Flora of the Canadian Arctic Archipelago

S.G. Aiken, M.J. Dallwitz, L.L. Consaul, C.L. McJannet, L.J. Gillespie, R.L. Boles, G.W. Argus, J.M. Gillett, P.J. Scott, R. Elven, M.C. LeBlanc, A.K. Brysting and H. Solstad

Phippsia algida (Sol.) R. Br.

Poaceae, grass family.

Chlor. Melvill. 27. 1823.

Agrostis algida Sol. in Phipps, Voy. North Pole 200. 1774. Catabrosa algida (Sol.) Fr., Nov. fl. suec. mant. 3: 174. 1843.

Type: Described from Svalbard, Spitzbergen (BM).

Phippsia algida f. vestita Holmb., Bot. Not. 130. 1924. Catabrosa algida f. vestita (Holmb.) Polunin, Bull. Natl. Mus. Canada 92 (Biol. Ser. 24): 62. 1940.

Vegetative morphology. Plants perennial herbs; (2–)3.5–15 cm high; caespitose; not vegetatively proliferating by bulbils or fragmentation. Ground-level or under-ground stems not developed horizontally or vertically. Aerial stems circular or oval in cross-section; glabrous. Leaves distributed along the stems, or in a basal tuft (appearing basal in plants less than 5 cm high, cauline in plants 12–20 cm); alternate; simple; marcescent. Prophylls 4–10 mm long; with smooth veins; lacking pronounced keels (veins prominent, but not keeled). Petioles absent. Sheaths with the margins fused only in the lower part; glabrous; collars present. Ligules present; (0.3–)0.5–1(–1.6) mm long; membranous; glabrous; triangular; apices acute; entire. Blades 6–28 mm long; 1.2–3 mm wide (when flat). Blades appressed to the stem, or spreading; folded in bud; linear; flat, or folded; veins parallel; midvein conspicuously larger than the lateral veins, or midvein similar in size to other veins in the leaf; bulliform cells in distinct rows on either side of the midvein. Blades adaxial surface glabrous. Blades abaxial surface glabrous.

Reproductive morphology. Flowering stems present. Flowering culm nodes not rooting at the lower nodes (culms often forming a flat mat against the tundra early in the season, becoming erect at anthesis); not exposed. Flag leaf sheaths not inflated (uppermost culm leaf sheath somewhat inflated). Inflorescence paniculate; dense; linear, or ovate (branches appressed usually, or spreading, particularly at anthesis); (0.5–)1–2(–3) cm long; 3–7 mm wide; main axis glabrous. Number of inflorescence branches at lowest node 3–8. Inflorescence primary branches (0.9–)3–8(–9.5) mm long; glabrous; with appressed secondary branches. Spikelets pedicellate; disarticulating at the base of the spikelet; laterally compressed; lanceolate to ovate; (1.5–)3–7(–10) mm long; 0.5–0.8 mm wide. Florets per spikelet 1. Glumes caducous (falling early). First glume 0.02–0.03 × the length of the second glume; 0.2–0.35 × spikelet length; 0.05–0.2 mm long; ovate; glabrous; margins glabrous; veins 0; apex obtuse. Second glume 0.4 × as long as the spikelet or less; shorter than the lowest floret; ovate; 0.3–0.6 mm long; glabrous; veins 0. Lemma 1.3–1.6(–1.8) mm long; ovate (broadly so); rounded on the back (floret laterally compressed, but not strongly keeled); surface dull; surface glabrous, or sparsely scabrous (on the lower half, hairs if present are on the veins); veins 1–3. Lemma apex acute, or rounded; entire, or lacerate; glabrous, or ciliate (sparsely); awnless. Palea well developed; 1.1–1.3 mm long (almost as long as the lemma); with glabrous veins. Perianth reduced to lodicules. Stamens 3. Anthers (0.3–)0.5–0.7 mm long (only 1–2 anthers). Gynoecia superior. Carpels syncarpous; 3. Styles 2. Ovules 1. Fruit sessile. Fruit dry; a caryopsis; indehiscent. Fruit 1.2–1.9 mm long (broadly ovate-obovate in P. algida, narrowly ovate in P. concinna). Seeds 1.

