A reassessment of the Neotropical Cyathea pungens complex (Cyatheaceae)
Abstract
Cyathea pungens, a widespread Neotropical scaly tree fern, is reassessed taxonomically. Based on extensive field observations in South America and the Caribbean, and comparative studies of herbarium specimens and online resources, we define C. pungens as a mainly Amazonian species that extends into the Caribbean but neither beyond west of the Andes nor into the Brazilian Mata Atlantica. Records from west of the Colombian Cordillera Oriental belong to other species, which differ slightly but consistently in the laminar indument (i.e., abundance and color of bullate squamules, and presence of hairs variable vs constant) and prevailing pinnule shape (with only obtuse tips vs acute and obtuse tips). The weak differences in the specimens are generally corroborated by different appearances in the field (i.e., shape of the blades, and number of pinnae) and preferred elevational ranges. The distributions of C. pungens and C. oblonga overlap in the area of the Guyana shield, but C. oblonga is also found along the Caribbean coast and northern lowlands of Colombia and Venezuela. Both species are rarely found above 1000 m and are largely replaced in the mountains by C. klotzschiana (Colombian Cordillera Oriental, Sierra Nevada de Santa Marta and Venezuelan coastal ranges) and C. pastazensis (Colombian Cordillera Central and Occidental as well as eastern Andean slopes of Ecuador). To this group further belongs C. dombeyi from the Andes of Peru and Bolivia, and the reinstated C. pycnocarpa and C. floribunda from Peru and Bolivia. Several additional species are tentatively placed in this group: the northern Andean C. squamata, the Guayanan C. neblinae, Brazilian C. praecincta and Andean C. werffii.
References
Allaire J.J. 2012. UseR! The R User Conference (useR! 2011) held on August 16-18 2011 at the University of Warwick in Coventry, UK. Book of contributed abstracts, compiled 28 Aug 2011. Available from https://de.scribd.com/document/649443739/Allaire-2012 [accessed 7 Apr. 2025].
Barrington D.S. 1976. New taxa and nomenclatural changes in the genus Trichipteris (Cyatheaceae). Rhodora 78 (813): 1–5. Available from https://www.jstor.org/stable/23311305 [accessed 26 Feb. 2025].
Barrington D.S. 1978. A revision of the genus Trichipteris. Contributions from the Gray Herbarium of Harvard University 208: 3–93. https://doi.org/10.5962/p.336446
Berry P.E., Holst B.K. & Yatskievych K. 1995. Introduction. In: Berry P.E., Holst B.K. & Yatskievych K. (eds) Flora of the Venezuelan Guayana, Introduction, Vol 1: xv–xx. Missouri Botanical Garden Press, St. Louis.
Cárdenas D., Rodríguez W.D., García N., Sua S., Lehnert M. & Giraldo F. 2019. Libro Rojo de Plantas de Colombia Vol. 7. Helechos Arborescentes. Instituto Amazónico de Investigaciones Científicas SINCHI, Bogotá.
Carvalho F.A., Salino A. & Zartman C.E. 2012. New country and regional records from the Brazilian side of Neblina Massif. American Fern Journal 102: 228–232. https://doi.org/10.1640/0002-8444-102.3.228
Cremers G. & Boudrie M. 2007. Les Ptéridophytes des Guyanes: les spécimens de référence depuis Aublet (1775) à nos jours. Le Journal de Botanique 40: 3–111. https://doi.org/10.3406/jobot.2007.2039
Cronquist A., Takhtajan. A. & Zimmermann W. 1966. On the higher taxa of Embryobionta. Taxon 15: 129–134. https://doi.org/10.2307/1217531
Desvaux N.A. 1827. Prodrome de la famille des fougères. Mémoires de la Société Linnéenne de Paris 6: 171–337.
Domin C. 1929a. Pteridophyta. Soustavný přehled žijících i vyhynulých kaproďorostů. Česká akademie věd a umění, Prague.
