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{{Speciesbox
{{Speciesbox
| image = Ormia ochracea (gravid female).jpg
| image = Ormia ochracea (gravid female).jpg
| taxon = Ormia ochracea
| genus = Ormia
| species = ochracea
| authority = ([[Jacques-Marie-Frangile Bigot|Bigot]], 1889)
| display_parents = 3
| synonyms = *''Phasiopteryx montana'' <small>Townsend, 1912</small>
| authority = ([[Jacques-Marie-Frangile Bigot|Bigot]], 1889)<ref name="Bigot1889b">{{cite journal |last= Bigot |first= J. M. F. |date= 1889 |title= Dipteres nouveaux ou peu connus. 34e partie, XLII: Diagnoses de nouvelles especes|journal= Annales de la Société Entomologique de France |volume= 8 |issue= 6 |pages = 253–270 }}</ref>
*''Pyrrosia ochracea''<small> Bigot, 1889</small>
| synonyms = *''Phasiopteryx montana'' <small>[[Charles Henry Tyler Townsend|Townsend]], 1912</small>
*''Pyrrosia ochracea'' <small>[[Jacques-Marie-Frangile Bigot|Bigot]], 1889</small><ref name="Bigot1889b"/>
}}
}}


'''''Ormia ochracea''''' is a small yellow [[nocturnal]] [[fly]] in the family [[Tachinidae]]. It is notable for its parasitism of [[Cricket (insect)|crickets]] and its exceptionally acute directional hearing. The female is attracted to the song of the male cricket and deposits larvae on or around him, as was discovered in 1975 by the zoologist [[William H. Cade]].<ref name=":0">{{cite journal | last=Cade | first=W. | title=Acoustically Orienting Parasitoids: Fly Phonotaxis to Cricket Song | journal=Science | publisher=American Association for the Advancement of Science (AAAS) | volume=190 | issue=4221 | date=1975-12-26 | issn=0036-8075 | doi=10.1126/science.190.4221.1312 | pages=1312–1313}}</ref>
'''''Ormia ochracea''''' is a small yellow [[nocturnal]] [[fly]] in the family [[Tachinidae]].<ref name="OHara2020">{{cite web |last1=O’Hara |first1=James E. |last2=Shannon |first2=J. Henderson |last3=D. Monty |first3=Wood |title=World Checklist of the Tachinidae |url=http://www.nadsdiptera.org/Tach/WorldTachs/Checklist/Tachchlist_ver2.1.pdf |website=Tachinidae Resources |access-date=28 February 2022 |date=5 March 2020}}</ref> It is notable for its parasitism of [[Cricket (insect)|crickets]] and its exceptionally acute directional hearing. The female is attracted to the song of the male cricket and deposits larvae on or around him, as was discovered in 1975 by the zoologist [[William H. Cade]].<ref name=":0">{{cite journal | last=Cade | first=W. | title=Acoustically Orienting Parasitoids: Fly Phonotaxis to Cricket Song | journal=Science | publisher=American Association for the Advancement of Science (AAAS) | volume=190 | issue=4221 | date=1975-12-26 | issn=0036-8075 | doi=10.1126/science.190.4221.1312 | pages=1312–1313| s2cid=85233362 }}</ref>


''Ormia ochracea'' is a [[model organism]] in [[sound localization]] experiments because of its unique "ears", which are complex structures inside the fly's [[prothorax]] near the bases of its front legs. The fly is too small for the time difference of sound arriving at the two ears to be calculated in the usual way, yet it can determine the direction of sound sources with exquisite precision. The [[tympanic membrane]]s of opposite ears are directly connected mechanically, allowing resolution of nanosecond time differences<ref>{{cite journal | last=Miles | first=R. N. | last2=Robert | first2=D. | last3=Hoy | first3=R. R. | title=Mechanically coupled ears for directional hearing in the parasitoid fly Ormia ochracea | journal=The Journal of the Acoustical Society of America | publisher=Acoustical Society of America (ASA) | volume=98 | issue=6 | year=1995 | issn=0001-4966 | doi=10.1121/1.413830 | pages=3059–3070|pmid=8550933}}</ref><ref>{{cite journal | last=Robert | first=D. | last2=Miles | first2=R.N. | last3=Hoy | first3=R.R. | title=Directional hearing by mechanical coupling in the parasitoid fly Ormia ochracea | journal=Journal of Comparative Physiology A | publisher=Springer Science and Business Media LLC | volume=179 | issue=1 | year=1996 | issn=0340-7594 | doi=10.1007/bf00193432 | pages=29–44|pmid=8965258}}</ref> and requiring a new neural coding strategy. Various research groups have designed low-noise differential microphones inspired by ''O. ochracea''’s directionally sensitive hearing system.
''Ormia ochracea'' is a [[model organism]] in [[sound localization]] experiments because of its unique "ears", which are complex structures inside the fly's [[prothorax]] near the bases of its front legs. The fly is too small for the time difference of sound arriving at the two ears to be calculated in the usual way, yet it can determine the direction of sound sources with exquisite precision. The [[tympanic membrane]]s of opposite ears are directly connected mechanically, allowing resolution of nanosecond time differences<ref>{{cite journal | last1=Miles | first1=R. N. | last2=Robert | first2=D. | last3=Hoy | first3=R. R. | title=Mechanically coupled ears for directional hearing in the parasitoid fly Ormia ochracea | journal=The Journal of the Acoustical Society of America | publisher=Acoustical Society of America (ASA) | volume=98 | issue=6 | year=1995 | issn=0001-4966 | doi=10.1121/1.413830 | pages=3059–3070|pmid=8550933}}</ref><ref>{{cite journal | last1=Robert | first1=D. | last2=Miles | first2=R.N. | last3=Hoy | first3=R.R. | title=Directional hearing by mechanical coupling in the parasitoid fly Ormia ochracea | journal=Journal of Comparative Physiology A | publisher=Springer Science and Business Media LLC | volume=179 | issue=1 | year=1996 | issn=0340-7594 | doi=10.1007/bf00193432 | pages=29–44|pmid=8965258| s2cid=21452506 }}</ref> and requiring a new [[neural coding]] strategy. Various research groups have designed low-noise differential microphones inspired by ''O. ochracea''’s directionally sensitive hearing system.


