RE RE RT en EE ST NS EE SEEN : ~ PE HORE TARR Rey ee ces ax Noe Ty ye owhne nl Mefeerod WV I pees gab See ees STE ae RAY ‘ DPN RN TENG Be TE SON ST SS ONT STEEN TREN NES SE ’ . . Sine, Oe . 5 see a Yh ead a RL ors ee Vag ey RY Pwned “re wes, Bie wre my se ; er : ; 3 7 ~ e,! Yt ‘ 7 a AK Yessy SAGE EN 3 Bedrta CG Lae At A SN A ne vy YS NS ant ry’ et Pie a Nseniy . aie Se ry . 9 TN ae ie LAr Sea Yd ere ae ; Ne Ses *2 me YN et aw wt > NS aren : ¢ . yank XN ss yy? Ps Bie | Sa eS CPN eS Porm “pense eRe Or tamer DS SOA ra at ont ag a 4 ‘ ma ne : Amer ett, SN oe cons EY ted —, . Pe’, Seg eS EA ey 8, ame oe We ee re erent tae Sin te eh cl eae Ae Nal a LIBRARY FHE NEW YORK BOTANICAL GARDEN. BRONX, NEW YORK 10458 TAXONOMICAL REVISIONS _ | 23. U Death ; ; Map NA REPUBLIK INDONESIA REPUBLIC OF INDONESIA LEMBAGA ILMU PENGETAHUAN INDONESIA (L.LP.I.) INDONESIAN INSTITUTE OF SCIENCES FLORA MALESIANA BEING AN ILLUSTRATED SYSTEMATIC ACCOUNT OF THE MALESIAN FLORA / INCLUDING KEYS FOR DETERMINATION | DIAGNOSTIC DESCRIPTIONS | REFERENCES TO THE LITERATURE|/SYNONYMY/| AND DISTRIBUTION | AND NOTES ON THE ECOLOGY OF ITS WILD AND COMMONLY CULTIVATED PLANTS PUBLISHED UNDER THE AUSPICES OF LEMBAGA BIOLOGI NASIONAL BOTANIC GARDENS OF INDONESIA / BOGOR / JAVA AND OF THE RIJKSHERBARIUM / LEYDEN / NETHERLANDS PREPARED ON AN INTERNATIONAL CO-OPERATIVE BASIS UNDER THE SUPERVISION OF SEVERAL DIRECTORS OF BOTANIC GARDENS/ KEEPERS OF HERBARIA AND VARIOUS PROMINENT BOTANISTS FOR THE PROMOTION OF BOTANICAL SCIENCE AND THE CULTURAL ADVANCEMENT OF THE PEOPLES OF SOUTH-EASTERN ASIA TO THE SOUTHWEST PACIFIC REGION SERIES II PTERIDOPHYTA VOLUME 1 GENERAL EDITORS: Dr C. G. G. J. VAN STEENIS DIRECTOR OF THE FOUNDATION ‘FLORA MALESIANA’ & Dr R. E.HOLTTUM PUBLISHED BY MARTINUS NIJHOFF / DR W. JUNK PUBLISHERS THE HAGUE / BOSTON / LONDON 1959-1982 529 My Foo COPYRIGHT 1982 vA All rights reserved, including the right to reproduce this book or parts thereof in any form ISBN 90 247 2653 0 PUBLICATION DATES Part 1 10 Dec. 1959 Part 2 11 Nov. 1963 Part 3 20 April 1971 Part 4 15 Dec. 1978 Part 5 1 March 1982 Printed in the Netherlands CONTENTS A ETE SEL, REG ee eee: Gee Ree - ee a a ee me cei ee pee (3) RECTED LS Rare rth Wak CA al WwS, a Wis Lak rset icchcceke yeh Kgsane eit aniickasuecie, Gees (5) memenere yy Teil). EIOItin: fo). Steps is Milage net tia peapfanns ie LOS serve na aenee ee (7) GENERAL CHAPTERS muroduciony note: Dy iho HOKU. . ass) ee ee a eg eo see, Bee Sas. i went a, Aaclesian Pieridophytes by R.E.Holttum.. . . 2. ¢ 2 ciel cs Sco Se eS ee ii ite GO LUADGY Gai Re leas GlGiil —5 5 9 ecw oe nouc ou. Gonlmae cee o ono See ili eVCLANKCYUNO:l.to)Picropsiad by, Re Be Holttum = 6 4 sess, « Se 2 et ee ix GencralkeyeNo-2 to-Pteropsida by R-B- Holttummems = 5 - = = - = Ge 6s eee ee ee: Xii iKeys to the, genera of -Pteropsida by R. E. Holttum = =... ......0+%2+4s5:24.-48 XV PIDMOLTADLYIOY Rove SP ELOLUEUI: cer crac OS. ow 2s a ee eae eo ha) ore ee aeons Seals XXil TAXONOMICAL REVISIONS in alphabetical sequence Gyatheaccae bys EE. -HOlttum Fa fines es es ee eee ee 65 GicicheniaccacibyoR. BHOlttuni = so sos, 2a ey oy A ee eo 1 WSOCTACCOCID Ys Ac EA Gry AIStOM ye wet to oe renee ance ek cede eee) oe, 62 Pamsacaoroup by KU, Kramer o's 22. es, & Soe Se ee ee ee ee 177 Lomariopsis group by R. E. Holttum & E.Hennipman .............2.2.4... 255 Selnzacaccde by RB. Holttum one. sb hs eee ee ee ee. 37 mueiypicridaceae by. RE: Holttum’., ... 2-2.) 252.6 a PS eee eee 331 ADDENDA Addenda, corrigenda et emendanda by R. E. Holttum & K.U. Kramer ......... 561, 565 INDEX Index to scientific plant names by Mrs. M. J. van Steenis-Kruseman ...........-. 567 Dedicated to the memory of C.F. A. CHRISTENSEN DEDICATION CARL FREDRICK ALBERT CHRISTENSEN (1872-1942) was the founder of modern fern taxonomy. To appreciate the scope of his work, it is necessary to understand the confusions of thought on the subject which persisted through the 19th century and were still evident in the summary prepared (by Diets) for ENGLER & PRANTL’s Pflanzenfamilien in 1899. CHRISTENSEN’S first great work was his Index Filicum (1905-6) in which he listed all known fern binomials and also rele- gated many to synonymy. In the main he adopted the classification and nomenclature of DiEts. While preparing the Index he came to realize that many generic concepts accepted in the Index were unnatural or confused. This was especially evident in the great complex of species which he listed under the name Dryopteris. He next made a study of the tropical American species of that complex, and in so doing discovered how to separate them into natural groups (1913, 1920). At the time I first made contact with him (about 1925) he had begun to study ferns of the Old World tropics. I maintained a regular correspondence with him from 1925 to 1940, and sent him many specimens for identification. I also met him in Europe in 1930, 1934 and 1938 and had long discussions with him. I benefited from his wisdom also indirectly through the publications of R. C. CHING, who studied with CHRISTENSEN in 1929-1932 and applied CHRISTENSEN’S ideas to Chinese and Indian ferns in an important series of papers in the 1930s. CHRISTENSEN’s identifica- tions of my collections and his comments upon them were the basis on which my own work was built; in the present Series of Flora Malesiana I have tried to extend his methods and his ideas to a much wider range of species than he could have encountered. To him I am profoundly grateful, and I am concerned also to acknowledge my debt, through him, to some perceptive earlier workers, notably G. H. METTENIUS and JOHN SMITH. The objectives of any scheme of biological classification are to show natural relationships and to provide a means for the identification of individual organisms. It has sometimes been suggested that only the latter objective is important, and that a ‘practical’ scheme is all that is needed. The history of fern classification has shown that artificial schemes, made without thought as to relationships, do not work; and distribution-maps based on such schemes are meaningless. Fern classification as understood today should be based not only on gross-morphological characters but also on microscopical characters pertaining to the fern’s anatomy, indument, spores, gameto- phytes, efc., and on cytotaxonomy. There can be no doubt that existing ferns have originated through a process of evolution. They have therefore an inbuilt classification, and our object is to find it; the nearer we get to it, the nearer we are to the practical aspect of taxonomy. Fossils provide no clear evidence of the progress of the evolution of the great majority of leptosporangiate ferns. In most cases our only evidence for this is the natural relationships shown by taxonomy. We now have reached the stage at which most Malesian species can be allocated to definable natural groups which may have generic rank; most genera can also be associated in groups which appear to be natural; but it is often not yet clear how groups of genera are inter-related. For example, within the family Thelypteridaceae 1 cannot see a definite pattern of inter-relationships between the groups of genera which I have recognized. For a better understanding of this wider problem, genera throughout the tropics need to be taken into consideration; such an undertaking is beyond the scope of Flora Malesiana Series II, but I believe that this Series has presented a great deal of new evidence on which wider considerations may be based. Some botanists appear to think that a Flora is not the place for discussion of such questions. I disagree with that idea. Floras and taxonomic monographs always appear to account for every- thing, owing to the nature of their presentation. But in fact there are always gaps and uncertain- ties, especially in tropical Floras; some indication of this should be given; no classification can be final. (7) FLORA MALESIANA Even within groups of ferns already dealt with in Flora Malesiana, much more information is needed. For example, existing specimens do not show clearly the distinctions between species in the genus Plesioneuron (Thelypteridaceae). There is a great need for new collections made by persons who have specialist knowledge and are prepared to undertake prolonged critical search. After the publication of my book on the ferns of Malaya, Betry MOLESworRTH ALLEN, by per- sistent collecting, discovered nearly twenty additional species including representatives of three additional genera. The genera of LINNAEUS, which should be the basis of fern classification, are very crudely defined, and are only useful through agreement as to their typification. He did not notice indusia, upon which J. E. SMITH (1793) was the first to base new generic concepts, but SMITH also was not a critical observer. Within a few years, several other authors extended his observations and proposed new generic names, some not well distinguished, and in 1801 Swartz and BERNHARDI noted (more exactly than SmiTH) the differences between annulate (or gyrate) and exannulate sporangia. These observations were collated by Swartz in his Synopsis Filicum (1806) where he separated the genera of Osmundaceae, Schizaeaceae and Gleicheniaceae as spuriously gyrate, distinguishing the annulate genera (Polypodiaceae) solely by the form of indusia and the form and position of sori where indusia were lacking. An extreme example of the artificial nature of some genera proposed at this period is Belvisia MiRBEL (1802) which, in addition to the recognized type B. spicata (L.) MirBeEL (Polypodiaceae) included species now allocated to Actiniopteris, Schizaea, Asplenium and Ceratopteris. Simultaneous with SwARTZ, SCHKUHR was producing the first good series of illustrations of ferns (1804-1809). When one makes a drawing of a plant with the intention of accuracy, one often notices hitherto neglected characters. This was true of SCHKUHR, who observed and com- mented upon many details, especially of hairs, which have, since CHRISTENSEN, become recog- nized as important key characters. He noticed the jointed (septate) hairs of Ctenitis villosa (L.) CopPEL., and portrayed accurately the equally long unicellular hairs on the indusia of a species of Christella, though he did not comment on the latter. In some cases he illustrated venation clearly and accurately, in others indistinctly or casually. Some of SCHKUHR’s drawings were made from living plants, but most were from dried specimens. HOoKeErR and GREVILLE’s Icones Filicum (1827-1831) was the next illustrated work. The plates were better executed (by GREVILLE) than SCHKUHR’s and one can also detect a gradual increase of interest in detail as the series progressed. For example, plate 5 depicts Ceterach pedunculosum and plate 6 Grammitis decurrens, but in neither case are any details of venation shown; both species are now placed in the genus Colysis Prest (Polypodiaceae). Plate 125 shows Polypodium irioides, with enlarged details of venation well observed. Simultaneous with Hooker and GREVILLE, BLUME (1829-1830) was publishing the wonderful plates 1-65 of his Flora Javae, Filices, in which details are, on the whole, even more carefully dealt with. His subdivision of Polypodium is important. H. ScHoTrT, at the imperial palace of Sch6nbrunn, had living fern plants in his care and pub- lished (1834) a short series of excellent engravings illustrating new genera, showing much more detail than Hooker and GREVILLE; some of these were certainly based on living plants, in parti- cular his Nephrodium which shows very exactly capitate hairs and the elongate unicellular glands which are present on the stalks of sporangia, noted by no-one else until I made drawings of them in Singapore in 1943 (published 1971). Between 1840 and 1851 G. Kunze published a series of illustrations which he regarded as a continuation of SCHKUHR’s. He was in charge of the Botanic Garden at Leipzig, in which was the best collection of living ferns in Europe (soon to be sur- passed by Kew). His successor at Leipzig, G. H. METTENIUS, inherited KUNze’s living collections (8) Dedication and herbarium, and on their basis prepared a new system of classification of ferns (1856) with plates often showing new details. He subsequently prepared monographs describing all known species of several major genera, after which he began observations on the collections from Malesia in the Rijksherbarium. He had previously misinterpreted some of BLUME’s species through not having seen authentic specimens, and corrected some of them in the Ann. Mus. Bot. Lugd.-Bat.; he also incorporated new basic observations on several genera. While engaged on the latter work, he died of cholera in 1866 at the age of 42. Had he lived longer, he would have changed the course of pteridological thinking; I will revert to him later. C. B. PRESL was given the task of describing the collections made by HAENKE on the Malaspina Expedition; these included many specimens of ferns from the Philippines (described in 1825). As a result PRESL became interested in the classification of ferns, and realized that characters other than those of sori needed to be taken into account. In 1836 he published Tentamen Pterido- graphiae, comprising a new scheme of classification in which the arrangements of vascular tissue, and of venation, had an important place. His work is illustrated by many small drawings showing details of venation in relation to sori, in most cases quite accurately. His later publication (1848) showing arrangement of vascular strands in the stipes of ferns is not so well observed. PRESL’s emphasis on venation led him to associate together species of very diverse relationship, but it was a beginning of new thought. At the same time JOHN SMITH of Kew had been taking an interest in the cultivated tropical ferns in his charge, many raised (as at Leipzig) from spores from herbarium specimens. He was in touch with ROBERT BRowN, who had made some original observations on the venation of ferns when describing his own Australian specimens and also some collected by HorSFIELD in Java. SMITH was also in touch with FRANCIS BAUER, the Kew botanical artist, and supplied him with living fern plants and herbarium specimens, from which BAUER prepared a beautiful set of forty plates. These were submitted to W. J. HOOKER (then at Glasgow) who arranged for them to be published and added more, prepared by W. H. Fitcu, 120 plates in all, finalized after HOOKER came himself to Kew. Many of the genera are those of PRESL, but twenty were newly named and described by JOHN SMITH. SMITH himself had prepared a new scheme of classification indepen- dently of PresL, finding much agreement between their ideas when the Tentamen appeared; he collated his nomenclature with PREsL’s and his scheme was published by Hooker in 1841-1843. JOHN SMITH continued to study ferns, and to add to the collection of living plants at Kew. By 1865, when he was obliged through failing sight to retire, he had seen more than 1000 species of ferns in a living condition, of which he published a list in 1866. The final summary of his ideas, resulting from continued observation of living plants, appeared in 1875 and will be considered later. Having published Genera Filicum, HooKER planned Species Filicum, in which he proposed to describe all known species of ferns. For this, he had to re-consider the question of classification, and concluded that Pres had proposed too many genera; the result was that HOOKER’s genera in Species Filicum are almost the same as those of SWARTZ. The work was published in five volumes over a period of twenty years (1844-1864); Hooker planned to follow it with a summary in one volume, to be called Synopsis Filicum. He died just as the first part of the latter was printed. J. D. Hooker, who succeeded his father at Kew, engaged J. G. BAKER to continue the Synopsis on the lines planned, and this was completed in 1868 (second edition, with many additions by BAKER, in 1874). In 1891 BAKER published a summary of new ferns discovered since 1874, still with the same set of genera. HOOKER’s Species Filicum was illustrated by 304 excellent plates prepared by W. H. FitTcH (often two species on one plate). These show clearly and accurately details of venation and (9) FLORA MALESIANA indusia, but rarely any smaller structures. In his descriptions HOOKER rarely described details of hairs or scales. He thought such details unimportant; his main objective (see Vol. 3, p. 3) was ‘to assist the tyro in the verification of genera and species .. . natural habit is often a safer guide than minute microscopic characters’. He placed most exindusiate species of Thelypteroid, Tectarioid and Dryopteroid ferns in the genus Polypodium, but some in Gymnogramme and Meniscium; Dictyocline was merged with Hemionitis. He could not understand how JOHN SMITH could believe Brainea to be closely related to Blechnum, though it differs from Blechnum only in the absence of indusia. He placed Brainea between Gymnogramme (which included the diverse genera Selliguea and Syngramma) and Meniscium. His refusal to examine details led him to include in one species specimens which show great diversity in what are now considered to be significant characters. He united most of FEE’s species of Lomariopsis (including also Terato- phyllum MeEtv.), thus including several distinct Malesian species in Acrostichum sorbifolium L., of which the type came from the West Indies. His confusions in the synonymy of Thelypteroid ferns are very numerous, and can only be understood by reference to his herbarium. BAKER’s descriptions of ferns discovered after HOOKER’s death are even less satisfactory than HOOKER’s and often do not serve to identify specimens with any certainty. The remaining authors who proposed new schemes of classification were FEE (1852) and T. Moore (1857). FEe’s works were all admirably illustrated and his numerous plates show many significant details, but not always accurately. For example, in tab. XXI A, fig. 2 he was the first to show a transverse section of the stipe of Pleocnemia (sensu HOLTTUM 1974), but the accom- panying figure of venation in an allied species (fig. 1) is not accurate and fails to show the distinc- tive sinus-teeth. Neither FEE nor any earlier author (so far as I have observed) shows the distinc- tive row of four cells on one side of the sporangia of leptosporangiate ferns. FEE attempted to use the number of cells in the annulus as a generic character, but this is rarely practicable. His scheme is more elaborate than PRESL’s but is no nearer to a natural arrangement by present standards. He has Phegopteris as a genus separate from Polypodium, but in the same group of genera, not with its true allies, which are in other groups. Under the tribe Acrosticheae he has an astonishing diversity of genera. THOMAS Moore’s scheme is accompanied by good small drawings to show diagnostic characters. For example, he shows the difference in venation between Stenochlaena and Lomariop- sis, not noticed by FEE. But his scheme only differs in minor features from that of PRESL. MILDE in 1866 made important observations of scales and stipe-anatomy showing a clear distinction between Asplenium and Athyrium (including Diplazium); he elaborated these in 1870. METTENIUs had noted that previous attempts to distinguish these genera were unsatisfactory, and Hooker denied that any clear distinction was possible (and in 1928 Bower still copied HOOKER’s statement). In my judgement (HoLTTUM 1947) Asplenium and Athyrium are not very closely related. R. H. Beppome did not propose a new scheme of classification, but during the years 1856-1882 he made a more intensive field study of ferns in a tropical region than any previous author. He was critical of HooKEr’s genera and made some minor alterations in them for purposes of his Handbook (1883, with Supplement 1892), though still accepting the main scheme (his preface hinted that more needed changing). His work covered the Malay Peninsula and so is important for Flora Malesiana (he also accepted HooKER’s misidentification of some Indian ferns with species in Java). JOHN SmiTH’s Historia Filicum (1875) proposed a new scheme based on much study of living plants subsequent to his first one (1841). He did not use a microscope, and rarely refers to details of structure of sporangia, scales efc., but from observation of his plants he did learn much that (10) Dedication HOooKER never understood. I will refer later to some of his insights in a discussion of the work of Diets. SMITH and MetTeENtus, both observers of living plants, were the only authors of their period who (apparently independently) separated Phegopteris, Dictyopteris and other terrestrial exindusiate ferns from the alliance of Polypodium and transferred them to one including Aspidium. Both authors maintained separate genera for the exindusiate ferns, but JoHN SMITH admitted that probably some species were placed in Phegopteris and Dictyopteris merely because the only known specimens had old sori from which indusia might have fallen. Presumably he still thought the idea of uniting indusiate and exindusiate species in one genus too revolutionary. It should be noted that both METTENIUS and SmiTH had a mixture of Thelypteroid, Tectarioid and Dryopteroid ferns in their genera, whether indusiate or not; and SmitH kept Meniscium (Thelypteridaceae) far from his Aspidioid ferns. The major advance in thinking was that ind- usiate and exindusiate species could be closely allied; this was something HOoKER refused to consider. In this connection, the history of Pleocnemia leuzeana (GAUD.) PRESL is instructive. GAUDICHAUD described the species (from the Moluccas) in the genus Polypodium because its sori were exindusiate. PREsL founded a new genus based on the combination of a particular vein- pattern with circular exindusiate sori. Later CUMING collected specimens in Luzon which were similar in venation and general aspect, but some of them had indusiate sori. HooKER, who had illustrated the genus Pleocnemia as exindusiate (Gen. Fil. t. 70A, copied from Pres) published drawings made from two of CuMING’s specimens, one sterile and one showing indusiate sori (t. 97), and stated that this gave him an opportunity to correct his previous ‘error’ in reporting that P. leuzeana was exindusiate. But CUMING made four different collections (all seen by HOOKER) which are now known to represent three distinct species, two of them indusiate, one exindusiate, all different from the type specimen of P. Jeuzeana. HOOKER assumed that some specimens had lost the indusia which they originally possessed. F&E had specimens of the same collections from CumING, and speculated (1852, p. 311) on the strange fact that different plants of the same species could have, or lack, indusia. BEDDOME, examining plants of Pleocnemia from N.E. India which are in fact exindusiate (as seen from young sori) and belong to a species different from all three in the Philippines, thought that his Indian specimens must have lost their indusia and figured a fertile leaflet from a Philippine specimen (Ferns Br. India, t. 134). COPELAND in 1960 (p. 310) still only recognized one species in the Philippines, noting that the indusia are ‘sometimes fugacious’. Recent collections from Mt Makiling, at the foot of which CoPpELAND spent several years, confirm that CumING’s three species are distinct. The fronds are very large, so that only small parts can be put on herbarium sheets, and the stipe-scales (usually absent from herbaria) are distinctive. But herbarium specimens do show enough peculiar details if one knows what to look for, and the sum of these characters is sufficient to indicate that these species (and some others) form a genus distinct from Tectaria (to which CopELAND referred P. Jeuzeana), though the venation-pattern of Pleocnemia does occur in some species of the former. The sinus-teeth, which project out of the plane of the frond and are very conspicuous in living plants, were not noted by anyone except GAUDICHAUD until I re-defined the genus (HoLTTUM 1951, 1974); there are also distinctive glands (noted by METTENIUS but not by others). The petiolar vascular structure, also peculiar, was figured by FEE (1852, t. 21A fig. 2) and mentioned by no-one else. It was details such as the presence and nature of scales, hairs and glands on pinnae that METTENIUs noted; these have subsequently been found to be significant diagnostically, and they give METTENIUS’s specific descriptions a significance that is often lacking in BAKER’s. METTENIUS maintained large genera, perhaps (like CHRISTENSEN at a later time) because he did not want to publish new binomials until he was more sure of them; he subdivided his large genera much more (11) FLORA MALESIANA intelligently than Hooker, and made improvements in subdivision in his works published in 1864. DIELs erred in ignoring several important observations made by METTENIUS. The situation near the end of the century was that in most cases clear distinctions between groups of genera, and often even between genera now known not to be closely related, had not been discovered. This was due to a failure to understand that similar structures, whether of venation or sori, could have come into existence along different evolutionary lines. It is very clear that this is true of a simple type of anastomosis, seen in such genera as Acrostichum (s.str.), Pteris, Elaphoglossum, Lomagramma, Taenitis, Lindsaea and Hemionitis; in Pteris, Lindsaea and Elaphoglossum most species have free veins. The vein-pattern in Tectaria (Aspidium of Ind. Fil. 1905) and Microsorium (a segregate from Polypodium) is closely similar; in Malaya I found that up to 1955 a species of Tectaria had been included by all authors in Polypodium. It is also evident that acrostichoid ferns belong to several different alliances; and the acrostichoid condition is not exactly definable, so that authors disagreed in assigning genera to it. The sori of Davallia and Microlepia are very similar, but in other respects the plants are very different. An extreme case is Heterogonium PRESL, which I believe to be a natural genus (HOLTTUM 1949); some species have indusia, some not; some species have free veins, some have anastomoses; some have separate indusiate sori, some are acrostichoid. So the problem is to look for characters which may be a better guide to relationship than vein- patterns and sori. MILDE had shown the way by distinguishing between Diplazium and Asplenium on the basis of scales combined with vascular anatomy of the stipe. SmirH had noted that Polypodiaceae (s.str.) and the Davallia group of genera have a creeping caudex with stipes jointed to the dorsal surface of it; he coined the term Eremobrya for ferns of this habit; other ferns he called Desmobrya. The two terms were first defined in 1855. By this standard the ferns included by CHING (1940) and HoLtrum (1947) in Grammitidaceae are separate from Polypodiaceae. METTENIUS also found that the spores of the two groups differ (see below on DiE.s 1899). HERMANN CunrisT (1833-1933) was a lawyer who throughout a long life was actively interested in plants. He began to publish papers on ferns in 1890, and in 1897 produced Die Farnkrduter der Erde, an attempt to give a more balanced view of the more important species throughout the world than HooKeEr and BAKER. He recognized the nature of the problem stated in the preceding paragraph, but did not manage to do much towards solving it. He placed Aspidium and Phegop- teris (still separate genera, on the model of METTENIUS) in a family Aspidiaceae, distinct from Polypodiaceae, but under both Aspidium and Phegopteris had a great mixture of species not closely allied. In Polypodiaceae, tribe Acrosticheae, he had much confusion, especially in the genus Stenochlaena (see HOLTTUM 1978, pp. 261, 266); some of this was copied by DieELs. His later work also showed lack of critical insight. In his monograph of Elaphoglossum (1899) he tried to subdivide the genus on characters of venation, but did not examine the veins carefully and the result is confusion; in his paper of 1907 on the Philippine species of Dryopteris (the com- posite genus of Ind. Fil. 1905) he did not make good descriptions nor understand relationships between species. He did not know of MILDe’s work. The century closed with the volume of ENGLER & PRANTL’s Pflanzenfamilien covering vascular cryptogams, in which DiELs dealt with almost all the ferns (1899-1900). His Polypodiaceae con- sisted of nine tribes. He transferred several genera of the tribe Acrosticheae of some previous authors to Aspidieae, but mixed together Aspidioid and Polypodioid species under Polybotrya. He placed the Polypodioid genus Platycerium in Acrosticheae. He united Phegopteris with Aspidium but had a great mixture of species in it; his treatment only adds more confusion to an already very confused situation. He did understand MILDe’s work, but he failed to notice some important observations made by METTENIUS and JoHN SmiTH, of which the following are three examples. (12) Dedication 1. Gleicheniaceae. PRESL based his primary division of Gleichenia (sens.lat.) on the position of the sori on the veins, stating that in Eu-Gleichenia the sori were terminal, in all other cases dorsal on the veins. This division was copied by HOOKER, CHRIST and DIELS; but in 1856 METTENIUS had stated that the sori are not terminal in Eu-Gleichenia, and had repeated this in 1863. In the latter paper he divided G/eichenia into three subgenera, stating that two of them agreed in scales and in sporangia, the third (Dicranopteris) differing in both these structures. This was ignored by DiELs, who did not cite the paper of 1863 and mixed together in one subgenus species of Dicranop- teris with those which differed both in scales and in sporangia. When preparing an account of the family for Flora Malesiana (HOLTTUM 1959) I failed to notice METTENIUS 1863 and repeated his observations, differing only in the recognition of Dicranopteris as a genus distinct from Gleichenia, the latter having three subgenera; this is certainly the important division. 2. Stenochlaena and Lomariopsis. In 1875 (p. 140) JOHN SMITH stated the distinctions between these two genera (he had united them in 1841 and subsequently discovered the difference through observation of living plants). METTENIUS still included them in the same genus (1869, in a post- humous paper edited by KuHN) but in separate sections, and he established a new genus Terato- phyllum, distinct from both, with two species. DiELs united Stenochlaena, Lomariopsis and one species of Teratophyllum in one genus (in the tribe Asplenieae) which he divided into two sections: Eu-Stenochlaena comprising the whole of Lomariopsis and Teratophyllum aculeatum (BL.) METT., and Cafraria, which consisted only of S. tenuifolia; the latter differs from the type species S. palustris (BURM.) BEDD. in having bipinnate fertile fronds and in little else. This is an absurdly unnatural division. DieELs included the second species of Teratophyllum (T. articulatum (J.SM.) MetT.) in Polybotrya (tribe Aspidieae). 3. Grammitidaceae. This family was recognized as distinct by CHING in 1940; for fuller details see also HoLtrumM 1947 and 1955. Diets placed all species of the family in Polypodium sect. Eu- Polypodium, mixing them indiscriminately with true Polypodium species, except Prosaptia PRESL which he included in Davallia. BLUME in 1830, though retaining them in Polypodium, had already distinguished these ferns as ‘spurious’ in that genus. METTENIUS (1866) distinguished them in Polypodium as Div. 1, Sphaerosporeae, placing true Polypodium in Div. 2, Nephrosporeae; he did not mention Prosaptia in this paper, but had previously placed it with Davallia. As above noted, JoHN SMITH placed most Grammitoid ferns in his division Desmobrya, and thus separated them from Polypodium, but somehow he included Prosaptia (with the closely related Cryptosorus) in Eremobrya; he did however note that their sori were very different from those of Davallia. For some reason which is not at present understood, Grammitoid ferns are difficult to maintain in cultivation, and not one of them appears in JOHN SMITH’s list of species which he had seen alive; this probably accounts for his mistake in placing Prosaptia with Polypodium. METTENIUS always noted the peculiar hairs on plants of Grammitidaceae, and also the hairs on scales and setae on sporangia, where these occur (true Polypodiaceae never have these characters). When he died in 1866 he was just beginning to see the significance of such structures. Towards the middle of the 19th century academic botanists realized that taxonomic study, of the limited and formal kind which still prevailed, did not deal with important aspects of the life of plants. So they started ‘scientific botany’, but they made the mistake of thinking that taxonomy was an out-dated activity; many such botanists still persist in that mistake. What was needed was a better taxonomy, not its abandonment. This was especially true of tropical plants in general, and most ferns are tropical; significant facts about these plants had often not been put on record, or if recorded (such as the hairs figured by ScHKUHR) had not been understood. As ‘scientific’ botany diverged more and more from taxonomy, the shortcomings of the latter were less and less understood. A factor in this process was, and still is, the binomial system of nomenclature. (13) FLORA MALESIANA Valid names consist of two parts, a generic name and a specific epithet. Thus one must know the correct genus if one wishes to describe a new species. But in the case of tropical plants, which were very little known to earlier authors, it was impossible to be sure of generic concepts, which changed with increasing knowledge. Thus the binomial system, in theory, imposed an impossible condition for naming new species. In practice, this situation was avoided by allowing taxonomists to make the best guess they could, with permission afterwards to change the generic name if later knowledge so indicated. Morphologists rightly wished to study plant-structures not mentioned in taxonomic descriptions; taxonomists were slow to realize the need for this as a help to better taxonomy. An outstanding exception was METTENIUS, who published important works on lateral buds on ferns (1860), on the morphology and anatomy of Angiopteris in comparison with other ferns (1863A) and on Hymenophyllaceae (1864B). Morphologists who have not a wide knowledge of taxonomy are apt to think that any species is representative of the generic name it bears, and thus are liable to have erroneous ideas about genera (especially where such genera are still not clearly defined), and may be misled into making wrong comparisons or invalid statements about phylogeny. In view of the above discussion on the history of taxonomic study of the leptosporangiate ferns, it is evident that most 19th century taxonomy was an inadequate guide to morphologists. The most important morphologists were GorBEL and Bower. Bower began his studies in the 1880s, mainly on the more primitive ferns. When he came to his summary on the leptosporangiate ferns (1928) he quoted GOEBEL’s com- parison of their study to wandering in a dark and trackless forest, but he did not know enough about existing information which could have provided him with some guiding light. He did not know of the work of MiLpE and discussed the possible evolution of the sorus of Asplenium by reference to a species of Diplazium. He discussed Stenochlaena, which he interpreted according to the confused statement by CurisT, and described the anatomy of the rhizome, but the material he described belonged to a species of Lomariopsis, as he could have learned from JOHN SMITH. He placed Phyllitis ina group separate from Asplenium, not knowing that natural hybrids between the two existed. He accepted Curist’s comparison of Elaphoglossum with Syngramma, though the resemblance between the two is very superficial. He accepted the idea that the sorus of Microlepia was marginal in phyletic origin, but did not realize that this might also be true of Cyathea and Dryopteris. He insisted that Deparia was a natural genus, though each of the species included in it shows an alliance to a different group of ferns. He had not looked at CHRISTENSEN’S dismemberment of ‘Dryopteris’ and accepted a phyletic sequence (fig. 663 on p. 132) which derives the vein-pattern of Bolbitis (then still included in Leptochilus) from the condition of a Thelypteroid fern. But he did have a better understanding of the Gymnogrammoid ferns. GoeBEL had far more understanding than Bower, having spent at least two periods of study in Java (BOWER never went to the tropics), but he did not keep in touch with CHRISTENSEN’s work. As above noted, CHRISTENSEN made a systematic study of all the tropical American species included in the comprehensive Dryopteris of Index Filicum. In so doing he followed the example of METTENIUs in looking for details of dermal appendages, but more critically and more con- sistently than MeTTENIUus had done; he had also a much wider range of species to examine. In this process he discovered that the many species could be separated into groups according to the nature of their hairs; he rightly insisted that groups distinguished in this way show also many other differences of a less easily definable character. CHING (1936, p. 243) added the distinctive character of vascular anatomy of the stipe of Thelypteroid ferns, in which they constantly differ from Crenitis and Dryopteris (s.str.), as indeed METTENIUS had noted in his discussion of Aspidium in 1864. CHRISTENSEN (1911) expressed the opinion that some of the groups he had distinguished should be regarded as good genera, but he retained them in Dryopteris because he wished to (14) Dedication examine species of the Old World before publishing new combinations. This work of CHRISTENSEN’S was a turning-point in fern taxonomy. R. C. CHING applied CHRISTENSEN’s ideas to ferns of southeast Asia, clearing up much previous confusion. But Thelypteridaceae are far more abundant and more diversified in Malesia than in mainland Asia. When writing my book on the ferns of Malaya (1955) I adopted CHING’s generic concepts but stated (p. 236) that the resulting arrangement was not a natural one. I made new observations, especially of glands and hairs on sporangia (some not then published) but could not see my way to a good re-arrangement on the basis of the limited number of species in Malaya. It was only when I looked at all species in Malesia, mainland Asia and the Pacific (and also many previously unnamed collections), noting in detail structures not mentioned in earlier descriptions, that I was able to see how to improve on CHING’s scheme, and to establish new genera peculiar to the Old World. CHING’s work and mine (presented in the present volume) are built on CHRISTENSEN’s methods and on his insights. CHRISTENSEN subsequently identified a number of collections of ferns from Malesia and Asia (including my Kinabalu ferns, 1934) and wrote a fern flora of Madagascar (1932); in so doing he examined a large number of type specimens which had not been well described and published new information about them. The nomenclature in my book of 1955 was largely dependent on his observations on types. In 1939 he contributed a chapter on the classification of ferns to VERDOORN’S Manual of Pteridology. This contains many new ideas, the result of his wide-ranging studies; from it one can judge the progress made since 1905, largely due to his own work and thought. His last work was a fern flora of Samoa, published after his death (1943). Owing to the stress of the war situation, no adequate obituary notice was published. Pteridology in Malesia. The first considerable field work was on the ferns of Java, summarized by BLumeE in 1828 and elaborated with excellent illustrations in 1829-30 (additional plates were published in 1847 and 1851). Little more was published until RACIBORSKI went to Bogor and undertook new field studies in West Java, summarized in his book of 1898; his descriptions are better than most of their time and his ecological information is valuable. A few years later, VAN ALDERWERELT VAN ROSENBURGH began fern studies covering the whole of Malesia by collating all existing descriptions, most of them too brief or too inaccurate to form a good basis for the keys which he prepared. In the main he followed the nomenclature of Index Filicum, but he wanted more clear-cut distinctions between genera as a better guide to identification, and so he adopted an artificial system. He reverted to a comprehensive tribe Acrosticheae, and a tribe Phegopterideae widely separated from Aspidieae; he revived Pleocnemia Pres and included in it some Thelyp- teroid ferns. After completing his Handbook (1908) he continued a critical study of the specimens in the herbarium at Bogor, including many new collections, also plants in cultivation. He pub- lished descriptions of these in a series of papers, those up to 1917 being summarized in his Supplement. These new and amplified descriptions show a careful examination of much detail not previously recorded, including observations on spores ; many of his new species are still recognized in the present work. In Malaya H. N. RIpLey made large new collections in the years 1888-1911, but his published work on them (1926) is very uncritical; his generic and specific descriptions are confused and sometimes inaccurate; the names he wrote on herbarium specimens at Kew are often wrong. Thus his statements on distribution of species are also often wrong. It is sometimes impossible to know the basis of such statements because often he did not write names on herbarium sheets in Singapore; I re-arranged the sheets without noting in which covers RIDLEY had placed them. RIDLEY’s work on ferns is therefore usually ignored in Flora Malesiana except for his new names. E. B. CopELAND (1873-1964) began a study of ferns in 1893 but soon specialized in plant (15) FLORA MALESIANA physiology. He went to the Philippines in 1903 and during the years to 1917 made extensive field studies of ferns, also naming and describing collections made by others from the Philippines and other parts of Malesia. Between 1917 and 1928 he was concerned with rice cultivation in Cali- fornia; after that most of his active life was devoted to ferns. I have elsewhere (HOLTTUM 1973) summarized his work, which culminated in his Genera Filicum (1947). His observations on Philip- pine ferns, based on the same classification, were not published until 1960. He was the first person to understand that HooKker’s genera Cyathea, Hemitelia and Alsophila were unnatural, but his revised scheme for Cyatheaceae in 1947 was little better because he did not examine the detailed structure of scales. Similarly, in dealing with Thelypteridaceae (which he did not recognize as a distinct family) he did not look carefully at hairs and glands; his descriptions of species are little better than BAKER’s. His floristic work suffered also because he did not see the types of many of the older species and misconstrued some of them; this however does not excuse his failure to distinguish between Sphaerostephanos penniger (HooK.) HoLTtTuM and Pneumatopteris truncata (Poir.) Ho_trum (as named in the present work). Yet in Hymenophyllaceae he did make very careful detailed observations of structure, which were very fully illustrated (COPELAND 1933, 1937, 1938). His families Pteridaceae and Aspidiaceae of 1947 are both confused mixtures which are still not fully disentangled. In general, CopELAND’s failure was due to not looking for signifi- cant characters. His statement that the generic separation of Gymnocarpium dryopteris NEWM. from Phegopteris connectilis (MiCHx) WATT was hardly possible is an illustration of this. Simultaneously with CopELAND’s work in the Philippines, C. A. BACKER (from 1905) was making large collections of ferns, as part of his general herbarium of the flora of Java. He collaborated with O. PosrHumus, who had specialized on ferns and had made many collections in several other islands also, in the production of a fern flora of Java (1939). The nomenclature follows that of the third supplement of Index Filicum. In general, this is a considerable advance on VAN ALDERWERELT, though the descriptions of species in complex genera are not as good as VAN ALDERWERELT’S later ones. In Cyatheaceae the genera are not well distinguished and the specific descriptions are very inadequate; the authors could have learned much from METTENIUS (1863B). In Dryopteris no attempt is made to separate Thelypteroid species from the rest. The Grammitoid ferns are not separated from Polypodioids. The citation of synonyms is often un- critical. The work is of value mainly for its ecological information, but where species have been confused (e.g. under Dryopteris uliginosa) ecological information is also confused. The present situation. At the beginning of this volume (p. ii, 1959) I presented a tentative list of genera, remarking that much new information would arise in the course of study in the produc- tion of the present Flora, and that new ideas on classification would probably emerge. I refrained from assigning the majority of genera of leptosporangiate ferns to families. In part 2 (1963) I showed Dicksonia and Cyathea to be much more nearly allied than I had thought in 1959. The inter-relations between genera there presented still appear sound, but the assignment of genera to families is still uncertain. In part 3 (1971) K. U. KRAMER presented a major revision of the genera of the Lindsaea group, clarifying distinctions between them and making a new subdivision of Lindsaea. Part 4, on the Lomariopsis group (as delimited by me in 1947) included much new information, especially on Elaphoglossum and Bolbitis, with a revised account of my earlier work on the other genera. The present part attempts a new subdivision and conspectus of the Thelyp- teris group of genera, which is so sharply distinct from other groups that I judge it to deserve family status. This decision commits us in some measure to the ultimate recognition of other families, but for this we still need more evidence. I think that the elaborate arrangement of Picu1 SERMOLLI (1977) is premature, though it is much nearer to being natural than COPELAND’S of 1947. (16) Dedication We need more information about significant characters, and to be useful they must be avail- able for all species; but no-one can tell in advance which characters will be significant. Since MANTON’s book of 1950 the observation of chromosome numbers in a great range of ferns has provided important new evidence; but chromosome number by itself, without evidence of con- formity with some quite different characters, can be misleading, and the proportion with species with known chromosome numbers is still relatively small in some genera and not easily aug- mented. As a result of the work by MANTON and others it has become clear that phenomena like hybridization and polyploidization — formerly regarded to be extremely rare in ferns — com- monly occur also in tropical ferns. The impact of cytotaxonomic work on fern classification has recently been thoughtfully dealt with by Lovis (1977) and WALKER (1979). KLEKOwSKI (1979) has contributed much to our knowledge of the reproductive biology of the ferns also in relation to polyploidy. Morphologists can provide useful suggestions for characters of possible significance. Mor- phology and taxonomy are interdependent; morphology without a good taxonomy may arrive at wrong conclusions; taxonomy without the stimulus of morphology may miss important dis- tinguishing characters. Features which have recently been shown to be of great significance for fern classification are stomata (VAN COTTHEM, 1970) and spores (LUGARDON, 1971). Gameto- phytes, formerly a neglected item, are important and often indicate relationships within the larger groups of ferns. In the end, a practical taxonomy must rest on a limited number of characters which are observable without very elaborate equipment, which is one reason why uninformed academic botanists regard it as unscientific. I hope and believe that this Flora is producing new contribu- tions to that end. Work now in progress for further instalments of Flora Malesiana Series II is as follows. Dr E. HENNIPMAN, with collaborators, has begun a study of Polypodiaceae (s.str.); Prof. K. IWATSUKI is making progress with Hymenophyllaceae; Mr G. J. DE JONCHEERE is working on the Davallia group; Dr B. S. CROXALL is studying the complexities of Grammitidaceae; Prof. K. U. KRAMER has started on Preris, which seems to me to be an isolated genus; Prof. T. C. CHAMBERS has made a world monograph of the genus Blechnum and it is hoped that he will be able to deal with the Blechnum group for Flora Malesiana; Mr A. C. JERMy is working on the complex genus Selaginella. Of the other genera, Dennstaedtia is of basic importance. It is more diversified in the Philip- pines, New Guinea and the Pacific than in any other part of the world and, as the fronds are very large, existing herbarium material often does not give full information about them; more field work, by people who know what to look for, is needed. The most complex groups still not dealt with are those of Tectaria and Athyrium, the latter being very difficult, with need of much new observation, especially of scales. I have made studies of two genera of the Tectaria group and propose to continue with that group if I am able to do so. The Adiantum group (as listed in 1959) is complex, but not so well developed in Malesia as in drier climates, and is more dependent on studies of plants in such climates than are most other groups. This is true also of the Dryopteris group, which is mainly temperate in distribution. December 1979 R. E. HoLtruM (17) FLORA MALESIANA REFERENCES ALDERWERELT VAN ROSENBURGH, C. W. R. K. VAN: 1908. Malayan Ferns: Handbook to the determination of the ferns of the Malayan Islands. Landsdrukkerij, Batavia. — 1917. Ibid. Supplement. BACKER, C. A. & O. PostHuMus. 1939. Varenflora voor Java. ’s-Lands Plantentuin, Buitenzorg. BAKER, J. G. 1891. A summary of new ferns discovered since 1874. Ann. Bot. 5: 181-221, 301- 332, 455-500. BEDDOME, R. H. 1883. Handbook to the ferns of British India and the Malay Peninsula. Thacker, Spink & Co., Calcutta. — 1892. Ibid. Supplement. BERNHARDI, J. J. 1801. Tentamen alterum filices in genera redigendi, in Schrader, Journ. Bot. 1800, 2: 121-130. BLuME, C. L. 1828. Enumeratio Plantarum Javae. Van Leeuwen, Leiden. — 1829-1830. Flora Javae; Filices, t. 1-65. H. Remy, Bruxelles. Bower, F. O. 1928. The Ferns, vol. III. Cambridge University Press. CHING, R. C. 1936. A revision of the Chinese and Sikkim-Himalayan Dryopteris, I. Thelypteris. Bull. Fan Mem. Inst. Biol. Bot. 6: 237-350. —— 1940. On natural classification in the family ‘Polypodiaceae’. Sunyatsenia 5: 201-268. CurisT, H. 1897. Die Farnkréuter der Erde. G. Fischer, Jena. —— 1899. Monographie des Genus Elaphoglossum. Neue Denkschr. allg. Schweiz. Ges. Naturwiss. 36: 1-159. —— 1907. The Philippine species of Dryopteris. Philip. J. Sci. Bot. 2: 189-217. CHRISTENSEN, C. 1905-1906. Index Filicum. Hagerup, Copenhagen. 1911. On a natural classification of the species of Dryopteris, in Biologiske Arbejder, tilegnede Eug. Warming, ed. L. K. Rosenvinge. Hagerup, Copenhagen. — 1913. A monograph of the genus Dryopteris, Part 1. K. Dansk Vid. Selsk. Skr. VII, 10 (2): 55-282. — 1920. Ibid. Part 2. /.c. VIII, 6 (1): 1-132. —— 1932. The Pteridophyta of Madagascar. Dansk Bot. Ark. 7: 1-253, pl. 1-80. —— 1939. Filicinae. Chapter XX in Fr. Verdoorn, Manual of Pteridology. M. Nijhoff, The Hague. — 1943. A revision of the Pteridophyta of Samoa. Bernice P. Bishop Museum, Bulletin 177. Honolulu. — &R. E. Horttum. 1934. The ferns of Mt Kinabalu. Gard. Bull. Str. Settl. 7: 191-324, pl. 51-62. COPELAND, E. B. 1933. Trichomanes. Philip. J. Sci. 51: 119-280, t. 1-61. — 1937. Hymenophyllum. /.c. 64: 1-188, t. 1-189. — 1938. Genera Hymenophyllacearum. /.c. 67: 1-110, t. 1-11. —— 1947. Genera Filicum. Chronica Botanica, Waltham, Mass. — 1958-1960. Fern Flora of the Philippines. Institute of Science and Technology, Monograph 6, Manila. CoTTHEM, W. VAN. 1970. Comparative morphological study of the stomata in the Filicopsida. Bull. Jard. Bot. Nat. Belg. 40: 81-239. Fre, A. L. A. 1852. Mémoires sur la famille des Fougéres, V. Genera Filicum. Paris & Stras- bourg. (18) Dedication Ho.ttvuM, R. E. 1932. On Stenochlaena, Lomariopsis and Teratophyllum in the Malayan Region. Gard. Bull. Str. Settl. 5: 245-312. —— 1947. A revised classification of Leptosporangiate Ferns. Journ. Linn. Soc. Bot. 53: 123- 158. — 1951. The fern-genus Pleocnemia Pres]. Reinwardtia 1: 171-189. — 1955. A Revised Flora of Malaya, vol. 2, Ferns of Malaya. Government Printer, Singapore. — 1959. Flora Malesiana, Ser. II, vol. 1, pt 1, Introduction and Gleicheniaceae. — 1963. Ibid. pt 2, Cyatheaceae. — 1973. Copeland’s contribution to fern taxonomy. Philippine Agriculturist 57: 17-20. — 1974. The fern-genus Pleocnemia. Kew Bull. 29: 341-357. —— 1978. Flora Malesiana, Ser. II, vol. 1, pt 4, The Lomariopsis Group. Hooker, W. J. 1838-1842. Genera Filicum. G. Bohn, London. — 1844-1864. Species Filicum, vol. 1-4, W. Pamplin; vol. 5, Dulau. London. — & J. G. BAKER. 1865-1868. Synopsis Filicum. R. Hardwicke, London. —— & —— 1874. Ibid. ed. 2. — &R. K. GREVILLE. 1827-1831. Icones Filicum; vol. 1, 1827-28; vol. 2, 1829-31. Treuttel & Wiirtz, London. KLeEKowskKI Jr, E. J. 1979. The genetics and reproductive biology of ferns, in Dyer, A. F. (ed.), The experimental biology of ferns. Exp. Bot. 14: 133-170. Kunn, M. 1869. Filices. Ann. Mus. Bot. Lugd.-Bat. 4: 276-300. KunzZE, G. 1840-1851. Die Farrnkrauter in kolorirten Abbildungen (2 volumes). E. Fleischer, Leipzig. Lovis, J. D. 1977. Evolutionary patterns and processes in ferns. Adv. Bot. Res. 4: 229-415. LuGARDON, B. 1971. Contribution a la connaissance de la morphogénése et de la structure des parois sporales chez les filicinées isosporées. Thése, Toulouse, 257 pp., 51 tab. MANTON, I. 1950. Problems of cytology and evolution in the Pteridophyta. Cambridge University Press. METTENIUS, G. 1856. Filices horti botanici Lipsiensis. L. Voss, Leipzig. — 1860. Uber Seitenknospen bei Farnen (reprint at Kew; origin ?). — 1863A. Uber den Bau von Angiopteris. Abhandl. math. phys. Cl. K. Sachs. Gew. Wiss. 6: 501-570, t. i-x. —— 1863B. Filices, praesertim Indicae et Japonicae. Ann. Mus. Bot. Lugd.-Bat. 1: 46-58. —— 1864A. Ibid., para altera; /.c. 1: 222-241. — 1864B. Uber die Hymenophyllaceae. Abhandl. math. phys. Cl. K. Sachs. Gew. Wiss. 7: 403-504, t. i-v. — 1866. Filices; praesertim Indicae et Japonicae. Ann. Mus. Bot. Lugd.-Bat. 2: 219. MILpE, J. 1866. Das Genus Athyrium. Bot. Zeit. 24: 373-376. — 1870. Uber Athyrium, Asplenium und Verwandte. Bot. Zeit. 28: 329-337, 345-354, 370- S71. Moorg, T. 1857. Index Filicum, I. Synopsis of the genera of ferns. W. Pamplin, London. PICHI SERMOLLI, R. E. G. 1977. Tentamen Pteridophytorum genera in taxonomicum ordinem redigendi. Webbia 31: 313-512. PRESL, C. B. 1825. Pteridophyta, in Reliquiae Haenkeanae, fasc. 1: 14-84. Prague. — 1836. Tentamen Pteridographiae seu Genera Filicacearum. Prague. RIDLEY, H. N. 1926. The ferns of the Malay Peninsula. J. Mal. Br. R. As. Soc. 4: 1-121. SCHKUHR, C. 1804-1809. Vier und zwanzigste Klasse der Linnéischen Pflanzensystems oder Kryptogamische Gewichse, vol. 1-3. Wittenberg. (19) FLORA MALESIANA Scuott, H. 1834. Genera Filicum. Wallishauser, Vienna. SmiTH, JAMES E. 1793. Tentamen botanicum de Filicum generibus dorsiferarum. Mem. Acad. Turin 5: 401-422. SMITH, JoHN. 1841-1843. An arrangement and definition of the genera of ferns, in Hook. J. Bot. 4: 38-70, 147-198; in Hook. Lond. J. Bot. 1: 419-438, 659-668; 2: 378-394. — 1855. Filices, in Seemann, Botany of the voyage of H.M.S. Herald: 226-244. L. Reeve, London. —— 1866. Ferns, British & Foreign. Hardwicke, London. —— 1875. Historia Filicum. Macmillan, London. SWARTZ, O. 1801. Genera et Species Filicum ordine systematico redactarum, in Schrader, Journ. Bot. 1800, 2: 1-110. — 1806. Synopsis Filicum. Kiliae. WALKER, T. G. 1979. The cytogenetics of ferns, in Dyer, A. F. (ed.), The experimental biology of ferns. Exp. Bot. 14: 87-132. (20) ly ENTRODUC TORY ‘NOTE The work of preparation of a new survey of all the Pteridophytes of Malaysia will occupy a considerable period. It is proposed to publish this work in parts, as studies of particular families or genera are completed, but it is impossible to plan in advance the precise sequence of these studies. It is anticipated that the new information to be recorded, and new ideas based upon it, will throw a good deal of new light on the delimitation of genera, and upon the inter-relationships of genera, especially among the ferns, which are by far the largest of the major groups concerned. Therefore one cannot now predict what final scheme of classification will emerge. But it is necessary to have some sort of conspectus at the start, as a preliminary survey of the ground to be covered, and as a guide for those who wish to consult the parts of the work as they appear. I have therefore drawn up a list of the major groups, with the genera in each, and also a series of keys to the genera of ferns. The nomenclature of the major groups, the generic concepts, and the keys, must all be regarded as tentative. As regards the major groups, there are differences of opinion as to the status of each and as to the most appropriate names. I hope at least that the significance of the names here used is clear. As regards the ferns, there is still no general agreement as to the subdivision of the vast and heterogeneous assemblage formerly known as Polypodiaceae, though the more primitive families are clearly defined. In the conspectus which follows, I have arranged the genera of Polypodiaceae sens. lat. in Groups which seem to me to be natural, but I have given no formal names to the Groups, nor specified their status, except for Polypodiaceae sens. strict. and Grammitidaceae, which (with the exception of a few genera) are also accepted by COPELAND. The fern specimens in the Herbarium at Bogor are now arranged according to these Groups. Probably the Groups will be modified in any final re-assessment at the conclusion of their treatment in ‘Flora Malesiana’. So far as possible, genera within a family or Group will be treated together. There are two alternative General Keys, which lead to families or Groups of genera. In the first key a principal distinction is between epiphytes and terrestrial ferns; in the second key this distinction is not used. The delimitation of some genera of ferns is still not clear, and the generic names are to be taken as tentative. In general, they are those accepted in COPELAND’S Genera Filicum (1947), but there are some changes. Critical literature on fern taxonomy up to the year 1934 is very fully listed in CHRISTENSEN’S Index Filicum and its three Supplements. Some important books and papers subsequently published and dealing with Malaysian ferns are listed after the keys to genera. R. E. Horrrum 2. LIST OF MALAYSIAN, PTERIDOPHYTES PSILOTOPSIDA Colysis Loxoscaphe PSILOTALES Crypsinus Athyrium Group Dendroconche Anisocampium Psilotaceae Dendroglossa Athyrium Est orm Diblemma Callipteris Tmesipteridaceae Dipteris Cornopteris Tmesipteris Drymoglossum Cystopteris Drynaria Diplaziopsis LYCOPSIDA Goniophlebium piplaziers LYCOPODIALES Grammatopteridium Dryoathyrium : Holcosorus Lycopodiaceae Holostachyum Blechnum Group Lycopodium Lecanopteris Blechnum Selaginellaceae Lemmaphyllum Brainea Selaginella Lepisorus Doodia Isoetaceae Leptochilus Weed ee Isoetes Loxogramme Davallia Group Merinthosorus Araiostegia SPHENOPSIDA Microsorium Davallia EQUISETALES Myrmecophila Davallodes j Neocheiropteris Humata Equisetaceae : : : Oleandropsis Leucostegia Equisetum Paragramma Parasorus PTEROPSIDA Photinopteris Scyphularia 6 Platycerium Trogostolon pianist Polypodiopsis Dennstaedtia Group Ophioglossaceae Pycnoloma Dennstaedtia Botrychium Pyrrosia Histiopteris Helminthostachys Selliguea Hypolepis Ophioglossum Thayeria Microlepia MARATTIALES Thylacopteris Monachosorum Warattinceae Grammitidaceae Orthiopteris : ; Acrosorus Paesia Angiopteris a : . Calymmodon Pteridium Christensenia i Macroglossum Ctenopteris Dicksonia Group Marattia Grammitis — Cibotium Nematopteris Culcita FILICALES Oreogrammitis Cystodium Osmundaceae Prosaptia Dicksonia Leptopteris Scleroglossum Dryopteris Group Osmunda Xiphopteris Acrophorus Schizaeaceae REMAINING GENERA OF FERNS Currania Lygodium (in Groups, alphabetically) Diacalpe Schizaea Adiantum Group Didymochlaena Gleicheniaceae Adiantum Dryopteris _ Dicranopteris Anogramma Gymnocarpium Gleichenia Ceratopteris Peranema Hymenophyllaceae Cerosora Polystichum Cheilanthes Polystichopsis Hymenophyllum ; Stenolepia ‘ Coniogramme p Trichomanes : = ‘ Doryopteris Lindsaea Group Matoniaceae Hemionitis Isoloma Matonia Notholaena Lindsaea Phanerosorus Onychium Sphenomeris Cyatheaceae Pellaea Tapeinidium Cyathea Pityrogramma Lomariopsis Group Polypodiaceae (s.str.) Syngramma Arthrobotrya Aglaomorpha Schizolepton Bolbitis Arthromeris Taenitis Egenolfia Belvisia Asplenium Group Elaphoglossum Cheiropleuria Asplenium Lomagramma Christiopteris Diplora Lomariopsis II Morphology of ferns Teratophyllum Ctenitis Dictyocline Thysanosoria Cyclopeltis Sphaerostephanos Nephrolepis Group Dryopolystichum Stegnogramma Arthropteris Hemigramma Thelypteris Nephrolepis Heterogonium Vittaria Group Oleandra Hypodematium Antrophyum Plagiogyria Group SESH Monogramma Plagiogyria Luerssenia Vaginularia : Pleocnemia Vittaria Pteris Group Psomiocarpa Acrostichum Pteridrys HETEROSPOROUS FERNS Hemipteris Quercifilix Marsileaceae = aia Stenosemia Marsilea mEIS Tectaria +6; Schizostege oak Stenochlaena Thelypteris Group Salvinia Tectaria Group Ampelopteris Arcypteris Cyclosorus THE MORP HOVMOGY-OFeFERNS A fern plant consists of a stem, bearing leaves and roots. The leaves (or some of them) bear dehiscent sporangia, each sporangium containing unicellular spores, which are in most cases wind-dispersed. A spore germinates to produce a small green plant called a prothallus. The prothallus bears sexual organs (archegonia and antheridia). After fertilization by an antherozoid, the female cell in an archegonium grows to form a new fern plant. The life cycle of a fern thus has two phases, asexual (the fern plant) and sexual (the prothallus). These phases are also called the sporophyte and the gametophyte. The sporophyte is much longer-lived, larger and more diversified than the gametophyte, and its characters are mainly used in taxonomy. The following statement deals with the parts of the sporophyte in turn, with discussion of the kinds of modification of each which occur, and of special terminology. Finally, a note on the gametophyte will be given, including reference to the not infrequent condition in which the sexual process is omitted. Stem. (a) Shape, size, and habit of growth——A fern stem may be long and creeping or climbing, in which case it is usually called a rhizome, or it may be short and compact, in which case it is often called a stock, rootstock or caudex. If it grows erect, as in tree-ferns, with a tuft of leaves at its apex, it is called a trunk. A creeping stem or rhizome may be dorsiventral or radial in construction. In the former case the leaves (or their stalks) are borne on the upper surface (often in two alternate rows, some- times more than two), the roots entirely or mainly on the lower surface; the internal vascular structure corresponds to this external differentiation. In a radially constructed rhizome leaves and roots are borne on all sides (though of course the leaf-stalks all grow upwards), and the vascular system shows a corresponding radial symmetry. A short stem or rootstock, bearing crowded leaves, is usually radial in construction. Such a stem may be quite erect, but more often it is more or less decumbent; the presence of a per- sistently erect rootstock is sometimes an important diagnostic character. Tree-ferns, and others in which the erect rootstock grows to an appreciable height above the surface of the ground, have numerous roots which may form a close covering on the lower part of the stem, thus giving stability to the plant. (b) Branching.—In a few cases this is by simple dichotomy (the apical growing-point dividing into two equal parts). More frequently branches arise in association with the bases of leaf- stalks, usually on the outside. The method of branching may be important taxonomically; it has been too little studied. (c) Hairs, Bristles and Scales.—The young parts of a stem are in almost all cases protected either by scales or hairs; similar scales or hairs also cover the very young leaves before they uncurl. Such scales and hairs are more or less persistent on older parts of stem and leaves. HI FLORA MALESIANA [ser. II, vol F Hairs consist of a single cell or of a single row of cells, and in different ferns are of characteristic length and thickness; the thicker ones are sometimes quite rigid and bristle-like. True bristles, which are more than one cell thick at the base, but round in section, also occur in some cases. Scales are flat plates of cells, one cell thick; the details of their structure (especially characters of edges, base and apex, and the presence or absence of superficial outgrowths) are often very important taxonomically and may need microscopic examination. A peltate scale is one attached at some point on its surface, not on its edge. Where lateral cell-walls of a scale are thickened so as to form a distinct lattice-work pattern, the scale is called clathrate; the upper and lower walls may or may not be translucent. (d) Internal Structure.-—The most important internal structure is the vascular tissue. The vascular strands which pass into leaves and roots are connected to the vascular system of the stem; the latter is called a stele, or if at any cross section it consists of more than one part, the parts are called meristeles. The simplest kind of stele is a protostele; in this there is one single solid strand of xylem with phloem around it, the whole surrounded by an endodermis. In some cases the xylem of a protostele is not solid, but has a core of non-vascular tissue; such a structure is called a medullated protostele. In some long-creeping rhizomes (e.g. Micro- lepia) the stele is a hollow cylinder, with phloem and endodermis both inside and outside the xylem; this is called a solenostele or siphonostele. In this case there is usually a gap in the stele where the vascular supply to a leaf is attached; such a gap is called a /eaf-gap. Rhizomes of this type are usually dorsiventral. In Davallia, the solenostele does not consist of a continuous hollow cylinder, but the cylinder (not circular in section) is composed of a network, most of the gaps in which are not leaf-gaps. This may be called a dissected solenostele; in the case of Davallia it is dorsiventral, the ventral meristele being broader than the rest and bearing roots. In stems of radial structure bearing many leaves close together, the stele usually forms a hollow cylinder in which there are many leaf-gaps, these gaps being like the meshes in a network. Such a stele is called a dictyostele. In a few ferns there are additional steles internal to the principal stele (e.g. Matonia, some species of Preris); also in a few there is a cortical system of stem-bundles outside the principal stele (e.g. Stenochlaena). Besides vascular tissue, most fern- stems contain thick-walled tissues which serve to give mechanical strength; the presence (or absence) and distribution of such tissue may be important. Roots. Roots are all adventitious. The primary root of a fern-embryo, having no power of increasing in thickness, is soon inadequate to supply the needs of the growing plant, and further roots must grow from the stem (in some cases also from leaf-bases). Little use has been made of root characters for taxonomic purposes, but there is considerable diversity of root-structure among ferns as a whole. In some species of Hymenophyllaceae there appear to be no roots, the rhizome being covered with hairs which function as root-hairs. Leaves. VEGETATIVE CHARACTERS. Fern-leaves are usually called fronds; their stalks or petioles are called stipes; the blade or flat green part of a leaf is called the /amina. The lamina is usually divided into parts called /eaflets; the axes on which leaflets are borne are called rachises (rhachis is the more correct spelling, but rachis is in general use in English). Fronds may be sterile (lacking sporangia), or fertile (bearing sporangia); sterile and fertile fronds may be alike in form, or dissimilar (dimorphous). (a) Stipes —A stipe may be jointed or articulated at its junction with the stem, or in some cases it is articulated to an outgrowth from the stem called a phyllopodium (e.g. Elaphoglossum, Oleandra); but more commonly it is not articulated. Externally a stipe usually bears hairs or scales like those of the rhizome or caudex, but often smaller; sometimes the scales are borne at the ends of warts or thorns. In some cases where the rhizome bears only scales, the stipe (and rachis) may also bear hairs. The character of the hairs, whether unicellular or multicellular (with transverse septa), and whether terminating in a glandular cell, is always important. In a few ferns the stipe has slime-glands, producing mucilage when the fronds are very young, and elongate aerophores, the latter often in association with much-reduced leaflets. In any case, there is usually a pale and more or less raised line along each side of the stipe. This line is also nV’ Dec. 1959] 7 _ Morphology of ferns an aerophore; there are stomata on it, and internally the tissue is thin-walled, with air-spaces between the cells. The line is sometimes not continuous but broken, and sometimes doubled (Cyathea). Internally the number and arrangement of the vascular bundles in a stipe is important; often the number and arrangement change between the base and apex. (b) Branching of fronds.—Fronds may be simple (consisting of a continuous lamina, which may be entire or /obed in various ways), or they may consist of two to many leaflets which are borne by the rachis or its branches. The commonest arrangement is for the main rachis (continuing the line of the stipe) to be almost straight, bearing leaflets (stalked or not, articulate or not) along its sides; such a frond is said to be pinnate, and the leaflets are called pinnae. Like a simple frond, a pinna may be entire or lobed; if pinnately lobed, the leaflet is pinnatifid, if palmately lobed, palmatifid. Where the rachis bears branches, these are usually arranged in a pinnate manner; if each such primary branch bears leaflets, the frond is said to be bipinnate, and the leaflets are called pinnules. If each primary branch bears secondary branches pinnately arranged, and the secondary branches bear leaflets, the frond is said to be fripinnate, and the leaflets are of the third order; fronds may also be quadripinnate. Pinnae or rachis-branches are usually a/ternate on the two sides of a main rachis, but in some cases they are regularly almost exactly opposite (e.g. Gleichenia). The edge of a leaflet on the side towards the apex of the frond is called acroscopic, that towards the base of the frond is called basiscopic. The terms upper and lower are used for the two surfaces of the leaflets; the upper surface is that facing the stem-apex (also called the adaxial surface, the lower surface being abaxial). There are some cases in which the stipe is not continued into a single straight rachis, but forks into two equal branches at its apex; such a dichotomous branching is seen in fronds of young plants of Lygodium. The dichotomy may be repeated, in which case a fan-shaped frond results (Schizaea dichotoma, Dipteris lobbiana); or only the outer branch at each dichotomy may fork again, in which case pedate branching results. The ultimate branching of large fronds or of their veins is also often dichotomous, and one can trace a transition from dichotomous to pinnate branching in descending a frond to its base; the pinnate condition can thus be regarded as a development from the dichotomous one. Pseudo-dichotomy occurs in a few cases (notably in Gleicheniaceae and in Lygodium); in these a rachis bears a pair of opposite branches and then stops growing, its dormant apex remaining between the branches. (c) Shape of rachises and of junctions between them and with leaflets——In addition to the characters of superficial hairs and scales, and of internal vascular systems, rachises of ferns provide characters of another kind which are useful taxonomically. These characters relate to the way in which the structure of a smaller rachis is adjusted to that of a larger one where the two join, and ultimately to the relationship of a lateral leaflet to the rachis which bears it. Some rachises are rather deeply grooved on the upper surface, and where a smaller rachis joins a larger one, the groove of the latter is opened to admit that of the former. In some such cases (e.g. Dryopteris, Athyrium, Pteris) the midrib of a leaflet is also grooved, and a rachis- groove is open to admit the midrib-groove of a lateral leaflet; the edge of the leaflet is then more or less decurrent down the side of the rachis. In other cases the thickened basiscopic edge of a leaflet is decurrent on the edge of the rachis-groove (e.g. Lindsaea). Other rachises are not or only slightly grooved on the upper surface and there is no opening of the groove of a main rachis to admit that of a smaller one (e.g. Thelypteris, Ctenitis). In such cases the base of the lamina of a leaflet is decurrent on the side of the rachis, sometimes forming a small continuous ridge or wing. Characters such as this are constant throughout a genus, or even throughout a group of genera, and are often valuable diagnostically, especially in sterile fronds. I believe also that they may provide valuable evidence for those who are seeking a natural classification of ferns. (d) Venation.—Veins mark the course of the vascular bundles in a leaf; they are usually evident on the surface, as the surface tissue over a vein is different from that over the rest of the lamina. The pattern of venation is always important taxonomically. In fronds which are thick and fleshy or leathery the veins are sometimes quite invisible on the surface, or only the Vv FLORA MALESIANA [ser. II, vol. 11 larger ones may be prominent; the pattern of venation can then be seen by clearing the frond (boiling it and then treating it with a bleaching agent; or boiling alone may be sufficient, due to replacement of internal air by water). In large much-branched fronds the ultimate leaflets are usually small, with few veins in each, and these veins commonly end in lobes or teeth at the edge of the lamina. Such veins are free; a vein may fork once or more times, but the branches do not unite again at any point. Where leaflets are very small, the veins are often dichotomously branched, while larger leaflets usually have a midrib or costa with lateral veins which are simple or forked. In larger leaflets the vein-pattern may be more complex, still with free veins. The sides of a leaflet may be deeply lobed (pinnatifid) with pinnate vein-groups in each lobe; the main vein of a lobe is then called a costule (e.g. Thelypteris, Cyathea). In a large number of ferns with large leaflets, the veins join neighbouring veins after branching, thus forming areas of lamina surrounded by veins; such areas are called areo/es, and veins which unite with others are said to anastomose. The anastomosis may consist only in the formation of a single series of areoles along costae or costules (the rest of the veins ending freely) or it may form a more or less elaborate network occupying almost the whole lamina. In the latter case there are several possible patterns (e.g. Cyclosorus, Tectaria, Goniophlebium, Acrostichum). One useful character is the presence or absence of small veins which end freely inside the areoles. At the ends of some veins are water-excreting pores or hydathodes, often evident on the upper surface of the lamina as distinctive spots (round or elongate) which in some cases may ultimately become covered with white scales due to deposits of salts left after the water originally holding them has evaporated. Such hydathodes may provide useful diagnostic characters (e.g. Pyrrosia, Grammitis, Coniogramme). (e) Surface characters of lamina.—The lower surface of the lamina may be glaucous (covered with a pale blue-green waxy layer); this character often disappears if specimens are subject to much heat in drying. The lower surface in other cases may be more or less completely covered with a layer of white or yellow loose waxy powder; this is excreted from glandular hairs (e.g. Pityrogramma). Hairs on the lamina are always important, and often need to be examined with a microscope for the structure to be clearly evident. The nature and arrangement of stomata may in some cases be significant (e.g. Schizaea). The patterns of thickenings on walls of epidermal cells may be characteristic (or of the single cell-layer of filmy ferns); and in some ferns there are narrow spicular cells containing silica (e.g. Vittaria). In some ferns there are false veins, which are lines along which surface cells are more or less elongate and sometimes devoid of stomata, simulating the surface appearance of veins but with no underlying vascular tissue (e.g. Angiopteris, Trichomanes sect. Crepidomanes). (f) Polymorphism of fronds.