Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Rhagoletis completa
(walnut husk fly)

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Datasheet

Rhagoletis completa (walnut husk fly)

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TitleLine drawing of adult R. completa
Caption
Copyright©CABI BioScience
Line drawing of adult R. completa©CABI BioScience

Identity

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Preferred Scientific Name

  • Rhagoletis completa Cresson

Preferred Common Name

  • walnut husk fly

Other Scientific Names

  • Rhagoletis suavis completa Cresson
  • Rhagoletis suavis var. completa Cresson

International Common Names

  • English: husk maggot
  • French: mouche des brous du noyer

Local Common Names

  • North America: husk maggot
  • Germany: Amerikanische Walnussschalen-Fliege; Fliege, Amerikanische Walnussschalen

EPPO code

  • RHAGCO (Rhagoletis completa)

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Diptera
  •                         Family: Tephritidae
  •                             Genus: Rhagoletis
  •                                 Species: Rhagoletis completa

Notes on Taxonomy and Nomenclature

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R. completa was originally described as a subspecies of R. suavis but these have been recognized as separate species since the 1930s (see Foote et al., 1993).

Description

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Adult

Diagnostic features of the genus are as follows (characters extracted from key to North American genera of Tephritidae by Foote et al., 1993): Head with two pairs orbital setae; posterior pair reclinate. Gena with only short anterior setae. First flagellomere (third antennal segment) at least slightly pointed at the apex. Thorax with dorsocentral setae closer to level of anterior supra-alar setae than transverse suture. Scutellum not swollen or shiny. Wing with cells bm and bcu of similar depth; bcu with a short acute extension. Crossvein R-M near middle of cell dm.

This species may be identified using the Diptera key in the Crop Protection Compendium taxonomic identification aid. For full details of its separation from other North American species, see Foote et al. (1993).

The main features of R. completa are as follows: thorax and abdomen predominantly pale yellow to orange. Wing with a distinct subbasal crossband; discal and preapical crossbands usually separate. Mediotergite dark marked (either entirely dark brown or with a pair of brown stripes). For more comprehensive details see Foote et al. (1993).

Larva

Diagnosis of genus by Elson-Harris (White and Elson-Harris, 1994): Antennal sensory organ with a short basal segment and cone-shaped distal segment; maxillary sensory organ flat, with well defined sensilla surrounded by small cuticular folds; stomal sensory organ rounded, with a peg-like sensilla; large, preoral teeth near base of stomal sensory organ; no preoral lobes; oral ridges in 5-13 short, unserrated rows; no accessory plates. Stout spinules forming discontinuous rows on almost all segments. Anterior spiracles with 7-35 stout tubules. Posterior spiracular slits 3-8 times as long as broad, with 3-16 short, branched spiracular hairs. Anal lobes large, protuberant with well defined tubercles and sensilla.

There is no modern description of the larva of this species (in the sense of SEM studies being applied). However, Steyskal (1973) provided a key for the separation of the larvae of three species found in walnut. Any Rhagoletis larvae found in walnuts and having the following features is likely to be this species: 16-21 tubules, in each anterior spiracle; upper and lower slits of the posterior spiracles at about 60 degrees to each other. See key to larvae in White and Elson-Harris (1994), which used a combination of host and fragmentary morphological data.

Distribution

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R. completa is native to southern and central USA; adventive in western USA since the 1920s (Bush, 1966). Also adventive in southern Europe since the early 1990s (Ciampolini and Trematerra, 1992; Merz, 1994; Seljak and Zezlina, 1999).

