Efnisyfirlit

• Front Matter
• Preface to the Fifth Edition
• Preface to the Fourth Edition
• Preface to the Third Edition
• Preface to the Second Edition
• Preface and Acknowledgments for First Edition
• About the Companion Website
• Chapter 1 The Importance, Diversity and Conservation of Insects
• 1.1 WHAT IS ENTOMOLOGY?
• 1.2 The importance of insects
• Box 1.1 Citizen entomologists—community participation
• 1.3 Insect biodiversity
• 1.3.1 The described taxonomic richness of insects
• 1.3.2 The estimated taxonomic richness of insects
• 1.3.3 The location of insect species richness
• 1.3.4 Some reasons for insect species richness
• Fig. 1.1 Speciescape, in which the size of individual organisms is approximately proportional to the number of described species in the higher taxon that it represents.
• 1.4 Naming and classification of insects
• Table 1.1 Taxonomic categories (obligatory categories are shown in bold).
• 1.5 Insects in popular culture and commerce
• Box 1.2 Butterfly houses
• 1.6 Culturing insects
• 1.7 Insect conservation
• Box 1.3 Tramp ants and biodiversity
• Box 1.4 Conservation of the large blue butterfly
• 1.8 Insects as food
• 1.8.1 Insects as human food: entomophagy
• Box 1.5 Palmageddon? Weevils in the palms
• Fig. 1.2 Mopane moths and host tree. (a) Larva of G. belina on mopane (Colophospermum mopane) leaves. (b) Adult of Gonimbrasia belina. (c) Distribution of mopane woodland in southern Africa.
• Fig. 1.3 A delicacy of the Australian Aborigines—a witchety (or witjuti) grub, a caterpillar of a wood moth (Lepidoptera: Cossidae) that feeds on the roots and stems of witjuti bushes (certain Acacia species).
• 1.8.2 Insects as feed for domesticated animals
• Further Reading
• Chapter 2 External Anatomy
• 2.1 The cuticle
• Fig. 2.1 The general structure of insect cuticle; the enlargement above shows details of the epicuticle.
• Fig. 2.2 Structure of part of a chitin chain, showing two linked units of N-acetyl-D-glucosamine.
• Fig. 2.3 The ultrastructure of cuticle (from a transmission electron micrograph). (a) The arrangement of chitin microfibrils in a helicoidal array produces characteristic (though artefactual) parabolic patterns. (b) Diagram of how the rotation of microfibrils produces a lamellar effect owing to microfibrils being either aligned or non-aligned to the plane of sectioning.
• Fig. 2.4 A specialized worker, or replete, of the honeypot ant, Camponotus inflatus (Hymenoptera: Formicidae), which holds honey in its distensible abdomen and acts as a food store for the colony. The arthrodial membrane between tergal plates is depicted to the right in its unfolded and folded conditions.
• Fig. 2.5 The cuticular pores and ducts on the venter of an adult female of the citrus mealybug, Planococcus citri (Hemiptera: Pseudococcidae). Enlargements depict the ultrastructure of the wax glands and the various wax secretions (arrowed) associated with three types of cuticular structure: (a) a trilocular pore; (b) a tubular duct; and (c) a multilocular pore. Curled filaments of wax from the trilocular pores form a protective body-covering and prevent contamination with their own sugary excreta, or honeydew; long, hollow, and shorter curled filaments from the tubular ducts and multilocular pores, respectively, form the ovisac.
• Fig. 2.6 The four basic types of cuticular protuberances: (a) a multicellular spine; (b) a seta, or trichoid sensillum; (c) acanthae; and (d) microtrichia.
• 2.1.1 Colour production
• 2.2 Segmentation and tagmosis
• Fig. 2.7 Types of body segmentation. (a) Primary segmentation, as seen in soft-bodied larvae of some insects. (b) Simple secondary segmentation. (c) More-derived secondary segmentation. (d) Longitudinal section of dorsum of the thorax of winged insects, in which the acrotergites of the second and third segments have enlarged to become the postnota.
• Fig. 2.8 The major body axes and the relationship of parts of the appendages to the body, shown for a sepsid fly.
• 2.3 The head
• Fig. 2.9 Lateral view of the head of a generalized pterygote insect.
• 2.3.1 Mouthparts
• Fig. 2.10 Frontal view of the head and dissected mouthparts of an adult European earwig, Forficula auricularia (Dermaptera: Forficulidae). Note that the head is prognathous and thus a gular plate, or gula, occurs in the ventral neck region.
• Fig. 2.11 Frontal view of the head of a worker honey bee, Apis mellifera (Hymenoptera: Apidae), with transverse section of proboscis showing how the “tongue” (fused labial glossae) is enclosed within the sucking tube formed from the maxillary galae and labial palps.
• Fig. 2.12 Mouthparts of a white butterfly, Pieris sp. (Lepidoptera: Pieridae). (a) Positions of the proboscis showing, from left to right, at rest, with proximal region uncoiling, with distal region uncoiling, and fully extended with tip in two of many possible different positions due to flexing at the “knee bend”. (b) Lateral view of proboscis musculature. (c) Transverse section of the proboscis in the proximal region.
• Fig. 2.13 Mouthparts of female mosquito in: (a) frontal view; (b) transverse section.
• Fig. 2.14 Mouthparts of adult Diptera. (a) House fly, Musca (Muscidae). (b) Stable fly, Stomoxys (Muscidae).
• Fig. 2.15 Head and mouthparts of a thrips, Thrips australis (Thysanoptera: Thripidae). (a) Dorsal view of head showing mouthparts through prothorax. (b) Transverse section through proboscis. The plane of the transverse section is indicated by the dashed line in (a).
• Fig. 2.16 Head and mouthparts of a sucking louse, Pediculus (Psocodea: Anoplura: Pediculidae). (a) Longitudinal section of head (nervous system omitted). (b) Transverse section through eversible proboscis. The plane of the transverse section is indicated by the dashed line in (a).
• Fig. 2.17 Head and mouthparts of a human flea, Pulex irritans (Siphonaptera: Pulicidae). (a) Lateral view of head. (b) Transverse section through mouthparts. The plane of the transverse section is indicated by the dashed line in (a).
• 2.3.2 Cephalic sensory structures
• Fig. 2.18 The mouthparts and feeding currents of a mosquito larva of Anopheles quadrimaculatus (Diptera: Culicidae). (a) The larva floating just below the water surface, with head rotated through 180° relative to its body (which is dorsum-up so that the spiracular plate near the abdominal apex is in direct contact with the air). (b) Viewed from above, showing the venter of the head and the feeding current generated by setal brushes on the labrum (direction of water movement and paths taken by surface particles are indicated by arrows and dotted lines, respectively). (c) Lateral view showing the particle-rich water being drawn into the preoral cavity between the mandibles and maxillae, and its downward expulsion as the outward current.
• Fig. 2.19 Some types of insect antennae: (a) filiform—linear and slender; (b) moniliform—like a string of beads; (c) clavate or capitate—distinctly clubbed; (d) serrate—saw-like; (e) pectinate—comb-like; (f) flabellate—fan-shaped; (g) geniculate—elbowed; (h) plumose—bearing whorls of setae; and (i) aristate—with enlarged third segment bearing a bristle.
• 2.4 The thorax
• Fig. 2.20 Diagrammatic lateral view of a wing-bearing thoracic segment, showing the typical sclerites and their subdivisions.
• 2.4.1 Legs
• Fig. 2.21 The hind leg of a cockroach, Periplaneta americana (Blattodea: Blattidae), with enlargement showing ventral surface of pretarsus and last tarsomere.
• 2.4.2 Wings
• Fig. 2.22 Nomenclature for the main areas, folds and margins of a generalized insect wing.
• Fig. 2.23 A generalized wing of a neopteran insect (any living, winged insect, other than Ephemeroptera and Odonata), showing the articulation and the Kukalová-Peck nomenclatural scheme of wing venation. Notation is as follows: AA, anal anterior; AP, anal posterior; Ax, axillary sclerite; C, costa; CA, costa anterior; CP, costa posterior; CuA, cubitus anterior; CuP, cubitus posterior; hm, humeral vein; JA, jugal anterior; MA, media anterior; m-cu, cross-vein between medial and cubital areas; MP, media posterior; PC, precosta; R, radius; RA, radius anterior; r-m, cross-vein between radial and median areas; RP, radius posterior; ScA, subcosta anterior; ScP, subcosta posterior. Branches of the anterior and posterior sector of each vein are numbered, e.g. CuA1−4.
• Fig. 2.24 The left wings of a range of insects showing some of the major wing modifications: (a) fore wing of a butterfly of Danaus (Lepidoptera: Nymphalidae); (b) fore wing of a dragonfly of Urothemis (Odonata: Anisoptera: Libellulidae); (c) fore wing or tegmen of a cockroach of Periplaneta (Blattodea: Blattidae); (d) fore wing or elytron of a beetle of Anomala (Coleoptera: Scarabaeidae); (e) fore wing or hemelytron of a mirid bug (Hemiptera: Heteroptera: Miridae), showing three wing areas—the membrane, corium and clavus; (f) fore wing and haltere of a fly of Bibio (Diptera: Bibionidae). Nomenclatural scheme of venation consistent with that depicted in Fig. 2.23; that of (b) after J.W.H. Trueman, unpublished.
• 2.5 The abdomen
• 2.5.1 Terminalia
• Fig. 2.25 The female abdomen and ovipositor. (a) Lateral view of the abdomen of an adult tussock moth (Lepidoptera: Lymantriidae), showing the substitutional ovipositor formed from the extensible terminal segments. (b) Lateral view of a generalized orthopteroid ovipositor composed of appendages of segments 8 and 9. (c) Transverse section through the ovipositor of a katydid (Orthoptera: Tettigoniidae). T1 − T10, terga of first to tenth segments; S2 − S8, sterna of second to eighth segments.
• Fig. 2.26 (left) Male external genitalia. (a) Abdominal segment 9 of the bristletail Machilis variabilis (Archaeognatha: Machilidae). (b) Aedeagus of a click beetle (Coleoptera: Elateridae).