—In many ferns, the fronds of young plants have a distinctive shape different from that of fronds of mature plants; such young plants may offer useful diagnostic characters. Some high-climbing ferns of the Lomariopsis Group have leaves of distinctive shape on those parts of the plant which are near the ground (always in moist shady evergreen forest) on rocks or tree-trunks; such leaves are called bathyphylls, and the leaves of high-climbing parts of the same plants are called acrophylls. FERTILE LEAVES. (a) Sori.—In most ferns sporangia are borne in distinct groups called sori. In some cases the sori spread along the veins to such an extent that they can hardly be called groups of sporangia, and this leads to the acrostichoid state (see below). Many sori are protected by indusia, which are thin outgrowths from the surface of a frond. The sori of other ferns are exindusiate, but in some of these the sori are protected by being produced in depressions or grooves, or by being covered when young by paraphyses (hairs of various forms, or scales, borne among the sporangia). The position and shape of sori and of their indusia (if present) are always important taxonomically. The older schemes of classification, and also that of Hooker and BAKER (Synopsis Filicum, 1868) were based entirely on these characters. But species closely related in every other respect may differ in presence or absence of an indusium, or in shape of sori (especially if these lack indusia) and a natural classification must take such facts VI Bec. 1959] te Morphology of ferns — into consideration. The Orders Ophioglossales and Marattiales are quite different from Filicales in the form and arrangement of their sporangia (see statement on sporangia below); further remarks here refer to Filicales. A sorus may be more or less circular, or elongate. If circular, it may be at the end of a free vein (at the edge of the lamina or not) or seated upon a free vein, or at the junction of veins in a network, or at the end of a free vein enclosed in an areole surrounded by other veins. In any of these cases it may be protected by an indusium, which may be pocket-shaped (at- tached at base and sides), or kidney-shaped, or circular and attached by the edge or the centre, or cup-shaped, or of other shapes, or it may have no indusium. If the sorus is at the end of a vein, it may be protected by the thin reflexed edge of the lamina, or by two more or less equal outgrowths from upper and lower surface (Dicksonia), the two outgrowths sometimes more or less joined to form a protective funnel or cup (Trichomanes, Dennstaedtia). Sori which are elongate may spread along free veins, or along veins which anastomose, or they may spread along the margin, joining the ends of veins which in a sterile leaflet would be free (fusion- sori or coenosori) or they may lie close to the costa of a leaflet (Blechnum). Marginal fusion- sori are sometimes protected by an inner indusium (Lindsaea), sometimes also by the reflexed margin (Preridium), but if the margin is reflexed, the inner indusium may be lacking (Pteris). It is especially exindusiate sori which spread along veins away from the margin, often to a different extent in closely related species. The above survey of soral form is not exhaustive, but is intended as an indication of the possibilities and as a guide in using the keys which follow. Details of individual soral forms will be given in the taxonomic treatment of the families and genera. (b) Dimorphism of leaves.—In many ferns the fertile fronds differ in shape from sterile ones. In such cases the fertile fronds often have a lamina of reduced size; this reduction may be slight or it may be so considerable that the sterile and fertile fronds are quite different in aspect. Fertile fronds also often have longer stipes than sterile, and in some cases this may be the chief difference between the two. (c) The Acrostichoid state.—Where sori spread along all the veins of a fertile leaflet and the leaflet is of reduced size as compared with a sterile one of the same species, the ripe sporangia may be so close together that they entirely cover the lower surface of the fertile leaflet (they may also grow from the surface of the lamina between the veins); this is called the acrostichoid condition, the name being taken from the Acrostichum, in which only the upper leaflets are fertile in this way. Formerly all acrostichoid ferns were included in the genus Acrostichum, but the acrostichoid state has certainly arisen along several different evolutionary lines, and a genus based on it alone is a very unnatural one. In some acrostichoid ferns there is an additional vascular system close to the lower surface of the lamina, in addition to the normal system found in sterile fronds of the same species. Sporangia. (a) Ophioglossales. —Here the sporangia are large, spherical, opening by slits, and are attached to spike-like or branched outgrowths from the base of the lamina; the fertile part of the frond is thus not fern-like in aspect. (b) Marattiales—In this Order the sporangia are also large, more or less laterally joined together in linear or circular groups on the surface of the lamina (along veins or at vein- junctions); they do not have an annulus comparable with that of members of the Order Filicales. (c) Filicales—In the great majority of this order the sporangia have a basically similar structural plan, in which dehiscence is caused by contraction on drying of a more or less com- plete ring of cells (the annulus) which have inner walls thickened but outer walls thin; there is also a particular place (stomium) where rupture occurs. The more primitive families (Osmunda- ceae, Schizaeaceae, Gleicheniaceae) have a less specialized development of the annulus, that of Osmundaceae being the least specialized (its annulus is not ring-shaped). In Hymenophyllaceae, Cyatheaceae, Plagiogyria and some other genera the annulus is complete and ob/ique in its position on the sporangium; in the great majority of genera the annulus is almost vertical and incom- plete, being broken by the stalk, but even in these cases the structure of the sporangium is not per- VII FLORA MALESIANA fectly symmetrical when divided along the plane of the annulus. In some members of the Adiantum Group the cells of the annulus are broad and more or less uneven. A detailed study of the development and structure of sporangia has been made in comparatively few fern-genera. Spores. Fern spores are aiways produced in groups of four (tetrads), each tetrad normally the result of the meiotic divisions of one spore-mother-cell. A spore may have either of two distinct shapes, monolete (or bilateral), and trilete (or tetrahedral). Monolete spores are more or less bean-shaped (like a Phaseolus seed), with an angle along the straight edge where the spore is in contact with the similarly angled edge of another spore of the tetrad; in each tetrad there are two such pairs. Trilete spores meet together on three faces, and at the angles between them, all four spores meeting at the centre of a tetrad. Usually all species in a genus have spores of the same shape, but there are certainly some genera in which both shapes of spores occur (e.g. Dicranopteris), and I have seen evidence that even within a single species there may be spores of both kinds. The inner layer of cells in the wall of a sporangium (called the tapetum) breaks down during the development of the tetrads of spores, its substance being absorbed by the spores during their development. In some cases part of the substance of the tapetum forms an external covering on each spore, known as a perispore. The perispore is usually more or less folded into rather irregular wing-like structures, or sometimes into more regular spines. Other genera of ferns lack a perispore, and then the wall of the spore itself may be variously sculptured into a more or less complex pattern of warts or ridges. The presence or absence of a perispore, its structure if present, or the wall-characters where there is no perispore, are always of taxonomic importance as well as the actual size of the spores. In the case of hybrids, where normal meiosis does not occur, there are often shrivelled empty spores, and the presence of such is always significant. In the case of apogamous ferns (see note on the gametophyte below) the spores are of at least two kinds, large functional ones and smaller ones which are not functional (for a detailed statement, see MANTON, Problems of Cytology and Evolution in the Pteridophyta, Cambridge, 1950). Gametophyte. In Ophioglossales the gametophytes are subterranean and saprophytic, and obtain their nutriment through the activity of an endophytic fungus. In all other ferns the gametophytes are green, and in the vast majority of cases they are more or less heart-shaped, with a growing-point in the sinus between the two lobes; they are thickened in a median area which bears the rhizoids, antheridia and archegonia on the lower surface. Distinctive characters are provided by shape of the whole prothallus (in some cases this is asymmetric or elongate), presence of superficial hairs of different kinds, colour and septation of rhizoids, and especially in details of structure of archegonia and antheridia. In a few cases the gametophyte is more or less filamentous (Schizaea, Hymenophyllaceae), or irregularly lobed (Vittaria and allied genera). In apogamous ferns, no sexual process occurs. The prothallus is developed from a diploid spore; it bears antheridia but no archegonia, and gives rise to a new sporophyte by vegetative budding. The diploid antherozoids of such a prothallus may fertilize haploid archegonia of a sexual prothallus of an allied species, the result being a triploid hybrid sporophyte. Such a hybrid is normally sterile, but may develop vegetative means of propagation not involving the formation of a prothallus; and some such hybrids have become apogamous. Higher polyploid plants are also not uncommon among ferns, and in such cases cytological evidence is of great taxonomic value. Heterosporous Ferns. The two families Salviniaceae and Marsileaceae are very different from other ferns in many respects. They have spores of two kinds, large and small, in separate sporangia. The small spores produce very small gametophytes which are male, and the large spores produce larger female gametophytes. Both kinds of sporangia are formed inside closed structures called sporocarps, borne by the leaves. These ferns are all aquatic, and they are sometimes collectively called Hydropterideae or Hydropteridales, but the two families are not closely related, and probably had quite different evolutionary histories. VIII 4. GENERAL KEY No 1 to PTEROPSIDA 1. Aquatic plants. 2. Plants floating. Leaves small, simple or bilobed sev, Swryrelaee Salviniaceae 2. Plants rooted in earth or on rocks. Leaves larger, more divided. 3. Leaves 4-partite. Sporocarps attached to stipes. ee : Marsileaceae 3. Leaves not 4-partite. Sporangia singly or in sori on lower surface ‘of lamina. 4. Sporangia borne singly, protected by reflexed edges of narrow lamina . Adiantum Group 4. Sporangia grouped in sori, on lower surface of lamina, not protected by reflexed edges. 5. Fern of stream-beds in deep shade. Fronds pinnatifid, sori without indusia Polypodiaceae 5. Fern of open swamps. Fronds bipinnatifid, sori indusiate. . . . . Thelypteris 1. Land plants or epiphytes. 6. Epiphytes. Group 7. Fronds simple, not over 2 mm wide, with a single vein, or with a few simple lateral soriferous veins close to the main vein . . FEM MORK Vittaria 7. Fronds branched, or if simple with a more * complex venation. Group 8. Lamina one cell thick apart from midribs of segments. . . . . . Hymenophyllaceae 8. Lamina throughout more than one cell thick. 9. Sporangia embedded in slender cylindrical appendages attached to surface of frond. Ophioglossaceae 9. Sporangia not so arranged. 10. Sori not indusiate (sometimes otherwise protected). 11. Sporangia not acrostichoid. 12. Sori superficial (not in pockets or grooves). 13. Fronds simple, pinnatifid or pinnate; if pinnate, pinnae not articulate to rachis. 14. Veins much anastomosing . . : Le, Sire Son. tee APolypodiaceac 14. Veins not or slightly anastomosings 15. Frond and stipe + hairy; spores trilete at Seat lct ae Grammitidaceae 15. Frond and stipe not hairy; spores monolete . . . . . . . Polypodiaceae 13. Fronds pinnate, pinnae articulate to rachis. (6MRinnacventires 1) Mone 2). 32) Pe, ease a es ee BPolypodiaceae 16. Pinnae lobed . . : Nephrolepis Group 12. Sori in pockets or grooves ‘(which are sometimes marginal). 17. Sori in pockets or depressions, + circular. 18. Veins anastomosing; or, if free, fronds not hairy . . . . . . Polypodiaceae LSsaVeins: tree, tronds more or less hairy =) 5: -senoneeeee Grammitidaceae 17. Sori elongate, in grooves. 19. Grooves all evenly oblique to costa . . . . . . Polypodiaceae 19. Grooves marginal or parallel to margin, or uneven in direction, sometimes anastomosing. 20. Scales entirely opaque, usually brown . . : Grammitidaceae 20. Scales nearly black, strongly clathrate, lumina of cells translucent Vittaria Group 11. Sporangia acrostichoid, covering entirely part or whole of a frond. 21. Veins much anastomosing; spores without perispore eos ae . . Polypodiaceae 21. Veins free or slightly anastomosing near edge; perispore present . Lomariopsis Group 10. Sori indusiate. 22. Sori elongate along veins . . . . . . . . +. +. +. +. ~~. ~~ Asplenium Group 22. Sori otherwise. 23. Sori elongate along edge of lamina. 24. Pinnae articulate to rachis . . SE a ERT eae Nephrolepis Group 24. Pinnae (if any) not articulate to rachis. 25. Rhizome protostelic . . - ATES te GRR ae Lindsaea Group 25. Rhizome with more complex vascular system , Oe BUTE Davallia Group 23. Sori otherwise. 26. Fronds articulate to rhizome. 27. Sori at ends of veins, near edge of lamina. P8ePinnae! (if present) not joimted stomrachis.. 5) S80) -) eee Davallia Group 28. Pinnae jointed to rachis . . ys ORAS.» HOSE) PE Nephrolepis Group 27. Sori close to costa of the simple frond 22> .. he elvia omiie ..aNephrolepis*Group 26. Fronds not articulate to rhizome . . . Nephrolepis Group 6. Terrestrial plants, or climbers starting from the exons or rock- plants. 29. High-climbing rhizome starting from the ground. 30. Rhizome not dorsiventral; veins anastomosing in a narrow series of costal areoles (seen at apex of pinna) Ae leo Peet od ce) LRAT OS Shy ee Pteris 30. Rhizome dorsiventral; veins either free or much anastomosing . . Lomariopsis 29. Terrestrial or rock plants. Group Group Ix FLORA MALESIANA [ser: II, vole 31. Caudex massive, erect; stipes succulent, with stipule-like outgrowths at their bases; bases of pinnae swollen Pe eat eras ee ge Loo oe ee Marattiaceae 31. Not these characters. 32. Lamina one or two cells thick apart from midribs of segments of lamina; no stomata. 33. Sporangia attached to elongating slender receptacles in funnel-shaped pockets at ends of veins. Hymenophyllaceae 33. Sporangia attached to surface of veins . Ole Vheile. Bees rs eee Osmundaceae 32. Lamina throughout more than two cells thick; stomata present. 34. Caudex or rhizome at apex, and bases of stipes, hairy or bristly (no flat scales) or apparently naked. 35. Rootstock massive, erect (in a few cases tree-like) or more or less decumbent, radially or- ganized, its apex above ground, bearing a close group of fronds. 36. Fronds simply pinnate; apex of caudex not densely hairy. 37. Aerophores at bases of stipes (sometimes also at bases of ae Plagiogyria Group 37. Aerophores lacking : fey Osmundaceae 36. Fronds more amply divided; apex of caudex densely ‘hairy : Dicksonia Group 35. Rootstock otherwise, usually entirely below ground. 38. Fertile part of frond not leaf-like, erect and attached to base of leafy part Ophioglossaceae 38. Fertile part of frond leaf-like, sometimes reduced in size as compared with sterile. 39. Fronds palmately divided; leaflets 3 or 5; veins anastomosing . . Marattiaceae 39. Fronds otherwise. 40. Veins much anastomosing, with free veins in the areoles. . . . Polypodiaceae 40. Veins in most cases free; where anastomosing, no free veins in areoles. 41. Sori quite superficial, on lower surface of lamina, or in a marginal groove. 42. Sori indusiate; fronds fan-shaped or slender and trailing . . . Matoniaceae 42. Sori not indusiate. 43. Fronds repeatedly pseudo-dichotomous, with a dormant apex between each pair of branches = «0 =. 3) Se Boe ie. Cy Soe Boe © Gleicheniaceae tae Fronds otherwise. . sa Sale: Law 2 Adiantum Group Sori at ends of veins or on 1 special appendages. Sporangia on special appendages which are at ends of veins of leaflets or attached near apex of frond or of its branches. . . Sol ter Schizaeaceae 44. Sporangia in sori at ends of single veins or uniting ends of several veins, not on special appendages . . : Dennstaedtia Group 34. Caudex or rhizome at apex, and ‘bases of stipes (at least when young) scaly. 45. Sporangia acrostichoid. 46. Rhizome dorsiventral, creeping on rocks. 47. Veins free, or if anastomosing the free veins almost all pointing outwards; spores with perispore - : Lomariopsis Group 47. Veins much anastomosing ‘with free veins in areoles pointing all ways; no perispore. Polypodiaceae 46. Rhizome not dorsiventral, often massive, bearing a tuft of fronds at its apex. 48. Only the upper pinnae fertile; veins much anastomosing, no free veins in areoles. Pteris Group 48. Whole frond fertile; in sterile frond veins free or anastomosing otherwise. 49. Veins free in sterile fronds, or a single row of costal areoles present. SOs Bertiestrond simplyapimnate. ) =e eee cc) ee Blechnum Group 50. Fertile frond bipinnate . . Ty tate Sink She Tectaria Group 49. Veins much anastomosing in sterile fronds 2 its Fa ae Tectaria Group - Sporangia not acrostichoid. . Sorus along edge of lamina, continuous or nearly so. s Edge of lamina reflexed, protecting sori. 53. Rachis grooved on upper surface, groove open to admit groove of midrib of pinna. Pteris Group 53. Rachis not so grooved (if grooved, edge of lamina may be decurrent on edge of groove). Adiantum Group 52. Edge of lamina not reflexed; sorus protected by indusium attached below it, opening fowaLrdsvedeesotslaminay 2, %) -irayl eens tee Meera eee Lindsaea Group 51. Sorus otherwise. 54. Sorus elongate, continuous along each side of costa of pinna ; Blechnum Group 54. Sorus otherwise. 55. Sporangia on surface of reflexed marginal lobes . . . . . Adiantum Group 55. Sporangia not on such lobes. 56. Sorus along veins (at least some of them). 57. Sorus indusiate. Pec. 1959] General key No. 1 to Pteropsida 58. Sorus symmetrically divided by line of vein. 59. Rachis grooved, groove open to admit groove of branch; scales lacking superficial Haws, = Dryopteris Group 59. Rachis somewhat srooved, groove not open to admit groove of pinna; scales with superficial hairs . . CW Ridtysbor e% Thelypteris Group 58. Sorus asymmetric, or on one side of 1 vein. 60. Sori along outer veins of costular or costal areoles . . . Blechnum Group 60. Sori otherwise. 61. Two strands in stipe, uniting upwards to form a single X-shaped strand. Asplenium Group 61. Two strands in stipe, uniting upwards to form a single U-shaped strand. Athyrium Group 57. Sorus not indusiate. 62. Sori spreading along all veins of lower surface. 63. Slender unicellular hairs present on frond and on scales . Thelypteris Group 63:3 Slender: unicellular shairsvlacking eens 9 -e Ae e Adiantum Group 62. Sori not spreading along all veins. 64. Several vascular bundles in stipe. . . a be Tectaria Group 64. Two vascular bundles at base of stipe, uniting upwards. 65. Waxy powder on lower surface of lamina x (rime e Seer. Adiantum Group G5s. No waxyepowder present... 225 ees ese ee) eee one Athyrium Group 56. Sori not along veins. 66. Sori at ends of veins, at or close to edge of lamina, each in the base of a cup, or protect- ed by an indusium attached below it or by the refiexed edge of the lamina. 67. Sori each in the base of a cup. ey Dennstaedtia Group 67. Sori protected by indusia or by edge of lamina. 68. Sori protected by indusia opening outwards. 692, Pinnae,articulatestoprachis) gre), fs sb act see 2 Nephrolepis Group 69. Pinnae not articulate . . vy Tie gs Lindsaea Group 68. Sori protected by reflexed edge of lamina. 70. An inner indusium also present; rachis sess groove open to admit groove of midrib of pinna. . : Pteris Group 70. An inner indusium lacking: rachis not grooved, or if grooved, edge of pinna decur- rent on edge of groove . . a: Adiantum Group 66. Sori not at ends of veins, or if so not ‘close to edge of lamina. 71. Rhizome dorsiventral. 72. Sori without indusium. 73. Fronds lacking dormant apices . . . emi. ©; Polypodium Group 73. Fronds always having some dormant apices oil Sieeet @ scan Gleichemacese 72. Sori indusiate. 74. Fronds simple . . i de doocwl. sheet. er eee NephrolepisiGronp 74. Fronds pinnately branched wit). qailesl. see Se eee Davallia Group 71. Rhizome not dorsiventral. 75. Fronds simple and jointed at base, or pinnate with pinnae jointed to rachis. Nephrolepis Group 75. Fronds otherwise. 76. Tree-ferns; sporangia with complete oblique annulus; many vascular bundles in Stipe: 2: i Cyatheaceae 76. Not tree- ferns; annulus vertical, ‘interrupted; vascular bundles in a simple ring (except Pleocnemia). 77. Rachis grooved, groove open to admit groove of branch-rachis or pinna. 78. Several vascular bundles in stipe. . . . . Dryopteris Group 78. Two bundles, joining to form one of U-shape . give: Athyrium Group 77. Rachis not grooved, or if grooved groove not open to admit groove of branch. 79. Hairs (if present) multicellular; scales lacking superficial hairs or glands. 80. Several vascular bundles in stipe . . . .-. . . .- Tectaria Group 80. Two bundles, joining to U-shape . . ; Athyrium Group 79. Hairs unicellular; scales bearing superficial hairs or glands Thelypteris Group 5. GENERALSREY Nor24oePRTEROPSI DA 1. Aquatic plants. 2. Plants floating; leaves small, simple or bilobed. . . 3) ep RE, Salviniaceae 2. Plants rooted in earth or on rocks; leaves larger, more divided. 3. Leaves 4-partite; sporocarps attached to stipes cae : Marsileaceae 3. Leaves not 4-partite; sporangia singly or in sori on lower surface of lamina. 4. Sporangia borne singly, protected by reflexed edges of narrow lamina . Adiantum Group 4. Sporangia grouped in sori, on lower surface of lamina, not protected by reflexed edges. 5. Fern of stream-beds in deep shade; fronds pinnatifid, sori without indusia Polypodiaceae 5. Fern of open swamps; fronds bipinnatifid, sori indusiate. . . . Thelypteris Group 1. Land plants or epiphytes. 6. Stipe containing 3 or more vascular bundles throughout. 7. Rhizome in most cases dorsiventral, stipes jointed to it or to phyllopodia borne by rhizome. 8. Sori of various form (in some cases acrostichoid), not indusiate. 9. Fronds pinnate; pinnae articulate to rachis, veins free wile Bn ES Nephrolepis Group 9. Fronds rarely pinnate; pinnae (if present) not articulate to rachis, or if so veins reticulate. 10. Acrostichoid; spores with perispore; veins free or uniting near margin Lomariopsis Group 10. Acrostichoid or not; no perispore; where acrostichoid, veins much anastomosing. Polypodiaceae 8. Sori indusiate. 11. Fronds simple. 12. Sori near midrib of frond . . . Wale MEAL. Nephrolepis Group pe Sori at ends of veins, solitary or joining many VEINS: MAL SOs We Davallia Group Fronds more or less branched. me Rimnaemointedatoprachisit |eciaaha stl coe Ghee cunt) eee ne Nephrolepis Group 13. Pinnae not jointed to rachis. . ileieg -£ Davaliia Group 7. Rhizome not dorsiventral, or if so the stipes not jointed to. it. 14. Stipes containing numerous bundles in a complex pattern, not a simple ring, in cross section. 15. Rootstock massive, short or tall; if creeping, subterranean. 16. Mangrove plants; distal pinnae acrostichoid, lower ones sterile . . . Pteris Group 16. Not mangrove plants; not acrostichoid. 17. Stock and stipes Se woody tissue; no stipules at bases of stipes. Hiss WSIS JOS 5 aoe iy eC RAMS €thch (0eet a Og Es ag SORA eB A Cyatheaceae 18. Veins anastomosing . . . . Tectaria Group 17. Stock and stipes without woody tissue; stipules ‘present at bases of stipes. Marattiaceae 15. Rootstock slender, long-creeping or high-climbing ATA died Be Pa: Pteris Group 14. Stipes containing a simple open ring of bundles. 19. Sporangia on a separate branch attached at base of lamina, or to surface of lamina; no annulus. Ophioglossaceae 19. Sporangia not on a separate branch of frond; annulus present. 20. Fronds pinnate or bipinnate, leaflets jointed to rachis. 21. Fronds in close tufts, not spaced on a climbing rhizome; sori separate and indusiate. Nephrolepis Group 21. Fronds spaced on a climbing rhizome; sori not indusiate, in most cases acrostichoid. Lomariopsis Group 20. Leaflets (if present) not jointed to rachis. 22; Fronds simple, sori oblique, exindusiate . . . . . . . . =. . BPolypodiaceae 22. Fronds otherwise. 23. Sori elongate, either close to midrib or to edge. 24 eS OnMCLOSERtOMMIdnI Digan). rss ree ay Oe, Devi) ea Blechnum Group PASSO closesto) edges.) =) - b pe bis Seley Ae Te Nephrolepis Group 23. Sori not elongate near midhibe or ‘edge. 25. Sporangia acrostichoid, edge of lamina in some cases reflexed. 26. Veins of sterile leaflets free. Oi eEronds“bipinnate, ¢ sa. 7 2Gthetn anes tach Ieee Tectaria Group 27. Fronds simply pinnate. 28. Rhizome creeping, dorsiventral A slg ool poo ae Lomariopsis Group At, Uinecomte One, walellhy gyacinyel 2995 6 4 5 6 6 4 & Blechnum Group 26. Veins of sterile leaflets anastomosing. POmiNhizomercteepinge. Gorsiventraly 1) ee ba) ae Lomariopsis Group PO ARNIZOMeZerech, enotaGorsiyentralie. se eebuey 2) se aa Tectaria Group 25. Sporangia not acrostichoid. 30. Rachis grooved, groove open to admit branches or midribs of leaflets. Dryopteris Group XII General key No. 2 to Pteropsida 30. Rachis not grooved, its wings (if any) confluent with edges of lamina. Tectaria Group 6. Stipe containing one vascular bundle, or two at the base joining upwards to form a single strand of various shape. 31. Rhizome-apex and bases of stipes hairy or apparently glabrous, not scaly. 32. Fronds repeatedly pseudo-dichotomous, a dormant apex between pairs of branches. Gleicheniaceae 32. Fronds otherwise branched. 33. Apices of some or all primary rachis-branches dormant. 34. Rachis climbing and twining; sporangia solitary : 2 tS £ Schizaeaceae 34. Rachis trailing, not twining; several sporangia in a sorus, indusiate 3 0R Matoniaceae 33. Apices of primary rachis-branches not dormant (apex of main rachis sometimes periodically dormant). 35. Sori at ends of veins, either in cups or protected by two equal or subequal flaps. Roary serus;slamina one Cell thick sect ls ss eee Hymenophyllaceae 36. Not filmy, lamina thicker. 37. Massive rootstock, prostrate or erect and trunk-like, densely covered with long hairs. Dicksonia Group Bversiender ‘creepins rhizome; hairs, short) 794. #2 Ss) 2 SoS Dennstaedtia Group 35. Sori otherwise. 38. Bases of stipes bearing prominent aerophores; fronds dimorphous, fertile with narrow acrostichoid pinnae . . = tee... <, .Plagiogyria 38. Bases of stipes lacking acrophores; fertile fronds otherwise: 39. Sporangia large, round, lacking an annulus, on deeply dissected pinnae Osmundaceae 39. Sporangia otherwise. 40. Sporangia variously arranged on lower surface of fronds, usually along the veins, or in a submarginal groove, not indusiate. 41. Sori elongate . . . id re ns eee oe Adiantum Group 41. Sori small and round, or - acrostichoid. cqeg tk Reece oe ee ee Se OLY POCIAceae 40. Sporangia arranged otherwise. 42. Annulus almost apical; fronds simple or branched dichotomously; sporangia on special appendages ts WON Ae CA NS eo eee Schizaeaceae 42. Annulus and frond otherwise. 43. Fronds pedately branched; sori superficial, indusiate. . . . . Matoniaceae 43. Fronds pinnately branched; sori at ends of veins, indusiate or ‘not. Dennstaedtia Group 31. Rhizome-apex and bases of stipes scaly. 44. Sporangia on surface of reflexed marginal lobes . . . ... . Adiantum Group 44. Sporangia not on such lobes. 45. Sori elongate along the veins of the lower surface. 46. Sori indusiate. 47. Scales clathrate; the 2 vascular bundles at base of stipe uniting upwards to an X-shape. Asplenium Group 47. Scales not clathrate; the 2 vascular bundles at base of stipe uniting upwards to a U-shape. 48. Unicellular slender hairs lacking . . . Ba ae Oh ee ee Athyrium Group 48. Unicellular slender hairs on frond and on scales A ee ee Thelypteris Group 46. Sori not indusiate. 49. Fronds simple; sori in grooves (or rarely superficial); epiphytes or rock-plants. Vittaria Group 49. Fronds usually not simple; sori not in grooves; not epiphytes or rock-plants. 50. Sori not occupying whole length of veins. 51. Lower surface of lamina not covered with powder . . . . . Athyrium Group 51. Lower surface of lamina covered with powder . . .. . . Adiantum Group 50. Sori occupying whole length of veins. 52. Slender unicellular hairs lacking on lamina and scales . . . . Adiantum Group 52. Slender unicellular hairs present on lamina and scales . . . . Thelypteris Group 45. Sori not elongate along the veins of the lower surface (in some cases elongate along a sub- marginal vein). 53. Sori in marginal or submarginal grooves, or superficial and forming continuous lines parallel to midrib. SA: aScalesuclathrate: oS p56 se ees see wc. eae eS) hee | te i ariagGroap 545,Scales: notuclathrate. 4. tim At. fee Pace ue lee, ta ee PG Lanne 53. Sori otherwise. 55. Sori at ends of veins (uniting several veins or not) protected by reflexed edge of lamina. 56. Rachis grooved on upper surface, groove opening to admit groove of midrib of pinna. Pteris Group XI FLORA MALESIANA XIV 56. Rachis not grooved in this way . . . oo. . . . . . . Adiantum Group 55. Sori not at ends of veins or not so protected. 57. Lower surface of frond more or less covered with white or yellow powder. Adiantum Group 57. Lower surface of frond not so covered. 58. Sori submarginal, at ends of veins, in small projecting cups, or protected by an indusium attached on the side remote from the margin. 59. Sori in’ cups: projecting sirom) margin) 9 ee Dennstaedtia Group 59. Sori not in such cups. 60. Sori always solitary at ends of single veins; scales of rhizome bearing papillae on marginal cells; rhizome not protostelic. . . . . Davallia Group 60. Sori often uniting ends of several veins; ‘scales lacking papillae; rhizome protostelic. Lindsaea Group 58. Sori otherwise. 61. Main rachis of fronds periodic in growth; thicket-forming ferns . Gleicheniaceae 61. Main rachis of fronds continuous in growth; not thicket-forming ferns. 62. Small epiphytes (occasionally on rocks); sori superficial or immersed in pockets in substance of the lamina, not indusiate . . . . ... . Grammitidaceae 62. Not epiphytes; sori in most cases indusiate. 63. Hairs on fronds unicellular (in rare cases very long hairs may consist of more than one cell). 64. Scales confined to swollen basal part of stipe; no superficial hairs or glands on scales. Tectaria Group 64. Scales not so confined, bearing superficial unicellular hairs or glands. Thelypteris Group 63. Hairs on fronds multicellular, cells short . . . . .. . Athyrium Group ace YS TO THE*GENERA)» OF PTEROPSIDA OPHIOGLOSSACEAE 1. Sporangia in two rows, embedded in an almost terete spike . . . . . . Ophioglossum 1. Sporangia on branches of the fertile segment of a frond. 2. Fertile segment of frond compact, with many short branches; sterile segment tripartite, each part with few leaflets . . . . Helminthostachys 2. Fertile segment of frond amply branched with spreading branches: sterile segment (in Malaysian species) pinnately branched with many small divisions Suchet? twice sine sboayciium MARATTIACEAE 1. Sporangia in each group along the veins near margins of leaflets; veins free. PEOraneia ih ‘cach eroup) united laterallysymereeres Smee) Se Rll ieee Marattia 2. Sporangia in each group almost free. 3. Fronds bipinnate; sporangia in each group commonly 8-12, less often to 20 . Angiopteris 3. Fronds simply pinnate; sporangia in each group much more numerous . . Macroglossum 1. Sporangia in each group united laterally to form a small circle, the circular groups scattered over the Rieiacesnveins; anastomosing’ . 4. 5. pe ey ed ers Rea ee Christensenia OSMUNDACEAE 1. Fertile pinnae quite different from sterile, lacking a green lamina; lamina‘of sterile pinnae not trans- lucent BIG og . . Osmunda 1. Fertile pinnae not different from sterile in “shape; lamina very thin and ‘translucent Leptopteris SCHIZAEACEAE 1. Fronds of adult plant dichotomously branched or simple, the fertile lobes at the end of a frond or of its branches. . . so Sette 1. Fronds of adult plants scandent (rachis twining) with very short primary “rachis- branches bearing featvasccondary, (branches 5 oe ace oe he sae! gee Be ee a re Lygodium GLEICHENIACEAE 1. Rhizome and resting apices of fronds bearing multicellular hairs which are branched near the base; veins always at least twice forked . . . ; Dicranopteris 1. Rhizome and resting apices of fronds bearing flat scales, rest of fronds usually also stellate hairs (one cell to each ray); veins simple or forked once ®° oie beetle alee Gleichenia HYMENOPHYLLACEAE* 1. Lips of indusium always well developed, broader and longer than the hollow basal part; receptacle usually much shorter than the lips of the indusium (an elongate receptacle in Meringium PRESL). Hymenophyllum 1. Indusium tubular or trumpet-shaped, sometimes with two-lipped mouth; receptacle usually elongating considerably vand» protruding iromvold soni ere. - eeeecne ene ee Trichemanes MATONIACEAE 1. Fronds erect, branching pedate-dichotomous . . . . Matonia 1. Fronds drooping, elongate, pinnately branched with apices “of some branches dormant. Phanerosorus CYATHAEACEAE Note. Probably all species will be united in the genus Cyathea (see HoLTtuM, Kew Bulletin 1957, pp. 41-45). POLYPODIACEAE 1. Rhizome bristly or hairy, stipes not jointed to it; terrestrial. 2. Fronds dimorphous; fertile fronds narrow, acrostichoid . . .... . Cheiropleuria 2. Fronds uniform; sori small and round . a sprtiveast i, dead, 2a) cae Dipteris 1. Rhizome scaly, stipes usually jointed to it; mostly epiphytes. 3. Young sori protected by umbrella-shaped paraphyses. 4. Sporangia acrostichoid on a constricted apical part of the frond . . . . . Belvisia 4. Sporangia in separate sori, or whole fertile frond narrow and acrostichoid. 5. Sori somewhat elongate, close to and parallel with edge of lamina . . . Paragramma * For a much fuller subdivision of this family, see COPELAND, Genera Filicum (1947) pp. 31-44. XV FLORA MALESIANA [ser. Il, vol} i 5. Sori otherwise. 6. Fronds small, fertile ones (or fertile parts) contracted. . . . . Lemmaphyllum 65 Fronds larger; not dimorphousis. 26 cae oo) ee eee _ Lepisorus and Neocheiropteris 3. Young sori not so protected. 7. Fronds bearing stellate hairs. 8. Fronds simple, entire. 9. Sori continuous along margin of narrow fertile frond or between margin and midrib. Drymoglossum 9. Sori separate, round, variously distributed . . . Pyrrosia 8. Fronds branched dichotomously, sporangia acrostichoid. on - part of surface ‘of fertile frond. Platycerium 7. Fronds not bearing stellate hairs. . ier at the ends of free veins (veins conspicuous). . All veins free ae 3S eye! 6 Ge le ano AOA heel > i ee Thylacopteris . Some veins anastomosing badltis . . . . . Goniophlebium 1 ‘Sori nat at ends of free veins (veins often inconspicuous). 12. Veins anastomosing to form only one series of areoles along costae . . Polypodiopsis 12. Veins more copiously anastomosing. 13. Rhizome swollen and ant-inhabited. 14. Rhizome not scaly; sori on distinct small lobes Sot Wo ee ee Lecanopteris 14. Rhizome copiously scaly; sori not on distinct lobes . . . . . . + Myrmecophila 13. Rhizome not ant-inhabited. 15. Fertile fronds (or fertile parts) acrostichoid, usually much reduced as compared with sterile. 16. Fronds pinnate or pinnatifid, lower pinnae or lobes always sterile. I75 Bronds pinnate; sterile pmnae jointed toyrachis 4 =< = = =. © « Photinopteris 17. Fronds pinnatifid . . eT ro cet en eee ee igniniiacaaTe 16. Fertile fronds wholly fertile. i Teron Sinonyube (inl 5 75 6 6 = 6 « oa 6 6 o oa -<% Christiopteris 18. Fronds simple. 19. Fronds thin, all veins visible. 20. Fronds c. 10 cm long; terrestrial ‘ Sg eke. ee Dendroglossa 20. Fronds c. 30 cm long; rock- plants or epiphytes » « « « » « » SMeptochilus 19. Fronds coriaceous, at most main veins visible. oh Sterile fronds narrow. NORIO?) Fee FF Oleandropsis Sterile fronds circular to broadly lanceolate. o Sterile fronds circular ; SEER SSW. (Eire . Pycnoloma 22. Sterile fronds longer than wide, apex acute eae Te te Grammatopteridium 15. Fertile fronds (or fertile parts) not acrostichoid. 23. Fronds pinnate or pinnatifid, pinnae or segments jointed to rachis. 24. Separate short humus-collecting fronds present . . . . . . . +. . Drynaria 24. Separate humus-collecting fronds lacking. 25. Bases of fronds broad, humus-collecting. 26; Eronds) on ‘special! branches of rhizomes. (2) 2:02) 93 41-05-55 Belhaven 26:) Kronds; not ionvspecial branches) = 95 9.) 4 = 4.5 4 © one Aglaomorpha 25. Bases of fronds not humus-collecting. 27. Fronds dimorphous En ey, Oe We he olga claniate | AR in oe Da Sule iS Holostachyum 27. Fronds not dimorphous . . Shoe Clee Arthromeris 23. Fronds simple or pinnatifid, segments not jointed ‘to rachis. 25. separate humus-collecting fronds present... = 2 9. 4 «4 ee Dendroconche 28. Separate humus-collecting fronds lacking. 29. Sori elongate, oblique, parallel to main lateral veins. 30. Fronds thin, all veins visible . . . 7 SS) eee Colysis 30. Fronds fleshy or leathery, at most main veins visible. 31. Main lateral veins distinct; scales not clathrate . . . . .. . Selliguea 31. Main lateral veins not distinct; scales clathrate . . . . . . . Loxogramme 29. Sori otherwise. S25 Sorivelongate, near margins ol thim fronds eyes) re) | enenennne Diblemma 32. Sori otherwise. 33. Fronds very narrow, not dimorphous; sori elongate, close to costa . Holcosorus 33. Fronds otherwise; sori round. 34. Fronds always coriaceous; edges usually notched; scales not clathrate. Crypsinus 34. Fronds not always coriaceous; edges not notched; scales clathrate Microsorium XVI Dec. 1959] ; Keys to the genera’ of Pteropsida GRAMMITIDACEAE Fronds simple and entire or with somewhat undulate rene a Sori round or obliquely elliptical, separate . . anes Grammitis 2. Sori elongate parallel to costa and edge, or confluent _ near apex of frond. 3. Sori in grooves between margin and costa. 4. Fronds almost terete apart from widened apical fertile part . . . . . Nematopteris 4. Fronds of about uniform width throughout . . . . . . . . . +. ~ Seleroglossum 3. Sori more or less confluent near apex, not in grooves . . . . . . . QOreogrammitis 1. Fronds pinnately lobed to bipinnately lobed. 5. One vein (sometimes forked) and one sorus on each lamina-lobe. 6. Fertile lobes flat. feesoOnl supericial . 7... Ee ee ets es ga ee a ONLCTAS 7. Sori deeply sunk in submarginal. pockets <0 Sea a ae ee ea ee Prosaptia 6. Fertile lobes partly reflexed, protecting the sori. 8. Lobes thin, only basiscopic margin reflexed . . . . . Calymmodon 8. Lobes coriaceous, both margins reflexed towards apex and. enclosing the sorus Acrosorus 5. Veins in lobes pinnately branched, each lobe with more than one sorus. 9. Sori superficial, or sunk in cavities perpendicular to the surface . . . . Ctenopteris 9. Sori deeply sunk in submarginal pockets not perpendicular to the surface Oe. Prosaptia REMAINING GENERA OF FERNS ADIANTUM Group 1. Water plants; sporangia borne singly and protected by reflexed edges of the lamina. Ceratopteris 1. Land plants; sporangia in sori or more or less acrostichoid. 2. Rigid hairs or bristles on rhizome and bases of stipes. 3. Fronds bipinnate, leaflets more or less lobed . . . . Sp Ss senkiseks me lesa ECL OSOEA 3. Fronds simple or simply pinnate, or palmate with entire leaflets. 4. Veins anastomosing only near edge of lamina; fronds simple or ae paraphyses (if present) with distinctive apical cell opr . . . Syngramma 4. Veins anastomosing throughout lamina; ‘fronds pinnate, trilobed or simple: paraphyses always abundant, hair-like, of many cells, apical one not different from the rest. 5. Sori broad, elongate, parallel to costa and edge, or more or less spreading along the veins. Taenitis 5. Sori in submarginal grooves, edges of grooves of equal thickness (Schizoloma sensu COPEL.). Schizolepton 2. Scales on rhizome and bases of stipes. €. Lower surface of fronds covered with white or yellow powder. 7. Sori along whole length of veins on lower surface . . Sa ty ohn oc , .ityRogramma 7. Sori at ends of veins, protected Dy reflexed lobes of margin sal ca Lesiceeis mip Cnelanees 6. Lower surface of fronds not covered with such powder. 8. Sori elongate (continuous or broken) along margins of lamina and protected by reflexed margin. 9. Fronds much dissected, the ultimate lamina-lobes small and connected bya narrow wing. 10. Fertile lobes much broader than sterile, the whole of each margin bearing a broad thin reflexed indusium. . aes of Onyehrom 10. Fertile lobes not much broader than sterile, reflexed margin not ‘continuous along larger fertile lobes nor very broad . . . . . Cheilanthes 9. Fronds, if much branched, having quite distinct leaflets not joined by a wing. 11. Fronds simple and lobed, or deeply pinnatifid; sori quite continuous along edge Doryopteris 11. Fronds branched with distinct stalked leaflets EAL tad tare eer teed te llaea 8. Sori otherwise. 12. Sori elongate along all or most of the veins. 13. Fronds simple . . a ee el Re pe Re ee een ee EICTNIONINEES 13. Fronds pinnate or bipinnate Re ee en oe ree a ee OF ey pe a OT rae ne 12. Sori otherwise. 14. Sori on lower surface, spreading a little along the veins . . . . . . Anogramma 14. Sori at ends of veins. 15. Sori on surface of reflexed marginal flaps . . . . Adiantum 15. Sori at ends of veins, sometimes more or less protected by reflexed marginal flaps Notholaena ASPLENIUM Group 1. Sori short, on one-veined ultimate segments of much-divided fronds . » 0 aatLoxoscaphe 1. Sori distinctly elongate along the veins; segments of frond usually with more than one vein.. XVII FLORA MALESIANA [ser. II, vol. 1 2. Sori on adjacent veins ae towards each other, a raised line remaining between the sori when they are ripe . . Diplora 2. Sori all facing the s same way except | ina 1 few species which have simple fronds and then the raised lines between, adjacentysonmlacking. "2-959 = ee ATHYRIUM Group 1. Veins free. 2. Groove of rachis open to admit grooves of branch rachises and midribs of leaflets. 3. At least some sori elongate along veins. 4. Sori not indusiate; fleshy outgrowths present at bases of pinnae on upper surface. Cornopteris 4. Sori indusiate; no such outgrowths. 5. Double sori (on both sides of a vein) not connected at their distal ends. . Diplazium 5. Double sori (at least some of them) connected at their distal ends; usually some reniform and J-shaped sori present Se a ie eee Tinea AAA Eee 8 ITT 3. Sori not elongate along veins. . 5 os me | ae OUST OMLCEIS 2. Groove of rachis not open to admit grooves of branches pea enc” ae Dryoathyrium 1. Veins anastomosing. 6. Sori almost circular, indusium reniform . . . . . - . . . . . +. Anisocampium 6. Sori elongate along veins. 7. Veins anastomosing regularly, about as in Cyclosorus; sori on nearly all veins of fertile frond. Callipteris 7. Veins anastomosing less regularly, the areoles adjacent to costae large; sori never so abundant. 8. Indusium rather firm, its edge reflexed at maturity . . oes Diplazium 8. Indusium thin, sausage-shaped, breaking at maturity, its edge not reflexed . . Diplaziopsis BLECHNUM Group 1. Sori not acrostichoid, or if apparently so, a thin indusium present all along edge of fertile leaflets. 2. Sori continuous along each side of the costa of a leaflet, a sterile lamina present beyond the sorus or not = -; Phebe Blechnum 2. Sori not continuous throughout length of a leaflet: one sorus ‘to ‘each areole along the costa. 3. Small ferns, simply pinnate; edges of pinnae sharply toothed . . 2 ee One 3. Large ferns, at least bipinnatifid; edges of pinnae not sharply toothed . . . Woodwardia . Sori acrostichoid (no thin reflexed indusium on fertile leaflets) . . . . . . +. Brainea DAVALLIA Group 1. Rhizome bearing hairs and scales; marginal cells of scales bearing papillose outgrowths Leucostegia 1. Rhizome bearing scales only; marginal cells of scales not bearing papillose outgrowths. 2. Fronds simple. 3. Sori elongate along the margin, protected by continuous indusia . . . . ._ . Parasorus 3. Sori singly at ends of veins. 4. Indusium attached at base only (or slightly above base) . . . . . . . +. Humata 4. Indusium attached along sides as well as base . . . . . . . +. +. ~~. + Seyphularia 2. Fronds pinnately branched. 5. Fronds more or less copiously persistently hairy; lamina thin; indusium small, of various shapes. Dayallodes 5. Fronds not hairy when mature. 6. Indusium attacheé along base and whole of sides. 7. Fronds trifoliate or simply pinnate with elongate narrow pinnae . . . . Scyphularia 7. Fronds more copiously branched . . . . Dayvallia 6. Indusium attached at base only, or also a little above the base, not along whole length of both sides. 8. Fronds much-branched, thin. . EE LO ON ES REY AUTESS LCE 8. Fronds of various form, lamina coriaceous or fleshy. 8: Scales: with lone*acicular tips 21'S SP Pe ee eee Be seales tacking lone acicular tips "3. 2 Sw DENNSTAEDTIA Group 1. Sori in cups at ends of veins. Paleai7Omemssicndcr creeping, Hairy” 2 ese ge) ve Ge ee ee a ee ts Dennstaedtia 2. Rhizome stout, erect, scaly. . . a ret a re ae Orthiopteris 1. Sori at ends of veins, variously protected, ‘not in cups. 3. Sori lacking protection; glandular hairs mixed with sporangia . . Monachosorum 3. Sori with indusium or covered when young by reflexed lobe of edge of lamina; no glandular hairs with sporangia. 4. Sori distinctly superficial, each at the end of a single vein and protected by an indusium which is XVIII Dec. 1959] Keys to the genera of Pteropsida pocket-shaped (attached by base and sides), opening towards margin. . . . M@éicrolepia 4. Sori more or less protected by reflexed edge (or lobes of edge) of lamina, with or without an inner indusium also. 5. Sori almost continuous along edges of lamina. 6. Veins anastomosing; NO’ Inner Indust yaes a) Ga Coslce)l) a/oc3 Oe) cee aeeishiopteris 6. Veins free; inner indusium present es , ‘ Pteridium 5. Sori singly at ends of veins, or uniting a few v veins, not continuous along all edges of lamina. PRINTING USIiINT fe O5s 5. Ge sie eee REE ise ole Soa 27) |S Se Hypolepis REET CINOUSINM Pleseht, .. .. .o iP eMC eel a ye Gee) a ee Paesia DICKSONIA Group 1. Fertile leaflets distinctly narrower than sterile. 2. Fertile pinnules deeply lobed; massive tree-ferns . . . . . . . «.. . Dicksonia 2. Fertile pinnules not lobed; stem short, creeping : SSVI Ona Wed FAal?: Cystodium 1. Fertile leaflets little different from sterile in size and shape. 3. Pinnules deeply lobed almost at right angles to costa, not pinnate ft Leste} Cibotium 3. Pinnules pinnate, tertiary leaflets small, very oblique and lobed. . . . . . . Culcita DRYOPTERIS Group 1. Fronds hairy as well as scaly on costae, costules and rachises; hairs septate, coarse, different from the scales. 2. Sorus when young spherical and quite enclosed by indusium which when ripe breaks to expose sporangia. BMESGOUIBONEGIStiINCL StalIkKS = \.) 3 oo « be Ce ee ee os ee ee | erAanema BESCIMINSESSIGGe tt gee) Ecc ne roa 2 eee cy Oe eee Sy es sek aco Diacalpe 2. Sorus otherwise; indusium attached on basiscopic side of sorus. 4. Hairs sparse; seales broad’ and thin’ %.) *..'-.- “2.4 | a bee, ce. oe, ReIOp ONES 4. Hairs copious; scales thick, elongate . . salle Stenolepia . Fronds scaly; no hairs as distinct from scales (some ‘scales may be narrow and almost hair-like). a Pinnules jointed to rachis; sori longer than wide . . . . . . . +. . Didymochlaena 5. Pinnules not jointed to rachis; sori not elongate. 6. Several vascular bundles in stipe. 7. Basal acroscopic leaflet or lobe of middle pinnae distinctly nearer to main rachis than basal basiscopic leaflet or lobe; fronds never simply pinnate. 8. Fronds elongate, basal pinnae not much enlarged on basiscopic side; rachis often bearing a bud. Polystichum 8. Fronds broadly deltoid, basal pinnae much enlarged on basiscopic side; rachis lacking buds. Polystichopsis 7. Basal acroscopic leaflet or lobe of middle pinnae not distinctly nearer to main rachis than basis- copic leaflet or lobe; or fronds simply pinnate throughout. 9. Indusium peltate; teeth on edges of lamina often with stiff slender points. . Polystichum 9. Indusium reniform; teeth not so produced . . ... . . Dryopteris faiwo vascular bundles in ibase of stipe) =) )6 "seh alesis. e- pee Gymnocarpium and Currania LINDSAEA Group 1. Fronds simply pinnate, veins free, pinnae numerous, spreading, jointed to rachis. . Isoloma 1. Fronds otherwise. 2. Pinnae or lobes elongate, bearing many sori, each at the end of a single vein; lamina rather thick. Tapeinidium 2. Pinnae or lobes, if elongate, bearing a sorus connecting the ends of several veins; lamina thinner. 3. Fronds finely divided, the ultimate divisions small and connected with each other by wings; sori solitary at apices of ultimate divisions . . . . . Sphenomeris 3. Fronds variously branched; leaflets distinct, each with several sori or a more or less continuous miieeinal SOruUS. ‘.. 5. .< Veins: free=ssterile’ frond? much divided". 92). © 5A. 19.1. OG Oe eee 6. Veins anastomosing. 7. Fronds deltoid, the basal basiscopic lobe of basal pinnae largest . . . . Stenosemia 7. Fronds otherwise, basal basiscopic lobes of basal pinnae not largest. 8. Fronds small, trifoliate; apical leaflet largest, commonly 5 cm long . . . Quercifilix 8. Fronds larger and in most cases more divided. 9. Sterile fronds simple to deeply pinnatifid, not truly pinnate . . . . Hemigramma 9. Sterile fronds with at least one pair of free pinnae, usually several pairs Heterogonium 5. Fertile leaflets not acrostichoid. 10. Two vascular bundles in base of stipe, uniting upwards to form a single bundle; scales confined to swollen bases of stipes . . . . . Hypodematium 10. More than two vascular bundles throughout ‘length of stipe; ‘scales not so confined. 11. Veins free. 12. Basal basiscopic vein of a vein-group springing from the costule. 13. Indusia peltate Paes : TED AEP ey ae ere! Bee Dryopolystichum 13. Indusia reniform or absent. 14. Fronds usually much longer than wide; basiscopic margin of lamina-lobes not thickened. Ctenitis XX Dec. 1959] Keys to the genera of Pteropsida 14. Fronds usually about as long as wide; basiscopic margin of lamina-lobes thickened. Lastreopsis 12. Basal basiscopic vein of a vein-group springing directly from the costa. 15. Basal pinnae with basal basiscopic lobes longest... . . Tectaria 15. Basal pinnae with basal basiscopic lobes or pinnules shorter ‘than middle ones. Heterogonium 11. Veins anastomosing. 16. Sori large, indusiate, terminal on a free vein, the receptacle elongate, fertile fronds narrow. Luerssenia 16. Sori various, indusiate or not, on free or netted veins; if indusiate, the receptacle not elongate, or fertile fronds not contracted as compared with sterile. 17. Basal pinnae deeply lobed, basal basiscopic lobe not largest; few free veins in areoles. Heterogonium 17. Basal pinnae lobed or not; if lobed, basal basiscopic lobe largest; many free veins in areoles. Tectaria THELYPTERIS Group ieeEronassbearing, many buds On Tachis; 50 - eesoa ce Weiner e em scl Ampelopteris 1. Fronds lacking buds. 2. Veins free (in some cases basal veins of adjacent groups just meet at the sinus). Thelypteris 2. Veins anastomosing. 3. Sori elongate along veins. 4. No indusia. 5. Venation as in Cyclosorus; veins from adjacent costules uniting to form a single excurrent vein. Stegnogramma 5. Venation irregularly SES COS: often with additional enclosed areoles . . Dictyocline 4. Indusia present. . : Sphaerostephanos 3. Sori not elongate, or slightly so if without indusia \(Abacepieris FéE, Haplodictyune PRESL). Cyclosorus VITTARIA Group 1. Frond very small, with one vein only; sorus near apex, along the vein ee Monogramma 1. Frond with lateral veins as well as a main vein. 2. Frond linear, sori one or more, on eas free branches, which run close to and parallel with main vein . . olen el (EATS b.8Vaginularia 2. Fronds otherwise; veins anastomosing ‘at least near “the margin. 3. Sori linear, in marginal grooves or superficial and parallel to margin; veins forming one series of areoles, anastomosing only near the margin aA . . Wittaria 3. Sori along veins, variously disposed; veins anastomosing “copiously throughout the lamina. Antrophyum MARSILEACEAE Solemeculisay arise EN? PRAVEEN ee ans Gs Jee ee) oe ee oe ne cee Vearsilea SALVINIACEAE 1. Leaves less than 1 mm long, bilobed, with one lobe submerged . . . ete Azolla 1. Leaves at least 1 cm long, simple, floating with upper surface fully exposed . . . Salvinia 7. BIBLIOGRAPHY A list of books and papers dealing with the taxonomy of Malaysian ferns, published subsequent to Christensen, Index Filicum, Suppl. 3 (1934) Aston, A. H. G., Fern notes II (New Guinea ferns). J. Bot. 77 (1939) 288-290. —, Undescribed ferns from New Guinea. Nova Guinea n.s. 4 (1940) 109-112, pl. 4-10; also in J. Bot. 78 (1940) 225-229, no plates. —, Some undescribed ferns from New Guinea and Ambon. Nova Guinea n.s. 7 (1956) 1-3. Aston, A. H. G. & R. E. Ho_trum, Notes on taxonomy and nomenclature in the genus Lygodium. Reinwardtia 5 (1959) 11-22. BACKER, C. A., Fam. Hymenophyllaceae—Equise- taceae. Beknopte Flora van Java (em.ed.) pts 1-2 (1940) fam. I-XV. BACKER, C. A. & O. PostHuUMUS, Varenflora voor Java. Buitenzorg, 1939. CHING, R. C., On the genus Onychium Kaulf. from the Far Orient. Lingn. Se. J. 13 (1934) 493-501. —, A revision of the compound leaved Polysticha and other related species in the continental Asia including Japan and Formosa. Sinensia 5 (1934) 23-91, 18 pl., 2 fig. —, On the genus Hypodematium Kunze. Sunyat- senia 3 (1935) 3-15. —, On the genus Pyrrosia Mirbel. Bull. Chin. Bot. Soc. 1 (1935) 36-72. —, On the genera Stegnogramma Bl. and Lepto- gramma J.Sm. Sinensia 7 (1936) 89-112. —, Arevisionof the Chinese and Sikkim-Himalayan Dryopteris with reference to some species of neighbouring regions, 1. Bull. Fan Mem. Inst. Biol. 6 (1936) 237-352. —, Ditto, 2-5. Bull. Fan Mem. Inst. Biol. 8 (1938) 157-268. —, Ditto, 6-9, l.c. 275-334, pl. 6-7. —, Ditto, 10. l.c. 363-507. —, The studies of Chinese Ferns XXXV. op. cit. 11 (1941) 79-82. —, On natural classification of the family “‘Poly- podiaceae’’. Sunyatsenia 5 (1940) 201-268, table. —, On the genus Gleichenia Smith. Sunyatsenia 5 (1940) 269-288. —, The Studies of Chinese Ferns, XX XI. Aleurito- pteris Fée. Hong Kong Nat. 10 (1941) 194-204. . CHRISTENSEN, C., New and noteworthy Papuan ferns. Brittonia 2 (1937) 265-317. —, Taxonomic Fern-studies, III. Revision of the genera and species of ferns described by A. J. Cavanilles. Dansk Bot. Ark. 9, no 3 (1937) 1-32. —, Ditto, IV. Revision of the Bornean and New Guinean ferns collected by O. Beccari and described by V. Cesati and J. G. Baker. l.c. 33-52. XXII —, Ditto, V. Descriptions of 36 new species of ferns. l.c. 53-73, pl. 1-6. —, Filicinae (Ch. XX) in Verdoorn, Manual of Pteridology. The Hague, 1938. CHRISTENSEN, C. & R. C. CHING, Pteridrys, a new fern genus from tropical Asia. Bull. Fan Mem. Inst. Biol. 5 (1934) 125-148, pl. 11-20. CLAUSEN, R. T., A Monograph of the Ophioglos- saceae. Mem. Torrey Bot. Cl. 19, no 2 (1938) 1-177, fig. 1-33. COPELAND, E. B., New or interesting Philippine ferns, VIII. Philip. J. Sc. 56 (1935) 97-110, pl. 1-14. —, Additional ferns of Kinabalu. Philip. J. Sc. 56 (1935) 471-483, pl. 1-10. —, The Philippine ferns collected by J. B. Steere. Philip. J. Sc. 60 (1936) 19-25. —, Solomon Island ferns. Philip. J. Sc. 60 (1936) 99-117, pl. 1-23. —, Hymenophyllum. Philip. J. Sc. 64 (1937) 1-188, pl. 1-89. —, Genera Hymenophyllacearum. Philip. J. Sc. 67 (1938) 1-110, pl. 1-11. —, Fern evolution in Antarctica. Philip. J. Sc. 70 (1939) 157-188, 2 maps. —, Oleandrid ferns (Davalliaceae) of New Guinea. Philip. J. Sc. 73 (1940) 345-357, pl. 1-10. —, Notes on Hymenophyllaceae. Philip. J. Sc. 73 (1941) 457-469, pl. 1-4. —, Holttumia, genus novum. Philip. J. Sc. 74 (1941) 153-156, pl. 1. —, Gleicheniaceae of New Guinea. Philip. J. Sc. 75 (1941) 347-361, pl. 1-6. —, Miscellaneous ferns of New Guinea. Philip. J. Sc. 76 (1941) 23-25. —, Filicum novarum CXXVIII diagnoses. Un. Cal. Publ. Bot. 18 (1942) 217-226. —, Ferns of the Second Archbold Expedition to New Guinea. J. Arn. Arb. 24 (1943) 440-443. —, Genera Filicum. Waltham, Mass., 1947. —, Cyathea in New Guinea. Philip. J. Sc. 77 (1947) 95-125, pl. 1-15. —, Pteridaceae of New Guinea. Philip. J. Sc. 78 (1950) 5-41, pl. 1-6. —, Aspleniaceae and Blechnaceae of New Guinea. Philip. J. Sc. 78 (1950) 207-229, pl. 1-6. —, Aspidiaceae of New Guinea. Philip. J. Sc. 78 (1951) 389-475, pl. 1-44. —, Grammitis. Philip. J. Sc. 80 (1952) 93-276, pl. 1-6. —, New Philippine ferns, IX. Philip. J. Sc. 81 (1952) 1-47, pl. 1-26. —, Grammitidaceae of New Guinea. Philip. J. Sc. 81 (1952) 81-119, pl. 1-10. —, New Philippine ferns, X. Philip. J. Sc. 83 (1954) 97-101, pl. 1-5. —, New Philippine ferns, XI. Philip. J. Sc. 84 (1955) 161-165, pl. 1-2. Bibliography Donk, M. A., Notes on Malesian ferns, I. On the genus Lemmaphyllum Presi. Reinwardtia 2 (1954) 403-410. Ho.ttum, R. E., The tree-ferns of the Malay Peninsula. Gard. Bull. Str. Settl. 8 (1935) 293-320, pl. 29-36. —, Notes on Malayan Ferns, with descriptions of five new species. Gard. Bull. Str. Settl. 9 (1937) 119-138. —, The genus Lomagramma. Gard. Bull. Str. Settl. 9 (1937) 190-221, pl. 8-16. , A redefinition of the genus Teratophyllum. Gard. Bull. Str. Settl. 9 (1938) 355-362, pl. 28-30. —, The fern genus Diplazium in the Malay Penin- sula. Gard. Bull. Str. Settl. 11 (1940) 74-108, fig. 1-6. —, New species of vascular plants from the Malay Peninsula (Filicales). Gard. Bull. Str. Settl. 11 (1947) 267-274, fig. 1. —, A revised classification of Leptosporangiate ferns. J. Linn. Soc. Bot. 53 (1947) 123-158. —, The classification of ferns. Biol. Reviews 24 (1949) 267-296, 1 fig. —, The fern-genus Heterogonium Presl. Sarawak Mus. J. 5 (1949) 156-166, 2 fig. —, The selection of type-species of some old genera of ferns. Gard. Bull. Sing. 12 (1949) 303-306. , Further notes on the fern-genus Heterogonium Presl. Reinwardtia 1 (1950) 27-31. —. The fern-genus Pleocnemia Presl. Reinwardtia 1 (1951) 171-189, 20 fig. —, The fern-genus Arcypteris Underwood. Rein- wardtia 1 (1951) 191-196, 3 fig. —, Anew fern from Malaya. Gard. Bull. Sing. 14 (1953) 8. —, A new Malayan Athyrium. Kew Bull. 1953, 545 (1954). —, Arevised Flora of Malaya, II. Ferns of Malaya. Singapore, 1954 (1955). —, Some additional species of Heterogonium. Reinwardtia 3 (1955) 269-274. —, On the taxonomic subdivision of the Gleichenia- ceae, with descriptions of new Malaysian species and varieties. Reinwardtia 4 (1957) 257-280. —, Morphology, growth-habit and classification in the family Gleicheniaceae. Phytomorpholo- gy 7 (1957) 168-184, 6 fig. —, The scales of Cyatheaceae. Kew Bull. 1957, 41-45. —, Notes on Malaysian Ferns, with descriptions of a new genus and new species. Kew Bull. 1958, no 3 (1959) 447-455. KRAMER, K. U., A new genus of Lindsaeoid ferns. Acta Bot. Neerl. 6 (1957) 599-601, 1 fig. Nakal, T., New classification of Gleicheniales, etc. Bull. Nat. Sc. Mus. Tokyo no 29 (1950) 1-71. PICHI-SERMOLLI, R. E. G., The nomenclature of some fern genera. Webbia 9 (1953) 387-454. —, Names and types of fern-genera, 1. Hymeno- phyllopsidaceae, Loxsomaceae, Schizaeaceae. Webbia 12 (1956) 1-40. —, Ditto, 2. Angiopteridaceae, Marattiaceae, Danaeaceae, Kaulfussiaceae, Matoniaceae, Parkeriaceae, Adiantaceae. Webbia 12 (1957) 339-373. PostHuMus, O., On some Malayan ferns. Proc. Kon. Ak. Wet. A’dam 39 (1936) 823-827. —, Malayan Fern Studies, I. The synonymy and distribution of the Java-ferns. Verh. Kon. Ak. Wet. A’dam sect. 2, 36, no 5 (1937) 1-67. —, Ditto, II. On the fern-flora of Australia, and its relation to that of neighbouring countries, especially of the Malay Archipelago. Verh. Kon. Ak. Wet. A’dam sect. 2, 37, no 5 (1938) 3-35. —, Ditto, II. The ferns of the Lesser Sunda Islands. Ann. Jard. Bot. Btzg, hors sér. (1943) 35-113. REnSCcH, I., Farne und Bdrlappe der Sunda-Ex- pedition Rensch. Hedwigia 74 (1934) 224-256, pl. VII. SELLING, O. H., Studies in the recent and fossil species of Schizaea, with particular reference to their spore characters. Act. Hort. Gotob. 16 (1944) 1-112, pl. 1-5. —, Two new species of Schizaea and their affi- nities. Svensk Bot. Tidskr. 40 (1946) 273-283, fig. 1-11. —, Further studies in Schizaea. Svensk Bot. Tidskr. 41 (1947) 431-450, fig. 1-14. WAGNER, W. H. & D. F. GreTHER, The Pterido- phytes of the Admiralty Islands. Un. Cal. Publ. Bot. 23 (1948) 17-110, pl. 5-25. XXIII ‘a i bg vinegar Leer SPSS fare = a0- WIA as eer ad (ete ie an) er oe | m o 2 ane me ews a Sh = or i b elie) .@ 4-- +h wet al i > ; é eZ a het et ie CUT Mn “= roe .?, } a Piven ye) A’ / * - . KINGS: - 2 Btegl 4 y 5 i { ( a i v0 4i* : yee 7 — aie wins ¥ . yee Za ' 1 ss > = een 3 ¥ f = + . , 4 .- 3 - ann +f “SS : mY f 3 Le if q v1 Ss f e ~ F ~ * 50 s 4 - : ‘ ; q ’ 7a - =. 2 ( - 7 3% oes ti 4 ‘ ‘ 47 - - Ake ' = 3 - Ths), tery. Cease rand ai subse iv bi ad tnd ' rf vy . fod “ha, « ) ie aad ? a 39 a Fb qe'l <= i ia? P tR-i igrehyin 2 Stabs] ye Seaver! wis ek ANS many ioe Ey eld Fe] —=iiqta bois ene te Tot : ate ») calhats a’ ww \ J A = ‘7 t ‘ % A. 7 4. on, . ahs, 4SAY thy Ch orfates «. yp PEED po, oo a a a? = » Forraiare ccilg ete? wows > 4 a. 2 Ai Ww ae ea olt bt Sg) ray arte 9G AD) sed =A eet fab ena, vey adie . A ee ak aie at _ ss ame ary heey? Par pv aA tz ) er a ~*~ = I> sie pad Bite Nat 2341 98)7 16 jive af Me 2 a ie 1s ow Le GLEICHENIACEAE (R. E. Holttum, Kew) Rhizome relatively slender, creeping, protostelic (solenostelic only in Dicranopteris pectinata (WILLD.) UND. of tropical America), in Stromatopteris bearing erect irregularly dichotomous branches which bear the fronds, in all other cases bearing fronds directly; young parts covered with peltate fringed scales (scales otherwise in Stromatopteris) or branched hairs. Fronds unbranched in Stromatopteris, in all other cases branched in fully developed plants, the main rachis bearing a series of pairs of branches, its apex periodically dormant while each successive pair of branches develops; each primary branch often bearing a pair of secondary branches and a permanently dormant apex between them, the process some- times repeated several times; ultimate branches either bipinnatifid or pinnatifid, the lamina (whether of an ultimate branch, or leaflet of an ultimate branch) cut almost to the costa; veins in lamina-segments pinnate, branches simple or forked, free (in some cases apparently joining a thickened non-vascular margin). Sori of 2-15 or more sporangia, attached to a small receptacle on the surface of a vein (except in Stromatopteris, where each sorus is spread along part of both branches of a forked vein), never at the end of a vein, all sporangia in one sorus developing simultaneously; branched hairs or scales often present with sporangia but no indusium. Sporangia with complete oblique annulus, dehiscing vertically, containing c. 200-800 or more spores. Spores monolete or trilete, smooth, translucent, colourless. Gametophyte (not known in Stromatopteris) at first cordate, then ribbon-like with heavy midrib, finally branching at apex; rhizoids stiff, abundant, usually reddish-brown; two-celled glandular hairs developed by many species in as- sociation with archegonia and also on margin; antheridia comparatively large and complex in structure (some more so than others); archegonia with long necks (longest in Gleichenia subg. Gleichenia) directed towards apex of prothallus; no cases of apogamy observed. Distribution. Throughout the wetter parts of the tropics and subtropics, and in south temperate regions. Three genera: Stromatopteris (monotypic, in New Caledonia), Gleichenia (3 subgenera, c. 150 species, in the tropics mainly on mountains, at lower altitudes in southern temperate regions), and Dicranopteris (2 subgenera, c. 10 species, mainly tropical, at low and moderate altitudes). Dicranopteris is much more polymorphic in Malaysia than in any other part of the tropics. (The genus Platyzoma is excluded from the family; see HoLTTUM in Kew Bulletin no 3, 1956, p. 551). Fossils. The fragmentary nature of the earlier fossils ascribed by various authors to the family makes judgment upon them difficult. The form and arrangement of sporangia is the best criterion; on this basis paleozoic fossils named Oligocarpia have been assigned to Gleicheniaceae, but they do not always show details of structure of the individual sporangia clearly. Some have been found on fronds of the Pecopteris type (not unlike Gleichenia of today), some on the rather different Sphenopteris, but none show pseudo-dichotomy of the kind now universal in the family. It may be that some of these fossils represent members of the family before periodic growth and pseudo-dichotomy developed. Gleichenia gracilis ZiGNo, from the Jurassic of northern Italy, has been more confidently referred to the family by paleobotanists; but the main branches of the frond are not opposite, and the irregular forking of the smaller branches looks more like true dichotomy than pseudo-dichotomy (judging from ZIGNO’s figure); there are no sori. Wealden fossils from Belgium show anatomical structure of rhizome and petioles as in living Glei- chenia. In somewhat later Cretaceous rocks of various parts of the world, but especially in West Green- land, are fossils which show every character (except scales, which have not been seen) of existing members of Gleichenia subg. Diplopterygium (HEER, Flora Fossilis Arctica, II[) and some are very like subg. Gleichenia, with more or less transitional stages between the two conditions which are not shown by living members of the family. Some of them show permanent dormancy of a lateral branch of a frond, but none show the condition either of Gleichenia subg. Mertensia or of Dicranopteris, which have what I regard as the most highly developed forms of branching. Some Greenland fossils show many sporangia in a sorus, a character not shown by any living member of the family having similar leaf-form, and TuTIN (Ann. Bot. Lond. 46, 1932, p. 503-508) has described Gleicheniopsis with numerous small (1) 2 FLORA MALESIANA [ser. Il, yoru sporangia each containing less than 32 spores; the latter were not attached to any forked axis. The only fossil known to me which corresponds in leaf-form with Dicranopteris is Gleichenia han- tonensis WANKLYN from Eocene beds at Bournemouth (southern England). This was the subject of a reconstruction, copied by other authors, by GARDNER and ETTINGSHAUSEN (British Eocene Flora, I fig. 28) which (on the same frond as typical Dicranopteris ultimate branches) incorporated accessory branches as in living D. linearis (not shown in the fossils) and also some curious hook-bearing leaflets which are associated with the Dicranopteris fossil leaflets but nowhere attached to them. Disregarding these unwarranted additions, Gleichenia hantonensis agrees with Dicranopteris in branching, venation and sori (position, shape and number of sporangia); if hairs could be found, agreement with Dicranop- teris would be complete. The Greenland and Bournemouth fossils offer evidence hard to reconcile with COPELAND’S opinion that the family is of antarctic origin (Genera Filicum, p. 26). Ecology. Allspecies are sun-ferns, and most form thickets, to which they are adapted by their creeping rhizomes, and by the indefinite growth in length of the fronds (at least in the wet tropics). Periodic dormancy of the apex of the main rachis permits periodic upward growth of the rachis-apex, unen- cumbered by branches; it may thus pass between any other leaves or branches which may be above it, and then develop its new pair of leafy branches in a fully exposed position while the apex itself is again dormant. The most efficient members of the family from this standpoint are species of Dicranopteris and of Gleichenia subg. Mertensia, because by repeated forking their lateral branch-systems produce a spreading fan-shaped arrangement, but it is notable that in Dicranopteris a reversion to an effectively pinnate form of branching has occurred, seen at its full development in D. speciosa (PR.) HOLTT. Most members of the family are pioneer plants, establishing themselves on bare ground, sometimes in fully exposed places, talus, earthslides, precipices, rocks (fig. 3), steep ridges (fig. 2, 10), often on poor rocky or leached soils. Prothalli (which are often very abundant) need a little shelter, but a young sporophyte can quickly spread by means of its rhizome, and Dicranopteris linearis, for example, can grow in places where few other plants can establish themselves. A Dicranopteris thicket, once formed, can persist for a long time (fig. 11, 13), unless tree seedlings have established themselves in it at an early stage (shade of trees weakens or kills Dicranopteris plants unless they can climb); such thickets have been greatly encouraged by man-made clearings of forest. On the edge of forest, some of the larger members of the family can climb to a considerable height. At high altitudes on mountains, where slower-growing alpine scrub occurs, smaller species of the family are often abundant and form lower thickets. They may grow in association with Sphagnum, or in dry rocky places. An important character is that their demands of mineral substances are small. Vegetative morphology. Stromatopteris is peculiar in its erect rhizome-branches, in its scales and hairs, and in having sori spread along both branches of a vein. NAKAI (Bull. Nat. Sc. Mus. Tokyo n. 29, 1950) has proposed its separation as a distinct subfamily, and I consider this separation fully justified. The genus does not occur in Malaysia. It is probably to be regarded as a reduced relic of an otherwise extinct group. As above noted (under Ecology), the fronds of other members of the family are all branched (except in young or stunted plants), the branching being pinnate in plan, with dormancy, either periodic or permanent, of apices of various orders. Fig. 1, 7, 12. The branch-patterns are in some cases complex, and are characteristic in the different genera and subgenera (for a comparative account, see HOLTTUM, Phytomorphology 7, 1957, 168-184). FILARsKy attempted a detailed analysis of these branch-patterns (Ann. Hist.-Nat. Mus. Nat. Hungarici 20, 1923, 1-23; ibid. 21, 1924, 163-170) but the results seem to me confused. He did not distinguish clearly between Dicranopteris and Gleichenia subgen. Mertensia of the present treatment, nor between Platyzoma and Gleichenia subgen. Gleichenia. The form of branching shown in his fig. 12 and 24 is one I have never seen (shorter branches on the same side at successive unequal forkings). The major divisions of the family are also distinguished by their dermal appendages. Those of Stro- matopteris are quite peculiar (scales, only on rhizome, not peltate, and long simple hairs also on rhizome). The rest of the family have either fringed peltate scales and stellate hairs with unicellular rays (Gleichenia) or branched hairs of complex structure, true scales being absent (Dicranopteris, fig. 14). The venation in Gleichenia is pinnate in each segment of the lamina, lateral veins being forked (fig. 8c) except in subg. Gleichenia (where the area of the segment is very small). In Dicranopteris the lateral veins are at least twice forked. Fig. 15a, c. The sori are always attached on the surface of a vein, the sporangia attached to a small raised receptacle. In Gleichenia subg. Gleichenia the veins are very short, and are not visible unless the lamina is cleared; in many taxonomic works the sori are said to be ter- minal on the veins in this subgenus, but cleared specimens show that they are not. The arrangement of sporangia in a sorus, in both genera, has been discussed and illustrated by Bower (The Ferns, vol. 2, 203-206, fig. 476, 486-489). In some species of Gleichenia subg. Gleichenia the sori are in depressions in the substance of the lamina (e.g. G. peltophora CopPeEL.); in others they are protected by the reflexed margins of the lamina and by outgrowths of tissue from the costa (G. vulcanica BL., fig. 1d—e). NAKAI (/.c.), following PRESL, attempted to use these characters to establish separate genera, but the species G. microphylla R.Br. is intermediate. Sporangia and spores. The structure of sporangia has been fully described by Bower (i.c.). Both trilete and monolete spores occur in the family. NAKAI (/.c.) proposed a basic division of the family Dec. 1959] . Gleicheniaceae (Holttum) ; 3 on spore-form, but in so doing he supposed that only trilete spores occur in his restricted genus Di- cranopteris (D. linearis and its near allies); in fact closely related species of this group have spores of different forms, and a division of the family on spore-form is certainly unnatural. Gametophyte. The most recent and most complete account of gametophytes in the family is by A. G. Stroxey (Bull. Torrey Bot. Club 77, 1950, 323-339) and includes references to earlier accounts. Species of all genera and subgenera here recognized were studied. Dr Stokey’s conclusion is that the gametophyte in all cases shows many primitive characters, but there is little significant difference within the family to indicate that one part is more primitive than the rest. Gleichenia subg. Gleichenia shows specialized characters (notably the long neck of the archegonium) and on characters of the gametophyte is judged to be further removed from the rest of the family than they are from each other. Two-celled hairs of a peculiar nature have an origin similar to that of the larger hairs on prothalli of Cyatheaceae; apart from these hairs, the prothalli of Gleicheniaceae most resemble those of Dipteris. Cytology. MANTON & SLEDGE (Phil. Trans. Roy. Soc. B, 238, p. 143, pl. 4) report for Dicranopteris linearis from Ceylon both n = 39 and n = 78; from Singapore, for what is now recognized as D. cur- ranii COPEL., n = 39. MANTON reports verbally that a plant of D. linearis sent from Singapore and cul- tivated at Kew is a Sterile triploid hybrid. MEHRA & SINGH also report n = 39 for D. linearis from Northern India (Curr. Sc. 25, 1956, 168), and n = 56 for Hicriopteris glauca (probably Gleichenia gigantea WALL., which is the common Himalayan species of this group), a species of Gleichenia subg. Diplopterygium. BROWNLIE (Trans. R. Soc. N.Z. 85, 1958, 213-214) reports n = 20 for G. (subg. Glei- chenia) microphylla R. Br. and n = 34 for G. (subg. Mertensia) cunninghamii. T. G. WALKER reports (personal communication) n = 34 for two species of subg. Mertensia from Jamaica. Anatomy. The rhizome has a simple protostelic structure except in Dicranopteris pectinata (see Bower, /.c.). The rachis has a single C-shaped vascular strand; all the outer tissues in the rachis are thick-walled and when mature form a very strong protection for the vascular strand. Such protection is important in fronds which continue to grow in length for a long period. Economic importance. HEYNE (Nutt. Pl. N.I. 1927, p. 97) records the following uses for parts of plants of this family; the chief species used are Dicranopteris linearis and D. curranii. The rachises of mature fronds are tied into bundles and the bundles used in making the fences of a certain kind of fish trap; they will last two years when immersed in sea-water. Parts of the rachis of a large frond (the largest are produced by D. curranii) when suitably split make excellent pens for writing Arabic characters (I learned of this use also in Singapore). The vascular strands of stipe and rachis are separated and used for special kinds of fine plaited work, being strong and pliable. Dicranopteris thickets may be useful as preventing erosion, but they are troublesome to the forester when they prevent regeneration of tree-seedlings in forest clearings. As the rhizomes are almost or quite superficial, they are exposed when the thicket is cut down, and usually one such cutting is enough to kill almost all of the plants. Fronds are sometimes cut — when in absence of other suitable material — to provide light shade for transplanted seedlings. Taxonomy. I published a statement on taxonomy, with discussion of the present arrangement, in 1957 (Reinwardtia 4, p. 257-280). CopELAND (Gen. Fil. 1947) divided the Malaysian members into four genera. It seems to me, however, that Dicranopteris is so different from the rest that a main division should indicate this difference, and I therefore recognize two genera, Gleichenia and Dicranopteris, the former with three subgenera which correspond to COPELAND’s genera. For one of the latter he used the name Hicriopteris PRESL, but the type species of that genus is a Dicranopteris (described as D. speciosa in the present work). I have therefore adopted the subgeneric name Diplopterygium, first proposed (as a sectional name) for this group of ferns by DreLs (in ENGLER & PRANTL, Pfl. Fam. 1, 4, p. 350-356). For the other subgenus the name Mertensia is available; it was first used in this rank by Hooker, though with a larger content. This name was first proposed by WILLDENOw in 1804 as a generic name to cover all known members of the family with larger divisions of the lamina than the original G. polypodioides (THUNB.) SMITH; but as a generic name it was antedated by Mertensia RoTH (Boraginaceae) and so it is illegitimate. In 1806 BERNHARDI published the generic name Dicranopteris, citing under it only one species of Mertensia WILLD., viz Polypodium dichotomum THUNB., which thus becomes the type species of Dicranopteris. I have typified Mertensia by the species M. truncata WILLD. (Reinwardtia 4, 1957, 261). Both Mertensia and Dicranopteris are used here in a more restricted sense than intended by some earlier authors. COPELAND used the generic name Sticherus PRESL for subg. Mertensia of the present account. This name was established by Pres for two species of which he had seen no specimens, with a brief and confused description, and the lapse of the name is not to be regretted. KEY TO THE GENERA 1. Young parts of plants protected by fringed peltate scales and stellate hairs. Sori of 2-5 sporangia. Veins simple or once forked. . ai gate 1. Gleichenia 1. Young parts of plants protected by branched hairs of various s form, scales lacking. Sori of 8-15 or more sporangia. Veins forked at least twice... = 9 -..4) ©) « «..