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{{Short description|Genus of parasitic insects}}
{{Italic title}}
{{Automatic taxobox
{{Taxobox
| image = Female of Trichogramma dendrolimi on egg of armyworm (Noctuidae), photo was taken by Dr Victor Fursov.jpg
⚫
| name = ''Trichogramma''
| image_caption = Female ''Trichogramma dendrolimi'' on egg of armyworm (Noctuidae)
| regnum = [[Animal]]ia
⚫
| taxon = Trichogramma
| phylum = [[Arthropod]]a
| authority = Westwood, 1833
| classis = [[Insect]]a
| display_parents = 3
| ordo = [[Hymenoptera]]
| subordo = [[Apocrita]]
| superfamilia = [[Chalcidoidea]]
| familia = [[Trichogrammatidae]]
| genus = ''''' Trichogramma '''''
| subdivision_ranks = Species
| subdivision_ranks = Species
| subdivision =
| subdivision = 230+, see text
230+, see text
}}
}}


'''''Trichogramma''''' is a genus of minute [[polyphagous]] wasps that are [[endoparasitoid]]s of insect eggs.<ref name=Flanders1960>{{cite journal | last1 = Flanders | first1 = S | last2 = Quednau | first2 = W | year = 1960 | title = Taxonomy of the genus ''Trichogramma'' (Hymenoptera, Chalcidoidea, Trichogrammatidae) | url = | journal = BioControl | volume = 5 | issue = 4| pages = 285–294 | doi=10.1007/bf02372951}}</ref> ''Trichogramma'' is one of around 80 genera from the family [[Trichogrammatidae]], with over 200 species worldwide.<ref name=consoli>Consoli FL, Parra JRP, Zucchi RA (2010) 'Egg Parasitoids in Agroecosystems with Emphasis on Trichogramma.' (Springer).</ref><ref name=knutson>Knutson A (2005) 'The ''Trichogramma'' Manual: A guide to the use of ''Trichogramma'' for Biological Control with Special Reference to Augmentative Releases for Control of Bollworm and Budworm in Cotton.' (Texas Agricultural Extension Service).</ref><ref name=sumer>{{cite journal | last1 = Sumer | first1 = F | last2 = Tuncbilek | first2 = AS | last3 = Oztemiz | first3 = S | last4 = Pintureau | first4 = B | last5 = Rugman-Jones | first5 = P | last6 = Stouthamer | first6 = R | year = 2009 | title = A molecular key to the common species of ''Trichogramma'' of the Mediterranean region | url =https://cloudfront.escholarship.org/dist/prd/content/qt66b2f9k0/qt66b2f9k0.pdf | journal = BioControl | volume = 54 | issue = 5| pages = 617–624 | doi=10.1007/s10526-009-9219-8}}</ref>
'''''Trichogramma''''' is a genus of minute [[polyphagous]] wasps that are [[endoparasitoid]]s of insect eggs.<ref name=Flanders1960>{{cite journal | last1 = Flanders | first1 = S | last2 = Quednau | first2 = W | year = 1960 | title = Taxonomy of the genus ''Trichogramma'' (Hymenoptera, Chalcidoidea, Trichogrammatidae) | journal = BioControl | volume = 5 | issue = 4| pages = 285–294 | doi=10.1007/bf02372951| s2cid = 34509414 }}</ref> ''Trichogramma'' is one of around 80 genera from the family [[Trichogrammatidae]], with over 200 species worldwide.<ref name=consoli>Consoli FL, Parra JRP, Zucchi RA (2010) 'Egg Parasitoids in Agroecosystems with Emphasis on Trichogramma.' (Springer).</ref><ref name=knutson>Knutson A (2005) 'The ''Trichogramma'' Manual: A guide to the use of ''Trichogramma'' for Biological Control with Special Reference to Augmentative Releases for Control of Bollworm and Budworm in Cotton.' (Texas Agricultural Extension Service).</ref><ref name=sumer>{{cite journal | last1 = Sumer | first1 = F | last2 = Tuncbilek | first2 = AS | last3 = Oztemiz | first3 = S | last4 = Pintureau | first4 = B | last5 = Rugman-Jones | first5 = P | last6 = Stouthamer | first6 = R | year = 2009 | title = A molecular key to the common species of ''Trichogramma'' of the Mediterranean region | url =https://cloudfront.escholarship.org/dist/prd/content/qt66b2f9k0/qt66b2f9k0.pdf | journal = BioControl | volume = 54 | issue = 5| pages = 617–624 | doi=10.1007/s10526-009-9219-8| doi-access = free | bibcode = 2009BioCo..54..617S }}</ref>


Although several groups of egg parasitoids are commonly employed for [[biological control]] throughout the world, ''Trichogramma'' spp. have been the most extensively studied.<ref name=upadhay>Upadhyay RK, Mukerji KG, Chamola BP (2001) 'Biocontrol potential and its Exploitation in Sustainable Agriculture: Insect Pests.' (Kluwer Academic/ Plenum Publishers).</ref> More than a thousand papers have been published on ''Trichogramma'' species, and they are the most used biological control agents in the world.<ref name=knuston>Knutson A (2005) 'The Trichogramma Manual: A guide to the use of Trichogramma for Biological Control with Special Reference to Augmentative Releases for Control of bollworm and Budworm in Cotton.' (Texas Agricultural Extension Service).</ref> ''Trichogramma'' spp. have less than 10,000 neurons, approaching the size limit of how small an insect can be, determined by how few neurons they can fit in their central nervous systems, yet exhibiting a complex behavior to sustain their lives.{{cn|date=April 2019}}
Although several groups of egg parasitoids are commonly employed for [[biological control]] throughout the world, ''Trichogramma'' spp. have been the most extensively studied.<ref name=upadhay>Upadhyay RK, Mukerji KG, Chamola BP (2001) 'Biocontrol potential and its Exploitation in Sustainable Agriculture: Insect Pests.' (Kluwer Academic/ Plenum Publishers).</ref> More than a thousand papers have been published on ''Trichogramma'' species, and they are the most used biological control agents in the world.<ref name=knuston>Knutson A (2005) 'The Trichogramma Manual: A guide to the use of Trichogramma for Biological Control with Special Reference to Augmentative Releases for Control of bollworm and Budworm in Cotton.' (Texas Agricultural Extension Service).</ref>

''Trichogramma'' spp. are also of interest in [[neuroscience]] research, having fewer than 10,000 [[neuron]]s, approaching the theoretical lower limit of the size of an insect brain, yet exhibiting complex behaviors to sustain their lives. <ref>{{cite journal |last1=van der Woude |first1=Emma |last2=Smid |first2=Hans M. |title=Maximized complexity in miniaturized brains: morphology and distribution of octopaminergic, dopaminergic and serotonergic neurons in the parasitic wasp, Trichogramma evanescens |journal=Cell and Tissue Research |date=September 2017 |volume=369 |issue=3 |pages=477–496 |doi=10.1007/s00441-017-2642-8|pmid=28597098 |pmc=5579201 |doi-access=free }}</ref>