• Further Reading
• Chapter 3 Internal Anatomy and Physiology
• 3.1 Muscles and locomotion
• Fig. 3.1 Dissections of: (a) a female American cockroach, Periplaneta americana (Blattodea: Blattidae): and (b) a male black field cricket, Teleogryllus commodus (Orthoptera: Gryllidae). The fat body and most of the tracheae have been removed; most details of the nervous system are not shown.
• 3.1.1 Muscles
• 3.1.2 Muscle attachments
• Fig. 3.2 Muscle attachments to body wall: (a) tonofibrillae traversing the epidermis from the muscle to the cuticle (b) a muscle attachment in an adult beetle of Chrysobothrus femorata (Coleoptera: Buprestidae) (c) a multicellular apodeme with a muscle attached to one of its thread-like, cuticular “tendons” or apophyses.
• 3.1.3 Crawling, wriggling, swimming and walking
• Fig. 3.3 A ground beetle (Coleoptera: Carabidae: Carabus) walking in the direction of the dashed line. The three blackened legs are those in contact with the ground in the two positions illustrated—(a) is followed by (b).
• 3.1.4 Flight
• Fig. 3.4 Direct flight mechanisms: thorax during (a) upstroke and (b) downstroke of the wings. Indirect flight mechanisms: thorax during (c) upstroke and (d) downstroke of the wings. Stippled muscles are those contracting in each illustration.
• 3.2 The nervous system and co-ordination
• Fig. 3.5 Diagram of a simple reflex mechanism of an insect. The arrows show the paths of nerve impulses along nerve fibres (axons and dendrites). The ganglion, with its outer cortex and inner neuropile, is shown on the right.
• Fig. 3.6 The central nervous system of various insects showing the diversity of arrangement of ganglia in the ventral nerve cord. Varying degrees of fusion of ganglia occur from the least to the most specialized: (a) three separate thoracic and eight abdominal ganglia, as in Dictyopterus (Coleoptera: Lycidae) and Pulex (Siphonaptera: Pulicidae); (b) three thoracic and six abdominal ganglia, as in Blatta (Blattodea: Blattidae) and Chironomus (Diptera: Chironomidae); (c) two thoracic and considerable abdominal fusion of ganglia, as in Crabro and Eucera (Hymenoptera: Crabronidae and Anthophoridae); (d) highly fused, with one thoracic and no abdominal ganglia, as in Musca, Calliphora and Lucilia (Diptera: Muscidae and Calliphoridae); (e) extreme fusion, with no separate suboesophageal ganglion, as in Hydrometra (Hemiptera: Hydrometridae) and Rhizotrogus (Scarabaeidae).
• Fig. 3.7 Mediolongitudinal section of an immature cockroach of Periplaneta americana (Blattodea: Blattidae) showing internal organs and tissues.
• 3.3 The endocrine system and the function of hormones
• 3.3.1 Endocrine centres
• Neurosecretory cells
• Fig. 3.8 The main endocrine centres in a generalized insect.
• Box 3.1 Molecular genetic techniques and their application to neuropeptide research
• Corpora cardiaca
• Prothoracic glands
• Corpora allata
• Inka cells
• 3.3.2 Hormones
• 3.4 The circulatory system
• 3.4.1 Haemolymph
• Table 3.1 Examples of some important insect physiological processes mediated by neuropeptides; note that usually only one, of often multiple, function of each neuropeptide is listed.
• 3.4.2 Circulation
• Fig. 3.9 Schematic diagram of a well-developed circulatory system: (a) longitudinal section through body; (b) transverse section of the abdomen; (c) transverse section of the thorax. Arrows indicate directions of haemolymph flow.
• 3.4.3 Protection and defence by the haemolymph
• 3.5 The tracheal system and gas exchange
• Fig. 3.10 Schematic diagram of a generalized tracheal system seen in a transverse section of the body at the level of a pair of abdominal spiracles. Enlargements show: (a) an atriate spiracle with closing valve at inner end of atrium; (b) tracheoles running to a muscle fibre.
• Fig. 3.11 Some basic variations in the open (a–c) and closed (d–f) tracheal systems of insects. (a) Simple tracheae with valved spiracles, as in cockroaches. (b) Tracheae with mechanically ventilated air sacs, as in honey bees. (c) Metapneustic system with only the terminal spiracles functional, as in mosquito larvae. (d) Entirely closed tracheal system with cutaneous gas exchange, as in most endoparasitic larvae. (e) Closed tracheal system with abdominal tracheal gills, as in mayfly nymphs. (f) Closed tracheal system with rectal tracheal gills, as in dragonfly nymphs.
• Box 3.2 Tracheal hypertrophy in mealworms at low oxygen concentrations
• 3.5.1 Diffusion and ventilation
• 3.6 The gut, digestion and nutrition
• Fig. 3.12 The four major categories of insect-feeding specialization. Many insects are typical of one category, but others cross two categories (or more, as in generalist cockroaches).
• Box 3.3 The filter chamber of Hemiptera
• 3.6.1 Structure of the gut
• Fig. 3.13 Generalized insect alimentary canal showing division into three regions. The cuticular lining of the foregut and hindgut are indicated by thicker black lines.
• Fig. 3.14 Preoral and anterior foregut morphology in: (a) a generalized orthopteroid insect; and (b) a xylem-feeding cicada. Musculature of the mouthparts and the (a) pharyngeal or (b) cibarial pump are indicated but not fully labelled. Contraction of the respective dilator muscles causes dilation of the pharynx or cibarium and fluid is drawn into the pump chamber. Relaxation of these muscles results in elastic return of the pharynx or cibarial walls and the expulsion of food upwards into the oesophagus.
• 3.6.2 Saliva and food ingestion
• Fig. 3.15 Longitudinal section through the anterior body of a caterpillar of the small white, small cabbage white or cabbage white butterfly, Pieris rapae (Lepidoptera: Pieridae). Note the thickened epidermal layer lining the midgut.
• 3.6.3 Digestion of food
• Fig. 3.16 Generalized scheme of the endo–ectoperitrophic circulation of digestive enzymes in the midgut.
• 3.6.4 The fat body
• 3.6.5 Nutrition and microorganisms
• 3.7 The excretory system and waste disposal
• 3.7.1 The Malpighian tubules and rectum
• Fig. 3.17 Schematic diagram of a generalized excretory system showing the path of elimination of wastes.
• Box 3.4 Cryptonephric systems*
• Fig. 3.18 Schematic diagram of the organs in the excretory system of the desert locust, Schistocerca gregaria (Orthoptera: Acrididae). Only a few of the >100 Malpighian tubules are drawn. (a) Transverse section of one Malpighian tubule showing probable transport of ions, water and other substances between the surrounding haemolymph and the tubule lumen; active processes are indicated by solid arrows and passive processes by dashed arrows. (b) Diagram illustrating the movements of solutes and water in the rectal pad cells during fluid resorption from the rectal lumen. Pathways of water movement are represented by open arrows, and solute movements by black arrows. Ions are actively transported from the rectal lumen (compartment 1) to the adjacent cell cytoplasm (compartment 2) and then to the intercellular spaces (compartment 3). Mitochondria are positioned so as to provide the energy for this active ion transport. Fluid in the spaces is hyperosmotic (higher ion concentration) to the rectal lumen and draws water by osmosis from the lumen via the septate junctions between the cells. Water thus moves from compartment 1 to 3 to 4 and finally to 5, the haemolymph in the haemocoel.
• 3.7.2 Nitrogen excretion
• Fig. 3.19 Molecules of the three common nitrogenous excretory products. The high N : H ratio of uric acid relative to both ammonia and urea means that less water is used for uric acid synthesis (as hydrogen atoms are derived ultimately from water).
• 3.8 Reproductive organs
• Fig. 3.20 Comparison of generalized: (a) female reproductive system; and (b) male reproductive system.
• Table 3.2 The corresponding female and male reproductive organs of insects
• 3.8.1 The female reproductive system
• 3.8.2 The male reproductive system
• Further Reading
• Chapter 4 Sensory Systems and Behaviour
• 4.1 Mechanical stimuli
• 4.1.1 Tactile mechanoreception
• Fig. 4.1 Longitudinal section of a trichoid sensillum showing the arrangement of the three associated cells.
• 4.1.2 Position mechanoreception (proprioceptors)
• 4.1.3 Sound reception
• Fig. 4.2 Proprioceptors: (a) sensilla of a hair plate located at a joint, showing how the hairs are stimulated by contacting adjacent cuticle; (b) campaniform sensillum on the haltere of a fly.
• Non-tympanal vibration reception
• Fig. 4.3 Longitudinal section of a scolopidium, the basic unit of a chordotonal organ.
• Tympanal reception
• Fig. 4.4 Tympanal organs of a katydid, Decticus (Orthoptera: Tettigoniidae): (a) transverse section through the fore legs and prothorax to show the acoustic spiracles and tracheae; (b) transverse section through the base of the fore tibia; (c) longitudinal breakaway view of the fore tibia.
• Box 4.1 Aural location of host by a parasitoid fly
• 4.1.4 Sound production
• Fig. 4.5 The singing burrow of a mole cricket, Scapteriscus acletus (Orthoptera: Gryllotalpidae), in which the singing male sits with his head in the bulb and tegmina raised across the throat of the horn. The depth markers in the burrow are in centimetres, with the burrow being just over 4 cm deep.
• 4.2 Thermal stimuli
• 4.2.1 Thermoreception
• 4.2.2 Thermoregulation
• Behavioural thermoregulation (ectothermy)
• Physiological thermoregulation (endothermy)
• 4.3 Chemical stimuli
• 4.3.1 Chemoreception
• Fig. 4.6 The antennae of a male moth of Trictena atripalpis (Lepidoptera: Hepialidae): (a) anterior view of head showing tripectinate antennae of this species; (b) cross-section through an antenna showing the three branches; (c) enlargement of tip of the outer branch of one pectination showing olfactory sensilla.
• Box 4.2 Reception of communication molecules
• Box 4.3 The electroantennogram
• 4.3.2 Semiochemicals: pheromones
• Sex pheromones
• Fig. 4.7 Location of pheromone-emitting female by male moth tacking upwind. The pheromone trail forms a somewhat discontinuous plume because of turbulence, intermittent release, and other factors.