== Distribution ==
== Distribution ==
''Ormia ochracea'' is native to the southeastern United States, including states such as [[Texas]] and [[Florida]].<ref>{{Cite journal|last=Sabrosky|first=C.W.|date=1953|title=Taxonomy and host relations of the tribe Ormiini in the Western Hemisphere (Diptera, Larvaevoridae)|journal=Proceedings of the Entomological Society of Washington|volume=55|pages=167–183}}</ref> ''O. ochracea'' is also found throughout North America, South America, and the Caribbean,<ref name=":1">{{Cite journal|last=Lehmann|first=Gerlind U.C.|date=January 2003|title=Review of Biogeography, Host Range and Evolution of Acoustic Hunting in Ormiini (Insecta, Diptera, Tachinidae), Parasitoids of Night-calling Bushcrickets and Crickets (Insecta, Orthoptera, Ensifera)|url=https://semanticscholar.org/paper/d2fb7956a41bda9d43625e7079f6cf83301f3b52|journal=Zoologischer Anzeiger - A Journal of Comparative Zoology|volume=242|issue=2|pages=107–120|doi=10.1078/0044-5231-00091|issn=0044-5231}}</ref> though its exact range is not known.
''Ormia ochracea'' is native to the southeastern United States, including states such as [[Texas]] and [[Florida]].<ref name="Sabrosky1953">{{Cite journal|last=Sabrosky|first=C.W.|date=1953|title=Taxonomy and host relations of the tribe Ormiini in the Western Hemisphere (Diptera, Larvaevoridae)|journal=Proceedings of the Entomological Society of Washington|volume=55|pages=167–183}}</ref> ''O. ochracea'' is also found throughout North America, South America, and the Caribbean,<ref name=":1">{{Cite journal|last=Lehmann|first=Gerlind U.C.|date=January 2003|title=Review of Biogeography, Host Range and Evolution of Acoustic Hunting in Ormiini (Insecta, Diptera, Tachinidae), Parasitoids of Night-calling Bushcrickets and Crickets (Insecta, Orthoptera, Ensifera)|journal=Zoologischer Anzeiger - A Journal of Comparative Zoology|volume=242|issue=2|pages=107–120|doi=10.1078/0044-5231-00091|s2cid=85839051 |issn=0044-5231}}</ref> though its exact range is not known.


