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A Synopsis of the Fern Genus Pteris in Japan, Ryukyu, and Taiwan By

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A Synopsis of the Fern Genus Pteris in Japan, Ryukyu, and Taiwan By Bot. Mag. Tokyo 79: 283-292 (June 25, 1966) A Synopsis of the Fern Genus Pteris in Japan, Ryukyu, and Taiwan by Wang-Chueng SHIEH* ReceivedMarch 2, 1966 Pteris is an old genus which Linnaeus (1753,Sp. Pl. 2: 1073)established to include 12 species. According to Copeland (1947,Gen. Fil. 60), there are about 280 species predominantly distributed in the tropics and subtropics of the world. I have studied taxonomically the species of this genus indigenous in the chain of islands from Hokkaidoto Taiwan which includes Honsyu, Sikoku, Kyusyu, Yakusima, Amami-Osima, Okinawa, and many other islands, and attempted to set up a new arrangement of infra-generic grouping of the species. In my opinion the most stable and important character to divide this genus is the branching manner of the frond, namely pinnating or triparting. The venation, on the contrary, is unstable in some extent. The traditional classification of this genus has long been based on this character. Presl (1836,Tent. Pter. 143) divided it into three genera-Pteris characterized by free venation, Cam~teria by forming a single series of costal areoles, and Lithobrochia by showing a more ample reticulate venation. Christensen (1906, Ind. Fil.) also used the same criterion and divided it into three subgenera-Eupteris, Campteria, and Lithobrochia. However, later, he (1932, Pter. Madagasc. 126) recognized that such division is unnatural and suggested that the venation must be used as a character for species, not for super specific groups. Walker (1962,Evolution 16: 27) 'commented that Christensen's division had certain practical uses, but it was purely artificial and in no way truly represented the biological situation, because the distinction among the three subgenera is liable to break down. Walker's view may be correct, although he did not propose any new classification, as he mentioned the costal arches are inconstant in Pteris biaurita. I also observed furthermore the same case in P. linearis and P. tokioi. In connection with P. linearis, Ching (1933, Lingnan Sci. Journ. 12: 568) illustrated the range of variation in venation of this species. Besides these two characters, the followings are also important, and accepted in my study. They are : the scales, margin of the pinnae, relation between the vein tips and the margin, presence or absence of the false-veins in mesophyll, dimorphism of the frond, base of the pinnae, etc. For the purpose of studying, I have examined all the herbarium specimens of Pteris available in the Botanical Institute, Faculty of Science, Kyoto University, Kyoto (KYO), Botanical Institute, Faculty of Science, University of Tokyo (TI), and Botanical Institute, Faculty of Agriculture, University of Tokyo (TIA). This work was carried out under the direction of Prof. H. Ito of Tokyo University of Education, to whom I wish to express my sincere appreciation for his kind guidance and en- couragement. I am much obliged to the directors and curators of all the abovecited herbaria for permitting me to use their facilities. A special debt of thanks is owed to Dr. M. Tagawa of Kyoto University for his valuable suggestions. Grateful acknowlege- ment is due to National Council on Science Department of Chinese Government for * BotanicalInstitute , Facultyof Science,Tokyo University of Education,Tokyo, Japan. 284 Bot. Mag. Tokyo Vol. 79 supplying me full expences for two years and working this study possible in Tokyo. Key to the subdivisions of Pteris 1. Frond pinnate or bipinnatifid .......................................... Subgen. 1. Pteris 2. Frond pinnate or rarely bipinnatifid, terminal pinna usually undivided, with or without false-veins in mesophyll ...........................................Sect. 1. Pteris 3. Margin not entire, pinnae distinctly dimorphous or rarely not 4. Base of pinnae cordate, scales pale brown densely on stipe and rachis..... ..........................................................Subsect. 3. Pteris 4. Base of pinnae obtuse or acute, scales brown mainly on base of stipe 5. Without false-vein in mesophyll .................... Subsect. 1. Creticae 5. With false-veins in mesophyll ....................... Subsect. 2. Cadierii 3. Margin entire, pinnae little dimorphous 6. Vein tips not reach to margin, stipe-scales pale brown belt-shaped ......... ...................................................... Subsect. 4. Formosae 6. Vein tips resch to margin, stipe-scales brown cuneate ...................... .................................................... Subsect. 5. Longipinnae 2. Frond bipinnatifid, terminal pinna pinnatifid, little dimorphism between sterile and fertile pinnules, without false-vein in mesophyll .............. Sect 2. Campteria 7. Pinnules entire, vein tips reach to margin, margin of stipe-scales with thin membrane................................................ Subsect. 2. Campteria 7. Pinnules not entire, vein tips not reach to margin or rarely do, margin of stipe- scales without thin membrane......................... Subsect. 1. Fxcelsae 1. Frond triparted with each division bipinnatifid, margin not entire ...................... ............................................................... Subgen. 2. Tripedipteris 8. Veins anastomose in various degrees, tips reach to margin .... Sect. 2. Tripedipteris 8. Veins all free, tips not reach to margin ...................... Sect. 1. Hypsopodium Subgenus 1. Pteris Frons parvula vel ampla ovata vel oblonga pinnata vel bipinnatifida non tripartita, pinnis margine integris vel dentatis, costis ad basin supra spinulosis vel non, venis omnibus solutis vel in gradis variis anastomosantibus, pinnis leviter vel dare dimorphis, falsivenis in mesophylla vel non. Typus : Pteris longifolia L. Sectio 1. Pteris Frons pinnata vel bipinnatifida, pinna terminali usitate indivisa, pinnis lateralibus leviter vel dare dimorphic, falsivenis in mesophylla vel non. Typus : Pteris longifolia L. Subsectio 1. Creticae Shieh, subsect. nov. Frons pinnata vel rare bipinnatifida, pinnis leviter vel pluris saepe dimorphis margine dentatis, venis ultimis fere ad margines non attingentibus. Typus : Pteris cretica L. Key to the species 1. Frond bipinnatifid, apex of sterile segment obtuse .................... (6) P. ensi formic 1. Frond pinnate, apex of sterile segment acute, caudate or tapering 2. Frond palmate, lateral pinnae 1-2 pairs 3. Sterile pinna less than 0.7 cm wide, apex tapering............. (1) P. dactylina 3. Sterile pinna more than 1.0 cm wide, apex acute 4. Margin dentate............................................. (2) P. deltodon 4. Margin spinose-serrate ....................................(3) P. nipponica 2. Frond pinnate, lateral pinnae more than 3 pairs 5. Upper pinnae decurrent .........................................(5) P. calcarea 5. Upper pinnae not decurrent 6. Apex of sterile pinna acute............................... (3) P. nipponica 6. Apex of sterile pi_nna caudate or tapering June, 1966 SHIER, W.-C. 285 7. Sterile pinna less than 2 cm wide ........... ( 4 ) P. cretica (var. cretica) 7. Sterile pinna mone than 2 cm wide ............. (4a) P. cretica var. laeta (1) Pteris dactylina Hook., Sp. Fil. 2: 160, t. 130-A. 1858 (based on specimens from eastern Nepal and Sikkim-Himalaya)-Ching, Ic. Fil. Sin. 3: t. 141. 1935. Hab. Taiwan. Distr. north India, west China. (2) Pteris deltodon Bak. Journ. Bot. 26: 226. 1888 (based on Faber no. 1010 from Mt. Omei, 3500 ft., Szechuan, China)-Hu et Ching, Ic. Fil. Sin. 1: t. 32, 65. 1930- Tagawa in Acta Phytotax. Geobot. 14: 45. 1950-in Journ. Jap. Bot. 36: 207. 1961- P. trifoliata Christ in Bull. Herb. Boiss. 7: 7. 1899 (type from Yunnan, China). Hab. Japan : Pref. Kumamoto ; Taiwan. Distr. China. (3) Pteris nipponica Shieh, sp. nov. Rhizoma breviter repens, apice squamis castaneis linearisubulatis integris ad 4 mm longis dense obtectum. Frondes caespitosae. Stipites graciles glabri straminei vel subtus praecipue castanei, supra sulcati, basi squamis castaneis linearisubulatis ad 5 mm longis parce obtecti; stipitibus frondum sterilium 11-30 cm longis, fertilium 14-50 cm longis. Laminae steriles coriariae, utrinque glabrae fusco-virides vel interdum variegatae palmate tridivisae vel pinnate 5-7 divisae, pinnis infimis saepe longe ornatis, pinnis lateralibus a se 1.5-6 cm remotis, inferioribus generaliter bipartitis, partibus posterioribus brevioribus quam anterioribus, pinnis terminalibus lineari-lanceolatis vel linearibus 7-23 cm longis 1.7-2.7 cm latis, apice acutis basi cuneatis sessilibus vel petiolulatis, petiolulis 0.5-3.0 cm longis, pinnis lateralibus brevioribus quam pinnis terminalibus 5-16 cm longis 1.5-2.7 cm latis polymorpis (ovatis, ovato-oblongis, vel lanceolatis), ascendentibus subfalcatis, apice acutis vel acuminatis basi acutis vel obtusis sessilibus vel brevissime petiolulatis, margine irregulariter argute denticulatis, costis pinnarum utrinque elevatis supra sulcatis, venis a costis angulo 55-70° divaricatis simplicibus vel furcatis. Lamina fertiles pinnatae laminarum sterilium similes utrinque glabrae fusco-virides vel interdum variegatae, pinnis linearibus 0.5-1.3 cm latis apice acuminatis serrato-denticulatis, pinnis terminalibus 13-30 cm longis basi sessilibus vel petiolulatis, petiolulis 0.5-3.0 cm longis, pinnis lateralibus 11-24 cm longis basi acuminatis sessilibus plus minus decurrentibus, inferioribus 11-20 cm longis basi sessilibus vel brevissime petiolulatis, non decurrentibus, interdum bipartitis, partibus posterioribus 4-10 cm longis, costis pinnarum utrinque elevatis supra tenuiter sulcatis, venis a costis
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