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The
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...
Euglossini, in 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 '' ...
, commonly known as orchid bees or euglossine bees, are the only group of corbiculate bees whose non-parasitic members do not all possess eusocial behavior.


Description

Most of the tribe's species are solitary, though a few are communal, or exhibit simple forms of eusociality. There are about 200 described species, distributed in five genera: ''
Euglossa ''Euglossa'' is a genus of orchid bees (Euglossini). Like all their close relatives, they are native to the Neotropics; an introduced population exists in Florida. They are typically bright metallic blue, green, coppery, or golden. ''Euglossa in ...
'', ''
Eulaema ''Eulaema'' is a genus of large-bodied euglossine bees that occur primarily in the Neotropics. They are robust brown or black bees, hairy or velvety, and often striped with yellow or orange, typically resembling bumblebees. They lack metallic co ...
'', ''
Eufriesea ''Eufriesea'' is a genus of euglossine bees. Like all orchid bees, they are restricted to the Neotropics. All species range from entirely to at least partially metallic (the face and/or tegulae), though much of the body in some species may be b ...
'', ''
Exaerete ''Exaerete'' is a genus of euglossine bees found from Mexico to northern Argentina. Like all orchid bees, they are restricted to the Neotropics. All but one species is metallic green, and they are cleptoparasites in the nests of other euglossine ...
'' and the monotypic ''
Aglae ''Aglae'' is a genus of Euglossini, euglossine bees, with the only described species ''Aglae caerulea''. Like all orchid bees, it is restricted to the Neotropics. They are metallic blue. This species, like the genus ''Exaerete'', is a nest paras ...
''. All exclusively occur in South or Central America (though one species, ''
Euglossa dilemma ''Euglossa dilemma'', the green orchid bee or dilemma orchid bee, is a species of solitary euglossine bee native to a broad area of Central America, and recently introduced to Florida in the United States. It was first detected in Broward County ...
'', has become established in 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 ...
). The genera ''Exaerete'' and ''Aglae'' are kleptoparasites in the nests of other orchid bees. All except ''Eulaema'' are characterized by brilliant metallic coloration, primarily green, gold, and blue. Females gather pollen and nectar as food from a variety of plants, and resins, mud and other materials for nest building. Some of the same food plants are also used by the males, which leave the nest upon hatching and do not return.Williams & Whitten, 1983


