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39. Polypodiaceae

水龙骨科 shui long gu ke

Authors: Zhang Xianchun, Shugang Lu, Youxing Lin, Qi Xinping, Shannjye Moore, Prof. Fuwu Xing, Wang Faguo, Peter H. Hovenkamp, Michael G. Gilbert, Hans P. Nooteboom, Barbara S. Parris, Christopher Haufler, Masahiro Kato & Alan R. Smith

Drynaria delavayi

Credit: Harvard University Herbaria

Plants mostly epiphytic and epilithic, a few terrestrial. Rhizomes shortly to long creeping, dictyostelic, bearing scales. Fronds monomorphic or dimorphic, mostly simple to pinnatifid or 1-pinnate (uncommonly more divided); stipes cleanly abscising near their bases or not (most grammitids), leaving short phyllopodia; veins often anastomosing or reticulate, sometimes with included veinlets, or veins free (most grammitids); indument various, of scales, hairs, or glands. Sori abaxial (rarely marginal), orbicular to oblong or elliptic, occasionally elongate, or sporangia acrostichoid, sometimes deeply embedded, sori exindusiate, sometimes covered by caducous scales (soral paraphyses) when young; sporangia with 1-3-rowed, usually long stalks, frequently with paraphyses on sporangia or on receptacle; spores hyaline to yellowish, reniform, and monolete (non-grammitids), or greenish and globose-tetrahedral, trilete (most grammitids); perine various, usually thin, not strongly winged or cristate. Mostly x = 35, 36, 37.

More than 50 genera and ca. 1,200 species: pantropical, a few temperate; 39 genera and 267 species (82 endemic) in China.

The Polypodiaceae s.s., as often recognized is paraphyletic, because it excludes the grammitids. Generic boundaries need clarification. Polypodiaceae contains large wholly neotropical and wholly paleotropical clades.

Molecular data show that several families recognized in FRPS, i.e., Drynariaceae, Grammitidaceae, Gymnogrammitidaceae, Loxogrammaceae, Platyceriaceae, and Pleurosoriopsidaceae, all nest within the Polypodiaceae. Five lineages are defined that can be treated as subfamilies (see Christenhusz et al., Phytotaxa 19: 18-19. 2011) as follows: Loxogrammoideae (Loxogrammaceae): genus no. 1; Drynarioideae (Crypsinoideae, Drynariaceae, Gymnogrammitidaceae): genera nos. 2-8; Platycerioideae (Platyceriaceae, Pyrrosioideae): genera nos. 9-10; Microsoroideae (incl. Crypsinoideae, Lepisorioideae): genera nos. 11-25; Polypodioideae (incl. Grammitidaceae, Pleurosoriopsidaceae): genera nos. 26-39. The status of several genera in the Microsoroideae is controversial (Kreier et al., Molec. Phylogen. Evol. 48: 1155-1167. 2008). Microsorum is still polyphyletic, and some new genera are to be established.

Ching Ren-chang, Fu Shu-hsia, Wang Chu-hao & Shing Gung-hsia. 1959. Gymnogrammitis. In: Ching Ren-chang, ed., Fl. Reipubl. Popularis Sin. 2: 284-285; Wu Shiew-hung. 1999. Pleurosoriopsidaceae. In: Wu Shiew-hung, ed., Fl. Reipubl. Popularis Sin. 4(2): 154-156; Lin Youxing, Lu Shugang & Shi Lei. 2000. Polypodiaceae. In: Lin Youxing, ed., Fl. Reipubl. Popularis Sin. 6(2): 7-266, 346-349; Zhang Xianchun. 2000. Drynariacae, Platyceriaceae, Grammitidaceae, and Loxogrammaceae. In: Lin Youxing, ed., Fl. Reipubl. Popularis Sin. 6(2): 267-335.


1 Fronds with stellate hairs, often stellate-tomentose at least when young   (2)
+ Fronds with scales, unbranched hairs, glandular hairs, and/or forked hairs, or glabrous   (3)
       
2 (1) Fronds monomorphic or weakly dimorphic, fronds not with base adpressed to substrate, fertile fronds similar in form to sterile fronds differing only in relative width, usually simple, rarely hastate or pedately divided.   9 Pyrrosia
+ Fronds strongly dimorphic, basal fronds with basal part tightly adpressed to substrate, foliage/fertile fronds dichotomously lobed.   10 Platycerium
       
3 (1) Specialized humus-collecting fronds or widened frond bases present   (4)
+ Specialized humus-collecting fronds or lamina bases absent   (5)
       
4 (3) Fronds usually strongly dimorphic with specialized short brown sessile humus-collecting fronds and longer pinnatisect fertile fronds.   4 Drynaria
+ Fronds monomorphic, fronds with broad humus-collecting base and distal deeply pinnatisect fertile portion.   2 Aglaomorpha
       
5 (3) Nectaries present along costa   (6)
+ Nectaries absent   (7)
       
