Ecology of Freshwater Fish 2006: 15: 96103 Printed in Singapore All rights reserved

2006 The Authors Journal compilation 2006 Blackwell Munksgaard

ECOLOGY OF FRESHWATER FISH

Fish assemblages as inuenced by environmental factors in streams in protected areas of the Czech Republic
ka K. Fish assemblages as inuenced by environmental Humpl M, Pivnic factors in streams in protected areas of the Czech Republic. Ecology of Freshwater Fish 2006: 15: 96103. Blackwell Munksgaard, 2006 Abstract Three streams of comparable size located in different landscape-protected areas were selected for studying the effect of environmental factors on sh assemblages using indirect (detrended correspondence analysis, DCA) and direct (canonical correspondence analysis, CCA) gradient analysis. DCA of species showed well a gradient of assemblage changes in the longitudinal prole. DCA of sites stressed the variability between the sh assemblages of the three streams. This pattern was then conrmed by the highly signicant between-stream CCA. In the within-site CCA, environmental factors explained 50.7% variability for presenceabsence data and 58.3% for the relative abundance data. The analysis revealed that number of ponds and land use are the most inuential factors of the strongest environmental gradient. However, in the partial CCAs, factor substratum type explained the largest proportion of the variability affecting sh in their habitat choice. Generally, presence absence and relative abundance data of sh gave similar results in both DCA and CCA analyses; the same environmental factors proved to be important in both data type analyses. The environmental factors explain more variability than the regional (between-stream) one. The total proportion of variability explained by the presenceabsence data analysis was 71.9% and in the relative abundance analysis even 80.8%. The environmental factors measured during the eld survey explain 2.1- and 3.4-times more assemblages variability than factors measured from a hydrological map.
ka M. Humpl, K. Pivnic
Institute for Environmental Studies, Faculty of Science, Charles University, Prague, Czech Republic

Key words: mid-sized streams; presence absence and relative abundance data; factors measured in situ and on maps; detrended correspondence analysis; canonical correspondence analysis M. Humpl, Institute for Environmental Studies, tska Faculty of Science, Charles University, Bena 2, 128 01, Prague 2, Czech Republic; e-mail: humpl@email.cz Accepted for publication November 14, 2005

Un resumen en espan ol se incluye detra s del texto principal de este art culo.
Introduction

Investigations into seminatural stream ecosystems are still more frequently oriented to the assessment of the relationships between species and the environment (Angermeier & Davideanu 2004). Gradient changes in longitudinal proles of streams are commonly observed together with gradients of environmental factors (Schlosser 1982; Oberdorff et al. 1993, 2001; Lyons 1996; Belanger 1997; Angermeier & Winston 1999; Marsh-Matthews & Matthews 2000; Moyle et al. 2003). 96

Variability in sh assemblages is generally inuenced by landscape features (Angermeier & Winston 1999), by the stream size or elevation (Schlosser 1982; Lyons 1996; Brown 2000) or by stocking of economically important shes (the brown trout, grayling, carp, ka & Humpl 2004). and pike) (Pivnic Both indirect and direct multivariate gradient analyses are now frequently used to identify and explain how environmental gradients affect the distribution of shes (Godinho et al. 2000; Ero s et al. 2003; Moyle et al. 2003) and their simultaneous use has often yielded the most appropriate results (Maret doi: 10.1111/j.1600-0633.2006.00126.x

Fish assemblages and environmental factors et al. 1997; Angermeier & Winston 1999; Ero s et al. 2003). The statistical robustness of these methods has milauer 2002). been evaluated (ter Braak & S Headwater reaches are highly variable, while more downstream environmental conditions, if not anthropogenically inuenced, are generally less extreme (Ostrand & Wilde 2002). Evaluation of the impact of environmental variables on sh in Protected Areas (PA), which have little human inuence, seems therefore more simple. However, construction of ponds, management with salmonids in headwaters or with carp more downstream cannot be overlooked. In this paper we analyse sh assemblages in three icka na streams (the Tepla , Luz Nisa and Ostruz rivers) located in different Protected Areas where human inuence is minimised by law. We hypothesise that: (i) ponds and stream management do not signicantly change the sh assemblages; (ii) using presenceabsence or relative abundance data of sh and measuring the environmental factors directly in situ or from a hydrological map, inuences the variability of sh assemblages equally.
Material and methods Study sites
Table 1. The basic information on the streams. Tepla Stream source altitude (m.a.s.l) Stream mouth altitude (m.a.s.l) Stream length (km) Mean stream slope (mkm)1) Mean ow in the stream mouth (m3s)1) Higher order stream Sites shed 784 370 64 6.47 3.1 e Ohr 13 Nisa icka Luz 765 235 55 9.64 5.4 Odra 8 na Ostruz 938 452 39 12.46 1.2 Otava 10

