Hydrobiologia 389: 183–191, 1998.

© 1998 Kluwer Academic Publishers. Printed in the Netherlands.

183

Oligochaete drift in a large river (French Upper Rhône):

the effect of life cycle and discharge

Bernard Cellot & Jacques Juget

ESA CNRS 5023 ‘Ecologie des Eaux Douces et des Grands Fleuves’ Université Claude Bernard, Lyon 1, 69622
Villeurbanne cedex, France. Fax: [+33] 4 72 43 11 41; E-mail: cellot@univ-lyon1.fr.

Received 20 April 1998; in revised form 16 November 1998; accepted 30 November 1998

Key words: drift, oligochaetes, spatio-temporal variation, large river

Abstract
In the framework of a macroinvertebrate drift study, the oligochaete community was analysed over one year (1989)
at six sites located in a partly impounded reach of the Upper Rhône River (France). Drift composition between sites
was somewhat affected by the environmental conditions. These differences were caused by significant deviations
in drift rates (P < 0.05) of endobenthic oligochaetes and non-swimming Naididae. There was no significant
difference (P > 0.05) among the swimming Naididae. As the latter strongly dominated, global spatial differences
in the drift were weak. Temporal changes constituted the main source of variability in drifting oligochaetes. Life
cycles and reproductive ecology of four major swimming Naidid species,Nais elinguis, Stylaria lacustris, Nais
christinae, and Nais barbata, explained most of the drift structure over the year. Hydrological fluctuations also
affected drift structure, both for swimming and non-swimming species. Taxon richness and relative abundance in
the drift were very similar to those obtained in the river benthos from a reference reach previously studied. There
was a lack of stygobiont and stygophilous species that are frequent in area of groundwater seepage. Biological
and environmental implications of oligochaete drift are stressed. The drift phenomenon is appropriated to provide
ecological informations on oligochaete communities in fast flowing and deep, large rivers.

