Science of the Total Environment 454–455 (2013) 40–50

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Science of the Total Environment

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Pressures at larger spatial scales strongly influence the ecological status of heavily
modified river water bodies in Germany
Jochem Kail ⁎, Christian Wolter
Leibniz-Institute of Freshwater Ecology and Inland Fisheries, Müggelseedamm 310, 12587 Berlin, Germany

H I G H L I G H T S

• Importance of pressures in Heavily Modified Water Bodies (HMWB) was quantified.

• Large scale pressures are generally more important than reach scale habitat pressures.
• Fish metrics equally influenced by large scale and reach scale pressures
• Ecological status of macroinvertebrate mainly influenced by large scale pressures
• Large scale pressures must be considered for management and restoration of HMWB.

a r t i c l e i n f o a b s t r a c t

Article history: River biota are influenced by anthropogenic pressures that operate at different spatial scales. Understanding
Received 29 May 2012 which pressures at which spatial scales affect biota is essential to manage and restore degraded rivers. In Eu-
Received in revised form 14 February 2013 rope, many river reaches were designated as Heavily Modified Water Bodies (HMWB) according to the Euro-
Accepted 28 February 2013
pean Water Framework Directive (WFD), where the ecological potential might mainly be determined by
Available online 26 March 2013
pressures at larger spatial scales outside the HMWB (e.g. hydromorphological alterations at the river network
Keywords:
and land use at the catchment scale). In Germany, hydromorphological alterations and diffuse pollution were
EU-Water Framework Directive the main pressures. Therefore, the three objectives of this study were to (i) identify the hydromorphological
Fish pressures at the site, reach, and river network scale, and land use categories at the catchment scale which sig-
Macroinvertebrates nificantly affect the ecological status of HMWB in Germany, (ii) quantify the relative importance of these
Rehabilitation pressures at different spatial scales, and (iii) analyse the differences in response between fish and
Restoration macroinvertebrates. The results indicated that: (i) At the reach scale, fish were most strongly influenced by
Heavily Modified Water Bodies channel-bank conditions whilst the naturalness of channel-planform was the best proxy for the ecological
status of macroinvertebrates. At the catchment scale, urbanization was the most detrimental land use.
(ii) The pressures at larger spatial scales (catchment land use and hydromorphological alterations in the
river network) generally were more important than hydromorphological alterations at the reach scale.
(iii) Fish were affected equally by both, hydromorphological alterations at the reach scale and large-scale
pressures whereas the latter were far more important for the ecological status of macroinvertebrates. In con-
clusion, these results indicated that large-scale pressures may often limit the efficiency of reach-scale resto-
ration, especially for macroinvertebrates, even in the absence of saprobic pollution, and have to be considered
for the management and restoration of HMWB in Germany and comparable degraded river reaches.
© 2013 Elsevier B.V. All rights reserved.

1. Introduction
River ecosystems are influenced by natural controls and anthropo-
genic pressures that operate at different spatial scales (Hynes, 1975).
Pressures at large spatial scales, especially catchment land use, can be
more important in shaping invertebrate and fish communities com-
pared to pressures at smaller spatial scales (Allan et al., 1997; Black
⁎ Corresponding author. Tel.: +49 228 44 636 33; fax: +49 228 44 636 32.
E-mail addresses: jochem.kail@igb-berlin.de (J. Kail), wolter@igb-berlin.de
(C. Wolter).
et al., 2004; Hughes et al., 2008; Roth et al., 1996; Stephenson and
Morin, 2009). However, other studies showed that pressures at the
reach scale are of similar importance or that macroinvertebrates and
fish were even more strongly influenced by riparian land use and
instream habitat conditions at smaller, especially the reach scale (Feld
and Hering, 2007; Lammert and Allan, 1999; Moerke and Lamberti,
2006; Roy et al., 2003; Sponseller et al., 2001; Verdonschot, 2009). Be-
sides the catchment and reach scale, pressures at a third spatial scale –
the hydromorphological state of the adjacent river network up to a
few kilometres upstream of sampling sites – significantly affect the
local ecological status of macroinvertebrates (Kail and Hering, 2009).

0048-9697/$ – see front matter © 2013 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.scitotenv.2013.02.096
 J. Kail, C. Wolter / Science of the Total Environment 454–455 (2013) 40–50
Disentangling which pressures at which spatial scales significantly
affects stream biota and analysing differences in response to distur-
bances between biological groups is essential to manage and restore de-
graded rivers effectively. In near-natural rivers, macroinvertebrates'
composition equally depends on environmental variables at different
spatial scales (Death and Joy, 2004; Weigel et al., 2003), whereas
for fish, reach-scale variables were more important compared to
catchment-scale natural controls (Wang et al., 2003). Some studies
showed decreasing influence of reach-scale habitat conditions with in-
creasing catchment land use pressure (Wang et al., 2003). This shift in
importance might indicate that in a highly degraded environment,
large-scale pressures (e.g. hydromorphological alterations at the river
network and land use at the catchment scale) are more important for
macroinvertebrates whereas fish equally depend on pressures at differ-
ent spatial scales. As a consequence, in heavily degraded catchments,
especially macroinvertebrate assemblages might be mainly shaped by
pressures at larger spatial scales and thus, restoration projects which
are limited to the reach scale potentially are prone to failure if pressures
at larger scales are not adequately considered.
In Europe, the objectives of river management and restoration are
mainly set by the European Water Framework Directive (2000/60/
EEC of 22 December 2000; WFD) which requires that all EU waters
reach good ecological status (GES) at the latest by 2027. However,
river reaches can be designated as Heavily Modified Water Bodies
(HMWB) if applying the respective hydromorphological measures
to reach GES would significantly affect water uses (e.g. navigation,
flood protection, hydropower generation) that cannot be achieved
by other means which are technically feasible and not disproportion-
ately costly. The environmental objectives for HMWB can be lowered
to good ecological potential (GEP) which corresponds to the state
that results from applying all hydromorphological measures that do
not significantly affect these specified water uses. A substantial part
of the river water bodies in Europe (13.2%) has been designated as
HMWB, especially in Germany (39%) and other Central European
countries like Belgium (54%) and the Netherlands (94%) (European
Environment Agency, 2011).
Given the potential influence of pressures at larger spatial scales de-
scribed above, an essential question for HMWB management is to what
extent the ecological status depends on pressures at larger spatial scales
outside the HMWB that hardly can be addressed by reach scale restora-
tion measures. For two reasons, the most important pressures that have
to be considered for HMWB management are hydromorphological al-
terations at the reach and river network scale and diffuse pollution at
the catchment scale: First, saprobic pollution was no longer a main
pressure in German rivers (BMU, 2005). Second, regardless if the envi-
ronmental objective for HMWB is lowered due to hydromorphological
alterations that cannot be enhanced without having a significant nega-
tive effect on the specified water uses, HMWB have to reach a good
chemical status according to the WFD. Therefore, the pressures and spa-
tial scales considered in this study were hydromorphological alterations
at the site, reach, and river network scale, and land use at the catchment
scale as a proxy for diffuse pollution and hydrological alterations. The
three main objectives of this study were to (i) identify the specific
hydromorphological pressures at the site, reach, and river network
scale, and land use categories at the catchment scale which significantly
affect the ecological status of HMWB in Germany, (ii) quantify the
relative importance of these pressures at the different spatial
scales, and (iii) analyse differences in response to these pressures be-
tween two organism groups – fish and macroinvertebrates – and
stream types.
It was hypothesized that in HMWB (i) pressures at larger spatial
scales have a significant effect on the ecological status besides reach-
scale hydromorphology, (ii) the relative importance of large-scale pres-
sures is especially high for macroinvertebrates whereas fish might be
equally affected by pressures at different spatial scales, (iii) there are
stream-type-specific differences.
41
2. Methods
2.1. Study area and selection of water bodies
Three German federal states were selected as study area because
comparatively high-quality hydromorphological data were available
for large parts of the river network (Fig. 1). In Germany, 25 natural
stream types have been defined based on natural controls (e.g. geol-
ogy, catchment size, topography, altitude) and biota of natural refer-
ence sites (Pottgiesser and Sommerhäuser, 2008). Biological data
were available for degraded to heavily degraded streams and rivers,
covering most of the stream types that occurred in the study area
(15 out of 18) (Table 1). The macroinvertebrate sampling sites were
located in all three federal states (Fig. 1A), whereas the fish sampling
sites were all located in the federal state of Northrhine-Westphalia
(Fig. 1B). The dataset intentionally included HMWB and water bodies
in a moderate hydromorphological state only. Near-natural or refer-
ence sites were not considered since the objective was to assess the
effect of pressures and to discuss the possible effects of reach-scale
restoration projects in HMWB where not all hydromorphological
measures to reach GES can be applied and which cannot be restored
to near-natural conditions (see description in introduction, WFD Arti-
cle 4(3), CIS (2003) page 11).
2.2. Biological data
Data on 317 macroinvertebrate and 142 fish sampling sites, one in
each water body, were extracted from a database on WFD monitoring
data that was compiled by the University of Essen from the German fed-
eral states (Rolauffs et al., 2011). The samples were analysed using Euro-
pean standard methods developed for the WFD to assess the ecological
status because (i) the assessment methods are based on biological metrics
that describe community function and are better related to anthropogenic
pressures than species composition (Feld and Hering, 2007) which is
more dependent on regional biogeographic and historical differences as
well as regional species pools, and (ii) the metrics are part of the official
national assessment method for the ecological status of water bodies
and the success of restoration measures according to the WFD and
hence, are of special interest for river management. The few samples
not classified as good or high in the PERLODES module saprobic pollution
(http://www.fliessgewaesserbewertung.de/en/, accessed 01.05.2012)
were excluded for two reasons: First, HMWB have to reach good chemical
status according to the WFD. Second, organic pollution may mask the ef-
fect of other pressures like hydromorphological alterations and diffuse
pollution (Grosch et al., 2000; Kail et al., 2012).
All invertebrate sites were sampled and assessed with the PERLODES
method, which essentially corresponds to the AQEM method (Hering
et al., 2004). Sampling is based on a multi-habitat sampling strategy,
with a sample being composed of 20 spatially stratified sampling
units. The three main pressures considered in individual modules are
saprobic pollution, acidification, and “general degradation”. The module
general degradation is a multi-metric index, a continuous variable that
ranges from 1 (high ecological status) to 0 (bad ecological status), and
is calculated as weighted mean of 4–5 single core metrics that differ
depending on the stream type. In general, the core metrics used de-
scribe species composition and abundance, richness and diversity, func-
tional aspects, and the ratio between sensitive and tolerant taxa. The
latter is usually assessed by the German Fauna Index which is based
on a stream-type-specific list of indicator taxa for hydromorphological
degradation and makes up 50% of the final multi-metric index (Lorenz
et al., 2004). The core metrics in PERLODES are selected based on the
relations with hydromorphological degradation and land use impacts,
i.e. the module reflects the pressures investigated in this study, and
therefore, was used as the response variable for macroinvertebrates
(for more information see http://www.fliessgewaesserbewertung.de/
en/, accessed 01.05.2012).
42 J. Kail, C. Wolter / Science of the Total Environment 454–455 (2013) 40–50

