Megachilidae is a
cosmopolitan family
In biogeography, cosmopolitan distribution is the term for the range of a taxon that extends across all or most of the world in appropriate habitats. Such a taxon, usually a species, is said to exhibit cosmopolitanism or cosmopolitism. The extr ...
of mostly solitary
bees. Both that their pollen-carrying structure (called a ''
scopa
''Scopa'' (; literally "broom") is an Italian card game, and one of the three major national card games in Italy, the others being '' Briscola'' and ''Tresette''. It is also popular in Argentina and Brazil, brought in by Italian immigrants, most ...
'') is restricted to the ventral surface of the abdomen (rather than mostly or exclusively on the hind legs as in other bee families), and their typically elongated
labrum is characteristic of this family.
[ C. D. Michener (2007) ''The Bees of the World'', 2nd Edition, pg. 122, Johns Hopkins University Press.] Megachilid genera are most commonly known as
mason bee
Mason bee is a name now commonly used for species of bees in the genus ''Osmia'', of the family Megachilidae. Mason bees are named for their habit of using mud or other "masonry" products in constructing their nests, which are made in naturally ...
s and
leafcutter bees
The genus ''Megachile'' is a cosmopolitan group of solitary bees, often called leafcutter bees or leafcutting bees; it also includes the called resin bees and mortar bees. While other genera within the family Megachilidae may chew leaves or pet ...
, reflecting the materials from which they build their nest cells (soil or leaves, respectively); a few collect plant or animal hairs and fibers, and are called carder bees, while others use plant resins in nest construction and are correspondingly called resin bees. All species feed on nectar and pollen, but a few are
kleptoparasite
Kleptoparasitism (etymologically, parasitism by theft) is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, which can mean when fo ...
s (informally called "
cuckoo bee
The term cuckoo bee is used for a variety of different bee lineages which have evolved the kleptoparasitism, kleptoparasitic behaviour of laying their eggs in the nests of other bees, reminiscent of the behavior of cuckoo birds. The name is perhap ...
s"), feeding on pollen collected by other megachilid bees. Parasitic species do not possess scopae. The motion of Megachilidae in the reproductive structures of flowers is energetic and swimming-like; this agitation releases large amounts of
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 ...
.
Life cycle
Nonparasitic species
Nonparasitic Megachilidae typically divide their nests into cells. Each cell receives a supply of food (pollen or a pollen/nectar mix) and an egg; after finding a suitable spot (often near where she emerged), a female starts building a first cell, stocks it, and oviposits. She builds a wall that separates the completed cell from the next one. The larva hatches from the egg and consumes the food supply. After moulting a few times, it spins a cocoon and pupates. It emerges from the nest as an adult. Males die shortly after mating, but females survive for another few weeks, during which they build new nests.
Nests are often built in natural or artificial cavities. Some embed individual cells in a mass of clay or resin attached to a wall, rock surface, or plant stem. Nest cavities are often linear, for example in hollow plant stems, but not always (snail shells are used by some ''Osmia'', and some species readily use irregular cavities).
Parasitic species
Some genera of megachilids are
brood parasite
Brood parasites are animals that rely on others to raise their young. The strategy appears among birds, insects and fish. The brood parasite manipulates a host, either of the same or of another species, to raise its young as if it were its own ...
s, so have no ventral scopa (e.g. ''
Stelis'' and ''
Coelioxys''). They often parasitize related taxa. They typically enter the nest before it is sealed and lay their eggs in a cell. After hatching, the parasite larva kills the host larva, unless the female parasite has already done so, and then consumes the provisions. Parasitic species are of equal size or smaller than their victims. In 1921, the journal ''
American Museum Novitates
''American Museum Novitates'' is a peer-reviewed academic journal published by the American Museum of Natural History. It was established in 1921. According to the ''Journal Citation Reports'', the journal has a 2013 impact factor
The impact f ...
'' published a preliminary report on parasitic megachilid bees of the western United States.
