THELYPTERIDACEAE (in part) [DRAFT]

6. PHEGOPTERIS (C. Presl) Fιe, Gen. Fil. 242. 1852, emend. Ching in Acta Phytotax. Sin. 8: 312. 1963.

卵果蕨属  luan guo jue shu

Polypodium [par.] 2 Phegopteris C. Presl, Tent. Pterid. 179. 1836; Dryopteris subg. Phegopteris (C. Presl) C. Christensen; Phegopteris sect. Euphegopteris H. Ito [??valid]; Polypodium sect. Phegopteris C. Presl[??is this the same as the basionym]; Thelypteris sect. Phegopteris (C. Presl) K. Iwatsuki; Thelypteris subg. Phegopteris (C. Presl) Ching.

Plants mid- and small-sized, terrestrial. Rhizome long-creeping or short and erect, densely covered with brown scales and whitish acicular hairs. Fronds remote or clustered; stipe stramineous, shiny, slender, base scaly; scales brown, lanceolate and sparsely long hairy along margins; lamina bipinnatifid or pinnate-pinnatifid, ovate-triangular or narrowly lanceolate; pinnae connected to each other by a narrow wing along rachis, or proximal 1–3 pairs free, proximal pinnae shortened or basal pair only slightly shortened, or proximal several pairs gradually reduced to auricles; veins pinnate, lateral veins simple or forked, veinlets reaching margins; lamina herbaceous or soft papery, with whitish acicular hairs on both surfaces, rachis, costae, and costules, rounded and raised on both sides and similarly with dense acicular hairs, sometimes mixed with a few forked hairs, with more brownish hairs and lanceolate, ciliate scales abaxially. Sori round to oblong, borne above middle of ultimate veins, exindusiate or indusia very small and vestigial; sporangia often with a few short acicular hairs or capitate hairs near annulus. Spores bilateral, reniform, perispores winged, thin, and transparent, granulate on surfaces. x = 30.

Type species: Phegopteris polypodioides (Linnaeus) Fιe (= Polypodium phegopteris Linnaeus) = Phegopteris connectilis (Michaux) Watt.

Phegopteris is monophyletic, and differs from Pseudophegopteris in its smaller stature, stramineous stipes, laminae usually triangular or narrowly lanceolate, proximal pinna bases decurrent and adnate to each other by a rachis wing, veinlets reaching margin, sparsely ciliate scales on abaxial side of rachises, and costae sparsely ciliate. Phegopteris is primarily north-temperate and circumboreal, while Pseudophegopteris is tropical and subtropical, and restricted to the Paleotropics.

Four species; three species in China (one endemic).

1a.     Rhizome short, erect; lamina lanceolate, pinnae connected to each other by narrow wings; most pinnae below middle of blade gradually shortened, proximal pair of pinnae reduced to auricles  3. P. decursive-pinnata

1b.     Rhizome long creeping; lamina ± triangular, proximal 1–3 pairs of pinnae free, not tapering proximally, or basal pair slightly shortened, distal pinnae connected to each other by narrow rachis wing.

2a.    Lamina triangular, length and width subequal or length slightly longer than width, proximal pair of pinnae largest and often reflexed downward ..................................................................  1. P. connectilis

2b.    Lamina narrowly triangular, length almost 2 times width, proximal 1–3 pairs of pinnae free, proximal pair slightly shortened, spreading or slightly ascending ......................................................  2. P. tibetica

 

1. Phegopteris connectilis (Michaux) Watt, Canad. Naturalist Geol., n.s., 3: 29. 1866[??3(2): 159. 1866; issued ?1867].

卵果蕨  luan guo jue

Polypodium connectile Michaux, Fl. Bor.-Amer. 2: 271. 1803; Dryopteris phegopteris (Linnaeus) C. Christensen; Phegopteris polypodioides Fιe; P. vulgaris Mettenius; Polypodium phegopteris Linnaeus; Thelypteris phegopteris (Linnaeus) Slosson.

Plants 25–40 cm tall. Rhizome long creeping, with bright brown, ovate-lanceolate thin scales at apex. Fronds remote; stipe dark brown at base, stramineous distally, 15–30 cm, sparsely scaly, nearly smooth; lamina bipinnatifid, deltoid, 13–20 Χ 10–18 cm, acuminate and pinnatifid at apex. Pinnae ca. 10 pairs, usually opposite, spreading, lanceolate, 5–9 Χ 1–2 cm, basal pair largest, bases slightly or not tapering, free from second pair of pinnae, slightly deflexed, acuminate at apices; segments oblong, entire, undulate, or lobed along margins, rounded or obtuse at apices; distal pinnae gradually shortened, bases connected along rachises by triangular wings. Veins pinnate, lateral veins simple or occasionally forked. Fronds herbaceous or papery, drying gray-green or yellowish green, with sparse gray-white acicular hairs on both surfaces, ± with small scales along rachises and costae; scales brownish, ovate-lanceolate and ciliate along margins. Sori ovate-round or round, borne at or near ends of ultimate veins and close to margins. Sporangia with 1 or 2 hairs near annulus. 2n = 60. 90.

Forest understories; 1200–3100 m. Guizhou, Heilongjiang, Henan, Jilin, Liaoning, Shaanxi, Sichuan, Yunnan [Widely distributed in temperate regions of the northern hemisphere, south to mountains of central Asia and the Himalaya.

The type is from North America.