A) B)

Fig. 1. Left: location of Germany (dark grey) in Europe. Right: two maps of the sampling sites in Germany. A: The macroinvertebrate (MIV) sites located in three federal states
(Northrhine-Westfalia in dark grey, Rhineland-Palatinate and Saxony in light grey). B: The fish sites solely located in Northrhine-Westfalia (NRW). Open symbols: sites only
used for grouping of stream types. Filled symbols: sites also used for analysis of pressures at multiple scales.

All fish samplings were conducted as single pass, boat or wading rivers, respectively, with an absolute maximum cumulative fishing
electrofishing (DC generator powered gears). The length fished varied length of 10 km in large streams (Dußling et al., 2004; Dußling,
with the wetted river width and the sampling method ranging from 2009). A further sampling requirement was a sufficient number of
40 to 100 times the average width in wadeable streams and boatable fish caught, which had to be at least 30 times the number of species

Table 1
German stream types occurring in the study area for which biological data were available. All stream types which show no significant differences in sample site communities
(ANOSIM, p > 0.15) and were grouped for further analysis are marked (M = macroinvertebrates, F = fish). Catchment sizes: streams = 10–100 km2, rivers = 100–10,000 km2.

German stream types (codes: names) Groups of stream types Number of MIV samples Number of fish samples

5: Small coarse substrate dominated siliceous highland riversMF Lower-mountain streams 43 23

5.1: Small fine substrate dominated siliceous highland rivers
6: Small fine substrate dominated calcareous highland riversF
7: Small coarse substrate dominated calcareous highland riversF

9: Mid-sized fine to coarse substrate dominated siliceous highland riversMF Lower-mountain rivers 76 23
9.1: Mid-sized fine to coarse substrate dominated calcareous highland riversF
9.2: Large highland riversMF

14: Small sand-dominated lowland riversMF Lowland and supra-regional streams 76 53

16: Small gravel-dominated lowland riversMF
18: Small loess and loam-dominated lowland riversMF
11: Small, organic substrate-dominated riversMF
19: Small streams in riverine floodplainsMF

15: Mid-sized and large sand and loam-dominated lowland riversMF Lowland rivers 33 23
17: Mid-sized and large gravel-dominated lowland rivers
12: Mid-sized and large organic substrate-dominated riversF
 J. Kail, C. Wolter / Science of the Total Environment 454–455 (2013) 40–50 43

