MARATTIACEAE [Draft]

合囊蕨科  he nang jue ke

He Zhaorong (和兆荣); Maarten J. M. Christenhusz

Ferns, terrestrial (rarely epiphytic, sometimes in streams), evergreen. Rhizome erect, ascending, or creeping, amylaceous[??starchy], polycyclostelic to dictyostelic. Fronds monomorphic and long-lived or dimorphic (Danaea) and fertile ones short-lived, simple or compound, 1–4-pinnate or (pedate-)palmate (Christensenia), small (ca. 20 cm) to very large (up to at least 6 m). Stipe fleshy, with a pair of stipulelike appendages at base often with conspicuous lenticels. Pulvini at base of fronds, at nodes, at base of pinnae and in some species at frondless nodes along stipe. Veins free, simple or bifurcate, or reticulate (Christensenia), false veins present between true veins or absent. Tissues mucilaginous. Indument of multicellular uniseriate hairs and basifixed or peltate scales. Sori lacking true indusia, usually with paraphyses, superficial, sunken or stalked, elongate along veins, bilateral or radial. Sporangia partially or completely fused into synangia, dehiscing by a slit or pore. Spores trilete or monolete, with bulliform, muriform, or stelliform ornamentation. Prothallia very large, thalloid, photosynthetic.

Six genera and ??? species: pantropical, Danaea, Eupodium, and Marattia in the Neotropics, Angiopteris, Christensenia, and Ptisana in the Paleotropics; three genera and ca. 24 species in China (13 endemic).

1a.       Fronds simple, pedately lobed to palmately compound; venation reticulate; synangium radial .  1. Christensenia

1b.       Fronds pinnate (sometimes simple when plants juvenile); venation free, simple or bifurcate; synangium ovate.

2a.       Sporangia fully fused into synangia, synangia bivalved; (Taiwan) ...........................  2. Ptisana

2b.       Sporangia fused basally but free apically, synangia composed of globose sporangia  3. Angiopteris

1. CHRISTENSENIA Maxon, Proc. Biol. Soc. Wash. 18: 239. 1905.

天星蕨属  tian xing jue shu

Kaulfussia Blume, Enum. Pl. Javae 2: 260. 1828, not Dennstedt (1818), nor Nees (1820); Macrostoma Griffith.

Plants terrestrial. Rhizome creeping. Stipe with pulvini at base of segments. Lamina simple (and then usually pedately lobed) to palmately compound, base rounded to pedate, margin simple to broadly crenate, apex acute to acuminate. Venation reticulate. Stomata large, raised, round, permanently open, densely distributed on abaxial frond surfaces, visible to naked eye. Scales flattened, peltate with lobed margins on short stalks, hairs glandular, mixed with scattered uniseriate simple hairs. Sori ± in rows on either side of main veins, appearing scattered. Sporangia fully fused into circular, sessile synangia, sporangium dehiscing by a vertical slit on inside of synangium. Spores monolete, ellipsoid, exospore long spinulose, spines simple. 2n = 80, 160.[??x]

One species: patchily distributed from NE India, Indochina, and Indonesia to the Solomon Islands.

1. Christensenia aesculifolia (Blume) Maxon, Proc. Biol. Soc. Wash. 18: 240. 1905.

Chinese  pinyin

Aspidium aesculifolium Blume, Enum. Pl. Javae 2: 143. 1828; Christensenia assamica (Griffith) Ching; C. lobbiana (de Vriese) Rolleri; Kaulfussia aesculifolia (Blume) Blume; K. assamica Griffith; K. korthalsii de Vriese; K. lobbiana de Vriese.

Plants up to 80 cm tall. Rhizome creeping (to suberect), short, fleshy, scaly; scales brown with red spots, large, rounded. Stipe up to 36(–50) cm. Lamina simple or pedately lobed to palmate, middle pinnae largest, 9–25 × 3–15 cm, bases cuneate, margins entire to undulate, apices acute to acuminate, distal pinnae (if present) smaller, inequilateral and subrounded at base, margins entire, lobed, or broadly crenate, usually undulate, apices acute to acuminate. Synangia radially arranged, circular, made up of 8–12 sporangia.

