taxonID	type	description	language	source
163C91782808BC38FDAEBA37F35450DE.taxon	description	Description Leaves shallowly bipinnate-pinnatifid to partially tripinnate with narrow to closed sinuses; costae adaxially ± flattened, often green alate, at least with a green edge along each side, veins excurrent into the segment margins; blades often appearing glabrous, without multicellular hairs or hairs <1 mm long, often replaced by unicellular trichomidia, if hairs longer then tortuous and appressed, small bullate and flat squamules may cocurr in varying quantities and persistencies, colors ranging from whitish-gray, yellowish to dark brown, concolorous to bicolorous with white tips, never castaneaous; sori exindusiate (small irregularly shaped scales subtending the receptacle may occur), paraphyses relatively short (shorter than sporangia to slightly longer than closed sporangia), their tips often hyaline, fragile and missing in mature sori, never contorted; sporangia relatively few, ca 20 – 30 per sorus, dark orange-brown.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782806BC3BFD9CBB5BF5E05065.taxon	description	Fig. 3	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782806BC3BFD9CBB5BF5E05065.taxon	materials_examined	– Type: PERU – Huánuco • “ In Peruvia, ” Cochero; 1780; J. Dombey s. n.; lectotype: P [P 00642343]!, designated by Lehnert 2016: 36; isolectotypes: B [B 20 0000123 b]!, P [P 00642344, P 00642345]!.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782806BC3BFD9CBB5BF5E05065.taxon	materials_examined	– Type: PERU – Huánuco • Prov. Huánuco, Ventanilla de Cassapi; Jul. 1829; E. Poeppig s. n.; holotype: LZ [destroyed]; lectotype: W [W 0053351]!, designated by Lehnert 2016; isolectotypes: LE [LE 00008089] image!, MO [MO- 1858413] image!, P [P 00642342]!, W [W 0053352]!.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782806BC3BFD9CBB5BF5E05065.taxon	materials_examined	– Type: PERU • “ Ad Tarma ”; Ruíz Herb. 66; lectotype: B [B 20 0000123 a]!, designated by Lehnert 2016; isolectotypes: GH [fragment of B] n. v., NY [00148737, fragment of B]!.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782806BC3BFD9CBB5BF5E05065.taxon	etymology	Etymology The specific epithet honors Joseph Dombey (1742 – 1794), French botanist and the collector of the type specimen.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782806BC3BFD9CBB5BF5E05065.taxon	materials_examined	Selected material studied PERU – Amazonas, Prov. Bagua • Distrito Imaza, Yamayakat; 5 ° 03 ′ 20 ″ S, 78 ° 20 ′ 23 ″ W; 400 m a. s. l.; 18 Nov. 1996; R. Vasquez, P. Stern, R. Rojas & R. Aguilar 21824; MO, UC. – Huánuco, Prov. Leoncio Prado • Near confluence of Río Cayumba with Huallaga; 875 m a. s. l.; 14 Oct. 1936; Y. Mexia 8293; UC • Tingo Maria; 700 m a. s. l.; 2 Sep. 1956; R. M. Tryon & A. F. Tryon 5256; BM. – Pasco, Prov. Oxapampa • Dist. Villa Rica, zona de amortiguamiento del Parque Nacional Yanachaga-Chemillén; 10 ° 39 ′ 17 ″ S, 75 ° 10 ′ 39 ″ W; 1290 m a. s. l.; 15 Apr. 2006; A. Monteagudo et al. 11882; MO. – San Martin, Prov. San Martin • near Tarapoto; 6 ° 26.832 ′ S, 76 ° 17.741 ′ W; 1050 – 1100 m a. s. l.; 3 Nov. 2010; M. Lehnert 1998; HAL, USM • In montibus secus flumen Mayo, prope Tarapoto; Jul. – Aug. 1856; R. Spruce 4715 [excluded syntypes of Alsophila floribunda]; B [B 20 0000122], BM [BM 000777032 (mix with C. pungens)], P [P 00642346 (mix with C. pungens), P 00642351]. BOLIVIA – La Paz, Prov. Larecaja • 6 – 10 km E of Consata along new road; 15 ° 15 ′ S, 68 ° 30 ′ W; 1350 – 1400 m a. s. l.; 14 Dec. 1981; C. R. Sperling et al. 5437; BM, GH, MO, US. – Cochabamba, Prov. Chapare • 159 km antigua carretera Cochabamba-Villa Tunari; 17 ° 05 ′ S, 65 ° 31 ′ W; 700 m a. s. l.; 7 Sep. 1997; M. Kessler 8221; GOET, UC.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782806BC3BFD9CBB5BF5E05065.taxon	description	Description Trunks 0.5 – 2.4 (– 4.0) m tall, 5.0 – 12.0 (– 15.0) cm diam., straight, with persistent old petiole bases, spiny; apices hidden between petioles; adventitious buds absent. Leaves to 270 cm long, erect to weakly arching. Petioles 60 – 100 cm long, aculeate, prickles 3 – 5 mm long, dark brown; pneumathodes to 10 × 1 mm, inconspicuously brown in dried material, whitish in fresh material; without adventitious (aphlebioid) pinnae at the petiole bases; petiole scales lanceolate, 25.0 – 32.0 × 2.5 – 4.0 mm, their tips straight to falcate, twisted, concordantly to discordantly bicolorous, shiny dark brown, with whitish margins and regular dark brown teeth; colors mostly 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, white in general aspect, soon caduceus, persistent between prickles. Blades 100 – 160 × 60 – 100 cm, bipinnate-pinnatifid, elliptic to obovate, chartaceous to subcoriaceous; dark olive-green adaxially, often blackish when dried, olive-green abaxially; apices ± gradually reduced. Rachises inermous or proximally aculeate, stramineous to yellowish brown on both sides; adaxially with antrorsely curved uniseriate hairs 0.5 – 1.0 mm long, abaxially glabrous except for scurf remnants, containing appressed, white trichomoidia and dissected squamules (0.2 – 0.5 mm). Largest pinnae 50 – 65 cm long, pinnae 10 – 15 pairs per leaf, sessile to stalked to 0.5 cm, alternate, ascending, distally broadly green alate, the distal segments decurrently adnate before ending in pinnatifid terminal segments; basal pinna pairs much smaller than medial pinnae, reflexed. Costae inermous, to 2.0 – 3.0 mm wide, adaxially with antrorsely curved uniseriate hairs 0.5 – 1.0 mm long, abaxially with scurf like on the rachises and scattered pale brown squamules, insertions into rachises abaxially weakly swollen, each with an inconspicuous planar pneumathode, dark brown, elliptic, to 2.0 × 1.0 mm, area around it often black in dried specimens. Largest pinnules 7.5 – 9.0 (– 13.5) × (1.3 –) 1.5 – 1.8 (– 2.0) cm, sessile to subsessile, articulate, alternate, 1.5 – 2.0 cm between the stalks, linear-oblong, incised to ½ or more of their width, bases truncate to weakly cordate, tapering from the middle to attenuate tips; costules with flat lanceolate bicolorous scales to 3 × 2 mm and bullate squamules to 0.5 – 1.0 mm long with entire margins and long acute tips, varying in color from tan to deep brown, usually with white tips; costules basally with a black ring going all around their bases (abscission layer); segments oblong, to 5.0 – 15.0 × 3.0 (– 7.0) mm, patent to ascending, straight to distally weakly falcate, with entire to weakly dentate margins, tips rounded to obtuse, rarely acute and then margins weakly dentate; basal segments opposite to alternate, the lowest ones not remote from each other, sinuses triangular, acute, 1.0 – 2.0 mm wide, sometimes occluded; veins prominent abaxially and adaxially, midveins strongly so and adaxially ridged, lateral veins ending at 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 glabrous or with squamules and trichomidia like on the costules (squamules on midveins darker or more strongly bicolorous than those on the costules), trichomidia sometimes also between the veins; sterile and fertile veins mostly simple, rarely forked. Sori (0.8 –) 1.0 – 1.5 mm diam., medial and parallel to the midveins, or tapering to a subproximal position towards the segment tips (V-shape), on the back of veins, indusia absent; receptacles globose to ellipsoid, 0.3 – 0.4 mm diam.; paraphyses few to numerous, hyaline, tan to brown, shorter than sporangia (0.2 – 0.3 mm). Spores not examined.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782806BC3BFD9CBB5BF5E05065.taxon	distribution	Distribution and ecology Peru and Bolivia (Fig. 2 A) at elevations of 700 – 2000 m a. s. l. in the understory of humid mountain forests.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782806BC3BFD9CBB5BF5E05065.taxon	discussion	Remarks Cyathea dombeyi and C. pungens differ significantly in the overall appearance of the plants and the shape of the pinnules (compare Figs 3 and 8). In fresh material with intact sori, the different position of the soral lines in these two species is even more evident, which are closer to the midveins in C. dombeyi and closer to the margins in C. pungens. The type of Alsophila floribunda (Spruce 4715) was identified as a mixture of different species by Barrington (1978); the specimen P 00642346 has a corresponding label, excluding the upper left pinna as belonging to C. pungens (as Trichipteris procera) and referring the rest correctly to C. dombeyi. However, the Kew material belongs to a third morphospecies with remotely postioned, blunt pinnules and abundant whitish bullate squamules below on veins and costules, which is clearly the condition described by Hooker & Baker (1874) for A. floribunda. We separate this taxon from C. dombeyi and transfer with it another synonym, Alsophila bulligera Rosenst. More material of the type collection of A. floribunda (Spruce 4715) can be identified as C. pycnocarpa, another synonym of C. dombeyi here reinstated. This means that the otherwise reliable Richard Spruce mixed at least four invididuals, each representing a different morphotype (and likely true species), under one number.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782805BC3EFDC7BAE5F4505621.taxon	description	Fig. 4	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782805BC3EFDC7BAE5F4505621.taxon	materials_examined	– Type: PERU • San Martín, Prov. San Martín, Mt. Campana; Aug. 1855 (isolectotypes Dec. 1855); R. Spruce 4715; lectotype: K [K 000227562]!, designated by Lehnert 2016; isolectotypes: K [K 000227563]!, W [W-Rchb. 1889 - 0198979]!. Excluded syntypes are discussed below.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782805BC3EFDC7BAE5F4505621.taxon	materials_examined	– Type: BOLIVIA • La Paz, Prov. Larecaja, San Carlos (Mapiri Region); 850 m a. s. l.; 30 Mar. 1927; O. Buchtien 288; lectotype: S [S-R- 199] image!, designated by Lehnert 2016; isolectotypes: GH [01100756]!, NY [00148642]!, US [01100756]!.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782805BC3EFDC7BAE5F4505621.taxon	etymology	Etymology The epithet refers to the abundant sori, which used to be likened to flowers or fruits.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782805BC3EFDC7BAE5F4505621.taxon	materials_examined	Selected material studied ECUADOR – Prov. Morona-Santiago, Cantón Gualaquiza • Off main road south of Cualaquiza, at turn-off to Bomboiza; 3.45 ° S, 78.55 ° W; 800 m a. s. l.; 8 Feb. 1993; A. Fay & L. Fay 4219; MO. – Prov. Zamora-Chinchipe, Cantón El Pangui • Campamento Shaime (Shaimi) along Río Nangaritza, trail to the oil bird cave (“ cueva de los tayos ”); 4 ° 19 ′ S, 78 º 40 ′ W; 950 – 1050 m a. s. l.; 6 Nov. 2004; M. Lehnert 1524; GOET, QCA, UC. PERU – Amazonas, Prov. Bagua • Aramango, above Nueva Esperanza; 5.3558333 ° S, 78.3733334 ° W; 1271 m a. s. l.; 1 Nov. 2012; H. van der Werff, L. Valenzuela G., G. Shareva M. & A. Reyes Barrantes 24820; MO, UC. – Junín, Prov. Junín • Pichis Trail, Yapas; 10.834 ° S, 75.210 ° W; 1350 – 1600 m a. s. l.; 28 Jun. 1929; E. P. Killip & A. C. Smith 25536; NY, US. – San Martín, Prov. Rioja • Along road Rioja-Pedro Ruiz, in area with limestone; 5.674 ° S, 77.676 ° W; 1170 m a. s. l.; 23 Mar. 1998; H. van der Werff, B. Gray, R. Vásquez & R. Rojas 15511; MO, NY, UC. • Tarapoto; 1855 – 1856; R. Spruce 4724; E, K. BOLIVIA – Cochabamba, Prov. Carrasco • 149 km antigua carretera Cochabamba-Villa Tunari; 17.117 ° S, 65.567 ° W; 1000 m a. s. l.; 31 Aug. 1996; M. Kessler 7993 with J. Gonzales; T. Krömer, A. Acebey, B. Hibbits, & M. Sonnentag; LPB, UC. – La Paz, Prov. Franz Tamayo • Calabatea, 44 km SW of Apolo, ca 10 km S of Correo, drainage of Rio Yuyo, Transect 1; 14 ° 55 ′ S, 68 ° 20 ′ W; 1500 – 1550 m a. s. l.; 4 Jun. 1990; A. Gentry & St. G. Beck 70926; LPB, MO.