6 (5) Fronds pinnatisect, stipe winged to base; segments uniformly 10-15(-20) mm wide; sporangia in discrete, orbicular sori (Drynaria parishii).   4 Drynaria
+ Fronds pinnate, stipe and rachis terete; segments dimorphic, sterile segments 3.5-10 cm wide, fertile segments distal, 0.4-0.7 cm wide; sporangia acrostichoid, covering most of pinnule surface.   3 Photinopteris
       
7 (5) Fronds 2-pinnatifid to 4-pinnate   (8)
+ Fronds entire, pinnately divided, or 1-pinnate, rarely palmately to pedately divided   (10)
       
8 (7) Fronds finely 3- or 4-pinnate, sparsely scaly.   8 Gymnogrammitis
+ Fronds 2-pinnatifid, hairy   (9)
       
9 (8) Lamina 1.5-8 ?0.5-1.5 cm; sori linear-oblong, along veins.   27 Pleurosoriopsis
+ Lamina 5-25 ?2-4 cm; sori orbicular, 1 at base of pinnule (Themelium tenuisectum).   37 Themelium
       
10 (7) Sporangia acrostichoid, covering most of lamina surface   (11)
+ Sporangia in discrete sori or coenosori, not covering lamina surface   (13)
       
11 (10) Sporangia restricted to a distinctly narrowed apical extension of lamina (Belvisia).   19 Lepisorus
+ Sporangia covering whole of fertile lamina   (12)
       
12 (11) Lamina deeply 3-lobed or pinnatilobate.   6 Christopteris
+ Lamina entire.   25 Leptochilus
       
13 (10) Lamina hastate or palmately to pedately lobed   (14)
+ Lamina entire, pinnately lobed, or pinnate   (16)
       
14 (13) Scales opaque; paraphyses absent.   7 Selliguea
+ Scales entirely or partly clathrate; paraphyses present   (15)
       
15 (14) Lamina palmately 3-10-lobed, 12-30 cm; sori elongated.   16 Neocheiropteris
+ Lamina hastate to pedately 3-5-lobed, 4-7 cm; sori suborbicular (Lepisorus waltonii).   19 Lepisorus
       
16 (13) Stipe and/or lamina margins stiffly hirsute, lamina surface often with forked or glandular hairs; spores green   (17)
+ Stipe and/or lamina margins glabrous or scaly, less often pubescent; spores brown, yellow, or whitish (greenish at time of dispersal in Loxogramme)   (28)
       
17 (16) Lamina simple   (18)
+ Lamina pinnately lobed to bipinnatifid   (20)
       
18 (17) Sori linear, sunken in 2 grooves, parallel to midrib.   28 Scleroglossum
+ Sori orbicular to narrowly oblong, oblique to midrib, usually superficial or slightly sunken in lamina, rarely deeply sunken   (19)
       
19 (18) Rhizomes dorsiventral.   29 Oreogrammitis
+ Rhizomes radial.   30 Radiogrammitis
       
20 (17) Veins in pinnae simple or forked, sori 1(or 2) on each pinna; rhizomes radial   (21)
+ Veins in pinnae pinnately branched, sori 1 to several on each pinna; rhizomes radial or dorsiventral   (23)
       
21 (20) Sori protected by folded pinnae.   31 Calymmodon
+ Sori not protected by folded pinnae   (22)
       
22 (21) Frond hairs medium to dark reddish brown, simple, all or most hairs more than. 5 mm, up to 1.8 mm.   32 Micropolypodium
+ Frond hairs pale, simple or 1- or 2-forked, less than 0.5 mm.   33 Xiphopterella
       
23 (20) Pale yellowish brown simple glandular hairs and forked hairs with glandular branches,. 1-0.2 mm, on stipe and lamina, sometimes also on rhizomes scales; no other types of hairs present.   34 Chrysogrammitis
+ Pale yellowish brown simple glandular hairs and forked hairs with glandular branches,. 1-0.2 mm, absent from stipe, lamina, and rhizome scales; other types of hairs present   (24)
       
24 (23) Sori submarginal or marginal, or on abaxial surface of lamina, usually deeply sunken in lamina, sometimes superficial or very slightly sunken on abaxial surface of lamina; rhizomes dorsiventral, stipe articulate to rhizome, phyllopodia present; rhizome scales subclathrate to clathrate, with marginal hairs.   35 Prosaptia
+ Sori on abaxial surface of lamina, superficial or very slightly sunken; rhizomes radial or dorsiventral, stipe sometimes articulate to rhizome, phyllopodia sometimes present; rhizome scales sometimes subclathrate to clathrate, sometimes with marginal hairs   (25)
       
25 (24) Rhizomes dorsiventral, rhizome scales glabrous   (26)
+ Rhizomes radial, all or most rhizome scales with hairs at apex and/or margin   (27)
       
