Blattodea

Blattodea; Temporal range: Late Jurassic–Present PreꞒ Ꞓ O S D C P T J K Pg N Stem groups present since Late Carboniferous
	Domino cockroach Therea petiveriana
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Superorder:	Dictyoptera
Order:	Blattodea ; Wattenwyl, 1882
Superfamilies
	Corydioidea ; Blaberoidea ; Blattoidea (includes termites);
Synonyms
	Blattaria (cockroaches only);

Last updated November 15, 2024

Blattodea is an order of insects that contains cockroaches and termites.^[3] Formerly, termites were considered a separate order, Isoptera, but genetic and molecular evidence suggests they evolved from within the cockroach lineage, cladistically making them cockroaches as well.^[4] The Blattodea and the mantis (order Mantodea) are now all considered part of the superorder Dictyoptera. Blattodea includes approximately 4,400 species of cockroach in almost 500 genera, and about 3,000 species of termite in around 300 genera.

Termites are pale-coloured, soft-bodied eusocial insects that live in colonies, whereas cockroaches are darker-coloured (often brown), sclerotized, segmented insects. Within the colony, termites have a caste system, with a pair of mature reproductives, the king and the queen, and numerous sterile workers and soldiers. Cockroaches are not colonial but do have a tendency to aggregate and may be considered pre-social, as all adults are capable of breeding. Other similarities between the two groups include various social behaviours, trail following, kin recognition, and methods of communication.

Phylogeny and evolution

Cladistic analysis of five DNA sequences in 107 species representing all the termite subfamilies, all six cockroach families, including 22 of the 29 subfamilies, and five of the 15 mantis families (as out-groups) showed that the termites are nested within the cockroaches, and that the Cryptocercidae is a sister group to the termites. The mantids were shown to be the sister group to Blattodea.^[4]Cryptocercus also shares characteristics such as species of gut bacteria with the termites.^[5]

The cockroach families Lamproblattidae and Tryonicidae are not shown but are placed within the superfamily Blattoidea. The cockroach families Corydiidae and Ectobiidae were previously known as the Polyphagidae and Blattellidae.^[6]^[7]

The evolutionary relationships of the Blattodea (cockroaches and termites), based on Eggleton, Beccaloni & Inward (2007) and modified by Evangelista et al. 2019, are shown in the cladogram:^[8]^[9]

The cladogram shows the family Alienopteridae (originally assigned to its own order "Alienoptera") as sister to Mantodea; while it was reassigned to the extinct Blattodea superfamily Umenocoleoidea by Vršanský et al.,^[10] a more recent analysis places Alienopteridae and Umenocoleidae as sister taxa within Dictyoptera, and not within Blattodea.^[11]

Dictyoptera

Manipulatoridae (extinct)

	Alienopteridae (extinct)

	Mantodea (mantises)

Blattodea

Blaberoidea

Ectobiinae

Pseudophyllodromiinae

	Nyctiborinae

	Blattellinae

Blaberidae (giant cockroaches)

Ectobiidae (formerly Blattellidae)

Solumblattodea

Corydioidea

	Corydiidae (sand cockroaches, etc.)

	Nocticolidae (cave cockroaches, etc.)

Blattoidea

Blattoidae

	Tryonicidae

	Blattidae (Oriental, American and other cockroaches)

Cryptocercidae (brown-hooded cockroaches)

Termitoidae

Mastotermitidae

Euisoptera

	Archotermopsidae

	Hodotermitidae

	Serritermitidae

	Termitidae

(termites)

Diversity

Over 4,000 species of cockroaches are found in every corner of the globe with each continent having its own indigenous species. Most of these are omnivores or detritivores and live in a range of habitats such as among leaf litter, in rotting wood, in thick vegetation, in crevices, in cavities beneath bark, under logs and among debris. Some are arboreal, some live in caves and some are aquatic.^[12] A small number of species have taken to living in close proximity to humans in buildings, have been transported around the world by them, and are regarded as pests.^[13] Although some species harbour symbionts in their guts which facilitate cellulose digestion, many species also produce enzymes to digest cellulose independent of the symbionts.^[14]

Over 3,000 species of termite are found in all the continents except Antarctica. The greatest diversity is found in Africa and relatively few species inhabit Europe and North America. They are also detrivores and many species eat wood, having specialised guts with symbiotic protozoa to digest the cellulose. Termites have soft bodies and keep out of sight as far as possible. They can loosely be subdivided into dampwood, drywood and subterranean types. In general, dampwood termites inhabit coniferous forests, drywood termites inhabit hardwood forests and subterranean termites live in a wide variety of habitats.^[15]

