Aeschlimann J.P. (1983).
Notes on the variability of Microctonus aethiopoides Loan (Hymenoptera: Braconidae: Euphorinae).
Contributions of the American Entomological Institute 20: 329-335.
The authory describes some characteristics of Mediterranean biotypes of the parasitoid Microctonus aethiopoides Loan. Laboratory experiments and field observations have shown that that there are several geographical as well as host-associated biotypes of the parasite.
Aeschlimann J.P. (1989).
On the importance of assessing the quality of beneficial organisms mass-produced for use in biological control programmes against noxious plants and animals.
Boletin Sanidad Vegetal, Fuera de Serie No. 17: 377-382.
Examples are given illustrating the importance of continuous quality assessment of biological control agents, which have been mass-reared for field use, are described. Examples include the curculionid Rhinocyllus conicus which attacks thistle species and braconids in the genus Microctonus used to control curculionids.
Aeschlimann J.P. (1995). Lessons from post-release investigations in classical biological control: the case of Microctonus aethiopoides Loan (Hym., Braconidae) introduced into Australia and New Zealand for the biological control of Sitona discoideus Gyllenhal (Col., Curculionidae). Pp. 75-83 In: Biological Control: Benefits and Risks, H.M.T. Hokkanen and J.M. Lynch (Ed.) Cambridge University Press, Cambridge, UK.
Alaphilippe A., Elad Y., David D.R., Derridj S. and Gessler C. (2008).
Effects of a biocontrol agent of apple powdery mildew (Podosphaera leucotricha) on the host plant and on non-target organisms: an insect pest (Cydia pomonella) and a pathogen (Venturia inaequalis).
Biocontrol Science and Technology 18: 121-138
Non-target effects of sprayed applications of a potential biocontrol agent, an epiphytic yeast isolate called Y16, of apple powdery mildew Podosphaera leucotricha Ell. Et Ev.), on scab infections (Venturia inaequalis Cooke Winter), on codling moth oviposition and damage and apple quality were examined. The BCA affected neither conidia germination of V. inaequalis nor their penetration of the leaf tissue but suppressed the disease caused by this pathogen. The quantity of eggs laid by the codling moth during its second flight period on yeast treated trees was significantly different, but inconsistent from year to year, the differences attributed to year-to-year variation in environmental conditions, which may affect yeast survival and activity. A 2-month-long assay was conducted in the orchard during the codling moth's second flight period from mid-July until mid-September. The yeast treatment did not affect the damage caused by the codling moth to the fruits or any of the examined fruit quality parameters.
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.
Andersen M.C., Ewald M. and Northcott J. (2005).
Risk analysis and management decisions for weed biological control agents: Ecological theory and modelling results.
Biological Control 35: 330-337.
The risks posed by weed biological control agents, and a simple model of herbivorous insect movement and oviposition on two species of host plant, a target invasive plant species and a non-target native species, in simulated landscapes is discussed. The model shows that risks of non-target impacts may be influenced by movement behaviour of biological control agents in heterogeneous landscapes. The authors conclude that such models should be considered as part of a comprehensive strategy of risk assessment for proposed weed biological control agents.
Andreas J.E., Schwarzlander M. and Clerck-Floate R.d (2008).
The occurrence and potential relevance of post-release, nontarget attack by Mogulones cruciger, a biocontrol agent for Cynoglossum officinale in Canada.
Biological Control 46: 304-311
The root-mining weevil Mogulones cruciger, was approved and released in Canada, but was not approved for release in the United States, to control Cynoglossum officinale. Confamilial species co-occurring with C. officinale at six M. cruciger release sites in Alberta and British Columbia were assessed over a two year period. All four co-occurring species were attacked by the weevil to varying degrees, although attack was inconsistent between years and sites and nontarget species were attacked to a lesser degree than C. officinale. There was a positive relationship between the probability of nontarget attack and C. officinale attack rate by M. cruciger suggesting that the immigration of M. cruciger into the US may expose certain Boraginaceae to nontarget attack, but the risk to native species is unknown.
Araj S.A., Wratten S.D., Lister A.J. and Buckley H.L. (2006). Floral nectar affects longevity of the aphid parasitoid Aphidius ervi and its hyperparasitoid Dendrocerus aphidum. New Zealand Plant Protection 59: 178-183.
Arnett A.E. and Louda S.M. (2002).
Re-test of Rhinocyllus conicus host specificity, and the prediction of ecological risk in biological control.
Biological Conservation 106: 251-257.
Rhinocyllus conicus was released in North America to control exotic thistles. It is now reducing seed production by multiple native North American thistle species. It was hypothesized that host specificity of R. conicus has changed since pre-release testing, providing an explanation for the unexpected magnitude of the documented ecological effects, however, this was not the case. The authors concluded that accurate prediction of the potential level of impact on native host plants in the field requires further ecological information in addition to host specificity.
Ash G.J., Chung Y.R., McKenzie C. and Cother E.J. (2008).
A phylogenetic and pathogenic comparison of potential biocontrol agents for weeds in the family Alismataceae from Australia and Korea.
Australasian Plant Pathology 37: 402-405
Plants in the family Alismataceae are weeds of rice in Australia and Korea. Research programs have investigated the use of inundative plant-pathogenic fungi for biological control of these weeds. Recent studies have shown a close phylogenetic relationship between the organisms under investigation in the two countries. A survey of Alismataceae weeds in southern South Korea was carried out and fungal isolations were made from the diseased specimens. The isolates overlapped between those previously described in Korea as Plectosporium tabacinum and the newly named P. alismatis. The host range testing on australian weeds in the glasshouse showed that the isolates from Korea were less pathogenic than the Australian isolates. Therefore, although the isolates were phylogenetically related, the isolates from Korea did not show greater virulence or a wider or different host range than the Australian isolates.
Askew R.R. (1994). Parasitoids of leaf-mining Lepidoptera: what determines their host ranges? Pp. 177-202 In: Parasitoid community ecology, B.A. Hawkins and W. Sheehan (Ed.) Oxford University Press, Oxford
Askew R.R. and Shaw M.R. (1986). Parasitoid communities: their size, structure and development. Pp. 225-264 In: Insect parasitoids, J.K. Waage and D.J. Greathead (Ed.) Academic Press, London.
Asquith A. and Miramontes E. (2001). Alien parasitoids in native forests: the ichneumonoid wasp community in Hawaiian rainforest. Pp. 54-67 In: Balancing nature: assessing the impact of importing non-native biological control agents (an international perspective), J.A. Lockwood, F.G. Howarth and M. Purcell (Ed.) Entomological Society of America, Lanham, Maryland.
Aubert B. and Quilici S. (1983). Nouvel équilibre biologique observe à la Réunion sur les populations de psyllides après l’introduction et l’éstablissement d’hymenopteres chalcidiens. Fruits 38: 771-780.
Baars J.R. (2000).
Emphasizing behavioural host-range: the key to resolving ambiguous host-specificity results on Lantana camara L.
Proceedings of the X International Symposium on Biological Control of Weeds: 887-896
Candidate biological control agents presently under evaluation for release on L. camara in South Africa accept closely related native plant species. The results from host-range results of two natural enemies, Falconia intermedia (Hemiptera: Miridae) and Coelocephalapion sp. (Coleoptera: Brentidae) were compared to determine the influence these trials have on the interpretation of the accepted host-range. Results suggested that the natural host-range of a candidate biological control agent is best determined by focusing on behavioural factors influencing host acceptance. The implications of using trials that incorporate insect behaviour during host-specificity screening and risk analysis are discussed.
Babendreier D. and Bigler F. (2005).
How to assess non-target effects of polyphagous biological control agents: Trichogramma brassicae as a case study.
Pp. 603-610 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 risk assessment conducted for Trichogramma brassicae (Hymenoptera: Trichogrammatidae), an egg parasitoid used for control of the European corn borer in European countries is discussed. The main factors investigated were the potential of establishment, acceptance and parasitism of non-target butterflies under laboratory, field-cage and field conditions, the searching efficiency in non-target habitats, the dispersal capacities and the potential for effects on other natural enemies in maize. Although high parasitism of non-target butterflies and other natural enemies were observed under laboratory conditions, very few eggs of non-target species were attacked in the field, possibly because of low host searching efficiency, and limited parasitoid dispersal. It was concluded that the possibility of using invertebrate agents with a broad host range in inundative biological control should not be excluded, although a thorough environmental risk assessment should be performed prior to release.
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., Bigler F. and Kuhlmann U. (2006). Current status and constraints in the assessment of non-target effects. Pp. 1-13 In: Environmental impact of invertebrates for biological control of arthropods – methods and risk assessment, F. , F. Bigler, D. Babendreier and U. Kuhlmann (Ed.) CABI Publishing, Wallingford, UK
Babendreier D., Kuske S. and Bigler F. (2003).
Parasitism of non-target butterflies by Trichogramma brassicae Bezdenko (Hymenoptera: Trichogrammatidae) under field cage and field conditions.
Biological Control 26: 139-145.
The egg parasitoid Trichogramma brassicae has been inundatively released to control the European corn borer, Ostrinia nubilalis Hübner, in maize. Non-target parasitism of butterfly eggs by T. brassicae in field cages and under field conditions in Switzerland was investigated. Although the tested non-target butterflies were all attacked under semi-field and field conditions, it was concluded that effects on non-target butterflies due to mass released T. brassicae are minimal.
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.
Bailey, K.L., Pitt, W.M., Falk, S. and Derby, J. (2011). The effects of Phoma macrostoma on nontarget plant and target weed species. Biological Control 58: 379-386
Balciunas J.K. (2004).
Are mono-specific agents necessarily safe? The need for pre-release assessment of probable impact of candidate biocontrol agents, with some examples.
Proceedings of the XI International Symposium on Biological Control of Weeds: 252-257
Biosafety is now often considered more important than efficacy. However, even a highly specific agent can have unpredictable adverse impacts especially if it becomes abundant on the target, but fails to reduce target weed populations. However, pre-release consideration of the proposed agent's probable efficacy is receiving increased attention. This is usually done overseas, in the native range of both the target weed and candidate agent and approaches used are reviewed. Pre-release impact assessments can also be performed in quarantine. The results of two "dosage" trials conducted with a gall-making fly that is being considered as a biological control agent for Cape ivy (Delairea odorata) are described. Plants exposed to both low and high densities of gall flies, were smaller, and had fewer leaves than the ungalled controls. Pre-release evaluations of a candidate agent's potential impact should lead to fewer ineffective agents being released, thereby making weed biocontrol more efficient, and reducing the possibility of negative indirect impacts on non-targets.
Balciunas J.K. and Villegas B. (2007).
Laboratory and realized host ranges of Chaetorellia succinea (Diptera : Tephritidae), an unintentionally introduced natural enemy of yellow starthistle.
Environmental Entomology 36: 849-857
In 1999, Chaetorellia succinea (Costa) (Diptera: Tephritidae), an unintentional introduction from Greece, was considered for biocontrol of yellow starthistle, Centaurea solstitialis L., one of the worst weeds in the western United States. However, the host range of C. succinea had not been studied, and so the physiological host range was determined in the laboratory by exposing it under no-choice conditions to 14 potential Cardueae hosts. Two introduced weed species and the native American basketflower (Centaurea americana Nuttall) were found to be laboratory hosts, although yellow starthistle was highly preferred. Because Ch. succinea is already widespread throughout California, flower heads were collected from 24 potential host plant species in the field to determine the realized host range. Ch. succinea emerged only from the other two known hosts: Ce. melitensis and Ce. sulfurea. Our results suggest that American basketflower growing in the southwestern United States may also be at risk if Ch. succinea expands its range into that region.
Balciunas J.K., Burrows D.W. and Purcell M.F. (1996). Comparison of the physiological and realized host-ranges of a biological control agent from Australia for the control of the aquatic weed, Hydrilla verticillata. Biological Control 7: 148-158.
Bale, J. (2011).
Harmonization of regulations for invertebrate biocontrol agents in Europe: progress, problems and solutions.
Journal of Applied Entomology 135: 503–513.
This article reviews major developments in the regulation and environmental risk assessment of insect biocontrol agents in Europe over the last 10 years including: the fragmented pattern of regulation between countries, variation in information requirements for release licences, format and methods of environmental risk assessment for different taxonomic groups, use and updating of the European Plant Protection Organisation Positive List, sources of expert advice, communication between regulators, and options for the provision of international leadership to coordinate regulatory issues with biocontrol in Europe.
Bangsund D.A., Leistritz F.L. and Leitch J.A. (1999). Assessing economic impacts of biological control of weeds: the case of leafy spurge in the northern Great Plains of the United States. Journal of Environmental Management. 1999. 56: 1, 35-43 56: 35-43.
Barbosa P. (1998). Conservation biological control. Academic Press, London.
Barlow N.D. (1999).
Models in biological control: a field guide
Pp. 43-70 In: Theoretical approaches to biological control, B.A. Hawkins and H.V. Cornell (Ed.) Cambridge University Press UK.
This paper reviews a number of examples of models for biological control programmes, but notes that the same principles can be applied to non-target as wellas target species.
Barlow N.D. and Goldson S.L. (1993). A modelling analysis of the successful biological control of Sitona discoideus (Coleoptera: Curculionidae) by Microctonus aethiopoides (Hymenoptera: Braconidae) in New Zealand. Journal of Applied Ecology 30: 165-179
Barlow N.D., Barratt B.I.P., Ferguson C.M. and Barron M.C. (2004).
Using models to estimate parasitoid impacts on non-target host abundance.
Environmental Entomology 33: 941-948.
A method is described for estimating the impact of a parasitoid on the abundance of a nontarget host, using the intrinsic rate of host increase, the average abundance of the host in the presence of parasitism, and the estimated mortality caused by the parasitoid. The method is applied to the braconid Microctonus aethiopoides Loan, which is known to attack native weevils. The non-target host population was modelled using discrete Ricker or continuous logistic models, tuning the models to host population data in the presence of parasitism, then removing parasitism and determining the increase in predicted equilibrium host density. In an area where up to 30% parasitism of a nontarget host population has been recorded, the model estimated an 8% reduction of the nontarget host, but in another area, where the parasitoid has not established, the method was applied in reverse to predict the parasitoid's impact if it did establish. In this case, the model predicted a 30% suppression of population density, the host's intrinsic rate of increase, rm, accounting for this difference in predicted impact.
Barlow N.D., Goldson S.L. and McNeill M.R. (1994). A prospective model for the phenology of Microctonus hyperodae (Hymenoptera: Braconidae), a potential biological control agent of Argentine stem weevil in New Zealand. Biocontrol Science and Technology 4: 375-386.
Barlow N.D., Kean J.M. and Goldson S.L. (2002). Biological control lessons from modeling of New Zealand successes and failures. Proceedings of the First International Symposium on Biological Control of Arthropods: 105-107.
Barratt B.I.P. (1996). Biological control: Is it environmentally safe? Forest and Bird 282: 36-41.
Barratt B.I.P. (2002). Risks of Biological Control. Pp. 720-722 In: Encyclopaedia of Pest Management, D. Pimental (Ed.) Marcel Dekker Inc, New York.
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. and Kuhlmann U. (2005). Introduction: legislation and biological control of arthropods: challenges and opportunities. Pp. 683-685 In: Second International Symposium of Biological Control of Arthropods, M. Hoddle (Ed.) USDA Forest Service FHTET-2005-08.
Barratt B.I.P. and Moeed A. (2005).
Environmental safety of biological control: policy and practice in New Zealand.
Biological Control 35: 247-252.
The regulatory system for biological control agent introduction in NZ and the process by which biological control applications are received and processed is described. Two case studies of weed biological control agents which have been through the HSNO process, and the scientific issues that arose in considering the environmental safety of these agents are discussed.
Barratt B.I.P., Blossey B. and Hokkanen H.M.T. (2006).
Post-release evaluation of non-target effects of biological control agents.
Pp. 166-186 In: Environmental Impact of Arthropod Biological Control: Methods and Risk Assessment, U. Kuhlmann, F. Bigler and D. Babendreier (Ed.) CABI Bioscience, Delemont, Switzerland.
This paper deals with weed biological control agents, pathogens and parasitoids.
Barratt B.I.P., Evans A.A. and Ferguson C.M., (1997). Potential for control of Sitona lepidus Gyllenhal by Microctonus spp. New Zealand Plant Protection 50: 37-40
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., Evans A.A., Ferguson C.M., McNeill M.R. and Addison P. (2000).
Phenology of native weevils (Coleoptera: Curculionidae) in New Zealand pastures and parasitism by the introduced braconid, Microctonus aethiopoides Loan (Hymenoptera: Braconidae).
New Zealand Journal of Zoology 27: 93-110.
The phenology of native weevils pasture sites in Otago, Canterbury and Waikato was studied by monthly sampling to record reproductive status and incidence of parasitism by introduced braconid parasitoids in the genus Microctonus. Most parasitism occurred after the main reproductive period of entimine weevils in spring, but a putative second generation in some species might be more affected by parasitoid attack. A native rhytirhinine species, Steriphus variabilis, differed from the entimines because adults emerged in autumn and spring, and may be bivoltine.
Barratt B.I.P., Evans A.A., Ferguson C.M., McNeill M.R., Proffitt J.R. and Barker G.M. (1998).
Curculionoidea (Insecta: Coleoptera) of agricultural grassland and lucerne as potential non-target hosts of the parasitoids Microctonus aethiopoides Loan and Microctonus hyperodae Loan (Hymenoptera: Braconidae).
New Zealand Journal of Zoology 25: 47-63.
The paper describes a survey of the weevil fauna of pasture, lucerne and modified native grassland in parts of the southern South Island, Canterbury and the northern North Island of New Zealand, where the parasitoids Microctonus spp. are present to identify weevils with taxonomic and ecological affinities with the target hosts, and hence, potential non-target hosts.
Barratt B.I.P., Evans A.A., Stoltz D.B., Vinson S.B. and Easingwood R. (1999). Virus-like particles in the ovaries of Microctonus aethiopoides Loan (Hymenoptera: Braconidae), a parasitoid of adult weevils (Coleoptera: Curculionidae). Journal of Invertebrate Pathology 73: 182-188.
Barratt B.I.P., Ferguson C.M. and Evans A.A. (2001). Non-target effects of introduced biological control agents and some implications for New Zealand. Pp. 41-53 In: Balancing Nature: Assessing the Impact of Importing Non-Native Biological Control Agents (An International Perspective), J.A. Lockwood, F.G. Howarth and M.F. Purcell (Ed.) Thomas Say Publications, Maryland.
Barratt B.I.P., Ferguson C.M., Bixley A.S., Crook K.E., Barton D.M. and Johnstone P.D. (2007).
Field parasitism of nontarget weevil species (Coleoptera : Curculionidae) by the introduced biological control agent Microctonus aethiopoides Loan (Hymenoptera : Braconidae) over an altitude gradient.
Environmental Entomology 36: 826-839
The parasitoid, Microctonus aethiopoides Loan (Hymenoptera: Braconidae) was introduced into New Zealand in 1982 to control the alfalfa pest, Sitona discoideus Gyllenhal (Coleoptera: Curculionidae). Studies have shown that a number of nontarget weevil species are attacked in the field by this parasitoid. A field study was carried out over 6 years to investigate nontarget parasitism by M. aethiopoides over an altitudinal sequence from the target host habitat (alfalfa) into native grassland. Weevil densities were estimated, species identified, and dissections carried out to determine reproductive status and parasitism. Seven nontarget weevil species were found to be parasitized. Substantial nontarget parasitism was found at only one of the three locations, with up to 24% parasitism of a native weevil, Nicaeana fraudator Broun (Coleoptera: Curculionidae), recorded. Results are discussed in relation to weevil phenology.
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., Goldson S.L., Ferguson C.M., Phillips C.B. and Hannah D.J. (2000). Predicting the risk from biological control agent introductions: A New Zealand approach. Pp. 59-75 In: Nontarget effects of biological control introductions, P.A. Follett and J.J. Duan (Ed.) Kluwer Academic Publishers, Norwell, Massachusetts, USA.
Barratt B.I.P., Howarth F., Withers T.M., Kean J.M. and Ridley G. (2010).
Progress in risk assessment for classical biological control.
Biological Control 52: (3) 245-254.
Controversy in the 1980s about the biosafety of biological control created tension between biological control practitioners and those concerned about non-target impacts. Research has addressed a number of questions which have subsequently led to a greater understanding of risk assessment and biosafety. Advances in quarantine host range testing have improved our ability to predict post-release impacts; pre- and post-release studies are increasingly involving population models to estimate the population impact of introduced biological control agents; regulators have access to accumulating data from past introductions to validate earlier decisions. Progress in research and regulation of biological control are discussed with particular reference to Australasia.
Barratt B.I.P., Murney R., Easingwood R., Ward V.K. (2006).
Virus-like particles in the ovaries of Microctonus aethiopoides Loan (Hymenoptera: Braconidae): comparison of biotypes from Morocco and Europe.
Journal of Invertebrate Pathology 91: 13-18.
Virus-like particles (MaVLP) have been discovered in the ovarial epithelial cells of the solitary, koinobiont, endoparasitoid, Microctonus aethiopoides Loan (Hymenoptera: Braconidae) introduced to New Zealand originally from Morocco to control the lucerne pest Sitona discoideus Gyllenhal (Coleoptera: Curculionidae). MaVLP have been found in all females examined. It has been suggested, although not demonstrated, that like many other such VLP found in parasitoids, MaVLP might play a role in host immunosuppression. Since another biotype of M. aethiopoides from Ireland has been proposed for introduction to control the white clover pest, Sitona lepidus Gyllenhal, in New Zealand, it was considered that females from this biotype warranted transmission electron microscope examination for VLP. No VLP were observed in ovarian tissues of specimens collected from three diVerent locations in Ireland. Similarly, none were found in M. aethiopoides sourced from France, Wales, and Norway. These observations are discussed in relation to quarantine host speciWcity tests with the Irish biotype, which found that the host range of the Irish biotype is likely to be less extensive than that of the Moroccan biotype already in New Zealand.
Barratt B.I.P., Oberprieler R.G., Ferguson C.M. and Hardwick S. (2005).
Parasitism of the lucerne pest Sitona discoideus Gyllenhal (Coleoptera: Curculionidae) and non-target weevils by Microctonus aethiopoides Loan (Hymenoptera: Braconidae) in south-eastern Australia, with an assessment of the taxonomic affinities of non-target hosts of M. aethiopoides recorded from Australia and New Zealand.
Australian Journal of Entomology 44: 192-200.
A survey of weevils found in and near lucerne in south-eastern Australia was carried out to investigate whether similar non-target parasitism was occurring in Australia as in NZ. A single incidence of parasitism of a species of an Australian native weevil Prosayleus sp. by M. aethiopoides was recorded. No parasitism of any other weevil species was observed. The taxonomic affinities between Sitona and native Australian and New Zealand weevils are discussed, concluding that non-target host range in M. aethiopoides may be determined more by ecological factors than by taxonomic affinities among its hosts.
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.
Barratt B.I.P., Todd J.H., Ferguson C.M., Crook K., Burgess E.P.J., Barraclough E.I. and Malone L.A. (2013). Biosafety testing of genetically modified ryegrass (Lolium perenne L.) plants using a model for the optimum selection of test invertebrates. Environmental Entomology 42: 820-830
Barratt BIP, Oberprieler RG, Barton D, Mouna M, Stevens M, Alonso-Zarazaga MA, Vink CJ and Ferguson CM. (2012). Could research in the native range, and non-target host range in Australia, have helped predict host range of the parasitoid Microctonus aethiopoides Loan (Hymenoptera: Braconidae), a biological control agent introduced for Sitona discoideus Gyllenhal (Coleoptera: Curculionidae) in New Zealand? Biocontrol 57: 735-750.
Barratt, B.I.P. (2011).
Assessing safety of biological control introductions.
CAB Reviews: Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources 6 1-12
A review summarising the biosafety debate, and characterising the direct and indirect risks of biological control mainly for weeds and insect pests.
Barron M.C. (2007).
Retrospective modelling indicates minimal impact of non-target parasitism by Pteromalus puparum on red admiral butterfly (Bassaris gonerilla) abundance.
Biological Control 41: 53-63
There is anecdotal evidence that populations of the New Zealand endemic red admiral butterfly Bassaris gonerilla (F.) have declined since the early 1900s as a result of the introduction of the generalist pupal parasitoids Pteromalus puparum (L.) and Echthromorpha intricatoria (F.). A discrete-time model for B. gonerilla population dynamics was constructed which suggested that the impact of non-target parasitism by P. puparum has been minimal, but that parasitism by E. intricatoria was estimated to have caused 30% suppression of B. gonerilla abundance. The model suggested that the presence of an overwintering larval generation of B. gonerilla provides a temporal refuge from the high levels of E. intricatoria parasitism, assuming that parasitism rates are independent of B. gonerilla density. This assumption appears valid for P. puparum parasitism, but may not be so for E. intricatoria.
Barron M.C., Barlow N.D. and Wratten S.D. (2003).
Non-target parasitism of the endemic New Zealand red admiral butterfly (Bassaris gonerilla) by the introduced biological control agent Pteromalus puparum.
Biological Control 27: 329-335.
The New Zealand red admiral butterfly has long been recognised as a non-target host for the introduced biological control agent Pteromalus puparum but its impact has never been quantified. Data were collected to construct a partial life table for B. gonerilla. Egg parasitism by an unidentified Telenomus (scelionid) was 95%. P. puparum parasitized 14% of B. gonerilla pupae sampled. However, pupal parasitism by the self-introduced Echthromorpha intricatoria (F.) (Hymenoptera: Ichneumonidae), was higher at 26%. It is concluded that P. puparum has permanently enhanced mortality in B. gonerilla, but the level of mortality is low relative to egg parasitism by Telenomus sp. and pupal mortality due to E. intricatoria parasitism.
Barton J.E. (2004). How good are we at predicting the field host-range of fungal pathogens used for classical biological control of weeds? Biological control in agricultural IPM systems 31: 99-122
Barton J.E., Fowler S.V., Gianotti A.F., Winks C.J., de Beurs M., Arnold G.C. and Forrester G. (2007). Successful biological control of mist flower (Ageratina riparia) in New Zealand: Agent establishment, impact and benefits to the native flora. Biological Control 40: 370-385
Barton, J. (2012).
Predictability of pathogen host range in classical biological control of weeds: an update.
Biocontrol 57: 289-305
The author reports that to-date 28 species of fungi have been released as classical biological control agents against weeds world-wide. These pathogens have been reported infecting only six non-target plant species outdoors and all of these incidents were predicted. More non-target plant species developed disease symptoms in glasshouse tests than in the field.
Barton-Browne L. (1995). Ontogenetic changes in feeding behavior. Pp. 307-342 In: Regulatory Mechanisms in Insect Feeding, R.F. Chapman and G. de Boer (Ed.) Chapman and Hall.
Barton-Browne L. and Withers T.M. (2002). Time-dependent changes in the host-acceptance threshold of insects: implications for host specificity testing of candidate biological control agents. Biocontrol Science and Technology 12: 677-693.
Beard J.J. and Walter G.H. (2001). Host plant specificity in several species of generalist mite predators. Ecological Entomology 26: 562-570
Beckage N.E. and Gelman D.B. (2004). Wasp parasitoid disruption of host development: Implications for new biologically based strategies for insect control. Annual Review of Entomology 49: 299-330
Beirne B.P. (1975). Biological control attempts by introductions against pest insects in the field in Canada. The Canadian Entomologist 107: 225-236.
