Thelytoky
Guides
Aprostocetus
Aprostocetus is a large genus of parasitoid wasps in the family Eulophidae, established by Westwood in 1833. With approximately 800 described species, it has a global distribution across all major zoogeographic regions. Species are primarily ectoparasitoids or endoparasitoids of insects inhabiting plant galls, including gall midges (Cecidomyiidae), gall wasps (Cynipoidea), and other hosts spanning Hemiptera, Coleoptera, Lepidoptera, Blattodea, Orthoptera, and Neuroptera. Some species are phytophagous, inducing galls directly on plants. The genus is economically significant as a source of biological control agents for invasive pests.
Aridelus
Aridelus is a genus of braconid wasps in the subfamily Euphorinae, comprising approximately 40 described species with cosmopolitan distribution but greatest diversity in tropical regions. These wasps are solitary koinobiont endoparasitoids of heteropteran bugs, primarily targeting nymphs and adults of stink bugs (Pentatomidae) and related families. Several species have been investigated for biological control potential against agricultural pests, particularly Nezara viridula.
Balcha indica
Balcha indica is a solitary ectoparasitoid wasp in the family Eupelmidae, native to Asia and accidentally introduced to North America. It attacks larvae, prepupae, and pupae of the emerald ash borer (Agrilus planipennis), an invasive wood-boring beetle threatening ash trees. Under laboratory conditions, the species exhibits extended adult longevity (mean 59 days, maximum 117 days) and reproduces via thelytokous parthenogenesis. Development from egg to adult takes approximately 83 days at 25°C, suggesting one to two generations per year in temperate North American regions. The species has been recovered from field surveys in Michigan, Pennsylvania, Ohio, Ontario, Virginia, and West Virginia, where it functions as an adventitious biological control agent of this invasive pest.
Ceratina dallatorreana
Dalla Torre's ceratina
Ceratina dallatorreana is a small carpenter bee in the family Apidae, first described by Friese in 1896. The species exhibits a rare reproductive strategy among bees: it reproduces exclusively by parthenogenesis and never produces males. This bee has been documented across three continents—Africa, Europe and Northern Asia (excluding China), and North America—making it one of the more broadly distributed Ceratina species. Its wide geographic range and unique reproductive biology distinguish it from most other small carpenter bees.
Ceroplastes sinensis
Chinese Wax Scale
Ceroplastes sinensis is a cosmopolitan scale insect pest of commercial citrus and other woody plants. Native to Central or South America based on cladistic analysis, it has established populations across temperate regions including Australia, New Zealand, Mediterranean Europe, North America, and parts of Asia. The species produces a distinctive waxy covering and completes a single annual generation in temperate climates. It has become a significant agricultural pest, particularly in citrus groves, prompting development of integrated pest management sampling protocols and biological control efforts.
Coccus viridis
green scale, green coffee scale, coffee green scale
Coccus viridis is a soft scale insect (Hemiptera: Coccidae) and major agricultural pest with a wide tropical and subtropical distribution. It reproduces parthenogenetically via thelytoky, with females producing 50-600 eggs that hatch within minutes to hours beneath the mother's body. The species completes its life cycle in 47-51 days at 25°C, with three nymphal instars before adulthood; first-instar nymphs actively disperse as 'crawlers' while older instars and adults become sessile. It is polyphagous, feeding on phloem sap of over 200 plant genera in 72 families, with primary economic damage to coffee, guava, and cacao crops.
Cyphomyrmex
fungus-growing ants
Cyphomyrmex is a genus of small, drab-colored fungus-growing ants in the tribe Attini, found primarily in the Neotropics. These ants cultivate fungi in the tribe Leucocoprineae as their primary food source, with most species growing fungal nodules rather than full mycelial gardens. Colonies are typically monogynous and small, rarely exceeding 500 workers. The genus is divided into two species complexes: the strigatus complex (South America only) and the rimosus complex (southern North America to South America). Cyphomyrmex represents a basal lineage among attine ants and serves as sister group to Mycetophylax.
