Cirsium arvense

• Overview
• Interesting Facts
• Common Names
• Description
• Biology
• Taxonomy
• Similar Species
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• Notes

Overview

Herb. Cirsium arvense is an herbaceous perennial in the aster family . It occurs in nearly every upland herbaceous community within its range , and is a particular threat in grassland communities and riparian habitats . C. arvense is shade intolerant and can tolerate soils with up to 2% salt content . It grows on all but waterlogged, poorly aerated soils, including clay , clay loam, silt loam , sandy loam, sandy clay, sand dunes, gravel , limestone, and chalk , but not peat. It spreads primarily by vegetative means, and secondarily by seed. The seeds spread as a contaminant in agricultural seeds in hay and in cattle and horse droppings and on farm machinery. It produces an abundance of bristly-plumed seeds that are easily dispersed by the wind and they may also be transported by water. Nuzzo (1997) reports that American Indians purportedly used an infusion of C. arvense roots for mouth diseases. The Chippewa considered it to be a "tonic, diuretic, and astringent". Young shoots and roots "can be used in the same ways as asparagus," and were eaten in Russia and by Native Americans. The nectar of its flowers is also said to make good honey. Zouhar (2001) reports that the weed has been used by native people in the northeastern United States in remedies for worms and poison-ivy (Toxicodendron radicans) and was used to make a mouthwash for children, a treatment for tuberculosis, and a tonic for gastrointestinal ailments.

Interesting Facts

Common Names

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Description

Family Asteraceae

Annuals , biennials, perennials , subshrubs , shrubs , vines , or trees . Roots usually taproots , sometimes fibrous . Stems usually erect , sometimes prostrate to ascending (underground stems sometimes woody caudices or rhizomes, sometimes fleshy ) . Leaves usually alternate or opposite, sometimes in basal rosettes, rarely in whorls; rarely stipulate , usually petiolate , sometimes sessile, sometimes with bases decurrent onto stems; blades usually simple (margins sometimes 1 2+ times pinnatifid or palmatifid ), rarely compound . Inflorescences indeterminate heads (also called capitula) ; each head usually comprising a surrounding involucre of phyllaries (involucral bracts ), a receptacle, and (1 ) 5 300+ florets; individual heads sessile or each borne on a peduncle; heads borne singly or in usually determinate, rarely indeterminate, arrays (cymiform, corymbiform , racemiform , spiciform , etc. ) ; involucres sometimes subtended by calyculi (sing. calyculus) ; phyllaries borne in 1 5( 15+) series proximal to (i.e. , outside of or abaxial to) the florets ; receptacles usually flat to convex , sometimes conic or columnar , either paleate (bearing paleae or receptacular bracts that individually subtend some or all of the florets) or epaleate (lacking paleae) ; epaleate receptacles sometimes bristly or hairy or bearing subulate enations among the florets. Florets bisexual , pistillate , functionally staminate , or neuter (also called neutral) ; sepals highly modifed (instead of ordinary sepals, each ovary usually bears a pappus of bristles , awns , and/or scales , sometimes in combination within a single pappus) ; petals connate , corollas (3 ) 5-merous, ± actinomorphic or zygomorphic (one or both kinds in a single head, see descriptions of radiate , discoid , liguliflorous, disciform, and radiant following) ; stamens (4 ) 5, alternate with corolla lobes , filaments inserted on corollas, usually distinct , anthers introrse , usually connate and forming tubes around styles (rarely filaments connate and anthers distinct; e.g. , Heliantheae, Ambrosiinae) ; ovaries inferior, 2-carpellate, and 1-locular with 1 basally attached, anatropous ovule ; styles 1 in each bisexual, functionally staminate, or pistillate floret; each style usually ringed at base by a nectary , distally 2-branched with stigmatic papillae borne on adaxial face of each branch in 2 separate or contiguous lines or in 1 continuous band (styles usually not branched in functionally staminate florets), style branches apically truncate or appendaged beyond the stigmatic bands or lines, appendages usually papillate to hirsute distally on abaxial (or abaxial and adaxial) faces. Fruits (technically cypselae, historically called achenes) usually dry with relatively thick, tough pericarps, sometimes beaked (rostrate ) and/or winged (alate ), often dispersed with aid from pappi. Seeds 1 per fruit, exalbuminous ; embryos straight.

Genera ca. 1500, species ca. 23,000 (418 genera, 2413 species in the flora ) : nearly worldwide, especially rich in numbers of species and/or in numbers of plants in arid and semiarid regions of subtropical and lower to middle temperate latitudes .

Asteraceae (Compositae, "composites," or "comps") have long been recognized as a natural group, and circumscription of the group has never been controversial (although some authors have divided the traditional family into three or more families) . A. Cronquist (1981) placed Asteraceae as the only family in the order Asterales within subclass Asteridae, associated with the Gentianales, Rubiales, Dipsacales, and Calycerales and relatively distant from Campanulales. On recent molecular phylogenetic data, the Angiosperm Phylogeny Group (2003; see references there for details; classification abbreviated APGII hereafter) has suggested that Asteraceae are better treated as part of a more widely defined Asterales within the asterids II informal clade (or campanulid clade; see W. S. Judd and R. G. Olmstead 2004) . Judd and Olmstead summarized the higher-order relationships of Asteraceae as follows (in order of decreasing inclusiveness; synapomorphies in parentheses) : asterids (ovules unitegmic and tenuinucellate , iridoid chemistry) ; core asterids (sympetaly, stamen number equal to petal number, stamen epipetaly, mostly 2 3-carpellate gynoecia) ; campanulids (early sympetaly), comprising eight unassigned families plus Aquifoliales, which is sister to Dipsacales, Apiales, and Asterales (last three sharing frequently inferior ovaries, polyacetylenes) ; and Asterales, which appears to be sister to Dipsacales-Apiales (K . Bremer et al. 2004) . The order Asterales (valvate petals, lack of apotracheal parenchyma, storage of inulin , ellagic acid present, and, possibly, the presence of a plunger or brush pollen presentation mechanism) now includes the following families (fide APGII) : Alseuosmiaceae, Argophyllaceae, Calyceraceae, Campanulaceae (optionally including Lobeliaceae), Goodeniaceae, Menyanthaceae, Pentaphragmaceae, Phellinaceae, Rousseauaceae, and Stylidiaceae. Within Asterales, Asteraceae is part of a clade (corollas with more or less fused lateral veins joining midvein near lobe apices, thick integuments, no endosperm haustorium) with the Menyanthaceae (cosmopolitan with Southern Hemisphere genera) basal to a more nested clade (inferior ovaries, possibly connate anthers, pollen exine with bifurcating columellae) comprising Asteraceae, Goodeniaceae (mainly Australia), and Calyceraceae (South America), the last being the immediate sister to Asteraceae (highly modified, persistent calyces, corolla venation patterns , unilocular and uniovulate gynoecia, pollen with intercolpar depressions , specialized fruits) . Aggregation of flowers into heads with involucres appears to have been a parallel phenomenon in Calyceraceae and Asteraceae, given the determinate nature of the former and indeterminate (racemose) organization of the latter. Some traits typical of Asteraceae predate evolution of the family as a distinct clade. Relationships of Asteraceae and Calyceraceae have been discussed by M. H. G. Gustafsson and Bremer (1995) . Synapomorphies of the Asteraceae clade include: calyces modified to structures called pappi, anthers connate (forming tubes) and styles modified to function as brushes in a specialized pollen presentation mechanism, ovaries each containing a single basal ovule, and production of sesquiterpene lactones .

K. Bremer et al. (2004) gave an Early Cretaceous origin for the Asteridae and the basal campanulids, and a Late Cretaceous origin for the Asterales. Bremer and M. H. G. Gustafsson (1997) also hypothesized a Late Cretaceous ancestry of Asterales in East Gondwanaland (Australasia), with later expansion into West Gondwanaland (South America-Antarctica), where the Asteraceae originated before the final separation of South America and Antarctica. Similarly, M. L. DeVore and T. F. Stuessy (1995) argued that the close relationships of Asteraceae to Goodeniaceae and Calyceraceae, plus the basal position of Barnadesioideae K. Bremer & R. K. Jansen (Asteraceae), indicated a South America-Antarctica-Australia origin for the complex . After reviewing previous hypotheses, they proposed a late Eocene origin for the complex and suggested a South American origin for the Asteraceae based on the basal position of the South American Barnadesioideae (see also Stuessy et al. 1996, on Barnadesioideae origin in southern South America in the Oligocene ) and their sister relationship to Calyceraceae. Fossil pollen data (both Mutisieae and Asteroideae types notably Heliantheae in the broad sense among earliest reports) reviewed by A. Graham (1996) appear to indicate an Eocene origin for Asteraceae in South America, with migration to North America at least by the Oligocene, possibly as early as the late Eocene. More recently, M. S. Zavada and S. E. de Villiers (2000; and references therein) reported Asteraceae pollen (assignable to Mutisieae in the broad sense) from the Paleocene-Eocene of South Africa, suggesting an earlier, West Gondwana (southern Africa or Australia) origin for the family. Such data indicate that some tribes of Asteraceae may have arrived in North America via long-distance dispersal or island hopping well before closure of the isthmus of Panama. They also have a bearing on the possible times of radiation of some tribes in North America, particularly Heliantheae in the broad sense and Eupatorieae, which originated in the continent (including Mexico and parts of Central America), and those that came to North America from or through South America such as Mutisieae, Vernonieae, some Plucheeae, and Astereae. Other tribes, such as Cynareae, Cichorieae, some Gnaphalieae, and Anthemideae, may have reached North America from Eurasia , possibly via Beringia (or as Amphi-Atlantic disjuncts ), at a later time.