Bellows T.S. and Fisher T.W. (1999). Handbook of Biological Control: Principles and Applications of Biological Control. Academic Press, San Diego.
Benson J., Pasquale A., Van Driesche R. and Elkinton J.S. (2003). Assessment of risk posed by introduced braconid wasps to Pieris virginiensis, a native woodland butterfly in New England. Biological Control 26: 83-93
Berenbaum M.R. and Zangerl A.R. (1992). Genetics of physiological and behavioral resistance to host furanocoumarins in the parsnip webworm. Evolution 46: 1373-1384.
Berg G., Grosch R. and Scherwinski K. (2007).
Risk assessment for microbial antagonists: are there effects on non-target organisms?
Gesunde Pflanzen 59: 107-117
Biological control of phytopathogenic fungi using antagonistic microorganisms is potentially environmentally friendly but possible non-target effects on ecologically important soil-microbes need to be considered. Serratia plymuthica HRO-C48 and Streptomyces sp. HRO-71 were applied to control the pathogen Verticillium dahliae on strawberry and potato, and the bacterial strains Pseudomonas trivialis 3Re2-7, P. fluorescens L13-6-12, S. plymuthica 3Re4-18 and the fungal antagonists Trichoderma reesei G1/8 and T. viride G3/2 were introduced to control Rhizoctonia solani on lettuce and potato. After BCA treatment we did not observe any long-term effect on the plant-associated microbes in any tested pathosystem. Therefore, no sustainable risks could be seen for the indigenous micro-organisms.
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.
Berner D.K. (2010).
BLUP, a new paradigm in host-range determination.
Biological Control 53: 143-152
There has been increased focus on modernizing the approach from centrifugal phylogenetic testing to basing selection of test plants on molecular phylogeny rather than taxonomic classification. Mixed model equations (MME) and best linear unbiased predictors (BLUPs) have been used to determine the probable host-range of plant pathogens proposed for biological control of Russian thistle. The work focuses on evaluating disease severity on related plant species although the author describes how MME can be used with any biological weed control agent or target as long as the evaluation criterion is quantitative and variances and molecular genetic relationships among test species can be obtained. The author's objectives are to familiarize biological control researchers and regulators with some of the requirements and advantages of the MME and the use of the MME to construct test plant lists.
Berner D.K., Bruckart W.L., Cavin C.A., Michael J.L., Carter M.L. and Luster D.G. (2009).
Best linear unbiased prediction of host-range of the facultative parasite Colletotrichum gloeosporioides f. sp salsolae, a potential biological control agent of Russian thistle.
Biological Control 51(1): 158-168.
Russian thistle or tumbleweed (Salsola tragus L.) is an introduced, widely distributed invasive weed in N. America. The fungus Colletotrichum gloeosporioides (Penz.) Penz. & Sacc. in Penz. f. sp. salsolae (CGS) is a facultative parasite being considered for classical biological control of this weed. Host-range tests were conducted with 92 accessions from 19 families of plants including 62 genera and 120 species. Disease reaction data were combined with a relationship matrix derived from internal transcribed spacer DNA sequences and analyzed with mixed model equations to produce Best Linear Unbiased Predictors (BLUPs) for each species. Twenty-nine species (30 accessions) from seven closely-related Chenopodiaceae tribes had significant levels of disease severity as indicated by BLUPs, compared to six species determined to be susceptible with least squares means estimates. Of the 29 susceptible species, 10 native or commercially important species in N. America were identified as needing additional tests to determine the extent of any damage caused by infection.
Bigler F., Babendreier D. and Kuhlmann U. (2006). Environmental impact of arthropod biological control: methods and risk assessment. Pp. 288. CABI Publishing, Delemont, Switzerland.
Bigler, F., Babendreier, D. and van Lenteren, J.C. (2010). Risk Assessment and non-target effects of egg parasitoids in biological control. Pp. 413-442 In: Egg Parasitoids in Agroecosystems with Emphasis on Trichogramma, F. L. Consoli, J. R. P. Parra and R. A. Zucchi (Eds.) Springer, Dortrecht, Netherlands.
Blossey B. (1995). Host specificity screening of insect biological control agents as part of an environmental risk assessment. Pp. 84-89 In: Biological Control: Benefits and Risks, H.M.T. Hokkanen and J.M. Lynch (Ed.) Cambridge University Press, Cambridge, UK.
Blossey B. (1999). Before, during and after: the need for long-term monitoring in invasive plant species management. Biological Invasions 1: 301-311.
Blossey B. and Skinner L. (2000). Design and importance of post-release monitoring. Pp. 693-706 In: Proceedings of the X International Symposium on Biological Control of Weeds, N.R. Spencer (Ed.)
Boettner G.H., Elkinton J.S. and Boettner C.J. (2000).
Effects of a biological control introduction on three nontarget native species of saturniid moths.
Conservation Biology 14: 1798-1806.
The nontarget effects of a generalist parasitoid fly, Compsilura concinnata (Diptera: Tachinidae), that has been introduced as a biological control agent against 13 pest species were examined. Results suggested that reported declines of silk moth populations in New England may have been caused by C. concinnata.
Bourchier R.S. and McCarty L.S. (1995). Risk assessment of biological control (predators and parasitoids). Bulletin of the Entomological Society of Canada 27: 15.
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.
Briese D.T. (1999). Open field host-specificity tests: is "natural" good enough for risk assessment? Pp. 44-59 In: Host specificity testing in Australasia: towards improved assays for biological control, T.M. Withers, L. Barton-Browne and J. Stanley (Ed.) Scientific Publishing, Department of Natural Resources, Brisbane.
Briese D.T. (2003). The centrifugal phylogenetic method used to select plants for host-specificity testing of weed biological control agents: Can and should it be modernised? In: Improving the selection, testing and evaluation of weed biocontrol agents, H. S. Jacob and D. T. Briese (Ed.) CRC for Australian Weed Management Technical Series no. 7, Adelaide, Australia
Briese D.T. (2006). Can an a priori strategy be developed for biological control? The case for Onopordum spp. thistles in Australia. Australian Journal of Entomology 45: 317-323
Briese D.T. and Walker A. (2008).
Choosing the right plants to test: The host-specificity of Longitarsus sp (Coleoptera : Chrysomelidae) a potential biological control agent of Heliotropium amplexicaule.
Biological Control 44: 271-285
Using the case of the root-feeding flea beetle, Longitarsus sp., a candidate agent for biological control of Heliotropium amplexicaule in Australia, this paper describes a new protocol, based on phylogeny, and refined by ecological and biogeographic similarities. Taxonomic nomenclature is de-emphasized in favour of strict phylogenetic relationships and the use of so-called "safeguard species" is abandoned. The testing showed that adult feeding extended to plant species with up to five degrees of phylogenetic separation from H. amplexicaule, indicating that there would be a moderate risk that more distantly related plants suffer some feeding damage by adult Longitarsus sp. when they co-occur with infestations of the target weed that have large flea-beetle populations. Longitarsus sp. was able to complete its life-cycle on plants related to the target weed by two degrees of phylogenetic separation or less, leaving indigenous Heliotropium and Tournefortia species at some risk of colonisation. While these species had different life-histories and/or only slightly overlapped with the actual and potential range of the target weed, a minority of reviewers were concerned that insufficient information was available on the dispersal abilities of Longitarsus sp. to dismiss this risk. Release was therefore not approved, although this was not unexpected, as the assessment was based on factors that modified the effects of host range alone. The new protocols highlighted problems of an overreliance on taxonomic nomenclature as opposed to actual genetic relationships. However, they also directed attention to knowledge gaps in biogeography and agent biology that might refine the assessed risk.
Brodeur J. (2012). Host specificity in biological control: insights from opportunistic pathogens. Evolutionary Applications 5: 470-480.
Buckley Y.M., Rees M. and Paynter Q. (2004). Modelling integrated weed management of an invasive shrub in tropical Australia. Journal of Applied Ecology 41: 547-560
Butt T.M., Jackson C. and Magan N. (2002). Fungi as Biological Control Agents: Progress, Problems and Potential. CABI Publishing, Wallingford, U.K. 390 pp.
Byers R.A. and Kendall W.A. (1982). Effects of plant genotypes and root nodulation on growth and survival of Sitona spp. larvae. Environmental Entomology 11: 440-443.
Cagnotti C., Mc Kay F. and Gandolfo D. (2007).
Biology and host specificity of Plectonycha correntina Lacordaire (Chrysomelidae), a candidate for the biological control of Anredera cordifolia (Tenore) Steenis (Basellaceae).
African Entomology 15: 300-309
The paper described host range testing for Plectonycha correntina Lacordaire (Coleoptera: Chrysomelidae), a proposed biocontrol agent for the Neotropical perennial climber, Anredera cordifolia (Tenore) Steenis (Basellaceae), an environmental weed in Africa and Australasia. Larvae and adults feed on the leaves. The host range was evaluated by no-choice larval survival tests and adult feeding and oviposition choice tests using 16 test plant species. The results indicated that the host range of P. correntina is restricted to the Basellaceae, with A. cordifolia as its primary host, and so P. correntina was considered a safe and promising biocontrol agent for Madeira vine in countries such as Australia and New Zealand where no other Basellaceae occur.
Caltagirone L.E. (1981). Landmark examples in classical biological control. Annual Review of Entomology 26: 213-232.
Caltagirone L.E. and Huffaker C.B. (1980). Benefits and risks of using predators and parasites for controlling pests. Ecological Bulletin 31: 103-109.
Cameron P., Hill R.L., Bain J., Thomas W.P. (1993). Analysis of importations for biological control of insect pests and weeds in New Zealand. Biocontrol Science and Technology 3: 387-404.
Cameron P.J. and Walker G.P. (1989). Status of introduced larval parasitoids of tomato fruitworm. Proceedings of the New Zealand Plant Protection Conference 42: 229-232
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.
Cameron P.J., Hill R.L., Bain J. and Thomas W.P. (1989). A review of biological control of invertebrate pests and weeds in New Zealand 1874-1987. CAB International Wallingford, UK and DSIR, New Zealand.
Cameron P.J., Walker G.P. and Keller M.A. (1995). Introduction of Cotesia rubecula, a parasitoid of white butterfly. Proceedings of the New Zealand Plant Protection Conference 48: 345-347
Carruthers R.I. and D'Antonio C.M. (2005). Science and decision making in biological control of weeds: Benefits and risks of biological control. Biological Control 35: 181-182.
Carson R. (1963). Silent Spring. Hamish Hamilton, London.
Carson W.P., Hovick S.M., Baumert A.J., Bunker D.E. and Pendergast T.H. (2008).
Evaluating the post-release efficacy of invasive plant biocontrol by insects: a comprehensive approach.
Arthropod - Plant Interactions 2: 77-86
A program is proposed to evaluate the post-release phase of biocontrol programs that use insect herbivores to control invasive plant species. The authors argue that randomized release and non-release sites should be followed up to evaluate the degree of success or failure, including (1) the abundance of the biocontrol agent, (2) the impact of the biocontrol agent on the target plant species, (3) the potential for non-target effects (4) the response of native species and communities to a reduction in the invasive species and (5) experimental reductions of the biocontrol agent are required to eliminate the chance that the putative impact of the biocontrol agent is not confounded with other causes. Six scenarios are described in which a biocontrol agent may decrease the abundance or vigor of the target plant species but not lead to successful control where native communities re-establish.
Carvalheiro L.G., Buckley Y.M., Ventim R. and Memmott J. (2008).
Assessing indirect impacts of biological control agents on native biodiversity: a community-level approach.
Pp. 83-86 In: Proceedings of the XII International Symposium on Biological Control of Weeds, (Ed.) La Grande Motte, France, 22-27 April, 2007.
Apparent competition (competition due to shared natural enemies) has been neglected when considering possible impacts of biological control agents because of the difficulty in assessing and predicting indirect effects. In this paper the authors outline a methodology to predict and measure non-target impacts of biological control agents due to apparent competition.
Carvalheiro L.G., Buckley Y.M., Ventim R., Fowler S.V. and Memmott J. (2008).
Apparent competition can compromise the safety of highly specific biocontrol agents.
Ecology Letters 11: 690-700
Using food webs, the authors demonstrate that the use of a highly host-plant specific weed biocontrol agent, recently introduced into Australia, is associated with declines of local insect communities via indirect effects, most likely apparent competition. Both species richness and abundance in insect communities (seed herbivores and their parasitoids) were negatively correlated with the abundance of the biocontrol agent, Mesoclanis polana (Diptera: Tephritidae), a seed herbivore of Chrysanthemoides monilifera ssp. rotundata (Bitou). More investment is required in pre-release studies on the effectiveness of biocontrol agents, as well as in post-release studies assessing indirect impacts, to avoid or minimize the release of potentially damaging species.
Casas J., Swarbrick S. and Murdoch W.W. (2004).
Parasitoid behaviour: predicting field from laboratory.
Ecological Entomology 29: 657-665.
The basic components of Aphytis melinus's response to California red scale (Aonidiella aurantii) were studied in the laboratory and validated in the field. Laboratory studies predicted foraging behaviour in the field with variable success; potential explanations for observed mismatch between laboratory and field and its possible larger implications are discussed.
Catherine G.W., Schulthess F. and Stephane D. (2010.). An association between host acceptance and virulence status of different populations of Cotesia sesamiae , a braconid larval parasitoid of lepidopteran cereal stemborers in Kenya. Biological Control 54: 100-106
Causton C.E. (2004). Predicting the field host range of an introduced predator, Rodolia cardinalis Mulsant, in the Galapagos. Pp. 195-239 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
Chalak M., Hemerik L., van der Werf W., Ruijs A. and van Ierland E.C. (2010). On the risk of extinction of a wild plant species through spillover of a biological control agent: analysis of an ecosystem compartment model. Ecological Modelling 221: 1934-1943
Charles J.G. (1998). The settlement of fruit crop arthropod pests and their natural enemies in New Zealand: an historical guide to the future. Biocontrol News and Information 19: 47-58
Charles J.G. (2001). Introduction of a parasitoid for mealybug biocontrol: a case study under new environmental legislation. New Zealand Plant Protection 54: 37-41
Charles J.G. and Allan D.J. (2002). An ecological perspective to host-specificity testing of biocontrol agents. New Zealand Plant Protection 55: 37-41
Charles, J.G. (2011).
Using parasitoids to infer a native range for the obscure mealybug, Pseudococcus viburni, in South America.
Biocontrol 56: 155–161
An examination of the biogeographical origins and historical trade records provided an explanation as to why the obscure mealybug, Pseudococcus viburni (Signoret) (Hemiptera: Pseudococcidae), considered to be an American species, is not attacked by native parasitoids in the USA, whereas it is controlled in Europe by Acerophagus maculipennis (Mercet) (Encyrtidae) described from the Canary Islands (as Pseudophycus maculipennis). The hypothesis was supported that P. viburni and A. maculipennis are co-evolved Neotropical species, and that both were transported from S. America (probably Chile) to Europe via the Canary Islands possibly as early as the sixteenth century. Invasion of P. viburni into the USA occurred later, but without natural enemies. This explains why P. viburni in the USA is not attacked by native North American parasitoids and why A. maculipennis is not known to attack any mealybugs of Palaearctic origin. The hypothesis adds confidence that well conducted classical biocontrol programmes involving these taxa pose a low environmental risk to native, non-target fauna.
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.
Charudattan R. (2005). Ecological, practical, and political inputs into selection of weed targets: What makes a good biological control target? Biological Control 35: 183-196.
Chong J.-H. and Oetting R.D. (2007). Specificity of Anagyrus sp nov nr. sinope and Leptomastix dactylopii for six mealybug species. BioControl 52: 289-308
Civeyrel L. and Simberloff D. (1996). A tale of two snails: is the cure worse than the disease? Biodiversity and Conservation 5: 1231-1252.
Clarke A.R. (1995).
"Strains" and the classical biological control of insect pests.
Canadian Journal of Zoology 73: 1777-1790
The strategy of introducing two or more populations of the same species of beneficial agent to increase the genetic diversity of that species is reviewed. From the literature literature, cases of multiple introductions of conspecific populations against insect targets were listed and the effect of subsequent introductions on the outcome of the project was recorded. The analysis suggested that introducing two or more populations of the same species is less likely to result in enhanced success than if other species of natural enemies are sought for "normal" classical biological control (historical success rate 12-16%). It was considered from a reveiw of genetic theory that there is also no theoretical support for the continued introduction of strains.
Cock M.J.W., Murphy S.T., Kairo M.T.K., Thompson .E, Murphy R.J. and Francis A.W. (2016). Trends in the classical biological control of insect pests by insects: an update of the BIOCAT database. Biocontrol. doi:DOI 10.1007/s10526-016-9726-3.
Coetzee J.A., Byrne M.J., Hill M.P. and Center T.D. (2009).
Should the mirid, Eccritotarsus catarinensis (Heteroptera: Miridae), be considered for release against water hyacinth in the United States of America?
Biocontrol Science & Technology 19: 103-111
Eccritotarsus catarinensis (Carvalho) (Heteroptera: Miridae), damageswater hyacinth on the African continent, and was considered potentially useful in the USA where water hyacinth remains a problem. However, during host specificity trials, it developed on Pontederia cordata L. (pickerelweed), indigenous to the USA, although it did not establish on pickerelweed monocultures during South African field trials. The authors used models developed for South Africa using CLIMEX to predict whether the mirid will establish where water hyacinth and pickerelweed co-occur, but not where pickerelweed occurs in the absence of water hyacinth. The models suggest that the mirid's distribution will be limited by cold winter temperatures and insufficient thermal accumulation to the southern states of the USA, within the main distribution of water hyacinth. Itv was concluded that the benefits outweigh the minimal risk of damage to pickerelweed.
Coombs M. (2003). Post-release evaluation of Trichopoda giacomellii (Diptera: Tachinidae) for efficacy and non-target effects. Pp. 399-406 In: Proceedings of the 1st International Symposium on Biological Control of Arthropods, Honolulu, Hawaii, 14-18 January 2002, R. G. Van Driesche (Ed.) United States Department of Agriculture, Forest Service, Washington
Cory J.S. and Myers J.H. (2000).
Direct and indirect ecological effects of biological control.
Trends in Ecology and Evolution 15: 137-139.
The identification of potential impacts by risk and benefit analysis, host specificity testing, the impacts of biopesticides and the evolutionary stability of host range are discussed.
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.
Crawley M.J. (1989). Plant life history and the success of weed biological control projects. Pp. 17-26 In: Proceedings of the VII International Symposium on Biological Control of Weeds, E. S. Delfosse (Ed.) Rome, Italy. Istituto Sperimentale per la Patologia Vegetale (MAF).
Cristofaro M.De Biase A. and Smith . (2013). Field release of a prospective biological control agent of weeds, Ceratapion basicorne, to evaluate potential risk to a nontarget crop. Biological Control 64: 305-314.
Cullen J.M. (1989). Current problems in host-specificity screening. Pp. 27-36 In: Proceedings of the VII International Symposium on Biological Control of Weeds, E.S. Delfosse (Ed.) CSIRO Publications, Melbourne.
Cullen J.M. (1995). Predicting effectiveness: fact or fantasy? Pp. 103-109 In: Proceedings of the VIII International Symposium on Biological Control of Weeds, E. S. Delfosse and R. R. Scott (Ed.) Lincoln University, New Zealand, DSIR/CSIRO, Melbourne, Australia.
Cullen J.M. (1997).
Biological control and impacts on non-target species.
Pp. 195-201 In: Proceedings of the 50th New Zealand Plant Protection Conference, M. O'Callaghan (Ed.) New Zealand Plant Protection Society Inc.
Analysis of examples of non-target impacts of weed biological control agents suggested that in most cases impacts are limited, but the potential exists for serious impacts. A risk analysis approach is advocated. Lack of relevant research data is a major problem.
Cullen J.M. and Hopkins D.C. (1982).
Rearing, release and recovery of Microctonus aethiopoides Loan (Hymenoptera: Braconidae) imported for the control of Sitona discoideus Gyllenhal (Coleoptera: Curculionidae) in south eastern Australia.
Journal of the Australian Entomological Society 21: 279-284.
The braconid parasite Microctonus aethiopoides Loan was imported from Morocco in 1976 and released at sites in South Australia and New South Wales in 1977 and 1978 for the biological control of Sitona discoideus Gylh., a pest of lucerne and annual species of Medicago. Rearing, release and recovery methods are described, including techniques necessary to overcome the problems posed by aestivation of the host. The parasite has become established at several sites, and is a promising control agent with a high searching capacity and rapid rate of increase relative to its host.
Davies J.T., Ireson, J.E. and Allen G.R., (2005). The impact of gorse thrips, ryegrass competition, and simulated grazing on gorse seedling performance in a controlled environment. Biological Control 32: 280-286
Day M.D. (1999). Continuation trials: their use in assessing the host range of a potential biological control agent. Pp. 11-19 In: Host specificity testing in Australasia: towards improved assays for biological control, T.M. Withers, L. Barton-Brown and J. Stanley (Ed.) CRC for Tropical Pest Management, Brisbane.
DeBach P. (1974). Biological control by natural enemies. Cambridge University Press, London.
DeBach P. and Rosen D. (1991). Biological control by natural enemies. Cambridge University Press, Cambridge, UK.
DePrenger-Levin M.E., Grant TA., Dawson C. (2010).
Impacts of the introduced biocontrol agent, Rhinocyllus conicus (Coleoptera: Curculionidae), on the seed production and population dynamics of Cirsium ownbeyi (Asteraceae), a rare, native thistle.
Biological Control 55(2): 79-84.
This study evaluated non-target effects of Rhinocyllus conicus Frolich, on Cirsium ownbeyi S.L. Welsh, a rare, native and short-lived perennial thistle in northwestern Colorado, northeastern Utah, and southwestern Wyoming. C. ownbeyi is one of 22 known native hosts on which this introduced weevil has naturalized. The study population remained stable over the eight years of the study despite damage by thebeetle. The growth rate from was 1.03; however, large inter-year variation indicates this rare species is still vulnerable to local extirpation. The target species, Carduus nutans L. (musk thistle) is generally absent near the study plots, which may limit the population levels of R. conicus that can be sustained in this area. Although R. conicus utilizes C. ownbeyi as a host plant, the late flowering and the small size of the flower heads may limit the impact of R. conicus on C. ownbeyi.
Delalibera J.I. (2009). Biological Control of the Cassava Green Mite in Africa with Brazilian Isolates of the Fungal Pathogen Neozygites tanajoae. Pp. 259-269 In: Use of Microbes for Control and Eradication of Invasive Arthropods. A. E. Hajek, T. R. Glare and M. O'Callaghan (Eds.)
Delfosse E.S. (2005).
Risk and ethics in biological control.
Biological Control 35: 319-329.
Traditional risk analysis techniques are discussed and adapted for biological control. How people perceive risk is the key to understanding their attitude to risk. Criticisms of biological control relating to inadequate post-release monitoring are valid and the ethical responsibilities of scientists in this area are also discussed.
Denslow J.S. and D'Antonio C.M. (2005).
After biocontrol: Assessing indirect effects of insect releases.
Biological Control 35: 307-318.
Weeds in conservation land have become a focus of biological control projects where desired outcomes include both reduction of the target and indirect effects of increased diversity and abundance of native species and restoration of ecosystem services. Few quantitative assessments of the impacts of pest plant reduction on community composition or ecosystem processes were found and there was variation in the impacts of agent(s) across the invasive range of the target plant as well as variation in impacts on the invaded ecosystem. Most successful weed management programs integrated the use of biocontrol agents with other weed management strategies, especially modifications of disturbance and competing vegetation.
Desneux, N., Blahnik, R., Delebecque, C.J. and Heimpel, G.E. (2012).
Host phylogeny and specialisation in parasitoids.
Ecology Letters 5: 453-460
The authors build upon previous studies of preference- and performance-related traits on the host range of the aphid parasitoid Binodoxys communis (Hymenoptera: Braconidae) by mapping a series of these traits onto the phylogeny of the (aphid) host species. They found that both classes of traits showed phylogenetic conservatism with respect to host species.
Dhileepan K., Lockett C.J., Balu A., Murugesan S., Perovic D.J. and Taylor D.B.J. (2015). Life cycle and host range of Phycita sp. rejected for biological control of prickly acacia in Australia. Journal of Applied Entomology 139: 800-812.
Dhileepan K., Trevino M. and Raghu S. (2006).
Temporal patterns in incidence and abundance of Aconophora compressa (Hemiptera: Membracidae), a biological control agent for Lantana camara, on target and nontarget plants.
Environmental Entomology 34: 1001-1012
The membracid Aconophora compressa Walker, released in 1995 to control Lantana camara (Verbenaceae) in Australia, has since been collected on several nontarget plant species. A survey suggested that sustained populations of A. compressa were found only on the introduced nontarget ornamental Citharexylum spinosum (Verbenaceae) and the target weed L. camara. However, it was found on other nontarget plant species when populations on C. spinosum and L. camara were high, suggesting a spill-over effect. Some attack on nontarget plants could have been anticipated from host specificity studies done on this agent before release. This raises important issues about predicting risks posed by weed biological control agents and the need for long-term postintroduction monitoring on nontarget species.
Dhileepan K., Wilmot Senaratne K.A.D. and Raghu S. (2006). A systematic approach to biological control exploration and prioritisation for prickly acacia (Acacia nilotica spp. indica). Australian Journal of Entomology 45: 303-307
Drea J.J. (1993). Classical biological control - an endangered discipline? Pp. In: Biological Pollution: The control and impact of invasive exotic species, B.N. McNight (Ed.) Indiana Academy of Science, Indianapolis.
Duan J.J. and Messing R.H. (1996). Response of two Opiine fruit fly parasitoids (Hymenoptera: Braconidae) to the Lantana Gall fly (Diptera: Tephritidae). Environmental Entomology 25: 1428-1437.
Duan J.J. and Messing R.H. (1996). Risk analysis and decision-making in biological control - A case study with fruit fly parasitoids. Journal of Agriculture and Human Values 13: 1-10.
Duan J.J. and Messing R.H. (1996).
Effect of two Opiine parasitoids (Hymenoptera: Braconidae) introduced for fruit fly control on a native Hawaiian Tephritid, Trupanea dubautiae (Diptera: Tephritidae).
Biological Control 8: 177-184.
Diachasmimorpha longicaudata and Psyttalia fletcheri are opiine parasitoids introduced into Hawaii for control of Bactrocera dorsalis and Bactrocera cucurbitae, respectively. Both species have been mass-reared and released for research in augmentative biocontrol programs. The potential impact of mass-produced D. longicaudata and P. fletcheri on a native Hawaiian tephritid, Trupanea dubautiae, infesting the flowerheads of Dubautia raillardioides was investigated. The results demonstrated that biological control programs targeted against frugivorous tephritid pests by D. longicaudata and P. fletcheri have no harmful impact on T. dubautiae.
Duan J.J. and Messing R.H. (1997).
Biological control of fruit flies in Hawaii: factors affecting non-target risk analysis.
Agriculture and Human Values 14: 227-236.
