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Home , Ambrosia artemisiifolia, Dicoria (plant), Dicoria canescens, Epiblema strenuana, Ophraella communa, Ragweed

296 Chapter 43

43 L., Common ()

Alan K. Watson and Miron Teshler McGill University, Ste Anne de Bellevue, Québec

43.1 Pest Status bronchitis and . The complex of 22 proteins released from ragweed Common ragweed (herbe-à-poux), Ambrosia grains are some of the most powerful artemisiifolia L. (Asteraceae), a native antigens/ known (Bagarozzi and annual , occurs throughout Travis, 1998). Although ragweed pollen and is most abundant in can travel great distances, allergenic southern Ontario and Quebec and in the reactions are caused by A. artemisiifolia north-east and north-central US states growing very close to the susceptible (Bassett and Crompton, 1975). It is a individual. Within urban and suburban pioneer species that fl ourishes in disturbed regions, effective control of ragweed habitats such as along rights-of-way, in infestations would substantially lower the vacant lots and cultivated fi elds. Ambrosia incidence of allergenic reactions. Ambrosia artemisiifolia is often the most frequent artemisiifolia has rapidly become a serious species in these habitats, and forms dense agricultural weed, especially in vegetable linear populations along the fi rst metre of crops grown in the muck soils of south- roads and highways in eastern western Quebec and Ontario. Ambrosia (DiTommaso, 2004; Joly et al., 2011). There artemisiifolia seeds germinate in spring, is strong historical evidence that the the vegetative phase occurs from May to expansion of road networks has contrib- August, fl owering commences on the fi rst uted to the spread of A. artemisiifolia week of August and produces copious (Lavoie et al., 2007; Simard and Benoit, quantities of airborne pollen, and seeds are 2010). Moreover, this weed species is set in late summer or autumn. Individual highly plastic and very variable in size, leaf produce 3000 to 62,000 seeds that shape, infl orescence form, degree of can remain viable for 39 years or more hairiness and life-form strategy. Germin- when buried in the soil (Bassett and ation response for roadside populations of Crompton, 1975). A. artemisiifolia may be locally adaptive to salinity and allows A. artemisiifolia to emerge relatively early in spring thus 43.2 Background providing a competitive advantage over later emerging roadside plants (DiTommaso, Ambrosia artemisiifolia can easily be 2004). uprooted in most soil types; however, it Ambrosia artemisiifolia pollen is the can readily adapt to frequent mowing by primary cause of allergenic hay fever, quickly producing new stems and fl owers

© CAB International 2013. Biological Control Programmes in Canada 2001–2012 (eds P.G. Mason and D.R. Gillespie) Chapter 43 297

