Classification
 Nomenclature
Scientific Name:
Pteris tremula R.Br., Prodr. Fl. Nov. Holland. 154 (1810)
Synonymy:
Lectotype (selected by Green 1994): Port Jackson [Sydney, Australia], R. Brown Iter Austral. 4, 1802-5, BM 001044256!
  • = Pteris affinis A.Rich., Essai Fl. Nouv.-Zél., 81 (1832)
Lectotype (selected by Brownsey & Perrie 2020): Nouvelle Zélande [New Zealand], Baie des Iles [Bay of Islands], Astrolabe, Herb. Richard, P 01319309!
  • = Pteris tenuis A.Cunn., Companion Bot. Mag. 2: 365 (1837)
  • Pteris tremula var. tenuis (A.Cunn.) Domin, Biblioth. Bot. 20(85): 156 (1913)
Holotype: Vicinity of Wangaroa [Whangaroa], A. Cunningham, Nov. 1826, K 001057870!
Etymology:
From the Latin tremulus (shaking), a reference to the trembling appearance of the frond in a light breeze.
Vernacular Name(s):
shaking brake; tender brake; turawera
 Description

Rhizomes erect, scaly. Rhizome scales narrowly ovate, 2.5–9 mm long, 0.4–1 mm wide, golden brown, concolorous. Fronds 220–2440 mm long. Stipes 120–1170 mm long, red-brown proximally, yellow-brown or chestnut-brown distally, glabrous. Rachises green when young, darkening to chestnut-brown or yellow-brown, narrowly winged only in distal half, glabrous. Laminae deeply 2-pinnate-pinnatifid to deeply 3-pinnate-pinnatifid, 100–1270 mm long, 80–900 mm wide, ovate, mid-green adaxially, yellow-green abaxially, herbaceous or coriaceous, glabrous or with minute hairs along costae. Primary pinnae in 5–18 pairs below pinnatifid apex, not or slightly overlapping; the longest at or near base, 60–620 mm long, 42–240 mm wide, broadly ovate or ovate or narrowly elliptic, straight; pinna apices acute or acuminate, bases stalked. Longest secondary pinnae 23–215 mm long, 10–120 mm wide, ovate, sub-opposite; apices acute, bases short-stalked; basiscopic secondary pinnae on basal primary pinnae elongated. Longest tertiary segments 5–70 mm long, 2–35 mm wide, ovate to linear; apices acute to obtuse, margins entire, bases adnate or short-stalked; sometimes divided into quaternary segments 4–20 mm long, 1–4 mm wide, linear to oblong; apices obtuse, margins entire or shallowly serrate, bases adnate. Veins free. Sori continuous along sides of the ultimate segments.

 Recognition

Pteris tremula is distinguished by its erect rhizome, 2-pinnate-pinnatifid to almost 4-pinnate laminae, free veins, herbaceous texture, long narrow ultimate segments, and laminae lacking scales but sometimes with minute hairs along the costae. It is more divided, with more pairs of primary pinnae, than most other species in New Zealand. It could be confused with P. epaleata in overall morphology but differs significantly in having free veins rather than anastomosing venation.

 Distribution

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

South Island: Western Nelson, Sounds-Nelson, Marlborough, Westland, Canterbury.

Kermadec Islands, Three Kings Islands, Chatham Islands.

Altitudinal range: 0–550 m.

Pteris tremula occurs on the Kermadec Islands, Three Kings Islands, and in coastal and lowland areas of the North Island from Te Paki to Wellington, extending locally into montane districts. It is absent only from parts of the central North Island and Gisborne high country. It grows from near sea level, reaching 550 m in the Kāweka Ranges. In the South Island it occurs from north-west Nelson to the Marlborough Sounds, south to Banks Peninsula on the east coast, and to Punakaiki on the west coast, with an outlying population at the Paringa River mouth in south Westland. It reaches almost 250 m on Mt Burnett. It also occurs on the Chatham Islands.

Also Australia (Northern Territory, South Australia, Queensland, New South Wales, Australian Capital Territory, Victoria, Tasmania), Norfolk Island, Lord Howe Island, Vanuatu and Fiji. Widely cultivated internationally; naturalised in southern Africa and California (Crouch et al. 2011), parts of the Northern Hemisphere (Zhang et al. 2015), and in Tamil Nadu State, India (Fraser-Jenkins et al. 2017).

 Habitat

Pteris tremula is found under coastal forest and scrub, open broadleaved and beech forest, kānuka and mānuka scrub, and in open pine plantations. It also occurs on forest margins, under regenerating scrub, in reverting pasture, on wooded sand-dunes, at the base of cliffs, and as a weed in gardens. It favours streambanks, the base of waterfalls, river terraces, bush clearings, open forest floor, roadside banks, rocky hillsides, and swamp margins. It occurs on greywacke, limestone and scoria. It is sometimes common in heavily grazed areas because the fronds are unpalatable to stock.

 Biostatus
Indigenous (Non-endemic)
 Hybridisation

Pteris tremula is not known to hybridise with any other species in New Zealand.

 Cytology

n = c. 120 (Brownlie 1957).

 Notes

Pteris novae-zelandiae was described by Field (1906) from material received in a “parcel of ferns from Waikanae”. He compared it with P. tremula but noted that it differed most obviously in its growth form – spreading horizontally by the production of lateral fronds, rather than being erect. No original material has been located and it is impossible to determine the identity of the plant from the description alone. The name was not documented by either Cheeseman (1925) or Allan (1961) and, in any case, was not validly published because Field stated only that “it may be provisionally named Pteris novae-zelandiae”. Provisional names are not considered to be validly published (see Art. 36.1, Ex. 5).

 Bibliography
Brown, R. 1810: Prodromus Florae Novae Hollandiae et Insulae Van-Diemen. Johnson, London.
Brownlie, G. 1957: Cyto-taxonomic studies on New Zealand Pteridaceae. New Phytologist 56: 207–209.
Brownsey, P.J.; Perrie, L.R. 2020: Taxonomic notes on the New Zealand flora: lectotypes in Pteridaceae. New Zealand Journal of Botany 58(3): 245–254.
Brownsey, P.J.; Perrie, L.R. 2021: Pteridaceae. In: Breitwieser, I. (ed.) Flora of New Zealand — Ferns and Lycophytes. Fascicle 30. Manaaki Whenua Press, Lincoln.
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.
Crouch, N.R.; Klopper, R.R.; Burrows, J.E.; Burrows, S.M. 2011: Ferns of southern Africa. A comprehensive guide. Struik Nature, Cape Town.
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.
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Domin, K. 1913: Beiträge zur Flora und Pflanzengeographie Australiens. Bibliotheca Botanica 20(85): 1–239.
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Fraser-Jenkins, C.R.; Gandhi, K.N.; Kholia, B.S.; Benniamin, A. 2017: An annotated checklist of Indian Pteridophytes. Part 1. Lycopodiaceae to Thelypteridaceae. Bishen Singh Mahendra Pal Singh, Dehra Dun, India.
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