Diversity
North America has an estimated 630 different megachilid species, including ''
Megachile
The genus ''Megachile'' is a cosmopolitan group of solitary bees, often called leafcutter bees or leafcutting bees; it also includes the called resin bees and mortar bees. While other genera within the family Megachilidae may chew leaves or pet ...
'', ''
Osmia
Mason bee is a name now commonly used for species of bees in the genus ''Osmia'', of the family Megachilidae. Mason bees are named for their habit of using mud or other "masonry" products in constructing their nests, which are made in naturally ...
'', ''
Anthidium
''Anthidium'' is a genus of bees often called carder or potter bees, who use conifer resin, plant hairs, mud, or a mix of them to build nests. They are in the family Megachilidae which is cosmopolitan in distribution and made up of species that ...
'', ''
Hoplitis
''Hoplitis'' is a genus of bees in the family Megachilidae
Megachilidae is a cosmopolitan family of mostly solitary bees. Both that their pollen-carrying structure (called a '' scopa'') is restricted to the ventral surface of the abdomen ( ...
'', and ''
Chalicodoma
''Chalicodoma'' is a subgenus of the bee genus ''Megachile'' in the family Megachilidae
Megachilidae is a cosmopolitan family of mostly solitary bees. Both that their pollen-carrying structure (called a '' scopa'') is restricted to the vent ...
''. Most Megachilidae are native, and a few are introduced, accidentally and intentionally; globally the number of species identified exceeds 4,000.
Thus Megachilidae represent 15% to 20% of named species of bees.
The scientific name Megachilidae refers to the genus ''Megachile'', translating roughly as ''large lipped'' (Ancient Greek μέγᾰς (mégas, "big") + χεῖλος (kheîlos, "lip"); their "large lips" and strong jaws are well-suited for collection of nest building materials.
Most Megachilidae build their nests in above-ground cavities; they all are solitary bees. Their nesting habits means that in some studies of bee diversity, this bee family is most likely to be the one encountered, even though the many ground nesting bees are much greater in species numbers (~70% of all bee species are ground nesters). For example, in Krombein's trap-nesting survey (1967), almost all bees that nested in his offerings were Megachilid species—40 of 43 occupying bee species. (They were outnumbered in diversity by almost twice as many species of wasps (75) that utilized the nests).
Because they are (mostly) above-ground nesters and more commonly attracted to artificial nests, megachilid bees are also more commonly cultivated than ground nesting solitary bees. They accept nesting materials made from hollow stems, tubes, and blocks with preformed holes ("nest blocks"), and several megachilids have become important species for
agricultural / horticultural pollination. In North America these cultivated bees include the introduced ''Megachile rotundata'', (
alfalfa leafcutter bee
''Megachile rotundata'', the alfalfa leafcutting bee, is a European bee that has been introduced to various regions around the world. As a solitary bee species, it does not build colonies or store honey, but is a very efficient pollinator of alfa ...
s), used extensively in alfalfa pollination, and the western native and frequently raised ''
Osmia lignaria
''Osmia lignaria'', commonly known as the orchard mason bee or blue orchard bee, is a megachilid bee that makes nests in reeds and natural holes, creating individual cells for its brood that are separated by mud dividers. Unlike carpenter bees, ...
'' (the orchard mason bee or blue orchard bee), used in
orchard
An orchard is an intentional plantation of trees or shrubs that is maintained for food production. Orchards comprise fruit- or nut-producing trees which are generally grown for commercial production. Orchards are also sometimes a feature of larg ...
pollination. Other ''Osmia'' and ''Megachile'' species are also in commercial use in North America, Europe and Asia.
