Fairyfly
Encyclopedia
Mymaridae, commonly known as fairyflies or fairy wasps, is a family
of chalcid
wasp
s found in temperate
and tropical regions throughout the world. It contains around 100 genera and 1424 species. All of them are parasitoids of the eggs of other insect
s. Several species have been successfully used as biological pest control
agents.
Like most chalcid wasps, fairyflies are very tiny insects, averaging at only 0.5 to 1 mm (0.0196850393700787 to 0.0393700787401575 in) long. They include the world's smallest known insect
, with a body length of only 0.139 mm (0.00547244094488189 in).
They can be distinguished from other chalcid wasp
s by having an H-shaped pattern of sutures, known as trabeculae or carinae, below the frontmost ocelli and the inner eye margins. Rarely, the sutures can also extend behind the ocelli.
Fairyflies have long antennae
, at least as long as the head and the mesosoma
(middle part of the body). The antennal toruli (sockets of the antennae) are set high on the head and near the eye margins. They are separated by a distance of three to five times their own diameter. In contrast, the antennae of other chalcid wasps are separated only by one diameter. In females, the antennae are tipped with club-like segments known as clava. In males, the antennae are filiform (thread-like).
Though some fairyflies are wingless or possess only short wings, most fairyflies possess wings with long bristles (marginal setae) on the fringes. The forewings usually have hypochaeta. These are small bristles (setae) which point distinctly backwards on the ventral surface of the wing membrane. The hind wings are stalked (petiolate) and very narrow.
Because of their small sizes, fairyflies may sometimes be mistaken for members of the families Aphelinidae
and Trichogrammatidae
, but members of these other families can readily be distinguished by having much shorter antennae.
Fairyflies are found in temperate
and tropical regions throughout the world. The largest number of species can be found in tropical forests, with the greatest diversity in genera found in the Southern Hemisphere
(South America
, New Zealand
, and Australia
). In the Nearctic
(North America
), only around 28 (of around 100) genera and 120 (of around 1424) species are found.
Fairyflies can survive in all kinds of terrestrial habitats, from desert
s to rainforest
s. At least five species of fairyflies are also known to be aquatic
, inhabiting freshwater
ponds and streams. Among these is Caraphractus cinctus, which use their wings as paddles to swim around. They can remain underwater for as long as 15 days. Because of their small sizes, however, they have to exit the water by climbing up plant stems that jut through the surface, as they would not be able to break the surface tension
of the water otherwise.
s of egg
s of other insects. These are commonly eggs which are laid in concealed locations, like in plant tissues or underground. They do not seem to be species-specific when it comes to choosing hosts
. Some species are known to parasitize insects from several families of a single order
. Their most common hosts are insects belonging to the order Hemiptera
(true bugs), especially Auchenorrhyncha
(leafhopper
s, cicada
s, and allies) and Coccoidea (scale insects), but this might be because these groups are simply better studied. Other important host orders include Coleoptera (beetles), Diptera
(true flies), Odonata
(dragonflies
and allies), Psocoptera
(booklice and allies), and Thysanoptera (thrips). Hosts are known definitely for only a quarter of all known genera.
Fairyflies include the smallest known insect, Dicopomorpha echmepterygis
from Costa Rica
, which have males that are only 0.139 mm (0.00547244094488189 in) long. They do not have wings or eyes, their mouths are mere holes, and their antennae are simply spherical blobs. The ends of their legs have been modified into suction cups for clutching at females long enough to fertilize them. They are so small, their entire bodies are smaller than a single-celled Paramecium
. Four male individuals, lined up end-to-end, would just about encompass the width of a period
at the end of a typical printed sentence. The females of the species, however, are typical fairyflies, and are much larger.
Mating occurs immediately after emergence. No courting behaviors have been observed and each female is fertilized by only one male. Once fertilized, the males lose interest in the female and the female also loses interest in further mating. In Prestwichia aquatica, mating has been reported to occur prior to the emergence of females from the host eggs. In P. aquatica, Anagres incarnatus, A. armatus, and Anaphes nipponicus, females outnumber males in each brood; from two females for each male to twenty females for each male. In Polynema striaticorne, it is the opposite, with males outnumbering females by a ratio of three to one or more. In Anagrus atomus, A. frequens, A. optabilis, A. perforator, and Polynema euchariformes, females are able to reproduce without males (parthenogenesis
), though males may still sometimes be found within broods.
Adult lifespans of fairyflies are very short. In Stethynium, adults (males and females) can live for only 1 to 2 days. In Anagrus, depending on the species, it can range from 3 to 11 days. Each fertilized (or parthenogenic) female can lay a maximum of only about 100 eggs. Access to food can prolong lifespans and increase fecundity. In Gonatocerus, it has been shown that with the absence of hosts, females, can resorb eggs, retaining energy in order to live longer and increase the chances of finding hosts.
After emerging, females search rapidly for suitable host eggs by tapping their antennae over stems or barks of plants. When a telltale scar left by egg-laying insects is found, they will insert their antennae into the recess and check to see if the eggs are suitable. If they are, she will thrust her ovipositor
into all of the eggs and lay her own eggs inside in quick succession. She retains contact with each of the eggs with her antennae while doing this.
Most fairyflies require a sufficient amount of development in embryo
s inside the eggs before they attack it, as they can't mature if the eggs are too new or if the embryos inside are too advanced. Older host embryos are apparently harder to digest for the fairyfly larvae. However, there are exceptions. Some species of Polynema can attack embryos at various stages of development. They have been recorded to produce three successive generations in a single brood of the treehopper
Ceresa
.
All fairyflies possess ellipsoid eggs with a long tapering stalk. They develop rapidly once laid and can hatch in six hours to two days. Several generations may be produced in a year, often on different hosts.
Fairyflies have two to four larval stages, all apparently without functional spiracle
s or tracheae
. Metamorphosis
occurs completely within the host egg. They are peculiar for insects which exhibit complete metamorphosis (holometabolism) in that they produce two distinct kinds of larva
l instar
s before pupation. In some fairyflies like Anaphes, the first instar is a highly mobile "mymmariform" larva. The second instar, however, is a completely immobile sac-like larva without discernible segments, spines, or setae. In other fairyflies like Anagrus, this is reversed. The first instar is immobile, while the second instar is a very distinctive highly active "hystriobdellid" larva. At least one instar of the larvae is capable of overwintering
when laid in colder seasons.
entomologist Alexander Henry Haliday
. Haliday and two close friends and respected entomologists in their own right, John Curtis
and Francis Walker
, were influential in the early studies on Hymenoptera
in the 19th century.
Haliday originally described fairyflies as the tribe "Mymares" of the family "Chalcides". He based his descriptions on the type genus
Mymar, described by Curtis in 1829.
