Tachinidae

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Tachinidae
Temporal range: Eocene–Recent
Thomas Bresson - Tachina fera (by).JPG
Tachina fera
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
(unranked): Eremoneura
(unranked): Cyclorrhapha
Section: Schizophora
Subsection: Calyptratae
Superfamily: Oestroidea
Family: Tachinidae
Bigot, 1853 [1]
Subfamilies
Diversity
1,523 genera

The Tachinidae are a large and variable family of true flies within the insect order Diptera, with more than 8,200 known species and many more to be discovered. Over 1,300 species have been described in North America alone. Insects in this family commonly are called tachinid flies or simply tachinids. As far as is known, they all are protelean parasitoids, or occasionally parasites, of arthropods, usually other insects. The family is known from many habitats in all zoogeographical regions and is especially diverse in South America. [2]

Contents

"Tachinidae" by Harold Maxwell-Lefroy, 1909 10-Indian-Insect-Life - Harold Maxwell-Lefroy - Tachinidae.jpg
"Tachinidae" by Harold Maxwell-Lefroy, 1909

Life cycle

Reproductive strategies vary greatly between tachinid species, largely, but not always clearly, according to their respective life cycles. Many species are generalists rather than specialists. [3] Comparatively few are restricted to a single host species, so there is little tendency towards the close co-evolution one finds in the adaptations of many specialist species to their hosts, such as are typical of protelean parasitoids among the Hymenoptera.

Larvae (maggots) of most members of this family are parasitoids (developing inside a living host, ultimately killing it). In contrast a few are parasitic (not generally killing the host). Tachinid larvae feed on the host tissues, either after having been injected into the host by the parent, or penetrating the host from outside. Various species have different modes of oviposition and of host invasion. Typically, tachinid larvae are endoparasites (internal parasites) of caterpillars of butterflies and moths, or the eruciform larvae of sawflies. For example, they have been found to lay eggs in African sugarcane borer larva, a species of moth common in sub-Saharan Africa, [4] as well as the more northerly Arctic woolly bear moth. [5] However, some species attack adult beetles and some attack beetle larvae. Others attack various types of true bugs, and others attack grasshoppers; a few even attack centipedes. Also parasitised are bees, wasps and sawflies. [6]

Tachinid eggs (possibly Trichopoda pennipes) on Leptoglossus occidentalis Tachinid eggs on Leptoglossus.jpg
Tachinid eggs (possibly Trichopoda pennipes ) on Leptoglossus occidentalis

Oviposition and ovoviviparity

Probably the majority of female tachinids lay white, ovoid eggs with flat undersides onto the skin of the host insect. Imms [7] mentions the genera Gymnosoma , Thrixion, Winthemia, and Eutachina as examples. In a closely related strategy some genera are effectively ovoviviparous (some authorities prefer the term ovolarviparous [8] ) and deposit a hatching larva onto the host. For example, this occurs in Tachinidae species which parasitize the butterfly Danaus chrysippus in Ghana. [9] The free larvae immediately bore into the host's body. Illustrative genera include Exorista and Voria . Many tachinid eggs hatch quickly, having partly developed inside the mother's uterus, which is long and often coiled for retaining developing eggs. However, it is suggested that the primitive state probably is to stick unembryonated eggs to the surface of the host. [8]

Many other species inject eggs into the host's body, using the extensible, penetrating part of their ovipositor, sometimes called the oviscapt, which roughly translates to "egg digger". Species in the genera Ocyptera, Alophora , and Compsilura are examples.

Istocheta aldrichi egg on Popillia japonica (Japanese beetle) Istocheta aldrichi egg on Japanese beetle.jpg
Istocheta aldrichi egg on Popillia japonica (Japanese beetle)
Most tachinids are dull colored, resembling house flies Tachinidae.jpg
Most tachinids are dull colored, resembling house flies
Ormia ochracea, notable for its acute directional hearing Ormia ochracea (gravid female).jpg
Ormia ochracea , notable for its acute directional hearing

In many species only one egg is laid on or in any individual host, and accordingly such an egg tends to be large, as is typical for eggs laid in small numbers. They are large enough to be clearly visible if stuck onto the outside of the host, and they generally are so firmly stuck that eggs cannot be removed from the skin of the host without killing them. Furthermore, scientists have observed in studies with the host cabbage looper that being glued to the host insect helps maggots burrow into the larva, where they remain until fully developed. [11]

Yet another strategy of oviposition among some Tachinidae is to lay large numbers of small, darkly coloured eggs on the food plants of the host species. Sturmia, Zenillia, and Gonia are such genera.

