Mischocyttarus Flavitarsis
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''Mischocyttarus flavitarsis'' is a social
paper wasp Paper wasps are Vespidae, vespid wasps that gather fibers from dead wood and plant stems, which they mix with saliva, and use to construct nests made of gray or brown papery material. Some types of paper wasps are also sometimes called umbrell ...
found in western North America. Their nests can be found both in forests close to rivers or in close proximity to human life under the eaves of roofs. Despite the fact that ''M. flavitarsis ''nests are frequently in close contact with humans, ''M. flavitarsis'' typically will not sting, but rather ram into the threatening individual. Their colony cycle typically begins before May and will last until October. The queen will then seek a hibernation site for the winter. Perched near female hibernation sites are males with whom the female will mate. The males have claimed their territory by rubbing sternal brushes along the border of the site, leaving a chemical that deters other individuals from approaching. ''M. flavitarsis'' feed on
arthropods Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
, nectar, and animal carcasses and are often prey to birds, ants, and praying mantis.


Taxonomy and phylogenetics

Henri Louis Frédéric de Saussure Henri Louis Frédéric de Saussure (; ; 27 November 1829 – 20 February 1905) was a Swiss mineralogist and entomologist specialising in studies of Hymenoptera and Orthopteroid insects. He also was a prolific taxonomist. Biography Sauss ...
, a Swiss mineralogist and entomologist, classified ''M. flavitarsis'' in 1854. The genus ''
Mischocyttarus ''Mischocyttarus'' is a very large, primarily Neotropical genus of social wasps with a few species found also in the Nearctic region. It is the only member of the tribe Mischocyttarini; the asymmetrical tarsal lobes of ''Mischocyttarus'' separat ...
''is the only genus in the tribe Mischocyttarini and is the second largest genus of social wasps. ''Mischocyttarus ''species'' ''are recognizable by the internal and external lobes of the tarsal segments. Additionally, mature larvae have an appendix-like process on their first abdominal
sternum The sternum or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major blood vessels from injury. Sh ...
.


Description and identification


Species

Paper wasps typically have long and slender bodies with trailing hind legs. The color of their body is generally brownish. Specifically, ''M. flavitarsis'' is divided into five races based on coloration.


Nest

The nests of ''M. flavitarsis ''often hang from a 2–3 mm pedicle and are symmetrically round under the stem. Although rare, nests have been observed to be asymmetrically oblong or built on a vertical wall. The cells are sloping combs, and there are typically fewer than 300 cells per nest, making a relatively small nest per colony.


Distribution and habitat

''M. flavitarsis'' is found in the western United States and
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
, ranging from southern
California California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
to British Columbia. Typically, the wasps build nests in riparian forests at elevations of 1500 to 2700 meters. At lower elevations, the nests are located in deciduous and hard evergreens, and at higher elevations in conifers. Nests are typically found within 50 meters of rivers. M. flavitarsis is also known to nest in the built environment, particularly in protected spaces such as wall voids, the underside of eves, etc.


Colony cycle


Pre-emergence

Following
hibernation Hibernation is a state of minimal activity and metabolic depression undergone by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It most ...
, nests will be initiated between March and May by one or two foundresses or queens. During this period, cell construction is high as queens are laying eggs at a rate of 0.5 eggs per day. Construction by the queen slows down once eggs begin to hatch, but will increase again when
pupae A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
appear. If there are two foundresses for the colony, then one becomes the dominant egg-layer and leaves the nest less frequently than the other. Typically, the dominant queen will eat the eggs laid by the other foundress.


Emergence phase

Adults will begin to emerge about 60 days after eggs have been laid. Egg,
larvae A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
, and pupae stages each take around 20 days to complete development. Early emerging daughters help to construct the
natal NATAL or Natal may refer to: Places * Natal, Rio Grande do Norte, a city in Brazil * Natal, South Africa (disambiguation), a region in South Africa ** Natalia Republic, a former country (1839–1843) ** Colony of Natal, a former British colony ...
nest by building it larger and strengthening the stem from which it hangs. They also help to fan the
brood Brood may refer to: Nature * Brood, a collective term for offspring * Brooding, the incubation of bird eggs by their parents * Bee brood, the young of a beehive * Individual broods of North American Periodical Cicadas: ** Brood X, the largest bro ...
to keep it cool and defend the nest against predators. After the adults are 10–15 days old, they begin to forage for fiber,
arthropods Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
, and nectar. A queen continues to forage through this stage but will leave the nest more rarely as adults emerge. If queens do leave the nest, it is typically only for construction materials for the nest.