On limestone. Yunnan [India, Indonesia, Malaysia, Myanmar, Philippines, Vietnam; Pacific islands (Bismarck Archipelago, Solomon Islands)].

2. PTISANA Murdock, Taxon 57: 744. 2008.

合囊蕨属  he nang jue shu

Plants terrestrial. Rhizome erect. Stipe with pulvini at base of pinnae and at nodes. Lamina 1–4-pinnate, evenly divided throughout, primary division generally (sub-)opposite, terminal segments usually with a prominent suture at point of attachment; pinnae margins usually dentate, occasionally crenulate or entire. Venation free, simple or bifurcating. Scales peltate, often asymmetrical and appearing basifixed, larger scales stalked, scale cells elongate, hairs uniseriate, simple or branching. Sori on veins, marginal to medial. Sporangia fully fused into sessile synangia, outer walls composed of small, rounded cells, mature synangia not sulcate along septa in. Synangia comprised of two opposing rows of sporangia and subtended by rings of paraphyses or uniseriate hairs. Synangium opening as a unit, deeply cut, bivalved. Sporangia dehiscing by a vertical slit on inner surface of each sporangium valve. Spores monolete (rarely trilete or alete), exospores granular to rugose. 2n = 78, 156.[??x]

About 20–25 species: widely distributed in the Paleotropics with the highest diversity in New Guinea, formerly often referred to the genus Marattia (which is a genus restricted to the Neotropics and Hawaii); one species in China.

1. Ptisana pellucida (C. Presl) Murdock, Taxon 57: 747. 2008.

合囊蕨  he nang jue

Marattia pellucida C. Presl, Suppl. Tent. Pterid. 10. 1845; M. haenkeana C. Presl; M. vestita H. Christ.

Rhizome erect, globose, up to 40 cm tall. Fronds up to 2 × 1.5 m. Stipe shorter than lamina, ca. 2 cm in diam., covered with cortical spines and irregularly toothed, narrowly lanceolate brown scales. Lamina oblong in outline, tripinnate, herbaceous to papery; pinnae up to 60 cm; ultimate pinnules narrowly elliptic, 6–10 × 1–1.5 cm, bases cuneate, margins serrate, apices acuminate; veins free, 1.5–2 mm apart, simple, not forked; costae with small, castaneous, thin scales with bullate base. Synangia medial or supramedial, 1–2.2 mm, with up to 8- or 9-cellular uniseriate paraphyses, branched paraphyses rare.

Forests on slopes. Taiwan (Lan Yu, Taitung) [Philippines].

3. ANGIOPTERIS Hoffmann, Commentat. Soc. Regiae Sci. Gott. 12: 29. 1796, nom. cons., not Adanson (1763).

莲座蕨属  lian zuo jue shu [or 原始莲座蕨属 yuan shi lian zuo jue shu]

Archangiopteris H. Christ & Giesenhagen; Clementea Cavanilles; Macroglossum Copeland; Protangiopteris Hayata; Protomarattia Hayata; Psilodochea C. Presl.

Plants terrestrial (sometimes rooted in stream beds). Rhizome erect, ascending, or creeping. Lamina 1–4-pinnate, usually evenly divided throughout, occasionally with irregular division of some segments (probably a result of stress when lamina developed). Pulvini present at bases of segments, at frondless nodes on stipe in juvenile fronds, generally absent at maturity, but present at maturity in some species with creeping rhizomes; primary pinnae alternate or subopposite; veins free, simple or bifurcate, false veins absent or present between veins of varying length and extending from margins toward costae; uniseriate simple hairs present, not glandular, scales peltate but generally appearing basifixed due to asymmetry. Sori borne on veins, marginal, submarginal, or medial; sporangia ± free, fused at base into receptacles, sessile, bilateral, each with two opposing rows of sporangia; opening via a vertical slit on inner surface of each valve, apertures labiate, with distinct patch of specialized thickly walled cells at apex of each sporangium. Spores trilete (rarely monolete or alete), exospores granular to spinulose, spines simple or branched. 2n = 80, 160.[??x]

About 30–40 species: widely distributed in the Paleotropics, from Madagascar to the S Pacific islands; introduced and naturalized in Hawaii, Jamaica, and Central America; ca. 22 species in China (13 endemic).

