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flat-stem spikerush, flat-stem spikesedge, unknown

Matted spikerush, mudflat spikesedge, éléocharide intermédiaire

Habit Plants perennial, mat-forming; rhizomes evident, short to rather long, 2–3 mm thick, firm, cortex persistent, internodes very short to 2 mm, scales often decaying to coarse fibers, 5–12 mm, papery. Plants annual, densely tufted; rhizomes present, inconspicuous, erect or ascending, 0.5 mm thick, internodes less than 2 mm, scales disintegrating, 3 mm, membranous-translucent.
Culms

subterete to greatly compressed, to 5 times as wide as thick, often with 2–12 ridges, 8–45 cm × (0.2–)0.5–1.8 mm, firm to hard, spongy.

often arching or recurved or reclining, their spikelets usually all fruiting simultaneously, long to very short in 1 tuft, subterete, 4–40 cm × 0.2–0.5 mm, soft.

Leaves

distal leaf sheaths persistent, not splitting, proximally red, distally green to stramineous, papery, apex usually red-brown, broadly obtuse to subtruncate, callose, tooth absent.

distal leaf sheaths persistent and evident or disintegrating (fugaceous) and not evident, proximally green or stramineous, brown, reddish, or purple, distally stramineous to green or colorless, papery to membranous, apex subacute to acuminate, sometimes with linear blade to 1 mm.

Spikelets

ovoid, 4–8 × 2–4 mm, apex acute;

proximal scale clasping 3/4 of culm to amplexicaulous, entire;

subproximal scale empty or with flower;

floral scales spreading in fruit, 20–60, 6 per mm of rachilla, medium brown to very dark brown, sometimes stramineous or colorless, midrib region often paler, ovate, 2–3(–4) × 1–1.5 mm, apex acute to acuminate, the apical colorless hyaline region mostly longer than wide and 0.6–1.2 mm, in proximal part of spikelet 2-fid to shallowly cut, rarely entire, in distal part often entire, carinate in distal part of spikelet.

basal spikelets absent (often some spikelets subsessile); never proliferous, ovoid, terete, 2–10 × 1–2 mm, apex acute;

proximal scale empty or with a flower (usually in the same plant), deciduous or some persistent, amplexicaulous, similar to other scales (except apex sometimes rounded);

subproximal scale with a flower;

floral scales spiraled, 5–30, 5–6 per mm of rachilla, pale brown to colorless, often spotted brown, ovate, 1.5–2 × 1 mm, thinly membranous to translucent, midribs often green, evident to obscure, apex acute.

Flowers

perianth bristles 0–5, stramineous, length variable, to equaling achene, obscurely to clearly retrorsely spinulose;

stamens 3;

anthers yellow- to orange-brown, 0.7–2 mm;

styles 3-fid or some 2-fid.

perianth bristles 6–7, pale brown to stramineous or whitish, slender, equaling to exceeding tubercles or rarely rudimentary to less than 1/2 achene length;

spinules sparse, retrorse, sharply acute;

stamens 3 (sometimes fewer?);

anthers 0.2–0.5 mm;

styles 3-fid.

Achenes

falling with scales or some persistent, yellow-brown or dark brown, obovoid to obpyriform, nearly equilaterally to compressed-trigonous or usually some biconvex, angles evident or obscure, 0.8–1.1 × 0.6–0.8 mm, neck very short, finely rugulose and sometimes finely (or coarsely) cancellate at 10–20X with 14–20 blunt horizontal ridges in a vertical series.

golden brown, narrowly obpyriform, proximally markedly narrowed, compressed-trigonous, angles slightly prominent, 0.9–1 × 0.6–0.75 mm, finely reticulate or pitted or papillose at 20–30X, apex usually constricted proximal to tubercle.

Tubercles

brown (to whitish), pyramidal, usually depressed, often rudimentary, 0.15–0.3 × 0.2–0.35 mm.

brown, narrowly pyramidal to linear, trigonous, 0.3–0.4 × 0.1–0.25 mm.

2n

= 22.

Eleocharis compressa

Eleocharis intermedia

Phenology Fruiting (spring–)summer–fall.
Habitat Fresh wet, often marly places along streams, lakeshores, tidal meadows, disturbed areas
Elevation 10–100 m (0–300 ft)
Distribution
from FNA
AL; AR; CO; DC; IA; IL; IN; KS; KY; MD; MI; MN; MO; MS; ND; NE; NM; NY; OH; OK; PA; SD; TN; TX; VA; WI; WV; BC; MB; ON; QC; SK
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from FNA
CT; IL; IN; MA; MD; ME; MI; MN; MT; NH; NJ; NY; OH; PA; TN; VA; VT; WI; WV; NB; ON; QC
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Discussion

Varieties 2 (2 in the flora).

