Diphasiastrum thyoides (Willd.) Holub (1975: 108)

Øllgaard, Benjamin, 2020, Lycopodiaceae in Colombia: Subfamilies Lycopodioideae and Lycopodielloideae, Phytotaxa 433 (3), pp. 195-224 : 202-204

Diphasiastrum thyoides (Willd.) Holub (1975: 108) .— Fig. 2A .

Lycopodium thyoides Willdenow (1810: 18) .— Lycopodium complanatum L. var. tropicum Spring , in Martius (1840: 116).— Lycopodium complanatum L. var. adpressifolium Spring (1842: 102) .— Lycopodium complanatum L. var. thyoides (Willd.) Christ (1900: 42) . Type:— VENEZUELA: Silla de Caracas, Humboldt s. n. (B-Willd. 19352, holotype).

Lycopodium comptonioides Desvaux (1827: 185) excl. synonyms. Type:— BRAZIL: Locis montosis Brasilii, Herbier de A. N. Desvaux, Donnée par Mme Vve Lavallée en 1896. With Desvaux’s label and annotation (P holotype).

Lycopodium complanatum L. var. validum Weatherby (1910: 414) .— Syntypes (Fide Mickel & Beitel 1988):— MEXICO: Chiapas, Between Cristóbal Las Casas and Huistán & Seler 2273 (GH, K); Ghiesbrecht 600 (GH); Oaxaca, Cerro San Felipe, González & Conzatti 889 (GH); Veracruz, Region d’Orizaba, Bourgeau 3159 p. p. (GH, M, US); Hidalgo, Trinidad, Pringle 11856 (GH, L, MO).

Plants with creeping, trailing to scandent rhizomes, usually above ground, or hanging over banks. Rhizomes terete, 1.2–2.5 mm in diam. excl. leaves. Rhizome leaves relatively distant, borne in irregular spirals, or subverticillate, subulate, appressed to ascending. Aerial shoots arising from the rhizomes in a dorso-lateral position, ascending to erect, 10–50 cm tall, with vegetative portions to ca 30 cm tall. Main upright axis terete to somewhat flattened, bearing lateral, flattened, fan-shaped branchlet systems. Ultimate branchlets flattened, dorsiventral, anisophyllous, 1.5–3 mm wide incl. leaves, with trimorphic, decussate leaves in 4 ranks. Upper, median branchlet leaves with pointed, subulate to acicular, appressed, 1–2 mm long, free blades, and a conspicuous, ca 0.4–0.6 mm wide, prominently decurrent base. Lateral branclet leaves bilaterally compressed, long-decurrent, 2.5–7 mm long incl. bases, the free blades 1–3 mm long, appressed to spreading, acuminate to long-pointed, the leaf bases 0.6–1.5 mm wide, with almost parallel to distinctly diverging margins, often curved down. Ventral leaves inconspicuous, acicular, without decurrent base, 1–2 mm long. Peduncles terminating main erect axis or major axes of upper branchlet systems, 10–25 cm long, terete, with rather distant, subulate, appressed leaves, each bearing 4–9 pedicellate strobili. Strobili 1.5–5 cm long, 2–4 mm in diam. incl. sporophylls, often with protracted sterile tips. Sporophylls usually borne in alternating whorls of 3, forming 6 longitudinal ranks, subpeltate, with a basiscopic, median wing on the stalk, with broadly deltoid-ovate, long-cuspidate, ca 2–3 mm long, 1.5–2 mm wide exterior face, with almost entire, broadly membranous margins. Sporangia 1.5–2 mm in diam., with side walls of epidermis cells evenly sinuate. Spores densely reticulate on all faces.

Distribution: Throughout moist mountainous regions of Tropical America, south to northern and central Argentina.

Habitats: Clearings, road banks, open habitats, and secondary scrub in upper montane forest, and in the lowest páramos, alt. 1700–3700 m.

Notes: The present application of the name Diphasiastrum thyoides follows the “ L. thyoides -complex” of Wilce (1965: 155–158). I have not attempted to treat the infraspecific variation. The Colombian material referred to this species is variable and may belong to more than one taxon, but also external factors apparently affect the growth habit of the individuals.

