Cockroach

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Cockroach

Common household roaches A) German cockroach B) American cockroach C) Australian cockroach D&E) Oriental cockroach ( & )

Scientific classification Kingdom: Phylum: Animalia Arthropoda

Subphylum: Class: Subclass: Infraclass: Superorder: Order:

Hexapodaa Insecta Pterygota Neoptera Dictyoptera Blattaria Families

Blaberidae Blattellidae Blattidae Cryptocercidae Polyphagidae Nocticolidae Tryonicidae Lamproblattidae Cockroaches are insects of the order Blattaria or Blattodea, of which about 30 species out of 4,500 total are associated with human habitations. About four species are well known as pests.[1][2] Among the best-known pest species are the American cockroach, Periplaneta americana, which is about 30 millimetres (1.2 in) long, the German cockroach, Blattella germanica, about 15 millimetres (0.59 in) long, the Asian cockroach, Blattella asahinai, also about 15 millimetres (0.59 in) in length, and the Oriental cockroach, Blatta orientalis, about 25 millimetres (0.98 in). Tropical cockroaches are often much bigger, and extinct cockroach relatives and 'roachoids' such as the Carboniferous Archimylacris and the Permian Apthoroblattina were not as large as the biggest modern species.

Contents
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1 Etymology 2 Notable species 3 Evolutionary history and relationships 4 Behavior 5 Description o 5.1 Eggs and egg capsules o 5.2 Sounds 6 Physiology o 6.1 Digestive tract o 6.2 Tracheae and breathing o 6.3 Reproduction o 6.4 Hardiness 7 Role as pests 8 Cultural references 9 References o 9.1 Notes o 9.2 Bibliography 10 External links

[edit] Etymology
The name cockroach comes from the Spanish word cucaracha, "chafer", "beetle", from cuca, "kind of caterpillar." The scientific name derives from the Latinized Greek name for the insect (Doric Greek: , bltta; Ionic and Attic Greek: , bltte' The English form cockroach is a folk etymology reanalysis of the Spanish word into meaningful native parts, although cock referred to a rooster and a roach is a type of fish.

[edit] Notable species

Blattella germanica, German cockroach Blaptica dubia, South American/Peruvian Dubia cockroach Blatta orientalis, Oriental cockroach Blattella asahinai, Asian cockroach Blaberus craniifer, true death's head cockroach Blaberus discoidalis, discoid cockroach or false death's head Eurycotis floridana, Florida woods cockroach Gromphadorhina portentosa, Madagascar hissing cockroach laxta granicollis, Bark cockroach Parcoblatta pennsylvanica, Pennsylvania woods cockroach Periplaneta americana, American cockroach Periplaneta australasiae, Australian cockroach Periplaneta brunnea, black Mississippi cockroach Periplaneta fuliginosa, smokybrown cockroach Pycnoscelus surinamensis, Surinam cockroach

Supella longipalpa, brown-banded cockroach

[edit] Evolutionary history and relationships

A 40-50 million year old cockroach in Baltic amber

Blattaria

Polyphagidae Cryptocercidae Blattidae Blattellidae Blaberidae phylogeny of the

A proposed families.[3]

Mantodea, Isoptera, and Blattaria are usually combined by entomologists into a higher group called Dictyoptera. Current evidence strongly suggests that termites evolved directly from true cockroaches, and many authors now consider termites to be an epifamily of cockroaches, [4][5] as Blattaria excluding Isoptera is not a monophyletic group.[6] Historically, the name Blattaria has been used largely interchangeably with the name Blattodea, and this name is used for the order by the current world catalogue, the Blattodea Species File Online. Another name, Blattoptera has come into use for this same paraphyletic group.[7] These earliest cockroach-like fossils ("Blattopterans" or "roachids") are from the Carboniferous period between 354295 million years ago. However, these fossils differ from modern cockroaches in having long external ovipositors and are the ancestors of mantises as well as modern cockroaches. The first fossils of modern cockroaches with internal ovipositors appear in the early Cretaceous.

[edit] Behavior

A cockroach soon after ecdysis.

A bush cockroach (Ellipsidion australe).

Cockroaches live in a wide range of environments around the world. Pest species of cockroaches adapt readily to a variety of environments, but prefer warm conditions found within buildings. Many tropical species prefer even warmer environments and do not fare well in the average household. The spines on the legs were earlier considered to be sensory, but observations of their locomotion 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.[8][9] Cockroaches leave chemical trails in their feces as well as emitting airborne pheromones for swarming and mating. These chemical trails transmit bacteria on surfaces.[citation needed] Other cockroaches will follow these trails to discover sources of food and water, and also discover

where other cockroaches are hiding. Thus, cockroaches can exhibit emergent behavior,[10] 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.[11] Research has shown that group-based decision-making is responsible for complex behavior such as resource allocation. In a study where 50 cockroaches were placed in a dish with three shelters with a capacity for 40 insects in each, the insects arranged themselves in two shelters with 25 insects in each, leaving the third shelter empty. When the capacity of the shelters was increased to more than 50 insects per shelter, all of the cockroaches arranged themselves in one shelter. Researchers found a balance between cooperation and competition exists in the group decisionmaking behavior found in cockroaches. The models used in this research can also explain the group dynamics of other insects and animals.[10] Cockroaches are mainly nocturnal and will run away when exposed to light. A peculiar exception is the Asian cockroach, which is attracted to light. Another study tested the hypothesis that cockroaches use just two pieces of information to decide where to go under those conditions: how dark it is and how many other cockroaches there are. The study conducted by Jos Halloy and colleagues at the Free University of Brussels and other European institutions created a set of tiny robots that appear to the roaches as other roaches and can thus alter the roaches' perception of critical mass. The robots were also specially scented so that they would be accepted by the real roaches.[12] Additionally, researchers at Tohoku University engaged in a classical conditioning experiment with cockroaches and discovered that the insects were able to associate the scent of vanilla and peppermint with a sugar treat.[13]