na 20022003, on the Ostruz in 20012002 and on the icka Luz Nisa in 2000. In total, 3312 specimens of 18 sh species were caught in Tepla , 1283 specimens of icka 15 sh species in Luz Nisa and 1848 specimens of na nine sh species in Ostruz River. Fished sections were minimally 100 m long. In all cases sh were collected by at least a three-person team using the Honda GX 120 backpack electroshing gear (1.8 kVA; Radomir Bendar, Olomouc, Czech Republic). The stream sections were shed by wading in two passes with comparable effort (3040 min). The sh relative abundance (number of caught sh per 100 m2) is given in Table 2.
Data analysis

Study sites were located on three mid-sized (four to ve order) streams in PA the Tepla River in PA icka Slavkov Forest, the Luz Nisa River in PA Jizera umava. na Mountains, and the Ostruz River in PA S They are situated in the West, North and Southwest Bohemia respectively (Fig. 1). The basic information on the streams studied is in Table 1. The major parts of the streams pass through highlands with prevailing forest stands and permanent grasslands. Human activities are minimised in these watersheds.
Sampling regime

In many cases, only the species presenceabsence data are available when temporal variability of older and recent data is to be evaluated. Thus it is very helpful to know the differences of assemblage patterns when the presenceabsence and/or the relative abundance data are used. The effect of extreme values was minimised by transforming the species relative abundance data by log (x + 1) and the inuence of rare species by the evaluation only of species found at least twice. Altogether 19 species were used in the analyses (Table 3).
Table 2. The sh relative abundance expressed as the number of caught sh per 100 m2. Site no. 1 2 3 4 5 6 7 8 9 10 11 12 13 Tepla 16.80 4.64 46.66 45.01 15.26 12.56 8.55 5.66 8.21 18.65 18.69 3.80 2.13 S 2 4 5 11 12 3 5 5 5 9 8 7 6 Nisa icka Luz 22.81 5.24 7.72 8.09 1.29 4.39 7.69 10.99 S 1 5 6 6 4 6 8 7 na Ostruz 44.17 16.07 30.38 30.83 31.30 17.89 30.52 33.62 26.23 18.70 S 3 2 4 3 4 5 7 6 4 4

Thirty-one sites regularly distributed in longitudinal proles were selected to cover all the important habitats. The sites on the Tepla were sampled in

Fig. 1. Map of Czech Republic showing the location of studied streams.

The sites are arranged according to their increasing distance from the source (S, the total number of species).

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 ka Humpl & Pivnic

Table 3. The list of sh species, their scientic and common names and the abbreviations used. Common Abbreviation name Anan Babt Caca Cogo Cyca Eslu Gogo Gyce Lapl Lece Lele Lolo Onmy Pe Ruru Satr Safo Thth Titi European eel Stone loach Crucian carp Bullhead Carp Pike Gudgeon Ruffe Brook lamprey Chub Dace Burbot Rainbow trout Perch Roach Brown trout Brook trout Grayling Tench icka Luz Nisa na Tepla Ostrunz x x x x x x x x x x x x x x x x x x x

Scientic name Anguilla anguilla L. Barbatula barbatula L. Carassius carassius L. Cottus gobio L. Cyprinus carpio L. Esox lucius L. Gobio gobio L. Gymnocephalus cernuus L. Lampetra planeri Bloch 1784 Leuciscus cephalus L. Leuciscus leuciscus L. Lota lota L. Oncorhynchus mykiss Walbaum 1792 Perca uviatilis L. Rutilus rutilus L. Salmo trutta m. fario L. Salvelinus fontinalis Mitchill 1814 Thymallus thymallus L. Tinca tinca L.