== Sensation ==
''Trichogramma'' have highly developed [[chemosensory]] organs due to their need to discriminate host from nonhost in a crowded environment.<ref name=knutson/> Zhang et al. 1979 finds 13 [[sensillum|sensilla]] types on the [[antenna (insect)|antenna]]e, [[eye (insect)|eye]]s, [[mouthparts (insect)|mouthparts]], [[wing (insect)|wing]], [[leg (insect)|leg]], and external [[genitalia (insect)|genitalia]] of ''[[Trichogramma dendrolimi|T. dendrolimi]]''. This is considered to generalize to the entire genus, and there may be more still undiscovered.<ref name="Zang-et-al-2021"/>


== Parasitism ==
== Parasitism ==
To locate host eggs, adult females use chemical and visual signals, such as egg shape and colour.<ref name=knutson/> After she finds a suitable egg, an experienced female attempts to determine if the egg has previously been parasitized, using her ovipositor and antennal drumming (tapping on the egg surface). Females also use antennal drumming to determine the size and quality of the target egg, which determines the number of eggs the female will insert.<ref>{{cite journal |last= Klomp |first=H |last2=Teerink |first2=B.J.|last3=Wei |first3=Chun Ma |date=1979 |title=Discrimination Between Parasitized and Unparasitized Hosts in the Egg Parasite ''Trichogramma embryophagum'' (Hym.=Trichogrammatidae)=a Matter of Learning and Forgetting |journal=Netherlands Journal of Zoology |volume=30 |issue=2 |pages=254–27 |doi=10.1163/002829679X00412 |issn=0028-2960 }}</ref> A single female can parasitize up to 10 host eggs a day.
To locate host eggs, adult females use chemical and visual signals, such as egg shape and colour.<ref name=knutson/> After she finds a suitable egg, an experienced female attempts to determine if the egg has previously been parasitized, using her ovipositor and antennal drumming (tapping on the egg surface). Females also use antennal drumming to determine the size and quality of the target egg, which determines the number of eggs the female will insert.<ref>{{cite journal |last1= Klomp |first1=H |last2=Teerink |first2=B.J.|last3=Wei |first3=Chun Ma |date=1979 |title=Discrimination Between Parasitized and Unparasitized Hosts in the Egg Parasite ''Trichogramma embryophagum'' (Hym.: Trichogrammatidae): a Matter of Learning and Forgetting |journal=Netherlands Journal of Zoology |volume=30 |issue=2 |pages=254–27 |doi=10.1163/002829679X00412 |issn=0028-2960 }}</ref> A single female can parasitize up to 10 host eggs a day.


== Identification ==
== Identification ==
''Trichogramma'' wasps are small and very uniform in structure, which causes difficulty in identifying the separate species.<ref name=nag>{{cite journal | last1 = Nagarkatti | first1 = S | last2 = Nagaraja | first2 = H | year = 1977 | title = Biosystematics of ''Trichogramma'' and ''Trichogrammatoidea'' species | url = | journal = Annual Review of Entomology | volume = 22 | issue = | pages = 157–176 | doi=10.1146/annurev.en.22.010177.001105}}</ref><ref name=thomson>{{cite journal | last1 = Thomson | first1 = LJ | last2 = Rundle | first2 = BJ | last3 = Carew | first3 = ME | last4 = Hoffmann | first4 = AA | year = 2003 | title = Identification and characterization of ''Trichogramma'' species from south-eastern Australia using the internal transcribed spacer 2 (ITS-2) region of the ribosomal gene complex | url = | journal = Entomologia Experimentalis et Applicata | volume = 106 | issue = 3| pages = 235–240 | doi=10.1046/j.1570-7458.2003.00029.x}}</ref> As females are all relatively similar, taxonomists rely upon examination of males to tell the different species apart, using features of their antennae and genitalia.<ref name=nagar>{{cite journal | last1 = Nagarkatti | first1 = S | last2 = Nagaraja | first2 = H | year = 1971 | title = Redescriptions of some known species of ''Trichogramma'' (Hym., Trichogrammatidae), showing the importance of the male genitalia as a diagnostic character | url = | journal = Bulletin of Entomological Research | volume = 61 | issue = | pages = 13–31 | doi=10.1017/s0007485300057412}}</ref><ref name=polaszek>{{cite journal | last1 = Polaszek | first1 = A | last2 = Rugman-Jones | first2 = P | last3 = Stouthamer | first3 = R | last4 = Hernandez-Suarez | first4 = E | last5 = Cabello | first5 = T | last6 = Pino Pérez | first6 = M | year = 2012 | title = Molecular and morphological diagnoses of five species of ''Trichogramma'': biological control agents of ''Chrysodeixis chalcites'' (Lepidoptera: Noctuidae) and ''Tuta absoluta'' (Lepidoptera: Gelechiidae) in the Canary Islands | url = | journal = BioControl | volume = 57 | issue = | pages = 21–35 | doi=10.1007/s10526-011-9361-y}}</ref>
''Trichogramma'' wasps are small and very uniform in structure, which causes difficulty in identifying the separate species.<ref name=nag>{{cite journal | last1 = Nagarkatti | first1 = S | last2 = Nagaraja | first2 = H | year = 1977 | title = Biosystematics of ''Trichogramma'' and ''Trichogrammatoidea'' species | journal = Annual Review of Entomology | volume = 22 | pages = 157–176 | doi=10.1146/annurev.en.22.010177.001105}}</ref><ref name=thomson>{{cite journal | last1 = Thomson | first1 = LJ | last2 = Rundle | first2 = BJ | last3 = Carew | first3 = ME | last4 = Hoffmann | first4 = AA | year = 2003 | title = Identification and characterization of ''Trichogramma'' species from south-eastern Australia using the internal transcribed spacer 2 (ITS-2) region of the ribosomal gene complex | journal = Entomologia Experimentalis et Applicata | volume = 106 | issue = 3| pages = 235–240 | doi=10.1046/j.1570-7458.2003.00029.x| bibcode = 2003EEApp.106..235T | s2cid = 84163027 }}</ref> As females are all relatively similar, taxonomists rely upon examination of males to tell the different species apart, using features of their antennae and genitalia.<ref name=nagar>{{cite journal | last1 = Nagarkatti | first1 = S | last2 = Nagaraja | first2 = H | year = 1971 | title = Redescriptions of some known species of ''Trichogramma'' (Hym., Trichogrammatidae), showing the importance of the male genitalia as a diagnostic character | journal = Bulletin of Entomological Research | volume = 61 | pages = 13–31 | doi=10.1017/s0007485300057412}}</ref><ref name=polaszek>{{cite journal | last1 = Polaszek | first1 = A | last2 = Rugman-Jones | first2 = P | last3 = Stouthamer | first3 = R | last4 = Hernandez-Suarez | first4 = E | last5 = Cabello | first5 = T | last6 = Pino Pérez | first6 = M | year = 2012 | title = Molecular and morphological diagnoses of five species of ''Trichogramma'': biological control agents of ''Chrysodeixis chalcites'' (Lepidoptera: Noctuidae) and ''Tuta absoluta'' (Lepidoptera: Gelechiidae) in the Canary Islands | journal = BioControl | volume = 57 | issue = 1 | pages = 21–35 | doi=10.1007/s10526-011-9361-y| bibcode = 2012BioCo..57...21P | s2cid = 18701180 }}</ref>


