The Polystichum Homepage

Welcome to the website for the hollyferns, members of the fern genus Polystichum and allies.
This website is maintained for the use of hollyfern enthusiasts and scholars
throughout the world by Dave Barrington at the University of Vermont in the United States.

Polystichum Homepage

Portrait of the Genus


Species List

Diversity Map

Interactive Keys


Cytology (PDF)

Barrington Web Page
(with publications)

Morphological Characters

Morphological Analysis

Lab Protocols


David Barrington, University of Vermont

November 9, 2013


Recent phylogenetic and taxonomic work allows the certain placement of Polystichum in a monophyletic group including Cyrtomium s.s. (excluding Cyrtomium section Balansae, which comprises polystichums, see Li et al. 2007, etc.), Phanerophlebia, Dryopteris, and Arachniodes (Schuettpelz and Pryer, 2007). Within this assemblage, Polystichum, Cyrtomium s.s., and Phanerophlebia form a monophyletic group, which includes at least the genera listed below marked with asterisks. At this writing, our evidence (i.e. the larger tree on this website) suggests that Cyrtomium s.s. is the sister genus to Polystichum. This latter group has been informally called the polystichoid ferns by recent authors (including Driscoll and Barrington 2007 and Lu et al. 2007). However, there is no formal taxonomic basis for this usage, which in fact relates to a much broader group of ferns (details below). A modern subfamilial taxonomy for the Dryopteridaceae is needed. Though I am far from done mining the nineteenth-century literature for supergeneric names relevant to Polystichum, here are my notes to date – on the work of Mettenius and Todaro.

The Polystichoideae of Mettenius, G.F., 1858. Über einige Farngattungen. IV. Phegopteris und Aspidium. H. L. Brönner, Frankfurt A.M.

Mettenius was perhaps the first to use the name Polystichoideae. He characterizes the group as having what Christensen was to call in his Monograph of Dryopteris anadromous venation, including as examples species belonging to the current genera Dryopteris, Polystichum, Mensicium, Cyrtomium, and Arachniodes as well as Fadyenia and Adenoderris. Based on this set of examples, Mettenius’s taxon includes taxa now placed in the Thelypteridaceae, Tectariaceae, and Dryopteridaceae (sensu Alan Smith, 2006). In both Phegopteris and Aspidium Mettenius distinguished groups of species that he characterized as Polystichoideae, Sub-polystichoideae, Phegopteroideae, and Cyatheoideae. Thus, his approach was to articificially divide the indusiate from the exindusiate species, then organize them in groups by leaf architecture, with a largely parallel approach in the two major groups.

The Polysticheae of Todaro (1864, 1866).

Tribus Polysticheae Tod., Flora sicula exiccata [sic]. 1864. see Syn. Pl. Acot. Vasc. 33. Jan-Sep 1866.

Todaro did not describe the tribe in 1866. What happened in 1864 I do not know for want of that publication. The 1866 publication has three species of plants in the tribe, two are Dryopteris species in the modern sense, which he puts in Polystichum. The third is in Cystopteris. From this the tribe Polysticheae appears to be broadly defined to cover the modern Dryopteridaceae, i.e. it represents roughly the same part of fern diversity as Mettenius’s Polystichoideae. Thus, to be used for the polystichoid ferns, it would have to be emended – but it depends on what species are mentioned in the 1864 publication. It’s possible that no polystichums were included in the original species list, which might create problems.


Based on our own work (Driscoll and Barrington 2007, Li et al. 2008, McHenery et al. 2013), and that of Schuettpelz and Pryer 2007 and Liu et al., 2007, the following genera, among those immediately allied to Polysitchum, only Polystichum, Phanerophlebia, and Cyrtomium should be retained.

A Catalog of Generic Names Relevant to Polystichum, Phanerophlebia, and Cyrtomium

The genera in this list with asterisks should be treated as Polystichum species based on molecular evidence (their retention would render one of the above genera artificial). The genera with a sword (†) are candidate synonyms of Polystichum without molecular confirmation.