Examples from both classical and augmentative biological control of fruit fly pests (Tephritidae) in Hawaii were used to address non-target risks of fruit fly parasitoids (Braconidae). A lack of host-specificity testing of parasitoids with non-target species has raised concerns about their impact on non-pest fruit flies, including some flies introduced for weed biological control endemic Hawaiian species. For assessing susceptibility of a non-target species to parasitoids, behavioural tests are as important as suitability tests. Experimental factors, such as host-exposure substrate, absence or presence of preferred hosts, and laboratory vs. natural conditions, were shown to affect the results of host-specificity tests and risk analysis.
Duan J.J. and Messing R.H. (1998).
Effect of Tetrastichus giffardianus (Hymenoptera: Eulophidae) on nontarget flowerhead-feeding tephritids (Diptera: Tephritidae).
Biological Control 27: 1022-1028.
Laboratory tests and field surveys were conducted in Hawaii to evaluate the impact of a deliberately introduced fruit fly parasitoid, Tetrastichus giffardianus, on 2 nontarget flowerhead-feeding tephritid flies, Trupanea dubautiae and Ensina sonchi. In the laboratory, T. giffardianus were able to parasitize late instars of both T. dubautiae and E. sonchi when they were dissected out of the flower heads, however, no T. dubautiae and few E. sonchi were attacked by T. giffardianus when presented in their respective host flowerheads. So although late instars of both T. dubautiae and E. sonchi are suitable for the physiological development of T. giffardianus progeny, the parasitoid is unlikely to affect either species under natural conditions because the host microhabitats are not suitable for gravid parasitoids to find and oviposit in the fly larvae.
Duan J.J. and Messing R.H. (1999). Evaluating nontarget effects of classical biological control: fruit fly parasitoids in Hawaii as a case study. Pp. 95-109 In: Nontarget effects of biological control introductions, P.A. Follett and J.J. Duan (Ed.) Kluwer Academic Publishers, Norwell, Massachusetts, USA.
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.
Duan J.J., Ahmad M., Joshi K. and Messing R.H. (1996).
Evaluation of the impact of the fruit fly parasitoid Diachasmimorpha longicaudata (Hymenoptera: Braconidae) on a nontarget tephritid, Eutreta xanthochaeta (Diptera: Tephritidae).
Biological Control 8: 58-64.
Field studies were carried out in Hawaii to investigate the effects of the release of Biosteres longicaudatus on Eutreta xanthochaeta, a biological control agent of the weed Lantana camara. In field cages, Bactrocera dorsalis was presented on guavas and lantana twigs with galls. The presence of B. dorsalis or food had minimal effect on visiting and probing of B. longicaudatus and rates of parasitism were very low. It was concluded that an increase in the number of B. longicaudatus will not result in significant losses of E. xanthochaeta or influence the biological control of L. camara.
Duan J.J., Purcell M.F. and Messing R.H. (1996). Parasitoids of non-target tephritid flies in Hawaii: implications for biological control of fruit fly pests. Entomophaga 41: 245-256.
Dymock J.J. (1987). Population changes of the seedfly, Pegohylemyia jacobaeae (Diptera: Anthomyiidae) introduced for biological control of ragwort. New Zealand Journal of Zoology 14:337-342.
ERMA New Zealand (1998). Annotated methodology for the consideration of applications for hazardous substances and new organisms under the HSNO Act 1996. ERMA New Zealand, Wellington, New Zealand. 28 pp.
ERMA New Zealand (2000). Pseudaphycus maculipennis for the control of the obscure mealybug (Pseudococcus viburni). Evaluation and Review Report.
Edwards P. (1999). The use of choice tests in host-specificity testing of herbivorous insects. Pp. 35-43 In: Host specificity testing in Australasia: towards improved assays for biological control, T.M. Withers, L. Barton-Browne and J. Stanley (Ed.) Scientific Publishing, Department of Natural Resources, Brisbane.
Ehler L.E. (1990). Environmental impact of introduced biological control agents: implications for agricultural biotechnology. Pp. 85-96 In: Risk assessment in agricultural biotechnology, J.J. Marois and G. Breuning (Ed.) California Division of Agriculture and Natural Resources, Oakland.
Ehler L.E. (1999). Critical issues related to nontarget effects in classical biological control of insects. Pp. 3-13 In: Nontarget effects of biological control introductions, P.A. Follett and J.J. Duan (Ed.) Kluwer Academic Publishers, Norwell, Massachusetts, USA.
Elkinton J.S., Parry D. and Boettner G.H. (2006).
Implicating an introduced generalist parasitoid in the invasive browntail moth's enigmatic demise.
Ecology 87: 2664-2672.
The parasitoid Compsilura concinnata (tachinid) is considered an example of biological control gone wrong. It was introduced for gypsy moth, Lymantria dispar, and now attacks more than 180 species of native Lepidoptera in North America. It failed to control gypsy moth, data suggest that parasitism by C. concinnata is the cause of the near extinction of another exotic lepidopteran, the browntail moth (Euproctis chrysorrhoea). Despite this beneficial role played by C. concinnata in controlling the browntail moth, the authors do not advocate introduction of generalist biological control agents, which can have unpredictable and far-reaching impacts.
Elkinton, J.S. and Boettner, C.J. (2012).
Benefits and harm caused by the introduced generalist tachinid, Compsilura concinnata, in North America.
Biocontrol 57: 277–288
Compsilura concinnata (Meigen) is a highly generalist tachinid parasitoid that was introduced in the USA to control gypsy moth and browntail moth. The impact on gypsy moth was thought to be minor, although research with experimentally created populations of gypsy moth suggests that it may be more important than previously realized. Studies on browntail moth suggest that C. concinnata is probably the main reason browntail moth disappeared from most of its former range in North America. Research on giant silk moths suggests that C. concinnata has become the major source of mortality among several species and may be responsible for the notable decline in their densities that has occurred over the last century. C. concinnata is considered an example of a generalist natural enemy that should be avoided in classical biological control introductions, yet in the case of browntail moth its effect has been extremely beneficial.
Etzel L.K. and Legner E.F. (1999). Culture and colonisation. Pp. 125-197 In: Handbook of Biological Control: Principles and Applications of Biological Control, T.S. Bellows and T.W. Fisher (Ed.) Academic Press.
European and Mediterranean Plant Protection Organization (EPPO) (2010). Import and release of non-indigenous biological control agents. Bulletin OEPP/EPPO Bulletin 40: 335-344
Evans A.A., Barratt B.I.P. and Emberson R.M. (1997).
Field cage and laboratory parasitism of Nicaeana cervina by Microctonus aethiopoides.
Pp. 223-226 In: Proceedings of the 50th New Zealand Plant Protection Conference, M.R. O'Callaghan (Ed.) New Zealand Plant Protection Society Inc.
A study was carried out to compare parasitism of the NZ native weevil Nicaeana cervina Broun by Microctonus aethiopoides Loan in field cage versus laboratory conditions. Total parasitism was 40-55% and 15% in laboratory and field cages, respectively. The level of parasitism obtained in the field cages was similar to that recorded in a natural population nearby.
Evans A.A., Barratt B.I.P. and Ferguson C.M. (1994). Susceptibility of legume and Hieracium spp. seedlings to feeding by native broad-nosed weevils (Coleoptera: Curculionidae). Pp. 206-209 In: Proceedings of the 47th New Zealand Plant Protection Conference, A.J. Popay (Ed.) Waitangi Hotel, Pahia, N.Z., New Zealand Plant Protection Society Inc.
Evans W.E. and England S. (1996).
Indirect interactions in biological control of insects: pests and natural enemies in alfalfa.
Ecological Applications 6: 920-930.
Trophic interaction between alfalfa weevil, Hypera postica, and its parasitoid Bathyplectes curculionis, along with pea aphid (which produce honeydew, of value to the parasitoid) and ladybird beetles which feed on aphids and H. postica larvae. The results emphasise the complexity of these interactions.
FAO (1996). Code of conduct for the import and release of exotic biological control agents. Food and Agriculture Organisation. International Standards for Phytosanitary Measures No 3.
Faria L. de B., Umbanhowar J. and McCann K.S. (2008).
The long-term and transient implications of multiple predators in biocontrol.
Theoretical Ecology 1: 45-53
The authors explore the role of multiple predators on the transient and long-term dynamic outcomes of biological control. Theory indicates that specialist predators ought to promote less stable long-term biological control than generalists, while generalists readily drive suppression of nontarget prey species. However, these results showed that the combination of specialists and generalists acted synergistically to promote safe biological control. The results also suggested that endemic generalist predators, not introduced generalist predators, may often be responsible for the suppression and elimination of nontarget species. This final result demands empirical attention.
Fauvergue X., Malausa J.-C., Giuge L. and Courchamp F. (2007).
Invading parasitoids suffer no Alee effect: a manipulative field experiment.
Ecology 88: 2392-2403
Populations released for biological control are similar to fortuitous invading populations and may therefore suffer from Allee effects, and since experimental manipulation of initial population size is possible, a unique opportunity to test for the occurrence of Allee effects is provided. The initial size of 45 populations of a parasitoid wasp introduced for the biological control of a phytophagous insect was measured monitored for three years. Results suggested an absence of Allee effects but clear negative density dependence instead: (1) the probability of establishment after three years was not affected by initial population size; (2) net reproductive rate was highest at low parasitoid density and high host density; (3) the sex ratio, reflecting the proportion of virgin females, did not increase at low density, suggesting that low densities did not impede matefinding; (4) the depression of host populations did not depend upon the number of parasitoids introduced.
Ferguson C.M., Barratt B.I.P. and Cresswell A.S. (1999). Field parasitism of the weed biological control agent Rhinocyllus conicus by the introduced braconid, Microctonus aethiopoides. Pp. 275 (abstract) In: Proceedings of the 52nd New Zealand Plant Protection Society Conference, M.R. O'Callaghan (Ed.) New Zealand Plant Protection Society.
Ferguson C.M., Cresswell A.S., Barratt B.I.P. and Evans A.A. (1998). Non-target parasitism of the weed biological control agent, Rhinocyllus conicus Froelich (Coleoptera: Curculionidae) by Microctonus aethiopoides Loan (Hymenoptera: Braconidae). Pp. 517-524 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, Australia.
Ferguson C.M., Kean J.M., Barton D.M. and Barratt B.I.P. (2016).
Ecological mechanisms for non-target parasitism by the Moroccan ecotype of Microctonus aethiopoides Loan (Hymenoptera: Braconidae) in native grassland.
Biological Control 96: 28-38.
The Moroccan ecotype of the braconid parasitoid Microctonus aethiopoides was introduced into New Zealand for biological control of the lucerne pest Sitona discoideus. The parasitoid also attacks several non-target native weevil species found in pasture and also to a lesser extent in native tussock grassland. We carried out a series of laboratory and field experiments, and population modelling to investigate whether the parasitoids were established at low levels on native weevils in tussock grassland, whether S. discoideus was able to survive and support parasitoid development away from lucerne, its normal host plant, or whether parasitism was occurring as a result of spillover from agricultural environments. We found that S. discoideus was able to survive and support parasitoid development on white clover which is commonly found in native grassland. However, the levels of parasitism in weevil species in tussock grassland appeared to be constrained, at least in part, by low temperatures limiting the number of parasitoid generations possible per year and by the frequency of sub-zero temperatures that caused pupal mortality. Projected climate change might reduce this constraint and the implications of this are discussed.
Ferguson C.M., Moeed A., Barratt B.I.P., Hill R.L. and Kean J.M. (2007).
BCANZ - Biological Control Agents introduced to New Zealand.
This database contains information on the biological control agents that have been introduced to New Zealand to help manage weed and invertebrate pests. The database currently contains records for 720 introductions of 518 biological control agents against 126 targets (25 weeds and 101 invertebrates). This information is being constantly updated.
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.
Fernandez G.C.J. (1992). Residual analysis and data transformation: Important tools in statistical analysis. Horticultural Science 27: 297-300.
Field R.P. and Darby S.M. (1991).
Host specificity of the parasitoid, Sphecophaga vesparum (Curtis) (Hymenoptera: Ichneumonidae), a potential biological control agent of the social wasps, Vespula germanica (Fabricius) and V. vulgaris (Linnaeus) (Hymenoptera: Vespidae) in Australia.
New Zealand Journal of Zoology 18: 193-197
Choice, non-choice, and host location tests using Sphecophaga vesparum indicated that brood of some Australian native Polistes, Ropalidia, and Trigona species would not be at risk from releases of the parasitoid in Australia. S. vesparum was approved for release in Australia and released in metropolitan Melbourne (Victoria) in December 1989, to act as a biological control agent against Vespula species
Follett P.A., Johnson M.T. and Jones V.P. (1999). Parasitoid drift in Hawaiian pentatomids. Pp. 77-93 In: Nontarget effects of biological control introductions, P.A. Follett and J.J. Duan (Ed.) Kluwer Academic Publishers, Norwell, Massachusetts, USA.
Fornasari L., Turner C.E. and Andres L.A. (1991). Eustenopus villosus (Coleoptera: Curculionidae) for biological control of yellow starthistle (Asteraceae: cardueae) in north America. Environmental Entomology 20: 1187-1194.
Forno I.W., Kassulke R.C. and Harley K.L.S. (1992). Host specificity and aspects of the biology of Calligrapha pantherina (Col.:Chrysomelidae), a biological control agent of Sida acuta (Malvaceae) and S. rhombifolia in Australia. Entomophaga 37: 409-417.
Fowler S.V. (2000). Trivial and political reasons for the failure of classical biocontrol of weeds: a personal view. Pp. 169-172 In: Proceedings of the International Symposium on Biological Control of Weeds, N.R. Spencer (Ed.) Bozeman, Montana.
Fowler S.V. and Griffin D. (1995). The effect of multi-species herbivory on shoot growth in gorse Ulex europaeus. Pp. 579-584 In: Proceedings of the VIIIth International Symposium on Biological Control of Weeds, E.S. Delfosse (Ed.) Christchurch, New Zealand
Fowler S.V., Barreto R., Dodd S., Macedo D.M., Paynter Q., Pedrosa-Macedo J.H., Pereira O.L., Peterson P., Smith L.Waipara N., Winks C.J. and Forrester G. (2013). Tradescantia fluminensis, an exotic weed affecting native forest regeneration in New Zealand: Ecological surveys, safety tests and releases of four biocontrol agents from Brazil. Biological Control 64: 323-329.
Fowler S.V., Gourlay A.H., Hill R.L. and Withers T. (2003).
Safety in New Zealand weed biocontrol: a retrospective analysis of host-specificity testing and the predictability of impacts on non-target plants.
Pp. 265–270 in Proceedings of the XI International Symposium on Biological Control of Weeds, Canberra, Australia, 2003, J.M. Cullen, D.T. Briese, D.J. Kriticos, W.M. Lonsdale, L. Morin and J.K. Scott (Ed.).
A retrospective analysis showed that all weed biocontrol agents released in New Zealand were subjected to generally appropriate host-range tests, although there were several examples where significant plant species were not tested. The results have been used to focus field surveys on the most likely non-target plant species to be attacked by biocontrol agents in New Zealand. For example, Tyria jacobaeae (cinnabar moth) did feed on some Senecio species in the original host-range tests, so the occasional field attack on native New Zealand fireweeds such as S. minimus was predictable. To date, this is the only weed biocontrol agent in New Zealand (of the total of 32 established in the field since 1929) that has been recorded attacking a native non-target plant species in the field. There were two cases where test results did not predict potentially substantial non-target impacts: Bruchidius villosus (broom seed beetle) and Cydia succedana (gorse pod moth), attacking seed of nontarget, exotic Fabaceae. Limited replication and duration of tests, were among possible explanations for the failure to predict these impacts.
Fowler S.V., Syrett P. and Hill R.L. (2000).
Success and safety in the biological control of environmental weeds in New Zealand.
Austral Ecology 25: 553-562.
Weed biological control agents have been recorded attacking non-target plants in NZ and elsewhere, but the effects are usually minor and/or transitory. Probably only two cases, worldwide, will result in significant damage to non-target plants both of which predictable from host range testing. For NZ programmes a full/partial success rate of 83% was calculated. Costs of biocontrol programmes against some NZ weeds can be reduced by using Australian research.
Fowler, S.V., Paynter, Q., Dodd, S. and Groenteman, R. (2012). How can ecologists help practitioners minimize non-target effects in weed biocontrol? Journal of Applied Ecology 49: 307-310
Frank J.H. (1998). How risky is biological control? Comment. Ecology 79: 1829-1834.
Froud K.J. and Stevens P.S. (2004). Importation biological control of Heliothrips haemorrhoidalis (Thysanoptera: Thripidae) by Thripobius semiluteus (Hymenoptera: Eulophidae) in New Zealand - a case study of non-target host and environmental risk assessment. Pp. 90-102 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, Morganstown, West Virginia, USA
Froud K.J. and Stevens P.S., (1998). Parasitism of Heliothrips haemorrhoidalis and two non-target thrips by Thripobius semiluteus (Hymenoptera; Eulophidae) in quarantine. Pp. 526-529 In: Pest Management - Future Challenges. Proceedings of the 6th Australasian Applied Entomological Research Conference, M.P. Zalucki, R.A.I. Drew and G.G. White (Ed.) Brisbane Australia.
Frye, M.J., Lake, E.C. and Hough-Goldstein, J. (2010).
Field host-specificity of the mile-a-minute weevil, Rhinoncomimus latipes Korotyaev (Coleoptera: Curculionidae).
Biological Control 55: 234-240
The authors hypothesized that Rhinoncomimus latipes (Coleoptera: Curculionidae), the biological control agent released against mile-a-minute weed, Persicaria perfoliata (Polygonaceae), would not feed or oviposit on nontarget plants in a two-phase, open field setting. Whereas prerelease studies showed feeding at low levels on 9 of the 13 plant species tested here, under open field conditions R. latipes did not feed on any nontarget plant species and dispersed from these plants. In an open field setting, where the weevil was able to use its full range of host-selection behaviors, there was no observed risk of nontarget effects for any species tested
Fuester R.W., Kenis M., Swan K.S., Kingsley P.C., López-Vaamonde C. and Hérard F. (2001). Host Range of Aphantorhaphopsis samarensis (Diptera: Tachinidae), a larval parasite of the gypsy moth (Lepidoptera: Lymantriidae). Environmental Entomology 25: 332-340.
Futuyma D.J. (1999). Potential evolution of host range in herbivorous insects. Pp. 42-53 In: Proceedings of the X International Symposium of Biological Control of Weeds, N.R. Spencer (Ed.).
Futuyma D.J. (2000). Potential evolution of host range in herbivorous insects. Pp. 42-53 In: Host-specificity testing of exotic arthropod biological control agents: the biological basis for improvement in safety, R.G. Van Driesche, T. Heard, A.S. McClay and R. Reardon (Ed.) USDA Forest Service Bulletin, Morgantown, West Virginia, USA.
Gariepy T., Kuhlmann U., Gillott C. and Erlandson M. (2008).
A large-scale comparison of conventional and molecular methods for the evaluation of host-parasitoid associations in non-target risk-assessment studies.
Journal of Applied Ecology 45: 708-715
Host rearing and dissection are used to define the ecological host range of candidate biological control agents and assess host-specificity of parasitoids, however, molecular methods may also be useful, e.g. Lygus plant bugs, host rearing, dissection and multiplex polymerase chain reaction (PCR) analysis were compared for estimation of parasitism levels and parasitoid species composition in field-collected target and non-target Miridae. Parasitism levels estimated by conventional and molecular methods were similar but molecular analysis could detect parasitoids earlier than dissection and rearing. Molecular methods can provide more complete, parasitoid species composition information because the results are not confounded by the host and parasitoid mortality encountered in rearing. However, detection of a parasitoid in a host does not necessarily indicate survival to the adult stage. Beyond agent identification, molecular diagnostics can facilitate and expedite pre- and post-release studies on the ecological host range of parasitoids, potential non-target effects, host-parasitoid associations and trophic interactions.
Gassman A. and Louda S.M. (2001). Rhinocyllus conicus: initial evaluation and subsequent ecological impacts in North America. Pp. 147-183 In: Evaluating indirect ecological effects of biological control, E. Wajnberg, J.K. Scott and P.C. Quimby (Ed.) CABI Publishing, Wallingford, Oxon., UK.
Gassmann A. and Schroeder D. (1995). The search for effective biological control agents in Europe: History and lessons from leafy spurge (Euphorbia esula L.) and cypress spurge (E. cyperissias L.). Biological Control 5: 466-472
Gassmann A., Tosevski I. and Skinner L. (2008).
Use of native range surveys to determine the potential host range of arthropod herbivores for biological control of two related weed species, Rhamnus cathartica and Frangula alnus.
Biological Control 45: 11-20
The buckthorn species, Rhamnus cathartica and Frangula alnus have become invasive in North America. The key question for biocontrol of teghse species was whether they are distantly enough related that they would not share the same arthropod complex in Europe, and, if so, which arthropod species would be less likely to use native North American buckthorns as hosts. Sampling in Europe indicated that the arthropod-species richness is higher on R. cathartica than on F. alnus and includes more species that are presumed to be host-specific at the species or genus level. At least 12 arthropod species were found exclusively on Rhamnus, some of which may be specific to R. cathartica and only one species was found exclusively on F. alnus.
Gerard P.J., McNeill M.R., Barratt B.I.P. and Whiteman S.A. (2006). Rationale for release of the irish strain of Microctonus aethiopoides for biocontrol of clover root weevil. New Zealand Plant Protection 59: 285-289.
Gerber E., Hinz H.L., Blossey B. and Bacher S. (2004).
Two shoot miners as potential biological control agents for garlic mustard: should both be released?
Proceedings of the XI International Symposium on Biological Control of Weeds: 108-112
Two shoot-mining weevils, Ceutorhynchus alliariae and C. roberti, both potential biological control agents for Alliaria petiolata in North America, show high temporal and spatial niche overlap. The comparison of attack levels as an indirect estimate of their potential to damage garlic mustard showed that C. alliariae was equally as effective in attacking garlic mustard alone as in combination with C. roberti, infact under experimental conditions, C. alliariae alone reached higher infestation levels than the mixed species but did not result in a higher impact on garlic mustard. Replicated releases of different combinations of the two species would provide a unique opportunity to test the conclusions from our pre-release investigations.
Gilbert G.S. and Webb C.O. (2007).
Phylogenetic signal in plant pathogen-host range.
Proceedings of the National Academy of Sciences of the United States of America 104: 4979-4983
Susceptibility of plant species plant pathogens is poorly understood. Most fungal pathogens are usually polyphagous but most plant species in a local community are resistant to any given pathogen. The probablility that a pathogen can infect two plant species decreases continuously with phylogenetic distance between the plants. This allows prediction of the likely host range of plant pathogens in a local community. The results suggest that the rate of spread and ecological impacts of a disease through a natural plant community depends on the phylogenetic structure of the community itself and that current regulatory approaches strongly underestimate the local risks of global movement of plant pathogens or their hosts.
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.
Godfray H.C.J. (1994). Parasitoids: Behavioural and Evolutionary Ecology. Princeton University Press, Princeton. 473 pp.
Goldson S.L. and Phillips C.B. (1990). Biological control in pasture and lucerne and the requirements for futher responsible introduction of entomophagous insects. Bulletin of the Entomological Society of New Zealand 10: 63-74.
Goldson S.L., Barratt B.I.P., Barlow N.D. and Phillips C.B. (1998). What is a safe biological control agent? Pp. 530-538 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.
Goldson S.L., Frampton E.R. and Ridley G.S. (2010.).
The effects of legislation and policy in New Zealand and Australia on biosecurity and arthropod biological control research and development.
Biological Control 52: 241-244.
The authors highlight some of the differences between legislation, policy and what science can deliver relating to biological control and biosecurity in New Zealand and Australia. They also discuss some of the inconsistencies and impracticalities in their implementation with a focus on examples from arthropod biological control.
Goldson S.L., McNeill M.R. and Proffitt J.R. (2003). Negative effects of strain hybridisation on the biocontrol agent Microctonus aethiopoides. New Zealand Plant Protection 57: 138-142.
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.
Goldson S.L., McNeill M.R., Proffitt J.R., Barker G.M., Addison P.J., Barratt B.I.P. and Ferguson C.M. (1993). Systematic mass rearing and release of Microctonus hyperodae (Hym.: Braconidae, Euphorinae), a parasitoid of the Argentine stem weevil Listronotus bonariensis (Col.: Curculionidae) and records of its establishment in New Zealand. Entomophaga 38: 1-10.
Goldson S.L., Phillips C.B., McNeill M.R. and Barlow N.D. (1997). The potential of parasitoid strains in biological control: observations to date on Microctonus spp. intraspecific variation in New Zealand. Agriculture, Ecosystems and Environment 64: 115-124.
Goldson S.L., Proffitt J.R. and Baird D.B. (1998). Establishment and phenology of the parasitoid Microctonus hyperodae (Hymenoptera: Braconidae) in New Zealand. Environmental Entomology 27: 1386-1392.
Goldson S.L., Proffitt J.R. and McNeill M.R. (1990). Seasonal biology and ecology in New Zealand of Microctonus aethiopoides (Hymenoptera: Braconidae), a parasitoid of Sitona spp. (Coleoptera: Curculionidae), with special emphasis on atypical behaviour. Journal of Applied Ecology 27: 703-722.
Goldson S.L., Proffitt J.R. and Muscroft-Taylor K.E. (1993). The economic value of achieving biological control of Sitona discoideus. Pp. 45-60 In: Plant Protection: Costs, Benefits and Trade Implications, D.M. Suckling and A.J. Popay (Ed.) New Zealand Plant Protection Society Inc.
Goolsby J.A., Makinson J.R., Hartley D.M., Zonneveld R. and Wright A.D. (2004).
Pre-release evaluation and host-range testing of Floracarus perrepae (Eriophyidae) genotypes for biological control of Old World climbing fern.
Proceedings of the XI International Symposium on Biological Control of Weeds: 113-116
As part of a biological control program for Lygodium microphyllum, an invasive climbing fern in Florida, surveys for natural enemies were conducted in the fern's native range (Australia, Asia and Oceania). Twenty-two herbivores were identified including an eriophyid mite, Floracarus perrepae Knihinicki & Boczek. Using molecular diagnostics a plant population from Cape York, Queensland was found to be an exact match with the invasive populations in Florida for the two chloroplast DNA sequences analyzed. Pre-release field impact studies revealed that F. perrepae caused more than 50% impact on L. microphyllum biomass production over a two-year period. Several genotypes of the mite were screened for their acceptance of the invasive Florida genotype, and the populations from Cape York and Thailand performed best and came from fern genotypes that were most closely related to the Florida genotype.
Goolsby J.A., Van Klinken R.D. and Palmer W.A. (2006). Maximising the contribution of native-range studies towards the identification and prioritisation of weed biocontrol agents. Australian Journal of Entomology 45: 276-286
Gourlay A.H., Wittenberg R., Hill R.L., Spiers A.G. and Fowler S.V. (2000). The biological control programme against Clematis vitalba in New Zealand. Pp. 709-718 In: Proceedings of the X International Symposium on Biological Control of Weeds, N. R. Spencer (Ed.) Bozeman, Montana, USA Montana State University.
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.
Grandgirard J., Hoddle M.S., Petit J.N., Roderick G.K. and Davies N. (2008).
Engineering an invasion: classical biological control of the glassy-winged sharpshooter, Homalodisca vitripennis, by the egg parasitoid Gonatocerus ashmeadi in Tahiti and Moorea, French Polynesia.
Biological Invasions 10: 135-148
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.
Greathead D.J. (1971). A review of biological control in the Ethiopian region. Commonwealth Institute of Biological Control Technical Communication 5: 1-162.
