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Lindsaea Dryander

Terrestrial, rupestral, scandent or ipiphytic ferns of small to medium size. Rhizome short- to long-creeping, protostelic with a solid fibrovascular bundle with internal phloem, (Lindsaeoid protostele), or open in some small epiphytes, bearing basally attached, concolourous, narrow, elongate to acicular scales, some reduced to simple hairs. Fronds stipitate, the stipe with a single V- or U-shaped vascular bundle, abaxially terete or angled, adaxially grooved upwards, the grooves and edges continuous with those of the higher order branches, the lamina membranous to leathery, pinnately to bipinnate or decompound, the apical pinnae gradually reduced, or +/- the same as the lateral pinnae, strongly anadromous, ultimate free segments +/- equal-sided, with or without a costa or sostule, or dimidiate, the veins free, simple or 1 - several-times forked, or anastomosing without free included veinlets in the areoles, terminating short of the margin. Sori submarginal, on the upper and outer margins if the pinnules dimidiate, or on the non-basal margins if the pinnules equal-sided, terminal on a single vein or more commonly uniting 2 or more veins, the indusium attached basally, round or reinform on a single free vein or elongate and attached over several veins, free at the sides or not, opening towards the apical margin, parapyses 2-celled, fugacious; sporangia with a triseriate stalk, the annulus longitudinal, interrupted, of c.12-15 thickened cells. Spores trilete or less commonly monolete, smooth.

Subg. Lindsaea: terestrial with a radial stele

Sect. Schizoloma: rhizome essentially terrestrial, short- to moderately long-creeping, the stele radially symmetric or nearly so; lamina 2(-3)-pinnate, with the upper pinnae gradually reduced (no terminal pinna like the lateral), or pinnate with equal sided pinnules and anastomosing veins, or both; axes angular or terete; sori continuous or interrupted or interrupted by incisions of the margin; spores trilete.

Sect. Temnolindsaea: Rhizome essentially terrestrial, short-creeping, the stele radially symmetric or nearly so; lamina bipinnate with a terminal pinna like the laterals (pinnate in juvenile plants); axes abaxially 1-2-angluar; pinnules dimidiate; veins free; sori interrupted by incisions of the margin; spores trilete.

Sect. Synaphlebium: rhizome essentially terrestrial, short- to moderately long-creeping, the stele radially symmetric or nearly so; lamina pinnate, or bipinnate with a terminal pinna like the lateral; axes abaxially angular; pinnules dimidiate; veins anastomosing; sori entire or interrupted by incisions of the margin; spores trilete.

Sect. Osmolindsaea: rhizome essentially terrestrial, short-creeping, the stele radially symmetric or nearly so; lamina pinnate, the upper pinnae gradually reduced; axes abaxially rounded; pinnules dimidiate; veins free; sori interrupted by incisions of the margin (except in reduced forms); spores monolete.

Sect. Psammolindsaea: rhizome essentially terrestrial, short- to moderately long-creeping, the stele radially symmetric or nearly so lamina pinnate with a free terminal pinna like the laterals; axes abaxially rounded: pinnules dimidiate, not articulte to the rachis; veins free; sori continuously; spores trilete.

Sect. Steolindsaea: rhizome essentially terrestrial, short- to moderately long-creeping, the stele radially symmetric or nearly so; lamina pinnate with reduced upper pinnae, the terminal pinna unlike the laterals, the basal pinnae also reduced; axes abaxially bi-angular; pinnules dimidiate; veins free; sori continuous or interrupted by incisions of the margin; spores trilete.

Subg. Odontoloma: ephipytic with a dorsiventral stele

Sect: Odontoloma: rhizome epiphytic, long-creeping, +/- persistently scaly and dull, the stele strongly dorsiventral with a closed xylem strand; lamina pinnate with gradually reduced upper pinnae; axes abaxially angular; pinnules dimidiate; veins free; sori interrupted by incisions of the margin; spores trilete.

Sect. Lindsaenium: rhizome epiphytic, long-creeping, +/- persistently scaly and dull, the stele strongly dorsiventral with a closed xylem strand; lamina bipinnate with a terminal pinna like the laterals, or pinnate; axes abaxially angular; pinnules dimidiate; veins free or irregularly and infrequently anastomosing; sori interrupted by incisions of the margin; spores trilete.

