Classification
 Nomenclature
Scientific Name:
Microsorum pustulatum (G.Forst.) Copel., Gen. Fil. 196 (1947) – as Microsorium pustulatum
Synonymy:
  • Polypodium pustulatum G.Forst., Fl. Ins. Austr. 81 (1786)
  • Phymatosorus pustulatus (G.Forst.) Large, Braggins & P.S.Green, New Zealand J. Bot. 30: 207 (1992) – as Phymatosorus pustulatum
  • Zealandia pustulata (G.Forst.) Testo & A.R.Field, Syst. Bot. 44: 14 (2019)
Lectotype selected by Pichi Sermolli 1951: New Zealand, G. Forster, BM 000066256!
  • = Polypodium scandens Labill., Nov. Holl. Pl. 2, 91, t.240 (1807) nom. illeg., non Polypodium scandens G.Forst. 1786
  • Polypodium diversifolium Willd., Sp. Pl. 5(1), 166 (1810) nom. nov. pro Polypodium scandens Labill. 1807 (non Polypodium scandens G.Forst. 1786)
  • Microsorum diversifolium (Willd.) Copel., Univ. Calif. Publ. Bot. 16: 114 (1929) – as Microsorium diversifolium
  • Pleopeltis diversifolia (Willd.) Melvaine, Proc. Linn. Soc. New South Wales 61: 120 (1936)
  • Phymatodes diversifolium (Willd.) Pic.Serm., Webbia 8: 222 (1951)
  • Phymatosorus diversifolius (Willd.) Pic.Serm., Webbia 28: 459 (1973)
Lectotype (selected by Brownsey & Perrie 2012): Nova Hollandia, [Tasmania], J.J.H.de Labillardière, FI 004209 (image!)
  • = Polypodium billardierei R.Br., Prodr. Fl. Nov. Holland. 147 (1810)
  • Phymatodes billardierei (R.Br.) C.Presl, Tent. Pterid. 196 (1836)
  • Chrysopteris billardierei (R.Br.) Link, Fil. Spec. 123 (1841)
  • Drynaria billardierei (R.Br.) J.Sm., Companion Bot. Mag. New Ser. 2: 14 (1846)
  • Pleopeltis billardierei (R.Br.) T.Moore, Index Fil. Ixxviii (1857)
  • Polypodium scandens var. billardierei (R.Br.) F.Muell., Veg. Chatham Isl. 69 (1864)
Lectotype (selected by Tindale 1961): Port Dalrymple, [Tasmania], R. Brown Iter Austral. 11, Jan. 1804, BM 001038373!
Etymology:
From the Latin pustulatus (having pustules), a reference to the impressed sori forming bulges on the upper lamina surface.
Vernacular Name(s):
hound's tongue; kōwaowao; pāraharaha
 Description

Epiphytic and terrestrial; creeping or climbing fern.  Rhizomes long-creeping, 3–11 mm diameter, scaly.   Rhizome scales clathrate, ovate, 3–7 mm long, 1–2 mm wide, mostly appressed but with the tips sometimes squarrose, blackish-brown, entire.  Fronds 45–750 mm long.  Stipes 10–350 mm long, pale to chocolate brown, glabrous or with very scattered scales.  Laminae very varied, from undivided or variously lobed to deeply 1-pinnatifid; undivided laminae (fertile) narrowly elliptic, 35–220 mm long, 6–30 mm wide, or (sterile) longer and broader, up to 260 mm long and 70 mm wide; lobed or pinnatifid laminae (fertile) ovate to elliptic or broadly ovate to broadly elliptic, 50–425 mm long, 35–350 mm wide, or (sterile) up to 420 mm long and 300 mm wide; lamina extending 5–40 mm below the lowest lobe, bright glossy green, coriaceous, glabrous or with scattered scales on the costae.  Lamina lobes (fertile) in 1–15 pairs, 25–185 mm long, 4–30 mm wide, or (sterile) less numerous and wider, in up to 10 pairs and up to 60 mm wide; straight, acute to acuminate, margins entire or wavy, widest about the middle.  Veins reticulate, forming 2–3 series of areoles between costa and lobe margin; hydathodes conspicuous on adaxial surface of lamina.  Sori round or rarely elliptic, 2.5–5 mm wide, impressed into lamina forming low bulges on adaxial surface, in 1 row either side of costa, medial or closer to the lobe margin; paraphyses absent; exindusiate.

 Recognition

This is the commonest and most variable of the three species of Microsorum in New Zealand.  It can be distinguished by its thick rhizomes (> 4 mm diameter) with appressed blackish-brown scales, its very variable fronds (Brownsey & Smith-Dodsworth 2000), coriaceous and bright glossy green laminae, conspicuous veins and hydathodes on the upper lamina surface, and broad lamina lobes.

Occasional aberrant forms are found - either crested at the apices or with 2-pinnatifid laminae.

Only one subspecies is present in New Zealand.  Microsorum pustulatum subsp. howense (Tindale et P.S.Green) Bostock is recognised as endemic to Lord Howe Island (Green 1994).  It is distinguished by having rhizome scales which are abruptly acuminate and 1.5-3.3 mm wide, laminae which are usually divided to the rachis into adnate pinnae, and sori which are deeply sunken into the lamina and usually either submarginal or about one third of the way to the costa.

 Distribution

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

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

Kermadec Islands, Three Kings Islands, Chatham Islands, Solander Island, Stewart Island, Antipodes Islands, Auckland Islands.

Altitudinal range: 0 - 900 m.

Abundant throughout the North Island from near sea-level to c. 700 m; more common in coastal and lowland regions of the South Island, extending locally to c. 900 m but largely absent from inland high country regions and much of Otago.  

Also Australia (Queensland, NSW, ACT, Victoria, Tasmania), Norfolk Island.

 Habitat

Creeping and often covering extensive areas on the ground, growing over rocks or fallen logs, on cliffs and banks, or epiphytic on native and naturalised trees.  Occurs in coastal to montane forest, pine forest, scrub, scoria and open areas, usually in slightly drier habitats, extending into subalpine scrub in the southern South Island.

 Biostatus
Indigenous (Non-endemic)
Number of subspecific taxa in New Zealand within Microsorum pustulatum (G.Forst.) Copel.
CategoryNumber
Indigenous (Non-endemic)1
Total1
 Cytology

n = 37 (Brownlie 1954, as Microsorium diversifolium).

 Notes

This species has been widely known as Phymatosorus diversifolius (e.g. Brownsey & Smith-Dodsworth 2000), Microsorum diversifolium (e.g. Crookes 1963) or Phymatodes diversifolium (e.g. Allan 1961), combinations based on Polypodium diversifolium Willd. collected in Tasmania by Labillardière.  An earlier basionym, Polypodium pustulatum G.Forst., was for many years considered a "species dubia" because of the inadequate type specimen lacking a rhizome, which could not be distinguished as either diversifolium or novae-zealandiae (Pichi Sermolli 1951).  However, on the basis of spore morphology Large et al. (1992a) showed conclusively that the specimen is the same as the former, and that the name P. pustulatum should take priority.  Hence the species is now recognised as Microsorum pustulatum (Large et al. 1992a, 1992b).

Robert Brown’s Polypodium billardieri, and names based on his type from Port Dalrymple, are later synonyms.  The name Polypodium phymatodes L. used by Richard (1832) is a misidentification.

 Bibliography
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Brownlie, G. 1958: Chromosome numbers in New Zealand ferns. Transactions of the Royal Society of New Zealand 85: 213–216.
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