The Eucharitidae are a family of parasitic

wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...

s. Eucharitid wasps are members of the

superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...

Chalcidoidea Chalcid wasps (, , for their metallic colour) are insects within the superfamily Chalcidoidea, part of the order Hymenoptera. The superfamily contains some 22,500 known species, and an estimated total diversity of more than 500,000 species, m ...

and consist of four subfamilies: Akapalinae, Eucharitinae, Gollumiellinae, and Oraseminae. Most of the 42 genera and >400 species of Eucharitidae are members of the subfamilies Oraseminae and Eucharitinae, and are found in tropical regions of the world.Heraty, John. Eucharitidae. Hymenopteran Systematics, University of California, Riverside (2002): Web. 16 Sep. 2011. Eucharitids are specialized

parasitoids In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...

ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...

s, meaning each species is usually only parasitic of one genus of ant. Furthermore, they are one of the few parasitoids that have been able to use ants as hosts, despite ants’ effective defense systems against most parasitoids.Lachaud, Jean-Paul and Perez-Lachaud, Gavriela. Impact of natural parasitism by two eucharitid wasps on a potential biocontrol agent ant in southeastern Mexico. Biological Control 48 (2009): 92-99.Heraty, J.M. Biology and importance of two eucharitid parasites of ''Wasmannia'' and ''Solenopsis''. (1994): Pages 104-120 in Williams, D. (ed), Exotic Ants: Biology, Impact and Control of Introduced Species. Boulder, CO: Westview Press. 332 pp. Eucharitid parasitism occurs year-round, with a majority of it occurring during hot and humid months. However, the amount of parasitism that occurs depends primarily on the size of the ant colony and the number of host

pupae A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...

in them, and not on the season.

Life cycle

Female eucharitids

oviposit The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...

rows of eggs into plant tissue, such as leaves and stems, away from ant colonies.Varone, L. and Briano, J. Bionomics of Orasema simplex (Hymenoptera: Eucharitidae), a parasitoid of Solenopsis fire ants (Hymenoptera: Formicidae) in Argentina. Biological Control 48 (2009): 204-209. The eggs are a translucent white and are about 0.19 mm long and 0.08 mm wide. They are elliptical and flat on one side. As the eggs mature, they turn a brown color and ten days after oviposition, they hatch. The larvae are solely responsible for their entry into the ant colony and the

parasitism Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The en ...

of their host. They are 0.13 mm long and are able to travel several inches on the leaf but do not leave the egg cluster. After six to seven days, they attach themselves to foraging ants heading back to their

brood Brood may refer to: Nature * Brood, a collective term for offspring * Brooding, the incubation of bird eggs by their parents * Bee brood, the young of a beehive * Individual broods of North American periodical cicadas: ** Brood X, the largest br ...

; however, sometimes they will attach themselves to other insects, using them as

intermediate host In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include ...

s. Once in the brood, the larvae will attach to their host larva. Some Eucharitidae are external

parasites Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The en ...

while others are internal parasites; however, all eucharitid species finish their development as ectoparasites. Limited feeding on the host occurs until the host pupates, after which, most of it is consumed by the wasp. Usually, only one parasite per host is found, but in some cases,

superparasitism Superparasitism is a form of parasitism in which the host (typically an insect larva such as a caterpillar) is attacked more than once by a single species of parasitoid. Multiparasitism or coinfection, on the other hand, occurs when the host has b ...

occurs, and two to four wasps will attach to, and emerge from one host. Once the wasps emerge, the ant colony grooms and feeds them as if they were part of the ants’ brood. In some instances, worker ants have been observed assisting the wasps to emerge from their host. The wasps gain acceptance in these ways, and the ants show no signs of aggression because the wasps acquire their host's odor upon entry into the colony. By mimicking the odor of their host, eucharitid wasps are able to keep themselves safe until the scent wears off, at which point they leave the ant colony and begin mating. Adult wasps emerge from the ant nest in the morning; the males emerge before the females. In most cases, the males swarm one to two feet above the nest, and as soon as the females emerge, mating occurs. However, the males of certain species, such as ''Kapala terminalis'', calmly wait on foliage surrounding the nest until the females emerge. Many times, the males will begin mating with the females before they have a chance to take flight, and in some instances, mating will occur while the wasps are still inside the ants' nest.Clausen, C.P. The habits of the Eucharidae. Psyche 48 (1941): 57-69. After mating, the females lay all their eggs in one day. The egg capacity of each female is 1000 to 10000.

