PLAGIOGYRIACEAE [Draft]

瘤足蕨科  liu zu jue ke

Zhang Xianchun (张宪春)[1]; Hans P. Nooteboom[2]

Ferns terrestrial, small to medium-sized, 10–150(–200) cm tall, without scales. Rhizomes dictyostelic, erect, suberect, or short-decumbent, stout, thickened; stipe bases persistent. Roots blackish, sparsely branched, wiry, regularly arranged on stem, one beside each stipe base. Fronds dimorphic, bearing minute, uniseriate, multicellular, gland-tipped, mucilage-secreting trichomes. Stipes round, ovate, triangular, or tetragonal in cross section, bases swollen; vascular bundle 1 per stipe, expanded or divided into three meristeles in enlarged base; rachises winged or not, adaxially sulcate, abaxially semiterete, flat, or carinate. Aerophores variously present on stipes and rachises, cushion like or elongate and hornlike with blunt apex. Sterile fronds herbaceous, firmly so, or subleathery. Lamina 1-pinnate, pinnatifid, or pinnatilobed, apices coadunate, pinnalike, pinnatilobed, or confluent. Pinnae sessile, short-stalked, or adnate to rachises, falcate to linear-lanceolate, widest near base or below middle, glabrous or abaxially sparsely glandular-pubescent, margins entire to serrulate, apices minutely serrulate to dentate; veins simple or paired at base, mostly 1- or 2- forked, reaching pinna margins, distinct on both surfaces. Fertile fronds erect in middle of plant, with proportionally longer stipes and shorter lamina. Lamina pinnate or deeply pinnatifid. Pinnae sessile, short-stalked, or adnate to rachises; apical pinna usually pinnatifid or pinnalike, margins entire, subentire, or erosely toothed, glabrous or with few short glandular hairs, scarious; tissue above the veinlets hygroscopic and reversible, functioning as indusium (“false indusium”), scarious margins covering sori in juvenile stage, recurved at maturity; veins free, 1- or 2-forked, solitary or paired at base. Sori borne on swollen vein tips, extending slightly, confluent, ascrostichoid, almost wholly covering abaxial surface of pinnae when ripe. Paraphyses numerous, mixed with sporangia, uniseriate, orange to dark brown, multicellular, glandlike, slightly longer than sporangia, apically subcapitate. Sporangia with a complete oblique annulus, not interrupted by stalk, laterally dehiscent. Spores 64 per sporangium, tetrahedral; aperture trilete; arms of aperture ca. 3/4 radius; surface ornamentation irregular, coarse, tuberculate with coalescent papillae and rodlets. x = 65 (sexual)

One genus and ca. 10 species: mainly in E and SE Asia, one species in tropical America: eight species (one endemic) in China.

Ching Ren-chang, Fu Shu-hsia, Wang Chu-hag & Shing Gung-hsia. 1959. Plagiogyriaceae. In: Chien Sung-shu & Chun Woon-young, ed., Fl. Reipubl. Popularis Sin. 2: 88–105.

1. PLAGIOGYRIA (Kunze) Mettenius, Abh. Senckenberg. Naturf. Ges. 2: 268. 1858.

瘤足蕨属  liu zu jue shu

Morphological characters and geographic distribution are the same as those of the family.

1a. Sterile lamina pinnate, most pinnae cuneate, rounded, truncate, or attenuate at base.

2a. Sterile lamina glaucous abaxially      2. P. glauca

2b. Sterile lamina not glaucous abaxially.

3a. Aerophores on stipes and rachises elongate and hornlike with blunt apex      1. P. pycnophylla

3b. Aerophores on stipes and rachises cushion like

4a. Veins usually simple, or as many simple as forked    3. P. euphlebia

4b. Veins usually paired at base or forked above base  4. P. japonica

1b. Sterile lamina pinnate to pinnatifid or pinnatilobed, pinnae usually adnate to rachises.

5a. Sterile lamina glaucous abaxially      5. P. assurgens

5b. Sterile lamina not glaucous abaxially.

