoma nasus and
Phoxinus phoxinus. According to Verneaux (1981), this
fish belongs to the fish communities of the middle and
lower reaches of rivers, from the mesorhithron to the
epipotamon.
Particular biological character&tics. There are practically no publications on the biology of the species. The
related species from the Danube are better known (Banarescu, 1964) and could provide biological analogies
in terms of the description of the habitat type (Hensel,
1979), morphological variation (Smirnov, 1971; Holcik,
1979), breeding parameters (Bastl, 1981), growth and
diet (Geracopol, 1970; Makara & Stranai, 1980). This
species is active at night, lives close to the bottom in
large rivers and is easily overlooked (Spillmann, 1961).
Its rarity, already reported from the start of the century, makes it likely that its populations have always
had low densities, especially as it is a species with a
particularly low fecundity for a percid (Boutitie,
1984b). It is recognized as being sensitive to overall
habitat degradation (Verneaux, 1981), but the precise
"C2.,-';"
',
,"........
~ / .'~
"""~3
...
,~..,"
."
.~:,-ql--C"':-.
;-"?
,, %.,--,
...,~g
; ,.o.r
/
;
.a"
,"",........ v ....
•
147
I
l"
;
t
"
/ " ....... "~"
.,' I'"'\ ~ '; .:.....:" ;'" ./".r....
Presence of B. fluviatilis
,
,
,
,
1'
f
'
a
~.,
-~..
"~,-,
:' .....~. " '-'~ .-"
- "
;,
,~ :,~.. ,,
,':: ,-" ' .,', ""
,':7
:'" },~:-~>4-" k
".
It,
#
~.
A b s e n c e of B. fluviatilis
c"" -/ .... "",,
Presence of B. fluviatilis
I
'
in30to 100%of
f""~
in 1 to 30 % of
.....
A b s e n c e of B. fluviatilis
. ....
"-,.
_:',
"',,,,--, " "
-.: .....
.
-..
• ~
Non sampled areas
../'"
.,, ,,, ,..T
-
!,_ "~'.'""
~,
:',.
•
:
('
,', "-~ . . . . . .
~
,."" . . ~ "
-"
';
50 km
'
'
•
;,
'
£
'
-
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'
,, ".
i
,.,.. '
,,
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d
~".,
"'":",
",.
,
'
"',,,"
!...~..:
.,,4":'-"
:.,
.-"
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.)
,<':~
. . - ~ , '"~~
"l,,;
- ' l
,~¢
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".--..-
'~ '~
:',
t
, ""(
"., ,,'-: i',..
£., ,-..
.
..
.," "k
..,.~. .... _.._~:.
, c"-.v"'L'3:'.-',".-. ".-~-'~
~
•
',
•
,''-
i ': ,,'
/
" " . ....... ' " "
~,".'i..""........
'
~.-..,,.,
<
"j
"', " " ' . ' " , .
,.." i,
~,---"" ...... "' !":, "'-j
y "
,>.-.,.,
,
"",,; . . . . . . - ' " ' "
" ......
..~-<~.~"
-
/
~
..
A.",~?"
........ ""
k"
~'
. . . . . . . . . ..,,--" . "-._g?-_
.
,'
,
,,~ ..... ~, ,"
" ...-<
.,:'~ "--.4,,
"...,
..,~-~.
"
'
-
4
' i -°
.o._.o...~or
-._._.
Blennius fluviatilis
Fig. 7. Distribution map of the freshwater blenny Blenniusfluv&tilis. Main river courses are shown as lines. Small water courses
are pooled in surface drainage units.
causes of its decline are not clear (Boutitie, 1984a). The
disappearance of rime-pool sequences in the major
rivers is still the most obvious explanation. First, nonsilted bottoms are required for spawning and egg survival and, secondly, calm water areas are essential for
the surface-dwelling planktivorous larval stages (Leger
& Stankovitch, 1921). The related species from the
Danube are considered to be the percids most sensitive
to changes in current speed caused by river management (Zauner, 1988). This species is not highly
esteemed by anglers and does not seem to have been
artificially managed up till now. Its territoriality causes
problems for aquarium rearing (Perrin, 1988).
Blenniusfluviatilis Asso, 1801
Status and geographical distribution. Blennius fluviatilis
belongs to a genus of marine origin (Roule, 1935b).
This species occurs both in rivers with a direct connection to the Mediterranean and also, outside our sampling network, as isolated populations on the rocky
shores of some alpine lakes (Bourget and Annecy)
(Leger, 1943; Allardi & Keith, 1991). Outside of the
Mediterranean basin, the species has been recorded for
a long time from a station in the mid-Garonne (Roule,
1904; Spillmann, 1961; Allardi & Keith, 1991). Its presence in the Canal du Midi, which provides a connection between the Aude and the Garonne, indicates that
it could have entered the Garonne basin by this route.
This rare fish (28 sections) occurs essentially in three
main river types: in small coastal rivers on mainland
France and Corsica, in some sections of major rivers
(Rh6ne, Aude) and in some tributaries flowing directly
into the Rh6ne or into the Lac du Bourget (Fig. 7).
This distribution differs from that published recently by
148
Allardi and Keith (1991), in which it is recorded in the
Ard6che, Is6re and Sa6ne, but these records require
confirmation. The species lists of Kiener (1985) extend
its distribution in coastal rivers to the lower parts of
the most important rivers situated to the west of the
Aude.
Our information confirms that it is well represented
in Corsica (15 sections) as shown in recent studies (A1maqa, 1988; Roche, 1988), whereas it was formerly
considered as rare (Roule, 1933, 1935a).
This fish is always uncommon in the Rh6ne. For
example, in the upper Rh6ne it only occurred in 6% of
fish catches and only as a few individuals per catch,
and has not increased in numbers in the last 10 years of
monitoring (Persat, 1988). The oldest record in the
Rh6ne basin is from the Lac du Bourget and in the
Rh6ne next to it (Leger, 1943). Recent information
would therefore seem to indicate an expansion in its
range within this basin although its preference for relatively deep water could have led to it being overlooked
in inventories from the start of the century. In addition, its numbers are always low and suggest that the
river is not a very favourable site for this species, which
according to our results and the conditions recorded in
the additional sites provided by Allardi & Keith (1991)
prefers smaller rivers and lakes.
Habitat and accompanying species. The habitat characteristics indicate a preference for the lower courses
(great distance from the source, low mean gradient, low
altitude), although the largest rivers do not seem to be
included since the best represented width class is 2-5 m.
The mean discharges are varied (1-1421 m 3 s l), but
these sections nevertheless have rather high summer
temperatures (from 19.6 to 23°C).
This fish belongs to a fish community of 48 species of
which nine are non-native. The number of dominant
accompanying species is only two (Anguilla anguilla
and Salmo trutta) because the majority of sections are
located on the island of Corsica. It does not figure in
the potential communities of Verneaux (1981).
Within our study, which only takes into account running waters, it only appears in two sections of small
streams situated at either end of the eastern coastal
plain of the island of Corsica (Fig. 8). The presence of
A. fasciatus on the island, where it occurs in all the
lagoons along the east coast, has been known since the
start of the 19th century (Kiener & Schachter, 1974;
Kiener, 1985; Almaqa, 1988). As the lagoon habitat is at
the limits of the freshwater environment, this species
has not usually been included in studies of the fish populations of the river network of the island (Roule,
1933; Roche, 1988).
Its distribution, extending to Sardinia, where it is
considered as rare (Almaqa, 1988), and to nearby
coastal Italy, probably results from possible connections between these three regions at periods of lowered
sea level during the glaciations. Its occurrence in
Tunisia in inland oases is a relict of former ancient
coastlines (Villwock, 1970).
Records from the coasts of mainland France
(Arnoult, 1957; Moreau, 1881) have never been confirmed by later surveys (Spillmann, 1961; Kiener &
Schachter, 1974). This absence could be the result of
the sensitivity of this fish not only to temperatures
(Kiener & Schachter, 1974), but also to the depth of
the submarine trench that separates the French coast
from Corsica and forms a barrier to dispersion. Despite
its tolerance of salinity variations, the species could be
excluded from the marine fauna by competition (Villwock, 1970) or because of its poor swimming ability
(Kiener, 1985).
Habitat and accompanying species. A. fasciatus, a lagoon
species, is a member of the euryhaline fish community
(Anguilla anguilla, Liza aurata, Atherina mochon,
Dicentrachus labrax and Mugil cephalus) together with
one non-native species (Gambusia affinis). It is absent
from the potential communities of Verneaux (1981),
which exclude estuarine zones. The Corsican populations of A. fasciatus use macrophyte stands as a refuge
and form shoals when they make excursions into open
water (Kiener & Schachter, 1974).
