The cockroaches are an ancient group, dating back at least as far as the Carboniferous period, some 320 million years ago. Those early ancestors however lacked the internal ovipositors of modern roaches. Cockroaches are somewhat generalized insects without special adaptations like the sucking mouthparts of aphids and other true bugs; they have chewing mouthparts and are likely among the most primitive of living Neopteran insects. They are common and hardy insects, and can tolerate a wide range of environments from Arctic cold to tropical heat. Tropical cockroaches are often much bigger than temperate species, and, contrary to popular belief, extinct cockroach relatives (Blattoptera) and 'roachoids' such as the Carboniferous Archimylacris and the Permian Apthoroblattina were not as large as the biggest modern species.
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 dirty pests, though the great majority of species are inoffensive and live in a wide range of habitats around the world.
Taxonomy and evolution
Cockroaches are members of the order Blattodea, which includes the termites, a group of insects once thought to be separate from cockroaches. Currently, 4,600 species and over 460 genera are described worldwide. The name "cockroach" comes from the Spanish word for cockroach, cucaracha, transformed by 1620s English folk etymology into "cock" and "roach". The scientific name derives from the Latin blatta, "an insect that shuns the light", which in classical Latin was applied to not only cockroaches, but also mantids.
Historically, the name Blattaria was used largely interchangeably with the name Blattodea, but whilst the former name was used to refer to 'true' cockroaches exclusively, the latter also includes the termites. The current catalogue of world cockroach species uses the name Blattodea for the group. Another name, Blattoptera, is also sometimes used to refer to extinct cockroach relatives. The earliest cockroach-like fossils ("blattopterans" or "roachids") are from the Carboniferous period 320 million years ago, as are fossil roachoid nymphs.
Since the 19th century, scientists believed that cockroaches were an ancient group of insects that had a Devonian origin, according to one hypothesis. Fossil roachoids that lived during that time differ from modern cockroaches in that they had long external ovipositors and are the ancestors of mantises, as well as modern cockroaches. As the body, hind wings and mouthparts are not preserved in fossils frequently, the relationship of these roachoids and modern cockroaches remains disputed. The first fossils of modern cockroaches with internal ovipositors appeared in the early Cretaceous. A recent phylogenetic analysis suggests that cockroaches originated at least in the Jurassic.
The evolutionary relationships of the Blattodea (cockroaches and termites) shown in the cladogram are based on Eggleton, Beccaloni and Inward (2007). 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.
Termites were previously regarded as a separate order Isoptera to cockroaches. However, recent genetic evidence strongly suggests that they evolved directly from 'true' cockroaches, and many authors now place them as an "epifamily" of Blattodea. This evidence supported a hypothesis suggested in 1934 that termites are closely related to the wood-eating cockroaches (genus Cryptocercus). This hypothesis was originally based on similarity of the symbiotic gut flagellates in termites regarded as living fossils and wood-eating cockroaches. Additional evidence emerged when F. A. McKittrick (1965) noted similar morphological characteristics between some termites and cockroach nymphs. The similarities among these cockroaches and termites have led some scientists to reclassify termites as a single family, the Termitidae, within the order Blattodea. Other scientists have taken a more conservative approach, proposing to retain the termites as the Termitoidae, an epifamily within the order. Such a measure preserves the classification of termites at family level and below.
Most species of cockroach are about the size of a thumbnail, but several species are bigger. The world's heaviest cockroach is the Australian giant burrowing cockroach Macropanesthia rhinoceros, which can reach 9 cm (3.5 in) in length and weigh more than 30 g (1.1 oz). Comparable in size is the Central American giant cockroach Blaberus giganteus. The longest cockroach species is , which can reach 97 mm (3.8 in) in length and 45 mm (1.8 in) across. A Central and South American species, , has the largest wingspan of up to 185 mm (7.3 in).
Cockroaches are generalized insects, with few special adaptations, and may be among the most primitive living Neopteran insects. They have a relatively small head and a broad, flattened body, and most species are reddish-brown to dark brown. They have large compound eyes, two ocelli, and long, flexible antennae. The mouthparts are on the underside of the head and include generalized chewing mandibles, salivary glands and various touch and taste receptors.
The body is divided into a thorax of three segments and a ten-segmented abdomen. The external surface has a tough exoskeleton which contains calcium carbonate and protects the inner organs and provides attachment to muscles. It is coated with wax to repel water. The wings are attached to the second and third thoracic segments. The tegmina, or first pair of wings, are tough and protective, lying as a shield on top of the membranous hind wings, which are used in flight. All four wings have branching longitudinal veins, and multiple cross-veins.
The three pairs of legs are sturdy, with large coxae and five claws each. They are attached to each of the three thoracic segments. The front legs are the shortest and the hind legs the longest, providing the main propulsive power when the insect runs. The spines on the legs were earlier considered to be sensory, but observations of the insect's gait on sand and wire meshes have demonstrated that they help in locomotion on difficult terrain. The structures have been used as inspiration for robotic legs.
The abdomen has ten segments, each with a pair of spiracles for respiration. Segment ten bears a pair of cerci, a pair of anal styles, the anus and the external genitalia. Males have an aedeagus through which they secrete sperm during copulation and females have spermatheca for storing sperm and an ovipositor through which the ootheca are laid.