6a. Rachises of sterile lamina abaxially semiterete to trigonous    1. P. adnata

6b. Rachises of sterile lamina abaxially carinate or flattened

7a. Stipes distally tetragonal in cross section      7. P. stenoptera

7b. Stipes distally triangular in cross section       8. P. falcata

1. Plagiogyria pycnophylla (Kunze) Mettenius, Abh. Senckenberg. Naturf. Ges. 2: 272, t. 4, f. 22. 1858.

密叶瘤足蕨  mi ye liu zu jue

Lomaria pycnophylla Kunze, Bot. Zeitung (Berlin) 6: 143. 1848; Plagiogyria coerulescens Ching; P. communis Ching; P. decrescens Ching; P. gigantea Ching; P. glauca var. virescens C. Christensen; P. lanuginosa Ching; P. lineata Ching; P. minguingensis R. D. Dixit & A. Das; P. pycnophylla var. mixta Copeland; P. pycnophylla var. remota H. Christ; P. scandens Mettenius; P. simulans Ching; P. taliensis Ching; P. tetraptera W. M. Chu & J. J. He; P. virescens (C. Christensen) Ching; P. wilhelmensis Nakaike.

Stipes 6–60 cm on sterile fronds, 10–70 cm on fertile fronds, distally tetragonal, terete, ovate, or triangular in cross-section; vascular bundle U-shaped; stipes and rachises glabrous to hairy, sometimes densely hairy; aerosphere elongate and hornlike with blunt apex. Sterile lamina pinnate, not glaucous, 15–100 Χ 6–40 cm, apically with a pinnatilobed segment or a pinnalike segment; rachises abaxially flattened, semiterete, or grooved. Pinnae 20–50 pairs; proximal pinnae sessile to short-stalked, bases truncate or rounded, subdeflexed or not, gradually abbreviated or slightly shorter, some becoming vestigial on stipes; middle pinnae sessile, base truncate or rounded; veins simple or some 1-forked. Fertile lamina pinnate, 20–70 Χ 4–20 cm; rachises abaxially flattened, semiterete, or grooved, sometimes winged. Pinnae 16–40 pairs, sessile or short-stalked; proximal pinnae gradually abbreviated or some pairs aerosphere like or vestigial. Paraphyses few, shed early, brown or dark brown. Spores yellow to brown, not obviously tuberculate.

Forests, wet areas; 1200–3500 m. Sichuan, Xizang, Yunnan [Bhutan, E India, Indonesia, Malaysia (including Borneo), N Myanmar, Nepal, New Guinea, Philippines (Luzon), Sikkim].

2. Plagiogyria glauca (Blume) Mettenius, Abh. Senckenberg. Naturf. Ges. 2: 273. 1858.

粉背瘤足蕨  fen bei liu zu jue

Lomaria glauca Blume, Enum. Pl. Javae, 204. 1828; Blechnum papuanum Brause; Plagiogyria brausei Nakaike; P. distanta R. D. Dixit & A. Das; P. formosana Nakai; P. formosana var. angustata Nakaike; P. glauca subsp. formosana (Nakai) Nakaike; P. glauca var. nana (Copeland) C. Christensen; P. glauca var. philippinensis H. Christ; P. glaucescens Ching; P. glaucescens var. arguta Ching; P. media Ching; P. nana Copeland; P. papuana (Brause) Alston (1939), not C. Christensen (1937); Struthiopteris papuana (Brause) Ching.

Stipes 3–55 cm on sterile fronds, 5–75 cm on fertile fronds, distally tetragonal, round, or ovate in cross section; vascular bundle U-shaped; stipes and rachises glabrous to hairy, sometimes densely hairy; aerosphere elongate and hornlike with blunt apex. Sterile lamina pinnate, glaucous abaxially, 7–110 Χ 2–30 cm, apically with a pinnatilobed segment or a pinnalike segment; rachises abaxially semiterete to flattened, or grooved. Pinnae 13–43 pairs; proximal pinnae sessile to short-stalked, base truncate or rounded, not deflexed to subdeflexed, abruptly abbreviated or not, some becoming vestiges on stipes; middle pinnae sessile, base truncate or rounded; veins as many simple as forked. Fertile lamina pinnate, 11–77 Χ 4–24 cm; rachises abaxially semiterete, flattened, or grooved. Pinnae 13–43 pairs, short-stalked or sessile; proximal pinnae gradually abbreviated or some pairs becoming aerosphere like vestiges. Paraphyses few, shed early, dark brown or yellow. Spores yellow to brown, not obviously tuberculate.