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782805BC3EFDC7BAE5F4505621.taxon	description	Description Trunks 0.5 – 3 m tall, 4.0 – 15.0 cm diam., straight to decumbent, covered with old petiole bases, due to these sparsely to strongly aculeate; apices hidden between petioles; adventitious buds absent. Leaves 125 – 170 cm long, held with petioles erect to ascending, blades ± planar, weakly arching. Petioles 36 – 46 cm long, dark yellowish brown to stramineous, basally brown to dark brown, sparsely aculeate, prickles to 2 mm long; 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 scurf a tomentum of small squamellae 0.2 – 0.4 mm long, tan with brown parts, brown in general aspect, appearing as small flakes on the epidermis, persistent but easily abraded. Petiole scales 11.0 – 25.0 × 2.2 – 3.8 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 not persistent in distal petiole parts, only scattered her and on lower rachis. Blades to 115 × 83 cm, bipinnate-pinnatifid, ovate-elliptic to obovate, chartaceous; dark olive-green adaxially, 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 10 – 16 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 yellowish brown on both sides, rachises sparsely aculeate 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 whitish to pale brown bullate squamules, 0.5 – 1.0 × 0.5 mm with subulate tips, larger flat squamules absent; 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 41.5 cm long, sessile (appearing stalked if proximal basiscopic pinnule fallen off), pinnae patent to weakly ascending, distally narrowly to broadly green alate, the pinnatifid terminal segment shortly decurrent into the costae. Largest pinnules 45 – 60 × 11 – 14 mm, linear-oblong, incised to ¾ of their width (usually 2 mm between sinuses and costules), sessile, bases truncate to weakly cuneate, tips obtuse to long acute (rarely attenuate), largest segments 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 subopposite, 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 pale 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 midveins, indusia absent; receptacles globose to ellipsoid, 0.2 – 0.3 mm diam.; paraphyses few to numerous, hyaline, tan, shorter than sporangia (0.2 – 0.5 mm). Spores not examined.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782805BC3EFDC7BAE5F4505621.taxon	distribution	Distribution and ecology Southern Ecuador, Peru and Bolivia (Fig. 2 B) at elevations of ca 800 – 1550 m a. s. l. in the understory of humid mountain forests.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782805BC3EFDC7BAE5F4505621.taxon	discussion	Remarks Excluded syntypes of Alsophila floribunda belong to following species: Cyathea dombeyi: B [B 20 0000122]!, BM [BM 000777032 p. p. (mix with C. pungens): “ In montibus secus flumen Mayo, prope Tarapoto; Jul. – Aug. 1856 ”] image!, P [P 00642346 p. p. (mix with C. pungens)]!, P [P 00642351]! (with scales on rachis) Ɨ rounded segments, pointed pinnules, indument flakey, flat. Cyathea pungens: BM [BM 000777031, BM 000777032 p. p. (mix with C. dombeyi), BM 000777346: “ In montibus secus flumen Mayo, prope Tarapoto; Jul. – Aug. 1856 ”] image!, P [P 00642346 p. p. (mix with C. dombeyi): “ Julius 56 ”]!, P [P 00642349: “ In montibus secus flumen Mayo, prope Tarapoto; Jul. – Aug. 1856 ”]!, P [P 00642350]!, W [W-Rchb. 0053380: “ 4715 var, in monte Campana pr. Tarapoto, (…) Aug. 56 ”]! Ɨ pointed, sessile pinnules, hardly any indument. Cyathea pycnocarpa: “ 4715 forma pygmea, in m. Pingullu-Urku [ca 6.356 ° S, 76.688 ° W, ca 800 – 1500 m a. s. l.], Aug. 56 ”; P [P 00642347, P 00642348]! Ɨ with pale sub-bullate squamules in rachis axil, lower costules with dark brown bullate and flat squamules, the larger ones with white margins; petiole scales with relatively wide white margins, dark brown teeth scattered. The isolectotype has the original label by Spruce stating “ I have not seen this growing; two imperfect fronds were brot. [sic] me - they seemed to have been about 4 feet long ”. The amount and variation of the existing material suggests that Spruce has used more than these two original leaves. Described by Hooker as having sessile blunt pinnules incised ⅓ – ½, medial sori and abundant minute pale scales on the veins and rachis abaxially. The excluded type material differs in one or more of these characters: Cyathea pycnocarpa has almost subentire pinnules; Cyathea dombeyi has such deeply incised pinnules but with tapering tips and the squamules below are not as notably pale as in C. floribunda. This mixture probably answers the question how Spruce managed to generate so many duplicates of each number. In this particular case, the different collection dates and localities are clear indicators that several individuals were combined under one catalogue number. The type material of the synonym Alsophila bulligera lacks the petiole (as does that of Alsophila floribunda) but at least enough material is present to indicate a tapering blade base and a more or less gradually reduced blade apex. The type collections of both names appear very differenct at first glance, because most pinnules in A. floribunda are remote from each other whereas those of A. bulligera are close, with almost no gap between them. In pinnule length and distance as measured from costule to costule, however, both types are very similar. Fertile pinnules tend to be narrower than sterile ones in this taxon, and the different appearances stem from the fact that the material of A. floribunda is fully fertile, and that of A. bulligera predominantly sterile.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782800BC22FDBABC22F454534D.taxon	description	Figs 5 – 6	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782800BC22FDBABC22F454534D.taxon	etymology	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 AN 070 MEDI- 070; COAH. – 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. – 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. – 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. – Santander • Barrancabermeja en Valle del Magdalena, entre Sogamoso y El Rio Colorado; 4.315 ° N, 76.363 ° W; O. Haught 1422; GH, UC, US. VENEZUELA – Aragua • North slope, near La Cumbre, Parque Nacional; 900 m a. s. l.; 29 Nov. 1938; A. H. G. Alston 5790; MO • 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. – Carabobo • R. Aguada; [ca 10 ° 10 ′ N, 68 ° 09 ′ W]; 1500 m a. s. l.; 9 Jan. 1939; A. H. G. Alston 6278; MO • same locality as for preceding; A. H. G. Alston 6279; MO. – 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. – 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. – 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.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782800BC22FDBABC22F454534D.taxon	description	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.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782800BC22FDBABC22F454534D.taxon	distribution	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.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782800BC22FDBABC22F454534D.taxon	discussion	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.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281CBC23FDACB9CEF5CA57F5.taxon	materials_examined	– Type: VENEZUELA • Prov. Amazonas, Dept. Río Negro, 0 – 1 km E of Cerro de la Neblina Base Camp on Rio Mawarinum; 0 ° 50 ′ N, 66 ° 10 ′ W; 140 m a. s. l.; 2 Sep. 1984; R. L. Liesner & V. A. Funk 15781; holotype: UC [UC 1506573]!; isotype: MO [MO- 1873835]!.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281CBC23FDACB9CEF5CA57F5.taxon	etymology	Etymology The specific epithet refers to the type locality.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281CBC23FDACB9CEF5CA57F5.taxon	materials_examined	Selected material studied VENEZUELA – Amazonas, Dept. Río Negro • 0 to 1 km east of Cerro de La Neblina Base Camp on Río Mawarinuma; 0 º 50 ′ N, 66 º 10 ′ W; 140 m a. s. l.; 9 Feb. 1984; R. L. Liesner 15798; MO, L, NY, UC, US. – Bolívar, Dept. Bolívar • Riberas del Río Canaracuni (aguas negras) y selvas adyacentes, Expedición Proyecto I. R. N. R. G. a la cuenca alta del Rio Caura (Hoja NB- 20 - 14), convenio UNELLEZ-C. V. S. (TECHIN, C. A.); 4.450 ° N, 64.117 ° W; 13 Apr. 1988; B. Stergios 11892; MO, UC. BRAZIL – Amazonas, Mun. Santa Isabel do Rio Negro • Neblina massif; 1300 m a. s. l.; 24 Dec. 2003 – 5 Jan. 2004; F. A. Carvalho et al. 308; INPA • same locality as for preceding; F. A. Carvalho et al. 320; INPA.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281CBC23FDACB9CEF5CA57F5.taxon	description	Description Trunkless or trunks 0.3 m tall, decumbent, (1.0 –) 2.5 – 4.0 cm diam., with persistent old petiole bases, spiny; apices hidden between petioles; adventitious buds absent. Leaves to 90 (– 100) cm long, ca 3 – 5 in a crown; presumably arching. Petiole to 25 cm long with prickles 2 – 5 mm long, dark yellowish brown to dull orange-brown, sometimes basally darker brown; aerophores to 10 × 1 mm, inconspicuously brown in dried material, without remote (aphlebioid) pinnae at the petiole bases; petiole scales lanceolate, 8.0 – 9.0 × 2.5 mm, their tips straight to falcate, concordantly to discordantly bicolorous, shiny dark brown with whitish margins at petiole base, scales in distal half of the petiole gradually becoming paler with wider white margins, scales only sharply contrasted in scales near petiole base and on crosiers; 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 prickles. Blades 58 × 46 – 50 cm, pinnate-pinnatifid to bipinnate-pinnatifid, elliptic to ovate-lanceolate, chartaceous; blackish adaxially when dried (probably dark olive-green when fresh), olive-green abaxially; apices gradually reduced. Rachises dark stramineous to orange-brown on both sides, in basal half often with some strong prickles; adaxially with antrorsely curved uniseriate hairs 0.5 – 1.0 mm long, abaxially glabrous except for scurf remnants, containing appressed, white trichomoidia and dissected squamules (0.2 – 0.5 mm). Largest pinnae to 23 cm long, pinnae (6 –) 8 – 12 pairs per leaf (in fully bipinnate-pinnatifid blades, may be higher in less divided ones), subsessile to stalked to 1 cm, ascending, narrowly green alate, the distal segments adnate, decurrent into the costae, forming a pinnatisect section with long acute to attenuate tips. Costae inermous, to 1.5 – 2.0 mm wide, adaxially with antrorsely curved uniseriate hairs 0.5 – 1.0 mm long, abaxially glabrous or glabrescent with scurf like on the rachises, insertions into rachises abaxially weakly swollen, each with an inconspicuous planar pneumathodes, dark brown, elliptic, to 2.0 × 1.0 mm, area around it often black in dried specimens. Pinnules 1.3 – 6.2 × (0.4 –) 0.8 – 1.3 cm, smaller ones sessile, larger ones stalked 1.0 mm, articulate, alternate, ca 1.0 – 1.5 cm between costules, linear-oblong to weakly elliptic, bases notably cuneate, tapering from beyond the middle, tips blunt in entire pinnules, long actue to attenuate to long acute to attenuate in pinnatifid pinnules, also caudate in pinnitisect pinnules; costules with ephemeral tan to brown trichomidia and whitish squamules 0.5 – 1.0 mm long with entire margins andacute tips; costules basally with a black ring going all around their bases (abscission layer); segments oblong, to 7.0 × 3.0 mm, strongly ascending, distally weakly falcate, with entire to subentire margins, tips obtuse to acute and then margins weakly dentate; basal segments alternate, sinuses in pinnatifid pinnules triangular, acute, 1.0 – 2.0 mm wide, in pinnatisect to partially pinnate pinnules the lowest segments remote from each other, separated by angular sinusus; veins prominent abaxially and adaxially, midveins strongly so and adaxially ridged, lateral veins ending at segment margins; midveins yellowish brown abaxially and adaxially, lateral veins yellowish to greenish brown or blackish; adaxially glabrous except for occasional single hairs on the midveins, abaxially glabrous or with squamules and trichomidia like on the costules; sterile and fertile veins mostly forked in bipinnate or less dissected blades, simple and forked in more strongly dissected blades and larger pinnules (> 1 cm wide). Sori 0.6 – 0.8 mm diam., supramedial, parallel to the margins, on the back of veins, indusia absent; receptacles globose to ellipsoid, 0.2 mm diam.; paraphyses few hyaline, white, shorter than sporangia (0.2 – 0.3 mm). Spores not examined. See Smith (1990) for an illustration.