Introduction family Naididae are favoured by the predominance

The composition, intensity and causes of invertebrate
drift in running waters have received considerable
attention, especially for arthropods; several biotic
and abiotic factors are involved in the phenomenon
(see reviews by Waters, 1972; Bournaud & Thibault,
1973; Statzner et al., 1984; and Brittain & Eikeland,
1988). Very few data are available for other groups.
Oligochaetes are rarely taken into account at a species
level and, as noticed by Dumnicka (1996), they are
even ignored in drift studies. However, the oligochaete
community may often represent an important part of
the benthic macrofauna in impounded rivers as well
as in pristine streams (e.g. Simpson et al., 1986; Os-
wood, 1989; Petto & Humpesch, 1992; Bournaud et
al., 1996; Koetsier & Bryan, 1996).
Among the oligochaetes living in lotic environ-
ment, specific richness and abundance within the
of coarse substratum, in contrast with other families
such as the Tubificidae (Learner et al., 1978). Nai-
dids are primarily linked to the biotecton (i.e. epilithon
and periphyton). They may thus contribute a higher
drift component than other oligochaete families that
include endobenthic and subterranean species more
able to resist hydraulic disturbances. Furthermore,
the Naididae comprise swimming and non-swimming
species. Sexual and asexual reproduction also co-
occur or succeed each other in their populations; the
proportion of these two types of reproduction gov-
erns productivity and survival rates of the species
concerned (see references in the reviews by Loden,
1981; Giere & Pfannkuche, 1982). Such aspects pre-
sumably could play a role in the drift of oligochaete
communities, and conversely, drift ought to study eco-
logical requirements for some oligochaete populations
in large rivers.
184
Figure 1. Map of the French Upper Rhône River
Brégnier-Cordon, and location of the six sampling sites.
Several studies on the macroinvertebrate fauna in
the Upper Rhône River (France) have shown that
oligochaetes were numerous in the benthos (e.g. Cel-
lot & Bournaud, 1986; Tachet et al., 1991; Cogérino
et al., 1995; Bournaud et al., 1996). Particular at-
tention was paid to spatial dynamics of the drift in a
recent work (Cellot, 1996). In the present paper, we re-
examine the data focusing on oligochaete community
only. The main objective of this investigation was the
study of spatial composition and temporal changes of
oligochaete drift throughout a year. We also compare
drifting oligochaetes with those sampled by dredging
in a previously studied neighbouring reach of this fast
flowing and deep large river.
Material and methods
The study was carried out between river km 88 and
98, about mid-way between Geneva and Lyon (Fig-
ure 1). This reach, called Brégnier-Cordon, is part
of a stony-sandy glacial basin surrounded by calcare-
ous mountains. It was originally characterized by the
presence of several side-channels but was impounded
in 1984 by a hydroelectric power plant (Roux et al.,
1989; Amoros, 1991). The impoundment regulated the
by-pass section and, now only two side-arms are still
connected to the main channel. One side-arm located
in the by-pass section is a fast-flowing anastomosed
channel (minimum mean velocity > 0.2 m s−1 ), also
permanently connected at its upstream end. The other
side-arm located in the Total Rhône section is a more
lentic braided channel connected with the Rhône itself
at its upstream end only during spates (discharge >
1100 m3 s−1 ); at low water level (the mean annual dis-
charge is 445 m3 s−1 over the period 1920–1990, data
from ‘Compagnie Nationale du Rhône’), this side-
arm is mainly supplied by seepage from the alluvial
aquifer. More details of the study area are given in
Cellot (1996).
Six sampling sites were chosen along the main
river axis (Figure 1): three in the by-pass section and
three downstream from the power plant in the Total
Rhône section. To test the influence of the two side-
arm systems, sites were located in the single mouth
of each side-arm (2, 5), and in the main channel
about 0.8 km upstream (1, 4) and downstream (3, 6)
at
from each confluence. All the sites were located along
concave shorelines so that they were scoured by the
main water flow. As it has been shown in river drift
studies that samples at mid-depth gave a mean value
of drift in the water column (Cellot, 1989a,b), only
one net was placed at mid-depth at each site. The
drift net used was based on the design of Burton &
Flannagan (1976) which provides a good compromise
between filtration efficiency and sorting time. It had
a mouth diameter of 0.3 m, a length of 1.4 m and a
500-µm mesh. The mouth of the net was fitted with
an aluminium cone with a 0.02 m2 opening, and a
flowmeter was placed within the cone to determine
the volume of water filtered. Drift samples were col-
lected only between sunset and sunrise. This night
period is not critical because no change in the diel drift
of the oligochaete had been observed (Cellot, 1989b;
see also Dumnicka, 1996). Sampling was performed
throughout 1989 about every ten days; however, a
spate in March was sampled more often (i.e. three
times a week).
A total of 216 drift samples was thus collected (6
sites × 36 dates). All the oligochaetes were sorted,
except for spate samples to which the 1/4 subsam-
pling technique of Chavanon & Bournaud (1980) was
applied. They were identified to species level when-
ever possible, and they were counted. Taxon richness
and drift densities (i.e. number of individuals per