[edit] Description
Cockroaches are generally rather large insects. Most species are about the size of a thumbnail, but several species are bigger. The world's heaviest cockroach is the Australian giant burrowing cockroach, which can reach 9 centimetres (3.5 in) in length and weigh more than 30 grams (1.1 oz). Comparable in size is the Central American giant cockroach Blaberus giganteus, which grows to a similar length but is not as heavy. Cockroaches have a broad, flattened body and a relatively small head. They are generalized insects, with few special adaptations, and may be among the most primitive living neopteran insects. The mouthparts are on the underside of the head and include generalised chewing mandibles. They have large compound eyes, two ocelli, and long, flexible, antennae. The first pair of wings (the tegmina) are tough and protective, lying as a shield on top of the membranous hind wings. All four wings have branching longitudinal veins, and multiple cross-

veins. The legs are sturdy, with large coxae and five claws each. The abdomen has ten segments and several cerci.[14]
[edit] Eggs and egg capsules

Female Periplaneta fuliginosa with ootheca.

Hatched Ootheca

Female cockroaches are sometimes seen carrying egg cases on the end of their abdomen; the egg case of the German cockroach holds about 30 to 40 long, thin eggs, packed like frankfurters in the case called an ootheca. The eggs hatch 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 many to claim the existence of albino cockroaches. A female German cockroach carries an egg capsule containing around 40 eggs. She drops the capsule prior to hatching, though live births do occur in rare instances. Development from eggs to adults takes 3 to 4 months. Cockroaches live up to a year. 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 cockroach, however, can produce an extremely high number of eggs in a lifetime; in some cases a female needs to be impregnated only once to be able to lay eggs for the rest of her life.

[edit] Sounds

Aside from the famous hissing noise, some cockroaches (including a species in Florida) will make a chirping noise.[15]

[edit] Physiology
[edit] Digestive tract

Cockroaches are most common in tropical and subtropical climates. Some species are in close association with human dwellings and widely found around garbage or in the kitchen. Cockroaches are generally omnivorous with the exception of the wood-eating species such as Cryptocercus; these roaches are incapable of digesting cellulose themselves, but have symbiotic relationships with various protozoans and bacteria that digest the cellulose, allowing them to extract the nutrients. The similarity of these symbionts in the genus Cryptocercus to those in termites are such that it has been suggested that they are more closely related to termites than to other cockroaches,[16] and current research strongly supports this hypothesis of relationships.[17] All species studied so far carry the obligate mutualistic endosymbiont bacterium Blattabacterium, with the exception of Nocticola australiensise, an Australian cave dwelling species without eyes, pigment or wings, and which recent genetic studies indicates are very primitive cockroaches.[18][19]
[edit] Tracheae and breathing

Nymph of a cockroach, 3 mm in length.

Cockroaches, like all insects, breathe through a system of tubes called tracheae. The tracheae of insects are attached to the spiracles, excluding the head. Thus cockroaches, like all insects, are not dependent on the mouth and windpipe to breathe. The valves open when the CO2 level in the insect rises to a high level; then the CO2 diffuses out of the tracheae to the outside and fresh O2 diffuses in. Unlike in vertebrates that depend on blood for transporting O2 and CO2, the tracheal system brings the air directly to cells, the tracheal tubes branching continually like a tree until their finest divisions, tracheoles, are associated with each cell, allowing gaseous oxygen to

dissolve in the cytoplasm lying across the fine cuticle lining of the tracheole. CO2 diffuses out of the cell into the tracheole. While cockroaches do not have lungs and thus do not actively breathe in the vertebrate lung manner, in some very large species the body musculature may contract rhythmically to forcibly move air out and in the spiracles; this may be considered a form of breathing.[20]
[edit] 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.[14] 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 bodies, with or without an egg-case, until they hatch. At least one genus, Diploptera is fully viviparous.[14] Cockroach nymphs are generally similar to the adults, except for undeveloped wings and genitalia. Development is generally slow, and may take anywhere from a few months to over a year. The adults are also long-lived, and have been recorded as surviving for four years in the laboratory.[14]
[edit] Hardiness

Roach Leg

Cockroaches are among the hardiest insects on the planet. Some species are capable of remaining active for a month without food and are able to survive on limited resources like the glue from the back of postage stamps.[21] Some can go without air for 45 minutes. In one experiment, cockroaches were able to recover from being submerged underwater for half an hour.[22]

Ootheca of Periplaneta americana; Florianpolis, SC, Brazil.

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 6 to 15 times that for humans. However, they are not exceptionally radiation-resistant compared to other insects, such as the fruit fly.[23] 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, but lingering radioactive fallout would still be harmful.[24]

[edit] Role as pests

Cockroaches are one of the most commonly noted household pest insects.[25] They feed on human and pet food, and can leave an offensive odor.[26] They can also passively transport microbes on their body surfaces including those that are potentially dangerous to humans, particularly in environments such as hospitals.[27][28] Cockroaches have been shown to be linked with allergic reactions in humans.[29][30] One of the proteins that triggers allergic reactions has been identified as tropomyosin.[31] These allergens have also been found to be linked with asthma.[32] General preventive measures against household pests include keeping all food stored away in sealed containers, using garbage cans with a tight lid, frequent cleaning in the kitchen, and regular vacuuming. Any water leaks, such as dripping taps, should also be repaired. It is also helpful to seal off any entry points, such as holes around baseboards, in between kitchen cabinets, pipes, doors, and windows with some steel wool or copper mesh and some cement, putty or silicone caulk.

