title: Bumblebee

text: On this hypothesis, the molecular data suggest that the Bombini are 25 to 40 million years old, while the Meliponini (and thus the clade that includes the Bombini and Meliponini) are 81 to 96 million years old, about the same age as the corbiculate group.


title: Bumblebee

text: However, a more recent phylogeny using transcriptome data from 3,647 genes of ten corbiculate bee species supports the single origin of eusociality hypothesis in the corbiculate bees. They find that Bombini is in fact sister to Meliponini, corroborating that previous finding from Sophie Cardinal and Bryan Danforth (2011). However, Romiguier et al. (2015) shows that Bombini, Meliponini, and Apini form a monophyletic group, where Apini shares a most recent common ancestor with the Bombini and Meliponini clade, while Euglossini is most distantly related to all three, since it does not share the same most recent common ancestor as Bombini, Meliponini, and Apini. Thus, their analysis supports the single origin of eusociality hypothesis within the corbiculate bees, where eusociality evolved in the common ancestor of Bombini, Apini, and Meliponini.


title: Bumblebee

text: The fossil record for bees is limited, with around 14 species that might possibly be Bombini having been described by 2019. The only "Bombus" relatives in Bombini are the late Eocene "Calyptapis florissantensis" from the Florissant Formation, USA, and "Oligobombus cuspidatus" from the Bembridge Marls of the Isle of Wight. Two species of "Bombus" have been described from the Oligocene of Beşkonak, Bucak Turkey: "Bombus (Mendacibombus) beskonakensis" and "Bombus (Paraelectrobombus) patriciae". Both species were originally placed in genera considered at the time of description as outside of "Bombus", being initially named "Oligoapis beskonakensis" and "Paraelectrobombus patriciae" respectively, however reexaminiation of the fore-wings lead to both being considered as "Bombus" species In 2012 a fossil bumblebee from the Miocene was found in Germany's Randeck Maar and classified as "Bombus (Bombus) randeckensis". In 2014, another species, "Bombus cerdanyensis", was described from Late Miocene lacustrine beds of La Cerdanya, Spain, but not initially placed into any subgenus, The species "Bombus trophonius" was described in October 2017 and placed in "Bombus" subgenus "Cullumanobombus". A redescription of the Bombini fossil record by Dehon "et al" (2019) resulted in the synonymization of the genus "Oligoapis" with "Bombus" subgenus "Mendacibombus", and the placement of genus "Paraelectrobombus" as "Bombus" subgenus "Paraelectrobombus", rather than as a genus in Electrobombini. The subgenus "Cullumanobombus" was expanded to include not only "Bombus trophonius" but also "Bombus randeckensis" which was moved from subgenus "Bombus" and "Bombus pristinus", first described by Unger (1867). Within the subgenus "Melanobombus" only "Bombus cerdanyensis" is present from the fossil record. An additional three species, ""Bombus" luianus", ""Bombus" anacolus" and ""Bombus" dilectus" have been attributed to "Bombus" from the Middle Miocene Shanwang formation of China by Zhang, (1990) and Zhang "et al" (1994). Due to not being able to study Zhang's type specimens, but only illustrations of the fossils, Dehon "et al" did not place the three species within any specific subgenera, and considered all three as "species inquirenda", needing fuller re-examination. Two other species were not examined at all by Dehon "et al", "Bombus? crassipes" of the Late Miocene Krottensee deposits in the Czech Republic, and "Bombus proavus" from the Middle Miocene Latah Formation, USA.


title: Bumblebee

text: The genus "Bombus", the only one extant genus in the tribe Bombini, comprises over 250 species; for an overview of the differences between bumblebees and other bees and wasps, see characteristics of common wasps and bees. The genus has been divided variously into up to 49 subgenera, a degree of complexity criticised by Williams (2008). The cuckoo bumblebees "Psithyrus" have sometimes been treated as a separate genus but are now considered to be part of "Bombus", in one or more subgenera.


title: Bumblebee

text: Examples of "Bombus" species include "Bombus pauloensis", "Bombus dahlbomii", "Bombus fervidus", "Bombus lapidarius", "Bombus ruderatus", and "Bombus rupestris."


title: Bumblebee

text: Bumblebees vary in appearance, but are generally plump and densely furry. They are larger, broader and stouter-bodied than honeybees, and their abdomen tip is more rounded. Many species have broad bands of colour, the patterns helping to distinguish different species. Whereas honeybees have short tongues and therefore mainly pollinate open flowers, some bumblebee species have long tongues and collect nectar from flowers that are closed into a tube. Bumblebees have fewer stripes (or none), and usually have part of the body covered in black fur, while honeybees have many stripes including several grey stripes on the abdomen. Sizes are very variable even within species; the largest British species, "B. terrestris", has queens up to long, males up to long, and workers between long. The largest bumblebee species in the world is "B. dahlbomii" of Chile, up to about long, and described as "flying mice" and "a monstrous fluffy ginger beast".


title: Bumblebee

text: Bumblebees are typically found in temperate climates, and are often found at higher latitudes and altitudes than other bees, although a few lowland tropical species exist. A few species ("B. polaris" and "B. alpinus") range into very cold climates where other bees might not be found; "B. polaris" occurs in northern Ellesmere Island in the high Arctic, along with another bumblebee "B. hyperboreus", which parasitises its nest. This is the northernmost occurrence of any eusocial insect. One reason for their presence in cold places is that bumblebees can regulate their body temperature, via solar radiation, internal mechanisms of "shivering" and radiative cooling from the abdomen (called heterothermy). Other bees have similar physiology, but the mechanisms seem best developed and have been most studied in bumblebees. They adapt to higher elevations by extending their wing stroke amplitude. Bumblebees have a largely cosmopolitan distribution but are absent from Australia (apart from Tasmania where they have been introduced) and are found in Africa only north of the Sahara. More than a hundred years ago they were also introduced to New Zealand, where they play an important role as efficient pollinators.


title: Bumblebee

text: The bumblebee tongue (the proboscis) is a long, hairy structure that extends from a sheath-like modified maxilla. The primary action of the tongue is lapping, that is, repeated dipping of the tongue into liquid.


title: Bumblebee

text: The tip of the tongue probably acts as a suction cup and during lapping, nectar may be drawn up the proboscis by capillary action. When at rest or flying, the proboscis is kept folded under the head. The longer the tongue, the deeper the bumblebee can probe into a flower and bees probably learn from experience which flower source is best-suited to their tongue length. Bees with shorter proboscides, like "Bombus bifarius", have a more difficult time foraging nectar relative to other bumblebees with longer proboscides; to overcome this disadvantage, "B. bifarius" workers were observed to lick the back of spurs on the nectar duct, which resulted in a small reward.


title: Bumblebee

text: The exoskeleton of the abdomen is divided into plates called dorsal tergites and ventral sternites. Wax is secreted from glands on the abdomen and extruded between the sternites where it resembles flakes of dandruff. It is secreted by the queen when she starts a nest and by young workers. It is scraped from the abdomen by the legs, moulded until malleable and used in the construction of honeypots, to cover the eggs, to line empty cocoons for use as storage containers and sometimes to cover the exterior of the nest.


title: Bumblebee

text: The brightly coloured pile of the bumblebee is an aposematic (warning) signal, given that females can inflict a painful sting. Depending on the species and morph, the warning colours range from entirely black, to bright yellow, red, orange, white, and pink. Dipteran flies in the families Syrphidae (hoverflies), Asilidae (robber flies), Tabanidae (horseflies), Oestridae (bot or warble flies) and Bombyliidae (bee flies, such as "Bombylius major") all include Batesian mimics of bumblebees, resembling them closely enough to deceive at least some predators.


title: Bumblebee

text: Many species of "Bombus", including the group sometimes called "Psithyrus" (cuckoo bumblebees), have evolved Müllerian mimicry, where the different bumblebees in a region resemble each other, so that a young predator need only learn to avoid any of them once. For example, in California a group of bumblebees consists of largely black species including "B. californicus", "B. caliginosus", "B. vandykei", "B. vosnesenskii", "B. insularis" and "B. fernaldae". Other bees in California include a group of species all banded black and yellow. In each case, Müllerian mimicry provides the bees in the group with a selective advantage. In addition, parasitic (cuckoo) bumblebees resemble their hosts more closely than would be expected by chance, at least in areas like Europe where parasite-host co-speciation is common; but this too may be explained as Müllerian mimicry, rather than requiring the parasite's coloration to deceive the host (aggressive mimicry).


title: Bumblebee

text: Bumblebees are active under conditions during which honeybees stay at home, and can readily absorb heat from even weak sunshine. The thick pile created by long setae (bristles) acts as insulation to keep bumblebees warm in cold weather; species from cold climates have longer setae (and thus thicker insulation) than those from the tropics. The temperature of the flight muscles, which occupy much of the thorax, needs to be at least before flight can take place. The muscle temperature can be raised by shivering. It takes about five minutes for the muscles to reach this temperature at an air temperature of .


title: Bumblebee

text: The chill-coma temperature in relation to flying insects is the temperature at which flight muscles cannot be activated. Compared to honey bees and carpenter bees, bumblebees have the lowest chill-coma temperature. Of the bumblebees "Bombus bimaculatus" has the lowest at . However, bumblebees have been seen to fly in colder ambient temperatures. This discrepancy is likely because the chill-coma temperature was determined by tests done in a laboratory setting. However, bumblebees live in insulated shelters and can shiver to warm up before venturing into the cold.


title: Bumblebee

text: Bumblebees do not have ears, and it is not known whether or how well they can hear. However, they are sensitive to the vibrations made by sound travelling through wood or other materials.


title: Bumblebee

text: Bumblebees do not exhibit the "bee dances" used by honeybees to tell other workers the locations of food sources. Instead, when they return from a successful foraging expedition, they run excitedly around in the nest for several minutes before going out to forage once more. These bees may be offering some form of communication based on the buzzing sounds made by their wings, which may stimulate other bees to start foraging. Another stimulant to foraging activity is the level of food reserves in the colony. Bees monitor the amount of honey in the honeypots, and when little is left or when high-quality food is added, they are more likely to go out to forage.


title: Bumblebee

text: Bumblebees have been observed to partake in social learning. In a 2017 study involving "Bombus terrestris", bees were taught to complete an unnatural task of moving large objects to obtain a reward. Bees who first observed another bee complete the task were significantly more successful in learning the task than bees who observed the same action performed by a magnet, indicating the importance of social information. The bees did not copy one another exactly: in fact, the study suggested that the bees were instead attempting to emulate one another's goals.