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281CBC23FDACB9CEF5CA57F5.taxon	distribution	Distribution and ecology Venezuela and Brazil, mainly around Cerro de la Neblina except for one outlying record from the southern base of Cerro Sarisariñama, mountain forests and scrub vegetation; at elevations of 100 – 1400 m a. s. l.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281CBC23FDACB9CEF5CA57F5.taxon	discussion	Remarks This species was described as being only pinnate-pinnatifid to partially bipinnate; later records from Brazil (F. A. Caravalho et al. 320, INPA) have bipinnate-pinnatifid blades with stalked pinnules (Carvalho et al. 2012), ultimately giving Cyathea neblinae the same variability in laminar disscection as C. pungens s. str .. Both species are also practically identical in petiole scale shape and color, laminar texture and color of the leaf axes, thus C. neblinae might be interpreted as a special dwarf form of C. pungens. However, there are some constant characters that make C. neblinae visibly different: the blade apex seems to be always gradually reduced (vs mostly subconform in C. pungens); all plants described as C. neblinae (Smith 1990) have pinnules with more strongly cuneate bases than C. pungens, irrespective of their size and degree of further dissection. In parts of the blade that mark the transition between simply pinnate and fully bipinnate, C. neblinae has remote adnate segments that are separated wide, assymmetric angular sinuses, formed by the decurrently alate bases at the basiscopic side of the segments. In direct comparison, such transitory sinuses are closer and more symmetric in C. pungens.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281DBC27FD93BE75F24D518D.taxon	description	Figs 7 – 8	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281DBC27FD93BE75F24D518D.taxon	materials_examined	– Type: GUYANA • R. Schomburgk 1125; lectotype: K [K 000589887]!, designated by Barrington 1978; isolectotypes: B [B 20 0000323 a, B 20 0000325]!, BR [BR 0000006987886] image!, LE [LE 00008119] image!, NY [00148722, fragment of K] image!, P [P 00631669]!, US [00066269] image!.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281DBC27FD93BE75F24D518D.taxon	materials_examined	– Type: FRENCH GUIANA • “ In Gujana gallica, ” vicinity of Saül, ca 10 km NW from Eaux Claires; ca 03 ° 37 ′ N, 53 ° 12 ′ W; 400 m a. s. l.; G. S. Perrottet s. n.; lectotype: PR!, designated by Lehnert 2016; isolectotype: NY [0014874, fragment of PR] image!.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281DBC27FD93BE75F24D518D.taxon	materials_examined	– Type: GUYANA [BRITISH GUIANA] • Near Rockstone, in dense up-land forest; ca 05 ° 58 ′ 26 ″ N, 58 ° 31 ′ 09 ″ W; ca 100 – 120 m a. s. l.; 15 Jul. – 1 Aug. 1921; H. A. Gleason 830; holotype: US [00066252] image!; isotypes?: NY [00148678] image!, US [00715386] image!.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281DBC27FD93BE75F24D518D.taxon	etymology	Etymology The specific epithet refers to the shape of the pinnules.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281DBC27FD93BE75F24D518D.taxon	materials_examined	Selected material studied COLOMBIA – Antioquia • Vereda Providencia, Anorí, 500 m arriba de la carretera por el camino de la retroescabadora desde la casa de Doña María; 7.3483333 ° N, 75.0208333 ° W; 300 m a. s. l.; 15 Jun. 2011; J. Colorado 732; HUA. VENEZUELA – Amazonas, Dept. Río Negro • Cerro de la Neblina, Río Yatua, 6 – 8 km S of Camp 3; 1.03 ° N, 65.93 ° W; 1200 – 1600 m a. s. l.; 24 Dec. 1953; B. Maguire, J. J. Wurdack & G. S. Bunting 36870; US. – Bolivar, Mun. Bolivariano Angostura [= Raul Leoni] • 50 km al SW de Guaiquinima y 72 km al W de Karún; 5 ° 18 ′ N; 63 ° 59 ′ W; 230 m a. s. l.; A. Fernández & G. Aymard 4904; MO, UC. – Distrito Federal, Mun. Libertador • Cordillera de la Costa, al noreste de Guatire, excursion Fila Juan Torres-Fila Las Perdices por el Rio Guayabal hacia el pueblo Guayabal; 10.516667 ° N, 66.333333 ° W; 950 – 1000 m a. s. l.; 19 Feb. 1992; W. Meier 3415; UC. GUYANA – Cuyuni-Mazaruni • Mt. Maringma, slope below sub-summit plateau; 5.20461 ° N, 60.5766 ° W; 1370 m a. s. l.; 22 Jun. 2004; H. D. Clarke, C. Perry, E. Tripp, S. R. Stern & D. Gittens 11931; US. – Potaro-Siparuni • Iwokrama Rainforest Reserve, Karupukari-Annai Road, summit of unnamed peak at end of 5 km transect; 4.471 ° N, 58.788 ° W; 700 m a. s. l.; 19 Mar. 1997; H. D. Clarke, S. A. Mori & S. Heald 4120; US. – Upper Takutu-Upper Essequibo • saddle between tops of Two-Head Mt, Kanuku Mts; 4.206 ° N, 59.117 ° W; 580 m a. s. l.; 2 Feb. 1994; M. J. Jansen-Jacobs, B. J. H. ter Welle, A. Chanderbali, U. Raghoenandan & V. James 3535; NY, US. SURINAME – Marowijne • Marowijne [river], Nassau Mts, Plateau C; 4.83 ° N, 54.61 ° W; 500 – 550 m a. s. l.; 5 Feb. 2003; M. J. Jansen-Jacobs et al. 6556; U, UC, US. – Sipaliwini • Upper reaches of Grace Kreek, Tafelberg Mountain, headwaters of the Grace Kreek drainage; 3.893 ° N, 56.160 ° W; 700 – 790 m a. s. l.; 5 Jul. 1998; T. Hawkins 1892; L, MO, UC, US. FRENCH GUIANA – Cayenne • D. Z. de Crique Jupiter, bassin du Sinnamary; 4.06 ° N, 53.16 ° W; 100 m a. s. l.; 25 Apr. 1991; J. J. de Granville 11511; CAY, L, NY, UC, US. – Saint-Laurent-du-Maroni • Mont Atachi Bacca, Region de lʼInini; 3.55 ° N, 53.91666 ° W; 500 m a. s. l.; 14 Jan. 1989; G. Cremers 10227; MO, NY. BRAZIL – Amapá • Rivière Haut [= upper Río] Jari; 2.46 ° N, 54.76 ° W; 380 m a. s. l.; 19 Aug. 1993; J. J. de Granville, P. Acevedo-Rodríguez, A. Boyer & L. A. Hollenberg 12381; US. – Pará • Mun. Oriximiná Estação Ecológica Grão Pará, Córego próximo do acampamento; 1 ° 16 ′ 47.4 ″ N, 58 º 41 ′ 28.5 ″ W; 475 m a. s. l.; 30 Aug. 2008; S. Maciel 983; MCT, STU. – Roraima • Lower slopes of Serra da Lua, by Igarape Camarao; 2 ° 25 – 29 ′ N, 60 ° 11 – 14 ′ W; 23 Jan. 1969; G. T. Prance, J. R. Steward, F. Ramos & L. G. Farias 9400; F, NY.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281DBC27FD93BE75F24D518D.taxon	description	Description Trunks 0.8 – 2.0 (– 8.0?) m tall, straight to decumbent, 4.0 – 5.0 cm diam., with persistent old petiole bases, sparsely aculeate; apices hidden between petioles; adventitious buds absent. Leaves to 165 cm long, held with petioles erect to ascending, blades ± planar, weakly arching. Petioles 27 – 44 (– 53) cm long, sparsely to strongly aculeate, prickles 2 – 3 mm long, dark yellowish brown to stramineous (Andes), or dark reddish brown (Guayana shield); aerophores to 10 × 1 mm, inconspicuously brown in dried material, whitish in fresh material; without adventitious (aphlebioid) pinnae at the petiole bases; petiole scales 15.0 – 20.0 (– 23.0) × 2.5 – 4.0 (– 5.0) mm, 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, soon caduceus, persistent between spines. Blades 80 – 90 × 70 – 80 cm, bipinnate-pinnatifid, elliptic to obovate, chartaceous; dark green adaxially, often blackish when dried, olive green abaxially; (8 –) 10 – 12 (– 24?) pinna pairs, alternate, basal ones ½ – ⅓ the length of longest pinnae, weakly to strongly reflexed; apices abruptly reduced to non-conform apical sections, sometimes very broad and nearly gradually reduced. Leaf axes (rachises, costae and costules) stramineous to yellowish brown on both sides, often with a reddish tinge, inermous; adaxially with antrorsely curved uniseriate hairs 0.5 – 1.0 mm long (sparse to absent on costules), abaxially glabrous except for scurf remnants, containing appressed branched brown hairs (like on petioles), dark brown bullate squamules, 0.5 – 1.0 × 0.5 mm with subulate, rarely flattened or elongate tips (sparse to absent on rachis); junctions rachises / costae abaxially weakly swollen, each with an inconspicuous planar pneumathode, dark brown, elliptic, to 2.0 × 1.0 mm, area around it often black in dried specimens. Largest pinnae 37 – 45 cm long, sessile or stalked to 1.5 cm (proximal basiscopic pinnule reduced or fallen off), pinnae patent to weakly ascending, distally narrowly to broadly green alate, the pinnatifid terminal segment shortly decurrent into the costae. Largest pinnules 5.0 – 8.0 × 1.3 – 1.5 cm, linear-oblong, incised to ½ or more of their width (usually 2 mm between sinuses and costules), sessile, bases truncate to weakly cuneate, tapering from beyond the middle to obtuse to acute (rarely short-attenuate) tips; costules basally with a black ring going all around their bases (abscission layer), basally without aerophore; largest segments 8.0 – 10.0 × 2.5 – 4.5 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, acute, to 1.0 – 2.0 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 glabrous or with squamules and trichomidia like on the costules, trichomidia sometimes also between the veins; sterile and fertile veins mostly simple, rarely forked. Sori 0.8 – 1.0 mm diam., appearing ± medial when intact, closer to the margins when reduced to receptacles, parallel to the margins / midveins, on the back of veins, indusia absent; receptacles globose to ellipsoid, 0.2 – 0.3 mm diam.; paraphyses few to numerous, hyaline, white to tan, shorter than sporangia (0.2 – 0.3 mm). Spores not examined.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281DBC27FD93BE75F24D518D.taxon	distribution	Distribution and ecology Colombia, Venezuela, Guyana, Suriname, French Guiana, and northern Brazil (Fig. 2 B) at elevations of (100 –) 140 – 1400 (– 1600) m a. s. l., in lowland rain forests on terra firme and premontane forest with Caribbean influence.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178281DBC27FD93BE75F24D518D.taxon	discussion	Remarks The maximum reported height for the plant is 8 m (Maguire et al. 29929), which is exceptionally high for the species and likely erroneous. The highest number of pinna pairs is twice the average for the species and might have been intended as “ 24 pinnae in pairs ”. Cyathea oblonga was resurrected from the synonymy of a loosely conceived C. pungens by Cremers & Boudrie (2007) based on their field observations in French Guiana. Here, both species overlap in their distribution, and show consistent differences in the anatomy and distribution of the petiole scales (relatively narrow, shiny dark brown with white margins, all along the petioles without getting paler in C. oblonga vs petiole scales usually restricted to petiole bases or if reaching further up then becoming paler to almost white in C. pungens) and the laminar indument (bullate squamules and scurf remnants on costae and costules brown to dark brown, well represented vs scurf and bullate squamules pale brown to white, scant). Cyathea oblonga from higher elevations at the tablemountain bases differs from those in the lowlands of eastern Guayanan in having the axes abaxially not atropurpureous but stramineous to orange brown, and being stouter plants with thicker axes and larger pinnules. Plants here reported from northern Venezuela and the Andes of Colombia look more like the stouter tablemountain variety, and although having the diagnostic characters of that species (± medial sori, narrow petiole scales extending to the rachis), they may turn out to be aberrations of other taxa (mostly C. klotzschiana, presumably) after more scrutinous investigation in the field. Precociously fertile plants with less divided blades than regular fertile plants are the basis for the synonym Alsophila gleasonii. Here, also the petiole scales tend to be smaller, but full in proportion to the smaller size of the plants. These individuals look like non-sympatric dwarf species of Cyathea, such as C. pseudonanna Lellinger from Panama, which is best distinguished by the denser, stiffer hairs abaxially on the rachises and the bluish sheen of the adaxial laminar side in fresh material.