Ixodiphagus

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Ixodiphagus
Ixodiphagus hookeri

Ixodiphagus hookeri

Systematics
Class : Insects (Insecta)
Order : Hymenoptera (Hymenoptera)
Subordination : Waist Wasps (Apocrita)
Superfamily : Wood Wasps (Chalcidoidea)
Family : Encyrtidae
Genre : Ixodiphagus
Scientific name
Ixodiphagus
Howard , 1907
Ixodiphagus hookeri , female habit and enlarged genitals
Ixodiphagus hookeri , male habit and enlarged genitals
Ixodiphagus hookeri , egg-laying on a saturated nymph of the common wooden goat
Adult Ixodiphagus hookeri with dead nymph of the common wooden goat
Loophole in the dead nymph of the common ram

Ixodiphagus is a genus of tick parasitic wasps of the family Encyrtidae . The females lay their eggs in the larvae or nymphs of ticks. The hatching larvae feed ontheir hostas parasitoids . The suitability of various species for biological pest control has been studied in detail since the early 20th century, with the cosmopolitan species Ixodiphagus hookeri receiving by far the greatest attention.

description

The species of the genus Ixodiphagus are small, dark colored wasps, the external appearance of the other Chalcidoidea corresponds, but which have a strong and slightly flattened physique. The color of the head and body is brown, with a brass or purple tinge. The body length is between 0.9 and 1.6 millimeters in females and between 0.6 and 1.3 millimeters in males. The wingspan is well below two millimeters, while the front wings, when fully developed, are about 2.1 to 2.3 times as long as they are wide and translucent to opaque and dark. The wing vein on the edge is dotted. In some species the forewings are regressed, more often in males than females.

The head is flattened at the top and several times as wide as it is long. The front vertex appears straight or curved in profile and is about two fifths to two thirds the width of the head. It is usually trimmed with bristle-like setae that are located in conspicuous depressions. There are large complex eyes on the side and three ocelli on the head . The mandibles have only one tooth and have a blunt end. At the front of the head there are two antennas that are close together . They have a scapus, pedicellus and a funiculus divided into six limbs. The clavus is divided into two or three members, which are important for distinguishing between species and sexes. It happens that the members of the clavus are so clearly demarcated that its basal member acts like a seventh member of the funiculus.

The abdomen consists of eight segments in males and seven segments in females, the first of which is fused with the thorax so that only seven or six segments are visible. The thorax is slightly flattened dorsally and often has noticeable depressions with strong setae. The posterior edge of the pronotum is straight to strongly convex in shape. The mesoscutum and the scutellum are densely covered with strong or bristle-like setae. The gaster is slightly shorter than the thorax. The ovipositor is shorter than the middle tibia , but at least five times as long as the gonostyle and surrounded by broad shields. The gonostyle is sometimes extremely shortened and apparently absent. The legs are mostly brown, sometimes with yellow parts. They are often very strong, especially the anterior and posterior femora . Within a species, the males sometimes have noticeably stronger legs than the females.

The sexes can be distinguished by the structure of the antennae and by the genitals. The females have genitals that are complexly shaped and reshaped into an ovipositor , while those of the males are simply structured.

The genus Ixodiphagus differs from all other genera of the Encyrtidae by the combination of blunt mandibles with a tooth, dotted marginal vein of the forewings, small mesopleuron and short ovipositor between broad shields.

Way of life

All species in the genus Ixodiphagus are parasitoids . They lay their eggs in larvae or nymphs of various types of ticks , which serve as food for their larvae.

While a wide range of hosts has been given for Ixodiphagus hookeri and Ixodiphagus texanus , only a single host or none at all are known of some of the species described in the 21st century . The ticks parasitized by Ixodiphagus come from the following genera:

Almost all ticks parasitized by Ixodiphagus are parasites of mammals , and reports of finds in birds have been rare exceptions. In March 1957, during investigations by researchers from Naval Medical Research Unit Three (NAMRU-3) in Cairo, a tick nymph of the species Hyalomma rufipes was found on a wheatear that was on its way from Africa to Europe. About a month later dozens of wasps of the species Ixodiphagus theilerae left the nymph. In the following three years another four nymphs of Hyalomma rufipes parasitized by Ixodiphagus theilerae were discovered, three on Isabel wheatears and one on a common redstart . Haemaphysalis leporispalustris nymphs infested by Ixodiphagus texanus have been repeatedly found on rattle clamps in Oklahoma . They served as the basis for building a laboratory population of Ixodiphagus texanus . The species Ixodiphagus taiaroaensis , described in New Zealand in 2010 , was found in nymphs of ticks of the species Ixodes eudytides and Ixodes uriae in a breeding colony of Stewart's shear .

