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The Biology of Canadian weeds. 150 Erechtites
hieraciifolius (L.) Raf. ex DC.
Stephen J. Darbyshire1, Ardath Francis1, Antonio DiTommaso2, and David R. Clements3
Can. J. Plant Sci. Downloaded from pubs.aic.ca by Agriculture and Agri-food Canada on 06/04/12
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1
Agriculture 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: 729746. 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
(Fragariaananassa) 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, érechtite à feuilles d’épervière, 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: 729746. L’érechtite à feuilles d’épervière (Erechtites
hieraciifolius) est une herbacée annuelle de la famille des Astéracées indigène aux régions forestières de l’est de l’Amérique
du Nord, mais qui a été introduite dans certaines parties de l’Europe et du Pacifique. On la confond parfois à d’autres
adventices à rayons courts de la famille des Composées qui poussent au Canada tels le séneçon vulgaire (Senecio vulgaris)
et la vergette du Canada (Conyza canadensis). Espèce pionnière thérophyte, on en retrouve de grand peuplements aux
endroits qui ont subi de profondes perturbations comme un feu de forêt, cependant l’espèce colonise souvent des zones plus
restreintes comme les rivages, la lisière des boisés et les chablis. Les conditions du sol peuvent varier considérablement
quant à la concentration d’éléments nutritifs, la teneur en eau, le pH et la salinité. Au Canada, on la considère
généralement comme une adventice secondaire dans les jardins et les champs. L’érechtite à feuilles d’épervière se rencontre
le plus souvent dans les cultures comme le bleuet nain (Vaccinium spp.), la canneberge (Vaccinium macrocarpon), la fraise
(Fragariaananassa) et les plantes maraı̂chères, cependant, on la découvre à l’occasion dans d’autres grandes cultures et
des cultures fourragères. Elle engendre divers problèmes qui concourent à accroı̂tre les coûts de production ou à réduire le
rendement, notamment la résistance aux herbicides, la concurrence pour les ressources existantes et le fait de servir de
réservoir aux agents pathogènes des cultures. L’espèce est prisée pour ses vertus médicinales et on s’en sert pour soigner
divers maux. Les peuplements prennent parfois rapidement de l’ampleur à la suite des perturbations typiques au
aménagements paysagers, mais on peut habituellement venir à bout de E. hieraciifolius par les moyens de lutte chimique,
culturaux ou manuel.
Mots clés: Erechtites hieraciifolius, érechtite à feuilles d’épervière, American burnweed, pilewort, écologie des adventices,
biologie des mauvaises herbes
1. Name and Generic Status
Erechtites hieraciifolius (L.) Raf. ex DC. * Synonyms
(spelling not corrected): Senecio hieracifolius L.; E.
hieracifolia var. intermedia Fern.; E. praealta Raf.; E.
hieracifolia var. praealta (Raf.) Fern.; Eriophthalmia
Can. J. Plant Sci. (2012) 92: 729746 doi:10.4141/CJPS2012-003
hieracifolia (L.) Prov.; Neoceis hieracifolia (L.) Cass.
Common names: American burnweed (Darbyshire et al.
2000), eastern burnweed, fireweed (fire-weed), pilewort
(Darbyshire 2003), butterweed (Belcher 1956), white
fireweed (Fogg 1945); érechtite à feuilles d’épervière
Abbreviation: DALPT, days after late postemergence treatment
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730
CANADIAN JOURNAL OF PLANT SCIENCE
(Darbyshire et al. 2000), crève-à-yeux, crève-z-yeux
(Darbyshire 2003). European and Mediterranean
Plant Protection Organization (Bayer) code: EREHI.
Asteraceae (Compositae): aster family Asteracées
(Composées).
The name has often been spelled as ‘‘E. hieracifolia’’or ‘‘E. hieraciifolia’’. According to the International
Code of Botanical Nomenclature (McNeill et al. 2006),
Erechtites hieraciifolius is correct, the genus being
masculine in gender and an ‘‘i’’ being required between
the two elements (‘‘hieraci’’ and ‘‘folius’’) of the epithet
(i.e., a double ‘‘i’’). Augustin de Candolle (DC.) was
the first to publish the name E. hieraciifolius in 1838,
based on the earlier name and description, Senecio
hieraciifolius, by C. Linnaeus (1753). In doing so de
Candolle acknowledged that C. S. Rafinesque (Raf.)
had suggested this to him in a letter; the nomenclatural
authority may be given as ‘‘Raf. ex DC.’’ (as above) or
simply ‘‘DC.’’ (McNeill et al. 2006). For a more
complete list of synonyms see Belcher (1956).
This account deals with E. hieraciifolius var. hieraciifolius, the only variety present in Canada, and not with
the nearly pantropical weed E. hieraciifolius var. cacalioides (Fisch. ex Spreng.) Griseb. (see Section 2c), unless
explicitly mentioned.
2. Description and Account of Variation
(a) Description * The following description is of
E. hieraciifolius var. hieraciifolius, the variety present
in Canada and most of the United States. It shows
considerable phenotypic variability (Belcher 1956),
especially in plant size and leaf shape (Fig. 1). The
description here is based on material from Canadian
populations, supplemented with information from published descriptions (e.g., Fernald 1917; Kummer 1951;
Belcher 1956; Gleason and Cronquist 1991; Barkley and
Cronquist 1978; Barkley 2006).
Annual herb, (5) 50200 ( 300) cm tall; taproot
with fibrous secondary roots; stem erect, more or less
strongly ribbed (grooved), simple or branched toward
the top, glabrous to villose with translucent multicellular
hairs; leaves alternate, the lower (3) 620 cm long (1) 28 cm wide and diminishing in size upwardly,
mostly glabrous or margins ciliate with short hairs and
lower surface usually also with longer multicellular hairs
(especially along the principal veins), irregularly toothed
and lobed, with secondary veins extending to the
marginal gland-tipped teeth (Fig. 2); lower leaves elliptic
to broadly lanceolate, shallowly toothed to weakly acute
lobed, usually weakly petiolate; middle to upper leaves
lanceolate, acute, irregularly toothed, often deeply acute
lobed, sessile or clasping at the base, with distal leaves
bract-like.
The leaves are highly variable in size and shape. The
base may be attenuate to a distinct petiole (Fig. 1B;
Lloyd and Lloyd 1887), broadly sessile, or expanded and
clasping to auriculate (Fig. 1A). Several leaf forms are
often visible on the same plant and those with clasping
or auriculate bases tend to be larger toward the middle
of the stem.
Inflorescence varying from a single terminal capitulum (flower head) in depauperate specimens (Fig. 1C)
to compound, corymbose panicles usually with many
capitula (often 100 or more) on robust plants; capitula
subcylindric or flask-shaped (slightly swollen at the
base), (6.5) 1017 mm long, with a few linear
bracteoles at the base and upper pedicel (sometimes
collectively referred to as a calyculus); phyllaries
(involucral bracts) about 20 in a single row, linear
with a slender attenuated tip, (6) 1017 mm long and
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.
Fig. 2. Lower (abaxial) surface of leaf of Erechtites hieraciifolius showing secondary veins extending to the blade margins
(somewhat obscure in the black and white version of this
photo) and terminating in gland-tipped teeth. Translucent,
multicellular hairs present along the veins (and margins). Inset
shows a single hair.