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782819BC2AFDECBB8DF289542F.taxon	description	Fig. 9	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782819BC2AFDECBB8DF289542F.taxon	materials_examined	– Type: ECUADOR • Prov. Tungurahua, between Baños and Jivaría de Pintuc, Río Pastaza Valley; 1870 – 1874; A. Stübel 995 a; lectotype: B [B 20 0000321]!, designated by Lehnert 2016; isolectotype: NY [00148730, fragment of B]!. One excluded element (Stübel 988, B [B 20 0000320]!) represents Cyathea tortuosa R. C. Moran; other syntypes (Stübel 975, B [B 20 0000319 a]!; Stübel 876 a, B [B 20 0000319 b]!) are not assignable to species.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782819BC2AFDECBB8DF289542F.taxon	etymology	Etymology The specific epithet refers to the type locality, the Río Pastaza in Ecuador; today also a province of that country.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782819BC2AFDECBB8DF289542F.taxon	materials_examined	Selected material studied COLOMBIA – Antioquia • Mun. San Carlos, between San Rafael and San Carlos; 5.1544444 ° N, 75.0413889 ° W; 1600 m a. s. l.; 21 Feb. 2015; M. Lehnert 3162; BONN, HUA, Z • Mun. Anori, entre Barbosa y Medellín; 7.252 ° N; 75.052 ° W; Bro. Henri-Stanislas 1708; MO, US. – Caquetá • Municipio de Belén de los Andaquíes, camino Anfaquí, reserva Andaquí, La Profunda; 1 ° 39 ′ 39.5 ″ N, 75 ° 54 ′ 21.9 ″ W; 800 – 1000 m a. s. l.; 13 Mar. 2016; N. Castaño, D. Daly, E. Paki, J. A. Bustos, M. Lasso & A. Valencia 7714; COAH. – Cundinamarca • Yacopí; 5.4588111 ° N, 74.3080083 ° W; 1450 m a. s. l.; M. Morales 474; COL, FMB. – Meta • Municipio de Lejanias, vereda El Triunfo, ubicado a 3 km de las escuela; 3 ° 35 ′ 38.4 ″ N, 74 ° 05 ′ 52.3 ″ W; 1295 m a. s. l.; 21 Jul. 1998; R. López, J. Martínez, N. Cruz, M. A. Pinzón & R. Pinzón 4058; COAH. – Putumayo • Trocha de el Pepino al Rio Guineo, 1.084709 ° N, 76.707126 ° W; 1000 m a. s. l.; W. Hagemann & N. Leist 2051; COL. – Risaralda • Mun. Pereira, Hacienda Alejandría, Km 8 carretera Cerritos – La Virginia, extremos norte de parte ancha del valle del Rio Cauca, en bosque de galería a lo largo de la quebrada; 4.850 ° N, 75.867 ° W; 960 m a. s. l.; 8 Jul. 1995; P. Silverstone-Sopkin 7744; CUCV. – Tolima • Fresno; 1.426 ° N, 78.384 ° W; 1480 m a. s. l.; J. Cuatrecasas 9379; COL, F, US. ECUADOR – Prov. Morona-Santiago • Macas, roadside vegetation near town of Macas; 2 ° 19 ′ S, 78 ° 08 ′ W; 1100 m a. s. l.; 3 Jul. 1993; A. Fay & L. Fay 3957; MO, NY, QCNE • same collection data as for preceding; A. Fay & L. Fay 4066; MO, UC. – Prov. Napo • Cantón Archidona, road Hollín-Loreto, Km 17, near Río Hollín; 0 ° 41 ′ S, 77 ° 41 ′ W; 1100 m a. s. l.; 14 – 22 Feb. 1989; F. Hurtado & J. Shiguango 1624; MO, UC. – Prov. Pastaza • About 0.5 km E of El Puyo; [ca 01 ° 29 ′ 59 ″ S, 77 ° 58 ′ 43 ″ W]; [ca 1000 m a. s. l.]; 5 Oct. 1974; J. Hudson 845; MO, UC • on E side of El Puyo; 7 Oct. 1974; J. Hudson 896; MO, UC. – Prov. Sucumbios • Shishicho ridge, Alto Aguarico drainage, above (S of) Río Cofanes, W of Puerto Libre, NW of Lumbaqui, access from Río Sieguyo, below Shishicho Camp; 0 ° 12 ′ 01.3 ″ N, 77 ° 31 ′ 54.3 ″ W; 1500 – 1570 m a. s. l.; 13 Aug. 2001; R. Aguinda, N. Pitman & R. B. Foster 1302; F, UC. – Prov. Zamora-Chinchipe • Nangaritza canton, Parroquia Guayzimi, Campamento Militar Miazi, al sur del río Nangaritza; 4.27 ° S, 78.70 ° W; 1060 – 1000 m a. s. l.; 21 Oct. 1991; C. E. Cerón, M. Chango, V. Tapur & G. Aymard 16836; MO. PERU – Amazonas • Bagua, Soldado Oliva, carretera Bagua-Imaza; [ca 5.304 ° S, 78.388 ° W]; 660 m a. s. l.; 6 Feb. 1999; C. Díaz et al. 10642; MO.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782819BC2AFDECBB8DF289542F.taxon	description	Description Trunks to 3.5 m tall, straight, 5.0 – 8.0 cm diam. (records of 30 cm diam. likely include layer of adventitious roots), covered with old petiole bases, due to these sparsely to strongly aculeate, in lower parts petioles rotting; apices hidden between petioles; adventitious buds absent. Leaves to 275 cm long, held with petioles erect to ascending, lamina ± planar, weakly arching. Petioles 20 – 45 (– 59) cm long, sparsely aculeate, prickles to 3 mm long, dark yellowish brown to stramineous, sometimes basally darker brown; 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 – 32.0 × 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 the yellowish brown epidermis, persistent but easily abraded. Blades (110 –) 180 – 255 × 90 – 105 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 16 – 18 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 yellowish 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, costae and costules also with dark brown, erect shell-like to bullate squamules, 0.5 – 1.0 × 0.5 mm with subulate, rarely flattened or elongate tips, larger flat squamules to 2 × 1 mm rare or absent; junctions rachises / costae abaxially weakly swollen, each with an inconspicuous planar pneumathode, dark brown, elliptic, to 4.0 × 2.5 mm, area around it often black in dried specimens (foliar nectary). Largest pinnae 37 – 45 cm long, sessile (appearing stalked if proximal basiscopic pinnule fallen off), pinnae patent to weakly ascending, distally narrowly to broadly green alate, the pinnatifid terminal segment shortly decurrent into the costae. Largest pinnules 8.0 – 11.5 × 2.0 – 2.5 cm, linear-oblong to linear-lanceolate, incised to ½ or more of their width (usually 2 – 3 mm between sinuses and costules), sessile, bases on smaller ones truncate to weakly cuneate, in larger ones truncate to weakly cordate, smaller pinnules (especially in lower pinna) with obtuse to rounded tip, larger pinnules tapering from beyond the middle to short-attenuate tips; costules basally with a black ring going all around their bases (abscission layer); largest segments 12.0 – 15.0 × 2.5 – 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 glabrous except for some dark brown bullate squamules and trichomidia on midveins, absent between the veins; sterile and fertile veins mostly simple, regularly forked in the tips of lager segments. Sori 0.8 – 1.0 mm diam., medial to supramedial (sometimes appearing submarginal when sori still complete), parallel to the midvein or in triangular lines, on the back of veins or in fork of veins, indusia absent; receptacles globose to ellipsoid, 0.2 – 0.3 mm diam.; paraphyses few to numerous, hyaline, brown to tan, shorter than sporangia (0.2 – 0.3 mm). Spores not examined.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782819BC2AFDECBB8DF289542F.taxon	distribution	Distribution and ecology Colombia (Cordilleras Occidental and Central), Ecuador and northern Peru (eastern Andean slopes) (Fig. 2 B) at elevations of (310 –) 850 – 1600 m a. s. l., dubious records also from the western escarpment (Ecuador: El Oro; Colombia: Chocó); in the understory of lower montane forest, preferably in small valleys (quebradas) and near water courses.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782819BC2AFDECBB8DF289542F.taxon	discussion	Remarks The syntypes of Alsophila pastazensis have been regarded to represent one species, but we consider them to belong to at least two species, one of them Cyathea tortuosa R. C. Moran. This is supported by recent observations by the first author at the Jardin Botanico las Orquideas, a private reserve near Puyo, Ecuador. Both species grew only a few meters apart in the dense forest understory, but each with a specific habit, with C. pastazensis having a trunk covered in old petiole bases and funnel shaped crown, and C. tortuosa with a clean-shed trunk and patent leaves. Cyathea pastazensis is best described as a Cyathea floribunda with darker, concolorous squamules on the blades because both have sessile pinnae and round-tipped segments. However, C. pastazensis shows the typical leaf architecture of C. pungens with the pinnules of the lower pinnae being blunt-tipped whereas the pinnules of the distal pinnae are decidedly more acute (vs all equally blunt-tipped in C. floribunda, as observed so far). In the contact zone of the ranges of both species in southern Ecuador and northern Peru, it almost looks like a fluid transition between the two morphologies in a large shared range (Fig. 2 B); but as we interpret it, C. pastazensis extends south along the lowland forests of the Amazon basin whereas C. floribunda reaches north along the Andean foothills. Both Cyathea pastazensis and C. floribunda differ from C. pungens in soral lines that follow the midveins, thus forming parallel lines across the whole pinnule; in C. pungens, the soral lines are parallel to the segment margins, thus shaped in a zig-zag pattern across the whole pinnule. In C. pastazensis and C. floribunda, pinnae and pinnules are sessile whereas in C. pungens, at least the largest pinnae are clearly stalked, creating a visible gap between rachis and blade; pinnules tend to be remote from the costae, too, but this character is more clearly developed in eastern Amazonian population of C. pungens. Cyathea werffii grows in the same region (e. g., Baker 5699, MO, NY), but has smaller leaves with stalked lower pinnae (vs all sessile in C. pastazensis), smaller pinnules with blunt, often widened tips, appearing almost rectangular; it also has scarcer scaly indument on the axes than C. pastazensis.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782814BC2CFD93BE28F5A75628.taxon	materials_examined	– Type: BRAZIL • Bahia; Ilheos; ca 14 ° 10 ′ S, 53 ° 05 ′ W; 1817 – 1820; C. F. P. Martius 391; lectotype: B [B 20 0000200]!, chosen by Barrington 1978; isolectotypes: B [B 20 0000198, B 20 0000199, B 20 0000201]!, BR [BR 0000006987541, labelled “ Ilheos, Luschnatt ”] image!, HAL [HAL- 0086574]!, K [K 000589886]!, L [L. 1284712, L. 1284714]!, LE [LE 00008123] image!, M!, MO [MO- 255859]!, NY [00148753, fragment of B, 00148755, 00148756] image!, P [P 00631777, P 00631778, P 00631779]!.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782814BC2CFD93BE28F5A75628.taxon	etymology	Etymology The specific epithet translates as ʻwell girdledʼ, which may either refer to the tight, string-like arrangement of the sori or the many deep sinuses along each side of a pinnule.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782814BC2CFD93BE28F5A75628.taxon	materials_examined	Selected material studied BRAZIL – Alagoas • Poço DʼAnta, ca 16 – 19 km NNW of Muricí by road, Mata de Muricí; 9 ° 14 ′ S, 35 ° 53 ′ W; 550 – 600 m a. s. l.; 14 May 2001; W. W. Thomas 12419; MO, NY. – Bahia • Arataca, Serra do Peito-de-Moça; 15 ° 12 ′ 10 ″ S, 39 ° 24 ′ 29 ″ W; 900 m a. s. l.; 12 Aug. 2009; P. L. R. de Moraes, H. van der Werff, L. Daneu & R. Perdiz 2737; HUEFS, RB • Camacã, RPPN Serra Bonita; 28 Nov. 2014; P. L. R. de Moraes, H. van der Werff & L. Daneu 4254; HRCB • Almadina, Serra do Corcovado; 13 Dec. 2014; P. L. R. de Moraes, H. van der Werff & L. Daneu 4553; HRCB • Belmonte, Fazenda Taquara; 16 Feb. 2015; P. L. R. de Moraes, H. van der Werff & L. Daneu 4810; HRCB • Una Reserva Biologica de Una, near Rio Maruim; 14 Nov. 2000; I. Fernades, S. C. Sant ʼ Ana & M. Caravallo 1498; NY • Barro Preto, ca 13.5 km on street from Barro Preto to the access to Serra da Pedra Lascada; 14 ° 46 ′ 22 ″ S, 39 ° 32 ′ 16 ″ W; 960 m a. s. l.; 13 Aug. 2003; P. Fiaschi, A. M. Amorim, S. C. Sant ʼ Ana & J. L. Paixão 1574; NY.