14.06 PUCCINELLIA Parl.
Jerrold I. Davis
Laurie L. Consaul

Plants annual, biennial, or perennial; usually cespitose, sometimes weakly or strongly stoloniferous and mat-forming. Culms 2–100 cm, erect or decumbent, sometimes geniculate; internodes hollow. Sheaths open to the base or nearly so; auricles absent; ligules membranous, acute to truncate, entire or erose; blades flat, folded, or involute. Inflorescences terminal panicles, open to contracted; branches smooth or scabrous, some branches longer than 1 cm; pedicels usually longer than 3 mm, thinner than 0.5 mm. Spikelets pedicellate, subterete to weakly laterally compressed, with 2–10 florets; disarticulation above the glumes, beneath the florets. Glumes usually unequal, sometimes subequal to equal, usually distinctly shorter than the lowest lemma in the spikelets, sometimes only slightly shorter, rarely longer, membranous, rounded or weakly keeled, veins obscure or prominent, apices unawned; lower glumes 1(3)-veined; upper glumes (1)3(5)-veined; calluses blunt, glabrous or pubescent; lemmas membranous to slightly or distinctly coriaceous, glabrous or pubescent, pubescence sometimes restricted to the bases of the veins, rounded or weakly keeled, at least distally, (3)5(7)-veined, veins obscure to prominent, more or less parallel distally, usually not extending to the apices, lateral veins sometimes reduced, apical margins with or without scabrules, apices usually acute to truncate, sometimes acuminate, entire or serrate to erose, unawned; paleas subequal to the lemmas, scarious or membranous distally, 2-veined, veins terminating at or beyond midlength; lodicules 2, free, glabrous; anthers 3; ovaries glabrous. Caryopses shorter than the lemmas, concealed at maturity, oblong, terete to dorsally flattened, falling free or with the palea or both the lemma and palea attached; hila oblong, about 1/3 or less the length of the caryopses. x = 7. Named for Benedetto Puccinelli (1808–1850), an Italian botanist.

Puccinellia, a genus of approximately 120 species, is most abundant in the middle and high latitudes of the Northern Hemisphere. There are 21 species in the Flora region, of which 3 are introduced. Ten are confined to the arctic, four are circumarctic and two are transberingian. Most species of Puccinellia are halophytes, either in coastal habitats or in saline or otherwise mineralized soils of interior habitats. Polyploidy, selfing, and hybridization are widespread in the genus, and many of the species boundaries are controversial. Several of the species with Arctic distributions have received different taxonomic treatments in North America and Eurasia.

The angle of the panicle branches (whether erect, ascending, etc.) refers to their position when the caryopses are mature. Lemma measurements should be made on the lowest lemma in the spikelets. Principal features of the lemmas, as used in the key and descriptions, are as follows. Scabrules (short, pointed hairs, similar in form to those that occur on the pedicels and inflorescence branches of many species of Puccinellia, and generally requiring magnification to observe) often occur along the distal margins of the lemmas. When present, they may be few and irregularly scattered, with gaps between them that are either wider than the individual scabrules (e.g., in some P. pumila), or arranged in a continuous palisade-like row that lacks gaps (e.g., in P. distans). Independent of the presence or absence of scabrules, the lemma margins may be entire (e.g., in P. pumila and P. distans) or serrate to erose (e.g., in P. andersonii, and P. vahliana).

SELECTED REFERENCES Argus, G.W. and K.M. Pryer. 1990. Rare Vascular Plants in Canada: Our Natural Heritage. Canadian Museum of Nature, Ottawa, Ontario, Canada. 191 pp.; Consaul, L.L. and L.J. Gillespie. 2001. A re-evaluation of species limits in Canadian Arctic island Puccinellia (Poaceae): Resolving key characters. Canad. J. Bot. 79:927–956; Consaul, L.L., L.J. Gillespie, and K.I. MacInnes. [in press]. Addition to the flora of Canada? A specimen from the Arctic Archipelago, Northwest Territories links two allopatric species of alkali grass; Davis, J.I. 1983. Phenotypic plasticity and the selection of taxonomic characters in Puccinellia (Poaceae). Syst. Bot. 8:341–353; Fernald, M.L. and G.A. Weatherby. 1916. The genus Puccinellia in eastern North America. Rhodora 18:1–23; Porsild, A.E. 1964. Illustrated Flora of the Canadian Arctic Archipelago, ed. 2, rev. Bulletin of the National Museum of Canada No. 146 [Biological Series No. 50]. R. Duhamel, Queen’s Printer, Ottawa, Ontario, Canada. 218 pp.; Scribner, F.L. and E.D. Merrill. 1910. The grasses of Alaska. Contr. U.S. Natl. Herb. 133:47–92; Sørensen, T.J. 1953. A revision of the Greenland species of Puccinellia Parl. with contributions to our knowledge of the arctic Puccinellia flora in general. Meddel. Grønl. 136:1–169; Sørensen, T.J. 1955. Puccinellia agrostidea, Puccinellia bruggemannii, Puccinellia poacea. Pp. 78–82 in A.E. Porsild. The Vascular Plants of the Western Canadian Arctic Archipelago. Bulletin of the National Museum of Canada No. 135 [Biological Series No. 45]. E. Cloutier, Queen’s Printer, Ottawa, Ontario, Canada. 226 pp.; Swallen, J.R. 1944. The Alaskan species of Puccinellia. J. Wash. Acad. Sci. 34:16–32;Tsvelev, N.N. 1995. Puccinellia. Pp. 237–263 in J.G. Packer (ed., English edition). Flora of the Russian Arctic, vol. 1, trans. G.C.D. Griffiths. University of Alberta Press, Edmonton, Alberta, Canada. 330 pp. [English translation of A.I. Tolmachev (ed.). 1964. Arkticheskaya Flora SSSR, vol. 2. Nauka, Leningrad [St. Petersburg], Russia. 272 pp.]

 

The interactive dichotomous key and interactive multientry key are currently not available.

 

1. Plants stoloniferous perennials, forming low, often extensive mats; most plants lacking inflorescences, the spikelets, when present, usually not producing mature pollen or caryopses ... P. phryganodes
1. Plants annual, biennial, or cespitose perennials, sometimes stoloniferous but not mat-forming; plants reproducing sexually, forming mature pollen and caryopses ... 2
2. Lemmas slightly to markedly coriaceous for most or all of their length; plants of temperate regions ... 3
2. Lemmas mostly membranous or herbaceous, apical margins sometimes hyaline; plants of temperate and arctic regions ... 5
3. Lemmas with hyaline apical margins; lemma midveins prominent ... P. rupestris
3. Lemmas with coriaceous apical margins; lemma midveins obscure ... 4
4. Lemmas 1.8–3 mm long; lower branches of the panicles ascending to erect, spikelet-bearing nearly to the base; anthers 0.6–1 mm long ... P. fasciculata
4. Lemmas 3–5 mm long; lower branches of the panicles erect to descending, spikelet-bearing from about midlength; anthers 1.5–2.6 mm long ... P. maritima
5. Plants annual, of temperate regions ... 6
5. Plants perennial, of temperate and arctic regions ... 7
6. Lemma apices acute; lemmas 2.5–4 mm long, veins glabrous or hairy, particularly on the basal 1/2, short (about 0.1 mm) hairs sparsely and evenly distributed between the veins ... P. simplex
6. Lemma apices obtuse to truncate; lemmas 1.8–2.2 mm long, veins densely hairy on the basal 1/2–3/4, glabrous between the veins ... P. parishii
7. Palea veins with curly, intertwined hairs proximally, scabrous distally; plants of arctic regions ... 8
7. Palea veins glabrous, shortly ciliate, or with fewer than 5 longer hairs proximally, never with curly intertwined hairs, scabrous or smooth distally; plants of temperate and arctic regions ... 12
8. Pedicels smooth; apical margins of the lemmas smooth, veins obscure or distinct ... 9
8. Pedicels scabrous; apical margins of the lemmas scabrous, sometimes minutely so, veins obscure ... 10
9. Lower glumes 2/3 to nearly as long as the adjacent lemmas; culms 5–15 cm; panicles 2–4 cm, anthers 0.8–1.5 mm ... P. vahliana
9. Lower glumes usually less than 2/3 as long as the adjacent lemmas, culms 15–40 cm; panicles 5–8 cm, anthers 1.5–2.5 mm ... P. wrightii
10. Culms 50–65 cm tall; panicles 15–30 cm long ... P. groenlandica
10. Culms 5–35 cm tall; panicles 1–13 cm long ... 11
11. Lemmas 3.5–5.2 mm long; panicles (4)5–13 cm long ... P. angustata
11. Lemmas 2.8–3.8 mm long; panicles 1–4 cm long ... P. bruggemannii
12. Lemma margins smooth or with a few scabrules at and near the apices ... 13
12. Lemma margins densely scabrous at and near the apices ... 17
13. Lemmas 2–2.5 mm long, usually purple with whitish margins, veins distinct, apices obtuse to truncate; lemmas and palea veins smooth and glabrous; pedicels smooth ... P. tenella
13. Lemmas 2.4–4.6 mm long, variously colored, margins not white, veins obscure to distinct, apices acute to truncate; lemmas and palea veins glabrous or hairy on the lower portion, often scabrous distally; pedicels smooth or scabrous ... 14
14. Lemmas glabrous or with a few hairs on the lower portion of the veins; lemma apices entire; plants of temperate regions or the low arctic, but not of the high arctic ... 15
14. Lemmas usually sparsely to moderately hairy, particularly on the vein bases, sometimes glabrous; lemma apices entire, irregularly serrate, or erose; plants of the low and high arctic ... 16
15. Pedicels scabrous; palea veins scabrous distally; anthers 1–2 mm long; plants not littoral ... P. lemmonii (in part)
15. Pedicels smooth or with a few scattered scabrules; palea veins smooth or with a few scabrules distally; anthers 0.5–1.2 mm long; plants littoral ... P. pumila
16. Panicles with (2)3–5 branches at the lowest node; lemmas 2.5–3.7 mm long, veins obscure to distinct, apices entire or slightly erose; anthers 1.2–2.2 mm long ... P. arctica (in part)
16. Panicles usually with 2 branches at the lowest node; lemmas 3–4.5 mm long, veins obscure, apices irregularly serrate or erose; anthers 0.8–1.2 mm long ... P. andersonii (in part)
17. Lemmas 1.5–2.2 mm long, apices widely obtuse to truncate; anthers 0.4–0.8 mm long; lower panicle branches horizontal to descending ... P. distans
17. Lemmas 2–5 mm long, apices usually acute to obtuse, occasionally acuminate or rounded; anthers 0.5–2.2 mm long; lower panicle branches erect to descending ... 18
18. Lemma apices irregularly serrate or erose; lemmas 3–4.5 mm long; anthers 0.8–1.2 mm long ... P. andersonii (in part)
18. Lemma apices entire or slightly erose; lemmas 2–4.5(5) mm long; anthers 0.5–2.2 mm long ... 19
19. Pedicels smooth or with a few scattered scabrules; lemmas glabrous or with a few hairs on the lower 1/2, principally along the veins; anthers 1.5–2 mm long; plants restricted to mineralized springs in California ... P. howellii
19. Pedicels smooth to uniformly scabrous; lemmas glabrous or sparsely to moderately hairy on the lower 1/2; anthers 0.5–2.2 mm long; plants of varied habitats, including hot springs ... 20
20. Lemma midveins often extending to the apical margins; lemma apices acute; lemmas mostly smooth, midveins often slightly scabrous distally; leaf blades involute, 1.2–1.9 mm wide when flattened; leaves concentrated at the base of plant; plants of inland, temperate habitats ... P. lemmonii (in part)
20. Lemma midveins usually not extending to the margins; lemma apices usually acute to obtuse, occasionally acuminate; lemmas scabrous or smooth distally; leaf blades involute or flat, 0.5–6 mm wide when flat, leaves ranging from nearly all basal to evenly distributed along the culms; plants of coastal and inland habitats in temperate and arctic regions ... 21
21. Culms 10–100 cm tall; lower glumes 0.5–1.6 mm long; plants usually growing south of 65º N latitude ... 22
21. Culms 6–30(40) cm tall; lower glumes 0.8–2.5 mm long; plants usually growing north of 65º N latitude ... 23
22. Pedicel epidermal cells not tumid, pedicels uniformly scabrous; lower branches of the panicles erect to descending; lemmas (2)2.2–3(3.5) mm long; plants usually of interior habitats, occasionally of coastal habitats ... P. nuttalliana (in part)
22. Pedicel epidermal cells often tumid, pedicels sparsely to densely scabrous; lower branches of the panicles usually erect to ascending, occasionally spreading to descending; lemmas (2.2)3–4.5(5) mm long; plants of coastal habitats ... P. nutkaensis
23. Anthers 1.2–2.2 mm long; lateral margins of the lemmas often inrolled ... P. arctica (in part)
23. Anthers 0.6–1.2 mm long; lateral margins of the lemmas usually not inrolled ... 24
24. Lemmas 2.8–4 mm long; panicles usually barely exserted from the sheaths ... P. vaginata
24. Lemmas 2–2.8 mm long; panicles usually distinctly exserted from the sheaths ... P. nuttalliana (in part)

 

1. Puccinellia phryganodes (Trin.) Scribn. & Merr.
Goose Grass, Puccinellie Rampante, Puccinellie Trompeuse

Plants perennial; stoloniferous, often forming extensive low mats. Culms 2–15 cm, erect or decumbent. Ligules 0.4–1.5 mm, acute, obtuse, or truncate, entire; blades 0.4–2.2 mm wide when flat, 0.2–1 mm in diam-eter when involute. Panicles usually not developed, if devel-oped, 1–7 cm, diffuse, lower branches ascending, spikelets usually confined to the distal 1/3; pedicels scabrous, usually with tumid epidermal cells. Spikelets 6–9 mm, with 3–6(7) florets. Glumes rounded or slightly keeled over the back, veins obscure or distinct, apices acute or obtuse; lower glumes 1.5–2.2 mm; upper glumes 2.3–2.8 mm; calluses glabrous or almost so; lemmas 3.2–3.8(4.5) mm, membranous, usually glabrous, occasionally sparsely hairy, backs rounded, 5-veined, veins obscure, not extending to the margins, apices acute or rounded, apical margins whitish, hyaline, smooth, entire; palea veins glabrous; anthers (1.5)2–2.5 mm, almost always indehiscent, mature pollen rarely produced. Caryopses rarely developed. 2n = 21, 28.

Puccinellia phryganodes is a widespread and common circumpolar arctic species that grows on seashores at or near the high tide line, in wet saline meadows, and in saline or brackish marshes.

 

2. Puccinellia rupestris (With.) Fernald & Weath.
Stiff Saltmarsh Grass

Plants annual or biennial; tufted, not mat-forming. Culms 5–50 cm, erect to decumbent. Ligules 1–2 mm, obtuse to truncate, entire; blades 2–6 cm wide, flat or folded. Panicles 2–10 cm, contracted at matur-ity, lower branches ascending, spikelet-bearing nearly to the base; pedicels scabrous, sometimes only slightly so, sometimes with tumid epidermal cells. Spikelets 4–9 mm, with 2–6 florets. Glumes rounded to keeled over the back, midveins obscure or prominent, extending to the apices, often scabrous distally, lateral veins obscure, apices acute to obtuse; lower glumes 1–2 mm; upper glumes 2–3.2 mm; calluses with a few hairs; lemmas 2.5–4 mm, somewhat coriaceous for most of their length, glabrous or sparsely hairy on the lower 1/2, particularly along the veins, backs rounded to keeled, 5-veined, midveins and sometimes all veins prominent, midveins extending to the margin, often excurrent, apical margins hyaline, smooth to densely scabrous, apices acute to obtuse, entire; palea veins shortly ciliate proximally, shortly ciliate to scabrous distally; anthers 0.8–1.2 mm. 2n = 42.

Puccinellia rupestris grows in coastal and noncoastal habitats in Eurasia; North American collections were apparently introduced in ballast.

 

3. Puccinellia fasciculata (Torr.) E.P. Bicknell
Borrer’s Saltmarsh Grass

Plants short-lived perennials; cespitose, not mat-forming. Culms 10–65 cm, usually decumbent and geniculate, sometimes erect. Ligules 1–2 mm, obtuse to truncate, entire; blades 2–7 mm wide, flat, folded, or involute. Panicles 4–16 cm, compact to diffuse at maturity, linear to pyramidal, lower branches ascending to erect, spikelet-bearing nearly to the base; pedicels slightly to densely scabrous, often with tumid epidermal cells. Spikelets 3–6 mm, with 2–6 florets. Glumes rounded over the back, veins prominent to obscure, apices acute to obtuse; lower glumes 0.8–1.6 mm; upper glumes 1.2–2.3 mm; calluses with a few hairs; lemmas 1.8–3 mm, slightly to markedly coriaceous throughout, glabrous or with a few hairs near the base, backs rounded, 5-veined, veins obscure, midveins usually excurrent, sometimes ending at the margins, apices acute to obtuse, entire, apical margins smooth or with a few scattered scabrules; palea veins glabrous proximally, scabrous to shortly hispid near midlength, scabrous distally; anthers 0.6–1 mm. 2n = 28.

Puccinellia fasciculata is native to Europe. In the Flora region, it is found principally along the east coast, but it is also established at a few sites in Arizona and Utah, and has been reported from Nevada. All occurrences in the Flora region are probably the result of human introductions.

 

4. Puccinellia maritima (Huds.) Parl.
Common Saltmarsh Grass, Puccinellie Maritime

Plants perennial; cespitose, often stoloniferous, not mat-forming. Culms 20–100 cm, erect to decumbent. Ligules 1–3.5 mm, obtuse to truncate, entire; blades 2–4.4 mm wide, flat to involute. Panicles 3–30 cm, compact to diffuse at maturity, lower branches erect to descending, spikelet-bearing from about midlength; pedicels usually densely scabrous, occasionally with only a few scattered scabrules, often with tumid epidermal cells. Spikelets 5.5–13 mm, with 4–9 florets. Glumes rounded over the back, veins obscure, apices acute to obtuse; lower glumes 2–3.4 mm; upper glumes 3–4.5 mm; calluses with few to many hairs; lemmas 3–5 mm, slightly to markedly coriaceous throughout, sparsely to densely hairy in the lower 1/2, principally along the veins, backs rounded, 5-veined, veins obscure, usually not extending to the margins, sometimes the midvein doing so, apical margins usually smooth, sometimes with a few scattered scabrules, occasionally densely scabrous, apices acute to obtuse, entire; palea veins ciliate proximally, scabrous to short-ciliate distally; anthers 1.5–2.6 mm. 2n = 14–77, usually 56.

Puccinellia maritima grows in coastal environments in North America and Greenland. It is native to Europe; most or all occurrences in the Flora region are probably the result of human introduction.

 

5. Puccinellia simplex Scribn.
Western Alkali Grass

Plants annual; not mat-forming. Culms usually erect, 2–25 cm. Ligules 1–3 mm, acute to obtuse, entire; blades 0.7–2 mm wide, flat to involute. Panicles 1–18 cm, compact, mostly linear at maturity, primary branches usually spikelet-bearing to the base, lower branches erect; pedicels densely scabrous or with only a few scattered scabrules, often also with a few hairs, often with tumid epidermal cells. Spikelets 3.5–8 mm, with 2–7 florets. Glumes rounded to weakly keeled over the back, veins obscure to prominent, apices acute; lower glumes 1.3–2 mm; upper glumes 2.3–3 mm; calluses hairy; lemmas 2.5–4 mm, mostly herbaceous, usually with about 0.1 mm hairs distributed sparsely and evenly between the veins, longer hairs also usually present along the veins and near the base, basal hairs often longest, twisted and tangled, backs rounded, sometimes keeled distally, 5-veined, veins usually obscure, sometimes prominent, midveins sometimes reaching the margins, other veins usually not doing so, apical margins often hyaline, smooth or with a few scattered scabrules, apices acute, entire; palea veins with hairs, hairs on the proximal portion longer, twisted and somewhat tangled, hairs on the distal 2/3 usually short and straight; anthers 0.2–0.5 mm. 2n = 56.

Puccinellia simplex is widespread in, and mostly confined to, saline soils of central California. The records from Utah probably reflect introductions.

 

6. Puccinellia parishii Hitchc.
Parish’s Alkali Grass

Plants annual; not mat-forming. Culms 3–22 cm, erect. Leaves basally concentrated; ligules 1–2 mm, obtuse to truncate, entire; blades 0.2–1.2 mm wide, flat to involute. Panicles 1–8.5 cm, compact to diffuse at maturity, lower branches erect to descending, usually spikelet-bearing to the base; pedicels densely scabrous or with a few scattered scabrules, often with tumid epidermal cells. Spikelets 3.5–5 mm, with 2–7 florets. Glumes rounded over the back, veins obscure to prominent, apices acute to obtuse; lower glumes 1–2 mm; upper glumes 1.8–2.2 mm; calluses hairy; lemmas 1.8–2.2 mm, mostly herbaceous, densely hairy over the proximal 1/2–3/4 of the veins, glabrous between the veins, backs rounded, 5-veined, veins obscure to prominent, not extending to the margins, apical margins hyaline, smooth or with a few scattered scabrules, apices obtuse to truncate, entire; palea veins glabrous proximally, hairy at midlength, glabrous or scabrous-ciliate distally; anthers 0.4–0.5 mm. 2n = 14.

Puccinellia parishii grows in saline seepage areas in California, Arizona, and New Mexico.

 

7. Puccinellia vahliana (Liebm.) Scribn. & Merr.
Vahl’s Alkali Grass, Puccinellie de Vahl

Plants perennial; cespitose, not mat-forming. Culms 5–15 cm, erect. Ligules 1–2.5 mm, acute to obtuse, entire; blades 2–8 mm wide, flat or folded. Panicles 2–4 cm, usually con-tracted and dense, sometimes slightly diffuse at maturity, lowest node usually with long and short branches, lower branches erect to ascending, spikelets usually confined to the distal 2/3; pedicels glabrous and smooth, lacking tumid epidermal cells or with very small tumid epidermal cells. Spikelets 3.8–6.5 mm, with 2–4(5) florets. Glumes broadly ovate, enfolding the bases of the lower lemmas, rounded over the back, veins obscure or distinct, apices acute to obtuse; lower glumes 2–3.5 mm, at least 2/3 as long as the adjacent lemmas; upper glumes 2.4–4 mm; rachilla internodes abruptly broadened at the point of attachment to the lemmas, less than 0.09 mm thick; calluses with a few hairs; lemmas 3–5.2 mm, usually herbaceous and mostly purplish, sometimes mem-branous with purple veins, basal 1/2 hairy over and between the veins, backs rounded, 5-veined, veins obscure or distinct, often dark purple, not extending to the margins, apical margins hyaline, often yellowish, smooth, apices acute, entire, becoming erose with age; palea veins with curly, intertwined hairs on the proximal portion, scabrous distally; anthers 0.8–1.5 mm. 2n = 14.

Puccinellia vahliana is an arctic species that is circumpolar, except in the Beringian region. In the Flora region, it extends from Alaska through northern Canada to Greenland. It is generally non-halophytic, growing in calcareous gravel, sand, clay, or moss of imperfectly drained moist areas, and on seepage slopes from near sea level to 700 m, or, rarely, in seasonally dry, turfy sites. It is often a pioneering species in moist clay and silt by alpine brooks, ephemeral lakes, glacial runoff streams, and on snowbeds. The roots of this species and P. wrightii are characteristically thicker and more tightly curled than those of other Puccinellia species. It sometimes hybridizes with Phippsia algida.

 

8. Puccinellia wrightii (Scribn. & Merr.) Tzvelev
Wright’s Alkali Grass

Plants perennial; cespitose, not mat-forming. Culms 15–40 cm, erect. Ligules 1.5–3 mm, acute to obtuse, entire; blades 2–8 mm wide, flat or folded. Panicles 5–8 cm, open and diffuse at maturity, lowest node usually with long branches, lower branches ascending, hori-zontal, or descending, spikelets usually confined to the distal 1/3; pedicels glabrous and smooth, lacking tumid epidermal cells. Spikelets 4–7 mm, with 4–5 florets. Glumes narrow, not enfolding the base of the lower lemmas, rounded over the back, veins obscure or distinct, apices acute to obtuse; lower glumes 1.7–3 mm, usually less than 2/3 as long as the adjacent lemmas; upper glumes 2.5–4 mm; rachilla internodes abruptly or gradually broadened to the point of attachment with the lemmas, at least the lowest internode usually more than 0.09 mm thick; calluses with a few hairs; lemmas 4–5 mm, mostly herbaceous, hairy along and between the veins on the proximal 1/2, backs rounded, 5-veined, veins obscure or distinct, not extending to the margins, apical margins hyaline, smooth, apices acute, entire, becoming erose with age; palea veins with curly, intertwined hairs proximally, these hairs rarely lacking, scabrous distally; anthers 1.5–2.5 mm. 2n = 14.

Puccinellia wrightii is an uncommon arctic species. Its range extends from the Chukotka Peninsula in the Russian Far East to western Alaska. Like P. vahliana, its roots are characteristically thicker and more tightly curled than those of other Puccinellia species.

 

9. Puccinellia groenlandica T.J. Sørensen
Greenland Alkali Grass

Plants perennial; cespitose, not mat-forming. Culms 50–65 cm, erect to decumbent. Ligules 2.5–3.3 mm, obtuse to truncate, entire; blades 1.8–3.2 mm wide, flat. Panicles 15–30 cm, diffuse at maturity, lower branches ascending to horizontal, spike-lets usually confined to the distal 2/3; pedicels uniformly scabrous, lacking tumid epidermal cells. Spikelets 7–10 mm, with 4–7 florets. Glumes rounded over the back, veins obscure, apices acute to obtuse; lower glumes 1.8–2.5 mm, up to approximately 1/2 as long as the adjacent lemmas; upper glumes 2.5–3 mm; rachilla internodes slightly and gradually broadened to the point of attachment with the lemmas, at least the lowest internodes usually more than 0.09 mm thick; calluses with a few hairs; lemmas 3–4.4 mm, mostly herbaceous, hairy on the proximal 1/2, principally along the veins, backs rounded or keeled distally, 5-veined, veins obscure, not extending to the margins, apical margins hyaline, uniformly scabrous, apices obtuse to acute, entire; palea veins with curly, intertwined hairs proximally, scabrous distally; anthers 1.1–1.5 mm. 2n = 56.

Puccinellia groenlandica grows in littoral and nonlittoral environments. It is endemic to Greenland.

 

10. Puccinellia angustata (R. Br.) E.L. Rand & Redfield
Tall Alkali Grass, Puccinellie Étroite

Plants perennial; cespitose, not mat-forming. Culms 10–35 cm, erect to decumbent. Ligules (0.8)1–3(4) mm, acute, obtuse, or truncate, entire; blades usually involute and 0.5–1.2 mm in diameter, sometimes flat and 0.5–3 mm wide. Panicles (4)5–10(13)cm, usually con-tracted, occasionally diffuse at maturity, lower nodes with 2–3(4) branches, lower branches erect to ascending, spikelets usually confined to the distal 1/2; pedicels scabrous, lacking tumid epidermal cells. Spikelets 4–10 mm, with 3–5(6) florets. Glumes rounded over the back, veins obscure, apices acute; lower glumes 1.5–2.8 mm, usually less than 1/2 as long as the adjacent lemmas; upper glumes 2.2–4.2 mm; rachilla internodes slightly and gradually broadened to the point of attachment with the lemmas, at least the lowest internode usually more than 0.09 mm thick; calluses with a few hairs; lemmas 3.5–5.2 mm, herbaceous, often purplish, hairy on the proximal 1/2 along the veins, sometimes also between the veins, backs rounded, 5-veined, veins obscure, not extending to the margins, apical margins hyaline, scabrous, apices acute, entire or erose; palea veins with curly, intertwined hairs proximally, scabrous distally; anthers (0.6) 0.8–1.1(1.5) mm. 2n = 42.

Puccinellia angustata is a common and widespread arctic species that grows in disturbed silty or sandy sediments. It is usually non-littoral, but when in coastal areas it grows above the influence of high tide.

 

11. Puccinellia bruggemannii T.J. Sørensen
Bruggemann’s Alkali Grass

Plants perennial; cespitose, not mat-forming. Culms 5–12 cm, erect to decumbent. Ligules 0.8–2 mm, acute or obtuse, entire; blades usually involute and 0.5–1.1 mm in diameter, sometimes flat and 0.7–2.5 mm wide. Panicles 1–4 cm, con-tracted at maturity, lowest node with 2(3) branches, lower branches erect to ascending, spikelets usually confined to the distal 1/3; pedicels scabrous, lacking tumid epidermal cells. Spikelets 3.5–6.5 mm, with 2–4 florets. Glumes rounded over the back, veins obscure, apices acute to obtuse; lower glumes 0.7–2 mm, less than 1/2(2/3) as long as the adjacent lemmas; upper glumes 1.5–2.8 mm, broadly elliptic; rachilla internodes slightly and gradually broadened to the point of attachment with the lemmas, at least the lowest internode usually more than 0.09 mm thick; calluses with a few hairs; lemmas 2.8–3.8 mm, herbaceous, hairy on the lower 1/2 along and between the veins, backs rounded, 5-veined, veins obscure, not extending to the margins, apical margins herbaceous, scabrous or scabridulous, apices acute or somewhat obtuse, entire or erose, slightly incurved; palea veins with curly, intertwined hairs proximally, scabrous distally; anthers 0.7–1.1 mm. 2n = 28.

Puccinellia bruggemannii is restricted to arctic islands of Canada and northern Greenland. In Canada, it is a widespread yet local northern, western, and central arctic island species. It is found in calcareous, barren, gravelly, sandy, or silty sites, and is sometimes coastal. Although P. bruggemannii has been reported as non-littoral (Porsild 1964) and probably non-halophilous (Sørensen 1955), specimens keying to this species have been found near the sea coast: for example, paratype specimens collected on Beechey Island and in the vicinity of salt springs on Axel Heiberg Island.

Puccinellia bruggemannii sometimes superficially resembles Poa abbreviata or small Poa glauca in the field, because of its small, dense inflorescence.

 

12. Puccinellia tenella (Lange) Holmb. ex Porsild
Tundra Alkali Grass

Plants perennial; cespitose, not mat-forming. Culms 3–16 cm, erect. Ligules 0.5–1.7 mm, acute, obtuse, or truncate, entire; blades usually involute and 0.4–0.7 mm in diameter, occasionally flat and 0.5–1.5 mm wide. Panicles 1.6–5.5 cm, usually contracted, sometimes diffuse, lower branches erect or ascending, spikelets usually confined to the distal 1/2; pedicels smooth, with tumid epidermal cells. Spikelets 3.5–7 mm, with 3–6 florets. Glumes slightly keeled over the back, veins distinct, apices acuminate to acute; lower glumes 0.7–1.3 mm; upper glumes 1.3–1.8 mm; calluses glabrous or with 5 or fewer hairs shorter than 0.1 mm; lemmas 2–2.5 mm, herbaceous, usually purple with whitish margins, smooth, glabrous or with a few hairs on the bases of the lateral veins, backs rounded, 5-veined, veins distinct, not extending to the margins, apical margins smooth, apices obtuse to truncate, entire or slightly erose; palea veins smooth, glabrous; anthers 0.6–0.9 mm. 2n = 14.

Puccinellia tenella is a halophytic, circumpolar subarctic and low arctic species. It is found above the high tide zone on sandy spits, in salt marshes, in silty soils, and among granitic rocks to 30 m above sea level. The above description applies to P. tenella subsp. langeana (Berlin) Tzevlev, the only subspecies in the Flora region. Puccinellia alaskana Scribn. & Merr., considered a subspecies of P. langeana (Berlin) T.J. Sørensen ex Hultén [= P. tenella] by Sørensen (1953), has acute lemmas and hairs on the palea. It is discussed under P. pumila.

 

13. Puccinellia lemmonii (Vasey) Scribn.
Lemmon’s Alkali Grass

Plants perennial; cespitose, not mat-forming. Culms 5–40 cm, usually erect. Leaves basally concentrated; ligules 0.8–2.2 mm, obtuse to acute, mostly entire, sometimes slightly erose; blades involute, 1.2–1.9 mm wide when flattened. Panicles 2–18 cm, compact to diffuse at maturity, lower branches ascending to descending, usually spikelet-bearing to the base; pedicels scabrous, lacking tumid epidermal cells. Spikelets 3.5–8 mm, with 2–6 florets. Glumes rounded over the back, veins obscure, apices acute to obtuse; lower glumes 0.7–1.5 mm; upper glumes 1.4–3 mm; calluses with a few hairs; lemmas 2.4–4 mm, herbaceous, mostly smooth, usually glabrous, sometimes with a few hairs near the base, principally along the veins, backs usually rounded, sometimes weakly keeled distally, 5-veined, veins obscure, midveins often slightly scabrous and prominent in the distal 1/2, often extending to the apical margins, lateral veins not extending to the margins, apical margins ranging from smooth to scabrous, entire, not white, apices acute, entire; palea veins glabrous or shortly ciliate proximally, uniformly scabrous distally; anthers 1–2 mm. 2n = 14.

Puccinellia lemmonii grows in non-littoral saline environments in the western portion of the contiguous United States. Reports from Saskatchewan are probably based on depauperate specimens of P. nuttalliana.

 

14. Puccinellia pumila (Vasey) Hitchc.
Smooth Alkali Grass, Puccinellie Naine

Plants perennial; usually ces-pitose, occasionally appearing rhizomatous or stoloniferous after rooting at the nodes of buried stems, infrequently sto-loniferous, not mat-forming. Culms 8–40 cm, erect to decumbent. Ligules 0.8–2.5 mm, obtuse to truncate, entire; blades 1–3 mm wide, flat to involute. Panicles 3–20 cm, dense to diffuse at maturity, lower branches ascending to descending, spikelets borne from near the bases or confined to the distal 2/3; pedicels smooth or with a few scattered scabrules, often with tumid epidermal cells distally. Spikelets 4–9 mm, with 3–7 florets. Glumes rounded over the back, veins obscure to distinct, apices acute to obtuse; lower glumes 1.4–2(4) mm; upper glumes 2–3(9) mm; calluses glabrous or with a few hairs; lemmas 2.5–4.6 mm, herbaceous, glabrous or with a few hairs on the vein bases, backs rounded, 5-veined, veins obscure, not extending to the margins, apical margins usually smooth, occasionally with a few scattered scabrules, entire, not white, apices acute to obtuse, entire; palea veins glabrous, smooth or with a few scabrules distally; anthers 0.5–1.2 mm. 2n = 14 [for Puccinellia alaskana], 42, 56.

Puccinellia pumila is primarily North American, growing on the Pacific, Arctic, and Atlantic coasts. It also grows in Kamchatka, Russia (Tsvelev 1995). It generally grows in sand and among stones in protected intertidal environments. A few specimens with exceptionally long glumes and lemmas were treated by Fernald and Weatherby (1916) as P. paupercula var. longiglumis Fernald & Weath.; they are regarded here as representing extremes of P. pumila.

Puccinellia alaskana Scribn. & Merr., here included in P. pumila, was considered a subspecies of P. langeana (Berlin) T.J. Sørensen ex Hultén [= P. tenella] by Sørensen (1953), but more closely resembles P. pumila. It differs morphologically from P. pumila mainly in its relatively distinct lemma veins. It also differs from most specimens of P. pumila in having smaller lemmas (2.5–3 mm) and anthers (0.5–0.9 mm), and in being diploid. It represents the Aleutian Islands component of the geographic distribution given for P. pumila. Its status is currently under investigation. Molecular data obtained as this volume went to press (Consaul et al. [in prep.]) tend to support recognition of P. alaskana as a distinct species.

 

15. Puccinellia arctica (Hook.) Fernald & Weath.
Arctic Alkali Grass

Plants perennial; cespitose. Culms 10–30(40) cm, erect. Leaves basally concentrated; ligules 0.9–3 mm, acute, obtuse, or truncate, entire, margins decurrent; blades usually flat and 0.5–2.2 mm wide, some-times involute and 0.2–1.6 mm in diameter. Panicles 3–11 cm, diffuse or contracted at maturity, lowest nodes with (2)3–5 branches, lower branches ascending to horizontal, spikelets usually confined to the distal 2/3; pedicels scabrous, without tumid epidermal cells. Spikelets 4.5–7(9.5) mm, with (2)3–6(8) florets. Glumes rounded over the back, veins distinct or obscure, lateral margins often inrolled, apices acute to obtuse; lower glumes 0.8–2.1(2.5) mm; upper glumes 1.8–3 mm; calluses with a few hairs; lemmas 2.5–3.7 mm, herbaceous or membranous, often translucent, often purplish, hairy, particularly on the bases of the veins, backs rounded, 5-veined, veins obscure to distinct, midveins scabrous or smooth distally, sometimes extending to the apical margin, sometimes excurrent, lateral veins not extending to the margins, lateral margins often inrolled, apical margins often hyaline and yellowish, scabrous, entire or slightly erose, apices acute to obtuse; palea veins glabrous, smooth proximally, scabrous from midlength or just below midlength to the apices; anthers 1.2–2.2 mm. 2n = 14.

Puccinellia arctica is restricted to the North American arctic, where it grows in silt, clay, and sandy substrates near the coast, and on alkaline, sparsely vegetated soils further inland. As treated here, it includes three entities that are sometimes treated as distinct species: P. arctica sensu stricto, P. poacea T.J. Sørensen, and P. agrostidea T.J. Sørensen. Puccinellia arctica sensu stricto is restricted to the southwestern arctic, P. poacea to the high arctic (Ellesmere and Axel Heiberg islands), and P. agrostidea to the southwestern arctic and possibly also Ellesmere Island. There are no morphological characters known for distinguishing these three entities. The first two may differ from the latter by the relatively frequent presence of small scabrules along the upper lemma midvein, slightly more distinct veins, and frequent yellowish margins to the lemma. The taxonomic validity of these characters was not completely understood at the time this treatment was written, but molecular analyses being conducted as this volume went to press (Consaul et al. [in prep.]) suggest that this group is best represented as a single species, P. arctica. Argus and Pryer (1990) stated that all three entities are rare in Canada.

 

16. Puccinellia andersonii Swallen
Anderson’s Alkali Grass

Plants perennial; cespitose, not mat-forming. Culms 10–25 cm, usually decumbent or genic-ulate. Ligules 1–2.8(3.3) mm, acute, obtuse, or truncate, entire or slightly erose; blades usually involute and 0.5–1 mm in diameter, sometimes flat and 0.8–2 mm wide. Panicles (3.5) 5–8 cm, diffuse or contracted at maturity, lowest node almost always with 2 branches, lower branches ascending to horizontal in fruit, spikelets usually confined to the distal 1/3; pedicels smooth or scabrous, with tumid epidermal cells. Spikelets 5–7(9.5) mm, with (2)4–5(7) florets. Glumes rounded over the back, veins obscure, apices acute; lower glumes 1–2 mm; upper glumes 2–3 mm, often borne distinctly above the lower glumes; calluses glabrous or hairy; lemmas (3) 3.2–4(4.5) mm, herbaceous or membranous, glabrous or sparsely hairy on the bases of the veins, backs rounded, 5-veined, veins obscure, not extending to the margins, apical margins smooth or scabrous, not white, apices usually acute, occasionally rounded, irregularly serrate or erose; palea veins glabrous, smooth proximally, scabrous distally; anthers 0.8–1.2 mm. 2n = 56.

Puccinellia andersonii is a widespread, coastal arctic species. It grows near the tideline and on otherwise barren, reworked marine sediments of eroded flood plains. Its decumbent growth form often gives it an unhealthy appearance. It is unique among Puccinellia species in the Flora region in having blunt, rather than pointed, scabrules in the apical region of its lemmas.

 

17. Puccinellia distans (Jacq.) Parl.
European Alkali Grass, Puccinellie à Fleurs Distantes

Plants perennial; cespitose, not mat-forming. Culms 5–60 cm, erect to decumbent. Ligules 0.8–1.2 mm, obtuse to truncate, usually entire; blades 1–7 mm wide, flat to involute. Panicles 2.5–20 cm, diffuse at maturity, lower branches horizontal to descending, spikelets usually confined to the distal 2/3; pedicels scabrous, lacking tumid epidermal cells. Spikelets 2.5–7 mm, with 2–7 florets. Glumes rounded over the back, veins obscure, apices acute to truncate; lower glumes 0.4–1.3 mm; upper glumes 0.9–1.8 mm; calluses with a few hairs; lemmas 1.5–2(2.2) mm, mostly herbaceous, glabrous or sparsely hairy on the lower 1/2, principally along the veins, backs rounded, 5-veined, veins obscure, not extending to the margins, apical margins hyaline and often yellowish, uniformly and densely scabrous, apices widely obtuse to truncate, entire; palea veins shortly ciliate proximally, glabrous, sometimes scabrous distally; anthers 0.4–0.8 mm. 2n = 14, 28, 42.

Puccinellia distans is a Eurasian native, reportedly introduced in North America, where it is widespread, particularly as a weed in non-littoral environments, including the margins of salted roads. It is also found occasionally in coastal environments.

A specimen in the Smithsonian Institution attributed to Puccinellia tenuiflora (Griseb.) Scribn. & Merr. by Scribner and Merrill (1910) is a robust example of P. distans; others of this taxon in North America remain to be investigated. Puccinellia hauptiana (Trin. ex V.I. Krecz.) Kitag. has been reduced to P. distans subsp. hauptiana (Trin. ex V.I. Krecz.) W.E. Hughes. This taxon may represent a tetraploid component of P. distans; several specimens identified as P. hauptiana in Eurasia have a tetraploid chromosome count. Specimens from Alaska, the Yukon, and Saskatchewan, identified as P. hauptiana on the basis of their relatively narrow leaves (1–2 mm wide) and small anthers (0.5–0.6 mm), appear to be native in these regions and require further study with regard to their relationship with P. distans.

 

18. Puccinellia howellii J.I. Davis
Howell’s Alkali Grass

Plants perennial; cespitose, not mat-forming. Culms 7–40 cm, erect to ascending. Ligules 1.5–2.7 mm, obtuse, entire or minutely and irregularly serrate; blades involute, 1.4–2.2 mm wide when flattened. Panicles 2–13 cm, compact to diffuse at maturity, lower branches erect to descending, spikelet-bearing from near the base or confined to the distal 1/2; pedicels smooth or with a few scattered scabrules, sometimes with tumid epidermal cells. Spikelets 3–8 mm, with (1)2–5 florets. Glumes rounded or weakly keeled over the back, veins obscure, apices acute to obtuse; lower glumes 0.8–1.9 mm; upper glumes 1.7–2.5 mm; calluses with a few hairs; lemmas 2.4–3.3 mm, herbaceous, glabrous or with a few hairs near the base, principally along the veins, backs rounded or weakly keeled distally, 5-veined, veins obscure, not extending to the margins, apical margins uniformly and densely scabrous, apices acute to obtuse, entire; palea veins smooth and glabrous proximally, smooth or scabrous distally; anthers 1.5–2 mm. 2n = unknown.

Puccinellia howellii is known only from the type locality in Shasta County, California, where it is a dominant element of the vegetation associated with a group of three mineralized seeps. Isozyme profiles suggest that it is a polyploid.

 

19. Puccinellia nuttalliana (Schult.) Hitchc.
Nuttall’s Alkali Grass

Plants perennial; cespitose, not mat-forming. Culms 10–100 cm, usually erect. Leaves either concentrated at the base or distributed along the culms; ligules 1–3 mm, obtuse, usually entire, sometimes slightly erose; blades 1–4 mm wide, flat to involute. Panicles 5–30 cm, compact to diffuse at maturity, usually distinctly exserted from the sheaths, lower branches usually erect to diverging, occasionally descending, spikelet-bearing from the base or on the distal 2/3; pedicels scabrous, lacking tumid epidermal cells. Spikelets 3.5–9 mm, with 2–7 florets. Glumes rounded over the back, veins obscure, apices acute to obtuse; lower glumes 0.5–1.5 mm, usually less than 1/2 as long as the adjacent lemmas; upper glumes 1–2.8 mm; rachilla internodes slightly and gradually broadened to the point of attachment with the lemmas; calluses with a few hairs; lemmas (2)2.2–3(3.5) mm, herbaceous, glabrous or sparsely hairy on the proximal 1/2, principally along the veins, backs rounded, 5-veined, veins obscure, not extending to the margins, smooth distally, lateral margins inrolled or not, apical margins uniformly and densely scabrous, apices acute to obtuse, entire; palea veins glabrous, short-ciliate, or with a few long hairs proximally, smooth or scabrous distally; anthers 0.6–2 mm. 2n = 28, 42, 56.

Puccinellia nuttalliana is a widespread and variable species, restricted to the Flora region. It grows principally in the interior, but is also found in coastal settings, where it is difficult to distinguish from P. nutkaensis. Northern, primarily boreal or southern arctic populations with relatively short lemmas and anthers (2–2.8 mm and 0.6–0.9 mm, respectively), and with a few long hairs on the lower palea veins, have sometimes been recognized as P. borealis Swallen.

 

20. Puccinellia nutkaensis (J. Presl) Fernald & Weath.
Alaska Alkali Grass, Pacific Alkali Grass, Puccinellie Brillante

Plants perennial; cespitose, occasionally appearing rhizom-atous or stoloniferous after rooting at the nodes of buried stems, not mat-forming. Culms 10–90 cm, usually erect, sometimes decumbent. Leaves usually distributed evenly along the culms; ligules 1–3 mm, obtuse to truncate, entire; blades 1.5–6 mm wide when flat, flat to involute. Panicles 5–30 cm, compact to diffuse at maturity, lower branches usually erect to ascending, occasionally spreading to descending, spikelet-bearing from near the base or the spikelets confined to the distal 1/2; pedicels from sparsely to densely scabrous, epidermal cells often tumid. Spikelets 3.5–12 mm, with 3–7 florets. Glumes rounded over the back, veins obscure, apices acute to truncate; lower glumes 1–1.6 mm; upper glumes 2–3 mm; calluses with a few hairs; lemmas (2.2)3–4.5(5) mm, herbaceous, glabrous or sparsely hairy on the proximal 1/2, principally along the veins, backs rounded, 5-veined, veins obscure, not extending to the margins, midveins smooth distally, apical margins uniformly and densely scabrous, apices usually acute to obtuse, sometimes acuminate, entire; palea veins glabrous or with short hairs proximally, scabrous distally; anthers 0.5–1.4 mm. 2n = 42, 56.

Puccinellia nutkaensis grows in coastal habitats of continental North America and Greenland, generally in sand and stones in protected intertidal environments. It is variable in form, ranging from diminutive plants that resemble P. pumila to tall, erect plants, often with dense or open inflorescences, resembling P. nuttalliana. Larger plants on the Pacific coast have been called P. grandis Swallen, and those on the Atlantic coast P. lucida Fernald & Weath., but there are many plants of intermediate stature.

 

21. Puccinellia vaginata (Lange) Fernald & Weath.
Sheathed Alkali Grass, Puccinellie Engainée

Plants perennial; cespitose, not mat-forming. Culms (6)9–20 (35) cm, erect or decumbent. Leaves basally concentrated on small plants, distributed along the culms in larger plants; ligules 1–3 mm, acute, obtuse, or truncate, entire; blades 1–2 mm wide when flat, 0.5–1.6 mm in diameter when involute. Panicles (3)6–12(14) cm, diffuse or contracted and usually barely exserted from the sheaths at maturity, lower branches ascending to horizontal, spikelets usually confined to the distal 2/3; pedicels slightly scabrous, epidermal cells tumid. Spikelets 4–8 mm, with (2)4–5(6) florets. Glumes rounded over the back, veins distinct or obscure, margins not inrolled, apices broadly acute or obtuse; lower glumes 1.3–2.1 mm; upper glumes 1.3–2.6(3.4) mm; calluses with a few hairs; lemmas 2.8–4 mm, herbaceous, thin, often translucent, bases hairy, particularly on the veins, backs rounded, 5-veined, veins obscure, not extending to the margins, lateral margins usually not inrolled, apical margins uniformly and densely scabrous, apices broadly acute or obtuse, entire or slightly erose; palea veins glabrous proximally, scabrous distally; anthers 0.7–1.1(1.2) mm. 2n = 56.

Puccinellia vaginata is a widespread North American arctic species. It is locally common in Greenland to about 78º N latitude; it is uncommon in Canada, where it is found primarily in the low arctic, rarely in the high arctic. One specimen is reported from eastern Siberia. It grows on coastal marine sediments and on eroding, raised marine sediments inland, and forms large tussocks near bird cliffs.