Cyathea divergens var. divergens

Lehnert, Marcus, 2014, Do you know Cyathea divergens (Cyatheaceae-Polypodiopsida)?, Phytotaxa 161 (1), pp. 1-42 : 10-39

Cyathea divergens var. divergens Figs. 2A , 4 .

Cyathea equestris Kunze (1834: 100) . Type:— PERU. Huánuco: Pampayaco, Cerro de Cristobal , July 1829, Poeppig s.n. (holotype LZ, destroyed,-fragment K,-photo GH)

Cyathea globularis Presl (1849 [1851]: 30), from description. Type :— COLOMBIA. Prov. unknown: “Nova Grenada,” without date, Linden s.n. (holotype not located).

Cyathea calva Karsten (1869: 175) .—Type: VENEZUELA. Mérida: Sierra de Mérida, Escuque, 1000 m, without date, H. Karsten s.n. (holotype B-fragment NY,-fragment US)

Cyathea petiolulata Karsten (1869: 163) . Alsophila petiolulata Mettenius (1864: 263) , nom. nud. Type: — VENEZUELA. Mérida: Mérida, 2000–2500 m, without date, Karsten s.n. (holotype not located). Authentic specimen: Mérida, Karsten 135 (B).

Cyathea equestris var. boconensis Karsten (1856: 456) . Cyathea petiolulata var. boconensis (H.Karst.) Karsten (1869: 164) .— Type: VENEZUELA. Mérida: Sierra de Mérida, Paramo de Bocono, without date, Karsten s.n. (holotype not located, W?).

Alsophila subaspera H.Christ in Pittier (1901: 43). Cyathea subaspera (H.Christ) Domin (1929: 263) . Type:— COSTA RICA. San José: “Forets de Copey,” 1800 m, Tonduz 11787 (Lectotype P, here designated, isolectotypes A, GH, NY, US). Excluded syntypes: Tonduz 11802 (BR-fragment NY, P), Tonduz 12183 (NY).

Cyathea pelliculosa Christ (1904: 946) .— Type : COSTA RICA. Prov. unknown: Without locality, without date, Wercklé 45 (lectotype P, here designated).

Cyathea petiolulata var. pastoensis Hieronymus (1904: 437) . Type:— COLOMBIA. Putumayo: “Prope Altaquer et San Pablo, Cordillera del Pasto,” without date, Lehmann 81 (holotype not located).

Cyathea divergens var. minor Rosenstock (1925: 2) . Type:— COSTA RICA. Alajuela: La Palma, without date, Brade & Brade 108 (lectotype not located, chosen by Tryon 1976: 54, isolectotype US). An excluded element (Brade & Brade 853) belongs to Cyathea gracilis Grisebach (1864: 704) ( Tryon 1976: 54) .

Cyathea divergens var. hirta Losch (1950: 20) . Type:— COSTA RICA. Cartago: Chirripo Grande, without date, Kupper 1265 (holotype M).

Trunks to 6(–10) m tall, 6–10 cm diameter, with old petiole bases when smaller than 2.5 m, without them when larger; upper parts covered in concolorous brown to bicolorous scales, similar to petiole scales; frond scars circular to weakly elliptic; trunk apices hidden in fascicles of the youngest petioles; adventitious buds lacking. Fronds to 630 cm long, arching, distally drooping, often to the ground. Petioles to 200 cm long, muricate to aculeate with spines to 5 mm long, stramineous to tan or brown; without adventitious (aphlebioid) pinnae at the petiole bases. Petiole scales narrowly lanceolate to lanceolate, (12–)20–30 × 3–4 mm, their tips weakly twisted, the brown to dark brown centre sharply set against the white to stamineous margins, usually not shiny; petiole scurf well developed, white to stramineous, consisting of several types of squamules and small scales; mainly erect white lanceolate squamules 0.5–2.0 mm long with darkened insertion, with (in larger plants, especially Colombian material) or without (smaller plants, prevailing in Costarican and Venezuelan material) irregularly set dark brown to black marginal teeth, smaller concolorous white squamules occur scattered as undercoat. Laminae to 430 × 190 cm, bipinnate-pinnatifid to tripinnate, firm chartaceous to subcoriaceous, widest at the middle, apex gradually reduced and long drooping; medium green adaxially, usually plumbeous when dried, pale green abaxially. Largest pinnae 60–95 cm long, 15–20(–29?) pairs per frond, long-stalked 3–6(–11) cm, distally narrowly green-alate, the distal segments decurrently adnate. Frond axes stramineous to brown, pluricellular hairs only adaxially on costules and distal parts of costae and rachises, hairs 0.5–1.0 mm long, whitish to tan, abaxially glabrescent with white to brown scurf consisting of small squamules 0.2–1.0 mm long, unlike those of the petiole, usually strongly dissected to fimbriate, persisting in junctures of costae with costules and rachises; costules sometimes also with small ovatelanceolate scales 2–3 mm long, dark brown with the white margins irregularly streaked brown; costae muricate, rarely more than 3–4 mm wide. Largest pinnules (10.0–)14.0–17.5 × (1.5–)2.5–3.5(–4.5) cm, short- to long-stalked (5.0–10.0 mm), alternate, (0.5–)1.0– 2.5 cm between the stalks, long triangular to oblong, truncate at base, tapering from beyond the middle (rarely in large pinnules from the base) to acute or short attenuate tips, the segments weakly falcate with finely crenate margins and rounded to obtuse, rarely acute tips; basal segments alternately placed, sometimes remote from each other, sinuses wide (2–3 mm) and obtuse; sterile pinnules usually broader than fertile ones. Veins glabrous abaxially except for few white unicellular trichomidia; with small flattish, brown, ovate squamules with elongated tips and finely dissected dark brown squamules, sometimes single subbullate brown squamules on the midribs of segments distally; sterile veins forked once to twice or simple, fertile veins forked. Sori 1.0(–1.5) mm diameter, proximal to submedial, indusium sphaeropteroid, with umbo, tan, translucent, fragile but persisting as a cup; paraphyses of the same length or shorter than sporangia. Spores not examined.

Distribution and habitat: — Costa Rica, Panama, Colombia, Venezuela, Ecuador to central Peru, in moist tropical montane forests at 1000–2770 m.

Etymology: —The epithet refers to the drooping fronds of the species, thus having an orientation divergent to the trunk (Latin divergere = going different ways; vergere = bending over).

Additional specimens examined: — COSTA RICA. Cartago: Finca La Esperanza, ca. 3 km E of Muñeco and 2 km SW of Navarro , 1200–1500 m, 13 July 1970, Lellinger & White III 1120 ( F) ; 1 km W of Pacayas along road # CR 230 , 1620 m, 9 April 1967, Lent 787 ( F) . Puntarenas: Upper Río Burú , 2100 m, 20 August 1987, Gómez et al. 21696 ( AAU) . San José: 25 km N of San Isidro del General twoards Cerro de La Muerte , 1800 m, December 1967, Gastony & Gastony 755 ( F) .— PANAMA. Chiriquí: Rio Piedra Candela, road to Las Mellisas , 1800–2000 m, 01 May 1973, McAlpin 2192 ( GH) ; vicinity of Casita Alta, Volcan de Chiriqui, 1800–2000 m, 28 June–2 July 1938, Woodson, Allen & Seibert 922 ( GH) ; above the Río Caldera beyond Bajo Mono, in vicintiy of Boquete , 1800 m, 17 January 1970, Wilbur et al. 11077 ( F) . Panamá: Cerro Jefe, 100 m al N de la torre, 1000 m, 11 September 1990, Valdespino & Aranda 247 ( AAU) .— COLOMBIA. Antioquia: Jardin, Vereda La Mesenia, sector La Orqueta , 05°30'15"N, 75°53'20"W, 2200–2300 m, 1 November 2009, Rodriguez et al. ( NY) ; San Antonio del Prado , August 1943, Daniel 307 ( GH) ; Urrao, Vereda Calles, Parque Nacional Natural "Las Orquideas", right slope of Quebrada La Honda , 06º32'N, 76º19'W, 1330–1400 m, 7 May 1993, Cogollo et al. 6162 ( MO, UC) ; Antadó, valley of Río San Jorge, Paramilo National Park , 07º15'N, 75º55'W, 1560 m, 3 January 1993, Gentry et al. 78832 ( MO, UC) . Caldas: Canaan, south of Salento , 8 August 1922, Hazen 9693 ( GH) . Cauca: Popayán, Rio Molino, vereda Santa Barbara, 1950 m, 15 July 2007, Munar & Ceballos 152 ( COL) . Cundinamarca: En el kilometro 14.1 de la carretera Fómeque a Chingaza , 2770 m, 28 April 1978, Acosta Arteaga 351 ( MO) ; Guaduas, Vereda La Campeona, camino a La Cabaña-Cordillera Montefrio , 1900 m, 20 July 1991, Linares & Devia 1945 ( COL) . El Valle: La Cumbra, Cordillera Occidental ; bushy summit of west peak, 2100–2400 m, 11–18 September 1922, Killip 11392 ( GH, US) . Huila: La Argentina, Quebrada del Pueblo , 1850 m, 26 September 1988, Lozano et al. 3996 ( COL) . Magdalena: Santa Marta, 1898–1901, Smith 2225 ( UC) . Nariño: Pasto, Finca de Chimayoy , Km 11 Pasto-Chachagüi , 01º23'N, 77º16'W, 2650 m, 27 July 1991, Ramírez-P. & Cuayal-M. 3941 ( UC) . Norte de Santander: Loso and vicinity ( N of Toledo), 2200–2400 m, 6–7 March 1927, Killip & Smith 20407 ( COL, GH) . Quindio : "Provincia de Mariquita: Quindio", 1800 m, January 1852, Herb. Triana 653 ( COL) . Risaralda: Santuario, Vereda Las Colonias , margen derecha del Rio San Rafael , 2500 m, 26 February 1987, Torres et al. 2314 ( COL) . Valle de Cauca: Río Calima watershed, El Cairo, between Darién and Mediacanoa, 1650–1750 m, 6–7 January 1943, Cuatrecasas 13919 ( F) ; km 19 on road from Cali to Buenaventura , 8 August 1976, Barrington 500 ( COL) ; Argelia, Vereda Las Brisas, finca San Jorge, 1950 m, 23 January 1987, Franco et al. 1724 ( COL) .— VENEZUELA. Lara: Crespo, Quebrada de Oro , 1500 m, 7 November 1982, Smith V9366 ( UC) ; Palavecino , sector urbanizado, 09º55'N, 69º17'W, 500 m, 28 September 1984, Rivero R. 722 ( UC) . Mérida: Mérida, along trail below first stop above base of teleferico, 1 August 1990, Conant 4449 ( LSC) . Portuguesa: Portuguesa, 50 km WNW of Guanare by air, 09º28'N, 69º55'W, 1300–1500 m, 15 March 1982, Liesner et al. 12772 ( UC) . Tachira: Quebrada La Lejia , S of Quebrada Azul, 07º30'N, 72º24'W, 2150–2300 m, 25 July 1979, Steyermark & Liesner 118552 ( UC) . Trujillo: En las cabeceras de Vitú, Cerro El Zamuro, Quebrada El Limón , 09°28'N, 70°24'W, 1800–1900 m, 24 November 1988, Ortega & van der Werff 2291 ( AAU, UC) . Yaracuy: Districto Bruzual: Serrania de Aroa, fila entre los rios Tejar y Coritico , 14–15 km al NNE de Urachiche , 3 km al NE de Caserio Buenos Aires, 10º14'40"N, 68º59'30"W, 1350–1500 m, 1 March 1981, Steyermark et al. 124820 ( GH) .— ECUADOR. Carchi: Espejo, El Gualtal, Faldas del Cerro Golondrina Hembra, 00°51'N, 78°07'W, 2450 m, 22 August 1998, Palacios & Clark 12617 ( QCNE, UC) . Napo: Quijos, Reserva Ecologica Antisana, Cordillera de los Guacamayos , sector oriente, cruce del oleoducto de la companía ARCO, entre El Mirador y Camino de la Virgen, 00°38'S, 77°51'W, 2300 m, 12–14 January 1999, Vargas & Navarrete 3489 ( MO, QCNE) . Tungurahua: Río Verde , 5 June 1968, Harling, Storm & Ström 10178 ( GH, UC) .— PERU. Oxapampa: Dist. Villa Rica, centro poblado Palma ( Centro Bocaz ), 10º39'24"S, 75º09'42"W, 1736–1800 m, 16 January 2005, Mellado L. et al. 2629 ( MO) , Centro Bocaz, road and trench to Purus , 10º38'S, 75º11'W, 1590 m, 19 September 2003, Perea et al. 376 ( MO, UC) .

Remarks:— The specimens of Cyathea divergens var. divergens show a considerable variation in pinnule shape. Some have mostly oblong-lanceolate pinnules as they are generally found in the outer parts of the blade (margins and apex) and none of the clearly triangular ones ( Fig. 4E ).

Poeppig (1836: 189) reported from the type locality in Peru that the drooping fronds usually reach to the ground, which can be corroborated for Costa Rica (personal observation). In the same source Poeppig also reports that C. divergens is usually found in more open, lower vegegation and under drier conditions than the sympatric species of Cyathea .

Other species similar in leaf division are C. ebenina Karsten (1856: 461) , C. gibbosa ( Klotzsch 1844: 542) Domin (1929: 262) and C. gracilis Grisebach (1864: 704) and C. tuerckheimii Maxon (1909: 4) , but these species have dark brown, atropurpureous or even blackish axes and smooth costae (vs. frond axes mostly pale, yellowish or stramineous, only rarely light brown, the costae muricate or at least verrucate to some extend in C. divergens ). All display the entire variability in pinnules shape found in C. divergens but each extreme is restricted to one species and correlated with other characters: Cyathea ebenina has predominantly oblong pinnules but is recognized by the orange petiole scale margins; Cyathea gibbosa has mostly acute, falcate segments but lacks indusia and persistent petiole scurf; Cyathea gracilis has triangular to oblong pinnules with crenulate margins but ovate-lanceolate, dark brown scales and lacks scurf entirely. Cyathea maxonii Underw. ex Maxon (1909b: 82) from Costa Rica and Panama can be easily distinguished from the sympatric C. divergens by having patent-arching fronds that are not drooping to the ground, ± sessile pinnules and the absence of pale colors in the indument (scurf always brown, rather sparse, petioles scales brown to partially black in C. maxonii vs. scurf whitish when dry, petiole scales with pale margins in C. divergens ) ( Figs. 2A, C ).

Cyathea tuerckheimii is no longer considered a variety of C. divergens ( Tryon 1976) . Besides differing in similar characters as the newly recognized Cyathea divergens var. sipaliwiniana from var. divergens (i.e., scurf morphology; different variability in pinnules size, shape and stalks; peripheral geographic distribution), C. tuerckheimii also differs from C. divergens var. divergens in the color of the petiole scales (centres more reddish brown in C. tuerckheimii vs. brown to blackish in C. divergens ) and the softer laminar texture (firm herbaceous vs. subcoriaceous). Furthermore, the pinnules of C. tuerckheimii are positioned more closely to each other than in C. divergens (largest ones touching to overlapping vs. usually with gap between them). See C. tuerckheimii for further discussion.

Cyathea divergens var. sipaliwiniana Lehnert , var. nov.

Differt a var. divergente squamulis petiolorum obscurioribus cum dentibus pallidioribus.

Type: — SURINAME. Sipaliwini: Grace Camp, Tafelberg Mountain , N side of Grace Kreek Camp, 1 km N of Grace Falls , 03º54'32"N, 56º12'44"W, 800–820 m, 4 July 1998, Hawkins 1877 (holotype UC, isotype MO) .