Domin C. 1929b. The Pteridophyta of the Island of Dominica, With Notes on Various Ferns from Tropical America. Rozpravy Královské české společnosti nauk, Třída matematicko-přírodovědecká, Nová řada, číslo 2 / Memoirs of the Royal Czech Society of Sciences, Division of Natural History and Mathematics, New Series, No. 2., Dr Ed. Grégr. & Son, Prague.
Domin C. 1929c. New ferns from Tropical America and the West Indies. Bulletin of Miscellaneous Information (Royal Gardens, Kew) 7: 215–222. https://doi.org/10.2307/4113535
Domin C. 1930. The species of the genus Cyathea J.E.Sm. Acta Botanica Bohemica 9: 85–174.
Farwell O.A. 1931. Fern Notes II. Ferns in the Herbarium of Parke, Davis & co. American Midland Naturalist 12: 231–311. https://doi.org/10.2307/2420088
Hieronymus G.H.E.W. 1906. Plantae Stübelianae. Pteridophyta. Hedwigia 45: 215–232.
Hooker W.J. & Baker J.G. 1874. Synopsis Filicum, 2nd Ed. R. Hardwicke, London. Available from http://biodiversitylibrary.org/page/28796458 [accessed 26 Feb. 2025].
Kaulfuss G.F. 1827. Das Wesen der Farrenkräuter: besonders ihrer Fruchttheile zugleich mit Rücksicht auf systematische Anordnung betrachtet und mit einer Darstellung der Entwickelung der Pteris serrulata aus dem Samen begleitet. Erste Hälfte (Vol. 1). C. Cnobloch, Leipzig.
Klotzsch J.F. 1844. Beiträge zu einer Flora der Aequinoctial-gegenden der neuen Welt. Filices. Linnaea 18: 516–556.
Klotzsch J.F. 1847. Beiträge zu einer Flora der Aequinoctial-gegenden der neuen Welt. Filices. Linnaea 20: 337–445.
Klotzsch J.F. 1852. Einige neue Baumfarrn. Allgemeine Gartenzeitung (Otto & Dietrich) 30: 41–43.
Kunze G. 1834. Synopsis plantarum cryptogamicarum ab Eduardo Poeppig in Cuba insaula et America merdionali collectarum. Linnaea 9: 1–111. https://doi.org/10.5962/bhl.title.51054
Kunze G. 1839. Centuria Quarta. Flora 22 (1, Beibl.): 1–64.
Kunze G. 1846. Die Farrnkräuter in kolorirten Abbildungen naturgetreu erläutert und beschrieben, 1. Band, Taf. 81–90. E. Fleischer, Leipzig. https://doi.org/10.5962/bhl.title.69313
Lehnert M. 2008. On the identification of Cyathea pallescens (Sodiro) Domin: typifications, reinstatements and new descriptions of Neotropical tree ferns. Botanical Journal of the Linnean Society 158 (4): 621–649. https://doi.org/10.1111/j.1095-8339.2008.00887.x
Lehnert M. 2009. Resolving the Cyathea caracasana complex. Stuttgarter Beiträge für Naturkunde A, Neue Serie 2: 409–445. Available from https://www.zobodat.at/pdf/Stuttgarter-Beitraege-Naturkunde_NS_2_A_0409-0445.pdf [accessed 26 Feb. 2025].