== Life history ==
== Life history ==
''Ormia ochracea'' has the full life cycle of egg, larvae, pupa, and adult. Once a female fly finds a suitable host, she deposits planidia (first [[instar]] larvae) which then quickly burrow into the host.<ref name=":3">{{Cite journal|last=Wineriter|first=S. A.|last2=Walker|first2=T. J.|date=December 1990|title=Rearing phonotactic parasitoid flies [Diptera: Tachinidae, ormiini, ormia spp.]|journal=Entomophaga|volume=35|issue=4|pages=621–632|doi=10.1007/bf02375096|issn=0013-8959}}</ref> The [[Planidium|planidia]] develop within the body of the field cricket host, embedding initially in muscle before migrating into the abdomen.<ref name=":4">{{Cite journal|last=Adamo|first=S.A.|last2=Robert|first2=D.|last3=Hoy|first3=R.R.|date=March 1995|title=Effects of a tachinid parasitoid, Ormia ochracea, on the behaviour and reproduction of its male and female field cricket hosts (Gryllus spp)|journal=Journal of Insect Physiology|volume=41|issue=3|pages=269–277|doi=10.1016/0022-1910(94)00095-x|issn=0022-1910}}</ref> The larvae molt within the host's abdomen and feed primarily on the host's muscle and fat. ''O. ochracea'' larvae typically complete development and emerge after about 7 days, which subsequently kills the host. The larvae pupate and emerge as adult flies approximately 2 weeks after emerging from the host.<ref name=":3" />
''Ormia ochracea'' has the full life cycle of egg, larvae, pupa, and adult. Once a female fly finds a suitable host, she deposits planidia (first [[instar]] larvae) which then quickly burrow into the host.<ref name=":3">{{Cite journal|last1=Wineriter|first1=S. A.|last2=Walker|first2=T. J.|date=December 1990|title=Rearing phonotactic parasitoid flies [Diptera: Tachinidae, ormiini, ormia spp.]|journal=Entomophaga|volume=35|issue=4|pages=621–632|doi=10.1007/bf02375096|s2cid=27347119 |issn=0013-8959}}</ref> The [[Planidium|planidia]] develop within the body of the field cricket host, embedding initially in muscle before migrating into the abdomen.<ref name=":4">{{Cite journal|last1=Adamo|first1=S.A.|last2=Robert|first2=D.|last3=Hoy|first3=R.R.|date=March 1995|title=Effects of a tachinid parasitoid, Ormia ochracea, on the behaviour and reproduction of its male and female field cricket hosts (Gryllus spp)|journal=Journal of Insect Physiology|volume=41|issue=3|pages=269–277|doi=10.1016/0022-1910(94)00095-x|issn=0022-1910|doi-access=free}}</ref> The larvae molt within the host's abdomen and feed primarily on the host's muscle and fat. ''O. ochracea'' larvae typically complete development and emerge after about 7 days, which subsequently kills the host. The larvae pupate and emerge as adult flies approximately 2 weeks after emerging from the host.<ref name=":3" />
[[File:Fall Field Cricket (Gryllus pennsylvanicus) - Kitchener, Ontario 2018-06-28.jpg|left|thumb|field cricket is the common host for ''O. ochracea'']]
[[File:Fall Field Cricket (Gryllus pennsylvanicus) - Kitchener, Ontario 2018-06-28.jpg|left|thumb|field cricket is the common host for ''O. ochracea'']]


== Food resources ==
== Food resources ==
''O. ochracea'' is a [[parasitoid]] known to prey on several species of ''[[Gryllus]]'' field crickets including ''[[Gryllus integer]], Gryllus rubens, Gryllus texensis,'' and ''Gryllus firmus.''<ref name=":1" /> Flies have been observed responding to various cricket songs, but seem to be limited to the family [[Gryllidae]]. The natural host of the fly may vary by location. Larvae of ''O. ochracea'' exhibit highest survival in its natural host and limited survival in other potential host species.<ref>{{Cite journal|last=Thomson|first=Ian R.|last2=Vincent|first2=Crystal M.|last3=Bertram|first3=Susan M.|date=March 2012|title=Success of the Parasitoid FlyOrmia ochracea(Diptera: Tachinidae) on Natural and Unnatural Cricket Hosts|journal=Florida Entomologist|volume=95|issue=1|pages=43–48|doi=10.1653/024.095.0108|issn=0015-4040|doi-access=free}}</ref>
''O. ochracea'' is a [[parasitoid]] known to prey on several species of ''[[Gryllus]]'' field crickets including ''[[Gryllus integer]], Gryllus rubens, Gryllus texensis,'' and ''Gryllus firmus.''<ref name=":1" /> Flies have been observed responding to various cricket songs, but seem to be limited to the family [[Gryllidae]]. The natural host of the fly may vary by location. Larvae of ''O. ochracea'' exhibit highest survival in its natural host and limited survival in other potential host species.<ref>{{Cite journal|last1=Thomson|first1=Ian R.|last2=Vincent|first2=Crystal M.|last3=Bertram|first3=Susan M.|date=March 2012|title=Success of the Parasitoid FlyOrmia ochracea(Diptera: Tachinidae) on Natural and Unnatural Cricket Hosts|journal=Florida Entomologist|volume=95|issue=1|pages=43–48|doi=10.1653/024.095.0108|issn=0015-4040|doi-access=free}}</ref>


