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Cyathea Sm.

Terrestrial, medium-size to large, mostly arborescent, rarely climbing (clinging by roots to a supporting tree). Rhizome an erect, stout caudex bearing a radial tuft of fronds at the apex, dictyostelic, supported by strong bands of sclerenchyma, the young parts and bases of stipe bearing basally attached, variously fringed scales. Fronds short- to long-stipitate, the stipe with numerous small vascular bundles arranged in two adaxial and one abaxial series, with discontinuous pneumathodes (sometimes in 2-3 rows), rachises adaxially raised but not continuous with each other, the lamina pinnate, mostly bipinnate-pinnatifid, sometimes tripinnate, the basal pinnae +/- reduced, the veins in the ultimate pinnules pinnately arranged from a prominent costule, simple, forked or pinnate in each lamina lobe; the sterile and fertile fronds iso-, di- or subdimorphic. Sori intramarginal or medial on the ultimate veins, usually at the fork of a vein, the receptacle erect, domed, to globular or calvate, exindusiate, or with an indusim attached completely around the receptacle and covering the young sorus, rupturing irregularly or to form an even-edged cup, or attached on the costular side of the receptacle (hemitelioid), often minute and hidden by sorus, or as separate scales around the base of the receptacle, +/- covering the young sorus, sporangia shor-stalked, annulus oblique, stomium lateral, paraphyses simple, multiseptate. Spores trilete, translucent, smooth to papillose.

Subgenus Cyathea: stipe scales flabelloid

Sect. Cyathea: Stipe scales flabelloid or marginate, several cells thick at the base; pinnules generally deeply lobed and the costules not widely spaced, the basal basiscopic vein rarely from the costa; sometimes with crisped, appressed hairs beneath; indusium cup-shaped, hemitelioid or lacking.

Sect. : Stipe scales flabelloid or marginate, several cells thick at the base; pinnules usually dimorphic, generally deeply lobed and the costules not widely spaced, the basal basiscopic vein rarely from the costa; axes very dark; hairs lacking beneath; indusium lacking.

Subgenus Sphaeropteris: stipes scales setiferous

Sect. Sphaeropteris: costules close, deeply lobed

Subsect. Sphaeropteris: Stipe scales conform or setiferous, flat and few cells thick, or as stiff, setate bristles; pinnules deeply lobed and the costa not widely spaced, the basal basiscopic vein from the costule, free tertiary leaflets few; thick spreading hairs may be present; indusium +/- cup-shaped or lacking.

Subsect. Fourniera: Stipe scales conform on setiferous, flat and few cells thick, throughtout; pinnules deeply lobed and the costa not widely spaced, the basal basiscopic vein from the costule, free tertiary leaflets many; fringed scales but no thick spreading hairs may be present; indusium lacking, young sorus protected by overlapping scales.

Sect. Schizocaena: costules widely spaced, shallowly lobed

Subsect. Schizocaena: stipe scales conform or setiferous, one cell thick throughtout; pinnules not deeply lobed, costules widely spaced, basal basiscopic vein always from costa; thick, spreading hairs may be present; indusium +/- cup-shaped or lacking.

Subsect. Sarcopholis: stipe scales conform or setiferous, thick and fleshy at the base, upcurved, tapering and flat distally; pinnules not deeply lobed, costules widely spaced, basal basiscopic vein always from costa; thick, spreading hairs may be present; indusium +/- cup-shaped or lacking.

