Bombus terricola
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''Bombus terricola'', the yellow-banded bumblebee, is a species of bee in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
''
Bombus 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 group in the tribe Bombini, though a few extinct related gener ...
''. It is native to southern
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 ...
and the east and midwest of the United States. It possesses complex behavioral traits, such as the ability to adapt to a queenless nest, choose which flower to visit, and regulate its temperature to fly during cold weather. It was at one time a common species, but has declined in numbers since the late 1990s, likely due to urban development and parasite infection. It is a good pollinator of wild flowers and crops such as alfalfa, potatoes, raspberries, and cranberries.


Taxonomy and phylogeny

''B. terricola'' belongs to 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 ...
, which consists of
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
s, bees,
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s, and sawflies. ''B. terricola'' belongs to 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 encompasses
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 group in the tribe Bombini, though a few extinct related gener ...
s,
honey bees 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 ...
, stingless bees, and more. Within this, it is part of the genus ''Bombus,'' which consists of bumblebees. Kirby first defined this species in 1837. Bombus occidentalis has been speculated to be a subspecies of ''B. terricola'', but most experts now agree that it is its own distinct species. ''B. terricola'' is also closely related to ''B. affinis'' both phylogenetically and in terms of pheromone signalling. Oftentimes, the ''B. terricola'' is so similar to ''B. affinis'' that members of ''B. affinis'' can invade and dominate entire ''B. terricola'' nests without the hosts knowing.


Description

The yellow-banded bumblebee is black and yellowish-tan, and has a characteristic fringe of short yellow-brown hairs on its fifth abdominal segment. The queen is about long. The front half of the thorax is yellowish-brown, as are segments 2, 3 and 4 and the sides of segment 6 of the abdomen. The other parts of the thorax and abdomen are black. The worker is similar in appearance to the queen but smaller at a length of . The male is intermediate in size, being long. In the male, abdominal segments 2, 3, and 7 are yellowish-brown as are usually the sides of abdominal segment 6.


Distribution and habitat

''Bombus terricola'' occupies the eastern and Midwestern parts of the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
as well as southern
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 ...
. They are known to occupy a wide range of habitats including urban areas, meadows, grasslands, wetlands, woodlands, and farmlands. They can also occupy alpine meadows to lowland tropical forests.


Colony cycle

In ''B. terricola,'' there are three phases of colony development. The first phase, known as colony initiation, begins when a solitary queen starts to produce her first workers by laying
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
eggs. This leads to further
eusociality 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 genera ...
within the colony and the queen's continued efforts to produce more
worker bees The working class (or labouring class) comprises those engaged in manual-labour occupations or industrial work, who are remunerated via waged or salaried contracts. Working-class occupations (see also " Designation of workers by collar colo ...
. The emergence of workers is essential for colony growth. The onset of the second phase, known as the switch point, is when the queen stops laying
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
eggs and starts making haploid eggs to produce male bees. During the third phase, the workers exhibit overt aggression towards each other and towards the queen. The beginning of the third phase is known as the competition point. Reciprocal
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 (, ...
also occurs during this third stage.


Behavior


Reproduction

In ''Bombus terricola'', there are female-biased investment ratios. The workers of the colony try to bias the sex ratios to be 3:1, in favor of the worker bees, which are female. The workers attempt to bias the sex ratios so that they can benefit for their own gene propagation; however, the queen tries to bring the sex ratio back to a favorable 1:1 of males and females for her own benefit as well. This process is referred to conflict between kin and is commonly seen in bees. ''B. terricola'' queens usually have a single mating opportunity in one mating flight with multiple males; the queen then stores all of the
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, whi ...
in a
spermatheca The spermatheca (pronounced plural: spermathecae ), also called receptaculum seminis (plural: receptacula seminis), is an organ of the female reproductive tract in insects, e.g. ants, bees, some molluscs, oligochaeta worms and certain other ...
, from which only one sperm will get to fertilize her egg.


Orphaning

Sometimes, some ''B. terricola'' colonies become orphaned. If there are feeble and weak queens, the queen has a decreased ability to fertilize eggs or cannot fertilize them at all. With declining queen vigor, worker-laid male eggs became more common. Furthermore, the highest proportion of worker-laid male eggs was observed in smaller colonies. Due to the absence of a strong queen leader, these male-dominated colonies attempt to get young, strong queens back because that would be in the best interest and benefit for the colony.


Foraging

''Bombus terricola'' congregate in nectar rich areas. They can discern which flowers have been previously visited by other bees or are depleted of resources such as nectar or pollen. They usually visit consecutive flowers in one direction and do not return to previous flowers. They tend to go to areas that are rich in resources and usually do not visit these areas again once the bees deplete the resources. Instead, it has been found that ''B. terricola'' expend energy to fly to new locations of food sources. Although the flight costs energy, they expend it for future profit in finding areas abundant in resources. The mechanism of their
foraging Foraging is searching for wild food resources. It affects an animal's Fitness (biology), fitness because it plays an important role in an animal's ability to survive and reproduce. Optimal foraging theory, Foraging theory is a branch of behaviora ...
beyond this information is currently unknown; it is also unknown whether they do not return to previously visited sites due to memory of the flowers or memory of landmarks (such as a nearby tree).


Flower pollination

''B. terricola'' foragers are highly selective about the
flowers A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
they
pollinate Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
. They can visit about 12–21 flowers per minute. The amount, quality, and availability of
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
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 ...
are the primary qualities that ''B. terricola'' use to determine which flowers to pollinate. These bees can determine the quality of pollen grain from a distance, but the method as to how this is accomplished is still not yet known. Individual foragers vary in their speed, flight, directionality, and erratic movements. Some bees hover over flowers but never land on them. Flower visitation rates depend on the number of rewarding or non-rewarding flowers they encountered on their flights. Rewarding flowers are ones that contain much pollen or nectar. One study noted that ''B. terricola'' bypass flowers that they have previously visited and only foraged a small amount of pollen from them. Finally, the bees release their pollen by emitting one to four sharp buzzing sounds of one second each. While buzzing, ''B. terricola'' also rotate clockwise or counterclockwise.