The bases of a tribal classification within Asteraceae were established in the nineteenth century, primarily through the work of H. Cassini (especially in articles scattered through the 61 volumes of F. Cuvier 1816 1845; Cassini included synopses of his tribes as part of his entry for Zoegea, i.e., zyégée in French; the articles have been collected in three volumes by R. M. King and H. W. Dawson 1975), C. F. Lessing (1832), A. P. de Candolle (1828 1838, 1836 1838), and, particularly, G. Bentham (1873) . In the twentieth century, the tribal system of Cassini, as elaborated by Bentham, was widely followed with only slight modifications (see S. Carlquist 1976; A. Cronquist 1955, 1977; C. Jeffrey 1978; G. Wagenitz 1976b; see also J. Small 1919 and, for alternate views on Heliantheae-Eupatorieae, H. Robinson 1996) .

A molecular phylogenetic study by R. K. Jansen and J. D. Palmer (1987) established that a South American clade (later named Barnadesioideae) is basal within Asteraceae. Both cladistic morphologic analyses (e.g., K. Bremer 1994, 1996) and mostly chloroplast-DNA molecular phylogenies (e.g., Jansen et al. 1991, 1992; K. J. Kim et al. 1992; Kim and Jansen 1995; R. J. Bayer and J. R. Starr 1998; P. K. Eldenäs et al. 1999; B . G. Baldwin et al. 2002) have deepened our knowledge of tribal interrelationships within Asteraceae and led to the recent proposal of a phylogenetic classification for the family with 10 subfamilies and 35 tribes (J. L. Panero and V. A. Funk 2002) .

Treatment of Asteraceae here differs from some of the recently proposed classifications in that some groups continue to be traditionally circumscribed (e.g., Mutisieae in the broad sense, Heliantheae in the broad sense, including Helenieae and excluding Eupatorieae) . Where appropriate and so far as practicable, new taxonomies are acknowledged in our discussions of individual tribes (which see) . In North America, the following subfamilies and tribes, as defined by J. L. Panero and V. A. Funk (2002), are represented (tribes with no native representatives are marked by asterisks ) : Mutisioideae-Mutisieae in the strict sense, Gochnatioideae-Gochnatieae, and Hecastocleioideae-Hecastocleideae (all included in Mutisieae here, which see), Carduoideae (Cardueae = Cynareae), Cichorioideae (*Arctoteae, Cichorieae, Vernonieae), and Asteroideae [Senecioneae, *Calenduleae, Gnaphalieae, Anthemideae, Astereae, Plucheeae, *Inuleae, Eupatorieae, and the following segregates of Heliantheae in the broad sense (all treated here within or as subtribes of a fairly traditionally circumscribed Heliantheae) : Bahieae, Chaenactideae, Coreopsideae, Helenieae, Heliantheae in the strict sense, Madieae, *Millereae, Perityleae, Polymnieae, and Tageteae) ].

Asa Gray produced the first broadly influential floristic synthesis of North American Asteraceae. Other authors who made important contributions to floristics of North American Asteraceae in the nineteenth and first half of the twentieth centuries were S. F. Blake, N. L. Britton, R. S. Ferris, M. L. Fernald, E. L. Greene, H. M. Hall, M. E. Jones, D. D. Keck, P. A. Rydberg, J. K. Small, and S. Watson. Some of those authors had narrower concepts of genera and species than had their predecessors and they freely recognized new taxa in Asteraceae (mostly genera and species) . Floristics of North American Asteraceae in the second half of the twentieth century was especially influenced by A. Cronquist (e.g., 1955, 1980, 1994; H. A. Gleason and Cronquist 1991), who usually favored traditional generic circumscriptions.

In the last 20 years or so, developments in molecular systematics have led to revisions of generic limits in some tribes of Asteraceae and, sometimes, to a return to generic concepts that had been suggested earlier but largely ignored. More or less worldwide, taxonomies in some tribes or parts of tribes have included segregate genera that have been revived or newly published. Most of the innovations will be summarized in the forthcoming Asterales volume of K. Kubitzki et al. (1990+) . The generic circumscriptions adopted here incorporate recent taxonomic findings relevant to North America, insofar as our contributors have accepted them. As a result, many of the genera treated herein have never been presented in a major flora before, and some species are included within genera with which they were not associated traditionally. Thus, the Flora brings together much new knowledge and many new names . In most instances, circumscriptions of species have turned out to be conventional. So far as practicable, recently named species from North America have been accounted for within relevant treatments herein.

With 418 genera and 2413 species (Table 1), Asteraceae is, numerically, the largest family in the flora of North America north of Mexico. Members of the family are found in diverse habitats , from the High Arctic tundra and polar deserts to the Sonoran warm-desert scrub , and from alpine habitats to salt marshes. Asteraceae are particularly conspicuous elements of warm-desert and intermountain grasslands, as well as of desert scrubs, notably the intermountain desert scrub where Artemisia dominates (M. G. Barbour and N. L. Christensen 1993) . Among other conspicuous species, members of Solidago and Symphyotrichum form a very showy part of the fall flowering in eastern North America, and members of Heliantheae sometimes produce striking displays in the American West (e.g., Gaillardia spp. , Lasthenia spp., members of Madiinae) .

Much has been published, not only on systematics (at various levels), but on biology , chemistry, and economic and medical uses of Asteraceae worldwide, particularly in proceedings (from conferences and symposia) edited by V. H. Heywood et al. (1977), T. J. Mabry and G. Wagenitz (1990), and D. J. N. Hind et al. (1995, 1996) .

Relatively few North American species of Asteraceae are economically important or widely used ethnobotanically. The only major Asteraceae crop of North American origin is the sunflower, Helianthus annuus, which is valued for its seed oil and is appreciated in the horticultural trade. Other crop plants from native species worth mention are Helianthus tuberosus, the Jerusalem artichoke, and Parthenium argentatum, the guayule, a source of rubber. Echinacea spp. are touted as health plants. Members of several genera of Asteraceae native to the flora are grown for their ornamental value, notably species of Coreopsis (tickseeds), Echinacea (coneflowers), Helianthus (sunflowers), Liatris (blazingstars and gayfeathers), Rudbeckia (black-eyed Susans), Solidago (goldenrods), and Symphyotrichum ("asters" of the trade) .

Many species of Asteraceae have been introduced into North America, mainly from Europe and Asia, some deliberately for medicines, foods, or horticulture , others accidentally (often with seeds or other agricultural products or by other means) . Few, if any, of the introduced taxa are thought to be noxious at the continental level, but some (e.g., Acroptilon) are considered noxious in large parts of their ranges within the flora. Taraxacum officinale is a common lawn weed that (in terms of dollars spent and herbicides applied in weed control) has an economic and ecologic impact disproportionate to the actual harm it causes; other weedy introduced Asteraceae are of little economic consequence. Some native Asteraceae are toxic to cattle and other livestock and are therefore considered weeds. And some native species of open habitats (e.g., Symphyotrichum pilosum) are often considered weeds because they invade fields left fallow. Ragweeds (especially Ambrosia artemisiifolia and A. trifida) range over nearly the whole continent and their wind-blown pollens cause late-summer allergic reactions (hayfever) for a large number of people. Because ragweeds have a large impact on human health, they have a significant, negative economic impact.

In contrast to Orchidaceae, for which a wealth of excellent, well-illustrated popular books are available, few popular field guides on Asteraceae of North America have been published. The guide by T. M. Antonio and S. Masi (2001) deserves notice for its maps, color photographs, and useful information.

Composites (members of Asteraceae) share some unusual morphologic traits and some morphologic terms are used in particular ways as applied here to them.

For treatments of composites here, "perennials" are herbaceous and differ from annuals and biennials in living longer than two years and differ from subshrubs, shrubs, and trees in not developing woody aerial stems.

In most composites, leaf venation comprises a midrib plus more or less equal lateral nerves or veins; such leaves are described as pinnately nerved. Venation in leaf blades of some composites often consists of a midrib plus relatively strong lateral veins that diverge at or just distal to bases of blades. Such leaves are described as 3-nerved, 3( 5) -nerved, 5-nerved, etc., and, as appropriate, the phrases "from bases" or "distal to bases" may be added for clarification.

Composites often have subsessile to sessile or sunken glandular hairs that consist of multicellular bases supporting globular elements that usually contain resinous or sticky substances. Such structures have been called glands , glandular hairs, glandular trichomes, punctae, resin dots, and so on. Sometimes, the glands are embedded in epidermal depressions or pits. Epidermes with glands more or less sunk into or embedded within the surface have been called glandular-punctate and/or punctate-glandular. The glands may be colorless (translucent ) or yellowish to dark brown or orange and are sometimes more prominent on dried specimens than in living plants. In keys and descriptions here, gland-dotted refers to the presence of such glandular hairs, whether sessile or in depressions or pits (as appropriate, "in pits" or "sessile" may be added for clarification) .

Inflorescences of composites are called heads (or capitula, sing. capitulum) . Heads may be borne singly (i.e., not clearly associated with other heads on the same plant) or associated in arrays. The arrays of heads on composites correspond to arrays of individual flowers (inflorescences) on plants of other families; arrays of heads are sometimes called capitulescences . Terms for architectural structures of arrays of heads are parallel to terms for kinds of inflorescences: cymiform, corymbiform, paniculiform , racemiform, spiciform, thyrsiform, etc.