Distribution and habitat
Cockroaches are abundant throughout the world and live in a wide range of environments, especially in the tropics and subtropics. Cockroaches can withstand extremely cold 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. In North America, 50 species separated into five families are found throughout the continent. 450 species are found in Australia. Only about four widespread species are commonly regarded as pests.
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.
Cockroaches are social insects; a large number of species are either gregarious or inclined to aggregate, and a slightly smaller number exhibit parental care. It used to be thought that cockroaches aggregated because they were reacting to environmental cues, but it is now believed that pheromones are involved in these behaviors. Some species secrete these in their feces with gut microbial symbionts being involved, while others use glands located on their mandibles. Pheromones produced by the cuticle may enable cockroaches to distinguish between different populations of cockroach by odor. The behaviors involved have been studied in only a few species, but German cockroaches leave fecal trails with an odor gradient. Other cockroaches follow such trails to discover sources of food and water, and where other cockroaches are hiding. Thus, cockroaches have emergent behavior, in which group or swarm behavior emerges from a simple set of individual interactions.
Daily rhythms may also be regulated by a complex set of hormonal controls of which only a small subset have been understood. In 2005, the role of one of these proteins, pigment dispersing factor (PDF), was isolated and found to be a key mediator in the circadian rhythms of the cockroach.
Pest species adapt readily to a variety of environments, but prefer warm conditions found within buildings. Many tropical species prefer even warmer environments. Cockroaches are mainly nocturnal and run away when exposed to light. An exception to this is the Asian cockroach, which flies mostly at night but is attracted to brightly lit surfaces and pale colors.
Gregarious cockroaches display collective decision-making when choosing food sources. When a sufficient number of individuals (a "quorum") exploits a food source, this signals to newcomer cockroaches that they should stay there longer rather than leave for elsewhere. Other mathematical models have been developed to explain aggregation dynamics and conspecific recognition.
Cooperation and competition are balanced in cockroach group decision-making behavior.
Cockroaches appear to use just two pieces of information to decide where to go, namely how dark it is and how many other cockroaches there are. A study used specially-scented roach-sized robots that appear to the roaches as real to demonstrate that once there are enough insects in a place to form a critical mass, the roaches accepted the collective decision on where to hide, even if this was an unusually lit place.
When reared in isolation, German cockroaches show behavior that is different from behavior when reared in a group. In one study, isolated cockroaches were less likely to leave their shelters and explore, spent less time eating, interacted less with conspecifics when exposed to them, and took longer to recognize receptive females. Because these changes occurred in many contexts, the authors suggested them as constituting a behavioral syndrome. These effects might have been due either to reduced metabolic and developmental rates in isolated individuals or the fact that the isolated individuals had not had a training period to learn about what others were like via their antennae.
Individual American cockroaches appear to have consistently different "personalities" regarding how they seek shelter. In addition, group personality is not simply the sum of individual choices, but reflects conformity and collective decision-making.
The gregarious German and American cockroaches have elaborate social structure, chemical signalling, and "social herd" characteristics. Lihoreau and his fellow researchers stated:
|“||The social biology of domiciliary cockroaches ... can be characterized by a common shelter, overlapping generations, non-closure of groups, equal reproductive potential of group members, an absence of task specialization, high levels of social dependence, central place foraging, social information transfer, kin recognition, and a meta-population structure.||”|
Some species make a hissing noise while other cockroaches make a chirping noise. The Madagascar hissing cockroach produces its sound through the modified spiracles on the fourth abdominal segment. Several different hisses are produced, including disturbance sounds, produced by adults and larger nymphs; and aggressive, courtship and copulatory sounds produced by adult males. has a stridulatory organ between its thorax and abdomen, but the purpose of the sound produced is unclear.
Several Australian species practice acoustic and vibration behavior as an aspect of courtship. They have been observed producing hisses and whistles from air forced through the spiracles. Furthermore, in the presence of a potential mate, some cockroaches tap the substrate in a rhythmic, repetitive manner. Acoustic signals may be of greater prevalence amongst perching species, particularly those that live on low vegetation in Australia's tropics.
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. 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.
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, and current research strongly supports this hypothesis about their relationships. All species studied so far carry the obligate mutualistic endosymbiont bacterium Blattabacterium, with the exception of , an Australian cave-dwelling species without eyes, pigment or wings, which recent genetic studies indicate is a very primitive cockroach. 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. australiensise subsequently lost its symbionts, or alternatively this hypothesis will need to be re-examined.
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.
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.
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.
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. 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.
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.
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, and have survived for as much as four years in the laboratory.
Cockroaches are among the hardiest insects. Some species are capable of remaining active for a month without food and are able to survive on limited resources, such as the glue from the back of postage stamps. Some can go without air for 45 minutes. Japanese cockroach (Periplaneta japonica) nymphs, which hibernate in cold winters, survived twelve hours at −5 °C to −8 °C in laboratory experiments.
Experiments on decapitated specimens of several species of cockroach found a variety of behavioral functionality remained, including shock avoidance and escape behavior, although many insects other than cockroaches are also able to survive decapitation, and popular claims of the longevity of headless cockroaches do not appear to be based on published research. The severed head is able to survive and wave its antennae for several hours, or longer when refrigerated and given nutrients.
It is popularly suggested that cockroaches will "inherit the earth" if humanity destroys itself in a nuclear war. Cockroaches do indeed have a much higher radiation resistance than vertebrates, with the lethal dose perhaps six to 15 times that for humans. However, they are not exceptionally radiation-resistant compared to other insects, such as the fruit fly.
