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Annotated bibliography

Laboratory host range testing

Entomophagous biological control agents

References

Althoff D.M. (2003). Does parasitoid attack strategy influence host specificity? A test with New World braconids. Ecological Entomology 28: 500-502.
Parasitoid attack strategy has been divided into koinobiosis and idiobiosis, based on the arrest of host development and the intimacy of larval contact. Comparisons from specific host communities have shown that koinobionts are more host specific than idiobionts. Koinobiont genera utilised fewer host families than idiobionts, suggesting that parasitoid attack strategy may direct the evolution of host specificity throughout the evolutionary history of parasitoid lineages.

Babendreier D., Bigler F. and Kuhlmann U. (2005). Methods Used to Assess Non-target Effects of Invertebrate Biological Control Agents of Arthropod Pests. BioControl 50: 821-870.
An overview of methods currently applied in the study of non-target effects in biological control of arthropod pests. It provides the first step towards the ultimate goal of devising guidelines for the appropriate methods that should be universally applied for the assessment and minimisation of potential non-target effects. The topics reviewed include host specificity (including field surveys, selection of non-target test species and testing protocols), post-release studies, competition, overwintering and dispersal.

Babendreier D., Kuske S. and Bigler F. (2003). Non-target host acceptance and parasitism by Trichogramma brassicae Bezdenko (Hymenoptera: Trichogrammatidae) in the laboratory. Biological Control 26: 128-138.
Case study of host range testing Trichogramma brassicae Bezdenko using eggs of 23 non-target lepidopteran species including nine butterflies endangered in Switzerland to parasitoids under no-choice conditions in the laboratory. The results show that T. brassicae parasitizes a number of non-target lepidopteran eggs belonging to different families.

Barratt B.I.P. (2004). Microctonus parasitoids and New Zealand weevils: comparing laboratory estimates of host ranges to realized host ranges. Pp. 103-120 In: Assessing host ranges for parasitoids and predators used for classical biological control: A guide to best practice, R.G. Van Driesche and R. Reardon (Ed.) USDA Forest Service, Morgantown, West Virginia.

Barratt B.I.P. and Johnstone P.D. (2001). Factors affecting parasitism by Microctonus aethiopoides Loan (Hymenoptera: Braconidae) and parasitoid development in natural and novel host species. Bulletin of Entomological Research 91: 245-253.
A laboratory study of aspects of parasitoid host acceptance, suitability and physiological regulation in natural and novel host species was carried out to investigate the degree of variability encountered with different hosts and to determine the value of such observations in host range determination. The study uses Microctonus aethiopoides Loan as a model braconid parasitoid. It was concluded that laboratory observations can provide useful information on the compatibility between host and parasitoid which can complement traditional host range tests to predict field host range.

Barratt B.I.P., Evans A.A. and Johnstone P.D. (1996). Effect of the ratios of Listronotus bonariensis and Sitona discoideus (Coleoptera: Curculionidae) to their respective parasitoids Microctonus hyperodae and Microctonus aethiopoides (Hymenoptera: Braconidae), on parasitism, host oviposition and feeding in the laboratory. Bulletin of Entomological Research 86: 101-108.
Laboratory experiments were carried out to investigate the effect of host-parasitoid ratio, and exposure time on host survival, parasitism, oviposition and feeding. Over the ranges studied, increasing parasite number, and to a greater extent, period of exposure of parasitoids to their hosts increased parasitism levels. Effects of parasitoid exposure on host fecundity and feeding was assessed.

