Bombus muscorum
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''Bombus muscorum'', commonly known as the large carder bee or moss carder bee, is a species of
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 ...
in 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 ...
. The species is found throughout
Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago a ...
in fragmented populations, but is most commonly found in the
British Isles The British Isles are a group of islands in the North Atlantic Ocean off the north-western coast of continental Europe, consisting of the islands of Great Britain, Ireland, the Isle of Man, the Inner and Outer Hebrides, the Northern Isles, ...
. ''B. muscorum'' is a
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 ...
insect. The queen is monandrous, mating with only one male after leaving a mature nest to found its own. Males mate territorially and the species is susceptible to inbreeding and bottlenecks. The species builds its nests on or just under the ground in open grassland and forages very close to the nest. In recent years, populations have significantly declined due to loss of natural habitat. ''B. muscorum'' is currently listed as vulnerable in Europe by the European Red List of Bees.


Taxonomy

''B. muscorum'' was one of the many insect species originally described in 1758 by
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
in the 10th edition of ''Systema Naturae.'' It was given the
binomial name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
''Apis muscorum.'' The species name ''muscorum'' is the genitive plural of the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
''mūscus,'' meaning moss. ''B. muscorum'' is part 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 ...
, the third largest order of insects, which includes bees, wasps, ants, and sawflies. It is part of the subfamily
Apinae The Apinae are the subfamily that includes the majority of bees in the family Apidae. It includes the familiar " corbiculate" (pollen basket) bees—bumblebees, honey bees, orchid bees, stingless bees, Africanized bees, and the extinct genus '' ...
, which contains the majority of species within 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 ...
. It is part of the tribe
Bombini The Bombini are a tribe of large bristly apid bees which feed on pollen or nectar. Many species are social, forming nests of up to a few hundred individuals; other species, formerly classified as ''Psithyrus'' cuckoo bees, are brood parasites ...
, which contains a single living genus
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 ...
, consisting entirely of bumblebees. Within ''
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 ...
'', ''B. muscorum'' is part of the subgenus '' Thoracobombus'', which includes closely relates species such ''
Bombus pauloensis ''Bombus pauloensis'' is a neotropical bumblebee, formerly known as ''Bombus atratus'', that is found throughout regions of South America, including Colombia, Ecuador, Brazil, and Argentina. It lives in social colonies that include a founder que ...
,
Bombus dahlbomii ''Bombus dahlbomii'', also known as the moscardón, is a species of bumblebee endemic to southern South American temperate forests. ''B. dahlbomii'' is one of the largest bee species in the world, with matured queens growing up to long.King, A. ...
,
Bombus fervidus ''Bombus fervidus'', the golden northern bumble bee or yellow bumblebee, is a species of bumblebee native to North America. It has a yellow-colored abdomen and thorax. Its range includes the North American continent, excluding much of the souther ...
,
Bombus morio ''Bombus morio'' is one of the few bumblebee species found in South America. These bees reside mainly in the forests of Brazil, nesting on the surface of the ground. They are one of the biggest species of bumblebee and are important pollinators. ...
,
Bombus pensylvanicus ''Bombus pensylvanicus'', the American bumblebee, is a threatened species of bumblebee native to North America. It occurs in eastern Canada, throughout much of the Eastern United States, and much of Mexico.Hatfield, R., et al. 2015''Bombus pensyl ...
,'' and ''
Bombus transversalis ''Bombus transversalis'' is a bumblebee specifically native to the Amazon Basin. It is most notable for its surface level colonies which are built by the workers on the rainforest floor. Unlike its relatives, ''B. transversalis'' is able to thriv ...
.'' The species is further differentiated into at least seven different
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
. These include ''B. muscorum sladeni'' and ''B. muscorum smithianu,'' found in the
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the continental mainland. It comprises England, Scotland, Wales and North ...
and differentiated on the basis of coat color. Certain forms of this variable taxon are considered separate species by some authors, including ''B. bannitus'' and ''B. pereziellus''.


Description

''B. muscorum'' is characterized by its oblong face and long tongue. The bristles of the species are ginger in color and cover the head, thorax, and abdomen. The abdomen, however, can have a darker, brownish coloring. In some subspecies, black bristles are intermixed with the ginger bristles that characterize the species. Some subspecies have an entirely black
ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Th ...
side. The species is similar in appearance to the more common ''
Bombus pascuorum ''Bombus pascuorum'', the common carder bee, is a species of bumblebee present in most of Europe in a wide variety of habitats such as meadows, pastures, waste ground, ditches and wikt:embankment, embankments, roads, and field margins, as well ...
.'' The queen has a body length 17–19 mm and a wingspan of 32–35 mm. Workers have a body length of 10–16 mm and a wingspan of 26–29 mm. Males have a body length of 13–15 mm and a wingspan of 26–29 mm.


