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Miller, R.R., Svejcar, T.J., Rose, J.A. and McInnis, M.L. (1994) development, water relations, and carbon allocation of heart-podded hoary cress. Agronomy Journal 86, 487–491. Mulligan, G.A. and Findlay, J.N. (1974) The biology of Canadian weeds. 3. Cardaria draba, C. chalepensis, and C. pubescens. Canadian Journal of Plant Science 54, 149–160. Redfern, M. and Shirley, P. (2012) Checklist of British Galls. Available at: http://www.british-galls. org.uk/index.htm (accessed 2 February 2012). Selleck, G.W. (1965) An ecological study of lens- and globe-podded hoary cresses in Saskatchewan. Weeds 13, 1–5. Sheley, R.L. and Stivers, J. (1999) Hoary Cress (Whitetop). In: Sheley, R.L. and Pertroff, J.K. (eds) Biology and Management of Noxious Rangeland Weeds. Oregon State University Press, Corvallis, Oregon, pp. 401–407. Skinner, K., Smith, L. and Rice, P. (2000) Using noxious weed lists to prioritize targets for developing weed management strategies. Weed Science 48, 640–644. Van den Berg, C. and van de Sande, J.C. (1999) Ceutorhynchus turbatus new for the Netherlands (Coleoptera: Curculionidae). Entomologische Berichten 59, 157–159.

51 vulgare Lam., Oxeye Daisy ()

Alec S. McClay,1 Sonja Stutz2 and Urs Schaffner2 1McClay Ecoscience, Sherwood Park, Alberta; 2CABI, Delémont, Switzerland

51.1 Pest Status by seed, and individual expand by rooting at the stem bases and by Lamarck (=Chrys- spread (Clements et al., 2004). Leuc- anthemum leucanthemum L.) (Asteraceae) anthemum vulgare was reported to be is a shallow-rooted perennial herb native to naturalized in Quebec by the 18th century . It has a short creeping rootstock (Lavoie et al., 2012). It is widespread that gives rise to erect, simple or slightly across , occurring in all provinces branched stems usually up to 90 cm in and in the Yukon Territory (Clements et al., height, generally one to two per plant but 2004; Bennett and Mulder, 2009) and also sometimes forming larger clumps. Leaves throughout the USA, although less frequent are spatulate or obovate at the base of the in southern states (Strother, 2006; USDA- plant and ligulate higher on the stem. The NRCS, 2012). Some authors separate the fl ower head is a daisy, 2.5–7.5 cm in tetraploid Leucanthemum ircutianum DC. diameter, with a yellow central disk and from the diploid L. vulgare, and AFLP white rays. Leucanthemum vulgare spreads analysis of European material suggests that

© CAB International 2013. Biological Control Programmes in Canada 2001–2012 (eds P.G. Mason and D.R. Gillespie) 338 Chapter 51