The first description of a ''Trichogramma'' species was in North America in 1871, by Charles V. Riley. He described the tiny wasps that emerged from eggs of the viceroy butterfly as ''Trichogramma minutum''.<ref name=knutson/> In [[Taxonomy (biology)|taxonomy]], original specimens are very important, as they are the basis of reference for subsequent descriptions of species. The original specimens, however, were lost. Riley also described a second species in 1879 as ''Trichogramma pretiosum'', but these specimens were also lost. To correct these errors, entomologists returned to the areas where Riley originally found the species and obtained [[neotype]] specimens of ''T. minutum'' and ''T. pretiosum''. These specimens are now preserved properly in the [[United States National Museum]].<ref name=knutson/> Currently, the number of ''Trichogramma'' species is over 200, but as of 1960, only some 40 species of ''Trichogramma'' had been described.<ref>{{Cite web | url=http://www.nhm.ac.uk/research-curation/research/projects/chalcidoids/ | title=Chalcids}}</ref>
The first description of a ''Trichogramma'' species was in North America in 1871, by Charles V. Riley. He described the tiny wasps that emerged from eggs of the viceroy butterfly as ''Trichogramma minutum''.<ref name=knutson/> In [[Taxonomy (biology)|taxonomy]], original specimens are very important, as they are the basis of reference for subsequent descriptions of species. The original specimens, however, were lost. Riley also described a second species in 1879 as ''Trichogramma pretiosum'', but these specimens were also lost. To correct these errors, entomologists returned to the areas where Riley originally found the species and obtained [[neotype]] specimens of ''T. minutum'' and ''T. pretiosum''. These specimens are now preserved properly in the [[United States National Museum]].<ref name=knutson/> Currently, the number of ''Trichogramma'' species is over 200, but as of 1960, only some 40 species of ''Trichogramma'' had been described.<ref>{{Cite web | url=http://www.nhm.ac.uk/research-curation/research/projects/chalcidoids/ | title=Chalcids}}</ref>


==''Wolbachia'' in ''Trichogramma''==
==''Wolbachia'' in ''Trichogramma''==
''[[Wolbachia]]'' is a widespread bacterial genus that infects insects' organs, most commonly the reproductive organs.<ref name = Grenier>{{cite journal | last1 = Grenier | first1 = Simon | year = 1998 | title = Successful horizontal transfer of ''Wolbachia'' symbionts between Trichogramma wasps | journal = Proceedings of the Royal Society of London B: Biological Sciences | volume = 265 | issue = 1404 | pages = 1441–1445 | doi=10.1098/rspb.1998.0455|display-authors=etal| pmc = 1689218 }}</ref> ''Wolbachia'' has been observed to alter the host’s reproductive success upon infection.<ref name="Grenier"/> Through a series of manipulations, ''Wolbachia-''infected hosts transmit this intracellular bacterium to uninfected individuals.<ref name="Grenier"/><ref name=Huigens2004>{{cite journal | last1 = Huigens | first1 = M. E. | year = 2004 | title = Natural interspecific and intraspecific horizontal transfer of parthenogenesis–inducing wolbachia in trichogramma wasps | journal = Proceedings of the Royal Society of London B: Biological Sciences | volume = 271 | issue = 1538 | pages = 509–515 | doi=10.1098/rspb.2003.2640| pmid = 15129961 |display-authors=etal| pmc = 1691627 }}</ref> These manipulations include male killing (increasing ratio of infected females that can reproduce), feminization (males become fertile females), [[parthenogenesis]], and [[cytoplasmic incompatibility]].<ref name=Huigens2004/> Horizontal transfer of parthenogenesis-inducing ''Wolbachia'', which has been observed in ''Trichogramma'' wasps, causes infected females to asexually produce fertile females and nonfunctional males.<ref name=Bourtzis>{{cite journal | last1= Bourtzis | first1 = Kostas | last2= O'Neill | first2= Scott | year = 1998 | title = ''Wolbachia'' Infections and Arthropod Reproduction | journal = BioScience | volume = 48 | issue = 4 | pages = 287–293 | doi=10.2307/1313355| jstor = 1313355 }}</ref> The effects of this include potential speciation of ''Trichogramma'', if ''Wolbachia'' is maintained long enough for genetic divergence to occur and for a new species of asexual wasps to become reproductively isolated.<ref name="Bourtzis"/>
''[[Wolbachia]]'' is a widespread bacterial genus that infects insects' organs, most commonly the reproductive organs.<ref name = Grenier>{{cite journal | last1 = Grenier | first1 = Simon | year = 1998 | title = Successful horizontal transfer of ''Wolbachia'' symbionts between Trichogramma wasps | journal = Proceedings of the Royal Society of London B: Biological Sciences | volume = 265 | issue = 1404 | pages = 1441–1445 | doi=10.1098/rspb.1998.0455|display-authors=etal| pmc = 1689218 }}</ref> ''Wolbachia'' has been observed to alter the host's reproductive success upon infection.<ref name="Grenier"/> Through a series of manipulations, ''Wolbachia-''infected hosts transmit this intracellular bacterium to uninfected individuals.<ref name="Grenier"/><ref name=Huigens2004>{{cite journal | last1 = Huigens | first1 = M. E. | year = 2004 | title = Natural interspecific and intraspecific horizontal transfer of parthenogenesis–inducing wolbachia in trichogramma wasps | journal = Proceedings of the Royal Society of London B: Biological Sciences | volume = 271 | issue = 1538 | pages = 509–515 | doi=10.1098/rspb.2003.2640| pmid = 15129961 |display-authors=etal| pmc = 1691627 }}</ref> These manipulations include male killing (increasing ratio of infected females that can reproduce), feminization (males become fertile females), [[parthenogenesis]], and [[cytoplasmic incompatibility]].<ref name=Huigens2004/> Horizontal transfer of parthenogenesis-inducing ''Wolbachia'', which has been observed in ''Trichogramma'' wasps, causes infected females to asexually produce fertile females and nonfunctional males.<ref name=Bourtzis>{{cite journal | last1= Bourtzis | first1 = Kostas | last2= O'Neill | first2= Scott | year = 1998 | title = ''Wolbachia'' Infections and Arthropod Reproduction | journal = BioScience | volume = 48 | issue = 4 | pages = 287–293 | doi=10.2307/1313355| jstor = 1313355 | doi-access = free }}</ref> The effects of this include potential speciation of ''Trichogramma'', if ''Wolbachia'' is maintained long enough for genetic divergence to occur and for a new species of asexual wasps to become reproductively isolated.<ref name="Bourtzis"/>