Distribution and habitat

''B. muscorum'' is widely distributed throughout Eurasia. Populations appear from Ireland in the West to
Mongolia Mongolia; Mongolian script: , , ; lit. "Mongol Nation" or "State of Mongolia" () is a landlocked country in East Asia, bordered by Russia to the north and China to the south. It covers an area of , with a population of just 3.3 million, ...
in the East. The species has been observed as far north as Scandinavia and Russia and has been spotted as far south as Crete. Populations, however, are fairly rare in all locations. Populations in the United Kingdom have declined sharply since the 1970s. Once widely distributed throughout the British Isles, distributions are now fragmented and predominantly coastal. Populations of other species, such as ''
Bombus pascuorum ''Bombus pascuorum'', the common carder bee, is a species of bumblebee present in most of Europe in a wide variety of habitats such as meadows, pastures, waste ground, ditches and wikt:embankment, embankments, roads, and field margins, as well ...
'', appear to be replacing ''B. muscorum'' in some parts of Northern Britain. The species is still relatively abundant in Northern Scotland. ''B. muscorum'' is classified as vulnerable in Europe by the IUCN. ''B. muscorum'' inhabits moors, grasslands, and salt marshes, where it builds its nest above ground. In the British Isles, ''B. muscorum'' nests in open landscapes near coastal areas in the lowlands. Populations are found in moorland and machair in the North. In the South, populations are found in coastal marshes, shingle, and calcareous grasslands. Nests have been found near open regions of fallowed land and man-made ditches. Colonies are small, numbering between 20 and 100 workers.


Diet

''B. muscorum'' is polylectic, the diet of the species depends on the surrounding area. The species has a strong preference for flowers of the families
Fabaceae The Fabaceae or Leguminosae,International Code of Nomenc ...
,
Scrophulariaceae The Scrophulariaceae are a family of flowering plants, commonly known as the figwort family. The plants are annual and perennial herbs, as well as shrubs. Flowers have bilateral (zygomorphic) or rarely radial (actinomorphic) symmetry. The Scr ...
,
Lamiaceae The Lamiaceae ( ) or Labiatae are a family of flowering plants commonly known as the mint, deadnettle or sage family. Many of the plants are aromatic in all parts and include widely used culinary herbs like basil, mint, rosemary, sage, savory ...
, and
Asteraceae The family Asteraceae, alternatively Compositae, consists of over 32,000 known species of flowering plants in over 1,900 genera within the order Asterales. Commonly referred to as the aster, daisy, composite, or sunflower family, Compositae w ...
. Common food sources include
clover Clover or trefoil are common names for plants of the genus ''Trifolium'' (from Latin ''tres'' 'three' + ''folium'' 'leaf'), consisting of about 300 species of flowering plants in the legume or pea family Fabaceae originating in Europe. The genus ...
,
bird's-foot trefoil ''Lotus corniculatus'' is a flowering plant in the pea family Fabaceae, native to grasslands in temperate Eurasia and North Africa. Common names include common bird's-foot trefoil, eggs and bacon, birdsfoot deervetch, and just bird's-foot trefo ...
,
vetches ''Vicia'' is a genus of over 240 species of flowering plants that are part of the legume family ( Fabaceae), and which are commonly known as vetches. Member species are native to Europe, North America, South America, Asia and Africa. Some othe ...
, and
thistle Thistle is the common name of a group of flowering plants characterised by leaves with sharp prickles on the margins, mostly in the family Asteraceae. Prickles can also occur all over the planton the stem and on the flat parts of the leaves. ...
s. Flowers with long corollas are especially dependent on the long-tongued species.


Colony cycle

A number of new queens are produced in the last brood of the summer colony. These queens mate and then hibernate for an average of 9 months. From March to May, these queens emerge from hibernation to search for nesting sites. After a suitable site is found, the queen lays a small batch of diploid eggs. Once these eggs hatch, the queen tends to the larvae until they are grown and pupate. These pupa emerge as workers. From June onward, the queen will produce a steady supply of workers. From July to September, new males and queens are present and mate.


Reproduction


Haplodiploidy

''B. muscorum'', like many bees, exhibit
haplodiploidy Haplodiploidy is a sex-determination system in which males develop from unfertilized eggs and are haploid, and females develop from fertilized eggs and are diploid. Haplodiploidy is sometimes called arrhenotoky. Haplodiploidy determines the sex ...
. The
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 ...
queens produce diploid workers and new queens from fertilized eggs.
Haploid 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 ...
males are produced from unfertilized eggs. Workers are also able to lay unfertilized eggs that develop into males. Haploid males produce identical haploid sperm while diploid females produce genetically variant haploid eggs through meiosis. Sex is determined under a single-locus complementary sex determination (sl-CSD) system, where multiple alleles at a single locus determine the sex of an individual. Sex locus
heterozygotes Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mo ...
develop as females, while
hemizygous Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mo ...
and
homozygous Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mo ...
eggs develop as haploid and diploid males.


