PTERIDIACEAE [Draft]

蕨科  jue ke

Liao Wenbo (廖文波), Ding Mingyan (丁明艳), Wu Shiewhung (吴兆洪); Jefferson Prado

Ferns, terrestrial, large to medium-sized (0.6–2 m tall). Rhizome long-creeping, vascular system a complex dissected double solenostele, densely clothed with rusty yellow or chestnut articulate hairs, without scales. Fronds monomorphic; stipe well-developed, with one to several vascular bundles, branch buds present near stipe base; lamina often ovate, ovate-oblong, or ovate-triangular, 20–120 ื 10–100 cm, leathery or papery, abaxially ฑ puberulent, rarely glabrescent, adaxially glabrous, 3- or 4-pinnate, broadly lobed (Pteridium) or narrowly lobed (Paesia). Venation free; veins forked. Sori submarginal, linear, on a vascular commissure joining apices of veins; indusia formed by two layers; outer layer consisting of thin reflexed margin of lamina, persistent, conspicuous; inner layer thinner, not conspicuous, sometimes nearly degenerate, interrupted at sinuses and absent at segment apex. Spores trilete (Pteridium) or monolete (Paesia), smooth or slightly gemmulate.

Two genera: widely distributed in the tropics; both genera occur in China [??number of species in the world]; seven species in China (four endemic).

[??move to Dennstaedtiaceae]

1a.     Stipe with several vascular bundles; spores trilete ................................................  1. Pteridium

1b.     Stipe with only one vascular bundle; spores monolete ...............................................  2. Paesia

1. PTERIDIUM Gleditsch ex Scopoli, Fl. Carniol. 169. 1760, nom. cons.

蕨属  jue shu

Plants terrestrial, robust. Rhizome brownish black, thick, ca. 5 mm in diam., hypogeal, densely clothed with rusty yellow to castaneous hairs. Fronds 50–130 cm; lamina large, often ovate or ovate-triangular, leathery to papery, abaxially ฑ pilose, adaxially glabrous or sparsely hairy, 3- or 4-pinnate-pinnatifid, costae of ultimate pinnae closed with fine cinereous-brown hairs, rarely glabrescent; rachises grooved adaxially, glabrous or pubescent; pinnae subopposite or alternate, petiolulate, basal pinnae pair larger than other pinnae, triangular; venation of ultimate pinnae pinnate, veins 2-forked, adaxially conspicuously prominent. Sori submarginal, linear, paraphyses not present; indusia formed by two layers; outer layer consisting of thin reflexed margin of lamina, persistent; inner layer thinner, vestigial, not conspicuous, sometimes nearly degenerate; sporangia long pedicellate; spores trilete, with thin nipple-shaped protuberances. x = 13.

About 13 species: widely distributed worldwide, but mainly in the tropics; six species in China (three endemic).

1a.     Rachises and costae glabrous in adaxial grooves.

2a.    Ultimate segments entire, broadly lanceolate to oblong, near each other or spaces not wider than their own width; widely distributed ..................................................  1. P. aquilinum var. latiusculum

2b.    Ultimate segments entire, linear, widely spaced, often by 1–2 ื their own width.

3a.     Fronds leathery when dried; ultimate segments linear, ca. 2.5 mm wide or wider, abaxially midvein hairy .........................................................................................................  2. P. esculentum

3b.     Fronds papery when dried; ultimate segments broadly linear, ca. 4 mm wide, abaxially nearly glabrous except on midvein ..................................................................................  3. P. falcatum

1b.     Rachises and costae densely hairy in adaxial grooves (at least when young).

4a.    Fronds subleathery; secondary pinnae lanceolate, up to 8 ื 1.5 cm ..............  4. P. revolutum

4b.    Fronds papery or leathery; secondary pinnae narrowly linear or linear-lanceolate, 13–20 ื ca. 3 cm.

