The Biology of Canadian weeds. 150 Erechtites hieraciifolius (L.) Raf. ex DC.
Stephen J. Darbyshire1, Ardath Francis1, Antonio DiTommaso2, and David R. Clements3
1Agriculture and Agri-Food Canada, Eastern Cereal and Oilseed Research Centre, 960 Carling Ave., Wm. Saunders Bldg. #49, Ottawa, Ontario, Canada K1A 0C6 (e-mail: [email protected]); 2Department of Crop and Soil Sciences, 903 Bradfield Hall, Cornell University, Ithaca, NY 14853, USA; and 3Biology Department, Trinity Western University, 7600 Glover Road, Langley, British Columbia, Canada V2Y 1Y1. Received 10 January 2012, accepted 3 March 2012.
Darbyshire, S. J., Francis, A., DiTommaso, A. and Clements, D. R. 2012. The Biology of Canadian weeds. 150 Erechtites hieraciifolius (L.) Raf. ex DC. Can. J. Plant Sci. 92: 729 746. Erechtites hieraciifolius, American burnweed, is a herbaceous annual in the Asteraceae native to forest zones of eastern North America, but introduced to parts of Europe and the Pacific region. Confusion sometimes arises in distinguishing it from other weedy rayless composites in Canada, such as Senecio vulgaris (common groundsel) and Conyza canadensis (Canada fleabane). A pioneer therophyte species of forest zones, it occurs in large numbers when associated with major disturbances such as forest fires, but is also common in areas of smaller scale disturbance such as shores, forest edges and wind-throws. Soil conditions may vary greatly in nutrient content, moisture content, pH and salinity. In Canada, it is generally considered a minor garden and agricultural weed. It is most common in crops such as lowbush blueberry (Vaccinium spp.), cranberry (Vaccinium macrocarpon), strawberry (Fragaria ananassa) and vegetable crops; however, it occasionally occurs in other field and forage crops. Impacts include a variety of problems contributing to increased production costs or crop yield losses, including herbicide resistance, resource competition, and as a reservoir harbouring crop pathogens. It has been valued as a medicinal herb for treating a variety of ailments. Populations may grow rapidly in response to disturbances typical of managed landscapes, but E. hieraciifolius can usually be effectively controlled by chemical, cultural or manual weed control tactics.
Key words: Erechtites hieraciifolius, American burnweed, e´ rechtite a` feuilles d’e´ pervie` re, pilewort, weed ecology, weed biology
Darbyshire, S. J., Francis, A., DiTommaso, A. et Clements, D. R. 2012. La biologie des mauvaises herbes au Canada. 150 Erechtites hieraciifolius (L.) Raf. ex DC. Can. J. Plant Sci. 92: 729 746. L’e´ rechtite a` feuilles d’e´ pervie` re (Erechtites hieraciifolius) est une herbace´ e annuelle de la famille des Aste´ race´ es indige` ne aux re´ gions forestie` res de l’est de l’Ame´ rique du Nord, mais qui a e´ te´ introduite dans certaines parties de l’Europe et du Pacifique. On la confond parfois a` d’autres adventices a` rayons courts de la famille des Compose´ es qui poussent au Canada tels le se´ nec¸ on vulgaire (Senecio vulgaris) For personal use only. et la vergette du Canada (Conyza canadensis). Espe` ce pionnie` re the´ rophyte, on en retrouve de grand peuplements aux endroits qui ont subi de profondes perturbations comme un feu de foreˆt, cependant l’espe` ce colonise souvent des zones plus restreintes comme les rivages, la lisie` re des boise´ s et les chablis. Les conditions du sol peuvent varier conside´ rablement quant a` la concentration d’e´ le´ ments nutritifs, la teneur en eau, le pH et la salinite´ . Au Canada, on la conside` re ge´ ne´ ralement comme une adventice secondaire dans les jardins et les champs. L’e´ rechtite a` feuilles d’e´ pervie` re se rencontre le plus souvent dans les cultures comme le bleuet nain (Vaccinium spp.), la canneberge (Vaccinium macrocarpon), la fraise (Fragaria ananassa) et les plantes maraıˆche` res, cependant, on la de´ couvre a` l’occasion dans d’autres grandes cultures et des cultures fourrage` res. Elle engendre divers proble` mes qui concourent a` accroıˆtre les couˆts de production ou a` re´ duire le rendement, notamment la re´ sistance aux herbicides, la concurrence pour les ressources existantes et le fait de servir de re´ servoir aux agents pathoge` nes des cultures. L’espe` ce est prise´ e pour ses vertus me´ dicinales et on s’en sert pour soigner divers maux. Les peuplements prennent parfois rapidement de l’ampleur a` la suite des perturbations typiques au ame´ nagements paysagers, mais on peut habituellement venir a` bout de E. hieraciifolius par les moyens de lutte chimique, culturaux ou manuel.
Mots cle´s: Erechtites hieraciifolius,e´ rechtite a` feuilles d’e´ pervie` re, American burnweed, pilewort, e´ cologie des adventices, Can. J. Plant Sci. Downloaded from pubs.aic.ca by Agriculture and Agri-food Canada on 06/04/12 biologie des mauvaises herbes
hieracifolia (L.) Prov.; Neoceis hieracifolia (L.) Cass. Common names: American burnweed (Darbyshire et al. 1. Name and Generic Status 2000), eastern burnweed, fireweed (fire-weed), pilewort Erechtites hieraciifolius (L.) Raf. ex DC. * Synonyms (Darbyshire 2003), butterweed (Belcher 1956), white (spelling not corrected): Senecio hieracifolius L.; E. fireweed (Fogg 1945); e´rechtite a` feuilles d’e´pervie`re hieracifolia var. intermedia Fern.; E. praealta Raf.; E. hieracifolia var. praealta (Raf.) Fern.; Eriophthalmia Abbreviation: DALPT, days after late postemergence treatment
Can. J. Plant Sci. (2012) 92: 729 746 doi:10.4141/CJPS2012-003 729 730 CANADIAN JOURNAL OF PLANT SCIENCE
(Darbyshire et al. 2000), cre` ve-a` -yeux, cre` ve-z-yeux Belcher 1956; Gleason and Cronquist 1991; Barkley and (Darbyshire 2003). European and Mediterranean Cronquist 1978; Barkley 2006). Plant Protection Organization (Bayer) code: EREHI. Annual herb, (5 )50 200 ( 300) cm tall; taproot Asteraceae (Compositae): aster family Asterace´ es with fibrous secondary roots; stem erect, more or less (Compose´ es). strongly ribbed (grooved), simple or branched toward The name has often been spelled as ‘‘E. hieracifo- the top, glabrous to villose with translucent multicellular lia’’or ‘‘E. hieraciifolia’’. According to the International hairs; leaves alternate, the lower (3 )6 20 cm long Code of Botanical Nomenclature (McNeill et al. 2006), (1 )2 8 cm wide and diminishing in size upwardly, Erechtites hieraciifolius is correct, the genus being mostly glabrous or margins ciliate with short hairs and masculine in gender and an ‘‘i’’ being required between lower surface usually also with longer multicellular hairs the two elements (‘‘hieraci’’ and ‘‘folius’’) of the epithet (especially along the principal veins), irregularly toothed (i.e., a double ‘‘i’’). Augustin de Candolle (DC.) was and lobed, with secondary veins extending to the the first to publish the name E. hieraciifolius in 1838, marginal gland-tipped teeth (Fig. 2); lower leaves elliptic based on the earlier name and description, Senecio to broadly lanceolate, shallowly toothed to weakly acute hieraciifolius, by C. Linnaeus (1753). In doing so de lobed, usually weakly petiolate; middle to upper leaves Candolle acknowledged that C. S. Rafinesque (Raf.) lanceolate, acute, irregularly toothed, often deeply acute had suggested this to him in a letter; the nomenclatural lobed, sessile or clasping at the base, with distal leaves authority may be given as ‘‘Raf. ex DC.’’ (as above) or bract-like. simply ‘‘DC.’’ (McNeill et al. 2006). For a more The leaves are highly variable in size and shape. The complete list of synonyms see Belcher (1956). base may be attenuate to a distinct petiole (Fig. 1B; This account deals with E. hieraciifolius var. hieracii- Lloyd and Lloyd 1887), broadly sessile, or expanded and folius, the only variety present in Canada, and not with clasping to auriculate (Fig. 1A). Several leaf forms are the nearly pantropical weed E. hieraciifolius var. caca- often visible on the same plant and those with clasping lioides (Fisch. ex Spreng.) Griseb. (see Section 2c), unless or auriculate bases tend to be larger toward the middle explicitly mentioned. of the stem. Inflorescence varying from a single terminal capitu- 2. Description and Account of Variation lum (flower head) in depauperate specimens (Fig. 1C) (a) Description * The following description is of to compound, corymbose panicles usually with many E. hieraciifolius var. hieraciifolius, the variety present capitula (often 100 or more) on robust plants; capitula in Canada and most of the United States. It shows subcylindric or flask-shaped (slightly swollen at the considerable phenotypic variability (Belcher 1956), base), (6.5 )10 17 mm long, with a few linear especially in plant size and leaf shape (Fig. 1). The bracteoles at the base and upper pedicel (sometimes description here is based on material from Canadian collectively referred to as a calyculus); phyllaries