A suite of megachilid rely on plant resins for nest construction. These "resin bees" are typically smaller than honey bees, and effective pollinators, although the hard glue-like resins can complicate management of other tunnel nesting bees. Carder bees, ''Anthidium'', are unique for using plant fibers; there are 80 to 90 species of them in North America. Ironically, a non-native is best known—''A. manicatum'', the European wool carder bee, was accidentally introduced to the Americas in the late '60s and has now spread across the continent. It has been described as "... perhaps the most widely distributed unmanaged bee species in the world."
Like most ''Anthidium'', rather than cutting leaves or petals, ''A. manicatum'' scrapes the hairs from leaves to use for nesting material. It is atypical because the male is larger than the female and constantly on patrol, protecting a "harem" by chasing and even attacking all interlopers including honey and bumble bees, its tail equipped with multiple prongs that can knife in between the segments of almost any intruder.
Neither the introduced ''Anthidium'' nor its American cousins are considered parasites, only territorial and at times aggressive, though some genera are, including ''
Coelioxys'' (
kleptoparasite
Kleptoparasitism (etymologically, parasitism by theft) is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, which can mean when fo ...
s mostly of leafcutter bees), and ''
Stelis'' (kleptoparasites of leafcutter and mason bees).
While some Megachilidae are extensively studied for their commercial possibilities (or impacts), others are studied by
naturalists. ''Chalicodoma'' mason bees, not commonly cultivated, are known through extended observation and writings in the last half of the 19th century by
Jean-Henri Fabre
Jean-Henri Casimir Fabre (21 December 1823 – 11 October 1915) was a French naturalist, entomologist, and author known for the lively style of his popular books on the lives of insects.
Biography
Fabre was born on 21 December 1823 in Saint-L ...
, with his writings made further famous by his English translator
Alexander Teixeira de Mattos
Alexander Louis Teixeira de Mattos (April 9, 1865 – December 5, 1921), known as Alexander Teixeira de Mattos, was a Dutch-English journalist, literary critic and publisher, who gained his greatest fame as a translator.
Early life
The Teix ...
("The Mason Bee"); Fabre wrote many observations, including of other Megachilidae, from his home in France, and his writings inspired many future researchers and enthusiasts, from
Charles Darwin
Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended fr ...
to
Gerald Durrell
Gerald Malcolm Durrell, (7 January 1925 – 30 January 1995) was a British naturalist, writer, zookeeper, conservationist, and television presenter. He founded the Durrell Wildlife Conservation Trust and the Jersey Zoo on the Channel Island o ...
. ''Chalicodoma'' typically uses grit rather than mud in nest construction, along with other differences.
Leafcutter bee by Bernhard plank.jpg, A leafcutter bee cutting a leaf
Megachile-pjt.jpg, ''Megachile sp.'' with cut leaf
Leafcutting 1 6431.JPG, Leaves cut by ''Megachile'' sp.
Leaf_cutter_bee_nests.jpg, Nests of ''Megachile'' bees from Bangalore, India.
Leaf_cutter_bee_nest.jpg, Dissected nest of a ''Megachile'' bee
Cut_leaf_bits_from_a_Leaf-cutter_Bee.jpg, Dissected cut leaf bits from a nest.
Evolution and taxonomy
The fossil record for megachilid bees is poor, but a
Middle Eocene
The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', "dawn ...
dicotyledonous leaf shows definite semicircular cutouts along its margin, implying that leaf-cutting bees existed at that time. Multiply-cut leaves and rare body fossils from the
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
of Germany and the
Paleocene
The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
of France suggest that Megachilinae began cutting leaves early in their evolution. Phylogenetic analysis yields an age consistent with this Eocene origin for the group.
*Subfamily Fideliinae
**Tribe
Pararhophitini
***''
Pararhophites''
**Tribe
Fideliini
***''
Fidelia''
***''
Neofidelia''
*Subfamily
Megachilinae
**Tribe
Lithurgini
***''
Lithurgus''
***''
Microthurge''
***''
Trichothurgus''
**Tribe
Osmiini
***''
Afroheriades''
***''
Ashmeadiella''
***''
Atoposmia''
***''
Bekilia''
***''
Chelostoma''
***''
Haetosmia''
***''
Heriades'' species have narrow abdominal bands. They resemble small ''
Osmia
Mason bee is a name now commonly used for species of bees in the genus ''Osmia'', of the family Megachilidae. Mason bees are named for their habit of using mud or other "masonry" products in constructing their nests, which are made in naturally ...