100 m3 of filtered water) were calculated. A drift
rate (i.e. number of individuals per 10 h) were com-
puted and log-transformed for each sample. These
latter data were examined by means of a multivari-
ate analysis: centred Principal Component Analysis,
i.e. cPCA (Dolédec & Chessel, 1991). Ordination
methods provide factorial scores (i.e. new synthetic
variables) that may be used for mapping. In addition,
between-class cPCA was performed to know the part
of the total inertia (i.e. variability) of the data table
that was linked to either space or time (Dolédec &
Chessel, 1989, 1991). The 6 sites and the 36 dates
were thus used here as classes, respectively. This pro-
cedure takes into account the data cumulated by class
and introduces the principles of an analysis of vari-
ance (Lebreton et al., 1991). Analyses and graphics
were carried out using the ADE (Analyse de Don-
nées Ecologiques) software, a package for multivariate
analysis and graphical display for environmental data
(Thioulouse et al., 1995).
Results and discussion
Composition and differences between sites
In total, 32 taxa were collected in drift, including the
unidentified Enchytraeidae (see Appendix 1). As Nai-
dids dominated greatly in richness as well as in abun-
dance, three main categories of taxa of well-known
ecological significance were first considered:
(i) swimming Naididae (Dero digitata, Nais barbata,
Nais christinae, Nais elinguis, Nais pardalis, Nais
pseudobtusa, Nais simplex, and Stylaria lacustris),
(ii) non-swimming Naididae (including the 14 other
species of Naididae), and
(iii) endobenthic species or endobenthos (i.e. all other
taxa excluding the Naididae).
General parameters concerning the whole oligo-
chaete community and its composition according to
the previous categories are given as an annual aver-
age for each of the six sampled sites (Table 1). Based
on the amount of water sampled which largely varied
between sites, mean drift densities were much higher
in the side-arms (sites 2 and 5) but they were similar
at the main channel sites (1, 3, 4 and 6). The between-
site differences in both taxon richness and drift rate
showed a common pattern; high mean richness was
rather associated with high mean drift rate, and low
mean richness was associated with low mean drift rate
(site 5, where the volume of water filtered was both
185
lower and more stable). The coefficients of variation
of the means (from 53 to 66% for taxon richness,
and from 133 to 169% for drift rates) reflected similar
temporal variability whatever the site, despite a great
disparity in filtered water volumes between sites (from
20 to 139%). Based on the drift rates, the average pro-
portion of swimming Naididae was somewhat lower
at sites 1–3 (by-pass section) than at sites 4–6 (Total
Rhône section), and it was especially low at site 2
and high at site 5. This was clearly reversed for the
non-swimming Naididae as well as endobenthos.
Such spatial differences in drift composition were
statistically compared by Student’s t-tests performed
on taxon richness and log-transformed drift rates for
the three categories previously described (Table 2).
Significant differences were often found between two
sites at least one of each was a side-arm site (i.e.
2 or 5). They especially occurred at high levels of
significance for total Rhône side-arm (site 5) which
is the site most isolated from discharge fluctuations.
However, all these differences concerned only the
richness (taxa) and the drift rates of non-swimming
oligochaetes (non-swimming Naididae and endoben-
thos); in contrast, swimming Naididae showed no
significant difference whatever the site. Taking into ac-
count the disparity in the water volume filtered by nets
between sites (see Table 1), this last result suggests
a greater active drift (i.e. drift largely independent
of local hydraulic factors) of the swimming species
than of species unable to swim. Furthermore, the lack
of significant differences between sites located up-
stream and downstream from the side-arm confluences
(paired sites 1–3 and 4–6, Table 2), confirmed that
side-arm drift had no influence on oligochaete drift in
the main channel of this large river (Cellot, 1996). As
Naididae and particularly swimming ones dominated
the drift, differences between sites appeared globally
weak.
At a multivariate level, this was highlighted by
the results of the simple cPCA (Figure 2). On the
factorial map of the major axes (F1 and F2 which
explained 32 and 27% of the total variability, respec-
tively), the mean positions of the samples according to
the site were very close to the axis origin. Only the by-
pass section sites (1–3) were slightly separated from
the Total Rhône section sites (4–6). However, each
mean position of sites resulted from samples widely
dispersed as shown by the star graphs. In addition,
the inertia analysis (between-site and between-date
cPCA) showed that the spatial differences expressed
just only 8% of the total variabiltity of the data ta-
186
Table 1. Mean parameters (±SD expressed as a percentage of the mean, for n=36), and average relative
composition of drifting oligochaetes sampled at six sites during 1989 in the Brégnier-Cordon area, Upper
Rhône River