Earlier attempts of classification by Walker treated the group as a genus, and classified all other known fairyflies under it as subgenera. Walker (who was infamous for his shortcomings in systematic nomenclature
) later conceded to Haliday's classification in a letter in 1839 and requested assistance from Haliday in classifying the chalcid wasps collected by Charles Darwin
on his voyage
on the HMS Beagle
. "Mymares", as well as other "tribes" were elevated to the family rank
(as Mymaridae) by Haliday in 1839.
Haliday described fairyflies as "the very atoms of the order Hymenoptera" and remarked on the beauty of their wings when viewed under the microscope. These characteristics of fairyflies also made them popular to entomologists and microscopists of the late 19th century and the early 20th century. The beautifully mounted fairyfly specimens of the early 20th century English
microscopist Fred Enock is possibly the most famous of the collections.
(Chalcidoidea). There are approximately 1424 species of fairyflies grouped in about 100 genera. The largest genera are Anagrus, Anaphes, Gonatocerus, and Polynema, which comprise approximately half of all known species. They are the most commonly encountered fairyflies, followed by Alaptus, Camptoptera, Erythmelus, Ooctonus, and Stethynium, which make up a further quarter of known species. Mymaridae are considered to be monophyletic, but their exact relationships with other chalcid wasps remain unclear.
No commonly accepted subfamilies have been acknowledged, but there are two notable proposals. Annecke and Doutte (1961) proposed the subfamilies Alaptinae and Mymarinae based on the morphology of the gaster
. Peck et al. (1964) proposed the subfamilies Gonatocerinae and Mymarinae based on the number of segments (tarsomeres) in the tarsi. Both systems included further tribal categories. A fossil subfamily was also proposed for a genus recovered from Canadian amber.
Of the extant fairyfly genera, the genus Eustochomorpha is the most primitive. For the list of subordinate taxa, see the section below.
The generic name Mymar (and by extension, Mymaridae) was derived by Curtis from the Greek word μΰμαρ (mumar), an Aeolian
variant of the standard Greek μῶμος (mōmos, "spot" or "blot", also "blame", "reproach", or "disgrace").
of various crop pests. They are especially valued for their ability to locate the eggs of their hosts. Of the fairyflies, the cosmopolitan genus Anagrus is the most promising, as it can parasitize a wide range of hosts. There are at least ten known instances of successful introductions of fairyflies to control pests.
In Hawaii
, Anagrus optabilis has been successfully used to control the sugarcane planthopper (Perkinsiella saccharicida). In South Africa
, Anaphes nitens was introduced very successfully to control the eucalyptus snout beetle (Gonipterus scutellatus
).
Anagrus epos has also previously been used effectively against the grape leafhopper (Erythroneura elegans) and the variegated grape leafhopper (Erythroneura variabilis). At first it met limited success as the wasps died in winter for lack of a host. The planting of wild blackberries
(Rubus
spp.) near vineyards in California brought along with it the blackberry leafhopper (Dikrella cruentata). Though not pests, D. cruentata proved to be sufficient hosts for A. epos in winter, allowing them to survive into the next year. They have also been shown capable of overwintering in prune leafhoppers (Edwardsiana prunicola).
Gonatocerus triguttatus, Gonatocerus tuberculifemur, and Anagrus epos (which is probably a species complex
) were also studied for possible use in controlling glassy-winged sharpshooter
s (Homalodisca vitripennis) in California
. Gonatocerus triguttatus and Gonatocerus ashmeadi were eventually introduced in California in 2000. It proved to be very successful, causing a ~90% decline in populations of glassy-winged sharpshooters.
In 2005, Gonatocerus ashmeadi was used against invasive glassy-winged sharpshooters in Tahiti
and Moorea
of the French Polynesia
. Again, it was very effective, bringing down the pest density by ~95% in just a year after introduction. It was subsequently also released in the Society Islands
, the Marquesas, and Austral
, where similar results were obtained against glassy-winged sharpshooters.
In Australia
and Israel
, members of the genus Stethynium are being investigated as possible biological control agents for gall-forming crop pests, particularly Ophelimus maskelli.
amber inclusions, copal
, and compression fossil
s. Fossils of fairyflies have been found from the Early Cretaceous
up to the Miocene
epoch
. It is, in fact, the only family of chalcid wasps definitely known to date back to the Cretaceous
period.
Paleogene
and Neogene
fossils of fairyflies were first described in 1901 by Fernand Anatole Meunier. He described fossil fairyflies from Baltic amber
, most of them from the Eocene
(55 to 37 mya). In 1973, Richard L. Doutt described several species from the Burdigalian
(20 to 15 mya) amber of Mexico
. In 1983, Csaba Thuróczy described another species from Baltic amber, this time dating to the Oligocene
(33 to 23 mya). And in 2011, John T. Huber and Dale Greenwalt described fairyfly fossils from the oil shale
s of the Kishenehn Formation (Lutetian
age) of Montana
. These comprised two new genera and six species.
Cretaceous fairyflies are much rarer. In 1975, Carl M. Yoshimoto described four genera of fairyflies from the Cretaceous of Canada
. In 2011, John T. Huber and George Poinar, Jr.
described the genus Myanmymar
from Burmese amber. Dating back to the Upper Albian
age (about 100 mya) of the Early Cretaceous
, it is the oldest known fairyfly (and chalcid wasp). They are surprisingly very similar to modern genera, though with a greater number of flagellar segments and longer forewing veins. The characteristics of the fossil (taking into account Yoshimoto's earlier discoveries) led them to conclude that fairfylies either may have existed much earlier than Myanmymar, or that they may have diversified rapidly during that time period.
listed below. Allomymar
and Metanthemus has been transferred to the family Aphelinidae
. The fossil genus Protooctonus has been transferred to the family Mymarommatidae
, and is now considered to be a synonym
of Archaeromma. Shillingsworthia is also excluded, as it was a tongue-in-cheek hypothetical concept of a species from the planet Jupiter
, "described" by Alexandre Arsène Girault
in 1920 to disparage his colleague Johann Francis Illingworth.
Genera marked with † are extinct.
genera classified under Mymaridae:
Despite their relative abundance, fairyflies are unpopular among modern insect collectors because of the great difficulty in collecting them. As one of the least known insect families, there is still a large amount of information that are waiting to be discovered for fairyflies. This is an area of entomology where an amateur naturalist can still make significant contributions.
In the 19th century, Curtis described the methods by which Haliday collected fairyflies as thus:
The best modern collection method is using Malaise trap
s. It requires little maintenance and can collect insects in great abundance. Other effective methods include yellow pan trapping, sweep nets, and suction trapping. Direct collection from leaf litter with Berlese funnels can also result in specimens that can not be collected by other means.
Rearing is also another method that can bring the most rewards. This can be done with wild host eggs or laboratory prepared host eggs that are exposed in suitable habitats outside. They can then be taken in after a suitable amount of time has passed and examined for developing fairyflies. With this method, it is possible to observe the life history and determine the hosts of particular species of discovered fairyflies.