« « 2. Dicranopteris. + FLORA MALESIANA [ser. II, vol. 12 1. GLEICHENIA SMITH, Mem. Ac. Turin 5 (1793) 419, non NEcK. 1790, nom. cons. — For synonyms see the subgenera.—Fig. 1-10. Rhizome dichotomously branched, protostelic, near the apex protected by peltate scales. Fronds of mature plants of indefinite growth in length (except sometimes at high altitudes), bearing primary branches in pairs, the apex of the main rachis dormant during the development of each pair of primary branches, the dormant apex in some cases protected by a pair of stipule-like leaflets of distinctive form (such stipular leaflets less often present in conjunction with dormant apices of lateral branch-systems); primary branches often each bearing a pair of secondary branches with a usually dormant apex between them, the process sometimes repeated to produce ultimate branches of fourth or fifth orders; dormant apices protected by peltate scales which (with two exceptions) are fringed by outgrowths from the marginal cells; u/timate branches either simply pinnatifid or bipinnatifid; /amina in all cases lobed almost to the costa, the veins in each segment pinnately branched; lateral veins simple or once forked; costae, costules and veins when young protected by small fringed peltate scales and by stellate hairs, sometimes glabrescent when mature; sori one or several to each segment of the lamina, upon the lateral veins, not terminal upon them (on the acroscopic branch of a forked vein), exindusiate, each consisting of 2-5 large sporangia; paraphyses, in the form of small stellate hairs or small scales with long marginal hairs, often present with the sporangia; annulus complete and oblique, dehiscing dorsally. Spores monolete or trilete, smooth and translucent, 256 or more in each sporangium. KEY LO DME SRE CIES 1. Ultimate branches (branches on each side of an ultimate dormant apex) bipinnatifid. 2. Segments of the lamina not much longer than wide; one sorus on each segment. (subg. Gleichenia). 3. Fully developed pinnule-lobes not deeply concave beneath (the edges only slightly revolute). Tissues adjacent to costa not swollen. Sporangia when young in a circular depression in the lamina. 4. Edge of depression occupied by sorus distinctly raised. Small scales on lower surface of lamina. 1. G. peltophora 4. Edges of depression occupied by sorus not raised. No small scales on lower surface of lamina. 2. G. microphylla 3. Fully developed pinnule-lobes deeply concave beneath, the distal edges strongly revolute. Tissue adjacent to the costa more or less swollen. Sporangia not in a depression in the lamina. 5. Costae persistently scaly; tissue adjacent to the costa slightly swollen. Sporangia often 3. 3. G. yulcanica 5. Costae of fully developed fronds usually quite glabrous; tissue agent to the costa much swollen. Sporangia 2. . : F 4. G. dicarpa 2. Segments of the lamina elongate, the costule bearing several forked veins, each of which may bear a sorus on its acroscopic branch. (subg. Diplopterygium). 6. Scales on dormant apex of rachis entire, to 10 by 3 mm. Segments of lamina at c. 45° to costae. 5. G. laevissima 6. Scales on dormant apex of rachis fringed with hairs or setae, or with a broad translucent margin. Segments of lamina almost at right angles to costae. 7. Scales on dormant apex of rachis 1 mm wide or more, edges fringed with spreading hairs to 0.5 mm long, or with translucent edge bearing fine short hairs. 8. Rachis-branches persistently quite covered with scales, or with a mixture of scales and hairs, on lower surface. 9. Rachis-branches covered almost entirely with thin brown scales 4-5 mm long and nearly 1 mm wide, stellate hairs few. . . . 6. G. paleacea 9. Rachis-branches covered with a dense felt of stellate hairs, and also with + abundant scales. 10. Pinnules commonly 12-16 by 2-21/, cm. Upper surface of the lamina “not swollen between the veins. Stipular leaflets present . . . . . . 7. G. yolubilis 10. Pinnules commonly 6-9 by 0.9-1.3 cm. Upper surface of the lamina much swollen between Dec. 1959] Gleicheniaceae (Holttum) 5 the veins. No stipular leaflets .. 8. G. bullata 8. Rachis-branches not persistently quite covered with scales nor r with ¢ a felt of stellate hairs on the lower surface. 11. Very few persistent scales on the lower surface of rachises and costae. Pinnules to 22 by 3.5 cm. 9. G. longissima 11. Many persistent dark scales on rachises and costae. Pinnules to 12 by 2 cm 10. G. clemensiae 7. Scales on dormant apex of rachis narrower, their edges bearing short stiff oblique setae. 12. Segments of the lamina near base of each pinnule constricted at the base (i.e. widening just a little above the constriction), joined together by a very narrow wing of even width along the costa. 13. Many such segments (20 or more pairs) on each pinnule. 14. Lamina rigid; segments of lamina ae mm wide; scales of dormant apex uniformly dark DIOWDs = |. . . . 11. G. angustiloba 14. Lamina thin; segments of lamina 31), mm wide; scales of dormant apex pale with dark edges. 13. G. deflexa 13. Few (at most 7-8) such segments on each pinnule. 15. Pinna-rachises more or less persistently scaly; costules 31/,-41/, mm apart. 14. G. sordida 15. Pinna-rachises sparsely scaly, soon glabrous; costules 5!/,-7 mm apart. 12. G. elmeri 12. Segments of lamina near base of each pinnule not thus constricted (or not more than one segment on the largest pinnules). 16. Stipular leaflets having lobes similar to pinnule-segments, not narrow and acuminate. Scales on rachis-apex not dark-fringed. . . eye 15. G. norrisii 16. Stipular leaflets having very narrow acuminate lobes quite different from normal segments of pinnules. Scales on rachis-apex dark-fringed. 17. Distal half of each rachis-branch strongly zig-zag, the pinnules deflexed so that each is IimUINem with thempanrt. ofethesrachisebeyondeit sea) Hen t eee = 16. G. matthewii 17. Distal half of each rachis-branch not thus zig-zag. I, Praimwes ClO x7 2 Cal, 4 ee ERC. eer Poel Ur, 17. G. brevipinnula 18. Pinnules commonly 20 by 3-4 cm. 19. Dark rigid stellate hairs abundantly persistent on the lower surface of veins and lamina. 18. G. blotiana 19. Stellate hairs light brown, not abundant nor Been on the lower surface of veins and lamina. ee Ce eee 19. G. conversa Ultimate branches simply, pinnatiide Gee Mertensia): Sh Stipule-like leaflets, different from ordinary segments of the lamina, present at the branchings of the main rachis (on fronds of mature size). 21. Segments of lamina at right angles to the costa, or more or less deflexed. Lower surface not con- spicuously glaucous. 22. Segments of lamina deflexed (lower ones much so), thick, with edges much revolute when dry. Main rachis-branches once or twice forked . . . rae es 20. G. reflexipinnula 22. Segments of lamina not or little deflexed. Main rachis- branches often more than twice forked. 23. Forkings of rachis-branches equal or nearly so, not alternately unequal with larger branches at successive forks forming almost a straight line. 24. Scales on costules very abundant, spreading so as almost to cover the lower surface of the lamina. Veins on lower surface strongly raised . . . 21. G. venosa 24. Scales on costules not persistently covering the lower surface of ‘the lamina. Veins not strongly raised ‘ : 22. G. truncata 23. Forkings of rachis- branches unequal, ‘the larger branch at successive > forks alternately to left and right, axes of these larger branches in almost siraight line. . . . 23. G. milnei 21. Segments of lamina distinctly oblique-ascending. Lower surfaces distinctly glaucous. 25. Primary branch-systems 5 times forked on large fronds, branches of all orders except the first leafy; no stellate hairs on the lower surface of veins . . 8 24. G. oceanica 25. Primary branch- ~systems twice forked; stellate hairs (or at least their bases) persistent on the lower surface of veins . . a: 25. G. erecta 20. Stipule-like leaflets absent (sometimes present in G. lohers var. aoe. 26. Margins of segments of lamina not distinctly toothed. 27. Only the ultimate branches fully leafy (penultimate without lamina or with some scattered segments). 28s Witimate, branches: to20)cmylongieeie . 9 linea ecu ees Senn eeeee2O.8G. opulchra 28. Ultimate branches 30-40 cm long. 29. Segments of the lamina to 25 mm long, costules aye mm apart. Primary rachis-branches usually 3 times forked ii ‘ 27. G. pseudoscandens 29. Segments of lamina to 17 mm long, ‘costules 4 mm apart. Primary rachis-branches usually twice forked 5 2142-98285). G.walstonii 27. At least the penultimate ‘branches fully leafy, lower ones ‘sometimes also. 6 FLORA MALESIANA (ser. Ik volar 30. Lamina-segments rarely more than twice as long as the distance between bases of adjacent costules. 31. Segments 8-12 mm long, costules 4-5 mm apart. Rachis-branches of Ist and 2nd orders 3-4 cm long. Scales on the main as ek oe brown, more than | mm wide at base; scales on the costae rusty brown 3 aS 29. G. vestita 31. Segments 3-6 mm long, costules 3 mm apart. Rachis- branches of ist and 2nd orders 1—2 cm long. Scales on the main peut very dark, narrower; scales on the costae dark at base, paler distally 30. G. bolanica 30. Lamina-segments much 1 more than twice as Jong as the distance between adjacent costules. 32. Scales on the costae only a few cells wide at the base, about half their length consisting of a hair, these hairs crisped and entangled » ids : 31. G. hispida 32. Scales on the costae with short hair-points, their bases wider and their apices not entangled. 33. Costules at 60°-70° to costae : : 25. G. erecta 33. Costules at almost a right angle to the costae. 34. Ultimate branches 25-45 cm long, eee cm wide; costules on ultimate branches 4-5 mm aparte yee. . . 32. G. brassii 34. Ultimate branches smaller, ‘costules on ultimate branches 3-4 mm apart. 33. G. loheri 26. Margins of the segments of the lamina distinctly toothed, at least towards the apex. 35. Margins toothed almost to base of segments. Costules at c. 45° to costae. 34. G. flabellata 35. Margins toothed only near apex of segments. Costules at wider angle to costae. 35. G. hirta 1. Subgenus Gleichenia Hott. Reinwardtia 4 (1957) 262.—Calymella Pres_, Tent. Pterid. (1836) 48; CHING, Sunyatsenia 5 (1940) 287; Naxkal, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 40.—Gleicheniastrum Pres_, Abh. (K.) Bo6hm. Ges. Wiss. M.-N. Cl. 5 (1848) 338; NAKAI, /.c. 42.—Gleichenia subg. Eugleichenia Diets in E. & P. Pfl. Fam. 1, 4 (1900) 355.—Gleichenia; CopeEL. Gen. Fil. (1947) 26.—Fig. 1-3. Young plants first producing determinate bipinnatifid fronds, the largest of these as large as the branches of later fronds and sometimes fertile (plants in exposed places at high altitudes sometimes producing only fronds of this kind); then fronds bearing pairs of primary branches like the fronds of the first stage; in some cases the primary branches bearing one or more pairs of determinate bipinnatifid secondary branches, or the secondary branches leafless and bearing pairs of tertiary branches, the ultimate branches always determinate and _ bipin- natifid. Leaflets of bipinnatifid fronds or of ultimate branches lobed almost to the costa, lobes hardly longer than wide; veins in each lobe pinnately branched but without a conspicuous costule, lateral veins simple. One sorus on each lobe, upon the basal acroscopic vein, superficial or sunk in the substance of the lamina. Distr. About 10 spp. in tropical and southern Africa, the Mascarene Islands, Malaysia and Austra- lasia (not in Ceylon and India). Ecol. In Malaysia only on mountains, in exposed places, usually in sandy or acid, peaty soils, sometimes in association with members of the other subgenera. 1. Gleichenia peltophora CopeL. Philip. J. Sc. 40 (1929) 292, t. 1.—G. circinnata var. borneensis BAK. J. Bot. 17 (1879) 37.—G. borneensis C. Cur. Gard. Bull. S. S. 7 (1934) 211.—Calymella bor- neensis CHING, Sunyatsenia 5 (1940) 288. var. peltophora. Rhizome-scales ovate, dark, entire. Determinate bipinnatifid fronds to 40 cm high (including stipe), often fertile; branched fronds bearing 1 or few pairs of branches; primary branches bipinnatifid- determinate or once forked with a permanently dormant apex in the fork, the primary branch always bearing leaflets (where primary branches are not forked, the main rachis may be leafy below junction with branches); sca/es on resting apex of rachis to 11/, by 1 mm, ovate, entire, convex, shining, very dark brown with narrow pale edge; similar but smaller scales scattered on lower sur- faces of rachises and less abundantly on costae; ultimate branches 20-30 cm long, bearing many leaflets, costae of adjacent leaflets 4-6 mm apart; leaflets 2-5 cm long, 2!/,-4 mm wide; lobes c. 114 mm wide at base, gradually narrowed to rounded apex, edges slightly reflexed, lower sur- face slightly concave, glaucous, bearing scattered circular scales 0.2-0.3 mm diameter. Sori, if present, each sunk in a circular depression in the substance of the lamina, the lower surface of the lamina somewhat raised round the edge of the depression which occupies about half the width of the base of a lobe. Type: Copeland s.n., 1 May 1917, Mt Matutum, Mindanao, 1600 m (H. Copel.). Dec. 1959] Gleicheniaceae (Holttum) T Fig. 1. Gleichenia microphylla R. Br. a. Lower surface of part of a pinnule, showing one mature sorus, and two soral depressions from which sporangia have been removed, x 10, 5. one lobe of lamina, cleared to show veins and position of sorus, 13, c. a scale, x 50. — G. vulcanica BL. d. Part of pinnule _ from below (scales removed), x 10, e. same cleared to show veins and position of sorus, x 13. —G. dicarpa R. Br., f. Lower surface of part of pinnule, « 10, g. same cleared to show veins, 13, A. upper surface of cleared pinnule showing position of sori. — Diagrams of various stages of branching which may occur during development of a single plant of G. vulcanica BL., i. First stage: plants in exposed places or at high altitudes may never develop further, j. transition stage, k. primary branches simple (corresponding to condition of subg. Diplopterygium), 1. primary branches once forked, m. primary branches twice forked, n. primary branch proliferous beyond the first fork. Distr. Malaysia: N. Borneo (Mt Kinabalu), Philippines (Mindanao), South Central Celebes, West New Guinea (several localities). Ecol. In open places on mountain summits or high ridges, 1500-2500 m; on Mt Kinabalu in association with G. vulcanica and Dipteris novo- guineensis POSTH. var. schizolepis C. CHR. ex HoLtrT. Reinwardtia 4 (1957) 262. Scales on rhizome short-fringed; scales on costae bearing a few rigid dark brown marginal hairs; scales on lower surface of lamina replaced by dark red-brown stellate hairs; fronds of the only speci- men unbranched, with leaflets 2!4 cm long, sterile. Distr. Borneo (Sarawak: Mt Murud), 2500 m, once found. 2. Gleichenia microphylla R.Br. Prod. (1810) 161; y. A. v. R. Handb. Suppl. (1917) 80, incl. var. semivestita v. A. v. R.—G. semivestita LABILL. Sert. Austro-Cal. (1824) 8, t. 11.—Calymella microphylla PResL, Tent. Pter. (1836) 49.— Gleicheniastrum microphyllum Pres_, Abh. (K.) Bohm. Ges. Wiss. M.-N. Cl. 5 (1848) 338; NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 45, incl. var. semivestitum NAKAI.—Calymella circinnata (7G. circinnata Sw.) CHING, Sunyatsenia 5 (1940) 288.—Calymella semivestita CHING, I.c.—Glei- cheniastrum lowei NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 44.—Fig. la-c, 2. Rhizome to 3 mm diameter, the young parts covered with narrow dark brown rigid fringed scales. Fronds on young plants _bipinnatifid, determinate, rarely fertile, on old plants usually FLORA MALESIANA [ser. Tl, volaet Fig. 2. Gleichenia microphylla R. Br. on edge of dwarf forest on wet sandy soil near summit of Kedah Peak, Malaya, 1200 m (HoLtrum, 1953). Dec. 1959] Gleicheniaceae (Holttum) i 9 branched, the primary branches sometimes forked with leaflets below the fork, rarely proli- ferous beyond the fork and then not bearing leaflets below the fork; secondary branches sometimes also forked. Sca/es on resting apices of rachis narrow, very dark, with pale thin marginal hairs towards the base and rigid dark setae near apex; rachises and costae more or less densely clothed with very small dark scales, their edges bearing rigid concolorous setae. Ultimate branches 12—25 by 4-5 cm; leaflets 7-9 mm apart, 2!4-4 mm wide; lobes of /amina 1—1.8 mm long, 0.7—1.5 mm wide at the base, apex rounded, edges usually a little reflexed; lower surface slightly concave, not glaucous. Sorus (if present) at first embedded in a circular cavity which occupies half the width of the lobe, edges of cavity not raised; sporangia 3-4 (rarely 5), surrounded by brown hairs not longer than the ripe sporangia. Type: Robert Brown, Wales (BM; dupl. at K). Distr. From Annam through Malaysia to Australia, New Caledonia, and New Zealand; in Malaysia: Sumatra, Malay Peninsula, Lingga Arch., Borneo, and Moluccas (Ambon). Ecol. In open places in sandy or thin peaty soil on summits or exposed ridges of mountains at 750-1800 m, especially on sandstone. At Fraser’s Hill, Malay Peninsula, 1250 m, this species occurs in open places on a quartzite ridge, not on the neighbouring granite; it occurs on the summits of some isolated granite mountains (Mt Ophir, G. Belumut). It is abundant from 1000 m upwards in open Leptospermum forest, with Sphagnum, in thin peaty soil on the sandstone mountain G. Jerai or Kedah Peak. Note. For a note on the type specimen of G. circinnata, see note under 4. G. dicarpa. Port Jackson, N.S. 3. Gleichenia vuleanica BL. En. Pl. Jav. (1828) 251; Racis. Fl. Btzg 1 (1898) 10; v. A. v. R. Handb. (1909) 56; BACKER & PosTH. Varenfl. Java (1939) 353.—Calymella vulcanica PRESL, Abh. (K.) B6hm. Ges. Wiss. M.-N. Cl. 5 (1848) 338; CHING, Sunyatsenia 5 (1940) 287; NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 42.—G. dicarpa var. vulcanica CHRIST, Ann. Jard. Bot. Btzg 15 (1898) 75.—G. squamosissima COPEL. Philip. J. Sc. 75 (1941) 348, pl. 1.—Calymella Squamosissima NAKAI, |.c.—Fig. 1d—e, i-n, 3. Similar in habit to G. microphylla, differing as follows: leaflets commonly 4-5 mm apart; seg- ments of lamina less than 1 mm long and wide; lower surface glaucous, deeply concave, with edges much reflexed and the surface adjacent to the costa also swollen; many small thin scales, with slender crisped marginal hairs, all along costules; sporangia 2 or 3, not in a depression in the lamina. At very high altitudes often only un- branched fronds, very densely scaly, are produced. Type: Blume, Java (L). Distr. Malaysia: Sumatra, Malay Peninsula (3 localities), W. Java, Borneo, Philippines (Mindanao, Mindoro), Celebes, and New Guinea. Ecol. Abundant in open scrub (on both dry stony and wet ground) on volcanic mountains throughout W. Java and Sumatra, 1800-3600 m, at these altitudes apparently less abundant on granite mountains in Borneo; densely scaly at highest elevations both in N. Sumatra and in New Guinea (G. sqguamosissima COPEL.). Fig.3. Gleichenia vulcanica BL. in the high mountains of Mt Goh Lembuh, Gajo Lands, N. Sumatra, sheltered in rock crevices, c. 3000 m (1937). 4. Gleichenia dicarpa R.Br. Prod. (1810) 161; C. Cur. Ark. Bot. 9% (1910) 33.—Calymella dicarpa PRESL, Abh. (K.) BOhm. Ges. Wiss. M.-N. Cl. 5 (1848) 338.—Calymella circinnata (non G. circinnata Sw.) CHING, Sunyatsenia 5 (1940) 288. —Fig. 1f-h. Like G. vulcanica, but the /eaflets not copiously scaly (costae almost or quite glabrous when mature), the aperture of the concave lower surface of each segment of the lamina reduced to much less than half of the area enclosed by the outline of the segment as seen from below, owing to much swelling of the costal tissue; sporangia 2, filling the aperture. 10 FLORA MALESIANA [ser. Il, voleaie Type: Robert Brown, Tasmania (BM; dupl. New Guinea, occurring at 750-850 m, in open at K). forest or rocky places. Distr. Australia, New Caledonia, in Malaysia: Note. Though typical specimens of G. vulcanica New Guinea, Philippines (Mindanao). in Java differ markedly from typical specimens of Ecol. On Normanby and Rossel Islands, SE. _ G. dicarpa in Australia, some specimens on moun- ale unnl Fig. 4. a. Diagram of branching in Gleichenia subg. Diplopterygium. — G. longissima BL., 6. Part of rachis-branch bearing two pinnules, x 2/3, c. segment of lamina, lower surface, showing veins and sori, x 4, d. resting apex of main rachis and bases of branches, with stipular leaflets, nat. size. — Scales and hairs. — G. longissima BL., e. A single scale from apex of rachis, x 8, f. edge of scale, x 133, g. stellate hair, x 33.—G. conversav. A. v. R.,h. A single scale, x 8, i. edge of scale, x 133, j. stellate hair 3336 Dec. 1959] Gleicheniaceae (Holttum) 11 tains in Mindanao and New Guinea seem some- what intermediate. Ecological distinctions between the two have not been studied. CHRISTENSEN, in his notes on specimens in the herbarium of Swartz at Stockholm (/.c.) stated that the type specimen of G. circinnata was identical with G. dicarpa R.Br. With his original description SWARTZ gave no precise locality, but in his ‘Synopsis Filicum’ he stated that the species was from Botany Bay, near Sydney. The only specimen labelled G. circinnata in SWARTZ’s herbarium from that locality is certainly not G. dicarpa, though it has only two sporangia on each lamina-segment; the segments are not pouch- shaped in the manner of G. dicarpa. One can only conclude that CHRISTENSEN did not make a careful examination of the specimen. However, the Botany Bay specimen is not certainly the original on which the species G. circinnata was based (furthermore, the original description in- cluded the phrase capsulis quaternis, not true of the Botany Bay specimen). I therefore refrain from reverting to CHRISTENSEN’S earlier identifica- tion of G. microphylla R. BR. with G. circinnata Sw. After the publication of CHRISTENSEN’s note, some botanists used the name G. circinnata to replace G. dicarpa, but it seems clear that this was an error, and I| therefore restore R. BROWN’s name. 2. Subgenus Diplopterygium HoLtt. Reinwardtia 4 (1957) 261.—Gleichenia subg. Mertensia sect. Diplopterygium DiELs in E. & P. Pfl. Fam. 1, 4 (1900) 353.—Gleichenia subg. Mertensia § 1 HooK. Sp. Fil. 1 (1844) 4.—Gleichenia subg. Mertensia sect. Dicranopteris v. A. v. R. Handb. (1909) 57, p.p.—Dicranopteris UND. Bull. Torr. Bot. Cl. 34 (1907) 249, p.p.—Sticherus § Hicriopteris C. CHR. in Verdoorn, Man. Pterid. (1939) 530.— Hicriopteris (non PRESL) CHING, Sunyatsenia 5 (1940) 277; CopeL. Gen. Fil. (1947) 28.—Mesosorus Hassk. Fil. Jav. 1 (1856) 2, p.p.—Diplopterygium NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 47.—Fig. 4-6. Fronds of a young plant producing pairs of bipinnatifid branches, with periodic dormancy of the apex, from an early stage, the lateral branches of such fronds of immature plants small and sterile; periodic dormancy of the apex of the main rachis, and no other dormancy, occurring on fronds of mature plants; lowest leaflets on the branches often deltoid and bipinnatifid (or with deltoid-bipin- natifid lowest branchlets), forming stipule-like structures which protect the dormant apex of the main rachis; remaining leaflets very deeply pinnatifid, segments of the lamina oblong, each with a costa bearing on either side several once-forked veins. Sori on the acroscopic branches of the veins, several to each segment of the lamina. Distr. More than 20 spp. in NE. India, Burma and Indo-China, China, Japan, Malaysia, Polynesia, Hawaii, and tropical America (1 sp.). This subgenus is far more diversified in Malaysia than else- where. Ecol. All species occur on mountains, some descending to only 600 m. All may form dense thickets on edges of forest or other open places; fronds can climb to a height of 6-7 m, if trees suitable for support are present. Note. Some fossils of Cretaceous age in West Greenland (lat. 70-71°N) have exactly the characters of this subgenus (habit of branching, venation, position of sori, nature of sporangia). 5. Gleichenia laevissima CHRisT, Bull. Ac. Inst. Geogr. Bot. III, 11 (1902) 268; v. A. v. R. Handb. (1909) 795; Suppl. (1917) 82.—Hicriopteris laevissima CHING, Sunyatsenia 5 (1940) 280.— Diplopterygium laevissimum NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 52. Scales on rachis-apex 10 mm long, nearly 3 mm wide, acuminate, entire; rest of frond quite glabrous except for very short simple hairs on lower surface of veins and lamina of young fronds and soral hairs. Primary rachis-branches to at least 70 cm long, lowest leaflets not stipule-like; leaflets at an angle of about 45° to rachis, largest 17-20 cm long, 214-3 cm wide, costae 214-314 cm apart; lamina lobed almost to the costa, costules at about 45° to the costa and 314-414 mm apart; segments of lamina 2-2!4 mm wide above the base, narrowed gradually towards the apex; lower surface not glaucous, veins only slightly raised; upper surface not raised along margins of costae, veins raised only near their bases. Sporangia usually 4, pale, with very slender pale hairs. Type: Bodinier 1295, Kouy-yang (P). Distr. China (Kweichow, Chekiang, Yunnan, Fukien), Formosa, and Malaysia: Philippines (Luzon). Ecol. In the mountains of Luzon, c. 2000 m. [ser. Tly volute \ ky \ Nth {i tui 14, a < Z. < n ca] 4 < < a4 e) 4 fL, Fig. 5. Gleichenia longissima BL. on edge of forest, Taiping Hills, Malaya, c. 800 m (HOLTTUM, 1952). Dec. 1959] Gleicheniaceae (Holttum) — 13 6. Gleichenia paleacea (CopeL.) Ho.ttr. Rein- wardtia 4 (1957) 265.—Hicriopteris paleacea CopeEL. Philip. J. Sc. 81 (1952) 3. Scales on main rachis-apex not seen; sca/es on primary branches and costae beneath very copious, light brown, larger ones 4-5 mm long and 1 mm wide, those on costae commonly 2 mm long and more than 0.5 mm wide, thin, fringe fine, close, concolorous; smaller scales narrower with long fine fringe and hair-tips; no persistent stellate hairs On veins except in sori. Primary rachis- branch more than 100 cm long (collector’s note); costae 214 cm apart, at right angles to rachis; largest pinnules 15-16 cm long, -244—3 cm wide, lobed almost to the costa, basal lobes slightly reduced and very close to the rachis; costules c. 4 mm apart, very slightly oblique; /amina- segments thin but firm, hardly narrowed above the base, edges slightly reflexed when dry so that sinuses between segments are 1-2 mm wide; lower surface glaucous, veins distinct and slightly raised; lamina on each side of the costa on upper surface raised as in G. gigantea WALL. and some long hairs persistent in the groove, veins slightly sunk in upper surface of lamina when dry. Sori commonly of 3 sporangia, with numerous pale hairs nearly 1 mm long. Type: Elmer 9902 (Herb. Copel.; dupl. at BM, Keebor is. -P): Distr. Malaysia: Philippines (Negros: Cuernos Mts), once collected. Note. This is near to G. longissima BL., but differs in persistent abundant scaliness. A collec- tion of G. longissima from Canlaon Volcano, Negros (MERRILL 607) shows very scaly young parts of a frond, but the scales are much narrower and the old parts are glabrous. 7. Gleichenia volubilis JUNGH. Java 1 (1853) 592, 664; v. A. v. R. Handb. Suppl. (1917) 83,p.p.; BACKER & PostH. Varenfl. Java (1939) 256.— Mertensia arachnoides Hassk. in Hook. J. Bot. Kew Misc. 7 (1855) 332.—Mesosorus arachnoides Hassk. Fil. Jav. 1 (1856) 6.—G. arachnoides METT. in Mig. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 47; Racis. Fl. Btzg 1 (1898) 11; v. A. v. R. Handb. (1909) 58 (arachnoidea); non G. arachnoidea CUNN. 1844.—G. glauca var. arachnoides C. CHR. Ind. Fil. (1905) 320.—Hicriopteris volubilis CHING, Sunyatsenia 5 (1940) 280.—Diplopterygium volu- bile NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 55. var. volubilis. Scales on main rachis-apex nearly black, to at least 6 mm long and 114 mm wide, gradually narrowed towards apex, edges closely fringed with fine spreading pale hairs (or with a broad, thin translucent border bearing short hairs) ; similar but smaller scales on lower surfaces of branch- tachises, costae and costules (smallest scales not dark), these surfaces also densely covered with a persistent felt of pale stellate hairs each less than 0.5 mm diameter, old costae showing the Taised points of attachment of former scales; stellate hairs on costules and lower surface of veins rusty brown. Primary rachis-branches to at least 150 cm long and 30 cm wide; stipule-like leaflets present and deeply lobed; pinnules 12-16 cm long, 2—3 cm wide, costules 314-4 mm apart, at right angles to costae; /amina-segments firm, slightly narrowed above the base, their edges reflexed when dry, lower surface not (or very slightly?) glaucous, veins strongly raised; upper surface of rachis covered with stellate hairs as lower surface together with very narrow pale ciliate scales; smaller pale scales and hairs at first in groove of upper surface of costae (lamina slightly raised on each side of the groove); veins on upper surface paler than lamina and slightly raised in dried specimens (lamina between veins sometimes slightly raised but not swollen); sporan- gia surrounded by a group of crisped red-brown hairs. Type: Junghuhn s.n., 1839, G. Gedeh (L). Distr. Malaysia: Java, Sumatra, and Central Celebes (one collection). Ecol. Forming dense thickets in open places on ridges, and on edges of forest, often very abundant, 1800-3000 m. Note. The Celebes specimen has smaller pin- nules than normal in Java (10-12 cm long, 114 cm wide), with thinner lamina and veins hardly prominent beneath. var. peninsularis HoLtr. Reinwardtia 4 (1957) 265. Scales on lower surfaces of rachises and cosiae rusty brown like the stellate hairs, not black. Type: F. M. S. Museum s.n., June 1917 (K). Distr. Malaysia: Malay Peninsula (G. Bintang on Kedah-Perak boundary), once collected. 8. Gleichenia bullata Moore, Ind. Fil. (1862) 374; C. Cur. Gard. Bull. S.S. 7 (1934) 212.—G. volubilis (non JUNGH.) v. A. v. R. Handb. Suppl. (1917) 83, p.p.—RHicriopteris bullata CHING, Sunyatsenia 5 (1940) 279. Differs from G. volubilis JUNGH. as follows: scales on rachises, costae and costules much more abundant, those on costules often pale with a dark base; stipular leaflets lacking; primary rachis-branches shorter, pinnules commonly 1.3— 1.5 cm apart, 6-9 cm long, 0.9-1.3 cm wide; veins on upper surface of Jamina (when dry) much sunken, the surface between them swollen. Type: Low, Mt Kinabalu, 7000 ft (K). Distr. Malaysia: Borneo (Sarawak: Mts Dulit and Tibang; N. Borneo: Mt Kinabalu), New Guinea (Mt Dayman). Ecol. Forming thickets 1700-3000 m. Note. The bullate character of the upper surface of the lamina, and the scaliness, distinguish this from dwarfed specimens of G. volubilis which may have pinnules no larger than those normal in G. bullata. like G. volubilis, 9. Gleichenia longissima BL. En. Pl. Jav. (1828) 250; Racis. Fl. Btzg 1 (1898) 10; BACKER & 14 FLORA MALESIANA [ser. rhe vol. |! PostH. Varenfl. Java (1939) 256; HoLtrT. Rev. FI. Mal. 2 (1955) 67.—G. excelsa J. SM. ex HOOK. Sp. Fil. 1 (1844) 5, t. 4B.—G. glauca v. A. v. R. Handb. (1908) 58 et al. p.p.—Hicriopteris longis- sima CHING, Sunyatsenia 5 (1940) 280.—Diplop- terygium longissimum NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 53.—Fig. 4b-g, 5. Scales on resting apex of rachis as in G. volubilis JUNGH.; young expanding parts of fronds covered with loose indumentum of narrow rust-coloured scales and lax stellate hairs, a few such scaies and hairs sometimes persistent on lower surface of costae and costules, mature rachises quite smooth and glabrous. Primary rachis-branches to about 200 cm long and 40 cm wide; largest pin- nules 15-22 cm long, 2144-314 cm wide, costae 2.8-3'4 cm apart; lamina thin, lobed almost to costa, costules 4-5 mm apart, slightly oblique to costa; segments of lamina slightly narrowed above the base, edges slightly reflexed when dry; lower surface glaucous, veins slender and slightly raised; upper surface of rachis and costae glabrous, lamina slightly raised on each side of costae; veins on upper surface slightly raised near base, rarely also distally, very slender; sori usually of 3 sporangia, surrounded by crisped brown hairs. Type: Blume, Java (L; dupl. at K, P). Distr. Southern China to Indo-China through Malaysia to Melanesia (Fiji, Tahiti), in Malaysia: not collected in the Lesser Sunda Islands E of Flores, Celebes, the Moluccas, and New Guinea. Ecol. Forming thickets in clearings and on edges of forest, more or less persisting in open secondary forest; 750-1800 m. RACIBORSKI states that this species may climb very high on trees. Note. This is the most widely distributed species in the subgenus. In Java and Sumatra it is replaced at higher altitudes by G. volubilis; a comparison of the two species near the transition zone needs to be made. 10. Gleichenia clemensiae (CoPEL.) HoLtTtT. comb. nov.—Hicriopteris clemensiae CopEL. Un. Cal. Publ. Bot. 18 (1942) 217.—G. papuana Hovtt. Reinwardtia 4 (1957) 266. var. clemensiae. Scales on resting apex of main rachis dark brown, shining, 4 mm long, more than 1 mm wide, edges bearing spreading hairs to 0.5 mm long; lower surface of rachises and costae bearing numerous persistent scales like those on the resting apex but smaller, dark and shining with pale edges and fringe, those on the costae commonly 1144 mm long and 14 mm wide, old costae minutely warty from the raised bases of former scales; scales on lower surface of costules abundant, very small, laxly fringed; no hairs on veins, apart from sori; upper surface of rachis rather persistently covered with very narrow scales and long lax hairs, upper surface of costae hairy near the base only. Rachis- branches 120 cm long; pinnules spreading at right angles, 2!4 cm apart, largest 12 cm long, 2 cm wide; costules 2!4-3 mm apart; lamina thin, veins slightly prominent on lower surface, not on upper; soricommonly of 3 sporangia, with crisped paraphyses. Type: Clemens 41227, Morobe, Matap, 5000- 6000 ft (H. Copel.). Distr. Malaysia: SE. New Guinea (Morobe, and Milne Bay Distr.: Mt Dayman). Ecol. Scrambling to 2-3 m, forming dense tangles on edges of mossy forest, 1600-2250 m. var. membranacea (HOLTT.) comb. nov.—G. papuana var. membranacea HOLTT. Reinwardtia 4 (1957) 266. Scales on rachis-apex pale and very thin near the edges which are very shortly fringed; scales on rachises and costae more abundant, dark at the base only. Type: Brass 24763 (BM). Distr. Malaysia: New Guinea (Goodenough I.). Ecol. Plentiful in openings in forest, 1600 m. 11. Gleichenia angustiloba HoLtr. Reinwardtia 4 (1957) 263. Rhizome 7 mm diam., young parts covered with shining dark brown scales 5-7 by 144 mm, old parts warty from bases of former scales; stipes 6-7 mm diam., the base at first scaly like the rhizome and then warty, upper part covered with small scales like those of the rachises, later asperulous. Primary rachis-branches 100 cm long or more; stipular leaflets broadly deltoid, to 4 cm long, pinnatifid, lowest lobes deeply and narrowly lobed again; scales on apex of main rachis dark brown, shining, narrow, 3-4 mm long, the edges bearing concolorous oblique rigid setae; rachises and costae, and lower surfaces of costules more or less persistently covered with smaller scales of various sizes and red-brown stellate hairs. Pin- nules to 15 cm long and 3-314 cm wide, 2—3'4 cm apart, the distal ones distinctly deflexed, the lower ones at right angles to rachis; all segments of the lamina c. 2% mm wide, separated by wide sinuses, many of the lower segments being constricted at the base and joined laterally by a very narrow wing along the costa (wing about 0.2 mm wide); lowest segments forming distinct separate leaflets; costules 4 mm apart; veins prominent on both surfaces; sori of 4 sporangia with long red-brown crisped hairs, no other hairs on the veins of mature fronds. Type: Brass 4960, Mt Tafa (BM, dupl. at K, Bo, US). Distr. Malaysia: E. New Guinea (Mt Tafa, and on Asaro-Mairi Divide, Goroka Subdistr.). Ecol. Altitude 2400 m; ‘Common in native rest clearings in ridge-crest forest; conspicuous, rambling amongst forest fringe bushes and small trees; fronds 4-5 m with 6-8 or more pairs of large spreading pinnae”’ (BRAss). 12. Gleichenia elmeri CopeL. in Elmer, Leaf. Philip. Bot. 3 (1910) 799.—Diplopterygium elmeri NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 49. Scales on rachis-apex not seen; scales on costae sparse; scales on costules at first abundant, Dec. 1959] Gleicheniaceae (Holttum) = a 15 longest 1-1'14 mm long, narrow, rusty brown with well-spaced rigid spreading marginal hairs; very small scales and stellate hairs present on veins and with sporangia, hair-branches stiff, c. 0.2 mm long; dark brown short clavate hairs abundant on costules and less so on sides of costae. Pinnae 150 cm long (fide COPELAND); largest pinnules 25 cm long, 344-4 cm wide, 4144-5 cm apart; costules 514-7 mm apart, at right angles to costa; several segments of lamina constricted at base, widely separated and joined by a wing 14 mm wide along the costa; lower surface pale green, veins slightly raised; upper surface of lamina along sides of costae wrinkled when dry, probably somewhat raised in living fronds; veins not raised on upper surface except near base; sporangia commonly 3-5, less often 6 or 7, not always completely covering lower surface (as reported in original description); numerous pale hairs, longer than sporangia, attached to the receptacle. Type: Elmer 11423 (H. Copel.; dupl. at K, WS2eb, LP): Distr. Malaysia: Philippines (Mindanao: Mt Apo), once collected. 13. Gleichenia deflexa HoLTT. Reinwardtia 4 (1957) addendum p. 280. Scales on apex of main rachis light brown with darker obliquely setose edges; branch rachises and lower parts of costae at first covered on both surfaces with a close felt of small setose scales and stellate hairs and also with numerous narrow scales (those on rachis 5 mm long, 0.2 mm wide), setae of scales and hairs of upper surface all rigid and rather dark, of smaller scales and hairs on lower surface pale and lax; lower surface of costules scaly and hairy near their bases like the costae; lower surface of veins bearing persistent scattered stellate hairs with slender pale rays. Rachis-branches 120 cm or more long, fertile ones to 35 cm wide, sterile sometimes only 18 cm wide, all pinnules distinctly deflexed, making angles of about 75° with the rachis; costae on large fertile rachis-branches 414 cm apart, on smaller sterile branches 214-314 cm apart; fertile pinnules 16-20 cm long, widest part 3!4-3.8 cm wide, costules 4-5 mm apart; /amina thin, veins raised near their bases on both surfaces when dry; 20 or more pairs of segments on each larger pin- nule constricted at the base, connected by a very narrow wing along the costa, c. 314 mm wide above the base, lowest segment sometimes quite free and shortly stalked; sori of 3-5 sporangia; stipular leaflets to 4 cm long, bipinnatifid, segments c. 1 mm wide, acuminate. Type: Brass 27171, Ferguson Isl. (L). Distr. Malaysia: SE. New Guinea (Normanby and Fergusson Islands). Ecol. Scrambling to 7 m high in rather open forest, 800-850 m. 14. Gleichenia sordida CopeL. in Elmer, Leafl. Philip. Bot. 3 (1910) 798.—G. novoguineensis BrRAusE, Bot. Jahrb. 56 (1920) 210.—Hicriopteris novoguineensis COPEL. Philip. J. Sc. 75 (1941) 358. —Hicriopteris astrotricha Cope. /.c.—Diplo- pterygium novoguineense NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 54.—G. sumatrana Ho.tr. Reinwardtia 4 (1957) 264. Scales on dormant apex of rachis 8-10 mm long, 0.5 mm wide, light brown with dark edges bearing oblique setae; rachises and costae more or less persistently covered with a felt of stellate hairs (all dark brown or some rusty brown) and some narrow scales to 4 mm long; lower surfaces of costules and veins more or less persistently covered with stellate hairs. Rachis-branches 120-200 cm long, costae 214-414 cm apart; largest pinnules 15-20 cm long, 2!4-3 cm wide, slightly deflexed or not; costules 314-414 mm apart; 2-8 basal segments of lamina on each pinnule constricted at the base and connected by a very narrow wing along the costa, rest separated by narrow sinuses; lamina rather thin, not glaucous beneath; veins raised on lower surface, sometimes not on upper surface; sori of 3-5 sporangia; stipular leaflets broadly deltoid, 214-414 cm long, their lobes c. 1 mm wide. Type: Elmer 11423a, Mt Apo, Mindanao (H. Copel.; dupl. at US, BM). Distr. Malaysia: Sumatra, Malaya, Celebes, Moluccas (Halmaheira, Batjan, Morotai), Philip- pines (Mindanao), New Guinea, and Solomons. Ecol. High-climbing in open forest and on forest-edge, apparently not forming thickets, 1000-2000 m. 15. Gleichenia norrisii METT. in Kuhn, Linnaea 36 (1869) 165; v. A. v. R. Handb. (1908) 58; C. Cur. Gard. Bull. S.S. 7 (1934) 212, incl. var. floccigera C. CHR.; Hottrt. Rev. Fl. Mal. 2 (1955) 67.—Hicriopteris norrisii CHING, Sunyatsenia 5 (1940) 280.—Diplopterygium norrisii NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 54.—Fig. 6. Scales on apex of main rachis 2-3 mm long, narrow, medium brown, edges bearing short oblique concolorous setae; costae and costules when mature quite glabrous or bearing a few scales near the base; young rachises, costae and costules bearing scattered very narrow brown scales, small setose scales and stellate hairs with rusty brown rays; stellate hairs at first present on veins but not persistent. Rachis-branches 100 cm or more long; largest pinnules to 20 cm long and 3 cm wide, the costae 4-5 cm apart, distinctly deflexed, distal pinnules more acutely deflexed than basal ones; lobes of lowest pinnules usually only slightly enlarged and not stipuliform, but stipule-like leaflets, with broad blunt lobes, sometimes produced; costules 414-514 mm apart; lamina light green, drying light olive green without reddish tinge, thin, lower surface more or less glaucous in young plants, often not appreciably so in older ones; segments of lamina only slightly narrowed above the base, separated by sinuses not over 1 mm wide, apices broadly rounded, lowest segments much reduced but not separated from the rest; veins slender, distinctly raised on lower surface, slightly on upper surface; sori FLORA MALESIANA [ser. Ui, -volaaie Fig. 6. Gleichenia norrisii METT. on edge of forest, Penang, c. 650 m; one pair of rachis-branches showing widely spaced and deflexed pinnules (HOLTTUM, 1926). Dec. 1959] Gleicheniaceae (Holttum) Sed : bz commonly of 3-5 sporangia, with paraphyses consisting of small scales bearing long, pale, marginal hairs. Type: Norris, Malay Peninsula (B?; dupl. at K, BM, P). Distr. Malaysia: Malay Peninsula, Sumatra (Bencoolen), and N. Borneo. Ecol. In clearings and on edge of forest, 650— 1250 m. On Penang Hill this species is abundant at 650-750 m, below the lower limit of altitude of G. longissima; above 750 m the latter, which forms denser thickets because of its closer pinnules, is the common species. Note. CHRISTENSEN’S var. floccigera was based on immature fronds which had not lost their indumentum. The development of stipule-like leaflets varies, but they never have the narrow caudate lobes found in most other species. 16. Gleichenia matthewii HoLtr. Reinwardtia 4 (1957) 265. Scales of main rachis-apex hardly 14 mm wide, edges with their setae shining dark brown, middle part paler; costae bearing a few scales near the base, the frond otherwise glabrous. Rachis- branches 70 cm or more long, strongly flexuous above the basal part; all pinnules deflexed, the upper ones at an angle to 45° so that each is in line with the portion of the rachis beyond it, the uppermost grading into the spreading lobes of the terminal lamina; largest pinnules 12 cm long and 2'4 cm wide, the costae 4 cm apart; costules 414 mm apart; /amina firm, glaucous beneath, lobed to 1-114 mm from the costa, segments oblong, contiguous, almost truncate at the apex and often slightly retuse; no sori seen; stipular leaflets deltoid, 2 cm long, narrowly lobed. Type: Matthew s.n., 31 Jan. 1912, G. Sing- galang (K). Distr. Malaysia: Central Sumatra (Mt Sing- galang), twice collected, altitude 1800 m. 17. Gleichenia brevipinnula HoL_tr. Reinwardtia 4 (1957) 264. Scales on apex of rachis c. 5 mm long, hardly 4 mm wide, brown, with almost black shining edges bearing oblique setae; rachises rather persistently covered with very dark rigid stellate hairs with a few narrow scales; scattered similar hairs on costae and costules beneath, and scattered somewhat paler stellate hairs on lower surface of veins. Main rachis 5 mm diameter; branches 70 cm long; costae c. 2 cm apart, pinnules to 10 cm long and 2 cm wide, not or slightly deflexed; costules 3-314 mm apart; /amina cut down to 1 mm from the costa, very firm, sinuses very narrow, edges of segments slightly toothed near apices at the ends of the veins, lower surface strongly glaucous; veins raised on both surfaces; stipular leaflets 31 cm long, broadly deltoid, all lobes with very narrow caudate tips, lower lobes deeply and very obliquely lobed again; segments of sub-basal pinnules often with caudate tips like the stipular leaflets; sori of 3 or 4 sporangia. Type: Bell 2042, Pueh Range, Sarawak (BM). Distr. Malaysia: Borneo (Sarawak & N.Borneo). Ecol. On ridges and summits of mountains, in dwarf forest or scrub, 1250-2500 m. Fronds are reported up to 214 m long. 18. Gleichenia blotiana C. Cur. Bull. Mus. Hist. Nat. Paris Il, 6 (1934) 103.—Hicriopteris blotiana CHING, Sunyatsenia 5 (1940) 279.—Diploptery- gium blotianum NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 49. Scales on dormant apex of rachis nearly black, edges bearing oblique black setae; scales on growing main rachis usually with a paler median band; rachis-branches when young bearing many dark stellate hairs with rigid rays (so close that hairs touch each other but do not completely obscure surface of rachis) and also very narrow ciliate scales 2 mm long and some long lax multi- cellular hairs; a similar indumentum on upper surface of costae at first, later deciduous; in- dumentum on lower surface of costae and costules mostly of dark rigid stellate hairs, with a few scales at bases of costae; dark stellate hairs scattered abundantly on veins and lamina beneath, persistent. Main rachis-branches c. 150 cm long, the lowest pinnules stipule-like, 314 cm long, deltoid, with very narrow deeply pinnatifid basal lobes; pinnules commonly 20 cm long, 3-4 cm wide, the base of the costa distinctly deflexed as in G. norrisii; costae 4-5 cm apart, costules c. 5 mm apart; segments of lamina 3!4-4 mm wide, rather thin, their apices broadly rounded and sometimes retuse; veins distinctly raised on upper surface and slightly so below; sporangia 3-4 in each sorus. Type: Pételot 3900, near Chapa, Indo-China (BM; dupl. at P). Distr. Indo-China and S. China, in Malaysia: Malay Peninsula, once collected. Ecol. In the Malay Peninsula on edge of forest in a thicket with G. Jongissima, at 1500 m. 19. Gleichenia conversa v. A. v. R. Bull. Jard. Bot. Btzg II, ». 20 (1915) 17; Handb. Suppl. (1917) 81; BACKER & Postu. Varenfl. Java (1939) 255.—Diplopterygium conversum NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 49.—Fig. 4h-j. Differs from G. norrisii as follows: scales on rachis-apex brown with almost black edges; some rather light brown stellate hairs often persistent on costules and sides of costae; largest pinnules to 25 cm long and 4 cm wide; distal pinnules not appreciably deflexed; segments of lamina separated by sinuses about 2 mm wide; lamina and rachises drying rather red-brown, lamina usually thicker than in G. norrisii; veins usually not at all raised on upper surface (slightly so in specimens with thin lamina); stipular leaflets well developed, the lobes narrow. Type: Hasskarl, Kandang Badak, G. Gedeh (Bo; dupl. at L). Distr. South Malaysia: Java and Lesser Sunda Islands (Flores). Ecol. In open places in forest and on forest- edges, 900—2400 m. 18 FLORA MALESIANA [ser. II, vol. 1? 3. Subgenus Mertensia Hook. Sp. Fil. 1 (1844) 6, pro § 2; Hottr. Reinwardtia 4 (1957) 266.—Mertensia WILLD. Kongl. Vet. Ak. Nya Handl. 25 (1804) 163, p.p., non RotH 1797.— Gleichenia subg. Mertensia sect. Holopterygium Dies in E. & P. Pfl. Fam. 1, 4 (1900) 353.—Dicranopteris (non BERNH.) UND. Bull. Torr. Bot. Cl. 34 (1907) 249, p.p.—-Gleichenia subg. Mertensia sect. Dicranopteris v. A. v. R. Handb. (1908) 56, p.p.—Sticherus PRESL, Tent. Pterid. (1836) 51; CHING, Sunyatsenia 5 (1940) 281; CopeL. Gen. Fil. (1947) 27; NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 7.— Sticherus § Eu-Sticherus C. Cur. in Verdoorn, Man. Pterid. (1938) 530.— Mesosorus Hassk. Fil. Jav. 1 (1856) 2, p.p.—F ig. 7-10. Primary rachis-branches each ending in a dormant apex which lies in the angle between a pair of secondary branches; secondary branches behaving similarly, the process usually repeated to produce pseudo-dichotomous branching of several orders; ultimate branches simply deeply pinnatifid, the venation of segments of the lamina and sori as in subg. Diplopterygium; penultimate branches (often also + TTT Fig. 7. Diagrams showing branching habit of species of Gleichenia subg. Mertensia. a. G. truncata (WILLD.) Spr., b. G. milnei BAK., c. G. hispida METT., d. G. hirta BL., e. G. vestita BL. Bect959] or) Gleicheniaceae (Holttum) 19 branches of lower orders) more or less completely provided with a deeply pinnatifid lamina like that of the ultimate branches. Notes. This subgenus includes far more species than any other major division of the family, in all continents, and its distribution is mainly south of the equator. Most of the species are rather small in size, as compared with representatives of subg. Diplopterygium, and could not compete as thicket- forming ferns either with the latter or with Dicranopteris. The only Malaysian species which can so compete is G. truncata, and it is also the only species of subg. Mertensia which occurs at sea-level in our region. The others are mountain plants occurring in scrub and dwarf forest of high ridges or in open grassy or rocky places, which are comparable with the habitats of related species in south temperate regions. As in other divisions of the family, the characters and distribution of scales and hairs are important diagnostically in this subgenus. The number of times the lateral branch-systems are forked (the number of orders of forking) is probably important, but shows considerable variation within a species ac- cording to the age of the plant and to environmental conditions, and these cannot be fully judged from dried specimens. A more important kind of character is the relative length of branches of the first and ultimate orders. But in some species one frond will have branch-systems forked to 2 orders with long ultimate branches, other fronds (even part of the same frond) may be forked to 3 orders with much shorter ultimate branches. The glaucous character of the lower surface of the lamina may be significant, but is easily destroyed by heat in drying. In all these characters, and in all species, more field study is needed to establish distinctions between species more clearly. Judging from anatomical characters, CHRYSLER (Am. J. Bot. 31, 1944, 483-491) has argued that subg. (eer ‘oF Fig. 8. Gleichenia truncata (WILLD.) Spr. a. Stipular leaflets at main branching, nat. size, b. lower surface of part of frond, x 4. — G. vestita BL., c. Lower surface of a large segment of lamina, x 4, d. a single scale from costa, x 8. — G. hispida METT., e. Lower surface of part of frond, showing sori and hair-pointed scales, x 8, f. same with scales and sporangia removed, g. a single scale from costa, xX 22. -G. hirta BL. h. Lower surface of part of frond, x 4, i. tip of a segment of the lamina, showing toothed edge, j. a scale from the costa, x 20. 20 FLORA MALESIANA [ser. IL -yvolaat Mertensia is the most primitive division of the genus. But from the point of view of sporangia and sori it is exactly in equality with subg. Diplopterygium, and in leaf-form it is clearly more highly or- ganized (in having a branching-pattern dependent on a series of permanently dormant apices). It seems to me significant that among fossils none are clearly referable to this subgenus, whereas there are abun- dant fossils having a close resemblance to subg. Diplopterygium. An apparent exception is Gleichenites gracilis ZIGNO. But the branching-pattern of this fossil is very irregular and there is no clear evidence of pseudo-dichotomy; I think that its resemblance to subg. Mertensia is superficial. 20. Gleichenia reflexipinnula C. Cur. Brittonia 2 (1937) 27.—Sticherus reflexipinnula CoPEL. Philip. Ie SeseaCl94)) 355% Rhizome 5 mm diameter, covered when young with dark brown shining acuminate scales hardly 1 mm wide at the base, when old closely warty; stipe 30 cm or more long, scaly and then warty like the rhizome near the base; main rachis bearing 2 or more pairs of primary branches, with deeply pinnatifid stipular leaflets 2-4 cm long at bases of branches; primary branches once or _ twice forked, leafy down to the junction with the main rachis, not proliferous beyond the first fork; angle of ultimate forks rather less than a right angle; ultimate branches to c. 20 cm long and 314-4 cm wide (shorter if primary branch is twice forked), penultimate branches 314-9 cm long; scales on resting apex of main rachis 314 mm long, less than 1 mm wide, acuminate but not hair- pointed, rather thin, medium brown with short sparse fringe of paler hairs; scales on main rachis persistent, abundant, thin, pale and long-fringed; scales on lower surface of costae abundant, largest 2 mm long, 14 mm wide, thin, pale, long- fringed; scales on costules similar and smaller, abundant, ranging down to stellate hairs; no stellate hairs on veins; /Jamina lobed almost to the costa, middle and upper segments slightly deflexed, lower ones much deflexed, costules 3144 mm apart; segments gradually narrowed to apex (which is sometimes retuse), coriaceous, edges strongly revolute when dry, veins hardly raised on either surface; sporangia usually 4, sori close together. Type: Brass 4847, Mt Tafa (BM; dupl. at Bo, US). Distr. Malaysia: E. New Guinea (Mt Tafa, Mt Dayman). Ecol. Forest-edge, subscandent to 2 m (Mt Dayman), plentiful in summit clearing (Mt Tafa), 2230-2700 m. 21. Gleichenia venosa (CoPEL.) HOLTT., comb. nov. —Sticherus venosus CoPEL. Philip. J. Sc. 75 (1941) 3565 ple 5. Habit of G. truncata and of similar size (rachis- branches of second order sometimes only partly leafy); segments of lamina thicker, with edges sometimes much reflexed when dried, the veins on the lower surface strongly raised; scales of apex of main rachis similar to those of G. truncata; scales on costae abundant and persistent, dark rusty-brown, to 1144 mm long and more than 4 mm wide, fringed; scales on costules similar but smaller, very abundant, spreading and closely overlapping so as completely to cover lower surface of lamina; smaller scales also present throughout on rachises, persistent as a continuous rusty covering at and just below the forks. Type: Brass 12348, Idenburg River (H. Copel.; dupl. at Bo, BM, L). Distr. Malaysia: New Guinea. Ecol. Thickets in open places, scrambling to 2-3 m, 1600-2700 m. 22. Gleichenia truncata (WILLD.) Spr. Syst. Veg. ed. 16, 4 (1827) 25; HoLtr. Reinwardtia 4 (1957) 271.—See for further synonyms under the varieties. —Fig. 7a, 8a—b, 9. KEY TO THE VARTERIESs 1. Additional stipular leaflets present about 1 cm below main and first lateral forks of rachis, as well as just above forks 2. var. bracteata 1. Such additional stipular leaflets lacking. 2. Costae persistently densely scaly on the upper Giighes 5 So 6 5 5 4. var. involuta 2. Costae at most sparingly and not persistently scaly on the upper surface. 3. Main rachis-branches several times forked. Scales on costae beneath very small or nar- row and acuminate. 4. Scales on costae sparse and very small. 1. var. truncata 4. Scales on costae 114-2 mm long, 0.3-0.6 mm wide, acuminate 5. var. celebica 3. Main rachis-branches 2 or 3 times forked. Scales on costae beneath rather abundant, 1-114 mm long, 0.7—1 mm wide. 3. var. plumaeformis 1. var. truncata.—Mertensia truncata WILLD. Kongl. Vet. Ak. Nya Handl. 25 (1804) 169, t. V, f. A.—Mertensia laevigata WILLD. Sp. Pl. 5 (1810) 75.—Sticherus laevigatus PRESL, Tent. Pterid. (1836) 52.—G. laevigata Hook. Sp. Fil. 1 (1844) 10; Racts. Fl. Btzg 1 (1898) 11; v. A. v. R. Handb. (1908) 59; BAcKER & PostH. Varenfi. Java (1939) 255; Hoxtr. Rev. Fl. Mal. 2 (1955) 71.—Sticherus truncatus NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 20.—Sticherus myriapoda NAKAI Zc) 125 f 12 Main rachis 8 mm or more diameter near the base, often high-climbing with many pairs of branches; primary rachis-branches several times almost equally forked, the angle of forking about a right angle, of ultimate forks sometimes more than 90°; basal segment of primary rachis-branch leafless (apart from stipular leaflets), branches of second and later orders usually all leafy, length of each branch from one fork to the next commonly 10—15(—20) cm; stipular leaflets usually present at base of primary branch and at its first fork, Dec. 1959] a4 TS y sae ie ’ Gleicheniaceae (Holttum) 21 Fig. 9. Gleichenia truncata (WILLD.) SPR. on Taiping Hills, Malaya, by roadside, c. 1200 m (HOLTTUM, 1952). deltoid and deeply lobed (basal lobes again lobed), at base of primary rachis-branch to 3 cm long, at next fork 114 cm long; lamina cut almost to the costa into lobes at right angles to the costa; lobes rather irregular in length even on same branch, longest on lowest branches, commonly 2-314 cm long (to 5 cm) and about 2!4 mm wide (to 3 mm), costules 3-4 mm apart; lower surface not glaucous, veins dark when dry, slightly raised; veins on upper surface not raised when dry, concolorous; sori of 3-5 sporangia, surrounded by pale hairs. Scales on smaller dormant apices mostly very small, often not more than 1 mm long, rusty brown, of varying shape, fringed; on larger apices also some very narrow scales; sca- les on costae very small, long-fringed, at first rather abundant; very short simple hairs, often shrivelled and black when old, present on lower surface of costules, veins and lamina. Type: Herb. Willd. (B). Distr. Throughout Malaysia, apparently most abundant in the west. Ecol. On edges of forest, often climbing to a considerable height, 0-1600 m, growing with varieties of Dicranopteris linearis, also in thickets at higher altitudes. Note. The original specimen of Mertensia truncata WILLD. had incompletely expanded lamina-lobes, which thus appeared to be truncate. Sticherus myriapoda NAKAI was described from a plant in the Riouw Archipelago differing only from typical var. truncata in having a twining main rachis. Such a condition has also been observed in one plant in Johore, but is not general in this variety. Experimental culture is desirable to decide whether the twining condition is genetic- ally controlled. The Johore plant had rachises twining both to the left and to right, and the length of rachis making one complete turn round the support was about 20 cm. NAKAI described a closer spiral. 2. var. bracteata (BL. ex Hook. & BAKER) Ho.tr. Reinwardtia 4 (1957) 271.—G. bracteata BL. ex Hoox. & BAKER, Syn. Fil. (1865) 14 (as synonym but with descr.)—G. /aevigata var. bracteata v. A. v. R. Handb. Suppl. (1917) 85. Lamina-segments commonly less than 2 mm wide, 10-18 mm long; additional stipular leafiets present about 1 cm below main and first lateral forks of rachis, on the outside. Type: Blume, Java (L; dupl. at K). Distr. Malaysia: Java, Flores, S. Celebes. Ecol. On edges of forest, apparently abundant at 1000-1600 m; extremes of altitude recorded 600 and 2000 m. 3. var. plumaeformis (PRESL) HOoLTr. Rein- wardtia 4 (1957) 272.—Mertensia plumaeformis PresL, Abh. (K.) B6hm. Ges. Wiss. M.-N. Cl. V, 22 FLORA MALESIANA 5 (1848) 338; Epim. Bot. (1851) 24, t. 15.— Sticherus plumaeformis NAKAI, Bull. Nat. Soc. Mus. Tokyo n. 29 (1950) 25. Primary rachis-branches once or twice forked, 3-6 cm to first fork ‘this part almost or quite leafless), ultimate branches up to 25-35 cm long; costae usually bearing rather numerous, thin, broad, almost entire scales 1-144 mm long, 0.7-1 mm wide, on lower surface. Type: Cuming 377, Malacca (PRc; dupl. at K, BM;)P) EE): Distr. Malaysia: Sumatra, Malay Peninsula, Borneo (?). Ecol. In open places on mountains, 1200-1800 m. As seen by me in W. Sumatra this variety does not form long climbing fronds. 4. var. involuta HoLTT. Reinwardtia 4 (1957) 272. Primary rachis-branches 3 times forked; ultimate branches sometimes much longer than lower ones; ultimate forks forming an angle of rather less than 90°; upper surfaces of costae densely scaly, scales persistent, 1 mm long, pale-fringed; lower surfaces of costae bearing scales ¥% mm long; edges of dried Jamina-lobes much revolute. Type: J. Winkler; Rosenst. Fil. Sum. Exsic. 182 (L; dupl. at BM, P). Distr. Malaysia: N. Sumatra (Karo Plateau). 5. var. celebica HOLTT., var. nov. Primary rachis-branches 3 times equally forked, ultimate and penultimate branches leafy, angle of ultimate forks less than a right angle, segments of lamina c. 2 cm long; sca/es on lower surface of costae copious, acuminate, fringed, c. 114-2 mm long, 0.3-0.6 mm wide. Distr. Malaysia: Central 2600-3000 m (once collected). Celebes, altitude 23. Gleichenia milnei BAKER, Syn. Fil. (1874) 449; Hook. Ic. Pl. t. 1602; HoL_tr. Reinwardtia 4 (1957) 270.—G. kajewskii Core. Philip. J. Sc. 60 (1936) 102, pl. 6.—Sticherus milnei CHING, Sunyatsenia 5 (1940) 284.—Sticherus kajewskii CopEL. Gen. Fil. (1947) 27.—Fig. 7b. General aspect of frond as in G. truncata (in its wide forkings and narrow lamina-segments at right angles to the costa), but at each fork (apart from forks of main rachis) one branch larger than the other, larger branches alternately to left and right and successive larger branches almost in line with each other; ultimate branches, and branches of 2—3 lower orders leafy; leafy branches in general 214-314 cm wide (lower ones to 414 cm), lamina-segments 2-3 mm wide, costules 3-5 mm apart; scales as in G. truncata; additional stipular leaflets often present outside the lower forks, as in G. truncata var. bracteata but nearer the forks. Type: Milne 341, New Hebrides (K). Distr. Malaysia: Celebes,? Philippines (?Luzon), Moluccas, New Guinea, Admiralty Islands, Solomon Islands, New Hebrides. Ecol. Altitude 100-1400 m. [ser. II, vol. 11 24. Gleichenia oceanica KUHN, Verh. K. K. Zool.- Bot. Ges. Wien 19 (1869) 583; non CuHRistT, Ann. Jard. Bot. Btzg 15 (1898) 76; v. A. v. R. Handb. (1908) 60; Suppl. (1917) 85, 497; CopeL. Univ. Cal. Publ. Bot. 12 (1931) 388.—Sticherus oceanicus St JOHN, Occ. Pap. Bish. Mus. 17 (1942) 81. Habit and scaliness of G. truncata, but the forkings of the rachis-branches much less than a right angle (ultimate forks often less than 45°), the lamina glaucous beneath, its segments distinct- ly oblique and commonly less than 2 cm long (often not more than 114 cm); primary rachis- branches of large fronds 5 times forked, with deltoid stipular leaflets at their base, of smaller fronds sometimes only 3 times forked and lacking stipular leaflets. Type: Herus 66, Aneityum, New Hebrides (B). Distr. Melanesia (New Hebrides, Fiji, Samoa), may occur in East Malaysia. Note. The SARASIN specimens from Celebes referred to this species by CHRisT (/.c.) are G. hispida and G. hirta var. paleacea; 1 have not seen the WARBURG specimen. 25. Gleichenia erecta C. Cur. Brittonia 2 (1937) 269; HoLtr. Reinwardtia 4 (1957) 268.—Stricherus erectus COPEL. Philip. J. Sc. 75 (1941) 353.— Sticherus habbemensis Cope. l.c. 355, pl. 3. Rhizome 2!4-4 mm diameter, young parts covered with thin brown ciliate scales to 3 mm long and | mm wide, old parts closely warty; stipe at first scaly, then warty, near the base, like the rhizome, rest smooth, 12-40 cm long; frond bearing one or more pairs of primary branches which are simple or once or twice forked, sometimes proliferous beyond the first pair of secondary branches; if twice forked, the first-order branch 2-3 cm long, bearing a stipular lobed leaflet 15 mm long and sometimes | or 2 other lamina-segments but not fully leafy, the second-order branch 3-6 cm long, fully leafy; angle of ultimate forks less than 45°; ultimate branches 12-30 cm _ long, largest /amina-segments commonly 15-23 mm long, sometimes to 40 mm, rather coriaceous, entire, edges much reflexed when dry, abruptly narrowed just above the base, the rest 2—-2'4 mm wide when flattened, tapering to apex, lower surface glaucous; veins not raised on either surface; costules 3-4 mm apart (lowest ones on large fronds to 5 mm apart), basal part at 60—70° to the costa, distal part often faleate when dry; scales on resting apices of main rachis like those on base of stipe but more finely fringed; some persistent scales on lower surface of costae, 1! mm long, 14 mm wide, shortly hair-pointed, edges fringed at least towards the apex (these scales sometimes very pale with a darker base); scales on costules very small and long-fringed; pale- rayed stellate hairs on veins, their red-brown bases sometimes persistent; sori of 3-4 sporangia. Type: Brass 4591, Murray Pass, Wharton Range, Papua (BM; dupl. at K, Bo, US). Distr. Malaysia: New Guinea. Ecol. In open places, in peaty grassland, and on wet clay-soil of land-slips, 1000-3225 m. Dec. 1959] 26. Gleichenia pulchra (CopeL.) Ho.ttr. Rein- wardtia 4 (1957) 271.—Sticherus pulcher COPEL. Philips. Sc: 75° (1941).355, pl. 4. Main rachis slender, bearing several pairs of branches, each primary branch bearing up to at least 3 pairs of secondary branches, each secondary branch simple or once forked; only ultimate branches leafy, except sometimes where a second- ary branch is forked, in which case the penul- timate branch may bear a few lamina-lobes; ultimate branches to 20 cm long and 2!4 cm wide (to 3 cm wide if a simple secondary branch), lamina lobed almost to the costa, segments very slightly oblique, 3 mm wide above the base, costules 4 mm apart; veins slightly raised on lower surface, on upper surface only at the base of each vein; lower surface glaucous when living (not clearly so on dried specimens available); scales on main dormant apices small, on lower surface of costae 14-1 mm long, firm, dark brown with a pale fringe; scales on lower surface of costules abundant, small, long-fringed; many small rusty- brown stellate hairs on lower surface of veins and on lamina. Type: Brass 12351, Idenburg River (H. Copel.; dupl. at BM, L). Distr. Malaysia: New Guinea (NW. part, once collected, SE. part, doubtful). Ecol. Scrambling in mossy forest, 1650 m. 27. Gleichenia pseudoscandens v. A. v. R. Nova Guinea 14 (1924) 24.—Sticherus pseudoscandens CopEL. Philip. J. Sc. 75 (1941) 356. Fronds to 4 m long, with several pairs of primary branches; primary branch-systems usually with 3 orders of forking, the primary branch sometimes proliferous beyond its first pairs of branches, only the ultimate branches leafy; ulti- mate branches to 40 cm long, lower ones 10—20 cm long; costules 5'4-6 mm apart, at right angles to the costa or nearly so; segments of lamina 17-25 mm long, 3 mm wide above the base, /amina firm, drying dark, veins not raised on either sur- face; scales on resting apex of main rachis 4-5 mm long, rigid, dark, very narrow, with short oblique marginal setae; scales on lower surface of costae dark and small, almost entire, mostly deciduous leaving the surface warty; very short hairs copious on lower surface of lamina and on costules; some hairs with the sporangia. Type: Lam 1935, Doorman summit (U; dupl. at L, Bo, BM). Distr. Malaysia: NW. New Guinea, once collected. Ecol. Altitude 2480 m. 28. Gleichenia alstonii HoL_tr. Reinwardtia 4 (1957) 267. Main rachis 4 mm diameter; primary branch- systems usually with forking of two orders, only the ultimate branches leafy; primary branches rarely proliferous beyond the first fork; branches of first order 4-5 cm long, of second order 6-7 em long; ultimate branches 30-35 cm _ long, 314 cm wide, the basal 10-15 mm sometimes Gleicheniaceae (Holttum) 23 bare of lamina; costules 4 mm apart, slightly oblique; segments of /amina c. 17 mm long, 3 mm wide above the base, thin, apices entire or nearly so, veins slightly prominent on both surfaces; sori of 3-4 sporangia. Scales on dormant apex of rachis 4 mm long, 0.7 mm wide, narrowly acumi- nate, brown, with short spreading pale marginal hairs near base, dark oblique sparse setae towards apex; scales on lower surface of costae scattered, dark, 1144 mm long, 0.2 mm wide, edges with sparse oblique dark hairs; scales on lower surface of costules pale, very narrow; scattered pale lax stellate hairs on lower surface of veins. Type: Alston 14981, Lae Pondom (BM). Distr. Malaysia: N. Sumatra (Tapanuli), once collected. Fig. 10. Gleichenia vestita BL. on ridge with ericoid ridge forest from Poka Pindjang to Rante Mario in the Latimodjong Range, Central Celebes, c. 3000 m (1937). 29. Gleichenia vestita BL. En. Pl. Jav. (1828) 249; Racis. Fl. Btzg 1 (1898) 11; v. A. v. R. Handb. (1908) 61; Suppl. (1917) 86; BACKER & PosTH. Varenfl. Java (1939) 255.—Sticherus vestitus CHING, Sunyatsenia 5 (1940) 285.—Fig. Te, 8c—d, 10. Rachis of a well-grown frond bearing several 24 FLORA MALESIANA [ser.: Il, ovolal pairs of primary branches, usually without stipular leaflets at their base; primary branch-systems usually forked to 2 orders (rarely 3), primary branch not proliferous beyond its first fork; all branches leafy, down to junction with main rachis; angle of forks about 45°; ultimate branches 15-25 cm long, lower ones 5—7 cm long; /amina cut down to within 1 mm of costa, segments distinctly oblique, thin but very rigid, 8-12 mm long, not narrowed immediately above the base but tapering very gradually, then abruptly to broadly pointed or rounded entire apex, edges not usually much reflexed, lower surface glaucous; costules 4-5 mm apart; veins slightly raised on both surfaces; sori of 3-5 sporangia, without evident paraphyses. Scales on dormant apex of main rachis firm, rusty to rather dark brown, 4-5 mm long, more than | mm wide, gradually narrowed to hair point, edges rather sparsely obliquely ciliate towards apex; scales on lower surface of costae abundant and persistent, spreading, rusty brown, 0.6-1.2 mm wide, 2—3 mm long, ciliate towards hair-pointed apex; smaller scales few, stellate hairs quite lacking; very short reddish simple hairs frequent on costules and veins beneath. Type: Blume, Java (L; dupl. at K). Distr. Malaysia: Sumatra, Java, S. Celebes. Ecol. In open alpine forest and scrub, 2300-— 2800 m. Note. At high elevations and in exposed places, fronds may be smaller than above described, with branches of first and second order only 2-3 cm long. 30. Gleichenia bolanica RosENsT. in Fedde, Rep. 12 (1913) 162; v. A. v. R. Handb. Suppl. (1917) 86.—G. monticola RiDL. Trans. Linn. Soc. Bot. 9 (1916) 252.—G. subulata v. A. v. R. Nova Guinea 14 (1924) 23.—Sticherus bolanicus COPEL. Philip. J. Sc. 75 (1941) 352.—Sticherus monticola NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 23. Lateral branch-systems of well-grown fronds with forks to 4th order; primary branches not normally proliferous beyond first fork nor bearing stipular leaflets at their base; ultimate branches 7-20 cm long, 6-10 mm wide, lower branches 1-2 cm long; usually only ultimate and penultimate branches leafy, but lower ones sometimes partly leafy; /amina cut almost down to the costa, segments apparently triangular because of their much-reflexed edges, 3 mm wide at the base, rather thick and very rigid when dry, lower sur- face glaucous with slightly raised veins; costules slightly oblique; sori often very crowded, of 3-6 sporangia. Scales on dormant apex of main rachis rigid, dark and shining, 3-4 mm long, to 0.7 mm wide, apex not hair-pointed, marginal hairs short, oblique, rigid; scales on lower surface of costae abundant, to 2 mm long, not hair- pointed, base 0.50.7 mm wide, brown at base and pale distally, edges bearing short hairs which are sometimes very rigid and dark brown; small similar scales or stellate hairs on costules and veins. Type: Keysser B. 14, Mt Bolan, New Guinea (B). Distr. Malaysia: New Guinea. Ecol. In open scrub or rocky places, fronds to 2 m or more long, 2800-3950 m. 31. Gleichenia hispida Metr. ex KUHN, Verh. K. K. Zool.-Bot. Ges. Wien 25 (1875) 600; RAcis. Fl. Btzg 1 (1898) 12; v. A. v. R. Handb. (1908) 61; BACKER & PostH. Varenfl. Java (1939) 255; Hoxtr. Reinwardtia 40 (1957) 270.—G. koordersii CuHrRist, Ann. Jard. Bot. Btzg 15 (1898) 76, pl. 13, f. 1.—Sticherus caudatus COPEL. Philip. J. Sc. 75 (1941) 354, pl. 2.—Sticherus hispidus Corer. Gen. Fil. (1947) 27.—Sticherus pinnatus COPEL. Philip. J. Sc. 83 (1954) 98, pl. 3.— Fig. 7c, 8e-g. Well-grown fronds with several pairs of primary branches which are not proliferous beyond their first fork and have no stipular leaflets; \ateral branch-systems with forks to 2nd or 3rd order; branches of first order leafless or only partly leafy, rest fully leafy; first order branches 214-6 cm long, second order branches 5-12 cm long; from second-order fork to end of ultimate branches (whether 2 or 3 forks present) 20-30 cm, the third fork (if present) about the middle of this length; /amina cut almost down to the costa, the segments slightly oblique, coriaceous, stiff and rigid when dry with much-recurved edges, longest segments on lower branches to 3!4 cm long, on ultimate branches sometimes only | cm long, effective width when dry 114-214 mm, costules 3-4 mm apart; lower surface of lamina more or less glaucous, veins dark and slightly prominent, not raised on upper surface; soriof 3 or 4sporangia with rather long crisped rusty hairs. Scales on dormant apex of main rachis rusty brown, 5-7 mm long, 0.5 mm wide, narrowed above the base and for the most part very narrow, hair-pointed; scales on lower surface of costae only a few cells wide at the base, the hair-point forming at least half of the length, hair-points crisped and entangled with those of other scales; scales on costules similar and smaller, grading to stellate hairs on veins and edges. Type: Jagor 558, G. Galungung (B; not seen). Syntype: De Vries, Ternate (dupl. at U). Distr. Malaysia: Sumatra, Java, Bali, N. Ce- lebes, Ternate, and Philippines (Negros, Luzon). Ecol. In thickets in open places, 1000-2200 m. Note. Isolated plants in exposed places may have fertile fronds much smaller and less branched than those described; Sticherus pinnatus COPEL. appears to represent a condition of this species with unbranched fertile fronds. 32. Gleichenia brassii C. Cur. Brittonia 2 (1937) 271.—Sticherus brassii NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 15. Rhizome 6 mm diameter, strongly warty from bases of old scales, young parts scaly, scales 5 mm long, 1 mm wide, ciliate; stipes to 100 cm long, persistently scaly throughout but only warty near the base, scales of all sizes with rather copious pale fringe, the longest ones hair-pointed; main rachis to about 200 cm long, bearing several pairs Dec. 1959] Gleicheniaceae (Holttum) 25 of primary branches which lack stipular leaflets; lateral branch-systems with forks of 2 orders, ultimate and penultimate branches fully leafy, ultimate forks less than 90°; first and second order branches 12-15 cm long, ultimate branches 30-40 cm long and 4-414 cm wide; costules about 5 mm apart, at right angles to costae; segments of lamina 3 mm wide above the base, firm, edges more or less revolute when dry, lower surface not glaucous when dry; veins slightly prominent on lower surface, not on upper surface except their bases; sori of 3-4 sporangia without evident para- physes. Scales on dormant apex of main rachis like those on rhizome but thinner, edge shortly ciliate, apex a short hair; scales on lower surface of costae spreading, 114-2 mm long, 0.5 mm wide, thin, brown, darker at the base, shortly hair- pointed, closely fringed with spreading pale hairs; scales on lower surface of costules abundant, 14-1 mm long, rusty brown with long pale fringe; upper surface of costae raised and terete, bearing persistent small pale long-fringed scales, rest of upper surface glabrous; lower surface of veins bearing many very short simple hairs. Type: Brass 4922, Mt Tafa (BM; dupl. at Bo). Distr. Malaysia: New Guinea (Mts Arfak and Tafa). Ecol. Scrambling in open places in forest, 1550-2400 m. 33. Gleichenia loheri CHRIST, Bull. Herb. Boiss. II, 6 (1906) 1009; v. A. v. R. Handb. (1908) 796; Ho.ttr. Reinwardtia 4 (1957) 272.—Sticherus loheri CoreL. Gen. Fil. (1947) 27.