Trunks to 8 m tall, to 12 cm diameter, presumably without old petiole bases; upper parts covered in bicolorous scales, similar to petiole scales. Fronds 150–600 cm long. Petioles to 200 cm long, muricate to shortly aculeate with spines to 2.5 mm long, tan to brown, basally darker; without adventitious (aphlebioid) pinnae at the petiole bases; petiole scales narrowly lanceolate, 15–20(–25) × 23 mm, their tips straight, concordantly bicolorous, the brown to dark brown centre sharply set against the white to yellowish margin, usually shiny; petiole scurf well developed, whitish to pale brown, consisting of several types of squamules; mainly erect to ovate to lanceolate squamules 0.5–1.0 mm long, with a brown to tan centre, whitish to yellowish margins and irregularly set, small pale brown marginal teeth. Laminae (150–)250–400 × 50–140 cm, bipinnate-pinnatifid, firm chartaceous, apex gradually reduced and long drooping; dark green adaxially, usually blackish when dried, pale green abaxially. Pinnae to 70(–80) cm long, 15(–20) pairs per frond, short to long stalked (1.8–)2.0–3.0 cm, distally narrowly green alate, the distal segments decurrently adnate. Frond axes brown to dark brown, hairy only adaxially on costules and distal parts of costae and rachises, hairs 0.5–1.0 mm long, whitish to tan, without hairs abaxially, but with tan to brown scurf consisting of small squamules 0.2–1.0 mm long, like those of the petiole, usually strongly dissected to fimbriate, persisting in junctures of costae with costules and rachis; costae smooth to weakly muricate, rarely more than 3–4 mm wide. Largest pinnules (7.0–)10.0–11.5 × 1.8–2.2(–2.5) cm, alternate, stalked 4–5 mm, 1.5–2.3 cm between the stalks, long triangular to oblong, truncate to weakly cordate at base, tapering from beyond the middle to short attenuate tips, the segments patent to weakly falcate with finely crenate margins and rounded to obtuse tips; basal segments alternately placed, the lowest ones sometimes remote from each other, sinuses narrow to wide 2.0(– 4.0) mm, obtuse. Veins mostly glabrous adaxially except for occasionally single multicellular hairs on the midrib; abaxially with few white multicellular hairs on and sometimes between the veins, brown unicellular trichomidia and small squamules; the latter vary from auburn to brown, flattish, ovate with elongated tips to dark brown, finely dissected squamules, sometimes single subbullate brown squamules on the midribs of segments distally; sterile veins forked once or simple, fertile veins forked. Sori 1.0– 1.5 mm diameter, proximal to subproximal, indusium sphaeropteroid, with umbo, tan, translucent, fragile but persisting as a cup; paraphyses as long as or shorter than sporangia. Spores not examined.

Distribution and habitat: —Eastern Venezuela, Guyana and Suriname, in mixed hardwood forests along streams at 800–2050 m.

Etymology: —The epithet refers to the province where the type locality is situated.

Additional specimens examined:— VENEZUELA. Bolivar : Ptari-tepui, just NE of "Cave Rock", 4 November 1944, Steyermark 59811 ( F); Meseta del Jaua , Cerro Jaua , cumbre, porcion sur-oeste, al borde del tributario del Río Marajano , 04°48'50"N, 64°34´10"W, 1750–1800 m, 22–28 February 1974, Steyermark, Espinoza & Brewer-Carias 109395 ( US); Sarven-tepui, 1900–2050 m, 13 January 1953, Wurdack 34115 ( US) .— GUYANA. Region Potaro-Siparuni : Pakaraima Mountains, Mount Wokomung , summit ridge of Ka-mie-wah pinnacle NE to S pinnacle, "Little Ayanganna", 05º04'N, 59º52'W, 1550–1650 m, 17 November 1993, Henkel et al. 4486 ( US) .

Remarks:— Cyathea divergens var. sipaliwiniana differs from the eponymous variety in its larger scurf squamules, which have daker centres (brown to dark brown with paler margins in var. sipaliwiniana vs. in the same pale color as the margins or slightly darker in var. divergens ). This makes the whole appearance of the petiole scurf darker and fuzzier. In the main population in the Andes, the largest scurf squamules would have dark brown marginal teeth whereas their bodies remain uniformly pale; it is the other way around in var. sipaliwiniana . Cyathea divergens var. sipaliwiniana also has its largest pinnules shorter and more shortly stalked than var. divergens on average (to 11 cm with 4–5 mm stalk vs. to 17.5 cm long with 5–10 mm stalk). Furthermore, the soral position is less variable (proximal to subproximal vs. proximal to submedial), which is especially evident when just the receptacles are left. Together with its occurrence in a different biogeographic region, these characters seem to justify the recognition of Cyathea divergens var. sipaliwiniana at the rank of a variety.

Cyathea ebenina Karsten (1856: 461) . Type:— VENEZUELA. Aragua: Between Caracas and Puerto Cabello , Karsten s.n. (holotype not located, W?, isotype B [labeled „Caracas“]). Figs. 2E , 5 .

Trunks to 4 m tall, 3.5–4.0 cm diameter, without old petiole bases, most parts covered in concolorous to bicolorous scales, similar to petiole scales; epidermis dark brown to blackish, with few vermillion, 0.3–1.0 mm long, elliptic pneumathodes, often obscured by scales; adventitious roots only near the trunk bases; apices not hidden in fascicle of petioles, unfolding only one crosier at the time. Petioles (37–) 50–150 cm long, inermous, dark castaneous to atropurpureous, basally black; without adventitious (aphlebioid) pinnae at the petiole bases; petiole scales lanceolate to ovate-lanceolate, (12–)15–20 × 35 mm, their tips straight to falcate, shiny orange-brown, discordantly bicolorous with a darker brown to castaneous central stripe; petiole scurf absent. Fronds to 200–300 cm long, strongly arching. Laminae to 100–250 × 50–70 cm, bipinnate-pinnatifid, firm herbaceous to chartaceous, apex gradually reduced and drooping; glossy dark green adaxially, often yellowish when dried, bright green abaxially. Pinnae to 25–35 cm long, 8–12 pairs per frond, short to long stalked 0.4–2.0 cm, distally narrowly green alate, the distal segments decurrently adnate. Rachises and costae dark-castaneous to black, costules pale brown to yellowish, hairy only adaxially on costules, costae and distal parts of rachises, hairs 0.5–1.0 mm long, tan to brown, without hairs abaxially; costae smooth, rarely more than 2 mm wide; insertions of costae into rachises with two pale brown to yellowish pneumathodes, either separate, each to 2 mm long, or contiguous to 5 mm. Largest pinnules 6.0–9.5 × 1.8–2.5 cm, the largest ones stalked 3.0–5.0 mm, alternate, 0.5–2.2 cm between the stalks, ovate lanceolate, truncate to weakly cordate at base, tapering from beyond the middle to short attenuate tips; the yellowish to brown stalks and costules contrasting with the atropurpureous to blackish costae, the segments weakly falcate with finely to coarsely crenate margins and rounded to obtuse, rarely short acute tips; basal segments opposite to alternate, the lowest ones not remote from each other, sinuses narrow 2.0–3.0 mm, if wider then enclosed by segment tips, acute. Veins pale brown to yellowish, contiguous with the identically coloured hyaline segment margin, mostly glabrous adaxially except for occasionally single multicellular hairs on the midrib; abaxially glabrous except for occasional tan to castaneous unicellular trichomoidia on the veins; sterile veins simple or forked, fertile veins forked. Sori (1.0–)1.2–2.0 mm diameter, subproximal to medial, deep brown, indusia deeply urceolate to subshaeropteroid, sometimes sphaeropteroid with weak apical umbo, tan to auburn, not translucent, fragile but persisting as a fragmented cup, receptacles globose, to 0.6 mm diameter; paraphyses few, hyaline, shorter than sporangia (ca. 0.2 mm). Spores not examined.

Distribution and habitat: — Colombia, Venezuela, Ecuador and northern Peru at (1000–) 1600–2450 m; growing terrestrial, mainly in the understory of perhumid montane forests.

Etymology: —The epithet refers to the shiny ebony-color of the petioles and frond axes.

Additional specimens examined:— COLOMBIA. Antioquia: Ciudad Bolivar, Corregimiento San Bernardo de los Farallones , cerro San Nicolas-Farallones del Citará , sector El Aserradero , 05°46'09"N, 76°02'11"W, 2100– 2250 m, 11 September 2008, Rodriguez et al. 4637 ( NY). Huila: Cordillera Oriental, on bank of Río Negro , ESE of Baraya, 8700 ft, 31 October 1944, Little 8905 (US) .— VENEZUELA. Portuguesa: Region of Cerro Córdoba , 16–17 km E of Chabasaquen, 09°26–27' N , 69°54–55'W, 1600–1620 m, 30 October 1982, Steyermark, Liesner & Stergios 126826 ( UC) .— ECUADOR. Tungurahua: Río Verde , 6 June 1972, Harling, Storm & Ström 10178 ( F, US). Zamora-Chinchipe: Estación Cientifica San Francisco, above refuge, along trail T1 study plot A 2 , 03°59´33"S, 79°04'15"W, 2550 m, 29–30 September 2003, Lehnert 937 ( GOET, QCA, UC); Reserva Tapichalaca, study plot B3 , near Ventanillas , 04°29'S, 79°07'W, 2520 m, 2 November 2007, Lehnert 1055 ( GOET, QCA, UC) .— PERU. Amazonas: Bongara, on road to Pomacocha , 28 km above Puente Ingenio, 2200 m, 7 October 1964, Hutchinson & Wright 6817 ( UC). Cajamarca: San Ignacio , San José de Lourdes , locality Picorana , 04°58'00"S, 78°53'01"W, 2350–2450 m, 3 December 1998, Campos, Zurita & Camizan 5898 ( UC) .

Remarks: —The shiny black petioles, rachises and costae that contrast strongly with the greenish to brown costules and the mainly orange-brown petiole scales are the best distinguishing characters of Cyathea ebenina . In C. gracilis , the petioles and frond axes vary more in colour and are not so intensely contrasting with scales and costules. Cyathea kalbreyeri ( Fig. 10 ) agrees with C. ebenina in the frond structure (i.e., few, relatively short pinnae with large pinnules with oblique segments and shiny blackish frond axes; Fig. 2E ), but differs in the exindusiate sori (vs. sori with sphaeropteroid indusia in C. ebenina ).

Cyathea ebenina is a very attractive species, but due to its small size, low population densities and preference of shady understories, it is often overlooked. It may thus be anticipated that is occurs even further south in the Andes. Suitable habitats extend into Bolivia, e.g. Parque Nacional Madidi, Dept. La Paz.

Cyathea farinosa (H.Karst.) Domin (1929: 262) . Alsophila farinosa Mettenius (1864: 262) , nom. nud. Alsophila farinosa Karsten (1869: 163) . Type:— VENEZUELA. Aragua: “Prope Colonia Tovar, Caracas,“ without date, H. Karsten s.n. (holotype B-fragment GH, isotypes M, GH n.v.-photo M, US [labeled “ Karsten 49 ”]). Fig. 6 .

Trunks (0.3–)1.0–4.0 m tall, 5.0– 7.5 cm diameter, straight to decumbent, with persistent old petiole bases, spiny or not; adventitious buds not reported. Fronds to 250 cm long, patent to ascending. Petioles more than 35 cm long, inermous to spiny, spines 3–5 mm long, brown to atropurpureous; lenticels to 10 × 1 mm, inconspicuously brown in dried material; without adventitious (aphlebioid) pinnae at the petiole bases; petiole scales lanceolate, 14.0–26.0 × (1.5–)2.5–4.0 mm, their tips straight, shiny auburn to dark brown, concordantly bicolorous with narrow, pale to golden brown margins, colors often sharply contrasted; petiole scurf a matted tomentum of small branched hairs and dissected squamules 0.2–0.4 mm long, yellowish white with brown parts, grayish white in general aspect, soon caduceus, persistent between spines; sometimes small erect brown squamules to 0.8 mm long between lower petioles scales. Laminae ca. 150–200 × 80–100 cm, bipinnate-pinnatifid, chartaceous, dark olive-green adaxially, often blackish when dried, olive-green abaxially, apices gradually reduced. Largest pinnae 40–50 cm long, 12–20 pairs per frond, stalked to 2.5 cm, distally not green-alate, the pinnatifid terminal segments shortly decurrent into the costae. Frond axes brown to atropurpureous on both sides; completely glabrous abaxially except for scurf remnants, containing appressed, white trichomoid squamules (0.2–0.5 mm), appressed brown squamules with white ciliate margins (0.4–0.6 mm) and erect, subbullate brown squamules with entire margins (0.5–0.8 mm); hairy only adaxially on costules, costae and distal parts of rachises, hairs 1.0– 1.5 mm long, tan to brown; costae smooth, 3–4 mm wide; insertions of costae into rachises weakly swollen, each with two inconspicuous planar pneumathodes, orange brown, elliptic, to 2.0 × 1.0 mm. Pinnules (4.5–)6.0–10.0 × (1.1–) 1.7–2.3 cm, the largest ones stalked 10–14 mm, alternate, 1.5–2.0(–2.6) cm between the stalks, linearly oblong to lanceolate, incised to 1/ 2 or more of their width, bases truncate to weakly cordate, in larger pinnules (> 2 cm wide) sometimes with basiscopical auricles; pinnules tapering from beyond the middle to long acuminate to short-attenuate tips; the brown stalks articulate, with a black ring going all around their bases (abscission layer), their bases with an orange brown, elliptic pneumathode to 2.5 × 0.6 mm, inconspicuous and often hidden in folds in dried specimens; segments oblong to deltate, patent to ascending, with entire to weakly dentate margins; tips weakly falcate, obtuse to acute, or often at least appearing so due to a keel formed by the revolute margins; basal segments usually opposite, the lowest ones not remote from each other, sinuses triangular, acute and narrow (1.0–2.0 mm). Veins weakly prominent abaxially and adaxially, ending at segment margins; midveins yellowish brown abaxially and adaxially, lateral veins yellowish brown adaxially, yellowish or not different from the lamina in color abaxially; glabrous adaxially except for occasional single hairs on the midveins, abaxially glabrous to scurfy, with white to tan, tortuous hairs on the veins, easily abraded, also with auburn to castaneous squamules with entire margins, subbullate to bullate ones to 0.6 mm long mainly on lateral veins and distal midveins, larger flattish ones to 2 × 1 mm on costules and proximally on midveins; sterile and fertile veins simple or forked. Sori (0.8–)1.0– 1.2 mm diameter, medial, indusia absent, but with ephemeral brown scales to 1.5 mm long subtending the receptacles; receptacles globose to ellipsoid, 0.2–0.4 mm diameter; paraphyses few to numerous, hyaline, white, longer than sporangia (0.6–0.8 mm). Spores not examined.

Distribution and habitat: — Venezuela, Cordillera de la Costa, in cloud forests at 1000–2200(–2700) m.

Additional specimens examined: — VENEZUELA. Aragua: Loma del Hierro, SE of Las Tejerías , W of road Las Terías-Tiara, above the Estación de Peste , 10º10'N, 67º09'W, 1000–1350 m, 29 November 1998, Meier & Speckmeier 4278 ( UC); Colonia Tovar and vicinity, 2100–2700 m, 26 December 1921, Pittier 9995 ( US) . Distrito Federal: Border between Distrito Federal and Miranda, P. N . El Avila, Cordillera de la Costa, NE of Guatire , trail to La Sabana , 10º32'N, 66º28'W, 1450–1550 m, 29 February 2001, Meier & Hérnandez-Bretón 10023 ( UC). Falcón: Sierra San Luis, above Santa Maria, 1080 m, 2 January 1979, Flora Falcón ( HW, TR, BV, ES) 302 ( UC). Miranda : Headwaters of quebrada Chacaito, 2200 m, 28 March 1992, Meier & Groeger 1963 ( UC). Tachira: Along road from La Honda dam to La Fundacíon, 4 August 1990, Conant 4462 ( LSC). Vargas: Cordillera de la Costa, Serranía del Litoral , Monumento Natural Pico Codazzi, road Arcos de la Colonia Tovar-Purto Cruz, 1 km from Arco, Urbanización Residencial Jengibrillar, Av. La Neblina, Sector La Neblina , end of Calle 2 , 10º26'N, 67º14'W, 2000– 2100 m, 6 February 1999, Mostacero, Leal & Grau 89 ( UC). Yaracuy: Nirgua / San Felipe, NW of Nirgua, Montaña El Zapatero, SW part, Fila Jaiguao, caserio Mundo Nuevo, N slope, 10º14'N, 68º38'30''W, 1100–1400 m, 28 December 2000, Meier, Bergold & Speckmeier 7828 ( UC) .

Remarks: —The name Cyathea farinosa is here applied to the species previously known as C. gibbosa . The name C. gibbosa rightfully applies to the species hitherto known as C. kalbreyeri .

The specimen “Flora Falcon 302” has small sori (0.8 mm) with only few sporangia (10–15) and rather few, short paraphyses. In contrast, luxuriant plants of Cyathea farinosa have a regular veil of paraphyses in sori that contain about 35–40 sporangia. Indument and receptacle size as well as the dissection pattern of the fronds are, however, in both cases the same. Meier & Hernández-Bretón 10023 has very small pinnules that resemble those of C. venezuelensis A.R.Sm. ex Lellinger (1987[1988]: 94); they are nonetheless characteristically long stalked.

