Bombus Vancouverensis
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''Bombus vancouverensis'' is a common species of
eusocial Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
bumblebee of the subgenus ''Pyrobombus''. ''B. vancouverensis'' inhabits mountainous regions of western North America, where it has long been considered as a synonym of ''
Bombus bifarius ''Bombus bifarius'', the two-form bumblebee, is a species of eusocial bumblebee of the subgenus ''Pyrobombus''. ''B. bifarius'' inhabits mountainous regions of western North America, primarily the states of Colorado and Utah. Its common name refe ...
'', and essentially all of the literature on ''bifarius'' refers instead to ''vancouverensis''. ''B. vancouverensis'' has been identified as one of the two species of bumblebee observed to use
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s in kin recognition. The other is the
frigid bumblebee ''Bombus frigidus'', the frigid bumblebee, is a rare species of bumblebee largely found in Canada and parts of the United States. These bees have adapted to their cold environment by being able to keep their internal temperature within a certai ...
, ''Bombus frigidus''.


Taxonomy

''Bombus vancouverensis'' has two recognized subspecies: *''Bombus vancouverensis vancouverensis'' Cresson, 1878 - limited to
British Columbia British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, ...
*''Bombus vancouverensis nearcticus'' Handlirsch, 1888 - widespread in the United States and Canada ''Bombus vancouverensis'' was first described by
Ezra Townsend Cresson Ezra Townsend Cresson, also Ezra Townsend senior (18 June 1838, in Byberry – 19 April 1926, in Swarthmore) was an American entomologist who specialised in the Hymenoptera order of insects. He wrote ''Synopsis of the families and genera of the H ...
in the 1878 ''Proceedings of the Academy of Natural Sciences of Philadelphia''. It is a member of the order
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
and the family
Apidae Apidae is the largest family within the superfamily Apoidea, containing at least 5700 species of bees. The family includes some of the most commonly seen bees, including bumblebees and honey bees, but also includes stingless bees (also used for ...
, which also includes orchid bees, honey bees, and bumblebees. A subspecies, originally named ''Bombus nearcticus'', was named in 1888. ''B. vancouverensis'' expresses varying color forms, with the two most observed being a red-tailed "bifarius" form and a black-tailed ''nearcticus'' form, historically both considered to belong to the related species ''
Bombus bifarius ''Bombus bifarius'', the two-form bumblebee, is a species of eusocial bumblebee of the subgenus ''Pyrobombus''. ''B. bifarius'' inhabits mountainous regions of western North America, primarily the states of Colorado and Utah. Its common name refe ...
''. Individuals showing the red-tailed and black-tailed forms can both be found from Utah and Wyoming north to western Canada, Alaska, California, Oregon, Idaho, Montana, and Washington, though the black-tailed form is generally far more common. Because of differences in genetic structuring between these populations in various geographic locations, there had been debate as early as 2013 suggesting these two major color pattern polymorphisms might represent more than one biological species.Lozier, J.D., Strange, J.P. & Koch, J.B. (2013) Landscape heterogeneity predicts gene flow in a widespread polymorphic bumble bee, ''Bombus bifarius'' (Hymenoptera: Apidae). Conservation Genetics, 14, 1099–1110. As of 2020, the genetic differentiation between ''bifarius'' and ''vancouverensis'' has been confirmed and characterized, with true ''bifarius'' being exclusively red-tailed, and much more geographically-restricted, while ''vancouverensis'' is polymorphic and very widely distributed.


Description and identification

''Bombus vancouverensis'' has a relatively small body size ranging from for workers and for queens, with short, even hair covering their bodies. ''B. vancouverensis'' individuals express multiple color forms; however, many similarities exist between these color variants. Hair on the faces of ''B. vancouverensis'' individuals is usually yellow or white in color and sometimes exhibits black coloration on the top of the head. In at least the lowermost third of the
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...
, there is also black coloration. The hindlegs and
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
baskets can be a brownish-orange or black, depending on whether metasomal
tergite A ''tergum'' (Latin for "the back"; plural ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'mar ...
(abdominal segment) 3 is black or not. In the red-tailed color variant, metasomal tergites 2 and 3 are red, while in the black-tailed color variant, metasomal tergites 2 and 3 are black.


Sexual dimorphism

Males are similar in size to female workers, ranging from . Their eyes are also similar in size and shape to those of their female counterparts. Colorations on their bodies are similar to those of workers and the queen; however, T3 and T6 are most frequently black in males and can vary between black, red, and yellow in workers and queens.


Nests

''Bombus vancouverensis'' nests are made underground or on the surface of the ground. These nests are often small and are made up of one singular open chamber. ''B. vancouverensis'' can also make their nests in abandoned rodent nests.


