Beetles

The Coleoptera, or beetles, includes many commonly encountered insects such as ladybird beetles (family Coccinellidae), click beetles (Elateridae), scarabs (Scarabaeidae), and fireflies (Lampyridae). They live throughout the world (except Antarctica), but are most speciose in the tropics.

The oldest beetle fossils are from the Lower Permian (about 265 million years old; Ponomarenko, 1995); since then the group has diversified into many different forms. They range in size from minute featherwing beetles (Ptiliidae), adults of which are as small as 0.3 mm long, to the giant Goliath and Hercules beetles (Scarabaeidae), which can be well over 15 cm. While most species are phytophagous, many are predacious, or fungivores, or are parasitoids. They communicate to one another in many ways, either by use of chemicals (e.g. pheromones), sounds (e.g. stridulation), or by visual means (e.g. fireflies). They live in rainforest canopies, the driest deserts, in lakes, and above tree lines on mountains.

In one sense the most unusual property of beetles is not some aspect of their structure or natural history, but their sheer number. There are more known species of Coleoptera than any other group of organisms, with over 350,000 described species.

The most distinctive feature of beetles is the hardening of the forewings into elytra; it is from this that they get their formal name (koleos - sheath, pteron - wing). The elytra serve to protect the more delicate hind wings, as well as the dorsal surface of the abdomen, and may have been a key factor allowing them to exploit narrow passageways (for example, in leaf litter and under bark). During flight the forewings are opened enough to allow the hind wings to unfold and function:  

Other derived characteristics of beetles are:

• hind wings folded under elytra, with reduced venation
• hind two thoracic segments (mesothorax + metathorax = pterothorax) broadly connected with abdomen, so that the primary functional units of body are head / prothorax / pterothorax + abdomen, rather than the more typical head / thorax / abdomen of many other insects.
• genitalia retracted into abdomen
• adult antenna with 11 articles

Beetles are holometabolous insects, normally with adecticous, exarate pupae. Most species have chewing mouthparts. There is a gula present on the undersurface of the head.

The four living suborders of beetles diverged from one another in the Permian and early Triassic, and are substantially different from one another. Adults differ in the structure of the prothorax, hind wing, abdomen, ovary, testes, and so on. The major differences are summarized in a table.

Polyphaga is by far the largest suborder, containing 85% of the known species, including rove beetles, scarabs, stag beetles, metallic wood-boring beetles, click beetles, fireflies, blister beetles, mealworms, ladybirds, leaf beetles, longhorn beetles, and weevils. Many are phytophagous. Adephaga includes ground beetles, tiger beetles, predacious diving beetles, and whirligig beetles; most adephagans are predacious. Myxophaga is a small suborder, containing less than 100 known species, whose members are small or minute, and associated with hygropetric habitats, drift material, or interstitial habitats among sand grains. Archostemata contains several families of beetles, most associated with wood; members of this family are somewhat similar to some of the earliest, Paleozoic beetle fossils.