Phegopteris connectilis is similar to P. hexagonoptera (Michaux) Fιe, which occurs in North America only. In P. connectilis, the proximal pinnae taper to their bases, the proximal and subbasal pinnae are connected by decurrent wings along the rachis, the laminae are thinly herbaceous and sparsely shortly hairy on both surfaces, and the lateral veins are mostly simple. Phegopteris hexagonoptera has the proximal pair of pinnae connected to the next pair by a wing along the rachis, shorter laminar hairs (less than 0.25 mm vs. mostly 0.3–0.5 mm or longer in P. connectilis), and ultimate veins forked or pinnate.

2. Phegopteris tibetica Ching, Fl. Xizang. 1: 161. 1983.

西藏卵果蕨  xi zang luan guo jue

Plants ca. 45 cm tall. Rhizome not seen. Stipe dark brown at base, distally stramineous, 24–27 cm, with triangular-lanceolate scales, rounded abaxially, nearly smooth or with short hairs, grooved and with acicular hairs adaxially; lamina oblong, 18–20 Χ 10–14 cm, base not decurrent, pinnate-pinnatifid, apex acuminate and pinnatifid; pinnae 13–15 pairs, proximal 2 or 3 pairs free or slightly obliquely spreading, 5–7 Χ ca. 2 cm (basal one pair slightly shortened), oblong-lanceolate, slightly tapering to base, ± adnate to rachis, pinnatifid; segments oblong, entire along margins (or occasionally undulate-crenate on long segments), sparsely ciliate, rounded-obtuse at apices; pinnae above middle gradually shortened, linear-lanceolate, bases decurrent and connected to each other along rachis by narrow wings. Veins pinnate, lateral veins forked or simple and reaching margins. Fronds herbaceous, drying deep green, rachises raised abaxially, adaxially with grooves with dense acicular hairs, rachises and costae with acicular hairs on both sides and mixed with a few forked hairs, with sparse brownish narrowly lanceolate scales abaxially, scales sparsely long ciliate along margins, nearly glabrous between veins on both surfaces. Sori round or nearly so, borne subterminally on ultimate veins and close to margins, exindusiate. Sporangia each occasionally with 1 or 2 hairs near annulus.

* Understories of Abies forests; ca. 3600 m. Xizang (Bomi).

Phegopteris tibetica is most similar to P. connectilis, but the laminar length is 2 times the width, proximal 2 or 3 pairs of pinnae usually not connected at bases by a rachis wing, basal pair of pinnae slightly shortened, and laminae glabrous between veins on both surfaces.

3. Phegopteris decursive-pinnata (H. C. Hall) Fιe, Gen. Filic. 242. 1852 [??1850].

延羽卵果蕨  yan yu luan guo jue

Polypodium decursive-pinnatum H. C. Hall, Nieuwe Verhdl. Nederl. Inst. 5: 204. 1836; Aspidium decursive-pinnatum (H. C. Hall) Kunze; Dryopteris decursive-pinnata (H. C. Hall) Kuntze; Lastrea decurrens J. Smith; L. decursive-pinnata (H. C. Hall) J. Smith, [‘decursivo-pinnata’]; Nephrodium decursive-pinnatum (H. C. Hall) Baker, [decursivo pinnatum’]; Thelypteris decursive-pinnata (H. C. Hall) Ching.

Plants 30–60 cm tall. Rhizome short and erect, including base of stipe with reddish brown, ciliate, narrowly lanceolate scales. Fronds clustered; stipe stramineous, 10–25 cm; lamina lanceolate, 20–50 Χ 5–12 cm, gradually tapering to base, bipinnatifid or 1-pinnate and toothed along margins, acuminate and pinnatifid at apex; pinnae 20–30 pairs, alternate, obliquely spreading, middle ones largest, narrowly lanceolate, 2.5–6 Χ ca. 1 cm, bases broad and decurrent, connected by round auricles or triangular wings between pinnae, pinnatifid up to 1/3–1/2 of distance to costules, acuminate at apices; segments obliquely spreading, ovate-triangular, entire, obtuse at apices; pinnae gradually shortened to both ends, basal one pair of pinnae often shortened into small auricles. Veins pinnate, lateral veins simple and reaching margins. Fronds herbaceous, along rachis, costae, and veins with whitish unicellular acicular short hairs on both sides, abaxial sides with forked and stellate hairs, abaxial[??ok] sides of rachises and costae with sparse brownish, hairlike or lanceolate, ciliate scales. Sori nearly round, borne at or near ends of ultimate veins, 2 or 3 per segment, sometimes with bunched[??] stalked forked hairs in center when young. Sporangia each sometimes with 1 or 2 short hairs near annulus. 2n = 60, 90, 120.

Along rivers on fluvial plains, lower montane regions of hills, forest understories by roadsides; sea level up to 2000 m. Widely distributed in subtropical regions of China [rewrite as a list of provinces], to the North reaching S Henan and Qinling mountains of Shaanxi, East to the plains region of Taiwan, West to Sichuan, Guizhou and E-S and E, Yunnan [Japan, S Korea, N Vietnam].

Phegopteris decursive-pinnata is very different with other species of the genus in having short, erect rhizomes and narrowly lanceolate laminae, but can spread by long-creeping rhizomes. Because of these characters, Iwatsuki removed it from Thelypteris sect. Phegopteris and treated it and some species that we place in Pseudophegopteris, as well as other species, in Thelypteris sect. Lastrea. Three ploidal levels are known, but these numbers do not clearly correlate with differing morphologies.

 

8. PSEUDOPHEGOPTERIS Ching, Acta Phytotax. Sin. 16(4): 12. 1936 [??8: 313. 1963].

紫柄蕨属  zi bing jue shu

Dryopteris subg. Phegopteris Ching group 4[??okay]; Phegopteris sect. Lastrea H. Itτ; Thelypteris sect. Lastrella K. Iwatsuki.