Differences are difficult to observe on herbarium specimens; species of Angiopteris differ chiefly in habit, size, and general appearance.

The historical variation in species delimitation in Angiopteris has made nomenclature in this genus highly unstable. Moreover, the characters distinguishing species of Angiopteris are usually difficult to observe on herbarium specimens, which are fragmentary and usually lack information on stipule, stipe, frond size, and general habit. Therefore, many of the published names are difficult to interpret, because type specimens are often small frond fragments. The majority of species listed here, therefore, may not be readily identifiable in herbaria. Most species are relatively scarce (or at least rarely collected), and it should, therefore, be noted that the majority of Chinese specimens probably belong the following taxa: Angiopteris evecta, A. fokiensis, A. somai, and A. wallichiana. The diversity of this genus in Hainan and Yunnan is far from understood; it is possible that hybridization or allopolyploidy plays a part in the complexity of Angiopteris. Molecular work and cultivation experiments are needed to solve the complex taxonomy of this genus, but the morphological diversity is far greater than the genetic diversity in Angiopteris. Nucleotide sequence data suggest that there may be only a small number of species with highly plastic morphology. Even though perhaps not satisfactory, the list of species below should be seen as an approximate consensus.

1a.       Rhizome dorsiventral, long creeping; lamina of mature plants once pinnate, rarely bipinnate; stipes with 1–4(–7) frondless pulvini in mature fronds; synangia medial, elongate (Archangiopteris).

2a.       Lamina once to twice pinnate, usually with 10–12 pairs of lateral pinnae ........  1. A. bipinnata

2b.       Lamina once pinnate, pinnae usually with 2–7 pairs of lateral pinnae.

3a.       Frondless pulvini 4 or 5(–7) per stipe; sori 3–4 mm, paraphyses long and dense  2. A. chingii

3b.       Frondless pulvini only 1 per stipe; sori shorter than 3 mm; paraphyses short and sparse.

4b.       Exospore with spinose ornamentation.

5a.       False veins present; lamina ca. 1.5 m, with 9–12 pairs of pinnae .......................  6. A. itoi

5b.       False veins absent; lamina 0.5–1.2 m, with 4–6 pairs of pinnae
                                                                                                         7. A. subrotundata

4a.       Exospore with rodlike ornamentation.

6a.       Pinnae subopposite ............................................................................  3. A. danaeoides

6b.       Pinnae clearly alternate, at least in apical part of lamina.

7a.       Pinna margins acutely dentate ...........................................................  4. A. tonkinensis

7b.       Pinna margins entire, undulate-crenate toward apex ..................................  5. A. somai

1b.       Rhizome erect, subglobose; lamina of mature plants usually bipinnate, rarely once pinnate, or tripinnate at base; stipes without pulvini in mature fronds; synangia usually restricted to margins (Angiopteris s.s.).

8a.       Stipe smooth.

9a.       False veins absent.

10a.     Mature lamina very large, 2–4.5 m, bipinnate or tripinnate ......................  21. A. esculenta

10b.     Mature lamina smaller, 0.8–1.2 m, usually bipinnate, sometimes once pinnate  22. A. sparsisora

9b.       False veins present.

11a.     False veins long, extending over halfway to costa.

12a.     Mature lamina up to 2 m, false veins extending ± halfway between margin and costa  17. A. wallichiana

12b.     Mature lamina longer than 2 m, usually reaching 4–5 m in length, false veins nearly reaching costa  18. A. evecta

11b.     False veins short, not extending much beyond synangium.

13a.     Veins dense, ca. 20 per cm ..........................................................  19. A. caudatiformis

13b.     Veins sparse, ca. 10 per cm ...............................................................  20. A. helferiana

8b.       Stipe tuberculate.

14a.     False veins absent.

15a.     Mature lamina less than 2 m.

16a.     Veins dense, 20–22 veins per cm ..................................................  8. A. confertinervia

16b.     Veins normally spaced, 10–19 veins per cm ...............................................  9. A. wangii

15b.     Mature lamina larger than 2 m.

17a.     Mature pinnae 1–2 cm wide .................................................................  10. A. fokiensis

17b.     Mature pinnae 3–4 cm wide .............................................................  11. A. hokouensis

14b.     False veins present.