Mating system

''B. muscorum'' is thought to be a
monandrous In botanical terms, monandrous simply means to have a single stamen. In orchids A distinction between monandrous and other flowers is particularly relevant in the classification of orchids. The monandrous orchids form a clade consisting of the s ...
species; the queen mates only once with a single male to start a new colony. This monandrous behavior decreases the amount of genetic variation present in a single colony relative to that of a polygynous or
polyandrous Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wives" ...
species. As a result, ''B. muscorum'' has an increased susceptibility to the effects of inbreeding. Queens may possess a distinctive odor that signals relatedness and prevents interbreeding. Studies have shown, however, that mating appears to be random.


Nest surveillance

Male mating strategies of bumblebees can be grouped into four broad categories: patrolling, racing, territorial, and cruising. Males of ''B. muscorum'' exhibit territorial strategies for mating. In this strategy, males choose a perch outside of a mature nest and pursue potential mates that it spots. Males compete with one another for preferred perches, each attempting to find a spot close to the nest entrance. When another bee flies too close to a perched male, the male sets off in pursuit for several seconds, seeking to either chase away a potential competitor or mount a potential mate. After the chase, one of the males will land back at the perch close to the nest.


Inbreeding

As a result of small population sizes and haplodiploidy, ''B. muscorum'' has an increased susceptibility to inbreeding. As sex is determined at a single locus, the overall fitness of a population is directly related to the number of different alleles at the sex locus, which in turn is related to the size and isolation of the population. When males mate with related queens, diploid males with reduced fertility and immune response are produced. 10 out of 14 ''B. muscorum'' populations in the
Hebrides The Hebrides (; gd, Innse Gall, ; non, Suðreyjar, "southern isles") are an archipelago off the west coast of the Scottish mainland. The islands fall into two main groups, based on their proximity to the mainland: the Inner and Outer Hebrid ...
showed significant signs of recent
bottlenecking Bottleneck literally refers to the narrowed portion (neck) of a bottle near its opening, which limit the rate of outflow, and may describe any object of a similar shape. The literal neck of a bottle was originally used to play what is now known as ...
. In recent years, diploid males have come to represent a considerable proportion of males within populations in the British Isles, suggesting an increase in inbreeding among populations.


Behavior


Foraging

''B. muscorum'' exhibit small foraging ranges, having been recorded going no further than 500 m from their nest to forage. Foraging, however, often takes place within 100 m of the nest. Individuals are often observed foraging at the same spots between 50 and 200 m away from the nest. This variation in individual flight range is attributed to an individual's body size. ''B. muscorum'' has been described as doorstep foragers, due to the tendency of workers to use food resources near the nest more frequently than the workers of other species. This restricted radius may be an important factor in the decline of the species.


Nesting

''B. muscorum'' build its nest on or just under the ground. It cards together collected moss and dry grass to cover the nest. It is this behavior that gives ''B. muscorum'' the name moss carder bee. It rarely, if ever, crosses sea barriers greater than 10 km to establish a nesting site''.'' Once a nest is established, the bee is notoriously aggressive, readily attacking intruders that are too close to the nest, which they bite and sting simultaneously.


Parasitism


''Crithidia bombi''

''B. muscorum'' may contract infections of ''Crithidia bombi'', a trypanosome parasite, from infected nest mates or from others while foraging on contaminated flowers. Infections have been linked to reduced individual and colony fitness, but the specific relationship between ''Crithidia bombi'' and ''B. muscorum'' is complex. The presence of ''Crithidia bombi'' has been found to be higher in populations with lower genetic diversity. As ''B. muscorum'' populations continue to lose heterozygosity, the impact of parasitism increases, pushing already threatened populations closer to extinction.


Human impact


Habitat destruction

Due to recent decades of agricultural intensification in Europe, the natural habitat of ''B. muscorum'' has been largely diminished. Many of the permanent flower-rich areas where the bee commonly forages have been destroyed by increased commercial farming. ''B. muscorum'' has poor dispersal ability compared to other species of bumblebee, making it more vulnerable to habitat loss. As a result, populations have decreased dramatically in recent decades.


Status

''B. muscorum'' is currently listed as vulnerable in Europe by the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
. Populations are diminishing in Britain and the species is currently in a species recovery program. Decreasing populations are evident in multiple European countries.
Estonia Estonia, formally the Republic of Estonia, is a country by the Baltic Sea in Northern Europe. It is bordered to the north by the Gulf of Finland across from Finland, to the west by the sea across from Sweden, to the south by Latvia, a ...
lists the species as vulnerable. Germany and the Netherlands consider the species to be endangered.


Sea walls

''B. muscorum'' is often found on sea wall flood defenses managed by the
Environment Agency The Environment Agency (EA) is a non-departmental public body, established in 1996 and sponsored by the United Kingdom government's Department for Environment, Food and Rural Affairs, with responsibilities relating to the protection and enha ...
. Nests can be found on the flat grassy area between the sea and the borrowdyke, a brackish pond behind the sea wall. As grasslands on sea walls are predominantly unimproved, they have become an important habitat for rare insect species such as ''B. muscorum''. Current research is examining how these sea walls can be maintained to preserve rare populations.


References

{{Taxonbar, from=Q1352185 Bumblebees Hymenoptera of Europe Bees described in 1758 Taxa named by Carl Linnaeus