26 (25) Hairs on stipe up to 0.4 mm.   36 Ctenopterella
+ Hairs on stipe up to 2 mm.   37 Themelium
       
27 (25) Vein endings without hydathodes on adaxial surface of lamina, frond hairs dark reddish brown.   38 Dasygrammitis
+ Vein endings with hydathodes on adaxial surface of lamina, frond hairs pale to medium reddish brown.   39 Tomophyllum
       
28 (16) Sori or coenosori elongate to linear   (29)
+ Sori orbicular to elliptic, occasionally in confluent pairs   (33)
       
29 (28) Sori parallel to costa in long coenosori, often discontinuous   (30)
+ Sori discrete, at angle to costa and parallel to lateral veins   (31)
       
30 (29) Rhizome relatively stout, (1-)1.5-5 mm in diam., densely scaly, with closely spaced fronds; fronds monomorphic, usually with dark, clathrate scales.   19 Lepisorus
+ Lamina usually dimorphic, without scales.   20 Lemmaphyllum
       
31 (29) Rhizome scales basifixed; spores greenish at time of dispersal.   1 Loxogramme
+ Rhizome scales pseudopeltate or peltate; spores brown or whitish   (32)
       
32 (31) Lamina leathery; rhizome scales opaque; spores brown (Selliguea feei).   7 Selliguea
+ Lamina herbaceous; rhizome scales clathrate; spores whitish.   25 Leptochilus
       
33 (28) Lamina entire   (34)
+ Lamina pinnately divided or pinnate   (42)
       
34 (33) Plants climbing; rhizome scales bearing a tuft of long, stiff, needlelike, reddish brown hairs abaxially near their center.   17 Tricholepidium
+ Plants epiphytic, terrestrial, or climbing; rhizome scales glabrous, occasionally bearing a tuft of short, soft, brown hairs adaxially at point of attachment   (35)
       
35 (34) Lamina with many conspicuous reddish linear scales on both surfaces.   21 Caobangia
+ Lamina with scales absent or short and inconspicuous   (36)
       
36 (35) Sori arranged in 1 line on each side of costa, discrete or sometimes ?merging into longitudinal coenosori   (37)
+ Sori scattered or arranged into 2-4 ?straight lines, never merging into coenosori   (39)
       
37 (36) Rhizome scales opaque.   7 Selliguea
+ Rhizome scales clathrate   (38)
       
38 (37) Rhizome relatively stout, (1-)1.5-5 mm in diam., densely scaly, with closely spaced fronds; fronds monomorphic, herbaceous or papery.   19 Lepisorus
+ Rhizome slender, up to 1.5 mm in diam., sparsely scaly, with widely spaced fronds; fronds often dimorphic (monomorphic in Lemmaphyllum rostratum), fleshy to thinly leathery.   20 Lemmaphyllum
       
39 (36) Paraphyses absent   (40)
+ Paraphyses present   (41)
       
40 (39) Fronds articulate; rhizome scales peltate, apex obtuse (Phymatosorus lanceus).   22 Phymatosorus
+ Fronds not articulate; rhizome scales pseudopeltate, apex acute.   24 Microsorum
       
41 (39) Plants terrestrial; sori arranged into (1 or)2-4 lines on each side of midrib.   18 Neolepisorus
+ Plants climbing; sori scattered over lamina, not in defined lines.   23 Lepidomicrosorium
       
42 (33) Lamina pinnate, at least lower part of rachis terete   (43)
+ Lamina pinnatisect to pinnatifid   (46)
       
43 (42) Rhizome scales opaque.   5 Arthromeris
+ Rhizome scales clathrate   (44)
       
44 (43) Lateral pinnae articulate to rachis.   11 Goniophlebium
+ Lateral pinnae not articulate at base   (45)
       
45 (44) Frond not articulate at base; young sori with paraphyses present; pinna margins toothed or crenate.   13 Polypodiastrum
+ Frond articulate at base; paraphyses absent; pinna margins entire (Phymatosorus cuspidatus).   22 Phymatosorus
       
46 (42) Veins free, without areoles along costa   (47)
+ Veins anastomosing to form areoles along costa or rachis   (48)
       
47 (46) Veinlets pinnate; scales on rhizome thick, not clathrate.   26 Polypodium
+ Veinlets forked; scales on rhizome thin, pellucid, clathrate.   12 Metapolypodium
       
48 (46) Lamina with thick multicellular hairs on both surfaces; sporangium setose.   15 Himalayopteris
+ Lamina glabrous, finely pubescent, or minutely scaly; sporangium glabrous   (49)
       
49 (48) Rhizome scales opaque.   7 Selliguea
+ Rhizome scales clathrate   (50)
       
50 (49) Frond not articulate at base; young sori with paraphyses present; pinna margins toothed or serrate.   14 Polypodiodes
+ Frond articulate at base; paraphyses absent; pinna margins entire.   22 Phymatosorus

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