Anthidium manicatum

Anthidium manicatum
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Megachilidae
Genus:	Anthidium
Species:	A. manicatum
Binomial name
Anthidium manicatum (Linnaeus, 1758)
Synonyms
See text

Anthidium manicatum, commonly called the European wool carder bee, is a species of bee in the family Megachilidae, the leaf-cutter bees or mason bees.

They get the name 'carder' from their behaviour of scraping hair from leaves such as lamb's ears (Stachys byzantina). They carry this hair bundled beneath their bodies to be used as a nest lining. Like other members of the tribe Anthidiini, these bees do not cut leaves or petals as is typical for megachilids.

Taxonomy and phylogeny

This species belongs to the order Hymenoptera, which consists of organisms such as ants, bees and wasps, and the superfamily Apoidea, which is more specific to bees and wasps.

Description and identification

Detail of a male Anthidium manicatum

Anthidium manicatum is originally an Old World bee. It has a wingspan of approximately 20 millimetres (0.79 in), with a body length of about 11–13 mm (0.43–0.51 in) for females, and 14–17 mm (0.55–0.67 in) for males. The males are substantially larger than females.

A. manictum are black and covered with yellow-grey hairs. Their faces and abdomen are covered in yellow spots. Male A. manicatum have a black head and thorax, coated with short yellowish brown hairs. The cheeks below the antennae, a small spot behind each eye, a bilobate spot on the clypeus, and the mandibles (except the apex) are yellow. The wings are dusky in color. The abdomen is black with grey hairs, with a band of brown hairs at each segment’s apex as well as along their lateral margins. There is a yellow spot on each side of the segments except the 7th. A second pair of spots is often seen on the disks of the 4th and 5th segments. There is a spine on each side of the 6th and 7th segments at the apex, the 7th having a third thin spine in the middle. The legs exhibit variegation of yellow and are covered with grey hairs. Female A. manicatum are smaller in size than the males, but have a similar color pattern. The abdominal spots are smaller and the apex is rounded. Female legs are almost completely black, with very small yellow spots. The anterior sides of the tarsal segments of each leg of female A. manicatum have fine, soft and small white-colored hairs. This pilosity occurs in most species of Anthidium in the Western Palearctic region.

Distribution and habitat

Anthidium manicatum is found in parts of Europe, Asia, North Africa, and North America. It has also recently been documented in the Canary Islands, and South American countries such as Argentina, Brazil, Paraguay, and Uruguay. This species is now present in New Zealand.

This insect was accidentally introduced into the United States from Europe sometime prior to 1963, when it was discovered in New York State. 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.

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

Being a member of the Anthidiini tribe of megachilid bees, A. manicatum engages in highly elaborate nesting behavior. These bees construct their nests in preexisting cavities, using the trichomes of wooly plants. Female 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 preexisting cavity. Inside the cavity, the bees fashion the trichome ball into cells, where they deposit an egg as well as 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 in order to minimize the nest’s exposure to parasites and predators. This may also be to avoid nest usurpation by other female A. manicatum.

Materials

Besides trichomes, other materials used by 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 anti-microbial 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 species Betonica and 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.

Behaviour

Anthidium manicatum asleep in a flower

Males are highly aggressive against other males of this species, as well as other visitors to the flowers in its territory. They will also defend Anthidium manicatum females, although they do harass them by holding them immobile and repeatedly attempting to mate.

Females collect "down" from such plants as lamb's ears (Stachys byzantina). They scrape the hairs from the leaves and carry them back to their nests bundled beneath their bodies. There it is used as a lining for their nest cavities. Both males and females hover near flowers similar to flies in the family Syrphidae.

When females mate only once in their lifetime, there is an increased selection pressure amongst males to be the first to locate receptive females. Because of this, behavioral adaptations have evolved in males of other species, such as digging through soil in search of emerging females. As such, when females are receptive to males shortly after emergence there is a selection pressure on males to find females immediately after their emergence. However, female A. manicatum mate repeatedly, not just once. Males copulate with the same female repeatedly, both during the same visit as well as subsequent visits by the female to his territory. Studies show that in A. manicatum, there appears to be no selection pressure on the males to be the first to copulate, and hence no (or very little) selection pressure to emerge before females.

Mating Behavior

The mating system of A. manicatum is unlike those of most other bees. Females exhibit polyandry and continuously mate throughout their reproductive life. The interval of time between copulations amongst different males can be as short as 35 seconds in length. Males exhibit resource defense polygyny. A. manicatum display extreme polyandry, which is linked to male territoriality and resource defense of flowering plants. Males claim patches of floral plants, ward off conspecific males and other resource competitors, and mate with the females who 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) utilize an alternative ‘sneaking’ tactic. These unfit males receive fewer copulation opportunities than females.

Resource defense

The territorial mating behavior of male A. manicatum occurs when foraging resources are amassed, allowing for monopolization and defense of territories by individual males.

Resource defense, as exhibited by male A. manicatum, has been thought to benefit females by reducing foraging competition for pollen and nectar. If females forage in sites that are being defended by males and the cost of additional matings is low for them, then male resource defense and female polyandry may coevolve.

Male aggression

Male A. manicatum display highly aggressive behavior. In the process of obtaining and defending flowering plant territories, they frequently attack conspecific males and heterospecific pollinators. A. manicatum sometimes lethally injure the pollinators as a result of their aggressiveness. The territorial behavior of males develops after a period of flight without any localization.

Unlike most other species of Hymenoptera, male A. manicatum are larger than females in size. 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.

Late male sperm precedence

Patterns of sperm use by the females determine the benefits of resource defense 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 defense 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 defense strategy will be favored.