Transmission of the bacterium through horizontal transfer has been observed within the same species and among different species of ''Trichogramma'', including ''T. kaykai'', ''T. deion, T. pretiosum,'' and'' T. atopovirilia''; however, limitations to transmission exist.<ref name=Huigens2004/> ''In vitro'' successful horizontal transfer is uncommon within ''Trichogramma'', which suggests that the density of ''Wolbachia'' must be relatively high inside of the hosts' ovaries.<ref name=Huigens2004/> Cytoplasmic incompatibility of the host and bacterium can also be the source of this unsuccessful transfer in-vitro.<ref name=Huigens2004/> These limitations ''in vitro'' suggest that in nature, horizontal transfer by parthenogenesis-inducing ''Wolbachia'' may be a difficult and rare phenomenon. However, when looking at the ''Wolbachia''-host associations, the ''Trichogramma-Wolbachia'' form a monophyletic group based on several ''Wolbachia''-specific genes, which may be explained by horizontal transfer of ''Wolbachia'' between different species.<ref name=Huigens2004/> Therefore, although interspecific horizontal transfer of ''Wolbachia'' is limited ''in vitro'', it is likely to occur quite frequently in nature and is not well understood yet.
Transmission of the bacterium through horizontal transfer has been observed within the same species and among different species of ''Trichogramma'', including ''T. kaykai'', ''T. deion, T. pretiosum,'' and'' T. atopovirilia''; however, limitations to transmission exist.<ref name=Huigens2004/> ''In vitro'' successful horizontal transfer is uncommon within ''Trichogramma'', which suggests that the density of ''Wolbachia'' must be relatively high inside of the hosts' ovaries.<ref name=Huigens2004/> Cytoplasmic incompatibility of the host and bacterium can also be the source of this unsuccessful transfer in-vitro.<ref name=Huigens2004/> These limitations ''in vitro'' suggest that in nature, horizontal transfer by parthenogenesis-inducing ''Wolbachia'' may be a difficult and rare phenomenon. However, when looking at the ''Wolbachia''-host associations, the ''Trichogramma-Wolbachia'' form a monophyletic group based on several ''Wolbachia''-specific genes, which may be explained by horizontal transfer of ''Wolbachia'' between different species.<ref name=Huigens2004/> Therefore, although interspecific horizontal transfer of ''Wolbachia'' is limited ''in vitro'', it is likely to occur quite frequently in nature and is not well understood yet.


The effects of ''Wolbachia'' in ''Trichogramma'' have several evolutionary implications. Commonly, uninfected wasps are unable to breed with infected wasps.<ref name=":0">{{Cite journal|title = Evolutionary consequences of Wolbachia infections|journal = Trends in Genetics|date = 2003-04-01|issn = 0168-9525|pmid = 12683975|pages = 217–223|volume = 19|issue = 4|doi = 10.1016/S0168-9525(03)00024-6|first = Sylvain|last = Charlat|first2 = Gregory D. D.|last2 = Hurst|first3 = Hervé|last3 = Merçot}}</ref> Many generations of reproductive isolation of these different groups may result in speciation.<ref name=":0"/> In addition, some hosts can evolve with a dependency on ''Wolbachia'' for core reproductive functions, such as [[oogenesis]], so that eventually an infection is a requirement for successful reproduction.<ref name=":0" /> Finally, ''Wolbachia'' can influence gender determination in its hosts so that more females are successfully born. This results in a reversal in [[sexual selection]], where females must compete for male mates, which has evolutionary implications as it exposes different phenotypes to natural selection.<ref name=":0" />
The effects of ''Wolbachia'' in ''Trichogramma'' have several evolutionary implications. Commonly, uninfected wasps are unable to breed with infected wasps.<ref name=":0">{{Cite journal|title = Evolutionary consequences of Wolbachia infections|journal = Trends in Genetics|date = 2003-04-01|issn = 0168-9525|pmid = 12683975|pages = 217–223|volume = 19|issue = 4|doi = 10.1016/S0168-9525(03)00024-6|first1 = Sylvain|last1 = Charlat|first2 = Gregory D. D.|last2 = Hurst|first3 = Hervé|last3 = Merçot}}</ref> Many generations of reproductive isolation of these different groups may result in speciation.<ref name=":0"/> In addition, some hosts can evolve with a dependency on ''Wolbachia'' for core reproductive functions, such as [[oogenesis]], so that eventually an infection is a requirement for successful reproduction.<ref name=":0" /> Finally, ''Wolbachia'' can influence gender determination in its hosts so that more females are successfully born. This results in a reversal in [[sexual selection]], where females must compete for male mates, which has evolutionary implications as it exposes different phenotypes to natural selection.<ref name=":0" />


== Biological control ==
== Biological control ==
''Trichogramma'' spp. have been used for control of lepidopteran pests for many years. They can be considered the ''Drosophila'' of the parasitoid world, as they have been used for inundative releases and much understanding today comes from experiments with these wasps.<ref name=smith>Smith SM (1996) Biological control with Trichogramma: advances, successes, and potential of their use. In 'Annual Review of Entomology' pp. 375-406.</ref><ref>BURGIO G., MAINI S., 1995.- Control of European corn borer in sweet corn by ''Trichogramma brassicae'' Bezd. (Hym., Trichogrammatidae).- Journal of Applied Entomology, 119 (1): 83-87.</ref>
''Trichogramma'' spp. have been used for control of [[lepidoptera]]n pests for many years. They can be considered the ''[[Drosophila]]'' of the parasitoid world, as they have been used for inundative releases and much understanding today comes from experiments with these wasps.<ref name="Smith-1996">{{cite journal | last=Smith | first=Sandy M. | title=Biological Control with ''Trichogramma'': Advances, Successes, and Potential of Their Use | journal=[[Annual Review of Entomology]] | publisher=[[Annual Reviews (publisher)|Annual Reviews]] | volume=41 | issue=1 | year=1996 | issn=0066-4170 | doi=10.1146/annurev.en.41.010196.002111 | pages=375–406 | pmid=15012334 | s2cid=33808315}} {{S2CID|96421784}}.</ref><ref>BURGIO G., MAINI S., 1995.- Control of European corn borer in sweet corn by ''Trichogramma brassicae'' Bezd. (Hym., Trichogrammatidae).- Journal of Applied Entomology, 119 (1): 83-87.</ref>


Entomologists in the early 1900s began to rear ''Trichogramma'' spp. for biological control. ''T. minutum'' is one of the most commonly found species in Europe and was first mass reared in 1926 on eggs of ''[[Sitotroga cerealella]]''.<ref name=flanders>{{cite journal | last1 = Flanders | first1 = SE | year = 1930 | title = Mass Production of Egg Parasites of the Genus ''Trichogramma'' | url = | journal = Hilgardia | volume = 4 | issue = 16| pages = 465–501 | doi=10.3733/hilg.v04n16p465}}</ref> ''T. minutum'' has been investigated as a method of biological control of the ''Choristoneura fumiferana'', a major pest of spruce and fir forests.<ref name="smith2">Smith, S.M.; Hubbes, M.; Carrow, J.R. 1986. [http://onlinelibrary.wiley.com/doi/10.1111/j.1439-0418.1986.tb00830.x/full#references Factors affecting inundative releases of ''Trichogramma minutum'' Ril. against the spruce budworm]. J. Appl. Entomol. 101(1):29–39.</ref>
Entomologists in the early 1900s began to rear ''Trichogramma'' spp. for biological control. ''T. minutum'' is one of the most commonly found species in Europe and was first mass reared in 1926 on eggs of ''[[Sitotroga cerealella]]''.<ref name=flanders>{{cite journal | last1 = Flanders | first1 = SE | year = 1930 | title = Mass Production of Egg Parasites of the Genus ''Trichogramma'' | journal = Hilgardia | volume = 4 | issue = 16| pages = 465–501 | doi=10.3733/hilg.v04n16p465| doi-access = free }}</ref> ''T. minutum'' has been investigated as a method of biological control of the ''Choristoneura fumiferana'', a major pest of spruce and fir forests.<ref name="smith2">Smith, S.M.; Hubbes, M.; Carrow, J.R. 1986. [http://onlinelibrary.wiley.com/doi/10.1111/j.1439-0418.1986.tb00830.x/full#references Factors affecting inundative releases of ''Trichogramma minutum'' Ril. against the spruce budworm]. J. Appl. Entomol. 101(1):29–39.</ref>