—Sticherus perpaleaceus CopeEL. Philip. J. Sc. 81 (1952) 3. var. loheri. Lateral branch-systems with forks of 1, 2 or 3 orders, ultimate and penultimate branches fully leafy, angle of ultimate fork much less than a right angle; with forks of 1 or 2 orders, penultimate branches 5—7 cm long, ultimate branches 17-23 cm long; with forks of 3 orders, ultimate and penultimate branches together 17-20 cm long; ultimate branches 114-2.2 cm wide, costules 3-314 mm apart, at right angles to costa or very slightly oblique, /amina cut almost down to the costa, segments not abruptly narrowed above the base, edges when dry somewhat reflexed, apex rounded and entire, texture firm but not coriaceous, veins distinctly raised on lower surface, slightly so above, lower surface not glaucous when dry; sori of 3-4 sporangia. Scales on apex of main rachis 5 mm long, 1 mm wide, thin, medium rusty brown, hair-pointed, edges with oblique slender hairs; scales on lower surface of costae copious and spreading, the largest 2 mm long, 0.8 mm wide, shortly hair-pointed; scales on costules very small, long-fringed, pale, mixed with long-armed stellate hairs; upper surface of costae raised and terete, more or less persistently covered with small pale long-fringed interlacing scales; very short simple hairs abundant on lower surface of veins and occasionally also on lamina. Type: Loher s.n., 7 Feb. 1906, Mt Banaho (P). Distr. Malaysia: Philippines (Luzon). var. major HoLtr. Reinwardtia 4 (1957) 272. Larger than typical form of species; ultimate branches 20-30 cm long, 2'4—31/4 cm wide, costules 3-4 mm apart; 3-lobed stipular leaflets 15 mm long sometimes present. Type: Biinnemeijer 11965, Celebes (Bo; dupl. Alea). Distr. Malaysia: S. Celebes, N. Borneo, and Philippines (Negros). Ecol. Altitude 2000-2500 m. 34. Gleichenia flabellata R. Br. Prod. (1810) 161; Hook. Sp. Fil. 1 (1844) 6; Domin, Bibl. Bot. 85 (1915) 204; v. A. v. R. Handb. (1908) 60; K. Scu. & Laur. Fl. Schutzgeb. (1901) 144.—Sticherus flabellatus H. St. JOHN, Occ. Pap. Bish. Mus. 17 (1942) 81. Stipe to 60 cm long, scaly at the base; main rachis bearing 2 or more pairs of branches; lateral branch-systems compact, forked to 3 orders at short intervals (lower branches com- monly 1 cm long, penultimate 2—3 cm long), ulti- mate and penultimate branches leafy, angle of forks less than 45°; ultimate branches 12-15 cm long, 2—3(-4) cm wide; costules 4 mm apart, at an angle of 45° to the costa; /amina cut almost to the costa, segments 214 mm wide above the base, edges toothed almost to the base, apex broadly pointed, veins very oblique and raised on both surfaces, lower surface not glaucous. Scales on apex of main rachis 4 mm long, shining medium brown, acuminate, fringe copious, long, lax; scales on lower surface of costae few and persistent only near base, narrow, hair-pointed and long-fringed; smaller scales or long-armed pale stellate hairs occasional on costules near base of branch, surfaces otherwise glabrous. Type: Robert Brown, Port Jackson, N. S. Wales (BM; dupl. at K). Distr. E. & SE. Australia, New Caledonia, New Zealand, in Malaysia: E. New Guinea (SCHUMANN & LAUTERBACH /.c.). 35. Gleichenia hirta BL. En. Pl. Jav. (1828) 250; v. A. vy. R. Handb. (1908) 60; Suppl. (1917) 85; Ho tr. Reinwardtia 4 (1957) 268. —See for further synonyms under the varieties.— Fig. 7d, 8h-j. Main rachis bearing several pairs of branches, not high-climbing; lateral branch-systems forked in 2-4 orders (rarely 5), the first-order branches in most varieties not normally proliferous beyond their first fork; ultimate branches always much longer than those of lower order, angle of ultimate forks less than 45°; /amina thin, cut almost down to the costa into thin oblique segments, edges distinctly toothed towards apices of segments, lower surface more or less glaucous (glaucous character destroyed by heat in drying). Scales on apex of main rachis to 5 mm long, less than 1 mm wide, fringed with short hairs; scales on costae abundant or not, spreading, 114-2 mm long, less than 14 mm wide, thin and rather pale, not 26 FLORA MALESIANA conspicuously hair-pointed, edges fringed with slender hairs; smaller scales and stellate hairs also present on costae, costules and veins, or quite [ser. IE vor lacking; sori with raised receptacle, conspicuous paraphyses present or not. KEY TO THE VARIETIES 1. Lower surface of costae of fully expanded ultimate branches scaly near the base only or more generally, the scales all about equal in size with rather rigid fringing hairs; no scales on costules nor on veins except when young. 2. Scales 3-4 cells wide at the base. Ultimate branches to 40 cm long and 344 cm wide. 2. Scales wider. Ultimate branches rarely over 25 by 214 cm. 3. Costules 4414 mm apart 3. Costules 314-4 mm apart 2. var. amoena 3. var. paleacea 4. var. candida . Lower surface of costae persistently scaly almost throughout, the scales of various sizes, often with lax fringe; smaller scales also persistent on costules and sometimes scales or hairs on veins. 4. Segments of the lamina not abruptly narrowed above the base, separated by narrow sinuses. 5. Scales on costules abundantly long-fringed. 6. Ultimate branches of well-grown fronds 2-3 cm wide; primary rachis-branches not always proli- ferous beyond first fork 1. var. hirta 6. Ultimate branches 114-214 cm wide; primary rachis- branches always proliferous beyond first fork. 5. Scales on costules with very few long marginal hairs 5. var. amboinensis 6. var. virescens 4. Segments of lamina c. 2 mm wide except at the very base, separated by | sinuses 1-114 mm wide. 1. var. hirta.x—Dicranopteris dolosa COPEL. in Perk. Frag. Fl. Philip. (1905) 193, 4. 4 fic.—G. dolosa C. Cur. Ind. Fil. (1906) 664; v. A. v. R. Handb. (1908) 62; Suppl. (1917) 85.—Sticherus hirtus CHING, Sunyatsenia 5 (1940) 203 (err. hirsutus). Lateral branch-systems forked in 3 or 4 orders; ultimate branches 15—22 cm long, 2-214 cm wide (rarely to 3 cm); penultimate branches fully leafy and often wider than ultimate ones; costules 3 mm apart; scales on lower surface of costae copious, persistent, of many sizes together, very long- fringed; pale stellate hairs present on veins. Type: Reinwardt, Tidore (L; dupl. at BM). Distr. Malaysia: Moluccas and Philippines (Luzon, Mindoro, Mindanao), 1200-1800 m. 2. var. amoena (v. A. v. R.) Hott. Reinwardtia 4 (1957) 269.—G. amoena v. A. v. R. Bull. Jard. Bot. Btzg II, . 23 (1916) 12; Handb. Suppl. (1917) 497—Sticherus amoenus NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) .—G. peninsularis CopeL. Un. Cal. Publ. Bot.12 (1931) 387.—Sticherus peninsularis CHING, Sunyatsenia 5 (1940) 284.— G: lurta; Hottrr. Rev. Fl. Mal. 2 (1955) 7- Lateral branch-systems forked in 1-3 orders; penultimate branches fully or partly leafy, branches of lower order not leafy; ultimate branches 25-40 cm long, 2!4-314 cm wide; costules 4-414 mm apart; /amina-segments 3—-3\/ mm wide above the base, strongly glaucous beneath; sca/es on lower surface of costae. narrow (fringe of hairs longer than width of scale), very sparse and near base of costae only, mature fronds otherwise glabrous (stellate hairs on veins when young). Type: ateP): Distr. Malaysia: Sumatra, Lingga Archipelago, Malay Peninsula. Teysmann 16628, Lingga (Bo; dupl. 7. var. lanuginosa Ecol. Altitude 700-1600 m; scrambling or trailing in lightly shaded places or edge of forest, not forming dense thickets; fronds usually not over 200 cm long. 3. var. paleacea (BAK.) C. Cur. Gard.Bull. S. S. 7 (1934) 212; Hottr. Reinwardtia 4 (1957) 269.—G. vestita var. paleacea BAK. J. Bot. 17 (1879) 38.—G. hallieri CuHrist, Ann. Jard. Bot. Btzg 20 (1905) 138; v. A. v. R. Handb. (1908) 61.— G. barbula C. Cur. Dansk Bot. Ark. 9, 3 (1937) 67.—Sticherus hallieri NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 18.—Sticherus barbulus NAKAI, byes, Mia} Lateral branch-systems usually forked in 2 or 3 orders (to 5 orders in type specimen of G. barbula); if in 2 orders penultimate branches incompletely leafy and ultimate branches to 30 cm long, other- wise ultimate branches 15—24 cm long and 2-214 cm wide; costules 4-4!4 mm apart; scales on mature fronds confined to costae and all alike (not mixed with smaller ones) with rather stiff fringing hairs as long as width of scale. Type: Burbidge, N. Borneo (K). Distr. Malaysia: Borneo, Lesser Sunda Islands (Bali), Celebes (Menado), 1100-3000 m. Ecol. On Mt Kinabalu on open places in ridge forest. 4. var. candida (RoseNstT.) Hott. Reinwardtia 4 (1957) 269.—G. candida RosENstT. in Fedde, Rep. 5 (1908) 33; v. A. v. R. Handb. (1908) 796.— Sticherus hirtus var. candidus NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 19. Similar to var. paleacea, but costules 314-4 mm apart, lamina of firmer texture. Type: Werner 72, Mt Gelu (B; dupl. at BM, US, EE Distr. East Malaysia: New Guinea, Admiralty Islands, Solomon Islands. Dec. 1959] Ecol. On edges of forest and in light shade, sometimes forming dense thickets to 3 m high, 700-2150 m. 5. var. amboinensis (v. A. v. R.) Hott. Rein- wardtia 4 (1957) 269.—G. amboinensis v. A. v. R. Bull. Dép. Agr. Ind. Néerl. 7. 18 (1908) 3; Handb. (1908) 62; Suppl. (1917) 85 (as a synonym). Like var. hirta in scaliness; ultimate branches 1144-2144 cm wide; primary rachis-branch always proliferous beyond first fork and bearing 2 or more pairs of secondary branches. Type: Teysmann, Amboyna (Bo; dupl. at L). Distr. Malaysia: S. Central Celebes, Moluccas (Ambon, Buru, Waigeo Isl.). Ecol. Climbing to 4 m in open places in mountain forest. 6. var. virescens (HIERON.) HoLtr. Reinwardtia 4 (1957) 270.—G. dolosa var. virescens HIERON. ex BRAUSE, Bot. Jahrb. 56 (1920) 209.—Sticherus hirtus var. virescens NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 19. Similar to var. hirta, but scales on costules with very few long marginal hairs. Gleicheniaceae (Holttum) pe | Type: Ledermann 9935 (B). Distr. Malaysia: E. New Guinea. Note. Collector of type specimen stated that lower surface of lamina was white (i.e. glaucous) when fresh. 7. var. lanuginosa (v. A. v. R.) comb. nov.—G. ornamentalis ROSENST. Nova Guinea 8 (1912) 715; v. A. v. R. Handb. Suppl. (1917) 85.—G. ornamentalis var. lanuginosa vy. A. v. R. Nova Guinea 14 (1924) 23.—Sticherus lamianus COPEL. Philip. J. Sc. 75 (1941) 356, pl. 6.—G. hirta var. ornamentalis (ROSENST.) HoLtr. Reinwardtia 4 (1957) 269. Lateral branch-systems commonly forked to 4 orders; costae persistently and densely scaly, scales as in var. hirta; costules 3-4 mm apart, segments of lamina 2 mm wide, thus separated by rather wide sinuses. Type: Lam 1945, Doormantop, 3480 m (U; dupl. at Bo, US, L). Distr. Malaysia: New Guinea. Ecol. Altitude 800-2500 m; specimens from the higher altitudes are the most densely scaly. 2. DICRANOPTERIS BERNHARDI in Schrad. Neu. J. Bot. 1, 2 (1806) 38; UNDERW. Bull. Torr. Bot. Cl. 34 (1907) 244, p.p.; CHING, Sunyatsenia 5 (1940) 272; CopeL. Gen. Fil. (1947) 28; NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 56; HoLtr. Reinwardtia 4 (1957) 273.—Mertensia WILLD. Kongl. Vet. Ak. Nya Handl. 25 (1804) 163, p.p.; PRESL, Tent. Pterid. (1836) 50, p.p.; non RoTH, 1797 (Borag.).—Gleichenia subg. Mertensia, § 1 and § IV, Hook. Sp. Fil. 1 (1844) 11, 12.—Hicriopteris PRESL, Epim. Bot. (1851) 26, non CHING, nec COPEL.—Gleichenia subg. Mertensia sect. Heteropterygium Diets in E. & P. Pfl. Fam. 1, 4 (1900) 355.—Gleichenia subg. Mertensia sect. Dicranopteris v. A. v. R. Handb. (1908) 56, p.p.—Mesosorus Hassk. Fil. Jav. 1 (1856) 2, p.p.—Fig. 11-16. Indumentum on rhizome, dormant apices, and on other parts of fronds con- sisting of multiseptate hairs which have outgrowths from cells near the base and sometimes also from other cells. Apex of main rachis of fronds resting periodically while each pair of lateral branch-systems develops; primary rachis-branches repeatedly branched pseudo-dichotomously (forked), the apex between each pair of branches usually permanently dormant, the members of a pair equal or unequal; a short stipule-like lobed leaflet usually present at the base of each primary rachis-branch, these leaflets growing upwards and protecting the temporarily dormant apex of the main rachis (smaller stipule-like leaflets sometimes also present at forks of the lateral branch-systems); a pair of accessory branches, bearing a lamina like that of the ultimate branches, present at some of the forks of the lateral branch-systems, on the outside of the fork (usually attached just above it) and deflexed; apart from stipular leaflets and accessory branches, only the ultimate branches leafy. Ultimate branches bearing throughout a deeply pinnatifid /amina, the segments of the lamina usually entire, each with costule bearing lateral veins which are forked at least twice; sori one to each vein-group on an acroscopic branch (rarely also on a basiscopic branch), each of 8-15 or . 500 m (MEIJER, [ser. Il; voles INN we f ' / | f < Z < DN | sa) 4 < ~S = SS Z| me | eo) 4 ew i i] Ly Fig. 11. Dicranopteris curranii CopeL. Harau gorge, near Pajakumbuh, W. Sumatra, c Dec. 1959] | Gleicheniaceae (Holttum) 29 more sporangia without paraphyses, the sporangia smaller than in Gleichenia; spores monolete or trilete. Distr. and Ecol. Thicket-forming ferns, abundant in open places throughout the wetter parts of the tropics and subtropics, especially characteristic of equatorial lowlands (the species of Gleichenia being almost exclusively mountain plants in Malaysia). Notes. There are two well-marked subgenera, Dicranopteris (pantropic) and Acropterygium (one species, in tropical America; no accessory branches, rhizome with solenostele). The subgenus Dicranopteris is much more diversified in Malaysia than in any other part of the world. Asmall group of species may be clearly distinguished by their monolete spores. Almost all other specimens are at present regarded as belonging to a polymorphic species, D. linearis. Some of the varieties of D. linearis are more clearly distinct than others, and probably should be recognized as species. They /, 11K Zz vil my \ ‘ .* C d © Fig. 12. Diagrams showing variations of branching-habit in Dicranopteris. a. Basic habit, found in D. linearis (BuRM. f.) UND. var. linearis, D. pubigera (BL.) NAKAI, and D. curranii COPEL., b. accessory branches present at ultimate forks, as in D. linearis var. montana HOLtT. and var. tetraphylla (Ros.) NAKAI, c. alternate unequal forking, as in D. linearis var. subpectinata (CurRIstT) HoLtt., var. alternans (Me_tt.) Hottt., and var. subspeciosa Ho.tr., d. special development of c, found only in D. speciosa (PRESL) HOLTT. 30 FLORA MALESIANA [ser. IT, volyatz Fig. 13. Dicranopteris brake in an old crater swamp at c. 1200 m, Gajo Lands, N. Sumatra, surrounded by pole wood forest consisting mostly of J/ex cymosa (1937). need, however, to be more clearly characterized from field studies. The existence of a triploid hybrid (see note below on cytology) indicates that it is probably impossible to refer every plant to a clearly defined variety. Four varieties of D. linearis have the common character of invariably producing accessory branches at the bases of ultimate branches; in other characters they differ considerably from each other and do not form a natural group. Another variety having this character occurs commonly in west tropical Africa, but no such variety has been found in east Africa. Cytology (observations by I. MANTON). Haploid chromosome number of D. curranii CopeL. (Singa- pore) is 39. D. linearis (apparently the normal form of the species) in Ceylon is either diploid (n = 39) or tetraploid (n = 78); a plant of D. linearis sent to Kew from Singapore is a sterile triploid. KEY TO THE SPECIES 1. Spores monolete. The two branches at each fork equal; either lowland plants with ultimate branches 9-12 cm wide, or mountain plants. First fork of each vein at its very base. 2. Lamina not very thick nor with the veins grooved on the lower surface. Dec. 1959] Gleicheniaceae (Holttum) ¢ 31 60. Except in fig. a and b the cell of attachment is gS black.— a. Hair from base of costa. — D. clemensiae Ho.tr. b. One of the rigid, shining hairs from lower surface of costa, c. thin-walled hair from costule. — D. pubigera (BL.) NAKAI. d-e. Hairs from costule, lower surface. — D. linearis (BURM. f.) UND., hairs from lower surface of costules: Fig. 14. Hairs of Dicranopteris, all x D. curranii COPEL. f. var. subferruginea (H1IERON.) NAKAI, g. var. linearis, h. var. sf subspeciosa HOutTrt., i. var. alternans (Metr.) HoLtr. 3. Ultimate branches commonly 40 cm long and 9-12 cm wide; lower surface almost glabrous. 1. D. curranii 3. Ultimate branches not over 7 cm wide; lower surface of costae and costules rather persistently hairy. 4. Ultimate branches 30-55 cm long, 5—7 cm wide; costae at first with many shining hairs (1-1) mm long) throughout . 2451 DB). pee 4. Ultimate branches 15-25 cm long, D qe 5 cm wide: costae with such hairs near base only. 2. Lamina very thick, the veins grooved on the lower surface 3. D. pubigera 3. D. pubigera 1. Spores trilete. Branches at a fer’ in many cases unequal; ultimate branches to c. 6 cm wide. Veins never grooved on lower surface; tis. ork of each vein distinctly above the base. 5. At each fork, except primary and ultimate ones, one branch leafy and without accessory branch, the other branch not leafy and with an accessory branch 4. D. speciosa 5. At each fork, except (in most cases) an ultimate one, a pair of accessory branches present. 1. Dicranopteris curranii CopeL. Philip. J. Sc. 81 (1952) 4; Hottr. Reinwardtia 4 (1957) 274.—Glei- chenia hermannii var. venosa BL. incl. also var. tenera BL. En. Pl. Jav. (1828) 249.—Gleichenia dichotoma var. tenera METT. in Mig. Ann. Mus. Bot. Lugd. Bat. 1 (1863) 50.—Gleichenia dichotoma var. malayana Curist, Ann. Jard. Bot. Btzg 15 (1898) 77.—Gleichenia linearis var. malayana v. A. v. R. Handb. (1908) 59; Suppl. (1917) 84; Hottr. Rev. Fl. Mal. 2 (1955) 70.—D. lessonii NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 61, guoad specim., excl. basion. Mertensia lessonii A. RicH.—Fig. 11, 12a, 14a, 16. Fronds large, strongly erect but not greatly elongate nor high-climbing; primary rachis- branches few times equally forked; ultimate branches commonly 40 cm long and 12 cm wide, 5. D. linearis accessory branches at penultimate forks commonly 20-30 cm long and 7—10 cm wide; costules 5—7 mm apart; Jamina-segments 314-4144 mm wide above the base, texture firm, lower surface glaucous, usually quite glabrous apart from minute simple hairs on the veins, often pinkish on costae and lower parts of costules; veins slender, slightly prominent on lower surface but not on upper; stipular leaflets to 3 cm long at bases of primary branches; spores monolete. Type: Curran, For. Bur. 19265, Laguna Prov., Luzon (H. Copel.; dupl. at US, P). Distr. Malaysia: Sumatra, Malay Peninsula, W. Java, Lesser Sunda Islands (Flores), Borneo, Celebes, Philippines (Luzon). Ecol. In thickets with varieties of D. linearis, from sealevel up to 1500 m. 32 FLORA MALESIANA [ser. II, vol. 12 Note. Gleichenia weatherbyi FosBERG (Am. Fern J. 40, 1950, 140), from the Caroline Islands, is very near D. curranii, with similar spores, but is even larger, and much more hairy. 2. Dicranopteris clemensiae Hott. Reinwardtia 4 (1957) 275.—Fig. 14b-c. Main rachis 7 mm or more diameter; primary branches twice equally forked, copious dark red rigid hairs very persistent near the furks; ultimate branches 30-55 cm long, 5—7 cm wide, the lowest outer segments of the lamina sometimes enlarged and more or less lobed; accessory branches at penultimate forks 16-18 cm long, 5 cm wide; stipular leaflets at base of primary branches 3 cm long, lobes 3 mm wide; costules of ultimate branches 5—6 mm apart; /amina-segments 314 mm wide above the base, thin but firm; veins usually 3 times forked, first fork very close to the costule, very slightly prominent on both surfaces: stout dark red multicellular hairs (1-114 mm long, with several basal branches) at first abundant all along lower surface of costae, at length deciduous leay- ing costae perceptibly rough; thinner and paler, dull rusty-brown, hairs at first abundant on costules, slightly crisped and with short unicellular branches near base only; sori of 10-15 sporangia, often two on a vein-group (on each of the outer- most branches); spores monolete, c. 32 16 yu. Type: Clemens 28745, Mt Kinabalu (BM; dupl. at Bo, K, L, US). Distr. Malaysia: N. Borneo (once collected, 1600 m). 3. Dicranopteris pubigera (BL.) NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 68; Hott. Rein- wardtia 4 (1957) 274.—Gleichenia hermannii var. pubigera Bu. En. Pl. Jav. (1828) 249.—Gleichenia dichotoma var. pubigera Metr. in Mig. Ann. Mus. Bot. Lugd. Bat. 1 (1863) 50.—Gleichenia linearis var. pubigera v. A. vy. R. Handb. Suppl. (1917) 85.—Mertensia spissa (non FEE) var. pu- bigera NaAKal, Bot. Mag. Tokyo 39 (1925) 180.— Fig. 12a, 14d—e, 15a-b. Primary rachis-branches 2-4 times equally or subequally forked; ultimate branches 15-25 cm long, 3-8 cm wide; costules 414-6 mm apart; lamina-segments c. 4 mm wide when flattened, thick and rigid when dry and usually with edges reflexed; lower surface not glaucous, the costules at first covered rather densely with coarse dull rusty flexuous hairs and also some rigid shining hairs; veins grooved on the lower surface when the lamina is very thick, on thirner leaves hardly grooved, pale but not raised on the upper surface; spores monolete. Type: Blume, Java (L; dupl. at K, P). Distr. Malaysia: Sumatra, Java, and Lesser Sunda Islands (Bali, Lombok, Flores). Ecol. Abundant in open scrub and in clearings in mountain forest, forming dense thickets, 1100-3000 m. Note. The extreme forms of this species, one with very thick lamina with veins grooved on the lower surface, the other with thinner lamina and veins not distinctly grooved, appear very different (the former is larger as well as thicker) but there are intermediates. The differences may in part be connected with altitude and exposure; no ob- servations have been made. Fig. 15. Dicranopteris pubigera (BL.) NAKAI. a. Lower surface of part of frond (veins grooved), < 4, b. a main fork with stipular leaflets and ac- cessory branches, * 1%. — D. speciosa (PRESL) Ho itr. c. Part of frond to show venation, x 2, d. main fork showing stipular leaflets, «x %. 4. Dicranopteris speciosa (PRESL) HoLtr. Rein- wardtia 4 (1957) 273.—Hicriopteris speciosa PRESL, Epim. Bot. (1851) 27.—Gleichenia opposita v. A. v. R. Bull: Jard. Bot. Btzg I, 72 TGs ise Ho tt. Rev. Fl. Mal. 2 (1955) 70, f. 14 F.—Glei- chenia parallela RiDL. J. Mal. Br. R. As. Soc. 4 (1926) 3.—D. opposita NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 68.—Fig. 12d, 15¢e—-d. Primary rachis-branches several times very unequally forked; at each fork the smaller branch unbranched and leafy throughout, commonly 20-25 cm long and 4-5 cm wide, with no accessory branch, the larger branch continuing almost the same straight line as the larger branch from the previous fork and bearing an accessory branch 12-15 cm long and 314 cm wide (this accessory branch almost opposite the simple leafy branch of the fork); ultimate fork formed by two normal leafy branches without accessory branches; irregularities from this basic scheme occasionally present; lJamina-segments 31,4 mm wide, flat and rather rigid, separated by very narrow sinuses, apices often retuse; veins distinctly prominent on the upper surface; lower surface more or less glaucous, the costules when young rather densely covered with fine pale entangled woolly hairs, some similar hairs on veins; some dark redbrown, rigid, shining hairs, like those on resting apices, also at first present on costae and costules; sori as in D. linearis; spores trilete. Type: collector unknown, ‘“‘Pendschab”’ (PRc). Perhaps from Penang. Distr. Malaysia: Sumatra, Malay Peninsula, Moluccas (Sula Isl.: P. Mangoli). ——————— Dec. 1959] Gleicheniaceae (Holttum) 35 Ecol. Growing with D. linearis on the edges of forest, 100-600 m. Note. The remarkable pseudo-pinnate lateral branch-systems of this species were described in detail by PresL, but because he compared these branches to Gleichenia glauca (THUNB.) HOOK. later authors thought he was describing a species of Gleichenia subg. Diplopterygium and conse- quently used the generic name Hicriopteris in a wrong sense. 5. Dicranopteris linearis (BURM. f.) UNDERW. Bull. Torr. Bot. Cl. 34 (1907) 249; Ho.ttr. Rein- wardtia 4 (1957) 275.—Polypodium dichotomum THUNB. FI. Jap. (1784) 338, t. 37.—Gleichenia dichotoma Hook. Sp. Fil. 1 (1844) 6; Racts. Fl. Btzg 1 (1898) 13.—Fig. 12, 14f-i. KEY 10, DHE VARIETIES 1. Accessory branches not always present at ultimate forks. 2. Branches at each fork of a lateral branch-system approximately equal; forked 1-3 times. lateral branch-systems 3. Costules less than 5 mm apart, their lower surface with some persistent hairs. MeeLAIrS not persistent om lower surface OlevelnS -)ee 2 4) 6 4 8 6 6 Le var. linearis 4. Hairs persistent along lower surface of veins. 5. Hairs long, very finely woolly and entangled; no distinct separate hairs present. 2. var. ferruginea 5. Hairs coarse, shorter, more or less entangled on costules, some distinctly separate on veins. 3. var. subferruginea 3. Costules 5-7 mm apart, their lower surface glabrous. 6. Lamina thick and gid 6. Lamina thin 2. Branches at successive forks alternately unequal; forked at least 3 times. 7. Lower surface quite glabrous and persistently pale glaucous; 4. var. rigida Sane 5. var. latiloba lateral branch- -systems on well-grown plants lamina thin 6. var. subpectinata 7. Lower surface more or less persistently hairy, less strongly glaucous (often losing glaucous character on drying); lamina firm. 8. Hairs very slender, their long branches much crisped and entangled; no separate short hairs. 9. Hairs copious, at first covering whole lower surface, persistent along the veins, rusty when dry; edges of lamina strongly reflexed when dry . 2. var. ferruginea 9. Hairs not covering whole lower surface, persistent only. on “costules, pale when dry; edges of lamina not much reflexed when dry. 10. Angle of secondary and later forks not more than a right angle 10. Angle of secondary and later forks more than a right angle . . . 8. 7. var. subspeciosa var. inaequalis 8. Hairs coarser and shorter, some more or less crisped and entangled but separate short hairs also present. 11. Hairs confined to bases of some of the costules 1. var. linearis 11. Hairs more abundant on costules, and some also on veins. 12. Hairs persistently rust-coloured, copious all along veins. ns 12. Hairs pale on dried specimens, sparse on the veins; W. Malaysia 3. var. subferruginea 9. var. alternans 1. Accessory branches always present at ultimate forks. 13. Accessory branches, especially the lower ones, distinctly below the fork; branches at successive forks alternately somewhat unequal . 10. var. demota 13. Accessory branches opposite the fork or distinctly above it; branching usually equal at all forks. 14. Ratio of length to width of branches ee Die 214-3 mm wide 1; costules c. 3 mm apart, segments of lamina 11. var. tetraphylla 14. Ratio of length to width greater: costules more widely ‘separated. 15. Costules on ultimate branches 5-7 mm apart; texture subcoriaceous; lower surface quite glabrous; accessory branches at ultimate forks almost opposite the fork . . . 12. var. montana 15. Costules on ultimate branches 314-414 mm apart; texture thin; some rusty hairs often on lower surface; accessory branches at ultimate forks 3-4 mm above the fork 1. var. linearis.—Polypodium lineare Burn. f. Fl. Ind. (1768) 235, t. 67 f. 2.—Gleichenia herman- nii R. Br. Prod. (1810) 161.—Gleichenia dichotoma var. normalis MetT. in Mig. Ann. Mus. Bot. Lugd. Bat. 1 (1863) 50; Racts. FI. Btzg 1 (1898) 13.—Mertensia pteridifolia PRest, Abh. (K.) Bohm. Ges. Wiss. M.-N. Cl. V, 5 (1848) 339; Epim. Bot. (1851) 23, t. 14.—Gleichenia linearis CLARKE, Trans. Linn. Soc. Bot. 1 (1880) 428; v. A. v. R. Handb. (1908) 59 pro var. normalis; Suppl. 13. var. altissima (1917) 84, ditto; Hottr. Rev. Fl. Mal. 2 (1955) 70, ditto.—Fig. 12a, 14g, 16. Primary rachis-branches commonly 2 or 3 times forked, the two branches at all forks equal or nearly so (not regularly alternately unequal); stipular leaflets at bases of primary branches c. 1 cm long, lobed near the base only; ultimate branches 15-25 cm long, 4-6 cm wide; /amina- segments 214-3 mm wide, separated by rather wide sinuses, texture very firm; lower surface slightly 34 FLORA MALESIANA [ser. Il, volets glaucous; veins slightly prominent on lower surface and bearing very short simple hairs, not prominent on upper surface; some persistent much- branched rusty hairs on costules; spores trilete. Type: Ceylon, Herb. Delessert (G). Distr. Tropical and subtropical Africa, Asia, Malaysia, Australasia, and Polynesia. Ecol. Thicket-forming but not high-climbing, from sealevel to 1400 m. Note. Polypodium dichotomum THUNB. was described from Japan, and as noted by NAKAI (/.c.) is not quite identical with Polypodium lineare BuRM. (BURMAN’S specimens, preserved at Ge- neva, are from Ceylon). I have therefore ranked it as a Dicranopteris linearis var. dichotoma (Rein- wardtia 4, 1957, 277). The lamina of the fronds is thinner than in var. linearis and the veins are prominent on the upper surface. This may be connected with the fact that (at least in part of its range) the fronds of var. dichotoma only last for one season, dying in the winter, whereas those of var. linearis grow for an indefinite period which is usually not climatically limited. Var. dichotoma apparently occurs also in China and probably in the Sikkim region; field studies in these regions are needed to establish clear distinctions between var. linearis and var. dichotoma. 2. var. ferruginea (BL.) Ho_tr. Reinwardtia 4 (1957) 278.—Gleichenia ferruginea Bu. En. Pl. Jav. (1828) 249; Metr. Ann. Mus. Bot. Lugd. Bat. 1 (1863) 50.—Gleichenia dichotoma_ var. ferruginea Racis. Fl. Btzg 1 (1898) 13.—Gleichenia linearis var. ferruginea v. A. v. R. Handb. (1908) 59; Suppl. (1917) 85.—‘Dicranopteris ferruginea’ CopeL. Philip. J. Sc. 75 (1941) 349, not Mertensia ferruginea Desv. 1811. Lateral branch-systems of small plants sym- metrically forked twice, on larger plants forked to 3 or more orders, branches at successive forks alternately unequal; ultimate branches 15—20 cm long, 3-5 cm wide; costules 4-5 mm apart; seg- ments of lamina brittle when dry, with recurved edges, when flattened c. 214 mm wide above the base, lower surface covered with very fine floccose entangled pale rusty hairs (c. 10 w diam.) which are attached mainly along the veins, the veins thus persistently hairy on old fronds; stiff hairs present on dormant apices only; upper surface smooth, the veins very distinct and slightly raised when dry; accessory branches not normally present at the ultimate forks, but the lowest basi- scopic lobes of the ultimate branches usually larger than the rest, their edges more or less lobed. Type: Blume, Java (L). Distr. Malaysia: Java, Celebes, Moluccas, New Guinea. Ecol. Altitude 1000-2500 m. 3. var. subferruginea (HIERON.) NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 66.—Gleichenia linearis var. subferruginea HIERON. ex BRAUSE, Bot. Jahrb. 56 (1920) 209.—Fig. 14f. Like var. linearis but perhaps more often with unequal branches at the more distal forkings, veins somewhat prominent on upper surface, veins on the lower surface rather persistently hairy, the hairs rusty, rather coarse (diam. c. 25 yw) and much-branched, not finely woolly nor forming a continuous entangled web as in var. ferruginea. Type: Ledermann 6926, New Guinea (B). Distr. Queensland, Polynesia (Fiji, Samoa), and East Malaysia: Celebes, Moluccas, New Guinea. Ecol. From sealevel to 1200 m. Note. It is probable that this variety is much more abundant than var. linearis in New Guinea and regions further eastwards. The hairs on the lower surface of the veins are of the same general character as those of var. linearis, but much more copious; they are more abundant on plants at higher altitudes than near sealevel, and possibly abundance of hairs also depends on degree of exposure. 4. var. rigida (BL.) Hottr. Reinwardtia 4 (1957) 277.—Gleichenia hermannii var. rigida BL. En. Pl. Jav. (1828) 249.—Mertensia crassifolia Pres_, Abh. (K.) Béhm. Ges. Wiss. M.-N. Cl. V, 5 (1848) 339; Epim. Bot. (1851) 23, t. 13.— Gleichenia dichotoma var. rigida METT. in Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 50.—Glei- chenia warburgii Curist, Ann. Jard. Bot. Btzg 15 (4898) 78.—Gleichenia linearis var. rigida vy. A. v. R. Handb. (1908) 59; Suppl. (1917) 84.—Glei- chenia crassifolia CopeL. Philip. J. Sc. 1 (1906) Suppl. I 257.—D. crassifolia NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 57. (Non Mertensia rigida Kze 1834, D. rigida NAKAI). Lateral branch-systems once or twice forked, members of all forks equal or nearly so; ultimate branches 15-25 cm long, 3—5!4 cm wide; costules 5-6 mm apart; segments of lamina 4-5 mm wide above the base, thick and very rigid when dry with edges slightly reflexed, apices not or slightly retuse; lower surface quite glabrous from an early stage; veins broad, slightly prominent on lower surface, distinctly so on upper surface; spores trilete. Type: Reinwardt, Tidore (L). Distr. Malaysia: Moluccas (type from Tidore), Celebes, and Philippines (Mindanao, Luzon, Negros). Ecol. Altitude 1500-3000 m. Note. MEeETTENIUS (and apparently also RACI- BORSKI) referred some specimens of D. pubigera from Java to this variety of D. linearis. 5. var. latiloba HoLtr. Reinwardtia 4 (1957) 277. Lateral branch-systems forked to about 4th order, branches at all forks subequal; stipular leaflets at bases of primary branches to 4 cm long, deeply and broadly lobed; ultimate branches c. 20 cm long, to 9 cm wide, the lowest segments of the lamina somewhat enlarged and deflexed but not pinnately lobed; costules c. 7 mm apart; segments 4-5 mm wide above the base, firm but not very thick, edges slightly sinuous, only reflexed when dry; lower surface quite glabrous apart from minute simple hairs; veins sometimes Dec. 1959] ; Gleicheniaceae (Holttum) 35 Fig. 16. Thicket of Dicranopteris on edge of forest, Singapore. D. curranii CopPeEL. (large fronds) to left and right; D. linearis (BURM. f.) UND. var. linearis in centre; var. subpectinata (CHR.) HOLTT. to right of centre, below (HoLtTrum, pale and distinctly prominent on the upper surface, concolorous and only slightly prominent on lower surface. Type: Merrill 975, Luzon (US; dupl. at P, U, F). Distr. Malaysia: Philippines (Luzon, Negros, Mindanao), Celebes (?). Ecol. Altitude 1000-1600 m. 6. var. subpectinata (CHrRist) Ho.tr. Rein- wardtia 4 (1957) 277.—Gleichenia subpectinata CurisT, Bot. Tidsskr. 24 (1901) 111.—Gleichenia Pteridifolia (non Mertensia pteridifolia PRESL) RIpL. J. Mal. Br. R. As. Soc. 4 (1926) 4.—D. warburgii (non Gleichenia warburgii CHRIST) sensu NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 70.—Gleichenia linearis var. alternans (non METT.) Ho tr. Rev. Fl. Mal. 2 (1955) 70 and frontispiece. —Fig. 12c, 16. Lateral branch-systems several times forked, branches at successive forks alternately unequal, successive larger branches forming almost a Straight line; ultimate branches 9-15 cm long, 2-314 cm wide (sterile ones sometimes to 18 by 6 cm); /amina thin, lower surface pale glaucous and quite glabrous when mature; segments of 1925). lamina gradually and rather evenly tapering from base to apex, the sinuses thus narrowly triangular; costules 4414 mm apart; veins slightly raised on both surfaces, concolorous with lamina; the smaller accessory branches usually attached 3-5 mm above the forks (sometimes lacking or reduced at penultimate forks). Type: Schmidt, Koh Chang Isl. (P). Distr. Lower Siam (type from Koh Chang Island), and Malaysia: Sumatra, Malay Peninsula, Lingga Archipelago, Banka, Borneo. Ecol. On edges of forest, in thickets with var. linearis and other varieties, climbing higher than var. linearis, from sealevel up to 700 m. 7. var. subspeciosa Hott. Reinwardtia 4 (1957) 278.—Fig. 12c, 14h. Resembling D. speciosa in shape, texture etc. of ultimate branches and in hairiness, but lateral branch-systems branched as in var. alternans. Type: Topping 1516, Mt Kinabalu, Kiau (US; dupl. at SING). Distr. Malaysia: Sumatra, Malay Peninsula, Borneo, and Philippines (Mindanao, Mindoro, Luzon). 36 FLORA MALESIANA [ser. II, vol. 11, Dec. 1959] Ecol. On edges of forest, in thickets with other varieties, 0-1400 m. 8. var. inaequalis (ROSENST.) HoLTr. Rein- wardtia 4 (1957) 278. — Gleichexia linearis var. inaequalis ROSENST. in Fedde, Rep. 13 (1915) 212; vy. A. v. R. Handb. Suppl. (1917) 85. Shape and venation of /amina-segments, and hairiness, as in var. subspeciosa, differing as follows in branching: forking in lateral branch-systems alternately very unequal, the angle of the forks much more than a right angle (sometimes nearly 180°), the smaller branch at a fork simple or once forked and sometimes lacking anaccessory branch, the accessory branch on the larger branch at- tached 5-10 mm above the base of that branch. Type: J. Winkler 114, Batakerland, Sumatra (H. Selim Birger, S). Distr. Malaysia: Sumatra, Malay Peninsula (three collections in all). Ecol. Altitude c. 1400 m. 9. yar. alternans (MetTT.) HoLtr. Reinwardtia 4 (1957) 278.—Gleichenia dichotoma var. alternans Me_etr. in Mig. Ann. Mus. Bot. Lugd. Bat. 1 (1863) 51.—Gleichenia linearis var. alternans v. A. v. R. Handb. Suppl. (1917) 84.—Fig. 12c, 14i. Very similar to var. linearis in texture, shape, and hairiness of the ultimate branches, but lateral branch-systems several times forked, the branches at successive forks alternately unequal about as in var. subpectinata, differing from the latter in larger somewhat hairy ultimate branches of much firmer texture. Type: Korthals, Sumatra (L). Distr. Malaysia: Sumatra, Malay Peninsula, Banka, Borneo. Ecol. Apparently not common, but needs field study; perhaps only a growth-form of var. linearis; from sealevel up to 500 m. 10. var. demota Ho.tr. Reinwardtia 4 (1957) Did Lateral branch-systems several times forked, branches at successive forks alternately unequal; accessory branches normally present at all ulti- mate forks; ultimate branches to 18 cm long, 3-6 (rarely to 7) cm wide; costules 5 mm apart; lamina thin, lower surface glabrous and probably glaucous; veins distinctly prominent on upper surface and slightly so on lower surface; accessory branches at the lower forks attached distinctly below the fork. Type: Clemens 29535, Mt Kinabalu, 5600 ft (K; dupl. at BM, Bo, L, US). Distr. Malaysia: N. Borneo, Ceram (doubt- ful), New Guinea. Ecol. Altitude 1600-2250 m. 11. var. tetraphylla (ROsENST.) NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 67.—Gleichenia linearis var. tetraphylla ROSENST. in Fedde, Rep. 13 (1914) 213; v. A. v. R. Handb. Suppl. (1917) 84. —Fig. 12b. Lateral branch-systems several times forked, branches at successive forks alternately somewhat unequal; penultimate branches less than 1 mm diam. when dry; accessory branches always present at bases of ultimate branches; ratio of length to width of ultimate and accessory branches about 2:1; ultimate branches to 12 cm long and 7 cm wide, costules 3 mm apart, /amina very thin and strongly glaucous on the lower surface, quite glabrous; veins very slender, distinctly prominent on both surfaces. Type: J. Winkler 136, Batakerland, Sumatra (H. Selim Birger, S). Distr. Indo-China, Hainan, Kwangtung, in Malaysia: Sumatra. Ecol. Altitude 1250 m. Note. The specimens from China have ultimate branches proportionately rather narrower than those from Sumatra, but are otherwise similar. 12. var. montana HoLtr. Reinwardtia 4 (1957) 276.—Gleichenia linearis var. montana HOLTT. Rev. Fl. Mal. 2 (1955) 69, descr. angl.—Fig. 12b. Lateral branch-systems several times equally forked; ultimate branches 15-25 cm long, 314-6 cm wide; costules (5—)6-7 mm apart; /amina coriaceous, glabrous and glaucous on the lower surface, veins rather strongly prominent on both surfaces; accessory branches, about half as long as the ultimate branches, always present at the bases of ultimate branches. Type: Molesworth Allen 2720 (SING). Distr. Ceylon, S. India, Sikkim, in Malaysia: Sumatra, Malay Peninsula, Java, Borneo, and Moluccas (Ternate). Ecol. On the edges of forest and in clearings, with other varieties of D. Jinearis and with other members of the family; 1000-1600 m. 13. var. altissima Ho.Ltr. Reinwardtia 4 (1957) 276.—Gleichenia linearis var. altissima WHOLtr. Rev. Fl. Mal. 2 (1955) 69, descr. angl. Sometimes very high-climbing, the main rachis to 10 mm diameter; lateral branch-systems equally forked 4 or 5 times; ultimate branches to 15 cm long and 3 cm wide; costules 314-414 mm apart; /amina thin, lower surface glaucous, glabrous apart from scattered rusty hairs on lower surface of costules; veins prominent on upper surface, not on lower surface; accessory branches always present at bases of ultimate branches. Type: Corner 31447, Johore, Malay Peninsula (SING; dupl. at K, L). Distr. Malaysia: Malay Peninsula, Philip- pines (Palawan, Luzon), Moluccas (Talaud), New Guinea, and Solomon Islands. Ecol. At low altitudes, climbing on edges of forest. SCHIZAEACEAE (R. E. Holttum, Kew) Rhizome usually short-creeping with closely-placed fronds, less often wide- creeping or somewhat erect, the young parts covered with thick septate hairs (except Mohria, not Malaysian), structure dorsiventral or radial, vascular strand in Malaysian genera a protostele (medullated in Schizaea). Fronds of very varied structure, their branching showing varying gradations from dichotomous to pinnate; veins usually free; sporangia borne on specialized segments of the fronds (sorophores) except in the non-Malaysian Mohria. Sorophores at the ends of veins of fertile leaflets (Lygodium), or in small pinnate groups at the apex of a frond or of its branches (Schizaea), or confined to special branches of the frond (Anemia, not Malaysian). Sporangia arising marginally but becoming superficial due to subsequent extra-marginal growths, large, borne on short massive stalks or sessile, with an almost apical annulus of a single row of elongate thickened cells, dehiscing on a line from annulus to base. Spores trilete or monolete (Schizaea only), without perispore, the surface usually sculptured. Gametophytes filamentous in Schizaea, thalloid in other genera, symmetrical or not. Distribution. The Malaysian genera Schizaea and Lygodium are pantropic with a few outlying species of both in temperate regions (U.S.A., S. Africa, Chile, Japan, and New Zealand). Anemia has its main distribution in tropical America, with a few species in Africa and one in southern India. Mohria is confined to southern and eastern Africa and the Mascarene Islands. Fossils. The older fossils belong to extinct genera, which are quite different in frond-form from living members of the family; their relationship to these living members is shown only by their sporangia and spores. The Upper Carboniferous genus Senftenbergia has been most recently and fully described by RADFORTH (Trans. R. Soc. Edinb. 59, 1938, 385-396; ibid. 1939, 745-761). The fronds were bipinnate- tripinnatifid, comparable in size with those of Cyathea of today, with sterile and fertile parts alike in shape of lamina; the rhizome is not known. The sporangia, attached to the edge of the lamina by very short stalks, were similar in shape to those of the living genus Anemia but differed in having an annulus of 2-5 rows of cells, in at least one case not sharply differentiated from the rest of the sporangial wall. The Jurassic genus K/ukia had fronds of similar form, but the sporangia had an annulus of a single row of cells, much as in Anemia. C. F. REED has made the most recent summary of fossil forms in the family (Bol. Soc. Brot. 21, 1947, 71-197); he includes also the genera Tempskya, Acrostichopteris, Pelletiera, and Schizaeopsis, from Upper Jurassic and Cretaceous horizons, but sporangia of these have not been seen. Fossils of the living genera Anemia, Lygodium, and Schizaea have been found in late Cretaceous and early Tertiary rocks in Europe and North America (REED, /.c.; CHANDLER, Bull. Br. Mus. Nat. Hist., Geol., 2, 1955, 291-314; SELLING, Act. Hort. Gotob. 16, 1944, 1-112); fossils of Schizaea have also been found in Eocene of SE. Australia and Pliocene in New Guinea (CooKsoN, Proc. R. Soc. Victoria n.s. 69, 1957, 41). Fossils of Anemia and Lygodium include good leaf-impressions and sporangia; those of Schizaea are confined to spores. One species of Schizaea is also known from fossils of Quaternary age in Hawaii. Ecology. Lygodium plants are twining climbers, mainly in secondary vegetation (fig. 6, 10), producing fertile leaflets on parts exposed to the brightest light (in most cases, to full sunlight). Fronds of the larger species may climb to a height of 10 m, but others only to 2-3 m. In regions where there is a prolonged dry season the plants may be confined to wet ground. L. microphyllum, L. polystachyum and L. salici- folium have caducous leaflets which fall when old. Whether plants are ever quite bare of leaflets in dry seasons is not recorded; certainly the rhizomes have no power of resisting considerable drought, and most plants are evergreen. L. japonicum occurs outside Malaysia in decidedly seasonal climates, and in Malaysia does not occur in the uniformly wet region of Sumatra, Malaya and Borneo. L. polystachyum, native of the seasonal climate of the region from northern Malaya to Burma and Indochina, is only found in shady forest; possibly this is also true of L. merrillii. All species of Lygodium in Malaysia are lowland plants, the highest altitude record being little over 1000 m, apart from L. japonicum (to 2500 m). Schizaea plants are all small, and occur in poor acid sandy or peaty soils where the vegetation is rather open (fig. 1, 2), apart from S. digitata, which always occurs in light to moderate shade. S. fistulosa is a high mountain plant, S. malaccana on exposed ridges at moderate elevations, the other species in low country. S. inopinata occurs only on limestone. S. wagneri and S. spirophylla are reported as growing in moss-cushions on trees. Vegetative morphology. Both Lygodium and Schizaea are specialized, in quite different ways; 38 FLORA MALESIANA [ser. Il, volaae Fig. 1. Unusually dense stand of Schizaea dichotoma (L.) SM. on very poor sandy podsol, under shrubs near Pasir Pandjang, west coast of Borneo (DUNSELMAN, 1936). as above indicated, they are adapted to peculiar habitat-conditions. It is probably significant that neither is represented by fossils earlier than late Cretaceous or Eocene. The sorophores of both genera (and of Anemia) are perhaps reduced modifications of the lobes of fertile leaflets of ancestral ferns with fronds like those of Senftenbergia. The photosynthetic function of such lobes is transferred in both genera to a lamina developed from the wing of the rachis which occurs throughout the family (see further notes under the genera); such development is much more extensive in Lygodium than in Schizaea (for a discussion of transference of function, see CORNER in Journ. Linn. Soc. Bot. 56, 1958, 33-48). Fronds of Schizaea are either unbranched (the ultimate condition of reduction) or symmetrically dichotomously branched. Fronds of young plants of Lygodium are dichotomously branched, but later fronds develop an elongate sympodial climbing rachis by means of a series of alternate unequal dichoto- mies. The short branch at each dichotomy of a climbing rachis has a pair of opposite secondary branches beyond which it is dormant (HoLTTUM, Phytomorphology 7, 1957, 152). The dormant apices may grow if there is injury to a distal part of the frond. Similar, but not identical, dormancy occurs in the families Gleicheniaceae and Matoniaceae. Periodic dormancy of apices occurs in Dennstaedtia, Hypolepis and Paesia, which are perhaps derivatives of Schizaeaceae. PRANTL investigated the development of sorophores in Schizaeaceae, and stated that the sporangia were marginal in origin (Untersuch. z. Morph. der Gefasskrypt. II, 1881). Diets doubted PRANTL’s observations (in Engler, Pflanzenfam. 1, 4, 1900, 356-372), but later workers confirmed them (BOWER, The Ferns 2, 1926, 163-165). The lamina of the sorophores, and the separate indusia for each of the sporangia in Lygodium, are later extra-marginal developments. Sporangia and spores. The sporangia are the most distinctive feature of the family; they are well illustrated by PRANTL for all genera and his drawings have been frequently copied. The sporangia are large, almost or quite sessile, and all have an almost apical annulus with a longitudinal line of dehiscence. Those of Lygodium are peculiar in having an asymmetrically placed lateral attachment. The spores are in most cases rather elaborately sculptured on the surface, and are often specifically diagnostic (see further discussion under the genera). Gametophyte. The gametophyte of Schizaea is filamentous, with antheridia and archegonia on special short branches; some cells have an endophytic fungus. Other genera have more normal gameto- phytes, sometimes with asymmetric growth; their antheridia are large, and they show some other primitive features (for references up to 1926, see Bower, /.c. 170). Most observations of gametophytes have been of non-Malaysian species. Dec. 1959] | Schizaeaceae (Holttum) 39 Cytology. The only records are by MANTON & SLEDGE (Phil. Trans. R. Soc., B, 238, 1954, 142-143) and Lovis (Nature 181, 1958, 1085). Lygodium scandens (L. microphyllum of present work) in Ceylon had n = 30. L. circinnatum in Ceylon had n = 58, but a plant in cultivation at Kew had n = 29. A plant of L. japonicum at Kew also had n = 58. The basic number in Anemia is 38; a naturalized plant in Ceylon was tetraploid, a plant in cultivation at Kew diploid. Schizaea asperula WAkeF. in New Zealand had n = 77 (Lovis, /.c.); the same number was also found by Lovis (unpublished) for S. dichotoma in New Zealand. In Ceylon Lovis found that S. digitata had a very high number (n = 325 + 30). These figures indicate that polyploidy is not uncommon in the family, and that the extreme reduction of plant-form in Schizaea may be associated with high polyploidy. Anatomy. The fullest account of anatomy in the family is by PRANTL (J/.c.); later works are cited by Bower. The rhizome of Lygodium has a solid protostele; the rachis of the climbing frond has also a compact vascular strand (not C-shaped), with very large tracheids in the xylem, no doubt in adaptation to its habit (a slender twining rachis carrying many leaflets, which thus needs a vascular system of capacity large in proportion to its area of cross-section). The rhizome of Schizaea has a more or less medullated protostele, and the stipe has a small compact vascular strand in which the xylem is reduced, more or less 3-armed as seen in cross-section. In both genera there is considerable development of sclerenchyma. The genera Anemia and Mohria have a more complex vascular anatomy. Schizaea shows specialization in the arrangement of stomata, the details of this varying from species to species. Economic importance. The tough slender climbing rachises of Lygodium find various uses, either in their natural form, or prepared by splitting for finer purposes. They are used as a substitute for cord (e.g. for tying sheaves of rice), for plaiting into hats, bracelets, erc., for fastening the rims of sieves, and in other ways. There are records of the use of several species of Lygodium, and also of Schizaea dichotoma, for a great variety of medicinal purposes (see BURKILL, Dict. Econ. Prod. Mal. Pen. 1378, 1975; also HEYNE, Nutt. Pl. Ned. Ind. ed. 2, 96, 97), but no critical study of such uses has been made. Very young leaves of Lygodium microphyllum and L. circinnatum are eaten in Java (OCHsE, Veg. D.E.I. 1931, 655— 657). The plants are also used (especially Lygodium) in magical ceremonies connected with house-building, rice culture, fishing, efc. Taxonomy. The early history of the taxonomy of the genus Lygodium is very complex, for various reasons. LINNAEUS began badly by including references to three distinct species under his Ophioglossum scandens (the basis of Lygodium scandens Sw.); and he was unfortunate in having a poor specimen of a sterile frond of an immature plant on which to base his O. fexuosum. In the years immediately following 1800, several authors were independently studying specimens of Lygodium, and the following generic names were given: Lygodium Sw., Ugena Cav., Ramondia MirBeL, Odontopteris BERNH., Gisopteris BERNH. and Hydroglossum WILLD. (all in 1801), Cteisium Micux (1803) and Vallifilix THOUARS (1809). The following names were also given to Schizaea: Lophidium Ricu. (1792) and Ripidium BERNH. (1801). For a full bibliographic statement on these genera, and a discussion of their typification, see PICHI- SERMOLLI, Webbia 12 (1955) 4-36. In the case of Lygodium, there is so much variation in leaflet-form, due to (a) age of plant, (b) en- vironmental conditions, (c) height above ground from which specimen is taken, (and probably also to polyploidy and hybridization) that, even with ample material, it is not easy to define specific limits, and there was much confusion in the use of names by earlier authors. Different forms of the same species received different names, while in other cases two quite different species were confused under one name. WILLDENOw based his Hydroglossum pinnatifidum on two specimens, one sterile and one fertile, belonging to two quite distinct species, and his name, transferred to Lygodium, was subsequently used by different authors for both these species. For a detailed discussion of this subject, see ALSTON & Hoxrtrum, Reinwardtia 5 (1959) 11—22. The only good monograph of the whole family is that of PRANTL (/.c., 1881), whose very thorough morphological study (of material then available) laid a sound basis on which others, working on various aspects of the family and with new material, could build. Diets (in Engler, Pflanzenfam.) followed PRANTL with little alteration, as did also CHRISTENSEN (Index Filicum, 1905). As regards nomenclature, PRANTL did not look fully into the typification of L. scandens (L.) Sw. and he did not follow some of our modern rules. Also he did not have good material (in a few cases he had no material) of some of the less common Malaysian species. Nakal published a survey of the whole family in 1937 (J. Jap. Bot. 13, 139-154). He devided it into the three families Schizaeaceae, Lygodiaceae and Anemiaceae (including Mohria in the last) and also raised some infra-generic groups to generic rank. This process was carried further by C. F. REED (Bol. Soc. Brot. 21, 1947, 71-197) who raised the rank of the whole group to that of an Order, Schizaeales, with families for the fossil as well as living representatives. He separated Mohria as a family distinct from Anemia, and raised further subdivisions of Lygodium and Schizaea to generic rank, but made no critical contribution to the understanding of species, nor any new basic morphological study. COPELAND (Gen. Fil. 1947) made little change from CHRISTENSEN’S arrangement. KEY TO THE GENERA 1. Fronds simple and linear, or dichotomously branched with linear branches (which in some non- Malaysian species are joined laterally), the sorophores borne laterally near the apex of the frond 40 FLORA MALESIANA [ser. Il; volo or of its branches; no indusia. Spores monolete. . . . « 1. Schizaea 1. Fronds of young plants dichotomously branched, of older plants of indefinite growth with twining rachis, the rachis-branches variously branched and bearing leaflets which have sorophores at the ends of their veins; each sporangium protected by a separate indusium. Spores trilete 2. Lygodium 1. SCHIZAEA Sm. Mem. Ac. Turin 5 (1793) 419, nom. cons.—Lophidium Ricu. Act. Soc. Hist. Nat. Paris 1 (1792) 114.—Actinostachys WALL. ex Hook. Gen. Fil. (1842) t. 111; REED, Bol. Soc. Brot. 21 (1947) 130.—Microschizaea REED, /.c. 133.—Fig. 1-4. Rhizome creeping or suberect, the young parts, and bases of stipes, covered with coarse septate hairs, vascular system a medullated protostele. Stipes erect, slender, narrowly winged towards apex in most species. Frond simple or dicho- tomously branched, lamina reduced to a narrow wing bearing a single (rarely double) row of stomata on the lower surface; two-celled glandular hairs frequent on the surface of fronds, the basal cells persistent and often forming small warts, distal cells shrivelling or falling when old or dried. Sorophores pinnately arranged at the apex of a frond or of its branches, each sorophore with a median ridge on its lower (abaxial) surface, the sporangia attached to the sides of the ridge, the reflexed edge of the lamina protecting them. Sporangia not quite symmetrically ovoid or ellipsoid, sessile, with distal annulus of a single row of cells; spores pale, monolete, the surface variously sculptured. Distr. Pantropic, comprising c. 30 spp., widely distributed also in temperate regions of the southern hemisphere (S. Africa, Chile, New Zealand, Tasmania) but only in N. America in the northern hemisphere. Morph. PRANTL referred to the whole fertile apex of a frond of Schizaea (or of a branch of S. dicho- toma) as a sorophore; he called the lateral appendages /aciniae. But it seems probable that each lacinia is homologous with a sorophore of Lygodium, and the term sorophore is therefore here used for what PRANTL called a lacinia. Photosynthetic tissue in Schizaea is reduced to the wings on each side of the axis of a simple frond, or of the branches of S. dichotoma. The regular row of stomata associated with this photosynthetic tissue (a double row in S. inopinata) corresponds to a similar less regular row along each side of the stipe and rachis of many ferns. In Lygodium, there are irregularly scattered stomata on both sides of the narrow rachis-wings. Spores. SELLING published a very full account of the spores of all species known to him in 1944 (Medd. Géteb. Bot. Tradg. 16, 1-112, p. 1-5). He also discussed the later-known species of the group of S. digitata (Svensk Bot. Tidskr. 41, 1947, 431-450). Fossil spores are also known (see remarks on fossils, supra). KEY TO THE SPECIES ieeirondssrepeatediy dichotomous? ails) se) es) ee) eco tee atcle ele) Syne. ee 1. S. dichotoma 1. Fronds simple. 2. Sorophores 214-5 cm long, all attached close together at apex of frond; frond 2 mm or more wide. 3. Sporangia in four rows on the sorophores; one row of stomata on each wing of the frond. 2. S. digitata 3. Sporangia in two rows on sorophores; stomata in two rows on each wing of the frond. 3. S. inopinata 2. Sorophores much shorter, fronds narrower. 4. Edges of sorophores smooth and glabrous. 5. Fronds more than 1 mm wide, with distinct flat wing on each side of costa; no hairs with sporangia. 6. Costa very prominent on lower surface; stomata in each row very close together 2. S. digitata 6. Costa hardly prominent on lower surface; stomata in each row rather widely spaced. 4. S. spirophylla 5. Fronds less than 1 mm wide, wing very slightly developed; hairs present with sporangia. 5. S. wagneri 4. Edges of sorophores irregularly lobed, the lobes bearing coarse hairs. 7. Sorophores attached along the distal 10-20 mm of the axis of the frond; upper surface of frond deeply grooved, lower surface evenly rounded and bearing superficial rows of stomata. 6. S. fistulosa 7. Sorophores attached along the distal 5-10 mm of the axis of the frond; upper surface of frond slightly grooved, stomata intwo slight grooves on the distinctly flattened lower surface 7.S. malaccana Dec. 1959] Schizaeaceae (Holttum) ~— 41 1. Schizaea dichotoma (L.) SM. Mem. Ac. Turin 5 (798) 422. t.09; Buc Ens) Pls Javs (1828) 2555 Hook. & GrReV. Ic. Fil. (1827) t. 17; BEDp. Ferns S. India (1863) t. 65; Handb. (1883) 452; PRANTL, Unters. Morph. Gefasskr. 2 (1881) 138; Racis. Pterid. Buit. (1898) 6; v. A. v. R. Mal. Fig. 2. Schizaea dichotoma (L.) SM. near Kepa- hiang, Bencoolen, S. Sumatra (DE VooGD). Ferns (1908) 116; Merr. Int. Rumph. (1917) 69; BACKER & Postu. Varenfl. Java (1939) 256, fig. 66; Hoxttr. Ferns Mal. (1955) 50, fig. 6.— Acrostichum dichotomum LINNE, Sp. PI. (1753) 1068. —Osmunda dichotoma Spr. in Schrader, J. Bot. (1799) pt 2, 268.—Ripidium dichotomum BERNH. in Schrader, J. Bot. 1800, pt 2 (1801) 127, t. 2, f. 3.—S. forsteri Spr. Anleit. 3 (1804) 57.—S. cristata WILLD. Sp. Pl. 5 (1810) 88.—S. biroi RICHTER, Math. Termeszet. Ertesito 29 (1911) 1074; TROLL, Flora 128 (1933) 339, fig. 1.—S. copelandica RicuTeER, /.c.—Fig. 1, 2, 4a-d. Rhizome 3-6 cm below surface of ground, creeping, sometimes to 6 cm or more long, densely covered with coarse shining brown hairs 2-3 mm long. Stipes commonly 15-30 cm long (extremes 10-50 cm), narrowly winged towards apex; frond commonly 10-20 cm long and wide, dichoto- mously branched 2-8 times, the basal branches like the stipe, the distal ones gradually with wider wings and 1-114 mm wide, lacking a prominent costa on the lower surface; all parts with scattered small projections which are the bases of glandular hairs; sorophores occupying the distal 3-5 mm of each branch of the frond, 5-10 pairs, the lowest 3-4 mm long, upper ones smaller, edges hairy; sporangia in two rows, mixed with conspicuous long brown hairs; spores smooth or minutely granular. Type: Petiver, Gaz. t. 70, f. 12 (drawing of a specimen from Cochinchina). Distr. Mascarene Isl., Ceylon and S. India, Burma, Siam, Indochina, throughout Malaysia except for East Java and Lesser Sunda Isl., to Tahiti, Australia, and New Zealand. Ecol. In lightly shaded places, or sometimes in forest, often (always?) in sandy ground, sealevel to 1000 m, rarely abundant. Note. Small plants like those named S. biroi and S. copelandica by RICHTER are not uncommon, and have been found at many places near larger plants. As pointed out by TROLL (/.c.) the small little-branched fronds of these small plants are usually fertile, whereas some fronds of much larger plants are sometimes sterile. I have however examined a very large number of specimens and have failed to find any sharp distinction between those with little-branched and much-branched fronds. If one picks out individual specimens, one can separate fronds with long and with short ultimate branches; but in some cases fronds from one collection may show ultimate branches of very diverse length. Vern. Tatagar payong, Kedayan, pirangas, Murut, oemiar, biak, E. New Guinea, paku tjakar ajam, radja hantu, Banka, rumput bulu merak, Billiton, silaju, Sum. Uses. HEYNE records medicinal use (in Billiton) for coughs and affections of the throat and also in childbirth. 2. Schizaea digitata (L.) Sw. Syn. Fil. (1806) 150, 380, t. 4 f. 1; Bu. En. Pl. Jav. (1828) 255; Bepp. Ferns S. India (1864) t. 268; Handb. (1883) 452; PRANTL, Unters. Morph. Gefasskr. 2 (1881) 133, t. 5 f. 83; CLARKE, Trans. Linn. Soc. Bot. 1 (1880) 583; Racts. Pterid. Buit. (1898) 7: v. A. v. R. Mal. Ferns (1908) 116; BACKER & PostH. Varenfl. Java (1939) 257, f. 66; SELLING, Svensk. Bot. Tidskr. 41 (1947) 431-450; Ho tr. Ferns Mal. (1955) 51, f. 7.—Acrostichum digitatum LINneE, Sp. Pl. (1753) 1068.—Actinostachys digitata WALL. ex REED, Bol. Soc. Brot. 21 (1947) 130.— Fig. 3a-e. 42 FLORA MALESIANA [ser. Il; vole Rhizome very short, creeping or suberect, 3-4 cm below surface of ground, bearing many crowded fronds, apex clothed with brown hairs under 2 mm long. Fronds erect, unbranched, grass-like, 20-35 cm long, base (stipe) slender and triquetrous, rest winged, greatest width 24 mm, r, Fig. 3. Schizaea digitata (L.) Sw. a. Habit, x \4, b. sorophores, nat. size, c. detail of soro- phore, x 5, d. spore, x 300, e. detail of lower surface showing stomata, x 5.—S. inopinata SELLING. f. Detail of sorophore, x 18, g. spore, x 300, A. flattened part of exospore, x 200, i. lower surface of leaf, showing stomata, x 5 (a, e For. Bot. Burma 7670, bB-c MATTHEW s.n. (SING), f SYNGE S606, g SYMINGTON CF 37414, h after SELLING 1946). costa very prominent on lower surface of winged portion and slightly grooved on upper surface, 2- celled glandular hairs abundant on lower surface of wing, stomata in a close even single row on each side of costa. Sorophores all attached very close together (apparently digitate), 5-18, all about equal in length, commonly 214-5 cm long (on stunted plants sometimes shorter), little over 1 mm wide, edges thin, entire, glabrous; sporangia small, apparently in four rows and completely covering lower surface of sorophore; spores small, finely and evenly obliquely striate. Type: Herb. Hermann, Ceylon (BM). Distr. Ceylon, NE. India, Siam, Indochina, Micronesia; in Malaysia: throughout except for East Java and Lesser Sunda Isl. Ecol. In lightly shaded forest, rubber estates, etc., sealevel to 1200 m, rarely very abundant. Notes. The species was formerly credited with a much wider distribution, due to confusion with other species. SELLING has distinguished most of the latter, and has given a comparative survey of the group and of individual distributions (/.c.). Actinostachys boninensis NAKAI (J. Jap. Bot. 13, 1937, 140) appears to differ from S. digitata only in the greater number of sorophores (to 30) which are shorter (8-40 mm long); I have seen no specimens. 3. Schizaea inopinata SELLING, Svensk Bot. Tidskr. 40 (1946) 274, f. 1-7; Hottr. Ferns Mal. (1955) 52.—Actinostachys inopinata REED, Bol. Soc. Brot. 21 (1947) 130.—Fig. 3f-i. In habit like S. digitata, differing as follows: fronds to 244 mm wide, with a double (occasional- ly triple) row of stomata on each side of the costa on the lower surface, the wings thicker and more rigid and the edges reflexed on drying, the costa not so strongly raised and rather variable; sporangia much larger, in two rows; spores much larger, with broad irregular longitudinal ridges. Type: Henderson 19460, Gua Tipus, Chigar Perah, Pahang (SING, K). Distr. Micronesia, in Malaysia: Sumatra, Malaya, Borneo, Philippines (Bohol), W. New Guinea. Ecol. On limestone crags, sealevel to 300 m. Note. FosBerG, correctly reporting the oc- currence of the species in Micronesia, considered it conspecific with S. ponapensis HOSOKAWA (Am. Fern J. 40, 1950, 145). I have examined the isotype of S. ponapensis in the Arnold Arboretum Herbarium, and find that, apart from its much smaller size (fronds to 8 cm long, sorophores to 8 mm long) it differs in having a hardly raised costa with a rather widely-spaced single row of stomata oneach side of it. The specimen corresponds well with the description of S. spirophylla TROLL. and I have placed S. ponapensis as a synonym of that species. I saw no spores. 4. Schizaea spirophylla TRoLL, Flora 128 (1937) 343, fig. 2-6.—S. ponapensis HOSOKAWA, Trans. Nat. Hist. Soc. Formosa 31 (1941) 39. Rhizome short, apex covered with dark hairs. Fronds unbranched, sometimes twisted, 4-8 cm mec; 1959] | Schizaeaceae (Holttum) 43 Fig. 4. Schizaea dichotoma (L.) SM. a. Habit, x ?/3, b. rhizome, x 2/3, ¢. sorophores, x 7, d. lower surface of leaf, x 14.—S. wagneri SELLING. e. Habit, x 4/,, f. detail of sorophore, x 14, g. lower sur- face of leaf, x 27.—S. malaccana BAKER. h. Sorophores, x */;, i. lower surface of leaf showing stomata, x 27.—S. fistulosa LaBILL. j. Sorophores, X 4/3, k. detail of sorophore, x 7, /. lower surface of leaf, showing stomata, x 27 (a Hose 199, b BRAss 8583, c-d FMS 199, e-g GRETHER & WAGNER 4177, h-i SFN 1100, j-7 CLEMENS 10729). long, about 114 mm wide, costa very slightly rows (sometimes apparently in 4 rows near the prominent on the lower surface, with a distinct middle); spores as in S. digitata. flat wing on each side of it; stomata in one row Type: Troll, Ambon (M). on each side of the costa, rather widely spaced Distr. Micronesia (Ponape); in Malaysia: in the rows; sorophores 1-3, 4-8 mm long (some- Moluccas (Ambon). times longer?), glabrous, the sporangia in two Ecol. Growing in moss-cushions on trees. 44 FLORA MALESIANA [ser. I, volosi4 Note. See note under S. inopinata. The twisting of the fronds of S. spirophylla is probably not a constant character; such twisting is also common, but not universal, in S. digitata. Stunted plants of the latter may superficially resemble S. spiro- phylia but differ in costa and stomata and in more crowded sporangia. 5. Schizaea wagneri SELLING, Svensk Bot. Tidskr. 40 (1946) 278, f. 8-11; Hottr. Ferns Mal. (1955) 52.—Actinostachys wagneri REED, Bol. Soc. Brot. 21 (1947) 131.—S. paucijuga Hoitr. Gard. Bull. Sing. 11 (1947) 267.—Fig. 4e-g. Rhizome short, apex clothed with slender brown hairs 114 mm long. Fronds simple, 6—20 cm long, base terete, upper part winged and in all 14-34 mm wide, with rather broad midrib prominent on the lower surface, the stomata rather widely spaced in one row close to each side of the midrib. Sorophores 2-5, 7-15 mm long, edges smooth and glabrous, sporangia in two rows, mixed with brown hairs; spores finely verrucose. Type: Grether & Wagner 4177, summit of Mt Tjajiak, 600 m, Manus Isl., Admiralty Is. (S-PA, dupl. at K). Distr. Malaysia: Malaya (P. Rumbia in Perak, Singapore) Borneo, Ambon, W. New Guinea, and Admiralty Is. Ecol. “Epiphytic in mosses on stumps and bases of trees; abundant at one place, but ex- ceedingly inconspicuous” (GRETHER & WAGNER). In forest near sea (Western New Guinea and Borneo). 6. Schizaea fistulosa LABILL. Novy. Holl. Pl. Spec. 2 (1806) 103, t. 250 f. 3; PRANTL, Unters. Morph. Gefasskr. 2 (1881) 135, excl. var. malaccana and var. robusta; v. A. v. R. Mal. Ferns (1908) 116; C. Cur. & Hott. Gard. Bull. S. S. 7 (1934) 210.— Microschizaea fistulosa REED, Bol. Soc. Brot. 21 (1947) 134.—S. propinqua A. CUNN. in Hook. Comp. Bot. Mag. 2 (1836) 362.—S. australis Gaupb. Ann. Sc. Nat. Bot. 5 (1825) 98.—S. chilensis Putt. Linnaea 30 (1859-60) 207.—Fig. 4j-l. Rhizome short-creeping, young parts clothed with shining brown hairs 2-3 mm long; stipes very crowded; fronds unbranched, 9-18 (rarely to 30) cm long below the fertile part, width to about 1 mm, upper surface rather deeply grooved, lower surface almost evenly rounded and bearing two rows of stomata which are not in grooves; surfaces bearing scattered glandular hairs the bases of which are slightly prominent. Sorophores all about equal, arranged in a distinctly pinnate manner along the distal 10-20 mm of the axis of the frond, 8-20 pairs, lowest often forked, 4-6 mm long, edges much reflexed and bearing many coarse forward-pointing hairs; sporangia in two rows, without hairs; spores smooth. Type: Labillardiére, Australia (F?; not seen). Distr. Madagascar, Australia, Tasmania, New Zealand, Fiji, New Caledonia, Chile; in Malaysia: Borneo (Mt Kinabalu), New Guinea. Ecol. In alpine bogs at 2400-3750 m, and on fine rock-screes. Note. Some New Guinea specimens have been distributed as S. papuana BRAUSE, which is here placed as a synonym of S. malaccana. 7. Schizaea malaccana BAK. Syn. Fil. (1868) 428; BEDDOME, Ferns Br. India (1870) t. 255; Handb. (1883) 452; TANsLEY & Cuick, Ann. Bot. 17 (1903) 493-510; v. A. v. R. Mal. Ferns (1908) 116; Hoxttr. Ferns Mal. (1955) 52, fig. 8.—S. fistulosa var. malaccana PRANTL, Unters. Morph. Gefasskr. 2 (1881) 136.—S.papuana BRAUSE, Bot. Jahrb. 56 (1920) 211.—Microschizaea malaccana REED, Bol. Soc. Brot. 21 (1947) 134.—Fig. 4h-i. var. malaccana. Habit of S. fistulosa, differing as follows: hairs on rhizome pale brown, fronds 6-15 cm long, less than 1 mm wide, apical fertile part of axis c. 5mm long, upper surface of frond nearly flat or shal- lowly grooved, lower surface when dry with two small grooves in which the stomata are situated; sorophores 4-10 pairs, lowest 4-5 mm long, upper ones shorter. Type: Cuming 379, Mt Ophir, Malaya (K, BM). Distr. Malaysia: Malaya, Borneo, Ambon, W. New Guinea. Ecol. In open mossy places on mountain ridges, or in moss cushions on trees at 800-2000 m; also in swamp forest in Sarawak at lower altitudes. var. robustior C. Cur. Gard. Bull. S.S. 7 (1934) 210.—S. hallieri RicHTER, Med. Rijksherb. n. 28 (1916) 24, t. 1 f. 5, etc.; Math. Naturw. Ber. Ungarn 31 (1916) 24, 28, t. 1 f. 5 etce.—Mi- croschizaea hallieri REED, Bol. Soc. Brot. 21 (1947) 134. Larger than var. malaccana; fronds to 25 cm long, 1 mm or more wide; sorophores to 15 pairs, spread along 10 mm of the axis, lowest sorophores 5-10 mm long. Type: Clemens 10919, Mt Kinabalu, Borneo (BM, Bo). Distr. Malaysia: Malaya (Gunong Tahan), Borneo, W. New Guinea. 2. LYGODIUM Swartz in Schrader, J. Bot. 1800 pt 2 (Nov.—Dec. 1801) 106, nom. cons.; PRANTL, Unters. Morph. Gefasskr. 2 (1881) 60; CopELAND, Gen. Fil. (1947) 23; PICHI- SERMOLLI, Webbia 12 (1956) 10, preprint (1955).—Ramondia MIRBEL, Bull. Soc. Philom. Paris 2 (Feb.—Mar. 1801) 179.—Ugena Cavan. Ic. Descr. Pl. 6 (Oct. 1801) 73.—Odontopteris BERNH. in Schrader, J. Bot. 1800 pt 2 (Nov.—Dec. 1801) 127, t. 2 f. 4.—Ripidium BERNH. /.c. 127, t. 2 f. 3.—Gisopteris BERNH. /.c. 129, Dec. 1959] Schizaeaceae (Holttum) 45 t. 2 f. 1—Hydroglossum WitLp. Abh. Kurfiirstl. Mainz. Ak. Niitzl. Wiss. Erfurt 2, pt 4 (1802) 13, 20.—Hugona Cav. ex Roemer, Arch. Bot. 2 (1801-02) 486.— Cteisium Micux, Fl. Bor. Am. 2 (1803) 275.—Vallifilix THouARS, Gen. Nov. Madag. (1808) 1.—Lygodictyon J. SMitH in Hook. Gen. Fil. (1842) t. 111 B.— Fig. 5-15. Rhizome creeping, below ground surface, protostelic, short with fronds very close together or longer with spaced fronds, young parts densely covered with rather thick rigid multiseptate hairs, branching dichotomous; fronds borne in two rows on upper surface of rhizome, roots mainly from lower surface. Fronds of young plants erect, once or twice dichotomously branched and bearing usually palmately lobed leaflets; fronds of older plants with slender elongate twining rachises formed by a succession of very unequal dichotomies, at least the upper part of the rachis (except in L. polystachyum) bearing two narrow wings towards the adaxial side, the surface between the wings flat or slightly raised and papillose; all branch rachises and stalks of leaflets similarly winged (fig. 13d), the wings always interrupted to join with those of a lateral branch; primary rachis-branches always short, usually hardly developed, ending in dormant apices which are covered with hairs (such apices proliferous if the main rachis beyond them is injured), each primary branch bearing a pair of secondary branches which bear the leaflets; sterile /eaflets (or their lobes) with costa and oblique lateral veins which are 1-3 times forked (anastomosing in a few species), edges entire or serrate (pinnatifid only in L. polystachyum); fertile leaflets often with contracted lamina, bearing narrow sorophores spreading from the edges of the lamina at the ends of most of the veins; edges of sorophores serrate, the main vein in each sorophore bearing alternate short lateral veins each of which bears a single sporangium protected by a separate indusium attached along the vein and opening forwards; sporangia oblong-ovoid with a short lateral stalk, the annulus at the narrower end which is directed away from the margin of the sorophore, splitting longitudinally when ripe; spores trilete, pale, variously sculptured on the surface, lacking perispore. Gametophyte thalloid, sometimes asymmetric; antheridia larger and more complex in structure than in most leptosporangiate ferns. Distr. Pantropic, comprising c. 40 spp., also extratropical southwards in New Zealand and S. Africa, northwards in Japan and in eastern U.S.A. to Massachusetts. Morph. Owing to the peculiar structure of the climbing leaves, it is difficult to apply the usual descrip- tive terms pinna and pinnule to them, especially where the branching of the leafy parts is dichotomous. The following terms are here used. The climbing rachis of the frond (sympodial in structure but for convenience considered as a unit) bears alternate short primary branches, each ending in a dormant apex and bearing a pair of apparently opposite secondary branches. The secondary branches may bear leaflets or tertiary branches pinnately arranged, or they may be once or more times dichotomously branched. The narrow wings on rachis-branches of all orders, those of the ultimate branches joined to the edge of the lamina, correspond with the wings which are the only lamina in fronds of Schizaea, but do not have the very regular single rows of stomata found in Schizaea (there are irregularly scattered stomata on both sides of a wing in Lygodium). If the sorophores of Lygodium are regarded as homologous with the fertile leaflet-lobes of the paleozoic fossil genus Senftenbergia, the lamina of Lygodium may be con- sidered as a specialized development of the rachis-wing consequent on the reduction of the original lamina-lobes to sorophores. The dormant apices of the primary rachis-branches are covered with septate hairs. In a group of Malaysian species these hairs have swollen bases, each base formed of a mass of cells; these species are L. borneense, L. longifolium, L. auriculatum, L. trifurcatum, and L. dimorphum. So far as I know, this type of hair has not hitherto been reported in Lygodium. Taxon. There are four pairs of species which in some measure intergrade. These need experimental study in cultivation to discover how much variation is due to environmental conditions, and also cyto- 46 FLORA MALESIANA [ser. Il, vole logical study. It may be that natural hybridization occurs, and as tetraploids as well as diploids have been already discovered in L. japonicum and L. circinnatum they may occur also in other species, leading to the possibility of the formation of sterile triploids; apogamy has however not yet been discovered in the genus. The pairs of species which intergrade are: L. flexuosum and L. japonicum; L. flexuosum and L. salicifolium; L. borneense and L. auriculatum; L. dimorphum and L. trifurcatum. The following species are very distinct: L. polystacnyum, L. microphyllum, L. circinnatum, L. merrillii, and L. versteegii. KEY TO THE SPECIES 1. Secondary rachis-branches pinnate, leaflets 10-15 on each side. Sterile leaflets evenly pinnatifid throughout 1. L. polystachyum 1. Secondary rachis- branches pinnate with fewer leaflets, or r dichotomous. Sterile leaflets simple, palmate, or lobed near the base only. 2. Venation of sterile leaflets free. 3. Primary rachis-branches 4-10 mm long below the pair of secondary branches. Rhizome wide- creeping, fronds distinctly spaced. 