Anthidium manicatum | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hymenoptera |
Family: | Megachilidae |
Genus: | Anthidium |
Species: | A. manicatum |
Binomial name | |
Anthidium manicatum | |
Synonyms | |
See text |
Anthidium manicatum, commonly called the European wool carder bee, [1] is a species of bee in the family Megachilidae, the leaf-cutter bees or mason bees. [2]
They get the name "carder" from their behaviour of scraping hair from leaves [3] such as lamb's ears ( Stachys byzantina ). They carry this hair bundled beneath their bodies to be used as a nest lining. [4] Like other members of the tribe Anthidiini, these bees do not cut leaves or petals as is typical for megachilids. [5] The males engage in territorial behaviour, aggressively chasing other males and pollinators from their territory. [6]
This bee is native to Europe, Asia, and North Africa. It has recently been seen in regions of South America, New Zealand, and the Canary Islands. They are generalists, and do not seem to prefer any plant genera for foraging, although in New Zealand, native plants are visited less frequently by this bee species. [7]
It was accidentally introduced into North America from Europe some time in the mid-20th century, first seen in 1963 near Ithaca, New York, and since then an invasive pest. [6]
This species belongs to the family Megachilidae and 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. [8]
Anthidium manicatum is originally an Old World bee. It has a wingspan around 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. [9] The males are substantially larger than females. [1]
A. manicatum bees are black and covered with yellow-grey hairs. Their faces and abdomens are covered in yellow spots. A male A. manicatum has 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 the lateral margins. [10] This feature distinguishes male A. manicatum from New World Anthidium species. [11] A yellow spot is on each side of the segments except the seventh. A second pair of spots is often seen on the disks of the fourth and fifth segments. A spine occurs on each side of the sixth and seventh segments at the apex, the seventh having a third thin spine in the middle. The legs exhibit variegation of yellow and are covered with grey hairs. A female A. manicatum is smaller in size than the male, but has 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. [10] 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. [12]
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). [11]
A. 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. [13] [14] As of 2006, this species is also now established in New Zealand. [15]
This insect was accidentally introduced into the United States from Europe sometime prior to 1963, when it was discovered in New York. [16] 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. [17] This species' tendency to occupy ready-made nesting sites, usually movable objects, allows it to spread to new locations easily. [11]
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, [4] where they can be seen from May to September. [4] A. manicatum is the only species of the genus Anthidium that can be found in England. [18]
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. 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. [19]
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, the bundle is used as a lining for their nest cavities. [1] [20]
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. [19]
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 . [21]
The mating system of A. manicatum is unlike those of most other bees. Females exhibit polyandry and continuously mate throughout their reproductive lives. [22] The interval of time between copulations amongst different males can be as short as 35 seconds in length. [23] 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. [22]
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. [24]
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. [22] Resource defence, as exhibited by male A. manicatum bees, has been thought to benefit females by reducing foraging competition for pollen and nectar. [22] The rate a female visits a territory is highly correlated with the number of flowers in that area. [23] If females forage in sites that are being defended by males and the cost of additional matings is low for female members, then male resource defence and female polyandry may co-evolve. [22]
Males of A. manicatum display highly aggressive behaviour, [25] both to conspecifics and other visitors to the flowers in its territory. [1] In the process of obtaining and defending flowering plant territories, males regularly patrol their territories and attack conspecific males and heterospecific pollinators. This typically consists of brief, aggressive "tackles", but they sometimes lethally injure the pollinators as a result of their aggression. [19] The territorial behaviour of males develops after a period of flight without any localisation in search of a suitable flower patch. [25] They will also defend conspecific females, [14] although they do harass them by holding them immobile and repeatedly attempting to mate. [2]
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. [26] This is due to the females' polyandrous behaviour, and can also be attributed to a phenomenon called "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. [22]
'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. Although females do have a spermatheca for long-term sperm storage, more recent mates tend to father the brood she produces. 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 defence in A. manicatum males. [22]
A. manicatum bees consume 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 [1] with Old World origins. [1] Both males and females can maintain a precise static hover near flowers similar to flies in the family Syrphidae. [27]
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. [25] Male size has been found to correlate with mating success. Smaller males implement alternative mating tactics if they cannot defeat larger males for territory. [19]
The mating behavior of a male A. manicatum can be determined by its relative body size compared 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). [28] Hence the number of copulations a male obtains is positively correlated with territory quality as well as male size. [22]
Behavioral ecology, also spelled behavioural ecology, is the study of the evolutionary basis for animal behavior due to ecological pressures. Behavioral ecology emerged from ethology after Niko Tinbergen outlined four questions to address when studying animal behaviors: What are the proximate causes, ontogeny, survival value, and phylogeny of a behavior?