Characteristics

Termites are eusocial insects that live in colonies. They have a caste system, with a king and queen in each colony and many non-reproductive workers. The workers forage for food which they bring back to the colony to feed the reproductives and the developing young.^[16] Cockroaches are also social insects but do not live in colonies, and all adults are able to reproduce. Some species form aggregations, others show an inclination to aggregate, and some exhibit parental care of their offspring.^[17]

Cockroaches and termites have striking similarities in behaviour which they likely inherited from their common ancestor. These include an attraction to warm and humid places, thigmotaxis, burrowing, substrate manipulation, hygienic behaviour, food sharing, cannibalism, excretion behaviour, vibrational communication, kin recognition, trail following, allogrooming, care of the brood, cropping of antennae and certain mating behaviours.^[18] In some of these behaviours, there are marked similarities between termites and juvenile, but not adult, cockroaches. During the evolution of eusociality, the individuals need to share a desire to group together. Juvenile cockroaches have a tendency to aggregate while adults often compete aggressively with each other for space and resources. Similarly, grooming and being groomed is common in termite colonies but allogrooming is not a behaviour generally engaged in by cockroaches although individuals groom themselves.^[18] An exception to this is the cockroach Cryptocercus, which seems to be more closely related to the termites than to other cockroaches. ^[19] Here juveniles groom each other and also groom adults.^[18]

Both groups are also affected by their social environments. A single termite, kept alone, has a significantly decreased level of vigour and a shorter lifespan than when two are kept together. An isolated cockroach nymph may grow at less than half the rate of grouped individuals, and has a poorer life expectancy.^[18]

Both termites and cockroaches engage in coprophagy, the consumption of fecal pellets. Adult termite workers forage and bring food back to the nest where they pass it to the reproductives and young either by mouth or by anus, providing the whole of their nutritional needs in this manner. Young cockroaches are ineffective foragers, seldom straying from their hiding places, and obtain much of their nourishment from eating the fecal pellets of larger individuals. From these they acquire the microbial flora that helps them to digest their food.^[18]

A single cockroach family, the Cryptocercidae, and one primitive species of termite, Mastotermes darwiniensis , share such characteristics as the segmental origin of certain female reproductive structures, and the fact that both deposit their eggs in the oothecae that are typical of cockroaches.^[20]

Cockroaches

American cockroach American-cockroach.jpg — American cockroach

Arthropods similar to living cockroaches dominated the insect communities of the Carboniferous period. Modern crown group cockroaches radiated from them by the middle of the Mesozoic,^[21] with the first appearance of the extant family Corydiidae during the Late Jurassic.^[1] This group of insects are nocturnal, only foraging for food and water at night. They are not considered eusocial because their populations are not divided into different caste systems; however, they are still social creatures and can live in groups with over a million individuals.^[22] The cockroach is flattened dorsolaterally and is roughly oval with a shield-like plate, the pronotum, covering its thorax and posterior region of the head. The antennae are many-segmented, long and slender, and the mouthparts are adapted for chewing. The forewings are normally leathery and the hind wings membranous. The coxae of the legs are flattened to enable the femurs to fit neatly against them when folded. Cockroaches are hemimetabolous; there is no pupal stage and the nymphs resemble the adults apart from their size and the absence of wings.^[21] Female cockroaches produce an egg sac known as an ootheca and can hold anywhere from 12-25 eggs depending upon the species.^[23] Some species display parenting behavior, whereas other species have nothing to do with the young. In most species, growth to maturity takes three to four months,^[24] but in a few species, the nymph stage can last for several years. The main factors affecting the duration of the nymph stage are seasonal differences, and the amount of nutrients received in the diet.^[25]

Chemical communication

As in most insect species, cockroaches communicate with one another by the release of pheromones. It has also been discovered that cockroaches release hydrocarbons from their body that are transferred through interactions of the antennae. These hydrocarbons can aid in cockroach communication and can even tell whether an individual is a member of its kin or not to prevent inbreeding. Cockroaches that have been isolated in a lab setting have shown extreme behavioral effects and are less stimulated by these hydrocarbons and pheromones, possibly suggesting a group environment is required for development of these communication skills.^[22]

Termites

All species of termite are to some degree eusocial, and the members of a colony are differentiated into caste systems. The majority of termite populations consist of the worker caste, which are responsible for foraging, nest building, grooming, and brood care. The soldier caste has one responsibility, which is to protect the nest from predators and other competitors. Soldiers have highly developed mandibles as well as many exocrine glands that can secrete multiple defensive substances harmful to predators.^[26]