Fragrance collection

Male orchid bees have uniquely modified legs which are used to collect and store different volatile compounds (often
ester In chemistry, an ester is a compound derived from an oxoacid (organic or inorganic) in which at least one hydroxyl group () is replaced by an alkoxy group (), as in the substitution reaction of a carboxylic acid and an alcohol. Glycerides ar ...
s) throughout their lives, primarily from
orchids Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Along with the Asteraceae, they are one of the two largest families of flowering ...
in the subtribes
Stanhopeinae Stanhopeinae is a subtribe of plants in the tribe Cymbidieae. The subtribe in the strict sense, have viscidia and stipes that are thin and strap-like, they are adapted for attachment to edge of the bee's scutellum or to a leg. Pseudobulbs are u ...
and Catasetinae, where all species are exclusively pollinated by euglossine males. These orchids do not produce nectar, and hide the pollen on a single
anther The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...
under an anther cap; orchids are not visited by females, as females require both nectar and pollen as food provisions for their offspring, and visit other types of plants to obtain these resources. The whole
pollinarium A pollinium (plural pollinia) is a coherent mass of pollen grains in a plant that are the product of only one anther, but are transferred, during pollination, as a single unit. This is regularly seen in plants such as orchids and many species of mil ...
becomes attached to the male as it leaves the flower. Several flowers from other plant families are also visited by the bees: ''
Spathiphyllum ''Spathiphyllum'' is a genus of about 47 species of monocotyledonous flowering plants in the family Araceae, native to tropical regions of the Americas and southeastern Asia. Certain species of ''Spathiphyllum'' are commonly known as spath or pea ...
'' and ''
Anthurium ''Anthurium'' (; Schott, 1829) is a genus of about 1,000Mantovani, A. and T. E. Pereira. (2005)''Anthurium'' (section ''Urospadix''; subsection ''Flavescentiviridia'').''Rodriguesia'' 56(88), 145–60. species of flowering plants, the largest ...
'' (
Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe (or leaf-like bract). Also ...
), ''
Drymonia ''Drymonia'' is a genus of moths of the family Notodontidae erected by Jacob Hübner in 1819. It consists of the following species: * ''Drymonia dodonaea'' (Denis & Schiffermüller, 1775) * ''Drymonia dodonides'' (Staudinger, 1887) * ''Drymonia ...
'' and '' Gloxinia'' (
Gesneriaceae Gesneriaceae, the gesneriad family, is a family of flowering plants consisting of about 152 genera and ca. 3,540 species in the tropics and subtropics of the Old World (almost all Didymocarpoideae) and the New World (most Gesnerioideae), with ...
), ''
Cyphomandra ''Cyphomandra'' was a formerly accepted genus in the plant family Solanaceae (the nightshades and relative). It used to contain about 35 species native to the Americas from Mexico southwards to Northern Argentina. Recent authors have treated ...
'' (
Solanaceae The Solanaceae , or nightshades, are a family of flowering plants that ranges from annual and perennial herbs to vines, lianas, epiphytes, shrubs, and trees, and includes a number of agricultural crops, medicinal plants, spices, weeds, and orn ...
), and ''
Dalechampia ''Dalechampia'' is a genus of plant of the family Euphorbiaceae and of the monogeneric subtribe Dalechampiinae. It is widespread across lowland tropical areas (generally below 2,000 m ASL) primarily in the Americas with smaller numbers of spec ...
'' (
Euphorbiaceae Euphorbiaceae, the spurge family, is a large family of flowering plants. In English, they are also commonly called euphorbias, which is also the name of a genus in the family. Most spurges, such as ''Euphorbia paralias'', are herbs, but some, e ...
) contain one or more species that attract male euglossines. The chemicals are picked up using special brushes on the forelegs, transferred from there by rubbing the brushes against combs on the middle legs, and finally these combs are pressed into grooves on the dorsal edge of the hind legs, squeezing the chemicals past the waxy hairs which block the opening of the groove, and into a sponge-like cavity inside the hind tibia. The accumulated "fragrances" are evidently released by the males at their display sites in the forest understory, where matings are known to take place. The accumulated volatiles were long believed to be used by males as a pheromone to attract females; however, female attraction to male odors or to orchid fragrances has never been demonstrated in behavioral experiments. Instead, it is now thought that the function of the male odors is to signal male 'genetic quality' to females, because great effort must be expended by males to collect orchid fragrances and thus only the most fit males could gather complex odor mixes. This would constitute an unusual example of Zahavi's
handicap principle The handicap principle is a hypothesis proposed by the biologist Amotz Zahavi to explain how evolution may lead to "honest" or reliable signalling between animals which have an obvious motivation to bluff or deceive each other. It suggests that ...
, analogous to the male peacock's tail. The relationship between male euglossine bees and volatile chemicals is essentially unique in the animal kingdom. Scientists use single synthetic compounds as bait to attract and collect males for study; among them are many familiar flavorings and odors considered appealing to humans (e.g.,
methyl salicylate Methyl salicylate (oil of wintergreen or wintergreen oil) is an organic compound with the formula C8H8O3. It is the methyl ester of salicylic acid. It is a colorless, viscous liquid with a sweet, fruity odor reminiscent of root beer, but often a ...
,
eugenol Eugenol is an allyl chain-substituted guaiacol, a member of the allylbenzene class of chemical compounds. It is a colorless to pale yellow, aromatic oily liquid extracted from certain essential oils especially from clove, nutmeg, cinnamon, basil ...
,
cineole Eucalyptol is a monoterpenoid. A colorless liquid, it is a bicyclic ether. Eucalyptol has a fresh mint-like smell and a spicy, cooling taste. It is insoluble in water, but miscible with organic solvents. Eucalyptol makes up ~70% - 90% of eucal ...
,
benzyl acetate Benzyl acetate is an organic ester with the molecular formula . It is formed by the condensation of benzyl alcohol and acetic acid. Similar to most other esters, it possesses a sweet and pleasant aroma, owing to which, it finds applications in pe ...
,
methyl benzoate Methyl benzoate is an organic compound. It is an ester with the chemical formula C6H5CO2CH3. It is a colorless liquid that is poorly soluble in water, but miscible with organic solvents. Methyl benzoate has a pleasant smell, strongly reminisce ...
,
methyl cinnamate Methyl cinnamate is the methyl ester of cinnamic acid and is a white or transparent solid with a strong, aromatic odor. It is found naturally in a variety of plants, including in fruits, like strawberry, and some culinary spices, such as Sichuan p ...
), and others which are not (e.g.,
skatole Skatole or 3-methylindole is an organic compound belonging to the indole family. It occurs naturally in the feces of mammals and birds and is the primary contributor to fecal odor. In low concentrations, it has a flowery smell and is found in seve ...
). It is also important to note that resource 'hot spots' wax and wane throughout the year as plants bloom and die, largely due to temporal changes, particularly between the changing of seasons. This often shifts euglossine bee preferences for certain chemicals over others. For '' Euglossa imperialis'', studies have shown that there is a significant trend in chemical preference for cineole during later times in the year as opposed to methyl salicylate. In the local fragrance environment, a shift in the wind direction is another factor which may also cause another fragrance 'hot spot' to be included in the odor plume for euglossine bees. Neotropical orchids themselves often exhibit elaborate adaptations involving highly specific placement of pollen packets (
pollinia A pollinium (plural pollinia) is a coherent mass of pollen grains in a plant that are the product of only one anther, but are transferred, during pollination, as a single unit. This is regularly seen in plants such as orchids and many species of mil ...
) on the bodies of the male orchid bees; the specificity of their placement ensures that
cross-pollination 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, ...
only occurs between orchids of the same species. Different orchid bee males are attracted to different chemicals, so there is also some specificity regarding which orchid bees visit which types of orchid. The early description of this pollination system was by
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 ...
, though at the time, he believed the bees were females.Darwin & Appleton 1877 Not all orchids utilize euglossines as pollen vectors, of course; among the other types of insects exploited are other types of bees, wasps, flies, ants, and moths. The male of '' Eufriesea purpurata'' is highly unusual among insects in seeking out and collecting large quantities of insecticide. Dressler (1967) discovered ''E. purpurata'' collecting
aldrin Aldrin is an organochlorine insecticide that was widely used until the 1990s, when it was banned in most countries. Aldrin is a member of the so-called "classic organochlorines" (COC) group of pesticides. COCs enjoyed a very sharp rise in popular ...
and Roberts (1982) observed them collecting
DDT Dichlorodiphenyltrichloroethane, commonly known as DDT, is a colorless, tasteless, and almost odorless crystalline chemical compound, an organochloride. Originally developed as an insecticide, it became infamous for its environmental impacts. ...
Insect Behavior Mathews and Mathews 2010, p. 352 in huge amounts from houses in
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
, amounting to several percent of the bee's weight, without suffering any harm from the activity.