Anatomy

Most eucharitids are between 2.0 and 5.4 mm long and vary from metallic colors to black. They have 13 or fewer segments on their antennae, but some have up to 26.Heraty, J.M. Family Eucharitidae. (1995) Pages 309-314. in Gauld, I.D.; Hanson, P., Hymenoptera of Costa Rica. Oxford: Oxford University Press. ?? The main anatomical characteristics of eucharitid wasps are: *The prepectus lies in the same plane as, and fused, to

pronotum The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum (dorsal), the prosternum (ventral), and the propleuron (lateral) on e ...

. *The

is small and hidden by the head

dorsally Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position provi ...

. *The gaster is usually relatively small with a very long

petiole Petiole may refer to: *Petiole (botany), the stalk of a leaf, attaching the blade to the stem *Petiole (insect anatomy) In entomology, petiole is the technical term for the narrow waist of some hymenopteran insects, especially ants, bees, and ...

. *The forewing marginal vein is moderately long and stigmal, and the postmarginal veins are extremely short. *Moderately large, about 2.0 to 5.4 mm in length.Pitkin, B. R. Dr. “Eucharitidae”. Natural History Museum (2004) Universal Chalcidoidea Database Notes on families. Web. 30 Sep. 2011.

Subfamilies

Oraseminae

Oraseminae are parasitic on the ant genus ''Pheidole'' (Myrmicinae), as well as the

fire ant Fire ants are several species of ants in the genus ''Solenopsis'', which includes over 200 species. ''Solenopsis'' are stinging ants, and most of their common names reflect this, for example, ginger ants and tropical fire ants. Many of the nam ...

in southern

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

and the little red fire ant in the

Caribbean The Caribbean ( , ; ; ; ) is a region in the middle of the Americas centered around the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean, mostly overlapping with the West Indies. Bordered by North America to the north, Central America ...

. They are distributed worldwide in

tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...

regions, such as

Costa Rica Costa Rica, officially the Republic of Costa Rica, is a country in Central America. It borders Nicaragua to the north, the Caribbean Sea to the northeast, Panama to the southeast, and the Pacific Ocean to the southwest, as well as Maritime bo ...

Ecuador Ecuador, officially the Republic of Ecuador, is a country in northwestern South America, bordered by Colombia on the north, Peru on the east and south, and the Pacific Ocean on the west. It also includes the Galápagos Province which contain ...

Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...

, and Argentina. They are also native to

Texas Texas ( , ; or ) is the most populous U.S. state, state in the South Central United States, South Central region of the United States. It borders Louisiana to the east, Arkansas to the northeast, Oklahoma to the north, New Mexico to the we ...

Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...

Massachusetts Massachusetts ( ; ), officially the Commonwealth of Massachusetts, is a U.S. state, state in the New England region of the Northeastern United States. It borders the Atlantic Ocean and the Gulf of Maine to its east, Connecticut and Rhode ...

Delaware Delaware ( ) is a U.S. state, state in the Mid-Atlantic (United States), Mid-Atlantic and South Atlantic states, South Atlantic regions of the United States. It borders Maryland to its south and west, Pennsylvania to its north, New Jersey ...

,Ashmead, W.H. Studies on North American Chalcididae, with descriptions of new species from Florida. Transactions of the American Entomological Society 12 (1885 0: 10-14. and Colorado. An orasemine female punctures the plant tissue on which she will oviposit, and places an egg inside of each incision. The eggs are deposited on the edges of the underside of leaves, and are sometimes found in the buds and stems of the plant. They have been known to oviposit on blueberry leaves, tea leaves (Das), oak leaves, olive leaves, mango leaves, flower heads, and banana fingers. Species of Eucharitidae are consistent in their choice of leaf for oviposition. Because orasemine eggs are laid on plants that are not always visited by foraging ants, the larvae will most often attach to

intermediate hosts In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include a ...

,Das, G.M. Preliminary studies on the biology of ''Orasema assectator'' Kerrich (Hymenoptera: Eucharitidae) parasitic on ''Pheidole'' and causing damage to leaves of tea in Assam. Bulletin of Entomological Research 54 (1963): 393-398. or “insect prey” that will eventually be devoured by ants. Sometimes, however, foraging ants are present and there is no need for an intermediate host. Upon arriving at the ants’ brood, the larva burrows into the host's

thorax The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main di ...

and feeds there. The wasp feeds until the host larva pupates; after the host pupates, the wasp will resume feeding until it emerges from its host. The average time of development from planidia to adult is 29.5 days; the average pupal stage is 8.2 days. Because orasemine larvae, pupae, and adults are easily distinguished from their host,Vander Meer, R.K., Jouvenaz, D.P. and Wojcik, D.P. Chemical mimicry in a parasitoid (Hymenoptera: Eucharitidae) of fire ants (Hymenoptera: Formicidae). Journal of Chemical Ecology 15 (1989): 2247-2261. the wasps disguise themselves by passively obtaining the odor of the fire ants. After a few days in the nest, the odor wears off and the ants begin to notice the wasps are not a part of their brood. At this point, the wasps leave the nest to mate and lay eggs. Though orasemines have a high fertility rate, only a small percentage of eggs survive to adulthood.