In radiate heads, peripheral florets (ray florets) in one or more series have corollas with zygomorphic limbs and may be pistillate, or styliferous and sterile , or neuter; the central florets (disc florets) in radiate heads have ± actinomorphic corollas and may be bisexual or functionally staminate. In liguliflorous heads, all florets are bisexual and (usually) fertile and have zygomorphic corollas (ligulate florets) ; liguliflorous heads are characteristic of Cichorieae and are found in no other composites. In discoid heads, all florets have ± actinomorphic corollas and all are either bisexual and fertile or all are either functionally staminate or pistillate (in monoecious or dioecious taxa, e.g., Baccharis spp.) . In disciform heads, all florets have ± actinomorphic corollas, and peripheral florets (in one or more series) are usually pistillate and usually have relatively slender (often filiform ) corollas. Such peripheral pistillate florets are generally thought to be derived by reduction from ray florets, and plants with disciform heads are generally thought to be derived from ancestors with radiate heads. The central florets of disciform heads are usually bisexual, sometimes functionally staminate. By tradition and for simplicity, both the peripheral, pistillate florets and the inner, bisexual or functionally staminate florets in disciform heads may be referred to as "disc" florets. In radiant heads, all florets have ± actinomorphic corollas and the peripheral florets usually have much enlarged corollas and may be bisexual, pistillate, or neuter; the central florets of radiant heads are usually bisexual. Some composites have peripheral, bisexual florets with slightly to strongly zygomorphic corollas (e.g., some members of Chaenactis, Lessingia, Thymophylla, et al.) ; heads of such plants do not quite conform to any of the five types just described and such heads may be referred to as "quasi-radiate" or "quasi-radiant." Some florets in heads of some Mutisieae have 2-lipped corollas and those heads may be called "quasi-radiate" or "quasi-liguliflorous." The term eradiate is used to refer collectively to discoid, disciform, and radiant heads.

Heads with all florets of one sexual form (bisexual, pistillate, or functionally staminate) are called homogamous (discoid and liguliflorous heads are homogamous , some radiant heads may be homogamous) and heads with florets of two or more sexual forms are called heterogamous (radiate and disciform heads are heterogamous, some radiant heads may be heterogamous) .

Phyllaries collectively constitute an involucre, usually number 5 21( 50+), usually are unequal (outermost usually shorter than the inner), and usually are arranged ± imbricately (overlapping like shingles) in 3 5( 15+), usually ± spiral series. Sometimes, the phyllaries are ± equal in 1 2 series; they are rarely wanting (e.g., Psilocarphus spp.) . Phyllaries may be herbaceous or chartaceous to scarious and are often medially herbaceous with chartaceous to scarious borders and/or apices. The phyllaries "proper" are sometimes immediately subtended by a calyculus (pl. calyculi) of (1 ) 3 15+ distinct, usually shorter bractlets in 1( 3+) series (e.g., Coreopsis spp., Taraxacum spp.) .

Receptacles may bear paleae (i.e., some or all florets are individually subtended by a bractlet called a palea or receptacular bract) . Collectively paleae have been called "chaff" and paleate receptacles have been described as "chaffy." Receptacles that bear paleae are referred to as paleate and receptacles that never bear paleae are referred to as epaleate. Epaleate receptacles sometimes bear subulate enations (e.g., some Gaillardia spp.) or bristles or subulate to linear scales (e.g., some Cynareae), or fine hairs (e.g., some Anthemideae) . Epaleate receptacles (and paleate receptacles that have shed their paleae) may be smooth or pitted (alveolate , foveolate, etc.) .

The terms tube, throat, and limb have been variously used in descriptions of corollas of composites. Here, in ± actinomorphic corollas of bisexual and functionally staminate disc florets, the tube is the part of the corolla proximal to the insertion of the staminal filaments, and the limb is the part that is distal to insertion of the filaments. The limb comprises, proximally, the throat and, distally, the lobes. The distinction between tube and throat hinges on insertion of filaments, not on external morphology.

The relatively flat portion of a corolla of a ligulate floret from a liguliflorous head (i.e., members of Cichorieae) is called a ligule; it terminates in 5 teeth or lobes. The relatively flat portion of a corolla of a ray floret is called a lamina; it terminates in 0 3( 4) teeth or lobes. More or less bilabiate corollas are characteristic of some members of Mutisieae and are seldom found in members of other tribes.

Fruits of composites have been called "achenes" because they resemble true achenes. Achenes are dry, hard, single-seeded fruits derived from unicarpellate, superior ovaries. Ovaries of composites are bicarpellate and inferior. Fruits derived from ovaries of composites are called cypselae (sing. cypsela, a term coined by C. de Mirbel in 1815) . Morphology of an ovary of a composite at flowering is often markedly different from the morphology of the mature fruit (cypsela) derived from that ovary. References to cypselae in keys and descriptions here almost always refer to mature fruits, not to ovaries at flowering.

Shapes of cypselae have been used in distinguishing among species, genera, and even subtribes of composites. In most composites, cypselae are ± isodiametric in cross section . In some composites, cypselae are characteristically ± lenticular to elliptic in cross section. Such cypselae are said to be compressed (or laterally flattened) if the longer axis of the cross section is ± parallel to a radius of the head (e.g., Verbesina spp.) . Cypselae are said to be obcompressed (or radially flattened) if the shorter axis of the cross section is ± parallel to a radius of the head (e.g., Coreopsis spp.) .

In composites, pappi (sing. pappus) are found where calyces are usually found on inferior ovaries; pappi have been shown to be greatly modified calyces. They show a great range of diversity and are often diagnostic for recognition of taxa, especially at rank of genus and below. The forms of individual pappus elements intergrade . For keys and descriptions here, the following distinctions are made: cross sections of bristles and awns are ± circular or polygonal and have the longer diameter of the cross section no more than 3 times the shorter diameter. Pappus elements with "flatter" cross sections (i.e., longer diameter more than 3 times the shorter diameter) are called scales, regardless of relative overall lengths and widths of the elements. As used here, "subulate scale" and "setiform scale" mean much the same as "flattened bristle" of some authors. Pliable to stiff pappus bristles with diameters less than ca. 50 µm are called fine bristles; pliable to stiff bristles with diameters 50 100 µm are called coarse bristles. Rigid pappus elements with ± circular or polygonal cross sections greater than 100 µm in diameter are called awns. Bristles, awns, and scales may be smooth or finely to coarsely barbed or plumose . A scale of a pappus may terminate in one or more bristlelike or awnlike appendages; such scales are said to be aristate.

In keys and descriptions, "pappus" and "pappi" usually refer to structures found on cypselae (mature fruits), not to "immature pappi" of ovaries at flowering. Sometimes pappi of ovaries that do not form fruits (e.g., in functionally staminate florets of some tarweeds) may be taxonomically useful and may be referred to in descriptions and keys.

Following is a synoptic key to tribes into which genera of composites of the flora area are placed. Keys to genera within each tribe will be found in the accounts of the individual tribes. Because some traits in the key to tribes and in keys to genera within tribes may be difficult to assess, we have also provided a key to artificial groups of composites and keys to genera within those artificial groups. Those keys will be found following the key to tribes.

In the following key, "radiate heads" have ray florets; "eradiate heads" lack ray florets and may be disciform, discoid, or radiant. Ray florets have zygomorphic corollas with laminae ; the laminae may be showy (as in some species of Helianthus) or inconspicuous (as in some species of Erigeron) . Usually, we have included plants with inconspicuous ray laminae in keys to genera of both radiate and eradiate groups.

Some plants have questionably paleate or epaleate receptacles. Epaleate receptacles of some plants are notably pitted and have fimbriate to deeply lacerate pit borders ; such receptacles have sometimes been interpreted as paleate. Plants with notably lacerate pit borders are usually keyed here as both paleate and epaleate.

Some plants with pappi of conspicuous bristles often have the bristles subtended by minute, inconspicuous scales. Although such plants technically belong to groups with pappi "wholly, or partially, of awns or scales," they are usually also keyed here in groups characterized as having pappi "wholly of bristles," because the scales are easily overlooked. As well, some pappus elements are borderline between being called subulate or setiform scales or being called "flattened bristles." Consequently, some plants that technically belong to groups with pappi of scales are keyed both in groups with pappi "wholly of bristles" and in groups with pappi "wholly, or partially, of awns or scales."[1]

Genus Cirsium

Annuals , biennials, or perennials , 5-400 cm, spiny . Stems (1-several) erect , branched or simple , sometimes narrowly spiny-winged. Leaves basal and cauline; finely bristly-dentate to coarsely dentate or 1-3 times pinnately lobed , teeth and lobes bristly-tipped, faces green and glabrous or densely gray-canescent, usually eglandular . Heads discoid , borne singly, terminal and in distal axils, or in racemiform , spiciform , subcapitate , paniculiform , or corymbiform arrays. ( Peduncles with ± reduced leaflike bracts.) Involucres cylindric to ovoid or spheric, (1-6 ×) 1-8 cm. Phyllaries many in 5-20 series, subequal or weakly to strongly, outer and middle with bases appressed and apices spreading to erect, usually spine-tipped, innermost usually with erect, flat, often twisted, entire or dentate, usually spineless apices (distal portion of phyllary midveins in many species with elongate , glutinous resin gland , usually milky in fresh material but dark brown to black when dry) . Receptacles flat to convex , epaleate, covered with tawny to white bristles or setiform scales . Florets 25-200+; corollas white to pink, red, yellow or purple, ± bilateral , tubes long, slender, distally bent, throats short, abruptly expanded. cylindric, lobes linear ; (filaments distinct ) anther bases sharply short-tailed, apical appendages linear-oblong; style tips elongate (as measured in descriptions including the slightly swollen nodes, long cylindric fused portions of style branches and very short distinct portions) . Cypselae ovoid, ± compressed , with apical rims, smooth , not ribbed , glabrous, basal attachment scars slightly angled ; pappi persistent or falling in rings , in 3-5 series of many flattened, plumose bristles or plumose, setiform scales (longer bristles shorter than corollas except in C. foliosum and C. arvense) . x = 17.

