Phlegmariurus crassus (Willd.) B. Øllgaard (2012b: 13)

Øllgaard, Benjamin & Testo, Weston, 2021, The Lycopodiaceae of Panamá, Phytotaxa 526 (1), pp. 1-66 : 14-15

. Phlegmariurus crassus (Willd.) B. Øllgaard (2012b: 13) .—Fig. 8

Lycopodium crassum Willdenow (1810: 50) .— Urostachys crassus (Willd.) Nessel (1939: 75) .— Huperzia crassa (Willd.) Rothmaler (1944: 60) . Type:— ECUADOR: Antisana, Humboldt & Bonpland 2263 (B-Willd. no. 19417 holotype; isotypes BM, P).

Lycopodium erythraeum Spring (1849: 7) .— Huperzia erythraea (Spring) Trevisan (1874: 249) .— Urostachys erythraeus (Spring) Nessel (1939: 74) . Type:— ECUADOR: Antisana, Hartweg 1472 (lectotype K, designated by Øllgaard (1988: 54); BM, G, GL, P isotypes).

Urostachys crassus (Willd.) Nessel var. herteri Nessel (1934: 180) . Type:—The lectotype should probably be chosen among the specimens in BONN [Herb. Nessel 123], all of which belong to Phlegmariurus crassus . See Øllgaard (1988: 55) for a comment.

Urostachys bonae-voluntatis Herter (1954: 112) .— Lycopodium bonae-voluntatis (Herter) Morton (1964: 72) . Type:— ECUADOR: Mojanda, Mille 95 ( US holotype).

Urostachys pilgerianus Nessel (1940 a: 161) .— Huperzia pilgeriana (Nessel) Holub (1985: 75) . Type:— PERU: Cordillere, 1901, Rautenstock et Mann s.n. (BONN [Herb. Nessel 124] holotype). See Øllgaard (1994: 34) for discussion of the type.

Urostachys springii Nessel (1940 a: 161) .— Huperzia springii (Nessel) Holub (1985: 77) . Type:— ECUADOR: Ecuador, Jameson 103 (BONN [Herb. Nessel 125 lectotype], designated by Øllgaard 1988: 54). The lectotype is different from specimens in G and OXF with the same collection number.

Illustrations: — Øllgaard (1988: Fig. 8C-D).

Plants erect from a prostrate to ascending base, forming loose to dense clumps with basal, prostrate-ascending, rooting shoots (these often absent in collections), and erect, finger-like aerial shoots, at least to 40 cm tall, at least to 7 times dichotomous. Erect shoots homophyllous or almost so, equally thick throughout, or slightly tapering upward, 7–14 mm in diameter including leaves, sporangiate from 5–20 cm above the base. Stems excluding leaves (2–) 3–5 mm thick at the base, sometimes tapering to 2–3 mm thick, usually completely concealed by leaves, except in shaded divisions or individuals. Leaves uniform throughout, or slightly reduced upward, borne in often irregular, alternating whorls of 4–7, these 1–3(–4) mm apart, forming 8–14 obscure or oblique to regular longitudinal ranks, densely crowded, usually closely imbricate, straight (or slightly upward curved in shaded divisions), linear-lanceolate to lanceolate, or triangular-lanceolate, narrowly to broadly acute, (6–)7–11 × (1.5–) 2–2.5 mm, upward sometimes reduced to 6–8 mm long with a prominent, short to long decurrent basal swelling (air sac) causing a bulging or perpendicular appearance of the very leaf base in sporangiate leaves, amphistomatic, adaxially concave to slightly convex, with a raised veinal ridge near the base, abaxially convex and rounded or with a prominent veinal ridge, slightly to strongly rugose by protruding, blister-like epidermal cells, with smooth to slightly rugose, usually sclerified margins, green to brick red or dark red, usually strongly pruinous. Sporangia 1.5–2.5 mm wide.

Habitats:— Páramos, 3000–3300 m elev.

­­­ Distribution:— Costa Rica to the Andes of Venezuela south to Bolivia.

­­­ Notes:—As here delimited, Phlegmariurus crassus is a widely distributed polymorphic species. Some of the Central Americal plants may merit status as a distinct taxon. Specimens from Mexico, Guatemala, and Hispaniola commonly referred this name apparently are a different species.

This species differs from Phlegmariurus talamancanus by the heteroblastic shoots, the usually rugose abaxial leaf epidermis, by the smaller and more appressed, pruinose leaves. In Central America it also generally has narrower shoots than P. talamancanus .

­­­ Specimens­­­examined:—Puntarenas/Bocas­­­del­­­Toro: ­­­ Cordillera de Talamanca, Cerro Echandí, on international border, 3160 m, Davidse et al. 23848 (AAU, MO). ­­­ Páramo of Cerro Echandi, CR/ Panamá borderline, 3000–3162 m, Gómez et al. 22676 (MO). Cerro Fábrega, Parque Nacional La Amistad, scrub paramo, 3250 m, Klitgaard et al. 822 (AAU, PMA). ­­­ Cerro Fabrega and ridge to the North, 14 km NE (straight line) of Estación Pittier, 3300 m, Monro & Knapp 5159 (PMA). Cordillera de Talamanca, Cerro Fábrega, main peak of Fábrega Massif, 3100–3300 m, Davidse et al. 25289 (AAU, F, MO). ­­­ Between Bine and Itamut peaks, Fábrega Massif, 3200 m, Gómez et al. 22565 (AAU).