Flora of the Canadian Arctic Archipelago
| Flora of the Canadian Arctic Archipelago | |
English: Creeping goose grass, creeping alkali grass,
French: Puccinellie rampante, puccinellie trompeuse,
Inuktitut: Nakiruat, Iviit, ivisuka, ivitsuskaka.
Poaceae, Grass family.
Published in Contrib. U.S. Natl. Herb. 13: 78. 1910.
Type: U.S.A.: Alaska, "Kotzebue-Sund", leg. Eschscholtz 46. Holotype: LE-TRIN-2773.04.
Synonymy. Poa phryganodes Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, sér. 6, Sci. Math. 1: 389. 1831.
Glyceria phryganoides (Trin.) Fr. Novit. Fl. Suec. Mant. 3: Add. 176. 1843.
Atropis phryganodes (Trin.) Steffen, Beih. Bot. Centralbl. 44: 330. 1928.
Atropis phryganodes (Trin.) Krecz. in Komarov, Fl. SSSR, 2: 480. 1934 (later isonym).
Phippsia phryganodes (Trin.) A. Löve and D. Löve, Bot. Not. 128: 500. 1976.
Vegetative morphology. Plants 1–10(–15) cm high; perennial herbs; not caespitose (rarely forming somewhat tufted plants that flower). Only fibrous roots present. Ground level or underground stems horizontal, or absent (rarely in plants that are loosely caespitose and produce inflorescences); stoloniferous (with zig-zag stolons and extravaginal branching along the trailing stolons which makes this species unusual and conspicuous among the arctic grasses. Plants form loose turfs, with stolons arising from the leaf axils and rooting at the lower nodes. They also develop plantlets that detach readily); elongate. Aerial stems decumbent, or prostrate (or procumbent); not filiform. Leaves present; mostly along the stolons; alternate; marcescent. Petioles absent. Sheaths present; with the margins fused only in the lower part; glabrous; sheath collars present. Ligules present; 0.7–1.5 mm long (culm ligules; 0.4–0.9 mm long, stolon ligules to 1.5 mm long); membranous; glabrous; transversely oblong. Ligule apices acute; erose. Leaf blades simple. Leaves grass-like. Blades 10–35 mm long, 0.4–0.8 mm wide (when rolled), spreading (usually), rolled in bud, linear, without auricles (ligules slightly decurrent), involute, veins parallel, midvein conspicuously larger than the lateral veins (due to heavy sclerenchyma at abaxial surface opposite the vein). Blade adaxial surface glabrous. Blade abaxial surface glabrous.
Reproductive morphology. Flowering stems two or more per plant. Flowering stems circular or oval in cross section. Flowering stems rooting at the lower nodes; culm nodes not exposed (on flowering culms), or becoming exposed (along stolons); culm nodes number visible 0–2. Inflorescences paniculate (relatively rare); diffuse; lanceolate, or pyramidal; 1–4 cm long; 5–20 mm wide. Inflorescences main axis glabrous. Number of inflorescence branches at lowest node 2–3. Inflorescence primary branches 1–14 mm long; glabrous; with appressed secondary branches, or with spreading secondary branches. Spikelets disarticulating above the glumes (usually withering early, as the plants are sterile); lanceolate, or ovate; 6–9 mm long; 1.4–3.4 mm wide. Florets per spikelet 3–6(–7). Two glumes present. First glume 0.65–0.8 × the length of the second glume; 0.2–0.3 × spikelet length; 1.5–2.2 mm long; oblong; glabrous; margins glabrous; veins 2–3; apex obtuse. Second glume 0.4 × as long as the spikelet or less; shorter than the lowest floret; 2.3–2.8 mm long. Second glume ovate. Second glume glabrous (sometimes appearing so under low magnification), or with trichomes (sometimes small and infrequent); margins glabrous; veins 3. Rachilla not pronounced between the florets; extending beyond the uppermost floret; internode 0.9–1.1 mm long; internode glabrous. Callus differentiated. Lemma oblong, or lanceolate; 3.2–3.8(–4.5) mm long; rounded on the back; surface dull; surface glabrous (slightly emarginate, the margins thinner than the body of the lemma and whitish); veins 4–6 (faint); apex rounded; apex entire; apex glabrous; awnless. Palea well developed; 2.9–3.7 mm long; veins glabrous. Flowers bilaterally symmetrical (zygomorphic). Perianth represented by lodicules. Stamens present (preanthesis), or absent ((1.5-)2–2.5 indehiscent, without normal pollen); 3. Ovary superior; carpels 3; syncarpous. Styles 2. Placentation basal. Ovules per ovary 1 (rarely, if ever). Fruit sessile; dry; a caryopsis; indehiscent. Seeds 1.
Chromosome information. 2n = 21, 28 (not in the Flora region).
Puccinellia phryganodes (Trin.) Scribn. and Merr. subsp. neoarctica (A. Löve and D. Löve) comb. nov. needed:
2n (3x) = 21. Jørgensen et al. (1958, Greenland); Mosquin and Hayley (1966, northern Canada); Löve and Löve (1981a, northern Canada); Dalgaard (1989, western Greenland). This is the only taxon in the Canadian Arctic Islands.