'', but they are
oligolectic
The term oligolecty is used in pollination ecology to refer to bees that exhibit a narrow, specialized preference for pollen sources, typically to a single family or genus of flowering plants. The preference may occasionally extend broadly to mult ...
(specialized on a few subfamilies of
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 ...
) and use resin from conifers, as well as plant fibers and sand, as cell wall material.
***''
Hofferia''
***''
Hoplitis
''Hoplitis'' is a genus of bees in the family Megachilidae
Megachilidae is a cosmopolitan family of mostly solitary bees. Both that their pollen-carrying structure (called a '' scopa'') is restricted to the ventral surface of the abdomen ( ...
''
***''
Hoplosmia''
***''
Noteriades''
***''
Ochreriades
Ochreriades is a suprageneric lineage within Megachilidae. This genus can be considered the sister lineage "to a clade consisting of the “core” Osmiini, the tribe Megachilini and the genera ''Pseudoheriades''and ''Afroheriades."''Litman, Jess ...
''
***''
Osmia
Mason bee is a name now commonly used for species of bees in the genus ''Osmia'', of the family Megachilidae. Mason bees are named for their habit of using mud or other "masonry" products in constructing their nests, which are made in naturally ...
''
***''
Othinosmia''
***''
Protosmia
''Protosmia'' is a genus of subgenus ''Chelostomopsis'' in the family Megachilidae. There are more than 30 described species in ''Protosmia''.
Species
These 31 species belong to the genus ''Protosmia'':
* ''Protosmia asensioi'' Griswold & Parke ...
''
***''
Pseudoheriades''
***''
Stenoheriades''
***''
Stenosmia''
***''
Wainia''
***''
Xeroheriades''
**Tribe
Anthidiini
Anthidiini is a tribe of insects in the family Megachilidae. There are at least 40 genera and 840 described species in Anthidiini.S There is strong evidence that the tribe is monophyletic.
Genera
* '' Acedanthidium'' Michener, 2000
* '' Afranth ...
***''
Acedanthidium''
***''
Afranthidium''
***''
Afrostelis''
***''
Anthidiellum''
***''
Anthidioma''
***''
Anthidium
''Anthidium'' is a genus of bees often called carder or potter bees, who use conifer resin, plant hairs, mud, or a mix of them to build nests. They are in the family Megachilidae which is cosmopolitan in distribution and made up of species that ...
''
***''
Anthodioctes''
***''
Apianthidium''
***''
Aspidosmia''
***''
Austrostelis''
***''
Aztecanthidium''
***''
Bathanthidium
''Bathanthidium'' is a genus of bees belonging to the family Megachilidae. The species of this genus are found in Southeastern Asia.
Species
*''Bathanthidium atriceps''
*''Bathanthidium barkamense''
*''Bathanthidium bicolor''
*''Bathanthidiu ...
''
***''
Benanthis''
***''
Cyphanthidium''
***''
Dianthidium''
***''
Duckeanthidium''
***''
Eoanthidium''
***''
Epanthidium''
***''
Euaspis
''Euaspis'' is a genus of bees belonging to the family Megachilidae.
The species of this genus are found in Africa and Southeastern Asia.
Species:
* ''Euaspis aequicarinata'' Pasteels, 1980
* ''Euaspis basalis'' (Ritsema, 1874)
References ''
***''
Hoplostelis''
***''
Hypanthidioides''
***''
Hypanthidium
''Hypanthidium'' is a genus of bees belonging to the family Megachilidae
Megachilidae is a cosmopolitan family of mostly solitary bees. Both that their pollen-carrying structure (called a ''scopa'') is restricted to the ventral surface of th ...
