1

Macrophyte distribution in the River Vils (Oberpfalz, Bavaria)

Alexander Kohler1, Esther Sonntag1, Matthias Köder1, Karin Pall2, Uwe Veit1, Georg-

Heinrich Zeltner1 & Georg A. Janauer3

With 6 figures and 1 table in the text

Abstract: The macrophyte vegetation of the eutrophic River Vils (Oberpfalz, Bavaria) was surveyed and mapped in summer 1999. Regarding carbonate content, the river is of medium to rather hard. The macrophyte diversity is relatively high. In the Vils 26 tracheophytic hydrophytes, 24 tracheophytic amphiphytes and 37 helophytic species were found. The picture is dominated by eutrophent elements like Potamogeton pectinatus, Ceratophyllum demensum, Sparganium emersum and others. Floristic specialities in the Vils are the hybrids

Potamogeton x fluitans and Potamogeton x schreberi. The distribution diagram shows patterns for several species in the river course. According to changes in the floristic composition and river-morphological structures, the river could be divided into five reaches

(A-E). These reaches were characterized regarding Relative Plant Mass (RPM) and species numbers as well as Mean Mass Index and Relative Area Length. The RPM values in combination with species numbers revealed important new aspects regarding biodiversity and the dominance of macrophyte species. Additionally the macrophyte distribution across the river was described by five transects.

1 University of Hohenheim, Institute of Landscape and Plant Ecology (320), D - 70593 Stuttgart, Germany 2 Systema, Bio- und Management Consulting GmbH, Bensasteig 8, A – 1140 Wien, Austria 3 University of Vienna, Institute of Ecology and Conservation Biology, Department of Hydrobotany, Althanstrasse 14, A – 1090 Vienna, Austria 2

Introduction

The eutrophic, carbonate-rich River Vils is a tributary to the in the German catchment. Industrial and urban pollution as well as diffuse loading from agricultural areas through which the river flows, results in a critically loaded system over long stretches. Some reaches have improved due to construction of sewage treatment plants. Several barrages and other management impacts have changed the river bed of the Vils to a great extent. In spite of these anthropogenic influences, the river still has a remarkable and especially rich macrophyte vegetation. This was surveyed for the first time in Summer 1990 (JUNGE et al. 1991) and the survey was repeated by our team in Summer 2000 (SONNTAG et al. 2000). The following problems were investigated:

- What is the macrophyte composition in a eutrophic and turbid running water?

- Are there species with a distinctive distribution pattern along the river and is it

possible to differentiate floristic, ecological river zones?

- In which way can the vegetation of the river with its five reaches be characterized

quantitatively by means of the parameters Mean Mass Index, Relative Plant mass and

species numbers?

- Which additional information does the investigation of the macrophytic distribution

across the river provide concerning the characterization of the vegetation?

Survey area

The Vils is a typical river in a hilly landscape of medium elevation. The catchment area is

1096 km2 and comprises the "Oberpfälzer Hügelland", which is dominated by sandstones

(Keuper, Buntsandstein), and the "Nördliche and Mittlere Frankenalb" (Figure 1). South of the town of Amberg, the river runs through a deep cut valley in jurassic limestone. The 3 gradient is between 5 and 1.5‰. The main land use types are agriculture (47%) and forestry

(42%). From the source to Vilshofen, the saprobic quality is "critically loaded" (quality class

2-3) and from there on to the confluence with the Naab it is "fairly loaded" (Quality class 2)

(Figure 2, WWA-Info.1 1996, cited after SONNTAG et al. 2000). In the upper reaches the structural quality of the river is "near natural", and some reaches in the lower stretch of the river are of the same quality (Figure 2). Overall only 10% of the river is of near-natural quality.

The total water hardness of the Vils is between 10 and 19.8° DH ("medium hard" to "rather hard" water, Höll, 1996). Water hardness, as well as carbonate hardness (8-16° K.H.), rises in the jurassic stretch. Electrical conductivity rises down river from 370-595 µS.cm-1. The pH is close to 8.0, which is typical for water rich in hydrogen-carbonate (SONNTAG et al. 2000).

In several stretches, the aquatic vegetation was cut with electrical underwater scythes. After a severe decrease in the aquatic vegetation only a stretch in the middle of the river channel is now being cut.

Methods

The entire river from the source to its confluence with the Naab (87 km length) was surveyed following the method of KOHLER (1978). The quantitative assay is described by JANAUER et al. (1993), KOHLER & JANAUER (1995), PALL & JANAUER (1995). A more detailed description was given by SONNTAG et al. (2000).

Results

Occurrence of macrophytes in the River Vils 4

Macrophyte taxa and growth forms are collated in Table 1. The floristic list is divided into hydrophytes, amphiphytes and helophytes.