The habit of Cyathea farinosa is probably identical to many other species of the genus, with relatively slender trunks that are covered in old petiole bases and with the fronds more ascending rather than drooping, like C. pungens (Willdenow 1810: 206) Domin (1929: 263) . This is deducted from the thickness of the rachises and the angles between them and the costae. This habit would differ greatly from that of C. gibbosa (= C. kalbreyeri auct .), which is similar in its frond dissection, but has thicker trunks that bear very long fronds that are supported over adjacent vegetation. Cyathea farinosa further differs in its abaxially stramineous to yellow axes (vs. dark brown to atropurpureous in C. gibbosa ), the indument of bullate squamules and peltately attached brown, ovate lanceolate scales (vs. indument absent to sparse, lacking larger scales or bullate squamules) and the typical white tortuous hairs (present and soon caduceus in both species, but much more abundant in C. farinosa ).

Cyathea gibbosa (Klotzsch) Domin (1929: 262) . Alsophila gibbosa Klotzsch (1844: 542) . Sphaeropteris gibbosa

(Klotzsch) Tryon (1970: 20). Trichipteris gibbosa (Klotzsch) Barrington (1976: 3) . Type:— GUYANA. “Im

britischen Guiana,” without date, Schomburgk 1124 (holotype B-fragment GH,-fragment NY, isotypes BM, BR, K,

P). Figs. 7 , 8 .

Cyathea surukunensis Marcano (1990: 220) . Type:— VENEZUELA. Bolivar: Near Río Surukún , near Peray-tepuy , 985 m, 16 March 1987, Marcano 209 (holotype VEN, isotypes P, UC).

Trunks (0.35–)0.50–2.00(–4.00) m tall, 12–20 cm diameter, old petiole bases not shed but rotting off the trunk, upper trunk parts clothed in concolorous brown to auburn scales, similar to petiole scales, epidermis dark brown to blackish, mostly obscured by scales; sheath of adventitious roots at the bases usually thin/not massive; apices hidden in fascicle of petioles, unfolding only one crosier at a time. Petioles to 300 cm long, inermous to muricate, rarely with few short spines to 3 mm long, pale to dark brown, sometimes atropurpureous or basally black; without adventitious (aphlebioid) pinnae at the petiole bases. Petiole scales narrowly lanceolate 20–25(–28) × 1.2–2.0(– 2.2) mm, their tips straight or weakly twisted, shiny auburn to dark reddish brown, concordantly bicolorous with very narrow, inconspicuous yellowish to white margins; petiole scurf of small crested auburn to reddish brown squamules only on crosiers and unfolding fronds, usually absent in most parts when fronds unfolded. Fronds 200– 500(–700) cm long; in forest understories strongly ascending as crosiers until finding support on branches of surrounding trees, laxly drooping after that; freely pendent on cliffs with short petioles; pinnae start to unfold only when rachis is fully expanded. Laminae 180–500 × 150 cm, bipinnate-pinnatifid, firm herbaceous to chartaceous, apex gradually to abruptly reduced, non-conform, drooping; glossy dark green adaxially, often plumbeous when dried, paler green abaxially. Pinnae to 75 cm long, 12–16(–20) pairs per frond, long stalked to 5.0 cm, distally narrowly green-alate, the distal segments decurrently adnate. Frond axes vinaceous to reddish brown adaxially, pale brown to brown, rarely stramineous or yellowish abaxially, in smaller plants sometimes dark castaneous to atropurpureous on both sides; completely glabrous abaxially, hairy only adaxially on costules, costae and distal parts of rachises, hairs 0.5–1.0 mm long, tan to brown; costae smooth, 3–4 mm wide; insertions of costae into rachises swollen, with one inconspicuous pneumathode in the color of the costae, elliptic to 3 mm long. Pinnules 10.0–16.0 × 1.8–3.0(–3.8) cm, the largest ones stalked 3.0–5.0 mm, alternate, 0.5–3.0 cm between the stalks, linearly oblong to long triangular, truncate to weakly cordate at bases, tapering from beyond the middle to long acuminate to short attenuate tips; the yellowish to brown stalks articulate, with a dark ring around their insertions; the segments patent to weakly ascending with finely to coarsely crenate margins, especially at their shortly acute, falcate tips; basal segments usually opposite to alternate, the lowest ones not remote from each other, sinuses narrow, 0.5–1.0 mm wide, rarely in larger pinnules to 5 mm wide, acute. Veins pale brown to yellowish, contiguous with the identically coloured hyaline segment margins, mostly glabrous adaxially except for occasionally single multicellular hairs on the midribs; abaxially glabrous except for occasional dark brown subbullate squamules to 0.5 mm long on the veins; sterile veins forked or simple, fertile veins forked. Sori (0.7–)1.0– 1.2 mm diameter, inframedial to medial, deep brown, indusia absent, receptacles globose, 0.3–0.4 mm diameter; paraphyses of the same length as or shorter than sporangia (0.3–0.6 mm). Spores not examined.

Distribution and habitat: — Colombia, Venezuela, Ecuador, Peru and Bolivia at 900–1450 m, in the understory of moist montane forests, probably restricted to limestone and sandstone. May also occur in Brazil, e.g. in the Neblina Range at the border to Venezuela and the Serránia de Huanchaca at the border to Bolivia.

Additional specimens examined: — COLOMBIA. Antioquia: San Luis, vía Medellin-Bogotá, quebrada La Tebaida , 06°08'N, 75°10'W, 1010–1060 m, 22 June 1987, Callejas et al. 4000 ( UC, US) . — VENEZUELA. Bolivar: 17 km E of El Pauji by road and 64 km W of Santa Elena by road, 4 km N of highway, Río Las Ahallas , 04°30'N, 61°30'W, 850 m, 31 October 1989, Liesner 19204 ( UC) ; Gran Sabana, ca. 10 km SW of Karaurin Tepui at junction of Río Karaurin and Río Asadon ( Río Sampa ), 05°19'N, 61°03'W, 900–1000 m, 22 April 1992, Liesner 23578 ( UC) . Lara: Morán, Caserio Altamira , 09°26'N, 69°47'W, 1200 m, 17 April 1991, Rivero 1155 C ( UC) . Portuguesa: Guanare, ESE of Paraíso de Chabasquén, along road to Cordoba, ca. 20–25 min from Chabasquén , 09°23'N, 69°54'W, 1300–1500 m, 11 June 1986, Smith et al. 1009 ( UC) ; Ospino, Cerro Sabana , 1200–1500 m, 12 July 1991, Ortega, Diáz & Rosales 3115 ( UC) ; Unda, 20 km NE of Chabasquen, ridge El Helechal , 09°03'N, 69°59'W, 1600– 2000 m, 9 August 1987, Ortega et al. 1746 ( UC) . Tachira: Lobatera, La Cazadora , 2000 m, 23 July 1987, van der Werff & Ortiz 5485 ( UC) ; Uribante, surroundings of camp Siberia ( CADAFE) , 08°01'N, 71°43'W, 1200 m, 20 November 1989, Ortega & van der Werff 2815 ( UC) . Trujillo: Boconó, P. N. Guaramacal, between camp in Quebrada Amarillo above the Río Amarillo ( W of bridge over river and Mesa de Cunaviche ) and a mesa-like area above karst outcrops, 1011215 N –1011252 N, 19-269899 E –19-369690 E, 1200– 1200 m, 4 January 2005, Dorr & Stergios 8850 ( UC) .— ECUADOR. Morona-Santiago: Gualaquiza. Hilltops near the town of Gualaquiza , 03°24'S, 78°34'W, 1100 m, 30 July 1997, Fay & Fay 4177 ( AAU, MO, QCNE, UC) . Sucumbios: Sinangoe Station, Shishicho ridge, Alto Aguarico drainage, above ( S of) Río Cofanes , W of Puerto Libre, NW of Lumbaqui, access from Rio Sieguyo , ridgeline trail above camp, 00°12'01.3"N, 77°31'54.3"W, 1300–1450 m, 15 August 2005, Aguinda, Pitman & Foster 1321 ( UC) . Zamora-Chinchipe: Hill about 2 km downstream from campamento Shaime along Río Nangaritza , 900–1200 m, 16 February 1998, van der Werff et al. 13047 ( AAU, QCNE) ; in the vicinity of the mining camp at the Río Tundaime , 03°34'44"S, 78°24'11"W, 1100–1400 m, 10 November 2008, van der Werff et al. 19421 ( UC) ; within 3 km of the town of Zamora , 04°03.5'S, 78°57.5'W, 1000 m, 11 July 1998, Fay & Fay 4386 ( AAU, MO, QCNE, UC) , 19 July 1998, Fay & Fay 4453 ( AAU, QCNE) , Fay & Fay 4477 ( AAU, QCNE) ; Hills and pastures immediately S and SE of Zamora, 04º04'S, 78º57'W, 1000–1250 m, 14 June 1988, Øllgaard et al. 74875 ( AAU, LOJA) .— PERU. Junin: Satipo, Gran Pajonal, E of Chequitavo on trail to Kotampaz , 10°45'S, 74°23'W, 1200 m, 5 April 1998, Smith 6749 ( UC) . San Martin: Zepelacio, near Moyobamba , 1200–1600 m, October 1933, Klug 3256 ( K) ; “ In monte Campana, prope Tarapoto , Peruvia orientalis,” August 1856, Spruce 4331 ( BM, K) , Spruce 4694 ( BM) ; along road Rioja-Pedro Ruiz, about bridge Serranoyacu , 05°40'26"S, 77°40'35"W, 1170 m, 6 March 2005, van der Werff et al. ( UC) .— BOLIVIA. La Paz: Larecaja, Region de Maipiri , San Carlos , ca. 15°20'S, ca. 68°15'W, 850 m, 10–19 April 1927, Buchtien 289 ( BM, K, NY, UC, US). Santa Cruz: Prov. Velasco , Parque Nacional Noel Kempff Mercado , Campamento Las Gamas , 14°48'41"S, 60°23'45"W, 900 m, 29 March 1997, Arroyo & Killeen 180 ( LPB, MO) .

Remarks: —The name Cyathea gibbosa is applied here to the species previously known as C. kalbreyeri . What was previously known as C. gibbosa is typified by C. farinosa . The true C. kalbreyeri represents a previously unrecognized species, which shows quite a different habit in the field than C. farinosa or C. gibbosa . The habit in the field makes Cyathea gibbosa (hitherto C. kalbreyeri ) unmistakable. The long, strongly ascending fronds are supported on high branches of trees, from where the distal parts hang free. The species does not invest much in the trunks, which are mostly less than 2 m tall (a report of a 7 m tall plant (Rivero 1155) may refer to the total height including the fronds). This can be seen as adaptation to growing in the understory where long leaves reach better light conditions quicker than a long trunk. Sometimes the fronds are freely hanging from cliffs, in which case the petioles are much shorter (personal observation). Many plants from the Venezuelan Cordillera de la Costa have rather narrow but long stalked pinnules.

Scattered through the Andes, Cyathea gibbosa is locally frequent but apparently absent over large stretches. The species is well known in southern Ecuador and northern Peru from areas with limestone and sandstone, but is absent from nearby quartz areas.

The synonym Cyathea surukunensis was described from the Venezuelan part of the Pakaraina massif bordering Guyana, which is remote from most known populations of C. gibbosa in the Andes, but close to the type locality. It differs apparently in the petiole scales (not seen; described as having cretaceous borders, which C. gibbosa usually lacks) and the paler petioles and rachises (yellowish brown to stramineous vs. dark brown to atropurpureous). I have come across Andean specimens of C. gibbosa with similar pale petioles and axes. They differ further from typical C. gibbosa in having spines on the petiole (vs. inermous), but have the same habit with short trunks and large ascending fronds. In contrast to the description of C. surukunensis , these plants lack cretaceous scales margins. Irregularly developed sori with partially aborted sporangia and spores of variable size indicate that this aberrant phenotype may be a hybrid. Similarly, C. surukunensis may represent another hybrid between C. gibbosa and an unknown species, in which case the name would deserve taxonomic recognition as a nothospecies.

Cyathea gracilis Grisebach (1864: 704) . Type :— JAMAICA. Fox Gap, without date, Purdie s.n. (holotype K). Fig. 9 .

Trunks 0.2–1.0(–2.0) m tall, 4–10(–15) cm diameter, with or without old petiole bases, upper parts covered in scales, similar to petiole scales; adventitious roots up near the apex; apex not hidden in fascicle of petioles. Fronds 150–350 cm long, drooping. Petioles to 150(–200) cm long, inermous to sparsely muricate, dark castaneous to atropurpureous, basally black; without adventitious (aphlebioid) pinnae at the petiole bases. Petiole scales lanceolate to ovate-lanceolate, 15–20(–25) × 2–3 mm, their tips undulated or twisted, shiny dark brown to atropurpureous, concolorous or bicolorous with a narrow, paler brown to yellowish margin; petiole scurf absent. Laminae to 250 × 50–80 cm, bipinnate-pinnatifid to tripinnate, firm herbaceous to chartaceous, apex gradually reduced and drooping; dark green adaxially, often blackish when dried, bright green abaxially. Pinnae to 45 cm long, 8–10 pairs per frond, long stalked 2.0–4.0(–5.0) cm, distally narrowly green alate, the distally segments decurrently adnate. Frond axes brown to castaneous, rarely pale brown or atropurpureous, often shiny when darkcolored, hairy only adaxially on costules and distal parts of costae and rachises, hairs 0.5–1.0 mm long, tan to brown, without hairs abaxially, scurf absent, rarely with small brown scales 0.5–1.0 mm long from the curled-up crosier stage persisting in junctures of costae with costules and rachis; costae smooth, rarely more than 3–4 mm wide, insertions of costae into rachises with two separate pneumathodes to 2 mm, in the color of the costae. Largest pinnules (7.0–)9.0–12.5(–14) × 1.8–3.0(–5.0) cm, stalked 3–6 mm, alternate, 1–2 cm between the stalks, long triangular to ovate lanceolate, truncate to weakly rounded at base, tapering from below the middle to short to long attenuate tips; the stalks and the costules not contrasting with the costae, of the same color or gradually transient to a paler brown; the segments patent to weakly oblique with finely to coarsely crenate margins and rounded to obtuse tips; basal segments opposite to alternate, the lowest ones sometimes remote from each other, sinuses narrow to wide 2–3(–5) mm, obtuse to rectangular. Veins pale brown to yellowish, contiguous with the identically colored hyaline segment margin, mostly glabrous adaxially except for occasionally single multicellular hairs on the midveins; abaxially glabrous except for occasional brown unicellular trichomidia and small, auburn to brown squamules; the latter vary from flattish to bullate with entire margins; sterile veins forked once or simple, fertile veins forked. Sori (1.0–)1.2–2.0 mm diameter, subproximal (to medial), indusia deeply urceolate to subsphaeropteroid, sometimes sphaeropteroid with weak apical umbo, tan to auburn, not translucent, fragile but persisting as fragmented cups, receptacles globose, to 0.6 mm diameter; paraphyses shorter than sporangia (ca. 0.2 mm). Spores not examined.

Distribution and habitat: — Costa Rica, Panama, Jamaica, Colombia, Ecuador and northern Peru at (770–) 1300– 2500 m; mainly growing epiphytic, sometimes terrestrial in open, humid places.