• Fig. 4.8 A pair of queen butterflies, Danaus gilippus (Lepidoptera: Nymphalidae: Danainae), showing aerial “hairpencilling” by the male. The male (above) has splayed hairpencils (at his abdominal apex) and is applying pheromone to the female (below).
• Aggregation pheromones
• Spacing pheromones
• Trail-marking pheromones
• Alarm pheromones
• 4.3.3 Semiochemicals: kairomones, allomones and synomones
• Kairomones
• Allomones
• Synomones
• 4.3.4 Carbon dioxide as a sensory cue
• 4.4 Insect vision
• 4.4.1 Dermal detection
• 4.4.2 Stemmata
• Box 4.4 Biological clocks
• 4.4.3 Ocelli
• 4.4.4 Compound eyes
• Fig. 4.9 Longitudinal sections through simple eyes: (a) a simple stemma of a lepidopteran larva; (b) a light-adapted median ocellus of a locust.
• Fig. 4.10 Details of the compound eye: (a) a cutaway view showing the arrangement of the ommatidia and the facets; (b) a single ommatidium with an enlargement of a transverse section.
• 4.4.5 Light production
• Fig. 4.11 The flash patterns of males of nine of Photinus firefly species (Coleoptera: Lampyridae), each of which generates a distinctive pattern of signals in order to elicit a response from their conspecific females.
• 4.5 Insect behaviour
• Further Reading
• Chapter 5 Reproduction
• 5.1 Bringing the sexes together
• Fig. 5.1 Males of the Arctic fly, Rhamphomyia nigrita (Diptera: Empididae), hunt for prey in swarms of Aedes mosquitoes (lower mid-right of drawing) and carry the prey to a specific visual marker of the swarm site (left-hand side of drawing). Swarms of both the empidids and the mosquitoes form near conspicuous landmarks, including refuse heaps or oil drums, which are common in parts of the tundra. Within the mating swarm (upper left), a male empidid rises towards a female hovering above, they pair, and the prey is transferred to the female; the mating pair alights (lower far right) and the female feeds as they copulate. Females appear to obtain food only via males and, as individual prey items are small, must mate repeatedly in order to obtain sufficient nutrients to develop a batch of eggs.
• 5.2 Courtship
• 5.3 Sexual selection
• Box 5.1 Courtship and mating in Mecoptera
• Fig. 5.2 Relationship between length of horn and body size (thorax width) of male scarabs of Onthophagus taurus.
• Fig. 5.3 Two males of Phytalmia mouldsi (Diptera: Tephritidae) fighting over access to the oviposition site at the larval substrate visited by females. These tropical rainforest flies thus have a resource-defence mating system.
• 5.4 Copulation
• Fig. 5.4 Posterior ends of a pair of copulating milkweed bugs, Oncopeltus fasciatus (Hemiptera: Lygaeidae). Mating commences with the pair facing in the same direction, then the male rotates his eighth abdominal segment (90°) and genital capsule (180°), erects the aedeagus and gains entry to the female’s genital chamber, before he swings around to face in the opposite direction. The bugs may copulate for several hours, during which they walk around with the female leading and the male walking backwards. (a) Lateral view of the terminal segments, showing the valves of the female’s ovipositor in the male genital chamber; (b) longitudinal section showing internal structures of the reproductive system, with the tip of the male’s aedeagus in the female’s spermatheca.
• Box 5.2 Mating in katydids and crickets
• 5.4.1 Nuptial feeding and other “gifts”
• Box 5.3 Cannibalistic mating in mantids
• Box 5.4 Puddling and gifts in Lepidoptera
• 5.5 Diversity in genitalic morphology
• Fig. 5.5 Species-specificity in part of the male genitalia of three sibling species of Drosophila (Diptera: Drosophilidae). The epandrial processes of tergite 9 in: (a) D. mauritiana; (b) D. simulans; (c) D. melanogaster.
• Fig. 5.6 Spermatophores lying within the bursae of the female reproductive tracts of moth species from four different genera (Lepidoptera: Noctuidae). The sperm leave via the narrow end of each spermatophore, which has been deposited so that its opening lies opposite the “seminal duct” leading to the spermatheca (not drawn). The bursa on the far right contains two spermatophores, indicating that the female has re-mated.
• Fig. 5.7 Males of three species of the water-strider genus Rheumatobates, showing species-specific antennal and leg modifications (mostly flexible setae). These non-genitalic male structures are specialized for contact with the female during mating, when the male rides on her back. Females of all species have a similar body form. (a) R. trulliger; (b) R. rileyi; (c) R. bergrothi.
• 5.6 Sperm storage, fertilization and sex determination
• 5.7 Sperm competition
• Box 5.5 Sperm precedence
• Box 5.6 Control of mating and oviposition in a blow fly
• Fig. 5.8 A copulating pair of stink or shield bugs of the genus Poecilometis (Hemiptera: Pentatomidae). Many heteropteran bugs engage in prolonged copulation, which prevents other males from inseminating the female until either she becomes non-receptive to further males or she lays the eggs fertilized by the “guarding” male.
• 5.8 Oviparity (egg-laying)
• Fig. 5.9 Oviposition by a South African ladybird beetle, Cheilomenes lunata (Coleoptera: Coccinellidae). The eggs adhere to the leaf surface because of a sticky secretion applied to each egg.
• Fig. 5.10 The generalized structure of a libelluloid dragonfly egg (Odonata: Corduliidae, Libellulidae). Libelluloid dragonflies oviposit into freshwater but always exophytically (i.e. outside of plant tissues). The endochorionic and exochorionic layers of the eggshell are separated by a distinct gap in some species. A gelatinous matrix may be present on the exochorion or as connecting strands between eggs.
• Box 5.7 Does mother know best?
• Box 5.8 Egg-tending fathers—the giant water bugs
• Fig. 5.11 A female of the parasitic wasp Megarhyssa nortoni (Hymenoptera: Ichneumonidae) probing a pine log with her very long ovipositor in search of a larva of the sirex wood wasp, Sirex noctilio (Hymenoptera: Siricidae). If a larva is located, she stings and paralyzes it before laying an egg on it.
• Fig. 5.12 Tip of the ovipositor of a female of the black field cricket, Teleogryllus commodus (Orthoptera: Gryllidae), split open to reveal the inside surface of the two halves of the ovipositor. Enlargements show: (a) posteriorly directed ovipositor scales; (b) distal group of sensilla.
• 5.9 Ovoviviparity and viviparity
• 5.10 Other modes of reproduction
• 5.10.1 Parthenogenesis, paedogenesis (pedogenesis) and neoteny
• 5.10.2 Hermaphroditism
• 5.10.3 Polyembryony
• 5.10.4 Reproductive effects of endosymbionts
• 5.11 Physiological control of reproduction
• 5.11.1 Vitellogenesis and its regulation
• Fig. 5.13 A schematic diagram of the hormonal regulation of reproductive events in insects. The transition from ecdysteroid production by the pre-adult prothoracic gland to the adult ovary varies between taxa.
• Further Reading
• Chapter 6 Insect Development and Life Histories
• 6.1 Growth
• Fig. 6.1 Schematic drawing of the life cycle of a non-biting midge (Diptera: Chironomidae, Chironomus), showing the various events and stages of insect development.
• 6.2 Life-history patterns and phases
• Fig. 6.2 The life cycle of a hemimetabolous insect, the southern green stink bug or green vegetable bug, Nezara viridula (Hemiptera: Pentatomidae), showing the eggs, nymphs of the five instars, and the adult bug on a tomato plant. This cosmopolitan and polyphagous bug is an important world pest of food and fibre crops.
• Fig. 6.3 The life cycle of a holometabolous insect, a bark beetle, Ips grandicollis (Coleoptera: Curculionidae: Scolytinae), showing the egg, the three larval instars, the pupa, and the adult beetle.
• 6.2.1 Embryonic phase
• Fig. 6.4 Stages in the development of the wings of the small white, small cabbage white, or cabbage white butterfly, Pieris rapae (Lepidoptera: Pieridae). A wing imaginal disc in: (a) a first-instar larva; (b) a second-instar larva; (c) a third-instar larva; and (d) a fourth-instar larva. The wing bud, as it appears: (e) if dissected out of the wing pocket; or (f) cut in cross-section in a fifth-instar larva.
• Fig. 6.5 Embryonic development of the scorpionfly Panorpodes paradoxa (Mecoptera: Panorpodidae): (a–c) schematic drawings of egg halves from which yolk has been removed to show position of embryo; (d–j) gross morphology of developing embryos at various ages. Age from oviposition: (a) 32 h; (b) 2 days; (c) 7 days; (d) 12 days; (e) 16 days; (f) 19 days; (g) 23 days; (h) 25 days; (i) 25–26 days; (j) full grown at 32 days.
• Box 6.1 Molecular insights into insect development
• 6.2.2 Larval or nymphal phase
• Fig. 6.6 Examples of larval types. Polypod larvae: (a) Lepidoptera: Sphingidae; (b) Lepidoptera: Geometridae; (c) Hymenoptera: Diprionidae. Oligopod larvae: (d) Neuroptera: Osmylidae; (e) Coleoptera: Carabidae; (f) Coleoptera: Scarabaeidae. Apod larvae: (g) Coleoptera: Scolytinae; (h) Diptera: Calliphoridae; (i) Hymenoptera: Vespidae.
• 6.2.3 Metamorphosis
• Fig. 6.7 Examples of pupal types. Exarate decticous pupae: (a) Megaloptera: Sialidae; (b) Mecoptera: Bittacidae. Exarate adecticous pupae: (c) Coleoptera: Dermestidae; (d) Hymenoptera: Vespidae; (e,f) Diptera: Calliphoridae, puparium and pupa within. Obtect adecticous pupae: (g) Lepidoptera: Cossidae; (h) Lepidoptera: Saturniidae; (i) Lepidoptera: Papilionidae, chrysalis; (j) Coleoptera: Coccinellidae.
• 6.2.4 Imaginal or adult phase
• Fig. 6.8 The nymphal–imaginal moult of a male dragonfly of Aeshna cyanea (Odonata: Aeshnidae). The final-instar nymph climbs out of the water prior to the shedding of its cuticle. The old cuticle splits mid-dorsally, the teneral adult frees itself, swallows air and must wait many hours for its wings to expand and dry.