x x x x x x x x x x x x x x x x x x

x x

We used an indirect ordination method of gradient analysis (detrended correspondence analysis, DCA) with the aim to identify the strongest gradient of assemblage composition independently of the environmental variables we measured. Based on the DCA sites ordination we also assessed between-stream differences in the sh assemblages. The direct gradient canonical correspondence analysis (CCA) was then used for the evaluation of the variability in assemblage structure in relation to the selected environmental factors. The regional variability of assemblages (between-stream variability) was accounted for in the rst CCA. Then in the second CCA (within-stream), the inuence of the measured environmental factors was shown, while the effect of between-stream variability was partialled out (eliminated). Step-wise forward selection of environmental variables available in CANOCO was used to choose the most inuential factors. The individual effect of each signicant environmental factor was detected in partial CCAs. In these analyses the residual effect of each individual factor was tested, while the effects of the other signicant factors were (together with the between-stream variability) partialled out as covariables. Multivariate statistical analyses were performed milauer using CANOCO version 4.5 (ter Braak & S 2002); for the correlation analysis STATISTICA software was used (STATSOFT Inc., Tulsa, OK, USA). Fifteen environmental factors measured both from a hydrological map of the Czech Republic (1:50,000) and in situ during the eld surveys, were available for the study. To reduce the high number of variables, 98

relationships among all the environmental factors were rst tested by correlation analysis using Spearmans rank order correlation coefcients (rs). Only the factors with rs < 0.3 (a < 0.05) were retained in the analyses. Four factors were eliminated (catchment area, stream order, number of settlements in the watershed and maximal depth). Eleven environmental factors were nally used for the analyses. Four of them were measured from a map, the other seven during eld surveys (Table 4). The number of ponds was counted in the catchment up to a distance of 5 km above the given site. We dened a pond as an articial reservoir of minimally one to maximally hundreds of hectares, fed mostly by a small stream. Ponds are used for rearing carp, together with tench and pike. In some ponds several species of coarse sh (gudgeon, perch, roach, stone loach, crucian carp and minnow) occur and can invade into streams, increasing their diversity. Mean stream width was the average of measurements made in three different positions situated in the beginning, middle and at the end of each shed section (ve times each). Mean depth was measured at the same positions, at each position ve measurements equally distributed in a diagonal prole were made. Current velocity was the average stream current velocity in the site measured using a simple oat. When evaluating the channel type we differentiated among natural channel (1), channel with adjusted banks (0.5) and completely regulated channel (0). The quality of the cover was assessed according to the presence or absence of the following: boulders and scoured banks (1), scoured banks only (0.5) and none (0). Substratum type is a ve-state variable which describes proportions (%) of all
Table 4. Environmental factors measured from map and during eld survey in situ. Environmental variable Map Distance from the source Elevation Stream slope Number of ponds Mean channel width Mean depth Current velocity Channel type Cover Substratum type Unit/categories km m.a.s.l. mkm)1 Value m m m s) 1 1/0.5/0 1/0.5/0 %Boulders %Stones %Gravel %Sand %Silt Permanent grassland Permanent grassland with trees Forest Abbreviation used km Elevation Slope Ponds WIDTH DEPTH VELOCITY CHANNEL COVER %BO %ST %GR %SA %SI PG PG+T

In situ

Land use

Fish assemblages and environmental factors individual bottom substratum categories on each site. Land use was classied as a three-state categorical variable containing permanent grassland (1,0,0), permanent grassland with trees (0,1,0) or forest (0,0,1).
Results Fish assemblage gradients (DCA)

Presenceabsence data The rst two ordination axes of DCA explain 35.2% variability of sh assemblages. On the rst axis (20.5%) the species sequence from left to right reected the gradient of assemblage change in the longitudinal prole. The second axis gradient (14.7%) reects the variability caused by unidentied environmental factors (Fig. 2a). To heighten the differences between the streams, the distribution of individual sites in the ordination space is arranged anew in Fig. 2b. This arrangement shows the na more prominent difference of the Ostruz River sites. na The salmonid character of the Ostruz River (Table 3) na plays a role; all the Ostruz sites are situated at the beginning of the longitudinal gradient reected by the rst axis. The less prominent difference in the position icka of the Tepla and Luz Nisa sites relates to the size and lower altitude of the watershed of both rivers.
(a)
4