Additional specimens examined:— COSTA RICA. Cartago: N and S slopes of ridge on E side of Río Grande de Orosi, opposite mouth of Río Humo , ca. 6 km upstream from Tapantí, 09°43'N, 83°47'W, 1500–1800 m, 25 November 1988, Grayum, Beach & Sleeper 4546 ( MO, UC) . Guanacaste: Reserva Santa Elena, Monteverde , 24 November 1998, Bittner 2296 ( AAU) . Heredia: Along Río San Rafael, Atlantic slope of Volcán Barva , 10°13'N, 84°05'W, 1500 m, 13 April 1990, Grayum 7020 ( AAU, MO) . Puntarenas, Santa Elena Rainforest, 6 km NE of Santa Elena, 10°21'N, 84°07'W, 1600–1725 m, 20 May 1996, Moran & Rohrbach 5874 ( NY) . Puntarenas- Alajuela: In and around the Monteverde Nature Reserve, mostly on the Pacific watershed , 10º18'N, 84º47'W, 1450–1650 m, 31 October–2 November 1975, Burger & Baker 9716 ( US) . San José: Finca Haberl near San José , 22 December 1912, Brade & Brade 108 ( NY, UC) ; above La Hondura , 1510 m, 28 December 1971, Gastony & Gastony 772 ( F) .— PANAMA. Panamá: Veraguas, near summit of Cerro Arizona, above Santa Fé , 1400 m, 23 April 1984, Hammel & Kress 8564 ( F) . Veraguas: Cerro Tute , summit, ca. 3 km N of Santa Fé, 08°31'N, 81°06'W, 1453 m, 7 February 1992, Moran 4065 ( MO, UC) .— JAMAICA. St. Andrew Parish, Mt. Horeb trail, Fairy Glade , 1250 m, 25 June 1967, Crosby, Hespenheide & Anderson 316 ( UC) ; near Cinchona , 1495 m, 20 March 1913, Watt 184 ( UC) ; below New Haven Gap , 1500–1600 m, 3 October 1920, Maxon & Killip 944 ( F, GH) . — COLOMBIA. Antioquia: Río Guatape , 2347 m, 25 February 1880, Kalbreyer 1465 ( COL) ; 8 km W of Valdiva , 1700 m, 11 March 1972, Madison 803 ( GH) ; Ciudad Bolivar, Corregimiento San Bernardo de Farallones, Cerro San Nicolas , Farallones del Citará , sector La Peña , 05°45'25.11"N, 76°03'10.72"W, 2750 m, 10 November 2008, Rodriguez et al. 4677 ( NY) Chocó: San José del Palmar, Cerro del Torrá , E slope, mesa below the peak, 2500 m, 21 August 1992, Silverstone-Sopkin et al. 4593 ( UC) , 25 August 1992, Silverstone-Sopkin et al. 4701 ( UC) .— ECUADOR. Carchi: Approx. 6 km above Maldonado, just below Puente de Palo , transect 2250-1, 00°54'N, 78°06'W, 2275 m, 24 May 1997, Boyle & Bradford 1900 ( AAU, MO) ; border area between Provs. Carchi and Esmeraldas, about 20 km past Lita on road Lita-Alto Tambo , 550 m, 25 June 1995, van der Werff, Gray & Tipaz 11967 ( QCNE, MO, UC) . Morona-Santiago: Limon-Indanza; Cordillera del Condor, Centro Shuar Warints , copper mining concession area, about 8 km SW of Warints, 03°10'59"S, 78°17'57"W, 1800 m, 8 October 2006, Neill et al. 14111 ( MO, UC, QCNE) . Napo: Carretera Baeza-Mera, cerca P. N. Antisana , 00º38.9'S, 77º48.2'W, 9 July 2002, Lehnert 183 ( GOET, QCA, UC) ; Cerro Huacamayos, on road Baeza-Tena, ca. 34 km from Baeza , 00º41'S, 77º50'W, 2000 m, 9– 10 August 1980, Øllgaard, Roth & Sperling 35827 ( AAU, QCA) . Pastaza: Road N of Mera, towards Río Anzu, Km 6.7, 01º27'S, 78º04'W, 1480 m, 9 December 1997, Øllgaard & Navarrete 2801 ( AAU, QCA) ; Mera, road and muletracks to approx. 4 km N of the village (along Río San Jorge and Río Tigre ), 01°35'S, 77°53'W, 1200 m, 1–2 September 1976, Øllgaard & Balslev 9201 ( AAU, F) . Zamora-Chinchipe: Nangaritza; faldas de la Cordillera del Condor, sendero desde Pachicutza hacia "El Hito", 04°07'S, 78°34'W, 1500–1600 m, 6 December 1994, Palacios & Neill 6551 ( MO, QCNE, UC) .— PERU. Amazonas: Bongara, Laguna de Pomacochas , 2550 m, 27 March 1998, van der Werff et al. 15823 ( UC) . Pasco: Oxapampa, Dist. Palcazú, San Francisco de Pichanaz , 10º29'16"S, 75º03'58"W, 770 m, 28 February 2004, Mellado, Monteagudo & Rodriguez 894 ( MO) .

Remarks: —The distinction between Cyathea gracilis and C. ebenina has often been based solely on the colour of the pinnule stalks and its contrast to the costae. Since colours are highly influenced by subjective perception, the two species have often been confounded. Nevertheless, both are distinct and ecologically separated. Cyathea gracilis is the only scaly tree fern that regularly grows as an epiphyte; rarely it occurs on road cuts or shaded cliffs where its fronds may hang free, since the trunk is often shorter than the frond length. Trunks are usually less than 1 m tall, fertile fronds are developed from a trunk size of 20 cm on. The colours of trunk, petiole scales and petiole epidermis are more or less monochrome brown to dark brown, often weakly shiny. Opposed to this, C. ebenina is always found as a terrestrial in the forest understory. Its adaptation to this shady environment is to develop a tall trunk quickly. It does this by developing a thin trunk with spirally arranged fronds instead of distinct whorls. The trunks are usually not thicker than a broomstick and inconspicuous in the field, although they are covered in shiny scales that are bright orange brown with a weakly to strongly contrasting, darker central stripe. The same scales are found at the bases of the petioles, where they strongly contrast with the atropurpureous to black, shiny epidermis. Conditioned by their growth habits, both species have only few green fronds (ca. 3–5).

One specimen from Colombia, Dept. Chocó (Silverstone-Sopkin et al. 4593, UC) may represent a luxuriant form of Cyathea gracilis with shiny dark castaneous frond axes, broad, orange scale margins and long stalked pinnae and pinnules (to 5.0 and 2.0 cm long, resp.). The general colour resembles more C. ebenina , but the pinnule stalks are not contrasting with the costae, a character of C. gracilis . The sori are mainly medial, well developed (to 1.5 mm diameter), with functional sporangia and well-formed spores, arguing against a hybrid origin with both mentioned species as parents.

Another plant from roughly the same area ( Colombia, Dept. Valle, Gentry et al. 53210, UC; first indentified as C. divergens ) has similarly long-stalked large pinnules (14 × 5 cm) and almost concolorous blackish brown petiole scales with narrow pale brown margins. Here, the general coloration resembles more Cyathea gracilis (dark brown frond axes with non-contrasting pinnule stalks), but the proximal placement of the sori and the scale colour are reminiscent of C. aemula and C. divergens . However, the pale petiole scurf characteristic of these species is missing. The aborted sori of this specimen may be an indication of hybrid origin, potentially between C. gracilis and either C. aemula or C. divergens .

Cyathea hemiepiphytica Moran (1995: 56) . Type:— ECUADOR. Esmeraldas: Road Lita-Alto Tambo, km 18, perhumid forest on plateau, 00.47' N 78.30 ' W, 900 m, 13 January 1991, Øllgaard et al. 98772 (holotype AAU, isotypes K, QCA, QCNE, UC, US) .

Plants terrestrial, hemiepiphytic or epiphytic. Trunks to 5 m tall, ca. 4 cm diameter; without old petiole bases, most parts covered in bicolorous brown to dark brown scales with paler margins, similar to petiole scales; epidermis dark brown to blackish, mostly obscured by scales; adventitious roots all along the trunk; apices not hidden in fascicle of petioles, unfolding only one crosier at the time. Fronds to 200 cm long, distantly placed along the trunk, patent, weakly arching. Petioles 30–70 cm long, inermous to verrucate, dark brown to dark castaneous, basally black; without adventitious (aphlebioid) pinnae at the petiole bases. Petiole scales lanceolate, 15–20(–22) × 3 mm, their tips straight or weakly twisted, shiny auburn to dark brown, discordantly bicolorous with yellowish to white margins, with brown marginal streaks and black insertions; petiole scurf of small dissected auburn to pale brown appressed squamules, erect multicellular hairs 0.2–0.5 mm long and appressed unicellular trichomoidia deciduous, usually absent in most parts if fronds unfolded. Laminae to ca. 100 × 80–90 cm, triangular, bipinnate-pinnatifid, firm herbaceous to chartaceous, apices gradually reduced; glossy dark green adaxially, often plumbeous when dried, pale grey-green abaxially. Pinnae to 45 cm long, ca. 8 pairs per frond, sessile to short stalked to 0.4–1.5 cm, distally narrowly green alate, the distal segments decurrently adnate. Frond axes brown to dark brown or castaneous, adaxially darker than abaxially; hairy adaxially on costules, costae, central and distal parts of rachises, hairs 0.5–1.0 mm long, tan to brown; abaxially completely glabrous or short pubescent with white multicellular hairs, costae smooth, 2–3 mm wide; insertions of costae into rachises weakly swollen abaxially, with two well separated inconspicuous pneumathodes, dark brown, the apical one elliptic to 23 × 1 mm, the basal one isodiametric to 1 mm. Largest pinnules 5.0–9.0 × 1.5–1.8(–2.0) cm, sessile to stalked to 3 mm, alternate, 0.5–1.5 cm between the stalks, linearly oblong, truncate to weakly cordate at bases, tapering from beyond the middle to short attenuate tips; the yellowish to brown stalks articulate, with a dark ring around their insertions; the segments patent to weakly ascending with finely crenate margins, especially at their obtuse tips; basal segments usually opposite, the lowest ones not remote from each other, sinuses occluded by overlapping segment margins, or very narrow, 0–0.2 mm wide, acute. Veins pale brown to yellowish, contiguous with the identically coloured hyaline segment margins; glabrous adaxially, abaxially glabrous except for occasional dark brown subbullate squamules to 0.5 mm long on the veins; sterile and fertile veins simple, rarely forked. Sori 0.7–1.0 mm diameter, proximal to subproximal, sitting on the back of a vein, orange-brown, indusia absent, receptacle globose, 0.2–0.3 mm diameter, paraphyses much shorter than sporangia (0.2–0.3 mm), straight, hyaline, white to pale brown. Spores not examined.

Distribution and habitat:— On the western Andean slopes of southern Colombia and northern Ecuador at 550–1950 m, in wet tropical montane forests of the Chocó region.

Additional specimens examined:— COLOMBIA. Chocó: San José del Palmar, 2 km E of San José del Palmar , 1550–1650 m, 27 March 1975, Lellinger & de la Sota, 767 ( COL, US). San José del Palmar , Cerro del Torrá , E slope, trench above heliport, 1920–1950 m, 8 September 1992, Silverstone-Sopkin et al. 4192 ( UC). Nariño: Ricaurte , El Palmar, bank of Guiza river , 1372 m, 18 August 1965, Soejarto 1449 ( COL, GH, US) .— ECUADOR. Carchi: San Juán valley, 4 hour walk below Chical, at Ortiz ranch between Peñas Blancas and El Pailón, known locally as Gualtal , above San Juán , 01°02'N, 78°15'W, 1230–1250 m, 11 June 1997, Boyle , Butler & Lloyd 1991 ( AAU, QCNE); San Marcos de los Coaiqueres, on trail towards Gualpi Bajo, 01°06'N, 78°17'W, 1000 m, 8 February 1989, Øllgaard et al. 57508, AAU, QCA); border area between Prov. Carchi and Esmeraldas, about 20 km past Lita on road Lita-Alto Tambo, 00°54'N, 78°30'W, 550 m, 24 June 1991, van der Werff, Gray & Tipáz 11951 ( MO, UC). Esmeraldas: Road Lita-Alto Tambo, Km 17.5, entrance of Reserva Ecologica Cotacachi-Cayapa, 00°53'N, 78°83'W, 820 m, 5 November 1998, Øllgaard & Navarrete, 105424, ( AAU, QCNE), Km 16.4, Río Anchayacu , 00°53'W, 78°32'W, 820 m, 6 November 1998, Øllgaard & Navarrete 105470 ( AAU), Km 18, on plateau, 00°53'N, 78°83'W, 820 m, 14 January 1995, Øllgaard, Korning & Krogstrup 98772 ( AAU) .

Remarks: — Cyathea hemiepiphytica starts its life as a terrestrial plant, but its trunk usually inclines and leans against a tree where it climbs further with its adventitious roots ( Moran 1995). Apart from the distinctive habit with distantly placed fronds and triangular laminae, C. hemiepiphytica is easily recognized by the rounded, closely placed segments of the pinnules and the small, proximal sori that are sitting on the back of unforked veins. Cyathea gibbosa (hitherto C. kalbreyeri ) and C. ulei have medial to subproximal sori. Cyathea gibbosa further has wider, usually triangular sinuses between the segments and the fertile veins are forked. The outline of the pinnules of C. hemiepiphytica with the relatively short, round segments resembles more C. ebenina , C. ulei and C. tuerckheimii , rather than C. divergens , C. gracilis or C. aemula , whose segments are often relatively long and have acute tips.

Cyathea × jurgensenii Fournier (1872: 135), stat nov.

Cyathea fulva ( Martens & Galeotti 1842: 78) Fée (1857: 34) × C. tuerckheimii E.Fourn. = Cyathea divergens var. tuerckheimii (Fourn.) R.M.Tryon × C. fulva (M.Mart. & Galeotti) Fée ( Tryon 1976: 90) . Type:— MEXICO. Without locality, without date, Jürgensen 874 (holotype P, isotypes NY [fragment], US).

Trunks to 2 m tall, to 8 cm diameter; other trunk characters unknown, but presumably similar to C. tuerckheimii . Fronds to 400 cm long, arching. Petioles 40 cm long or more, strongly aculeate, brown, proximally dark brown; without adventitious (aphlebioid) pinnae at the petiole bases; petiole scales narrow lanceolate to lanceolate, 18–25 × 1.8–2.5 mm, bicolorous brown with narrow pale brown to whitish margins; petiole scurf well developed and persistent but not dense, erect squamules 0.5–2.0(–3.0) mm long, the larger ones lanceolate, uniformly brown to dark-brown or with a tan to white margins, smaller ones ovate to round, uniformly brown, with fimbriate to setulose margins; undercoat of smaller appressed, hair-like squamules sparse or lacking. Laminae to 350 × 100– 130 cm, bipinnate-pinnatifid to almost tripinnate, firm herbaceous to chartaceous, apices gradually reduced, not drooping; dark green adaxially, but not blackish when dried, grey green abaxially. Largest pinnae 50–65 cm long, stalked to 2.0–4.0 cm, distally narrowly green alate, the distal pinnules/segments decurrently adnate. Frond axes pale brown to brown, costae and costules sometimes stramineous; hairs only adaxially on costules, costae and distal parts of rachises; hairs 0.5–1.0 mm long, whitish to tan; frond axes abaxially with sparse brown scurf consisting of scattered squamules 0.2–0.6 mm long, larger ones ovate-lanceolate, flat with dentate or short fimbriate margins, smaller ones narrow and strongly dissected, grading into short branched hairs; scurf persisting in junctures of costae with costules and rachises; costae muricate to smooth, 2–4 mm wide. Pinnules (10–)11–15 × (1.3)– 2.2–3.5 cm, the largest ones subsessile to short stalked (1–5 mm), alternate, 0.6–1.5(–2.4) cm between the stalks, oblong to long triangular, truncate to weakly cordate at bases, tapering from the middle, rarely from the base, to short attenuate tips, the segments weakly falcate, margins finely crenate to crenate-serrate, tips obtuse to acute, rarely rounded; basal segments alternate to opposite, sometimes remote from each other but connected by a green wing; sinuses narrow (1–2 mm), obtuse to acute; sterile pinnules not or only slightly broader than fertile ones. Veins adaxially glabrous, abaxially glabrous or with single white multicellular hairs to 1 mm long and some small flattish, brown to dark brown ovate squamules with elongated tips to 2 mm long; few finely dissected squamules present, dark brown, often appearing as hairs with short branches; subbullate or bullate squamules lacking; sterile veins forked or simple, fertile veins forked. Sori 0.6–1.0 mm diameter, proximal to submedial, indusia sphaeropteroid, with umbo, tan, translucent, often all fragments persisting; paraphyses shorter than or of the same length as sporangia; sori may be irregularly distributed and quite often aborted (empty indusia). Spores not examined.

Distribution and habitat: — Mexico, El Salvador and Honduras, at 1300–2000 m in moist montane forests; to be expected from Belize and Guatemala.

Additional specimens examined:— MEXICO. Chiapas: La Trintaria, E of Laguna Tzikaw, Monte Bello National Park, 16°04'43"N, 91°39'32"W, 1300 m, 13 May 1973, Breedlove 35209 ( F, MO) ; without locality, 1907, Munch s.n. ( BM) . Veracruz: 20 km antes de Huatusco, carretera Puente Nacional a Huatusco, Nevling & Gómez- Pompa 2443 ( GH) .— EL SALVADOR. Santa Ana: Volcán Santa Ana, 2000 m, 5 April 1987, Seiler 276 ( F) .— HONDURAS. Francisco Morazán: Above San Juancito, San Juancito mountains , 1800 m, 24 March 1951, Williams 17543 ( F) .

Remarks: —The morphology of the scurf indicates that Cyathea × jurgensenii is a hybrid between C. tuerckheimii and C. fulva . The low density and the dark brown colour found in the squamules are referable to C. fulva , while the general shape and the white colour of the squamules are typical of C. tuerckheimii . Also, sori are often aborted and the indusia are inflated and empty, or appear as hoses of unexpanded tissue. The hybrid was recognized by Tryon (1976) and discussed on one specimen, but was not connected with the type of C. × jurgensenii.

