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Pros and Cons of Biological Control 409<br />

recent studies indicated that the establishment of exotic ladybeetles may<br />

have adverse affects on native coccinellids. In South Dakota, the abundance<br />

of Coccinella transversoguttata richardsoni and Adalia bipunctata was<br />

approximately 20 times lower after the establishment of an exotic coccinellid,<br />

C. septempunctata (Elliott et al. 1996). Evidence is building to indicate<br />

that H. axyridis may be having similar adverse effects on native Coccinellidae.<br />

Over a 13-year period, Brown and Miller (1998) monitored the<br />

abundance of various coccinellid species in apple orchards. The abundance<br />

of native coccinellids decreased after the establishment and rapid rise to<br />

dominance of the exotics, C. septempunctata and H. axyridis (Brown and<br />

Miller 1998). A 9-year study of the abundance of various Coccinellidae in an<br />

agricultural landscape showed a decrease in the abundance of Brachiacantha<br />

ursina, Cycloneda munda and Chilocorus stigma after the establishment<br />

of H. axyridis (Colunga-Garcia and Gage 1998). Similarly, a field study conducted<br />

in the major citrus-producing regions of Florida over 5 years showed<br />

that the introduced H. axyridis has increased in abundance while the formerly<br />

dominant ladybeetle Cycloneda sanguinea has declined (Michaud<br />

2002b). The author concluded that competitive displacement of C. sanguinea<br />

by H. axyridis may be in progress in this citrus ecosystem in Florida.<br />

Recently, a study conducted on the coccinellid community inhabiting potato<br />

crops in northern Maine over 31 years showed dramatic changes. Prior to<br />

the arrival of the exotic species, ladybeetle communities were comprised<br />

almost exclusively of the two native ladybeetles, Coccinella transversoguttata<br />

and Hippodamia tredecimpunctata. Starting 1980, the exotic C. septempunctata<br />

became permanently established and quickly began to dominate the<br />

ladybeetle community (Alyokhin and Sewell 2004). Two other exotic species,<br />

H. axyridis and Propylea quatuordecimpunctata, became prominent members<br />

of the ladybeetle community in 1995 and 1996. Altogether, these studies<br />

conducted in the US demonstrate the profound effects that the exotic natural<br />

enemy H. axyridis may have on the abundance of native coccinellid<br />

species.<br />

To date, research has concentrated on the effect of common natural enemies<br />

of agricultural crops because of fears regarding the loss of biocontrol<br />

function. As a consequence, there is a paucity of studies on the potential<br />

adverse effects of H. axyridis on coccinellids in non-agricultural settings or<br />

generally on less common coccinellids. In addition, few studies have been<br />

conducted on other non-target insects. In one of those few studies, Koch et al.<br />

(2003) recently identified H. axyridis as a potential hazard to immature<br />

Monarch butterflies, Danaus plexippus. In laboratory and field-cage studies,<br />

eggs and larvae of D. plexippus incurred significant predation by H. axyridis<br />

adults and larvae.A follow-up study showed that this effect is mediated by the<br />

presence of aphids, the preferred prey of H. axyridis. However, although predation<br />

on the monarch decreased in the presence of aphids, some monarch<br />

larvae were nevertheless consumed (Koch et al. 2005).

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