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Desmonomata is an infraorder of oribatid mites (beetle mites) within the suborder Oribatida. The group is notable for its exceptionally high incidence of thelytokous parthenogenesis: four of seven families are completely female-producing, and a fifth is nearly so. This reproductive mode appears obligate and ancient, with speciation and evolutionary radiation having occurred without biparental reproduction.
Diplolepis
gall wasps, rose gall wasps
Diplolepis is a genus of gall-inducing wasps in the family Diplolepididae, most commonly associated with roses (Rosa spp.). These small wasps are notable for inducing complex, often conspicuous galls on their host plants through chemical manipulation of plant tissues. The genus includes both sexual and parthenogenetic species, with some like D. rosae reproducing primarily by thelytokous parthenogenesis. Diplolepis species exhibit intricate life cycles involving alternation between sexual and agamic generations, and their galls support diverse communities of parasitoids and inquilines.
Diplolepis rosae
Mossy Rose Gall Wasp, Rose Bedeguar Gall Wasp, Robin's Pincushion Gall Wasp
A gall-inducing wasp in the family Diplolepididae that forms distinctive mossy, pincushion-like galls on wild roses, primarily Rosa canina and Rosa arvensis. The species is notable for its predominantly parthenogenetic reproduction, with fewer than 5% males in most populations. Females lay up to 60 eggs in developing rose buds, inducing chemically-driven plant tissue distortion that creates the characteristic bedeguar gall. The wasp has been introduced to North America alongside cultivated roses.
Disholcaspis canescens
Round Honeydew Gall Wasp
Disholcaspis canescens is a species of gall wasp in the family Cynipidae, known for inducing distinctive galls on oak trees. Like other members of the genus Disholcaspis, this species produces round, detachable galls on oak twigs. The genus is notable for having eliminated males from the life cycle in all known species, with females reproducing parthenogenetically and producing only female offspring. The specific epithet 'canescens' refers to a grayish or hoary appearance, though the precise application to this species' galls or wasps is not detailed in available sources.
Disholcaspis prehensa
clasping twig gall wasp
Disholcaspis prehensa is a gall-inducing cynipid wasp native to California that induces distinctive galls on scrub oak and leather oak. The species exhibits heterogony, alternating between sexual and asexual generations that produce two morphologically different gall types. The asexual generation forms conspicuous mushroom-shaped twig galls with clasping bases, while the sexual generation produces small, hidden bud galls.
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formosa
Greenhouse Whitefly Parasitoid Wasp
Encarsia formosa is a minute chalcidoid wasp in the family Aphelinidae, renowned as one of the first biological control agents used commercially for greenhouse pest management beginning in the 1920s. Females are approximately 0.6 mm in length and exhibit a distinctive black body with yellow abdomen and opalescent wings. The species reproduces asexually via thelytoky induced by Wolbachia bacterial infection; males are produced but are incapable of inseminating females. This parasitoid attacks at least 15 whitefly species, with primary hosts including Trialeurodes vaporariorum (greenhouse whitefly) and Bemisia tabaci (silverleaf whitefly).
Entedoninae
Entedoninae is a subfamily of chalcid wasps in the family Eulophidae comprising over 90 genera. Members are primarily parasitoids with documented associations across diverse host groups including gall-inducing insects, leafminers, sawflies, thrips, and ants. The subfamily exhibits considerable morphological and ecological diversity, with species found in tropical to temperate habitats worldwide.