For personal use only. populations, supplemented with information from pub- (involucral bracts) about 20 in a single row, linear lished descriptions (e.g., Fernald 1917; Kummer 1951; with a slender attenuated tip, (6 )10 17 mm long and Can. J. Plant Sci. Downloaded from pubs.aic.ca by Agriculture and Agri-food Canada on 06/04/12
Fig. 1. Erechtites hieraciifolius: A. Drawing from Hermann (1705) with leaves lobed and clasping to auriculate (note lower leaves with attenuate bases); B. Drawing from Millspaugh (1887) showing upper stem with reduced dentate leaves and linear inflorescence bracts and bracteoles, a middle leaf with attenuate (more or less sessile) base; C. Diminutive mature plant with a single capitulum. DARBYSHIRE ET AL. * ERECHTITES HIERACIIFOLIUS (L.) RAF. EX DC. 731
centre (Fig. 3C); pappus of copious white hairs, about 5 10 times as long as cypsela, becoming readily detached (Fig. 3A, C); naked seeds (i.e., cypsela without the ovary wall) mostly dark blue to blackish with a whitish zone at the base, testa finely verrucose (Fig. 3D). As is typical in the Asteraceae, the dispersal unit (diaspore) is a cypsela, defined as a type of fruit formed from a single-seeded inferior ovary (Fig. 3C). The seed, technically defined as a ripened ovule, is left only when the ovary wall is removed (Fig. 3D). Except in Section 8C, where the distinction between fruit and seed becomes important, the term ‘‘seed’’ will be used to refer to the entire fruit. Cotyledons (Fig. 3B) round to oval, entire or faintly Fig. 2. Lower (abaxial) surface of leaf of Erechtites hieraciifo- lobed or emarginate, 6 9( 10) (3.5 )5 9 mm, lius showing secondary veins extending to the blade margins (somewhat obscure in the black and white version of this usually red-tinged beneath, a few scattered bristly short photo) and terminating in gland-tipped teeth. Translucent, hairs above and on the usually purple principal veins multicellular hairs present along the veins (and margins). Inset beneath, and a granular bloom on the lower surface. shows a single hair. Chromosome counts on plants of E. hieraciifolius at Ottawa, ON, were determined as n 20 by G. A. 0.5 1.5 mm wide, becoming strongly reflexed with age; Mulligan in 1966 (collection number 3118; voucher ligulate (ray) florets lacking but florets heterogamous, DAO 857169). This is consistent with published counts the outer florets pistillate, 10 100 , corollas whitish to of n 20, 2n 40 (Cooper 1936; Ornduff et al. 1963; yellow, tubular filiform, 4 5 deltate lobes, erect; disc Coleman 1982). florets 10 20 ( 50 ), mostly bisexual and fertile, inner sometimes functionally staminate, corollas whit- ish to pale yellow or pinkish, 4 5 erect to spreading (b) Distinguishing Features * In eastern Canada, Erech- deltate lobes; receptacle naked, 5 8.5 mm diam.; tites hieraciifolius is distinguished from most other cypselae (fruits) narrowly cylindric to oblong, 2 3mm composites by its many flowered (20 150), relatively long, brown with 8 12 paler ribs, strigose (rarely large (10 17 mm) capitula with the outer pistillate florets glabrous) between the ribs, the apex with a conspicuous lacking ligules (rays), and the small (2 3 mm long)
For personal use only. collar and persistent style base protruding from the seeds with copious white pappus hairs. Superficially, Can. J. Plant Sci. Downloaded from pubs.aic.ca by Agriculture and Agri-food Canada on 06/04/12
Fig. 3. Erechtites hieraciifolius: A. Seed with pappus of copious hairs; B. Seedling at two leaf stage; C. Cypselae, showing brown, ribbed and pubescent wall of fruit; D. Seeds, showing bluish-black, finely verrucose testa. 732 CANADIAN JOURNAL OF PLANT SCIENCE
it somewhat resembles the annual Canada fleabane, A third variety is present in the tropics, var. caca- Conyza canadensis (L.) Cronq., whose usually simple lioides, and found in the West Indies and from Mexico stalks reach comparable heights, and are topped by through Central America to northern Argentina and in branched inflorescences of numerous small capitula southeast Asia and the East Indies (Belcher 1956). This (about 3 4 mm long) with phyllaries of unequal lengths variety differs in its longer bracteoles which extend from and outer pistillate florets obscurely ligulate, and tiny one-third to one-half the length of the involucre, and by seeds (1 1.5 mm long). The stems of Canada fleabane the presence of multicellular hairs on the bracteoles and are covered with simple hairs bent upward (adpressed), phyllaries (Belcher 1956). the leaves are crowded and numerous, linear and In addition to treating megalocarpus and cacalioides attenuated in shape; the leaf margins, when not entire, as distinct species, Fernald (1917, 1950) recognized three are often more coarsely toothed with small teeth lacking intraspecific taxa within E. hieraciifolius in eastern glandular tips. North America [var. hieraciifolius ( var. typica), var. The widespread species Senecio vulgaris L. (common praealtus (Raf.) Fernald, and var. intermedius Fernald], groundsel) is another annual species which usually lacks based primarily on leaf size and shape. Few have ligulate florets. It may be distinguished from E. hier- adopted this intraspecific treatment pointing out that aciifolius by: its deeply lobed mid-stem leaves (appearing single populations may contain forms typical of all three coarsely pinnatifid), with the lobes obtuse rather than varieties (e.g., Deam 1940) and that leaf form and size acute; the black tips of the calycular bracts (and are affected by the growing conditions (Fogg 1945; sometimes also the phyllaries); smaller capitula about Belcher 1956). 5 8 mm long; and, the smaller seeds, 1.5 2( 2.5) mm long. The annual rayless aster [Brachyactis ciliata Ledeb.; Symphyotrichum ciliatum (Ledeb.) G. L. (d) Illustrations * Various aspects of E. hieraciifolius Nesom] also has small capitula (5 11 mm long) and are illustrated in Figs. 1, 2, and 3. Variation in leaf shape seeds (1.5 2.5 mm long), but there are multiple rows of is shown in the different elements of Fig. 1. Cypselae, phyllaries and the leaves are linear to oblanceolate with seeds and seedlings are illustrated in Fig. 3. Other useful entire and usually ciliate margins. illustrations may be found in Pammel et al. (1913), Fogg Several native species of Packera ( Senecio sensu (1945), Cronquist (1955), Muenscher (1955), Barkley lato) in eastern Canada may also be confused with E. (2006), Neal and Derr (2005), Jung and Chung (2010), hieraciifolius, such as the rayless P. indecora (Greene) Trewatha (2010), and USDA-NRCS (2011). A´ .Lo¨ ve & D. Lo¨ ve, and the occasionally rayless P. paupercula (Michx.) A´ .Lo¨ ve & D. Lo¨ ve. These per- 3. Economic importance ennial species have smaller seeds (52 mm), lower leaves (a) Detrimental * Various authors include it among that are distinctly spatulate with a long petiole (usually the agricultural or forestry weeds in Canada (e.g.,
For personal use only. as long as or longer than the blade), and the mid-stem Provancher 1862; Dalaire 1904; Clark and Fletcher leaves somewhat lyrate. 1909; Groh and Frankton 1949; Wright 1950; Darbyshire 2003), although it is a pest of secondary importance. It is a relatively minor weed in cranberry (c) Intra-specific Variation * This species is highly (Vaccinium macrocarpon Aiton) and blueberry (Vacci- polymorphic throughout its range (Fernald 1917, 1950; nium spp.) fields in Nova Scotia, Prince Edward Island Belcher 1956). Although several entities have been and New Brunswick where it is usually easily controlled variously recognized taxonomically, as varieties or with broad-spectrum herbicides or sometimes hand distinct species, the variation is continuous within and pulling (J. Calder, personal communication, Nova between the forms (Deam 1940; Belcher 1956; Seymour Scotia Department of Agriculture, Truro, NS; G. 1982). Most of the striking variation in plant size and Graham, personal communication, New Brunswick leaf size and shape is attributable to phenotypic Department of Agriculture, Aquaculture and Fisheries, responses to varying conditions (Belcher 1956). Two Fredericton, NB). Wild blueberry crops may be parti- North American varieties have been commonly recog- cularly susceptible to infestations since periodic burning