Several macrophyte species of the Vils River are noted in the "red list of Germany". Butomus umbellatus, Groenlandia densa, Potamogeton alpinus, Potamogeton berchtholdii, P. nodosus,

P. perfoliatus, and Eleocharis ovata and Matteuccia struthiopteris are endangered in Bavaria on level 3 ("endangered", Rote Liste gefährdeter Pflanzen Deutschland, 1996) (all authors of the species names are claimed in T. 1). The hybrids of Potamogeton, P. x fluitans and

P. x schreberi are probably rare taxa (all authors of the species names are claimed in T. 1).

Little is known about their distribution and habitat conditions in Southern Germany and they should be looked for in other water bodies. P. x schreberi was found in the Vils near

Hahnbach in 1936 (GLÜCK 1936) (all authors of the species names are claimed in Table 1).

Regarding neophytic species in the Vils, Elodea canadensis is the only agriophytic neophyte among the hydrophytes. The ubiquitous and spreading Elodea nuttallii was not (yet) found in the Vils by 1999 (all authors of the species names are claimed in T. 1). In the riparian reach

Arch angelica and Impatiens glandulifera are notable neophytic plants.

Pattern-building macrophyte species

Some macrophyte species show distinctive distribution patterns in the run of the river (Figure

3). The highly eutraphent bleustophyte, Ceratophyllum demersum (indicator value 3.18,

SCHNEIDER 2000) is widely distributed but it is missing in the upper reaches and near the mouth (from survey stretch 109 down river). Sagittaria sagittifolia is also eutraphent (IW

2.98), occurring down river of survey stretch 19 but is missing near the mouth. Potamogeton nodosus (highly eutraphent: IW 3.10) has two centres of occurrence. Ranunculus fluitans (IW

3.00) is found below the confluence of the Rosenbach into the town of Amberg. Several hydrophytic species are missing in the uppermost reach. These are Myriophyllum spicatum 5

(IW 2.83), P. pectinatus (IW: 2.88) and E. canadensis (IW 2.55). Callitriche hamulata (IW

1.80), P. crispus (IW 2.88) and Nuphar lutea (IW 3.15). In the lowermost reaches of the river.

Berula erecta (IW 2.65) occurs only downriver of survey stretch 1.09. P. berchtholdii (IW

2.40) has its focus of occurrence in the lower half of the river.

A direct ecological relationship between habitat parameters and the pattern of occurrence of species is not possible, because the number of chemical data are not sufficient. Yet, based on the changes in floristic composition and those of river morphology, the Vils could be divided into five reaches (A – E, SONNTAG et al. 2000). These reaches will be compared regarding their Relative Plant Mass (RPM)-diagrams, which only contain "higher" hydrophytes and amphiphytes (tracheophytic species). A more detailed comparison regarding Mean Mass

Indices and other aspects is given by SONNTAG et al. (2000).

Mean Mass Index and Distribution Ratio "d"

In the entire river only five species occur with high MMO-and MMT-values (>3, Figure 4).

These species are hydrophytic: M. spicatum, E. canadensis, and P. pectinatus, and the amphiphytes Phalaris arundinacea and Sparganium emersum (type 1, JANAUER et al. 1993).

Clumped occurrence (type 2) is typical for P. perfoliatus, Ranunculus trichophyllus, R. peltatus and some amphiphytes like Glyceria maxima, Sparganium erectum and B. erecta.

According to their respective MMO-/MMT-values P. lucens, G. densa, B. umbellatus,

Schoenoplectus lacustris, Alisma plantago-aquatica, P. x fluitans, M. verticillatum, P. alpinus are rare taxa. This is not surprising regarding oligo-traphent species like G. densa (IW 1.83) and P. alpinus (IW 1.55) in a river with eutrophic conditions. Eutraphent taxa like B. umbellatus (IW 2.98) and P. lucens (IW 2.65) are dominant in other eutrophic rivers, e.g. in the province of Schonen in Southern Sweden (KOHLER et al. 2000, SIPOS et al. 2000,

SONNTAG et al. 1999). 6

A quantitative expression for the proportion of individual river reaches inhabited by certain species is the Distribution Ratio (which is numerically equivalent to the Relative Area-Length

Lr, Figure 2). A wide distribution in the river (> 50%) is typical for the hydrophytes M. spicatum, P. pectinatus, E. canadensis and C. hamulata, and the amphiphytes P. arundinacea and S. emersum. Values between 40-50% are reached by P. crispus, C. demersum, L. minor,

P. berchtholdii, N. lutea, Spirodela polyrhiza, Callitriche cophocarpa, Myosotis scorpioides and S. sagittifolia, and the haptophyte, Fontinalis antipyretica. The distribution ratios (d) of all other species are much lower.