Relative abundance data The rst two DCA axes explain 42.5% of species composition in sh assemblages (Fig. 3a). The rst axis (32.7%) reects a gradient of assemblage change in the longitudinal prole similar to the previous analysis. As in the previous analysis (Fig. 2a), the brook lamprey (Lampetra planeri Bloch 1784), brown trout (Salmo trutta m. fario L.) and bullhead (Cottus gobio L.), followed by the rainbow trout (Oncorhynchus mykiss Walbaum 1792), burbot (Lota lota L.) and grayling (Thymallus thymallus L.) represent typical species of the upper parts of the streams (Fig. 2a). The second axis (9.8%) gradient again is not denitively understood from the species data only. In the sites distribution diagram (Fig. 3b), the more na prominent difference of the Ostruz River sites can again be observed; sites are located in the beginning of the longitudinal gradient within the rst axis gradient. icka The separation of the Tepla and Luz Nisa River sites is more extensive than in the presenceabsence data analysis (Fig. 2b). The longest gradient of the DCA was 3.109 SD (standard deviation units of species turnover) for the

(a)

Gyce Anan

Cogo

DCA axis 2

Satr Thth Lapl Lolo

Eslu Ruru Babt Safo Pefl Gogo

Cyca Caca

DCA axis 2

Anan Lele Titi Lece Gyce

Pefl Cogo Lele Ruru Lolo Titi Satr Lece Cyca Eslu Thth Caca Lapl Safo Babt Gogo

2 2.0

DCA axis 1

Onmy

Onmy

2 2.5

DCA axis 1

(b)

(b)

DCA axis 2

0.5 DCA axis 2 1.0

0.5

0.5

DCA axis 1

3.5

DCA axis 1

4.0

Fig. 2. (a) Detrended correspondence analysis (DCA) diagram shows the ordination of species in assemblages using presence absence data of 19 species found in the 31 sites. The rst axis explains 20.5% and the second axis 14.7%. (b) DCA diagram shows the icka na , Luz Nisa, h Ostruz . distribution of sites Tepla

Fig. 3. (a) Detrended correspondence analysis (DCA) using relative abundance data of 19 species found in the 31 sites. The rst axis explains 32.7% and the second axis 9.8%. (b) DCA diagram icka , Luz Nisa, h shows the distribution of sites Tepla na Ostruz .

99

 ka Humpl & Pivnic presenceabsence and 3.657 SD for the species relative abundance data which indicates there are species that show a unimodal response along the gradient in both data types. These results enabled us to use CCA for the direct gradient analyses.
Fish assemblages and environmental factors (CCA)
CCA axis 2 0.8
Gyce Lapl Anan elevation Ruru Titi Eslu

Cogo

% SA F Satr % GR PG + T

% BO

Lele

Differences between streams Presenceabsence data. Canonical correspondence analysis conrmed statistically highly signicant differences (P < 0.0001) between sh assemblages of the individual streams. In total, 21.2% of sh assemblage variability is explained by this pattern, as it roughly summarises differences in environmental conditions of individual streams. Relative abundance data. Statistically highly signicant differences (P < 0.0020) in sh assemblages between the three streams were conrmed also for the relative abundance data. In total, 22.5% variability in sh assemblages is explained. Variability in relation to the environmental factors Data of all three streams were evaluated together and the between-stream variability (21.2% for the presenceabsence and 22.5% for the relative abundance data) was partialled out. Presenceabsence data. The high statistical signicance of the nal CCA ordination model (P < 0.0001) was conrmed by the Monte Carlo permutation test (Fig. 4). The rst two ordination axes explain 26.7% variability in sh assemblages. Number of ponds and land use (category permanent grasslands) are highly correlated with the rst ordination axis (Table 5) and represent the most important environmental gradients related to the sh assemblage structure. Substratum type (categories %sand and %stones) and elevation are the factors most correlating with the second axis. Substratum type (%stones) is the factor most correlating with the third axis (not shown in the diagram). Statistically non-signicant environmental factors (stream slope, stream width, depth, current velocity and cover) were identied in the stepwise forward selection process available in CANOCO and were not used in the model. All six signicant (P < 0.05) environmental factors (distance from the source, elevation, number of ponds, channel type, substratum type and land use) further explain 50.7% of total sh assemblage variability; 21.2% variability was already explained by the differences between the streams. The individual effects of environmental factors were: substratum type 100