Species in Papuasia

Key to the sections and subsections:

1a. Stipe scales flabelloid, several rows of marginal cells of different texture, often erose; hairs on the lower surface if present , crisped and appressed; pinnules in most cases deeply lobed, the basal basiscopic vein rarely from the costa; indusium in some cases hemitelioid or incompletely surrounding the receptacle (subg. Cyathea) 2
Stipe scales setiferous, with dark marginal bristles; hairs on lower surface if present rather thick and spreading; where pinnules are shallowly lobed, basiscopic vein always from costa; indusia complete or lacking, or formed by separate scales (in some cases imperfect, then not hemitelioid) (subg. Sphaeropteris)

3

2a. Indusiate (indusia in some cases very small), or if exindusiate, hairy on lower surface of pinna-rachis; axes not very dark, little dimorphism between sterile and fertile pinnules Sect. Cyathea
Exidusiate; axes very dark, not hairy beneath; fertile and sterile pinnules usually very dimorphous Sect. Gymosphaera
3a. Costules not widely spaced (rarely over 4 mm apart in pinnules 10 cm long); pinnules usually more than 10 cm long, lobed almost or quite to costa throughout, or fully pinnate (sect. Sphaeropteris)

4

Costules widely spaced (at least 4 mm apart except where pinnules are under 4 cm long), pinnules mostly less than 10 cm long, not lobed to within 1 mm of the costa except near the base; basal basiscopic vein always from costa (sect. Schizochaena)

5

4a. Free tertiary leaflets few; indusia present or absent; sori never covered with overlapping scales Sect. Sphaeropteris subsect. Sphaeropteris
Free tertiary leaflets many; no indusia, the sori covered by overlapping scales Sect Sphaeropteris subsect. Fourniera
5a. Scales of stipe 1 cell thick throughout (not in Papusia) Sect. Schizocaena subsect. Schizocaena
Scales of stipe thick and fleshy at the base, tapering and flat distally Sect. Schizocaena subsect. Sarcopholis

Subgen Cyathea sect. Cyathea

Species in Papuasia

1a. Pinnules not over 30 mm long; segments constricted at the base or the lower ones quite free 2
Pinnules longer, or if under 30 mm long, then without free basal segments 9
2a. Lower surface of pinna-rachis bearing crisped hairs 3
Lower surface of pinna-rachis not hairy 7
3a. Sori lacking indusia C. lepidoclada
Sori indusiate, the indusium cup-shaped or incompletely surrounding the receptacle (hemitelioid) 4
4a. Indusium hemitelioid C. nothofagorum
Indusium cup-shaped 5
5a. Bullate scales present on pinna-rachis and/or costae 6
Bullate scales lacking from pinna-rachis and/or costae C. hunsteiniana
6a. Pinnules to c. 30 mm long C. microphylloides
Pinnules to c.15 mm long C. perpelvigera
7a. Largest pinnae 8.5 cm long; pinnules to 11 mm long C. arfakensis
Largest pinnae 22-26 cm long; pinnules 23-30 mm long 8
8a. Stipe 30 cm long; pinna-rachis glabrescent; sori 4-5 to each tertiary leaflet C. ledermannii
Stipe 10 cm long, pinna-rachis covered with sma;ll bullate scales; a single sorus at the base of each tertiary leaflet C. hooglandii
9a. (1) Indusium an entire cup with an even edge at maturity of sorus, if fragile sometimes breaking later 10
Indusium otherwise or lacking 19
10a. Lower surface of lamina segments almost or quite covered with scales or hairs 11
Lower surface of lamina segments not so-covered 12
11a. Small scales abundant on lower surface of veins; costal scales mostly bearing dark setae C. percrassa
Crisped hairs on lower surface of veins; costal scales bearing many slender crisped marginal hairs C. vandersenii
12a. Lower surface of lamina strongly glaucous C. pruinosa
Lower surface of lamina not glaucous 13
13a. Bullate scales present on pinna-rachis and/or costae 14
Bullate scales present on pinna-rachis and/or costae 14
14a. No fine crisped hairs on lower surface of pinna-rachis, nor on the edges of its scales C. pvcnoneura
Fine crisped hairs present on pinna-rachis or on its scales 15
15a. Indusial cup shallow; small scales on pinna-rachis bearing very fine hairs; no coarse crisped hairs; bullate scales only on distal part of pinna-rachis C. rigens
Indusial cup deep; crisped hairs, not small fringed scales, also bullate scales present on pinna-rachis C. everta
16a. Lower surface of pinna-rachis covered with crisped hairs C. cincinnata
Lower surface of pinna-rachis lacking such hairs 17
17a. Pinna-rachis almost glabrescent on lower surface C. subtripinnata
Pinna-rachis persistently sclay on lower surface 18
18a. Scales on pinna-rachis mostly narrow, 4-5 mm long, with long flexuous marginal setae; stipe over 20 cm long C. costalisora
Scales on pinna-rachis mostly very small, thin, short-fringed, mostly not setiferous, forming a continuous felt C. coactilis
19a. (9) Indusium at maturity an almost flat disk, symmetric or not, in some cases hidden by the mature sorus, in some cases the residual part of a complete indusium of which the thin apical part falls away at maturity. 20
Indusium otherwise or lacking 28
20a. Indusium a very narrow wing around the base of the receptacle 21
Indusium ultimately a regular or irregular disc, almost or quite as big as the base of the sorus; if irregular then usually the base of a formerly complete indusium 23
21a. Scales on costae bullate-based C. parva
Scales on costae flat or lacking, not bullate 22
22a. Broad flat scales with a few marginal setae on costae. C. wengiensis
Broad flat setiferosus scales lacking on costae C. batjanensis
23a. Indusium at maturity a disc with a fairly even edge 24
Indusium at first covering sorus, the apical part very thin and caducous, an irregular disc remaining on the old sorus 26