parasitoidbiological-controlgall-associatedleafminer-parasitoidChalcidoideaHymenopteraEulophidaeEntedoninaetropical-forestagricultural-pest-controlsawfly-parasitoidthrips-parasitoidant-associatedrice-pestCecidomyiidaeTenthredinidaePontaniaClosterocerusCeranisusAfronymphaAmetallonChrysonotomyiaOmphale-genus-groupcanopyunderstoreyecological-plasticityBangladeshPanamaAfricaNew-ZealandEuropeSenegalSouth-AfricaDemocratic-Republic-of-the-CongoUgandaendoparasitoidlarval-parasitoidegg-parasitoidhost-specificityforest-habitatdegraded-habitatmid-altitude-rainforestlowland-rainforesttropical-Americaclassical-biological-controlaugmentative-biological-controlintegrated-pest-managementrice-hispaDicladispa-armigeragall-waspsawfly-gallleafminerant-symbiosisFormicidaeKhaya-senegalensisOchna-pulchramorphological-taxonomymolecular-taxonomy28S-rDNACOICytBnew-species-descriptiongeneric-revisionhost-rangeparasitoid-complexfood-webecosystem-servicenatural-enemypest-managemententomophagoushyperparasitoidinquilinecurculionidApion-minimumPhyllocolpa-leucostictaChrysocharisEntedonAsecodesPleurotroppopsisAfrotroppopsisZaommomentedonApleurotropisKratoysmaPlatocharisSchizocharisParahorismenusAtullyaTropicharisChrysochuriscruymirabilisoryzamyntoreminpashaicarinatumdussiaelongicaudatamachaeriaeunimaculataclavigergorgonaenseauripunctatagalbinalaeviscutaphenacapsiacecvoraelongatapontaniaeheyeririsbecimilletiaenewbyiFörster-1856Chalcidoidea-phylogenybiodiversityspecies-richnesscryptic-speciesmolecular-diagnosticshost-plant-associationhabitat-specificitycanopy-faunaunderstorey-faunaforest-stratificationtropical-biodiversityAfrotropicalNeotropicalPalaearcticNearcticOrientalAustralasianPacificHolarcticbiogeographytaxonomic-revisionannotated-keycataloguetype-specimenmuseum-collectionslide-mounted-specimensmorphometricsgenitaliawing-venationantennaeclypeusmandiblesmalar-spacescapepedicelflagellumgastervertexsuturecolor-patternpale-femoradark-gastersexual-dimorphismmalefemaleovipositoremergencerearingquarantinefield-collectionforeign-explorationaugmentative-releasepesticide-resistancesustainable-agriculturefood-securityrice-productioncereal-pestvegetable-pestfruit-pestornamental-pestforest-pesturban-pestinvasive-speciesnative-rangeintroduced-rangeestablishmentspreadimpactnon-target-effectshost-specificityrisk-assessmentenvironmental-safetyregulatory-approvalresearch-symposiumproceedingspeer-reviewedZootaxaZooKeysInsect-Systematics-&-EvolutionAustralian-Journal-of-EntomologyFar-Eastern-EntomologistRussian-Entomological-JournalBelgian-Journal-of-EntomologyFlorida-EntomologistPan-Pacific-EntomologistEntomological-NewsProceedings-of-the-Hawaiian-Entomological-SocietyTransactions-of-the-American-Entomological-SocietyJournal-of-Natural-HistoryZoological-Journal-of-the-Linnean-SocietyBioControlJournal-of-Economic-EntomologyEntomologia-Experimentalis-et-ApplicataEnvironmental-EntomologyOecologiaCladisticsMolecular-Phylogenetics-and-EvolutionHeredityCanadian-EntomologistProceedings-of-the-Entomological-Society-of-WashingtonAnnals-of-the-Entomological-Society-of-AmericaBulletin-of-Entomological-ResearchNeotropical-EntomologyJournal-of-Asia-Pacific-EntomologyTurkish-Journal-of-ZoologyPolish-Journal-of-EntomologyNorth-Western-Journal-of-ZoologyEntomologica-AmericanaEntomologische-BerichtenSahlbergiaDugesianaActa-Zoologica-LilloanaFolia-Entomologica-MexicanaRevista-de-la-Sociedad-Entomologica-ArgentinaBoletin-de-la-Asociacion-Espanola-de-EntomologiaEntomologicheskoe-ObozrenieEntomological-ReviewZoological-Institute-RASProceedings-of-the-Zoological-Institute-of-the-Russian-Academy-of-SciencesTaiwan-Agricultural-Research-InstituteCouncil-of-AgricultureUniversity-of-CaliforniaRiversideEntomology-Research-MuseumCalifornia-Department-of-Food-and-AgriculturePierce's-Disease-Control-Programbiological-control-programintegrated-pest-management-programresearch-progress-reportannual-summarytechnical-reportsymposium-proceedingsconference-abstractposter-presentationinvited-papermonographbook-chapteridentification-guideillustrated-keyannotated-cataloguetype-materialvoucher-specimencollection-curationtaxonomysystematicsphylogeneticsmolecular-systematicsDNA-barcodingspecies-complexsibling-speciesspecies-boundariesgeographic-variationpopulation-geneticsgene-flowreproductive-isolationhost-racebiotypestrainlaboratory-rearinginsectarymass-productionreleasemonitoringimpact-assessmentefficacyeconomic-evaluationcost-benefit-analysisfarmer-adoptionextensioneducationpolicyregulationinternational-cooperationcapacity-buildingtrainingworkshopfield-daydemonstrationon-farm-researchparticipatory-researchfarmer-field-schoolsustainable-intensificationagroecologyconservation-biological-controlhabitat-managementfloral-resourcesrefugebuffer-zonelandscape-complexityconnectivityspillovernatural-enemy-diversityfood-web-structuretrophic-interactionsmultitrophicindirect-effectstrait-mediateddensity-dependentinverse-density-dependentfunctional-responsenumerical-responsesearching-efficiencyhandling-tim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messene
Eupelmus messene is a tiny parasitoid wasp in the family Eupelmidae. Females are brachypterous (short-winged) and reproduce via thelytoky, producing only daughters. The species is a specialized idiobiont ectoparasitoid of the gall wasp Aulacidea hieracii, attacking both concealed and exposed larvae and pupae of its host. Females possess a remarkably thin, flexible, and mobile ovipositor capable of drilling through hard substrates, including polystyrene Petri dish walls in laboratory observations.
Gronotoma
Gronotoma is a genus of parasitoid wasps in the family Figitidae (subfamily Eucoilinae). Species within this genus are primarily known as parasitoids of leaf-mining flies (Diptera: Agromyzidae), with documented hosts including Liriomyza trifolii, L. huidobrensis, L. sativae, L. bryoniae, Melanagromyza tomaterae, and Japanagromyza tokunagai. Several species, particularly G. micromorpha, have been extensively studied for their potential as biological control agents in agricultural systems. The genus exhibits thelytokous parthenogenesis in at least some species and shows flexibility in host stage utilization.
Leptocybe
blue gum chalcid wasp, eucalyptus gall wasp
Leptocybe is a monotypic genus of gall wasps in the family Eulophidae, containing the single species L. invasa. Native to Queensland, Australia, this genus has become globally significant as an invasive pest of Eucalyptus plantations. The wasp induces spherical galls on leaf buds and young tissues of multiple Eucalyptus species, causing substantial economic damage to forestry operations worldwide. The genus is characterized by extremely small body size, thelytokous parthenogenetic reproduction, and rapid population expansion in introduced ranges.
Lysiphlebus
Lysiphlebus is a genus of small parasitoid wasps in the family Braconidae, subfamily Aphidiinae. Species in this genus are endoparasitoids of aphids, with females laying eggs inside aphid hosts where larvae develop and eventually emerge from mummified aphid remains. The genus exhibits remarkable reproductive diversity, including both sexual and asexual lineages, with some species capable of facultative sex. Several species, particularly L. testaceipes, are important biological control agents used in integrated pest management programs worldwide.
Micromalthus debilis
Telephone-pole Beetle
Micromalthus debilis is a small beetle native to eastern North America and the sole living representative of the family Micromalthidae, often considered a 'living fossil'. The species exhibits one of the most unusual reproductive strategies in Coleoptera: obligate paedogenesis, where reproduction occurs exclusively through parthenogenetic female larvae without production of fertile adults. Adults are vestigial, sterile, and rarely encountered in nature; they can be artificially induced through heat treatment but cannot reproduce. The species is associated with decaying wood and fungal near-red-rot habitats.
Naupactus leucoloma
white-fringed weevil, white-fringed beetle
Naupactus leucoloma is a broad-nosed weevil native to South America that has become a globally distributed agricultural pest. Adults are flightless, parthenogenetic females outside their native range, with males occurring only rarely in Argentina, Uruguay, and southern Brazil. The species is notable for its extreme polyphagy, having been documented feeding on over 385 plant species. Larval root-feeding causes substantially more economic damage than adult folivory.