The cockroach's ability to withstand radiation better than human beings can be explained through the cell cycle. Cells are most vulnerable to the effects of radiation when they are dividing. A cockroach's cells divide only once each time it molts, which is weekly at most in a juvenile roach. Since not all cockroaches would be molting at the same time, many would be unaffected by an acute burst of radiation, although lingering radioactive fallout would still be harmful.
Relationship with humans
In research and education
Because of their ease of rearing and resilience, cockroaches have been used as insect models in the laboratory, particularly in the fields of neurobiology, reproductive physiology and social behavior. The cockroach is a convenient insect to study as it is large and simple to raise in a laboratory environment. This makes it suitable both for research and for school and undergraduate biology studies. It can be used in experiments on topics such as learning, sexual pheromones, spatial orientation, aggression, activity rhythms and the biological clock, and behavioral ecology. Research conducted in 2014 suggests that humans fear cockroaches the most, even more than mosquitoes, due to an evolutionary aversion.
The Blattodea include some thirty species of cockroaches associated with humans; these species are atypical of the thousands of species in the order. They feed on human and pet food and can leave an offensive odor. They can passively transport pathogenic microbes on their body surfaces, particularly in environments such as hospitals. Cockroaches are linked with allergic reactions in humans. One of the proteins that trigger allergic reactions is tropomyosin. These allergens are also linked with asthma. About 60% of asthma patients in Chicago are also sensitive to cockroach allergens. Studies similar to this have been done globally and all the results are similar. Cockroaches can live for from a few days to up to a month without food, so just because no cockroaches are visible in a home does not mean they are not there. Approximately 20-48% of homes with no visible sign of cockroaches have detectable cockroach allergens in dust.
Many remedies have been tried in the search for control of the major pest species of cockroaches, which are resilient and fast-breeding. Household chemicals like sodium bicarbonate (baking soda) have been suggested, without evidence for their effectiveness. Garden herbs including bay, catnip, mint, cucumber, and garlic have been proposed as repellents. Poisoned bait containing hydramethylnon or fipronil, and boric acid powder is effective on adults. Baits with egg killers are also quite effective at reducing the cockroach population. Alternatively, insecticides containing deltamethrin or pyrethrin are very effective. In Singapore and Malaysia, taxi drivers use pandan leaves to repel cockroaches in their vehicles.
Few parasites and predators are effective for biological control of cockroaches. Parasitoidal wasps such as Ampulex wasps sting nerve ganglia in the cockroach's thorax, temporarily paralyzing the victim, allowing the wasp to deliver an incapacitating sting into the cockroach's brain. The wasp clips the antennae with its mandibles and drinks some hemolymph before dragging the prey to a burrow, where an egg (rarely two) is laid on it. The wasp larva feeds on the subdued living cockroach. Another wasp which is considered a promising candidate for biological control is the ensign wasp Evania appendigaster which attacks cockroach oothecae to lay a single egg inside. Ongoing research is still developing technologies allowing for mass-rearing these wasps for application releases.
Cockroaches can be trapped in a deep, smooth-walled jar baited with food inside, placed so that cockroaches can reach the opening, for example with a ramp of card or twigs on the outside. An inch or so of water or stale beer (by itself a cockroach attractant) in the jar can be used to drown any insects thus captured. The method works well with the American cockroach, but less so with the German cockroach.
A study conducted by scientists at Purdue University concluded that the most common cockroaches in the US, Australia and Europe were able to develop a “cross resistance” to multiple types of pesticide. This contradicted previous understanding that the animals can develop resistance against one pesticide at a time. The scientists suggested that cockroaches will no longer be easily controlled using a diverse spectrum of chemical pesticides and that a mix of other means, such as traps and better sanitation, will need to be employed.
Although considered disgusting in Western culture, cockroaches are eaten in many places around the world. Whereas household pest cockroaches may carry bacteria and viruses, cockroaches bred under laboratory conditions can be used to prepare nutritious food. In Mexico and Thailand, the heads and legs are removed, and the remainder may be boiled, sautéed, grilled, dried or diced. In China, cockroaches have become popular as medicine and cockroach farming is rising with over 100 farms. The cockroaches are fried twice in a wok of hot oil, which makes them crispy with soft innards that are like cottage cheese. Fried cockroaches are ground and sold as pills for stomach, heart and liver diseases. A cockroach recipe from Formosa (Taiwan) specifies salting and frying cockroaches after removing the head and entrails.
In traditional and homeopathic medicine
In China, cockroaches are raised in large quantities for medicinal purposes.
While a small minority of cockroaches are associated with human habitats and viewed as repugnant by many people, a few species are of conservation concern. The Lord Howe Island wood-feeding cockroach () is listed as endangered by the New South Wales Scientific Committee, but the cockroach may be extinct on Lord Howe Island itself. The introduction of rats, the spread of Rhodes grass (Chloris gayana) and fires are possible reasons for their scarcity. Two species are currently listed as endangered and critically endangered by the IUCN Red List, and Nocticola gerlachi. Both cockroaches have a restricted distribution and are threatened by habitat loss and rising sea levels. Only 600 Delosia ornata adults and 300 nymphs are known to exist, and these are threatened by a hotel development. No action has been taken to save the two cockroach species, but protecting their natural habitats may prevent their extinction. In the former Soviet Union, cockroach populations have been declining at an alarming rate; this may be exaggerated, or the phenomenon may be temporary or cyclic. One species of roach, Simandoa conserfariam, is considered extinct in the wild.
