Flora of the Canadian Arctic Archipelago
English: Arctic fireweed, broad-leaved willow herb, dwarf fireweed, river beauty,
French: Épilobe à feuilles larges,
Inuktitut: Paunnat; paunnait (Baffin Island); boundnak or paunnaq (northern Quebec).
Onagraceae, Fireweed family.
Published in Hort. Brit., ed. 2, 198. 1830.
Type: Described from Siberia.
Synonymy. Epilobium latifolium L., Sp. Pl. 347. 1753.
Chamaenerion latifolium (L.) Th. Fr. and Lange, Fl. Dan. 49: t. 2901. 1877.
Chamaenerion subdentatum Rydberg, Fl. Rocky Mountains. 585. 1917.
Chamerion subdentatum (Rydberg) Löve and Löve, Bot. Not. 128: 516. 1976.
Vegetative morphology. Plants 5–30 cm high; perennial herbs. Only fibrous roots present. Ground level or underground stems horizontal; rhizomatous; elongate (in old plants), or compact; 2–9 mm wide. Aerial stems developed; ascending, or decumbent. Aerial stem trichomes retrorse. Leaves present; distributed along the stems; alternate, or opposite; dying annually and non-persistent (killed at the end of each growing season, arctic island plants). Petioles absent. Leaf blade bases obtuse, or cuneate. Blades (10–)25–45(–65) mm long, (3–)10–15(–22) mm wide, spreading, lanceolate (broadly or narrowly), flat, veins pinnate or appearing single-veined. Blade adaxial surface glabrous or glabrescent, hairs simple. Blade abaxial surface glabrous (to the naked eye) or glabrescent or hairy, hairs puberulent (if applicable), hairs sparse or moderately dense or very dense (hairs glabrescent), hairs white, hairs curved. Blade margins entire, glabrous; apices acuminate, or acute.
Reproductive morphology. Flowering stems two or more per plant; with leaves. Flowering stems hairy. Flowering stems puberulent. Flowering stem hairs simple; shorter than the diameter of the flowering stem; white or translucent. Inflorescences racemose (leafy); diffuse; elongating as the fruit matures. Pedicels present; with non-glandular hairs. Flowers per inflorescence (1–)3–5(–8); large (largest flowers in the Arctic islands). Sepals conventional; 4; free; (1.5–)2.5–3.5(–5.5) mm long; (11–)14–18 mm wide; purple (dark, intense colour). Calyx hairy (in bud), or glabrous (with age). Calyx hairs puberulent; non-glandular; white or translucent. Petals conventional; free; 4; white (rarely), or pink, or purple (pale); obovate and spatulate (ulu-shaped); unlobed; 15–25 mm long; 0–20(–22) mm wide (4 mm wide at the base). Stamens 8; stamen filaments glabrous. Anthers purple; 1.4–2 mm long. Nectaries present (near the base of the anthers). Ovary inferior; carpels 4; syncarpous. Ovaries hairy; puberulent, or pubescent. Ovary hairs very dense; white. Styles 1; 4–7(–8) mm long (elongating after anthesis); straight; basal portion smooth. Stigmas per ovary 4. Stigma lobes 2–3 mm long. Placentation axile. Ovules per ovary very numerous. Fruit stalked; stalk 5–25 mm long; without calyx persisting; dry; a capsule; elongate-cylindrical; purple, or brown (grey); 45–70 mm long; 2–3.5 mm wide; hairy, or glabrescent (puberulent when young; losing hairs with age); surface appearing veinless; not distinctly flattened; dehiscent; splitting to the base into separate segments; teeth 4. Seeds numerous; 1.8–2 mm long (with a tuft of yellowish hairs to 10 mm long); brown, or yellowish; surfaces smooth.
Chromosome information. 2n = 36, or 54, or 72.
