Suggestions for unifying the terminology in biological control

This paper gives suggestions forunifying the terminology in biological controlacross different research disciplines, such asbiological control of arthropods, weeds andplant pathogens. It is suggested that use ofthe term `biological control' is restricted tothe use of living organisms. Four strategiesof biological control are outlined anddefined: (1) Classical biological control, (2) Inoculation biological control, (3) Inundationbiological control, and (4) Conservationbiological control. It is proposed to usethese four terms as defined, and avoid usageof the term `augmentation'. Terms for specificprocesses and modes of action (for example,`parasitoid' and `competitor') can be definedby usage within the different biologicalcontrol disciplines. Microbial control usuallyindicates biological control of invertebratesusing microbes and, as such, is a subdivisionof biological control. Use of additionalauxiliary terms such as biopesticide isdiscussed.     

Mowing during rainfall enhances the control of Cirsium arvense

Pastoral farmers in New Zealand have described dramatic demises in populations of the weed Cirsium arvense (L.) Scop., a perennial herb indigenous to Eurasia, following its mowing during rainfall. To test the hypotheses (1) that the mowing of C. arvense during rainfall increases the control of this weed and (2) that the causal organism in this ‘mowing-in-the-rain’ effect is the vascular wilt fungus Verticillium dahliae, two series of field experiments were carried out in C. arvense-infested pastures in New Zealand, one in autumn 2008 (Experiment series 1, 9 farms), and another in spring/summer 2008–2009 (Experiment series 2, 12 farms). The effect of mowing in the rain as compared to mowing in the dry was to reduce the % ground cover of the thistle in the spring following treatment by 21 and 32% in Experiment series 1 and 2, respectively. Correlations of this ‘rain versus dry’ effect with the incidence of V. dahliae in the subterranean parts of C. arvense shoots sampled in each of the two field experiment series provided no statistical evidence that the effect increased with V. dahliae incidence. Thus these experiments provide no support for the hypothesis that V. dahliae is the biological mechanism for the ‘mowing-in-the-rain’ effect. Nevertheless, they do support mowing during rainfall as a simple and effective management tactic for C. arvense.     

Host-specificity of Uromyces heliotropii, a fungal agent for the biological control of common heliotrope (Heliotropium europaeum) in Australia

Common heliotrope, Heliotropium europaeum (Boraginaceae), a summer annual, is a serious weed of pastures in Australia causing poisoning in sheep and other grazing animals. The weed is native to Mediterranean and Middle Eastern regions of Eurasia where it is attacked by several arthropods and pathogens including the rust fungus Uromyces heliotropii. The host-specificity of a virulent and effective strain of the rust from Turkey has been tested by inoculating 96 plants of importance to the Australasian region using both microscopic and macroscopic observations of the reaction of host and non-host plants. The test plants included several members of the Boraginaceae and related families as well as species of Heliotropium native to Australia. The infection was mostly limited to European species of Heliotropium. U. heliotropii was thus found to represent a safe introduction and has recently been introduced to Australia for the biological control of common heliotrope.     

Host specificity of Metriona elatior, a potential biological control agent of tropical soda apple, Solanum viarum, in the USA

The leaf beetle Metriona elatior from Brazil-Argentina was screened in the Florida (USA)State quarantine facility as a potential biological control agent of tropical soda apple, Solanum viarum, a recently arrived weed species. Multiple-choice host-specificity tests were conducted in small cages (60 cm × 60 cm × 60 cm) using 95 plant species in 29 families. Adults fed heavily on the main target weed (S. viarum), and on turkey berry,Solanum torvum (noxious weed of Asiatic origin); fed moderately on red soda apple, Solanum capsicoides (weed of South American origin), and eggplant, Solanum melongena (economic crop); and fed lightly on aquatic soda apple, Solanum tampicense (weed of Mexican-Caribbean-Central American origin), and onsilverleaf nightshade, Solanum elaeagnifolium(native weed widely distributed). M.elatior adults laid 84 to 97% of their egg masses onS. viarum, and 3 to 16% on S. melongena. Non-choice host-specificity tests were also conducted in quarantine in which M. elatior adults and neonate larvae were exposed to 17 and 19 plant species, respectively. Tests with the neonates indicate that this insect was able to complete its development on S. viarum, S. torvum, S. melongena, and S. capsicoides. Although some adult feeding and oviposition occurred on S.melongena in quarantine on potted plants in small cages, no feeding or oviposition by M. elatiorwas observed in field experiments conducted in Brazil. Surveys in unsprayed S. melongena fields in Argentina and Brazil indicated that M. elatioris not a pest of S. melongena in South America. The evidence obtained from the South-American field surveys, Brazil open-field experiments, and Florida quarantine host specificity tests indicate that M. elatior causes significant feeding damage toS. viarum, and does not represent a threat to S. melongena crops in the USA. Therefore an application for permission to releaseM. elatior against S. viarum in the USA was submitted in October 1998. This revised version was published online in July 2006 with corrections to the Cover Date.     