Chromosome information. 2n = 28. 28 (4x). - Nannfeldt (1937); Flovik (1938, 1940 Svalbard); Sørensen and Westergaard in Löve and Löve (1948 Greenland?); Knaben (1950 southern Norway); Holmen (1952 Greenland); Sokolovskaya (1955); Löve and Löve (1956b Iceland); Jørgensen et al. (1958 Greenland); Bowden (1960a northern Canada); Sokolovskaya and Strelkova (1960); Hedberg (1962a northern Canada); Zhukova (1966, 1969 north eastern Asia, 1980 southern Chukotka); Mosquin and Hayley (1966 northern Canada); Knaben and Engelskjøn (1967 northern Norway); Johnson and Packer (1968 northwestern Alaska ); Knaben (1968 Alaska); Zhukova and Petrovsky (1972 north eastern Asia); Packer and McPherson (1974 northern Alaska); Vestre in Engelskjøn (1979 southern Norway); Dalgaard (1988 western Greenland).

Distribution. Northern hemisphere distribution: circumpolar; Greenland, Canada. High arctic. Range in the Canadian Arctic Archipelago widespread. Arctic Islands: Baffin, Devon, Ellesmere, Axel Heiberg, Parry Islands (Bathurst, Prince Patrick), Cornwallis, Banks, Victoria, Prince of Wales, Somerset, Southampton, Coats (Ellef RIngnes, Meighen).

Ecology and habitat. Substrates: hummocks, snow patches, river terraces, tundra (and disturbed habitats), ridges, seashores; imperfectly drained moist areas, or on seepage slopes (and seepage areas), or dry (CAN 30699); acidic, or calcareous (e.g. alkaline till), or nitrophilous; rocks (sandstone and its rubble), gravel, sand, silt, clay, till; peat (collected in bogs around lakes, CAN 503755). Habitats: Above high tide line. This is one of the first grasses to flower and set seed, which may contribute to it being an early colonizer of disturbed zones, both those made by human activities and those associated with permanent or late lying snow beds. Phippsia algida is highly nitrophilous, can tolerate highly alkaline soils, peat, imperfectly drained silts, and clays, such as mudflats in ephemeral river channels. It is found along lake shores at the base of colluvial slopes, in sandy and marshy places, generally on neutral to weakly alkaline terrain.

Taxon as an environmental indicator. This species is an early colonizer of disturbed soil. It occurs on otherwise bare ground around the town of Iqaluit, Baffin Island, particularly close to human habitation by the shoreline. In this town, when Phippsia was found away from the shoreline the area had been built up with material brought in from near the beach. Phippsia was observed 350 m above sea level on Melville Island as the only vascular plant present in a rocky area where even lichens were sparse. On Axel Heiberg Island, the Sawtooth Range of Ellesmere Island, and on Lower Savage Island, this species was found in 1999 in moist meadows and in snow patch areas up to 550–800 m above sea level. These areas are not recently disturbed and are also populated with Festuca, Braya, Saxifraga cernua, S. hirculus, Poa abbreviata and an abundance of mosses.
This species is usually perennial, but may be annual in some extreme environments having sufficient growing season for seed production in a single year. Weber (1999, personal communication) has found annual plants in the Rocky Mountains of Colorado.