=== Host-finding ===
=== Host-finding ===
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==== Learning ====
==== Learning ====
''O. ochracea'' have been shown to adjust their preference for host songs after exposure to different songs in the laboratory. In a 2011 study, flies that were previously exposed to the ''G. lineaticeps'' song chose the ''G. lineaticeps'' song over the ''G. integer'' song, and vice versa.<ref>{{Cite journal|last=Paur|first=Jennifer|last2=Gray|first2=David A.|date=October 2011|title=Individual consistency, learning and memory in a parasitoid fly, Ormia ochracea|journal=Animal Behaviour|volume=82|issue=4|pages=825–830|doi=10.1016/j.anbehav.2011.07.017|issn=0003-3472}}</ref> This preference was very short term. ''O. ochracea'''s flexible learning capabilities may have been critical in expanding its host and geographical range.
''O. ochracea'' have been shown to adjust their preference for host songs after exposure to different songs in the laboratory. In a 2011 study, flies that were previously exposed to the ''G. lineaticeps'' song chose the ''G. lineaticeps'' song over the ''G. integer'' song, and vice versa.<ref>{{Cite journal|last1=Paur|first1=Jennifer|last2=Gray|first2=David A.|date=October 2011|title=Individual consistency, learning and memory in a parasitoid fly, Ormia ochracea|journal=Animal Behaviour|volume=82|issue=4|pages=825–830|doi=10.1016/j.anbehav.2011.07.017|s2cid=18427803 |issn=0003-3472}}</ref> This preference was very short term. ''O. ochracea'''s flexible learning capabilities may have been critical in expanding its host and geographical range.


=== Effects of infestation on host behavior ===
=== Effects of infestation on host behavior ===
''O. ochracea'' infestation has been shown to affect the behavior and reproduction of host field crickets.<ref name=":4" /><ref name=":5">{{Cite journal|last=Beckers|first=Oliver M.|last2=Wagner|first2=William E.|date=April 2013|title=Parasitoid infestation changes female mating preferences|journal=Animal Behaviour|volume=85|issue=4|pages=791–796|doi=10.1016/j.anbehav.2013.01.025|pmc=3857093|pmid=24347669|issn=0003-3472|url=http://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1240&context=bioscifacpub}}</ref> Early in the infestation period, non-reproductive behavior is largely unimpaired because the parasitic larvae do not consume the digestive system or central nervous system of the host.<ref name=":4" /> After the larvae migrate to the host's abdomen, the host's mating, egg-laying, and fighting ability decline, most likely due to tissue damage caused by the larvae.<ref name=":4" /> Additionally, infestation of female crickets alters their mating preferences. ''[[Gryllus lineaticeps]]'' females normally prefer to respond to male songs with intermediate chirp rates over those with slow chirp rates, but females parasitized by ''O. ochracea'' show no preference between chrip rates.<ref name=":5" /> Reduced selectivity in infested female ''G. lineaticeps'' may be adaptive, as a female may be more likely to reproduce before being killed by the parasitoids if they are less selective.
''O. ochracea'' infestation has been shown to affect the behavior and reproduction of host field crickets.<ref name=":4" /><ref name=":5">{{Cite journal|last1=Beckers|first1=Oliver M.|last2=Wagner|first2=William E.|date=April 2013|title=Parasitoid infestation changes female mating preferences|journal=Animal Behaviour|volume=85|issue=4|pages=791–796|doi=10.1016/j.anbehav.2013.01.025|pmc=3857093|pmid=24347669|issn=0003-3472|url=http://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1240&context=bioscifacpub}}</ref> Early in the infestation period, non-reproductive behavior is largely unimpaired because the parasitic larvae do not consume the digestive system or central nervous system of the host.<ref name=":4" /> After the larvae migrate to the host's abdomen, the host's mating, egg-laying, and fighting ability decline, most likely due to tissue damage caused by the larvae.<ref name=":4" /> Additionally, infestation of female crickets alters their mating preferences. ''[[Gryllus lineaticeps]]'' females normally prefer to respond to male songs with intermediate chirp rates over those with slow chirp rates, but females parasitized by ''O. ochracea'' show no preference between chirp rates.<ref name=":5" /> Reduced selectivity in infested female ''G. lineaticeps'' may be adaptive, as a female may be more likely to reproduce before being killed by the parasitoids if they are less selective.


=== Host defenses ===
=== Host defenses ===
Some species of [[Cricket (insect)|cricket]] which are parasitzed by ''O. ochracea'' have evolved methods to avoid infestation. For example, some members of the prey cricket ''[[Teleogryllus oceanicus]]'' have a mutation called flat wing, in which the sound-producing structures of the male forewings are erased.<ref>{{cite journal | last=Pascoal | first=Sonia | last2=Cezard | first2=Timothee | last3=Eik-Nes | first3=Aasta | last4=Gharbi | first4=Karim | last5=Majewska | first5=Jagoda | last6=Payne | first6=Elizabeth | last7=Ritchie | first7=Michael G. | last8=Zuk | first8=Marlene | last9=Bailey | first9=Nathan W. |display-authors=5| title=Rapid Convergent Evolution in Wild Crickets | journal=Current Biology | publisher=Elsevier BV | volume=24 | issue=12 | year=2014 | issn=0960-9822 | doi=10.1016/j.cub.2014.04.053 | pages=1369–1374}}</ref> The flat wing was first observed in 2003 on the [[Hawaii]]an island of [[Kauai]], and was also found on neighbouring [[Oahu]] two years later. Genetic studies of crickets from each island show that the mutations arose from different genomic variations.
Some species of [[Cricket (insect)|cricket]] which are parasitzed by ''O. ochracea'' have evolved methods to avoid infestation. For example, some members of the prey cricket ''[[Teleogryllus oceanicus]]'' have a mutation called flat wing, in which the sound-producing structures of the male forewings are erased.<ref>{{cite journal | last1=Pascoal | first1=Sonia | last2=Cezard | first2=Timothee | last3=Eik-Nes | first3=Aasta | last4=Gharbi | first4=Karim | last5=Majewska | first5=Jagoda | last6=Payne | first6=Elizabeth | last7=Ritchie | first7=Michael G. | last8=Zuk | first8=Marlene | last9=Bailey | first9=Nathan W. |display-authors=5| title=Rapid Convergent Evolution in Wild Crickets | journal=Current Biology | publisher=Elsevier BV | volume=24 | issue=12 | year=2014 | issn=0960-9822 | doi=10.1016/j.cub.2014.04.053 | pages=1369–1374| pmid=24881880 | doi-access=free }}</ref> The flat wing was first observed in 2003 on the [[Hawaii]]an island of [[Kauai]], and was also found on neighbouring [[Oahu]] two years later. Genetic studies of crickets from each island show that the mutations arose from different genomic variations.