Nine species of ''Trichogramma'' are produced commercially in insectaries around the world, with 30 countries releasing them. ''Trichogramma'' wasps are used for control on numerous crops and plants; these include cotton, sugarcane, vegetables, sugarbeets, orchards, and forests.<ref name=hassan>{{cite journal | last1 = Hassan | first1 = SA | year = 1993 | title = The mass rearing and utilization of ''Trichogramma'' to control lepidopterous pests: Achievements and outlook | url = | journal = Pesticide Science | volume = 37 | issue = 4| pages = 387–391 | doi=10.1002/ps.2780370412}}</ref> Some of the pests controlled include cotton bollworm (''[[Helicoverpa armigera]]''), codling moth (''[[Cydia pomonella]]''), lightbrown apple moth (''[[Epiphyas postvittana]]''), and European corn borer (''[[Ostrinia nubilalis]]'').
Nine species of ''Trichogramma'' are produced commercially in insectaries around the world, with 30 countries releasing them. ''Trichogramma'' wasps are used for control on numerous crops and plants; these include cotton, sugarcane, vegetables, sugarbeets, orchards, and forests.<ref name=hassan>{{cite journal | last1 = Hassan | first1 = SA | year = 1993 | title = The mass rearing and utilization of ''Trichogramma'' to control lepidopterous pests: Achievements and outlook | journal = Pesticide Science | volume = 37 | issue = 4| pages = 387–391 | doi=10.1002/ps.2780370412}}</ref> Some of the pests controlled include cotton bollworm (''[[Helicoverpa armigera]]''), codling moth (''[[Cydia pomonella]]''), lightbrown apple moth (''[[Epiphyas postvittana]]''), and European corn borer (''[[Ostrinia nubilalis]]'').


''Trichogramma'' species vary in their host specificity. This can lead to nontarget hosts being parasitized. This, in turn, can cause problems by reducing the amount of parasitism of the target host, and depending on the rate of parasitism, nontarget effects could be significant on nontarget host populations. Research is being done on the use of ''Trichogramma'' wasps to control populations of spruce bud moth (''[[Zeiraphera canadensis]])'', which damages white spruce trees.<ref>Turgeon, Jean J. [http://pubs.cif-ifc.org/doi/pdf/10.5558/tfc68614-5 "Status of research on the development of management tactics and strategies for the spruce bud moth in white spruce plantations"]. ''The Forestry Chronicle''. '''68''' (5): 614–622. [[Digital object identifier|doi]]:[https://doi.org/10.5558%2Ftfc68614-5 10.5558/tfc68614-5].</ref>
''Trichogramma'' species vary in their host specificity. This can lead to nontarget hosts being parasitized. This, in turn, can cause problems by reducing the amount of parasitism of the target host, and depending on the rate of parasitism, nontarget effects could be significant on nontarget host populations. Research is being done on the use of ''Trichogramma'' wasps to control populations of spruce bud moth (''[[Zeiraphera canadensis]])'', which damages white spruce trees.<ref>Turgeon, Jean J. [http://pubs.cif-ifc.org/doi/pdf/10.5558/tfc68614-5 "Status of research on the development of management tactics and strategies for the spruce bud moth in white spruce plantations"]. ''The Forestry Chronicle''. '''68''' (5): 614–622. [[Digital object identifier|doi]]:[https://doi.org/10.5558%2Ftfc68614-5 10.5558/tfc68614-5].</ref>

''Trichogramma'' began to be seriously used in the 1990s in China. Since then some applications have fallen out of use due to the rise of [[Bt crop|''Bt'' crops]] because ''Bt'' is also toxic to the parasitoid. Future expansion of ''Bt'' in China is expected, and this threatens some uses of ''Trichogramma'', however for some crops/pests it remains the better option and so is expected to continue instead of expanded ''Bt'' in those applications. ''Trichogramma'' will be especially necessary for [[pesticide resistance management|resistance management]] if [[Bt maize|''Bt'' maize/''Bt'' corn]] is widely adopted.<ref name="Zang-et-al-2021">{{cite journal | last1=Zang | first1=Lian-Sheng | last2=Wang | first2=Su | last3=Zhang | first3=Fan | last4=Desneux | first4=Nicolas | title=Biological Control with Trichogramma in China: History, Present Status, and Perspectives | journal=[[Annual Review of Entomology]] | publisher=[[Annual Reviews (publisher)|Annual Reviews]] | volume=66 | issue=1 | date=2021-01-07 | issn=0066-4170 | doi=10.1146/annurev-ento-060120-091620 | pages=463–484| pmid=32976724 | s2cid=221938883 }}</ref>

In 2021 the [[National Trust]] in England embarked on a trial of using ''[[Trichogramma evanescens]]'', which parasitises clothes moth eggs, in conjunction with [[pheromone]]s to control [[common clothes moth]]s, which cause serious damage to carpets, furniture, clothing and other wool and silk objects in historic buildings.<ref>{{cite web | title=Conservation at Blickling Hall, Norfolk | website=National Trust |year=2022 | url=https://www.nationaltrust.org.uk/visit/norfolk/blickling-estate/our-work-in-the-house-at-blickling }}</ref> The trial was abandoned in 2023; while the microwasps performed well at reducing moth populations in combination with pheromones, they were no better than pheromones alone. It was suspected that the high interiors may not have been suitable and the Trust may continue to use the wasps in smaller stores or where they can be sited close to a known infestation source.<ref>{{cite press release| title='Surprising' clothes moths slump of nearly 40% across National Trust houses, possibly helped by record heat and drought, charity's annual insect pests report finds| publisher=National Trust | date=13 February 2023 | url=https://www.nationaltrust.org.uk/services/media/pests-in-properties-report-2023}}</ref><ref>{{Cite web|url=https://www.webwire.com/ViewPressRel.asp?aId=300600|title=‘Surprising’ clothes moths slump of nearly 40% across National Trust houses, possibly helped by record heat and drought, charity’s annual insect pests report finds|website=WebWire}}</ref>


=== Species used ===
=== Species used ===
Line 48: Line 52:
==''T. pretiosum''==
==''T. pretiosum''==