Safo Pefl Lolo

Cyca Caca ponds

% SI

Thth CHANNEL Lece km % ST Babt

PG Gogo

Onmy

1.0 0.6

CCA axis 1

1.0

Fig. 4. Canonical correspondence analysis (CCA) ordination diagram showing the effect of signicant (P < 0.05) environmental factors on the structure of sh assemblages (presenceabsence data). Environmental variables measured in situ are in capital letters, variables measured from the map are in lower case letters. The rst axis explains 14.8% and the second axis 11.9%

(19.96%, P < 0.0017), land use (6.86%, P < 0.0178), number of ponds (5.87%, P < 0.0003), elevation (4.19%, P < 0.0042), distance from the source (4.30%, P < 0.0039) and channel type (4.03%, P < 0.0079). Relative abundance data. The statistical signicance of the CCA ordination model (P < 0.0001) was conrmed as well (Fig. 5). The rst two ordination axes explain 36.6% variability in sh assemblages. Similar to the previous analysis, number of ponds and land use
Table 5. Inter-set correlations of signicant (P < 0.05) environmental factors with the rst three ordination axes of the nal canonical correspondence analysis (CCA) (presenceabsence data). CCA axis 1 km Elevation Ponds CHANNEL %BO %ST %GR %SA %SI PG PG+T F )0.30 0.28 0.68 )0.36 0.08 0.02 )0.26 0.02 0.22 0.50 )0.29 )0.19 CCA axis 2 )0.32 0.39 )0.07 )0.22 0.28 )0.36 0.09 0.43 )0.02 )0.25 )0.09 0.32 CCA axis 3 )0.29 0.20 )0.24 )0.19 0.00 0.40 )0.17 )0.24 )0.13 )0.06 0.10 )0.04

For denition of variables see Table 4.

Fish assemblages and environmental factors All six signicant (P < 0.05) environmental factors (distance from the source, elevation, number of ponds, channel type, substratum type, and land use) further explain 58.3% of total sh assemblage variability; 22.5% variability was already explained by the between-stream differences. The individual effects of environmental factors were: substratum type (16.31%, P < 0.0006), land use (7.80%, P < 0.0072), channel type (6.22%, P < 0.0003), number of ponds (3.92%, P < 0.0011), elevation (2.55%, P < 0.0234) and distance from the source (2.50%, P < 0.0288).
Discussion
0.8

Lolo Gyce Eslu Pefl Ruru

CCA axis 2

% BO Cogo

Anan % ST Satr PG + T Lapl Thth % GR % SI Babt Safo PG Lece Onmy Lele Gogo Caca Cyca ponds Titi

% SA F

elevation

km CHANNEL

0.6

CCA axis 1

1.0

Fig. 5. Canonical correspondence analysis (CCA) showing the effect of signicant (P < 0.05) environmental factors on the structure of sh assemblages (relative abundance data). Environmental variables measured in situ are in capital letters, variables measured from the map are in lower case letters. The rst axis explains 23.7% and the second axis 12.9%

