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26. Thelypteridaceae

金星蕨科 jin xing jue ke

Authors: Youxing Lin, Li Zhongyang, Kunio Iwatsuki & Alan R. Smith

Plants terrestrial or on rocks. Rhizomes stout, dictyostele radially symmetrical, branched or not, erect, ascending, or long creeping, with scales at apices; scales basiflexed, lanceolate or nearly ovate, brown, thick, luminae elongate, usually with grayish white short setae on dorsal side or ciliate along margins. Fronds clustered, approximate, or remote; stipes slender, stramineous, not articulate, with 2 crescent-shaped vascular bundles at base, usually scaly at bases, distally ± with grayish white unicellular acicular hairs, rarely with multicellular long hairs or stellate hairs. Fronds monomorphic, rarely subdimorphic, oblong-lanceolate or oblanceolate, sometimes ovate or ovate-triangular, usually pinnate-pinnatifid, sometimes 3- or 4-pinnate-pinnatifid, rarely 1-pinnate; pinnae symmetrical at bases; costae grooved adaxially but grooves not confluent with rachial grooves, or raised and with dense grayish acicular hairs, with expanded tuberculate aerophores at bases of pinnae. Laminae herbaceous or papery, sometimes somewhat leathery, green or dark brown-green when dry, both sides (particularly rachises, costae, and main veins adaxially) with grayish white unicellular acicular hairs, rarely glabrous, usually with orange or reddish orange, stalked or sessile spherical or club-shaped glands, occasionally small scaly along rachises and costae abaxially. Sori orbicular, oblong, or shortly linear, dorsifixed on veins, indusiate or exindusiate; indusia orbicular-reniform, fixed by deep notch, most ± hairy, persistent or hidden in sori, caducous, or not concentrated into sori but scattered along reticulate veins and exindusiate. Sporangia long stalked, usually with hairs or glandular hairs below annuli and at distal end of sporangial stalks. Spores bilateral, rarely tetrahedral, tuberculate, echinate, granular, or usually with a winged perispore. Prothalli green, cordate or narrowly cordate, usually with broad wings, symmetrical, usually with hairs or glands. x = 27-36 (lacking 28).

About 20 genera and ca. 1,000 species, more at lower elevations, very few tropical species above 4500 m: widespread in all tropical and subtropical zones of the world, less common in temperate zone, particularly more common in Asia; 18 genera (one endemic) and 199 species (102 endemic) in China.

The family* is very natural and is distinguished from others by having grayish white unicellular acicular hairs and pubescence throughout the plant. However, there are many different viewpoints about generic circumscription in the family. Ching recognized 18 genera (including Hypodematium) in his 1963 treatment (Acta Phytotax. Sin. 8: 289-335); soon afterward, in 1978 (Ching, Acta Phytotax. Sin. 16(3): 12-13), the number of recognized genera in China grew to 20 (Hypodematium was removed and placed in its own family). In 1971, Holttum subdivided this family in the Paleotropics into 23 genera (Blumea 19(1): 17-52). In 1977, Pichi Sermolli, mainly following Holttum, circumscribed other genera for a total of 32 genera (Webbia 3(2): 213-512). In 1990, A. R. Smith divided the family into five genera (in Kramer & Green, Fam. Gen. Vasc. Pl. 1: 263-272), i.e., Thelypteris (including five subgenera), Phegopteris, Pseudophegopteris, Macrothelypteris, and Cyclosorus (including 20 subgenera). Of the many systems, those of Holttum and Pichi Sermolli divide the family most finely, with the greatest number of genera. Holttum (loc. cit.) segregated the following genera from Cyclosorus s.l.: Amphineuron, Christella, Pneumatopteris, and Sphaerostephanos. Recognition of these genera was based on several characters, including whether the proximal pinnae were shortened or not, and whether the sporangia and sporangial stalks bore hairs or glandular hairs. Holttum (loc. cit.) also segregated several genera (e.g., Parathelypteris and Coryphopteris) from the classical Thelypteris s.l., the free-veined thelypteroids, by characters that included rhizome habit, laminar glands, and chromosome base number. Smith (loc. cit.: 265) noted: "Although many of Holttum’s genera seem natural (i.e., monophyletic), a combination of characters must be used to circumscribe them. Some of the characters concern minute glands and hairs and require 30 × magnification or greater for observation. Others require that complete specimens be at hand (including lower part of blade and stem). Even then, identification to genus may be difficult, as generic lines are not always sharp." The question of generic delimitation within the family needs further study, but for now we adopt Ching’s system (1978), modified from Ching (1963).**

*In 1940, Ching established many new families (Sunyatsenia 5(4): 237), including Thelypteridaceae with 12 genera, but the names of these new families are nomina nuda and were not therefore validly published (Melbourne Code, Art. 38.1(a)). Only in 1970 were these families formally published by Pichi Sermolli.