Cockroaches were known and considered repellent but useful in medicines in Classical times. An insect named in Greek "σίλφη" (silphe) has been identified with the cockroach. It is mentioned by Aristotle, saying that it sheds its skin; it is described as foul-smelling in Aristophanes' play Peace; Euenus called it a pest of book collections, being "page-eating, destructive, black-bodied" in his Analect. Virgil named the cockroach "Lucifuga" ("one that avoids light"). Pliny the Elder recorded the use of "Blatta" in various medicines; he describes the insect as disgusting, and as seeking out dark corners to avoid the light. Dioscorides recorded the use of the "Silphe", ground up with oil, as a remedy for earache.
Lafcadio Hearn (1850–1904) asserted that "For tetanus cockroach tea is given. I do not know how many cockroaches go to make up the cup; but I find that faith in this remedy is strong among many of the American population of New Orleans. A poultice of boiled cockroaches is placed over the wound." He adds that cockroaches are eaten, fried with garlic, for indigestion.
Several cockroach species, such as Blaptica dubia, are raised as food for insectivorous pets. A few cockroach species are raised as pets, most commonly the giant Madagascar hissing cockroach, Gromphadorhina portentosa. Whilst the hissing cockroaches may be the most commonly kept species, there are many species that are kept by cockroach enthusiasts; there is even a specialist society: the Blattodea Culture Group (BCG), which was a thriving organisation for about 15 years although now appears to be dormant. The BCG provided a source of literature for people interested in rearing cockroaches which was otherwise limited to either scientific papers, or general insect books, or books covering a variety of exotic pets; in the absence of an inclusive book one member published Introduction to Rearing Cockroaches which still appears to be the only book dedicated to rearing cockroaches.
Cockroaches have been used for space tests. A cockroach given the name Nadezhda was sent into space by Russian scientists as part of a Foton-M mission, during which she mated, and later became the first terrestrial animal to produce offspring that had been conceived in space.
Because of their long association with humans, cockroaches are frequently referred to in popular culture. In Western culture, cockroaches are often depicted as dirty pests. In a 1750–1752 journal, Peter Osbeck noted that cockroaches were frequently seen and found their way to the bakeries, after the sailing ship Gothenburg ran aground and was destroyed by rocks.
Donald Harington's satirical novel (Harcourt, 1989) imagines a community of "roosterroaches" in a mythical Ozark town where the insects are named after their human counterparts. Madonna has famously quoted, "I am a survivor. I am like a cockroach, you just can't get rid of me." An urban legend maintains that cockroaches are immortal.
- Beccaloni, G. W. (2014). "Cockroach Species File Online. Version 5.0".
- "Blattodea (Cockroaches & Termites)". CSIRO Entomology. Retrieved 21 November 2015.
- Harper, Douglas. "Cockroach". Online Etymology Dictionary.
- Gordh, G.; Headrick, D. H. (2009). A Dictionary of Entomology (2nd ed.). Wallingford: CABI. p. 200. ISBN 978-1-84593-542-9.
- Lewis, Charlton T.; Short, Charles. "Blatta". Perseus Digital Library. Tufts University. Retrieved 26 October 2015.
- Grimaldi, D. (1997). "A fossil mantis (Insecta, Mantodea) in Cretaceous amber of New Jersey: with comments on the early history of the Dictyoptera". American Museum Novitates. 3204: 1–11.
- Garwood, R.; Sutton, M. (2010). "X-ray micro-tomography of Carboniferous stem-Dictyoptera: new insights into early insects". Biology Letters. 6 (5): 699–702. doi:10.1098/rsbl.2010.0199. PMC 2936155. PMID 20392720.
- Grimaldi, David; Engel, Michael S. (2005). Evolution of the Insects. Cambridge University Press. p. 1. ISBN 978-0-521-82149-0.
- Garwood, R.; Ross, A.; Sotty, D.; Chabard, D.; Charbonnier, S.; Sutton, M.; Withers, P.J.; Butler, R.J. (2012). "Tomographic Reconstruction of Neopterous Carboniferous Insect Nymphs". PLoS ONE. 7 (9): e45779. Bibcode:2012PLoSO...745779G. doi:10.1371/journal.pone.0045779. PMC 3458060. PMID 23049858.
- Legendre, F.; Nel, A.; Svenson, G. J.; Robillard, T.; Pellens, R.; Grandcolas, P. (2015). "Phylogeny of Dictyoptera: Dating the Origin of Cockroaches, Praying Mantises and Termites with Molecular Data and Controlled Fossil Evidence". PLoS ONE. 10 (7): e0130127. Bibcode:2015PLoSO..1030127L. doi:10.1371/journal.pone.0130127. PMC 4511787. PMID 26200914.
- Inward, D.; Beccaloni, G.; Eggleton, P. (2007). "Death of an order: a comprehensive molecular phylogenetic study confirms that termites are eusocial cockroaches". Biology Letters. 3 (3): 331–335. doi:10.1098/rsbl.2007.0102. PMC 2464702. PMID 17412673.
- Beccaloni, G.; Eggleton, P. (2013). Order Blattodea. In: Zhang; Z.-Q.; Survey of Taxonomic Richness (eds.). "Animal Biodiversity: An Outline of Higher-level Classification" (PDF). Zootaxa. 3703 (1): 46–48. doi:10.11646/zootaxa.3703.1.10.