Particular biological characteristics. Freeman et al.
(1990) showed, in a moderate-sized southern Spanish
river, that this benthic and territorial species requires
non-silted pebbles on which to site its nests. In addition
it needs a minimum water depth at the period of low
water where it can find lower temperatures. It is held in
low esteem by anglers, but is sometimes used as a bait.
Its unusual reproductive behaviour and ornamental
properties make it interesting as an aquarium specimen
and for ethologists (Wickler, 1957).
Cyprinodontidae
Aphaniusfasciatus Cuvier & Valenciennes, 1821
Status and geographical distribution: This member of
the Cyprinodontidae is not particularly restricted to fresh
water. Its present distribution is strictly related to the
brackish waters of coastal lagoons with salinities of
between 10 and 35 g litre ~ (Kiener, 1985).
Particular biological character&tics. The fact that G.
affinis is always dominant numerically over A. fasciatus
suggests that there is a competitive relationship between
these two ecologically related species. Nevertheless, a
study carried out in Spain on a local species of Aphanius
(Aphanius iberus) has shown that there is no dietary
overlap with G. affinis since A. iberus feeds deeper in the
water column (Vargas & Sostoa, 1991). According to
these authors, the advantage of G. affinis resides in a
better utilization of the various microhabitats.
There have been numerous studies on this species
around the Mediterranean. The main subjects covered
are: its capacity to withstand extreme conditions
(Kiener, 1985), its sexual dimorphism (Kiener &
Schachter, 1974; Boumaiza, 1980), variation in breeding parameters (Boumaiza et al., 1979; Penaz & Zaki,
1985), its morphological polymorphism (Kiener &
149
Presence of A. fasciatus
in 25 % of the included sections
Absence ofA. fasciams
Non sampled areas
I , 50km
I
Aphanius fasciatus
Fig. 8. Distribution map of the toothcarp Aphaniusfasciatus. Small river courses are pooled in surface drainage units. This species
does not occur in main water courses.
Schachter, 1974; Boumaiza, 1980; Villwock, 1981)and
enzyme polymorphism (Villwock, 1982; Comparini et
al., 1984).
A. fasciatus is of small size, reputedly has toxic flesh
and does not appear to have been managed by man up
till now, but its low requirements and ornamental qualities make it interesting as an aquarium species. It
inhabits lagoons which have been the site of major
piscicultural developments during the last 15 years, but
nothing is known of their effects on A. fasciatus.
Clupeidae
A losa f ailax rhodanensis (Roule, 1924)
Status and geographical distribution. Alosa fallax belongs
to a genus of euryhaline migratory riverine Clupeidae.
Because of the morphologic plasticity of the species, it
is difficult to base the identification of the Mediterranean
subspecies Alosa fallax rhodanensis on morphological
criteria. Individuals of this subspecies have a much faster
growth and attain sexual maturity later than their
Atlantic counterparts, which gives them a larger size
when they enter rivers on migration. Douchement (1981)
interpreted these distinctions as an adaptation by Alosa
fallax rhodanensis to a migration of longer distance.
The same author also considered that the migrations
into rivers of Alosa alosa and A. fallax are controlled
by homing behaviour, leading to genetic isolation of
populations. However, his morphometric analysis was
unable to distinguish clearly between the Mediterranean populations of Alosa fallax occurring in the
150
T. C h a n g e u x , D. P o n t
(-...
/
g
;
0"='. ",
>~., ....;o..,
.. .... ~ '..,"
,.-" ,~., "~ ~ ",,..--'"
.:
L.,.--#'- /
;
°
r""--~'
,'-"
: :;
,,"
";
Presence o f A. f a l l a x
•
: " v - - , ',. ~," /~]e ~,..
~.
~',,
t., "',.
~.~.,
-~ L.,-"
t
50 km ~
r ....
,,.,.-.
I
~.', -,,
..
;.
~.......
"-".,
~.
,'".
.."~,~
-
~
,,.~\~
."?"
.......;'7~..,"
"..":;."-:"
z .... J"."~
~' ..7 ......
.,:-...~ ....... , ~
/",4
,~
".
i
"~.: :"
~,"
,,,,
,"
c"
•
",,, - ' " " "
:" ~"""'-"
;' ~
, ,.L,~
'....L.':---...%
~, "-'...""
,..
...," " , j
"'-,,;-..".,
.Y, ; :'-.,.,
c".,
. .
,-~
"~-'/.\
" "J,~
~",,'..-.',. *'"
.,
"....
",.'--..
'-.
/ ........ ~,
]"
-- ,- ,.
"
i
..,:"'.d
Absence of A l o s a sp,
/- ,-"
~j
'-. ~',
.,' ...
,.,,- ,,,,.¢:~:..~, ,,
. ....- ~..
c
( ....
T.'"
)
,-.'" ,,. . . . S"~""-r,, ~..~
in 33 to100 % of the included sections
Presence o f A l o s a a l o s a
:'.
"
....
J -'". '~ /
./
" ,.'~:
"""
,',
~..--'.-'
,...'~-;;~
., ~/
-
' ,"
:."h
.,...
t"" #m-"
"~.- "--~"
....:.,-..
"
~ /
-
.,,-
">':, ...-." -.,,',..,..
'
~;/-"......
...." , , ,
,," ,'" ",
..:
'....
""
.. ....
3_ .," ~ i
~ ....
/'.-, ..' "~;
......."'"
. .....
""
"
" ":
,."
.', ...-.........:?
:.-, ',.. ~ .~."
--."; .,>
"~"
-...
Alosa fallax
Fig. 9, Distribution map of the twaite shad Alosa f a l l a x and allis shad Alosa alosa. Main river courses are shown as lines. These
species do not occur in small water courses.
Aude from those of the Rh6ne, although homing has
been established for the USA, and has been demonstrated indirectly in France (Douchement, 1981).
This rare species (10 sections) only appears in major
river courses where there is a permanent direct connection with the sea (coastal rivers, lower Rh6ne) except in
the case of the Ard6che (Fig. 9). The distribution of
A l o s a a l o s a is also given (Fig. 2) because the difficulties
in distinguishing between the two species and the very
precise information given by Douchement (1981) on
breeding individuals in the Var make us think that
there has been confusion and that A. f a l l a x is also present in the Aude and Argens. A l o s a a l o s a has always
been considered to be much rarer and less widespread
than A. f a l l a x (Leger e t al., 1945; Douchement, 1981;
Zylberblat et al., 1991).
The construction of insurmountable dams on the
Rh6ne from 1952 onwards prevented its migration
(Rameye e t al., 1976) and led to its distribution
range being restricted to the lower 60 km (Larinier e t
al., 1978; Petit, 1979). The recent record of this species
in the Ard6che, which was formerly an important
breeding site (Gallois, 1947a), is only based on
information provided by fishermen and needs confirming.
and accompanying
species. The quantitative
variables show a preference for the lower reaches with
a high temperature and which are frequently subject to
serious pollution. Roule (1929) and Quignard (1978)
have noted a preferential orientation of the migratory
routes towards the warmest river courses.
Habitat
In the sections where A. fallax is present, the total
cumulative number of species is 37, of which nine are
non-native. The most frequent in decreasing order are:
Anguilla anguilla, Ictalurus melas, Scardinius erythrophthalmus, Alburnus alburnus, Stizostedion lucioperca,
Perca fluviatilis, Rutilus rutilus, Leuciscus cephalus and
Tinca tinca. It does not figure among the potential
communities of Verneaux (1981).
Particular biological characteristics. Of all the endemic
fishes included in this study, A. fallax is the only one
subject to fisheries exploitation. Estimated catches in
1988 for the whole of the Rh6ne were under 4 tonnes
(Changeux & Zylberblat, 1989; Zylberblat et al., 1991),
compared with 34 tonnes or more before 1944 (Rameye
et al., 1976; Patt6e, 1988). Nevertheless the economic
importance of this fish has not been judged sufficient to
maintain the population by artificial stocking, due to
the difficulties of artificial rearing (Gallois, 1946a,b
1947b, c; Hoestlandt, 1948). Studies of the variety of A.
fallax occurring in the Nile show that the Mediterranean subspecies seem to be able to adapt to a more
varied bottom substrate and depth for reproduction
than its Atlantic counterpart (Ivanovic, 1977).