• 6.3 Process and control of moulting
• Fig. 6.9 Schematic diagram of the classical view of endocrine control of the epidermal processes that occur in moulting and metamorphosis in a holometabolous insect. This scheme simplifies the complexity of ecdysteroid and JH secretion, and does not indicate the influence of neuropeptides such as eclosion hormone. JH, juvenile hormone; PTTH, prothoracicotropic hormone.
• Fig. 6.10 Diagrammatic view of the changing activities of the epidermis during the fourth and fifth larval instars and prepupal (= pharate pupal) development in the tobacco hornworm, Manduca sexta (Lepidoptera: Sphingidae), in relation to the hormonal environment. The dots in the epidermal cells represent granules of the blue pigment insecticyanin. ETH, ecdysis triggering hormone; EH, eclosion hormone; JH, juvenile hormone; EPI, EXO, ENDO, deposition of pupal epicuticle, exocuticle and endocuticle, respectively. The numbers on the x-axis represent days.
• 6.4 Voltinism
• Fig. 6.11 A flow chart of the events prior to metamorphosis that determine body size in the tobacco hornworm, Manduca sexta (Lepidoptera: Sphingidae). During the last larval instar, there are three physiological decision points. The final size of the insect is determined by the amount of growth that occurs in the intervals between these three conditional events. JH, juvenile hormone; PTTH, prothoracicotropic hormone.
• 6.5 Diapause
• 6.6 Dealing with environmental extremes
• 6.6.1 Cold
• Freeze tolerance
• Freeze avoidance
• Chill tolerance
• Chill susceptibility
• Opportunistic survival
• 6.6.2 Heat
• 6.6.3 Aridity
• 6.7 Migration
• 6.8 Polymorphism and polyphenism
• 6.8.1 Genetic polymorphism
• 6.8.2 Environmental polymorphism, or polyphenism
• 6.9 Age-grading
• 6.9.1 Age-grading of immature insects
• Fig. 6.12 Growth and development in a marine midge, Telmatogeton (Diptera: Chironomidae), showing increases in: (a) head capsule length; (b) mandible length; and (c) body length, between the four larval instars (I–IV). The dots and horizontal lines above each histogram represent the means and standard deviations of measurements for each instar. Note that the lengths of the sclerotized head and mandible fall into discrete size classes representing each instar, whereas body length is an unreliable indicator of instar number, especially for separating the third- and fourth-instar larvae.
• 6.9.2 Age-grading of adult insects
• 6.10 Environmental effects on development
• 6.10.1 Temperature
• Fig. 6.13 Age-specific oviposition rates of three predators of cotton pests, Chrysopa sp. (Neuroptera: Chrysopidae), Micromus tasmaniae (Neuroptera: Hemerobiidae) and Nabis kinbergii (Hemiptera: Nabidae), based on physiological time above respective development thresholds of 10.5°C, −2.9°C and 11.3°C. D°, day-degrees.
• Box 6.2 Calculation of day-degrees
• 6.10.2 Photoperiod
• 6.10.3 Humidity
• 6.10.4 Mutagens and toxins
• 6.10.5 Biotic effects
• Fig. 6.14 Solitary and gregarious females of the migratory locust, Locusta migratoria (Orthoptera: Acrididae). The solitaria adults have a pronounced pronotal crest and the femora are larger relative to the body and wing than in the gregaria adults. Intermediate morphologies occur in the transiens (transient stage), during the transformation from solitaria to gregaria or the reverse.
• Further Reading
• Chapter 7 Insect Systematics: Phylogeny and Classification
• 7.1 Systematics
• 7.1.1 Phylogenetic methods
• Fig. 7.1 A cladogram showing the relationships of four species, A, B, C and D, and examples of: (a) the three monophyletic groups; (b) two of the four possible (ABC, ABD, ACD, BCD) paraphyletic groups; and (c) one of the four possible (AC, AD, BC and BD) polyphyletic groups that could be recognized based on this cladogram.
• Molecular phylogenetics
• 7.1.2 Taxonomy and classification
• How entomologists recognize insect species
• Box 7.1 Gonipterus weevils—recognition of a species complex
• Box 7.2 Integrative taxonomy of woodroaches
• Box 7.3 DNA barcoding and species discovery
• 7.2 The extant hexapoda
• 7.3 Informal group Entognatha: collembola (springtails), diplura (diplurans) and protura (proturans)
• 7.3.1 Order Collembola (springtails) (see also Taxobox 1)
• 7.3.2 Order Diplura (diplurans) (see also Taxobox 1)
• 7.3.3 Order Protura (proturans) (see also Taxobox 1)
• 7.4 Class insecta (true insects)
• 7.4.1 Apterygote Insecta (former Thysanura sensu lato)
• Order Archaeognatha (Microcoryphia; archaeognathans or bristletails) (see also Taxobox 2)
• Fig. 7.2 Cladogram of postulated relationships of extant hexapods, based on combined morphological and nucleotide sequence data. Dashed lines indicate uncertain relationships or alternative hypotheses. Thysanura sensu lato refers to Thysanura in the broad sense. An expanded concept is depicted for each of two orders—Blattodea (including termites) and Psocodea (former Psocoptera and Phthiraptera)—but intraordinal relationships are shown simplified (see Fig. 7.4 and Fig. 7.5 for full details).
• Order Zygentoma (silverfish) (see also Taxobox 3)
• 7.4.2 Pterygota
• Division Palaeoptera
• Fig. 7.3 The three possible relationships of Ephemeroptera, Odonata and Neoptera. The top tree shows a monophyletic Palaeoptera, as indicated by the vertical line on the top right.
• Order Ephemeroptera (mayflies) (see also Taxobox 4)
• Order Odonata (damselflies and dragonflies) (see also Taxobox 5)
• Division Neoptera
• Subdivision Polyneoptera (or Orthopteroid–Plecopteroid assemblage)
• Order Plecoptera (stoneflies) (see also Taxobox 6)
• Fig. 7.4 Cladogram of postulated relationships within Dictyoptera, based on combined morphological and nucleotide sequence data. The revised concept of order Blattodea includes the termites, which are given the rank of epifamily (-oidae) as Termitoidae; similarly the woodroaches (Cryptocercoidae: Cryptocercidae).
• Order Dermaptera (earwigs) (see also Taxobox 7)
• Order Zoraptera (zorapterans or angel insects) (see also Taxobox 8)
• Order Orthoptera (grasshoppers, locusts, katydids and crickets) (see also Taxobox 9)
• Order Embioptera (= Embiidina, Embiodea) (embiopterans or webspinners) (see also Taxobox 10)
• Order Phasmatodea (phasmids, stick-insects or walking sticks) (see also Taxobox 11)
• Order Grylloblattodea (= Grylloblattaria, Notoptera) (grylloblattids, ice crawlers or rock crawlers) (see also Taxobox 12)
• Order Mantophasmatodea (heelwalkers) (see also Taxobox 13)
• Order Mantodea (mantids, mantises or praying mantids) (see also Taxobox 14)
• Order Blattodea (cockroaches and termites) (see also Taxobox 15 and Taxobox 16)
• Epifamily Termitoidae (former order Isoptera; termites, “white ants”) (see also Taxobox 16)
• Subdivision Paraneoptera (Acercaria, or Hemipteroid assemblage)
• Order Psocodea (bark lice, book lice, chewing lice and sucking lice) (see also Taxobox 17 and Taxobox 18)
• Fig. 7.5 Cladogram of postulated relationships among suborders of Psocodea, with Condylognatha as the sister group. The depicted hypothesis of relationships suggests two origins of parasitism in the order.
• Order Thysanoptera (thrips) (see also Taxobox 19)
• Order Hemiptera (bugs, moss bugs, cicadas, leafhoppers, planthoppers, spittle bugs, treehoppers, aphids, jumping plant lice, scale insects and whiteflies) (see also Box 10.2 and Taxobox 20)
• Fig. 7.6 Cladogram of postulated relationships within Hemiptera, based on combined morphological and nucleotide sequence data. The dashed line and italicized name indicate the paraphyly of Homoptera.
• Subdivision Holometabola (= Endopterygota)
• Fig. 7.7 Two competing hypotheses for the relationships of Antliophora. (a) Based on nucleotide sequence data, including ribosomal genes, supported by morphology. (b) Based on nucleotide sequence data from single-copy protein-coding genes.
• Orders Neuroptera (lacewings, owlflies, antlions), and Megaloptera (alderflies, dobsonflies, fishflies) and Raphidioptera (snakeflies) (see also Box 10.4 and Taxobox 21)
• Order Coleoptera (beetles) (see also Box 10.3 and Taxobox 22)
• Order Strepsiptera (strepsipterans) (see also Taxobox 23)
• Order Diptera (true flies) (see also Box 10.1 and Taxobox 24)
• Order Mecoptera (scorpionflies, hangingflies and snowfleas) (see also Taxobox 25)
• Order Siphonaptera (fleas) (see also Taxobox 26)
• Order Trichoptera (caddisflies) (see also Box 10.4 and Taxobox 27)
• Order Lepidoptera (butterflies and moths) (see also Taxobox 28)
• Order Hymenoptera (ants, bees, wasps, sawflies and wood wasps) (see also Taxobox 29)
• Fig. 7.8 Cladogram of postulated relationships of selected lepidopteran higher taxa, based on molecular data.
• Fig. 7.9 Simplified tree of postulated relationships among higher taxa of Hymenoptera based on molecular and morphological data. Three species-poor taxa (Cephoidea, Xiphydrioidea and Ceraphronoidea) with unstable positions have been omitted; Xyeloidea and Diaprioidea may be paraphyletic, as indicated by double lines in the tree. Sensu stricto (abbreviated s.s.) means in the restricted sense.
• Further Reading
• Chapter 8 Insect Evolution and Biogeography
• 8.1 Relationships of the hexapoda to other arthropoda
• Fig. 8.1 One possible evolutionary scenario for Pancrustacea (Crustacea + Hexapoda), in which hexapods evolved from an ancestor shared with Remipedia (today in coastal aquifers) and Cephalocarida (today in benthic intertidal). Note that none of the depicted relationships are fully supported by all available molecular and morphological data.