2n (4x) = 36. Small (1968, Alaska, western USA, western Canada, 14 counts); Johnson and Packer (1968, northwestern Alaska); Sokolovskaya (1968, northeastern Asia); Krogulevich (1971, Siberia); Zhukova and Tikhonova (1973, Chukotka); Belaeva and Siplivinsky (1976a, southern and northern Siberia);
2n (6x) = 54. Mosquin and Hayley (1966, northern Canada); Small (1968, northern and western Canada, two counts, one arctic);
2n (8x) = 72. Johansen (1929a, 1929b); Böcher (1938a, Greenland, 2n = about 72); Harmsen, in Löve and Löve (1948, northern Europe); Löve and Löve (1948, northern Europe; 1956, Iceland; 1982a, Arctic Canada); Böcher and Larsen (1950, Greenland); Harmsen, in Jørgensen et al. (1958, Greenland); Böcher (1962); Small (1968, Iceland, Canada, seven counts); Sokolovskaya (1970, northeastern Russia); Zhukova and Petrovsky (1971, northeastern Asia); Krogulevich (1976a, northern Siberia; 1978, southern and northern Siberia); Dawe and Murray, in Löve (1979, central Alaska; 1981).
Ploidy levels recorded 4x, 6x, and 8x.
Indigenous knowledge. Though there is no special season to pick this plant, year-old plants are best (Anon 1984). All parts of the plant (leaves, flowers and seed pods) are edible. They can be eaten raw or mixed with fat, or blood and oil. The leaves are good cooked and taste somewhat like spinach. The flowers of the willow-herb may be eaten raw as a salad, and the fleshy leaves are edible when cooked. The flowers are delicious when mixed with crowberries, blood, and oil. The long seed pod is also edible before it becomes woody.
These plants can also be used to make tea. The tea is good for stomach aches and to help make a person stronger after they had bled a lot. As well the leaves can be chewed to help stop a nose bleed (George Kappinanq, personal communication, reported in Mallory and Aiken 2004). The whole plant, except the roots, was boiled or chewed to relieve general stomach ache given to people who had intestinal problems to the point where they had blood in their stools if they ate amniq (lamp moss). Lamp moss was used to absorb excess acid in the stomach, causing heartburn, but could result in other complications (Ootoova et al. 2001).
Other uses include fuel for fires and insulation for qammait (Ootoova et al. 2001).
In Greenland, the dwarf fireweed is called niviaqsiaq, "young girl", and it is the national flower. Also in Greenland, the fresh leaves and the flowers are occasionally eaten raw with seal blubber (Porsild 1953).
Ecology and habitat. Substrates: snow patches (runoff areas), along streams, river terraces, ridges, cliffs, seashores, flood plains (river bars, screes, along brooks); imperfectly drained moist areas, seepage slopes, solifluction slopes, dry, moderately well-drained areas; gravel, sand, silt, clay, moss (rarely); with low organic content (usually). Common or even abundant throughout the Arctic in sandy and gravelly, well-watered soils, such as gravel bars or flood plains (Porsild 1957). Sandy fluvial material (CAN 312086); gravel beach (CAN 342823); well-drained shallow braided sandy stream terraces; dry clay ridge (CAN 270534); loose rock slope (CAN 393566); wet gravelly flood plain; snow patch runoff areas (CAN 582436); pioneer species on moderately to well-drained sand bar (CAN 502468); alkaline silty diamicton (CAN 502562).
Mosquin (1966) speculated that, during the last ice age, the diploid populations probably survived all along the southern margins of the ice, and it was these eastern diploids which recolonised much of northeastern Canada. The large unglaciated regions of Alaska were probably the main areas from which recolonisation occurred across the Canadian Arctic. The species has aggressive colonising properties and freely invades man-made clearings, roadsides, logged and burnt-over lands, where it may become extremely abundant for many years. It thrives best under conditions of highly reduced competition.
North American distribution. Alaska, Yukon, Northwest Territories Islands, continental Northwest Territories, Nunavut Islands, continental Nunavut, northern Quebec, Labrador. Range in the Canadian Arctic Archipelago widespread. Common. Arctic, alpine. Arctic islands: Baffin, Devon, Ellesmere, Axel Heiberg, Parry islands (Eglington, Emerald, Melville), Banks, Victoria, Somerset, King William, Southampton, Coats (Big, Digges and Mill islands, Boothia and Melville peninsulas).