Barratt B.I.P., Evans A.A., Ferguson C.M., Barker G.M., McNeill M.R. and Phillips C.B. (1997). Laboratory nontarget host range of the introduced parasitoids Microctonus aethiopoides and Microctonus hyperodae (Hymenoptera: Braconidae) compared with field parasitism in New Zealand. Environmental Entomology 26: 694-702.
Laboratory host specificity of Microctonus aethiopoides Loan and Microctonus hyperodae Loan, braconids already imported to control weevil pests, was compared with actual field parasitism. NZ native, introduced, and beneficial species were tested. M. aethiopoides oviposited in 11 of the 12 species to which it was exposed and successfully parasitized 9 species. M. hyperodae oviposited in 5 of the 11 species to which it was exposed and developed successfully in 4 species. Higher percentage parasitism was recorded with M. aethiopoides than with M. hyperodae. Field collections of weevils indicated that 10 New Zealand native species and 3 other nontarget species, including the weed biological control agent Rhinocyllus conicus (Froehlich), were parasitized by M. aethiopoides. M. hyperodae has been found parasitizing only native species. Field parasitism levels in the field of >70% have been recorded for M. aethiopoides and <5% for M. hyperodae. The results of this study suggest that laboratory host range testing is indicative of nontarget parasitism in the field.

Barratt B.I.P., Ferguson C.M., McNeill M.R. and Goldson S.L. (1999). Parasitoid host specificity testing to predict host range. Pp. 70-83 In: Host specificity testing in Australasia: towards improved assays for biological control, T.M. Withers, L. Barton-Browne and J.N. Stanley (Ed.) CRC for Tropical Pest Management, Brisbane, Australia.

Barratt B.I.P., Phillips C.B., Ferguson C.M. and Goldson S.L. (2002). Predicting non-target impacts of parasitoids: where to from here? Pp. 378-386 In: Proceedings of First International Symposium on Biological Control of Arthropods, R. Van Driesche (Ed.) Forest Health Technology Enterprise Team, Morgantown, West Virginia.

Barratt B.I.P., Todd J. and Malone L.A. (2016). Selecting non-target species for arthropod biological control agent host range testing: evaluation of a novel method. Biological Control 93: 84-92.

Berndt L., Withers T.M., Mansfield S. and Hoare R.J.B. (2009). Non-target species selection for host range testing of Cotesia urabae. New Zealand Plant Protection 62: 168-173
The Australian solitary larval endoparasitoid Cotesia urabae (Hymenoptera: Braconidae) is a promising biocontrol agent for Uraba lugens. A non-target species list was compiled for host range testing. The endemic species Celama parvitis is the sole New Zealand representative of the Nolinae and was highest priority. The next most closely related subfamily is the Arctiinae, in which New Zealand has four endemic species (Metacrias huttoni, M. erichrysa, M. strategica and Nyctemera annulata) and one introduced biological control agent (Tyria jacobaeae). The merits of including other Lepidoptera are discussed.

Berndt L.A., Mansfield S. and Withers T.M. (2007). A method for host range testing of a biological control agent for Uraba lugens. New Zealand Plant Protection 60: 286-290.
Uraba lugens (gum leaf skeletoniser) is a serious pest of Eucalyptus spp. in Australia and is now well established in the greater Auckland region of New Zealand. Two parasitoid species are under consideration as potential biological control agents and this paper describes host range testing methods developed using one of these species (Cotesia urabae) against two non-target species, Helicoverpa armigera and Spodoptera litura. Using sequential no-choice tests clear preferences were observed for U. lugens over both non-target test species. Although some females did attempt to attack the non-target species, no evidence of parasitism was observed when reared or dissected. This method elucidated both behavioural responses and physiological development of C. urabae, and it is proposed to be a suitable host range testing method for full evaluation of this species.

Bigler F., Babendreier D. and Kuhlmann U. (2006). Environmental impact of arthropod biological control: methods and risk assessment. Pp. 288. CABI Publishing, Delemont, Switzerland.