• 8.2 The antiquity of insects
• 8.2.1 The insect fossil record
• Box 8.1 The difficulties with dating
• Fig. 8.2 The geological history of insects in relation to plant evolution. Taxa that contain only fossils are indicated by the symbol †. The record for extant orders is based on definite members of the crown group and does not include stem-group fossils; dashed lines indicate uncertainty in placement in the crown group. Thus, this chart does not include records of most early insect radiations; for example “roachoid” fossils occur in the Palaeozoic but are not part of the more narrowly defined Dictyoptera and Blattodea. Protura and Siphonaptera are not shown due to inadequacy of their fossil record; Isoptera is part of Blattodea. The placement of Rhyniognatha is unknown. (Insect fossil records have been interpreted from primary sources and after Grimaldi & Engel 2005; the date for the start of each geological period is from the International Commission on Stratiography; the Tertiary often is divided into the Palaeogene (66–23 Ma) and the Neogene (23–2.6 Ma.)
• Box 8.2 There were giants—evolution of insect gigantism
• Fig. 8.3 Reconstruction of Stenodictya lobata (Palaeodictyoptera: Dictyoneuridae).
• 8.2.2 Living insect distributions as evidence for antiquity
• 8.3 Were the first insects aquatic or terrestrial?
• Fig. 8.4 Stylized tracheal system. (a) Oxygen uptake through invagination. (b) Invagination closed, with tracheal gas exchange through gill. 1, indicates point of invagination of the tracheal system; 2, indicates point for oxygen uptake; 3, indicates point for oxygen delivery, such as muscles.
• 8.4 Evolution of wings
• Fig. 8.5 Appendages of hypothetical primitive Palaeozoic (left of each diagram) and modern (right of each diagram) pterygotes (winged insects). (a) Thoracic segment of adult showing generalized condition of appendages. (b) Dorsal view of nymphal morphology.
• 8.5 Evolution of metamorphosis
• 8.6 Insect diversification
• 8.7 Insect biogeography
• 8.8 Insect evolution in the pacific
• Fig. 8.6 Area cladogram showing phylogenetic relationships of hypothetical insect taxa with taxon names replaced by their areas of endemism in the Hawai′ian archipelago. The pattern of colonization and speciation of the insects on the islands is depicted by arrows showing the sequence and direction of events: A, founding; B, diversification within an island; C, back-colonization event. myo, million years old. Dashed line denotes extinct lineage.
• Further Reading
• Chapter 9 Ground-Dwelling Insects
• 9.1 Insects of litter and soil
• Fig. 9.1 Diagrammatic view of a soil profile showing some typical litter and soil insects and other hexapods. Note that organisms living on the soil surface and in litter have longer legs than those found deeper in the ground. Organisms occurring deep in the soil usually are legless or have reduced legs; they are unpigmented and often blind. The organisms depicted are: (1) worker of a wood ant (Hymenoptera: Formicidae); (2) springtail (Collembola: Isotomidae); (3) ground beetle (Coleoptera: Carabidae); (4) rove beetle (Coleoptera: Staphylinidae) eating a springtail; (5) larva of a crane fly (Diptera: Tipulidae); (6) japygid dipluran (Diplura: Japygidae) attacking a smaller campodeid dipluran; (7) pupa of a ground beetle (Coleoptera: Carabidae); (8) bristletail (Archaeognatha: Machilidae); (9) female earwig (Dermaptera: Labiduridae) tending her eggs; (10) wireworm, larva of a tenebrionid beetle (Coleoptera: Tenebrionidae); (11) larva of a robber fly (Diptera: Asilidae); (12) larva of a soldier fly (Diptera: Stratiomyidae); (13) springtail (Collembola: Isotomidae); (14) larva of a weevil (Coleoptera: Curculionidae); (15) larva of a muscid fly (Diptera: Muscidae); (16) proturan (Protura: Sinentomidae); (17) springtail (Collembola: Isotomidae); (18) larva of a March fly (Diptera: Bibionidae); (19) larva of a scarab beetle (Coleoptera: Scarabaeidae). (Individual organisms after various sources, especially Eisenbeis & Wichard 1987.)
• Box 9.1 Soldier flies can recycle waste
• Fig. 9.2 Fossorial fore legs of: (a) a mole cricket of Gryllotalpa (Orthoptera: Gryllotalpidae); (b) a nymphal periodical cicada of Magicicada (Hemiptera: Cicadidae); and (c) a scarab beetle of Canthon (Coleoptera: Scarabaeidae). ((a) After Frost 1959; (b) after Snodgrass 1967; (c) after Richards & Davies 1977.)
• Box 9.2 Antimicrobial tactics to protect the brood of ground-nesting wasps
• 9.1.1 Root-feeding insects
• Box 9.3 Ecosystem engineering by southern African termites
• Box 9.4 Ground pearls
• 9.2 Insects and dead trees or decaying wood
• Fig. 9.3 A plume-shaped tunnel excavated by the bark beetle Scolytus unispinosus (Coleoptera: Curculionidae: Scolytinae), showing eggs at the ends of a number of galleries; enlargement shows an adult beetle.
• Fig. 9.4 Underside of the thorax of the beetle Henoticus serratus (Coleoptera: Cryptophagidae), showing the depressions, called mycangia, which the beetle uses to transport fungal material that inoculates new substrate on recently burnt wood.
• 9.3 Insects and dung
• Fig. 9.5 Interactions among the arthropod groups that are common in cattle dung: the larvae of dung-feeding flies (from several families) feed on microorganisms; early-instar larvae of mixed-diet flies feed on microorganisms, but in later instars become predatory; larvae of predatory flies feed only on other insects; adult dung-feeding beetles (mainly Scarabaeidae) probably feed mostly on microorganisms in the fluids of fresh dung, whereas their larvae breakdown the plant fibres in ingested dung with the help of symbiotic gut bacteria; predatory beetles (such as Staphylinidae) feed on other insects, especially immature stages of flies; fungus-feeding beetles colonize pats in their later stages of decomposition after fungi have developed; larvae of wasps are mostly parasitoids on dung-breeding flies.
• Fig. 9.6 A pair of dung beetles of Onthophagus gazella (Coleoptera: Scarabaeidae) filling in the tunnels that they have excavated below a dung pad. The inset shows an individual dung ball within which beetle development takes place: (a) egg; (b) larva, which feeds on the dung; (c) pupa; and (d) adult just prior to emergence.
• 9.4 Insect–carrion interactions
• 9.5 Insect–fungal interactions
• 9.5.1 Fungivorous insects
• 9.5.2 Fungus farming by leaf-cutter ants
• Fig. 9.7 The fungus gardens of the leaf-cutter ant, Atta cephalotes (Formicidae), require a constant supply of leaves. (a) A medium-sized worker, called a media, cuts a leaf with its serrated mandibles, while a minor worker guards the media from a parasitic phorid fly (Apocephalus) that lays its eggs on living ants. (b) A guarding minor hitchhikes on a leaf fragment carried by a media.
• 9.5.3 Fungus cultivation by termites
• 9.6 Cavernicolous insects
• 9.7 Environmental monitoring using ground-dwelling hexapods
• Further Reading
• Chapter 10 Aquatic Insects
• 10.1 Taxonomic distribution and terminology
• Box 10.1 Aquatic immature Diptera (true flies)
• Box 10.2 Aquatic Hemiptera (true bugs)
• Box 10.3 Aquatic Coleoptera (beetles)
• 10.2 The evolution of aquatic lifestyles
• Box 10.4 Aquatic Neuropterida
• Aquatic Neuroptera (lacewings and spongillaflies)
• Aquatic Megaloptera (alderflies, dobsonflies, fishflies)
• 10.3 Aquatic insects and their oxygen supplies
• 10.3.1 The physical properties of oxygen
• 10.3.2 Gaseous exchange in aquatic insects
• 10.3.3 Oxygen uptake with a closed tracheal system
• Fig. 10.1 A stonefly nymph (Plecoptera: Gripopterygidae) showing filamentous anal gills.
• 10.3.4 Oxygen uptake with an open spiracular system
• Fig. 10.2 The life cycle of the mosquito Culex pipiens (Diptera: Culicidae): (a) adult emerging from its pupal exuviae at the water surface; (b) adult female ovipositing, with her eggs adhering together as a floating raft; (c) larvae obtaining oxygen at the water surface via their siphons; (d) pupa suspended from the water meniscus, with its respiratory horn in contact with the atmosphere.
• Fig. 10.3 A male water beetle of Dytiscus (Coleoptera: Dytiscidae) replenishing its store of air at the water surface. Below is a transverse section of the beetle’s abdomen showing the large air store below the elytra and the tracheae opening into this air space. Note that the tarsi of the fore legs are dilated to form adhesive pads that are used to hold the female during copulation.
• 10.3.5 Behavioural ventilation
• Fig. 10.4 Dorsal (left) and ventral (right) views of the larva of Edwardsina polymorpha (Diptera: Blephariceridae); the venter has suckers, which the larva uses to adhere to rock surfaces in fast-flowing water.
• 10.4 The aquatic environment
• 10.4.1 Lotic adaptations
• Box 10.5 Aquatic–terrestrial insect fluxes
• Fig. 10.5 Portable larval cases of representative families of caddisflies (Trichoptera): (a) Helicopsychidae; (b) Philorheithridae; (c) and (d) Leptoceridae.
• Fig. 10.6 A caddisfly larva (Trichoptera: Hydropsychidae) in its retreat; the silk net is used to catch food.
• 10.4.2 Lentic adaptations
• Fig. 10.7 The whirligig beetle, Gyretes (Coleoptera: Gyrinidae), swimming on the water surface. Note that the divided compound eye allows the beetle to see both above and below water simultaneously; hydrofuge hairs on the margin of the elytra repel water.
• 10.5 Environmental monitoring using aquatic insects
• 10.6 Functional feeding groups
• 10.7 Insects of temporary waterbodies
• 10.8 Insects of the marine, intertidal and littoral zones
• Further Reading
• Chapter 11 Insects and Plants
• 11.1 Coevolutionary interactions between insects and plants
• Box 11.1 Figs and fig wasps
• 11.2 Phytophagy (or herbivory)
• Box 11.2 The grape phylloxera
• 11.2.1 Induced defences
• 11.2.2 Leaf chewing
• Fig. 11.1 Christmas beetles of Anoplognathus (Coleoptera: Scarabaeidae) on the chewed foliage of a eucalypt tree (Myrtaceae).