Harris (1989), Foote et al. (1993) and CABI (1997) all provide distribution maps.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Last updated: 26 Nov 2020
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Europe

AustriaPresent, Few occurrences
BelgiumPresent, Localized
Bosnia and HerzegovinaPresent
CroatiaPresent
CzechiaPresent, Localized
FrancePresent, Localized
-CorsicaPresent, Localized
GermanyPresent, Localized
HungaryPresent, Localized
ItalyPresent, Localized
NetherlandsPresent, Few occurrences
PolandPresent
SlovakiaPresent, Few occurrences
SloveniaPresent, Localized1997
SpainPresent, Few occurrencesGirona.
SwitzerlandPresent, Widespread1986
United KingdomAbsent, Confirmed absent by survey

North America

CanadaPresent, Localized
-British ColumbiaPresent, Localized
MexicoPresent, Localized
United StatesPresent
-ArizonaPresent
-CaliforniaPresent, Few occurrences
-ColoradoPresent
-IdahoPresent
-IowaPresent
-KansasPresent
-MinnesotaPresent
-MississippiPresent
-MissouriPresent
-NebraskaPresent
-NevadaPresent
-New MexicoPresent
-OklahomaPresent
-OregonPresent
-TexasPresent
-UtahPresent
-WashingtonPresent

Oceania

New ZealandAbsent, Confirmed absent by survey

Risk of Introduction

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R. completa poses a small quarantine risk for temperate regions. However, now that it is established in Switzerland and Italy, this species has been removed from the European and Mediterranean Plant Protection Organization (EPPO) A1 list. However, there are no practical measures to prevent its spread and it is not considered to be a very important pest, so EPPO recently decided not to add it to its A2 list. It has not been individually considered to be a quarantine pest by any other Regional Plant Protection Organization.

Habitat

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Areas with suitable hosts and climate.

Hosts/Species Affected

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R. completa is mainly associated with Juglans species. It is also known to attack Prunus persica (peach) (Bush, 1966).

Growth Stages

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Fruiting stage

Symptoms

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Attacked fruit will be pitted by oviposition punctures, around which some discoloration usually occurs.

List of Symptoms/Signs

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SignLife StagesType
Fruit / discoloration
Fruit / extensive mould
Fruit / gummosis
Fruit / internal feeding
Fruit / lesions: black or brown
Fruit / lesions: scab or pitting
Fruit / obvious exit hole
Fruit / odour
Fruit / ooze

Biology and Ecology

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Most Rhagoletis species have a similar biology as detailed by Christenson and Foote (1960): eggs are laid below the skin of the host fruit and hatch after 3-7 days; the larvae usually feed for 2-5 weeks; pupariation is in the soil under the host plant and this is the normal overwintering stage; adults may live for up to 40 days under field conditions. In Oregon, adults emerge from late June to mid-August or sometimes early September; peak oviposition is in late August (Kasana and Aliniazee, 1996).

For development rates at various temperatures see Kasana and AliNiazee (1994).

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Biosteres sublaevis Parasite Larvae
Coptera occidentalis Parasite
Trybliographa sp. Parasite Larvae

Notes on Natural Enemies

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The braconid, Biosteres sublaevis, and a eucoilid of the genus Trybliographa, were found to cause varying degrees of natural parasitism in New Mexico (Legner and Goeden, 1987).

Means of Movement and Dispersal

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Adult flight and the transport of infected fruit are the major means of movement and dispersal to previously uninfected areas. However, Rhagoletis species are not known to fly more than a short distance (Fletcher, 1989). In international trade, the major means of dispersal to previously uninfested areas is the transport of fruit containing live larvae. There is also a risk from the transport of puparia in soil or packaging with plants which have already fruited.

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Clothing, footwear and possessionsFruit in case or handbag. Yes
Containers and packaging - woodOf fruit cargo. Yes
Land vehiclesLorries, aeroplanes and perhaps ships, with fruit cargo. Yes
MailFruit in post. Yes
Soil, sand and gravelRisk of puparia in soil. Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Fruits (inc. pods) eggs; larvae Yes Pest or symptoms usually visible to the naked eye
Growing medium accompanying plants pupae Yes Pest or symptoms usually visible to the naked eye
Plant parts not known to carry the pest in trade/transport
Bark
Bulbs/Tubers/Corms/Rhizomes
Flowers/Inflorescences/Cones/Calyx
Leaves
Roots
Seedlings/Micropropagated plants
Stems (above ground)/Shoots/Trunks/Branches
True seeds (inc. grain)
Wood