Species ca. 200: North America, Eurasia , n Africa.

Only three genera in Cynareae are represented by native species in the New World, and of these Cirsium is by far the most widely distributed and diverse . Native species of Cirsium range from sea level to alpine and from boreal regions of Canada to the tropics of Central America. Members of the genus occur in a myriad of habitats including swamps , meadows, forests , prairies, sand dunes, and deserts.

Preliminary molecular phylogenetic studies by D. G. Kelch and B . G. Baldwin (2003) indicated that this diversity is the product of a rapid evolutionary diversification based upon a single initial introduction from Eurasia. Relationships among the North American species are apparently complex , and molecular studies have only begun to provide an outline of phylogeny for these plants . Although there has been a remarkable evolutionary and morphologic diversification in North American Cirsium, it has not been accompanied by very much divergence in the base sequences of genes commonly used to elucidate phylogenetic relationships. This suggests either that the diversification has been very rapid or that genetic markers in North American Cirsium mutate more slowly than in most other lineages .

Chromosomal diversification has accompanied the morphologic radiation of North American Cirsium. Many New World Cirsium species share the chromosomal base number of x = 17 that also predominates in most Eurasian species. Among the North American thistles, however, is a mostly descending dysploid series with chromosome numbers ranging from n = 18 to n = 10. Very few instances of polyploidy are known among New World Cirsium.

Cirsium species of remarkably different morphologies often are able to hybridize . Although in some hybrid combinations fertility is reduced, in others the formation of complex hybrid swarms indicates a lack of breeding barriers and the potential for emergence of novel character combinations. In the absence of adequate sampling and field observations, hybrids may go unrecognized, treated as distinct taxa or as variants of non-hybrid taxa, or left occupying the indeterminate folders of herbaria. In other cases hybridization has been invoked without much evidence as an explanation for Cirsium variants encountered in herbaria or in the field. Hybrid combinations are listed herein when evidence is convincing. Additional hybrids are likely to be found where the ranges of Cirsium species overlap. I have seen no documentation of hybridization between native American Cirsium species and introduced Eurasian taxa.

Much of the geographic range currently occupied by New World Cirsium species was greatly affected by the events of the Quaternary . Large areas were glaciated and other areas were vastly different during glacial episodes. The ancestors of thistles that currently occupy the high mountains of western North America were undoubtedly displaced elevationally and/or latitudinally during the recurrent glacial and interglacial episodes of the Pleistocene . Taxa that are currently isolated may have been in contact during glacial episodes with the opportunity for hybridization and genetic interchange. Episodes of prehistoric hybridization may have led to some of the character combinations found in modern American thistles, particularly in the western half of the continent. Current isolation and localized selection or genetic drift apparently have promoted differentiation of populations separated on mountaintop islands.

One of the most challenging aspects for a taxonomist studying New World Cirsium is the presence of species complexes that are apparently evolutionary works in progress. Some of the thistles, especially in the mountainous western part of North America, are frustratingly polymorphic with much overlapping variability and intergradation of characters. Early taxonomists, basing their work on a limited sampling of the morphologic diversity, named many of the forms as species, and the literature is rife with species names . The infilling that results from more collectors visiting more localities within the ranges of these complexes has blurred the boundaries between many of the proposed species and often added forms that do not "fit" the characteristics of named species. As I faced the challenges of preparing this treatment, I recognized that maintaining some of the named entities as species would, for consistency, require a further proliferation of species names. I have chosen to go the other way. Instead of proposing yet more ill-defined microspecies, I have chosen to recognize that the groups in question are rapidly evolving, only partially differentiated assemblages of races that have not reached the level of stability that is usually associated with the concept of species. Certainly much of such variation within the genus deserves a level of taxonomic recognition, or at least should be mentioned, but for those assemblages I think it much more prudent to recognize varieties -- entities that may be expected to freely intergrade -- rather than species.

Many problems remain to be worked out in North American Cirsium. Further investigation will undoubtedly reveal the need for refinement or major revision within some of the species groups. Studies that focus on variation within and among populations and on the biological basis for the variations are much needed. The field is open and the challenges are many.

Preparation of a workable key to Cirsium species has been frustratingly difficult. Extensive and overlapping ranges of variation in morphologic characteristics often require that a species be keyed two or more times. The resulting key is longer and more complex than I would prefer, and I have no doubt ignored, overlooked, or been completely unaware of variants that will not key out. Caveat clavitor!

The reputation of Cirsium has suffered greatly as a result of the introduction to North America of a few invasive weedy species from Eurasia. Cirsium vulgare (bull thistle) and C. arvense (Canada thistle€”a misnomer) have long been despised as noxious weeds . In recent years C. palustre (European swamp thistle) has joined their ranks . Additionally, weedy Eurasian species of Carduus, Onopordum, Centaurea, etc. , add to the public perception that all thistles are bad. Most North American native Cirsium are not at all weedy, and many are strikingly attractive plants. All are spiny plants that command respect, but they deserve a better reputation as one of North America€™s evolutionary success stories.

Native Cirsium species have come under threat from biocontrol programs instituted to suppress populations of weedy introduced thistles. Beginning in 1968 the seedhead weevil Rhinocyllus conicus has been widely introduced in various areas of the United States and Canada, primarily to control weedy species of Carduus. S. M. Louda et al. (1997) reported that R. conicus has crossed over to several native species of Cirsium. They observed that the number of viable cypselae in infested heads was greatly reduced; e.g. , heads of C. canescens infested by R. conicus produced 14.1 percent of the number of viable cypselae as in uninfested heads. Not all taxa are impacted as much as C. canescens, particularly those with later flowering phenology (Louda 1998) . R. W. Pemberton (2000) reported that 22 Cirsium taxa in North America are known hosts of R. conicus. I suspect that the number is higher. During my field work I have observed that the heads of many Cirsium species are heavily parasitized, although I have not determined which of these are infested by R. conicus and which by native seedhead parasites. The long-term impacts of R. conicus and other biocontrol agents on native thistles, particularly rare taxa, remain to be determined.[2]

Physical Description

Habitat

Disturbed sites, fields , pastures, roadsides, forest openings; 0-2600 m ; introduced .

Nuzzo (1997) cites that C. arvense occurs in nearly every upland herbaceous community within its range , and is a particular threat in prairie communities and riparian habitats . Throughout its range it is common on roadsides, in oldfields, croplands, and pastures, in deep, well-aerated, mesic soils. In eastern North America, it occasionally occurs in relatively dry habitats, including sand dunes and sandy fields, as well as on the edges of wet habitat, including stream banks, lakeshores, cleared swamps , muskegs and ditches. It is shade intolerant . It grows on all but waterlogged, poorly aerated soils, including clay , clay loam, silt loam , sandy loam, sandy clay, sand dunes, gravel , limestone, and chalk , but not peat. Zouhar (2001) reports that it can tolerate soils with up to 2% salt content . It grows best between 0 - 32 °Celsius. It tolerates annual precipitation ranging from 305-1015 mm per year and grows best with 400-750 mm of precipitation per year.

Typically found at an altitude of 0 to 2,848 meters (0 to 9,344 feet).[3]

Biome: Agricultural areas, disturbed areas, riparian zones, urban areas, wetlands.

Ecology: Nuzzo (1997) states that C. arvense threatens natural communities by directly competing with and displacing native vegetation, decreasing species diversity , and changing the structure and composition of some habitats . Species diversity in an "undisturbed" Colorado grassland was inversely proportional to the relative frequency of C. arvense. It presents an economic threat to farmers and ranchers. Infestations reduce crop yield through competition for water, nutrients and minerals, and through interference with harvest . In Canada, the major impact of C. arvense is in agricultural land, and in natural areas that have been disturbed or are undergoing restoration . In the United States, it is a host for bean aphid and stalk borer , insects that affect corn and tomatoes, and for sod-web worm, which damages corn. In Bulgaria, C. arvense is a host for the cucumber mosaic virus. In addition to reducing forage and pasture production , it may scratch grazing animals, resulting in small infections . Zouhar (2001) reports that it has been identified as a management problem in many national parks and on TNC (The Nature Conservancy) preserves in the upper Midwest, the Great Plains states, and the Pacific Northwest. Infestations of C. arvense may contribute to the elimination of endangered and/or endemic plant species, such as the Colorado butterfly plant in Wyoming.

Biology

Reproduction

Nuzzo (1997) states that the weed spreads primarily by vegetative means (by its root ), and secondarily by seed. The root system can be extensive, growing horizontally as much as 6 m in one season , and individual roots live up to two years. Most patches spread at the rate of 1-2 m/year. Under good growing conditions, female plants produce an average of 29 flowering shoots/square meter, each with an average of 41 heads/shoot and 59 seeds/head. A single plant produces an average of 1500 and up to 5300 seeds. Multiple plants produced 100-64,300 viable seeds/m2 in Australia and up to 30,200/m2 in Holland.