in the reference fish community. The absolute minimum total number Kail and Hering (2009). Twenty-five parameters (Table 2) are recorded
of fish for any assessment is 101 (Dußling et al., 2004; Dußling, 2009). for 100 m channel segments and compared to natural conditions, not
The national assessment method for fish (fiBS) is reference based only for single reaches but continuously along large parts of the river
(Dußling et al., 2004). For each river system, unique reference fish com- network. For larger rivers, a slightly different set of parameters is
munities have been developed using historical data, reference sites, and mapped (Table 2) and the length of surveyed river segments depends
expert judgement (e.g., Wolter et al., 2005). These fish faunistic refer- on channel width ranging from 100 m (width 5–10 m) to 1 km
ences are compared to the actual fish assemblages obtained at the sam- (width > 160 m) (LAWA, 2002). Possible assessment results for each
pling sites. The quality criteria of the WFD have been translated into six parameter range from unchanged (only minor deviations from the refer-
metric groups (inventory of species and ecological guilds, abundance of ence conditions, class 1) to heavily degraded (class 7). The assessment re-
species and guilds, natural reproduction, migratory fish, fish region, and sults of the 25 parameters are aggregated to six indices, which are further
dominant species) with a total of 18 single metrics (Dußling et al., combined to three scores to assess the channel bed, banks, and floodplain,
2004). Metric scores range 5, 3, and 1 for the “high”, “good”, and “mod- and an overall assessment score, all by calculating the arithmetic mean
erate or worse” ecological status, respectively. (Table 2). Assessment results of parameters describing pressures (e.g.
In fiBS, there is no biological metric which was explicitly developed to bed-fixation) are only included if they do not improve the indices. Data
assess the effect of hydromorphological alterations and land use on fish. on the following 11 parameters, indices, and scores were available to de-
Therefore, the fiBS metrics related best to these anthropogenic pressures scribe the hydromorphological state and floodplain conditions (Table 2):
were identified in a preliminary analysis as described below (Section 2.5) planform parameter (which is a good proxy for the hydromorphological
and used as response variables for fish in this study. state, Kail and Hering, 2009), the six indices (channel pattern, longitudinal
profile, channel bed features, cross-section, channel bank features, flood-
2.3. Abiotic data on hydromorphological alterations and land use plain conditions), the three final assessment scores for channel bed, bank,
and floodplain, and the overall assessment score.
The hydromorphological data were compiled from regional author- The CORINE Land Cover 2000 (CLC2000) vector dataset was used to
ities and were recorded with methods essentially corresponding to describe the overall catchment characteristics and the potential influ-
Länderarbeitsgemeinschaft Wasser (LAWA, 2000) briefly described by ence of pressures like urbanization and agricultural land use. It is
based on satellite remote sensing images on a scale of 1:100,000 with
a grid size of 25 m and a minimum size of land-use polygons of 25 ha.
Table 2
Grouping of the parameters of the hydromorphological quality assessment for calculat-
The catchment upstream of the sampling sites was demarcated auto-
ing the six indices and the channel bed, banks, and floodplain scores. Parameters which matically and checked visually based on a digital elevation model
are only mapped for large rivers are marked with an asterisk. using GIS, and the percentage cover of the 33 CORINE land-use catego-
Channel bed
ries occurring in the sub-catchments was calculated and aggregated to
Channel pattern four more general categories with increasing land-use pressure
Planform (woody vegetation, extensive agriculture, intensive agriculture, urban
Erosion at bends areas) and an additional category “other” (Table 3) which covers b2%
Bars
of the area in 95% of the catchments and 4.8% at maximum.
Features indicating natural channel dynamics (e.g. large wood, islands,
widening)
Longitudinal profile 2.4. Spatial scales and pressures
Artificial barriers limiting continuity of flow, sediment and migration for biota
(e.g. weirs) Based on literature findings, hydromorphological alterations were
Culverts
Artificial impoundments
considered at the reach and river-network scale as well as land use
Riffles and steps at the catchment scale. Further sub-dividing the reach and river-
Flow-diversity network scale resulted in a total of 7 spatial scales that were investi-
Depth-variability gated in this study, which are numbered in the following in accor-
Flow diversion for hydropower*
dance with Fig. 2.
Channel bed features
Dominant substrate At the reach scale, it was further distinguished between (1) the site
Bed-fixation scale, i.e. the hydromorphological mapping section where a biological
Substrate-diversity sampling site was located, which was the smallest spatial resolution of
Bed features (e.g. scour and backwater pools, rapids, cascades) the hydromorphological data and where length depends on channel
Alteration of river bed (e.g. navigation, groins, sediment placement)*
width (compare to Section 2.3, Table 4), and (2) a longer reach up-
Channel banks and downstream of the biological sampling sites comprising several
Cross-section mapping sections and better describing the hydromorphological alter-
Cross-section form ations in the home range of mobile species like fish (Table 4). The length
Cross-section depth
of the reach scale was set to 100 times channel width, which corre-
Bank erosion (indicating widening of channel)
Cross-section width variability sponds to the upper limit of the fishing-section length in Dußling
Bridges (2009) (compare to Section 2.2, Table 4).
Widening/narrowing (artificial)* At the river-network scale, it was further distinguished between
Channel bank features the river network (including tributaries) (3) upstream and (4) down-
Riparian vegetation
stream of the fish sampling sites, whereas only the upstream river
Revetment/bank protection
Bank features (e.g. large wood, cutbanks) network was considered for the macroinvertebrate sampling sites
Alteration of river banks (e.g. hydropower peaking)* since Kail and Hering (2009) found no significant conditional effect
of the hydromorphological alterations in the downstream river net-
Floodplain
work on the ecological status of invertebrates. It was further distin-
Floodplain
Land-use guished between the up- and downstream river network just to the
Riparian buffer-strip next migration barrier (3a, 4a), describing the hydromorphological
Features hindering lateral channel migration (e.g. roads, dumping sites, fish- conditions in a section that can be reached and used as habitat by mo-
farms) bile species like fish, and including reaches up- and downstream of
Floodplain features (e.g. oxbow lakes, high-flow channels)*
existing migration barriers (3b, 4b) up to the maximum distance
44 J. Kail, C. Wolter / Science of the Total Environment 454–455 (2013) 40–50

Table 3 Table 4
Grouping of the 33 CORINE land-use categories to four more general categories and an Length of the river segments considered at the different spatial scales depending on
additional category “other”. Category name and CORINE category code and name are channel width.
given.
Channel width Site length Reach length Up- and downstream river
Urban (m) (km) (km) network length (km)
111 Continuous urban fabric
b5 0.1 0.5 2.5
112 Discontinuous urban fabric
5–10 0.1 0.7 2.5
121 Industrial or commercial units
10–20 0.2 1.5 5.0
122 Road and rail networks and associated land
20–40 0.5 3.0 7.5
123 Port areas
124 Airports
131 Mineral extraction sites
132 Dump sites
133 Construction sites arithmetic means are continuous variables ranging between 1 and 7.
141 Green urban areas In addition, the percentage of the impounded reaches was quantified
142 Sport and leisure facilities for one of the spatial scales (upstream river network 3b). At the
Intensive agriculture
(5) catchment scale, the percentage cover of the four land-use catego-
211 Non-irrigated arable land
221 Vineyards ries in the upstream catchment of the sampling sites were used as a
222 Fruit trees and berry plantations proxy for e.g. hydrological changes and fine sediment input to quan-
242 Complex cultivation patterns tify the land-use pressure.
Extensive agriculture
242 Complex cultivation patterns
231 Pastures 2.5. Preliminary analyses
243 Land principally occupied by agriculture, with significant areas of natural
vegetation
321 Natural grasslands (usually agriculturally used in study area, e.g. sheap Two preliminary analyses were conducted prior to the data analysis:
grazing) First, stream types were grouped based on the sample site communities.
322 Moors and heathland (usually agriculturally used in study area) Second, the fiBS fish metrics related best to the anthropogenic pressures
Woody vegetation investigated in this study were identified as mentioned in Section 2.2.
311 Broad-leaved forest
312 Coniferous forest
313 Mixed forest 2.5.1. Grouping of stream types
324 Transitional woodland-shrub
Other
A stream-type-specific analysis was conducted for two reasons:
331 Beaches, dunes, sands First, catchment land use, which is one of the pressures considered
332 Bare rocks in this study, is strongly constrained by and related to natural controls
333 Sparsely vegetated areas at the catchment scale (e.g. geology, climate) and hence, the effect of
411 Inland marshes
catchment land use on biota may be confounded with natural
412 Peat bogs
511 Water courses stream-type-specific differences (Allan and Johnson, 1997; Richards
512 Water bodies et al., 1996; Yates and Bailey, 2010). Second, it was hypothesized
521 Coastal lagoons that different stream types respond differentially to anthropogenic
522 Estuaries pressures. However, stream types could be pooled for further analysis
523 Sea and ocean
because natural stream-type-specific differences might vanish in de-
graded rivers and many of these anthropogenic changes must be con-
sidered irreversible at least in an engineering or management time
given in Table 4. As at the site and reach scale, the length of the up- frame. For example, past peat-cutting and decomposition due to
and downstream river network considered at these spatial scales draining has reduced the organic substrate of many peatland streams
depended on channel width (Table 4) and was set to 2.5 km for rivers (stream type 11, Table 1) which now are often very similar to
with a channel width b10 m based on empirical findings on the effect sand-bed streams (stream type 14, Table 1) even in the absence of
of the upstream river network on invertebrates in lower-mountain other land-use and hydromorphological pressures. All stream types
streams (Kail and Hering, 2009). The length of the river network which showed no significant differences in sample site communities
scale for rivers with a channel width > 10 m was based on expert (ANOSIM, p > 0.15) were grouped for further analysis which resulted
judgement. A mean score of the hydromorphological mapping sec- in four classes: Lower-mountain streams, lower-mountain rivers, low-
tions at these spatial scales (1, 2, 3a, 3b, 4a, 4b) was calculated for land streams, and lowland rivers (Table 1). These four classes have
each of the 11 hydromorphological variables, i.e. the resulting been separately treated and analysed for the macroinvertebrate and