Plants mid-sized, terrestrial. Rhizome short and erect, long-creeping to ascending, apex with brown lanceolate scales. Fronds clustered, sparse or remote; stipe castaneous or red-brown, less often stramineous or tan, shiny, base sometimes with whitish acicular hairs, sometimes mixed with stellate hairs, distally often glabrescent and smooth; lamina pinnate-pinnatifid to bipinnate-pinnatifid, lanceolate, oblong-lanceolate, oblong, or ovate, tapering or not to base, acuminate and pinnatifid at apex; pinnae usually opposite or subopposite, spreading or obliquely spreading, pinnae below middle adnate, sessile or shortly stalked, lanceolate or triangular-lanceolate, bases often broadened, truncate, symmetrical or not, sometimes hastate, acuminate at apices; basiscopic segments sometimes longer than acroscopic segments, and basal segments on both sides sometimes prolonged; costae raised on both sides, usually same color as stipes and rachises or lighter colored, abaxial sides glabrous or with whitish acicular hairs, hairy adaxially; veins free, lateral veins simple or forked, each with a clavate hydathode at end and not reaching margin. Sori oblong, ovate, or nearly round, borne at middle or above middle, exindusiate. Sporangia glabrous or with short hairs just below annulus. Spores bilateral, round-reniform, perispores thin and transparent, reticulate or foveolate on surfaces, exine smooth. x = 31.

About 25 species, tropical and subtropical Asia, east to the Pacific islands, to the west reaching to western Africa; 12 species in China (four endemic).

1a.     Stipes stramineous, bases including abaxial sides of rachises and costae with whitish acicular hairs mixed with irregularly forked or stellate hairs.

2a.    Stipes same size or slightly shorter than laminae, 15–30 cm; pinnae with sparse acicular hairs adaxially      1. P. levingei

2b.    Stipes equal to 1/4 of laminae in length, laminae 8–10 cm; pinnae except costae with sparse acicular hairs adaxially, otherwise glabrous .......................................................................  2. P. brevipes

1b.     Stipes red-castaneous or castaneous, if stramineous or brown-stramineous, then bases of stipe, abaxial sides of rachis and costae never with mixed acicular hairs.

3a.    Basal pair of pinnules or segments on proximal pinnae, particularly basiscopic one, clearly longer than adjacent one and also lobed or incised, pinna base asymmetrically hastate.

4a.     Laminae not tapering to base, i.e., proximal 1 or 2 pairs of pinnae largest; pinnae all sessile     3. P. yigongensis

4b.     Laminae ± tapering to base, i.e., proximal 1 or 2 pairs of pinnae smaller than distal ones.

5a.     Pinnae below middle shortly stalked ....................................................  4. P. tibetana

5b.     Pinnae all sessile.

6a.    Rhizome long-creeping; stipes nearly same length or slightly shorter than laminae; laminae papery, drying brownish green, glabrous on both surfaces ...............................  5. P. aurita

6b.    Rhizome shortly creeping to ascending; stipes much shorter than laminae; laminae thinly herbaceous, drying yellowish green, with fine acicular hairs on both surfaces .....  6. P. subaurita

3b.    Basal pair of pinnules or segments on proximal pinnae same shape and size as distal pinnules/segments on same pinna, or at most slightly inflated; pinnae symmetrically hastate at base, or seemingly not hastate.

7a.     Stipes stramineous, occasionally tan; plants 90–120 cm tall; laminae 60–80 Χ 20–30 cm; segments toothed along margins ..................................................................................  12. P. microstegia

7b.     Stipes red-castaneous, castaneous-brown, or reddish brown, never stramineous nor brown-stramineous; segments entire along margins.

8a.     Laminae less than 20 cm wide, pinnate-pinnatifid.

9a.    Sori near end of acroscopic veinlet of a forked pair and closer to margin; sporangia lacking hairs; pinnae subglabrous on abaxial surfaces between veins .................  10. P. zayuensis

9b.    Sori at middle of ultimate veins, between costae and segment margins; sporangia with 1 or 2 seta(e) below annulus; pinnae with sparse fine acicular hairs on abaxial surfaces between veins      11. P. rectangularis

8b.     Laminae wider than 20 cm, pinnate-pinnatisect with pinnules often toothed or lobed, or 2-pinnate-pinnatifid.

10a.   Laminae broadly ovate, proximal pair of pinnae largest, 30–45 Χ 13–20 cm, pinnules pinnatifid nearly to costules ............................................................................  9. P. yunkweiensis

10b.   Laminae oblong or oblong-lanceolate, proximal pair of pinnae same size as ones above or slightly shorter, 10–20 Χ 2.5–7 cm; pinnules pinnatifid to 1/2 of distance to costule.

11a.   Rhizome long-creeping; laminae with sparse acicular short hairs or sometimes glabrous on abaxial surfaces; sporangia glabrous ............................................  7. P. pyrrhorhachis

11b.   Rhizome short, ascending; laminae with dense acicular short hairs on abaxial surfaces; sporangia each with 2 or more hairs ......................................................  8. P. hirtirachis

 

1. Pseudophegopteris levingei (C. B. Clarke) Ching, Acta Phytotax. Sin. 8: 314. 1963.

星毛紫柄蕨  xing mao zi bing jue

Gymnogramma levingei C. B. Clarke, Trans. Linn. Soc. London, Bot. 1: 95. 1880 [??Ann. Bot. (Oxford) 5: 483. 1891]; Dryopteris levingei (C. B. Clarke) C. Christensen; D. purdomii C. Christensen; Phegopteris levingei (C. B. Clarke) Tagawa; Thelypteris levingei (C. B. Clarke) Ching.