Normally, only the king and queen termite reproduce; the other castes are all sterile. There are two classes of reproductives: primary reproductives and neotenic reproductives. The primary reproductives class is responsible for colony creation and is characterized by compound eyes, wing marks (spots where wings once were before shedding), and defined sclerotization. Neotenic reproductives can develop from within the colony usually when one of the primary reproductives has died, or can develop in addition to the queen.^[27] neotenic reproductives can experience two different phenotypes, one with wings and one without. If neotenics are winged they will fly away from the parental colony, pair up and form a new colony, but if they are wingless they will remain within the parental colony. The different developmental routes taken by these two morphs are usually dependent upon food availability in the colony, or varying levels of parasitism within the colony.^[27] The caste into which any particular nymph will develop begins to become apparent among the late instars; at this time, potential reproductives will begin to show an increase in the size of the gonadal region.^[26]

Termite colonies may be arboreal, mound-like or subterranean, with primitive termites nesting completely inside enclosed structures such as stumps or logs. Nest construction is largely from the termites' own faecal matter, other materials being chewed vegetable fibre, which makes a weak carton-like but waterproof substance, and soil, which makes a strong substance, but which is subject to erosion by water. Aerial nests are connected to the ground by enclosed passageways; the soft-bodied, blind workers of most species live permanently in their protected environments and do not venture into the open air.^[28] Trinervitermes trinervoides is an exception to this, with workers foraging in small groups on the surface at night, secreting noxious terpenes to deter predators.^[29] The nests are complex structures, and tunnels link them to the foraging areas.^[28] In Africa, termite mounds can be as large as nine meters tall and thirty meters in diameter, producing an area of increased fertility and creating a small hotspot for biodiversity.^[30]^[31]

Related Research Articles

Termites are a group of detritophagous eusocial insects which consume a variety of decaying plant material, generally in the form of wood, leaf litter, and soil humus. They are distinguished by their moniliform antennae and the soft-bodied and often unpigmented worker caste for which they have been commonly termed "white ants"; however, they are not ants, to which they are only distantly related. About 2,972 extant species are currently described, 2,105 of which are members of the family Termitidae.

<span class="mw-page-title-main">Trophallaxis</span> Transfer of food between members of a community through stomodeal or proctodeal means

Trophallaxis is the transfer of food or other fluids among members of a community through mouth-to-mouth (stomodeal) or anus-to-mouth (proctodeal) feeding. Along with nutrients, trophallaxis can involve the transfer of molecules such as pheromones, organisms such as symbionts, and information to serve as a form of communication. Trophallaxis is used by some birds, gray wolves, vampire bats, and is most highly developed in eusocial insects such as ants, wasps, bees, and termites.

Dictyoptera is an insect superorder that includes two extant orders of polyneopterous insects: the order Blattodea and the order Mantodea (mantises). All modern Dictyoptera have short ovipositors and typically lay oothecae. The oldest fossils of Dictyoptera from the Late Carboniferous, referred to as "roachoids" have long ovipositors and did not lay oothecae. The oldest modern oothecae-laying dictyopterans date to the Late Triassic.

Cryptocercus is a genus of Dictyoptera and the sole member of its own family Cryptocercidae. Species are known as wood roaches or brown-hooded cockroaches. These roaches are subsocial, their young requiring considerable parental interaction. They also share wood-digesting gut bacteria types with wood-eating termites, and are therefore seen as evidence of a close genetic relationship, that termites are essentially evolved from social cockroaches.

Corydiidae, previously known as Polyphagidae, is a family of the order Blattodea (cockroaches). Many are known as sand cockroaches. The family is divided into five subfamilies, comprising some 40 genera. One prominent species is the desert cockroach, Arenivaga investigata.

Cockroaches are insects belonging to the order Blattodea (Blattaria). About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known pests.

Reticulitermes flavipes, the eastern subterranean termite, is the most common termite found in North America. These termites are the most economically important wood destroying insects in the United States and are classified as pests. They feed on cellulose material such as the structural wood in buildings, wooden fixtures, paper, books, and cotton. A mature colony can range from 20,000 workers to as high as 5 million workers and the primary queen of the colony lays 5,000 to 10,000 eggs per year to add to this total.

Sociality is the degree to which individuals in an animal population tend to associate in social groups (gregariousness) and form cooperative societies.

