Phlegmariurus reflexus ­­­ (Lamarck) B. Øllgaard 2012

2. Phlegmariurus reflexus ­­­ (Lamarck) B. Øllgaard (2012a: 481). Fig. 11A

Lycopodium reflexum Lamarck (1792: 653).— Plananthus reflexus (Lamarck) Palisot de Beauvois (1805: 100).— Urostachys reflexus (Lamarck) Herter (1922: 249).— Huperzia reflexa (Lamarck) Trevisan (1874: 248). Type:— MARTINIQUE: herb. Lamarck, Comm. Joseph Martin s. n. (P holotype).

Lycopodium bifidum Willdenow (1810: 53).— Lycopodium reflexum Lamarck var. majus Spring (1842: 26).— Huperzia reflexa (Lamarck) Trevisan var. bifida (Willdenow) Trevisan (1874: 248).— Urostachys bifidus (Willdenow) Nessel (1939: 110).— Huperzia bifida (Willdenow) Holub (1985: 71). Type:— VENEZUELA: Cuchilla de Guajana Guajana, Humboldt & Bonpland 474 (B-Willdenow 19421 holotype; P-Humboldt isotype).

Lycopodium reversum Presl (1825: 82). — Urostachys reversus (Presl) Herter (1949a: 78). Type:— ECUADOR: Guayaquil, Haenke s. n. (PRC holotype).

Lycopodium reflexum Lamarck var. densifolium Baker (1887: 11).— Lycopodium densifolium (Baker) Underwood & Lloyd (1906: 106).— Urostachys reflexus (Lamarck) Herter var. densifolius (Baker) Nessel (1927: 394).— Urostachys densifolius (Underwood & Lloyd) Herter (1949a: 58).—Type:— COLOMBIA: Popayan, Hartweg 1480 (G, K); Moritz 2266 (= 226b: HBG, K); BRAZIL: Glaziou 15797 (K); syntypes.

Lycopodium pearcei Baker (1887: 14).— Urostachys pearcei (Baker) Nessel (1939: 73).— Huperzia pearcei (Baker) Holub (1985: 75). Type:— BOLIVIA: Santa Cruz, Feb. 1864 (K holotype; BONN-herb. Nessel 116 isotype).

Urostachys jergii Nessel (1935: 70). Type:— PUERTO RICO: Sintenis 6346 (BONN-Nessel 229 holotype; NY isotype).

Lycopodium mexiae Copeland (1941: 294).— Huperzia mexiae (Copeland) Rolleri and Deferrari (1988: 156). Type:— PERU: Huánuco, Churubamba, trail Cotirarda–Mercedes, 1875 m, Mexia 8193 a (UC holotype; F, GH, K, MICH isotypes).

Urostachys leptodon Herter (1954: 120–121).— Lycopodium leptodon (Herter) Morton (1964: 72).— Huperzia leptodon (Herter) Rolleri & Deferrari (1988: 156). Type:— COLOMBIA: Guasca, Bro. Ariste-Joseph A132 (US 888340 holotype).

Urostachys stellae-polaris Herter (1954: 121).— Lycopodium stellae-polaris (Herter) Morton (1964: 72). Type:— COLOMBIA: Cundinamarca, Guayabetal to Monte Redondo, SE of Quetame, 1300–1500 m, Pennell 1801 (US holotype; GH, NY isotypes).

Lycopodium spongiosum Rolleri (1985: 6).— Huperzia spongiosa (Rolleri) Rolleri & Deferrari (1988: 157). Type:— COLOMBIA: Cundinamarca, Chocontá–Guateque highway, 2900 m, Haught 5906 (US holotype, COL, F, NY, S isotypes)

Plants erect or ascending from a decumbent base, soft, usually forming small loose clumps, 10–30 (–40) cm tall. Shoots homophyllous, almost equally thick throughout, 7–15 mm in diameter including leaves. Stems excluding leaves 1.5–3 (–4) mm thick at base, sometimes tapering to 1–1.5 mm thick, ridged by decurrent leaves or almost smooth, sporangiate from 2–12 cm above the base and upward, usually 2–5 times dichotomous. Leaves borne in alternating, often irregular and oblique, whorls of (6–) 7–8 (–9), these 0.7–2.5 mm apart, forming 12–18 longitudinal ranks, ascending to spreading or sharply reflexed, straight to strongly recurved, linear-subulate, widest just above the base, 4–8 × 0.5–1 (–1.2) mm, softly herbaceous to subcoriaceous, adaxially convex, or concave near the base, abaxially flat, or slightly concave to convex, with obscure to somewhat prominent vein, with flat to revolute, very sparsely to densely denticulate to short-ciliolate margins. Leaf bases often somewhat decurrent. Sporangia 1–1.5 mm wide.

­­­ Distribution: Throughout humid mountainous regions of Tropical America.

­­­ Habitats: Terrestrial, erect or ascending herbs. Commonly pioneers on road banks and other open, humid situations, generally in montane forest, 370–3400 m elev.

­­­ Notes: Phlegmariurus reflexus is widespread in the Neotropics. It is a frequent pioneer on moist, disturbed ground, but may also occur in peat bogs, or even, by chance, as a low epiphyte. It is related to P. eversus.

Phlegmariurus reflexus is highly variable with respect to leaf and stem size, direction and crowding of leaves, and leaf margin characters. Part of the variation may reflect growth conditions, but often several different forms may be found growing intermixed in the same habitat, indicating that genetic differences exist. The variation patterns are complex and in need of a detailed study. The different morphological forms of P. reflexus, as well as other closely related species, often occur together on large road banks or landslides, and here may be compared under uniform growth conditions. In such populations the individuals may exhibit subtle or obvious differences. Neighbouring and more distant populations may exhibit similar variation and thus taxonomic recognition may be indicated. However, often the differences are not matched by other individuals, neither in the same nor in other populations, and therefore the plants may have the character of individual aberrants. Some of these are intermediate between other forms and may be hybrids, without exhibiting hybrid features such as abortive spores or irregular meioses.

The following variety of small dimensions stands somewhat apart, and was earlier recognized taxonomically, but this idea has been abandoned because it is an exaggerated simplification of the variation.