''Xylocopa virginica'', sometimes referred to as the eastern carpenter bee, extends through the eastern United States and into Canada. They are

sympatric In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species s ...

with ''

Xylocopa micans ''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 State ...

'' 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.

Description and identification

The bee is similar in size to

bumblebee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only Extant taxon, extant group in the tribe Bombini, though a few extinct r ...

s, but has a glossy, mostly black body with a slight metallic purple tint. ''X. virginica'' males and females have generally the same mass, but can be differentiated visually by the male's longer body and the female's wider head. The males also have a white spot on their face. Additionally, the males have larger thoracic volumes for given masses. Females of different social standing can also be told apart based on morphology. Primary females are larger than secondary or tertiary

females Female (symbol: ♀) is the sex of an organism that produces the large non-motile ova (egg cells), the type of gamete (sex cell) that fuses with the male gamete during sexual reproduction. A female has larger gametes than a male. Females ...

, and also have more mandibular and wing wear. ''X. virginica'' have distinctive maxillae that are adapted to performing perforations on

corolla Corolla may refer to: *Corolla (botany), the petals of a flower, considered as a unit *Toyota Corolla, an automobile model name *Corolla (headgear) A ''corolla'' is an ancient headdress in the form of a small circlet or crown.maxillae are sharp and wedge-shaped, allowing them to split the side of corolla tubes externally to access the nectar. Eastern carpenter bees also have galae on their maxillae that are shaped like large, flat blades. Bees with sharp galae can use these to further aid in penetrating the corolla tubes.

Taxonomy and phylogeny

''X. virginica'' belongs to the genus ''

Xylocopa Carpenter bees are species in the genus ''Xylocopa'' of the subfamily Xylocopinae. The genus includes some 500 bees in 31 subgenera. The common name "carpenter bee" derives from their nesting behavior; nearly all species burrow into hard plant ma ...

'', which consists of over 400 species worldwide, in the subgenus ''Xylocopoides'', which contains only 5 New World species, including ''

Xylocopa californica The California carpenter bee, ''Xylocopa californica'', is a species of carpenter bee in the order Hymenoptera, and it is native to western North America. Distribution There are approximately 400 species worldwide of the genus ''Xylocopa.''LeB ...

'', which also occurs in the U.S.

Distribution

''X. virginica'' is found throughout much of North America east of the

Rocky Mountains The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in straight-line distance from the northernmost part of western Canada, to New Mexico ...

and at least as far north as

Nebraska Nebraska () is a state in the Midwestern region of the United States. It is bordered by South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Kansas to the south; Colorado to the sout ...

, southern

Ontario Ontario ( ; ) is one of the thirteen provinces and territories of Canada.Ontario is located in the geographic eastern half of Canada, but it has historically and politically been considered to be part of Central Canada. Located in Central Ca ...

, and

Maine Maine () is a state in the New England and Northeastern regions of the United States. It borders New Hampshire to the west, the Gulf of Maine to the southeast, and the Canadian provinces of New Brunswick and Quebec to the northeast and nor ...

Nesting

''X. virginica'' build their nests in wood,

bamboo Bamboos are a diverse group of evergreen perennial flowering plants making up the subfamily Bambusoideae of the grass family Poaceae. Giant bamboos are the largest members of the grass family. The origin of the word "bamboo" is uncertain, ...

culms,

agave ''Agave'' (; ; ) is a genus of monocots native to the hot and arid regions of the Americas and the Caribbean, although some ''Agave'' species are also native to tropical areas of North America, such as Mexico. The genus is primarily known for ...

stalks, and other comparable materials, but they prefer to nest in milled

pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family (biology), family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanic ...

or cedar lumber. The nests are built by scraping wood shavings off of the wall. These shavings are then used to create partitions between nesting cells. The entrance cuts into the wood perpendicular to the grain, but they are built parallel beyond the entrance. These nests may be either social, containing groups of two to five females, or solitary. Social nests are more common, despite the fact that brood productivity is actually lower when females choose to nest together. Because ''X. virginica'' builds its nests in wood structures, it is common for it to nest in constructed furniture or buildings. ''X. virginica'' is the most common large carpenter bee in eastern North America, and it nests in small groups, so nests are fairly commonly encountered. The nests are usually round and typically have one to four tunnels. They have multiple branches, with each adult female living and laying eggs in a separate branch but females sharing one common entrance. Because the nests are costly to build, it is common for females to reuse old nests. Hibernaculum sweepings