Fig. 2. Schematic representation of the five spatial scales considered in the study.
 J. Kail, C. Wolter / Science of the Total Environment 454–455 (2013) 40–50 45

fish sampling sites, referred to as eight datasets in the following. Some
stream types showed significant differences (e.g. stream type 17:
gravel dominated lowland rivers) but had to be excluded due to a
low number of samples.
2.5.2. Selection of fish metrics
In a preliminary ordination analysis, the fiBS metrics related best
to the pressures investigated have been identified and selected as re-
sponse variables for fish in this study. For each of the four fish
datasets, only metrics were pre-selected which were measured on
an interval scale and where the data range included at least 5 differ-
ent values (Table 5). Since the length of the gradient in the Detrended
Correspondence Analyses (DCA) was ≪3 SD in all four fish datasets,
standardized Redundancy Analysis (RDA) was used following Braak
and Smilauer (2002). A selected set of 16 hydromorphological and
land-use variables was used to investigate the effect on the fiBS
metrics to avoid overfitting that occurs when the number of environ-
mental variables reaches or exceeds the number of samples. The 16
variables comprise the three final hydromorphological assessment
scores for channel bed, bank, and floodplain at the four spatial scales
1, 2, 3b, and 4b (12 variables) plus the four catchment land-use cate-
gories. Four metrics were selected for each fish dataset which were
related best to the anthropogenic pressures (variance explained by
the environmental variables being highest), as far as possible one
metric for each of the four fiBS metric groups (Table 5).
2.6. Data analysis
Some sampling sites had to be excluded since the distance to the
source was less than the length of the upstream river-network scale
3b (Fig. 2, Table 4). Nevertheless, it was possible to calculate all
hydromorphological and land-use variables for 228 macroinvertebrate
and 122 fish sampling sites, respectively (Table 1). All variables were
transformed to fit the normal distribution if necessary since this is an as-
sumption in linear regression analyses, and since the environmental
variables were measured in different units, they were standardized to
zero mean and unit variance to have equal weight in the ordination
analyses.
The statistical analysis had two main steps: a variable reduction sep-
arately at each spatial scale, and the analysis including the significant
variables of all spatial scales together. Firstly, the variable reduction
was performed separately for each spatial scale in each of the eight
datasets using the 11 hydromorphological variables at the reach and
river-network scales (1, 2, 3a, 4a, 4b, 12 variables for 3b) and the 4
land-use variables at the catchment scale. In the four invertebrate
datasets, the score of the PERLODES module general degradation was
used as a single response variable in multiple linear regression (MLR)
with forward stepwise selection. In each of the four fish datasets, the re-
spective fiBS metrics (Table 5) were used together as four response var-
iables in Redundancy Analyses (RDA). All environmental variables were
identified which had a significant conditional effect when included in
the statistical models (p b 0.05) using the F-test in MLR and a Monte
Carlo test with 499 permutations in RDA to test for significance. The
variance explained by these significant variables at each spatial scale
was quantified using adjusted r2 values for MLR and the sum of signifi-
cant conditional effects (sum of canonical eigenvalues) in RDA. The
river-network scales up to the next migration barrier (3a, 4a Fig. 2)
and including reaches up to the maximum distance given in Table 4
(3b, 4b Fig. 2) partly overlapped, and hence, the hydromorphological
state was similar and co-correlated in all datasets (t-test, p b 0.05),
except the two fish datasets in the lower-mountain and lowland riv-
ers. Therefore, only the river-network spatial scale better related to
the biological metrics (higher variance explained) was considered
for further analyses (3a or 3b, 4a or 4b, Fig. 2). Secondly, all signifi-
cant variables of each of the five spatial scales (1, 2, 3, 4, 5 Fig. 2)
were entered together in statistical models to investigate combina-
tions or interactions of anthropogenic pressures at different spatial
scales (shared variance) and the unique variance explained by each
spatial scale (partial correlation). Semi-partial correlation coeffi-
cients of MLR models and partial RDA were used to quantify the
unique and shared variance explained at each spatial scale on the
PERLODES score general degradation and the selected four fiBS fish
metrics, respectively. The total number of predictors at the five spa-
tial scales used in the eight models ranged between 3 and 10
(Table 6), depending on the number of variables that had a signifi-
cant conditional effect at each single spatial scale (p b 0.05).

Table 5
fiBS fish metrics in the four fish datasets which were measured on an interval scale and show a clear gradient in at least one of the four datasets. In fiBS, core species and
stream-type-specific species are distinguished, having a relative abundance of ≥5% and ≥1%, respectively in the site-specific reference assemblage. Moreover, the metrics related
best to the pressures and used in further analysis of the four fish datasets are marked (LMS = lower-mountain streams, LMR = lower-mountain rivers, LLS = lowland streams,
LLR = lowland rivers).

Metric Description LMS LMR LLS LLR

Inventory of species and guilds

CoreN Number of core species missing
CoreP Share of core species missing on total number of core species in reference
TypeN Number of stream type specific species missing x
TypeP Share of stream type specific species missing on total number of core species in reference x x
GuildPA Mean fiBS assessment result for presence/absence of all guilds (ranging from 1 to 5)

Relative abundance of species and guilds (R = deviation from reference conditions)

CoreAB(R) Core species
Rheo(R) Rheophilic species x x x x
Litho(R) Lithophilic species x
Psammo(R) Psammophilic species x
Phyto(R) Phytophilic species
Invert(R) Invertivore species
Omni(R) Omnivore species
GuildAB Mean fiBS assessment result for abundance of all guilds (ranging from 1 to 5)

Fish region
FRI Fish Region Index (mean indicator value of the sampled species for fish region, presence of potamal fish results in high values for FRI) x x x x

Dominant species
CSI Core Species Index (number of core species with relative abundance >5% in sample divided by total number of core species in reference) x x x
CDI Community Dominance Index (sum of relative abundance of the two most abundant fish species)
46 J. Kail, C. Wolter / Science of the Total Environment 454–455 (2013) 40–50

Table 6
Mean values and coefficients of variation (CV) of all variables with a significant conditional effect (p b 0.05) (bold) in separate statistical models for each spatial scale in the eight
datasets (LMS = lower-mountain streams, LMR = lower-mountain rivers, LLS = lowland streams, LLR = lowland rivers). In addition, in the first line of each spatial scale, the total
variance explained by the variables that have a significant conditional effect at the respective spatial scale is given. Only the underlined variables were included in the statistical
models where all spatial scales were considered.