Germination may be affected by ecotype, temperature , day length , depth of seed burial, substrate stratification , and seed freshness. Seeds from "male" plants are smaller and percent germination is lower. Seeds germinate best in warm temperatures 20 - 40 degrees Celsius, with alternating light and dark periods. At lower temperatures germination is aided by high light intensity . Germination at higher temperatures can help ensure that maximum germination takes place during warmer periods of the year. Seeds are somewhat tolerant of heat, and some were still viable after 10 minutes at 102 degrees Celsius and 2 minutes at 262 degrees Celsius, although viability was decreased at these temperatures compared to unheated controls . The seeds germinate over a wide range of soil moisture.

Duration: Biennial, Perennial

Growth

Culture: Space 15-18" apart.

Soil: Minimum pH: 5.1 • Maximum pH: 9.0

Sunlight: Sun Exposure: Full Sun .

Moisture: Drought Tolerance: High

Temperature: Cold Hardiness: 3a, 3b, 4a, 4b, 5a, 5b, 6a, 6b, 7a. (map)

Taxonomy

Unambiguous Synonyms

1. Breea arvensis (L.) Less.
2. Carduus arvensis (L.) Robson
3. Cephalonoplos arvense (L.) Fourr.
4. Cirsium arvense var. argenteum (Vest) Fiori
5. Cirsium arvense var. horridum Wimmer and Grab.
6. Cirsium arvense var. integrifolium Wimmer and Grab.
7. Cirsium arvense var. mite Wimmer and Grab.
8. Cirsium arvense var. vestitum Wimmer and Grab.
9. Cirsium incanum (Gmel.) Fisch.
10. Cirsium setosum (Willd.) Bess. Ex Bieb.
11. Cnicus arvensis (L.) Hoffm.
12. Serratula arvensis L.

Notes

Name Status: Accepted Name . Latest taxonomic scrutiny: 15-Mar-2000.

Place of publication : Fl. carniol. ed. 2, 2:126. 1772

Name verified on 25-May-2007 by ARS Systematic Botanists. Last updated: 25-May-2007.

Similar Species

Members of the genus Cirsium

ZipcodeZoo has pages for 763 species, subspecies, varieties, forms, and cultivars in this genus. Here are just 100 of them:

C. abukumense · C. abyssinicum · C. acanthodontum · C. acantholepis · C. acarna · C. acaule · C. acaulescens · C. acaule acaule · C. acaule gregarium · C. acaulos · C. acrolepis · C. adjaricum · C. aduncum · C. affine · C. afrum · C. aggregatum · C. aidzuense · C. alatum · C. alberti · C. albescens · C. albicans · C. albidum · C. albowianum · C. aleutrense · C. allionii · C. alpestre · C. alpicola · C. alpinum · C. alpis-lunae · C. alsophilum · C. altissimum (Tall Thistle Cirsium Altissimum) · C. amani · C. ambiguum · C. amblylepis (Mt. Tamalpais Thistle) · C. americanum · C. amplexifolium · C. amplum · C. anartiolepis · C. anatolicum · C. andersonii (Anderson's Thistle) · C. andrewsii (Franciscan Thistle) · C. anglicum · C. angustifolium · C. aomorense · C. apendiculatum · C. apoense · C. appendiculatum · C. arachnoideum · C. araneans (Jeweled Thistle) · C. araricum · C. arctioides · C. arcuum (Powderpuff Thistle) · C. arenesi · C. argentum · C. argillosum · C. argyracanthum · C. argyrancanthum · C. aridum (Cedar Rim Thistle) · C. arisanense · C. aristatum · C. aristitans · C. arizonicum (Arizona Thistle) · C. arizonicum var. arizonicum (Arizona Thistle) · C. arizonicum var. bipinnatum · C. arizonicum var. nidulum (Arizona Thistle) · C. arizonicum var. rothrockii · C. arizonicum var. tenuisectum · C. armatum · C. armenum · C. arvense (Californian Thistle) · C. arvense albiflorum · C. arvensis · C. ashinokuraense · C. ashiuense · C. asiaticum · C. aspinellum · C. austrinum · C. autareticum · C. babanum · C. balearicum · C. barnebyi (Barneby's Thistle) · C. baytopae · C. belingschanicum · C. bertolonii · C. bicentenariale · C. bipinnatum · C. bipontinum · C. bitchuense · C. bohemicum · C. bolocephalum · C. boluense · C. boninense · C. borderi · C. boreale · C. borealinipponense · C. botryodes · C. boujarti · C. boujartii · C. bourgaeanum · C. bourgaei

