Cyathea klotzschiana Domin, 1930

Cyathea klotzschiana Domin View in CoL

Figs 5–6 View Fig View Fig

Cyathea klotzschiana Domin ( Domin 1930: 128) View in CoL . – Replaced synonym: Alsophila obtusa Klotzsch ( Klotzsch 1852: 41) . – Cyathea obtusata Domin ( Domin 1929a: 263) View in CoL , nom. illeg., not Cyathea obtusata Rosenst. ( Rosenstock 1917: 1) View in CoL . – Trichipteris obtusa (Klotzsch) R.M.Tryon ( Tryon 1970: 46) View in CoL .

– Type: VENEZUELA • Without locality, cultivated by Allardt, Berlin; 1852; H. Wagener s.n.; holotype: B [ B 20 0000322 ]!; putative isotype: NY [ 4004118 , labelled “ H. Karsten ”, fragment ex B] image! .

Etymology

The epithet of the replacement name honors the author of the original name, Johann Friedrich Klotzsch (1805–1860), German botanist, curator and director of the Botanical Museum at Berlin Botanical Garden.

Selected material studied.

COLOMBIA – Boyacá • Puerto Boyacá , Vereda La Cunchalita, sitio El Laurel ; 1200 m a.s.l.; 25 Sep. 1996; O. Rangel 13602; COL. – Cundinamarca • Medina, Vereda Periquito ; ca 04°52′11″ N, 74°31′28″ W; 1600 m a.s.l.; 13 Jun. 2017; A. Nassar Arboleda AN070 MEDI-070; COAH. GoogleMaps – Magdalena • Santa Marta Alto, Río Buritaca , Alto de Mira , por el camino a la Quebrada Julepia ; 11.225° N, 74.136° W; 700 m a.s.l.; S. Madriñán 185; COL, FMB, MO. GoogleMaps – Meta • Municipio de Cuabarral, sendero hacia Reserva Natural Las Palmeras ; 3.832° N, 73.888° W; 1230 m a.s.l.; 21 Nov. 2021; J.C. Castro, F. Giraldo & J. Londoño 1700; COAH, HUA. GoogleMaps – Norte de Santander • Hoya del Río Margua, quebrada del Río Negro ; 11.045° N, 73.924° W; 1200–1300 m a.s.l.; J. Cuatrecasas 12921; COL, F, US. GoogleMaps – Santander • Barrancabermeja en Valle del Magdalena, entre Sogamoso y El Rio Colorado ; 4.315° N, 76.363° W; O. Haught 1422; GH, UC, US GoogleMaps .

VENEZUELA – Aragua • North slope, near La Cumbre , Parque Nacional ; 900 m a.s.l.; 29 Nov. 1938; A.H.G. Alston 5790; MO GoogleMaps • Rancho Grande in mountains above Maracay, trail left of road; (toward water) from research station; 10.350° N, 67.685° W; 1000 m a.s.l.; 2 Mar. 1970; R.A.White & N.B. White 19708; DUKE, MICH. GoogleMaps – Carabobo • R. Aguada ; [ca 10°10′ N, 68°09′ W]; 1500 m a.s.l.; 9 Jan. 1939; A.H.G. Alston 6278; MO GoogleMaps • same locality as for preceding; A.H.G. Alston 6279; MO. GoogleMaps – Distrito Federal • Along old road between ‘ Portachueloʼ and ‘ Peñitaʼ (Petaquire) and Carayaca , between Colonia Tovar-Junquito road and Hacienda El Limon , 6–8 mi. below junction of Junquito-Colonia Tovar road; 1300–1500 m a.s.l.; 12 Feb. 1966; J.A. Steyermark & M. Rabe 94779; US. GoogleMaps – Miranda • Ocumare ; ca 10°21′23″ N, 67°43′10″ W; 800 m a.s.l.; 2 Apr. 1937; H.F. Pittier 13955; F, US. GoogleMaps – Yaracuy • En los canelitos cerca del caserio de San José , Aroa; [ca 10°25′ N, 68°53′ W]; ca 200–600 m a.s.l.; 3–4 Jul. 1953; L. Aristeguieta & F. Pannier 1886; US GoogleMaps .

