Flora of the Canadian Arctic Archipelago
English: Pasque-flower, prairie smoke, prairie-crocus,
Ranunculaceae, Buttercup family.
Published in Linnaea 15: 58. 1841.
Type: Described from Siberia.
Synonymy. Pulsatilla patens (L.) Mill. subsp. multifida (Pritz.) Zämelis, Acta Hort. Bot. Univ. Latv. 1: 98. 1926.
Pulsatilla nuttalliana (DC.) Bercht. and J. Presl subsp. multifida (Pritz.) Aichele and Schwegler Feddes Repert. 60, 1–3: 81. 1957.
Pulsatilla patens (L.) Mill. subsp. hirsutissima Zämelis, Acta Hort. Bot. Univ. Latv. 1: 98 1926.
Pulsatilla hirsutissima (Pursh) Britton, nom. illegit., Ann. New York Acad. Sci. 6: 217. 1891.
Pulsatilla ludoviciana (Nutt.) A.Heller, nom. illegit., Cat. Enum. Amer. Pl., ed. 2, 4. 1900.
Vegetative morphology. Plants 10–15(–25) cm high (to 40–60 cm tall on continental North America); perennial herbs; not caespitose. Only fibrous roots present (fibrous roots from a branching caudex present). Ground level or underground stems vertical (stout); caudex; elongate, or compact; 5–10 mm wide. Ground level or underground stems scales present (and densely villous, aboveground at the base of the plant); 2–5. Caudex present. Aerial stems erect (or spreading). Aerial stem trichomes spreading, or erect. Leaves mainly basal, or basal in a rosette (of (30)5–8(-10) leaves); erect; alternate; dying annually and non-persistent (leaf blades), or marcescent (leaf petioles that build-up around the caudex). Stipules absent (expanded petiole bases and scales at the base of the plant may appear stipule-like, but they are separate organs). Petioles present (basal leaves), or absent (flowering stem leaves); (0–)45–50 mm long (Arctic island specimens, to 130 mm long in continental North America specimens); winged (expanded, at the base); hairy; villous. Petiole hairs longer than the diameter of the petiole (long and silky); spreading, or erect; straight, or curved. Leaf blades compound. Blades (15–)20–40 mm long (Arctic island specimens, to 60–60 cm long in specimens from continental North America), 15–20(–55) mm wide, spreading, straight, veins palmate. Blade adaxial surface dull, without sessile glands, hairy, hairs villous, hairs simple or stellate, hairs sparse, hairs white, or translucent. Blade abaxial surface dull, without sessile glands or glandular hairs, hairy or glabrous (rarely), hairs villous, hairs moderately dense, hairs white, hairs straight (sometimes twisted together into tufts), hairs spreading or erect. Blade margins deeply divided, glabrous (or with a few sparse hairs from the blade surface); degree of incision 65–75%; apices acuminate. Leaflet arrangement palmate (each leaflet dichotomously dissected). Leaflets 3; 10–20(–60) mm long; 15–20 mm wide (for the three leaflets, individual divisions 1–4 mm wide); linear (blade bipinnate, the leaflets pinnately divided, margins dichotomously dissected throughout into long, linear segments; the stem-leaves deeply cleft into linear lobes). Apical leaflet base not distinctly stipitate (terminal leaflet, obovate in outline, (2.5-)3–5 mm long, base narrowly cuneate).
Reproductive morphology. Flowering stems conspicuously taller than the leaves; with leaves. Flowering stems hairy. Flowering stems villous. Flowering stem hairs simple; shorter than the diameter of the flowering stem (for most of the length), or longer than the diameter of the flowering stem (below the flower); white or translucent. Flowers solitary. Involucral bracts present (three, one-tiered). Flowers large. Sepals conventional. Epicalyx absent. Sepals 5–7(–8); free; 8–12(–15) mm long; (10–)20–45 mm wide; purple (usually), or red, or blue, or yellow (after pressing); petaloid; non-accrescent. Calyx hairy (abaxially). Calyx hairs villous; non-glandular; white or translucent. Petals absent (but the sepals are coloured and resemble petals). Stamens (50–)60–85 (150–200 in the south); stamen filaments glabrous. Anthers yellow; long-cylindrical; (0.8–)1.1–1.3(–1.8) mm long. Nectaries present (at the base of the tepals). Receptacle 2–3(–6) mm high. Ovary carpels 30–45; apocarpous. Stipes 0.05–0.1 mm long. Ovaries hairy; villous. Ovary hairs very dense; white; straight. Styles 2–3 mm long (when receptive; becoming 23–28 mm long, in fruit). Stigmas per ovary 1. Ovules per ovary 1. Fruit stalk 9–20 mm long; dry; an aggregate of achenes; ellipsoid, or obovate; brown; 30–45 mm long; 40–70 mm wide (achenes (2.5-)4–5 × 0.8–1.2(-1.5) body ellipsoid to obovoid, not winged villous, beak curved 20–40 mm long-villous, plumose); hairy; not distinctly flattened; indehiscent. Styles modified and persisting (plumose); remaining straight; persisting in fruit 23–38 mm long. Seeds 1.
Chromosome information. 2n = 16 and 32.
2n (2x) = 16. Bormann and Beatty (1955, Alaska), Heimburger (1959, Alaska; 1962); Rothfels et al. (1966); Taylor and Brockman (1966, western Canada); Zhukova (1966, 1969, northeastern Asia); Löve et al. (1971, western North America); Zhukova and Tikhonova (1971, Chukotka); Kartashova and Malakhova (1972, Siberia); Zhukova and Petrovsky (1972, northeastern Asia); Belaeva and Siplivinsky (1975, 1976, southern and northern Siberia); Dawe and Murray, in Löve (1979, Alaska); Zhukova and Petrovsky (1980, 1987b, northeastern Asia); Zhukova (1980, southern Chukotka); Rostovtseva (1981) Krogulevich and Rostovtseva (1984, Siberia); Yurtsev and Zhukova (1982, northern Siberia); Löve and Löve (1982, central Canada); Volkova and Ulanova (1986, northeastern Asia).
