Megachilidae

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Megachilidae
Temporal range: Middle Eocene � recent, 45�0 Ma
Pre??OSDCPTJKPgN
Anthidium manicatum male.jpg
Male European wool carder bee, Anthidium manicatum
Scientific classification e
Kingdom: Animalia
Clade: Euarthropoda
Class: Insecta
Order: Hymenoptera
Suborder: Apocrita
Superfamily: Apoidea
Family: Megachilidae
Subfamilies
Fideliinae
Megachilinae

A leaf-cutter bee showing abdominal scopa

Leaves showing cuts by a leafcutter bee

Megachilidae is a cosmopolitan family of mostly solitary bees whose pollen-carrying
structure (called a scopa) is restricted to the ventral surface of the abdomen
(rather than mostly or exclusively on the hind legs as in other bee families).
Megachilid genera are most commonly known as mason bees and leafcutter bees,
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 cleptoparasites (informally called "cuckoo bees"), 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.

Contents
1 Lifecycle
1.1 Nonparasitic species
1.2 Parasitic species
2 Diversity
3 Evolution and taxonomy
4 References
5 External links
Lifecycle
Nonparasitic species

Lifecycle
The lifecycle of nonparasitic Megachilidae is typically that nests are built,
divided 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
cells in a mass of clay or resin attached to a
Nest cavities are often linear, for example in
(snail shells are used by some Osmia, and some
cavities).
cavities. Some embed individual
wall, rock surface, or plant stem.
hollow plant stems, but not always
species readily use irregular

Parasitic species
Some genera of megachilids are brood parasites, 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 published a
preliminary report on parasitic megachilid bees of the western United States.[1]

Diversity
North America has an estimated 630 different megachilid species, including
Megachile, Osmia, Anthidium, Hoplitis, and Chalicodoma. Most Megachilidae are
native, and a few are introduced, accidentally and intentionally; globally the
number of species identified exceeds 4,000.[2][3] 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 �???? (m�gas, �big�) + ?e???? (khe�los, �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 specie numbers (~70% 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).[4]

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 bees), used extensively in alfalfa pollination, and the western native
and frequently raised Osmia lignaria (the orchard mason bee or blue orchard bee),
used in orchard pollination. Other Osmia and Megachile species are also in
commercial use in North America, Europe and Asia.

Many Megachilidae not cultivated extend the diversity of this family, separated
typically by their choice of building materials. 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.� [5] Like most Anthidium, rather
than cutting leaves or petals, A. manicatum scrapes the hairs from leaves to use
for nesting material.[6] 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 most any intruder.[7]

Neither the introduced Anthidium nor its American cousins are considered parasites,
only territorial and at times aggressive. But some Megachilidae are, including
Coelioxys, a cleptoparasite mostly of leafcutter bees, and Stelis, a cleptoparasite
of leafcutter and mason bees.

While some Megachilidae are extensively studied for their commercial possibilities
(or impacts), others are studied by happenstance. 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, with his writings made further
famous by his English translator Alexander Teixeira de Mattos (The Mason Bee).
Chalicodoma typically uses grit rather than mud in nest construction, along with
other differences. (Fabre wrote many observations, including of other Megachilidae,
from his home in France; his writings inspired many future researchers and
enthusiasts, from Charles Darwin to Gerald Durrell.)

Evolution and taxonomy

Carder bee (Anthidium manicatum), female

Male Haetosmia vechti, foraging on Heliotropium in Mevo Horon, West Bank

Coelioxys acanthura, female

Osmia ribifloris

Male Anthidium florentinum visiting Lantana

The fossil record for megachilid bees is poor, but a Middle Eocene dicotyledonous
leaf shows definite semicircular cutouts along its margin, implying that leaf-
cutting bees existed at that time.[8] Multiply-cut leaves and rare body fossils
from the Eocene of Germany and the Paleocene of France suggest that Megachilinae
began cutting leaves early in their evolution.[9] Phylogenetic analysis yields an
age consistent with this Eocene origin for the group.[10]