• 11.2.3 Plant mining and boring
• Fig. 11.2 Leaf mines: (a) linear-blotch mine of Agromyza aristata (Diptera: Agromyzidae) in leaf of an elm, Ulmus americana (Ulmaceae); (b) linear mine of Chromatomyia primulae (Agromyzidae) in leaf of a primula, Primula vulgaris (Primulaceae); (c) linear-blotch mine of Chromatomyia gentianella (Agromyzidae) in leaf of a gentian, Gentiana acaulis (Gentianaceae); (d) linear mine of Phytomyza senecionis (Agromyzidae) in leaf of a ragwort, Senecio nemorensis (Asteraceae); (e) blotch mines of the apple leaf miner, Lyonetia prunifoliella (Lepidoptera: Lyonetiidae), in leaf of apple, Malus sp. (Rosaceae); (f) linear mine of Phyllocnistis populiella (Lepidoptera: Gracillariidae) in leaf of poplar, Populus (Salicaceae); (g) blotch mines of jarrah leaf miner, Perthida glyphopa (Lepidoptera: Incurvariidae), in leaf of jarrah, Eucalyptus marginata (Myrtaceae).
• Fig. 11.3 Plant borers: (a) larvae of the European corn borer, Ostrinia nubilalis (Lepidoptera: Pyralidae), tunnelling in a corn stalk; (b) a larva of the codling moth, Cydia pomonella (Lepidoptera: Tortricidae), inside an apple.
• 11.2.4 Sap sucking
• Box 11.3 Insects and the wood of live trees
• Fig. 11.4 Feeding in phytophagous Hemiptera: (a) penetration of plant tissue by a mirid bug, showing bending of the labium as the stylets enter the plant; (b) transverse section through a eucalypt leaf gall containing a feeding nymph of a scale insect, Apiomorpha (Eriococcidae); (c) enlargement of the feeding site of (b), showing multiple stylet tracks (formed of solidifying saliva), resulting from probing of the parenchyma.
• 11.2.5 Gall induction
• Fig. 11.5 A variety of insect-induced galls: (a) two coccoid galls, each formed by a female of Apiomorpha munita (Hemiptera: Eriococcidae) on the stem of Eucalyptus melliodora; (b) a cluster of galls, each containing a male of A. munita on E. melliodora; (c) three oak cynipid galls formed by Cynips quercusfolii (Hymenoptera: Cynipidae) on a leaf of Quercus sp.; (d) rose bedeguar galls formed by Diplolepis rosae (Hymenoptera: Cynipidae) on Rosa sp.; (e) a leaf petiole of lombardy poplar, Populus nigra, galled by the aphid Pemphigus spirothecae (Hemiptera: Aphididae); (f) three psyllid galls, each induced by a nymph of Glycaspis sp. (Hemiptera: Psyllidae) on a eucalypt leaf; (g) willow bean galls of the sawfly Pontania proxima (Hymenoptera: Tenthredinidae) on a leaf of Salix sp.
• 11.2.6 Seed predation
• 11.2.7 Insects as biological control agents for weeds
• Box 11.4 Salvinia and phytophagous weevils
• 11.3 Insects and plant reproductive biology
• 11.3.1 Pollination
• Fig. 11.6 Anatomy and pollination of a tea-tree flower, Leptospermum (Myrtaceae): (a) diagram of a flower showing the parts; (b) a jewel beetle, Stigmodera sp. (Coleoptera: Buprestidae), feeding from a flower.
• Fig. 11.7 A male hawk moth of Xanthopan morganii praedicta (Lepidoptera: Sphingidae) feeding from the long floral spur of a Malagasy star orchid, Angraecum sesquipedale: (a) full insertion of the moth’s proboscis; (b) upward flight during withdrawal of the proboscis with the orchid pollinium attached.
• 11.3.2 Myrmecochory: seed dispersal by ants
• Fig. 11.8 An ant of Rhytidoponera tasmaniensis (Hymenoptera: Formicidae) carrying a seed of Dillwynia juniperina (Fabaceae) by its elaiosome (seed appendage).
• 11.4 Insects that live mutualistically in specialized plant structures
• 11.4.1 Ant–plant interactions involving domatia
• 11.4.2 Phytotelmata: plant-held water containers
• Fig. 11.9 Two myrmecophytes showing the domatia (hollow chambers) that house ants, and the food resources available to the ants. (a) A Neotropical bull’s-horn acacia, Acacia sphaerocephala (Fabaceae), with hollow thorns, food bodies and extrafloral nectaries (EFNs), which are used by the resident Pseudomyrmex ants. (b) A hollow swollen internode of Kibara (Monimiaceae) with scale insects of Myzolecanium kibarae (Hemiptera: Coccidae), which excrete honeydew that is eaten by the resident ants, Anonychomyrma scrutator.
• Fig. 11.10 A tuber of the epiphytic myrmecophyte Myrmecodia beccarii (Rubiaceae), cut open to show the chambers inhabited by ants. Ants live in smooth-walled chambers and deposit their refuse in warted tunnels, from which nutrients are absorbed by the plant.
• Fig. 11.11 A pitcher of Nepenthes (Nepenthaceae) cut open to show fly (Diptera) inquilines in the fluid: (clockwise from the top left) two mosquito larvae, a mosquito pupa, two chironomid midge larvae, a small maggot and a large rat-tailed maggot.
• Further Reading
• Chapter 12 Insect Societies
• 12.1 Subsociality in insects
• 12.1.1 Aggregation
• 12.1.2 Parental care as a social behaviour
• Parental care without nesting
• Parental care with solitary nesting
• Parental care with communal nesting
• Subsocial aphids and thrips
• Fig. 12.1 First-instar nymphs of the subsocial aphid Pseudoregma alexanderi (Hemiptera: Hormaphidinae): (a) pseudoscorpion-like soldier; (b) normal nymph.
• Quasisociality and semisociality
• 12.2 Eusociality in insects
• 12.2.1 Hymenopterans showing primitive eusociality
• Fig. 12.2 Cladogram showing proposed relationships among selected aculeate Hymenoptera to depict the multiple origins of sociality (SOL, solitary; SUB, subsocial; EU, eusocial). Apoidea includes all bees and some wasp families. Relationships within non-social aculeates are not depicted. Possible non-monophyletic families are shown in quotes on a dashed branch.
• 12.2.2 Hymenopterans showing specialized eusociality: wasps and bees
• Colony and castes in eusocial wasps and bees
• Fig. 12.3 Worker bees from three eusocial genera, from left, Bombus (bumble bees), Apis (honey bees) and Trigona (stingless bees) (all in Apidae), superficially resemble each other in morphology, but they differ in size and ecology, including their pollination preferences and level of eusociality.
• Fig. 12.4 The hind leg of a worker honey bee, Apis mellifera (Hymenoptera: Apidae): (a) outer surface showing corbicula, or pollen basket (consisting of a depression fringed by stiff setae), on the tibia, and the press on the basitarsus that pushes the pollen into the basket; (b) the inner surface with the combs and rakes that manipulate pollen into the press prior to packing.
• Fig. 12.5 Development of the honey bee, Apis mellifera (Hymenoptera: Apidae), showing the factors that determine differentiation of the queen-laid eggs into drones, workers, and queens (on the left), and the approximate developmental times (in days) and stages for drones, workers and queens (on the right).
• Box 12.1 The dance language of bees
• Nest construction in eusocial wasps
• Box 12.2 Africanized honey bees
• Fig. 12.6 The nest of the common European wasp, Vespula vulgaris (Hymenoptera: Vespidae): (a) initial stages (1–5) of nest construction by the queen (the embryonic phase of the colony’s life); (b) a mature nest.
• Nesting in honey bees
• 12.2.3 Specialized hymenopterans: ants
• Colony and castes in ants
• Nesting in ants
• Box 12.3 Colony collapse disorder
• Fig. 12.7 Weaver ants of Oecophylla making a nest by pulling together leaves and binding them with silk produced by larvae that are held in the mandibles of worker ants.
• 12.2.4 Termitoidae (former order Isoptera, termites)
• Colony and castes in termites
• Fig. 12.8 Developmental pathways of the termite Nasutitermes exitiosus (Termitidae). Heavy arrows indicate the main lines of development, light arrows indicate the minor lines. A, alate; E, egg; L, larva; LL, large larva; LPS, large presoldier; LS, large soldier; LW, large worker; N, nymph; SL, small larva; SPS, small presoldier; SS, small soldier; SW, small worker. The numbers indicate the stages.
• Nesting in termites
• Fig. 12.9 The Formosan subterranean termite, Coptotermes formosanus (Blattodea: Termitoidae: Rhinotermitidae): worker (left) and soldier (right).
• 12.2.5 Eusocial ambrosia beetles (Coleoptera: Curculionidae)
• Fig. 12.10 A “magnetic” mound of the debris-feeding termite Amitermes meridionalis (Termitidae) showing: (a) the north–south view, and (b) the east–west view.
• 12.3 Inquilines and parasites of social insects
• Fig. 12.11 Section through the mound nest of the African fungus-farming termite Macrotermes natalensis (Termitidae) showing how air circulating in a series of passageways maintains favourable culture conditions for the fungus at the bottom of the nest (a) and for the termite brood (b). Measurements of temperature and carbon dioxide are shown in the boxes for the following locations: (a) the fungus combs; (b) the brood chambers; (c) the attic; (d) the upper part of a ridge channel; (e) the lower part of a ridge channel; and (f) the cellar.
• 12.4 Evolution and maintenance of eusociality
• 12.4.1 The origins of eusociality in Hymenoptera
• Fig. 12.12 Relatedness of a given worker to other possible occupants of a hive.
• 12.4.2 The origins of eusociality in termites
• 12.4.3 Maintenance of eusociality—the police state
• 12.5 Success of social insects
• Box 12.4 Social insects as urban pests
• Further Reading
• Chapter 13 Insect Predation and Parasitism
• Scorpionfly feeding on a butterfly pupa.
• 13.1 Prey/host location
• Fig. 13.1 The basic spectrum of predator foraging and prey defence strategies, varying according to costs and benefits in both time and energy.
