• Wita Wardani Herbarium Bogoriense (BO)
  • Bayu Adjie BRIN
  • Kusumadewi Sri Yulita BRIN
  • Andi Salamah UI



Athyrium nigripes complex, Athyriaceae, Java, clustering


Athyrium nigripes is a highly variable species described and recorded mostly from Java and few from surrounding islands. There are different arguments on its delimitation and hence its extent of distribution. We attempted to elucidate disagreement on delimitation of four related species described from Java: Athyrium nigripes, Aspidium costale, At. pulcherrimum and At. nitidulum with phenetic approach. The analysis formed four clusters; cluster I consist of the large sized and include type specimen of At. pulcherrimum and Asp. costale, cluster II for plants with standard form and include type specimen of At. nigripes, cluster III only for serrated plants and cluster IV contain the At. nitidulum form. Since Asp. costale clustered in group I, this species is more closely related to At. pulcherrimum than to At. nigripes nested in group II.  Large size and degree of division indicate major clustering within the complex species.


Alderwerelt van Rosenburgh, C.R.W.K. van. 1908. Malayan Ferns. Handbook to the Determination of the Ferns of the Malayan Islands. Landsdrukkerij, Batavia. P. 435.

Alderwerelt van Rosenburgh, C.R.W.K. van. 1917. Malayan Ferns and Fern Allies. Handbook to the Determination of the Ferns of the Malayan Islands. Supplement 1. Landsdrukkerij, Batavia. p. 278-281.

Alderwerelt van Rosenburgh, C.R.W.K. van. 1914. New or Interesting Malayan Ferns 6. Bull. Jard. Bot. Buitenzorg ser. 2 (16): 3.

Backer, C.A. & Posthumus, O. 1939. Varenflora voor Java: 137. ’s Lands Plantenuin, Buitenzorg.

Blume, C.L. Von 1828. Enumeratio plantarum Javae et insularum adjacentium minus cognitarium vel novarum ex herbaria Reinwardtii, Kuhlii, Hasseltii et Blumii, 2. Hartmann, The Hague, p. 162, 170.

Christensen, C. 1906. Index Filicum. Apud H. Hagerup, Hafniae.

Copeland, E.B. 1913. Notes on some Javan ferns. Phillip.J.Sci.,C.8:139-155

Fraser-Jenkins, C. R., Gandhi, K. N. & Kholia, B. S. 2018. An annotated checklist of Indian Pteridophytes Part-2 (Woodsiaceae to Dryopteridaceae). Bishen Singh Mahendra Pal Singh. Dehra Dun (India).

Hooker, W.J.1860. Species Fillicum Vol. 3. William Pamplin. London. P.222, 227

Hooker, W.J. & Baker, J.G. 1874. Synopsis of Filicum: or, A Synopsis of All Known Fern, including the Osmundaceae, Schizaeaceae, Marattiaceae, and Ophioglossaceae Chiefly derived from the Kew herbarium) Accompanied by figures representing the essential characters of each genus. 2nd Edition. Robert Hardwicke, London. p.227

Hope, C.W. 1902. The ferns of Northwest India. Jour. Bombay Nat. Hist. Soc. 14:119

Kunze, G. 1848. In Filices Javae Zollingerianas observationes continuatae Kunzii, Lipsiensis. Bot. Zeit. (Berlin) 6:191

Liu, Y.C., Chiou, W.-L. & Kato, M. 2011. Molecular Phylogeny and Taxonomy of the Fern Genus Anisocampium (Athyriaceae). Taxon 60: 824-830.

Mettenius, G. 1859. Uber Einige Farngattungen. IV. Asplenium Tafel IV-VI. Abh. Senckenberg. Naturf. Ges. 3:237

Moore, T. 1857. Index Filicum: A Synopsis, With Characters, of the Genera, and an Enumeration of the Species of Ferns, with Synonyms, References. Vol. 49. Pamplin, London.

PPG I. 2016. A community-derived classification for extant lycophyte and ferns. J. Syst. Evol. 54(6): 563-603

Raciborski, M.1898. Die Pteridophyten der Flora von Buitenzorg. Buchhandlung und Druckerei vormals E.J. Brill. Leiden. p.229, 230

Rothfels, C.J., Sundue, M.A., Kuo, L.-Y, Larsson, A., Kato, M., Schuettpelz, E. & Pryer, K.M. 2012. A revised family-level classification for eupolypod II ferns (Polypodiidae: Polypodiales). Taxon 61(3): 515–533.

Sledge, W.A. 1956. The nomenclature and taxonomy of Athyrium nigripes (Bl.) Moore, A. solenopteris (Kunze) Moore and A. praetermissum sp. nov. Annals and Magazine of Natural History. 9(102): 453-464

Swofford, D.L. 2003. PAUP*: Pyhlogenetic analysis using parsimony (*and other methods), version 4. Sinauer Associates. Sunderland.

Wang, Z.R., He, Z.R. & Kato, M. 2013. Athyriaceae. In: Wu, Z.Y. Raven, P.H. & Hong, D.Y. (eds.). Flora of China, vols 2-3 (Pteridophytes). Science Press, Beijing and Missouri Botanical Garden Press, St Louis, MO, pp. 418-534.

Wei, R., Ebihara, A., Zhu, Y.-M., Zhao, C.-F., Hennequin, S. & Zhang, X.-C. 2018. A total-evidence phylogeny of the lady fern genus Athyrium Roth. (Athyriaceae) with a new infrageneric classification. Mol. Phylog. Evol. 119: 25-36




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