  • *Acropelta Nakai, Bull. Nation. Sc. Mus. (Tokyo) 33:5. 1953. Type species: Polystichum omeiense C.Chr., Ind. Fil. 67. 1905; 585. 1906. nom. nov. for Aspidium carvifolium Bak. JoB. 1888:288 (non A. carvifolium Kze. 1851).
  • *Adenoderris J. Sm., Historia Filicum 222. 1875. Type species: Adenoderris glandulosa (Hooker & Greville) J. Sm. , based on Aspidium glandulosum Hooker & Greville, Ic. Fil. t. 140. See McHenry, M., M. Sundue, and D. Barrington 2013. The genus Adenoderris  is artificial. Taxon 62:000-000.
  • *Aetopteron Ehrhart, Beitr. 4: 148. 1789. nom. nud. [aff. Fraser-Jenkins, 1997]. based on Polypodium aculeatum L. See House, H.D. 1920. The genus Aetopteron, Ehrhart. Amer. Fern J. 10:88-89.
  • Amblia C. Presl, Tent. Pterid. 184. 1836. Type species: Amblia juglandifolia (Humboldt & Bonpland ex Willd.)Presl, based on Polypodium juglandifolium Humboldt & Bonpland ex Willd., Sp. Pl. ed. 4 5:195. 1810. [exindusiate Phanerophlebia]
  • *Cyrtogonellum Ching, Bulletin of the Fan Memorial Institute of Biology 8(5): 327-328. 1938. Type species: Cyrtogonellum fraxinellum (H. Christ) Ching, Bull. Fan Mem. Inst. Biol. Bot. 8: 329. 1938, based on Aspidium fraxinellum Christ, Bull. Herb. Boiss. 7:15. 1899.
  • *Cyrtomidictyum Ching, Bull. Fan Mem. Inst. Biol., Bot. 10: 182. 1940. Type species: Cyrtomidictyum lepidocaulon(Hook.)Ching, based on Aspidium lepidocaulon Hooker, Sp. Fil. 4: 12. 1862. (note: has same type as Polystichum section Cyrtomiopsis Tagawa.)
  • Cyrtomium C. Presl, Tent. Pterid. 86. 1836. Type species [first listed]: Cyrtomium falcatum (L.f.) C. Presl, Tent. Pterid. 86. 1836--based on Polypodium falcatum L.f., Suppl. 446. 1781.
  • *Hemesteum Leveillé., Flore de Kouy-Tcheou Fougeres 496. 1915. Type species [first listed]: Hemesteum craspedosorum (Maxim.) Leveillé, based on Aspidium craspedosorum Maxim., Bull. Ac. St. Pétersb. 15:231. Mél. Biol. 7:341.1871. = Polystichum craspedosorum (Maxim.)Diels.
  • *Hypopeltis Michx. ex Bory, Bél. Voy. Bot. 2. 63. 1833. Type species: Hypopeltis lobulata Bory, Exp. Scientif. de Morée 32. 286. 1832. = Polystichum aculeatum (L.) Roth.
  • †Papuapteris C. Chr., Brittonia 2:300 f.2. 1937. Type species: Papuapteris lineare C. Chr. (New Guinea) Likely a Polystichum segregate – resembles American and Asian alpine species (but certainly a separate origin from these)
  • Phanerophlebia C. Presl, Tent. Pterid. 84. 1836. Type species: Phanerophlebia nobilis (Schecht. & Chsm.) C. Presl, based on Aspidium nobile Schlecht. and Cham., Linn. 5: 610. 1830.
  • Phanerophlebiopsis Ching, Acta Phyotax. Sin. 10:115. 1965. Type species: Phanerophlebiopsis tsiangiana Ching, Acta Phytotax. Sin. 10(2): 116, t. 21, f.5-7. 1965 & Bull. Bot. Res. 7(1): 72. 1986. Species subject to molecular analysis to date are allied to Arachniodes (Liu et al., 2007. International Journal of Plant Science 168: 1311–1323) , and the genus is treated as a synonym of Arachniodes in the Flora of China.  The type species is probably an Arachniodes as well.  However at least one species historically placed in Phanerophlebiopsis (Phanerophlebiopsis hookeriana [C. Presl] Fraser-Jenkins) is a Polystichum (Li et al., J. Plant Research 121: 19–26.)
  • *Plecosorus Fée, Gen. Fil. 150. Pl. 13. 1850-1852. Type species: Plecosorus speciosissimus (A.Br. ex Kunze) Fée, based on Cheilanthes speciosissima A.Br. ex Kunze, Anal. 35 t. 23. 1837. = Polystichum speciosissimum(A. Braun ex Kunze) Tryon & Tryon.
  • *Ptilopteris Hance, Journal of Botany, British & Foreign 1884:138-139. 1884. Type: Ptilopteris hancockii Hance, JoB. 1884:139. 1884. = Polystichum hancockii(Hance)Diels, Nat. Pfl. 1(4):191. 1899.
  • *Sorolepidium Christ, Bot. Gaz.51:350. 1911. emend. Ching, Acta Phytotax. Sin. 16(3):15. 1978. Type species: Sorolepidium glaciale (H. Christ) H. Christ, based on Polystichum glaciale H. Christ, Bull. Soc. Fr. 52. Mém. I:28. 1905.