''T. pretiosum'' is the most widely distributed species in North America.<ref name=knutson/> It is a more generalized parasitoid, able to parasitise a range of different species. It has been the focus of many research studies and has been successfully reared on 18 genera of Lepidoptera. ''T. pretiosum'' was introduced into Australia in the 1970s as part of the Ord River Irrigation Area IPM scheme.<ref name=davies>{{cite journal | last1 = Davies | first1 = AP | last2 = Zalucki | first2 = MP | year = 2008 | title = Collection of ''Trichogramma'' Westwood (Hymenoptera: Trichogrammatidae) from tropical northern Australia: a survey of egg parasitoids for potential pest insect biological control in regions of proposed agricultural expansion | url = | journal = Australian Journal of Entomology | volume = 47 | issue = 2| pages = 160–167 | doi=10.1111/j.1440-6055.2008.00644.x}}</ref><ref name=davieset>{{cite journal | last1 = Davies | first1 = AP | last2 = Pufke | first2 = US | last3 = Zalucki | first3 = MP | year = 2011 | title = Spatio-temporal variation in ''Helicoverpa'' egg parasitism by ''Trichogramma'' in a tropical Bt-transgenic cotton landscape | url = | journal = Agricultural and Forest Entomology | volume = 13 | issue = 3| pages = 247–258 | doi=10.1111/j.1461-9563.2010.00512.x}}</ref>
''T. pretiosum'' is the most widely distributed species in North America.<ref name=knutson/> It is a more generalized parasitoid, able to parasitise a range of different species. It has been the focus of many research studies and has been successfully reared on 18 genera of Lepidoptera. ''T. pretiosum'' was introduced into Australia in the 1970s as part of the Ord River Irrigation Area IPM scheme.<ref name=davies>{{cite journal | last1 = Davies | first1 = AP | last2 = Zalucki | first2 = MP | year = 2008 | title = Collection of ''Trichogramma'' Westwood (Hymenoptera: Trichogrammatidae) from tropical northern Australia: a survey of egg parasitoids for potential pest insect biological control in regions of proposed agricultural expansion | journal = Australian Journal of Entomology | volume = 47 | issue = 2| pages = 160–167 | doi=10.1111/j.1440-6055.2008.00644.x}}</ref><ref name=davieset>{{cite journal | last1 = Davies | first1 = AP | last2 = Pufke | first2 = US | last3 = Zalucki | first3 = MP | year = 2011 | title = Spatio-temporal variation in ''Helicoverpa'' egg parasitism by ''Trichogramma'' in a tropical Bt-transgenic cotton landscape | journal = Agricultural and Forest Entomology | volume = 13 | issue = 3| pages = 247–258 | doi=10.1111/j.1461-9563.2010.00512.x| s2cid = 83477611 }}</ref>


==''T. carverae''==
==''T. carverae''==


''[[Trichogramma carverae]]'' is mainly used for [[light brown apple moth]] and [[codling moth]] control, and is predominately used in orchards.<ref name=llewellyn>Llewellyn R (2002) ''The good bug book: beneficial organisms commercially available in Australia and New Zealand for biological pest control.' (Integrated Pest Management Pty Ltd).</ref> In Australia, ''T. carverae'' is used for biological control of light brown apple moth in vineyards. Though Australia has its own native ''Trichogramma'' species, not much work has been undertaken to use them commercially for biological control within Australia.<ref name=glenn>{{cite journal | last1 = Glenn | first1 = DC | last2 = Hercus | first2 = MJ | last3 = Hoffmann | first3 = AA | year = 1997 | title = Characterizing ''Trichogramma'' (Hymenoptera: Trichogrammatidae) species for biocontrol of light brown apple moth (Lepidoptera: Tortricidae) in grapevines in Australia | url = | journal = Annals of the Entomological Society of America | volume = 90 | issue = 2| pages = 128–137 | doi = 10.1093/aesa/90.2.128 }}</ref>
''[[Trichogramma carverae]]'' is mainly used for [[light brown apple moth]] and [[codling moth]] control, and is predominately used in orchards.<ref name=llewellyn>Llewellyn R (2002) ''The good bug book: beneficial organisms commercially available in Australia and New Zealand for biological pest control.' (Integrated Pest Management Pty Ltd).</ref> In Australia, ''T. carverae'' is used for biological control of light brown apple moth in vineyards. Though Australia has its own native ''Trichogramma'' species, not much work has been undertaken to use them commercially for biological control within Australia.<ref name=glenn>{{cite journal | last1 = Glenn | first1 = DC | last2 = Hercus | first2 = MJ | last3 = Hoffmann | first3 = AA | year = 1997 | title = Characterizing ''Trichogramma'' (Hymenoptera: Trichogrammatidae) species for biocontrol of light brown apple moth (Lepidoptera: Tortricidae) in grapevines in Australia | journal = Annals of the Entomological Society of America | volume = 90 | issue = 2| pages = 128–137 | doi = 10.1093/aesa/90.2.128 | doi-access = free }}</ref>


Light brown apple moth is common throughout Australia and is polyphagous on more than 80 native and introduced species. The larvae cause the most damage, especially to grape berries, as their feeding provides sites for bunch rot to occur.<ref>{{cite journal | last1 = Glenn | first1 = DC | last2 = Hoffmann | first2 = AA | year = 1997 | title = Developing a commercially viable system for biological control of light brown apple moth (Lepidoptera: Tortricidae) in grapes using endemic ''Trichogramma'' (Hymenoptera: Trichogrammatidae) | url = | journal = Journal of Economic Entomology | volume = 90 | issue = 2| pages = 370–382 | doi = 10.1093/jee/90.2.370 }}</ref> Losses in the crops can amount up to $2000/ha in one season. It is very predominant in areas such as the [[Yarra Valley]]. Insecticide use is not a choice method for most growers, who prefer a more natural means of controlling pests. As a result, ''Trichogramma'' wasps were considered a good candidate for biological control, even more so as the moth larvae are difficult to control with insecticide. Moreover, light brown apple moths are relatively vulnerable to egg parasitism, with their eggs being laid in masses of 20-50 on the upper surfaces of basal leaves in grapevines.
Light brown apple moth is common throughout Australia and is polyphagous on more than 80 native and [[introduced species]]. The larvae cause the most damage, especially to grape berries, as their feeding provides sites for bunch rot to occur.<ref>{{cite journal | last1 = Glenn | first1 = DC | last2 = Hoffmann | first2 = AA | year = 1997 | title = Developing a commercially viable system for biological control of light brown apple moth (Lepidoptera: Tortricidae) in grapes using endemic ''Trichogramma'' (Hymenoptera: Trichogrammatidae) | journal = Journal of Economic Entomology | volume = 90 | issue = 2| pages = 370–382 | doi = 10.1093/jee/90.2.370 | doi-access = free }}</ref> Losses in the crops can amount up to $2000/ha in one season. It is very predominant in areas such as the [[Yarra Valley]]. Insecticide use is not a choice method for most growers, who prefer a more natural means of controlling pests. As a result, ''Trichogramma'' wasps were considered a good candidate for biological control, even more so as the moth larvae are difficult to control with insecticide. Moreover, light brown apple moths are relatively vulnerable to egg parasitism, with their eggs being laid in masses of 20–50 on the upper surfaces of basal leaves in grapevines.