Can. J. Plant Sci. Downloaded from pubs.aic.ca by Agriculture and Agri-food Canada on 06/04/12 nized: var. hieraciifolius and var. megalocarpus (Fern.) is often used as a tool for pruning and weed manage- Cronq. The latter is a form found only in coastal salt ment (Penney et al. 2008). Data on labels of herbarium marshes of New England (Barkley 2006), which differs specimens collected in Canada indicate that it sometimes from the typical form in its more succulent leaves and occurs in a wide variety of other crops, including stems, a receptacle 8 mm in diameter, and larger seeds oat (Avena sativa L.), barley (Hordeum vulgare L.), (4 5 mm) with 16 20 nerves (Barkley 2006). Although maize (Zea mays L.), strawberry [Fragaria ananassa Fernald (1950) considered it a distinct species, other (Weston) Duch. ex Rozier], onion (Allium cepa L.) and authors generally treat it as ‘‘merely a well-marked carrot (Daucus carota L.), as well as fodder crops ecotype of saline coastal marshes’’ with the morpholo- (Medicago sativa L.) and mixed pastures. gical variation between the two forms being continuous It is sometimes considered a weed in the eastern (Cronquist 1946; Belcher 1956; Barkley 2006). United States (e.g., Pammel et al. 1913; Runnels and DARBYSHIRE ET AL. * ERECHTITES HIERACIIFOLIUS (L.) RAF. EX DC. 733
Schaffner 1931; Muenscher 1955). As in Canada, E. enough, and never interfering with crops, this is one of hieraciifolius is present in wild blueberry fields in our most useless and disreputable-looking weeds, very Maine (D. Yarborough, personal communication, Uni- unattractive in appearance, and ill-scented besides.’’ versity of Maine, Orono, ME) and cranberry fields in Vegetation management resources are not always ex- New Jersey (Anonymous, undated), where, in high pended on an economic basis and such perceptions of densities, plants can obstruct harvesting equipment ‘‘weeds’’ can have a tremendous impact on our relation- (D. Yarborough, personal communication, University ships with, and actions on, plants and landscapes. of Maine, Orono, ME). As an adventive outside its native distribution E. hieraciifolius var. hieraciifolius,is a weed in the Pacific region (see Section 4) and central (b) Beneficial * As a medicinal plant in North America, Europe (Belcher 1956; Voss 1996). It has also become a the use of E. hieraciifolius has had a complex and serious agricultural weed in Hawaii, invading sugar confusing history which Lloyd and Lloyd (1887) review cane (Saccharum officinarum L.) plantations in that extensively. These authors indicate that ‘‘fireweed oil’’ state (Hanson 1962; Sund 1964). Haselwood and of North American commerce was usually derived from Motter (1966) stated that, in Hawaii, it was, ‘‘One of other species (not E. hieraciifolius), but maintain that E. the most abundant weeds in moist cultivated areas; hieraciifolius oil has distinct benefits. Erichsen-Brown also a weed in pastures and rangelands.’’ Although (1979) cited a number of early North American sources E. hieraciifolius is naturalised on all of the major on medicinal uses of the plant in treatment for Hawaiian islands (HEAR 2008), it is currently not haemorrhage, wounds, skin diseases, dysentery, cholera, as abundant as E. valerianifolius (Link ex Spreng.) and as a purgative and emetic; as a source of a blue dye DC. (F. and K. Starr, personal communication, US for cotton and wool; and, its use by Algonquin native Geological Survey, HI). peoples to treat poison ivy [Toxicodendron radicans (L.) It has been reported a weed in pine plantations in the Kuntz] and poison sumac [T. vernix (L.) Kuntz] southern United States (Britt et al. 1990; Gardiner et al. poisoning. Hale (1880) recommends its use in the 1991); as an increasingly important reservoir host for treatment of dysentery, menstrual disorders and gonor- crop pathogens including insects, fungi, nematodes and rhoea. Millspaugh (1887) considered the plant’s volatile viruses (see Section 13a, b); as a weed of cultivated lands oil to be the principal active ingredient in a tincture (Runnels and Schaffner 1931; Trewatha 2010); as a weed produced from whole fresh flowering plants for use as of pastures and forage grasses (Pammel et al. 1913; an emetic and in treatment of such conditions as eczema, Muenscher 1955; Stephenson and Rechcigl 1991), and diarrhoea, haemorrhages and piles, but noted that turf grass (Busey and Johnston 2006; Trewatha 2010). nausea and other adverse reactions could result from It is also reported as a problem weed in the container use of this tincture. In the early 20th century its use as a nursery industry (Neal and Derr 2005). medicinal herb was sufficiently popular that collectors
For personal use only. It is considered a potential weed in Australia where it were paid 2 3 cents per pound (Georgia 1914). is under quarantine (AQIS 2011). In Hungary it is Many authors comment on the unpleasant or rank considered an invasive pest (Csisza´ r 2006). odour of the plant. In spite of this Fernald et al. (1958) The presence of alkaloids (see also Section 7c) causes suggest it could be used as a salad or potherb. The hepatotoxicity if the plants are consumed in sufficient plant’s odour does not seem to be consistently distaste- quantity (Burrows and Tyrl 2001). The plant is fre- ful to all people. quently reported to have a noxious smell and taste, so it In the Andes of South America the leaves and flowers is unlikely that persons or livestock would voluntarily (var. cacalioides) have been used in folk medicine as a consume sufficient quantities to be at serious risk. blood depurative and the roots to treat cardiac disease However, the overall similarity between E. hieraciifolius (Lorenzo et al. 2001). The antioxidant and radical and some even more toxic Senecio species may cause scavenging properties of nine species of Asteraceae difficulties through pest mis-identification and inap- from Bolivia were investigated by Parejo et al. (2003); propriate management decisions. although some activity was exhibited by E. hieraciifolius, As implied by the French common name ‘‘cre` ve-z- other species appeared more promising as sources of
Can. J. Plant Sci. Downloaded from pubs.aic.ca by Agriculture and Agri-food Canada on 06/04/12 yeux’’, the pappus hairs were thought to pose a serious compounds for industrial and medical use. threat to the eyes (Provancher 1862). The flocculent In Japanese trials, E. hieraciifolius has been found to pappus bristles may well cause eye irritation when be among the most efficient of the plants tested at manually threshing grain crops heavily contaminated assimilating atmospheric nitrogen dioxide (NO2)withas with E. hieraciifolius, especially in relatively stagnant air much as 10% of its total organic nitrogen content spaces. The pappus bristles have scattered antrorse derived from this source (Morikawa et al. 1998). Such spicules which may contribute to the abrasion of soft plants have the potential to act as important sinks for tissues such as eyes and lungs. anthropogenic nitrogen oxides (Morikawa et al 1992; From a sociocultural perspective, Harper (1944) Morikawa 1996). Morikawa et al. (2003) proposed that indicates that it is greatly offensive to aesthetic sensi- ‘‘green walls’’ using such nitrogen dioxide-philic plants bilities, saying that, in Alabama, ‘‘Although harmless could be set up around buildings and on highway 734 CANADIAN JOURNAL OF PLANT SCIENCE
corridors to help sequester pollutants from car emissions E. hieraciifolius in Canada, but data on herbarium or other sources. specimen labels indicate the species will grow in a wide range of substrate conditions. Soil types reported are rather general, but include sand, clay, loam, rock, and (c) Legislation * Erechtites hieraciifolius is not listed in gravel. It is found in moist to saturated soils of shores, any federal or provincial noxious weed legislation in flood plains, fresh and salt marshes, bogs, dune slacks, Canada; and is similarly absent from federal and state gravel bars and ditches, but also occurs in dry or well- lists in the United States. The state of New York has drained substrates such as dry sands, talus slopes, and listed E. hieraciifolius var. megalocarpus as endangered granite balds. A wide range of substrate pH is tolerated in that state (USDA-NRCS 2011). Australia restricts as indicated by the collections from peat bogs to importation of E. hieraciifolius where it is considered a limestone cliffs and talus. Although nutrient poor soils potential threat and is regulated as a quarantine pest are readily colonized, Wilson and Shure (1993) found E. (AQIS 2011). hieraciifolius biomass production was greater at har- vested forest sites to which fertilizer was added. 4. Geographic Distribution More detailed analysis of soil types is found in some In North America, Erechtites hieraciifolius var. hieracii- United States sources. The substratum in a pine forest in folius is found primarily in deciduous forest regions of South Carolina, where E. hieraciifolius was among early eastern Canada from the Maritime Provinces to western successional species, consisted of a silty sand down to Ontario, and in the United States from New England 1.2 m underlain by a medium to fine clayey sand to 7.6 west to Minnesota in the north, and south to Florida m; and the upper soil layers averaged 87% sand, 6.4% and eastern Texas. The distribution in Canada is shown silt and 6.3% clay-sized particles (Hunt and Shure in Fig. 4, based on 837 specimens in Canadian herbaria. 1980). Following treatment of the soil with waste water References to its presence in Saskatchewan or Alberta in the above area, soil organic matter decreased from 3.5 are almost certainly erroneous (Boivin 1972; Harms to 1.5% and cation exchange capacity from an average 2006), and probably based on the report by Hooker of 55 meq 100 g 1 in the top 5 cm to 28 meq 100 g 1 at (1834). No specimen was seen to confirm literature 10 cm and 10 meq 100 g 1 below 15 cm; but there were reports by various authors (e.g., Gleason and Cronquist otherwise no differences in the soil profile of treated and 1991) of occurrence in Newfoundland (Meades et al. untreated areas except for an increase in soil moisture 2000; Barkley 2006). It has been reported as introduced from 8% in untreated areas to up to 15% in treated to Washington State, apparently based on a single areas (Hunt and Shure 1980). Forest top-soils in eastern collection from Seattle (Cronquist 1955), and as spor- Tennessee containing substantial seed banks of E. adically adventive on the West Coast (Barkley and hieraciifolius were found to have 3.2 5.3% organic Cronquist 1978; Barkley 2006). It is not known to occur matter, 4 11 mgg 1 phosphorus, 21 26% clay, 28