Eleocharis compressa is extremely variable. After E. bifida and E. occulta are segregated from E. compressa (type from Ohio) and E. acutisquamata (type from Texas), the remaining plants intergrade completely and are best treated as varieties of E. compressa (S. G. Smith 2001). Because some plants of E. compressa approach E. elliptica some authors have treated E. compressa as a synonym of E. elliptica. Putative E. compressa × E. erythropoda hybrids from southeastern Ontario have been described (P. M. Catling 1994), and I have observed similar putative hybrids in the field in southeastern Wisconsin. Introgression from E. erythropoda is suggested by the presence in many plants of E. compressa of some 2-fid styles and biconvex achenes and some or rarely all entire floral scales, including the type. Some collections of E. compressa from the District of Columbia region and Tennessee have distinctly cancellate achenes as is common in E. tenuis var. verrucosa. The earlier name E. acuminata (Muhlenberg) Nees, name confused (H. K. Svenson 1932), is often given as a synonym of E. compressa (S. G. Smith 2001).

Both 2n = 24 and 2n = 26 cytotypes for E. compressa from south-central United States have been reported (L. J. Harms 1972). Fertile artificial hybrids between these two cytotypes were also reported, as well as artificial E. compressa (2n = 36) × E. elliptica (2n = 38) hybrids with meiotic pairing and pollen stainability more than 90 percent. I have not seen voucher specimens for those artificial hybrids or for any of the published chromosone number reports for E. compressa, except for a count of 2n = 24 (presumably made by L. J. Harms) that is given on a herbarium specimen label of E. compressa var. acutisquamata from Kansas (S. G. Smith 2001).

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

The flowers are mostly cleistogamous, with stigmas and stamens remaining enclosed in the floral scales except sometimes on the shortest culms. We have not seen specimens to confirm literature reports of E. intermedia from Iowa.

Eleocharis intermedia perhaps should be placed in sect. Intermediae (Svenson) Zinserling following I. Kukkonen (1990). H. K. Svenson (1929) placed it in his E. ser. Intermediae (name invalid) but later (1937, 1957) placed it in his E. subser. Truncatae (name invalid), and M. S. González-E. and P. M. Peterson (1997) placed it in E. subser. Truncatae. However, E. intermedia differs from all other members of ser. Eleocharis in its annual, tufted habit without evident, creeping rhizomes, culms varying in one tuft from long to very short, often with some spikelets subsessile, all spikelets of a tuft fruiting simultaneously, and the proximal scale often subtending a flower. It was placed in ser. Tenuissimae because of a resemblance to some other species of that series; closer observation reveals that it differs substantially in its achenes or habit from other members of ser. Tenuissimae. It rather closely resembles E. flavescens var. olivacea and E. geniculata in 8a2a. ser. Maculosae as well as small plants of species in 8a2b. ser. Ovatae.

Eleocharis macounii Fernald had been treated as a distinct species, or included within E. intermedia, E. obtusa, or E. ovata in the broad sense (including E. obtusa). P. M. Catling and S. G. Hay (1993) presented morphologic evidence suggesting that E. macounii Fernald is a hybrid between E. intermedia and E. obtusa, and named it E. ×macounii (Fernald) P. M. Catling & S. G. Hay. It is known only from the type collection from 1893 and a second collection from 1987, both from along the Gatineau River north of Ottawa in southwestern Quebec. Its habit is very similar to that of E. intermedia, from which it differs mainly in its achenes lenticular to trigonous in the same plant, the two or three angles prominent, keeled, costate, the surface smooth or very finely reticulate or pitted at 30X, and tubercles 0.2–0.5 mm wide. It produces few achenes, whereas E. intermedia and E. obtusa usually produce abundant achenes. In the protologue M. L. Fernald (1899) compared E. macounii with the Eurasian E. carniolica W. D. J. Koch. H. K. Svenson (1929, 1937) compared it with both E. intermedia and the Eurasian E. multicaulis (Smith) Smith and later (1957) placed it in synonymy under E. obtusa. Pending further research, it seems best to treat the two known collections of E. macounii as probable hybrids between E. intermedia and either E. obtusa or E. ovata; all three species are sympatric in the region where the collections of E. macounii were made (P. M. Catling and S. G. Hay 1993; B. M. H. Larson and P. M. Catling 1996).

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

Key
1. Culms compressed, 2–5 times as wide as thick, 0.5–1.8 mm wide.
var. compressa
1. Culms subterete to slightly compressed, not more than 2 times as wide as thick, (0.2–)0.5–1 mm wide.
var. acutisquamata
Source FNA vol. 23. FNA vol. 23, p. 96.
Parent taxa Cyperaceae > Eleocharis > subg. Eleocharis > sect. Eleocharis > ser. Eleocharis Cyperaceae > Eleocharis > subg. Eleocharis > sect. Eleocharis > ser. Tenuissimae
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. 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. 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
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. 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. 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
Subordinate taxa
E. compressa var. acutisquamata, E. compressa var. compressa
Synonyms E. elliptica var. compressa Scirpus intermedius, E. intermedia var. habereri, E. reclinata
Name authority Sullivant: Amer. J. Sci. Arts 42: 50. (1842) Schultes: Mant. 2: 91. (1824)
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