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782814BC2CFD93BE28F5A75628.taxon	description	Description Trunks 0.7 – 1.5 m tall, ca 8 – 10 cm diam., with old petiole bases, straight to decumbent; without adventitious buds. Petioles ca 35 – 50 cm long, inermous to sparsely muricate, brown to dark purpureous, matte; scurf absent. Petiole scales lanceolate, 14.0 – 20.0 × 4.0 – 4.5 mm, their tips straight, concolorous yellowish to white, sometimes with dark central stipe, auburn to dark brown. Leaves to 230 cm long; patent and distally arching. Blades to ca 180 × 90 – 120 cm, bipinnate-pinnatifid, firm herbaceous to chartaceous; dark shiny green adaxially, often blackish when dried, pale olive green abaxially; apices abruptly reduced to a non-conform section. Pinnae to 35 – 65 cm long, number of pairs per leaf ca 8 – 10, stalked 1.0 – 2.5 cm, patent to ascending, basal pinnae ca ⅔ the length of longest pinnae, patent to weakly reflexed; pinnae distally green alate; distal pinnules free, the pinnatifid terminal segments shortly decurrent into the costae. Leaf axes brown to carnose on both sides, the costules abaxially often blackish; with whitish to brown, antrorsely curved hairs to 0.8 mm long adaxially, abaxially glabrescent with scattered, appressed, tan to brown trichomidia; abaxially with auburn to orange-brown squamules with entire margins and white tips, bullate or flat, 0.4 – 1.0 mm long; costae inermous, 1.5 – 3.0 mm wide; insertions of costae into rachises not or only abaxially weakly swollen, without clearly developed pneumathodes, usually blackened in dried specimens. Pinnules to 7.0 – 11.5 × 1.5 – 2.0 cm, short stalked to 3 mm, alternate, 1.5 – 3.0 cm between the stalks; linear-oblong, bases truncate to weakly cordate, long attenuate tips, and with finely serrate margins; incised to ½ or more of their width, with acute sinuses to 2 mm wide, weakly dimorphic, sterile part of pinnae and pinnules wider with narrower, often occluded sinuses; the carnose to brown stalks inarticulate, with a black ring at their bases (abscission layer?), without pneumathodes; segments oblong to deltate, patent to ascending, with subentire to finely crenulate margins, in the latter case weakly revolute; tips straight to weakly falcate, rounded to obtuse; basal segments opposite to alternate, the lowest ones not remote from each other. Veins planar, carnose to dark brown abaxially and adaxially, ending shortly before the segment margins, becoming wider and reddish adaxially; glabrous adaxially, abaxially with appressed, tan trichomidia and thin, hyaline, twisted hairs to 1 mm long, with auburn to orange-brown bullate squamules with entire margins, sometimes larger, flattish squamules to 2 × 1 mm proximally on midveins and on costules; no hairs between veins; sterile simple, fertile veins forked, rarely simple. Sori 0.8 (– 1.0) mm diam., inframarginal to marginal, parallel to the margins, usually in the forks of veins, indusia absent; receptacles globose to ellipsoid, 0.3 – 0.4 mm diam., without subtending scales; paraphyses few, hyaline, tan to reddish brown, shorter than the sporangia (0.2 mm). Spores with smooth exospore, finely pitted (Gastony 1979).	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782814BC2CFD93BE28F5A75628.taxon	distribution	Distribution and ecology Northeastern Brazil at elevations of 550 – 960 m a. s. l., in the understory of Mata Atlântica.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782814BC2CFD93BE28F5A75628.taxon	discussion	Remarks Cyathea praecincta has been often misidentified as C. pungens when petiole scales are missing (mostly concolorous white in C. praecincta vs strongly bicolorous with white margins in C. pungens), but the whole appearance and coloration of its specimens differs strongly from true C. pungens. Dried specimens have the blades bicolorous, with blackish adaxial side and olive to grayish green abaxial side (vs ± the same color in C. pungens, varying from pale green to brown depending on preservation) and the axes are a homogenous reddish brown to atropurpureus (vs yellowish to pale brown in C. pungens). Cyathea praecincta has many fertile veins forked while they are generally simple in C. pungens. According to our data, both species have clearly separated dstributions (Figs 1 – 2). From Alagoas, we have seen only images of specimens without petiole (W. W. Thomas 12419, MO, NY), which may be C. macrocarpa (C. Presl) Domin. The specimens have the lowest segments slightly smaller than the following in the pinnules, and the margins slightly dentate, which matches more C. macrocarpa than C. praecincta. Cyathea macrocarpa is scatteredly distributed from northeastern Brazil to the Guayana shield (Lehnert 2011). Both species occur together, have submarginal sori and very pale petiole scales, and they are often not distinguishable from photos alone (https: // www. inaturalist. org / observations / 103467779). A closer look at the sori allows a clear separation as C. praecincta has no indusium, which is easily discernable due to the sparse paraphyses, whereas C. macrocarpa has a dense mass of long persisting paraphyses that hide a well developed hemitelioid indusium.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782812BC13FD85BC2BF54A53F3.taxon	description	Figs 10, 13 B	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782812BC13FD85BC2BF54A53F3.taxon	etymology	Etymology The specific epithet refers to the sharp stinging prickles that are usually present on the petioles of this species. Selected material studied HAITI – Nord • Plaisance; [ca 19.596973 ° N, 72.466326 ° W]; 609 m a. s. l.; 27 Aug. 1903; G. V. Nash 887; NY, US. DOMINICAN REPUBLIC – Barahona • Paradis vicinity; [ca 18.149 ° N, 71.107 ° W]; 0 – 800 m a. s. l.; 28 Jan. 1922; W. L. Abbott 1580; US. – Pacificador • Vicinity of San Francisco de Macoris, Lo Bracito; [ca 19.326 ° N, 70.259 ° W]; 400 – 1000 m a. s. l.; 5 Apr. 1922; W. L. Abbott 2030; US. – Samaná • Monte Negro, about 10 miles N of Sánchez, Samaná Peninsula; ca 19.260 ° N, 69.591 ° W; 300 – 400 m a. s. l.; 17 Mar. 1969; A. H. Liogier 14453; NY. PUERTO RICO [USA] – Río Grande • Luquillo Forest, Rt. 191, km 5.2; [ca 18.334 ° N, 65.764 ° W]; 286 m a. s. l.; 22 Nov. 1973; D. S. Conant 670; F, MO, US, VT. GUADELOUPE – Basse-Terre • Marecageuse de Lamentin; [ca 16.292 ° N, 61.635 ° W]; 5 – 12 m a. s. l.; Jun. 1897; A. Duss 3882; F, NY, US. COLOMBIA – Amazonas • Leticia, Quebrada Tacana, 22.5 km NNW cabecera municipal; 4.022 ° S, 69.999 ° W; 3 Aug. 2000; J. C. Arias-G. 828; COAH, COL, HUA. – Caquetá • San José del Fragua Inspeción de policia de Puerto Bello, resguardo San Miguel; 1 ° 08 ′ 67 ″ N, 76 ° 16 ′ 14 ″ W; 200 m a. s. l.; 4 Oct. 2015; L. F. Giraldo 3575; COAH, HUA. – Cauca • Piamonte, aguas arriba de la desembocadura del Indayaco en el Caquetá, margen derecha; 1.02 ° N, 76.48 ° W; 400 m a. s. l.; 19 Oct. 1996; R. Sánchez 3232; COAH. – Guaviare • Municipio de San Juán del Guaviare, vereda Los Alpes, finca El Provenir, Serrania La Lindosa, 02 ° 31 ′ 51 ″ N, 72 ° 48 ′ 39 ″ W; 310 m a. s. l.; 22 Jul. 2017; D. Cárdenas, N. Marín, M. Holguín & G. Holguín 48784; COAH. – Meta • Uribe, PNN Tinigua, vereda Aires del Meta; 2.583 ° N, 74.333 ° W; 23 Apr. 2002; M. Gaitán 7; COAH. – Putumayo • Puerto Leguízamo, Río Putumayo en Puerto Ospina; 0.229 ° N, 75.938 ° W; 230 m a. s. l.; J. Cuatrecasas 10578; COL, F, US. VENEZUELA – Amazonas • Rio Yatuá; 1.17508 ° N, 66.0572 ° W; 105 m a. s. l.; K. M. Redden et al. 3539; US. – Bolívar • Dtto. Sifontes, concesión Minera Oro Uno, 7 km al NO de la Clarita; [ca 06 ° 13 ′ N, 61 ° 27 ′ W]; 180 m a. s. l.; 3 Aug. 1985; G. Aymard et al. 3874; MO. – Delta Amacuro • Venezuela-British Guiana frontier, Sierra Imataca, upstream from San Victor, past Quebrada Piedradero and Río Matanaima; [ca 01.665 ° N, 62.420 ° W]; 65 – 80 m a. s. l.; 1 Nov. 1960; J. A. Steyermark 87203; MO, NY, US, VEN. – Zulia • Dtto. Bolivar, Cuenca del Embalse Burro Negro; (Pueblo Biejo), sector entre Quiros-El Pensado y el pie de Cerro Socopo, en el area aprox. 10 km en linea recta al este de Churuguarita; 10.169 ° N, 71.0459 ° W; 250 – 300 m a. s. l.; G. S. Bunting 9515; UC. TRINIDAD AND TOBAGO – Trinidad • Causa road, gully, near 3 mile post; 25 May 1928; W. E. Broadway 6958; F, MO • Without locality; 1878 – 1880; A. Fendler 112; MO, NY, US. GUYANA – Barima-Waini • Barima River, Northwest District; [ca 8.33 ° N, 59.83 ° W]; [ca 110 m a. s. l.]; 19 Mar. 1923; J. S. de la Cruz 3417; MO, NY, US. – Cuyuni-Mazaruni • Arawak Matope, Cuyuni River; [ca 06.51 ° N, 58.91 ° W]; 91 m a. s. l.; 20 Jul. 1933; T. G. Tutin 403; US. – Demerara-Mahaica • Cuyuni-Mazaruni, Paruima to Conoch Tipu, S of village ± 1 km; 5.98333 ° N, 61.05 ° W; 530 – 610 m a. s. l.; 17 May 1990; T. McDowell & D. Gopaul 2624 a; US. – Upper Takutu-Upper Essequibo • Upper Essequibo Region, Rewa River, near Camp 2 at foot of Spider Mountains; 3 ° 07 ′ 59 ″ N, 58 ° 31 ′ 59 ″ W; 220 m a. s. l.; 13 Sep. 1999; M. J. Jansen-Jacobs 5881; L [formerly U], NY, UC, US. SURINAME – Commewijne • Perica R., Capoerica Ridge, near km 3; ca 05.758 ° N, 54.973 ° W; 10 m a. s. l.; 27 Jan. 1954; J. C. Lindeman 5405; US. – Para • Upper Para Creek to Suriname River, ca 8 km E of Zanderij on road to Phedra; ca 5.488 ° N, 55.156 ° W; 10 m a. s. l.; 12 Feb. 1961; R. M. Tryon Jr. & K. U. Kramer 5618; US. – Sipaliwini • Inselberg Talouakem – Massif des Tumuc-Humac; 2 ° 28 ′ 59 ″ N, 54 ° 45 ′ 00 ″ W; 250 m a. s. l.; 8 Aug. 1993; J. J. de Granville et al. 12126; L [formerly U], US. FRENCH GUIANA – Cayenne • D. Z. de Crique Jupiter, Bassin du Sinnamary; 4.0675 ° N, 53.1619 ° W; 80 m a. s. l.; 27 Apr. 1991; J. J. de Granville, C. S. Roesel & L. Brothers 11592; US. – Saint-Laurent-du-Maroni • W face of Roche Koutou ʻinselbergʼ, Upper Maroni Basin; 3.1444 ° N, 54.05 ° W; 170 m a. s. l.; 19 Aug. 1987; J. - J. de Granville, L. Allorge, W. J. Hahn & M. Hoff 9464; F. ECUADOR – Napo • Río Lagarto Cocha, near Redondo Cocha; 0 ° 33 ′ S, 75 ° 53 ′ W; 190 m a. s. l.; 15 Jun. 1983; J. E. Lawesson, T. LaessØe & P. M. JØrgensen 44394; AAU, MO, QCA. – Orellana • Parque Nacional Yasuni, Km 92.5 rd Pompeya-Iro; 0 ° 53.370 ′ S, 76 ° 13.520 ′ W; 250 – 300 m a. s. l.; 12 May 1997; M. J. Macía & A. P. Yánez 323; MO, QCA. – Pastaza • Oil exploration camp Chichirota, on the Río Bobonaza; 2 ° 22 ′ S, 76 ° 40 ′ W; 300 m a. s. l.; 26 Jul. 1980; B. Øllgaard, E. Asanza-C., J. Brandbyge, S. Roth & C. Sperling 35264; AAU, F, QCA, UC, US. – Sucumbios • Alto Río Aguarico, Río Chingual, ridge just before Ching Chingual bridge, between Rio Recodo and Rio Chingual, new trail toward Bermejo, from new road to Tulcán above Puerto Libre; 0 ° 15 ′ 22 ″ N, 77 ° 28 ′ 25 ″ W; 700 – 800 m a. s. l.; 7 Jul. 2000; R. Aguinda, M. Metz & T. Theim 805; F, UC. – Zamora-Chinchipe • Cordillera del Condor, Miazi, flood plain forest along Rio Nangaritza. Transect # 3; 4 ° 18 ′ S, 78 ° 40 ′ W; 850 m a. s. l.; 28 Jul. 1993; A. Gentry 80578; MO. PERU – Amazonas, Prov. Condorcanqui • Dist. Imaza, Comunidad Nativa de Yamayakat; 5.057 ° S, 78.338 ° W; 350 m a. s. l.; Mar. 2002; R. Bonino 345; MO. – Huanuco, Prov. Leoncio Prado • Dtto. José Crespo y Castillo, carretera a Cotomono, cerca a Aucayacu; ca 08.950 ° S, 76.1176 ° W; 500 – 580 m a. s. l.; J. Schunke V. 10507; MO, UC. – Loreto, Prov. Iqiuitos • San Juan, río Tigre; 2 ° 35 ′ S, 75 ° 40 ′ W; 245 m a. s. l.; 17 Mar. 1987; W. H. Lewis, M. Elvin-Lewis, D. Fast & J. Campos de la Cruz 12863; MO. – Madre de Dios, Prov. Tambopata • 30 air km or 70 – 80 river km SSW of Puerto Maldonado at effluence to La Torre (Río DʼOrbigny) / Río Tambopata (SE bank), Tambopata Nature Reserve; 12 ° 49 ′ S, 69 ° 17 ′ W; 260 m a. s. l.; 16 Apr. 1980; P. J. Barbour 4795; MO, UC. – Pasco, Prov. Oxapampa • Dist. Palcazú, San Pedro de Pichanaz-Azulis, Reserva Comunal Yanesha; 10 ° 28 ′ 40 ″ S, 75 ° 06 ′ 10 ″ W; 670 m a. s. l.; 25 Feb. 2004; L. F. Mellado N. 0733; MO, USM. – Puno, Prov. Carabaya • Río Tavara base camp; 13 ° 21 ′ S, 69 ° 40 ′ W; 400 m a. s. l.; 19 May 1992; A. Gentry, C. Reynel, R. Ortiz & P. Nuñez 76848; MO, NY, UC. – San Martin, Prov. Mariscal Caceres • Distrito Tocache Nuevo, Puerto Pizana (right margin of Río Huallaga); 350 m a. s. l.; 1 Jun. 1974; V. J. Schunke 6866; UC. • Prov. San Martin • In montibus secus flumen Mayo, prope Tarapoto; Jul. – Aug. 1856; R. Spruce 4715 [excluded syntypes of Alsophila floribunda, mix with C. dombeyi]; BM [BM 000777031, BM 000777032 (mix with C. dombeyi), BM 000777346], P [P 00642346 p. p., P 00642349, P 00642350] • In monte Campana prope Tarapoto; Aug. 1856; R. Spruce 4715; W [W-Rchb. 0053380]. – Ucayali • Col. Prado, along new road from Parque Nacional Alexander von Humboldt to Puerto Inca, ca 5 km E of main Pucallpa-Tingo Maria road at Km 86; 8 ° 17 ′ S, 74 ° 58 ′ W; 500 m a. s. l.; 5 Nov. 1980; T. B. Croat 51061; MO, UC. • Coronel Portillo, Calleria, camino a la altura del