Equisetum diffusum D. Don, Prod. Fl. Nepalensis 19. 1825.

E. diffusum var. paucidentatum Page, Fern Gaz. 11: 31. 1974.
E. mekongense Page, Fern Gaz. 11: 36. 1974.

Description:

Stems homophyadic, 24-65 cm tall (m 41) with internodes 1.7-5.0 cm long (m 3.3) and 1.5-2.5 mm in diameter (m 1.8), having 6-10 ridges (m 8.3), these being grooved to flat-topped. Internally there is only carinal collenchyma, and the chlorenchyma is continuous around the stem. The central canal is absent to obscure, with only partial breakdown of the pith tissue. Endodermis outer common.

Sheaths urceolate, 4-7 mm long (m 5), 2.0-4.5 mm wide (m 3.4), with teeth 2-4 mm long (m 3.1), narrow, and dark, the stem ridge grooves continuing up the center of each sheath segment and to the end of the teeth.

Branches ascending, in regular whorls, or irregular, or sometimes absent on coniferous stems, with the first internode about equaling the subtending stem sheath, 3-7 mm long (m 4.8). Ridges 4, these prominently grooved and bearing a silica profile of 2 rows of closely set, blocky swateeth. Sheath segments grooved, the grooves continuing into the triangular, narrowly pointed teeth, commissure to 0.6 mm long, with prominent furrow of anchor cells. Valleys rounded, with stomata scattered throughout. Silica pilules covering stomata and distinctly outlining them. Stoma lined with a single row of somewhat rod-like pilules. Mamillae transversely aligned, usually distinct. Branches solid.

Cones 25-60 mm long (m 39.5), narrow, on peduncles 20-30 mm long (m 25.3).

Rhizome shiny, brown, covered with hairs, occasionally bearing tubers.

Spores 30-41 µm in diameter (m 34).

Gametophytes unknown.

Type:

Collected by Wallich in Nepal before 1825. There are in BM two specimens collected by Wallich in Nepal, one with the year 1820, the other with the number 398. Since that number is from his List of Plants, it is probably not a collection number. Hauke (1978) designates the specimen collected by Wallich in Nepal in 1820 (BM!) as lectotype for E. diffusum.

Seasonality:

Coning July to November.

Distribution:

Himalaya mountains from Simla (India) to Tibet, Sikkim, Burma, Assam, Bhutan, Nepal, and adjacent China (Kweichow, Kwangsi, Szechuan, Yunnan). 450 to 3500 m altitude.

Ecology:

Open but shady, wet places; stream sides, mossy falls, wet meadows, bogs, seepage slopes.

Discussion:

This species, endemic in the Himalayas, is unlike all but E. bogotense in that the stems are seen above ground all year round. Coning is generally confined to the latter half of the year, but occasional coning specimens have been found at other times (Yunnan, China, 16 April 1915. Forrest 6935 [BM]; Garhwal, India. 18 June, 1902. Inayat 26049 [DD]). Hauke visited a population near Simla on 24 February 1974 and could find no coning specimens, but did find some unbranched stems with dried cones still hanging on tht top. Also at that time Hauke found a tuber producing a new plant. (This confirms the report of Mehra and Bir, 1959, of tubers in E. diffusum). Kingdon Ward, on the label of a specimen collected in North Burma on 30 Jan. 1953 (#20453, BM) notes "the new plants are just starting again". Thus it would appear that there is an annual cycle, but the plants are not deciduous. Page (1974) recognized a variety, E. diffusum var. paucidentatum, on the basis of branch internode ridges "lacking both the longitudinal groove and ridge tubercles, or nearly so", and cites specimens from Simla and Garhwal, India, and from Nepal. I saw similar variation in specimens from Kweichow, China, and occasionally on other specimens with the main axis poorly developed and the branches larger than normal (when the main axis aborts, branches assume some stem characters in other species of Equisetum). In a genus notorious for its morphological plasticity, a variable character such as extent of development of ridge silica (and consequent emphasis of the groove) does not seem a sufficient basis for the recognition of a taxonomic variety. A similar variation in silicification is found in E. sylvaticum, and E. pratense.

Page (1974) described a new species of horsetail, E. mekongense, with vegetative stems like E. diffusum but heterophyadic like E. arvense. Micromorphologically it is indistinguishable from E. diffusum. On p. 41-42, after describing its micromorphology, which resembles that of E. diffusum, he says "The combination of these micromorphological features with the presence of two-keeled sheath teeth allow E. mekongense to be clearly distinguished from both E. arvense and E. diffusum even in the absence of information on the type of fertile shoot." Since E. diffusum has two-keeled sheath teeth (and a commis sure with anchorcells just like that in Page's Fig. 9e) Hauke is unaware of any character other than the fertile shoot by which E. mekongense differs from E. diffusum. The description of E. mekongense on p. 36 primarily emphasizes the heterophyadic character, and Page says (p. 37), "I have seen only one collection of this unusual species, which superficially gives the appearance of young vegetative shoots of E. diffusum connected to ephemeral spring shoots of the E. arvense type." Page earlier (1972a) had downgraded the importance of heterophyadic vs. homophyadic shoots as a taxonomic character at the sectional level in Equisetum. Hauke thinks it loses its significance here as well in light of several observations. 1) there was recently reported by Van Hoek (1976) the occurrence of heterophyadic shoots in E. fluviatile, apparently resulting from disturbance related to transplantation. 2) E. pratense normally shows considerable variation in the degree of dimorphism of its coniferous stems - at times they are little different from the vegetative other than being unbranched, at other times they are quite distinctive. 3) The sheaths immediately below the cone of E. diffusum are commonly enlarged, a character associated with coning stems in heterophyadic species. 4) The coniferous stems of E. mekongense are described as "ephemeral", but since this species is known only from the type specimen, the persistence of the coniferous stems cannot be determined. Coniferous stems of E. sylvaticum collected when young also appear "ephemeral", but after the spores are shed they become green, branched, and persistent.

Equisetum mekongense Page appears to me to be an example of not only one character taxonomy, but also one specimen taxonomy. Hauke believes that the type specimen of E. mekongense (on the basis of Page's description, photograph, and a general knowledge of the genus Equisetum) is merely an aberrant specimen of E. diffusum and so place it in synonymy. (The only specimen of E. diffusum Hauke saw coning in April was also from Yunnan, the month and region of collection of E. mekongense).

Hauke observed that specimens of E. diffusum from Kweichow, China, bear some resemblance to E. palustre. Page pointed out that a specimen of E. diffusum from Yunnan had micromorphology tending toward that of E. palustre.


HAUKE, R.L. (1978)
A taxonomic monograph of the genus Equisetum subgenus Equisetum.
Nova Hedwigia 30, p385.

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