- Cleveland, L.R.; Hall, S.K.; Sanders, E.P.; Collier, J. (1934). "The Wood-Feeding Roach Cryptocercus, Its Protozoa, and the Symbiosis between Protozoa and Roach". Memoirs of the American Academy of Arts and Sciences. 17 (2): 185–382. doi:10.1093/aesa/28.2.216.
- McKittrick, F.A. (1965). "A contribution to the understanding of cockroach-termite affinities". Annals of the Entomological Society of America. 58 (1): 18–22. doi:10.1093/aesa/58.1.18. PMID 5834489.
- Eggleton, Paul; Beccaloni, George; Inward, Daegan (2007). "Response to Lo et al". Biology Letters. 3 (5): 564–565. doi:10.1098/rsbl.2007.0367. PMC 2391203.
- Lo, Nathan; Engel, Michael S.; Cameron, Stephen; Nalepa, Christine A.; Tokuda, Gaku; Grimaldi, David; Kitade, Osamu; Krishna, Kumar; Klass, Klaus-Dieter; Maekawa, Kiyoto; Miura, Toru; Thompson, Graham J. (2007). "Comment. Save Isoptera: A comment on Inward et al". Biology Letters. 3 (5): 562–563. doi:10.1098/rsbl.2007.0264. PMC 2391185. PMID 17698448.
- "Pet facts: giant burrowing cockroaches". Australian Broadcasting Corporation. Archived from the original on 18 February 2014. Retrieved 3 December 2005.
- Huang, C.Y.; Sabree, Z.L.; Moran, N.A. (2012). "Genome Sequence of Blattabacterium sp. Strain BGIGA, Endosymbiont of the Blaberus giganteus Cockroach". Journal of Bacteriology. 194 (16): 4450–4451. doi:10.1128/jb.00789-12. PMC 3416254. PMID 22843586.
- Guinness World Records. "Guinness World Records: World's Largest Cockroach".
- "Natural History Museum: Cockroaches hit the shelves". Natural History Museum. May 2006. Archived from the original on 19 August 2006. Retrieved 23 November 2015.
- "Diversity of Life: Cockroach anatomy". Biology4ISC. Archived from the original on 8 December 2015. Retrieved 8 November 2015.
- Hoell, H. V.; Doyen, J. T.; Purcell, A. H. (1998). Introduction to Insect Biology and Diversity (2nd ed.). Oxford University Press. pp. 362–364. ISBN 978-0-19-510033-4.
- Ritzmann, Roy E.; Quinn, Roger D.; Fischer, Martin S. (2004). "Convergent evolution and locomotion through complex terrain by insects, vertebrates and robots" (PDF). Arthropod Structure & Development. 33 (3): 361–379. doi:10.1016/j.asd.2004.05.001. PMID 18089044.
- Spagna, J.C.; Goldman, D.I.; Lin, P-C.; Koditschek, D.E.; Full, Robert J. (2007). "Distributed mechanical feedback control of rapid running on challenging terrain". Bioinspiration & Biomimetics. 2 (1): 9–18. doi:10.1088/1748-3182/2/1/002. PMID 17671322.
- Meyer, J. "Blattodea". General Entomology. University of North Carolina. Retrieved 9 November 2015.
- Mohs, K.; McGee, I. (2007). Animal planet: the most extreme bugs (1st ed.). John Wiley & Sons. p. 35. ISBN 978-0-7879-8663-6.
- "Cockroaches: Order Blattodea". Australian Museum. January 13, 2012. Retrieved November 10, 2015.
- Valles, S. M.; Koehler, P. G.; Brenner, R. J. (1999). "Comparative insecticide susceptibility and detoxification enzyme activities among pestiferous blattodea" (PDF). Comp Infibous Biochem Physiol C Pharmacol Toxicol Endocrinol. 124 (3): 227–232. doi:10.1016/S0742-8413(99)00076-6. PMID 10661713.
- Schal, C; Hamilton, R. L. (1990). "Integrated suppression of synanthropic cockroaches" (PDF). Annu. Rev. Entomol. 35: 521–551. doi:10.1146/annurev.en.35.010190.002513. PMID 2405773.
- Bell, William J.; Roth, Louis M.; Nalepa, Christine A. (2007). Cockroaches: Ecology, Behavior, and Natural History. JHU Press. pp. 55–58. ISBN 978-0-8018-8616-4.
- Costa, James T. (2006). The Other Insect Societies. Harvard University Press. p. 148. ISBN 978-0-674-02163-1.
- Lihoreau, M.; Costa, J.T.; Rivault, C. (November 2012). "The social biology of domiciliary cockroaches: colony structure, kin recognition and collective decisions". Insectes Sociaux. 59 (4): 445–452. doi:10.1007/s00040-012-0234-x.
- Hamasaka, Yasutaka; Mohrherr, C. J.; Predel, R.; Wegener, C. (22 December 2005). "Chronobiological analysis and mass spectrometric characterization of pigment-dispersing factor in the cockroach Leucophaea maderae". The Journal of Insect Science. 5 (43): 43. doi:10.1093/jis/5.1.43. PMC 1615250. PMID 17119625.
- Rust, M. K. (2007). "Cockroaches". University of California Integrated Pest Management Program. University of California. Retrieved 24 November 2015.
- Richman, Dina L. (1 June 2014). "Asian cockroach". featured Creatures. University of Florida. Retrieved 4 November 2015.