24a.

Pinna-rachis hairy on the lower surface, at least distally C. albidosquamata
Pinna-rachis not hairy on lower surface 25
25a. Scales on stipe dark; scales on pinna-rachis bullate. C. catillifera
Scales on stipe pale; scales on pinna-rachis not bullate, the larger ones dwith marginal setae C horridula
26a. Pinna-rachis hairy on lower surface, at lest distally C. tenuicaulis
Pinna-rachis not hairy on lower surface 27
27a. Fronds simply pinnate, pinnae to 7x1.8cm, lobed to c.2 mm from costa C. klossii
Fronds bipinnate C. rubiginosa
28a. (19) Indusium covering sorus to maturity, then breaking and persistent 29
Indusium hemitelioid (sometimes almost covering sorus to maturity, sometimes very small), or lacking 41
29a. Pinna-rachis conspicuously hairy and scaly on lower surface C. geluensis
Pinna-rachis not hairy, though sometimes scaly on lower surface 30
30a. Lamina very rigid, the small tertiary lobes with the edges much reflexed so that the sori are almost enclosed 31
Lamina not very rigid with strongly reflexed edges 32
31a. Larger tertiary leaflets lobed, each with 3-5 sori C. macgregorii
Tertiary leaflets not lobed, each with 1-2 sori C. gleichenioides
32a. Pinnules to 2.5 cm long, less than 10 mm wide C. imbricata
Pinnules at least 40 x 10mm, in most cases most larger 33
33a. Largest pinnules more than 100 x 20mm; costules 5 mm or more apart C. insulana
Largest pinnules not over 100 x 20mm; costules 304.5 mm apart 34
34a. Pinnules c.45 x 10mm, very rigid; stipe scales rigid, c. 40x1mm C. pseudomuelleri
Pinnules commonly more than 65 x 15mm; stipe scales otherwise 35
35a. Costae and costules densely scaly; scales mostly setiferous, not bullate; inna-rachis persistently covered with very small setiferous scales 36
Costae and costules bearing scales that arem ostly not setiferous; pinna-rachis glabrescent or its scales not seiferous C. foersteri
36a. Veins bearing scales on the lower surface 37
Veins lacking scales on lower surface 40
37a. Stipe 5-15cm, the lowest pinnae 5-12 cm long 39
Stipe 50 cm or more long C. archboldii