5a.     Fronds papery; secondary pinnae narrowly linear, ca. 13 mm, adaxially pubescent  5. P. lineare

5b.     Fronds herbaceous; secondary pinnae linear-lanceolate, ca. 13 cm, adaxially pilose  6. P. yunnanense

1. Pteridium aquilinum (Linnaeus) Kuhn var. latiusculum (Desvaux) Underwood ex A. Heller, Cat. N. Amer. Pl. (ed. 3), 17. 1909.

 jue

Pteris latiuscula Desvaux, M้m. Soc. Linn. Paris 6(3): 303. 1827; Pteridium aqualinum f. glabrum Tardieu & C. Christensen; P. aqualinum var. japonicum Nakai; P. aquilinum subsp. latiusculum (Desvaux) Hult้n; P. japonicum Tardieu & C. Christensen; P. latiusculum (Desvaux) Hieronymus ex Fries; Pteris lanuginosa Sprengel (1821), not Bory ex Willdenow (1810).

Plants up to 2 m tall. Rhizome long-creeping, densely castaneous hairy. Fronds erect, 0.5–1.5 m (sometimes larger in shaded forms); stipe brown, up to 40 cm, woody, basally with short brown hairs, apically glabrous; lamina deltoid to oblong-ovate in outline when pressed, leathery, 3- or 4-pinnate-pinnatifid; basal pinnae as long as to 1/2 length of lamina; rachises pale brown, thinly pubescent, becoming glabrous; lobes present between two adjacent segments; pinnae ascending or horizontal, ovate-deltoid to oblong, up to 40 ื 15 cm, acute; pinnules or segments linear to oblong, glabrous except for pinnule margins and midvein, deeply pinnatifid in larger pinnae, shortly bluntly or long caudate. Sori elongate, submarginal on ultimate segments; outer indusium ca. 0.5 mm wide, membranous, ciliate, inner indusium vestigial and fimbriate. 2n = 104.

Sunny slopes and edges of forests; 200–800 m. Throughout China, but mostly in S China including Taiwan [Japan (including Ryukyus); Europe, North America].

The type of Pteridium aquilinum var. latiusculum is from Canada (Newfoundland and St. Pierre).

The typical variety is distributed in Africa and Europe.

2. Pteridium esculentum (G. Forster) Cokayne in Engler, Vegt. Der Erde 14: 71, 308. 1921 [??Rep. Bot. Surv. Tongariro Nat. Park 34. 1908].

食蕨  shi jue

Pteris esculenta G. Forster, Pl. Escul. 74. 1786.

Plants ca. 1 m tall. Rhizome long-creeping. Fronds firmly leathery when dried; stipe yellowish brown, shiny, 40–50 cm, abaxially sulcate; lamina oblong-triangular, 50–60 cm, base rounded-cuneate, 60–80 cm wide, 3-pinnate-pinnatifid, apically acuminate; pinnae 15–18 pairs, alternate, stipitate (ca. 2 cm), slightly obliquely spreading, oblong-lanceolate, up to 35 ื 11–15 cm, base cordate or subtruncate, apex long-acuminate, caudate; pinnules ca. 30 pairs per pinna, alternate, sessile, broadly lanceolate, 5.5–7 ื 1.5–2.5 cm, base truncate, spreading, apex caudate (1.5–2 cm) to acuminate; segments 12–14 pairs per pinna, linear, lateral ones ca. 1 cm ื 2–2.5 mm, gradually wider upward, alternate, spreading, connate with costules, margins reflexed, serrulate, apex obtuse; middle pinnules similar in shape to lower ones but narrower, apex caudate; apical pinnules linear, up to 3 cm ื ca. 3 mm, not pinnate (entire); apical pinnae lanceolate, pinnate; segments linear, apically obtuse; veins not conspicuous, hairy abaxially, veinlets somewhat grooved; rachises, costae, and costules shiny, glabrous, adaxially grooved; rachises yellowish brown, costae and costules same color as lamina (green).