L. ircutianum is a hybrid of L. vulgare and industrial and other non-crop areas in another, unknown, diploid parent (Ober- Canada. Clements et al. (2004) report prieler et al., 2011). Both diploids and several other herbicides that are effective tetraploids occur in on L. vulgare but point out that most of (Clements et al., 2004), but preliminary these will also eliminate legumes and other results on a number of North American desirable broad-leafed in pastures. populations suggest that most of them are Fertilizing pastures can also be effective in diploid (Stutz et al., 2012). suppressing L. vulgare by stimulating the Leucanthemum vulgare is primarily a growth of grasses and other competing weed of pastures, rangelands and roadside vegetation (Olson and Wallander, 1999; areas, and can form very dense, extensive Clements et al., 2004). populations in pastures, where it is Leucanthemum is a European of generally avoided by grazing cattle, and 41 species, most of which, except for the promotes soil erosion because of its widespread L. ircutianum and L. vulgare, shallow root system. Overgrazing often have quite limited distributions in Europe promotes infestations of L. vulgare (Greiner et al., 2012). It belongs to the tribe (Clements et al., 2004). It is usually not and the subtribe Leuc- considered a problem in annual crops, antheminae, which has no native North although it was the fourth most abundant American species (Oberprieler et al., 2009), weed species in a survey of spring cereals so L. vulgare is taxonomically well isolated in New Brunswick in 1986–1987 (Thomas from potential native non-target species. et al., 1994). It is of major concern as a The main non-target taxon of concern is contaminant in grass seed production in the popular horticultural Shasta daisy, the Peace River region of British Columbia Leucanthemum × superbum (Bergmans ex (British Columbia Ministry of Agriculture, J.W. Ingram) D.H. Kent (Asteraceae), a Food and Fisheries, 2002) and cannot be hybrid introduced by the famous American separated from seed of timothy, Phleum plant breeder Luther Burbank in 1901 pratense L. (Poaceae) (Manitoba Agri- (Hawke, 2007). According to Burbank culture, Food and Rural Initiatives, 2012). (1914) its parentage included three Euro- Leucanthemum vulgare is an invasive pean Leucanthemum species, L. vulgare, L. species of concern in natural areas, such as maximum (Ramond) DC. and L. lacustre Riding Mountain National Park, Manitoba (Brot.) Samp. (Asteraceae) and the (Otfi nowski et al., 2007) and Yellowstone Japanese Nipponanthemum nipponicum National Park, USA (Olliff et al., 2001). It is (Franch. ex Maxim.) Kitam (Asteraceae). listed as a provincial noxious weed in However, given that Burbank ‘kept no Alberta, Saskatchewan and Manitoba, and systematic records’ and had ‘unorthodox as noxious in the Cariboo, North Okanagan, views of heredity’ (Crow, 2001), the true Thompson-Nicola and Peace River regions parentage of Shasta daisy must be of British Columbia. Leucanthemum considered unclear. For example, Tahara vulgare is a ‘primary noxious weed seed’ (1921) disputes Burbank’s identifi cation of under the Canada Seeds Act, and there is a N. nipponicum and suggests that the zero tolerance for oxeye daisy seed in most species he used was probably Arct- grades of most seed species sold in Canada anthemum arcticum (L.) Tzvelev (Minister of Justice, 2005). (Asteraceae). Numerous varieties of Shasta daisy are available, most of which fall within L. × superbum but a few of which 51.2 Background are probably selections of L. vulgare (Hawke, 2007). It will be necessary to test a Several herbicides containing amino- range of Shasta daisy to assess pyralid or aminopyralid + metsulfuron- possible risk to this horticultural plant methyl are currently registered for control from potential biological control agents for of L. vulgare in rangeland, pasture, L. vulgare. Chapter 51 339

