Rhizome long-creeping, 2-4 mm diam., covered in red-brown hairs up to 2.5 mm long, stipes arising at intervals of 9-20 cm (abundant near growing tip, sparse elsewhere). Stipes 8-60 cm long, 1.5- 5 mm diam., red-brown at very base, becoming pale red-brown or yellow-brown above, covered at base in red-brown hairs, up to 3 mm long, becoming sparse above and paler. Laminae ovate or broadly ovate, 20-120 × 15-100 cm, bipinnate at apex, tripinnate or quadripinnate below. Rachis red-brown or yellow-brown at base becoming green at apex, sparsely covered in pale brown hairs like those of stipe. 1ry pinnae in 22-30 pairs, opposite or subopposite, the lowest arising at 20-50° to stem, upper ones at 70-90°, the longest near the base 15- 55 × 8.5-27 cm; lowest ones 5-24 cm apart, middle ones 1.5-8 cm apart; upper ones narrowly ovate or + parallel-sided, lower ones ovate or broadly ovate, narrowly winged. 2ry pinnae narrowly ovate or ± parallel-sided, broadly winged, 5-16 × 2-5.3 cm, those on the lower pinnae decreasing markedly in length along the pinnae. 3ry pinnae narrowly ovate or parallel-sided, 1-3 × 0.5-1.2 cm. 4ry pinnae (when present) 0.25-0.8 × 0.15-0.4 cm. Veins reaching margin at tooth apex. Hairs: glistening colourless non-glandular hairs on underside of lamina, costae and midribs, absent from lamina margin; brown-tinged hairs on upper surfaces, abundant on midribs; 0.3-1 mm long (mostly c. 0.5 mm), those on pinna midribs and costae often longer. Sori: one on acroscopic edge of each ultimate segment, originating away from margin; protected at maturity by slightly reflexed pinnule margin which also often bears a few short hairs at this point; hairs also often present in sorus (0.3- 0.5 mm long). Spores very pale under light microscope, perispores of inter-connecting flattened projections, (32)36-41(44) × (21)22-28(31) mcm (12 populations). See Fig. 1C, 2C, 4B, 13, and 18E.

Terrestrial ferns; rhizomes long-creeping, solenostelic; fronds often large, bipinnate or more compound; veins free; scales absent; hairs present on at least some part of lamina, rachis, stipe or rhizome, often abundant, glandular or nonglandular, sometimes modified on stipe and rachis into recurved spines; sorus ± round, ranging in position from marginal and terminal on a vein to submarginal and not quite terminating the vein; indusium a reflexed laminal flap, sometimes welldeveloped protecting the marginal sorus, more usually only partially reflexed, occasionally totally lacking in species with submarginal sori; spores monolete; chromosome number n = 28, 29, (39?), 52, 98, 104 (Brownsey 1983).

Rhizome long-creeping, (3)4-8 mm diam., densely covered in pale brown hairs near growing apex, more scattered and becoming red-brown elsewhere, giving rise to stipes at intervals of 10-15 cm. Stipes (15)20-100 cm long, (2)5-15 mm diam., redbrown at base, becoming pale red-brown or yellowbrown above, bearing red-brown hairs at very base, soon replaced by colourless glandular and nonglandular hairs (up to 5 mm long on uncoiling fronds); two dark, prominent, vertical bands on opposite sides of stipe. Laminae broadly ovate or elliptic, (20)35-135 × (15)35-110 cm, bipinnate at apex, 4 or 5-pinnate at base. Rachis yellow-brown at base, green at apex, bearing colourless glandular and non-glandular hairs (up to 3 mm long). 1ry pinnae in 15-30 pairs, opposite or sub-opposite, lower pair arising at 30-50° to stem; longest pair below middle, usually basal, (11)20-70 × (7)14-45 cm; lowest ones (5) 10-40 cm apart, middle ones 2-5 cm apart; upper ones narrowly ovate, lower ones ovate. 2ry pinnae ovate, (4.5)7-30 × 3-15 cm, those on the lower pinnae decreasing markedly in length along the pinnae. 3ry pinnae ovate, (1.6)2-8 × (0.7)1- 3.5 cm, midrib winged. 4ry pinnae narrowly ovate, 0.7-1.6 × 0.2-0.6 cm, shallowly incised on smaller specimens, divided into 4-5 pairs of ultimate segments on larger specimens. Veins reaching margin at a tooth apex, or sometimes ending just short of margin. Hairs: colourless glandular and non-glandular hairs on midribs and veins of both lamina surfaces but absent from margins, 0.3-1.5 mm long. Sori on upper margin of each ultimate segment, situated at vein endings, protected by obvious reflexed flaps (green at base, membranous at apex, incised); protected from earliest stages. Spores very pale under light microscope, perispores of interconnecting flattened projections, (35)38-39(42) × (21)22-25(29) mcm (5 populations). See Fig. 1D, 2D, 4C, 16, and 18G.