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Further Reading

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• Canada Thistle (Cirsium arvense L.) Washington State Noxious Weed Control Board. Identification/Description; Photographs; Introduction History; Impacts; Life Cycle; Habitat; Distribution; Controls; Special Note: References
• Canada Thistle (Cirsium arvense) USDA. APHIS. Cooperative Agricultural Pest Survey. National Agricultural Pest Information System.Photographs; Distribution; Special Note: Links to other sites
• Canada Thistle (PDF | 224 KB) lberta Invasive Plants Council (Canada). Identification/Description; Photographs; Illustrations; Controls
• Canada Thistle (Thunhorst and Swearingen, 1997)
• Canada Thistle - Best Management Practices Weed Profile (PDF | 217 KB) Colorado State Parks.Identification/Description; Illustrations; Introduction History; Impacts; Habitat; Distribution; Controls; Special Note: References
• Canada Thistle - Biological Control of Invasive Plants in the Eastern United States USDA. Forest Service. Taxonomy; Identification/Description; Photographs; Distribution; Introduction History; Impacts; Life Cycle; Habitat; Controls
• Canada Thistle - Classical Biological Control of Weeds, Biology of Target Weeds Agriculture and Agri-Food Canada. Lethbridge Research Centre.Identification/Description; Photographs; Life Cycle;Controls; Special Note: References
• Canada Thistle - Guide to Weeds in British Columbia British Columbia Ministry of Agriculture Food and Fisheries (Canada). Identification/Description; Photographs; Illustrations; Impacts; Life Cycle; Habitat; Distribution; Dispersion; Controls
• Canada Thistle - Idaho's Noxious Weeds Idaho Association of Soil Conservation Districts. Idaho OnePlan. Identification/Description; Photographs; Introduction History; Distribution; Controls
• Canada Thistle - Invasive Alien Plant Species of Virginia (PDF | 98 KB) Virginia Department of Conservation and Recreation. Natural Heritage Program and Virginia Native Plant Society. Identification/Description; Illustrations; Impacts; Life Cycle; Habitat; Distribution; Controls
• Canada Thistle - Invasive Exotics (PDF | 49 KB) Bowman's Hill Wildflower Preserve (Pennsylvania).Illustrations; Impacts; Dispersion; Controls
• Canada Thistle - Invasive Plants of Ohio Ohio Department of Natural Resources. Natural Areas and Preserves.Identification/Description; Photographs; Introduction History; Impacts; Habitat; Distribution; Controls
• Canada Thistle - Missouri Vegetation Management Manual Missouri Department of Conservation.Identification/Description; Illustrations; Impacts; Life Cycle; Habitat; Distribution; Controls
• Canada Thistle - Noxious Weeds of Nebraska (PDF | 377 KB) University of Nebraska - Lincoln. Cooperative Extension. Identification/Description; Photographs; Introduction History; Life Cycle; Distribution; Dispersion; Controls
• Canada Thistle - Ohio Perennial and Biennial Weed Guide Ohio State University. Ohio Agricultural Research and Development Center. Identification/Description; Photographs; Introduction History
• Canada Thistle - Vegetation Management Guideline Illinois Nature Preserves Commission. Identification-Description; Photographs; Impacts; Life cycle; Distribution; Controls
• Canada Thistle - Virginia Tech Weed Identification Guide Virginia Polytechnic Institute and State University. Plant Pathology, Physiology, and Weed Science. Identification/Description; Photographs; Life Cycle
• Canada Thistle - Wisconsin Botanical Information System Wisconsin State Herbarium; University of Wisconsin-Madison. Taxonomy; Photographs; Habitat; Distribution
• Canada Thistle Colorado State University. Cooperative Extension. Identification/Description; Illustrations; Impacts; Life Cycle; Habitat; Dispersion; Controls
• Canada Thistle Colorado Weed Management Association.Identification/Description; Photographs; Life Cycle; Habitat; Distribution; Dispersion
• Canada Thistle Invasive.org. Photographs; Special Note: Resources
• Canada Thistle Montana Weed Control Association. Identification/Description; Photographs; Introduction History; Impacts; Life Cycle; Habitat; Dispersion; Controls
• Canada Thistle Official Control Program Kansas Department of Agriculture. Plant Protection and Weed Control Program. Identification/Description; Dispersion; Controls
• Canada Thistle Plant Conservation Alliance. Alien Plant Working Group. Identification/Description; Photographs; Introduction History; Impacts; Habitat; Distribution; Dispersion; Controls
• Canada Thistle State of Colorado. Larimer County. Weed Control District. Identification/Description; Photographs; Controls
• Canada Thistle Wanted Poster--Dead, Not Alive! (PDF | 211KB) University of Nevada - Reno. Cooperative Extension. Identification/Description; Photographs
• Canada Thistle Wisconsin Department of Natural Resources. Identification/Distribution; Photographs; Introduction History; Impacts; Habitat; Distribution; Control
• Canada thistle - Invasive Plant Species of Alaska (PDF | 49 KB) University of Alaska - Anchorage. Alaska Natural Heritage Program. Identification/Description; Photographs; Impacts; Life Cycle; Distribution; Dispersion; Controls
• Canada thistle - Noxious Weed Information Series (PDF | 274 KB) State of Colorado. Jefferson County. Weed and Pest Management Program. Identification/Description; Photographs; Habitat; Controls
• Candian Thistle - BioBullies (Western Pennsylvania) Natural Biodiversity. Identification/Description; Photographs; Introduction History; Impacts; Life Cycle; Habitat; Controls
• Carnegie Institution of Washington publication. Washington, Carnegie Institution of Washington, 1902- ENG url p. 144, p. 378, p. 59.
• Caroli Linnaei Opera. Editio prima critica, plena, ad editiones veras exacta, textum nullo rei detrimento contractum locosque editionum discrepantes exhibens. Vol. 2., Systema vegetabilium; libros diagnostico-botanicos co Lipsiae, Sumptum fecit O. Wigand, 1835 [i.e. 1835-1840] LAT url p. 784, p. 784.
• Catalog of Ohio vascular plants: arranged according to the phyletic classification: with notes on the geographical distribution in the state, based mainly on specimens in the State Herbarium, Botanical Laboratory, the Ohio State Uni by John H. Schaffner Columbus: Ohio State University, 1914. ENG url p. 235.
• Catalogue des plantes vasculaires indigènes, ou généralement cultivées en Corse suivant l'ordre adopté dans la flore de France de mm. Grenier et Godron avec l'indication des stations et des époques de floraison, par L.J.A. de C. de Marsilly, avec le conco Paris, G. Masson, 1872. FRE url p. 86.
• Catalogue of scientific papers, 1800-1900. Compiled by the Royal Society of London. London, C.J. Clay and Sons, 1867-1902 [etc.] Cambridge, Cambridge University Press, 1914-25. ENG url p. 39, p. 497, p. 818, p. 863.
• Catalogue of the books, manuscripts, maps and drawings in the British Museum (Natural History) London: BM(NH), 1903-1940. ENG url p. 1737.
• Catalogue raisonn des champignons des Pays-Bas / par C.A.J.A. Oudemans. Amsterdam: J. Mller, 1904 [i.e. 1905] FRE url p. 192.
• Catalogus dipterorum hucusque descriptorum auctore Dr. C. Kertesz. Lipsiae: G. Engelmann, 1902. LAT url p. 144.
• Chinese Academy of Sciences. 1959 –. Flora reipublicae popularis sinicae. (F China)
• Cirsium arvense (Canada Thistle) Nature Conservancy. Global Invasive Species Initiative. Identification/Description; Photographs; Impacts; Life Cycle; Habitat; Distribution; Dispersion; Controls; Special Note: Element Stewardship Abstract with excellent set of references
• Cirsium arvense (Canada thistle) - Invasive Plant Atlas of New England (IPANE) University of Connecticut. Ecology and Evolutionary Biology. Taxonomy; Identification/Description; Photographs; Introduction History; Distribution; Dispersion; Controls
• Cirsium arvense (L.) Scop. (Canada Thistle) - Noxious and Nuisance Plant Management Information System (PMIS) United States Army Corps of Engineers. Engineer Research and Development Center. Environmental Laboratory. Identification/Description; Photographs; Introduction History; Impacts; Distribution
• Cirsium arvense (L.) Scop. - Germplasm Resources Information Network (GRIN) USDA. ARS. National Genetic Resources Program. Taxonomy
• Cirsium arvense (herb) - ISSG Global Invasive Species Database World Conservation Union. Invasive Species Specialist Group. Taxonomy; Identification/Description; Photographs; Impacts; Life Cycle; Habitat; Distribution; Controls
• Cirsium arvense - Fire Effects Information System (FEIS) USDA. Forest Service. Rocky Mountain Research Station. Fire Sciences Laboratory. Taxonomy; Identification/Description; Photographs; Introduction History; Impacts; Life Cycle; Habitat; Distribution; Controls; Special Note: References
• Cirsium arvense - Species Abstracts of Highly Disruptive Exotic Plants at Effigy Mounds National Monument DOI. Northern Prairie Wildlife Research Center. Identification/Description; Life Cycle; Habitat; Dispersion; Controls
• Cirsium arvense Article Citation Search - AGRICOLA Database USDA. National Agricultural Library. Research; Special Note: NAL Catalog Search (resources)
• Cirsium arvense California Invasive Plant Council. Identification/Description; Photographs; Introduction History; Impacts; Life Cycle; Distribution; Dispersion; Controls
• Cirsium arvense Discover Life. Identification/Description; Photographs; Introduction History; Impacts; Life Cycle; Habitat; Distribution; Dispersion; Controls
• Cirsium genus - EncycloWeedia California Department of Food and Agriculture.Identification/Description; Photographs; Habitat; Distribution; Dispersion; Controls
• Coleoptera Neerlandica: de schildvleugelige insecten van Nederland en het aangrenzend gebied / door Éd. Everts. s'Gravenhage: Nijhoff, 1898-1922. DUT url p. 515, p. 519, p. 609.
• Comparative morphology and biology of the fungi, mycetozoa and bacteria / by A. de Bary; the authorized English translation by Henry E. F. Garnsey; rev. by Isaac Bayley Balfour. Oxford: Clarendon Press, 1887. ENG url p. 359.
• Compendio della flora italiana; ossia, Manuale per la determinazione delle piante che trovansi selvatiche od inselvatichite nell'Italia e nelle isole adiacenti. Torino, E. Loescher, 1882. ITA url p. 524.
• Compte rendu / Association franaise pour l'avancement des sciences. Paris, Secretariat de lAssociation. FRE url p. 157.
• Comptes rendus hebdomadaires des s©ances de l'Acad©mie des sciences. Paris: publi©s avec le concours du Centre national de la recherche scientifique par MM. les secr©taires perp©tuels: -1965. FRE url p. 917.
• Comptes rendus hebdomadaires des séances de l'Académie des sciences. Paris: publiés avec le concours du Centre national de la recherche scientifique par MM. les secrétaires perpétuels: -1965. FRE url p. 925.
• Control Practices for Canada Thistle Purdue University. Botany and Plant Pathology. Photographs; Life Cycle; Controls
• Czerepanov, S. K. 1995. Vascular plants of Russia and adjacent states (the former USSR). (L USSR)
• Danmarks fauna, Biller. København: G. E. C. Gad, 1908- DAN url p. 133, p. 322.
• Dansk plantevaekst / af Eug. Warming. København og Kristiania: Gyldendal, 1906-1909. DAN url p. 212, p. 217, p. 279, p. 280, p. 295, p. 33, p. 58, p. 61, p. 80, p. 82, p. 86, p. 89.
• Davis, P. H., ed. 1965 –1988. Flora of Turkey and the east Aegean islands. (F Turk)
• De montium inter Vistritium et Nissam fluvios sitorum flora / auctor Richard Sadebeck. Vratislaviae: Typ. officinae A. Neumanni, 1864. LAT url p. 34.
• Denkschriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Classe. Wien: Aus der Kaiserlich-K©niglichen Hof- und Staatsdruckerei, 1850-1918. GER url p. 155, p. 220, p. 221, p. 244.
• Deutsche botanische Monatsschrift. [Sondershausen: Verlag der Redaktion, 1883-1912]. GER url p. 16.
• Dictionary of biological equivalents, German-English, by Ernst Artschwager Baltimore, The Williams & Wilkins company, 1930. ENG url p. 26, p. 75.
• Die Flora des Puschlav (Bezirk Bernina, Kanton Graubünden) von H. Brockmann-Jerosch. Leipzig, W. Engelmann, 1907. GER url p. 225.
• Die Flora von Kärnten. Klagenfurt, Kleinmayr, 1853. GER url p. 129.
• Die Gartenwelt. Berlin: G. Schmidt, [1897- GER url p. 239.
• Die Gross-Schmetterlinge der Erde: eine systematische Bearbeitung der bis jetzt bekannten Gross-Schmetterlinge / von Professor Dr. Adalbert Seitz. Stuttgart: Alfred Kernen, 1909- GER url p. 200.
• Die Lehre von der Pflanzenzelle / von Wilh. Hofmeister. Leipzig: Engelmann, 1867. GER url p. 423, p. 646.
• Die Lepidopteren der Schweiz, von Prof. Dr. Heinrich Frey. Leipzig, W. Engelmann, 1880. GER url p. 296.
• Die Natürlichen Pflanzenfamilien nebst ihren Gattungen und wichtigeren Arten, insbesondere den Nutzpflanzen, unter Mitwirkung zahlreicher hervorragender Fachgelehrten begründet von A. Engler und K. Prantl, fortgesetzt von A. Engler Leipzig, W. Engelmann, 1887-1909. GER url p. 27.
• Die Ostsudeten; eine floristische Skizze. Brünn, Landesdurch-forschungs-Komemission, 1914. GER url p. 119, p. 64, p. 67, p. 79.
• Die Pflanzen und Raupen Deutschlands: Versuch einer lepidopterologischen Botanik / von O. Wilde. Berlin: E.S. Mittler, 1860-1861. GER url p. 179, p. 321.