== Enemies ==
== Enemies ==
Because field crickets commonly sing at night, ''O. ochracea'' are susceptible to predation by [[bat]]s. Studies have shown that ''O. ochracea'' has evolved an acoustic startle response to bat-like ultrasound, a response very similar to that of female crickets.<ref name=":2">{{Cite journal|last=Rosen|first=M. J.|last2=Levin|first2=E. C.|last3=Hoy|first3=R. R.|date=2009-11-27|title=The cost of assuming the life history of a host: acoustic startle in the parasitoid fly Ormia ochracea|journal=Journal of Experimental Biology|volume=212|issue=24|pages=4056–4064|doi=10.1242/jeb.033183|pmid=19946084|pmc=2784737|issn=0022-0949}}</ref> ''O. ochracea'' has also been shown to demonstrate a sharp response boundary between the frequencies of cricket song and bat ultrasound.<ref name=":2" />
Because field crickets commonly sing at night, ''O. ochracea'' are susceptible to predation by [[bat]]s. Studies have shown that ''O. ochracea'' has evolved an acoustic startle response to bat-like ultrasound, a response very similar to that of female crickets.<ref name=":2">{{Cite journal|last1=Rosen|first1=M. J.|last2=Levin|first2=E. C.|last3=Hoy|first3=R. R.|date=2009-11-27|title=The cost of assuming the life history of a host: acoustic startle in the parasitoid fly Ormia ochracea|journal=Journal of Experimental Biology|volume=212|issue=24|pages=4056–4064|doi=10.1242/jeb.033183|pmid=19946084|pmc=2784737|issn=0022-0949}}</ref> ''O. ochracea'' has also been shown to demonstrate a sharp response boundary between the frequencies of cricket song and bat ultrasound.<ref name=":2" />


== Physiology ==
== Physiology ==


=== Directional hearing ===
=== Directional hearing ===
In order for an animal to localize sound, it must be able to detect minute differences in intensity and time between the arrival of the sound to the ear closer to the source and the ear further from the source. ''O. ochracea'' displays a remarkable ability to localize sound despite the incredibly small distance (450-520μm) between its acoustic sensory organs.<ref name=":6">{{Cite journal|last=Miles|first=R. N.|last2=Robert|first2=D.|last3=Hoy|first3=R. R.|date=December 1995|title=Mechanically coupled ears for directional hearing in the parasitoid fly Ormia ochracea|journal=The Journal of the Acoustical Society of America|volume=98|issue=6|pages=3059–3070|doi=10.1121/1.413830|pmid=8550933|issn=0001-4966}}</ref> Its sound localization ability is facilitated by a cuticular structure which joins its ears, mechanically coupling their motion and magnifying interaural differences by a factor of about 20.<ref name=":6" /> Prior to ''O. ochracea'' no similar mechanism of auditory localization had been described.
In order for an animal to localize sound, it must be able to detect minute differences in intensity and time between the arrival of the sound to the ear closer to the source and the ear further from the source. ''O. ochracea'' displays a remarkable ability to localize sound despite the incredibly small distance (450-520μm) between its acoustic sensory organs.<ref name=":6">{{Cite journal|last1=Miles|first1=R. N.|last2=Robert|first2=D.|last3=Hoy|first3=R. R.|date=December 1995|title=Mechanically coupled ears for directional hearing in the parasitoid fly Ormia ochracea|journal=The Journal of the Acoustical Society of America|volume=98|issue=6|pages=3059–3070|doi=10.1121/1.413830|pmid=8550933|issn=0001-4966}}</ref> Its sound localization ability is facilitated by a cuticular structure which joins its ears, mechanically coupling their motion and magnifying interaural differences by a factor of about 20.<ref name=":6" /> Prior to ''O. ochracea'' no similar mechanism of auditory localization had been described.