0.51.5 mm wide, becoming strongly reflexed with age;
ligulate (ray) florets lacking but florets heterogamous,
the outer florets pistillate, 10100, corollas whitish to
yellow, tubular filiform, 45 deltate lobes, erect; disc
florets 1020 ( 50), mostly bisexual and fertile,
inner sometimes functionally staminate, corollas whitish to pale yellow or pinkish, 45 erect to spreading
deltate lobes; receptacle naked, 58.5 mm diam.;
cypselae (fruits) narrowly cylindric to oblong, 23 mm
long, brown with 812 paler ribs, strigose (rarely
glabrous) between the ribs, the apex with a conspicuous
collar and persistent style base protruding from the
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centre (Fig. 3C); pappus of copious white hairs, about
510 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
lobed or emarginate, 69 (10) (3.5) 59 mm,
usually red-tinged beneath, a few scattered bristly short
hairs above and on the usually purple principal veins
beneath, and a granular bloom on the lower surface.
Chromosome counts on plants of E. hieraciifolius at
Ottawa, ON, were determined as n 20 by G. A.
Mulligan in 1966 (collection number 3118; voucher
DAO 857169). This is consistent with published counts
of n 20, 2n 40 (Cooper 1936; Ornduff et al. 1963;
Coleman 1982).
(b) Distinguishing Features * In eastern Canada, Erechtites hieraciifolius is distinguished from most other
composites by its many flowered (20150), relatively
large (1017 mm) capitula with the outer pistillate florets
lacking ligules (rays), and the small (23 mm long)
seeds with copious white pappus hairs. Superficially,
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.
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it somewhat resembles the annual Canada fleabane,
Conyza canadensis (L.) Cronq., whose usually simple
stalks reach comparable heights, and are topped by
branched inflorescences of numerous small capitula
(about 34 mm long) with phyllaries of unequal lengths
and outer pistillate florets obscurely ligulate, and tiny
seeds (11.5 mm long). The stems of Canada fleabane
are covered with simple hairs bent upward (adpressed),
the leaves are crowded and numerous, linear and
attenuated in shape; the leaf margins, when not entire,
are often more coarsely toothed with small teeth lacking
glandular tips.
The widespread species Senecio vulgaris L. (common
groundsel) is another annual species which usually lacks
ligulate florets. It may be distinguished from E. hieraciifolius by: its deeply lobed mid-stem leaves (appearing
coarsely pinnatifid), with the lobes obtuse rather than
acute; the black tips of the calycular bracts (and
sometimes also the phyllaries); smaller capitula about
58 mm long; and, the smaller seeds, 1.52 ( 2.5) mm
long. The annual rayless aster [Brachyactis ciliata
Ledeb.; Symphyotrichum ciliatum (Ledeb.) G. L.
Nesom] also has small capitula (511 mm long) and
seeds (1.52.5 mm long), but there are multiple rows of
phyllaries and the leaves are linear to oblanceolate with
entire and usually ciliate margins.
Several native species of Packera (Senecio sensu
lato) in eastern Canada may also be confused with E.
hieraciifolius, such as the rayless P. indecora (Greene)
Á. Löve & D. Löve, and the occasionally rayless P.
paupercula (Michx.) Á. Löve & D. Löve. These perennial species have smaller seeds (52 mm), lower leaves
that are distinctly spatulate with a long petiole (usually
as long as or longer than the blade), and the mid-stem
leaves somewhat lyrate.
(c) Intra-specific Variation * This species is highly
polymorphic throughout its range (Fernald 1917, 1950;
Belcher 1956). Although several entities have been
variously recognized taxonomically, as varieties or
distinct species, the variation is continuous within and
between the forms (Deam 1940; Belcher 1956; Seymour
1982). Most of the striking variation in plant size and
leaf size and shape is attributable to phenotypic
responses to varying conditions (Belcher 1956). Two
North American varieties have been commonly recognized: var. hieraciifolius and var. megalocarpus (Fern.)
Cronq. The latter is a form found only in coastal salt
marshes of New England (Barkley 2006), which differs
from the typical form in its more succulent leaves and
stems, a receptacle 8 mm in diameter, and larger seeds
(45 mm) with 1620 nerves (Barkley 2006). Although
Fernald (1950) considered it a distinct species, other
authors generally treat it as ‘‘merely a well-marked
ecotype of saline coastal marshes’’ with the morphological variation between the two forms being continuous
(Cronquist 1946; Belcher 1956; Barkley 2006).
A third variety is present in the tropics, var. cacalioides, and found in the West Indies and from Mexico
through Central America to northern Argentina and in
southeast Asia and the East Indies (Belcher 1956). This
variety differs in its longer bracteoles which extend from
one-third to one-half the length of the involucre, and by
the presence of multicellular hairs on the bracteoles and
phyllaries (Belcher 1956).
In addition to treating megalocarpus and cacalioides
as distinct species, Fernald (1917, 1950) recognized three
intraspecific taxa within E. hieraciifolius in eastern
North America [var. hieraciifolius ( var. typica), var.
praealtus (Raf.) Fernald, and var. intermedius Fernald],
based primarily on leaf size and shape. Few have
adopted this intraspecific treatment pointing out that
single populations may contain forms typical of all three
varieties (e.g., Deam 1940) and that leaf form and size
are affected by the growing conditions (Fogg 1945;
Belcher 1956).
(d) Illustrations * Various aspects of E. hieraciifolius
are illustrated in Figs. 1, 2, and 3. Variation in leaf shape
is shown in the different elements of Fig. 1. Cypselae,
seeds and seedlings are illustrated in Fig. 3. Other useful
illustrations may be found in Pammel et al. (1913), Fogg
(1945), Cronquist (1955), Muenscher (1955), Barkley
(2006), Neal and Derr (2005), Jung and Chung (2010),
Trewatha (2010), and USDA-NRCS (2011).
3. Economic importance
(a) Detrimental * Various authors include it among
the agricultural or forestry weeds in Canada (e.g.,
Provancher 1862; Dalaire 1904; Clark and Fletcher
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
(Vaccinium macrocarpon Aiton) and blueberry (Vaccinium spp.) fields in Nova Scotia, Prince Edward Island
and New Brunswick where it is usually easily controlled
with broad-spectrum herbicides or sometimes hand
pulling (J. Calder, personal communication, Nova
Scotia Department of Agriculture, Truro, NS; G.
Graham, personal communication, New Brunswick
Department of Agriculture, Aquaculture and Fisheries,
Fredericton, NB). Wild blueberry crops may be particularly susceptible to infestations since periodic burning
is often used as a tool for pruning and weed management (Penney et al. 2008). Data on labels of herbarium
specimens collected in Canada indicate that it sometimes
occurs in a wide variety of other crops, including
oat (Avena sativa L.), barley (Hordeum vulgare L.),
maize (Zea mays L.), strawberry [Fragariaananassa
(Weston) Duch. ex Rozier], onion (Allium cepa L.) and
carrot (Daucus carota L.), as well as fodder crops
(Medicago sativa L.) and mixed pastures.