Variable Description Fish datasets Macroinvertebrate datasets

Each first line: sum of conditional effects (%) Each first line: adjusted r2 (%)
Following lines: mean (CV%) Following lines: mean (CV%)

LMS LMR LLS LLR LMS LMR LLS LLR

1 Site scale 57 29 9 11 9 0 17 55
S_BANK Bank score 4.4 (31) 4.7 (32) 5.0 (29) 5.2 (25) 5.3 (28) 4.8 (28) 5.3 (24) 5.3 (24)
S_FLOOD Floodplain score 5.3 (24) 5.4 (20) 5.6 (20) 5.2 (29) 6.0 (16) 5.5 (30) 5.7 (16) 5.4 (26)
S_IBED Channel bed features index 4.0 (35) 3.7 (38) 4.6 (27) 5.0 (23) 4.5 (30) 4.3 (33) 4.8 (28) 5.0 (21)
S_IPLAN Planform index 4.7 (34) 4.2 (42) 5.7 (20) 5.4 (26) 5.3 (21) 4.5 (35) 5.8 (17) 5.5 (20)

2 Reach scale 46 25 8 15 9 0 16 46
R_BANK Bank score 4.4 (27) 4.8 (26) 5.0 (27) 5.0 (25) 5.3 (22) 4.8 (23) 5.1 (23) 5.3 (24)
R_FLOOD Floodplain score 5.3 (24) 5.3 (18) 5.5 (17) 5.1 (30) 6.0 (14) 5.3 (24) 5.5 (14) 5.3 (25)
R_IBED Channel bed features index 4.0 (31) 3.9 (29) 4.4 (26) 4.9 (23) 4.5 (26) 4.3 (27) 4.7 (27) 5.0 (22)
R_ICROSS Cross-section index 4.3 (27) 4.5 (29) 5.0 (26) 5.0 (23) 5.0 (27) 4.8 (25) 5.0 (23) 5.0 (26)
R_IPLAN Planform index 4.5 (31) 4.3 (31) 5.7 (19) 5.2 (27) 5.3 (15) 4.4 (31) 5.7 (16) 5.4 (20)

3 Upstream river network scale

3a Only up to next migration barrier 26 22 12 24 19 6 17 18

UPD_IBED Channel bed features index 4.0 (34) 3.8 (27) 4.4 (23) 4.9 (20) 4.3 (31) 4.2 (27) 4.5 (26) 5.1 (19)
UPD_ICROSS Cross-section index 4.2 (30) 4.5 (27) 4.9 (24) 4.9 (22) 4.8 (31) 4.9 (26) 4.9 (25) 5.0 (22)
UPD_IPLAN Planform index 4.4 (32) 4.3 (31) 5.6 (18) 5.4 (18) 5.1 (17) 4.4 (33) 5.6 (17) 5.3 (16)

3b Whole upstream river network scale 25 19 9 22 27 12 26 52

UP_IPROF Longitudinal profile index 4.5 (29) 4.8 (19) 5.0 (22) 5.4 (16) 4.6 (23) 4.8 (20) 5.1 (19) 5.7 (15)
UP_IBED Channel bed features index 4.2 (29) 4.3 (19) 4.4 (24) 5.1 (19) 4.4 (28) 4.2 (24) 4.6 (24) 4.9 (20)
UP_ICROSS Cross-section index 4.3 (27) 4.7 (15) 4.9 (24) 5.0 (18) 4.8 (28) 4.7 (20) 5.0 (22) 4.9 (22)
UP_IPLAN Planform index 4.7 (24) 4.6 (18) 5.6 (18) 5.4 (19) 5.1 (16) 4.4 (27) 5.7 (16) 5.3 (15)
UP_IMP % impounded reaches 10.1 (171) 18.7 (90) 7.2 (247) 16.6 (163) 4.4 (189) 16.0 (126) 5.8 (258) 16.5 (162)

4 Downstream river network scale

4a Only down to next migration barrier 24 19 20 0

DD_IPAT Channel pattern index 4.7 (25) 4.9 (25) 5.4 (20) 5.3 (20)
DD_IBED Channel bed features index 4.3 (31) 4.2 (23) 4.6 (20) 4.9 (21)

4b Whole downstream river network scale 0 0 16 14

D_IBED Channel bed features index 4.3 (26) 4.2 (24) 4.6 (20) 4.9 (21)

5 Catchment scale 30 18 5 11 28 29 0 12
URBAN % urban land use 10.5 (73) 12.8 (68) 6.7 (142) 9.3 (44) 9.4 (104) 8.8 (68) 10.8 (110) 7.1 (34)
IAGRI % intensive agriculture 52.3 (65) 26.4 (91) 72.4 (23) 72.6 (14) 13.6 (120) 22.3 (80) 70.8 (24) 72.9 (15)
EAGRI % extensive agriculture 12.3 (166) 21.4 (68) 8.2 (78) 7.3 (105) 22.8 (72) 24.3 (54) 6.3 (81) 8.0 (85)
FOREST % forest 24.9 (93) 38.8 (49) 12.7 (108) 10.5 (42) 53.8 (42) 43.9 (34) 12.0 (91) 11.7 (59)

3. Results
3.1. Identification of the most important hydromorphological variables
and land use categories at the five spatial scales
The hydromorphological variables and catchment land-use catego-
ries which best explained the ecological status differed between the
five spatial scales and two organism groups. At the site and reach
scale, fish metrics were most strongly influenced by bank conditions
(S_BANK, R_BANK) whereas the naturalness of channel planform
(S_IPLAN) was the best predictor for the ecological status of inverte-
brates at the site scale (Table 6). There were no such clear differences
between the two organism groups at the upstream river-network
scale, where different hydromorphological variables had a significant
conditional effect on the fish and invertebrate metrics. For the fish met-
rics, the hydromorphological conditions in the river network just up-
and down to the next migration barrier (3a, 4a, Fig. 2) were a better pre-
dictor compared to the state in the whole river network (3b, 4b, Fig. 2),
with 4–33% higher explained variance for the upstream river network
and having the only significant effect and 25% higher explained variance
in 3 out of 4 fish datasets in the downstream river network (Table 6). In
contrast, for macroinvertebrates the hydromorphological state of the
upstream river network including impounded reaches upstream of mi-
gration barriers (3b, Fig. 2) was a markedly better predictor for the eco-
logical status compared to scale 3a (42–100% higher explained variance
in single scale regression models) (Table 6). Catchment land use, espe-
cially % urban area, had a significant effect on fish metrics in all four
datasets, and agricultural land use was an additional good predictor
for the ecological status of invertebrates in the lower-mountain region.
In contrast, catchment land use was related weakly or not significantly
to the PERLODES score in the lowlands.
The absolute variance of the biological metrics explained by the
anthropogenic pressures investigated markedly differed between
the eight datasets (Fig. 3). The environmental variables investigated
explained about half (43–59%) up to 75% of the variance in 5 out of
8 datasets. In three datasets (lowland-stream fish and inverts and