In searching for suitable hosts, Ixodiphagus hookeri follows the olfactory perception of carbon dioxide and odorous substances emitted by potential mammalian hosts of their preferred tick nymphs. The wasps can differentiate between suitable and unsuitable mammal species. The droppings of the common trestle were only attractive at a distance of up to an inch. Experiments with odorous substances dissolved in n- hexane at various stages of development of the Amblyomma variegatum tick showed a strong attraction, and dummies of hosts were penetrated with the ovipositor . At close range, the wasps follow visual cues such as size and shape in order to distinguish suitable from unsuitable hosts and to determine the number of eggs to be laid in a host.

After egg-laying in a tick larva, the eggs of Ixodiphagus hookeri are taken into the nymph stage. Only when the infected nymph of the tick takes its blood meal from a host does the development of the larvae begin, which feed on the meal and the entire internal organs of the nymph. The number of eggs laid in a host varies and is dependent on the size of the host. The development of the larva takes weeks to months and depends on the climatic conditions. In the case of Ixodiphagus hookeri in temperate zones, a complex sequence of delayed development and resting phases of the larva ensures that the adult wasps hatch at the time of the maximum tick population. At the end of development, the larvae in the front area of ​​the host align themselves parallel to the longitudinal axis and pupate in this position. The entire body of the host can be filled with pupae.

After development is complete, adult wasps hatch from the pupae and gnaw a hole in the back, at the weakest point, for the excursion into the shell of the tick nymph. Adult wasps can only be found for a short period of three to five weeks, in northern summer between late July and late September. Among the hatched wasps, female wasps are in the majority. Mating takes place immediately after hatching and lasts only seconds, then the female goes in search of a suitable host. Both sexes do not eat anything and have a lifespan of a few days.

distribution

Species of Ixodiphagus occur worldwide (except in Antarctica ). Ixodiphagus hookeri has the widest distribution and has been reported from all continents except Australia and Antarctica. The other species have smaller distribution areas or have only been identified once. Several species have been described for the Indian subcontinent , and the description of six species for Costa Rica suggests numerous other undescribed or Ixodiphagus hookeri species.

Biological pest control

Robert A. Cooley at Rocky Mountain Laboratories , 1938

Ticks are mandatory blood-sucking parasites of vertebrates. Most species infest mammals including farm animals and humans. They are major vectors of a wide variety of pathogens, including those causing Rocky Mountain typhus , tularemia , early summer meningoencephalitis, and Lyme disease . Since their vector property was discovered, there has been great interest in controlling ticks. Ixodiphagus hookeri and Ixodiphagus texanus were examined for the possibility of their use for biological pest control just a few years after their first description .

For the targeted control of ticks, in particular Dermacentor andersoni and Dermacentor variabilis , Ixodiphagus hookeri were bred in large numbers in the Rocky Mountain Laboratories in the 1920s under the direction of Robert A. Cooley . The aim of this breed was to establish self-reproducing populations in nature and thus to eradicate Dermacentor andersoni and other ticks as vectors of zoonoses . The first attempts were very promising, also with regard to the wintering of the parasite.

However, the research did not lead to a successful use in tick control. The failure of such attempts is possibly due to the strong adaptation of the populations to local conditions and to certain hosts, which make the use of imported wasps of other origins fail. Measures such as the use of acaricides appear to be unsuitable because of the poisonous effect on the rest of the fauna and humans, the high costs and the observed development of resistance . Current research is aimed at the use of acaropathogenic fungi and nematodes , but is not yet operational.