Site 1 2 3 4 5 6

Mean water volume 193 60 175 160 42 205

(m3 filtered by nets) (±71%) (±139%) (±66%) (±64%) (±20%) (±86%)

Mean drift density 44 159 61 47 97 50

(no. of ind. 100 m−3 ) (±165%) (±180%) (±181%) (±173%) (±162%) (±173%)

Mean taxon richness 5.0 6.9 6.6 5.3 3.2 5.7

(no. of taxa) (±62%) (±53%) (±57%) (±66%) (±61%) (±64%)

Mean drift rate 46 72 64 64 34 90

(no. of ind. 10 h−1 ) (±134%) (±133%) (±149%) (±169%) (±159%) (±166%)

Swimming Naididae 73.0 69.9 75.4 85.8 94.3 84.8

(average%)
Non-swimming Naididae 24.7 25.4 20.6 12.2 5.4 10.7
(average%)
Endobenthos (average%) 2.3 4.7 4.0 2.0 0.3 4.5

Table 2. Results of the Student’s t-test between six sites (paired

sites), on mean richness (Taxa) and log-transformed drift rates of
oligochaete groups (Swimming Naididae, Non-swimming Naidi-
dae, and Endobenthos) in the Brégnier-Cordon area, Upper Rhône
River

Paired Taxa Swimming Non- Endobenthos

sites Naididae swimming
Naididae

1–2 2.251∗ 0.980 2.008∗ 0.717

1–3 1.447 0.587 1.052 0.785
1–4 0.221 0.400 1.070 0.341
1–5 2.304∗ 0.598 4.242∗∗∗ 3.912∗∗∗
1–6 0.600 1.485 0.545 2.186∗
2–3 0.632 0.380 0.605 0.100
2–4 1.952 0.498 2.777∗∗ 0.948
2–5 4.946∗∗∗ 1.565 6.167∗∗∗ 3.364∗∗∗
2–6 1.557 0.620 2.182∗ 2.345∗
Figure 2. Factorial map of the samples (small squares) and mean
3–4 1.198 0.144 2.212∗ 1.028
position of the six sites (star graphs) on the first two axes (F1 and
3–5 3.784∗∗∗ 1.171 5.654∗∗∗ 3.811∗∗∗ F2, with the highest eigenvalues; see the window) from a centred
3–6 0.835 0.947 1.623 1.321 Principal Component Analysis (cPCA) of oligochaete drift over one
4–5 2.481∗ 0.951 3.283∗∗ 2.967∗∗ year in the Brégnier-Cordon area, Upper Rhône River.
4–6 0.369 1.019 0.508 1.243
5–6 2.898∗ 1.618 3.660∗∗∗ 4.979∗∗∗

Significant differences: ∗ P < 0.05, ∗∗ P < 0.01 and ∗∗∗ P < ble while the temporal differences accounted for more
0.001.
than 56%. Hence, the results emphasized mostly
temporal changes in drifting oligochaete community.
 187

Figure 3. Temporal changes over 1989 in mean (±SE) sample

scores by section on the first two factorial axes (F1 and F2) of a
centred Principal Component Analysis (cPCA) on oligochaete drift
collected in the Brégnier-Cordon area, Upper Rhône River. Avail-
able daily discharges in the two sections and river temperature (data
from the Compagnie Nationale du Rhône).