==Species==
== Selected species ==
* ''[[Trichogramma aomoriense]]''
[[List of Trichogramma species|List of ''Trichogramma'' species]]
* ''[[Trichogramma atopovirilia]]''
* ''[[Trichogramma brassicae]]''
* ''[[Trichogramma brevicapillum]]''
* ''[[Trichogramma carverae]]''
* ''[[Trichogramma chilonis]]''
* ''[[Trichogramma deion]]''
* ''[[Trichogramma dendrolimi]]''
* ''[[Trichogramma evanescens]]''
* ''[[Trichogramma exiguum]]''
* ''[[Trichogramma falx]]''
* ''[[Trichogramma fuentesi]]''
* ''[[Trichogramma funiculatum]]''
* ''[[Trichogramma japonicum]]''
* ''[[Trichogramma maori]]''
* ''[[Trichogramma minutum]]''
* ''[[Trichogramma nubilale]]''
* ''[[Trichogramma papilonis]]''
* ''[[Trichogramma platneri]]''
* ''[[Trichogramma pretiosum]]''
* ''[[Trichogramma siddiqi]]''
* ''[[Trichogramma thalense]]''
* ''[[Trichogramma valentinei]]''
* ''[[Trichogramma yawarae]]''


== References ==
== References ==
Line 94: Line 75:


[[Category:Trichogrammatidae]]
[[Category:Trichogrammatidae]]
[[Category:Insects acting as insect pest control agents]]
[[Category:Insects used as insect pest control agents]]
[[Category:Biological pest control wasps]]
[[Category:Biological pest control wasps]]
[[Category:Chalcidoidea genera]]
[[Category:Hymenoptera genera]]

Latest revision as of 11:33, 27 November 2024

Trichogramma
Female Trichogramma dendrolimi on egg of armyworm (Noctuidae)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Trichogrammatidae
Subfamily: Trichogrammatinae
Tribe: Trichogrammatini
Genus: Trichogramma
Westwood, 1833
Species

230+, see text

Trichogramma is a genus of minute polyphagous wasps that are endoparasitoids of insect eggs.[1] Trichogramma is one of around 80 genera from the family Trichogrammatidae, with over 200 species worldwide.[2][3][4]

Although several groups of egg parasitoids are commonly employed for biological control throughout the world, Trichogramma spp. have been the most extensively studied.[5] More than a thousand papers have been published on Trichogramma species, and they are the most used biological control agents in the world.[6]

Trichogramma spp. are also of interest in neuroscience research, having fewer than 10,000 neurons, approaching the theoretical lower limit of the size of an insect brain, yet exhibiting complex behaviors to sustain their lives. [7]

Sensation

[edit]

Trichogramma have highly developed chemosensory organs due to their need to discriminate host from nonhost in a crowded environment.[3] Zhang et al. 1979 finds 13 sensilla types on the antennae, eyes, mouthparts, wing, leg, and external genitalia of T. dendrolimi. This is considered to generalize to the entire genus, and there may be more still undiscovered.[8]

Parasitism

[edit]

To locate host eggs, adult females use chemical and visual signals, such as egg shape and colour.[3] After she finds a suitable egg, an experienced female attempts to determine if the egg has previously been parasitized, using her ovipositor and antennal drumming (tapping on the egg surface). Females also use antennal drumming to determine the size and quality of the target egg, which determines the number of eggs the female will insert.[9] A single female can parasitize up to 10 host eggs a day.

Identification

[edit]

Trichogramma wasps are small and very uniform in structure, which causes difficulty in identifying the separate species.[10][11] As females are all relatively similar, taxonomists rely upon examination of males to tell the different species apart, using features of their antennae and genitalia.[12][13]

The first description of a Trichogramma species was in North America in 1871, by Charles V. Riley. He described the tiny wasps that emerged from eggs of the viceroy butterfly as Trichogramma minutum.[3] In taxonomy, original specimens are very important, as they are the basis of reference for subsequent descriptions of species. The original specimens, however, were lost. Riley also described a second species in 1879 as Trichogramma pretiosum, but these specimens were also lost. To correct these errors, entomologists returned to the areas where Riley originally found the species and obtained neotype specimens of T. minutum and T. pretiosum. These specimens are now preserved properly in the United States National Museum.[3] Currently, the number of Trichogramma species is over 200, but as of 1960, only some 40 species of Trichogramma had been described.[14]

Wolbachia in Trichogramma

[edit]

Wolbachia is a widespread bacterial genus that infects insects' organs, most commonly the reproductive organs.[15] Wolbachia has been observed to alter the host's reproductive success upon infection.[15] Through a series of manipulations, Wolbachia-infected hosts transmit this intracellular bacterium to uninfected individuals.[15][16] These manipulations include male killing (increasing ratio of infected females that can reproduce), feminization (males become fertile females), parthenogenesis, and cytoplasmic incompatibility.[16] Horizontal transfer of parthenogenesis-inducing Wolbachia, which has been observed in Trichogramma wasps, causes infected females to asexually produce fertile females and nonfunctional males.[17] The effects of this include potential speciation of Trichogramma, if Wolbachia is maintained long enough for genetic divergence to occur and for a new species of asexual wasps to become reproductively isolated.[17]

Transmission of the bacterium through horizontal transfer has been observed within the same species and among different species of Trichogramma, including T. kaykai, T. deion, T. pretiosum, and T. atopovirilia; however, limitations to transmission exist.[16] In vitro successful horizontal transfer is uncommon within Trichogramma, which suggests that the density of Wolbachia must be relatively high inside of the hosts' ovaries.[16] Cytoplasmic incompatibility of the host and bacterium can also be the source of this unsuccessful transfer in-vitro.[16] These limitations in vitro suggest that in nature, horizontal transfer by parthenogenesis-inducing Wolbachia may be a difficult and rare phenomenon. However, when looking at the Wolbachia-host associations, the Trichogramma-Wolbachia form a monophyletic group based on several Wolbachia-specific genes, which may be explained by horizontal transfer of Wolbachia between different species.[16] Therefore, although interspecific horizontal transfer of Wolbachia is limited in vitro, it is likely to occur quite frequently in nature and is not well understood yet.

The effects of Wolbachia in Trichogramma have several evolutionary implications. Commonly, uninfected wasps are unable to breed with infected wasps.[18] Many generations of reproductive isolation of these different groups may result in speciation.[18] In addition, some hosts can evolve with a dependency on Wolbachia for core reproductive functions, such as oogenesis, so that eventually an infection is a requirement for successful reproduction.[18] Finally, Wolbachia can influence gender determination in its hosts so that more females are successfully born. This results in a reversal in sexual selection, where females must compete for male mates, which has evolutionary implications as it exposes different phenotypes to natural selection.[18]

Biological control

[edit]

Trichogramma spp. have been used for control of lepidopteran pests for many years. They can be considered the Drosophila of the parasitoid world, as they have been used for inundative releases and much understanding today comes from experiments with these wasps.[19][20]

Entomologists in the early 1900s began to rear Trichogramma spp. for biological control. T. minutum is one of the most commonly found species in Europe and was first mass reared in 1926 on eggs of Sitotroga cerealella.[21] T. minutum has been investigated as a method of biological control of the Choristoneura fumiferana, a major pest of spruce and fir forests.[22]

Nine species of Trichogramma are produced commercially in insectaries around the world, with 30 countries releasing them. Trichogramma wasps are used for control on numerous crops and plants; these include cotton, sugarcane, vegetables, sugarbeets, orchards, and forests.[23] Some of the pests controlled include cotton bollworm (Helicoverpa armigera), codling moth (Cydia pomonella), lightbrown apple moth (Epiphyas postvittana), and European corn borer (Ostrinia nubilalis).