Monterrosa & Monro (2008) cited Seiler 276 (F) as specimen for Cyathea tuerckheimii (as C. divergens var. tuerckheimii ).

Cyathea kalbreyeri (Baker) Domin (1929: 262) . Alsophila kalbreyeri Baker (1894: 9) , nom. nov. for Alsophila podophylla Baker (1881: 202) not Alsophila podophylla Hooker (1857: 334) . Alsophila kalbreyeri Christensen (1905: 44) , nom. superfl. Trichipteris kalbreyeri (Baker) Tryon (1970: 45) . Type:— COLOMBIA. Antioquia: San Carlos, [ca. 06°11'N, 74° 59'W], 4500 ft, 27 January 1880, Kalbreyer 1375 (holotype K, isotype B-fragment GH n.v.-fragment NY). Fig. 10 .

Trunks to 5 m tall, slender. Fronds ca. 2 m long, stiff, ascending, presumably distally drooping-arching. Petiole scales lustrous brown. Laminae ovate-elliptic with gradually reduced apex, basal pinnae ca. half the length of the largest pinnae, notably reflexed. Rachises glossy castaneous, adaxially glabrescent with few brown multicellular hairs less than 1 mm long, abaxially glabrous. Pinnae to 65 cm long, deltate-lanceolate, stalked to 5.5 cm, gradually reduced to a pinnatifid section, less than 10 pairs per frond. Largest pinnules to 150 × 25 mm, lanceolate, basally cuneate to round, often inequilateral, stalked 6–12 mm, apically attenuate, costules yellowish green, raised adaxially and abaxially, abaxially glabrous, adaxially glabrous or with few multicellular hairs to 1 mm long. Segments oblong to elongate, to 20 × 8 mm, ascending, straight to weakly falcate, with round to obtuse apices, sinuses very narrow, usually occluded, segment margins subentire along the sinuses, crenulate to serrulate at the apices; midveins weakly raised, yellowish, with few multicellular hairs adaxially, otherwise glabrous, lateral veins weakly raised to flat, green, forked, rarely simple. Sori medial to inframedial, exindusiate, 0.8–1.0 mm diameter, sitting in forks of veins or rarely on a branch beyond the fork, receptacles round, 0.3–0.4 mm diameter, paraphyses short (ca. 0.2 mm), not surpassing sporangia. Spores not examined.

Distribution and habitat: —Only known from the inter-Andean valley in Dept. Antioquia, Colombia, at 1500 m in moist montane rainforest with Caribbean influence.

Remarks: —The type specimens lack petiole material, but it can be said with certainty that it does not represent the species commonly recognized as Cyathea kalbreyeri (Barrington 1978) . The trunk is described on the label as “thin, erect 10–15 ft ”and the fronds as “stiff, 6 ft ”; both do not match the observations made in the field of recently collected plants, which have very long fronds to 7 m long and shorter, rather thick trunks. The rachis and the costae of the type material are dark castaneous to blackish and shiny and contrast with the yellowish green costules, resembling in this regard C. ebenina . The segments are ascending to falcate and round to obtuse, not acute as it is to be expected of pinnules of that size in what is currently addressed as C. kalbreyeri .

I have come to the conclusion that Cyathea kalbreyeri represents a distinct species previously unrecognized in taxonomic treatments and until now is only known from the type locality. The available material differs from C. ebenina mainly in the indusia (exindusiate in C. kalbreyeri vs. indusia sphaeropteroid and persisiting as fragments in C. ebenina ) and the petiole scales (reported as dark brown vs. orange-brown, concolorous or with dark brown central stripe).

The revision of the types of other species similar to Cyathea kalbreyeri reveals that what currently is known under this name should accurately bear the name C. gibbosa . Plants that are currently recognized as C. gibbosa (Barrington 1978) are correctly called C. farinosa .

Cyathea kalbreyeri resembles C. nigripes ( Christensen (1905: 45) Domin (1929: 263) and C. nodulifera Moran (1991: 88) , two rare species that may be present at the type locality. Cyathea nigripes differs in having mainly sessile pinnules with truncate to cordate bases (vs. stalked with mostly cuneate base in C. kalbreyeri ). Cyathea nodulifera differs in the presence of prominent aerophores at the bases of costae and costules (vs. weakly developed, flat). Cyathea kalbreyeri further resembles C. aemula from the eastern Andean escarpment in southern Colombia and Ecuador. The main differences lie in the length of the paraphyses (short in C. kalbreyeri vs. long and contorted in C. aemula ), the presence of scurf (absent vs. white on petioles and frond axes) and coloration of the frond axes (dark vs. pale).

Cyathea latevagans (Baker) Domin (1929: 262) . Alsophila latevagans Baker (1881: 203) . Trichipteris latevagans (Baker) Tryon (1970: 45) . Type:— COLOMBIA. Antioquia: Without locality, without date, Kalbreyer 1327 (holotype K). Figs. 11 , 12 .

Trunkless, rhizome short-creeping or short-ascending, covered with old petiole bases. Petioles to 40 cm long, inermous or sparsely verrucate, dark atropurpureous to black; petioles basally with two lines of large elliptic to ovate, pale brown to stramineous aerophores to 5.0 × 1.0– 2.5 mm; without adventitious (aphlebioid) pinnae at the bases; petiole scales broadly lanceolate to ovate, 5–10(–12) × 3.0–4.0(–4.5) mm, their tips straight, shiny dark brown, concordantly bicolorous with very narrow, inconspicuous yellowish to pale brown margins, pseudopeltately attached; petiole scurf absent. Fronds 80–200 cm long, scrambling or dangling. Laminae 56–130 × 14–22 cm, long triangular to lanceolate, pinnate-pinnatifid, firm herbaceous to chartaceous, with truncate bases and gradually reduced, long tapering apices; glossy dark green adaxially, often blackish when dried, paler green abaxially. Frond axes vinaceous to black on both sides; completely glabrous abaxially, hairy only adaxially on costae and distal parts of rachises, hairs 0.5–1.0 mm long, tan to brown, sometimes whitish, then to 2.0 mm long and sparser; insertions of costae into rachises swollen, with one conspicuous pneumathode, golden brown, elliptic ca. 2 × 1 mm. Pinnae 9.0– 17.0 × 1.8–3.0(–3.8) cm, (20–)22–28(–30) pairs per frond, alternate to opposite, the largest ones stalked 6–15 mm, to 2.2–4.5 cm between the stalks, long triangular to oblong, pinnatifid to almost to the costae, truncate to weakly rounded at bases, tapering from the middle to short acuminate to short attenuate tips; the brown stalks articulate; the segments patent to weakly ascending with finely crenate to serrate margins, especially at their obtuse, weakly falcate tips; basal segments usually opposite, the lowest ones not remote from each other; sinuses narrow, 1(–2) mm wide, acute; veins yellowish brown to blackish, contiguous with the identically coloured segment margins; each insertion of the segments’ midveins with the costae weakly raised, bearing a small elliptic pneumathode 2–3 mm long; segments adaxially with few to many curved, whitish to tan, erect multicellular hairs 0.5–1.0 mm long, mainly on the veins near the margins, sometimes hairs equally distributed on and between veins, or completely glabrous; abaxially glabrous except for unicellular trichomoidia, tan to reddish brown, ca. 0.2 mm long and some broadly ovate to almost round, pseudopeltate, auburn to brown scales to 2 mm long, distally on the midveins also some dark brown, bullate squamules; sterile veins forked or simple, fertile veins forked. Sori 0.7–1.0(–1.2) mm diameter, proximal to subproximal, in fork of veins, deep brown, indusia absent, receptacles globose, 0.2–0.3 mm diameter, sometimes supported by a small scale like those on the veins; paraphyses of the same length as or shorter than sporangia (0.3–0.5 mm), straight, brown to red. Spores with verrucate exospore and tuberculate perispore ( Gastony 1979).

Distribution and habitat: — Colombia, Ecuador and Peru in moist tropical montane forests, in open places at 1200–3415 m.

Additional specimens examined:— COLOMBIA. Putumayo: Belmira, Vereda El Yerbal, Los Patos, left margin of Río Chico, Alto de Sabanazo , 06°35'N, 75°32'W, 2700–3000 m, 25 April 1995, Echeverri et al. 459 ( UC) ; Ciudad Bolivar, Corregimiento San Bernardo de los Farallones , Farallones de Citará , sector La Peña, 05°45'25"N, 76°03'11"W, 2750 m, 10 November 2008, Rodriguez-W. et al. 4663 ( NY) . Chocó: Carretera Ansermanuevo-San José del Palmar , limite con el Valle del Cauca, Alto del Galápago , 2100 m, 28 August 1980, Forero et al. 2157 ( COL) ; San José del Palmar , 2750 m, 16 August 1992, Silverstone-Sopkin et al. 4437 ( UC) .— ECUADOR. Carchi: Above Maldonado , 2500 m, 2 August 1993, van der Werff 10831 ( AAU, MO, QCNE, UC) . Zamora-Chinchipe: In the vicinity of the mining camp at Río Tundaime , along trail to mine drilling sites, 03°34'44"S, 78°24'11"W, 1200–1400 m, 11 November 2008, van der Werff et al. 19454 ( MO, UC) .— PERU. Cuzco: Paucartambo, Kosñipata, Trocha Unión , Km 2, 13°06'49"S, 71°36'26"W, 3415 m, 29 July 2003, Garcia-C. et al. 104 ( CUZ, DAV, MO, USM, WFU) . San Martin: Rioja, along road Rioja-Pedro Ruiz, El Mirador, 05°40'29"S, 77°46'25"W, 1850 m, 26 March 2002, van der Werff et al. 15705 ( MO) .

Remarks: — Cyathea latevagans is unmistakable with its long-stalked, pinnatifid pinnae and the dark, shiny frond axes. Other species with similar shiny axes have more complex, i.e. bipinnate-pinnatifid fronds. There are many trunkless species of Cyathea with simply pinnate to pinnate-pinnatifid fronds, but their frond axes are not shiny and the laminae are often pubescent. Additionally, C. latevagans stands out by the presence of the pseudopeltate, almost round scales on the costae; these scales are so far not known from any other species in the genus. Furthermore, the laminae are shiny dark green above and glaucous beneath, a character that is lost in dried specimens.

Cyathea latevagans is possibly related to C. ulei and C. gibbosa . All three have almost glabrous, smooth frond axes that are often shiny and dark and their laminar texture is firm chartaceous to coriaceous. However, the coloration of the fronds of C. latevagans most resembles that of the indusiate C. ebenina . I suspect that there lies more information about the true relationships of the species mentioned here in characters that contribute to the overall similarity than in the presence of indusia.

Cyathea macrocarpa (C.Presl) Domin (1929: 264) . Hemitelia macrocarpa Presl (1847: 41) . Amphicosmia

macrocarpa (C.Presl) Moore (1857: 60) . Type:— BRAZIL. Bahia: Without locality, without date, Blanchet 17

(lectotype PRC, designated by Lehnert 2011: 54). Fig. 13B .

Hemitelia megalosora Trevisan (1851: 164) , nom. nud. (Blanchet 3227 is cited, see below).

Cyathea moricandiana Moore (1861: 272) , nom. nud. (Blanchet 3227 is cited, see below).

Hemitelia moricandiana Hooker (1865: 30) . Type:— BRAZIL. Bahia: Without locality, without date, Blanchet 3227 (lectotype PRC, designated by Lehnert 2011: 54, fragment F, fragment HB n.v, isolectoytpe K-fragment NY,-fragment RB n.v.).

Trunks to 4 m tall, slender, without old petiole bases. Fronds to 210 cm long, arching, distally drooping. Petioles 35 cm long or more, sparsely muricate, dark-stramineous to brown or auburn, matte, without scurf; petiole scales narrowly lanceolate to lanceolate, 8.0–18.0 × 1.5–2.0 mm, concolorous, yellowish white to pale stramineous or tan, basally cuneate, broadly attached, apically long acute, differentiated margins very narrow, only 1–2 cell rows wide, not exerted, without setae or teeth, although sometimes margins tattered with loose cell rows that look like ciliae. Laminae to 130 × 65 cm, bipinnate-pinnatifid, matte dark-green adaxially, pale olive-green abaxially; apices gradually reduced. Rachises inermous, brown to dark stramineous abaxially and adaxially; with scattered brown multicellular hairs to 0.6 mm long adaxially, glabrous abaxially. Pinnae to 35 cm long, short to long stalked to 4 cm, ca. 8 pairs per frond, alternate, weakly ascending; distally green–alate, distal segments strongly decurrently adnate. Costae to 2 mm wide, inermous, dark stramineous to brown abaxially and adaxially; adaxially with few tan to brown multicellular hairs to 0.5 mm long, abaxially glabrous; junctures of costae and rachises weakly swollen, bearing abaxially each one circular and one elliptic, raised pneumathode to 2–3 × 2 mm, yellowish brown to brown, rather conspicuous. Pinnules to 90 × 25 mm, sessile to short stalked to 2 mm, rarely [ Ecuador, Peru] long stalked to 5 mm, 1–2 cm between the stalks/costules, linear-oblong to lanceolate, cuneate basally, rarely truncate or [ Ecuador] weakly cordate with reflexed segments, acute to attenuate apically, ending in shallowly crenate tips; costules pale brown to stramineous, strongly prominent adaxially, weakly so abaxially, glabrous on both sides except for narrowly lanceolate scales to 1–2 mm long, papery, whitish with entire margins; costules basally with inconspicuous pneumathodes, to 1.0 × 0.4 mm. Segments to 12 × 5 mm, deltate, incised only 1/2 to 2/3 towards the costules, sessile, adnate, never remote, weakly ascending, distally weakly falcate, tips obtuse to rounded; sinuses wide, deltate and acute, to 3 mm wide; margins crenulate to finely serrate, in proximal segments basally entire; midveins weakly protruding, brown; lateral veins brown, planar on both sides; sterile veins simple or forked, fertile veins forked. Sori 1.0– 1.2 mm diameter, supramedial to inframarginal, more or less parallel to the margins, usually in the fork of a vein; indusia hemitelioid, large, arching half over the sori, glabrous, tan, papery, matte, firm, margins with short dark teeth; receptacles globose to elliptic, 0.3–0.5 mm long, paraphyses tan to brown, twisted, much longer than the sporangia (0.4–0.6 mm long). Spores not examined.

Distribution and habitat: — Venezuela, Guyana, French Guiana, Ecuador, Peru and Brazil at 50–1100(–2000) m, in lowland rainforest and moist premontane forests.

Selected specimens examined: — VENEZUELA. Amazonas: Atures, Sierra Maiqualida, NW sector, along an upper tributary of Caño Iguana , 05°30'N, 65°15'W, 2000 m, 28 February–3 March 1991, Berry, Huber & Rosales 4884 ( UC) .— GUYANA. Cuyuni-Mazaruni: Paruima, 9 km W, 05º49'36"N, 61º08'40"W, 800 m, 4 July 1997, Clarke 5331 (US). Mabura: West Pibiri compartment, base line back to logging road, 05º20'N, 58º30'W, 17 November 1992, Ek 532 ( UC) . Potaro-Siparuni: Pakaraima Mts, upper Ireng River , 0–300 m upstream from Kurutuik Falls, 05º10'N, 60º13'W, 800 m, 22 October 1994, Henkel, Williams & Mutchnick 6026 ( NY) ; Mt. Ayanganna , east face, plateau above first escarpment, 1 km N of camp, 05º20'19"N, 59º56'46"W, 1100 m, 13 June 2001, Clarke et al. 9135 ( NY) , Kaieteur falls, forest between the falls and Kaieteur summit, 05°11'N, 59°29'W, 550 m, 10 August 1991, Kvist et al. 117 ( US) .— FRENCH GUIANA. Vallée de la crique Paul, au sud-este de la piste "Ostoréro", á 2.5 km au nord de St. - Elie ( Guyane ), 04º50'N, 53º17'W, 110 m, September 1998, Boudrie 3024 ( UC) ; affluents of Crique Amadis-2, Bassin de la Mana, 55 km SSE of Saint-Laurent-du Maroni , 05º02'N, 53º52'W, 50 m, 16 April 2005, Boudrie 4062 ( UC) ; Crique Iracoumpay , 05º39'N, 53º37'W, 2 July 1990, Cremers & Hoff 11215 ( MO) . Province unknown: Without locality, without date, Appun 193 (K-fragment NY, paratype of Hemitelia moricandiana ) .— ECUADOR. Morona-Santiago: Trail Makuma-Mutinza, eastern foothills of Cordillera Cutucú , 02°10'S, 77°44'W, 660–750 m, 17 November 1999, Øllgaard & Navarrete 1486 ( AAU, QCA) .— PERU. Pasco: Oxapampa, Dist. Palcazú, Reserva Comunal Yanesha , Comunidad nativa 7 de Junio , Sector Pampa Hermosa , 10º11'41"S, 75º19'44"W, 410 m, 13 November 2005, Monteagudo et al. 11467 ( MO, USM) .— BRAZIL. Bahia: Municipio de Una, Reserva Biologica de Una , 14 November 2000, Fernandes, Sant'Ana & Carvalho 1499 ( NY) .