Quantitative resistance of Cirsium arvense to root bud infection by Puccinia punctiformis

A method is presented to test clones of Cirsium arvense for resistance to root bud infection by the rust fungus Puccinia punctiformis. Root buds were stained and cleared to detect mycelium of the rust. The fraction of root buds infected was determined and used as a measure of resistance. Clones of C. arvense, collected from three sites, were tested for resistance to infection. Variation in resistance was determined between and within sites. The results are discussed with a view to using P. punctiformis to control C. arvense.     

Field assessment,in Greece and Russia,of the facultative saprophytic fungus,Colletotrichum salsolae,for biological control of Russian thistle (Salsola tragus)

Russian thistle (Salsola tragus, tumbleweed, RT) is a problematic invasive weed in the United States (U.S.) and is a target of biological control efforts. The facultative saprophytic fungus Colletotrichum salsolae (CS) kills RT plants in greenhouse tests and is specific to Salsola spp., which are not native in the U.S. However, the effectiveness of CS in controlling RT has not been previously demonstrated. The objectives of this study were to determine in field tests: (1) disease progress of CS in time; (2) the relationship of disease progress to rainfall and temperature; (3) the effect of CS on RT plant density. Field tests were established in Serres and Kozani, Greece and Taman and Tuzla, Russia with isolates of the pathogen collected in the respective countries. Solid inoculum was prepared by asceptically inoculating sterile mixtures of grain and grain hulls with axenic cultures of CS. Spore suspensions used in Russia were prepared by blending pure sporulating cultures of CS with distilled water and diluting the suspension to 10⁶ conidia per ml. Six field plots, each subdivided into 36 subplots, of an RT infested field in Serres, Greece were inoculated on October 23, 2006 by placing about 300 g of solid inoculum in the center of each plot. Four field plots, similarly subdivided, in each of two fields at Kozani, Greece were inoculated in the same way on October 1, 2010. RT density was counted and recorded in each sub-plot prior to inoculation and in September in each of 2 years following inoculation. Disease incidence and/or severity in each sub-plot were recorded at about 7-days intervals after inoculation. Rainfall and temperature data, from inoculation until 40–55 days after inoculation, were collected and recorded at Serres and at the Kozani airport meteorological station. Disease progressed rapidly at both sites and was correlated with cumulative rainfall. By 2 years after inoculation, RT had been eliminated from the Serres site and one field in Kozani. In the other Kozani field, RT density declined to 0–25% from original densities of about 80% in large areas of the field. RT plants in Taman and Tuzla, Russia were inoculated either with 250 g of grain inoculum or with a suspension of 10⁶ conidia sprayed onto each plant until runoff. The proportion of diseased tissue reached 1.0 by 55 days after inoculation in both sites. Non-inoculated plants that were near inoculated plants became diseased quickly and reached the same disease severity as inoculated plants. Disease severity was correlated with cumulative rainfall but not temperature. This pathogen and inoculation procedure offers a low-cost solution to RT infestations. Since CS is specific to Salsola spp., this effective biological control is also environmentally friendly.     

Laboratory life history and field observations of Poliopaschia lithochlora (Lower) (Lepidoptera: Pyralidae), a potential biological control agent for Melaleuca quinquenervia (Myrtaceae)

Abstract  Melaleuca quinquenervia (Cav.) S.T. Blake, Australian broad-leaved paperbark, has become a serious weed in southern Florida. Poliopaschia lithochlora (Lower) is a promising candidate as a potential biological control agent, and this study describes laboratory and field observations of the life history of this moth. Eggs are laid in small batches, mainly on the surface of leaves, and larvae are voracious leaf feeders, concealed in tubes that are usually found in small colonies attached to leaves and stems. Larvae move from these tubes to feed on surrounding leaves, and saplings and suckers are frequently defoliated. Prepupae form sealed bulbs in the larval tubes in which they pupate. Adult females are mainly active and oviposit at night. Development from egg to adult occurs in approximately 80 d. Field populations appear to be regulated by several egg and larval parasites. Because this moth severely damages saplings and suckers of M .  quinquenervia , prefers low-lying humid sites, and can be successfully mass reared, it is rated highly for its potential as a biological control agent.     