Temporal changes

Figure 3 presents the F1 and F2 mean scores of the

samples by section over the year, together with river
temperature, and mean daily discharges in the To-
tal Rhône section and in the by-pass section with a
pre-scheduled seasonal base flow. The mean scores
fluctuated very similarly between the two sections,
both on F1 and on F2. This clearly indicated that
main temporal changes in drifting community struc-
ture were independent of spatial aspect because of low
spatial differences.
The first axis was not related to the river dis-
charge (Figure 3). It opposed two periods in the year:
January–May (positive mean scores) from the other
months and especially summer months (negative mean
scores). This was largely due to four taxa of swim-
ming Naididae: Nais elinguis, Stylaria lacustris, Nais
christinae, Nais barbata. In average, there was an
evident substitution of dominance over the year be-
tween Nais elinguis and the other species concerned
Figure 4. Temporal changes over 1989 in mean (±SE) drift rate of
the main species discriminated: (a) on F1, the first axis of the cPCA;
and (b) on F2, the second axis of the cPCA (see Figure 3 and text).
Note the log-scales.
(Figure 4a). Nais elinguis strongly dominated in the
initial period (with 99% of its total drift rate from Jan-
uary to May) while Stylaria lacustris, Nais christinae
and Nais barbata dominated during the second period
of the year, and these species peaked at the time of
summer solstice. In contrast, the second axis appeared
linked to the river flow regime which was rather insta-
ble, particularly from March (the first spate sampled)
to June (Figure 3). There was a broad similarity be-
tween the curve of mean F2 scores and the hydrograph
of the Total Rhône section, especially. However, mean
188
Figure 5. Comparison between mean (+SE) drift rate and mean per-
centage of sexual individuals for Nais elinguis from March to May
1989 in the Brégnier-Cordon area ,Upper Rhône River. #= absence
of sexual individuals; + = first appearance of sexual individuals (as
a very low percentage).
daily discharge whatever the section was significantly
correlated with log-transformed mean total drift rates
(P < 0.05), and no significant relationship occurred
for taxon richness (P > 0.05). This ‘size effect’ was
mostly due to Nais elinguis, the dominant taxon (see
Figure 4a), and to several non-swimming species as
Nais bretscheri, Chaetogaster diaphanus, Nais com-
munis and Limnodrilus hoffmeisteri, that were also
discriminated on this second axis. Nais bretscheri
and Chaetogaster diaphanus, for instance, occurred
in higher mean numbers rather at the time of strong
discharge fluctuations (Figure 4b). Thus, the sea-
sonal drift of oligochaetes were partly explained by
hydrological conditions, as has usually been found
for other invertebrate groups (e.g. O’Hop & Wallace,
1983). If temporal changes appeared as a passive re-
sponse of both swimming and non-swimming species
to hydrological instability, they were primarily the
expression of seasonal dynamics of some swimming
Naididae such as Nais elinguis, Stylaria lacustris, Nais
christinae and Nais barbata.
The sexual or asexual type of the individuals was
determinated for these latter species. Spring is consid-
ered as a key period in the life cycle of Nais elinguis
in running waters (Learner et al., 1978; Paoletti &
Sambugar, 1984; Martinez-Ansemil, 1990; Juget &
Lafont, 1994). Stylaria lacustris, Nais christinae and
Nais barbata were very scarce during this period. The
study of Nais elinguis from March to May showed
(Figure 5) that spring drift was initially sustained by
asexual reproduction (accelerated paratomy) which
enhanced the drift rate of this species as a possible
result of the increase in benthic productivity. The ap-
pearance of a majority (> 50%) of sexual individuals
in mid-April corresponded precisely with low drift
rates (Figure 5), and these rates remained very low
during the rest of the year until December (see Fig-
ure 4a). The shift of asexual to sexual reproduction
entails cocoon production that allows survival during
environmental stress or adverse conditions (Learner
et al., 1978; Loden, 1981). Coincidence of active
drift of mature individuals of Nais elinguis with the
spring population explosion might be also explained
as dispersal strategy. Nais elinguis might therefore
be classified as species intolerant of summer-autumn
conditions in the Upper Rhône River. The presence
of a few sexual individuals of Stylaria lacustris (4
ind.) and Nais christinae (2 ind.) in autumn might
also be viewed as a survival strategy for these two
thermophilous species that may not tolerate cold win-
ter conditions. The same remark can apply for Nais
barbata, a species never found as sexually mature
individuals in this study but very capable of sex-
ual reproduction mainly in autumn according to the
literature data (Learner et al., 1978).
A drift-benthos comparison
The taxa and their relative abundances recorded in
the present drift study are given in Appendix 1. They
are compared with a reference bottom community that
had been collected in 1981–82 by sediment dredg-
ing across a similar impounded river reach, called
Jons, located 60 km downstream from the Brégnier-
Cordon area (Berly, 1989; Tachet et al., 1991). The
two reaches had relatively the same physical, hydro-
logical, and hydrochemical features (see Cellot, et al.,
1994; Bournaud et al., 1996).
Comparison between the two lists (Appendix 1)
showed that species number and abundance of Nai-
didae dominated the drift and the benthos (22 vs. 20
species, and 95 vs. 89% of total abundances, respec-
tively) in a similar manner. These results resemble
those obtained by Dumnicka (1996) during a simul-
taneous drift-benthos study in the impounded River
Raba, a tributary of River Vistula. Among the other
families, Propappus volki is interesting (Appendix