Diet

''Bombus vancouverensis'' forages for pollen and nectar from the following plants: ''
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Ericameria ''Ericameria'' is a genus of North American shrubs in the family Asteraceae. ''Ericameria'' is known by the common names goldenbush, rabbitbrush, turpentine bush, and rabbitbush. Most are shrubs but one species ''( E. parishii)'' can reach tree ...
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Helenium ''Helenium'' is a genus of annuals and herbaceous perennial plants in the family Asteraceae, native to the Americas. They bear yellow or orange daisy-like composite flowers. A number of these species (particularly ''Helenium autumnale'') have ...
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Melilotus ''Melilotus'', known as melilot, sweet clover, and kumoniga (from the Cumans),Bulgarian Folk Customs, Mercia MacDermott, pg 27 is a genus in the family Fabaceae (the same family that also includes the ''Trifolium'' clovers). Members are known ...
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Monardella ''Monardella'' is a genus of approximately 40 species of annual and perennial plants native to western North America from British Columbia to northwestern Mexico. They are grown for their highly aromatic foliage, which in some species is used for ...
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Penstemon ''Penstemon'' , the beardtongues, is a large genus of roughly 250 species of flowering plants native mostly to the Nearctic, but with a few species also found in the North American portion of the Neotropics. It is the largest genus of flowering ...
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Ribes ''Ribes'' is a genus of about 200 known species of flowering plants, most of them native to the temperate regions of the Northern Hemisphere. The various species are known as currants or gooseberries, and some are cultivated for their edible f ...
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Senecio ''Senecio'' is a genus of flowering plants in the daisy family (Asteraceae) that includes ragworts and groundsels. Variously circumscribed taxonomically, the genus ''Senecio'' is one of the largest genera of flowering plants. Description Morp ...
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Solidago ''Solidago'', commonly called goldenrods, is a genus of about 100 to 120''Solidago''.
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'', and ''
Symphoricarpos ''Symphoricarpos'', commonly known as the snowberry, waxberry, or ghostberry, is a small genus of about 15 species of deciduous shrubs in the honeysuckle family, Caprifoliaceae. With the exception of the Chinese coralberry, '' S. sinensis'', wh ...
''.


Distribution and habitat

''Bombus vancouverensis'' is found across mountainous regions of western North America. In the United States, it has been found in parts of
Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U.S., ...
,
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 ...
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Idaho Idaho ( ) is a state in the Pacific Northwest region of the Western United States. To the north, it shares a small portion of the Canada–United States border with the province of British Columbia. It borders the states of Montana and Wyom ...
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Montana Montana () is a state in the Mountain West division of the Western United States. It is bordered by Idaho to the west, North Dakota and South Dakota to the east, Wyoming to the south, and the Canadian provinces of Alberta, British Columbi ...
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Nevada Nevada ( ; ) is a U.S. state, state in the Western United States, Western region of the United States. It is bordered by Oregon to the northwest, Idaho to the northeast, California to the west, Arizona to the southeast, and Utah to the east. N ...
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Oregon Oregon () is a U.S. state, state in the Pacific Northwest region of the Western United States. The Columbia River delineates much of Oregon's northern boundary with Washington (state), Washington, while the Snake River delineates much of it ...
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South Dakota South Dakota (; Sioux language, Sioux: , ) is a U.S. state in the West North Central states, North Central region of the United States. It is also part of the Great Plains. South Dakota is named after the Lakota people, Lakota and Dakota peo ...
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Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to it ...
,
Washington Washington commonly refers to: * Washington (state), United States * Washington, D.C., the capital of the United States ** A metonym for the federal government of the United States ** Washington metropolitan area, the metropolitan area centered on ...
, and
Wyoming Wyoming () is a U.S. state, state in the Mountain states, Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the south ...
. In Canada, it has been found in
Alberta Alberta ( ) is one of the thirteen provinces and territories of Canada. It is part of Western Canada and is one of the three prairie provinces. Alberta is bordered by British Columbia to the west, Saskatchewan to the east, the Northwest Ter ...
,
British Columbia British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, ...
,
Manitoba Manitoba ( ) is a Provinces and territories of Canada, province of Canada at the Centre of Canada, longitudinal centre of the country. It is Canada's Population of Canada by province and territory, fifth-most populous province, with a population o ...
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Northwest Territories The Northwest Territories (abbreviated ''NT'' or ''NWT''; french: Territoires du Nord-Ouest, formerly ''North-Western Territory'' and ''North-West Territories'' and namely shortened as ''Northwest Territory'') is a federal territory of Canada. ...
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Saskatchewan Saskatchewan ( ; ) is a Provinces and territories of Canada, province in Western Canada, western Canada, bordered on the west by Alberta, on the north by the Northwest Territories, on the east by Manitoba, to the northeast by Nunavut, and on t ...
, and
Yukon Yukon (; ; formerly called Yukon Territory and also referred to as the Yukon) is the smallest and westernmost of Canada's three territories. It also is the second-least populated province or territory in Canada, with a population of 43,964 as ...
. ''B. vancouverensis'' makes its home in various habitats, including open grassy fields, parks, prairies, shrubs, and mountainous areas.


