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General News

TEAMwork against Leafy This biologically based Integrated Pest ness (diversity) increased by averages of Spurge Management (IPM) programme focuses on 47% and 27%, respectively. researching and demonstrating effective, Multi-Species Grazing Biological control programmes for leafy affordable and ecologically sustainable spurge (Euphorbia esula complex) in North leafy spurge management techniques. The TLS is demonstrating that a combination of America have made significant progress cooperative programme stresses teamwork, multi-species grazing and biocontrol works since BNI’s last report on the problematic resulting in a vast network of partnerships well as an effective spurge management Eurasian perennial [BNI 18(3),65N(Sep- between the two USDA agencies, land tool. The concept is simple: use sheep to tember 1997)]. grant universities and numerous other local, graze heavy, dense patches of spurge, thus Although 15 biological control agents for state and federal entities. Fields of study giving flea increased opportunities leafy spurge have been released in the USA include (but are not limited to) classical to establish populations in the resultant since 1963, two in particular are well estab- biological control, multi-species grazing, thinner stands of spurge. For this demon- lished throughout the continental ‘spurge herbicide use, cultural controls (such as stration, TLS selected a site that originally belt’ in the northern Great Plains (North tillage, reseeding and burning), the integra- featured an extremely dense, widespread and South Dakota, Montana, Wyoming and tion of various control tools, socio-eco- spurge infestation (more than 50% of the the Prairie Provinces of Canada). The two nomic impacts, the application of GPS/GIS total demonstration area was infested with European flea beetles – nigris- and hyperspectral imaging technologies to spurge exceeding 200 stems/m²). In just 3 cutis and Aphthona lacertosa, first released weed management, and foreign exploration years, the combination of multi-species in the USA in 1988 and 1994, respectively to investigate new leafy spurge biocontrol grazing and biocontrol has reduced spurge – have established large populations at agents. densities by 31-50%; native vegetation and desirable grasses are responding by rees- numerous locations across a wide geo- Data collected by TLS programme partici- tablishing in areas formerly occupied by graphic area, and are now providing signif- pants in the past 3 years are especially spurge. Based on previous research, signif- icant levels of leafy spurge control. Perhaps promising, and suggest that widespread icant reductions are expected to occur in the more importantly, this build-up of numbers control of leafy spurge with biologically 4th and 5th years (i.e. 2001 and 2002) of the and sites is allowing for easy collection and based IPM strategies is no longer a question demonstration. The demonstration clearly redistribution of flea beetles, which further of ‘if’ but ‘when’. This is a brief summary shows the economic and environmental promotes the concept of biological control. of some TLS research efforts: advantages offered by combining the two The past 2 years have been especially Biocontrol biologically based IPM strategies. exciting for those involved with leafy Flea establishment has improved Herbicides spurge management. Although reductions dramatically the past 3-5 years, primarily in spurge densities due to flea beetle Herbicide use is declining in areas where because people now know more about activity have been documented at many biologically based IPM programmes have using biocontrol and have access to large locations for several years, never before been implemented. Ward County, located numbers of . A few years ago, have there been so many dramatic reduc- in northwestern North Dakota, provides an releases of 100-500 flea beetles were tions spread across such a wide area. At excellent example. Ten years ago, weed common, with roughly one-third of all some sites, extremely heavy infestations managers there invested considerable releases successfully establishing a popula- were completely eliminated. Biocontrol resources (both fiscal and personnel) into tion; releases of 3000 flea beetles are now worked so well in some places, in fact, that spraying 8000 of the 12,000 acres [3250/ considered the minimum, and establish- collection efforts were hampered: people 4850 ha] infested with spurge within the ment success has more than doubled. In an returned to sites that had historically pro- county. Last year, after just 3 years of effort to quantify flea beetle establishment, duced large numbers of flea beetles to dis- aggressive biocontrol efforts, the county population expansion and the resultant cover that both the spurge and flea beetles had 9500 acres [3850 ha] of spurge (a 20% impact on leafy spurge, TLS established were gone. This phenomenon can be reduction in total acreage) and used herbi- 264 ‘inventory and assessment’ sites in expected to become more common as flea cides on just 400 acres [160 ha] (a 95% 1998. Each site was inventoried – i.e. beetle populations increase and more sites reduction). Similar examples can be extensive data regarding soil type, mois- are established, and it emphasizes the need expected from across the region as weed ture, topography, species composition, etc. to educate people about properly managing managers learn how to use biological con- were collected – prior to being seeded with their biocontrol resource. trol as a stand-alone tool and in combina- 6000 Aphthona flea beetles (3000 A. lacer- tion with other tools. A major development in leafy spurge man- tosa and 3000 A. nigriscutis). Data col- agement occurred in 1997, when the US lected from these sites in the summer of In addition to creating cooperative partner- Department of Agriculture (USDA) Agri- 2000 document an establishment success ships and funding various research and cultural Research Service joined forces rate of 85-95%, a seven-fold increase in demonstration projects, TEAM Leafy with a sister agency, the USDA & flea beetle numbers and spurge canopy Spurge also stresses information and edu- Plant Health Inspection Service, to create cover reductions in the range 40-95%. In cation. Although a variety of tools (press ‘TEAM Leafy Spurge’ (TLS). addition, grass production and species rich- releases, magazine articles, pamphlets,

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CDs, etc.) are used to increase awareness of The weed also is a prolific seed producer, stock, possessing pyrrolizidine alkaloid the problem and potential solutions, TLS thus contributing to its continued spread levels which are much higher than those has discovered that low tech, grass-roots and invasion into new areas. Cattle and found in another toxic range weed, tansy efforts such as field day events are wildlife generally avoid grazing on Dalma- ragwort (Senecio jacobaea). Normally, extremely valuable in regard to teaching tian toadflax. Cattlemen in BC have listed livestock avoid feeding on green hound- people about biocontrol and IPM. the weed as their third priority for control stongue, but problems may arise once the after the knapweeds (Centaurea spp.) and plant dies back in late summer or fall, or if The programme’s information and educa- houndstongue, because of the loss in it happens to get into hay. tion efforts will be highlighted at grazing potential of toadflax infested lands. ‘Spurgefest II’, set for June 2001. The 3- As with Dalmatian toadflax, options for day event will include a leafy spurge sym- The options for control of Dalmatian toad- houndstongue control are limited, hence, a posium, tours of TLS research and demon- flax are few. Chemical treatment is uneco- biocontrol programme was initiated in stration sites, and demonstrations of flea nomical, difficult because of the plant’s 1988 with European exploration for poten- beetle collection and redistribution tech- deep roots and waxy leaves, and potentially tial agents. The root weevil, Mogulones niques. Its precursor, ‘Spurgefest ’99’, was environmentally damaging when applied to cruciger, was the first agent to be released wildly successful, capturing a considerable large weed stands on grasslands. Further- in Canada after 9 years of host specificity amount of media interest across the more, habitats where toadflax often grows, testing. Since the initial releases in 1997, northern Great Plains and drawing partici- such as coarse-textured soils or near water, the weevil has established at all sites so far pants from across the USA and Canada. restrict the use of effective herbicides (e.g. (i.e. 67 by the end of 1999) and already is picloram). Mechanical control (i.e. hand- showing evidence of reducing hound- For additional information about TEAM pulling or mowing) also has not proven fea- stongue density at the oldest sites. Moni- Leafy Spurge or Spurgefest II, see the TLS sible. Biological control is considered the toring of target and any potential nontarget website at: best long-term control option. impact will be of upcoming emphasis in our http://www.team.ars.usda.gov programme. or email: [email protected] A European stem-boring weevil, Mecinus janthinus, first released in 1991 in Canada By: Rose De Clerck-Floate, By: Steve Merritt, TEAM Leafy Spurge against Dalmatian toadflax, has established Agriculture and Agri-Food Canada, technology transfer specialist, USDA-ARS well and has produced major attack on the Lethbridge Research Centre, Northern Plains Agricultural Research weed in southern BC. Monitoring of sev- 5403 - 1 Avenue South, P.O. Box 3000, Laboratory, 1500 N. Central Ave., Sidney, eral 5- to 6-year-old release sites in 2000 Lethbridge, Alberta, Canada, T1J 4B1 MT 59270, USA revealed 100% attack of toadflax shoots for Email: [email protected] Email: [email protected] at least 50 m around the original release Fax: +1 406 433 5038 points. In some instances, this level of   attack was evident for several hundred metres. Most of the damage is attributed to Azolla Biocontrol in South spring feeding on shoot tips by mass- Weevils’ Success against Africa emerging adults, thus causing significant Canadian Rangeland stunting and a complete loss of flowering Azolla filiculoides or red water fern is a free Weeds on reproductive shoots. The ’s ability floating aquatic fern native to South to cause such impressive damage to toad- America. It was first recorded in South Progress on projects involving the classical flax was not predicted from earlier Euro- Africa in 1948. For many years the fern was biocontrol of two rangeland weeds, Dalma- pean studies. Although it has been too soon confined to small streams and farm dams in tian toadflax (Linaria dalmatica)and to detect actual reductions in toadflax den- the Colesburg area in the centre of the houndstongue (Cynoglossum officinale), sity at most sites, we continue to monitor country. However, the lack of natural ene- was significant in 2000. Both projects permanent plots to document any changes mies and the presence of enriched waters involve success with relatively recent intro- in vegetation. have contributed to its inevitable spread by ductions of European insects, which were man, waterfowl and floods to many sites screened by the CABI Bioscience Centre in Houndstongue around the country. By 1998 the weed had Switzerland with funding from Canadian been recorded at some 176 sites. and US sponsors. Houndstongue is a biennial or short-lived perennial weed of mountainous rangelands The increasing abundance of A. filiculoides Dalmatian Toadflax in northwestern North America. Originally in conservation, agricultural, recreational from Eurasia, the weed is thought to have This perennial with the bright yellow, snap- and suburban areas over the last 10 years is been accidentally introduced in the 1800s dragon-shaped flowers is an invasive weed cause for great concern. Among the major and has since spread considerably. The of grasslands, open forests, roadsides and consequences of the dense mats (5-30 cm weed thrives particularly in forest openings waste areas of western North America. thick) of the weed on still and slow-moving created through logging or mining activi- Originally introduced as an ornamental water bodies in South Africa are: reduced ties, sometimes forming dense monocul- from eastern Europe in the 1900s, Dalma- quality of drinking water caused by bad tures in these habitats. tian toadflax has since become a serious odour, colour and turbidity; increase in problem in Canada, particularly in the In British Columbia houndstongue is a waterborne, water-based and water-related southern interior of British Columbia (BC) major concern to cattlemen because it hin- diseases; increased siltation of rivers and and contiguous areas of southwest Alberta, ders establishment of forage on new pas- dams; reduced water surface for recreation where it currently infests thousands of hec- tures and its barbed seeds or ‘burrs’ attach (fishing, swimming and water-skiing) and tares of range and forest lands. An exten- to cattle, causing irritation, potential reduc- water transport; deterioration of aquatic sive root system and strong, early spring tions in auction price of , and a neg- biodiversity; clogging of irrigation pumps; growth allows Dalmatian toadflax to out- ative impact on the rancher’s reputation. drowning of livestock; and reduced water compete surrounding rangeland vegetation. Houndstongue also is highly toxic to live- flow in irrigation canals. News 3N