1). This species of a genus recently transferred from
the Enchytraeidae to the new family the Propappidae
(Coates, 1986), indicates active flow from under-
ground to surface waters (Lafont et al., 1992, 1996).
Propappus volki occurred in superficial stony-sandy
sediments of the Rhône as well as in other large Eu-
ropean rivers (Bird, 1982, who cited many records
from Central and Eastern Europe; Petto & Humpesch,
1992). Its absence from the drift paradoxically ex-
cludes it from the group of stygoxen species and
puts it in the same category as stygobious taxa (e.g.
species in the genera Trichodrilus, Dorydrilus, Rhya-
codrilus, Haber) frequently recorded from interstitial
hyporheic waters of the Upper Rhône River (Juget,
1984; Juget & Dumnicka, 1986) but absent from
the present material. In comparison with Propappus
volki, Stylodrilus lemani was among the most drifting
endobenthic taxa (Appendix 1); this Lumbriculidae
occurred usually in superficial sediments with lower
densities than Propappus volki and was never found in
hyporheic waters of the French Upper Rhône (Juget,
1984). Despite some other small divergences, there
was a good overlap in taxa identities between the two
lists (with 25 common taxa), and the Spearman rank
correlation gave a significant relationship between the
drift and the benthos (rs = 0.64, P < 0.001).
Finally, we are fully aware that these drift-benthos
data were not strickly comparable. They were com-
pared here because benthos data are very difficult to
obtain in fast flowing and deep, large rivers, which
become an overwhelming effort especially if only
one group of macroinvertebrates is concerned. Fur-
thermore, this comparison demonstrates the ability
of standard drift nets (i.e. those used to collect the
aquatic macrofauna with a low clogging effect) to
retain the oligochaetes, even the tiny species such
as Amphichaeta leydigii, Chaetogaster diastrophus,
Chaetogaster langi, Chaetogaster setosus (Appendix
1, individual body length of all these species found
in the drift were < 2mm), hence pointing out the
reliability of our drift sampling.
Conclusion
The very high predominance in the drift of Naididae
that principally include microdriles linked to biotec-
ton contrasted with the low occurrence of endobenthic
oligochaetes. The lack of hyporheic-interstitial fauna
in the drifting assemblage should be also emphasized.
Such results provides evidence of the specificity of the
189
drift method in describing the abilities of oligochaetes
to react to hydraulic disturbances affecting their en-
vironment. The absence from the drift of Propappus
volki, a species frequently recorded in coarse surface
sediments and groundwater flows, also tends to show
drift as a means of determining whether a species be-
longs to the stygoxen fauna (excluding stygobiont and
stygophilous species).
Drift rates indicated some ecophysiological char-
acteristics of certain species or groups. The swimming
Naididae were the least sensitive to hydraulic variation
(by-pass and Total Rhône sections) or habitat diversity
(main channel and more or less connected side-arms).
The capacity of naidids to swim is due to undulation
movements of their body associated with flexibility of
the hair setae. This mobility is high and well known
in Stylaria lacustris. It is probably the cause of drift in
this species as well as in other swimming species, such
as Nais elinguis which was by far the most dominant
taxa in the drifting oligochaete community. However,
drift dynamics depended primarily on the life cycles of
four naidid populations. The successive drift over the
year of Nais elinguis, then of Stylaria lacustris, Nais
christinae and Nais barbata, appears to be partly con-
sistent with literature data on the reproductive ecology
of benthic populations. Moreover, temporal changes
in the drift structure were also related to hydrolog-
ical fluctuations, and involved both swimming and
non-swimming species. Thus, biological and environ-
mental factors in oligochaete drift appear to be equally
important. Drift is an appropriate method to study the
ecological requirements of oligochaete populations of
fast flowing and deep, large rivers.
Acknowledgments
We especially thank M. Lafont and two anonymous re-
viewers for comments on the manuscript, J. C. Rostan
for important help in the field and E. Pattée for linguis-
tic advice. Financial support was provided by PIREN-
CNRS (Programme Interdisciplinaire de Recherches
sur l’Environnement).
190