Relative Plant Mass and Species Numbers

The Relative Plant Mass (RPM)-values were calculated only for tracheophytic hydrophytes and amphiphytes. RPM-values >10% are only reached by the amphiphytes P. arundinacea, S. emersum and by the hydrophyte, P. pectinatus (Figure 5). Upon comparison, the river reaches

A – E show remarkable differences regarding the dominance and diversity of the species. In river zone A, where no tracheophytes occurred, the amphiphytic P. arundinacea dominated

(48% RPM). In river zone B the dominant species is the hydrophytic E. canadensis (28%), followed by P. arundinacea (19% RPM). River zone C has no highly dominant species.

P. arundinacea, P. pectinatus and S. emersum reach values between 13%-10%. In river zone D, S. emersum dominates (23% RPM), followed by P. arundinacea, M. spicatum and

E. canadensis (12-10% RPM). The lower reach of the river (zone E) is dominated by

B. erecta (16% RPM).

A highly interesting result is the species number of hydrophytes and amphiphytes in the river and in the floristic-ecological river zones A – E.

26 hydrophytic species out of a total of 50 recorded is a very high value regarding conditions in running waters in Southern Germany. With respect to the fact that only 10% of the Vils is 7 structurally close to natural conditions, this is a remarkable fact. In Southern Sweden, where the river structure is rarely ever influenced, but the rivers are similarly eutrophic, species numbers are between 61 (Kävlingean, hydrophytes 30), and 51 (Björkaän, hydrophytic species: 21, SIPOS 2001). River zones A – E show remarkable differences in species numbers in the field. In river zone, E only eight species occur, which comprise no tracheophytes.

Species richness increases down-river (zone B: 26, zone C: 38 species), whereas the highest number of tracheophytic hydrophytes is reached in zone C, where 20 taxa were found.

In addition to the longitudinal survey of the Vils five transects were mapped. Transects of one metre width and 1 x 0.4-1.5 m aerial units were placed in characteristic and/or floristically interesting locations. Within the survey units of the belt transect the cover (%) of the macrophyte species present was estimated, which is graphically displayed in a bar diagram

(Figure 6).

• the first transect was in the upper reach of survey stretch 80. The river is 3 m wide and a

maximum of 0.275 m deep. The substrate is sandy, the flow is slow with little turbulence.

only a few helophytes can live on the deep banks. Persicaria lapathifolia grows on the

bank. R. peltatus forms thick swathes in the middle of the river (cover 75-100%).

Neophytic E. canadensis is ubiquitous over the whole width of the river bed with

changing cover. P. crispus grows in the southern half of the river only (cover up to 50%).

• The 2nd transect was located in survey stretch 2.20 m north of the bridge at -

Sorghof. The river is 9 m wide and 0.9 m deep. The flow is slow and the substrate is

gravel. M. spicatum is distributed over the whole transect in all depths, E. canadensis

occurs only near the banks.

• Transect 3 was located in survey stretch 34. The river is 10.4 m wide. The side is close to

the road between sewage treatment plants at Hallenbach and Kümmersbuch. The substrate

in the centre of the channel is gravel, closer to the banks it is muddy. The southern bank 8

was filled in with large gravel. Maximum depth is 1.35 m. Flow is slow. Amphiphytic

S. emersum is found across the whole transect, except in the fresh fill-in in the south. High

cover is reached by P. crispus, P. pectinatus (up to 100%), and E. canadensis (up to 75%).

• Transect 4 was located in survey stretch 46, 100 m south of the bridge in Traßelberg. The

banks are constructed with rip-rap, the river bottom is comprised of gravel and / or stones.

The river 11 m wide and maximum depth is 0.48 m. Flow is slow. M. spicatum and the

haptophyte, F. antipyretica grow almost across the whole transect. R. fluitans is dominant

in the middle of the channel (cover 25-75%).

• The last transect was in the lower reach of the Vils, survey stretch 113, in Schmidmühlen.

River width is 15.5 m, maximum depth is 0.7 m. Turbulent flow is found over gravel and

stone substrates. In the river centre C. hamulata occurs with medium and high cover. All

other species grow closer to the banks. There B. erecta and S. emersum can reach high

cover (50-75%).

Discussion

The Vils represents a eutrophic, calcium, hydrogen-carbonate rich river of medium elevation.

The species inventory is a mirror to the conditions imposed by water chemistry and the eutrophic background. Significant eutraphent species like P. pectinatus, C. demersum,

S. sagittifolia, S. emersum, and S. erectum dominate the vegetation over large stretches. The analysis of the aquatic vegetation was surprising. In comparison with other eutrophic rivers, rich in calcium hydrogen carbonate in southern Sweden (SIPOS et al., 2000, SONNTAG et al.,

1999), the species richness of the Vils is of quite the same order. The Rivers Björkanon and the Kavlingeön showed species numbers (tracheophytic hydrophytes and amphiphytes) of 51, and 61 taxa, respectively. In the Vils, 50 species occurred. Tracheophytic species are 26 taxa in the Vils, 21 taxa in Björkaän and 30 taxa in Kävlingeän. This is remarkable, because the 9

Vils has only 10% of river reaches in near natural conditions regarding river structure and morphology. In comparison the two rivers in southern Sweden are natural to near-natural over most of their length.

