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Brownsey, P.J. 1977: A taxonomic revision of the New Zealand species of Asplenium. New Zealand Journal of Botany 15(1): 39–86.

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Brownsey, P.J. 1977: A taxonomic revision of the New Zealand species of Asplenium. New Zealand Journal of Botany 15(1): 39–86.
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Rhizome short, stout, erect, bearing ovate scales up to 15 × 5 mm. Stipes 5-30 cm long, brown on underside, green above, stout, covered in small brown ovate scales. Laminae lanceolate to elliptic, 15-120 × 7-30 cm, bi- to tripinnate, sometimes bearing bulbils. Raches green, scaly, prominently grooved. Pinnae 15—30 (or more) pairs, ovate to narrowly ovate, acuminate, shortly stalked, 3-20 × 1-5 cm, scaly on underside, basal pair pointing downwards when fresh. Secondary pinnae sessile or shortly stalked, very narrowly elliptic to ovate or elliptic, obtuse, deeply serrate or sometimes almost pinnate, decreasing in size from base to apex, basal acroscopic pinnule often enlarged (up to 4 × 1 cm). Ultimate pinnules narrowly oblong, ± entire to crenate-serrate, up to 1 cm long. Sori numerous, broad, submarginal, 2-4 mm long. Perispore of spores irregularly ridged (Fig. 30K-N).
Very similar to subsp. bulbijerum but differing in the following characters. Stipe scales with apices drawn into long filiform projections (Fig. 17K-O). Laminae dark green, rarely bulbiferous. Spores (33)41-48(55) mcm long, (21)28-32(37) mcm wide (3 populations).
Rhizome short, erect, bearing dark brown subulate scales with long filiform apices up to 6 × 0.5 mm (Fig. 4M-S). Stipes 1-10 cm long, green, slender, lacking scales. Laminae linear, 7-25 × 1-2.5 cm, light green, flaccid and often prostrate, pinnate. Raches green, weak, lacking scales, frequently extending beyond the uppermost pinnae and rooting at their apices. Pinnae 8-25 pairs, gradually decreasing in size from base to apex, fanshaped, crenate-dentate, cuneate at base, lowest 5-15 × 5-20 mm. Sod radiating along veins, up to 6 mm long. Spores (37)43-51(62) mcm long, (30)36-41(51) mcm wide (8 populations), perispore with numerous crests and ridges (Fig. 29C-D).
Rhizome short, stout, erect, bearing dark brown subulate or ovate scales up to 20 × 2 mm. Stipes 5-20 cm (or more) long, brown on underside, green above, flaccid or firm and erect, sparingly covered in small subulate or ovate scales. Laminae lanceolate to elliptic, 10-90 (or more) × 4-25 cm, dull green, thick, leathery and either limp and pendulous or stiff and erect, pinnate to bipinnate. Raches green, sparingly scaly. Pinnae in 5-20 (or more) pairs, linear to narrowly ovate, acuminate, long stalked, 2-15 × 0.5-2 cm; degree of dissection very variable, sometimes only divided into very short obtuse segments, sometimes pinnate with linear acute pinnules up to 15 × 2 mm. Sori submarginal, linear, 2-10 mm long.
Rhizome bearing subulate scales. Stipes 5-20 cm (or more) long, flaccid, bearing small subulate scales with long filiform apices (Fig. 24A-E). Laminae lanceolate to elliptic, 15-90 (or more) × 5-25 cm, leathery, limp and pendulous. Pinnae 5-20 (or more) pairs, linear, 5-15 × 0.5-2 cm. Pinnules very variable in length, from oblong and obtuse to linear and acute, up to 15 × 2 mm. Basal acroscopic pinnule occasionally much longer than that next to it. Sori 2-10mm long. Spores (31)36-44(50) mcim long, (19)23-27(33) mcm wide (10 populations), perispore with a prominent wing but lacking ridges (Fig. 29M-N). Plant usually epiphytic, but sometimes terrestrial.
Rhizome bearing ovate scales with very thick cell walls. Stipes 5-12 cm long, erect, firm, bearing narrowly to broadly ovate scales with very thick cell walls (Fig. 24F-I). Laminae oblong to elliptic, 10-30 × 4-15 cm, stiff, erect. Pinnae 5-12 pairs, linear to narrowly ovate, 20-80 × 8-16 mm. Pinnules oblong and obtuse to linear and acute, up to 7 × 2 mm. Basal acroscopic pinnule frequently much longer than that next to it, up to 4 cm long, and itself pinnatifid. Sori 2-7 mm long. Spores (38)43-49(54) mcm long, (20)26-31(38) mcm wide (10 populations); perispore sometimes winged, usually lacking ridges (Fig. 29 O-P). Plant of coastal rocks.
