The Remaining Endopterygote Orders

FIGURE 10.7. Caraboidea. (A) A ground beetle, Calosoma sycophanta (Carabidae); (B) C. sycophanta larva; (C) a tiger beetle, Cicindela sexguttata (Carabidae); (D) Peltodytes edentulus (Haliplidae); (E) a diving beetle, Dytiscus verticalis (Dytiscidae); (F) Dytiscus sp. larva; (G) a whirligig beetle, Dineutes americanus (Gyrinidae); and (H) D. americanus larva [A, B, from L. A. Swan and C. S. Papp, 1972, The Common Insects of North America. Copyright 1972 by L. A. Swan and C. S. Papp. Reprinted by permission of Harper & Row, Publishers, Inc. C, D, from R. H. Arnett, Jr., 1968, The Beetles of the United States (A Manual for Identification). By permission of the author. F, from E. S. Dillon and L. S. Dillon, 1972, A Manual of Common Beetles of Eastern N. America. By permission of Dover Publications, New York. H, from A. G. Boving and F. C. Craighead, 1930, An illustrated synopsis of the principal larval forms of the order Coleoptera, Entomol. Am. XI:1-351. Published by the Brooklyn Entomological Society. By permission of the New York Entomological Society.]

movable, the testes are not tubular and coiled, and the ovarioles are acrotrophic. The legs of larvae are either four-segmented (without tarsus) plus single claw, vestigial, or absent. The larval mandibles may or may not have a molar area.

Within the Polyphaga six major series (evolutionary lines) can be recognized (Figure 10.6). Crowson (1960) suggested that the first adaptive radiation of the Polyphaga occurred in the Triassic when three ancestral stocks had their origins: the staphyliniform, eucinetoid, and dermestoid. The former gave rise to the modern Staphyliniformia. The eucinetoid group evolved into the Eucinetiformia, Scarabaeiformia, and Elateriformia, and the dermestoid group was ancestral to the Bostrichiformia and Cucujiformia. The latter series includes more than one half of the total beetle species and can therefore be considered as the most highly evolved group within the order.

FIGURE 10.8. Hydrophiloidea. Hydrophilus triangularis (Hydrophilidae).

Series Staphyliniformia

Three superfamilies are included in the series Staphyliniformia: the Histeroidea, Hy-drophiloidea, and Staphylinoidea. The Hydrophiloidea appear to be the most primitive group.

Superfamily Hydrophiloidea

More than 80% of the 2400 species included in the Hydrophiloidea belong to the family HYDROPHILIDAE (Figure 10.8), whose members are mainly aquatic beetles somewhat similar to Dytiscidae. However, adults are scavengers rather than predators, do not usually rest head downward at the surface of the pond, and, when swimming, move their legs alternately rather than synchronously. The larvae are predaceous. Some hydrophilids are terrestrial, but restricted to damp places, for example, in decaying plant material or dung.

Superfamily Histeroidea

Almost all of the approximately 3000 species of Histeroidea are placed in the family HISTERIDAE (Figure 10.9), both the adults and larvae of which feed on other insects. These small, usually shiny black beetles with short elytra (which leave the last one or two abdominal segments exposed) are found under bark, in rotting animal or vegetable matter including dung, or in ant nests, where they produce appeasement substances over their bodies to gain acceptance by their hosts.

FIGURE 10.9. Histeroidea. Margarinotus immunis (Histeridae). [From R. H. Arnett, Jr., 1968, The Beetles of the United States (A Manual for Identification). By permission of the author.]

FIGURE 10.10. Staphylinoidea. (A) The hairy rove beetle, Staphylinus maxillosus villosus (Staphylinidae); and (B) an antlike stone beetle, Euconnus clavipes (Scydmaenidae). [From R. H. Arnett, Jr., 1968, The Beetles of the United States (A Manual for Identification).By permission of the author.]