Quebrada Mojaral, este del Río Utiquina y Quebrada Pumayacu; 8 ° 09.13 ′ S; 74 ° 15.48 ′ W; 150 – 175 m a. s. l.; J. Schunke Vigo & J. G. Graham 15500; F, UC. BRAZIL – Acre • Município de Rio Branco, BR 364, estrada Rio Branco - Porto Velho, km 11 da magem ramal que da acesso ao lugar Belo Jardim; 25 Oct. 1980; C. A. Cid Ferreira & B. W. Nelson 3053; NY, US. – Amazonas • Coari, Província Petrolífera de Urucu, arredores do pólo Arara, igarapé da Onça; 4.864454 ° S, 65.298671 ° W; 75 m a. s. l.; 2007; M. R. Pietrobom-Silva 6928; RB. – Ceará • Serra do Araripe; [ca 07.389 ° S, 40.216 ° W]; v. Luetzelburg 25816 a; UC. – Goias • Jatai; [ca 17 ° 53 ′ 26.8 ″ S, 51 ° 45 ′ 01.4 ″ W]; [ca 670 m a. s. l.]; A. Macedo 1518; US. – Mato Grosso • Municipio de Porto dos Gauchos, bacia rio Patelão (afl. rio dos Peixes), ca 115 km W de Porto Atlantico (Rio Teles Pires); 11.5 ° S; 56.75 ° W; 350 – 400 m a. s. l.; 16 Jul. 1991; P. G. Windisch & W. Oliveira 6493; US. – Mato Grosso do Sul • Munic. Rio Verde de Mato Grosso, Serra Pimenteira, Cachoeira Anhumas, Fazenda Quartel; 18.9166667 ° S, 54.8833333 ° W; 350 m a. s. l.; 23 Feb. 1994; M. R. Silva & C. E. Rodrigues 1298; MO. – Pará • Serra dos Carajás, “ Azul ” near camp at Serra Norte; 5.983 ° S, 50.467 ° W; 646 m a. s. l.; 8 Dec. 1981; D. C. Daly 1836; MO • Alenquer, Floresta Estadual do Paru, Trilha T 3; [ca 00.947 ° S, 53.243 ° W]; 53 m a. s. l.; 11 Dec. 2008; S. Maciel 1541; BONN, MCT • “ Tanaii, ad Rio Acara, juxta Para ”; [ca 01.563 ° S, 1.563 ° W]; [ca 15 m a. s. l.]; Sep. 1849; R. Spruce 22; K, TCD. – Rondônia • Rio dos Pacaás Novos, above the first cachoeira; between the river and the base of the Serra dos Pacaás Novos; 10 ° 58 ′ 46 ″ S, 63 ° 46 ′ 10 ″ W; 220 m a. s. l.; 22 Mar. 1978; W. R. Anderson 12236; F, NY. BOLIVIA – Beni, Prov. Vaca Diez • 3 km E of Riberalta on road to Guayamerín, then 2 km SE on side road; 11 ° 00 ′ S, 66 ° 05 ′ W; 230 m a. s. l.; 19 May 1982; J. C. Solomon 7700; MO UC. – Cochabamba, Prov. Carrasco • Projecto Valle del Sacta, 241 km W of Santa Cruz, 219 km E of Cochabamba off new road between Cochabamba and Santa Cruz; 17 ° 12 ′ S, 64 ° 43 ′ W; 290 m a. s. l.; 13 Jul. 1989; A. Fay & L. Fay 2357; LPB, MO. – La Paz, Prov. Abel Iturralde • Parque Nacional Madidi, Pampas del Heath, along Río Heath, camp Maronal; 12 ° 51 ′ S, 68 ° 48 ′ W; 204 m a. s. l.; 9 Nov. 2004; J. Gonzales, H. Nenzen, J. Choque & J. C. Cespedes 4711; LPB, UC. – Pando, Prov. Manuripi • Along Rio Madre de Dios, 3 km W of Humaita; 12 ° 01 ′ S, 68 ° 18 ′ W; 150 m a. s. l.; 30 Aug. 1985; M. Nee 31660; LPB, NY, MO. – Santa Cruz, Prov. Velasco • Huanchaca I, d 1, Huanchaca; 13 ° 54 ′ S, 60 ° 48 ′ W; 650 m a. s. l.; L. Arroyo P., T. J. Killeen, J. Wellens, H. Gonzáles & P. Solíz 669; MO, UC, UCSZ.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782812BC13FD85BC2BF54A53F3.taxon	description	Description Trunks 0.5 – 4.0 m tall, 4.0 – 8.0 (– 15.0) cm diam., straight to decumbent, with persistent old petiole bases, spiny; apices hidden between petioles; adventitious buds absent. Leaves to 250 (– 300) cm long; smaller ones erect, in funnel-shaped crown, larger ones strongly arching. Petioles to 65 cm long, aculeate, prickles 3 – 5 mm long, dark yellowish brown to stramineous, sometimes basally darker brown; aerophores to 10 × 1 mm, inconspicuously brown in dried material, whitish in fresh material; without remote (aphlebioid) pinnae at the petiole bases; petiole scales lanceolate, 15.0 – 25.0 × 2.5 – 4.0 (– 6.0) mm, their tips straight to falcate, concordantly to discordantly bicolorous, shiny dark brown, with pale brown to whitish margins, scales in distal half of the petiole almost concolorous white except for brown spots near the base; colors only sharply contrasted in scales near petiole base and on crosiers; 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 prickles. Blades 100 – 180 × 60 – 100 cm, bipinnate-pinnatifid, elliptic to obovate, chartaceous; dark olive-green adaxially, often blackish when dried, olive-green abaxially; apices abruptly reduced to non-conform apical sections, sometimes very broad and almost gradually reduced. Rachises stramineous to yellowish brown on both sides, in basal half often with some strong prickles; adaxially with antrorsely curved uniseriate hairs 0.5 – 1.0 mm long, abaxially glabrous except for scurf remnants, containing appressed, white trichomoidia and dissected squamules (0.2 – 0.5 mm). Largest pinnae 35 – 60 cm long, pinnae 8 – 12 pairs per leaf, mostly stalked to 1.5 – 2.5 cm, ascending, distally narrowly to broadly green alate, the distal segments free, the pinnatifid terminal segments shortly decurrent into the costae. Costae inermous, to 1.5 – 2.0 mm wide, adaxially with antrorsely curved uniseriate hairs 0.5 – 1.0 mm long, abaxially glabrous or glabrescent with scurf like on the rachises, insertions into rachises abaxially weakly swollen, each with an inconspicuous planar pneumathodes, dark brown, elliptic, to 2.0 × 1.0 mm, area around it often black in dried specimens. Pinnules (3.5 –) 5.0 – 8.0 (– 10.0) × (1.0 –) 1.5 – 1.8 (– 2.0) cm, smaller ones sessile, larger ones stalked 1.0 – 1.5 (– 3.0) mm, articulate, alternate, 1.5 – 2.0 cm between the stalks, linear-oblong, incised to ½ or more of their width, bases truncate to weakly cuneate, tapering from beyond the middle to short acute to short attenuate tips; costules with ephemeral tan to brown trichomidia and tan to brown squamules 0.5 – 1.0 mm long with entire margins and short acute tips, varying in color from white over tan, concolorous or with paler tips; costules basally with a black ring going all around their bases (abscission layer); segments oblong, to 5.0 – 12.0 × 3.0 – 5.0 mm, patent to ascending, distally weakly falcate, with entire to subentire margins, tips rounded to obtuse, rarely acute and then margins weakly dentate; basal segments opposite to alternate, the lowest ones not remote from each other, sinuses triangular, acute, 1.0 – 2.0 mm wide, sometimes closed; veins prominent abaxially and adaxially, midveins strongly so and adaxially ridged, lateral veins ending at segment margins; midveins yellowish brown abaxially and adaxially, lateral veins yellowish to greenish brown or blackish; adaxially glabrous except for occasional single hairs on the midveins, abaxially glabrous or with squamules and trichomidia like on the costules, trichomidia sometimes also between the veins; sterile and fertile veins mostly simple, rarely forked in larger pinnules (> 2 cm wide). Sori 0.6 – 0.8 (– 1.0) mm diam., supramedial to inframarginal, 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, white, shorter than sporangia (0.2 – 0.3 mm). Spores not examined.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782812BC13FD85BC2BF54A53F3.taxon	distribution	Distribution and ecology Haiti, Dominican Republic, Puerto Rico, Guadeloupe, Trinidad & Tobago, Amazonian lowland of Colombia, Venezuela, Suriname, French Guiana, Ecuador, Peru, Brazil, and Bolivia at elevations of ca 5 – 1100 m a. s. l., in shade along creeks or in sun on swampy soils, flourishing in tree fall gaps and small clearings; in the Caribbean mostly near mangroves and in limestone areas, often in shaded sinkholes.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782812BC13FD85BC2BF54A53F3.taxon	discussion	Remarks The exact locations where the types of Polypodium pungens and P. procerum, both attributed to J. C. Hoffmannsegg, were collected are unknown. Hoffmannsegg did not travel to Brazil personally but had specimens collected and sent from there by paid collectors (Stresemann 1950). Although an accomplished botanist, Hoffmannsegg seemed to have had his main focus on animals during that time, and documentation of the Brazilian plant material in the literature is scarce. However, most of the plants seem to have been collected by Friedrich Wilhelm Sieber, who was a handyman of Hoffmannsegg, and his itinerary has been researched for the bird specimens he collected (Stresemann 1950). Sieber was almost exclusively active in todayʼs state of Pará, along Rio Amazonas between Obidos and Gurupá, and near “ Rio Preto ”, which refers most likely to the town of São Benito do Rio Preto in neighboring state Maranhão. One sheet of the type of P. pungens bears a slip with the fragmentary information that it is “ fern herb growing low, but if in dry forest at river (Sieber) ”, and the type of P. procerum “ fern herb, is a real tree, in shape of a ‘ palmeiroʼ, in forest at river, its height was more than 6 ells (Sieber) ” [personal translation from German], which are vague but still fitting descriptions of the aspect of Cyathea pungens. We may conclude that the types of P. procerum and P. pungens had been collected by Sieber, either in the state of Pará or Maranhão. One of the slips on the type of P. pungens (B 20 0000318) bears a little scribble below the location “ Brazil ” that may be deciphered as “ Coto ”; there is a location of that name in Maranhão east of São Benito do Rio Preto, which is one of the documented areas that Sieber visited. However, this particular specimen was stored in the Herbarium Link, does not bear any reference to Sieber or Hoffmannsegg, and was only aligned with the Willdenow specimen (B-W 19716) by Hieronymus. The publication of the species names in 1810 is presumably a result from Sieber′s last shipment from Brazil, which arrived in 1809 (Stresemann 1950). This seems rather incredibly fast for the 19 th century, but Hoffmannsegg had a network of keen and active collaborators in Europe who delivered determinations and accounts quickly after receiving specimens (Stresemann 1950). The other providers of Brazilian material sent specimens between 1801 and 1807. If the type specimen of P. pungens had been among them, it likely would have been published sooner. Also, these earlier shipments came from Bahia and Rio de Janeiro, from where C. pungens has only been erroneously reported, based on confusion with similar exindusiate species like C. praecincta (Kunze) Domin or C. atrovirens (Langsd. & Fisch.) Domin. We conclude that the location of the type of Cyathea pungens can be put rather confidently in the lower Amazonas region. Cyathea pungens is widely distributed in the Amazonian lowland (Zuquim et al. 2008), and even with the narrower definition, C. pungens remains quite variable. Most specimens fall into a form with short stalked, asymmetrically based pinnules (including the type of the species) and a form with sessile, ± symmetrical based pinnules (this can be tentatively aligned with the type of the synonym Alsophila infesta). The former seems to be more frequently encountered in eastern Amazonia-Guayana whereas the latter seems to be more dominant in western Amazonia and the Andean foothills; the map (Fig. 1), however, shows no clear regional clustering. Furthermore, small prematurely fertile plants that differ from regular mature plants in having no trunk and only pinnate-pinnatifid to partially bipinnate leaves can be found along the eastern distributional margin, where the tropical lowland rainforests border savannahs and cerrado vegetation, and in the Guayana region (Berry et al. 1995). A fourth morphotype seems to be restricted to open swamps; it is characterized by having small blades, short elongate pinnules with truncate bases and obtuse to short acute tips, and the scant squamules on the costules abaxially are mostly dark brown, all interpreted as adaptations to increased sun exposure; it still has stalked pinnae and often still visibly, albeit short-stalked pinnules. We dubbed these forms as “ regular ”, ” sessile ”, “ precocious ”, and “ swamp ”, respectively, but give them no taxonomic rank pending further study, preferably with molecular data included. In the Caribbean region, including the Lesser Antilles, Trinidad & Tobago, Puerto Rico and Hispaniola, but not on Cuba or Jamaica, there is a variant of C. pungens with both pinnae and pinnules sessile (vs at least lower pinnae stalked in continental South America) but with the