Greathead D.J. (1995). Benefits and risks of classical biological control. Pp. 53-63 In: Biological Control: benefits and Risks, H.M.T. Hokkanen and J. Lynch (Ed.) Cambridge University Press, Cambridge, UK.
Greathead D.J. and Greathead A.H. (1992). Biological control of insect pests by insect parasitoids and predators: the BIOCAT database. Biocontrol News and Information 13: 61N-68N
Gripenberg, S., Hamer, N.I.A., Brereton, T.O.M., Roy, D.B. and Lewis, O.T. (2011).
A novel parasitoid and a declining butterfly: cause or coincidence?
Ecological Entomology 36: 271-281
The small tortoiseshell butterfly (Aglais urticae L.) declined sharply in the U.K. between 2003 and 2008, coinciding with the arrival and spread of a parasitoid, Sturmia bella Meig. (Diptera: Tachinidae), which specialises on nymphalid butterflies. Data from a large-scale butterfly monitoring scheme, and collections of larvae were to assess parasitoid incidence and parasitism frequency. Similar data were compiled for a related butterfly (Inachis io) which is also parasitised by S. bella but which is not declining. Sturmia bella was present in 26% and 15% of the larval groups of A. urticae and I. io, respectively, and now kills more individuals of A. urticae (but not I. io) than any native parasitoid. Results indicated that S. bella causes host mortality in addition to that caused by native parasitoids and that S. bella may be playing a role in the recent decline of A. urticae. Other potential drivers of trends in the abundance of this butterfly may be present.
Groenteman R., Kelly D., Fowler S.V. and Bourdot G.W. (2008).
Factors affecting oviposition rate in the weevil Rhinocyllus conicus on non-target Carduus spp. in New Zealand.
Pp. 87-90 In: Proceedings of the XII International Symposium on Biological Control of Weeds, La Grande Motte, France, 22-27 April, 2007
Rhinocyllus conicus (Froehlich) (Coleoptera: Curculionoidae), oviposits on developing thistle flower buds and larvae feeding on the receptacle prevents seed development. The weevil attacks several thistle species, but prefers nodding thistle, Carduus nutans L. The effects of plant characteristics on oviposition preference and/or the size of emerging adult weevils were examined on three Carduus species. The results showed that larger, higher seed heads on larger plants were preferred for oviposition and larger seed heads supported the development of larger adults. Nodding thistle flowers over an extended period of time but the two winged thistle species offer additional oviposition opportunities three to four weeks before nodding thistle flowers. The adults emerging from the winged thistle species are likely to establish a second generation, enabling this normally univoltine weevil to sustain seasonally prolonged attack on nodding thistle.
Groenteman, R., Fowler, S.V. and Sullivan, J.J. (2011).
St. John's wort beetles would not have been introduced to New Zealand now: A retrospective host range test of New Zealand's most successful weed biocontrol agents.
Biological Control 57: 50-58
St. John's wort, Hypericum perforatum, was a serious weed in New Zealand (NZ) pastures in the 1930s. Chrysolina hyperici and C. quadrigemina, were introduced to NZ in 1943 and 1965, respectively. Earlier host specificity testing in Australia was deemed sufficient for approval for release in NZ. A review of worldwide reports suggested that St. John's wort beetles will attack a range of Hypericum species in the field. After a series of laboratory tests conducted to simulate modern host-range-testing protocols the authors concluded that the two Chrysolina species would not have been approved for introduction to NZ under current risk assessment protocols, and that NZ would have missed out on one of its greatest biocontrol success stories. No evidence for impacts on the populations of indigenous congeners has been recorded. Better procedures are required to predict the realized host-range of an agent from the potential range in contained host-range testing.
Grosskopf G., Smith L.A. and Syrett P. (2002). Host range of Cheilosia urbana (Meigen) and Cheilosia psilophthalma (Becker) (Diptera: Syrphidae), candidates for the biological control of invasive alien hawkweeds (Hieracium spp., Asteraceae) in New Zealand. Biological Control 24: 7-19
Grosskopf G., Wilson L.M. and Littlefield J.L. (2008).
Host-range investigations of potential biological control agents of alien invasive hawkweeds (Hieracium spp.) in the USA and Canada: an overview.
Proceedings of the XII International Symposium on Biological Control of Weeds, La Grande Motte, France, 22-27 April, 2007. pp552-557
Several European Hieracium species, e.g. Hieracium caespitosum Dumort. and Hieracium aurantiacum L., are noxious weeds in North America. A project for the biological control of alien invasive hawkweeds has therefore been initiated in 2000. Five European insect species investigated before their release in New Zealand and two additional gall wasps have been tested on North American test plants. The stolon-tip galling cynipid, Aulacidea subterminalis Niblett (Hym., Cynipidae) proved to be the most specific candidate attacking four Hieracium spp. in the subgenus Pilosella. The authors describe the results of their host-specificity tests.
Grundy T.P. (1989). An economic evaluation of biological control of rose-grain aphid in New Zealand. Agribusiness & Economics Research Unit, Lincoln College, Canterbury. 200 pp.
Gurney W., Crowley P. and Nisbet R. (1992). Locking life-cycles onto seasons: Circle-map models of population dynamics and local adaptation. Journal of Mathematical Biology 30: 251-279.
Haines M.L., Martin J.-F., Emberson R.M., Syrett P., Withers T.M. and Worner S.P. (2007).
Can sibling species explain the broadening of the host range of the broom seed beetle, Bruchidius villosus (F.) (Coleoptera : Chrysomelidae) in New Zealand?
New Zealand Entomologist 30: 5-11
Following introduction into New Zealand for biological control of Scotch broom, Cytisus scoparius, the broom seed beetle, Bruchidius villosus, was found utilising tagasaste, Chamaecytisus palmensis, which was not predicted by host range testing. One possible explanation for these inconsistencies is that more than one species is included within the current concept of B. villosus. However, sequence data from the mitochrondrial gene COI showed a low level of sequence polymorphism (0.8%) between individuals of B. villosus suggesting that B. villosus is a single species with a broader host range than was predicted by host range tests.
Haines M.L., Syrett P., Emberson R.M., Withers T.M., Fowler S.V. and Worner S.P. (2004).
Ruling out a host-range expansion as the cause of the unpredicted non-target attack on tagasaste (Chamaecytisus proliferus) by Bruchidius villosus.
Proceedings of the XI International Symposium on Biological Control of Weeds: 271-276
This paper describes an investigation of the original host-testing procedures. Despite showing a strong preference for Scotch broom, the beetles tested in this study accepted Chamaecytisus proliferus for oviposition allowing us to rule out the possibility that a host range expansion has occurred.
Halpen S.L. and Underwood N. (2006). Approaches for testing herbivore effects on plant population dynamics. Journal of Applied Ecology 43: 922–929
Harley K.L.S. and Forno I.W. (1992). Biological control of weeds. A handbook for practitioners and students. Inkata Press, Melbourne. 74 pp.
Harris P. (1990). Environmental impact of introduced biological control agents. Pp. 289-300 In: Critical issues in biological control, M. Mackauer, L.E. Ehler and J. Roland (Ed.) Intercept, Andover, Hampshire, UK.
Harris P. (1991). Classical biological control of weeds: its definition, selection of effective agents, and administrative-political problems. Canadian Entomologist 123: 827-849.
Harris P. and Zwölfer H. (1968). Screening of phytophagous insects for biological control of weeds. Canadian Entomologist 100: 295-303.
Harrison L., Moeed A. and Sheppard A. (2005).
Regulation of the release of biological control agents of arthopods in New Zealand and Australia.
Pp. 715-725 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.
Regulation of biological control agents in New Zealand is legislated by the Hazardous Substances and New Organisms Act 1996 and administered by the Environmental Risk Management Authority (ERMA New Zealand). In Australia the Department of the Environment and Heritage and the Agriculture Fisheries and Forestry Australia - Australian Quarantine Inspection Service jointly regulate the import, testing and release of biological control agents under the Quarantine Act 1908, Wildlife Protection (Regulation of Exports and Imports) Act 1982 and Biological Control Act 1984. The 2 regulatory systems are compared in this paper highlighting the pivotal role of information from the host specificity testing in the decision making process and the valuable opportunity for researchers to interact with the public.
Harvey C.D., Alameen K.M, and Griffin C.T. (2012). The impact of entomopathogenic nematodes on a non-target, service-providing longhorn beetle is limited by targeted application when controlling forestry pest Hylobius abietis. Biological Control 62: 173-182.
Harwood J.D. and Obrycki J.J. (2005).
Quantifying aphid predation rates of generalist predators in the field.
European Journal of Entomology 102: 335-350.
Over 100 investigations have utilized gut-content analysis to estimate aphid predation rates by predators including gut dissection, radio-labelling of prey, dissection of faecal samples, electrophoresis, stable isotope analysis and use of polyclonal antisera, monoclonal antibodies. Advances in molecular biology have enabled the detection of species-specific DNA sequences and use of these applications to quantify predation by aphidophagous predators is reviewed.
Hatcher P.E. (1995). Three-way interactions between plant pathogenic fungi, herbivorous insects and their host plants. Biological Reviews 70: 639-694
Hatcher P.E. and Melander B. (2003). Combining physical, cultural and biological methods: prospects for integrated non-chemical weed management strategies. Weed Research 43: 303-322
Hawkins B.A. and Cornell H.V. (1994). Maximum parasitism rates and successful biological control. Science 266: 1886.
Hawkins B.A. and Cornell H.V. (1999). Theoretical approaches to biological control. Cambridge University Press, Cambridge, UK. 412 pp.
Hawkins B.A. and Marino P.C. (1997).
The colonization of native phytophagous insects in North America by exotic parasitoids.
Oecologia 112: 566.
Classical biological control could have a major environmental cost if introduced natural enemies colonize and disrupt native systems. Impacts were evaluated based on the extent to which exotics have acquired native hosts. The ability of six biological and ecological variables to predict whether or not a parasitoid will move onto natives was evaluated. It was concluded that given the quality of the data available either now or in the foreseeable future, coupled with inherent stochasticity in host shifts by parasitoids, there are no rules of thumb to assist biological control workers in evaluating if an introduced parasitoid will colonize native insect communities.
Haye T., Goulet H., Mason P.G. and Kuhlmann U. (2005).
Does fundamental host range match ecological host range? A retrospective case study of a Lygus plant bug parasitoid.
Biological Control 35: 55-67.
Using the retrospective case study of Peristenus digoneutis (Hymenoptera: Braconidae) introduced in the United States for biological control of native Lygus plant bugs (Hemiptera: Miridae), laboratory and field studies were conducted in the area of origin to evaluate whether the fundamental host range of P. digoneutis matches its ecological host range. To confirm the validity of the fundamental host range, the ecological host range of P. digoneutis in the area of origin was investigated. Peristenus digoneutis was reared from 10 hosts, including three Lygus species and seven non-target hosts from the subfamily Mirinae. Despite the fact that laboratory tests demonstrated a high parasitism level in non-targets, ecological assessments in both North America and Europe suggest a much lower impact of P. digoneutis on non-target mirids. It was concluded that ecological host range studies in the area of origin provide useful supplementary data for interpreting pre-release laboratory host range testing.
Haye T., Kuhlmann U., Goulet H. and Mason P.G. (2006).
Controlling Lygus plant bugs (Heteroptera : Miridae) with European Peristenus relictus (Hymenoptera : Braconidae) in Canada - risky or not ?
Bulletin of Entomological Research 96: 187-196
The European Peristenus relictus Loan (syn. P. stygicus) has been considered for biological control of Lygus plant bugs native to Canada. Field and laboratory studies were carried out to compare fundamental with ecological host range.
Haye T., van Achterberg C., Goulet H., Barratt B.I.P. and Kuhlmann U. (2006).
Potential for classical biological control of the potato bug Closterotomus norwegicus (Hemiptera: Miridae): description, parasitism and host specificity of Peristenus closterotomae sp. n. (Hymenoptera: Braconidae).
Bulletin of Entomological Research 96: 421–431
The potato bug, Closterotomus norwegicus (Gmelin) (Hemiptera: Miridae) is an introduced pest of lucerne, white clover and lotus seed crops in New Zealand and a key pest of pistachios in California, USA. A total of eight parasitoids, including six from the genus Peristenus (Hymenoptera: Braconidae) and two hyperparasitoids from the genus Mesochorus (Hymenoptera: Ichneumonidae), were reared from C. norwegicus nymphs collected in northern Germany. With a proportion of more than 85% of all C. norwegicus parasitoids, Peristenus closterotomae (Hymenoptera: Braconidae), a new species, was the most dominant parasitoid, whereas other parasitoid species only occurred sporadically. Parasitism caused by P. closterotomae was on average 24% (maximum 77%). To assess the host specificity of parasitoids associated with C. norwegicus, the parasitoid complexes of various Miridae occurring simultaneously with C. norwegicus were studied. Peristenus closterotomae was frequently reared from Calocoris affinis (Herrich-Schaeffer), and a few specimens were reared from Calocoris roseomaculatus (De Geer) and the meadow plant bug, Leptopterna dolobrata (Linnaeus) (all Hemiptera: Miridae). The remaining primary parasitoids associated with C. norwegicus were found to be dominant in hosts other than C. norwegicus. Whether nymphal parasitoids may potentially be used in a classical biological control initiative against the potato bug in countries where it is introduced and considered to be a pest is discussed.
Heard T.A. (2000). Concepts in insect host-plant selection behavior and their application to host specificity testing. Pp. 1-10 In: Host-specificity testing of exotic arthropod biological control agents: the biological basis for improvement in safety, R.G. Van Driesche, T. Heard, A.S. McClay and R. Reardon (Ed.) USDA Forest Service Bulletin, Morgantown, West Virginia, USA.
Heard T.A. and Van Klinken R.D. (1998). An analysis of test designs for host range determination of insects for biological control of weeds. Pp. 539-546 In: Proceedings of the Sixth Australasian Applied Entomological Research Conference, M.P. Zalucki, R.A.I. Drew and G.G. White (Ed.) Brisbane, University of Queensland.
Henneman M.L. and Memmott J. (2001). Infiltration of a Hawaiian community by introduced biological control agents. Science 293: 1314-1316.
Herting B. and Simmonds F.J. (1972). A Catalogue of Parasites and Predators of Terrestrial Arthropods. Section A. Host or prey/enemy. Vol. II. Homoptera. CIBC. 210 pp.
Hill M.G. (1988). Analysis of the biological control of Mythimna separata (Lepidoptera: Noctuidae) by Apanteles ruficrus (Braconidae: Hymenoptera) in New Zealand. Journal of Applied Ecology 25: 197-208
Hill M.P. and Hulley P.E. (1995).
Host-range extension by native parasitoids to weed biocontrol agents introduced to South Africa.
Biological Control 5: 297-302.
Host range extension by native parasitoids to insect biocontrol agents of weeds in South Africa were examined. All host range extensions were from native herbivores and occurred within 3 years of release. Poorly concealed endophytic agents were most susceptible to attack, whereas exposed feeders were relatively free from attack.
Hill R.L. (1977). Parasite helps control armyworm. New Zealand Journal of Agriculture 134: 21-23
Hill R.L. (1982). Seasonal patterns of phytophage activity on gorse (Ulex europaeus) and host plant quality. Pp. 237-242. In: Proceedings of the 5th International Symposium on Insect-Plant Relationships, J.H. Visser and A.K. Minks (Ed.) Wageningen, The Netherlands 1-4 March 1982.
Hill R.L. (1999). Minimising uncertainty - in support of no-choice tests. Pp. 1-10 In: Host specificity testing in Australasia: towards improved assays for biological control, W.T.M., L. Barton Browne and J. N. Stanley (Ed.) CRC for Tropical Pest Management, Brisbane, Australia.
Hill R.L. (2008). Application to release from containment a beetle, Lema obscura F. (Chrysomelidae), for the biological control of the weed tradescantia (Tradescantia fluminensis). http://www.ermanz.govt.nz/appfiles/execsumm/pdf/NOR07001-002.pdf
Hill R.L. and Gourlay A.H. (2002). Host-range testing, introduction, and establishment of Cydia succedana (Lepidoptera: Tortricidae) for biological control of gorse, Ulex europaeus L., in New Zealand. Biological Control 25: 173-186
Hill R.L. and O'Donnell D.J. (1991). Reproductive isolation between Tetranychus lintearius and two related mites, T. urticae and T. turkestani (Acarina: Tetranychidae). Experimental and Applied Acarology 11: 241-251
Hill R.L. and Sandrey R.A. (1986). The costs and benefits of gorse Proceedings of the New Zealand Weed and Pest Control Conference 39: 70-73
Hill R.L., Cumber R.A. and Allan D.J. (1985). Parasitoids introduced to control larvae of the Noctuidae (Lepidoptera) in New Zealand (1968-1978). DSIR Entomology Division, 24 p.
Hill R.L., Gordon A.J. and Neser S. (2000). The potential role of Bruchophagus acaciae (Cameron) (Hymenoptera: Eurytomidae) in the integrated control of Acacia species in South Africa. Pp. 919-929 In: Proceedings of the X International Symposium on Biological Control of Weeds, N.R. Spencer (Ed.) Montana State University, Bozeman, Montana, USA.
Hill R.L., Gourlay, A.H. and Wigley P. (1989). The introduction of gorse spider mite Tetranychus lintearius for biological control of gorse. Proceedings of the 42nd New Zealand Weed and Pest Control Conference: 137–139.
Hill R.L., Gourlay, A.H. and Winks, C.J. (1993). Choosing gorse spider mite strains to improve establishment in different climates. In: Proceedings of the 6th Australasian Grasslands Invertebrate Ecology Conference, R.A Prestidge (Ed.). AgResearch, Hamilton, New Zealand.
Hill R.L., Markin G.P., Gourlay A.H., Fowler S.V. and Yoshioka E. (2001). Host range, release, and establishment of Sericothrips staphylinus Haliday (Thysanoptera: Thripidae) as a biological control agent for gorse, Ulex europaeus L. (Fabaceae), in New Zealand and Hawaii. Biological Control 21: 63-74
Hill R.L., O'Donnell D.J., Gourlay A.H. and Speed C.B. (1995). The suitability of Agonopterix ulicetella (Lepidoptera: Oecophoridae) as a biological control agent for Ulex europaeus Fabaceae: Genisteae) in New Zealand. Biocontrol Science and Technology 5: 3-10.
Hill R.L., Wittenberg R. and Gourlay A.H. (2001). Biology and host range of Phytomyza vitalbae and its establishment for the biological control of Clematis vitalba in New Zealand. Biocontrol Science and Technology 11: 459-473
Hoagland R.E., Weaver M.A. and Boyette C.D. (2007).
Myrothecium verrucaria fungus: a bioherbicide and strategies to reduce its non-target risks.
Allelopathy Journal 19: 179-192
The herbicidal activity of an M. verrucaria (MV) strain originally isolated from sicklepod (Senna obtusifolia) was evaluated against kudzu (Pueraria lobata) and several other weeds. Tests showed high levels of efficacy of MV and a large range of nontarget, young, woody plant species from several plant families ranged from non-susceptible to moderately susceptible. Although MV possesses desirable bioherbicidal traits, this isolate also produces undesirable mycotoxins, i.e. trichothecenes. Future approaches to possibly reduce or eliminate these mycotoxins are discussed.
Hoddle M. (2004). Restoring balance using exotic species to control invasive exotic species. Conservation Biology 18: 38-49
Hoddle M.S. (2004). Analysis of fauna in the receiving area for the purpose of identifying native species that exotic natural enemies may potentially attack. Pp. 24-39 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.
Hoelmer K.A., Schuster D.J. and Ciomperlik M.A. (2008).
Indigenous parasitoids of Bemisia in the USA and potential for non-target impacts of exotic parasitoid introductions.
Pp. 307-324 In: Classical biological Control of Bemisia tabaci in the United States - A review of interagency research and implementation, J. Gould, K. A. Hoelmer and J. Goolsby (Ed.) Springer, Dordtrecht
Surveys to document the presence and species composition of native natural enemies were conducted prior to the introduction of non-indigenous agents against sweetpotato whitefly, Bemisia tabaci biotype B, in the USA. The greatest diversity of native parasitoid species attacking B. tabaci was reported in Florida, where there was most diversity of invasive whitefly species established in Florida. Only two or three parasitoid species were responsible for the majority of parasitism of B. tabaci within any given region of the USA. The predominant species attacking B. tabaci prior to the introduction of new Palearctic parasitoid species were Eretmocerus tejanus (in Texas), Eretmocerus eremicus (Arizona and California), Eretmocerus sp. (undescribed, southeast USA), Encarsia pergandiella/Enc. tabacivora (southeastern USA and Texas), and Encarsia luteola (southwestern USA). Surveys up to 2001 (California) and 2003 (Texas), showed that the exotic species that were introduced have remained limited to their intended target.
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.
Hoffmeister T.S., Babendreier D. and Wajnberg E. (2006). Statistical tools to improve the quality of experiments and data analysis for assessing non-target effects. Pp. 222-240 In: Environmental impact of invertebrates for biological control of arthropods: methods and risk assessment, F. Bigler, D. Babendreier and U. Kuhlmann (Ed.) CABI Publishing, Wallingford, Oxford.
Hokkanen H.M.T. and Lynch J.M. (1995). Biological Control: Benefits and Risks. Cambridge University Press, Cambridge, UK. 304pp.
Hokkanen H.M.T. and Pimentel D. (1989). New associations in biological control: Theory and practice. Canadian Entomologist 121: 829-840
Holt R.D. and Hochberg M.E. (1997). When is biological control evolutionarily stable (or is it)? Ecology 78: 1673-1683.
Holt R.D. and Hochberg M.E. (2001). Indirect interactions, community modules and biological control: a theoretical perspective. Pp. 13-37 In: Evaluating indirect ecological effects of biological control, E. Wajnberg, J. K. Scott and P. C. Quimby (Ed.) CABI Publishing, Wallingford, Oxon., UK
Hoogendorn M. and Heimpel G.E. (2003). PCR-Based gut content analysis of insect predators: A field study. Pp. 91-97 In: Proceedings of the 1st International Symposium on Biological Control of Arthropods, R. Van Driesche (Ed.) Forest Health Technology Enterprise Team, Morgantown, West Virginia.
Hope K.J. and Olckers T. (2011). 2011. Gargaphia decoris (Hemiptera: Tingidae) from two South American provenances are equally safe for release against the invasive tree, Solanum mauritianum (Solanaceae). African Entomology 19: 106-112.
Hopper K.R. (1995). Potential impacts on threatened and endangered insect species in the United States from introductions of parasitic Hymenoptera for the control of insect pests. Pp. 64-74 In: Biological Control: Benefits and Risks, H.M.T. Hokkanen and J.M. Lynch (Ed.) Cambridge University Press, Cambridge, UK.
Hopper K.R. (1998). Assessing and improving the safety of introductions for biological control. Pp. 501-510 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.
Hopper K.R. (2001). Research needs concerning non-target impacts of biological control introductions. Pp. 39-56 In: Evaluating indirect ecological effects of biological control, E. Wajnberg, J.K. Scott and P.C. Quimby (Ed.) CABI Publishing, Wallingford, Oxon., UK.
Hopper K.R. and Roush R.T. (1993).
Mate finding, dispersal, number released, and the success of biological control introductions.
Ecological Entomology 18: 321-331.
Published data were analysed and a mathematical model of the population dynamics of introduced parasitoids were used to investigate the reason for failure of biological control introductions. Allee effects result in small populations becoming extinct because low densities lead to failure to mate, a male-biased sex ratio, and sometimes extinction. For many groups of parasitoids the proportion of populations that established increased with the number of parasitoids per release and the total number released. An analysis of past introductions and the reaction-diffusion model both suggested a threshold of about 1000 insects per release to ensure establishment of introduced parasitoids.
Hopper K.R. and Wajnberg E. (2006). The risks of interbreeding and methods for determination. In press In: Environmental Impact of Arthropod Biological Control: Methods and Risk Assessment, U. Kuhlmann, F. Bigler and D. Babendreier (Ed.) CABI Bioscience, Delemont, Switzerland.
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.
Hopper K.R., Farias A.M.I., Woolley J.B., Heraty J.M. and Britch S.C. (2005).
Genetics: relation of local populations to the whole "species" - implications for host range tests
Pp. 665-671 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 literature on variation in host specificity among populations and sibling species of parasitoids is reviewed and the evolution and genetics of host specificity in Aphelinus varipes and A. albipodus (Hymenoptera: Aphelinidae) discussed. The take-home lessons for biological control are: parasitoids in what appears to be a single species, but collected from widely different geographical regions or from different host species, may differ greatly in host specificity and thus should be tested separately; and allopatric sibling species with different patterns of host use may introgress if placed in sympatry, which could lead to evolutionary changes in host use.
Howarth F.G. (1983). Classical biological control: panacea or Pandora's box. Proceedings of the Hawaiian Entomological Society 24: 239-244.
Howarth F.G. (1991). Environmental impacts of classical biological control. Annual Review of Entomology 36: 489-509.
Howarth F.G. (1992). Environmental impact of species purposefully introduced for biological control of pests. Pacific Science 46: 388-389.
Howarth F.G. (2001). Environmental issues concerning the importation of non-indigenous biological control agents. Pp. 70-99 In: Balancing nature: assessing the impact of importing non-native biological control agents (an international perspective), J.A. Lockwood, F.G. Howarth and M. Purcell (Ed.) Entomological Society of America, Lanham, Maryland.
Howarth F.G. and Ramsay G.W. (1991). The conservation of island insects and their habitats. Pp. 71-107 In: The Conservation of Insects and their Habitat, N.M. Collins and J.A. Thomas (Ed.) Academic Press, London.
Hufbauer R.A. (2002). Evidence for nonadaptive radiation in parasitoid virulence following a biological control introduction. Ecological Applications 12: 66-78.
Hufbauer R.A. and Roderick G.K. (2005).
Microevolution in biological control: Mechanisms, patterns, and processes.
Biological Control 35: 227-239.
The four fundamental processes of microevolution are discussed in relation to how they interact in the context of biological control. The types of experiments that can address questions are discussed and ways of using microevolution to define risks, and enhance efficacy and safety of biological control.
Hunt E.J., Kuhlmann U., Sheppard A., Qin T.-K., Barratt B.I.P., Harrison L., Mason P.G., Parker D. and Goolsby J. (2008).
Review of invertebrate biological control agent regulation in Australia, New Zealand, Canada and the USA: recommendations for a harmonised European regulatory system.
Journal of Applied Entomology 132: 89-123
In this paper the current regulatory processes operating in Australia, New Zealand, Canada and the USA are reviewed with a view to allowing countries of Europe to benefit from the years of experience that these countries have in IBCA regulation. Recommendations are made based on features of the regulatory processes in each of the four countries that work well and that could be adopted to generate a workable regulatory system in Europe.
Ireson J.E., Gourlay A.H., Holloway R.J., Chatterton W.S., Foster S.D. and Kwong R.M. (2008).
Host specificity, establishment and dispersal of the gorse thrips, Sericothrips staphylinus Haliday (Thysanoptera: Thripidae), a biological control agent for gorse, Ulex europaeus L. (Fabaceae), in Australia.