Impact

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In North America, R. completa is a pest of various Juglans species but it has rarely been recorded on European walnut (J. regia). In Switzerland, it attacks the mesocarp of J. regia, and in the case of severe infestation may also damage the pericarp and the nut itself (R. Mani, Switzerland [address available from CABI], personal communication, 2001). It therefore becomes a problem for walnut fruit production. In 1991, 50% of harvested walnuts in some Italian orchards were infested by R. completa (Ciampolini and Trematerra, 1992). In Oregon, Kasana and Aliniazee (1996) recorded up to 95% infestation on untreated trees.

In California it causes two types of damage. In populations causing normal or late-season infestations, larval feeding causes the whole husk or large portions of it to turn black, seriously reducing the value of nuts sold in the shell although the kernel is undamaged. Attack by populations involved in early infestations impedes the maturation of the kernel and results in shrivelled nuts or empty shells (Hislop et al., 1981).

Diagnosis

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EPPO (2011) describes a diagnostic protocol for R. completa infesting walnuts.

Detection and Inspection

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Traps have been developed which capture both sexes, based on visual, or visual plus odour, attraction. They are coated in sticky material and are usually either flat-surfaced and coloured fluorescent yellow to elicit a supernormal foliage response (see Riedl et al., 1989), or spherical and dark-coloured to represent a fruit (green spheres are best, see Riedl and Hislop, 1985); traps which combine both foliage and fruit attraction can also be used. The odour comes from protein hydrolysate or other substances emitting ammonia, such as ammonium acetate. See Boller and Prokopy (1976) and Economopoulos (1989) for a discussion of these traps.

Similarities to Other Species/Conditions

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This species should not be mistaken for any other provided the diagnostic notes (see Morphology) are applied carefully. If in doubt consult Foote et al. (1993). R. suavis is an eastern USA species which is also associated with Juglans species, and in the past has been confused with R. completa (see Distribution Section).

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Upon detection, fallen and infected fruit must be removed and destroyed. If possible, wild and abandoned host trees should also be destroyed. Boller and Prokopy (1976) note that systemic organophosphates, such as dimethoate, are highly effective against most species, killing eggs, larvae and adults. Belanger et al. (1985) discussed the use of pyrethroids, but these were only of use when pest activity was low. More environmentally acceptable techniques have been tried; namely bait sprays (insecticide plus ammonia source) which can be applied as a spot treatment; soil application of insecticide to destroy pupae; and juvenile hormone analogues which can be applied to the soil (Boller and Prokopy, 1976). The IPM of walnut pests, including R. completa, was discussed by Haley and Baker (1982). Early harvest can be used to avoid attack (Yokoyama and Miller, 1996). Some varieties may be more susceptible to attack than others. Shelton and Anderson (1990) compared four varieties and found that Hartley suffered less damage than early leafing varieties.

Consignments of fruits from countries where these pests occur should be inspected for symptoms of infestation and those suspected should be cut open in order to look for larvae. European and Mediterranean Plant Protection Organization (EPPO) recommended (OEPP/EPPO, 1990), when only the North American origin was under consideration, that such fruits should come from an area where R. completa does not occur or from a place of production subject to growing season inspection. Fruits may also be treated in transit by cold treatment (for example, 40-42 days at -0.6°C) (FAO, 1983); however, Yokoyama and Miller (1996) found that low temperature storage could not be used as a quarantine treatment alone for some crops (e.g. walnuts) because the fly tolerated the treatment better than the host. Ethylene dibromide was previously widely used as a fumigant but is now generally withdrawn because of its carcinogenicity. Treatment methods against fruit flies are currently under review within EPPO and as part of an inter-RPPO programme.