Plants 60–80 cm tall. Rhizome long-creeping, with reddish brown, broadly lanceolate scales and whitish acicular hairs. Fronds remote; stipe stramineous, 15–30 cm, with sparse similar scales on proximal parts and denser whitish acicular hairs and few irregularly forked stellate hairs; lamina pinnate-pinnatisect, lanceolate or oblong-lanceolate, 35–60 Χ 5–15 cm, slightly tapering at base, acuminate and pinnatifid at apex; pinnae up to ca. 20 pairs, opposite, sessile, pinnae above middle approximate, proximal 2 or 3 pairs remote from each other, gradually shortened downward, basal pair of pinnae smallest, 2–5 cm, others 3–8(–11) Χ 1.5–2.5 cm, lanceolate, bases truncate, subsymmetrical, pinnatisect nearly to costae, acuminate at apices; segments 8–15 pairs per pinna, opposite, obliquely spreading, approximate, oblong, 1–1.3 Χ 0.3–0.5 cm, slightly broader at base, connected to each other by a narrow wing, entire, crenate, or occasionally lobed along margins of larger segments, obtuse at apices. Veins visible on both sides, lateral veins simple or forked, 5–7 per segment, basal pair arising from above bases of main veins. Laminae herbaceous, drying dark green, abaxial surfaces with dense whitish acicular hairs and a few stellate short hairs along costae and veins, also ± hairy between veins, adaxial surfaces with sparse acicular hairs, hairs denser along rachises. Sori nearly round or oblong, 3–5(–7) pairs per segment, borne at or above middle of ultimate veins and closer to margin. Sporangia each with 1–3 hairs ca. 0.2 mm below annulus. 2n = 124.

Forest understories beside streams or in thickets; 1300–2900 m. SE Gansu, SC Shaanxi, C Sichuan, E and S Xizang, NW Yunnan [NE India, Kashmir].

The type is from Kashmir.

Pseudophegopteris levingei is similar to P. bukoensis (Tagawa) Holttum, but the latter is larger in outline, with laminae 2-pinnate-pinnatifid, pinnae and pinnules oblong, pinnules incised to 2/3 of distance to costules, and costae with more stellate hairs abaxially.

2. Pseudophegopteris brevipes Ching & S. K. Wu, Fl. Xizang. 1: 163. 1983.

短柄紫柄蕨  duan bing zi bing jue

Plants 50–60 cm tall. Rhizome long-creeping, including base of stipe with sparse whitish long hairs and reddish brown scales; scales ovate-lanceolate, thinly membranous, apices hairlike, hairy on back and margins. Fronds remote; stipe stramineous, 8–20 cm, with sparse whitish acicular and a few stellate hairs, hairs denser in grooves adaxially; lamina 40–45 Χ 12–16 cm, gradually tapering to base, pinnate-pinnatifid, pinnatifid and acuminate at apex; pinnae ca. 20 pairs, opposite or subopposite, spreading, sessile, proximal 2 pairs of pinnae slightly shortened, basal pair shortest, ovate-lanceolate, 3.5–5 cm, above ones lanceolate, 6–8 Χ ca. 2 cm, bases widest and subsymmetrical, truncate, pinnatisect nearly to costae, acuminate at apices and entire; segments up to ca. 12 pairs, approximate, spreading, oblong, 8–9 Χ ca. 4.5 cm, hairy along margins, obtuse at apices. Veins slender, visible adaxially, lateral veins forked, 4 or 5 pairs per segment, basal pair arising from above bases of costae. Laminae thinly herbaceous, drying deep green or dark brown-green, with sparse whitish hairs, hairs denser along costae and veins and mixed with a few irregularly forked stellate hairs, adaxial surfaces with sparse acicular hairs along costae, rachises stramineous, sparsely hairy. Sori oblong, 2–4 pairs per segment, borne near ends of forked veinlets and close to margins. Sporangia each with hairs below annulus.

* In rock crevices in valleys; ca. 2300 m. SE Xizang (Chayu).

3. Pseudophegopteris yigongensis Ching, Fl. Xizang. 1: 165. 1983.

易贡紫柄蕨  yi gong zi bing jue

Plants 60–75 cm tall. Rhizome long-creeping, with lanceolate hairy scales. Fronds remote; stipe blackish brown at base, brown-stramineous and shiny distally, 30–40 cm, bases villous, distally glabrous; lamina narrowly oblong or narrowly lanceolate, 25–35 Χ 6–10 cm, base not tapering, pinnate-pinnatifid (basal pinnules on proximal pair of pinnae often pinnate), pinnatifid and acuminate at apex; pinnae 9–12 pairs, opposite, spreading, bases decurrent and connected, middle pinnae ± decurrent and adnate to rachis but not connected to each other, proximal 1 or 2 pairs of pinnae largest, sessile and not decurrent, 7–9 Χ ca. 4.5 cm, bases asymmetrical, pinnatifid, long acuminate at apices; basiscopic pinna segments longer than acroscopic ones, particularly basal basiscopic one, up to 3–4 cm, pinnatifid, other segments entire. Veins visible on both sides, lateral veins simple or forked, 4–6 pairs per segment. Laminae thinly herbaceous, drying yellowish green, abaxially villous along veins, nearly glabrous adaxially, rachises stramineous and sparsely villous. Sori oblong, borne at middle or above middle of ultimate veinlets, 2–4 pairs per segment. Sporangia each with 2–4 acicular hairs below annulus.

* Understories of dense forests, ca. 2500 m. SE Xizang (Bomi, Yigong).

Pseudophegopteris yigongensis is somewhat similar to P. levingei, differing in the fewer lateral pinnae, proximal two pairs much larger than distal ones, and laminae sparingly villous abaxially.