Preservation is a problem for fairyflies (and other small chalcid wasps). Their tiny sizes require special methods. Specimens have to be dried, if collected wet, e.g. if ethyl alcohol is used as the killing agent
. Drying can make specimens extremely brittle, so additional care should also be taken not to disintegrate them. Mounting specimens (preferably in permanent slides) is also time-consuming and requires a fair amount of practice. They are gummed or glued into cards, as they can not be mounted on pins like larger insects.
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
of chalcid
Chalcid wasp
Chalcid wasps belong to the insect order Hymenoptera, and are one of the largest groups within the order, with some 22,000 known species, and an estimated total diversity of anywhere from 60,000 to more than 500,000 species, meaning the vast majority have yet to be discovered and described.Most of...
wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...
s found in temperate
Temperate
In geography, temperate or tepid latitudes of the globe lie between the tropics and the polar circles. The changes in these regions between summer and winter are generally relatively moderate, rather than extreme hot or cold...
and tropical regions throughout the world. It contains around 100 genera and 1424 species. All of them are parasitoids of the eggs of other insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s. Several species have been successfully used as biological pest control
Biological pest control
Biological control of pests in agriculture is a method of controlling pests that relies on predation, parasitism, herbivory, or other natural mechanisms...
agents.
Like most chalcid wasps, fairyflies are very tiny insects, averaging at only 0.5 to 1 mm (0.0196850393700787 to 0.0393700787401575 in) long. They include the world's smallest known insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
, with a body length of only 0.139 mm (0.00547244094488189 in).
Description
Fairyflies are very small insects. They have body lengths in between 0.13 to 5.4 mm (0.00511811023622047 to 0.21259842519685 in), averaging at 0.5 to 1 mm (0.0196850393700787 to 0.0393700787401575 in). Their bodies are usually nonmetallic and colored black, brown, or yellow.They can be distinguished from other chalcid wasp
Chalcid wasp
Chalcid wasps belong to the insect order Hymenoptera, and are one of the largest groups within the order, with some 22,000 known species, and an estimated total diversity of anywhere from 60,000 to more than 500,000 species, meaning the vast majority have yet to be discovered and described.Most of...
s by having an H-shaped pattern of sutures, known as trabeculae or carinae, below the frontmost ocelli and the inner eye margins. Rarely, the sutures can also extend behind the ocelli.
Fairyflies have long antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
, at least as long as the head and the mesosoma
Mesosoma
The mesosoma is the middle part of the body, or tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the metasoma. It bears the legs, and, in the case of winged insects, the wings....
(middle part of the body). The antennal toruli (sockets of the antennae) are set high on the head and near the eye margins. They are separated by a distance of three to five times their own diameter. In contrast, the antennae of other chalcid wasps are separated only by one diameter. In females, the antennae are tipped with club-like segments known as clava. In males, the antennae are filiform (thread-like).
Though some fairyflies are wingless or possess only short wings, most fairyflies possess wings with long bristles (marginal setae) on the fringes. The forewings usually have hypochaeta. These are small bristles (setae) which point distinctly backwards on the ventral surface of the wing membrane. The hind wings are stalked (petiolate) and very narrow.
Because of their small sizes, fairyflies may sometimes be mistaken for members of the families Aphelinidae
Aphelinidae
Aphelinidae is a moderate-sized family of tiny parasitic wasps, with some 1160 described species in some 35 genera. These minute insects are challenging to study as they deteriorate rapidly after death unless extreme care is taken , making identification of most museum specimens difficult...
and Trichogrammatidae
Trichogrammatidae
The family Trichogrammatidae are tiny wasps in the Chalcidoidea that include some of the smallest of all insects, with most species having adults less than 1 mm in length. There are over 840 species in ca. 80 genera worldwide. Trichogrammatids parasitize the eggs of many different orders of insects...
, but members of these other families can readily be distinguished by having much shorter antennae.
Distribution and habitat
Fairyflies are some of the most common chalcid wasp, but because of their minute sizes, they are seldom noticed by humans. This apparent invisibility, their delicate bodies, and their hair-fringed wings have earned them their common name 'fairyfly'.Fairyflies are found in temperate
Temperate
In geography, temperate or tepid latitudes of the globe lie between the tropics and the polar circles. The changes in these regions between summer and winter are generally relatively moderate, rather than extreme hot or cold...
and tropical regions throughout the world. The largest number of species can be found in tropical forests, with the greatest diversity in genera found in the Southern Hemisphere
Southern Hemisphere
The Southern Hemisphere is the part of Earth that lies south of the equator. The word hemisphere literally means 'half ball' or "half sphere"...
(South America
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...
, New Zealand
New Zealand
New Zealand is an island country in the south-western Pacific Ocean comprising two main landmasses and numerous smaller islands. The country is situated some east of Australia across the Tasman Sea, and roughly south of the Pacific island nations of New Caledonia, Fiji, and Tonga...
, and Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...
). In the Nearctic
Nearctic
The Nearctic is one of the eight terrestrial ecozones dividing the Earth's land surface.The Nearctic ecozone covers most of North America, including Greenland and the highlands of Mexico...
(North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...
), only around 28 (of around 100) genera and 120 (of around 1424) species are found.
Fairyflies can survive in all kinds of terrestrial habitats, from desert
Desert
A desert is a landscape or region that receives an extremely low amount of precipitation, less than enough to support growth of most plants. Most deserts have an average annual precipitation of less than...
s to rainforest
Rainforest
Rainforests are forests characterized by high rainfall, with definitions based on a minimum normal annual rainfall of 1750-2000 mm...
s. At least five species of fairyflies are also known to be aquatic
Aquatic animal
An aquatic animal is an animal, either vertebrate or invertebrate, which lives in water for most or all of its life. It may breathe air or extract its oxygen from that dissolved in water through specialised organs called gills, or directly through its skin. Natural environments and the animals that...
, inhabiting freshwater
Freshwater
Fresh water is naturally occurring water on the Earth's surface in ice sheets, ice caps, glaciers, bogs, ponds, lakes, rivers and streams, and underground as groundwater in aquifers and underground streams. Fresh water is generally characterized by having low concentrations of dissolved salts and...
ponds and streams. Among these is Caraphractus cinctus, which use their wings as paddles to swim around. They can remain underwater for as long as 15 days. Because of their small sizes, however, they have to exit the water by climbing up plant stems that jut through the surface, as they would not be able to break the surface tension
Surface tension
Surface tension is a property of the surface of a liquid that allows it to resist an external force. It is revealed, for example, in floating of some objects on the surface of water, even though they are denser than water, and in the ability of some insects to run on the water surface...
of the water otherwise.
Ecology
All known fairyflies are parasitoidParasitoid
A parasitoid is an organism that spends a significant portion of its life history attached to or within a single host organism in a relationship that is in essence parasitic; unlike a true parasite, however, it ultimately sterilises or kills, and sometimes consumes, the host...
s of egg
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...
s of other insects. These are commonly eggs which are laid in concealed locations, like in plant tissues or underground. They do not seem to be species-specific when it comes to choosing hosts
Host (biology)
In biology, a host is an organism that harbors a parasite, or a mutual or commensal symbiont, typically providing nourishment and shelter. In botany, a host plant is one that supplies food resources and substrate for certain insects or other fauna...