Plants of host species transported with roots from countries where these pests occur should be free from soil, or the soil should be treated against puparia. The plants should not carry fruits. Such plants may indeed be prohibited from importation.

References

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Aluja, M., Guillén, L., Rull, J., Höhn, H., Frey, J., Graf, B., Samietz, J., 2011. Is the alpine divide becoming more permeable to biological invasions? - Insights on the invasion and establishment of the Walnut Husk Fly, Rhagoletis completa (Diptera: Tephritidae) in Switzerland. Bulletin of Entomological Research, 101(4), 451-465. doi: 10.1017/S0007485311000010

Armstrong JW; Couey HM, 1989. Control; fruit disinfestation; fumigation, heat and cold. In: Robinson AS, Hooper G, eds. Fruit Flies; their Biology, Natural Enemies and Control. World Crop Pests. Amsterdam, Netherlands: Elsevier, 3(B):411-424.

Belanger A; Bostanian NJ; Rivard I, 1985. Apple maggot (Diptera: Trypetidae) control with insecticides and their residues in and on apples. Journal of Economic Entomology, 78(2):463-466.

Berlocher, S. H., 1984. Genetic Changes Coinciding with the Colonization of California by the Walnut Husk Fly, Rhagoletis completa. Evolution, 38(4), 906-918. doi: 10.1111/j.1558-5646.1984.tb00361.x

Bjelis M, 2008. Fruit flies from the family Rhagoletis (Tephritidae) in Croatia. (Vocne muhe iz roda Rhagoletis (Tephritidae) u Hrvatskoj.) Glasilo Biljne Zastite, 8(1):25-28.

Boller EF; Prokopy RJ, 1976. Bionomics and management of Rhagoletis. In: Smith RF, Mittler TE, Smith CN, ed. Annual review of entomology. Volume 21. Annual Reviews Inc. Palo Alto, California, USA, 223-246.

Boyce, A. M., 1934. Bionomics of the walnut husk fly (Rhagoletis completa). Hilgardia, 8, 363-579.

Bush GL, 1966. The taxonomy, cytology and evolution of the genus Rhagoletis in North America (Diptera: Tephritidae). Bulletin of the Museum of Comparative Zoology, 134:431-526.

CABI/EPPO, 1997. Rhagoletis completa Cresson. Distribution Maps of Plant Pests, No. 337, rev. ed. Wallingford, UK: CAB International.

CABI/EPPO, 1998. Distribution maps of quarantine pests for Europe (edited by Smith IM, Charles LMF). Wallingford, UK: CAB International, xviii + 768 pp.

CABI/EPPO, 2014. Rhagoletis completa. [Distribution map]. Distribution Maps of Plant Pests, No.December. Wallingford, UK: CABI, Map 337 (2nd revision).

Chen, Y. H., Berlocher, S. H., Opp, S. B., Roderick, G. K., 2010. Post-colonization temporal genetic variation of an introduced fly, Rhagoletis completa. Genetica, 138(9/10), 1059-1075. doi: 10.1007/s10709-010-9491-7

Chen, Y. H., Opp, S. B., Berlocher, S. H., Roderick, G. K., 2006. Are bottlenecks associated with colonization? Genetic diversity and diapause variation of native and introduced Rhagoletis completa populations. Oecologia, 149(4), 656-667. doi: 10.1007/s00442-006-0482-4

Christenson LD; Foote RH, 1960. Biology of fruit flies. Annual Review of Entomology, 5:171-192.

Ciampolini M; Trematerra P, 1992. Widespread occurrence of walnut fly (Rhagoletis completa Cresson) in northern Italy. Informatore Agrario, 48(48):52-56

Drosopoulou, E., Koeppler, K., Kounatidis, I., Nakou, I., Papadopoulos, N. T., Bourtzis, K., Mavragani-Tsipidou, P., 2010. Genetic and cytogenetic analysis of the walnut-husk fly (Diptera: Tephritidae). Annals of the Entomological Society of America, 103(6), 1003-1011. doi: 10.1603/AN10059

Duso, C., Lago, G. dal, 2006. Life cycle, phenology and economic importance of the walnut husk fly Rhagoletis completa Cresson (Diptera: Tephritidae) in northern Italy. Annales de la Société Entomologique de France, 42(2), 245-254.

