Scientific Name:
Pneumatopteris pennigera (G.Forst.) Holttum, Blumea 21: 305 (1973)
  • Polypodium pennigerum G.Forst., Fl. Ins. Austr. 82 (1786)
  • Aspidium pennigerum (G.Forst.) Sw., J. Bot. (Schrader) 1800(2): 34 (1801)
  • Nephrodium pennigerum (G.Forst.) C.Presl, Reliq. Haenk. 1, 35 (1825)
  • Polystichum pennigerum (G.Forst.) Gaudich. in Freycinet, Voy. Uranie, Bot. 328 (1828)
  • Lastrea pennigera (G.Forst.) C.Presl, Tent. Pterid. 76 (1836)
  • Goniopteris pennigera (G.Forst.) J.Sm., J. Bot. (Hooker) 4: 54 (1841)
  • Goniopteris forsteri T.Moore, Index Fil. 99 (1858) nom. nov. pro Aspidium pennigerum (G.Forst.) Sw. 1801
  • Dryopteris pennigera (G.Forst.) C.Chr., Index Filic. 283 (1905)
  • Cyclosorus pennigerus (G.Forst.) Ching, Bull. Fan Mem. Inst. Biol. 10: 247 (1941)
  • Thelypteris pennigera (G.Forst.) Allan, Fl. New Zealand 1, 51 (1961)
Lectotype (chosen by Nicolson & Fosberg 2003): no locality, Forster, UPS-T 24698 (n.v.)
  • = Phegopteris cunninghamii Mett., Fil. Hort. Bot. Lips. 84 (1856)
Holotype: Cult. Leipzig ex New Zealand (n.v., see Holttum 1977)
  • = Aspidium novae-zeelandiae Ettingsh., Denkschr. Kaiserl. Akad. Wiss., Wien. Math.-Naturwiss. Kl. 23: 103 (1864)
Lectotype (selected by Brownsey & Perrie 2016): Nova Zeelandia [New Zealand], Hügel s.n., s.d., W 0052715 (!online)
  • = Polypodium pennigerum var. giganteum Colenso, Trans. & Proc. New Zealand Inst. 14: 339 (1882)
Lectotype (selected by Brownsey & Perrie 2016): New Zealand, Manawatu River, W. Colenso s.n., s.d., AK 142518! (two pinnae only)
  • = Polypodium pennigerum var. hamiltonii Colenso, Trans. & Proc. New Zealand Inst. 14: 338 (1882)
  • Dryopteris pennigera var. hamiltoni (Colenso) Cheeseman, Man. New Zealand Fl., ed. 2, 36 (1925)
  • Cyclosorus pennigerus var. hamiltoni (Colenso) Crookes in Dobbie, New Zealand Ferns ed. 4, 262 (1951)
Lectotype (chosen by Allan 1961): Kereru, A. Hamilton, Herb. W. Colenso, WELT P003350! (isolectotypes? AK 142511, 221795!)
  • = Polypodium subsimilis Colenso, Trans. & Proc. New Zealand Inst. 20: 233 (1888)
Lectotype (chosen by Allan 1961): 70 mile Bush, Herb. W. Colenso, WELT P003349!
From the Latin pennigerus (with feathery leaves), a reference to the dissection of the fronds.
Vernacular Name(s):
feather fern; gully fern; pākauroharoha; piupiu