Rhizome long-creeping, 1-2.25 mm diam., abundantly covered in dark red-brown hairs 2- 3.5 mm long, giving rise to stipes at intervals of 3- 7 cm. Stipes 5-50(60) cm long, 0.75-1.25 mm diam., dark red-brown, bearing a few scattered redbrown hairs at base. Laminae narrowly elliptic to elliptic or ovate, (15)25-60(95) × 9-30(40) cm, rather stiff and harsh to the touch, bipinnate at apex to almost tripinnate at base (truly tripinnate in largest specimens). Rachis dark red-brown becoming paler only at the very apex, bearing scattered pale brown hairs. lry pinnae in 20-40 pairs (more in very large specimens), opposite or subopposite, arising at c. 90 to rachis, the longest at or below middle 5-20 × 1.7-6 cm; lowest pinnae (2)4- 12(20) cm apart, middle pinnae (1)2-5(6) cm apart, narrowly ovate or triangular, midrib narrowly winged throughout; lowest pair orientated at 90° to plane of frond in largest specimens. 2ry pinnae sessile or shortly stalked, ovate to narrowly ovate, longest 0.8-3(4) × 0.4-0.7(1) cm, doubly serrate with 3-9 primary serrations on each side of longest 2ry pinna (divided to the midrib in tripinnate forms); those on lower 1ry pinnae ± equal in length, only decreasing in size towards the very apex of each pinna. Veins reaching margin at a sinus (emarginate). Hairs virtually absent on upper surface of frond, a few scattered pale-brown non-glandular hairs on underside of costae and midribs. Sori in 2-8 pairs on longest 2ry pinnae, marginal, terminating veins on sides of ultimate segments, protected by reflexed, incised membranous flaps. Spores appearing dark brown under light microscope with reticulate perispore lacking obvious projections (28)31-34(40) × (18)19-21(24) urn (6 populations). See Fig. 2B, 3A, 5, and 18A.

Rhizome long-creeping, 1-2 mm diam., covered in red-brown hairs 2-3.5 mm long, giving rise to stipes at intervals of 1.5-9 cm. Stipes 10-45 cm long, 1.5-3.5 mm diam., dark purple-red to redbrown, bearing red-brown hairs up to 3 mm long, numerous at base, more scattered above, becoming replaced by abundant colourless glandular hairs which are very variable in length but generally shorter than the brown hairs. Laminae angularovate, 20-60(90) × 12.5-35(50) cm, pinnate at apex, tripinnate at base (almost quadripinnate in largest specimens), exuding white milky substance from upper surface with age. Rachis red-brown at base, becoming pale yellow brown at apex, densely covered in short glandular hairs (0.1-0.8 mm) and more scattered longer red-brown hairs (up to 2 mm long, but mostly c. 0.5 mm). lry pinnae in 20-35 pairs ± tapering pinnatifid terminal portion, opposite or subopposite, lower arising at 35-80°, upper ones at nearly 90°, longest ones below the middle and usually basal 6.5-38 × 4-15 cm; lowest ones 3-15(20) cm apart, middle ones 1-5 cm apart; upper ones narrowly ovate or triangular to almost linear, lowest ovate or triangular, all with long tapering pinnatifid terminal portion. 2ry pinnae narrowly ovate, longest 2.4-9.0 × 0.9-3.0 cm, those on the lower pinnae decreasing markedly in length along the pinnae. 3ry pinnae ovate, 0.5-1.6 × 0.3-0.75 cm, divided into 2-4 pairs of ultimate segments, broadly winged. Veins reaching margin at a tooth apex, often very slightly excurrent. Hairs: lamina margins, undersurfaces, veins, and midribs densely covered in short colourless glandular hairs (0.1- 0.3 mm long on laminae, up to 0.4 mm on midribs), interspersed with occasional non-glandular hairs of same length; equal mixture of short colourless glandular and non-glandular hairs on upper surface; occasional longer red-brown non-glandular hairs on midribs. Sori: usually one on acroscopic edge of each ultimate segment, sometimes second on basiscopic edge, originating away from margin, virtually unprotected. Spores very pale under light microscope, perispores of interconnecting flattened projections, (25)28-35(39) × (17)18-21(23) mcm (8 populations). See Fig. IB, 2A, 3D, 7B, and 18B.