Experience with the species inventory of eutrophic rivers makes it probable that they are a type of aquatic biotopes, which are not only characterised by high species numbers, but rather comprise ecosystems in which a number of rare and even endangered taxa have found a refuge. In the Vils several species were found, which are in the red list of Germany and

Bayern (ROTE LISTE GEFÄHRDETER PFLANZEN DEUTSCHLANDS, 1996). Among them is the genus Potamogeton. It must be stated also, that in the Vils, Potamogeton hybrids are occurring for which there is no clear present knowledge regarding scarcity, endangerment, and habitat conditions, which should be investigated in the future. P. x fluitans and P. x schreberi are the two hybrids to be named for the Vils.

In the Vils, as in other rivers, several species show distinct patterns of distribution along the river course. In originally oligotrophic rivers the distribution pattern of species is often an expression of different trophic conditions. In eutrophic rivers, the distribution pattern cannot be explained by habitat related parameters. This may be due, in part, to a lack of sufficient ecological data. In running waters the absence and / or the distribution of species may have different reasons than those based on habitat parameters (e.g. CAIRNS 1974).

Although the Vils can be compared with eutrophic rivers in southern Sweden in terms of its florisitic compositions (e.g. occurrence of common eutrophic elements) definite floristic differences can be seen. A typical, highly dominant amphiphytic "Leitart" (characteristic species) Butomus umbellatus found in the rivers of the province of Schonen, was detected in the Vils only as a single individual. Another typical amphiphyte, S. latifolium, occurring in

Swedish rivers, is totally absent in the Vils. 10

Some macrophyte species show distribution patterns in the river (Figure 3). No tracheophytic hydrophytes were present in the upper reaches near the source of the river. The distribution pattern of Ceratophyllum demersum, a highly eutraphent species, can be correlated with the high nutrient levels in the river reaches dominated by it. The occurrence of Sagittaria sagittifolia may be explained by impounded reaches and very fine sediments in the river bed.

A very strict differentiation according to floristic-ecological river zones was not done in the

Vils, because many of the distribution patterns could not be explicitly explained, even though five river zones (A – E) were distinguished. Several parameters were found to be characteristic (species number, relative plant mass, etc.).

River zone A is characterised by the absence of tracheophytic hydrophytes and the dominance of amphiphytes. Very low discharges and, in part, extremely strong shading are the most probable reasons for this phenomenon. The species number is very low (8) in this reach near the source. The amphiophyte, Phalaris arundinacea dominates with 48% RPM. The adjacent river zone B already favours hydrophytes, among which Elodea canadensis is dominant (28%

RPM). The total species number increases to 26 (12 hydrophytic) species. River zone C is the most species rich of the Vils: here 38 species occur, among which are 20 tracheophytic hydrophytes. The RPM diagram shows that no highly dominant species occur here and the respective ratios of the taxa vary across a wide range. The high biodiversity of this zone can be explained by the relatively high structural and morphological diversity in the numerous survey stretches in association with close to natural conditions. In river zones D and E the species number decreases again (total: 37, hydrophytic 30 species, total 17, hydrophytic 15 taxa, respectively). Zone D is dominated by Sparganium emersum, zone E by Berula erecta.

The examples discussed above show that the RPM diagrams in combination with species numbers of tracheophytic, amphiphytes and hydrophytes are a valuable new instrument to evaluate and compare individual river reaches and to characterise different zones in a 11 qualitative and quantitative way. The Relative Area Length (which is equivalent numerically to the distribution ratio d) and the Mean Mass Indices have been found to be useful parameters for comparison of running waters and their ecological zones. All these parameters are tools for describing dominance, diversity, Mass Ratios and areas of macrophytes in rivers.

They are not only a scientific tool for describing the ecology , but they are also useful for management tasks regarding biotopes and species in rivers. Long-term studies have shown that with the help of these numerical tools, definite statements about the effects of river pollution human impacts and restoration measures in rivers can be made (PALL et al. 1999,

VEIT et al. 1997).

Conclusion

The macrophyte vegetation of the eutrophic River Vils (Oberpfalz, Bayern) was surveyed and mapped in Summer 1999. Regarding carbonate content the river is of medium to rather hard.

Due to changes in river morphology over the pasted centuries only a few stretches are close to natural regarding the run of the river and its structural features.