Life cycle

In ''X. virginica,'' mating occurs only once a year, in the spring. Eggs are laid in July, starting farthest from the exit hole, and by about August and mid-September, larval development has completed and all the pupae have become adults. Researchers suggest that there is a mechanism that synchronizes the emergence time of young that are laid at different times by causing the younger eggs to develop faster. This mechanism prevents bees that would emerge sooner from removing their siblings and decreasing their potential competition. Bees that have newly emerged have a soft cuticle and white wings. The wings later transition to brown, then to a bluish black. They can fly 3–4 days after emergence, but they remain in their nest for at least two weeks, consuming nectar but not pollen. The juveniles begin the next mating cycle the following spring, so one generation develops in a year. Females begin to exhibit signs of senescence around July. The indicative behavior includes resting in flowers, remaining in the nest, or even just falling to the ground from flight. Older individuals also crawl, avoid taking flight, and do not struggle when handled by humans. The old bees die by early August, the same time that juveniles emerge from brood cells. Due to the simultaneous nature of expiration of old bees and emergence of new ones, there is little overlap between generations, except for some females that survive a second winter.

Behavior

''X. virginica'' is not a solitary bee species, but it is not truly social either. The weak form of sociality they exhibit, with one female doing the majority of the work and caring for her sisters, may be a transitional step in the evolution of sociality.

Dominance hierarchy

Female ''X. virginica'' can have solitary nests, but they usually nest in social groups. The social order of ''X. virginica'' is broken into three groups: primary, secondary, and tertiary. Primary females act as the dominant within a nest and are in charge of reproduction, providing food for the larvae, and laying all the eggs. This is different from many bee species in which there is a queen that focuses her energy solely on laying eggs while relying on provisions provide by subordinate bees. Secondary females may sometimes participate in

oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...

, and they reinforce this potential role by helping provide for the larvae or performing nest maintenance. Tertiary females rely on the provisions provided by primary females and quietly await overwintering while remaining inactive. Studies have shown that primary females are usually the bees that have overwintered twice, while tertiary bees have only overwintered once. Tertiary bees will most likely survive a second overwintering and develop further to become primary female the following year. Secondary bees may survive a second winter, but that is unlikely if they actively forage after their first

overwintering Overwintering is the process by which some organisms pass through or wait out the winter season, or pass through that period of the year when "winter" conditions (cold or sub-zero temperatures, ice, snow, limited food supplies) make normal act ...

Division of labor among the sexes

Not all females do the same work in a social nest. This is evident based on the varying levels of wear on the wings and

mandibles In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...

of females of various social standing. Although many nests have more than one female, there is a division of labor between the older and younger females. During nesting time, only the older females are responsible for nesting duties such as digging, excavating the cells, lining the cells, collecting food, and

ovipositing The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...

. Evidence of this activity can be found in their worn mandibles. Young females rarely leave the nest and guard the entrance while the older females work, resulting in unworn wings and mandibles in the younger females. Additionally, ''X. virginica'' is the only known species in which one-year-old females cohabit the nest with two-year-old females that do all the labor. Males often spend long periods of time hovering, flying, or in fast pursuit of intruders, while female flight activity is usually very directed, such as flights to flowers and food sites. Larger females have an advantage because they can carry larger amounts of pollen or nectar back to the nest and can fly longer distances. Xylocopa virginica pollinia

Diet

''X. virginica'' survive mostly on nectar and pollen. Newly emerged bees do not have food stored in their nest, but they are occasionally brought nectar. ''X. virginica'' use their maxillae to penetrate the corolla of plants and reach the nectar stores, a behavior known as

nectar robbing Nectar robbing is a foraging behavior utilized by some organisms that feed on floral nectar, carried out by feeding from holes bitten in flowers, rather than by entering through the flowers' natural openings. "Nectar robbers" usually feed in this ...

. This happens when the bee pierces the corollas of long-tubed flowers, thus accessing nectar without making contact with the

anthers The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...

and bypassing

pollination Pollination is the transfer of pollen from an Stamen, anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by Anemophily, wind. Pollinating agents can ...

. In some plants this reduces fruit production and seed number. In other plants, defensive mechanisms allow pollination to occur despite perforation of the corolla.

Mating behavior

Each nest usually has one mated individual. Mating occurs in April and is often accompanied by a bobbing dance that involves about a dozen males and only a few females. Males require female activity, specifically flight, in mating. Occasionally before mating, the couple will face each other and hover for a few minutes. When the male contacts the female, he mounts her back and attempts to push his abdomen under hers. Copulation then occurs, and it is almost always followed by more mating attempts. If, during copulation, the female lands, the couple will disengage and the male will hover waiting for the female to take flight again; however, although the males almost always disengage and pause copulation when the female lands, there have been instances recorded in which the males will hold on to the female with all six legs and flap his wings in an attempt to lift her back into the air. Larger males are usually more successful in mating. Because of their competitive advantage due to their size, males will likely claim a territory near female nest sites. Smaller males will stay at

foraging Foraging is searching for wild food resources. It affects an animal's fitness because it plays an important role in an animal's ability to survive and reproduce. Foraging theory is a branch of behavioral ecology that studies the foraging behavi ...

sites or other areas they think females may pass so they can mate with reduced competition.