Natural enemies of L. vulgare in North sidered as potential biological control America have not been surveyed in detail. agents based on records of their restricted Guillet and Arnason (1995) found the host range: the root-mining polyphagous species Argyrotaenia velu- aeratana (Pierce & Metcalfe) tinana Walk. and Sparganothis sulfureana and D. baixerasana Trematerra, the shoot- (Clemens) (: ) feed- mining D. consortana Stephens ing on L. vulgare fl ower heads around (Lepidoptera, Tortricidae), the root-feeding Ottawa, Ontario. The fl ower-head feeding weevils Cyphocleonus trisulcatus (Herbst) weevil Microplontus campestris (Gyllen- and Diplapion stolidum (Germar) (Cole- hal) (=Ceutorhynchus campestris Gyllen- optera, Brentidae) and the fl ower-head hal) (Coleoptera: Curculionidae) has been attacking fl y neesii Meigen collected in Ontario on several occasions (Diptera, ). since 1971 (Anderson and Korotyaev, Initial in-depth studies on the biology, 2004). Macrosiphoniella leucanthemi host-range and impact of biological control Ferrari and M. sanborni (Gillette) (Hem- candidates focused on D. aeratana, which iptera: Aphididae) are recorded feeding occurs throughout western and central on L. vulgare in North America (Miller Europe. The larvae of D. aeratana feed and Stoetzel, 1997; Stoetzel and Miller, mainly inside the roots, where they also 1999). overwinter. Around March–April, they leave the roots and pupate in the soil. Adult D. aeratana, which have brownish 51.3 Biological Control Agents forewings and a wingspan of 12–16 mm, fl y in May and June. They live for Foreign explorations started in 2008 with approximately 1 week and females lay literature and fi eld surveys. In total, at least about 100 eggs. First results from no-choice 80 , 7 nematodes and 16 fungi were larval development tests indicated that the found to be associated with oxeye daisy in fundamental larval host range of D. its native range (Schaffner et al., 2008). aeratana is mainly restricted to the genus Particularly well represented is the genus Leucanthemum, the larvae being able to Dichrorampha (Lepidoptera, Tortricidae), develop on L. vulgare, L. ircutianum and which appears to have undergone speci- on Shasta daisies. A few larvae were also ation on L. vulgare and related plants; in found on other test plant species, but it total, some 15 Dichrorampha spp. have remains to be shown whether the larvae been recorded from Leucanthemum spp. were indeed D. aeratana or whether they (Razowski, 2003). The fl ower head-attack- belonged to other Dichrorampha spp. ing weevil M. campestris was originally (Stutz et al., 2012). Under multiple-choice identifi ed as a potential candidate bio- conditions, L. vulgare and L. ircutianum logical control agent. Adults of this species were attacked signifi cantly more often than start feeding in late April, and mating and Shasta daisies (S. Stutz and U. Schaffner, oviposition into the fl ower heads was Delémont, Switzerland, 2012, unpublished observed from mid-May onwards. Obser- data). vations made during the fi eld surveys and The root-feeding weevil C. trisulcatus, in the laboratory confi rmed that larval which has a scattered distribution through- feeding is restricted to the receptacle and out Europe (Dieckmann, 1983), was also that seeds were not attacked. As a identifi ed as a promising potential bio- consequence, attack by M. campestris was logical control agent in the initial literature shown to have no or minimal impact on survey (Schaffner et al., 2008). Larvae feed seed output of oxeye daisy. Microplontus externally on roots of L. vulgare and L. campestris was therefore dropped from the ircutianum. This species appears to be list of potential biological control agents. quite rare in Europe, and most site records To date, six European species are con- mentioned in the literature are more than 340 Chapter 51

50 years old. However, in 2011 and 2012, 51.4 Evaluation of Biological Control C. trisulcatus was found at several locations in southern Germany and No biological control agents have been southern France, allowing establishment of released to date in Canada or the USA. laboratory rearing and initiation of investigations on the biology of this species. 51.5 Future Needs The fl ower head-attacking fl y T. neesii is very common throughout Europe (White, Future work should include: 1988) and was frequently found during fi eld surveys. Larvae feed in the receptacle 1. Additional molecular and genetic stud- and on the developing seeds, thereby ies to clarify the identity of invasive reducing seed output (Robinson, 2008). Leucanthemum populations in North pupate in the fl ower heads America and the relationships between and adults emerge in summer. This species oxeye and Shasta daisy; has one generation per year, and over- 2. Host-specifi city testing with the candi- wintering occurs in the adult stage. Usually date biological control agents D. aeratana, only seed heads collected from plants C. trisulcatus, T. neesii and D. stolidum; fl owering in May or June were infested; 3. Assessing the potential impact on L. vul- seed heads collected later in the year from gare of all candidate biological control plants that had regrown after mowing were agents; not infested. 4. Locating large fi eld populations of D. The root-feeding weevil D. stolidum is baixerasana and D. consortana. considered to be a specialist on L. vulgare (Dieckmann, 1977). During fi eld surveys in southern France, Switzerland, southern Acknowledgements Germany, Czech Republic and southern Poland, L. vulgare plants infested by D. Work on D. aeratana, C. trisulcatus, D. stolidum were found at several locations. stolidum and T. neesii in Switzerland was Adults emerging from fi eld-collected roots funded by the British Columbia Ministry of in July were transferred into plastic Forests, Lands and Natural Resource cylinders containing a cut rosette of L. Operations and the Montana Noxious vulgare. The food was regularly changed Weed Trust Fund, through Montana State and the rosettes checked for eggs. Adults University. We thank J. Gaskin, USDA-ARS hibernated and started to lay eggs in April Sidney, Montana, for conducting the (S. Stutz and U. Schaffner, 2012, unpub- molecular work on Leucanthemum vulgare lished results). and related species.