Diatomaceous earth applied as a fine powder works very well to eliminate cockroaches as long as it remains in place and dry. Diatomaceous earth is harmless to humans and feels like talcum powder. Most insects, including bed bugs, are vulnerable to it. Some cockroaches have been known to live up to three months without food and a month without water. Frequently living outdoors, although preferring warm climates and considered "cold intolerant," they are resilient enough to survive occasional freezing temperatures. This makes them difficult to eradicate once they have infested an area. There are numerous parasites and predators of cockroaches, but few of them have proven to be highly effective for biological control of pest species. Wasps in the family Evaniidae are perhaps the most effective insect predators, as they attack the egg cases, and wasps in the family Ampulicidae are predators on adult and nymphal cockroaches (e.g., Ampulex compressa). The house centipede is probably the most effective control agent of cockroaches, though many homeowners find the centipedes themselves objectionable. Ampulex wasps sting the roach more than once and in a specific way. The first sting is directed at nerve ganglia in the cockroach's thorax; temporarily paralyzing the victim for 25 minutes, which is more than enough time for the wasp to deliver a second sting. The second sting is directed into a region of the cockroach's brain that controls the escape reflex, among other things.[33] When the cockroach has recovered from the first sting, it makes no attempt to flee. The wasp clips the antennae with its mandibles and drinks some of the hemolymph before walking backwards and dragging the roach by its clipped antennae to a burrow, where an egg will be laid upon it. The wasp larva feeds on the subdued, living cockroach. Bait stations, gels containing hydramethylnon or fipronil, as well as boric acid powder, are toxic to cockroaches.[34] Baits with egg killers are also quite effective at reducing the cockroach population. Additionally, pest control products containing deltamethrin or pyrethrin are very effective.[34] In Singapore and Malaysia, taxi drivers use Pandan leaves as a cockroach repellent in their vehicles.[35] An inexpensive roach trap can easily be made from a deep smooth-walled jar with some roach food inside, placed with the top of the jar touching a wall or with sticks leading up to the top, so that the roaches can reach the opening. Once inside, they cannot climb back out. An inch or so of water or stale beer (by itself a roach attractant) will ensure they drown. The method works well with the American cockroach but less so with the German cockroach.[36] A bit of Vaseline can be smeared on the inside of the jar to enhance slipperiness. The method is sometimes called the "Vegas roach trap" after it was popularized by a Las Vegas-based TV station. This version of the trap uses coffee grounds and water.[37] Some of the earliest writings about cockroaches encouraged their use as medicine. Pedanius Dioscorides (1st century), Abu Hanifa ad-Dainuri (9th century), and Kamal al-Din al-Damiri (14th century ) all offered medicines that either suggest grinding them up with oil or boiling them.

Cockroaches are but one of many insects whose nutritional excellence has been underestimated or scorned, according to The Eat-A-Bug Cookbook, which also reports (on p. 66) that Lafcadio Hearn found that many New Orleanians had great faith in a remedy of boiled cockroach tea. Pest control is cited as one of the reasons for reduced populations of cockroaches in ex-USSR countries.[38]

[edit] Cultural references

Main article: Cultural references to cockroaches

Because of their long, persistent association with humans, cockroaches are frequently referred to in art, literature, folk tales and theater and film. In Western culture, cockroaches are often depicted as vile and dirty pests. Their size, long antennae, shiny appearance and spiny legs make them disgusting to many humans, sometimes even to the point of phobic responses.[39][40] This is borne out in many depictions of cockroaches, from political versions of the song La Cucaracha where political opponents are compared to cockroaches, through the 1982 movie Creepshow and TV shows such as the X-Files, to the Hutu extremists' reference to the Tutsi minority as cockroaches during the Rwandan Genocide in 1994 and the controversial cartoons published in the "Iran weekly magazine" in 2006 which implied a comparison between Iranian Azeris and cockroaches. In the movie Men in Black a giant alien cockroach is shown as a predator who eats a farmer and then uses his skin to disguise itself as a human being. In Oliver Twist, the children, Mr. Bumble, and Widow Corney sing about feeding Oliver cockroaches in a canister. Awardwinning computer and video game series Fallout takes place in a post-nuclear war universe, in which enlarged, irradiated cockroaches are present as early enemies. This is a nod to the notion of their nuclear fortitude. Also, in Franz Kafka's The Metamorphosis, a man, Gregor, is transformed overnight into a monstrous insect with cockroach-like features. He views himself as repulsive in his new identity. Not all depictions of cockroaches are purely negative, however. Twilight of the Cockroaches depicts the extermination of cockroaches as a holocaust, and presents a happy ending as the pregnant lead cockroach, Naomi, escapes to mother many generations. In the Pixar film Wall-E, a cockroach that has survived all humanity is the best friend of the lead character (a robot), and waits patiently on him to return. The same cockroach survives getting squished twice. In the film Joe's Apartment, the cockroaches help the titular hero, and the narrator of the book archy and mehitabel is a sympathetic cockroach. In the book Revolt of the Cockroach People, an autobiographical novel by Oscar Zeta Acosta, cockroaches are used as a metaphor for oppressed and downtrodden minorities in US society in the 1960s and 70s. The image of cockroaches as resilient also leads people to compare themselves to cockroaches. Madonna has famously quoted, "I am a survivor. I am like a cockroach, you just can't get rid of me."[41] "Cockroach", or some variant of it is also used as a nickname, for example Boxing coach Freddie Roach, who was nicknamed La Cucaracha (The Cockroach) when he was still competing as a fighter. The album The Lonesome Crowded West by rock group Modest Mouse features a song with the title and lyric "Doin' The Cockroach". In the Netherlands 'Zaza the cockroach' becomes a buddy of the boy called Pluk in a popular Dutch book for children: 'Pluk van de Petteflet', written by Annie M.G. Schmidt. In Suzanne Collins's Underland Chronicles series, giant cockroaches are allies of