4. Pseudophegopteris tibetana Ching & S. K. Wu, Fl. Xizang. 1: 164. 1983.

西藏紫柄蕨  xi zang zi bing jue

Plants 85–90 cm tall. Rhizome long-creeping, with sparse dark brown lanceolate scales. Fronds remote; stipe red-castaneous at base, shiny, 15–40 cm, base with a few scales and acicular hairs; lamina lanceolate, (15–) 45–50 Χ (4–)12–16 cm, slightly tapering proximally, pinnate-pinnatifid (pinnules on proximal pair of pinnae often lobed), pinnatifid and long acuminate at apex; pinnae 4–17 pairs, ± opposite or subopposite, pinnae above middle ± adnate to rachis, pinnae below middle shortly stalked, proximal 1 or 2 pairs of pinnae obliquely triangular, (2–)7–7.5 Χ 1.5–3 cm, bases asymmetrical, pinnatifid, acuminate at apices; basiscopic segments largest, oblong-lanceolate, ca. 2 Χ 0.4–1 cm, pinnatifid, other segments entire; distal pinnae lanceolate, up to 8 Χ 2–2.2 cm, symmetrical, pinnatifid to 1/2 distance to costae or less, acuminate at apices; segments oblong, entire. Veins visible abaxially, lateral veins forked or simple, 2–6 pairs per segment, basal pair arising from bases of costae. Fronds papery, drying yellowish green, abaxial surfaces with sparse hairs along veins and margins, rachises stramineous to brown, sparsely villous, hairs denser adaxially. Sori oblong, borne above middle of ultimate veinlets, 2–4 pairs per segment. Sporangia each with a seta below annulus.

* Understories of dense forests; ca. 2010 m. SE Xizang.

The distinctness of Pseudophegopteris tibetana relative to P. aurita and P. yigongensis needs more study.

5. Pseudophegopteris aurita (Hooker) Ching, Acta Phytotax. Sin. 8: 314. 1963.

耳状紫柄蕨  er zhuang zi bing jue

Gymnogramma aurita Hooker, Icon. Pl. 10: t. 974. 1854; Dryopteris distans (Hooker) Kuntze var. coreana H. Christ [??not in Tropicos]; Phegopteris aurita (Hooker) J. Smith; Polypodium auritum (Hooker) E. J. Lowe; Thelypteris aurita (Hooker) Ching.

Plants 40–100 cm tall. Rhizome long-creeping, with scales at apex; scales brown, narrowly lanceolate, 3–4 mm, ciliate along margins. Fronds remote; stipe red-castaneous or brownish at least toward base, shiny, 20–60 cm; lamina ovate-lanceolate, 20–70 Χ 15–30 cm, slightly tapering to base, pinnate-pinnatifid, pinnatifid and acuminate at apex; pinnae 10–18 pairs, opposite, spreading, sessile, proximal 1 or 2 pairs of pinnae slightly shortened, lanceolate, distal pinnae 7–15 Χ 2–4 cm, bases hastate, asymmetrical, pinnatifid nearly to costae, acuminate at apices, pinnae above middle subsymmetrical at base and adnate to rachis; segments (10–)15–20 pairs, spreading, basiscopic segments of pinnae longer than acroscopic ones, basal pair largest, particularly basiscopic one more oblique, lanceolate, 2.5–4 Χ 0.7–1 cm, margins pinnately lobed or crenate, acuminate at apices, acroscopic ones shorter, parallel to rachis, oblong, 1–2 cm, entire or shallowly undulate, obtuse at apices; veins visible abaxially, lateral veins forked or simple, 5–7 pairs per segment, basal pair arising from bases of costules. Laminae thickly herbaceous, drying brown-green, with short hairs along both sides of costae or only adaxially, laminae otherwise glabrous, rachises smooth abaxially, with dense short hairs adaxially. Sori oblong or sometimes ovate-round, borne above middle of ultimate veinlets, far from costules, 2–5 pairs per segment. Sporangia each with hairs below annulus. 2n = 62, 124.

Forest understories beside streams on high mountains; 1200–2000 m. Chongqing, C Fujian, C Guizhou, S and W Jiangxi, SE Xizang, W Yunnan [Bhutan, NE India, Indonesia, Japan, Malaysia, N Myanmar, Papua New Guinea, Philippines, N Vietnam].

The type is from NE India (Khasia, Meghalaya).

A widespread species, Pseudophegopteris aurita varies considerably in frond size, and hence in laminar dissection, over its range. Specimens outside of China tend to be considerably larger.

6. Pseudophegopteris subaurita (Tagawa) Ching, Acta Phytotax. Sin. 8: 315. 1963.

光轴紫柄蕨  guang zhou zi bing jue

Dryopteris subaurita Tagawa, Acta Phytotax. Geobot. 1: 157. 1932; Phegopteris subaurita (Tagawa) Tagawa; Thelypteris subaurita (Tagawa) Ching.

Plants 50–120 cm tall. Rhizome shortly creeping to ascending, apex and stipe base with dense brown linear-lanceolate hairy scales. Fronds clustered or slightly spaced; stipe red-castaneous at least at base, 10–35 cm, abaxially sparsely hairy, adaxially with dense hairs along groove; lamina oblong-lanceolate, 40–55(–100) Χ 15–30 cm, tapering at base, pinnate-pinnatifid, pinnatifid and long acuminate at apex; pinnae 15–20 pairs, opposite, obliquely spreading, sessile, basal pair of pinnae smallest, narrowly triangular, 2–6 Χ 3–6 cm, more distal pinnae gradually longer, second pair 8–15 Χ ca. 2 cm (excluding basalmost segments), triangular-lanceolate, bases abruptly widened up to 7 cm, asymmetrically hastate, pinnatifid nearly to costules, acuminate at apices; segments 15–25 pairs per pinna, opposite, obliquely spreading, basal pair clearly longer than distal pairs, particularly basiscopic one pinnatifid and more oblique, lanceolate, up to ca. 4 Χ ca. 1.2 cm, tapering to an acute apex, acroscopic one parallel to rachis, up to 4 cm, more distal segments up to 3 Χ ca. 0.8 cm, oblong, margins crenate to lobed, distal segments entire, obtuse or subacute at apices; veins visible abaxially, lateral veins simple or forked on proximal larger segments, basal pair arising from bases of costae. Fronds herbaceous, drying yellowish green, with fine acicular hairs on both surfaces, hairs along costae and veins denser abaxially. Sori nearly round to oblong, borne at middle or above middle (closer to margin) of ultimate veins. Sporangia each often with 1 or 2 short hairs below annulus. 2n = 62.

Forest understories beside streams, open areas in thickets; 200–1000 m. N and S Taiwan [Japan (Ryukyu Islands)].

The type is from Taiwan.

Pseudophegopteris subaurita is closely related to P. pyrrhorhachis and to P. aurita, but seemingly absent from mainland China.

7. Pseudophegopteris pyrrhorhachis (Kunze) Ching, Acta Phytotax. Sin. 8: 315. 1963[??p. 313].

紫柄蕨 zi bing jue

Plants 80–100 cm tall. Rhizome long-creeping, scaly at apex. Fronds approximate or sparse; stipe red-castaneous, shiny, 20–40 cm, base with short hairs and few lanceolate scales, distally glabrous; lamina oblong-lanceolate, 60–70 Χ 20–35 cm, base somewhat tapered, pinnate-pinnatifid, acuminate at apex; pinnae 15–20 pairs, opposite, sessile, proximal pinnae narrowly lanceolate, middle pinnae larger, 13–20 Χ 2.5–5 cm, bases slightly broadened, rounded-truncate, proximal 1–3 pairs sometimes slightly shortened, pinnatifid, shortly acuminate at apices; pinnules 15–25 pairs, opposite, spreading, lanceolate and slightly falcate, 1.5–2.5 Χ 0.5–0.8 cm, bases slightly broadened and adnate to costae, connected to each other by a narrow wing, lobed to 1/2 distance to costae, shortly acuminate at apices; segments triangular-oblong, acuminate and entire at apices. Veins hidden, pinnate on segments, 2–4 pairs per segment, basal pair arising from above base of costules. Fronds herbaceous, drying dark brown-green, abaxial surfaces with sparse short acicular hairs, hairs along costae, costules, and veins denser, adaxially with short setae along costules and veins; rachises and costae reddish brown, glabrous or sparsely shortly hairy. Sori nearly round or ovate, 1 or 2 per segment, borne above middle of veinlets and close to margins, arranged in an irregular row on each side of costules. Sporangia glabrous or each with 1 or 2 hairs below annulus. 2n = 62, 124, 186.

Forest understories beside streams; 800–2400 m. Widely distributed in provinces and regions south of Changjiang [just list the provinces], east to Taiwan, reaching southwest to Yunnan, northwest to S Gansu, north to Henan [Bhutan, N India, Myanmar, Nepal, Sri Lanka, Vietnam].

The type is from Nilgiris, India.

1a.     Laminae abaxially with dense short acicular hairs along costae, costules, and veins, sparsely hairy between veins                                                                                                                                             .    7a. var. pyrrhorhachis

1b.     Laminae abaxially glabrous or at most with extremely short capitate hairs along rachises, costae, and costules, glabrous between veins. ..............................................................................  7b. var. glabrata

7a. Pseudophegopteris pyrrhorhachis var. pyrrhorhachis.

紫柄蕨  zi bing jue

Polypodium pyrrhorhachis Kunze, Linnaea 24: 257. 1851; [??Dryopteris brunnea C. Christensen]; Dryopteris laterepens (E. W. Trotter ex Hope) C. Christensen; Macrothelypteris pyrrhorhachis (Kunze) Pichi Sermolli; Nephrodium brunneum Handel-Mazzetti; Phegopteris pyrrhorhachis (Kunze) Tagawa; Polypodium distans D. Don [??not Kaulfuss] var. adnatum C. B. Clarke; P. laterepens E. W. Trotter ex Hope; Thelypteris brunnea (Handel-Mazzetti[??C. Christensen]) Ching; T. pyrrorhachis (Kunze) Nayar [??C. M. Kuo].

Laminae abaxially with dense short acicular hairs along costae, costules, and veins, often hairy between veins.

Forest understories beside streams; 800–2400 m. Widely distributed in provinces [list them] and regions south of Changjiang, east to Taiwan, the southwest to Yunnan, the northwest to S Gansu, north to Henan [Bhutan, N India, Myanmar, Nepal, Sri Lanka, Vietnam].

7b. Pseudophegopteris pyrrhorhachis var. glabrata (C. B. Clarke) Holttum, Blumea 17: 24. 1969.

光叶紫柄蕨 guang ye zi bing jue

Polypodium distans var. glabrata C. B. Clarke, Trans. Linn. Soc. London, Bot. 1: 544. 1880; Phegopteris distans (D. Don) Mettenius var. glabrata Beddome.

Lamina abaxially glabrous or with extremely short capitate hairs along rachises, costae, and veins, lacking hairs between veins.

In forests along streams; up to 3000 m. N Guizhou, W Hubei, Sichuan, Yunnan [N. India, Myanmar, SW regions of Himalaya].

The type is from Sikkim.