References

Anderson, R.S. and Korotyaev, B.A. (2004) Some Palaearctic weevils in the subfamily Ceutorhynchinae (Coleoptera, Curculionidae) recently discovered in North America. The Canadian Entomologist 136, 233–239. Bennett, B.A. and Mulder, R.S. (2009) Natives gone wild: climate change and a history of a Yukon invasion. In: Darbyshire, S.J. and Prasad, R. (eds) Proceedings of the 2008 Weeds Across Borders Conference, May 27–30, 2008, Banff, Alberta, pp. 235–248. British Columbia Ministry of Agriculture Food and Fisheries (2002) Guide to Weeds in British Columbia. Open Learning Agency Marketing Department, Burnaby, British Columbia. Burbank, L. (1914) The Shasta Daisy: how a troublesome weed was re-made into a beautiful fl ower. In: Whitson, J., John, R. and Williams, H.S. (eds) Luther Burbank: His Methods and Discoveries and Their Practical Application. Luther Burbank Press, New York and London, pp. 7–38. Chapter 51 341

Clements, D.R., Cole, D.E., Darbyshire, S., King, J. and McClay, A. (2004) The biology of Canadian weeds. 128. Leucanthemum vulgare Lam. Canadian Journal of Plant Science 84, 343–363. Crow, J.F. (2001) Plant Breeding Giants: Burbank, the Artist; Vavilov, the Scientist. Genetics 158, 1391–1395. Dieckmann, L. (1977) Beiträge zur Insektenfauna der DDR: Coleoptera–Curculionidae (Apioninae). Beiträge zur Entomologie 27, 7–143. Dieckmann, L. (1983) Beiträge zur Insektenfauna der DDR: Coleoptera – Curculionidae (Tanymecinae, Leptopiinae, Cleoninae, Tanyrhynchinae, Cossoninae, Raymondionyminae, Bagoinae, Tanysphyrinae). Beiträge zur Entomologie 33, 257–381. Greiner, R., Vogt, R. and Oberprieler, C. (2012) Phylogenetic studies in the polyploid complex of the genus Leucanthemum Mill. (Compositae, Anthemideae) based on cpDNA sequence variation. Plant Systematics and Evolution 298, 1407–1414. Guillet, G. and Arnason, J.T. (1995) Some phytophagous insects found on hirta and leucanthemum in the Ottawa/Hull area. Proceedings of the Entomological Society of Ontario 126, 95–97. Hawke, R.G. (2007) A Report on Leucanthemum × superbum and Related Daisies. Chicago Botanic Garden Plant Evaluation Notes Issue 30. Chicago Botanic Garden, Glencoe, Illinois. Lavoie, C., Saint-Louis, A., Guay, G., Groeneveld, E. and Villeneuve, P. (2012) Naturalization of exotic plant species in north-eastern North America: trends and detection capacity. Diversity and Distributions 18, 180–190. Manitoba Agriculture Food and Rural Initiatives (2012) Timothy Seed Production. Available at: https://www.gov.mb.ca/agriculture/crops/forages/bjb00s05.html (accessed 22 October 2012). Miller, G.L. and Stoetzel, M.B. (1997) Aphids associated with in the . Florida Entomologist 80, 218–239. Minister of Justice (2005) Weed Seeds Order. Available at: http://laws-lois.justice.gc.ca/PDF/SOR- 2005-220.pdf (accessed 22 October 2012). Oberprieler, C., Himmelreich, S., Källersjö, M., Vallès, J., Watson, L.E. and Vogt, R. (2009) Anthemideae. In: Funk, V., Susanna, A., Stuessy, T. and Bayer, R. (eds) Systematics, Evolution, and Biogeography of the Compositae. International Association for Plant , Vienna, pp. 631–666. Oberprieler, C., Eder, C., Meister, J. and Vogt, R. (2011) AFLP fi ngerprinting suggests an allopolyploid origin of two members of the Leucanthemum vulgare aggregate (Compositae, Anthemideae) in central Europe. Nordic Journal of Botany 29, 370–377. Olliff, T., Renkin, R., McClure, C., Miller, P., Price, D., Reinhart, D. and Whipple, J. (2001) Managing a complex exotic vegetation program in Yellowstone National Park. Western North American Naturalist 61, 347–358. Olson, B.E. and Wallander, R.T. (1999) Oxeye daisy. In: Sheley, R.L. and Petroff, J.K. (eds) Biology and Management of Noxious Rangeland Weeds. Oregon State University Press, Corvallis, Oregon, pp. 282–289. Otfi nowski, R., Kenkel, N.C., Dixon, P. and Wilmshurst, J.F. (2007) Integrating climate and trait models to predict the invasiveness of exotic plants in Canada’s Riding Mountain National Park. Canadian Journal of Plant Science 87, 1001–1012. Razowski, J. (2003) Tortricidae (Lepidoptera) of Europe: Olethreutinae, Vol. 2. František Slamka, Bratislava, Slovakia. Robinson, J. (2008) The evolution of fl ower size and fl owering behaviour in plants: the role of pollination and pre-dispersal seed predation. MPhil thesis, The University of Southampton, UK. Schaffner, U., Gundelwein, F., Grosskopf, G. and Häfl iger, P. (2008) Prospects for the biological control of oxeye daisy, Leucanthemum vulgare. Annual Report. CAB International, Delémont, Switzerland. Stoetzel, M.B. and Miller, G.L. (1999) Macrosiphoniella leucanthemi (Homoptera: Aphididae): new records and redescriptions of the apterous and alate viviparous females. Entomological News 110, 45–50. Strother, J.L. (2006) Leucanthemum Miller. In: Flora of North America Editorial Committee (ed.) Flora of North America North of Mexico, Vol. 19. Magnoliophyta: Asteridae, Part 6: Asteraceae, Part 1. Oxford University Press, New York, pp. 557–559. Stutz, S., Tateno, A., Hinz, H.L. and Schaffner, U. (2012) Annual Report 2011: Prospects for the biological control of oxeye daisy, Leucanthemum vulgare. CAB International, Delémont, Switzerland. 342 Chapter 52