Types of flowers

''B. terricola'' have been known to forage on
milkweed ''Asclepias'' is a genus of herbaceous, perennial, flowering plants known as milkweeds, named for their latex, a milky substance containing cardiac glycosides termed cardenolides, exuded where cells are damaged. Most species are toxic to humans ...
(''Aslepia syriaca''),
jewelweed ''Impatiens'' is a genus of more than 1,000 species of flowering plants, widely distributed throughout the Northern Hemisphere and the tropics. Together with the genus ''Hydrocera'' (one species), ''Impatiens'' make up the family Balsaminaceae. ...
(''Impatiens biflora''), and
fireweed ''Chamaenerion angustifolium'' is a perennial herbaceous flowering plant in the willowherb family Onagraceae. It is known in North America as fireweed, in some parts of Canada as great willowherb, in Britain and Ireland as rosebay willowherb. ...
(''Epilobium angustifolium''). ''B. terricola'' exhibit consistent foraging behavior and tend to visit the same flowers repeatedly, especially if these flowers were rewarding in the past in terms of the amount of
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
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 ...
they carried.


Temperature regulation

Bumblebees 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 group in the tribe Bombini, though a few extinct related gener ...
can fly at a wide range of temperatures, including normal air temperature or below freezing point. They must have body temperatures of at least 29-30 degrees
Celsius The degree Celsius is the unit of temperature on the Celsius scale (originally known as the centigrade scale outside Sweden), one of two temperature scales used in the International System of Units (SI), the other being the Kelvin scale. The ...
before they are able to free fly independently in the air. Like honeybees, they produce heat before their flight. To raise their body temperature, specifically their
thoracic 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 crea ...
temperature, these bees must expend a lot of energy. These bees can visit dispersed flowers within a short span of time at either high or low air temperatures. Sometimes at low temperatures, these bees do not have enough heat to maintain continuous flight. Consequently, landing on flowers warms them up so that their thoracic temperature remains high enough for flight. Since they use up a lot of energy for this
thermoregulation Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...
, they take a lot of nectar from flowers that are most readily available to use as energy storage. By maintaining a high thoracic temperature, ''B. terricola'' can look for new sources of food and have the advantage to fly away quickly if they are at risk for predation.


Interactions with other species


Parasites

There are a few
parasites Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted structurally to this way of lif ...
that have been found to affect the populations of ''Bombus terricola''. The
larva 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 ...
, ''
Physocephala ''Physocephala'' is a genus of flies from the family Conopidae. Species *'' P. antiqua'' ( Wiedemann, 1830) *'' P. biguttata'' von Röder, 1883 *'' P. burgessi'' (Williston, 1882) *'' P. chrysorrhoea'' ( Meigen, 1824) *'' P. curticornis'' Krà ...
'', infects about twelve percent of the worker population of ''B. terricola'' from July to August in
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 ...
,
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 ...
. There is another parasite that is known to affect this population: the
fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
''Nosema bombi''. Classified as a
fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
, ''Nosema bombi'' is a small,
unicellular A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...
parasite that is known to infect bumblebees. It may also be one of the factors that is leading to the decline in numbers of ''Bombus terricola''. There is a high prevalence of this parasite specifically in ''Bombus terricola'' populations.


Population decline

Similar to other species of the ''Bombus''
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
, ''B. terricola'' has been declining in numbers. A specific study in
Illinois Illinois ( ) is a U.S. state, state in the Midwestern United States, Midwestern United States. Its largest metropolitan areas include the Chicago metropolitan area, and the Metro East section, of Greater St. Louis. Other smaller metropolita ...
shows that there was a severe decline in the middle of the 20th century from 1940-1960. In the state of Illinois specifically, intensive
farming Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people to ...
and urban development have taken away the natural landscapes and habitats that these bees would normally occupy. This decline in ''B. terricola'' corresponded with the huge growth in
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people to ...
and urban development. Furthermore, the ''Nosema bombi'' parasite may also contribute to the decline in populations of ''B. terricola'' as they particularly attack these bees.


Conservation status

Like several other
North American North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Ca ...
species in its subgenus ''
Bombus 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 group in the tribe Bombini, though a few extinct related gener ...
'', the yellow-banded
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 group in the tribe Bombini, though a few extinct related gener ...
has suffered sharp declines in numbers since the mid-1990s.The Xerces Society for Insect Conservation has placed ''Bombus terricola'' on their "Red List" of endangered bees. The yellow-banded bumblebee has disappeared over large parts of its range, but is still present in Vermont, New Hampshire, and Maine, where the population appears to be stabilizing and in the
Great Smoky Mountains The Great Smoky Mountains (, ''Equa Dutsusdu Dodalv'') are a mountain range rising along the Tennessee–North Carolina border in the southeastern United States. They are a subrange of the Appalachian Mountains, and form part of the Blue Ridge ...
, where the population seems to be expanding. It is not clear whether this is because it has developed some resistance to the parasite, ''Nosema bombi'', or whether the parasite has not yet spread into these parts. The rusty-patched bumblebee (''Bombus affinis'') has suffered an even steeper decline. It will become clearer over time whether populations of these bees will remain viable or whether the species will become extinct.


References


External links

{{Authority control Bumblebees Hymenoptera of North America Insects of Canada Insects of the United States Insects described in 1837