Taxa outside the Canadian Arctic Archipelago:
Puccinellia phryganodes (Trin.) Scribn. and Merr. subsp. phryganodes
2n (4x) = 28. Zhukova (1967a, Northeastern Asia); Johnson and Packer (1968, Northwestern Alaska); Packer and McPherson (1974, northern Alaska); Zhukova and Petrovsky (1976, western Chukotka, this count could belong to subsp. sibirica);
Puccinellia phryganodes (Trin.) Scribn. and Merr. subsp. sibirica (Hada and A. Löve) comb. nov. needed
2n (3x) = 21. Nygren, in Löve and Löve (1948, northern Europe, 2n = 20, changed to 2n = 28 by Löve and Löve, 1975, to fit their concept); Engelskjøn (1979, northern Norway); Borgen and Elven (1983, northern Norway, two counts); 2n (4x) = 28. Sokolovskaya (1955), Nygren, in Jørgensen et al. (1958); Sokolovskaya and Strelkova (1960); Zhukova (1967a).
Puccinellia phryganodes (Trin.) Scribn. and Merr. subsp. vilfoidea (Andersson) Tzvelev
2n (4x) = 28. Flovik (1938, 1940 Svalbard); Engelskjøn (1979, Bear Island).
Diploid counts that have been reported are of Puccinellia geniculata (V.I.Krecz.) Hulten from Russia, which has been treated as a subspecies of P. phryganodes (P. phryganodes (Trin.) Scribn. subsp. geniculata (V.I.Krecz) Tzvelev), but which we are here treating as a separate species, following Elven et al. (2005). These authors note that one occurrence of this species in Alaska has been mapped by Hultén (1968a) in the Norton Sound area, which should be reinvestigated.
Ploidy levels recorded 3x, 4x.
Taxon as an environmental indicator. This species is found in areas usually covered by high tides or spray from the sea. Plants usually brownish red where salt dries onto the plants between tides. Where freshwater runoff washes out the salt on the plants between tides plants may be vivid green.
Ecology and habitat. Substrates: wet meadows (saline), marshes (saline or brackish), seashores; imperfectly drained moist areas, dry (specimen collected from dry clay (CAN 204374), moderately well-drained areas; rocks, gravel, sand, silt, clay; halophytic. Common on clay, silty and sandy shores and between boulders on coasts that are undisturbed by ice thrust. The taxon grows locally on recently eroded, Holocene marine sediments inland from the coast. Among the halophytic grasses this taxon has the most northern distribution. In the Low Arctic, for example around Iqaluit, it forms dense saline meadows that are often vivid orange with green streaks through them, where freshwater runoff between tides washes out the salt. In the High Arctic, it may be present as a few scattered runners.
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, coastal. Arctic islands: Baffin, Devon, Ellesmere, Axel Heiberg, Parry islands (Bathurst, Loughead, Melville and Prince Patrick), Cornwallis, Banks, Victoria, Somerset, King William, Southampton, Coats (Prince Charles and Resolution Islands, Melville Peninsula).
Northern hemisphere distribution. Circumpolar. Northern Fennoscandian, Kanin–Pechora, Svalbard – Franz Joseph Land, Polar Ural – Novaya Zemlya, Taimyr – Severnaya Zemlya, Anabar–Olenyok, Kharaulakh, Yana–Kolyma, 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. Bowden (1961) considered this taxon to be a sterile, triploid hybrid of unknown parentage. Sørensen (1953) recognised four variants of P. phryganodes, based on pedicels thickened or not thickened, floret numbers per spikelet, lemma lengths, and epidermal cell features. He concluded that Canadian material is mainly the "Greenland type" (= P. phryganodes subsp. neoarctica), but generally with better developed pistils. Tzvelev (1976) recognised various subspecies, but remarked that none of them is morphologically clearly separated, and they may not deserve recognition. The description by Hughes and Halliday (1980) differs from the one given here, in the following characters: "spikelets not more than 4 florets; lower glume 1.2–1.7 mm, upper glume 1.5–2.3 mm; lemma 1.5–2.5 mm". The Hughes and Halliday (1980) description is based on northern Norwegian plants, i.e., triploids with even more reduced florets and anthers (or the pale shadows of very clearly aborted anthers) (R. Elven, personal communication, 1999).
In the Panarctic Flora checklist, Elven et al. (2003) followed the concept of Tzvelev (1976) on the variation in the 'phryganodes' aggregate, noting that the morphological differences seem constant (albeit small) and are partly associated with ploidy differences. The entities they suggest represented major geographical allopatric or parapatric races, and qualified as subspecies in their concepts. This treatment suggested the following subspecies:
Puccinellia phryganodes (Trin.) Scribn. and Merr. subsp. phryganodes as most probably a tetraploid, 2n = 28, that occurs in the Russian Far East and Alaska. The plants investigated of this entity in W and NW Alaska are also fully pollen-fertile, in opposition to most or all plants elsewhere. We have investigated plants also from the type region ("Kotzebue Sund"). The W-NW Alaskan entity (phryganodes s. str.) is tetraploid (2n = 28) (Elven, personal communication, 2005).