Megachile rotundata, the alfalfa leafcutting bee, is a European bee that has been introduced to various regions around the world. As a solitary bee species, it does not build colonies or store honey, but is a very efficient pollinator of alfalfa, carrots, other vegetables, and some fruits. Because of this, farmers often use M. rotundata as a pollination aid by distributing M. rotundata prepupae around their crops. Each female constructs and provisions her own nest, which is built in old trees or log tunnels. Being a leafcutter bee, these nests are lined with cut leaves. These bees feed on pollen and nectar and display sexual dimorphism. This species has been known to bite and sting, but it poses no overall danger unless it is threatened or harmed, and its sting has been described as half as painful as a honey bee's.
Megachilidae is a cosmopolitan family of mostly solitary bees. Characteristic traits of this family are the restriction of their pollen-carrying structure to the ventral surface of the abdomen, and their typically elongated labrum. Megachilid genera are most commonly known as mason bees and leafcutter bees, reflecting the materials from which they build their nest cells ; a few collect plant or animal hairs and fibers, and are called carder bees, while others use plant resins in nest construction and are correspondingly called resin bees. All species feed on nectar and pollen, but a few are kleptoparasites, feeding on pollen collected by other megachilid bees. Parasitic species do not possess scopae. The motion of Megachilidae in the reproductive structures of flowers is energetic and swimming-like; this agitation releases large amounts of pollen.
Mason bee is a name now commonly used for species of bees in the genus Osmia, of the family Megachilidae. Mason bees are named for their habit of using mud or other "masonry" products in constructing their nests, which are made in naturally occurring gaps such as between cracks in stones or other small dark cavities. When available, some species preferentially use hollow stems or holes in wood made by wood-boring insects.
Osmia cornifrons, also known as the horned-face bee, is a species of solitary bee indigenous to Northern Asia. Physically, this species of bee is recognized for its horn-like extensions originating from its lower face. Populations of O. cornifrons have been recorded in multiple locations, including Japan, Korea, China, and Russia. O. cornifrons are more docile as compared to other species of bees and are less prone to sting when aggravated.
Xylocopa virginica, sometimes referred to as the eastern carpenter bee, extends through the eastern United States and into Canada. They are sympatric with Xylocopa micans in much of southeastern United States. They nest in various types of wood and eat pollen and nectar. In X. virginica, dominant females do not focus solely on egg-laying, as in other bee species considered to have "queens". Instead, dominant X. virginica females are responsible for a full gamut of activities including reproduction, foraging, and nest construction, whereas subordinate bees may engage in little activity outside of guarding the nest.
Anthidium is a genus of bees often called carder or potter bees, who use conifer resin, plant hairs, mud, or a mix of them to build nests. They are in the family Megachilidae which is cosmopolitan in distribution and made up of species that are mostly solitary bees with pollen-carrying scopa that are only located on the ventral surface of the abdomen. Other bee families have the pollen-carrying structures on the hind legs. Typically species of Anthidium feed their brood on pollen and nectar from plants. Anthidium florentinum is distinguished from most of its relatives by yellow or brick-red thoracic bands. They fly all summer and make the nests in holes in the ground, walls or trees, with hairs plucked from plants.
Osmia bicornis is a species of mason bee, and is known as the red mason bee due to its covering of dense gingery hair. It is a solitary bee that nests in holes or stems and is polylectic, meaning it forages pollen from various different flowering plants. These bees can be seen aggregating together and nests in preexisting hollows, choosing not to excavate their own. These bees are not aggressive; they will only sting if handled very roughly and are safe to be closely observed by children. Females only mate once, usually with closely related males. Further, females can determine the sex ratio of their offspring based on their body size, where larger females will invest more in diploid females eggs than small bees. These bees also have trichromatic colour vision and are important pollinators in agriculture.
