Cyathea lockwoodiana (Windisch) Lellinger (1984: 57)

Lehnert, Marcus, 2016, A synopsis of the exindusiate species of Cyathea (Cyatheaceae-Polypodiopsida) with bipinnate-pinnatifid or more complex fronds, with a revision of the C. lasiosora complex, Phytotaxa 243 (1), pp. 1-53 : 19-21

21. Cyathea lockwoodiana (Windisch) Lellinger (1984: 57) . ( Fig. 4 ). Sphaeropteris lockwoodiana Windisch (1976: 57) . Type:— COLOMBIA. Vaupés: Río Macaya (afluente Río Apaporis), Cerro Chiribiquete, 1700–2100 ft, 24 July 1943, R.E. Schultes 5635 (holotype GH-00022119–00022122!, isotypes US-00066158!/-00066159!).

Trunks to 2.5(–4.0) m tall, 5–8 cm diameter, covered with old petiole bases, due to these spiny, sometimes petioles bases rotting off in older trunk parts, revealing dark gray-brown frond scars 3.0–3.5 × 2.0– 2.5 cm; epidermis densely covered with brown, narrowly lanceolate scales similar to petiole scales; apices hidden between petiole bases; without adventitious buds. Fronds to 200 cm long, erect to arching. Petioles to 50–65 cm long, proximally aculeate with spines to 3 mm long, dark brown to dark purpureous, rarely blackish, weakly shiny, with dense scurf consisting of appressed reddish brown trichomidia and branched hairs, also some white, erect, lanceolate squamules to 2 mm long grading into multicellular hairs; petioles basally without lenticels, or not detectible in dried material. Petiole scales long-lanceolate, to 30.0 × 1.5–3.0 mm, relatively thin-textured, bases weakly cordate, basifix to pseudopeltately attached, straight to falcate, apices long attenuate, strongly undulate and twisted; scales concolorous, auburn to orange-brown, differentiated margins persistent, the cell rows not or only weakly exerted, ending in short teeth. Laminae to ca. 150 × 80 cm, bipinnate-pinnatifid, firm-chartaceous to subcoriaceous, matte, dark-green adaxially, often blackish when dried, dark olive-green abaxially; apices gradually reduced. Rhachises sparsely muricate in proximal parts, otherwise inermous, brown to dark purpureous abaxially and adaxially, with whitish to tan multicellular hairs to 2.0 mm long, antrorsely curved adaxially, abaxially spreading, persistent, leaving the epidermis rough if abraded; also with brown, narrowly lanceolate scales to 3.0 × 0.5 mm. Largest pinnae 40–55 cm long, subsessile or stalked to 1.0 cm, ca. 10– 12 pairs per frond, patent to ascending, alternate, inarticulate, distally narrowly green-alate, distal segments simply adnate before ending in a pinnatifid apical section; basal pinna pairs not much smaller than than the medial pinnae, weakly reflexed. Costae 2.0– 2.5 mm wide, inermous, dark ochre to brown or orange-brown abaxially, darker adaxially, with appressed, whitish to tan, multicellular hairs to 2.0 mm long adaxially, with scurf of appressed reddish brown trichomidia and branched hairs as well as spreading white uniseriate hairs to 1.5–2.0 mm long abaxially; junctures of costae and rhachises not swollen, abaxially often black when dried, each with an inconspicuous, planar to weakly protruding, elliptic aerophore, to 2 × 1 mm. Largest pinnules 70–100 × 13–17 mm, sessile to subsessile (stalked to 1 mm), inarticulate, 1.3–2.0(–2.2) cm between the stalks/costules, lanceolate to linear-oblong, truncate to cuneate basally, long-acute to attenuate apically with serrate to crenulate margins; costules dark carnose to atropurpureous adaxially and abaxially, proximally often darker; adaxially strongly prominent, ridged, and densely hairy with whitish to tan, spreading to appressed, multicellular hairs to 1.5 mm long, abaxially weakly to strongly prominent, densely and persistently scurfy like the costules, with many erect, white hairs to 2.0 mm long, also with few tan to brown, flat to subbullate squamules to 2.0 mm long, with flat to hair-like apices; costules basally without pneumathodes, blackened in dried specimens. Largest segments 9.0–11.0 × 3.0–4.0 mm, sessile, adnate, ascending, distally straight, tips obtuse, proximal segments alternate to subopposite, usually a bit shorter than following segments, sometimes remote and then strongly crenate; sinuses acute to rectangular, ca. 1.0–2.0 mm wide; margins crenulate to crenate; margins not differently incised in proximal segments of a pinnule; veins planar to weakly protruding adaxially and abaxially, dark carnose to atropurpureous or blackish, ending in the margins; veins adaxially and abaxially with many erect, white to yellowish white, multicellular hairs to 2.0 mm long on a them, also along the margins but not between the veins; midveins with few, white to light brown bullate squamules to 2.0 × 0.5 mm; sterile veins simple or forked, fertile veins forked. Sori 1.0(–1.2) mm diameter, medial, in the fork of veins, mature dark orange-brown; indusia lacking; receptacles globose, 0.2–0.3 mm diameter, paraphyses numerous, heavily contorted, whitish to tan, longer than the sporangia (0.6–0.8 mm long) and persisting in over-mature sori. Spores not examined.