8. Pseudophegopteris hirtirachis (C. Christensen) Holttum, Blumea 17: 22. 1969.

密毛紫柄蕨  mi mao zi bing jue

Dryopteris hirtirachis C. Christensen in H. Lιveillι, Fl. Kouy-Tcheou 49. 1915; Dryopteris christii H. Lιveillι (1915), not C. Christensen (1905), Pseudophegopteris pyrrhorhachis (Kunze) Ching var. hirtirachis Ching; Thelypteris brunnea (Handel-Mazzetti) Ching var. hirtirachis (C. Christensen) Ching.

Plants up to 1 m. Rhizome ascending, stipe base with brown lanceolate scales. Fronds somewhat clustered; stipe red-castaneous, shiny, 20–55 cm, base with scales and whitish hairs; lamina oblong, mostly 50–80 Χ 15–35 cm, slightly tapering at base, 2-pinnate-pinnatifid, acuminate and pinnatifid at apex; pinnae opposite to subopposite, sessile, lanceolate, middle pinnae largest, 9–20 cm or more, mostly 2–6 cm wide, slightly asymmetrical, acute at apices, proximal 1 or 2 pairs shortened, proximal pair 3–10 cm, pinnatifid or pinnatisect; pinnules 12–20 pairs per pinna, opposite, spreading, basiscopic ones on proximal pinnae slightly longer than acroscopic ones, 0.8–3.5 Χ 0.5–0.8 cm, bases broadened and adnate to costae, connected to each other by narrow wings, shortly acuminate to obtuse at apices, basal pair of pinnules same size or slightly longer, toothed to pinnatifid to 1/2 distance to costules; segments triangular-oblong, entire to dentate, obtuse to acute at apices. Veins visible on both sides, lateral veins simple or forked, mostly 4–15 or more pairs per pinnule, basal pair arising above bases of costules. Laminae herbaceous, drying deep green or brownish green, abaxial surfaces with dense short whitish hairs along costae, costules, veins, and between veins, adaxially with appressed thick short hairs along rachises, costae, and costules, rachises castaneous or red-castaneous and with sparse to dense, short, spreading hairs abaxially. Sori nearly round to oblong, near ends of ultimate veinlets. Sporangia each usually with 2 or more short hairs below annulus.

In forest understories along streams; 1500–2000 m. N Guangdong, N Guangxi, S Guizhou, C Sichuan, Taiwan, W Yunnan [NE India].

The type is from Pin-fa, Guizhou (Guiding).

Pseudophegopteris hirtirachis is similar to P. pyrrhorhachis (Kunze) Ching in laminar outline, but the rhizomes are shorter and ascending, the rachises bear short, spreading hairs abaxially, the costules and costae bear dense whitish short acicular hairs abaxially; sporangia with 2 or more short hairs.

9. Pseudophegopteris yunkweiensis (Ching) Ching, Acta Phytotax. Sin. 6: 315. 1963.

云贵紫柄蕨  yun gui zi bing jue

Thelypteris yunkweiensis Ching, Bull. Fan Mem. Inst. Biol. Bot. 6: 274. 1936; Phegopteris yunkweiensis (Ching) Tagawa.

Plants up to 1.5 m. tall. Rhizome stout (short and ascending?, not seen). Stipe red-castaneous, shiny, 40–55 cm, base with dense brown, ovate-lanceolate, hairy scales, distally glabrous; lamina ovate, ca. 100 Χ 40–60 cm, not tapering to base, 2-pinnate-pinnatifid, apex acuminate and pinnatifid; pinnae 10 or more pairs, opposite, or distal ones subopposite and more oblique, sessile or proximal ones shortly stalked, proximal pair largest, 30–45 Χ 13–20 cm, oblong-lanceolate, bases symmetrical or basal several pinnules shorted acroscopically, pinnate-pinnatifid, acuminate at apices, basiscopic pinnules not elongate relative to more distal pinnules; pinnules 15–25 pairs per pinnae, 2.5–3 cm apart, alternate, spreading, mostly free from each other, sessile or narrowly adnate to costae, distalmost ones connected by a narrow wing, lanceolate, 6–13 Χ 1.4–2.4 cm, bases truncate, pinnatifid 3/4–4/5 distance to costules, acute at apices; segments 15–20 pairs, falcate-lanceolate, margins crenate or entire distally, acute or obtuse at apices; veins visible on both sides, especially adaxially, lateral veins forked, 4–7 pairs per segment. Laminae herbaceous, drying dark green, rachises and costae reddish, costules stramineous, abaxially with sparse acicular hairs or glabrescent along rachises, costae, and costules, with denser acicular hairs adaxially along costae and costules, glabrous on both surfaces on and between veins. Sori nearly round or slightly oblong, borne at middle of lateral veins between costae and margins, 3–6 pairs per segment. Sporangia glabrous.

Forest understories beside streams. SW Guizhou (Zhunyi), SE Yunnan (Mengzi) [N Vietnam].

The lectotype (chosen by Holttum, 1969) is from Vietnam (Tonkin, Chapa).

10. Pseudophegopteris zayuensis Ching & S. K. Wu, Fl. Xizang. 1: 164. 1983, [‘zayu$Uensis’].

察隅紫柄蕨  cha yu zi bing jue

Plants ca. 85 cm tall. Rhizome not seen. Stipe reddish brown, shiny, ca. 30 cm, scaly at base; scales reddish brown, linear-lanceolate, glabrous, or occasionally with a few long hairs dorsally, including rachises with sparse whitish acicular hairs; lamina up to ca. 55 Χ 16 cm, slightly tapering at base, pinnate-pinnatifid, shortly acuminate and pinnatifid at apex; pinnae ca. 25 pairs, opposite or subopposite, proximal 1 or 2 pairs sessile, spreading, proximal pair shortest, ca. 5 cm, middle pinnae ca. 8 Χ 1.5–2 cm, linear-lanceolate, bases truncate and symmetrical, pinnatifid up to 3/4 of distance to costae, acute at apices; segments ca. 20 pairs per pinna, rectangular, 7–8 Χ 4–5 mm, entire. Veins visible abaxially, 4 or 5 pairs per segment, basal pair arising from above base of costules. Laminae thinly papery, drying green, subglabrous on both surfaces between veins, costae at bases red-castaneous abaxially, distally stramineous, shiny, with sparse acicular hairs. Sori round, borne near tips of acroscopic vein of forked lateral vein pair and close to margin. Sporangia glabrous.