humans in the Underland, and they and a toddler named Margaret (a.k.a. Boots, or "the princess," as the cockroaches call her) love each other.

Bee
From Wikipedia, the free encyclopedia Jump to: navigation, search For other uses, see Bee (disambiguation).

Bees

Osmia ribifloris

Scientific classification Kingdom: Phylum: Class: Order: Animalia Arthropoda Insecta Hymenoptera

Suborder: Superfamily: (unranked):

Apocrita Apoidea Anthophila Families

Andrenidae Apidae Colletidae Dasypodaidae Halictidae Megachilidae Meganomiidae Melittidae Stenotritidae

Synonyms

Apiformes Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila. There are nearly 20,000 known species of bees in seven to nine recognized families,[1] though many are undescribed and the actual number is probably higher. They are found on every continent except Antarctica, in every habitat on the planet that contains insect-pollinated flowering plants. Bees are adapted for feeding on nectar and pollen, the former primarily as an energy source and the latter primarily for protein and other nutrients. Most pollen is used as food for larvae. Bees have a long proboscis (a complex "tongue") that enables them to obtain the nectar from flowers. They have antennae almost universally made up of 13 segments in males and 12 in females, as is typical for the superfamily. Bees all have two pairs of wings, the hind pair being the smaller of the two; in a very few species, one sex or caste has relatively short wings that make flight difficult or impossible, but none are wingless. The smallest bee is Trigona minima, a stingless bee whose workers are about 2.1 mm (5/64") long. The largest bee in the world is Megachile pluto, a leafcutter bee whose females can attain a

length of 39 mm (1.5"). Members of the family Halictidae, or sweat bees, are the most common type of bee in the Northern Hemisphere, though they are small and often mistaken for wasps or flies. The best-known bee species is the European honey bee, which, as its name suggests, produces honey, as do a few other types of bee. Human management of this species is known as beekeeping or apiculture. Bees are the favorite meal of Merops apiaster, the bee-eater bird. Other common predators are kingbirds, mockingbirds, beewolves, and dragonflies.

Contents
[hide]

1 Pollination o 1.1 Pollinator decline 2 Evolution 3 Eusocial and semisocial bees o 3.1 Bumblebees o 3.2 Stingless bees o 3.3 Honey bees o 3.4 Africanized honey bee 4 Solitary and communal bees 5 Cleptoparasitic bees 6 Nocturnal bees 7 Flight 8 Bees and humans 9 See also 10 External links 11 References 12 Further reading

Pollination
See also: List of crop plants pollinated by bees

Bees play an important role in pollinating flowering plants, and are the major type of pollinator in ecosystems that contain flowering plants. Bees either focus on gathering nectar or on gathering pollen depending on demand, especially in social species. Bees gathering nectar may accomplish pollination, but bees that are deliberately gathering pollen are more efficient pollinators. It is estimated that one third of the human food supply depends on insect pollination, most of which is accomplished by bees, especially the domesticated European honey bee. Contract pollination has overtaken the role of honey production for beekeepers in many countries. Monoculture and the massive decline of many bee species (both wild and domesticated) have increasingly caused honey bee keepers to become migratory so that bees can be concentrated in seasonally varying high-demand areas of pollination.

Honey bee (Apis mellifera) collecting pollen

Most bees are fuzzy and carry an electrostatic charge, which aids in the adherence of pollen. Female bees periodically stop foraging and groom themselves to pack the pollen into the scopa, which is on the legs in most bees, and on the ventral abdomen on others, and modified into specialized pollen baskets on the legs of honey bees and their relatives. Many bees are opportunistic foragers, and will gather pollen from a variety of plants, while others are oligolectic, gathering pollen from only one or a few types of plant. A small number of plants produce nutritious floral oils rather than pollen, which are gathered and used by oligolectic bees. One small subgroup of stingless bees, called "vulture bees," is specialized to feed on carrion, and these are the only bees that do not use plant products as food. Pollen and nectar are usually combined together to form a "provision mass", which is often soupy, but can be firm. It is formed into various shapes (typically spheroid), and stored in a small chamber (a "cell"), with the egg deposited on the mass. The cell is typically sealed after the egg is laid, and the adult and larva never interact directly (a system called "mass provisioning"). In New Zealand scientists discovered that three genera of native bees have evolved to open flower buds of the native mistletoe Peraxilla tetrapetala. The buds cannot open themselves but are visited by birds such as the tui and bellbird which twist the top of the ripe bud. That action releases a mechanism which causes the petals to suddenly spring open, giving access to the nectar and pollen. However, when observing the native bees in the Canterbury province in the South Island, the scientists were astonished to see the bees biting the top off the buds, then pushing with their legs, occasionally popping open the buds to allow the bees to harvest the nectar and pollen, and therefore aid in the pollination of the mistletoe which is in decline in New Zealand. Nowhere else in the world have bees demonstrated ability to open explosive birdadapted flowers.[2] Visiting flowers can be a dangerous occupation. Many assassin bugs and crab spiders hide in flowers to capture unwary bees. Other bees are lost to birds in flight. Insecticides used on blooming plants kill many bees, both by direct poisoning and by contamination of their food supply. A honey bee queen may lay 2000 eggs per day during spring buildup, but she also must lay 1000 to 1500 eggs per day during the foraging season, mostly to replace daily casualties, most of which are workers dying of old age. Among solitary and primitively social bees,