below the cutting height (Vincent and grammes in , eastern and Ahmin, 1985; Patracchini et al., 2011). eastern , with variable results (Julien Repeated mowing can reduce pollen pro- and Griffi ths, 1998; Zhou et al., 2009). duction but will not reduce the seed bank Tarachidia candefacta Hübner (Lepi- (Simard and Benoit, 2011). Various herbi- doptera: Noctuidae) and cides and tank mixtures have provided suturalis Fabricius (Coleoptera: Chrys- control of A. artemisiifolia in maize, Zea omelidae) are two natural enemies from mays L. (Poaceae), and , Glycine Canada and the USA that were introduced max (L.) Merr. (Fabaceae), crops, but popu- into the former Soviet Union in 1966. They lations of A. artemisiifolia have developed became established and provided control resistance to herbicides having the follow- of the invasive A. artemisiifolia (Reznik, ing sites of action: Photosystem II inhibi- 1991; Goeden and Teerink, 1993). Of great tors, ALS inhibitors, PPO inhibitors, ureas recent interest, populations of T. cande- and amides and glycines (Heap, 1997; facta have recently begun to expand, many Patzoldt et al., 2001; Saint-Louis et al., years after introduction in southern Russia 2005), thus restricting control options. (Poltavsky et al., 2008). Zygogramma Several herbicides, including 2,4-D suturalis leaf were also introduced ((2,4-dichlorophenoxy) acetic acid), MCPA into Croatia, but failed to become ((2 methyl-4-chlorophenoxy) acetic acid) established (Igrc et al., 1995). Releases of Z. and dicamba (3,6-dichloro-2-methoxy- suturalis in China in 1985, both from benzoic acid), have been the mainstay of A. Canada and from the former Soviet Union, artemisiifolia control strategies along road- established in some locations, but failed in ways and marginal areas in most com- others (Wan et al., 1995). In 1990, Z. munities. However, herbicide use for the suturalis was introduced into Australia control of A. artemisiifolia in urban areas from the USA but failed to establish (Julien has declined in recent years due to the and Griffi ths, 1998). implementation of municipal and pro- Ambrosia artemisiifolia is presently vincial legislation throughout Canada that considered under good control in south- bans or severely restricts herbicide use. eastern Queensland and in northern New Currently, foliar application of sodium South Wales and is regarded as an chloride solutions is being used in cities outstanding success in Australia (Palmer et and municipalities of Quebec (Grégoire et al., 2010); but the success is not due to the al., 2002; Watson, 2008). Ambrosia introduction of specifi c biological control artemisiifolia is very sensitive to desic- agents against A. artemisiifolia. From 1980 cation and dries up whereas surrounding to 1984, three biological control agents, plants are not impacted. including the leaf-feeding chrysomelid Zygogramma bicolorata Pallister (Coleoptera: Chrysomelidae), the sap- 43.3 Biological Control Agents sucking bug concinna (Stål) (: ) and the tip- 43.3.1 galling strenuana (Walker) (: ) from Over 400 species have been known were introduced into Australia for the to attack A. artemisiifolia (Harris and biological control of parthenium weed, Piper, 1970) and more than 30 of these L. (Asteraceae) have been examined as potential biological (McFadyen, 1992). Parthenium hystero- control agents for A. artemisiifolia phorus is very closely related to the genus (Kovalev, 1970). Ambrosia artemisiifolia is Ambrosia and coincidentally it was an exotic invasive weed in Australia, observed that these three insects also Europe and central Asia. There have been attacked A. artemisiifolia. Epiblema strenu- several classical biological control pro- ana was the most effective biological 298 Chapter 43