DISCUSSION
Status of the species
Of all the species with a restricted distribution recorded
from French running waters flowing into the Mediterranean, Aphanius iberus and Valencia hispanica have
only been reported once, without subsequent confirmation. In addition the existence of Cottus petiti (Bacescu
& Bacescu-Mester, 1964) as a true species has still to be
confirmed.
Of the seven species or subspecies considered in this
study, only Zingel asper shows a distribution restricted
to the Rh6ne basin. It is the only one in our study area
that can be classified as being endemic sensu stricto,
since the other species also occur in other countries
around the Mediterranean basin. The distribution
range of two (B. meridionalis and L. souffia) also
extends to north and eastern Europe.
The status of the seven endemic species as isolated
reproductive communities is debatable, at least for four
being able to hybridize with related coexisting species:
the hybrid A. fallax × A. alosa accounts for 10% of the
French population according to Douchement (1981),
but the degree of hybridization seems to vary from one
basin to another and its extent is debatable (Tavemy,
1991; Boisneau et al., 1992); species of the genera Leuciscus and Chondrostoma can hybridize with other
Cyprinidae including species belonging to other genera
(Collares-Pereira, 1989); B. meridionalis crosses in the
wild with B. barbus (Berrebi et al., 1986, 1988; Philippart & Berrebi, 1990).
Although the hybrids are not always fertile, the
transfer of genes from one species to another is possible. This is particularly recorded between B. meridionalis and B. barbus. Thus, in the absence of reproductive
151
genetic isolation, the integrity of these species can be
altered by the introduction of new species or by
modification of spawning sites.
According to Banarescu (1990), the species included
in this study do not all have the same zoogeographical
status in terms of their relation to fresh water. The
cyprinids L. souffia, B. meridionalis and C. toxostoma
belong, together with the percid Z. asper, to the group
of 'primary freshwater fish' which are strictly restricted
to fresh water and generally spread by overland routes.
A. fasciatus is a 'secondary freshwater fish', and has a
distribution related to the sea, as do B. fluviatilis and
A. fallax, which are classed as 'peripheral and vicarious
freshwater fishes'.
Geographical distribution and species abundance
The distribution maps and the abundance values
obtained for the French Mediterranean catchment can be
considered as correct, with the exception of two species
which occur in habitats located at the margins of our
study area (A. fasciatus in the lagoon environment and
B. fluviatilis in large alpine lakes).
As evidence of the evolutionary success of the
Cyprinidae in fresh waters, the three representatives of
this family are all common or very common, whereas
all the other species are rare (Appendix 1). The
cyprinids belong to three different genera (Barbus, Leuciscus and Chondrostoma). There are no genera containing more than one endemic species within the study
area, which, together with the low level of genetic isolation between species, suggests that a minimum degree
of taxonomic separation is needed for endemicism to
appear within this area.
In addition, these endemic species diffel- from sympatric species of the same genus by having smaller
maximum lengths (30 cm for B. meridionalis compared
with 80 cm for B. barbus, 30 cm for C. toxostoma compared with 52 cm for C. nasus, and 20 cm for L. souffia
compared with 65 cm for L. cephalus, or 30 cm for
Leuciscus leuciscus). This finding is in agreement with
the principles relating distribution to body size--smaller
sizes occurring in species with restricted distributions
(Peters, 1983), situated at lower altitudes (Linsley,
1966). However, this suggests, in response to Gaston
and Lawton (1990), that in freshwater fish the relation
between extent of distribution and body size must be
studied by using the genus as the unit.
Three main zoogeographical regions are distinguished: the island of Corsica, the northern mainland
region situated to the north of the confluence of the
Is6re and to the south a southern mainland region
comprising the coastal rivers. Because of its isolation,
Corsica is only populated by secondary species (A. fasciatus) or by peripheral and vicarious freshwater fish
(B. fluviatilis and A. fallax) for which the sea is no
obstacle to dispersion. In contrast, the northern mainland region, which is distant from the sea, is populated
almost entirely by primary species. The only exception,
B. fluviatilis, occurs here as a land-locked population
mainly in alpine lakes. The southern mainland region is
152
distinguished from the above region by the presence of
B. meridionalis. Its coastal situation means that its
endemic community is enriched by a peripheral species
of marine origin (A. fallax), although at the start of the
century this species ran up the rivers to spawn as far as
the northern mainland region (Kreitmann, 1932). The
catchments of the coastal rivers form a series of zoogeographical islands remarkable for their primary
species: none of these rivers has been colonized by Z.
asper and L. souffia remains absent from rivers situated
to the west of the H6rault.
Habitat and accompanying species
Habitat preferences could only be established for the
three most abundant species. These occur both in main
river courses (mapped linearly) and in small tributaries
(mapped on an area basis), whereas the other species
show a preference for main river courses (Z. asper, A.
fallax and B. fluviatilis) or small water courses draining
into coastal lagoons (A. fasciatus).
B. meridionalis, L. souffia and C. toxostoma succeed
one another along the longitudinal gradient, by occupying the upper, middle and lower reaches, respectively
(correlated with reduced gradient, increased width and
distance from the source). As most variables change
along this longitudinal gradient, it is difficult to distinguish between what is related to the position of the
species along the gradient and what is the effect of any
given variable.
The absence of some endemic species in the definition of the potential communities of Verneaux (1981)
means that this method cannot be applied to the entire
study area. An additional approach would be needed
to include the special zoogeographical features of the
south of France and Corsica. C. toxostoma does not
change its relative position in the south of France since
it is placed in accordance with the potential communities. In contrast, in the south of the study area, L.
souffia is placed in communities further upstream and
certainly does occur in areas higher up-river.
Analysis of changes in distribution and abundance
The decreases in both distribution and abundance of
A. fallax and Z. asper are certain, according to our
definition, but it is difficult to establish for sure that
there has been an increase in the distribution range
of L. souffia or B. fluviatilis. Similarly, it remains uncertain whether there has been a decline in the populations of C. toxostoma and B. meridionalis and any
changes in the abundance of A. fasciatus are unknown.
According to the Agence de Bassin Rh6ne M6diterran6e Corse (RMC) (1988) and Patt6e (1988) human
actions are largely responsible for these changes.
The reduction in migration routes by the construction
of dams that form obstacles to fish movements is the
reason for the decline in the stocks of A. fallax. However, the effects of dams on the entire fish community
are poorly known as there is very little information on
the importance of fish movements in the life cycles of
other species, particularly the endemics.
Habitat destruction following river management and
sand and gravel extraction from within river courses is
probably responsible for the disappearance of Z. asper.
Water abstraction, leading to seasonal drying out
of rivers, is a practice that is becoming increasingly
widespread in areas with a Mediterranean climate. This
causes the extinction of those species that are not
adapted and, if the fish are specially adapted, it can
accentuate the effects of pollution by decreasing dilution and favouring sedimentation. This is the case for
B. meridionalis, which is still sensitive to water quality
even though it can resist very low water levels.
Organic, chemical and thermal pollution affect many
water courses (37% of sections are in categories 2, 3 or
4). The presence of four major cities (Lyon, Marseille,
Nice and Grenoble) leads to heavy organic loads.
There are several major industrial centres associated
with these cities (Rh6ne valley south of Lyon, Golfe de
Fos near Marseille, alpine valleys around Grenoble,
etc.) that discharge nitrogen, phosphorus, organochlorides and heavy metals (Cu, Hg, Cr, Pb), to which must
be added the radioelements discharged by various
nuclear research centres. Seven nuclear and fossil-fuel
burning power plants heat the water of the Rh6ne by
1.5 to 4°C, depending on the season. The influence of
background levels of this (chronic) pollution is not
known in detail. Studies carried out in the Rh6ne basin
have shown that: (1) it favours the most tolerant
species in the community (L. cephalus, R. rutilus and
A. alburnus (Patt6e, 1988)); (2) it leads to bioaccumulation of pollutants that render the fish unfit for human
consumption (commercial fishing has been prohibited
upstream of Lyon); (3) it modifies metabolism (excess
detoxifying enzyme levels in the liver (Monod et al.,
1986)); (4) it leads to more abundant mucus secretion
(Patt6e, 1988); (5) it renders fish more susceptible to
disease (Kiener, 1985); (6) it can increase mortality
among young stages (Patt6e, 1988) and (7) it can lead
to siltation of gravel beds needed for spawning. In contrast, there are great risks of accidental mortality
by chemical pollution (nine accidents reported on the
Rh6ne in the last 20 years (Agence de Bassin RMC,
1988). The flushing out of sediments accumulated at
the bottom of reservoirs led to massive fish kills every
3 years up until 1978. The enormous tourist influx into
the Mediterranean region leads to increased sewage
loads during the summer when river levels are at their
lowest, which accentuates problems associated with
eutrophication. The same is also true of the intensification of aquacultural activities in coastal lagoons.