- Lihoreau, Mathieu; Deneubourg, Jean-Louis; Rivault, Colette (2010). "Collective foraging decision in a gregarious insect". Behavioral Ecology and Sociobiology. 64 (10): 1577–1587. doi:10.1007/s00265-010-0971-7.
- Ame, Jean-Marc; Rivault, Colette; Deneubourg, Jean-Louis (Oct 2004). "Cockroach aggregation based on strain odour recognition". Animal Behaviour. 68 (4): 793–801. doi:10.1016/j.anbehav.2004.01.009.
- Jeanson, Raphael; Rivault, Colette; Deneubourg, Jean-Louis; Blanco, Stephane; Fournier, Richard; Jost, Christian; Theraulaz, Guy (Jan 2005). "Self-organized aggregation in cockroaches". Animal Behaviour. 69 (1): 169–180. doi:10.1016/j.anbehav.2004.02.009.
- Lemonick, Michael D. (15 November 2007). "Robotic Roaches Do the Trick". Time Magazine. Retrieved 24 November 2015.
- Lihoreau, Mathieu; Brepson, Loïc; Rivault, Colette (2009). "The weight of the clan: Even in insects, social isolation can induce a behavioural syndrome". Behavioural Processes. 82 (1): 81–84. doi:10.1016/j.beproc.2009.03.008. PMID 19615616.
- Planas-Sitjà, Isaac; Deneubourg, Jean-Louis; Gibon, Céline; Sempo, Grégory (2015). "Group personality during collective decision-making: a multi-level approach" (PDF). Proc. R. Soc. B. 282 (1802): 20142515. doi:10.1098/rspb.2014.2515. PMC 4344149. PMID 25652834.
- Morell, Virginia (3 February 2015). "Even cockroaches have personalities". Science. doi:10.1126/science.aaa7797. Retrieved 19 February 2015.
- Nelson, Margaret C. (1979). "Sound production in the cockroach, Gromphadorhina portentosa: The sound-producing apparatus". Journal of Comparative Physiology. 132 (1): 27–38. doi:10.1007/BF00617729.
- Guthrie, D.M. (1966). "Sound production and reception in a cockroach". Journal of Experimental Biology. 45: 321–328.
- Rentz, David (2014). A Guide to the Cockroaches of Australia. CSIRO Publishing. ISBN 978-0-643-10320-7.
- Bell, William J.; Adiyodi, K. G. (1981). American Cockroach. Springer. p. 4. ISBN 978-0-412-16140-7.
- Slaytor, Michael (1992). "Cellulose digestion in termites and cockroaches: What role do symbionts play?". Comparative Biochemistry and Physiology B. 103 (4): 775–784. doi:10.1016/0305-0491(92)90194-V.
- Eggleton, P. (2001). "Termites and trees: a review of recent advances in termite phylogenetics". Insectes Sociaux. 48 (3): 187–193. doi:10.1007/PL00001766.
- Lo, N.; Bandi, C.; Watanabe, H.; Nalepa, C.; Beninati, T. (2003). "Evidence for Cocladogenesis Between Diverse Dictyopteran Lineages and Their Intracellular Endosymbionts". Molecular Biology and Evolution. 20 (6): 907–13. doi:10.1093/molbev/msg097. PMID 12716997.
- Leung, Chee Chee (22 March 2007). "Cave may hold missing link". The Age. Retrieved 24 November 2015.
- Lo, N.; Beninati, T; Stone, F.; Walker, J.; Sacchi, L. (2007). "Cockroaches that lack Blattabacterium endosymbionts: The phylogenetically divergent genus Nocticola". Biology Letters. 3 (3): 327–30. doi:10.1098/rsbl.2006.0614. PMC 2464682. PMID 17376757.
- "The Cockroach FAQ". University of Massachusetts. Retrieved 24 November 2015.
- Mullen, Gary; Durden, Lance, eds. (2002). Medical and Veterinary Entomology. Amsterdam: Academic Press. p. 32. ISBN 978-0-12-510451-7.
- Tanaka, K.; Tanaka, S. (1997). "Winter Survival and Freeze Tolerance in a Northern Cockroach, Periplaneta japonica (Blattidae: Dictyoptera)". Zoological Science. 14 (5): 849–853. doi:10.2108/zsj.14.849.
- Berenbaum, May (30 September 2009). The Earwig's Tail: A Modern Bestiary of Multi-legged Legends. Harvard University Press. pp. 53–54. ISBN 978-0-674-03540-9.
- Choi, Charles (15 March 2007). "Fact or fiction?: a cockroach can live without its head". Scientific American. Scientific American. Retrieved 27 December 2013.
- Kruszelnicki, Karl S. (23 February 2006). "Cockroaches and Radiation". ABC Science. Retrieved 24 November 2015.
- Bell, W. J. (2012) . The Laboratory Cockroach: Experiments in cockroach anatomy, physiology and behavior. Springer. ISBN 978-94-011-9726-7.
- "Cockroaches: The insect we are programmed to fear". BBC. 18 September 2014.
- Gullan, P. J.; Cranston, P. S. (2014). The Insects: An Outline of Entomology. Wiley. p. 508. ISBN 978-1-118-84615-5.
- Brenner, R.J.; Koehler, P.; Patterson, R.S. (1987). "Health Implications of Cockroach Infestations". Infestations in Med. 4 (8): 349–355.
