Cyathea pungens (Willd.) Domin, 1929

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, pp. 1-57 : 29-36

Cyathea pungens (Willd.) Domin

Figs 10 , 13B

Cyathea pungens (Willd.) Domin ( Domin 1929a: 263) . – Polypodium pungens Willd. ( Willdenow 1810: 206) . – Alsophila pungens (Willd.) C.Presl ( Presl 1836: 61) . – Trichipteris pungens (Willd.) R.M.Tryon ( Tryon 1970: 46) .

– Type: BRAZIL • “ Brasilia ”; J.C. von Hoffmannsegg s.n.; holotype: B [ B-W 19716-011 , B-W 19716-012 , B-W 19716-013 , 3 sheets, Herb. Willdenow]!; isotypes: B [ B 20 0000318 , Herb. Link]!, NY [ 00148762 , fragment of B], US [ 00066283 , fragment of B] image! .

Polypodium procerum Willd. ( Willdenow 1810: 206) . – Alsophila procera (Willd.) Desv. ( Desvaux 1827: 319) . – Cyathea willdenowiana Domin ( Domin 1930: 171) . – Trichipteris procera (Willd.) R.M.Tryon ( Tryon 1970: 46) .

– Type: BRAZIL • “ Brasilia ”; J.C. von Hoffmannsegg s.n.; holotype: B [ B-W 19717-011 , B-W 19717-012 , B-W 19717-013 , 3 sheets, Herb. Willdenow]!; isotypes: NY [ 00148758 , fragment of B], US [ 00066279 , fragment of B]!.

Alsophila infesta Kunze ( Kunze 1834: 98) . – Cyathea infesta (Kunze) Domin ( Domin 1930: 125) . – Trichipteris infesta (Kunze) R.M.Tryon ( Tryon 1970: 45) . – Alsophila phalerata var. infesta (Kunze) Farw. ( Farwell 1931: 250) .

– Type: PERU • Huánuco, “ Peruviae ad Miss. Tocache fluv. Huallgae vicinam”; Aug. 1830; E. Poeppig s.n.; holotype: LZ [destroyed]; lectotype: W [ W0056018 , W0056019 , 2 sheets]!, designated by Lehnert 2016; isolectotypes: BM [ BM000937693 , fragment of W]!, MO [ MO-1855206 ] image!.

Alsophila subaculeata Splitg. ( Splitgerber 1840: 430) .

– Type: SURINAME • Canawappibo; Dec. 1837; F.L. Splitgerber s.n.; lectotype: L [ L.1285146 , L.1285147 , 2 sheets]!, first step designated by Barrington 1978, second step designated here; isolectotypes: NY [fragment of P] n.v. (fide Barrington 1978), US [ 00066287 , fragment of P] image!.

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 [ BM000777031 , BM000777032 (mix with C. dombeyi ), BM000777346 ], P [ P00642346 p.p., P00642349 , P00642350 ] • 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 25816a; 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 , d1, 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 .

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.

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.

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 , 2A ).

Figures

Fig. 10. Cyathea pungens (Willd.) Domin. A. Petiole with scurf and scales (Tuomisto et al. 16679, BONN). B. Petiole scale from petiole base, showing only thin white differentiated margin (Tuomisto et al. 16679, BONN). C. Pinna, silhouette (S. Maciel 1541, BONN). D. Fertile pinnule, abaxially (E. Poeppig s.n., W0056018). E. Fertile segment, abaxially (S. Maciel 1541, BONN).

Fig. 13. Cyathea werffii R.C.Moran in the field (Chiriaco, Prov. Bagua, Dept. Amazonas, Peru). A. Plant with upright fully bipinnate-pinnatifid leaves; note smaller and less dissected leaves left and right coming from the same rhizome. B. Comparison of soral lines of C. werffii (bottom) with “sessile” C. pungens (Willd.) Domin. Note the characteristic darkened spots at the base of the costules in both species. (photos © M. Lehnert)

Fig. 1. Distribution of Cyathea pungens (Wild.) Domin and its different forms; most likely area of origin of the type deduced from literature research. Gaps in Peru and Brazil presumably reflect collection biases, whereas the absence in central and south Bolivia and northern parts of Colombia and Venezuela probably represent true absence.

Fig. 2. Distribution of taxa of the Cyathea pungens complex except C. pungens (Wild.) Domin s. str. A. Cyathea dombeyi (Desv.) Lellinger (blue triangles), C. pycnocarpa (Kunze) Domin (green dots), C. squamata (Klotzsch) Domin (black squares), C. werffii R.C.Moran (magenta dots). B. Cyathea floribunda (Hook. & Baker) Domin (black squares), C. klotzschiana Domin (black dots), C. oblonga Domin (orange diamonds), C. pastazensis (Hieron.) Domin (blue triangles), C. praecincta (Klotzsch) Domin (green dots); note that the ranges of C. klotzschiana in the north and C. floribunda in the south do not overlap but are separated by C. pastazensis.