It is sometimes considered a weed in the eastern
United States (e.g., Pammel et al. 1913; Runnels and
Schaffner 1931; Muenscher 1955). As in Canada, E.
hieraciifolius is present in wild blueberry fields in
Maine (D. Yarborough, personal communication, University of Maine, Orono, ME) and cranberry fields in
New Jersey (Anonymous, undated), where, in high
densities, plants can obstruct harvesting equipment
(D. Yarborough, personal communication, University
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
Europe (Belcher 1956; Voss 1996). It has also become a
serious agricultural weed in Hawaii, invading sugar
cane (Saccharum officinarum L.) plantations in that
state (Hanson 1962; Sund 1964). Haselwood and
Motter (1966) stated that, in Hawaii, it was, ‘‘One of
the most abundant weeds in moist cultivated areas;
also a weed in pastures and rangelands.’’ Although
E. hieraciifolius is naturalised on all of the major
Hawaiian islands (HEAR 2008), it is currently not
as abundant as E. valerianifolius (Link ex Spreng.)
DC. (F. and K. Starr, personal communication, US
Geological Survey, HI).
It has been reported a weed in pine plantations in the
southern United States (Britt et al. 1990; Gardiner et al.
1991); as an increasingly important reservoir host for
crop pathogens including insects, fungi, nematodes and
viruses (see Section 13a, b); as a weed of cultivated lands
(Runnels and Schaffner 1931; Trewatha 2010); as a weed
of pastures and forage grasses (Pammel et al. 1913;
Muenscher 1955; Stephenson and Rechcigl 1991), and
turf grass (Busey and Johnston 2006; Trewatha 2010).
It is also reported as a problem weed in the container
nursery industry (Neal and Derr 2005).
It is considered a potential weed in Australia where it
is under quarantine (AQIS 2011). In Hungary it is
considered an invasive pest (Csiszár 2006).
The presence of alkaloids (see also Section 7c) causes
hepatotoxicity if the plants are consumed in sufficient
quantity (Burrows and Tyrl 2001). The plant is frequently reported to have a noxious smell and taste, so it
is unlikely that persons or livestock would voluntarily
consume sufficient quantities to be at serious risk.
However, the overall similarity between E. hieraciifolius
and some even more toxic Senecio species may cause
difficulties through pest mis-identification and inappropriate management decisions.
As implied by the French common name ‘‘crève-zyeux’’, the pappus hairs were thought to pose a serious
threat to the eyes (Provancher 1862). The flocculent
pappus bristles may well cause eye irritation when
manually threshing grain crops heavily contaminated
with E. hieraciifolius, especially in relatively stagnant air
spaces. The pappus bristles have scattered antrorse
spicules which may contribute to the abrasion of soft
tissues such as eyes and lungs.
From a sociocultural perspective, Harper (1944)
indicates that it is greatly offensive to aesthetic sensibilities, saying that, in Alabama, ‘‘Although harmless
733
enough, and never interfering with crops, this is one of
our most useless and disreputable-looking weeds, very
unattractive in appearance, and ill-scented besides.’’
Vegetation management resources are not always expended on an economic basis and such perceptions of
‘‘weeds’’ can have a tremendous impact on our relationships with, and actions on, plants and landscapes.
(b) Beneficial * As a medicinal plant in North America,
the use of E. hieraciifolius has had a complex and
confusing history which Lloyd and Lloyd (1887) review
extensively. These authors indicate that ‘‘fireweed oil’’
of North American commerce was usually derived from
other species (not E. hieraciifolius), but maintain that E.
hieraciifolius oil has distinct benefits. Erichsen-Brown
(1979) cited a number of early North American sources
on medicinal uses of the plant in treatment for
haemorrhage, wounds, skin diseases, dysentery, cholera,
and as a purgative and emetic; as a source of a blue dye
for cotton and wool; and, its use by Algonquin native
peoples to treat poison ivy [Toxicodendron radicans (L.)
Kuntz] and poison sumac [T. vernix (L.) Kuntz]
poisoning. Hale (1880) recommends its use in the
treatment of dysentery, menstrual disorders and gonorrhoea. Millspaugh (1887) considered the plant’s volatile
oil to be the principal active ingredient in a tincture
produced from whole fresh flowering plants for use as
an emetic and in treatment of such conditions as eczema,
diarrhoea, haemorrhages and piles, but noted that
nausea and other adverse reactions could result from
use of this tincture. In the early 20th century its use as a
medicinal herb was sufficiently popular that collectors
were paid 23 cents per pound (Georgia 1914).
Many authors comment on the unpleasant or rank
odour of the plant. In spite of this Fernald et al. (1958)
suggest it could be used as a salad or potherb. The
plant’s odour does not seem to be consistently distasteful to all people.
In the Andes of South America the leaves and flowers
(var. cacalioides) have been used in folk medicine as a
blood depurative and the roots to treat cardiac disease
(Lorenzo et al. 2001). The antioxidant and radical
scavenging properties of nine species of Asteraceae
from Bolivia were investigated by Parejo et al. (2003);
although some activity was exhibited by E. hieraciifolius,
other species appeared more promising as sources of
compounds for industrial and medical use.
In Japanese trials, E. hieraciifolius has been found to
be among the most efficient of the plants tested at
assimilating atmospheric nitrogen dioxide (NO2) with as
much as 10% of its total organic nitrogen content
derived from this source (Morikawa et al. 1998). Such
plants have the potential to act as important sinks for
anthropogenic nitrogen oxides (Morikawa et al 1992;
Morikawa 1996). Morikawa et al. (2003) proposed that
‘‘green walls’’ using such nitrogen dioxide-philic plants
could be set up around buildings and on highway
734
corridors to help sequester pollutants from car emissions
or other sources.
(c) Legislation * Erechtites hieraciifolius is not listed in
any federal or provincial noxious weed legislation in
Canada; and is similarly absent from federal and state
lists in the United States. The state of New York has
listed E. hieraciifolius var. megalocarpus as endangered
in that state (USDA-NRCS 2011). Australia restricts
importation of E. hieraciifolius where it is considered a
potential threat and is regulated as a quarantine pest
(AQIS 2011).
4. Geographic Distribution
In North America, Erechtites hieraciifolius var. hieraciifolius is found primarily in deciduous forest regions of
eastern Canada from the Maritime Provinces to western
Ontario, and in the United States from New England
west to Minnesota in the north, and south to Florida
and eastern Texas. The distribution in Canada is shown
in Fig. 4, based on 837 specimens in Canadian herbaria.
References to its presence in Saskatchewan or Alberta
are almost certainly erroneous (Boivin 1972; Harms
2006), and probably based on the report by Hooker
(1834). No specimen was seen to confirm literature
reports by various authors (e.g., Gleason and Cronquist
1991) of occurrence in Newfoundland (Meades et al.
2000; Barkley 2006). It has been reported as introduced
to Washington State, apparently based on a single
collection from Seattle (Cronquist 1955), and as sporadically adventive on the West Coast (Barkley and
Cronquist 1978; Barkley 2006). It is not known to occur
in British Columbia (Douglas et al. 1998; Barkley 2006).
Elsewhere in the world, E. hieraciifolius var. hieraciifolius is known from the West Indies (Belcher 1956) and
has been introduced to central Europe (Belcher 1956;
Tutin 1976), Hawaii (Belcher 1956), Japan (Walker
1976; Kamada and Nakagoshi 1990), China (Chien
and Ku 1998), Taiwan (Wu et al. 2004; Jung and Chung
2010), Korea (Shin and Braun 2000), and New Zealand
(de Lange 1995).
