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Megachile campanulae

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Bellflower resin bee
Male M. campanulae
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Megachilidae
Subfamily: Megachilinae
Tribe: Megachilini
Genus: Megachile
Subgenus: Chelostomoides
Species: M. campanulae
Binomial name
Megachile campanulae
(Robertson, 1903)
Subspecies
  • M. c. wilmingtoni
    (Mitchell, 1924)
  • M. c. campanulae
    (Robertson, 1903)
Distribution of M. campanulae
Synonyms

Oligotropus campanulae
Robertson, 1903
Oligotropus wilmingtoni
Mitchell, 1924
Megachile (Chelostomoides) campanulae var. wilmingtoni
Mitchell, 1937

Megachile campanulae, known as the bellflower resin bee, is a species of bee in the family Megachilidae. Described in 1903, these solitary bees are endemic to eastern North America. Studies in 2013 placed them among one of the first insect species documented in scientific literature to use synthetic materials for making nests. They considered a mason bees, which is a common descriptor of bees in several families, including Megachilidae. While the members of the genus Megachile are frequently also referred to as leafcutter bees, M campanulae is a member of the subgenus Chelostomoides. These resin bees have mandibles lacking cutting edges. They do not construct nests from cut leaves, but rather from plant resins and other materials. Females lay eggs in nests constructed with individual cell compartments for each egg. Once hatched, the eggs progress through larval stages and subsequently will overwinter as pupae. The bees are susceptible to parasitism from several other bee species, which act as brood parasites. They are medium sized bees and the female adults typically larger than the males. They are important pollinators of numerous native plant species throughout their range.

Taxonomy and naming

Megachile campanulae was originally described in 1903 under the name Oligotropus campanulae by Charles Robertson, an American entomologist from Carlinville, Illinois. Megachile translates from Greek mega (μεγας) 'large' + cheil- (χειλ) 'lip'. In Latin, campanulae translates as "small bell". M. campanulae has been documented to frequent flowers in the genus Campanula, several species of which are commonly referred to as bellflowers. Subspecies include M. campanulae campanulae and M. campanulae wilmingtoni. The genus Megachile is a cosmopolitan group of solitary bees, often called leafcutter bees and resin bees. It is one of the largest genera of bees, with 1520 species in 56 subgenera worldwide.

Life cycle and behavior

As a member of the subgenus Chelostomoides, M. campanulae are mason bees. This means that they use plant resins, mud, and pebbles for nest construction. Typically the females build small nests in pre-existing holes in trees, fences, or plant stems. They will also nest in artificial "nest-traps" or "bee-blocks." They build the nests as a long single column of cells, along a tube. The deepest cells are constructed first, at which point the female lays a single egg sequentially in each cell. The cells are partitioned and sealed using the construction substance noted above. Pollen, sometimes mixed with nectar, provisioned in the cell will nourish the larvae when hatched. The bees are polylectic, meaning the larvae are fed from a variety of pollen sources. Subsequently, after a few stages of molting, the larva spins a cocoon and pupates. They will overwinter as pupae. After several months, the bee will emerge in its adult form.

The typically smaller males emerge in advance of females. They will die shortly after mating. The female bees survive for another few weeks, during which time they build new nests and gather necessary provisions. Adult bees are active from April to September throughout most of the range. In Florida, that have been collected as early as February and as late as November. Flight times are typically May–October in cooler climates of their range.

Resin bee larva and a plug from a resin bee nest

Solitary bees, such as M. campanulae, do not form colonies. While social insects (ants, yellow jackets, honeybees) work in colonies, leafcutter and resin bees work independently building nests. Similar to honeybees, female bees perform nearly all essential tasks of brood rearing. M. campanulae does not produce honey, but does perform other important beneficial tasks, pollinating crops and wild plants. Although they can produce a mild sting, less intense than that from a honeybee, they are considered nonaggressive. Bees in the family Megachilidae carry pollen on the underside of their abdomen. Unlike honeybees, they do not have pollen baskets on their hind legs. Most bees in the genus are small to medium in size, although M. pluto at 38 mm is regarded as the largest bee in the world. Many bees in the genus are referred to as leafcutters. However, the mandibles of M. campanulae lack cutting edges; it belongs to the subgenus Chelostomoides, which use mud or resins to build.