(category permanent grasslands) are the factors most correlating with the rst ordination axis (Table 6). These factors represent the most important environmental gradients related to the structure of assemblages. Distance from the source and channel type are the factors most correlating with the second ordination axis. Substratum type (%stones) and elevation are the factors most correlating with the third axis (not shown in the diagram). Statistically non-signicant environmental factors identied in the stepwise forward selection process were the same as in the previous model (stream slope, stream width, depth, current velocity and cover).
Table 6. Inter-set correlations of signicant (P < 0.05) environmental factors with the rst three ordination axes of the nal canonical correspondence analysis (CCA) (relative abundance data). CCA axis 1 km Elevation Ponds CHANNEL %BO %ST %GR %SA %SI PG PG+T F )0.39 0.35 0.72 )0.39 )0.23 )0.11 )0.13 )0.01 0.41 0.73 )0.49 )0.15 CCA axis 2 )0.36 0.34 )0.31 )0.44 0.16 0.14 )0.17 0.33 )0.23 )0.42 0.09 0.27 CCA axis 3 0.42 )0.52 )0.08 )0.21 0.00 0.64 )0.21 )0.27 )0.45 0.02 0.16 )0.18

For denition of variables see Table 4.

As can be seen in the diagrams, the rst two axes in the indirect gradient analysis (DCA) always represent the two most inuential environmental gradients of sh assemblage variability (Figs 2 and 3). From the sh species sequence on the rst ordination axis it can be reasoned that this axis reects the stream longitudinal gradient of sh assemblage change. This was evident similarly for both presenceabsence and relative abundance data. The brook lamprey, bullhead and brown trout are species that occur nearest to the stream source. All the other species appear successively in the lower parts. At the very end of the gradient, species such as common carp (Cyprinus carpio L.), crucian carp (Carassius carassius L.), tench (Tinca tinca L.), and roach (Rutilus rutilus L.) appear. Their occurrence is often inuenced by their escaping from carp ponds or by their regular stocking by shermen. A continuum in assemblage composition in a longitudinal prole was also observed in a Central European submontane stream (Ero s et al. 2003). Marsh-Matthews & Matthews (2000) observed signicant changes in assemblage composition (summarised as a sample score on two axes of DCA) as a function of local and broad geographical factors in a Midwestern stream of the USA. The variability between the streams, already detected by DCA (Figs 2b and 3b), was conrmed by the direct gradient method (CCA); this pattern was similar in both types of data analysed. There was no important difference, even though the relative abundance gives extra information on the similarity of assemblages (Poff & Allan 1995). During the direct gradient analysis (within-stream CCA), environmental variables having signicant effect (P < 0.05) on the sh assemblages were always selected. Having analysed the presenceabsence data it was found that sh assemblage structure is most signicantly affected by number of ponds and land use. These environmental factors are the most closely correlated with the rst ordination axis, so they affect the principal gradient forming the sh assemblages most intensely (Fig. 4, Table 5). The second strongest 101