. Some species are known to parasitize insects from several families of a single order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...
. Their most common hosts are insects belonging to the order Hemiptera
Hemiptera
Hemiptera is an order of insects most often known as the true bugs , comprising around 50,000–80,000 species of cicadas, aphids, planthoppers, leafhoppers, shield bugs, and others...
(true bugs), especially Auchenorrhyncha
Auchenorrhyncha
The Auchenorrhyncha is the suborder of the Hemiptera which contains most of the familiar members of what was called the Homoptera - groups such as cicadas, leafhoppers, treehoppers, planthoppers, and spittlebugs. The aphids and scale insects are the other well-known "Homoptera", and they are in...
(leafhopper
Leafhopper
Leafhopper is a common name applied to any species from the family Cicadellidae. Leafhoppers, colloquially known as hoppers, are minute plant-feeding insects in the superfamily Membracoidea in the order Hemiptera...
s, cicada
Cicada
A cicada is an insect of the order Hemiptera, suborder Auchenorrhyncha , in the superfamily Cicadoidea, with large eyes wide apart on the head and usually transparent, well-veined wings. There are about 2,500 species of cicada around the world, and many of them remain unclassified...
s, and allies) and Coccoidea (scale insects), but this might be because these groups are simply better studied. Other important host orders include Coleoptera (beetles), Diptera
Diptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...
(true flies), Odonata
Odonata
Odonata is an order of insects, encompassing dragonflies and damselflies . The word dragonfly is also sometimes used to refer to all Odonata, but the back-formation odonate is a more correct English name for the group as a whole...
(dragonflies
Dragonfly
A dragonfly is a winged insect belonging to the order Odonata, the suborder Epiprocta or, in the strict sense, the infraorder Anisoptera . It is characterized by large multifaceted eyes, two pairs of strong transparent wings, and an elongated body...
and allies), Psocoptera
Psocoptera
Psocoptera are an order of insects that are commonly known as booklice, barklice or barkflies. They first appeared in the Permian period, 295–248 million years ago. They are often regarded as the most primitive of the hemipteroids. Their name originates from the Greek word psokos meaning...
(booklice and allies), and Thysanoptera (thrips). Hosts are known definitely for only a quarter of all known genera.
Fairyflies include the smallest known insect, Dicopomorpha echmepterygis
Dicopomorpha echmepterygis
Dicopomorpha echmepterygis is a parasitic wasp in the family Mymaridae. The males of this species are the smallest of all known insects. They are blind and wingless and may be no more than 0.139 mm in length . Females are 40% larger...
from Costa Rica
Costa Rica
Costa Rica , officially the Republic of Costa Rica is a multilingual, multiethnic and multicultural country in Central America, bordered by Nicaragua to the north, Panama to the southeast, the Pacific Ocean to the west and the Caribbean Sea to the east....
, which have males that are only 0.139 mm (0.00547244094488189 in) long. They do not have wings or eyes, their mouths are mere holes, and their antennae are simply spherical blobs. The ends of their legs have been modified into suction cups for clutching at females long enough to fertilize them. They are so small, their entire bodies are smaller than a single-celled Paramecium
Paramecium
Paramecium is a group of unicellular ciliate protozoa, which are commonly studied as a representative of the ciliate group, and range from about 0.05 to 0.35 mm in length. Simple cilia cover the body, which allow the cell to move with a synchronous motion at speeds of approximately 12 body...
. Four male individuals, lined up end-to-end, would just about encompass the width of a period
Full stop
A full stop is the punctuation mark commonly placed at the end of sentences. In American English, the term used for this punctuation is period. In the 21st century, it is often also called a dot by young people...
at the end of a typical printed sentence. The females of the species, however, are typical fairyflies, and are much larger.
Life cycle
Very little is known of the life histories of fairyflies as only a few species have been observed extensively. They are usually solitary, but can sometimes be gregarious.Mating occurs immediately after emergence. No courting behaviors have been observed and each female is fertilized by only one male. Once fertilized, the males lose interest in the female and the female also loses interest in further mating. In Prestwichia aquatica, mating has been reported to occur prior to the emergence of females from the host eggs. In P. aquatica, Anagres incarnatus, A. armatus, and Anaphes nipponicus, females outnumber males in each brood; from two females for each male to twenty females for each male. In Polynema striaticorne, it is the opposite, with males outnumbering females by a ratio of three to one or more. In Anagrus atomus, A. frequens, A. optabilis, A. perforator, and Polynema euchariformes, females are able to reproduce without males (parthenogenesis
Parthenogenesis
Parthenogenesis is a form of asexual reproduction found in females, where growth and development of embryos occur without fertilization by a male...
), though males may still sometimes be found within broods.
Adult lifespans of fairyflies are very short. In Stethynium, adults (males and females) can live for only 1 to 2 days. In Anagrus, depending on the species, it can range from 3 to 11 days. Each fertilized (or parthenogenic) female can lay a maximum of only about 100 eggs. Access to food can prolong lifespans and increase fecundity. In Gonatocerus, it has been shown that with the absence of hosts, females, can resorb eggs, retaining energy in order to live longer and increase the chances of finding hosts.
After emerging, females search rapidly for suitable host eggs by tapping their antennae over stems or barks of plants. When a telltale scar left by egg-laying insects is found, they will insert their antennae into the recess and check to see if the eggs are suitable. If they are, she will thrust her ovipositor
Ovipositor
The ovipositor is an organ used by some animals for oviposition, i.e., the laying of eggs. It consists of a maximum of three pairs of appendages formed to transmit the egg, to prepare a place for it, and to place it properly...
into all of the eggs and lay her own eggs inside in quick succession. She retains contact with each of the eggs with her antennae while doing this.
Most fairyflies require a sufficient amount of development in embryo
Embryo
An embryo is a multicellular diploid eukaryote in its earliest stage of development, from the time of first cell division until birth, hatching, or germination...
s inside the eggs before they attack it, as they can't mature if the eggs are too new or if the embryos inside are too advanced. Older host embryos are apparently harder to digest for the fairyfly larvae. However, there are exceptions. Some species of Polynema can attack embryos at various stages of development. They have been recorded to produce three successive generations in a single brood of the treehopper
Treehopper
Treehoppers and thorn bugs are members of the family Membracidae, a group of insects related to the cicadas and the leafhoppers. There are about 3,200 known species of treehoppers in over 600 genera...
Ceresa
Ceresa
Ceresa, commonly known as buffalo treehoppers, is a genus of treehoppers. It contains about 16 species.*Ceresa albescens*Ceresa albidosparsa*Ceresa alta*Ceresa ancora*Ceresa basalis*Ceresa borealis*Ceresa diceros...
.
