Anthidium manicatum

This bee is in the Anthidium genus, known as the carder bees or potter bees, and consisting of about 80 species. Anthidium is part of the family Megachilidae, which includes the leafcutter bees and mason bees in addition to the carder bees. ==Description and identification==

Anthidium manicatum has a wingspan around , with a body length of about for females, and for males. The anterior sides of the tarsal segments of each leg of a female have fine, soft and small white-colored hairs. This pilosity occurs in most species of Anthidium in the Western Palearctic region. Males of A. manicatum resemble A. maculosum in appearance. The two species have similar spiniform pygidia, as well as largely rounded sixth sterna (although that of A. manicatum is more so). ==Distribution and habitat==

A. manicatum is native to parts of Europe, Asia, and North Africa. It is now also found in North America, South American countries such as Argentina, Brazil, Paraguay, and Uruguay, the Canary Islands, and New Zealand (where it was found to be established in 2006). This insect was accidentally introduced into the United States from Europe sometime prior to 1963, when it was discovered in New York. It has since spread from the northeastern U.S. and southeastern Canada across the United States to California, where it was first collected in 2007. This species' tendency to occupy ready-made nesting sites, usually movable objects, allows it to spread to new locations easily. In Europe, this species is normally found in gardens, fields, and meadows in the southern part of Wales and England, but is localized in other places within the United Kingdom, where they can be seen from May to September. A. manicatum is the only species of the genus Anthidium that can be found in England. ==Nesting==

, used by A. manicatum'' for nest construction. Being a member of the Anthidiini tribe of megachilid bees, A. manicatum engages in highly elaborate nesting behavior. These bees construct their nests in pre-existing cavities, using the trichomes of wooly plants such as lamb's ears (Stachys byzantina). Females of A. manicatum use their mandibles, which are sharply toothed, to remove trichomes from the stems and leaves of various plants. They then roll up the trichomes into a ball and bring them to a pre-existing cavity. Inside the cavity, the bees fashion the trichome ball into cells, where they deposit an egg and a provisioning mass consisting of nectar and pollen. The female creates several cells in a cavity. Once finished, she seals the entrance to the cavity with a terminal plug, which consists of inorganic and organic materials that she brings to the nest. Females tend to build their nests at high locations. This may be to minimize the nest's exposure to parasites and predators. This may also be to avoid nest usurpation by other females of A. manicatum. Materials Besides trichomes, other materials used by a female A. manicatum for building brood cells include mud, stones, resin, and leaves. Some of the plant materials that are collected are hydrophobic, a feature that may serve an antimicrobial function for the nest. Females smear a plant substrate, plant extrafloral trichome secretions, on brood cells. The primary material used to build the cells are plant hairs, or "wool" (hence the name "wool carder bee"), that is collected from the stems and leaves of plants. Females largely use the hairs of Lamiaceae, especially those of the genus Betonica or Stachys. Additionally, females use specialized hair-like structures on the exterior of their tarsi to absorb the secretions of the plant hairs to apply onto the brood cells. These secretions are obtained from different species, such as Anthirrinum, Crepis, and Pelargonium. ==Behavior==

Mating behavior The mating system of A. manicatum is unlike those of most other bees. Females exhibit polyandry and continuously mate throughout their reproductive lives. Males exhibit resource defense polygyny. A. manicatum displays extreme polyandry, which is linked to male territoriality and resource defence of flowering plants. Males claim patches of floral plants, aggressively ward off conspecific males, bees, and other resource competitors, and mate with the females that forage in their territories. Copulations occur repeatedly and regularly in both sexes. Males that are unable to defend their own territory (usually because of their relatively small size) use an alternative "sneaking" tactic. These unfit males receive fewer copulation opportunities than females. Such mating and territorial behaviour in bees has also been observed in Anthidiellum notatum, Anthidiellum perplexum, and Anthidium banningense. However, males of the genus Anthidiellum chase away intruders rather than physically attacking them, so their aggressive behaviour differs significantly. Resource defence and aggression The territorial mating behaviour of males of A. manicatum occurs when foraging resources are amassed, allowing for monopolization and defence of territories by individual males. They will also defend conspecific females, Late male sperm precedence Apparently, no selection pressure exists on males of A. manicatum to be the first to copulate, hence no (or very little) selection pressure arises for them to emerge before females. This is due to the females' polyandrous behaviour, and can also be attributed to the phenomenon of late male sperm precedence. Patterns of sperm use by the females determine the benefits of resource defence for males. If the female uses only the sperm from her first copulation in a breeding season, then selection will not favor the 'sit-and-wait' strategy of resource defence for the males over strategies that are pre-emptive, such as patrolling nest sites. However, if delayed mating can still ensure a high probability of procreating, then the resource defence strategy will be favoured. ==Sexual dimorphism==

Unlike most other species of Hymenoptera, male A. manicatum bees are larger than females in size, displaying male-biased sexual dimorphism. The selection for larger size in males may have resulted due to their aggressive territorial behavior and subsequent differential mating success. Although it does not tear other males apart, the abdominal armature of a male A. manicatum may have been developed due to territorial aggression, rather than for mating purposes. Hence the number of copulations a male obtains is positively correlated with territory quality as well as male size. ==Subspecies==

The following subspecies are recognized: • A. m. barbarum Lepeletier, 1841 • A. m. gribodoi Schwarz and Gusenleitner, 2003 • A. m. manicatum (Linnaeus, 1758) ==References==