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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 in 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 the irrational fear of cockroaches is called katsaridaphobia taxonomy and evolution a turkestan cockroach periplaneta lateralis next to an approximately 108 million year old fossil cockroach from the cretaceous period a 40 to 50 million year old cockroach in baltic amber eocene cockroaches are members of the superorder dictyoptera which includes the termites and mantids 4 a group of insects once thought to be separate from cockroaches currently 4 600 species and over 460 genera are described worldwide 5 6 the name cockroach comes from the spanish word for cockroach cucaracha transformed by 1620s english folk etymology into cock and roach 7 the scientific name derives from the latin blatta an insect that shuns the light which in classical latin was applied not only to cockroaches but also to mantids 8 9 historically the name blattaria was used largely interchangeably with the name blattodea but whilst blattaria was used to refer to true cockroaches exclusively the blattodea also includes the termites the current catalogue of world cockroach species uses the name blattodea for the group 5 another name blattoptera is also sometimes used to refer to extinct cockroach relatives 10 the earliest cockroach like fossils blattopterans or roachoids are from the carboniferous period 320 million years ago 11 12 13 fossil roachoids are considered the common ancestor of both mantises and modern cockroaches and are distinguished from the latter by the presence of a long external ovipositor as the body hind wings and mouthparts are not preserved in fossils frequently the relationship of these roachoids and modern cockroaches remains disputed the earliest definitive fossils of modern crown group cockroaches specifically corydiidae are known from the late jurassic strata of russia and the kimmeridgian aged karabastau formation of kazakhstan 14 15 the evolutionary relationships of the blattodea cockroaches and termites shown in the cladogram are based on inward beccaloni and eggleton 2007 16 the cockroach families anaplectidae 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 17 dictyoptera blattodea blattoidea termitoidae termites termitidae rhinotermitidae kalotermitidae archotermopsidae hodotermitidae mastotermitidae cryptocercoidae cryptocercidae brown hooded cockroaches blattoidae blattidae oriental american and other cockroaches blaberoidea blaberidae giant cockroaches ectobiidae part ectobiidae part corydioidea corydiidae sand cockroaches etc nocticolidae cave cockroaches etc mantodea mantises 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 16 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 18 additional evidence emerged when f a mckittrick 1965 noted similar morphological characteristics between some termites and cockroach nymphs 19 the similarities among these cockroaches and termites have led some scientists to reclassify termites as a single family the termitidae within the order blattodea 16 20 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 21 description domino cockroach therea petiveriana normally found in india most species of cockroach are about the size of a thumbnail but several species are notably larger the world s heaviest cockroach is the australian giant burrowing cockroach macropanesthia rhinoceros which can reach 8 centimetres 3 in in length and weigh up to 35 grams 1 2 oz 22 comparable in size is the central american giant cockroach blaberus giganteus 23 the longest cockroach species is megaloblatta longipennis which can reach 97 mm 3 7 8 in in length and 45 mm 1 3 4 in across 24 a central and south american species megaloblatta blaberoides has the largest wingspan of up to 185 mm 7 1 4 in 25 at the other end of the size scale attaphila cockroaches that live with leaf cutter ants include some of the world s smallest species growing to about 3 5 mm in length 26 head of periplaneta americana 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 27 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 this protects the inner organs and provides attachment to muscles this external exoskeleton 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 these lay as a shield on top of the membranous hind wings which are used in flight all four wings have branching longitudinal veins as well as multiple cross veins 28 the three pairs of legs are sturdy with large coxae and five claws each 28 they are attached to each of the three thoracic segments of these the front legs are the shortest and the hind legs the longest providing the main propulsive power when the insect runs 27 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 29 30 the abdomen has ten segments each having a pair of spiracles for respiration in addition to the spiracles the final segment consists of 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 while females have spermatheca for storing sperm and an ovipositor through which the oothecae are laid 27 distribution and habitat cockroaches are abundant throughout the world and live in a wide range of environments especially in the tropics and subtropics 31 in north america 50 species separated into five families are found throughout the continent 31 450 species are found in australia 32 only about four widespread species are commonly regarded as pests 33 34 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 by doing this cockroaches can remain submerged for up to 40 minutes 35 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 behavior a cockroach soon after ecdysis cockroaches are social insects a large number of species are either gregarious or inclined to aggregate and a slightly smaller number exhibit parental care 37 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 37 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 38 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 39 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 40 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 41 collective decision making 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 42 other mathematical models have been developed to explain aggregation dynamics and conspecific recognition 43 44 45 cooperation and competition are balanced in cockroach group decision making behavior 38 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 seem real to the roaches 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 46 social behavior 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 among males 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 47 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 48 49 the gregarious german and american cockroaches have elaborate social structure chemical signaling and social herd characteristics lihoreau and his fellow researchers stated 38 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 38 there is evidence that a few species of group living roaches in the genera melyroidea and aclavoidea may exhibit a reproductive division of labor which if confirmed would make these the only genuinely eusocial lineage known among roaches in contrast to the subsocial members of the genus cryptocercus 50 sounds some species make a buzzing noise while other cockroaches make a chirping noise gromphadorhina species and archiblatta hoeveni produce sound through the modified spiracles on the fourth abdominal segment in the former species several different hisses are produced including disturbance sounds produced by adults and larger nymphs and aggressive courtship and copulatory sounds produced by adult males 51 henschoutedenia epilamproides has a stridulatory organ between its thorax and abdomen but the purpose of the sound produced is unclear 52 several australian species practice acoustic and vibration behaviour 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 53 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 endosymb...
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