Lehnert M. 2011. Species of Cyathea in America related to the western Pacific species C. decurrens. Phytotaxa 26: 39–59. https://doi.org/10.11646/phytotaxa.26.1.6
Lehnert M. 2014. Do you know Cyathea divergens (Cyatheaceae-Polypodiopsida)? Phytotaxa 161 (1): 1–42. https://doi.org/10.11646/phytotaxa.161.1.1
Lehnert M. 2016. A synopsis of the exindusiate species of Cyathea (Cyatheaceae-Polypodiopsida) with bipinnate-pinnatifid or more complex fronds, with a revision of the C. lasiosora complex. Phytotaxa 243 (1): 1–53. https://doi.org/10.11646/phytotaxa.243.1.1
Lehnert M. & Coritico F.P. 2018. The genus Dicksonia (Dicksoniaceae-Cyatheales) in western Malesia. Blumea – Biodiversity, Evolution and Biogeography of Plants 63 (3): 268–278. https://doi.org/10.3767/blumea.2018.63.03.02
Lehnert M. & Cámara-Leret R. 2018. Dicksonia utteridgei, a new species of hairy tree fern (Dicksoniaceae-Cyatheales) from New Guinea. Blumea – Biodiversity, Evolution and Biogeography of Plants 63 (2): 140–143. https://doi.org/10.3767/blumea.2018.63.02.09
Lehnert M. & Cámara-Leret R. 2019. New species of scaly tree ferns (Cyatheaceae) from New Guinea, and new combinations for the family for Malesia. Kew Bulletin 74: 46. https://doi.org/10.1007/s12225-019-9823-4
Lehnert M. & Tejedor A. 2016. Three new scaly tree fern species (Cyathea-Cyatheaceae) from the Amotape-Huancabamba Zone. American Fern Journal 106 (3): 175–190. https://doi.org/10.1640/AFJ-D-16-00002.1
Lehnert M. & Weigand A. 2013. A proposal to distinguish several taxa in the Brazilian tree fern Cyathea corcovadensis (Cyatheaceae). Phytotaxa 155 (1): 35–49. https://doi.org/10.11646/phytotaxa.155.1.3
Lehnert M. & Weigand A. 2017. A synopsis of the Neotropical species of Cyathea (Cyatheaceae; Polypodiopsida) with bipinnate fronds. Brittonia 69: 71–90. https://doi.org/10.1007/s12228-016-9445-1
Lehnert M., Rodríguez Duque W.D., Giraldo Gallego L.F. & Tejedor A. 2021. The scaly tree ferns allied to Cyathea multiflora (Cyatheaceae) in Colombia and neighboring countries. American Fern Journal 111 (4): 251–307. https://doi.org/10.1640/0002-8444-111.4.251
Lellinger D.B. 1987 [1988]. The disposition of Trichopteris (Cyatheaceae). American Fern Journal 77 (3): 90–94. https://doi.org/10.2307/1547497
Lellinger D.B. 2002. A modern multilingual glossary for taxonomic pteridology. Pteridologia 3: 1–264. https://doi.org/10.5962/bhl.title.124209
Leunis J. 1877. Synopsis der Pflanzenkunde (Ed. 2) 3. Specielle Botanik, Kryptogamen. Hahn, Hannover.
Loiseau O., Weigand A., Noben S., Rolland J., Silvestro D., Kessler M., Lehnert M. & Salamin N. 2019. Slowly but surely: gradual diversification and phenotypic evolution in hyper-diverse tree fern family Cyatheaceae. Annals of Botany 125 (1): 93–103. https://doi.org/10.1093/aob/mcz145
Maxon W.R. 1925. New Tropical American ferns: II. American Fern Journal 15 (2): 54–57. https://doi.org/10.2307/1544002
Mettenius G. 1856. In: Hohenacker R.F. (ed.) Filices Lechlerianae, chilenses ac peruanae (Vol. 1). L. Voss, Leipzig.
Moran R.C. 1991. Eight new species of tree ferns (Cyathea, Cyatheaceae) from the American tropics and three new combinations. Novon 1 (2): 88–104. https://doi.org/10.2307/3391634
Noben S. & Lehnert M. 2013. The genus Dicksonia (Dicksoniaceae) in the western Pacific. Phytotaxa 155 (1): 23–34. https://doi.org/10.11646/phytotaxa.155.1.2
PPG 1. 2016. Pteridophyte Phylogeny Group. A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54 (6): 563–603. https://doi.org/10.1111/jse.12229
Presl C. 1836. Tentamen Pteridographiae. A. Haase, Prague.
Presl C. 1851 [1849]. Epimeliae Botanicae. A. Haase, Prague.
Rosenstock E. 1917. Filices palaeotropicae novae Herbarii Lugduno-Batavi. Mededeelingen vanʼs Rijks-Herbarium. Leiden 31: 1–8. Available from https://repository.naturalis.nl/pub/508337/MRHL1917031001001.pdf [accessed 26 Feb. 2025].