Sect. Penna-arborea: rhizome epiphytic, long-creeping, deciduously scaly and lustrous, thestele strongly dorsiventral with a open xylem strand; lamina pinnate with gradually reduced upper pinnae or a large terminal segment; axes abaxially angular; pinnules dimidiate; veins free or anastomosing; sori interrupted by incisions of the margin; spores trilete.

Species in Papuasia

1a. Lamina pinnate and deeply pinnatifid, or bipinnate or more compound, without a conform terminal pinna, the upper pinnae gradually reduced L. media
Lamina simply pinnate, or if more compound then with a conform terminal pinna sharply set off from the lateral pinnae. 2
2a. Ultimate free divisions not, or only for a very short basal portion dimidiate 3
Ultimate pinnules dimidiate 6
3a. Veins anastomosing (L. ensifolia)

4

Veins quite free 5
4a. Upper pinnae gradually and strongly narrowed, gradually confluent into the terminal segment, or at least some small pinnae connected to it (sometimes bipinnate) L. ensifolia ssp. Agatii
Upper pinnae little reduced, the lamina with a free,conform terminal pinna L. ensifloia ssp. Ensifolia
5a. Pinnules articulate with the rachis L. queriniana
Pinnules continuous with the rachis L. walkerae
6a. Rhizome long-creeping, epiphytic or scandent, with remote leaves; stele dorsiventrally symmetric 7
Rhizome short-creepiong, terrestrial, or exceptionally longer and sometimes epiphytic; stele radially symmetric or nearly so 33
7a. Rhizome wiry, not over 1.3 mm diameter, deciduously scaly, eventually naked, dark brown to blackish, polished, with an open vascular strand; lamina simply pinnate but the pinnae sometimes deeply incised. 8
Rhizome stouter, 1.5-2 mm diameter, if thinner then not polished when naked, with a closed, strongly dosiventral vascular strand; lamina simply pinnate or bipinnate 13
8a. Veins anastomosing; larger pinnules 1.5-3 cm long; few or no upper pinnae gradually reduced 9
Veins free, or if anastomosing then the larges pinnules less than 1.5 cm long; upper pinnae gradually and strongly reduced 10
9a. Terminal segment narrow, lanceolate or caudate L. werneri
Terminal segment broad, obtuse, cuneate or flabellate 42
10a. Pinnules deeply pinnatifid, the segments connected by wings 0.25 mm - 0.5 mm wide L. roemeriana
Pinnules shallowly incised, or if more deeply incised then the lobes much more broadly connected1 (L. pulchella)

11

11a. Uninerval sori with a straight or convex base; indusium falling short of the margin by its own width or less, veins free or anastomosing, scales reddish brown, with a well-developed uniseriate apex L. pulchella var. lomotosora
Uninrval sori mostly with a very concave base; indusium falling short of the margin by its own width or more (except sometimes at the incisions); veins free; scales golden brown, apically very shortly uniseriate 12
12a. Pinnules herbaceous, mostly green when dry, 4.5 - 8 mm long c.twice as long as wide, the major incisions less than 0.5 mm deep; adaxial and abaxial sides of stipe not discolourous L. pulchella var. pulchella
Pinnules herbaceous, mostly brown when dry, the larger ones 7-12 mm long, up to twice as long as wide, the major incisions 1-3 mm deep; stipe faces not discolourous L. pulchella var. blanda
12c. Pinnules herbaceous to subcoriaceous, 10-16 mm long, 2.5-3 times as long as wide, incisions 1(-2) mm deep; adaxial face of petiole much paler than the abaxial side L. pulchella var. falcata
13a. Lamina simply pinnate 14
Lamina of most, or at least some fronds bipinnate 28
14a. Veins anastomosing 15
Veins free 20
15a. Upper pinnae not reduced, apex of a free or almost free terminal pinna 16
Pinnae gradually reduced, the apex narrowed and lobed 18
16a. Venation of lateral pinnae free, several anastomoses in the larger terminal pinna; sorus remote from the margin by 1-2 times its width; apical pinna +/- flabellate L. jarrettiana
Venation of lateral pinnae always with at least some annastomosis; sorus remote from the margin by less than its width; apical pinna flabellate or narrow-cuneate 17
17a. Sori 2-3-nerval; pinna translucent; terminal pinna narrow-cuneate, or with 2 narrow cuneate divisions L. modesta
Sori 4 - plurinerval; pinnae opaque; terminal pinnule flabellate L. obscura
18a. Indusium reaching the margin or nearly so 19
Indusium not reaching the margin by a distance equal to its width or more L. pulchra
19a. Major pinnules c. 20-25 to a side, their deepest incisions reaching down to 1/3 of their width, the larger pinnules 2-2.5 times as long as wide L. chrysolepis
Major pinnules c.30-45 to a side, their deepest incisions reaching down to 1/2 of their width, the larger pinnules 3 times as long as wide L. salomonis
20a. Upper pinnae not reduced, apex of a free or almost free flabellate terminal pinna L. jarrettiana
Pinnae gradually reduced, the apex narrowed and lobed