In South Africa, no herbicides are registered Animal, Plant & Environmental Sciences, Florida Fast Food for use on red water fern, and this, in con- University of the Witwatersrand, junction with the fact that its rapid rate of Private Bag 3, Wits 2050, South Africa Our ‘Florida Stripper’ [BNI 21(3), 57N increase renders manual and mechanical Email: [email protected] (September 2000)], the melaleuca weevil control ineffective, suggested that biolog-  Oxyops vitiosa, has developed a taste for an ical control was the most suitable option for artificial diet. This does not mean it will be this weed. The frond-feeding weevil Sten- invading hamburger bars, but it does make opelmus rufinasus was collected from Taste of Its Own it much easier to rear. Ultimately, the suc- Azolla caroliniana in Florida (USA) and Medicine? cess of a biocontrol programme relies on imported into quarantine in South Africa in both having appropriate agents, and being Search for Rhinocyllus conicus references in late 1995. Following host specificity testing, able to mass rear and release them. This recent literature, and you are likely to come the weevil was released in December 1997. rearing technique includes both a diet and a up with a host of publications discussing the The first release, of 900 weevils, was made pupation medium. Developed by USDA nontarget effects of this introduced weevil on a one-hectare dam in a bird sanctuary in scientists at the ARS Invasive Plant on native thistles in North America. How- Pretoria, which was 100% covered by the Research Laboratory in Fort Lauderdale, it ever, down in New Zealand, it is a different weed. By February 1998 (2 months later) gives a real boost to the programme for bio- story, for there the weevil has itself become the red water fern mat had collapsed, and control of Melaleuca quinquenervia in the subject of nontarget attack. some 30,000 weevils were reared from one Florida. 2 m² sample of decaying material. The braconid Microctonus aethiopoides was introduced to New Zealand in 1982 as The rearing technique has two key features: To date the weevils have been released a biocontrol agent for the weevil Sitona dis- first, it contains the right balance of nutri- (usually in batches of 100 adults) at some coides on lucerne, and has been useful in ents (including sucrose, glucose, maize 110 sites throughout South Africa. The reducing damage in this important crop. starch, vitamins and minerals) and feeding information that we have on these sites is Now, however, it has been found attacking stimulants (M. quinquenervia leaf extracts) that the weevil has been responsible for R. conicus in some agricultural environ- to support the complete development of the clearing 72 of them completely. For the ments. Rhinocyllus conicus was released in insect from egg to adult; second, a substrate remaining 38 sites, either the weed has been New Zealand as a biocontrol agent for nod- is described that provides a relatively high washed away during flooding, or we have ding thistle (Carduus nutans), and in con- rate of pupation. Although larval develop- not revisited them, or it is too early to tell. junction with other agents often provides ment requires nothing more than the right All of the sites that have cleared have done excellent control. The results of prelimi- nutrients, the pupae are more choosy. The so in less than one year. In addition to this, nary studies suggest that M. aethiopoides is substrate that works best included a mixture the weevils have migrated to other sites, likely to have limited impact on R. conicus of sand and water with an absorbent mate- sometimes up to 80 km away from the in the field, but this is not the first instance rial such as crushed florist’s foam or peat release site. We are uncertain if the weevils of this parasitoid exhibiting a catholic taste moss; such a mixture retains enough mois- have been transported on weed by water- for hosts in New Zealand. It has previously ture and allows enough air exchange for fowl, or if there has been short distance hop been recorded from three other weevil spe- successful pupal development. Artificially dispersal onto other dams of the weed, or if cies in the field here (Nonotus albicans and reared weevils have already been released it is as a result of long-range dispersal by two Irenimus spp.) with parasitism rates of at sites infested with melaleuca in south the adults. At around 40% of the sites the 40% or more. Florida and appear to be performing well. weedhasreturnedupto2yearsafterthe initial clearing. Interestingly the weevils The nontarget effects recorded with Micro- Contact: Gregory S. Wheeler, ARS Inva- have located 90% of these and the weed is ctonus aethiopoides contrast with another sive Plant Research laboratory, Ft. Lauder- under control. species of the , M. hyperodae, intro- dale,FL33314,USA duced more recently for control of another Email: [email protected] A thorough cost benefit analysis of this weevil pest of lucerne, Listronotus bon- Fax: +1 954 476 9169 project has been completed and is in the ariensis. This second parasitoid has so far  process of being published; the preliminary proved to be restricted to its target host. The benefit to cost ratio is 1200:1. difference reflects at least in part the increase in awareness of environmental risk There are several interesting aspects to this Mist Flower Flagging? that has occurred in recent years: greater project. Firstly, this appears to be a new rigour in testing procedures meant that the association: although A. filiculoides has Mist flower (Ageratina riparia) spreads at a narrow host range of M. hyperodae was been recorded as native to the southwestern phenomenal rate, and in New Zealand the predicted with a good degree of reliability. USA, this weevil is associated with A. public has been engaged to help to monitor caroliniana in Florida. Secondly the speed Sources: Watch out, Microctonus is about. this. From the results of a questionnaire, with which weevils were able to control Lincoln, New Zealand; Manaaki Whenua – researchers at Landcare Research have even the largest mats in the most eutrophied Landcare Research. What’s new in biolog- drawnupamapofitscurrentdistribution and, by identifying the environmental con- waters. Thirdly, the weevils have been able ical control of weeds, No. 16 (November ditions in which it grows, have also drawn to locate red water fern mats up to 18 2000), p.8. months after the original mats had col- Internet: http://www.landcare.cri.nz up a ‘worst-case’ map showing the areas it lapsed. It remains to be seen if the current Lynch, L.D.; Thomas, M.B. (2000) Non- could potentially invade. Presently most level of control will be sustained over the target effects in the biocontrol of insects common in the north of North Island, it long-term. with insects, nematodes and microbial could spread to other parts of North Island and the top of South Island. However, with agents: the evidence. BNI 21(4), 117N- Contact: Andrew McConnachie, the help of two exotic biocontrol agents 130N. Ecophysiological Studies Research Pro- with a track record of success in Hawaii, its gramme (ESRP),  spread may soon be slowed if not halted. 4N BiocontrolNews and Information 2001 Vol. 22 No. 1

The white smut Entyloma ageratinae was for effective control is well underway [BNI Email: [email protected] the first agent to be released against mist 20(4), 108N-109N (December 1999)]. In Fax: +61 8 93336646 flower in New Zealand [BNI 20(4),107N the last 2 years two defoliating biocontrol  (December 1999)], and now an insect agents have been released against it by agent has also been approved for release by researchers from CSIRO (Commonwealth the Environmental Risk Management Scientific and Industrial Research Organi- Mimosa invisa: a Growing Authority. A shipment of the gall fly sation) and the Cooperative Research Menace in South India Procecidochares alani has been imported Centre (CRC) for Weed Management Sys- Mimosa invisa, a noxious fast growing from Hawaii, and releases are expected to tems. invasive weed of South American origin begin this summer. It is hoped the insect has recently emerged as a new threat to nat- will complement the striking success of the The leafhopper Zygina sp. was released in ural forests, forest plantations and agricul- pathogen. May 1999 across southern Australia. It suc- cessfully oversummered at the majority of tural systems in Kerala State in southwest The smut, which was released in November sites. At two research sites the insect has India. The weed is an annual but wherever 1998, has been spreading even faster than spread 300 m from the release site after 18 water is available year-round it can also its mist flower host. It has been recorded up months. Schools and community groups grow as a biennial. Mimosa invisa has a to 56 kilometres from release sites in the are involved in rearing and release of the scrambling stem bearing four or five rows north of North Island. In the Waitakere insect. It has now been released at over 200 of sharp prickles. Leaves are small, bipin- Ranges, the spread has been so effective sites, and is beginning to impact on the nate and fluorescent green in colour. The that by December 1999 no uninfected areas weed at many sites, where the onset of inflorescence is a condensed spike (capi- could be found for studies into long-term senescence is advanced by several weeks. tate) which is pinkish in colour. The weed vegetation changes. Even more unexpect- Studies are underway to monitor reserve produces a large number of seeds which edly, an intrepid pathologist who ventured accumulation in the tubers. have a long period of viability. It is helio- out to Great Barrier Island to release the phytic in adaptation and cannot grow well smut found it had beaten her there. Leafhopper releases were followed in 2000 under closed canopy. Mimosa invisa is Although it may have arrived on the wind by releases of the rust Puccinia myrsiphylli. moderately drought resistant. The fact that (the nearest release site is on Waiheke There are high hopes for this pathogen, as it it can invade and cover the ground com- Island, 77 km away), Jane Fröhlich sug- causes considerable damage to the creeper pletely, competing with other plants, gests it is equally likely to have been taken in its native South Africa. It was released at smothering herbaceous growth implies there by an unwitting human carrier. 50 sites across all states of southern Aus- habitat degradation and loss of biodiversity. tralia from mid July until the plants began Mimosa invisa growing areas are impene- There are also encouraging signs that the to senesce with the onset of the hot summer trable because of the characteristic thick fungus is causing significant damage to the weather in November. The rust established growth and the stem being armed with weed. At the nine sites where the fungus well at most sites, and the epidemic devel- sharp prickles. It is known to be toxic to was released, 30-50% of the mature leaves oped steadily until the onset of senescence. cattle. last summer were destroyed in the first Spread, though, was slow from the artifi- wave of attack, and 30-90% of the regrowth cially inoculated release areas. At one site, A preliminary survey conducted in Kerala that season met a similar fate. Monitoring the rust spread about 30 m within 4 months indicated that M. invisa is widespread in the will continue this summer. However, there of release, while at another site along the central and southern parts of the State. Of are signs that the defoliation at release sites side of the road it spread as far as 100 m. It the 52 sites surveyed in seven districts in is already allowing the recovery of some caused premature defoliation of the the State, over 50% were heavily infested. native plants, including orchids and ferns, cladodes at the release points at these two In central Kerala, natural moist deciduous that had been choked out by mist flower. sites. The slowness of spread may be forests are heavily infested. In evergreen and semi-evergreen forests infestation is Sources: History sometimes repeats. Lin- related to the microclimate, which is char- seen only on the fringes where canopy is coln, New Zealand; Manaaki Whenua – acterized by very little wind movement partially or fully open. Among forest plan- Landcare Research. Patua te otaota – Weed because the weed tends to grow below tations, teak is seriously affected. The weed clippings. Biological control of weeds other shrubs. However, teliospores (sexual grows luxuriantly on roadsides and fallow annual review 1999/2000, p. 3. spores) have been produced in abundance. lands. Mimosa invisa is found to overgrow Drawing up enemy lines. Lincoln, New This autumn (February/March) researchers and smother Mikania micrantha in most Zealand; Manaaki Whenua – Landcare will be able to see if and how the rust reap- parts of Kerala. It appears that at the present Research. What’s new in biological control pears at the release sites, and field trials rate of growth and spread, M. invisa may of weeds, No. 16 (November 2000), pp.5-6. have been set up to monitor the spread and even exceed Mikania micrantha in posing Internet: http://www.landcare.cri.nz epidemic development of the rust over the next 2 years. serious threat to natural forests, forest plan- Contact: Jane Fröhlich, Landcare Research, tations and agricultural systems in Kerala. Private Bag 92170, Mt Albert, Auckland, Contacts: Louise Morin, CRC for Weed The potential combined impact of these New Zealand Management Systems, CSIRO Ento- two weeds cannot be over-estimated. Email: [email protected] mology, GPO Box 1700, Canberra, ACT Mechanical control of M. invisa by manual Fax: +64 9 849 7093 2601, Australia Email: [email protected] weeding is difficult and labour intensive.  Fax:+61262464362 Moreover, the weed can sprout vigorously from the cut base soon after the onset of Rust for Bridal Creeper Tim Woodburn, CRC for Weed Manage- monsoon. An attempt to control it using 2,4 ment Systems, CSIRO Entomology, D and dinitrobutyl phenol (denoseb) in Bridal creeper (Asparagus asparagoides)is Western Australian Laboratory, Private Brazil was not very successful. According one of Australia’s most damaging and per- Bag 5, PO Wembley, Western Australia to Rachel McFadyen, biological control sistent environmental weeds, and a search 6913, Australia using insect enemies is showing success in News 5N