4. Secondary branches simply pinnate. Leaflets articulate at the base and ultimately deciduous. 4. Secondary branches amply bipinnate. Leaflets not articulate at the base 2. L. microphyllum 3. L. japonicum 3. Primary rachis-branches hardly elongated, the pair of secondary branches thus almost sessile on the main rachis. Rhizome short, fronds close together. 5. Secondary rachis-branches regularly pinnate, normally with 3-5 leaflets on each side of the axis (young or depauperate fronds may have fewer leaflets). 6. Leaflets all about equal and all stalked (terminal one sometimes geminate), not auricled or branched at the base, or rarely with short spreading basal lobes; their junction with the lamina . leaflet-stalks thickened at 4. L. salicifolium 6. Lateral leaflets larger towards base of secondary branch, smaller distal ones sessile, basal ones stalked and usually auricled or with obliquely genau lobes, or with subsidiary leaflets below them; leaflet-stalks not thickened at apex . 5. L. flexuosum 5. Secondary branches simple or dichotomously branched one or more times (in some cases, by alternate unequal dichotomy, subpinnate with 1-3 lateral leaflets). Leaflets simple, forked, or palmately branched, never thickened at apex of stalk. 7. Leaflets strongly cordate on outer side at base. 8. Lamina of fertile leaflets reduced to a narrow wing along midrib and bases of veins, each vein usually with an apical sorophore. 9. Fertile tertiary branches evenly deltoid in outline with quaternary leaflets of increasing length towards the base. Sterile leaflets strongly auriculate-cordate at base . 6. L. dimorphum 9. Fertile leaflets not deltoid in outline, 10-15 mm wide (including sorophores), sometimes with 1 or 2 long branches at the base. Sterile leaflets more or less cordate at the base. 7. L. trifurcatum 8. Lamina of fertile leaflets not reduced to a narrow wing along midrib and veins. 8. L. auriculatum 7. Leaflets not strongly cordate at base on outer side. 10. Secondary branches once forked (rarely simple, or one branch forked again). Leaflets always simple. Spores smooth 9. L. borneense 10. Secondary branches at least twice forked (or Tower branches bearing single 3—5-lobed leaflets), sometimes sub-pinnate with 1—2 tertiary members on each side and a terminal leaflet or pair of leaflets. Leaflets often deeply 2-lobed to palmate. Spores verrucose. 11. Margins of sterile leaflets serrate, not much thickened, veins ending in the teeth. Spores coarsely and irregularly verrucose. Dormant apices of primary branches not sunk, hairs with swollen base 10. L. longifolium 11. Margins of sterile leaflets entire, thickened but not ‘vascular, the raised veins joining the thickened edge. Spores finely and evenly verrucose. Dormant apices sunk, hairs lacking swollen bases : 2. Venation of sterile leaflets retienlate: 11. L. circinnatum 12. Secondary rachis-branches (at least the upper ones) elongate and pinnate. 13. Leaflets on stalks to 3 cm long, palmately lobed or equally bilobed (basal secondary branches may bear only one palmately lobed leaflet on a longer stalk) 13. Leaflets on much shorter stalks, simple or the apical ones forked 12. L. merrillii 14. L. reticulatum 12. Secondary branches bearing 3 leaflets (lateral ones sometimes forked), all arising 1-3 mm from dormant apex of primary rachis-branch 1. Lygodium polystachyum WALL. ex Moore, Gard. Hook. Sec. Cent. (1861) t. 76; Ferns Br: India (1868) t. 300; Handb. (1883) 458, f. 284; Chron. (1859) 671; Syn. Fil. (1868) 438; BEDDOME, 13. L. versteegii v. A. v. R. Mal. Ferns (1908) 113; Copev. Philip. J. ‘Sc. Bot. 4 (1909) 19; Tarp. & Go Car El: Gén. I.-C. 7 (1939) 40; Hottr. Ferns Mal. (1955) 56, f. 10; ALSTON & HoLtTrT. Reinwardtia 5 (1959) Dec. 1959] Schizaeaceae (Holttum) 47 11. — Hydroglossum pinnatifidum WiLLD. Abh. Kurf. Mainz. Ak. Wiss. Erfurt 2, pt 4 (1802) 21, p.p.—L. pinnatifidum (non (WILLD.) Sw.) PRANTL, Unters. Morph. Gefasskr. 2 (1881) 83, t. 1 f. 11.—Fig. 5c, 8a-—c. Fig. 5. Spores of Lygodium. a. L. longifolium (WILLD.) Sw., b. L. borneense v. A. v. R., c. L. polystachyum WALL. ex Moore, d. L. circinnatum (Burm. f.) Sw., e-f. L. microphyllum (CaAv.) R. Br., lateral and upper view. All x 330. Rhizome short-creeping, 3-5 mm _ diameter, densely clothed with spreading black hairs. Juvenile fronds once or twice dichotomous; stipes up to 30 cm long to the first dichotomy, brown, withshort very slender hairs mixed with thick longer multicellular ones; ultimate branches leafy like the secondary rachis-branches of climbing fronds. Rachis of climbing fronds 214 mm dia- meter, shortly hairy, not winged; primary rachis- branches very short, ending in a dormant apex covered with brown hairs (the apices of lower primary branches sometimes proliferous); second- ary rachis-branches 20-30 cm long, not winged, shortly hairy, bearing 10-15 leaflets on each side and a similar terminal leaflet; sterile leaflets 3144-714 cm long, 114-2 cm wide, apex rather abruptly narrowed and rounded, base truncate or cordate, jointed to a hairy stalk 1-4 mm long, sides lobed half-way to the costa or rather more, lobes 4-5 mm wide, oblong with rounded apex, each lobe with a sinuous costule bearing oblique forked lateral veins, costa and costules bearing scattered stiff hairs on both surfaces, the costa also with shorter hairs; fertile leaflets like the sterile but the distal half or more of each lobe narrowed to about 2 mm wide, the narrow part (sorophore) 4-10 mm long, bearing sporangia on the under surface; indusia bearing scattered stiff hairs; spores finely and evenly verrucose. Type: Wallich 177, Penang (K). Distr. Assam, Burma, Siam, Yunnan, Indo- china; in Malaysia: northern half of Malay Peninsula. Ecol. In lowland forest, climbing trees to a considerable height; in the Malay Peninsula especially on limestone except in the extreme north (Kedah) where it is locally common apparently primary forest. (Further sor limestone provides a drier habitat than ott rocks, and on it occur a number of species whic have their main distribution in the seasona. climate north of Malaya.) 2. Lygodium microphyllum (CaAv.) R. Br. Prod. FI. Nov. Holl. (1810) 162; Br. En. Pl. Jav. (1828) 253; CLARKE, Trans. Linn. Soc. Bot. 1 (1880) 583; Bepp. Handb. (1883) 455, t. 282; ALSTON & Hott. Reinwardtia 5 (1959) 12.—Ugena micro- phyllatGAve ice Descra Elon sO) a76: sty 5955 C. Cur. Dansk Bot. Ark. 9, n. 3 (1937) 30.—L. scandens Sw. in Schrader, J. Bot. 1800 pt 2 (1801) 106, p.p. excl. syn. LINN.; BEDD. Ferns S. India (1863) t. 61; Hoox. Syn. Fil. (1868) 437, p.p.; PRANTL, Unters. Morph. Gefasskr. 2 (1881) 81, t. 6 f. 101; CHrRist, Farnkr. Erde (1897) 354, f. 1116; Racts. Pterid. Buit. (1898) 8; v. A. v. R. Mal. Ferns (1908) 113; Philip. J. Sc. 11 (1916) Bot. 116; HEyNne, Nutt. Pl. (1927) 97; BAck. Krakatoa (1929) 254; OcHsE & BAKH. Veg. DIE-E 9s) "6o7- Burke Dict. 2) C1935)" 1378: BACKER & PostTH. Varenfl. Java (1939) 258; OGatTaA, Ic. Fil. Jap. 7 (1936) t. 324; Hott. Ferns Mal. (1955) 58, f. 12.—Ophioglossum filiforme Roxs. Calc. J. Nat. Hist. 4 (1844) 476, t. 26 f. 3.—L. scandens var. microphyllum (CAV.) Luerss. J. Mus. Godeffr. 6 (1874) 4.—L. scandens var. intermedium Ces. Att. Ac. Sc. Fis. Nat. Napoli 7 (1876) 33.—Fig. 5e-f, 6, 7. Rhizome wide-creeping, dichotomously branch- ed, 2!14 mm diameter, densely clothed with short spreading brownish-black hairs. Juvenile fronds small, commonly once dichotomous (the stipe distinctly winged below the dichotomy), each branch bearing a 4-lobed leaflet not jointed at the base, lobes 3—5 cm long and c. 5 mm wide, thin, glabrous, edges crenately toothed (teeth larger towards apex where veins are unbranched). Rachis of climbing fronds glabrous, commonly 2-3 m long, hardly 114 mm diameter; primary branches 4 mm or more long, ending in a dormant apex covered with dark brown hairs; secondary rachis-branches pinnate, in all to about 15 cm 48 FLORA MALESIANA [ser. IT, volxaes Kee RIT AES Ant are Uy: eee ts Fig. 6. Lygodium microphyllum (Cav.) R. Br. and L. salicifolium PR. (larger leaflets on left) on edge ofa thicket in the Botanic Gardens, Singapore (HOLTTUM). Dec. 1959] Fill Q Schizaeaceae (Holttum) 49 Fig. 7. Lygodium microphyllum (Cav.) R. Br. a. Fertile leaf, x %4, b. sterile leaf, x 34, c. rhizome, x 34, d. detail of base of leaflet, x 7 (a FloyD NGF 5566, b-d cult. Kew from Ceylon). long, with 3-6 stalked leaflets on each side (stalks 2-4 mm long) and a similar or geminate terminal leaflet; /eaflets quite glabrous, mostly ovate (sterile leaflets often elongate with broader base on young plants), 1-4 cm long (sterile ones sometimes to 6 cm), 6-18 mm wide, edges of sterile ones minutely crenate, a joint always present at base of blade, where the wing which in other species connects stalk and lamina is constricted; fertile leaflets usually shorter than sterile but with lamina hardly narrowed, soro- phores 4-6 mm long; spores witha raised reticulum on the outer surface. Type: Née, Luzon (Ma). Distr. Tropical Africa, SE. Asia (north to Bengal and Hong Kong and the Riu Kiu Isl.), Melanesia (Solomon Isl., New Caledonia), N. and E. Australia south to N. S. Wales; in Malaysia: throughout, but few records from the Lesser Sunda Isl. (certainly to Flores). 50 FLORA MALESIANA [ser. II, vol. 1} VV. SS Wd me We Ny SEA t) a ERS ‘S ot ie Seay Y% : ee, Wy : ee NWMWn, * ea UN BSNS Ak ae LL py Wn Zo Bigs SSN “any Su MONA Sy Se NTN), Lys Fig. 8. Lygodium polystachyum WALL. ex Moore. a. Sterile leaf, x #/;, b. fertile leaflet, x */;, c. ditto, detail, x 214.—L. japonicum (THUNB.) Sw. d. Fertile leaf, x #/;, e. sterile leaf, x 3/,, f. rhizome, x 4/5 (a BALANSA 168, b-c MATTHEW s.n., d, f cult. Kew, e SAVINIERRE 71). Dec. 1959] Schizaeaceae (Holttum) . 51 Ecol. Edges of secondary forest, or climbing woody plants in open places, sometimes as a weed; in clay soil, or in swamps in regions subject to a dry season; from the lowlands to c. 1300 m. Vern. Ribu-ribu, M, sélada, capey papua, capay alus, M, paku kawat, Sum., paku hata béjas (bias), paku hata leutik, S, nitong-puti, Tag., paku rambat, Bali, sickey, Luhea, gomoha papua, Ternate, paku kawa, Ambon, paloge, N. Celebes. Uses. Native medicine (leaves macerated and mixed with lime for open wound), magic; young leaves edible; rachises of old leaves used for string and for plaiting. 3. Lygodium japonicum (THUNB.) Sw. in Schrader, J. Bot. 1800, pt 2 (1801) 106; Bepp. Ferns S. India (1863) 21, t. 64: CLARKE, Trans. Linn. Soc. Bot. | (1880) 584; PRANTL, Unters. Morph. Gefasskr. 2 (1881) 68, t. 1 f. 10, 15; CHrist, Farnkr. Erde (1897) 355, 356, f. 1122; Racis. Pterid. Buit. (1898) 8; Hore, J. Bomb. Nat. Hist. Soc. 15 (1903) 106; v. A. v. R. Mal. Ferns (1908) 114; Merk. FI. Manila (1912) 60; Doin, Bibl. Bot. 20, Heft 85 (1914) 211, f. 50; HaAtnes, Bot. Bih. & Or. 6 (1924) 1210; OcarTa, Ic. Fil. Jap. 7 (1936) t. 322; BACKER & PostH. Varenfl. Java (1939) 259; ALSTON & HoLtr. Reinwardtia 5 (1959) 14.— Ophioglossum japonicum THUNB. Fl. Jap. (1784) 328.—Hydroglossum japonicum (THUNB.) WILLD. Abh. Kurf. Mainz. Ak. Wiss. Erfurt 2, pt 4 (1802) 26.—L. dissectum Desv. Mag. Ges, Naturf. Fr. Berl. 5 (1811) 308.—L. microstachyum Desvy. lI.c.; NAKAI, Bot. Mag. Tokyo 39 (1925) 182; OGaTa, Ic. Fil. Jap. 7 (1936) t. 323.—L. pubescens KAULF. En. Fil. (1824) 47, t. 1 f. 4.—L. chaerophylloides Desv. Mém. Soc. Linn. Paris 6 (1827) 205.—L. cochinchinense Desy. ibid. 206.—L. tenue BL. En. Pl. Jav. (1828) 254.—L. microphyllum Link, Hort. Berol. 2 (1833) 141.—L. japonicum f. elongata v. A. v. R. Bull. Jard. Bot. Btzg II, n. 1 (1911) 10, t. 3; Mal. Ferns Suppl. (1917) 117.— L. japonicum var. microstachya (DESV.) TARD. & C. Cur. Fl. Gén. I.-C. 7 (1939) 38.—L. mearnsii CopEL. Philip. J. Sc. 3 (1908) Bot. 37.—Fig. 8d-f. Rhizome wide-creeping, dichotomously branch- ed, 2-5 mm diameter, densely clothed with dark brown hairs, fronds commonly 5-10 mm apart. Juvenile fronds erect, the first branching an unequal dichotomy (always?), the two main branches of large fronds bipinnate, deltoid in outline, with palmatisect leaflets, their margins doubly serrate. Rachis of climbing fronds hardly 2 mm diameter, glabrous apart from minute hairs on the flattened adaxial surface between the Narrow wings; primary rachis-branches 3-10 mm long, the dormant apex covered with pale hairs; secondary branches of fronds on young or stunted plants pinnate, on well-grown fronds bipinnate or tripinnate, deltoid in outline, commonly 12 cm long and wide, rachises densely short-hairy on the upper surface and bearing fewer longer hairs elsewhere; sterile tertiary /eaflets of lower rachis- branches palmate with 5—7 lobes, the middle lobe much longer than the laterals, tertiary leaflets higher up the leaf trilobed with an elongate middle lobe or pinnate with small oblique and often lobed quaternary leaflets and a usually deltoid-pinnatisect terminal leaflet about 3 cm long, edges acutely biserrate, apex obtuse or subacute; stalks of leaflets to 3 mm long, never articulate or thickened at the apex; costae usually bearing long scattered hairs, veins and surfaces usually glabrous but sometimes short- hairy; fertile secondary branches tripinnate, the leaflets smaller than sterile ones, sorophores 2-12 mm long; indusia glabrous or with a few hairs if the lamina is hairy; spores finely verrucose. Type: Herb. Thunberg, Japan (Ups). Distr. Ceylon, from Himalayas (Kashmir eastwards) to Chekiang in N. China, Korea, Japan (Nagasaki) and southwards to Siam and Indochina and southern China, naturalized in Florida and Texas; in Malaysia: Banka, Central and East Java, Celebes, Philippines, Moluccas (Ambon, Ternate, Banda Is!., Sula Isl.), Lesser Sunda Isl. (to Timor), and New Guinea. Ecol. Climbing in secondary vegetation, at altitudes up to 2550 m; only found native in regions with a pronounced dry season, during which fronds perhaps die (no records in Malaysia); absent from Sumatra, Borneo and the Malay Peninsula (except perhaps the extreme north). Small forms are not always clearly distinct from L. flexuosum. Vern. Pakis kémbang, J, paku areuj, hata kawat, S, nito, nitong puti, Tag., madik silai, durhawa, babar, talsiga, Alor. 4. Lygodium salicifolium Pres_, Suppl. Pterid. (1845) 102, p.p. excl. pl. WALLICH p.p. and syn. RHEEDE; PRANTL, Unters. Morph. Gefadsskr. 2 (1881) 79; v. A. v. R. Mal. Ferns (1908) 113; Bull. Jard. Bot. Btzg III, 5 (1922) 213; BACKER & Postu. Varenfl. Java (1939) 258; TARD. &. C. Cur. FI. Gén. I.-C. 7, 2 (1939) 41; ALSTON & Ho.ttr. Reinwardtia 5 (1959) 14.—L. kingii CopEL. Philip. J. Sc. 6 (1911) Bot. 68; v. A. v. R. Mal. Ferns Suppl. (1917) 117.—L. pinnatifidum sensu RAc. Fl. Buit. (1898) 7, p.p.—L. flexuosum sensu Hott. Ferns Mal. (1955) 57, p.p.—Fig. 6, 10, 13a—b. Rhizome and juvenile fronds as in L. flexuosum, except that the rachis is thickened at its junction with the midribs of the lobes of the leaflets. Rachis of climbing fronds to 2 mm diameter, to 10 m long; primary rachis-branches always very short (hardly measurable), ending in a dormant apex covered with brown hairs; secondary rachis- branches normally pinnate, rarely somewhat bipinnate and then the tertiary branches bearing one or more pairs of short spreading lateral leaflets (jointed at the base) and a large terminal one; secondary branch-system usually consisting of about 4 (rarely to 6) leaflets on each side, and a terminal deeply bilobed leaflet (or a pair of leaflets), all leaflets of about equal size and all stalked, the stalks 2-10 mm long and thickened at junction with lamina (old leaflets sometimes deciduous but not regularly so as in L. micro- phyllum); leaflets 4- 15 cmlong, 14-2 cm wide, acute 2 FLORA MALESIANA [ser. Tl, voliat= Fig. 9. Lygodium dimorphum Cork. a. Part of fertile leaf, x %, b-c. parts of sterile leaves, x 3%, d. detail of a, x 3.—L. flexuosum (L.) Sw. e. Sterile leaf, x %, f. fertile leaf, x % (@ LABILLARDIERE s.n., b-c CARR 12479, d PEEKEL 6, e Hose 5024, f HALLER s.n. from Java). Dec. 1959] 7 Schizaeaceae (Holttum) 53 and attenuate or subobtuse, edges of sterile leaflets finely crenate-serrate, base truncate to cordate, lamina thicker than in L. flexuosum; upper surface of costae more or less hairy es- pecially towards the base, lower surface often glabrous, veins usually glabrous; sorophores 2-5 mm long, usually constricted at the base, often with hairs on upper surface of midrib; indusia glabrous; spores finely verrucose. Fig. 10. ‘Columns’ of Lygodium salicifolium PR. on scattered pole trees of mostly Ilex cymosa, surrounded by a dense ground cover of ferns and sedges, Nepenthes, and orchids, in an old crater swamp in the Gajo Lands, N. Sumatra, c. 1200 m (1937). Type: Cuming 365, Singapore (W, K). Distr. Assam, Siam, Indochina to Yunnan, Formosa, south-east to New Guinea and Micro- nesia; in Malaysia: Sumatra, Malay Peninsula, Banka, W. Java, Borneo, and New Guinea. Ecol. In open secondary vegetation, sometimes in wet places, in the low country and to 1200 m; reported in teak forest in West Java. Note. There are specimens which are inter- mediate between this species and L. flexuosum; they may be hybrids. L. salicifolium occurs only in regions with a short dry season, whereas L. flexuosum will tolerate a longer dry season and has a wider distribution. In Burma and Assam very large forms of both species occur. Vern. Hata, S, mintuh, Dayak, paku kawat, Sum., akar sidin, M. 5. Lygodium flexuosum (L.) Sw. in Schrader, J. Bot. 1800, pt 2 (1801) 106; ibid. 1801, pt 2 (1802) 304; Pres_, Suppl. Pterid. (1845) 100; Bepp. Ferns S. India (1863) t. 63; CLARKE, Trans. Linn. Soc. Bot. 1 (1880) 584; PRANTL, Unters. Morph. Gefasskr. 2 (1881) 72, p.p.; Bepp. Handb. (1883) 457, f. 283; v. A. v. R. Mal. Ferns (1908) 114; Merr. Fl. Manila (1912) 61; Domin, Bibl. Bot. 20, Heft 85 (1914) 209, f. 49; Haines, Bot. Bih. & Or. 6 (1924) 1211; Back. Onkr. Suiker 7 (1928) 1, t. 1; Burk. Dict. 2 (1935) 1378: "C."Cur: Dansk Bot. ‘Ark: 9, ‘pt 3 (1937) 30; BACKER & PosTH. Varenfl. Java (1939) 259; Hott. Ferns Mal. (1955) 57, p.p.; ALSTON & Ho.ttr. Reinwardtia 5 (1959) 15.—Ophioglossum flexuosum LINNE, Sp. Pl. (1753) 1063.—Ophio- glossum scandens LINNE, Sp. Pl. (1753) 1063, p.p. —Ramondia flexuosa (L.) Mire. Bull. Soc. Philom. Paris 2 (Feb.—Mar. 1800) 179, t. 12 f. 3.—Aydro- glossum flexuosum (L.) WILLD. Abh. Kurf. Mainz. Ak. Wiss. Erfurt 2 (1802) 23, t. 1 f. 3.—Hydro- glossum pinnatifidum WiLLD. ibid. 21, p.p.—L. pinnatifidum Sw. in Schrader, J. Bot. 1801, pt 2 (1803) 303; Hook. Syn. Fil. (1868) 438, p.p.— L. semibipinnatum R. Br. Prod. Fl. Nov. Holl. (1810) 162.—L. serrulatum Bi. En. Pl. Jav. (1828) 254.—L. flexuosum var. setulosum TARD. & C. Cur. Fl. Gén. I.-C. 7 (1939) 39.—Fig. 9e-f. Rhizome short-creeping and densely covered with roots, the stipes very close together; apex of rhizome covered with dark brown to nearly black hairs. Juvenile fronds once or twice dichotomous, each branch bearing a single leaflet which is deeply palmately 3-7-lobed, the lobes almost equal, the base of the whole leaflet cordate, edges serrate and sometimes crenately lobed. Rachis of scandent fronds narrowly winged, flattened and puberulous on the upper surface between the wings; primary rachis-branches up to 3 mm long (lower ones longest), dormant apex covered with pale brown hairs; secondary rachis-branches pinnate to somewhat bipinnate, narrowly ovate to deltoid in outline, commonly about 15 cm long and 8 cm wide; sterile leaflets of lower branches palmate, often 5-lobed, base strongly cordate; higher secondary branches bearing 3-5 (some- times to 7) leaflets on each side and an apical one, the apical and lower leaflets asymmetric or more or less lobed at the base, the lowest often with 2 or 3 (exceptionally to 6) separate quaternary leaflets at its base; sterile leaflets 3-10 cm long, 8-15 mm wide above the lobed base, apex sub- acute, edges serrate, lower leaflets stalked, upper sessile, lamina rather thin; costae usually bearing scattered long hairs, less often densely short- hairy, veins often with scattered short hairs on the lower surface, the lamina sometimes similarly hairy; fertile leaflets smaller than sterile, soro- phores 3-5 mm long (rarely up to 10 mm), at the apices of small triangular lobes; indusia glabrous or with a few hairs like those of the lower surface of the lamina; spores finely verrucose. 54 FLORA MALESIANA [ser. TL vole Type: Hermann, Ceylon (BM). Distr. Ceylon, from the Himalayas (Dehra Dun eastwards) to southern China, Hong Kong, Riu Kiu Isl., south and south-east to Melanesia and northern Queensland, throughout Malaysia. Ecol. In open places, climbing on shrubs, in teak and bamboo forest, in low country and to 1000 m, not in shady evergreen forest. In very dry or exposed places the veins and lamina are often rather copiously hairy. Vern. Ribu-ribu gajah, ribu-ribu bésar, ikat sidin, M, paku ribu-ribu, Asahan, hata kémbang, J, durhawa, Alor, nito, Tag., Visc., tatan, Orokawa Horata (N.G.), zangi, Orokawa Mumuni. Uses. For tying rice sheaves; in native medicine for skin diseases and fever. 6. Lygodium dimorphum Cope-. Philip. J. Sc. 6 (1911) Bot. 67; RosENsT. in Fedde, Rep. 10 (1912) 343; v. A. v. R. Mal. Ferns Suppl. (1917) 116; Philip. J. Sc. 11 (1916) Bot. 116; ALSTON & Hoitr. Reinwardtia 5 (1959) 18.—L. flexuo- sum [non (L.) Sw.] GAuUDICH. in Freyc. Voy. Bot. (1826) 298.—L. circinnatum var. trifurcatum Curist, Monsunia | (1900) 93.—L. novoguineense RosensTt. in Fedde, Rep. 9 (1911) 427.—L. trifurcatum sensu v. A. v. R. Mal. Ferns (1908) 112, 802, p.p.—Fig. 9a-d. Rhizome short-creeping, bearing fronds very close together, apex densely covered with almost black hairs. Juvenile fronds once dichotomous, each branch bearing a deeply palmatisect leaflet, lobes subequal, base cordate, edges rather ir- regularly serrate and somewhat thickened. Rachis of climbing fronds hardly 2 mm diameter, glabrous or nearly so; primary rachis-branches very short, ending in a somewhat projecting dormant bud covered with light brown hairs having slightly swollen bases; secondary rachis-branches bearing sterile leaflets unbranched or more commonly once dichotomous, those bearing fertile leaflets usually sub-pinnate with a few tertiary branches; sterile leaflets 10-18 cm long, simple or forked (less often 3-lobed), usually strongly cordate and auriculate at the base on one side (sometimes with a separate rounded leaflet replacing the cordate base), when forked the lamina lobed to within 1 cm of the base, leaflets or lobes 1-2 cm wide, tapering, acute, irregularly doubly serrate, edge somewhat thicken- ed, surfaces glabrous, sometimes sparingly warty; fertile leaflets usually with the lamina reduced to a narrow wing (0.2 mm wide) along the costae and along each vein and its branches; tertiary fertile branches deltoid in outline (5 cm or more wide at the base), with quaternary leaf- lets of increasing length below each terminal one; sorophores 2-4 mm long, indusia glabrous; spores minutely verrucose on an unevenly undulating surface (always?). Type: C. King 134, Papua (Micu, Bo). Distr. Malaysia: Celebes(?), Moluccas(Ambon, Rawak), New Guinea. Ecol. “Climbing small trees to a height of about 12 feet. Quite common in low wet places along the coast’? (RUSSELL, on specimen from E. New Guinea). Collections have also been made inland, at altitudes up to 1000 m. Vern. Cana, Motuan, gailei, Bragi, paku kawa, Ambon. Uses. Used for making arm- and leg-bands. 7. Lygodium trifurcatum BAK. in Hook. Syn. Fil. (1868) 437; v. A. v. R. Mal. Ferns (1908) 112, 802, p.p.; WAGNER & GRETHER, Un. Cal. Publ. Bot. 23 (1948) 27, t. 8; ALSTON & HoLtTrT. Rein- wardtia 5 (1959) 17. Scandent rachis about 114 mm diameter; primary rachis-branches very short, hairs on the dormant apex having swollen bases; sterile secondary rachis-branches once or twice equally or sub- equally dichotomous (12-20 mm to first dicho- tomy), the leaflets simple or very deeply bilobed, 10-20 cm long, 1!4-2'4 cm wide, the outer base of each more or less cordate (rarely strongly auriculate), edges irregularly serrate and slightly thickened; fertile secondary branches usually sub-pinnate with flexuous axis bearing two lateral and one terminal leaflets, sometimes twice symmetrically dichotomous; fertile leaflets usually deeply bilobed or geminate, one or both members often having a shorter lobe commonly 9-15 cm long and up to 13 mm wide including sorophores, lamina reduced to a wing along the costa and along each vein-group (sometimes two adjacent vein-groups with a common lamina) the soro- Phores thus in groups of 2—5; sorophores usually 2-3 mm long; spores coarsely and unevenly verrucose, the warts often confluent. Type: Milne 511, Solomon Isl. (K). Distr. Melanesia (Solomon Isl. and New Hebrides); in Malaysia: Admiralty Isl., Louisiades. Ecol. ‘Climbing abundantly in brackish marsh” (GRETHER & WAGNER 3997, Admiralty Islands). Note. The specimen of GRETHER & WAGNER has fertile branches more like those of L. dimor- phum than is usual in L. trifurcatum, and has one branch intermediate between sterile and fertile; the fully sterile leaflets are smaller than normal in L. dimorphum and only slightly cordate at the base. The two species are very closely allied, and it may be that in the Admiralty Islands, where their areas of distribution overlap, there has been hybridization. 8. Lygodium auriculatum (WILLD.) ALSTON, Rein- wardtia 5 (1959) 16.—Ugena semihastata Cav. Ic. Descr. Pl. 6 (1801) 74, t. 594, nomen. illegit., excl. syn. REICHARD & RuMPH.; C. B. Ros. Philip. J. Sc. 6 (1911) Bot. 97; C. Cur. Dansk Bot. Ark. 9, pt. 3 (1937) 29.—Hydroglossum auriculatum WILLD. Sp. Pl. 5 (1810) 84.—L. semi- hastatum Dresv. Mém. Soc. Linn. Paris 6 (1827) 203, nom. illeg.; Hoox. Syn. Fil. (1868) 437; v. A. v. R. Mal. Ferns (1908) 111, excl. pl. bor- neensis; Mal. Ferns Suppl. (1917) 115; C. Cur. Ind. Fil. Suppl. I (1913) 119.—L. circinnatum var. semihastatum Foss. Am. Fern J. 40 (1941) 142.—L. flexuosum [non (L.) Sw.] PRANTL, Unters. Morph. Gefasskr. (1881) 73, p.p. quoad syn. Cav. and WILLD.—Fig. 11. Dec. 1959] Schizaeaceae (Holttum) g 55 *s oe ay Fig. 11. Lygodium auriculatum (WILLD.) ALSTON. a. Habit, x 24, b. hair, x 65 (a-b LE Roy TopPING 1287). Rhizome short-creeping, bearing fronds close together, its apex and bases of stipes densely covered with dark hairs. Juvenile fronds once dichotomous, each branch bearing a palmatisect leaflet, usually 5-lobed, with truncate base, edges closely and irregularly serrate. Rachis of climbing fronds hardly 2 mm diameter, usually glabrous; primary rachis-branches very short, dormant apex covered with pale brown hairs having swollen bases; secondary rachis-branches rarely bearing a simple leaflet, most commonly once dichotomous, one branch with a simple, one with a forked leaflet, less often each branch with a simple leaflet; sterile leaflets 12-20 cm long, 12-30 mm wide, subacute, edges not thickened, very shallowly serrate, base usually asymmetric and strongly cordate-auriculate on the outer, rarely on both sides, costae glabrous except near the base on upper surface; lamina of fertile leaflets 12-20 mm (rarely to 30 mm) wide, sorophores 3-9 mm long, constricted at the base, at the apices of short triangular lobes of the lamina; indusia glabrous; spores irregularly warty, variable as between different specimens, in some cases resembling those of L. /ongifolium, in others with many smaller warts of variable size. Type: Née, Luzon (Ma). Distr. Indochina, Micronesia; in Malaysia: E. Borneo, Philippines (Polillo, Luzon, Mindoro, Samar. Mindanao), to 600 m altitude. Note. The Micronesian specimens seen (from Guam) are all smaller than Philippine ones, and their secondary branches are regularly twice dichotomous; they may constitute a distinct local race (specimens of Née from Marianas not seen). It may be that this species intergrades with L. borneense in Borneo. Vern. Nito, Tag. Uses. Climbing rachises used for weaving, making hats, and magic (contra-poison) bracelets. 9. Lygodium borneense v. A. v. R. Bull. Jard. Bot. Btzg Il, m. 20 (1915) 29; Mal. Ferns Suppl. (1917) 115; CopeL. Sarawak Mus. J. 2 (1917) 303; Hottr. J. R. As. Soc. Mal. Br. 6 (1928) 16 with fig.; Ferns Mal. (1955) 56.—L. semi- hastatum sensu v. A. v. R. Mal. Ferns (1908) 111, p.p. quoad pl. borneenses.—L. borneense if samarindae v. A. v. R. Bull. Jard. Bot. Btzg II, n. 20 (1915) 29; Mal. Ferns Suppl. (1917) 116.—Fig. 5b, 13c-e. Rachis of scandent fronds glabrous, up to 2 mm diameter; primary rachis-branches very short, dormant apex covered with pale hairs having swollen bases; secondary rachis-branches rarely unbranched, normally once dichotomous (10-20 mm long below the dichotomy), each branch 56 FLORA MALESIANA [ser. Il, -voleat= ALL Me aaa: , Z\ WTMIT EIS TT ry sO : EVD wd ey ww att Oo ae ei ay Pe * x) K ¢ rn) SLL UTR \ Drprea Fig. 12. Lygodium longifolium (WILLD.) Sw. a. Habit, fertile, x 2/3, b. sterile, < 2/3, c. leaf detail showing edge, x 3 (a KiNG's coll. 259, 6 MoTLEy s.n., c Hose 5034). Dec. 1959] Schizaeaceae (Holttum) Fig. 13. Lygodium salicifolium PR. a. Habit, x 2/3, b. detail, « 10.—L. borneense v. A. v. R. c. Habit, x #/,, d. detail of branching, x 21/,, e. ditto, x 4/3 (@ BERKHOUT s.n., b BRASS 5686, c SFN 18656, d-e ENDERT 2057). with a simple leaflet or rarely one leaflet double, rarely the secondary branch-system sub-pinnate with two lateral and one terminal leaflets; sterile leaflets 20-35 cm long (on upper parts of frond smaller), 314-5 cm wide, margin more or less distinctly serrulate and slightly thickened, base cuneate (rarely cordate on the outer base), surfaces quite glabrous except the upper surface of the costa towards its base, lamina not verrucose when dry; fertile leaflets similar to sterile but slightly smaller, 10-30 cm long, 12-40 mm wide (smallest on upper branches), sorophores 4-10 mm long, usually about 2 mm apart, at the apices of small triangular projections of the lamina; spores quite smooth. Type: Teysmann, Borneo (Bo). Distr. Malaysia: Malaya (SE. Johore only), Sumatra (Mentawai Isl.), Borneo (many localities), Talaud Isl. Ecol. In light places in freshwater swamp- forest; in Sarawak twice reported in the neigh- bourhood of limestone hills but certainly occurring also not in the vicinity of limestone; one specimen from Sarawak, found growing with Jmperata on sandy ground, has unusually small leaflets. Note. This species has a branching habit closely similar to that of ZL. auriculatum. L. borneense usually differs from L. auriculatum in the larger size and cuneate base of its leaflets, but some Bornean specimens seem to be intermediate in 58 FLORA MALESIANA [ser. Il, volar = er x ae * Vy ey Em ZAI AY q i DAIWA. STII IIL SM Fig. 14. Lygodium circinnatum (BurRM. f.) Sw. a. Part of sterile leaf, x 24, b-c. parts of fertile leaves, showing different kinds of branching, x 24, d. veins of sterile leaf, x 3, e. ditto, leaf edge, x 9 (a, d-e CLEMENS 9487, b ZOLLINGER 169, c Lady DALHOUSIE s.n.). Dec. 1959] the latter character. So far as observed, L. bor- neense seems to be consistent in its smooth spores. 10. Lygodium longifolium (WILLD.) Sw. in Schra- der, J. Bot. 1801, pt 2 (1803) 305; ALston & Ho tr. Reinwardtia 5 (1959) 19.—Hydroglossum longifolium WiLLp. Abh. Kurf. Mainz. Ak. Wiss. Erfurt 2, pt 4 (1802) 22, t. 2.—L. digitatum PRESL, Rel. Haenk. 1 (1825) 73 (?); v. A. v. R. Mal. Ferns (1908) 112; Hoitr. Ferns Mal. (1955) 55.— L. dichotomum (non (CAv.) Sw.) BEepDpb. Ferns S. India (1863) t. 62.—L. teysmannii v. A. v. R. Bull. Dép. Agr. Ind. Néerl. 18 (1908) 5; Mal. Ferns (1908) 111, 801; Mal. Ferns Suppl. (1917) 115.—L. circinnatum var. cristatum vy. A. v. R. Bull. Dép. Agr. Ind. Néerl. 7. 18 (1908) 5; Mal. Ferns (1908) 112, 802.—L. derivatum v. A. v. R. Bull. Jard. Bot. Btzg III, 5 (1922) 213.— Fig. 5a, 12. Rhizome short-creeping, its apex and bases of stipes covered with shining black hairs. Juvenile fronds once or twice dichotomous, leaflets palma- tely divided with 4~7 subequal lobes, base often more or less cordate, lobes to about 18 cm long and 18 mm wide, acuminate, edges shallowly serrate, a vein ending in each tooth. Scandent frond to about 4 m long, rachis to 2 mm diameter; primary rachis-branches very short, with a dor- mant apex covered with brown hairs having small swollen bases; secondary rachis-branches 1-3 times dichotomous or sub-pinnate (lowest ones sometimes unbranched and bearing large 6-lobed leaflets); sterile leaflets composed of 2-4 subequal lobes 15 cm or more long and c. 15 mm wide, the sinuses between the lobes reaching to 15 mm from the base of the leaflet, edges regularly ser- rate, not or little thickened, base cuneate to cordate, surfaces glabrous and usually not warty when dried; fertile secondary branches \-3 times dichotomous or (if the dichotomies are unequal) more or less distinctly pinnate with two dicho- tomous tertiary branches (tertiary branches may rarely have three separate leaflets); fertile leaflets simple or more usually consisting of two subequal lobes united at the base, lamina 3-10 mm wide, sorophores commonly 2—3 mm long, less often to 6 mm; spores coarsely and irregularly verru- cose. Type: Herb. Willdenow, Malabar (B). Distr. Southern India, Hainan; in Malaysia: Malaya, Riouw and Lingga Islands, Sumatra, Borneo, Luzon (doubtful). Ecol. Edges of forest, probably in more ex- posed places than L. circinnatum, and not at- taining so large a size as that species. 11. Lygodium circinnatum (BurRM. f.) Sw. Syn. Bil= (1806) 1533 Br. Eno Plt Jay. (1828) 253: PRANTL, Unters. Morph. Gefasskr. 2 (1881) 64; BEDD. Handb. (1883) 455; v. A. v. R. Mal. Ferns (1908) 111; C. B. Ros. Philip. J. Sc. 6 (1911) Bot. 102; v. A. v. R. Philip. J. Sc. 11 (1916) Bot. 116; Merr. Int. Rumph. (1917) 69; v. A. v. R. Mal. Ferns Suppl. (1917) 115; W. H. Brown, Bull. Bur. For. Philip. n. 19 (1919) t. 4; ibid. Schizaeaceae (Holttum) 59 n. 22 (1920) 328, pl. IV; HEYNE, Nutt. Pl. (1927) 96; Back. Krakatoa (1929) 253; Burk. Dict. 2 (1935) 1378; BACKER & PostH. Varenfl. Java (1939) 258, f. 67; HoLtr. Ferns Mal. (1955)55, f.9; ALsTon & HoLtr. Reinwardtia 5 (1959) 20. —Ophioglossum circinnatum Burm. f. Fl. Ind. (1768) 228.—Ophioglossum pedatum Burm. f. ibid. 227, t. 66 f. 1. —Ugena dichotoma Cav. Ic. Descr. Pl. 6 (1801) 74, t. 594 f. 2; C. Cur. Dansk Bot. Ark. 9, pt 3 (1937) 30.—L. pedatum (BuURM. f.) Sw. Syn. Fil. (1806) 154; Merr. Philip. J. Sc. 19 (1921) 336.—Hydroglossum circinnatum (BURM. f.) WiLtLp. Abh. Kurf. Mainz. Ak. Wiss. Erfurt 2, pt 4 (1802) 24.—Hydroglossum pedatum (BurM. f.) WILLD. ibid. 25.—L. dichotomum (Cav.) Sw. Syn. Fil. (1806) 154; Hook. & GREV. omni 183i) Sd HOOK. ec BAK SYyt. bil (1868) 437; Racts. Pterid. Buit. (1898) 8.—Ophio- glossum furcatum Roxs. Calc. J. Nat. Hist. 4 (1844) 478.—L. basilanicum Curist, Philip. J. Sc. 2 (1907) Bot. 179; v. A. v. R. Mal. Ferns (1908) 802.—L. circinnatum var. monstruosum v. A. v. R. Bull. Dép. Agr. Ind. Néerl. n. 18 (1908) 5; Mal. Ferns (1908) 112; Mal. Ferns Suppl. (1917) 115.—Fig. 5d, 14. Rhizome short-creeping, bearing stipes very close together, its apex and bases of stipes densely covered with black hairs. Juvenile fronds once dichotomous, each branch bearing a _ pedato- palmatisect leaflet, lobes usually 4 or 5, subequal, to about 25 by 3!4 cm, the midrib of an outer lobe arising near base of the next inner lobe, edges entire, often somewhat crisped, pale and much thickened (translucent when living), apices acute to acuminate, surfaces glabrous but conspicuously warty when dry (not when living), veins uniting with the thickened margin. Rachis of climbing frond to about 10 m long, 2-5 mm diameter, glabrous; primary rachis-branches very short, with sunken dormant apex covered with pale hairs which are not thickened at the base; second- ary rachis-branches unbranched and 2-6 cm long, or once dichotomous with each branch 1-2 cm long beyond the fork; sterile leaflets usually with 2-6 subequal diverging lobes which are separate to within 2 cm from the base, entire, margin pale and thickened, base cuneate or truncate, surfaces nearly always warty when dry; fertile secondary rachis-branches unbranched or 1—3 times dichotomous (rarely sub-pinnate); fertile leaflets usually sessile in pairs at the ends of the ultimate branches, or members of a pair partly fused at the base, less often 3—5S-lobed (always so if the secondary rachis is unbranched), lamina more or less reduced and commonly 3-6 mm wide, rarely less than 2 mm or approximating in width to the sterile leaflet-lobes; sorophores 2-5 mm long, sessile; spores finely and evenly ver- rucose. Type: Java, herb. Burman (G, not seen). Distr. Ceylon, NE. India to southern China, Siam and Nicobar Isl. to Micronesia, the New Hebrides and Solomons; throughout Malaysia. Ecol. In lightly shaded places in primary or secondary forest, in the lowlands and to 1500 m 60 FLORA MALESIANA [ser. IT, voleais (Oe "58 Aha —— Ann. Rep. Brit. N. Guinea 1897-8 (1898) 149, nomen]; Kew Bull. (1899) 122; SADEBECK in E. & P. Pfl. Fam. 1, 4 (1901) 776; PFEIFFER, Ann. Mo. Bot. Gard. 9(1922) 211. Submerged aquatic. Stock 3—4-lobed. Leaves numerous, 5-714 cm by 3 mm, recurved, terete towards the apex, flattened lower down, abruptly dilated at base. Dilated base hyaline, c. 1 cm long and broad. Upper part of leaves dark green. Stomata few. Ligule broadly cordate. Velum none. Sporangia oblong, 6 by 4 mm, pale brown. Megaspores c. 800 w in diam., deeply and ir- regularly warted and reticulate on the outer face, warted on the inner faces, with a strongly marked triradiate marking. Distr. Malaysia: E. New Guinea (Mts Scratch- ley and Albert Edward), twice found. Ecol. Shallows of an alpine lake, 3000-3680 m. Note. BAKER’S statement that the megaspores are smooth between the triradiate ridges is incorrect. CYATHEACEAE (R. E. Holttum, Kew) Caudex massive, usually erect and unbranched, where prostrate not dorsiventral in structure; fronds arranged on caudex in spiral series; vascular system of caudex a hollow cylinder with gaps corresponding with leaf-bases, in some cases small medullary bundles also present; a cylinder of very hard sclerenchyma, with gaps at leaf-bases, present both inside and outside the vascular cylinder (but absent in Cibotium), the surfaces of the sclerenchyma covered with cubical cells containing silica; tangentially arranged sieve-tubes present in the phloem as well as longitudi- nal ones. Stipes of Cyathea containing numerous small vascular strands arranged in 3 series (fig. 6), these strands more or less united in smaller axes of Cyathea- fronds and also in larger axes of other genera (fig. 31f, 33d); stipe-bases per- sistent, or sooner or later caducous leaving a pattern of scars on the caudex; pneumathodes present along each side of stipe, in a single discontinuous or almost continuous row, or in 2—3 rows close together, the row joining upwards to a similar row on the basiscopic side of the first pinna, a + circular pneumathode at the base of the pinna beginning the row on the main rachis to the next pinna. Dermal appendages on fronds: multiseptate hairs only, or both hairs and scales (Cyathea); if both, the hairs often confined to the adaxial surface of the fronds. Fronds in most cases bipinnate-tripinnatifid, with varying gradations to tripinnate, in a few cases simply pinnate, in Cu/cita 3-4-pinnate; pinnules almost symmetrical at the base except in Culcita; veins normally free except in Cyathea capitata and in the genus Cnemidaria (trop. America). Sori either terminal on veins and protected by an inner indusium as well as by the more or less reflexed edges of a small lobe of the lamina (outer indusium), or apparently not terminal on veins and not near the edge of the lamina, with indusia of various form or without indusia; receptacle of various shape, in all cases containing vascular tissue which in the case of Cyathea represents the termination of a short vein; stalks of sporangia short or long, 4 or more cells in transverse section, annulus more or less oblique, with a more or less clearly defined lateral stomium; spores trilete, surfaces smooth or variously sculptured; multiseptate paraphyses, of a single row of cells (terminal cell glandular or not) or scale-like at the base, present with sporangia. Distribution. Throughout the wetter parts of the tropics, especially on mountains; a few species just north of the tropics, more south of the tropics especially in Australasia. As here construed, 9 genera, of which 5 are Malaysian: Cyathea (pantropic, at least 600 spp.); Cnemidaria (limited to species with simply pinnate fronds, anastomosing veins and distinctive spores, tropical America, 10 spp.); Lophosoria (tropical America, monotypic); Dicksonia (tropics and southern subtropics in Malaysia, Australasia, America, St Helena, c. 25 spp.); Cystodium (Malaysia, monotypic); Thyrsopteris (Juan Fernandez, monotypic); Culcita (subg. Culcita in Azores and tropical America; subg. Calochlaena in Malaysia and Australasia; in all c. 7 spp.); Cibotium (SE. Asia, Malaysia, Hawaii, Central America, c. 12 spp.); Metaxya (tropical S. America, monotypic). Fossils. SEwARD gave a summary of knowledge to 1920 (Fossil Plants 2, 365-375). T. M. HARRIS has recently published a fully illustrated account of some Jurassic frond-fossils which he includes in the family Dicksoniaceae (The Yorkshire Jurassic Flora, 1, 1961, 140-181), referring them to the genera Dicksonia, Coniopteris and Kylikipteris. Owing to the fragmentary nature of the fossils it is very difficult to judge how they compare with existing ferns. In my judgement, the fossil most like living Dicksonia is Coniopteris hymenophylloides (BRONGN.) SEWARD; C. murrayana BRONGN. is perhaps more like Culcita. The Jurassic fossils most resembling Cyathea in form of sterile leaflets are placed in the genus Kylikip- teris; their fertile leaflets have sori at the ends of veins on a reduced lamina, and seem more like those of Thyrsopteris than Dicksonia. Kylikipteris looks like a possible Cyathea-ancestor. Though no fossils with Cyathea-like sori have been found in the Yorkshire Jurassic, HARRIS described a genus Aspidistes which has sori and sporangia resembling Dryopteris or Thelypteris, spores trilete (known in a few species of (65) [ser. II, vol. 1? , Papua, 2840 m (L. J. Brass, 1933). 1v1ision FLORA MALESIANA o =I = = se) oO a o = o oO As) n o a. £ n 2. vo oO 2 nan qi cS) oD AS 3 ° [a jo!) x se O O ba S N ~ 8 0 ue) q a re se O ides C 10 athea gleichen Gy, Murray Pass, Wharton Range, Central D Fig Dec. 1963] iy ¥ CYATHEACEAE (Holttum) ne : 67 Fig. 2. Cyathea gleichenioides C. CHR., same locality as fig. 1. Trunks have been blackened by a recent grass fire (L. J. Brass). Thelypteris but not in Dryopteris) and spherical unicellular glands (common in Thelypteris). Aspidistes looks like an early Thelypteris, and Thelypteris has several features in common with Cyathea; but one would have expected Cyathea to have existed prior to Thelypteris. Fossil tree-fern trunks of Lower Cretaceous have also been called Coniopteris; they have leaf-scars and vascular system comparable with those of Cyathea and Dicksonia. OGURA described fossil tree-ferns from Upper Jurassic and Cretaceous rocks of Japan and Korea (J. Fac. Sc. Univ. Tokyo III, 1, 1927, 351-380, pl. 2-8). BANCROFT, describing a fossil Cyatheoid stem from the late Tertiary of East Africa (New Phytol. 31, 1932, 241-253) pointed out that OGuRA’s fossils differed in some respects from existing Cyatheaceae. K. JAcos described impres- sions of parts of a tree-fern stem from middle Jurassic of NE. India, but the pattern of vascular strands in the leaf-scars is not clearly preserved. He gives references to other descriptions of fossils of presumed Cyatheaceous affinity (Proc. Ind. Ac. Sc. 6, sect. B, 1938, 73-90). Ecology. Most species are forest plants, with varying degrees of tolerance of exposure to direct sunlight and to drying wind. In Western Malaysia Cyathea moluccana, C. squamulata and C. glabra occur only in quite shady forest and will not tolerate exposure; C. /atebrosa is most vigorous where it has more light and will tolerate almost full exposure of its crown to the sun; C. contaminans is only vigorous where its crown is fully exposed, though its roots need shade. It is probably significant that C. contaminans, flourishing in clearings in the forest, is more widely distributed than any other Ma- laysian species. On high mountains in New Guinea a few species (notably C. macgregorii) can tolerate the full exposure of open grassland and will tolerate periodic burning of the grass (fig. 1, 2). Vegetative morphology. The majority of species are arborescent, and the habit of growth very similar in all genera. The height to which the trunk will grow varies from species to species; full records of this are not available. The lower part of the trunk has many adventitious roots, which become entangled and form a rigid covering of increasing thickness, supporting the base of the trunk; the cover of roots at the base of an old tree-fern is many times thicker than the original trunk. Some species produce branches, either near the base of the trunk or higher up; in the former case there will be a small cluster of trunks, in the latter the main trunk will have lateral crowns of small leaves upon it (fig. 3-4). SCHOUTE reported an exceptional case in which 33 such lateral branches on one plant formed many roots which coalesced with those of the main trunk, each lateral branch growing upwards and forming a separate trunk; the result was a cluster of trunks all growing out from one great mass of roots, and the nature of the branching could only be seen by removing the roots (Ann. Jard. Bot. Btzg 20, 1906, 198-207). ScHoutE also published a detailed study of four cases in which trunks of tree-ferns appeared to branch by bifurcation, remarking that this condition is rare and may be due to injury (Rec. Trav. Bot. Néerl. 11, 1914, 95-192, t. 5-21). He noted that at each bifurcation is an ‘Angularblatt’, as in ferns with dicho- tomously branched rhizomes, and he suggested that perhaps there is no sharp distinction between such bifurcation and lateral branching in which the branch occurs on one side of a leaf-base. Some species of Cibotium and Culcita, and also Cystodium sorbifolium, have prostrate stems with 68 FLORA MALESIANA Ch aS Fig. 3. Cyathea contaminans (WALL.) CopEL. with branching of upper part of trunk. Mt Telemojo, Central Java (P. ARENS). —Fig. 4. The branched trunk shown in fig. 3 with fronds removed. indefinite apical growth; these prostrate stems are massive, with leaves close together, never long-creeping which is a distinction from Dennstaedtia, a genus united to Dicksonia by Hooker. The indefinite horizon- tal growth of these stems contrasts with the vertical growth of the arborescent species, which sooner or later outgrow their mechanical strength if they do not earlier succumb to other injuries. Cyathea biformis has a very slender trunk which supports itself by adventitious roots clinging to a tree-trunk; its fronds are more widely spaced than in arborescent species. The diameter of the trunk varies considerably, from the very slender C. biformis just mentioned to the massive C. contaminans and allied species. The fronds usually break near the base when they are old, their bases persisting for a longer or shorter period; in many cases they are ultimately shed, leaving distinctive scars on the trunk (the scars sometimes later covered by roots). The shape and arrangement of the scars depends on various factors, one being rate of growth of the trunk. A fast-growing trunk will have rather widely spaced frond-scars which are vertically elongated; a slow-growing trunk will have frond-scars closely placed and almost circular (fig. 6). Frond-scars seem always to be in vertical orthosti- chies, and in several spiral parastichies. ScHourTe (/.c.) described the arrangement of leaf-scars on the branching trunks he examined, and especially the way in which the pattern on the branches is related to that on the parent trunk. In the uniform conditions of the forest of Western Malaysia, new fronds ap- pear singly, but probably they are more abundant following wetter periods. At Tjibodas, West Java, JAAG made observations on seven young plants of Cyathea contaminans (Alsophila glauca) and found that the average time between development of successive fronds varied from 25 to 28 days, and the life of a single frond from 165 to 200 days; the number of fronds on a single plant varied from 6 to 10, and the time taken for a complete renewal of the whole crown of fronds from 182 to 243 days. An old plant with trunk 10 m tall bore about 12 fronds and the mean time between unfolding of new fronds was 21 days (Mitteil. Naturf. Ges. Schaffhausen 12, 1943, 211-217). It is remarkable that fronds thus ap- Dec. 1963] | o3 CYATHEACEAE (Holttum) iy 69 pearing singly leave ultimately the scars of their bases in regular alternate whorls. In New Guinea, Hooc- LAND has observed that some species produce their fronds in whorls, those of one whorl being simul- taneous. More observations on phyllotaxis and on rate of growth are needed. There is also the con- sideration that plants of the same species growing under more or less favourable conditions may vary considerably in the size of trunk and of fronds. Few Malaysian species have such massive trunks as the Australian C. australis (R. Br.) Domin and Dicksonia antarctica, which both have fronds in many orthostichies. JAAG observed three plants of Dicksonia blumei at Tjibodas. Each crown usually consisted of 16-18 fronds, each new frond appearing at an average interval of 22—27 days, the life of a single frond being 185-191 days. The pneumathodes which occur along each side of the stipe and rachis often afford distinctive char- acters in Cyathea (fig. 7) but are usually not seen in herbarium specimens, which shrink along the line of thin-walled tissue. In young fronds the pneumathode has a continuous epidermis containing stomata; later the epidermis ruptures and the cells of the underlying tissue become more or less separated from each other, sometimes having peg-like outgrowths. In shape of frond, shape of leaflets and external form of rachis-branches, Cu/cita and Thyrsopteris differ from the other genera. These differences are summarized as follows. Frond-form: in Culcita and Thyrsopteris broadly deltoid, 3—4-pinnate, stipe always long; in other genera elliptical, usually bipinnate- tripinnatifid, lower pinnae always somewhat reduced, sometimes much so and then the stipe very short. Shape of leaflets: in Culcita and Thyrsopteris very asymmetric at base (broad on acroscopic side), with a gradual reduction from largest to smallest; in other genera nearly symmetrical at base, usually with many leaflets (pinnules) on each pinna of approximately equal size. External form of rachis-branches: in Culcita and Thyrsopteris upper surface grooved (fig. 34c), the groove open to admit grooves of smaller branches and of midribs of leaflets (which are similarly grooved), edge of lamina separately decurrent on side of rachis-branch; in other genera upper surfaces raised (or at most slightly grooved), midribs of leaflets also raised (fig. 18b). It may be noted that in ali these characters there is more or less complete agreement between Culcita and the Dryopteris-Athyrium group of genera, and between the other genera and the Thelypteroid ferns. Dermal appendages. In Dicksonia, Cystodium, Culcita and Cibotium the dermal appendages are all simple septate hairs, the longest often quite thick at the base; the characters of the hairs (rigid or flaccid, long or short, varied colour) especially on the stipe, are always important diagnostically. In Cyathea there are always septate hairs (sometimes branched near the base) on the upper (adaxial) surface of stipe and rachises, these hairs rather crisped and antrorse; in most species there are no hairs on the lower surfaces. All species of Cyathea have scales on the lower surfaces, in size decreasing from those on the base of the stipe to those on costules of pinnule-lobes. The genus may be divided into two subgenera, subg. Cyathea having flabelloid, subg. Sphaeropteris setiferous scales (fig. 8). Where hairs occur on lower surfaces in subg. Sphaeropteris they are rather thick and straight, much as in Dicksonia; in subg. Cyathea they are crisped and more or less appressed. For further notes on scales, see Cyathea. Sori. In Dicksonia, Cystodium, Culcita and Cibotium the sorus is at the end of a vein (or of the acros- copic branch of a vein) near the margin; it is protected by a small reflexed marginal lobe (the outer in- dusium) and by an inner indusium which shows varying degrees of difference from the outer indusium. The inner indusium is more or less fused to the side of the receptacle remote from the margin (fig. 31b, c). The surface of the receptacle is not very prominent, but spreads at right angles to the end of the vein. In Cyathea the sorus is usually seated at the fork of a vein, well away from the margin; or where the veins are not branched, it is apparently in the middle of a vein. There is always vascular tissue in the receptacle, and this is the end of a short branch-vein. The receptacle is prominent, more or less spherical or clubshaped. The indusium is of very varied form, with also varying degrees of reduction, and is sometimes lacking. The Cyathea sorus may be compared to Dicksonia by considering the form called Hemitelia (fig. 9c), which has an indusium attached to the base of the receptacle on the side remote from the margin (there is never an indusium attached only on the marginal side). The beginning of the de- velopment of such a sorus and of a sorus of Dicksonia are identical; the receptacle appears to be on the true leaf-margin, with an outgrowth on upper and lower sides. In Dicksonia the two outgrowths develop almost equally, in Cyathea very unequally so that the sorus is ultimately far from the margin, and new veins are needed to supply the additional marginal area of leaf-lamina. The derivation of the other types of Cyathea sorus from this one is described under Cyathea. In Dennstaedtia, formerly included in Dicksonia, the receptacle is more or less columnar and free, at the end of the vein, and is surrounded by almost completely fused outer and inner indusia; it thus differs from both Dicksonia and Cyathea. Sporangia and spores. Sporangia are not very large, having in most cases 64 spores. The annulus is in all cases more or less oblique and usually indurated at the base where it passes the stalk. For details, see Bower, The Ferns 2 (1926) 266, 282, 301 and fig. 5. The stalk is in all cases rather massive, consist- ing of 4-7 rows of cells. In genera with the Dicksonia type of sorus, where the receptacle is not prominent, sporangia have rather long stalks; in Cyathea and Thyrsopteris, where the receptacle is prominent, spo- rangia-stalks are short. Spores are in all cases trilete, and the sculpturing of the surface varies consid- erably, though it is always slight in Cyathea. The spores of Cnemidaria (confined to C. horrida and re- 70 FLORA MALESIANA [ser.; Il, vole lated species with simply pinnate fronds and anastomosing veins) have an almost spherical cavity in the thickened lateral walls (alternating with the trilete ridges). This tropical American group have the most distinctive spores in the family, and should rank as a separate genus. ol Fig. 5. Cyathea capensis (L. f.) SM. a. Sporangium, outer face showing complete annulus and short stalk, b. same, lateral view to show stomium.—C. brownii DoMIN (Alsophila excelsa R. BR.). c. Sporangium, inner face, d. same, lateral view, e. transverse section of stalk. All x 100 (after F. O. Bower, The Ferns, 2, fig. 563). —Fig. 6. Cyathea contaminans (WALL.) CopPEL. Trunk showing scars after abscission of stipes. Mt Bukit Tungul, W. Java (L. VAN DER PUL). Gametophyte. STOKEY gave a comparative account of gametophytes of all genera except Metaxya and Cystodium (Bot. Gaz. 90, 1930, 1-45). Mature prothalli are longer, with more massive cushion, and with greater tendency to fork, than prothalli of the majority of more specialized ferns. Multicellular hairs of peculiar origin occur abundantly in Cyathea, rarely and late in Lophosoria, not in the other genera. These hairs are in origin like those only of Gleichenia and Loxsoma (see STOKEY & ATKINSON, Phyto- morphology 6, 1956, 260); they are longer in Cyathea than in Gleichenia and lack terminal glandular cells. Antheridia in all cases are relatively primitive, with wall of 5 cells; those of Thyrsopteris are largest and least symmetrical. Archegonia have rather long necks, longer in the Dicksonia group of genera than in Cyathea. Cytology. Chromosome numbers in the genera Dicksonia, Culcita and Cibotium have been recorded by MANTON (J. Linn. Soc. Bot. 56, 1958, 84) and in Cyathea by MANTON & SLEDGE (Phil. Trans. R. Soc. B, 238, 1953, 137) and by MANTON (Appendix to HoL_trum, Rey. Fl. Malaya 2, 1954, 623); Prof. MANTON also permits me to report unpublished observations on Culcita and Dr T. G. WALKER on Cnemidaria. No observations are yet available for Thyrsopteris, Lophosoria, Metaxya and Cystodium. The numbers are: Dicksonia, n = 65 (3 spp.); Cibotium, n = 68 (2 spp.); Cyathea, n = 69 (several spp.); Cnemidaria horrida, n = 69; Culcita macrocarpa, n = 66 approx.; Culcita dubia, n = 58. Anatomy. The first critical account of anatomy in this family was by METTENTIUs, in the course of his study of Angiopteris (Abh. M.—Ph. K1. K. Sachs. Ges. Wiss. 6, 1863, 525-531, t. V.) The most recent full account of the anatomy of members of Cyatheaceae is by OGURA (J. Fac. Sc. Imp. Univ. Tokyo, Bot. 1, 1927, 141-350). U. SEN has recently completed a new anatomical study of the family, summarized in HoOLTTUM & SEN Dec. 1963] ‘ved CY ATHEACEAE (Holttum ) a 2 71 (Phytomorphology 11, 1961, 406-420). The vascular structure, with its accompanying sclerotic tissue, is very similar in all Malaysian genera except Cibotium; it is most fully developed in Cyathea, to which the following notes apply. As seen in a transverse section of the trunk, there are several meristeles, with gaps between them, together forming a hollow cylinder, the gaps corresponding to leaf-bases. On the outer and inner sides of each meristele are plates of very hard sclerotic tissue. Small vascular strands arise from the margins of the gaps and supply the leaves, and in Cyathea there are also small medullary bundles which anastomose with each other and with the meristeles. Distinctive ‘cubical cells’ form a more or less continuous layer surrounding each mass of sclerenchyma; their walls adjacent to each other and to the sclerenchyma are much thickened, and they contain crystals which appear to be silica. The sclerotic tissue, with its cubical cells, is lacking in Cibotium. In the phloem are tangentially elongated cells, in structure like the longitudinal sieve-tubes; such cells are only otherwise known to occur in Os- munda. The pattern of arrangement of the numerous vascular strands in the stipe of Cyathea is distinc- tive; in other genera they are more or less joined. In the smaller axes of the frond the pattern is progres- sively simplified. The stomata of Cibotium show more complex developmental stages than those of the other genera. Economic importance. OcCHSE & BAKHUIZEN VAN DEN BRINK reported the use of coiled young fronds of Cyathea contaminans and C. junghuhniana (mis-named C. latebrosa) as food, also the pith of young parts of the trunk of the former species (Vegetables D. E. I., 1931, 212-215). Other species (perhaps all) are similarly edible; HOOGLAND notes this of some from the mountains of New Guinea. The pith of trunks was formerly eaten by Maoris in New Zealand. The common name in Java and Sumatra for the larger tree-ferns, Pakis (or Paku) tiang (tiyang, teehang), indicates the use of the trunks as posts; this name does not seem to have been noted in the Malay Peninsula. The sclerenchyma of most tree-fern trunks is exceedingly hard and durable, and provides nearly all the mechanical strength when they are used as posts. It also provides an interesting pattern when cut in different ways, and this effect is used in the construction of ornamental objects in various parts of the world. In North Borneo I noted old tree-fern trunks, hollowed out, in use as bee-hives around Dusun houses. On Mt Patuha, W. Java, hollowed tree- fern trunks are filled with carbide gas for making booms on New Year’s eve. The masses of adventitious roots at the bases of Cyathea trunks are used in orchid culture, either as solid slabs (cut with a saw) or broken, in potting mixtures. Taxonomy. BERNHARDI (in Schrader, Neues J. Bot. 1, ii, 1806, 1-204) attempted a classification of ferns according to the form and position of the annulus of a sporangium, proposing a division into Helicogyratae (including Cyathea and Dicksonia), Cathetogyratae (majority of leptosporangiate ferns), Pseudogyratae (including Gleichenia) and Agyratae. PRESL (Tentamen, 1836) varied this by associating Gleichenia and Cyathea (sens. lat.)in Helicogyratae and placing Dicksonia (under the name Balantium) in Cathetogyratae. HOOKER (Sp. Fil., 1844 and Syn. Fil., 1868) arranged all ferns in seven suborders, all genera here treated being included in suborder Polypodiaceae; they are divided as tribe Cyatheae (Cyathea, s.l.) and tribe Dicksoniae (all other genera, also some additional ones). METTENIUS (Fil. Hort. Bot. Lips., 1856) arranged all ferns in eight orders, of which the second was Cyatheaceae, which corresponded exactly with the present arrangement with the addition of Matonia (METTENIUS used the name Balantium in place of Dicksonia). Curist (Farnkr. d. Erde, 1897), Diets (in Engl. & Prantl, Pflanzenfam. 1, Abt. 4, 1899, 113-139) and CHRISTENSEN (Ind. Fil., 1905) adopted a family Cyatheaceae with the same content as the order Cyatheaceae of METTENIUS, with omission of Matonia. Bower, however, believed that Cyathea and its near allies should be associated closely with Gleicheniaceae, as one of the more primitive elements of the series Superficiales, while he placed Dicksonia and allies in the series Marginales, regarding the separation of the two as ‘long overdue’ (The Ferns 2, 1926, 326). This idea was followed by CuHrIs- TENSEN in 1938 (in Verdoorn, Man. Pterid., 532, 533), where he recognized two families, Dicksoniaceae and Cyatheaceae. BowER’s arrangement involves the assumption that primitive Cyathea, like Gleichenia, was exindusiate, so that indusia in his Cyatheaceae are a new development, not homologous with the inner indusium of Dicksonia (l.c. 304). In this he disagreed with GogBeL (Flora 105, 1913, 45), who regarded the indusium of Hemitelia (now included in Cyathea) as strictly homologous with the inner indusium of Dicksonia. COPELAND (Gen. Fil., 1947) included Dicksonia and allies in a family Pteridaceae, associating with them Lindsaea, Dennstaedtia, etc., while maintaining Cyathea s./. in a separate family Cyatheaceae. HOLTTUM & SEN have published a discussion of the whole question (Phytomorphology 11, 1961, 406-420), with the conclusion that GoEBEL’s contention was correct; they give a new subdivision of the family Cyatheaceae, as here constituted, based partly on new evidence. This subdivision is sum- marized as follows. CONSPECTUS OF THE FAMILY Subfamily Cyatheoideae. Fronds normally bipinnate with lower pinnae more or less reduced; pinnules almost symmetrical; upper surfaces of costae and pinna-rachis raised (or, if grooved, the groove of a major axis not open to admit that of a minor one borne upon it); sori terminal on veins or on lower surface of veins, indusiate or not; dermal appendages hairs or scales or both; cubical cells present in association with sclerenchyma; stomata with single subsidiary cell. UZ FLORA MALESIANA [ser: I voliate Tribe Cyatheae. Scales and hairs present as dermal appendages; sori superficial, indusiate or not; cubical cells in continuous layer on surfaces of sclerenchyma. Fronds mostly bipinnate; veins almost always free; spores with thin walls of uniform thickness, smooth or papillose; indusium various or lacking . aber . . 1. Cyathea Fronds simply pinnate with anastomosing veins; spores with ‘wall much thickened, a spherical hollow in the middle of each face; indusium hemitelioid (c. 10 spp., tropical America) (Cnemidaria) Tribe Lophosorieae. Hairs only as dermal appendages; sori superficial, no indusia; cubical cells singly in association with sclerenchyma’ (lsp;.. tropical@Amenica)) Go °05 F750 SPR ie tee ae gant en tance ra ea Tribe Dicksonieae. Hairs only as dermal appendages; sori marginal, protected by slightly modified marginal lobe of lamina (outer indusium) and a thinner inner indusium; receptacle of sorus fused to inner indusium. Fronds bipinnate with deeply lobed ah a or ars stem usually a thick erect trunk; cubical cellsasin Cyatheae .. . : . . . 2. Dicksonia Fronds bipinnate with simple pinnules; stem prostrate; veondition of enbical cells not known. 3. Cystodium Subfamily Thyrsopteridoideae. Fronds 3—4-pinnate, lowest pinnae largest; leaflets asymmetric; upper surface of axes and of leaflet- midribs grooved, grooves of major axes open to admit those of minor ones; sori at ends of veins; cubical cells present; stomata with single subsidiary cell. Tribe Thyrsopterideae. Fertile and sterile parts of frond strongly dimophous (lamina much reduced in fertile part); receptacle of sorus columnar with sporangia all round it, indusium ultimately a shallow uniform cup; stem massive, erect; cubical cells scattered (1 sp., Juan Fernandez Is). . . .. . . . . (Thyrsopteris) Tribe Culciteae. Fertile and sterile parts of frond not greatly dimorphous; receptacle of sorus fused to inner indusium (as in Dicksonia); inner indusium thinner than outer, the two slightly joined together at the base; stem prostrate or erect; cubical cells in a continuous layer as in Dicksonia. . .. . . . 5. Culcita Subfamily Cibotioideae. Fronds normally bipinnate; pinnules almost symmetrical; upper surfaces of pinna-rachises and costae raised; sori terminal on veins, shape much as in Dicksonia but with outer and inner indusia both unlike the lamina of the frond, lacking chlorophyll and lacking intercellular spaces; sclerenchyma and cubical celisilackinis:stomatal withi3isubsidiary, cells 34. 5+. =. -aem QueIey nc) ee suse Or DO tina Subfamily Metaxyoideae. Fronds simply pinnate, pinnae lobed on young plants only; upper surface of rachis and of midribs of pinnae grooved, groove of rachis open to admit grooves of pinna-midribs; sori superficial on lower surface of veins, usually more than one to a vein, no indusium; sclerenchyma and cubical cells lacking; stomata with 3)subsidiary. cells) (1 sp:, tropical America)... «3% >...» « «.- + = = (Metaxga)) ARTIFICIAL KEY TO MALAYSIAN GENERA 1. Upper surface of costae of ultimate leaflets raised; fronds + elliptical, mostly bipinnate. 2 Youngspartsiot plant protected’ by scales‘and hairs}. .-. 2. .)- - = 1): o)e 9) eeeele@yatnes 2. Young parts of plant protected by hairs only. 3. Outer indusium not distinct from rest of lamina. 4. Pinna-rachis raised on upper surface; fertile pinnules deeply lobed. . . . . . . 2. Dicksonia ae Pinna-rachis grooved on upper surface; fertile pinnules not lobed. . . . . . . 3. Cystodium Outer indusium quite distinct from lamina of leaflet. . . . ;-.-. 4. @ibotum ile are surface of costae of ultimate leaflets grooved, the groove decurrent into the groove of the supporting rachis; fronds deltoid in outline, 3-4-pinnate ..........~... . #S. €uleita 1.CYATHEA SMITH, Mem. Ac. Turin 5 (1793) 416; Swartz, Syn. Fil. (1806) 139, 364; KAULF. En. Fil. Chamisso (1824) 254; Pres_, Tent. Pterid. (1836) 54; Hook. Gen. Fil. (1839) t. 23; Sp. Fil. 1 (1844) 14; Syn. Fil. (1865) 16; J. Smiru, Lond. J. Bot. 1 (1842) 659-668; Hist. Fil. (1875) 244; Curist, Farnkr. Erde (1897) 10, 317; DIELS in E. & P. Pfl. Fam. 1, 4 (1899) 123; CopEL. Philip. J. Sc. 3 (1909) Bot. 353; Gen. Fil. (1947) 95.—Sphaeropteris BERNH. in Schrader, J. Bot. 1800, ii (1801) Dec. 1963] CYATHEACEAE (Holttum ) 73 Fig. 7. On left Cyathea orientalis (KUNZE) Moore, young frond showing widely-spaced short pneuma- thodes. On right C. incisoserrata CopEL., bases of stipes showing almost continuous and often double rows of pneumathodes. Cult. R. B. G. Kew (R. VAN CREVEL, 1961). 122.—Hemitelia R. Br. Prod. (1810) 158, p.p. (excl. H. horrida).—Alsophila R. Br. /.c.—Chnoophora KAULF. En. Fil. Chamisso (1824) 250.—Gymnosphaera BL. En. Pl. Jav. (1828) 242; CopeL. Gen. Fil. (1947) 98.—Disphenia PResL, Tent. Pterid. (1836) 55.—Schizocaena J. SM. in Hook. Gen. Fil (1838) t. 2; COPEL. Gen. Fil. (1947) 99.—Amphicosmia GARDNER, Lond. J. Bot. 1 (1842) 441.— Dichorexia PrReEsL, Abh. K. B6hm. Ges. Wiss. V, 5 (1848) 55.—Fourniera BOMMER, Bull. Soc. Bot. France 20 (1873) xix.—Eatoniopsis BoMMER, /.c.—Thysanobotrya v. A. v. R. Bull. Jard. Bot. Btzg II, n. 28 (1918) 66.—Fig. 1-30. Trunk always erect, short or tall. Scales present on lower (abaxial) and lateral surfaces of axes of frond, at least in early stages of growth, larger ones on stipe and rachis sometimes borne on spine-like outgrowths; fairs always on upper surfaces of all but smallest axes, antrorse, sometimes branched; hairs of various kinds some- times on lower surfaces of axes and veins, rarely on lamina between veins. Pneu- mathodes present in a discontinuous line (or 2-3 lines close together) along each side of stipe and rachis, in subg. Cyathea converging downwards on each side of base of stipe and there often deeply excavated at maturity (fig. 7, 12). Fronds + elliptical, lower pinnae always smaller than middle ones, sometimes gradually much reduced and then the stipe short; pinnae normally pinnate-bipinnatifid, in a few cases simple, in a few cases fully bipinnate; pinnules almost symmetrical at the base, many on each pinna subequal, distal ones more or less abruptly decreas- ing; upper surface of pinna-rachis and costa raised; veins simple or branched, 74 FLORA MALESIANA (ser. Lip volume lower ones usually once forked, sometimes pinnate where pinnule-segments are deeply lobed. Sori usually at the fork of a vein, or seated on a simple vein, a branch of the vein always entering the receptacle; indusium either attached all round base of receptacle and covering young sorus, opening to form a firm-edged cup or open- ing by irregular rupture, or attached on costular side of receptacle (hemitelioid) and of varying size, in some cases quite hidden by mature sorus, or lacking; receptacle erect, + club-shaped to spherical; sporangia many, always short- stalked; paraphyses usually present as multicellular hairs, sometimes flat and several cells wide at base; in some species of swhg. Sphaeropteris scales present round base of receptacle, more or less covering young sporangia; spores thin- walled, smooth or papillose. Type-species: Cyathea arborea (L.) SM. (tropical America). Distr. & Ecol. See under the sections. Morph. Dermal appendages. Scales on the stipes of Cyathea are of two kinds, and these appear to provide the best subdivision of the genus, at least in Malaysia. The two types of scale are called flabelloid and setiferous (see Ho_ttruM, Kew Bull. 1957, 41-45; Hottrum & SEN, Phytomorphology 11, 1961, 406-420). Flabelloid scales (fig. 8a, b) have a broad median portion consisting of longitudinally elongated cells with all walls thickened, and edges, of varying width, consisting of thin-walled cells diverging fan-wise outwards, with irregularly projecting marginal cells, some of them sometimes thick-walled and dark, often flexuous (fig. 9a, b). The scales develop at the apex of more or less massive (multicellular) out- growths from the surface of the stipe and at right angles to these outgrowths (thus parallel to the surface of the stipe); the base of a scale is peltate, with a narrow part encircling the supporting outgrowth on the Fig. 8. Cyathea oinops Hassk. a. Scale from costa of pinnule, showing median band of dark thick-walled cells and flabelloid margins of thin-walled cells with a few very thick-walled setae, x 70.—C. incisoser- rata CopgL. b. Part of flabelloid margin of scale from stipe, < 100.—C. squamulata (BL.) COPEL. c. Apical part of scale from costa of a pinnule, showing setiferous (not flabelloid) edge, < 70, c’. two very small scales from costa, x 70. basiscopic side. The outgrowths become very large and spine-like in some species, in others they are quite small. As one proceeds from the base of the stipe to the ultimate axes, the scales become progressi- vely smaller, and their character changes in ways characteristic of individual species. These scales pro- vide some of the most important diagnostic characters in Cyathea. Setiferous scales (fig. 8c, c’) also develop at the apex of outgrowths from the surface of the stipe, but not at right angles to the outgrowths; the base of a scale widens more or less abruptly from the apex of the outgrowth, and in C. sangirensis may be seen transitions from stout erect hairs (like those of Dicksonia) to setiferous scales. All cells in setiferous scales are longitudinally elongate and all have walls Dec. 1963] CYATHEACEAE (Holttum) = 75 Fig. 9. Different types of indusia in subg. Cyathea. a. C. batjanensis (CHRIST) COPEL.; indusium a small disc covered by mature sorus.—b. C. javanica BL.; indusium saucer-shaped, often asymmetric.—c. C. heterochlamydea CoreL.; indusium attached on costular side of receptacle, covering part of base of mature sorus.—d. C. oinops HASskK.; indusium covering sorus almost to maturity, but open on side remote from costule (two indusia cut to show receptacle, costular scales also shown).—e. C. orientalis (KUNZE) Moore; indusium a complete rather deep cup.—f. C. crenulata BL.; indusium very fragile, covering sorus completely to maturity, then breaking irregularly and in part disappearing (a, < 6; b-f, LS: Trunk to 24% m, 16 cm g, bearing up to 40 fronds 60-105 cm long. Stipe 20-30 cm, when dry light red-brown and warty after fall of scales, base densely covered with shining firm twisted brown scales to 50 by 2 mm, edges bearing ir- regular concolorous short setae; above the base more or less densely covered with small scales as rachis. Rachis covered beneath with a close felt of very small pale scales which have fiexuous marginal hairs, also with larger scales: (a) elongate flat pale scales to 1 mm wide having many brown setae near apices and often pale hairs near bases, (b) very narrow pale thin flexuous scales with slender marginal hairs; all scales more or less caducous with age. Longest pinnae 10-16 cm. Longest pinnules 30 by 8 mm, lobed almost or quite to costa; costules 2 mm apart, veins 6-7 pairs; lamina-segments very rigid, their edges much reflexed and inrolled, entire or slightly crenate. Sori filling cavity of lower surface of a segment; no indusium. Scales and hairs: lower surface of pinna-rachis and costae as main rachis but the flat scales smaller or lacking; on costules small brown bullate scales with long crisped marginal hairs; hairs on upper surface of costae pale, with a few on costules. Type specimen: Brass 9116, Lake Habbema, West Central New Guinea (A; dupl. at MICH, BOVE UE): Distr. Malaysia: New Guinea. Ecol. At 3225 m, ‘along streams of open grassland, in drier shrubberies, never in forest’ (BRASs). Note. This appears to be distinct in its narrow stipe-scales and small size of frond; also in the close felt of small woolly scales on lower surfaces of all rachises, though this is more or less ca- ducous. In other characters not clearly distin- guishable from C. atrox var. inermis. 126. Cyathea atrox C. Cur. Brittonia 2 (1937) 215; CopeL. Philip. J. Sc) 77 9@S4i) eis — Fig. 1, 17, 24, 25. var. atrox. Trunk to 6 or 7 m, bearing 20-30 fronds 125-175 cm long. Stipe 30-50 cm, spiny, spines dark, shining, 3-5 mm; scales near base of stipe to 50 by 3-6 mm, shining brown, firm but not very rigid, edges bearing irregular concolorous or pale hairs or setae; distal part of stipe glabrescent, light brown when dry, warty and thorny. Lower pinnae slightly reduced; longest 30-45 cm. Largest pinnules commonly 65-80 by 13-15 mm, excep- tionally to 100 by 20 mm, almost all segments separately adnate to costa (a few only on pinnules of upper pinnae) and constricted at base, 1-2 basal ones quite free; costules commonly 3 mm apart, on largest fronds to 4 mm; veins 8—10(—12) pairs, pale and raised on lower surface; lamina- segments rigid, edges reflexed and more or less inrolled, on distal segments entire, on largest ones rather deeply crenate, the basal free segments often deeply lobed. Sori nearer to costules than edge; no indusium; paraphyses short. Scales and hairs: pinna-rachis rather pale, warty, glabrescent, bearing residual scales as stipe but smaller; on lower surface of costae some elongate flat brown scales with dark marginal setae (these mostly caducous) and very small pale scales bearing long pale or dark marginal hairs; on costules many small brown bullate scales bearing crisped pale or dark marginal hairs; similar scales sometimes on lower surface of veins. Type specimen: Brass 4596, Murray Pass, Wharton Range, Papua (A; dupl. at BM, BO, BRI). Distr. Malaysia: E. New Guinea. Ecol. On edges of forest or in grassland, 2800-3600 m. 4 Notes. Most collections are reported as having many fronds, but two as having fronds in distinct age-groups, 6 or 7 (in a whorl) in each age-group. One of these (HOOGLAND & SCHODDE 7165) is unusually large, with pinnules 100 mm long. Dec. 1963] CYATHEACEAE (Holttum) 129 Fig. 24. Cyathea atrox C. CHR. var. inermis HOLTTUM in peaty grassland, Eastern Highlands, Territory of New Guinea, 3000 m (R. D. HOOGLAND). 130 FLORA MALESIANA fser. Il, vou var. inermis HoLtrum, var. nov.