For personal use only. in British Columbia (Douglas et al. 1998; Barkley 2006). 43% silt and 31 51% sand, with a pH of 5.2 6.0 Elsewhere in the world, E. hieraciifolius var. hieracii- (Farmer et al. 1982). The predominant soil types in folius is known from the West Indies (Belcher 1956) and eastern mixed forest areas where E. hieraciifolius has been introduced to central Europe (Belcher 1956; appears following fires or forest clearance are loams Tutin 1976), Hawaii (Belcher 1956), Japan (Walker and silt loams originating from acid shales, sandstones 1976; Kamada and Nakagoshi 1990), China (Chien and limestones, often acidic with an average soil pH of and Ku 1998), Taiwan (Wu et al. 2004; Jung and Chung about 4.5 (Coxe et al. 2006). In the wetland complex of 2010), Korea (Shin and Braun 2000), and New Zealand the Delaware River (NJ), E. hieraciifolius was found in (de Lange 1995). shrub forest habitats on alluvial and aeolian deposits (Leck et al. 1988). 5. Habitat (a) Climatic Requirements* Climatic conditions within the native distribution of E. hieraciifolius var. hieracii- (c) Communities in Which the Species Occurs * In folius vary from humid continental to maritime. The Canada, E. hieraciifolius occurs in boreal (southern
Can. J. Plant Sci. Downloaded from pubs.aic.ca by Agriculture and Agri-food Canada on 06/04/12 plant extends over a wide latitudinal range in eastern part), Great Lakes/St. Laurence, Acadian and Caroli- North America, from southern boreal (9498N) to nian forest regions (Fig. 4). The species usually occurs in subtropical regions with considerable variation in aver- abundance and high density in places of recent major age temperature and day length hours. In Canada, it is disturbance, although small scale disturbances giving not known from areas with a growing season of less rise to suitable microsites are colonized by small than about 160 d, or average precipitation of less than numbers of plants. Disturbances may result from fires about 70 cm. (Masters et al. 1996; Nakagoshi et al. 2003; Hutchinson et al. 2005), forest cutting (Boring et al. 1981; Masters et al. 1996; Landenberger and McGraw 2004; Mou et al. (b) Substratum * There is little information in the 2005; Abella 2010), herbicide application (Kochenderfer literature on the edaphic requirements and tolerances of and Wendel 1983; Luken et al. 1994), cultivation DARBYSHIRE ET AL. * ERECHTITES HIERACIIFOLIUS (L.) RAF. EX DC. 735
Fig. 4. Map of distribution of Erechtites hieraciifolius in Canada, based on 837 herbarium specimens from ACAD (35), CAN (72), DAO (120), HAM (27), LKHD (2), MT (146), MTMG (37), NSAC (7), OAC (24), QFA (142), QK (27), QUE (61), TRT (75), TRTE (29), UNB (10), WAT (16), and WLU (7). Herbarium abbreviations follow Holmgren et al. (1990).
(Fig. 5), flooding or hydrological changes (Shull 1914; the water level subsided (herbarium specimen label Leck et al. 1988; herbarium specimen label data), data, OAC). erosion (Peterson et al. 1990), or other disturbances Smaller scale disturbances also provide microsites (Wagner 1966; Hunt and Shure 1980; Roman et al. where reduced competition within a broader commu- 1984). In hemlock [Tsuga canadensis (L.) Carrie` re] nity allows small populations of E. hieraciifolius to forests in New England damaged by an imported establish; e.g., tree falls and wind-throws (Peterson and adelgid [Adelges tsugae (Annand)], it is among oppor- Pickett 1990; Peterson et al. 1990; Peterson and Pickett tunistic species said to have ‘‘invaded’’ the stands (Orwig 1995). and Foster 1998). It is a poor competitor (Boring et al. 1981) and Peterson et al. (1990) found that it 6. History was negatively correlated (r 0.45, P 0.018) with As a native plant of deciduous and mixed forest areas, total cover (primarily other plant species). E. hieraciifolius has presumably been present in those habitats since the re-vegetation of de-glaciated areas of For personal use only. Population explosions may occur under these and other conditions where competition is reduced, but southern Canada after the end of the last Ice Age. In the generally subside rapidly with successional progression early 19th century, when clearing of the great eastern to less open environments. Shores of lakes and rivers, forests of North America had begun in earnest, E. habitats characterized by natural disturbance, are hieraciifolius had become known as ‘‘fire-weed’’ because of its abundant occurrence in areas of forest clearance, common locations for E. hieraciifolius as are sites especially where fires had occurred (Pursh 1814; Eaton associated with beaver activities causing cyclical flood- 1824; Torrey 1843). William Barton (1818) said that it ing and drainage. It is found in a wide range of was ‘‘one of the commonest weeds [around Philadel- anthropogenic habitats where vegetation is regularly phia], growing almost every where, even on roofs’’. disturbed or maintained at early successional stages, such as roadsides, railway lines, ditches, energy corri- 7. Growth and Development dors, quarries, tree plantations, cultivated land, turf, (a) Morphology* All annual plants are considered flower beds, or ruderal sites. It grows in coniferous, therophytes, and this life-form is highly developed in deciduous or mixed forests, although usually in places
Can. J. Plant Sci. Downloaded from pubs.aic.ca by Agriculture and Agri-food Canada on 06/04/12 E. hieraciifolius. The species can rapidly respond to receiving some direct sunlight such as openings, edges, take advantage of favourable growth conditions, rapidly or along trails. A variety of wetland habitats are also producing large individuals in dense populations suitable, including shores, wet thickets, wet forests, wet (Fig. 5). The phenotypic plasticity of its growth also prairies, marshes, bogs and fens (Fernald 1950; Roman promotes maximum reproductive effort under sub- et al. 1984; Gleason and Cronquist 1991; Voss 1996; optimal conditions. Depending on the availability of Murphy et al. 2009; herbarium specimen label data). A moisture, nutrients and light, plants may reach maturity 1975 collection of E. hieraciifolius in southern Ontario at heights from a few centimetres and bearing only a came from a Spaghnum moss (Spaghnum sp.) mat single capitulum, to more than 2 m with a few hundred around a lake which had previously been flooded and capitula (Fogg 1945; Csisza´ r 2006; Fig. 1). The plant is was then invaded by ‘‘hundreds of these weeds’’ after capable of considerable morphological plasticity (see 736 CANADIAN JOURNAL OF PLANT SCIENCE
(1944), was later shown to be a mixture of two compounds, senecionine and seneciphylline (Adams and Gianturco 1956). Bohlmann and Abraham (1980) found a syringyl alcohol derivative and sesquiterpenes in the roots of E. hieraciifolius. In the re-growth following a forest clear-cut in North Carolina, E. hieraciifolius was found to contain 2.699 0.05, 0.2590.01, 6.9790.75, 0.9890.05, and 0.3790.03 percent dry mass, N, P, K, Ca, and Mg, respectively (Boring et al. 1981). The N, P, and K values were the highest detected among all herbaceous species and most woody species (significantly higher values of N were detected only in the sprouts of two tree species). An analysis of oil extracted from aerial parts of E. hieraciifolius in Brazil found that 11 compounds con- stituted 94% of the oil, a-phellandarene (41.3%) and r-cymene (22.2%) being the main components (Lemos et al. 1998). Somewhat different results were obtained from an analysis of fresh leaves and stems of plants from Bolivia (Lorenzo et al. 2001). In the latter analysis, 22 components were identified as composing 93.3% of the oil, of which a-pinene was the main component (48%), followed by myrcene and (E)-b-ocimene (14% each), but only traces of a-phellandrene and r-cymene were found. Nutrients leached from mature leaves of E. hieracii- folius subjected to simulated rain were at rates of: 0.03 2 1 mg 0.1 m h NH4 (ammonium ion); 2.4 K; 1.68 Ca; 0.31 Mg; 0.26 NO3 (nitrate ion); and 0.93 P (Haines et al. 1985a). No significant difference in the rate of nutrient leaching was detected at different treatment pH levels (2.5 5.5). Phosphorus was appar- Fig. 5. A dense population of Erechtites hieraciifolius in New ently absorbed from the simulated rain water rather