Grasslands, lightly shaded or open places; 1200–3800 m. Taiwan, Xizang, Yunnan [E India, Indonesia, Malaysia (Borneo), N Myanmar, New Guinea, Philippines (Luzon, Mindanao); SW Pacific Islands (S New Ireland, Solomon Islands)].

3. Plagiogyria euphlebia (Kunze) Mettenius, Abh. Senckenberg. Naturf. Ges. 2: 274. 1858.        

华中瘤足蕨  hua zhong liu zu jue

Lomaria euphlebia Kunze, Bot. Zeitung (Berlin) 6: 521. 1848; Lomariopsis triquetra (Wallich ex Mettenius) Ettingshausen; Plagiogyria attenuata Ching; P. chinensis Ching; P. christii Copeland; P. elongata R. D. Dixit & A. Das; P. euphlebia var. grandis (Copeland) De Vol; P. euphlebia var. triquetra (Wallich ex Mettenius) Ching; P. grandis Copeland; P. integripinna Ching; P. koidzumii Tagawa; P. maxima C. Christensen; P. triquetra Wallich ex Mettenius; Stenochlaena henryi H. Christ.

Stipes (7–)25–35(–70) cm on sterile fronds, (9–)40–75 cm on fertile fronds, distally ovate or tetragonal, rarely round, in cross section; vascular bundle V-shaped to U-shaped; stipes and rachises glabrous to hairy, sometimes densely hairy; aerosphere cushion like. Sterile lamina pinnate, not glaucous, (11–)20–60(–135) Χ (7–)12–30(–40) cm, apically with a pinnalike segment; rachises abaxially usually semiterete. Pinnae 7–25 pairs; proximal pinnae short-stalked or rarely sessile, base usually equilateral, cuneate, rounded, or attenuate, not deflexed, not abbreviated or basal pair slightly shorter; middle pinnae short-stalked or sessile, base cuneate or rounded; veins mostly simple to as many simple as forked. Fertile lamina pinnate, (9–)20–50(–80) Χ (3–)10–20(–33) cm; rachises abaxially semiterete, rarely flattened or grooved. Pinnae 7–25 pairs, short-stalked or sessile; proximal pinnae not abbreviated or basal pair slightly shorter. Paraphyses many, brown. Spores yellow or brown, with reddish tubercles. 2n = 260.

Forests; near sea level to 2500 m, usually 600–1500 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [India, Japan, Myanmar, Nepal, Philippines, Vietnam].

4. Plagiogyria japonica Nakai, Bot. Mag. (Tokyo) 42: 206. 1928.

华东瘤足蕨  hua dong liu zu jie

Plagiogyria adnata (Blume) Beddome var. distans Rosenstock; P. caudifolia Ching; P. hainanensis Ching; P. intermedia Copeland; P. japonica var. pseudojaponica (Nakaike) K. Iwatsuki; P. liankwangensis Ching; P. pseudojaponica Nakaike.

Stipes (7–)20–40 cm on sterile fronds, 40–55 cm on fertile fronds, distally tetragonal, round, or ovate in cross section; vascular bundle V-shaped; stipes and rachises glabrous; aerosphere cushion like, present only on stipe base. Sterile lamina pinnate, not glaucous, (12–)20–43 Χ (4–)10–20 cm, apically with a pinnalike segment; rachises abaxially semiterete, flattened, or grooved. Pinnae (8–)12–20(–30) pairs; proximal pinnae sessile, base usually inequilateral, rounded, or cuneate, not deflexed to subdeflexed, slightly shortened; middle pinnae short-stalked, sessile, or rarely acroscopically adnate to rachises, base rounded or cuneate; distal pinnae inequilaterally adnate to rachises, basiscopically decurrent to rounded; veins mostly paired at base or forked above base. Fertile lamina pinnate, 23–50 Χ 6–17 cm; rachises abaxially flattened or semiterete. Pinnae 9–19 pairs, short-stalked; proximal pinnae longest. Paraphyses many, brown or yellow. Spores yellow, with reddish tubercles. 2n = 260.