18a.     Pinnules oblanceolate, widest above middle, (sub-)opposite .............  12. A. oblanceolata

18b.     Pinnules lanceolate to elliptic, widest at or below middle, subopposite to alternate.

19a.     False veins long, reaching at least halfway to costa.

20a.     Pinnule margins sharply dentate ....................................................  13. A. acutidentata

20b.     Pinnule margins entire, undulate ............................................................  14. A. remota

19b.     False veins short, reaching up to 1/3 to costa, often not beyond synangia.

21a.     Pinnae with more than 15 pinnules; sori less than 1 mm; lamina up to 2.5 m  15. A. cochinchinensis

21b.     Pinnae with less than 15 pinnules; sori 1–6 mm; lamina 2–4 m
.....................................................................................................  16. A. dianyuecola

1. Angiopteris bipinnata (Ching) J. M. Camus, Proc. Int. Symp. Syst. Pteridol. 35. 1989.

二回原始莲座蕨  er hui yuan shi lian zuo jue

Archangiopteris bipinnata Ching, Icon. Filic. Sin. 5: t. 203. 1958.

Lamina once pinnate to bipinnate, 80–50 cm; stipe 60–70 cm, with 1 frondless pulvinus, and 10–12 pairs of pinnae; pinnae 16–19 × 6–7 cm; pinnules 0–7 pairs, broadly lanceolate, 7–10 × 2.5–2.8 cm, bases cuneate, margins subentire, undulate, apices caudate. Veins clear, false veins absent. Sori medial, ca. 4 mm from margin, extending to costule, 1–2 cm, composed of 20–40 sporangia. Exospore with forked rodlike ornamentation.

* Forests; 1100–1300 m. Yunnan (Maguan, Malipo).

2. Angiopteris chingii J. M. Camus, Proc. Int. Symp. Syst. Pteridol. 35. 1989.

河口原始莲座蕨  he kou yuan shi lian zuo jue

Archangiopteris hokouensis Ching, Icon. Filic. Sin. 5: t. 204. 1958, not Angiopteris hokouensis Ching (1959).

Lamina once pinnate, 50–85 cm; stipe ca. 50 cm, with 4 or 5(–7) frondless pulvini; pinnae 2 or 3 (sub-)opposite pairs, elliptic, 15–20 × 5–7 cm, bases cuneate, margins serrulate-crenulate, apices caudate. Veins clear, false veins absent. Sori medial, ca. 1 cm; paraphyses longer than sporangium.

* Forests; 100–500 m. Yunnan (Hekou, Jinping).

3. Angiopteris danaeoides Christenhusz, nom. nov.

尾叶原始莲座蕨  wei ye yuan shi lian zuo jue

Replaced synonym: Archangiopteris caudata Ching, Icon. Filic. Sin. 5: t. 208. 1958, not Angiopteris caudata de Vriese (1853); Angiopteris recaudata Ching, nom. illeg.

Lamina once pinnate, 50–85 cm; stipe 20–50 cm, with 1 frondless pulvinus; pinnae 2 or 3 (sub-)opposite pairs, elliptic, 10–30 × 6–11 cm, bases cuneate, margins serrulate-crenulate, apices long caudate. Veins clear, false veins absent. Sori medial, ca. 1 cm; paraphyses longer than sporangium. Exospore with rodlike ornamentation.

* Guangxi.

4. Angiopteris tonkinensis (Hayata) J. M. Camus, Proc. Int. Symp. Syst. Pteridol. 34. 1989.

尖叶原始莲座蕨  jian ye yuan shi lian zuo jue

Protomarattia tonkinensis Hayata, Bot. Gaz. 67: 88. 1919; Archangiopteris tamdaoensis Hayata; A. tonkinensis (Hayata) Ching; Protangiopteris tamdaoensis (Hayata) Hayata.

Lamina once pinnate, 50–85 cm; stipe 40–45 cm, with 1 frondless pulvinus; pinnae 2–4 alternate pairs, elliptic, 20–25 × 4–5 cm, bases cuneate, margins sharply serrate, apices caudate. Veins clear, false veins absent. Sori medial, ca. 0.7–1 cm; paraphyses shorter than sporangium. Exospore with rodlike ornamentation.

Hainan [N Vietnam].