• Die Pflanzendecke Österreich-Ungarns. Auf Grund fremder und eigener Forschungen geschildert, von Dr. August Edler von Hayek. Hrsg. mit einem Druckkostenbeitrag der Kaiserlichen Akademie der Wissenschaften in Wien. Leipzig, F. Deuticke, 1916- [i.e. 1914] GER url p. 103, p. 124, p. 147, p. 150, p. 259, p. 331, p. 342, p. 374, p. 506.
• Die Pflanzenfeinde aus der Klasse der Insekten. Ein nach Pflanzenfamilien geordnetes Handbuch sämmtlicher auf den einheimischen Pflanzen bisher beobachteten Insekten zum Gebrauch für Entomologen, Insektensammler, Botaniker, Land- und Forstwirthe un von J. Stuttgart, J. Hoffmann, 1874. GER url p. 110, p. 113, p. 14, p. 144, p. 372, p. 373, p. 374, p. 375, p. 376, p. 378, p. 379, p. 381, p. 383, p. 405, p. 407, p. 61, p. 787.
• Die Pilze (Fungi) von Tirol, Vorarlberg und Liechtenstein. Unter Beistand von K.W. v. Dalla Torre und Ludwig Grafen v. Sarnthein bearb. von Paul Magnus. Innsbruck, Verlag der Wagner'schen Universitts-Buchh., 1905. GER url p. 19, p. 562, p. 77.
• Die Pusztenflora der grossen ungarischen Tiefebene / von Franz Woenig; mit einer farbigen Beilage und zahlreichen Pflanzenbildern im Text von Maler Ernst Kiesling; nach des Verfassers Tode herausgegeben von E.S. Zürn. Leipzig: Verlag von Carl Meyers Graphischem Institut, 1899. GER url p. 97.
• Die Schmetterlinge Deutschlands mit besonderer Berücksichtigung ihrer Biologie, von Dr. Karl Eckstein Stuttgart, K.G. Lutz' Verlag[1913]-33. GER url p. 73.
• Die Schmetterlinge Europas. Stuttgart, E. Schweizerbart, 1908-1910. GER url p. 290.
• Die bisher bekannten Pflanzen Galiziens und der Bukowina, von Josef Armin Knapp. Wien, W. Braumüller, 1872. GER url p. 425.
• Diptera danica: genera and species of flies hitherto found in Denmark / by William Lundbeck. Copenhagen: G.E.C. Gad, 1907-27. DAN url p. 269.
• Dr. Johannes Leunis Synopsis der thierkunde. Ein handbuch für höhere lehranstalten und für alle, welche sich wissenschaftlich mit der naturgeschichte der thiere beschäftigen wollen. Hannover, Hahnsche buchhandlung, 1883-86. GER url p. 232.
• Dr. L. Rabenhorst's Kryptogamen-Flora von Deutschland, Oesterreich und der Schweiz Leipzig: E. Kummer, 1884-1966. GER url p. 190, p. 410, p. 421, p. 441, p. 555, p. 674.
• Element Stewardship Abstract for Cirsium arvense (Nuzzo, 1997)
• Encke, F. et al. 1984. Zander: Handwörterbuch der Pflanzennamen, 13. Auflage. (Zander ed13)
• Entomological news. [Philadelphia]American Entomological Society, 1925- ENG url p. 112, p. 115, p. 122, p. 123, p. 16, p. 79.
• Entomologische Nachrichten. Berlin [etc.], R. Friedländer [etc.]. GER url p. 107, p. 194, p. 196, p. 197, p. 223, p. 224, p. 330, p. 332, p. 360, p. 7.
• Enumeratio plantarum Transsilvaniae: exhiben: stirpes phanerogamas sponte crescentes atque frequentius cultas, cryptogamas vasculares, charceas, etiam muscos hepaticasque / auctore Phil. Johanne Ferdinando Schur. Vindobonae: Apud G. Braumhuller, 1866. GER url p. 424.
• Enumeratio plantarum anno 1890 in Caucaso lectarum: additis nonnullis speciebus a claris viris H. Lojka, G. Radde, N. Seidlitz et fratr. Brotherus in eadem ditione lectis / S. Sommier et E. Levier, cum tabulis lithographicis XLIX ab E. Levier et C. Cuisi Petropoli: Typ. Imp. akademii nauk, 1900. FRE url p. 267.
• Essai sur les Myodaires / par le Docteur J. B. Robineau-Desvoidy. Paris: Imprimé par autorisation du Roi a l'Imprimerie Royal, 1830. FRE url p. 771.
• Experiment station record. Washington: G.P.O., 1889-1946. ENG url p. 1179, p. 149, p. 332, p. 335, p. 542, p. 555, p. 70, p. 727, p. 8.
• FNA Editorial Committee. 1993 –. Flora of North America. (F NAmer)
• Fauna Austriaca: die Fliegen (Diptera) Nach der analytischen Methode bearb., mit der Characteristik almmilicher europäischer Gattungen, der Beechraibung aller in Deutschland vorkommenden Arten und der Aufzahlung aller von J. Rudolph Schiner. Wien: C. Gerolds Sohn, 1862-64. GER url p. 132, p. 317.
• Fauna austriaca. Die Käfer. Nach der analytischen Methode bearb. von Ludwig Redtenbacher. Wien, C. Gerold's Sohn, 1874 [i.e. 1871-74] GER url p. 445.
• Femhundra afbildningar af mera allmänt förekommande svenska växter: för skolan och hemmet / med text af N. J. Andersson. Stockholm: Z. Haeggström, 1870. SWE url p. 4.
• Fifty years of botany; golden jubilee volume of the Botanical Society of America, edited by William Campbell Steere. New York, McGraw-Hill, 1958. ENG url p. 406, p. 416.
• Flora Berolinensis, sive, Enumeratio plantarum circa Berolinum sponte crescentium secundum familias naturales disposita / auctore Carolo Sigismundo Kunth. Berolini: Duncker et Humblot, 1838. LAT url p. 367.
• Flora Einsidlensis: systematische Aufzählung der in Einsiedeln freiwachsenden und häufiger cultivirten Gefässpflanzen / von Thomas A. Bruhin. [Einsiedeln; K. und N. Benziger, 1864]. GER url p. 34.
• Flora Italica; sistens plantas in Italia et in insulis circumstantibus sponte nascentes. Bononiae, R. Masi, 1833-1854. LAT url p. 31, p. 32.
• Flora Peoriana; the vegetation in the climate of middle Illinois. Peoria, Ill., J. W. Franks, 1887. ENG url p. 38.
• Flora de las Islas Baleares, seguida de un diccionario de los nombres baleares, castellanos y botánicos de las plantas espontáneas y de las cultivadas. Palma, P. J. Gelabert, 1879-1881. SPA url p. 616.
• Flora de las islas Baleares, seguida de un diccionario de los nombres baleares, castellanos y botánicos, de las plantas espontáneas y de las cultivadas. Palma, Estab. tip. de P.J. Gelabert, 1870-1881. SPA url p. 616.
• Flora del Modenese e del Reggiano / dei Professori G. Gibelli e R. Pirotta. Modena: Tip. di G.T. Vincenzi, 1882. ITA url p. 95.
• Flora der Nordfriesischen Inseln / von Paul Knuth. Kiel; Lipsius & Tischer, 1895. GER url p. 80.
• Flora der Schweiz: zum Gebrauche auf Exkursionen, in Schulen und beim Selbstunterricht / von Hans Schinz und Robert Keller. Zürich: A. Raustein, 1909-1914. GER url p. 224.
• Flora der Umgebung Lemsals und Laudohns; zwei Beiträge zur Flora Livlands, von A. Rapp. Hrsg. und mit einer phytogeographischen Einleitung versehen von J. Klinge. Riga, W.F. Häcker, 1895. GER url p. 12, p. 35, p. 69, p. 89.
• Flora der gefürsteten Grafshaft Tirol, des Landes Vorarlberg und des Fürstenthumes Liechtenstein. Nach eigenen und fremden Beobachtungen, Sammlungen und den Litteraturquellen bearb. von K.W. v. Dalla Torre und Ludwig Grafen von Sarnthein. Innsbruck, Verlag der Wagner'-schen Universitäts-Buchhandlung, 1900-1913. GER url p. 19, p. 22, p. 77.
• Flora der ostfriesischen Inseln. Norden und Norderney, H. Braams, 1881. GER url p. 87.
• Flora melitensis nova [by] S. Sommier et A. Caruana Gatto. Firenze, Stabilimento Pellas, 1915. ITA url p. 189, p. 455.
• Flora of Ann Arbor and vicinity. Ann Arbor, [Mich.]: Courier Steam Print. House, 1876. ENG url p. 116, p. 99.
• Flora of Berkshire County, Massachusetts. Boston, 1922. ENG url p. 363.
• Flora of Illinois, containing keys for identification of flowering plants and ferns. Notre Dame, Ind., University of Notre Dame Press, 1963. ENG url p. 268.
• Flora of Indiana, by Charles C. Deam. Indianapolis, Wm. B. Burford printing co., contractor for state printing and binding, 1940. ENG url p. 1001, p. 1002.
• Flora of Lancaster County; being descriptions of the seed- plants growing naturally in Lancaster County, Pennsylvanial. By John Kunkel Small and Joel Jackson Carter. New York, 1913. ENG url p. 311.
• Flora of Middlesex County, Massachusetts, by L. L. Dame and F. S. Collins. Malden, Middlesex Institute, 1888. ENG url p. 57.
• Flora of New Bedford and the shores of Buzzards Bay, with a procession of the flowers. New Bedford [Mass.]E. Anthony, 1911. ENG url p. 55.
• Flora of Vermont. List of ferns and seed plants growing without cultivation. Prepared by Vermont Botanical Club. Burlington, Vt., Free Press Print. Co., 1915. ENG url p. 253.
• Flora of southeastern Washington and adjacent Idaho, by Charles V. Piper and R. Kent Beattie Lancaster, Pa., Press of the New era printing company, 1914. ENG url p. 259, p. 260.
• Flora of the Indiana dunes, a handbook of the flowering plants and ferns of the lake Michigan Coast of Indiana and of the Calumet District, by Donald Culross Peattie. Chicago, Field Museum of Natural History, 1930. ENG url p. 397.
• Flora of the Miami Valley, Ohio /c by A.P. Morgan. Dayton, Ohio: Literary Union, 1878. ENG url p. 62.
• Flora of the northwest coast, including the area west of the summit of the Cascade Mountains, from the forty-ninth parallel south to the Calapooia Mountains on the south border of Lane County, Oregon. By Charles V. Piper and R. Kent Beattie. Lancaster, Pa., Press of the New Era Printing Company, 1915. ENG url p. 394, p. 395.
• Flora of the state of Washington. By Charles V. Piper. Washington, Govt. Print. Off., 1906. ENG url p. 609.
• Flora padovana; ossia, Prospetto floristico e fitogeografico delle piante vascolari indigene inselvatichite o largamente coltivate crescenti nella provincia di Padova; con notizie storico-bibliografiche sulle fonti d Padova, Prem. Soc. coop. tip., 1909-1914. ITA url p. 646, p. 675, p. 694.
• Flora von Bern; systematische Uebersicht der in der Gegend von Bern wildwachsenden und allgemein cultivirten Phanerogamen und Gefässkryptogamen, von L. Fischer. Bern, Huber, 1888. GER url p. 287.
• Flora von Halle, mit nherer Bercksichtigung der Umgegend von Weissenfels, Naumburg, Freiburg, Bibra, Nebra, Querfurt, Allstedt, Artern, Eisleben, Hettstedt, Sandersleben, Aschersleben, Stassfurt, Bernburg, Kthen, Dess Halle, Anton, 1848-1856. GER url p. 259.
• Flora von Nieder-Österreich. Handbuch zur Bestimmung sämmtlicher in diesem Kronlande und den angrenzenden Gebieten wildwachsenden, häufig gebauten und verwildert vorkommenden Samenpflanzen und Führer zu weiteren botanischen Forsc Wein, C. Gerold, 1890-93. GER url p. 1237, p. 1250, p. 45, p. 53, p. 54, p. 61, p. 66.
• Flora von Ober-Oesterreich, oder, Systematische Uebersicht aller in diesem Kronlande wildwachsenden oder im Freien gebauten Samenpflanzen. [Wien, 1862] GER url p. 1036, p. 1068.
• Flora von Schaffhausen / bearbeitet von Jakob Meister. Schaffhausen: Buchdr. von H. Meier, 1887. GER url p. 110.
• Flora von Schlesien, preussischen und österreichischen Antheils, oder, Vom oberen Oder- und Weichsel-Quellen-Gebiet. Nach natürlichen Familien, mit Hinweisung auf das Linnéische System. Breslau, F. Hirt, 1857. GER url p. 281, p. 413.
• Flora von Tirol. Ein Verzeichniss der in Tirol und Vorarlberg wild wachsenden und häesiger gebauten Gefässpflanzen. Mit Berücksichtigung ihrer Verbreitung und örtlichen Verhältnisse verfasst und nach Koch's Synopsis d von Frz. Freih. v. Hausmann. Innsbruck, Wagner, 1851. GER url p. 477.
• Flora von Wien; eine Aufzählung der in den Umgebungen Wiens wild wachsenden oder im grossen gebauten Gefässpflanzen, nebst einer pflanzen-geografischen Uebersicht. Wien, F. Beck, 1868. GER url p. 55.
• Flore de la Côte-d'Or: avec déterminations par les parties souterraines / par Ch. Royer. Paris: F. Savy, 1881-1883. FRE url p. 348, p. 396, p. 397.
• Flore de la Suisse / par Hans Schinz et Robert Keller. Lausanne: F. Rouge, 1909. FRE url p. 615.
• Floristicka istrazivanja po jugoistocnoj hrvatskoj. U Zagrebu, Tisak Kr. Zemaljske Tiskare, 1915. SCR url p. 39.
• Fr. Berge's Schmetterlingsbuch nach dem gegenwärtigen Stande der Lepidopterologie neu bearb. und hrsg. von Professor Dr. H. Rebel Stuttgart, E. Schweizerbart, 1910. GER url p. 443.
• Führer zu den wissenschaftlichen Exkursionen des II. Internationalen Botanischen Kongresses, Wien, 1905. Wien: Selbstverlage des Organisationskomitees, 1905. GER url p. 153, p. 49, p. 7.
• Gartenflora. Erlangen: F. Enke, 1852-1940. GER url p. 232.
• Great Basin naturalist memoirs. [Provo, Utah]Brigham Young University, 1976-1992. ENG url p. 171, p. 79.
• Growth of plants; twenty years' research at Boyce Thompson Institute. New York, Reinhold Pub. Corp., 1948. ENG url p. 40.
• Haandbog i den danske flora / af Joh. Lange. Kjöbenhaven: C.A. Reitzel, 1851. DAN url p. 470.
• Handbook of British fungi, with full descriptions of all the species, and illustrations of the genera. By M. C. Cooke, M.A. London, Macmillan and co., 1871. ENG url p. 452.
• Handbuch der systematischen botanik, mit besonderer berücksichtigung der arzneipflanzen. Bd. 1. Leipzig, H. Haessel, 1879. GER url p. 137, p. 190, p. 242.
• Handwörterbuch der naturwissenschaften / hrsg. von prof. dr. E. Korschelt (zoologie) prof. dr. G. Linck (mineralogie und geologie) prof. dr. F. Oltmanns (botanik) prof. dr. K. Schaum (chemie) prof. dr. H. Th. Simon (physi Jena: G. Fischer, 1912-1915. GER url p. 977.
• Hedwigia. 18 1879 Dresden: Verlag und Druck von C. Heinrich. GER url p. 135.
• Histoire naturelle des insectes. Genera des coléoptères, ou exposé méthodique et critique de tous les genres proposés jusquici dans cet orde dinsectes, par m. Th. Lacordaire. Paris: Librairie encyclopédique de Roret, 1854-76. FRE url p. 75.
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• Petrak, F. 1917. Die nordamerikanischen Arten der Gattung Cirsium. Beih. Bot. Centralbl. 35(2): 223-567.