Neochrysocharis
Neochrysocharis is a genus of small parasitoid wasps in the family Eulophidae. Species in this genus are endoparasitoids of leafmining insects, particularly agromyzid flies. Some species exhibit host-feeding behavior and possess both thelytokous (asexual) and arrhenotokous (sexual) reproductive strains. Multiple species have been evaluated as biological control agents for invasive leafminer pests in agricultural systems.
Neochrysocharis formosa
Neochrysocharis formosa is a small parasitoid wasp in the family Eulophidae. It is a larval parasitoid of leafminers (Agromyzidae) and has been recorded as a parasitoid of Tuta absoluta (Gelechiidae) and Diprion pini (Diprionidae). The species exhibits host-feeding behavior and is widely used as a biological control agent in agricultural systems. Both thelytokous and arrhenotokous strains exist, with thelytokous strains showing superior biocontrol potential. Sex ratios are typically female-biased.
Otiorhynchus sulcatus
black vine weevil, vine weevil, strawberry weevil, cyclamen weevil
Otiorhynchus sulcatus is a parthenogenetic weevil native to the Palearctic region that has become a globally significant horticultural pest. Adults are flightless, nocturnal, and feed on foliage, producing characteristic notching damage along leaf margins. Larvae develop belowground, feeding on roots, corms, and rhizomes, causing severe plant damage or death. The species reproduces exclusively through thelytokous parthenogenesis; males are absent from all populations. Adults exhibit strong visual and thigmotactic preferences for refuge selection, favoring dark colors (black, blue), tall silhouettes, and basal entrance openings.
Phyllonorycter emberizaepenella
Large Midget
A small leaf-mining moth in the family Gracillariidae with a wingspan of 9–10 mm. It reproduces exclusively by thelytokous parthenogenesis, with no males known to exist. Females nevertheless display diurnal calling behavior and produce sex pheromone compounds, despite the absence of mate attraction. The species is widespread across Europe and has been introduced to North America.
Phytomyza plantaginis
Plantain leafminer
Phytomyza plantaginis is a leaf-mining fly in the family Agromyzidae. It is notable as the second documented case of parthenogenetic reproduction (thelytoky) in its family. Populations in Australia exhibit variation in sex ratio, with some areas having only females and others maintaining bisexual populations with female-biased ratios. The species is associated with Plantago host plants.
Platygastrinae
Platygastrinae is a subfamily of minute parasitoid wasps in the family Platygastridae. Members are characterized by their extremely small size, often under 2 mm, and specialized parasitoid biology targeting gall midges (Cecidomyiidae). The subfamily includes approximately 50+ genera, with Synopeas, Leptacis, and Platygaster being among the most studied. Many species exhibit elongated metasomas, though the functional significance of this trait remains unknown. Several species have been documented as biological control agents for agricultural pests, including the recently confirmed parasitoid of soybean gall midge, Synopeas ruficoxum.
Platythyrea
Platythyrea is a genus of predatory ants in the subfamily Ponerinae and the sole member of the tribe Platythyreini. Workers range from 4–20 mm in size. The genus exhibits remarkable reproductive diversity: most species have gamergates (reproductive female workers), though P. conradti has ergatoid queens instead. Some species with gamergates also produce alate queens. The genus has a worldwide distribution in tropical and subtropical regions.
Platythyrea punctata
Common Silvery Hunter Ant
Platythyrea punctata is a neotropical ant species distinguished by its remarkable reproductive flexibility. Most populations reproduce through thelytokous parthenogenesis, where unmated workers produce female offspring from unfertilized eggs, resulting in colonies with clonal genetic structure. The species also exhibits facultative sexual reproduction, with occasional queens, gamergates (mated reproductive workers), and males appearing in some populations. This combination of alternative reproductive tactics is unique among ants and has made the species a model for studying social evolution, division of labor, and the transition between sexual and asexual reproduction.