5. Angiopteris somai (Hayata) Makino & Nemoto, Fl. Japan, 1563. 1925.

原始莲座蕨  yuan shi lian zuo jue [or 台湾原始莲座蕨 tai wan yuan shi lian zuo jue]

Archangiopteris somai Hayata, Icon. Pl. Formosan. 5: 256. 1915; A. henryi H. Christ & Giesenhagen; A. henryi var. somai (Hayata) Tagawa; A. latipinna Ching; Angiopteris henryi (H. Christ & Giesenhagen) J. M. Camus (date?), not Hieronymus (1919); Protangiopteris somai (Hayata) Hayata.

Lamina once pinnate, 50–85 cm; stipe 20–70 cm, with 1(or 2) frondless pulvinus; pinnae 4–6, alternate to subopposite, (ob-)lanceolate, (15–)22–25(–28) × 3–6.5 cm, bases cuneate, margins undulate, crenate to dentate, apices caudate to acuminate, false veins absent. Sori medial, 1–2 cm, composed of up to 170 sporangia; paraphyses present, longer than sporangium.

* Forests; 1100–1500 m. Taiwan, Yunnan.

Typical populations of Angiopteris somai in Taiwan differ from the Yunnan specimens (described as Archangiopteris henryi) in the length of pinnae apices and the degree of crenation; however, this does not warrant recognition at the species level.

6. Angiopteris itoi (W. C. Shieh) J. M. Camus, Proc. Int. Symp. Syst. Pteridol. 35. 1989.

伊藤氏原始莲座蕨  yi teng shi yuan shi lian zuo jue

Archangiopteris itoi W. C. Shieh, J. Jap. Bot. 45: 165. 1970.

Lamina once pinnate, ca. 150 cm; stipe 60–100 cm, with 1 frondless pulvinus; pinnae 9–12 pairs, oblong, 25–30 × 3–3.5 cm, bases cuneate, margins undulate, apices caudate. Veins clear, false veins short. Sori medial, ca. 1 cm, with 40–90 sporangia; multicellular paraphyses present. Exospores with spinulose ornamentation.

* Taiwan.

7. Angiopteris subrotundata (Ching) Christenhusz, comb. nov.

圆基原始莲座蕨  yuan ji yuan shi lian zuo jue

Basionym: Archangiopteris subrotundata Ching, Icon. Filic. Sin. 5: t. 206. 1958.

Lamina once pinnate, 50–120 cm; stipe 18–70 cm, with 1 frondless pulvinus; pinnae 4–6 pairs, alternate, elliptic, 10–30 × 2.5–7.5 cm, bases cuneate, margins entire-undulate, apices caudate. Veins clear, false veins absent. Sori medial, 0.3–2 cm; paraphyses longer than sporangium. Exospores with spinose ornamentation.

* Yunnan (Maguan, Malipo, Xichou).

8. Angiopteris confertinervia Ching ex C. Christensen & Tardieu, Notul. Syst. (Paris) 5: 11. 1935.

密脉莲座蕨  mi mai lian zue jue

Lamina bipinnate, 1.5–2 m; stipe tuberculate; pinnae ca. 60 × 20–25 cm, with 12–20 pairs of pinnules; pinnules 4–10 × 1.5–3 cm, bases rounded to truncate, margins serrate, apices shortly acuminate. Veins clear, 20–22 lateral veins per cm, false veins absent. Sori ca. 0.5 mm from margin, composed of 8–12 sporangia.

Forests; ca. 200 m. Yunnan (Hekou) [N Vietnam].

9. Angiopteris wangii Ching in Chien & Chun, Fl. Reipubl. Popularis Sin. 2: 30. 1959.

王氏莲座蕨  wang shi lian zuo jue

Lamina bipinnate, dark green, 1–2 m; stipe tuberculate; pinnae narrowly ovate, 40–60 × 15–18 cm, with 25–27 pairs of pinnules; pinnules 12–15 × 3–5 cm, bases rounded to truncate, margins regularly serrate, apices acuminate, curved acroscopically. Veins clear, false veins absent. Sori 1–3 mm from margin, 2–4 mm, composed of 18–20 sporangia.

* Broad-leaved forests; 400–1600 m. Yunnan (Hekou, Malipo).