- Rivault, C.; Cloarec, A.; Guyader, A. Le (1993). "Bacterial load of cockroaches in relation to urban environment". Epidemiology and Infection. 110 (2): 317–325. doi:10.1017/S0950268800068254. PMC 2272268. PMID 8472775.
- Elgderi, R.M.; Ghenghesh, K.S.; Berbash, N. (2006). "Carriage by the German cockroach (Blattella germanica) of multiple-antibiotic-resistant bacteria that are potentially pathogenic to humans, in hospitals and households in Tripoli, Libya". Ann Trop Med Parasitol. 100 (1): 55–62. doi:10.1179/136485906X78463. PMID 16417714.
- Bernton, H.S.; Brown, H. (1964). "Insect Allergy Preliminary Studies of the Cockroach". J. Allergy. 35 (506–513): 506–13. doi:10.1016/0021-8707(64)90082-6. PMID 14226309.
- Kutrup, B (2003). "Cockroach Infestation in Some Hospitals in Trabzon, Turkey" (PDF). Turk. J. Zool. 27: 73–77.
- Santos, A. B.; Chapman, M. D.; Aalberse, R. C.; Vailes, L. D.; Ferriani, V. P.; Oliver, C.; Rizzo, M. C.; Naspitz, C. K.; et al. (1999). "Cockroach allergens and asthma in Brazil: identification of tropomyosin as a major allergen with potential cross-reactivity with mite and shrimp allergens". The Journal of Allergy and Clinical Immunology. 104 (2): 329–337. doi:10.1016/S0091-6749(99)70375-1. PMID 10452753.
- Kang, B.; Vellody, D.; Homburger, H.; Yunginger, J. W. (1979). "Cockroach cause of allergic asthma. Its specificity and immunologic profile". The Journal of Allergy and Clinical Immunology. 63 (2): 80–86. doi:10.1016/0091-6749(79)90196-9. PMID 83332.
- Eggleston, P.A.; Arruda, L.K. (2001). "Ecology and elimination of cockroaches and allergens in the home". Journal of Allergy and Clinical Immunology. 107 (3): S422–S429. doi:10.1067/mai.2001.113671. PMID 11242603.
- Helmer, Hendrik (10 November 2017). "Experience: A cockroach got stuck in my ear". Theguardian.com. Retrieved 10 November 2017.
- "Doctors Pull Cockroach From Australian Man's Ear (PHOTO)". Huffingtonpost.com. 10 January 2014. Retrieved 10 November 2017 – via Huff Post.
- "Doctor pulls 2cm long cockroach from Hendrik Helmer's ear". Independent.co.uk. 10 January 2014. Retrieved 10 November 2017.
- "Man has 2cm cockroach removed from ear". Abc.net.au. 10 January 2014. Retrieved 10 November 2017.
- Malhotra, Priya (23 August 2011). "The Common Ailment of Cockroach-in-Ear". Huffingtonpost.com. Retrieved 10 November 2017.
- "Best Home Remedies To Kill And Control Cockroaches – Baking Soda". HRT.whw1.com. Retrieved 2015-06-20.
- "Best Home Remedies To Kill And Control Cockroaches – Catnip". HRT.whw1.com. Retrieved 20 June 2015.
- "Cockroaches". Alamance County Department of Environmental Health. Archived from the original on March 12, 2009. Retrieved 11 May 2008.
- Li J. and Ho S.H. Pandan leaves (Pandanus amaryllifolius Roxb.) As A Natural Cockroach Repellent. Proceedings of the 9th National Undergraduate Research Opportunities Programme (13 September 2003).
- Paterson Fox, Eduardo Gonçalves; Bressan-Nascimento, Suzete; Eizemberg, Roberto (September 2009). "Notes on the Biology and Behaviour of the Jewel Wasp, Ampulex compressa (Fabricius, 1781) (Hymenoptera; Ampulicidae), in the Laboratory, Including First Record of Gregarious Reproduction". Entomological News. 120 (4): 430–437. doi:10.3157/021.120.0412.
- Fox, Eduardo G P; BUYS, SANDOR CRISTIANO; MALLET, JACE-NIR REIS DOS SANTOS; BRESSAN-NASCIMENTO, SUZETE (3 August 2006). "On the morphology of the juvenile stages of Ampulex compressa (Fabricius 1781) (Hymenoptera, Ampulicidae)" (PDF). Zootaxa. 1279 (1): 43. doi:10.11646/zootaxa.1279.1.2. hdl:11449/69026.
- Piper, Ross (2007). Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals. Greenwood Press.
- Fox, Eduardo G P (10 May 2011). "Evania appendigaster Development". YouTube.
- Fox, Eduardo Gonçalves Paterson; Solis, Daniel Russ; Rossi, Mônica Lanzoni; Eizemberg, Roberto; Taveira, Luiz Pilize; Bressan-Nascimento, Suzete (June 2012). "The preimaginal stages of the ensign wasp Evania appendigaster (Hymenoptera, Evaniidae), a cockroach egg predator". Invertebrate Biology. 131 (2): 133–143. doi:10.1111/j.1744-7410.2012.00261.x.
- Bressan-Nascimento, S.; Oliveira, D.M.P.; Fox, E.G.P. (December 2008). "Thermal requirements for the embryonic development of Periplaneta americana (L.) (Dictyoptera: Blattidae) with potential application in mass-rearing of egg parasitoids". Biological Control. 47 (3): 268–272. doi:10.1016/j.biocontrol.2008.09.001.