''
***''
Icteranthidium''
***''
Indanthidium''
***''
Larinostelis''
***''
Notanthidium''
***''
Pachyanthidium''
***''
Paranthidium''
***''
Plesianthidium''
***''
Pseudoanthidium''
***''
Rhodanthidium''
***''
Serapista''
***''
Stelis''
Panzer and related genera (stelidine bees) are
kleptoparasite
Kleptoparasitism (etymologically, parasitism by theft) is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, which can mean when fo ...
s on other Megachilidae. They belong to the tribe
Anthidiini
Anthidiini is a tribe of insects in the family Megachilidae. There are at least 40 genera and 840 described species in Anthidiini.S There is strong evidence that the tribe is monophyletic.
Genera
* '' Acedanthidium'' Michener, 2000
* '' Afranth ...
. Bees in the subgenus ''
Heterostelis'' are parasitic on ''
Trachusa''.
***''
Trachusa''
***''
Trachusoides''
***''
Xenostelis''
**Tribe
Dioxyini
Dioxyini is a tribe of cuckoo bees in the family Megachilidae. There are eight genera in Dioxyini, comprising 36 species.
Genera
* '' Aglaoapis'' Cameron, 1901
* '' Allodioxys'' Popov, 1947
* '' Dioxys'' Lepeletier & Serville, 1825
* ''Ensliniana ...
***''
Aglaoapis
''Aglaoapis'' is a genus of cuckoo bee belonging to the family Megachilidae
Megachilidae is a cosmopolitan family of mostly solitary bees. Both that their pollen-carrying structure (called a '' scopa'') is restricted to the ventral surface ...
''
***''
Allodioxys''
***''
Dioxys'' is a brood parasite of ''
Megachile
The genus ''Megachile'' is a cosmopolitan group of solitary bees, often called leafcutter bees or leafcutting bees; it also includes the called resin bees and mortar bees. While other genera within the family Megachilidae may chew leaves or pet ...
'', ''
Anthidium
''Anthidium'' is a genus of bees often called carder or potter bees, who use conifer resin, plant hairs, mud, or a mix of them to build nests. They are in the family Megachilidae which is cosmopolitan in distribution and made up of species that ...
'', and ''
Osmia
Mason bee is a name now commonly used for species of bees in the genus ''Osmia'', of the family Megachilidae. Mason bees are named for their habit of using mud or other "masonry" products in constructing their nests, which are made in naturally ...
''.
***''
Ensliniana
The cuckoo bee genus ''Ensliniana'' contains only one species, ''Ensliniana bidentata'', which is found in the Mediterranean and Middle East
The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly e ...
''
***''
Eudioxys''
***''
Metadioxys''
***''
Paradioxys''
***''
Prodioxys''
**Tribe
Megachilini
***''
Coelioxys'' is a brood parasite of ''
Megachile
The genus ''Megachile'' is a cosmopolitan group of solitary bees, often called leafcutter bees or leafcutting bees; it also includes the called resin bees and mortar bees. While other genera within the family Megachilidae may chew leaves or pet ...
''. Females have a pointed conic abdominal apex (tip); males have several spikes on their apices.
***''
Megachile
The genus ''Megachile'' is a cosmopolitan group of solitary bees, often called leafcutter bees or leafcutting bees; it also includes the called resin bees and mortar bees. While other genera within the family Megachilidae may chew leaves or pet ...
''
***''
Radoszkowskiana''
**''
incertae sedis
' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
''
***''
Neochalicodoma''
***''
Stellenigris''
References
External links
Palaearctic Osmiine Beeson the
UF /
IFAS Featured Creatures Web site
Online Guides to all the eastern North American MegachilidaeBugguide. Information and photos of Megachilidae of North America*
{{Taxonbar, from=Q530200
Bee families
Cenozoic insects
Extant Eocene first appearances