10. Angiopteris fokiensis Hieronymus, Hedwigia 61(3): 175. 1919.

福建莲座蕨  fu jian lian zuo jue

Angiopteris angustipinnula Ching; A. attenuata Ching, nom. illeg.; A. jiangxiensis Ching & J. F. Cheng; A. kwangsiensis Ching; A. lingii Ching; A. longipetiolata Ching; A. muralis Ching; A. officinalis Ching; A. omeiensis Ching; A. petiolulata Ching; A. shanyuanensis Ching; A. sinica Ching; A. subcordata Ching; A. tenera Ching.

Lamina bipinnate, 2–4 m; stipe tuberculate; pinnae 50–60 × 20–60 cm, with 35–40 pairs of pinnules; pinnules lanceolate, 7–9 × 1–1.8 cm, bases rounded to truncate, margins regularly serrate, apices acuminate, acroscopically curved. Veins clear, false veins absent. Sori 0.5–1 mm from margin, ca. 1 mm, composed of 8–10 sporangia.

* Broad-leaved forests; 400–1600 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangxi, Sichuan, Yunnan, Zhejiang.

11. Angiopteris hokouensis Ching in Chien & Chun, Fl. Reipubl. Popularis Sin. 2: 31. 1959.

河口莲座蕨  he kou lian zuo jue

Angiopteris brevicaudata Ching; A. cartilaginea Ching; A. consimilis Ching; A. crassa Ching; A. crenata Ching; A. fengii Ching; A. garbongensis Ching; A. grossodentata Ching; A. lateterminalis Ching; A. latipinnula Ching; A. parvipinnula Ching; A. taweishanensis Ching.

Lamina bipinnate, light green, 2–4 m; stipe tuberculate; pinnae narrowly ovate, ca. 60 × 6–20 cm, with 9–11 pairs of pinnules; pinnules 12–15 × 3–5 cm, bases rounded to truncate, margins regularly serrate, apices acuminate, curved acroscopically. Veins clear, false veins absent. Sori 1–3 mm from margin, 2–4 mm, composed of 18–24 sporangia.

Broad-leaved forests; 400–1600 m. Guangxi, Yunnan [Vietnam].

12. Angiopteris oblanceolata Ching & Chu H. Wang, Acta Phytotax. Sin. 8: 159. 1959.

倒披针莲座蕨  dao pi zhen lian zuo jue

Lamina bipinnate; stipe tuberculate; pinnae 60–70 cm, up to 30 cm wide, with 12–14 pairs of (sub-)opposite pinnules; pinnules oblanceolate, 12–18 × ca. 2 cm, bases rounded, margins densely shallowly crenate, apices acuminate. Veins clear abaxially, obscure adaxially; false veins short, just surpassing sori. Sori 0.3–0.5 mm from margin, 0.5–0.7 mm, composed of 10–12 sporangia.

* Forests. Hainan.

13. Angiopteris acutidentata Ching, Acta Phytotax. Sin. 8: 157. 1959.

尖齿莲座蕨  jian chi lian zuo jue

Lamina bipinnate, 1–2 m; stipe tuberculate; pinnae 50–60 × 20–25 cm, with 16–18 pairs of pinnules; pinnules 10–20 × 1.5–2 cm, bases asymmetrical, rounded, margins sharply serrate, apices acuminate. Veins clear, up to 15 lateral veins per cm; false veins long, ± halfway from margin to costule. Sori ca. 0.5 mm from margin, ca. 1 mm, composed of 10–12 sporangia. Exospore with reticulate ornamentation.

* Hainan.

14. Angiopteris remota Ching & Chu H. Wang, Acta Phytotax. Sin. 8: 159. 1959.

疏叶莲座蕨  shu ye lian zuo jue

Lamina bipinnate; stipe tuberculate; pinnae ca. 60 × 15–22 cm, with ca. 15 pairs of pinnules; pinnules 13–15 × 1.5–2 cm, bases rounded, margins nearly entire, slightly undulate, apices acuminate. Veins clear, up to 15 lateral veins per cm; false veins long, up to ± halfway from margin to costule. Sori ca. 0.5 mm from margin, ca. 1 mm, composed of 10–12 sporangia.

* Hainan.