21

22a. Pinnules incised far beyond the middle, the lamina as a wing along each vein; sori 1-2-nerval 23
Pinnules subentire to incised to the middle, sori 1 - plurinerval 24
23a. Basal pinnule-lobes only bifid L. rosenstockii
Pinnules more dissected L. versteegii
24a. Basal pinnules inserted on the edge of the adaxial face of the rachis L. microstegia
Basal pinnules inserted below the adaxial face of the rachis (L. reopens)

25

25a. Rhizome delicate, less than 1 mm diameter L. reopens var. deliculata
Rhizome of full grown plants (1.5-)2-3 mm diameter 26
26a. Indusium with a very concave base, +/- reinform; most sori 1-nerval and round, some 2-nerval and crescent-shaped; margin very regularly incised to somewhat beyond the level of the receptacle L. repens var. sessilis
Indusium with a weakly concave, straight or somewhat convex base; sori 1 - plurinerval; margin incised almost to receptacle or more deeply 27
27a. Sori almost marginal, the indusium reaching to slightly surpassing the margin; pinnules elongate, to 3-4 times as long as wide; incisions of margin reaching or slightly surpassing receptacle L. repens var. cheilosora
Sori distinctly intramarginal, the indusim falling short of the margin; pinnules 2-3 times as long as wide; margin incised considerably beyond receptacle L. repens var. lingulata
28a. Secondary axes abaxially rounded L. monocarpa
Secondary axes biangular and/or sulcate, at least in the upper half 29
29a. Pinnules coriaceous (veins sometimes casually anastomosing) L. rigida
Pinnules herbaceous to chartaceous 30
30a. Pinnules shallowly incised, at most to the receptacle or slightly beyond, never to the middle 31
Pinnules incised considerable beyond the middle, the lamina as a wing along each vein 32
31a. Adaxial groove of primary rachis broad, occupying 1/2 of its width or more; most pinnules fertile only near the apex, always bipinnate; indusium not reaching the margin by less than its width; (veins sometimes casually anastomosing, if regularly anastomosing go to 42) L. rigida
Adaxial groove of primary rachis narrow, occupying much less than 1/2 of its width; most pinnules entirely fertile; not rarely simply pinnate leaves present alongside fertile ones; indusium short of the margin by 1.5-3-times its width; (veins always free) L. microstegia
32a. Pinnules more dissected, all lobes at least once forked; segments narrow, 0.2-0.3 mm wide L. versteegii
Basal pinnule-lobes only bifid, the outer lobes often not forked; segments narrow, 0.6-2 mm wide L. rosenstockii
33a. Veins free 34
Veins anastomosing 42
34a. Simply pinnate; rachis abaxially rounded or narrow-rounded; spores monolete L. odorata
Simply pinnate with an abaxially biangular rachis or bi-(subri-) pinnate; spores trilete 35
35a. Simply pinnate 36
Bipinnate 40
36a. Pinnae entire, or if incised then the inner incisions of the larger pinnules not going beyond the middle (L. lucida) 37
Pinnae incised far beyond the middle 38
37a. Stipe 3-15 cm long, 0.2-0.5 the length of the lamina; indusium 0.3-0.4 mm wide, at least in the inner sori almost or quite reaching the margin L. lucida ssp. Lucida
Stipe 1-2(-4) cm long, c. 0.05 - 0.07 the length of the laming; indusium 0.2 mm wide, more strongly intramarginal L. lucida ssp. Brevipes
38a. Pinnules 2-2.5 times as long as wide (L. bakeri)