A review of white rust (Puccinia horiana Henn.) disease on chrysanthemum and the potential for its biological control with Verticillium lecanii (Zimm.) Viégas

The biology, aetiology and epidemiology of Puccinia horiana, the cause of white rust disease of chrysanthemum (Chrysanthemum spp.) is reviewed in relation to current environmental, cultural and chemical methods for its control. Importantly, basidiospore release, germination and infection can take as little as 5 h at optimum r.h. (96%) and temperature (between 17–24°C). Recent developments using the fungus Verticillium lecanii for the control of insects on glasshouse-grown all-year-round chrysanthemums rely upon the maintenance of r.h. during night periods in excess of 95%, thus predisposing plants to white rust attack. However, V. lecanii is unusual in that it can also parasitise spores and fruiting structures of a range of rust fungi including P. horiana. This mycoparasitic ability is also reviewed, and against this background, the potential for an integrated insect and white rust control programme on all-year-round chrysanthemums is assessed.     

The potential for biological control of invasive alien aquatic weeds in Europe: a review

A retrospective analysis shows that invasive, alien, free-floating and emergent aquatic weeds in Europe are good targets for classical biological control, and that genus-specific chrysomelid and curculionid beetles offer the most potential. Ludwigia spp., Azolla filiculoides, Lemna minuta, Crassula helmsii and Hydrocotyle ranunculoides should be prioritised as targets. Fungal pathogens have been under-utilised as classical agents but, whilst they may have some potential against free-floating weeds, they appear to be poor candidates against submerged species, although the suitability of arthropod agents against these difficult targets still merits investigation. The use of indigenous pathogens as inundative agents (mycoherbicides) shows some promise.     

Classical biological control program for the mealybug Oracella acuta in Guangdong Province,China

Abstract The mealybug Oracella acuta, native to the southeastern US, was accidentally introduced into slash pine plantations in Guangdong Province in China in 1988. A classical biological control program was initiated in 1995, and the parasitoids Allotropa oracellae, Acerophaus coccois, and Zarhopalus debarri were imported from the US. A total of 19 972 parasitized mealybugs were shipped to China from 1996–2004, from which 15 430 wasps emerged, 12 933 of which were the three target species. Efforts to establish a mass-rearing program for the parasitoids in China failed. Five field release sites were established, and 6 020 parasitoids were released. Only 118 individuals of the three imported species were collected during establishment checks, although several wasps were collected 1–2 years after the last parasitoid release. Over 2 000 Anagyrus dactylopii, a cosmopolitan parasitoid, emerged from the parasitized mealybugs collected, a majority from the Taishan area near the site of the original introduction of O. acuta. To date the imported parasitoids have failed to establish, and natural enemies have not noticeably reduced mealybug populations.     

Climatic analysis to determine where to collect and release Puccinia jaceae var. solstitialis for biological control of yellow starthistle

Puccinia jaceae var. solstitialis is an autoecious rust fungus that is native to areas of Afro-Eurasia with a Mediterranean climate. An isolate collected near Sivas, Turkey was released for classical biological control of yellow starthistle (YST), which is an invasive alien weed in California, USA. The fungus has been released throughout California, but long-term establishment rates are generally low, apparently because this ecotype is not well adapted to the climate where the weed is most invasive. Using a site with excellent establishment as a target, the Match Climates function in CLIMEX climate modeling software identified similar sites in and around the San Francisco Bay Area, east to the Central Valley and Sierra foothills, and along the coast of Southern California. Similar sites in other states include Walla Walla, Washington, Pendelton, Oregon and Salt Lake City, Utah. A Compare Locations model based primarily on experimentally measured temperature and humidity requirements of the rust produced similar results. Using Sacramento, California, which is in the center of YST distribution, as a target, the Match Climates function predicted that the best locations to search for rust accessions to use in California are near Tunis, Tunisia, Foggia, Italy, Khalkis, Greece, Kayseri, Turkey, and possibly Constantine, Algeria. This generally agrees with the prediction of a Compare Locations model based on the geographic distribution of YST in California. Climatic factors that limit the long-term establishment of the fungus are likely to be summer heat and/or dry stress and short dew periods.     

The biology and host specificity of the onion weed rust, Puccinia barbeyi, a potentially useful agent for biological control in Australia

Onion weed, Asphodelus fistulosus L., (Liliaceae) a weed of Mediterranean and Middle Eastern origin is widespread in southern Australia where it invades pastures making them unsuitable for grazing. A program of research is underway to discover natural enemies of this plant and to study their possible role in the biological control of onion weed. A rust fungus Puccinia barbeyi (Roum.) Magnus has been found to severely attack A. fistulosus . Observations on the biology of the rust confirmed that it is monoecious and microcyclic and multiplied essentially by aecial and telial stages, although occasionally urediniospores also appeared among teliospores. Several members of the Liliaceae exposed to the aeciospores of the rust remained unattacked indicating that it is most probably specific to Asphodelus spp. and thus its potential for the biological control of A. fistulosus in Australia should be studied further.     