Forests, ravines, rocky places; 100–1800 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [Japan, Korea (Cheju Island)].

5. Plagiogyria assurgens H. Christ, Bull. Soc. Bot. Ital. 1901: 293. 1901.

峨嵋瘤足蕨 e mei liu zu jue

Blechnum faberi C. Christensen; Lomaria deflexa Baker (1888), not Col.?? (1844); Spicanta deflexa (??) Kuntze.

Stipes 8–20 cm on sterile fronds, 15–45 cm on fertile fronds, distally tetragonal in cross section; vascular bundle U-shaped; stipes and rachises glabrous; aerosphere cushion like, present only on stipe bases. Sterile lamina pinnatifid, glaucous abaxially, 23–65 Χ 10–20 cm, apex confluent, terminating in a pinnatilobed segment; rachises abaxially trigonous or carinate. Pinnae 22–47 pairs; proximal pinnae adnate to rachises, deflexed, abruptly abbreviated; middle pinnae acroscopically adnate to rachises, basiscopically rounded; distal pinnae subequilaterally adnate to rachises, basiscopically rounded; veins usually paired at base. Fertile lamina deeply pinnatifid, 28–60 Χ 7–14 cm; rachises abaxially carinate. Pinnae 20–35 pairs, base broadly adnate to rachises; proximal 1–4 pairs of pinnae much shorter than others, deflexed, often sterile. Paraphyses few, shed early, brown. Spores yellow, with reddish tubercles.

* Humus-rich forests, mountain slopes; 1200–2500 m. Chongqing, Sichuan, Yunnan.

6. Plagiogyria adnata (Blume) Beddome, Ferns Brit. India 1: t. 51. 1865.

瘤足蕨 liu zu jue

Lomaria adnata Blume, Enum. Pl. Javae, fasc. 2: 205. 1828; L. brooksii Alderwerelt van Rosenburgh; Plagiogyria adnata var. condensata H. Christ; P. adnata var. reflexa C. Christensen & Tardieu; P. adnata var. yakushimensis (K. Satτ) K. Iwatsuki; P. distinctissima Ching; P. meghalayensis R. D. Dixit & A. Das; P. rankanensis Hayata; P. rankanensis var. yakushimensis (K. Satτ) Nakaike; P. subadnata Ching; P. wulingshanensis C. M. Zhang & S. F. Wu; P. yakumonticola Nakaike; P. yakushimensis K. Satτ; P. yunnanensis Ching; Struthiopteris brooksii (Alderwerelt van Rosenburgh) Ching.

Stipes (4.5–)15–25(–35) cm on sterile fronds, (17–)40–50(–65) cm on fertile fronds, distally tetragonal or rarely triangular in cross section; vascular bundle V-shaped; stipes and rachises glabrous; aerosphere cushion like, present only on stipe bases. Sterile lamina pinnatifid, not glaucous, (8–)15–25(–36) Χ (3–)5–15(–26) cm, apex confluent, terminating in a pinnatilobed segment; rachises abaxially semiterete to trigonous. Pinnae (8–)13–20(–35) pairs; proximal pinnae adnate to rachises, not deflexed to subdeflexed, not abbreviated or slightly shortened; middle pinnae acroscopically adnate to rachises, basiscopically rounded; distal pinnae subequilaterally adnate to rachises, basiscopically decurrent; veins mostly paired at base, forked above base, or rarely simple. Fertile lamina pinnate, (8–)16–26(–45) Χ (2–)4–10(–16) cm; rachises abaxially flattened or grooved. Pinnae (9–)13–17(–24) pairs, sessile or short-stalked; proximal pinnae short-stalked, not abbreviated. Paraphyses many or few, rarely shed early, yellow or brown. Spores yellow, with reddish tubercles. 2n = 260.

Forests, wet areas, mountain slopes; near sea level to 2000 m, common above 500 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [E India, Indonesia (Java, Sumatra), Japan, Malaysia (including Borneo), Myanmar, Philippines (Luzon), Thailand, Vietnam].

