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Contents - LAC Biosafety

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feeding with no significant impact to occasional large-scale outbreaks resulting<br />

in massive tree mortality may occur in natural tropical forests. However, the<br />

frequency and severity of pest damage is greater in plantations as summarised in<br />

Chapter 5 and described in detail in Chapter 10. Chapter 4 also showed that the<br />

most common insect outbreaks in natural forests occurred in high-density<br />

stands approaching monoculture.<br />

A detailed analysis of the plantation effect on pest incidence in tropical tree<br />

species was made by Nair (2001a). He compared the pest incidence in natural<br />

forests and plantations of several species for which relevant published literature<br />

was available—Eucalyptus spp., Gmelina arborea, Hevea brasiliensis, Swietenia<br />

macrophylla and Tectona grandis, and found that all of them suffered greater pest<br />

damage in plantations. In a meta-analysis of 54 individual studies reported in<br />

the literature, Jactel et al. (2005) also concluded that, overall, forest monocultures<br />

are more prone to pest infestation than more diverse forests. Thus the<br />

greater pest incidence in plantations is an undisputed scientific fact.<br />

Two main hypotheses have been proposed to explain the lower pest incidence<br />

in natural forests – the ‘enemies hypothesis’ and the ‘resource concentration<br />

hypothesis’ (Root, 1973; Carson et al., 2004). Recently, Nair (unpublished)<br />

proposed a third hypothesis called the ‘pest evolution hypothesis’.<br />

8.2.1 Enemies hypothesis<br />

According to the enemies hypothesis, the lower pest incidence in the<br />

mixed-species stand is due to greater action of the pests’ natural enemies. This is<br />

thought to be facilitated by the diverse plant community providing (1) alternative<br />

prey or hosts on which the natural enemies can sustain themselves and build up<br />

during periods when the pest is not present in the habitat, (2) a better supply of<br />

food such as pollen, nectar and honeydew for the natural enemies that enhances<br />

their fecundity and longevity and therefore overall effectiveness and (3) greater<br />

variation in microhabitats and microclimate that provides a larger variety of<br />

shelters for natural enemies. The increased natural enemy effectiveness<br />

therefore is thought to prevent pest build-up in the natural forest.<br />

8.2.2 Resource concentration hypothesis<br />

8.2 Do plantations suffer greater pest damage? 135<br />

According to the resource concentration hypothesis (Root, 1973), also<br />

called host concentration hypothesis (Carson et al., 2004), monoculture favours<br />

pest build-up by providing (1) a larger absolute supply of food resources,<br />

(2) greater ease in host location due to the physical proximity of the host trees<br />

and absence of interfering non-host volatiles and (3) reduced dispersal of the<br />

pests from the host patch. Arresting the dispersal, i.e. curbing the tendency of<br />

the herbivores that arrive on a clump of host plants to leave the area, appears to

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