however, lifetime reproduction is among the lowest of all insects, as it is common for females of such species to produce fewer than 25 offspring. The population value of bees depends partly on the individual efficiency of the bees, but also on the population itself. Thus while bumblebees have been found to be about ten times more efficient pollinators on cucurbits, the total efficiency of a colony of honey bees is much greater due to greater numbers. Likewise during early spring orchard blossoms, bumblebee populations are limited to only a few queens, and thus are not significant pollinators of early fruit.
Pollinator decline

Morphology of a female honey bee

From 1972 to 2006, there was a dramatic reduction in the number of feral honey bees in the US, which are now almost absent.[3] At the same time there was a significant though somewhat gradual decline in the number of colonies maintained by beekeepers. This decline includes the cumulative losses from all factors, such as urbanization, pesticide use, tracheal and Varroa mites, and commercial beekeepers' retiring and going out of business. However, in late 2006 and early 2007 the rate of attrition reached new proportions, and the term colony collapse disorder was coined to describe the sudden disappearances.[4] After several years of research and concern, a team of scientists headed by Jerry Bromenshenk published a paper in October 2010 saying that a new DNA-based virus, invertebrate iridescent virus or IIV6, and the fungus Nosema ceranae were found in every killed colony the group studied. In their study they found that neither agent alone seemed deadly, but a combination of the virus and Nosema ceraneae was always 100% fatal. Bromenshenk said it is not yet clear whether one condition weakens the bees enough to be finished off by the second, or whether they somehow compound the others destructive power. "They're co-factors, thats all we can say at the moment. Theyre both present in all these collapsed colonies."[5][6][7] Investigations into the phenomenon had occurred amidst great concern over the nature and extent of the losses.[8] In 2009 some reports from the US suggested that 1/3 of the honey bee colonies did not survive the winter,[9] though normal winter losses are known to be around 25%.[10] Apart from colony collapse disorder, many of the losses outside the US have also been attributed to other causes. Pesticides used to treat seeds, such as Clothianidin and Imidacloprid, have been

considered prime suspects.[11] Other species of bees such as mason bees are increasingly cultured and used to meet the agricultural pollination need.[12] Native pollinators include bumblebees and solitary bees, which often survive in refuges in wild areas away from agricultural spraying, but may still be poisoned in massive spray programs for mosquitoes, gypsy moths, or other insect pests. Although pesticide use remains a concern, the major problem for wild pollinator populations is the loss of the flower-rich habitat on which they depend for food.[citation needed] Throughout the northern hemisphere, the last 70 or so years have seen an intensification of agricultural systems, which has decreased the abundance and diversity of wild flowers.[citation needed] Legislation such as the UK's Bees Act 1980 is designed to stop the decline of bees.[13]

Evolution
Bees, like ants, are a specialized form of wasp. The ancestors of bees were wasps in the family Crabronidae, and therefore predators of other insects. The switch from insect prey to pollen may have resulted from the consumption of prey insects which were flower visitors and were partially covered with pollen when they were fed to the wasp larvae. This same evolutionary scenario has also occurred within the vespoid wasps, where the group known as "pollen wasps" also evolved from predatory ancestors. Up until recently, the oldest non-compression bee fossil had been Cretotrigona prisca in New Jersey amber and of Cretaceous age, a meliponine. A recently reported bee fossil, of the genus Melittosphex, is considered "an extinct lineage of pollencollecting Apoidea sister to the modern bees", and dates from the early Cretaceous (~100 mya).[14] Derived features of its morphology ("apomorphies") place it clearly within the bees, but it retains two unmodified ancestral traits ("plesiomorphies") of the legs (two mid-tibial spurs, and a slender hind basitarsus), indicative of its transitional status. The earliest animal-pollinated flowers were pollinated by insects such as beetles, so the syndrome of insect pollination was well established before bees first appeared. The novelty is that bees are specialized as pollination agents, with behavioral and physical modifications that specifically enhance pollination, and are generally more efficient at the task than any other pollinating insect such as beetles, flies, butterflies and pollen wasps. The appearance of such floral specialists is believed to have driven the adaptive radiation of the angiosperms, and, in turn, the bees themselves. Among living bee groups, the "short-tongued" bee family Colletidae has traditionally been considered the most "primitive", and sister taxon to the remainder of the bees. In the 21st century, however, some researchers have claimed that the Dasypodaidae is the basal group, the short, wasp-like mouthparts of colletids being the result of convergent evolution, rather than indicative of a plesiomorphic condition.[1] This subject is still under debate, and the phylogenetic relationships among bee families are poorly understood.
See also: characteristics of common wasps and bees