"Roachoids", also known as "Roachids", "Blattoids" or Eoblattodea, are members of the stem group of Dictyoptera. They generally resemble cockroaches, but most members, unlike modern dictyopterans, have generally long external ovipositors, and are thought not to have laid ootheca like modern dictyopterans.

Task allocation and partitioning is the way that tasks are chosen, assigned, subdivided, and coordinated within a colony of social insects. Task allocation and partitioning gives rise to the division of labor often observed in social insect colonies, whereby individuals specialize on different tasks within the colony. Communication is closely related to the ability to allocate tasks among individuals within a group. This entry focuses exclusively on social insects. For information on human task allocation and partitioning, see division of labour, task analysis, and workflow.

Eusociality is the highest level of organization of sociality. It is defined by the following characteristics: cooperative brood care, overlapping generations within a colony of adults, and a division of labor into reproductive and non-reproductive groups. The division of labor creates specialized behavioral groups within an animal society, sometimes called castes. Eusociality is distinguished from all other social systems because individuals of at least one caste usually lose the ability to perform behaviors characteristic of individuals in another caste. Eusocial colonies can be viewed as superorganisms.

<i>Therea petiveriana</i> Species of cockroach

Therea petiveriana, variously called the desert cockroach, seven-spotted cockroach, domino cockroach, or Indian domino cockroach, is a species of crepuscular cockroach found in southern India. They are members of a basal group within the cockroaches. This somewhat roundish and contrastingly marked cockroach is mainly found on the ground in scrub forest habitats where they may burrow under leaf litter or loose soil during the heat of the day.

Nasutitermes corniger is a species of arboreal termite that is endemic to the neotropics. It is very closely related to Nasutitermes ephratae. The species has been studied relatively intensively, particularly on Barro Colorado Island, Panama. These studies and others have shown that the termite interacts with many different organisms including a bat that roosts in its nest and various species of ants that cohabit with the termite.

Blattabacterium is a genus of obligate mutualistic endosymbiont bacteria that are believed to inhabit all species of cockroach studied to date, with the exception of the genus Nocticola. The genus' presence in the termite Mastotermes darwiniensis led to speculation, later confirmed, that termites and cockroaches are evolutionarily linked.

Eusociality evolved repeatedly in different orders of animals, notably termites and the Hymenoptera. This 'true sociality' in animals, in which sterile individuals work to further the reproductive success of others, is found in termites, ambrosia beetles, gall-dwelling aphids, thrips, marine sponge-dwelling shrimp, naked mole-rats, and many genera in the insect order Hymenoptera. The fact that eusociality has evolved so often in the Hymenoptera, but remains rare throughout the rest of the animal kingdom, has made its evolution a topic of debate among evolutionary biologists. Eusocial organisms at first appear to behave in stark contrast with simple interpretations of Darwinian evolution: passing on one's genes to the next generation, or fitness, is a central idea in evolutionary biology.

Corydioidea is a superfamily of insects in the order Blattodea containing the cockroaches and termites. It contains two extant families, Corydiidae and Nocticolidae, comprising about fifty genera and two hundred and fifty species, along with the extinct family Liberiblattinidae. Members of this superfamily are found worldwide, mostly in hot, arid habitats.

Social immunity is any antiparasite defence mounted for the benefit of individuals other than the actor. For parasites, the frequent contact, high population density and low genetic variability makes social groups of organisms a promising target for infection: this has driven the evolution of collective and cooperative anti-parasite mechanisms that both prevent the establishment of and reduce the damage of diseases among group members. Social immune mechanisms range from the prophylactic, such as burying beetles smearing their carcasses with antimicrobials or termites fumigating their nests with naphthalene, to the active defenses seen in the imprisoning of parasitic beetles by honeybees or by the miniature 'hitchhiking' leafcutter ants which travel on larger worker's leaves to fight off parasitoid flies. Whilst many specific social immune mechanisms had been studied in relative isolation, it was not until Sylvia Cremer et al.'s 2007 paper "Social Immunity" that the topic was seriously considered. Empirical and theoretical work in social immunity continues to reveal not only new mechanisms of protection but also implications for understanding of the evolution of group living and polyandry.

Alienopteridae is an extinct family of dictyopterans, known from the Early Cretaceous to the Eocene epoch. They are noted for their unusual combination of features not found in other dictyopterans.

Cryptocercus punctulatus, known generally as brown-hooded cockroach, is a species of cockroach in the family Cryptocercidae. Other common names include the woodroach, wingless wood roach, and eastern wood-eating cockroach. It is found in North America.

The social networks of colonies of social insects have been extensively studied as model biological networks.

References

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