Dolichovespula maculata

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Bald-faced hornet
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Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Vespidae
Subfamily: Vespinae
Genus: Dolichovespula
Species:
D. maculata
Binomial name
Dolichovespula maculata
Synonyms
  • Vespa maculata

Dolichovespula maculata is a species of wasp in the genus Dolichovespula and a member of the eusocial, cosmopolitan family Vespidae. It is known by many colloquial names, primarily bald-faced hornet, but also including bald-faced aerial yellowjacket, bald-faced wasp, bald hornet, white-faced hornet, blackjacket, white-tailed hornet, spruce wasp, and bull wasp. Technically a species of yellowjacket wasp, it is not one of the true hornets, which are in the genus Vespa . Colonies contain 400 to 700 workers, the largest recorded colony size in its genus, Dolichovespula. [1] It builds a characteristic large hanging paper nest up to 58 cm (23 in) in length. Workers aggressively defend their nest by repeatedly stinging invaders. [2]

Contents

The bald-faced hornet is distributed throughout the United States and southern Canada, but is most common in the Southeastern United States. Males in this species are haploid and females are diploid. Worker females can, therefore, lay eggs that develop into males.

Taxonomy and phylogenetics

The bald-faced hornet gets its name from the characteristic white markings on its face, as the word "bald" in English is derived from the word "piebald". [3] It was first described by Carl Linnaeus in his 1763 Centuria Insectorum . D. maculata is part of the cosmopolitan family Vespidae, in the genus Dolichovespula. Its black and white coloring differentiates it from its mostly black and yellow congenerics. [1]

Description

The bald-faced hornet is distinguished from other yellowjackets by its striking black-and-white coloring. It has a white or "bald-faced" head, which is the source of its colloquial name. This wasp also has three white stripes at the end of its body and is notably larger than other species of Dolichovespula, as adults average about 19 mm (0.75 in) in length. [4] Queen and worker wasps have similar morphologies. Queens are always larger than workers in their colonies, though size distributions can vary in different nests, and workers in one colony might be as large as a queen in a different one. [1]

D. maculata creates egg-shaped, paper nests up to 360 mm (14 in) in diameter and 580 mm (23 in) in length. Nests are layered hexagonal combs covered by a mottled gray paper envelope. Bald-faced hornets create this paper envelope by collecting and chewing naturally occurring fibers. The wood fiber mixes with their saliva to become a pulpy substance that they can then form into place. [4]

Distribution

Dolichovespula maculata distribution Dolichovespula maculata distribution.jpg
Dolichovespula maculata distribution

The bald-faced hornet lives in North America, including Canada, the Rocky Mountains, the western coast of the United States, and most of the eastern United States. It is most common in the Southeastern United States. [3]

D. maculata is found in forested areas and in vegetation in urban areas. Nests are generally located in trees and bushes, but occasionally are found under rock overhangs or the sides of buildings. Vertical distribution of nests has been recorded from heights of 0.3 to 20 m (1 ft 0 in to 65 ft 7 in) above ground level. [5]

Behavior

Bald-faced hornets are omnivorous, and considered to be beneficial due to their predation of flies, caterpillars, and spiders. Their aggressive defensive nature, though, makes them a threat to humans who wander too close to a nest or when a nest is constructed too close to human habitation. They vigorously defend the nest, with workers stinging repeatedly, as is common among social bees and wasps. The bald-faced hornet has a unique defense in that it can squirt or spray venom from the stinger into the eyes of vertebrate nest intruders. The venom causes immediate watering of the eyes and temporary blindness. [4]

Colony cycle

The life cycle of a colony can be divided into the founding stage, the ergonomic stage, and the reproductive stage. [6] Colonies show annual cycling. New nests are generally founded during spring and early summer by a single queen, though temporal specifics vary depending on location. In Washington, nest initiation occurs during mid-May, and workers emerge during mid-June. Large-cell building starts during mid-July, and the first queens emerge during mid-August. The colony terminates during mid-September, for a life cycle around 4 months (122 days). [5] Lower latitudes correlate with longer life cycles. In Indiana, colonies have been observed to begin in early May and terminate in late September, a life cycle of 5 months (153 days), and in Central California, nests are initiated as early as the end of March. These nests survive between 155 and 170 days. [5] Active colonies have been observed in central Pennsylvania as late as mid-October. On October 28, 2022, an active colony was encountered on the Middle Fork Willamette River in western Oregon.

