Annual Review of Entomology - Volume 44, 1999

The traditional use of insects as food continues to be widespread in tropical and subtropical countries and to provide significant nutritional, economic and ecological benefits for rural communities. Westerners should become more aware of the fact that their bias against insects as food has an adverse impact, resulting in a gradual reduction in the use of insects without replacement of lost nutrition and other benefits.

Over the last four decades, a number of arthropod-borne infections have been recognized for the first time. Some have become of considerable public health importance, such as dengue hemorrhagic fever (DHF), and others are spreading geographically and their incidence is increasing. There has been an important recrudescence of several long-known vector-borne diseases. Malaria, leishmaniasis, dengue, and plague have resurged in numerous foci, in some cases where they were thought to be under effective control. In most instances, the appearance of new diseases and syndromes and the resurgence of old can be associated with ecological changes that have favored increased vector densities. Dam construction, irrigation and other development projects, urbanization, and deforestation have all resulted in changes in vector population densities that appear to have enabled the emergence of new diseases and the resurgence of old diseases. Greatly increased human travel has spread infectious agents, introducing them into areas in which they had been hitherto absent. It is essential to understand the factors that caused increased vector densities and hence the transmission of disease to prevent the emergence and resurgence of more diseases, as well as to serve as a basis for effective control.

Pigeonpea (Cajanus cajan) is an important crop in semi-arid tropical and subtropical farming systems, providing high quality vegetable protein, animal feed, and firewood. Insect pests feeding on flowers, pods, and seeds are the most important biotic constraint affecting pigeonpea yields. This review summarizes the biology and ecology of the three most important groups of pests: flower- and pod-feeding Lepidoptera, pod-sucking Hemiptera, and seed-feeding Diptera and Hymenoptera. Recent research investigating the complex interactions among pigeonpea, its key pests, and their natural enemies is also reviewed. These relationships have implications on the pest status of individual species and on possible control strategies. Pigeonpea pest management research has focused until recently on the identification and development of resistant cultivars and on chemical control. Future research must focus on environmentally sound pest management strategies that are compatible with the needs and limitations of pigeonpea farmers. Several priority areas for research are suggested.

In this review, we use the wing veins of dipteran insects as potential models for understanding the evolution of development. We briefly discuss previous work in this field and examine the genetic complexity of wing formation, discussing the genes involved in wing formation and their roles in Drosophila wing development and vein formation. Furthermore, patterns of wing vein formation, addition, and reduction are discussed as they occur throughout the Diptera. Using the phyletic phenocopy paradigm, we draw attention to many wing vein morphologies that phenocopy various wing mutants in Drosophila melanogaster. The systematic issues of the nature of characters, homology, and the role of modern developmental approaches to evolutionary studies, which has recently become important, can be addressed from the perspective of the wing. We argue that further developmental evolutionary studies, and the interpretation of data therefrom, must be conducted within the context of a well-supported phylogeny of the organisms under study.

The African mosquito species Anopheles gambiae sensu lato s.l. and Anopheles funestus rank among the world's most efficient vectors of human malaria. Their unique bionomics, particularly their anthropophilic, endophagic and endophilic characters, guarantee a strong mosquito-host interaction, favorable to malaria transmission. Olfactory cues govern the various behaviors of female mosquitoes and here we review the role of semiochemicals in the life history of African malaria vectors. Recent evidence points towards the existence of human-specific kairomones affecting host-seeking A. gambiae s.l., and efforts are under way to identify the volatiles mediating this behavior. Based on examples from other Culicidae spp., it is argued that there is good reason to assume that mating, sugar feeding, and oviposition behavior in Afrotropical malaria vectors may also be mediated by semiochemicals. It is foreseen that increased knowledge of odor-mediated behaviors will be applied in the development of novel sampling techniques and possibly alternative methods of intervention to control malaria.

This review summarizes the literature about pathogens and predators of ticks and their potential use as biocontrol agents published since the beginning of this century. In nature, many bacteria, fungi, spiders, ants, beetles, rodents, birds, and other living things contribute significantly toward limiting tick populations, as do, for instance, the grooming activities of hosts. Experiments with the most promising potential tick biocontrol agents—especially fungi of the genera Beauveria and Metarhizium and nematodes in the families Steinernematidae and Heterorhabditidae, as well as oxpeckers—are described.