Sunny gaps on slopes; ca. 1000 m. SW Guangxi (Longjin, Napo), Hainan (Diaoluoshan) [widely distributed in Indochina[??countries], Indonesia, Malaysia; Australia, New Zealand, South-Pacific Islands].

The type of Pteridium esculentum is from the Society Islands.

3. Pteridium falcatum Ching, Acta Bot. Austro Sin. 2: 2: 1986.

镰羽蕨  lian yu jue

Plants ca. 2 m tall. Rhizome creeping. Fronds papery when dried; stipe brownish black, acroscopically reddish brown to brown-stramineous, slightly lustrous, up to 1 m, ca. 5 mm in diam. at base, adaxially grooved; lamina pale green, narrowly triangular-ovate, ca. 1 m, base ca. 50 cm wide, abaxially with light cinereous-brown hairs along costae, adaxially glabrous, base rounded-cuneate, 3-pinnate-pinnatifid, apex acuminate; pinnae ca. 6 pairs, opposite or subopposite; basal pinnae ovate-triangular, base rounded-cuneate, stalked (ca. 6 cm), 2-pinnate-pinnatifid (1-pinnate apically), apex acuminate; pinnules ca. 10 pairs per pinna, alternate, slightly obliquely spreading; basal pair ca. 22 ื 7–8 cm, pinnate, narrowly triangular, base truncate, stalked (ca. 1 cm), apex long caudate (2.5–3 cm); segments ca. 18 pairs per pinnule, alternate, spreading, falcate-lanceolate, up to 3 ื 3–4 cm, apex acute; lower ones nearly separate from costules or slightly connected to costules, basal ones generally triangular auriculate, entire, costule without ala [??wings], only ultimate segments, slightly expanded on both sides; veins close to each other, anadromous, conspicuous on both surfaces; rachises and costae glabrous, costules glabrescent, adaxially deeply grooved, glabrous, straw-colored.

* Sunny slopes. E Guangxi (Hexian).

4. Pteridium revolutum (Blume) Nakai, Bot. Mag. (Tokyo) 39: 109. 1925.

毛轴蕨  mao zhou jue

Plants up to 1 m tall. Rhizome long-creeping. Fronds subleathery when dried, margins often revolute; stipe straw-colored or brown, 35–50 cm, 5–8 mm in diam. at base, adaxially grooved, densely clothed with pallid hairs when young, glabrous when old; lamina broadly triangular or ovate-triangular, 30–80 ื 30–50 cm, 3-pinnate-pinnatifid, acuminate; pinnae 4–6 pairs, opposite, decumbent, stalked (2–3 cm), oblong, base subtruncate, apex acuminate; basal pinnae slightly triangular, 20–30 ื 10–15 cm, stalked (2–3 cm), 2-pinnate-pinnatifid; pinnules 12–18 pairs per pinna, opposite or alternate, spreading, sessile, base not adnate to costule, lanceolate, 6–8 ื 1–1.5 cm, base truncate, deeply pinnate, apex shortly caudate-acuminate; segments ca. 20 pairs per pinnule, opposite or alternate, slightly obliquely spreading, lanceolate-falcate, ca. 8 mm, base ca. 3 mm wide, connate, often entire, apex obtuse or acute; apical pinnae 2-pinnate-pinnatifid, lanceolate; segments approximate, with pallid or light brown hairs abaxially; veins prominent abaxially, grooved adaxially; rachises, costae, and costules approximate, with pallid or light brown hairs or tuberculate, glabrescent.

Sunny slopes, open shaded forests; 600–3000 m. Gansu (Wenxian), Guangdong, Guangxi, Guizhou, Hubei (Badong, Gucheng), Hunan (Qianyang), Jiangxi (Jinggangshan), Shaanxi (S slopes of Qinling), Sichuan, Taiwan, Xizang (Jiazhongka, Yadong), Yunnan, [??add]Zhejiang [widely distributed in tropical and subtropical regions of Asia].