All fairyflies possess ellipsoid eggs with a long tapering stalk. They develop rapidly once laid and can hatch in six hours to two days. Several generations may be produced in a year, often on different hosts.
Fairyflies have two to four larval stages, all apparently without functional spiracle
Spiracle
Spiracles are openings on the surface of some animals that usually lead to respiratory systems.-Vertebrates:The spiracle is a small hole behind each eye that opens to the mouth in some fishes. In the primitive jawless fish the first gill opening immediately behind the mouth is essentially similar...
s or tracheae
Invertebrate trachea
The invertebrate trachea refers to the open respiratory system composed of spiracles, tracheae, and tracheoles that terrestrial arthropods have to transport metabolic gases to and from tissues....
. Metamorphosis
Metamorphosis
Metamorphosis is a biological process by which an animal physically develops after birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation...
occurs completely within the host egg. They are peculiar for insects which exhibit complete metamorphosis (holometabolism) in that they produce two distinct kinds of larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
l instar
Instar
An instar is a developmental stage of arthropods, such as insects, between each molt , until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, or...
s before pupation. In some fairyflies like Anaphes, the first instar is a highly mobile "mymmariform" larva. The second instar, however, is a completely immobile sac-like larva without discernible segments, spines, or setae. In other fairyflies like Anagrus, this is reversed. The first instar is immobile, while the second instar is a very distinctive highly active "hystriobdellid" larva. At least one instar of the larvae is capable of overwintering
Hibernation
Hibernation is a state of inactivity and metabolic depression in animals, characterized by lower body temperature, slower breathing, and lower metabolic rate. Hibernating animals conserve food, especially during winter when food supplies are limited, tapping energy reserves, body fat, at a slow rate...
when laid in colder seasons.
History
The family Mymaridae was first established in 1833 by the IrishIrish people
The Irish people are an ethnic group who originate in Ireland, an island in northwestern Europe. Ireland has been populated for around 9,000 years , with the Irish people's earliest ancestors recorded having legends of being descended from groups such as the Nemedians, Fomorians, Fir Bolg, Tuatha...
entomologist Alexander Henry Haliday
Alexander Henry Haliday
Alexander Henry Haliday, also known as Enrico Alessandro Haliday and Alexis Heinrich Haliday sometimes Halliday , was an Irish entomologist. He is primarily known for his work on Hymenoptera, Diptera and Thysanoptera, but Haliday worked on all insect orders and on many aspects of entomology.Haliday...
. Haliday and two close friends and respected entomologists in their own right, John Curtis
John Curtis (entomologist)
John Curtis was an English entomologist and illustrator.-Biography:Curtis was born in Norwich and learned his engraving skills in the workshop of his father, Charles Morgan Curtis...
and Francis Walker
Francis Walker (entomologist)
Francis Walker was an English entomologist. He was one of the most prolific authors in entomology, and stirred controversy during his later life as his publications resulted in a huge number of junior synonyms....
, were influential in the early studies on Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
in the 19th century.
Haliday originally described fairyflies as the tribe "Mymares" of the family "Chalcides". He based his descriptions on the type genus
Type genus
In biological classification, a type genus is a representative genus, as with regard to a biological family. The term and concept is used much more often and much more formally in zoology than it is in botany, and the definition is dependent on the nomenclatural Code that applies:* In zoological...
Mymar, described by Curtis in 1829.
Earlier attempts of classification by Walker treated the group as a genus, and classified all other known fairyflies under it as subgenera. Walker (who was infamous for his shortcomings in systematic nomenclature
Binomial nomenclature
Binomial nomenclature is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, although they can be based on words from other languages...
) later conceded to Haliday's classification in a letter in 1839 and requested assistance from Haliday in classifying the chalcid wasps collected by Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...
on his voyage
Second voyage of HMS Beagle
The second voyage of HMS Beagle, from 27 December 1831 to 2 October 1836, was the second survey expedition of HMS Beagle, under captain Robert FitzRoy who had taken over command of the ship on its first voyage after her previous captain committed suicide...
on the HMS Beagle
HMS Beagle
HMS Beagle was a Cherokee-class 10-gun brig-sloop of the Royal Navy. She was launched on 11 May 1820 from the Woolwich Dockyard on the River Thames, at a cost of £7,803. In July of that year she took part in a fleet review celebrating the coronation of King George IV of the United Kingdom in which...
. "Mymares", as well as other "tribes" were elevated to the family rank
Taxonomic rank
In biological classification, rank is the level in a taxonomic hierarchy. Examples of taxonomic ranks are species, genus, family, and class. Each rank subsumes under it a number of less general categories...
(as Mymaridae) by Haliday in 1839.
Haliday described fairyflies as "the very atoms of the order Hymenoptera" and remarked on the beauty of their wings when viewed under the microscope. These characteristics of fairyflies also made them popular to entomologists and microscopists of the late 19th century and the early 20th century. The beautifully mounted fairyfly specimens of the early 20th century English
English people
The English are a nation and ethnic group native to England, who speak English. The English identity is of early mediaeval origin, when they were known in Old English as the Anglecynn. England is now a country of the United Kingdom, and the majority of English people in England are British Citizens...
microscopist Fred Enock is possibly the most famous of the collections.
Taxonomy and etymology
The family Mymaridae is the most primitive member of the chalcid wasp superfamilySuperfamily
The term superfamily is used to describe several different concepts in different scientific fields:* Superfamily and Superfamily : a level of biological classification; redirects to Taxonomic rank...
(Chalcidoidea). There are approximately 1424 species of fairyflies grouped in about 100 genera. The largest genera are Anagrus, Anaphes, Gonatocerus, and Polynema, which comprise approximately half of all known species. They are the most commonly encountered fairyflies, followed by Alaptus, Camptoptera, Erythmelus, Ooctonus, and Stethynium, which make up a further quarter of known species. Mymaridae are considered to be monophyletic, but their exact relationships with other chalcid wasps remain unclear.
No commonly accepted subfamilies have been acknowledged, but there are two notable proposals. Annecke and Doutte (1961) proposed the subfamilies Alaptinae and Mymarinae based on the morphology of the gaster
Gaster
The gaster is the bulbous posterior portion of the metasoma found in Apocrita Hymenoptera . This begins with abdominal segment III on most ants, but some make a postpetiole out of segment III, in which case the gaster begins with abdominal segment IV....
. Peck et al. (1964) proposed the subfamilies Gonatocerinae and Mymarinae based on the number of segments (tarsomeres) in the tarsi. Both systems included further tribal categories. A fossil subfamily was also proposed for a genus recovered from Canadian amber.
Of the extant fairyfly genera, the genus Eustochomorpha is the most primitive. For the list of subordinate taxa, see the section below.
The generic name Mymar (and by extension, Mymaridae) was derived by Curtis from the Greek word μΰμαρ (mumar), an Aeolian
Aeolians
The Aeolians were one of the four major ancient Greek tribes comprising Ancient Greeks. Their name derives from Aeolus, the mythical ancestor of the Aeolic branch and son of Hellen, the mythical patriarch of the Greek nation...
variant of the standard Greek μῶμος (mōmos, "spot" or "blot", also "blame", "reproach", or "disgrace").