Distribution and habitat: — Panama, Colombia and Venezuela, in evergreen and semideciduous lowland and premontane forest at 0–1300 m, often occurring on sandstone.

Additional specimens examined: — PANAMA. Darien: Alturas de Nique, above Cana mine, 07°45’N, 77°40’W, 600–800 m, 2 March 1992, G. McPherson 12182 ( MO!, UC!). Panamá: Río La Maestra, 0–25 m, 4 December 1936, P. H. Allen 14 ( GH!).— COLOMBIA. Antioquia: San Luis, 16 km SW de las partidas a San Luis, sobre la via Medellin- Bogotá, Vereda La Josefina , 06°00’N, 74°50’W, 800 m, 26 June 1991, R. Callejas et al. 4200 ( COL) ; San Carlos, Corregimiento Alto de Samana, Vereda Miraflores, finca “El Desespero,” bosques a 2 horas de camino NE de la finca, en el camino a “Jardin”, 06º05’N, 74º50’W, 750–890 m, 26 October 1989, R. Callejas et al. 8583 ( UC!, US!). Santander: Between Girón and Lebrija, 1300 m, 28 February 1939, A. H. G. Alston 7360 ( BM!, MO!). Vaupes: Upper Apaporis Basin, Río Macaya , Cerro Chiribiquete, 365–640 m, 24 July 1943, R. E. Schultes 5635 ( GH!, US!).— VENEZUELA. Apure: Reserva Forestal San Camilo, Rio Nulita , 300 m, 5 April 1968, J. A. Steyermark et al. 101901 ( GH). Barinas: 0.5 km SW of dam site on Río Caparo , 31 km ESE of Santa Barbara, 07º41’N, 71º28’W, 100–300 m, 10 March 1980, R. Liesner & A. González 9328 ( UC!). Mérida: Tovar, Los Giros near El Osa, ca. 5 km from previous location, 08º27’N, 77º44’W, 700 m, 26 July 1983, H. van der Werff & R. Ortiz 5721 ( UC!). Tachira: Vertientes del valle situado inmediatamente al oeste de Ayari, 200 m, 27 August 1966, J. A. Steyermark & M. Rabe 96651 ( GH) ; SE facing slopes of Cerro of Cuchilla La Pabellana, W of San Joaquina de Navay , 07º37’30”N, 71º47’00”W, 250–350 m, 06 November 1979, J. A. Steyermark et al. 19379 ( MO!, UC!). Zulia: Colón, surroundings of Casigua, sector Las Cruces, 18–20 km SSW of the village and to the pozo T- 221, 250 m, 20 February 1979, G. S. Bunting 6696 ( UC!) ; Catatumbo, along road Machiques-La Fría and Río de Oro , between Km 5–20, 100–200 m, 19 February 1982, G. S. Bunting 10762 ( UC!) ; 3 km E of the Río de Oro settlement, 09º18’N, 72º28’W, 100–250 m, 26–28 June 1980, G. Davidse et al. 18625 ( NY!, UC!) ; ca. 55 km SW of Machiques by air, Aricuaisa (Ariguaisa), pie de monte on Río Aricuaisa , 09º36’N, 72º54’W, 100–250 m, 24–25 March 1982, R. Liesner & A. González 13140 ( UC!) ; 6 km ENE of Río de Oro , 09º06’N, 72º52’W, 100–350 m, 28 March 1982, R. Liesner & A. González 13303 ( UC!) ; above Camp 2, Perija Exploration Company, on Río Lora , 14 December 1922, H. Pittier 10945 ( GH!) .