Symbiotic or parasitic microorganisms

The presence of bacteria of the Wolbachia pipientis group in the common wood tick has been demonstrated in several studies . These bacteria are widespread in insects , but have also been found in claw carriers and nematodes . Their observed influence ranges from obligatory mutualism in filariae , which are dependent on the presence of Wolbachia for development and reproduction , to commensalism to parasitism , with the prevalence ranging from low to complete, depending on the species. In infected insects, Wolbachia was able to demonstrate the diverse influence on the reproduction of their hosts, such as causing parthenogenesis , higher mortality or feminization of male individuals and cytoplasmic incompatibilities, but also strengthening the immune system. The prevalence of ticks is low and the route of infection has long been unclear.

Ixodiphagus hookeri are almost 100 percent infected with a strain of Wolbachia pipientis , whose close relatives have also been found in other hymenoptera . In a molecular genetic investigation of unfeeded nymphs in a natural French population of the common woodbuck, it was found that nymphs either did not contain any DNA from Ixodiphagus hookeri and Wolbachia pipientis , i.e. were free from parasites, or that both DNAs could be detected. Alleged infections of ticks with Wolbachia found in the past were probably an infestation with the infected larvae of Ixodiphagus hookeri . Nothing is known about the specific effects of the infection with Wolbachia pipiens on Ixodiphagus hookeri .

Among the various genres in ticks microorganisms detected, a possible interaction with their presence for Ixodiphagus is the cause, is one among various rickettsial also Arsenophonus nasoniae . Corresponding research has so far focused on Ixodiphagus hookeri .

Systematics and taxonomy

Internal system

Ixodiphagus texanus was first described by Leland Ossian Howard in 1907 . For this he established the initially monotypical genus Ixodiphagus .

The genus Ixodiphagus contains the following species:

In 1980, John S. Noyes, in his review of the Neotropical genera of the Encyrtidae, mentioned a collection specimen from the Natural History Museum in London, which belongs to an undescribed species from Trinidad .

External system

The genus Ixidophagus was with the synonymisation the genera Australzaomma and Hunterellus the only genus of the tribe Ixodiphagini Howard , 1908. The tribe Ixodiphagini forms with its be sister Encyrtini , Echthroplexiellini , Discodini and Oobiini the subfamily Encyrtinae Walker , 1837. The types of other Triben parasitize Hemiptera (Hemiptera), insect eggs from Schnabelkerfen, butterflies (Lepidoptera), beetles (Coleoptera) and reticulated winged birds (Neuroptera). The subfamily Encyrtinae sale in the family Encyrtidae Walker , 1837, with more than 3,000 species one of the largest of nineteen families of the superfamily chalcids (Chalcidoidea) is. Almost all species of the Encyrtidae are parasites of insects , spiders , mites or ticks, the Ixodiphagini are the only parasites of ticks.

Synonyms

  • Hunterellus Howard , 1908 with the type species Hunterellus hookeri Howard
  • Australzaomma Girault , 1925 with the type species Australzaomma brunnea Girault

In 1908 Howard described another species, Hunterellus hookeri , for which he established the genus Hunterellus . For Hunterellus and Ixodiphagus he established the tribe Ixodiphagini. Robert A. Cooley suspected in 1929 that Ixodiphagus texanus , Hunterellus hookeri and a third species were synonyms. Arthur B. Gahan confirmed the species status for the two species described by Howard in 1934 and also retained the genera. Wladimir A. Trjapitzin synonymized Hunterellus with Ixodiphagus in 1985 .

The Australian species Australzaomma brunnea , described by Alexandre A. Girault in 1925, was placed in the genus Ixodiphagus by John S. Noyes and Mohammad Hayat in 1984 , making the monotypical genus Australzaomma synonymous with Ixodiphagus .