Plagiogyria japonica and P. adnata hybridize. Specimens of P. japonica with acroscopically adnate pinnae may key to P. adnata. Fronds of P. japonica usually end in a segment similar to a short lateral pinna, while fronds of P. adnata have pinnatifid apices.

7. Plagiogyria stenoptera (Hance) Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 282. 1899.

耳形瘤足蕨  er xing liu zu jue

Blechnum stenopterum Hance, J. Bot. 21: 268. 1883; Lomaria stenoptera (Hance) Baker; Plagiogyria argutissima H. Christ; P. henryi H. Christ; P. petelotii Copeland.

Stipes (3–)4–8(–12) cm on sterile fronds, (9–)11–22(–28) cm on fertile fronds, distally tetragonal in cross section; vascular bundle V-shaped; stipes and rachises glabrous; aerosphere cushion like, present only on stipe bases. Sterile lamina pinnatifid, not glaucous, 20–60 Χ 4–20 cm, apex confluent, terminating in a pinnatilobed segment; rachises abaxially flattened or carinate distally, sometimes with crisp wings. Pinnae 25–35 pairs; proximal pinnae reduced to auricles, middle pinnae adnate to rachises, basiscopically straight or rounded in large fronds; distal pinnae subequilaterally adnate to rachises, basiscopically decurrent; veins mostly forked above base. Fertile lamina pinnate, (10–)35–55(–75) Χ 4–8(–12) cm; rachises abaxially flattened to grooved, sometimes winged. Pinnae 15–30 pairs, subsessile or short-stalked; some proximal pinnae becoming aerosphere like vestiges or replaced by auriculate pinnae. Paraphyses absent. Spores yellow, with reddish tubercles.

Densely forested mountains, moist ravines, rocky slopes, 500–2500 m. Guangxi, Guizhou, Hubei, Hunan, Sichuan, Taiwan, Yunnan [Japan (Yakushima), Philippines (Luzon), Vietnam].

"Plagiogyria stenoptera var. major Ching" (Acta Phytotax. Sin. 7: 152.1958) belongs in synonymy here, however, it was not validly published as no type specimen was designated.

 

8. Plagiogyria falcata Copeland, Philipp. J. Sci. 2: 133. 1907.

镰羽瘤足蕨 liang yu liu zu jue

Lomaria matthewii H. Christ ex C. G. Matthew; Plagiogyria adnata (Blume) Beddome var. angustata Rosenstock; P. angustipinna Ching; P. chekiangensis P. L. Chiu; P. dentimarginata J. F. Cheng; P. dunnii Copeland; P. hayatana Makino; P. tenuifolia Copeland.

Stipes (6–)12–20(–30) cm on sterile fronds, 20–45 cm on fertile fronds, distally sharply triangular in cross section; vascular bundle V-shaped; stipes and rachises glabrous; aerosphere cushion like, present only on stipe bases. Sterile lamina pinnatifid, not glaucous, (10–)20–40(–55) Χ (3–)5–13(–17) cm, apex confluent, terminating in a pinnatilobed segment; rachises abaxially carinate. Pinnae (12–)35–55 pairs; proximal pinnae adnate to rachises, usually deflexed, not abbreviated or abruptly abbreviated; middle pinnae adnate to rachises, basiscopically straight or rounded; distal pinnae subequilaterally adnate to rachises, basiscopically straight or rounded; veins paired at base and forked above, paired or branched at base with both branches forked above, or rarely mostly simple. Fertile lamina deeply pinnatifid, 20–50 Χ 3–6(–10) cm; rachises abaxially carinate, sometimes winged. Pinnae (15–)35–50 pairs, sessile, broadly adnate to rachises; proximal (lower) pinnae not abbreviated or gradually abbreviated, sometimes not well developed. Paraphyses few, often shed early or absent, yellow. Spores yellow, with reddish tubercles.

Densely forested ravines, under shady rocks; near seal level to 2200 m, usually 500–1500 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Jiangxi, Taiwan, Zhejiang [Philippines].


[1] Herbarium, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China.

[2] Rijksherbarium/Hortus Botanicus, P.O. Box 9514, 2300 RA Leiden, The Netherlands.