Economopoulos AP, 1989. Control; use of traps based on color and/or shape. In: Robinson AS, Hooper G, eds. Fruit Flies; Their Biology, Natural Enemies and Control. World Crop Pests 3(B): 315-327. Amsterdam, Netherlands: Elsevier.

EPPO, 1990. Specific quarantine requirements. EPPO Technical Documents, No. 1008. Paris, France: European and Mediterranean Plant Protection Organization.

EPPO, 2011. EPPO Reporting Service. EPPO Reporting Service. Paris, France: EPPO. http://archives.eppo.org/EPPOReporting/Reporting_Archives.htm

EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm

EPPO, 2016. EPPO Global database (available online). Paris, France: EPPO. https://gd.eppo.int/

European and Mediterranean Plant Protection Organization, 2011. Diagnostics: Rhagoletis completa. Bulletin OEPP/EPPO Bulletin, 41(3):357-362. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1365-2338

FAO, 1983. International plant quarantine treatment manual. FAO Plant Production and Protection Paper No. 50. Rome, Italy: FAO.

Fletcher BS, 1989. Ecology; movements of tephritid fruit flies. In: Robinson AS, Hooper G, eds. Fruit Flies; Their Biology, Natural Enemies and Control. World Crop Pests, 3(B). Amsterdam, Netherlands: Elsevier, 209-219.

Foote RH, 1981. The genus Rhagoletis Loew south of the United States (Diptera: Tephritidae). Technical Bulletin, Science and Education Administration, United States Department of Agriculture, No. 1607:iv + 75 pp.

Foote RH; Blanc FL; Norrbom AL, 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Ithaca, USA: Comstock.

Guillén, L., Aluja, M., Rull, J., Höhn, H., Schwizer, T., Samietz, J., 2011. Influence of walnut cultivar on infestation by Rhagoletis completa: behavioural and management implications. Entomologia Experimentalis et Applicata, 140(3), 207-217. doi: 10.1111/j.1570-7458.2011.01157.x

Haley MJ; Baker L, ed. , 1982. Integrated pest management for walnuts. Integrated pest management for walnuts. University of California Statewide Integrated Pest Management Project Berkeley USA, 96 pp.

Harris EJ, 1989. Pest status; Hawaiian Islands and North America, In: Robinson AS, Hooper G, eds. Fruit Flies; their Biology, Natural Enemies and Control. World Crop Pests. Amsterdam, Holland: Elsevier, 3(A):73-81.

Hislop RG; Riedl H; Joos JL, 1981. Control of the walnut husk fly with pyrethroids and bait. California Agriculture, 35(9/10):23-25

Hood, G. R., Egan, S. P., Feder, J. L., 2012. Evidence for sexual isolation as a prezygotic barrier to gene flow between morphologically divergent species of Rhagoletis fruit flies. Ecological Entomology, 37(6), 521-528. doi: 10.1111/j.1365-2311.2012.01392.x

IPPC, 2013. First report of Rhagoletis completa. IPPC Official Pest Report, No. HUN-02/1. Rome, Italy: FAO. https://www.ippc.int/

IPPC, 2016. First suspicion of Rhagoletis completa on Juglans regia in a private garden. IPPC Official Pest Report, No. NLD-46/1. https://www.ippc.int/en/

Kasana A; AliNiazee MT, 1994. Effect of constant temperatures on development of the walnut husk fly, Rhagoletis completa. Entomologia Experimentalis et Applicata, 73(3):247-254

Kasana A; AliNiazee MT, 1996. Seasonal phenology of the walnut husk fly, Rhagoletis completa Cresson (Diptera: Tephritidae). Canadian Entomologist, 128(3):377-390; 29 ref.