Boyd E.A. and Hoddle M.S. (2007). Host specificity testing of Gonatocerus spp. egg-parasitoids used in a classical biological control program against Homalodisca vitripennis: a retrospective analysis for non-target impacts in southern California. Biological Control 43: 56-70
A host specificity testing protocol was developed for estimating potential physiological and ecological risk to non-target species posed by species of the egg parasitoids Gonatocerus spp. using choice and no-choice host options. The solitary Gonatocerus ashmeadi Girault and gregarious G. fasciatus Girault (Hymenoptera: Mymaridae) are non-native egg-parasitoids of the exotic Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae), and were introduced for classical biological control California, USA. The parasitoids' physiological and ecological host ranges were estimated on three non-target indigenous sharpshooters and results were compared with observed non-target impacts in the field. Laboratory tests with G. ashmeadi revealed Homalodisca liturata Ball (Hemiptera: Cicadellidae) eggs were a physiologically and ecologically acceptable host; Graphocephala atropunctata (Signoret) and Draeculacephala minerva Ball (Hemiptera: Cicadellidae) eggs were not acceptable hosts. Tests with G. fasciatus showed both H. liturata and D. minerva, but not G. atropunctata eggs, were physiologically acceptable hosts. Only H. liturata eggs were determined to be an ecologically acceptable host for G. fasciatus. Field surveys failed to find parasitism of G. atropunctata or D. minerva eggs by either G. ashmeadi or G. fasciatus.

Cameron P.J. and Walker G.P. (1997). Host specificity of Cotesia rubecula and Cotesia plutellae, parasitoids of white butterfly and diamondback moth. Pp. 236-241 In: Proceedings of the 50th New Zealand Plant Protection Conference, M. O'Callaghan (Ed.) NZ Plant Protection Society Inc.
Cotesia rubecula and C. plutellae were assessed as potential biological control agents for Pieris rapae and Plutella xylostella, respectively, in NZ. Host specificity was evaluated by rearing collections of Lepidoptera from natural parasitoid habitats overseas, and by laboratory testing of their host preferences for related Lepidoptera and species from brassica habitats. C. rubecula showed strong preferences for P. rapae and developed in no other species, whereas although C. plutellae demonstrated preferences for P. xylostella in oviposition rate and suitability for development, it could develop in several other Lepidoptera in the laboratory.

Charles, J.G. and Dugdale, J.S. (2011). Non-target species selection for host-range testing of Mastrus ridens. New Zealand Entomologist 34: 45-51
This paper describes the approach taken to selecting non-target species for host-range testing of Mastrus ridens (= M. ridibundus auct.) (Hymenoptera: Ichneumonidae), a proposed biocontrol agent for codling moth. Cydia panatella (Lepidoptera: Tortricidae) in New Zealand. An initial list of potential hosts was developed, derived from a combination of phylogenetic/taxonomic affinity to codling moth, ecological similarity to codling moth, and 'safeguard' or environmental considerations. Species selected are listed in the paper.

Couch K.M., Cresswell A.S., Barratt B.I.P. and Evans A.A. (1997). Implications of host weevil circadian activity for parasitism by Microctonus aethiopoides (Hymenoptera: Braconidae). Pp. 227-231 In: Proceedings of the 50th New Zealand Plant Protection Conference, M. O'Callaghan (Ed.) New Zealand Plant Protection Society Inc.
A laboratory investigation was carried out to determine whether diurnally active non-target weevils may be more susceptible to parasitism than nocturnally active weevils, since it was thought that Microctonus aethiopoides Loan oviposits in its target host primarily during light periods.

Duan J.J. and Messing R.H. (1999). Effects of origin and experience on patterns of host acceptance by the opiine parasitoid Diachasmimorpha tryoni. Ecological Entomology 24: 284-291.
Parasitoid acceptance of less-preferred hosts or host-substrate complexes may be more amenable to conditioning through prior experience (i.e. learning) than preferred host-substrate complexes. The relevance of these findings to host range expansion of parasitoids used in fruit fly biological control is discussed.

Ferkovich S.M. and Blumberg D. (1995). Acceptance of six atypical host species for oviposition by Microplitis croceipes (Hymenoptera: Braconidae). Israel Journal of Entomology 29: 123-131.
Comparisons were made of the acceptance for oviposition by the endoparasitoid, Microplitis croceipes of 6 atypical lepidopteran hosts with 2 typical hosts, Helicoverpa zea and Heliothis virescens. The atypical hosts were Spodoptera frugiperda, S. exigua, Plodia interpunctella, Trichoplusia ni, Galleria mellonella and Plutella xylostella. The acceptability of the atypical hosts for parasitoid oviposition was investigated after treatment of host larvae with H. zea haemolymph, frass, and both. S. frugiperda larvae were significantly more acceptable for oviposition by parasitoid females than the other atypical hosts, when untreated. The H. zea frass plus haemolymph treatment increased the mean number of eggs laid/host across all 6 atypical species.