Superfamily Staphylinoidea

The very large superfamily Staphylinoidea contains nearly 40,000 species, adults of which are characterized by unusually short elytra that leave about half of the abdomen visible. Some 30,000 of these species belong to the family STAPHYLINIDAE (rove beetles) (Figure 10.10A), a group of very diverse habits. Most appear to be carnivorous or saprophagous, though precise details of their feeding habits are not known. They occur in many places, for example, in decaying animal or vegetable matter, under stones or bark, on flowers, under seaweed, in moss or fungi, and in the nests of birds, mammals, and social insects. The PSELAPHIDAE (about 5000 species) closely resemble rove beetles both morphologically and in habits. They are found, moreover, in similar habitats. Adult SCYDMAENIDAE (2000 species), in contrast to those of the two families above, have fully developed elytra. These small to minute, hairy predators are somewhat antlike in appearance (Figure 10.10B), being found under stones, in humus, or sometimes in ant nests. The ANISOTOMIDAE (LEIODIDAE) form a family of about 2000 species of beetles that are found in decaying organic matter, under bark, etc. A number of species have become adapted for a cave-dwelling existence.

Series Eucinetiformia

Superfamily Eucinetoidea

About 90% of the nearly 700 species in this group belong to the SCIRTIDAE (= HELODIDAE), a worldwide family with greatest diversity in the temperate regions. Adults are usually found on vegetation near water, while larvae occupy a variety of still-water habitats and occasionally wet, rotting wood where they are filter feeders on algae, diatoms, fungi, and other organic material.

Series Scarabaeiformia

Superfamily Scarabaeoidea

Of the nearly 28,000 species in the superfamily Scarabaeoidea, about 25,000 are included in the family SCARABAEIDAE. Adults may be recognized by the lamellate terminal

FIGURE 10.10. Staphylinoidea. (A) The hairy rove beetle, Staphylinus maxillosus villosus (Staphylinidae); and (B) an antlike stone beetle, Euconnus clavipes (Scydmaenidae). [From R. H. Arnett, Jr., 1968, The Beetles of the United States (A Manual for Identification).By permission of the author.]

FIGURE 10.11. Scarabaeoidea. (A) Amalerhinoceros beetle, Dynastes tityus (Scarabaeidae); (B) a May beetle, Phyllophaga rugosa (Scarabaeidae); and (C) the giant stag beetle, Lucanus elaphus (Lucanidae). [From L. A. Swan and C. S. Papp, 1972, The Common Insects of North America. Copyright 1972 by L. A. Swan and C. S. Papp. Reprinted by permission of Harper & Row Publishers, Inc.]

FIGURE 10.11. Scarabaeoidea. (A) Amalerhinoceros beetle, Dynastes tityus (Scarabaeidae); (B) a May beetle, Phyllophaga rugosa (Scarabaeidae); and (C) the giant stag beetle, Lucanus elaphus (Lucanidae). [From L. A. Swan and C. S. Papp, 1972, The Common Insects of North America. Copyright 1972 by L. A. Swan and C. S. Papp. Reprinted by permission of Harper & Row Publishers, Inc.]

segments of their antennae. Many common beetles belong to the family, including dung beetles (SCARABAEINAE = COPRINAE) (Figure 24.3), cockchafers (May or June bugs) (MELOLONTHINAE) (Figure 10.11B), shining leaf chafers (known as Christmas beetles in Australia) (RUTELINAE), and the large and striking elephant (rhinoceros) beetles (DYNASTINAE) (Figure 10.11A). Most scarabaeids feed on decaying organic matter, especially dung, in both adult and juvenile stages, though there are many variations of this theme. Larvae of some species feed underground on plant roots and a few live in termite nests. Adults frequently feed on nectar, foliage, or fruit, or they do not feed at all. The family LUCANIDAE (stag beetles) (Figure 10.11C) includes about 1200 species in which the adults are sexually dimorphic. The mandibles of males are enormously enlarged, though the significance of this is not understood. Larvae generally feed on rotting wood. Adults are mainly nectar, occasionally foliage, feeders. The PASSALIDAE form a mainly tropical, forest-dwelling family of about 500 species. Large numbers of these beetles are often found in the same log, and it appears that the adults assist in feeding the larvae by partially chewing the rotting wood beforehand. The GEOTRUPIDAE (600 species) is a worldwide family found especially in drier regions. The beetles, which often have conspicuous horns on the head and prothorax, dig burrows that they provision with food (usually fungi, but some species use dung or decaying organic material) prior to egg laying.

Superfamily Dascilloidea

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