BLUMEA VOL. 23, No. 1, 1976 18 Coryphopteris Gymnopoda (Baker

BLUMEA VOL. No. 1976 18 23, 1, of Mt. Kinabalu Coryphopteris gymnopoda (Baker) Holttum. Plant growing on the Kamborangah ridge at this R. E. 2200m, showing the arborescent habit characteristic of genus (phot. Holttum, 1972). BLUMEA 23 (1976) 18—47 Studies in the family Thelypteridaceae X. Thegenus Coryphopteris R.E. Holttum Summary known of this with for and for identification. All species genus are described, synonymy each a key Seventeen new species are described: C. andersonii, C. andreae, C. arthrotricha, C. atjehensis, C. athyrioides, C. borealis, C. brevipilosa, C. hubrechtensis, C. iwatsukii, C. kolombangarae, C. meiobasis, C. microlepigera, C. seemannii, C. C. tahanensis, C. C. vitiensis. Allother C. subbipinnata, tanggamensis, species, except hirsutipes, and C. C. viscosa are transferred from other to Coryphopteris. The following names are pectiniformis , , genera A. D. indochinensis D. A. reduced to synonymy: Dryopteris megalocarpa v. v. R., Chr., subviscosa v. v. R., C. kinabaluensis linearis D. supravillosa Chr., D. Copel., D. Copel.,D. villosipes Gepp, D. rigidifolia v. A. v. R.; Lastrea ridleyi Bedd., L. robinsonii Ridl.; Nephrodium macgregorii Bak.; Thelypteris angulariloba Ching, T. simozawae Tagawa, T. herbacea Holttum, T. subglanduligera Ching. New varieties are described in C. viscosa, C. gymnopoda, C. pectiniformis, and C. pubirachis. Coryphopteris Coryphopteris Holttum, Blumea 19 (1971) 33. — Parathelypteris sect. Melanostipes Ching, Acta Phytotax. Sinica 8 (1963) 301, p.p. Caudex erect, commonly 15—30 cm tall, bearing roots throughout as in Cyathea, hairs stipes dark throughout or paler distally, in a few species bearing spreading septate with rather broad which near the base, always scales in most species lack acicular hairs. Lamina commonly 20—40 cm long, rarely to 80 cm, apex never pinna-like; lowest pinnae not or little reduced (several pairs gradually much reduced in C. fasciculata and narrowed towards their and wider C. squamipes), bases often at their middle than other basal lobeoften and pinnae, acroscopic enlarged dentatewith some forked veins, sometimes free; aerophores at bases of pinnae somewhat swollen; pinnae always deeply lobed, one basal both basal or more lobes sometimes free; veins in lobes simple or rarely once forked, veins to edge above base of sinus; lower surface of rachis, costae, costules, and veins reduced scales acicular hairs always bearing (smallest filiform); spreading usually present on lower surface of axes of frond (sometimes replaced by short capitate hairs), in some hairs which be sessile resinous cases longer may septate; glands (which may collapse on abundant lower drying) in many species ± on surfaces; costa always grooved on upper surface; hairs on upper surface of rachis and costae always acicular, normally unicellular, few sessile acicular hairs in a species septate; glands or capitate or sometimes present on veins. upper surface between Sori usually with rather large indusia (absent in C. engleriana) which be distal may glandular or hairy, sori (rarely all) sometimes ± asymmetric as in hairs Athyrium; sporangia short-stalked, never with glands or near annulus, sometimes IX in —68. *) Kalikasan 4, 1 (1975) 47 ") 50 Gloucester Court, Kew Road, Richmond, Surrey, TW9 3EA, England. BLUMEA VOL. No. 20 23, 1, 1976 this with a sessile gland on the stalk (in C. badia non-glandular hairs of several cells in continuous and a few position); spores usually pale, translucent, with a ± wing cross- wings (in C. klossii many small wings). I.e. Type species: Coryphopteris viscosa (Baker) Holttum, 1971. Distribution: mountains of the Malayan region (except Java and the Lesser Sunda Islands) and some island-groups of the Pacific including New Caledonia; N. E. India to southern China and Japan. Fl. Cytology: 11=32 (1(C. pectiniformis; Manton in Holttum, Rev. Malaya 2, 1955: 624, fig. 8.). I with In 1971 (I.e. pp. 18, 19) compared Thelypteridaceae Cyathea (sensu lato), pointing in out similarities. Assuming the relationship to be a real one, the following characters in of Coryphopteris maybe regarded as primitive: erect caudex; presence some species almost sessile without septateacicular hairs on upper surface of rachis and costae; sporangia annulus. Plants of all small appendages near species are in stature, with simply pinnate all fronds (partly bipinnate in C. habbemensisand C. stereophylla in New Guinea) and, as in than that of But of all the Old Thelypteridaceae, a much simpler vascular system Cyathea. World members of look like — a Thelypteridaceae they to me most a primitive group has the buthas uniform group which doubtless changed in course of evolution preserved a arborescent habit. In the New World and Africa the comparable genus is Amauropelta of which similar in habit Kunze, most species are very to Coryphopteris though they and mostly have gradually reduced lower pinnae a few have developed a prostrate rhizome. distinct shown for Amauropelta certainly represents a quite evolutionary line, as the other Old World of