Plant Ecology and Evolution 152(3): 521-530, doi: 10.5091/plecevo.2019.1545
Taxonomic novelties in Cyclodium (Dryopteridaceae) and a key to the species with free veins
expand article infoAmabily Bohn, Fernando B. Matos, Paulo H. Labiak
‡ Universidade Federal do Paraná, Dept. de Botânica. Caixa Postal 19031, 81531-980, Curitiba, Paraná, Brazil
Open Access

BackgroundCyclodium is a neotropical fern genus comprising about ten species. Most species are found in northern South America, and the foothills of the Guiana Shield is an important region for species diversification. Our phylogenetic and taxonomic studies of the genus demonstrated the need to describe a new species and to recognize a variety at species level.

Methods – This study is based on herbarium specimens from CAY, HUA, INPA, MBM, NY, P, RB, UC, UFP, UPCB, and US. Morphological characters were analyzed using standard procedures. The indumentum and spores of the new species were studied using a scanning electron microscope. Species delimitation is proposed based on our preliminary phylogenetic studies, as well as on morphology and geographical distribution.

Key resultsCyclodium alansmithii Bohn & Labiak is recognized as a new species, described, and illustrated. The most similar species is Cyclodium inerme (Fée) A.R.Sm., from which it differs by ovate-lanceolate and bicolored scales, reduced fronds, truncate pinna bases, and non-ciliate indusia. Cyclodium alansmithii is currently assessed as Endangered (EN) using IUCN criteria, but more fieldwork and herbarium studies are necessary to establish a more accurate conservation assessment. Cyclodium trianae (Mett.) A.R.Sm. var. chocoense A.R.Sm. is here elevated to species rank. A key to species of Cyclodium with free veins is provided.

Amazon, biodiversity, ferns, Guayana Shield, polybotryoids, taxonomy


  • Berry P.E., Holst B.K., Yatskievych K. (1995) Flora of the Venezuelan Guayana vol. 1: Introduction. Portland, Timber Press.
  • Christensen C. (1913) A monograph of the genus Dryopteris. Part I. The tropical American pinnatifid bipinnatifid species. Kongelige Danske Videnskabernes Selskabs Skrifter. Naturvidenskabelig og Mathematisk Afdelung, ser. 7 10: 55–282.
  • Christensen C. (1920) A monograph of the genus Dryopteris. Part II. The tropical American bipinnate-decompound species. Kongelige Danske Videnskabernes Selskabs Skrifter. Naturvidenskabelig og Mathematisk Afdelung, ser. 8 6: 1–132.
  • Cremers G., Kramer K.U., Moran R.C., Smith A.R. (1993) Dryopteridaceae. In: Görts-van Rijn A.R.A. (ed.) Flora of the Guianas. Series B: Ferns and Fern allies fasc. 6: 3–65. Koenigstein, Koeltz Scientific Books.
  • Engels M.E., Canestraro B.K. (2017) ×Cyclobotrya: A new hybrid genus between Cyclodium and Polybotrya (Dryopteridaceae) from the Brazilian Amazon. Brittonia 69(3): 307–312.
  • Funk V.A., Berry P.E. (2005) Chapter 4.3: The Guiana Shield. In: Krupnick G., Kress W.J. (eds) Plant conservation: A natural history approach: 76–79. Chicago, University of Chicago Press.
  • Garcia P.A., Salino A. (2008) Dryopteridaceae (Polypodiopsida) no estado de Minas Gerais, Brasil. Lundiana 9(1): 3–27.
  • IUCN (2001) IUCN Red List Categories and Criteria, v.3.1. Second edition. Gland, Switzerland and Cambridge, United Kingdom, IUCN Species Survival Commission [online]. Available at [accessed 5 Aug. 2018].
  • Moran R.C. (1987) Monograph of the neotropical fern genus Polybotrya (Dryopteridaceae). Illinois Natural History Survey Bulletin 34: 1–138.
  • Moran R.C. (1991a) Monograph of the neotropical fern genus Stigmatopteris (Dryopteridaceae). Annals of the Missouri Botanical Garden 78(4): 857–914.
  • Moran R.C. (1991b) Eight new species of tree ferns (Cyathea, Cyatheaceae) from the American Tropics and three new combinations. Novon 1(2): 88–104.
  • Moran R.C. (1992) The Potato Fern. Fiddlehead Forum 19(3): 18–20.
  • Moran R.C. (1995) The importance of mountains to pteridophytes, with emphasis on Neotropical montane forests. In: Churchill S.P., Balslev H., Forero E., Luteyn J.L. (eds) Biodiversity and conservation of Neotropical montane forests: Proceedings of the Neotropical Montane Forest Biodiversity and Conservation Symposium, the New York Botanical Garden, 21–26 June 1993: 359–363. New York, New York Botanical Garden.
  • Moran R.C., Labiak P.H. (2015) Phylogeny of the polybotryoid fern clade (Dryopteridaceae). International Journal of Plant Sciences 176(9): 880–891.
  • Mori S.A., Cremers G., Gracie C., Granville J.-J. de, Hoff M., Mitchell J.D. (1997) Vascular plants of central French Guiana. Part 1. Bronx, The New York Botanical Garden.
  • Murakami N., Moran R.C. (1993) Monograph of the Neotropical species of Asplenium sect. Hymenasplenium (Aspleniaceae). Annals of the Missouri Botanical Garden 80(1): 1–38.
  • Øllgaard B. (1988) Part I: Lycopodiaceae. In: Harling G., Andersson L. (eds) Flora of Ecuador 33: 1–155. Arlöv, Berlings.
  • PPG I (2016) A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54(6): 563–603.
  • Quantum GIS Development Team (2013) Quantum GIS Geographic Information System. Open Source Geospatial Foundation Project. Available at [accessed 5 Aug. 2018].
  • Smith A.R., Moran R.C. (1995) Cyclodium. In: Moran R.C., Riba R. (eds) Flora Mesoamericana vol. 1, Psilotaceae a Salviniaceae: 211–212. Mexico, Universidad Nacional Autónoma de México.
  • Smith A.R., Mickel J.T., Moran R.C. (1995) Dryopteridaceae. In: Steyermark J.A., Berry P.E., Holst B.K. (eds) Flora of the Venezuelan Guayana vol. 2: 73–128. Hong Kong, Timber Press.
  • Thiers B. (2018, continuously updated). Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Available at [accessed 19 Dec. 2018].
  • Tryon R.M., Stolze R.G. (1991) Pteridophyta of Peru, Part IV, Dryopteridaceae. Fieldiana, Botany, New Series 27: 1–176.