literature

Web links

Commons : Ixodiphagus  - collection of images, videos and audio files

Individual evidence

  1. ↑ a b c d e f g h i j k John S. Noyes: Encyrtidae of Costa Rica (Hymenoptera: Chalcidoidea) 3. Subfamily Encyrtinae: Encyrtini, Echthroplexiellini, Discodini, Oobiini and Ixodiphagini, parasitoids associated with bugs (Hemiptera), insect eggs (Hemiptera, Lepidoptera, Coleoptera, Neuroptera) and ticks (Acari) . Memoirs of the American Entomological Institute Volume 84. Gainesville, FL 2010, ISBN 978-1-887988-28-5 , pp. 649-660.
  2. ↑ a b c M. S. Quaraishi: Morphology of Two Chalcidoid Parasites of Ticks, Hunterellus hookeri Howard, 1908, and Ixodiphagus texanus Howard, 1907 . The American Midland Naturalist, 59, 2, pp. 489-504. doi : 10.2307 / 2422494
  3. ↑ a b c Fanuel A. Demas et al .: Cattle and Amblyomma variegatum Odors Used in Host Habitat and Host Finding by the Tick Parasitoid, Ixodiphagus hookeri . In: Journal of Chemical Ecology 2000, Volume 26, No. 4, pp. 1079-1093, doi : 10.1023 / A: 1005497201074 .
  4. ^ A b c Herbert P. Wood : Notes on the Life History of the Tick Parasite Hunterellus hookeri Howard . In: Journal of Economic Entomology 1911, Volume 4, No. 5, pp. 425-431, digitizedhttp: //vorlage_digitalisat.test/1%3D~GB%3D~IA%3Djournalofeconomi04ento~MDZ%3D%0A~SZ%3Dn489~ double-sided%3D~LT%3D~PUR%3D .
  5. ↑ a b Robert L. Zuparko: Annotated Checklist of California Encyrtidae (Hymenoptera) . Zootaxa 2015, Volume 4017, pp. 1–126, doi : 10.11646 / zootaxa.4017.1.1 .
  6. ↑ a b c d Jana Collatz et al .: A hidden beneficial: biology of the tick-wasp Ixodiphagus hookeri in Germany . In: Journal of Applied Entomology 2011, Volume 135, pp. 351-358, doi : 10.1111 / j.1439-0418.2010.01560.x .
  7. ↑ Renjie Hu, Kerwin E. Hyland and James H Oliver: A review on the use of Ixodiphagus wasps (Hymenoptera: Encyrtidae) as natural enemies for the control of ticks (Acari: Ixodidae) . In: Systematic and Applied Acarology 1998, Volume 3, pp. 19-28.
  8. ^ A b Robert A. Cooley : A Search for Tick Parasites in South Africa . In: Onderstepoort Journal of Veterinary Science and Animal Industry 1934, Volume 3, No. 1, pp. 23-42, digitizedhttp: //vorlage_digitalisat.test/1%3Dhttps%3A%2F%2Frepository.up.ac.za%2Fbitstream%2Fhandle%2F2263%2F48340%2F2cooley1934.pdf%3Bsequence%3D1~GB%3D~IA%3D~MDZ% 3D% 0A ~ SZ% 3D ~ double-sided% 3D ~ LT% 3D ~ PUR% 3D .
  9. ↑ Marcos Antônio Bezerra Santos et al .: Larvae of Ixodiphagus wasps (Hymenoptera: Encyrtidae) in Rhipicephalus sanguineus sensu lato ticks (Acari: Ixodidae) from Brazil . In: Ticks and Tick-borne Diseases 2017, Volume 8, No. 4, pp. 564-566, doi : 10.1016 / j.ttbdis.2017.03.004 .
  10. Jump up ↑ Makram N. Kaiser and Harry Hoogstraal: Hunterellus theileri Fiedler (Encyrtidae, Chalcidoidea) Parasitizing an African Hyalomma Tick on a Migrant Bird in Egypt . In: Journal of Parasitology 1958, Vol. 44, No. 4, p. 392, JSTOR 3274322 .
