Flora of China
  FOC Vol.2-3 (2013)              | Family List  Polypodiaceae  PDF     | Link Font:+ Big | - Small
   39. Polypodiaceae  
水龙骨科   shui long gu ke
Authors: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


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.

1Fronds 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    石韦属
+Fronds strongly dimorphic, basal fronds with basal part tightly adpressed to substrate, foliage/fertile fronds dichotomously lobed.10  Platycerium    鹿角蕨属
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    槲蕨属
+Fronds monomorphic, fronds with broad humus-collecting base and distal deeply pinnatisect fertile portion.2  Aglaomorpha    连珠蕨属
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    槲蕨属
+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    顶育蕨属
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    雨蕨属
+Fronds 2-pinnatifid, hairy(9)
9(8)Lamina 1.5-8 ?0.5-1.5 cm; sori linear-oblong, along veins.27  Pleurosoriopsis    睫毛蕨属
+Lamina 5-25 ?2-4 cm; sori orbicular, 1 at base of pinnule (Themelium tenuisectum).37  Themelium    蒿蕨属
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    瓦韦属
+Sporangia covering whole of fertile lamina(12)
12(11)Lamina deeply 3-lobed or pinnatilobate.6  Christopteris    戟蕨属
+Lamina entire.25  Leptochilus    薄唇蕨属
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    修蕨属
+Scales entirely or partly clathrate; paraphyses present(15)
15(14)Lamina palmately 3-10-lobed, 12-30 cm; sori elongated.16  Neocheiropteris    扇蕨属
+Lamina hastate to pedately 3-5-lobed, 4-7 cm; sori suborbicular (Lepisorus waltonii).19  Lepisorus    瓦韦属
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    革舌蕨属
+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    滨禾蕨属
+Rhizomes radial.30  Radiogrammitis    辐禾蕨属
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    荷包蕨属
+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    锯蕨属
+Frond hairs pale, simple or 1- or 2-forked, less than 0.5 mm.33  Xiphopterella    剑羽蕨属
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    金禾蕨属
+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 sc35  Prosaptia    穴子蕨属
+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 hair(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    小蒿蕨属
+Hairs on stipe up to 2 mm.37  Themelium    蒿蕨属
27(25)Vein endings without hydathodes on adaxial surface of lamina, frond hairs dark reddish brown.38  Dasygrammitis    毛禾蕨属
+Vein endings with hydathodes on adaxial surface of lamina, frond hairs pale to medium reddish brown.39  Tomophyllum    裂禾蕨属
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    瓦韦属
+Lamina usually dimorphic, without scales.20  Lemmaphyllum    伏石蕨属
31(29)Rhizome scales basifixed; spores greenish at time of dispersal.1  Loxogramme    剑蕨属
+Rhizome scales pseudopeltate or peltate; spores brown or whitish(32)
32(31)Lamina leathery; rhizome scales opaque; spores brown (Selliguea feei).7  Selliguea    修蕨属
+Lamina herbaceous; rhizome scales clathrate; spores whitish.25  Leptochilus    薄唇蕨属
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    毛鳞蕨属
+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    高平蕨属
+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    修蕨属
+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    瓦韦属
+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    伏石蕨属
39(36)Paraphyses absent(40)
+Paraphyses present(41)
40(39)Fronds articulate; rhizome scales peltate, apex obtuse (Phymatosorus lanceus).22  Phymatosorus    瘤蕨属
+Fronds not articulate; rhizome scales pseudopeltate, apex acute.24  Microsorum    星蕨属
41(39)Plants terrestrial; sori arranged into (1 or)2-4 lines on each side of midrib.18  Neolepisorus    盾蕨属
+Plants climbing; sori scattered over lamina, not in defined lines.23  Lepidomicrosorium    鳞果星蕨属
42(33)Lamina pinnate, at least lower part of rachis terete(43)
+Lamina pinnatisect to pinnatifid(46)
43(42)Rhizome scales opaque.5  Arthromeris    节肢蕨属
+Rhizome scales clathrate(44)
44(43)Lateral pinnae articulate to rachis.11  Goniophlebium    棱脉蕨属
+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    拟水龙骨属
+Frond articulate at base; paraphyses absent; pinna margins entire (Phymatosorus cuspidatus).22  Phymatosorus    瘤蕨属
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    多足蕨属
+Veinlets forked; scales on rhizome thin, pellucid, clathrate.12  Metapolypodium    篦齿蕨属
48(46)Lamina with thick multicellular hairs on both surfaces; sporangium setose.15  Himalayopteris    锡金假瘤蕨属
+Lamina glabrous, finely pubescent, or minutely scaly; sporangium glabrous(49)
49(48)Rhizome scales opaque.7  Selliguea    修蕨属
+Rhizome scales clathrate(50)
50(49)Frond not articulate at base; young sori with paraphyses present; pinna margins toothed or serrate.14  Polypodiodes    水龙骨属
+Frond articulate at base; paraphyses absent; pinna margins entire.22  Phymatosorus    瘤蕨属
   List of lower taxa