All honey bees live in colonies where the worker bees will sting intruders as a form of defense, and alarmed bees will release a pheromone that stimulates the attack response in other bees. The different species of honey bees are distinguished from all other bee species (and virtually all other Hymenoptera) by the possession of small barbs on the sting, but these barbs are found only in the worker bees. The sting and associated venom sac are also modified so as to pull free of the body once lodged (autotomy), and the sting apparatus has its own musculature and ganglion which allow it to keep delivering venom once detached. The worker bee dies after the stinger is torn from its body. As with other forms of life, warnings are given before an attack is launched. In the case of some honey bees species in the wild, this takes the form of a ‘Mexican wave’ which spreads as a ripple across a layer of bees densely packed on the surface of a comb when a threat is perceived, and consists of bees momentarily arching their bodies and flicking their wings.
It is presumed that this complex apparatus, including the barbs on the sting, evolved specifically in response to predation by vertebrates, as the barbs do not usually function (and the sting apparatus does not detach) unless the sting is embedded in fleshy tissue. While the sting can also penetrate the flexible exoskeletal joints in appendages of other insects (and is used in fights between queens), in the case of Apis cerana defense against other insects such as predatory wasps is usually performed by surrounding the intruder with a mass of defending worker bees, who vibrate their muscles so vigorously that it raises the temperature of the intruder to a lethal level. It was previously thought that the heat alone was responsible for killing intruding wasps, but recent experiments have demonstrated that it is the increased temperature in combination with increased carbon dioxide levels within the ball that produces the lethal effect. This phenomenon is also used to kill a queen perceived as intruding or defective, an action known to beekeepers as balling the queen, named for the ball of bees formed.
A honey bee that is away from the hive foraging for nectar or pollen will rarely sting, except when stepped on or roughly handled. Honey bees will actively seek out and sting when they perceive the hive to be threatened, often being alerted to this by the release of attack pheromones.
Although it is widely believed that a worker honey bee can sting only once, this is a partial misconception: although the stinger is in fact barbed so that it lodges in the victim’s skin, tearing loose from the bee’s abdomen and leading to its death in minutes, this only happens if the skin of the victim is sufficiently thick, such as a mammal’s. The bee’s sting is speculated to have evolved for inter-bee combat between members of different hives, and the barbs serve to improve penetration of the chitinous plates of another insect’s exoskeleton. When bees sting elastic-skinned mammals, the barbs become a hazard to the bees as described above. Honey bees are the only hymenoptera with a strongly barbed sting, though yellow jackets and some other wasps have small barbs.
The sting’s injection of apitoxin into the victim is accompanied by the release of alarm pheromones, a process which is accelerated if the bee is fatally injured. Release of alarm pheromones near a hive or swarm may attract other bees to the location, where they will likewise exhibit defensive behaviors until there is no longer a threat, typically because the victim has either fled or been killed. (Note: A true swarm is not hostile; it has deserted its hive and has no comb or young to defend.) These pheromones do not dissipate or wash off quickly, and if their target enters water, bees will resume their attack as soon as it leaves the water.
The larger drone bees, the males, do not have stings. The female worker bees are the only ones that can sting, and their stinger is a modified ovipositor. The queen bee has a smooth stinger and can, if need be, sting skin-bearing creatures multiple times, but the queen does not leave the hive under normal conditions. Her sting is not for defense of the hive; she only uses it for dispatching rival queens, ideally before they can finish pupating. Queen breeders who handle multiple queens and have the queen odor on their hands are sometimes stung by a queen.
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