typical scant hair and pale squamules on the blades. At first glance, this form is in stark contrast with the Amazonian population and may merit formal recognition below species rank; in lack of an already published name, we refer to it simply as “ Caribbean ” C. pungens until further studies can clarify its exact status. It is not only found at lower elevations in swamps, which is a typical biotope for the species, but also in sinkholes in limestone areas. We keep C. ruttenbergiae from Puerto Rico (Tejedor & Areces-Berazain 2018, 2021) as separate from Caribbean C. pungens for now, although some of its distinguishing features (i. e., stalked pinnae and pinnules) also occur in continental C. pungens. Its bicolorous petiole scales and the pale laminar indument clearly align it with this species, but the branched fertile veins and the general pinnule shape (especially the truncate sinuses between the segments) of C. ruttenbergiae are more reminiscent of C. muricata Willd. from the Lesser Antilles. In the Caribbean, Cyathea pungens is less frequent than previous accounts suggested because of widespread confusion with Cyathea aspera (L.) Sw. and its segregates. The latter group is often identical in texture and dissection of the blade to the C. pungens group, but generally has concolorous plain brown to castaneous scales on petioles and blade (C. pungens has always some elements with whitish color, e. g., petiole scale margins and laminar squamules). The central Brazilian specimen Windisch & Oliveira 6493 (US) has exceptionally wide pinnules to 25 mm across with forked fertile veins and medial sori. Regarding the indument and the stalked pinnae, it can be confidently put under C. pungens proper and none of the segregates here proposed, except maybe for Cyathea ruttenbergiae (Tejedor & Areces-Berazain 2018). Some collections from French Guiana (i. e., G. Cremers 6700, G. Cremers 7386 are plants of the regular type but have many short, stout hairs on costae and costules abaxially. Another collection from the same area (J. - J. de Granville et al. 9526) is slightly less hairy than these but still hairier than regular C. pungens, and also having dark, white tipped bullate squamules. Petiole scales are not preserved in this specimen, but the general pinnule shape strongly speaks for C. pungens. These aberrants may either constitute their own variant or their own unrecognized hybrid, though we do not have a suggestion for the potential other parent; an involvement of Cyathea oblonga would explain the squamule color but not the hairiness. The specimen Schunke Vigo & Graham 15500 from the Peruvian lowland is a very atypical plant of the “ swamp ” Cyathea pungens. Due to the small segments, the soral lines appear not to follow the margins in a zig-zag-pattern, as they should, but to be parallel to the midveins. Because of this, a smaller leaf at UC was determined as C. werffii by the first author, but the duplicate at F has a trunk and clearly surpasses the size of that species. Cyathea werffii from the Andean foothills of Ecuador and northern Peru is similar to precociously fertile C. pungens. Both species have been found growing closely together in northern Peru (Chiriaco, Dist. Baeza, Prov. Amazonas; Figs 1, 2 A).	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178282DBC16FDE8BA7CF563519B.taxon	description	Figs 11, 12 A	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178282DBC16FDE8BA7CF563519B.taxon	materials_examined	– Type: PERU • Huánuco, Pampayacu; Jul. 1829; E. F. Poeppig s. n.; holotype: LZ [destroyed]; lectotype: B [B 20 0000124, labelled “ Poeppig 201 ”]!, designated by Lehnert 2016.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178282DBC16FDE8BA7CF563519B.taxon	materials_examined	– Type: PERU • San Martin, Tarapoto, “ by rocky streams ”; 1855 – 56; R. Spruce 4717; lectotype: K [K 000227601]!, first step designated by Lehnert 2016, second step designated here; isolecotypes: B [B 20 0000362, fragment]!, NY [00148759, fragment of B] image!, US [00066280] image!.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178282DBC16FDE8BA7CF563519B.taxon	etymology	Etymology The specific epithet refers to the small size of the sori.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178282DBC16FDE8BA7CF563519B.taxon	materials_examined	Selected material studied PERU – San Martín • San Martin, path up to the telephone antenna above Tarapoto, (Alto Ahuashiyacu), along Tarapoto; 6.45 ° S, 76.30 ° W; 1333 m a. s. l.; 10 Aug. 2002; M. J. M. Christenhusz 2081; GOET, TUR • “ In m. Pingullu-Urku ” [Cerro Pingulla?]; [ca 06.356 ° S, 76.689 ° W]; [ca 800 – 1500 m a. s. l.]; Aug. 1856; R. Spruce 4715 forma pygmea [excluded syntypes of Alsophila floribunda]; P [P 00642347, P 00642348]. – Cuzco • Quispicanchi, Marcapata-Quinze Mil road, quebrada of tributary of Río Azara; 13 ° 27.2 ′ S, 70 ° 54.5 ′ W; 1570 m a. s. l.; 23 Oct. 2002; M. Lehnert 458; GOET, NY, UC, USM. – Madre de Dios • Prov. Manu, Parque Nacional Manu Rio Manu, Rio Sotileja; 11.67 ° S, 71.92 ° W; 400 – 500 m, 1 Oct. 1986; R. B. Foster & B. dʼAchille 11548; US. BOLIVIA – La Paz • Nor Yungas, PN Cotapata, a 600 m de la Estación Tunquini; 16.191 ° S, 67.869 ° W; 1690 m a. s. l.; 22 Aug. 1998; A. Portugal, I. Jiménez & C. Rojas 221; GOET, LPB, UC.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178282DBC16FDE8BA7CF563519B.taxon	description	Description Trunks absent or to 1.5 (– 3.0) m tall, slender, 6 – 7 (– 10) cm diam., straight to decumbent, with persistent old petiole bases, inermous to muricate; apices hidden between petioles; adventitious buds absent. Leaves to 150 cm long; weakly arching to strongly arching. Petioles to 55 cm long, inermous to sparsely muricate, prickles 1 – 2 mm long, dark yellowish brown to stramineous, sometimes basally darker brown; aerophores to 10 × 1 mm, inconspicuously brown in dried material, whitish in fresh material; without adventitious (aphlebioid) pinnae at the petiole bases; petiole scales lanceolate, to 16 × 1.5 – 2.5 (– 3.0) mm, their tips straight, weakly twisted, concordantly to discordantly bicolorous, shiny medium to dark brown with whitish margins, scales in distal half of the petiole almost concolorous white except for brown spots near the base; colors only sharply contrasted in scales near petiole base and on crosiers; petiole scurf sparse tomentum of small branched hairs and dissected squamules 0.2 – 0.4 mm long, yellowish white with brown parts, grayish white in general aspect, easily abraded. Blades 80 – 95 × 45 cm, mostly bipinnate, chartaceous; shiny dark green adaxially, often blackish when dried, olive-green abaxially; apices abruptly reduced to non-conform apical sections, sometimes very broad and almost gradually reduced. Rachises stramineous to yellowish brown on both sides, in basal half often with some scattered prickles less than 2 mm long; adaxially with antrorsely curved uniseriate hairs 0.5 – 1.0 mm long, abaxially glabrous except for scurf remnants, containing a mix of appressed, white trichomoidia, dissected squamules (0.2 – 0.5 mm) and larger filiform scales to 20 mm long. Largest pinnae 23 – 39 cm long, pinnae 5 – 7 pairs per leaf, sessile or stalked to 1.5 – 2.5 cm, ascending, distally narrowly to broadly green alate, the distal segments sessile, adnate, the pinnatifid terminal section long decurrent into the costae. Costae inermous, to 1.5 – 2.0 mm wide, adaxially with antrorsely curved uniseriate hairs 0.5 – 1.0 mm long, abaxially glabrescent with scurf like on the rachises, insertions into rachises retaining larger, filiform scales from crozier, insertions abaxially weakly swollen, each with an inconspicuous planar pneumathodes, dark brown, elliptic, to 2.0 × 1.0 mm, area around it often black in dried specimens. Largest pinnules (3.5 –) 5.2 – 7.8 × (1.0 –) 1.2 cm, sessile, articulate, alternate, 1.5 cm between the stalks, oblong-lanceolate to triangular-ovate, subentire to strongly crenate, incised to ⅕ of their width, bases truncate to weakly cuneate, tapering from beyond the middle to short acute to short attenuate tips; basal lobes pronounced to auricles; costules adaxially glabrous or with few hairs like on the costae, abaxially with ephemeral tan to light brown trichomidia and mostly concolorous, tan to yellowish brown, flat and bullate squamules 0.5 – 1.0 mm long, with short subulate tips; costules basally with a black ring going all around their bases (abscission layer); segments not differentiated, outlined only by pinnate venation and short marginal lobes; veins weakly prominent abaxially and adaxially, lateral veins ending at lobe margins; midveins yellowish brown abaxially and adaxially, lateral veins yellowish to greenish brown or blackish; adaxially glabrous except for occasional single hairs on the midveins, abaxially without hairs, with squamules and trichomidia like on the costules; sterile and fertile veins simple. Sori 0.6 – 0.8 (– 1.0) mm diam., medial to subcostal, parallel to the miveins or in tight triangular lines, indusia absent; receptacles globose to ellipsoid, 0.2 – 0.3 mm diam.; paraphyses few, hyaline, white, shorter than sporangia (0.2 – 0.3 mm). Spores not examined.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178282DBC16FDE8BA7CF563519B.taxon	distribution	Distribution and ecology Peru and western Bolivia at elevations of (400 –) 500 – 1690 m a. s. l., mostly in montane forests, observed along rocky creeks in shaded gullies (quebradas).	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C9178282DBC16FDE8BA7CF563519B.taxon	discussion	Remarks A relatively complete leaf of this species was included in the type material of A. floribunda (Spruce 4715 “ forma pygmea ”; isotypes P 00642347, P 00642348). The type of Alsophila pterorachis (Spruce 4717) has serrate pinnule margins, which is atypical for C. pycnocarpa. It is placed here because of the matching overall appearance, pinnule shape and petiole scale morphology. Nevertheless, it may just represent a precociously fertile plant of a different, larger bipinnate-pinnatifid species. With its overhanging leaves and subentire pinnules, Cyathea pycnocarpa is a relatively distinct species in the Andes, but not well understood until now. It had been subsumed under either Cyathea pungens (because of the small sori and the general resemblance) or C. dombeyi (because of the presence of small brown teeth in the pale scale margins). The material was long interpreted as precociously fertile plants of C. pungens s. lat. As fragmentary specimen, Cyathea pycnocarpa bears a strong resemblance to the Brazilian C. dichromatolepis, where the pinnae are similarly coarsely dissected with entire pinnules and a large pinnatifid apical section. This Brazilian endemic has larger, wider petiole scales with broader white margins. It also reaches larger sizes than C. pycnocarpa, which can already be deduced from the thickness of the petioles and leaf axes. In the field, both species are easily distinguished because C. pycnocarpa has only 5 – 7 pinna pairs and blades with truncate bases whereas C. dichromatolepis can have twice the number of pinnae and blades with tapering bases. Foster & dʼAchille 11548 (F, US) from Madre de Dios, Peru, is a very large specimen with most of the laminar indument relatively dark. Because of the dark squamules, it could be mistaken for C. tortuosa, but that species has white tortuous hairs on the blade abaxially and a dense villous hairs cover on the axes abaxially, which are both absent in this specimen.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782828BC17FDD8BB9AF3545000.taxon	materials_examined	– Type: PUERTO RICO • Río Grande, El Yunque National Forest, route 191, km 4.8; 18.338 ° N, 65.764 ° W; 210 m a. s. l.; 30 Jul. 2017; F. Areces 1160; holotype: UPRRP!, isotype: NY!.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782828BC17FDD8BB9AF3545000.taxon	etymology	Etymology The epithet honors Kathy Ruttenberg, US American artist and sculptor.