title: Bumblebee

text: Nest size depends on species of bumblebee. Most form colonies of between 50 and 400 individuals, but colonies have been documented as small as ~20 individuals and as large as 1700. These nests are small compared to honeybee hives, which hold about 50,000 bees. Many species nest underground, choosing old rodent burrows or sheltered places, and avoiding places that receive direct sunlight that could result in overheating. Other species make nests above ground, whether in thick grass or in holes in trees. A bumblebee nest is not organised into hexagonal combs like that of a honeybee; the cells are instead clustered together untidily. The workers remove dead bees or larvae from the nest and deposit them outside the nest entrance, helping to prevent disease. Nests in temperate regions last only for a single season and do not survive the winter.


title: Bumblebee

text: In the early spring, the queen comes out of diapause and finds a suitable place to create her colony. Then she builds wax cells in which to lay her eggs which were fertilised the previous year. The eggs that hatch develop into female workers, and in time, the queen populates the colony, with workers feeding the young and performing other duties similar to honeybee workers. In temperate zones, young queens (gynes) leave the nest in the autumn and mate, often more than once, with males (drones) that are forcibly driven out of the colony. The drones and workers die as the weather turns colder; the young queens feed intensively to build up stores of fat for the winter. They survive in a resting state (diapause), generally below ground, until the weather warms up in the spring with the early bumblebee being the species that is among the first to emerge. Many species of bumblebee follow this general trend within the year. "Bombus pensylvanicus" is a species that follows this type of colony cycle. For this species the cycle begins in February, reproduction starts in July or August, and ends in the winter months. The queen remains in hibernation until spring of the following year in order to optimize conditions to search for a nest.


title: Bumblebee

text: In fertilised queens, the ovaries only become active when the queen starts to lay. An egg passes along the oviduct to the vagina where there is a chamber called the spermatheca, in which the sperm from the mating is stored. Depending on need, she may allow her egg to be fertilised. Unfertilised eggs become haploid males; fertilised eggs grow into diploid females and queens. The hormones that stimulate the development of the ovaries are suppressed in female worker bees, while the queen remains dominant.


title: Bumblebee

text: To develop, the larvae must be fed both nectar for carbohydrates and pollen for protein. Bumblebees feed nectar to the larvae by chewing a small hole in the brood cell into which they regurgitate nectar. Larvae are fed pollen in one of two ways, depending on the bumblebee species. Pocket-making bumblebees create pockets of pollen at the base of the brood-cell clump from which the larvae feed themselves. Pollen-storing bumblebees keep pollen in separate wax pots and feed it to the larvae.


title: Bumblebee

text: After the emergence of the first or second group of offspring, workers take over the task of foraging and the queen spends most of her time laying eggs and caring for larvae. The colony grows progressively larger and eventually begins to produce males and new queens.


title: Bumblebee

text: Bumblebee workers can lay unfertilised haploid eggs (with only a single set of chromosomes) that develop into viable male bumblebees. Only fertilised queens can lay diploid eggs (one set of chromosomes from a drone, one from the queen) that mature into workers and new queens.


title: Bumblebee

text: In a young colony, the queen minimises reproductive competition from workers by suppressing their egg-laying through physical aggression and pheromones. Worker policing leads to nearly all eggs laid by workers being eaten. Thus, the queen is usually the mother of all of the first males laid. Workers eventually begin to lay male eggs later in the season when the queen's ability to suppress their reproduction diminishes. Because of the reproductive competition between workers and the queen, bumblebees are considered "primitively eusocial".


title: Bumblebee

text: Although a large majority of bumblebees follow such monogynous colony cycles that only involve one queen, some select "Bombus" species (such as "Bombus pauloensis") will spend part of their life cycle in a polygynous phase (have multiple queens in one nest during these periods of polygyny).


title: Bumblebee

text: Bumblebees generally visit flowers that exhibit the bee pollination syndrome and these patches of flowers may be up to 1–2 km from their colony. They tend to visit the same patches of flowers every day, as long as they continue to find nectar and pollen there, a habit known as pollinator or flower constancy. While foraging, bumblebees can reach ground speeds of up to .


title: Bumblebee

text: Bumblebees use a combination of colour and spatial relationships to learn which flowers to forage from. They can also detect both the presence and the pattern of electric fields on flowers, which occur due to atmospheric electricity, and take a while to leak away into the ground. They use this information to find out if a flower has been recently visited by another bee. Bumblebees can detect the temperature of flowers, as well as which parts of the flower are hotter or cooler and use this information to recognise flowers. After arriving at a flower, they extract nectar using their long tongues ("glossae") and store it in their crops. Many species of bumblebees also exhibit "nectar robbing": instead of inserting the mouthparts into the flower in the normal way, these bees bite directly through the base of the corolla to extract nectar, avoiding pollen transfer.


title: Bumblebee

text: Pollen is removed from flowers deliberately or incidentally by bumblebees. Incidental removal occurs when bumblebees come in contact with the anthers of a flower while collecting nectar. When it enters a flower, the bumblebee's body hairs receive a dusting of pollen from the anthers. In queens and workers this is then groomed into the corbiculae (pollen baskets) on the hind legs where it can be seen as bulging masses that may contain as many as a million pollen grains. Male bumblebees do not have corbiculae and do not purposively collect pollen. Bumblebees are also capable of buzz pollination, in which they dislodge pollen from the anthers by creating a resonant vibration with their flight muscles.


title: Bumblebee

text: In at least some species, once a bumblebee has visited a flower, it leaves a scent mark on it. This scent mark deters bumblebees from visiting that flower until the scent degrades. This scent mark is a general chemical bouquet that bumblebees leave behind in different locations (e.g. nest, neutral, and food sites), and they learn to use this bouquet to identify both rewarding and unrewarding flowers, and may be able to identify who else has visited a flower. Bumblebees rely on this chemical bouquet more when the flower has a high handling time, that is, where it takes a longer time for the bee to find the nectar once inside the flower.


title: Bumblebee

text: Once they have collected nectar and pollen, female workers return to the nest and deposit the harvest into brood cells, or into wax cells for storage. Unlike honeybees, bumblebees only store a few days' worth of food, so are much more vulnerable to food shortages. Male bumblebees collect only nectar and do so to feed themselves. They may visit quite different flowers from the workers because of their different nutritional needs.


title: Bumblebee

text: Bees beat their wings about 200 times a second. Their thorax muscles do not contract on each nerve firing, but rather vibrate like a plucked rubber band. This is efficient, since it lets the system consisting of muscle and wing operate at its resonant frequency, leading to low energy consumption. Further, it is necessary, since insect motor nerves generally cannot fire 200 times per second. These types of muscles are called asynchronous muscles and are found in the insect wing systems in families such as Hymenoptera, Diptera, Coleoptera, and Hemiptera. Bumblebees must warm up their bodies considerably to get airborne at low ambient temperatures. Bumblebees can reach an internal thoracic temperature of 30 °C (86 °F) using this method.


title: Bumblebee

text: Bumblebees of the subgenus "Psithyrus" (known as 'cuckoo bumblebees', and formerly considered a separate genus) are brood parasites, sometimes called kleptoparasites, in the colonies of other bumblebees, and have lost the ability to collect pollen. Before finding and invading a host colony, a "Psithyrus" female, such as that of the "Psithyrus" species of "B. sylvestris", feeds directly from flowers. Once she has infiltrated a host colony, the "Psithyrus" female kills or subdues the queen of that colony, and uses pheromones and physical attacks to force the workers of that colony to feed her and her young. Usually, cuckoo bumblebees can be described as queen-intolerant inquilines, since the host queen is often killed to enable the parasite to produce more offspring, though some species, such as "B. bohemicus", actually enjoy increased success when they leave the host queen alive.


title: Bumblebee

text: The female "Psithyrus" has a number of morphological adaptations for combat, such as larger mandibles, a tough cuticle and a larger venom sac that increase her chances of taking over a nest. Upon emerging from their cocoons, the "Psithyrus" males and females disperse and mate. The males do not survive the winter but, like nonparasitic bumblebee queens, "Psithyrus" females find suitable locations to spend the winter and enter diapause after mating. They usually emerge from hibernation later than their host species. Each species of cuckoo bee has a specific host species, which it may physically resemble. In the case of the parasitism of "B. terrestris" by "B. (Psithyrus) vestalis", genetic analysis of individuals captured in the wild showed that about 42% of the host species' nests at a single location had "[lost] their fight against their parasite".


title: Bumblebee

text: Queen and worker bumblebees can sting. Unlike in honeybees, a bumblebee's stinger lacks barbs, so the bee can sting repeatedly without leaving the stinger in the wound and thereby injuring itself. Bumblebee species are not normally aggressive, but may sting in defence of their nest, or if harmed. Female cuckoo bumblebees aggressively attack host colony members, and sting the host queen, but ignore other animals unless disturbed.


title: Bumblebee

text: The sting is painful to humans, and not medically significant in most cases, although it may trigger an allergic reaction in susceptible individuals.


title: Bumblebee

text: Bumblebees, despite their ability to sting, are eaten by certain predators. Nests may be dug up by badgers and eaten whole, including any adults present. Adults are preyed upon by robber flies and beewolves in North America. In Europe, birds including bee-eaters and shrikes capture adult bumblebees on the wing; smaller birds such as great tits also occasionally learn to take bumblebees, while camouflaged crab spiders catch them as they visit flowers.


title: Bumblebee

text: The great grey shrike is able to detect flying bumblebees up to away; once captured, the sting is removed by repeatedly squeezing the insect with the mandibles and wiping the abdomen on a branch. The European honey buzzard follows flying bees back to their nest, digs out the nest with its feet, and eats larvae, pupae and adults as it finds them.


title: Bumblebee

text: Bumblebees are parasitised by tracheal mites, "Locustacarus buchneri"; protozoans including "Crithidia bombi" and "Apicystis bombi"; and microsporidians including "Nosema bombi" and "Nosema ceranae". The tree bumblebee "B. hypnorum" has spread into the United Kingdom despite hosting high levels of a nematode that normally interferes with queen bees' attempts to establish colonies. Deformed wing virus has been found to affect 11% of bumblebees in Great Britain.


title: Bumblebee

text: Female bee moths ("Aphomia sociella") prefer to lay their eggs in bumblebee nests. The "A. sociella" larvae will then feed on the eggs, larvae, and pupae left unprotected by the bumblebees, sometimes destroying large parts of the nest.


title: Bumblebee

text: Bumblebees are important pollinators of both crops and wildflowers. Because bumblebees do not overwinter the entire colony, they do not stockpile honey, and therefore are not useful as honey producers. Bumblebees are increasingly cultured for agricultural use as pollinators, among other reasons because they can pollinate plants such as tomato in greenhouses by buzz pollination whereas other pollinators cannot. Commercial production began in 1987, when Roland De Jonghe founded the Biobest company; in 1988 they produced enough nests to pollinate 40 hectares of tomatoes. The industry grew quickly, starting with other companies in the Netherlands. Bumblebee nests, mainly of buff-tailed bumblebees, are produced in at least 30 factories around the world; over a million nests are grown annually in Europe; Turkey is a major producer.