Founding stage

The colony is founded by a single overwintered, inseminated queen. She rears the first generation of workers on her own until they are functional. Colonies pass through the foundation over a typical period of 23–24 days. After the queen lays her eggs, 6 days are needed for them to hatch. They grow as larvae for 8 days, then an additional 9–10 days to mature into adult workers. [5]

Ergonomic and reproductive stage

During the ergonomic stage, the colony's activities are concerned with cell building and worker production. The queen devotes herself entirely to laying eggs, while the workers take over all other necessary housekeeping tasks. Sometime before the midpoint of colony life, the colony begins to invest energy in producing reproductives, marking the transition into the reproductive stage, during which both workers and reproductives are raised. Production of both castes limits the size of the workforce, but an early switching time is highly adaptive for social-wasp colonies in limiting the risk of total loss. [6] In a sampling of 50 colonies taken in Maryland in 1977, workers were produced from mid-April to early October, and reproductives were produced from mid-July through the end of November. Onset of male output usually precedes that of queen output in social wasps, but D. maculata is an exception. [6]

Energy investment by workers required to produce reproductives is considerable. Newly emerged males and queens are no more functional than pupae; both depend heavily on solid prey brought in by workers, thus competing with larvae for food resources. As a result, worker quantity must remain high to maintain reproductive output. Workers must maintain food stores and defend the nest, and colonies whose work force diminishes too early in the colony's life cycle can suffer a greater overall loss in reproductives. [6]

Social organization

Caste structure and distribution

D. maculata male on goldenrod 2919baldf.w.jpg
D. maculata male on goldenrod

A colony is divided into haploid males, female workers, and the queen. All females are born with reproductive capacities. Dolichovespula is unique from its sister group Vespula in that some of the workers create haploid offspring that develop into males. [1] Caste systems are determined by larval feeding regimens. G. T. Felippotti et al examined caste distribution among females in five small-cell colonies and six large-cell colonies. Small-cell colonies had one queen and 17–21 female workers. Large-cell colonies had two to six queens and 10–52 workers. Morphological comparisons revealed that queens are always larger than workers in the same colonies. [7]

Cuticular hydrocarbons

Cuticular hydrocarbons serve as a barrier to moisture diffusion, so prevent dehydration in wasps. Cuticular hydrocarbon profiles vary over species and nests, and they are a proposed mechanism for nest-mate recognition. Worker and queen cuticular lipids have similar components, but their distributions differ dramatically, implying that cuticular hydrocarbons also play a role in caste differentiation. [8]

Cuticular hydrocarbon profile and dimorphism among castes

The following lipid profiles were determined from specimens collected in the summer and early fall of northeastern Georgia in 1989. The dominant hydrocarbons recovered from the cuticles of workers were n-alkanes and methyl- and dimethyl-branched alkanes with a C27 or a C29 backbone. The major lipids and their distributions in workers were: n-heptacosane (28%), 11-13-methylheptacosane (15%) 3,13-dimethylheptacosane (11%), and 13 and 15-methylnonacosane (10%). Nonacosene composed 34% of cuticular lipids in the queen. The average chain length of identified cuticular lipids is 28.0 in queens and 27.5 in workers. [8]

Kin selection

Genetic relatedness within various classes

In D. maculata, queens mate with only one male, which results in a relatedness of 0.75 between workers, whereas the relatedness between the queen and worker is only 0.5. [4]

Kin recognition and discrimination

Gynes in D. maculata have the ability to discriminate between fragments of natal comb and foreign comb. Recognition does not depend upon presence of viable brood in the comb fragment. The physical nature of cues mediating natal comb recognition is unknown, though some researchers propose that distinct cuticular hydrocarbon profiles allow wasps to recognize nestmates. [8] [9]

Worker-queen conflict

D. maculata is characterized by low paternity, worker reproduction, and queen-worker conflict. Divergent genetic interests between workers and their queen cause intranest struggle for control, which disrupts social organization. Because of haplodiploidy, workers are unable to mate, but their unfertilized eggs become males. Workers and their queens are most related to their own sons. Natural selection then favors those workers that produce their own sons rather than rearing the queen's brood. In a sampling of seven D. maculata nests, 20.9% of males were produced by workers. The percentage of males that were workers' sons did not correlate with the time during which nests were collected or colony size. Because worker relatedness is so high in D. maculata, workers are more related to other workers' sons than to the queen's own sons, so worker policing of egg production does not occur. An explanation for the queen's near monopoly on male production is that worker production is costly, so reduces total colony reproduction. The cost toward worker production acts as a selective pressure on the workers, so they are more likely to exercise reproductive restraint.