the example by its peculiar spores. Among genera family as few have is such habit recognized by me in 1971, species ofarborescent habit, and in none in of The which alone in present a majority species. genus Trigonospora, Thelypteridaceae has caudex has trilete spores, uniformly a short erect which never attains any considerable rocks in height. This is doubtless dueto the peculiar habitat ofall species, which grow on stream-beds liable with and beside where plants are to periodic flooding swiftly-flowing water. The African Menisorus is related and has a similar habitat. The habitat of Coryphopteris is also distinctivebut quite different, being confined to the of — rather low (sometimes mossy) forest of the crests ridges of higher mountains (1000 mineral- 3000 m altitude) where the soil is much leached and plants needto be adapted to a that deficient water-supply and probably also to high acidity. It is notable no species ash occur in Java, where all high mountains are volcanoes and where volcanic replenishes all the mineral nutrients in almost soils. It may be that Coryphopteris owes its survival to this adaptation, which allows its species to flourish under conditions where few other of where mountains of plants can compete. In the Pacific, species Coryphopteris only occur sufficient height and age provide the right soil conditions (Solomons, Fiji, Samoa, New Caledonia, Marquesas, Tahiti, Austral Islands). The with their 47 species here recognized are nearly all Malesian, greatest diversity where there the of in New Guinea, is greatest development high mountains. It is quite been col- possible that further new species exist in all the islands; these plants have rarely lected with sufficient attention to detail. In mainland Asia there is only one variable species (C. hirsutipes) which is widely distributed, and C. petelotii which appears to be a distinct local ally in Tonkin. But in mainland Asia also there other species included in and Parathelypteris by Ching; these have slender creeping rhizomes, but sporangia spores similar of lamina the to those Coryphopteris. The sessile glands on the of two genera have rather different them a aspect, but to distinguish clearly would need more recondite techniques than are available to the herbarium botanist. I do not doubt that Parathelypteris R. E. Holttum: Coryphopteris 21 and related (as typified by P. glanduligera Ching) Coryphopteris are genera. The American species Thelypteris simulata (Davenp.) Nieuwl., certainly close to P. glanduligera, has & A. Amer. Fern agreeing with the n=32 (R. M. Tryon, Journ. 63, 1973: 68), only of and observation on a species Coryphopteris. But Parathelypteris beddomei (Baker) Ching P. nipponica (Fr. & Sav.) Ching have a different frond-form and the former has a different chromosomenumber n=3i. Coryphopteris plants mostly grow in places which are only accessible by much physical and of exist in visited effort, probably most those that are places never yet by botanical collectors. Anyone who wished to make a general study of them would need to be very also have for his energetic and considerable resources travel at disposal. But some localities be where they exist (notably in Malaya) are easily accessible and it is to hoped that their be peculiar habitat-conditions, and their response to such conditions, may studied. Owing to their peculiar needs, which are not yet understood, no plants have ever made new growth when transplanted to Kew. Attempts to establish them in cultivation would have to be made near a natural habitat, and perhaps would only be successful when some of of the habitat the the more important characteristics are understood, especially acidity and mineral content of the soil water. C. in sometimes the At least one species, badia, grows (always?) moss-cushions, on branches of small trees in the dwarf moss-forest of high ridges, and plants of several The of badia covered species have been found as epiphytes. bases stipes of C. are with from and the slender hairs which seem indistinguishable root-hairs, probably have same function; they are slender, shrivelled but glossy golden brown when dry, with usually C. in few irregular septa (the sporangia of badia are also peculiar having non-glandular hairs of several cells on their stalks). It is possible that 'root-hairs' occur on the bases of when the I noted this stipes of other species plants grow in moss, though have not in The much almost white and herbarium specimens. stouter multiseptate hairs, opaque, few other do which occur at the bases of stipes in a species (see key) perhaps not have function of C. similar but hairs the same as the hairs badia; shorter occur on the upper of fronds of some of these parts plants. Plants of this have much in genus the same general aspect all species, whetherliving or as dried specimens, and it is not easy to see how to distinguish clearly between species, distinct exist. Basal though undoubtedly many species pinnae are always deflexed (this is characters not an artefact due to drying for the herbarium).

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