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114<br />
R. CAMPOS-HERRERA ET AL.<br />
(1997) noted the apparent usefulness of composted animal manure for conservation<br />
biocontrol after showing that long-term applications of manure increased EPNs <strong>by</strong><br />
3-fold. Animal manure mulches likely increase availability of insect prey <strong>and</strong> were<br />
also shown to decrease the prevalence of some nematophagous fungi that prey on<br />
EPNs (Jaffee, Ferris, Stapleton, Norton, & Muldoon, 1994; Duncan et al., 2007). If a<br />
number of physical properties can be identified that consistently affect EPN spatial<br />
patterns then this information might be used to incrementally improve biological<br />
control achieved <strong>by</strong> either augmented or endemic EPNs.<br />
Underst<strong>and</strong>ing the basis of relationships between EPNs <strong>and</strong> other variables in<br />
the field can require a substantial research investment. Whereas EPN associations<br />
with some variables have a causative basis, others arise indirectly through the effects<br />
of unmeasured hidden variables. Therefore, identifying those variables that are most<br />
likely to modulate EPN prevalence or behavior in predictable ways is important. A<br />
comparison of the results of surveys using different sample criteria can be helpful in<br />
selecting variables of interest.<br />
Campos-Herrera et al. (2007) found that soil moisture was positively associated<br />
with EPN recovery frequency in a survey of 100 undisturbed sites (natural areas <strong>and</strong><br />
field borders) in La Rioja, Spain. However, in a temporal survey of 18 agricultural<br />
fields with different management regimes <strong>and</strong> bordering natural areas, EPNs were<br />
unrelated to soil moisture but inversely related to soil disturbance (Campos-Herrera et<br />
al., 2008). Because greater irrigation was employed in tilled annual cropping systems<br />
with few EPNs than in natural areas or <strong>org</strong>anically grown perennial crops with<br />
numerous EPNs, Campos Herrera et al. (2008) speculated that soil disturbance is more<br />
important than moisture in regulating EPN spatial patterns. Similarly, Campos-Herrera<br />
et al. (2008) reported linear correlations between EPNs <strong>and</strong> certain soil characteristics<br />
that were also studied <strong>by</strong> Alumai, Grewal, Hoy, <strong>and</strong> Willough<strong>by</strong>, (2006) <strong>and</strong> Duncan<br />
et al. (unpublished). Differences in the survey protocols among these studies suggest<br />
the possibility that some of the correlations between soil properties <strong>and</strong> EPNs are due<br />
to the influence of soil texture on both EPNs <strong>and</strong> certain chemical properties (Fig. 5).<br />
Because greater s<strong>and</strong> content of soil generally favors EPNs, chemical properties<br />
associated in some manner with s<strong>and</strong> content would likely be similarly associated with<br />
EPNs. The relationships measured in Ohio <strong>and</strong> Florida (shown in small font in Fig. 5)<br />
are strikingly similar. Organic matter, K + <strong>and</strong> Mg ++ tended to be negatively associated<br />
with both EPNs <strong>and</strong> percentage s<strong>and</strong> whereas P was positively associated with both<br />
variables. In contrast, these relationships had exactly opposite trends in the Spanish<br />
survey. Nevertheless, soil chemicals in each of the three surveys were associated with<br />
EPNs in the same manner that they were associated with percentage s<strong>and</strong>. The Ohio<br />
<strong>and</strong> Florida surveys occurred on several golf courses <strong>and</strong> within a single citrus<br />
orchard, respectively. Thus, cultural practices were similar among sites within each of<br />
these surveys, <strong>and</strong> the effects of porosity <strong>and</strong> soil colloid surfaces on leaching of some<br />
chemicals <strong>and</strong> the adsorption of others operated on similar levels of nutrient inputs.<br />
However, the Spanish survey compared EPN populations in natural areas <strong>and</strong> in a<br />
variety of different annual <strong>and</strong> perennial cropping systems. The very different levels of<br />
inputs between the Spanish sample sites might have caused different relationships<br />
between chemicals <strong>and</strong> percentage s<strong>and</strong> <strong>and</strong>, therefore, with EPNs than seen in Ohio<br />
or Florida.