== Scientific significance ==
== Scientific significance ==
Several researchers have reported the construction of microphones inspired by the hearing system of ''O. ochracea.'' In 2009, R.N. Miles et al. designed and created a low-noise differential microphone inspired by the unique hearing system of ''O. ochracea'', for use in hearing aids''.'' The design of their microphone diaphragm, which measures 1 x 2&nbsp;mm<sup>2</sup>, is based on the mechanically coupled ears of ''O. ochracea.'' Their microphone was found to have lower noise than commercially available hearing aid microphones while minimizing distance between sensors.<ref>{{Cite book|title=A low-noise differential microphone inspired by the ears of the parasitoid fly Ormia ochracea|last=Miles, R.N. Su, Q. Cui, W. Shetye, M. Degertekin, F.L. Bicen, B. Garcia, C. Jones, S. Hall, N.|date=April 2009|publisher=Acoustical Society of America|oclc=678236270}}</ref> In April 2015, a group from the [[University of Strathclyde]] and the MRC/CSO Institute for Hearing Research (IHR) announced that it had created a microphone based on ''O. ochracea'''s hearing system, and had been awarded a £430,000 grant by the U.K. [[Engineering and Physical Sciences Research Council]] to build and test the hearing aid for three years.<ref>{{cite web | last=Clark| first=Liat | title=Parasitic flies inspire potential revolution in hearing aids | website=WIRED UK | date=2015-04-21 | url=https://www.wired.co.uk/article/parasitic-flies-inspire-hearing-aid | access-date=2021-11-11}}</ref>
Several researchers have reported the construction of microphones inspired by the hearing system of ''O. ochracea.'' In 2009, R.N. Miles et al. designed and created a low-noise differential microphone inspired by the unique hearing system of ''O. ochracea'', for use in hearing aids''.'' The design of their microphone diaphragm, which measures 1 x 2&nbsp;mm<sup>2</sup>, is based on the mechanically coupled ears of ''O. ochracea.'' Their microphone was found to have lower noise than commercially available hearing aid microphones while minimizing distance between sensors.<ref>{{Cite book|title=A low-noise differential microphone inspired by the ears of the parasitoid fly Ormia ochracea|last=Miles, R.N. Su, Q. Cui, W. Shetye, M. Degertekin, F.L. Bicen, B. Garcia, C. Jones, S. Hall, N.|date=April 2009|publisher=Acoustical Society of America|oclc=678236270}}</ref> In April 2015, a group from the [[University of Strathclyde]] and the MRC/CSO Institute for Hearing Research (IHR) announced that it had created a microphone based on ''O. ochracea'''s hearing system, and had been awarded a £430,000 grant by the U.K. [[Engineering and Physical Sciences Research Council]] to build and test the hearing aid for three years.<ref>{{cite magazine | last=Clark| first=Liat | title=Parasitic flies inspire potential revolution in hearing aids | magazine=WIRED UK | date=2015-04-21 | url=https://www.wired.co.uk/article/parasitic-flies-inspire-hearing-aid | access-date=2021-11-11}}</ref>


==References==
==References==
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[[Category:Insects described in 1889]]
[[Category:Insects described in 1889]]
[[Category:Diptera of North America]]
[[Category:Diptera of North America]]
[[Category:Diptera of South America]]

Latest revision as of 12:55, 25 October 2023

Ormia ochracea
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Family: Tachinidae
Subfamily: Tachininae
Tribe: Ormiini
Genus: Ormia
Species:
O. ochracea
Binomial name
Ormia ochracea
(Bigot, 1889)[1]
Synonyms

Ormia ochracea is a small yellow nocturnal fly in the family Tachinidae.[2] It is notable for its parasitism of crickets and its exceptionally acute directional hearing. The female is attracted to the song of the male cricket and deposits larvae on or around him, as was discovered in 1975 by the zoologist William H. Cade.[3]

Ormia ochracea is a model organism in sound localization experiments because of its unique "ears", which are complex structures inside the fly's prothorax near the bases of its front legs. The fly is too small for the time difference of sound arriving at the two ears to be calculated in the usual way, yet it can determine the direction of sound sources with exquisite precision. The tympanic membranes of opposite ears are directly connected mechanically, allowing resolution of nanosecond time differences[4][5] and requiring a new neural coding strategy. Various research groups have designed low-noise differential microphones inspired by O. ochracea’s directionally sensitive hearing system.

Distribution

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Ormia ochracea is native to the southeastern United States, including states such as Texas and Florida.[6] O. ochracea is also found throughout North America, South America, and the Caribbean,[7] though its exact range is not known.