Rhizoma paleis ovatis cum parietibus cellularum crassissimis vestitum. Stipes 5-12 cm longus, erectus, firmus, paleis anguste vel late ovatis cum parietibus cellularum crassissimis vestitus (Fig. 24F-I). Lamina oblonga vel elliptica, 10-30 × 4-15 cm rigida, erecta. Pinnae 5-12 jugatae, lineares vel anguste ovatae, 20-80 × 8-16 mm. Pinnulae oblongae obtusaeque vel lineares acutaeque, usque ad 7 × 2 mm. Pinnula basalis acroscopica multo longior quam pinnula vicina, usque ad 4 cm longa, pinnatifida. Sori 2-7 mm longi. Sporae (38)43-49(54) mcm longae, (20)26- 31(38) mcm latae; perisporae interdum alatae, plerumque non cristatae. Numerus somaticus chromosomatum 144; in meiosi paria 72 observantur. Habitat in rupibus orae.
Rhizome short, erect, bearing numerous brown, ovate, acuminate scales up to 10 × 2 mm. Stipes 2-10 cm long, pale brown below, green above, densely covered in small, subulate to narrowly ovate scales with filiform apices (Fig. 17T-Z). Laminae lanceolate to rhombic, 4—25 × 1-15 cm, dark green, thin, normally bipinnate but often pinnate when young and almost tripinnate in well-grown specimens. Raches green, slender, and very scaly. Pinnae 5-15 pairs, very narrowly ovate to ovate, obtuse to acuminate, long-stalked, 1-8 × 0.5-2 cm, basal pair pointing upwards when fresh. Pinnules stalked, 3-10 pairs, linear to suborbicular, crenate to deeply incised (or almost pinnate), 3-12 × 3-10 mm, tending to lie at 90° to plane of frond in well-grown specimens. Sori sub-marginal on narrow pinnules, remote from margins on broad segments, 1-3 mm long. Spores (25)31-37(42) mcm long, (19)23-27(33) mcm wide (9 populations), perispore very prominently winged and ridged (Fig. 30G-H).
Rhizome creeping, up to 15 cm long, pale green with a few scattered scales, stoloniferous. Stipes 6-20 cm long, pale green above, brown below, deeply grooved, covered in small, very dark, triangular to ovate scales which have thick cell walls (Fig. 17F-J). Laminae lanceolate to elliptic, 15-60 × 7-20 cm, light green, glossy above, thin, bipinnate. Raches green, scaly, prominently grooved. Pinnae 12-20 pairs, lanceolate to narrowly ovate, acuminate, stalked, 3-10 × 0.5-2.5 cm, scaly on underside; lower pinnae themselves pinnate, upper ones pinnatifid. Pinnules sessile or shortly stalked, elliptic, often deeply serrate, 10-20 × 5-15 mm. Sori 3-10 mm long, nearer mid-vein than margin. Sporangia orangebrown. Spores (30)38-42(50) mcm long, (20)24-29(35) mcm wide (4 populations), perispore very pale and not prominently patterned (Fig. 30E-F).
Rhizome stout, often forming a hard woody mass above ground, bearing pale brown, shiny, ovate, acuminate scales up to 30 × 7 mm. Stipes 8-20 cm long, dark brown, stout, densely covered in narrow scales with very long filiform apices (Fig. 9K-O). Laminae oblong to elliptic, 18-100 × 10-35 cm, dark green and glossy above, pinnate. Raches brown below, green above, stout, slightly ridged, scaly. Pinnae 4-15 pairs, lanceolate to narrowly oblong or ovate, acuminate, crenateserrate to ± entire, cuneate at base, 4-15 × 1-3 cm, frequently covered in very small hair-like scales on the underside. Sori up to 20 mm long, not reaching lamina edge. Spores (27)33-38(44) mcm long, (18)20- 24(26) mcm wide (5 populations), perispore a complex reticulate network (Fig. 29A-B).