Eusocial and semisocial bees

A honey bee swarm

Bees may be solitary or may live in various types of communities. The most advanced of these are eusocial colonies[15] found among the honey bees, bumblebees, and stingless bees. Sociality, of several different types, is believed to have evolved separately many times within the bees. In some species, groups of cohabiting females may be sisters, and if there is a division of labor within the group, then they are considered semisocial. If, in addition to a division of labor, the group consists of a mother and her daughters, then the group is called eusocial. The mother is considered the "queen" and the daughters are "workers". These castes may be purely behavioral alternatives, in which case the system is considered "primitively eusocial" (similar to many paper wasps), and if the castes are morphologically discrete, then the system is "highly eusocial". There are many more species of primitively eusocial bees than highly eusocial bees, but they have rarely been studied. The biology of most such species is almost completely unknown. The vast majority are in the family Halictidae, or "sweat bees". Colonies are typically small, with a dozen or fewer workers, on average. The only physical difference between queens and workers is average size, if they differ at all. Most species have a single season colony cycle, even in the tropics, and only mated females (future queens, or "gynes") hibernate (called diapause). A few species have long active seasons and attain colony sizes in the hundreds. The orchid bees include a number of primitively eusocial species with similar biology. Certain species of allodapine bees (relatives of carpenter bees) also have primitively eusocial colonies, with unusual levels of interaction between the adult bees and the developing brood. This is "progressive provisioning"; a larva's food is supplied gradually as it develops. This system is also seen in honey bees and some bumblebees.

Highly eusocial bees live in colonies. Each colony has a single queen, many workers and, at certain stages in the colony cycle, drones. When humans provide the nest, it is called a hive. Honey bee hives can contain up to 40,000 bees at their annual peak, which occurs in the spring, but usually have fewer.
Bumblebees Main article: Bumblebee

Bumblebees (Bombus terrestris, Bombus pratorum, et al.) are eusocial in a manner quite similar to the eusocial Vespidae such as hornets. The queen initiates a nest on her own (unlike queens of honey bees and stingless bees which start nests via swarms in the company of a large worker force). Bumblebee colonies typically have from 50 to 200 bees at peak population, which occurs in mid to late summer. Nest architecture is simple, limited by the size of the nest cavity (preexisting), and colonies are rarely perennial. Bumblebee queens sometimes seek winter safety in honey bee hives, where they are sometimes found dead in the spring by beekeepers, presumably stung to death by the honey bees. It is unknown whether any survive winter in such an environment. Bumblebees are one of the more important wild pollinators, but have declined significantly in recent decades. In the UK, 2 species have become nationally extinct during the last 75 years while others have been placed on the UK Biodiversity Action Plan as priority species in recognition of the need for conservation action. In 2006 a new charity, the Bumblebee Conservation Trust, was established in order to coordinate efforts to conserve remaining populations through conservation and education.
Stingless bees Main article: Stingless bee

Stingless bees are very diverse in behavior, but all are highly eusocial. They practise mass provisioning, complex nest architecture, and perennial colonies.
Honey bees

A European honey bee extracts nectar from an Aster flower

Main article: Honey bee

The true honey bees (genus Apis) have arguably the most complex social behavior among the bees. The European (or Western) honey bee, Apis mellifera, is the best known bee species and one of the best known of all insects.
Africanized honey bee Main article: Africanized bee

Africanized bees, also called killer bees, are a hybrid strain of Apis mellifera derived from experiments by Warwick Estevam Kerr to cross European and African honey bees. Several queen bees escaped from his laboratory in South America and have spread throughout the Americas. Africanized honey bees are more defensive than European honey bees.

Solitary and communal bees

Most other bees, including familiar species of bee such as the Eastern carpenter bee (Xylocopa virginica), alfalfa leafcutter bee (Megachile rotundata), orchard mason bee (Osmia lignaria) and the hornfaced bee (Osmia cornifrons) are solitary in the sense that every female is fertile, and typically inhabits a nest she constructs herself. There are no worker bees for these species. Solitary bees typically produce neither honey nor beeswax. They are immune from acarine and Varroa mites, but have their own unique parasites, pests and diseases (see also diseases of the honey bee). Solitary bees are important pollinators, and pollen is gathered for provisioning the nest with food for their brood. Often it is mixed with nectar to form a paste-like consistency. Some solitary bees have very advanced types of pollen-carrying structures on their bodies. A very few species of solitary bees are being increasingly cultured for commercial pollination.

A solitary bee, Anthidium florentinum (family Megachilidae), visiting Lantana

Solitary bees are often oligoleges, in that they only gather pollen from one or a few species/genera of plants (unlike honey bees and bumblebees which are generalists). No known

bees are nectar specialists; many oligolectic bees will visit multiple plants for nectar, but there are no bees which visit only one plant for nectar while also gathering pollen from many different sources. Specialist pollinators also include bee species which gather floral oils instead of pollen, and male orchid bees, which gather aromatic compounds from orchids (one of the only cases where male bees are effective pollinators). In a very few cases only one species of bee can effectively pollinate a plant species, and some plants are endangered at least in part because their pollinator is dying off. There is, however, a pronounced tendency for oligolectic bees to be associated with common, widespread plants which are visited by multiple pollinators (e.g., there are some 40 oligoleges associated with creosote bush in the US desert southwest,[16] and a similar pattern is seen in sunflowers, asters, mesquite, etc.) Solitary bees create nests in hollow reeds or twigs, holes in wood, or, most commonly, in tunnels in the ground. The female typically creates a compartment (a "cell") with an egg and some provisions for the resulting larva, then seals it off. A nest may consist of numerous cells. When the nest is in wood, usually the last (those closer to the entrance) contain eggs that will become males. The adult does not provide care for the brood once the egg is laid, and usually dies after making one or more nests. The males typically emerge first and are ready for mating when the females emerge. Providing nest boxes for solitary bees is increasingly popular for gardeners. Solitary bees are either stingless or very unlikely to sting (only in self defense, if ever). While solitary females each make individual nests, some species are gregarious, preferring to make nests near others of the same species, giving the appearance to the casual observer that they are social. Large groups of solitary bee nests are called aggregations, to distinguish them from colonies. In some species, multiple females share a common nest, but each makes and provisions her own cells independently. This type of group is called "communal" and is not uncommon. The primary advantage appears to be that a nest entrance is easier to defend from predators and parasites when there are multiple females using that same entrance on a regular basis.