Rhizoma repens, 1 -2 mm diametro, pilis rufescentibus 2-3.5 mm longis, stipites 1.5-9 cm distantes efferens. Stipes 10-45 cm longus, 1.5-3.5 mm diametro, atropurpureus vel rufescens, pilis rufescentibus usque ad 3 mm longos basi multis supra sparsis et pilis multis brevioribus glanduliferis hyalinis interspersis. Lamina angulato — ovata, 20-60(90) × 12.5-35(50) cm, apice pinnata, basi tripinnata (grandissimae laminae fere quadripinnatae), humorem lacteam pagina superiore maturitate exsudans. Rhachis basi rufescens, apice fulva, pilis multis brevibus glanduliferis (0.1-0.8 mm) et pilis sparsis rufescentibus longioribus usque ad 2 mm longos (plerumque c. 0.5 mm). 1ry pinnae 20-35-jugatae cum apice pinnatifido acuminato, oppositae vel suboppositae, inferiores ad angulum 35-80° patentes, superiores fere 90° patentes; longissimae infra medium et plerumque basi repertae, 6.5-38 × 4- 15 cm; infimae 3—15(20) cm distantes, medianae 1-5 cm distantes; superiores anguste ovatae vel triangulatae vel fere lineares, infimae ovatae vel triangulatae, omnes apicibus longis pinnatifidis acuminatis. 2ry pinnae angulare ovatae, longissimae 2.4-9.0 × 0.9-3.0 cm, inferiorum lrpinnarum apicem versus longe valde decrescentes. 3ry pinnae ovatae, 0.5-1.6 × 0.3-0.75 cm, in 2-4 segmenta binata divisae, late alatae. Venae marginem in apicibus dentium attingentes, saepe parum excurrentes. Margines, paginae inferiores costaeque pinnarum pilis multis brevibus hyalinis glanduliferis, pilis sparsis setosis interspersis, dense vestiti (0.1-0.3 mm longis in laminis, usque ad 0.4 mm in costis); paginae superiores pilis brevibus hyalinis glanduliferis setosisque pariter vestitae; costae pilis sparsis setosis rufescentibus longioribus vestitae. Sori 1-2 in pinnulis ultimis, procul a margine exorientes, reflexis pinnularutn marginibus fere non obtecti. Sporae pallidae, perisporae projecturis complanatis interconnexis, (25)28-35(39) × (17)18-21(23)mcm.

Rhizome subterranean, long-creeping, 1.5-3 mm diam., glabrous or bearing scattered pale brown hairs (especially near growing point), stipes arising at intervals of 1.5-13 cm. Stipes 5-35 cm long,1.5- 3 mm diam., red-brown below, pale brown above, a few pale or reddish brown hairs along stipe. Laminae rhombic, ovate or broadly ovate, 15-40(70) × 8-25 cm, membranous to slightly coriaceous, normally tripinnate (bipinnate in smallest specimens) at apex to quadripinnate or sometimes almost 5- pinnate at base. Rachis pale brown to yellowish green, bearing pale brown or colourless non-glandular hairs c. 1 mm long. 1ry pinnae in 20-25 pairs, the lowest arising at 30-50° to stem, upper ones at 70-90°, opposite below, subopposite or alternate above, the longest below middle 7-26 × 3.5-13 cm; lowest ones 3.5-15 cm apart, middle ones 1.5-5 cm apart; upper ones narrowly ovate, lower ones ovate, midrib narrowly winged towards apices. 2ry pinnae narrowly ovate to ovate, midribs broadly winged, 2.5-8 × 1-3.5 cm, those on the lower pinnae decreasing in length along the pinnae. 3ry pinnae 0.5-2 × 0.3-1 cm, 2-6 deep serrations per pinnule. 4ry pinnae 0.1-0.5 × 0.1-0.4 mm, sometimes deeply divided. Hairs: glistening colourless hairs scattered along midribs of pinnae and pinnules, on both surfaces, up to 1 mm long. Veins reaching margin at tooth apex. Sori: one on each ultimate segment, though often absent from lowest two pairs of pinnae, originating away from margin, unprotected at maturity or partially covered by a green reflexed marginal flap. Spores very pale under light microscope, perispores of inter-connecting flattened projections, (28)31-34(38) × (18)20-22(26) mcm (5 populations). See Fig. 3B, 11, and 18D.