Kin selection

Research has shown that, regardless of sex, ''X. virginica'' show more aggression toward non-nestmates than nestmates, indicating that they can recognize each other. By living in social groups with

inclusive fitness In evolutionary biology, inclusive fitness is one of two metrics of evolutionary success as defined by W. D. Hamilton in 1964: * Personal fitness is the number of offspring that an individual begets (regardless of who rescues/rears/supports th ...

, the bees can raise offspring with the help of the nest community rather than as a solitary effort. The ability of ''X. virginica'' to recognize nestmates allows primaries and secondaries to exclude tertiary bees from their nests. Tertiary bees are a burden on resources because they do not perform any useful activities, but they benefit from the food and shelter provided by the primary females.

Defense

Territorial behavior in males

Males will establish territories near an active nest entrance to protect the colony and seek mating opportunities. For males that are near the nest entrance, their boundaries are usually linear and several meters long. For males that are farther from the exit, their boundaries are usually in the shape of a square and shorter in length. Males can stay in one territory for as long as two weeks. Although they do most of their foraging and resting during the night, they take small breaks throughout the day as well. After these breaks, they often have to fight off intruders that have taken advantage of their absence. Flights near the nest are usually uniform and involve much hovering. Flights protecting a bee's territory can be as short as a few minutes, but may extend beyond an hour. Males will not react to another bee unless the other is flying at high speed. When other individuals hover near the nest, it is unlikely that the male will pursue, whereas if another male comes into a territory at a high speed, the territorial male will give chase. When males patrolling the entrance of a nest are confronted with either dead or living Eastern carpenter bees suspended from a thread and dangled within the male's territory, the male does not respond when the bee is suspended and motionless, whether it is living or dead—even though ''X. virginica'' are capable of recognizing other individuals of their species. However, when the suspended bee is released and allowed to fly in the male's territory or is swung through the territory on the thread, the territorial male pursues it.

Parasites

There is one common species of bombyliid flies known to parasitize the larvae of ''Xylocopa virginica'': ''

Xenox tigrinus The tiger bee fly, ''Xenox tigrinus'', is an insect of the family Bombyliidae (bee flies) found in the eastern United States and southern Ontario. It formerly went by the name ''Anthrax tigrinus''. The distinctive wing pattern may resemble tiger s ...

''.

Marking

Eastern carpenter bees have mandibular glands that are known to produce a marking chemical in ''X. hirsutissima'' that functions as a nest marker or for female attraction. The glands are present in both males and females, but they produce no marking substance. However, ''X. virginica'' does have a Dufour's gland that is used to deposit a scent on a flower immediately following nectar collection. The scent, composed of hydrocarbons and esters, discourages ''X. virginica'', as well as other bee species, from returning to that same flower.

Stinging

The male bee is unable to sting because the stinger is simply a modified ovipositor (which males lack by definition), though they will commonly approach human beings and buzz loudly around them or fly close to them. The female, on the other hand, is capable of stinging; while the pain level of these stings is not well-documented, researchers have testified that ''X. virginica'' will sting if roughly handled. As the stinger is not barbed, a female can sting multiple times.

Human importance

Agriculture

''X. virginica'' visits many different kinds of flowers in order to gather pollen and nectar to bring back to the nest for larvae. Most of the plants they visit are wild grown or grown for decorative value; however, they can be good pollinators of blueberry crops. Their active seasons are quite long, and they forage on a wide variety of plant species. Also, because the start of their activity season is dependent on temperature, it is easy for greenhouse workers to manipulate the beginning of foraging activity. However, in comparison to species such as the

honey bee A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosm ...

, their smaller nest makes them less powerful as pollinators.

Destructive behavior

Because ''X. virginica'' builds its nests in various types of wood, it presents the disadvantage of weakening wood in manmade structures. They are also able to produce an excrement upon exiting their tunnels that may splash on the sides of buildings and negatively affect the aesthetic appeal of that structure. However, when weighed against the benefits ''X. virginica'' have as pollinators, the costs of its destructive behavior are insignificant. ''X. virginica'' offer benefits in the form of pollination for fruits, vegetables, legumes, and flower crops. Although the pollination strengths of ''X. virginica'' are secondary to that of the bumblebees and honey bees, the contribution is great enough to overlook destructive tendencies. ''X. virginica'' avoid entrances that are stained white, which is a possible solution to keeping them out of unwanted areas.

References

External links

OSU Agricultural Extension Fact Sheet - HYG-2074-06

{{Taxonbar, from=Q2149448 Xylocopinae Hymenoptera of North America Insects of Canada Insects of the United States Fauna of the Eastern United States Fauna of the Great Lakes region (North America) Ecology of the Appalachian Mountains Insects described in 1771 Taxa named by Carl Linnaeus