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Many basic and applied studies in insect ecology have focused on the proximate and ultimate responses of insect populations to their physical and chemical environment (2, 15, 21, 27, 32, 74, 82, 111, 112). From an economic perspective, macro- and microclimatic factors can influence the stress that insect populations inflict on plants and the efficacy of management tactics. For above-ground insects, the mechanisms of behavioral response to environmental factors are often observable, if not always apparent to the researcher. However, this is not typically the situation with soil insects. As a result, field studies of soil insects often quantify only the consequences of behavior while the behaviors themselves remain hidden within the soil matrix (14, 103, 105).
Soil ecology research has been productive at the ecosystem level on such topics as nutrient cycling (18), arthropod regulation of micro- and meso-fauna in below-ground detrital food webs (75), impact of microfauna on soil genesis and structure (87), rhizosphere dynamics (17), and energy dynamics of soil systems (79). These examples highlight the importance of multidisciplinary approaches to research programs that unite expertise in insect ecology, soil physics, chemistry, and microbiology as well as systems analysis and modeling (87). Considerable interest also exists in the relationships within soil communities, but these studies have focused primarily upon nonagricultural systems (71, 108) and on the more abundant microarthropod members of the soil fauna ( 106-109).
Ecological, morphological, and physiological adaptations of nonagricultural soil arthropods have been discussed in the literature (8, 10, 26, 59); however, insects that are agricultural pests primarily in their immature soil-inhabiting stages have often been studied in detail only in their more accessible adult stage. Although the mobile adult stages of soil pests often determine initial habitat and host selection, a considerable proportion of subsequent host and habitat selection is performed by immatures in the soil, if host or habitat quality deteriorates over time.
A major obstacle to the study of soil insect ecology has been the inability to follow soil insect movement and feeding behavior in situ (3, 14, 33, 34, 103, 105). It is critically important in these studies to minimize the disturbance of the soil system through experimental manipulations. R. L. Rabb (cited in 103) notes that the greatest problem with studies of soil insects is that the system is altered through its study. Also, research workers often fail to consider dominant mass and energy transport mechanisms in soil ecosystems. Differences in above- and below-ground environments may alter soil insect sensitivity (over ecological and evolutionary time) to shifting environmental conditions, the movement of chemical cues from potential food sources to soil herbivores, and the mechanisms for soil insect host-finding behavior when compared to terrestrial organisms.
In this review we briefly outline several basic principles of soil physics as they relate to soil insect movement and host-finding behavior, to provide a general understanding of the environment in which soil macroarthropods exist. We then selectively review the entomological literature in light of these principles to stress the need to evaluate soil insect behavior within the soil matrix when trying to understand the underlying mechanisms that produce observable behavior. Finally we briefly discuss the importance of behavior in the management of soil insects.