15. Angiopteris cochinchinensis de Vriese, Monogr. Marattiac. 23. 1853.

琼越莲座蕨  qiong yue lian zuo jue [or 宽昭莲座蕨 kuan zhao lian zuo jue, 亚全缘莲座蕨 ya quan yuan lian zue jue, 长尾莲座蕨 chang wei lian zue jue]

Angiopteris caudipinna Ching; A. howii Ching & Chu H. Wang; A. subintegra Ching; A. venulosa Ching.

Lamina bipinnate, 1–2.5 m; stipe tuberculate; pinnae 40–70 × 15–35 cm, with 10–15 pairs of pinnules; pinnules lanceolate 12–18 × 1.5–2.8 cm, bases rounded to cuneate, margins dentate to shallowly crenate (to nearly entire), apices acute to caudate. Veins usually clear, sometimes obscure adaxially, ca. 10–15 lateral veins per cm; false veins short, not surpassing sori or up to 1/3 toward costule. Sori 0.3–0.5 mm from margin, 0.5–1.5 mm, composed of 7–15 sporangia.

Hainan [Vietnam].

16. Angiopteris dianyuecola Z. R. He & W. M. Chu, Fl. Yunnan. 20: 720. 2006.

滇越莲座蕨  dian yue lian zuo jue

Angiopteris repandula de Vriese var. latemarginata C. Christensen & Tardieu.

Lamina bipinnate, 2–4 m; stipe tuberculate; pinnae ovate-lanceolate, 100–240 × 100–150 cm, with 15–25 pairs of (sub-)opposite pinnules; pinnules oblong, ovate-lanceolate, or oblanceolate, 5–8 × 1–2 cm, bases rounded to cuneate, margins serrate, apices short acuminate to caudate. Veins clear abaxially, false veins present. Sori 2–4 mm from margin, 1–6 mm, composed of 14–20 sporangia.

Forests; ca. 100 m. Yunnan (Hekou) [Vietnam].

17. Angiopteris wallichiana C. Presl, Suppl. Tent. Pterid.: 22. 1845 (lectotype, designated here: K!, Wallich 187).

西藏莲座蕨  xi zang lian zuo jue [or 海金沙叶莲座蕨 hai jin sha ye lian zuo jue, 海南莲座蕨 hai nan lian zue jue]

Angiopteris fibrillosa Ching & Y. X. Ling; A. formosa Ching; A. hainanensis Ching; A. henryi Hieronymus; A. lobulata Ching; A. lygodiifolia Rosenstock; A. medogensis Ching & Y. X. Ling; A. rahaoensis Ching; A. sakurai Hieronymus; A. taiwanensis Ching.

Lamina bipinnate, 1–2 m; stipe smooth, streaked; pinnae 35–70 × 15–30 cm, with 15–18 pairs of spreading pinnules; pinnules lanceolate, 5–15 × 1–2 cm, bases rounded to cuneate, margins crenulate-entire to serrate, apices acuminate. Veins clear, sparse, ca. 10 lateral veins per cm; false veins present, usually longer than sori, extending halfway from margin to costule. Sori 0.3–1.1 mm from margin, 1–1.5 mm, with 8–18 sporangia.

Forests along streams; 300–1500 m. Hainan (Danxian), Taiwan, SE Xizang (Moto), Yunnan (Yingjiang) [N India, Japan, Nepal].

18. Angiopteris evecta (G. Forster) Hoffmann, Commentat. Soc. Reg. Sci. Gott. 12: 29. 1796.

兰屿莲座蕨  lan yu lian zuo jue [or 云南莲座蕨 yun nan lian zue jue]

Polypodium evectum G. Forster; Angiopteris acrocarpa de Vriese; A. alata Nadeaud; A. badia Ching; A. commutata de Vriese; A. durvilleana de Vriese; A. evecta var. alata H. Christ; A. oldhamii Hieronymus; A. palmiformis (Cavanilles) C. Christensen; A. pingpienensis Ching; A. robusta Ching; A. yunnanensis Hieronymus; Clementea palmiformis Cavanilles.