Stingless bees (Apidae: Meliponini) are common visitors to flowering plants in the tropics, but evidence for their importance and effectiveness as crop pollinators is lacking for most plant species. They are known to visit the flowers of ∼90 crop species. They were confirmed to be effective and important pollinators of 9 species. They may make a contribution to the pollination of ∼60 other species, but there is insufficient information to determine their overall effectiveness or importance. They have been recorded from another 20 crops, but other evidence suggests that they do not have an important role because these plants are pollinated by other means. The strengths and limitations of stingless bees as crop pollinators are discussed. Aspects of their biology that impact on their potential for crop pollination are reviewed, including generalized flower visiting behavior of colonies, floral constancy of individual bees, flight range, and the importance of natural vegetation for maintaining local populations.

The family Anthocoridae (Hemiptera:Heteroptera) contains between 400 and 600 species distributed worldwide, chiefly on the continents but also on oceanic islands. They are small (1.4–4.5 mm) and common to a wide variety of habitats. Many are found in cryptic habitats such as galls, several widespread genera are surface feeders on small arthropods (Anthocoris, Orius, and Tetraphleps), and others can be found in ant nests and, especially, under bark. Wing polymorphism is common in this family, often associated with the cryptic habit. Most known species are predaceous, though some take plant food as well (e.g. Orius insidiosus, Orius pallidicornis). A few of these are believed to be entirely phytophagous (Paratriphleps laeviusculus). Their small size and often generalized feeding habits have resulted in about 30 introduced species, mostly accidental. A few have been introduced deliberately as biological control agents (Anthocoris spp., Montandoniola moraguesi, O. insidiosus, Orius tristicolor, and Tetraphleps spp.). Most nonindigenous species seem to have been distributed as a result of human activities, especially commerce. The predaceous habits of many Anthocoridae have attracted the attention of researchers who work in agroecosystems. Integrated pest management programs often include these predators, which has given us greater knowledge about these species than those found in natural ecosystems. Exciting discoveries about the attractiveness to these bugs of certain volatile plant and arthropod compounds are opening new areas of investigation into their chemical ecology. The reactions of these tiny predators will surely become better understood as a result.

For those arthropod species adapted for living below the soil surface, the soil is a refuge from the biotic and abiotic perturbations existing above ground. Convergent morphological, physiological, and behavioral adaptations of epedaphic, euedaphic, and hemiedaphic arthropods to selective aspects of subterranean existence are examined in light of overlapping ecological niches. The abiotic impact of the soil environment and its relationship to arthropod evolution, radiation, and ecology are discussed as well. Specific areas addressed include the invasion of land by marine arthropods, the impact of morphology on arthropod mobility, osmoregulatory/respiratory systems, and defensive strategies.

Baculoviruses, among other insect viruses, are regarded as safe and selective bioinsecticides, restricted to invertebrates. They have been used worldwide against many insect pests, mainly Lepidoptera. Their application as microbial pesticides, however, has not met their potential to control pests in crops, forests, and pastures, with the exception of the nuclear polyhedrosis virus of the soybean caterpillar (Anticarsia gemmatalis), which is used on approximately 1 million ha annually in Brazil. Problems that have limited expansion of baculovirus use include narrow host range, slow killing speed, technical and economical difficulties for in vitro commercial production, timing of application based on frequent host population monitoring, variability of field efficacy due to climatic conditions, and farmers' attitudes toward pest control, which have been based on application of fast-killing chemical insecticides. Farmer education regarding use of biological insecticides and their characteristics is considered one of the major actions necessary for increased use of baculoviruses. Strategies to counteract some of the limitations of baculoviruses, especially their slow killing activity, have been investigated and are promising. These include the use of chemical or biological substances added to virus formulations and genetic engineering of the viruses themselves to express insect toxins or hormones. Such strategies can enhance viral activity and increase speed of kill as well as reduce larval feeding activity. The use of baculoviruses against Lepidoptera is reviewed, with the utilization of the nuclear polyhedrosis virus of A. gemmatalis in Brazil serving as a case-study.