Fig. 3. Relative variance explained by the five different spatial scales on fish and invertebrate metrics and shared variance of the site/reach and catchment/upstream scale. The
absolute variance explained by all spatial scales is given above the stacked bars. Spatial scales with a significant conditional effect using forward stepwise selection are marked
with an asterisk (p b 0.05).
lower-mountain river inverts) the explained absolute variance was
lower (about one third) indicating that other, not measured variables
more strongly influenced the ecological status here.
3.2. Relative importance of pressures at the five spatial scales and
differences between biological groups and stream types
The pressures at larger spatial scales (catchment land use and
hydromorphological alterations in the river network) had a significant
effect on the ecological status in 6 out of 8 datasets, whereas local pres-
sures (hydromorphological alterations at the site and reach scale) only
in 3 out of 8 datasets (see asterisks in Fig. 3). To quantify the importance
of hydromorphological alterations and land-use pressure at the five
spatial scales investigated relative to each other, the relative variance
explained at the different spatial scales was calculated (Fig. 3).
The relative importance of the pressures at the five spatial scales dif-
fered between fish and invertebrates (Fig. 3). For fish, the unique vari-
ance explained by hydromorphological alterations at the site and reach
scale was about one third (27–34%) and similar to the unique variance
explained by pressures at larger spatial scales (hydromorphological
PERLODES score module general degradation
(ecological status of macroinvertebrates)
1.0
0.8
0.6
0.4
0.2
0.0
0 5 10 15
Share of urban land-use in upstream catchment (%)
Fig. 4. Scatterplot of the ecological status of macroinvertebrates vs. the share of urban
land use in the upstream catchment in the lower-mountain rivers dataset. The plan-
form index ranging from 1 (high) to 7 (bad) and the 90% quantile regression line
and equation are given.
J. Kail, C. Wolter / Science of the Total Environment 454–455 (2013) 40–50 47
alterations at the river-network scale and catchment land use, 22–
35%). In contrast, for macroinvertebrates the unique variance explained
by pressures at larger spatial scales (39–100%) was much higher com-
pared to those at the site and reach scale (0–8%). Two datasets showed
the reverse pattern: for lowland-stream fish anthropogenic pressures
at the site and reach scale were of low importance, whilst for
lowland-river macroinvertebrates, the unique variance explained at
the site/reach scale was much higher compared to the other
macroinvertebrate datasets (Fig. 3).
Stream types strongly differed with respect to the relative impor-
tance of the spatial scales (Fig. 3). In particular, the lower-mountain
river macroinvertebrate dataset showed a unique pattern since
none of the variables at the site and reach scale had a significant effect
on the ecological status even when the spatial scales were considered
in separate regression models (Table 6).
The only parameters with a significant conditional effect in this
dataset were the share of impounded reaches in the upstream river net-
work (UP_IMP) and the share of urban land-use in the upstream catch-
ment (URBAN). The latter showed a clear wedge-shaped relation with
the ecological status (Fig. 4), which is typical for variables that act as
limiting factor or bottleneck. Even sites with a natural channel planform
(high planform index S_IPLAN) had a bad ecological status (PERLODES
score b 0.2). The best ecological status that could be expected at a
Planform index given level of urbanization was estimated based on the linear 90%
at sampling sites
(1 high to 7 bad) quantile regression line. Sample sites with a share of urban land use in
1
2 the upstream catchment larger than about 15% did not reach good eco-
3 logical status (PERLODES score > 0.6).
4
5
y = 1.025 - 0.028 x 6
4. Discussion
7
The first objective of the study was to identify the specific
hydromorphological variables at the site, reach, and river network
scale, and land use categories at the catchment scale which most strongly
affect the ecological status of fish and invertebrates in HMWB.
At the site and reach scale, the ecological status of fish was most
strongly related to the habitat conditions at the river banks, indicating
20 25 30 that rehabilitating river banks may be an appropriate measure for fish
in HMWB. However, this might have been an artefact due to the stan-
dard sampling protocol using electric fishing because fish are caught
most efficiently at littoral sites providing cover (Cowx, 1991), and
cover like macrophytes, large wood, overhanging terrestrial vegeta-
tion, and undercut banks is predominantly found at near-natural
48 J. Kail, C. Wolter / Science of the Total Environment 454–455 (2013) 40–50
river banks. For the ecological status of macroinvertebrates, river
planform was the best proxy, which is consistent with the results of
Kail and Hering (2009).
At the river network scale, the ecological status of fish was affected
more strongly by hydromorphological alterations up- and down-
stream just to the next migration barrier (3a, 4a Fig. 2) compared to
the whole river network scales (3b, 4b, Fig. 2), which seemed reason-
able since fish are principally mobile and use habitats at larger spatial
scales but only as far as they are connected longitudinally. These dif-
ferences potentially are higher (i) if a substantial part of the river net-
work considered at the spatial scales 3 and 4 is located behind the
migration barriers, and (ii) if the hydromorphological state at the
two spatial scales differs. This did hold true for the upstream river net-
work scale 3 in most fish datasets, where the river network considered
at scale 3b was 1.8 to 3.8 times longer compared to scale 3a, and hence,
the hydromorphological state at the two spatial scales was not or only
weakly correlated (r 2 b 0.21). In contrast, in the downstream river
network, scale 4b and 4a largely overlapped (4b only 1.0 to 1.9 times
longer than 4a) and the hydromorphological states were strongly
correlated (r 2 0.64 to 0.94). However, the four fish datasets showed
a reverse pattern and the differences were more pronounced in the
downstream and relatively small in the upstream river network
(Table 6), which indicates that the ecological status of fish is also af-
fected by the river reaches upstream of the migration barriers besides
the habitat conditions up to the dams.
In contrast to fish, macroinvertebrates were affected more strongly
by hydromorphological alterations of the upstream river network in-
cluding reaches upstream of migration barriers (3b Fig. 2). However,
the length of the impounded reaches in the upstream river network
only had a significant effect (p b 0.05) in 1 out of 4 datasets (p =
0.39–0.49 in other datasets), indicating that the impoundments them-
selves may be of varying importance and not necessarily the most im-
portant pressure in the upstream river network. There are several
other possible reasons for the higher influence of scale 3b on
macroinvertebrates: The poor hydromorphological state of the up-
stream river network might affect the input of organic matter such as
leaves and large wood from riparian areas (Moerke and Lamberti,
2006), cause changes of physico-chemical variables like water tempera-
ture (Nelson and Palmer, 2007) or decrease the re-colonization potential
due to missing source populations (Brown and Swan, 2010). However, it
is not possible to directly infer causal relationships from the statistical
analyses. The hydromorphological state could also be a proxy for land
use in the riparian zones upstream and its detrimental effects on the
stream ecosystem like the input of nutrients, fine sediment, and pesti-
cides caused by agriculture (Omernik et al., 1981; Weijters et al.,
2009). Studies which consider both, the hydromorphological state and
the adjacent land use in buffers along the river network could help to
disentangle the effect of these two pressures at the river-network scale.
At the catchment scale, urban land cover had a significant effect on
the ecological status of HMWB in most of the datasets (6 out of 8),
which is consistent with many other studies showing that urbaniza-
tion is the most detrimental catchment land use for fish and
macroinvertebrates (Allan et al., 1997; Black et al., 2004; Hughes
et al., 2008; Roth et al., 1996; Stephenson and Morin, 2009). Urban
land use is a surrogate rather than a pressure per se, and it might affect
ecological status through different pathways, e.g. water quality or
quantity (Burcher et al., 2007). Although sampling sites affected by
saprobic pollution have been excluded from this study, other pollut-