title: Bumblebee

text: Bumblebees are Northern Hemisphere animals. When red clover was introduced as a crop to New Zealand in the nineteenth century, it was found to have no local pollinators, and clover seed had accordingly to be imported each year. Four species of bumblebee from the United Kingdom were therefore imported as pollinators. In 1885 and 1886 the Canterbury Acclimatization Society brought in 442 queens, of which 93 survived and quickly multiplied. As planned, red clover was soon being produced from locally-grown seed. Bumblebees are also reared commercially to pollinate tomatoes grown in greenhouses. The New Zealand population of buff-tailed bumblebees began colonising Tasmania, away, after being introduced there in 1992 under unclear circumstances.


title: Bumblebee

text: Some concerns exist about the impact of the international trade in mass-produced bumblebee colonies. Evidence from Japan and South America indicates bumblebees can escape and naturalise in new environments, causing damage to native pollinators. Greater use of native pollinators, such as "Bombus ignitus" in China and Japan, has occurred as a result. In addition, mounting evidence indicates mass-produced bumblebees may also carry diseases, harmful to wild bumblebees and honeybees.


title: Bumblebee

text: In Canada and Sweden it has been shown that growing a mosaic of different crops encourages bumblebees and provides higher yields than does a monoculture of oilseed rape, despite the fact that the bees were attracted to the crop.


title: Bumblebee

text: Bumblebee species are declining in Europe, North America, and Asia due to a number of factors, including land-use change that reduces their food plants. In North America, pathogens are possibly having a stronger negative effect especially for the subgenus "Bombus". A major impact on bumblebees was caused by the mechanisation of agriculture, accelerated by the urgent need to increase food production during the Second World War. Small farms depended on horses to pull implements and carts. The horses were fed on clover and hay, both of which were permanently grown on a typical farm. Little artificial fertiliser was used. Farms thus provided flowering clover and flower-rich meadows, favouring bumblebees. Mechanisation removed the need for horses and most of the clover; artificial fertilisers encouraged the growth of taller grasses, outcompeting the meadow flowers. Most of the flowers, and the bumblebees that fed on them, disappeared from Britain by the early 1980s. The last native British short-haired bumblebee was captured near Dungeness in 1988. This significant increase in pesticide and fertilizer use associated with the industrialization of agriculture has had adverse effects on the genus "Bombus". The bees are directly exposed to the chemicals in two ways: by consuming nectar that has been directly treated with pesticide, or through physical contact with treated plants and flowers. The species "Bombus hortorum" in particular has been found to be affected by the pesticides; their brood development has been reduced and their memory has been negatively affected. Additionally, pesticide use negatively affects colony development and size.


title: Bumblebee

text: Bumblebees are in danger in many developed countries due to habitat destruction and collateral pesticide damage. The European Food Safety Authority ruled that three neonicotinoid pesticides (clothianidin, imidacloprid, and thiamethoxam) presented a high risk for bees. While most work on neonicotinoid toxicity has looked at honeybees, a study on "B. terrestris" showed that "field-realistic" levels of imidacloprid significantly reduced growth rate and cut production of new queens by 85%, implying a "considerable negative effect" on wild bumblebee populations throughout the developed world.


title: Bumblebee

text: However, in another study, following chronic exposure to field-realistic levels of the neonicotinoid pesticide thiamethoxam, colony weight gain was not affected, nor were the number or mass of sexuals produced. Low levels of neonicotinoids can reduce the number of bumblebees in a colony by as much as 55%, and cause dysfunction in the bumblebees' brains. The Bumblebee Conservation Trust considers this evidence of reduced brain function "particularly alarming given that bumblebees rely upon their intelligence to go about their daily tasks." A study on "B. terrestris" had results that suggests that use of neonicotinoid pesticides can affect how well bumblebees are able to forage and pollinate. Bee colonies that had been affected by the pesticide released more foragers and collected more pollen than bees who had not been dosed with neonicotinoid. Although the bees affected by the pesticide were able to collect more pollen, they took a longer amount of time doing so.


title: Bumblebee

text: Of 19 species of native nestmaking bumblebees and six species of cuckoo bumblebees formerly widespread in Britain, three have been extirpated, eight are in serious decline, and only six remain widespread. Similar declines have been reported in Ireland, with four species designated endangered, and another two considered vulnerable to extinction. A decline in bumblebee numbers could cause large-scale changes to the countryside, resulting from inadequate pollination of certain plants.


title: Bumblebee

text: Some bumblebees native to North America are also vanishing, such as "Bombus balteatus", "Bombus terricola", "Bombus affinis", and "Bombus occidentalis", and one, "Bombus franklini", may be extinct. In South America, "Bombus bellicosus" was extirpated in the northern limit of its distribution range, probably due to intense land use and climate change effects.


title: Bumblebee

text: In 2006 the bumblebee researcher Dave Goulson founded a registered charity, the Bumblebee Conservation Trust, to prevent the extinction "of any of the UK's bumblebees." In 2009 and 2010, the Trust attempted to reintroduce the short-haired bumblebee, "Bombus subterraneus", which had become extinct in Britain, from the British-derived populations surviving in New Zealand from their introduction there a century earlier. From 2011 the Trust, in partnership with Natural England, Hymettus and the RSPB, has reintroduced short-haired bumblebee queens from Skåne in southern Sweden to restored flower-rich meadows at Dungeness in Kent. The queens were checked for mites and American foulbrood disease. Agri-environment schemes spread across the neighbouring area of Romney Marsh have been set up to provide over 800 hectares of additional flower-rich habitat for the bees. By the summer of 2013, workers of the species were found near the release zone, proving that nests had been established. The restored habitat has produced a revival in at least five "Schedule 41 priority" species: the ruderal bumblebee, "Bombus ruderatus"; the red-shanked carder bee, "Bombus ruderarius"; the shrill carder bee, "Bombus sylvarum"; the brown-banded carder bee, "Bombus humilis" and the moss carder bee, "Bombus muscorum".


title: Bumblebee

text: The world's first bumblebee sanctuary was established at Vane Farm in the Loch Leven National Nature Reserve in Scotland in 2008. In 2011, London's Natural History Museum led the establishment of an International Union for Conservation of Nature Bumblebee Specialist Group, chaired by Dr. Paul H. Williams, to assess the threat status of bumblebee species worldwide using Red List criteria.


title: Bumblebee

text: Bumblebee conservation is in its infancy in many parts of the world, but with the realization of the important part they play in pollination of crops, efforts are being made to manage farmland better. Enhancing the wild bee population can be done by the planting of wildflower strips, and in New Zealand, bee nesting boxes have achieved some success, perhaps because there are few burrowing mammals to provide potential nesting sites in that country.


title: Bumblebee

text: According to 20th-century folklore, the laws of aerodynamics prove the bumblebee should be incapable of flight, as it does not have the capacity (in terms of wing size or beats per second) to achieve flight with the degree of wing loading necessary.


title: Bumblebee

text: The origin of this claim has been difficult to pin down with any certainty. John H. McMasters recounted an anecdote about an unnamed Swiss aerodynamicist at a dinner party who performed some rough calculations and concluded, presumably in jest, that according to the equations, bumblebees cannot fly. In later years, McMasters backed away from this origin, suggesting there could be multiple sources, and the earliest he has found was a reference in the 1934 book "" by French entomologist Antoine Magnan (1881–1938); they had applied the equations of air resistance to insects and found their flight was impossible, but "One shouldn't be surprised that the results of the calculations don't square with reality".


title: Bumblebee

text: Magnan refers to his assistant André Sainte-Laguë. Some credit physicist Ludwig Prandtl (1875–1953) of the University of Göttingen in Germany with popularizing the idea. Others say Swiss gas dynamicist Jakob Ackeret (1898–1981) did the calculations.


title: Bumblebee

text: The calculations that purported to show that bumblebees cannot fly are based upon a simplified linear treatment of oscillating aerofoils. The method assumes small amplitude oscillations without flow separation. This ignores the effect of dynamic stall (an airflow separation inducing a large vortex above the wing), which briefly produces several times the lift of the aerofoil in regular flight. More sophisticated aerodynamic analysis shows the bumblebee can fly because its wings encounter dynamic stall in every oscillation cycle.


title: Bumblebee

text: Additionally, John Maynard Smith, a noted biologist with a strong background in aeronautics, has pointed out that bumblebees would not be expected to sustain flight, as they would need to generate too much power given their tiny wing area. However, in aerodynamics experiments with other insects, he found that viscosity at the scale of small insects meant even their small wings can move a very large volume of air relative to their size, and this reduces the power required to sustain flight by an order of magnitude.


title: Bumblebee

text: The orchestral interlude "Flight of the Bumblebee" was composed (c. 1900) by Nikolai Rimsky-Korsakov. It represents the turning of Prince Guidon into a bumblebee so he can fly away to visit his father, Tsar Saltan, in the opera "The Tale of Tsar Saltan", although the music may reflect the flight of a bluebottle rather than a bumblebee. The music inspired Walt Disney to feature a bumblebee in his 1940 animated musical "Fantasia" and have it sound as if it were flying in all parts of the theater. This early attempt at "surround sound" was excluded from the film in later showings.


title: Bumblebee

text: In 1599, during the reign of Queen Elizabeth I, someone, possibly Tailboys Dymoke, published "Caltha Poetarum: Or The Bumble Bee", under the pseudonym "T. Cutwode". This was one of nine books censored under the Bishop's Ban issued by the Archbishop of Canterbury John Whitgift and the Bishop of London Richard Bancroft.


title: Bumblebee

text: Emily Dickinson made a bumblebee the subject of her parody of Isaac Watts's well-known poem about honeybees, "How Doth the Little Busy Bee" (1715). Where Watts wrote "How skilfully she builds her cell! How neat she spreads the wax!", Dickinson's poem, "The Bumble-Bee's Religion" (1881), begins "His little Hearse-like Figure / Unto itself a Dirge / To a delusive Lilac / The vanity divulge / Of Industry and Morals / And every righteous thing / For the divine Perdition / of Idleness and Spring." The letter was said to have enclosed a dead bee.


title: Bumblebee

text: The entomologist Otto Plath wrote "Bumblebees and Their Ways" in 1934. His daughter, the poet Sylvia Plath, wrote a group of poems about bees late in 1962, within four months of her suicide, transforming her father's interest into her poetry.


title: Bumblebee

text: The scientist and illustrator Moses Harris (1731–1785) painted accurate watercolour drawings of bumblebees in his "An Exposition of English Insects Including the Several Classes of Neuroptera, Hymenoptera, & Diptera, or Bees, Flies, & Libellulae (1776–80)".


title: Bumblebee

text: Bumblebees appear as characters, often eponymously, in children's books. The surname Dumbledore in the Harry Potter series (1997–2007) is an old name for bumblebee. J. K. Rowling said the name "seemed to suit the headmaster, because one of his passions is music and I imagined him walking around humming to himself". J. R. R. Tolkien, in his poem "Errantry", also used the name Dumbledor, but for a large bee-like creature.