Hyperparasitoids are secondary insect parasitoids that develop at the expense of a primary parasitoid, thereby representing a highly evolved fourth trophic level. This review evaluates multitrophic relationships and hyperparasitoid ecology. First, hyperparasitoid communities of various taxa of phytophagous and predacious insects are described. Second, specific patterns of hyperparasitoid community organization and hyperparasitoid ecology are described in detail, using the aphid-parasitoid–hyperparasitoid food web as a model system. Aphid hyperparasitoid communities consist of ecto- and endohyperparasitoids, with ectohyperparasitoids being less host specific than endohyperparasitoids. Lifetime fecundity and intrinsic rate of increase of hyperparasitoids are generally lower than those of their primary hosts. Aphid ectohyperparasitoids search randomly for hosts and do not use specific cues, whereas endohyperparasitoids gain information that originates from host plants or hosts for long-range search. Interactions with adult primary parasitoids do not influence hyperparasitoid searches, but aphid-attending ants typically prevent successful hyperparasitoid foraging. Impact of hyperparasitism on biological control is reviewed.

Insects respond to crowding in a variety of ways that are usually exemplified by rapid changes in behavior and culminate in enduring long-term morphological and/or chromatic responses. A common feature of both short-term and long-term effects is that they are graded, dependent not only on density but also on the duration and on phase history of the maternal generation. Because of their exoskeletons, which are persistent for the duration of each instar and endure throughout adult life, overt changes in morphology or coloration are restricted to the molting period and shortly afterward, when cuticular hardening and pigmentation are expressed. Changes in internal organs or metabolism elicited by population density, being independent of integumental constraints, are not restricted to the molting period, but the temporal difference between internal and external responses is not of fundamental significance. Intraspecific responses to the presence of sibling insects are of apparent ecological significance and often involve directional movement and/or migration. They are mediated via the sensory system, involve signal transduction, and elicit downstream biochemical and physiological changes.

Cassava (Manihot esculenta) occupies a uniquely important position as a food security crop for smallholder farmers in areas of the tropics where climate, soils, or societal stresses constrain production. Given its reliability and productivity, cassava is the most important locally produced food in a third of the world's low-income, food-deficit countries. It is the fourth most important source of carbohydrates for human consumption in the tropics, after rice, sugar, and maize. World production of cassava from 1994–1996 averaged 166 million tons/year grown on 16.6 million hectares (ha), for an average yield of 9.9 tons/ha. Approximately 57% is used for human consumption, 32% for animal feed and industrial purposes, and 11% is waste. Africa accounts for 51.3% of the production; Asia, 29.4%; and Latin America, 19.3%. The area planted to cassava in Africa, Asia, and Latin America is 10.3, 3.7, and 2.6 million ha, respectively.

Mate choice theory has become a major field of research in behavioral ecology. Tree crickets provide excellent opportunities for studying the diversity and variability of mate choice. The evidence for female mate choice in tree crickets is reviewed, and broad comparisons with other orthopteran groups are made. The evidence shows that female choice may occur during several different stages of mating and may target several different criteria. Song preferences are perhaps dominated by stabilizing preferences for the cues of species recognition, but there is a growing body of evidence for directional preferences based on sensory biases or mate quality. Mate rejection during courtship and forms of postcopulatory choice may favor males, based both on phenotypic quality and on the amount of nutritious courtship gifts they provide, and may differ with the value of mating incentives. Understanding the balance and trade-offs between different forms of mate choice may help in understanding their evolutionary causes.