Remarks: —The most noteworthy feature of Cyathea lockwoodiana is the abundant persistent prapaphyses that are much longer than the sporangia and heavily contorted in their distal half. This character separates it from C. pilosissima , C. senilis and C. parianensis , which have spreading persistent hairs on the leaf axes and relatively pale laminar squamules like C. lockwoodiana but which have their paraphyses not contorted.Among them, C. senilis further matches C. lockwoodiana in having reddish brown scurf on the petioles but differs in its concolorous laminar squamules (tan to pale brown with attenuate to subulate tips in C. lockwoodiana vs. whitish bullate squamules with flattened dark brown tips in C. senilis ). The appearance in the field matches between C. lockwoodiana and C. pilosissima ( Fig. 2 ), and presumably also with C. parianensis and C. senilis .

The group formed by Cyathea lasiosora , C. wendlandii , C. schlimii and C. pauciflora is characterized by having dark brown to reddish brown squamellar indument on the laminae (vs. squamellar indument tan to pale brown in C. lockwoodiana ) but lacking spreading hairs here (not counting appressed hairs on the adaxial side of distal petiole parts vs. spreading hairs extending to petiole bases). Cyathea lasiosora has distally contorted paraphyses just like C. lockwoodiana , but these are relatively few and tufted apically on the receptacles (vs. densely packed and equally distributed on the receptacles in C. lockwoodiana ) and broken off in older dried material (vs. persistent). Furthermore, the contorted paraphyses are an artefact from drying in C. lasiosora and apparently straight in fresh material (see discussion under that species). The spreading hairs on the abaxial surfaces of the leaf axes in C. lockwoodiana are tan to reddish, relatively long (to 2 mm) and generally dense, persisting and leave a scabrous surface if abraded (vs. hairs on the abaxial sides of the leaf axes white, relatively short, ± 1 mm long in C. lasiosora and the other species of its group).

One population in the eastern Andean foothills of southern Ecuador and northern Peru is intermediate between C. lockwoodiana and C. lasiosora and cannot be placed in neither species confidently. These are separated here as C. calamitatis .

Figures

FIGURE 2. Cyathea spp. in the field: A1–D1. Trunk apex; A2–D2. Lamina apex; showing different expressions of abruptly reduced apices; A3–D3. Details of rhachichses, B–D at junctures with costae, showing presence and extend of pubescence and alae (D3). A. Cyathea pilosissima (Dept. San Martin, Peru). B. Cyathea calamitatis, sp. nov. (Prov. Morona-Santiago, Ecuador). C. Cyathea lasiosora (Dept. San Martin, Peru). D. Cyathea tortuosa (Prov. Zamora-Chinchipe, Ecuador).

FIGURE 4. Cyathea lockwoodiana. A. Lower petiole, showing scales. B. Junction of rhachis and costa abaxially. C. Pinnule, adaxially. (All from Callejas et al. 4200, COL)