38

38a. Stipe with pale brown scales and short spines; setiferous scales common on veins C. archboldii var. archboldii
Stipe with dark brown scales and spines to 2.5 cm long; scales on veins few and not setiferous C. archboldii var. horrida
39a. Larger scales on pinna-rachis pale; veins dark and raised on lower surface C. foersteri
Larger scales on pinna-rachis with dark median band; veins concolorous, not raised below C. nigrolineata
40a. Larger scales on pinna rachis pale C. foersteri
Larger scales on pinna rachis with a dark median band C. nigrolineata
41a. (28) Indusium lacking 42
Indusium hemitelioid, of varying size, sometimes hidden by sorus 46
42a. Fronds simple pinnate, or bipinnate with small pinnules lobed halfway to costa 43
Fronds bipinnate, pinnules more deeply lobed 44
43a. Fronds simply pinnate C. ascendens
Fronds bipinnate, pinnules c.30 mm long C. recurvata
44a. Pinna-rachis densely covered with long tangled pale crisped hairs and narrow scales on the lower surface C. eriophora
Pinna-rachis rather sparsely hairy on the lower surface 45
45a. Sori near costules; dark hairs present on costules beneath C. gregaria
Sori medial or nearly so; no dark hairs on costules beneath C. macgillivrayi
46a. (41) Lamina segments, both sterile and fertile, lobed at least half-way to costule, the veins often pinnate in the lobes 47
Sterile lamina segments not deeply lobed, the lateral veins in the segments not pinnate 48
47a. Paraphyses longer than sporangia; thick hairs abundant on lower surface of costae and costules C. croftii
Paraphyses shorter than sporangia; thick hairs few or sometimes lacking on lower surface of costae and costules C. decurrens
48a. Pinna-rachis densely hairy throughout lower surface C. cucullifera
Pinna rachis hairy at most on the distal part (may be scaly throughout) 49
49a. Indusium entirely brown and rather firm, entirely covering sorus at maturity, breaking only when old 50
Indusium in part pale and fragile, breaking at maturity, or not entirely covering sorus 51
50a. Stipe scales c.50 mm long, shiny brown; costae almost glabrous; pinnules c.50 mm long C. muelleri
Stipe scales 25-30 mm long, pale; costae very scaly; pinnules to 100 mm long C. pachyrrhachis
51a. Indusium visible as a scale backing costule, not entirely hidden by sorus 52
Indusium very small, +/- hidden by sporangia 53
52a. Indusium more than a semicircle, concave towards the sorus and sometimes covering part of the sorus to maturity C dicksonioides
Indusium not more than a semicircle, reflexed against costule at maturity C. semiamplectens
53a. Pinnules commonly more than 20 mm wide C. glaberrima
Pinnules commonly not more than 20 mm wide 54
54a. Pinna-rachis rather densely covered with bullate scales 55
Pinna rachis not so-covered 56
55a. Pinnules c.30x 10mm C. physolepidota
Pinnules c.90x 19mm C. alta
56a. Scales on costae and costules all flat, or at most convex 57
Scales on costules bullate, some bullate scales often on costae 59
57a. Lower pinnules stalked; no hairs on pinna-rachis and costae 58
Lower pinnules sessile; hairs present on pinna-rachis and costae C. wengiensis
58a. Lowest 1-2 pairs of segments quite free and articulate; lamina thick and rigid C. kanehirae
Lowest segmetns, several pairs, contracted at the base but not free; lamina not thick C. nigropaleata
59a. Stipe 20-25 cm long, the basal pinnae gradually reduced in size; pinnules 10-15 mm wide, costules 2-3.5 mm apart C. media
Stipe c.90 cm long, with a pair of reduced basal pinnae c.15 cm above the base; pinnules 15-20 mm wide, costules c.5 mm apart C. solomonensis