Colony termination

Foraging for the nest continues through August, and the adults that emerge at this time are typically males and non-worker females. However, egg-laying begins to slow down as temperatures begin to drop. The nest will be abandoned by ''M. flavitarsis'' between September and November as the females begin to seek shelter for hibernation.


Behavior


Dominance hierarchy

Female ''M. flavitarsis'' are capable of laying eggs, so dominance among females is established through physical attacks and
oophagy Oophagy ( ) sometimes ovophagy, literally "egg eating", is the practice of embryos feeding on eggs produced by the ovary while still inside the mother's uterus. The word oophagy is formed from the classical Greek (, "egg") and classical Greek (, ...
. The dominant female becomes queen and will eat the eggs of subordinates on the same day they are laid to direct reproduction.


Mating behavior

During the summer season, males can be found in female foraging areas attempting to mate. However, as fall approaches, males will begin to set up
leks A lek is an aggregation of male animals gathered to engage in competitive displays and courtship rituals, known as lekking, to entice visiting females which are surveying prospective partners with which to mate. A lek can also indicate an avail ...
near female hibernation sites. Fall mating occurs during September and October. Males will choose perch sites near areas frequented by females, sometimes changing sites up to 3 times per day. However, after six to seven days of waiting, the male will then remain at his perch site for the rest of the 6–7 weeks of mating. Once a site is chosen, the male will drag his abdomen along the surface of the perch site for scent marking. When a female comes to mate, she lands on the perch site. The male will then either lunge or walk slowly toward the female. The male mounts the female for approximately 8 seconds and extends its curled antennae and draws the female antennae upward in a behavior called lassoing. Following mating, females will walk over the surface of the perch site, dragging their abdomens, before flying off.


Male defense of perch site

When another wasp attempts to land on an occupied perch site, the owner is alerted to the intruder. If the intruder does not fly away right away, then the owner will lunge and grapple the other wasp. Typically the two wasps will fight on the perch site and often fall off the site and land on the ground to finish fighting. Following the fight, the owner returns to its perch site, and again, rubs its abdomen to secrete its scent and ward off future intruders.


Nest usurpation

A common practice of ''M. flavitarsis'' is nest
usurpation A usurper is an illegitimate or controversial claimant to power, often but not always in a monarchy. In other words, one who takes the power of a country, city, or established region for oneself, without any formal or legal right to claim it as ...
, in which the queen is displaced by a subordinate daughter or foreign wasp, which will then become the queen of the colony. The replacement queen is defined as whoever becomes the primary egg-layer in the nest. Typically, nest usurpation takes place during the pre-emergence phase or early emergence phase because there are fewer adult wasps to deter the usurper. Attempts occur because the queen is away from the nest and disappears, or there is only one foundress to overthrow. When a usurper attacks, the two wasps fight by biting, using antennation, and grappling each other attempting to sting each other. If the queen is usurped, then sometimes the older daughters will follow the queen when she flees to start a new nest. Daughters that hatch following usurpation will work for the usurper-queen.


Foraging

''M. flavitarsis'' will often forage along a stream in search of plant fibers to construct their nest, arthropods and nectar for feeding, and water for cooling the nest. Time spent away from the nest during foraging varies depending on what the wasp is looking for. When looking for pulp, a wasp will spend about 5 minutes away from the nest. A wasp searches for arthropods and nectar for about 15 minutes. Typically in the early stages of the nest when there are fewer adult wasps, they are away from the nest for a shorter amount of time.


Prey detection

''M. flavitarsis'' wasps are generalist predators, meaning their cues come from the many different insects and plants they feed on. Therefore, they use both visual and olfactory cues to find their prey.