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782828BC17FDD8BB9AF3545000.taxon	materials_examined	Selected material studied PUERTO RICO [USA] • Luquillo Mts., Rt 988 (Sabana Rd.); D. S. Conant 4171 with J. DeCamp; VT.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782828BC17FDD8BB9AF3545000.taxon	description	Description (diagnostic characters) Trunk to 2 m tall, 5 – 7 cm diam.; leaves to 190 cm long, ascending-arching, petioles to 60 cm long, petiole scales ovate-lanceolate, 10.0 – 15.0 × 4.0 – 5.0 mm, shiny, concordantly bicolorous brown with lighter translucent yellowish to cream margins, fragile, often abraded; blades to 90 × 70 cm; 6 – 7 (– 8) pinna pairs, basal ones strongly reflexed ca ½ the length of the longest pinnae; apices abruptly reduced. Pinnae to 45 cm long, ascending, stalked to 2.5 cm, alternate; pinnules to 12 × 2.5 cm, stalked to 4 mm, inarticulate, 2 – 3 cm between the stalks, elliptic-lanceolate, bases asymmetric, with a gap or missing segment on one side, tips attenuate with crenulate margins; costules green on both sides, abaxially strongly prominent, ridged, with sparse tortuous white hairs to 1 mm and flat, lanceolate whitish to beige squamules to 1.5 mm. Segments to 15 × 6 mm, basal segments strongly asymmetric, if slightly remote then connected by wide green wing; sinuses oblong to acute to 3.0 mm wide, never occluded; margins crenulate; midveins with few, tortuous, white hairs to 1 mm; veins simple or forked; sori submarginal. (Full description in Tejedor & Areces-Berazain 2018.)	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782828BC17FDD8BB9AF3545000.taxon	distribution	Distribution and ecology Endemic to Puerto Rico, in the understory of perhumid rainforests.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782828BC17FDD8BB9AF3545000.taxon	discussion	Remarks Almost identical to the regular form of Cyathea pungens proper, but seems to have fewer pinna pairs per leaf, with basal pinnae not as much reduced as in C. pungens and larger pinnules (to 12 × 2.5 cm in C. ruttenbergiae vs mostly 5 – 8 × 1.5 – 1.8 cm in C. pungens); due to the larger pinnules, also fertile veins are often forked (vs generally simple in C. pungens). Most notable are the blunt sinuses between the lowest segments of the largest pinnules; the very few samples of C. pungens that have pinnules wider than 2 cm still have acute sinuses between the lowest segments and not the blunt ones of C. ruttenbergiae.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782829BC19FDEABA02F5C550B2.taxon	description	Fig. 12 B	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782829BC19FDEABA02F5C550B2.taxon	etymology	Etymology The specific epithet refers to the scaly indument found on the leaves. Selected material studied COLOMBIA – Tolima • Mariquita; 1200 m a. s. l.; Jan. 1848; J. L. Linden 1021; W. – Cundinamarca • La Palma, Murca valley, Cordillera de Helicon, 10 km SE of Gachalá; 2300 m a. s. l.; 30 Sep. 1944; M. L. Grant 10322; US. – Norte de Santander • Cordillera Oriental, region del Sarare, entre el Alto del Loro y el Alto de Santa Ines; 1800 – 2200 m a. s. l.; 18 – 21 Oct. 1941; J. Cuatrecasas, R. E. Schultes & E. Smith 12513; F, GH. – Locality unknown • H. Karsten s. n.; PRC. VENEZUELA – Aragua • La Victoria-Colonia Tovar rd., 11.5 km N of Pie de Cerro; 10 ° 22 ′ N, 67 ° 20 ′ W; 1950 m a. s. l.; 14 Jan. 1982; J. Luteyn, S. Mori, N. Holmgren & J. A. Steyermark 8247; NY. – Distrito Federal • Cerro El Avila, S slopes; 10 º 32.6 ′ N, 66 º 52.5 ′ W; 2000 – 2050 m a. s. l.; 11 Nov. 1991; W. Meier 902; UC. – Mérida • Near summit of Cerro San Isidro, above La Carbonera; 2430 – 2745 m a. s. l.; 22 Apr. 1944; J. A. Steyermark 56029; GH, US. – Miranda • headwaters of the Quebrada Chacaito; 10 º 33 ′ N, 66 º 52 ′ W; 1990 m a. s. l.; 28 Feb. 1992; W. Meier 1763; UC. – Vargas • Monumento Natural Pico Codazzi, road Colonia Tovar-La Victoria, sector Matalpo, road Los Colonos-Pto. Cruz, 2.5 km behind Arco, SE of tourist center Villa Bahareque border with Edo. Aragua; 10 º 26 ′ N, 67 º 13.5 ′ W; 1850 – 1950 m a. s. l.; 5 Aug. 1999; J. Mostacero & R. Castillo 261; UC. – Yaracuy • Nirgua, Cerro Azul (Tucuragua), near the limit with Edo. Cojedes, mainly of peak S of Hacienda Venezolano; 9 ° 59 ′ N, 68 ° 37 ′ W; 1400 – 1690 m a. s. l.; 24 Jan. 1999; W. Meier & O. Kunert 4591; UC. TRINIDAD & TOBAGO – Trinidad • Port of Spain; [ca 10 ° 40 ′ 32 ″ N, 61 ° 29 ′ 27 ″ W]; Apr. 1874; C. E. O. Kuntze 1684; NY.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782829BC19FDEABA02F5C550B2.taxon	description	Description Trunks 0.1 – 3.0 m tall, 5 – 7 cm diam., presumably covered with old petiole bases, due to these generally spiny; apices hidden between petiole bases; without adventitious buds. Leaves 120 – 200 cm long. Petioles 20 – 40 cm long, muricate to weakly aculeate with prickles to 2 mm long, dark brown to blackish, rarely dark stramineous, opaque, basaly with persistent scurf consisting of appressed reddish brown trichomidia and erect, white, lanceolate to ovate squamules to 1 mm long with subentire to erose margins; hairs absent; petioles basally with a discontinuous line of distant aerophores to 6.0 × 1.0 mm on each side, gray brown and inconspicuous in dried material. Petiole scales long lanceolate, to 15.0 – 25.0 × (1.5 –) 2.0 – 3.5 (– 4.0) mm, rather thin textured, bases cordate, pseudopeltately attached, apices long attenuate, straight to falcate, often strongly undulate; proximal scales concordantly to discordantly bicolorous, auburn to yellowish brown with white margins, distal ones often almost concolorous stramineous to white with a dark apical streak; differentiated margins without setae or teeth, usually persistent. Blades to 180 × 120 – 140 cm, bipinnate-pinnatifid, firm herbaceous, matte, dark-green adaxially, dark olive green abaxially; apices gradually reduced. Rachises inermous to sparsely verrucate proximally, dark brown abaxially and adaxially; pubescent with tan to brown multicellular hairs 1.0 – 1.5 mm long, antrorsely curved and persistent adaxially, abaxially glabrescent with few thin hairs and appressed trichomidia, leaving the cortex smooth. Largest pinnae 24 – 40 cm long, pinnae subsessile or stalked to 1.0 (– 1.7) cm, 15 – 20 pairs per leaf, patent to weakly ascending, alternate, inarticulate, distally narrowly green-alate, distal segments simply adnate before ending in a pinnatifid apical section; basal pairs smaller than than the medial pinnae, reflexed. Costae to 1.5 – 2.0 mm wide, inermous to sparsely verrucate, dark brown to grayish brown abaxially, darker adaxially; glabrous abaxially, with tan to brown, antrorsely curved multicellular hairs 0.5 – 1.0 mm long adaxially; junctures of costae and rachises not swollen, abaxially often black when dried, each with an inconspicuous planar, elliptic aerophore, to 2 × 1 mm, dark orange-brown. Largest pinnules 45 – 70 × 11 – 17 mm, short-stalked to 3 mm, inarticulate, 1.5 – 2.0 (– 2.2) cm between the stalks / costules, pinnules lanceolate, rarely linear-oblong, truncate to cordate basally, long-acute to attenuate apically with subentire to weakly crenulate margins; costules dark carnose to dark grayish brown adaxially and abaxially, proximally often darker brown; adaxially strongly prominent, ridged, and densely hairy with tan to brown, antrorsely curved multicellular hairs to 1.0 mm long, abaxially weakly prominent, glabrous or glabrescent with appressed, tan to brown unicellular trichomidia and thin tortuous hairs; also with white to brown squamules, bullate ones to 1.0 mm long, flat lanceolate ones to 3 × 2 mm, with entire margins and short-attenuate tips; costules basally without pneumathodes. Segments to 7.0 – 10.0 × 3.0 – 4.0 mm, rarely remote and then connected by laminar tissue, patent to strongly ascending, distally falcate, tips obtuse to rounded, proximal segments alternate to subopposite, usually a bit shorter than following segments; sinuses acute to obtuse, to 1.0 (– 2.0) mm wide; margins subentire to crenate; margins not differently incised in proximal segments of a pinnule; veins planar on both sides or weakly protruding adaxially, dark carnose to dark grayish brown; veins adaxially glabrous except for some multicellular hairs to 0.5 mm long on the midveins, abaxially glabrous except for some appressed trichomidia; midveins with some pure white to light brown squamules, mostly bullate ones to 1.0 × 0.5 mm; sterile and fertile veins simple or forked. Sori 1.0 – 1.2 mm diam., medial to supramedial, indusia lacking; receptacles globose, 0.3 – 0.4 mm diam., paraphyses few to numerous, stiff, hyaline, tan to brown, shorter than the sporangia (0.2 – 0.3 mm long). Spores tetrahedral-globose, pale yellow to white, appearing hyaline.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782829BC19FDEABA02F5C550B2.taxon	distribution	Distribution and ecology Magdalena Valley and Cordillera Oriental of Colombia, Cordillera de la Costa in Venezuela at elevations of 1200 – 2430 (– 2740) m a. s. l.; one sterile specimen recorded from Trinidad (Barrington 1978).	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782829BC19FDEABA02F5C550B2.taxon	discussion	Remarks Cyathea squamata has only rarely been confused with C. pungens, despite sharing the diagnostic characters of being exindusiate, having bicolorous brown petiole scales with white margins, and whitish bullate squamules on the blade. Cyathea squamata differs in having more pinna pairs per blade, generally smaller pinnules with notable short stalks and cordate bases, by which means the blade appears more delicate and finely dissected than in C. pungens. Moreover, C. squamata has a mixture of larger and smaller squamules on the axes and veins abaxially, giving it an untidy appearance, whereas the squamules in C. pungens are so small and scarce that they are hardly noticeable.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782827BC1BFDAEBABDF334569E.taxon	description	Fig. 13	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782827BC1BFDAEBABDF334569E.taxon	materials_examined	– 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 [UC 1564628]!.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782827BC1BFDAEBABDF334569E.taxon	etymology	Etymology The specific epithet honors Henk van der Werff, botanist at the Missouri Botanical Garden and collector of the type specimen.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782827BC1BFDAEBABDF334569E.taxon	materials_examined	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. – 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. ECUADOR – Morona-Santiago • Along road between Santiago and Río Morona; 2 º 58 ′ 24 ″ S, 77 º 49 ′ 36 ″ W; 322 m a. s. l.; 10 Jul. 2004; T. B. Croat et al. 90750; MO • 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 • 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 • 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 • 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. 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.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782827BC1BFDAEBABDF334569E.taxon	description	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.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782827BC1BFDAEBABDF334569E.taxon	distribution	Distribution and ecology Southern Colombia, Ecuador and northern Peru (Fig. 2 A) at elevations of 150 – 1400 m a. s. l., in deeply shaded forest on steep rocky slopes.	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
163C91782827BC1BFDAEBABDF334569E.taxon	discussion	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. 13 A), 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). 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. 2 A), has many biogeographic ties with the Guayana highlands (Lehnert & Tejedor 2016).	en	Lehnert, Marcus, Tejedor, Adrian, Kessler, Michael, Rodríguez Duque, Wilson D., Gallego, Luis Fernando Giraldo (2025): A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae). European Journal of Taxonomy 988: 1-57, DOI: 10.5852/ejt.2025.988.2883, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2883/13079