Gilbert L.E. and Morrison L.W. (1997). Patterns of host specificity in Pseudacteon parasitoid flies (Diptera: Phoridae) that attack Solenopsis fire ants (Hymenoptera: Formicidae). Environmental Entomology 26: 1149-1154.
Pseudacteon spp. that parasitize Solenopsis invicta in South America are not present in the introduced range of this pest species in the USA. Sequential host specificity tests were conducted with 4 South American Pseudacteon species to investigate the degree to which these species attack the native North American S. geminata. Three species showed little interest in ovipositing on S. geminata, but P. curvatus oviposited on S. geminata readily, but there was no larval development. Methods for assaying host specificity and the biocontrol potential of these insects are discussed.

Goldson S.L., McNeill M.R., Phillips C.B. and Proffitt J.R. (1992). Host specificity testing and suitability of the parasitoid Microctonus hyperodae (Hym.: Braconidae, Euphorinae) as a biological control agent of Listronotus bonariensis (Col.: Curculionidae) in New Zealand. Entomophaga 37: 483-498.
Microctonus hyperodae was imported from South America as a potential biological control agent of the adult stage of the pest weevil Listronotus bonariensis. Four non-target weevils were found to sustain some M. hyperodae development but in all but Irenimus aequalis, parasitoid development was impeded, with up to 50% of the larvae becoming encapsulated. I. aequalis was not considered to be threatened by M. hyperodae as this weevil is now recognised as a minor pest. In view of its relatively oligophagous behaviour, the parasitoid was recommended as suitable for release.

Goldson S.L., McNeill M.R., Proffitt J.R. and Barratt B.I.P. (2005). Host specificity testing and suitability of a European biotype of the braconid parasitoid Microctonus aethiopoides Loan as a biological control agent against Sitona lepidus (Coleoptera: Curculionidae) in New Zealand. Biocontrol Science and Technology 15: 791-813.
The paper described host specificity testing for European biotypes of Microctonus aethiopoides Loan. Choice and no-choice tests were carried out. European M. aethiopoides was able to develop in the native weevils Irenimus aequalis, Nicaeana cervina, Catoptes cuspidatus, Protolobus porculus and Steriphus variabilis with parasitism rates of 13, 28, 2, 7 and 8%, respectively. These levels were significantly less than those in the corresponding S. lepidus control. It was concluded that the ecological impact of the European biotype is likely to be less severe than those already exhibited by the Moroccan M. aethiopoides.

Grandgirard J., Hoddle M.S., Petit J.N., Percy D.M. and Roderick G.K. (2006). Pre-introductory risk assessment studies of Gonatocerus ashmeadi (Hymenoptera: Mymaridae) for use as a classical biological control agent against Homalodisca vitripennis (Hemiptera: Cicadellidae) in the Society Islands of French Polynesia. Biocontrol Science & Technology 17: 809-822
Homalodisca vitripennis (Germar)( Hemiptera: Cicadellidae) invaded French Polynesia in 1999. A classical biological control program against H. vitripennis was initiated in 2004 aiming to introduce the exotic egg parasitoid Gonatocerus ashmeadi (Girault) (Hymenoptera: Mymaridae) to the Society Islands archipelago. The primary risk of H. vitripennis is its potential to vector the lethal plant bacterium, Xylella fastidiosa, although its presence in French Polynesia has not yet been demonstrated. Studies assessing the risk of to native cicadellids showed at least 14 cicadellid species were present and the risk to these species from non-taget attack was assessed by examining their phylogenetic relationships to known hosts of G. ashmeadi, their similarity in body size, egg laying biology, and ecology. It was concluded that none of the potential non-taget species were at risk of attack because none are in the tribe Proconiini, all were very small and, appeared to lay tiny single eggs, deposited on the undersides of leaves of trees. These results persuaded the French Polynesian Government that the benefits of establishing G. ashmeadi for H. vitripennis control outweighed the risks. Releases of G. ashmeadi in Tahiti began in May 2005.