Puccinellia phryganodes (Trin.) Scribn. and Merr. subsp. neoarctica (Á. Löve and D. Löve) comb. nov. needed, a triploid, 2n = 21, that occurs in Svalbard, Greenland, and Canada.
Puccinellia phryganodes (Trin.) Scribn. and Merr. subsp. sibirica (Hadač and Á. Löve) comb. nov. needed, Type, European Russia: Waigatsch, Sinus Warnek, 14.07.1907, leg. O. Ekstam (S) holotype, a taxon with triploids and tetraploids that occurs from Norway to the Russian Far East.
Puccinellia phryganodes (Trin.) Scribn. and Merr. subsp. vilfoidea (Andersson) Tzvelev, in Tolm. Fl. Arct. URSS 2: 186. 1964. Described from Svalbard.
Puccinellia phryganodes (Trin.) Scribn. and Merr. subsp. geniculata (Turcz. ex V.I.Krecz.) Tzvelev, Novosti Sist. Vyssh. Rast. 8: 77. 1971. This is a diploid taxon that occurs in Alaska and may be species-level distinct. In 2005, Elven et al. (2005) treated this taxon as a separate species.
Following Elven et al. (2003) we draw attention to this discussion, but as some of the combinations have not been published, we do not follow it yet (March 2005).
Jefferies and Gottlieb (1983) studied genetic variation within and between populations of Puccinellia phryganodes. Because they assumed that the taxon is a sterile triploid, they expected that the level of genetic variation would be low, both within and between populations. They examined the electrophoretic mobilities of isozymes of 12 systems in plants from three widely separated populations in Arctic Canada (La Prouse Bay, Manitoba, 58°4'N, 94°4'W; Tuktoyaktuk, N.W.T. 69°27'N, 133°00'W; and Alexandra Fiord, Ellesmere Island, Nunavut 78°53'N, 75°55'W) and found a high level of variability both within and between populations. They suggested that the unexpected discovery of significant amounts of genetic variability indicated that sexual processes occur in this species.
Siira and Merila (1985), working in Finland, found that around the coast of Bothnian Bay, in the Baltic Sea area, the populations of P. phryganodes had decreased greatly during the previous 20 years. They noted that the species has "a narrow ecological amplitude and poor colonization ability", and suggested that in some areas, continuous land uplift causes fairly rapid changes in the ecological conditions, and primary succession is not always the same. They also noted that human activity had influenced the nature of the shorelines studied.
Illustrations. • Habitat. Large salt-marsh completely dominated by Puccinellia phryganodes. The shallow pools are caused by sea ice freezing on the ground in winter and lifting up turfs during spring tides. Norway, Finnmark, Neiden. 9 June, 1983. Photograph by R. Elven. • Habitat: Dorset. Light green coloured plants growing in a limited saline meadow where runoff stream enters the ocean at the end of the valley suburb. Nunavut, Baffin Island, Cape Dorset. 5 August, 2005. Aiken. No voucher. • Habitat: salt effects. Plants growing in a tidal meadow. Those with green leaves are in a fresh water run-off where the salt is rinsed off between tides. Those that are not rinsed turn orange brown from salt-burn which occurs when the salt water dries on the plants. Nunavut, Baffin Island, Iqaluit, near the airport. 1986. Aiken. No voucher. • Stolons on beach: Dorset. Braching stolons radiating from a central plant on beach in area often covered at high tide. Baffin Island, Cape Dorset. 2 August, 2005. Aiken. No voucher. Scale bar in cm. • Habitat. Mud flat early in the season with grey stolons of previous season's growth and in the centre of the plants, new season's green growth developing. Southern Quebec, Raglan Mine, Douglas Harbour, southwest fiord. Photograph J. Cayouette. • Stolon (runner). Aboveground horizontal stem, stolon, with branches forming at the base of sheaths but appearing inside the sheath of the leaf below. Norway, Finnmark, Porsanger, Valddat. 31 July, 1977. Photograph by R. Elven. Voucher at TROM. • Plant in Habitat. Saline meadow dominated mostly by sterile plants of Puccinellia phryganodes. The plant was developing inflorescences where it was growing over the rock (next image). Nunavut, Southampton Island, Coral Harbour. Aiken and Brysting 01–076. CAN. • Close-up of inflorescence. Inflorescence of a species that does not often flower. Spikelets have numerous florets at anthesis, and relatively large anthers, (1.5-) 2–2.5 mm long, that are without normal pollen. Nunavut, Southampton Island, Coral Harbour. Aiken and Brysting 01–076. CAN. • Spikelet drawing. From Sorensen (1953). Reproduced with permission from Meddeleser om Gr�nland. • 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.
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