title: Bumblebee

text: Among the many books for younger children are "Bumble the Bee" by Yvon Douran and Tony Neal (2014); "Bertie Bumble Bee" by K. I. Al-Ghani (2012); "Ben the Bumble Bee: How do bees make honey?" by Romessa Awadalla (2015); "Bumble Bee Bob Has a Big Butt" by Papa Campbell (2012); "Buzz, Buzz, Buzz! Went Bumble-bee" by Colin West (1997); "Bumble Bee" by Margaret Wise Brown (2000); "How the Bumble Came to Bee" by Paul and Ella Quarry (2012); "The Adventures of Professor Bumble and the Bumble Bees" by Stephen Brailovsky (2010). Among Beatrix Potter's "little books", Babbity Bumble and other members of her nest appear in "The Tale of Mrs. Tittlemouse" (1910).


title: Thunderbolts (comics)

text: The Thunderbolts are a fictional antihero team appearing in American comic books published by Marvel Comics. The team consists mostly of reformed supervillains.


title: Thunderbolts (comics)

text: The Thunderbolts first appeared in "The Incredible Hulk" #449 (January 1997) and were created by Kurt Busiek and Mark Bagley.


title: Thunderbolts (comics)

text: The Thunderbolts were first presented, both to readers and to the Marvel Universe, as a group of super-powered figures who became heroes to help protect the world when the Avengers were declared dead after the events of the 1996 "Onslaught" crossover. The final page of the first issue of their comic book, however, revealed that the Thunderbolts were actually the Masters of Evil in disguise, a surprise twist carefully guarded by Marvel.


title: Thunderbolts (comics)

text: In subsequent storylines, the group rejects their leader Baron Helmut Zemo and attempts to become heroes in their own right, eventually under the leadership of the Avenger Hawkeye.


title: Thunderbolts (comics)

text: Themes of redemption and the nature of heroism are often featured in Thunderbolts comics. The book has also garnered critical praise for its use of secondary characters from other Marvel Comics and its use of continuity-themed storytelling.


title: Thunderbolts (comics)

text: The Thunderbolts was an original concept created for Marvel by Busiek and Bagley. Most of the characters used in the final concept were reimagined versions of existing Marvel characters, with additional original characters for the series developed by Busiek and designed by Bagley. The pair also created the new heroic identities for the Masters of Evil. Busiek recalled:


title: Thunderbolts (comics)

text: The actual origin of "Thunderbolts" came when I used to live in New Jersey and drive to New England to visit my parents. To keep myself awake, I'd give myself books to write, and work out about two to three years of continuity. One trip, I assigned myself "Avengers", and came up with the plan that the Masters of Evil would ultimately conquer them by posing as new heroes and slowly replacing them. At the time, I thought it was a neat idea, and filed it away.


title: Thunderbolts (comics)

text: While Busiek's original reluctance was because he deemed readers would not have liked replacing the established characters and then having the new ones being villains – "Doing that with one character, like what Marv Wolfman did with Terra, made a great sting, but doing it on a team-wide scale wouldn't work." – seeing a world devoid of many heroes following "Onslaught" made him revive the concept. The Thunderbolts first appeared as a team in "The Incredible Hulk" #449 (January 1997), written by Peter David and illustrated by Mike Deodato. Originally intended to be a similar team known as the "Echelon," the synchronization of the plans led to the Thunderbolts being used instead as a "teaser" for their own series. No mention was made of the connection between the Thunderbolts and the Masters of Evil in this appearance, save perhaps for the Hulk almost recognizing Meteorite's voice (having fought Moonstone before). The team also appeared in a one-shot called "Tales of the Marvel Universe". The twist would not be revealed until the first issue of their own series.


title: Thunderbolts (comics)

text: Soon after the publication of "The Incredible Hulk" #449, the team's own series premiered. The first issue, cover dated April 1997, was played largely as a straight superhero story, until the revelation of the Thunderbolts' true nature on the last page of the comic. This is considered one of the most well-conceived plot twists in the history of American comic books, with "Wizard" magazine readers voting it "Comics' Greatest Moment of 1997" and later, in 1998, placing it at #11 on a list of "The 25 Greatest Comic Moments Ever". Marvel managed to keep the secret of the Thunderbolts' true villainous identities tightly under wraps before the book launched. When word got out, the first issue sold out so quickly that Marvel not only offered a second printing, but also did a "mini-trade paperback" collecting the first two issues. Fabian Nicieza replaced Busiek in issue #34. Patrick Zircher, after a couple of fill-ins, replaced Bagley in issue #51.


title: Thunderbolts (comics)

text: Despite critical acclaim, the book was reformatted with "Thunderbolts" #76 (March 2003), removing the entire cast and creative team and replacing it with a brand new set of characters, along with a new writer, John Arcudi. The move was done in part due to Marvel Editor-in-Chief Joe Quesada's desire to emulate the success he had with "X-Force", which was reformatted with a new cast of characters and status quo that was successful in sales and popularity. However, the new direction for the series—an underground fighting circuit that employed predominantly newly created super-villain characters—was a commercial failure and canceled after six issues.


title: Thunderbolts (comics)

text: In 2004, Marvel Comics launched a limited series titled "Avengers/Thunderbolts", which continued one year after the events of issue #75. The limited series ran for the same number of issues as the reformatted Thunderbolts arc.


title: Thunderbolts (comics)

text: Soon after the completion of "Avengers/Thunderbolts", Marvel Comics launched a second series featuring the characters with "New Thunderbolts" #1 (January 2005). The storyline continued the events from "Avengers/Thunderbolts" as well as the fall-out of "Avengers Disassembled" and returned to the original series concept, though with a roster that lacked many fan favorites (such as Baron Zemo, Moonstone, and Techno). With the combination of the eighty-one issues of the first series and the first eighteen issues of "New Thunderbolts", the series reverted to its original numbering with "Thunderbolts" #100.


title: Thunderbolts (comics)

text: "Thunderbolts" #110 saw another change to the direction of the series, with writer Warren Ellis introducing a new team of Thunderbolts, villains working for the government, tasked with capturing unregistered superheroes. Ellis has stated that he chose to approach the series "gently, but directly from a political agenda" and the relaunch was closely tied to Marvel's commercially successful Civil War event, with the team serving as a dark reflection of the event's controversial ending. The Thunderbolts also feature in the Spider-Man storyline "New Ways to Die", which is the first proper showdown between him and the team.


title: Thunderbolts (comics)

text: Ellis stepped aside in issue #121 and was replaced by Christos Gage, who wrote three one-shots and a four-issue tie-in with Secret Invasion. At the 2008 San Diego Comic Con, Andy Diggle was announced as the new ongoing writer, starting in issue #126 (November 2008) with a two issue story "Burning Down the House" which cleared the way for the introduction of a new team line-up. This team debuted in "Thunderbolts" #128-129, a story that dealt with "Dark Reign", the Secret Invasion aftermath, which was followed by "Magnum Opus", a 4-issue crossover with "Deadpool" (vol. 2). Miguel Sepulvida took over art duties with "Thunderbolts" #133 and Jeff Parker became the new writer with issue #138. Parker then piloted the title through the end of "Dark Reign", featuring a crossover with the Agents of Atlas team he was also writing, and into "Siege", following which the team was revamped again. Parker announced that "the status quo of the team undergoes a major overhaul for the new era to come. It's going to synthesize a lot of what readers like about recent history and re-instill some elements from the early days of the book."


title: Thunderbolts (comics)

text: The Heroic Age team debuted in "Thunderbolts" #144 with a new main artist, Kev Walker. The title crossed over with "Avengers Academy" in issue #147, which was bookended by "Avengers Academy" #3 and #4. The series then went on to cross over with the Daredevil storyline "Shadowland" in issues #148-149, with artist Declan Shalvey stepping in for the two issues. The series then crossed over with the "Fear Itself" storyline in issues #158-163.


title: Thunderbolts (comics)

text: The "Thunderbolts" comic book was renamed "Dark Avengers" beginning with issue #175, but the creative team remained unchanged. Dark Avengers ended with issue #190.


title: Thunderbolts (comics)

text: As part of Marvel NOW!, a new Thunderbolts series was launched featuring a new team composed of Red Hulk, Deadpool, Elektra, Venom, and Punisher. This series ended in October 2014 with issue #32.


title: Thunderbolts (comics)

text: Baron Helmut Zemo summoned several of his former allies from the prior incarnations of the Masters of Evil during a rescue attempt of Goliath (the former bodyguard to Baron Heinrich Zemo). The summoned members included Beetle, Fixer, Moonstone and Screaming Mimi. Zemo took the accidental gathering as an omen, and decided to reform the Masters of Evil and attack the Avengers. Before they could strike, the Avengers, and the Fantastic Four were apparently killed by the villain Onslaught.


title: Thunderbolts (comics)

text: The death of the Avengers and Fantastic Four created an opportunity for Zemo and the Masters of Evil. Zemo realized that the world needed superpowered champions, and that his team could fill that need. By posing as superheroes, the Masters of Evil could gain the public trust and build a position of power that rivaled the status of the Avengers. Once they had gained the public's faith, Zemo believed they could gain access to all the secrets of the Avengers and S.H.I.E.L.D. once they were in ultimate power. Zemo then planned to sell the secrets they found to the criminal underworld.


title: Thunderbolts (comics)

text: The villains adopted new heroic costumes and codenames. Baron Zemo became the patriotic American Citizen V, patterned after a WWII hero his father killed. Former Human Torch and Spider-Man foe Beetle became MACH-1, using a new high-tech suit designed by Fixer. Fixer became the gadget-wielding Techno. Long time Avengers enemy Goliath became the powerhouse Atlas. Screaming Mimi became Songbird, her damaged abilities aided by technology Zemo had adapted from ex-Master Klaw. Moonstone was secretly freed from the Vault and added to the team by Zemo, who extracted a promise of loyalty from her. She was to be Zemo's personal enforcer against any betrayal committed by the others. She took the alias Meteorite. Calling themselves the Thunderbolts, the six new 'heroes' were ready for action.


title: Thunderbolts (comics)

text: The team found tremendous success as superheroic champions. The public began to think of the Thunderbolts as heroes. After several adventures, some of the villains began to think of themselves the same way. Dallas Riordan, an aide to the Mayor of New York, befriended the new heroes.


title: Thunderbolts (comics)

text: Jolt, an Asian American teenage girl whose entire family was killed by Onslaught and was experimented on by Arnim Zola, soon joined the team after she came to the Baxter Building seeking the help of the FF. Jolt, however, was not a supervillain, with the young girl honestly believing that her new friends were heroes (Zemo used the opportunity to create the team's new history). Soon after the addition of Jolt, Techno's neck was broken in battle with the Elements of Doom. Techno then seemingly transferred his mind into an android body built from his tech-pack.