Poratia
Poratia is a genus of minute polydesmidan millipedes in the family Pyrgodesmidae, characterized by obligate parthenogenetic reproduction via thelytoky. The genus includes Poratia salvator, a small species (3.5 mm length, 0.5 mm width, 19 body segments, brownish-yellow coloration) originally described from El Salvador and recorded from Brazil. Populations exhibit ecological plasticity, inhabiting both strictly terrestrial and periodically inundated environments. Reproduction is independent of males, with extremely skewed sex ratios (approximately 1 male:140 females) and flavobacteria-mediated parthenogenesis demonstrated at the genus level.
Pulvinaria vitis
Cottony Grape Scale, Cottony Scale
Pulvinaria vitis is a thelytokous cottony scale insect native to Europe, now established across much of the Northern Hemisphere and beyond. It is a univoltine pest of grape and peach, with adult females overwintering on host bark and producing large ovisacs containing thousands of eggs in spring. The species exhibits distinctive discontinuous growth and feeding patterns, with three nymphal instars and limited mobility after the first moult.
Pygostolus
Pygostolus is a genus of braconid wasps in the subfamily Blacinae, comprising approximately eight described species. The genus is best known through studies of P. falcatus, an endoparasitoid of weevils in the genus Sitona. These wasps exhibit thelytokous parthenogenesis and develop as solitary endoparasitoids within the haemocoele of their hosts. The genus has been investigated for biological control applications, particularly against sweetclover weevil (Sitona cylindricollis), though effectiveness is limited by phenological asynchrony and diapause.
Strumigenys membranifera
Bare Mustache Ant
Strumigenys membranifera is a minute trap-jaw ant species native to the Old World tropics that has become a globally distributed tramp species. It is one of only six known Strumigenys species exhibiting thelytokous parthenogenesis, where virgin queens produce diploid female offspring from unfertilized eggs without sperm involvement. Queens retain a functional spermatheca and complete reproductive system despite rarely mating, with males produced only exceptionally. The species has been recorded from numerous regions including Taiwan, Greece (Crete), the Galápagos Islands, and across Asia, reflecting its capacity for human-mediated dispersal and establishment in new environments without mating requirements.
Strumigenys rogeri
Roger's Pygmy Snapping Ant
Strumigenys rogeri is a small dacetine ant species first described by Emery in 1890. It is notable for exhibiting thelytokous parthenogenesis in Taiwan, where virgin queens produce female offspring without mating. This reproductive mode, combined with a short development time of approximately 39 days, may contribute to its success as a tramp ant species. The species belongs to a genus of over 850 species characterized by distinctive trap-jaw mandibles.
Tetrastichinae
Tetrastichinae is one of the largest subfamilies of Eulophidae, containing over 100 genera and nearly 3,000 species of minute chalcid wasps. Members exhibit exceptionally diverse biology: most are parasitoids attacking hosts across 10 insect orders and over 100 families, including nematodes, mites, and spider eggs. Some species are phytophagous (typically as inquilines in galls), gall formers, or inquilines. Endoparasitism predominates over ectoparasitism, with both solitary and gregarious forms known; gregarious species may produce over 2,000 individuals from a single host. Reproduction is often by thelytokous parthenogenesis, though arrhenotoky occurs in some taxa.
Trichadenotecnum castum
common barklouse
Trichadenotecnum castum is a uniparental barklouse species in the family Psocidae, described by Betz in 1983. It is one of three obligately parthenogenetic species derived from the biparental ancestor T. alexanderae. Populations are composed exclusively of females that reproduce via obligate thelytokous parthenogenesis. The species belongs to the T. alexanderae species complex, a group of closely related barklice distinguished primarily by reproductive mode and subtle morphological differences.
Trichadenotecnum merum
common barklouse
Trichadenotecnum merum is a uniparental barklouse species described in 1983 as part of the Trichadenotecnum alexanderae species complex. It reproduces exclusively through obligatory parthenogenesis (thelytoky), with populations composed entirely of females. The species was established through mating tests, life history observations, and morphological analysis across the geographic range of the species complex. As a member of the family Psocidae, it belongs to a group commonly known as common barklice.