Remarks: — Cyathea macrocarpa has an unusual distribution as it occurs sporadically in the Andean foothills, the Guyana Highlands and northeastern Brazil. Windisch (1978) doubted the correctness of its locus classicus in the state of Bahia, Brazil, but the species was recently recollected there. The range supposedly reflects a preference for a special substrate, because it coincides with known occurrences of sandstone in South America ( Givnish et al. 2004). Enclosed in the geographical range of C. macrocarpa is the only known occurrence of C. rufescens (Mettenius ex Kuhn 1869: 159) Domin (1929: 264) in northern Peru. That species differs in having sessile pinnules (vs. largest pinnules at least short-stalked in C. macrocarpa ) and darker petiole scales (reddish brown vs. yellowish white to stramineous).

Cyathea macrosora (Baker) Domin (1929: 263) . Alsophila macrosora Baker in Im Thurn (1886: 211). Hemitelia macrosora (Baker) Jenman (1898: 45) . Sphaeropteris macrosora (Baker) Windisch (1973: 372) . Type:— GUYANA. Mount Roraima, 5750 ft, Im Thurn 87 (holotype K, isotype US). Fig. 13A .

Trunks erect, to 8 m tall, usually less than 2 m, slender, to 6 cm diameter, without old petiole bases. Fronds to 250 cm long, arching, distally drooping. Petioles 60–100 cm long, inermous to scabrous, dark brown to blackish, rarely distally dark stramineous, matte, without scurf, with a discontinuous line of pneumathodes along each side, to 5 × 1 mm, orange-brown when dried; scaly only around the base; petiole scales narrowly lanceolate to almost linear, 15.0–25.0 × 0.5–1.0 mm, concordantly to discordantly bicolorous, dark brown to auburn with white margins, basally cuneate, truncate or hastulate (with two flaring auricles), broadly attached, apically long acute, differentiated margins fragile, very narrow, mostly 1–2 cell rows wide, only near the base wider and recognizable, without setae or teeth, but with few to many, exerted, tortuous ciliae. Laminae to 130 × 80–100 cm, bipinnatepinnatifid, matte dark-green, blackish when dried adaxially, pale olive-green abaxially; apices gradually reduced. Rachises inermous, dark brown to atropurpureous abaxially and adaxially; with scattered brown multicellular hairs to 0.6(–1.0) mm long adaxially, glabrous abaxially. Largest pinnae 40–50 cm long, stalked 1.5–3.5 cm, ca. 18 pairs per frond, alternate, patent; distally notably green–alate, distal segments free to decurrently adnate. Costae 1.5–2.5 mm wide, inermous, dark brown to atropurpureous abaxially and adaxially, rarely dark straminoeus; adaxially moderately hairy with tan to brown multicellular hairs to 1.0 mm long, abaxially glabrous; junctures of costae and rachises swollen, bearing abaxially each one elliptic to lunular, planar pneumathode to 2–3(–5) × 1.0– 1.5 mm, colored like the adjacent tissue, inconspicuous. Largest pinnules 45–80(–90) × 9–20(–25) mm, stalked 3–5 mm, 1.0– 2.5 cm between the stalks/costules, linear-oblong to lanceolate, truncate to cordate basally, acute to attenuate apically, ending in shallowly crenate tips; costules in the same color as the costae, brown to dark stramineous, strongly prominent adaxially, weakly so abaxially, adaxially glabrous to hairy with multicellular hairs 0.5–1.0 mm long, glabrous abaxially except for lanceolate scales to 1–2 mm long, firm, shiny dark brown to atropurpureous, with dentate to shortly fimbriate margins; costules/stalks basally with inconspicuous pneumathodes, to 1.0 × 0.4 mm and a dark, often blackish incisions around their bases (abscission layer). Segments to 8–14 × 3–5 mm, deltate to oblong, incised 1/3 to 3/4 towards the costules, sessile, adnate, usually not remote, if so then connected by laminar tissue, patent to weakly ascending, distally weakly falcate, tips obtuse to acute; sinuses narrowly to widely deltate, acute, 1–3 mm wide; margins crenulate to finely serrate, usually in proximal segments basally entire; midveins weakly protruding, brown; lateral veins brown, planar on both sides; sterile and fertile veins simple or forked. Sori 1.0– 1.2 mm diameter, supramedial to inframarginal, more or less parallel to the margins, in the fork or on the back of a vein; indusia hemitelioid, small, variously developed on one frond, reaching 1/5–1/3 around the receptacle, covered by the sori, glabrous, tan, shiny, firm, margins entire; receptacles globose to elliptic, 0.3–0.4 mm long, paraphyses numerous, tan to brown, twisted, barely to much longer than the sporangia (0.4–0.6 mm long). Spores not examined.

Distribution and habitat: — Colombia, Venezuela, Guyana, French Guiana and Brazil, in moist lowland and premontane forest at 100–1700 m.

Additional specimens examined:— COLOMBIA. Amazonas: Corregimiento de Puerto Santander, resguardo indígena Nonuya de Villa Azul , 00°39'13.8"S, 72°04'27.2"W, 100 m, 29 June 2005, Castaño-A. et al. 404 ( COL) .— VENEZUELA. Amazonas: Río Negro, Camp XI, Cerro de La Neblina , 6.2 km NNE Pico Phelps (= Neblina ) (20.5 km ENE Neblina Base Camp), along trail into Cañon Grande , down ridge NW of camp, 00º51'45"N, 65º58'52"W, 1300–1390 m, 27 February 1985, Beitel 85305 ( UC) ; 0–3 km southeast of San Carlos de Rio Negro , 1°51'N, 67°03'W, 120 m, 21 January 1980, Liesner 8558 ( US) . Bolivar: Piar, at top of Salto Aicha neat E base of Uaipan-tepui , 05º38'N, 62º32'W, 1100 m, 27–28 November 1982, Davidse & Huber 22926 ( NY, UC) ; Amarauay-tepui, summit of W edge E of Auyan-tepui , W of Aparamán-tepui, 05º55'N, 62º15'W, 1030 m, 27 May 1986, Holst, Steyermark & Liesner 2988 ( UC) ; 1–2 km NE of El Pauji, Río Cabass , 800–900 m, 7 November 1985, Liesner 19651 ( UC) ; summit of Amaruay-tepui , S side E half, 05º55'N, 62º13'W, 50–1100 m, 11 May 1986, Liesner & Holst 20828, 20833 ( UC) ; Cerro Guaiquinima, Río Paragua, Intermediate Camp , 1200 m, 14 January 1952, B. Maguire 33103 ( UC) ; vicinity Mission Santa Teresita de Kavayanén , 1200–1300 m, 11 December 1952, Maguire & Wurdack 33765 ( UC) ; Río Tehuanen camp between Kavanayén and Ptari-tepui , 1240 m, 12–13 August 1970, Moore et al. 9707 ( UC) ; SW facing slopes of Chimantá tepui (Torono-tepui), above valley of Río Tirica , 1000–1700 m, 16 May 1953, Steyermark 75443 ( UC) .— FRENCH GUIANA. Mont Chauve , 03º49'N, 52º44'W, 100 m, 16 April 1997, Cremers & Crozier 15045 ( NY) .— PERU. Pasco: Oxapampa, along road Chatarra-Pto. Bermudéz , 10º30'S, 75º03'W, 700 m, 9 July 2003, van der Werff et al. 18171 ( MO) , Dist. Palcazú, Reserva Comunal Yanesha, Comunidad Nativa Lomalinda-Laguna , sector Nueva Aldea , 10º23'43"S, 75º05'12"W, 620–680 m, 14 October 2005, Monteagudo et al. 10588 ( MO, USM) .— BRAZIL. Amazonas: Manaus-Porto Velho Highway ( BR319 ) behind road camp, km 240, 21 November 1973, Lleras, Holley & Moneiro P 19583 ( NY) .

Remarks: — Windisch (1978) recognized two additional varieties in Cyathea macrosora, var. reginae and var. vaupensis , which are now treated as separate species: C. traillii ( Baker 1891: 188) Domin (1929: 264) and C. vaupensis ( Windisch 1973: 374) Lehnert (2011: 55) . Cyathea macrosora is characterized by the variable bicolored scales and the general absence of scurf. Related species like C. rufescens ( Kuhn 1869: 159) Domin (1929: 264) and C. macrocarpa differ in their larger, clearly visible indusia ( Lehnert 2011 b).

Cyathea meridensis Karsten (1869: 141) . Type:— VENEZUELA. Mérida: “Habitat in silvis montanis Meridensibus altitudine 2000 m ”, Karsten s.n. (holotype not located, W?). Fig. 2D . Cyathea firma Mettenius ex Kuhn (1869: 163) , not Cyathea firma Baker (1877: 161) Domin (1929: 263) . Cyathea calva var.

firma (Kuhn) Domin (1930: 102) . Type:— VENEZUELA. Mérida: Without locality, without date, Funck & Schlim 1228

( holotype P-fragment US, isotype BR) .

Trunks 0.5–10 m tall, 5–12 cm in diameter, with or without old petiole bases, aapices hidden in fascicles of petioles. Fronds to 340 cm long, arching to drooping. Petioles 50–200 cm long, inermous to verrucate or shortly aculeate, brown to castaneous; scurf dark brown to castaneous, a dense matted tomentum. Petiole scales 15–25 × 2.5–3.5 mm, lanceolate to ovate-lanceolate, concordantly to discordantly bicolorous, with dark brown to blackish centres and narrow orange to golden brown margins. Laminae to 200 × 150 cm, bipinnate-pinnatifid to tripinnate, coriaceous. Largest pinnae (16–) 47–75 cm long, alternate, 8–15 pairs per frond, stalked to (0.3–) 0.5–3.4 cm. Pinnules to 130 × 40 mm, triangular to oblong, tapering from the base or beyond the middle to acute or attenuate tips, subsessile to long-stalked; segments weakly ascending, their tips obtuse to round, margins flat to revolute, crenate to entire. Veins forked or simple, fertile veins forked. Sori 1.0– 1.4 mm diameter, subproximal, indusia subsphaeropteroid to sphaeropteroid; paraphyses few, thin, hyaline, of the same length as the sporangia (0.5 mm).

For full description see Lehnert (2009b).

Distribution and habitat: — Colombia, Venezuela, Ecuador and northern Peru, in moist montane forests at 1200–2500(–2870) m.

Selected specimen examined: — ECUADOR. Zamora-Chinchipe: Parque Nacional Podocarpus, plot C7 ( AG Gradstein FOR 402 ), Quebrada Los Lumos , along trail on ridge with white cross along the street, 03°59.379'S, 79°07.791'W, 2575 m, 7 January 2011, Lehnert 2338 (LOJA, QCA STU) .

Remarks:— Three varieties are recognized, of which the Cyathea meridensis var. meridensis and var. obtecta Lehnert (2009b: 421) may have long stalked, triangular pinnules ( Lehnert 2009b). The selected specimen represents the first record of var. meridensis in the south of Ecuador ( Fig. 2D ), which differs from the var. nana ( Lehnert 2009b: 420) and var. obtecta in its larger stature with massive trunks that shed the old petiole bases early (vs. stature smaller and old petiole bases persistent in vars. nana and obtecta)

Cyathea nodulifera Moran (1991: 88) . Type:— PANAMA. Veraguas: Cerro Tute, ca. 2 km N of Santa Fé, along ridge trail to summit, N of Esculea Primaria Alto de Piedra , 900–1000 m, 5 February 1988, Moran 4030 (holotype MO, isotypes PMA, UC, US) .

Trunks stout, to 0.5 m tall, otherwise unknown. Fronds to 300 cm long, presumably arching. Petioles 25–45 cm long, aculeate with few strong spines 1–5 mm long, atropurpureous to blackish, shiny, with dense scurf consisting of appressed, brown squamules to 0.6(–1.0) mm long; petioles basally without lenticels; only proximally scaly. Petiole scales narrowly lanceolate, 10–20 × 2–3 mm, shiny, concolorous brown or with lighter brown margins, bases weakly cordate, pseudopeltately attached, apices acute, straight, weakly undulate but not twisted; differentiated margins fragile, often abraded, the cell rows strongly exerted. Laminae to 110 cm wide, presumably broadly elliptic; bipinnate-pinnatifid, firm chartaceous to subcoriaceous, opaque, a rich green adaxially and abaxially; apices gradually reduced. Rachises inermous, atropurpureous to blackish abaxially and adaxially; distal parts adaxially pubescent with tan to brown multicellular hairs to 1.0 mm long, abaxially with dark brown scurf, consisting of appressed trichomidia and dissected squamules. Pinnae to 55 cm long, 12 pairs per frond, patent, stalked to 3–5 cm, alternate, inarticulate, distally narrowly green-alate, distal segments simply to decurrently adnate before ending in a pinnatifid apical section. Costae to 3.0 mm wide, inermous, atropurpureous to blackish on both sides; adaxially with tan, antrorsely curved, multicellular hairs to 1.0 mm long, abaxially with scurf like on the rachises; junctures of costae and rachises abaxially weakly swollen, each with one or two conspicuous, strongly protruding, circular aerophore to 2.0 mm diameter, pale brown. Largest pinnules 150–230 × 25–32 mm, stalked to 6 mm, inarticulate, 2–3 cm between the stalks, long lanceolate, bases cordate, tips attenuate with crenulate margins; costules carnose to ochre or yellowish green on both sides, adaxially strongly prominent, ridged densely hairy with tan to brown, antrorsely curved multicellular hairs to 1.0 mm long, abaxially weakly to strongly prominent, glabrous, with flat to subbullate, circular to ovate-acuminate, golden, squamules with finely erose-denticulate margins to 1.0(–2.0) × 0.5(–1.0) mm; costules basally with one or two strongly raised (1–2 mm high) pneumathodes, light brown, conspicious. Segments to 15 × 5–6 mm, sessile, adnate, patent to ascending, falcate, tips rounded to obtuse, proximal segments alternate to opposite, usually shorter than the following segments, never remote; sinuses acute to 0.5–1.0 mm wide, sometimes occluded; margins crenulate; margins not differently incised in proximal segments of a pinnule; veins protruding adaxially and abaxially, midveins adaxially ridged, dark stramineous to yellowish green, ending shortly before the margins, adaxially widened, deep orange-brown; veins glabrous on both sides except for some small squamules like on the costules; without bullate squamules; sterile veins simple or forked, fertile veins forked. Sori 0.8 mm diameter, inframedial to subproximal, parallel to the margins (weak triangular pattern), in the fork or above the fork of veins, mature dark orange-brown; indusia lacking; receptacles hemispherical to elliptic, 0.2–0.3 mm diameter, not subtended by a scale, paraphyses few, straight, reddish, shorter than the sporangia (0.2 mm long). Spores not examined.

Distribution and habitat: — Costa Rica and Panama at 900–1150 m, in wet forests.

Additional specimens examined:— COSTA RICA. Cartago: Turrialba, Jicotea , 09°46'48"N, 083°32'24"W (9.7800000, -83.5400000), 1000 m, 22 June 1995, Rivera & Rojas-A. 2537 ( MO) . Heredia: Between Río Peje and upper Río Guácimo , Atlantic slope of Volcán Barva , 10°16'48"N, 084°04'48"W (10.2800000, -84.0800000), 950– 1150 m, 11 November 1986, Grayum, Herrera-Ch. & Santana 7800 ( MO) .— PANAMA. Without locality, Pittier 5641 ( MO) .

Remarks:— Cyathea nodulifera is similar to C. kalbreyeri , from which it mainly differs in the prominent pneumathodes at the bases of costae and costules. Specimens from Ecuador identified as C. nodulifera actually belong to C. hemiepiphytica .

Cyathea traillii (Baker) Domin (1929: 264) . Hemitelia traillii Baker (1891: 188) . Type:— BRAZIL. Amazonas:

Rio Mahues (Maués), 30 April 1874, Traill 1384 (holotype K-fragment US, isotypes GH, P-fragment F). Fig. 12C .

Hemitelia pumila Maxon (1946: 439) . Type:— COLOMBIA. Caqueta: Cerro del Castillo, upper Apaporis basin, near confluence of the Ajuju and Macaya , Schultes 5664 (holotype US) .