Economic importance
Fairyflies have been used for the biological pest controlBiological pest control
Biological control of pests in agriculture is a method of controlling pests that relies on predation, parasitism, herbivory, or other natural mechanisms...
of various crop pests. They are especially valued for their ability to locate the eggs of their hosts. Of the fairyflies, the cosmopolitan genus Anagrus is the most promising, as it can parasitize a wide range of hosts. There are at least ten known instances of successful introductions of fairyflies to control pests.
In Hawaii
Hawaii
Hawaii is the newest of the 50 U.S. states , and is the only U.S. state made up entirely of islands. It is the northernmost island group in Polynesia, occupying most of an archipelago in the central Pacific Ocean, southwest of the continental United States, southeast of Japan, and northeast of...
, Anagrus optabilis has been successfully used to control the sugarcane planthopper (Perkinsiella saccharicida). In South Africa
South Africa
The Republic of South Africa is a country in southern Africa. Located at the southern tip of Africa, it is divided into nine provinces, with of coastline on the Atlantic and Indian oceans...
, Anaphes nitens was introduced very successfully to control the eucalyptus snout beetle (Gonipterus scutellatus
Gonipterus scutellatus
Gonipterus scutellatus is a species of weevil in the Curculionidae family. It is commonly known as the eucalyptus snout beetle, the eucalyptus weevil or the gum tree weevil. It feeds and breeds on Eucalyptus trees and is endemic to Australia.-Description:This weevil is greyish-brown with a light...
).
Anagrus epos has also previously been used effectively against the grape leafhopper (Erythroneura elegans) and the variegated grape leafhopper (Erythroneura variabilis). At first it met limited success as the wasps died in winter for lack of a host. The planting of wild blackberries
Rubus
Rubus is a large genus of flowering plants in the rose family, Rosaceae, subfamily Rosoideae. Raspberries, blackberries, and dewberries are common, widely distributed members of the genus. Most of these plants have woody stems with prickles like roses; spines, bristles, and gland-tipped hairs are...
(Rubus
Rubus
Rubus is a large genus of flowering plants in the rose family, Rosaceae, subfamily Rosoideae. Raspberries, blackberries, and dewberries are common, widely distributed members of the genus. Most of these plants have woody stems with prickles like roses; spines, bristles, and gland-tipped hairs are...
spp.) near vineyards in California brought along with it the blackberry leafhopper (Dikrella cruentata). Though not pests, D. cruentata proved to be sufficient hosts for A. epos in winter, allowing them to survive into the next year. They have also been shown capable of overwintering in prune leafhoppers (Edwardsiana prunicola).
Gonatocerus triguttatus, Gonatocerus tuberculifemur, and Anagrus epos (which is probably a species complex
Species complex
A species complex is a group of closely related species, where the exact demarcation between species is often unclear or cryptic owing to their recent and usually still incomplete reproductive isolation. Ring species, superspecies and cryptic species complex are example of species complex...
) were also studied for possible use in controlling glassy-winged sharpshooter
Glassy-winged sharpshooter
The glassy-winged sharpshooter is a large leafhopper insect from the family Cicadellidae, similar to other species of sharpshooter.-Description:These sharpshooters are about in length...
s (Homalodisca vitripennis) in California
California
California is a state located on the West Coast of the United States. It is by far the most populous U.S. state, and the third-largest by land area...
. Gonatocerus triguttatus and Gonatocerus ashmeadi were eventually introduced in California in 2000. It proved to be very successful, causing a ~90% decline in populations of glassy-winged sharpshooters.
In 2005, Gonatocerus ashmeadi was used against invasive glassy-winged sharpshooters in Tahiti
Tahiti
Tahiti is the largest island in the Windward group of French Polynesia, located in the archipelago of the Society Islands in the southern Pacific Ocean. It is the economic, cultural and political centre of French Polynesia. The island was formed from volcanic activity and is high and mountainous...
and Moorea
Moorea
Moʻorea is a high island in French Polynesia, part of the Society Islands, 17 km northwest of Tahiti. Its position is . Moʻorea means "yellow lizard" in Tahitian...
of the French Polynesia
French Polynesia
French Polynesia is an overseas country of the French Republic . It is made up of several groups of Polynesian islands, the most famous island being Tahiti in the Society Islands group, which is also the most populous island and the seat of the capital of the territory...
. Again, it was very effective, bringing down the pest density by ~95% in just a year after introduction. It was subsequently also released in the Society Islands
Society Islands
The Society Islands are a group of islands in the South Pacific Ocean. They are politically part of French Polynesia. The archipelago is generally believed to have been named by Captain James Cook in honor of the Royal Society, the sponsor of the first British scientific survey of the islands;...
, the Marquesas, and Austral
Austral
Austral may refer to:*Austral Islands, the southernmost group of islands in French Polynesia*Austral language, the language of the Austral Islands*Austral, New South Wales, a suburb of Sydney, Australia*Argentine austral, a former currency of Argentina...
, where similar results were obtained against glassy-winged sharpshooters.
In Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...
and Israel
Israel
The State of Israel is a parliamentary republic located in the Middle East, along the eastern shore of the Mediterranean Sea...
, members of the genus Stethynium are being investigated as possible biological control agents for gall-forming crop pests, particularly Ophelimus maskelli.
Fossil record
Fairyflies are well-represented in fossilFossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...
amber inclusions, copal
Copal
Copal is a name given to tree resin that is particularly identified with the aromatic resins used by the cultures of pre-Columbian Mesoamerica as ceremonially burned incense and other purposes...
, and compression fossil
Compression fossil
A compression fossil is a fossil preserved in sedimentary rock that has undergone physical compression. While it is uncommon to find animals preserved as good compression fossils, it is very common to find plants preserved this way...
s. Fossils of fairyflies have been found from the Early Cretaceous
Early Cretaceous
The Early Cretaceous or the Lower Cretaceous , is the earlier or lower of the two major divisions of the Cretaceous...
up to the Miocene
Miocene
The Miocene is a geological epoch of the Neogene Period and extends from about . The Miocene was named by Sir Charles Lyell. Its name comes from the Greek words and and means "less recent" because it has 18% fewer modern sea invertebrates than the Pliocene. The Miocene follows the Oligocene...
epoch
Epoch (geology)
An epoch is a subdivision of the geologic timescale based on rock layering. In order, the higher subdivisions are periods, eras and eons. We are currently living in the Holocene epoch...
. It is, in fact, the only family of chalcid wasps definitely known to date back to the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...
period.
