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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 2 (artificial)

1 Veins partly or almost fully connected   (2)
+ Veins free   (6)
2 (1) Veins reticulate and forming regular areoles; sori scattered and attached along reticulate veins.   18 Dictyocline
+ Veins partly joining; sori orbicular or shortly linear   (3)
3 (2) Veins meniscioid.   17 Pronephrium
+ Veins goniopteroid   (4)
4 (3) Plants of indeterminate growth, usually with gemmae in pinna axils, these gemmae potentially developing into new plants; laminae usually with mixed simple and stellate hairs.   16 Ampelopteris
+ Plants of determinate growth; laminae with only simple hairs   (5)
5 (4) Segments with an elongate translucent membrane below sinus; laminae usually with orange or reddish orange, spherical or clavate glands abaxially; sori orbicular, indusiate.   14 Cyclosorus
+ Segments lacking translucent membranes below sinuses; laminae without glands abaxially; sori shortly linear, exindusiate.   15 Stegnogramma
6 (1) Tuberculate aerophores present at pinna bases abaxially   (7)
+ Tuberculate aerophores lacking at pinna bases abaxially   (8)
7 (6) Proximal pair of veinlets on segments or acroscopic veinlet reaching sinus, others reaching margin above sinus; laminae with only acicular hairs on both surfaces.   12 Pseudocyclosorus
+ Veinlets on segments all reaching margins above sinus; laminae ?with mixed hooked long hairs, except with acicular hairs.   9 Cyclogramma
8 (6) Segments each with a cartilaginous ridge at bottom of adjacent sinus and with a transparent membranous line below that   (9)
+ Segments without cartilaginous ridge at bottom of sinus, lacking a translucent membranous line below that   (10)
9 (8) Plants ?with acicular hairs; pinnae pinnatifid nearly to costae; segments falcate-lanceolate, proximal pair of veinlets only reaching nearby line of transparent membrane below sinus or to margins above sinus; sori close to costules.   11 Glaphyropteridopsis
+ Plants glabrous throughout; pinnae pinnatifid to 1/2 of distance to costules; segments triangular; proximal 3 pairs of veinlets reaching transparent membranous margins below sinus but not connected; sori not close to costules.   13 Mesopteris
10 (8) Laminae ovate-triangular or triangular, 3- or 4-pinnate; proximal pair of pinnae largest; veinlets not reaching margins; throughout with multicellular acicular hairs.   5 Macrothelypteris
+ Laminae oblong or broadly lanceolate, most often pinnate-pinnatifid, to 3-pinnate-pinnatifid, rarely simple or bipinnatifid; proximal pair of pinnae similar in size or smaller than more distal ones; throughout with unicellular acicular hairs   (11)
11 (10) Costae densely covered with persistent grayish white acicular hairs adaxially   (12)
+ Costae without unicellular acicular hairs adaxially (occasionally with sparse caducous hairs)   (14)
12 (11) Costae rounded and raised adaxially; sporangia each sometimes with one multicellular hair expanded at tip on distal part of sporangial stalks.   4 Metathelypteris
+ Costae grooved adaxially; sporangial stalks sometimes each with a sessile spherical gland on distal part or sporangia each with one seta near annulus   (13)
13 (12) Veins not expanded at ends, reaching margins; sori orbicular, indusiate; sporangial stalks sometimes with 1-3 sessile spherical glands on distal parts.   3 Parathelypteris
+ Veins with minute expanded hydathodes at ends and not reaching margins; sori oblong or shortly linear, exindusiate; sporangia each usually with a seta near annulus.   7 Craspedosorus
14 (11) Rachises and costae with few small scales   (15)
+ Rachises and costae without scales   (16)
15 (14) Scales on rachises and costae with glands at apices, without cilia; pinnae sessile at bases, not adnate to rachises.   2 Oreopteris
+ Scales on rachises and costae without glands at tips but ciliate below tips; pinna bases adnate to rachises and decurrent.   6 Phegopteris
16 (14) Plants of marshes; veinlets forked; sori indusiate; sporangia each with several short setae near top of annulus.   1 Thelypteris
+ Plants terrestrial; veinlets simple; sori exindusiate; sporangia each with several acicular hairs   (17)
17 (16) Sori orbicular; veinlets not reaching margins.   8 Pseudophegopteris
+ Sori oblong or shortly linear; veinlets reaching margins.   10 Leptogramma

List of Keys

  • List of lower taxa


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