Alosa is the only endemic fish that is exploited by
commercial fishermen. The fishing effort using fixed net
installations is declining continuously as is the quantity
of catches (Rameye et aL, 1976; Kiener, 1985;
Changeux & Zylberblat, 1989, 1993). Populations are
currently at a very low level. The promotion of a new
type of exploitation of Alosa sp. by angling needs to
be monitored (Zylberblat et al., 1991). Angling for B.
meridionalis is popular (Berrebi et al., 1988), whereas L.
souffia and C. toxostoma are only of minor importance
for anglers. The possibility of B. fluviatilis and A. fasciatus being exploited as aquarium fish is a threat that
must be considered in the future.
The responsibility of man for introducing new species
likely to compete with endemic species concerns A. fasciatus, which is outnumbered by G. affinis in lagoon
environments, and C. toxostoma, which has declined
in favour of C. nasus in the Sa6ne. The first case is an
example of the inter-continental introductions dating
from the start of the century which also include Lepomis gibbosus, Ictalurus melas, Micropterus salmoides,
Oncorhynchus mykiss, Salvelinus fontinalis and Salvelinus
namaycush. The second case results from the current
extension of the distribution of species from the Danube
(Stizostedion lucioperca, Silurus glanis, C. nasus, etc.)
towards western Europe, that has been accelerated
by man by the construction of connections between
basins.
Legal status of species and conservation measures
Two types of laws that provide lists of protected
species are applicable in France.
International laws ratified by France
The Washington Convention (3 March 1973) does not
list any of our endemic species.
The Berne Convention (19 September 1979) distinguishes (i) strictly protected species for which conservation measures are needed including habitat protection,
and for which capture and possession are prohibited;
(ii) species needing special management, but whose populations may be exploited. All of our endemic species
are affected by one or other of these measures, Z. asper
and V. hispanica having the status of strictly protected
species.
The directives of the European Communities (21
May 1992) reiterate the Berne convention lists and
define those species that should be subject to special
management (B. meridionalis and A. fallax). B. fluviatilis is omitted from the EC lists.
French laws
The decree of 8 December 1988 prohibits the destruction or collection of eggs or the alteration or degradation of the breeding sites of four of our seven endemic
species. A. fasciatus, L. souffia and C. toxostoma do not
benefit from any protection measure at the national level.
The IUCN Red Data Book (Keith et al., 1992) has
recently established a reference status for endangered
species within France (Appendix 1). All the endemic
species, except C. petiti, appear in this list:
L. souffia is considered as rare within France as it
is endemic to the south-east. Our results confirm
this status and show that this species is relatively
insensitive to man-made changes. It is unlikely
therefore to be endangered;
C. toxostoma is considered as endangered because of
possible competition from C. nasus, which is
spreading towards the west in other French river
153
basins. Our results cannot either support or
refute this status;
B. meridionalis is considered as rare, because of the
disjointed nature of its distribution range. Our
results cannot either support or refute this status.
However, this species is certainly not extending
its range.
Z. asper is considered as threatened with extinction.
Our results confirm this status.
B. fluviatilis is considered as vulnerable because it is
particularly sensitive to habitat changes and
excessive water abstraction. Our results cannot
either support or refute this status. This status
does nevertheless agree with the declines
recorded in other countries (Vinyoles et al.,
1991).
Aphanius fasciatus is considered as rare because it is
very localized within France. Our results confirm
this status.
Alosa fallax is considered as vulnerable. Our results
confirm this status. In contrast to Atlantic populations, the Rh6ne shad has not benefited from
the programmes in favour of migratory fish
recently developed in France. Activities are currently in progress to allow shad to get past the
first few dams blocking its migration on the Aude
at Moussoulens (Desjames, 1988), on the H6rault
at Agde (Cruz, 1989), on the Vidourle at Mas de
Terre de Port (Bourgogne, 1988), on the Rh6ne
at Beaucaire (Zylberblat et al., 1991) and on the
Argens at Roquebrune (Anon., 1988).
In proportion to the number of species present in
the basin, those in RhSne-Mediterranean-Corsica are
the most endangered' with 46% of the species present
listed in Keith et al. (1992). Our results also show that
the majority of these species occur in the lower reaches
of the river, which are the habitats most affected by
human activities. These species are therefore particularly
vulnerable.
ACKNOWLEDGEMENTS
We would like to thank J. F. Perrin for giving us his
old maps of fish distribution. This work was supported
by the Agence de l'Eau Rh6ne M6diterran6e Corse. It
forms part of the Programme lnterdisciplinaire de
Recherche sur l'Environnement of the Centre National
de Recherche Scientifique entitled 'Environmental
factors, spatial habitat use and spatio-temporal structure
of fish populations'.
REFERENCES
Agence de Bassin RhSne Mediterran6e Corse (RMC) (1988).
Qualit~ du fleuve Rh6ne, synthOse des connaissances. Agence
de Bassin RMC/Minist~re de l'Environnement/D616gation
de bassin Rh6ne M6diterran6e Corse.
Allardi, J. & Keith, P. (1991). Atlas pr61iminaire des poissons
d'eau douce de France. In Patrimoines Naturels, No. 4, ed.
Secr6tariat de la Faune et de la Flore. Mus4um National
154
d'Histoire Naturelle de Paris/Conseil Sup6rieur de la P6che/
Centre National du Machinisme Agricole du G6nie Rural,
des Eaux et des For~ts/Minist6re de l'Environnement. Paris
84-85, 104-5,118-19, 200-201.
Alldredge, J. R. & Ratti, J. T. (1985). Comparison of some
statistical techniques for analysis of resource selection. J.
Wildl. Manage., 50, 157-65.
Alldredge, J. R. & Ratti, J. T. (1992). Further comparison of
some statistical techniques for analysis of resource selection. J. Wildl. Manage., 56, 1-9.
Almaqa, C. (1988). Compte rendu de la table ronde: les poissons des iles de la M6diterran6e occidentale (Bal6ares,
Corse, Sardaigne, Sicile). Bull Ecol., 19, 471-2.
Almaqa, C. (1990). Neogene circum-mediterranean paleogeography and euro-mediterranean Barbus biogeography.
Arq. Mus. Boc. (Nova Serie), 41,585-611.
Anon. (1984). Dossier A: 6tude g6n6rale. Sch6ma de vocation
piscicole et halieutique de la Dr6me. Minist6re de l'Environnement/Direction D6partementale de l'Agriculture et de
la For6t de la Dr6me/Conseil Sup6rieur de la P6che, Lyon.
Anon. (1988). Rapport de synth6se. Sch6ma d6partemental
de vocation piscicole du Var. Direction D6partementale de
l'Ariculture et de la For~t du Var/Minist6re de l'Environnement/Conseil Sup6rieur de la P6che, Montpellier.
Anon. (1989). Guide de l'agent pr61eveur. Minist6re de l'Environnement/Conseil Sup~rieur de la P6che/Centre National
du Machinisme Agricole du G6nie Rural, des Eaux et des
For6ts.
Arnoult, J. (1957). Sur quelques poissons rares et peu connus
des eaux douces de France. Bull. Mus. natn. Hist. nat.
( 2Ome sOrie) , 29, 464-6.
Aubenton, F. D., Daget, J. & Spillmann, J. (1970-1971).
Classification num6rique des blageons Leuciscus (Telestes)
souffia (Pisces, Cyprinidae), 86me note. Bull. Mus. hath.
Hist. nat. (2dine skrie), 42, 839-48.
Bacescu, M. & Bacescu-Mester, L. (1964). Cottus petiti sp. n.,
un chabot nouveau r6colt6 en France. Consid6rations
zoog6ographiques et donn6e comparatives sur d'autres Cottus d'Europe. Vie et Mileu, Suppl. 17, 431-46.
Bailey, R. G. (1989). Explanatory supplement to ecoregions
map of the continents. Environ. Conserv., 16, 307-9.
Banarescu, P. (1964). Pisces-Osteichthes. In Fauna Republicii
Populare Romine, No. 13. Academia Republicii Populare
Romine, Bucharest.