Footnotes


References

* (1877)
''The Various Contrivances by which Orchids are Fertilized by Insects''
* (1983): Orchid floral fragrances and male euglossine bees: methods and advances in the last sesquidecade. ''Biol. Bull.'' 164: 355–395. * Engel, Michael S. (1999): The first fossil ''Euglossa'' and phylogeny of the orchid bees (Hymenoptera: Apidae; Euglossini). ''American Museum Novitates'' 3272: 1–14
PDF
* (2004): Abejas De Orquídeas De La América Tropical: Biología y Guía De Campo / Orchid Bees of Tropical America: Biology and Field Guide. Santo Domingo, Costa Rica: INBio. In Spanish and English. . * (2004) Odor Compound Detection in Male Euglossine Bees. ''Journal of Chemical Ecology'' 29: 253–257. * (2005): Juggling with volatiles: fragrance exposure by displaying male orchid bees. ''Journal of Comparative Physiology A'' 191:575–581. * (2006): Species-specific attraction to pheromonal analogues in orchid bees. ''Behavioral Ecology and Sociobiology'' 60: 833–843.


External links



.
Abstract about ''Euglossa paisa'', ''Zootaxa'' 1065: 51–60 (2005)

Video showing Euglossini Orchid Bees collecting fragrance from ''Mormodes badia'', a Mexican Orchid


on the UF / IFAS Featured Creatures website. {{Taxonbar, from=Q1521051 Apinae Bee tribes Insects of Central America Hymenoptera of North America Hymenoptera of South America Orchid pollinators