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Effects of Invasive Non-Native Species on the Native Biodiversity in the River Rhine 261<br />

Basel, the number of stonefly species declined from 13 to four between 1910<br />

and 1990, and those of mayflies from 19 to 13 (Küry 1994).<br />

The decline of the freshwater fauna in the river Rhine is linked to extensive<br />

habitat deterioration caused by channelisation and flow regulation by weirs,<br />

stream fragmentation, organic pollution from land-use activities, toxic contaminants<br />

from municipal and industrial sources, and interactions with an<br />

increasing number of non-native species (Streit 1992; Baur and Ringeis 2002;<br />

Van der Velde et al. 2002; Nehring 2003). Since the industrial revolution and<br />

the construction of sewage systems, domestic and industrial pollution have<br />

led to a gradual deterioration in water quality, and this from the second half of<br />

the 19th century to the end of the 1960s. Water quality was very poor during<br />

the period 1950–1970, with low oxygen levels, serious eutrophication, high<br />

chemical and organic pollution loads, salination caused by French potassium<br />

mines and mining water from brown coal mines in Germany, and thermal<br />

pollution (Rhine river water temperature has risen by approximately 2 °C<br />

above its natural value; Admiraal et al. 1993).<br />

Faunal diversity in the river Rhine was lowest in the late 1960s,when levels of<br />

toxicants were highest and oxygen levels extremely low (Kinzelbach 1972; Streit<br />

1992). During the period 1970–1986, waste water treatment plants were constructed<br />

along the river,resulting in improvements of water quality including an<br />

increase in oxygen levels and a reduction of some heavy metals and organic pesticides.<br />

Also, faunal diversity began to recover (Admiraal et al. 1993). Driven<br />

partly by the toxic spill following the Sandoz accident (see below), ministers<br />

from riparian countries decided in 1986 to establish the Rhine Action Programme.One<br />

of its aims is the restoration of the river ecosystem.<br />

Haas et al. (2002) described three successional phases in the development<br />

of benthic communities in the German section of the Rhine, following the<br />

extreme toxic and organic contamination which the river has known in earlier<br />

times.<br />

1. From 1970 to 1986, the aquatic community was species-poor and still in an<br />

early recovery. Because of the remaining organic pollution, only sewageresistant<br />

taxa such as the leech Erpobdella octoculata, the isopod Asellus<br />

aquaticus, the snail Radix ovata, sponges, chironomids and oligochaetes<br />

occurred. The non-native zebra mussel Dreissena polymorpha started to<br />

colonise hard substrates. However, the major Sandoz industrial accident<br />

near Basel in 1986, when runoff from water used in firefighting carried<br />

nearly 30 t of toxic chemicals (insecticides, fungicides and herbicides) into<br />

the Rhine, caused serious damage to the flora and fauna over hundreds of<br />

kilometres, resetting the recovery process. In 1987, benthic faunal densities<br />

were still close to zero (Den Hartog et al. 1992).Yet, D. polymorpha was able<br />

to quickly recolonise the Rhine following the Sandoz spill because of the<br />

immigration of pelagic larvae from unaffected sites.<br />

2. In 1987 and 1988, the non-native amphipod Corophium curvispinum<br />

(=Chelicorophium curvispinum), and the Asiatic clams Corbicula fluminea

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