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408<br />

D. Babendreier<br />

strated that parasitism of non-target species under field conditions is zero or<br />

restricted to a few meters from the release field (Orr et al. 2000; Babendreier<br />

et al. 2003a, 2003b). All these studies concluded that non-target effects of the<br />

biological control agent have been negligible, which is remarkable in light of<br />

the fact that such effects were presumed in these cases.Although still no comprehensive<br />

answer is possible on whether we have missed many non-target<br />

effects, evidence is increasing from the abovementioned studies and others<br />

that at least the more serious effects would have been detected.<br />

23.4 Harmonia axyridis, a Case Study<br />

Already in the section above it was mentioned that especially the polyphagous<br />

predators have the potential to cause serious non-target effects if introduced<br />

into foreign countries. The multicoloured Asian ladybeetle, Harmonia axyridis,<br />

is native to large parts of Asia and definitely is a polyphagous predator.<br />

It is a voracious feeder, preying mostly on aphids but also on immature stages<br />

of various other insects – psyllids, butterflies and aphid predators, including<br />

other ladybeetles. Moreover, cannibalism is often observed in adults and larvae<br />

of H. axyridis, which consume conspecific eggs and sometimes smaller<br />

larvae. Due to its effects on native coccinellids in the US and its recent invasion<br />

in Europe, H. axyridis has attracted quite some attention. Many studies<br />

have already been conducted and, thus, H. axyridis may serve as a case study<br />

to be examined in greater detail.<br />

Since H. axyridis feeds voraciously on aphids and has strong dispersal<br />

capacities, it was released as a classical biological control agent in the United<br />

States already in 1916. However, these early introductions failed and periodic<br />

releases continued until the 1980s. First established populations were documented<br />

only in 1988 and, subsequently, the beetle spread rapidly across North<br />

America. Currently, H. axyridis occurs throughout much of the continental<br />

United States and also in southern Canada. The ability of H. axyridis to feed<br />

on many non-target herbivores, intraguild prey and conspecifics may contribute<br />

to its success to invade new ecosystems but also to control aphid numbers.<br />

After the introduction in North America, it provided control of pests in<br />

several systems. For instance, H. axyridis provides effective biological control<br />

of Aphis spiraecola in apple orchards (Brown 2004), and the biological control<br />

of several citrus pests may also be benefiting from the establishment of H.<br />

axyridis (Michaud 2002a). Harmonia axyridis has also been utilized successfully<br />

in augmentative biological control in Asia, Europe and North America. In<br />

Europe, releases were first undertaken in 1982, i.e. before the negative impact<br />

had emerged, but continued until only a few years ago.<br />

However, the same attributes responsible for the effectiveness of H.<br />

axyridis against aphids may also cause less desirable effects. A number of

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