1.3.4. Some reasons for insect species richness

Whatever the global estimate is, insects surely are remarkably speciose. This high species richness has been attributed to several factors. The small size of insects, a limitation imposed by their method of gas exchange via tracheae, is an important determinant. Many more niches exist in any given environment for small organ- isms than for large organisms. Thus, a single acacia tree, that provides one meal to a giraffe, may support the complete life cycle of dozens of insect species; a lycaenid butterfly larva chews the leaves, a bug sucks the stem sap, a longicorn beetle bores into the wood, a midge galls the flower buds, a bruchid beetle destroys the seeds, a mealybug sucks the root sap, and several wasp species parasitize each host-specific phytophage. An adjacent acacia of a different species feeds the same giraffe but may have a very different suite of phytophagous insects. The environment can be said to be more fine-grained from an insect perspective compared to that of a mammal or bird.

Small size alone is insufficient to allow exploitation of this environmental heterogeneity, since organisms must be capable of recognizing and responding to environmental differences. Insects have highly organized sensory and neuro-motor systems more comparable to those of vertebrate animals than other invertebrates. However, insects differ from vertebrates both in size and in how they respond to environmental change. Generally, vertebrate animals are longer lived than insects and individuals can adapt to change by some degree of learning. Insects, on the other hand, normally respond to, or cope with, altered conditions (e.g. the application of insecticides to their host plant) by genetic change between generations (e.g. leading to insecticide- resistant insects). High genetic heterogeneity or elasticity within insect species allows persistence in the face of environmental change. Persistence exposes species to processes that promote speciation, predominantly involving phases of range expansion and/or subsequent fragmentation. Stochastic processes (genetic drift) and/or selection pressures provide the genetic alterations that may become fixed in spatially or temporally isolated populations.

Insects possess characteristics that expose them to other potential diversifying influences that enhance their species richness. Interactions between certain groups of insects and other organisms, such as plants in the case of herbivorous insects, or hosts for parasitic insects, may promote the genetic diversification of eater and eaten. These interactions are often called coevolutionary and are discussed in more detail in Chapters 11 and 13. The reciprocal nature of such interactions may speed up evolutionary change in one or both partners or sets of partners, perhaps even leading to major radiations in certain groups. Such a scenario involves increasing specialization of insects at least on plant hosts. Evidence from phylogenetic studies suggests that this has happened — but also that generalists may arise from within a specialist radiation, perhaps after some plant chemical barrier has been overcome. Waves of specialization followed by breakthrough and radiation must have been a major factor in promoting the high species richness of phytophagous insects.

Another explanation for the high species numbers of insects is the role of sexual selection in the diversification of many insects. The propensity of insects to become isolated in small populations (because of the fine scale of their activities) in combination with sexual selection (section 5.3) may lead to rapid alteration in intraspecific communication. When (or if ) the isolated population rejoins the larger parental population, altered sexual signaling deters hybridization and the identity of each population (incipient species) is maintained in sympatry. This mechanism is seen to be much more rapid than genetic drift or other forms of selection, and need involve little if any differentiation in terms of ecology or non-sexual morphology and behavior.

Comparisons amongst and between insects and their close relatives suggest reasons for insect diversity. We can ask what are the shared characteristics of the most speciose insect orders, the Coleoptera, Hymenoptera, Diptera, and Lepidoptera? Which features of insects do other arthropods, such as arachnids (spiders, mites, scorpions, and their allies) lack? No simple explanation emerges from such comparisons; probably design features, flexible life-cycle patterns and feeding habits play a part (some of these factors are explored in Chapter 8). In contrast to the most speciose insect groups, arachnids lack winged flight, complete transformation of body form during development (metamorphosis) and dependence on specific food organisms, and are not phytophagous. Exceptionally, mites, the most diverse and abundant of arachnids, have many very specific associations with other living organisms.

High persistence of species or lineages or the numerical abundance of individual species are considered as indicators of insect success. However, insects differ from vertebrates by at least one popular measure of success: body size. Miniaturization is the insect success story: most insects have body lengths of 1–10 mm, with a body length around 0.3 mm of mymarid wasps (parasitic on eggs of insects) being unexceptional. At the other extreme, the greatest wingspan of a living insect belongs to the tropical American owlet moth, Thysania agrippina (Noctuidae), with a span of up to 30 cm, although fossils show that some insects were appreciably larger than their extant relatives. For example, an Upper Carboniferous silverfish, Ramsdelepidion schusteri (Zygentoma), had a body length of 6 cm compared to a modern maximum of less than 2 cm. The wingspans of many Carboniferous insects exceeded 45 cm, and a Permian dragonfly, Meganeuropsis americana (Protodonata), had a wingspan of 71 cm. Notably amongst these large insects, the great size comes predominantly with a narrow, elongate body, although one of the heaviest extant insects, the 16 cm long hercules beetle Dynastes hercules (Scarabaeidae), is an exception in having a bulky body.

Barriers to large size include the inability of the tracheal system to diffuse gases across extended distances from active muscles to and from the external environment (Box 3.2). Further elaborations of the tracheal system would jeopardize water balance in a large insect. Most large insects are narrow and have not greatly extended the maximum distance between the external oxygen source and the muscular site of gaseous exchange, compared with smaller insects. A possible explanation for the gigantism of some Palaeozoic insects is considered in section 8.2.1.

In summary, many insect radiations probably depended upon (a) the small size of individuals, combined with (b) short generation time, (c) sensory and neuro-motor sophistication, (d) evolutionary inter- actions with plants and other organisms, (e) metamorphosis, and (f ) mobile winged adults. The substantial time since the origin of each major insect group has allowed many opportunities for lineage diversification (Chapter 8). Present-day species diversity results from either higher rates of speciation (for which there is limited evidence) and/or lower rates of species extinction (higher persistence) than other organisms. The high species richness seen in some (but not all) groups in the tropics may result from the combination of higher rates of species formation with high accumulation in equable climates.

Chapter 1