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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 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 3 millimeter cockroach nymph female validiblatta fuliginos...
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