* Understories of broad-leaved forests; ca. 2100 m. S Xizang.

This species is known only from the type.

Pseudophegopteris zayuensis is closely related to P. tibetana, differing primarily by the basiscopic basal pinnule on proximal pinnae not being strongly elongate and the glabrous sporangia. More collections are needed to evaluate the importance of these differences.

11. Pseudophegopteris rectangularis (Zollinger) Holttum, Blumea 17: 19. 1969.

对生紫柄蕨 dui sheng zi bing jue

Polypodium rectangulare Zollinger, Syst. Verz. 37, 46[??48]. 1854; Dryopteris moussetii Rosenstock; D. oppositipinna (Alderwerelt) C. Christensen; Phegopteris oppositipinna Alderwerelt; Polypodium distans D. Don [??Kaulfuss] var. minor C. B. Clarke; Pseudophegopteris oppositipinna (Alderwerelt) Ching; Thelypteris oppositipinna (Alderwerelt) Ching; T. rectangularis (Zollinger) B. K. Nayar & S. Kaur.

Plants 40–70 cm tall. Rhizome short, suberect. Fronds clustered; stipe red-castaneous, shiny throughout, 10–30 cm, with short spreading acicular hairs, base with narrowly lanceolate, brownish, sparsely hairy, thin scales; lamina narrowly oblong-lanceolate, 30–50 Χ 7–12 cm, slightly tapering and with pinnae more spaced proximally, pinnate-pinnatifid; pinnae ca. 20 pairs, opposite to subopposite, spreading, sessile and distal ones becoming narrowly adnate to rachis, proximal 1–3 pairs slightly shortened, up to 3 cm, basal segments of proximal pinnae not significantly elongate or lobed, distal pinnae lanceolate, 4–7 Χ 0.8–1.8 cm, pinnately lobed to 3/4 of distance to costae, acuminate at apices; segments 12–20 pairs per pinna, obliquely spreading, oblong-triangular, entire along margins, obtuse at apices. Veins visible, lateral veins simple or occasionally forked, 3–5 pairs per segment, basal pair arising from above base of costules, acroscopic one of a pair running toward sinus but not reaching margin. Laminae herbaceous, drying greenish, adaxial surfaces nearly smooth except for hairs along costae, abaxial surfaces with sparse to many acicular hairs to 0.5 mm, hairs along costae denser, also sometimes with a few capitate short hairs between veins, rachises red-castaneous and with denser acicular hairs. Sori nearly round, borne at middle or near tips of lateral veins, 2–4 pairs per segment. Sporangia each usually with 1 or 2 acicular hairs below annulus. 2n = 124.

Forest understories beside streams; 1000–1500 m. N Guangxi, SE Xizang, SE and W Yunnan [NE India, Indonesia, Malaysia].

The type is from Indonesia (Java).

12. Pseudophegopteris microstegia (Hooker) Ching, Fl. Xizang. 1: 162. 1983.

禾杆紫柄蕨  he gan zi bing jue

Nephrodium microstegium Hooker, Sp. Fil. 1: 119. 1862; Pseudophegopteris pallida (Ching) Ching; Thelypteris brunnea (C. Christensen) Ching var. pallida Ching.

Plants 90–120 cm tall. Rhizome long-creeping, with sparse, appressed, brownish, broadly lanceolate scales. Fronds remote; stipe stramineous, occasionally brown-stramineous, 30–40 cm, base with sparse brownish lanceolate scales, distally glabrous; lamina 60–80 Χ 20–30 cm, tapering proximally, pinnate-pinnatifid, pinnatifid and acuminate at apex; pinnae 20–25 pairs, subopposite, obliquely spreading or spreading, sessile, proximal pinnae linear-lanceolate, 10–15 Χ 2–3 cm, bases truncate, acute at apices, proximal 2 or 3 pairs sometimes slightly shortened, 7–10 cm, pinnatifid nearly to costae; segments ca. 25 pairs per pinna, opposite, spreading, basal pair of segments same shape and size as more distal ones, oblong, 1–1.5 Χ ca. 0.5 cm, coarsely dentate or entire along margins, sparsely ciliate, obtuse or truncate at apices. Veins visible abaxially, lateral veins forked, 6 or 7 pairs per segment, basal pair arising from above base of costules. Laminae thinly herbaceous, drying greenish, both surfaces with short hairs along costae and veins, hairs denser adaxially, rachises stramineous, adaxially densely hirsute along groove. Sori round to slightly oblong, borne at middle of acroscopic vein of lateral vein pair. Sporangia glabrous.

Understories of evergreen broad-leaved forests; 2300–2400 m. Chongqing (Nanchuan), C Sichuan, S and SE Xizang, NW and SE Yunnan [NE India].

The type is from NE India, Khasia.

Pseudophegopteris microstegia is very similar to P. pyrrhorhachis (and treated as a synonym of that by Holttum, 1969), differing mainly in the stramineous stipes and rachises and laminae abaxially glabrous except on rachises and costae. The purported differences need further study.