ants like toxic substances might have impaired water quality. More-
over, impervious cover and changed runoff conditions increase
peak-discharges, flushing frequencies and hydraulic stress (Archer
et al., 2010). Increased peak flows are known to impact biota negative-
ly (Coleman et al., 2011), and changes in discharge conditions may
even limit invertebrate assemblages (Konrad et al., 2008). Finally,
urban land use in the upstream catchment may be a proxy for pres-
sures at smaller spatial scales like the hydromorphological alterations
or buffer land use along the river network. However, pressures at both
spatial scales (upstream catchment and upstream river network) had a
significant conditional effect in 3 out of 8 datasets (Fig. 3), indicating
that they were not just co-correlated and affected the ecological status
of the HMWB through different pathways. More extensive datasets
would be needed to identify the key pressures which have to be targeted
by river restoration and management, including data on a larger set of
pressures for which urban land served as proxy.
The second objective was to assess the relative importance of
hydromorphological alterations at the site, reach, and river network
scale as well as catchment land use for the ecological status of
HMWB. As hypothesized, the pressures at larger spatial scales, espe-
cially catchment land use, had a significant conditional effect on the
ecological status of HMWB in most datasets (6 out of 8). In the two
lowland-river macroinvertebrate datasets, the missing effect of
urban and agricultural land use in the catchment possibly resulted
from the small differences between sampling sites and corresponding
low coefficients of variation, which were b34% for agricultural land
use in both datasets and in addition for urban land use in the
lowland-river dataset compared to 68–120% in the lower-mountain
datasets (Table 6). In contrast, the hydromorphological alterations at
the site and reach scale had no significant conditional effect in 5 out of
8 datasets. This might have been simply due to differences in the varia-
bility of the predictors at the different spatial scales, since the strength
of statistical relationships also depends on the length of the gradients,
which was potentially low at the site and reach scale due to the exclu-
sion of near-natural sampling sites. However, the coefficients of varia-
tion of the hydromorphological variables at the site and reach scale
even were significantly higher compared to the variables at the river
network scales (t-test, p b 0.01, n = 168, Table 6). These results indi-
cate that pressures at larger spatial scales like catchment land use and
the hydromorphological alterations in the up- and downstream river
network have to be considered for the management of HMWB in
Germany and comparable rivers in Central Europe.
The importance of the river network scale depends on the length of
the HMWB since the shorter the HMWB, the higher is the share of river
reaches which are influenced by an adjacent up- and downstream river
network that is located outside the HMWB. For example, given the
smallest length of the river network scale used in this study (2.5 km),
the ecological status of all river reaches in HMWB being b 5 km in
length, and a substantial part (50%) of HMWB 10 km in length is
influenced by the river network outside the HMWB. Therefore, espe-
cially short HMWB, which comprised a large part of the HMWB investi-
gated in this study (32% b 5 km, 66% b 10 km), cannot be managed
and restored without considering adjacent water bodies. Rather than
implementing measures in the short HMWB, it is probably much
more efficient to improve the ecological status of the adjacent water
bodies which influence the HMWB and define its ecological potential.
These results on the relative importance should be confirmed by
more extensive studies for three reasons. First, a substantial part of
the explained variance in the eight datasets (about one third to
half) could not be uniquely attributed to one of the spatial scales, sim-
ilar to other studies where the shared variance was 8–34% (Weigel
et al., 2003), 30% (Moerke and Lamberti, 2006), and 28–50% (Feld
and Hering, 2007), and hence it is not possible to finally assess the
relative importance of the pressures at the different spatial scales.
Second, the importance of the pressures at different spatial scales rel-
ative to each other was investigated in this study. The absolute vari-
ance explained by the pressures investigated, and hence, their
absolute importance was low (about 1/3) in 3 out of 8 datasets. This
might have been due to confounding factors like fish stocking or
methodological reasons like the study design (short gradient), the
metrics used, and the heterogeneous data originating from different
regional authorities, which potentially resulted in higher variability
and scatter. Moreover, it could be that other important pressures
not considered in this study affected the ecological status in these
 J. Kail, C. Wolter / Science of the Total Environment 454–455 (2013) 40–50
three datasets. Third, three of the fish datasets were comparably small
(n = 23) since samples were grouped by stream types to investigate
stream-type specific differences. However, overfitting did not occur
and the number of samples was 2.1 to 10.6 times the number of pres-
sure variables. These values were similar to the share of samples and
environmental variables in comparable studies, ranging from 1.4 to
15.7 (Black et al., 2004; Feld and Hering, 2007; Hughes et al., 2008;
Moerke and Lamberti, 2006; Verdonschot, 2009; Weigel et al.,
2003). Nevertheless, regional peculiarities and outliers potentially
have a marked influence on the ordination results.
The third objective was to investigate possible differences be-
tween the relative importance of pressures at different spatial scales
in HMWB for the ecological status of fish and macroinvertebrates.
As hypothesized, in the degraded to heavily degraded water bodies
investigated in this study, the pressures at larger spatial scales
(hydromorphological alterations at the river-network scale and
catchment land use) were far more important than local habitat
conditions for macroinvertebrates, whereas the ecological status
of fish equally depended on the hydromorphological alterations at
the site and reach scale. In less degraded or near-natural rivers,
macroinvertebrates' composition was found to depend equally on en-
vironmental variables at different spatial scales (Death and Joy, 2004;
Feld and Hering, 2007; Weigel et al., 2003), whereas for fish,
reach-scale variables were more important compared to catchment-
scale natural controls (Wang et al., 2003). These differences between
studies in less degraded or natural vs. the results of this study in highly
degraded rivers support the hypothesis of Wang et al. (2003) that the
importance of pressures at the catchment scale increases in disturbed
rivers. This indicates that for fish, restoration measures to improve the
hydromorphological state at the reach scale that are applicable in
HMWB could be of similar importance to measures at larger spatial
scales, but they probably would have only very minor effects on the eco-
logical status of macroinvertebrates, even if the water bodies reach
good chemical status in respect to saprobic pollution.
The results of this study indicate that pressures at larger spatial
scales (catchment land use and hydromorphological alterations in the
upstream river network) are of similar importance for the ecological
status of fish, and far more important for macroinvertebrates compared
to reach-scale hydromorphological alterations in German HMWB. The
high importance of catchment land use and the hydromorphological al-
terations in the upstream river network might have been one of the rea-
sons for the small effect of reach-scale hydromorphological restoration
measures on macroinvertebrates observed in German rivers (Jähnig
et al., 2010) and other countries (Palmer et al., 2010). This study
assessed the relative importance of different pressures at different spa-
tial scales. Even though it was not possible to infer causal relationships
from these analyses, important conclusions for river management can
be drawn. The results indicated that the ecological potential of short
HMWB b 5 km in length is also defined by adjacent water bodies and
it is probably much more efficient to implement restoration measures
outside these HMWB without affecting the uses within the HMWB.
More extensive studies are needed to identify the causal pressures con-
sidering a larger set of variables to describe the pressures in more detail
(e.g. with data on water quality parameters like the concentration of
toxic substances, pesticides, and nutrients, fine sediment loads, hydrol-
ogy and hydraulics) and larger sample sizes. In addition, the main