2n (4x) = 32. Löve (1954a, 1954b); Löve and Solbrig (1964a, Canada).
Ploidy levels recorded 2x/4x.
Indigenous knowledge. Native Americans, to the south, used fresh leaves to treat rheumatism and neuralgia; crushed leaves for poultices; pulverised leaves to smell to ease headaches; and made decoctions of roots for lung problems (Moerman 1986).
Ecology and habitat. Substrates: gravel, sand; with low organic content. Sandy well-drained soil.
North American distribution. Alaska, Yukon, Northwest Territories Islands, continental Northwest Territories. Range in the Canadian Arctic Archipelago limited. Rare. Alpine, boreal. Arctic islands: Banks, Victoria.
Northern hemisphere distribution. North American, or Cordilleran (western), or Siberian. YamalGydan, Taimyr Severnaya Zemlya, YanaKolyma, West Chukotka, Wrangel Island, South Chukotka, North Alaska Yukon, Central Canada.
General notes. Dutton et al. (1997), in the Flora of North America, treated this taxon as Anemone patens var. multifida. They were aware that this taxon is often placed in the genus Pulsatilla Miller on the basis of the long plumose achene beaks but claimed that recent phylogentic analyses of Anemone in the broad sense indicated that it should be subsumed within Anemone (Hoot et al. 1994). They commented that Anemone patens var. multifida has frequently been recognised as a subspecies of A. patens. A close relative of this taxon Anemone patens var. wolfgangiana is the floral emblem of Manitoba and the State Flower of North Dakota.
Elven et al. (2003) treated this taxon as Pulsatilla patens (L.) Mill. subsp. multifida (Pritz.) Zämelis.
Hultén (1944, p. 739) commented, "I can see no essential difference between the Siberian plant and the American and must consequently refer both to the same subspecies [i.e., subsp. multifida]. I find it impossible to regard the different races of Anemone patens as distinct species as Juzupczuk does in Fl. S.S.S.R. Where they meet, intermediate forms occur and they are clearly geographic races". This concept has been followed in Elven et al. (2003).
Petrovsky, in Tolmatchew (1971a), discussed the relations among the Asian and North American plants but not their relation to the European - western Siberian P. patens. Other authors, like Zämelis (1926), Hultén and Fries (1986), and Lindell, in Jonsell (2001a), have considered these entities as widely distributed geographical races (i.e., subspecies) of one wide species for which P. patens is the oldest name. That approach was followed by Elven et al. (2003).
This taxon and its related varieties show considerable variation, which is considered to be environmentally induced.
Two varieties were recognised by Dutton et al. (1997) in northwestern North America, both within the Arctic. Variety multifida was mapped for arctic parts of northeastern Alaska, northern Yukon, the Hudson Bay area in Manitoba, Ontario, and Quebec, and northernmost Ungava Peninsula. Variety saxicola occurs in the western Cordilleran and reaches the arctic in northern Alaska. Several assumed independently inherited differential characters are given. However, in general, var. saxicola occurs almost entirely within the range of var. multifida. Elven et al. (2003) compared Alaskan material annotated, as the two varieties found no very clear differences, and proposed that the varieties be treated as synonyms.
Illustrations. • Flower. Herbarium specimen showing side view of immature flower. Several divided involucral bracts in a single tier are covered in long hairs. Yukon, Snag Airport, Deadmann's Hill, 62°24'N, 140°22'W. 20 May, 1945. G.A. Noel. CAN 58050. • Close-up of flower. Herbarium specimen of flower that has faded from blue to yellow in pressing. Centre of the flower shows a ring of numerous anthers with yellow tips surrounding carpel. Carpels have dark tips and are covered with white silky hairs. Note the involucral bracts cut into linear divisions. Yukon, Lake Laberge. 10 June, 1919. W.E. Cockfield. CAN 58035. • Close-up of fruit. Herbarium specimen of pressed fruiting head. Achenes are numerous and have long modified styles. Yukon, Dawson. 2 June, 1914. Alice Eastwood 125. CAN 58045. • Arctic Island Distribution.
This publication is available on the internet (posted May 2011) and on CD-ROM (published in 2007). These versions are identical in content, except that the errata page for CD-ROM is accessible on the main index page of the web version.
Recommended citation for the web-based version of this publication: Aiken, S.G., Dallwitz, M.J., Consaul, L.L., McJannet, C.L., Boles, R.L., Argus, G.W., Gillett, J.M., Scott, P.J., Elven, R., LeBlanc, M.C., Gillespie, L.J., Brysting, A.K., Solstad, H., and Harris, J.G. 2007. Flora of the Canadian Arctic Archipelago: Descriptions, Illustrations, Identification, and Information Retrieval. NRC Research Press, National Research Council of Canada, Ottawa. http://nature.ca/aaflora/data, accessed on DATE.
Recommended citation for the CD-ROM version of this publication: Aiken, S.G., Dallwitz, M.J., Consaul, L.L., McJannet, C.L., Boles, R.L., Argus, G.W., Gillett, J.M., Scott, P.J., Elven, R., LeBlanc, M.C., Gillespie, L.J., Brysting, A.K., Solstad, H., and Harris, J.G. 2007. Flora of the Canadian Arctic Archipelago: Descriptions, Illustrations, Identification, and Information Retrieval. [CD-ROM] NRC Research Press, National Research Council of Canada, Ottawa..