Banarescu, P. (1989a). The freshwater fishes of Europe. In
General introduction to fishes Acipenseriformes, ed. J. Holcik. Aula-Verlag. Wiesbaden, pp. 88-107.
Banarescu, P. (1989b). Vicariant pattern and dispersal in European freshwater fishes. Spixiana (Miinchen), 12, 91-103.
Banarescu, P. (1990). General distribution and dispersal of
freshwater animals. In Zoogeography of fresh waters, No.
1, ed. P. Banarescu. Aula-Verlag. Wiesbaden.
Bastl, I. (1981). Zur Fruchtbarkeit des Strebers, Zingel streher. Folia Zool. (Brno), 30, 177-80.
Berrebi, P. (1990). An example of multi-field knowledge-sharing on the european level - - the Barbus roundtable in
Montpellier (France), July 1989. Arch. Hydrobiol., 117, 383.
Berrebi, P., Lamy, G., Cattaneo-Berrebi, G. & Renno, J. F.
(1988). Variabilit6 g6n6tique de Barbus meridionalis Risso
(Cyprinidae): une esp6ce quasi monomorphe. Bull. Fr.
P~che Piscic., 310, 77-84.
Berrebi, P., Le Brun, N., Renaud, F. & Lambert, A. (1986).
Hybridation intersp6cifique de deux Cyprinidae (genre Barbus). Cons6quence sur la sp6cificit6 parasitaire de Dilpozoon
gracile (Monogenea). In Actes du Colloque National du
CNRS, 'Biologie des Populations" ed. Claude Bernard
(Lyon 1) University Institut d'Analyse des Syst6mes
biologiques et socio-6conomiques, Lyon, pp. 179-80.
Blaive, P. (1934). Une nouvelle station de barbeau m6ridional
(Barbus meridionalis Risso) en dauphin6. Ann. Univ. Grenoble, 11, 141~i.
Boisneau, P., Mennesson-Boisneau, C. & Guyomard, R.
(1992). Electrophoretic identity between allis shad Alosa
alosa (L), and twaite shad A. fallax (Lacepede). J. Fish
Biol., 40, 731-8.
Boumaiza, M. (1980). Dimorphisme sexuel et polymorphisme
d'Aphanius fasciatus Nardo, 1827 (Pisces, Cyprinodontidae). Bull. Off. natn. PYch., Tunisie, 4, 83-143.
Boumaiza, M., Quignard, J. P. & Ktari, M. H. (1979). Contribution h la biologie de la reproduction d'Aphaniusfasciatus Nardo, 1827 (Pisces, Cyprinodontidae) de Tunisie. Bull,
Off. natn. P~ch., Tunisie, 3, 221-40.
Bourgogne (1988). Rapport de synth~se. Sch6ma d6partemental de vocation piscicole du Gard. Direction D6partementale de l'Agriculture et de la For~t du Gard/Minist6re de
l'Environnement/Conseil Sup6rieur de la P~che, Montpellier.
Boutitie, F. (1984a). Biologic et r6partition de l'Apron, poisson du bassin du Rh6ne menac6 d'extinction. Rapport
d'6tude. Service R6gional d'Am6nagement des Eaux
Rh6ne-Alpes/Plan piscicole de la Dr6me.
Boutitie, F. (1984b). L'apron Zingel asper (L.), Percidae - poisson rare menac6 de disparition (biologie-r6partitionhabitat). DEA, Claude Bernard (Lyon 1) University, Lyon.
BRGM (1980). Carte g~ologique de la France et de la marge
continentale 6 I'~chelle de 1/1 500 O00dme et notice explicative. Bureau de Recherches G6ologiques et Mini6res,
Orl6ans.
Brossier, P. (1990). Logiciel de transfert d'entitOs cartographiques. Maison de la G6ographie, Montpellier.
Changeux, T. & Zylberblat, M. (1989). La pYche professionnelle et amateur aux engins. Schema de vocation piscicole du
fleuve Rh6ne, No. 7. Minist~re de l'Environnement/D~16gation de Bassin Rh6ne Mediterranee Corse/Service de la
Navigation Rh6ne Sa6ne.
Changeux, T. & Zylberblat, M. (1993). Analysis of the fishing
gear fishery statistics in the Rh6ne River Basin. Bull. Ft.
P~che Piscic., 330, 245-94.
Chappaz, R. (1986). Etude piscicole de la retenue de Sainte
Croix-Fontaine l'6v~que. Dissertation, Provence-Marseille
St Charles University.
Chappaz, R., Brun, G. & Olivari, G. (1989). Donn6es nouvelles sur la biologie et l'6cologie d'une esp6ce de poisson
Cyprinid6 peu 6tudi6 Chondrostoma toxostoma (Vallot,
1836). Comparaison avec Chondrostoma nasus (L., 1766).
C. R. Hebd. Seances Acad. Sci. (III), 309, 181-6.
Cohn, R., Hodge, B., Holt, J. et al. (1987 90). Adobe Illustrator® Software, Adobe, California.
Colette, B. (1977). Systematics and zoogeography of the
fishes of the family of Percidae. J. Fish. Res. Bd Can., 34,
1450-63.
Collares-Pereira, M. J. (1989). Hybridization in european
cyprinids: evolutionary potential of unisexual population.
In Evolution and ecology of unisexual vertebrates, ed. R. M.
Dawley & J. P. Bogart. New York State Museum, Albany,
New York, pp. 281-8.
Comparini, A., Scattolin, N. & Rodino, E. (1984). Genetic
differentiation among some populations of the cyprinodont
Aphanius fasciatus Nardo. In 15th Congr. Soc. Ital, Biol.
Mar. (28 Sept. 83). Trieste, (Italy). Nova Thalassia, 261-8.
Cruz, O. (1989). Rapport de synth6se. Sch6ma d~partemental
de vocation piscicole de l'H6rault. Direction D6partementale de l'Agriculture et de la For~t de l'H6rault/Minist~re
de l'EnvironnementJConseil Sup6rieur de la P~che, Montpellier.
Daget, J. & Bauchot, M.-L. (1976). Les probl~mes de l'esp~ce
chez les t616ost6ens. In Les problOmes de l'espOce dans de
monde animal, ed. C. Bocquet, J. Genermont & M. Lamotte. Soci6t6 Zoologique de France. Paris, pp. 17-27.
Desjames (1988). Rapport de synth6se. Sch6ma d6partemental
de vocation piscicole de l'Aude. Direction D6partementale
de l'Agriculture et de la For~t de l'Aude/Minist6re de l'Environnement/Conseil Sup6rieur de la P~che, Montpellier.
Status o f French riverine Mediterranean fish
Doadrio, I. (1990). Phylogenetic relationships and classification of western palaeartic species of the genus Barbus (Ostreichthyes, Cyprinidae). Aquat. Living Resour., 3, 265-82.
Dorier, A. (1957). R6partition du barbeau m6ridional dans le
sud-est de la France. Tray. Lab. Hydrobiol. Pisc. Grenoble,
48--49, 141-9.
Douchement, C. (1981). Les aloses des fleuves franqais Alosa
fallax Lacdp6de, 1803 et Alosa alosa Linnd, 1758-Biom6trie,
6cobiologie: autonomie des populations. Dissertation,
Languedoc (Montpellier) University.
Dupias, G. & Rey, P. (1985). Carte des r~gions Ocologiques de
la France 1/1000 O00Ome, document pour un zonage des rkgions phyto-Ocologiques. CNRS, Toulouse.
Filipoic, D. & Jankovic, D. (1978). Relation between the bottom fauna and fish diet of the upland stream of east Serbia
(abstract only). Acta Biol. Iugosl. (E. Ichtyol. ), 10, 29-40.
Freeman, M. C., Vinolas, I., Grossman, G. D. & Desostoa,
A. (1990). Microhabitat use by Blenniusfluviatilis in the rio
Matarrafia, Spain. Freshwat. Biol., 24, 335-45.
Gallois, G. (1946a). L'alose du Rh6ne. Bull. Fr. Piscic., 142,
5-14.
Gallois, G. (1946b). L'alose du Rh6ne. Bull. Fr. Piscic., 143,
72-9.
Gallois, G. (1947a). Am6nagement du Rh6ne vivarais par la
Compagnie Nationale du Rh6ne, ses incidences sur
l'6conomie piscicole, mesures tendant ~t les attdnuer. Bull.
Fr. Piscic., 146, 25-34.
Gallois, G. (1947b). L'alose du Rh6ne. Bull. Fr. Piscic., 144,
130-43.