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, but they
are oligolectic (specialized on a few subfamilies of Asteraceae) and use resin from
conifers, as well as plant fibers and sand, as cell wall material.
Hofferia
Hoplitis
Hoplosmia
Noteriades
Ochreriades
Osmia
Othinosmia
Protosmia
Pseudoheriades
Stenoheriades
Stenosmia
Wainia
Xeroheriades
Tribe Anthidiini
Acedanthidium
Afranthidium
Afrostelis
Anthidiellum
Anthidioma
Anthidium
Anthodioctes
Apianthidium
Aspidosmia
Austrostelis
Aztecanthidium
Bathanthidium
Benanthis
Cyphanthidium
Dianthidium
Duckeanthidium
Eoanthidium
Epanthidium
Euaspis
Gnathanthidium
Hoplostelis
Hypanthidioides
Hypanthidium
Icteranthidium
Indanthidium
Larinostelis
Neanthidium
Notanthidium
Pachyanthidium
Paranthidium
Plesianthidium
Pseudoanthidium
Rhodanthidium
Serapista
Stelis Panzer and related genera (stelidine bees) are cleptoparasites on other
Megachilidae. They belong to the tribe Anthidiini. Bees in the subgenus
Heterostelis are parasitic on Trachusa.
Trachusa
Trachusoides
Xenostelis
Tribe Dioxyini
Aglaoapis
Allodioxys
Dioxys is a brood parasite of Megachile, Anthidium, and Osmia.
Ensliniana
Eudioxys
Metadioxys
Paradioxys
Prodioxys
Tribe Megachilini
Coelioxys is a brood parasite of Megachile. Females have a pointed conic abdominal
apex (tip); males have several spikes on their apices.
Megachile
Radoszkowskiana
incertae sedis
Neochalicodoma
Stellenigris
References
Cockerell, Theodore; Lutz, Frank Eugene (1 December 1921). "Some parasitic
megachilid bees of the western United States" (PDF). American Museum Novitates. 21.
Retrieved 5 October 2014.
BugGuide. "Megachilidae". BugGuide. Retrieved 29 September 2017.
>630 spp. in 18 genera of 4 tribes and 2 subfamilies in our area, ~4,100 spp. in
~80 genera of 11 tribes and 4 subfamilies worldwide(1)
Krombein, Karl V. (1967). Trap-nesting wasps and bees: life histories, nests, and
associates. Smithsonian Press. p. 2. Retrieved 29 September 2017.
Gonzalez, V. H. and Griswold, T. L. (1 June 2013). "Wool carder bees of the genus
Anthidium in the Western Hemisphere (Hymenoptera: Megachilidae): diversity, host
plant associations, phylogeny, and biogeography". Zoological Journal of the Linnean
Society. 168 (2): 221�425. doi:10.1111/zoj.12017. Retrieved 29 September 2017.
Michener, C. D. (2000). Bees of the World. Baltimore: Johns Hopkins University
Press. p. 913.
Honey Bee Suite. "Native pollinator - wool carder bee". honeybeesuite.com. Rusty
Burlew. Retrieved 29 September 2017.
Victor H. (June 2008). Phylogeny and Classification of the Bee Tribe Megachilini
(Hymenoptera: Apoidea, Megachilidae), with Emphasis on the Genus Megachile. UMI
3316016. ProQuest / University of Kansas (PhD thesis). pp. 54�56. ISBN 978-0-549-
68378-0.
Wedmann, Soni a; Wappler, Torsten; Engel, Michael S. (June 2009). "Direct and
indirect fossil records of megachilid bees from the Paleogene of Central Europe
(Hymenoptera: Megachilidae)". Naturwissenschaften. 96 (6): 703�712.
doi:10.1007/s00114-009-0525-x. PMID 19296064.
Cardinal, Sophie; Danforth, Bryan N. (January 2013). "Bees diversified in the age
of eudicots". Proceedings of the Royal Society B: Biological Sciences. 280:
20122686. doi:10.1098/rspb.2012.2686. PMC 3574388?Freely accessible. PMID 23363629.
External links
Wikimedia Commons has media related to Megachilidae.
Palaearctic Osmiine Bees
Leafcutting bees on the UF / IFAS Featured Creatures Web site
Online Guides to all the eastern North American Megachilidae
Bugguide. Information and photos of Megachilidae of North America
[show] v t e
Extant Hymenopteran families
Taxon identifiers
Wd: Q530200 ADW: Megachilidae BugGuide: 84 EoL: 7500 EPPO: 1MEGCF Fossilworks:
72088 GBIF: 7911 ITIS: 154365 NCBI: 124286
Categories: MegachilidaeBee familiesCenozoic insectsExtant Eocene first appearances
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