title: Thunderbolts (comics)

text: Just as Zemo's plans were about to come to fruition, everything unravelled. To the astonishment of the entire world, the Fantastic Four and the Avengers returned. Faced with the return of the lost heroes, Zemo revealed the true nature of the Thunderbolts to the world. Ostensibly, he did this to ensure the loyalty of the team by ruining their chances of becoming heroes.


title: Thunderbolts (comics)

text: The Thunderbolts (minus the android Techno) turned on Zemo for his betrayal. In the ensuing battle, Zemo and Techno used a mind control device to turn the Avengers and Fantastic Four against the remaining Thunderbolts, who, with the help of the size-changing Atlas- as well as Iron Man, who had designed his armour to make himself more resistant to mind control- ultimately rallied and freed the other heroes. Together, they defeated Zemo and Techno. Unbeknownst to his teammates, Atlas helped the wounded Zemo escape, while Techno fled under his own power.


title: Thunderbolts (comics)

text: Amidst this chaos, Meteorite decided to return to her old identity. She altered her costume and changed her codename back to Moonstone. After a brief stop-over in an alternate dimension, the team learned that Moonstone had no intention of reforming and becoming a superhero. She told them she only turned against Zemo out of self-preservation.


title: Thunderbolts (comics)

text: Now fugitives, new members soon joined the Thunderbolts. These members included the former Avenger Hawkeye, and later a young African American hero named Charcoal who had previously fought the team, having been created by a "Create a character" contest in "Wizard Magazine". Hawkeye convinced his new teammates they would be pardoned if MACH-1, who had murdered someone as the Beetle (as the only member of the team who had ever actually "killed" anyone), turned himself in to authorities. The team considered joining the mysterious Crimson Cowl's new Masters of Evil instead. Ultimately, the Thunderbolts followed Hawkeye's advice. MACH-1 turned himself in and pleaded guilty to murder. Even with MACH-1's surrender, the U.S. Government refused to pardon the group.


title: Thunderbolts (comics)

text: The Thunderbolts then defeated the Crimson Cowl's Masters of Evil and took over their headquarters. After unmasking the Crimson Cowl, the team discovered that she was their old friend Dallas Riordan. In truth, Riordan was not the Crimson Cowl. The real Crimson Cowl was Justine Hammer. Riordan had been framed by Hammer to take the fall for the Cowl's crimes. While Riordan was not the Crimson Cowl, she did have her own secret identity. Riordan was actually the new Citizen V, leader of the secret V-Battalion. Riordan decided to keep her secret to avoid exposure of the group. She was sent to jail for the Crimson Cowl's crimes, but was later rescued by the V-Battalion.


title: Thunderbolts (comics)

text: While exploring their new headquarters, the Thunderbolts discovered Ogre. Ogre was a former member of the villainous Factor Three, the original owners of the base. Factor Three made Ogre the new base caretaker after they disbanded. This also left him with custody of Humus Sapien, a dangerous mutant teenager that Factor Three had kidnapped and placed in suspended animation.


title: Thunderbolts (comics)

text: Ogre was accepted as a member of the Thunderbolts. Soon after, Techno attacked him, placed him in stasis, and assumed his identity. At the same time, MACH-1 was freed from prison in exchange for stealing some top secret weapons technology from evil industrialist Justin Hammer. MACH-1 returned to the Thunderbolts after gaining his freedom thanks to Hawkeye making a deal with the Commission that had released him that he would continue officially serving his sentence while returning to the team. Upon his return, Techno upgraded MACH-1's armor, and used another device to change his features so that others wouldn't realize he was the same person as the original identity. MACH-1 was now MACH-2.


title: Thunderbolts (comics)

text: During this time, the Thunderbolts joined forces with the Avengers against the latest plan of Count Nefaria, as he intended to detonate an ionic bomb, which will transform millions of people into an ionic state which he can then control, perceiving it as the best way to guarantee that he receives the respect that he feels he deserves. The two teams are drawn into this plan when Nefaria uses his new control of ionic energy to take control of the heroes Wonder Man and Atlas, intending to use them to kill the Avengers. Despite his power, he is stopped by the combined efforts of the Avengers, the Thunderbolts, and Madame Masque, Masque using a weapon she had developed to disrupt Nefaria's own ionic energy so that the heroes can defeat him.


title: Thunderbolts (comics)

text: Meanwhile, Moonstone found herself going through changes. She fell in love with Hawkeye. Soon after, she learned that the spirit of the Kree Moonstone that powered her tried to make her a more honest person. This conflicted with her naturally amoral personality, and slowly made her insane.


title: Thunderbolts (comics)

text: The Thunderbolts faced many more challenges. Henry Gyrich sought to destroy the team and Hawkeye. Gyrich changed a brainwashed Jack Monroe into the new Scourge. The Scourge then attempted to assassinate the Thunderbolts one by one. First he killed Jolt. He then traveled to South America and apparently killed Baron Zemo. After this, Scourge broke into the Thunderbolts headquarters and destroyed the robotic Techno. He then killed Atlas by allowing the giant to implode into a storm of ionic energy.


title: Thunderbolts (comics)

text: All four characters apparently killed by Scourge had survived in some form: the robotic Techno had recovered Jolt's body and used her electrical powers to resurrect her from the dead, even while he "died", imbuing her with the knowledge that Hawkeye had failed to get the team pardoned in the process. The Thunderbolts were upset with Hawkeye, but, on unmasking Monroe, decided that confronting Gyrich was more important.


title: Thunderbolts (comics)

text: While all this happened, Val Cooper had gathered her own army of heroes she named the Redeemers. This team included a new Citizen V, Atlas' brother Smuggler, and Fixer, who had only copied his mind into his tech-pack as a contingency. The original Norbert P. Ebersol survived his injuries and recovered in secret while the robotic Techno continued on as a Thunderbolt in his place. Leila Davis, the wife of the Ringer, used an updated version of the Beetle armour.


title: Thunderbolts (comics)

text: The Redeemers helped the Thunderbolts battle Gyrich, who had acquired experimental nanite technology. Gyrich wanted to use the nanites to kill off all heroes and villains on Earth. Gyrich's scheme was foiled. It was soon revealed that he had been infected with nanites, and had been secretly manipulated by Baron Strucker of the terrorist group HYDRA. Hawkeye tried to use this information as blackmail to get the Thunderbolts pardoned. Gyrich countered that he would tell the public himself about HYDRA's scheme. Ultimately, Gyrich agreed to stay quiet and give the Thunderbolts their pardon. In return, Hawkeye turned himself in for aiding the fugitive heroes. Hawkeye went to prison and the team disbanded.


title: Thunderbolts (comics)

text: Eventually, all the others murdered by Jack Monroe returned from the dead. Baron Zemo's mind had been transferred into the comatose body of the man whose role as Citizen V he had usurped in the first place.


title: Thunderbolts (comics)

text: Later, after a teleportation accident, Zemo's mind was transferred into Techno's mechanical "Tech-Pack", which had also cybernetically replaced the broken segment of Techno's real body's spine. Atlas was later raised from the dead after a merger with Riordan, who had been crippled in battle with the Crimson Cowl.


title: Thunderbolts (comics)

text: Jolt and Charcoal, the only Thunderbolts without criminal records, were assimilated into the Redeemers under the leadership of Captain America and the Zemo-possessed Citizen V. The Redeemers were promptly slaughtered by the Thunderbolts' deadliest foe, the powerful supervillain Graviton with Citizen V, Fixer (who ran away), and Jolt (who re-formed her electric form) as the only survivors of the massacre, although Smuggler and Screamer have since resurfaced.


title: Thunderbolts (comics)

text: The Thunderbolts reformed to defeat Graviton. During the fight, several of the team members present (Fixer, Jolt, Moonstone, Jenkins as MACH-3 and the merged Atlas/Dallas Riordan, along with Zemo's mind — accidentally transferred into Fixer's tech-pack by the teleportation) were transported to Counter-Earth, the same parallel Earth the Avengers and Fantastic Four were sent to after their final battle with Onslaught. The Thunderbolts met Counter-Earth versions of Heinrich Zemo, Helmut Zemo and the first Moonstone, the last of which was known as Phantom Eagle.


title: Thunderbolts (comics)

text: Under duress — Zemo being able to disable his ability to walk at will — Fixer transferred Zemo's mind from "Tech-Pack" into the body of Zemo's counterpart. Zemo then killed the Counter-Earth version of his father. Soon after, the Thunderbolts stopped the Nazi Germany of Counter-Earth from taking control of all of Counter-Earth's computers. Zemo convinced the team to remain and help rebuild Counter-Earth. The team reluctantly agreed and based themselves in the mobile Counter-Earth Attilan. Then, Moonstone stole the mentally unstable Phantom Eagle's moonstone for herself, boosting her powers to godlike levels.


title: Thunderbolts (comics)

text: Back on Earth-616, many things happened. Hawkeye escaped from prison alongside several supervillains just as S.H.I.E.L.D. contacted him with an offer to be freed from prison. Industrialist Justin Hammer died. His daughter Justine (the Crimson Cowl) discovered her father had exposed every supervillain he ever employed to a poison that enslaved their minds. This included members of the Crimson Cowl's Masters of Evil. However, the villain Plantman had helped create the poison and was the only one who could activate it.


title: Thunderbolts (comics)

text: With the telepathic terrorist Mentallo serving as a middleman, Hawkeye tried to help Plantman in order to give him to Crimson Cowl. During the escape, Hawkeye watched helplessly as Plantman murdered a prison guard. Before Crimson Cowl could kill Hawkeye or take Plantman, they were rescued by Songbird.


title: Thunderbolts (comics)

text: Hawkeye and Songbird then formed a second group of Thunderbolts. He explained to the Crimson Cowl's Masters of Evil that if the Crimson Cowl was not stopped, they would all become her slaves. Plantman, using the codename Blackheath, was their first new member. Most of the members of Crimson Cowl's Masters of Evil also joined, including: Cardinal (now Harrier); Gypsy Moth (now Skein); Man-Killer (now Amazon); and Cyclone, who did not change his codename.


title: Thunderbolts (comics)

text: These new Thunderbolts were eventually captured by the Crimson Cowl (who was helped by Cyclone). Crimson Cowl vivisected Plantman. Soon after, Plantman mutated into a plant creature that neutralized the mind-control poison. The group was then sent to the V-Battalion's base. The Counter-Earth group returned to Earth at the V-Battalion base through a rift in space. The new and old teammates were reunited. Jolt stayed on Counter-Earth and joined the Young Allies. Closing the rift between Earth and Counter-Earth destroyed the V-Battalion's base.