  11. ↑ Harry Hoogstraal and Makram N. Kaiser: Records of Hunterellus theilerae Fiedler (Encyrtidae, Chalcidoidea) Parasitizing Hyalomma Ticks on Birds Migrating Through Egypt . In: Annals of the Entomological Society of America 1961, Vol. 54, No. 4, pp. 616-617, doi : 10.1093 / aesa / 54.4.616 .
  12. ↑ Jerry L. Bowman, Thomas M. Logan and Jakie A. Hair: Host suitability of Ixodiphagus texanus Howard on five species of hard ticks . In: Journal of Agricultural Entomology 1986, Volume 3, No. 1, pp. 1-9, digitizedhttp: //vorlage_digitalisat.test/1%3Dhttp%3A%2F%2Fscentsoc.org%2FVolumes%2FJAE%2Fv3%2F1%2F00031001.pdf~GB%3D~IA%3D~MDZ%3D%0A~SZ%3D~ double-sided% 3D ~ LT% 3D ~ PUR% 3D .
  13. ^ A b Allen CG Heath and Rachel P. Cane : A new species of Ixodiphagus (Hymenoptera: Chalcidoidea: Encyrtidae) parasitizing seabird ticks in New Zealand . In: New Zealand Journal of Zoology 2010, Volume 37, No. 2, pp. 147-155, doi : 10.1080 / 03014223.2010.482973 .
  14. ↑ Fanuel A. Demas et al .: Visual Evaluation and Recognition of Hosts by the Tick Parasitoid, Ixodiphagus hookeri (Hymenoptera: Encyrtidae) . In: Journal of Insect Behavior 2002, Volume 15, No. 4, pp. 477-494, doi : 10.1023 / A: 1016377132585 .
  15. ↑ Jana Collatz et al .: Being a parasitoid of parasites: host finding in the tick wasp Ixodiphagus hookeri by odours from mammals . In: Entomologia Experimentalis et Applicata 2010, Volume 134, No. 2, pp. 131-137, doi : 10.1111 / j.1570-7458.2009.00943.x .
  16. ↑ Keiji Takasu et al .: Host Recognition by the Tick Parasitoid Ixodiphagus hookeri (Hymenoptera: Encyrtidae) . In: Environmental Entomology 2003, Volume 32, No. 3, pp. 614-617, doi : 10.1603 / 0046-225X-32.3.614 .
  17. ↑ Renjie Hu and Kerwin E. Hyland: Effects of the Feeding Process of Ixodes scapularis (Acari: Ixodidae) on Embryonic Development of its Parasitoid, Ixodiphagus hookeri (Hymenoptera: Encyrtidae) . In: Journal of Medical Entomology 1998, Volume 35, No. 6, pp. 1050-1053, doi : 10.1093 / jmedent / 35.6.1050 .
  18. Jump up ↑ Esther N. Mwangi et al .: Predators, parasitoids and pathogens of ticks: A review . In: Biocontrol Science and Technology 1991, Volume 1, No. 3, pp. 147-156, doi : 10.1080 / 09583159109355195 .
  19. ^ Moses M. Cole: Biological Control of Ticks by the Use of Hymenopterous Parasites. A review . WHO / EBL 43.65. World Health Organization, Geneva 1965, digitizedhttp: //vorlage_digitalisat.test/1%3Dhttp%3A%2F%2Fwww.nhm.ac.uk%2Fresources%2Fresearch-curation%2Fprojects%2Fchalcidoids%2Fpdf_X%2FCole965.pdf~GB%3D~IA%3D~MDZ% 3D% 0A ~ SZ% 3D ~ double-sided% 3D ~ LT% 3D ~ PUR% 3D .
  20. ↑ Wladimir A. Trjapitzin and Enrique Ruíz Cancino: Los encírtidos del género Ixodiphagus Howard (Hymenoptera: Chalcidoidea: Encyrtidae), parasitoides de garrapatas (Acarina: Ixodidae) . In: Biotam 1996, Volume 8, No. 1, pp. 9-20.