Workers in reproductive nests may kill their queen so they can reproduce. When researchers examined a collection of 19 D. maculata nests during the reproductive phase of their life cycle, they found that 14 nests did not have a queen. Matricide might occur after a sufficient workers have been raised and queen-destined eggs have been laid. However, matricide has not been directly observed and other causes of death are possible.

Diet

Diet in D. maculata varies depending on an individual's life cycle stage and geographic location. Adult yellowjackets, like true hornets, are carnivorous, and prey upon several insect types. They have been observed consuming meat, spiders, fruit, and insects. Adults also drink flower nectar, which they feed to their larvae. [10]

Parasites

Female bee moths ( Aphomia sociella ) have been known to lay their eggs in bald-faced hornet nests. The hatched larvae then proceed to feed on the eggs, larvae, and pupae left unprotected by the wasps, sometimes destroying large parts of the nest as they tunnel throughout looking for food. [11]

Life cycle

Each spring, queens that matured and were fertilized at the end of the previous season begin new colonies. A queen selects a location for her nest, begins building it, lays a first batch of eggs, and feeds this first group of larvae. These become workers and assume the chore of expanding the nest. They chew up wood, which mixes with a starch in their saliva. They then spread it around with their mandibles and legs, and it dries into a papery structure. The workers guard the nest and feed on nectar, tree sap and fruit pulp (particularly that of apples). They also prey on insects and other arthropods, chewing them up and feeding them to the larvae. They have been known to scavenge raw meat. In late summer and early fall, the queen begins to lay eggs that become drones and new queens. After pupation, these fertile males and females fly off to mate. Fertilized queens then overwinter and start new colonies during the next year. Males and workers die in the end of the cycle. The old queen, if not killed by workers, dies with them around mid-autumn.

Related Research Articles

<span class="mw-page-title-main">Hornet</span> Genus of eusocial wasp

Hornets are the largest of the eusocial wasps, and are similar in appearance to their close relatives yellowjackets. Some species can reach up to 5.5 cm (2.2 in) in length. They are distinguished from other vespine wasps by the relatively large top margin of the head. Worldwide, 22 species of Vespa are recognized. Most species only occur in the tropics of Asia, though the European hornet, is widely distributed throughout Europe, Russia, North America, and north-eastern Asia. Wasps native to North America in the genus Dolichovespula are commonly referred to as hornets, but are actually yellowjackets.

<span class="mw-page-title-main">Yellowjacket</span> Common name for two genera of wasps

Yellowjacket or yellow jacket is the common name in North America for predatory social wasps of the genera Vespula and Dolichovespula. Members of these genera are known simply as "wasps" in other English-speaking countries. Most of these are black and yellow like the eastern yellowjacket and the aerial yellowjacket ; some are black and white like the bald-faced hornet. Some have an abdomen with a red background color instead of black. They can be identified by their distinctive markings, their occurrence only in colonies, and a characteristic, rapid, side-to-side flight pattern prior to landing. All females are capable of stinging. Yellowjackets are important predators of pest insects.

<i>Polistes</i> Genus of insects

Wasps of the cosmopolitan genus Polistes are the most familiar of the polistine wasps, and are the most common type of paper wasp in North America. Walter Ebeling coined the vernacular name "umbrella wasps" for this genus in 1975 to distinguish it from other types of paper wasp, in reference to the form of their nests. It is also the single largest genus within the family Vespidae, with over 300 recognized species and subspecies. Their innate preferences for nest-building sites leads them to commonly build nests on human habitation, where they can be very unwelcome; although generally not aggressive, they can be provoked into defending their nests. All species are predatory, and they may consume large numbers of caterpillars, in which respect they are generally considered beneficial.

