Cyathea werffii R.C.Moran

Cyathea werffii R.C.Moran View in CoL

Fig. 13 View Fig

Cyathea werffii R.C.Moran ( Moran 1991: 94, fig.5).

– Type: ECUADOR • Morona-Santiago, along new road Mendez-Morona , km 55–62; 800 m a.s.l.; 23 Aug. 1989; H. van der Werff & E. Gudiño 11386; holotype: MO [ MO-288184 ]!; isotypes: AAU!, QCNE [ QCNE-179 ]!, UC [ UC1564628 ] !.

Etymology

The specific epithet honors Henk van der Werff, botanist at the Missouri Botanical Garden and collector of the type specimen.

Selected material studied

COLOMBIA – Caquetá • Municipio de Belén de los Andaquies, Parque Natural Municipial Andaqui , cerro de Aguacata y flanco oriental hacia el sector La Mina ; 1°39′17.5″ N, 75°54′29.1″ W; 1200–1450 m a.s.l.; N. Castaño-A. et al. 9197; COAH. GoogleMaps – Putomayo • Municipio de Orito, Santuario de Flora de Plantas Medicinales Ingi-Andé ( Nuestra Tierra ); 0°41′06.2″ N, 77°03′07.89″ W; 944 m a.s.l.; 26 Sep. 2015; W.D. Rodríguez, D. Cárdenas, N. Marin & J. Restrepo 9354; COAH GoogleMaps .

ECUADOR – Morona-Santiago • Along road between Santiago and Río Morona ; 2º58′24″ S, 77º49′36″ W; 322 m a.s.l.; 10 Jul. 2004; T.B. Croat et al. 90750; MO GoogleMaps • Comunidad Shuar de Mutints, faldas orientales de la Cordillera de Cutucú ; 2°11′ S, 77°44′ W; 600 m a.s.l.; 10 Sep. 1995; H. Navarrete 1214; AAU, QCA GoogleMaps • Comunidad Shuar de Mutints, faldas orientales de la Cordillera de Cutucú ; 2°11′ S, 77°44′ W; 600 m a.s.l.; 10 Sep. 1995; H. Navarrete 1248; AAU, QCA GoogleMaps • Road 3.8 km N of Santa Susana de Chiviasa (NE of Limón ); 2°50′ S, 78°23′ W; 1380 m a.s.l.; 19 Mar. 1997; B. Øllgaard & H. Navarrete 2506; AAU, QCA GoogleMaps • Road Patuca-Santiago, km 23 from Santiago ; 3°01′S, 78°10′W; 830 m a.s.l.; 19 Mar. 1997; B. Øllgaard & H. Navarrete 2492; AAU, QCA GoogleMaps .

PERU – Amazonas • Prov. Bagua, ca 40–43 km (by road) NE of Chiriaco ; [1050–2400 ft a.s.l.]; 7 Nov. 1978; P.J. Barbour 4516; US, USM • Puerto Nazareth , 25 kms – Olmos; 540 m a.s.l.; 22 Dec. 1970; H. Ellenberg 3489; GOET, UC. – Ucayali • Prov. Coronel Portillo [Aguaytia], Rio Chino near Sinchono , between Tingo Maria and Pucallpa ; [ca 9°08′30″ S 75°46′ W]; 1400 m a.s.l.; 2 Aug. 1948; P. Aguilar 904; F GoogleMaps .