title: Thunderbolts (comics)

text: In the aftermath, Atlas and Dallas were split into separate bodies, with Dallas retaining the remainder of Atlas' ionic power, allowing her to walk again with enhanced agility and strength. Zemo convinced Hawkeye that he wanted to reform and help the world instead of ruling it. Hawkeye, Amazon and Skein left the team. MACH-3 and Harrier returned to prison. Zemo then revealed to the team that he had lied to Hawkeye and that he still wanted to conquer the world, only to save it from itself. The "Thunderbolts" comic then shifted focus for six issues.


title: Thunderbolts (comics)

text: In 2004, the six issue "Avengers/Thunderbolts" limited series was launched, picking up a year after the events of "Thunderbolts" #75. Zemo led the Thunderbolts (now including Dallas Riordan, under the codename Vantage) in an attempt to drain the powers of all superhumans on Earth, using Moonstone. They fought the Avengers, including former Thunderbolt Hawkeye. The Avenger Iron Man infiltrated the Thunderbolts disguised as Cobalt Man. Eventually, all the power absorbed by Moonstone caused her to snap. Jolt returned from Counter-Earth to help stop Moonstone. Finally, Iron Man convinced Hawkeye to lobotomize Moonstone to save the planet.


title: Thunderbolts (comics)

text: Zemo vowed revenge against the Thunderbolts and the Avengers for putting Moonstone into a coma. Jolt returned to Counter-Earth. A depowered Blackheath returned to prison. The Fixer fled. Vantage retired to a government job. Songbird was offered reserve membership in the Avengers but turned it down. MACH-3 was paroled from prison and decided to form a new team of Thunderbolts.


title: Thunderbolts (comics)

text: Marvel subsequently launched "New Thunderbolts" #1. MACH-3 (now called MACH-IV), Atlas and Songbird were now a part of the new Thunderbolts. The team's new recruits included Photon, Speed Demon, Joystick, Blizzard II and the Radioactive Man. The new team has battled Atlantean superhuman terrorist group the Fathom Five and Baron Strucker's HYDRA organization, which funded the team's return.


title: Thunderbolts (comics)

text: In "Purple Reign", Swordsman, along with his master, the Purple Man, plotted to enslave New York City by drugging the water supply with the Purple Man's pheromones, which allowed him to control his victims. While the Thunderbolts eventually defeated the Purple Man, before they could interrogate him he was teleported away by his boss, Baron Zemo, who tortured him by inducing rigor mortis in his body and sent him back to prison where his powers would be nullified.


title: Thunderbolts (comics)

text: Later, Hank Pym and Warbird offered to pardon the members of the Thunderbolts (who still had outstanding legal problems), if the Thunderbolts would attack and humiliate the New Avengers in public. However, it was later revealed that Pym and Warbird had been blackmailed into doing so.


title: Thunderbolts (comics)

text: Only Spider-Woman survived unscathed from the Thunderbolts' sneak attack, beating Joystick senseless. The rest of the team was beaten back before the Thunderbolts left. Songbird told Captain America that the Thunderbolts could beat the New Avengers senseless anytime they wished. In the end, it was revealed that Baron Zemo spearheaded the attack. He did this to humiliate Captain America but also to see how far the Thunderbolts would go for the chance at being pardoned.


title: Thunderbolts (comics)

text: Meanwhile, new threats were rising as Fixer resurfaced and recruited both MACH-IV and Blizzard to work with him on a top secret project, which was also run by Zemo. Meanwhile, Speed Demon was confronted by the new female Doctor Spectrum, who was out to reform the Squadron Sinister and take over the world. In the end, Speed Demon quit the Thunderbolts to join Dr. Spectrum while Nighthawk, the former Defender and member of the original Squadron Sinister, was offered membership on the Thunderbolts by Songbird.


title: Thunderbolts (comics)

text: Soon after Nighthawk joined the team, the Thunderbolts discovered that Photon's awareness of all space and time brought with it a vision of the destruction of the universe at his hands. As they tried to come to terms with this, they were attacked by a Moonstone puppeteered by Zemo to kill Genis-Vell. When the initial strike failed, the remainder of Zemo's team revealed themselves to the Thunderbolts. Zemo explained that he had used the Moonstones to accelerate Genis' return from death, and in the process made the mistake of siphoning energy from the beginning and end of time itself, caused by inexperience with his Moonstones, creating a link between Genis and the universe that threatened to end existence. Unable to find a way to save both Genis and the universe, he concluded that the only solution was Genis's death.


title: Thunderbolts (comics)

text: Unwilling to accept this Songbird led the team in combat against Zemo's group. To prevent the other Thunderbolts from intervening, Zemo revealed that Atlas' brother Smuggler had survived Graviton's massacre of the Redeemers, trapped in the Darkforce dimension. Using the prospect of his release to make Atlas stop the other Thunderbolts from interfering, Zemo bested Genis in battle and, apologizing for both his mistake and the necessary solution, sliced Genis' body into pieces and scattered them through both time and the Darkforce dimension to prevent Genis returning from the dead a third time. He then fully released Smuggler. In the aftermath, Zemo—his face apparently fully healed from Moonstone's attack—and Songbird became allies and lovers.


title: Thunderbolts (comics)

text: The new Thunderbolts engaged and defeated Quicksand in a battle in Denver during the early days of the superhero Civil War. After this, they were summoned to Washington where they met with Iron Man, Mister Fantastic, and Yellowjacket. The three heroes, all supporting the Superhuman Registration Act, informed Zemo that they wanted the Thunderbolts to hunt down supervillains and recruit them to the Pro-Registration cause, which would be their chance at redemption. Unknown to Iron Man, the Thunderbolts had been doing this in secret for three weeks.


title: Thunderbolts (comics)

text: Zemo's "Thunderbolt Army" grew rapidly, the team vastly expanding. It now included dozens of other supervillains, including most notably Doctor Octopus, the Wrecker, and Ox. The new team dispersed to battle super-villains, capture them and offer them a choice: join the Thunderbolts or go to prison. Of course, they all chose to join the Thunderbolts. Baron Zemo convinced Captain America not to stop him from battling the Grandmaster, while Nighthawk was revealed as being a spy for the Squadron Sinister inside Captain America's Secret Avengers. Zemo then informed Songbird that in the coming battle, he knew that she would betray him, which she had been planning to do all along, in revenge for his killing of Photon, and he would sacrifice himself to save the world.


title: Thunderbolts (comics)

text: Zemo then saved the Wellspring of Power from the Grandmaster, who planned to use it for his own ends. Believing that all of his visions were subject to the flow of time, and that nothing was set in stone, Zemo defeated the Grandmaster, and boasted to his teammates that the power was now all his and theirs. He insisted that he would use it to help the world, despite the consequences for doing so. Songbird, who had temporarily lost her powers during the final battle, was told by Zemo, "now is when your betrayal would have come." The vision of her betrayal turned out to be somewhat correct after all, though. Although she could not use her super-sound, Songbird used a simple opera note to crack the moonstones, sending Zemo into a whirlwind of cosmic time/space. In his final words before he was completely sucked into the vacuum, he screamed out that he would never have hurt a world he worked so hard to save.


title: Thunderbolts (comics)

text: Some of the Army of Thunderbolts, consisting of Venom, Lady Deathstrike, Taskmaster, Bullseye, Jester, Jack O'Lantern join Songbird assembled as the Pro-Registration side's task force to hunt down Anti-Registration heroes at the end of "Civil War" #4, but never officially saw any real combat. The Jester and Jack O'Lantern were later killed by the Punisher. The two were sent to hunt down and capture Spider-Man when he attempted to leave Iron Man's Pro-Registration army.


title: Thunderbolts (comics)

text: Bullseye, Taskmaster, and Lady Deathstrike were part of the final battle of Civil War fighting alongside the Pro-Registration side, with Taskmaster wounding Mr. Fantastic when he sought to kill Invisible Woman (a member of the Anti-Registration forces). At the end of the fight, Taskmaster, Bullseye, and Deathstrike were sent to the Negative Zone prison; but Bullseye escaped before he could be sent, and Taskmaster was freed by Deadpool en route to the portal. Deathstrike was either released or somehow escaped on her own. She would go on to battle the X-Men during the events of "", a story arc in the X-Men related books.


title: Thunderbolts (comics)

text: After the events surrounding the Wellspring, the current team disbanded. MACH-IV and the Fixer were offered jobs from the Commission on Superhuman Activities. Blizzard was released from jail and left the team. Atlas was de-ionized after his encounter with the Wellspring but left catatonic and trapped in his enlarged state while Smuggler, his suit destroyed, took care of him. Joystick was imprisoned for her traitorous actions during the Wellspring debacle and Speed Demon ran away to avoid arrest. Only Songbird, Moonstone, Swordsman and the Radioactive Man, remained on the team.


title: Thunderbolts (comics)

text: In the events of "Thunderbolts: Zemo - Born Better" mini-series, Baron Zemo found himself lost in time and in Europe, forced to witness the lives and deaths of the previous generations of "Baron Zemos", while being violently thrust forward in time at random points of his adventures. The journey forward in time was a prolonged "moment of clarity" experience for Zemo, who saw his family's true history and not the rose-colored version his father and grandfather had taught him. When he ultimately returned to the present day, Zemo discovered that a distant cousin had been responsible for his return to the present, though at a horrific cost of causing Zemo to jump forward from era to era every time his cousin murdered a family member who shared the same blood as Helmut. When Helmut discovered that his cousin rescued Zemo just so he could kill his infamous relative, Helmut refused to resist, a move that caused his cousin to attempt to take his own life before Zemo convinced him to put down his gun and stop the cycle of violence.


title: Thunderbolts (comics)

text: "Thunderbolts" #110 featured a new creative team (writer Warren Ellis and artist Mike Deodato) and a new roster and direction for the team. The team was formed under the control of Norman Osborn, and the majority of the roster was made up of villains wishing to redeem themselves.


title: Thunderbolts (comics)

text: After the Marvel Comics "Civil War" event, the new Thunderbolts were tasked with tracking down and arresting individuals avoiding the Superhuman Registration Act. Despite the new line-up, many of the themes of the original series continued; in particular, Songbird's decision to cripple Bullseye in retaliation for the villain crippling the super-hero Jack Flag, Norman Osborn's own struggle for redemption as he claimed, and the growing friendship between Songbird and Radioactive Man. Moonstone was made team leader, a move that proved to be disastrous and ultimately led to Songbird resuming control over the team and expanding the group's missions to involve regular super-hero missions besides hunting down unregistered super-heroes.


title: Thunderbolts (comics)

text: Due to lengthy delays towards the end of Ellis's run, several one-shot specials and a "Penance" limited series were launched to fill in the gap. The "Penance" limited series saw the hero formerly known as Speedball carrying out a complex scheme hatched to gain revenge against Nitro, the man who murdered his teammates the New Warriors, which put him at odds with his new teammates on the Thunderbolts. Several specials advanced other plotlines, such as the mysterious death of Songbird's mother and Swordsman betraying the Thunderbolts by allying himself with Arnim Zola to resurrect his sister, after realizing that Osborn would not use his cloning technology to resurrect Andrea.


