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toxic effects detected on the survival or reproduction of
F. candida
or
X. griesea
(Sims and
Martin, 1997) and no effects of purified Cry2A on
P. saber
(Sims, 1997).
Only in a few studies have negative effects of
Bt
proteins on microarthropods been
reported: Wandeler et al. (2002) found that
P. scaber
fed significantly less on
Bt
maize
expressing Cry1Ab than on non-
Bt
maize during a 20-day feeding trial in the labora-
tory in soil microcosms; Griffiths et al. (2006) reported lower collembolan abundance
and higher mite populations in soil cultivated with
Bt
maize expressing Cry1Ab than in
soil cultivated with non-
Bt
maize in the laboratory; and Bakonyi et al. (2006) found that
there were species-specific effects in feeding preference when the collembolans
F. candida
,
Heteromurus nitidus,
and
Sinella coeca
were fed dried leaves of
Bt
(Cry1Ab) or non-
Bt
maize
in laboratory feeding tests—
F. candida
preferred non-
Bt
maize as a food source over
Bt
maize, but there was no difference in feeding preference for
H. nitidus
and
S. coeca
. Cortet
et al. (2007) detected a minor negative effect of
Bt
maize (Cry1Ab) cultivation on micro-
arthropod abundance (mites and collembolans) in high-clay soils in field trials; however,
agricultural practices had a comparable or greater effect on microarthropod abundance
than the
Bt
crop. Debeljak et al. (2007) also reported a lower abundance of collembolans
in field soil cultivated in
Bt
maize (Cry1Ab) but only at one site and only in early fall; no
difference in functional groups of Collembola was detected.
of Bt crops on microarthropods
In general, cultivation of transgenic
Bt
crops,
Bt
plant biomass, or purified
Bt
proteins,
including Cry1Ab, Cry1Ac, and Cry3Bb1, has had little to no effect on most microarthro-
pods tested. When effects have been reported, they were minor, and the microarthropods
were often more affected by temporal differences in sampling time, agricultural practices,
plant varietal differences, or other biotic and abiotic factors not related to a particular
Bt
protein. Although most studies have not shown a consistent pattern of
Bt
effects on micro-
arthropods, longer-term field experiments would improve the current understanding
Bt
effects on the abundance and diversity of microarthropods in the soil environment (e.g.,
Theissen and Russell, 2009). Future research studies should focus on multitrophic-level
interactions and continue to evaluate the effects of new and different types of
Bt
crops on
nontarget microarthropods in the soil.
8.4.6
Effects of the cultivation of Bt crops on nontarget
Lepidopteran larvae and aquatic insects
Although most insects are not generally classified as soil organisms per se, many insect
orders (e.g., Lepidoptera, Coleoptera, and Hymenoptera) have a larval stage that lives in
or feeds in close proximity to
Bt
plant leaves and other residues,
Bt
pollen, or Cry proteins
in soil. Thus, it is possible for nontarget insects, including pollinators, prey and predator
species, and biocontrol agents, to be affected by the cultivation of
Bt
crops, even if they
do not feed on the GM crop directly. As the number of studies evaluating the effects of
Bt
crops on nontarget insects are too numerous to review in this chapter, only the nontarget
effects of the cultivation of
Bt
crops on selected species of Lepidoptera and aquatic insects
are summarized here (
Table 8.9)
. More information on the nontarget effects of the cultiva-
tion of
Bt
crops on a variety of insect species can be found in several reviews (including
those of Romeis et al., 2006; Marvier et al., 2007; Thies and Devare, 2007; Duan et al., 2008;
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