Many tachinids are important natural enemies of major insect pests, and some species actually are used in biological pest control; for example, some species of tachinid flies have been introduced into North America from their native lands as biocontrols to suppress populations of alien pests. [12] Conversely, certain tachinid flies that prey on useful insects are themselves considered as pests; they can present troublesome problems in the sericulture industry by attacking silkworm larvae. One particularly notorious silkworm pest is the Uzi fly ( Exorista bombycis ).

Another reproductive strategy is to leave the eggs in the host's environment; for example, the female might lay on leaves, where the host is likely to ingest them. Some tachinids that are parasitoids of stem-boring caterpillars deposit eggs outside the host's burrow, letting the first instar larvae do the work of finding the host for themselves. In other species, the maggots use an ambush technique, waiting for the host to pass and then attacking it and burrowing into its body.

Adult tachinids are not parasitic, but either do not feed at all or visit flowers, decaying matter, or similar sources of energy to sustain themselves until they have concluded their procreative activities. Their non-parasitic behaviour after eclosion from the pupa is what justifies the application of the term "protelean".

Description

Tachinid flies mating 20170403 TachinidFly.jpg
Tachinid flies mating

Tachinid flies are extremely varied in appearance. Some adult flies may be brilliantly colored and resemble blow-flies (family Calliphoridae). Most however are rather drab, some resembling house flies. However, tachinid flies commonly are more bristly and more robust. Also, they usually have a characteristic appearance. They have three-segmented antennae, a diagnostically prominent postscutellum bulging beneath the scutellum (a segment of the mesonotum). They are aristate flies, and the arista usually is bare, though sometimes plumose. The calypters (small flaps above the halteres) are usually very large. Their fourth long vein bends away sharply.

Adult flies feed on flowers and nectar from aphids and scale insects. As many species typically feed on pollen, they can be important pollinators of some plants, especially at higher elevations in mountains where bees are relatively few.

The taxonomy of this family presents many difficulties. It is largely based on morphological characters of the adult flies, but also on reproductive habits and on the immature stage.

As biological pest control

Some tachinid flies parasitize pest species. This has allowed them to be used as biological control agents by farmers. Some Tachinidae are generalists; for instance, Compsilura concinnata uses, at least, 200 different hosts, and they are not safe to be used as biological controls. Others are more specialized and are safer; for instance, Istocheta aldrichi , which only attacks the Japanese beetle. [13] [14] [15]

Evolution

This clade appears to have originated in the middle Eocene. [16] The oldest known putatively tachinid fossil ( Lithexorista ) dates from the Eocene Green River Formation in Wyoming. [17] [18] [19]

See also

Related Research Articles

<span class="mw-page-title-main">Fly</span> Order of insects

Flies are insects of the order Diptera, the name being derived from the Greek δι- di- "two", and πτερόν pteron "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced mechanosensory organs known as halteres, which act as high-speed sensors of rotational movement and allow dipterans to perform advanced aerobatics. Diptera is a large order containing an estimated 1,000,000 species including horse-flies, crane flies, hoverflies, mosquitoes and others, although only about 125,000 species have been described.

<span class="mw-page-title-main">Parasitoid</span> Organism that lives with its host and kills it

In evolutionary ecology, a parasitoid is an organism that lives in close association with its host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionary strategies within parasitism, distinguished by the fatal prognosis for the host, which makes the strategy close to predation.

<span class="mw-page-title-main">Ovoviviparity</span> Gestation type

Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop inside eggs that remain in the mother's body until they are ready to hatch.

<span class="mw-page-title-main">Oestroidea</span> Superfamily of flies

Oestroidea is a superfamily of Calyptratae including the blow flies, bot flies, flesh flies, and their relatives. It occurs worldwide and has about 15,000 described species.

<span class="mw-page-title-main">Parasitoid wasp</span> Group of wasps

Parasitoid wasps are a large group of hymenopteran superfamilies, with all but the wood wasps (Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, sooner or later causing the death of these hosts. Different species specialise in hosts from different insect orders, most often Lepidoptera, though some select beetles, flies, or bugs; the spider wasps (Pompilidae) exclusively attack spiders. More rarely, parasitoid wasps may use plant seeds as hosts, such as Torymus druparum.

<i>Delia</i> (fly) Genus of flies

Delia flies are members of the Anthomyiidae family within the superfamily Muscoidae. The identification of different species of Delia can be very difficult for non-specialists as the diagnostic characteristics used for immature and/or female specimens may be inconsistent between species. Past taxonomic keys were not as comprehensive in their identification of Delia specimens; they were either too reliant on genetic characteristics, focused solely on a specific life stage, or were focused only on certain species. However current taxonomic keys aim to be more thorough by not only including morphological diagnostics for males, females, and immature specimens of various species, but also their genetic make-up or molecular barcode.

<i>Senostoma</i> Genus of flies

Senostoma is a genus of parasitoid tachinid flies in the family Tachinidae. Endemic to Australasia, the flies are medium-sized, bristly, and long-legged.

