You also want an ePaper? Increase the reach of your titles
YUMPU automatically turns print PDFs into web optimized ePapers that Google loves.
106<br />
R. CAMPOS-HERRERA ET AL.<br />
Heterorhabditis <strong>and</strong> Steinernema, respectively. The bacteria are carried in the nematode<br />
intestine <strong>and</strong> released into the host upon infection (Poinar, 1990). Insect death usually<br />
occurs within 72 h of infection, <strong>and</strong> the nematodes feed, develop to adults, <strong>and</strong><br />
reproduce within the host, often completing 2–3 generations, before producing a new<br />
generation of specialized IJs, which emerge from the cadaver <strong>and</strong> seek new insect hosts<br />
(Poinar, 1990). Because of their established safety to non-target <strong>org</strong>anisms, EPNs are<br />
exempt from pesticide registration with the Environmental Protection Agency in the<br />
United States <strong>and</strong> are similarly exempt in many other countries.<br />
EPNs have been used effectively in classical biological control programs. For<br />
example, Steinernema scapterisci was discovered in Uruguay, near the presumed<br />
center of origin of the mole cricket (Scapteriscus spp.). The nematode was<br />
introduced into Florida where it is now established <strong>and</strong>, in combination with<br />
introduced parasitoids, provides effective biological control of several invasive mole<br />
cricket species (Nguyen & Smart, 1990; Adjei, Smart, Frank, & Leppla, 2006). Most<br />
commonly, however, EPNs are used for augmentation biological control where they<br />
function as biopesticides <strong>and</strong> treatment efficacy is expected to be of short duration,<br />
typically a matter of 1–2 weeks (McCoy et al., 2000; Duncan et al., 2003, 2007). In<br />
such programs, EPNs function in the same manner as non-persistent chemical<br />
pesticides <strong>and</strong> are applied repeatedly, as needed.<br />
Surveys have shown that some naturally occurring EPNs are more abundant in<br />
undisturbed compared to intensively farmed habitats (Campos-Herrera et al., 2008), <strong>and</strong><br />
that use of various animal manure mulches can increase the prevalence of some EPNs<br />
(Bednarek & Gaugler, 1997; Duncan et al., 2007). Such observations suggest the<br />
possibility of developing conservation biological control tactics <strong>by</strong> which agricultural or<br />
other managed habitats are modified in ways that enhance natural control of arthropods<br />
<strong>by</strong> EPNs. However, reliable <strong>and</strong> effective conservation biological control tactics are as<br />
yet unknown <strong>and</strong> remain a matter of speculation (Lewis, Campbell, & Gaugler, 1998;<br />
Stuart, Barbercheck, Grewal, Taylor, & Hoy, 2006).<br />
In the Florida citrus industry, Steinernema carpocapsae was the first nematode to<br />
be developed commercially for root weevil control (Schroeder, 1987; Figueroa &<br />
Roman, 1990; Smith, 1994) but a further discovered species, S. riobrave (formerly S.<br />
riobravis) (Cabanillas, Poinar, & Raulston, 1994), was found to cause greater D.<br />
abbreviatus mortality (Schroeder, 1994; Duncan et al., 1996; Bullock et al., 1999) with<br />
some field studies reporting ~90% suppression (Duncan & McCoy, 1996; Duncan et<br />
al., 1996; Bullock et al., 1999). However, estimates of the efficacy <strong>and</strong> profitability of<br />
using EPNs for weevil control in citrus vary widely <strong>and</strong> probably reflect variation in<br />
factors such as product quality, application rates, suitability of edaphic conditions for<br />
EPNs, <strong>and</strong> experimental methods (Adair, 1994; Duncan et al., 1996; Duncan, Graham,<br />
& Zellers, 2002; Duncan et al., 2003; 2007; Bullock et al., 1999; Stansly et al., 1997;<br />
McCoy et al., 2000; McCoy, Stuart, Duncan, & Nguyen, 2002). Currently, S. riobrave<br />
is marketed in Florida under the br<strong>and</strong> name Bio Vector 355 (Becker Underwood Inc.,<br />
Ames, IA) <strong>and</strong>, in 1999, approximately 19,000 ha of citrus were treated with this<br />
product to control citrus root weevils (Dimock, personal communication).<br />
Despite the evident <strong>and</strong> long-recognized potential of EPNs for insect pest<br />
management <strong>and</strong> the commercial development of effective EPN products for<br />
augmentation, their market penetration <strong>and</strong> incorporation into broadly applied IPM