Colony cycle

Colonies regenerate annually, with queens emerging from hibernation in the early spring to found one colony per queen. After emerging, queens produce the first wave of workers, initiating the
eusocial Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
phase. In the eusocial phase, exponential growth of the colony occurs and workers help queens expand the size of the colony. Eventually, the eusocial phase gives way to the reproductive phase, in which the offspring produced are sexual offspring. These offspring then leave the nest and mate with non-nestmates, and young queens that have been inseminated enter hibernation until the following spring when they found their new colony.


Behavior


Mating

''Bombus vancouverensis'' queens only mate once during the colony cycle, preferably with a non-related male. Often, the duration of mating interactions between males and ''B. vancouverensis'' queens are long, lasting up to forty-five minutes.


Kin recognition and inbreeding avoidance

''Bombus vancouverensis'' and '' B. frigidus'' prefer to mate with non-nestmates, so they have evolved a number of strategies to determine which individuals are kin and which are not. For example, environmental cues like proximity to the nest may indicate that a reproductive female is related. However, not all encounters fall under the category of environmental cues; ''B. vancouverensis'' is one of the few bumblebee species that can also determine kin using individually-borne methods. For example, males exhibit a behavior called “patrolling”, in which they mark specific paths with
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s and “patrol” these paths hoping to encounter a reproductive female that was attracted to the scent. Reproductive females are able to sense these pheromones and are attracted to them.


Mate guarding

''Bombus vancouverensis'' males often compete for access to females, which has caused them to evolve strategies to ensure that they successfully complete the mating process with a chosen queen. That mating interactions are prolonged indicates that males “guard” their mate to ensure other males cannot copulate with her. This behavior is costly, however, since prolonged mating with one queen means that males overall engage in fewer mating interactions. Additionally, mate guarding immobilizes the mating pair, making them more susceptible to predatory attacks.


Thermoregulation in the nest

Keeping the nest environment at an optimal temperature is a crucial aspect of survival, especially for developing
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 ...
. Studies have shown that ''B. vancouverensis'' workers are capable of regulating nest temperature; they do this in a number of ways, namely by changing the rates at which they perform specific behaviors of wing fanning and brood cell incubation. During wing fanning, individuals flap their wings quickly as a means of cooling down their surroundings. Brood incubation involves a coiling of the worker's body around a brood cell and contracting her muscles so as to transfer heat to her surroundings. Broadly, as the temperature of the nest increases, the rate of brood cell incubation decreases and wing fanning behaviors increase within the nest. However, these behaviors are not universally exhibited by individuals at the same temperatures; some individuals begin incubating at higher temperatures than others, and some begin fanning their wings at lower temperatures than others.


Foraging

Studies have shown that ''B. vancouverensis'' forage in random directions when foraging for pollen and nectar. These bees have mechanisms by which they can recognize which flowers they have already visited. The mechanism appears to be one by which the bee sees open flowers and senses how much pollen is still available in them. Their ultimate strategy for pollen foraging, whether it be through area-restricted searching behavior or moving between flowers near them regardless of quality of bloom, is variable depending on the conditions under which ''B. vancouverensis'' is foraging. For example, in '' P. gracilis'' blooms, the quality of bloom can be assessed by ''B. vancouverensis'' at relatively low cost to the individual, so moving to nearby neighboring flowers instead of area-restricted searching yields more energy intake per energy input.


Buzz pollination

''Bombus vancouverensis'', like many other bumblebees, gathers pollen from flowers in a process called
buzz pollination Buzz pollination or sonication is a technique used by some bees, such as solitary bees to release pollen which is more or less firmly held by the anthers. The anthers of buzz-pollinated plant species are typically tubular, with an opening at onl ...
. During buzz pollination, the bee “sonicates” the anthers of the flowers, thereby shaking the pollen from the anthers to be collected in the individual's pollen sacs. Because of their small size, ''B. vancouverensis'' individuals are known to hang upside-down on the flower when engaging in buzz pollination.


Nectar foraging

''Bombus vancouverensis'' workers have short proboscides (tongue-like structures), which makes nectar foraging more difficult for them relative to bumblebees with longer proboscides. Therefore, in order to effectively retrieve nectar from plants, ''B. vancouverensis'' individuals often use their short proboscides to lick along the back of nectar ducts of the plant in question. This yields a smaller reward than the rewards obtained with larger proboscides, but not small enough to make nectar foraging ineffective.


Foraging task specialization

Most workers of ''B. vancouverensis'' forage for both nectar and pollen; however, there has been evidence of specialization within foragers, with some collecting only nectar and others collecting only pollen throughout their whole careers as foragers. Studies have shown that foragers that specialize in collecting either nectar or pollen are able to gather more food than those that did not specialize.


References

{{taxonbar, from=Q101211048 Bumblebees Hymenoptera of North America Insects described in 1878 Taxobox binomials not recognized by IUCN