Rhizome stout, ascending, bearing brown subulate to narrowly triangular scales up to 25 × 2 mm. Stipes 3-20 cm long, pale brown at base and on underside, green elsewhere, covered with scales similar to but smaller than those of the rhizome (Fig. 4F-L). Laminae lanceolate to elliptic, 4—40 × 2—20 cm, dark shiny green to dull grey-green, pinnate to bipinnate. Raches normally green, occasionally brown on the underside, often grooved, scaly. Pinnae 2-16 pairs, ovate to narrowly oblong, obtuse to acuminate, serrate to ± entire, stalked, 1-10 × 0.5-5 cm, often covered on both surfaces with tiny scales. Lowermost pinnae normally at least partially lobed or divided at the base, sometimes completely pinnate; pinnules stalked, lanceolate to elliptic, obtuse, serrate to entire, up to 3 × 1.5 cm. Sori up to 10 mm long, not reaching lamina edge. Spores (36)40-54(60) mcm long, (23)27-35(40) mcm wide (13 populations), perispore prominently winged and ridged (Fig. 29K-L).
Rhizome stout, often forming a hard woody mass above ground, bearing brown, shiny, ovate, acuminate scales up to 15 × 4 mm. Stipes 2-30 cm long, dark brown at base, green above, stout, covered in ovate or narrowly triangular scales with acute or acuminate apices. Laminae ovate to lanceolate, 4-40 × 2-20 cm, dull green, very thick and fleshy, pinnate. Raches green, stout, prominently grooved and scaly. Pinnae 4-18 pairs, narrowly ovate to oblong, obtuse to acute, crenate-serrate to ± entire, cuneate at base, 10-100 × 7-25 mm, scaly on the underside. Sori up to 10 mm long, not reaching to lamina edge. Perispore of spores slightly winged but with few crests or ridges (Fig. 29E-H).
Generally a smaller plant than subsp. obtusatum. Stipes 2-8 cm long, covered in narrowly triangular scales (Fig. 9F-J). Laminae lanceolate to ovate, 4-20 × 2-7 cm. Pinnae in 4-18 pairs, 10-40 × 7-12mm. Spores (39)45-52(60) mcm long, (25)29- 33(38) mcm wide (6 populations).
Planta plerumque parvior quam subsp. obtusatum. Stipites 2-8 cm longi, paleis anguste triangularibus vestiti (Fig. 9F-J). Laminae lanceolatae vel ovatae, 4-20 × 2-7 cm. Pinnae 4-18 jugatae, 10-40 × 7-12 mm. Sporae (39)45-52(60) mcm longae, (25)29- 33(38) mcm latae (Fig. 29G-H). Numerus somaticus chromosomatum 288; in meiosi paria 144 observantur (Fig. 31E).
Rhizome stout, short creeping, densely covered in red-brown, narrowly triangular scales up to 10 × lmm. Stipes 10-30 cm long, dark brown, still", densely covered in scales similar to but smaller than those of the rhizome (Fig. 4A-E). Laminae lanceolate, 25-50 (or more) × 10-20 cm, dark green and glossy above, paler and dull below, frequently pendulous, pinnate. Raches dark chocolate brown, very scaly. Pinnae 25 (or more) pairs, narrowly angular-ovate to ovate, sometimes with a large rounded basal acroscopic lobe, acuminate, doubly serrate, 5-10 × 1—2 cm, scaly and with prominent veins on underside. Sori often slightly curved away from the midrib, up to 2 mm long. Spores (26)30-37 (40) mcm long, (15)18-21(25) mcm wide (6 populations), perispore with a few very prominent ridges (Fig. 30A-B).
Rhizome short, stout, erect, bearing dark brown subulate scales up to 20 × 2 mm. Stipes 5—15 cm long, brown on underside, green above, densely covered in subulate scales with filiform apices (Fig. 17P-S). Laminae ovate to narrowly ovate, 10-25 × 4-12 cm, dark green, relatively thin, normally tripinnate. Raches green, very scaly, slightly grooved. Pinnae 10-15 crowded and overlapping pairs, ovate to narrowly ovate, sub-acute, stalked, 2-8 × 1-4 cm. Secondary pinnae stalked, ovate, 10-20 × 10-15 mm, again pinnate or pinnatifid. Ultimate segments linear, acute or sub-acute, up to 8 mm long. Pinnae and pinnules not flattened in one plane but spreading in three dimensions. Sori 2-4 mm long, submarginal. Spores (38)44-49(55) mcm long, (27)32-36(40) mcm wide (6 populations), perispore prominently winged and ridged (Fig. 30I-J).