0.8

 ka Humpl & Pivnic gradient represented by the second ordination axis is best depicted by substratum type and elevation. The partial analyses (CCA) showed that substratum type individually explains the variability in sh assemblage structure to the greatest extent. The occurrence of many sh species in streams is dependent on the characteristic habitat type, where the proportions of the individual categories of substratum type are its important components (Lyons 1989; Doughty & Maitland 1994; Pires et al. 1999). Having analysed the relative abundance data it was found that the sh assemblage structure is also signicantly affected by the number of ponds and land use. These environmental factors correlate well with the rst ordination axis (Fig. 5, Table 6). Distance from the source and channel type explained well the second strongest gradient, which is represented by the second ordination axis. The partial CCA analyses showed that substratum type explains the greatest variability in sh assemblages just like in the presenceabsence data analysis. The three most inuential factors explaining the variability to the greatest extent (substratum type, land use and channel type) were measured in situ. It was shown that the direct gradient analysis also gave very consistent results for both types of data. The same environmental factors appeared to be important both in the presenceabsence and the relative abundance data analyses. The factors having no signicant effect on the assemblages were stream slope, stream width, depth, current velocity and cover. Analyses of both the presenceabsence and the relative abundance data showed that number of ponds is one of the most important environmental factors icka inuencing sh assemblages in the Tepla , Luz na Nisa and Ostruz rivers. This has already been ka 1998; ascertained in many Czech streams (Pivnic ka & Humpl 2004). Other important factors Pivnic explaining a large proportion of the variability are land use, substratum type and channel type. Generally, it is known that the presence of sh or their relative abundance is often related to the distinct types of habitat including bottom substratum type (Lyons 1989; Doughty & Maitland 1994; Pires et al. 1999; Angermeier & Davideanu 2004) and land use (Schlosser 1991; Maret et al. 1997; Angermeier & Winston 1999; Wang et al. 2001). The signicant effect of channel type has also been observed for relative abundance data (Lyons 1996; Angermeier & Davideanu 2004). It was conrmed that sh assemblages vary more signicantly according to the changes in environmental factors (50.7%/58.3%) than according to the regional (between-stream) variability (21.2%/22.5%) for presenceabsence/relative abundance data respectively. This was also conrmed by Marsh-Matthews & 102 Matthews (2000) who demonstrated that the assemblage composition may vary most as a function of environmental and landscape factors. Generally, presenceabsence and relative abundance data of sh gave similar results in both DCA and CCA analyses; the same environmental factors proved to be important in both data type analyses. The overall variability explained in the presenceabsence data analysis was 71.9% (50.7 + 21.2); in the relative abundance analysis it was even 80.8% (58.3 + 22.5). Therefore, summarising results of both types of data analysed we can say that the environmental factors explain sh assemblage variability slightly more effectively together with the relative abundance data than with the presenceabsence data. The environmental factors measured directly in situ explain 2.1-times more variability (map 14.4%, in situ 30.9%) and with the relative abundance data even 3.4times more (map 9.0%, in situ 30.3%). The factors measured from the map give only an average picture about the environment, whereas the values measured directly in situ reect the actual state of the inuence of environment on sh. The role of the factors measured in a map should, however, not be underestimated. Poff & Allan (1995) pointed out that largescale patterns are best detected with coarse-grain data. For ecological communities, relative abundance data provide ne-grain information because they emphasise local peaks in species performance, while species presenceabsence data emphasise a coarser grain of environmental tolerance (Rahel 1990). The results can be used for the prediction of the structure and species composition of sh assemblages. Expected changes can be assessed on the basis of the measurable environmental variables. As far as sh management affects the majority of streams, we can also assess the effect of stocking on species diversity and relative abundance. Both types of information are important for ecologists, conservationists and sh managers.
Resumen
1. Tres r os de taman o comparable localizados en diferentes a reas de paisaje protegido de la Repu blica Checa fueron seleccionados para estudiar el efecto de factores ambientales sobre los ensamblajes de peces. Para ello, utilizando ana lisis de gradientes indirectos (DCA) y directos (CCA). 2. El DCA para las especies enfatizo la variabilidad entre los ensamblajes de peces de los tres r os. Este patro n fue conrmado por un CCA altamente signicativo. Para la variabilidad dentro de la localidad, un CCA revelo que los factores ambientales explicaron un 50.7% para datos de presencia-ausencia y un 58.3% para las abundancias relativas. 3. Los ana lisis revelaron que el nu mero de pozas y el uso del suelo fueron los factores de mayor inuencia en el gradiente ambiental. Sin embargo, en el CCA parcial, el tipo de sustrato