Rhizome long-creeping, 1-2 mm diam., covered in dark red-brown hairs up to 4 mm long, giving rise to stipes at intervals of 2-8 cm. Stipes 4.5- 30 cm long, 0.7-2.5(3) mm diam., dark purple-red to red-brown, bearing abundant red-brown nonglandular hairs (to 3 mm long) and shorter redbrown glandular hairs. Laminae narrowly ovate to ovate, (6)15-55(65) × (2)5-25 cm, pinnate or bipinnate at apex, tripinnate at base (occasionally bipinnate in very small specimens). Rachis red-brown at base becoming pale yellow-brown only near apex, densely covered in red-brown hairs of various lengths with some widely-spaced non-glandular hairs up to 3 mm long but the majority shorter (< 1 mm) and either glandular or non-glandular. 1ry pinnae in 17-25 pairs + pinnatifid apex, opposite or subopposite, the lower arising at (30)40- 70(90)°, upper ones at nearly 90°, longest ones below middle 2.5-19 cm; lowest pairs 3.5-12 cm apart, middle ones 1-5 cm; upper ones parallel sided or narrowly ovate, lower ovate or triangular. 2ry pinnae ovate or narrowly ovate, longest 0.8-4 × 0.4- 1.7 cm, those on the lower pinnae decreasing markedly in length along the pinnae. 3ry pinnae 0.3-0.8 × 0.15-0.5 cm, slightly incised, bearing 1-3 pairs sori (deeply incised in very large specimens). Veins reaching the margin at a tooth apex. Hairs: brown bristly hairs (mostly 0.25-0.5 mm) abundant on margins and both surfaces (especially upper) of lamina (often rather pale in young specimens), interspersed with pale glandular and non-glandular hairs on lamina undersurface (few also on top surface); pinna midribs similar to rachis with longer red-brown bristly hairs and some red-brown short glandular and non-glandular hairs. Sori in pairs on ultimate pinnules, originating away from margin, unprotected. Spores very pale under light microscope, perispores of interconnecting flattened projections, (25)28-35(39) × (17)18-21(23) urn (5 populations). See Fig. 1A, 3C, 7A, and 18F.

Rhizome long-creeping, 2-2.5 cm diam., covered in red-brown hairs. Stipes 7-23(45) cm long, 1- 3.5 mm diam., dark red-brown, bearing abundant colourless or pale-brown glandular hairs up to 2 mm long. Laminae narrowly ovate to ovate, 15-55 × 10-35 cm, bipinnate at apex, tripinnate at base. Rachis dark red-brown at base becoming paler above and often green at apex, densely covered in colourless or pale-brown glandular hairs up to 1.5 mm long. 1ry pinnae in 15-30 pairs + pinnatifid apex, opposite or alternate, lower arising at (20)30- 70°, upper ones at 70-90°, longest ones below the middle 5-20 × 2-9 cm; lowest pairs 3-10 cm apart, middle ones 1.5-5 cm apart; upper ones parallelsided or narrowly ovate, lower ones ovate or narrowly ovate. 2ry pinnae ovate to ± parallel-sided, longest 0.5-4.5 × 1-2 cm, those on the lower pinnae decreasing markedly in length along the pinnae. 3ry pinnae up to 0.6 × 1.0 cm, deeply incised on larger specimens, bearing 1-3 pairs of sori. Veins reaching margin at a tooth apex. Hairs: colourless glandular hairs (0.3-0.7 mm on laminae, up to 1 mm on midribs) interspersed with a few bristly colourless hairs densely covering both surfaces of lamina, lamina margins, and pinna midribs and costae. Sori in 1-3 pairs on ultimate pinnules, originating away from margin, slightly protected by reflexed margins of pinnules. Spores very pale under light microscope, perispores of inter-connecting flattened projections, (28)31-34(36) × (18)20- 22(25) mcm (5 populations). See Fig. 4A, 10, and 18C.

Rhizoma repens, 2-2.5 cm diametro, pilis rufescentibus. Stipes 7-23(45) cm longus, 1-3.5 mm diametro, rufescens, pilis multis glanduliferis hyalinis vel ferrugineis usque ad 2 mm longos. Lamina anguste ovata vel ovata, 15—55 × 10-35 cm, apice bipinnata, basi tripinnata. Rhachis basi rufescens, in medio ferruginea, apice viridis, pilis multis glanduliferis hyalinis vel ferrugineis usque ad 1.5 mm longos. 1ry pinnae 15-30-jugatae cum apice pinnatifido, oppositae vel alternae, inferiores ad angulum (20)30-70°patentes, superiores ad angulum 70- 90° patentes, longissimae infra medium 5-20 × 2-9 cm; infimae 3—10 cm distantes, medianae 1.5—5 cm distantes; superiores lineares vel anguste ovatae, inferiores ovatae vel anguste ovatae. 2ry pinnae ovatae vel ± lineares, longissimae 0.5-4.5 × 1-2 cm, inferiorum 1ry pinnarum apicem versus longe valde decrescentes. 3ry pinnae usque ad 0.6 × 1.0 cm, profunde incisae, soris 1-3-jugatis. Venae marginem attingentes, in apicibus dentium terminantes. Margines, paginae costaeque pinnarum pilis multis glanduliferis hyalinis (0.3-0.7 mm longis in laminis, usque ad 1 mm in costis), pilis paucis setosis hyalinis interspersis, dense vestiti. Sori 1-3-jugati, in pinnulis ultimis, procul a margine exorientes, retlexis pinnularum marginibus partim obtecti. Sporae pallidae, perisporae projecturis complanatis interconnexis, (28)31-34(36) × (18)20- 22(25) mcm.