Cleptoparasitic bees

Bombus vestalis, a cuckoo bee parasite of the bumblebee Bombus terrestris

Cleptoparasitic bees, commonly called "cuckoo bees" because their behavior is similar to cuckoo birds, occur in several bee families, though the name is technically best applied to the apid subfamily Nomadinae. Females of these bees lack pollen collecting structures (the scopa) and do not construct their own nests. They typically enter the nests of pollen collecting species, and lay their eggs in cells provisioned by the host bee. When the cuckoo bee larva hatches it consumes the host larva's pollen ball, and if the female cleptoparasite has not already done so, kills and eats the host larva. In a few cases where the hosts are social species, the cleptoparasite remains in the host nest and lays many eggs, sometimes even killing the host queen and replacing her. Many cleptoparasitic bees are closely related to, and resemble, their hosts in looks and size, (i.e., the Bombus subgenus Psithyrus, which are parasitic bumblebees that infiltrate nests of species in other subgenera of Bombus). This common pattern gave rise to the ecological principle known as "Emery's Rule". Others parasitize bees in different families, like Townsendiella, a nomadine apid, one species of which is a cleptoparasite of the dasypodaid genus Hesperapis, while the other species in the same genus attack halictid bees.

Nocturnal bees
Four bee families (Andrenidae, Colletidae, Halictidae, and Apidae) contain some species that are crepuscular (these may be either the vespertine or matinal type). These bees have greatly enlarged ocelli, which are extremely sensitive to light and dark, though incapable of forming images. Many are pollinators of flowers that themselves are crepuscular, such as evening primroses, and some live in desert habitats where daytime temperatures are extremely high.

Flight

Bee in mid air flight carrying pollen in pollen basket

In his 1934 French book Le vol des insectes, M. Magnan wrote that he and a M. Saint-Lague had applied the equations of air resistance to bumblebees and found that their flight could not be explained by fixed-wing calculations, but that "One shouldn't be surprised that the results of the calculations don't square with reality".[17] This has led to a common misconception that bees "violate aerodynamic theory", but in fact it merely confirms that bees do not engage in fixedwing flight, and that their flight is explained by other mechanics, such as those used by helicopters.[18]

In 1996 Charlie Ellington at Cambridge University showed that vortices created by many insects wings and non-linear effects were a vital source of lift;[19] vortices and non-linear phenomena are notoriously difficult areas of hydrodynamics, which has made for slow progress in theoretical understanding of insect flight. In 2005, Michael Dickinson and his Caltech colleagues studied honey bee flight with the assistance of high-speed cinematography[20] and a giant robotic mock-up of a bee wing.[21] Their analysis revealed that sufficient lift was generated by "the unconventional combination of short, choppy wing strokes, a rapid rotation of the wing as it flops over and reverses direction, and a very fast wing-beat frequency". Wing-beat frequency normally increases as size decreases, but as the bee's wing beat covers such a small arc, it flaps approximately 230 times per second, faster than a fruitfly (200 times per second) which is 80 times smaller.[22]

Bees and humans

Bee larvae as food in Java

Bees figure prominently in mythology and have been used by political theorists as a model for human society. Journalist Bee Wilson states that the image of a community of honey bees "occurs from ancient to modern times, in Aristotle and Plato; in Virgil and Seneca; in Erasmus and Shakespeare; Tolstoy, as well as by social theorists Bernard Mandeville and Karl Marx."[23] Despite the honey bee's painful sting and the stereotype of insects as pests, bees are generally held in high regard. This is most likely due to their usefulness as pollinators and as producers of honey, their social nature, and their reputation for diligence. Bees are one of the few insects regularly used on advertisements, being used to illustrate honey and foods made with honey (such as Honey Nut Cheerios). In ancient Egypt, the bee was seen to symbolize the lands of Lower Egypt, with the Pharaoh being referred to as "He of Sedge and Bee" (the sedge representing Upper Egypt). In North America, yellowjackets and hornets, especially when encountered as flying pests, are often misidentified as bees, despite numerous differences between them. Although a bee sting can be deadly to those with allergies, virtually all bee species are nonaggressive if undisturbed and many cannot sting at all. Humans are often a greater danger to

bees, as bees can be affected or even harmed by encounters with toxic chemicals in the environment (see also bees and toxic chemicals). In Indonesia bee larvae are eaten as a companion to rice, after being mixed with shredded coconut "meat", wrapped in banana leaves, and steamed.