In spite of these changes the macrophyte diversity is relatively high in the Vils and comes close to eutrophic rivers in southern Sweden, which are structurally intact and close to natural conditions. In the Vils 26 tracheophytic hydrophytes, 24 tracheophytic amphiphytes and 37 helophytic species were found. The picture is dominated by eutraphent elements like

Potamogeton pectinatus, Ceratophyllum demersum, Sparganium emersum and others. A few oligotraphent species like Potamogeton alpinus and Groenlandia densa were also detected.

Floristic specialities in the Vils are the hybrids Potamogeton x fluitans, and P. x schreberi.

The distribution diagram (Figure 3) shows patterns for several species in the river course. The mass development of Ceratophyllum demersum, Potamogeton nodosus and Ranunculus fluitans is probably based on higher nutrient levels. Sagittaria sagittifolia is focused in 12 impoundments with fine sediments and avoids reaches with sandy or gravel bed-sediments.

Berula erecta occurs in turbulent, quick flowing and in shallow river reaches near the mouth.

According to changes in the floristic composition and river-morphological structures, the river could be divided into five reaches (A – E). These reaches were characterised regarding

Relative Plant Mass (RPM) and species numbers for tracheophytic amphiphytes and hydrophytes. The macrophyte distribution across the river was described by 5 transects.

Important results were found using the RPM values and species number. The relationships of dominance and biodiversity of macrophytes were much different in five river zones, where the highest species number was found in zone C. However, in contrast to the other floristic zones, the RPM diagrams showed no real dominants here. The high species richness in this river reach may be due to the very near natural morphological and structural characteristics of the river.

The methodology regarding the survey of the aquatic vegetation and the preparation of numerical derivatives of field data (KOHLER & JANAUER 1995) also proved successful for the eutrophic Vils River. The RPM values in combination with species numbers revealed important new aspects regarding biodiversity and the dominance of macrophyte species (see

SIPOS 2001).

Acknowledgements

The authors received generous support by many institutions. The Wasserwirtschaftsamt

Amberg is thanked for accommodation, boats and numerous background data on the hydrology and physical and chemical conditions of the river. Specials thanks go to Mr.

Viehauser and his colleagues of Flussmeisterei Amberg, who supported us all the time in the best way possible. The investigation was financially supported by the Geschwister-Stauder-

Schenkung. 13

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Virologie, Biologie. 7. Aufl. - de Gruyter Verlag Berlin, 592 pp.

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Charakterisierung und vergleichenden Beurteilung der Gewässervegetation. - Ber. Inst.

Landschafts- Pflanzenökologie Univ. Hohenheim 2: 59-70.

JUNGE, M., MELZER, A. & ZIMMERMANN, S. (1991, n. p.): Submerse Makrophytenvegetation

der Vils/Opf. - Endbericht über die Untersuchungen in der Vegetationsperiode 1990.

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KOHLER, A. & JANAUER, G. A. (1995): Zur Methodik der Untersuchung von aquatischen

Makrophyten in Fließgewässern. - In: STEINBERG, CH., BERNHARDT, H. & KLAPPER, H.

(Hrsg.): Handbuch Angewandte Limnologie. VIII-1.1.3. Ecomed Verlag.

KOHLER, A., SIPOS, V., SONNTAG, E., PENKSZA, K., POZZI, D., VEIT, U. & BJÖRK, S. (2000):

Makrophyten-Verbreitung und Standortqualität im eutrophen Björka-Kävlinge-Fluss

(Skåne, Südschweden). - Limnologica 30: 281-298.

PALL, K. & JANAUER, G. A. (1995): Die Makrophyten-Vegetation von Flußstauen am Beispiel

der Donau zwischen Fluß-km 2552,0 und 2511,8 in der Bundesrepublik Deutschland. -

Arch. Hydrobiol. Suppl. 101, Large Rivers 9/2: 91-109. 14

PALL, K., JANAUER, G. A. & DOKULIL, M. (1999): Sanierung der Alten Donau in Wien -

Entwicklung der Makrophytenbestände. - Deutsche Gesellschaft für Limnologie,

Tagungsber. 1998 Klagenfurt: 269-277.

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Bonn-Bad Godesberg (Hrsg.). 744 pp.

SCHNEIDER, S. (2000): Entwicklung eines Makrophytenindex zur Trophieindikation in

Fließgewässern. - Shaker Verlag, Aachen, 182 pp.

SIPOS, V., KOHLER, A. & BJÖRK, S. (2000): Makrophyten-Vegetation und Standorte im

eutrophen Björka-Fluß (Südschweden). - Bot. Jahrb. Syst. 122: 93-152.

SIPOS, V. (2001): Makrophyten-Vegetation und Standorte in eutrophen und humosen

Fließgewässern - Beispiele aus Südschweden und Ungarn. Diss. Univ. Hohenheim.

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Makrophyten-Vegetation und Standorte im eutrophen Kävlinge-Fluß (Skåne,

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Oberpfalz, Amberg, 4 pp. 15

Figures

Fig. 1: Examination area of the River Vils with its catchment area and the natural land

units.