—Fig. 24, 25. Stipes yverrucosus, vix spinosus; paleae Sstipitis 6-10 cm longae, ad 6 mm latae; pinnulae vulgo ad 45 & 12 mm; costulae 2\% mm inter se distantes; rhaches pinnarum subtus paleis minutis ciliatis pal- lidis vestitae. Stipe with numerous conical warts 14 mm high; stipe-scales 60-100 mm long, to 6 mm wide; pinnules commonly to 45 by 12 mm, edges strongly reflexed and inrolled, costules 21% mm apart; pinna-rachis often with very small pale fringed scale. Type specimen: HoOGLAND & SCHODDE 7457, NE. New Guinea, Western Highlands (L). Ecol. ““Commonest species in tree-fern grass- land’’, at 2600-2900 m. A specimen from 3500 m (BrRAss 31013) has darker, more rigid, twisted stipe-scales to 60 mm long. Note. Two collections are reported as bearing 22-30 fronds spirally arranged, one (SCHODDE 1762) as having fronds in two whorls of 6 or 7. 127. Cyathea sangirensis (CHRIST) CopeL. Philip. J. Sc. 4 (1909) Bot. 37.—Alsophila concinna BAK. Syn. Fil. ed. 2 (1874) 459, non C. concinna (BAK.) Jenm. 1891.—Alsophila polyphlebia Bax. J. Linn. Soc. Bot. 15 (1876) 104, non C. polyphlebia BAK. 1883.—Alsophila sangirensis CHRIST in Warb. Monsunia 1 (1900) 90.—Alsophila_ scaberula Curist in K. Sch. & Laut. Fl. Deut. Schutzgeb. Siidsee (1901) 110; v. A. v R. Handb. (1908) 35.— Fig. 25. Cyathea atrox C. Cur. var. inermis HoLttuM on the Sugarloaf complex, Western Highlands (R. D. HOOGLAND). Dec. 1963] CYATHEACEAE (Holttum) 131 C. scabriseta CopPEL. Philip. J. Sc. 9 (1914) Bot. 2; ibid. 77 (1947) 111.—Alsophila okiana v. A. Vv. R. Bull. Jard. Bot. Btzg II, n. 23 (1916) 4; Handb. Suppl. (1917) 494.—Alsophila rumphiana v. A. V. R. Philip. J. Sc. 11 (1916) Bot. 104; Handb. Suppl. (1917) 491.—Alsophila scabriseta v. A.V. R. Handb. Suppl. (1917) 73.—C. rumphiana MERR. Interpr. Rumph. Herb. Amb. (1917) 63.— Alsophila buruensis ROSENST. Med. Rijksherb. n. 31 (1917) 1.—C. okiana v. A. v. R. Bull. Jard. Bot. Btzg II, n. 28 (1918) 14.—Alsophila scaberulipes v. A. v. R. Nova Guinea 14 (1924) 2.—C. eminens Donn, Pterid. (1929) 262 (new name for Alsophila concinna BAK.); CopeL. Philip. J. Sc. 77 (1947) 116.—C. aruensis Domin, Pterid. (1929) 262 (new name for A/sophila polyphlebia BaAK.); Co- PEL. Philip. J. Sc. 77 (1947) 115.—C. scaberula Domin, Pterid. (1929) 263; Cope. Philip. J. Sc. 77 (1947) 109.—C. brassii Cope. J. Arn. Arb. 10 (1929) 175; Philip. J. Sc. 77 (1947) 111.—C. buruensis Domtin, Acta Bot. Bohem. 9 (1930) 102. — C. scaberulipes Doin, I.c. 174. Trunk stout, to at least 8 m. Stipe long (ap- parently to 100 cm), pale, finely warty, persistently scaly near base; scales pale, thin but firm, finely acuminate, to c. 30 by 2 mm, edges closely set with dark setae. Lamina to 300 cm or more long. Main rachis beneath pale and finely warty, some- times with residual spreading very narrow se- tiferous scales or hairs. Pinnae: largest about 75 cm long, lowest not greatly reduced. Pinnules: largest 120 by 25 mm, sessile, lowest 1 or 2 seg- ments usually free, rest of pinnule lobed almost to costa; costules 4-414 mm apart; veins commonly 15 pairs in largest segments; segments of lamina thin, strongly crenate (the largest crenae, where a vein is forked, slightly notched). Sori rather near costules; no indusium; paraphyses slender, dark at base and pale distally, alittle longer than sporangia; a ring of very narrow ciliate pale scales round base of sorus. Scales and hairs: pinna-rachis minutely warty, glabrescent or with very small pale scales which do not bear setae; costal scales small, pale, appressed, more or less elongate, bearing pale short marginal hairs; costular scales pale bullate; a few stout pale hairs rarely present near apices of costae and costules on lower surface, these normally lacking; upper surface of costules and veins bearing scattered stout pale hairs, and similar hairs on edges of segments at apices of some veins (these latter early caducous). Type specimen: WaARBURG 16605, Sangir (= Sangihe) I. (P?; dupl. at B). Distr. Malaysia: Moluccas (Sangihe I. (NE of Celebes), Buru, Ambon, Aru Is), throughout New Guinea, and Louisiades (Sudest I.). Ecol. A large lowland tree-fern, in secondary growth and in open places in forest; often near rivers. In Sudest I. common at junction of rain- forest with mangrove (BRASS). Note. MerrRILL (/.c.) has suggested that Palmifilix alba RumMeH. (Herb. Amb. 6, 63) is to be regarded as representing this species, but the data given by RUMPHIUS are not adequate for a certain identification. 128. Cyathea lunulata (Forst.) Copec. Bull. Bern. P. Bish. Mus. 59 (1929) 37; C. Cur. ibid. 177 (1943) 29.—Polypodium lunulatum Forst. Prod. (1786) 83, n. 456.—Alsophila lunulata R. Br. Prod. (1810) 158; Hook. Sp. Fil 1 (1844) 51; Syn. Fil. (1868) 41, p.p.; CARR. in Seem. FI. Vit. (1873) 333.—Alsophila vitiensis CARR. in Seem. Fl. Vit. (1873) 170.—Alsophila naumannii KUHN, Forschungsr. Gazelle 4 (1889) Farne 13; v. A. v. R. Handb. Suppl. (1917) 59.—C. naumannii Domin, Pterid. (1929) 263.—C. vitiensis DOMIN, Acta Bot. Bohem. 9 (1930) 170. Stipe smooth or finely warty; scales to 30 by 114 mm, pale, thin, with a few dark marginal setae. Pinnules to at least 120 by 20 mm; costules 314-414 mm apart; segments of lamina distinctly crenate- serrate; veins to 15 pairs. Sori exindusiate; pa- raphyses shorter than sporangia, usually dark when dry. Scales and hairs: pinna-rachis pale, beneath smooth or slightly prickly, quite glabres- cent; costae bearing stout pale hairs distally on lower surface, scales mostly deciduous except a few very small ones; costules bearing similar hairs and also scales which are bullate and more or less fringed, or on lower pinnules may be setiferous; minute hairs sometimes visible on lower surface of veins; hairs on upper surface of costules rare. Type specimen: Forster, Pacific (Géttingen; dupl. at BM). Distr. Solomon Is, New Hebrides, Tonga, Fiji Samoa, in Malaysia: Bismarck Arch. Ecol. Apparently a large tree-fern of low country. Note. The type specimen is labelled “In Societatis Ins.?”; but a distribution extending to Tahiti seems improbable. 129. Cyathea elmeri Cope-. Philip. J. Sc. 4 (1909) Bot. 54; Fern Fl. Philip. 2 (1960) 233.—Alsophila comosa (non WALL.) CHRIST, Ann. Jard. Bot. Btzg 15 (1898) 80.—Alsophila latebrosa (non WALL.) var. major CurisT, Philip. J. Sc. 2 (1907) Bot. 183.—Alsophila elmeri CopeL. in Elmer, Leafi. Philip. Bot. 2 (1908) 419; v. A. v. R. Handb. Suppl. (1917) 66.—Alsophila christii Vv. A. v. R. Handb. (1908) 42 (non Sop. 1908); Suppl. (1917) 69, 70.—C. dimorphotricha CopEL. in Elmer, Leafl. Philip. Bot. 5 (1913) 1681; Fern Fi. Philip. 2 (1960) 230.—Alsophila subcomosa C. Cure. Ind. Fil. Suppl. 1 (1913) 5.—Alsophila dimorphotricha v. A. v. R. Handb. Suppl. (1917) 61.—C. subco- mosa Domin, Pterid. (1929) 263.—Alsophila fe- nicis [non (CoPEL.) C. Cur.] Post. in Holthuis & Lam, Blumea 5 (1942) 153, p.p.—C. argyrolepis CopeL. Philip. J. Sc. 81 (1952) 17; Fern Fl. Philip. 2 (1960) 227. Stipe 30 cm or more, the base densely scaly, closely warty when scales have fallen; scales 25 by 114 mm, medium brown, thin but firm, shining, acuminate, edges throughout closely set with short concolorous setae. Main rachis, lower surface, pale, closely warty, glabrescent. Pinnae to at least 60 cm long. Pinnules 90-130 by 13-25 mm, sessile, lowest 2 segments of the largest free, 132 FLORA MALESIANA [ser. II, vol. 1? next few segments separately adnate to costa, rest (whole of smaller pinnules) lobed nearly to costa: costules 3—414 mm apart; veins 9-12 pairs; lamina-segments rather thin, strongly crenate- serrate, sinuses narrow. Sori near costules; no indusium; paraphyses dark, not usually longer than sporangia. Scales and hairs: pinna-rachis beneath pale, finely warty, glabrescent, the smallest residual scales pale with a short pale fringe, larger ones with setiferous edges; on costae near base narrow scales with long dark marginal setae, on distal half more or less abundant stout pale spread- ing hairs; on costules bullate-based broad scales, the larger bearing dark setae near apex, the distal ones wholly bullate and lacking setae; hairs also more or less abundant on costules; on upper surface of costules no hairs, or in some cases a few. Type specimen: Ermer 9457, Cuernos Mts, Negros (MICH; dupl. at US, BO, F, K, P, L, U; SYD): Distr. Malaysia: Philippines (Mindanao, Ley- te, Negros, Biliran), Talaud Is, and N. Celebes. Ecol. In forest at 500-1400 m. Notes. The specimen from Talaud Is (LAM 3319), listed as Alsophila fenicis by HOLTHUIS & Lam, appears to lack hairs on lower surface of costae, but its setiferous scales are like those of C. elmeri, not of C. sangirensis. The type specimen of C. argyrolepis, from an exposed place on Camiguin de Mindanao, has small pinnules (to 90 by 11 mm) and is more scaly than normal. C. haenkei (PR.) MeErRR. from Guam is very nearly allied. 130. Cyathea tomentosa (BL.) ZoLL. & Mor. in Moritzi, Syst. Verz. (1846) 108.—Chnoophora tomentosa Bui. En Pl. Jay. (1828) 244.—Chnoo- Phora lanuginosa JUNGH. Tijd. Nat. Gesch. & Physiol. 8 (1841) 372; Flora 30 (1847) 522.— Alsophila tomentosa Hook. Sp. Fil. 1 (1844) 55; Racis. Fl. Btzg 1 (1898) 32; v. A. v. R. Handb. (1908) 43; Suppl. (1917) 493; BACKER & PosTuH. Varenfl. Java (1939) 30.—Alsophila lanuginosa Pr. Epim. Bot. (1851) 29.—Alsophila crinita (non Hook.) v. A. v. R. Handb. (1908) 40. Trunk to 15 m, older parts showing leaf-scars 4-414 cm g. Stipe to at least 50 cm, densely scaly throughout, spines short; scales on stipe light brown, shining, firm, 30-45 by 1-3 mm, edges bearing close dark setae throughout. Lamina to 250 cm long; main rachis rather closely warty with more or less persistent smaller scales. Pinnae: largest to 70 cm or more long. Pinnules to 110 by 20 mm, of plants in exposed places sometimes only 60 by 10 mm wide; basal 1 or 2 segments of lowest pinnules quite free, then 1-2 pairs sep- arately adnate to costa, rest of pinnule lobed almost to costa, apex acuminate; costules 3-4 mm apart; veins 10-12 pairs; lamina-segments very firm, edges usually reflexed when dry, crenate. Sori nearer costule than edge; no indusium; pa- raphyses slender, not longer than sporangia. Scales and hairs: pinna-rachis closely warty and more or less persistently scaly on lower surface, scales strongly setiferous, the larger ones long and narrow; costae densely scaly throughout, scales near base long and narrow, flat, strongly seti- ferous (setae long and dark), grading to smaller long-fringed scales towards the apex; costules densely covered with pale elongate pale-fringed scales, the hairs of the fringes crisped and entan- gled, at least the distal scales witha bullate base; stout spreading pale hairs present on lower sur- face towards apices of costae and costules and scattered on lower surface of veins; a few also on upper surface of costules; hairs on upper surface of pinna-rachis always pale. Type specimen: BLumMeE, Mt Gedeh, Java (L). Distr. Malaysia: Java, Lesser Sunda Is (Flores). Ecol. At 2200 m and over, in ridge forest and in Open swampy places in gullies (abundant near hot springs at 2200 m on Mt Gedeh, West Java). Note. C. crinita (HOOK.) CopEL., of Ceylon and South India, is very closely related to C. tomentosa. It occurs at lower elevations, is less scaly generally, the pinnules are wider, the lamina less rigid, the hairs on scales of costules straighter and less entangled. Specimens from Java formerly recorded as C. crinita are C. tomentosa; if the two species should be united, C. tomentosa is the older name. 131. Cyathea magna CopeL. Un. Cal. Publ. Bot. 18 (1942) 218; Philip. J. Sc. 77 (1947) 110, pl. 7.— Alsophila tomentosa var. novoguineensis ROSENST. in Fedde, Rep. 5 (1908) 34.—C. ordinata COPEL. Philip. J. Sc. 77 (1947) 109. Trunk to 8 m; fronds 8-14, spirally arranged. Stipe 40-90 cm, medium brown when dry, near base more or less spiny, spines 3-5 mm long, finely warty between the spines; large scales abun- dant and persistent near base only, pale or in part medium brown, firm and shining, to 50 by 114(-2) mm, edges closely set throughout with short dark setae; rest of stipe + persistently covered with smaller pale to brown scales, the larger freely dark-setiferous, the smallest fringed with pale hairs. Lamina to nearly 300 cm long; main rachis on lower surface light brown when dry, rather closely warty and scaly as upper part of stipe. Pinnae: largest 90 cm long. Pinnules mostly rather stiffly spreading almost at right angles to pinna-rachis, largest 100-140 by 14-20 mm, sessile, acuminate, lobed almost to the costa (lowest lobe sometimes just free but adnate by its lamina); costules of large fronds 4 mm apart, of smaller ones 3 mm; veins 10-12 pairs, pale and prominent beneath, not raised above; segments of lamina thick, stiffly spreading when dry, edges often rather much reflexed on drying, finely crenate to almost entire. Sori nearer costule than edge; indusium lacking; paraphyses not longer than sporangia, no scales round base of sorus. Scales and hairs: pinna-rachis beneath pale, scaly as main rachis; costae beneath densely scaly, scales near base like the larger ones of pinna-rachis, flat and elongate, strongly setiferous, smaller and narrow scales bearing long flexuous dark marginal setae present almost throughout (smallest have pale CYATHEACEAE (Holttum) 133 Dec. 1963] marginal hairs); costules densely scaly, scales near base narrow flat and bearing long dark setae, rest light brown with + bullate base and long pale flexuous marginal hairs; erect hairs present distally on costules and on lower surface of veins; upper surface of costules not hairy; hairs on upper surface of pinna-rachis dark except towards apex of frond. Type specimen: Brass 11278, Bele R., 18 km NE of Lake Habbema, 2250 m, W. New Guinea (A; dupl. at MICH, BO, K, BM, L, UC). Distr. Malaysia: New Guinea. Ecol. In open places in forest or in grassland, 1700-2750 m. Note. This species is closely allied to C. tomentosa of Java, differing in narrower and firmer stipe scales, more rigid pinnules, and a greater preponderance of small scales bearing long dark setae, these giving a darker aspect; the hairs on upper surface of pinna-rachises are also dark. 132. Cyathea pilulifera CopeL. Un. Cal. Publ. Bot. 18 (1942) 219; Philip. J. Sc. 77 (1947) 112, folk, Ye Stipe 50 cm, basal part copiously spiny, spines dark, to 5 mm; scales pale, to 25 by 2 mm, edges bearing sparse dark setae or concolorous hairs; main rachis spiny, glabrescent; pinna-rachis sparsely spiny or warty. Lamina c. 100 cm long; longest pinnae 45 cm. Pinnules to 110 by 20 mm, lowest segment just free, rest lobed almost to costa; costules 4 mm apart, distinctly curved; lamina-segments firm, close, edges crenate; veins 12-14 pairs, mostly forked, lowest basiscopic vein sometimes from costa. Sori exindusiate; paraphyses not longer than sporangia; scales present round base of receptacle. Scales and hairs on lower surface of costae, costules and veins: flat pale scales with many long dark setae present near bases of costae and smaller ones dis- tally and on costules; pale bullate scales with some setae present on distal part of costae and on costules and veins; stout pale hairs abundant on distal part of costae and on costules and veins; upper surface of costules and veins glabrous. Type specimen: BrAss 11492, 18 km NE. of Lake Habbema, 2200 m, W. New Guinea (A; dupl. at BM, BO, MICH). Distr. Malaysia: Moluccas (Ceram), Guinea, and Louisiades. Ecol. At 1250-2750 m; at some lower elevations reported in forest with tall trunk, at higher ele- vations in open places (the type in young secondary growth on old garden land, with short trunk). New 133. Cyathea curranii Cope. Philip. J. Sc. 3 (1909) Bot. 356; ibid. 4 (1910) Bot. 52; Fern Fl. Philip. 2 (1960) 231.— Alsophila curranii Cy Grins Iie lel Soysylk We (MNS) Se awe ANS we TRE Handb. Suppl. (1917) 70. Trunk 3 m, 20 cm @; leaf-scars 3 cm o. Stipe 35 cm, pale, warty, bearing light brown shining scales to 60 by 4 mm, their edges bearing rather few concolorous setae. Lamina 100 cm long. Lower pinnae long-stalked, reduced and deflexed, largest 35 cm. Pinnules to 90 by 13 mm, lowest segments almost free; costules 3 mm apart; segments of lamina rigid, nearly entire, glaucous beneath; veins 8-10 pairs. Sori exindusiate, paraphyses present. Scales and hairs: pinna-rachis beneath closely warty (warts dark), with a few residual narrow light brown scales, their edges bearing concolorous setae; costae glabrescent; costules and veins beneath bearing scattered pale bullate scales and many very short hairs (probably bases of fallen scales); long hairs absent. Type specimen: CURRAN & MerritT FB 7925, Mt Banajao, 2000 m, Luzon (MICH; dupl. at PSUs): Distr. Malaysia: Philippines (Luzon). Note. Specimens seen are in poor condition and do not include largest pinnae. The nearest relationship seems to be to C. mertensiana (KUNZE) CopeL. of the Bonin Is and C. aeneifolia (v. A. v. R.) Domin, and allied species of New Guinea and the Pacific. 134. Cyathea aeneifolia (v. A. v. R.) Domin, Acta Bot. Bohem. 9 (1930) 174; CopeL. Philip. J. Sc. 77 (1947) 113.—Alsophila aeneifolia v. A. v. R. Nova Guinea 14 (1924) 3.—Alsophila aeneifolia var. subglauca v. A. Vv. R. l.c. 4.—C. curvipinnula C. Cur. Brittonia 2 (1937) 276; CopeL. Philip. J. Sc. 77 (1947) 116. Trunk to 414 m. Fronds 9-12, 175-250 cm long. Stipe 50-60 cm, strongly spiny and scaly; spines dark and shining, to 6 mm; largest scales 30-60 by 3-5 mm, thin but firm and flat when young, rather pale shining brown, edges bearing scattered concolorous hairs or dark setae; rachis short- spiny or warty, bearing similar but smaller scales which are mostly caducous. Lower pinnae slightly reduced and deflexed, largest pinnae 30-60 cm. Largest pinnules 70-120 by 15-25 mm, sessile, lobed nearly to costa, a few lower segments almost or quite free; costules 3-414 mm apart; lamina- segments very firm, almost entire, glaucous be- neath; veins 10-12 pairs. Sori medial on veins, no indusium; paraphyses numerous, as long as spo- rangia; some narrow pale scales surrounding base of receptacle. Scales and hairs: pinna-rachis bearing -- caducous pale brown scales to 151 mm, also small pale fringed scales; throughout lower surface of costae small fringed pale scales, with some elongate setiferous scales near base and bullate ones towards apex; on costules pale bullate hair-tipped scales, often with crisped marginal hairs or sometimes setae; on veins beneath small pale bullate scales more or less abundant, with lax hairs like tips of scales at apices of veins; no hairs on upper surface of costules and veins. Type specimen: Lam 1751, W. New Guinea, near foot of Doormantop, in sunny ravine, 3240 m (BO; dupl. at US, L, U). Distr. Malaysia: New Guinea. Ecol. On forest edges or in grass-land, 2840— 3240 m. LAM 1805, distinguished as var. subglauca v. A. v. R., was collected in a shady valley at 3200 m, and has larger thinner fronds than the type but 134 FLORA MALESIANA [ser. DL, youl is otherwise similar. This species is nearly allied to C. pilulifera CopEL., but the latter has abundant long hairs on costae, costules and veins, and has been found at lower altitudes. Perhaps the two should be united. var. melanacantha (CoPEL.) HOLTTUM, var. nov.— C. melanacantha CopEL. Un. Cal. Publ. Bot. 18 (1942) 219; Philip. J. Sc. 77 (1948) 114, pl. 10. Stipe-scales to 15 by 2 mm, dark brown. Type specimen: Brass 9311, Lake Habbema, W. New Guinea (A; dupl. at BM, BO, UC, L, MICH). Ecol. Occasional in forest-edge, at 3225 m. var. macrophylla HOLTTUM, var. nov. A typo speciei differt: frondibus ad 425 cm longis, stipitibus 200 cm longis inclusis; paleis stipitis ad 50 < 4 mm; pinnis maximis 70 cm longis, pinnulis maximis 135 * 21 mm; costulis 5 mm inter se distantibus. Type specimen: HOOGLAND & SCHODDE 7209, Western Highlands, NE. New Guinea (L). Ecol. In cloud-forest at 2900 m. 135. Cyathea tenggerensis (ROSENST.) DOMIN, Ac- ta Bot. Bohem. 9 (1930) 165.—Alsophila haenkei var. angustata HAssk. in Hook. J. Bot. Kew Misc. 7 (1855) 326.—Alsophila glauca (BL.) HOOK. p.p.: v. A. v. R. Handb. (1908) 41; Suppl. (1917) 68; BACKER & PostH. Varenfl. Java (1939) 28.— Alsophila tenggerensis ROSENST. Med. Rijksherb. n. 31 (1917) 1. Stipe to at least 60 cm, warty, persistently scaly near base; stipe-scales to 45 by 214-314 mm, shining pale brown, firm, setiferous. Pinnae to at least 50 cm long. Pinnules to 110 by 16 mm, sessile, lobed almost to costa with | or 2 basal segments almost free; costules 214-3 mm apart; lamina-segments firm, edges -+ reflexed when dry, lower surface probably glaucous; veins 10—12 pairs. Sori exindusiate, nearer to costule than to edge, pale paraphyses present. Scales and hairs: pinna- rachis closely warty beneath, with some residual pale scales to 7 by 14 mm, setiferous; costae with similar shorter elongate scales near base, for the most part covered with rather large pale brown bullate scales which have a few concolorous marginal hairs; near apex of costae a few long pale hairs; costules bearing similar bullate scales and occasionally hairs; no long hairs on veins but very short appressed hairs (bases of former scales ?) often conspicuous; hairs on upper surface of costules rare. Type specimen: HAsskKARL, Java (L?; not seen). Cited by RosENsTOCK: ZOLLINGER 2541, Mt Tengger, E. Java (L; dupl. at P). Distr. Malaysia: East Java, Lesser Sunda Is (Flores), South Celebes. Ecol. In open places at 1500-2300 m; locally abundant on Mt Tengger. Note. This species was established by ROSEN- STOCK by reference to the description of Also- Phila haenkei var. angustata HASsK., without further description; HASSKARL’s specimen, for which he cites no locality, is therefore the type. The above description is largely based on several later col- lections from Mt Tengger. 136. Cyathea persquamulifera (v. A. v. R.) Domin, Acta Bot. Bohem. 9 (1930) 146.—C. contaminans var. persquamulifera v. A. v. R. Bull. Jard. Bot. Btzg II, n.28 (1918) 13.—Alsophila persquamulifera v. A. v. R. ibid. III, 2 (1920) 130. Stipe scaly throughout, no long spines; basal scales to 30 by 1 mm, medium brown, shining, strongly setiferous (on a young frond). Main rachis closely warty beneath. Pinnae to at least 60 cm long, lower ones rather long-stalked. Pinnules to 135 by 25 mm, | or 2 basal segments free or contacted at base, rest of pinnule cut nearly to costa, sessile, caudate-acuminate; costules 314-4 mm apart; segments firm, edges often + reflexed when dry, crenate; veins to at least 12 pairs, lowest from costa or base of costule. Sori nearer costule than margin; no indusium, thin paraphyses not longer than sporangia. Scales and hairs: pinna- rachis beneath bearing very small fringed pale scales and copious narrow light brown strongly setiferous ones up to 10 mm long; costae at base as pinna-rachis, the larger scales 5-6 mm long, grading to very narrow setiferous scales and some bullate hair-pointed scales, also many very long (2 mm) stout brown hairs except near base; costules bearing copious similar hairs, sometimes narrow pale scales 2-3 mm long with marginal hairs or setae, always hair-pointed pale bullate scales; veins with a few stout erect hairs on lower surface; upper surface of costules and veins gla- brous. Type specimen: BUNNEMEIJER 961, Mt Talamau, 2300 m, Sumatra (BO; dupl. at K, L). Distr. Malaysia: Central Sumatra, throughout Java. Ecol. On mountains at 1500-2500 m. 137. Cyathea sarasinorum Hoittum, Kew Bull. 16 (1962) 61.—Alsophila contaminans var. longe- paleata Curist, Ann. Jard. Bot. Btzg 19 (1904) 42. —Alsophila glauca var. longepaleata v. A. Vv. R. Handb. (1908) 41; Suppl. (1917) 69. Main rachis pale beneath, with dark shining warts bearing dark shining setiferous scales to 15 by 1 mm. Pinnae to 67 cm long. Pinnules to 90 by 18 mm, lowest 1 or 2 lamina-segments constricted at base, rest of pinnule lobed almost to costa, apex rather shortly acuminate; costules 4 mm apart; segments of lamina thin, oblique, their rounded ends curved forwards, edges crenulate; veins 9-10 pairs. Sori near costules, exindusiate. Scales on costae beneath near base very narrow, dark shining brown, with many concolorous setae, smaller distally; pale spreading hairs also present towards apex of costae and on costules, on lower surface; scales on costules dark narrow, setiferous, also pale bullate scales bearing setae near their tips. "Type specimen: F. & P. SARASIN 2105, Si- baronga-Riicken, Central Celebes (BAS). Distr. Malaysia: Central Celebes (one collec- tion). Dec. 1963 ] CYATHEACEAE (Holttum) ge 138. Cyathea angiensis (GepP) DomIN, Acta Bot. Bohem. 9 (1930) 90.—Alsophila angiensis GEPP in Gibbs, Arfak (1917) 69. Like C. contaminans (WALL.) COPEL., but rather small; pinnules 100-150 by 16-20 mm; costules 314 mm apart, veins flat or impressed on lower surface; lamina-segments very firm, edges almost entire to crenate, glaucous beneath; more or less abundant pale flat setiferous scales present on lower surface of costae and costules; long pale spreading hairs more or less abundant towards apex of costae and on costules and veins on lower surface, rarely on upper surface of costules; pa- raphyses pale, longer than sporangia. Type specimen: Gipps 5968, Angi Lakes, 7000 ft, W. New Guinea (BM; K, P, BO). Distr. Malaysia: Moluccas Guinea. Ecol. Mountains, 600-2200 m. Note. A specimen from Sepik District, Ter- ritory of New Guinea (DARBYSHIRE & HooG- LAND 8191) bears the following information: “fronds immediately fully deciduous, the leaf- scars in distinct orthostiches (8) ...12 fronds in 3 whorls of 4, the outer whorl old, the middle whorl young fertile, the upper whorl not yet fully expanded, within each whorl the fronds very closely of the same age’’. DARBYSHIRE 384 has similar information, but fronds in whorls of 5. If I have correctly identified these specimens, the following characters distinguish this species from C. contaminans: fronds in whorls of 4 or 5, old fronds immediately and fully caducous. The re- maining specimens referred to C. angiensis do not bear such information. See also under 140. C. contaminans. (Buru), New 139. Cyathea verrucosa HoLttuM, Kew Bull. 16 (1962) 63. Stipe rather pale, closely warty throughout, persistently scaly near base only; scales to 40 by 3 mm, thin, pale, edges strongly dark-setiferous; main rachis, lower surface, glabrescent, closely warty (warts to 1 mm high). Pinnae to at least 60 cm long, lower ones with stalks 6 cm long. Pinnules to 110 by 20 mn, sessile or the lowest stalked, caudate-acuminate, basal 1-2 segments free or nearly so, rest cut almost to costa; costules 314-4 mm apart; lamina-segments rigid, edges minutely crenate, sinuses narrow; veins to 12 pairs. Sori at about 14 distance from costule to edge, marked by a depression on the upper surface, exindusiate; paraphyses pale, as long as sporangia, no scales round base of sorus. Scales and hairs: pinna- rachis beneath rather closely warty, bearing scattered pale setiferous scales to 10 by 1144 mm and very small pale setiferous or short-fringed scales; costae at first bearing many narrow pale setiferous scales to 2 mm long, these mostly caducous, also much shorter pale scales which are setiferous or the smallest pale-fringed, not bullate, distal half of costae bearing copious long pale spreading hairs with few scales; costules at first bearing many narrow pale setiferous scales to 2 mm long, also flat ovate-acute setiferous or fringed scales, and long pale spreading hairs; veins beneath bearing stout erect hairs (2 or 3 on a vein) and very short hairs which appear to be bases of fallen scales; upper surface of costules and veins glabrous. Type specimen: MATTHEW s.n., 3.2.1912, Mt Merapi, 5000 ft, Sumatra (K). Distr. Malaysia: Central Sumatra (two col- lections). Ecol. In open places in forest at 1600-1900 m. Note. Closely related to C. contaminans but much more densely scaly, the stipe closely warty instead of rather sparsely spiny. 140. Cyathea contaminans (WALL. ex Hook.) CopeL. Philip. J. Sc. 4 (1909) Bot. 60; v. A. v. R. Bull. Jard. Bot. Btzg II, n. 28 (1918) 13; Copet. Philip. J. Sc. 77 (1949) 115; Hottrum, Rev. FI. Mal. 2 (1954) 119; CopeL. Fern FI. Philip. 2 (1960) 230.—Polypodium contaminans WALL. Cat. (1828) n. 320, nomen.—Chnoophora glauca BL. En. Pl. Jav. (1828) 243 (non C. glauca Bory, 1804). —Alsophila glauca J. Sm. J. Bot. 3 (1841) 419; Bepp. Handb. (1883) 12; v. A. v. R. Handb. (1908) 41 (incl. var. celebica, var. squamulata, var. densa, var. setulosa, and var. microloba); Koorp.-ScHum. Syst. Verz. 1, 2 (1912) 5; Ro- SENST. Hedwigia 56 (1915) 349, incl. var. tricho- carpa ROSENST.; Vv. A. v. R. Handb. Suppl. (1917) 69 (incl. var. squamulosa); Bull. Jard. Bot. Btzg III, 2 (1920) 129; BACKER & PostH. Varenfl. Java (1939) 28.—Alsophila contaminans WALL. ex Hook. Sp. Fil. 1 (1844) 52, t. 18, f. 2; HaAssxk. in Hook. J. Bot. Kew Misc. 7 (1855) 323 (incl. var. robusta, var. squamulata, var. densa, var. microloba and var. setulosa HAssk.); BEDD. Ferns Br. Ind. (1865) pl. 85; Hook. Syn. Fil. (1866) 41; Scott, Trans. Linn. Soc. 30 (1874) 35; CuHrist, Verh. Nat. Ges. Basel 11 (1895) 199, incl. var. celebica Curist; Farnkr. Erde (1897) 327; Ann. Jard. Bot. Btzg 15 (1898) 79, pl. XIII f. 2; Diets in E. & P. Pfi. Fam. 1, 4 (1899) 136, p.p.—Alsophila acuta PresL, Abh. k. BOhm. Ges. Wiss. V, 5 (1848) 343. —Alsophila smithiana PREsL, I.c. 342.—Alsophila clementis Core. Philip. J. Sc. 1, Suppl. 2 (1906) Bot. 143.—C. clementis Copet. ibid. 4 (1909) Bot. 59; Fern Fl. Philip. 2 (1960) 230.—Fig. 3, 6, 22, 26. Trunk often very tall and much thickened by adventitious roots at base, only when old showing leaf-scars in upper part. Stipe to 100 cm long, glaucous, purplish towards the base, usually strongly spiny, at first scaly throughout, per- sistently so near the base; scales of all sizes up to 45 by 3 mm, pale brown, very thin, edges bearing close dark short setae. Main rachis pale, spiny, at first scaly as stipe, later + glabrescent. Pinnae: lowest somewhat reduced and with stalks to 10 cm long, largest 60 cm. Pinnules to c. 150 by 30 mm, often smaller, lowest distinctly stalked, largest with 1-2 pairs of basal segments more or less free, rest of pinnule lobed almost to the costa; costules commonly 4414 mm apart, rarely to 5 mm; segments of lamina firm, glaucous beneath, edges + crenate-serrate; veins commonly 12 pairs. 136 FLORA MALESIANA {ser. Il, volte Fig. 26. Cyathea contaminans (WALL.) CoPEL. in Tjibodas mountain garden (VAN STEENIS). Sori near costules, lacking indusia; paraphyses pale, not longer than sporangia. Scales and hairs: pinna-rachis more or less spiny, pale, glabrescent; lower surface of costae at first bearing scattered pale setiferous scales to 3 by 144 mm, shorter distally, these in most cases early caducous leaving the costae glabrous; costular scales small, + ovate, pale-fringed, mostly caducous, not bullate; a few hairs normally present towards apex of pinnules on both costae and costules, their abundance varying much; in a few localities (notably Mt Kinabalu, also in New Guinea) stout erect hairs, in variable number, may be present on lower surface of veins. Type specimen: WALLICH 320, Penang (K). Distr. Throughout Malaysia, in Peninsular India as far N as Mergui. Specimens so named from Hong Kong are C. /epifera (J. SM.) COPEL. Ecol. In clearings and open places in forest, especially near streams, 200 to 1600 m, often very abundant. Notes. This is the most widespread species of Cyathea in Malaysia, having no close allies at low elevations in Western Malaysia. In New Guinea there are several allied species, of which C. angiensis (GEPP) DOMIN appears to be nearest; it may perhaps better be united to C. contaminans. The several varieties of C. contaminans which have been described appear mostly to be due to the influence of environmental factors, or to be Dec. 1963] CYATHEACEAE (Holttum) 137 young, and therefore unusually scaly, at the time of collection. I have only seen one specimen of this species from NE. India; it was collected by Scott, who noted (/.c.) that typical A/sophila contaminans occurred there only at low altitudes. He distin- guished var. brunoniana (A. brunoniana HOoK.), which is common in Sikkim at higher altitudes and has smooth (not spiny) stipes and copiously scaly costae and costules. I prefer to regard this as a distinct species, which must be called Cyathea brunoniana (HooK.) CLARKE & BAK., though this name was actually used by CLARKE & BAKER for a quite different species. I hope to publish else- where descriptions of this and other species of the mainland of Asia. 141. Cyathea lepifera (J. SM.) Cope. Philip. J. Sc. 4 (1909) Bot. 40; ibid. 56 (1935) 98, pl. 3, fig. 4-7; Fern FI. Philip. 2 (1960) 228.—Alsophila lepifera J. SM. ex Hook. Sp. Fil. 1 (1844) 54; CurisT, Bull. Herb. Boiss. 6 (1898) 137, incl. var. congesta CHRIST; Vv. A. v. R. Handb. (1908) 39; Suppl. (1917) 65.—Alsophila pustulosa CHRIST, Bull. Herb. Boiss. Il, 1 (1901) 1019; Naxkat, Bot. Mag. Tokyo 41 (1927) 73; Ito, Fil. Jap. Ill. (1944) pl. 453.—Alsophila calocoma CurIsT, Philip. J. Sc. 2 (1907) Bot. 182; v. A. v. R. Handb. (1908) 789.—C. pustulosa CopeL. Philip. J. Sc. 4 (1909) Bot. 51.—C. calocoma CopPEL. /.c. 53; Fern Fl. Philip. 2 (1960) 229.—C. umbrosa COPEL. Philip. J. Sc. 56 (1935) 98, pl. 3, fig. 1-3.—(?) C. pteridioides Cope. l.c. 98, pl. 4; Fern FI. Philip. 2 (1960) 229. Stipe 16 cm or more, almost wholly covered with scales, not spiny (warty when scales have fallen); scales pale, thin, to 40 by 24 mm wide at base, apical part narrowly acuminate, edges throughout closely setiferous, narrow apical part sometimes entirely brown; smaller scales on stipe very narrow (under 14 mm wide), brown, with concolorous marginal setae. Rachis and pinna- rachis closely conspicuously warty, pale, more or less completely glabrescent, warts darker, to nearly 1 mm high. Pinnae: lowest somewhat reduced; largest to at least 80 cm long. Pinnules 100-150 by 15-22 mm, caudate-acuminate, sessile, largest with a few free or separately adnate basal segments, the rest lobed almost to the costa; costules 3-314 mm apart; veins 12-14 pairs; lamina-segments firm, entire or nearly so, glaucous beneath. Sori near costules; no indusia; paraphyses pale, longer than sporangia, some in form of narrow scales. Scales and hairs: pinna-rachis bearing some persistent appressed small pale fringed scales and occasionally longer spreading setiferous scales; costae beneath more or less densely scaly, basal scales elongate, narrow, pale with dark setae, distal ones smaller with pale marginal hairs, all flat, also some very small pale fringed scales throughout; at least the apical part of each costa also bearing stout pale hairs beneath; costules beneath bearing pale flat ovate to elongate scales bearing short pale marginal hairs, also more or less abundant stout spreading pale hairs; upper surface of costules lacking hairs. Type specimen: CuMING 180, Luzon (K; dupl. at BM, F, A, BO). Distr. Ryukyu Is, Formosa, Kwangtung, Hong Kong, in Malaysia: Philippines (Luzon, Panay, Mindoro, Babuyan). Ecol. In the Philippines a mountain species, but herbarium labels bear little ecological in- formation. I have been unable to find the type specimen of C. pteridioides CoPeL.; it is not in COPELAND’s herbarium. 142. Cyathea atrospinosa HoL_trum, Kew Bull. 16 (1962) 52. Trunk 6 m; fronds 350 cm long, in whorls of 4, 5 or 6, usually two whorls green at one time. Stipe to 125 cm long, lower part armed with dark spines to 8 mm long, also densely covered with scales which are up to 30 by 5 mm, thin, pale or pale brown, their edges bearing concolorous short hairs or dark setae. Rachis pale and glabrescent on lower surface, with scattered small thorns. Pinnae to 80 cm long. Pinnules to 120 by 25-30 mm, lowest on stalks up to 5 mm, lobed almost to the costa with the 1—2 lowest segments more or less free; costules 314-4 mm apart; lamina-seg- ments crenate, apices rounded, sinuses between them about 1 mm wide; veins 13-17 pairs. Sori exindusiate; paraphyses numerous, longer than sporangia. Scales and hairs: pinna-rachises pale beneath, glabrescent or with scattered setiferous scales; scales near base of costae shining brown, ovate, flat, long-setiferous, towards apices paler fringed scales; costules bearing pale brown flat fringed scales, and a few similar smaller ones also on lower surface of veins. Type specimen: HOoOGLAND & PULLEN 6090, Western Highlands District, 2650 m, NE. New Guinea (K). Distr. Malaysia: E. New Guinea. Ecol. Inmountain forest, on limestone (always? ) at c. 2400-2850 m. 143. Cyathea fugax v. A. v. R. Bull. Jard. Bot. Btzg Il, m. 7 (1912) 8: Handb. Suppl. (1917) 34; CopEL. Philip. J. Sc. 77 (1947) 116. Trunk to 10 m; fronds up to 300 cm long; leaf-scars 4 cm 9, in alternate whorls of 5 (BRAss 25585, Normanby I.). Stipe to at least 40 cm, spiny and rather densely scaly for most of its length; spines 2 mm or more long; scales pale, thin, to 25 by 2 mm, bearing many dark marginal setae near apices. Pinnae to 70 cm or more long. Pinnules to 145 by 35 mm, lowest 1—2 segments free, then some separately adnate, rest of pinnule cut nearly to costa, apex caudate-acuminate; costules 414 mm apart; lamina-segments rigid, glaucous beneath, little over 3 mm wide, almost entire; veins to 20 pairs, basiscopic basal vein from costa. Sori nearer costules than edge, ex- indusiate, with a ring of narrow fringed scales round base of receptacle; long paraphyses also present. Scales and hairs: pinna-rachis pale, bear- ing scattered short thorns, glabrescent; on costae 138 FLORA MALESIANA [ser. II, vol. 7 near base pale scales (apex dark or not) bearing dark flexuous setae, grading to smaller pale fringed scales without setae (or with a few) on costules and on veins; small scales on veins with short marginal hairs; very small hairs present on lower surface between veins; no hairs on upper surface of costules. Type specimen: CoPpLAND KING 215, Papua (BO; dupl. at MICH, SYD). Distr. Malaysia: E. New Guinea and adjacent islands. Ecol. In wet ground, secondary forest and open places in forest in low country up to 1400 m. Note. Details of trunk and stipe are taken from specimens other than type. Under type number at Sydney and Brisbane are also specimens of C. contaminans (WALL. ex HooKk.) COpPEL., which differ notably in scales from the type. 3b. Subsection Fourniera (BOMMER) HOLTTUM, stat. nov.—Fourniera BOMMER, Bull. Soc. Bot. Fr. 20 (1873) x1x.—Fig. 27. Type species: Cyathea novae-caledoniae (METT.) COPEL. Distr. The centre of distribution is New Guinea, with eastward and south extension to Samoa, New Caledonia and NE. Australia, westward and north to W. Java, Celebes, North Borneo (also Pulau Tio- man) and Philippines. Taxon. A distinction between species with long stipes and those with short stipes (the latter having gradually reduced lower pinnae) appears to be valid, but some collections do not include the stipe and so are not easy to place. More field study is needed in Eastern New Guinea. Ecol. These appear all to be forest species. KEY TO THE SPECIES 1. Pinnules 30-40 mm long; lower pinnae gradually reduced. 2. Tertiary leaflets on stalks almost 1 mm long. Pinnae to 25 cm long. Bullate scales present on costules. 144. C. carrii 2. Tertiary leaflets almost sessile. Pinnae to 45 cm long. No bullate scales on costules. 145. C. womersleyi 1. Pinnules commonly at least 70 mm long, in most cases much longer. 3. Lower pinnae gradually reduced, lowest 11 cm long near base of stipe. 146. C. auriculifera 3. Lower pinnae not thus reduced (sometimes 1-2 pairs of isolated small pinnae at base of stipe). 4. Bullate scales present on costae and costules. 5. Pinnules to 110 by 22 mm. Soral scales small, not covering sorus at maturity. 147. C. teysmannii 5. Pinnules to about 65 by 16 mm. Soral scales broad, covering sorus at maturity. 4. Bullate scales lacking. 148. C. aciculosa 6. Veins and lower surface of lamina bearing ee fringed ee scales which cover lower surface of leaflets 149. C. celebica 6. Veins bearing at most small separate scales on lower surface. 7. Scales absent from lower surface of veins . 150. C. tripinnata 7. Scales present on lower surface of veins, not long- fringed 7 nor r covering lower surface of leaflets. 144. Cyathea carrii HoLttum, Kew Bull. 16 (1962) 53.—Fig. 27. Stipe c. 7 cm, covered with a felt of very small dull scales, the larger ones setiferous, surface where exposed dark and smooth; larger scales near base to 30 by hardly 1 mm, edges bearing short dark setae, apex a long dark seta; lower surface of rachis dark brown, smooth or finely war- ty, glabrescent or with sparse minute dull scales. Lowest pinnae c. 3 cm long, succeeding ones gradually longer, up to 25 cm long. Pinnules to 32 by 7 mm, fully pinnate; tertiary leaflets to c. 12 pairs, distinctly stalked (stalks to almost 1 mm long and 2 mm apart), to 4 by a little more than 1 mm, edges sinuous, lowest sometimes 151. C. macrophylla deeply lobed at the base; veins 4-5 pairs. Sori up to 6 on each tertiary leaflet, covered to ripeness by broad pale lacerate overlapping scales which give the appearance of a complete indusium. Scales and hairs: pinna-rachis beneath dark, bearing many very small pale dull scales, some small bullate scales and scattered long narrow pale or partly dark closely setiferous scales; scales of pinnule-rachis (costa) small, brown, shining, mostly bullate and ending in a seta, some nearly flat and slightly elongate, rarely with lat- eral setae; scales of midribs of leaflets shining dark brown, bullate, acuminate, similar scales sometimes also on lower surface of veins; upper surface of tertiary leaflets glabrous. Dec. 1963] , CYATHEACEAE (Holttum) a Fig. 27. Cyathea carrii HOLTTUM.a. Single pinnule, upper surface, x 2, b. part of pinnule, lower surface, showing scales and sori, 6, c. sorus, showing covering of overlapping scales, x 20, d. scales from costule, < 30 (CARR 13526). Type specimen: CARR 13526, in forest, Boridi, 5000 ft, Papua (K; dupl. at BM, L). Distr. Malaysia: E. New Guinea (one col- lection). 145. Cyathea womersleyi Ho_ttrum, Kew Bull. 16 (1962) 63. Trunk to 8 m, fronds to 10, suberect, to 300 cm long, with pinnae gradually reduced to the base. Stipe 8-20 cm, sparsely spiny (spines to 214 mm), densely and persistently covered with scales which are pale, rather dull and soft, with a dark red seta at apex and no others, 10-20 by less than 1 mm, smaller ones 4-5 by !/; mm; main rachis persistently scaly beneath, smaller scales all with setiform apex, some elongate ones with pale edges bearing dark setae. Pinnae: lowest 8 cm long, longest 45 cm. Pinnules to 40 by 10 mm, almost fully pinnate; costules (midribs of tertiary leaf- lets) 2 mm apart; tertiary leaflets mostly 4-5 mm long, hardly stalked, slightly oblique, entire, apex rounded, base unequal, veins 6-7 pairs, lowest leaflets on largest pinnules to 7 mm long, bearing at base 1 or 2 pairs of 4th order leaflets which are round and entire, 1 mm long. Sori covered by overlapping fringed scales. Scales and hairs: lower surface of pinna-rachis densely scaly, some scales dark,shining,elongate with pale sparsely fringed or setiferous edges, some smaller, light brown and sub-bullate with setiform apex; near base of costae dark shining ovate to elongate scales with pale edges bearing some dark setae, then light brown nearly flat roundish scales with short setiform apex to pale hair-tip; on costules a few scales which usually lack setae but may bear marginal hairs; small scales sometimes on lower surface of veins. Type specimen: WoOMERSLEY & MILLAR NGF 8470, Skindewai, 5400 ft, Morobe District, NE. New Guinea (K; dupl. at L, BO, A, SYD). Distr. Malaysia: E. New Guinea (3 collec- tions). Ecol. At 1700-2400 m; “‘common through the rain-forest’’ (BRASS), in mixed Nothofagus forest (SCHODDE). 146. Cyathea auriculifera Cope. Philip. J. Sc. 6 (1911) Bot. 364; ibid. 77 (1947) 107. Trunk to 3 m; fronds 220 cm long, lower pinnae gradually reduced. Stipe 10 cm, copiously spiny (spines to 3 mm long), also densely covered with pale scales 10-55 mm long, larger ones twisted and rather straight, edges closely set with short dark setae; main rachis beneath rather closely spiny throughout, covered with a dark brown felt of very small mostly setiferous scales and at first with elongate pale scales attached to the thorns. Pinnae: lowest 11 cm long, largest 54 cm. Pinnules slightly overlapping, to 70 by 15 mm, fully pinnate, bases of tertiary leaflets 3 mm apart; rertiary leaflets mostly stalked, largest 9 by 214 mm, several pairs near base of pinnule having 1 or 2 pairs of free quaternary leaflets, acroscopic basal tertiary leaflets deflexed across pinna-rachis; veins to 9 pairs, those in quaternary leaflets pin- nately branched, rest mostly forked. Sori nearer to costules than to edge, covered with overlapping fringed scales until nearly ripe, fringe of these scales longer than on costal scales. Scales and hairs: pinna-rachis scaly beneath as main rachis; costae copiously scaly, larger scales 1 mm long, ovate-acute, dark and shining with pale edges bearing dark setae, smaller ones entirely pale, either with setae or fringing hairs, no bullate scales; costules and veins beneath bearing similar smaller scales; upper surface of costules glabrous. Type specimen: C. KING 227, Goodenough Bay, 1200 m, Papua (MICH; fragm. at BM). Distr. Malaysia: New Guinea and Louisiades. Ecol. In mountain forest or transition to mossy forest, on mainland 1200-2600 m, on the islands 750-1000 m. 147. Cyathea teysmannii Cope. Philip. J. Sc. 4 (1909) Bot. 51 (new name for C. celebica v. A. v. R.); v. A. v. R. Handb. Suppl. (1917) 39.— Hemitelia truncata (non BRACK.) CHRIST, Ann. Jard. Bot. Btzg 15 (1898) 81; in Warburg, Mon- sunia (1900) 91.—C. celebica v. A. Vv. R. Bull. Dép. Agr. Ind. Néerl. 18 (1908) 2, non BL. 1828; Handb. (1908) 26. Differs from C. tripinnata in the presence of distinctly bullate pale scales on the costules, and in thinner and smaller soral scales which do not 140 FLORA MALESIANA [ser. I, yolue cover the fully developed sporangia. The speci- mens do not give information about characters of the stipe and its scales, which may also be distinctive. Type specimen: TEYSMANN 13681 (BO). Distr. Malaysia: SW. Celebes. Ecol. At c. 1000 m altitude. 148. Cyathea aciculosa CopeL. Philip. J. Sc. 60 (1936) 104, pl. 9.—C. arachnoidea (non Hoox.) GRETHER & WAGNER, Un. Cal. Publ. Bot. 23 (1948) 43. Stipe 18 cm long, dark, smooth, covered with minute scales, also near base with pale scales 25 by less than 1 mm, bearing dark marginal setae near apices. Pinnae: lowest 10 cm long, largest 50 cm. Pinnules to 65 by 16 mm, fully pinnate; tertiary leaflets to 20 pairs, to 7 by almost 2 mm, larger ones crenate; veins 7 pairs. Sori covered with overlapping pale scales; no true indusium. Scales and hairs: pinna-rachis warty, shining, bearing many very small pale short-fringed scales and a few long narrow setiferous ones, these latter also on bases of costae; most scales on costae pale, bullate, also very small pale scales. Type specimen: Brass 2887, San Christoval, 900 m, Solomon Is (MICH; dupl. at L, BRI). Distr. Solomon Is, in Malaysia: Admiralty Is. Ecol. In forest, to c. 1000 m. Note. This species is very near C. truncata BRACK.) CopPeEL. of Fiji and Samoa, differing in the much larger indusial scales. 149. Cyathea celebica BL. En. Pl. Jav. (1828) 245 (not v. A. v. R. 1908); TINDALE, Contr. N.S.W. Nat. Herb. 2 (1956) 338, p.p.—Alsophila celebica Metr. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 53; v. A. v. R. Handb. (1908) 42.—C. arachnoidea Hook. Syn. Fil (1865) 24; v. A. v. R. Handb. (1908) 26; Suppl. (1917) 38.—Alsophila truncata var. Sagittata CHRIST, Bull. Herb. Boiss. II, 1 (1901) 458.—Alsophila truncata var. nivea CHRIST, K. Sco. & Laut. Nachtr. (1905) 36; v. A. v. R. Handb. (1908) 42.—C. quadripinnatifida CopeEL. Un. Cal. Publ. Bot. 18 (1942) 218; Philip. J. Sc. 77 (1947) 108, pl. 6. Stipe to 100 cm long, dark, bearing slender dark spines to 3 mm long, near base bearing persistent light brown scales 20-40 mm long, mostly not over 1 mm wide, firm and shining, edges bearing close short oblique concolorous setae, rest of stipe covered closely with small interlacing setiferous or fringed scales. Pinnae: small pinnae (5-8 cm long) sometimes present near base of stipe (seen only in two specimens); largest pinnae to 70 cm long. Pinnules 90-140 by 17-28 mm, fully pinnate, the lower tertiary leafiets stalked; costules (stalks of tertiary leaflets) 3-5 mm apart; tertiary leaflets to 15 by 214-5 mm, sometimes dilated at the base, the largest deeply lobed near the base, the basal lobes sometimes forming free quaternary leaflets. Sori medial, protected when young by overlapping very thin finely fringed scales. Scales and hairs: pinna-rachis and costae beneath almost covered with very small irregular pale scales bearing short marginal hairs and some larger scales bearing setae; costules at first bearing small ovate scales bearing setae, lower surface of costules and veins covered throughout with small pale fringed scales which interlace and completely cover lower surface of lamina. Type specimen: REINWARDT, Ternate (L; dupl. at BM). Distr. Queensland, in Malaysia: New Guinea, Moluccas (Ternate, Ambon), ? Celebes. Ecol. At 100-1750 m. Note. The type of C. quadripinnatifida was in open forest in a ravine and had 4 fronds 600 cm long (including stipe of 100 cm); these do not show reduced basal pinnae. 150. Cyathea tripinnata Cope. Philip. J. Sc. 1, Suppl. 4 (1906) Bot. 251; ibid. 4 (1909) Bot. 40; v. A. v. R. Handb. (1908) 788; Suppl. (1917) 39; Ho.Lttum, Rev. FI. Mal. 2 (1954) 120; Copet. Fern Fl. Philip. 1 (1960) 208.—C. densisora v. A. v. R. Bull. Jard. Bot. Btzg III,2 (1920) 138; CopeE-. Fern Fl. Philip. 2 (1960) 208. —C. leucostegia CopeEL. Philip. J. Sc. 38 (1929) 130; Fern FI. Philip. 2 (1960) 209.—C. leytensis CopEL. Philip J. Sc. 38 (1929) 131; Fern FI. Philip. 2 (1960) 209. —C. arachnoidea (non HooK.) BACKER & POSTH. Varenfl. Java (1939) 25. Trunk to 4or 5m. Stipe to at least 40 cm, dark, bearing scattered sharp spines 1-3 mm long, covered almost throughout by a felt of very small setiferous scales; basal scales to 25 by 1 mm, thin and soft, matted together, edges with some dark setae, apex a dark seta. Pinnae: lowest 20-30 cm long, largest to 60 cm. Pinnules 90-140 by 17—25 mm, fully pinnate, lower tertiary leaflets distinctly stalked; costules 4-6 mm apart; tertiary leaflets to 15 by 3!4mm, the larger ones deeply lobed at the base, edges crenate; veins to 9 pairs, those in basal lobes pinnate. Sori near costules, covered to maturity by overlapping pale thin scales. Scales and hairs: lower surface of pinna-rachis covered with minute pale fringed scales; costae bearing similar scales with some longer narrow ones bearing a fringe of hairs or dark setae; on costules ovate flat brown to pale scales, setiferous or fringed; on veins no scales. Type specimen: CoPELAND 2068, Mt Mariveles, in extinct crater, 900 m, Luzon (MICH; dupl. at LUIS), 12, WK, SYAD. SHAY) Distr. Malaysia: West Java, Pulau Tioman (E off Malaya), N. Borneo, Philippines (Luzon to Mindanao), Moluccas (Ambon). Ecol. In forest, 250-1700 m; the smaller forms, represented by C. densisora, C. leytensis and C. leucostegia probably in more exposed places. 151. Cyathea macrophylla Domin, Acta Bot. Bohem. 9 (1930) 133 (mew name for Hemitelia ledermannii BRAUSE); CopEL. Philip. J. Sc. 77 (1947) 108.—Hemitelia ledermannii BRAUSE, Bot. Jahrb. 56 (1920) 60. Differs from C. tripinnata in firmer texture of lamina and in presence of more or less abundant small scales on lower surface of veins, these scales Dec. 1963] CYATHEACEAE (Holttum) 141 bearing short dark setae or short pale hairs, never long-fringed nor covering lower surface of lamina. foliolis tertiariis maximis 18 mm longis, omnino profunde lobatis, segmentis infimis interdum liberis Type specimen: LEDERMANN 12533, Sepik et stipitulatis. Region, E. New Guinea (B). Type specimen: WomerRSLEY NGF 13959, Mo- Distr. Malaysia: New Guinea. robe Distr., NE. New Guinea (K). Ecol. In forest, from sea-level to 1500 m. Ecol. “Short tree-fern in under-storey of the forest. Trunk not more than 2 feet tall. 5 feet’, 2000 m. ay Fronds var. quadripinnata HOLTTUM, var. nov. A typo speciei differt: stipitibus c. 10 cm longis; 4. Section Schizocaena (J. SM.) HOLTTUM, stat. nov.—Schizocaena J. SM. in Hook. Gen. Fil. (1838) t. 2; Lond. J. Bot. 1 (1842) 661, p.p.; CopEL. Gen. Fil. (1947) 99, p.p.—Fig. 8c, 28-30. Type species: Schizocaena brunonis J. SM. = Cyathea moluccana R. Br. Distr. Malaysia and Polynesia. Taxon. The division into two subsections is perhaps not sharp, but extreme examples of subsect. Sarcopholis are strikingly different from species of subsect. Schizocaena from Western Malaysia. The characteristic feature of the section is the position of the basal basiscopic vein of each vein group; this vein springs from the costa, not from the costule (or in C. moluccana from the midrib of the pinna). This condition is correlated with the relatively shallow lobing of the pinnules; similar lobing in subg. Cyathea sect. Gymnosphaera is usually not associated with this vein-character. The species C. sinuata Hook. and C. hookeri THw., of Ceylon, included here by J. SmitH and Cope- LAND, have flabelloid scales; in my opinion their nearest relatives are in the group of C. borbonica DEsv. of Madagascar. 4a. Subsection Schizocaena Distr. Malaysia, except Moluccas and New Guinea.—Fig. 8c, 28-29. Taxon. In Borneo and the Philippines species of this subsection are often difficult to characterize learly, appearing to vary in size and in distribution of scales and hairs on axes. KEY TO THE SPECIES 1. Fronds simply pinnate, pinnae entire or at most serrate-crenate. 2. Apex of frond a deltoid deeply lobed lamina. Outer veins of each group joining to form a single excurrent vein. Pinnae sessile, base truncate or subcordate . aa 2 1S22 CC eapiata 2. Apex of frond a pinna of same shape as other pinnae. No anastomosis. Pinnae usually stalked. 3. Pinnae not over 15 mm wide, base narrowly cuneate, stalkes to 12 mm long. Sori in 1-2 rows on each side of costa, fully indusiate : si 153. C. angustipinna 3. Pinnae 2-4 cm wide, base rather broadly cuneate, ‘stalked o or not. Sori i in fully fertile fronds in more — 2 rows, indusiate or not. . Pinnae not long-acuminate, upper usually sessile . - Pinnae long-acuminate, all stalked . 1. Fronds simply pinnate with deeply lobed pinnae, or bipinnate. 5. Fronds simply pinnate with deeply lobed pinnae, the largest sometimes with free pinnules at their base. 6. Lower surface of pinna-midribs covered near base with long hairs; no free pinnules. 156. C. deminuens 6. Lower surface of pinna-midrib lacking hairs; some free pinnules on largest pinnae. 7. Indusium present (sometimes hidden by mature sorus). 8. Lower pinnae little narrowed at base, free leaflets as zone as lobes, apex not long-acuminate; pinnae commonly 252 Cmielon tar. w. = = ce 157-2@>altermans 8. Lower pinnae narrowed to base so that free leaflets a are , very small, apex long-acuminate; pinnae to 18 cm long : 158. C. binuangensis 7. Indusium lacking. 9. Pinnae commonly 25 cm long; no long pale hairs on rachis . 9. Pinnae much shorter; main rachis bearing long pale hairs . 5. Fronds amply bipinnate. 10. Pinnules entire or with crenate edges. 11. Indusia present; pinnules sessile 11. Indusia lacking; pinnules stalked 10. Pinnules distinctly lobed. 12. Sori indusiate. 13. Pinnules of larger pinnae not lobed more than 34, to costa throughout, no free basal pinnules, Or sometimes one on lowest pinnule of lower pinnae. 154. C. moluccana 155. C. arthropoda 157. C. alternans 159. C. elliptica 157. C. alternans 160. C. obliqua 142 FLORA MALESIANA [ser.. TI; vols 14. Bullate scales lacking on costules . 14. Bullate scales present on costules. 161. C. integra 15. Basal pinnules of middle pinnae with stalks to at least 4 mm long; pinnules lobed less than 1, way to costa 162. C. stipitipinnula 15. Basal pinnules sessile or nearly SO; ypiinules lobed more than half way to costa. 16. Segments of pinnules acute, falcate; largest pinnules 20-30 mm wide. 163. C. zamboangana 16. Segments of pinnules rounded; largest pinnules not over 20 mm wide. 17. Indusium a disc hidden by sorus . 17. Indusium complete, breaking and persistent at ‘maturity. 18. Lower surface of costae densely scaly . 18. Lower surface of costae not densely scaly . 5 13. Pinnules of larger pinnae always with one or more free basal segments. 164. C. discophora 165. C. megalosora 166. C. suluensis 19. Copious long hairs on lower surface of veins as well as costules. 20. About half the tertiary segments free or separately adnate to costa on largest pinnules. Pin- nules to 100 by 20 mm. Costae not densely scaly. . . i to, MULGHE C. robinsonii 20. One or two pairs tertiary segments free on largest pinnules. Pinnules to 80 by 12 mm. Costae ieee scaly. Costal scales not setiferous. Pinnules sessile . 5 Costal scales strongly setiferous. 19. No hairs on lower surface of veins. 165. C. megalosora Lower pinnules with stalks 2-3 mm long. 168. C. senex 22. Free basal segments of largest pinnules deeply lobed. Rachis and pinna-rachis densely persistently scaly, scales small, sub-bullate, setiferous . 169. C. sibuyanensis 22. Free basal segments entire. Rachises glabrescent . 23. Pinnules sessile, to c. 65 by 15 mm, often smaller. . . 23. Pinnules stalked (stalks to 8 mm), to 100 by 25 mm . 12. Sori without indusia. 170. C. philippinensis 171. C. assimilis 24. Long-spreading hairs abundant on lower surface of rachis and/or pinna-rachis, often also on costae. 25. Costules and veins bearing hairs like those of costae on lower surface. 26. Pinnules to 110 mm long, cut 24-34, to costa; costules 5 mm apart . 172. C. trichodesma 26. Pinnules to 65 mm long, cut to within 1 mm of costa; costules 3-314 mm apart. 173. C. wallacei 25. Costules and veins lacking hairs on lower surface. 27. Hairs of lower surface confined, or almost confined, to main rachis . 159. C. elliptica 27. Hairs of lower surface present on pinna-rachis and costae, few or none on main rachis. 174. C. trichophora 24. Long spreading hairs lacking on lower surface of rachis and pinna-rachis. 28. Largest pinnules with a free segment at the base; pinnules on stalks to 4 mm or more long; texture firm . 28. Largest pinnules lacking. a free segment at the base, ‘almost sessile. 175. C. polypoda 29. Sori on 3-4 pairs of basal veins in each group only, not on distal veins, at maturity confluent. 176. C. obscura 29. Sori on almost all veins, not confluent at maturity. 30. Bullate scales lacking on costae and costules. Pinnules cut to less than 1 mm from costa; stout erect hairs abundant on upper surface of costules and veins . 177. C. agatheti 30. Bullate scales present on costae and costules. Pinnules less deeply lobed; hairs on upper surface of costules few 152. Cyathea capitata Corer. Philip. J. Sc. 12 (1917) Bot. 49; C. Cur. & HoLtrum, Gard. Bull. S. S. 7 (1934) 199, 218.—Schizocaena capitata CopEL. Gen. Fil. (1947) 99.—Fig. 28f-h. Trunk \—3 m, bearing c. 12 fronds. Stipe dark, smooth, at least 40 cm; basal scales pale brown, firm, to 25 by 3-4 mm, rather thick at the base, edges bearing rather irregular concolorous setae; pneumathodes 9-15 mm long, rather widely spaced. Rachis dark to medium brown, smooth; lamina 100 cm or more long, simply pinnate, apex of frond not like a pinna but broadly deltoid and deeply lobed, lobes grading to upper pinnae. Pinnae c. 40 pairs, sessile, jointed to rachis, largest 15-19 by 2-3 cm wide at base, lower ones some- what smaller, edges entire except near apex, base .178. C. squamulata truncate to cordate, in the latter case more or less auricled on both sides, apex short-acuminate, crenate. Veins pinnate in each group, usually with 3 pairs of veinlets, outer veinlets of each group always anastomosing with outer veinlets of ad- jacent groups to form a single excurrent vein; basal basiscopic vein of each group springing separately from the costa. Sori usually in 2 rows on each side of the costa (3-4 sori on each vein- group) or in narrower pinnae only one row; indusium thin and translucent, at first covering sorus, later breaking irregularly and more or less persistent. Type specimen: CLEMENS 11033, Mt Kinabalu, N. Borneo (MICH; dupl. at A, UC, K). Distr. Malaysia: Borneo (Sarawak: Mt Dec. 1963] Murud; N. Borneo: Mt Kinabalu). Ecol. Near a waterfall and in wet ground near a stream, in forest, 1400-2100 m. 153. Cyathea angustipinna Ho_trumM, Kew Bull. 16 (1962) 52. Stipe 30 cm, smooth, scaly near base, scales pale, firm and shining, to c. 20 by 114 mm, edges bearing copious short dark setae. Frond 70 cm long, simply pinnate; pinnae about 18 pairs, jointed to rachis; terminal pinna like the rest, usually with a small rudiment beside it. Pinnae stalked to 12 mm or more (lowest longest), to 12 cm long, fertile ones 1—1.2 cm wide, sterile to 1.6 cm, bases narrow- ly cuneate (of lower ones somewhat asymmetric) apices shortly acuminate, edges entire except to- wards apices where they are crenate; veins common- ly in groups of three, middle one sometimes forked, basiscopic one separately attached to costa. Sori 2 or 3 on each vein-group (middle vein not always soriferous), in two rather uneven rows on each side of the costa; indusium thin and pale, covering young sorus and breaking later, more or less persistent as a disc around base of old sorus; costae usually quite glabrous on both surfaces. Type specimen: RICHARDS 1675, Mt Dulit, 1200 m, Sarawak (K). Distr. Malaysia: collections). Ecol. Sandy bank of a stream, near waterfall, in shade; trunk 50 cm. Borneo (Sarawak, two 154. Cyathea moluccana R. Br. in Desv. Mém. Soc. Linn. Paris 6 (1827) 322; v. A. v. R. Handb. (1908) 15; CopeL. Philip. J. Sc. 4 (1909) Bot. 32; v. A. v. R. Bull. Jard. Bot. Btzg I, n. 28 (1918) 12.—Schizocaena brunonis J. SM. ex Hook. Gen. Fil. (1838) t. 2; J. Sm. Hist. Fil. (1875) 243.— C. brunonis WALL. ex Hook. Sp. Fil. 1 (1844) 15; Syn. Fil. (1865) 16; Bepp. Ferns Br. Ind. (1865) pl. 87; Handb. (1883) 5; CuHrist, Farnkr. Erde (1897) 318; Diets in E. & P. Pfl. Fam. 1, 4 (1899) 127; CopeL. Sarawak Mus. J. 2 (1917) 345, 347; HoL_trum, Rev. Fl. Mal. 2 (1954) 117.— C. pinnata Roxs. Calc. J. Nat. Hist. 4 (1844) 517; Fl. Ind. ed. Clarke (1874) 762.—Schizocaena gaudichaudii Fre, Gen. Fil. (1852) 354.—C. pseudobrunonis CoreL. Philip. J. Sc. 12 (1917) Bot. 50.—C. fuscopaleata CopeL. I.c. 50.—C. kinabaluensis Cope. /.c. 51; C. Cur. Gard. Bull. S.S. 7 (1934) 218.—Schizocaena moluccana Co- PEL. Gen. Fil. (1947) 99.—Schizocaena kinaba- luensis COPEL. I.c. 99.—Fig. 28a-e. Trunk usually not over 50 cm tall. Stipe common- ly 20-30 cm, dark, scaly near base and finely warty when scales have fallen; scales medium brown, firm, 15-30 by 14-3 mm, edges bearing setae little darker than scale. Lamina to 150 cm or more long (largest frond reported, stipe with lamina 308 cm), simply pinnate, apical pinna usually like the rest (sometimes with a rudiment at its base), all pinnae articulate to rachis. Pinnae stalked or the upper ones sessile (stalks variable in length, the lowest 5-10 mm), 12—28 by 2-4 cm, edges parallel for most of their length, base asym- CYATHEACEAE (Holttum) ze hi metric (rounded on acroscopic side, broadly cuneate on basiscopic), apex shortly acuminate and crenate, edges otherwise entire, rarely slightly lobed with one lobe to each vein-group; veins in groups of 3 from the costa (basiscopic one usually separate from the other two), the median one forked once or twice to give a group of 3-6 (rarely to 10) veins at the edge, veins all free and all ending close to the margin, or sometimes the acroscopic vein ending in a sorus or joining with another vein. Sori in 1—3 (rarely more) rows on each side of the midrib, commonly 4—6 on each vein-group, exceptionally to 10 (in such cases the edge usually lobed), covered when young by a thin translucent indusium which breaks at ma- turity and is more or less persistent, or in some cases the indusium forming a disc which is hidden by the mature sporangia; lower surfaces of rachis, costae and veins usuaily glabrous, occasionally a few persistent small ciliate or setiferous scales present. Type specimen: C. SmitH, Moluccas (BM). Distr. Malaysia: Central Sumatra, Malay Peninsula, Lingga, Borneo (excluding south and south-west), South & Central (?) Celebes, Mo- luccas (Ceram, Ambon). Ecol. In forests, 0-900 m. Notes. Fronds bearing imperfect indusia appear to be most common in Borneo, but occur also in the Malay Peninsula. COPELAND described the three species C. pseudobrunonis, C. fuscopaleata and C. kinabaluensis as lacking indusia, but I have not found a specimen totally devoid of indusia, though in old specimens only small fragments remain (VAN ALDERWERELT made the same observation, 1918 /.c.). COPELAND thought that species could also be distinguished by size and colour of stipe-scales, but, after seeing a very large number of specimens, I cannot see any clear distinction into groups based on such charac- ters. COPELAND’s fourth species, C. arthropoda, is here regarded as distinct in shape of pinnae. R. Brown described this species, but did not name it, in 1810 (Prod. Fl. N. Holl. 158); DEsvAUXx supplied a name, copying the information provided by Brown. The type must be the specimen seen from the Moluccas by Brown, though it is not named by him. 155. Cyathea arthropoda Copev. Philip. J. Sc. 6 (1911) Bot. 134, t. 13; v. A. v. R. Handb. Suppl. (1917) 22.—Schizocaena arthropoda Copel. Gen. Fil. (1947) 99. Fronds simply pinnate; lamina to c. 70 cm long, apical pinna like the rest. Pinnae jointed to rachis, always stalked (stalks of upper pinnae c. 5 mm, of lower ones 12-15 mm), 12-20 cm long, fertile to 214 cm wide, sterile to 314 cm, base almost equally cuneate, apex caudate-acuminate (cauda to 4.cm long), sides for the most part not parallel. Sori in 1-3 irregular rows on each side of the costa; indusium a narrow irregular ring, hidden by sporangia, or lacking. Type specimen: Brooxs 8, Bungo Range, Sarawak (MICH; dupl. at BM). 144 FLORA MALESIANA [ser Ik voles Fig. 28. Cyathea moluccana R. Br. a. Middle part of rachis with pinnae, * 14, b. apex of frond, x 4, c. part of pinna, showing venation and sori, * 114, d. scale from stipe, < 4, e. edge of stipe-scale, * 40. —C. capitata Core. f. Middle part of rachis with pinnae, < 14, g. apex of frond, 14, A. part of pinna showing venation and sori, * 114 (a—e CUMING 378, f CLEMENS 27959, g JAcoBs 5790, h CLEMENS 11033). Dec. 1963] CYATHEACEAE (Holttum) 145 Distr. Malaysia: Sarawak (neighbourhood of Kuching). Ecol. In lowland forest. Note. This seems to be the most distinct of the ““exindusiate’” species described by CopELAND from Sarawak (see C. moluccana). Field study is necessary to decide whether or not it is connected by intermediate with typical C. moluccana, which occurs in the same district. 156. Cyathea deminuens HOLTTUM, nom. nov.— Alsophila parvifolia Hotttum, J. Mal. Br. R. As. Soc. 6 (1928) 19. Trunk to 75 cm. Stipe to 15 cm; scales at base pale brown, rather firm, 10-15 mm by 114 mm, edges closely set with dark setae. Lamina to 65 cm long, simply pinnate, pinnae pinnatifid, lower ones gradually reduced, lowest 314 cm long. Rachis densely covered beneath throughout with stout pale spreading hairs to 3 mm long. Largest pinnae 9 by 2 cm, lobed 24 towards the costa; costules 5 mm apart; veins 6—7 pairs, basal basiscopic vein from costa; lower surface of costae hairy towards base, and a few hairs on lower costules, no bullate scales seen; a few long hairs present on upper surface of costules and veins. Sori at about one third distance from costule to edge; no indusia; long paraphyses, with dark walls between the cells, present. Type specimen: BoDEN KLoss 14579, Siberut, Mentawai Is (S; dupl. at BO; K). Distr. Malaysia: Sumatra (two collections, the second from S. Sumatra). Ecol. In lowland forest. 157. Cyathea alternans (WALL. ex HooK.) PRESL, Abh. K. B6hm. Ges. Wiss. V, 5 (1848) 347; Bepp. Handb. Suppl. (1892) 2; v. A. v. R. Handb. (1908) 17; Rip. J. Mal. Br. R. As. Soc. 4 (1926) 5, incl. var. serrata RIDL.; Domin, Acta Bot. Bohem. 9 (1930) 90, incl. var. sarawakensis (HOOK.) Domin and var. lobbiana (HooK.) DoMIN; HOLTTUM Rey. FI. Mal. 2 (1954) 119.—Polypodium alternans WALL. Cat. (1829) n. 329, nom. nud.—Hemitelia alternans Hook. Ic. Pl. (1844) t. 622; Sp. Fil. 1 (1844) 29.— Amphicosmia alternans Moore, Ind. Fil. (1857) civ; Bepp. WHandb. (1883) 10.—C. lobbiana Hoox. Syn. Fil. (1865) 24.—C. sarawakensis Hoox. lc. 23; Hose, J. Str. Br. R. As. Soc. 32 (1899) 35.—Alsophila alternans Hook. Syn. Fil. (1866) 41; Bepp. Ferns Br. Ind. (1867) t. 236.—Schizocaena alternans J. Sm. Hist. Fil. (1875) 244; Coper. Gen. Fil. (1947) 99.—Alsophila janseniana v. A. v. R. Bull. Jard. Bot. Btzg III, 5 (1922) 179.—C. janseniana Domin, Acta Bot. Bohem. 9 (1930) 126. Trunk usually less than 2 m. Stipe to 60 cm, dark, persistently scaly near base, otherwise glabrescent and smooth; scales medium brown, firm, shining, to 30 by 2 mm, edges closely seti- ferous. Pinnae articulate to rachis, lowest some- what reduced, largest commonly 25 by 4-5 cm, sometimes to 40 by 9 cm (in type of C. sarawaken- sis 50 by 16 cm), deeply lobed throughout or with few to many of the lobes separately joined by the more or less contracted base of their lamina to the axis of the pinna, rarely the lowest one or two forming true pinnules with lamina quite free; costules of lobed pinnae 7-9 mm apart, of pinnae in which the lobes are free up to 15 mm apart; lobes which are not free usually entire and rounded at apex, free lobes acute at the apex or with margins sometimes broadly crenate, one crenation to each vein-group. Veins in the narrower, lobed, pinnae to c. 10 pairs in each lobe, individual veins forked Once or twice; in free lobes the veins in small lateral pinnate groups. Sori usually in one row on each side of the costule of a lobe of a pinna, in the largest free lobes occasionally an incomplete second row; indusium varying greatly, in a mi- nority of cases completely covering the young sorus and persistent, in most cases forming a disc of irregular width covered by the mature sorus, sometimes only detectable as small fragments; long pale paraphyses present. Scales on lower surface of pinna-midribs, costules and veins usual- ly rather sparse, elongate, narrow and strongly setiferous, pale to medium brown, in some cases also bullate scales on costules; in some cases stout pale hairs present on distal parts of costae and costules and a few on veins. Type specimen: WALLICH 329, Penang (K). Distr. Malaysia: Sumatra (Karo plateau, Benkulen Distr.), Malay Peninsula (Penang, Pe- rak, Negri Sembilan, Pahang, Trengganu, Kelan- ton), Sarawak and North Borneo. Ecol. In forest,often near streams,at 300-1300 m. Note. This species is usually found growing in association with C. moluccana R. Br. and always within the range of C. moluccana. The variable C. alternans has the appearance of being a series of hybrids between C. moluccana and either C. squamulata or C. ridleyi (which are bipinnate and exindusiate) with many possible combinations of characters of pinna-shape and of indusia and scales and hairs. The type specimen of C. alternans has pinnae to about 614 cm wide, with 3 pairs of free adnate lobes, a few lobes longer than adjacent ones, as in the type of Alsophila janseniana v. A. v. R.; the sori have disc-shaped indusia, as shown in Hooker, Ic. Pl. t. 622, though Hooker stated later (Syn. Fil. 41) that ‘“‘a depression formed by the sorus on the lobe was mistaken by the artist for an involucre”’. Hairs occur on the lower sur- face of costules in some Peninsular specimens (especially Mou. Nur 11269); I have not seen any on Bornean ones. Bullate scales appear more common in Peninsular than in Bornean specimens. 158. Cyathea binuangensis v. A. v. R. Bull. Jard. Bot. Btzg III, 2 (1920) 136; CopeL. Fern Fl. Philip. 2 (1960) 206. Stipe 20 cm, densely scaly at base only; scales pale, thin, to 20 by 2 mm, edges bearing concol- orous hairs. Lamina 50 cm long, simply pinnate. Pinnae: lowest slightly reduced; largest 18 by 4 cm, lobed to 3-4 mm from costa except at base where one segment is usually almost free; bases of pinnae conspicuously narrowed (free segment always smaller than rest), apex rather long-acu- minate; lobes falcate, almost entire, their costules 146 FLORA MALESIANA [ser. II, vol: 12 to 9 mm apart; veins to about 10 pairs, mostly forked. Sori medial; indusium thin, breaking and persistent; paraphyses very dark. Scales on costae of pinnae very few, very narrow, pale, bearing sparse short concolorous hairs; no hairs seen on lower surface of pinnae, on upper surface present only near base of costae. Type specimen: RamMos & EDANOo, BS 28779, Mt Binuang, Luzon (BO; dupl. at US, MICH, K). Distr. Malaysia: Philippines (Luzon, one collection). Note. This specimen has the aspect of a pre- maturely fertile plant of C. integra J. SM. ex HOOK., but it lacks the characteristic scales of that species. It has the same degree of division of the frond as C. alternans (WALL.) PRESL, but is very different in shape of pinnae, and in paraphyses. It might be a hybrid of C. integra, but I cannot suggest what other parent is likely. 159. Cyathea elliptica Cope. Philip. J. Sc. 12 (1917) Bot. 51.—C. subbipinnata Cope . ibid. 56 (1935) 471, pl. 1.—C. holttumii Corer. I.c. 472, pl. 2.—Gymnosphaera subbipinnata CopeL. Gen. Fil. (1947) 99.—Gymnosphaera holttumii COPEL. es 99: Stipe scaly throughout, also bearing on abaxial surface some long spreading hairs, the latter some- times very abundant and to 10 mm long; scales medium to light brown, shining, edges setiferous; main rachis similarly scaly and hairy, scales some- times deciduous. Pinnae: lower ones not greatly reduced, largest 30-45 cm long. Largest pinnules 4.5-6.5 cm long, 10-14 mm wide, sessile, lobed half way to costa; costules 3!4-4 mm apart; veins to 5 pairs; lamina-segments rather thin, entire, ends rounded. Sori medial; no indusia; paraphyses not longer than sporangia. Scales and hairs: lower surface of pinna-rachis lacking hairs or a few present near the base; costae bearing sparse narrow pale scales and sometimes a few hairs; costules bearing pale hair-pointed bullate scales. Type specimen: CLEMENS 10859, Mt Kinabalu, N. Borneo, (MICH; dupl. at UC). Distr. Malaysia: North Borneo. Ecol. In forest, 900-1800 m. Note. The type-collection of C. holttumii has much more abundant hairs on all parts of the frond than the other specimens referred to this species. The type of C. subbipinnata was collected at 1800 m, and is small, with largest pinnae only 10 cm long, having only one or two pairs of small free pinnules; the specimen is old and has lost most of its scales and hairs. It seems doubtful whether C. elliptica should be maintained as distinct from 174. C. trichophora COPEL. 160. Cyathea obliqua CopeL. in Elmer, Leafl. Philip. Bot. 4 (1911) 1150.—Al/sophila obliqua C. Cur. Ind. Fil. Suppl. 1 (1913) 5; v. A. v. R. Handb. Suppl. (1917) 54.—Gymnosphaera obliqua CopreL. Gen. Fil. (1947) 98; Fern Fl. Philip. 2 (1960) 236. Stipe unknown. Pinnae to 30 cm long. Pinnules to 72 by 10 mm, the lowest on stalks to 5 mm long, articulate to rachis, base very unequally cuneate (narrow on basiscopic side), apex caudate-acu- minate, edges broadly crenate, lamina rather thin; veins in pinnate groups each with up to 4 pairs, lowest basiscopic vein of each group springing from the costa. Sori 1-3 on each vein-group; no indusia; paraphyses as long as sporangia. Scales and hairs: pinna-rachis glabrous on lower surface; costal scales rather sparse, flat and ovate-acute to bullate and acuminate, light brown, most with a short fringe of pale hairs, some bearing dark setae. Type specimen: ELMER 12354, Sibuyan L., Philippines (MICH; dupl. at K, A, BO, US, FI, S-PAQ SYD se): Distr. Malaysia: Philippines once collected), at 600 m. (Sibuyan LI., 161. Cyathea integra J. Sm. ex Hook. Ic. PI. (1844) t. 638, incl. also var. petiolata HooK. lI.c. t. 638, fig. 2; Sp. Fil. 1 (1844) 26; Syn. Fil. (1865) 23; v. A. v. R. Handb. (1908) 20 (not Suppl. 25, which is C. urdanetensis COPEL.); COPEL. Philip. J. Sc. 4 (1909) Bot. 35; Fern Fl. Philip. 2 (1960) 200.—Trichopteris falcata LLANOS, Fragm. PI. Filip. (1851) 111 (fide MERRILL, Sp. Blanc. 1918, 41).—C. hypocrateriformis v. A. v. R. Bull. Jard. Bot. Btzg II, n. 7 (1912) 9; Handb. Suppl. (1917) 39; CopEL. in Elmer, Leafl. Philip. Bot. 5 (1913) 1680.—C. bulusanensis CopeEL. in Elmer, Leafl. Philip. Bot. 9 (1920) 3109; Fern FI. Philip. 2 (1960) 201.—C. arguta CopeL. Philip. J. Sc. 38 (1929) 133; Fern Fl. Philip. 2 (1960) 203.— C. breviloba Cope. Philip. J. Sc. 81 (1952) 13; Fern FI. Philip. 2 (1960) 201.—Fig. 29a, b. Stipe 20-40 cm, base with spines 2 mm long; scales at base of stipe thin, pale, edges closely setiferous, to 25 by 114 mm; also above base a more or less persistent cover of very small pale fringed or setiferous scales. Main rachis medium brown when dry, in basal part bearing scattered short spines, otherwise smooth and glabrescent. Pinnae: lowest c. 15 cm long, largest 60 cm. Pinnules on larger pinnae 80-120 by 15-25 mm, sessile or lowest stalked to 2 mm (rarely to 4 mm), apex acuminate (not caudate), in lowest pinnules on lower pinnae the basal segment sometimes just free, otherwise whole pinnule lobed to 2-3 mm from costa; costules 6 mm apart; veins 6—8 pairs; lamina-segments firm, edges almost entire or distinctly crenate towards apices, apex bluntly to acutely pointed at end of falcate costule. Sori medial; indusium at first complete, thin and trans- lucent but firm, breaking and persisting at ma- turity; paraphyses abundant, as long as sporangia, pale. Scales and hairs: pinna-rachis beneath gla- brescent, smooth or sparsely warty, residual scales small, pale, fringed or some with setae; on costae near base narrow pale scales with dark marginal setae, smaller ones with irregular fringe of pale hairs; on costules scales like smaller ones on costae, not bullate; stout spreading hairs rarely present on lower surface of costae and costules near apex of pinnule, not on upper surface of costules. Type specimen: CumING 120, Luzon (K; dupl. at US). Dec. 1963] CYATHEACEAE (Holttum) Fig. 29. Cyathea integra J. SM. a. Part of pinnule, upper surface, x 2, b. lower surface showing scales and sori, X 6.