  21. ↑ AJ Davis: Bibliography of the Ixodiphagini (Hymenoptera, Chalcidoidea, Encyrtidae), parasites of ticks (Acari, Ixodidae), with notes on their biology . In: Tijdschrift voor Entomologie 1986, Volume 129, No. 6, pp. 181-190, digitizedhttp: //vorlage_digitalisat.test/1%3D~GB%3D~IA%3Dtijdschriftvoore128129198586nede~MDZ%3D%0A~SZ%3Dn511~ double-sided%3D~LT%3D~PUR%3D .
  22. ^ A b M. S. Mani : Studies on Indian parasitic Hymenoptera I. In: Indian Journal of Entomology 1941, Volume 3, No. 1, pp. 25-36, digitizedhttp: //vorlage_digitalisat.test/1%3Dhttp%3A%2F%2Fwww.nhm.ac.uk%2Fresources%2Fresearch-curation%2Fprojects%2Fchalcidoids%2Fpdf_X%2FMani941.pdf~GB%3D~IA%3D~MDZ% 3D% 0A ~ SZ% 3D ~ double-sided% 3D ~ LT% 3D ~ PUR% 3D .
  23. ↑ a b G. Geevarghese : A new species of chalcid (Hymenoptera: Encyrtidae), parasitizing Haemaphysalis bispinosa from Karnataka, India . In: Oriental Insects 1977, Volume 11, No. 1, pp. 49-52, digitizedhttp: //vorlage_digitalisat.test/1%3Dhttp%3A%2F%2Fwww.nhm.ac.uk%2Fresources%2Fresearch-curation%2Fprojects%2Fchalcidoids%2Fpdf_X%2FGeevar977.pdf~GB%3D~IA%3D~MDZ% 3D% 0A ~ SZ% 3D ~ double-sided% 3D ~ LT% 3D ~ PUR% 3D .
  24. ↑ a b Mohammad Hayat and Sarfrazul Islam Kazmi : On some Encyrtidae (Hymenoptera: Chalcidoidea) from Namdapha Tiger Reserve in Arunachal Pradesh, India . In: Zootaxa 2011, Volume 2830, No. 1, doi : 10.11646 / zootaxa.2830.1.1 (not viewed).
  25. ^ A b Mohammad Hayat and Kamalanathan Veenakumari: Description of four new species of brachypterous Encyrtidae (Hymenoptera: Chalcidoidea) from India . In: Zootaxa 2015, Volume 3990, No. 2, pp. 259-271, doi : 10.11646 / zootaxa.3990.2.6 .
  26. ^ A b F. Larrousse , Arthur G. King and SB Wolbach : The Overwintering in Massachusetts of Ixodiphagus caucurtei . In: Science 1928, Volume 67, No. 1735, pp. 351-353, doi : 10.1126 / science.67.1735.351 .
  27. ↑ Fred A. Morton: Quantity Production of Tick Parasites . In: Montana State Board of Entomology (Ed.): Seventh Biennual Report. 1927-1928 . The Tribune, Great Falls, MT 1928, pp. 32-35, digitizedhttp: //vorlage_digitalisat.test/1%3D~GB%3D~IA%3Dbiennialre19271928mont~MDZ%3D%0A~SZ%3Dn34~ double-sided%3D~LT%3D~PUR%3D .
  28. ^ JR Parker and WJ Butler: Tick ​​Parasite Liberation in Montana During 1928 . In: Montana State Board of Entomology (Ed.): Seventh Biennual Report. 1927-1928 . The Tribune, Great Falls, MT 1928, pp. 35-38, digitizedhttp: //vorlage_digitalisat.test/1%3D~GB%3D~IA%3Dbiennialre19271928mont~MDZ%3D%0A~SZ%3Dn37~ double-sided%3D~LT%3D~PUR%3D .
  29. ^ Edward F. Knipling and Carrol Dayton Steelman: Feasibility of Controlling Ixodes scapularis Ticks (Acari: Ixodidae), the Vector of Lyme Disease, by Parasitoid Augmentation . In: Journal of Medical Entomology 2000, Volume 37, No. 5, pp. 645-652, doi : 10.1603 / 0022-2585-37.5.645 .
  30. ↑ Kathrin Hartelt et al .: Biological control of the tick Ixodes ricinus with entomopathogenic fungi and nematodes: Preliminary results from laboratory experiments . In: International Journal of Medical Microbiology 2008, Volume 298, Supplement 1, pp. 314-320, doi : 10.1016 / j.ijmm.2007.10.003 .