Queensland (Australia), and attempts to particularly polyphagous, fecund and ellite DNA contains highly repetitive non- control this weed through biological means hardy population of B. tabaci was identi- coding sequences that accumulate muta- are highly warranted in India. fied in the USA in the early 1990s, which tions quickly as compared with other parts proved also to be a vector for a new of the genome. This has the potential to By:K.V.Sankaran, arsenal of viruses. It spread rapidly give better information about geneflow Kerala Forest Research Institute, through the New World, and to Europe, between populations and to distinguish Peechi – 680 653, Thrissur Dist., Australia, Southeast Asia and the Pacific. strains in rapidly evolving systems. Kerala, India It was designated B. tabaci biotype-B Email: sankaran @kfri.org based on esterase profiles (while the B. Conveniently for this morphologically Fax: +91 487 282249 tabaci population native to the south- indistinct group, RAPD banding patterns of  westernUSAandnorthwesternMexico Eretmocerus and species differ. was designated biotype-A). In 1994, bio- These unique patterns were found, by com- type-B was given species status as B. paring them with minute species differ- DNA Fingerprinting: ences in antenna morphology, to corre- Pointing the Way argentifolii on the basis of allozyme and RAPD-PCR (random amplified polymor- spond directly to distinct species within the two genera. In this instance, RAPD is a Finding biocontrol agents, whether for old phic DNA – polymerase chain reaction) useful quarantine tool for making prelimi- or new pests, may superficially seem a analysis, and viral transmission, morpho- nary separations of geographic populations straightforward business, but in practice it logical and mating studies. However, the or strains of Eretmocerus, and possibly new is very complex. Factors that make it so status of the biotype/species has remained species. However, there is not always include (a) incomplete of both contentious, as indeed has that of B. tabaci agreement between traditional taxonomic pests and natural enemies; (b) wide native as a whole. Many studies of many kinds and DNA techniques. RAPD analysis at the distribution of pests; and (c) limited infor- have indicated wide variation in numerous Mission laboratory consistently distin- mation on associations between natural parameters between different B. tabaci guished between several geographically enemies and the target and its related spe- populations, and possibly more than isolated collections of Encarsia sophia (= cies. Unravelling which are the best natural would be expected within many species, E. transvena) that could not be distin- enemies to introduce, and how many spe- yet it has not been possible to establish a guished by morphological methods. cies are needed is a time-consuming and reliable phylogeny. Recent DNA analyses therefore costly process, yet projects are (PCR amplification and sequencing of two DNA sequencing (of the D2 expansion generally severely limited by both. Projects mitochondrial markers (portions of the region of the 28s rDNA gene) of Encarsia also have a habit of coming to an untimely 16S ribosomal subunit and cytochrome species has been carried out by Chris Bab- end: once the agents are out, it can be hard oxidase I genes, COI) and one of the ribos- cock & John Heraty (University of Cali- to persuade funding agencies that their omal internal transcribed spacers (ITS1)) fornia at Riverside, USA). Sequencing (of progress needs to be monitored. for B. tabaci from different locations the ITS 1 and 2, mitochondrial CO1 and D2 around the world have provided the first Molecular biologists have developed a andD3regions)ofEncarsia and molecular evidence of important genetic range of techniques, based on multiplying Eretmocerus parasitoids of B. tabaci and divergence between geographically iso- Trialeurodes vaporariorum in Australia up small variations in DNA to detectable lated populations2,3. The authors argue levels, to allow them to distinguish between has been carried out by Paul De Barro & that it is more realistic to consider B. Felice Driver (CSIRO Entomology, Aus- species, and between populations of the tabaci as a whole as a species complex, for same species. An account of some of these tralia). The aim of these studies is to suc- any alternative would require the descrip- techniques is given in a previous news cessfully characterize several species and tion of a separate species for each unique item1, and more are being developed. They species-groups of these parasitoids, as well population. form a powerful battery of investigative as contribute to phylogeny reconstruction. tools with a lot to contribute if applied in a The taxonomy of whitefly parasitoids pro- Their studies have shown a strong relation- targeted intervention way in combination vides another challenge. Some of the prob- ship between morphological characters and with other biocontrol research methods. lems were outlined by Andrew Polaszek in molecular phylogenetic structure. This Here we look at what kind of information an earlier news article4. A number of teams work is continuing at Imperial College, UK DNA analysis can provide (drawing partic- are now using DNA techniques to clarify (Shahab Manzari, Andrew Polaszek, ularly on work with whiteflies). We see the taxonomy of some groups. RAPD-PCR Robert Belshaw & Donald Quicke), how this can inform decision-making in markers were first developed to help sort focusing especially on the Encarsia inaron biocontrol projects. At this time, we are on out the large number of foreign collections species group. As with the Mission work, a learning curve, and the usefulness and made as part of an interagency whitefly sequence data have already shown the pres- limitations of DNA techniques are still programme (involving the US Department ence of undescribed species in the inaron- becoming apparent. However as they begin of Agriculture, Agricultural Research group which are morphologically to be more widely understood, they will be Service (USDA-ARS) and Animal and extremely difficult to distinguish. used more often and more effectively. Plant Health Inspection Service (USDA- Contacts: Paul De Barro, Project Leader, APHIS), state agencies and university Taxonomic Tool Whitefly Research, CSIRO Entomology, researchers). The USDA-APHIS team 120 Meiers Road, Indooroopilly, Qld 4068, In the last ten years, the whitefly Bemisia based at Mission, Texas has conducted Australia tabaci has become a serious worldwide studies using classical taxonomy backed up Email: [email protected] pest, and one of the foremost whitefly vec- by RAPD-PCR analysis of Encarsia and Fax: +61 7 3214 2885 tors in field and glasshouse crops. It is as a Eretmocerus parasitoids from around the vector that it poses the most severe threat, world5,6. Work is also being conducted by John A. Goolsby, as it takes very few individuals to infect USDA-ARS researchers at Fargo, North ARS-Australian Biological Control Lab, and devastate a field with viral disease Dakota, who are using satellite DNA 120 Meiers Rd., Qld 4068, Australia (many of which are little studied as yet). A sequences as species-specific markers. Sat- Email: [email protected] 6N BiocontrolNews and Information 2001 Vol. 22 No. 1 or Don Vacek & Matt Ciomperlik, APHIS, weary traveller in their time, but probably Morphological analysis indicated the Mission Plant Protection Center, never before a footsore exploratory ento- Spanish whiteflies to be a non-biotype-B, P.O. Box 2140 Mission, mologist. For the search had taken 5 years, and those from Texas to be biotype-B, TX, USA 78573, USA and during this time, although the biolog- while those from Thailand were identified Email: [email protected] ical control efforts had continued in tandem as ‘B. tabaci species complex’. Molecular with the searches, they were hampered by classification (based on sequencing of the Andrew Polaszek, the uncertainties. DNA analysis would not mitochondrial COI gene) also suggested Unit of Parasitoid Systematics, have obviated the need for all the explora- that the Spanish material was a distinct Dept of Biology, tion, nor for climate matching that directed local variant. However, comparison with Imperial College at Silwood Park, Ascot, the searches, nor for the morphometric similar analyses of other B. tabaci popula- Berkshire SL5 7PY, UK studies that finally clarified the taxonomy. tions indicated that it fitted into a clade con- Email: [email protected] But by using DNA techniques in combina- taining reference B-biotypes from several Fax: +44 207 594339 tion with conventional ones, not only might locations, including Texas. Molecular iden- Forensic Biogeography taxonomic uncertainties be resolved more tification of the Thailand whiteflies placed easily, but promising areas to focus on for them in a Far East-Southeast Asian clade. DNA analysis has potential for helping in natural enemy exploration may be pin- The Eretmocerus data from the Australian one of the great conundrums of biogeog- pointed more quickly. study3 suggests that the species of effective raphy and classical biocontrol – where spe- Eretmocerus species from Spain, Pakistan, cies originated and how they have moved, Taking up the Bemisia tabaci story again, the Middle East and Australia are all very naturally or otherwise, around the globe. we can see how. Although, based on spe- closely related. These results can be used in By analysing the genetic variation in popu- cies and natural enemy diversity, the centre combination with other criteria such as cli- lations of a pest in its indigenous range and of origin of B. tabaci was believed to be mate matching to make decisions on prior- comparing them to samples from its intro- either the Indian subcontinent or the Middle ities for surveys and testing of potential duced range, it is often possible to pinpoint East/Africa, there was no indication where agents. where the introduced material originated. biotype-B had sprung from. Host plant Diadegma Dilemma For an introduced pest with a very large affiliations suggested perhaps an origin home range, this is probably a good place to somewhere in the Eastern Hemisphere, and The area of origin of natural enemies may start the search for a biocontrol agent, as this was supported by preliminary DNA be far from apparent. It is not uncommon classical theory tells us that this is where sequence analysis (of the mitochondrial for a natural enemy to appear serendipi- effective co-evolved natural enemies are 16S ribosomal gene). Recent studies using tously, having been introduced to a new most likely to be found. DNA techniques have made a significant area along with the pest. The diamondback contribution to unravelling the convoluted moth (DBM), Plutella xylostella,isthe The area of origin of the target pest is not biogeographical relationships within the most important pest of crucifers world- always easy to pinpoint, particularly for species and of its parasitoids. In two of wide. It is notorious for developing pesti- pandemic species. For example, cypress 2,3 these , putting ITS1, 16S and COI pro- cide resistance and, therefore, biological aphid (Cinara cupressivora) was intro- files into analytical models gave consistent control has been widely and successfully duced to Africa in the late 1980s, with results that distinguished New and Old deployed in highland growing areas in immediate and devastating impact on plan- World populations, and separated Old Asia. The most important parasitoid is an tation and smallholder trees throughout the World populations into a number of groups, introduced ichneumonid wasp of European continent. Classical biological control was including Africa; Sahel and Sahel-like origin, Diadegma semiclausum.Thispara- deemed to have the best chance of pro- regions; Australia; and several distinct sitoid is now the target of a new biocontrol viding an effective long-term solution, but clades in Asia. Biotype-B was found to fit attempt in East Africa. there was a catch. Cypress trees are grown with the ‘Sahel’ group, and this provided all over the world, in subtropical and tem- the first definitive molecular evidence to Earlier efforts by the GTZ/ICIPE (Gesells- perate climates, and exactly where the pest support the hypothesis that biotype-B is an chaft für Technische Zusammenarbeit/ in Africa had come from was a mystery. To introduction into the New World from the International Centre for Insect Physiology add to the confusion, the identity of the spe- Old World. It further suggested an area of and Ecology) IPM Horticulture Project in cies in Africa was unclear (indeed, origi- origin in the eastern Mediterranean/North cooperation with a number of national nally thought to be Cinara cupressi, it was Africa. This fitted with features of its research organizations in eastern and later described as a new species). Begin- ecology, and with observations that only southern Africa – and collections by others ning in 1992, CABI and collaborating sci- populations from the ‘Sahel’ group pro- – have consistently yielded an apparently entists from many countries searched duced the phytotoxic symptoms associated indigenous parasitoid identified as D. semi- exhaustively (and exhaustedly) on cypress with biotype-B in the New World. Further, clausum. However, parasitization rates trees throughout North and Central the molecular data supports the hypothesis were very much lower (<15%) than the ones America, and from the Atlantic coast of that the unique physiological changes reported from Asia. Owing to this, and as western Europe, through Central Europe, induced through the feeding of biotype-B is there is no record of an introduction to this the Mediterranean, North Africa and the a recently evolved trait. region of Africa, doubt always persisted Middle East to Pakistan. Eventually, the about the correctness of the identification. probable source was tracked down to Syria, In another study5, DNA analysis and clas- Now, molecular taxonomic methods are to cypress trees nestling below the ruins of sical taxonomy were demonstrated to be a being employed to solve the problem. The castles built during the Crusades many hun- powerful combination. They helped to African material has recently been declared dreds of years ago, which proved to be the establish the identities and predict the puta- identical with Diadegma mollipla, a potato centre of origin of C. cupressivora tive centres of diversity of both B. tabaci tuber moth (Phthorimaea operculella) (although it is now also found in the hosts and their parasitoids, using material parasitoid7. Preliminary molecular work southern Mediterranean region). Doubtless, from southern Spain, Thailand and Texas (a confirms this separation from D. semi- the castle walls have sheltered many a target site for the biocontrol programme). clausum: material collected from two popu- News 7N lations in Kenya and one each from Ethiopia pathogenic to most populations of the weed the area of origin of the subspecies as Sri and Tanzania can be distinguished from the from India. This illustrates the importance Lanka, surveys have also been carried out D. semiclausum introduced to Taiwan by of using DNA analysis in combination with in India, Vietnam and China on other their mitochondrial DNA. other techniques, in this case with host Ligustrum species, since La Réunion has no testing. The results indicated that the most desirable congenerics and only two other Molecular taxonomy will also be used to pathogenic isolate is a new association, native members of the same family. This help keep track of the establishment and information that would not have been approach should maximize the chances of spread of D. semiclausum and its impact on apparent if either technique had been used finding suitable biocontrol agents in this the local species once an introduction has in isolation. instance. been made. For this purpose, a simple and cheap yet safe method will need to be Contact: Sean Murphy, CABI Bioscience Contact: Richard Milne, developed as thousands of samples from all UK Centre, Silwood Park, Buckhurst Road, Department of Plant Sciences, over the region will have to be screened. Ascot, SL5 7TA, UK University of St Andrews, [Progress in this field with other pest spe- Email: [email protected] St.Andrews,FifeKY169TH,UK cies is described below.] Finally, as the tax- Fax: +44 1491 829123 Email: [email protected] onomy of the genus Diadegma is so Less Testing notoriously difficult, the project will also Needless to say, the Bemisia story has attempt to make a contribution towards the something to contribute here. The molec- The Mikania project illustrates a decision- molecular classification of species. ular analysis of Bemisia hosts described making and potentially time-saving use of above was supplemented with morpholog- the AFLP results. Although the Trinidadian Bernhard Löhr and Barbara Wagener are ical and RAPD-PCR analyses of parasitoids Puccinia spegazziniii isolate was highly requesting fresh material from anywhere in from these host populations. The new pathogenic to most samples (indeed, more the world for inclusion in this study. knowledge gained about the biogeograph- so than to the host from which it was iso- Contact: Bernhard Löhr ical relationships of the host meant that lated), some Assam populations proved or Barbara Wagener, resources could be focused on the Spanish resistant. These were also separated by International Centre of Insect Physiology material as likely to provide the most co- AFLP analysis, and all proved susceptible 5 and Ecology, P.O. Box 30772, evolved parasitoid strains . In particular, to a Mexican isolate of another pathogen, Nairobi, Kenya attention was focused on a strain of Dietelia portoricensis. Traditionally, the Email: [email protected] Eretmocerus mundus. However, although only way of testing for promising host – or [email protected] this strain of E. mundus established suc- natural enemy associations is to conduct cessfully and seems to be spreading, the independent matching of potential natural Qualified Match-Making most successful introduced parasitoid in the enemies for the entire invasive population. It is important not to allow the results of Lower Rio Grande Valley, Texas appears to Given the practical and financial con- DNA analysis alone dictate the path of a be E. hayati from Pakistan. This parasitoid straints of most biocontrol programmes, biocontrol project. AFLP (amplified frag- attacks B. tabaci from the Asia group of such comprehensive testing was and is ment length polymorphism) is proving a populations, which are quite distinct from seldom done, with the result was that aber- particularly useful technique to distinguish those found in the Middle East. In this latter rant populations were and are missed. between populations of the same species, case, climate matching appears to have However, using AFLP analysis, it is now and give a measure of the relatedness of been more important than co-evolution. possible to identify ‘representative’ popula- them. It was used to analyse the relatedness Contact: John Goolsby (details above) tions and ‘hot spots’ of genetic variation. of weed populations in a CABI Bioscience/ or Kim Alan Hoelmer, The outcome of this is that natural enemy Kerala Forest Research Institute (KFRI) USDA, ARS, testing can be done on a more representative project (funded by the UK Department for European Biological Control Laboratory, selection of the pest in its introduced range, International Development, DFID) to Campus International de Baillarguet, and, depending on the genetic homogeneity develop a biocontrol strategy for Mikania CS 90013, Montferrier-sur-Lez, 34988 St. of the introduced material, the extent of the micrantha (mile-a-minute) in India. Gely du Fesc Cedex, France testing necessary may be reduced. In the Mikania micrantha is a Neotropical weed Email: [email protected] case of Mikania, material from Assam with an indigenous range extending from Fax:+33 499 62 30 49 shows variation which testing protocols Mexico to Paraguay. In India, it is an inva- need to take into account, but that from the sive weed of tropical moist forest regions. It Too fine a focus for exploration for natural Western Ghats is more homogenous. is a serious pest of tree crop/agroforestry enemies may be undesirable for other rea- systems in the Western Ghats and is sons. The specificity requirements of the Rationalizing culturing and host testing spreading through Kerala towards Karna- recipient country (which may be affected was seen as a priority by the US whitefly taka. It is also having a severe impact on tea by the pest status of the target organism) programme. This programme was unique production in Assam in northeastern India. may dictate otherwise. For example, if the in the availability of so many species of nat- AFLP analysis indicated the Indian weed to target site is an island with no other indige- ural enemies from intensive directed for- be of Central American origin. Surveys for nous members of the same genus, or the eign exploration, combined with extensive pathogens revealed only mildly pathogenic target pest is causing catastrophic damage, rearing facilities. Exploration for natural species to be present in India, while 29 spe- a less than host-specific natural enemy enemies was carried out by staff of the cies were collected in Brazil, Mexico, might be acceptable to its authorities, and a USDA-ARS European Biological Control Trinidad and Costa Rica, and four were wider search area may be appropriate. The Laboratory in Montpellier (France) considered to have potential as biocontrol former scenario is illustrated by a current together with other collaborating scientists, agents.Ofthese,therustPuccinia spe- CABI Bioscience biocontrol programme at sites around the world selected on the gazzinii was identified as most promising, against the invasive privet species Ligus- basis of climate matching with target areas yet laboratory host testing of 11 strains trum robustum ssp. walkeri in La Réunion. in the USA. USDA-ARS scientists at Mis- indicated one from Trinidad to be the most Although molecular taxonomy confirmed sion, Texas developed a unique quarantine 8N BiocontrolNews and Information 2001 Vol. 22 No. 1 protocol, integrating DNA (RAPD-PCR) necessary. In an Australian study, molec- machinery vs. growth chamber availability) analysis and morphologically-based sys- ular methods were used to help provide a would help determine the most practical tematics, to ensure “the maximum amount structure within which the morphological method for that situation. of species diversity with a minimum systematist can operate. On the basis of this, amount of duplication in cultures”6.This they developed a user-friendly morpholog- So far, the new method identifies two intro- allowedthemtoassessavastarrayofcan- ical key to Encarsia species that removes duced European parasitoids: Peristenus didate parasitoid populations/biotypes on a the need for any molecular analysis. digoneutis, which attacks the tarnished large scale simultaneously. plant bug, Lygus lineolaris, a pest of many DNA techniques can also be used to obtain crops, and P. conradi, which attacks the Foreign collections were categorized in reliable information on the dynamics of par- alfalfa plant bug, Adelphocoris lineolatus, quarantine by plant type, collection site and asitoid populations in the field. The Mission and it can distinguish them from the native macrotaxonomic characters of both hosts laboratory conducted RAPD-PCR analysis P. pallipes. The Newark laboratory intro- and parasitoids. Only parasitoids of ‘the on parasitoid material recovered from unin- duced both European species, and they Bemisia tabaci complex’ were accepted. oculated plots after agent release in Imperial have shown that the wasps are permanently Eretmocerus and Encarsia were separated Valley, California. The most promising established and are spreading in the north- on the basis of pupal and adult female char- agents in this desert climate appeared to be eastern USA. The bugs are pests of crops acters. Individuals from each accession Eretmocerus emiratus, E. hayati, E. mundus grown for seed, vegetables, fruits, cotton were then characterized using RAPD-PCR and an Eretmocerus species from Ethiopia, and seedling trees throughout the USA, and (primers CO4 and A10). Typically, mate- while Encarsia transvena from Pakistan annually cause tens of millions of dollars in rial was characterized by both methods also looked promising. Morphological char- losses and control costs. Research is contin- within three days, and unique accessions acters were used to determine that exotic uing to determine whether other related weresetupinpureculturesonlocalB. Eretmocerus were in the majority (80%) at parasite species can also be identified with tabaci, while duplicates of existing cultures a time of normally peak native population the new technique. DNA probes do not were combined with them to increase their levels. Encarsia populations also peaked, invariably supply all the answers, but pro- genetic diversity. USDA-ARS scientists and DNA analysis of recoveries was used to vide another tool in the taxonomic kit. In used this process to allow them to evaluate show that all E. transvena recoveredinthe this project, determining the phylogenetic multiple species on a large-scale. First, plots were of the newly released Pakistani relationships of the various species of Per- fecundity of candidate agents was assessed strain, replacing the Spanish strain of the istenus is one of the continuing objectives. on hosts on selected crop plants. Promising same species, which was not capable of In cases where clear morphological differ- parasitoids were reared and cage released such population increase during the summer ences between species exist, their relation- onto the same crops to measure parasitism months in this region. ships have been confirmed by DNA under field conditions. All species information. More work on the most Keeping Track approved for release were tested by field closely related species is needed to advance release in an establishment evaluation. Biocontrol of some other key crop pests in further toward the ultimate goal of phyloge- Then the ‘sentinel plant’ technique was the USA is receiving a helping hand from netic reconstruction, a process requiring used to test whether populations became DNA fingerprinting. Cornell University appropriate quantities of fresh material of widely established. In total, 38 exotic and scientists, in collaboration with ARS all species in the genus, and the time to per- 8 two native parasitoids were evaluated . researchers at the Beneficial Insects Intro- form the analyses. Establishing Results duction Research Laboratory in Newark, Delaware have modified an AFLP method Contact: William H. Day, The role of DNA techniques in biocontrol for identifying parasitoid larvae within their ARS Beneficial Insects Introduction programmes is not limited to the pre- pest host9. This method can even detect and Research Laboratory, release phase. Probes are now being devel- identify early stages of parasitoid larvae Newark, DE19713-3814, USA oped to make post-release monitoring of which are difficult to distinguish by con- Email: [email protected] natural enemies more effective. However, ventional means. A similar procedure was Fax: +1 302 737 6780 morphological analyses continue to pro- developed and used for blackflies Kelley Tilmon, vide the framework within which the more (Simulium spp.) infected with Onchocerca Entomology Department, expensive and time-consuming molecular in the 1980s, and this was an important Cornell University, Ithaca, methods can be targeted. development for assessing the prevalence NY 14853 USA of river blindness. Email: [email protected] DNA analysis has been used in the Bemisia Fax: +1 607 255 0939 programme to help evaluate the success of At a stroke, this does away with some the released agents and conclusively iden- lengthy and painstaking monitoring proce- The technique is not limited to insects, for tify field recoveries of cryptic species/pop- dures. Once adult parasitoids have been ARS scientists elsewhere have developed ulations in Texas, desert valleys of Arizona reared in the laboratory and identified to similar methods to monitor different strains and California, and the San Joaquin Valley, species (a process which can take months), of weed pathogens following their release California. For example, species-specific DNA probes can be developed. Then there into the environment as biocontrol agents. probes developed from satellite DNA were is no longer any need to go through the Methods are emerging for detecting and used to identify Eretmocerus species from lengthy ‘rearing out’ of parasitoids before identifying several isolates of Myrothecium material collected post-release5. However, they can be identified. Estimating per- verucarria, a soil fungus that kills morning the cost of RAPD analysis is the limiting centage parasitism no longer requires the glories, which plague sugarcane and other factor, and to overcome this and time limi- skilled dissection of large numbers of field- crops. In field studies, spraying redroot- tations, research has been initiated to collected hosts. However, it is important to and smallflower-morning glories with an develop specific DNA probes from the sat- note that the DNA evaluations also require oil-based carrier containing Myrothecium ellite DNA for development into a ‘squash special equipment, skill, and time; the spores proved as lethal to these weeds as blot’ kit. Such refinements are not always resources of a given laboratory (DNA the herbicide atrazine. News 9N