Legner EF; Goeden RD, 1987. Larval parasitism of Rhagoletis completa (Diptera: Tephritidae) on Juglans microcarpa (Juglandaceae) in western Texas and southeastern New Mexico. Proceedings of the Entomological Society of Washington, 89(4):739-743

McPheron, B. A., Steck, G. J., 1996. Fruit Fly Pests: A World Assessment of Their Biology and Management. USA: St. Lucie Press.https://agris.fao.org/agris-search/search.do?recordID=US9617737

Merz B, 1994. Insecta Helvetica, Fauna, 10: Diptera Tephritidae. Neuchatel, Switzerland; Centre Suisse de Cartographie de la Faune (CSCF), 198 pp.

Ostojic I; Zovko M; Petrovic D, 2014. First record of walnut husk fly Rhagoletis completa (Cresson, 1929) in Bosnia and Herzegovina. (Prvi nalaz orahove muhe Rhagoletis completa (Cresson, 1929) u Bosni i Hercegovini.) Radovi Poljoprivrednog Fakulteta Univerziteta u Sarajevu (Works of the Faculty of Agriculture University of Sarajevo), 59(64(1)):121-126.

Riedl H; Barnett WW; Coates WW; Coviello R; Joos J; Olson WH, 1989. Walnut husk fly (Diptera: Tephritidae): evaluation of traps for timing of control measures and for damage predictions. Journal of Economic Entomology, 82(4):1191-1196

Riedl H; Hislop R, 1985. Visual attraction of the walnut husk fly (Diptera: Tephritidae) to color rectangles and spheres. Environmental Entomology, 14(6):810-814

Rull J; Tadeo E; Aluja M; Guillen L; Egan SP; Feder JL, 2012. Hybridization and sequential components of reproductive isolation between parapatric walnut-infesting sister species Rhagoletis completa and Rhagoletis zoqui. Biological Journal of the Linnean Society, 107(4):886-898. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1095-8312

Rull, J., Aluja, M., Tadeo, E., Guillen, L., Egan, S., Glover, M., Feder, J. L., 2013. Distribution, host plant affiliation, phenology, and phylogeny of walnut-infesting Rhagoletis flies (Diptera: Tephritidae) in Mexico. Biological Journal of the Linnean Society, 110(4), 765-779. doi: 10.1111/bij.12157

Sarles, L., Boullis, A., Fassotte, B., Lognay, G., Verhaeghe, A., Francis, F., Verheggen, F. J., 2017. Identification of walnut husk (Juglans regia L.) volatiles and the behavioural response of the invasive Walnut Husk Fly, Rhagoletis completa Cresson. Pest Management Science, 73(10), 2100-2104. doi: 10.1002/ps.4584

Seljak G; Zezlina I, 1999. Appearance and distribution of walnut husk fly (Rhagoletis completa Cresson) in Slovenia. Zbornik predavanj in referatov 4. Slovenskega Posvetovanja o Varstvu Rastlin v Portoroz^hacek~u od 3. do 4. Marca 1999., 231-238; 7 ref.

Shelton MD; Anderson JL, 1990. Walnut cultivars: evidence for differential susceptibility to insect pests. Fruit Varieties Journal, 44(4):179-182

Steyskal GC, 1973. Distinguishing characters of the walnut husk maggots of the genus Rhagoletis (Diptera, Tephritidae). Cooperative Economic Insect Report, 23: 522.

Tuba K; Schuler H; Stauffer C; Lakatos F, 2012. First record of the walnut husk fly (Rhagoletis completa Cresson 1929) (Diptera: Tephritidae) in Hungary. (A nyugati dióburok-fúrólégy (Rhagoletis completa Cresson 1929 - Diptera: Tephritidae) megjelenése magyarországon.) Növényvédelem, 48(9):419-424.