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52 Linaria dalmatica (L.) Miller, Dalmatian Toadfl ax (Plantaginaceae)

Rosemarie A. De Clerck-Floate1 and Susan C. Turner2 1Agriculture and Agri-Food Canada, Lethbridge, Alberta; 2British Columbia Ministry of Forests, Lands and Natural Resource Operations, Kamloops, British Columbia

52.1 Pest Status et al., 2010), suggesting that it is still spreading and may become a serious weed Dalmatian toadfl ax, Linaria dalmatica (L.) in northern latitudes of Canada as the Mill. (Plantaginaceae), introduced to climate warms. eastern North America from Europe at the Linaria dalmatica continues to be of turn of the 19th century (Alex, 1962; greatest concern in western North America, Vujnovic and Wein, 1997), now occurs where this perennial invades rangelands, widely in Canada and the USA (Wilson et rights-of-way and natural areas. It is al., 2005; United States Department of currently listed as ‘noxious’ in British Agriculture, Natural Resource Conser- Columbia (British Columbia Ministry of vation Service, 2012). In Canada, there are Agriculture, 2012), Alberta (Alberta anecdotal reports of L. dalmatica being Agriculture and Rural Development, 2012) planted as an ornamental at the Central and Manitoba (Government of Manitoba, Experimental Farm in Ottawa, Ontario in 2012). Its shallow, creeping roots, early 1901 (Macoun, 1908; Alex, 1962), however, seasonal growth, prolifi c seed production the fi rst voucher specimen in Canada was (Lajeunesse et al., 1993; Vujnovic and not collected until 1933 in Edmonton, Wein, 1997), but also a high specifi c leaf Alberta (Alex, 1962). More recently, a fi rst- area (Maron and Marler, 2008) are thought time occurrence of L. dalmatica in the to contribute to the invasiveness and Yukon Territory has been reported (Bennett competitiveness of L. dalmatica. Other

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