The DNA fingerprinting technique will Contact: Dave Moore, CABI Bioscience To return for a final time to the USDA help biocontrol scientists keep close tabs on UK Centre, Bakeham Lane, Egham, Bemisia biocontrol programme, what the spore growth and spread, host range and TW20 9TY makes it particularly striking is that DNA effectiveness of different strains of biocon- Email: [email protected] techniques were used at all stages of the trol pathogens such as Myrothecium fol- Fax: +44 1491 829100 programme, to inform decisions, to lowing release. In this way, it can give increase efficiency, and to contribute to the genetic evidence linking a specific micro- Although there are too many strategic scientific basis. As a result of introductions bial release to a specific disease seen in applications of DNA analysis to cover here, of Eretmocerus mundus and E. hayati,sen- target weeds. It also reveals the spread of we include one that has particular signifi- tinel plant sampling indicated a dramatic biocontrol microbes and demonstrates their cance for disease quarantine and manage- increase in the numbers of introduced effectiveness in reducing invasive weed ment. In Canada, probes are being Eretmocerus spp. Before releases of these, populations. developed (US patent #5792611) to iden- native E. tejanus formed more than 95% of tify tree root rots and scleroderris canker in Contact: Jan Suszkiw, USDA – ARS, recoveries, yet within three months, exotic the absence of symptoms. One problem populations had risen to 85% of the Information Staff, with such diseases is that they may remain 5601 Sunnyside Avenue, Mailstop 5129, Eretmocerus spp. recovered. Evaluations latent and asymptomatic for long periods, of their impact on B. tabaci populations are Beltsville, MD20705, USA during which time they are hard to detect Email: [email protected] now underway, and are expected to confirm but can still spread. Often, by the time that they are making a significant contribu- Fax: +1 301 504 1641 symptoms are visible, it is too late to 6 Douglas G. Luster, tion to biocontrol . The authors sum up manage the disease. The DNA probes will “We hope these results will encourage USDA – ARS Foreign Disease – have a two-fold application: first, to ensure Weed Science Research Unit, other biological control programs to the health of nursery seedlings before develop predictive methods and test their 1301 Ditto Ave, Ft. Detrick, planting, which will help to prevent the MD 21702-5023, USA predictions in field settings. The informa- spread of established diseases; second, to tion gathered… might further the theoret- Email: [email protected] monitor imported stock for diseases, which Fax: +1 301 619 2880 ical aspects of our science and in turn will be a useful quarantine tool, enabling increase the likelihood of success in future Strategic Designs forest pathogens to be detected and diag- biological control programs”. nosed at the point of entry and thus prevent Programmes in future may be able to use their introduction. Sources information from DNA analysis in formu- 1Reid, A.; Murphy, S. (1999) Biogeo- Contact: Richard Hamelin, lating biocontrol and IPM management graphic fine-tuning helps weed biocontrol. strategies. One project that intends to capi- Natural Resources Canada, BNI 20(2), 51N-54N. talize on this is aimed at developing an inte- Laurentian Forestry Center, grated strategy for management of the 1055 rue du P.E.P.S., 2 Frohlich, D.R.; Torres-Jerez, I.; Bedford, coconut mite, Aceria guerreronis. This Ste-Foy, Quebec, Canada G1V 4C7 I.D.; Markham, P.G.; Brown, J.K. (1999) pest, which is of undetermined origin, Email: [email protected] A phylogeographical analysis of the causes significant problems for coconut Fax: +1 418 648 5849 Bemisia tabaci species complex based on growers around the world; crop losses of mitochondrial markers. Molecular Ecology Picking Winners more than 30% have been reported in the 8, 1683-1691. Caribbean. It has long been a problem in Finally, in a world where funding is hard to 3De Barro, P.J.; Driver, F.; Trueman, the Americas, from where it was first come by, being able to convince a potential J.W.H.; Curran, J. (2000) Phylogenetic recorded, and more recently from Africa donor that you have a success story waiting relationship of world populations of (while coconuts are probably indigenous to to happen could be what clinches the deal. Bemisia tabaci (Gennadius) using ribos- Melanesia). The mite is unlikely to have Molecular techniques can sometimes iden- omal ITS1. Molecular Phylogenetics and been transported on coconut as it is not tify in advance certain situations where Evolution 16, 29-36. found on the mature nut, which is both the classical biocontrol is likely to be suc- 4 natural mode of dispersal and the form cessful. Where there is little variation Polaszek, A. (1999) Identification of transported by humans. It is therefore within a population, host-specific natural whitefly parasitoids: some advice. BNI assumed that the mite’s original host enemies are less likely to be challenged by 20(4), 117N-119N. belongs to the indigenous flora of the resistant varieties and therefore stand more 5Kirk, A.A.; Lacey, L.A.; Brown, J.K; Americas. However, outbreaks in Asia (Sri chance of succeeding. For example, plant Ciomperlik, M.A.; Goolsby, J.A.; Vacek, Lanka) have only been reported recently, cytologists at Leicester University proved D.C.; Wendel, L.E.; Napompeth, B. (2000) and it is unclear whether these have fol- that all the invasive Japanese knotweed Variation in the Bemisia tabaci s.l. species lowed a recent introduction, or indicate the (Fallopia japonica) in the UK (and prob- complex (Hemiptera: Aleyrodidae) and its breakdown of natural population regula- ably USA and Europe) appears to be same natural enemies leading to successful bio- tion; outbreaks tend to occur in pockets male-sterile clone, which would make it logical control of Bemisia biotype B in the separated from each other by long dis- extremely susceptible to biological control USA. Bulletin of Entomological Research tances. This issue could be addressed by – as well as providing the press with oppor- 90, 317-327. comparing DNA samples from populations tunities for headlines such as ‘Largest 6 in the Caribbean and Africa with multiple female on earth set to swamp Britain’! Goolsby, J.A.; Ciomperlik, M.A.; Kirk, samples from Sri Lanka. If the origin of the A.A.; Jones, W.A.; Legaspi, B.C., Jr.; mite in Sri Lanka can be ascertained, an Contact: Richard Shaw, CABI UK Centre, Legaspi, J.C.; Ruiz, R.A.; Vacek, D.C.; IPM strategy can be designed specifically Silwood Park, Buckhurst Road, Wendel, L.E. (2000) Predictive and empir- either to deal with a newly introduced pest, Ascot, SL5 7TA, UK ical evaluation for parasitoids of Bemisia or to identify and correct whatever has Email: [email protected] tabaci (biotype ‘B’) based on morpholog- caused the upsurge. Fax: +44 1491 829123 ical and molecular systematics. In: Austin, 10N BiocontrolNews and Information 2001 Vol. 22 No. 1