For personal use only. Jersey, 21 August 2008, Mr. Peter Craig in photo. The 1.6-ha than leached. The relatively high rate of nutrient field was treated with two applications of glyphosate (about a leaching from the leaves was attributed to the wett- 1-month interval), and then no-till planted with a variety of ability of adaxial leaf surfaces (Haines et al. 1985b) (see native grassland species. It is unknown whether the growth of Section 11). Tissue necrosis resulted from exposure to E. hieraciifolius was the result of a soil seed bank, seed rain pH levels below 2.5 (Haines et al. 1980). from an outside source, or contamination of the planted seed. The complex nitrogen metabolism has been exten- sively studied by Japanese researchers. Of the 217 plant Section 2) allowing a wide range of growth responses to species studied in laboratory experiments, Morikawa environmental conditions. et al. (1998) observed that E. hieraciifolius had one of the 1 highest NO2 assimilation rates at 5.7 mg g dry weight, representing 10% of the total nitrogen content. Average (b) Perennation * A typical therophyte, E. hieraciifolius levels of nitrate (NO3 ); nitrite (NO2 ) and ammonium is a ‘‘summer’’ annual over-wintering only as seeds. It (NH4 ) ions detected in the leaves of seedlings of E. has been postulated that some plants may behave as hieraciifolius by capillary electrophoresis were 16.8
Can. J. Plant Sci. Downloaded from pubs.aic.ca by Agriculture and Agri-food Canada on 06/04/12 winter annuals in southerly regions where winters are mmol, 8.2 nmol and 1.0 mmol per gram of fresh tissue, milder than in Canada. In Kentucky, Baskin and Baskin respectively (Kawamura et al. 1996). Of the six plant (1996) found that 1% of freshly collected seeds germi- species studied by Kawamura et al. (1996), E. hieracii- nated immediately in an unheated greenhouse. They also folius contained the smallest quantity (in relation to wet observed some plants of such a large size that it seemed weight) of NH4 , and the largest quantity (in relation to unlikely their growth could have been attained from chlorophyll content) of NO2 : The ability of E. hier- spring germinating seeds. aciifolius to convert NO3 to nitrous oxide (N2O) was demonstrated by Hakata et al. (2003), although it was one of the least efficient of the 17 species studied. The (c) Physiology and Biochemistry * The alkaloid hier- capacity of various plant species to convert NO3 to acifoline, reported from E. hieraciifolius by Manske N2O (i.e., N2O emission) appears to be inversely DARBYSHIRE ET AL. * ERECHTITES HIERACIIFOLIUS (L.) RAF. EX DC. 737
proportional to their ability to assimilate NO2 (Hakata either no viability (Telewski and Zeevaart 2002) or as et al. 2003). much as 89% germination after 8 yr (Baskin and Baskin 1996). Some studies on germinating or recovering buried seeds from forest soils have found few seeds (Oosting (d) Phenology * Seeds mostly germinate in the spring and Humphreys 1940; Livingston and Allesio 1968; (see Section 8c) and somewhat later with increasing Leck et al. 1988; Matlack and Good 1990; Schiffman latitude. Flowering may begin as early as late July, and Johnson 1992), in spite of the potential for but seed maturation occurs mostly through mid-August immigration of wind-borne seeds. The low numbers of to early October (herbarium specimen label data). seeds and distributional patchiness in mature forest soils Robertson (1928) reported the flowering period in found by these studies suggests that long-term seed Illinois as Aug. 20 to Sep. 25. The basal leaves often banks are not formed. However, other studies have begin to wither as flowering commences (Barkley 2006). detected large numbers of buried E. hieraciifolius seeds germinating from forest soils (Farmer et al. 1982; Mou et al. 2005; Schelling and McCarthy 2007). Differences (e) Mycorrhiza * The roots of E. hieraciifolius plants in the results of studies measuring germination may be germinated from forest topsoils (from eastern Tennes- due in part to experimental conditions and the cyclical see) were extensively colonized by endomycorrhizae conditional dormancy exhibited by E. hieraciifolius with hyphae and vesicles present in all samples (Farmer (Baskin and Baskin 1996). Direct observation of ‘‘seeds’’ et al. 1982). in soil samples may be obfuscated by the rapid disin- tegration of the brown, ribbed fruit wall of the cypsela 8. Reproduction which reveals a finely verrucose, dark blue to blackish (a) Floral Biology * The capitula of E. hieraciifolius testa (Baskin and Baskin 1996; Fig. 3D). Although still lack ligulate florets to attract pollinators and are viable, the naked seeds (ripened ovules) are very probably primarily autogamous. Thirteen species of different in appearance and can be easily overlooked mostly hymenopteran insect visitors were reported by or mis-identified. Robertson (1928) in Illinois; however, they are unlikely Few studies have been conducted in Canada. The to be effective pollinators. plant did not germinate from any of the soil samples The cytology, histology and ontogeny of micro- and from mature deciduous forests at Mont St. Hilaire, QC megasporogenesis, and early embryo development in E. (Leckie et al. 2000), although herbarium specimens have hieraciifolius were studied by Cooper (1936). Wagner been collected indicating its presence throughout the (1966) studied the effects of radiation on embryo area (herbarium specimen label data). abortion rates (100% atB1.5 Gy d 1) in a field study Profuse germination of E. hieraciifolius is often noted in Brookhaven, NY. in conjunction with water draw-downs, such as drained
For personal use only. beaver ponds, emergent shorelines and flood plains (herbarium specimen label data). At Cold Spring (b) Seed Production and Dispersal * The pappus of Harbor, NY, abundant germination of E. hieraciifolius, copious large hairs at the apex of the seed (Fig. 3A) along with more than 140 other species of plants, was facilitates wind dispersal (Fogg 1945; Ohtsuka 1998). noted on the bed of a recently drained millpond (Shull Large plants are capable of producing thousands of 1914). Shull (1914) tried to artificially recreate similar seeds (see Section 2). In Hungary, Csisza´ r (2006) found conditions in the laboratory, but did not observe any that an average of 32 390 seeds were produced per plant, germination of E. hieraciifolius. In a Connecticut salt although growing conditions were not indicated. In a marsh, Roman et al. (1984) found that E. hieraciifolius forest area of NY, Wagner (1965) measured the seed germinated well in areas where the soil water salinity rain of E. hieraciifolius from late summer to early was near 0.5% throughout the growing season. autumn at 36 145 seeds ha 1. In North Carolina, Oosting and Humphreys (1940) In agricultural settings, E. hieraciifolius may also be examined seed germination from soil samples along a dispersed through mechanical harvesting equipment plant succession transect from cultivated fields to
Can. J. Plant Sci. Downloaded from pubs.aic.ca by Agriculture and Agri-food Canada on 06/04/12 (J. Calder, personal communication, Nova Scotia De- climax oak-hickory forest; 10 sites were considered to partment of Agriculture, Truro, NS; D. Yarborough, represent more than 200 yr of forest growth. No seeds personal communication, University of Maine, Orono, of E. hieraciifolius germinated from soil collected at ME). sites representing pre-afforestation communities, and only small numbers of seedlings were detected at forested sites. Although no data are given on the (c) Seed Banks, Seed Viability and Germination * The above-ground presence of E. hieraciifolius in any of results of various observations and experiments on seed the vegetation age classes, Oosting and Humphreys bank formation of E. hieraciifolius are incongruous and (1940) state that it is improbable that seeds immigrating difficult to interpret. Germination tests of exhumed from elsewhere would be able to readily penetrate litter seeds, artificially buried for long periods, have shown layers into the soil layers sampled. Livingston and 738 CANADIAN JOURNAL OF PLANT SCIENCE