**(1) Holttum set up the genus Amphineuron with mixed characters in 1971, which was also adopted by Ching in 1978. Among the 12 recognized species in the genus, three occur in China: the type species A. opulentum, which is similar to Cyclosorus in venation and lemon-yellow glands; A. immersum, which is similar to Parathelypteris species in venation; and A. tonkinensis, which has already been removed to a new genus, Mesopteris. As Holttum himself considered Amphineuron to be a provisional treatment, this genus is not adopted here.

(2) Trichoneuron Ching (Acta Phytotax. Sin. 10: 118, pl. 22. 1967) was based on a single gathering, and the collection locality was unknown. Later, this species was rediscovered by Prof. W. M. Chu in Pingbian, SE Yunnan. According to Prof. Chu’s study, it belongs to the genus Lastreopsis (Dryopteridaceae). It is, therefore, not included here.

Using only venation, the family may be divided into 3 tribes: 1. Tribe Thelypterideae Ching: Veins free; with two subtypes: (1) all veinlets reaching margins or nearly reaching margins above sinuses between segments, with the bottom of sinuses not cartilaginous (e.g., Parathelypteris); (2) proximal pair of veinlets from adjacent segments reaching cartilaginous sinuses but not united; or acroscopic veinlet of basal pair of veinlets reaching cartilaginous sinus, basiscopic veinlet reaching margin above sinus (e.g., Pseudocyclosorus and Mesopteris); 2. Tribe Goniopterideae Ching: Veins partly combined; at least proximal pair of veinlets from adjacent segments united to form triangular areoles and this vein union producing a long or short excurrent veinlet; additional areoles may be produced in the same row, by subsequent vein unions of veinlets; excurrent veinlets may connect with more distal vein unions, or with a translucent line leading to a sinus, thus forming oblique rhomboid areoles (goniopteroid venation) (e.g., Cyclosorus, Ampelopteris), or squarish areoles (meniscioid venation) (e.g., Pronephrium, the areoles without included veinlets; 3. Tribe Dictyoclineae Ching: Veinlets between lateral veins all combined into irregular squarish or pentagonal areoles and each areole sometimes with simple or forked included veinlets.

The following taxon is excluded from the present treatment, pending further research: Thelypteris calvata Ching (Bull. Fan Mem. Inst. Biol. Bot. ser. 2, 1: 313. 1949), described from Guangdong.

Shing Kunghsia, Chiu Peishi, Yao Guanhu & Lin Youxin. 1999. Thelypteridaceae. In: Shing Kunghsia, ed., Fl. Reipubl. Popularis Sin. 4(1): 15-317, 319-353.