THE GENUS POLYSTICHUM Roth, Tent. Fl. Germ. 3:31, 69. 1799. Nom. cons.

Type species Polystichum lonchitis (L.) Roth [because it is so designated as part of the designation of Polystichum as a nomen. conserv. in the 1988 International Code of Botanical Nomenclature, see Frazer-Jenkins (1997) for detailed commentary on typification].

SECTIONS AND SERIES (These have not been evaluated phylogenetically.)

  • sect. Achroloma Tagawa 1940: type species P. nepalense (Spreng.) C.Chr. (India, Yunnan, Szechuan, and Taiwan)
  • sect. Aculeata Christ, Über die australen Polystichum Arten. Ark. Bot. 4(12) 1-5 1905. Type species: "A[spidium]. aculeatum" (Europe and western Asia). First listed species is Polystichum orbiculare Desv. [sic], Mag. V. 321. 1811— a misspelling and incorrect generic placement of Aspidium orbiculatum Desv., Mag. V. 321. 1811 = Polystichum orbiculatum (Desv.) Remy & Fée (Mexico to Bolivia)
  • sect. Adenolepia Daigobo 1972: type species P. formosanum Rost. (Ryukyu and Taiwan)
  • sect. Afra Roux 1998: type species P. transkeiense W.B.G. Jacobsen (southern Africa)
  • sect. Cirrhosa Roux 1998: type species P. macleae(Baker)Diels (southern Africa)
  • sect. Crucifilix Tagawa 1940: type species P. tripteron (Kunze) C. Presl (Japan, eastern Siberia, southern Saghalien, Korea, and China)
  • sect. Cyrtomiopsis Tagawa 1940: type species P. lepidocaulon (Hook.) J.Sm. (Japan and Taiwan)
  • sect. Cyrtomiphlebium (of Polypodium) W.J. Hooker, Sp. Fil. 5:15. 1864: type species Phegopteris dubia Karst. = Polystichum dubium (Karst.)Diels
  • sect. Duropolystichum Fraser-Jenkins 1997: type species P. squarrosum (Don)Fée (Sino-Himalayan region)
  • sect. Haplopolystichum Tagawa 1940, emend. Daigobo 1972: type species P. deltodon(Bak.)Diels (Philippines, Japan, Taiwan, Szechuan, Yunnan, Hupeh, Kwangsi)
    • ser. Deltodontes L.L.Xiang ex Ching 1994
    • ser. Hecatoptera L.L. Xiang 1994
  • sect. Hypopeltis (Michx. ex Bory)T. Moore, Ind. Fil., lxxxiv. 1857. Type species (based on Bory, not Moore who does not indicate a type): P. aculeatum (L.) Roth (Europe)
  • sect. Insularum Roux 1998: type species P. drepanum (Sw.) C.Presl (Madeira only)
  • sect. Lacera Roux 1998: type species P. falcinellum (Sw.) C.Presl (Madeira only)
  • sect. Lasiopolystichum Daigobo 1972: type species P. lachenense (Hook.)Bedd. (Japan, Taiwan, China and Himalayas, Aleutian Islands)
    • ser. Moupinensia H.S.Kung & L.B.Zhang 1996
    • ser. Sinensia H.S.Kung & L.B.Zhang 1996
  • sect. Macropolystichum Daigobo 1972: type species P. eximium (Mett. ex Kuhn) C.Chr. (Ceylon to southeastern China, Tonkin, Taiwan, and Japan [Yaku Is.])
    • ser. Simplicipinnata Ching ex w. M. Chu et Z. R. He 2001: type species P. prionolepis Hayata (Taiwan)
    • ser. Bipinnata [as Bipinnatum] Ching ex w. M. Chu et Z. R. He 2001: type species P. eximium (Mett. ex Kuhn) C. Chr.
      • subser. Decurrentia Ching ex W. M. Chu et Z. R. He 2001: type species P. chunii Ching
      • subser. Disjuncta Ching ex W. M. Chu et Z. R. He 2001: type species P. eximium (Mett. ex Kuhn) C. Chr. [nom. superfl., = subser. bipinnata?]
  • sect. Mastigopteris Tagawa 1940: type species P. craspedosorum(Maxim.)Diels (Japan, Korea, north and central China, southern and eastern Manchuria, eastern Siberia)
  • sect. Metapolystichum Tagawa 1940: type species P. setiferum (Forst.) Moore ex Woyn. (Europe, northern Africa)
    • ser. Brauniana H.S. Kung & L.B. Zhang 1996
    • ser. Exauricula H.S. Kung & L.B. Zhang 1996
    • ser. Linearia H.S. Kung & L.B. Zhang 1996
    • ser. Makinoiana H.S. Kung & L.B. Zhang 1996
    • ser. Punctifera H.S. Kung & L.B. Zhang 1996
  • sect. Micropolystichum Daigobo, 1972: type species P. thomsonii (Hook.) Bedd. (North India, Szechuan, Yunnan, and Taiwan)
    • ser. Basigemmifera W. M. Chu et Z. R. He 2001: type species P. capillipes (Bak.)Diels (western China).
    • ser. Exgemmifera W. M. Chu et Z. R. He 2001: type species P. thomsoni (Hook. f.) Bedd. (China, Taiwan)
  • sect. Neopolystichum Ching ex L. B. Zhang et H. S. Kung, 1999. Type species P. grandifrons C. Chr. nom. nov. for P. grande Ching 1931 non Fée 1857
  • sect. Polystichum (Eupolystichum Diels emend. Tagawa 1940): type species P. lonchitis (L.) Roth (circumboreal)
  • sect. Prionolepia Daigobo 1972: type species P. biaristatum (Bl.) Moore (Java, Ceylon, Assam, Khassya, Burma, Yunnan, Annan, and Taiwan)
  • sect. Pungentia Roux 1998: type species P. pungens (Kaulf.) C.Presl (southern Africa)
  • sect. Scleropolystichum Daigobo 1972: type species P. aculeatum (L.) Roth (Europe)
  • sect. Sorolepidium (H. Christ) Tagawa 1940 (genus Sorolepidium H. Christ, Bot. Gaz. 51:360. 1911): type species P. duthiei (Hope) C.Chr. (Northwest India, Yunnan, Szechuan, and Taiwan in alpine regions) [Note that the type of the section is different from the type of the genus Sorolepidium.]
  • sect. Sphaenopolystichum Ching ex W. M. Chu et Z. R. He 2001: type P. omeiense C. Chr. (Southwest China).
  • sect. Stenopolystichum Daigobo 1972: type species P. stenophyllum Christ (North India, East Tibet, Yunnan, Szechuan, and Taiwan)
  • sect. Uncata Roux 1998: type species P. zambesiacum Schelpe (south-central Africa)
  • sect. Xiphopolystichum Daigobo, 1972: type species P. tsus-simense (Hook.) J. Sm. (Japan, Korea, Taiwan, and China)

