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Classification Kingdom of the Animal exapodsstand out among all other invertebratesfor being far and away the most diverse group of animals on Earth, the only inverte- bratesto fly, and the only terrestrial invertebratesto undergo indirect developmentor completemetamorphosis. The arthropod subphylum Hexapoda comprisesthe classInsecta and three other small, closelyrelated, wingless, insect-likegroups: Collembola, protura, and Diplura. The Hexapodaare united on the basisof a distinct body plan of a head,3-segmented thorax, and 11-segmentedabdomery 3 pairs of thoricic legs, a single pair of antennae,3 setsof "jaws" (mandibles,maxillae, and labium), an aerial gasexchange system composed of tracheaeand spiracles,Malpighian tubules formed as proctodeal (ectodermal)evaginations, and, among the Pterygota, wings (Box 22A). The presence of a tho- rax fixed at 3 segments, each with a pair of walking legs, is a unique slmapomorphy for the Hexapoda. Classificationof The Animal Other synapomorphies include the presence of a Kingdom(Metazoa) large fat body (mainly concentrated in the abdo- men), and fusion of the second maxillae to form a Non-Bilateria* Lophophorata' lowerlip (thelabium). (a.k.a.the diploblasts) PHYLUM PHORONIDA PHYLUM PORIFERA PHYLUM BRYOZOA Hexapods evolved on land; groups inhabit- pHYrDv pLAcozoA PHYLUM BRACHIOPODA ing aquatic environments today have secondarily PHYLUM CNIDARIA EcpysozoA invaded those habitats through behavioral adap- PHYLUM CTENOPHORA Nematoida tations and modifications of their aerial gas ex- PHYLUM Bilateria NEMATODA change systems. The earliest undisputed fossils of PHYLUM NEMATOMORPHA (a.k.a.the triploblasts) hexapods are early Devonian (412Ma). However, Scalidophora PHYLUM XENACOELOMORPHA PHYLUM KINORHYNCHr'. there are Silurian trace fossils that are very hexa- Protostomia PHYLUM PRIAPULA pod-like, and molecular clock data suggest an PHYLUM CHAETOGNATHA PHYLUM LOBICIFERA Early Ordovician origin for Hexapods at about Sptneura Panarthropoda 479 millionyears ago and PHYLUM PLATYHELMINTHES an early Silurian origin PHYLUM TARDIGRADA PHYLUM GASTROTRICHA about M7 million years ago for Insecta. PHYLUM ONYCHOPHORA PHYLUM RHOMBOZOA The most spectacular evolutionary PHYLUM ARTHROPODA radiation PHYLUM ORTHONECTIDA suBpHvLUMcRusrncen* among the Hexapoda (in fact among all eukary- PHYLUM NEMERTEA SUAPHYLUMH€XAPObA otic life) has, of course, been within the insects, PHYLUM MOLLUSCA SUBPHYLUM MYRIAPODA which inhabit nearly every PHYLUM ANNELIDA conceivable terrestrial SUBPHYLUM CHELICERATA PHYLUM ENTOPROCTA and freshwater habitat and, less commonly, even Deuterostomia PHYLUM CYCLIOPHORA the sea surface and the marine littoral iegion. PHYLUM ECHINODERMATA Gnathifera PHYLUM HEMICHORDATA lnsects are also found in such unlikely places as oil PHYLUM GNATHOSTOMULIDA PHYLUM CHORDATA swamps and seeps, sulfur springs, glacial streams, PHYLUM MICROGNATHOZOA and brine ponds. They often live where few other PHYLUM ROTIFERA "Paraphyletic group This chapterhas been revised by Wendy Moore. 844 Chapter Twenty-Two Insects are not only diverse, but also incredibly BOX22A Characteristicsof the abundant. For every human alive, there are an estimat- SubphylumHexapoda ed 200 million insects. Howard Ensign Evans estimat- ed that an acre of ordinary English pasture supports (plus 1. Body composed of 20 true somites acron) an astonishin g 248,375,000 springtails and 17,825,000 organized as a head (6 somites), thorax (3 somites) beetles. In tropical rain forests, insects can constitute and abdomerl (11 sopites). Due to fusion of (dry somites, these body segments are not always 40% of the total animal biomass weight), and the externallyobvious. biomass of the ants can be far greater than that of the 2. Head segments bear the following structures (from combined mammal fauna (up to 75"/" of the total ani- anterior to posterior):compound eyes and ocelli; mal biomass). A single colony of the African driver ant antennae; clypeolabrum; mandibles; maxillae,labi- Anomma wilaerthi may contain as many as 22 million um (fused second maxillae).Ocelli (and compound workers. Based on his research in the tropics, biodi- eyes) are secondarily lost in some groups. versitv sleuth Terrv Erwin has calculated that there 3. Legs uniramous; present on the three thoracic seg- are about 3.2 x 108individual arthropods per hectare, ments of adults; legs composed of 6 articles: coxa, representing more than 60,000 species, in the west- pretarsus; trochanter, femur, tibia, tarsus, tarsus ern Amazon. In Maryland, a single population of the often subdivided; pretarsus typically clawed mound-building ant Formicn exsectoidescomprised 73 4. Gas exchange by spiracles and tracheae nests covering an area of 10 acres and containing ap- 5. Gut with gastric (digestive)ceca proximately 