‘Late male sperm precedence’, or LMSP, is an experimental situation in which two males mate in sequence with a female and the second male will hold paternity of over 50% of the female’s offspring. This phenomenon might be caused by a variety of mechanisms, including sperm digestion or removal by the female, removal by the next male, or stratification of sperm from different males in the sperm storage organ of the female (i.e. the last sperm in is the first to come out). Studies have shown that A. manicatum males that copulate late in a sequence have a greater than average chance of paternity of the female’s eggs. Late male sperm precedence may have fostered the evolution of resource defense in A. manicatum males.

Sexual size dimorphism

A. manicatum displays male-biased sexual size dimorphism, which is uncommon amongst bees. Male size has been found to correlate with mating success. Smaller males will implement alternative mating tactics if they cannot defeat larger males for territory.

The mating behavior of male A. manicatum can be determined by relative body size to other conspecific males. Territory owners are larger in size than wanderers, and copulate with females more frequently as well. The number of copulations a male territory owner can achieve varies based on the size of the territory – males with larger territories generally achieve more copulations than those with smaller territories. This difference may be due to the energy expenditure of smaller bees to defend their territory from larger males, the inability of small males to mate on their territory, or female choice of larger males (regardless of territory size). Hence the number of copulations a male obtains is positively correlated with territory quality as well as male size.

Diet

Anthidium manicatum consumes the pollen from flowers of varying families. They are thus considered to be generalists. They visit garden flowers and weeds preferring blue flowers that have long throats with Old World origins.

Subspecies [1]

• Anthidium manicatum manicatum (Linnaeus, 1758)
• Anthidium manicatum barbarum Lepeletier, 1841
• Anthidium manicatum gribodoi Schwarz and Gusenleitner, 2003

Synonyms [2]

• Apis manicata Linnaeus, 1758
• Apis pervigil Harris, 1776
• Apis maculata Fabricius, 1781
• Apis fulvipes de Villers, 1789 (homonym)
• Apis modesta Christ, 1791
• Apis amoenita Christ, 1791
• Apis uncata Schrank, 1802
• Anthidium maculatum Latreille, 1806 (homonym)
• Anthidium marginatum Latreille, 1809
• Anthidium obtusatum Lepeletier, 1841
• Anthidium productum Lepeletier, 1841
• Anthidium manicatum var. nigrithorax Dalla Torre, 1877
• Anthidium manicatum var. fasciatum Schirmer, 1915
• Anthidium manicatum var. nasicolle Friese, 1917
• Anthidium manicatum var. luteus Gribodo, 1924 (homonym)
• Anthidium manicatum subcrenulata Alfken, 1930
• Anthidium manicatum cyrenaica van der Zanden, 1992 (homonym)

See also

• Pollinator
• Stingless Bees
• Mason Bees

Further reading

• Ascher, John S. (2001). "Hylaeus hyalinatus Smith, a European bee new to North America, with notes on other adventive bees (Hymenoptera: Apoidea)". Proceedings of the Entomological Society of Washington. 103: 184–190.
• Donovan, B. J. (2007). "Apoidea (Insecta: Hymenoptera)". Fauna of New Zealand. Vol. 57.
• Donovan, B. J., G. Burnip & B. McCarthy (2006). "The wool-carder bee, Anthidium manicatum, from the Old World, is newly established in New Zealand". New Zealand Beekeeper. 14 (5): 11–12, 25–26.{{cite journal}}: CS1 maint: multiple names: authors list (link)
• Hoebeke, E. R. & A. G. Wheeler, Jr. (1999). "Anthidium oblongatum (Illiger): an Old World bee (Hymenoptera: Megachilidae) new to North America, and new North American records for another adventive species, A. manicatum (L.)". Special Publication. 24. University of Kansas Natural History Museum: 21–24.{{cite journal}}: CS1 maint: multiple names: authors list (link)
• Payette, A. (2001). "Première mention de l'abeille adventice Anthidium manicatum (Linné) (Hymenoptera : Megachilidae) pour le Québec". Fabreries. 26: 87–97.
• Severinghaus Lucia Liu, Barbara Harris Kurtak & George C. Eickwort (1981). "The reproductive behavior of Anthidium manicatum (Hymenoptera: Megachilidae) and the significance of size for territorial males". Behavioral Ecology and Sociobiology. 9 (1): 51–58. doi:10.1007/BF00299853. JSTOR 4599410.
• Smith, I. (1991). "Anthidium manicatum (Hymenoptera: Megachilidae), an interesting new Canadian record". Proceeding of Entomological Society of Ontario. 122: 105–108.
• Starks, P. T. & H. K. Reeve (1999). "Condition-based alternative reproductive tactics in the wool-carder bee, Anthidium manicatum" (PDF). Ethology Ecology & Evolution. 11: 71–75. doi:10.1080/08927014.1999.9522843.
• Wirtz, Peter, Michael Szabados, Horst Pethig & John Plant (1988). "An extreme case of interspecific territoriality: male Anthidium manicatum (Hymenoptera: Megachilidae) wound and kill intruders". Ethology. 78 (2): 159–167. doi:10.1111/j.1439-0310.1988.tb00227.x.{{cite journal}}: CS1 maint: multiple names: authors list (link)
• Zavortink, T. J. & S. S. Shanks (2008). "Anthidium manicatum (Linnaeus) (Hymenoptera: Megachilidae) in California". Pan-Pacific Entomologist. 84 (3): 238–241. doi:10.3956/2007-47.1.

External links

• Images
• Information on leafcutter bees from Colorado State University