Sphaeropteris macrosora (Baker) Windisch var. reginae Windisch (1973: 374) . Cyathea macrosora (Baker) Domin var. reginae (Windisch) Smith (1990: 25) . Cyathea reginae (Windisch) Smith (2006: 426) . Type:— COLOMBIA. Vaupés: Raudal Jerijerimo, Río Apaporis, Schultes & Cabrera 13448 (holotype GH).

Trunks erect to decumbent, to 4 m tall, usually less than 2 m, slender, 4–5 cm diameter, without old petiole bases; epidermis dark brown, covered in narrowly lanceolate scales 15–30 × 2 mm, bicolorous brown with yellowishwhite to white margins, strongly twisted, frond scars to 50 × 25 mm, light brown, below with large oval lenticels, orange brown to vermillion, to 15 × 3 mm; apex not hidden in fascicle of petioles, elongated, unexpanded crosiers visible; without adventitious buds. Fronds to 200 cm long, arching, distally drooping; apparently spirally arranged on the trunk. Petioles 55–70 cm long, inermous to scabrous, dark brown to blackish, matte, with scurf consisting of contorted multicellular hairs and small squamules with long, hair-like ciliae, forming a matted tomentum, usually pale brown to tan (if petiole scales bicolorous), sometimes white (if petiole scales concolorus whitish to stramineous), easily abraded, often completely lacking; with a discontinuous line of pneumathodes along each side, to 3 × 1 mm, brown to orange-brown when dried, inconspicuous; scaly near the bases or up to the middle; petiole scales narrowly lanceolate, to 12.0–15.0(–20.0) × 1.5–3.5 mm, concordantly to discordantly bicolorous, dark brown to auburn with white margins, or concolorous white to stramineous with occasional brown streaks; basally cuneate, truncate or hastulate (with two flaring auricles), broadly attached, apically long acute, differentiated margins fragile, very narrow, mostly 1–2 cell rows wide, only near the base wider and recognizable, without setae or teeth, but with few to many, exerted, tortuous ciliae. Laminae to 130 × 80–100 cm, bipinnate-pinnatifid; matte dark-green, blackish when dried adaxially, dark olive-green abaxially; apices gradually reduced. Rachises inermous, dark brown to atropurpureous abaxially and adaxially; with scattered brown multicellular hairs to 0.6(– 1.0) mm long adaxially, with white scurf of matted, tortuous hairs adaxially and abaxially. Pinnae to 35–50 cm long, stalked 1.5–3.0(–4.0) cm, alternate, patent to weakly ascending; distally notably green-alate, distal segments free before ending in pinnatifid apical sections; basal pinnae with the basal pinnules often missing in the basiscopic half. Costae 2.0– 2.5 mm wide, inermous, dark brown to atropurpureous abaxially and adaxially; adaxially moderately hairy with tan to brown multicellular hairs to 1.0 mm long, abaxially glabrous, with remnants of white scurf on both sides; junctures of costae and rachises swollen, bearing abaxially each one or two elliptic to lunular, planar pneumathodes 2–3 × 1.0– 1.5 mm, colored like the adjacent tissue, inconspicuous. Largest pinnules 90–125 × (18–)20–25(–35) mm, articulate, stalked 2–4(–6) mm, 1.5–2.5 cm between the stalks/costules, lanceolate to long triangular, cordate to truncate basally, attenuate apically, ending in shallowly crenate tips; costules in the same color as the costae, blackish to dark brown, strongly prominent adaxially, weakly so abaxially, adaxially hairy with brown multicellular hairs 0.5–1.0 mm long, glabrous abaxially except for remants of white scurf and some lanceolate scales firm, dark brown to auburn, with paler, dentate to shortly fimbriate margins, flat ones to 3 mm long, bullate ones to 1 mm long; costules/stalks basally with inconspicuous pneumathodes, to 1.0 × 0.4 mm and a dark, often blackish incision around their bases (abscission layer). Segments to 9–12 × 3–5 mm, oblong, incised 3/ 4 towards the costules, sessile, adnate, not remote, weakly ascending, falcate, basal segmenst often reflexed; tips obtuse to round; sinuses narrowly to widely deltate, acute, 1–3 mm wide; margins crenulate to serrate, usually in proximal segments basally more strongly crenate; midveins weakly protruding, brown; lateral veins brown, planar on both sides; sterile and fertile veins simple or forked. Sori 1.0– 1.2 mm diameter, medial, more or less parallel to the margins, in the fork or on the back of a vein; indusia hemitelioid, reaching to 1/4 –1/5 around the receptacle, arching but not covering the sori, glabrous, brown, shiny, firm, margins finely dentate; receptacles globose to elliptic, 0.3–0.4 mm long, paraphyses numerous, tan to brown, twisted, barely to much longer than the sporangia (0.4–0.6 mm long). Spores not examined.

Distribution and habitat: — Costa Rica, Colombia, Venezuela, Guyana, Peru and Brazil, to be expected in eastern Ecuador, in wet lowland and premontane forests at 100–1000(–1670) m.

Selected specimens examined:— COSTA RICA. Puntarenas: Golfito, Playa Cacao, Punta Voladera , 08°37'35"N, 83°11'00"W, 100 m, 6 July 1998, Herrera & Rivera 7124 ( F) .— COLOMBIA. Vaupés: Río Apoporis, Jinogojé (at mouth of Río Pirapaná ) and vicinity, 00º15'N, 70º30'W, 700 ft, Schultes & Cabrera 16624 (US) ; headwaters of Caño Teemeña, Lobo Igarapé, Río Piraparaná (tributary of Río Apaporis ), 00º15'S –25' N , 70º30'W, 10 September 1952, Schultes & Cabrera 17343 ( NY, UC) .— VENEZUELA. Amazonas: Misión Río Mavaca , 02º26'N, 65º07'W, 185 m, 29 January 1991, Stergios & Yánez 15030a ( MO, UC) .— GUYANA. U . Takutu-U. Essequibo: Acarai Mountains, 01º20'N, 58º48'W, 300–600 m, 2 November 1996, Clarke 2861 ( NY); Wassarai Mountains , summit of highest peak, 14 km S of S. Kassikaityu R ., 01º31'40"N, 59º14'23"W, 1135 m, 8 September 1999, Clarke, Williams & Perry 8263 ( NY) . Cuyuni-Mazaruni: Pakaraima Mountains , 0.5 km NW of Imbaimadai settlement, 05º42'N, 60º17'W, 525–575 m, 17 November 1992, Hoffman 3406 ( NY) .— PERU. Amazonas: Condorcanqui, Distrito El Cenepa, Comunidad de Tutino , Cerro Tutino , 04º33´50"W, 78º12´15"W, 600 m, "Shinkushuk", 24 June 1997, Vasquez et al. 24141 ( MO, UC) . Loreto: Maynas Upper Río Itaya, about 14 km WSW of the village Carbajal 04º18'S 73º43'W, 100–200 m, 12 March 1996, Tuomisto et al. 10184, 10185, 10186 ( GOET, TUR) . Pasco: Oxapampa Dist. Palcazú, P. N . Yanachaga-Chemillén, surroundings of biological station Paujil , trench to view point, 10º19'S 75º15'W, 355 m, 8 May 2003, Monteagudo, Francis & Ortiz 5096 ( MO, UC) .— BRAZIL. Amazonas: Manaus-Porto Velho Highway, Km 250, 18 March 1974, Prance et al. 20702 ( MO, NY) .

Remarks: — Cyathea traillii is the most widely distributed species of the C. macrosora -alliance and is characterized by well-developed hemitelioid indusia and relatively broad bicolorous petiole scales. In comparison, C. vaupensis has practically no indusium; it is reduced to a small flap that is only visible if the whole receptacle is removed, but conglomerated paraphyses may form narrow structures that may be mistaken for small indusia in that species. The petiole scales of C. traillii are commonly 3 mm or wider, in C. macrosora and C. rufescens they are less than 2 mm wide; in C. macrocarpa they differ additionally in being concolorous whitish to stramineous (vs. bicolorous to almost concolorous brown). The Peruvian poplation of C. traillii usually has nearly concolorus petiole scales but clearly bicolorous scales on the trunk, which agrees with the Venezuelan population and separates them from C. macrocarpa .

Cyathea traillii is most similar to C. rufescens in the fine indument of petioles and laminae, especially in the white tortuous hairs found on the frond axes; that species, however has sessile, clearly pinnate pinnules (vs. always short stalked and pinnatisect to pinnatifid in C. traillii ).

Cyathea tuerckheimii Maxon (1909a: 4) . Cyathea divergens var. tuerckheimii (Maxon) Tryon (1976: 56) . Type:— GUATEMALA. Alta Verapaz: Coban, 1500 m, November 1907, Türckheim II 1645 (holotype US, isotypes BR, F, LE, MEXU, MO, NY, P, S, UC, US). Fig. 14 .

Trunks to 15(–18) m tall, 8–12 cm diameter, without old petiole bases; upper parts clothed in concolorous brown scales, similar to petiole scales; epidermis dark brown to blackish; frond scars round to elliptic, 36 × 24 cm, light brown to grey, with small round orange-brown pneumathodes below them; trunk apices hidden in fascicles of the youngest petioles; adventitious buds lacking. Fronds to 300 cm long, arching, distally not drooping. Petioles at least to 50 cm long, verrucate to muricate, not aculeate, stramineous to tan, proximally brown; without adventitious (aphlebioid) pinnae at the petiole bases; petiole scales narrow-lanceolate to lanceolate, 12–20 × 2.5–3.0 mm, concolorous shiny brown to auburn, or bicolorous with a thin white to yellowish margin, sharply set or getting gradually paler from the brown to dark brown centres towards the margins; petiole scurf well developed but not very persistent, white to stramineous, consisting of small erect squamules, oblong, the tips rounded, broadly truncate or lobed, 0.2–1.0 mm long, mainly whitish with brown base or body, without dark marginal cells; undercoat of smaller appressed squamules lacking. Laminae to 250 × 100–140 cm, bipinnate-pinnatifid, firm herbaceous to chartaceous, apices gradually reduced, not drooping; dark green adaxially, usually blackish when dried, pale green abaxially. Largest pinnae 50–70 cm long, ca. 15 pairs per frond, alternate, short stalked to 1.5(– 4.5) cm, distally narrowly green alate, the distal pinnules/segments decurrently adnate. Frond axes pale brown to brown, hairs only adaxially on costules, costae and distal parts of rachises, hairs 0.5–1.0 mm long, whitish to tan, abaxially with tan to light to brown scurf consisting of scattered small squamules 0.2–1.0 mm long, larger ones like those of the petiole, ovate-lanceolate, subentire to dentate or short fimbriate margins, smaller ones lobed to dissected with spinulose margins; scurf persisting in junctures of costae with costules and rachis; costules sometimes also with small ovate-lanceolate scales 1–2 mm long, dark brown with paler margins; costae muricate to smooth, rarely more than 2–3 mm wide. Largest pinnules 9.0–13.5 × (1.2–)2.0– 2.5 cm, subsessile to short stalked (5 mm), alternate, 1.2–1.4 cm between the stalks, oblong to long triangular, truncate to weakly cordate at base, tapering from beyond the middle to short attenuate tips, the segments weakly falcate with finely crenate margins and rounded to obtuse, rarely acute tips; basal segments alternate to opposite, never remote from each other, sinuses narrow (1–2 mm), obtuse to acute; sterile pinnules usually not broader than fertile ones. Veins adaxially glabrous, abaxially usually with few white multicellular hairs or with some appressed unicellular trichomidia instead & some small flattish, pale brown to brown ovate squamules with elongated tips to 2 mm long; finely dissected squamules absent; regularly with few subbullate to bullate brown squamules 1–2 mm long on the midribs of segments distally; sterile veins forked or simple, fertile veins forked. Sori 0.6–1.0 mm diameter, proximal to submedial, indusia sphaeropteroid, with umbo, tan, translucent, fragile but persisting as a cup; paraphyses as long as or shorter than sporangia. Spores not examined.

Distribution and habitat: — Mexico, Guatemala and Honduras, also El Salvador ( Mickel & Smith 2004, Monterrosa & Monro 2008) and Nicaragua ( Mickel & Smith 2004), in moist montane forests at 900–2000 m (to 2700 m fide Véliz & Vargas 2006). Not yet reported from Belize.

Additional specimens examined:— MEXICO. Chiapas: From El Triunfo (gap of mountains) along road to Finca Liquidambar , Sierra de Soconusco , 2100 m, 11 November 1945, Xolocotzi & Sharp 472 (US). Veracruz- Llave : About 8 km S of Misantla , ca. 750 m, 1 January 1974, Conant 840 ( LSC) .— GUATEMALA. Huehuetenango: Top of Cerro Chemalito, Sierra de los Cuchumatanes , 3.5 mi W of Santa Eulalaia, 3100–3150 m, 2 August 1942, Steyermark 49962 ( PR) . Alta Verapaz: Cobán , 1500 m, November 1907, Von Tuerckheim s.n. [Herb. Rosenstock #6] ( PR) . Verapaz : Guatemala Highway #17, ca. 35.1 mi S of Cobán and W of Purulhá , 8 January 1973, Alameda & Luteyn 1776 ( F) . El Quiche: Chunama District, between Pulay and San Juan Cotzal, 6200 ft, 1 July 1964, Proctor 25016 ( US). San Marcos : Between Canjulá and La Unión Juárez , near SE portion of Volcán Tacaná, 2000–3000 m, 22 February 1949, Steyermark 36394 ( F) ; near Aldea Fraternidad, between San Rafael Pie de Cuesta and Palo Gordo , W facing slope of the Sierra Madre Mountains , 1800–2400 m, 10–18 December 1963, Williams et al. 25670 ( UC) .— HONDURAS. Cortes: Parque Nacional El Cusuco, Sierra de Omoa , W of San Pedro Sula , 1534 m, Kelly 314 ( BM) .

Remarks: —With the separation of Cyathea ×jurgensenii, C. tuerckheimii appears morphologically more constant and clear-cut within its range than before. Tryon (1976) remarked that C. tuerckheimii is not distinct enough to merit distinction as a species from C. divergens . However, the characters of C. tuerckheimii contrast with those of the adjacent population of C. divergens var. divergens in Costa Rica, which indicates that they are better separated as two distinct species. Habitually, C. tuerckheimii grows up to 18 m tall ( Véliz & Vargas 2006) and has patent-arching fronds like many large species of Cyathea ( Tryon 1976, Véliz & Vargas 2006); Cyathea divergens var. divergens has shorter trunks to 6(–10) m tall and long-drooping fronds. The petiole scurf of C. tuerckheimii appears more flakey and scattered than in C. divergens , in which the petiole scurf initally completely covers the epidermis of the petiole. The similarity in the scurf is only superficial; a closer look reveals structural differences, above all the lack of an undercoat of finely dissected squamules on the petiole in C. tuerckheimiii , which is always present in C. divergens . In both species, the pale colour of the petiole indument is lost in wet conditions, since the dead cells soak up the water and become transclucent. Also, the petiole scales of C. tuerckheimii appear more reddish brown with relatively narrow white margins while those of C. divergens var. divergens are sometimes almost black and have broad, white to stramineous margins that often extend into the centre.

In some Mexican material of Cyathea tuerckheimii (including the type collection), few stellate hairs have been found among the petiole scurf squamules ( Fig. 13 ). These may represent contamination, because they are more typical of the genus Alsophila Brown (1810: 158) .

Cyathea ulei (H.Christ) Domin (1930: 168) . Alsophila ulei Christ (1905: 367) . Trichipteris ulei (H.Christ) Tryon (1970: 46) . Type:— PERU. Amazonas: Cerro de Ponasa, 1300 m, 1903, Ule 6901 (holotype P, isotype B). Cyathea subtropica Domin (1929: 263) , nom. nov. for Alsophila lechleri Mettenius (1859: 32) , not Cyathea lechleri Mettenius

(1859: 32). Trichipteris lechleri (Mett.) Tryon (1970: 45) . Type:— PERU. Puno: Tatanara, Lechler 2532 (holotype B

[Herb. Mettenius]).