- Bressan-Nascimento, S.; Fox, E.G.P.; Pilizi, L.G.T. (February 2010). "Effects of different temperatures on the life history of Evania appendigaster L. (Hymenoptera: Evaniidae), a solitary oothecal parasitoid of Periplaneta americana L. (Dictyoptera: Blattidae)". Biological Control. 52 (2): 104–109. doi:10.1016/j.biocontrol.2009.10.005.
- Herms, William Brodbeck (1915). Medical and Veterinary Entomology. MacMillan. p. 44.
- Wahlquist, Calla (2019-07-03). "Cockroaches could soon be almost impossible to kill with pesticides". The Guardian. ISSN 0261-3077. Retrieved 2019-07-04.
- Wahlquist, Calla (2019-07-03). "Cockroaches could soon be almost impossible to kill with pesticides". The Guardian. ISSN 0261-3077. Retrieved 2019-07-04.
- Marion Copeland . Cockroach (Pages 86 to 88). Published by Reaktion Books
- Ronald L. Taylor, Barbara J. Carter . Entertaining with Insects: Or, The Original Guide to Insect Cookery. Published by Woodbridge Press Publishing Company.
- David George Gordon . The Eat-a-bug Cookbook (Page 78). Published by Ten Speed Press.
- Demick, Barbara (15 October 2013). "Cockroach farms multiplying in China". Los Angeles Times. Retrieved 20 April 2018.
- Malcolm Moore . How to eat a cockroach: a Telegraph guide. Published by Washington Post.
- . How cockroaches could save lives. Published by BBC News.
- David McKenzie . Eating cockroaches in China: Healing and delicious?. Published by CNN.
- Richard Schweid . The Cockroach Papers: A Compendium of History and Lore (Page 69). Published by University of Chicago Press.
- Chen, Stephen (2018-04-19). "A giant indoor farm in China is breeding 6 billion cockroaches a year. Here's why". South China Morning Post. Retrieved 2018-04-20.
- Kmietowicz, Z. (1997-05-17). "Cockroaches blamed for high asthma rates". BMJ. 314 (7092): i. doi:10.1136/bmj.314.7092.1433i.
- "Lord Howe Island wood-feeding cockroach – endangered species listing". Office of Environment and Heritage. Government of New South Wales. 2011. Retrieved 10 November 2015.
- Gerlach, J. (2012). "Delosia ornata". IUCN Red List of Threatened Species. IUCN. 2012: e.T199490A2593559. doi:10.2305/IUCN.UK.2012.RLTS.T199490A2593559.en.
- Gerlach, J. (2012). "Nocticola gerlachi". IUCN Red List of Threatened Species. IUCN. 2012: e.T199508A2595807. doi:10.2305/IUCN.UK.2012.RLTS.T199508A2595807.en.
- "Исчезновение тараканов на Белгородчине не связано с радиацией". Bel.ru (in Russian). 4 December 2006. Archived from the original on 16 October 2013. Retrieved 24 November 2015.
- Anthon, Charles (1843). Smith, William (ed.). A Dictionary of Greek and Roman Antiquities (3rd American ed.). New York – Cincinnati – Chicago: American Book Company. p. 161.
- Lockyer, Norman (1871). Nature. Macmillan Journals. p. 27.
- Hearn, Lafcadio; Starr, S. Frederick (2001). Inventing New Orleans: Writings of Lafcadio Hearn. University Press of Mississippi. pp. 68–69. ISBN 978-1-57806-353-6.
- Wu, Hao; Appel, Arthur G.; Hu, Xing Ping (2013). "Instar Determination of Blaptica dubia (Blattodea: Blaberidae) Using Gaussian Mixture Models". Annals of the Entomological Society of America. 106 (3): 323–328. doi:10.1603/AN12131. ISSN 0013-8746.
- Mulder, Phil. "Madagascar Hissing Cockroaches: Information and Care" (PDF). Oklahoma 4-H Youth Development. Oklahoma State University. Retrieved 31 October 2013.
- "Blattodea Culture Group". Blattodea-culture-group.org. Retrieved 10 November 2017.
- Bragg, P.E. (1997) An Introduction to Rearing Cockroaches. P.E. Bragg, Ilkeston.
- ""Hope" the Russian cockroach gives birth to first space babies". RIA Novosti. 23 October 2007. Retrieved 24 November 2015.
- Berle, D. (2007). "Graded Exposure Therapy for Long-Standing Disgust-Related Cockroach Avoidance in an Older Male". Clinical Case Studies. 6 (4): 339–347. doi:10.1177/1534650106288965.
- Botella, C.M.; Juan, M. C.; Baños, R. M.; Alcañiz, M.; Guillén, V.; Rey, B. (2005). "Mixing Realities? An Application of Augmented Reality for the Treatment of Cockroach Phobia". CyberPsychology & Behavior. 8 (2): 162–171. doi:10.1089/cpb.2005.8.162. PMID 15938656.
- Klausnitzer, B. (1987). Insects: their biology and cultural history. New York: Universe Books. p. 42. ISBN 978-0-87663-666-4.
- Madonna. "Madonna". Thinkexist.com. Retrieved 29 April 2012.
I am a survivor. I am like a cockroach, you just can't get rid of me
- "Cockroaches are not radiation-proof and most are not pests". BBC Earth. Retrieved 1 June 2018.