title: Thunderbolts (comics)

text: Following the events of "Caged Angels", the various Thunderbolt specials (written by Christos Gage), and the Penance limited series, and guest appearances in "Moon Knight" and "The Amazing Spider-Man", the group is thrust into the events of "Secret Invasion", when Thunderbolt Mountain is attacked by the Skrull Khn'nr, also known as Skrull sleeper agent Captain Marvel, just as Arnim Zola drops off the newly resurrected (via cloning technology) Andrea Strucker.


title: Thunderbolts (comics)

text: Barely surviving the fight due to Khn'ner's inability to deal with the implanted memories inside of him belonging to Mar-Vell, the group went onto the offensive against the Skrull armada as they invaded Washington DC. During the fight, Andrea is teamed with her brother Swordsman, Moonstone and Bullseye under the fear that she is a Skrull spy sent to infiltrate the group. However, when Moonstone betrays Swordsman in order to join forces with the Skrulls in order to bait Andrea into revealing herself to be a Skrull, she is shocked to realize that she is a clone, culminating in Bullseye murdering her in order to save Moonstone. With Norman's help, Moonstone blames the Skrulls for Andrea's murder. After the Thunderbolts defeat the Skrull armada attacking Washington, D.C., as well as two Super-Skrulls, Norman begins to play up his and his team's role to the media, making them appear to be the force that is saving the Earth from Tony Stark's bungling. The Thunderbolts then journey to New York where they reluctantly aid Earth's heroes in their battle against the Skrulls.


title: Thunderbolts (comics)

text: After Songbird gets mad at Osborn for leaving her to deal with a dangerous Skrull who nearly killed her, Norman Osborn and Moonstone come to the agreement to get rid of Songbird and the other Thunderbolts whose morals do not match their own. Their plan comes into effect while Norman is in Washington during the "Dark Reign" storyline, preparing to assume control over S.H.I.E.L.D. The group deports Radioactive Man back to China after Norman has his work visa revoked, and Moonstone has Penance sent to a corrupt maximum security mental institution where he would be held prisoner for the rest of his natural life. Moonstone then aids Bullseye and Venom in attempting to kill Songbird, with Bullseye and Moonstone inadvertently and indirectly revealing to Songbird that Norman has given them permission to kill her. Though she neutralizes Moonstone and Bullseye, Venom attacks Songbird on the "Zeus" minijet; she barely survives the crash. Bullseye recovers and sneaks up on her after the wreck, but the Swordsman saves her and tells her to run away, while blowing up the downed craft to forge her death in the explosion.


title: Thunderbolts (comics)

text: Swordsman confronts Norman with the revelation that he is not being offered a position on the "Dark Avengers" team and that Norman had never planned on fulfilling his vow to resurrect the Thunderbolt's deceased sister or to give him a full presidential pardon after his contract with the Thunderbolts ended. Norman ends the confrontation by stabbing and defenestrating him.


title: Thunderbolts (comics)

text: With Bullseye, Moonstone, and Venom being transferred to Norman's new team the Dark Avengers, Osborn decided to create a new Thunderbolts roster, one that would provide themselves as assassins for H.A.M.M.E.R.


title: Thunderbolts (comics)

text: One of their first missions is an attempt to kill Deadpool after he tries to blackmail Osborn for Norman's theft of Skrull data. After Deadpool and Taskmaster thwart the hit, the Thunderbolts then travel to Madripoor to recruit Mister X as a team member. Osborn then adds a new incarnation of Scourge, who Norman seems to know. Songbird eventually returns, and he has his team try to kill her. Yelena abandons the team after Scourge takes over, and reveals to Songbird that she is, in fact, the original Natasha Romanoff working for Nick Fury. The two escape, only to end up leading Osborn to Nick Fury. Osborn orders the Thunderbolts to kill Black Widow and Songbird, and shoots Fury himself; however, "Fury" is revealed to be a Life Model Decoy. After Black Widow and Songbird escape, Scourge is revealed to be the former Super-Soldier Nuke.


title: Thunderbolts (comics)

text: Norman Osborn later places Grizzly on the Thunderbolts team at the time they fight the Agents of Atlas. The fight between the Thunderbolts and the Agents of Atlas rages on as the deadly chemicals causes both teams to retreat. During the battle Scourge is brainwashed into shooting when he sees Norman Osborn. As the Thunderbolts return to the Cube on Zeus, their pilots informs that there is an important message from Osborn, Displayed holographically, Osborn informs the Thunderbolts that there is an important assignment he needs them to do in Broxton, Oklahoma, but does not disclose the details. Upon seeing Osborns' face, "Scourge's" programming kicks him and attempts to shoot. The bullet goes right through the hologram and hits Headsman right in the head, horrifying his teammates.


title: Thunderbolts (comics)

text: During the "Siege" storyline, Norman Osborn sends his Thunderbolts to infiltrate Asgard and steal a weapon from the Asgardian armory that will turn the tide in his favor.


title: Thunderbolts (comics)

text: The weapon that the Thunderbolts stole from the Asgardian armory was actually Odin's spear. They were stopped by the Mighty Avengers but not before Scourge used the spear to sever U.S. Agent's left limbs. Most of the team is apprehended by authorities while Paladin and Ant-Man escape.


title: Thunderbolts (comics)

text: A brand new team of Thunderbolts appears in the aftermath of "Siege" as seen during the "Heroic Age" storyline.


title: Thunderbolts (comics)

text: Captain Steve Rogers brings Luke Cage to the Raft to assemble a new team of Thunderbolts that will not be related to the ones that Norman Osborn previously assembled and rather be criminals who work towards redemption again. They recruit Ghost (who betrayed Osborn), Moonstone (despite objections by long-standing member Songbird), Juggernaut (who is vouched for by Professor X), Crossbones (who is supposed to gravitate the rest of the Thunderbolts to Cage), and Man-Thing (who serves as the Thunderbolts' mode of transportation). No longer wearing a costume and now stuck in a wheelchair, U.S. Agent is appointed the new warden of the Raft. During their first training session, Luke Cage is attacked by Baron Helmut Zemo, who declares that he will take leadership of the Thunderbolts. He tells the Thunderbolts that they can serve him if they can escape from the Raft. Though Crossbones is more than willing to oblige, the rest are suspicious. As it turns out, it was a setup to test the team and the Raft's security measures. They are then deployed to deal with a group of Asgardian trolls.


title: Thunderbolts (comics)

text: After killing two of the trolls and capturing the third (who turned out to be a girl named Gunna who was raised by trolls), the group is reprimanded by Cage for the death of the two trolls. The Thunderbolts are sent out to search for a group of missing S.H.I.E.L.D. agents, who turn out to be mutated by the Terrigen Mist.


title: Thunderbolts (comics)

text: After slaying the mutated S.H.I.E.L.D. agents and sealing off the cave, the Thunderbolts return to the Raft for a check-up. The check-up reveals that Crossbones has not been mutated by the Terrigen Mist and that MACH-V will be out for weeks. The facility is soon visited by the staff and students of Avengers Academy. After getting Thunderbolts introduced to the students, the prison's power grid suddenly shut down due to an EMP, allowing the Raft's inmates to escape. Together with Warden Walker (U.S. Agent) and the staff of Avengers Academy, the Thunderbolts manage to detain all escaped prisoners. Some of Hank Pym's students attempted to get to Norman Osborn, so Luke Cage reprimands them.


title: Thunderbolts (comics)

text: During the "Shadowland" storyline, the Thunderbolts are assigned with rescuing a young boy who is held prisoner by the Hand, and bringing down their stronghold. While fighting the ninjas, Fixer is stabbed, while Songbird is taken down, giving Moonstone the opportunity to do things her way.


title: Thunderbolts (comics)

text: Following the dischargement of Crossbones, the Thunderbolts end up getting Hyperion as a new member when it comes to fighting giant monsters. He betrays the Thunderbolts while they are split up, destroying Man-Thing's lower body and legs with his atomic vision and leaving Moonstone and Songbird to drown while he takes the com device which controls the nanites in each Thunderbolts member. Hyperion activates it on Juggernaut to test it after the latter had saved the two drowning members, and Juggernaut responds by fighting back. Moonstone and Ghost join in to help Juggernaut upon which they take down Hyperion. They let Man-Thing touch him after a fallen Hyperion shows fear in front of them. The Thunderbolts next recruit Satana as his replacement when Luke Cage and Doctor Strange discover that the Raft is susceptible to magic.


title: Thunderbolts (comics)

text: With recent events involving the Thunderbolts, the team's advisory committee approve the creation of a Beta team, the Underbolts. As the Thunderbolts head toward a new mission in Eastern Europe, Songbird, Mach V, and Fixer select candidates of Raft inmates, with the chosen candidates being Troll, Shocker, Centurius, Boomerang and Mister Hyde.


title: Thunderbolts (comics)

text: After the mission in Europe as seen during the "Fear Itself" storyline, Juggernaut gets infected with nanites by Cage, who erroneously thought that he will attack the others. Juggernaut is left at the Raft while the alpha and beta team went on to a mission in Iraq, as zombies emerge from the area. In that moment, a Worthy hammer falls inside the prison and is taken immediately by Juggernaut (who feels also the calling of Cytorak). The hammer ends up transforming Kuurth: Breaker of Stone who ends up destroying the Raft, which led to most of its inmates escaping. The alpha and beta teams are called back to the raft to capture and control inmates, and work to clear the rubble. During the process, Centurius manages to find the controls for the beta team's nanites and disable them. The beta team decides to wait for a better time to escape, as they reason that more heroes may be on the way to help the raft. Meanwhile, Moonstone restores order to the partially collapsed women's wing of the prison, as Ghost helps the warden avert a crisis in a section of the prison where a malfunctioning failsafe is suffocating inmates. Crossbones, an ex-member of the Thunderbolts, with the help of another inmate, escapes the raft in the chaos. After order is restored, the alpha team, including Songbird and Mach V, is sent to subdue Juggernaut/Kuurth, with the beta team and Fixer held in reserve. To the team's surprise, Man-Thing becomes uncontrollable, and teleports away before they fight Juggernaut. Satana explains that he has reached the "next stage" of his evolution, but before she can finish explaining, Juggernaut arrives. The team tries to free Juggernaut from the hammer's influence, but fail and he escapes.


title: Thunderbolts (comics)

text: Upon being sent back in time to the World War II era, the Thunderbolts members end up passing themselves off as the American Thunderbolts when they end up helping the World War II era's versions of Captain America and Namor. As Centurius adapts the name of Doc Century for this time period, the American Thunderbolts continue their fight with the Nazis and the Human Torch androids that were created by Baron Heinrich Zemo.