Biological Control 45: 460-471
Sericothrips staphylinus was released as a biological control agent for Ulex europaeus in New Zealand and Hawaii following tests which showed it was narrowly oligophagous. To determine the suitability for release in Australia, further host specificity tests were conducted on Australian plants which confirming host specificity and it was released in Tasmania during January 2001. Releases of 10, 30, 90, 270 and 810 adults showed that establishment could be achieved with as few as 10 thrips. 250 thrips were chosen as the minimum number for release because this release size produced close to the maximum population growth. Surveys in early 2007 recovered S. staphylinus from 80% of 30 sites in Tasmania, but densities were low with no evidence of visible plant damage. The survey results indicated that S. staphylinus is a sedentary, latent species characterised by steady densities and low levels of damage to its host plant. Its efficacy as a biological control agent on gorse may be restricted primarily by 'bottom up' effects of plant quality limiting its rate of natural increase and an inability of the thrips to reach large, damaging populations under field conditions.
Jacas J.A., Urbaneja A. and Viñuela E. (2006).
History and future of introduction of exotic arthropod biological control agents in Spain: A dilemma?
BioControl 51: 1-30.
More success for IBCAs has been achieved with seasonal inoculative releases (50.0% of cases) than for classical biological control programs (17.1% of cases). Concerns about potential non-target effects but post-release evaluation has often been insufficient to draw any conclusions about them. Most of the biological control agents introduced in Spain were parasitoids (n = 53), and the remainder predators (n = 12). Only four parasitoids are considered monophagous. Using information from literature and the internet, the mean number of host species parasitized by parasitoids is 15.2, Therefore, polyphagy appears to be quite common among the IBCAs that have been introduced in Spain.
Jallow M.F.A. and Zalucki M.P. (1996). Within- and between-population variation in host-plant preference and specificity in Australian Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae). Australian Journal of Zoology 44: 503-519.
Jarvis P.J., Fowler S.V., Paynter Q. and Syrett P. (2006). Predicting the economic benefits and costs of introducing new biological control agents for Scotch broom Cytisus scoparius into New Zealand. Biological Control 39: 135-146
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.
Jetter K. (2005).
Economic framework for decision making in biological control.
Biological Control 35: 348-357.
A technique known as threshold cost/benefit analysis is presented and an example on how to apply this method is illustrated using the yellow starthistle biological control program. The results show that incorporating uncertainty into the analysis can have a significant impact on the decision to undertake a biological control program.
Johnson M.T., Follett P.A., Taylor A.D. and Jones V.P. (2005).
Impacts of biological control and invasive species on a non-target native Hawaiian insect.
Oecologia 142: 529-540.
Adverse impacts on endemic Hawaiian koa bug, Coleotichus blackburniae White (Hemiptera: Scutelleridae), by parasitoids introduced for control of the southern green stink bug, Nezara viridula (L.) (Hemiptera: Pentatomidae) were examined using life tables. Self-introduced generalist egg predators, had the greatest impacts on C. blackburniae populations. Effects of intentionally introduced parasitoids were relatively minor, although the tachinid T. pilipes showed potential for large impacts at individual sites. In retrospect, non-target attacks by biological control agents on C. blackburniae were predictable, but not the environmental range and magnitude of impacts.
Julien M.H., Skarratt B. and Maywald G.F. (1995). Potential geographical distribution of alligator weed and its biological control by Agasicles hygrophila. Journal of Aquatic Plant Management 2: 55-60
Julien, M. and Griffiths M.W. (1999). Biological Control of Weeds. A World Catalogue of Agents and their Target Weeds. CABI Publishing, Wallingford, UK. 223 p.
Kairo M.T.K., M.J.W. Cock. and M.M. Quinlan (2003).
An assessment of the use of the Code of Conduct for the Import and Release of Exotic Biological Control Agents (ISPM No. 3) since its endorsement as an international standard
Biocontrol News and Information 24: 27N
This review assesses the use of ISPM No. 3 since it became an international standard for making decisions about biological control agents. It was found that ISPM No. 3 or similar national procedures have been applied in most cases to support decisions regarding import and release of exotic biological control agents since 1996. It has provided a mechanism for formalizing current good practice and provided internationally accepted standards to countries with little experience in implementing biological control. Limitations in implementation of ISPM No. 3 included lack of technical capacity and appropriate quarantine facilities. Case studies for decisions by Kenya, Colombia, the Caribbean, Yemen, Samoa and Brazil are discussed to provide further insights into the use of ISPM No. 3 over its first seven years.
Kaufman L.V. and Wright M.G. (2009).
The impact of exotic parasitoids on populations of a native Hawaiian moth assessed using life table studies.
Oecologia 159: 295-304
This study investigated the impact of introduced Hymenoptera parasitoids on the Hawaiian moth Udea stellata (Butler) which has seven alien parasitoids associated with it. The study determined the relative contribution of the seven parasitoid species to the population dynamics of U. stellata. The factors found to contributed to total mortality were: disappearance (42.1%), death due to unknown reasons during rearing (16.5%) and parasitism (4.9%). Adventive parasitoids inflicted greater total larval mortality attributable to parasitism (97.0%) than purposely introduced species (3.0%).
Kaupp W.J., Barber K.N., Fick W.E., Ebling P.M., Ladd T.R. and Holmes .SB. (2011). Host-range testing of a mixture of two nucleopolyhedroviruses of Choristoneura fumiferana (Lepidoptera: Tortricidae). Canadian Entomologist 143: 165-177.
Kay N. and Hill R.L. (in press). The disintegration of the Scrophulariaceae and the biological control of Buddleja davidii. In: Proceedings of the XII International Symposium on Biological Control of Weeds, (Ed.) Montpellier, France.
Kean J.M. and Barlow N.D. (2000). Can host-parasitoid metapopulations explain successful biological control? Ecology 81: 2188-2197.
Kean J.M. and Kumarasinghe L. (2007). Predicting the seasonal phenology of fall webworm (Hyphantria cunea) in New Zealand. New Zealand Plant Protection 60: 279-285.
Kelleher J.S. and Hulme M.A. (1984). Biological control programmes against insects and weeds in Canada 1969-1980. CAB, Farnham Common, UK. 410 pp.
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
Kimberling D.N. (2004).
Lessons from history: predicting successes and risks of intentional introductions for arthropod biological control.
Biological Invasions 6: 301-318.
The US National Invasive Species Management Plan (2001) calls for better screening methods for biological control agent introductions. Literature searches were used to develop a database of 13 life history traits and 8 descriptive variables for 87 insect biological control species in the United States. Models for predicting success in controlling a target species and likelihood of nontarget effects were developed using logistic regression. The most important life history traits for predicting success included host specificity, whether the agent was a predator or parasitoid, and number of generations per year. Traits important for predicting nontarget effects included sex ratio of progeny and the presence of native natural enemies.
Kindlmann P. and Dixon A.F.G. (1999). Generation time ratios - determinants of prey abundance in insect predator-prey interactions. Biological Control 16: 133-138.
Kitt J.T. and Keller M.A. (1998). Host selection by Aphidius rosae Haliday (Hym., Braconidae) with respect to assessment of host specificity in biological control. Journal of Applied Entomology 122: 57-63
Kluge R.L. (2000).
The future of biological control of weeds with insects: no more 'paranoia', no more 'honeymoon'.
Proceedings of the X International Symposium on Biological Control of Weeds: 459-467
Two issues are considered in this paper: first the 'paranoia' about the threat of biocontrol agents to non-target plant species, and second, the 'honeymoon' regarding the lack of accountability for projects that failed to achieve their desired objectives. Proposals are made to deal with five current pressures that biological control of weeds is facing, the image of the discipline, host specificity verification, selection of candidates, funding and regulatory requirements.
Knudsen I.M.B., Hockenhull J., Jensen D.F., Gerhardson B., Hokeberg M., Tahvonen R., Teperi E., Sundheim L. and Henriksen B. (1997). Selection of biological control agents for controlling soil and seed-borne diseases in the field. European Journal of Plant Pathology 103: 775.
Knutson L. and Coulson J.R. (1997).
Procedures and policies in the USA regarding precautions in the introduction of classical biological control agents.
EPPO/CABI workshop on safety and efficacy of biological control in Europe 27: 133-142.
Scientists, administrators and others have long recognized the need to ensure that natural enemies of weeds do not attack commercially or horticulturally important plants and to ensure that natural enemies of insects do not attack beneficial species. Procedures for testing the host specificity of the natural enemies of weeds in their area of origin, before shipment to the country of release, have been developed to quite high levels of reliability but there is need for further improvements. All applications for permission to introduce biocontrol agents are examined by an Inter-agency Technical Advisory Committee for Biological Control of Weeds (TAGBCW) before import permits are issued by the relevant authority (USDA APHIS/PPQ). TAGBCW review includes study of research plans and of lists of host plants for testing host specificity. It is considered that there is relatively little need for host-range testing of most parasitoids because these are generally co-evolved and intimately related organisms that are restricted to one or a few host species. Concern about the potential impact of oligophagous predators on non-target organisms has increased recently and is a developing field of research. The import permit system in the USA is presented, and two suggestions for changes in European permit procedures are suggested.
Kriticos D.J. (2003). The roles of ecological models in evaluating weed biological control agents and projects. Pp. 69-74 In: Improving the selection, testing and evaluation of weed biological control agents. Proceedings of the CRC for Australian Weed Management biological control of weeds symposium and workshop, J. H. Spafford and D. T. Briese (Ed.) University of Western Australia, Perth, Australia, CRC for Australian Weed Management, Adelaide, Australia, 13 September 2002.
Kriticos D.J. and Randall R.P. (2001). A comparison of systems to analyse potential weed distributions. Pp. 61-79 in: Groves R.H., Panetta F.D. and Virtue J.G. ed. Weed Risk Assessment. CSIRO Publishing, Melbourne, Australia.
Kriticos D.J., Brown J.R., Radford I.D. and Nicholas M. (1999). Plant population ecology and biological control: Acacia nilotica as a case study. Biological Control 16
Krombein K.V., Hurd P.D., Smith D.R. and Burks B.D. (1979). Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, USA
Krugner R., Johnson M.W., Groves R.L. and Morse J.G. (2008).
Host specificity of Anagrus epos : a potential biological control agent of Homalodisca vitripennis.
Biocontrol 53: 439-449
Anagrus epos Girault (Hymenoptera: Mymaridae) is a candidate for a classical biological control program targeting the glassy-winged sharpshooter (GWSS), Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae), in California. Mass production of GWSS is expensive and labor-intensive and a factitious host that is more economical to produce is required. The use of a factitious host and potential nontarget effects of this generalist parasitoid are discussed.
Kuhlmann U., Mason P.G. and Greathead D.J. (1998).
Assessment of potential risks for introducing European Peristenus species as biological control agents of native Lygus species in North America: a cooperative approach.
Biocontrol News and Information 19: 83n-90n.
The pest status of Lygus species in North America and history of European collections and importations of Peristenus species into North America is described. Strategies and methods for host specificity testing of parasitoids are outlined and discussed in relation to the selection of non-target and native Lygus species for testing with Peristenus parasitoids. European Peristenus species are identified and their life histories outlined. It is concluded that cooperative research in Europe and North America is needed to assess the potential risks for the introduction of European Peristenus species for control of Lygus species in North America.
Kuhlmann U., Mason P.G., Hinz H.L., Blossey B., De Clerck-Floate R.A., Dosdall L.M., McCaffrey J.P., Schwarzlaender M., Olfert O., Brodeur J., Gassmann A., McClay A.S. and Wiedenmann R.N. (2006).
Avoiding conflicts between insect and weed biological control: selection of non-target species to assess host specificity of cabbage seedpod weevil parasitoids.
Journal of Applied Entomology 130: 129-141
Classical biological control of insect pests and weeds may lead to potential conflicts, where insect pests are closely related to weed biological control agents, e.g. the cabbage seedpod weevil, Ceutorhynchus obstrictus (Marsham) in North America, which belongs to the same subfamily, Ceutorhynchinae, as a number of agents introduced or proposed for introduction for invasive weed species. This paper describes a procedure to develop a non-target species test list for screening candidate entomophagous biological control agents of a herbivore pest insect to simultaneously evaluate non-target potential on weed biological control agents and other non-target species.
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.
Kuris A.M. (2003).
Did biological control cause extinction of the coconut moth, Levuana iridescens, in Fiji?
Biological Invasions 5: 133-141.
Biological control of Levuana iridescens, the endemic coconut moth of Fiji, was so successful that this pest of the copra crop had been reduced to almost undetectable levels by the tachinid fly, Bessa remota. This example is presented in the modern literature as the first and best documented extinction of a species due to biological control and portrayed as an example of the highly controversial practice of neoclassical biological control. However, the moth was likely not to be native to Fiji and might have spread to other island groups in the South Pacific, and may not be extinct. A strategy to search for L. iridescens populations is proposed and development of biological control of B. remota, using hyperparasitoids, is possible.
Kuske S., Babendreier D., Edwards P., Turlings T.C.J. and Bigler F. (2004).
Parasitism of non-target Lepidoptera by mass released Trichogramma brassicae and its implication for the larval parasitoid Lydella thompsoni.
Biocontrol 49: 1-19.
The release of high numbers of the egg parasitoid Trichogramma brassicae Bezd. (Hym. Trichogrammatidae) to control the European corn borer, Ostrinia nubilalis Hb. (Lep.: Crambidae) in maize has raised concerns about potential negative effects on native natural enemies, particularly Lydella thompsoni Herting (Dipt.: Tachinidae) Inundative releases of T. brassicae coincide with the oviposition period of the alternative hosts of the tachinid. T. brassicae upon which it relies in spring. Laboratory host specificity tests showed that the tachinid's two most abundant spring hosts are successfully parasitised by T. brassicae females in no-choice situations. Field surveys, however, showed that the two spring hosts escape parasitism since their eggs are well hidden or not attractive, and the study concluded that negative effects of inundative releases of T. brassicae on the native tachinid fly L. thompsoni, are unlikely.
Laing J.E. and Heraty J.M. (1981). Establishment in Canada of the parasite Apanteles pedias Nixon on the spotted tentiform leafminer, Phyllonorycter blancardella (F.). Environmental Entomology 10: 933-935.
Lesica P. and Hanna D. (2004).
Indirect effects of biological control on plant diversity vary across sites in Montana grasslands.
Conservation Biology 18: 444-454.
The hypothesis that biological control agents reduce the dominance of the target weed, increasing the native plant diversity was tested. Aphthona nigriscutis was released into grassland sites infested with Euphorbia esula L. on a nature reserve in Montana (U.S.A.) and compared with herbicide treatment. After 5 years, Aphthona release caused a 33-39% decline in Euphorbia aboveground biomass compared with controls at all sites. Other effects of the biocontrol depended on the site. Results suggested that biocontrol reductions in weed dominance was not always associated with increased species diversity. Monitoring of community-level effects should accompany biocontrol introductions on nature reserves.
Lockwood J.A. (1993).
Environmental issues involved in biological control of rangeland grasshoppers (Orthoptera: Acrididae) with exotic agents.
Environmental Entomology 22: 504-518.
Neoclassical biological control with a parasitic wasp and an entomophagous fungus from Australia is now being applied to rangeland grasshoppers in the western USA, and it is predicted that there may be a number of possible nontarget impacts. Adverse effects include competitive suppression or extinction of both native biological control agents and nontarget acridids. Suppression of nontarget acridids may result in loss of biological diversity, existing control of weed species, release of otherwise innocuous acridid species from competitive regulation, disruption of plant community structure, suppression of essential organisms vectored by grasshoppers, and disruption of food chains and other nutrient cycling processes. Given that the value of the rangeland resource depends on the largely unknown ecological processes that underlie its sustainable productivity, there are a number of management techniques that offer a greater probability of success with a markedly lower likelihood of ecological and economic disruption than does neoclassical biological control.
Lockwood J.A. (1996).
The ethics of biological control: Understanding the moral implications of our most powerful ecological technology.
Agriculture and Human Values 13: 2-19.
The system of environmental ethics developed by Johnson (1991) is used to analyse the moral implications of biological control. In this formulation of ethical analysis, species and ecosystems are morally relevant because they are not simply aggregates of individuals, so their processes, properties, and well-being interests are not reducible to the sum of their individual members. Following Johnson's thesis, species and ecosystems have morally relevant interests in surviving and maintaining themselves as integrated wholes with particular self-identities. It is evident that not all biological control efforts are ethically defensible. In general terms, natural biological control is most desirable, followed by augmentative strategies, classical approaches, and finally neoclassical biological control.
Lockwood J.A. (1997).
Competing values and moral imperatives: an overview of ethical issues in biological control.
Agriculture and Human Values 14: 205-210.
The perception and resolution of ethical issues relating to biological control appear to emerge from a set of factors that includes one's ethical viewpoint (anthropocentric or biocentric), agricultural system (industrial or sustainable), economic context (rich or poor), and power structure (expert or public). From this set of parameters at least five major ethical questions can be formulated: (1) How we should regulate and apply biological control given uncertainty regarding environmental impacts; (2) How we balance benefits of biological control to human and ecosystem well-being against the known and anticipated risks; (3) Who should be empowered to develop policies and make decisions; (4) How we can assure a more just distribution of benefits and costs associated with biological control technologies and (5) Whether biological control can be justified as a resource substitution for pesticides or is its ethical application only possible as part of a reconceptualization of agricultural production. These central questions and possible answers are presented in a varied set of provocative analyses by some of the leading thinkers and authorities in their fields.
Lockwood J.A. (2000). Nontarget effects of biological control: what are we trying to miss? Pp. 15-30 In: Nontarget effects of biological control introductions, P.A. Follett and J.J. Duan (Ed.) Kluwer Academic Publishers, Norwell, Massachusetts, USA.
Lockwood J.A., Howarth F.G. and Purcell M. (2001). Balancing nature: Assessing the impact of importing non-target biological control agents (An international perspective). Thomas Say Publications, Lanham, Maryland, USA. 130 pp.
Longworth J.F. (1987). Biological control in New Zealand: policy and procedures. New Zealand Entomologist 10: 1-7.
Loomans A. and Van Lenteren J.C. (2005).
Tools for environmental risk assessment of invertebrate biological control agents: a full and quick scan method.
Pp. 611-619 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 International Standard for Phytosanitary Measures No. 3 (ISPM3) offers a framework for risk assessment and focuses specifically on the shipment, import, export and release of biological control agents. The major challenge in developing risk assessment methodologies is to develop protocols and guidelines that will prevent serious mistakes through import and release of potentially harmful exotics, while at the same time still allowing safe forms of biological control to proceed. A risk assessment methodology for biological control agents should integrate information on the potential of an agent to establish, its abilities to disperse, its host range and its direct and indirect effects on non-targets. A comprehensive risk evaluation method (full scan) for new natural enemies is proposed and, 'quick scan' method for natural enemies already in use.
Loomans A.J.M., Van Lenteren J.C., Bigler F., Burgio G., Hokkanen H.M.T., Thomas M.B. and Editor: Enkegaard E. (2002).
Evaluating environmental risks of biological control introductions: how to select safe natural enemies?
Proceedings of the joint IOBC/WPRS Working Group "Integrated Control in Protected Crops, Temperate Climate" and IOBC/NRS "Greenhouse, Nursery, and Ornamental Landscape IPM Working Group" at Victoria (British Columbia), Canada, 6 9 May 2002; Bulletin OILB/SROP 25: 147-150.
Biological control of greenhouse pests has become a key component of sustainable horticulture in the world. No clear direct adverse effects have been found, but the potential nontarget effects of these releases have been little emphasized. The current situation with respect to selection procedures for importing, mass-rearing and releasing (new) exotic natural enemies is discussed.
Louda S.M. (1998).
Population growth of Rhinocyllus conicus (Coleoptera: Curculionidae) on two species of native thistles in Prairie.
Environmental Entomology 27: 834-841.
Rhinocyllus conicus is a flowerhead weevil deliberately introduced into the USA for the biological control of invasive exotic thistles in the genus Carduus. This study documents the course and magnitude of the weevil population expansion onto nontarget host plants Platte thistle and wavyleaf thistle, a later flowering native species. There is greater phenological synchrony of Platte thistle than wavyleaf thistle flowerhead development with R. conicus oviposition activity. The results suggest that pre-release studies should account for ecological characteristics, such as phenology.
Louda S.M. (1999). Negative ecological effects of the musk thistle biological control agent, Rhinocyllus conicus. Pp. 213-243 In: Nontarget effects of biological control introductions, P.A. Follett and J.J. Duan (Ed.) Kluwer Academic Publishers, Norwell, Massachusetts, USA.
Louda S.M. (2000).
Rhinocyllus conicus - insights to improve predictability and minimize risk of biological control of weeds.
Proceedings of the X International Symposium on Biological Control of Weeds: 187-193
This paper reviews information on the release of Rhinocyllus conicus to control Carduus spp. thistles in North America and suggests 8 lessons for future biological control efforts: (1) better a priori quantification of the occurrence and ecological effects of the weed; (2) improved ecological criteria to supplement the phylogenetic information used to select plants for pre-release testing; (3) increased assessment of potential direct and indirect effects when an agent looks promising but feeding tests suggest it is not strictly monophagous, (4) quantitative evaluation of the efficacy of the proposed biological agent (5) more evidence on alternative control methods; (6) expanded review, both prior to release and periodically afterward; (7) addition of post-release evaluations and redistribution control; and, finally, (8) a rethinking of the situations that qualify for the use of biological control releases.
Louda S.M. and Arnett A.E. (2000).
Predicting non-target ecological effects of biological control agents: evidence from Rhinocyllus conicus.
Proceedings of the X International Symposium on Biological Control of Weeds: 551-567
The significant non-target ecological effects of Rhinocyllus conicus on native species in the northcentral USA provides the opportunity to evaluate factors that might help predict direct non-target effects, and indirect effects mediated by trophic interactions. The relevance for biocontrol risk assessment of at least four important ecological relationships has emerged from these studies so far: (1) ecological and phylogenetic similarity of potential host plants; (2) synchrony of critical stages between insect and potential host plant(s), as well as acceptability; (3) population limiting processes of potential host plants; and, (4) overlap of feeding niche within the native guild of species dependent upon the host plants. In the selection of biocontrol agents, knowledge of the ecological relationships should help to quantify the risks inherent in deliberate introductions of new species.
Louda S.M. and O'Brien C.W. (2002). Unexpected ecological effects of distributing the exotic weevil, Larinus planus (F.), for the biological control of Canada thistle. Conservation Biology 16: 717-727.
Louda S.M., Kendall D., Connor J. and Simberloff D. (1997).
Ecological effects of an insect introduced for the biological control of weeds.
Science 277: 1088-1090.
The weevil Rhinocyllus conicus, introduced to control exotic thistles, has exhibited an increase in host range as well as continuing geographic expansion, in the USA and Canada. Weevils significantly reduce seed production of native thistles, and density of native tephritid flies was lower at high weevil density.
Louda S.M., Pemberton R.W., Johnson M.T. and Follett P.A. (2003).
Nontarget effects - the Achilles' heel of biological control? Retrospective analyses to reduce risk associated with biocontrol introductions.
Annual Review of Entomology 48: 365-396.
Ten projects with quantitative data on nontarget effects were reviewed and the patterns which emerged were discussed. The review lead to six recommendations: avoid using generalists or adventive species; expand host-specificity testing; incorporate more ecological information; consider ecological risk in target selection; prioritize agents; and pursue genetic data on adaptation.
Louda S.M., Rand T.A., Arnett A.E., McClay A.S., Shea K. and McEacherne A.K. (2005).
Evaluation of ecological risk to populations of a threatened plant from an invasive biocontrol insect.
Ecological Applications 15: 234-249.
The risk to the rare, Pitcher's thistle (Cirsium pitcheri) in North America from Rhinocyllus conicus, a biological control weevil now feeding on many native thistles, was evaluated. It was hypothesized that quantification of host specificity and potential phenological overlap between insect and plant would improve assessment of the magnitude of risk. In laboratory host specificity tests, we found no significant difference in R. conicus feeding or oviposition preference between the rare C. pitcheri and the targeted exotic weed (Carduus nutans) or between C. pitcheri and Platte thistle (C. canescens), a native North American species also known to be affected by R. conicus. Results of the study indicated that the weevil poses a serious risk to the threatened C. pitcheri, supporting the suggestion that ecological data can be used to improve the quantification of risk to native nontarget plant populations within the potential physiological host range of a biological control insect.
Louda S.M., Rand T.A., Russell F.L. and Arnett A.E. (2005).
Assessment of ecological risks in weed biocontrol: Input from retrospective ecological analyses.
Biological Control 35: 253-264.
Quantitative retrospective analyses of ongoing biocontrol projects provide a systematic strategy to evaluate and further develop ecological risk assessment. Analyses showed that host range and preference from host specificity tests are not sufficient to predict ecological impact if the introduced natural enemy is not strictly monophagous. The studies demonstrate that the environment influences and can alter host use and population growth, leading to higher than expected direct impacts on the less preferred native host species at several spatial scales, and that easily anticipated indirect effects can be both widespread and significant. It was concluded that intensive retrospective ecological studies provide some guidance for the prospective studies which are needed to assess candidate biological control agent dynamics and impacts
Louda S.M., Simberloff D., Boettner G., Connor J. and Kendall D. (1998). Insights from data on the nontarget effects of the flowerhead weevil. Biocontrol News and Information 19: 70N-71N.
Lynch L.D. and Ives A.R. (1999).
The use of population models in informing non-target risk assessment in biocontrol.
Aspects of Applied Biology 53: 181-188.
The use of models to illustrate risks of biological control to nontarget organisms is shown using two examples. The first model for classical biological control illustrates how the minimum density observed in the nontarget species follows a simple approximation, which is based on the carrying capacity for the target and the searching efficiency of the agent for the nontarget host. The second model looks at how augmentation methods of biocontrol may have impacts on the density of non-target.
Lynch L.D. and Thomas M.B. (2000). Nontarget effects in the biocontrol of insects with insects, nematodes and microbial agents: the evidence. Biocontrol News and Information 21: 117N-130N
Lynch L.D., Hokkanen H.M.T., Babendreier D., Bigler F., Burgio G., Gao Z.-H., Kuscke S., Loomans A., Menzler-Hokkanen I., Thomas M.B., Tommasini G., Waage J.K., Van Lenteren J.C. and Zeng Q.-Q. (2001). Insect biological control and non-target effects: a European perspective. Pp. 99-125 In: Evaluating indirect ecological effects of biological control, E. Wajnberg, J. K. Scott and P. C. Quimby (Ed.) CABI Publishing, Wallingford, Oxon., UK.
Mafokoane L.D., Zimmermann H.G. and Hill M.P. (2007).
Development of Cactoblastis cactorum (Berg) (Lepidoptera : pyralidae) on six north American Opuntia species.
African Entomology 15: 295-299
The recent arrival and spread of Cactoblastis cactorum in North America has raised concerns for the native Opuntia species. The host range of the moths was examined in South Africa. Results showed that although O. ficusindica is the preferred host for C. cactorum in South Africa, the moth is nevertheless able to utilize several other species of Opuntia as hosts.
Malone L.A., Todd J.H., Burgess E.P.J., Walter C., Wagner A. and Barratt B.I.P. (2010). Developing risk hypotheses and selecting species for assessing non-target impacts of GM trees with novel traits: the case of altered-lignin pine trees. Environmental Biosafety Research 9: 181-198.
Mansfield S. and Mills N.J. (2004). A comparison of methodologies for the assessment of host preference of the gregarious egg parasitoid Trichogramma platneri. Biological Control 29: 332-340.