Rhizome short, stout, erect, bearing red-brown, ovate, acuminate scales up to 25 × 4 mm. Stipes 15-50 cm long, green on upper surface, brown below, stout, covered in narrowly triangular scales with long filiform apices (Fig. 9P-R). Laminae lanceolate, 15-50 × 8-20 cm, dark green, thick and leathery, pinnate. Raches green, stout, prominently grooved and scaly. Pinnae 6-17 pairs, lanceolate, acuminate, often broadened and lobed towards cuneate base, deeply and regularly crenate-serrate, 5-10 × 1-2 cm, scaly on underside. Sori reaching the lamina margin in sinuses, not more than 10 mm long. Spores (38)48-53(61) mcm long, (24)28-33(42) mcm wide (5 populations), perispore slightly winged, but with few ridges (Fig. 29I-J).
Rhizome short, erect, bearing redbrown, linear attenuate scales. Stipes up to 15 cm long, green above, brown below, sparingly covered in small linear scales with long filiform apices (Fig. 24 O-P). Laminae oblong to elliptic, 15-90 × 10-25 cm, yellow-green, coriaceous, 3- to 4-pinnate. Raches green, almost lacking scales, prominently grooved. Pinnae ovate to narrowly ovate, acuminate, stalked, up to 12 × 5 cm. Secondary pinnae linear to lanceolate, up to 15 × 2 mm, often pinnatifid. Ultimate segments linear, subacute, slightly expanded in the region of the sori. Sori near tips of ultimate segments, solitary, broad, submarginal, c.2mm long. Spores (46)51-63(71) mcm long, (32)34-44(50) mcm wide (3 populations), perispore prominently winged, granular, but not conspicuously ridged (Fig. 30C-D).
Rhizome short, erect, bearing dark brown subulate scales up to 15 × 2 mm. Stipes 5-15 cm long, brown below, green above, covered in subulate to narrowly triangular scales with long filiform apices (Fig. 24J-N, Q-V). Laminae lanceolate to elliptic, 10-30 × 5-20 cm, dark green, leathery or very thick and fleshy, slightly drooping or stiff and upright, bi- to tripinnate. Raches green, scaly, prominently ridged. Pinnae 7-20 pairs, those at the base ovate, those above very narrowly ovate or elliptic, acute to acuminate, stalked, 3-10 × 0.5-3 cm, covered on underside in tiny scales. Upper pinnules linear, acute, entire and sessile; lower pinnules ovate to narrowly elliptic and pinnatifid or pinnate, up to 30 × 10 mm. Sori submarginal, 2-7 mm long. Spores (38)43-51(56) mcm long, (25)29-34(38) mcm wide (14 populations), perispore prominently winged and ridged (Fig. 30 O-R). 288 chromosomes in the sporophyte; 144 bivalents at meiosis.
Rhizoma breve, erectum, paleis fuscis subulatis usque ad 15 × 2 mm obtectum. Stipes 5—15 cm longus, infra fuscus, supra viridis, paleis subulatis vel anguste triangularibus cum apicibus longis filiformibus vestitus (Fig. 24J-N, Q-V). Lamina lanceolata vel elliptica, 10-30 × 5-20 cm, atroviridis, coriacea aut crassissima carnosaque, cernua aut rigida erectaque, bi- vel tri-pinnata. Rhachis viridis, squamata, manifeste canaliculata. Pinnae 7-20 jugatae, inferiores ovatae, superiores peranguste ovatae vel ellipticae, acutae vel acuminatae, 3-10 × 0.5-3 cm, paleis minutis infra vestitae. Pinnulae superiores lineares acutae integrae sessilesque; inferiores ovatae vel anguste ellipticae, pinnatifidae vel pinnatae, petiolatae, usque ad 30 X 10 mm. Sori submarginales, 2-7 mm longi. Sporae (38)43-51(56) mcm longae, (25)29-34(38) mcm latae, perisporae manifeste alatae cristataeque (Fig. 30 O-R). Numerus somaticus chromosomatum 288; in meiosi paria 144 observantur.
Laminae lanceolate to elliptic, 10-30 × 5-15 cm, thick and leathery, usually slightly drooping. Pinnae 8-20 pairs, those at the base ovate, those above very narrowly ovate or elliptic, often with long relatively undivided acuminate tips. Upper pinnules linear and entire, lower pinnules narrowly elliptic and pinnatifid to pinnate, up to 30 × 8 mm.
Laminae relatively short and broad, elliptic, 10-25 × 7-20 cm, very thick and fleshy, stiff and upright. Pinnae 7-15 pairs, those at the base ovate, those above narrowly ovate, acute. Upper pinnules linear and entire, lower pinnules ovate to elliptic and pinnatifid, up to 20 × 10 mm.