Xylocopa sonorina, the valley carpenter bee or Hawaiian carpenter bee, is a species of carpenter bee found from western Texas to northern California, and the eastern Pacific islands. Females are black while males are golden-brown with green eyes.
Anthidium florentinum, one of several European wool carder bees, is a territorial species of bee in the family Megachilidae, the leaf-cutter, carder, or mason bees.
Amegilla dawsoni, sometimes called the Dawson's burrowing bee, is a species of bee that nests by the thousands in arid claypans in Western Australia. It is a long tongued bee, of the tribe Anthophorini and genus Amegilla, the second largest genus in Anthophorini.
The California carpenter bee or Western carpenter bee, Xylocopa californica, is a species of carpenter bee in the order Hymenoptera, and it is native to western North America.
Anthidium oblongatum, the oblong woolcarder bee, is a species of bee in the family Megachilidae, the leaf-cutter, carder, or mason bees. It is native to Eurasia and north Africa, and has also been introduced to North America.
Anthidium maculosum is a species of bee in the family Megachilidae, the leaf-cutter, carder, or mason bees. It is a solitary bee where the males are territorial and the females take part in polyandry. The males of A. maculosum differ from most other males of bee species because the males are significantly larger than females. In addition, subordinate males that act as satellites are smaller than territory-owning males. This species can be found predominately in Mexico and the United States.
Lasioglossum zephyrus is a sweat bee of the family Halictidae, found in the U.S. and Canada. It appears in the literature primarily under the misspelling "zephyrum". It is considered a primitively eusocial bee, although it may be facultatively solitary. The species nests in burrows in the soil.
Polygyny is a mating system in which one male lives and mates with multiple females but each female only mates with a few males. Systems where several females mate with several males are defined either as promiscuity or polygynandry. Lek mating is frequently regarded as a form of polygyny, because one male mates with many females, but lek-based mating systems differ in that the male has no attachment to the females with whom he mates, and that mating females lack attachment to one another.
Eulaema meriana is a large-bodied bee species in the tribe Euglossini, otherwise known as the orchid bees. The species is a solitary bee and is native to tropical Central and South America. The male collects fragrances from orchid flowers, which it stores in hollows in its hind legs. Orchids can be deceptive by mimicking the form of a female and her sex pheromone, thus luring male bees or wasps. Pollination will take place as the males attempt to mate with the labellum, or the tip petal of the flower. Male E. meriana are territorial and have a particular perch on a tree trunk where it displays to attract a female. After mating, the female builds a nest with urn-shaped cells made with mud, feces, and plant resin, and provisions these with nectar and pollen before laying an egg in each. These bees also have complex foraging and wing buzzing behaviors and are part of a mimicry complex.
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Euglossa imperialis is a bee species in the family Apidae. It is considered to be one of the most important pollinators to many Neotropical orchid species in mainland tropical America. It is also one of the most common non-parasitic euglossine species in lowland Panama. E. imperialis, unlike many other bee species, is not a social bee in the sense that there is no apparent morphological or physiological division within the species to distinguish individual bees to be part of a worker or reproductive caste.
Xylocopa micans, also known as the southern carpenter bee, is a species of bee within Xylocopa, the genus of carpenter bees. The southern carpenter bee can be found mainly in the coastal and gulf regions of the southeastern United States, as well as Mexico and Guatemala. Like all Xylocopa bees, X. micans bees excavate nests in woody plant material. However, unlike its sympatric species Xylocopa virginica, X. micans has not been found to construct nest galleries in structural timbers of building, making it less of an economic nuisance to humans. Carpenter bees have a wide range of mating strategies between different species. The southern carpenter bee exhibits a polymorphic mating strategy, with its preferred method of mating changing as the season progresses from early spring to mid summer. Like most bees in its genus, the southern carpenter bee is considered a solitary bee because it does not live in colonies.
The European wool-carder bee was first discovered in North America in 1963 near Ithaca, New York, and since then, its impact has been felt from coast to coast.