Life span

Perennial, moderately long-lived.

Growth form

Solitary graminoid herb growing in small, dense tussocks with intravaginal branching (e.g., with no runners or stolons). Culms 2–8 cm, procumbent or spreading, glabrous, sometimes with a few small spinules just beneath the inflorescence. Base of shoots surrounded by yellowish white sheaths of previous years' leaves. Usually no prophylls (scaly leaves without a developed blade at base of leafy and reproductive shoots) or sometimes one. The entire plant has a bluish bloom.

Leaf

Leaves glaucous green, filiform (narrowly convolute) or rarely almost flat, with ribs due to sclerenchyma (strings of discontinuous strengthening tissue), smooth or with a few spinules in margin and along veins apically. Basal leaves 1.5–3 cm long, about 1/2 length of culm, 0.3–0.5 mm broad (when convolute) or up to 1 mm (when flat). Leaf sheaths open. Uppermost culm leaf (the 'flag leaf') blade 0.2–0.7(0.9) cm, shaped like the bow of a boat (broader than linear), attached in the lower 1/3 of the culm. Ligula very short (less than 1 mm), truncate.

Inflorescence and Flower

The units of the inflorescence of Poaceae are the spikelets, nearly always numerous in a panicle or spike-like inflorescence. Spikelets are composed of 2 glumes (bracts for the spikelet) and one or more flowers (the term used below) or rather floral units often named ‘florets’ because we do not know what is the exact flower. A flower or floret is composed of a lemma with 1 mid vein (probably the floral bract), a palea with 2 mid veins (either 2 fused bracteoles or perhaps 2 fused perianth leaves), 3 small organs called ‘lodiculae’ and essential in the opening of the flower at anthesis (possibly transformed perianth leaves or transformed stamens), 3 stamens (mostly), and a gynoecium of 2 fused carpels with 2 feathery stigmas and one seed.

Inflorescence an open, narrow, several-sided, pale lilac panicle 0.8–1.5 cm long, occupying less than 1/4 of culm length. Panicle branches short (< 5 mm), smooth or with a few short spinules, each branch with 1(2) spikelets; the entire inflorescence often with less than 5 spikelets. Spikelets 5–7 x 1.0–2.0 mm (awns included), with 2–3 flowers. Bracts (glumes and lemmas) with rounded backs. Glumes 2.0–3.5 mm, the lower shorter than the upper, acute to acuminate, with 1–3 indistinct veins, glabrous and shiny, the upper or both glumes often with slender fringes or hairs apically and along margins. Lemmas (excluding awn) 3.5–4.5 mm, with several more or less distinct veins, scabrous, especially in distal part, with a very scabrous awn 1.8–2.3 mm. Paleas scabrous both on and between veins. Anthers well developed, 0.5–1.1 mm.

Fruit

Fruit an achene (with one seed).

Reproduction

Reproduction sexual; no vegetative reproduction. Wind pollinated. Seed production abundant (R. Elven observ.) and germination about 23 % (Alsos et al. 2013).

Passive dispersal of fruits inside florets, but the scabrous awn may attach to birds and animals and also facilitate some wind dispersal.

Comparison

The fescues of Svalbard belong to several groups, all within the major section Festuca. The Festuca rubra group is characterized by both intravaginal and extravaginal branching, the latter resulting in rhizomatous mats, and by several prophylls (reduced leaves usually without or with short blades) at the base of the shoots. All the others have intravaginal branching only, resulting in dense tussocks without any runners, and only one or no prophyll. The tussocky species are divided on four groups: Festuca baffinensis, the F. brachyphylla group, F. ovina, and the F. vivipara group. Festuca baffinensis and the F. brachyphylla group are both distinguished from F. ovina (and its more southern relatives) by the short anthers (less than 1.2 mm), whereas F. ovina has anthers of ca. 2 mm). Festuca baffinensis differs from the F. brachyphylla group in its hairy culms and its one-sided dark purple panicle; the others have glabrous culms and many-sided and otherwise coloured panicles. The F. vivipara group differs from the other groups in reproduction by bulbils (vivipary).