Allessio (1968) reported similar results along a succes- communities in West Virginia, E. hieraciifolius seeds sional transect in Massachusetts from abandoned fields, germinated at rates of 0 55 seeds m 2 (McGraw 1987). through conifer plantations, to mixed woodland; only Unfortunately data were not presented on the vertical 4 seedlings were germinated from a total of 16 soil distribution (and therefore relative age) of germinating samples, one each from 4 mid- to late-succession seeds. At an upland site in West Virginia, Landenberger samples. In a southern Appalachian mature oak forest and McGraw (2004) studied seed banks at two forest (Montgomery Co., Virginia) with a low level of past sites which had been clear-cut 7 yr previously. They disturbance, E. hieraciifolius was among pioneer species found E. hieraciifolius to be among the most abundant that were either absent or present at very low densities seeds (relative to other species detected) in the soil in the (50.03 m 2) in the seed bank (Schiffman and Johnson open clear-cut and that seed densities declined in 1992). At sites where seeds were found in the soil (by proportion to other species and logarithmically in germination testing, but undetected in visual inspec- density across boundary between clear-cut and undis- tions), adult plants were also found in the forest turbed forest. community; however, no germination of E. hieraciifo- Baskin and Baskin (1996) found that in Kentucky lius was observed in soil samples from sites which about half of the freshly collected seeds were dormant lacked adult plants (suggesting limited immigration). At and the remainder only germinated under warmer disturbed and undisturbed mixed oak forest sites in the conditions. Unburied seeds which were winter-stratified central Appalachians (southeastern Ohio), Schelling germinated the following spring, while buried seeds (also and McCarthy (2007) germinated seeds of E. hieracii- stratified), germinated the April following exhumation. folius from 79% of their soil samples, where it occurred Buried seeds re-entered conditional dormancy each most frequently among the 70 species detected in the October, but 89% of seeds remained viable after 8 yr seed bank. Adult plants were also found in the above- of burial. Maximum germination occurred under ther- ground vegetation in the study area. A study of several moperiods of 30:15 and 35:208C (12:12 h) and a forest soil seed banks in eastern Tennessee (Farmer photoperiod of 14 h light:10 h dark, although consider- et al. 1982) found that E. hieraciifolius was one of five able germination also occurred in total darkness. In species germinating from 100% of the samples and Massachusetts, Lincoln (1983) found that 85% of trials, that it produced the second highest number of freshly harvested seed germinated in either light of seedlings (mean of 127 425 ha 1) of the 95 species dark conditions at 20 308C within 14 d following a detected, and, after 5 6 mo of growth, represented 60-d cold treatment. about 32 42% (depending on substrate trial) of the Although the vegetative community at the site of soil total biomass of all germinated plants. Germination provenance is not given, E. hieraciifolius germinated tests on litter and soil (to 5 cm depth) from a pine from most of the 96 pots using soil from Ona, Florida, (Pinus elliottii Englem.) plantation in South Carolina, in a greenhouse study (Stephenson and Rechcigl 1991). 2
For personal use only. found seedling densities of 26 m (Mou et al. 2005), It is not known whether germination was, in whole or in with almost all seedlings germinating from unscarified part, from fresh (previous season) or dormant (seed- soil samples. banked) seed, or whether seeds were from locally grown The unsuitability of shade conditions in mature plants or transported from elsewhere. Germination of closed-canopy forests, the scarcity of seeds in the litter seed tended to increase with soil pH (4.5 5.7) when and soil, and the rapid germination of E. hieraciifolius augmented with dolomite, reaching a maximum at pH following cool burns and litter removal might suggest 5.7 and then diminishing somewhat at pH 6.1 6.8 that some seeds will remain dormant in the substratum achieved with gypsum addition. Also, plants growing until surface conditions permit access to sunlight at low soil pH (B5.2) were smaller and less vigorous. (Glasgow and Matlack 2007), but might also suggest immigration of wind-borne seeds from neighbouring open areas (Matlack and Good 1990). For example, a (d) Vegetative Reproduction * Erechtites hieraciifolius study of buried seeds after clear-cutting of an ancient reproduces only by seed (Muenscher 1955). stand of Pinus strobus L. in Connecticut found that E.
Can. J. Plant Sci. Downloaded from pubs.aic.ca by Agriculture and Agri-food Canada on 06/04/12 hieraciifolius was not among plants germinated from 9. Hybrids buried seeds, and that its substantial presence in the No inter-specific hybrids have been reported. clearing arose from wind-dispersal of its seeds from open areas (Del Tredici 1977). Ecologists have come 10. Population Dynamics to similar conclusions in Japan, where its explosive The rapidity with which large numbers of plants of E. occurrence as an early pioneer of post-disturbance hieraciifolius appear immediately after major distur- forest succession is attributed primarily to anemo- bance, then quickly begin to decline over the following chory (wind dispersal) rather than seed banking 2 or 3 yr, represents a classic example of an early pioneer (Ohtsuka 1998). species in the ecological succession of plants. Pursh Most studies of soil seed communities have been (1814) noted the tendency of this plant to rapidly form conducted at upland sites. In a variety of wetland (bog) extensive monocultures in recently cleared forest areas, DARBYSHIRE ET AL. * ERECHTITES HIERACIIFOLIUS (L.) RAF. EX DC. 739
particularly where the woody debris was burned in situ. frequency) of E. hieraciifolius was 0.6 in un-thinned Farmer et al. (1982) observed densities of 49 220 103 control plots and 9.5 in plots 3 yr after thinning (Abella plants ha 1 when testing for seed germination from 2010). In a managed pine plantation in South Carolina, southern Appalachian forest soils. When they measured Mou et al. (2005), found the high importance value of E. the above-ground biomass 5 6 mo after germination hieraciifolius (6.3) attained in the first post-harvest (representing plant growth in the first season post- growing season declined rapidly in the following 4 yr disturbance), E. hieraciifolius comprised 32 42% of the (3.8, 0.5, 0, and 0.2, respectively). total plant biomass, more than twice as much as produced any of the other 94 species under any of the 11. Response to Herbicides and Other Chemicals soil condition treatments. In the first year of re- The leaf adaxial epidermis in E. hieraciifolius is essen- colonization after total vegetation removal by herbicide tially glabrous, lacks epicuticular wax, has a low water in a West Virginia forest, E. hieraciifolius comprised drop contact angle (about 708), and has a high water- 22% of the ground cover (Kochenderfer and Wendel holding capacity (Haines et al. 1985b). This wettability 1983). In the second year it had fallen to 2% cover and of the leaf surface promotes retention and penetration, was not detected in subsequent years. In a North and therefore effectiveness, of spray-applied foliar Carolina clear-cut forest, Boring et al. (1981) found herbicides. that E. hieraciifolius biomass production was 59 kg ha 1 Little work has been done in Canada to assess the during the season following cutting (i.e., removal of all response of E. hieraciifolius to herbicides. A single post marketable timber), which represented 13.6% of herbac- application of mesotrione at 101 g a.i. ha 1 early in the eous plant biomass and 3.4% of the total biomass season has provided good control of E. hieraciifolius in production by all woody and non-woody plants. It was New Brunswick cranberry fields (G. Graham, personal the only species making a substantial contribution to the communication, New Brunswick Department of Agri- re-growth biomass which was not detected in pre-cut culture, Aquaculture and Fisheries, Fredericton, NB). vegetation surveys. At a nearby study site, Elliott et al. Mesotione also controls E. hieraciifolius well in wild (1998) reported that, in the first year of revegetation blueberry fields in Maine (D. Yarborough, personal after a severe disturbance, E. hieraciifolius biomass communication, University of Maine, Orono, ME), production was 1820 kg ha 1 and represented 37.1% although application rates are usually higher than in of the biomass of the ground flora. A subsequent survey Canada. 15 yr later did not detect any E. hieraciifolius, but after The species was among the first plants for which 28 yr it was detected as a minor component (0.9 kg herbicide tolerance was reported. After less than 10 yr of ha 1, 0.2%). The occurrence of E. hieraciifolius in the heavy use of 2,4-D in sugar cane fields of Hawaii, more mature forest stage may be due to the creation of populations of E. hieraciifolius were showing reduced suitable microsites by local disturbances (Peterson et al. response to the ‘‘new’’ chemical control (Hanson 1956,