A; Dowton, M. (eds) : evolu- tera: Plutellidae). Bulletin of Entomological 9Tilmon, K.J; Danforth, B.N.; Day, W.H.; tion, biodiversity and biological control. Research 90, 375-389. Hoffmann, M.P. (2000) Determining para- Canberra, Australia; CSIRO Publishing, sitoid species composition in a host popula- pp. 347-358. 8Goolsby, J.A.; Ciomperlik, M.A.; tion: a molecular approach. Annals of the Entomological Society of America 93,640- 7Azidah, A.A.; Fitton, M.G.; Quicke, Legaspi, B.C., Jr.; Legaspi, J.C. ; Wendel, 647. D.L.J. (2000) Identification of Diadegma L.E. (1998) Laboratory and field evaluation species (Hymenoptera: Ichneumonidae, of exotic parasitoids of Bemisia tabaci (bio- Campopleginae) attacking the diamond- type ‘B’) in the Lower Rio Grande Valley back moth, Plutella xylostella (Lepidop- of Texas. Biological Control 12, 27-135. 

IPM Systems

This section traditionally deals with IPM argues, because they are designed to criticized both high-input screening and techniques compatible with biocontrol, appease the public rather than deal with the heavy-handed exclusion policies. They particularly in agriculture. As a diversion problem, and instead they tend to burden argued that the former has no measurable this quarter, we consider the community the child. He points to exclusion from impact on infestation levels (although there health sector, and how action at community school of allegedly infested children and are as yet no reliable data for these). They level is bringing about reductions in pesti- mass screening as pointless measures. He also pointed out that exclusion policies do cide use for head lice control. argues that quarantining has not been little to reduce spread in schools, and only shown to be effective (by the time head lice serve to increase the stigma of head lice Bringing Head Lice are detected the child may have had them infestations. for a month or more), and that more than Control up to Scratch The UK health authorities used to cover the half of the children identified as infested cost of routine screening of schoolchildren Head lice (Pediculus capitis)arenota may be misdiagnosed. for head lice, but discontinued this practice danger to health; they neither serve as vec- Exclusion of children from school because in the early 1980s declaring it to be largely tors nor do they generally cause direct harm they have ‘nits’ is a widespread practice in ineffective. Failure to establish an alterna- or disease. Yet their management is a head- the USA, although no distinction is made tive management strategy, though, has led ache for public health workers worldwide. between live eggs and inviable/hatched parents in the UK to blame a perceived Here, we look at why they are perceived as eggs. Such exclusion policies are discour- increase in head lice incidence in schools such a problem, reasons for the limits to the aged by the American Academy of Pediat- on the withdrawal of school nurse inspec- success of current control measures, and rics and the (US) National Association of tions. Ian Burgess notes that in most dis- how the limits are being addressed. School Nurses. Pollack argues that most tricts most children had been screened less Head lice are wingless, parasitic insects reported outbreaks of head lice in schools than once a year, if that. The little data col- adaptedtolivingsolelyontheheadhairof are no such thing: most ‘infestations’ he lected after inspections stopped suggested their human hosts. The claws on the ends of examines show no evidence of current (or that prevalence dropped, if anything, after their legs are adapted for grasping hair past) infestation, normal debris on the hair screening ceased! Although Belgian health shafts. They feed exclusively on blood, or scalp frequently being mistaken for lice workers have demonstrated that inspection taking several blood meals a day, and die of or eggs. In schools reporting more than based on finding nits gives a high level of dehydration and starvation within 2 days if 50% prevalence amongst its pupils, Pollack false positives, they and the UK nonprofit removed from the head. Head lice do not found actual prevalence rarely exceeded organization Community Hygiene Concern jump, fly or swim but they can move 1%, and sometimes he identified not a (CHC) both found that visual inspection swiftly along hair shafts. Transfer to a new single case. A study of specimens associ- also misses a large number of light, but con- host is almost exclusively by direct contact ated with diagnosis of infestation submitted tagious cases. In the absence of firm current with infested hair. Nymphal stages tend to by health care workers and lay people data the Stafford Group suggest that preva- stay on the head where they hatch, while found that less than 50% were head lice lence has not increased, only public aware- adults (particularly males) tend to migrate stages. Ian Burgess (Cambridge Medical ness – or perhaps over-awareness? where possible to maximize out-crossing. Entomology Centre, UK) says that diag- Conditions Right for Combing nosis of real infections has been consist- Why Scalps Crawl ently poor, and often based on paranoia and In Richard Pollack’s view, there is no con- Head lice are rather less contagious than the misunderstanding. He cites a survey of ceivable need to apply insecticides for head common cold and far less serious in terms school nurse managers attending the human lice within schools (and neither are such of health. Yet although revulsion to them louse control management courses in the ancillary measures as cutting hair, bagging may be groundless it has been the basis for UK in 1982. It found that 50% of the health clothes and not using shared protective most management strategies. It is particu- professionals interviewed (all of whom had sports equipment justified; indeed, the larly in schools, where reports of waves of been involved in screening for head lice in latter carries far greater risk than a head infestations are commonplace, that most schools) had never seen a live louse, and louse infestation.) However, many are control efforts are focused – often need- relied on the presence of nits to confirm an uncomfortable with the notion of parasites lessly and unsuccessfully in the view of infestation. A report by the ‘Stafford they can actually see or feel crawling about Richard Pollack (Harvard School of Public Group’ for the Consultants in Communi- on their heads or their children’s, so even if Health, USA). Control measures fail, he cable Disease Control (CCDCs) in the UK control cannot be justified on health News 11N grounds, there is still a demand for it. In In the USA, a different detection policy has Less contentious is the adverse impact of some cases, too, infestations are so chronic been developed for determining whether an poor application of products, which is often and severe that treatment is justified on infestation is active or a treatment is based on poor understanding of the life health grounds. But what or when to treat is working. The America Head Lice Informa- cycle of the head louse. Instructions may not obvious to the non-expert. Many health tion Center (AHLIC) educate health profes- not be followed precisely. Too little may be authorities and other extension services in sionals and parents that eggs are firmly used (products are perceived as expensive), the USA, the UK and elsewhere dissemi- attached at an angle to one side of the hair and/or the treatment may not be followed nate information by various means (leaf- shaft, and cannot be removed by blowing or up where users believe that all the head lice lets, videos, websites, etc.), but quality is flicking. Once nits have been confirmed to have been killed by the first treatment. variable and sometimes only increases con- be present, an active infestation is diag- Another ‘wave’ of head lice appearing fusion. Clear guidance is needed. nosed by a two-step process: all eggs are some weeks post-treatment is often blamed removed, then the hair is inspected to see if on a ‘new’ infestation (acquired from The first step is finding out whether or not new ones appear. another person) rather than the more likely someone is infested. There are strong advo- explanation of a resurgence from surviving cates for inspecting dry and wet hair, and No Certain Solutions eggs and lice, or misdiagnosis (with only the methods may be appropriate in different dead or hatched eggs present). Joan Sawyer situations. Both, though, rely on combing The question of quite how to kill the head says that AHLIC call it ‘the myth of rein- with a fine-toothed comb to locate any lice lice presents the next dilemma. The con- festation’. Misunderstandings about insec- or nits. Recent advances in this technique ventional approach is a synthetic insecti- ticide action are also reflected in attempts to have been due to the work of CHC, who in cidal application or series of applications. A apply prophylactic treatments. 1988 set up a community development pro- variety of products is registered in different When synthetic insecticides appear to fail, gramme, Bug Busting, to help parents of countries, commonly pyrethroids and people may turn to other treatments, and schoolchildren in the UK acquire head malathion but others include carbaryl or increasingly to ‘alternative’ remedies in the louse detection skills. They had already lindane. There are two problems with such misguided belief that because they are ‘nat- established the value of combing wet hair treatments. The first is the growing antip- ural’ they are inevitably safe. Yet amongst with a plastic fine-toothed comb for this athy of the public to synthetic pesticide use, various herbal remedies widely touted are purpose. Wet detection combing is now the with children a particular concern; the ingredients such as rosemary oil, which is most widely-recommended method in the second is the perception that the insecticide known to induce abortion. Even products UK for confirming presence or absence of treatments ‘don’t work’. Together these generally considered safe for some uses on active head lice. The CHC protocol com- have led to disenchantment. There are humans, such as tea tree oil and lavender, prises methodical combing of washed, still- reports of side-effects for all the synthetic may be applied at doses well in excess of wet, well-conditioned hair. The moisture insecticides currently registered, although those recognised as safe doses on children’s temporarily immobilizes lice, which can be proponents argue that the doses used in heads. Research on essential oils suggests comfortably combed out and then head lice products are so low as to be rea- that they act as contact nerve poisons in the inspected. Emphasis is also placed on sonably safe when used properly. same way as synthetic insecticides. The tracing infested contacts. CHC have con- efficacy of both synthetic and natural insec- There is evidence from a number of coun- ducted a series of case studies that have ticides relies in part on their lipid soluble tries for the development of resistance to demonstrated the value of wet Bug Busting characteristics, as this facilitates their entry the most commonly used pesticides, the to detect low-level infestation, and thus rec- into the body (through either insect cuticle pyrethroids and malathion, and Ian Burgess ommend it for parents anxious to know if a or human skin). Children are more sensitive says that places they have surveyed in the child is infested. However, Ian Burgess than adults to the toxic effects of pesticides UKinthepast6yearshavedoubleoreven argues that dry combing is equally accurate because they have incompletely-developed treble resistance. Resistance is likely to and is routinely practised by the Cambridge detoxification mechanisms and are growing increase in incidence and intensity with Medical Entomology Centre. The method rapidly. The active ingredients in most nat- time, but how often do treatments really has been used in several trials. He points ural remedies are monoterpenes, which as a fail? One common misconception at least out it is more practical (particularly where class are widespread in plants and are also partly to blame for perceived failure is the large numbers of heads are examined) found in some insects, and there is every expectation that an insecticide treatment because it is quicker and less messy. reason to suppose that they will also be sub- provides an instant fix. This is reinforced ject to resistance development in time. For For either method, generally no indication by labels on many products proclaiming example, there has been interest in coconut is given of how long should be spent on that they kill both lice and eggs in a single oil in the USA, and it may have efficacy as combing, and without adequate guidance, application, which most do not, and cer- a treatment against head lice populations detection combing by inexpert hands can tainly not reliably. Eggs are resistant to there. However, this effect may not be sus- lead to both false negatives and false posi- many of the registered products. A second tainable, and may not be mirrored in coun- tives. Richard Pollack advises using the application (recommended in the UK by tries where coconut oil is commonly presence of live lice or viable eggs as the the Stafford Group to be made on day 8) is present in hair products already. The use of sole basis to confirm an infestation, but intended to mop up the lice that hatch after sublethal levels of monoterpene com- notes that recognition of eggs as viable day 1. Some of the products provide a lim- pounds (as are likely to be present, for requires suitable magnification and some ited residual effect, designed to kill example, in tea tree oil shampoo) will training. Moving objects maybe also erro- hatching nymphs. Egg survival may be the enhance resistance development. neously identified as head lice. CHC sug- first sign of resistance, but it is unclear how gest that if parents are in doubt, ‘combings’ far widespread treatment failure is due to The lack of certainty about the efficacy and can be taken to a professional for identifica- this. However, the residual action of current toxic effects of synthetic and natural insecti- tion, but say that most parents, given the products, so hatching nymphs are exposed cide treatments is creating a vacuum, but the right information and combs, can identify to lessening levels of insecticide, is likely to challengeistocomeupwithaneffective them without this recourse. enhance resistance development. alternative. 12N BiocontrolNews and Information 2001 Vol. 22 No. 1