Northern hemisphere distribution. Circumpolar, or circumboreal (with a gap in northern Europe). Northern Iceland, Polar Ural Novaya Zemlya, Taimyr Severnaya Zemlya, AnabarOlenyok, Kharaulakh, West Chukotka, Wrangel Island, South Chukotka, East Chukotka, West Alaska, North Alaska Yukon, Central Canada, Labrador Hudson Bay, Ellesmere Land Peary Land, West Greenland, East Greenland.
General notes. Numerous infraspecific taxa of C. latifolium have been described from local regions (Haussknecht 1884, Böcher 1962, Munz 1965). Small (1968) reported that tetraploids of C. latifolium tend to have high percentages of 4-pored pollen, while the diploids usually have only 3-pored pollen. Small (1968) mapped the distribution of the races in the western hemisphere, indicating that all the collections from the Canadian Arctic Archipelago appear to be diploid. The races appear to form worldwide, allopatric ecogeographic phases. A high level of quadrivalent formation observed during tetraploid meiosis indicated that the tetraploids arose by autopolyploidy. Small (1968) stated that the tetraploids in C. latifolium are undoubtedly autotetraploids, as the polyploids and diploids are almost indistinguishable morphologically. The distinctiveness of C. latifolium within its genus make it highly unlikely that the tetraploids arose by allopolyploidy. Since distinctive phases are not found among the diploid populations, the origin of the tetraploid from hybridisation between different diploid races of C. latifolium is very unlikely. This led Small (1968) to recommend that the races not be given formal taxonomic recognition.
Mosquin and Small (1971, p. 680, 681) considered that C. angustifolium and C. latifolium represent an example of parallel evolution and that "although the two species are closely related, evidence indicates that they are isolated by ecological and genetic barriers. [They admitted that] F1 hybrids have been collected from transitional habitats (Böcher 1962) [but noted that hybrids are] extremely infrequent in view of the occurrence of widespread areas of sympatry... Attempts to cross the two species have given variable results (Pugsley 1960, Mosquin, unpubl.) but generally revealed strong barriers to F1 seed production coupled with marked F1 pollen sterility. While these barriers were not considered sufficiently well developed to have prevented introgression between the two species, [Mosquin and Small (1971) stated that] there iss no indication of this in the polyploids, which closely resemble their diploid progenitors, and do not possess characters attributable to introgression from their sister species.
Evidence for parallelism comes from five observations:
(1) Both species occupy comparably widespread ranges in the northern hemisphere...
(2) In each species diploids occupy the more northern and/or alpine areas, while polyploids occur in more southern and/or warmer regions.
(3) In each species autopolyploidy has been the mechanism of polyploidisation.
(4) Each species has retained the capacity to form high levels of quadrivalents at meiosis, unlike many other autopolyploids (Morrison and Rajhathy 1960).
(5) Within each species the diploids very closely resemble the polyploids."
Mosquin and Small (1971) suggested "the evolution of polyploidy has been accompanied by a common morphological change, namely an increase in frequency of extra pores in the pollen grains." (p. 682).
Doak (1991) conducted a 3-year experimental study of dwarf fireweed's responses to real and simulated attacks by a momphid caterpillar and found that the type and extent of plant responses changed with morphological and temporal scale. Damaged shoots showed effective compensation, producing between 89 and 583% more branch tissue than undamaged shoots, but as clumps of shoots are joined by perennial rhizomes; a single year of herbivore damage to a clump's shoots reduced net growth by more than 75% over a 3-year period. Clump form also changed following attack. In the year after damage, shoot numbers were reduced by 51%, while mean shoot size either increased or was unaffected. Further complicating the caterpillar's impact on its host was its inhibition of mammalian herbivory; simulated caterpillar damage reduced the probability of mammalian attack from 20 to 6%. Doak (1991) concluded that these complicated and very different changes following herbivory illustrate the need to study compensation mechanistically and in the context of plant's biotic environment in order to understand its importance and consequences.
