Diphasium jussiaei (Desv. ex Poiret) Presl ex Rothmaler (1944: 65)

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

Diphasium jussiaei (Desv. ex Poiret) Presl ex Rothmaler (1944: 65) [as jussieui].— Fig. 2B

Lycopodium jussiaei Desv. ex Poiret (1814: 543) .— Lycopodium scariosum Forst. var. jussieui (Desv. ex Poiret) Baker (1887: 29) . Type:— PERU: Perou, Herb. Jussieu 658 (P holotype!).

Lycopodium jussiaei Poiret var. microphyllum Poiret (1814: 543) . Type:—Amérique meridionale, Herb. Bonpland (P holotype!).

Lycopodium haenkei Presl (1825: 78) . Type :— PERU: In montosis Peruviae ad Huanocco, Haenke s. n. (holotype, PRC!; possible isotype, K!).

Lycopodium heterophyllum Sprengel (1827: 13) .— Lycopodium lindseaceum Spring (1838: 180) . Type:— PERU: Peruvia, Humboldt s. n. (holotype, B, Herb. Willd. 19425!).

Lycopodium holtonii Underwood & Lloyd (1906: 123) . Type :— COLOMBIA: In montibus juxta Bogotam, Holton 79 (holotype, NY; isotype, K!).

Lycopodium scariosum Forst. var. ramosissimum Nessel (1940: 172) .— Lycopodium canceris Herter (1949: 8) . Type:— JAMAICA: Bancroft 2420 c (holotype, BONN, Herb. Nessel!).

Plants creeping, trailing or hanging over banks. Main stems usually above ground, rooting at intervals, rigid, 2–4 mm thick, usually above ground. Leaves of main stems radially arranged, uniform, 3–5 mm long, ca 1 mm wide, linear-lanceolate, with broadly membranous upper part and apex (sometimes scarious), irregularly obtuse to acute, erose apex. Aerial shoot systems 5–75 cm tall, branched almost from the base, in large individuals with a main axis almost conform to the main stem, upward gradually changing to anisophyllous, bearing alternating, fan shaped branchlet systems. Branchlets dorsiventral, anisophyllous, flattened, 4–6 (–8) mm wide incl. leaves, with 2 dorso-lateral ranks of broad leaves and 2—3 indistinct ventral ranks of narrow leaves. Dorsolateral leaves flattened in the plane of the branchlet system, or inclinate to it, obliquely elliptic, acroscopic margin 2–3.5 × 1–1.5 mm (middle), upward and ventrally curved, subacute to mucronate to short hair-tipped, basiscopic margin short to long decurrent. Ventral leaves appressed, lanceolate-subulate, with membranous apices. Strobili terminating main aerial axes, or stronger branches, pedunculate or rarely sessile. Peduncles to 12 cm long (Brazilian material, longer elsewhere), simple or to twice forked, bearing 1–3 strobili, radially symmetrical, with distant, narrow membranous-tipped, appressed leaves. Strobili (1–) 3– 10 cm long, 4–6 mm in diam. incl. leaves. Sporophylls borne in alternating whorls of 4, forming 8 longitudinal ranks, subpeltate, with a basiscopic, median membranaceous wing on the stalk, with ovate, more or less acuminate exterior face, 4–6 mm long, ca 2 mm wide, with narrowly membranous, shallowly erose denticulate margins. Sporangia 1.5–2 mm in diam. Spores reticulate, with large, regular meshes on distal faces, and unornamented proximal faces.

Distribution: Jamaica, Hispaniola (fide Proctor 1985), Costa Rica, Panamá, Venezuela, the Andes south to Bolivia, Brazil (Itatiaia and Serra do Mar).

Habitats: Usually a vigorous scrambling to scandent plant, common in clearings and on road banks and open places in montane forest, 1700–4000 m elev. In relatively dry habitats the plants are smaller and sparsely branched. A highly variable species, apparently due to external factors.

Notes: Diphasium jussiaei is related to D. scariosum (G. Forster) Rothmaler (1944: 65) ( Indonesia to New Zealand), and to plants from Southern Argentina, Chile and Juan Fernandez recognized as D. gayanum (Remy) Holub (1985: 78) . The plants referred to Diphasium jussiaei are generally larger and coarser than those species. A modern revision of the group is desirable.

Dept. unknown: Mutis 3353 (K). Antioquia: Mun. Urrao, trail to Páramo de Frontino, 76°10’W 6°30’N, McPherson 13202 (AAU). Mun. Belmira, old road Belmira–Sta Rosa, 2900–3000 m, Arbelaez et al. 143 (AAU). And. Central Antioquia, 3000–4000 m, Mayor 43 (S). Páramo de Sonsón, above Sonsón, 2743–2930 m, Ewan 15704 (NO). Boyacá: Vado Hondo, Valle Río Cusiana, 2915 m, Cleef et al. 9234 (U). Road Sogamoso–Pajarito, near Badhondo, 2200 m, Bischler 1805 (G). Road Sogamoso–Pajarito, Río Cusiana, 2000 m, Bischler 1735 (G). Mun. Sutamarchan, Serranía Marchan, subparamo, 3000 m, Stancik 1110 (AAU). Caldas: Nevado del Ruiz, road Manizales–Hotel Termales, 3110 m, Santa & Hoyos 680 (AAU). Cauca: Prov. de Popoyan, 1844, Hartweg 1475 (G, K). Macizo Colombiano, Páramo de las Papas, between Boquerón and La Hoyola, 3200–3510 m, Idrobo et al. 3848 (AAU, G). Road Totoró–Inzá, W slope of Cord Central, towards paramo de Las Delicias, 3100 m, Santa et al. 975 (AAU). Puracé, road to Parque Nacional Puracé, 3250 m, Santa & Hoyos 720 (AAU, HUA). Volcán Puracé, Laguna de San Rafael. 3450 m, Cleef 2653 (AAU). El Tambo, 3060 m, Sneidern 1126 (S), 2198 (G). Cundinamarca: Bogotá, Sprague 182 (K). Páramo de la Cruz Verde, Alto de La Viga, 74°W 4°35’N, 3460 m, Cleef 2828 (U). Torca, 2630 m, Cleef 4 (AAU). Road Bogotá –La Galera, 6 km from Bogotá, 2900 m, Bischler 1260 (G). Páramo de Guasca, 3100 m, Bischler 1165 (G). Andes near Bogotá, 3000 m, Apollinaire s. n. (E). Zipaquirá, Alto del Chamicero, 3200 m, Santa 1021 (AAU). Cord. de Bogotá, 2700 m, Triana s. n. (G). Huila: Pitalito–Mocoa road, km 28–32, 2100–2240 m, Luteyn et al. 7558 (AAU, NY). Nariño: Pasto, 1846, Jameson 482 (E, G, K). Norte de Santander: Páramo de Fontibon, Pamplona, 2400 m, Alston 7284 (BM). Tolima: Ibagué, trail Juntas– Nevado del Tolima, 2850 m, Santa et al. 967 (AAU). Valle: Road Ansermanuevo– San José del Palmar, km 60–62, 1900 m, Luteyn et al. 7282 (AAU, NY).

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.