  31. ^ Richard S. Ostfeld et al .: Controlling Ticks and Tick-borne Zoonoses with Biological and Chemical Agents . In: BioScience 2006, Volume 56, No. 5, pp. 383-394, doi : 10.1641 / 0006-3568 (2006) 056 [0383: CTATZW] 2.0.CO; 2 .
  32. ↑ a b Ellen Tijsse-Klasen, Marieta Braks, Ernst-Jan Scholte, Hein Sprong: Parasites of vectors - Ixodiphagus hookeri and its Wolbachia symbionts in ticks in the Netherlands . In: Parasites & Vectors 2011, Volume 4, Article 228, doi : 10.1186 / 1756-3305-4-228 .
  33. ↑ a b Olivier Plantard et al .: Detection of Wolbachia in the Tick Ixodes ricinus is Due to the Presence of the Hymenoptera Endoparasitoid Ixodiphagus hookeri . In: PLOS ONE 2012, Volume 7, No. 1, Article e30692, doi : 10.1371 / journal.pone.0030692 .
  34. ↑ a b Geetha Subramanian et al .: Multiple tick-associated bacteria in Ixodes ricinus from Slovakia . In: Ticks and Tick-borne Diseases 2012, Volume 3, No. 5-6, pp. 406-410, doi : 10.1016 / j.ttbdis.2012.10.001 .
  35. ↑ Monika Bohacsova et al .: Arsenophonus nasoniae and Rickettsiae Infection of Ixodes ricinus Due to Parasitic Wasp Ixodiphagus hookeri . In: PLOS ONE 2016, Volume 11, No. 2, Article e0149950, doi : 10.1371 / journal.pone.0149950 .
  36. ↑ Jonathan Kagemann and Keith Clay: Effects of Infection by Arsenophonus and Rickettsia Bacteria on the Locomotive Ability of the Ticks Amblyomma americanum, Dermacentor variabilis, and Ixodes scapularis . In: Journal of Medical Entomology 2013, Volume 50, No. 1, pp. 155-162, doi : 10.1603 / ME12086 .
  37. ↑ Oleg Mediannikov et al .: Isolation of Arsenophonus nasoniae from Ixodes ricinus ticks in Slovakia . In: Ticks and Tick-borne Diseases 2012, Volume 3, No. 5-6, pp. 367-370, doi : 10.1016 / j.ttbdis.2012.10.016 .
  38. ↑ a b c Alexandre A. Girault : Notes and descriptions of Australian chalcid-flies III. (Hymenoptera) . In: Insecutor Inscritiae Menstruus 1925, Volume 13, No. 4-6, pp. 91-100, here pp. 96-97, digitizedhttp: //vorlage_digitalisat.test/1%3Dhttp%3A%2F%2Fwww.nhm.ac.uk%2Fresources%2Fresearch-curation%2Fprojects%2Fchalcidoids%2Fpdf_X%2FGiraul925b.pdf~GB%3D~IA%3D~MDZ% 3D% 0A ~ SZ% 3D ~ double-sided% 3D ~ LT% 3D ~ PUR% 3D .
  39. ↑ a b c d John S. Noyes and Mohammad Hayat: A review of the genera of Indo-Pacific Encyrtidae (Hymenoptera: Chalcidoidea) . In: Bulletin of the British Museum of Natural History (Entomology) 1984, Volume 48, No. 3, pp. 131-395, here p. 288, digitizedhttp: //vorlage_digitalisat.test/1%3D~GB%3D~IA%3Dbulletinofbritis48entolond~MDZ%3D%0A~SZ%3Dn308~ double-sided%3D~LT%3D~PUR%3D .
  40. ↑ Marija N. Nikolskaja : Новы ви паразит иксодовы клеще Ixodiphagus hirtus Nik., Sp. п. (Hymenoptera, Chalcidoidea) . In: Паразитол. сб. ЗИ А СССР ( Parazitologicheski Sbornik ) 1950, Volume 12, pp. 272-274 (English: A new Species of Parasite of Ixodid Ticks Ixodiphagus hirtus Nik., Sp. N. (Hymenoptera, Chalcidoidea) , not viewed).