Notes. Canadian specimens of Phippsia sometimes have lemmas with a few conspicuous hairs towards the base, but the lemmas never become "densely and stiffly hairy in the lower half" (Tutin 1980) as they do in the European P. concinna (Th. Fr.) Lindeb. That taxon is also distinguished by having a distinctly open panicle. No specimens assignable to P. concinna have been seeNorth Americaong Canadian records at CAN or DAO, although some specimens with slightly hairy lemmas have been so labeled. They would be better assigned to f. vestita Holmb.
Grulke and Bliss (1988) in a study of comparative life history characteristics of two High Arctic grasses noted that Phippsia algida is a ruderal species that colonizes bare soil in mesic sites, but also occurs across a broad range of habitats. Phippsia was observed to experience more frequent and severe drought stress at less extreme physical conditions than Puccinellia vaginata, the species it was being compared with. Drought stress and low temperatures reduced whole plant biomass of Phippsia of all ages in the study. Grulke and Bliss (1988) found that Phippsia was relatively short lived (living on average 23 years and a maximum of 34 years). They noted that the mean age of initial reproduction was relatively late when compared to temperate species, that is 17 + or - 6 years. Under all the field conditions monitored Phippsia had high, constant seed set. In short growing seasons, Phippsia continued high carbon allocation to reproduction, but also had high rates of plant mortality. The authors concluded that Phippsia algida has many characteristics of a ruderal and fills this role in a High Arctic ecosystem.
Bell and Bliss (1978) estimated that the roots of P. algida live 1.5 years. The first author has found plants of Phippsia to be one of the most difficult grasses to transplant to a greenhouse in the south, on three separate occasions. Although several plants were brought south they did not survive more than 2–3 weeks after being removed from an arctic environment. This species is possibly the first arctic grass to flower and set seed and appears to do so before mid-August in most habitats. On Herschel Island in July 1988, Phippsia had flowered and was setting seed before other grasses in the area were at anthesis.
In the former Soviet Union this species is considered to produce good pasture plants when fertilised (Tzvelev 1976). In Canada, the usually small plants, 5–10 cm high, are larger when they receive additional nutrients, see for example plants growing near the sewage lagoon in Iqaluit.
Aares et al. (2000) studied phylogeographic and taxonomic relations among 54 North Atlantic populations of Phippsia using isozyme techniques and genetically based morphological variation and found that their results supported the recognition of two distinct species, P. algida and P. concinna., the latter with at least two subspecies. They stated that both of these self-fertilizing arctic pioneer species were genetic allotetraploids almost without intra-populational variation. Also, the two species showed strikingly different phylogeographies in the North Atlantic region spite of their similarity in morphology, habitat ecology, mating system and dispersal ecology, and in spite of their present co-occurrence in many geographic areas, sometimes even in the same snowbeds. The same electrophoretic multilocus phenotype was observed in all populations of P. algida, and although this species showed considerable morphological variation, the variation was unstructured geographically. They concluded that P. algidia showed a pattern similar to other arctic species investigated in the North Atlantic region and that it has probably dispersed postglacially across the sea barriers among Greenland, Svalbard, Iceland, and Scandinavia. In contrast, P. concinna was virtually fixed for different multilocus phenotypes in the three main geographic areas analyzed (southern and northern Norway, Svalbard, Greenland) corresponding to fairly distinct divergence in morphology. The patterned suggested absence of postglacial among-area dispersal in this species and that it may have immigrated to the North Atlantic from different source areas and/or survived the last glaciation in situ.

Illustrations. • Habitat. This grass is an early colonizer of disturbed, often moist habitats. Quebec, Mine Raglan, 10 August, 1994, J. Cayouette. (DAO). • Plant in field. Phippsia algida. Growing on a mossy substrate. Canada. Quebec, Kangiqsujuak, 10 August, 1994, J.Cayouette. (DAO). • Plant habit. Plants growing on wet, calcareous silt in the flood plain of a stream. Ellesmere Island, Scoresby Bay 79°53'N, 71°33'W. Aiken 98–023. Photograph by Mollie MacCormac. • Close-up of plant. Plant growing in a wet hummocky meadow at base of glacier, ~450 m. Nunavut: Axel Heiberg Island. Large valley along upper Bukken River, 80°31.57'N, 92°21.83'W. Uncommon. 4 Aug. 1999. Photo by L. Consaul. L.J. Gillespie 6611, L.L. Consaul and R.J. Soreng. CAN. • Close-up of inflorescence. Post-anthesis inflorescence, with fruits forming. Nunavut, Axel Heiberg Island. Large valley along upper Bukken River, 80°31.57'N, 92°21.83'W. Wet hummocky meadow at base of glacier, ~450 m; uncommon. 4 Aug. 1999. Photo by L. Consaul. L.J. Gillespie 6611, L.L. Consaul and R.J. Soreng. CAN. • Close-up of inflorescence. Inflorescence approaching anthesis, with fluffy white stigmas exposed and solid pale yellow anthers beginning to show. The lemmas are red and the caducous glumes have already fallen. Nunavut, Axel Heiberg Island. Photo by L. Consaul, L.J. Gillespie, 6611, L.L. Consaul and R.J. Soreng. CAN. • Distribution map.