Paleogene
Paleogene
The Paleogene is a geologic period and system that began 65.5 ± 0.3 and ended 23.03 ± 0.05 million years ago and comprises the first part of the Cenozoic Era...
and Neogene
Neogene
The Neogene is a geologic period and system in the International Commission on Stratigraphy Geologic Timescale starting 23.03 ± 0.05 million years ago and ending 2.588 million years ago...
fossils of fairyflies were first described in 1901 by Fernand Anatole Meunier. He described fossil fairyflies from Baltic amber
Baltic amber
The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite, with about 80% of the world's known amber found there. It dates from 44 million years ago...
, most of them from the Eocene
Eocene
The Eocene Epoch, lasting from about 56 to 34 million years ago , is a major division of the geologic timescale and the second epoch of the Paleogene Period in the Cenozoic Era. The Eocene spans the time from the end of the Palaeocene Epoch to the beginning of the Oligocene Epoch. The start of the...
(55 to 37 mya). In 1973, Richard L. Doutt described several species from the Burdigalian
Burdigalian
The Burdigalian is, in the geologic timescale, an age or stage in the early Miocene. It spans the time between 20.43 ± 0.05 Ma and 15.97 ± 0.05 Ma...
(20 to 15 mya) amber of Mexico
Mexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...
. In 1983, Csaba Thuróczy described another species from Baltic amber, this time dating to the Oligocene
Oligocene
The Oligocene is a geologic epoch of the Paleogene Period and extends from about 34 million to 23 million years before the present . As with other older geologic periods, the rock beds that define the period are well identified but the exact dates of the start and end of the period are slightly...
(33 to 23 mya). And in 2011, John T. Huber and Dale Greenwalt described fairyfly fossils from the oil shale
Oil shale
Oil shale, an organic-rich fine-grained sedimentary rock, contains significant amounts of kerogen from which liquid hydrocarbons called shale oil can be produced...
s of the Kishenehn Formation (Lutetian
Lutetian
The Lutetian is, in the geologic timescale, a stage or age in the Eocene. It spans the time between and . The Lutetian is preceded by the Ypresian and is followed by the Bartonian. Together with the Bartonian it is sometimes referred to as the Middle Eocene subepoch...
age) of Montana
Montana
Montana is a state in the Western United States. The western third of Montana contains numerous mountain ranges. Smaller, "island ranges" are found in the central third of the state, for a total of 77 named ranges of the Rocky Mountains. This geographical fact is reflected in the state's name,...
. These comprised two new genera and six species.
Cretaceous fairyflies are much rarer. In 1975, Carl M. Yoshimoto described four genera of fairyflies from the Cretaceous of Canada
Canada
Canada is a North American country consisting of ten provinces and three territories. Located in the northern part of the continent, it extends from the Atlantic Ocean in the east to the Pacific Ocean in the west, and northward into the Arctic Ocean...
. In 2011, John T. Huber and George Poinar, Jr.
George Poinar, Jr.
George O. Poinar, Jr. is an entomologist and writer. He is known for popularizing the idea of extracting DNA from insects fossilized in amber, an idea which received widespread attention when adapted by Michael Crichton for the book and movie Jurassic Park.Poinar earned a B.S. and M.S. at Cornell...
described the genus Myanmymar
Myanmymar
Myanmymar is an extinct genus of fairyfly preserved in amber from Myanmar. It has only one species, Myanmymar aresconoides. It is dated to the Early Cretaceous, at least a hundred million years ago. As of 2011, it is the oldest known fossil mymarid.-Discovery and geologic time range:Myanmymar...
from Burmese amber. Dating back to the Upper Albian
Albian
The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous epoch/series. Its approximate time range is 112.0 ± 1.0 Ma to 99.6 ± 0.9 Ma...
age (about 100 mya) of the Early Cretaceous
Early Cretaceous
The Early Cretaceous or the Lower Cretaceous , is the earlier or lower of the two major divisions of the Cretaceous...
, it is the oldest known fairyfly (and chalcid wasp). They are surprisingly very similar to modern genera, though with a greater number of flagellar segments and longer forewing veins. The characteristics of the fossil (taking into account Yoshimoto's earlier discoveries) led them to conclude that fairfylies either may have existed much earlier than Myanmymar, or that they may have diversified rapidly during that time period.
Genera
Mymaridae includes the generaGenera
Genera is a commercial operating system and development environment for Lisp machines developed by Symbolics. It is essentially a fork of an earlier operating system originating on the MIT AI Lab's Lisp machines which Symbolics had used in common with LMI and Texas Instruments...
listed below. Allomymar
Allomymar
Allomymar is a little-known genus of chalcid wasps belonging to the family Aphelinidae. It is possible that Allomymar belongs in the genus Encarsia....
and Metanthemus has been transferred to the family Aphelinidae
Aphelinidae
Aphelinidae is a moderate-sized family of tiny parasitic wasps, with some 1160 described species in some 35 genera. These minute insects are challenging to study as they deteriorate rapidly after death unless extreme care is taken , making identification of most museum specimens difficult...
. The fossil genus Protooctonus has been transferred to the family Mymarommatidae
Mymarommatidae
Mymarommatidae is a very small family of microscopic hymenopteran insects. Only 10 living species in 1 genus have currently been described but they are known from all parts of the world...
, and is now considered to be a synonym
Synonym (taxonomy)
In scientific nomenclature, a synonym is a scientific name that is or was used for a taxon of organisms that also goes by a different scientific name. For example, Linnaeus was the first to give a scientific name to the Norway spruce, which he called Pinus abies...
of Archaeromma. Shillingsworthia is also excluded, as it was a tongue-in-cheek hypothetical concept of a species from the planet Jupiter
Jupiter
Jupiter is the fifth planet from the Sun and the largest planet within the Solar System. It is a gas giant with mass one-thousandth that of the Sun but is two and a half times the mass of all the other planets in our Solar System combined. Jupiter is classified as a gas giant along with Saturn,...
, "described" by Alexandre Arsène Girault
Alexandre Arsène Girault
Alexandre Arsène Girault was an American entomologist specialising in the study of chalcid wasps. An eccentric and controversial figure, Girault was also a prolific and dedicated entomologist...
in 1920 to disparage his colleague Johann Francis Illingworth.
Genera marked with † are extinct.
Extant genera
- Acanthomymar Subba Rao, 1970
- Acmopolynema Ogloblin,1946
- Acmotemnus Noyes & Valentine, 1989
- Agalmopolynema Ogloblin, 1960
- Alaptus Ferrière, 1930
- Allanagrus Noyes & Valentine, 1989
- Allarescon Noyes & Valentine, 1989
- Anagroidea Girault, 1915
- Anagrus Haliday, 1833
- Anaphes Haliday, 1833
- Anneckia Subba Rao, 1970
- Apoxypteron Noyes & Valentine, 1989
- AresconAresconArescon is a genus of fairyflies.. It contains the following species:* Arescon aspidioticola * Arescon clarkei Doutt, 1955* Arescon dallasi * Arescon dimidiata...