  41. ↑ Vladimir A. Trjapitzin : Переописание Ixodiphagus hirtus (Hyraenoptera, Encyrtidae) - паразита таежного клеща Ixodes persulcatus на Дальнем ВосССРем . In: Parazitologiia 1982, Volume 16, No. 6, pp. 489-493, digitized versionhttp: //vorlage_digitalisat.test/1%3Dhttps%3A%2F%2Fwww.zin.ru%2Fjournals%2Fparazitologiya%2Fcontent%2F1982%2Fprz_1982_6_10_Trjapitzin.pdf~GB%3D~IA%3D~MDZ%3D%0 3D ~ double-sided% 3D ~ LT% 3D ~ PUR% 3D (English: Redescription of Ixodiphagus hirtus (Hymenoptera, Encyrtidae), a parasite of the taiga tick Ixodes persulcatus in the Far East of the USSR ).
  42. ↑ a b c Leland Ossian Howard: Another Chalcidoid Parasite of a Tick . In: The Canadian Entomologist 1908, Volume XL, No. 7, pp. 239-241, digitizedhttp: //vorlage_digitalisat.test/1%3D~GB%3D~IA%3Dcanadianentomolo40ento~MDZ%3D%0A~SZ%3Dn259~ double-sided%3D~LT%3D~PUR%3D .
  43. ↑ Leland Ossian Howard : A Chalcidid Parasite of a Tick . In: Entomological News 1907, Volume XVIII, No. 9, pp. 375-378, Plate XIV, digitizedhttp: //vorlage_digitalisat.test/1%3D~GB%3D~IA%3Dentomologicalnew18acaduoft~MDZ%3D%0A~SZ%3Dn421~ double-sided%3D~LT%3D~PUR%3D .
  44. ^ OGH Fiedler: A new African tick parasite, Hunterellus theilerae, n.sp. In: Onderstepoort Journal of Veterinary Research 1953, Volume 26, No. 1, pp. 61-63, digitizedhttp: //vorlage_digitalisat.test/1%3Dhttps%3A%2F%2Frepository.up.ac.za%2Fbitstream%2Fhandle%2F2263%2F58706%2F4fiedler1953.pdf%3Fsequence%3D1~GB%3D~IA%3D~MDZ% 3D% 0A ~ SZ% 3D ~ double-sided% 3D ~ LT% 3D ~ PUR% 3D .
  45. ^ John S. Noyes: A review of the genera of Neotropical Encyrtidae (Hymenoptera: Chalcidoidea) . In: Bulletin of the British Museum of Natural History (Entomology) 1980, Volume 41, No. 2, pp. 107-253, here p. 204, digitizedhttp: //vorlage_digitalisat.test/1%3D~GB%3D~IA%3Dbulletinofbritis41entolond~MDZ%3D%0A~SZ%3Dn127~ double-sided%3D~LT%3D~PUR%3D .
  46. ^ John S. Noyes: Encyrtidae (Insecta: Hymenoptera) . Fauna of New Zealand 13. DSIR Science Information Publishing Center, Wellington 1988, ISBN 0-477-02517-X , pp. 9-12, digitizedhttp: //vorlage_digitalisat.test/1%3Dhttps%3A%2F%2Fwww.biotaxa.org%2Ffnz%2Farticle%2Fview%2F1753%2F2888~GB%3D~IA%3D~MDZ%3D%0A~SZ%3D~ double-sided% 3D ~ LT% 3D ~ PUR% 3D .
  47. ^ Arthur B. Gahan: On the Identities of Chalcidoid Tick Parasites (Hymenoptera) . In: Proceedings of the Entomological Society of Washington 1934, Vol. 36, No. 4, pp. 89-97, digitizedhttp: //vorlage_digitalisat.test/1%3D~GB%3D~IA%3Dproceedingsofent3536193334ento~MDZ%3D%0A~SZ%3Dn347~ double-sided%3D~LT%3D~PUR%3D .
  48. ↑ Wladimir A. Trjapitzin: Natural enemies of the Taiga tick . In: NA Filippowa (Ed.): The Taiga tick Ixodes persulcatus (Acarina, Ixodidae). Morphology, Systematics, Ecology, Medical importance . Nauky Leningrad, Leningrad 1985, pp. 334-341 (Russian).