Rosenstock E. 1928. Filices novae a cl. Dr. O. Buchtien in Bolivia collectae. VI. Repertorium Specierum Novarum Regni Vegetabilis 25 (4–6): 56–64. https://doi.org/10.1002/fedr.19280250408
Schwartsburd P.B., Becari-Viana I., Lopes L.R. & Lehnert M. 2015. A new hybrid and further taxonomic notes on Brazilian tree ferns (Cyatheaceae). Phytotaxa 231 (1): 42–52. https://doi.org/10.11646/phytotaxa.231.1.4
Smith A.R. 1990. Pteridophytes of the Venezuelan Guayana: new species. Annals of the Missouri Botanical Garden 77 (2): 249–273. https://doi.org/10.2307/2399539
Smith J.E. 1793. Tentament botanicum de filicum generibus dorsiferarum. Mémoires de l'Academie royale des Sciences (Turin) 5 (2): 401–422. Available from http://onlinebooks.library.upenn.edu/webbin/gutbook/lookup?num=39534 [accessed 7 Apr. 2025].
Splitgerber F.L. 1840. Enumaratio Filicum et Lycopodiacearum quas in Surinamo legit Splitgerber. Tijdschrift voor Natuurlijke Geschiedenis en Physiologie 7: 391–444.
Stresemann E. 1950. Die brasilianischen Vogelsammlungen des Grafen von Hoffmannsegg aus den Jahren 1800–1812. Bonner Zoologische Beiträge 1:43–51. Available from https://zoologicalbulletin.de/BzB_Volumes/Volume_01_2_3_4/126_143_BZB01_2_3_4_Stresemann_Erwin.PDF [accessed 26 Feb. 2025].
Tejedor A. & Calatayud G. 2017. Eleven new scaly tree ferns (Cyathea: Cyatheaceae) from Peru. American Fern Journal 107 (3): 156–191. https://doi.org/10.1640/0002-8444-107.3.156
Tejedor A. & Areces-Berazain F. 2018. Cyathea ruttenbergii, a new tree fern (Cyatheaceae, Polypodiopsida) from Puerto Rico. Phytotaxa 336 (3): 279–285. https://doi.org/10.11646/phytotaxa.336.3.6
Tejedor A. & Areces-Berazain F. 2021. An update on the distribution and affinities of Cyathea ruttenbergii (Cyatheaceae, Polypodiopsida). Phytotaxa 516 (1):108–110. https://doi.org/10.11646/phytotaxa.516.1.9
Thiers B. 2024 (continuously updated). Index Herbariorum: A Global Directory of Public Herbaria and Associated Staff. New York Botanical Garden’s Virtual Herbarium. Available from https://sweetgum.nybg.org/science/ih/ [accessed 7 Jun. 2022].
Tryon R.M. 1970. The classification of the Cyatheaceae. Contributions from the Gray Herbarium of Harvard University 200: 3–53. https://doi.org/10.5962/p.336422
Willdenow C.L. 1810. Species Plantarum, Ed. 4. G.C. Nauk, Berlin.
Zuquim G., Costa F.R.C., Prado J. & Tuomisto H. 2008. Guide to the Ferns and Lycophytes of REBIO Uatumã, Central Amazonia. nstituto Nacional de Pesquisas da Amazonia, Manaus.
Copyright (c) 2025 Marcus Lehnert, Adrian Tejedor, Michael Kessler, Wilson D. Rodríguez Duque, Luis Fernando Giraldo Gallego

This work is licensed under a Creative Commons Attribution 4.0 International License.
Creative Commons Copyright Notices
Authors who publish with this journal agree to the following terms:
- Authors retain copyright and grant the journal right of first publication with the work simultaneously licensed under a Creative Commons Attribution License (CC BY 4.0) that allows others to share the work with an acknowledgement of the work's authorship and initial publication in this journal.
- Authors are able to enter into separate, additional contractual arrangements for the non-exclusive distribution of the journal's published version of the work (e.g., post it to an institutional repository or publish it in a book), with an acknowledgement of its initial publication in this journal.
- Authors are NOT PERMITTED to post their submitted work online (e.g., in institutional repositories or on personal websites) prior to publication, as it may lead to nomenclatural issues.
