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Effects of Invasive Non-Native Species on the Native Biodiversity in the River Rhine 263<br />

50<br />

40<br />

30<br />

20<br />

10<br />

0<br />

1831-1850<br />

1851-1870<br />

1871-1890<br />

1891-1910<br />

1911-1930<br />

1931-1950<br />

1951-1970<br />

1971-1990<br />

1991-2005<br />

Cumulative number of species<br />

Fig. 15.2 Increasing number of non-native macroinvertebrate species colonising the<br />

river Rhine. Cumulative data are shown for periods of 20 years (note: the last bar<br />

includes data for only 15 years). The exponential model was fitted by least-squares<br />

regression (y=5.5936 x 10 –22 e 0.0265t , n=9, R 2 =0.98, t indicates the year). Data were<br />

obtained from Tittizer et al. (2000), Geitler et al. (2002) and Rey et al. (2004)<br />

duced by fishermen, another seven species (26 %) having been inadvertently<br />

introduced by the aquarium trade. Interestingly, among the phytoplankton,<br />

an ecologically important group, no non-native species have yet been<br />

observed in the Rhine (Nehring 2005).<br />

About one of two non-native aquatic species entering German rivers could<br />

spread over large areas, and about one of five non-native species have become<br />

invasive (Nehring 2003). In the Rhine delta in The Netherlands, the proportion<br />

of non-native species in the biodiversity of river channels and floodplain<br />

lakes ranges from 7–10 % among macrophytes to 5–12 % among macroinvertebrates<br />

and 17–19 % among fish (Van den Brink et al. 1996). In the Middle<br />

and Upper Rhine, non-native species represent 10–15 % of total species richness<br />

(Haas et al. 2002). Non-native species dominate in terms of total abundance<br />

and biomass, however, the values exceeding 80 % (Tittizer et al. 2000;<br />

Haas et al. 2002).<br />

Thus, species composition in the river Rhine has changed remarkably in<br />

the past four decades. Replacing characteristic riverine species, large numbers<br />

of euryoecious and non-native species, in particular macroinvertebrates and<br />

fish, have invaded this river system (e.g.Van den Brink et al. 1988, 1990). Some<br />

of the species entered the river via ports and estuaries, and then moved<br />

upstream whereas others moved downstream after entering via canals. Several<br />

of these species have penetrated into the larger, still-water expanses but

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