Fig. 2: Water quality (saprobity system) and structural quality of the River Vils according

to WWA-INFO (1996).

Fig. 3: Distribution diagram of hydrophytic, amphiphytic, helophytic and haptophytic

species from the spring (left side) to the mouth (right side) of the River Vils.

Fig. 4: Mean Mass Index (white: MMO, black: MMT) and „Relative Area Length“ (d)

for the complete River Vils.

Fig. 5: Relative Plant Mass (RPM) for the River Vils and its five reaches. Species with an

RPM value < 1 % are represented in the „residual“.

Fig. 6: Macrophytic distribution across the river at five different sites in the River Vils.

16

Tables

Tab. 1: Species list of the River Vils.

Fig. 1

Fig. 2

Fig. 3

Friedberger Ach Ground water ditches

1972 1972

40 40

30 30

20 20 9 12 RPM [%] RPM [%] 10 10

0 0 Ran tri Ran Pot cri Pot Ran tri Ran Pot col Pot Myr spiMyr Agr sto Agr Agr sto Agr Cha vul Cha Zan pal Cha his Cha Elo can Elo ere Ber Ber ere Ber Spa nat Spa Pha aru Pha Pha aru Pha Ran glu Ran Ver ana Ver residual Pot pec Pot Jun sub Pot pec Pot residual Gro den Gro Myo sco Myo Men aqu Men Nas oem Nas Nas oem Nas

1978 1978

40 40

30 30

20 20 9 11 RPM [%] RPM [%] 10 10

0 0 Gly flu Gly Pot cri Pot Ran tri Ran Ran tri Ran Jun art Ran flu Ran Pot col Pot Myr spiMyr obt Cal Agr sto Agr Agr sto Agr Zan pal Cha vul Cha Elo can Elo Ber ere Ber Elo can Elo ere Ber Ran glu Ran Pha aru Pha glu Ran Pha aru Pha Ver ana Ver Ver ana Ver Pot pec Pot residual Pot pec Pot Jun sub residual Pot ber Myo sco Myo Spa eee Spa Men aqu Men Men aqu Men Spa eee Spa Nas oem Nas Nas oem Nas

1982 1982

40 40

30 30

20 20

RPM [%] 10 RPM [%] 8 10 10

0 0 Gly flu Gly Ran tri Ran Ran tri Ran Pot cri Pot Jun art Pot col Pot Cal obt Cal Myr spiMyr Agr sto Agr Agr sto Agr Zan pal Elo can Elo his Cha Ber ere Ber Ber ere Ber can Elo Ran glu Ran Pha aru Pha Ran glu Ran Pha aru Pha Ver ana Ver Ver ana Ver residual Jun sub Pot pec Pot Pot pec Pot residual Pot ber Gro den Gro Gro den Gro Myo sco Myo Men aqu Men Men aqu Men Nas oem Nas Nas oem Nas

1987 1987

40 40

30 30

20 20

RPM [%] 8 RPM [%] 9 10 10

0 0 Ran tri Ran Ran tri Ran Pot cri Pot Jun art Pot col Pot Pot col Pot Myr spiMyr Cal obt Cal Agr sto Agr Agr sto Agr Cha vul Cha Zan pal Ber ere Ber Ber ere Ber can Elo Pha aru Pha Ran glu Ran Pha aru Pha Ver ana Ver Ver ana Ver Pot pec Pot Jun sub pec Pot residual residual Pot ber Gro den Gro Myo sco Myo Myo sco Myo Men aqu Men Spa eee Spa Lem min Lem Nas oem Nas Nas oem Nas

1992 1992

40 40

30 30

20 20 14

RPM [%] RPM [%] 16 10 10

0 0 Ran tri Ran Pot cri Pot Ran tri Ran Spi pol Spi Pot col Pot Myr spiMyr Cal obt Cal Cha vul Cha Zan pal Fon ant Elo can Elo Ber ere Ber Ber ere Ber Ran glu Ran Pha aru Pha Pha aru Pha Ver ana Ver Ver ana Ver Jun sub pec Pot residual residual Pot pec Pot Pot ber Myo sco Myo Men aqu Men Lem min Lem Lem min Lem Nas oem Nas Nas oem Nas

1996 1996

40 40

30 30

20 20

RPM [%] 8 RPM [%] 14 10 10

0 0 Pot cri Pot Ran tri Ran Pot col Pot Cal obt Cal spiMyr Agr sto Agr Myr verMyr Agr sto Agr Zan pal Elo can Elo Ber ere Ber can Elo Fon ant Ber ere Ber Ran glu Ran Ran glu Ran Pha aru Pha Pha aru Pha Ver ana Ver Ver ana Ver residual Jun sub residual Pot pec Pot Pot ber Myo sco Myo Spa eee Spa Men aqu Men Nas oem Nas Nas oem Nas Fig. 4 Myr spi Myr spi