Look to Our Roots Ogg says that a major problem with head ommended protocol. Others may become lice in the local community was exacer- over-keen on bug busting and family con- Grooming and ‘nit picking’ are practised by bated by health professionals giving dif- flict can ensue. All these are issues which our ape relatives, and were once part of ferent recommendations on control to proponents of the method recognise and human hygiene, a routine task undertaken parents, who became confused. A head lice have sought to deal with. CHC have specif- by older siblings or parents (and still task force to formulate a policy was formed ically developed methods for combing dif- remembered by some of us, although not that included health professionals (school ferent hair types and instructions for these necessarily fondly). Perhaps it is a casualty nurses, public health nurses, university are included in their Bug Busting kit. Joan of modern life, perhaps the advent of insec- educators), the state medical entomologist Sawyer of AHLIC argues that far from ticide treatments led to its demise, but since and child care programme administrators resenting the time that combing takes, par- 1995 its metamorphosis into Bug Busting (from the health department). The policy ents and health professionals are increas- as a remedy has occurred. CHC developed was announced before the beginning of the ingly aware of the even greater cost in time wet detection combing into a treatment, school year in autumn 1999. Parents were and money of relying on poor information which depends on systematic removal of urged to check their children’s hair before and ineffective or dangerous pesticides. head lice to break the life cycle. The Bug school started and school nurses inspected Busting method follows a prescribed pro- children during the first week of term, and The success of a bug busting programme tocol, twice weekly for two weeks. It is again mid-term and after the Christmas maydependonhoweffectivelythemes- based on three tenets: break (January 2000). School nurses agreed sage is disseminated through the commu- • ‘coated’ lice move more slowly and so that they would not recommend products or nity and, in the long term, on the degree to are more easily removed by combing treatment methods not proven to be effec- which the momentum can be maintained tive against head lice; this was a major once the initial wave of enthusiasm has • removing lice before they mature pre- achievement because there were many subsided. Recognizing this, CHC resource vents them spreading and reproducing advocates for mayonnaise and other home designated Bug Busting days on 31 January • co-ordinated community action helps remedies. The University of Nebraska and 31 October each year in the UK. They decrease the rate of reinfestation Cooperative Extension provided a video- say that success relies to a large extent on tape and fact sheets that were sent home the understanding by schools of their role in CHC produced a Bug Buster kit containing with children. generating coordinated, informed effort full instructions and the necessary comb, followed by a top-up each Bug Busting whichwaspilotedin1995andcommercial- The focus of the recommendations pro- day. Getting the message across would be ized in 1996. An improved model with a motes combing with a long-toothed metal helped by including head lice in the pri- modified comb became available early in comb over all other activities as the ‘alter- mary/first school science curriculum (in the 1999. Currently, the London School of native’ method of control to remove nits. UK, the national science curriculum at this Hygiene and Tropical Medicine (LSHTM) Where time is limited, people are encour- level currently does not include insects with is conducting a randomized controlled trial aged to nit comb above all else. Barbara incomplete metamorphosis – a no-nits comparing the effectiveness of the Bug Oggsaysthatthisreallywasacommunity policy with a difference). Buster kit with insecticidal medication. collaboration and worked well over a trial period. Head lice have not been eliminated, Although some good short-term results A number of organizations in the USA, but (and arguably more importantly) the have been reported, success levels vary. including the National Pediculosis Associ- school system is now seen to have a more How far, too, is initial success simply a ation (NPA) and the American Head Lice proactive policy and to be taking the neces- reflection of the focus on controlling head Information Center (AHLIC) promote sary steps to minimize the problem. An lice rather than the method employed? combing together with other measures for acceptable level of success was being main- Until recently, there have been few head lice control. However, the emphasis tained one year on: in the year 1999-2000, attempts to quantify the success of the US head lice policies placed on children head lice cases had dropped to 2000 from a method in comparison with other (insecti- being nit (rather than active lice) free means pre-programme level of 6700 (presumably cide) treatments. The mechanisms of long- that mechanical methods to resolve infesta- based on the presence or absence of nits). term sustainability of the CHC programme tion have taken a different path from those This programme set itself realistic goals, are currently undergoing evaluation in in the UK, with an emphasis on nit as well and undertook active education at the com- addition to being included in the 3-year as active lice removal. AHLIC have pro- munity level to achieve these. full-scale evaluation of head lice treatments duced a book and a video, ‘Head Lice to by the London School of Hygiene and Dead Lice’, which uses humour to get Testing Time Tropical Medicine begun in 1999. Other across the message. The programme is Advocates consider that bug busting offers studies comparing insecticide treatments based on a five-step battle plan, and an affordable head louse remedy, which is and bug busting in the UK have recently includes the optional use of (a pyrethroid) re-usable, does not require insecticides, published results. These varied in design; insecticide. Instructions are given on the clears lice that are resistant to insecticides, for example, in using professionals to apply correct technique for smothering hair with andpromotesselfrelianceinheadlouse treatments, or working through parents, and olive oil (which is left on overnight under a control. Detractors have some reservations the results are far from clear cut. However, rubber cap to smother the lice by blocking about details of current protocols, but more a trial in Bristol which looked at insecticide the spiracles), the correct days and method significantly consider that the bug busting resistance found 87% and 64% failure for for combing to remove nits and lice, and method is simply too time-consuming and permethrin (pyrethroid) and malathion, advice on nit picking on dry hair. demanding in commitment to be of value at respectively. A randomized controlled trial A modified ‘no nit’ policy was included in a community level. They also point to in North Wales, which compared the effi- a programme encouraging combing as a social constraints: some children’s hair is cacy of two malathion treatments with method of head lice removal that was not readily combed, and bug busting can CHC Bug Busting (using the 1996 comb) developed by the extension service of the cause pain to them. Some parents are found 78% and 38% success, respectively. University of Nebraska at Lincoln. Barbara unable or unwilling to comply with the rec- Arguments over the trials and their inter- News 13N pretation have focused not only on trial that covered all possible errors, worked duced by the programmes discussed in this designs and interpretation of results, but consistently for people unskilled at nit- article. CHC point out that the time for bug also on whether it is justifiable to recom- picking, and did not overtax the family. The busting depends on hair type and length but mend a head lice management strategy criterion for success that they used to eval- agree that poor explanation of how to comb based on either method. However, CHC are uate the protocol during the research phase by some information sources contributes to encouraged by the North Wales results, was that the entire family should be louse failure, which is then used to discredit bug which showed 38% success using the more and nit free 10 days after completion of the busting as a whole. CHC has produced a laborious 1996 comb without initial dem- programme. The CHC protocol includes a demonstration video, recommended by onstration, or the back-up they provide. follow-up Bug Busting 4 days after the last many UK health authorities, to introduce They point out that skill at Bug Busting treatment; if live lice are found, treatments Bug Busting. In this and printed resources, grows with familiarity, while lice resistance are repeated. The UK Department of Health they demonstrate clearly how to go about to insecticides increases with continued advises a wet combing check 3-5 days after detection and treatment combing for dif- use. A knowledge of Bug Busting, they using an insecticidal product. But even if a ferent hair types and lengths. argue, is empowering to a community, protocol seems foolproof, it won’t be: there Finally there is disagreement over the while it does not benefit from dependence will be a few who will not follow it strictly optimal comb design, and this is perpetrated on failing insecticides. for whatever reason, and a few will make in many leaflets, websites and other infor- mistakes. An important message to get mation products which do not clearly dis- Teasing Out the Snags across, especially in societies that expect tinguish between detection and treatment sure-fire solutions, is that bug busting is not What, then, are the current constraints to combs, or lice and nit combs. CHC raised a guaranteed fix, but it is a skill which bug busting and what are the keys to making funding to research the issue. It built on the improves with practice. AHLIC’s experi- it work? Areas to consider include: (a) com- results of field tests made in the first half of ence is that people are willing to admit mis- municating the message; (b) length of treat- the 20th century to select the precise slant takes and try again. ment; (c) hair lubricants and how to use on the leading edge of the new plastic Bug them; and (d) combs and how to use them. There are some robust disagreements about Buster comb that most effectively removed even newly hatched lice close to the scalp. According to Ian Burgess, Bug Busting is a hair lubricants and their use. Olive oil is Burgess agrees that a blunt-tipped plastic well-known and often-tried method, which favoured by Ian Burgess and Joan Sawyer, comb with appropriately and evenly spaced can and should work. When it fails, it does as it is fairly easily emulsifiable (and there- teeth works best for active lice removal. so for a number of reasons, probably most fore washed out afterwards), non-irritant Both CHC and Burgess say that the use of importantly, he says, because many users do and emollient (and so can be used on suf- inappropriate combs is a widespread reason not understand why they are doing what ferers with eczema and psoriasis). Burgess for failure. In the USA, metal toothed nit they are doing. They follow the method, as notes that it is available in the UK as a phar- combs are more popular, and these work they understand it, and are surprised and maceutical preparation. Sawyer describes it particularly well for removing nits. disappointed if it fails. Flawed information as “cheap, healthy and healing, with the Apparent discrepancies in advice may well from unreliable sources and word-of-mouth only side-effect of gorgeous hair”! How- be a reflection of the UK focus on breaking are at least partly to blame, but in order to ever, hair conditioner has the edge as a the lice cycle by removing hatched lice but get the message across it may be over-sim- familiar and attractive product to use, not worrying about nits, and the US focus plified. On the other hand, CHC maintain which CHC say makes it more immediately on removing nits as well as lice so children that more than 80% of people they deal with acceptable. They suggest that even the most can return to school. are able to understand and follow their stout-hearted parent might be put off by the instructions, and achieve good results. Clear thoughtoftryingtocontrolandcombsmall Putting Heads Together consistent information is crucial to this, and slippery children dripping oil all over their technical backstopping, as provided by their soft furnishings. Although there are disa- In conclusion, IPM of head lice may be Bug Busting days, is important for rein- greements over which is the more effective struggling a little at the moment, but there is forcing the message and contributes to the for allowing lice/nits to be extracted, and an urgent need for it to succeed. Ian Bur- sustainability of the approach. also over safety (and lack of testing for this gess argues that failure in head lice man- use), all parties report good success rates. agement begins at the primary level: Most protocols (including CHC’s) involve stakeholders from health professionals at combing twice-weekly for 2 weeks; a total Protocols rarely suggest how long combing all levels through public and community of four or at most five treatments. AHLIC is should take either for detection or for treat- workers of all description to affected indi- rare in advising a longer treatment period of ment, although the University of Nebraska- viduals and families would rather not know 3 weeks, with six combings over this Lincoln programme stresses that nit about head lice, or have to deal with them. period. The more common 2-week period combing should take precedence over all In addition, conflicting and confusing to remove live lice makes the method seem other anti-louse activities. Burgess suggests advice on how to deal with head lice is manageable, and CHC and AHLIC concur that detection combing may take 30 min- widespread. Barbara Ogg says that the suc- that this is an essential consideration. It is utes, treatment combing several hours. cess of the University of Nebraska-Lincoln not much point developing a protocol that is Given normal family pressures, this time is programme was rooted in forming a task too onerous for people even to attempt; but likelytobehardtofind,anditisunder- force so all ‘experts’ spoke with one voice, equally, advocate a protocol that frequently standable that bug busting programmes do a view firmly supported by John Simpson, fails and you quickly lose credibility. Con- not set out draconian regimes that will chairman of the ‘Stafford Group’. Clear, sideration of the head louse life cycle indi- probably be ignored, if not put people off consistent high-quality information based, cates that two weeks does not allow much altogether. However, if combing is not per- as CHC emphasize, on sound science and of a margin for error: miss a few lice, and formed adequately, the method is bound to practical approaches, is crucial. Parents you will soon be back to square one. fail. Successful combing is a technique need to be aware of the likely reasons if AHLIC say that in devising their strategy which needs to be demonstrated, and this is control fails, so they can consider how to they were determined to create a protocol addressed in information products pro- revise their practice to optimize the chances 14N BiocontrolNews and Information 2001 Vol. 22 No. 1 of success the next time. If this option is not Contacts/information: Dept. Immunology & Infectious Diseases, available, they are more likely to abandon American Head Lice Information Center Harvard School of Public Health, bug busting for some other remedy. Email: [email protected] 665 Huntington Avenue, Boston, Website: http://www.headliceinfo.com MA 02115-6021, USA The clear message coming through is that Email: [email protected] Ian Burgess, Medical Entomology Centre, combing for head lice management can Website: http://www.hsph.harvard.edu/ Cambridge Road, Fulbourn, and does work under a variety of protocols, headlice.html and this is very positive. Its sustainability Cambridge CB1 5EL, UK at the community level is being assessed. Email: [email protected] University of Nebraska-Lincoln However, the approach taken by agricul- Fax: +44 1223 882690 Website: http://www.ianr.unl.edu/ianr/ tural IPM of integrating a variety of Community Hygiene Concern (CHC), lanco/nviro/pest/lice.htm approaches to help overcome the short- 160 Inderwick Road, London N8 9JT, UK John Simpson, comings of individual components has Email: [email protected] Public Health Laboratory Service, limited application currently. Whether new Fax:+442082927208 Communicable Disease Surveillance Centre non-pesticidal methods will become avail- Website: http://www.nits.net/bugbusting South East, Lower Ground Floor, able remains to be seen, but other tech- The National Pediculosis Association 40 Eastbourne Terrace, niques would be useful. Ian Burgess Website: http://www.headlice.org/ London W2 3QR, UK observes that potential methods in the Fax: +44 207 725 3460 pipeline based on a non-neurotoxic phys- Richard J. Pollack, ical approach could fill one niche. Laboratory of Public Health Entomology, 