Rhizomes prostrate or erect, often forming short arborescent trunks up to 1 m tall, bearing scales at the apex. Rhizome scales ovate to narrowly ovate, 3–8 mm long, 1–4 mm wide, chestnut-brown, entire. Fronds 340–2070 mm long, rarely only 200 mm long at high elevations, arching upwards. Stipes 40–430, rarely up to 770 mm long, pale brown, scaly to almost glabrous. Laminae 1-pinnate to 2-pinnate-pinnatifid, narrowly elliptic or elliptic, gradually tapering to a pinnatifid apex, 270–1390, rarely up to 1500 mm long, 80–400 mm wide, rarely only 160 mm long and 50 mm wide at high elevations, mid-green on both surfaces, thin and herbaceous. Ovate or broadly ovate, pale brown scales on abaxial surface of costae; colourless or pale brown acicular hairs up to 0.2 mm long on both costa surfaces; colourless capitate hairs <0.1 mm long on abaxial surfaces of costae. Primary pinnae in 8–35 pairs below pinnatifid apex, widely spaced especially proximally, narrowly oblong to narrowly ovate; the longest at or near the middle, sessile or short-stalked, the longest 44–220 mm long, rarely only 27 mm long at high elevations, 11–34 mm wide; the basal pair greatly reduced, 10–70 mm long, auricled acroscopically. Primary pinnae divided ⅓ to ½, or rarely ⅔, to the midrib; ultimate segments (excluding proximal pair) 5–17 mm long, 3.5–7 mm wide, apices obtuse or rounded, margins entire or minutely serrate; the proximal pair of ultimate segments often longer than the others with the basal acroscopic segment occasionally divided to the costa and greatly extended and deeply lobed, up to 53 mm long and 24 mm wide. Basal veins in adjacent pinna segments joining, unbranched in each ultimate pinna segment. Sori round, in one row either side of midrib away from pinna margins; indusia absent.


Pneumatopteris pennigera is recognised by its erect rhizome, primary  pinnae divided ⅓ to ⅔ to the midrib with rounded segments, greatly reduced basal pinnae, veins in adjacent segments joining, indumentum on the abaxial surfaces comprising ovate scales and acicular and capitate hairs but lacking glands, and sori lacking indusia. The network of dark veins on the thin green fronds distinguishes Pneumatopteris pennigera from other common forest ferns.


North Island: Northland, Auckland, Volcanic Plateau, Gisborne, Taranaki, Southern North Island.

South Island. Western Nelson, Sounds-Nelson, Marlborough, Westland, Canterbury, Otago, Southland, Fiordland.

Three Kings Islands, Chatham Islands

Altitudinal range:  0–700 m.

Pneumatopteris pennigera occurs in lowland areas, extending locally to montane sites, throughout the North Island, growing from near sea level to about 700 m on Mt Maungatautari in the Waikato region. In the South Island it is largely confined to lowland areas in the northern half of the island, although it extends locally to Fiordland, Southland and the Otago Peninsula, and from near sea level to about 600 m on Banks Peninsula.  

Also Australia (Queensland, Victoria, Tasmania).


A terrestrial fern that occurs under kauri, podocarp, broadleaved and beech forest, under mānuka, kānuka and Salix spp., and in Carex secta swamps. It grows on the forest floor, in gullies, on banks, streamsides and alluvial terraces, usually in heavy shade and damp sites, but sometimes in more open areas.

Indigenous (Non-endemic)

n = 72 (Brownlie 1954, as Cyclosorus pennigerus).


Pneumatopteris pennigera is an atypical member of the genus in that it has ovate scales on the abaxial surface of the costae when young, the abaxial surface of the lamina is not pustular when dry, indusia are lacking, and there are no glands or hairs on the sporangia (Holttum 1977). Bostock (1998) noted that it (and P. costata) needed further study in relation to their generic placement.

Kunze (1850) evidently believed that Cunningham’s (1837) concept of Aspidium pennigerum differed from that of Swartz (1801) and he created a nomen novum for it – Aspidium cunninghamii Kunze, Linnaea 23: 225 (1850). However, neither Kunze (1850), nor Cunningham (1837), provided a description and the name is therefore a nomen nudum. In any case, Kunze’s name is illegitimate, being a later homonym of Aspidium cunninghamii Colenso (1843).