Description

Trunks to 6 m tall, straight to decumbent, 9.0–10.0 cm diam., covered with old petiole bases, due to these sparsely to strongly aculeate; apices hidden between petioles; adventitious buds absent. Leaves 150– 335 cm long, held with petioles erect to ascending, blades ± planar, weakly arching. Petioles 40–50 cm long, sparsely aculeate, prickles to 2 mm long, dark yellowish brown to stramineous, usually basally darker brown, often reddish; aerophores to 20 × 1 mm, in a ± continuous line on each side, inconspicuous in dried material, whitish in fresh material; without remote (aphlebioid) pinnae at the petiole bases; petiole scales 20.0–25.0 × 2.5–5.0 mm, lanceolate to ovate-lanceolate, tapering to linear tips, straight to falcate, weakly twisted, concordantly bicolorous, shiny dark brown to castaneous, with narrow, often abraded whitish margins, scales persistent in distal petiole parts, often reaching lower rachis, here only smaller than lower ones, remaining concordantly bicolorous (never paler or almost completely white); petiole scurf a tomentum of small branched clavate hairs 0.2–0.4 mm long, tan with brown parts, dark brown in general aspect, appearing as small dark dots or irregular stars on paler parts of the epidermis, persistent but easily abraded. Blades 110–290 × 90(–170) cm, bipinnate-pinnatifid, ovate-elliptic to obovate, chartaceous; dark olive-green adaxially, often blackish when dried, olive green abaxially; smaller plants with long petioles and ovate-elliptic blades with 8–10 pinna pairs, basal ones ca ½ the length of longest pinnae, larger plants with short petioles and basally tapering blades with 11–16(–20) pinna pairs, basal ones ca ⅓ the length of longest pinnae, patent to weakly reflexed; pinnae alternate; apices gradually reduced. Leaf axes (rachises, costae and costules) stramineous to dull orange-brown on both sides, sparsely aculate in lower half, otherwise inermous; adaxially with antrorsely curved uniseriate, reddish brown hairs 0.5–1.0 mm long, abaxially with scurf remnants and small, erect curved multicellular hairs to 0.5 mm, costae and costules also with brown to pale brown bullate squamules, 0.5–1.0 × 0.5 mm, often with white flattened or elongate tips, occasionally larger flat squamules to 2 × 1 mm present; junctions rachises/costae abaxially weakly swollen, each with an inconspicuous planar pneumathode, dark brown, elliptic, to 2.5–3.0 × 1.5 mm, area around it often black in dried specimens (foliar nectary). Largest pinnae to 45(–85?) cm long, sessile (appearing stalked if proximal basiscopic pinnule has fallen off), pinnae patent to weakly ascending, distally narrowly to broadly green alate, the pinnatifid terminal segment shortly decurrent into the costae. Largest pinnules 4.5–8.5 × 1.1–1.7 cm, linear-oblong, incised to ½ or more of their width (usually 2 mm between sinuses and costules), sessile, bases truncate to weakly cuneate, with obtuse to short acute tips; costules basally with a black spot or ring going all around their bases (abscission layer); largest segments 8.0–9.0 × 3.0–4.0 mm, oblong, patent to ascending, distally weakly falcate, with entire to weakly dentate margins, tips rounded to obtuse; basal segments alternate, the lowest ones not remote from each other, sinuses narrowly triangular to elliptic, acute, to 1.0 mm wide, sometimes closed; veins prominent abaxially and adaxially, midveins adaxially ridged, veins ending in cartilaginous segment margins; midveins yellowish brown abaxially and adaxially, lateral veins yellowish to greenish brown or blackish, glabrous adaxially except for occasional single hairs on the midveins, abaxially with some bicolorous bullate squamules and short multicellular hairs on midveins, absent between the veins; sterile and fertile veins simple. Sori 0.8–1.0 mm diam., medial to supramedial (sometimes appearing submarginal when sori still complete), parallel to the margins, on the back of veins, indusia absent; receptacles globose to ellipsoid, 0.2–0.3 mm diam.; paraphyses few to numerous, hyaline, brown to tan, of the same length as or shorter than sporangia (0.2–0.5 mm). Spores not examined.