Synthetic nest materials

In the wild, M. campanulae seal off their cells within the nest with natural resins found in plants and trees. In 2013, however, researchers at York University reported in the journal Ecosphere that the bees had used synthetics, including caulk, to seal the cells. Compositional analysis of these materials revealed calcium, titanium, and iron. They resembled polyurethane-based sealants typically used in building construction. It is not uncommon for insects to live inside found objects made of plastic. However, these findings are the first known of insects actually building nests with plastic. With help from citizen scientists in Toronto, over 200 nest boxes were placed throughout the city. Scanning electron microscopy, x-ray microanalysis, and infrared microscopy were employed in identification of polymeric specimens. Researchers suggest the bees' behavior is an example of adaptive behavior. Since some of the bees were free of parasites, these novel and possibly more robust methods of nest building may offer additional protection. Incorporation of plastic into the walls and sealants of the cell nests appeared to provide some protection against brood parasite invasion. However, use of these more readily available synthetic materials, in an urban setting, may be incidental. In fact, exposure of brood to polyurethane and polyethylene based plastics could be detrimental, as the Canadian team noted, since diffusion of moisture could be inhibited. Some of the brood specimens were heavily affected by mold growth. Additionally, synthetic materials in the nest might hinder the bees ability to move and breath. Toxin exposures and other effects of urbanization are well documented contributors towards pollinator decline in general.

Distribution and habitat

The range of M. campanulae covers a broad expanse of the eastern North American continent. They are native to southern Ontario. The range extends from this southeastern Canadian province, through the New England states to Florida. The range extends west, as far as Minnesota, Nebraska and Texas. A few reports of sightings further west are noted, including presence in Colorado and Montana.

Morphology and identification

Morphologically, they most resemble Megachile angelarum. The mandibles in these resin gathering bees have four dentate ridges are characteristically lacking cutting edges. Unlike M. angelarum females, they lack an apical fascia on T5. Females can also can be recognized by parallel sided metastoma. The female measures 10–12 millimetres (0.39–0.47 in) in length, while males measure 8–9 millimetres (0.31–0.35 in) in length. Females have subparallel to slightly converging compound eyes, while the males eyes are only very slightly convergent near the apex. There are also marked similarities in appearance between M. campanulae and M. exilis. The males of M. exilis have characteristically dilated and hollowed out front tarsi. The tarsi in M. campanulae are not modified and are otherwise unremarkable.

Females

M. campanulae females measure 10–12 millimeters (0.39–0.47 in) in length. They bear a strong resemblance to M. angelarum. However, the white apical fascia at T5 found on M angelarum is absent in M campanulae.

Head

The mandibles have 4 dentate ridges. They do not have the cutting edges found in leafcutter bees. Females can also can be recognized by parallel sided metastoma. They have subparallel to slightly converging compound eyes. The lateral ocelli are located close to the compound eyes than to the vertex margin. There is a distinct pair of tubercles located along the margin of the lower facial plate (clypeus). One tubercle is located on each side of the midline. These tubercles are denticulate along the lateral margin. Other facial features include gena more narrow than the compound eyes. There are punctures on the vertex and the gena. Those on the vertex are uniform in shape with rough edges, slightly separated from one another. On the gena, they are less roughened and closer together. The frons shows more coarse and closely arranged punctures. Around the eyes, there are fine punctures, closely arranged. There are narrow areas supraclypeal with some shining spaces, but also areas of coarse and deep punctuation. The pubescence on the head is short in length and pale in tone. It is less densely distributed than that on the face. The hairs are also more prominent around the antennae, gena, and inner orbits. They are more white in these areas. They become more yellow on the vertex. They are sparse on the clypeus. F1 is twice as wide as it is long. It is approximately half the length of the pedicel. It is shorter than F2, F3, flagellomeres. The apical flagellomete is 1.5 times as long.

Mesosoma

Along the lateral and posterior aspects, the pubescence is short, white, and sparsely distributed. They are more densely collected around the pronotal lobes. On the mesoscutum, they are pale, very short, and sparse in distribution. They are whiter and longer on the scutellum and they stand erect. Punctures across the mseoscutum and scutellum are coarse, deep, and closely arranged. The spaces between the mesoscutuem and scutellum are shiny and very narrow. Punctate markings are finer on the axilla, where they are closely arranged. The pleura demonstrate punctate markings which are coarse and deep. These are densely arranced anteriorly. The pleura are shiny between the punctate markings. Lateral aspects of the propodeum are dull. They are overall smooth, except some fine and shallow punctures. These are closely arranged. The basitarsi are short in comparison with the tibiae. They are also more narrow. There are yellow spurs present. There are minute and closely spaced punctures on the tegula. The basal aspects of the wings are glassy. The wing apices are less translucent. The wing veins are black.