Lamina pro ratione brevis lataque, elliptica, 10-25 × 7-20 cm, crassissima, carnosa, rigida, erecta. Pinnae 7-15 jugatae, inferiores ovatae, superiores anguste ovatae, acutae. Pinnulae superiores lineares integraeque, inferiores ovatae vel ellipticae pinnatifidaeque, usque ad 20 × 10 mm. Numerus somaticus chromosomatum 288; in meiosi paria 144 observantur.
Rhizome stout, erect, bearing dark brown subulate scales up to 5 mm long. Stipes 1-10 cm long, dark brown, shining, stiff, lacking scales except at the very base. Laminae linear, 5-30 cm long, dark green, sub-coriaceous, pinnate. Raches dark brown, shining, stiff, lacking scales. Pinnae sub-sessile, 15-25 (or more) pairs, oblong to ± orbicular, crenateserrate to ± entire, 2-10 × 1-5 mm. Sori up to 3 mm long.

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KEY TO THE NEW ZEALAND SPECIES OF ASPLENlUM

1
Fronds simply pinnate
2
Fronds 2 — or more pinnate
11
2
Fronds < 3 cm wide
3
Fronds > 3 cm wide
6
3
Stipe and rachis scaly; usually < 8 pairs pinnae
4
Stipe and rachis without scales; usually > 8 pairs pinnae
5
4
Fronds thick and fleshy; pinnae with scales on underside only; plant of exposed sea cliffs 4.
Fronds not thick and fleshy; pinnae often with scales on both surfaces; plant of calcareous soils
5
Stipe and rachis green; fronds lax and rooting at tip
Stipe and rachis brown; fronds stiff and erect
6
Rachis dark chocolate brown throughout; pinnae irregularly and doubly serrate
Rachis green, at least on the upper surface (sometimes turning pale brown when dried); pinnae entire or regularly serrate
7
7
Pinnae linear, > 10 times as long as broad
Pinnae < 10 times as long as broad
8
8
Pinnae prominently serrate; sori extending right to margin between serrations
Pinnae entire or minutely serrate; sori not reaching margin
9
9
Lowest pinnae usually lobed or with at least one distinct pinnule; sometimes with scales on both surfaces; plant of calcareous soils
Lowest pinnae not lobed or incised, lacking scales on upper surface
10
10
Rachis and foliar scales with very long hair-like apices (Fig. 9K-O); pinnae firm but not thick and fleshy, glossy above, accuminate to ± acute
Rachis and foliar scales ovate to narrowly ovate with short hair-like apices (Fig. 9A-J); pinnae thick and fleshy, dull above, obtuse to ± acute
11
Fronds bearing bulbils
Fronds not bearing bulbils
12
12
Rhizome creeping; fronds thin, glossy above
Rhizome not creeping; fronds dull above
13
13
Sori remote from margin, 4-10 mm long
Sori submarginal, or, if remote from margin, < 4 mm long
14
14
Fronds 3-4 pinnate at base; ultimate segments long and narrow, but broadened in region of sorus; plant of Kermadecs
Fronds 2-pinnate at base, or, if 3-4 pinnate, ultimate segments not broadened in region of sorus
15
15
Fronds 3-4 pinnate at base; ultimate segments linear, mostly < 1 mm wide; pinnae and pinnules crowded and overlapping; stipe stout
Fronds 2-3 pinnate at base; not with all above combination of characters
16
16
Fronds thin; sori 2-4 mm long
17
Fronds thick and leathery or fleshy; sori 2-10 mm long
17
Fronds with < 15 pairs pinnae (if c. 15 pairs pinnae, pinnules with distinct but very slender stalks)
Fronds with > 15 pairs pinnae (if c. 15 pairs pinnae, pinnules sessile or on very short broad stalks)
18
Spores* pale yellow, surface granular but with few or no prominent ridges (Fig. 29 M-P)
19
Spores* dark brownish yellow, surface with many very prominent ridges (Fig. 30 O-R)
20
19
Fronds limp and flaccid; plant usually epiphytic
Fronds firm and erect; plant of exposed sea cliffs, confined to east coast regions north of lat. 38°
20
Fronds thick and fleshy; plant of exposed sea cliffs
Fronds leathery, but not fleshy; plant of forest floor or rocky inland areas

Reproduced with permission from New Zealand Journal of Botany and The Royal Society of New Zealand Te Apārangi.

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7791dd2d-73e8-43a3-89ee-6424f8dd0840
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Names_Plants
16 March 2004
18 September 2007
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