Fish assemblages and environmental factors

explico la mayor proporcio n de la variabilidad que afecta a los peces en la eleccio n de ha bitat. 4. Generalmente los datos de presencia-ausencia y abundancia relativa produjeron resultados similares tanto en los ana lisis DCA como en los CCA; los mismos factores ambientales probaron ser importantes en los ana lisis de ambos tipos de datos. Los factores ambientales explicaron mas variabilidad que los regionales (entre r os). La proporcio n total de variabilidad explicada por el ana lisis de los datos de presencia-ausencia fue 71.9% mientras que para las abundancias relativas fue de 80.8%. Los factores ambientales medidos durante los muestreos de campo explicaron 2.1 y 3.4 veces mas variabilidad que los factores medidos sobre mapas hidrolo gicos. Lyons, J. 1989. Correspondence between the distribution of sh assemblages in Wisconsin streams and Omerniks ecoregion. American Midland Naturalist 122: 163182. Lyons, J. 1996. Patterns in the species composition of sh assemblages among Wisconsin streams. Environmental Biology of Fishes 45: 329346. Maret, T.R., Robinson, C.T. & Minshall, G.W. 1997. Fish assemblages and environmental correlates in least-disturbed streams of the Upper Snake River Basin. Transactions of the American Fisheries Society 126: 200216. Marsh-Matthews, E. & Matthews, W.J. 2000. Geographic, terrestrial, and aquatic factors: which most inuence structure of Midwestern USA stream sh assemblages? Ecology of Freshwater Fish 9: 1321. Moyle, P.B., Crain, P.K., Whitener, K. & Mount, J.F. 2003. Alien shes in natural streams: sh distribution, assemblage structure, and conservation in the Cosumnes River, California, USA. Environmental Biology of Fishes 68: 143162. Oberdorff, T., Guilbert, E. & Lucchetta, J.C. 1993. Patterns of sh species richness in the Seine River basin, France. Hydrobiologia 259: 157167. Oberdorff, T., Pont, D., Hugueny, B. & Chessel, D. 2001. A probabilistic model characterizing sh assemblages of French rivers: a framework for environmental assessment. Freshwater Biology 46: 399415. Ostrand, K.G. & Wilde, G.R. 2002. Seasonal and spatial variation in a prairie stream-sh assemblage. Ecology of Freshwater Fish 11: 137149. Pires, A.M., Cowx, I.G. & Coelho, M.M. 1999. Seasonal changes in sh communities structure of intermittent streams in the middle reaches of the Guadiana Basin, Portugal. Journal of Fish Biology 54: 235249. ka, K., 1998. The inuence of some parameters on sh Pivnic ivokla diversity in small streams of the Protected Area Kr tsko. R II:3134 (in Czech with Biodiverzita ichthyofauny C English summary). ka, K. & Humpl, M. 2004. Fish assemblages in the Elbe Pivnic River watershed species richness, frequency, and clusters. Acta Universitatis Carolinae Environmentalica 18: 107116. Poff, L.N. & Allan, J.D. 1995. Functional organisation of stream sh assemblages in relation to hydrological variability. Ecology 72: 606672. Rahel, F.J. 1990. The hierarchical nature of community persistence: a problem of scale. The American Naturalist 136: 328344. Schlosser, I.J. 1982. Fish community structure and function along two habitat gradients in a headwater stream. Ecological Monographs 52: 395414. Schlosser, I.J. 1991. Stream sh ecology: a landscape perspective. BioScience 41: 704712. Wang, L., Lyons, J., Kanehl, P. & Bannerman, R. 2001. Impact of urbanisation on stream habitat and sh across multiple spatial scales. Environmental Management 28: 255266.

Acknowledgements
ek during all eld We acknowledge the help of Dr Josef Kr z operations. The research was funded by the Grant Agency of the Academy of Science of the Czech Republic, Project no. A6093105.

References
Angermeier, P.L. & Davideanu, G. 2004. Using sh community to assess streams in Romania: initial development of an index of biotic integrity. Hydrobiologia 511: 6578. Angermeier, P.L. & Winston, M.R. 1999. Characterizing sh community diversity across Virginia landscapes: prerequisite for conservation. Ecological Applications 9: 335349. Belanger, S.E. 1997. Literature review and analysis of biological complexity in model stream ecosystems: inuence of size and experimental design. Ecotoxicology and Environmental Safety 36: 116. milauer, P. 2002. CANOCO reference ter Braak, C.J.F. & S manual and CanoDraw for Windows users guide: software for canonical community ordination (version 4.5). Ithaca, NY: Microcomputer Power. 500 pp. Brown, L.R. 2000. Fish communities and their associations with environmental variables, lower San Joaquin River Drainage, California. Environmental Biology of Fishes 57: 251269. Doughty, C.R. & Maitland, P.S. 1994. The ecology of the River Endrick: present status and change since 1960. Hydrobiologia 290: 139151. Ero s, T., Botta-Duka t, Z. & Grossman, G.D. 2003. Assemblage structure and habitat use of shes in a Central European submontane stream: a patched based approach. Ecology of Freshwater Fish 12: 141150. Godinho, F.N., Ferreira, M.T. & Santos, J.M. 2000. Variation in sh community composition along an Iberian river basin from low to high discharge: relative contributions of environmental and temporal variables. Ecology of Freshwater Fish 9: 2229.

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