Announcements

Are you producing a newsletter, holding a control projects is meant to differentiate this and monitoring for effects of biocontrol meeting, running an organization or rearing meeting from more basic research on nat- agents on nontarget organisms. a natural enemy that you want other biocon- ural enemy biology, such as is presented in trol workers to know about? Send us the the Entomophagous Insects Workshop. International focus is vital, and regional details and we will announce it in BNI. Focus will also be limited to the work on the coordinators have been identified who will use of parasitoids and predators, to the promote awareness of the meeting among First International exclusion of pathogens. This is largely to biological control workers in specific geo- Symposium on Biological permit the assembly of a smaller group. graphic areas. Control of A full day will be given to address pertinent Regional contacts: The first international symposium dedi- aspects of each of the following major Europe: Ulli Kuhlmann cated exclusively to biocontrol of arthro- topics: (U.Kuhlmann@ cabi-bioscience.ch) pods, will be held on 17-21 September North America (including Mexico): 2001 in Honolulu, Hawaii. It will bring 1. Classical biological control: Key issues Mark Hoddle ([email protected]) together biological control practitioners in the future expanded use of classical bio- South and Central America: from around the world to discuss interna- logical control; Better methods for coloni- Elizabeth De Nardo tional issues relating to the use of parasites zation, evaluation and monitoring of new ([email protected]) and predators against pest mites and natural enemies; Applications of molecular China: Dr Da-Wei Huang insects. Biological control of arthropods, methods to the processes of classical bio- ([email protected]) while in increasing demand, faces a series logical control; Modelling and theory as Australia, New Zealand and Oceania: of challenges that, unless addressed, will tools to clarify causes of success or failure Don Sands ([email protected]) lead to a substantial reduction in such work. of biocontrol projects. Middle East: Moshe Coll Better communication is needed among 2. Augmentative biological control: Suc- ([email protected]) scientists working in this area. Until now cesses in augmentative biological control; Africa: Peter Neuenschwander there has been no meeting that regularly Economics of production and use of reared ([email protected]) brings this group together in an atmosphere natural enemies (including storage and Volunteers are being sought as regional conducive to focused exchange of informa- shipping); Post-release dispersal, distribu- coordinators for Japan, Southeast Asia and tion. The Symposium on Biological Con- tion, and impact of augmented natural ene- Southern Asia (contact Roy Van Driesche, trol of Arthropods has as its goal to provide mies in field crops; Survey of actual and details below). for such meetings. Symposia will be held potential use in outdoor crops. every 4 years, using the same style as the highly successful meetings held by weed 3. Conservation of natural enemies in IPM Further information: biocontrol workers over the last 40 years. systems: Nectar feeding by parasitoids; Roy Van Driesche, Alternative hosts and habitat refuges for Dept. Entomology, The meeting will be limited to 250 people, parasitoids; Effects on natural enemies of Univ. of Massachusetts, so that the group is small enough to meet as using Bt crops in IPM systems; Pesticide Amherst, MA 01003, USA a whole body. The organizers hope to attract effects on natural enemies. Email: [email protected] largely if not exclusively people actually Website: http://www.isbca.ucr.edu/ conducting programmes of bio- 4. International examples of recent, impor- logical control. The focus on applied pest tant projects of classical biological control,  News 15N

Biocontrol Meeting Aphelinid and Email: [email protected] Focuses on Education Trichogrammatid Meeting Fax: +86 10 62565689 Website: http://www.ioz.ac.cn/zcd/ An international symposium ‘The Practice The systematics and biology of  of Biological Control: Importation and and Trichogrammatidae will be the subject Management of Natural Enemies and of a symposium to be held at the University Agents’ will be held on 2-5 August 2001 at of California, Riverside, USA on 18-19 African Entomology Montana State University in Bozeman, June 2001. The symposium will bring Congress USA. The symposium is intended for prac- together more than 20 of the world’s leading titioners in all disciplines of biological con- systematists, behaviorists and biological The 13th Entomological Congress, organ- trol, and is being sponsored by: the control specialists from China, India, Eng- ized by the Entomological Society of International Organization for Biological land, Europe, Canada and the USA. The Southern Africa in association with the Uni- Control, Nearctic Regional Section (IOBC- meeting will promote an exchange of ideas versity of Natal, will be held in Pietermar- NRS), the Experiment Station Committee across disciplines, stimulate greater interac- itzburg, KwaZulu-Natal, South Africa on 2- on Organization and Policy (ESCOP- tion among participants and, perhaps most 5 July 2001. This will provide a forum for BCWG), and the National Biological Con- importantly, provide a single venue for exchange of information and ideas relevant trol Institute (UDSD-APHIS-PPQ-CPHST). training students from various disciplines to entomologists of all persuasions and with interested in these parasitic wasps. particular emphasis on the needs of Africa. The science and application of biological Symposia will be held on a number of topics Parasitizing scale insects, whiteflies, control are the focus of greater public including: biodiversity and insect conserva- aphids, leafhoppers, Lepidoptera, and sev- appreciation and scrutiny than ever before. tion; biotechnology, insects and plants; eral other groups of insects, for biocontrol To develop and implement biological con- entomology and sustainable development; purposes these wasps rank among the top trol programmes, practitioners must be able forensic entomology; Hymenoptera; and ten most important taxa. While trichogram- to present the activities and goals of their insect pathology. The following workshops matids are important for augmentative con- discipline to diverse audiences, and hence are also planned: Final workshop of the trol programmes, aphelinids are primarily the focus of this symposium is on educa- Southern African Stem Borer Management used for classical biological control. The tion: what we have learned from the past Project; Insect rearing; Permits and legisla- taxonomic diversity and relationships of century of biological control efforts, and tion for collection of invertebrates; and Spa- both groups are poorly understood. Tricho- how we can apply this knowledge. It will tial data and the African entomologist. emphasize: a renewed focus on why bio- grammatidae are represented by 75 genera logical control should be of major consider- and 675 valid species, and Aphelinidae Contact: Professor Denis J. Brothers, ation in pest control; development of have 38 genera and more than 975 valid School of Botany and Zoology, and Centre effective education programmes about bio- species. However, little is known about for Environment & Development, logical control that target diverse audi- species of either family in most habitats. University of Natal, Pietermaritzburg, ences; a new set of biological control case Knowledge is especially poor in tropical Private Bag X01, Scottsville, histories that illustrate important successes; regions, where, it has been suggested, 3209 South Africa and a discussion of the issues that promote microparasitic wasps attacking early or Email: [email protected] and challenge the practice of biological cryptic life stages may be the dominant Fax: +27 0 33 260 5105 fauna. Both families exhibit peculiarities in control.  behaviour associated with host choice, Keynote sessions will cover: the need for competition and sex ratio distortion, which biological control; challenges to biological have made them model organisms for Fungal and Bacterial Plant control; success in biological control; numerous studies. Pathogens approaches to biological control of invasive Contact: Phyllis Crabtree, species; conservation of natural enemies The seventh workshop of the IOBC/ Department of Entomology, and antagonists; augmentation of natural WPRS Working Group ‘Biological Con- University of California, CA 92521, USA enemies and antagonists; size, accounta- trol of Fungal and Bacterial Plant Patho- Email: [email protected] bility and coordination of biological control gens’ will be held at Kusadasi, Turkey, in Fax: +1 909 787 3086 programmes. Panel discussions will cover: May 2002, organized by the Ege Univer- challenges to biological control; system-  sity at Izmir. The meeting will focus on the atics and biological control; approaches ‘Influence of abiotic and biotic factors on and methods used in applied biological Fifth International biocontrol agents’. Factors emphasized control; and coordinating biological control Conference of will include: microclimate, soil/substrate/ crop/fertilization, chemicals, saprophytes/ activities. Posters may be on any biological Hymenopterists control topic. nontarget microorganisms, and mesofauna, The 5th International Conference of their effects on and interaction with the Contact: Tim Kring, Hymenopterists will be held in Beijing, population dynamics/survival of antago- Univ. of Arkansas-Entomology, China from 22-26 July 2002. Nominations nists, on their biocontrol activity and on Cralley-Warren Research Laboratories, are sought for plenary speakers, and also the economics of biocontrol. 2601 N. Young Avenue, Fayetteville, ideas and organizers for symposia and spe- Contact: Yigal Elad, AR 72704, USA cialist discussion groups. Email: [email protected] Dept. of Plant Pathology, ARO, Fax: +1 501 575 3348 Contact: Chao-dong ZHU, The Volcani Center, Bet Dagan, Website: http://opal.msu.montana.edu/conf Institute of Zoology, 50250 Israel _services/biocontrol/index.htm Chinese Academy of Sciences, Email: [email protected] Beijing, Haidian, Fax +972 3 9683688 / 9683543 / 9604180 /  Zhongguancun Road 19#, P. R. China 9604180 16N BiocontrolNews and Information 2001 Vol. 22 No. 1