Lamina 2–5 m; stipe smooth; pinnae 60–80 cm, with 15–30 pairs of spreading pinnules; pinnules 7–20 × 0.9–3.5 cm, bases cordate, rounded to cuneate, margins crenulate to serrate, apices acuminate to caudate; Veins clear; false veins clear, extending nearly to costule. Sori marginal to ca. 1 mm from margin, ca. 2 mm, with 8–15 sporangia.

Broad-leaved forests, rainforest valleys, roadsides, slopes; 100–1200 m. W Guangxi, Taiwan (Lanyu), Yunnan  [New Guinea, Philippines, N Vietnam; Australia, Pacific islands (Melanesia, Polynesia)].

Angiopteris evecta is a widespread species with an unclear taxonomy.

19. Angiopteris caudatiformis Hieronymus, Hedwigia 61: 278. 1919.

披针莲座蕨  pi zhen lian zuo jue

Angiopteris crassifolia Ching (1959), not de Vriese (1852); A. crassiuscula Ching; A. magna Ching ex C. Christensen & Tardieu; A. majuscula Ching; A. megaphylla Ching; A. multijuga Ching; A. nuda Ching; A. pinnata Ching; A. vasta Ching.

Lamina bipinnate, 2–4 m; stipe smooth; pinnae 60–110 × 20–40 cm, with 14–18 pairs of pinnules; pinnules 15–25 × 1.5–3 cm, bases cordate, rounded, or truncate, margins usually serrate, apices acute to caudate. Veins dense; false veins present, rarely absent. Sori 1–2 mm from margin, 2–3 mm, composed of 12–18 sporangia.

Rainforests, broad-leaved forests; 300–1200 m. Yunnan [E Myanmar, Vietnam].

20. Angiopteris helferiana C. Presl, Suppl. Tent. Pterid.: 22. 1845.

楔基莲座蕨  xie ji lian zuo jue

Angiopteris latemarginata Ching; A. subcuneata Ching.

Lamina bipinnate, 2–3 m; stipe smooth; pinnae 60–80 × 20–30 cm, with 15–20 pairs of pinnules; pinnules 10–20 × 2–6 cm, bases cuneate, margins serrate to sharply serrate, apices acuminate to caudate. Veins sparse, ca. 10 per cm; false veins present, rarely absent. Sori 2–3 mm from margin, 1.5–3 mm, composed of 14–26 sporangia.

Seasonal rainforests, broad-leaved forests; 900–1400 m. Yunnan [Myanmar].

21. Angiopteris esculenta Ching, Bull. Fan Mem. Inst. Biol., Bot. 10: 1. 1940.

食用莲座蕨  shi yong lian zuo jue

Angiopteris acuta Ching; A. parvifolia Ching & Fu.

Lamina bipinnate, sometime tripinnate at base, 2–4.5 m; stipe smooth; pinnae 50–60 × ca. 20 cm, with 20–30 pairs of spreading pinnules; pinnules lanceolate, 7–15 × 1–1.8 cm, base cuneate, margins serrate, apices acute. Veins sparse, ca. 10 per cm, false veins absent. Sori almost marginal, with 6–8 sporangia.

* Xizang (Moto), Yunnan.

22. Angiopteris sparsisora Ching, Acta Phytotax. Sin. 20: 347. 1982.

法斗莲座蕨  fa dou lian zuo jue [or 疏脉莲座蕨 shu mai lian zuo jue]

Lamina pinnate to bipinnate, 85–130 × 60–70 cm, with up to 4 pairs of pinnae; stipe smooth; pinnae 45–65 × 18–70 cm, with 7–15 pairs of spreading pinnules; pinnules lanceolate, 7–16 × 1.5–2.5 cm, bases cuneate, margins serrate to crenulate, apices acute; Veins sparse, less than 10 per cm, false veins absent (or obscure). Sori 1–3 mm from margin, 1–2 mm, with 5–16 sporangia.

* Broad-leaved or open forests; 200–1600 m. Guangxi (Luocheng), Yunnan (Xichou).

It has been suggested that Angiopteris sparsisora is of hybrid origin, because it is putatively intermediate between the former genera Angiopteris and Archangiopteris. One of the reasons to merge the two genera is because a clear morphological distinction is difficult to provide.

[Add note re. synonym?: Angiopteris paucinervis W. M. Chu & Z. R. He, Acta Bot. Yunnan. 22: 399. 2000, nom. inval.]