Marohasy J. (1996).
Host shifts in biological weed control: real problems, semantic difficulties or poor science?
International Journal of Pest Management 42: 71-75.
Many biologists perceive organisms as constantly evolving and therefore consider the host plant ranges of biological control agents as likely to undergo adaptive changes should environmental conditions change, after successful biological control. However, despite the introduction of over 600 insect species from one geographic region to another for biological weed control during this century, there are relatively few documented cases of changes in host plant range. It is concluded that apparent additions to host range can be explained in terms of established behavioural concepts of preadaptation, threshold change resulting from host deprivation, and effects of experience (learning). The study highlights the inappropriate term 'host shift' and it is concluded that evidence from biological weed control contradicts some aspects of ecological and evolutionary theory.
Marohasy J. (1998).
The design and interpretation of host-specificity tests for weed biological control with particular reference to insect behaviour.
Biocontrol News and Information 19: 13-20.
Current host specificity procedures are reviewed and a new procedure proposed that takes into account mechanisms known to underlie the behavioural process of host plant finding and acceptance. This minimizes the risk of rejection of safe insect species or the release of potentially unsafe insect species for weed biological control. The period of time between the acceptance of the target weed and lower-ranked plant species by candidate biological control agents, can also be determined. This period may be as important a measure of specificity as the actual number of plant species susceptible to attack.
Mason P.G. and Kuhlmann U. (2002). Regulations are necessary for biological control agents. IOBC/wprs Bulletin 25: 165-171.
Mason P.G., Broadbent A.B., Whistlecraft J.W. and Gillespie D.R. (2011). Interpreting the host range of Peristenus digoneutis and Peristenus relictus (Hymenoptera: Braconidae) biological control agents of Lygus spp. (Hemiptera: Miridae) in North America. Biological Control 57: 94-102.
Mathenge, C.W., Holford, P., Hoffmann, J.H., Zimmermann, H.G., Spooner-Hart, R., Beattie, G.A.C. (2010). Determination of biotypes of Dactylopius tomentosus (Hemiptera: Dactylopiidae) and insights into the taxonomic relationships of their hosts, Cylindropuntia spp. Bulletin of Entomological Research 100: 3, 347-358
McClay A.S. and Balciunas J.K. (2005). The role of pre-release efficacy assessment in selecting classical biological control agents for weeds - applying the Anna Karenina principle. Biological Control 35: 197-207
McCoy E.D. and Frank J.H. (2010).
How should the risk associated with the introduction of biological control agents be estimated?
Agricultural and Forest Entomology 12: (1) 1-8.
Florida has a large burden of invasive species, and pre-release testing for nontarget effects has historically beeen inadequate. The authors suggest some ways in which balancing the risks and associated costs of releasing a biological control agent against the risks and associated costs of not releasing the agent may be improved. The precautionary principle applied to biological control falls short as a guide because it does not provide a prescription for action. Florida case histories illustrate the complexity and urgency related to developing such a prescription.
McEvoy P.B. (1996).
Host specificity and biological pest control.
BioScience 46: 401-405.
This review of host specificity and biological pest control is set out under the following headings: biological control organisms used to control weeds; biological control organisms used to control arthropods; and other aspects of biological control.
McEvoy P.B. and Coombs E.M. (1999). Why things bite back: unintended consequences of biological weed control. Pp. 167-194 In: Nontarget effects of biological control introductions, P.A. Follett and J.J. Duan (Ed.) Kluwer Academic Publishers, Norwell, Massachusetts, USA.
McEvoy P.B. and Coombs E.M. (1999). Biological control of plant invaders: regional patterns, field experiment and structured population models. Ecological Applications 9: 387-401
McEvoy P.B., Karacetin E. and Bruck D.J. (2008).
Can a pathogen provide insurance against host shifts by a biological control organism?
Pp. 37-42 In: Proceedings of the XII International Symposium on Biological Control of Weeds, (Ed.) La Grande Motte, France, 22-27 April, 2007.
The cinnabar moth, Tyria jacobaeae (L.) (Lepidoptera: Arctiidae is less effective than alternatives (such as the ragwort flea beetle Longitarsus jacobaeae (Waterhouse) Coleoptera: Chrysomelidae) for controlling ragwort, Senecio jacobaea L. (Asteraceae), and it attacks non-target plant species. It also carries a disease (a host-specific microsporidian Nosema tyriae). We used a life table response experiment to estimate the independent and interacting effects of Old World and New World host plant species (first trophic level) and the entomopathogen (third trophic level) on the life cycle and population growth of the cinnabar moth (second trophic level). We found the population growth rate of the cinnabar moth is sharply reduced on novel compared with conventional host plants by interacting effects of disease and malnutrition. Paradoxically, a pathogen of the cinnabar moth may enhance weed biological control by providing insurance against host shifts.
McFadyen R.E. (2004). Biological control: managing risks or strangling progress? Pp. 78-81 In: 14th Australian Weeds Conference. Weed management: balancing people, planet, profit, B. M. Sindel and S. B. Johnson (Ed.) Wagga Wagga, New South Wales, Australia, 6-9 September 2004
McFadyen R.E.C. (1998). Biological control of weeds. Annual Review of Entomology 43: 369-393.
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., Ferguson C.M., Bixley A.S. and Barratt B.I.P. (2009). Development of Microctonus aethiopoides Loan through multiple generations of novel weevil hosts in the laboratory. Pp. 617-618 in: Proceedings of the 3rd International Symposium on Biological Control of Arthropods, P.G. Mason, D.R. Gillespie and C. Vincent (Ed.) Christchurch, New Zealand.
McNeill M.R., Proffitt J.R., Gerard P.J. and Goldson S.L. (2006). Collections of Microctonus aethipopoides Loan (Hymenoptera: Braconidae) from Ireland. New Zealand Plant Protection 59: 290-296.
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.
McPartland J.M. and Nicholson J. (2003).
Using parasite databases to identify potential nontarget hosts of biological control organisms.
New Zealand Journal of Botany 41: 699-706
The authors propose that biocontrol researchers use internet-available databases to identify potential nontarget organisms that share parasites (biotrophic pathogens and pests) with target hosts, and add these organisms to test species lists. Marijuana (Cannabis sativa) has been targeted for biocontrol, and host range studies have focused upon the Moraceae. A list of Cannabis parasites was compared with database lists of pests and pathogens for hosts in the order Urticales. The databases revealed seven Cannabis biotrophic parasites that were shared by hosts in the family Urticaceae, one biotroph shared by a host in the Celtidaceae, and no biotrophs shared by hosts in the Moraceae, Cercropiaceae, or Ulmaceae. These results suggest that biocontrol host range studies of Cannabis parasites should focus on the Urticaceae and Celtidaceae as well as the Moraceae and that taxonomic relationships within the Uricales be reassessed.
Memmott J. (1999). Food webs as a tool for studying nontarget effects in biological control. Pp. 147-163 In: Nontarget effects of biological control introductions, P.A. Follett and J.J. Duan (Ed.) Kluwer Academic Publishers, Norwell, Massachusetts, USA.
Memmott J., Fowler S.V., Hill R.L. (1998). The effect of release size on the probability of establishment of biological control agents: Gorse thrips (Sericothrips staphylinus) released against gorse (Ulex europaeus) in New Zealand. Biocontrol Science and Technology 8: 103-115.
Memmott J., Martinez N.D. and Cohen J.E. (2000).
Predators, parasitoids and pathogens: species richness, trophic generality and body sizes in a natural food web.
Journal of Animal Ecology 69: 1-15.
A food web is presented which describes trophic interactions among the herbivores, parasitoids, predators and pathogens associated with broom, Cytisus scoparius (L.) Link. The web comprises a total of 154 taxa: one plant, 19 herbivores, 66 parasitoids, 60 predators, five omnivores and three pathogens. There are 370 trophic links between these taxa in the web. The taxa form 82 functionally distinct groups, called trophic species. Predators consumed more species than did parasitoids; externally feeding herbivores were most vulnerable and the concealed herbivores were least vulnerable. Miners were vulnerable to the most parasitoid species and externally feeding herbivores were the most vulnerable to predators. Relative sizes of predators and parasitoids are discussed.
Messing R.H. (1992). Biological control in island ecosystems: cornerstone of sustainable agriculture or threat to biological diversity. Pacific Science 46: 387-388.
Messing R.H. (2005).
Hawaii as a role model for comprehensive U.S. Biocontrol legislation: the best and the worst of it.
Pp. 686-691 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 USA currently has no comprehensive, integrated legislative or regulatory framework to manage the permitting of imported biological control agents. In contrast, the State of Hawaii has specific, detailed and exhaustive rules for obtaining import and release permits for natural enemies and this could serve as a useful model for national protocols, with coordinated scientific evaluation at several levels of specialization and input from a wide range of concerned parties. The author suggests that the best parts of the Hawaii system are captured, and the legalistic and bureaucratic aspects removed, then a thorough, streamlined, efficient, transparent, accountable and enabling regulatory framework could be put in place that would safeguard non-target species while facilitating biological control and environmentally sound pest management at the national level.
Messing R.H. and Xin-Geng W. (2008).
Competitor-free space mediates non-target impact of an introduced biological control agent.
Ecological Entomology 34: 107-113
The authors show that competitor-free space is a key mechanism maintaining an apparent host shift by an introduced biocontrol agent onto a non-target species.
Messing R.H., Roitberg B.D. and Brodeur J. (2006). Measuring and predicting indirect impacts of biological control, competition, displacement and secondary interactions. Pp. 64-77 In: Environmental impact of invertebrates for biological control of arthropods - methods and risk assessment, F. Bigler, D. Babendreier and U. Kuhlmann (Ed.) CABI Publishing, Wallingford, UK
Miller J.R. and Strickler K.L. (1984). Finding and accepting host plants. Pp. 127-157 In: Chemical Ecology of Insects, W.J. Bell and R.T. Cardé (Ed.) Chapman & Hall, London.
Miller M.L. and Aplet G.H. (2005).
Applying legal sunshine to the hidden regulation of biological control.
Biological Control 35: 358-365.
This article identifies a legal gap in current USDA policy concerning decisions about the review and release of biological pest control agents. Current practices do not provide sufficient information for biologists or an informed public to understand or evaluate policy decisions and environmental outcomes. The USDA needs to comply with federal law by making all relevant documents and data available on the internet. Federal law and policy requires that the USDA release all relevant information, and make it readily accessible to all interested parties.
Mills N.J. (2006). Accounting for differential success in the biological control of homopteran and lepidopteran pests. New Zealand Journal of Ecology 30: 61-72.
Mills N.J. and Getz W.M. (1996). Modelling the biological control of insect pests: a review of host-parasitoid models. Ecological Modelling 92: 121-143
Mills N.J. and Gutierrez A.P. (1999). Biological contro of insects: a tritrophic perspsective. Pp. 89-102 In: Theoretical approaches to biological control, B. A. Hawkins and H. V. Cornell (Ed.) Cambridge University Press, Cambridge, UK
Mills N.J. and Kean J.M. (2010). Modelling, behavioural studies and molecular approaches: Methodological contributions to biological control success. Biological Control (in press).
Moeed A., Hickson R. and Barratt B.I.P. (2006). Principles of environmental risk assessment of invertebrates in biological control of arthropods. Pp. 241-253 In: Environmental impact of arthropod biological control: methods and risk assessment., U. Kuhlmann, F. Bigler and D. Babendreier (Ed.) CABI Bioscience, Delemont, Switzerland.
Morehead S.A. and Feener D.H. (2000). An experimental test of potential host range in the ant parasitoid Apocephalus paraponerae. Ecological Entomology 25: 332–340.
Morin L. and Edwards P.B. (2006). Selection of biological control agents for bridal creeper: a retrospective review. Australian Journal of Entomology 45: 287-291
Morin L., Evans K.J. and Sheppard A.W. (2006). Selection of pathogen agents in weed biological control: critical issues and peculiarities in relation to arthropod agents. Australian Journal of Entomology 45: 349-365
Morin L., Hill R.L. and Matayoshi S. (1997). Hawaii's successful biological control strategy for mist flower (Ageratina riparia) - can it be transferred to New Zealand? Biocontrol News and Information 18: 77-88
Moriya S., Inoue K., Shiga M. and Mabuchi M. (1992).
Interspecific relationship between and introduced parasitoid, Torymus sinensis Kamijo, as a biological control agent of the chestnut gall wasp, Dryocosomus kuriphylus Yasumatsu, and an endemic parasitoid, T. beneficus Yasumatsu et Kamijo.
Acta Phytopathologia et Entomologica Hungarica 27: 479-483.
The interspecific relationship of a parasitoid of Dryocosmus kuriphilus that was endemic to Japan (Torymus beneficus) and an introduced control agent (T. sinensis) was investigated. The ovipositor sheath is longer in T. sinensis than in T. beneficus and 1990 they made up about 90% of the emerging parasitoids. In crossing experiments, females with ovipositor sheaths of intermediate length were observed and F1 females were fertile when backcrossed to males of the parent species. Females with intermediate ovipositors were observed in the field from 1984.
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.
Morrison-Lloyd W. (2006). Post-release host-specificity testing of Pseudacteon tricuspis, a phorid parasitoid of Solenopsis invicta fire ants. BioControl 51: 195-205.
Muller-Scharer, H. and Schaffner, U. (2008).
Classical biological control: exploiting enemy escape to manage plant invasions.
Biological Invasions 10: 859-874
Key issues considered to be important for safe, efficient, and successful classical biological control project are discussed. These include selection of effective control agents, host specificity of the biological control agents, implications of the genetic population structure of the target populations, and potential impact on native food webs. It is recommended that pre-release impact assessment should focus on how to reach high densities of the control agents, and aspects of tolerance to, and compensation of herbivory; more effort should be made to integrate and combine biological control with existing or potential management options.
Munro V.M.W. and Henderson I.F. (2002).
Nontarget effect of entomophagous biocontrol: shared parasitism between native lepidopteran parasitoids and the biocontrol agent Trigonospila brevifacies (Diptera: Tachinidae) in forest habitats.
Environmental Entomology 32: 388-396.
The parasitoid guild attacking Tortricidae on broadleaf/podocarp forests was studied at six sites in the central North Island. Host parasitoid interactions at a community level between native parasitoids and the introduced species Trigonospila brevifacies (Hardy) were investigated. T. brevifacies was numerically dominant in the tortricid parasitoid guild and it parasitized more species of Tortricidae than other parasitoids at the North Island forest sites surveyed. Only the introduced Australian canefruit pest Eutorna phaulocosma Meyrick (Lepidoptera: Oecophoridae) received a higher proportion of parasitism from T. brevifacies than any of the native Lepidoptera. All native parasitoid species were less abundant than T. brevifacies.
Murdoch W.W., Briggs C.J. and Nisbet R.M. (1996).
Competitive displacement and biological control in parasitoids: A model.
American Naturalist 148: 807-826.
California red scale is controlled in many areas by parasitoids of the genus Aphytis. In southern California, Aphytis lingnanensis provided inadequate control of red scale in inland valleys. Aphytis melinus was introduced, competitively displaced A. lingnanensis within a few red scale generations, and caused satisfactory biological control. Aphytis melinus is successful on smaller- sized scale than A. lingnanensis, and a stage-structured parasitoid-host model is presented in which the two parasitoid species are the same except for A. melinus's size-dependent advantage in sex allocation. This model can account for the competitive displacement of A. lingnanensis and the improvement in biological control. Aphytis melinus can also produce an additional female egg from larger red scale, and this increases its advantage in competition. The effects of other model parameters were discussed along with implications for a predictive theory of biological control.
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.
Nafus D.M. (1993). Movement of introduced biological control agents onto nontarget butterflies, Hypolimnas spp. (Lepidoptera: Nymphalidae). Environmental Entomology 22: 265-272.
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.
Nechols J.R., Kauffman W.C. and Schaefer P.W. (1992). Significance of host specificity in classical biological control. Pp. 41-52 In: Selection Criteria and Ecological Consequences of Importing Natural Enemies, W.C. Kaufmann and J.E. Nechols (Ed.) Entomological Society of America, Lanham, Maryland, USA.
Nowell D. and Maynard G.V. (2005). International guidelines for the export, shipment, import and release of biological control agents and other beneficial organisms (ISPM No. 3). Pp. 726-734 In: Second International Symposium on Biological Control of Arthropods, M. Hoddle (Ed.) Davos, Switzerland, USDA Forest Service.
O'Callaghan M. and Brownbridge M. (2009).
Environmental impacts of microbial control agents used for control of invasive pests.
In: Use of Microbes for Control and Eradication of Invasive Arthropods, Hajek, AE., Glare, TR., O'Callaghan, M. (Eds.) Progress in biological control Vol. 6: 305-327
Insect pathogens vary in key characteristics which determine their safety profile with respect to impacts on non-target species. Laboratory testing against beneficial species and post-application monitoring of impacts suggest that effects on non-target organisms, in comparison with other control methods are environmentally benign. Biopesticides can be attractive control options in many situations, and their use is likely to have minimal impact on beneficial and other non-target species. For example Bacillus thuringiensis, has not precipitated any major ecological disturbances, even when used in very intensive and prolonged eradication programmes.
O'Hanlon P.C., Briese D.T. and Peakall R. (2000).
Know your enemy: the use of molecular ecology in the Onopordum biological control project.
Proceedings of the X International Symposium on Biological Control of Weeds: 281-288
Accurate identification of the target weed(s) for a biological control project is critical to the success of a biological control project, particularly where the weed may comprise different biotypes or be part of a species complex. Molecular ecology provides tools for resolving the identity of weeds. An example is given with a hybrid swarm of Onopordum spp. in Australia. Molecular markers can be used to better understand the phylogeny of plant groups containing the target weed(s).
OECD E.D. (2003). Guidance for information requirements for regulation of invertebrates as biological control agents (IBCAs). Organisation for Economic Co-operation and Development. 19 pp.
Obrycki J.J. (1989).
Parasitization of native and exotic coccinellids by Dinocampus coccinellae (Schrank) (Hymenoptera: Braconidae).
Journal of the Kansas Entomological Society 62: 211-218.
The suitability of 3 introduced and 3 native (to the USA) coccinellids as hosts for the braconid Dinocampus coccinellae was examined in the laboratory at 22°C and LD 16:8. Adults of the Nearctic species (Coleomegilla maculata, Cycloneda munda and Hippodamia convergens) and one Palaearctic species, Coccinella septempunctata, were suitable hosts for D. coccinellae. The mean development time for the parasitoid ranged from 30 to 33.3 days in these hosts, while successful parasitism varied between 30 and 57%. Only 1.5% of D. coccinellae emerged from Propylea quattuordecimpunctata, development times being significantly longer (37 days) with this host. The parasitoid failed to develop in H. variegata.
Obrycki J.J., Elliott N.C. and Giles L.G. (1999). Coccinellid introductions: potential for and evaluation of nontarget effects. Pp. 127-145 In: Nontarget effects of biological control introductions, P.A. Follett and J.J. Duan (Ed.) Kluwer Academic Publishers, Norwell, Massachusetts, USA.
Olckers T. and Borea C. (2009).
Assessing the risks of releasing a sap-sucking lace bug, Gargaphia decoris , against the invasive tree Solanum mauritianum in New Zealand.
BioControl 54: 143-154
The South American tree Solanum mauritianum Scopoli (Solanaceae), a major environmental weed in South Africa and New Zealand, has been targeted for biological control, with releases of agents restricted to South Africa. The leaf-sucking lace bug, Gargaphia decoris Drake (Tingidae), has become established in South Africa with reports of severe damage. Host-specificity testing was carried out in South Africa in laboratory and open-field trials, with cultivated and native species of Solanum from New Zealand showed that none of the three native New Zealand Solanum species are acceptable as hosts. Some cultivars of S. melongena L. (eggplant) supported feeding, development and oviposition in the no-choice tests. Field trials and risk assessment indicate that the insect has a host range restricted to S. mauritianum. An application for permission to release G. decoris in New Zealand will be submitted to the regulatory authority.
Onstad D.W. and McManus M.L. (1996).
Risks of host range expansion by parasites of insects.
BioScience 46: 430-435.
The authors discuss the estimation of risks that biological control agents pose to nontarget species. There has been no demonstration that biological control agents have threatened, endangered or extirpated any native insect or arachnid species in the USA.
Orlinskii A.D. (1997). Precautions for and experiences with introduction of exotic biological control agents into the former USSR. Pp. 61-68 In: EPPO/CABI workshop on safety and efficacy of biological control in Europe, I.M. Smith (Ed.) Blackwell Science Ltd., Oxford.
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.
Palmer W.A. (2004).
Risk analyses of recent cases of non-target attack by potential biocontrol agents in Queensland.
Proceedings of the XI International Symposium on Biological Control of Weeds: 305-309
Examples are given of three insects considered for biocontrol of weeds in Queensland have potential or realized non-target risks associated with them. These examples are discussed within the general framework of risk analysis for introduced biocontrol agents in Australia.
Palmer W.A., Day M.D., Kunjithapatham D., Snow E.L. and Mackey A.P. (2004). Analysis of the non-target attack by the lantana sap-sucking bug, Aconophora compressa , and its implications for biological control in Australia. Pp. 341-344 In: 14th Australian Weeds Conference. Weed management: balancing people, planet, profit, B. M. Sindel and S. B. Johnson (Eds.) Wagga Wagga, New South Wales, Australia, 6-9 September 2004
Paraiso O., Kairo M.T.K., Hight S.D., Leppla N.C., Cuda J.P., Owens M. and Olexa M.T . (2013).
Opportunities for improving risk communication during the permitting process for entomophagous biological control agents: a review of current systems.
Biocontrol 58: 1-15.
Concerns about potentially irreversible non-target impacts from the importation and release of entomophagous biological control agents (BCAs) have resulted in increasingly stringent national import requirements by National Plant Protection Organizations worldwide. However, there is a divergence of opinions among regulators, researchers, environmentalists, and the general public on ways to appropriately manage associated risks. Implementation of a comprehensive and effective risk communication process might narrow the opinion gaps. Results from a comprehensive survey cnducted in the United States were used to describe communication habits of stakeholders involved in biological control and identify areas that are fundamental in an efficient process. In addition, this study critically reviews risk communication practices and how phytosanitary decisions are communicated in the permitting systems for entomophagous BCAs of several countries to identify risk communication tools used in an effective risk communication framework. The following barriers to efficient risk communication were identified: absence of a formalized risk communication process, undefined risk communication goals and target audiences, lack of credibility and objectivity of information sources, inefficiency of mode of distribution of messages, insufficient public participation, and lack of transparency of decision making processes. This paper suggests the creation and/or enhancement of modes of distribution of risk messages to increase coverage, understanding, and guidance. For instance, messages should be presented in different formats such as internet, brochures, and newspapers. Surveys, public meetings, and trainings/workshops are tools that can be used to characterize stakeholders’ diversity and develop risk messages specific to the targeted audience. Implementation of a participatory decision making process will increase stakeholder involvement and trust in the risk management plan. Development of practical mechanisms, such as public hearings will increase all stakeholders’ involvement in the risk assessment process. A clear framework describing how public comments will be incorporated in the decision making process should be implemented. Finally, to ensure a streamlined risk communication process, there must be consistency in the messages disseminated by federal, state, and local agencies.
Parker I.M., Simberloff D., Lonsdale W.M., Goodell K., Wonham M., Kareiva P.M., Williamson M.H., Von Holle B., Moyle P.B., Byers J.E. and Goldwasser L. (1999). Impact: toward a framework for understanding the ecological effects of invaders. Biological Invasions 1: 3-19.
Parry D. (2009).
Beyond Pandora's Box: quantitatively evaluating non-target effects of parasitoids in classical biological control.
Biological Invasions 11: 47-58
Howarth's paper (Proc Hawaii Entomol Soc 24:239-244, 1983) "Classical biological control: Panacea or Pandora's Box" raised awareness of the relative safety of introductions for classical biological control. Here, the author examines the potential for non-target effects among insect parasitoids. In response to the need for quantitative studies, three different techniques, quantitative food webs, life table analysis, and experimental populations are explored along with three approaches to ascertaining the strength of competitive interactions between native and introduced parasitoids.
Paton (1992). Legislation and its administration in the approval of agents for biological control in Australia. In: Proceedings of the VIII International Symposium on Biological Control of Weeds, E.S. Delfosse and R.R. Scott (Ed.)
Paynter Q., Martin N., Berry J., Hona S., Peterson P., Gourlay A.H., Wilson-Davey J., Smith L., Winks C. and Fowler S.V. (2008).
Non-target impacts of Phytomyza vitalbae a biological control agent of the European weed Clematis vitalba in New Zealand.
Biological Control 44: 248-258
The agromyzid leaf-mining fly Phytomyza vitalbae, which was introduced into New Zealand as a biological control agent of the invasive vine Clematis vitalba L. (old man's beard; Ranunculaceae) has been recorded attacking two native Clematis species in New Zealand, particularly C. foetida but at lower incidence and levels of attack than the target. No-choice starvation tests indicated that non-target attack was a "spillover" effect that is unlikely to have a major detrimental impact on the non-target plants. Our results show that the prevalence of spillover onto non-target species was underestimated in pre-release testing and we discuss how host-range testing might be improved in the light of these findings.
Paynter Q., Waipara N.W., Peterson P.G., Hona S.R., Fowler S.V., Gianotti A.F. and Wilkie P. (2006). The impact of two introduced biocontrol agents, Phytomyza vitalbae and Phoma clematidina, on Clematis vitalba in New Zealand. Biological Control 36: 350-357
Paynter Q.E., Fowler S.V., Gourlay A.H., Haines M.L., Harman H.M., Hona S.R., Peterson P.G., Smith L.A., Wilson-Davey, J.R.A., Winks, C.J. and Withers T.M. (2004). Safety in New Zealand Weed Biocontrol: A nationwide survey for impacts on non-target plants. New Zealand Plant Protection 57: 102-107
Pearson D.E. and Callaway R.M. (2005).
Indirect nontarget effects of host-specific biological control agents: Implications for biological control.
Biological Control 35: 288-298.
Recent case studies of indirect nontarget effects of biological control agents were evaluated in the context of theoretical work in community ecology. Although difficult to predict, all indirect nontarget effects of host specific biological control agents derived from the nature and strength of the interaction between the biological control agent and the pest. It was concluded that safeguarding against indirect nontarget effects of host-specific biological control agents depends on host specific and efficacious biological control agents.
Pemberton R.W. (2000). Predictable risk to native plants in weed biological control. Oecologia 125: 489-494.
Pemberton R.W. (2004).
Biological control safety within temporal and cultural contexts.
Proceedings of the XI International Symposium on Biological Control of Weeds: 245-246
An analysis of non-target plant use resulting from natural enemy introductions in continental USA, Hawaii and the Caribbean between 1902 and 1994 was carried out. Fourteen of the 117 agents introduced have adopted 45 native plants as developmental hosts. All but one of these plants are closely related to the target weeds. The non-target use was predictable, based on known host ranges of the insects in their native areas and host-specificity testing. The single case in which a plant unrelated to the target weed was adopted involves the lantana lacebug (Teleonemia scrupulosa introduced to Hawaii in 1902), which was thought to be a lantana specialist but apparently is not. Almost all insects adopting native plants were introduced between 1902 and 1972 when 20% (13/63) of the agents introduced have adopted native plants compared with only 1.8% (1/54) of the agents introduced between 1973 and 2002. None of the 117 introduced natural enemies have adopted agricultural plants.