Eucharitinae

Eucharitinae are parasites of poneromorph ants, Ectatommatinae, Ponerinae, and Formicinae, although one genus from Australia is parasitic on the

bulldog ant ''Myrmecia'' is a genus of ants first established by Danish zoologist Johan Christian Fabricius in 1804. The genus is a member of the subfamily Myrmeciinae of the family Formicidae. ''Myrmecia'' is a large genus of ants, comprising at least ...

. Female Eucharitinae bear up to 4500 eggs and begin oviposition soon after emerging from the nest. They deposit their eggs in groups of eight to 15 on plant buds, on the undersides of leaves or on fruit skin. Eucharitine larvae attach themselves to foraging ants and do not use an intermediate host. Some species of Eucharitinae, such as ''Kapala terminalis'' and other ''Kapala'' sp., have been known for their jumping capabilities. They stand erect on the plant on which they hatched, and without any distinguishable preparation, jump about 10 mm from the leaf onto a foraging ant. The larvae are external parasitoids of their hosts, and are not noticed due to their acquisition of the host’s odor. After the wasps are fully developed, they emerge in large numbers. The males swarm around the nest in wait of the females. Mating takes place immediately, and oviposition occurs soon after. A fossil genus, '' Palaeocharis'' is known from Eocene

Baltic amber Baltic amber or succinite is amber from the Baltic region, home of its largest known deposits. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that this forested region provided the re ...

Gollumiellinae

Gollumiellinae are unique in that they hook their eggs onto the plants and connect a ropey secretion to them, which stand erect. This acts as an attraction mechanism for ''

Paratrechina ''Paratrechina'' is one of seven ant genera (alongside '' Euprenolepis, Nylanderia, Paraparatrechina, Prenolepis, Pseudolasius,'' and '' Zatania'') in the ''Prenolepis'' genus-group from the subfamily Formicinae (tribe Lasiini). Six species ...

'' ants. Gollumiellinae larvae burrow into the hosts' thoraces and feed there. The rest of its life cycle is similar to the aforementioned life cycle of eucharitids. Two Indo‐Pacific genera are included here:Heraty J, Hawks D, Kostecki JS, Carmichael A (2004) Phylogeny and behaviour of the Gollumiellinae, a new subfamily of the ant‐parasitic Eucharitidae (Hymenoptera: Chalcidoidea). ''Systematic Entomology'' 29 544-559. * '' Anorasema'' Boucek, 1988 * '' Gollumiella'' Hedqvist, 1978

Akapalinae

This subfamily is monotypic, containing only the genus '' Akapala'' (Girault, 1934).

Biological control

Eucharitids are candidates for

biological control Biological control or biocontrol is a method of controlling pests, whether pest animals such as insects and mites, weeds, or pathogens affecting animals or plants by using other organisms. It relies on predation, parasitism, herbivory, or o ...

because each subfamily targets a specific ant genus. On the other hand, some eucharitids, including the genus ''Kapala'', are classified as

pests PESTS was an anonymous American activist group formed in 1986 to critique racism, tokenism, and exclusion in the art world. PESTS produced newsletters, posters, and other print material highlighting examples of discrimination in gallery represent ...

for several different reasons. First, a few days after

oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typica ...

occurs, the leaves dry up where each egg was located. Second, tea leaves with such marks reportedly do not have as strong of a flavor as tea leaves without these marks. Third, not all eucharitids have a significant effect on the size of their hosts' colony. ''Orasema'' species are sometimes used as means of biological control for many ants, including the fire ant and little fire ant, because

pesticides Pesticides are substances that are used to pest control, control pest (organism), pests. They include herbicides, insecticides, nematicides, fungicides, and many others (see table). The most common of these are herbicides, which account for a ...

can cause damage to the environment, and they do not always work on fire ants. In addition, at least two eucharitid species groups are specialized parasites of these ants. Heraty stated, “Leaving aside the philosophical problems associated with introducing any organism for biological control, I believe that species of ''Orasema'' do have potential as biological control agents and deserve more study.”

References

Additional references

UC Riverside Eucharitid Page
pdfs

External links

images of pinned specimens
Catalog of World Eucharitidae
. John Heraty. 2014 {{Authority control Chalcidoidea Apocrita families