5. Habitat
(a) Climatic Requirements* Climatic conditions within
the native distribution of E. hieraciifolius var. hieraciifolius vary from humid continental to maritime. The
plant extends over a wide latitudinal range in eastern
North America, from southern boreal (9498N) to
subtropical regions with considerable variation in average temperature and day length hours. In Canada, it is
not known from areas with a growing season of less
than about 160 d, or average precipitation of less than
about 70 cm.
(b) Substratum * There is little information in the
literature on the edaphic requirements and tolerances of
E. hieraciifolius in Canada, but data on herbarium
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
gravel. It is found in moist to saturated soils of shores,
flood plains, fresh and salt marshes, bogs, dune slacks,
gravel bars and ditches, but also occurs in dry or welldrained substrates such as dry sands, talus slopes, and
granite balds. A wide range of substrate pH is tolerated
as indicated by the collections from peat bogs to
limestone cliffs and talus. Although nutrient poor soils
are readily colonized, Wilson and Shure (1993) found E.
hieraciifolius biomass production was greater at harvested forest sites to which fertilizer was added.
More detailed analysis of soil types is found in some
United States sources. The substratum in a pine forest in
South Carolina, where E. hieraciifolius was among early
successional species, consisted of a silty sand down to
1.2 m underlain by a medium to fine clayey sand to 7.6
m; and the upper soil layers averaged 87% sand, 6.4%
silt and 6.3% clay-sized particles (Hunt and Shure
1980). Following treatment of the soil with waste water
in the above area, soil organic matter decreased from 3.5
to 1.5% and cation exchange capacity from an average
of 55 meq 100 g1 in the top 5 cm to 28 meq 100 g1 at
10 cm and 10 meq 100 g1 below 15 cm; but there were
otherwise no differences in the soil profile of treated and
untreated areas except for an increase in soil moisture
from 8% in untreated areas to up to 15% in treated
areas (Hunt and Shure 1980). Forest top-soils in eastern
Tennessee containing substantial seed banks of E.
hieraciifolius were found to have 3.25.3% organic
matter, 411 mg g1 phosphorus, 2126% clay, 28
43% silt and 3151% sand, with a pH of 5.26.0
(Farmer et al. 1982). The predominant soil types in
eastern mixed forest areas where E. hieraciifolius
appears following fires or forest clearance are loams
and silt loams originating from acid shales, sandstones
and limestones, often acidic with an average soil pH of
about 4.5 (Coxe et al. 2006). In the wetland complex of
the Delaware River (NJ), E. hieraciifolius was found in
shrub forest habitats on alluvial and aeolian deposits
(Leck et al. 1988).
(c) Communities in Which the Species Occurs * In
Canada, E. hieraciifolius occurs in boreal (southern
part), Great Lakes/St. Laurence, Acadian and Carolinian forest regions (Fig. 4). The species usually occurs in
abundance and high density in places of recent major
disturbance, although small scale disturbances giving
rise to suitable microsites are colonized by small
numbers of plants. Disturbances may result from fires
(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.
2005; Abella 2010), herbicide application (Kochenderfer
and Wendel 1983; Luken et al. 1994), cultivation
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;
Leck et al. 1988; herbarium specimen label data),
erosion (Peterson et al. 1990), or other disturbances
(Wagner 1966; Hunt and Shure 1980; Roman et al.
1984). In hemlock [Tsuga canadensis (L.) Carrière]
forests in New England damaged by an imported
adelgid [Adelges tsugae (Annand)], it is among opportunistic species said to have ‘‘invaded’’ the stands (Orwig
and Foster 1998). It is a poor competitor (Boring
et al. 1981) and Peterson et al. (1990) found that it
was negatively correlated (r0.45, P 0.018) with
total cover (primarily other plant species).
Population explosions may occur under these and
other conditions where competition is reduced, but
generally subside rapidly with successional progression
to less open environments. Shores of lakes and rivers,
habitats characterized by natural disturbance, are
common locations for E. hieraciifolius as are sites
associated with beaver activities causing cyclical flooding and drainage. It is found in a wide range of
anthropogenic habitats where vegetation is regularly
disturbed or maintained at early successional stages,
such as roadsides, railway lines, ditches, energy corridors, quarries, tree plantations, cultivated land, turf,
flower beds, or ruderal sites. It grows in coniferous,
deciduous or mixed forests, although usually in places
receiving some direct sunlight such as openings, edges,
or along trails. A variety of wetland habitats are also
suitable, including shores, wet thickets, wet forests, wet
prairies, marshes, bogs and fens (Fernald 1950; Roman
et al. 1984; Gleason and Cronquist 1991; Voss 1996;
Murphy et al. 2009; herbarium specimen label data). A
1975 collection of E. hieraciifolius in southern Ontario
came from a Spaghnum moss (Spaghnum sp.) mat
around a lake which had previously been flooded and
was then invaded by ‘‘hundreds of these weeds’’ after
the water level subsided (herbarium specimen label
data, OAC).
Smaller scale disturbances also provide microsites
where reduced competition within a broader community allows small populations of E. hieraciifolius to
establish; e.g., tree falls and wind-throws (Peterson and
Pickett 1990; Peterson et al. 1990; Peterson and Pickett
1995).
6. History
As a native plant of deciduous and mixed forest areas,
E. hieraciifolius has presumably been present in those
habitats since the re-vegetation of de-glaciated areas of
southern Canada after the end of the last Ice Age. In the
early 19th century, when clearing of the great eastern
forests of North America had begun in earnest, E.
hieraciifolius had become known as ‘‘fire-weed’’ because
of its abundant occurrence in areas of forest clearance,
especially where fires had occurred (Pursh 1814; Eaton
1824; Torrey 1843). William Barton (1818) said that it
was ‘‘one of the commonest weeds [around Philadelphia], growing almost every where, even on roofs’’.
7. Growth and Development
(a) Morphology* All annual plants are considered
therophytes, and this life-form is highly developed in
E. hieraciifolius. The species can rapidly respond to
take advantage of favourable growth conditions, rapidly
producing large individuals in dense populations
(Fig. 5). The phenotypic plasticity of its growth also
promotes maximum reproductive effort under suboptimal conditions. Depending on the availability of
moisture, nutrients and light, plants may reach maturity
at heights from a few centimetres and bearing only a
single capitulum, to more than 2 m with a few hundred
capitula (Fogg 1945; Csiszár 2006; Fig. 1). The plant is
capable of considerable morphological plasticity (see
736
Fig. 5. A dense population of Erechtites hieraciifolius in New
Jersey, 21 August 2008, Mr. Peter Craig in photo. The 1.6-ha
field was treated with two applications of glyphosate (about a
1-month interval), and then no-till planted with a variety of
native grassland species. It is unknown whether the growth of
E. hieraciifolius was the result of a soil seed bank, seed rain
from an outside source, or contamination of the planted seed.
Section 2) allowing a wide range of growth responses to
environmental conditions.
(b) Perennation * A typical therophyte, E. hieraciifolius
is a ‘‘summer’’ annual over-wintering only as seeds. It
has been postulated that some plants may behave as
winter annuals in southerly regions where winters are
milder than in Canada. In Kentucky, Baskin and Baskin
(1996) found that 1% of freshly collected seeds germinated immediately in an unheated greenhouse. They also
observed some plants of such a large size that it seemed
unlikely their growth could have been attained from
spring germinating seeds.