<span class="mw-page-title-main">Median wasp</span> Species of wasp

The median wasp is a species of social wasp of the family Vespidae found throughout Europe and Asia. It builds aerial paper nests often in shrubs or trees, and occasionally under the eaves of buildings. It is most common to see this wasp between May and October during its 3.3 month colony cycle. Behaviours of this wasp include nest defense, curling which is believed to function in brood incubation, and gastral vibration which is involved in larval feeding. The median wasp has a halplodiploid sex determination system that results in a high level of relatedness within the colony. This species is not usually aggressive but will sting if they feel their nest is threatened. Most foraging in the nest is done by the workers once the first ones reach adulthood. These workers forage for insects, nectar, and wood for nest construction in temperatures as low as 7 °C (45 °F). The median wasp is known to be occasionally affected by the fungus Cordyceps sphecocephala and the Cricket paralysis virus.

<i>Vespula germanica</i> Species of wasp

Vespula germanica, the European wasp, German wasp, or German yellowjacket, is a species of wasp found in much of the Northern Hemisphere, native to Europe, Northern Africa, and temperate Asia. It has spread and become well-established in many other places, including North America, South America, Australia, South Africa, and New Zealand. German wasps are part of the family Vespidae and are sometimes mistakenly referred to as paper wasps because they build grey paper nests, although strictly speaking, paper wasps are part of the subfamily Polistinae. In North America, they are also known as yellowjackets.

<i>Dolichovespula</i> Genus of wasps

Dolichovespula is a small genus of social wasps distributed widely throughout the Northern Hemisphere. The yellow and black members of the genus are known by the common name yellowjackets in North America, such as Dolichovespula norwegica, along with members of their sister genus Vespula. In a study on the nesting biology of Dolichovespula, a colony of D. maculata with 771 workers was reported as having the largest recorded population count.

<span class="mw-page-title-main">European paper wasp</span> Species of wasp

The European paper wasp is one of the most common and well-known species of social wasps in the genus Polistes. Its diet is more diverse than those of most Polistes species—many genera of insects versus mainly caterpillars in other Polistes—giving it superior survivability compared to other wasp species during a shortage of resources.

<span class="mw-page-title-main">Eastern yellowjacket</span> Species of insect

The eastern yellow jacket or eastern yellowjacket is a wasp found in eastern North America. Although most of their nests are subterranean, they are often considered a pest due to their nesting in recreational areas and buildings. This yellow jacket is a social insect, living in colonies of hundreds to thousands of individuals. Along with their subfamily, Vespinae, this species demonstrates supportive parental care for offspring, separation of reproductive and sterile castes, and overlapping generations. They aggressively defend their hives from threats and are known to inflict painful stings.

<i>Polistes annularis</i> Species of wasp

Polistes annularis is a species of paper wasp found throughout the eastern half of the United States. This species of red paper wasp is known for its large size and its red-and-black coloration and is variably referred to as a ringed paper wasp or jack Spaniard wasp. It builds its nest under overhangs near bodies of water that minimize the amount of sunlight penetration. It clusters its nests together in large aggregations, and consumes nectar and other insects. Its principal predator is the ant, although birds are also known to prey on it. Unlike other wasps, P. annularis is relatively robust in winter conditions, and has also been observed to store honey in advance of hibernation. This species has also been used as a model species to demonstrate the ability to use microsatellite markers in maternity assignment of social insects.

<i>Dolichovespula saxonica</i> Species of wasp

Dolichovespula saxonica, also known as the Saxon wasp, is a common social wasp found in the Palearctic region, specifically in large parts of Europe and in northern and central Asia. Although originally from continental Europe, D. saxonica has since colonised Britain, mainly in the south and east, but has been recorded as far north as East Lothian, Scotland. Most of their nests are above ground in trees and bushes, but they can also be found in buildings. Due to the proliferation of nests in urban areas and near residential homes, D. saxonica can be a pest for people. As a result, many human interventions are in place to remove Saxon nests. D. saxonica has been found to use chemical signaling in a lot of behaviours, such as alarm calls, fertility cues, and chemical trails.

<i>Dolichovespula norwegica</i> Species of wasp

The Norwegian wasp is a species of eusocial wasp. It is common in Scandinavia and can also be found in Scotland and other areas in Britain and Ireland. Often known for being a tree wasp, it nests in low branches and bushes and feeds on insects. It also obtains nectar from blueberry and snowberry flowers. Although D. norwegica is rarely considered a pest in the past, a few cases of pest problems relating to them have been reported. The species is not endangered.