Gallois, G. (1947c). L'alose du Rhone. Bull. Fr. Piscic., 145, 198.
Gaston, K. J. & Lawton, J. H. (1990). The population ecology of rare species. J. Fish Biol., 37A (Suppl.), 97-104.
Geracopol, O. (1970). Contribution to the biological study of
A. zingel from the Lower Danube. Hydrobiologia, 11,
143-53.
Grossman, G. D., de Sostoa, A., Freeman, M. C. & LobonCervia, J. (1987a). Microhabitats use in a mediterranean
riverine fish assemblage--fishes of the lower Matarrafia.
Oecologia (Berl.), 73, 490-500.
Grossman, G. D., de Sostoa, A., Freeman, M. C. & LobonCervia, J. (1987b). Microhabitats use in a mediterranean
riverine fish assemblage--fishes of the upper Matarrafia.
Oecologia (Berl. ), 73, 501-12.
Hensel, K. (1979). Geographical distribution of percid genera
Gymnocephalus and Zingel (Osteichthyes, Percidae) in Slovakia. Folia Zool. ( Brno), 21, 85-94.
Hesse, E. & Paris, P. (1924). Cours d'eau de la c6te d'or, 1.
Bassin de l'Ouche et de la Vouge. Institut de Pisciculture de
Grenoble, Grenoble.
Hesse, E. & Paris, P. (1927). Cours d'eau de la C6te d'or, 2.
Bassin de la Vingeanne, de la BOze et de la Tille. Institut de
Pisciculture de Grenoble, Grenoble.
Hillenius, D. (1965). Two new fishes for the fauna of Spain:
Leuciscus (= Telestes) souffia Risso, 1826 and Leuciscus leuciscus (Linnaeus, 1758). Beaufortia (Zoological Museum,
Amsterdam), 149, 1-4.
Hoestlandt, H. (1948). Fdcondation artificielle et incubation
de l'alose du Rh6ne: Paralosa rhodanensis Roule. Ann. Stat.
centre. Hydrobiol. appl., 2, 223-8.
Holcik, J. (1979). Supplements to the description of Zingel
streber (Osteichthyes, Percidae) with regard to its geographical variability. Folia Zool. (Brno), 28, 73-84.
Holcik, J. (1991). Fish introductions in Europe with particular reference to its central and eastern part. Can. J. Fish.
Aquat. Sci., 48, Suppl. 1, 13-23.
Huet, M. (1954). Biologie, profil en long et en travers des
eaux courantes. Bull. Fr. Piscic., 175, 41-53.
lilies, J. & Botosaneanu, M. (1963). Probl6mes et m6thodes
de la classification et de la zonation 6cologique des eaux
courantes, consid6r6es surtout du point de vue faunistique.
Ver. Int. Verein. Limnol., 12, 1-57.
155
Ivanovic, B. (1977). Sexual maturity and spawning of Alosa
fallax nilotica in lake Skadar. Ichthyologia, 9, 25-9.
Keith, P., Allardi, J. & Moutou, B. (1992). Livre rouge des
esp6ces menac6es de poissons d'eau douce de France. In
Patrimoines Naturels, No 10, ed. Secrdtariat de la Faune et
de la Flore. Mus6um National d'Histoire Naturelle de
Paris/Conseil Supdrieur de la P6che/Centre National du
Machinisme Agricole du G6nie Rural, des Eaux et des
For~ts/Minist6re de l'Environnement, Paris, pp. 111.
Kiener, A. (1985). Au fil de l'eau en pays mkditerranOen.
Aubanel, Avignon.
Kiener, A. & Schachter, D. (1974). Polymorphisme d'Aphanius fasciatus Nardo, 1827 (Poisson Cyprinodontidae) des
eaux saum~tre (Populations de Corse et de la lagune italienne de Comaccio). Bull. Mus. natn. Hist. nat. (3~me SHie),
142, 317-39.
Kreitmann, L. (1932). Les grandes lignes de l'dconomie piscicole du bassin franqais du Rh6ne. Tray. Lab. Hydrobiol.
Pisc. Grenoble, 24, 127-31.
Larinier, M., Rivier, B., Allardi, J. & Yrocherie, F. (1978).
Possibilitd de franchissement du seuil de Beaucaire par les
aloses de Rh6ne. Bull. Fr. Piscic., 268, 107-20.
Leger, L. (1901). Carte piscicole du dOpartement de l'Iskre avec
une notice explicative. Institut de Pisciculture de Grenoble,
Grenoble.
Leger, L. (1910). Sur la prdsence du barbeau m6ridional dans
les alpes du dauphind. Tray. Lab, Hydrobiol. Pisc.
Grenoble, 1, 1 7.
Leger, L. (1927). Carte piscicole du dOpartement de l'Ain avec
une notice explicative. Travaux du Laboratoire d'Hydrobiologie et de Pisciculture de l'Universit6 de Grenoble,
Grenoble.
Leger, L. (1943). Carte piscicole du dOpartement de la Savoie avec
une notice sur l'hydrographie et l~conomie piscicole des cours
d'eau et des lacs. Travaux du Laboratoire d'Hydrobiologie et
de Pisciculture de l'Universitd de Grenoble, Grenoble.
Leger, L., Burdin, M. G. & Arnaud, M. A. (1945). Carte piscicole du dOpartement du Rh6ne avec une notice sur l'hydrographie et l'dconomie piscicole des cours d'eau. Travaux du
Laboratoire d'Hydrobiologie et de Pisciculture de l'Universitd de Grenoble, Grenoble.
Leger, L. & Stankovitch, S. (1921). Fdcondation artificielle et
d6veloppement de l'apron Aspro asper L. Tray. Lab. Hydrobiol. Pisc. Grenoble, 13, 191-4.
Leger, L. (1937). Economie biologique g6n6rale des cours
d'eau alpins. Bull. Fr. Piscic., 109, 5-13.
Lelek, A. (1987). Threatened fishes of Europe. In The freshwater fishes of Europe, Aula-Verlag, Wiesbaden, pp. 343.
Linsley, L. (1966). Body size of poikilotherm vertebrates at
different latitudes. Evolution, 20, 456-65.
Maitland, P. S. (1981). Les poissons des lacs et rivi6res d'Europe en couleurs. In Multiguide nature. Bordas, ElsevierSdquoia, Bruxelle, pp. 255.
Makara, A. & Stranai, I. (1980). Notes on growth of the Zingel
streber and of the Z. zingel. Biologia ( Bratislava), 35, 595-9.
Mathias, P. (1921). Etude du genre Chondrostoma dans l'Europe occidentale et la r6gion circum-m6diterrandenne.
MOm. Soc. Zool. France, 1-2, 28, 52.
Monjuvent, G. (1984a). Quaternaire. In Synthkse gOologique
du sud-est de la France, stratigraphie et palOogeographie, ed.
S. Debrand-Passard & S. Courbouleix. Bureau de Recherches
G6ologiques et Minieres, Orldans, pp. 521-99.
Monjuvent, G. (1984b). Riss, pal6og6ographie au 1/1 500
0006me. In Synthdse gOologique du sud-est de la France,
Atlas, ed. S. Debrand-Passard & S. Courbouleix. Bureau
de Recherches Gdologiques et Minieres, Orl6ans, pp. Q2.
Monod, G., Devaux, A. & Rivi6re, J.-L. (1986). Une caractdrisation 6cotoxicologique des poissons du Rh6ne par leur
activit6 h6patique de biotransforrnation. In Actes du Colloque
National du CNRS, 'Biologie des populations', ed. Claude
Bernard (Lyon I) University IASBSE, Lyon, pp. 682-3.
156
Moreau, E. (1881). Manuel d'ichtyologie franfaise. Masson,
Paris.
Nelva, A. (1988). Origine et biog6ographie des deux Chondrostomes franqais: C. nasus et C. toxostoma (Pisces,
Cyprinidae). Cybium, 12, 287-99.
Nelva-Pasqual, A. (1985). Biog6ographie, d6mographie et
6cologie de Chondrostoma nasus nasus (L., 1758) (Hotu,
Poisson, Teleost6en, Cyprinid6). Dissertation, Claude
Bernard (Lyon I) University.
Neophitou (1987). A study of some autoecological parameters of southern barbel Barbus meridionalis R. in the
Rentina stream, Greece. J. Appl. IchthyoL, 3, 24-9.
Omeragic, N., Pocrnjic, Z. & Vukovic, T. (1980). Heat resistance of skeletal muscle of stream barbel Barbus meridionalis petenyi (abstract only). Acta Biol. Iugosl. (E. Ichtyol.),
12, 33-42.