• 13.1.1 Sitting and waiting
• Fig. 13.2 An antlion of Myrmeleon (Neuroptera: Myrmeleontidae): (a) larva in its pit in sand; (b) detail of dorsum of larva; (c) detail of ventral view of larval head showing how the maxilla fits against the grooved mandible to form a sucking tube.
• 13.1.2 Active foraging
• Random, or non-directional foraging
• Non-random, or directional foraging
• Chemicals
• Sound
• Light
• 13.1.3 Phoresy
• 13.2 Prey/host acceptance and manipulation
• 13.2.1 Prey manipulation by predators
• Fig. 13.3 Distal part of the leg of a mantid showing the opposing rows of spines that interlock when the tibia is drawn upwards against the femur.
• Host acceptance and manipulation by parasitoids
• Fig. 13.4 Ventrolateral view of the head of a dragonfly nymph (Odonata: Aeshnidae: Aeshna) showing the labial “mask”: (a) in folded position; and (b) extended during prey capture, with opposing hooks of the palpal lobes forming claw-like pincers.
• 13.2.3 Overcoming host immune responses
• Fig. 13.5 Encapsulation of a living larva of Apanteles (Hymenoptera: Braconidae) by the haemocytes of a caterpillar of Ephestia (Lepidoptera: Pyralidae).
• 13.3 Prey/host selection and specificity
• 13.3.1 Host use by parasitoids
• Fig. 13.6 Two examples of the ovipositional behaviour of hymenopteran hyperparasitoids of aphids: (a) endophagous Alloxysta victrix (Hymenoptera: Figitidae) ovipositing into a primary parasitoid inside a live aphid; (b) ectophagous Asaphes suspensus (Hymenoptera: Pteromalidae) ovipositing onto a primary parasitoid in a mummified aphid.
• 13.3.2 Host manipulation and development of parasitoids
• Fig. 13.7 Steps in host selection by the hyperparasitoid Alloxysta victrix (Hymenoptera: Figitidae).
• Box 13.1 Viruses, wasp parasitoids and host immunity
• 13.3.3 Patterns of host use and specificity in parasites
• Fig. 13.8 Comparisons of louse and host phylogenetic trees: (a) adherence to Fahrenholz’s rule; (b) independent speciation of the lice; (c) independent speciation of the hosts.
• 13.4 Population biology—predator/parasitoid and prey/host abundance
• Fig. 13.9 An example of the regular cycling of numbers of predators and their prey: the aquatic planktonic predator Chaoborus (Diptera: Chaoboridae) and its cladoceran prey Daphnia (Crustacea).
• 13.5 The evolutionary success of insect predation and parasitism
• Further Reading
• Chapter 14 Insect Defence
• An African ant-mimicking membracid bug, shown in side and dorsal view.
• Fig. 14.1 The basic spectrum of prey defence strategies and predator foraging, varying according to costs and benefits in both time and energy.
• 14.1 Defence by hiding
• Fig. 14.2 Pale and melanic (carbonaria) morphs of the peppered moth Biston betularia (Lepidoptera: Geometridae) resting on: (a) pale, lichen-covered trunks; and (b) dark trunks.
• Fig. 14.3 A leaf-mimicking katydid, Mimetica mortuifolia (Orthoptera: Tettigoniidae), in which the fore wing resembles a leaf even to the extent of leaf-like venation and spots resembling fungal mottling.
• 14.2 Secondary lines of defence
• Box 14.1 Avian predators as selective agents for insects
• Fig. 14.4 The eyed hawk moth, Smerinthus ocellatus (Lepidoptera: Sphingidae). (a) The brownish fore wings cover the hind wings of a resting moth. (b) When the moth is disturbed, the black and blue eyespots on the hind wings are revealed.
• 14.3 Mechanical defences
• Box 14.2 Backpack bugs—dressed to kill?
• 14.4 Chemical defences
• 14.1.1 Classification by function of defensive chemicals
• 14.4.2 The chemical nature of defensive compounds
• 14.4.3 Sources of defensive chemicals
• Box 14.3 Chemically protected eggs
• Fig. 14.5 The distasteful and warningly coloured caterpillars of the cinnabar moth, Tyria jacobaeae (Lepidoptera: Arctiidae), on ragwort, Senecio jacobaeae.
• 14.4.4 Organs of chemical defence
• Fig. 14.6 A caterpillar of the orchard swallowtail butterfly, Papilio aegeus (Lepidoptera: Papilionidae), with the osmeterium everted behind its head. Eversion of this glistening, bifid organ occurs when the larva is disturbed, and is accompanied by a pungent smell.
• Box 14.4 Insect binary chemical weapons
• 14.5 Defence by mimicry
• Fig. 14.7 An aggregation of sawfly larvae (Hymenoptera: Pergidae: Perga) on a eucalypt leaf. When disturbed, the larvae bend their abdomens in the air and exude droplets of sequestered eucalypt oil from their mouths.
• 14.5.1 Batesian mimicry
• 14.5.2 Müllerian mimicry
• Fig. 14.8 Three nymphalid butterflies that are Müllerian co-mimics in Florida: (a) the monarch or wanderer (Danaus plexippus); (b) the queen (D. gilippus); (c) the viceroy (Limenitis archippus).
• 14.5.3 Mimicry as a continuum
• 14.6 Collective defences in gregarious and social insects
• Fig. 14.9 Nest guarding by the European ant Camponotus (Colobopsis) truncatus (Hymenoptera: Formicidae): a minor worker approaching a soldier that is blocking a nest entrance with her plug-shaped head.
• Fig. 14.10 Defence by mandible snapping in termite soldiers (Blattodea: Termitoidae). (a) Head of a symmetric snapping soldier of Termes in which the long thin mandibles are pressed hard together (1) and thus bent inwards (2) before they slide violently across one another (3). (b) Head of an asymmetric snapping soldier of Homallotermes in which force is generated in the flexible left mandible by being pushed against the right one (1) until the right mandible slips under the left one to strike a violent blow (2).
• Fig. 14.11 Diagram of the major components of the venom apparatus of a social aculeate wasp.
• Fig. 14.12 Three ant mimics: (a) a fly (Diptera: Micropezidae: Badisis); (b) a bug (Hemiptera: Miridae: Phylinae); (c) a spider (Araneae: Clubionidae: Sphecotypus).
• Further Reading
• Chapter 15 Medical and Veterinary Entomology
• 15.1 Insects as causes and vectors of disease
• 15.2 Generalized disease cycles
• 15.3 Pathogens
• 15.3.1 Malaria
• The disease
• Box 15.1 Life cycle of Plasmodium
• Malaria epidemiology
• Vector distribution
• Vector abundance
• Vector survival rate
• Anthropophily of the vector
• Feeding interval
• Vector competence
• Box 15.2 Anopheles gambiae complex
• Vectorial capacity
• Control of malaria
• Box 15.3 Bed nets
• 15.3.2 Arboviruses
• Box 15.4 Dengue—an emerging insect-borne disease
• Box 15.5 West Nile virus—an arbovirus disease emergent in North America
• 15.3.3 Rickettsias and plague
• 15.3.4 Protists other than malaria
• Trypanosoma
• Leishmania
• Fig. 15.1 A tsetse fly, Glossina morsitans (Diptera: Glossinidae): (a) at the commencement of feeding; and (b) fully engorged with blood. Note that the tracheae are visible through the abdominal cuticle in (b).
• 15.3.5 Filariases
• Bancroftian and brugian filariasis
• Onchocerciasis
• 15.4 Forensic entomology
• Fig. 15.2 The stages of carcass (carrion) decomposition associated with a succession of arthropod groups in guinea-pig carcasses during spring in a woodland habitat in Perth, Australia. Variation in the thickness of each band indicates the approximate relative abundance within the groups at different times.
• 15.5 Insect nuisance and phobia
• Box 15.6 Bed bugs resurge
• 15.6 Venoms and allergens
• 15.6.1 Insect venoms
• 15.6.2 Blister and urtica (itch)-inducing insects
• 15.6.3 Insect allergenicity
• Further Reading
• Chapter 16 Pest Management
• 16.1 Insects as pests
• 16.1.1 Assessment of pest status
• Fig. 16.1 Schematic graphs of the fluctuations of theoretical insect populations in relation to their general equilibrium position (GEP), economic threshold (ET) and economic injury level (EIL). From comparison of the GEP with the ET and EIL, insect populations can be classified as: (a) non-economic pests if population densities never exceed the ET or EIL; (b) occasional pests if population densities exceed the ET and EIL only under special circumstances; (c) perennial pests if the GEP is close to the ET so that the ET and EIL are exceeded frequently; or (d) severe or key pests if population densities always are higher than the ET and EIL. In practice, as indicated here, control measures are instigated before the EIL is reached.
• 16.1.2 Why insects become pests
• Box 16.1 Exotic insect pests of crops in the United States
• Tephritid fruit flies
• Light brown apple moth (LBAM)
• Asian citrus psyllid and huanglongbing (HLB)(citrus greening)
• Box 16.2 Bemisia tabaci—a pest species complex
• 16.2 The effects of insecticides
• Box 16.3 The cottony-cushion scale
• 16.2.1 Insecticide resistance
• 16.3 Integrated pest management
• 16.4 Chemical control
• 16.4.1 Insecticides (chemical poisons)
• Box 16.4 Neonicotinoid insecticides
• 16.4.2 Insect growth regulators
• 16.4.3 Neuropeptides and insect control
• 16.4.4 RNA interference and insect control
• 16.5 Biological control
• Box 16.5 Taxonomy and biological control of the cassava mealybug
• Box 16.6 Glassy-winged sharpshooter biological control—a Pacific success
• 16.5.1 Arthropod natural enemies
• Fig. 16.2 Generalized life cycle of an egg parasitoid. A tiny female wasp of a Trichogramma species (Hymenoptera: Trichogrammatidae) oviposits into a lepidopteran egg; the wasp larva develops within the host egg, pupates, and emerges as an adult, often with the full life cycle taking only one week.