Antioquia: Alto Capiro, above Sonsón–Abejorral road, 2800 m, Ewan 15737 (NO, S, UC). Mun. Belmira, vereda El Yerbal, road to páramo, 75°40’W 6°40’N, 2730–2795 m, Gómez et al. 640 (AAU). Arauca: Sierra Nevada de Cocuy, Quebrada el Playon, 3100 m, Cleef 10176 (U). Boyacá: Cord. Oriental, Nevado del Cocuy, Valle de Cocuy, 3400–3500 m, Cuatrecasas 1649 (SI). Subpáramo 5 km NW of Belén, 3410 m, Cleef 2231 (AAU). Vado Hondo, Valle de Río Cusiana, 2915 m, Cleef et al. 9234A (U). Vereda La Buitrera, road Sogamoso–Pajarito, beyond crossroad to Pueblo Viejo, 3350 m, Jaramillo et al. 2836 (AAU, K). Caldas: Laguneta, Salento, 2800 m, Sneidern 3084 (AAU, MO, S). Cauca: Méndez, NE of Silvia, 3000–3100 m, Haught 5099 (G). El Tambo, 2800 m, Sneidern 2204 (G, S). El Tambo, 1700 m, Sneidern 1381 (K, S). W of Tambo, 1800 m, Haught 5273 (S). Chisquio, 1700 m, Asplund 10647 (S). César: Mun. Pueblo Bello, Nabusimake, first bigger ridge above valley between Nabusimake and Pantano, in direction Duitama. 2850 m, Stancik 164 (AAU). Cundinamarca: Cord. Bogotá, 2700 m, 1866, Triana s.n. (G). San Cristóbal, near Bogotá, April 1905, Apollinaire s. n. (E, G). Mont. E. E. prés Bogotá, 2800 m, 29 June 1876, André 973 (K, NY). Mun. Carupa, road to La Peña de Sumaga, 3200 m, Acosta-Arteaga et al. 565 (GH, MO, VEN). Zipaquirá–Pacho, Páramo de Guerrero, 3100 m, Murillo & Villareal 1355 (AAU). Páramo de Palacio, W flank, km 8 along road, 3230 m, Cleef 3646 (AAU). Páramo de Cruz Verde, Bogotá–Choachí, km 11.2, 3275 m, Cleef 3331 (AAU). Neusa Dam, Cordillera de La Senda, Río Cubillos Valley 1 km SE of Puente del Borracho, 3150 m, Cleef 6643 (AAU). Páramo de Sumapaz, near Lagunitas S of San Juan, Los Saladeros, 3555 m, Cleef 8414 (AAU). Mun. Guasca, Páramo de Guasca, 2730 m, García Barriga 6239 (AAU). Montserrate, above Bogotá, along high ridge of escarpment above Valle de Bogotá, 3140 m, Ewan 16897 (NO, UC). Above Usaquén, Alston 7468 (AAU, BM). Usaquén, N of Bogotá, 2560 m, Moore et al. 9855 (VEN). Sabana de Bogotá, May 1923, Pring 146 (AAU, MO). Páramo Guasco, near Bogotá, Maguire & Maguire 44024 (K, VEN). 30 km E of Bogotá, 3000 m, Schaek s. n. (BR). Mun. Villapinzón, Páramo La Calavera, 73°33’W 5°12’N, 3200 m, Stancik 418 (AAU). Guadalupe, Bogotá, Haught 5003 (S). 7 km NW of Zipaquirá, 3100–3200 m, Tryon & Tryon 6065 (K, S, UC). Guajira: Sierra Nevada de Santa Marta , Río San Miguel, 3100–3700 m, Cuadros & Gentry 2737 (AAU). Magdalena: Sierra de Perijá, Laguna de Juncos, 2200 m, Haught 4500 (S). Sierra Nevada de Santa Marta, 2400 m, Santa & Escobar 660 (AAU, QCA). Sierra Nevada de Santa Marta , road to crest of San Lorenzo, 2400 m, Santa & Escobar 638 (AAU). Yerbabuena–Cebolleta, 2000–2600 m, Romero Castañeda 7082 (AAU). Nariño: Mountains of Pasto, before 1846, Jameson 430 (E, G). Norte de Santantander/ Cesar: Cerro de Oroque, entre Abrego y Las Jurisdicciones, 3440–3750 m, García-Barriga & Jaramillo 19903 (U). Cerro Tablón, above Las Mesas on W slope of Volcán Doña Juana, 2700–2800 m, Ewan 16585 (NO). N slope of Volcán de Cumbal, via Laguna Bolsa, 2740–2980 m, Ewan 16141 (NO, S, UC). Norte de Santander: E slope of Páramo de Santurban, toward Mutiscua, 3600–3900 m, Killip & Smith 19605 (K, P). Santander: Ocaña, 2440–2740 m, 1853, Schlim 390 (G). Páramo de Las Puentes, above La Baja, 3500–3700 m, Killip & Smith 18209 (S). Valle: Cali–Loboquerro road, 2000–2195 m, Luteyn & Lebron-Luteyn 6981 (AAU). Hoya del Río Cali, Pichindé: Morro Pelado, 2300 m, Cuatrecasas 18152 (S).

Diphasium C. Presl ex Rothmaler (1944: 64) . Type:— Diphasium jussiaei (Poir.) Rothm. (= Lycopodium jussiaei Poir. ).

Lycopodium sect. Diphasium (C. Presl ex Rothm.) Øllgaard (1987: 172) ; basionym: Diphasium C. Presl ex Rothm. — Diphasium ser. Jussieua Rothmaler (1944: 65) .

Sporophytes with subterranean or creeping to subscandent main stems; branchlets dorsiventral, anisophyllous, with dimorphic leaves, two dorsolateral ranks of broad alternate leaves, and two or three ventral ranks of narrow membranous-tipped leaves; strobili pedunculate or sessile; sporophylls subpeltate; sporangial epidermal cells with sinuate, finely curled side walls; spores reticulate with large meshes and unornamented proximal faces; gametophytes obconic.

The genus Diphasium , with possibly 5 species, is known from the Greater Antilles, Central America, the Andes from Venezuela to Argentina and Chile, Brazil, the Phillipines, Australia, New Zealand and Tasmania.

Figures

FIGURE 2. A: Diphasiastrum thyoides (Ecuador: Holm-Nielsen et al. 4818, AAU), branchlets seen from above (left) and below (right). B: Diphasium jussiaei (Ecuador: Holm-Nielsen et al. 4670, AAU) branchlets seen from (left) and below (right). C: Lycopodium clavatum ssp. clavatum (Ecuador. Harling 4091, S). C1: Branchlet leaf. C2: Sporophyll. D: Lycopodium clavatum ssp. contiguum (Ecuador: Cazalet & Pennington 5409, US). D1: Sporophylls.