Appendix 1. References

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Brégnier-Cordon (this study) compared with those collected from tions for river system management. Rivers 2: 105–112.
the benthos by dredging in a reference reach (Jons) of the Upper Berly, A., 1989. Distribution spatio-temporelle des peuplements
Rhône River, see text. - = not found. macrobenthiques prélevés par dragage dans une station du Haut-
Rhône. Doc. Thesis, Univ. Lyon 1, 307 pp.
Drift Benthos Bird, G.J., 1982. Distribution, life cycle and population dynamics
Taxa % % of the aquatic enchytraeid Propappus volki (Oligochaeta) in an
(of 14 (of 2755 English chalkstream. Holarct. Ecol. 5: 67–75.
801 ind.) ind.) Bournaud, M. & M. Thibault, 1973. Etude bibliographique. La
dérive des organismes dans les eaux courantes. Ann. Hydrobiol.
Naididae 4: 11–49.
Amphichaeta leydigii Tauber 0.20 – Bournaud, M., B. Cellot, P. Richoux & A. Berrahou, 1996. Macroin-
vertebrate community structure and environmental characteris-
Chaetogaster diaphanus (Gruithuisen) 3.30 1.63
tics along a large river: congruity of patterns for identification to
Chaetogaster diastrophus (Gruithuisen) 0.61 0.44 species or family. J. N. Am. Benthol. Soc. 15: 232–253.
Chaetogaster langi Bretscher 0.04 0.04 Brittain, J. E. & T. J. Eikeland, 1988. Invertebrate drift – A review.
Chaetogaster setosus Svetlov 0.08 2.21 Hydrobiologia 166: 77–93.
Dero digitata (Müller) 0.01 – Burton, W. & J. F. Flannagan, 1976. An improved river drift
sampler. Fisheries and Marine Service, Research Development,
Nais alpina Sperber 2.11 9.80
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Nais barbata Müller 4.73 1.67 Cellot, B., 1989a. Macroinvertebrate movements in a large Euro-
Nais behningi Michaelsen 0.72 3.37 pean river. Freshwat. Biol. 22: 45–55.
Nais bretscheri Michaelsen 5.52 37.05 Cellot, B., 1989b. Rythme nycthéméral et distribution verticale de la
Nais christinae Kasprzak 3.70 0.07 dérive des macroinvertébrés benthiques dans une grande rivière
européenne. Arch. Hydrobiol. 115: 265–286.
Nais communis Piguet 1.12 1.42
Cellot, B., 1996. Influence of side-arms on aquatic macroinverte-
Nais elinguis Müller 46.60 26.05 brate drift in the main channel of a large river. Freshwat. Biol.
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