Website: http://www.agri.gov.il/Depts/ and P.B. McEvoy); Implementation (arti- sion (D.W. Tallamy); (3) The influence of IOBCPP/IOBCPP.html cles by A. McClay, R.W. Hansen, E.M. time dependent processes on the outcome of  Coombs et al., B. Villegas), and Moni- bioassays (T.M. Withers et al.); (4) Evolu- toring (articles by M.J. Pitcairn and D.A. tion of host range in herbivorous insects Pyke). Websites for weed biological con- (D.J. Futuyma); (5) Host specificity testing Weed Biocontrol trol are also provided. – why it is done and how it can be improved Publications (R. Dekker van Klinken); (6) Evaluting host Book 2: This 95 pp. publication comprises The USDA Forest Service has produced ranges – rationale, methodology and inter- the proceedings of the session: ‘Host specif- two books on weed biological control that icity testing of exotic arthropod biological pretation (D.P.A Sands & R.G. Van Drie- are now available to interested individuals. control agents – the biological basis for sche); (6) Host specificity assessments – Book 1: ‘Weed biocontrol: extended abstracts improvement in safety’, which was held case studies (U. Kuhlmann et al.). from the 1997 Interagency Noxious Weed during the X International Symposium on Symposium’ (58 pp.). This covers the fol- Biological Control of Weeds, Bozeman, Contact: Roy Van Driesche, University of lowing aspects of weed biological control: Montana, USA, July 1999 (eds:R.G.Van Massachusetts at Amherst, USA History of weed biological control (articles Driesche, T. Heard, A. McClay & R. Email: [email protected] by J.R. Coulson and L.A. Andres); Regula- Reardon). It covers: (1) Concepts of insect- or Richard Reardon, USDA Forest Service tion (articles by R.E. Pizel, G.P. Markin host plant selection behaviour and applica- Email: [email protected] and S. Stenquist): Safety (articles by Q. tion to host specificity testing (T.A. Heard); Paynter & J.L. Littlefield, J.K. Balciunas, (2) Physiological issues in host range expan- 

Conference Reports

IOBC Water Hyacinth hyacinth. From the presentations and dis- “The mission of the IOBC Working Group Meeting cussions the following ideas emerged for for the Biological and Integrated Control of further investigation. Water Hyacinth is to promote better man- The second Working Group Meeting for agement of water hyacinth through: the Biological and Integrated Control of • Investigate the impact of cold climates WaterHyacinthheldevery2yearsunder on the success of biological control. • facilitation of interactions, the auspices of the International Organiza- • dissemination of information, and • Use plant competition studies between tion for Biological and Integrated Control water hyacinth and other aquatic plants • identification of research needs. of Noxious Animals and Plants (IOBC) as an indication of how effective took place in Beijing, China on 9-12 This will be achieved by: particular agents are. October 2000. This meeting brought • holding a meeting every 2-3 years, together 31 delegates from 11 countries • Investigate the compatibility of the • publishing the meeting proceedings, a with the common purpose of identifying different control options that could be water hyacinth newsletter and main- suitable biological and integrated control used in integrated management. taining website, solutions for water hyacinth. • Select suitable locations and undertake • supporting activities that contribute to The meeting took the form of a series of integrated management of water better management of water hyacinth.” oral presentations, which included three hyacinth where biological control is The next meeting will be held in Uganda on keynote presentations covering the biolog- the base technique. ical control of water hyacinth using insects, the shores of Lake Victoria in early August the biological control of water hyacinth • Identify and conduct surveys in areas 2002. where detailed surveys for additional using pathogens both as classical biological By: Martin Hill, PPRI, South Africa control agents and as mycoherbicides, and natural enemies (both insects and  an overview of the current status of pathogens) have not been conducted in research on the weed in China. In addition, the region of origin of water hyacinth. Fifth Chromolaena there was a series of reviews from different • Research the interaction between countries around the world, several papers insect natural enemies and pathogen Workshop dealt with the need for additional biological natural enemies. The Fifth International Workshop on Bio- control agents, both insects and pathogens, logical Control and Management of Chro- and there were several papers dealing with • Make a thorough investigation into the molaena odorata was held at the Umhlanga aspects of the integrated control of the development of mycoherbicide for Protea Hotel near Durban, South Africa, on weed. These papers will be published by water hyacinth. 23-25 October 2000, with a post-workshop ACIAR, in a proceedings, which is The workshop closed with a general fieldtriptonorthernKwaZulu-Natalprov- expected to be available in the first half of meeting of the working group (the partici- ince from 26-28 October. It was organized 2001. pants). During the meeting the following by the Plant Protection Research Institute One of the roles of this working group is to mission statement was developed for the of the Agricultural Research Council of identify further research needs on water working group: South Africa (ARC-PPRI) in association News 17N with the International Organization for Bio- • biological control of chromolaena (10 candidate agents are being tested in logical Control of Noxious Animals and papers). South Africa. These include the leaf- Plants (IOBC), the provincial conservation mining agromyzid fly, Calycomyza A mid-workshop tour on 24 October took service (KZN Wildlife) and Dr R.N. eupatorivora, and the stem-boring wee- delegates to the ARC-PPRI laboratories at Muniappan of the University of Guam. vil, Lixus aemulus, both of which have Cedara to view the insects in quarantine This is the second time the workshop has been tested for host specificity to near- presently being cultured and tested as bio- been held in Africa (the first being the completion, with favourable results. control candidates on chromolaena and the Third Workshop, held in Abidjan in 1993). The stem tip-galling weevil Conotra- South American invasive Solanum mauri- chelus reticulatus and the root-boring It was attended by 20 international dele- tianum (bugweed). flea-beetle Longitarsus horni are also gates from 12 countries (India, the Philip- The workshop ended with an afternoon of: undergoing testing in South Africa. The pines, Indonesia, Papua New Guinea, ACIAR programme has also resulted in Guam (USA), Australia, Malaysia, Ghana, 1. Summarizing the workshop sessions. the establishment of centres of exper- Congo-Brazzaville, Cameroon, Benin, 2. Drawing up a list of biocontrol agents tise in most Southeast Asian countries, Swaziland) and 30 South African dele- established in, available from and for the testing and distribution of agents gates. These delegates included agricultur- wanted by the various countries, and on chromolaena. alists, water conservationists, biologists funding possibilities for regional bio- and nature conservationists. Major spon- 2. The progress made on unblocking the control programmes. sors of delegates included the Australian biocontrol programme on chromolaena Centre for International Agricultural 3. The proposal and discussion of recom- in West and Central Africa. Due to Research (ACIAR) (for Southeast Asia) mendations emanating from the work- perceived conflicts of interest over the and the Technical Centre for Agricultural shop. role of chromolaena in agriculture and Rural Co-operation (CTA) based in the here, outside of Ghana little progress This information has been published in the Netherlands (for West and Central Africa). has been made over the past decade on Chromolaena odorata Newsletter No. 14. biocontrol of chromolaena. Discussion The Workshop was opened by the Minister The post-workshop fieldtrip afforded dele- between parties representing these of Agriculture and Environmental Affairs gates the opportunity to see chromolaena in conflicting interests at the Fifth for KwaZulu-Natal Province, Mr Narend the field and its effect on renowned conser- Workshop resulted in a commitment to Singh. The keynote address was delivered vation areas such as the St Lucia estuary (a host a meeting within the next year by the Chief Executive of the World Wide World Heritage Site) and the Hluhluwe- bringing together all stakeholders in Fund for Nature (South Africa), Dr Ian Umfolozi Reserve, at the historical centre West and Central Africa. Macdonald, and was entitled “Chromo- of the effort to conserve both black and laena at the cutting edge, or, Why the battle In addition, several delegates who had not white rhinoceros. Control efforts on chro- to control Chromolaena is a flagship battle visited South Africa previously and thus molaena and other invasive plants, under- for the new millennium”. Dr Macdonald not seen the southern African form of chro- taken by KZN Wildlife and the Department outlined his belief that, since chromolaena molaena were struck by its different growth of Water Affair’s Working-for-Water Pro- has all the characteristics of one of the habit, morphology and ecology. That this gramme, were also outlined on the trip. world’s worst invasive plants, if it can be form of chromolaena has a different beaten, any invasive can be beaten. As an Two of the highlights of the workshop biology (e.g. susceptibility to fire) to that invasive of tropical, largely developing were: invading other areas of the world was also countries, the successful struggle against evident from research presented at the chromolaena is a good indicator that we can 1. The progress that has been made since workshop. get to grips with many of the underlying the Fourth Workshop (India, 1996) on Papers for the Proceedings are currently problems affecting conservation in the chromolaena biocontrol research and being collated for refereeing. The Proceed- tropics, which is the repository for much of implementation. This is largely due to ings will be published during the course of the world’s biodiversity. Dr Macdonald’s theproactiveroletakenbyACIARin 2001. optimism for such success came from the Southeast Asia, and the South African fact that the chromolaena workshops are and Ghanaian programmes. The By: Costas Zachariades, PPRI, South being held on a regular basis and that they tephritid stem-galling fly, Procecido- Africa chares connexa, is currently being dis- are bringing together on-the-ground  workers from the developing world; by the tributed widely in Southeast Asia, progress being made on biocontrol, a sus- where it is establishing readily and tainable rather than symptomatic form of causing significant damage. The defoli- IOBC Fungal and Bacterial control; and by the global groundswell of ating arctiid moth, Pareuchaetes pseu- Plant Pathogens awareness and action on invasive species. doinsulata, which has had an inconsist- Workshops ent history in terms of its success as a Thirty-three oral and two poster presenta- biocontrol agent around the world, is A workshop of the IOBC/WPRS Working tions were made. Presentations were spreading and causing significant dam- Group ‘Biological Control of Fungal and grouped into seven sessions with the fol- age in both Ghana and Sumatra (Indo- Bacterial Plant Pathogens’, held in lowing general topics: nesia). The nymphalid Actinote sp., Taormina, Sicily in September 2000, • country and regional reports (13 whose larvae defoliate chromolaena, focused on the biocontrol of foliar patho- papers), was rejected as an agent for South gens. Organized and chaired by Yigal Elad Africa because it fed on indigenous (Bet Dagan, Israel) in cooperation with • taxonomy, ecology and impacts of Mikania species, but has been released EFPP (the European Foundation of Plant chromolaena (5 papers), in Sumatra, as Mikania is invasive Pathology), it was attended by some 200 • impacts and management of chromo- there. Its establishment has yet to be people, and was devoted to the interaction laena (4 papers), confirmed. A number of promising new of biocontrol agents with foliar plant patho- 18N BiocontrolNews and Information 2001 Vol. 22 No. 1 gens, and to biological control and its cooperation, was attended by 108 people restraining the pathogenicity enzymes of mechanisms. Presentations included: an from 33 countries. Presentations and dis- the pathogen and its antigens, and arresting introduction to the subject and field use of a cussions focused on the use of biocontrol the production of multiplication propagules biocontrol preparation; induced resistance agents against soilborne and foliar patho- of the pathogens. It was noted that in many to control leaf pathogens by microbial inoc- gens of all kind of plants either in the open systems multiple modes of action are ulants that are applied to the root system; field or in greenhouses and at post harvest involved. Several commercial biocontrol strategies and application of biological con- stages, in the roots, stems, leaves, flowers, agents were presented. The improvement trol for a disease of stone fruits; and how it fruits and wood. It dealt with the use and of efficacy, integration of biocontrol agents is possible to cope with variability and implementation of biocontrol, various inconsistency of biocontrol. modes of action, the nature and use of among themselves and with other means of genes that originate from biocontrol agents, disease control were discussed with respect The sixth workshop of the Working Group, involvement of mycorrhizae and improve- to the improved implementation of biocon- held on 30 November-3 December 2000, ments of biocontrol activity. Modes of trol and the reduction of variability in the focused on ‘Biocontrol agents, modes of action that were dealt with can be catego- performance of biocontrol agents. action and their interaction with other rized as competition including space and means of control’. The meeting, organized nutrient exclusion, parasitism including By: Yigal Elad, Bet Dagan, Israel by Enrique Monte (Salamanca, Spain) and phages that are hosted by bacteria, antibi- Yigal Elad, with local and international osis, different modes of induced resistance,