<i>Dolichovespula sylvestris</i> Species of wasp

The tree wasp is a species of eusocial wasp in the family Vespidae, found in the temperate regions of Eurasia, particularly in western Europe. Despite being called the tree wasp, it builds both aerial and underground paper nests, and can be found in rural and urban habitats. D. sylvestris is a medium-sized wasp that has yellow and black stripes and a black dot in the center of its clypeus. It is most common to see this wasp between May and September during its 3.5 month colony cycle.

<i>Dolichovespula adulterina</i> Species of wasp

Dolichovespula adulterina is a species of parasitic social wasp found in the Palearctic region. D. adulterina feeds on a variety of foods, including insects, spiders, arthropods, meat, molluscs, fruit, nectar, and larval secretions. D. adulterina was formerly considered to be synonymous with D. arctica from the Holarctic region, but more recent research indicates that D. arctica is a separate species.

<i>Vespula squamosa</i> Species of wasp

Vespula squamosa, or the southern yellowjacket, is a social wasp. This species can be identified by its distinctive black and yellow patterning and orange queen. This species is typically found in eastern North America, and its territory extends as far south as Central America. Within these territories, they create enormous, multiple-comb nests. The colonies may be either annual or perennial depending on the climate, and in many perennial nests, polygyny takes place. In addition, this species uses pheromones both as a sexual attractant and an alarm signal. This species feeds on insects and animal carcasses; it does not produce honey. V. squamosa, a social insect, has developed a parasitic relationship with the species V. vidua and V. maculifrons. Due to their painful, venomous stings, the species is considered a pest.

<span class="mw-page-title-main">Worker policing</span> Insects destroying eggs not laid by queen

Worker policing is a behavior seen in colonies of social hymenopterans whereby worker females eat or remove eggs that have been laid by other workers rather than those laid by a queen. Worker policing ensures that the offspring of the queen will predominate in the group. In certain species of bees, ants and wasps, workers or the queen may also act aggressively towards fertile workers. Worker policing has been suggested as a form of coercion to promote the evolution of altruistic behavior in eusocial insect societies.

Apoica flavissima is a paper wasp found primarily in South America. The species is distinguishable by its light coloring, unique single comb nests, and nocturnal nature. A notable feature of this species is the size dimorphism between queens and workers. Unlike most Vespidae wasps, Apocia flavissima queens are smaller than their worker counterparts which results in unique intraspecies relationships.

<i>Dolichovespula arenaria</i> Species of wasp

Dolichovespula arenaria, also known as the common aerial yellowjacket, sandhills hornet, and common yellow hornet, is a species of wasp within the genus Dolichovespula widely distributed in the North American continent.

<i>Polistes semenowi</i> Species of wasp

Polistes semenowi is a species of paper wasp in the genus Polistes that is found in southeastern and southern central Europe, as well as central Asia, and was until 2017 erroneously known by the name Polistes sulcifer, while a different species was incorrectly believed to represent P. semenowi. It is one of only four known Polistes obligate social parasites, sometimes referred to as "cuckoo paper wasps", and its host is the congeneric species Polistes dominula. As an obligate social parasite, this species has lost the ability to build nests, and relies on the host workers to raise its brood. P. semenowi females use brute force, followed by chemical mimicry in order to successfully usurp a host nest and take over as the queen.

<i>Ropalidia revolutionalis</i> Species of wasp

Ropalidia revolutionalis, the stick-nest brown paper wasp, is a diurnal social wasp of the family Vespidae. They are known for the distinctive combs they make for their nests, and they are mostly found in Queensland, Australia in the areas of Brisbane and Townsville. They are an independent founding wasp species, and they build new nests each spring. They can be helpful because they control insect pests in gardens.

<i>Vespula atropilosa</i> Species of wasp

Vespula atropilosa, also known as the prairie yellowjacket, is a black and yellow social wasp that forms annual colonies. Vespula atropilosa was formerly a member of the Vespa family. Males have either xanthic (yellow) or melanic (black) abdominal coloring. The prairie yellowjacket can be found in the Western and Midwestern United States as well as parts of southern Canada. It builds its nests underground and prefers large open areas such as pastures and golf courses. Vespula atropilosa colonies are founded by a queen in the spring and grow most during mid-summer. It preys on other insects such as flies and grasshoppers but is not a scavenger like other wasps. Queens mate with multiple males, and males will fight each other for matings. V. atropilosa is a social wasp that practices altruism. Prairie yellowjackets are not considered pests but are commonly found in backyards in the Pacific Northwest. This species derives its common name from the prairies and grasslands where it can be found.

References

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