Subgen. Cyathea sect. Gymnosphaera

Species in Papuasia

1a. Fronds simply pinnate with entire pinnae (sterile), or bipinnate with few pinnae and almost entire sterile pinnules; in either case the true apex of the frond short and abortive , evident above the attachment of the uppermost pinna 2
Fronds at least bipinnate, with a normal apex 3
2a. Fertile pinnules lobed almost or quite to costa C. biformis
Fertile pinnules only slightly lobed C. scandens
3a. Scales on lower surface of costules bullate 4
Scales on lower surface of costules not bullate 6
4a. Bullate scales present on lower surface of veins of sterile pinnules C. hornei
Bullate scales lacking on veins 5
5a. Pinnules to 110 x 30mm, cut to 2 mm from costa; costules of sterile pinnules to 6 mm apart C. rubella
Pinnules to 70 x 20mm, cut to within 1 mm from costa; costules of sterile pinnules to 4.5 mm apart C. macgillivrayi
6a. Costal scales not setiferous C. acrostichoides
Costal scales bearing many setae C. schlechteri

Subg. Sphaeropteris sect. Sphaeropter is subsect. Sphaeropteris

Species in Papuasia

1a. Indusium present 2
Indusium lacking 4
2a. Base of stipe covered with thin, pale scalers to 35 x 1 mm C. whitmeei
Base of stipe bearing many bristles 10-20 mm long, spreading at right angles 3
3a. Slender hairs 1 mm long abundant on costae, costules and veins beneath C. pulcherrima
Such hairs lacking, at most much shorter hairs present towards the apex of pinnules C. procera
4a. Scales of stipe base to 50 mm or more long, shining medium brown, rigid edges bearing rather long concolorous setae; lamina very rigid, the edges stongly reflexed and inrolled 5
Scales of stipe base mostly shorter, often not rigid and pale, the edges mostly bearing close dark setae; edges of lamina rarely much reflexed, then not inrolled. 7
5a. Pinnules to 30 mm long; sacles on stipe base 2 mm wide; pinna-rachis rather persistently covered with a felt of small, pale fringed scales; scales on costusles bearing pale, crisped marginal hairs C. tomentosissima
Pinnules 45-80 mm long; scales on stipe base 3-5 mm wide; pinna-rachis not persistently or densely so-covered; scales on costules often bearing long, dark setae (C. atrox) 6
6a. Stipe bearing many dark shiny spines 3-6 mm long; pinnules to 80 mm long C. atrox var. atrox
Stipe lacking spines; pinnules commonly to 45 mm long. C. atrox var. inermis
7a. Upper part of stipe and all rachises finely and very closely warty; bullate scales always present on costules 8
Upper part of stipe and rachises thorny or conspicuously warty, the warts not very fine or close; bullate scales present or not 10
8a. Long pale hairs normally lacking on lower surface of costae and costules, present on upper surface of costules; no setiferous scales on costae C. felina
Long pale hairs always rather abundant distally on the lower surface of costae and costules, lacking or very few on upper surface of costules; setiferous scales present, at least near base of costules 9
9a. Bullate scales abundant on lower surfae of veins; stipe scales to 15 x 3mm, very think C. vittata
Bullate scales lacking or rare on lower surface of veins; stipe scales mostly longer and proportionately narrower C. lunulata
10a. Scales on constules abundant, copiously fringed with long, crisped hairs which become entangled and form a wolly covering C. magna
Scales on costules not thus fringed 11
11a. Bullate scales present on lower surface of costae and costules 12
Bullate scales lacking on lower surface of costae and costules 15
12a. Copious long hairs also on lower surface of veins C pilulifera
Long hairs lacking on lower surface of veins or only at vein tips (C. aenifolia)