Evaluation of four aphidiine parasitoids for biological control ofAphis gossypii

Four aphidiine parasitoid species (Hymenoptera: Braconidae) were evaluated with respect to their potential to controlAphis gossypii Glover (Homoptera: Aphididae) in glasshouse cucumbers. In a laboratory experiment thirty cotton aphids were offered to individual females for two hours.Aphidius matricariae Haliday parasitized less than six percent of the aphids and was ruled out as potential biological control agent.Ephedrus cerasicola Stary andLysiphlebus testaceipes Cresson parasitized 23 and 26 percent of the aphids, respectively.Aphidius colemani Viereck parasitized 72 to 80 percent of the aphids. With the latter three species, experiments were performed in small glasshouses with cucumbers (Cucumis sativus L. cv. ‘Aramon’). As in the laboratory testA. colemani performed best; significantly more colonies were found and parasitization rates in the colonies were higher byA. colemani than byE. cerasicola andL. testaceipes. Because of the good correspondence between laboratory and glasshouse experiments, it is suggested that bad performance of an aphid parasitoid species in a simple laboratory trial might be sufficient evidence to disregard this species for further tests.     

An analysis of historical trends in classical biological control of arthropods suggests need for a new centralized database in the USA

The documentation of biological control agents targeting arthropods in the United States has historically been subject to less regulation relative to weed biological control releases. This study reviews publicly available databases to track environmental releases of biological control agents targeting arthropods in the United States. It then presents available data for the states with the most releases between 1962 and 2005: Hawaii, California and Florida. These data indicate a clear decline in rates of introduction since 1982 or 1994, depending on the source. Existing record-keeping systems offer incomplete or inconsistent data for evaluation because they were designed with limited goals, attempt to capture excessive detail and are thus impractical, or are insufficiently resourced. Existing databases cannot be used to answer meaningful questions regarding non-target effects of introduced control agents. Current databases are inappropriately designed and insufficiently resourced to meet today's research and regulatory needs. We propose and describe a new database system for classical biological control of arthropods.     

外来入侵杂草传统生物防治的生态风险及其防范对策

长期大量实践说明,引进天敌防治外来入侵杂草的传统生物防治方法是治理外来入侵杂草的一条切实可行的有效途径,但对其潜在的生态风险——对本土生物的直接或间接不良影响不容忽视。利用传统评价方法预测候选天敌的生态风险存在缺陷,主要表现在：（1）寄主专一性测定过分依赖室内进行的生理寄主范围测定结果,对生态寄主范围（实际寄主范围）问题重视不够,后者指在新环境中的一系列物理和生物条件下的寄主利用预测;（2）在生理寄主范围测定中,过分依赖完成生长发育的可能性,对行为、遗传性状以及系统发育关系重视不够;（3）在风险评估中,过多强调对经济作物的风险,而对自然生态系统的风险重视不够。对此,建议：（1）鼓励对已释放的天敌进行回顾性跟踪研究,从而为杂草生物防治实践提供生态学理论支撑;（2）在运用生物防治手段对付外来入侵杂草实践中,建议采用“有害推论”的预防性原则,以避免在面临入侵生物重大威胁时草率做出释放天敌的决策;（3）在评估候选天敌风险中重视生态效应的风险评估。     

Tolypocladium cylindrosporum gams (deuteromycetes,moniliaceae) as a biological control agent against the window midge Sylvicola cinctus Fabr. (diptera: Anisopodidae)

Larvae of Sylvicola cinctus Fabr. were found killed by an entomopathogenic fungus Tolypocladium cylindrosporum Gams, in Vetlanda, Sweden. This is the first report of this fungus as an insect pathogen from Sweden. The fungus was cultured and sprayed on aspen log piles in an attempt to reduce population levels. In September 1992, when maximum abundance is expected, the area treated with T. cylindrosporum had significantly fewer midges than an untreated area. Possibilities for biological control are discussed.     

Host specificity testing,release and establishment of Urophora solstitialis (L.) (Diptera: Tephritidae), a potential biological control agent for Carduus nutans L., in Australia

The seed‐fly Urophora solstitialis was imported into Australia from southern France. Previous workers had shown that this seed‐fly limited its attack to selected members of the genus Carduus. Host specificity studies revealed that no native Australian plant tested from the family Asteraceae was a suitable host plant. Permission to release the fly was granted, and it was released in three regions of the New South Wales tablelands. The fly underwent a second generation at two of the release sites, and reasons for its failure to do so at the third are suggested. It has successfully established itself in all three regions, having completed a full seasonal cycle in the field.