Illustrations. • Habitat: Cape Dorset. Plants growing with Dryas in dense Low Arctic tundra. Nunavut, Baffin Island, Cape Dorset. 2 August, 2005. Aiken. No voucher. • Habitat. Plants line the pebble-boulder edge of a runoff stream. Nunavut, Ellesmere Island, MacDonald River Delta, north of Tanquary Camp (in the distance). 21 July, 1999. L.J. Gillespie 6472 and L.L. Consaul. Photograph by L.J. Gillespie. • Close-up of plant. Plants growing in pebble-sand bank with Poa hartzii. Nunavut, Ellesmere Island, MacDonald River Delta, north of Tanquary Camp. 21 July, 1999. L.J. Gillespie 6472 and L.L. Consaul. Photograph by L.J. Gillespie. • Plant habitat. Early colonising plants, less than 20 cm high, invading the disturbed habitat of a gravel flood plain. Nunavut, Ellesmere Island, Vendom Fiord. Aiken 93–015. CAN. • Habitat in rocks. Plants colonising cracks among rocks. Opingivik Island, 60° 56'N; 70°W. 3 August, 2006. Aiken. No voucher. • Close-up of plant. New season's growth with very reddish shoots. Previous season's leaves are gray "ghosts" on the ground around the new plant. Banks Island, Aulavik National Park, near Green Cabin. 29 June, 1999. Aiken 99–004. CAN. • Close-up of plant. Left, flower with 4 free lanceolate sepals, 4 narrowly obovate, free petals, 4 pre-anthesis swollen red anthers, and 4 anthers that have shed pollen. Right, flower with anthers that have shed pollen. Nunavut, Ellesmere Island, Vendom Fiord. Aiken 93–015. CAN. • Plants with white or pink petals. Adjacent plants having flowers with the more usual magenta purple petals and others with white petals. Nunavik, Kangiqsujuaq. 22 July, 2006. Aiken. No voucher. • Close-up of rhizome. Stout rhizome of dwarf fireweed. Yukon, Little Atlin Lake. 15 August, 1943. Raup and Correll 11306. CAN 281069. • Underside of flower. a) inferior ovary; b) flower pedicel; 4 narrow dark purplish red, free sepals, and 4 free pale purple petals attached at the top of the receptacle. Aiken and LeBlanc 2004. No voucher. • Surface view of flower. Deep reddish purple sepals, obovate petals, 8 anthers with filaments around the dark green nectary on the top of the gynoecium, and filaments larger at the base and a branching receptive stigma. Aiken and LeBlanc 2004. No voucher. • Side view of petal attachment. Deep reddish purple sepals, the base of an obovate petal, anther filament larger at the base and a trumpet shaped style dividing into 4 receptive stigmas. Aiken and LeBlanc 2004. No voucher. • Close-up of stigma. Close-up of stigma showing velcro-like surface that accepts the polllen. 24 July, 2005. Photograph by Kathy Thornhill. • Habitat with fruiting capsules. Fruiting capsules that have split into four units and are dispersing the numerous plumose seeds. Nunavut, Baffin Island, Iqaluit. 1997. Aiken. No voucher. • Mature capsules. Close-up of capsules formed from an inferior ovary. Note the prominent top to each capsule where the sepals, petals, and anthers of the flower were attached. Nunavut, Baffin Island, Iqaluit. September, 2005. Photograph by Kathy Thornhill. No voucher. • Close-up of opening capsule. Left, mature capsule; right, capsule beginning to open by splitting from the top. Nunavut, Baffin Island, Iqaluit. September, 2005. Photograph by Kathy Thornhill. No voucher. • Capsule beginning to open. Capsule splitting into the four carpels with the thin axis of the of the axile placentation (a) in the centre. Nunavut, Baffin Island, Iqaluit. September, 2005. Photograph by Kathy Thornhill. No voucher. • Capsule split to base. The four carpels (1–4) of the capsule have split to the base. The axis is to the right. A few of the seeds that were born on the axis are held on the carpel walls by the plumose tufts of the tops of the seeds. Nunavut, Baffin Island, Iqaluit. September, 2005. Photograph by Kathy Thornhill. No voucher. • 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..