Within the Festuca brachyphylla complex, F. hyperborea is distinguished by its very short and spoon-shaped blade of the "flag leaf" (the uppermost leaf on the culm); the others (and especially F. brachyphylla) have an extended blade. Festuca brachyphylla differs from F. edlundiae in its erect culms scabrous just beneath the panicle. In F. edlundiae the culms usually grows more in lateral direction or are appressed to the ground, and they are smooth beneath the panicle, at least in Svalbard. Another difference is that F. edlundiae is glaucous (with a bluish bloom) on leaves and culms and has a pale lilac panicle; F. brachyphylla is green (not glaucous) on leaves and culms and has an often dark purple panicle.

Habitat

Festuca edlundiae is a specialist of gravelly ridges and open patches in heaths. It is a weak competitor but able to colonize open patches where and when they appear. The main sites are ridges and gravelly outcrops, and also gravel plains along rivers. Most occurrences are in areas with circumneutral to basic soils.

Distribution

The range of Festuca edlundiae is in the middle and northern arctic tundra zones and the clearly continental to transitional sections. This is the most frequent species of the F. brachyphylla group in Svalbard. It is rather common in the fjord districts of Spitsbergen northwards from Van Keulenfjorden (Nathorst and Wedel Jarlsberg Lands) and Kvalhovden (on the east coast, Heer Land) to Reinsdyrflya north of Liefdefjorden (Haakon VII Land) and the Wijdefjorden and Sorgfjorden areas (Andrée Land and Ny-Friesland). It is conspicuously absent along the west coast, including the otherwise species-rich Kongsfjorden area. In the other islands, it is known from several sites in the more favourable parts of Nordaustlandet, from Barentsøya and from one site northwest on Edgeøya.

In a circumpolar context, this species was mapped by Aiken et al. (1995) as mainly North American and Greenlandic, with very few records from NE Asia and from Svalbard, i.e., amphi-Atlantic and amphi-Beringian. In North America it was mapped only from the northern parts of the Canadian Arctic Archipelago and in Greenland mainly from the north and northeast. Since then it has been found in Alaska (connecting the American and Asian range, see Cortés-Burns et al. 2009), on Wrangel Island in NE Asia (R. Elven & H. Solstad observ. 2005, Oslo herbarium), and in more localities in W Greenland. The Russian material is still insufficiently revised (the main treatments of Alexeev, e.g., in Tolmachev et al. 1995, are based on a misunderstanding of F. hyperborea and before the recognition of F. edlundiae). We have, however, seen no material of F. edlundiae from any part of European Russia or Siberia. The ancestral connection of this species in Svalbard is therefore probably to the west.

Comments

The Festuca brachyphylla complex includes 3 species in Svalbard, the tetraploids (2n = 28) F. edlundiae and F. hyperborea and the hexaploid (2n = 42) F. brachyphylla (for the tetraploid F. baffinensis, see that species). A diploid (2n = 14) representative of the complex, F. brevissima, is found in Beringia. It is likely that many of its species have a hybridogeneous polyploid origin; however, the evolutionary pathways have not been revealed. The species delimitations were made quite clear by Aiken et al. (1995) and subsequently in Svalbard by Fjellheim (1999), Fjellheim et al. (2001), Guldahl (1999), and Guldahl et al. (2001). These Svalbard studies showed that Rønning's treatment (Rønning 1961, 1972) as F. brachyphylla and F. hyperborea was untenable, like most other previous studies of the group (and especially Alexeev's in Russia, see, e.g., Alexeev in Tolmachev et al. 1995). These authors tried to fit three distinct species into a two-species model (F. brachyphylla–F. hyperborea). Guldahl et al. (2001) argued that especially F. brachyphylla was enzymatically similar to N Scandinavian F. ovina (diploid, 2n = 14) and that the polyploids in the F. brachyphylla complex might possibly track back (evolutionary through allopolyploidizations) to diploids in the F. brachyphylla complex, especially to Beringian F. brevissima, and to the F. ovina complex (several species in Europe and N Asia).

