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men 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 fuliginosa with ootheca captured adult and baby roaches around ootheca empty ootheca american cockroach oothecae parthenogenesis when female american cockroaches periplaneta americana are housed in groups this close association promotes parthenogenic reproduction 61 oothecae a type of egg mass are produced asexually 61 the parthenogenetic process by which eggs are produced in p americana is automixis 62 during automixis meiosis occurs but instead of giving rise to haploid gametes as ordinarily occurs diploid gametes are produced probably by terminal fusion that can then develop into female cockroaches hardiness 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 63 some can go without air for 45 minutes japanese cockroach crescispina japonica nymphs which hibernate in cold winters have survived twelve hours at 5 to 8 c 23 to 18 f in laboratory experiments 64 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 popular claims of the longevity of headless cockroaches do not appear to be based on published research 65 66 the severed head is able to survive and wave its antennae for several hours or longer when refrigerated and given nutrients 66 it is popularly suggested that cockroaches will inherit the earth if humanity destroys itself in a nuclear war while cockroaches do indeed have a much higher radiation resistance than vertebrates with a lethal dose perhaps six to 15 times that for humans they are not exceptionally radiation resistant compared to other insects such as the fruit fly 67 the cockroach s ability to withstand radiation has been explained through the cell cycle cells are most vulnerable to the effects of radiation while they are dividing a cockroach s cells divide only once each molting cycle which is weekly for the juvenile german cockroach 68 since not all cockroaches would be molting at the same time many would be unaffected by an acute burst of radiation although prolonged radiation would still be harmful 60 relationship with humans cockroaches in research periplaneta americana in an electrophysiology experiment cockroach climbing on a wall in japan 2018 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...
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