Bostock, P.D. 1998: Thelypteridaceae. In: Flora of Australia. Vol. 48. 327–358.
Brownlie, G. 1954: Introductory note to cyto-taxonomic studies of New Zealand ferns. Transactions of the Royal Society of New Zealand 82: 665–666.
Brownsey, P.J.; Perrie, L.R. 2016: Taxonomic notes on the New Zealand flora: lectotypes in the fern family Thelypteridaceae. New Zealand Journal of Botany 54(1): 87–91.
Brownsey, P.J.; Smith-Dodsworth, J.C. 2000: New Zealand ferns and allied plants. Edition 2. David Bateman, Auckland.
Cheeseman, T.F. 1925: Manual of the New Zealand Flora. Edition 2. Government Printer, Wellington.
Ching, R.C. 1941: New family and combinations of ferns. Bulletin of the Fan Memorial Institute of Biology 10: 235–256.
Christensen, C. 1905–1906: Index Filicum. Hagerup, Copenhagen.
Colenso, W. 1843: Description of some new ferns lately discovered in New Zealand. Tasmanian Journal of Natural Science 1: 375–379.
Colenso, W. 1882: A description of a few new plants from our New Zealand forests. Transactions and Proceedings of the New Zealand Institute 14: 329–341.
Colenso, W. 1888: On newly discovered and imperfectly known ferns of New Zealand, with critical observations. Transactions and Proceedings of the New Zealand Institute 20: 212–234.
Cunningham, A. 1837: Florae insularum Novae Zelandiae precursor; or a specimen of the botany of the islands of New Zealand. Companion to the Botanical Magazine 2: 222–233, 327–336, 358–378.
de Lange, P.J.; Rolfe, J.R.; Barkla J.W.; Courtney, S.P.; Champion, P.D.; Perrie, L.R.; Beadel, S.N.; Ford, K.A.; Breitwieser, I.; Schönberger, I.; Hindmarsh-Walls, R.; Heenan, P.B.; Ladley, K. 2018: Conservation status of New Zealand indigenous vascular plants, 2017. New Zealand Threat Classification Series. No. 22. [Not Threatened]
de Lange, P.J.; Rolfe, J.R.; Champion, P.D.; Courtney, S.P.; Heenan, P.B.; Barkla, J.W.; Cameron, E.K.; Norton, D.A.; Hitchmough, R.A. 2013: Conservation status of New Zealand indigenous vascular plants, 2012. New Zealand Threat Classification Series 3. Department of Conservation, Wellington. [Not Threatened]
Dobbie, H.B. 1951: New Zealand ferns. Edition 4. Whitcombe & Tombs, Auckland.
Ettingshausen, C.F. von 1864: Beiträge zur Kenntnis der Flächen-Skelete der Farrnkräuter. Denkschriften der Kaiserlichen Akademie der Wissenschaften, Wien. Mathematisch-Naturwissenschaftliche Klasse 23: 39–119.
Forster, J.G.A. 1786: Florulae Insularum Australium Prodromus. Dietrich, Göttingen.
Freycinet, H.L.C. de 1826–1830: Voyage autour du monde, enterpris par ordre du roi, exécuté sur les corvettes de S.M. l’Uranie et la Physicienne, pendant les anneés 1817, 1818, 1819 et 1820. Botanique. Pillet-ainé, Paris.
Holttum, R.E. 1973: Studies in the family Thelypteridaceae V. The genus Pneumatopteris Nakai. Blumea 21: 293–325.
Holttum, R.E. 1977: The family Thelypteridaceae in the Pacific and Australasia. Allertonia 1: 169–234.
Kunze, G. 1850: Index filicum. Gebauer, Halle.
Mettenius, G.H. 1856: Filices Horti Botanici Lipsiensis. Leopold Voss, Leipzig.
Moore, T. 1857–1862: Index Filicum. Pamplin, London.
Nicolson, D.H.; Fosberg, F.R. 2003: The Forsters and the Botany of the Second Cook Expedition (1772–1775). Regnum Vegetabile 139: 1–760.
Presl, C.B. 1825–1830: Reliquiae Haenkeanae, seu, descriptiones et icones plantarum, quas in America meridionali et boreali, in insulis Philippinis et Marianis collegit Thaddaeus Haenke. Vol. 1. Calve, Prague.
Presl, C.B. 1836: Tentamen Pteridographiae. Haase, Prague.
Smith, J. 1841: An arrangement and definition of the genera of ferns, with observations on the affinities of each genus. Journal of Botany (Hooker) 4: 38–70.
Swartz, O.P. 1801: Genera et species filicum ordine systematico redactarum. Journal für die Botanik (Schrader) 1800(2): 1–120.
Webb, C.J.; Edgar, E. 1999: Spelling New Zealand in the specific and infraspecific epithets of vascular plants. New Zealand Journal of Botany 37(1): 71–77. [as Aspidium novae-zeelandiae Ettingsh.]