Visual cues

It has been found that ''M. flavitarsis'' wasps often use visual cues to locate their prey on leaves. Often, the larvae of caterpillars will be eaten by ''M. flavitarsis'' if they are on a damaged plant leaf. A leaf that is partially eaten by a caterpillar attracts the wasps due to its irregular shape. The wasp is more likely to approach a damaged leave with holes in the middle of the leaf rather than along the edge.


Olfactory cues

''M. flavitarsis ''wasps also locate prey by smell. In one study, holes were made in a leaf and placed near a plant that had holes made in its leaves twelve days prior. Once caterpillar larvae were placed on the leaf, the time for the wasp to locate the larvae was recorded. It was found that ''M. flavitarsis'' are more likely to approach the freshly damaged leaf, which release mechanical odors that serve as cues for the wasps. In studies, these wasps are more likely to approach a leaf based on olfactory cues, rather than visual cues. This shows that presented
olfactory The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
stimuli are processed before the wasp processes visual stimuli.


Glandular influences

Male ''M. flavitarsis'' have larger sixth, seventh, and eighth sternal glands than other wasps, suggesting a role in secretion and possible sexual attractant. Secretion is suggested because the wasps will often rub their abdomens along the nest, spreading a highly repellant substance to deter ants. The use of secretion is also used by males when trying to attract a female because they will rub their abdomen along the perimeter of their perching territory. Near the openings of the glands are sternal brushes, which assist in rubbing the secreted substance along the surface. Sternal brushes occur only in the ''M. flavitarsis ''of the ''Mischocyttarus ''genus.


Hibernation

Due to the relatively cool and dry climate in which ''M. flavitarsis ''lives, it is the only species in the ''Mischocyttarus ''genus to hibernate and will seek shelter for the duration of the months of October through April. Typically, only female wasps will hibernate, with male wasps rarely being observed during the hibernation season. Female wasps will hibernate in cracks in eaves and roofs, cavities in electric light boxes, loose bark, logs, rocks, boxes, and attics. Rarely, females will emerge on warm winter days to fly in the sun before returning to hibernate for the duration of winter.


Differences in male and female behavior

Typically, female ''M. flavitarsis ''will remain with its nest for the duration of their life. On the other hand, males remain on the natal nest for only a few days. Following departure from the colony, they spend the rest of their lives attempting to attract a mate.


Interaction with other species


Diet

''M. flavitarsis ''forage for arthropods like flies and caterpillars to feed to larvae. They will also feed upon fresh meat and hides of animals or fish. Adult wasps will drink nectar for themselves, to feed to larvae, or to store in cells.


Predators

Birds such as flickers, acorn woodpeckers, Mexican jays, and flycatchers are typical predators of ''M. flavitarsis''. In addition to birds, ants will often try to invade the wasp nests. The wasps fight back by attempting to drive and throw the ants off the nest. They will also rub their abdomen on nest surfaces to deter ants from approaching. One final predator of ''M. flavitarsis'' is the praying mantis, which will eat perching males during mating season.


Defense

The defensive behavior of ''M. flavitarsis'' is dependent on which phase the colony cycle is in. If the nest is in the emergence phase or later, ''M. flavitarsis'' are more likely to sting a predator or intruder as a form of altruistic suicide because of colony size. However, typically the species is less prone to attack and will opt to ram the attacker as if to frighten it. When disturbed, ''M. flavitarsis ''will stand high on their mid and hind legs with their forelegs and abdomen raised and antennae forward. They then show displeasure by buzzing their wings to produce a sound audible from several meters away.


Parasites

''M. flavitarsis ''nests are often invaded by moths (''
Chalcoela iphitalis ''Chalcoela iphitalis'', the sooty-winged chalcoela, is a moth species of the family Crambidae that occurs throughout North America. They are seen as far south as California, Arizona and South Carolina and as far north as Ontario. Adults can be ...
'') at night because the wasps cannot see them. The moths migrate among the cells consuming wasp larvae and pupae. They will then lay their larvae, which spin cocoons in empty cells. ''M. flavitarsis'' do not attempt to rid the nest of the parasite. Instead, they will continue as a nest or abandon and renest elsewhere.


References

{{Taxonbar, from=Q14424291 Vespidae Hymenoptera of North America Insects of Canada Insects of the United States Insects described in 1854