12 million workers. Termites have colo- 6. With large fat body (mainlyconcentrated in nies of similar magnitudes. E. O. Wilson has calculated abdomen) that, at any given time, 101s(a million billion) ants are 7. Fused exoskeleton of head forms unique internal alive on Earth! tentorium In most parts of the world, insects are among the B. With ectodermally derived Malpighiantubules (proc- principal predators of other invertebrates. Insects are todeal evaginations) also key items in the diets of many terrestrial verte- 9. Gonopores open on the last abdominal segment, brates, and they play a major role as reducer-level or- or on abdominalsegment 7, B, or I ganisms (detritovores and decomposers) in food webs. 10. Gonochoristic; direct or indirect development Due to their sheer numbers, they constitute much of the matrix of terrestrial food webs. Their biomass and energy consumption exceed those of vertebrates in most terrestrial habitats. In deserts and in the tropics, animals or plants can exist. It is no exaggeration to say ants replace earthworms as the most abundant earth that insects rule the land. Their diversity and abun- movers (ants are nearly as important as earthworms dance defy imagination (Figures 22.1,-22.7). even in temperate regions). Termites are among the We do not know how many species of insects there chief decomposers of dead wood and leaf litter around are, or even how many have been described. Published the world, and without dung beetles African savan- estimates of the number of described species range nahs would be buried under the excrement of the tens from 890,000 to well over a million (we calculate of thousands of large grazingmammals. about926,990). An average of about 3,500 new species Without insects, life as we know it would ceaseto have been described annually since the publication exist. In fact,E. O. Wilson has stated, "so important of Linnaeus's SystemaNsturne in1758, although in re- are insects and other land-dwelling arthropods that if cent years the average has climbed to 7,000 new spe- all were to disappear, humanity probably could not cies annually. Estimates of the number of insect species last more than a few months." Eighty percent of the remaining to be described range from 3 million to 100 world's crop species, including food, medicine, and million. The Coleoptera (beetles), with an estimated fiber crops, rely on animal pollinators, nearly all of 380,000described species,is far and away the largest which are insects. Insects also play key roles in pol- insect order (more than a quarter of all animal species linating wild, native plants. Beekeeping began long are beetles). The beetle family Curculionidae (the wee- ago/ at least by 600 BC in the Nile Valley and prob- vils) contains about 65,000 described species (nearly ably well before that. The first migratory beekeep- 5% of all described animal species).The rich diversity ers were Egyptians who floated hives up and down of insects seems to have come about through a combi- the Nile to provide pollination services to floodplain nation of advantageous features, including the evolu- farmers while simultaneously producing a honey tionary exploitation of developmental genes working crop. Domestic honeybees (Apis mellifera),introduced on segmented and compartmen talize d bodies, coevo- to North America from Europe in the mid-1600s, are lution with plants (particularly the flowering plants), now the dominant pollinators of most food crops miniaturization, and the invention of flight. grown around the world, and they play some role PHYLUMARTHROPODA The Hexaooda:Insects and Their Kin 845 Figure 22.1 Representatives of the three orders of entognathous (noninsect) hexapods. (A) Anurida grana- na, a springtail (order Collembola). (B) Ptenothrix sp., a resulted in flowers with anatomy and scentsthat are springlail showing entognathous mouthparts. (C) A diplu- finely tuned to their insect partners. In exchangefor ran from New Zealand (order Diplura). (D) A proturan from pollination services,flowers provide insects with British Columbia (order Protura). food (nectar, pollen), shelter, and chemicals used by the insectsto produce such things as pheromones.In general,insect pollination is accomplishedcoinciden- in pollinatingS}% of the crop varieties grown in the tally, as the pollinators visit flowers for other reasons. United States(they are estimated to be directly re- But in a few cases,such as that of the yucca moths of sponsiblefor $10to $20billion in crops annually).l the American Southwesl (Tegiticulaspp.), the insects Interactions between insects and flowering plants actually gather up pollen and force it into the recep- have been going on for a very long time, beginning tive stigma of the flower, initiating pollination. The over 100 million years
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