title: Thunderbolts (comics)

text: From then on the book focuses on the team in the present (Luke Cage, MACH-V, Songbird, Ghost) versus the Underbolts team in the past (Centurius, Satana, Troll, Boomerang, Moonstone, Fixer). Eventually, jumping around in time, the team encounter themselves as the original Thunderbolts (1997) and ends with the renaming of the title into "Dark Avengers" although the storyline carries through.


title: Thunderbolts (comics)

text: A new team composed of Red Hulk, Deadpool, Elektra, Agent Venom, and Punisher debuted as a part of Marvel NOW! major relaunch. Written by Daniel Way with drawings by Steve Dillon. This incarnation is not a government-sponsored team.


title: Thunderbolts (comics)

text: Red Hulk assembles his incarnation of the Thunderbolts to be a strike team that is close to "Code Red." He has gathered Deadpool, Elektra, Punisher, and Agent Venom because "their conditions cannot be cured." Red Hulk has his Thunderbolts do things that are similar to X-Force. Red Hulk has obtained Samuel Sterns' body and hooks it up to a machine that emits Red Gamma Radiation onto Samuel Sterns. Deadpool later finds Samuel Sterns' as a Red Leader with no apparent powers as he expresses his view that the Punisher won't be pleased. Punisher finds Red Leader's body and shoots him between the eyes disrupting whatever plans Red Hulk has for Red Leader.


title: Thunderbolts (comics)

text: Red Hulk brings the Thunderbolts along to deal with gamma technology on the island of Kata Jaya where the technology was supplied by Madman. During the fight against Madman, Red Hulk took the Leader to a pipe line that feeds Gamma energy to Madman's lab. Red Hulk then absorbed the energy himself and then forced fed it to the Leader, bringing him back to life. Red Leader is revived, but doesn't possess his prior intellect. They do get into the Madman's bunker to find out that The Leader hid his intellect on the internet. Madman confronts them and Leader promises to unlock his intellect to his brother, but after whispering one secret word his brother drops dead—his brain not being able to handle all of the information.


title: Thunderbolts (comics)

text: The team then follows leads for the supplies of the gamma radiation technology to Kabul to discover Elektra's brother Orestez Natchios is creating an army of Crimson Dynamos powered by gamma radiation. Elektra runs off with her brother and claims to have killed him, but Punisher realizes she was lying and kills him himself. During this fiasco, the Red Leader secretly gains access to the internet and downloads his intellect off the internet despite his body not being able to physically contain it (he suffers from profuse nose bleeds).


title: Thunderbolts (comics)

text: The Team decides that they can't do only missions for Red Hulk, as none truly trust him, so a system of give and take missions is developed, in which the team does a mission for General Ross, then a mission for a random member. The first random name pulled for a member is Punisher, who chooses a mission to take out a crime family in New York. The story arc takes place during the crossover event Infinity. It turns out the Leader purposefully tricked the team into letting him pick the next mission, since he had no skin in the game, so he can escape. He wasn't expecting an alien invasion. While trying to escape the team's submarine, Red Leader begins to get his brain siphoned by Supergiant of Thanos' Black Order, but is saved by Red Hulk. He reluctantly releases Mercy and is forced by Red Hulk to trick her into destroying the Black Order's flag ship. Afterwards, Red Leader starts having visions of how to kill every member of the Team, but determines that something is wrong with his plan, there was an unseen variable that stopped him from succeeding. That variable presents itself mainly in the form of Abigail Mercy, who joined Ross's Thunderbolts team as a condition for her release.


title: Thunderbolts (comics)

text: Ghost Rider joins the team in order to stop Mercy, who is getting too powerful to control, but inadvertently drags the team to hell. Leader negotiates a contract with Mephisto to put him back in power of the Underworld by overthrowing the current leader of hell, Strong Guy, who gave up his soul to save his love. Strong Guy throws the fight, so the Thunderbolts escape and Mercy is dragged to hell.


title: Thunderbolts (comics)

text: Deadpool's name gets pulled next for a mission and he wants to kidnap actor Ryan Reynolds, but Venom asks to be given the mission, as he is leaving the team. He wants to let the symbiote take over and have the team fight it, which they do, to reassure him that if he would lose control of the symbiote, there would be someone that could stop him.


title: Thunderbolts (comics)

text: The team then travels to a haunted swamp in Honduras to find an ancient artifact. Leader is the only one who can translate the ancient language. During the boat ride, Leader whispers something cryptic from an ancient language and Johnny Blaze loses his powers and is killed by a tentacled monster. Leader then betrays the team by promising their river guide to help ambush the team, but then doubles crosses him and alerts the team, who them quickly dispose of the ambushers. Leader then tricks the team into entering the secret temple via booby trapped entrance. Ross survives and finds the power of the temple, an infant celestial head. Leader attempts to drink the Celestial's blood, but this causes his brain to explode. One of Ross' former squadmates, Mancuso, is waiting for him and powered by the Celestial. Mancuso warns him that if Ross continues his selfish ways he will be forced to use his cosmic powers to kill him. Ross realizes that he pushes his team too hard and constantly endangers them, so Mansuco teleports them back to before the briefing. Ross decides not to travel to Honduras.


title: Thunderbolts (comics)

text: Much later, the team tracked down Doctor Faustus. The Punisher wanted to kill him for murdering all the children in a high school. Red Hulk stopped him and wanted to recruit Faustus, but the Punisher does not like the idea and quits the team. While going to the refrigerator at his safe house, the Punisher found a bomb with a note saying "You don't quit on us. You're fired." Angered, the Punisher believes Red Hulk set the bomb and began hunting down his former teammates. He snipers Leader and Faustus. He decapitates Deadpool, who is not angry and actually flattered that Frank found him a threat. He steals magic regalia from Bloodstone and depowers Ghost Rider, but Johnny Blaze is actually glad to be freed of the demon. Punisher can't bring himself to kill Elektra, but does confess that he killed her brother. She then admits she knew he would kill her brother. Punisher then uses a customized Hulkbuster armor with a special gun that depowers Red Hulk temporarily. During a gruesome hand-to-hand combat between Ross and Castle, Hawkeye and his new team of Avengers arrest both of them. It is later revealed that Red Leader was alive and set the bomb knowing the Punisher will come after each member of the Thunderbolts and set a Life Model Decoy to fool the other Thunderbolts. The Avengers and Thunderbolts return to Kata Jaya and arrest Leader who is their new dictator. Ross then decides to disband the team.


title: Thunderbolts (comics)

text: The annual takes place before the team disbands and follow the team as they work up with W.A.N.D., S.H.I.E.L.D.'s paranormal research department, in order to take down an out of control Doctor Strange.


title: Thunderbolts (comics)

text: The Winter Soldier formed his own Thunderbolts team following the events of the "" storyline. Besides Winter Soldier, the membership consisted of Fixer, Atlas, Moonstone, MACH-X, and Kobik. Winter Soldier, Kobik, and MACH-X helped Fixer, Atlas, and Moonstone evade S.H.I.E.L.D. In exchange, they agree to form the Thunderbolts with the mission to make sure that S.H.I.E.L.D. never uses the Kobik Project again.


title: Thunderbolts (comics)

text: Over the course of this 12-issue run, Bucky's Thunderbolts team clashes with the All-New Inhumans, Squadron Supreme, mysterious alien invaders and others. They also frequently test the limits of Kobik's terrifying, reality-bending cosmic powers. The girl is the one who magically manifests the team's arctic headquarters and even resolves Fixer's time travel-loop conundrum that has lingered since the "Thunderbolts Throughout Time" storyline.


title: Thunderbolts (comics)

text: Following the "Civil War II" event, the team reunites with Songbird as they rescue Bucky from an altered, secretly fascist Captain America. She remains with the team, briefly reuniting her romance with Abe Jenkins (MACH-X). In the lead up to the "Secret Empire" event, Zemo attacks the Thunderbolts' base with the latest incarnation of the Masters of Evil. As that confrontation begins to unfold, Jolt is transported back from Counter Earth just in time to save Atlas from arctic exposure.


title: Thunderbolts (comics)

text: Later, Zemo makes a nefarious offer the original Thunderbolts members: give in to their darker natures and join the new rise of Hydra... or die. Abe and Melissa reject the offer, but Fixer, Moonstone, and Atlas ultimately leave the ruins of the HQ with Zemo's Masters of Evil. In the aftermath of the destruction, Songbird scours the snowy horizon for Abe, who is presumed dead, while Ghost—who has been spying on the Thunderbolts for some time—rescues a pocket-sized Jolt, who has shrunk due to her power exertion during the battle with the Masters of Evil.


title: Thunderbolts (comics)

text: After losing custody of Frank Castle in the fledgling country of Bagalia, Baron Zemo and Jigsaw flee to New York City. There, Zemo approaches the Mayor Wilson Fisk, in hopes of joining forces and finally taking on revenge on the Punisher. Fearing a public relations disaster, Mayor Fisk reluctantly agrees—and is promptly dismayed when Zemo reveals a new team of Thunderbolts at a mayoral press conference.


title: Thunderbolts (comics)

text: Mayor Fisk and Citizen V announce to a cheering crowd that the Thunderbolts will work with newly arrived U.N. forces (who are, in fact, Zemo's disguised Hydra soldiers) and a covert team of NYPD officers to rid the city of Punisher.


title: Thunderbolts (comics)

text: Zemo and his team are able to surprise Frank Castle once, but are thwarted by Black Widow. Moonstone and Mayor Fisk ultimately goad an over-confident Zemo into confronting Punisher again and "Citizen V" musters a rousing speech to rally spooked Hydra agents. The Thunderbolts then clash with Castle again—but discover that the vigilante has reluctantly partnered with a team of other street-level operatives, including Black Widow, Moon Knight, Rachel Cole-Alves, and even Frank's old Thunderbolts teammate, Ghost Rider. The battle is fierce and even though Frank shoots down Fixer (or possibly a robotic copy) and corners "Citizen V," Jigsaw intervenes... and the Thunderbolts escape with Rachel as a hostage. As they flee, Zemo challenges Frank to come retrieve her... alone.


title: Thunderbolts (comics)

text: The ploy doesn't work. Punisher and Zemo battle at Fisk Tower. Before Frank can kill Zemo, Ghost intervenes. Calling Zemo "a third-rate fascist with an inherited army," Ghost tells the defeated Zemo to return to Bagalia and wire him his money. Punisher wards Ghost off and Zemo is able to meet Mayor Fisk on the roof, hoping to rally a comeback. Instead, Mayor Fisk tells Zemo that he's in over his head. At the touch of Ghost's hand, Zemo is seemingly vaporized. Fisk thanks Ghost and tells him that they'll discuss "the details of their arrangement" later.


title: Thunderbolts (comics)

text: When the symbiote god Knull encircles the planet in symbiote goo and sets out to destroy every living thing on Earth, Mayor Wilson Fisk decides to put together his own team of superpowered operatives to beat back Knull's creatures and save the city. He secretly meets with eight of these operatives, all known criminals who have been working in the city without Kingpin's blessing: