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four-angle spikerush, square-stem spike-rush, square-stem spikesedge

Kamchatka spike-rush, éléocharide du kamtchatka

Habit Plants perennial; rhizomes 1.5–4 mm thick, soft to hard, longer internodes 3–8 cm, scales 5–10 mm, tubers absent. Plants perennial, mat-forming; rhizomes evident, sometimes vertical, long, 0.5–1.5 mm thick, soft, cortex often breaking loose, longer internodes 1–3 cm, scales fugaceous, 4–7 mm, thinly membranous, not fibrous.
Culms

acutely quadrangular, (30–)45–105 cm × (1–)2–5.4 mm, soft to firm, internally spongy, transverse septa incomplete;

plants never forming filiform, flaccid culms.

terete, often with to 10 blunt ridges when dry, 4–60 cm × 0.3–1.8 mm, soft to firm, internally spongy.

Leaves

distal leaf sheaths persistent, membranous, apex narrowly acute to acuminate, sometimes prolonged into a bladelike portion to 8 cm.

distal leaf sheaths persistent, not splitting, proximally red (to stramineous), distally red or stramineous or green, not callose, membranous, apex usually pale red, broadly obtuse to narrowly acuminate, tooth absent.

Spikelets

not proliferous, (15–)20–76 × 3–5(–6) mm;

rachilla joints bearing obscure winglike remnants of floral scales;

proximal scale empty, amplexicaulous, (1–)2.2–5.4 mm;

floral scales (28–)45–135, 2–3 per mm of rachilla, stramineous to pale brown, usually with pale to dark brown submarginal band, midrib region sometimes greenish, broadly obovate to ovate, (4–)4.5–6.2 × 2.8–5 mm, subcartilaginous, apex rounded to obtuse.

ovoid to lanceoloid, 4–20 × 3–6 mm, apex acute;

proximal scale amplexicaulous, entire;

subproximal scale with flower;

floral scales often spreading in fruit, 5–20, 1–2 per mm of rachilla, entirely blackish brown to red-brown, or midrib regions paler, broadly ovate, 2.5–5 × 2–2.5 mm, apex rounded to subacute, not carinate.

Flowers

perianth bristles 6–7, whitish to brown, slender, often markedly unequal, shorter than achene or some equalling tubercle, sparsely retrorsely spinulose to smooth;

anthers stramineous to red-brown, 2.3–2.9 mm;

styles 3-fid, sometimes 2-fid.

perianth bristles 0–6(–9), light brown to stramineous, slender to stout, often unequal, much shorter than to equaling achene;

stamens 3;

anthers yellow-brown to medium brown, 1.3–2.5 mm;

styles 2-fid.

Achenes

yellow or pale green to brown or purplish, biconvex, obovoid to obpyriform, 1.8–3 × 1.3–2 mm, almost smooth to markedly sculptured at 10–15X, each face with 19–38 rows of almost linear, transversely elongated cells, which are sometimes isodiametric at achene base, apex often constricted to neck 0.3–0.4 mm wide.

not persistent, yellow, stramineous, or medium brown, obovoid (mostly very broadly so), thickly biconvex, angles obscure, 1–1.5(–1.9) × 1–1.6 mm, apex truncate, neck absent or very short, finely reticulate at 20–30X, or rugulose with 20 or more horizontal ridges in a vertical series, or sometimes smooth.

Tubercles

dark brown or whitish, deltoid to high-pyramidal or lanceoloid, 0.7–1.5 × 0.4–1 mm, often spongy.

whitish, outline subrectangular or trapezoidal, sometimes broadly rounded, seldom pyramidal, (0.5–) 0.7–1.5 × (0.5–)0.8–1.4 mm, usually with vertical rows of depressions (cancellate).

2n

= 12.

Eleocharis quadrangulata

Eleocharis kamtschatica

Phenology Fruiting early summer–winter. Fruiting summer.
Habitat Shallow water of fresh lake and pond shores, marshes Brackish (to fresh?) marshes, meadows, ponds, inland in Asia
Elevation 10–600 m (0–2000 ft) 0–30 m (0–100 ft)
Distribution
from FNA
AR; CA; CT; DE; FL; GA; IL; IN; KY; LA; MA; MI; MO; MS; NC; NJ; NY; OH; OK; OR; PA; SC; TX; VA; WI; WV; ON; s to c Mexico
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from FNA
AK; BC; MB; NL; QC; Asia (China, Japan, Korea, Russia)
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Discussion

We have not seen voucher specimens for the reports of Eleocharis quadrangulata from Kansas. Plants with greenish achenes, longer bristles, and longer anthers than the average are known from Tennessee.

The tubercles of Eleocharis quadrangulata are often spongy as in E. obtusetrigona.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

The markedly different chromosome counts reported from North America (2n = 12, from the Queen Charlotte Islands, British Columbia) and from Asia (2n = 38–40, 42, 56) suggest that more than one taxon should be recognized within Eleocharis kamtschatica as currently broadly defined. The three similar species E. kamtschatica, E. komarovii, and E. sachalinensis have been recognized for Russia (I. D. Zinserling 1935); more recent authors have recognized only E. kamtschatica (T. V. Egorova 1981 and A. E. Kozhevnikov 1988, for the former U.S.S.R.; J. Ohwi 1965, for Japan; S.-O Strandhede 1967; E. Hultén 1968 for Alaska; Tang T. and Wang F. T. 1961, for China). Pending a taxonomic revision, the following five names, all from eastern Asia, should probably be treated as synonyms of E. kamtschatica: E. komarovii Zinserling, E. mitrata (Franchet & Savatier ex Makino, E. sachalinensis (Meinschauser) B. Fedtschenko, E. savatieri Svenson, and E. triflora Komarov, not E. triflora Boeckeler. In E. kamtschatica in the broad sense, the size and form of the achenes and tubercles, on which most of the probable synonyms listed above are based, and the form of the distal leaf sheath summits do not seem to be correlated with each other or with other characters. In North America E. kamtschatica is known only from coastal habitats; in Asia it also occurs inland, especially around hot springs.

North American plants of Eleocharis kamtschatica seem to be separable into two groups: (1) Smaller plants, with culms mostly shorter than 20 cm, spikelets 5–13 × 3–4 mm, and floral scales 2.5–4 mm are known from about 20 localities on the Pacific Coast from Kodiak Island, Alaska, to the Queen Charlotte Islands and Prince Rupert, British Columbia, as well as from Hudson Bay in Manitoba, James Bay and the Gulf of St. Lawrence in Quebec, and Labrador. These small plants include the voucher specimens of the 2n = 12 chromosome count from British Columbia, and seem to fit E. kamtschatica as described by I. D. Zinserling (1935). (2) More robust plants, with culms mostly 20–70 cm, spikelets 7–20 × 4–6 mm, and floral scales 4–5 mm, are known from about 12 localities from Norton Sound in west-central Alaska to central British Columbia. These robust plants fit the descriptions of E. sachalinensis by I. D. Zinserling(1935), of E. savatieri Svenson from Japan (H. K. Svenson 1939), and of E. kamtschatica as described by J. Ohwi (1965), and are very similar to several specimens from Japan that I have seen.

Because Eleocharis kamtschatica closely resembles E. uniglumis except for size of the tubercles, plants without achenes cannot be reliably identified to species. The floral scales and receptacles in E. kamtschatica are very sparsely placed, with only 1–2 scales per mm of rachilla, compared with 3–4 mm of rachilla in most North American plants of E. uniglumis. Some North American collections, e.g., from the Kobuk sand dunes in northwestern Alaska, as E. uniglumis, and James Bay in Quebec, as E. kamtschatica, are intermediate between E. kamtschatica and E. uniglumis in the relative sizes of the tubercles and achenes.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Source FNA vol. 23, p. 120. FNA vol. 23, p. 76.
Parent taxa Cyperaceae > Eleocharis > subg. Limnochloa Cyperaceae > Eleocharis > subg. Eleocharis > sect. Eleocharis > ser. Eleocharis
Sibling taxa
E. acicularis, E. aestuum, E. albida, E. ambigens, E. atropurpurea, E. baldwinii, E. bella, E. bernardina, E. bicolor, E. bifida, E. bolanderi, E. brachycarpa, E. brittonii, E. cancellata, E. cellulosa, E. coloradoensis, E. compressa, E. cylindrica, E. decumbens, E. diandra, E. elliptica, E. elongata, E. engelmannii, E. equisetoides, E. erythropoda, E. fallax, E. flavescens, E. geniculata, E. intermedia, E. interstincta, E. kamtschatica, E. lanceolata, E. macrostachya, E. mamillata, E. melanocarpa, E. microcarpa, E. minima, E. montana, E. montevidensis, E. nana, E. nigrescens, E. nitida, E. obtusa, E. obtusetrigona, E. occulta, E. ovata, E. pachycarpa, E. palustris, E. parishii, E. parvula, E. quinqueflora, E. radicans, E. ravenelii, E. retroflexa, E. reverchonii, E. robbinsii, E. rostellata, E. suksdorfiana, E. tenuis, E. torticulmis, E. tortilis, E. tricostata, E. tuberculosa, E. uniglumis, E. vivipara, E. wolfii
E. acicularis, E. aestuum, E. albida, E. ambigens, E. atropurpurea, E. baldwinii, E. bella, E. bernardina, E. bicolor, E. bifida, E. bolanderi, E. brachycarpa, E. brittonii, E. cancellata, E. cellulosa, E. coloradoensis, E. compressa, E. cylindrica, E. decumbens, E. diandra, E. elliptica, E. elongata, E. engelmannii, E. equisetoides, E. erythropoda, E. fallax, E. flavescens, E. geniculata, E. intermedia, E. interstincta, E. lanceolata, E. macrostachya, E. mamillata, E. melanocarpa, E. microcarpa, E. minima, E. montana, E. montevidensis, E. nana, E. nigrescens, E. nitida, E. obtusa, E. obtusetrigona, E. occulta, E. ovata, E. pachycarpa, E. palustris, E. parishii, E. parvula, E. quadrangulata, E. quinqueflora, E. radicans, E. ravenelii, E. retroflexa, E. reverchonii, E. robbinsii, E. rostellata, E. suksdorfiana, E. tenuis, E. torticulmis, E. tortilis, E. tricostata, E. tuberculosa, E. uniglumis, E. vivipara, E. wolfii
Synonyms Scirpus quadrangulatus, E. quadrangulata var. crassior, Scirpus albomarginatus, Scirpus marginatus Scirpus kamtschaticus
Name authority (Michaux) Roemer & Schultes: in J. J. Roemer et al., Syst. Veg. 2: 155. (1817) (C. A. Meyer) Komarov: Fl. Kamtschatka 1: 207. (1927)
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