—C. squamulata (BL.) CopeL. c. Part of sterile pinnule, lower surface, = 2, d. part of fertile pinnule showing sori and scales, < 6 (a—b SINCLAIR 9539, c SINCLAIR 10336, d KIAH 32179). Distr. Malaysia: Philippines (Luzon, Mindoro, Panay, Samar, Catanduanes, Basilan, Biliran, Mindanao). Ecol. In forest at low and medium altitudes; range reported 500-1200 m, but few specimens bear relevant information. Notes. With the original description of this species, HooKER cited first a specimen from Ambon, then CuminG 120 from Luzon; his figure was certainly prepared from the Luzon specimen, which I therefore regard as the type. The Ambon specimen is quite distinct in shape of lamina- segments and details of scales; it belongs to C. tripinnatifida Roxs. There is considerable variation in size, shape and marginal teeth of the segments of the pinnules in C. integra; there is also variation in the presence of hairs on the lower surface of costae and in length of stalks of lower pinnules. I have not ob- served clear correlation of such characters, and do not think that var. petiolata Hook. can be regarded as a distinct variety. 162. Cyathea stipitipinnula HoLtrum, Kew Bull. 16 (1962) 62. Stipe to more than 30 cm, medium dull brown, rather persistently scaly, warty where scales have fallen; larger scales to 25 by 3 mm, shining brown with paler edges bearing many dark setae, also minute pale irregularly fringed scales; main rachis smooth, glabrescent. Pinnae to 45 cm long. Pinnules almost at right angles to rachis, to 65 by 12 mm, lobed less than half way to costa, lobes rounded and entire, texture coriaceous, the lowest pinnules with cordate bases and on stalks to 4 mm long; costules 314-4 mm apart; veins 3-4 pairs, thick. Sori usually 3 to each lobe, medial; indusia pale, firm, complete, breaking irregularly and persistent; paraphyses dark. Scales and hairs: scales near base of costae ovate-acute, flat, light brown, with numerous crisped marginal hairs or the larger with some setae, grading to light brown bullate scales (often fringed near apices) on costules and on veins. Type specimen: CLEMENS 33156, Mt Kinabalu, N. Borneo (K; dupl. at BO, A, UC, L). Distr. Malaysia: N. Borneo (Mt Kinabalu, several collections). Ecol. In open places in forest, 1200-1500 m. 163. Cyathea zamboangana CopeL. Philip. do. SE 30 (1926) 325; Fern FI. Philip. 2 (1960) 201.— C. urdanetensis CopeL. Philip. J. Sc. 38 (1929) 132; Fern Fl. Philip. 2 (1960) 203.—C. integra (non 148 FLORA MALESIANA [ser. II, vole J. Sm. ex Hook.) v. A. v. R. Handb. Suppl. of C. megalosora CopeEL., and is intermediate in (ET), 23 soral characters between C. megalosora and the Trunk: \eaf-scars elliptic, 244 cm wide, in al- ternate whorls of 3. Stipe 30 cm, base spiny (spines slender, dark, abundant, 1-3 mm) and densely covered with scales; scales to c. 10 by 1 mm, mostly smaller, edges with close dark setae. Main rachis spiny, pale, glabrescent. Pinnae: lowest somewhat reduced, largest to 60 cm or more long. Pinnules on larger pinnae 100-130 by 25-35 mm, lower ones on stalks 3-4 mm long, one lowest segment sometimes just free, rest of pinnule lobed to about 214 mm from costa; costules 6 mm apart; veins to 8 pairs, mostly forked, a few with acroscopic braoch again forked; lamina-segments thin but firm, apices falcate, acute, edge some- times serrate near apex. Sori almost medial; indusium complete, globose, breaking and per- sistent at maturity. Scales and hairs: pinna-rachis pale, smooth, with some residual narrow pale setiferous scales; on costae scattered very small pale short-fringed scales and near base a few narrow pale setiferous scales, sometimes a few stout pale hairs near apex of pinnule; on costules pale bullate scales, sometimes with setae near apex, also sometimes stout pale hatrs near apices of segments; upper surface of costules lacking hairs. Type specimen: COPELAND 1646, near San Ramon, Mindanao (MICH). Distr. Malaysia: Philippines (Mindanao). Ecol. In forest, 500-800 m. 164. Cyathea discophora HOoLTTUM, Kew Bull. 16 (1962) 54. Stipe finely warty throughout, persistently scaly near base; scales pale, to 25 by 2 mm, edges closely set with short dark setae; rachis light brown, glabrescent, sparsely and finely warty. Pinnae to 50 cm long, pinnules rather widely spaced. Pinnules sessile, to 8 by 114 cm, lobed 24, towards the costa except at the very base, lowest segment not free; costules 4-414 mm apart; lamina-segments rather thin, edges cre- nulate, apex broad, sinuses narrow; veins 6—7 pairs. Sori medial; indusium at length a thin brown disc of irregular shape, sometimes excentric, completely covered by mature sorus; paraphyses dark, a little longer than sporangia. Scales and hairs: pinna-rachis glabrescent beneath; costae rather densely scaly, scales at base pale, flat, elongate with short rather stiff pale marginal hairs, grading through similar scales bullate at base to rather large pale bullate scales with or without marginal hairs near their acuminate apices; a few long pale hairs on costa near apex of pinnule; costules bearing large pale bullate acu- minate scales and few hairs; upper surface of costules glabrous. Type specimen: CLEMENS 31698, Mt Kinabalu, N. Borneo (B; dupl. at US, BO). Distr. Malaysia: N. Borneo (Mt Kinabalu, once collected). Ecol. In open place in forest, 2400 m. Note. This occurs within the altitudinal range exindusiate C. squamulata (BL.) COPEL. which occurs at lower altitudes. 165. Cyathea megalosora CopeL. Philip. J. Sc. 12 (1917) Bot. 54; C. Cur. Gard. Bull. S.S. 7 (1934) 221. Trunk to at least 2 m. Stipe c. 30 cm, pale to medium brown when dry, densely scaly, finely warty where scales have fallen; scales thin, pale, somewhat crisped, to c. 25 by 1144 mm, edges bearing rather sparse dark setae towards apex. Pinnae: lowest 20 cm long, largest 35 cm. Pinnules to 60 by 12 mm, almost sessile; lowest 1-2 seg- ments free, rest of pinnule lobed nearly to costa; costules 4-5 mm apart; veins 5—7 pairs; lamina- segments very firm, edges crenate, apices rounded, sinuses narrow. Sori medial; indusium firm, brown -translucent, quite covering sorus to maturity, breaking irregularly and persistent; paraphyses as long as sporangia. Scales and hairs: pinna- rachis persistently densely scaly on lower surface, scales long, pale, entire or nearly so, bases of smaller ones bullate; costae densely scaly on lower surface, scales elongate, pale, almost entire, not setiferous, on distal part of costae many long spreading hairs; costules bearing similar hairs, and scattered hairs on veins also; upper surface of costules and veins bearing long pale hairs. Type specimen: ToppinGc 1759, Mt Kinabalu, N. Borneo (US; dupl. at A, K, SING, S-PA). Distr. Malaysia: N. Borneo (Mt Kinabalu). Ecol. In mossy forest on ridges, 2200-2900 m. At the highest altitudes the lamina of fronds may be only 40 cm long, pinnae to 1014 by 214 cm, with about 5 pairs of free pinnules which are lobed half-way to the costa. 166. Cyathea suluensis BAK. J. Bot. 17 (1879) 65; v. A. v. R. Handb. (1908) 18; Cope. Philip. J. Sc. 4 (1909) Bot. 35; Fern Fl. Philip. 2 (1960) 202. —C. sessilipinnula Cope. Philip. J. Sc. 38 (1929) 134; Fern Fl. Philip. 2 (1960) 202. Stipe to more than 30 cm, minutely spiny to the base; no scales seen. Pinnae: lowest about 10 cm long, largest 30 cm. Pinnules on larger pinnae 50-70 by 13-18 mm, sessile, short-acuminate, lobed 14-34 towards the costa, no free basal segments; costules 4-5 mm apart; veins 4~7 pairs, strongly oblique; lamina-segments thin, slightly crenate near rounded apices, sinuses narrow. Sori medial; indusium complete, thin, pale, breaking irregularly and persistent. Scales and hairs: pinna-rachis beneath glabrescent, residual scales small, pale, short-fringed; on costae near base narrow flat pale dark-setiferous scales, throughout brown bullate scales, often with setae near apex, distally some stout pale hairs; on costules bullate scales, pale brown, and a few hairs; no hairs on upper surface of costules. Type specimen: BuRBIDGE s.n., 1877-78, Sulu Is (K). Distr. Malaysia: Philippines (Mindanao, Ba- silan, Sulu Is), Moluccas (Ternate ?). Dec. 1963] CYATHEACEAE (Holttum) uae Ecol. At c. 600 m. Note. Specimens from Ternate placed tenta- tively in this species are young and sterile; they agree in form of pinnules, venation and scales, but differ in having rather numerous stout hairs on the upper surface of costules. 167. Cyathea robinsonii Copet. Philip. J. Sc. 6 (1911) Bot. 145; v. A. v. R. Handb. Suppl. (1917) 30; CopeL. Fern Fl. Philip. 2 (1960) 203.—C. pseudoalbizzia Cope. Philip. J. Sc. 38 (1929) 135; Fern FI. Philip. 2 (1960) 203. Stipe to at least 40 cm, warty where scales have fallen; scales pale to brownish, shining, to 20 mm long, edges setiferous; main rachis finely warty, bearing some hairs throughout, densely hairy towards apex of frond, hairs mixed with small pale scales. Pinnae to 40 cm long. Pinnules to 100 by 20 mm, the lowest with stalks 5-10 mm long; at base of larger pinnules 1-2 pairs of free tertiary leaflets, then several pairs of segments separately adnate by narrow bases to costa, rest of pinnule lobed nearly to costa; costules on larger pinnules 5-7 mm apart; lamina-segments firm, dark on upper surface when dry, edges almost entire; veins 5—6 pairs. Sori medial; indusium at first complete, rather firm, breaking irregularly and persistent. Scales and hairs: pinna-rachis and costae densely hairy and scaly on lower surface, hairs spreading, pale, 2 mm long, scales narrow, pale, sometimes setiferous; on lower surface of costules and veins abundant pale spreading hairs, no scales. Type specimen: ROBINSON BS 9394, Mt Binuang, Luzon (MICH; dupl. at K, US, P, UC). Distr. Malaysia: Philippines (Luzon). Ecol. At 875-1150 m. Note. The type of C. pseudoalbizzia is small, with pinnules to 70 by 12 mm. 168. Cyathea senex v. A. v. R. Bull. Jard. Bot. Btzg II, n. 16 (1914) 4; Handb. Suppl. (1917) 34. Stipe densely scaly throughout, scales to 30 mm long, mostly not over 1 mm wide, pale, edges closely set with dark setae; main rachis similarly scaly near base only. Pinnae to 40 cm long; lowest pinnae somewhat reduced. Pinnules to 80 by 12 mm, lowest 1 or 2 segments almost free, rest of pinnule lobed to within 1 mm of costa; costules 314 mm apart; lamina-segments firm, dark above when dry, edges crenate, sinuses narrow; veins to 6 or 7 pairs. Sori medial, indusium at first thin and complete, breaking and persistent; paraphyses pale. Scales and hairs: pinna-rachis bearing pale spreading hairs and many small pale setiferous scales; scales at base of costae narrow, pale, seti- ferous, grading to acuminate bullate scales at apex, spreading hairs present throughout; a few bullate scales present on costules, with many hairs; erect pale hairs also present on lower surface of veins; thick curved hairs scattered on upper surface of costules and veins. Type specimen: MATTHEW 526-A, Mt Singga- lang, Sumatra (BO; dupl. at K). Distr. Malaysia: Central Sumatra. Ecol. At 1500-1800 m. Note. MATTHEW gave the number 526 also to specimens of C. sumatrana BAK., which are at Kew, from the same locality. 169. Cyathea sibuyanensis Cope. in Elmer, Leafl. Philip. Bot. 4 (1911) 1150; v. A. v. R. Handb. Suppl. (1917) 38; Copev. Fern Fl. Philip. 2 (1960) 204. Stipe not known. Main rachis closely warty on lower surface, bearing many very small dull scales. Pinnae to 35 cm long. Pinnules to 90 by mostly to 20 mm (on a lower pinna to 25 mm), lowest 1-2 pairs of segments quite free (lowest sometimes stalked) and deeply lobed at the base, then several segments contracted at the base and separately adnate to the costa, rest of pinnule lobed almost to the costa; costules 5-6 mm apart, decidedly oblique; lamina-segments firm, edges almost entire to crenate; veins 6—7 pairs, the lowest in lobes of basal segments pinnately branched. Sori medial; indusium pale and translucent, at first completely covering sorus, breaking and persistent. Scales and hairs: pinna-rachis closely warty and covered with very small dull scales, the largest setiferous, hairs also mixed with the scales, at least towards apex of pinna; costae scaly near base, hairy in apical half, scales mostly ovate-acute, hardly 1 mm long, brown, edges with a few setae or short hairs near tip, none bullate; costules bearing hairs on lower surface, but no hairs on veins; costules glabrous on upper surface. Type specimen: Evmer 12513, Mt Géiting- Giting, Sibuyan I. (MICH; dupl. at K, A, FI, US, BOR Siw sla): Distr. Malaysia: Philippines (Sibuyan, one collection). Ecol. Altitude 1450 m. 170. Cyathea philippinensis BAk. Ann. Bot. 5 (1891) 186; v. A. v. R. Handb. (1908) 16, 783; CopeL. Philip. J. Sc. 4 (1909) Bot. 111, incl. var. nuda CopEL.; Vv. A. v. R. Handb. Suppl. (1917) 23; Cope-. Fern FI. Philip. 2 (1960) 205.—C. bicolana CopeL. in Elmer, Leafl. Philip. Bot. 9 (Mar. 1920) 3108; Fern Fl. Philip. 2 (1960) 204.—C. ramosiana v. A. v. R. Bull. Jard. Bot. Btzg III, 2 (June 1920) 137; CopeL. Fern Fl. Philip. 2 (1960) 206.—C. heteroloba Copret. Philip. J. Sc. 38 (1929) 134; Fern Fl. Philip. 2 (1960) 204.—C. bontocensis Cope. Philip. J. Sc. 46 (1931) 209; Fern FI. Philip. 2 (1960) 205. Stipe 12-35 cm, base dark and warty; scales pale to brownish, to 25 by 2 mm, edges bearing concolorous setae. Lowest pinnae 4-10 cm long, largest to 30 cm. Pinnules to 65 by 15 mm, lowest 2-3 segments of larger pinnules free or nearly so, rest of pinnule lobed 14-34 towards costa; cos- tules 314-514 mm apart; lamina-segments entire, apex broadly rounded; veins 4-6 pairs. Sori nearer to costule than edge; indusium complete, thin, translucent, breaking and persistent. Scales and hairs: on lower surface of pinna-rachis narrow 150 FLORA MALESIANA [ser.- Tl) vel pale setiferous scales; on costae at base elongate flat scales with pale fringe or dark setae, grading to pale bullate scales, pale thick hairs also usually present near apex of costa; on costules bullate scales, some fringed near apices, and usually also a few hairs; no hairs on upper surface of costules and veins. Type specimen: ex Hort. Veitch, cult. Kew, origin Philippines, Feb. 1878 (K). Distr. Malaysia: Philippines (Luzon, Min- doro). Ecol. In mountain forest (only altitude records are 1400-1500 m). Note. The type specimen is a frond of a small cultivated plant, bearing pinnules only 20 by 6 mm, but in general shape and in scales and sori it re- sembles the type specimens of the other species above cited, from which the present description is prepared. 171. Cyathea assimilis Hoox. Syn. Fil. (1865) 24; v. A. v. R. Handb. (1908) 20; CopeL. Philip. J. Sc. 4 (1909) Bot. 49.—C. beccariana CESATI, Atti Ac. Napoli 7® (1876) 3.—C. dulitensis BAK. Kew Bull. (1896) 40; v. A. v. R. Handb. (1908) 16; CopeE-. Philip. J. Sc. 4 (1909) 33.—C. ampla CopPeEL. Philip. J. Sc. 6 (1911) Bot. 361; v. A. v. R. Handb. Suppl. (1917) 26; non HoLttrum, Rev. FI. Mal. 2 (1954) 135.—C. stipitulata CopeEL. Philip. J. Sc. 6 (1911) Bot. 362; v. A. v. R. Handb. Suppl. (1917) 29. Stipe to 65 cm, medium to dark brown, finely warty, persistently scaly near base; scales medium brown, shining, firm, 15-20 by 1-2 mm, edges closely setiferous. Lamina to almost 200 cm long; largest pinnae to 55 cm long, more or less distinctly articulate to rachis, the lower pinnae with stalks to 4cm long. Largest pinnules 80-90 by 25 mm, the lowest with stalks 4-8 mm, basal 1-2 lamina- segments quite free, next 1-2 pairs scmetimes free with adnate base, rest of pinnule lobed to 1-2 mm from costa, apex evenly attenuate; in somewhat smaller pinnules only the basal basiscopic seg- ment free, or no free segments; costules S-7 mm apart; lamina-segments firm, edges more or less crenate, apices rounded; veins 8—10 pairs. Sori medial; indusium at first quite covering sorus, pale and thin, breaking irregularly and persistent. Scales and hairs: pinna-rachis beneath glabrous or with a few small setiferous scales; costae be- neath near base bearing narrow strongly setiferous scales grading to small bullate setiferous scales distally and on costules; no hairs on lower surfaces of pinna-rachis, costae and costules. Type specimen: Loss s.n., 1857, hills, Sarawak K). Distr. Malaysia: S. Sumatra (Mt Dempo), Borneo (Sarawak). Ecol. Forests, 300—2000 m, the more coriaceous specimens in ridge-forest on sandstone. Notes. The type collection of C. dulitensis was a small plant with largest pinnae only just pinnate at base; the later collection of RICHARDS from the same locality has amply bipinnate fronds, agreeing in other characters, and probably re- presents the full development of the species. All specimens from Mt Dulit are much darker in all parts than those from elsewhere in Sarawak (e.g., the type of C. assimilis) and have a thicker lamina, but do not differ in other characters. The specimen from Mt Dempo in Sumatra is dark like those from Dulit. The type collection of C. stipitulata is an unusu- ally small specimen, with pinnules 70 by 17 mm, costules 4144 mm apart and veins 6—7 pairs; it was found on Mt Matang, near Kuching, at 300 m. In HoLtrum, Rev. FI. Mal. 2 (1954) 135, the name C. ampla is wrongly given to a specimen of C. polypoda which has unusually wide pinnules; this latter species is quite exindusiate. C. assimilis is closely related to C. philippinensis Bak. and appears to differ chiefly in the consistent- ly larger size of all parts of the frond. 172. Cyathea trichodesma (ScortT.) CopeL. Philip. J. Sc. 4 (1909) Bot. 55.—Alsophila trichodesma Scort. in Bedd. J. Bot. 25 (1887) 321; BeEpp. Handb. Suppl. (1892) 3; v. A. v. R. Handb. (1908) 35.—Alsophila margarethae SCHROET. ex Curist, Ann. Jard. Bot. Btzg 20 (1905) 136; v. A. v. R. Handb. (1908) 33.—C. margarethae Cope-. Philip. J. Sc. 4 (1909) Bot. 38; C. Cur. Gard. Bull. S.S. 7 (1934) 220.—C. burbidgei [non (BAK.) CopeL.] Ho_ttum, Rev. Fl. Mal.2 (1954) 124. Trunk slender, to 414 m. Stipe fairly long, near base densely scaly (not hairy), finely warty when scales have fallen; scales medium to light brown, shining, firm, rather dark brown when dried, to about 25 by 2 mm, edges closely set with short dark setae. Frond including stipe 2-314 m long. Pinnae: lowest slightly reduced, largest about 60 cm long. Pinnules commonly to 90 by 15 mm, largest seen 110 by 20 mm, nearly sessile, shortly acuminate, lobed to about 2 mm from costa, no free basal segments; costules 444-5 mm apart; veins 6-8 pairs; lamina-segments thin, crenate, sinuses narrow. Sori medial, often confluent at maturity; no indusium; paraphyses a little longer than sporangia. Scales and hairs: pinna-rachis, costae, costules and veins on lower surface bearing many pale spreading hairs 1-2 mm long; scales on costae and costules sparse, pale, some narrow and flat, some bullate, most bearing dark setae; hairs present on upper surface of costules and veins. Type specimen: SCORTECHINI s.n., Perak (BM). Distr. Malaysia: Malay Peninsula (central part), Borneo (Sarawak, N. Borneo). Ecol. Lowland forest, sometimes by rivers, to 1500 min N. Borneo. Notes. Bornean specimens are all smaller than those from the Malay Peninsula, the largest having pinnules to 70 by 15 mm; they also lack hairs on the upper surface of costules. In size, the Bornean specimens are nearer to C. trichophora, but the distribution of hairs is different. In the Malay Peninsula has been found a speci- men rather intermediate between C. trichodesma and C. alternans, both of which species were growing near it. Dec. 1963] CYATHEACEAE (Holttum) 151 173. Cyathea wallacei (MeTT. in KUHN) COPEL. Philip. J. Sc. 4 (1909) Bot. 48.—Alsophila wallacei Metrt. in Kuhn, Linnaea 36 (1869) 153; v. A. v. R. Handb. (1908) 36.—Alsophila burbidgei Bax. J. Bot. 17 (1879) 38; v. A. v. R. Handb. (1908) 33.— C. burbidgei CopeL. Philip. J. Sc. 4 (1909) Bot. 55; non HOLTTuM, Rev. Fl. Mal. 2 (1954) 124, which is C. trichodesma.—Gymnosphaera burbidgei Co- PEL. Gen. Fil. (1947) 99. Stipe 30 cm or more, pale and smooth above the base; scales at base light brown, firm, to 15 by 2 mm, setiferous; main rachis finely hairy on abaxial surface in apical part. Largest pinna seen 38 cm long. Pinnules to 65 by 13 mm, sessile, apex abruptly narrowed, lobed to within 1 mm of costa, lowest segment almost free; costules 3-314 mm apart; lamina-segments thin, slightly crenate, si- nuses narrow; veins 4-6 pairs, mostly simple. Sori medial, lacking indusia. Scales and hairs: lower surface of pinna-rachis, costae, costules and veins bearing pale spreading hairs 1 mm long; pale bullate scales present on costae and costules; upper surface of costules and veins bearing scattered long spreading hairs. Type specimen: WALLACE s.n., 1857, Borneo (original lost ?; dupl. at Kew). Distr. Malaysia: Borneo (Sarawak; N. Borneo). Ecol. In lowland forest, at least sometimes on poor sandstone soil. Notes. The pinnules of this species are more deeply cut than those of other species of sect. Schizocaena, and have closer costules; they have the aspect of sect. Sphaeropteris, but are much smaller than normal in that section, and have hairiness like that of some members of sect. Schizocaena. Alsophila wallacei was credited to METTENIUS by CHRISTENSEN, Ind. Fil., but it was described without author’s name in the original paper. 174. Cyathea trichophora CopeL. Philip. J. Sc. 6 (1911) Bot. 363.—C. poiensis CopEL. I.c. 362.— Alsophila poiensis v. A. v. R. Handb. Suppl. (1917) 56.—Alsophila trichophora v. A. v. R. L.c. 72.— C. mollis Corey. Philip. J. Sc. 12 (1917) Bot. 52; C. Cur. Gard. Bull. S.S. (1934) 220.—C. ramosii CopeL. Philip. J. Sc. 30 (1926) 325.—Alsophila ramosii C. Cur. Ind. Fil. Suppl. 3 (1934) 23.— C. bipinnatifida CoPeEL. Philip. J. Sc. 56 (1935) 97, pl. 2 (not C. bipinnatifida (BAK.) Domtn, 1929).— Gymnosphaera bipinnatifida CopEL. Gen. Fil. (1947) 99.—Gymnosphaera mollis CoPEL. l.c. 99.—Gymno- sphaera trichophora Cope. I.c. 99; Fern Fl. Philip. 2 (1960) 236. Trunk to 50 cm. Stipe 25-50 cm, at least the basal part persistently scaly, scales to 20 by 3 mm, light brown, shining, edges setiferous; main rachis bearing more or less abundant narrow pale seti- fercus scales and also spreading hairs 2 mm long. Pinnae: lowest reduced and deflexed, largest 25-30 cm long. Pinnules 30-55 by 10-14 mm, lobed half- way to costa; costules 314-4 mm apart; veins 3-5 pairs; lamina-segments thin, entire, ends rounded. Sori medial; no indusia; paraphyses not longer than sporangia. Hairs rather abundant on lower surface of pinna-rachis and costae, sometimes on costules; some pale bullate scales present on cos- tules. Type specimen: Ramos 949, Prov. Laguna, Luzon (MICH; dupl. at FI, UC). Distr. Malaysia: Philippines (throughout), Borneo (Sarawak and N. Borneo). Ecol. Apparently in low country forest, highest record 1200 m. Notes. The type of C. bipinnatifida (from Basilan I.) is from a young plant, with simply pinnate frond bearing few sori; the upper pinnae are closely similar to pinnules of other specimens referable to C. trichophora. It may be that C. tri- chophora and C. elliptica should be united; the latter has a different distribution of hairs and has apparently only been found at higher altitudes. 175. Cyathea polypoda BAK. Trans. Linn. Soc. II, Bot. 4 (1894) 250; v. A. v. R. Handb. (1908) 18; C. Cur. Gard. Bull. S.S. 7 (1934) 219; HoLttrum Rev. Fl. Mal. 2 (1954) 122.—C. kemberangana CopeEL. Philip. J. Sc. 12 (1917) Bot. 52; C. Cur. Gard. Bull. S.S. 7 (1934) 219.—Alsophila kem- berangana C. Cnr. Ind. Fil. Suppl. 3 (1934) 22.— Gymnosphaera dinagatensis CopeL. Philip. J. Sc. 81 (1952) 19, pl. 14; Fern FI. Philip. 2 (1960) 235. —C. ampla (non CopeEL.) HoLttum, Rev. Fl. Mal. 2 (1954) 125.—Gymnosphaera glabra (non BL.) CopeL. Fern FI. Philip. 2 (1960) 235. Trunk to 3 m, covered with persistent leaf-bases; small branches often borne on lower part of trunk. Stipe to 80 cm, pale (green when living) to rather dark, densely scaly near base, finely warty after scales have fallen; scales shining medium brown, firm, to 30 by 2 mm, edges bearing close con- colorous setae; rachis and pinna-rachis glabrescent on lower surface. Pinnae to 60 cm long, lower ones long-stalked, not greatly reduced. Pinnules com- monly 85 by 20 mm, sometimes to 110 by 27 mm, all stalked, stalks of lowest to 9 mm; basal 1-2 segments of lowest pinnules of largest pinnae quite free, then sometimes a pair with lamina adnate at base but free, rest of pinnule (whole of smaller pinnules) lobed to 1-2 mm from costa; costules 414-514 (sometimes to 644) mm apart; veins 7-9 pairs, forked, acroscopic branch some- times forked again; lamina-segments rather thick and rigid when dry, edges crenate, apices rounded, sinuses narrow except near base of largest pin- nules. Sori nearer to costule than to edge; no in- dusium; paraphyses a little longer than sporangia. Scales and hairs: small dark to medium brown setiferous scales near base of costae; bullate scales, often setiferous, on costules, all scales often early caducous. Type specimen: HAVILAND 1479, Mt Kinabalu, N. Borneo (K). Distr. Malaysia: Malay Peninsula, Borneo (Sarawak, N. Borneo), Philippines (Panay, Minda- nao). soe In open places on ridge-crests and sum- mits, 600-2200 m; specimens from the higher elevations are decidedly coriaceous. Note. BAKER described this species as indusiate; 152 he had a young frond on which bullate scales in some cases partly cover the immature sori. 176. Cyathea obscura (ScortT.) CopeL. Philip. J. Sc. 4 (1909) Bot. 37; Ho_trum, Rev. Fl. Mal. 2 (1954) 124.—Alsophila obscura Scort. in BEpp. J. Bot. 25 (1887) 321, t. 278, fig. 2; Handb. Suppl. (1892) 3; v. A. v. R. Handb. (1908) 34; Suppl. (1917) 57.—Alsophila subobscura v. A. v. R. Bull. Jard. Bot. Btzg II, n. 20 (1915) 1, t. 1; Handb. Suppl. (1917) 57.—C. bartlettii CopeL. Un. Cal. Publ. Bot. 14 (1929) 371.—C. pulchra CopEL. I.c. 372.—C. subobscura Domin, Pterid. (1929) 263.—Alsophila bartlettii C. Cur. Ind. Fil. Suppl. 3 (1934) 20.—Alsophila pulchra C. Cure. I.c. 23.— Gymnosphaera pulchra CopPeEL. Gen. Fil. (1947) 99. Stipe dark to medium brown, densely scaly towards the base, finely warty where scales have fallen; scales 20-40 by 2-314 mm, shining, pale brown, edges closely setiferous. Pinnae: lowest somewhat reduced (sometimes only 10 cm long), largest 50 cm long. Pinnules: largest commonly 60-70 by 12-13 mm, largest seen 80 by 15 mm, on stalks 1-2 mm long, apex shortly acuminate, edges lobed 14-24 distance to costa; costules 3144-4 mm apart; veins about 6 pairs; lamina- segments firm, edges almost entire, apices bluntly pointed and asymmetric. Sori medial, on about 3 pairs of basal veins only, becoming quite con- fluent at maturity; no indusium; paraphyses co- pious, pale, much longer than sporangia. Scales and hairs: main rachis and pinna-rachis minutely warty and glabrescent beneath or bearing small setiferous scales; costae near base bearing very narrow dark-setiferous scales; costules bearing pale bullate scales often with dark setae near their apices; no hairs on lower surface; no hairs on upper surface of costules and veins. Type specimen: SCORTECHINI S.7., Perak (BM; dupl. at K, SING). Distr. Malaysia: Sumatraand Malay Peninsula. Ecol. In forest, 900-1400 m. 177. Cyathea agatheti Ho_trum, Kew Bull. 16 (1962) 51. Trunk hardly 5 cm high. Stipe 35-75 cm, dark towards base which is covered with scales, slightly warty where scales have fallen, distal part and rachis pale (green when living), smooth and gla- brous; scales on base of stipe to 10 by 2-214 mm, light brown with somewhat paler edges which bear many rather long dark setae. Lamina 50-60 cm long; pinnae distinctly articulate to rachis, lowest somewhat reduced, largest 18-25 cm long. Pin- nules to 35 by 10 mm, abruptly narrowed at apex, lobed to within 1 mm of costa, lowest on stalks 1 mm long; costules 3 mm apart; lamina-segments thin, almost entire, sinuses narrow; veins 4—5 pairs, simple. Sori medial on veins; no indusia; para- physes dark, shorter than sporangia. Scales and hairs: lower surface of costae bearing scattered spreading pale hairs, and a very few hairs on costules; scales very few, only seen on young frond, narrow, pale, ciliate; upper surface of costae, costules and veins bearing scattered long spreading FLORA MALESIANA [ser. ly vole hairs in addition to the usual antrorse hairs on costae. Type specimen: KosTERMANS 12870, W. Kutai, E. Borneo (BO; dupl. at K, L). Distr. Malaysia: E. Borneo (one collection). Ecol. In Agathis-forest on water-logged white acid sand, 600 m. 178. Cyathea squamulata (BL.) Cope. Philip. J. Sc. 4 (1909) Bot. 37; HoLttrum, Rev. Fl. Mal. 2 (1954) 122, fig. 49.—Gymnosphaera squamulata BL. En. Pl. Jav. (1828) 243; CopreL. Gen. Fil. (1947) 99; Fern FI. Philip. 2 (1960) 235.—AI- sophila squamulata Hook. Sp. Fil. 1 (1844) 51, p.p.; Metr. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 52; Racis. Fl. Btzg 1 (1898) 33; v. A. v. R. Handb. (1908) 33; Suppl. (1917) 56.—Al/sophila comosa WALL. ex Hook. Sp. Fil. 1 (1844) 53; Syn. Fil. (1865) 41; Bepp. Ferns Br. Ind. (1865) pl. 84; Handb. (1883) 13.—Alsophila laeta KUNZE, Bot. Zeit. 4 (1846) 476, p.p.—Alsophila oligosora Mia. Verh. Kon. Ned. Inst. Wet. 3, pt 4 (1851) 43. —Alsophila ridleyi BAk. Ann. Bot. 8 (1894) 122; v. A. v. R. Handb. (1908) 32.—C. ridleyi COPEL. Philip. J. Sc. 4 (1909) Bot. 36.—C. brooksii CopeL. ibid. 6 (1911) Bot. 135, pl. 16, not C. brooksii MAXON, 1904.—C. paraphysata COPEL. I.c. 135, pl. 15.—Alsophila sarawakensis C. CHR. Ind. Fil. Suppl. (1913) 5.—Alsophila xantholepia v. A. v. R. Bull. Jard. Bot. Btzg II, n. 23 (1916) 1; Handb. Suppl. (1917) 489, not Alsophila xantholepis CHRIST, 1899.—Alsophila paraphysata v. A. v. R. Handb. Suppl. (1917) 58.—Alsophila allocota v. A. v. R. Bull. Jard. Bot. Btzg III, 5 (1922) 180.—Alsophila glabrescens v. A. Vv. R. Le. 181.—C. deuterobrooksii Cope. Philip. J. Sc. 38 (1929) 131.—C. allocota Domin, Acta Bot. Bohem. 9 (1930) 89.—C. glabrescens Domin, I.c. 119.— C. xanthina Domin, /.c. 172.—Alsophila xanthina C. Cure. Ind. Fil. Suppl. 3 (1934) 24.—Gymno- sphaera sarawakensis CopeL. Gen. Fil. (1947) 99. —Fig. 8c, 29c-d. Trunk to c. 2 m. Stipe 40-60 cm, densely and persistently scaiy for most of its length (main rachis sometimes also persistently scaly); scales firm, medium brown, largest 30 by 2-3 mm (rarely to 40 by 4 mm), edges closely set with dark setae. Frond to c. 150 cm long. Pinnae: lowest somewhat reduced, variable; largest 50 cm long. Pinnules commonly to 80 by 15 mm, largest seen 100 by 20 mm, lower ones on stalks 1-2 mm long, apex shortly acuminate, edges lobed 14-74 distance towards costa, no free basal segments but pinnules on exceptionally large pinnae some- times lobed nearly to costa at base; costules 314— 414 mm apart; veins 6-9 pairs, of smaller pinnules mostly simple, of larger ones mostly forked; lamina-segments rather thin, edges almost entire, apices rounded and asymmetric. Sori a little nearer to costule than to edge; no indusia; pale paraphyses usually longer than sporangia. Scales and hairs: pinna-rachis glabrescent beneath, finely warty, sometimes with residual small setiferous scales; on costae, near base, usually narrow flat brown strongly setiferous scales, grading to bul- Dec; 1963] CYATHEACEAE (Holttum) 153 late ones distally; on costules pale bullate scales, the larger ones acuminate and setiferous near apices; a few stout hairs on upper surface of costules. Type specimen: KUHL & VAN HASSELT, Pasir Ipis, W. Java (L). Distr. Malaysia: Sumatra, Malay Peninsula, Java, Borneo, S. Philippines (Sulu Arch.). Ecol. A small tree-fern of forest, not in open places, in lowlands and to c. 1500 m (rarely above 1000 m in the Peninsula); specimens from 2500 m have been found on Mt Kinabalu, N. Borneo, and are small, with exceptionally scaly rachis, but do not appear to differ significantly in other ways. Note. The type specimens of Alsophila comosa and A. ridleyi are both from Singapore I. The latter is only distinct in smaller size and less deeply lobed pinnules. Similar differences characterize the other species reduced to synonymy. The type collection of C. sqguamulata includes a small frond only 60 cm long in all, with pinnae to 8 cm long and very few free pinnules, but fertile. 4b. Subsection Sarcopholis Ho.ttum, subsect. nov.—Fig. 30. A subsectione Schizocaena differt: paleis stipitis carnosis ascendentibus, apices versus planis setiferisque. Type species: Cyathea rosenstockii BRAUSE. Distr. Malaysia: Moluccas and New Guinea to Polynesia. Taxon. Fleshy upcurved bases of stipe-scales (fig. 30) are the distinctive character of this subsection; distally these fleshy bases are more or less abruptly flattened to scales which have setiferous margins. The flat distal parts are often eroded from herbarium specimens; a careful morphological and developmental study from good fresh material is needed. Some species have the fleshy bases less developed, and I am not sure that there is a sharp distinction from subsect. Schizocaena. The scale-bases in C. pulcherrima CopEL. (subsect. Sphaeropteris) are comparable developments, but they are slender, rigid, and spread at right angles to the surface of the stipe. KEY TO THE SPECIES 1. Pinna-rachis hairy on the lower surface. 2. Scales on costae setiferous. 3. Few or no bullate scales. Pinnules lobed to 2 mm from costa . 3. Bullate scales present. Pinnules lobed to within 1 mm from costa . 2. Scales on costae not setiferous. 1. Pinna-rachis not hairy on lower surface. 4. Indusium quite lacking. 5. Pinnules almost sessile, to 28 mm wide . 5. Pinnules long-stalked, to 45 mm wide . 4. Indusium present. 6. Pinnules on stalks to 6 mm long. Stipe to 60 cm . ‘ 6. Pinnules sessile or on much shorter stalks. Stipe much shorter. 7. Pinnules to 35 mm wide, larger ones with lowest 1-2 lamina-segments free. No bullate scales. 185. C. inaequalis 7. Pinnules rarely over 25 mm wide, lowest segments not free. Bullate scales usually present on cos- tules. 8. Pinnules lobed only in basal half . 8. Pinnules deeply lobed throughout. 9. Pinnules not over 20 mm wide, lobed to c. 3 mm from costa. . eae 37-7 © werner 9. Pinnules on largest pinnae in most cases over 20 mm wide, lobed to 1-2 mm from costa. 10. Scales on costae and costules bullate, mostly entire. ...... . 188. C. insularum 10. Scales on costae mostly small and fringed or setiferous; bullate scales on costae and costules mostly fringed. 11. Pinnules of largest pinnae 22-28 mm wide. 12. Segments of lamina narrowed rather evenly from base to pointed apex, sinuses thus triangu- lar. Scales on costae not setiferous; bullate scales few. Lowest pinnae not greatly reduced. 189. C. tripinnatifida 12. Segments of lamina with rounded apex. Scales on costae often setiferous, bullate scales usually present. Lowest pinnae small, stipe very short . 190. C. runensis 11. Pinnules of largest pinnae less than 20 mm wide . 191. C. moseleyi 179. C. fusca . 180. C. setifera 181. C. rosenstockii 182. C. marginata 183. C. mesosora 184. C. papuana 186. C. parvipinna 154 FLORA MALESIANA [ser. II, vol. 1? 179. Cyathea fusca BAK. in Beccari, Malesia 3 (1886) 31; v. A. v. R. Handb. (1908) 19; Suppl. (1917) 25; CopeEL. Philip. J. Sc. 77 (1947) 100. Trunk to 2 or 3 m; fronds to 150 cm long. Stipe to 15 cm, bearing thick fleshy scales which have dark setae on their edges (at least when young); rachis bearing similar scales near base, also short hairs on lower surface. Pinnae: lowest c. 8 cm long, largest 40 cm. Pinnules to c. 90 by 20 mm, lobed to 2-3 mm from costa, lowest seg- ment not free; costules 5 mm apart; lamina- segments firm, almost entire, apices rounded; veins c. 6 pairs. Sori medial, indusiate; indusium complete, pale, breaking and largely persistent. Scales and hairs: pinna-rachis densely hairy on lower surface, hairs spreading, c. 14 mm long; some similar hairs on bases of costae; scales on costae very small, bearing long dark setae; a few similar scales on costules, none bullate; no hairs on upper surface of costules and veins. Type specimen: D’ALBERTIS s.n., Fly R., Papua (K). Distr. Malaysia: Eastern New Guinea. Ecol. Casual in undergrowth of forest near river (BRASS). Note. This species is very near C. werneri ROsENST., but appears to differ from it in presence of abundant hairs on lower surface of pinna-rachis and in absence of bullate scales. >] } | { iif H t i pctvatiesele: cone scbntnnsinndeedh Anette Snwke PERRIER eo 4 Fig. 30. Cyathea rosenstockii BRAusE. Base of stipe, showing fleshy scales; scale in cm (PULLE 509, Mt Perameles, 1100 m, BM). 180. Cyathea setifera Hottrum, Kew Bull. 16 (1962) 62. Stipe 10 cm, covered with thick dark pale-edged scales 20 by 1144 mm, their edges (at least near apex) bearing dark setae; also covered, between the large scales, with a thin dark felt of small scales of irregular shape, some with dark setae; scales of very young fronds very thick at their bases which have internal air-spaces. Rachis bearing scattered narrow dark scales 10 mm long and also thick hairs, and the remains of a felt of small scales as on the stipe. Pinnae: lowest about 4cm long, largest 40 cm. Pinnules to 70 by 16 mm, sessile, very shortly acuminate, lobed nearly to the costa; basal 1-2 segments contracted at base; costules 4-414 mm apart, at 60° to the costa; lamina-segments rigid but not very thick, entire or slightly crenulate, ends rounded; veins 6—7 pairs, strongly raised on lower surface, forked rather far from costule. Sori nearer to edge than to costules; indusium thin and pale, breaking and persistent; paraphyses dark, longer than sporangia. Scales and hairs: upper part of main rachis and pinna-rachis bearing thick pale hairs on iower surface, also scattered dark strongly setiferous scales of various sizes, to 5 mm long; costae densely covered beneath with elongate brown setiferous scales, grading to brown bullate scales setiferous near their apices; bullate scales distally on costae and on costules brown, pale-fringed at apices; hairs on upper surface of costules few (not more than | on a costule). Type specimen: MAIN & ADEN 1306, Morotai, N. Moluccas (BO; dupl. at K, L). Distr. Malaysia: Moluccas (Morotai), collection. Ecol. At 1000 m. one 181. Cyathea rosenstockii BRAUSE, Bot. Jahrb. 56 (1920) 49; Cope. Philip. J. Sc. 77 (1947) 101.— Fig. 30. Trunk 114-2 m, bearing 6-8 fronds. Stipe 10-15 cm, covered with thick ascending fleshy scales to 45 by 4 mm wide at base; main rachis almost smooth, glabrescent, pale. Pinnae: lowest 3-5 cm long, increasing upwards, largest 30 cm. Pinnules to 85 by 23 mm, sessile, lowest segment a little constricted at base, rest of pinnule lobed to 1 mm from costa; costules 5-544 mm apart, distinctly oblique; lamina-segments rigid, edges slightly crenate, sinuses wider in fertile than in sterile pinnules; veins 7-10 pairs, much raised on upper surface, less so beneath. Sori nearer costule than edge; indusium thin, at first complete, later breaking and sometimes almost disappearing. Scales and hairs: lower surface of pinna-rachis throughout bearing many pale thick crisped hairs; scales on lower surface of costae and costules dark, bullate, sometimes very few. Type specimen: LEDERMANN 11264, Sepik area, E. New Guinea (B). Distr. Malaysia: New Guinea. Ecol. In mossy forest, or rain forest, at 1300— 1750 m. Dec. 1963] 182. Cyathea marginata (BRAUSE) Domin, Acta Bot. Bohem. 9 (1930) 134.—Al/sophila marginata BraAuse, Bot. Jahrb. 56 (1920) 63. Trunk to 3 m tall; fronds 150-200 cm long. Stipe to at least 35 cm, warty, with one pair of reduced pinnae near base; stipe-scales not seen. Pinnae: lowest 18 cm long, largest 42 cm. Pinnules to 125 by 28 mm, sessile, acuminate, lobed to 2-3 mm from costa, lowest segment not free; costules 6-7 mm apart; lamina-segments rather thick and rigid, crenulate, apices rounded, sinuses narrow; veins 11-12 pairs. Sori medial; no indusium; re- ceptacle large. Scales and hairs: pinna-rachis beneath light brown, smooth, with a few persistent pale thin scales bearing irregular long flexuous setae; costal scales few, thin, ovate, bearing long dark setae; no scales seen on costules. Type specimen: LEDERMANN 12586, Sepik region, E. New Guinea (B). Distr. Malaysia: E. New Guinea (one collec- tion). Ecol. In mountain forest at 1400-1500 m. 183. Cyathea mesosora Ho_truM, Kew Bull. 16 (1962) 57. Trunk slender, to 3 m. Fronds few, 150-220 cm long. Stipe to 10 cm, dull, spines to 1 mm, rather abundant; scales to c. 15 by 114 mm, slightly thickened at base, dark, edges bearing irregular flexuous setae or hairs. Pinnae: lowest 3—5 cm long, simply pinnate (sometimes widely separated from next); largest 42 cm long. Pinnules widely spaced, to 110 by 45 mm, lowest on stalks to 8 mm long, lobed throughout to 3-4 mm from costa; costules 714-9 mm apart; lamina-segments very firm, edges crenate (more strongly towards apex) and thickened, apex rounded; veins 10—12 pairs on largest pinnules, mostly rather narrowly forked, strongly raised on lower surface. Sori medial, usually at the fork of a vein, sometimes one sorus in middle of each branch of lowest vein; no indusium; receptacle large, often very broad; paraphyses short, pale, slender. Scales and hairs: pinna-rachis beneath smooth and glabrous; no scales seen on lower surface of costae and costules; minute hairs (bases of former scales ?) sometimes abundant on lower surface of veins; no hairs on upper surface of costules and veins. Type specimen: CARR 15720, Lala River, Papua (BM; dupl. at K, L, MICH). Distr. Malaysia: Eastern New Guinea (three collections). Ecol. In forest at 1400-1750 m. 184. Cyathea papuana (RIDL.) v. A. v. R. Handb. Suppl. (1917) 487; Copev. Philip. J. Sc. 77 (1947) 121.—Alsophila papuana Riv. Trans. Linn. Soc. Il, Bot. (1916) 252.—Gymnosphaera papuana CopeL. Gen. Fil. (1947) 98. Stipe to 60 cm, reduced basal pinnae lacking; spines and scales as C. mesosora. Pinnae to more than 40 cm long. Pinnules to 90 by 22 mm, lobed to 3 mm from costa, lowest on stalks to 6 mm long; costules 5-7 mm apart; lamina-segments firm, edges thick, crenate; veins to 8 or 9 pairs, thick CYATHEACEAE (Holttum) 1655 and raised on lower surface. Sori medial, indusiate; indusium thin and translucent, at first complete, breaking and in part persistent; paraphyses short, thin, pale. Scales and hairs: residual scales on pinna-rachis and costae few, narrow with irregular long dark setae; no hairs on upper surface of costules and veins. Type specimen: BopEN KLoss, Mt Carstensz, Camp III, Jan. 1913, W. New Guinea (BM; dupl. at K). Distr. Malaysia: collections). Ecol. At 700-1100 m. West New Guinea (two 185. Cyathea inaequalis Ho_tTumM, Kew Bull. 16 (1962) 56. Trunk 4-6 m, 5-6 cm @g; leaf-scars in alternate whorls of 5. Stipe 24 cm, copiously thorny and scaly near the base; thorns | mm, acute; scales ascending, fleshy at their bases and thinner distally, to 20 by 114 mm, dark and shining except for the thin pale edges of the distal part which bear dark setae; scales above base of stipe very small, brown, the larger ones setiferous. Pinnae: lowest 8 cm long, pinnatifid, largest 50 cm long. Pinnules to 130 by 35 mm, lowest with stalks 3 mm long, apex acuminate, base very unequal (acroscopic segment much larger than basiscopic), 1-2 basal segments of largest pinnules free or nearly so, rest of pinnule lobed to 2 mm from costa; costules 71-814 mm apart; lamina-segments thin, almost entire, apex bluntly pointed and slightly falcate; veins to 9 pairs. Sori medial; indusia complete, thin, breaking and persistent. Scales and hairs: pinna-rachis minutely warty beneath, glabrescent; near base of costae a few small flat elongate seti- ferous scales; no scales seen on costules; no hairs on lower surface of pinnules, nor on upper surface of costules. Type specimen: Brass 23547, Mt Dayman, Milne Bay Distr., Papua (L; dupl. at A). Distr. Malaysia: E. New Guinea collected). Ecol. In rain forest ravine, at 700 m. (once 186. Cyathea parvipinna HOLTTUM, Kew Bull. 16 (1962) 60. Trunk 1 m tall. Stipe 20 cm; spines on stipe and basal part of rachis many, sharp, 1 mm; scales on stipe sparse, to 10 by 2 mm, thick at base, nar- rowed and thinner distally with paler edges bearing dark setae; also on stipe very small brown scales, the larger setiferous. Pinnae: lowest 31 by 1 cm, on stalks 8 mm long, lamina simple; largest 40 cm long. Pinnules to 90 by 18 mm, sessile, acuminate, apical half subentire, basal half gradually more deeply lobed, at base lobed more than half-way to costa, lobes almost entire, thin; costules 5 mm apart; veins in basal lobes 6-7 pairs. Sori medial; indusia thin, translucent, breaking and in part caducous; paraphyses pale. Scales and hairs: pinna-rachis glabrescent beneath, residual scales small, brown, sparingly setiferous; a few rigid brown setiferous scales at bases of costae; on distal part of costae small dark thick bullate scales; no scales seen on costules; no hairs on lower 156 FLORA MALESIANA [ser. Ik yole surface of costae and none on upper surface of costules. Type specimen: BrAss 25837, Normanby I., Papua (L; dupl. at K, US). Distr. Malaysia: E. New Guinea (once collected). Ecol. On banks of stream in forest at 270 m. 187. Cyathea werneri ROSENST. in Fedde, Rep. 5 (1908) 34; v. A. v. R. Handb. (1908) 786; CopeL. Philip. J. Sc. 77 (1947) 101.—C. kingii ROSENsT. in Fedde, Rep. 9 (1911) 422, non (CLARKE) Co- PEL. 1909; v. A. v. R. Handb. Suppl. (1917) 25; CopeEL. Philip. J. Sc. 77 (1947) 100. Trunk to 314 m, bearing c. 8 fronds to 2144 m long. Stipe 8-25 cm; scales to 20 by 144 mm, thick at base, pale and thin distally with setae on the edges. Pinnae: lowest 3-8 cm long, longest to at least 50 cm. Pinnules to 100 by 20 mm, sessile, acuminate, lobed to c. 3 mm from costa; costules 514-614 mm apart; lamina-segments almost entire, distinctly narrowed from the base so that sinuses are triangular. Sori medial; indusium complete, thin, breaking and persistent; paraphyses dark. Scales and hairs: pinna-rachis glabrescent beneath or with small residual scales, not hairs; scales on costae near base small, bearing some dark setae, usually grading to bullate scales distally and on costules; a few thick hairs sometimes near apex of costae on lower surface. Type specimen: WERNER 66, Damun, NE. New Guinea (S-PA; dupl. at P). Distr. Malaysia: S. & E. New Guinea. Ecol. In forest at 200-1200 m. Notes. WAKEFIELD has noted that plants are often fertile when juvenile (i.e. bearing only simply pinnate fronds). Some specimens lack bullate scales, and are then intermediate between this species and C. fusca; possibly the two should be united. 188. Cyathea insularum HoL_trum, Kew Bull. 16 (1962) 57. Trunk 3-5 m, 314-6 cm @ after decay of leaf- bases; leaf-scars 18-25 mm g, in 3 rather steep spirals. Fronds numerous, 165—200 cm long. Stipe 10-15 cm, near base covered with scales; scales 10-15 by 1 mm, dark and thick at the base, distal part dark with pale edges bearing dark setae; rest of stipe sparsely covered with very small pale short-fringed scales. Lowest pinnae less than 5 cm long, rest gradually larger, largest 45 cm. Largest pinnules 65-90 by 18-22 mm, sessile, acuminate, lobed to 1-2 mm from costa; costules 5-6 mm apart; lamina-segments slightly crenate towards obtusely pointed apex; veins 8—9 pairs. Sori medial; indusium complete, thin, pale, breaking and persistent. Scales and hairs: pinna-rachis below bearing small thin pale finely fringed scales; scales on costae throughout pale, bullate, rather large, mostly entire; similar scales on costules; a few thick hairs towards apex of costa on lower surface, no hairs on upper surface of costules. Type specimen: Brass 27419, Misima Island, Louisiades (L). Distr. Malaysia: Louisiade Arch. (on 4 islands). Ecol. In forest, near streams, at 100-350 m. 189. Cyathea tripinnatifida Roxs. Calc. J. Nat. Hist. 4 (1844) 518.—C. integra J. SM. ex Hook. Ic. Pl. (1844) t. 638, p.p. excl. ic.—C. nigrospinulosa v. A. v. R. Bull. Jard. Bot. Btzg II, n. 28 (1918) 15. Stipe to 25 cm (sometimes much shorter ?), bearing scattered thick scales to 20 by 114 mm, setiferous near their apices; stipe also covered with very small dull fringed scales. Pinnae to at least 55 cm long. Largest pinnules 90-110 by 20-28 mm, lobed to 114-2 mm from costa, sessile, short- acuminate; costules 6-7 mm apart; lamina- segments thin but firm, narrowed from the base to broadly pointed apex and so separated by tri- angular sinuses, edges crenate to almost entire; veins to 8 or 9 pairs. Sori nearer to costule than to edge; indusia rather thin, at first complete, break- ing and persistent; paraphyses dark, as long as sporangia. Scales and hairs: \ower surface of pinna-rachis bearing small pale fringed scales, hairs lacking; scales on costae small, fringed with pale hairs, not bullate; a few bullate scales on costules, sometimes caducous; a few thick hairs sometimes present on lower surface of costae and costules near apex of pinnule. Type specimen: Herb. Wallich n. 7076, Mo- luccas (CAL ?; dupl. at K, BM). Distr. Malaysia: Moluccas (Ambon). Ecol. In lowland forest. Notes. Hooker cited a specimen of this species, from Ambon, with his original description of C. integra, but his illustration was prepared from a Philippine specimen, which is thus taken as type of C. integra. Most specimens of C. tripinnatifida lack the stipe. One from Ambon, collected by REINWARDT (L) has a stipe at least 25 cm long, and does not show lower pinnae. 190. Cyathea runensis v. A. v. R. Bull. Dép. Agr. Ind. Néerl. 7. 18 (1908) 1; Handb. (1908) 22.— C. versteegii CHRIst, Nova Guinea 8 (1909) 161; v. A. v. R. Handb. Suppl. (1917) 25. Stipe short, warty; scales dull, dark, more than 1 cell thick at base not fleshy as in C. rosenstockii, Lowest pinnae short, gradually increasing upwards. largest at least 50 cm long. Pinnules to 120 by 25 mm (rarely to 30 mm), lowest segment of largest pinnules free or nearly so, rest lobed to 2 mm from costa; costules 6-7 mm apart; segments of lamina almost entire or slightly crenate, apices rounded, separated by sinuses 1-2 mm wide; veins to 8 pairs. Sori medial; indusium complete, thin, persistent, breaking irregularly; paraphyses dark, as long as sporangia. Scales and hairs: pinna-rachis glabrescent on lower surface, hairs on upper surface short and dark; scales on lower surface of costae small, brown, of rather irregular shape, edges set closely with short spreading rather stiff concolorous hairs or darker setae; bullate scales on costules few, small. Type specimen: TEYSMANN, Pulu Roon, W. New Guinea (BO; dupl. at L). Dec. 1963] CYATHEACEAE (Holttum) iS7 Distr.Malaysia:W.New Guinea, Bismarck Arch. Ecol. In lowland forest. Note. This species is very close to C. tripinna- tifida, and I am not sure of a clear distinction. It seems probable however that a short stipe is in- variable in C. runensis, that of C. tripinnatifida being longer, but very few specimens show this character. 191. Cyathea moseleyi BAK. J. Linn. Soc. Bot. 15 (1876) 104. Stipe short; stipe and lower part of rachis bearing scales to 20 by 114 mm, dark and some- what thickened in the middle with pale edges bearing dark setae. Pinnae: lowest gradually reduced, largest to at least 45 cm long. Largest pinnules to 80 by 18 mm, lobed to c. 1 mm from costa, sessile, acuminate; costules 5-6 mm apart; lamina-segments rather thin, edges more or less crenate, apices bluntly pointed; veins 8—10 pairs. Sori medial; indusium pale, thin, at first complete, breaking and persistent; paraphyses dark. Scales and hairs: pinna-rachis beneath smooth, usually with some residual very small dull brown fringed scales; small fringed scales present on lower surface of costae, with small fringed bullate scales, the latter also abundant on lower surface of costules; a few thick hairs present on upper sur- face of costules but not on lower surface. Type specimen: MoseLey, Admiralty Is (K). Distr. Malaysia: Admiralty Is, Bismarck Arch. Ecol. In lowland forest. Note. C. brackenridgei Metrt., of the Solomon Is, appears to be closely related to this species, but to differ in wider pinnules (to 24 mm wide) and in very abundant scales throughout the stipe and lower part of rachis. Doubtful species Alsophila hallieri RosENst. Med. Rijksherb. 7. 31 (1917) 2.—C. hallieri Domin, Acta Bot. Bohem. 9 (1930) 120. The specimens of HALLIER (735, 737, 738, 4726, 4727) cited with the description have not been found in the Rijksherbarium at Leiden. The brief description does not permit the species to be identified with certainty. Alsophila matthewii Curist, J. Linn. Soc. Bot. 39 (1909) 213.—C. matthewii Doin, Pterid. (1929) 263. I have only seen the specimen from the type collection at Kew; it is a small sterile frond, evidently from an immature plant, said to have been borne on a slender trunk 60 cm high. As the specimen was found in a much-collected locality (Mt Maquiling, Luzon, at 1100 m), it probably belongs to a species already known, possibly C. heterochlamydea CopeL., but I am very uncer- tain of this. Alsophila philippinense Hort. Veitch, Gard. Chron. II, 4 (1875) 179, nomen.—C. veitchiana Domin, Acta Bot. Bohem. 9 (1930) 168. A cultivated plant, described as follows: ‘“‘a handsome new fern, with regularly crimped pinnae from the Philippine Islands”. There is no specimen in Kew Herbarium, and no reference to the plant in Hortus Veitchii. Alsophila speciosa [non (MEYEN) PRESL] GOLD- MANN, Nova Acta 19, Suppl. I (1843) 465. The type of MEYEN’s species came from Brazil: GOLDMANN’s brief Latin description of a Phi- lippine fern is inadequate for identification. Cyathea adenochlamys CurisT, Bull. Herb. Boiss. II, 6 (1906) 1008; v. A. v. R. Handb. (1908) 785. I have not found the type specimen at Paris. Curist described the indusia as persistent, rigid, green, covered densely with pale sessile glands, opening transversely; I have not seen any Cyathea with such indusia. Cyathea grata Domtn, Acta Bot. Bohem. 9 (1930) 120, nomen. Under this name is cited C. frondosa ROSENST. in Fedde, Rep. 12 (1913) 163, but no such species is there described, nor have I found any other reference to C. frondosa ROSENST. Provenance New Guinea. Species not occurring in Malaysia Alsophila truncata BRACK. in Wilkes, U.S. Expl. Exped. 16 (1854) 289; v. A. v. R. Handb. (1908) 42. Malaysian specimens referred to this species are C. celebica BL. or an allied species. Cyathea aneitensis Hook. Syn. Fil. (1865) 26; v. A. v. R. Handb. (1908) 16. After citing specimens from Aneitijum, HOOKER added “‘Ternate, Herb. Hort. Calc.’ There is no Ternate specimen at Kew, and I have seen no Ma- laysian specimens referable to this species. Cyathea rumphii Desy. Mém. Soc. Linn. Paris 6 (1827) 323.—Polypodium arboreum Lour. FI. Coch. (1790) 831, non LINN. DESVAUX published a new name for LOUREIRO’s species, without description; like LouRErIRo, he also cited a plate by RUMPuHius. LouREIRO described a fern from Indo-China, not Ambon; neither his description, nor RuMputus’s plate is clearly identifiable. Therefore I cannot agree with MER- RILL’s suggestion that the name C. rumphii DEsv. should replace C. amboinensis (v. A. v. R.) MERR. (see Trans. Am. Phil. Soc. Philad. n.s. 24, 1935, Sif): Excluded from Cyathea Alsophila dielsii BRAUSE, Bot. Jahrb. 56 (1920) 67. —C. dielsii Domtn, Acta Bot. Bohem. 9 (1930) 111. The type specimen belongs to the genus Di- plazium. 158 FLORA MALESIANA [ser. Il, volume Alsophila dryopteroidea BRrAuseE, Bot. Jahrb. 56 (1920) 70.—C. atrispora Domin, Acta _ Bot. Bohem. 9 (1930) 95.—Dryopteris atrispora C. Cur. Brittonia 2 (1937) 296.—Lastrea dryopteroi- dea CopEL. Gen. Fil. (1947) 138. The type is a Thelypteroid fern; generic limits in this group need to be clarified. Alsophila gazellae KUHN, Forschungsr. Gazelle 4 (1889) 13.—C. gazellae Domin, Pterid. (1929) 262. I have seen the type specimen, and identified it as Pleocnemia cumingiana PRESL (see Reinwardtia 1, 1951, 188). Alsophila tristis (BL.) BL. ex Moore, Ind. Fil. (1857) 58, based on Aspidium triste BL. En. Pl. Alsophila warburgii Curist, Ann. Jard. Bot. Btzg 15 (1897) 80, from Celebes = Dryopteris sp. Alsophila xantholepis Curist ex Digs in E. & P. Pfl. Fam. 1, 4(1899) 138; v. A. v. R. Handb. (1908) 44.—C. xantholepis Domin, Pterid. (1929) 263; Acta Bot. Bohem. 9 (1930) 172 = Dryopteris sp. Cyathea aspidioides (BL.) ZoLt_. & Mor. Syst. Verz. (1846) 108 = Diacalpe aspidioides BL. Cyathea woodlarkensis CopeL. Philip. J. Sc. 9 (1914) Bot. 1; ibid. 77 (1947) 124.—Alsophila woodlarkensis C. Cur. Ind. Fil. Suppl. 2 (1917) 4. This is a large species of The/ypteris, apparently Jav. (1828) 169 = Stenolepia tristis (BL.) v. A. near 7. immersa (BL.) CHING. There are speci- v. R. Bull. Dép. Agr. Ind. Néerl. . 27 (1909) 45. mens in Herb. Copel. (MICH) and at Paris. 2. DICKSONIA L’HErRIT. Sert. Angl. (1788) 30, p.p.; Curist, Farnkr. Erde (1897) 313, p.p.; DieLs in E. & P. Pfl. Fam. 1, 4 (1899) 119; UNDERW. Mem. Torr. Bot. Cl. 6 (1899) 259, 278; C. Cur. Ind. Fil. (1905) xvi, 220; MAxon, J. Wash. Ac. Sc. 12 (1922) 454; Copet. Gen. Fil. (1947) 48.—Balantium KAuLr. En. Fil. Chamisso (1824) 288, p.p. typ.; PRESL, Tent. Pterid. (1836) 134 (excl. Balantium brownianum).— Dicksonia subg. Balantium Hook. Sp. Fil. 1 (1844) 66, p.p.—Dicksonia § Eudickso- nia HooK. & BAK. Syn. Fil. (1868) 50, p.p.—Fig. 31. Stem arborescent, bearing numerous fronds in several spiral series, the vascu- lar tissue supported both internally and externally with strong bands of scleren- chyma; leaf-bases usually persistent; young leaves, and at least the bases of stipes, persistently covered with long hairs; stipes usually short, vascular system complex, in two continuous or divided transverse arcs concave adaxially; pinnae numerous, the lower ones gradually reduced; pinnules usually bearing free deeply lobed tertiary leaflets which are dimorphous, sterile and fertile; upper surfaces of rachises and axes of leaflets raised, bearing rather stiff antrorse hairs, lower sur- faces variously hairy; fertile leaflets more deeply lobed than sterile and with re- duced lamina, each lobe bearing one sorus at the end of the vein or of the basal acroscopic branch if the vein is branched; sorieach protected by a reflexed marginal lobe of the lamina and by a somewhat thinner inner indusium joined to the re- ceptacle on its basiscopic side (fig. 31c); free surface of receptacle slightly elevated and distinctly elongated transverse to the end of the vein; paraphyses numerous, hair-like, multiseptate, each with a red-brown terminal cell; sporangia distinctly stalked, with complete oblique annulus indurated round the base, the stomium lateral; spores trilete, surfaces variously sculptured or almost smooth, exospore more or less thickened at the angles. Type species: Dicksonia arborescens HERItT. (St Helena). Distr. About 25 spp. St Helena; S. America and north to Mexico; New Zealand, E. Australia and Tasmania, New Caledonia, Samoa, Fiji; in Malaysia: New Guinea, Celebes, Philippines, N. Borneo, Java, and Sumatra. Morph. The trunk of Dicksonia is closely similar in form and anatomy to that of Cyathea, from which Dicksonia differs by the indument consisting entirely of hairs. For comment on the possible relationships between the very different soral forms in the two genera, see p. 69. Cytol. Chromosome counts have been made by BROWNLIE for two New Zealand species; both showed 159 Dec. 1963] n = 65 (New Phytol. 56, 1957, 207). MANTON found the same number for D. arborescens (type species of genus) in cultivation at Kew (J. Linn. Soc. Bot. 56, 1958, 84). Taxon. The history of the varied use of the generic name Dicksonia is summarized by MAxon, /.c. Earlier authors confused not only Dicksonia, Culcita and Cibotium, but also Dennstaedtia and other genera now recognized as distinct. Within the genus, the discrimination of species is difficult because the soral form is so constant and the variation in shape of tertiary leaflets as between those on larger or smaller pinnules in one species may be considerable. The character of the hairs on the stipe appears to be one of the clearest distinctions between species, as are scale-characters in Cyathea; but the differences between one hair and another are less easily defined than those between different scales. The species in New Guinea are particularly difficult to discriminate, and the present account can only be regarded as CYATHEACEAE (Holttum) tentative; more field observation is needed. KEY TO THE SPECIES 1. Base of stipe clothed with dark red spreading hairs 30-50 mm long; hairs of under-coat pale, slender or flaccid, much shorter. 2. Hairs of under-coat of stipe-base, and most hairs on lower surface of pinna-rachis and costae, pale, slender, rigid, spreading, not flaccid. Spores smooth . 1. D. blumei 2. Hairs of under-coat of stipe-base, and smaller hairs on lower eure of pinna- -rachis (in some cases also of costa) short, pale, flaccid, their lateral walls collapsed. Spores verrucose. | a= 3. Hairs on costae almost all flaccid, pale, antrorse, matted . 2. D. mollis 3. Hairs on costae mostly rigid, each cell evenly cylindrical. 4. Hairs on costae: some flaccid, some spreading, mostly pale . 4. Hairs on costae ascending, mostly dark red . . 3. D. sciurus 4. D. archboldii 1. Base of stipe clothed mainly with soft matted red-brown hairs, “with or r without a smaller number of stout rigid dark red hairs. 5. Some stout rigid spreading dark red hairs present near base of stipe, and scattered also on pinna- rachis; pinna-rachis mainly clothed with a close layer of pale matted flaccid hairs. 5. D. hieronymi 5. No stout rigid spreading dark red hairs mixed with softer hairs on stipe-base. 6. Pinna-rachis rather dark and + glabrescent; costae bearing flaccid pale hairs + mixed with thicker dark hairs with pale bases 6. D. grandis 6. Pinna-rachis brown, persistently hairy beneath; costae bearing entirely pale flaccid hairs. 1. Dicksonia blumei (KUNZE) Moore, Ind. Fil. (1860) 190; v. A. v. R. Handb. (1908) 47; BACKER & PostH. Varenfl. Java (1939) 22.—Balantium blumei Kunze, Bot. Zeit. 6 (1848) 214.—Ba- jantium chrysotrichum Hassk. Obs. Fil. Jay. 1 (1856) 53.—D. chrysotricha Moore, Ind. Fil. (1860) 190; Hook. & BAK. Syn. Fil. (1868) 50; Ractis. Fl. Btzg 1 (1898) 121; Diets in E. & P. Pfl. Fam. 1, 4 (1899) 121; Curist, Ann. Jard. Bot. Btzg 19 (1904) 41.—Fig. 31a—c. Trunk to 6 m; stipes to 60 cm or more, clothed at the base with spreading red-brown shining hairs 30-50 mm long, with an under-coat of much finer short pale hairs; upper part of stipe and main rachis dark, finely and closely warty, bearing sparse short pale hairs; lamina to 300 cm long; pinnae to 70 cm long, pinna-rachis bearing pale spreading hairs on lower surface, with a few red ones; pinnules to 100 by 15-20 mm; costules of tertiary leaflets 4 mm apart; largest fertile tertiary leaflets lobed throughout almost to the costule, with 4-5 pairs of soriferous lobes, the lowest lobes usually bilobulate with forked vein, sterile lobule not longer than fertile; largest sterile ter- tiary leaflets less deeply lobed, veins in lowest lobe pinnate, in rest forked or simple; hairs on lower 7. D. lanigera surface of costae and costules pale, firm (cell-walls not collapsed), spreading; spores almost smooth. Type specimen: ZOLLINGER 1894, Java (B?; dupl. at P, L); also cited SPORLEDER, Java. Distr. Malaysia: Sumatra (north to Karo Plateau), Java, Central Celebes (SARASIN 2030). Ecol. In mountain forest, 1500-2500 m. 2. Dicksonia mollis HoLTTUM, Kew Bull. 16 (1962) 64.—D. blumei [non (KUNZE) Moore] C. CHR. Gard. Bull. S.S. 7 (1934) 223; CopeL. Fern FI. Philip. 1 (1958) 84.—Fig. 31d-e. Differs from D. blumei as follows: hairs of un- der-coat on the stipe shorter, thicker, but flaccid with walls collapsed when dry; hairs on lower surface of pinna-rachis and costae more numerous, appressed, flaccid, mostly ascending; spores bear- ing conspicuous more or less coalescent warts. Type specimen: ELMER 9874, Dumaguete (Cuernos Mts) Negros Oriental, Negros (K; dupl. at L, P, MICH, US). Distr. Malaysia: NE. Borneo, Central Celebes (2), Philippines (Mindanao to S. Luzon). Ecol. Mountain forests. 1500-2000 m. The Celebes specimen, much smaller than those from FLORA MALESIANA [ser. Il; vole SS Fig. 31. Dicksonia blumei (KUNZE) Moore. a. Part of pinna, showing sterile and fertile pinnules, x 7/, b. part of pinnule, lower surface, showing sori and rigid hairs, < 6, c. section of sorus, * 8.—D. mollis Ho.ttuM. d. Part of sterile pinnule, lower surface, showing flaccid hairs, < 6, e. hairs from lower surface, < 30.—D. antarctica LABILL. f. Section of stipe (hairs omitted), < 2, g. section of pinna-rachis, < 4 (a DE VRIESE S.n., b—-c MATTHEW S.n , d SINCLAIR 9005, e ELMER 9874, f-g cult. R. B. G. Kew). Borneo and the Philippines, was found at 2900 m, in dry open vegetation (EYMA 959); it lacks a stipe. It agrees better with D. mollis than with New Guinea species (it is certainly not D. blumei), but without more complete material its status is doubtful. 3. Dicksonia sciurus C. Cur. Brittonia 2 (1937) 283; CopeEL. Philip. J. Sc. 78 (1949) 6. Stipe 35 cm, the base covered with spreading red-brown hairs 25-40 mm long, with an under- coat of short pale flaccid hairs; upper part of stipe and lower surface of main rachis more sparsely clothed with similar but shorter red-brown hairs and short flaccid pale ones; longest pinnae 60 cm long, rachis dark brown, its lower surface bearing rather sparse spreading reddish hairs and a more or less abraded cover of short flaccid pale hairs; pinnules to 150 mm long (on type not over 80 mm), costules of tertiary leaflets 4-5 mm apart, hairs on lower surface of costae and costules mostly pale, some flaccid and some spreading; tertiary leaflets to 20 mm long, fertile ones lobed almost to cos- tule, basal lobes of largest fertile leaflets having a bluntly-toothed sterile lobule exceeding the fertile one, sterile tertiary leaflets less deeply lobed than fertile, lobes mostly with pinnately arranged veins and bluntly toothed edges. Type specimen: Brass 4991, Mt Tafa, Papua (BM; dupl. at BO, MICH, BRI, UC). Distr. Malaysia: E. New Guinea (several collections). Ecol. Plentiful in ridge forest, at 1800-2400 m. 4. Dicksonia archboldii CoreL. Philip. J. Sc. 78 (1949) 6, pl. 1. Dec. 1963] Stipe 30-40 cm, base clothed with spreading rather soft red-brown hairs 25 mm or more long, with short flaccid pale hairs also; leafy part of frond to c. 300 cm long; lower pinnae gradually reduced, lowest 15 cm long, longest 65 cm, pinna- rachis beneath dark, clothed rather sparsely with dark red rigid hairs, the smallest ones with a pale inflated basal cell, also short pale flaccid hairs; pinnules to 140 mm long, costa covered beneath with rigid flexuous antrorse dark red hairs, some with a pale base, and on distal part a few pale flaccid ones; costules of tertiary leaflets 5-6 mm apart; sterile fertiary leaflets to 20 mm long, lobed to costule at base, lobes to 6 pairs, veins pinnate in each, edges toothed; fertile tertiary leaf- lets lobed throughout to costule, lobes to 5 pairs, each with a sorus, the basal ones also with an acute sterile tooth. Type specimen: Brass 10970, Lake Habbema, 2750 m, near Mt Wilhelmina, W. New Guinea (MICH; dupl. at L). Distr. Malaysia: West New Guinea, only known from the type collection and a much smaller one from Mt Arfak, Anggi Lakes (SLEv- MER & VINK BW 14193). Ecol. “Abundant in forest of lower slopes, stem 4 m high, 13 cm o under the leaves and thickened downwards; leaves 10, spreading’. Note. This is near D. sciurus C. Cur. (an earlier name) and the two should perhaps be united. The type of D. archboldii is larger than that of D. sciurus in all parts of the frond, and the hairs on the costae are mostly red and ascending; the fertile tertiary leaflets of D. sciurus have in some cases a larger sterile portion, but this varies ac- cording to position on the frond, and whole fronds of each type are not available for comparison. 5. Dicksonia hieronymi BRAUSE, Bot. Jahrb. 56 (1920) 48; C. Cur. Brittonia 2 (1937) 282; CopeEL. Philip. J. Sc. 78 (1949) 6. Stipe 20-30 cm, covered near the base with soft dull brown matted hairs with a few longer rigid shining dark red-brown ones; upper part of stipe and main rachis dark, rather persistently covered with appressed soft brown matted hairs with a few rigid shining ones; largest pinnae 30-60 cm long; pinna-rachis dark brown, hairy on under surface as main rachis; largest pinnules 70-120 mm long, costa pale distally, dark near base, covered beneath with flaccid + appressed light brown hairs, some with rigid dark apical portion; largest tertiary leaflets 10-20 mm long, fertile lobed to the costule, fertile lobes 3—5 pairs, the larger with forked vein and a sterile lobule which is rarely larger than the fertile one; sterile tertiary leaflets deeply lobed at base, less deeply upwards, larger lobes containing pinnately branched veins, edges of lobes almost entire. Type specimen: LEDERMANN 12851, Sepik re- gion, 1400-1500 m, E. New Guinea (B). Distr. Malaysia: New Guinea (several locali- ties) including Louisiade Arch. Ecol. In mossy forest, at 1400-3000 m; fronds usually few. CYATHEACEAE (Holttum) 161 6. Dicksonia grandis ROsENsT. in Fedde, Rep. 5 (1908) 34; CopeL. Philip. J. Sc. 78 (1949) 5.— D. schlechteri Brause, Bot. Jahrb. 49 (1912) 11: CopeL. Philip. J. Sc. 78 (1949) 6.—D. ledermannii BrAuse, Bot. Jahrb. 56 (1920) 46; CopeL. Philip. J. Sc. 78 (1949) 5. Stipe at least 30 cm, nearly black, lower part covered with fine soft light brown hairs, longest 25 mm, all of same texture; pinnae to 100 cm long, pinna-rachis beneath very dark, minutely warty, more or less glabrescent, with vestiges of short appressed hairs which are mostly lax and pale brown, with some firm and reddish (the latter sometimes with pale bases); pinnules to 140 mm long, costa hairy beneath as pinna-rachis, dark at base and paler distally; tertiary leaflets to 20 mm long, fertile ones deeply lobed, lobes to 5 pairs, each lobe of largest leaflets with a pinnate vein bearing a sorus on its basal acroscopic branch, of smaller leaflets sometimes with simple vein bearing a sorus; sterile tertiary leaflets lobed almost to costule at base, less deeply upwards, each lobe of larger ones containing a pinnately branched vein, edges of lobes almost entire. Type specimen: WERNER 79, in forest, 1000 m, Mt Gelu, E. New Guinea (also distributed as RosEnsT. Fil. Novoguin. exsic. 7. 50 (B; dupl. at ES eSUsSs Ue): Distr. Malaysia: New Guinea (several localities). Ecol. In forest, most specimens at 1000-1800 m; one (PULLE 939, Mt Hellwig) from 2500 m, a small plant but agreeing in hairiness. 7. Dicksonia lanigera HottrumM, Kew Bull. 16 (1962) 64. Stipe to at least 30 cm, densely hairy throughout, hairs uniform, soft, medium brown, to 15 mm, the shorter hairs of same texture, no thick firm red hairs present; /amina c. 150 cm long; rachis beneath red-brown when dry, covered with matted short hairs like those of the stipe; pinnae to 40 cm long, pinna-rachis persistently hairy as rachis on lower surface; largest pinnules 80 mm long, cos- tules 514 mm apart; costa beneath brown-hairy almost to apex; fertiary leaflets to 15 mm long, sterile ones deeply lobed near the base and less deeply upwards, larger lobes each with a pinnate vein, edges toothed; fertile tertiary leaflets deeply lobed throughout, vein in larger lobes forked, acroscopic branch bearing a sorus, sterile part of lobe not longer than fertile. Type specimen: PuLLte 1010, Mt Wichman, 3000 m, SW. New Guinea (L; dupl. at BM). Distr. Malaysia: W. & E. New Guinea. Ecol. In forest, at 2500-3000 m. Excluded from Dicksonia Dicksonia ampla Bax. J. Linn. Soc. Bot. 22 (1886) 223 = Dennstaedtia ampla (BAK.) BEDD. Dicksonia cuneata Hook. Sp. Fil. 1 (1844) 80 = Dennstaedtia cuneata (HOOK.) Moore. Dicksonia deltoidea Hook. Sp. Fil. 1 (1844) 80 = Dennstaedtia scabra (WALL.) Moore. 162 FLORA MALESIANA [ser. II, vol. 12 Dicksonia erythrorachis Curist, Ann. Jard. Bot. Dicksonia neglecta Fe, Gen. Fil. (1850-52) Btzg 15 (1897) 86 = Dennstaedtia erythrorachis 335 = Dennstaedtia smithii (HooK.) Moore. (CHRIST) DIELS. Dicksonia_ nephrolepioides Curist, Verh. Nat. Dicksonia flaccida (Forst.) Sw. in Schrader, J. Ges. Basel 11 (1895) 241 = Nephrolepis dicksonioi- Bot. 1800, ii (1801) 90 = Dennstaedtia flaccida des CHRIST. (Forst.) BERNH. Dicksonia remota Curist, Verh. Nat. Ges. Dicksonia glabrata Ces. Rend. Ac. Napoli 16 Basel 11 (1896) 423 = Dennstaedtia remota (1877) 24, 28 = Dennstaedtia glabrata (CEs.) (CurRisT) DIELs. C. CHR. Dicksonia rhombifolia Bax. J. Bot. 28 (1890) Dicksonia gomphophylla Bax. J. Linn. Soc. Bot. 105 = Dennstaedtia rhombifolia (Bak.) C. CHR. 22 (1886) 223 = Dennstaedtia gomphophylla : ; (BAK.) C. Cur. Dicksonia scabra WALL. ex Hook. Sp. Fil. 1 (1844) 80 = Dennstaedtia scabra (WALL.) MOorRE. Dicksonia japonica Sw. in Schrader, J. Bot. 1800, il (1801) 92 = Méicrolepia strigosa (THUNB.) PRESL. Dicksonia scandens Bui. En. Pl. Jav. (1828) 240 = Dennstaedtia scandens (BL.) Moore. Dicksonia smithii Hook. Sp. Fil. 1 (1844) 80 = Dicksonia kingii BEpp. Handb. Suppl. (1892) Dennstaedueaemi iit (HOO) Noort 6 = Orthiopteris kingii (BEDD.) HOLTTUM. Dicksonia strigosa THUNB. Trans. Linn. Soc. 2 Dicksonia linearis Cav. Descr. (1802) 274 = (4794) 341 = Microlepia strigosa (THUNB.) Tapeinidium pinnatum (CAv.) C. CHR. PRESL. Dicksonia moluccana BL. En. Pl. Jay. (1828) Dicksonia zippeliana KUNzE, Bot. Zeit. 3 (1845) 239 = Dennstaedtia moluccana (BL.) Moore. 838 = Dennstaedtia scandens (BL.) Moore. 3. CYSTODIUM J. Sm. in Hook. Gen. Fil. (1841) t. 96; Hist. Fil. (1875) 258.—Fig. 32. Stem massive, usually creeping, covered with long hairs. Fronds tufted, bipin- nate; stipe grooved on adaxial surface, vascular tissue arranged as in Dicksonia; upper surface of pinna-rachis antrorsely hairy and grooved, the groove not open to admit the grooves of midribs of pinnules; pinnules slightly dimorphous, sterile ones serrate, fertile ones with slightly narrower lamina and teeth enlarged to protect sori; veins in pinnules pinnately arranged, oblique, often forked in larger pinnules, simple in smaller ones except for the basal pair which may be pinnate; sori terminal on veins, receptacle slightly raised, round in section, tooth of lamina reflexed and enlarged, forming an outer indusium, inner indusium smaller and thinner, attached only at the base of the sorus; paraphyses abundant, as simple multiseptate hairs lacking a terminal glandular cell; sporangia stalked, stalk as long as body of the sporangium, annulus complete and slightly oblique but not indurated at base where it passes the stalk; spores trilete. Distr. Monotypic, Malaysia: from NE. Borneo to New Britain & Louisiades. Notes. The single species agrees with Dicksonia in hairs and their distribution, in arrangement of vascular tissue in the frond, and in form of sori; it differs from Dicksonia (1) in normally prostrate position of stem, (2) in grooved upper surface of rachis, pinna-rachis and costules, (3) in shape of pin- nules, which are crenate or serrate (not deeply lobed as in Dicksonia), (4) in circular transverse section of receptacle of sorus, (5) in smaller sporangia on longer stalks, and (6) almost vertical annulus, not indurated at the base. JOHN SMITH (/.c. 1875) stated that the pinnules are jointed to the rachis (a character to which he at- tached great importance), but they are not so. Dec. 1963] CYATHEACEAE (Holttum) C i2 Kee \Qiaz IS . tt, EWS mS — — hd ty Sy Fig. 32. Cystodium sorbifolium (SM.) J. SM. a. Middle part of rachis with pinnae, x 24, b. lower surface of part of pinna, near its base; sporangia removed from one sorus to show inner indusium, x 6, c. section of stipe, x 4, d. section of pinna-rachis, way to costa. 68. Reduced pinnae conspicuous, upper ones 1-3 cm long. 69. Pinnae 12 x 1.6 to 20 x 2. cm, gradually acuminate; lower surface of costae glabrous or with sparse short hairs oe 4 . . . 57.8. latebrosus 69. Pinnae to 14x 2.5cm, abruptly short- acuminate: low. er ‘surface of costae densely covered with appressed hairs . . ..... . . . . . 458.8. porphyricola 68. Reduced pinnae all very small. 70. Lower surface of rachis and costae almost glabrous . . . . 59.8. caulescens ps Lower surface of rachis and costae bearing hairs to | mm long. . Largest pinnae commonly 15-20cm long: hairs on rachis pale . . 21.S.hirsutus a Largest pinnae 7x 1.5cm; hairs onrachis brown . . . . . . 60.S.reconditus 67. Pinnae lobed conspicuously more than > way to costa. 72. Hairs on lower surface of costules antrorsely appressed. 73. Sori supramedial; pinnae to 2.5 cm wide. 74. Basal normal pinnae not narrowed at base . . . . 61.8. cyrtocaulos 74. Several pairs of lower pinnae much narrowed at their ‘bases . . 62.8. baramensis 73. Sori inframedial; pinnae to 1.5 cm wide -9 £Seyeoee Alera 63585 batekitessis 72. Hairs on lower surface of costules not antrorse. 75. Reduced pinnae all conspicuous, lowest 5 mm long . . . . . 64.8. subalpinus 75. Reduced pinnae all small, uppermost 5 mm long. 76. Dark hairs on both sides of rachis . . . ..... . =. - . 65.8. lobatus 76. Pallid hairs on both sides of rachis. 77. Lowest pinnae little narrowed at base; reduced pinnae evident . 66. S. ellipticus 77. Lower pinnae much narrowed at base; all reduced pinnae minute 67. S. foxworthyi 65. No appressed hairs on upper surface between veins. 78. Hairs on lower surface of costules copious, closely appressed. 79. Reduced pinnae all very small, basal normal pinnae narrowed at base. 80. Pinnae lobed less than > tow ardsicosta) ney 4ial (92) Seay. 9). = 6S: Ss iedrapurae 80. Pinnae lobed about ¢ Sige . . . 62.8. baramensis 79. Reduced pinnae conspicuous, many 10-15 1 mm 1 long: basal normal pinnae not narrowed at base ee. aut oe -. (69) Sc hbatacorum 78. Hairs on lower surface of costules short, ‘not appressed. 81. Sori indusiate; sporangia glandular, rarely with a seta. 82. Lowest normal pinnae narrowed at base Stes cguct BO 2A e Se aneestibass 82. Lowest normal pinnae not narrowed atbase . ... . . . . 57.8. latebrosus 81. Sori exindusiate; neither glands nor setae on sporangia. 83. Pinnae thin. lobed to2mmfromcosta ...... . . . . 71.S8.nudisorus 83. Pinnae subcoriaceous, lobed little more than 3 . «eat & 6722S. paripomains 61. Aerophores not or slightly elongate. 