Pennacchio F. and Strand M.R. (2006). Evolution of developmental strategies in parasitic Hymenoptera. Annual Review of Entomology 51: 233-258
Perdikis, D., Alomar, O. (2011).
Heteropteran predators and their role in biological control in agroecosystems.
Biological Control Special Issue: 59: 1-67
This issue contains 7 papers focusing on the importance of Heteropteran predators as biological control agents. The ecology and risks and benefits of using these predators in insect pest and weed control are also discussed.
Phillips C.B. (1996). Intraspecific variation in Microctonus hyperodae and M. aethiopoides (Hymenoptera: Braconidae); significance for their use as biological control agents. PhD Thesis, Department of Entomology, Lincoln University, Lincoln, New Zealand. 169 pp.
Phillips C.B., Cane R.P., Mee J., Chapman H.M., Hoelmer K.A. and Coutinot D. (2002).
Intraspecific variation in the ability of Microctonus aethiopoides (Hymenoptera: Braconidae) to parasitise Sitona lepidus (Coleoptera: Curculionidae).
New Zealand Journal of Agricultural Research 45: 295-303.
An experiment was conducted to compare the suitability of French and New Zealand Sitona lepidus (Coleoptera: Curculionidae) as hosts for a French biotype of Microctonus aethiopoides (Hymenoptera: Braconidae) Loan . This provided no evidence of S. lepidus intraspecific variation in host suitability for parasitism. However, amplification of inter simple sequence repeat (ISSR) regions of M. aethiopoides DNA demonstrated clear genetic differences between French and New Zealand M. aethiopoides. It was concluded that intraspecific variation in the ability of M. aethiopoides to evade the immune response of S. lepidus is the reason for the low levels of parasitism observed in New Zealand compared with Europe.
Phillips C.B., Iline I.L., Vink C.J., Winder L.M. and McNeill M.R. (2006). Methods to distinguish between the Microctonus aethiopoides strains that parasitise Sitona lepidus and Sitona discoideus. New Zealand Plant Protection 59: 1-6.
Phillips C.B., Proffitt J.R. and Goldson S.L. (1998). Potential to enhance the efficacy of Microctonus hyperodae Loan. Proceedings of the New Zealand Plant Protection Conference 51: 16-22.
Phillips C.B., Vink C.J., Blanchet A. and Hoelmer K.A. (2008).
Hosts are more important than destinations: what genetic variation in Microctonus aethiopoides (Hymenoptera: Braconidae) means for foreign exploration for natural enemies.
Molecular Phylogenetics and Evolution 49: 467-476
Nucleotide sequence data were generated from the gene regions COI, 16S, and arginine kinase to assess genetic variation within the parasitoid, Microctonus aethiopoides, reared from Sitona discoideus, S. hispidulus, and Hypera posticafrom locations in France. The results combined with previously published data from 14 countries show that M. aethiopoides genetic variation is more strongly correlated with host taxon than with sampling location. The results suggested that success rates and environmental safety in biological control would be improved by ensuring that parasitoids collected in the native range are reared from the same host species as the one being targeted for control in the region of introduction.
Pimentel D. (1980).
Environmental risks associated with biological control.
Ecological Bulletin 31: 11-24.
A review of the environmental effects of classical biological control programmes against pests and weeds under the headings: beneficial aspects of biological control; negative impacts of classical biological control; solutions; and future directions. Deals mainly with insects and other arthropods used as biological control agents, but microorganisms, molluscs and vertebrates are also included.
Pimentel D. (1995). Biotechnology: environmental impacts of introducing crops and biocontrol agents in North American agriculture. Pp. 13-29 In: Biological Control: Benefits and Risks, H.M.T. Hokkanen and J.M. Lynch (Ed.) Cambridge University Press, Cambridge, UK.
Pimentel D., Glenister C., Fast S. and Gallahan D. (1984). Environmental risks of biological pest controls. Oikos 42: 283-290.
Polis G.A. and Strong D.R. (1996). Food web complexity and community dynamics. The American Naturalist 147: 813-846.
Porter S.D., Valles S.M., Davis T.S., Briano J.A., Calcaterra L.A., Oi D.H. and Jenkins A. (2007).
Host specificity of the microsporidian pathogen Vairimorpha invictae at five field sites with infected Solenopsis invicta fire ant colonies in northern Argentina.
Florida Entomologist 90: 447-452
The microsporidian pathogen Vairimorpha invictae was evaluated for release in the United States as a biological control agent for imported fire ants. The host range of this pathogen was examined showing that Solenopsis invicta Buren fire ant colonies had high levels of infection (28-83%) at one site, but none were found in ants at other sites. The results of this study indicate that, in its native South American range, V. invictae is specific to Solenopsis fire ants.
Poulton J., Markwick N.P., Ward V.K. and Young V. (2007).
Host range testing of a nucleopolyhedrovirus of the lightbrown apple moth, Epiphyas postvittana.
New Zealand Plant Protection 60: 26-32
Epiphyas postvittana nucleopolyhedrovirus (EppoNPV) has potential as a biopesticide for control of lightbrown apple moth and non-target impacts were investigated. Eight non-target insect species from one hymenopteran and five lepidopteran families were inoculated with EppoNPV. Larval survival, growth rates, pupation and pupal weights were measured and larvae examined for virus. Few viral infections were found, growth and survival were compromised in virus-fed individuals in only one species, Tyria jacobaeae, where the majority of larvae had high microsporidal infections. EppoNPV polyhedra were found in only one larva, suggesting a very low likelihood of field infectivity.
Powell J.A. and Logan J.A. (2005). Insect seasonality: circle map analysis of temperature-driven life cycles. Theoretical Population Biology 67: 161-179.
Pratt P.D., Rayamajhi M.B., Center T.D., Tipping P.W. and Wheeler G.S. (2009). The ecological host range of an intentionally introduced herbivore: A comparison of predicted versus actual host use. Biological Control 49: 146-153.
Price P.W. (2000). Host plant resource quality, insect herbivores and biocontrol. Pp. 583-590 In: Proceedings of the Xth International Symposium on Biological Control of Weeds, N.R. Spencer (Ed.) Montana State University, Bozeman Montana, USA
Purcell M.F., Duan J.J. and Messing R.H. (1997). Response of three hymenopteran parasitoids for fruit fly control to a gall-forming tephritid, Procecidochares alani (Diptera: Tephritidae). Biological Control 9: 193-200.
Raghu S. and Van Klinken R.D. (2006). Refining the ecological basis for agent selection in weed biological control. Australian Journal of Entomology 45: 251-252
Raghu S., Dhileepan K. and Scanlan J.C. (2007).
Predicting risk and benefit a priori in biological control of invasive plant species: A systems modelling approach.
Ecological Modelling 208: 247-262
In this study a simulation model is used to predict the risks and benefits of introducing the chrysomelid beetle Charidotis auroguttata to manage the invasive liana Macfadyena unguis-cati in Australia. Preliminary host-specificity testing of this herbivore indicated that there was limited feeding on a non-target plant, although the non-target was only able to sustain some transitions of the life cycle of the herbivore. The model included herbivore, target and non-target life history and incorporates spillover from the target to the non-target. The model predicted that risk to the non-target became unacceptable when the ratio of target to non-target in a given patch ranged from 1:1 to 3:2. By considering risk and benefit simultaneously, we highlight how such a simulation modelling approach can assist in making more objective decisions on the value of releasing specialist herbivores as biological control agents.
Raghu S., Wilson J.R. and Dhileepan K. (2006). Refining the process of agent selection through understanding plant demography and plant response to herbivory. Australian Journal of Entomology 45: 308-316
Rand T.A. and Louda S.M. (2006).
Invasive insect abundance varies across the biogeographic distribution of a native host plant.
Ecological Applications 16: 877-890
The authors quantified the abundance of the introduced and now invasive biocontrol weevil, Rhinocyllus conicus, on a newly adopted native host plant, Cirsium canescens (Platte thistle), across the plant's distributional range using regression and structural equation analyses. R. conicus now occurs throughout the-majority of the range of C. canescens, and were highest in the center of the native plant's distribution where its coevolved, targeted weed host (Carduus nutans, musk thistle) is absent. In addition to biogeographic position, the only other consistent predictor of weevil densities across sites was the number of flower heads per C. canescens plant. The results were consistent with the hypothesis that exotic weevil abundance on C. canescens is related to the local availability of native floral resources. Results suggest that isolated, peripheral populations of C. canescens are likely to be critical for persistence of Platte thistle.
Rees M. and Hill R.L. (2001). Large-scale disturbances biological control and the dynamics of gorse populations. Journal of Applied Ecology 38: 364-377
Rees M. and Paynter Q. (1997). Biological control of Scotch broom: Modelling the determinants of abundance and the potential impact of introduced insect herbivores. Journal of Applied Ecology 34: 1203-1221
Retief E., Rooi C. van, and Breeyen A. den (2016). Environmental requirements and host-specificity of Puccinia eupatorii, a potential biocontrol agent of Campuloclinium macrocephalum, in South Africa. Australasian Plant Pathology 45: 135-144
Roberts L.I.N. (1986). The practice of biological control - implications for conservation, science and the community. The Weta, Entomological Society of NZ 9: 76-84.
Romeis J., Babendreier D., Wackers F.L. and Shanower T.G. (2005). Habitat and plant specificity of Trichogramma egg parasitoids - underlying mechanisms and implications. Basic and Applied Ecology 6: 215-236
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.
Rutledge C.E. and Wiedenmann R.N. (1999). Habitat preferences of three congeneric braconid parasitoids: implications for host-range testing in biological control. Biological Control 16: 144-154
Samways M.J. (1994). Insect Conservation Biology. Chapman & Hall, London. 358 pp.
Samways M.J. (1997). Classical biological control and biodiversity conservation: what risks are we prepared to accept? Biodiversity and Conservation 6: 1309-1316.
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.
Sands D.P.A. and Van Driesche R.G. (2000). Evaluating the host range of agents for biological control of arthropods: rationale, methodology and interpretation. Pp. 69-83 In: Host-specificity testing of exotic arthropod biological control agents: the biological basis for improvement in safety, R.G. Van Driesche, T. Heard, A.S. McClay and R. Reardon (Ed.) USDA Forest Service Bulletin, Morgantown, West Virginia, USA.
Sands D.P.A. and Van Driesche R.G. (2004). Using the scientific literature to estimate the host range of a biological control agent. Pp. 15-23 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.
Scherwinski K., Grosch R. and Berg G. (2007).
Root application of bacterial antagonists to field-grown lettuce: effects on disease suppression and non-target microorganisms.
Bulletin OILB/SROP 30: 257
This paper discusses biological control of the phytopathogenic fungus Rhizoctonia solani which reduced yield in agricultural and horticultural crops using naturally antagonistic bacteria. The two rhizobacteria Pseudomonas trivialis (3Re2-7) and Pseudomonas fluorescens (L13-6-12) as well as the potato endophyte Serratia plymuthica (3Re4-18) were selected as effective Rhizoctonia antagonists. The impact of these bacteria on non-target lettuce-associated microorganisms was assessed after their root application in two field trials in Germany and all resulted in a significant increase of the dry weight and a significant decrease of the disease severity. The microbial communities of the rhizosphere, the endorhiza and the endophyllosphere of field-grown lettuce were examined and only transient changes in the composition of the bacterial communities were found. The authors concluded that an environmentally friendly and efficient biocontrol strategy can be developed.
Schulten G.G.M. (1997). The FAO Code of Conduct for the import and release of exotic biological. Pp. 29-36 In: EPPO/CABI workshop on safety and efficacy of biological control in Europe, I.M. Smith (Ed.) Blackwell Science Ltd., Oxford.
Secord D. and Kareiva P. (1996).
Perils and pitfalls in the host specificity paradigm.
BioScience 46: 448-453.
The importance of host specificity in biological control agents is discussed in relation to unexpected host shifts; specificity assessment, evolution and indirect effects; and host specificity and cost/benefit analysis for biological control.
Seymour, C.L. and Veldtman, R. (2010).
Ecological role of control agent, and not just host-specificity, determine risks of biological control.
Austral Ecology 35: 704-711
Gall-formers are popular as biological control agents because they are host-specific and therefore considered low risk. However, galls can also be considered to be ecological engineers, because they provide nutritional resources for native invertebrates. The Authors tested whether native invertebrates had formed associations with the gall-forming fungus Uromycladium tepperianum, introduced into South Africa to control the Australian invasive alien tree Acacia saligna, by collecting U. tepperianum galls and monitoring emergence. A number of invertebrates had formed associations with the biological control agent, including the citrus pest, Thaumatotibia leucotreta (false codling moth). The study illustrated a case of a host-specific classical biological control agent providing resources for an economically significant crop pest. The authors concuded that although biological control agents are strictly vetted to ensure host-specificity, those that become abundant and can act as ecological engineers pose risks when native biota form associations with them, resulting in a number of possible cascading ecosystem effects. There could also be economic consequences when these associated species include agricultural pests. Potential ecological effects of biological control agents, should be considered in their selection.
Shea K., Kelly D., Sheppard A.W. and Woodburn T.L. (2005).
Context-dependent biological control of an invasive thistle.
Ecology 86: 3174-3181.
Success of biological control of Carduus nutans using insects that attack rosettes or developing seed heads, has varied in different parts of its invaded range. Here a demographic matrix models is used to compare populations in Australia and New Zealand, to explain differences. In a New Zealand population, rapid population growth of C. nutans is driven by early life history transitions. In an Australian population, fecundity of C. nutans is of reduced importance, and survivorship of rosettes plays an increased role. These differences suggest how biocontrol agents that are successful at providing control in one situation but may fail in another.
Shelby K.S. and Webb B.A. (1999). Polydnavirus-mediated suppression of insect immunity. Journal of Insect Physiology 45: 507-514
Sheppard A., Haines M.L. and Thomann T. (2006). Native-range research assists risk analysis for non-targets in weed biological control: the cautionary tale of the broom seed beetle. Australian Journal of Entomology 45: 292-297
Sheppard A.W. (1992). Predicting biological weed control. Trends in Ecology & Evolution 7: 290-291.
Sheppard A.W. (1999). Which test? A mini review of test usage in host specificity testing. Pp. 60-69 In: Host specificity testing in Australasia: towards improved assays for biological control, T.M. Withers, L. Barton-Browne and J. Stanley (Ed.) Scientific Publishing, Department of Natural Resources, Brisbane.
Sheppard A.W. (2003). Prioritising agents based on predicted efficacy: beyond the lottery approach. Pp. 11-21 In: Improving the selection, testing and evaluation of weed biocontrol agents, H. Spafford-Jacobs and D. T. Briese (Ed.) Cooperative Research Centre for Australian Weed Management, Adelaide, Australia
Sheppard A.W., Hill R.L., DeClerck-Floate R.A., McClay A., Olckers T., Quimby P.C. and Zimmermann H.G. (2003). A global review of risk-benefit-cost analysis for the introduction of classical weed biological control agents against weeds: a crisis in the making? Biocontrol News and Information 24: 91N-108N.
Sheppard A.W., Shaw R.H. and Sforza R. (2006).
Top 20 environmental weeds for classical biological control in Europe: a review of opportunities, regulations and other barriers to adoption.
Weed Research 46: 93-117
Despite more than 130 case histories in Europe against insect pests, no exotic classical biological control agent has been released in the EU against an alien invasive weed despite increasing numbers of exotic invasive plants being imported. The authors review possible European weed targets for classical biological control from ecological and socioeconomic perspectives. They also consider why classical biological control of European exotic plants remains untested, and the regulatory framework that surrounds such biological control activities within constituent countries of the EU and suggest how invasive exotic weeds in might be managed in the future in Europe.
Sheppard A.W., Van Klinken R.D. and Heard T.A. (2005).
Scientific advances in the analysis of direct risks of weed biological control agents to nontarget plants.
Biological Control 35: 215-226.
The strengths, weaknesses, and best practice for the different host specificity test types are now understood. Understanding the concept of fundamental host range and using this to maximize reliability in predicting field host specificity following release are still inconsistently understood or adopted. This needs to be consistently applied so the process of testing can follow a recognized process of risk analysis from hazard identification to uncertainty analysis based on the magnitude and likelihood of threats to non-targets. Modern molecular techniques are answering questions associated with subspecific variation in biological control agents with respect to host use and the chance of host shifts of agents following release. This review covers all these recent advances for the first time in one document, highlighting how inconsistent interpretation by biological control practitioners can be avoided.
Shiga M. (1999).
Classical biological control of the chestnut gall wasp, Dryocosmus kuriphilus: present status and interactions between an introduced parasitoid, Torymus sinensis, and native parasitoids.
Pp. 175-188 In: Biological invasions of ecosystem by pests and beneficial organisms, E. Yano, K. Matsuo, M. Shiyomi and D.A. Andow (Ed.) National Institute of Agro-Environmental Sciences, Tsukuba, Japan.
This research demonstrates a case of displacement of a native by and introduced parasitoid.
Simberloff D. (1991). Keystone species and community effects of biological control introductions. Pp. 1-19 In: Assessing ecological risks of biotechnology, L.R. Ginzburg (Ed.) Butterworth-Heinemann, Boston, Massachusetts, USA.
Simberloff D. (1992).
Conservation of pristine habitats and unintended effects of biological control.
Pp. 103-117 In: Selection Criteria and Ecological Consequences of Importing Natural Enemies, W.C. Kaufmann and J.E. Nechols (Ed.) Entomological Society of America, Lanham, Maryland, USA.
The effect of deliberate introductions of arthropods for biological control purposes on species in pristine habitats is examined. This paper was developed from a symposium held during the 1990 Annual Meeting of the Entomological Society of America in New Orleans, Louisiana.
Simberloff D. (2012). Risks of biological control for conservation purposes. Biocontrol 57: 263-276.
Simberloff D. and Stiling P. (1996).
How risky is biological control?
Ecology 77: 1965-1974.
Literature review of non-target effects of introduced biological control agents. There are few documented instances of damage to non-target organisms or the environment, however, this is not evidence that biological control is safe, because monitoring of non-target species is minimal. Predicting indirect effects is difficult, plus the fact that introduced species can disperse and evolve. Current regulation of introduced biological-control agents is considered insufficient. It is recommended that the likely impact of both the pest and its natural enemy on natural ecosystems and their species should be considered, not only on potential economic benefits.
Simberloff D. and Stiling P. (1996).
Risks of species introduced for biological control.
Biological Conservation 78: 185-192.
Biological control introductions have adversely affected nontarget native species, including some recent cases, and little monitoring of impacts is done, so known problems may be the tip of an iceberg. Regulations for officially releases for biological control are insufficient, and there are also freelance unregulated releases undertaken by private citizens. Cost-benefit analyses for conservation issues are difficult because it is hard to assign values to the loss of species or ecosystem functions. Cost-benefit analyses and risk assessments for biological control introductions would enforce consideration of many factors that now often receive cursory attention, and broaden public understanding of the issues.
Simberloff D. and Stiling P. (1998). How risky is biological control? Reply. Ecology 79: 1834-1836.
Sime K.R., Daane K.M., Wang X.G., Johnson M.W. and Messing R.H. (2008).
Evaluation of Fopius arisanus as a biological control agent for the olive fruit fly in California.
Agricultural and Forest Entomology 10, 423-431.
The egg-prepupal parasitoid Fopius arisanus (Hymenoptera: Braconidae) was evaluated in quarantine facilities as a potential biological control agent for the olive fruit fly Bactrocera oleae (Diptera: Tephritidae) in California, U.S.A. F. arisanus will not attack Tephritidae that feed in inflorescences or galls but may pose risks to native Tephritidae that feed in fruit. The broad host-range of F. arisanus with respect to fruit-feeding Tephritidae may preclude its introduction to California, as may its low fecundity and its intrinsic competitive superiority over larva l-pupal parasitoids, which include specialists on B. oleae that are currently being introduced to California. High rates of direct mortality, however, point to potential uses in augmentative biological control.
Simmonds F.J. and Bennett F.D. (1977). Biological control of agricultural pests. Pp. 464-472 In: Proc. 15th International Congress of Entomology.
Simmonds F.J., Franz J.M. and Sailer R.I. (1976). History of biological control. Pp. 788 In: Practice of biological control, C.B. Huffaker and P.S. Messenger (Ed.) Academic Press, New York.
Singer M.C. (1982). Quantification of host preference by manipulation of oviposition behaviour in the butterfly Euphydryas editha. Oecologia 52: 224-229.
Singer M.C. (1986). The definition and measurement of oviposition preference in plant-feeding insects. Pp. 65-94 In: Insect-Plant Interactions, J.R. Miller and T.A. Miller (Ed.) Springer-Verlag, New York.
Smith I.M. (1997). EPPO/CABI workshop on safety and efficacy of biological control in Europe. Blackwell Science Ltd., Oxford.
Solarz S.L. and Newman R.M. (1996). Oviposition specificity and behavior of the watermilfoil specialist Euhrychiopsis lecontei. Oecologia 106: 337-344.
Solter L.F. and Maddox J.V. (1998). Physiological host specificity of Microsporidia as an indicator of ecological host specificity. Journal of Invertebrate Pathology 71: 206-216.
Spafford-Jacob H. and Briese D.T. (2003). Improving the selection, testing and evaluation of weed biocontrol agents. CRC for Australian Weed Management Technical Series no. 7, Adelaide, Australia.
Spradberry P. (1970). Host finding by Rhyssa persuasoria L. an ichneumon parasite of siricid woodwasps. Animal Behaviour 18: 103-114
Standish R.J., Robertson A.W. and Williams P.A. (2001). The impact of an invasive weed Tradescantia fluminensis on native forest regeneration. Journal of Applied Ecology 38: 1253-1263
Stanley J.N. and Julien M.H. (1998). The need for post-release studies to improve risk assessments and decision making in classical biological control. Pp. 561-564 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.
Stanley M.C. and Fowler S.V. (2004).
Conflicts of interest associated with the biological control of weeds.
Proceedings of the XI International Symposium on Biological Control of Weeds: 322-340
The introduction of weed biological control agents may be delayed or prohibited where the plant targeted for control also has beneficial attributes. Conflicts fall into one of several categories: one or more groups value the target plant for economic and/or cultural use; non-target effects of biological control; those related to biocontrol programs against native plants; and ecological effects of successful biocontrol as a result of weed use by native biota. While industry-based conflicts dominate, there has been a shift towards conflicts associated with the ecological effects of weed biocontrol. The benefits of weeds to ecosystems, particularly where weeds provide resources for native fauna, are becoming an important part of cost-benefit analyses for weed biocontrol programs. Examples where weed biocontrol programs have been delayed because of economic and ecological conflicts are given. At present, weed biocontrol programs are usually initiated only when the risk of conflict is low. Where conflict does occur, communication and cost-benefit analyses are key to ensuring resolution is found. However, cost-benefit analyses, are expensive and time-consuming, causing substantial delays to weed biocontrol programs and ongoing environmental damage as a consequence of weed invasion.
Stiling P. (2004).
Biological control not on target.
Biological Invasions 6: 151-159.
Summary of previously recorded information on the diet breadth of natural enemies released to control insect pests worldwide. Of released biocontrol agents in North America, 48% were recorded as generalists, 29% attacked more than one species in a genus, 23% were specialists on the target pests. So many natural enemies released in biocontrol programs against insect pests have broad diets and non-target effects are likely. In North America many parasitoids attack multiple host genera and species, with an average of 5.8 genera and 7.3 species attacked, indicating broad agreement with data from biological control releases.
Stiling P. and Simberloff D. (1999). The frequency and strength of nontarget effects of invertebrate biological control agents of plant pests and weeds. Pp. 31-43 In: Nontarget effects of biological control introductions, P.A. Follett and J.J. Duan (Ed.) Kluwer Academic Publishers, Norwell, Massachusetts, USA.
Stoltz D.B. and Xu D. (1990). Polymorphism in polydnavirus genomes. Canadian Journal of Microbiology 36: 538-543.
Strand M.R. and Obrycki J.J. (1996).
Host specificity of insect parasitoids and predators.
BioScience 46: 422-429.
The authors discuss the host specificity of insect parasitoids and predators under the headings: categories of predators and parasitoids; use of predatory and parasitic insects in biological control; life histories of predators and parasitoids; host range determinants; and insect natural enemies: risks and benefits to biological control.
Strong D.R. and Pemberton R.W. (2001). Food webs, risks of alien enemies and reform of biological control. Pp. 57-79 In: Evaluating indirect ecological effects of biological control, E. Wajnberg, J.K. Scott and P.C. Quimby (Ed.) CABI Publishing, Wallingford, Oxon., UK.
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.
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Sutherst R.W., Maywald G.F. and Kriticos D.J. (2007). CLIMEX version 3. CD and User's Guide. Hearne Scientific Software, Melbourne, Australia.
Syrett P. (1996).
Insects for biological control of broom (Cytisus scoparius) in New Zealand.
Pp. 525-528 In: Proceedings of the 11th Australian Weeds Conference, Melbourne, Australia, 30 September - 3 October 1996, R. Shepherd (Ed.) Weed Science Society of Victoria Inc., Victoria, Australia.
A program to introduce insects for biological control of broom (Cytisus scoparius) in New Zealand began in 1981 and the progress in this programme is described.
Syrett P. and Emberson R.M. (1997). The natural host range of beetle species feeding on broom Cytisus scoparius (L.) Link (Fabaceae) in south-west Europe. Biocontrol Science and Technology 7: 309-326
Syrett P., Harman H.M. and Fowler S.V. (1995). Identification of risk to kowhai, a New Zealand native plant, Sophora microphylla Ait., from a potential biological control agent for broom, Cytisus scoparius (L.) Link. New Zealand Journal of Zoology 22: 305-309.
Tallamy D.W. (1999). Physiological issues in host range expansion. Pp. 11-26 In: Proceedings: Host specificity testing of exotic arthropod biological control agents: The biological basis for improvement in safety, N.R. Spencer (Ed.) Bozeman, Montana.
Taylor D.B.J., Heard T.A. and Jacob H.S. (2004). How effective is host testing at predicting non-target impacts of weed biological control agents in Australia? Pp. 91-94 In: 14th Australian Weeds Conference: Weed management: balancing people, planet, profit, B. M. Sindel and S. B. Johnson (Ed.) Wagga Wagga, New South Wales, Australia, 6-9 September 2004
Taylor D.B.J., Heard T.A., Paynter Q. and Spafford H. (2007).
Nontarget effects of a weed biological control agent on a native plant in Northern Australia.
Biological Control 42: 25-33
Pre-release laboratory tests predicted that Neurostrota gunniella, an agent released in Australia against Mimosa pigra, may occasionally use Neptunia spp. as hosts. However, it was not expected to persist on Neptunia spp., nor have a significant effect. N. gunniella has established widely and is now abundant on the target weed, which grows sympatrically with at least one species of Neptunia. Nontarget attack of Neptunia major in the field has been investigated, and although an average of 61% of N. major plants growing adjacent to M. pigra thickets had evidence of attack, this was relatively low. Where M. pigra was not present, use of N. major plants by N. gunniella was noticeably reduced or absent. Post-release results support the predictions made during prerelease studies of N. gunniella.