1a.. Costae approximate, with pallid or light brown hairs, glabrescent, (not verrucate); segment (pinnule) apices acute or shortly caudate-acuminate ..............................................................  4a. var. revolutum

1b.. Costae tuberculate (verrucate) abaxially; segment (pinnule) apices obtuse or rounded ......  4b. var. muricatulum

4a. Pteridium revolutum var. revolutum

毛轴蕨 (原变种)  mao zhou jue (yuan bian zhong)

Pteris revoluta Blume, Enum. Pl. Javae 214. 1828; Pteridium aquilinum (Linnaeus) Kuhn var. osmundoides H. Christ ex H. L้veill้, nom. nud.; P. aquilinum subsp. wightianum (J. Agardh) W. C. Shieh; P. aquilinum var. wightianum (J. Agardh) R. M. Tryon; P. capense (Thunberg) Krasser var. densa Nakai; Pteris densa Wallich, nom. nud.; P. recurvata Wallich var. wightiana J. Agardh; P. villosa F้e; P. wightiana Wallich, nom. nud.

Costae approximate, with pallid or light brown hairs, gradually more sparsely hairy; segment apices acute or shortly caudate-acuminate.

Sunny slopes, open shaded forests; 600–3000 m. Gansu (Wenxian), Guangdong, Guangxi, Guizhou, [??add]Henan, Hubei (Badong, Gucheng), Hunan (Qianyang), Jiangxi (Jinggangshan), Shaanxi (S slopes of Qinling), Sichuan, Taiwan, Xizang (Jiazhongka, Yadong), Yunnan [widely distributed in tropical and subtropical regions of Asia: India, Indochina [??countries], Indonesia, Malaysia, Philippines].

The type of Pteridium revolutum is from Indonesia (Dindygul).

4b. Pteridium revolutum var. muricatulum Ching & S. H. Wu, Acta Bot. Austro Sin. 2: 2. 1986.

糙轴蕨  cao zhou jue

Costae abaxially scabrous verrucose; segment apices obtuse or rounded.

* Slopes; 800–2700 m. Sichuan (Emei Shan, Lixian), Yunnan (Jinping, Mengzi).

The type of Pteridium revolutum var. muricatulum is from Yunnan (Jinping).

5. Pteridium lineare Ching, Acta Bot. Austro-Sin. 2: 3. 1986.

长羽蕨  chang yu jue

Plants up to 2 m tall. Rhizome creeping. Fronds papery when dried; stipe apically straw-colored, robust, ca. 0.8 m, up to 1 cm in diam., glabrous; lamina abaxially cinereous-green, adaxially green, triangular, ca. 1.2 m, abaxially with sparse pallid hairs or glabrescent, adaxially glabrous, 3-pinnate-pinnatifid; pinnae ca. 5 pairs; basal pinnae opposite, obliquely spreading, oblong-triangular, up to 90 ื 40–50 cm, base broadly cuneate, stalked (up to 9 cm), 2-pinnate-pinnatifid, apex acuminate; pinnules ca. 50 pairs, alternate, spreading, at nearly 90ฐ angle to costa, narrowly linear; basal basiscopic pinnules ca. 20 ื 3 cm, base truncate, sessile, apex acuminate; segments ca. 40 pairs per pinnule, spreading, linear-lanceolate, ฑ falcate, ca. 1.5–2 cm, base nearly symmetric, adnate to costules, not decurrent, separated by narrow sinuses, generally basal margins ฑ slightly lobed or undulate, entire upward, apex acute; basal pair close to costules and parallel to costules; basal acroscopic pinnules shorter, 1/3–1/2 as long as basiscopic pairs (0.5–1 cm); median pinnules conforming to basal pair, apical pairs shorter; veins forked twice from base, veinlets close to each other, abaxially conspicuous, adaxially somewhat grooved; costules, costae, and rachises adaxially grooved, often with sparse pallid hairs in grooves, abaxially sparsely hairy or glabrate.