Hoffmeister T.S. (2005). From design to analysis: effective statistical approaches for host range testing Pp. 672-682 In: Second International Symposium on Biological Control of Arthropods, Davos, Switzerland, 12-16 September, 2005, M.S. Hoddle (Ed.) United States Department of Agriculture, Forest Service, Washington.
The major goal of host range testing in biological control is to minimize the probability that released biological control agents have unwanted effects on populations of non-target hosts. This paper discusses common problems with experimental designs, emphasizes the need to decide on the statistical effect size that is biologically meaningful, and to determine the statistical power of the host range test employed. An overview is given of appropriate statistical approaches for analysing experiments on the host range of potential biological control agents.

Hopper K.R. and Wajnberg E. (2006). Risks of interbreeding between species used in biological control and native species, and methods for evaluating their occurrence and impact. Pp. 78-97 In: Environmental Impact of Arthropod Biological Control: Methods and Risk Assessment, U. Kuhlmann, F. Bigler and D. Babendreier (Ed.) CABI Bioscience, Delemont, Switzerland.

Jenner WH., Kuhlmann U. (2010). Refining the implementation of arthropod classical biological control. Journal fur Kulturpflanzen 62: 102-106.
The authors are using current biological control projects to tackle problems associated with estimating agent host specificity and risk assessment. These include key host range testing issues for arthropod biolocical control including methods for selection of non-target species, design and implementation of host specificity experiments, and extrapolation of laboratory results to a field context.

Kimber W., Glatz R., Caon G. and Roocke D. (2010). Diaeretus essigellae Stary and Zuparko (Hymenoptera: Braconidae: Aphidiini), a biological control for Monterey pine aphid, Essigella californica (Essig) (Hemiptera: Aphididae: Cinarini): host-specificity testing and historical context Australian Journal of Entomology 49: 377-387

Kuhlmann U., Schaffner U. and Mason P.G. (2005). Selection of non-target species for host specificity testing of entomophagous biological control agents. Pp. 566-583 In: Second International Symposium on Biological Control of Arthropods, Davos, Switzerland, 12-16 September, 2005, M.S. Hoddle (Ed.) United States Department of Agriculture, Forest Service, Washington.
Recommendations are given for setting up test species lists for arthropod biological control programmes are presented. Ecological similarities, phylogenetic/taxonomic affinities and safeguard considerations are applied to ecological host range information to develop an initial test list, which is then filtered by eliminating those with different spatial, temporal and morphological attributes and those species that are not readily obtained. The reduced test list is used for the actual testing but can be revised if new information indicates that more species should be included.

Kuhlmann U., Schaffner U. and Mason P.G. (2006). Selection of non-target species for host specificity testing. Pp. 15-37 In: Environmental impact of invertebrates for biological control of arthropods: methods and risk assessment F. Bigler, D. Babendreier and U. Kuhlmann (Ed.) CABI Publishing, Wallinford, Oxford.

McNeill M.R., Barratt B.I.P. and Evans A.A. (2000). Behavioural acceptability of Sitona lepidus (Coleoptera: Curculionidae) to the parasitoid Microctonus aethiopoides (Hymenoptera: Braconidae) using the pathogenic bacterium Serratia marcescens Bizio. Biocontrol Science and Technology 10: 205-214.
A method was developed to distinguish beween behavioural and physiological barriers to successful parasitism of a host. The insect pathogenic bacterium Serratia marcescens Bizio was placed on the ovipositor of the wasp and used as a marker for parasitoid ovipositor penetration.

McNeill M.R., Vittum R.J. and Jackson T.J. (2000). Serratia marcescens as a rapid indicator of Microctonus hyperodae oviposition activity in Listronotus maculiocollis and potential application of the technique to host-specificity testing. Entomologia Experimentalis et Applicata 95: 193-200.
Listronotus maculicollis (Dietz) (Coleoptera: Curculionidae) is a potential novel host of the braconid parasitoid Microctonus hyperodae Loan, but initial studies have shown that levels of parasitism are lower than in the natural host L. bonariensis (Kuschel). The incidence of ovipositor penetration by the parasitoid M. hyperodae into adult L. maculicollis was measured by immersing the ovipositor of the parasitoid in the facultative pathogen, Serratia marcescens Bizio. Adult weevils were then exposed to parasitoids rapid mortality used as an indicator of oviposition penetration. Application of bacteria to the parasitoid ovipositor provided a rapid, simple test for ovipositor penetration, which shows potential for separation of behavioural and physiological defence mechanisms in parasitoid/host range studies.