(c) Physiology and Biochemistry * The alkaloid hieracifoline, reported from E. hieraciifolius by Manske
(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 constituted 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. hieraciifolius subjected to simulated rain were at rates of: 0.03
mg 0.1 m 2 h1 NH
4 (ammonium ion); 2.4 K; 1.68
Ca; 0.31 Mg; 0.26 NO
3 (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 (2.55.5). Phosphorus was apparently absorbed from the simulated rain water rather
than leached. The relatively high rate of nutrient
leaching from the leaves was attributed to the wettability of adaxial leaf surfaces (Haines et al. 1985b) (see
Section 11). Tissue necrosis resulted from exposure to
pH levels below 2.5 (Haines et al. 1980).
The complex nitrogen metabolism has been extensively studied by Japanese researchers. Of the 217 plant
species studied in laboratory experiments, Morikawa
et al. (1998) observed that E. hieraciifolius had one of the
highest NO2 assimilation rates at 5.7 mg g1 dry weight,
representing 10% of the total nitrogen content. Average
levels of nitrate (NO
3 ); nitrite (NO2 ) and ammonium
(NH
)
ions
detected
in
the
leaves
of seedlings of E.
4
hieraciifolius by capillary electrophoresis were 16.8
mmol, 8.2 nmol and 1.0 mmol per gram of fresh tissue,
respectively (Kawamura et al. 1996). Of the six plant
species studied by Kawamura et al. (1996), E. hieraciifolius contained the smallest quantity (in relation to wet
weight) of NH4, and the largest quantity (in relation to
chlorophyll content) of NO
2 : The ability of E. hieraciifolius to convert NO
3 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
capacity of various plant species to convert NO
3 to
N2O (i.e., N2O emission) appears to be inversely
proportional to their ability to assimilate NO2 (Hakata
et al. 2003).
(d) Phenology * Seeds mostly germinate in the spring
(see Section 8c) and somewhat later with increasing
latitude. Flowering may begin as early as late July,
but seed maturation occurs mostly through mid-August
to early October (herbarium specimen label data).
Robertson (1928) reported the flowering period in
Illinois as Aug. 20 to Sep. 25. The basal leaves often
begin to wither as flowering commences (Barkley 2006).
(e) Mycorrhiza * The roots of E. hieraciifolius plants
germinated from forest topsoils (from eastern Tennessee) were extensively colonized by endomycorrhizae
with hyphae and vesicles present in all samples (Farmer
et al. 1982).
8. Reproduction
(a) Floral Biology * The capitula of E. hieraciifolius
lack ligulate florets to attract pollinators and are
probably primarily autogamous. Thirteen species of
mostly hymenopteran insect visitors were reported by
Robertson (1928) in Illinois; however, they are unlikely
to be effective pollinators.
The cytology, histology and ontogeny of micro- and
megasporogenesis, and early embryo development in E.
hieraciifolius were studied by Cooper (1936). Wagner
(1966) studied the effects of radiation on embryo
abortion rates (100% at B1.5 Gy d 1) in a field study
in Brookhaven, NY.
(b) Seed Production and Dispersal * The pappus of
copious large hairs at the apex of the seed (Fig. 3A)
facilitates wind dispersal (Fogg 1945; Ohtsuka 1998).
Large plants are capable of producing thousands of
seeds (see Section 2). In Hungary, Csiszár (2006) found
that an average of 32 390 seeds were produced per plant,
although growing conditions were not indicated. In a
forest area of NY, Wagner (1965) measured the seed
rain of E. hieraciifolius from late summer to early
autumn at 36 145 seeds ha1.
In agricultural settings, E. hieraciifolius may also be
dispersed through mechanical harvesting equipment
(J. Calder, personal communication, Nova Scotia Department of Agriculture, Truro, NS; D. Yarborough,
personal communication, University of Maine, Orono,
ME).
(c) Seed Banks, Seed Viability and Germination * The
results of various observations and experiments on seed
bank formation of E. hieraciifolius are incongruous and
difficult to interpret. Germination tests of exhumed
seeds, artificially buried for long periods, have shown
737
either no viability (Telewski and Zeevaart 2002) or as
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
and Humphreys 1940; Livingston and Allesio 1968;
Leck et al. 1988; Matlack and Good 1990; Schiffman
and Johnson 1992), in spite of the potential for
immigration of wind-borne seeds. The low numbers of
seeds and distributional patchiness in mature forest soils
found by these studies suggests that long-term seed
banks are not formed. However, other studies have
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
in the results of studies measuring germination may be
due in part to experimental conditions and the cyclical
conditional dormancy exhibited by E. hieraciifolius
(Baskin and Baskin 1996). Direct observation of ‘‘seeds’’
in soil samples may be obfuscated by the rapid disintegration of the brown, ribbed fruit wall of the cypsela
which reveals a finely verrucose, dark blue to blackish
testa (Baskin and Baskin 1996; Fig. 3D). Although still
viable, the naked seeds (ripened ovules) are very
different in appearance and can be easily overlooked
or mis-identified.
Few studies have been conducted in Canada. The
plant did not germinate from any of the soil samples
from mature deciduous forests at Mont St. Hilaire, QC
(Leckie et al. 2000), although herbarium specimens have
been collected indicating its presence throughout the
area (herbarium specimen label data).
Profuse germination of E. hieraciifolius is often noted
in conjunction with water draw-downs, such as drained
beaver ponds, emergent shorelines and flood plains
(herbarium specimen label data). At Cold Spring
Harbor, NY, abundant germination of E. hieraciifolius,
along with more than 140 other species of plants, was
noted on the bed of a recently drained millpond (Shull
1914). Shull (1914) tried to artificially recreate similar
conditions in the laboratory, but did not observe any
germination of E. hieraciifolius. In a Connecticut salt
marsh, Roman et al. (1984) found that E. hieraciifolius
germinated well in areas where the soil water salinity
was near 0.5% throughout the growing season.
In North Carolina, Oosting and Humphreys (1940)
examined seed germination from soil samples along a
plant succession transect from cultivated fields to
climax oak-hickory forest; 10 sites were considered to
represent more than 200 yr of forest growth. No seeds
of E. hieraciifolius germinated from soil collected at
sites representing pre-afforestation communities, and
only small numbers of seedlings were detected at
forested sites. Although no data are given on the
above-ground presence of E. hieraciifolius in any of
the vegetation age classes, Oosting and Humphreys
(1940) state that it is improbable that seeds immigrating
from elsewhere would be able to readily penetrate litter
layers into the soil layers sampled. Livingston and
738
Allessio (1968) reported similar results along a successional transect in Massachusetts from abandoned fields,
through conifer plantations, to mixed woodland; only
4 seedlings were germinated from a total of 16 soil
samples, one each from 4 mid- to late-succession
samples. In a southern Appalachian mature oak forest
(Montgomery Co., Virginia) with a low level of past
disturbance, E. hieraciifolius was among pioneer species
that were either absent or present at very low densities
( 50.03 m2) in the seed bank (Schiffman and Johnson
1992). At sites where seeds were found in the soil (by
germination testing, but undetected in visual inspections), adult plants were also found in the forest
community; however, no germination of E. hieraciifolius was observed in soil samples from sites which
lacked adult plants (suggesting limited immigration). At
disturbed and undisturbed mixed oak forest sites in the
central Appalachians (southeastern Ohio), Schelling
and McCarthy (2007) germinated seeds of E. hieraciifolius from 79% of their soil samples, where it occurred
most frequently among the 70 species detected in the
seed bank. Adult plants were also found in the aboveground vegetation in the study area. A study of several
forest soil seed banks in eastern Tennessee (Farmer
et al. 1982) found that E. hieraciifolius was one of five
species germinating from 100% of the samples and
trials, that it produced the second highest number of
seedlings (mean of 127 425 ha1) of the 95 species
detected, and, after 56 mo of growth, represented
about 3242% (depending on substrate trial) of the
total biomass of all germinated plants. Germination
tests on litter and soil (to 5 cm depth) from a pine
(Pinus elliottii Englem.) plantation in South Carolina,
found seedling densities of 26 m 2 (Mou et al. 2005),
with almost all seedlings germinating from unscarified
soil samples.