Trichogramma species vary in their host specificity. This can lead to nontarget hosts being parasitized. This, in turn, can cause problems by reducing the amount of parasitism of the target host, and depending on the rate of parasitism, nontarget effects could be significant on nontarget host populations. Research is being done on the use of Trichogramma wasps to control populations of spruce bud moth (Zeiraphera canadensis), which damages white spruce trees.[24]

Trichogramma began to be seriously used in the 1990s in China. Since then some applications have fallen out of use due to the rise of Bt crops because Bt is also toxic to the parasitoid. Future expansion of Bt in China is expected, and this threatens some uses of Trichogramma, however for some crops/pests it remains the better option and so is expected to continue instead of expanded Bt in those applications. Trichogramma will be especially necessary for resistance management if Bt maize/Bt corn is widely adopted.[8]

In 2021 the National Trust in England embarked on a trial of using Trichogramma evanescens, which parasitises clothes moth eggs, in conjunction with pheromones to control common clothes moths, which cause serious damage to carpets, furniture, clothing and other wool and silk objects in historic buildings.[25] The trial was abandoned in 2023; while the microwasps performed well at reducing moth populations in combination with pheromones, they were no better than pheromones alone. It was suspected that the high interiors may not have been suitable and the Trust may continue to use the wasps in smaller stores or where they can be sited close to a known infestation source.[26][27]

Species used

[edit]

The most commonly used species for biological control are T. atopovirilia, T. brevicapillum, T. deion, T. exiguum, T. fuentesi, T. minutum, T. nubilale, T. platneri, T. pretiosum, and T. thalense.[3]

T. pretiosum

[edit]

T. pretiosum is the most widely distributed species in North America.[3] It is a more generalized parasitoid, able to parasitise a range of different species. It has been the focus of many research studies and has been successfully reared on 18 genera of Lepidoptera. T. pretiosum was introduced into Australia in the 1970s as part of the Ord River Irrigation Area IPM scheme.[28][29]

T. carverae

[edit]

Trichogramma carverae is mainly used for light brown apple moth and codling moth control, and is predominately used in orchards.[30] In Australia, T. carverae is used for biological control of light brown apple moth in vineyards. Though Australia has its own native Trichogramma species, not much work has been undertaken to use them commercially for biological control within Australia.[31]

Light brown apple moth is common throughout Australia and is polyphagous on more than 80 native and introduced species. The larvae cause the most damage, especially to grape berries, as their feeding provides sites for bunch rot to occur.[32] Losses in the crops can amount up to $2000/ha in one season. It is very predominant in areas such as the Yarra Valley. Insecticide use is not a choice method for most growers, who prefer a more natural means of controlling pests. As a result, Trichogramma wasps were considered a good candidate for biological control, even more so as the moth larvae are difficult to control with insecticide. Moreover, light brown apple moths are relatively vulnerable to egg parasitism, with their eggs being laid in masses of 20–50 on the upper surfaces of basal leaves in grapevines.

Species

[edit]

List of Trichogramma species

References

[edit]
  1. ^ Flanders, S; Quednau, W (1960). "Taxonomy of the genus Trichogramma (Hymenoptera, Chalcidoidea, Trichogrammatidae)". BioControl. 5 (4): 285–294. doi:10.1007/bf02372951. S2CID 34509414.
  2. ^ Consoli FL, Parra JRP, Zucchi RA (2010) 'Egg Parasitoids in Agroecosystems with Emphasis on Trichogramma.' (Springer).
  3. ^ a b c d e f g Knutson A (2005) 'The Trichogramma Manual: A guide to the use of Trichogramma for Biological Control with Special Reference to Augmentative Releases for Control of Bollworm and Budworm in Cotton.' (Texas Agricultural Extension Service).
  4. ^ Sumer, F; Tuncbilek, AS; Oztemiz, S; Pintureau, B; Rugman-Jones, P; Stouthamer, R (2009). "A molecular key to the common species of Trichogramma of the Mediterranean region" (PDF). BioControl. 54 (5): 617–624. Bibcode:2009BioCo..54..617S. doi:10.1007/s10526-009-9219-8.
  5. ^ Upadhyay RK, Mukerji KG, Chamola BP (2001) 'Biocontrol potential and its Exploitation in Sustainable Agriculture: Insect Pests.' (Kluwer Academic/ Plenum Publishers).
  6. ^ Knutson A (2005) 'The Trichogramma Manual: A guide to the use of Trichogramma for Biological Control with Special Reference to Augmentative Releases for Control of bollworm and Budworm in Cotton.' (Texas Agricultural Extension Service).
  7. ^ van der Woude, Emma; Smid, Hans M. (September 2017). "Maximized complexity in miniaturized brains: morphology and distribution of octopaminergic, dopaminergic and serotonergic neurons in the parasitic wasp, Trichogramma evanescens". Cell and Tissue Research. 369 (3): 477–496. doi:10.1007/s00441-017-2642-8. PMC 5579201. PMID 28597098.
  8. ^ a b Zang, Lian-Sheng; Wang, Su; Zhang, Fan; Desneux, Nicolas (2021-01-07). "Biological Control with Trichogramma in China: History, Present Status, and Perspectives". Annual Review of Entomology. 66 (1). Annual Reviews: 463–484. doi:10.1146/annurev-ento-060120-091620. ISSN 0066-4170. PMID 32976724. S2CID 221938883.
  9. ^ Klomp, H; Teerink, B.J.; Wei, Chun Ma (1979). "Discrimination Between Parasitized and Unparasitized Hosts in the Egg Parasite Trichogramma embryophagum (Hym.: Trichogrammatidae): a Matter of Learning and Forgetting". Netherlands Journal of Zoology. 30 (2): 254–27. doi:10.1163/002829679X00412. ISSN 0028-2960.
  10. ^ Nagarkatti, S; Nagaraja, H (1977). "Biosystematics of Trichogramma and Trichogrammatoidea species". Annual Review of Entomology. 22: 157–176. doi:10.1146/annurev.en.22.010177.001105.
  11. ^ Thomson, LJ; Rundle, BJ; Carew, ME; Hoffmann, AA (2003). "Identification and characterization of Trichogramma species from south-eastern Australia using the internal transcribed spacer 2 (ITS-2) region of the ribosomal gene complex". Entomologia Experimentalis et Applicata. 106 (3): 235–240. Bibcode:2003EEApp.106..235T. doi:10.1046/j.1570-7458.2003.00029.x. S2CID 84163027.
  12. ^ Nagarkatti, S; Nagaraja, H (1971). "Redescriptions of some known species of Trichogramma (Hym., Trichogrammatidae), showing the importance of the male genitalia as a diagnostic character". Bulletin of Entomological Research. 61: 13–31. doi:10.1017/s0007485300057412.
  13. ^ Polaszek, A; Rugman-Jones, P; Stouthamer, R; Hernandez-Suarez, E; Cabello, T; Pino Pérez, M (2012). "Molecular and morphological diagnoses of five species of Trichogramma: biological control agents of Chrysodeixis chalcites (Lepidoptera: Noctuidae) and Tuta absoluta (Lepidoptera: Gelechiidae) in the Canary Islands". BioControl. 57 (1): 21–35. Bibcode:2012BioCo..57...21P. doi:10.1007/s10526-011-9361-y. S2CID 18701180.
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