<i>Tachina fera</i> Species of fly

Tachina fera is a species of fly in the genus Tachina of the family Tachinidae. It was first described by Carl Linnaeus in 1761.

<i>Gymnosoma clavatum</i> Species of fly

Gymnosoma clavatum is a species of tachinid flies in the genus Gymnosoma of the family Tachinidae.

<i>Prosena siberita</i> Species of fly

Prosena siberita is a species of fly in the family Tachinidae.

<span class="mw-page-title-main">Blondeliini</span> Tribe of flies

Blondeliini is a tribe of parasitic flies in the family Tachinidae. Larvae are parasitoids of other insects, mostly beetles and caterpillars. Although nearly cosmopolitan, its greatest diversity is in the New World and especially in South America.

<i>Cyzenis albicans</i> Species of fly

Cyzenis albicans is a species of fly in the family Tachinidae. A parasitoid, it lays its eggs on leaves of oak, maple, birch and other trees, so that when the leaves are consumed by the larvae of the host winter moth, the eggs hatch inside the larvae. The fly is native to Europe and Asia but has been introduced into North America as a biological control agent of the invasive winter moth.

<span class="mw-page-title-main">Charles Henry Tyler Townsend</span> American entomologist

Charles Henry Tyler Townsend was an American entomologist specializing in the study of tachinids (Tachinidae), a large and diverse family of flies (Diptera) with larvae that are parasitoids of other insects. He was perhaps the most prolific publisher of new tachinids, naming and describing some 3000 species and genera. He made important contributions to the biological control of insect pests and he was the first to identify the insect vector of a debilitating disease in Peru. Townsend was also a controversial figure and criticism of his approach to insect taxonomy continues to this day.

<i>Exorista larvarum</i> Species of fly

Exorista larvarum is a Palaearctic species of fly in the family Tachinidae.

<i>Compsilura concinnata</i> Species of fly

Compsilura concinnata is a parasitoid native to Europe that was introduced to North America in 1906 to control the population of an exotic forest, univoltine, spongy moth named Lymantria dispar. It is an endoparasitoid of larvae and lives with its host for most of its life. Eventually the parasitoid ends up killing the host and occasionally eating it. It attacks over 200 host species, mainly insects from the Orders: Coleoptera, Lepidoptera and Hymenoptera. Since this parasite has the ability to attack many different types of hosts, the organism has spilled over from the intended forest systems into other areas, like agricultural fields, affecting cabbage pests including the cabbage looper (Trichoplusia); the cabbage worm ; and even other invasive species such as the brown-tail moth. However, it also attacks native, non-pest insects such as the Cecropia moth and American moon moth.

<i>Archytas apicifer</i> Species of fly

Archytas apicifer is a medium to large sized Nearctic tachinid fly. The species name was authored by the German entomologist Johann Friedrich Jaennicke (1867) and presumably named after the Greek classical philosopher and mathematician Archytas. The larvae are parasites of several caterpillar species.

Lespesia archippivora is a species of tachinid fly, which, like all tachinids, are parasitoids of other arthropods. L. archippivora lives in the body of its host resulting in its death. This is not uncommon since it is estimated that about 10% of all insects are parasitoids. L. archippivora is a generalist and able to infect at least 25 lepidopteran species in addition to one species of Hymenoptera. It is common in North America and other species exist worldwide. One study suggests the species is bivoltine.

Sturmiopsis inferens is a species of fly in the family Tachinidae. It is native to Asia and is a parasitoid of various moth species whose larvae feed inside the stems of sugarcane, rice and other large grasses, including the Gurdaspur borer and the sugarcane shoot borer.

<i>Zelia vertebrata</i> Species of fly

Zelia vertebrata is a species of bristle fly in the family Tachinidae. It is a long-bodied fly with strong abdominal bristles and a distinctive abdominal pattern that resembles vertebrae. It has a widespread North American range, with records stretching from east to west from Washington to Maine, north to south from Québec to Florida. Its larval stage parasitizes beetles. It is most active during the day.

Exorista mella is a tachinid fly of the genus Ezorista within the family Tachinidae of the order Diptera. They are typically found in the United States and Canada. Within the U.S in the state of Arizona they have been found in both mountainous and agricultural regions. E. mella is a parasitoid fly, a polyphagous generalist which parasitizes a variety of hosts.