Life history

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Ormia ochracea has the full life cycle of egg, larvae, pupa, and adult. Once a female fly finds a suitable host, she deposits planidia (first instar larvae) which then quickly burrow into the host.[8] The planidia develop within the body of the field cricket host, embedding initially in muscle before migrating into the abdomen.[9] The larvae molt within the host's abdomen and feed primarily on the host's muscle and fat. O. ochracea larvae typically complete development and emerge after about 7 days, which subsequently kills the host. The larvae pupate and emerge as adult flies approximately 2 weeks after emerging from the host.[8]

field cricket is the common host for O. ochracea

Food resources

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O. ochracea is a parasitoid known to prey on several species of Gryllus field crickets including Gryllus integer, Gryllus rubens, Gryllus texensis, and Gryllus firmus.[7] Flies have been observed responding to various cricket songs, but seem to be limited to the family Gryllidae. The natural host of the fly may vary by location. Larvae of O. ochracea exhibit highest survival in its natural host and limited survival in other potential host species.[10]

Host-finding

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In 1975, William H. Cade experimentally demonstrated that Ormia ochracea uses the mating call of the field cricket as a means to locate its host. Cade placed dead crickets on top of speakers playing cricket songs and various control sounds and recorded the amount of time the fly spent on either the control or test speaker. He found that flies spent more time on the speakers playing cricket songs and observed that the flies would always deposit larvae on and around the speaker which was playing cricket songs.[3]

Learning

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O. ochracea have been shown to adjust their preference for host songs after exposure to different songs in the laboratory. In a 2011 study, flies that were previously exposed to the G. lineaticeps song chose the G. lineaticeps song over the G. integer song, and vice versa.[11] This preference was very short term. O. ochracea's flexible learning capabilities may have been critical in expanding its host and geographical range.

Effects of infestation on host behavior

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O. ochracea infestation has been shown to affect the behavior and reproduction of host field crickets.[9][12] Early in the infestation period, non-reproductive behavior is largely unimpaired because the parasitic larvae do not consume the digestive system or central nervous system of the host.[9] After the larvae migrate to the host's abdomen, the host's mating, egg-laying, and fighting ability decline, most likely due to tissue damage caused by the larvae.[9] Additionally, infestation of female crickets alters their mating preferences. Gryllus lineaticeps females normally prefer to respond to male songs with intermediate chirp rates over those with slow chirp rates, but females parasitized by O. ochracea show no preference between chirp rates.[12] Reduced selectivity in infested female G. lineaticeps may be adaptive, as a female may be more likely to reproduce before being killed by the parasitoids if they are less selective.

Host defenses

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Some species of cricket which are parasitzed by O. ochracea have evolved methods to avoid infestation. For example, some members of the prey cricket Teleogryllus oceanicus have a mutation called flat wing, in which the sound-producing structures of the male forewings are erased.[13] The flat wing was first observed in 2003 on the Hawaiian island of Kauai, and was also found on neighbouring Oahu two years later. Genetic studies of crickets from each island show that the mutations arose from different genomic variations.

Enemies

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Because field crickets commonly sing at night, O. ochracea are susceptible to predation by bats. Studies have shown that O. ochracea has evolved an acoustic startle response to bat-like ultrasound, a response very similar to that of female crickets.[14] O. ochracea has also been shown to demonstrate a sharp response boundary between the frequencies of cricket song and bat ultrasound.[14]

Physiology

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Directional hearing

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In order for an animal to localize sound, it must be able to detect minute differences in intensity and time between the arrival of the sound to the ear closer to the source and the ear further from the source. O. ochracea displays a remarkable ability to localize sound despite the incredibly small distance (450-520μm) between its acoustic sensory organs.[15] Its sound localization ability is facilitated by a cuticular structure which joins its ears, mechanically coupling their motion and magnifying interaural differences by a factor of about 20.[15] Prior to O. ochracea no similar mechanism of auditory localization had been described.

Scientific significance

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Several researchers have reported the construction of microphones inspired by the hearing system of O. ochracea. In 2009, R.N. Miles et al. designed and created a low-noise differential microphone inspired by the unique hearing system of O. ochracea, for use in hearing aids. The design of their microphone diaphragm, which measures 1 x 2 mm2, is based on the mechanically coupled ears of O. ochracea. Their microphone was found to have lower noise than commercially available hearing aid microphones while minimizing distance between sensors.[16] In April 2015, a group from the University of Strathclyde and the MRC/CSO Institute for Hearing Research (IHR) announced that it had created a microphone based on O. ochracea's hearing system, and had been awarded a £430,000 grant by the U.K. Engineering and Physical Sciences Research Council to build and test the hearing aid for three years.[17]