The subspecies Megachile campanulae wilmingtoni (Mitchell) is characterized by larger size 11–12 millimetres (0.43–0.47 in), dark pubescence present on the 6th tergum, and darker wings with a brownish tinge. In the female, hairs on the pollen carrying apparatus (scopa are black at the 6th sternal segment. Distribution is along the southeastern coast of the United States into Florida, where is assumes the predominant form.

Parasites and diseases

Stelis louisae is a nest parasite of Megachile campanulae

Megachile campanulae can be parasitized by a number of brood parasites, including Monodontomerus obscurus, a Chalcid wasp. The kleptoparasitic bee Stelis louisae has been found in the nests. Members of the genus Coelioxys are also known parasites of Megachile. Kleptoparasitic bees characteristically deposit their eggs in the nests of other bees. As this behavior is similar to that of cuckoo birds, such bees are referred to as cuckoo bees. These host-parasite relationships are complex and the relationships between M. campanulae and other parasitic species may not be well described.

Mold growth was shown to be problematic in the Toronto study, particularly when synthetic materials were incorporated in nest construction. Chalk brood is known to affect the related species, M. rotundata.

Human interaction

Bee boxes provide nesting locations for M. campanulae and other bees

More focus is being directed on interactions between humans and native pollinators, such as M. campanulae. These relationships are complex, involving issues of habitat loss, pesticides and toxin exposures, climate change, and other effects on the environment. The species M. campunulae pollinates of a wide array of flowers and crops. The significance of contributions from native pollinators is gaining increased attention in the wake of declines in managed bee populations. Such declines have received substantial press, especially in relation to colony collapse disorder. In 2013, Oregon Congressman Earl Blumenauer introduced H. R. 2692 the "Save America's Pollinators Act". In addition, an International Pollinator Initiative has been developed by the Food and Agriculture of the United Nations working group. A similar project, the North American Pollinator Protection Campaign might more directly address issues specific to M. campanuale.

On a smaller scale, human behaviors adversely affecting populations of bees, such as M. campanulae, can be mediated in other ways. Many native bee species can be managed with minimal equipment. M. campanulae will nest in simple bee boxes (pictured). These are constructed in simplest form by drilling holes in a block of wood. The wood is attached to a post or wall, ideally in an area receiving adequate sun. Holes of different diameter will be suited better for different bee species. Recommended diameters are between 2 to 10 mm (0.079 to 0.394 in), with a depth of around 18 cm (7.1 in). However, these recommendations are not specific to M. campanulae. Since native pollinators forage in an area within about 500 yards (460 m) of the nest, they can increase the productivity of a small garden.

Sparse literature has been devoted to effects of pesticides on M. campunulae specifically. However, in general pesticide exposure is detrimental to native bee populations. Bees can be harmed by numerous classes of pesticides including: insecticides, fungicides, herbicides, acaricides, rodenticides (coumarins).