control agent, and has reduced the size, pupation, the 3rd instar larvae spin loosely abundance and pollen production of woven cocoons on the upper or lower leaf ragweed to a greater extent than the two surfaces. Development time for the pupa other insects. stage is about 7 days and the total development time from oviposition to adult emergence is about 22 days (Welch, 1978). 43.3.1.1 Inundative biological control From 1995 to 1999, host-specifi city, life In Canada, Z. suturalis and table, biotic potential, mortality, feeding communa LeSage (Coleoptera: Chrys- potential and mass rearing studies of O. omelidae) are native natural enemies of A. communa were conducted on transplanted artemisiifolia. They have been the main ragweed plants (Teshler et al., 1996, 1999). focus of biological control studies at the Conclusions derived from these studies Macdonald Campus of McGill University suggested that O. communa was a very as inundative biological control agents in promising candidate for inundative bio- Quebec (Teshler et al., 2002). In North logical control of A. artemisiifolia. Char- America, under natural conditions, popu- acteristics of O. communa include: (i) being lation densities and impact of O. communa native to Quebec; (ii) having a restricted on A. artemisiifolia tend to be low, host range; (iii) causing signifi cant A. presumably because of strong attack by artemisiifolia damage, especially at the predators and parasitoids by the end of vulnerable seedling stage; and (iv) having a summer (Teshler et al., 2002). Thus, high intrinsic reproductive rate, which has inundative releases of Z. suturalis and O. facilitated its mass rearing under controlled communa early in the growing season conditions (Teshler et al., 2002; Dernovici (Teshler et al., 1996) would be a reasonable et al., 2006). In 1999 and 2000, inundative approach. However, the reduction or cage releases were conducted using various cessation of Z. suturalis oviposition on O. communa beetle-to-A. artemisiifolia and extensively damaged plants (as observed in O. communa egg-to-A. artemisiifolia ratios the former Soviet Union) and pupation in in carrot, Daucus carota L. subsp. sativa soil are the most important limitations for Schübl. & M. Martens (Apiaceae), cabbage, the mass-rearing of this species (Teshler et Brassica oleracea L. (Brassicaceae ), and al., 2002). Thus, O. communa became the soybean fi elds in Sherrington and St agent of choice. Isidore, Quebec. Four to fi ve O. communa is oligophagous beetles per A. artemisiifolia caused and feeds on various members of the complete defoliation and death of 4- to subtribe Ambrosiinae (Asteraceae). All 6-leaf-stage plants within 14 days. In 2001, developmental phases of this multivoltine small-scale open fi eld release trials were insect occur on A. artemisiifolia. Fertile conducted with O. communa in soybean adult females overwinter in soil debris and fi elds in St Isidore, Quebec. Larval feeding are observed, along with their eggs, on of O. communa signifi cantly reduced A. ragweed seedlings in early spring. Eggs are artemisiifolia growth when the insect generally deposited in clusters on the host infestation occurred early in the season plant and the development time of each of (Teshler et al., 2002). the three larvae instars is 3–4 days (LeSage, Ophraella communa was mass reared 1986). Larval instars are distinguished by on transplanted A. artemisiifolia plants in the size and colour of the head capsule the greenhouse. All development stages of (Welch, 1978). Neonate larvae generally O. communa can be produced on 6- to skeletonize leaves while adults and older 8-leaf-stage ragweed plants in the instar larvae can devour the entire leaves. greenhouse with a temperature of 24±4°C, Under favourable conditions, O. communa a relative humidity of 60±20% and a adults and larvae can completely defoliate photoperiod of 16:8 L:D conditions ragweed plants (LeSage, 1986). Prior to (Teshler et al., 1999). Approximately 40–60 Chapter 43 299

beetles (1:1 male:female ratio) and 100–200 mass rearing of O. communa was at 28°C larvae can be kept per rearing cage (Zhou et al., 2010). Initial release density (17.5×31×8 cm). Ophraella communa of O. communa adults for effective control pupae were collected and plants replaced of A. artemisiifolia in the fi eld in China bi-weekly. Pupae were placed into should be ≥1 beetles per plant at early 17.5×31×8 cm plastic containers and stored growth stages or ≥12 beetles per plant at under laboratory conditions until adults the later growth stages (Guo et al., 2011). emerged (Teshler et al., 2004). A multi- purpose device was then developed for the collection, short-term storage, transport 43.3.2 Pathogens and delivery of O. communa pupae or adults (Teshler et al., 2004). To achieve a In Canada, the incidence and biological successful A. artemisiifolia control through control potential of a forma specialis of large-scale releases of O. communa, mass Pustula tragopogonis (Pers.) Thines (= production techniques for O. communa Albugo tragopogonis (DC) Gray) (Albu- need to be improved. ginaceae) on A. artemisiifolia has been Mass production of O. communa on described (Hartmann and Watson, 1980). transplanted A. artemisiifolia plants in the When A. artemisiifolia is attacked by P. greenhouse is being compromised by high tragopogonis, considerable damage can labour costs, large space requirements and occur and signifi cant reduction in pollen high growth-facility costs. The collection and seed production occurs if systemic and handling methods of the specimens infection is achieved (Hartmann and need to be mechanized. A semi-artifi cial Watson, 1980); however, diffi culties in medium for rearing Coccinellidae spp. mass producing this white blister have (Daniel Coderre, Montreal, Quebec, 2012, so far prevented it from being produced pers. comm.) was tested and found not commercially (Teshler et al., 2002). In suitable for rearing larvae and adults of O. Quebec, a Phoma sp. (incertae sedis) was communa. The evaluation of several media discovered on A. artemisiifolia and evalu- used for rearing various herbivorous insect ated as a potential mycoherbicide candi- species and the incorporation of A. date (Brière et al., 1995). A combination of artemisiifolia leaf powder as a feeding this Phoma sp. and O. communa provided stimulant were conducted but limited pro- a synergistic effect and resulted in high gress has been achieved in fi nding an plant mortality (Teshler et al., 1996). artifi cial diet for the production of O. Unfortunately, this Phoma sp. isolate lost communa. its virulence and attempts to revive or re- Ophraella communa was accidentally isolate it from natural sites were unsuc- introduced in Japan in 1996 (Yamazaki et cessful (Teshler et al., 2002). al., 2000; Yamanaka et al., 2007) and in In Europe, A. artemisiifolia has once China in 2001 (Zhang et al., 2005). A mass- again become a major target for biological rearing programme was established in control with the recent approval of a China with the aim to implement inun- coordinated European research programme dative release of mass-reared O. communa on ‘Sustainable Management of Ambrosia in heavily A. artemisiifolia-infested habi- artemisiifolia in Europe’ (COST FA1203- tats (Zhou et al., 2009). Critical life table SMARTER). Currently, there are 18 insects data on the establishment potential of O. and fi ve fungal pathogens considered as communa in new environments with promising candidates for classical bio- diverse temperature regimes were gener- logical control (Gerber et al., 2011). How- ated (Zhou et al., 2010, 2011). Ophraella ever, it was noted that an inundative communa preferred moist microclimates approach will be necessary for A. with 75–90% relative humidity (Zhou et artemisiifolia-infested crop fi elds. al., 2009). The optimum temperature for Ophraella slobodkini Futuyma (Coleoptera: 300 Chapter 43