The unsuitability of shade conditions in mature
closed-canopy forests, the scarcity of seeds in the litter
and soil, and the rapid germination of E. hieraciifolius
following cool burns and litter removal might suggest
that some seeds will remain dormant in the substratum
until surface conditions permit access to sunlight
(Glasgow and Matlack 2007), but might also suggest
immigration of wind-borne seeds from neighbouring
open areas (Matlack and Good 1990). For example, a
study of buried seeds after clear-cutting of an ancient
stand of Pinus strobus L. in Connecticut found that E.
hieraciifolius was not among plants germinated from
buried seeds, and that its substantial presence in the
clearing arose from wind-dispersal of its seeds from
open areas (Del Tredici 1977). Ecologists have come
to similar conclusions in Japan, where its explosive
occurrence as an early pioneer of post-disturbance
forest succession is attributed primarily to anemochory (wind dispersal) rather than seed banking
(Ohtsuka 1998).
Most studies of soil seed communities have been
conducted at upland sites. In a variety of wetland (bog)
communities in West Virginia, E. hieraciifolius seeds
germinated at rates of 055 seeds m 2 (McGraw 1987).
Unfortunately data were not presented on the vertical
distribution (and therefore relative age) of germinating
seeds. At an upland site in West Virginia, Landenberger
and McGraw (2004) studied seed banks at two forest
sites which had been clear-cut 7 yr previously. They
found E. hieraciifolius to be among the most abundant
seeds (relative to other species detected) in the soil in the
open clear-cut and that seed densities declined in
proportion to other species and logarithmically in
density across boundary between clear-cut and undisturbed forest.
Baskin and Baskin (1996) found that in Kentucky
about half of the freshly collected seeds were dormant
and the remainder only germinated under warmer
conditions. Unburied seeds which were winter-stratified
germinated the following spring, while buried seeds (also
stratified), germinated the April following exhumation.
Buried seeds re-entered conditional dormancy each
October, but 89% of seeds remained viable after 8 yr
of burial. Maximum germination occurred under thermoperiods of 30:15 and 35:208C (12:12 h) and a
photoperiod of 14 h light:10 h dark, although considerable germination also occurred in total darkness. In
Massachusetts, Lincoln (1983) found that 85% of
freshly harvested seed germinated in either light of
dark conditions at 20308C within 14 d following a
60-d cold treatment.
Although the vegetative community at the site of soil
provenance is not given, E. hieraciifolius germinated
from most of the 96 pots using soil from Ona, Florida,
in a greenhouse study (Stephenson and Rechcigl 1991).
It is not known whether germination was, in whole or in
part, from fresh (previous season) or dormant (seedbanked) seed, or whether seeds were from locally grown
plants or transported from elsewhere. Germination of
seed tended to increase with soil pH (4.55.7) when
augmented with dolomite, reaching a maximum at pH
5.7 and then diminishing somewhat at pH 6.16.8
achieved with gypsum addition. Also, plants growing
at low soil pH ( B5.2) were smaller and less vigorous.
(d) Vegetative Reproduction * Erechtites hieraciifolius
reproduces only by seed (Muenscher 1955).
9. Hybrids
No inter-specific hybrids have been reported.
10. Population Dynamics
The rapidity with which large numbers of plants of E.
hieraciifolius appear immediately after major disturbance, then quickly begin to decline over the following
2 or 3 yr, represents a classic example of an early pioneer
species in the ecological succession of plants. Pursh
(1814) noted the tendency of this plant to rapidly form
extensive monocultures in recently cleared forest areas,
particularly where the woody debris was burned in situ.
Farmer et al. (1982) observed densities of 49220 103
plants ha1 when testing for seed germination from
southern Appalachian forest soils. When they measured
the above-ground biomass 56 mo after germination
(representing plant growth in the first season postdisturbance), E. hieraciifolius comprised 3242% of the
total plant biomass, more than twice as much as
produced any of the other 94 species under any of the
soil condition treatments. In the first year of recolonization after total vegetation removal by herbicide
in a West Virginia forest, E. hieraciifolius comprised
22% of the ground cover (Kochenderfer and Wendel
1983). In the second year it had fallen to 2% cover and
was not detected in subsequent years. In a North
Carolina clear-cut forest, Boring et al. (1981) found
that E. hieraciifolius biomass production was 59 kg ha 1
during the season following cutting (i.e., removal of all
marketable timber), which represented 13.6% of herbaceous plant biomass and 3.4% of the total biomass
production by all woody and non-woody plants. It was
the only species making a substantial contribution to the
re-growth biomass which was not detected in pre-cut
vegetation surveys. At a nearby study site, Elliott et al.
(1998) reported that, in the first year of revegetation
after a severe disturbance, E. hieraciifolius biomass
production was 1820 kg ha1 and represented 37.1%
of the biomass of the ground flora. A subsequent survey
15 yr later did not detect any E. hieraciifolius, but after
28 yr it was detected as a minor component (0.9 kg
ha1, 0.2%). The occurrence of E. hieraciifolius in the
more mature forest stage may be due to the creation of
suitable microsites by local disturbances (Peterson et al.
1990; Peterson and Pickett 1990).
At a woodland site in South Carolina, highly disturbed because of the regular spray disposal of wastewater, E. hieraciifolius growth was correlated to the
amount of disturbance (Hunt and Shure 1980). In the
high disturbance zone it reached a peak biomass
of about 125 g m 2 in June, whereas in the lower
disturbance zone a peak of about 50 g m 2 occurred in
August. As an important component of the plant
community and insect host, the growth of E. hieraciifolius had a significant effect on the temporal patterns and
productivity of arthropod herbivores (see Section
13aiii).
It is not clear whether the colonizing ability of E.
hieraciifolius is primarily the result of seed dispersal and
immigration or long-term seed banking (see Section 8c).
For example, seeds of E. hieraciifolius have been shown
to be short-lived or transitory in mixed-oak hardwood
clearings in the southern Appalachians where they are
abundant in the early-successional period, but rapidly
decline and are replaced by more shade-tolerant species
with the growth of over-story trees and gradual canopy
closure in the late-successional period (Elliott et al.