For personal use only. 1990; Peterson and Pickett 1990). 1962). At a woodland site in South Carolina, highly dis- Sulfometuron/hexazinone applied in granular form turbed because of the regular spray disposal of waste- was effective in early season control in pine plantations water, E. hieraciifolius growth was correlated to the (Pinus taeda L.) in the southeastern US (Gardiner et al. amount of disturbance (Hunt and Shure 1980). In the 1991). Sixty days after a late postemergence granule high disturbance zone it reached a peak biomass application of sulfometuron hexazinone at two rates of about 125 g m 2 in June, whereas in the lower (140.1 g a.i. ha 1 700.5 g a.i. ha 1 and 168.1 g a.i. disturbance zone a peak of about 50 g m 2 occurred in ha 1 840.6 g a.i. ha 1, respectively) 94% control of August. As an important component of the plant the two major herbaceous weed species [E. hieraciifolius community and insect host, the growth of E. hieraciifo- and Eupatorium capillifolium (Lam.) Small] was ob- lius had a significant effect on the temporal patterns and tained at a pine plantation site in Arkansas. By 120 days productivity of arthropod herbivores (see Section after a late postemergence, control from the two 13aiii). application rates decreased to 70 and 80%, respectively. It is not clear whether the colonizing ability of E. Several broad-spectrum specialty herbicides have
Can. J. Plant Sci. Downloaded from pubs.aic.ca by Agriculture and Agri-food Canada on 06/04/12 hieraciifolius is primarily the result of seed dispersal and been registered in the United States which are claimed immigration or long-term seed banking (see Section 8c). to control E. hieraciifolius in turf or other ornamental For example, seeds of E. hieraciifolius have been shown plantings. A mixture of clopyralid and triclopyr at a to be short-lived or transitory in mixed-oak hardwood recommended rate of 38 g a.e. ha 1 and 51 g a.e. ha 1, clearings in the southern Appalachians where they are respectively, is registered for postemergence control in abundant in the early-successional period, but rapidly turf (Dow AgroSciences 2008). A mixture of dimethe- decline and are replaced by more shade-tolerant species namid-P and pendimethalin at a recommended rate of with the growth of over-story trees and gradual canopy 1.7 kg a.i. ha 1 and 2.2 kg a.i. ha 1, respectively, is closure in the late-successional period (Elliott et al. registered for preemergence control in ornamental 1997). In managed pine plantations in Ohio, the plantings (BASF 2011). A mixture of dicamba, thien- importance value (sum of relative cover and relative carbazone-methyl and iodosulfuron-methyl-sodium at a 740 CANADIAN JOURNAL OF PLANT SCIENCE
recommended rate of 78 g a.i. ha 1, 4.9 g a.i. ha 1, and sap feeders and leaf strippers from 51 mg m 2 in 22.5 g a.i. ha 1, respectively, is registered for postemer- untreated areas to up to 252 mg m 2 in treated areas. gence control in warm-season grass turf (BAYER Sharp declines in the herbivore biomass corresponded to Undated). A mixture of soil-applied trifluralin and dieback of the E. hieraciifolius (and other herbaceous isoxaben at a recommended rate of 3.4 kg a.i. ha 1 plants) and increases in detritivore populations exploit- and 0.9 kg a.i. ha 1, respectively, is registered for ing nutrients available in the litter. preemergence control in turf, nursery and ornamental Larvae of Palthis asopialis (Guene´ e) are reported to plantings (Dow AgroSciences 2010). feed on E. hieraciifolius, and a variety of other plants For container-grown nursery crops, Neal and Derr (Robinson et al. 2010). This noctuid moth is known to (2005) provide a table on the efficacy of a number of occur only in extreme southern Ontario (Troubridge preemergence herbicides registered for use in ornamen- and Lafontaine 2004) and Quebec (J. D. Lafontaine, tal plantings in the US. Of the 11 herbicides tested, they personal communication, Agriculture and Agri-Food rate 4 as providing ‘‘good’’ control of E. hieraciifolius: Canada, Ottawa, ON). Larvae of Tyria jacobaeae L. flumioxazin (420 g a.i. ha 1); oxyfluorfen oxadiazon (Arctiidae), a biological control agent established in (2.2 1.1 kg a.i. ha 1); oxyfluorfen oryzalin (2.2 1.1 Canada against Senecio jacobaea L. (Harris et al. 1975), kg a.i. ha 1); and, oxyfluorfen pendimethalin (2.2 were reported to feed and mature on E. hieraciifolius in 1.1 kg a.i. ha 1). forced feeding trials (Bucher and Harris 1961). Outside North America, several species of moths have been 12. Response to Other Human Manipulation reported to include E. hieraciifolius in their host range Prior to the advent of chemical herbicides, Dalaire (Robinson et al. 2010), including Hypercompe icasia (1904) stated that control could be accomplished by Cramer (Arctiidae), Platyptilia molopias Meyrick (Pter- cultivation, hoeing and hand pulling. Even now in ophoridae), and Platphalonidia subolivacea Walsingham cranberry and blueberry fields of the Maritime pro- (Tortricidae). vinces control is often accomplished by hand pulling Evidence of leaf-mining insects is not uncommon on (G. Graham, personal communication, New Brunswick herbarium specimens of E. hieraciifolius from Canada. Department of Agriculture, Aquaculture and Fisheries, The microlepidoptera Phyllocnistis insignis Frey & Boll Fredericton, NB). Manual pulling and mowing, prior (Gracillariidae) has been reported on E. hieraciifolius in to seed production, has also been recommended for the US (Robinson et al. 2010). Although larvae control in the United States (Runnels and Schaffner produce track mines in a variety of herbaceous 1931; Muenscher 1955). Diligent sanitation of harvest- Asteraceae and the species has been reported from ing equipment appears to reduce spread in wild blue- several border States, it is not yet known to occur in berry fields in Maine (D. Yarborough, personal Canada. A lepidopteran miner on Erechtites was communication, University of Maine, Orono, ME). reported from Kentucky under the name Phyllocnistis
For personal use only. erechtitisella Chambers, but no description was pro- vided and the name cannot be used (Chambers 1878). 13. Response to Herbivory, Disease and Higher Reports of the Gracillariid Phyllonorycter insignis Plant Parasites (Walsingham) attacking Erechtites species are in error (a) Herbivory (J.-F. Landry, personal communication, Agriculture (i) Mammals, including both domestic and wild animals * and Agri-Food Canada, Ottawa, ON). No reports have been found indicating whether the In south Florida, E. hieraciifolius is among the many bitter leaves with their unpleasant odour are palatable species of Asteraceae suitable as hosts for the leaf miner to mammals. fly Phytobia maculosa (Malloch) [ Nemorimyza macu- losa (Malloch)] (Agromyzidae), whose larvae form large blotch mines on the leaves (Stegmaier 1967). In (ii) Birds and other vertebrates * No information was the vicinity of vegetable crops in Florida, the highly found. polyphagous serpentine leaf miners Liriomyza sativae Blanchard and L. trifolii (Burgess) (Agromyzidae) are
Can. J. Plant Sci. Downloaded from pubs.aic.ca by Agriculture and Agri-food Canada on 06/04/12 found on E. hieraciifolius (Genung et al. 1978; Genung (iii) Insects * The Chinese ladybeetle, Leis conformis 1981; Schuster et al. 1991). In Taiwan, E. hieraciifolius Boisd. (Coccinellidae: Coccinellini), imported into was found to be a preferred host of the leafminer L. Florida to combat the green citrus aphid, Aphis spir- trifolii (Chien and Ku 1998). Although none of these aecola Patch (Homoptera: Aphididae), has been found leaf-mining flies occur in the field in Canada, the latter is feeding on the blossoms of E. hieraciifolius; although a pest of greenhouse crops (Broadbent and Olthof 1995; entire blossoms including stamens and pistils were eaten, Dempewolf 2004; Lonsdale 2011). no larvae, pupae or eggs were observed (Watson and In Arkansas, E. hieraciifolius is host to the tarnished Thompson 1933). In a young pine forest in South plant bug, Lygus lineolaris P. Beauv. (Heteroptera: Carolina, increased biomass of E. hieraciifolius led to Miridae) (Young 1986). A gall midge, Neolasioptera sp. substantial increases in arthropod trophic guilds, such as (Diptera: Cecidomyiidae), has been reported on E. DARBYSHIRE ET AL. * ERECHTITES HIERACIIFOLIUS (L.) RAF. EX DC. 741