• 16.5.2 Microbial control
• Nematodes
• Fungi
• Bacteria
• Viruses
• Fig. 16.3 The mode of infection of insect larvae by baculoviruses. (a) A caterpillar of the cabbage looper, Trichoplusia ni (Lepidoptera: Noctuidae), ingests the viral inclusion bodies of a granulosis virus (called TnGV) with its food and the inclusion bodies dissolve in the alkaline midgut, releasing proteins that destroy the insect’s peritrophic membrane, allowing the virions access to the midgut epithelial cells. (b) A granulosis virus inclusion body with virion in longitudinal section. (c) A virion attaches to a microvillus of a midgut cell, where the nucleocapsid discards its envelope, enters the cell and moves to the nucleus in which the viral DNA replicates. The newly synthesized virions then invade the haemocoel of the caterpillar where viral inclusion bodies are formed in other tissues (not shown).
• 16.6 Host-plant resistance to insects
• 16.6.1 Genetic engineering of host-plant resistance and the potential problems
• Box 16.7 The Colorado potato beetle
• Fig. 16.4 Area of global planting of Bt crops annually and cumulative number of insect species showing field-evolved resistance associated with reduced efficacy of Bt crops. The area of Bt crops increased from 1.1 million hectares (ha) in 1996 to 66 million hectares in 2011. * Indicates possible underestimate pending publications reporting resistance.
• 16.7 Physical control
• 16.8 Cultural control
• 16.9 Pheromones and other insect attractants
• 16.10 Genetic manipulation of insect pests
• Further Reading
• Chapter 17 Insects in a Changing World
• 17.1 Models of change
• 17.1.1 Modelling climate and insect distributions
• Fig. 17.1 Flow diagram depicting the derivation of the “ecoclimatic index” (EI) as the product of the population growth index and four stress indices. The EI value describes the climatic favourability of a given location for a given species. Comparison of EI values allows different locations to be assessed for their relative suitability to a particular species.
• 17.1.2 Climate and historic insect range changes
• Box 17.1 Modelling distributions of fruit flies
• Box 17.2 Trouble brewing? A beetle threat to coffee
• 17.2 Economically significant insects under climate change
• Fig. 17.2 The complexity of interactions between some of the many environmental factors that affect the vector–pathogen–host epidemiological cycle.
• 17.2.1 Future agricultural health
• 17.2.2 Future animal health
• 17.2.3 Future human health
• 17.3 Implications of climate change for insect biodiversity and conservation
• 17.3.1 Range change
• 17.3.2 Temporal changes and asynchrony of mutual interactions
• 17.4 Global trade and insects
• Fig. 17.3 Interactions between factors associated with insect-pest homogenization. Inset: Icerya purchasi (Hemiptera: Monophlebidae), a global pest scale insect (see Box 16.3).
• Box 17.3 Global eucalypts and their pests
• Box 17.4 Alien insects change landscapes
• Box 17.5 Insects and biosecurity—an Australian perspective
• Further Reading
• Chapter 18 Methods in Entomology: Collecting, Preservation, Curation and Identification
• 18.1 Collection
• 18.1.1 Active collecting
• 18.1.2 Passive collecting
• Fig. 18.1 A diagrammatic pitfall trap cut away to show the in-ground cup filled with preserving fluid.
• 18.2 Preservation and curation
• 18.2.1 Dry preservation
• Killing and handling prior to dry mounting
• Pinning, staging, pointing, carding, spreading and setting
• Direct pinning
• Fig. 18.2 Pin positions for representative insects: (a) larger beetles (Coleoptera); (b) grasshoppers, katydids and crickets (Orthoptera); (c) larger flies (Diptera); (d) moths and butterflies (Lepidoptera); (e) wasps and sawflies (Hymenoptera); (f) lacewings (Neuroptera); (g) dragonflies and damselflies (Odonata), lateral view; (h) bugs, cicadas, leafhoppers and planthoppers (Hemiptera: Heteroptera, Cicadomorpha and Fulgoromorpha).
• Fig. 18.3 Correct and incorrect pinning: (a) insect in lateral view, correctly positioned; (b) too low on pin; (c) tilted on long axis, instead of horizontal; (d) insect in front view, correctly positioned; (e) too high on pin; (f) body tilted laterally and pin position incorrect. Handling insect specimens with entomological forceps: (g) placing specimen mount into foam or cork; (h) removing mount from foam or cork.
• Fig. 18.4 Micropinning with stage and cube mounts: (a) a small bug (Hemiptera) on a stage mount, with position of pin in thorax as shown in Fig. 18.2h; (b) moth (Lepidoptera) on a stage mount, with position of pin in thorax as shown in Fig. 18.2d; (c) mosquito (Diptera: Culicidae) on a cube mount, with thorax impaled laterally; (d) black fly (Diptera: Simuliidae) on a cube mount, with thorax impaled laterally.
• Micropinning (staging or double mounting)
• Pointing
• Fig. 18.5 Point mounts: (a) a small wasp; (b) a weevil; (c) an ant. Carding: (d) a beetle glued to a card mount.
• Carding
• Spreading and setting
• Fig. 18.6 Spreading of appendages prior to drying of specimens: (a) a beetle pinned to a foam sheet showing the spread antennae and legs held with pins; (b) setting board with mantid and butterfly showing spread wings held in place by pinned setting paper.
• 18.2.2 Fixing and wet preservation
• 18.2.3 Microscope slide mounting
• 18.2.4 Habitats, mounting and preservation of individual orders
• Archaeognatha (Microcoryphia; archaeognathans or bristletails)
• Blattodea (cockroaches or roaches)
• Blattodea: Termitoidae (former order Isoptera; termites, “white ants”)
• Coleoptera (beetles)
• Collembola (springtails)
• Dermaptera (earwigs)
• Diplura (diplurans)
• Diptera (true flies)
• Embioptera (Embiidina, Embiodea; embiopterans or webspinners)
• Ephemeroptera (mayflies)
• Grylloblattodea (Grylloblattaria or Notoptera; grylloblattids, ice crawlers or rock crawlers)
• Hemiptera (bugs)
• Hymenoptera (ants, bees, wasps, sawflies and wood wasps)
• Lepidoptera (butterflies and moths)
• Mantodea (mantids, mantises or praying mantids)
• Mantophasmatodea (heelwalkers)
• Mecoptera (hangingflies, scorpionflies and snowfleas)
• Megaloptera (alderflies, dobsonflies and fishflies)
• Neuroptera (lacewings, owlflies and antlions)
• Odonata (damselflies and dragonflies)
• Orthoptera (grasshoppers, locusts, katydids and crickets)
• Phasmatodea (phasmids, stick-insects or walking sticks)
• Plecoptera (stoneflies)
• Protura (proturans)
• Psocodea: “Phthiraptera” (chewing lice and sucking lice)
• Psocodea: “Psocoptera” (bark lice and book lice)
• Raphidioptera (snakeflies)
• Siphonaptera (fleas)
• Strepsiptera (strepsipterans)
• Thysanoptera (thrips)
• Trichoptera (caddisflies)
• Zoraptera (zorapterans or angel insects)
• Zygentoma (silverfish)
• 18.2.5 Curation
• Labelling
• Care of collections
• 18.3 Identification
• 18.3.1 Identification keys
• 18.3.2 Unofficial taxonomies and voucher specimens
• 18.3.3 DNA-based identifications and voucher specimens
• Further Reading
• Regional texts for identifying insects
• Africa
• Australia
• Europe
• The Americas
• Identification of immature insects
• Collecting and preserving methods
• Museum collections
• Taxoboxes
• Taxobox 1 Entognatha: non-insect hexapods (Collembola, Diplura and Protura)
• Collembola (springtails)
• Diplura (diplurans)
• Protura (proturans)
• Taxobox 2 Archaeognatha (Microcoryphia; archaeognathans or bristletails)
• Taxobox 3 Zygentoma (silverfish)
• Taxobox 4 Ephemeroptera (mayflies)
• Taxobox 5 Odonata (damselflies and dragonflies)
• Taxobox 6 Plecoptera (stoneflies)
• Taxobox 7 Dermaptera (earwigs)
• Taxobox 8 Zoraptera (zorapterans or angel insects)
• Taxobox 9 Orthoptera (grasshoppers, locusts, katydids and crickets)
• Taxobox 10 Embioptera (Embiidina, Emboidea; embiopterans or webspinners)
• Taxobox 11 Phasmatodea (phasmids, stick-insects or walking sticks)
• Taxobox 12 Grylloblattodea (Grylloblattaria or Notoptera; grylloblattids, ice crawlers or rock crawlers)
• Taxobox 13 Mantophasmatodea (heelwalkers)
• Taxobox 14 Mantodea (mantids, mantises or praying mantids)
• Taxobox 15 Blattodea: roach families (cockroaches or roaches)
• Taxobox 16 Blattodea: epifamily Termitoidae (former order Isoptera; termites, “white ants”)
• Taxobox 17 Psocodea: “Psocoptera” (bark lice and book lice)
• Taxobox 18 Psocodea: “Phthiraptera” (chewing lice and sucking lice)
• Taxobox 19 Thysanoptera (thrips)
• Taxobox 20 Hemiptera (bugs, moss bugs, cicadas, leafhoppers, planthoppers, spittle bugs, treehoppers, aphids, jumping plant lice, scale insects and whiteflies)
• Taxobox 21 Neuropterida: Neuroptera (lacewings, owlflies and antlions), Megaloptera (alderflies, dobsonflies and fishflies) and Raphidioptera (snakeflies)
• Neuroptera
• Megaloptera
• Raphidioptera
• Taxobox 22 Coleoptera (beetles)
• Taxobox 23 Strepsiptera (strepsipterans)
• Taxobox 24 Diptera (true flies)
• Taxobox 25 Mecoptera (hangingflies, scorpionflies and snowfleas)
• Taxobox 26 Siphonaptera (fleas)
• Taxobox 27 Trichoptera (caddisflies)
• Taxobox 28 Lepidoptera (butterflies and moths)
• Taxobox 29 Hymenoptera (ants, bees, wasps, sawflies and wood wasps)
• Back Matter
• Glossary
• Further Reading
• References
• Index
• Appendix: A Reference Guide to Orders
• Color Plates
• Plate 1
• Plate 2
• Plate 3
• Plate 4
• Plate 5
• Plate 6
• Plate 7
• Plate 8
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