Chrysomelidae) and the fungus, Septoria and fungal pathogens for host specifi city epambrosiae D.F. Farr, Sydowia (Myco- and effectiveness against A. artemisiifolia. sphaerellaceae) are candidate biological control agents for mass-rearing and repeated releases against ragweed (Gerber Acknowledgements et al., 2011). The future success of the biological control of A. artemisiifolia We thank Robert Anderson (Canadian worldwide may rely on successful opti- Museum of Nature, Ottawa) and Laurent mization of mass rearing techniques. LeSage (Canadian National Collection of Insects and Arachnids, Ottawa) for confi rming the taxonomic status of 43.4 Future Needs Ophraella communa; Daniel Coderre (Université du Québec à Montréal) for his Future work should include: collaboration in diet development, and Luc 1. Developing and optimizing mass rearing Brodeur (Phytodata Inc.) for the assistance for O. communa, O. slobodkini and S. in securing AAFC/NSERC Research epambrosiae; Partnership Program funding. 2. Evaluating additional candidate insects

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44 Centaurea diffusa Lamarck, Diffuse Knapweed, and Centaurea stoebe subsp. micranthos (S.G. Gmel. ex Gugler) Hayek, Spotted Knapweed (Asteraceae)

Rob S. Bourchier1 and Brian H. Van Hezewijk2 1Agriculture and Agri-Food Canada, Lethbridge, Alberta; 2Natural Resources Canada, Canadian Forest Service, Victoria, British Columbia

44.1 Project Status control in Canada with the fi rst agent, Urophora affi nis (L.) (Diptera: Tephritidae), Knapweeds, Centaurea diffusa Lamarck released in 1970 (Harris and Myers, 1984). and Centaurea stoebe subsp. micranthos The biology of all of the biological control (S.G. Gmel. ex Gugler) Hayek (=Centaurea agents for knapweeds in Canada was maculosa Lamarck) (Asteraceae), were reviewed in previous volumes of this series among the earliest targets for biological (Harris and Myers, 1984; Bourchier et al.,

© Her Majesty the Queen in Right of Canada, as represented by the Minister of Agriculture and Agri-Food Canada 2013