limiting pressures could be identified and recommendations for suc-
cessful restoration projects derived by experimental approaches and
by coupling different models to investigate the effect of pressures at dif-
ferent spatial scales on biota (Kiesel et al., 2009).
Acknowledgments
This study was funded as part of the iwrm-net project “FORECASTER”
by the German Federal Ministry for Education and Research (grant
number 02WM1031).
49
References
Allan JD, Johnson LB. Catchment-scale analysis of aquatic ecosystems. Freshw Biol
1997;37:107–11.
Allan JD, Erickson DL, Fay J. The influence of catchment land use on stream integrity
across multiple spatial scales. Freshw Biol 1997;37:149–61.
Archer DR, Climent-Soler D, Holman IP. Changes in discharge rise and fall rates applied
to impact assessment of catchment land use. Hydrol Res 2010;41:13–26.
Black RW, Munn MD, Plotnikoff RW. Using macroinvertebrates to identify biota-land
cover optima at multiple scales in the Pacific Northwest, USA. J N Am Benthol
Soc 2004;23:340–62.
BMU (Bundesministerium für Umwelt, Naturschutz und Reaktorsicherheit). Die
Wasserrahmenrichtlinie — Ergebnisse der Bestandsaufnahme 2004 in Deutschland.
http://www.umweltbundesamt.de/wasser/veroeffentlich/kostenlos.htm. 2005. [Last
access on 18th September 2012].
Braak CJF ter, Smilauer P. CANOCO Reference Manual and CanoDraw for Windows
User's Guide: Software for Canonical Community Ordination (version 4.5); 2002.
Brown BL, Swan CM. Dendritic network structure constrains metacommunity proper-
ties in riverine ecosystems. J Anim Ecol 2010;79:571–80.
Burcher CL, Valett HM, Benfield EF. The land-cover cascade: relationships coupling land
and water. Ecology 2007;88:228–42.
CIS. Identification and designation of heavily modified and artificial water bodies. Common
Implementation Strategy for the Water Framework Directive (2000/60/EC), Guidance
Document No. 4. Office for Official Publications of the European Communities; 2003.
Coleman JC, Miller MC, Mink FL. Hydrologic disturbance reduces biological integrity in
urban streams. Environ Monit Assess 2011;172:663–87.
Cowx IG. Catch effort sampling strategies: their application in freshwater fisheries
management. Oxford: Blackwell Scientific Publications; 1991.
Death RG, Joy MK. Invertebrate community structure in streams of the Manawatu-
Wanganui region, New Zealand: the roles of catchment versus reach scale influ-
ences. Freshw Biol 2004;49:982–97.
Dußling U. Handbuch zu fiBS. Schriftenr Verb Dtsch Fischereiverwaltungsbeamter
Fischereiwiss 2009;15:1-59.
Dußling U, Berg R, Klinger H, Wolter C. Assessing the ecological status of river systems
using fish assemblages. In: Steinberg C, Calmano W, Klapper H, Wilken RD, editors.
Handbuch Angewandte Limnologie VIII. Landsberg: Ecomed Verlagsgruppe; 2004.
p. 1-84.
European Environment Agency. Hydromorphology — zero draft for EEA 2012 state of
water assessment. EEA activity 1.4.3, Version: Zero draft, date: 22.11.2011, pre-
pared by Peter Kristensen and the European Topic Centre Inland, Coastal, Marine
Waters; 2011.
Feld C, Hering D. Community structure of function: effects of environmental stress on
benthic macroinvertebrates at different spatial scales. Freshw Biol 2007;52:
1380–99.
Grosch UA, Rennert B, Hilge V. Development and use of surface waters, and the fate of the
related fisheries in the Berlin area of Germany. Fish Manag Ecol 2000;7:179–88.
Hering D, Moog O, Sandin L, Verdonschot PFM. Overview and application of the AQEM
assessment system. Hydrobiologia 2004;516:1-20.
Hughes SJ, Ferreira T, Cortes RV. Hierarchical spatial patterns and drivers of change in
bentic macroinvertebrate communities in an intermittent Mediterranean river.
Aquat Conserv 2008;18:742–60.
Hynes HBN. The stream and its valley. Verh Proc Trav SIL 1975;19:1-15.
Jähnig SC, Brabec K, Buffagni A, Erba S, Lorenz AW, Ofenböck T, et al. A comparative
analysis of restoration measures and their effects on hydromorphology and
enthic invertebrates in 26 central and southern European rivers. J Appl Ecol
2010;47:671–80.
Kail J, Hering D. The influence of adjacent stream reaches on the local ecological status
of Central European mountain streams. River Res Appl 2009;25:537–50.
Kail J, Arle J, Jähnig SC. Limiting factors and thresholds for macroinvertebrate assemblages
in European rivers: Empirical evidence from three datasets on water quality, catch-
ment urbanization, and river restoration. Ecol Indic 2012;18:63–72.
Kiesel J, Hering D, Schmalz B, Fohrer N. A transdisciplinary approach for modelling
macro-invertebrate habitats in lowland streams. IAHS Publ 2009;328:24–33.
Konrad CP, Brasher MD, May JT. Assessing streamflow characteristics as limiting factors
on benthic invertebrate assemblages in streams across the western United States.
Freshw Biol 2008;53:1983–98.
Lammert M, Allan JD. Assessing biotic integrity of streams: effects of scale in measuring
the influence of land use/cover and habitat structure on fish and macroinvertebrates.
Environ Audit 1999;23:257–70.
LAWA. Gewässerstrukturgütebewertung in der Bundesrepublik Deutschland.
Verfahren für kleine und mittelgroße Fließgewässer. Schwerin (Germany):
Länderarbeitsgemeinschaft Wasser; 2000.
LAWA. Gewässerstrukturkartierung in der Bundesrepublik Deutschland. Verfahren für
mittelgroße bis große Fließgewässer. Schwerin (Germany): Länderarbeitsgemeinschaft
Wasser; 2002.
Lorenz A, Hering D, Feld C, Rolauffs P. A new method for assessing the impact of
hydromorphological degradation on the macroinvertebrate fauna of five German
stream types. Hydrobiologia 2004;516:107–27.
Moerke AH, Lamberti GA. Scale-dependent influences on water quality, habitat, and
fish communities in streams of the Kalamoazoo River Basin, Michigan (USA).
Aquat Sci 2006;68:193–205.
Nelson KC, Palmer MA. Stream temperature surges under urbanization and climate
change: data, models, and responses. J Am Water Resour Assoc 2007;43:440–52.
Omernik JM, Abernathy AR, Male LM. Stream nutrient levels and proximity of agricul-
tural and forest land to streams: some relationships. J Soil Water Conserv 1981;36:
227–31.
50 J. Kail, C. Wolter / Science of the Total Environment 454–455 (2013) 40–50
Palmer MA, Menninger H, Bernhardt ES. River restoration, habitat heterogeneity and
biodiversity: a failure of theory or practice? Freshw Biol 2010;55:205–22.
Pottgiesser T, Sommerhäuser M. Beschreibung und Bewertung der deutschen
Fließgewässertypen — Steckbriefe und Anhang. available from http://www.
wasserblick.net/servlet/is/18727/?highlight=flie%DFgew%E4ssertypen. 2008.
Richards C, Johnson LB, Host GE. Landscape-scale influences on stream habitat and
biota. Can J Fish Aquat Sci 1996;53(Suppl. 1):295–311.
Rolauffs P, Meier C, Hering D, Böhmer J, Schaumburg J, Schranz C, et al.
Weiterentwicklung biologischer Untersuchungsverfahren zur kohärenten Umsetzung
der EG-Wasserrahmenrichtlinie. Umweltbundesamt Project Report grant number
3707 28 201. Essen (Germany): University of Duisburg-Essen, Faculty of Biology,
Aquatic Ecology; 2011.
Roth NE, Allan JD, Erickson DL. Landscape influences on stream biotic integrity assessed
at multiple spatial scales. Landscape Ecol 1996;11:141–56.
Roy AH, Rosemund AD, Paul MJ, Leigh DS, Wallace JB. Stream macroinvertebrate re-
sponse to catchment urbanisation (Georgia, U.S.A.). Freshw Biol 2003;48:329–46.
Sponseller RA, Benfield EF, Valett HM. Relationships between land use, spatial scale and
stream macroinvertebrate communities. Freshw Biol 2001;46:1409–24.
Stephenson JM, Morin A. Covariation of stream community structure and biomass of
algae, invertebrates and fish with forest cover at multiple spatial scales. Freshw
Biol 2009;54:2139–54.
Verdonschot PFM. Impact of hydromorphology and spatial scale on macroinvertebrate
assembalge composition in streams. Integr Environ Assess Manag 2009;5:97-109.
Wang L, Lyons J, Rasmussen P, Seelbach P, Simon T, Wiley M, et al. Watershed, reach,
and riparian influences on stream fish assemblages in the Northern Lakes and
Forest Ecoregion, U.S.A. Can J Fish Aquat Sci 2003;60:491–505.
Weigel BM, Wang LZ, Rasmussen PW, Butcher JT, Stewart PM, Simon TP, et al. Relative
influence of variables at multiple spatial scales on stream macroinvertebrates in
the Northern Lakes and forest ecoregion, USA. Freshw Biol 2003;48:1440–61.
Weijters MJ, Janse JH, Alkemade R, Verhoeven JTA. Quantifying the effect of catchment
land use and water nutrient concentrations on freshwater river and stream
biodiversity. Aquat Conserv 2009;19:104–12.
Wolter C, Bischoff A, Wysujack K. The use of historical data to characterize
fish-faunistic reference conditions for large lowland rivers in northern Germany.
Arch Hydrobiol 2005;155:37–51. [Suppl.].
Yates AD, Bailey RC. Covarying patterns of macroinvertebrate and fish assemblages
along natural and human activity gradients: implications for bioassessment.
Hydrobiologia 2010;637:87-100.