McNeill M.R., Withers T.M. and Goldson S.L. (2005). Potential non-target impact of Microctonus aethiopoides Loan (Hymenoptera: Braconidae) on Cleopus japonicus Wingelmüller (Coleoptera: Curculionidae), a biocontrol agent for putative release to control the butterfly bush Buddleja davidii Franchet in New Zealand. Australian Journal of Entomology 44: 201-207.
Cleopus japonicus Wingelmüller (Coleoptera: Curculionidae) is being considered for release to control buddleia Buddleja davidii in New Zealand. As part of the pre-release testing, Moroccan and Irish biotypes of the solitary endoparasitoid Microctonus aethiopoides Loan (Hymenoptera: Braconidae) were evaluated for potential non-target impacts on adult C. japonicus should release occur. Parasitoid behavioural attraction was assessed using the pathenogenic bacterium Serratia marcescens (Enterobactereaceae), as an indicator of ovipositor penetration. Physiological suitability was assessed by comparing parasitism of C. japonicus with the natural hosts of the respective parasitoid biotypes. C. japonicus was found to be behaviourally acceptable to both Moroccan and Irish M. aethiopoides, however, it did not support development of either Moroccan or Irish M. aethiopoides biotypes suggesting that the impact of M. aethiopoides on field populations will be negligible.

Morrison L.W. and Porter S.D. (2006). Post-release host-specificity testing of Pseudacteon tricuspis, a phorid parasitoid of Solenopsis invicta fire ants. Biocontrol 51: 195-205.

Murray T.J., Withers T.M. and Mansfield S. (2010). Choice versus no-choice test interpretation and the role of biology and behavior in parasitoid host specificity tests. Biological Control 52: 2, 153-159.
The need to improve methods and interpretation of host specificity tests for arthropod natural enemies has been clearly identified. In this study, an established exotic host/parasitoid system was used to assess the outcomes and predictive accuracy of no-choice compared to paired choice tests within small laboratory arenas. Host acceptance by two egg parasitoids, Enoggera nassaui and Neopolycystus insectifurax (Pteromalidae), was interpreted in light of percent parasitism, offspring sex ratios and observed parasitoid behavior. Both test designs predicted that D. semipunctata is within the ecological host range of the two parasitoid species, whereas field evidence suggests this is a false positive result. Percent parasitism of all hosts was higher in no-choice compared to choice tests and was predictive of rank order of host preference in choice tests. Presence of the most preferred host did not increase attack on lower ranked hosts. The results supported the assertion that both no-choice and choice tests along with detailed behavioral studies should be conducted for interpretation of pre-release host specificity tests and prediction of field host range.

Neale C., Smith D., Beattie G.A.C. and Miles M. (1995). Importation, host specificity testing, rearing and release of three parasitoids of Phyllocnistis citrella Stainton (Lepidoptera: Gracillariidae) in eastern Australia. Journal of the Australian Entomological Society 34: 343-348.
Three parasitoids of the citrus pest Phyllocnistis citrella, Ageniaspis citricola, Citrostichus phyllocnistoides and Cirrospilus quadristriatus, were tested in quarantine against 17 other insect hosts with leafmining and gall-forming habits. Results showed they were restricted to P. citrella and releases were made in Queensland, New South Wales, South Australia and Victoria.

Orr C.J., Obrycki J.J. and Flanders R.V. (1992). Host acceptance behaviour of Dinocampus coccinellae (Hymenoptera: Braconidae). Annals of the Entomological Society of America 85: 722-730.
Describes observations of behaviour and parasitism of the braconid Dinocampus coccinellae exposed to a number of coccinellids.