Distribution and ecology

Colombia (Sierra Nevada de Sta. Marta, Cordillera Oriental) and Venezuela at elevations of 600–1700 m a.s.l., predominantly in lower montane forest with Caribbean influence.

Remarks

The type of Alsophila obtusa at Berlin has conflicting label information concerning the provenance (“Columbien” vs “ Venezuela ”) and the collection date (15 Jan. 1852 vs 23 Jul. 1855). The latter conflict may be interpreted as two separate samplings of the same cultivated plant, which were brought to Germany by H. Karsten. Other plants of this species were cultivated at Dresden Botanical Garden (1862, Anonymous s.n.; MO-3304121). The type consists of several pinnae, rachis parts and one petiole, but larger petiole scales or a blade apex are not preserved. The only fertile pinna is mounted face-down but the zig-zag-pattern of the sori can be recognized as impressions thorugh the laminar tissue. The type is also comparatively small and pale. Some representative collections that fill the gaps are Ortega Mendoza & Smith 2492 (NY), which matches the type in size and proportion of pinnae and pinnules, but has a darker, dull orange-brown color in petiole and rachis, and Meier et al. 11303 (UC), which seems to represent a stouter plant with pinnule tips not rounded but tapering to a tounge-shaped lobe.

Cyathea klotzschiana is distinguished in the field from the similar sympatric C. oblonga (both have dark castaneous petiole scales with narrow white margins and blunt pinnules, similar pinnae sizes and numbers) by the different soral lines (submarginal lines, or zig-zag-pattern in C. klotzschiana vs parallel to midveins in C. oblonga ) and color of the laminar indument (pale vs dark brown). Cyathea klotzschiana further has small erect, villose hairs on the costae and costules abaxially; C. oblonga has only brown appressed hairs on the veins and costules, mixed with short ribbon-like squamules.

Cyathea klotzschiana shows distinct variations in its key distinguishing characters related to age, environment, and region. The speciesʼ range spans across the Cordillera de la Costa and the Andes of Venezuela to Colombian Cordillera Oriental and Sierra Nevada de Santa Marta; most of these mountain ranges, though mostly contingent, fall into several subsets of ranges that act as separate biogeographic units, which is supported by the occurrence of local endemic species (e.g., C. barringtonii and C. venezuelensis ; Lehnert & Weigand 2017). This is reflected in regional morphotypes of Cyathea klotzschiana . For example, whereas plants from Serra do Aroa (Yaracuy state, Venezuela) have elongate blunt tipped pinnules with pale laminar squamules, which are here considered best fitting the type specimen, plants to the east in the Cordillera de la Costa have more bicolorous laminar squamules but do not differ in pinnule shape. Specimens from Santa Marta, Colombia, to the west, however, appear superficially different because of their exceptionally long pointed pinnules, but are just as pale-scaled as the plants from Serra do Aroa. Also, the short hairs abaxially on costae and costules can be lost in older leaves, which can make it hard to distinguish such plants of C. klotzschiana from C. pungens with sessile pinnules.

Among the reinstated taxa, Cyathea klotzschiana and C. floribunda are the most similar to each other regarding pinnule shape and laminar indument, with both often having a relatively dense strip of whitish bullate squamules on the costules abaxially. However, the faint leaf dimorphism of C. floribunda is not observed in C. klotzschiana . Both taxa are geographically separated over a large part of the northern Andes by the range of C. pastazensis .