Trunks erect to decumbent, (0.3–)0.5–4.0 m tall, (3–)6–8(–10) cm diameter, without old petiole bases; upper parts covered in concolorous brown to auburn scales, similar to petiole scales; epdermis dark brown to blackish, mostly obscured by scales; sheath of adventitious roots at the bases usually thin/not massive; apex hidden in fascicle of petioles, unfolding only one crosier at a time. Petioles to 90 cm long, inermous to verrucate, dark brown to castaneous, sometimes atropurpureous or basally black; with a discontinuous line of narrow white lenticels on each side when young, hardly seen in old and dried material; without adventitious (aphlebioid) pinnae at the petiole bases; petiole scales broadly lanceolate to long ovate, 10–13 × (4.5–)5.0–6.0 mm, their tips straight, shiny auburn to dark reddish brown, concordantly bicolorous with broad, pale to golden brown margins; colors transient, not sharply contrasted; petiole scurf of small reddish brown, branched hairs 0.2–0.4 mm long, only on crosiers and unfolding fronds, usually absent in most parts when fronds unfolded. Fronds to 240 cm long, arching to weakly drooping. Laminae to 150 × 80–110 cm, bipinnate to bipinnate-pinnatifid, firm herbaceous to chartaceous, apex gradually to abruptly reduced, non-conform; glossy dark green adaxially, often blackish when dried, olive green abaxially. Pinnae to 35–55 cm long, 8–12 pairs per frond, long stalked to (1.5–)2.0–5.0(–8.0) cm, distally narrowly green alate, the distal segments simply adnate. Frond axes brown to castaneous on both sides; completely glabrous abaxially, hairy only adaxially on costules, costae and distal parts of rachises; hairs 0.5–1.0 mm long, tan to brown; costae smooth, 2–3 mm wide; insertions of costae into rachises swollen, with one inconspicuous pneumathode in the color of the costae, elliptic to 2 mm long. Largest pinnules 6.0–10.5 × 1.1–2.0(–2.4) cm, linearly oblong, incised to 1/2 or less of their width, truncate to cuneate at their bases, fertile pinnules also weakly cordate, tapering from beyond the middle to long acuminate to short attenuate tips, the largest ones stalked to (1.0–)2.0–3.0 mm, alternate, (0.4–)1.5–2.0 cm between the stalks; the brown stalks articulate, with a dark ring around their insertions and each with a small elliptic pneumathode 1.0– 1.5 mm long at their base; the segments patent, rounded to truncate at tips, with crenulate to weakly dentate margins; tips sometimes falcate, with the acroscopic lobe usually supported by a lateral vein; basal segments usually opposite, the lowest ones not remote from each other, sinuses acute, narrow, 0.5–1.0 mm wide; segments with many pale brown to reddish unicellular trichomoidia evenly distributed on and between veins abaxially, often ephemeral. Veins pale brown to yellowish, prominent abaxially and adaxially, contiguous with the identically coloured hyaline segment margins, glabrous adaxially; abaxially glabrous except for occasional auburn to brown flat squamules to 1.0 mm long on the veins; sterile and fertile veins simple. Sori 1.0– 1.2 mm diameter, inframedial to subproximal, indusia absent, receptacles globose, 0.3–0.4 mm diameter; paraphyses of the same length as or longer than sporangia (0.5–0.6 mm). Spores with psilate perispore, weakly pitted near laesura ( Gastony 1979).

Distribution and habitat: — Colombia, Venezuela, Ecuador, Peru and Bolivia in moist tropical montane forests at 950–1800 m on the slopes of the eastern Andean and the table mountains of the Guayana shield.

Additional specimens examined:— COLOMBIA. Cauca: Santa Rosa, Serrania de los Churumbelos , Bota Caucana , 01°14'N, 76°31'W, 15 August 2002, Gonzales 903 ( COL) . Huila: Cordillera Oriental, SW of Alejandria at Rio Suaza , 1707 m, 17 September 1944, Little 8498 ( US) . Nariño: La Planada, Salazar Finca 7 km above Ricaurte, ca. 01°08'N, ca. 77°58'W, 1750 m, 27 November 1985, Gentry et al. 34995 ( COL, UC) ; Ricaurte, centro de cientificos a la Piña , 01°09'463"N, 77°58'590"W, 1800 m, 20 June 1999, Sánchez Baracaldo 23 ( UC) . Norte de Santander: Catatumbo, Cerro del Tirador , cerca de Las Mercedes , 1000 m, 19 May 1963, Bishler 2541 ( COL) .— VENEZUELA. Amazonas: Río Negro, Cerro Neblina , camp #7, 00°50'N, 65°58'W, 1850 m, 2 December 1988, Anderson 13451 ( UC) , S lopes of Cañon Grande , 00°55'N, 66°00'W, 1800 m, 30 November 1988, Croat 59424, 59455 ( UC) , valley near camp #5 at N base of Pico Cardenas, 00°49'N, 66°00'W, 1200 m, 14 April 1988, Gentry & Stein 46619 ( UC) , ridge at divide between Brazil and Venezuela , 26 km ENE of Neblina base camp, 00°53'N, 65°56'W, 2000 m, 16 April 1988, Plowman & Thomas 13602 ( UC) . Bolivar: Cedeno, Serranía de Maigualida , 20 km E of San José de Kayamá, 45 km N of Cerro Impacto, 06°19'N, 65°12'W, 1250 m, April 1993, Fernandez A. 5349 ( UC) . — ECUADOR. Morona-Santiago: Road Susana de Chivasa-Panecillo, Km 3–4, 02°55'S, 78°04'W, 1300 m, 19 November 2001, Øllgaard & Navarrete 2625 ( AAU, QCA) . Pastaza: Mera, rd and muletracks to approx. 4 km N of the village (along Río San Jorge and Río Tigra ), 01º35'S, 77º53'W, 1200 m, 1–2 September 1976, Øllgaard & Balslev 9102 ( QCA) ; road N of Mangayacu, Km 1.8 ( W of Mera), 01º26'S, 78º07'W, 1400 m, 13 November 1994, Øllgaard & Navarrete 105647 ( QCA) . Zamora-Chinchipe: Along road from Zamora to Romerillos along Río Jambué , 13.3 km E of Río Bombuscaro bridge in Zamora , 0.3 km E of Pituca , 04°08'03"S, 78º56'37"W, 1068 m, 21 July 2004, Croat 9176 B ( MO) ; Campamento Shaime (Shaimi) along Río Nangaritza , trail to the oil bird cave ("cueva de los tayos"), 04°19'S, 78°40'W, 950–1050 m, 7 November 2004, Lehnert 1520 ( GOET, QCA, UC) ; Miazi, junction Río Chumbiriatza to Río Nangaritza , 04°19'S, 78°40'W, 950–1200 m, 22 October 1995, Øllgaard, Bergmann & Ruíz 99307 ( AAU, QCNE) .— PERU. Cusco: Paucartambo, Dto. Cosñipata, Parque Nacional del Manu , entre Quebrada Quita Calzón e hito Km 164, 1180– 1100 m, 15 September 1991, León , Young & Huapaya 2940 ( LL n.v., MO, TEX n.v.). Pasco: Oxapampa , Dist. Palcazú , sector San Pedro de Pichanaz (cordillera San Matías ), 10º25'58"S, 75º00'21"W, 1030 m, 2 March 2004, Mellado-N. & Monteagudo 1169 ( MO) .— BOLIVIA, La Paz: Nor Yungas, Estación Biológica Tunquini , 16°11'S, 67°53'W, 1730 m, 20 September 2004, Eberhardt, Syszka & Pelka 546 ( GOET, LPB) ; Bautista Saavedra, Cerro Asunta Pata, entre Apolo y Charazani , 15°03'S, 68° 29'W, 1500 m, 22 June 2001, Kessler et al. 10198 ( LPB, UC) ; Larecaja, Hacienda Sumaco sobre el camino a Tipuani , 1400 m, January–February 1920, Buchtien 5299, 5300 ( GH, US) , 5301 ( UC, US) .

Remarks: — Cyathea ulei is sometimes treated as C. subtropica ( Holttum & Edwards 1983) , a name that poignantly refers to the preferred occurrence of the species between 1000–2000 m, in the subtropical elevational belt ( Holdridge 1947). Cyathea ulei is relatively variable in the size, the stalks and the incision of the pinnules, but can always be recognized by the basal veins that arise from the costules and are connivent to the sinuses, often ending blindly below them. This separates it especially from the morphologically closest species, C. hemiepiphytica , which usually has sessile to short-stalked pinnules. Furthermore, the pinnules of C. ulei are often incised only to ca. 1/5 towards the costules, which results in characteristic truncate lobes on the pinnules.

Cyathea vaupensis (Windisch) Lehnert (2011: 55) . Sphaeropteris macrosora (Baker) Windisch var. vaupensis Windisch (1973: 374) . Type:— COLOMBIA. Vaupés: Cerro Isibukuri, Río Kanari, Comisaria del Vaupes, Schultes & Cabrera 13407 (holotype GH).

Trunks erect, to 1 m tall, slender, to 6 cm diameter, without old petiole bases. Fronds at least to 150 cm long, arching, distally drooping. Petioles 55–100 cm long, inermous to scabrous, dark-brown to blackish, matte, with ephemeral scurf of matted, whitish lanceolate squamules to 0.5 mm long, with ciliate margins; with a discontinuous line of pneumathodes along each side, to 5 × 1 mm, orange-brown when dried; scaly to the full length, usually reaching up to central parts of the rachises; petiole scales narrowly lanceolate to almost linear, 10.0–15.0 × 0.5(– 1.0) mm, in upper parts of the petiole almost hair-like, concordantly to discordantly bicolorous, dark brown to auburn with white margins, broadest scales sometimes almost concolorous, basally cuneate, truncate or hastulate (with two flaring auricles), broadly attached, apically long acute, differentiated margins fragile, very narrow, mostly 1–2 cell rows wide, only near the base wider and recognizable, without setae or teeth, but with few to many, exerted, tortuous ciliae. Laminae to 100 × 70 cm, bipinnate-pinnatifid, firm chartaceous to subcoriaceous; matte dark-green, blackish when dried adaxially, pale olive-green abaxially; apices gradually reduced. Rachises inermous, dark brown to atropurpureous abaxially and adaxially; with scattered brown multicellular hairs to 0.6(– 1.0) mm long adaxially, with scurf and scales like on the petioles. Pinnae to 40–50 cm long, stalked 1.0–1.5(–2.0) cm, alternate, patent; distally notably green-alate, distal segments free to decurrently adnate. Costae 1.5–2.5 mm wide, inermous, dark brown to atropurpureous abaxially and adaxially; adaxially moderately hairy with tan to brown multicellular hairs to 1.0 mm long, abaxially glabrous; junctures of costae and rachises swollen, each bearing abaxially one elliptic to lunular, planar pneumathode 2–3(–5) × 1.0– 1.5 mm, colored like the adjacent tissue, inconspicuous. Largest pinnules to 125 × 12–20 mm, stalked 3–5 mm, 1.5–3.0 cm between the stalks/ costules, linear-oblong to lanceolate, truncate to cordate basally, long acute to attenuate apically, ending in shallowly crenate tips; costules in the same color as the costae, brown to dark atropurpureous, strongly prominent adaxially, weakly so abaxially, adaxially hairy with multicellular hairs 0.5–1.0 mm long, glabrous abaxially except for lanceolate to linear scales to 3.0 × 0.5 mm long, papery, matte whitish to tan, with dentate to shortly fimbriate margins; costules/stalks basally with inconspicuous pneumathodes, to 1.0 × 0.4 mm and dark, often blackish incisions around their bases (abscission layer). Segments to 15 × 3–5 mm, oblong, incised to 3/4 towards the costules, sessile, adnate, usually not remote, if so then connected by laminar tissue, patent to weakly ascending, distally weakly falcate, tips obtuse to acute; sinuses narrowly to widely deltate, acute, 1–3 mm wide; margins crenulate to finely serrate; midveins weakly protruding, brown; lateral veins brown, planar on both sides; sterile and fertile veins simple or forked. Sori 1.0– 1.2 mm diameter, supramedial to inframarginal, more or less parallel to the margins, in the fork or on the back of a vein; indusia hemitelioid, small, variously developed on one frond, reaching to 1/5–1/10 around the receptacle, covered by the sori, glabrous, tan, shiny, firm, margins entire; receptacles globose to elliptic, 0.3–0.4 mm long, paraphyses numerous, tan to brown, twisted, barely to much longer than the sporangia (0.4–0.6 mm long). Spores not examined.

Distribution and habitat: — Colombia, Venezuela and Peru in moist terra firme forests at 75– 500 m.

Additional specimens examined:— VENEZUELA. Amazonas: Río Negro, Km 11 NE of San Carlos de Río Negro along the road to Solano , 01º35'N, 67º02'W, 75 m, 24 June 1984, Davidse & Miller 26526 ( MO, UC) . Bolivar: Piar, Río Aparamán, affluent of Río Acanán near Yuray-merü rapids, 1.5 km S of SW corner of Amaruaztepui , 05º54'N, 62º15'W, 500 m, 22 April 1986, Liesner & Holst 20182 ( MO, UC) .— PERU. Loreto: Río Tigre , about 1.5 km E of the river, 03º57'S, 74º17'W, 100–200 m, 10 January 2005, Tuomisto et al. 13983 ( TUR), 1.5 km NE of the village San Andrés N of the river, just W of Cocha Brava , 03º44'S, 74º28'W, 100–200 m, 9 February 2005, Tuomisto et al. 14970 ( TUR) .

Remarks: —Easily recognized and distinguished from Cyathea divergens and most other species treated here by its ample cover of persistent, clearly bicolorous, narrow petiole scales, which reach up at least to the lower half of the rachises.

Figures

FIGURE 2. Field characters: A1–E1. Trunk apex; A2–E2. Rachises and costules abaxially, showing different colors and indument; A3–E3. Details of petioles with different types of scales and scurf. A. Cyathea divergens, Costa Rica (Lehnert 2642); white areas on petioles indicate dry scurf, which elsewhere became wet and translucent. B. Cyathea carolihenrici, Bolivia (Lehnert 607). C. Cyathea maxonii, Costa Rica (Lehnert 2658). D. Cyathea meridensis var. meridensis (D1, 2) and var. obtecta (D3), Ecuador (Lehnert 2338 & 1079, respectively). E. Cyathea ebenina, Ecuador (Lehnert 1055). (© Marcus Lehnert)

FIGURE 4. Cyathea divergens var. divergens A. Detail of petiole with fine scurf, partially abraded (right) (Churchill 3307, STU); B. Detail of scurf squamules, larger one whitish, smaller one light brown (Cogollo et al. 6162, UC); C. Petiole scales (Cogollo et al. 6162, UC); D. Pinna (Churchill 3307, STU); E. Large pinnule abaxially, fertile (Poeppig s.n., K); F. Small pinnule adaxially, fertile (Tonduz 11802, BR).

FIGURE 5. Cyathea ebenina. A. Pinna (Karsten s.n., B).

FIGURE 6. Cyathea farinosa. A. Pinna; B. Larger pinnule adaxially, fertile; C. Segment abaxially, fertile (all from Karsten s.n., B).

FIGURE 7. Cyathea gibbosa. A. Petiole scales (Lehnert 2106, STU); B. Pinna (Lehnert 2106, STU); C. Pinnule abaxially, fertile (Lehnert 2662, STU).

FIGURE 8. Cyathea gibbosa. A. Trunk, ca. 170 cm tall; apex of frond of neighbor plant seen at the right; with M. Kessler (Z). B. Fronds hanging over branch, total length ca. 7 m; arrows indicate an unfurling, still unsupported frond with its pinnae still coiled up tightly. Prov. Zamora-Chinchipe, Ecuador. (© Marcus Lehnert)

FIGURE 9. Cyathea gracilis. A. Pinna (Lehnert 183, STU); B. Smaller pinnule abaxially, fertile (Purdie s.n., K).

FIGURE 10. Cyathea kalbreyeri. A. Pinna. (Kalbreyer 1375, B).

FIGURE 11. Cyathea latevagans. A. Part of caudex with petioles and a few scales (right) (Garcia-C. et al. 104, MO); B. Distal part of frond, reconstructed from several sheets (Kalbreyer 1327, K); C. Pinna abaxially, fertile (Kalbreyer 1327, K); D. Laminar scale (Garcia-C. et al. 104, MO); D. Paraphyses on receptacle (Garcia-C. et al. 104, MO).

FIGURE 12. Cyathea latevagans growing in seepage area, Prov. Cuzco, Peru. (© Adrian Tejedor)

FIGURE 13. A. Cyathea macrosora. Large pinnule abaxially (Im Thurn 87, K). B. Cyathea macrocarpa. Large pinnule abaxially, fertile (Blanchet 3227, K). C. Cyathea traillii. Large pinnule abaxially, fertile (Traill 1384, K).

FIGURE 14. Cyathea tuerckheimii A. Petiole scurf, large and small squamule, plus stellate hair (probably contamination) (Türckheim II 1645, UC); B. Segment abaxially, showing complete indusia and brown squamules (Türckheim II 1645, US).