See also
Wikiquote has a collection of quotations related to: Bees

Wikimedia Commons has media related to: Apoidea

Apiology Bee and wasp stings Honey bee life cycle International Union for the Study of Social Insects Pesticide toxicity to bees Schmidt Sting Pain Index Starr sting pain scale Proboscis extension reflex

External links

All Living Things Images, identification guides, and maps of bees Bee Genera of the World Solitary Bees & Things Solitary Bees in British gardens Scientists identify the oldest known bee, a 100 million-year-old specimen preserved in amber North American species of bees at Bugguide Native Bees of North America

References
1. ^ Danforth BN, Sipes S, Fang J, Brady SG (October 2006). "The history of early bee diversification based on five genes plus morphology". Proc. Natl. Acad. Sci. U.S.A. 103 (41): 1511823. doi:10.1073/pnas.0604033103. PMC 1586180. PMID 17015826. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1586180. ^ "Native Bees With New Tricks". New Zealand Science Monthly. http://nzsm.spis.co.nz/article1699.htm. [dead link] Retrieved 16 June 2009. ^ Watanabe, M. (1994). "Pollination worries rise as honey bees decline". Science 265 (5176): 1170. doi:10.1126/science.265.5176.1170. PMID 17787573. ^ "Honey Bee Die-Off Alarms Beekeepers, Crop Growers and Researchers". Pennsylvania State University College of Agricultural Sciences. 2007-01-29. http://www.aginfo.psu.edu/News/07Jan/HoneyBees.htm. ^ "Scientists and Soldiers Solve a Bee Mystery"
ab

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^ "What a scientist didn't tell the New York Times about his study on bee deaths" ^ Jerry J. Bromenshenk, Colin B. Henderson, Charles H. Wick, Michael F. Stanford, Alan W. Zulich, Rabih E. Jabbour, Samir V. Deshpande, Patrick E. McCubbin, Robert A. Seccomb, Phillip M. Welch, Trevor Williams, David R. Firth, Evan Skowronski, Margaret M. Lehmann, Shan L. Bilimoria, Joanna Gress, Kevin W. Wanner, Robert A. Cramer Jr (06 Oct 2010). "Iridovirus and Microsporidian Linked to Honey Bee Colony Decline" (in English). PLoS ONE. http://www.plosone.org/article/info:doi/10.1371/journal.pone.0013181. ^ "Honey bees in US facing extinction", Telegraph 14 March 2007 ^ Fears for crops as shock figures from America show scale of bee catastrophe. Alison Benjamin. The Observer, 2 May 2010. ^ "Beekeepers Report Continued Heavy Losses From Colony Collapse Disorder". Sciencedaily.com. 200805-12. http://www.sciencedaily.com/releases/2008/05/080509111955.htm. Retrieved 2010-06-22. ^ German Consumer Protection Agency Bulletin June 9, 2008 ^ Mason bee from Everything.About. Retrieved 10 March 2009. ^ "Statute Law UK". Crown Copyright. March 1980. http://www.statutelaw.gov.uk/content.aspx?LegType=All+Legislation&title=Bees+Act&searchEnacted=0& extentMatchOnly=0&confersPower=0&blanketAmendment=0&sortAlpha=0&TYPE=QS&PageNumber=1& NavFrom=0&parentActiveTextDocId=1507973&ActiveTextDocId=1507973&filesize=17676. Retrieved 2009-01-20. ^ Poinar GO, Danforth BN (October 2006). "A fossil bee from Early Cretaceous Burmese amber". Science 314 (5799): 614. doi:10.1126/science.1134103. PMID 17068254. ^ Engel, Michael S. (2001-02-13). "Monophyly and Extensive Extinction of Advanced Eusocial Bees: Insights from an Unexpected Eocene Diversity". PNAS (National Academy of Sciences) 98 (4): 16611664. doi:10.1073/pnas.041600198. JSTOR 3054932. PMC 29313. PMID 11172007. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=29313. ^ Hurd, P.D. Jr., Linsley, E.G. (1975). "The principal Larrea bees of the southwestern United States.". Smithsonian Contributions to Zoology 193: 174. ^ Ingram, Jay The Barmaid's Brain, Aurum Press, 2001, pp. 91-92. ^ Cecil Adams (1990-05-04). "Is it aerodynamically impossible for bumblebees to fly?". The Straight Dope. http://www.straightdope.com/columns/read/1076/is-it-aerodynamically-impossible-for-bumblebees-tofly. Retrieved 2009-03-07. ^ Secrets of bee flight revealed, Phillips, Helen. 28 November 2005. Retrieved 2007-12-28. ^ http://www.newscientist.com/data/images/ns/av/dn8382.avi ^ Deciphering the Mystery of Bee Flight Caltech Media Relations. Nov. 29, 2005. Retrieved 2007, 4-7. ^ Douglas L. Altshuler, William B. Dickson, Jason T. Vance, Stephen P. Roberts, and Michael H. Dickinson (2005). "Short-amplitude high-frequency wing strokes determine the aerodynamics of honeybee flight". Proc Natl Acad Sci U S A. 102 (50): 1821318218. doi:10.1073/pnas.0506590102. PMC 1312389. PMID 16330767. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1312389. ^ Wilson, Bee (2004). The Hive: The Story Of The Honeybee. London, Great Britain: John Murray (publisher). ISBN 0719565987.

Further reading

O'Toole, Christopher, and Raw, Anthony. (1991). Bees of the World. Facts on File. Michener, Charles D. (2007). The Bees of the World, second edition. Baltimore: Johns Hopkins.

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Categories: Hymenoptera | Bees | Pollinators Hidden categories: All articles with dead external links | Articles with dead external links from September 2010 | Wikipedia indefinitely semi-protected pages | Articles with 'species' microformats | All articles with unsourced statements | Articles with unsourced statements from May 2011