Flora of North America species comparison

Eleocharis palustris is the most widespread and common species of the extremely difficult circumboreal “E. palustris complex,” which in North America comprises E. palustris, E. mamillata, E. macrostachya, E. erythropoda, E. uniglumis, E. kamtschatica, and E. ambigens. Two or more of these species have been combined by recent authors. The complex has been studied extensively only in northern Europe (S.-O. Strandhede 1965, 1966), where E. palustris, E. mamillata, and E. uniglumis are recognized (S.-O. Strandhede 1966). European studies and preliminary studies in North America by S.-O. Strandhede (1967) and L. J. Harms (1968) indicate that unstable chromosome structure and number as well as interspecific hybridization contribute to the taxonomic complexity of the E. palustris complex.

Eleocharis palustris is extremely variable worldwide. Recognition of infraspecific taxa outside northwestern Europe is premature. For northern Europe, S.-O. Strandhede (1966) recognized E. palustris subsp. palustris, with two varieties, for which the chromosome numbers 2n = (14–)16(–17) have been reported, and E. palustris subsp. vulgaris, without varieties, for which the chromosome numbers 2n = (33–)38–39(–40) have been reported. Eleocharis palustris subsp. vulgaris is morphologically intermediate between E. palustris and E. uniglumis and may be of hybrid origin. Its North American counterpart appears to be the polyploid populations of E. macrostachya (variants b and c, at least in part), as defined herein. For North America, S.-O. Strandhede (1967) and L. J. Harms (1968) recognized two “cytotypes” among the plants with the morphology of E. smallii, one with 2n = 16 (variant a below) and one with 2n = 36 (variant c below). Much of the variation in habit is undoubtedly because of modification of the phenotype by environmental conditions as described for Europe by S.-O. Strandhede (1966). The more robust plants are often emergent in open water and may be called “crassa” phenotypes; the more slender plants often grow in densely vegetated marshes and meadows and may be called “meadow” or “grassland” phenotypes. Intermediates between E. palustris variant b (below) and E. erythropoda are common in zones of sympatry.

At least 4 variants are notable in North America.

Variant a (Eleocharis smallii in the strict sense) has culms mostly 1–3 mm wide; distal leaf sheaths sometimes disintegrating, often splitting adaxially, summits often with red margins, apices obtuse to broadly acute; floral scales 3–4 mm; achenes to 1.5(–1.6) mm; culm stomates 39–48 µm (based on very few measurements). Reported chromosome numbers for which I have seen vouchers, all from Kansas, are 2n = 16, 17 (L. J. Harms 1968). The range of variant a is mostly northeastern, where it is known from elevations to 1700 m in Newfoundland, west to Manitoba and south to North Carolina, Kentucky, Missouri, and Kansas, with one collection from east-central Alaska.

Variant b is similar to variant a and intergrades with it. It has culms only 0.5–1.2 mm wide; distal leaf sheaths persistent and not splitting, summits usually with markedly red margins, markedly oblique when viewed from the side, apices acute to narrowly obtuse; and spikelets with proximal scale often clasping 3/4 of the culm. At least some of these slender plants may simply be meadow or grassland forms produced by the direct effects of unfavorable enviromental factors such as competition. Variant b is mostly sympatric with variant a; it is more common in the Southeast, where it is known south to Louisiana and Arkansas. Plants from the more southern part of the range are especially striking because of their extremely oblique, brightly red-margined sheath summits and proximal floral scales usually clasping to 3/4 of the culm.

Variant c may be called Eleocharis palustris var. vigens L. H. Bailey. The lectotype is from the shores of Lake Champlain in Vermont (S. G. Smith 2001). It is similar to unusually robust forms of variant a, from which it differs in that its achenes are 1.6–2 mm, culm stomates 52–65 µm, and floral scales mostly 3.5–4.5 mm. Because of its large achenes and stomates, variant c is assumed to be tetraploid with 2n = 36 (S.-O. Strandhede 1967; L. J. Harms 1968). Variant c apparently grows mostly as an emergent in open water to about 1 m deep. Its known range is northeastern, from Newfoundland and Labrador to Manitoba, south to New York, Michigan, Wisconsin, and Nebraska.

Variant d comprises most of the plants that cannot be placed in the preceding variants. Most of these plants closely resemble most specimens that I have seen from northern Eurasia and as described for Eleocharis palustris subsp. palustris by S.-O. Strandhede (1966). Variant d has distal leaf sheaths often splitting or disintegrating, the summit margins not reddish, and apices usually broadly obtuse. In North America variant d is mostly subarctic and boreal; it is known from Newfoundland and Labrador to Alaska, south to New York, Wisconsin, Minnesota, Iowa, New Mexico, and California. Some plants of variant d that have markedly narrow tubercles mostly much (to 2 times) higher than wide and narrow achenes only 0.9–1.1 mm wide may deserve taxonomic recognition; they are known from Manitoba west to British Columbia and Alaska, south to Colorado, Utah, and California. Specimens of variant d from scattered western localities from Alaska and Yukon south to California have floral scales 4–5 mm and achenes 1.6–1.9 mm and are very similar to variant c.

(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.)