White IM; Elson-Harris MM, 1994. Fruit flies of economic significance: their identification and bionomics. Wallingford, UK: CAB International. Reprint with addendum.

Yee, W. L., Goughnour, R. B., 2008. Host plant use by and new host records of apple maggot, western cherry fruit fly, and other Rhagoletis species (Diptera: Tephritidae) in western Washington state. Pan-Pacific Entomologist, 84(3), 179-193. doi: 10.3956/2007-49.1

Yee, W. L., Goughnour, R. B., 2016. Relative efficacies of sticky yellow rectangles against three Rhagoletis fly species (Diptera: Tephritidae) in Washington State and possible role of adhesives. Journal of the Entomological Society of British Columbia, 113, 49-70. https://journal.entsocbc.ca/index.php/journal/article/view/912/1000

Yokoyama VY; Miller GT, 1996. Response of walnut husk fly (Diptera: Tephritidae) to low temperature, irrigation, and pest-free period for exported stone fruits. Journal of Economic Entomology, 89(5):1186-1191; 7 ref.

Yokoyama, V. Y., Miller, G. T., 1993. Pest-free period for walnut husk fly (Diptera: Tephritidae) and host status of stone fruits for export to New Zealand. Journal of Economic Entomology, 86(6), 1766-1772. doi: 10.1093/jee/86.6.1766

Distribution References

Aluja M, Guillén L, Rull J, Höhn H, Frey J, Graf B, Samietz J, 2011. Is the alpine divide becoming more permeable to biological invasions? - Insights on the invasion and establishment of the Walnut Husk Fly, Rhagoletis completa (Diptera: Tephritidae) in Switzerland. Bulletin of Entomological Research. 101 (4), 451-465. DOI:10.1017/S0007485311000010

Berlocher S H, 1984. Genetic Changes Coinciding with the Colonization of California by the Walnut Husk Fly, Rhagoletis completa. Evolution. 38 (4), 906-918. DOI:10.1111/j.1558-5646.1984.tb00361.x

Bjeliš M, 2008. Fruit flies from the family Rhagoletis (Tephritidae) in Croatia. (Voćne muhe iz roda Rhagoletis (Tephritidae) u Hrvatskoj.). Glasilo Biljne Zaštite. 8 (1), 25-28.

Bouvet G, 2009. The walnut industry of the southeast faces the walnut fruit fly. (La filière noix du Sud-Est fait face à la mouche du brou.). Arboriculture Fruitière. 24-26.

Boyce A M, 1934. Bionomics of the walnut husk fly (Rhagoletis completa). Hilgardia. 363-579.

CABI, EPPO, 1997. Rhagoletis completa. [Distribution map]. In: Distribution Maps of Plant Pests, Wallingford, UK: CAB International. Map 337. DOI:10.1079/DMPP/20066600337

CABI, EPPO, 2014. Rhagoletis completa. [Distribution map]. In: Distribution Maps of Plant Pests, Wallingford, UK: CABI. Map 337 (2nd revisio. DOI:10.1079/DMPP/20153006586

CABI, Undated. Compendium record. Wallingford, UK: CABI

CABI, Undated a. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

Chen Y H, Berlocher S H, Opp S B, Roderick G K, 2010. Post-colonization temporal genetic variation of an introduced fly, Rhagoletis completa. Genetica. 138 (9/10), 1059-1075. DOI:10.1007/s10709-010-9491-7

Chen Y H, Opp S B, Berlocher S H, Roderick G K, 2006. Are bottlenecks associated with colonization? Genetic diversity and diapause variation of native and introduced Rhagoletis completa populations. Oecologia. 149 (4), 656-667. DOI:10.1007/s00442-006-0482-4

Ciampolini M, Trematerra P, 1992. Widespread occurrence of walnut fly (Rhagoletis completa Cresson) in northern Italy. (Diffusa presenza di mosca delle noci (Rhagoletis completa Cresson) nel nord Italia.). Informatore Agrario. 48 (48), 52-56.

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