Pot pec Pot pec

Elo can Elo can

Cal ham Cal ham

Pot cri Pot cri

Cer dem Cer dem

Lem min Lem min

Pot ber Pot ber

Nup lut Nup lut

Spi pol Spi pol

Cal cop Cal cop

Ran cir Ran cir

Ran tri Ran tri

Pot per Pot per

Gro den Gro den

Pha aru Pha aru

Spa eme Spa eme

Myo sco Myo sco

Sag sag Sag sag

Ver bec Ver bec

Cal pal Cal pal

Ror amp Ror amp

Gly max Gly max

Agr sto Agr sto

Spa ere Spa ere

Ber ere Ber ere

Ver a-a Ver a-a

Men aqu Men aqu

Ang arc Ang arc

But umb But umb

Fon ant Fon ant

12 3 4 5 0 0,25 0,5 0,75 1

MMT MMO d Fig. 5 Vils, all sections 30

H: 11+15=26 20 A: 10+14=24

G: 50

RPM (%) RPM 10

0 Pot cri Pot Spi pol Spi Per lap Per Nup lut Myr spiMyr Pot ber Pot Ber ere Ber Elo can Elo Cal cop Cal Pot pec Pot Ver bec Ver residual Pha aru Spa ere Gly max Gly Sag sag Myo sco Myo Cal ham Cal Ror amp Ror Cer dem Cer Lem min Spa eme

48 zone A 30

H: 0+0=0 20 A: 7+1=8

G: 8

RPM (%) RPM 10

0 Cal pal Cal Nas off sto Agr Gly dec Gly Ver bec Ver residual Pha aru Myo sco Myo

zone B 30 H: 6+6=12 20 A: 5+9=14 G: 26 10 RPM (%) RPM

0 Cal Spa ham eme Per lap Per Pot nat Pot Elo can Elo Ver bec Ver

zone C 30 H: 11+9=20 20 A: 11+6=17 G: 37 10 RPM (%) RPM

0 aru ere sag sco Cal Pha Spa Sag Myo Spa eme ham Cal pal Cal Ran cir Myr spiMyr Ver bec Ver nod Pot residual

zone D 30 H: 10+7=17 20 A: 8+12=20 G: 37 10 RPM (%) RPM

0 Gly sco Cal Ror Myo Spa max amp ham eme Pot cri Pot Spi pol Spi Myr spiMyr Pot ber Pot residual

zone E 30 H: 7+8=15 20 A: 8+7=15 G: 30 10 RPM (%) RPM

0 ere sco Ror Spa Myo Spa amp eme Myr spiMyr Pot per Pot Ber ere Ber Cal cop Cal Fig. 6

Transect 1 in section 80 S N

0 depth (m) 0,2 width (m) 0 123 Rumex aquaticus Epilobium roseum Persicaria lapathifolia Potamogeton crispus Callitriche hamulata Elodea canadensis Ranuculus peltatus

Transect 2 in section 2 WO

0 depth (m) 0,2 0,4 0,6 0,8 1,0 width (m) 0 123456789 Callitriche hamulata Phalaris arundinacea Carex acuta Elodea canadensis Potamogeton pectinatus Potamogeton berchtoldii Myriophyllum spicatum Sparganium emersum

Transect 3 insection 34 N S

0 depth (m) 0,2 0,4 0,6 0,8 1,0 1,2 1,4 width (m) 0 1234567891010,4 Spirodela polyrhiza Filipendula ulmaria Potamogeton crispus Ceratophyllum demersum Sagittaria sagittifolia Potamogeton pectinatus Elodea canadensis Sparganium emersum

Transect 4 in section 46 NO SW

0 depth (m) 0,2 0,4 width (m) 0 1234567891011 Phalaris arundinacea Potamogeton pectinatus Fontinalis antipyretica Myriophyllum spicatum Callitriche hamulata Sparganium emersum Agrostis stolonifera Ranunculus fluitans

Transect 5 in section 113 O/SO W/NW

0 depth (m) 0,2 0,4 0,6 width (m) 0 12345678910111213141515,5 Potamogeton berchtoldii Phalaris arundinacea Chlorophyta indet. Fontinalis antipyretica Sparganium emersum Berula erecta Callitriche hamulata

coverage (%) 0-25 25-50 50-75 75-100 Tab. 1 Species list of the River Vils (Oberpfalz, Bavaria)