The order Diptera (true flies) is one of the most species-rich and ecologically diverse clades of insects. The order probably arose in the Permian, and the main lineages of flies were present in the Triassic. A novel recent proposal suggests that Strepsiptera are the sister-order to Diptera. Within Diptera, evidence is convincing for the monophyly of Culicomorpha, Blephariceromorpha, and Tipulomorpha but weak for the monophyly of the other basal infraorders and for the relationships among them. The lower Diptera (Nematocera) is paraphyletic with respect to Brachycera, and morphological evidence suggests the sister-group of Brachycera lies in the Psychodomorpha. Recent analyses suggest Tipulomorpha are closer to the base of Brachycera than to the base of Diptera. Brachycera are undoubtedly monophyletic, but relationships between the basal lineages of this group are poorly understood. The monophyly of Stratiomyomorpha, Xylophagomorpha, Tabanomorpha, and Muscomorpha is well supported. Eremoneura, and its constituent clades Empidoidea and Cyclorrhapha, are monophyletic. The sister-group of Eremoneura is likely to be part or all of Asiloidea. Several viewpoints on the homology of the male genitalia of eremoneuran flies are discussed. Phylogenetic analyses suggest that lower Cyclorrhapha (Aschiza) are paraphyletic; however, schizophoran monophyly is well supported. The monophyly of Acalyptratae is not well-founded and the relationships between acalyptrate superfamilies remain obscure. Recent advances document the monophyly of the families of Calyptratae and the relationships among them. Areas critical to future advances in understanding dipteran phylogeny include the relationships among the basal infraorders of Diptera and Brachycera and the relationships between the superfamilies of acalyptrates. Progress in dipteran phylogenetics will accelerate with the exploration of novel data sources and the formulation of hypotheses in an explicitly quantitative framework.

Examination of the function, chemistry, and pharmacology of the voltage-gated insect sodium channel (ISC) reveals that the ISC closely resembles its vertebrate counterpart in electrophysiology and ion conductance, primary structure and allocation of all functional domains, and its pharmacological diversity and flexibility exhibited by the occurrence of different allosterically coupled receptor-binding sites for various neurotoxicants.

The toxicants include several groups of insecticides, namely DDT and its analogues, pyrethroids, N-alkylamides, and dihydropyrazoles, which affect channel gating and ion permeability.

Despite their similarity, the insect and vertebrate channels are pharmacologically distinguishable, as revealed by the responsiveness of the heterologously expressed Drosophila para clone to channel modifiers and blockers and the occurrence of the insect-selective sodium channel neurotoxins derived from arachnid venoms presently used for the design of recombinant baculovirus-mediated selective bioinsecticides.

The pharmacological specificity of the ISC may lead to the design of insect-selective toxicants, and its pharmacological flexibility may direct the use of ISC insecticides for resistance management. Insecticide resistance [such as knockdown resistance (KDR)] is acquired by natural selection and operated by increased metabolism, channel mutagenesis, or both.

The resistance issue can be dealt with in several ways. One is by simultaneous application of low doses of synergistic, allosterically coupled mixtures (thus delaying or preventing the onset of resistance). An alternative is to replace an insecticide to which resistance was acquired by channel mutation with a different ISC toxicant to which increased susceptibility was conferred by the same mutation. Such a possibility was exemplified by a significant increase in susceptibility to N-alkylamides, as well as an insect-selective neurotoxin revealed by KDR insects. Third, both of these methods can be combined. Thus owing to its pharmacological uniqueness, the ISC may serve as a high-priority target for future selective and resistance-manageable insecticides.

In this article we consider the role of epidemiological factors and transmission processes of insect-vectored viruses on the effectiveness of insecticides in a disease management program. We also discuss the use of insecticides within the broader framework of the chemical environment surrounding vectors, and how chemical-induced alterations in the behavior of vectors can influence transmission. Our analysis confirms the belief of Heathcote, who stated in 1973, that “no one method of control is likely to keep crops entirely free from virus infection and as many preventative measures should be taken as are economically justified.”

This investigation into the natural history and behavior of 81 species of cerambycid beetles suggests that reproductive behavior is correlated with the condition of the larval host: Adults of species whose larvae attack living trees tend to show behavioral differences from those that attack dying or dead hosts. Behavioral differences among species that are associated with larval host condition include: (a) choice of adult food source and whether adults feed at all; (b) mechanisms of mate location and the role of long-range pheromones; (c) vagility and dispersal behaviors of adults; (d) location of the mating site; and (e) duration of copulation.