References

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  1. ^ a b Bigot, J. M. F. (1889). "Dipteres nouveaux ou peu connus. 34e partie, XLII: Diagnoses de nouvelles especes". Annales de la Société Entomologique de France. 8 (6): 253–270.
  2. ^ O’Hara, James E.; Shannon, J. Henderson; D. Monty, Wood (5 March 2020). "World Checklist of the Tachinidae" (PDF). Tachinidae Resources. Retrieved 28 February 2022.
  3. ^ a b Cade, W. (1975-12-26). "Acoustically Orienting Parasitoids: Fly Phonotaxis to Cricket Song". Science. 190 (4221). American Association for the Advancement of Science (AAAS): 1312–1313. doi:10.1126/science.190.4221.1312. ISSN 0036-8075. S2CID 85233362.
  4. ^ Miles, R. N.; Robert, D.; Hoy, R. R. (1995). "Mechanically coupled ears for directional hearing in the parasitoid fly Ormia ochracea". The Journal of the Acoustical Society of America. 98 (6). Acoustical Society of America (ASA): 3059–3070. doi:10.1121/1.413830. ISSN 0001-4966. PMID 8550933.
  5. ^ Robert, D.; Miles, R.N.; Hoy, R.R. (1996). "Directional hearing by mechanical coupling in the parasitoid fly Ormia ochracea". Journal of Comparative Physiology A. 179 (1). Springer Science and Business Media LLC: 29–44. doi:10.1007/bf00193432. ISSN 0340-7594. PMID 8965258. S2CID 21452506.
  6. ^ Sabrosky, C.W. (1953). "Taxonomy and host relations of the tribe Ormiini in the Western Hemisphere (Diptera, Larvaevoridae)". Proceedings of the Entomological Society of Washington. 55: 167–183.
  7. ^ a b Lehmann, Gerlind U.C. (January 2003). "Review of Biogeography, Host Range and Evolution of Acoustic Hunting in Ormiini (Insecta, Diptera, Tachinidae), Parasitoids of Night-calling Bushcrickets and Crickets (Insecta, Orthoptera, Ensifera)". Zoologischer Anzeiger - A Journal of Comparative Zoology. 242 (2): 107–120. doi:10.1078/0044-5231-00091. ISSN 0044-5231. S2CID 85839051.
  8. ^ a b Wineriter, S. A.; Walker, T. J. (December 1990). "Rearing phonotactic parasitoid flies [Diptera: Tachinidae, ormiini, ormia spp.]". Entomophaga. 35 (4): 621–632. doi:10.1007/bf02375096. ISSN 0013-8959. S2CID 27347119.
  9. ^ a b c d Adamo, S.A.; Robert, D.; Hoy, R.R. (March 1995). "Effects of a tachinid parasitoid, Ormia ochracea, on the behaviour and reproduction of its male and female field cricket hosts (Gryllus spp)". Journal of Insect Physiology. 41 (3): 269–277. doi:10.1016/0022-1910(94)00095-x. ISSN 0022-1910.
  10. ^ Thomson, Ian R.; Vincent, Crystal M.; Bertram, Susan M. (March 2012). "Success of the Parasitoid FlyOrmia ochracea(Diptera: Tachinidae) on Natural and Unnatural Cricket Hosts". Florida Entomologist. 95 (1): 43–48. doi:10.1653/024.095.0108. ISSN 0015-4040.
  11. ^ Paur, Jennifer; Gray, David A. (October 2011). "Individual consistency, learning and memory in a parasitoid fly, Ormia ochracea". Animal Behaviour. 82 (4): 825–830. doi:10.1016/j.anbehav.2011.07.017. ISSN 0003-3472. S2CID 18427803.
  12. ^ a b Beckers, Oliver M.; Wagner, William E. (April 2013). "Parasitoid infestation changes female mating preferences". Animal Behaviour. 85 (4): 791–796. doi:10.1016/j.anbehav.2013.01.025. ISSN 0003-3472. PMC 3857093. PMID 24347669.
  13. ^ Pascoal, Sonia; Cezard, Timothee; Eik-Nes, Aasta; Gharbi, Karim; Majewska, Jagoda; et al. (2014). "Rapid Convergent Evolution in Wild Crickets". Current Biology. 24 (12). Elsevier BV: 1369–1374. doi:10.1016/j.cub.2014.04.053. ISSN 0960-9822. PMID 24881880.
  14. ^ a b Rosen, M. J.; Levin, E. C.; Hoy, R. R. (2009-11-27). "The cost of assuming the life history of a host: acoustic startle in the parasitoid fly Ormia ochracea". Journal of Experimental Biology. 212 (24): 4056–4064. doi:10.1242/jeb.033183. ISSN 0022-0949. PMC 2784737. PMID 19946084.
  15. ^ a b Miles, R. N.; Robert, D.; Hoy, R. R. (December 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. doi:10.1121/1.413830. ISSN 0001-4966. PMID 8550933.
  16. ^ Miles, R.N. Su, Q. Cui, W. Shetye, M. Degertekin, F.L. Bicen, B. Garcia, C. Jones, S. Hall, N. (April 2009). A low-noise differential microphone inspired by the ears of the parasitoid fly Ormia ochracea. Acoustical Society of America. OCLC 678236270.{{cite book}}: CS1 maint: multiple names: authors list (link)
  17. ^ Clark, Liat (2015-04-21). "Parasitic flies inspire potential revolution in hearing aids". WIRED UK. Retrieved 2021-11-11.
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