Notes

Contributors

• Bisby, F.A., Y.R. Roskov, M.A. Ruggiero, T.M. Orrell, L.E. Paglinawan, P.W. Brewer, N. Bailly, J. van Hertum, eds (2007). Species 2000 and ITIS Catalogue of Life: 2007 Annual Checklist. Species 2000: Reading, U.K.
• "Cirsium arvense". in Flora of North America Vol. 19, 20 and 21 Page 95, 96, 97, 102, 109, 110. Published by Oxford University Press. Online at EFloras.org.
• Global Biodiversity Information Facility. Accessed March 29, 2007. http://www.gbif.org Mediated distribution data from 2 providers.
• Light, Kris. East Tennessee Wildflowers
• National Invasive Species Information Center, National Agricultural Library, United States Department of Agriculture. Web Site. Accessed May 3, 2008.
• USDA, ARS, National Genetic Resources Program. Germplasm Resources Information Network - (GRIN) [Online Database]. National Germplasm Resources Laboratory, Beltsville, Maryland. URL (April 24, 2008)

Data Sources

Accessed through GBIF Data Portal November 18, 2007:

• Jyväskylä University Museum - The Section of Natural Sciences, Vascular plant collection of Jyvaskyla University Museum
• Marine Science Institute, UCSB, Paleobiology Database
• Missouri Botanical Garden, Missouri Botanical Garden
• Oregon State University, Vascular Plant Collection
• The Danish Biodiversity Information Facility, Botany registration database by Danish botanists
• The Swedish Museum of Natural History
• , Herbarium of Oskarshamn
• The Swedish Museum of Natural History
• , Lund Botanical Museum
• The Swedish Museum of Natural History
• , Plants
• UK National Biodiversity Network, Botanical Society of the British Isles - Vascular Plants Database
• UK National Biodiversity Network, Environment and Heritage Service - EHS Species Datasets
• UK National Biodiversity Network, Joint Nature Conservation Committee - Vegetation surveys of coastal shingle in Great Britain
• University of Alaska Museum of the North, University of Alaska Museum of the North Herbarium
• University of Washington Burke Museum, Vascular Plant Collection - University of Washington Herbarium
• Utah Valley State College
• , Utah Valley State College Herbarium

Identifiers

• Biodiversity Heritage Library NamebankID: 2657795
• Catalogue of Life Accepted Name Code: ITS-36335
• Global Biodiversity Information Facility Taxonkey: 13368305
• Globally Unique Identifier: urn:lsid:ipni.org:names:78011-3
• GRIN Nomen Number: 100755
• Integrated Taxonomic Information System (ITIS) Taxonomic Serial Number (TSN): 36335
• Natural Heritage Network Species Identifier: PDAST2E090
• U.S.D.A. Plant Symbol: SEAR12
• Zipcode Zoo Species Identifier: 12785

Footnotes

1. Theodore M. Barkley, Luc Brouillet, John L. Strother "Asteraceae". in Flora of North America Vol. 19, 20 and 21 Page 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 16, 70. Oxford University Press. Online at EFloras.org. [back]
2. David J. Keil "Cirsium". in Flora of North America Vol. 19, 20 and 21 Page 57, 66, 82, 83, 93, 95, 96, 97, 100, 102, 1. Oxford University Press. Online at EFloras.org. [back]
3. Mean = 91.180 meters (299.147 feet), Standard Deviation = 168.400 based on 20,000 observations. Altitude information for each observation from British Oceanographic Data Centre. [back]

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