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PREDATORY MITES IN IPM<br />

317<br />

of sex allocation, sex ratios approached half males, half females under r<strong>and</strong>om<br />

mating, whereas a female bias was observed under sib-mating. It is suggested that<br />

arrhenotoky is selected for when there is a substantial risk of high portion of<br />

unmated females in the population: pseudo-arrhenotoky may evolve <strong>by</strong><br />

external/environmental pressure, since it retains the possibility to reinstall lost genetic<br />

information in the maternally derived chromosome, <strong>by</strong> using the paternal chromosome<br />

as a template for DNA-repair. This precise control of sex allocation in phytoseiids is<br />

probably the significant mechanism <strong>by</strong> which these predators can regulate their<br />

density, depending on both phytoseiid female <strong>and</strong> prey densities. They can in fact<br />

adjust offspring sex ratio in response to the presence of conspecifics or their cues <strong>and</strong><br />

also to synchronize their population with that of prey (Nagelkerke & Sabelis, 1996).<br />

The prediction of optimal sex ratio <strong>by</strong> means of modeling is selectively advantageous<br />

when local mating groups vary in size <strong>and</strong> are usually small, as in the case of<br />

experimental laboratory studies on phytoseiids. At a larger spatial scale than the local<br />

mating group, the prediction appears to be less precise, may be due to operating <strong>and</strong><br />

interfering selection levels (Nagelkerke & Sabelis, 1998).<br />

3.3. Life Style Types<br />

The characterization <strong>and</strong> ranking of phytoseiid mites is a helpful <strong>and</strong> significant tool<br />

(McMurtry & Croft, 1997), allowing a rating of species based on some<br />

morphological, reproductive <strong>and</strong> developmental aspects. Furthermore, traits like<br />

feeding <strong>and</strong> diet needs (McMurtry & Rodriguez, 1987; Schausberger & Croft, 1999)<br />

<strong>and</strong> adaptation to certain foods (Castagnoli, Simoni, & Liguori, 2003), were<br />

considered to ascribe the predators to the different life styles. McMurtry <strong>and</strong> Croft<br />

(1997) considered that a four-types classification may be efficient to rank the<br />

different species of phytoseiids <strong>and</strong> gave emphasis to the biological control of spider<br />

mite pests (Table 2). However, new life style types might be identified.<br />

Recent studies aimed at estimating/identifying the more significant traits<br />

involved in phytoseiids rating <strong>and</strong> to possibly generate more stable classifications<br />

(Croft, Blackwood, & McMurtry, 2004). The mainly <strong>and</strong> first exploited traits/factors<br />

to rate phytoseiids were: (i) Feeding: the ability to prey <strong>and</strong> to feed on various prey<br />

<strong>and</strong> other food types (McMurtry & Rodriguez, 1987; McMurtry, 1992), a primary<br />

criterion considered for classification; (ii) External morphology: apparently, no<br />

strong correlation was found between body size <strong>and</strong> generalist–specialist phytoseiids<br />

(Schuster & Pritchard, 1963; Chant & Hansell, 1971) <strong>and</strong> more evidence is needed<br />

to establish the association between body size <strong>and</strong> different life styles (Croft et al.,<br />

2004); more correspondence was found in the evaluation of the adult dorsal shield<br />

setal length, in association with the feeding specialization (Sabelis & Bakker, 1992).<br />

Also, the mouthpart apparatus may be different between specialists <strong>and</strong> generalists<br />

(Flechtmann & McMurtry, 1992), but it is not clear to what extent this could work<br />

for a clear attribution to a life style; (iii) Biological parameters: research focused on<br />

reproduction, development <strong>and</strong> mortality; the specialist phytoseiids generally show<br />

intrinsic rates (Sabelis & Janssen, 1994) <strong>and</strong> sex ratio values higher than generalist<br />

(Nagelkerke & Sabelis, 1996) as well as shorter developmental times (Luh & Croft,

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