Description

Trunkless, or trunk to 0.3 m tall, with old petiole bases, 2–4 cm diam., without adventitious shoots. Leaves erect, to 120(–140) cm long. Petioles 35–70 cm (ca ⅓–½ the leaf length), inermous to sparingly muricate, dark brown to blackish at base, green in upper parts, drying to medium brown to yellowish brown; scurf absent to scant, with ephemeral, dust-like white squamellae. Petiole scales narrowly lanceolate, 3–7 × 1–2 mm, discordantly bicolorous, castaneous to brown with whitish margins, erose with some darker cells interspersed. Blades to 50(–100) × 60(–80) cm, obovate-elliptic, with 4-7 pinna pairs (in fully bipinnate-pinnatifid leaves, 15–20 pairs in simply pinnate leaves); apex abruptly reduced. Leaf axes inermous, abaxially with short hairs 0.2–0.4 mm long, scant ephemeral squamules, and small bicolorous lanceolate scales; costae green alate throughout. Largest pinnae 31(–40) × 9(–13) cm, notably stalked by 1–2.5 cm in lower half, sessile in upper parts; basal pinnae ca ½ the length of longest pinnae, stalked by 3 cm, weakly reflexed. Pinnules to 25(–62) × 12(–17) mm, elongate to oblong, smaller ones subentire, largest ones lobed more than half way (to 4 mm) to the costa; bases truncate, tips round to truncate (rarely attenuate in largest pinnules). Costules abaxially with dark brown spot at the base, with hairs like on the costae, with few to many dark brown bullate squamules to 1.0 mm long. Segments/lobes with entire margins, rounded tips. Veins without hairs, or stronger veins adaxially with single translucent or white hairs; sterile and fertile veins simple in bipinnate parts, also forked in simply pinnate parts. Sori ca 1.2 mm diam., ± medial, forming lines across the segments parallel to costules; receptacles ca 0.2 mm diam., paraphyses few, shorter than sporangia.

Distribution and ecology

Southern Colombia, Ecuador and northern Peru ( Fig. 2A View Fig ) at elevations of 150–1400 m a.s.l., in deeply shaded forest on steep rocky slopes.

Remarks

The blades of Cyathea werffii vary greatly between being pinnate-pinnatifid to fully bipinnate with slightly oblanceolate, blunt-tipped pinnules with entire to coarsely lobed margins. In the general variation of plant size and blade dissection, it comes close to C. pycnocarpa , with which it also shares comparatively small but abundant petiole scales that may have some darker cells in the white margins. Cyathea pycnocarpa differs in having lanceolate to weakly hastate pinnules (i.e., widest at the base and gradually tapering to an acute tip), the blade is basally truncate with the lowest pinnae patent and barely smaller than the longest pinnae (vs blades obovate-elliptic with the lowest pinnae smaller and reflexed in C. werffii ) and the blade apex gradually reduced (vs abruptly reduced). Having seen both species in the field, we can further add that the scaly indument of the axes and veins abaxially tends to be more evident and appear more whitish in C. pycnocarpa than in C. werffii . The laminar texture is thicker and heavier in the former species than in the latter, thus the leaves are arching in C. pycnocarpa and held upright in C. werffii ( Fig. 13A View Fig ), presumably forming a funnel-shape in larger plants.

The occurrence in Colombia ( Cárdenas et al. 2019) is only supported by aberrantly large specimens which look more like plants of C. tortuosa but with bicolorous petiole scales (vs concolorous in C. tortuosa ) and notably stalked pinnae (vs mostly sessile or nearly so). These plants could also be a local variant of C. pastazensis , whose small stature with long stalked pinnae may be an adaptation to the very shady locations where they were found.

The range of C. werffii is sometimes reported to reach southern Peru, but as far as we can confirm, these seem to base on erroneous determinations (also by the first author) of premature plants of other species, like C. pungens and C. tortuosa . We observed Cyathea werffii together with both species at the paratype location in Chiriaco, Bagua, northern Peru (Ellenberg 3489), which allowed us to distinguish between them more clearly ( Fig. 13 View Fig ).

Regarding the general size and shape of the pinnules, the soral line pattern, and composition and color combination of the fine indument, Cyathea werffii is practically identical to C. oblonga from the Guyana highlands, but that species has more numerous and persistent petiole scales that usually reach the lower half of the rachis, only sessile pinnae, and more pinna pairs per leaf (10–12 in C. oblonga vs 4–7 in C. werffii ). The similarity may base merely on convergence, but a closer relationship between these two disjunct taxa is also possible, as the Amotape-Huancabamba-zone, where C. werffii is endemic to ( Fig. 2A View Fig ), has many biogeographic ties with the Guayana highlands ( Lehnert & Tejedor 2016).