39

Pinnules 3-4 times as long as wide L. lapeyrousii ssp. Lapeyrousii
39a. Axes stramineus to medium brown; ultimate lobes 0.3-0.5 mm wide, at the sorus 0.5-1 mm wide; sori hardly ever binerval L. bakeri var. bakeri
Axes medium to dark brown; ultimate lobes 0.6 - 1 mm wide, 0.8 - 1.2 mm wide a the sorus; sori not rarely binerval L. bakeri var. pycnophylla
40a. Stipe and rachises abaxially keeled L. tenuifolia
Stipe and rachises abaxially biangular, rarly only obscurely so 41
41a. Basal primary pinnae strongly reduced L. kingii
Basal pinnae hardly or not reduced L. tetragona
42a. Sori of larger, fully fertile pinnules continuous 43
Sori of larger pinnules interrupted by incisions of the margin 45
43a. Upper pinnules little reduced, c.half as long as larger ones, the ultimate segment comparatively large, lanceolate, free or nearly so L. cultrata
Upper pinnules strongly reduced, some tooth-like ones connected with the narrow pinna-apex 44
44a. Pinnules not over twice as long as wide L. azurea
Pinnules 2.5-3- or more-times as long as wide L. papuana
45a. Terminal pinnule quite free, cuneate-flabellate or cuneater; rhizome not very short-creeping 46
Terminal segment narrow and connected by some tooth-like upper pinnules, or if free, triangular and acute. 48
46a. Venation of lateral pinnae free, several anastomoses in the larger terminal pinna; sorus remote from the margin by 1-2 times its width; apical pinna +/- flabellate L. jarrettiana
Venation of lateral pinnae always with at least some anastomosis; sorus remote from the margin by less than its width; apical pinna flabella or narrow-cuneate 47
47a. Sori 2-3 nerval; pinnae translucent; terminal pinna narrow-cuneate, or with 2 narrow cuneata divisions L. modesta
Sori 4- plurinerval; pinnae opaque; terminal pinnule flabellate L. obscura
48a. Pinnules opaque, hardly narrowed to the obliquely truncate apex, the outer margin distint, with an incision; pinnae rather suddenly narrrowed below the +/- caudate, pinnatifid apex; pinnule-bearing rachises abaxially brown, sulcate and pale-margined L. parallelogramma
Pinnules often translucent, with a subacute to rounded apex, and/0r distinctly narrowed towards the apex, or if truncate then the outer margin not incised; pinnae more gradually narrowed; pinnule bearing rachises various 49
49a. Sorus on the outer margin in most or all pinnules continuous with the upper most sorus of the upper margin. L. cultrata
Sorus or sori of the outer margin distince, or no distinct outer margin to pinnules developed 50
50a. At least the larger inner lobes of the pinnules truncate, with straight or concave outer margins 51
Lobes of pinnules with a convex outer margin 54
51a. Indusium almost or quite reaching the margin, 0.2-0.3 mm wide, or if wider, then the incisions on the pinnule-margin not reaching to the level of the receptacle. 52
Indusium falling short of the margin by 1-2-times its width, 0.3 mm or usually sider 53
52a. Most incisions of the margin not reaching the level of the receptacle, some reaching up to it; pinnules narrowed from base to apex L. pacifica
Most incisions of the margin reaching c. twice as far as the distance from the receptacle to the margin; pinnules apically +/- truncate. L. obtusa
53a. Lamina usually brownish or olivaceous when dry; lobes of pinnules hardly divergent, +/- contiguous; lamina simply pinnate or bipinnate with a few pinnae. L. obtusa
Lamina medium to dark green when dry; lobes of pinnules divergent, not contiguous; lamina often bipinnate with several pairs of pinnae L. harveyi
54a. Indusium almost or quite reaching the margin; at least some of the longer sori concave on the inner, covex on the outer side L. lobata
Indusim not reaching the margin, falling shrot of it by a distance equal to its width or more, straight on the inner side. L. harveyi