1997). In managed pine plantations in Ohio, the
importance value (sum of relative cover and relative
739
frequency) of E. hieraciifolius was 0.6 in un-thinned
control plots and 9.5 in plots 3 yr after thinning (Abella
2010). In a managed pine plantation in South Carolina,
Mou et al. (2005), found the high importance value of E.
hieraciifolius (6.3) attained in the first post-harvest
growing season declined rapidly in the following 4 yr
(3.8, 0.5, 0, and 0.2, respectively).
11. Response to Herbicides and Other Chemicals
The leaf adaxial epidermis in E. hieraciifolius is essentially glabrous, lacks epicuticular wax, has a low water
drop contact angle (about 708), and has a high waterholding capacity (Haines et al. 1985b). This wettability
of the leaf surface promotes retention and penetration,
and therefore effectiveness, of spray-applied foliar
herbicides.
Little work has been done in Canada to assess the
response of E. hieraciifolius to herbicides. A single post
application of mesotrione at 101 g a.i. ha 1 early in the
season has provided good control of E. hieraciifolius in
New Brunswick cranberry fields (G. Graham, personal
communication, New Brunswick Department of Agriculture, Aquaculture and Fisheries, Fredericton, NB).
Mesotione also controls E. hieraciifolius well in wild
blueberry fields in Maine (D. Yarborough, personal
communication, University of Maine, Orono, ME),
although application rates are usually higher than in
Canada.
The species was among the first plants for which
herbicide tolerance was reported. After less than 10 yr of
heavy use of 2,4-D in sugar cane fields of Hawaii,
populations of E. hieraciifolius were showing reduced
response to the ‘‘new’’ chemical control (Hanson 1956,
1962).
Sulfometuron/hexazinone applied in granular form
was effective in early season control in pine plantations
(Pinus taeda L.) in the southeastern US (Gardiner et al.
1991). Sixty days after a late postemergence granule
application of sulfometuronhexazinone at two rates
(140.1 g a.i. ha 1700.5 g a.i. ha1 and 168.1 g a.i.
ha 1840.6 g a.i. ha1, respectively) 94% control of
the two major herbaceous weed species [E. hieraciifolius
and Eupatorium capillifolium (Lam.) Small] was obtained at a pine plantation site in Arkansas. By 120 days
after a late postemergence, control from the two
application rates decreased to 70 and 80%, respectively.
Several broad-spectrum specialty herbicides have
been registered in the United States which are claimed
to control E. hieraciifolius in turf or other ornamental
plantings. A mixture of clopyralid and triclopyr at a
recommended rate of 38 g a.e. ha1 and 51 g a.e. ha1,
respectively, is registered for postemergence control in
turf (Dow AgroSciences 2008). A mixture of dimethenamid-P and pendimethalin at a recommended rate of
1.7 kg a.i. ha 1 and 2.2 kg a.i. ha 1, respectively, is
registered for preemergence control in ornamental
plantings (BASF 2011). A mixture of dicamba, thiencarbazone-methyl and iodosulfuron-methyl-sodium at a
740
recommended rate of 78 g a.i. ha 1, 4.9 g a.i. ha1, and
22.5 g a.i. ha1, respectively, is registered for postemergence control in warm-season grass turf (BAYER
Undated). A mixture of soil-applied trifluralin and
isoxaben at a recommended rate of 3.4 kg a.i. ha 1
and 0.9 kg a.i. ha 1, respectively, is registered for
preemergence control in turf, nursery and ornamental
plantings (Dow AgroSciences 2010).
For container-grown nursery crops, Neal and Derr
(2005) provide a table on the efficacy of a number of
preemergence herbicides registered for use in ornamental plantings in the US. Of the 11 herbicides tested, they
rate 4 as providing ‘‘good’’ control of E. hieraciifolius:
flumioxazin (420 g a.i. ha1); oxyfluorfenoxadiazon
(2.21.1 kg a.i. ha 1); oxyfluorfenoryzalin (2.21.1
kg a.i. ha 1); and, oxyfluorfenpendimethalin (2.2
1.1 kg a.i. ha1).
12. Response to Other Human Manipulation
Prior to the advent of chemical herbicides, Dalaire
(1904) stated that control could be accomplished by
cultivation, hoeing and hand pulling. Even now in
cranberry and blueberry fields of the Maritime provinces control is often accomplished by hand pulling
(G. Graham, personal communication, New Brunswick
Department of Agriculture, Aquaculture and Fisheries,
Fredericton, NB). Manual pulling and mowing, prior
to seed production, has also been recommended for
control in the United States (Runnels and Schaffner
1931; Muenscher 1955). Diligent sanitation of harvesting equipment appears to reduce spread in wild blueberry fields in Maine (D. Yarborough, personal
communication, University of Maine, Orono, ME).
13. Response to Herbivory, Disease and Higher
Plant Parasites
(a) Herbivory
(i) Mammals, including both domestic and wild animals *
No reports have been found indicating whether the
bitter leaves with their unpleasant odour are palatable
to mammals.
(ii) Birds and other vertebrates * No information was
found.
(iii) Insects * The Chinese ladybeetle, Leis conformis
Boisd. (Coccinellidae: Coccinellini), imported into
Florida to combat the green citrus aphid, Aphis spiraecola Patch (Homoptera: Aphididae), has been found
feeding on the blossoms of E. hieraciifolius; although
entire blossoms including stamens and pistils were eaten,
no larvae, pupae or eggs were observed (Watson and
Thompson 1933). In a young pine forest in South
Carolina, increased biomass of E. hieraciifolius led to
substantial increases in arthropod trophic guilds, such as
sap feeders and leaf strippers from 51 mg m 2 in
untreated areas to up to 252 mg m 2 in treated areas.
Sharp declines in the herbivore biomass corresponded to
dieback of the E. hieraciifolius (and other herbaceous
plants) and increases in detritivore populations exploiting nutrients available in the litter.
Larvae of Palthis asopialis (Guenée) are reported to
feed on E. hieraciifolius, and a variety of other plants
(Robinson et al. 2010). This noctuid moth is known to
occur only in extreme southern Ontario (Troubridge
and Lafontaine 2004) and Quebec (J. D. Lafontaine,
personal communication, Agriculture and Agri-Food
Canada, Ottawa, ON). Larvae of Tyria jacobaeae L.
(Arctiidae), a biological control agent established in
Canada against Senecio jacobaea L. (Harris et al. 1975),
were reported to feed and mature on E. hieraciifolius in
forced feeding trials (Bucher and Harris 1961). Outside
North America, several species of moths have been
reported to include E. hieraciifolius in their host range
(Robinson et al. 2010), including Hypercompe icasia
Cramer (Arctiidae), Platyptilia molopias Meyrick (Pterophoridae), and Platphalonidia subolivacea Walsingham
(Tortricidae).
Evidence of leaf-mining insects is not uncommon on
herbarium specimens of E. hieraciifolius from Canada.
The microlepidoptera Phyllocnistis insignis Frey & Boll
(Gracillariidae) has been reported on E. hieraciifolius in
the US (Robinson et al. 2010). Although larvae
produce track mines in a variety of herbaceous
Asteraceae and the species has been reported from
several border States, it is not yet known to occur in
Canada. A lepidopteran miner on Erechtites was
reported from Kentucky under the name Phyllocnistis
erechtitisella Chambers, but no description was provided and the name cannot be used (Chambers 1878).