* Acidic soils of wastelands; ca. 1000 m. W Yunnan (Mangshi).

6. Pteridium yunnanense Ching & S. H. Wu, Acta Bot. Austro Sin. 2: 4. 1986.

云南蕨  yun nan jue

Plants ca. 1 m tall. Rhizome creeping. Fronds herbaceous when dried; stipe apically light straw-colored, robust, ca. 30 cm, ca. 5 mm in diam., glabrous; lamina green, ovate-triangular, ca. 70 ื 70 cm, abaxially densely hairy (especially on costules), adaxially sparsely hairy, 4-pinnate-pinnatifid; basal pinnae largest, opposite, decumbent, oblong-triangular, up to 60 ื 22 cm, base broadly cuneate, long-stalked (ca. 6 cm), 3-pinnate-pinnatifid, apex acuminate; pinnules ca. 20 pairs, opposite or subopposite, spreading or decumbent, linear-lanceolate, ฑ falcate; basal ones ca. 13 ื 3 cm, base truncate, symmetrical, subsessile, 2-pinnate-pinnatifid, apex long acuminate; ultimate pinnules 20–23 pairs per pinnae, spreading, lanceolate, ca. 1.5 cm ื ca. 5 mm, base acuminate, connate with costules at base, symmetrical, separated by a wide sinus, pinnatifid; segments 6 or 7 pairs per pinnule, spreading, oblong, entire, basal pair larger; middle pinnules same size as basal ones, reduced in size upward; veins pinnate, veinlets 2-forked, conspicuous abaxially, grooved adaxially; costules, costae, and rachises adaxially deeply grooved, and costules with sparse pallid hairs, abaxially with sparse hairs or glabrate.

* About 1500 m. W Yunnan (Yingjiang).

2. PAESIA J. Saint-Hilaire, Voyage Dist. Diamans 1: 381. 1833.

曲轴蕨属  qu zhou jue shu

Plants terrestrial. Rhizome long-creeping, solenostelic, with chestnut hairs, without scales. Fronds distant, long-stalked; lamina pinnate, subleathery; rachises often zigzag, flexuous, ciliate; venation free. Sori submarginal, on a vascular commissure joining apices of veins; indusia double: formed by two layers, outer layer false, formed by reflexed edge of lamina, inner layer true, not conspicuous; annulus consisting of (12–)17–20 incrassate cells; spores monolete, laevigate. x = 13.

Widely distributed: tropical America, New Zealand to Tahiti, Philippines (Luzon), and Indonesia (Sumatra), ca. 14 species; only one species in China (Taiwan) (the one endemic).

1. Paesia taiwanensis Shieh, J. Jap. Bot. 45: 161. 1970

台湾曲轴蕨  tai wan qu zhou jue

Plants ca. 85 cm tall. Rhizome long-creeping, 1.5–4.5 mm in diam., hairy, hairs castaneous, lucid, bristlelike. Fronds distant, firmly herbaceous when dried; stipe brown, shiny, slender, 10–45 cm, hispid at base, muricate apically, slightly grooved adaxially; lamina green, ovate-oblong, 22–60 ื 11–25 cm, abaxially sparsely pubescent, adaxially glabrous, 3-pinnate-pinnatifid; pinnae numerous (ca. 15 pairs), alternate, decumbent, shortly stalked or subsessile, oblong-lanceolate, base subtruncate, apex acuminate; basal pair largest, 6–18 ื 2.5–9 cm; rachises zigzag, straw-colored, scabrous or glabrous; pinnules numerous, shortly stalked or sessile, oblong-lanceolate, 1–5 ื 0.5–0.8 cm, base subtruncate; ultimate pinnules 5–14 pairs, oblong or broadly subulate, to 8 ื 3 mm, 2–5-aristate on each side; veins simple or 2-forked. Sori absent at segment apex; indusia double, linear, membranous, entire, persistent.

* Taiwan (Taidong).