hieraciifolius in Florida, causing irregular stem swel- DC. [ E. betae (Vanha) Weltzien] (NY), Sphaerotheca lings 2 5 cm long; and, a flower-head gall produced by castagnei Le´ v. (VT), S. humuli (DC.) Burrill [ Podo- Asphondylia sp. (Cecidomyiidae) has been reported on sphaera macularis (Wallr.) U. Braun & S. Takam.] Erechtites sp. from an unspecified North American (eastern states, IA, IL, MN and MT), and S. macularis location (Gagne´ 1989). Infestation by the cosmopolitan (Wallr.) P. Magnus [ Podosphaera macularis (Wallr.) aphid species Brachycaudus helichrysi (Kaltenbach) U. Braun & S. Takam.] (MT, OH); root-rot fungus (Homoptera: Aphididae) was found on E. hieraciifolius Phymatotrichum omnivorum Duggar [ Phymatotri- in two Florida counties in 1997 (Halbert et al. 2000). chopsis omnivora (Duggar) Hennebert] (TX); the stem- Blatchley (1912) reported that a carabid beetle rot fungi Sclerotinia sclerotiorum (Lib.) de Bary (FL) [Anisodactylus terminatus (Say)] was often seen feeding and Sclerotium rolfsii Sacc. [ Athelia rolfsii (Curzi) C. on ripening seeds in Indiana. In Kagoshima City, C. Tu & Kimbr.] (FL); and, the leaf gall Synchytrium Japan, E. hieraciifolius was one of three plants infested erechtitis M. T. Cook (LA). In other parts of the world by an Asian moth, Nyctemera adversata (Schaller) Ramularia wisconsina was recently reported on E. (Lepidoptera: Arctiidae), with 72 92% of the larvae hieraciifolius in Korea (Shin and Braun 2000), Cercos- maturing in 27.5 d (Murakami et al. 1999). In host pora erechtitis in Venezuela, Septoria erechtitis from testing of the Lepidopteran Secusio extensa (Butler) as Cuba and the West Indies, and the leaf spot pathogen a biological control agent in Hawaii, E. hieraciifolius Mycosphaerella erechtitidina Petr. & Cif. was reported was found to be a poor host, with less than 2% of from Cuba, the Dominican Republic and the West larvae completing development on the plant (Ramadan Indies (Farr and Rossman 2010). et al. 2010). In a study of mildews and rusts affecting the Asteraceae (Kenneth and Palti 1984), Erechtites spp. were hosts to species of the downy mildews Bremia (iv) Nematodes and other invertebrates * Several species and Plasmopara, to the rust genus Puccinia,andto of root-knot nematodes have been reported on E. the powdery mildews Erysiphe and Sphaerotheca.In hieraciifolius, including Meloidogyne enterolobii Yang & Hawaii, E. hieraciifolius was among a majority of Eisenback ( M. mayaguensis Rammah & Hirschamm), plants tested which did not inhibit growth of the soil M. javanica (Treub) and M. incognita (Kofoid & White) fungus Phytophthora palmivora Butler (Powell and Ko in association with or proximity to agricultural crops 1986). (Carneiro et al. 2006; Rich et al. 2010). However, these are tropical and subtropical species and only the latter is present in Canada as a pest in greenhouse production (ii) Bacteria * In laboratory testing Pseudomonas systems (Ebsary and Eveleigh 1983). cichorii (Swingle) Stapp and P. viridiflava (Burkholder) Dowson were found to be pathogenic on E. hieraciifolius
For personal use only. (Tsuchiya et al. 1982). Both these bacterium species have (b) Diseases broad host ranges which include many crop species. (i) Fungi * There are few reports of fungi on E. hieraciifolius in Canada (Ginns 1986). The powdery mildew, Sphaerotheca fuliginea (Schltdl.) Pollacci [ (iii) Viruses * In Florida, E. hieraciifolius was identified Podosphaera fuliginea (Schltdl.) U. Braun & S. Takam.], as a host for the Bidens mottle potyvirus (BMV), which has been reported on E. hieraciifolius in Ontario was also infecting nearby lettuce (Lactuca sativa L.) and and Quebec (Parmelee 1977; DAOM specimens), endive (Cichorium endivia L.) crops (Purcifull and Zitter and Cicinobolus cesatii de Bary ( Ampelomyces quis- 1973). qualis Ces. ex Schlecht.), a parasitic fungus of powdery mildews, was reported by J. Dearness in 1895 on E. hieraciifolius collected in London, ON (Farr and (c) Higher Plant Parasites * No information has been Rossman 2010). found. A number of other phytopathogenic fungi have been
Can. J. Plant Sci. Downloaded from pubs.aic.ca by Agriculture and Agri-food Canada on 06/04/12 reported on E. hieraciifolius in the United States (Farr ACKNOWLEDGEMENTS and Rossman 2010) including downy mildews, Bremia J. Calder, G. Graham, J. D. Lafontaine, J.-F. Landry, lactucae Regel (PA), and Peronospora halstedii Farl. F. and K. Starr, and D. Yarborough are thanked for [ Plasmopara halstedii (Farl.) Berl. & de Toni] (MA, kindly sharing information and expertise. Peter Craig, of MD, NJ, TX and WI); leaf spot fungi Cercospora New Jersey, kindly provided the photograph in Fig. 5. erechtitis G. F. Atk. (AL, FL, TX), Phyllosticta Useful comments on the manuscript were provided by J. erechtitis F. L. Stevens & P. A. Young (HI), Ramularia Cayouette (Agriculture and Agri-Food Canada) and wisconsina H. C. Greene (WI), and Septoria erechtitis Mihai Costea (Wilfred Laurier University). We thank Ellis & Everh. (AK, DE, TX and WI); powdery the curators and staff of the following herbaria who mildews Erysiphe cichoracearum DC. [ Golovinomyces made material in their care available for study: ACAD, cichoracearum (DC.) V.P. Heluta] (MD), E. polygoni CAN, DAO, HAM, LKHD, MT, MTMG, NSAC, 742 CANADIAN JOURNAL OF PLANT SCIENCE
OAC, QFA, QK, QUE, TRT, TRTE, UNB, WAT, and jacobaeae (L.) (Lepidoptera: Arctiidae). Can. Entomol. 93: WLU [see Holmgren et al. (1990) for institutional 931 936. abbreviations]. Burrows, G. E. and Tyrl, R. J. 2001. Toxic plants of North America. Iowa State University Press, Ames, IA. viii 1342 pp. Abella, S. R. 2010. Thinning pine plantations to reestablish Busey, P. and Johnston, D. L. 2006. Impact of cultural factors oak openings species in northwestern Ohio. Environ. Manag. on weed populations in St. Augustinegrass turf. Weed Sci. 54: 46: 391 403. 961 967. Adams, R. and Gianturco, M. 1956. Senecio alkaloids. The Candolle, A. P. de 1837. Prodromus systematis naturalis. Part composition of hieracifoline and jacobine. J. Am. Chem. Soc. 6. Treuttel & Wurtz, Paris, France. 687 pp. 78: 398 400. ¨ Carneiro, R. G., Do Amaral Monaco, A. P., Moritz, M. P., Anonymous. Undated. Crop profile for cranberries in New Jersey. New Jersey Agricultural Experiment Station, Nakamura, K. C. and Scherer, A. 2006. Identification of Rutgers, The State University of New Jersey. [Online] Avail- Meloidogyne mayaguensis in guava and weeds, in loam soil, able: http://pestmanagement.rutgers.edu/njinpas/CropProfiles/ in Parand State. Nematologia Brasileira 30: 293 298 [in cranberryprofile.pdf [2012 Jan. 04]. Portuguese, English summary]. AQIS. 2011. Australian Quarantine Information Service Im- Chambers, V. T. 1878. Descriptions of new Tineina from port Conditions (ICON) Database. [Online] Available: http:// Texas, and others from more northern localities. Bull. U. S. www.daff.gov.au/aqis/import/icon-icd [2012 Jan. 04]. Geol. Geogr. Surv. Territories 4:79 106. Barkley, T. M. 2006. Pages 595 602 604 in Flora of North Chien, C.-C. and Ku, H.-C. 1998. The occurrence of Liriomyza America Editorial Committee, eds. Flora of North America trifolii (Diptera: Agromyzidae) and its parasitoids on fields of north of Mexico. Vol. 20. Magnoliophyta: Asteridae, part 7: Gerbera jamesonii. Chin. J. Entomol. 18: 187 196 [in Chinese, Asteraceae, part 2. Oxford University Press, New York, English abstract]. Oxford. Clark, G. H. and Fletcher, J. 1909. Farm weeds of Canada. 2nd Barkley, T. M. and Cronquist, A. 1978. Erechtites. N. Am. ed. Department of Agriculture, Ottawa, ON. 192 pp. Flora, Ser. II 10: 139 142. Coleman, J. R. 1982. Chromosome numbers of angiosperms Barton, W. P. C. 1818. Compendium florae Philadelphicae. collected in the state of Sa˜o Paulo. Rev. Brasil. Genet. 5: Vol. II. M. Carey and Son, Philadelphia, PA. 234 pp. 533 549. BASF. 2011. Freehand† 1.75 Herbicide. [Online] Avai- Cooper, G. O. 1936. Cytological investigations of Erechtites lable: http://betterturf.basf.us/products/freehand-1.75g-herbicide.html hieracifolia. Bot. Gaz. 98: 348 355. [2012 Jan. 04]. Coxe, R. B., Stephenson, S. L., Madarish, D. M. and Miller, Baskin, C. C. and Baskin, J. M. 1996. Role of temperature and G. W. 2006. Vascular flora of Fernow Experimental Forest light in the germination ecology of buried seeds of weedy and adjacent portions of Otter Creek Wilderness area. USDA species of disturbed forests. II. Erechtites hieracifolia. Can. J. Forest Service, Northeastern Research Station Gen. Techn. Bot. 74: 2002 2005. Rep. NE-344. 27 pp. BAYER. Undated. Celsius(TM) WG Herbicide specimen label, Cronquist, A. 1946. Notes on the Compositae of the north- US79925816B 100324A. [Online] Available: http://www.cdms. eastern United States. III. Inuleae and Senecioneae. Rhodora
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