Thomas H.Q., Zalom F.G. and Roush R.T. (2010). Laboratory and field evidence of post-release changes to the ecological host range of Diorhabda elongata: Has this improved biological control efficacy? Biological Control 53 (3): 353-359
Thomas M. and Waage J.K. (1996). Integration of biological control and host plant resistance breeding. A scientific and literature review. Technical Centre for Agricultural and Rural Cooperation (CTA), 99 pp.
Thomas M.B. and Willis A.J. (1998).
Biocontrol - risky but necessary?
Trends in Evolution and Ecology 13: 325-329.
Biocontrol advocates appear reluctant to accept the possibility that there could be side-effects associated with biocontrol, often refuting evidence of harmful effects. The biocontrol critics, although eager to provide evidence to the contrary, appear reluctant to propose any detailed, constructive criticisms or workable solutions. Suggestions for directions for future research that might help to resolve some of the problems.
Thompson W.R. and Simmonds F.J. (1964-1965). A Catalogue of the Parasites and Predators of Insect Pests. Commonwealth Agricultural Bureaux, Bucks, United Kingdom.
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.
Todd J.H., Ramankutty P., Barraclough E.I. and Malone L.A. (2008). A screening method for prioritizing non-target invertebrates for improved biosafety testing of transgenic crops. Environmental Biosafety Research 7: 35-56.
Toepfer S., Cabrera W.G., Eben A., Alvarez-Zagoya R., Haye T., Zhang F. and Kuhlmann, U. (2008).
A critical evaluation of host ranges of parasitoids of the subtribe Diabroticina (Coleoptera: Chrysomelidae: Galerucinae: Luperini) using field and laboratory host records.
Biocontrol Science and Technology 18: 483-504
The subtribe Diabroticina are New World Chrysomelidae including corn rootworms, cucumber beetles and other pests. The only parasitoids that consistently target and develop inside the beetle adults are Centistes gasseni Shaw, Centistes diabroticae Gahan (both Hym.: Braconidae), and Celatoria diabroticae Shimer, Celatoria compressa (Wulp), Celatoria bosqi Blanchard, and Celatoria setosa Coquillett (all Diptera: Tachinidae). The authors present new host records and data from laboratory host range tests showing that all tachinid and braconid species studied are considered to be specific at the level of subtribe. The realized and potential host range of Centistes diabroticae includes Acalymma species as well as species in the fucata and virgifera groups of Diabrotica. Celatoria compressa has the broadest realised range compared to the other species studied, since it was obtained from species in several genera of Diabroticina; and its potential host range may also include Old World Aulacophora species.
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.
Turlings T.C.J., Tumlinson J.H., Heath R.R., Proveaux, A.T. and Doolittle R.E. (1991). Isolation and identification of allelochemicals that attract the larval parasitoid, Cotesia marginiventris (Cresson), to the microhabitat of one of its hosts. Journal of Chemical Ecology 17: 2235-2251
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Unruh T.R. and Messing R.H. (1993).
Intraspecific biodiversity in Hymenoptera: implications for conservation and biological control.
Pp. 27-52 In: Hymenoptera and Biodiversity, J. LaSalle and I. D. Gauld (Ed.) CAB International, Wallingford; UK.
Selected genetic and biological attributes of the Hymenoptera which differ from those of other insect groups are discussed in relation to modes of reproduction, sex determination and gender allocation and sex ratio. Genetic variation and population viability, single locus variation and quantitative variation are considered. Genetic load and inbreeding depression are discussed in relation to haplodiploidy and thelytoky. Intraspecific variation in response to abiotic factors, host suitability and in susceptibility to toxins is considered in the final section.
Uygur S., Smith L.A., Uygur F.N., Cristofaro M. and Balciunas J.K. (2005). Field assessment in land of origin of host specificity, infestation rate and impact of Ceratapion basicorne a prospective biological control agent of yellow starthistle. BioControl 50: 525-541.
Vayssières J.F. and Wapshere A.J. (1983). Life-histories and host specificities of Ceutorhynchus geographicus (Goeze) and C. larvatus Schultze (Coleoptera: Curculionidae), potential biological control agents for Echium. Bulletin of Entomological Research 73: 431-440.
Vet L.E.M. and Dicke M. (1992). Ecology of infochemical use by natural enemies in a tritrophic context. Annual Review of Entomology 37: 141-172
Vet L.E.M. and Godfray H.C.J. (2008). Multitrophic interactions and parasitoids behavioural ecology. Pp. 231-253 In: Behavioural ecology of insect parasitoids: from theoretical approaches to field application, E. Wajnberg, C. Bernstein and J.J.M. Van Alphen (Ed.) Blackwell Publishing, Oxford, UK
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Vink C.J., Phillips C.B., Mitchell A.D., Winder L.M. and Cane R.P. (2003). Genetic variation in Microctonus aethiopoides (Hymenoptera: Braconidae). Biological Control 28: 251-264
Vinson S.B. (1990). How parasitoids deal with the immune system of their host: an overview. Archives of Insect Biochemistry and Physiology 13: 3-27.
Vinson S.B. (1998). The general host selection behavior of parasitoid Hymenoptera and a comparison of initial strategies utilized by larvaphagous and oophagous species. Biological Control 11: 79-96
Vitou J., Skuhrava M., Skuhravy V., Scott J.K. and Sheppard A.W. (2008).
The role of plant phenology in the host specificity of Gephyraulus raphanistri (Diptera: Cecidomyiidae) associated with Raphanus spp. (Brassicaceae).
European Journal of Entomology 105: 113-119
Recent host records for Gephyraulus raphanistri (Kieffer), a flower-gall midge, indicate that it is restricted to Raphanus raphanistrum throughout Europe. This study tested host specificity of G. raphanistri in the field in Europe by manipulating host plant phenology of actual and potential hosts in the genera Raphanus and Brassica as part of a risk assessment of the insect as a potential biological control agent of R. raphanistrum, one of the most important weeds of crops in Australia. The high field specificity of this gall midge was shown to be driven by the synchrony of oviposition and flower availability, not host physiological incompatibility or behavioural unacceptability. Commercially grown brassicas are not suitable hosts because in the field they differ in flowering phenology from Raphanus raphanistrum. The overlap in the flowering phenology of the crop and weed in Australia makes this insect unsuitable as a biological control agent.
Voegele J.M., Klingauf F. and Engelhardt T. (1989). Studies on the economic returns of biological pest control with a case study from Western Samoa. Gesunde Pflanzen 41: 255-258.
Waage J.K. (2001). Indirect ecological effects of biological control: the challenge and the opportunity. Pp. 1-12 In: Evaluating indirect ecological effects of biological control, E. Wajnberg, J.K. Scott and P.C. Quimby (Ed.) CABI Publishing, Wallingford, Oxon., UK.
Waipara N.W., Barton J., Smith L.A., Harman H.M., Winks C.J., Massey B., Wilkie J.P., Gianotti A.F. and Cripps, M.G. (2009).
Safety in New Zealand weed biocontrol: a nationwide pathogen survey for impacts on non-target plants.
New Zealand Plant Protection 62: 41-49.
Nationwide disease surveys conducted between 2000-2009 focused on species closely related to target weeds most at risk of attack. Disease damage attributable to biocontrol agents was observed on two non-target plants. Pustules of the blackberry rust, Phragmidium violaceum, were found on the endemic Rubus species, R. cissoides (bush lawyer, tataramoa) at one location as predicted from pre-release host range tests. No non-target damage was observed in the remaining case studies.
Wang Y., Ding J. and Guoan Z. (2008).
Gallerucida bifasciata (Coleoptera : Chrysomelidae), a potential biological control agent for Japanese knotweed (Fallopia japonica).
Biocontrol Science & Technology 18: 59-74
Japanese knotweed, Fallopia japonica (Houttuyn) Ronse Decraene (Polygonaceae) is an invasive weed in the United States and Europe. A leaf beetle, Gallerucida bifasciata (Coleoptera: Chrysomelidae) is an important natural enemy attacking this plant in Asia. However, its host range records were ambiguous. This study examined the beetle's host specificity through a set of choice and no-choice tests in the laboratory and field in its native China. Gallerucida bifasciata larvae were able to complete development on seven of 87 plant species in larval development tests, while adults fed and oviposited on 10 plants in no-choice tests. Multiple choice tests showed adults strongly preferred F. japonica, Persicaria perfoliata (L.) H. Gross and Polygonum multiflorum Thunb over all other plants. Open field tests and field surveys further revealed that these three species were in its ecological host range. The results of this study suggest that G. bifasciata is a potential promising agent for control of Japanese knotweed in the United States and Europe, although additional host specificity tests and risk assessment is required.
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Watson M.C., Withers T.M. and Hill R.L. (2009).
Two-phase open-field test to confirm host range of a biocontrol agent Cleopus japonicus.
New Zealand Plant Protection 62: 184-190.
The buddleia leaf weevil, Cleopus japonicus, was released in New Zealand in 2006 as a biological control agent for the weed Buddleja davidii. A field study investigated any impacts on six non-target plant species since release. C. japonicus strongly preferred B. davidii, but larvae were recorded on Verbascum virgatum and Scrophularia auriculata during the choice stage of the trial. Removing B. davidii plants resulted in adults feeding on the same two exotic species. Minor exploratory feeding was recorded on the natives Hebe speciosa and Myoporum laetum. These results confirm that laboratory tests have accurately predicted field host range.
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Host-range testing: safety and science.
Pp. 83-96 In: Microbial control of weeds, D.O. TeBeest (Ed.) Chapman and Hall Ltd, London, UK.
The review concludes that whilst phylogenetic based testing is useful it is not a completely predictable system. Genetic variation in hosts and pathogens must be taken into account in the design of future tests and this requires greater understanding of host specificity, adaptation and genetic diversity.
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A biological control feasibility study of the invasive weed-air potato, Dioscorea bulbifera L. (Dioscoreaceae): an effort to increase biological control transparency and safety.
Natural Areas Journal 27: 269-279
The invasive weed Dioscorea bulbifera L. threatens the biodiversity of many natural areas in the southeastern United States. The authors propose that this weed will be a relatively safe target for biocontrol because of taxonomic and geographic isolation from desirable native and economic plant species. The family Dioscoreaceae is poorly represented in North America, north of Mexico, and the two native species that are sympatric with the weed are from a different subgeneric taxon than the weed. The West Indian and northern Mexican species, while more diverse, are also assigned to different subgeneric taxa, and are geographically isolated from the northern range of the weed. Further research pre-release needs to better delimit the geographic origin of the weed's North American population within its large native range to aid in the detection of suitable natural enemies. This will ensure that potential conflicts and risks can be judged and addressed during the projects to ultimately produce safer, more acceptable agents for biological control.
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The potential for indirect effects between a weed, one of its biocontrol agents and native herbivores: A food web approach.
Biological Control 35: 299-306.
Constructing and analyzing food webs may be a valuable for assessing the post-release safety of control agents. How food webs can be used to generate testable hypotheses regarding indirect interactions between introduced agents and non-target species is shown using an exotic weed, bitou bush, Chrysanthemoides monilifera ssp. rotundata, and a key biocontrol agent for this weed in Australia, the tephritid fly, Mesoclanis polana. Food webs showed the interactions between plants, seed-feeding insects and their parasitoids.
Withers T. and Barton-Browne L. (1998). Possible causes of apparently indiscriminate oviposition in host specificity tests using phytophagous insects. Pp. 565-571 In: Pest Management - Future Challenges: Proceedings of the 6th Australasian Applied Entomological Research Conference, M. Zalucki, R. Drew and G. White (Ed.) Brisbane, The Cooperative Research Centre for Tropical Pest Management.
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Choice or no-choice tests? Effects of experimental design on the expression of host range.
Pp. 620-633 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.
Estimation of the host range of entomophagous biological control agents (parasitoids and predators) is complex. It is argued that initial test procedures should maximize the probability that the test species will be accepted for oviposition, particularly using no-choice tests. Sequential no-choice tests should only be used with caution as they have the potential to produce false negative results. Choice tests including the target host have the potential to mask the acceptability of lower ranked hosts, thereby producing false negative results. Examples where wider host ranges have been expressed in no-choice tests than in choice tests, and vice versa are presented. Sufficient variation exists that it is recommend that researchers routinely use both assay methods for host range testing of parasitoids and predators.
Withers T.M. (1997). Changes in plant attack over time in no-choice tests: an indicator of specificity. Pp. 214-217 In: Proceedings of the 50th New Zealand Plant Protection Conference, M. O'Callaghan (Ed.) Lincoln University, NZ., New Zealand Plant Protection Society Inc.
Withers T.M. (1998). Influence of plant species on host acceptance behaviour of the biocontrol agent Zymogramma bocolorata (Col.: Chrysomelidae). Biological Control 13: 55-62.
Withers T.M. (1999). Examining the hierarchy threshold model in a no-choice feeding assay. Entomologia Experimentalis et Applicata 91: 89-95.
Withers T.M. and Barton-Browne L. (2004). Behavioral and physiological processes affecting the outcome of host range testing. Pp. 40-55 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.
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.
Withers T.M., Barton-Browne L. and Stanley J. (1999). Host specificity testing in Australasia: towards improved assays for biological control. Pp. 98. CRC for Tropical Pest Management, Brisbane, Australia.
Withers T.M., Barton-Browne L. and Stanley J. (2000). How time-dependent processes can affect the outcome of assays. Pp. 27-41 In: Host-specificity testing of exotic arthropod biological control agents: the biological basis for improvement in safety, R.G. Van Driesche, T. Heard, A.S. McClay and R. Reardon (Ed.) USDA Forest Service Bulletin, Morgantown, West Virginia, USA.
Withers T.M., Carlson C.A. and Gresham B.A. (2013). Statistical tools to interpret risks that arise from rare events in host specificity testing. Biological Control 64: 177-185.
Withers T.M., Hill R.L., Paynter Q., Fowler S.V. and Gourlay A.H. (2008).
Post-release investigations into the field host range of the gorse pod moth Cydia succedana Denis & Schiffermuller (Lepidoptera : Tortricidae) in New Zealand.
New Zealand Entomologist 31: 67-76
The gorse pod moth Cydia succedana was released in New Zealand as a biological control agent against gorse Ulex europaeus L. in 1992 and is now widely established. Post-release evaluations of host range were undertaken using both laboratory assays and field collections on native and exotic plants related to gorse. Field surveys detected no attack on native New Zealand plant species. However, contrary to predictions based on pre-release host-range testing, several species of exotic Genisteae were shown to be hosts of C. succedana. Hypotheses to explain this unexpected non-target attack include a seasonal asynchrony between C. succedana and gorse flowering phenology, or that the original biocontrol introduction accidentally consisted of either two cryptic species or two populations with different physiological host range.
Withers T.M., McFadyen R.E. and Marohasy J. (2000). Importation protocols and risk assessment for weed biological control agents in Australia: The example of Carmenta nr ithacae. Pp. 195-214 In: Nontarget Effects of Biological Control, P.A. Follett and J.J. Duan (Ed.) Kluwer Academic Publishers, Norwell, MA.
Withers T.M., Potter K.J.B., Berndt L.A., Forgie S.A., Paynter Q.E. and Kriticos D.J. (2011). Risk posed by the invasive defoliator Uraba lugens to New Zealand native flora. Agricultural and Forest Entomology 13: 99–110.
Wright M.G., Hoffmann M.P., Kuhar T.P., Gardner J. and Pitcher S.A. (2005).
Evaluating risks of biological control introductions: A probabilistic risk-assessment approach.
Biological Control 35: 338-347.
Improved quantitative procedures for estimating potential non-target impacts of biological control agents are needed and a probabilistic risk-assessment approach is proposed. The procedure described uses precision trees to estimate risk based on probabilities that biological control agents will demonstrate predictable behaviour under specific conditions, based on their ecological characteristics. Trichogramma ostriniae, an egg parasitoid for Ostrina nubilalis in the US is used as case study to demonstrate the procedure, which potential for widespread use in quantifying non-target risk of biological control introductions prior to introductions being made.
Wu K., Center T.D., Yang C., Zhang J., Zhang J. and Ding J. (2013). Potential classical biological control of invasive Himalayan Yellow Raspberry, Rubus ellipticus (Rosaceae). Pacific Science 67: 59-80.
Wyckhuys K.A.G., Koch R.L., Kula R.R. and Heimpel G.E. (2009).
Potential exposure of a classical biological control agent of the soybean aphid, Aphis glycines, on non-target aphids in North America.
Biological Invasions 11: 857-871
In 2007, the Asian parasitoid Binodoxys communis (Hymenoptera: Braconidae) was released in North America for control of the exotic soybean aphid, Aphis glycines (Hemiptera: Aphididae). To estimate the risk of exposure of non-target aphids to B. communis, the authors merged assessments of temporal co-occurrence with projections of spatial overlap between B. communis and three native aphid species. The degree of temporal overlap depended greatly on location and the non-target aphid species. Geographic overlap between B. communis and native aphids based upon Climex modeling was used to assess the degree of geographic overlap between the parasitoid and non-target species. The authors were able to make broad statements regarding the ecological safety of current B. communis releases and their potential impact on native aphid species in North America.
Yanek M.L. and Raffa K.F. (2008).
Evaluation of Gypchek and its carrier on various Lepidoptera species under laboratory conditions.
Great Lakes Entomologist 41: 27-39
Gypchek is a gypsy moth-specific nucleopolyhedrosis virus applied in conjunction with Carrier 038-A, a surfactant and sunscreen. Carrier 038-A alone and with Gypchek was tested on an endangered species, the Karner blue butterfly ( Lycaeides melissa samuelis Nabokov), tobacco hornworm ( Manduca sexta L.), L. dispar , and seven other Lepidoptera species. Where there was evidence of nontarget effects with a high dose the authors continued with a method to approximate field application. Eight species were unaffected. Gypsy moth was sensitive to Gypchek as expected but not to its carrier, and L. m. samuelis showed putative effects which warranted further testing. With simulated field spray-deposition, there was no impact of carrier 038-A or Gypcheck with carrier on L. m. samuelis larvae. The authors further concluded that the partial canopy cover of oak savannas and partially asynchronous phenology with L. dispar would reduce exposure to L. m. samuelis .
Yara K., Sasawaki T. and Kunimi Y. (2010). Hybridization between introduced Torymus sinensis (Hymenoptera: Torymidae) and indigenous T. beneficus (late-spring strain), parasitoids of the Asian chestnut gall wasp Dryocosmus kuriphilus (Hymenoptera: Cynipidae). Biological Control 54: 14-18
Yodzis P. (1988). The indeterminancy of ecological interactions as perceived through perturbation experiments. Ecology 69: 508-515.
Zeddies J., Schaab R.P., Neuenschwander P. and Herren H.R. (2001). Economics of biological control of cassava mealybug in Africa. Pp. 209-219 In: Agricultural Economics, (Ed.) Elsevier Science B.V., Amsterdam, Netherlands.
Zimmermann G. (2007).
Review on safety of the entomopathogenic fungus Metarhizium anisopliae.
Biocontrol Science & Technology 17: 879-920
Metarhizium anisopliae (Metschn.) Sorokin is widely used for biocontrol of pest insects, and many commercial products are on the market or under development. This review summarises all relevant safety data for this fungus, which are necessary for the commercialisation and registration process. On the basis of the presented knowledge, M. anisopliae is considered to be safe with minimal risks to vertebrates, humans and the environment.
Zwölfer H. and Harris P. (1971). Host specificity determination of insects for biological control of weeds. Annual Review of Entomology 16: 159-178.
de Clercq, P., Mason, P. and Babendreier, D. (2011).
Benefits and risks of exotic biological control agents.
Biocontrol 56: 681-698
The use of exotic arthropods in biological control programs has yielded huge economic and ecological benefits. However, non-target effects of exotic biological control agents have been observed in a number of projects. Non-target effects range from very small effects, to massive effects on a large scale. This paper reviews both the benefits of biological control as well as the associated risks and an attempt is made at identifying the major challenges for assessing risks and for balancing benefits and risks. While sound risk assessment procedures preceding release of biological control agents are advocated, overly stringent regulations that would preclude promising agents from being developed must be avoided.
de Leon J.H., Neumann G., Follett P.A. and Hollingsworth R.G. (2010).
Molecular markers discriminate closely related species Encarsia diaspidicola and Encarsia berlesei (Hymenoptera: Aphelinidae): biocontrol candidate agents for white peach scale in Hawaii.
Journal of Economic Entomology 103: 3, 908-916.
The authors characterized Encarsia diapsidicola Silvestri and Encarsia berlesei Howard (Hymenoptera: Aphelinidae) by phylogenetic analysis of the cytochrome oxidase subunit I gene (COI) and intersimple sequence repeat-polymerase chain reaction (ISSR-PCR) DNA fingerprinting. These closely related parasitoids are candidate biological control agents for the white peach scale, Pseudaulacaspis pentagona Targioni-Tozetti (Hemiptera: Diaspididae), in Hawaii. Both molecular marker types successfully discriminated the two Encarsia spp., but the COI markers were considered useful to assess levels of parasitism in the field and to study competitive interactions between parasitoids.
de Nardo E.A.B. and Hopper K.R. (2004). Using the literature to evaluate parasitoid host ranges: a case study of Macrocentrus grandii (Hymenoptera: Braconidae) introduced into North America to control Ostrinia nubilalis (Lepidoptera: Crambidae). Biological Control 31: 280-295
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., Blossey B., Hoddle M., Lyon S. and Reardon R. (2002).
Biological control of invasive plants in the Eastern United States.
Forest Health Technology Enterprise Team, USDA Forest Service, Morgantown. CD-ROM
The purpose of this CD-ROM is to provide a reference guide for field workers and land managers concerning the historical and current status of the biological control of invasive plant species in the eastern USA. For each weed: pest status, nature of damage (economic and ecological), geographical distribution, background information (taxonomy, biology, related nativeplants in the eastern USA), history of biological control efforts in the eastern USA, recommendations for future work and references is given. The history of biological control efforts is divided into sections: area of origin of the weed, areas surveyed for natural enemies, natural enemies found, host range tests and results, releases made, and the biology and ecology of natural enemies.
van Driesche R.G. and Bellows T.S. (1996). Biological control. Chapman and Hall, New York. 539 pp.
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 Driesche R.G. and Hoddle M.S. (2000). Classical arthropod biological control: assessing success, step by step. Pp. 39-75 In: Biological control: Measures of success, G.M. Gurr and S.D. Wratten (Ed.) Kluwer Academic Publishers, Dordrecht, The Netherlands
van Driesche R.G. and Murray T.J. (2004). Parameters used in laboratory host range tests. Pp. 56-67 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.
van Driesche R.G. and Murray T.J. (2004). Overview of testing schemes and designs used to estimate host ranges. Pp. 68-89 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.
van Driesche R.G., Bellows T.S., Jr., Elkinton J.S., Gould J.R. and Ferro D.N. (1991).
The meaning of percentage parasitism revisited: solutions to the problem of accurately estimating total losses from parasitism.
Environmental Entomology 20: 1-7.
New analytical methods for obtaining stage-specific estimates of losses from parasitism for life-table, population dynamics, and evaluation studies are needed. Solutions considered include recruitment, stage frequency and death rate analyses. The rationale and methodology are presented and their usefulness for systems of varying types of biologies and sampling constraints is compared.
van Emden H.F. (2003). Conservation biological control: from theory to practice. In: Proceedings of the 1st International Symposium on Biological Control of Arthropods, R. Van Driesche (Ed.) United States Department of Agriculture Forest Service, Washington, USA.
van Halteren P. (1997). A code of conduct for the import and release of exotic biological control agents for Europe? Pp. 45-48 In: EPPO/CABI workshop on safety and efficacy of biological control in Europe, I.M. Smith (Ed.) Blackwell Science Ltd., Oxford.
van Klinken R.D. (2000). Host-specificity testing: why do we do it and how we can do it better. Pp. 54-68 In: Host-specificity testing of exotic arthropod biological control agents: the biological basis for improvement in safety, R.G. Van Driesche, T. Heard, A.S. McClay and R. Reardon (Ed.) USDA Forest Service Bulletin, Morgantown, West Virginia, USA.
van Klinken R.D. (2006). Biological control of Parkinsonia aculeata: what are we trying to achieve? Australian Journal of Entomology 45: 268-271
van Klinken R.D. and Edwards O.R. (2002). Is host specificity of weed biological control agents likely to evolve rapidly following establishment? Ecology Letters 5: 590-596.
van Klinken R.D. and Raghu S. (2006). A scientific approach to agent selection. Australian Journal of Entomology 45: 253-258
van Lenteren J.C. (1997). Benefits and risks of introducing exotic macro-biological control agents into Europe. Pp. 15-27 In: EPPO/CABI workshop on safety and efficacy of biological control in Europe, I.M. Smith (Ed.) Blackwell Science Ltd., Oxford.
van Lenteren J.C. (2000). Success in biological control of arthropods by augmentation of natural enemies. Pp. 77-103 In: Measures of Success in Biological Control, G. Gurr and S. D. Wratten (Ed.) Kluwer Academic Publishers, Dordrecht
van Lenteren J.C. (2006). The internet book of biological control http://www.unipa.it/iobc/downlaod/IOBC%20InternetBookBiCoVersion4October2006.pdf
van Lenteren J.C., Babendreier D., Bigler F. G. B., Hokkanen H.M.T., Kuske S., Loomans A.J.M., Menzler-Hokkanen I., Van Rijn P.C.J., Thomas M.B., Tommasini M.G. and Zeng Q.-Q. (2003).
Environmental risk assessment of exotic natural enemies used in inundative biological control.
BioControl 48: 3-38.
A methodology for risk assessment has been developed within the EU-financed project 'Evaluating Environmental Risks of Biological Control Introductions into Europe [ERBIC]' as a basis for regulation of import and release of exotic natural enemies used in inundative forms of biological control. This paper proposes a general framework of a risk assessment methodology for biological control agents, integrating information on the potential of an agent to establish, its abilities to disperse, its host range, and its direct and indirect effects on non-targets.
van Lenteren J.C., Bale J., Bigler F., Hokkanen H.M.T. and Loomans A.J.M. (2006).
Assessing risks of releasing exotic biological control agents of arthropod pests.
Annual Review of Entomology 51: 609-634.
This review summarizes documented nontarget effects of biological control agents and discusses the development and application of comprehensive and quick-scan environmental risk assessment methods.
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.
van Lenteren J.C., Loomans A.J.M., Babendreier D. and Bigler F. (2008).
Harmonia axyridis: an environmental risk assessment for Northwest Europe.
BioControl 53: 37-54
A recently designed, comprehensive risk evaluation method to evaluate the environmental risks of Harmonia axyridis showed that H. axyridis is a potentially risky species for Northwest Europe, because it is able to establish, it has a very wide host range including species from other insect orders and even beyond the class of Insecta. Its activities have resulted in the reduction of populations of native predators in North America, where it may develop as a pest of fruit. Current knowledge would lead to the conclusion that it should not have been released in Northwest Europe. In retrospect, the risks should have been sufficient to reject import and release of this species, but this was ignored. The case of Harmonia releases in Northwest Europe demonstrates an urgent need for harmonized, world-wide regulation of biological control agents, including an information system on risky natural enemy species.
van Veen F.J.F., Morris R.J. and Godfray H.C.J. (2006). Apparent competition, quantitative food webs, and the structure of phytophagous insect communities. Annual Review of Entomology 51: 187-208
van den Bosch, R., Messenger, P.S. and Gutierrez, A.P. (1982). An introduction to biological control. Intext Educational Publishers, Plenum Press, New York and London. Pp 247
Biological control agent release