Reports of the Gracillariid Phyllonorycter insignis
(Walsingham) attacking Erechtites species are in error
(J.-F. Landry, personal communication, Agriculture
and Agri-Food Canada, Ottawa, ON).
In south Florida, E. hieraciifolius is among the many
species of Asteraceae suitable as hosts for the leaf miner
fly Phytobia maculosa (Malloch) [ Nemorimyza maculosa (Malloch)] (Agromyzidae), whose larvae form
large blotch mines on the leaves (Stegmaier 1967). In
the vicinity of vegetable crops in Florida, the highly
polyphagous serpentine leaf miners Liriomyza sativae
Blanchard and L. trifolii (Burgess) (Agromyzidae) are
found on E. hieraciifolius (Genung et al. 1978; Genung
1981; Schuster et al. 1991). In Taiwan, E. hieraciifolius
was found to be a preferred host of the leafminer L.
trifolii (Chien and Ku 1998). Although none of these
leaf-mining flies occur in the field in Canada, the latter is
a pest of greenhouse crops (Broadbent and Olthof 1995;
Dempewolf 2004; Lonsdale 2011).
In Arkansas, E. hieraciifolius is host to the tarnished
plant bug, Lygus lineolaris P. Beauv. (Heteroptera:
Miridae) (Young 1986). A gall midge, Neolasioptera sp.
(Diptera: Cecidomyiidae), has been reported on E.
hieraciifolius in Florida, causing irregular stem swellings 25 cm long; and, a flower-head gall produced by
Asphondylia sp. (Cecidomyiidae) has been reported on
Erechtites sp. from an unspecified North American
location (Gagné 1989). Infestation by the cosmopolitan
aphid species Brachycaudus helichrysi (Kaltenbach)
(Homoptera: Aphididae) was found on E. hieraciifolius
in two Florida counties in 1997 (Halbert et al. 2000).
Blatchley (1912) reported that a carabid beetle
[Anisodactylus terminatus (Say)] was often seen feeding
on ripening seeds in Indiana. In Kagoshima City,
Japan, E. hieraciifolius was one of three plants infested
by an Asian moth, Nyctemera adversata (Schaller)
(Lepidoptera: Arctiidae), with 7292% of the larvae
maturing in 27.5 d (Murakami et al. 1999). In host
testing of the Lepidopteran Secusio extensa (Butler) as
a biological control agent in Hawaii, E. hieraciifolius
was found to be a poor host, with less than 2% of
larvae completing development on the plant (Ramadan
et al. 2010).
(iv) Nematodes and other invertebrates * Several species
of root-knot nematodes have been reported on E.
hieraciifolius, including Meloidogyne enterolobii Yang &
Eisenback ( M. mayaguensis Rammah & Hirschamm),
M. javanica (Treub) and M. incognita (Kofoid & White)
in association with or proximity to agricultural crops
(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
systems (Ebsary and Eveleigh 1983).
(b) Diseases
(i) Fungi * There are few reports of fungi on E.
hieraciifolius in Canada (Ginns 1986). The powdery
mildew, Sphaerotheca fuliginea (Schltdl.) Pollacci [ Podosphaera fuliginea (Schltdl.) U. Braun & S. Takam.],
has been reported on E. hieraciifolius in Ontario
and Quebec (Parmelee 1977; DAOM specimens),
and Cicinobolus cesatii de Bary (Ampelomyces quisqualis 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
Rossman 2010).
A number of other phytopathogenic fungi have been
reported on E. hieraciifolius in the United States (Farr
and Rossman 2010) including downy mildews, Bremia
lactucae Regel (PA), and Peronospora halstedii Farl.
[ Plasmopara halstedii (Farl.) Berl. & de Toni] (MA,
MD, NJ, TX and WI); leaf spot fungi Cercospora
erechtitis G. F. Atk. (AL, FL, TX), Phyllosticta
erechtitis F. L. Stevens & P. A. Young (HI), Ramularia
wisconsina H. C. Greene (WI), and Septoria erechtitis
Ellis & Everh. (AK, DE, TX and WI); powdery
mildews Erysiphe cichoracearum DC. [ Golovinomyces
cichoracearum (DC.) V.P. Heluta] (MD), E. polygoni
741
DC. [E. betae (Vanha) Weltzien] (NY), Sphaerotheca
castagnei Lév. (VT), S. humuli (DC.) Burrill [Podosphaera macularis (Wallr.) U. Braun & S. Takam.]
(eastern states, IA, IL, MN and MT), and S. macularis
(Wallr.) P. Magnus [ Podosphaera macularis (Wallr.)
U. Braun & S. Takam.] (MT, OH); root-rot fungus
Phymatotrichum omnivorum Duggar [ Phymatotrichopsis omnivora (Duggar) Hennebert] (TX); the stemrot fungi Sclerotinia sclerotiorum (Lib.) de Bary (FL)
and Sclerotium rolfsii Sacc. [Athelia rolfsii (Curzi) C.
C. Tu & Kimbr.] (FL); and, the leaf gall Synchytrium
erechtitis M. T. Cook (LA). In other parts of the world
Ramularia wisconsina was recently reported on E.
hieraciifolius in Korea (Shin and Braun 2000), Cercospora erechtitis in Venezuela, Septoria erechtitis from
Cuba and the West Indies, and the leaf spot pathogen
Mycosphaerella erechtitidina Petr. & Cif. was reported
from Cuba, the Dominican Republic and the West
Indies (Farr and Rossman 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
and Plasmopara, to the rust genus Puccinia, and to
the powdery mildews Erysiphe and Sphaerotheca. In
Hawaii, E. hieraciifolius was among a majority of
plants tested which did not inhibit growth of the soil
fungus Phytophthora palmivora Butler (Powell and Ko
1986).
(ii) Bacteria * In laboratory testing Pseudomonas
cichorii (Swingle) Stapp and P. viridiflava (Burkholder)
Dowson were found to be pathogenic on E. hieraciifolius
(Tsuchiya et al. 1982). Both these bacterium species have
broad host ranges which include many crop species.
(iii) Viruses * In Florida, E. hieraciifolius was identified
as a host for the Bidens mottle potyvirus (BMV), which
was also infecting nearby lettuce (Lactuca sativa L.) and
endive (Cichorium endivia L.) crops (Purcifull and Zitter
1973).
(c) Higher Plant Parasites * No information has been
found.
ACKNOWLEDGEMENTS
J. Calder, G. Graham, J. D. Lafontaine, J.-F. Landry,
F. and K. Starr, and D. Yarborough are thanked for
kindly sharing information and expertise. Peter Craig, of
New Jersey, kindly provided the photograph in Fig. 5.
Useful comments on the manuscript were provided by J.
Cayouette (Agriculture and Agri-Food Canada) and
Mihai Costea (Wilfred Laurier University). We thank
the curators and staff of the following herbaria who
made material in their care available for study: ACAD,
CAN, DAO, HAM, LKHD, MT, MTMG, NSAC,
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Cover:
A flower head (capitulum) of Erechtities hieraciifolius, a native
species in the Asteraceae which is a weed in various agricultural
and horticultural situations, and often goes through enormous
population swings. The photograph shows the rayless but heterogamous
florets, the outer florets being pistillate, the medial florets
bisexual and the innermost florets functionally staminate. The two
style arms terminate in a ring of hairs at the base of a conical
appendage formed by the fusion of papillose hairs.

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