References

  1. ↑ Systema Dipterorum: Tachinidae
  2. ↑ James E. O'Hara; Shannon J. Henderson (December 18, 2018). "World Genera of the Tachinidae (Diptera) and Their Regional Occurrence" (PDF).
  3. ↑ John O. Stireman (2006). "Tachinidae: Evolution, Behavior, and Ecology". Annual Review of Entomology. 51: 525–555. doi:10.1146/annurev.ento.51.110104.151133.
  4. ↑ Hastings, H.; Conling, D.E.; Graham, D.Y. (1988). "Notes on the natural host surveys and laboratory rearing of Goniozus natalensis Gordh (Hymenoptera: Bethylidae), a parasitoid of Eldana saccharina Walker (Lepidoptera: Pyralidae) larvae from Cyperus papyrus L. in Southern Africa" (PDF). Journal of the Entomological Society of Southern Africa. 51: 1.
  5. ↑ Morewood, W. Dean; Wood, D. Monty (2002). "Host utilization byExorista thula Wood (sp. nov.) and Chetogena gelida (Coquillett) (Diptera: Tachinidae), parasitoids of arctic Gynaephora species (Lepidoptera: Lymantriidae)". Polar Biology. 25 (8): 575–582. doi:10.1007/s00300-002-0382-y. S2CID   20312863.
  6. ↑ "Parasitic Flies - Ontario AppleIPM". www.omafra.gov.on.ca.
  7. ↑ Imms' General Textbook of Entomology: Volume 1: Structure, Physiology and Development Volume 2: Classification and Biology. Berlin: Springer. 1977. ISBN   978-0-412-61390-6.
  8. 1 2 Wood, D. M. 1987. Chapter 110. Tachinidae. Pp. 1193-1269 in McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. and D.M. Wood (eds.), Manual of Nearctic Diptera. Volume 2. Agriculture Canada Monograph 28: i-vi, 675-1332.
  9. ↑ Edmunds, Malcolm (1976-03-01). "Larval mortality and population regulation in the butterfly Danaus chrysippus in Ghana". Zoological Journal of the Linnean Society. 58 (2): 129–145. doi:10.1111/j.1096-3642.1976.tb00823.x. ISSN   0024-4082.
  10. ↑ Miles RN, Robert D, Hoy RR (1995). "Mechanically coupled ears for directional hearing in the parasitoid fly Ormia ochracea". The Journal of the Acoustical Society of America. 98 (6): 3059–3070. Bibcode:1995ASAJ...98.3059M. doi:10.1121/1.413830. PMID   8550933.
  11. ↑ Brubaker, R. W. (1968-02-01). "Seasonal Occurrence of Voria ruralis a Parasite of the Cabbage Looper, in Arizona, and Its Behavior and Development in Laboratory Culture1". Journal of Economic Entomology. 61 (1): 306–309. doi:10.1093/jee/61.1.306. ISSN   0022-0493.
  12. ↑ entomology.Wisconsin.edu. Compsilura concinnata, Parasitoid of Gypsy Moth
  13. ↑ O’Hara, J. Tachinidae Resources. Overview of the Tachinidae (Diptera)
  14. ↑ Grenier, S. (1988-04-01). "Applied biological control with Tachinid flies (Diptera, Tachinidae): A review". Anzeiger für Schädlingskunde, Pflanzenschutz, Umweltschutz. 61 (3): 49–56. doi:10.1007/BF01906254. ISSN   0340-7330. S2CID   10085920.
  15. ↑ Bugguide.net. Species Istocheta aldrichi - Winsome Fly
  16. ↑ Zhao, Zhe; Su, Tian-Juan; Chesters, Douglas; Wang, Shi-di; Ho, Simon Y. W.; Zhu, Chao-Dong; Chen, Xiao-lin; Zhang, Chun-Tian (2013). "The Mitochondrial Genome of Elodia flavipalpis Aldrich (Diptera: Tachinidae) and the Evolutionary Timescale of Tachinid Flies". PLOS ONE. 8 (4): e61814. Bibcode:2013PLoSO...861814Z. doi: 10.1371/journal.pone.0061814 . PMC   3634017 . PMID   23626734.
  17. ↑ Townsend, C.H.T. (1921). "Some new muscoid genera ancient and recent". Insecutor Inscitiae Menstruus. 9: 132–134. Retrieved 26 June 2023.
  18. ↑ Evenhuis, N.L. (1994). Catalogue of the fossil flies of the world (Insecta: Diptera). Leiden: Backhuys Publishers. pp. [i] + 1-600.
  19. ↑ Cerretti, Pierfilippo; Stireman, John O.; Pape, Thomas; O’Hara, James E.; Marinho, Marco A. T.; Rognes, Knut; Grimaldi, David A. (2017-08-23). Friedman, Matt (ed.). "First fossil of an oestroid fly (Diptera: Calyptratae: Oestroidea) and the dating of oestroid divergences". PLOS ONE. Public Library of Science (PLoS). 12 (8): e0182101. Bibcode:2017PLoSO..1282101C. doi: 10.1371/journal.pone.0182101 . ISSN   1932-6203. PMC   5568141 . PMID   28832610.
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