Rowbottom R.M., Allen G.R., Walker P.W. and Berndt L.A. (2013). Phenology, synchrony and host range of the Tasmanian population of Cotesia urabae introduced into New Zealand for the biocontrol of Uraba lugens. Biocontrol 58: 625-633.
The population dynamics of Cotesia urabae (Austin and Allen) (Braconidae: Microgastrinae), a biological control agent from Tasmania, and its eucalypt feeding host, Uraba lugens (Walker) (Lepidoptera: Nolidae) was investigated prior to its introduction to New Zealand in 2011. Previous host range testing on potential New Zealand non-targets determined C. urabae had some potential to attack an endemic species, Nyctemera annulata (Boisduval) (Lepidoptera: Arctiidae). A closely related species in Tasmania, Nyctemera amica, was thus investigated as a potential host along with the native host U. lugens, to better understand the host range of C. urabae and the synchrony with its host in Tasmania. Adult C. urabae emerged from pupal cocoons in the field during January which confirmed a five month window in which its host, the larvae of U. lugens, was absent in the field. Experiments using sentinel N. amica and U. lugens larvae, field collections of N. amica and of larvae of other Lepidopteran species during this five month time window detected no parasitism by C. urabae. In the laboratory, host specificity testing showed reduced attack rates and no resultant C. urabae eggs or developing larvae or any successful pupation of C. urabae larvae from attacked N. amica larvae. It was concluded that N. amica is most unlikely to be a host for C. urabae in Tasmania and no evidence of any other alternative host was found.

Sands D.P.A. (1993). Effects of confinement on parasitoid-host interactions: interpretation and assessment for biological control of arthropod pests. Pp. 196-199 In: Pest Control in Sustainable Agriculture, S.A. Corey, D.J. Dall and W.M. Milne (Ed.) CSIRO, Canberra, Australia.

Sands D.P.A. (1998). Guidelines for testing host specificity of agents for biological control of arthropod pests. Pp. 556-560 In: Pest Management - Future Challenges: Proceedings of the 6th Australasian Applied Entomological Research Conference, M. Zalucki, R. Drew and G. White (Ed.) The Cooperative Research Centre for Tropical Pest Management.

Sands D.P.A. and Coombs M.T. (1999). Evaluation of the Argentinean parasitoid, Trichopoda giacomelli (Diptera: Tachinidae), for biological control of Nezara viridula (Hemiptera: Pentatomidae) in Australia. Biological Control 15: 19-24.
Trichopoda giacomellii (Blanchard) (Diptera: Tachinidae), was evaluated prior to its release in Australia as a biological control agent for the green vegetable bug, Nezara viridula (L.) (Hemiptera: Pentatomidae). In no-choice host specificity studies, females of T. giacomellii were exposed in separate tests to selected representatives of indigenous Australian Hemiptera. In addition to the target N. viridula, only species of the 'pentatoma' group of Pentatomidae, Plautia affinis Dallas, Alciphron glaucas (Fabricius), and Glaucias amyoti (White), attracted oviposition and supported complete development by T. glacomellii. Other species attracted oviposition but parasitoids failed to develop, and others failed to attract oviposition.

Stufkens M.W., Farrell J.A. and Popay A.J. (1994). Quarantine host range tests on two exotic parasitoids imported for aphid control. Pp. 149-153 In: Proceedings of the 47th New Zealand Plant Protection Conference.
Aphidius sonchi was imported from Australia as a potential biological control agent for Hyperomyzus lactucae and Ephedrus cerasicola from Norway as a potential control agent of Myzus persicae. The rate of parasitism of 17-20 aphid species was determined. E. cerasicola parasitized 5 native aphids and was not released. A. sonchi did not attack any native aphid species or their close relatives, so release of this parasitoid was approved.

Todd J.H., Barratt B.I.P., Tooman L., Beggs J.R. and Malone L.A. (2015). Selecting non-target species for risk assessment of entomophagous biological control agents: Evaluation of the PRONTI decision-support tool. Biological Control 80: 77-88.