Pollination

M. campanulae have been documented to pollinate the following wildflowers:

See also

References

  1. "Megachile campanulae (Robertson, 1903)". Integrated Taxonomic Information System.
  2. ^ Mitchell, Theodore B (1962). The Bees of the Eastern United States II (PDF). Technical bulletin (North Carolina Agricultural Experiment Station). pp. 182–184. Retrieved 24 September 2014.
  3. ^ Sheffield, Cory S.; Ratti, Claudia; Packer, Laurence; Griswold, Terry (29 November 2011). "Megachile (Chelostomoides) campanulae (Robertson, 1903)". Canadian Journal of Arthropod Identification. York University. doi:10.3752/cjai.2011.18. ISSN 1911-2173. Retrieved 26 September 2014. Cite error: The named reference "Sheffield" was defined multiple times with different content (see the help page).
  4. Marlin, J. C. and W. E. LaBerge (2001), "The native bee fauna of Carlinville, Illinois, revisited after 75 years: a case for persistence", Conservation Ecology, 5 (1): 9
  5. ^ "Megachile campanulae (Robertson, 1903)". DiscoverLife.Org. Retrieved 20 September 2014.
  6. ^ "Genus Megachile – Leaf-cutter and Resin Bees". BugGuide.Net. Retrieved 25 September 2014.
  7. Sheffield, Cory S.; Ratti, Claudia; Packer, Laurence; Griswold, Terry (29 November 2011). "Leafcutter and Mason Bees of the Genus Megachile Latreille (Hymenoptera: Megachilidae) in Canada and Alaska". Canadian Journal of Arthropod Identification. York University. doi:10.3752/cjai.2011.18. ISSN 1911-2173. Retrieved 23 September 2014.
  8. ^ "Chelostomoides". UF/IFAS Entomology and Nematology Department. Univ of Florida. Retrieved 27 September 2014.
  9. Michener, Charles D. (June 1, 2000). The Bees of the World (1st ed.). The Johns Hopkins University Press. pp. 4–8. ISBN 0801861330.
  10. Mader, Eric; Spivak, Marla; Evans, Elaine (Feb 2010). Managing Alternative Pollinators: A Handbook for Beekeepers, Growers, and Conservationists (PDF). SARE Handbook 11, NRAES – 186. pp. 70–93. Retrieved 24 September 2014.
  11. Cranshaw, W.S. "Leafcutter Bees". Colorado State University Extension. Retrieved 21 September 2014.
  12. "Megachile bees Factsheet". BioNET-EAFRINET. Retrieved 27 September 2014.
  13. ^ "Encouraging Native Pollinators". Univ of Arkansas. Retrieved 5 October 2014.
  14. Cranshaw, Whitney; Redak, Richard (2013). Bugs Rule!: An Introduction to the World of Insects. Princeton University Press. p. 307. ISBN 9781400848928. {{cite book}}: |access-date= requires |url= (help)
  15. ^ "City bees line nests with plastic bags". University of Washington Conservation Magazine (Conservation This Week). February 13, 2014. Retrieved 5 October 2014.
  16. Goldman, Jason G (Jun 1, 2014). "Bees Living in Cities Are Building Their Homes with Plastic". Scientific American. 310 (6). Retrieved 20 September 2014.
  17. ^ Woollaston, Victoria (12 February 2014). "Urban bees are using PLASTIC to build hives – and it could stop parasites from infecting their nests". Daily Mail. UK. Retrieved 20 September 2014.
  18. "Study: Bees Using Plastic To Help Build Their Nests". CBS DC. February 5, 2014. Retrieved 21 September 2014. {{cite news}}: External link in |publisher= (help)
  19. ^ MacIvor, J Scott; Moore, Andrew E. (31 December 2013). "Bees collect polyurethane and polyethylene plastics as novel nest materials". Ecosphere. 4 (12). doi:10.1890/ES13-00308.1. Retrieved 21 September 2014.
  20. Vanbergen, Adam J; Insect Pollinators Initiative (2013). "Threats to an ecosystem service: pressures on pollinators" (PDF). Frontiers in Ecology and the Environment. doi:10.1890/120126. Retrieved 24 September 2014.
  21. Murray, Tom. "Species Megachile campanulae – Bellflower Resin Bee". BugGuide.net. Retrieved 20 September 2014.
  22. Scott, Virginia L.; Ascher, John S.; Griswold, Terry; Nufio, César R. (September 1, 2011). "The Bees of Colorado (Hymenoptera: Apoidea: Anthophila)" (PDF). Natural History Inventory of Colorado (23). University of Colorado Museum of Natural History. ISSN 0890-6882. Retrieved 24 October 2014.
  23. Macivor, J. Scott; Salehi, Baharak (1 August 2014). "Bee Species-Specific Nesting Material Attracts a Generalist Parasitoid: Implications for Co-occurring Bees in Nest Box Enhancements". Environmental Entomology. 43 (4): 1027–1033. doi:10.1603/EN13241.
  24. "Bees of the Week: genus Coelioxys". The bees needs. Retrieved 6 October 2014.
  25. James, Rosalind R. "Temperature and chalkbrood development in the alfalfa leafcutting bee,". Apidologie. 36 (1): 15–23. doi:10.1051/apido:2004065.
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  27. "Text of the Saving America's Pollinators Act of 2013". www.govtack.us. Retrieved 5 October 2014.
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  31. "Why worry about native bees?". Native Bee Conservancy. Retrieved 6 October 2014.

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