The cockroaches are an ancient group, with their ancestors, known as "roachoids", originating during the Carboniferous period, some 320 million years ago. Those early ancestors, however, lacked the internal ovipositors of modern roaches. Cockroaches are somewhat generalized insects lacking special adaptations (such as the sucking mouthparts of aphids and other true bugs); they have chewing mouthparts and are probably among the most primitive of living Neopteran insects. They are common and hardy insects capable of tolerating a wide range of climates, from Arctic cold to tropical heat. Tropical cockroaches are often much larger than temperate species.
Modern cockroaches are not considered to be a monophyletic group, as it has been found based on genetics that termites are deeply nested within the group, with some groups of cockroaches more closely related to termites than they are to other cockroaches, thus rendering Blattaria paraphyletic. Both cockroaches and termites are included into Blattodea.
Some species, such as the gregarious German cockroach, have an elaborate social structure involving common shelter, social dependence, information transfer and kin recognition. Cockroaches have appeared in human culture since classical antiquity. They are popularly depicted as large, dirty pests, although the majority of species are small and inoffensive and live in a wide range of habitats around the world.
Distribution and habitat
Cockroaches are abundant throughout the world and live in a wide range of environments, especially in the tropics and subtropics.[31] Cockroaches can withstand extremely low temperatures, allowing them to live in the Arctic. Some species are capable of surviving temperatures of −122 °C (−188 °F) by manufacturing an antifreeze made out of glycerol.[32] In North America, 50 species separated into five families are found throughout the continent.[31] 450 species are found in Australia.[33] Only about four widespread species are commonly regarded as pests.[34][35]
Cockroaches occupy a wide range of habitats. Many live in leaf litter, among the stems of matted vegetation, in rotting wood, in holes in stumps, in cavities under bark, under log piles and among debris. Some live in arid regions and have developed mechanisms to survive without access to water sources. Others are aquatic, living near the surface of water bodies, including bromeliad phytotelmata, and diving to forage for food. Most of these respire by piercing the water surface with the tip of the abdomen which acts as a snorkel, but some carry a bubble of air under their thoracic shield when they submerge. Others live in the forest canopy where they may be one of the main types of invertebrate present. Here they may hide during the day in crevices, among dead leaves, in bird and insect nests or among epiphytes, emerging at night to feed.[36]
Biology
Digestive tract
Cockroaches are generally omnivorous; the American cockroach (Periplaneta americana), for example, feeds on a great variety of foodstuffs including bread, fruit, leather, starch in book bindings, paper, glue, skin flakes, hair, dead insects and soiled clothing.[54] Many species of cockroach harbor in their gut symbiotic protozoans and bacteria which are able to digest cellulose. In many species, these symbionts may be essential if the insect is to utilize cellulose; however, some species secrete cellulase in their saliva, and the wood-eating cockroach, Panesthia cribrata, is able to survive indefinitely on a diet of crystallized cellulose while being free of microorganisms.[55]
The similarity of these symbionts in the genus Cryptocercus to those in termites are such that these cockroaches have been suggested to be more closely related to termites than to other cockroaches,[56] and current research strongly supports this hypothesis about their relationships.[57] All species studied so far carry the obligate mutualistic endosymbiont bacterium Blattabacterium, with the exception of Nocticola, an Australian cave-dwelling genus without eyes, pigment or wings, which recent genetic studies indicate is a very primitive cockroach.[58][59] It had previously been thought that all five families of cockroach were descended from a common ancestor that was infected with B. cuenoti. It may be that N. australiensis subsequently lost its symbionts, or alternatively this hypothesis will need to be re-examined.[59]
Tracheae and breathing
Like other insects, cockroaches breathe through a system of tubes called tracheae which are attached to openings called spiracles on all body segments. When the carbon dioxide level in the insect rises high enough, valves on the spiracles open and carbon dioxide diffuses out and oxygen diffuses in. The tracheal system branches repeatedly, the finest tracheoles bringing air directly to each cell, allowing gaseous exchange to take place.[60]
While cockroaches do not have lungs as do vertebrates, and can continue to respire if their heads are removed, in some very large species, the body musculature may contract rhythmically to forcibly move air in and out of the spiracles; this may be considered a form of breathing.[60]
Reproduction
Cockroaches use pheromones to attract mates, and the males practice courtship rituals, such as posturing and stridulation. Like many insects, cockroaches mate facing away from each other with their genitalia in contact, and copulation can be prolonged. A few species are known to be parthenogenetic, reproducing without the need for males.[28]
Female cockroaches are sometimes seen carrying egg cases on the end of their abdomens; the German cockroach holds about 30 to 40 long, thin eggs in a case called an ootheca. She drops the capsule prior to hatching, though live births do occur in rare instances. The egg capsule may take more than five hours to lay and is initially bright white in color. The eggs are hatched from the combined pressure of the hatchlings gulping air. The hatchlings are initially bright white nymphs and continue inflating themselves with air, becoming harder and darker within about four hours. Their transient white stage while hatching and later while molting has led to claims of albino cockroaches.[28] Development from eggs to adults takes three to four months. Cockroaches live up to a year, and the female may produce up to eight egg cases in a lifetime; in favorable conditions, she can produce 300 to 400 offspring. Other species of cockroaches, however, can produce far more eggs; in some cases a female needs to be impregnated only once to be able to lay eggs for the rest of her life.[28]
The female usually attaches the egg case to a substrate, inserts it into a suitably protective crevice, or carries it about until just before the eggs hatch. Some species, however, are ovoviviparous, keeping the eggs inside their body, with or without an egg case, until they hatch. At least one genus, Diploptera, is fully viviparous.[28]
Cockroaches have incomplete metamorphosis, meaning that the nymphs are generally similar to the adults, except for undeveloped wings and genitalia. Development is generally slow, and may take a few months to over a year. The adults are also long-lived; some have survived for as many as four years in the laboratory.[28]