Toepfer S., Zhang F. and Kuhlmann, U. (2009). Assessing host specificity of a classical biological control agent against western corn rootworm with a recently developed testing protocol. Biological Control. 51(1): 26-33.
The authors describe host range testing of Celatoria compressa (Wulp) (Diptera: Tachinidae) for control of Diabrotica virgifera virgifera LeConte (Coleoptera: Chrysomelidae: Galerucinae) in Europe. Nine potential non-target beetles were tested in no-choice tests, sequential no-choice tests, choice tests and sequential choice tests in small experimental arenas in a quarantine laboratory. The nine species were selected based on (1) ecological host range information of C. compressa, (2) ecological similarities to D. v. virgifera, (3) close phylogenetic/taxonomic relationships, (4) safeguard considerations, (5) morphological similarities, geographical distributions, overlap of temporal occurrences with D. v. virgifera and C. compressa, and (6) accessibility and availability. C. compressa only parasitized a few red pumpkin beetles, Aulacophora foveicollis (Chrysomelidae: Galerucinae), regardless of the presence or absence of D. v. virgifera but preferred D. v. virgifera (44.6% parasitized) over A. foveicollis (2.7%) in choice tests. The authors concluded that C. compressa has a fundamental host range restricted to the subtribes Diabroticina and Aulacophorina, and would therefore be unlikely to have a direct impact on indigenous species in Europe.

Withers T.M., Allen G.R. and Reid C.A.M. (2015). Selecting potential non-target species for host range testing of Eadya paropsidis. New Zealand Plant Protection 68: 179-186.

van Driesche R. (2004). Predicting host ranges of parasitoids and predacious insects - what are the issues? Pp. 1-3 In: Assessing host ranges for parasitoids and predators used for classical biological control: a guide to best practice, R. Van Driesche and R. Reardon (Ed.) USDA Forest Service, Morgantown, West Virginia.

van Driesche R. and Reardon R. (2004). Assessing host ranges for parasitoids and predators used for classical biological control: a guide to best practice. Pp. 243. USDA Forest Service, Morgantown, West Virginia.

van Driesche R.G. and Hoddle M. (1997). Should arthropod parasitoids and predators be subject to host range testing when used as biological control agents? Agriculture and Human Values 14: 211-226.
It is questioned whether agents introduced for arthropod biological control should be subjected to host range testing before release, and if so, are methods used for estimating host ranges of herbivorous arthropods appropriate, or are different approaches needed. Current examples in which host range testing has been employed for arthropod biological control are reviewed.

van Lenteren J.C., Bigler F., Burgio G., Hokkanen H.M.T. and Thomas M.B. (2002). Risks of importation and release of exotic biological control agents: how to determine host specificity. IOBC/wprs Bulletin 25: 281-284.
Many exotic natural enemies have been imported, mass reared and released as biological control agents for greenhouse pests. Negative effects of these releases for greenhouse biological control have not been reported yet. However, an increasing number of projects will be executed by persons not trained in identification, evaluation and release of biological control agents. A working group of OECD is developing a guidance document for registration requirements of exotic natural enemies. In this paper, the state of affairs concerning these developments is summarized.

van Lenteren J.C., Cock M., Hoffmeister T.S. and Sands D. (2005). Host ranges of natural enemies as an indicator of non-target risk. Pp. 584-592 In: Second International Symposium on Biological Control of Arthropods, Davos, Switzerland, 12-16 September, 2005, M.S. Hoddle (Ed.) United States Department of Agriculture, Forest Service, Washington.
The introduction of exotic natural enemies or mass release of biological control agents may lead to unwanted non-target effects, depending upon the host range of the biological control agent and the presence of non-target species in the area of release. Host specificity testing information is probably the most important and the easiest for regulators to use. A framework for stepwise host range testing with levels of increasing complexity that should allow avoidance of over- and underestimation of the host range is presented. The interpretation of data obtained from host range testing is discussed.

van Lenteren J.C., Cock M.J.W., Hoffmeister T.S. and Sands D.P.A. (2006). Host specificity in arthropod biological control, methods for testing and interpreting the data. Pp. 38-63. CAB Publishing, Delemont.