Species Abbreviation Occurrence Growth form

Hydrophytes Callitriche cophocarpa Sendtn. (Cal cop) z sr Callitriche hamulata Kütz ex W.D.J. Koch (Cal ham) z sr Ceratophyllum demersum L. (Cer dem) z mp Elodea canadensis Michx. (Elo can) z sr Groenlandia densa (l.) Fourr. (Gro den) z sr Lemna minor L. (Lem min) z ac Lemna gibba L. (Lem gib) z ac Myriophyllum spicatum L. (Myr spi) z sr Myriophyllum verticillatum L. (Myr ver) z sr Nuphar lutea (L.) Sibth. & Sm. (Nup lut) z fl Potamogeton alpinus Balb. (Pot alp) z sr Potamogeton berchtoldii Fieber (Pot ber) z sr Potamogeton x fluitans Roth (Pot xfl) z sr Potamogeton crispus L. (Pot cri) z sr Potamogeton lucens L. (Pot luc) z sr Potamogeton natans L. (Pot nat) z fl Potamogeton nodosus Poir. (Pot nod) z fl Potamogeton x schreberi G. Fisch. (Pot xsc) z fl Potamogeton perfoliatus L. (Pot per) z sr Potamogeton pectinatus L. (Pot pec) z sr Ranunculus fluitans Lam. (Ran flu) z sr Ranunculus peltatus Schrank (Ran pel) z sr/fl Ranunculus trichophyllus Chaix (Ran tri) z sr Ranunculus spec. (Ran spe) z Spirodela polyrhiza (L.) Schleid. (Spi pol) z ac

Amphiphytes Agrostis stolonifera L. (Agr sto) z am Alisma plantago-aquatica L. (Ali p-a) z am Angelica archangelica ssp. litoralis (Fr.) Thell. (Ang arc) z am, h Barbarea vulgaris R. Br. (Bar vul) z am, h Berula erecta (Huds.) Coville (Ber ere) z am Butomus umbellatus L. (But umb) z am Caltha palustris L. (Cal pal) z am Glyceria declinata Bréb. (Gly dec) z am Glyceria fluitans (L.) R. Br. (Gly flu) z am Glyceria maxima (Hartm.) Holmb. (Gly max) z am, h Mentha aquatica L. (Men aqu) z am Myosotis scorpioides L. (Myo sco) z am Nasturtium officinale R. Br. (Nas off) z am Persicaria amphibia (L.) Delarbre (Per amp) z am, f Persicaria lapathifolia (L.) Delarbre (Per lap) z am Phalaris arundinacea L. (Pha aru) z am, h Rorippa amphibia (L.) Besser (Ror amp) z am Rorippa palustris (L.) Besser (Ror pal) z am Sagittaria sagittifolia L. (Sag sag) z am Tab. 1 Schoenoplectus lacustris (L.) Palla (Sch lac) z am, h Sparganium emersum Rehmann (Spa eme) z am Sparganium erectum L. (Spa ere) z am, h Veronica anagallis-aquatica (Ver a-a) z am Veronica beccabunga (Ver bec) z am

Helophytes Bidens tripartita L. (Bid tri) z he Carex acuta L. (Car acu) z he Carex elata L. (Car ela) z he Carex buekii Wimm. (Car bue) z he Carex paniculata L. (Car pan) z he Eleocharis ovata (Roth) Roem. & Schult. (Ele ova) z he Epilobium hirsutum L. (Epi hir) z he Epilobium roseum Schreb. (Epi ros) z he Equisetum fluviatile L. (Equ flu) z he Eupatorium cannabinum L. (Eup can) z he Filipendula ulmaria (L.) Maxim. (Fil ulm) z he Gnaphalium uliginosum L. (Gna uli) z he Impatiens glandulifera Royle (Imp gla) z he Impatiens noli-tangere L. (Imp n-t) z he Iris pseudacorus L. (Iri pse) z he Juncus bufonius L. (Jun buf) z he Juncus effusus L. (Jun eff) z he Juncus inflexus L. (Jun inf) z he Lysimachia vulgaris L. (Lys vul) z he Lythrum salicaria L. (Lyt sal) z he Phragmites australis (Cav.) Trin. ex Steud. (Phr aus) z he Poa palustris L. (Poa pal) z he Poa trivialis L. (Poa tri) z he Ranunculus sceleratus L. (Ran sce) z he Rumex aquaticus L. (Rum aqu) z he Rumex hydrolapathum Huds. (Rum hyd) z he Rumex palustris Sm. (Rum pal) z he Scirpus sylvaticus L. (Sci syl) z he Scrophularia nodosa L. (Scr nod) z he Solanum dulcamara L. (Sol dul) z he Stachys palustris (Sta pal) z he Symphytum officinale L. (Sym off) z he Typha latifolia L. (Typ lat) z he Valeriana officinalis L. (Val off) z he

Haptophytes Chlorophyta indet (Chl ind) z Fontinalis antipyretica Hedw. (Fon ant) z

ac: acropleustophyte am: amphiphytes sp: submersed pleustophyte he: helophytes sr: submersed rhizophyte oe: other emergent species fl: floating-leaved rhizophyte