13

13a. Frond 175-250 cm long, stipe 50-60 cm long; largest pinnae 30-60 cm long; longest pinnules 7-12 cm long; costules 3-4.5 mm apart 14
Frond to 425 cm long, stipe 200 cm long; largest pinnae 70c. long; longest pinnules 13.5 cm long; costules 5 mm apart C. aenifolia var. macrophylla
14a. Stipe scales 30-60 x 3.5mm, pale brown C. enifolia var. aenifolia
Stipe to 15x2 mm long, dark brown C. aenifolia var. melanocantha
15a. Pale erect hairs abundant on lower surface of veins 16
Pale erect hairs absent or rare (then near apices of segments) on lower surfaces of veins 17
16a. Costae bearing many much-setiferous scales; pinnules to 20 mm wide C. engiensis
Costae bearing very few scales; pinnules commonly 20-30 mm wide C. contaminans
17a. Setiferous scales abundant on costae; small scales present on lower surface of veings; no stout hairs on lower surface of costae 18
Setiferous scales lacking or rare on lower surface of costae of mature fronds; no scales on lower surface of veins; stout pale hairs present near apices on lower surface of costae C. contaminans
18a. Stipe scales 5 mm wide; pinnules to 25 mm wide; veins 13-15 pairs C. atrospinosa
Stipe scales 2 mm wide; pinnules to 35 mm wide; veins to 20 pairs C. fugax

Subg. Shaeropteris sect. Sphaeropteris subsect. Fourniera

Species in Papuasia

1a. Pinnules 30-40 mm long; lower pinnae gradually reduced 2
Pinnules commonly at least 70 mm long; in most cases much longer 3
2a. Tertiary leaflets on stalks almost 1 mm long; pinnae to 25 cm long; bullate scales present on costules C. carrii
Tertiary leaflets almost sessile; pinnae to 4-5 cm long, no bullate scales present on costusles C. womerslevi
3a. Lower pinnae gradually reduced, the lowest c. 11 cm long near the base of the stipe C. auriculifera
Lower pinnae not thus reduced (sometimes 1-2 pairs of isolated small pinnae at the base of the stipe 4
4a. Bullate scales present on costae and costules 5
Bullate scales lacking 6
5a. Soral scales small, not covering sorus to maturity C. truncata
Soral scales large, covering sorus to maturity C. aciculosa
6a. Veins and lower surface of lamina bearing finely fringed interlacing scales which cover the lower surface of the leaflets C. celebica
Veins bearing at most all separate scales on lower surface (C. macrophylla)

7

7a. Stipe to at least 40 cm long; tertiary leaflets to 15 x 3.5mm, the larger ones deeply lobed at the base C. macrophylla var. macrophylla
Stipe c.10 cm long; largest tertiary leaflets 18 mm long, all deeply lobed C. macrophylla var. quadripinnata

Subg. Sphaeropteris sect. Schizocaena subsect. Sarcopholis

Species in Papuasia

1a. Pinna-rachis hairy on lower surface 2
Pinna-rachis not hairy on lower surface 3
2a. Scales on costae setiferous C. fusca
Scales on costae not setiferous C. rosenstockii
3a. Indusium quite lacking C. marginata
Indusium present 4
4a. Pinnules on stalks to 6 mm long; stipe to 60 cm long C. papuana
Pinnules sessile or on much shortened stalks; stipe much shorter 5
5a. Pinnules to 35 mm wide, the longer ones with lowest 1-2 lamina segments free; no bullate scales C. inaequalis
Pinnules rarely over 25 mm wide, lowest segments not free; bullate scales usually present on costules 6
6a. Pinnules lobed only in the basal half C. parvipinna
Pinnules deeply lobed throughout 7
7a. Pinnules not over 20 mm wide, lobed to c3 mm from costa C. werneri
Pinnules of largest pinnae in most cases over 20 mm wide, lobed to 1-2 mm from costa 8
8a. Scales of costae and costules bullate, mostly entire C. insularum
Scales on costae mostly small and fringed or setiferous; bullate scales on costae and costules mostly fringed 9
9a. Pinnules of largest pinnae 22-28 mm wide 10
Pinnules of largest pinnae less than 20 mm wide C. moseleyi
10a. Stipe greater than 25 cm long; all pinnules lobed to 1 mm from costa; costules 5-6 mm apart, veins c.10 pairs C. brackenridgei
Stipe shorter; lowest segments of pinnules free, the rest lobed to 2 mm from costa; costules 6-7 mm apart, veins c. 8 pairs C. runensis