Key 1

1 Veins partly or entirely reticulate   (2)
+ Veins free   (6)
2 (1) Veins entirely reticulate, with or without simple or forked included veinlets in areoles; sori scattered and attached along reticulate veins.   18 Dictyocline
+ Veins partly reticulate (venation goniopteroid or meniscioid), areoles produced all without included veinlets; sori orbicular or shortly linear   (3)
3 (2) Veins meniscioid, i.e., all veinlets joining into square or rectangular areoles; pinnae large, broadly lanceolate; sori orbicular when young and usually confluent when mature.   17 Pronephrium
+ Veins goniopteroid, i.e., proximal pair of veinlets from adjacent segments or one pair of veinlets between veinlets of pinna joined to form into triangular areoles and producing an excurrent veinlet from these vein unions; excurrent veinlet or a line of translucent membrane at its end running to a sinus, several other pairs of veinlets connected to this excurrent vein or translucent membrane; pinnae small, narrowly lanceolate or triangular-lanceolate   (4)
4 (3) Pinnae pinnatifid; one long or short translucent membranous line at bottom of sinus on segments connected with excurrent veinlet, veins except proximal pair from second to fifth pairs of veinlets connected with excurrent veinlets or translucent membranous line and forming rhombic areoles, other veinlets reaching margins above sinus; laminae usually with orange or reddish orange spherical or club-shaped glands abaxially; sori indusiate.   14 Cyclosorus
+ Pinnae pinnatilobate or subentire, without translucent membranous line below sinus; except proximal pair of veins, other several pairs of veinlets connected with excurrent veinlet or reaching margins above sinus; laminae without glands abaxially; sori exindusiate   (5)
5 (4) Plants with gemmae in pinna axils, these gemmae potentially developing into new plants; laminae covered with both simple and stellate hairs; sori orbicular or suborbicular; sporangia glabrous.   16 Ampelopteris
+ Plants lacking gemmae; laminae with only simple hairs; sori thick and shortly linear; sporangia each with several setae at top.   15 Stegnogramma
6 (1) Sori indusiate   (7)
+ Sori exindusiate   (12)
7 (6) Helophytes; lateral veins forked.   1 Thelypteris
+ Terrestrial plants; veinlets usually simple (occasionally forked)   (8)
8 (7) Costae glabrous adaxially (or occasionally with sparse caducous hairs); stipes with many scales on proximal parts.   2 Oreopteris
+ Costae with dense persistent acicular hairs adaxially; stipes with sparse scales on proximal parts   (9)
9 (8) Costae rounded and raised adaxially; ends of veinlets not reaching margin; indusia small, or sometimes not developed   (10)
+ Costae grooved adaxially; veinlets reaching margins; indusia large, brown   (11)
10 (9) Laminae oblong or broadly lanceolate, 2- or 3-pinnate, proximal pair of pinnae similar in shape and size or smaller than distal ones, throughout with unicellular hairs.   4 Metathelypteris
+ Laminae triangular-ovate, 3- or 4-pinnate, proximal pair largest, throughout with multicellular hairs.   5 Macrothelypteris
11 (9) Pinnae without tuberculate aerophores at pinna bases abaxially; proximal pairs of veins reaching margins above sinus; laminae herbaceous, abaxially usually with reddish orange spherical glands.   3 Parathelypteris
+ Pinnae each with a tuberculate aerophore at base abaxially; proximal pair of lateral veins from adjacent segments with acroscopic veinlet reaching cartilaginous bottom of sinus; laminae papery or leathery, abaxially without spherical glands.   12 Pseudocyclosorus
12 (6) Sori orbicular   (13)
+ Sori thick and shortly linear, or oblong   (16)
13 (12) Laminae ovate-triangular, 3-pinnate; veinlets forked, not reaching margins; laminae throughout with multicellular long hairs.   5 Macrothelypteris
+ Laminae narrowly oblong or lanceolate, pinnate-pinnatifid; veinlets simple and reaching margins; laminae throughout with unicellular short hairs   (14)
14 (13) Rachises at pinna bases abaxially with brown tuberculate aerophores; laminae ?brown when dry; segments without cartilaginous ridge at bottom of sinuses.   9 Cyclogramma
+ Rachises at pinna bases abaxially without brown tuberculate aerophores; laminae green when dry; segments with a cartilaginous ridge at bottom of sinuses   (15)
15 (14) Plants ?with acicular hairs; pinnae pinnatifid nearly to costae; segments falcate-lanceolate, proximal pair of veinlets nearly reaching translucent membrane or margin above sinus; sori close to costules.   11 Glaphyropteridopsis
+ Plants glabrous throughout; pinnae pinnatifid to 1/2 of distance to costae; segments triangular, proximal 3 pairs of veinlets reaching margin of translucent membrane below sinus but not joined; sori not close to costules.   13 Mesopteris
16 (12) Sori thick and shortly linear; veins simple; segments entire   (17)
+ Sori oblong or suborbicular; lateral veins ?forked; segments or ultimate pinnules pinnatifid   (18)
17 (16) Laminae brown or brownish green when dry, with full spreading acicular hairs on both surfaces; proximal 1 or 2 pairs of pinnae free, distal ones adnate to costae; veinlets expanded at ends of veins and reaching margin; sori attached on middle of veinlets; sporangia each with 2-6 acicular hairs at tops.   10 Leptogramma
+ Laminae green when dry, subglabrous on both surfaces; pinnae completely free from rachises; veinlets expanded at ends and not reaching margin; sori attached near tips of veinlets.   7 Craspedosorus
18 (16) Plants small; stipes stramineous, not polished; laminae ovate-triangular or narrowly lanceolate; lateral pinnae decurrent along both sides of rachises and connected to each other; rachises and costae with more lanceolate scales, scales ciliate along margins; veinlets reaching margin.   6 Phegopteris
+ Plants usually taller and larger; stipes reddish brown or brownish stramineous, polished; laminae oblong, rarely broadly lanceolate; lateral pinnae not decurrent and free from each other; rachises and costae without scales; veinlets not reaching margin.   8 Pseudophegopteris

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