Walker, 1846 - Australomymar Girault, 1929
- Baburia Hedqvist, 2004
- Bakkendorfia Mathot, 1966
- Boccacciomymar Triapitysn & Berezovskiy, 2007
- Borneomymar Huber, 2002
- Boudiennyia Girault, 1937
- Bruchomymar Ogloblin, 1939
- Caenomymar Yoshimoto, 1990
- Callodicopus Ogloblin, 1955
- Camptoptera Förster,1856
- Camptopteroides Viggiani, 1974
- Caraphractus Walker, 1846
- Ceratanaphes Noyes & Valentine, 1989
- Chaetomymar Ogloblin,1946
- Chrysoctonus Mathot,1966
- Cleruchoides Lin & Huber, 2007
- Cleruchus Enock,1909
- Cnecomymar Ogloblin, 1963
- Cremnomymar Ogloblin, 1952
- Cybomymar Noyes & Valentine, 1989
- Dicopomorpha Ogloblin,1955
- Dicopus Enock, 1909
- Dorya Noyes & Valentine, 1989
- Entrichopteris Yoshimoto, 1990
- Eofoersteria Mathot, 1966
- Erdosiella Soyka, 1956
- Erythmelus Enock, 1909
- Eubroncus Yoshimoto, Kozlov & Trjapitzin, 1972
- Eucleruchus Ogloblin, 1940
- Eustochomorpha Girault, 1915
- Eustochus Haliday, 1833
- Formicomymar Yoshimoto, 1990
- Gahanopsis Ogloblin,1946
- Ganomymar De Santis,1972
- Gonatocerus Nees, 1834
- Haplochaeta Noyes et Valentine, 1989
- Himopolynema Taguchi, 1977
- Idiocentrus Gahan, 1927
- Ischiodasys Noyes & Valentine, 1989
- Kalopolynema Ogloblin, 1960
- KikikiKikikiKikiki is a genus of fairyflies from Hawaii....
Huber & Beardsley, 2000 - Kompsomymar Lin & Huber, 2007
- Krokella Huber, 1993
- Kubja Subba Rao, 1984
- Litus Haliday,1833
- Macrocamptoptera Girault, 1910
- Malfattia Meunier, 1901
- Mimalaptus Noyes & Valentine, 1989
- Mymar Curtis,1829
- Mymarilla Westwood, 1879
- Myrmecomymar Yoshimoto, 1990
- Narayanella Subba Rao, 1976
- Neolitus Ogloblin, 1935
- Neomymar Crawford, 1913
- Neostethynium Ogloblin, 1964
- Neserythmelus Noyes & Valentine, 1989
- Nesomymar Valentine, 1971
- Nesopatasson Valentine, 1971
- Nesopolynema Ogloblin, 1952
- Notomymar Doutt & Yoshimoto, 1970
- Omyomymar Schauff, 1983
- Oncomymar Ogloblin, 1957
- Ooctonus Haliday,1833
- Palaeoneura Waterhouse, 1915
- Palaeopatasson Witsack, 1986
- Paracmotemnus Noyes & Valentine, 1989
- Paranaphoidea Girault, 1913
- Parapolynema Fidalgo, 1982
- Parastethynium Lin & Huber in Lin, Huber & La Salle, 2007
- Platyfrons Yoshimoto, 1990
- Platypolynema Ogloblin, 1960
- Platystethynium Ogloblin, 1946
- Polynema Haliday,1833
- Polynemoidea Girault, 1913
- Polynemula Ogloblin, 1967
- Prionaphes Hincks, 1961
- Pseudanaphes Noyes & Valentine, 1989
- Pseudocleruchus Donev & Huber, 2002
- Ptilomymar Annecke & Doutt, 1961
- Restisoma Yoshimoto, 1990
- Richteria Girault, 1920
- Schizophragma Ogloblin, 1949
- Scleromymar Noyes & Valentine, 1989
- Scolopsopteron Ogloblin, 1952
- Steganogaster Noyes & Valentine, 1989
- Stephanocampta Mathot,1966
- Stephanodes Enock,1909
- Stethynium Enock, 1909
- Tanyostethium Yoshimoto, 1990
- Tetrapolynema Ogloblin, 1946
- Zelanaphes Noyes & Valentine, 1989
Fossil genera
The following are the fossilFossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...
genera classified under Mymaridae:
- †Carpenteriana Yoshimoto, 1975
- †Enneagmus Yoshimoto, 1975
- †Eoanaphes Huber in Huber & Greenwalt, 2011
- †Eoeustochus Huber in Huber & Greenwalt, 2011
- †Macalpinia Yoshimoto, 1975
- †MyanmymarMyanmymarMyanmymar is an extinct genus of fairyfly preserved in amber from Myanmar. It has only one species, Myanmymar aresconoides. It is dated to the Early Cretaceous, at least a hundred million years ago. As of 2011, it is the oldest known fossil mymarid.-Discovery and geologic time range:Myanmymar...
Huber in Huber & Poinar, 2011 - †Triadomerus Yoshimoto, 1975
Collection and preservation
In the 19th century, Curtis described the methods by which Haliday collected fairyflies as thus:
The best modern collection method is using Malaise trap
Malaise trap
A Malaise trap is a large, tent-like structure used for trapping flying insects particularly Hymenoptera and Diptera. The trap is made of a material such as terylene netting and can be various colours. Insects fly into the tent wall and are funnelled into a collecting vessel attached to highest point...
s. It requires little maintenance and can collect insects in great abundance. Other effective methods include yellow pan trapping, sweep nets, and suction trapping. Direct collection from leaf litter with Berlese funnels can also result in specimens that can not be collected by other means.
Rearing is also another method that can bring the most rewards. This can be done with wild host eggs or laboratory prepared host eggs that are exposed in suitable habitats outside. They can then be taken in after a suitable amount of time has passed and examined for developing fairyflies. With this method, it is possible to observe the life history and determine the hosts of particular species of discovered fairyflies.
Preservation is a problem for fairyflies (and other small chalcid wasps). Their tiny sizes require special methods. Specimens have to be dried, if collected wet, e.g. if ethyl alcohol is used as the killing agent
Killing jar
A killing jar is a device used by entomologists to kill captured insects quickly and with minimum damage. The jar, typically glass, must be hermetically sealable and usually has a thin layer of hardened plaster of paris on the bottom to absorb the insecticide. The insecticide will then slowly...
. Drying can make specimens extremely brittle, so additional care should also be taken not to disintegrate them. Mounting specimens (preferably in permanent slides) is also time-consuming and requires a fair amount of practice. They are gummed or glued into cards, as they can not be mounted on pins like larger insects.
See also
- TrichogrammatidaeTrichogrammatidaeThe family Trichogrammatidae are tiny wasps in the Chalcidoidea that include some of the smallest of all insects, with most species having adults less than 1 mm in length. There are over 840 species in ca. 80 genera worldwide. Trichogrammatids parasitize the eggs of many different orders of insects...
- Biological pest controlBiological pest controlBiological control of pests in agriculture is a method of controlling pests that relies on predation, parasitism, herbivory, or other natural mechanisms...
- Parasitoid wasp