  • Driscoll, H. E. and D. S. Barrington, 2007. Origin of Hawaiian Polystichum (Dryopteridaceae) in the context of a world phylogeny. American Journal of Botany 94: 1413–1424.
  • Fraser-Jenkins, C. R. 1997. Himalayan Ferns (A Guide to Polystichum). International Booksellers, Dehra Dun, India.
  • Li, C. X., S. G. Lu, and D. S. Barrington. 2008. Phylogeny of Chinese Polystichum (Dryopteridaceae) based on chloroplast DNA sequence data (trnL-F and rps4-trnS). Journal of Plant Research. 121: 19-26.
  • Liu, H.M., X.C. Zhang, W.Wang, Y.L. Qiu, and Z.D. Chen. 2007. Molecular phylogeny of the fern family Dryopteridaceae inferred from chloroplast rbcL and atpB genes. Int. J. Plant Sci. 168: 1311-1323.
  • Schuettpelz, E. and K.M. Pryer, 2007. Fern phylogeny inferred from 400 leptosporangiate species and three plastid genes. Taxon 56: 1037-1050.
  • Smith, A.R., K.M. Pryer, E. Schuettpelz, P. Korall, H. Schneider, and P.G. wolf. 2006. A classification for extant ferns. Taxon 55: 705-731.

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