Pagney, P. (1988). Climats et cours d'eau de France. In Collection gkographie. Masson, Paris, pp. 248.
Paris, P. (1932). Faune de la Sa6ne moyenne, 26me note (Cyclostomes et Poissons). Bull. Scient. Bourgogne, 2, 47 54.
Patt6e, E. (1988). Fish and their environment in large European river ecosystems - - the Rh6ne. Sciences de I'eau, 7,
35-74.
Pedroli, J.-C., Zaugg, B. & Kirchhofer, A. (1991). Atlas de
distribution des poissons et cyclostomes de Suisse. In Documenta Faunistica Helvetiae, Centre suisse de cartographie et
de la faune, pp. 207.
Penaz, M. & Zaki, M. I. (1985). Cyprinodont fishes of lake
Mariut, Egypt. Folia Zool. (Brno), 34, 373-84.
Perrin, J. F. (1988). Maintien en aquarium de l'apron du
Rh6ne Zingel asper (L.), esp6ce menaq6e d'extinction.
Revue fr. Aquariol., 15, 17-20.
Persat, H. (1988). De la biologie des populations de l'Ombre
commun Thymallus thymallus (L. 1758) ~i la dynamique des
communaut6s dans un hydrosyst6me fluvial am6nag6, le
Haut-Rh6ne francais~lements pour un changement
d'6chelle. Dissertation, Claude Bernard (Lyon I) University.
Persat, H. & Berrebi, P. (1990). Relative ages of present population of Barbus barbus and Barbus meridionalis
(Cyprinidae) in southern France: preliminary considerations. Aquat. Living Resour., 3, 253-63.
Peters, R. H. (1983). The ecological implication of body size.
In Cambridge studies in ecology, ed. E. Beck, H. J. B. Briks
& E. F. Connors. Cambridge University Press, Cambridge,
pp. 329.
Petit (1979). Migration des aloses dans le bas-Rh6ne: dispositif de franchissement des obstacles et possibilit6 de reproduction au niveau de l'am6nagement de Beaucaire. DAA,
Ecole Nationale Sup6rieur d'Agronomie de Rennes.
Philippart, J.-C. & Berrebi, P. (1990). Experimental hybridization of Barbus barbus and Barbus meridionalis: physiological, morphological, and genetic aspects. Aquat. Living
Resour., 3, 325-32.
Pocrnjic, Z. & Omeragic, N. (1976). Temperature tolerance of
muscular tissue of barbel Barbus meridionalis petenyi in
various condition of thermal acclimatation and thermal
adaptation (abstract only). Rev. Tray. Inst. P~ches Marit.,
40, 163-78.
Quignard, J. P. (1978). Le Rh6ne et quelques problOmes concernant ses poissons ou histoire naturelle de ee fleuve. La
Sabranenque, Bagnols-sur-C6se.
Rameye, L., Kiener, A., Spillmann, C.-P. & J. B. (1976). Aspects de la biologie de l'alose du Rh6ne-P~che et difficult6s
croissante de ses migrations. Bull. Fr. Piscic., 263, 50-76.
Raveret-Wattel, C. (1913). L'acclimatation du barbeau m6ridional dans le Sig (Oued M6kina), Alg6rie. Bull Soc. Acclim., 60, 11.
Roche, B. (1988). Bilan des premiers inventaires ichtyologiques du r6seau hydrographique de la Corse. Bull.
Ecol., 19(2 3), 235-45.
Roule, L. (1904). La faune ichtyologique des Pyr6n6es
franqaises et du Sud-Ouest de la France. Bull. Stn Piscie. &
Hydrobiol. Univ. Toulouse, 2, 30-9.
Roule, L. (1929). La thermophilie des aloses. In Les poissons
et le monde vivant des eaux douces. Delagrave, Paris, pp.
161-79.
Roule, L. (1933). Le peuplement des cours d'eau de la Corse
en poisson. Bull. Fr. Piscic., 63, 61-2.
Roule, L. (1935a). Les blennies potamiques peri-m6diterran6enne. Ver. Int. Vet. Limnol., 7, 411-4.
Roule, L. (1935b). La gen6se des faunes d'eau douce d'apr6s
l'exemple de la blennie fluviatile. Arch. Mus. Natnl. Hist.
Nat., Vol. tricentenaire, 6dine sdrie, 11,457-60.
Smirnov, A. I. (1971). Morphological description of the
danubian percid Aspro zingel (L.) in the lower Danube. J.
Ichthyol., 11,812-4.
Soric, V. & Jankovic, D. (1989). Characteristics of growth
and sexual maturity of Barbus meridionalis. Acta Biol. lugosl. (E. Ichtyol.), 21, 27 37.
Souchon, Y. & Trocherie, F. (1990). Technical aspects of
French legislation dealing with the freshwater fishery (june
1984): 'Fisheries orientation schemes' and 'Fishery resources management plans'. In EIFA C Symposium, ed. W.
L. T. Van Densen, B. Steinmetz & R. H. Hughes. G6tebourg, pp. 190-214.
Spillmann, C. J. (1961). Poisson d'eau douce. In Faune de
France. Paul Lechevalier, Paris, pp. 304.
Spillmann, J. (1959). Note pr61iminaire sur la syst6matique de
Telestes souffia Risso, poisson de la famille des Cyprinidae.
Bull. Mus. natn. Hist. nat. (2dine s&ie), 31,491-8.
Spillmann, J. (1960). Sur la syst6matique de Telestes souffia
Risso--26me note. Bull. Mus. hath. Hist. nat. (20me skrie),
32, 411 14.
Spillmann, J. (1962). Sur la syst6matique de Telestes souffia
Risso, variation de certains caract6res num6riques et
metriques de l'esp6ce-36me note. Bull. Mus. hath. Hist. nat.
(2dine s&ie), 34, 435 52.
Spillmann, J. (1965a). Sur la syst6matique de Telestes soufJia
Risso, 6tude d'un lot de poisson du Paillon (Alpes Maritimes)-56me note. Bull. Mus. natn. Hist. nat. (2dine s&ie),
37, 962-5.
Spillmann, J. (1965b). Sur la syst6matique de Telestes souffia
Risso. Variation d'un caract~re m6trique chez une souche
transplant6e-46me note. Bull. Mus. natn. Hist. nat. (2dine
skrie), 37, 760-3.
Spillmann, J. (1970). Apropos d'une population de poissons
de la famille des Cyprinidae Leuciscus (Telestes) souffia
Risso, provenant de la Droubie, affluent de l'H6rault--76me
note. Bull. Mus. natn. Hist. nat. (2~me s&ie), 42, 170-4.
Spillmann, J. (1974). Comportement d'une population de blageons, Leuciscus (Telestes) souffia (Poisson, Cyprinidae)
61ev6s en eau stagnante. Bull. Mus. natn. Hist. nat. ( 3Ome
s&ie), 170, 1241-5.
Taverny, C. (1991). P~che, biologie, 6cologie des aloses dans
le syst6me Gironde Garonne-Dordogne. In Etudes
Ressources en eau, No. 4. Centre National du Machinisme
Agricole du G6nie Rural, des Eaux et des For6ts (Bordeaux)/Bordeaux University 1, Bordeaux.
Terofal, F. (1987). Les poissons d'eau douce. In Collection lu
nature en couleurs. France Loisirs, Paris, pp. 287.
Thienemann, A. (1950). Verbreitungsgeschichte der Siisswassertierwelt Europas--Versuch einer historishen Tiergeographie der europdischen Binnengewdsser, No. 13. E.
Sehweiserbart'sche Verlagsbuchhandlung (Erwin N/~gele),
Stuttgart, pp. 809.
Trutna, R. & Mone, C. (1990). Oracle Software. Oracle, Redwood Shores, CA.
Vargas, M. J. & de Sostoa, A. (1991). Diet composition and
prey preference of Aphanius iberus and Gambusia affinis in
Ebro delta (Spain). In 7th Int. lchthyol. Congr. Eur. IchthyoL
Un. (26-30 August 1991). Bull. zool. Mus. Univ. Amsterdam, 88.
Status o f French riverine Mediterranean fish
Verneaux, J. (1973). Cours d'eaux de Franche-comt6,
recherches 6cologiques sur le r6seau hydrographique du
Doubs (massif du Jura), essai de biotypologie. Dissertation
Franche Comt6 (Besanqon) University.