The resolvement of the Festuca brachyphylla species complex is rather recent. Until the 1950s all material was considered as F. brachyphylla, or in some regions even as F. ovina. The first indication of some variation at species level was Holmen's report of a tetraploid species in N Greenland (Holmen 1952; suggested as a new species, F. ‘hyperborea’,but not formally described). Holmen considered it different from both the mainly diploid F. ovina and the known hexaploid F. brachyphylla. Frederiksen (1977) showed that F. hyperborea was distinctly different from both, specified its characters, and chose as type a specimen from northernmost Greenland. This approach was (prematurely) applied by Rønning (1972) in mapping the Svalbard monocotyledons, and he mapped all small-grown plants as F. hyperborea, and all more tall-grown plants as F. brachyphylla. Both maps are misleading as nearly all records in both maps belong to F. edlundiae described by Aiken et al. (1995). This is the only widely distributed species of this group in Svalbard, as suggested by Elven & Elvebakk (1996) and shown by morphological, cytological and molecular means by Fjellheim (1999), Fjellheim et al. (2001), Guldahl (1999), and Guldahl et al. (2001).

At present, the Festuca species of this complex are fairly well resolved in Svalbard: the widespread species is F. edlundiae described by Aiken et al. (1995), whereas the otherwise major arctic species F. brachyphylla (mapped as common by Rønning 1972) has a very small, compact range and the high arctic F. hyperborea (mapped as common by Rønning 1972) is very rare, here as elsewhere in the Eurasian Arctic.

Literature

Aiken, S.G., Consaul, L.L. & Lefkovitch, L.P. 1995. Festuca edlundiae (Poaceae), a high arctic, new species compared enzymatically and morphologically with similar Festuca species. – Systematic Botany 20: 374–392.

Alsos, I.G., Müller, E. & Eidesen, P.B. 2013. Germinating seeds or bulbils in 87 of 113 tested Arctic species indicate potential for ex situ seed bank storage. – Polar Biology 36: 819–830. Doi 10.1007/s00300-013-1307-7.

Cortés-Burns, H., Carlson, M.L., Lipkin, R., Flagstad, L. & Yokel, D. 2009. Rare vascular plants of the North Slope. A review of the taxonomy, distribution, and ecology of 31 rare plant taxa that occur in Alaska's North Slope region. – BLM Alaska Technical Report 58.

Elven, R. & Elvebakk, A. 1996. Part 1. Vascular plants. – In: Elvebakk, A. & Prestrud, P. (eds.), A catalogue of Svalbard plants, fungi, algae, and cyanobacteria. – Norsk Polarinstitutts Skrifter 198: 9–55.

Fjellheim, S. 1999. RAPD DNA and morphological variation in seminiferous taxa of the Festuca brachyphylla complex (Poaceae) in Svalbard. – Cand. scient. thesis, Univ. Oslo, Oslo.

Fjellheim, S., Elven, R. & Brochmann, C. 2001. Molecules and morphology in concert. II. The Festuca brachyphylla complex (Poaceae) in Svalbard. – American Journal of Botany 88: 869–882.

Frederiksen, S. 1977. The Festuca brachyphylla group in Greenland. – Botaniska Notiser 130: 269–277.

Guldahl, A.S. 1999. The Festuca brachyphylla complex in Svalbard: enzymatic, chromosomal, and ecological variation. – Cand. scient. thesis, Univ. Oslo, Oslo.

Guldahl, A.S., Borgen, L. & Nordal, I. 2001. Variation in the Festuca brachyphylla (Poaceae) complex in Svalbard, elucidated by chromosome numbers and isozymes. – Botanical Journal of the Linnaean Society 137: 107–126.

Holmen, K. 1952. Cytological studies in the flora of Peary Land, north Greenland. – Meddelelser om Grønland 128(5). 40 pp.

Rønning, O.I. 1961. Some new contributions to the flora of Svalbard. – Norsk Polarinstitutts Skrifter 124. 20 pp.

Rønning, O.I. 1972. The distribution of the vascular cryptogams and monocotyledons in Svalbard. – Det Kongelige Norske Videnskabers Selskabs Skrifter 1972-24. 63 pp.

Tolmachev, A.I., Packer, J.G. & Griffiths, G.C.D. (eds.) 1995. Flora of the Russian Arctic. I. Lycopodiaceae–Gramineae. – Univ. Alberta Press, Edmonton.