Verneaux, J. (1981). Les poissons et la qualit6 des cours
d'eau. Ann. Scient. Univ. Franche-Comtk-Besanfon-Biol.
anim., 4, 33-41.
Villwock, W. (1970). Distribution, ecology and relationship
of Near East and Mediterranean cyprinodonts of the genus
Aphanius. J. ichtyol. CIESM, Rome, pp. 88-92.
Villwock, W. (1981). Distribution, ecology and intraspecific
variability of some external characters in Aphanius (Pisces:
Cyprinodontidae) - - a contribution to regressive evolution
in fish. Rapp. Comm. int. Met Medit., 27, 5.
Villwock, W. (1982). Supplement to Aphanius from the oasis
Azrag/Jordan and remarks on the taxonomy of Aphanius
iberus (Cyprinodontidae, Pisces) from oued Zousfana,
Igli/North West Algeria (in German). Mitt. Hamb. Zool.
Mus. Inst., 79, 267-71.
Vinyoles, D., de Sostoa, A., Casals, F. & Bianco, P. G.
157
(1991). The population of Blennius fluviatilis and its geographical distribution. In 7th Int. Ichthyol. Congr. Eur.
Ichthyol. Un. (26-30 August 1991). Bull. zool. Mus. Univ.
Amsterdam, 91.
Welcomme, R. L. (1985). River fisheries. FAO Tech. Pap,,
No. 262.
Wickler, W. (1957). Vergleichende Verhaltensstudien an
Grundfischen, I. Beitrage zur Biologie, besonder zur
Ethologie von Blenniusfluviatilis Asso 1801 im Vergleich zu
einigen anderen Bodenfischen. Z. Tierpsychol., 14, 393-428.
Zauner, G. (1988). Ecological requirement of Gymnocephalus
schraester (L. 1758), Zingel zingel (L. 1758) and Zingel streber (Siebold 1863) on the basis of the population in the
Danube impoundment of Altenw6rth (Austria). In 6th
Congr. Europ. Ichthyolog. Budapest.
Zylberblat, M., Solente, B., Le Hy, J. B. & Renoult, R.
(1991). Rapport de synth6se. Sch6ma de vocation piscicole
du fleuve Rhfne. Minist6re de l'Environnement, D616gation de Bassin Rh6ne M6diterran6e Corse, Service de la
Navigation Rh6ne Sa6ne.
A P P E N D I X 1 SPECIES LIST OF F R E S H W A T E R FISH IN THE
S T U D Y AREA.
Species
PETROMYZONIDAE
Lampetra fluvh~tilis (Linnaeus, 1758)
Lampetra planeri (Bloch, 1786)
Petromyzon marinus Linneaus, 1758
ACIPENSERIDAE
Acipenser sturio Linnaeus, 1758
ANGULLIDAE
Anguilla anguilla (Linneaus, 1758)
CLUPEIDAE
Alosa alosa (Linneaus, 1758)
Alosa fallax rhodanensis (Roule, 1924)
SALMONIDAE
Salmo trutta Linneaus, 1758
Salvelinus alpinus (Linneaus, 1758)
Salvelinus fontinalis (Mitchill, 1815)
Salvelinus namaycush (Walbaum, 1794)
Hucho hucho (Linneaus, 1758)
Oncorhynchus mykiss (Walbaum, 1792)
Coregonus sp (Linneaus, 1758)
Thymallus thymallus (Linneaus, 1758)
ESOCIDAE
Esox lucius Linneaus, 1758
CYPRINIDAE
Abramis brama (Linnaeus, 1758)
Alburnoides bipunctatus (Bloch, 1782)
Albumus alburnus (Linnaeus, 1758)
Barbus barbus (Linnaeus, 1758)
Barbus meridionalis Risso, 1826
Blicca bjoerkna (Linnaeus, 1758)
Carassius auratus (Linnaeus, 1758)
Carassius carassius (Linneaus, 1758)
Chondrostoma nasus (Linnaeus, 1758)
Chondrostoma toxostoma Vallot, 1836
Cyprinus carpio Linnaeus, 1758
Gobio gobio (Linnaeus, 1758)
Leueaspius delineatus (Heckel, 1843)
Leuciscus cephalus (Linnaeus, 1758)
Leuciscus leuciscus (Linnaeus, 1758)
Leuciscus souffia Risso, 1826
Abundance ~ Threat b
Status"
Range a
E
M
N
M
North-eastern Atlantic & Mediterranean
Northern Atlantic & Mediterranean
Extinct
Ex
M
CC
V
M
Northern Atlantic
R
R
E
V
M
E
Northern Atlantic
North-eastern Atlantic & Mediterranean
European
Northern Atlantic
North-Eastern American
Danubian
Norther-Western American
Holarctic
Holarctic
R
C
R
E
CC
R
C
R
R
CC
R
C
V
N
N
S
S
S
S
N
N
CC
V
N
Holarctic
N
N
N
N
E
N
A
A
C
E
A
N
A
N
European
European
European
European
Southern European
European
East Asian
Euro-Siberian
Central European
Southern European
Palaearctic
Holarctic
Central European
European
Euro-Siberian
Southern-European
CC
CC
CC
CC
CC
C
R
R
CC
C
CC
CC
R
CC
CC
CC
E
R
V
I
R
N
E
158
APPENDIX
CYPRINIDAE--eontd
Phoxinus phoxinus (Linnaeus, 1758)
Pseudorasbora parva (Schlegel, 1842)
Rhodeus amarus (Bloch, 1785)
Rutilus rutilus (Linnaeus, 1758)
Scardinius erythrophthalmus
(Linnaeus, 1758)
Tinca tmca (Linnaeus, 1758)
COBITIDAE
Cobitis taenia Linnaeus, 1758
Misgurnus fossilis (Linnaeus, 1758)
Noemacheilus barbatulus (Linnaeus, 1758)
SILURIDAE
Silurus glanis Linnaeus, 1758
PERCIDAE
Gymnocephalus cernua (Linnaeus, 1758)
Perca fluviatilis Linnaeus, 1758
Stizostedion lucioperca (Linnaeus, 1758)
Zingel asper (Linnaeus, 1758)
COTTIDAE
Cottus gobio Linneaus, 1758
CENTRARCHIDAE
Lepomis gibbosus (Linnaeus, 1758)
Micropterus salmoides (Lac6p6de, 1802)
GASTEROSTEIDAE
Gasterosteus aculeatus Linnaeus, 1758
Pungitius pungitius (Linneaus, 1758)
CYPRINODONTIDAE
Aphaniusfasciatus (Cuvier & Valenciennes, 1821)
POECILIIDAE
Gambusia affinis Baird & Girard, 1853
GADIDAE
Lota Iota (Linneaus, 1758)
ICTALURIDAE
Ictalurus melas (Rafinesque, 1820)
BLENNIIDAE
Blenniusfluviatilis Asso, 1801
1 --
contd
N
A
N
N
Holarctic
East Asian
Palaearctic
Euro-Siberian
N
N
European
Euro-Siberian
N
N
N
Holartic
European
Holarctic
R
C
Central European
C
CC
C
R
N
N
C
E
Euro-Siberian
Euro-Siberian
Central European
Rh6ne River
CC
N
European
CC
C
A
A
North American
C
R
N
N
Northern Atlantic
E
Circum-Mediterranean
A
N
Holarctic
A
E
Circum-Mediterranean
CC
R
C
CC
V
CC
CC
R
R
CC
R
E
E
E
R
C
C
V
C
R
V
aAbundance in three categories depending on number of occurrences in sections included in the database (R, _<30; C, 30-140; CC,
> 140).
bin accordance with IUCN rules according to Keith (1992) (Ex, species not recorded since 1961; E, species having disappeared
from a large part of their original range and whose numbers are reduced to a minimal threshold; these species are threatened with
extinction if the causes responsible for their current situation continue to act. V, species whose numbers are declining strongly because of unfavourable external factors; these species are likely to become E if the factors responsible for their vulnerability continue to act. R, species which are not immediately threatened with V or E, but whose populations are limited because of a
restricted geographical distribution, which exposes them to risks. I, species that may be E, V or R, but whose status is unclear
because of the lack of information).
"The origin of the various species according to Holcik's classification (1991) (N, natives; E, endemic, A, introduced by man; C,
introduced by means of inter-basin connections; S, maintained by artificial stocking).
aThe distribution of the species according to Banarescu's classifications (1964, 1989a,b).

current status of the riverine fishes of the french mediterranean basin

Transcription

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