84. Basal veins both passing to sides of sinus-membrane or uniting just below it, not or rarely forming a distinct excurrent vein. 85. Reduced pinnae all with distinct lamina; costules 2.5mm apart . 73.S.novae-britanniae 85. Reduced pinnae without distinct lamina; costules 4 mmapart .. . . 74.S.convergens 84. Basal veins uniting to form an excurrent vein, at least near base of pinna. 86. Sori supramedial; caudex a long-creeping rhizome > ces. ceeds Ss Seis 86. Sori medial or inframedial; caudex not long-creeping. 87. Pinnae to 2.5 cm long; dark hairs on upper surface of rachis . . . 76.8. sessilipinna 87. Not this combination of characters. 88. Lower surface of rachis glabrous or nearly so; hairs on lower surface of base of costae none or very short. 89. Reduced pinnae all very small. 90. Largest pinnae c.6cmlong .......-.-.- - - - - - 77.S.lobangensis 90. Largest pinnae at least 10cm long. : 91. Fertile pinnae to 16X2cm,lobed atmosts ... . .- . - 78.S.gymnorachis 91. Fertile pinnae to26x3cm,lobed? . . . . . 79.8. pullenii 89. Reduced pinnae: upper ones 1.5-2 cm long. lower gradually smaller. 92. Pinnae lobed toadepthof2mm ........ . . . . -80.S.gregarius 444 FLORA MALESIANA [ser. Il; voli 92. Pinnae lobed at least 3 wayatorcostal 4 = > i s . . « 57Slatebrosus 88. Lower surface of rachis and costae hairy throughout. 93. Pinnae crenate. 94. Pinnae not more than 4.5 x 1.4 cm; veins 4-5 pairs, I-13 pairs anastomosing. 95. Pinnae 5-6 pairs SAS Ree an SO BRS ee y SI Satephrophyllus 95. Pinnae 18 pairs . . . . . 17.8. kalkmanii 94. Sterile pinnae to 8 x 1.7 cm; 23 pairs of veins anastomosine:! sinus-membrane very short 20. S. lamii 93. Pinnae distinctly lobed. 96. Fronds dimorphous; sterile pinnae to 25 x 3.5 cm . . . . $2.8. pilososquamatus as Fronds not or little dimorphous, pinnae not over 2.5 cm wide. . Pinnae to 9.5X0.8-0.9cm . . . . . . .83.S. angustifolius a Pinnae proportionately wider, to at least 1. 5 cm n wide. 98. Lowest normal pinnae much narrowed at base . . . . . 24,8. heterocarpus 98. Lowest normal pinnae not narrowed at base. 99. Hairs | mm long on lower surface of rachis and costae; indusia with long hairs. 100. Pinnae lobed less than 1/2 way tocosta . . . . . . . . .84. S. efogensis 100. Pinnae lobed more than 1/2 way to costa. 101. Reduced pinnae CASmpairsie Ya 20 TE SS 85. Seidichnotnichoides 101. Reduced pinnae many pairs aa . . . 3. S. polycarpus 99. Hairs inmost cases less than 0.5 mm long on nilower surface of rachis and costae; glands present on indusia. 102. Lowest veins sometimes both touching sides of sinus-membrane; pinnae lobed 2/3-3/4 Hy as . . . 86.8. nakaikei 102. Lowest veins always aReCtomosine: pinnae lobed & ae 103. Pinnae rigid, lobes of larger ones toothed distally; reduced pinnae not over 8 pairs. 104. Reduced pinnae 4-5 pairs; upper surface of pinnae bearing appressed hairs between veins aah . . . . 87. S. woitapensis 104. Reduced pinnae 8 pairs; upper surface of pinnae lacking appressed hairs 88. S. ekutiensis 103. Pinnae thin, lobes not toothed; reduced pinnae 12-15 pairs. 105. Hairs on lower surface of costae not appressed. 106. Hairs on lower surface of costae all very short . . . . 57.8. latebrosus 106. Some hairs 0.5 mm or more long on lower surface of costae 66. S. ellipticus 105. Lower surface of costae covered with closely appressed hairs 58. S. porphyricola 35. Glands between veins on lower surface of pinnae lacking. 107. Glands present on lower surface of costules and veins. 108. Pinnae 3-4 pairs, to 1.1 x 0.5 cm, basal pinnae somewhat reduced . . . . 89.S. omatianus 108. Pinnae more numerous or larger, or several pairs reduced basal pinnae. 109. All pinnae with narrowly cuneate basiscopic base . . oe. ee ee). 44,8. aquatilis 109. Pinnae symmetrically broadly cuneate or truncate at base. 110. Few to many pairs of lower pinnae gradually reduced. 111. Pinnae subentire to crenate. 112. Pinnae to 3.5 x 1.0cm; pinna-lobes with strongly reflexed edges. 113. Sporangia setiferous; 12-14 pairs of lower pinnae gradually reduced . -. 90.8. alticola 113. Sporangia not setiferous; 4-5 pairs of lower pinnae gradually reduced . 91.8. rigidus 112. Pinnae larger, edges of lobes not strongly reflexed. 114. Sori exindusiate; lower surface of costae and costules glabrous 56. S. hastatopinnatus 114. Sori indusiate; short hairs abundant on lower surface of rachis and costae 2 92. S. arfakianus 111. Pinnae lobed 3-5 towards costa. 115. 1-4 pairs of basal pinnae gradually reduced . . . . . . . . +. 43.8. adenostegius 115. Many pairs of basal reduced pinnae. 116. 7-12 pairs of lower pinnae gradually reduced with subabrupt change to 12 pairs much omgilie 5 5 « . . . . . . 55.8. hispiduliformis 116. Similar transition but 2 2 6) pairs ae grail ihasal pinnae eee ee = 95>) Ssarchboldit 110. Transition to reduced basal pinnae (if any) abrupt or nearly so. 1981] THELYPTERIDACEAE (Holttum) 445 117. Reduced pinnae 0-3 pairs. 118. Pinna-lobes strongly concave beneath . .... . woah 28% & °91; Sarigidus 118. Pinnae-lobes not concave beneath. 119. Pinnae widened distally pe ee Tass AOR Aree ee LAS diversilobus 119. Pinnae not widened distally. 120. Caudex short-creeping; pinnae lobed c. § towards costa. 121. Fronds not dimorphous; sori supramedial. 122. Pinnae 3.5 X 1.4cm, narrowed evenly toapex . .. . . . . . 94.S. tibangensis 122. Pinnae larger, abruptly short-acuminate ....... . . . . 9§5.S.norrisii 121. Fronds dimorphous; sorimedial . . . . . . . . . . . 82.8. pilososquamatus 120. Caudex slender, erect; pinnae crenate ...... . .. . . .96.S.semimetralis 117. Reduced pinnae at least 4 pairs. 123. Aerophores elongate. 124. Pinnae crenate. 125. Hairs on lower surface of costae minute .... . . . . . 97.8. semicordatus 125. Hairs on lower surface of costae appressed . . .. . . . . . 98.S. pterosporus 124. Pinnae lobed at least 2/5 towards costa. 126. Hairs on lower surface of costules appressed fy ee ee Sie 9SaSspterosporus 126. Hairs on lower surface of costules not appressed. 127. Reduced pinnae 20-30 pairs; normal pinnae lobed less than > way to costa. 128. Reduced pinnae acuminate Lee Re es Pg BAe OE 99 Sssagittifolius 128. Reduced pinnae with rounded tips ayn: . . . 100.8. foliolosus 127. Reduced pinnae not more than 12 pairs; normal pinnae lobed ) way to costa or more deeply. 129. Pinnae lobed ; to costa. 130. Basal veins anastomosing; upper reduced pinnae 12mm long . . . 101.S. alpinus 130. Basal veins free; upper reduced pinnae 3 mm long . . . . 54,8. novoguineensis 129. Pinnae lobed about 2 to costa. 131. Pinnae c. 6.5 X 1.3 cm, lowest narrowed at base; indusia glabrous 102. S. polisianus 131. Pinnae c. 11 x 2.cm, lowest not narrowed at base; indusia bearing long hairs 103. S. hoalensis 123. Aerophores not elongate. 132. Sorielongate .. see PREP A ee eee > ey Pen O45 Sslarutensis 132. Sori not or little elongate. 133. Reduced pinnae c. 30 pairs, deflexed, overlapping. 134. Reduced pinnae acuminate; normal pinnae 20cmlong . . . . . 99.S. sagittifolius 134. Reduced pinnae with rounded tips; normal pinnae smaller . . . . 38.8. polyotis 133. Reduced pinnae not over 12 pairs, not overlapping. 135. Sori supramedial. 136. Pinnae lobed 3, sporangia setiferous . . oe. . we) 6. 105.8. loherianus 136. Pinnae lobed less than 3; sporangia glandular ORT se 7S sSsunitus 135. Sori not supramedial. 137. Pinnae lobed more than }. 138. Sori indusiate; sporangia with glands. 139. Largest reduced pinna 7 mm long; hairs on lower surface of costae not appressed 106. S. erectus 139. Largest reduced pinna 2 mm long; hairs on lower surface of costae appressed 98. S. pterosporus 138. Sori exindusiate; sporangia setiferous . . . . . . . . . 107.S.stresemannii 137. Pinnae crenate or lobed less than >. 140. Pinnae not over 1 cm wide; indusia very small or lacking . . . 17.8. kalkmanii 140. Pinnae more than 1 cm wide; indusia all distinct. 141. Some hairs on lower surface of rachis and costae | mm or more long. 142. Pinnae commonly more than 2 cm wide, crenate to a depth of | mm. 143. Apex of frond pinna-like, crenate to base . . . . . . 108. S. simplicifolius 143. Apex of frond deeply lobed at base « OR? SOR 2 eS A109SSsspenceni 142. Pinnae to 1.7 cm wide, distinctly lobed . . . . . . . . 84.8. efogensis 141. Hairs on lower surface of rachis and costae all very Short 144. Largest reduced pinnae 2 mm long » 2, oe 2) a ee 2 ee 98aSopterosporus 144. Largest reduced pinnae at least 7 mm long. 446 FLORA MALESIANA [ser. II, vol. 1° 145. Sporangia not setiferous. 146. Pinnae opposite; caudex 40 cm tall . oe. . . . . . . 110.8. mengenianus 146. Pinnae not opposite; caudex suberect SVs Se oe S05Ssoreganins 145. Sporangia setiferous . . soon A eee aye SMS Sconfertus 107. Glands lacking on lower surface of costules and \ veins. 147. Reduced pinnae c. 30 pairs. 148. Reduced pinnae acuminate. STAT a ce hae ee eee ease bene ep 299RSSsacittifolius 148. Reduced pinnae with rounded tips Via) bose toe Sk) .e 8 Veer QS8aSspolyotis 147. Reduced pinnae much fewer. 149. Sporangia bearing glands (sometimes a seta also). 150. Hairs on lower surface of costules appressed. 151. Aerophores elongate. 152. Several pairs of lower pinnae gradually reduced . o. . . . . 112.8. multiauriculatus 152. Transition to reduced pinnae + abrupt. 153. Upper reduced pinnae 12mm, lowest6mmlong ... . . . . . 113.S.stipellatus 153. All reduced pinnae very short. 154: *Pinnae lobed léssithania. = = < Aino) gb istesi ).. eee Soucy AGREES! pterosporus 154. Pinnae lobed more than 3. 155. Indusia large, firm . .watsis- 2. eee 3) eee SS lonitotine 155. Indusia small, thin. 156. Basal veins anastomosing, second acroscopic vein to sinus-membrane 115. S. hendersonii 156. Basal veins often connivent at sinus-membrane, next veins both to margin 116. S. posthumii 151. Aerophores not elongate. 157. Basal acroscopic vein ending beside sinus-membrane, basiscopic vein at margin above base of sinus. 158. Pinnae 15-20 pairs; reduced pinnae 4-7 pairs . oo. . . . . . . 117.8. inconspicuus 158. Pinnae to 30 pairs; reduced pinnae 2-3 pairs . . . . . .118.S. subulifolius 157. Basal veins both touching sinus-membrane or uniting below it. 159. Gradual transition to reduced pinnae at base of frond. 1605sindustaspresent’ 0 A seen Sea le eee es (eo 2a Saantakianns 160. Indusia absent eS: . oo. . . . . . 112.8. multiauriculatus 159. Abrupt transition to very ‘small reduced pinnae . oe... . . . . 98. S. pterosporus 150. Hairs on lower surface of costules not appressed. 161. Basal veins not connivent below sinus. 162. Pinnae deeply lobed. 163. Pinnae to 3.6 x 1.0cm; veins 3-4 pairs 8S 4 een ee Sea. Deo eS eSSmiyohersii 163. Pinnae to 10 x 1.5 cm; veins 9-10 sa . oe ee ee). 54.8. novoguineensis 162. Pinnae almost entire . . . . . . 120.8. mutabilis 161. Basal veins connivent below sinus or uniting to form an excurrent vein. 164. Pinnae to 4.2 x 1.1 cm; sori elongate with small indusia . . . . . . 4121.S.uaniensis 164. Pinnae in most cases much larger; sori otherwise. 165. Ten or more pairs of basal pinnae gradually reduced. 166. Pinna-margins sinuous or slightly crenate . . . . . . . 56.8. hastatopinnatus 166. Pinnae deeply lobed A ee . oe. ee ws). . 122.8. echinosporus 165. Abrupt transition to reduced pinnae if present. 167. Reduced pinnae 0-4 pairs on mature plants. 168. Pinnae almost entire, 3.5cm or more wide ... . . . . . 123.8. peltochlamys 168. Pinnae distinctly lobed, narrower. 169. Pinnae lobed more than uitiecte tlaniee: “Abid diad. oxv', Wer seegiaaass irayensis 169. Pinnae lobed less than}. . spike alot), Se RES SS jnornisit 167. Reduced pinnae at least 6 pairs on mature plants. 170. Sori supramedial. 171. Caudex a long-creeping rhizome ecb aivnd Je. 1c. cee A SaSamitus ijt Caudexeshontenectar), pace tees tes) tees Goes Sel eA SAI25SSalithophyllus 170. Sori medial or inframedial. 172. Pinnae crenate : PURtye AG abe) he ay Wenst Be sues LOSS sspenceri 172. Pinnae lobed at begi2 173. Pinnae lobed more than } Rea tye Ges (eee $590ae0.2> 391262 Sshamiferus 1981] THELYPTERIDACEAE (Holttum) 447 173. Pinnae lobed less than> . . . jucret. 10. tere See BSOFSZerecarius 149. Sporangia lacking glands, in most cases bearing setae. 174. Aerophores elongate. 175. Pinnae entire or slightly crenate; veins 18-20 pairs . . . . . . . 127. S. plurivenosus 175. Pinnae distinctly lobed; veins rarely more than 12 pairs. 176. Pinnae lobed less than > 177. Hairs on lower surface of costae and costules appressed. 178. Lower 12-15 pairs of pinnae gradually reduced, lowest 1 cm long 112. S. multiauriculatus 178. Transition to small pinnae at base of frond abrupt . . . . . . 128.S.squamatellus 177. Hairs on lower surface of costae and costules not appressed. 179. Reduced pinnae more than 12 pairs; sporangia not setiferous . . 55.S. hispiduliformis 179. Reduced pinnae to 5 pairs; sporangia setiferous Ay ovata Wie) ee 29S. canescens 176. Pinnae lobed more than 3. 180. Sporangia always with several setae. 181. Basal 2-3 pairs of pinnae somewhat narrowed at their bases. 182. Pinnae to 11 x 2.1 cm, lobed <3; hairs on lower surface of costules antrorse 130. S. suboppositus 182. Pinnae to 17 x 2.8cm, lobed more than 4; hairs on lower surface of costules erect 131. S. makassaricus 181. All pinnae in lower half of frond narrowed at their bases, lowest much narrowed 132. S. sarasinorum 180. Sporangia rarely with a seta. 183. Pinnae lobed 4; or more deeply; reduced pinnae all very small. 184. Lower surface of rachis glabrous vee . . . . 62.8. baramensis 184. Lower surface of rachis densely covered with erect brown hairs . 133. S. muluensis 183. Pinnae lobed at most 3; reduced pinnae 1-1.5 cm long . . . . . 126.8. hamiferus 174. Aerophores not elongate. 185. Sporangia not setiferous. 186. Pinnae not or little more than 5 cm long. 187. Pinnae crenate; sori inframedial. 188. Pinnae 10-20 pairs; 2-3 pairs of lower ones gradually reduced. 189. Caudex slender, erect; pinnae slightly crenate distally . . . . 134.S. telefominicus 189. Caudex very short; pinnae crenate throughout . . . . . . . 135.S.roemerianus 188. Pinnae to 30 pairs; abriet change to reduced ones at base . . . 136.S. hellwigensis 187. Pinnae lobed more than 2; sori supramedial at. hes «ate els 7iSsbenoitianus 186. Pinnae more than 10cm long. 190. No reduced pinnae; normal pinnae distinctly stalked . . . . . 138.8. neotoppingii 190. At least 5 pairs lower pinnae gradually or abruptly reduced; all pinnae sessile. 191. Sori elongate and indusiate. 192. Normal pinnae not or little more than 2 cm wide; upper reduced pinnae 4 mm long 139. S. oosorus 192. Normal pinnae commonly 2.5—3.5 cm wide; reduced pinnae much larger 104. S. larutensis 191. Sori not elongate, or if so exindusiate. 193. Abrupt transition to 6 pairs of very small basal pinnae . .. . . . . 140.S.rudis 193. Basal pinnae not very small. 194. Rachis quite glabrous on lower surface; pinnae almost entire . . 141.S.dimorphus 194. Rachis hairy on lower surface; pinnae crenate or lobed throughout 142. S. potamios 185. Sporangia setiferous. 195. Caudex terrestrial and long-creeping, or scandent. 196. Caudex terrestrial; indusia distinct : = laut, tee ee 1435S sinvisus 196. Caudex scandent; indusia very small or lacking. 197. Several pairs of lower pinnae gradually smaller and auricled on acroscopic base 144. S. austerus 197. Transition to basal reduced pinnae abrupt . . . . . . . . +. . +. 145.S. mundus 195. Caudex various, not long-creeping nor scandent. 198. Lower 1-S pairs of pinnae gradually reduced. 199. Pinnae to2cmlong;noindusia . . 50a. itd an = 6S obtusifohus 199. Pinnae to at least 4cm long; indusia present J 2” 5 eactldewiat Leoeee 435 Syadenostegius 448 FLORA MALESIANA [ser. II, vol. 1° 198. At least 3 pairs of reduced pinnae distinct from normal ones. 200. Pinnae lobed 3 way to costa or more deeply. 201. Reduced pinmae 20 pairs, all very small 201. Reduced pinnae to 12 pairs, all eee 202. Caudex short, erect; pinnae lobed 2 202. Caudex tall, slender, if free-standing; pinnae lobed 3 = 200. Pinnae lobed less than 3 way to costa. . 147. S. alcasidii 148. S. pilosissimus . 145. S. mundus 203. Pinnae to 3.5 cm long, slightly crenate 149. S. erwinii 203. Pinnae in most cases larger, deeply crenate or lobed. 204. Stipe 40-50 cm long; reduced pinnae 3 pairs, small 82. S. pilososquamatus 204. Stipe much shorter; reduced pinnae more numerous, conspicuous. 205. Pinnae crenate; no long hairs on lower surface of rachis and costae. 206. Caudex erect; largest fertile pinnae 17x 2.5 cm 206. Caudex short-creeping; largest pinnae 8 x 2cm . 150. S. atasripii 111. S. confertus 205. Pinnae distinctly lobed; long hairs usually present on lower surface of rachis and costae. 207. Stipe-scales 1 mm long; pinnae to 4 Icm 207. Stipe-scales 7 mm long; pinnae to 12 x 2cm 1. Sphaerostephanos debilis (METT.) HOLTTUM, comb. nov.— Phegopteris debilis METT. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 123, t. 6 f. 1; v.A.v.R. Handb. (1908) 501.— Dryopteris debilis (METT.) C. CHR. Ind. Fil. (1905) 260.— Cyclosorus debilis (METT.) COPEL. Gen. Fil. (1947) 142, non CHING 1941. — Thelypteris debilis (METT.) REED, Phytologia 17 (1968) 270. — Type: ZIPPELIUS, Amboina (L). Caudex short, erect; stipe 34cm long. Lamina 30-40 cm long, gradually attenuate both to base and apex; basal pinnae 5mm long; texture thin. Largest pinnae 3.0cm long, 0.7cm wide above base; base truncate, 1.0cm wide, auricled on acroscopic side, slightly dilated on basiscopic, distal part gradually narrowed to a rounded apex, edges slightly crenate; costules 2.5mm _ apart; veins 2 pairs, lower pair anastomosing; sparse hairs | mm long on lower surface of rachis, sparse minute hairs on costae and costules, small spheri- cal glands throughout; minute suberect hairs and glands between veins on upper surface. Sori in- framedial, small, exindusiate; sporangia bearing 1-2 short setae and sometimes a gland distally and a sessile gland on the stalk. Distr. Malesia: Moluccas (Amboina, without collector’s name at B, K, L, also one cult. Hort. Bog. at L); W. New Guinea. Ecol. In New Guinea on rocks by river at 100 m. Note. The New Guinea specimens (KANE- HIRA & HATUSIMA 12837) have lamina 30 cm long with 50 pairs of pinnae, largest pinnae 2.0 cm long, 0.4-0.5cm wide above base, basal pinnae 3 mm long; a few long hairs are present on lower surface of costae. 2. Sphaerostephanos sudesticus nov. Pinnae_ redactae 54cm longus; DCM Dast HOLTTUM, sp. verisimiliter nullae; lamina pinnae 15-jugatae, maximae 17 Xx late cuneatae, 3/5 costam versus . 151. S. albosetosus 152. S. wauensis lobatae, utrinque pilis minutis erectis glandulisque praeditae; venae 9-jugatae, basales solum anas- tomosantes; sori mediales, inferiores praesertim elongati, indusiis tenuibus breviter hirsutis et glandulosis tecti; sporangia setifera.— Type: BRASS 27772, Sudest Island, Louisiade Archipelago, terrestrial in openings in forest, alt. 2m (L). Caudex not known; stipe 47cm long, minutely hairy. Reduced pinnae not seen; if present, minute and few. Lamina 54cm long; pinnae 15 pairs, alternate; 3-4 pairs upper pinnae adnate to rachis with asymmetric bases; lowest pinnae 2.5cm wide. Rachis minutely hairy on lower surface, hairs on upper surface pale, less than 0.5 mm long. Middle pinnae to 17X2.2cm, base _ broadly cuneate, aerophores not swollen, apex with cauda 2cm long, edges lobed to 3.5-4mm from costa (less than 2/3), lobes slightly falcate; costules 5mm apart, at 60°; veins to 9 pairs, | pair anas- tomosing, next acroscopic vein to sinus-mem- brane; hairs on lower surface of costae 0.2mm long, slightly antrorse, a little longer distally, similar shorter hairs more sparse on costules, veins and between’ veins, glands present throughout; hairs on upper surface of costae 0.2 mm long, scattered hairs 0.5 mm long on cos- tules and veins, very short erect hairs and glands between veins. Sori medial, almost all elongate, lower ones longest; indusia very thin, rather large, bearing sparse short hairs and several glands; sporangia bearing I-2 short setae. Distr. Malesia: Papua New Guinea (Sudest I.), only known from the type. Ecol. Terrestrial in openings in forest at 2 m. 3. Sphaerostephanos polycarpus (BL.) COPEL. Un. Cal. Publ. Bot. 16 (1929) 60; Fern FI. Philip. (1960) 379. — Aspidium polycarpon BL. Enum. PI. Jav. (1828) 156.—Didymochlaena polycarpa (BL.) BAK. Syn. Fil. (1867) 248; RacisB. Fl. Btzg 1 (1898) 197.—Nephrodium polycarpum (BL.) 1981] KEYS. Pol. Cyath. Herb. Bung. (1873) vii; BEDD. Handb. Suppl. (1892) 74.— Mesochlaena_ poly- carpa (BL.) BEDD. Ferns Br. Ind. Suppl. (1876) 13; Handb. (1883) 199; v.A.v.R. Handb. (1908) 232.— Dryopteris polycarpa (BL.) CHRIST, Philip. J. Sci. 2 (1907) Bot. 202.— Cyclosorus polycarpus (BL.) HOLTTUM, Rev. FI. Malaya 2 (1955) 283.—Thelypteris polycarpa (BL.) K. TWAT. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 32.— Type: BLUME, Noesa Kambangan, Java (L). Aspidium heterodon BL. Enum. PI. Jav. (1828) 157. — Type: from Celebes, no collector cited (L). S. asplenioides J. SM. in Hook. Gen. Fil. (1839) 9245 KUNZE, Farnkr. 1 (1840) t. 11, 12.— Mesochlaena asplenioides J. SM. in Hook. J. Bot. 3 (1840) 18.— Stegnogramma asplenioides (J. SM.) FEE, Gen. Fil. (1852) 204.— Type: origin unrecorded (BM in Herb. J. Sm.). Mesochlaena javanica R. BR. ex METT. Fil. Hort. Lips. (1856) 96, t. 18 f. 13.— Aspidium jJavanicum MeEtTT. Farngatt. IV (1858) 103.— Nephrodium javanicum (METT.) Hook. Fil. Exot. (1859) t. 62. — Type: “Java” (not seen). Lastrea microchlamys DE VRIESE, Tijdschr. Wisk- Nat. Wet. Amst. | (1848) 155. — Type: Cult. Hort. Bot. Lugd.-Bat. (L). Nephrodium microchlamys BAK. J. Linn. Soc. Bot. 15 (1876) 107.—Type: MOSELEY, Chal- lenger Exp. Kei Isl. (K). Aspidium perakense BEDD. J. Bot. 26 (1888) 4.— Nephrodium perakense (BEDD.) BAK. Ann. Bot. 5 (1891) 319; BEDD. Handb. Suppl. (1892) 80. — Thelypteris perakensis (BEDD.) REED, Phy- tologia 17 (1968) 303.—Type: J. DAy, Perak, Birch’s Hill, on exposed rocks (K; E). Mesochlaena toppingii COPEL. Philip. J. Sci. 12 (1917) Bot. 57; C. CHR. Gard. Bull. Str. Settl. 7 (1934) 254. — S. toppingii (COPEL.) C. CHR. Ind. Fil. Suppl. III (1934) 172. — Type: TOPPING 1902, Mt Kinabalu (Herb. Am. Fern Soc.). S. unijugus COPEL. Philip. J. Sci. 60 (1936) 109, t. 15. — Type: BRASS 2692, Solomon Is., San Cris- tobal (UC; BRI, MICH). — Fig. 12a—. Caudex erect; stipe c. 10cm long, bearing many narrow scales 10mm long. Reduced pinnae 20 pairs or more, 2.5-3.5cm apart, lowest 1 cm long, uppermost 2.5-3cm, spreading, triangular with broad base dilated almost equally both sides and acute apex, edges of larger ones incised. Lamina above reduced pinnae 100-150 cm long; pinnae many pairs, rather close; basal pinnae not narrowed at base, aerophores slightly swollen. Largest pinnae commonly 25x 1.8cm, largest seen 35*2.5cm; base rather asymmetric, basal acroscopic lobe often elongate; edges lobed more than > way to costa; lobes slightly oblique, slightly falcate; costules commonly to 4mm apart; veins 12-15 pairs, basal pair anastomosing with excur- rent vein to sinus, I-2 pairs passing to sides of sinus-membrane (in smaller fronds sometimes THELYPTERIDACEAE (Holttum) 449 only acroscopic vein); lower surfaces bearing glands throughout, acicular hairs on rachis, costae and costules normally short, spreading, in some specimens to | mm long; acicular hairs on upper surface of costae to 1mm long, scattered similar hairs on costules and veins, between veins a vari- able number of glands and very short erect aci- cular hairs. Sori medial, elongate, commonly Imm long; indusia with many yellow glands; sporangia also bearing glands; spores with a moderate number of wings of varying size. Distr. Peninsular Thailand; throughout Malesia except Philippines north of 10° and Lesser Sunda Islands east of Lombok; Solomon Islands, Samoa. Ecol. In open not too dry places, in low country and to 1500 m. Notes. Quite small plants may be fertile; in these sometimes the second veins may both reach the margin above base of sinus. Specimens with long hairs on the lower surface do not differ from the normal condition in other ways, and length of hairs varies; there is no evident correlation of long hairs with drier habitats, but this is possible. Aspidium perakense BEDD. was based on young sterile plants growing on a rock; they are very hairy. DAy collected with them fragments of a fertile frond of an abnormal Christeila; this is the source of BEDDOME’S description of sori. Mesochlaena toppingii COPEL. was _ based on a small hairy specimen. S. unijugus was also based on a small specimen in which at most the second acroscopic vein passes to the sinus-mem- brane; the upper surface was almost or quite glandless but similar specimens from New Britain have many glands. Another specimen from the Solomon Islands, otherwise similar, has a fair num- ber of glands on the upper surface. 4. Sphaerostephanos eminens (BAK.) HOLTTUM, comb. nov. — Nephrodium eminens BAK. J. Bot. 18 (1880) 213.— Dryopteris eminens (BAK.) C. Cur. Ind. Fil. (1905) 263; v.A.v.R. Handb. (1908) 214.— Type: BECCARI 455, Sumatra, G. Sing- galang 1700 m (FI; K). Dryopteris pilososquamata_ var. obtusata v.A.v.R. Bull. Jard. Bot. Btzg II, 16 (1914) 12; Handb. Suppl. (1917) 182. — Type: MATTHEW 645, G. Singgalang (BO). Mesochlaena talamauensis v.A.v.R. Bull. Jard. Bot. Btzg II, 28 (1918) 34.—S. talamauensis (v.A.v.R.) C. CHR. Ind. Fil. Suppl. II (1934) 172. — Type: BUNNEMEIJER 873, Sumatra, Talamau, 1750 m (BO). Caudex erect or suberect. Stipe c. 50cm long, glabrous; base of stipe to first large pinna 110 cm, this part bearing 7 pairs widely-spaced reduced pinnae, upper ones broadly triangular, 2.2 cm long and wide, lowest hardly |cm long. Lamina to 90 cm long, pinnae 16 pairs, lowest almost sessile and slightly narrowed at base, middle ones with stalks 3-12 mm long; apex of frond almost pinna- 450 FLORA MALESIANA [ser. II, vol. 1° like; aerophores not swollen. Largest pinnae 30 x 3cm, base cuneate, apex long-acuminate, lobed about 2 way to costa, lobes distinctly falcate, blunt, entire; costules 7 mm apart at 60°; veins to 12 pairs, concolorous and prominent on lower surface but not above, basal 1-15 pairs anas- tomosing, 2 pairs to sides of sinus-membrane; pale curved hairs 0.2-0.5 mm long on lower surface of rachis and costae, shorter and somewhat antrorse on costules, glands (no hairs) on veins and surface between veins; hairs 0.5 mm long on upper surface of rachis and costae, much shorter on costules, glands present between veins. Sori medial, lower ones somewhat divergent, all except distal ones + elongate; indusia and sporangia bearing glands. Distr. Malesia: Middle to northern Sumatra, several collections. Notes. The length of pinna-stalks varies in different specimens; those of the type of M. talamauensis only 3mm, on BECCARIs type 12mm, on a specimen of MATTHEW’s from the same locality 5mm. YATES 699 from Brastagi is rather intermediate between this species and S. penniger, with sessile pinnae and _ setiferous sporangia. 5. Sphaerostephanos williamsii (COPEL.) HOLT- TUM, Kalikasan 4 (1975) 53.— Dryopteris willi- amsii COPEL. Brittonia 1 (1931) 67, pl. 1.— Thelypteris williamsii CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 255. — Lastrea williamsii COPEL. Gen. Fil. (1947) 140; Fern FI. Philip. (1960) 322.— Type: R. S. WILLIAMS 2216, Min- danao, San Ramon (UC; B, G, NY, US). Caudex short, erect. Stipe 10cm long, very- short-hairy, basal scales 2 mm long, ovate; base of stipe to first normal pinna 20-30mm. Reduced pinnae 3 pairs, not opposite, irregularly spaced, lowest 4mm long; transition to normal pinnae abrupt or with one pair intermediate. Lamina of type 30cm long (of NY isotype 40cm); pinnae 12-14 pairs, 1-2 lower ones narrowed at base; aerophores not enlarged. Largest pinnae of type 6.5 xX 1.lcm (of NY isotype 11 x 1.8cm) lobed to less than 1 mm from costa, apex acuminate; lobes oblique, slightly falcate; costules 2.5-3.5 mm apart, at 60°; veins 8-10 pairs, both basal veins ending above base of sinus; acicular hairs on lower surface of rachis and costae short, spread- ing; whole lower surface of pinna covered with short slender erect hairs and glands; upper surface minutely hairy and glandular throughout. Sori medial, exindusiate; sporangia bearing many small yellow glands, less often 2-3 short setae; spores with thin translucent wing and cross wings. Distr. Malesia: Philippines (Mindanao, at 150 m), only known from the type. 6. Sphaerostephanos stenodontus (COPEL.) HOLTTUM, Kalikasan 4 (1975) 56. — Cyclosorus stenodontus COPEL. Philip. J. Sci. 81 (1952) 28, t. 20; Fern Fl. Philip. (1960) 344. — Thelypteris stenodonta REED, Phytologia 17 (1968) 315.— Type: RAMOS & EDANO BS 30849, Panay (US). Caudex not known; base of stipe lacking. Reduced pinnae at least 5 pairs, lowest 3-4mm long; transition to normal pinnae subabrupt. Lamina 50cm long; pinnae 30 pairs, lower ones not narrowed at base. Largest pinnae 10cm long, 15cm wide at dilated base, short-acuminate, lobed 3/5 towards costa, lobes slightly oblique, not falcate; costules 2.5—3 mm apart, at more than 60°; veins 8-9 pairs, | pair anastomosing, next acros- copic vein to short sinus-membrane; hairs more than | mm long on lower surface of rachis, fewer with shorter ones on costae and costules, glands all over surface; scattered long hairs on upper surface of costules and veins, short slender ap- pressed hairs and a few glands between veins. Sori medial, lacking indusia, basal ones not divergent; sporangia bearing glands. Distr. Malesia: Philippines (Panay), only known from the type. 7. Sphaerostephanos flavoviridis HOLTTUM, sp. nov. Pinnae redactae 2-3-jugatae, superiores 1-2.cm longae; lamina 120cm longa; pinnae inferiores basi angustatae; pinnae maximae 21 X2.5 cm, profunde lobatae; venae 12-—14-jugatae, infimae anastomosantes; costae subtus pilis minutis praeditae; pagina inter venas utrinque copiose glandulosa; sori mediales, exindusiati; stipites sporangiorum pilis glanduliferis praediti. — Type: JERMY 3694, N.E. New Guinea, Butemu, Morobe Distr. on steep bank in open place (BM). Caudex not known. Stipe 30cm long, glabrous. Reduced pinnae variable, 2-3 pairs on 13cm length of rachis, lowest very small, uppermost 1-2 cm long. Lamina 120 cm long, somewhat yel- low-green when dry; basal pinnae much narrowed at base, next two pairs less so; aerophores not elongate. Middle pinnae to 21x2.5cm, almost sessile, base broadly cuneate to subtruncate, apex with cauda 3cm long, lobed to 2.5-3mm from costa, lobes hardly falcate, oblong; costules 5 mm apart, at more than 60°; veins 12-14 pairs, basal pair anastomosing with short excurrent vein to sinus, next pair to sides of short sinus-membrane; hairs on lower surface of rachis and costae minute, on costules 0.2 mm long, longer on sinus- membrane and edge, glands throughout lower surface; long antrorse hairs on upper surface of costae, short on costules and veins, glands and minute hairs between veins. Sori medial, basal ones divergent; no indusia seen; sporangia lack- ing glands or setae on body, hairs on stalks with glandular end-cell; spores with many small wings. Distr. Malesia: Papua New Guinea. Known only from type and another specimen from same locality. 1981] THELYPTERIDACEAE (Holttum) 451 8. Sphaerostephanos trimetralis HOLTTUM, sp. nov. Caudex erectus, brevis; stipes 10cm longus, e basi stipitis usque pinna normalis infima 150 cm; pinnae redactae 30-jugatae, late triangulares; lamina 150cm longa; pinnae maximae 14x 1.3. cm, crenatae; venae 6—7-jugatae; rachis cos- taeque subtus pilis 0.2mm _ longis adpressis praeditae; pagina inter venas utrinque glandulosa; sori mediales, indusia pilis brevibus et glandulis praedita; sporangia glandulifera. — Type: HOOGLAND 3391, Papua, Lake Koena (CANB; Is, IL) Caudex short, erect. Stipe 10cm long, minutely hairy, basal scales 7mm long, narrow; base of stipe to lowest normal pinna 150cm or more. Reduced pinnae c. 30 pairs, sub-opposite, 4-6 cm apart, broadly deltoid with symmetric base, upper ones 2X1.8cm, crenate, apex acute. Lamina 150cm long; pinnae subcoriaceous, drying light red-brown; aerophores not enlarged. Largest pinnae 14x 1.3 cm, base truncate, apex acuminate, edges crenate to depth of 1mm or little more; costules 3mm apart, at more than 60°; veins 6-7 pairs, 23 pairs anastomosing to form a Zig-zag excurrent vein, | pair to short sinus-membrane; lower surface of rachis, costae and costules covered with pale antrorse hairs 0.2mm long, a few short hairs and many glands between veins; hairs on upper surface of costae 0.5mm long, between veins short erect hairs and glands. Sori medial; indusia thin, bearing short hairs and glands; sporangia with glands; spores minutely papillose. Distr. Malesia: Papua New Guinea. known from the type. Ecol. Near shore of lake, in marshy vegetation of tall grasses and sedges, alt. 10 m. Only 9. Sphaerostephanos alatellus (CHRIST) HOLT- TUM, comb. nov.—Nephrodium alatellum CHRIST in K. Schum. & Laut. FI. Schutzgeb. (1900) 112.— Aspidium alatellum (CHRIST) CHRIST, Bull. Herb. Boiss. II, 1 (1901) 454.— Dryopteris alatella (CHRIST) C. CHR. Ind. Fil. (1905) 251; v.A.v.R. Handb. (1908) 210. — Cyclo- sorus alatellus (CHRIST) COPEL. Gen. Fil. (1947) 142; Philip. J. Sci. 78 (1951) 445, excl. syn. Dry- opteris stenobasis C. CHR., D. logavensis ROSENST. & D. pseudostenobasis COPEL. — Type: LAUTERBACH 2354, N.E. New Guinea, 600 m(B). Cyclosorus heterocarpus sensu HOLTTUM & Roy, Blumea 13 (1965) 134. — Fig. 13a—c. Caudex short, erect. Stipe 15cm long, basal scales thin, to 7 X 2 mm; base of stipe to first large pinna 40-60 cm. Reduced pinnae 4-6 pairs, very small. Lamina dimorphous, to 60 cm long; pinnae 20 pairs or more, texture thin, aerophores not elongate, basal pinnae somewhat narrowed at base. Largest sterile pinnae 20 x 2.6cm, fertile to 14x 1.4cm; base unequally broadly cuneate, apex short-acuminate, edges lobed 2/3-3/4 to costa, lobes slightly falcate with rounded tips; costules of sterile fronds 4.5-Smm apart, of fertile 3.5- 4.5 mm; veins 8-9 pairs (sterile) 6-8 pairs (fertile), basal pair spreading at a wide angle to costule and anastomosing with a short excurrent vein or sometimes meeting just below sinus; next veins both to edge; lower surface quite hairless or with some very short hairs on costa only, glands abundant; hairs on upper surface of costae short, very short and sparse on costules, between veins many glands. Sori medial; indusia small, glan- dular; sporangia bearing glands. Distr. Malesia: Widely in Eastern Guinea, in forest at 400-1300 m. Notes. CHRIST described only a fertile frond and noticed neither glands nor reduced pinnae. COPELAND confused this species with Am- phineuron attenuatum (O. KTZE) HOLTTUM in the Philippines (Fern FI. Philip. 341) and with Dryopteris logavensis ROSENST. (Amphineuron pseudostenobasis (COPEL.) HOLTTUM) in New Guinea. HOLTTUM and Roy misidentified a cul- tivated plant as Cyclosorus heterocarpus and reported a chromosome count 2n = 72. New 10. Sphaerostephanos HOLTTUM, sp. nov. Caudex erectus, 30-40cm altus; stipes usque 30cm longus; pinnae redactae c. 6-8-jugatae, superiores 1.5-2.5 cm longae, integrae, aerophorae vix 1mm longae; lamina 150 cm longa; pinnae maximae 25 X2.cm vel majores, usque 1/3 costam versus lobatae; venae 9-jugatae, 1-—13-jugatae anastomosantes; pagina utrinque inter venas glandulosa; sori inframediales, inferiores leviter divergentes et leviter elongati; indusia glabra, glandulosa; stipites sporangiorum pilis glanduli- feris praediti.— Type: BRASS 23557, Papua, Milne Bay District, Mt Dayman (BM; A, L, LAE). Caudex erect, 30-40cm tall, 4-5cm diam. (BRASS). Stipe to 30cm long, base covered with broad thin scales 10-15 mm long; base of stipe to first large pinna 70cm or more. Reduced pinnae 6-8 pairs, uppermost 1.5—2.5 cm long, broadly tri- angular, lowest less than 1mm long. Lamina 150cm long; lower 3-4 pairs normal pinnae gradually narrowed and more shallowly lobed towards base; aerophores hardly | mm long. Lar- gest pinnae of type 25x 2.0cm (of BRASS 23226 36x2.4cm), base subtruncate, apex caudate- acuminate (cauda 34cm), lobed 1/3-2/5 to costa, lobes slightly falcate; costules 4.5mm apart (to 6 mm in BRASS 23226) at more than 60°; veins to 9 pairs, 1-1; pairs anastomosing, 13-2 pairs to sinus- membrane; all lower surfaces glandular throughout, no acicular hairs; hairs on upper sur- face of costa 0.5 mm long, very short on costules, glands on all parts of surface. Sori inframedial, lower ones slightly divergent and slightly elongate; daymanianus 452 indusia thin, bearing glands only; sporangia lack- ing glands on body, hairs on stalks with terminal gland which is red in young sori. Distr. Malesia: Eastern New Guinea (Milne Bay and Morobe Districts), Bougainville. Ecol. In forest at 700-1500 m. Notes. PULLEN 7964 (Milne Bay Distr.) has pinnae to 2.6cm wide, lobed a little more than , with veins to 13 pairs. WOMERSLEY & MILLAR 8373 (Morobe Distr.) is reported to have had a caudex 150 cm tall; its sori are young and not elongate. CRAVEN & SCHODDE 277 from Bougainville had a caudex 60 cm tall, base of stipe to first large pinna 95 cm, fronds in all 3.25 m long; my record is that it has glands on body of sporangia, which needs checking. 11. Sphaerostephanos appendiculatus (BL.) HOLTTUM, Kalikasan 4 (1975) 62.—Gym- nogramme appendiculata BL. Enum. Pl. Jav (1828) addendum to p. 112; Fl. Jav. Fil. (1828) 92, t. 39. — Goniopteris appendiculata (BL.) PRESL, Tent. Pterid. (1836) t. 7, f. 11.— Aspidium pilosiusculum METT. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 233, nom. nov.— Nephrodium ap- pendiculatum (BL.) RAciIB. FI. Btzg 1 (1898) 186.— Dryopteris appendiculata (BL.) C. CHR. Ind. Fil. (1905) 252: v.A.v.R. Handb. (1908) 213; BACKER & POSTH. Varenfl. Java (1939) 52, f. 9.— Thelypteris appendiculata (BL.) REED, Phy- tologia 17 (1968) 260.— Type: BLUME, Java, Boerangrang (L). Dryopteris perakensis var. sumatrensis v.A.v.R. Bull. Jard. Bot. Btzg II, 11 (1913) 13.— Type: MATTHEW 516, Sumatra, Padang Pandjang (BO; K). Dryopteris brunnescens C. CHR. Bot. Jahrb. 66 (1933) 44. — Thelypteris brunnescens REED, Phy- tologia 17 (1968) 265.— Type: KJELLBERG 3534, S. W. Celebes, Rante Lemo (S-PA; BO). Caudex short-creeping. Stipe short, densely short-hairy; base of stipe to first large pinna 15- 40 cm. Reduced pinnae 6-12(-20) pairs, somewhat deflexed, lowest 3-S mm long, upper ones | cm long, broadly triangular with asymmetric base. Lamina 20-45(-70) cm long, reddish when young; apical lamina deeply lobed with gradual transition to upper pinnae, several pairs of which are adnate; free pinnae 10-20 pairs, oblique with upturned tips. Largest pinnae commonly to 8.5 x 1.5 cm (to 17 x 2.2 cm); base truncate and distinctly auricled on acroscopic side (bases of upper ones asym- metric), apex short-acuminate, lobed + way to costa (largest ones more than 3); lobes oblique with falcate obtusely angled tips; costules 3.5 mm apart in small frond, to 5 mm in large ones; veins 6-7(-9) pairs, one pair anastomosing, next pair to sinus-membrane; lower surface of costae and rachis bearing short and long spreading hairs, sparse on costules and veins, very short hairs and FLORA MALESIANA [ser. II, vol. 1° glands between veins; short hairs on upper surface of costae, very short erect hairs abundant between veins, usually some glands also. Sori medial, lower ones not divergent; indusia small with glands and short hairs; sporangia with several setae (to 10), sometimes also a gland. Distr. Malesia: Sumatra, Java, Lesser Sunda Is. (Bali, Flores), SW. & N. Celebes, Philippines (Luzon). Ecol. On rocks by streams or waterfalls at 400-1200 m. Notes. D. brunnescens was based on a small plant with pinnae to 4.5cm long. PRICE reports red young fronds on plants in Luzon but I have seen no such report from Java. 12. Sphaerostephanos pseudomegaphyllus (v.A.v.R.) HOLTTUM, comb. nov. — Dryopteris megaphylloides v.A.v.R. Bull. Jard. Bot. Btzg II, 20 (1915) 16, non ROSENST. 1913.— Dryopteris pseudomegaphylla v.A.v.R. Handb. Suppl. (1917) 180. — Type: BURCK, Sumatra, Padang Pandjang (BO). Aspidium megaphyllum var. hirsutum METT. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 233; MIQUEL, ibid. 4 (1869) 159. — Type: KORTHALS, Sumatra (lz): Differs from S. penniger: fronds smaller; pinnae 7x1 to 16x 2.2cm, lobed about 1/4-1/3 to costa; veins 8-10 pairs, 13-2 pairs anastomosing, 13 pairs to sides of sinus-membrane; dense spreading hairs to 0.5mm long on lower surface of rachis and costae; indusia densely hairy or with many glands and fewer hairs; sporangia copiously setiferous. Distr. Malesia: Sumatra (Padang Highlands). 13. Sphaerostephanos diversilobus (PRESL) HOLTTUM, Kalikasan 4 (1975) 57. — Nephrodium diversilobum PRESL, Epim. Bot. (1851) 47; HOLTTUM, Novit. Bot. Univ. Carol. Prag. 1968 (1969) 39.—Dryopteris diversiloba (PRESL) CHRIST, Philip. J. Sci. 2 (1907) Bot. 199, nomen tantum; COPEL. Philip. J. Sci. 40 (1929) 297. — Phegopteris diversiloba (PRESL) v.A.v.R. Handb. (1908) 503.—Cyclosorus diversilobus (PRESL) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 244; COPEL. Fern Fl. Philip. (1960) 357.— Thelypteris diversiloba (PRESL) REED, Phytolo- gia 17 (1968) 273. — Lectotype (HOLTTUM 1969): CUMING 102, Luzon (PRC; E, FI-W, G, K, LE). Goniopteris asymmetrica FEE, Gen. Fil. (1852) 253.— Thelypteris asymmetrica (FEE) REED, Phytologia 17 (1968) 261.—Specimens cited: CUMING 51, 102; FEE’s specimens of these num- bers not seen. Dryopteris acromanes CHRIST, Philip. J. Sci. 2 (1907) Bot. 200.— Phegopteris acromanes (CHRIST) v.A.v.R. Handb. (1908) 505. — Type not selected; specimens cited (which include one of CUMING 51) not seen. Caudex short, creeping. Stipe of sterile frond to 1981] 20 cm, of fertile to 30cm, sparsely hairy. Reduced pinnae 2 pairs, very small, seen in isotype at E only. Lamina 25 cm long; pinnae 7-9 pairs; basal pinnae gradually narrowed at base, basal basis- copic lobe missing. Fertile pinnae to 6.5 x 2.0cm, sterile to 8cm long, 3cm wide near apex; lower pinnae and sometimes others widened and more deeply lobed distally, some with irregularly elon- gate subapical lobes. Middle pinnae with = dilated base, abruptly short-pointed apex, lobed 2/3 to costa apart from widened apical part, lobes hardly falcate, normally entire; costules 4-5 mm apart; veins 7-8 pairs except in elongate distal lobes. | pair anastomosing, next acroscopic vein to sinus- membrane; lower surface of rachis and costae bearing short erect acicular hairs, also some longer ones, especially on sterile pinnae; short erect hairs between veins on both surfaces, glands few on both surfaces, those on lower surface usually confined to costules and veins. Sori medial; indusia thin, densely hairy with a few glands; sporangia with glands; spores with trans- lucent wing and cross-wings. Distr. Malesia: Philippines (Luzon, Panay), in forest at 300-1000 m. Notes. Some pinnae are irregularly lobed on most specimens; this suggests hybridization. One sterile frond of CUMING 51 at Kew has very long distal lobes which are crenate with pieocnemioid venation. The only specimen seen with reduced basal pinnae is one of CUMING 102 at Edinburgh. The specimens listed by CHRIST in 1907 under D. diversiloba represent several other species; he re-described PRESL’s species as D. acromanes from CUMING S51 and other specimens now ap- parently lost. 14. Sphaerostephanos’ dichrotrichus (COPEL.) HOLTTUM, comb. nov. — Dryopteris dichrotricha COPEL. Philip. J. Sci. 6 (1911) Bot. 74. — Thelyp- teris dichrotricha (COPEL.) REED, Phytologia 17 (1968) 272. — Type: C. KING 294, Papua “main- land opposite Samarai island’ (MICH: BRI, NSW). Caudex short-creeping. Stipe 40cm to first reduced pinna, dull reddish with pale hairs 0.5 mm or more long. Lamina 65 cm or more long: pinnae to 20 pairs, basal 1-3 pairs gradually reduced, lowest 14cm long: aerophores slightly swollen. Largest pinnae 182.0cm; base truncate, apex acuminate with entire cauda to 1.5 cm long; edges lobed +3 way to costa, lobes slightly falcate with round tips; costules 4.5-Smm apart, at 60° or more; veins 7 pairs, 1-1: pairs anastomosing, next +1 pair to sinus-membrane; rather sparse hairs 1mm long and many shorter ones on lower sur- face of rachis and costae, shorter on costules and veins, glands and short erect hairs between veins: minute erect hairs and glands between veins on upper surface. Sori medial; indusia thin, with many hairs, also glands; sporangia usually with THELYPTERIDACEAE (Holttum) 453 1-2 slender setae (rarely a gland) on body, on stalk a sessile gland; spores with a thin translucent wing and cross-wings. Distr. Malesia: extreme east of New Guinea, Admiralty Islands, at low altitudes. Notes. Some specimens have more erect hairs than glands on upper surface. In one (LAE 65333) there are 2 pairs of small pinnae near base of stipe and then a gap of 20cm. 15. Sphaerostephanos (COPEL.) HOLTTUM, comb. nov.— Dryopteris vestigiata COPEL. Un. Cal. Publ. Bot. 18 (1942) 220.— Cyclosorus vestigiatus (COPEL.) COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78 (1951) 446, pl. 27.— Thelypteris vestigiata (COPEL.) REED, Phytologia 17 (1968) 323.— Type: BRAsSs 12317, W. New Guinea, Idenburg River, 1700 m, in forest (MICH; BM, L). Caudex slender, erect, to 60cm tall. Stipe to 50cm long, glandular, hairless. Reduced pinnae 3 pairs, all very small, Scm apart. Lamina more than 100cm long; pinnae 40 pairs, not dimor- phous, basal pinnae somewhat narrowed at base, with stalks to 1 mm long, aerophores not swollen. Largest pinnae 21 x 2.2 cm, base broadly cuneate, apex caudate-acuminate (cauda 2.5-5.0 cm suben- tire) lobed to 2-2.5mm from costa, lobes hardly falcate with rounded tips; costules 44.5 mm apart at 60° or more; veins 9-11 pairs, 1 pair anas- tomosing, next veins both to edge: hairs on lower surface of rachis and costae 0.1 mm long (longer distally on costules), surface between veins copiously glandular; upper surface of costules and veins with scattered hairs like those on costae, between veins minute erect hairs and glands. Sori medial or little supramedial: indusia small, thin, with glands only; sporangia bearing glands on body and gland-tipped hairs on stalk; spores with many small wings. Distr. Malesia: Several collections from east- ern New Guinea at c. 1500 m. Note. Some specimens have quite hairless lower surface of rachis and base of costae. vestigiatus 16. Sphaerostephanos morotaiensis HOLTTUM, sp. nov. Caudex brevis, repens: stipes Scm longus; pinnae redactae usque 12-jugatae, patentes, superiores 1.5-2 cm longae: lamina 50cm longa: pinnae 16-jugatae, maximae 7.51.5 cm, subin- tegrae; venae 4-5-jugatae: pagina inter venas utrinque pilis minutis erectis glandulisque praedita; sori mediales; indusia brevi-pilosa; sporangia glandulifera. — Type: MAIN & ADEN 758, Morotai, Totodahu, 30m (K; BO, L, SING). Caudex short-creeping. Stipe S5cm_ long, minutely hairy; base of stipe to first normal pinnae 50cm. Reduced pinnae 1.5-2.5cm apart, to 12 pairs; upper ones 1.5-2cm long, spreading, broadly triangular, entire: lowest 4mm _ long. 454 Lamina 50cm long; pinnae 16 pairs, lower ones not narrowed at base nor auricled; aerophores not elongate. Largest pinnae 7.5 x 1.5cm; base trun- cate, sometimes a little dilated; apex short-acu- minate; edges subentire; costules 3—3.5 mm apart on fertile pinnae, 4.5mm on sterile; veins 4—5 pairs, 3-3; pairs anastomosing, sinus-membrane hardly detectable; whole lower surface (including rachis) covered with very short erect hairs and glands; hairs on upper surface of costae 0.3- 0.4 mm long, on costules and veins sparse, surface between veins with very short erect hairs and glands. Sori medial; indusia small with many short hairs; sporangia bearing glands. Distr. Malesia: Moluccas. Apart from the type and other specimens from Morotai, I refer here SA-AANAM 49 from Obi Island (L) which has larger crenate pinnae (to 18 x 3.0 cm), agreeing in other respects. 17. Sphaerostephanos kalkmanii HOLTTUM, sp. nov. Caudex erectus; pinnae redactae 4-jugatae, omnes minutae; pinnae normales 18-jugatae, usque 3.5X1.0cm, crenatae, costae costulaeque subtus pilis minutis appressis glandulisque praeditae; indusia parva vel nulla; sporangia nec glandulis nec setis instructa. —Type: KALKMAN 4070, W. New Guinea, Orion Mts, Tenma River, 1500 m (BM; LAE, L, SING). Caudex erect, to at least 15cm tall. Stipe 4- 5 cm long, dark, sparsely short-hairy, basal scales c. 4X 1mm; base of stipe to first large pinna 10-15 cm. Reduced pinnae to 4 pairs, all about 1mm long. Lamina 20cm long; pinnae 18 pairs, subcoriaceous, lower ones not narrowed at base; aerophores slightly swollen. Hairs on lower sur- face of rachis 0.3mm long, brown, on upper sur- face a little longer. Largest pinnae 2.6-3.5cm long, 0.7-1.0 cm wide, fertile ones sometimes, not always, smaller than sterile; base subtruncate to broadly cuneate; apex rather abruptly short-poin- ted; edges crenate, a little more deeply in fertile than in sterile pinnae; costules 2mm apart; veins all very oblique, 4-5 pairs, 13 pairs anastomosing, next vein to short sinus-membrane; lower surface of costae and costules covered with minute ap- pressed hairs, also glands, a few glands sometimes present on surface between veins; hairs on upper surface of costae to 0.5 mm long, on costules and veins shorter, sparse, a few glands sometimes present (?) on surface between veins. Sori medial, sometimes covering whole surface at maturity; indusia very small with short hairs and glands, sometimes lacking; sporangia bearing neither glands nor setae; spores minutely spinulose. Distr. Malesia: West New Guinea, 1500 m. Only known from type. Ecol. “In abandoned garden, rather common”. Note. Possibly nearest to the lowland species S. lamii, which has glands between veins on both FLORA MALESIANA [ser. II, volaale surfaces. S. lamii has longer pinnae, upper reduced pinnae 10mm long, and veins not very oblique. 18. Sphaerostephanos perglanduliferus (v.A.v.R.) HOLTTUM, Kalikasan 4 (1975) 59.— Dryopteris perglandulifera v.A.v.R. Bull. Jard. Bot. Btzg III, 2 (1920) 150.—Thelypteris perglandulifera (v.A.v.R.) REED, Phytologia 17 (1968) 303.— Type: BROOKS 281, Sumatra, Lebong Tandai, Benkoelen (BO). D. perglandulifera var. firmior v.A.v.R. l.c.— Type: DOCTERS VAN LEEUWEN-REYNVAAN 3284, Sumatra, Haboko (BO). Phegopteris pseudoarfakiana HOSOKAWA, Tr. Nat. Hist. Soc. Formosa 28 (1938) 147. — Menis- cium pseudoarfakianum (HOSOKAWA) HOSOK- AWA, ibid. 32 (1942) 286.— Type: HOSOKAWA, Caroline Is., Palau (TAI). Dryopteris arbuscula sensu KJELLBERG & C. CHR. Bot. Jahrb. 66 (1933) 43. Caudex massive, short, erect; stipe 5-15cm long, densely short-hairy; basal scales 10 mm long, narrow; base of stipe to first normal pinna 20- 50cm; reduced pinnae to 10 pairs, lowest 7 x 7mm, upper ones to 1.51.5 cm, base broadly cuneate, symmetric, apex obtusely pointed. Lamina of type 75cm long; transition at base to reduced pinnae subabrupt; lowest normal pinnae not auricled at base; aerophores not elongate. Largest pinnae of type 14x1.7cm (on small plants, fertile, 71cm), base broadly cuneate, apex short-acuminate, edges shallowly crenate; costules 3-4 mm apart, at c. 45° to costa; veins 6—7 pairs, 3 pairs anastomosing to form a slightly Zig-zag excurrent vein, | pair to sides of sinus- membrane; lower surface of rachis covered with slender appressed hairs less than 0.5mm _ long, hairs on costae minute, glands throughout lower surface; hairs on upper surface of rachis pale, 1mm long, on costae less than 0.5 mm, copious glands between veins. Sori medial; indusium bearing glands; sporangia with short setae and glands, a gland at end of short hair on sporangium- stalk. Distr. Malesia: S. Sumatra, Borneo, S.W. Cele- bes, Philippines (Negros) and Micronesia (Palau). Ecol. In rocky stream-beds at low to medium altitudes. Note. This is closely related to S. penniger, but appears to be distinct in its erect caudex (seen on several specimens) and narrow crenate pinnae; the stream-bed habitat is recorded for several specimens. 19. Sphaerostephanos veitchii HOLTTUM, AIl- lertonia 1 (1977) 204.—Type: J. G. VEITCH, Solomon Islands, San Cristobal (K). Nephrodium amboinense var. subglandulosum BAKER, J. Linn. Soc. Bot. 19 (1882) 295. — Type: R. B. COMINS, Solomon Islands (K). 1981] THELYPTERIDACEAE (Holttum) 455 Caudex short, creeping or suberect. Stipe of type lacking, of others 10-20cm, minutely hairy; base of stipe to first large pinna 35-60 cm. Reduced pinnae about 7 pairs, lowest 5 mm long, upper ones 1.5—2.5cm long and wide, broadly triangular with asymmetric base (wider on acros- copic side), crenate. Lamina 50-90 cm or more long; basal pinnae not narrowed at base; aero- phores not elongate. Largest pinnae of type 13 x 1.7cm, of others to 20 x 2cm; base broadly trun- cate to full width, sometimes a little dilated but not auricled; apex acuminate, not caudate; edges crenate to depth of 1-I.5mm; costules 4mm apart, at 60° to costa; veins 6 pairs, 23. pairs anastomosing to form a zig-zag excurrent vein, 15-2 pairs to sides of sinus-membrane; hairs on lower surface of rachis 0.4 mm long (much shorter on some plants from New Guinea), pale, curved, on costae and costules shorter, antrorse, many glands and short erect hairs between veins; upper surface of rachis covered with brown hairs more than 0.5mm long, similar hairs on costae and scattered on costules and veins, glands on surface between veins and sometimes short erect hairs. Sori medial, lower ones divergent, those on basal veins from adjacent costules sometimes confluent; indusia small, bearing short hairs and glands; sporangia bearing glands. Distr. Solomon Islands, and Malesia: Eastern New Guinea. Ecol. In the Solomons by streams in low country and to 800m; in New Guinea plants in forest at higher altitudes (to 1400 m) are smaller but otherwise not distinct. 20. Sphaerostephanos lamii HOLTTUM, sp. nov. Caudex brevis, repens vel suberectus; stipes 20cm longus; pinnae redactae 3-5-jugatae, superiores 10mm longae; lamina 30cm longa, pinnae 12-15-jugatae, dimorphae; pinnae Steriles usque 8.5X1.8cm, crenatae, pagina utrinque glandulifera; pinnae fertiles usque 6.0 1.2cm; sori inframediales; indusia parva, pilis brevibus glandulisque praedita; sporangia glandulifera. — Type: H. J. LAM 748, W. New Guinea, Prauwen bivak, 10m (L; BO). Caudex short, creeping or suberect; stipe 12- 20 cm long, short-hairy in groove; base of stipe to first normal pinna 30-40 cm; reduced pinnae 3-5 pairs, not opposite, upper ones to 10mm long, deflexed with slightly auricled asymmetric base, lowest 3mm long. Lamina 30cm long; pinnae 8-15 pairs, more or less dimorphous, basal pinnae not narrowed at base, not auricled; aerophores not elongate; rachis bearing stiff brown hairs to 1 mm long on both surfaces. Sterile pinnae to 8.5 x 1.8 cm, base truncate, apex short-acuminate, edges crenate to a depth of 1mm; costules 3.5-4mm apart, at 60° to costa; veins 5-6 pairs, 23 pairs anastomosing, one pair to sides of sinus-mem- brane; on lower surface hairs at base of costae very short, distally 0.2-0.3mm (but see note below), rest of lower surface of pinnae glabrous with sessile glands; hairs on upper surface of costae hardly 0.5 mm long, rest of upper surface glabrous or with some minute hairs, glands present throughout but fewer than on lower sur- face, sometimes lacking. Fertile pinnae to 6.0 x 1.2 cm, slightly crenate; veins 3-4 pairs; sori near costules, basal ones not divergent; indusia small, with very short hairs and a few glands; sporangia bearing glands. Distr. Malesia: W. New Guinea, several col- lections from Rouffaer and Mamberamo Rivers, at 10-200 m; Admiralties (Los Negros: GRETHER & WAGNER 4003); a distinct variety in Eastern New Guinea? Note. A specimen from 820m in the Southern Highlands of Eastern New Guinea differs in the presence of hairs 1 mm long on the lower surface of costae and scattered long hairs on costules and veins on the upper surface. 21. Sphaerostephanos hirsutus (KUNZE ex METT.) HOLTTUuUM, Kalikasan 4 (1975) 63. — Nephrodium hirsutum PRESL, Epim. Bot. (1851) 48, non D. DON 1825; HOOK. Sp. Fil. 4 (1862) 70 p.p. & t. 240B. — Aspidium hirsutum KUNZE ex METT. Farngatt. IV (1858) 107, nom. nov. — Dryopteris hirsuta (KUNZE ex METT.) O. KTZE, Rev. Gen. Pl. 2 (1891) 813; v.A.v.R. Handb. (1908) 214. — Dryopteris adenophora C. CHR. Ind. Fil. (1905) 251, nom. nov. superfl. — Cyclosorus adenophorus (C. CHR.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 242; CopEL. Fern FI. Philip. (1960) 343. — Thelypteris adenophora (C. CHR.) REED, Phytologia 17 (1968) 258.— Type: CUMING 82, Luzon (PRC; K, P). KEY TO THE VARIETIES 1. Pinnae commonly to 202.2cm; sporangia glanduliferous ae . a. var. hirsutus 1. Pinnae to 15*1.4cm; sporangia setiferous b. var. celebicus a. var. hirsutus Caudex short, erect, with many branches at its base; stipe short; base of stipe to first normal pinna 30-90 cm, basal scales broad, thin, not per- sistent; reduced pinnae to 12 pairs or more, each consisting of an aerophore with a very small rounded blade, uppermost less than 1 cm long. Lamina to 120cm long; transition at base to reduced pinnae abrupt or with an intermediate pair; lowest normal pinnae somewhat narrowed at their bases and sometimes slightly auricled; aerophores to Imm _ long. Largest pinnae commonly 12-20 x 1.5-2.2 cm (to 30 x 2.5 cm, teste COPELAND), base truncate and somewhat dilated; apex caudate-acuminate (cauda 1.5-2.5 cm long), lobed 2/5-3/5 towards costa; lobes distinctly fal- 456 FLORA MALESIANA [ser. II, vol. 1° cate with rounded or obtuse-pointed tips; costules 3-4 mm apart, at 60° or more to costa; veins 7-8 pairs, | pair anastomosing, next pair to sides of sinus-membrane; lower surface of rachis and cos- tae with a variable number of long coarse hairs (sometimes only distally on costae) with very short hairs and glands, costules similar, abundant glands between veins; hairs on upper surface of rachis and costae 1.5mm or more long, scattered hairs 1mm long on costules and veins, between veins abundant appressed hairs and a variable number of glands (often few). Sori medial, lower ones not divergent; indusia small. with glands and short hairs; glands on sporangia. Distr. Malesia: throughout forest, at 400-1200 m. Note. Pubescence in var. hirsutus is variable; it is most clearly distinguished from S. todayensis (which see), also variable in pubescence, by the small size of its basal reduced pinnae. A more nearly allied species is S. trichochlamys. Philippines, in b. var. celebicus HOLTTUM, var. nov. A typo speciei differt: pinnis usque 15 x 1.4cm; pagina inferiore inter venas pilis tenuibus ad- pressis vestita; indusiis nullis vel minutis; spo- rangiis 3-4 setis ornatis.— Type: POSTHUMUS 2743, S.W. Celebes, G. Bonthain, 1500m (BO). Distr. Malesia: S. W. and Central Celebes & Lesser Sunda Is. (Flores: JAAG 1647). 22. Sphaerostephanos acrostichoides (DESV.) HOLTTUM, comb. nov.— Nephrodium acrosti- choides DESV. Mém. Soc. Linn. Paris 6 (1827) 255. (Not Thelypteris acrostichoides (MICHX) NEWL. nom. inval.). — Type: Collector unnamed, Timor (P). Polystichum riedleanum GAUD. in Freyc. Voy. Ur. Phys. Bot. (1828) 327. — Dryopteris riedleana (GAUD.) v.A.v.R. Handb. (1908) 230.— Type: GAUDICHAUD, Timor (G). Nephrodium smithianum PRESL, Epim. Bot. (1851) 46; HOLTTUM, Novit. Bot. Inst. Bot. Univ. Carol. Prag. 1968 (1969) 41 excl. syn. Aspidium productum KAULF.—S. smithianus (PRESL) HOLTTuM, Kalikasan 4 (1975) 65. — Type: CUM- ING 279, Guimaras (PRC; B, BM, E, G, K, L). Dryopteris perpilifera v.A.v.R. Bull. Jard. Bot. Btzg II, 11 (1913) 12; Handb. Suppl. (1917) 183. — Thelypteris perpilifera (v.A.v.R.) REED, Phytolo- gia 17 (1968) 303. — Type: LAUTERBACH 68, N.E. New Guinea, Finschhafen (BO; BRI, L, SING). Dryopteris angusta COPEL. Philip. J. Sci. 9 (1914) Bot. 3; v.A.v.R. Handb. Suppl. (1917) 186. — Thelypteris angusta (COPEL.) REED, Phytologia 17 (1968) 259. — Type: C. KING 408, E. New Guinea (MICH; K, NSW). Dryopteris pseudoarbuscula yv.A.v.R. Philip. J. Sci. 11 (1916) Bot. 106; Handb. Suppl. (1917) 504.— Type: C. B. ROBINSON 1962, Amboina (BOskKe ea): Dryopteris riparia COPEL. Univ. Cal. Publ. Bot. 18 (1942) 221.—Cyclosorus riparia (COPEL.) COPEL. Gen. Fil. (1947) 143; Philip. J. Sct. 78 (1951) 456, pl. 37. — Thelypteris riparia (COPEL.) REED, Phytologia 17 (1968) 309. — Type: BRASS 13048, New Guinea, Idenburg River, 800m (UC; BM, BO, L, LAE). Cyclosorus boholensis COPEL. Philip. J. Sci. 81 (1952) 31, pl. 23; Fern FI. Philip. (1960) 348. — Thelypteris boholensis (COPEL.) REED, Phy- tologia 17 (1968) 204.— Type: RAMOS BS 42990, Bohol (UC; G). Cyclosorus reederi COPEL. Amer. Fern J. 43 (1953) 12.— Thelypteris reederi (COPEL.) REED, Phytologia 17 (1968) 308. — Type: REEDER 889, N.E. New Guinea, Finsch-hafen (US). Aspidium arbuscula sensu CHRIST, Ann. Jard. Bot. Btzg 15 (1898) 135.— Cyclosorus hispidula sensu COPEL. Philip. J. Sci. 78 (1951) 455. — Cyclo- sorus productus sensu COPEL. Fern Fl. Philip. (1960) 348, p.p. Caudex erect, not or little branched at the base. Base of stipe to lowest reduced pinna 3-10 cm, to lowest large pinna 10-30cm or more; at base of lamina a subabrupt transition to reduced pinnae; number of reduced pinnae varying with size of frond, larger ones spreading, triangular with a broad base. Lamina of smaller plants fertile at 20 cm long with 15 pairs pinnae, of large plants to 50cm long. Largest fertile pinnae of small plants 3-5 x 0.7-1.0 cm, of largest plants to 10 x 1.0 cm or larger; base broadly cuneate and + dilated on both sides, apex evenly tapered, edges lobed 1/3-1/2 way to costa; costules at less than 60° to costa, 2-2;mm apart in small plants; veins 3-4 pairs in small plants, 7-8 pairs in large ones, I-12 pairs anastomosing, next acroscopic vein to sinus membrane; lower surface of rachis with copious spreading pale hairs 1 mm long and shorter ones, long hairs less abundant on costae, lower surface between veins bearing copious glands and some erect short hairs; upper surface of rachis with pale hairs |mm long, shorter hairs on costae, also scattered on costules and veins, between veins copious short erect hairs and glands. Sori medial, on small pinnae covering lower surface; indusia hairy and glandular; sporangia bearing glands. Distr. Malesia: East Java, Lesser Sunda Is. (Sumba, Timor), Central & S.E. Celebes, Philip- pines (Bohol, Mindanao, Guimaras), Moluccas (Amboina), and New Guinea incl. Admiralty Is. Ecol. Usually in rocky stream-beds at 0-500 m (to 1500 m in Mindanao and New Guinea). In New Guinea plants of all sizes have been collected, but elsewhere none as large as the larger ones in New Guinea. Types of the various basionyms differ chiefly in size, agreeing well in frond-form, dilated bases of pinnae, pubescence and the distribution of glands. If the larger New Guinea plants (which apparently do not grow in stream-beds) are judged to represent a distinct species, the name Dry- 1981] THELYPTERIDACEAE (Holttum) 457 opteris perpilifera v.A.v.R. should be _ its basionym. 23. Sphaerostephanos melanorachis HOLTTUM, sp. nov. Stipes rachisque atrobrunneus; stipes 7cm longus; pinnae redactae 10-jugatae, superiores usque 1.7X1.5cm; lamina 55cm longa; pinnae dimorphae, steriles usque 13X1.9cm, _ fertiles minores, omnes c. 1/3 costam versus lobatae; venae 8-jugatae, 2-jugatae anastomosantes ; rachis costaeque subtus pilis brevibus antrorsis vestita; pagina inter venas utrinque glandulosa; sori mediales; indusia pilis paucis praedita.— Type: A. H. G. ALSTON 16927, Moluccas, Batjan, Mt Sibela 750 m (BM). Caudex short, apparently erect. Stipe 7cm, dark brown, minutely hairy; base of stipe to first large pinna 35-40cm; rachis also very dark. Reduced pinnae 10 pairs, upper ones 1.7 x 1.5 cm, triangular with asymmetric base, crenate, slightly deflexed, lowest 4-5 mm long. Lamina 55 cm long; pinnae 25 pairs, dimorphous; aerophores not elongate. Sterile pinnae to 13 cm long, 1.9 cm wide above truncate dilated base, apex caudate-acu- minate (cauda to 1.5cm); edges lobed to a depth of 3mm (c. 1/3), lobes slightly falcate; costules 4mm apart, to 60°; veins to 8 pairs, 2 pairs anas- tomosing, forming a zig-zag excurrent vein, | pair to sinus-membrane; lower surface of rachis bear- ing rather sparse pale appressed hairs 0.5mm long, those on costa also antrorse, 0.2 mm long, sparse short appressed hairs on costules and veins; glands on surface between veins; hairs on upper surface of rachis pale, | mm long, on costae a little shorter, similar hairs scattered on costules and veins, glands between veins. Fertile pinnae to 9.5cm long, 1.6cm wide at base, 1.2cm above base; sori medial; indusia rather small, thin, with a few short hairs; sporangia rarely with a gland. Distr. Malesia: Moluccas (Batjan, Mt Sibela), 750 m. 24. Sphaerostephanos heterocarpus (BL.) HOLT- TUM in Nayar & Kaur, Comp. to Bedd. (1974) 209.— Aspidium heterocarpon BL. Enum. PI. Jav. (1828) 155; MeETT. Ann. Mus. Bot. Lugd. — Bat. 1 (1864) 233.— Nephrodium heterocarpum (BL.) MOORE, Ind. Fil. (1858) 93; RACIB. FI. Btzg 1 (1898) 187.— Dryopteris heterocarpa (BL.) O. KTZE, Rev. Gen. Pl. 2 (1891) 813; v.A.v.R. Handb. (1908) 228; C. CHR. Gard. Bull. Str. Settl. 4 (1929) 380; BACKER & POSTH. Varenfl. Java (1939) 54.—Cyclosorus heterocarpus (BL.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 180; HOLTTUM, Rev. Fl. Malaya 2 (1955) 269, f. 155; TARD. FI. Gén. I.-C. 7, pt. 2 (1941) 385; COPEL. Fern Fl. Philip. (1960) 345, p.p.— Thelypteris heterocarpa (BL.) MORTON, Amer. Fern J. 49 (1959) 113.—Type: BLUME, Java, ad pedem Boerangrang (L, n. 908, 332-1158). Polypodium longifolium ROxB. Calc. J. Nat. Hist. 4 (1844) 462.— Type from Amboina (BR). Nephrodium pubescens BRACK. in Wilkes U.S. Expl. Exp. 16 (1854) 186, non DON 1825. — Type: U.S. Expl. Exp. Luzon, near Banos (US). Dryopteris bordenii CHRIST, Philip. J. Sci. 2 (1907) Bot. 204; v.A.v.R. Handb. (1908) 822. — Cyclosorus bordenii (CHRIST) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 243; COPEL. Fern Fl. Philip. (1960) 345, p.p.— Thelypteris bordenii (CHRIST) REED, Phytologia 17 (1968) 264.— S. heterocarpus var. bordenii HOLTTUM, Kalikasan 4 (1975) 67.—Lectotype (HOLTTUM 1975): BORDEN 1237, Luzon, Mt Mariveles (US). Dryopteris suprastrigosa ROSENST. Fed. Rep. 10 (1912) 335.—Cyclosorus suprastrigosus (ROSENST.) COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78 (1951) 446. — Thelypteris suprastrigosa (ROSENST.) REED, Phytologia 17 (1968) 318. — Type: Frau BAMLER 37, N.E. New Guinea, Sat- telberg (not seen; at BM is KEYSSER 37 from Sattelberg, named by ROSENST.). Dryopteris mixta ROSENST. Fed. Rep. 12 (1913) 172. — Lastrea mixta (ROSENST.) COPEL. Philip. J. Sci. 78 (1951) 426.— Thelypteris mixta (ROSENST.) REED, Phytologia 17 (1968) 293. — Type: KEYSSER S. 141, N.E. New Guinea, Sat- telberg (S-PA). — Fig. 1q. Caudex erect, rarely very tall, usually with several branches from base. Stipe 5cm or little more. Reduced pinnae commonly 10 pairs (dis- tance apart, and number, depending on size of plant), never less than 6 pairs, uppermost com- monly 10mm long, narrow, auricled, lowest very small. Lamina variable in length; pinnae com- monly 20-30 pairs; several pairs lower pinnae much narrowed towards their bases, basal one at most slightly auricled; aerophores slightly swollen. Largest pinnae 10-20cm long, 1.2-2.5cm wide, fertile sometimes narrower than sterile and with shorter hairs, lobed 5/8-3/4 towards costa; cos- tules 3-3; mm apart; veins 6-12 pairs, basal veins anastomosing at least in basal half of pinna with excurrent vein to sinus, second acroscopic vein sometimes touching side of sinus-membrane; hairs on lower surface of costae very short, erect, with scattered longer ones usually not over 0.5mm long, more sparse similar hairs on costules and veins, copious yellow glands and a variable num- ber of short erect hairs between veins; upper surface of costae bearing antrorse pale hairs to 1 mm long with scattered similar hairs on costules and veins; surfaces between veins bearing few to many appressed hairs 0.3-0.4 mm long, sometimes also glands which are never as abundant as on lower surface. Hairs on upper surface of rachis pale to brownish, curved, not over | mm long; on lower surface erect hairs as on costae. Sori medial, lower ones not divergent; indusia firm, usually with a few short hairs and glands; sporangia with glands near annulus. 458 FLORA MALESIANA [ser. Il, voluale Distr. Peninsular Thailand; Hainan, Hong Kong; throughout Malesia; North Queensland, Solomon Islands, New Hebrides, Fiji, Samoa. Ecol. In forest, low country to 1500 m. Notes. As here interpreted, this is a very variable species and varieties are not always sharply distinct; no formal names for them are here given. The following key attempts to dis- tinguish the various forms. KEY TO) THE LOCAL FORMS 1. Lower pinnae lobed at least 2/3 towards costa and at least 1.5 cm wide. 2. Glands none or rare on upper surface of pinnae. 3. Fertile pinnae of large plants 2cm wide (sterile ones sometimes wider); costules 4mm apart, second acroscopic vein usually touching side of sinus-membrane typical form 3. Fertile pinnae not over 1.5 cm wide; costules 3mm apart; second acroscopic vein always to margin (Dryopteris suprastrigosa ROSENSDID) ee) ee oe New Guinea 2. Glands + abundant on upper surface of pinnae. 4. Appressed hairs to 0.5mm _ long between veins on upper surface . Borneo 4. Short suberect hairs between veins on upper surface Kipel. eee. . Moluccas 1. Lower pinnae lobed not or little more than 1/2 way to costa, rarely more than 1.5 cm wide. 5. Glands and appressed hairs (sometimes dense) on upper surface . .Borneo (limestone) 5. Glands lacking on upper surface; appressed hairs variable, never dense. 6. Pinnae lobed distinctly more second acroscopic vein to margin. 7. Pinnae to 16 x 1.6 cm; costules almost at right angles to costa; pinnae more than 20 pairs; sori supramedial . Mindanao 7. Pinnae to 10 x 1.3 cm; costules at 60°; pinnae than 1/2; to 15 pairs; sori medial (Nephrodium pubescens BRACK., Dryopteris bordenii CHRIST) Eee? (oa ore ee Luzon 6. Pinnae lobed 1/2 or less deeply; second acroscopic vein usually ending beside sinus- membrane lowlands of W. Malesia The typical form occurs in mountain forest in Western Malesia and to Flores. In Malaya there seems to be no sharp distinction between the mountain form and the lowland one which comes at the end of the key. Plants like the lowland form of Malaya occur in Thailand, Hainan and Hong Kong, also in the lowlands of Borneo. BLUME distinguished varieties B and C in Java. I have found no authentic specimen of the former; var. C has rather large almost sterile fronds with sori on basal veins only and is not a distinct variety. The Borneo form with many glands and ap- pressed hairs on the upper surface occurs in Sarawak and Sabah at low to medium altitudes. The limestone Bornean form is abundant in the Gunong Mulu National Park; at higher altitudes and in more exposed positions fronds are smaller and very densely covered with hairs on the upper surface, but there seems to be no sharp distinction between these and plants with larger fronds less densely covered with hairs. In the Philippines neither the typical form nor the lowland form of Western Malesia has been found. From Palawan is one collection somewhat resembling the Borneo form; from Mindanao are two specimens (RAMOS & PASCASIO BS 34476, Surigao Province, and EDANO PNH 12581, Davao Province) which seem distinct and are keyed as the Mindanao form, but a third (COPELAND 1643, San Ramon, 500m) is nearer to the New Guinea form. In Luzon the form represented by Dry- opteris bordenii (small, with short fronds) appears common; in herbaria it has been confused with the large specimens of S. lobatus (COPEL.) HOLTTUM. Dryopteris suprastrigosa appears to be the common form in eastern New Guinea; few col- lections have been made in the west. Specimens from North Queensland are similar but less hairy on the upper surface of pinnae. D. mixta was described from a small fertile plant with most veins free but does not appear distinct in other characters. The Moluccan form is represented by BROOKS 18112 (Amboina) and KORNASSI 1222 (West Ceram). BACKER & POSTHUMUS cite Aspidium dimorphum KUNZE as a synonym. This was based on a specimen of JUNGHUHN’s from Mt Pangerango in W. Java; I have not found it. METTENIUS (l.c. 1864) described it briefly as a synonym of A. heterocarpon var. B BL. He stated that its aerophores were enlarged, for which reason I doubt whether it belongs here. 25. Sphaerostephanos isomorphus HOLTTUM, sp. nov. Stipes ignotus ; pinnae redactae 6-jugatae, omnes parvae; lamina 90 cm long; pinnae inferiores basi non angustatae; pinnae maximae 14.5 x 2.0 cm, profunde lobatae; venae 12-jugatae, infimae anas- tomosantes; costae subtus pilis erectis minutis vestitae; pagina inter venas utrinque glandulosa; indusia parva, glandulifera.-_Type: FLENLEY, ANU 2062, Papua New Guinea, W. Highlands, Lai valley (K). Caudex and stipe not known. Reduced pinnae 6 pairs, 4cm apart, all very small with enlarged aerophores; one intermediate pair of pinnae also present. Lamina 90 cm long; pinnae 40 pairs, light green when dry, firm; lower pinnae not narrowed at base. Largest pinnae 14.5 x 2.0 cm, sterile and 1981] fertile isomorphous; base truncate and slightly dilated; apex caudate-acuminate (cauda 22.5 cm), edges lobed to 2mm from costa, lobes slightly falcate; costules 3-3.5 mm apart, at more than 60°; veins to 12 pairs, basal pair only anastomosing with excurrent vein to short sinus-membrane, next veins both to edge; lower surface of rachis, costae and costules bearing very short erect hairs and glands, between veins many glands but no hairs; hairs on upper surface of costae pale, less than 1 mm long, scattered hairs 0.5 mm long on costules and veins, glands between veins. Sori medial, lower ones divergent, those on basal veins from adjacent costules often touching; indusia very small, glandular; sporangia bearing glands. Distr. Malesia: Papua New Guinea, only known from the type. 26. Sphaerostephanos dimidiolobatus HOLTTUM, sp. nov. Pinnae redactae usque 12-jugatae, superiores 2.5-3.0 cm longae; aerophori non elongati; pinnae normales usque 27 X2.3cm, dimidio costam ver- sus lobatae; venae 8—10-jugatae, infimae solum anastomosantes; costae costulaeque subtus glanduliferae, glandulae inter venas paucae; pagina superior inter venas glandulis pilisque minutis praedita; indusia parva; sporangia glandulifera. —Type: J.R. CRorr 324, New Ireland, 7km WNW of Taron, 700m (K; NSW). Stipe 22-27 cm long, almost glabrous, slightly flushed with red, scales thin, c. 10 x 2 mm; base of stipe to first normal pinna 50-55 cm; reduced pin- nae 8-12 pairs, uppermost 2.5-3.0cm_ long, broadly triangular with subtruncate base slightly auricled both sides, edges crenate, lowest reduced pinna | cm long. Lamina excluding basal pinnae c. 100cm long; basal pinnae slightly narrowed in basal third, basal lobes both slightly elongate; aerophores not elongate. Largest pinnae 20-27 cm long, to 2.3 cm wide; base truncate, not dilated nor auricled; apex narrowly acuminate, not caudate; edges lobed about half-way to costa (on one frond slightly more than half), lobes hardly falcate; cos- tules 4.5-S mm apart, at more than 60° to costa; veins 8-10 pairs, slender, slightly prominent when dry, basal pair at a broad angle to costule, anas- tomosing with an excurrent vein to the sinus, next pair usually both to sides of sinus-membrane; lower surface of rachis bearing very short hairs and some to 0.5mm long, costae minutely hairy near base only, with glands throughout, few short hairs and many glands on costules, few glands between veins; upper surface of rachis and costae bearing hairs 0.5mm long, minute hairs on cos- tules and veins, between veins a variable number of glands and very short erect hairs. Sori medial; indusia small, shrivelled, with a few short hairs and (?) glands; glands present on body of some sporangia, more commonly sessile on stalks of sporangia; spores not seen (sori old). THELYPTERIDACEAE (Holttum) 459 Distr. Malesia: Papua New Guinea (S. New Ireland), 2 collections. Ecol. In forest at 650-700 m. 27. Sphaerostephanos solutus HOLTTUM, sp. nov. — Aspidium hispidulum DECNE var. solutum MIQUEL, Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 160. — Aspidium hispidulum var. B METT. ibid. | (1864) 234. — Type: SPANOGHE (?), Timor (L, n. 908, 333-406). Base of frond lacking. Lamina of specimen 55cm long with 20 pairs pinnae. Largest pinnae: 19 x 2.1 cm, base truncate with acroscopic lobe of many pinnae elongate, apex caudate-acuminate, edges lobed to 2.5mm from costa, lobes falcate; costules 4-4.5mm apart, at 60°; veins 12 pairs, 1-15 pairs anastomosing, one vein or a pair to sinus-membrane; hairs on lower surface of costae and rachis sparse, erect, 0.3mm long, similar but shorter and more sparse on costules, glands also present; between veins glands and sparse short erect hairs; hairs on upper surface of costae 0.3mm long, similar hairs scattered on costules and veins, very short erect hairs and glands be- tween veins. Sori medial, lower ones not diver- gent; indusia thin, with a few glands and short hairs; sporangia with glands on body. Distr. Malesia: Lesser Sunda Is. (Timor), only known from the type. Note. The type bears the name Aspidium solutum n. sp. in BLUME’s hand, also Aspidium hispidulum var. B METT. and is evidently the one referred to by METTENIUS and MIQUEL, but they give ZIPPELIUS as collector whereas the speci- men here cited bears the collectors name SPANOGHE. A. hispidulum DECNE (P) =Christ- ella hispidula. 28. Sphaerostephanos hernaezii HOLTTUM, sp. nov. Caudex suberectus: stipes 10-15cm longus: pinnae redactae c. 6-jugatae, superiores 5-6mm longae; lamina 45cm longa; pinnae 9-jugatae, inferiores usque 11.5 x 2.5 cm, basin versus sensim angustatae, dimidio costam versus lobatae; cos- tulae 7mm inter se distantes; venae 9-jugatae; pagina utrinque pilis erectis brevibus glandulisque praedita; sori mediales, indusia pilis brevibus vestita. —Type: M. G. PRICE & B. E. HERNAEZ 75, Western Samar (K). Caudex suberect. Stipe 10-15 cm long, glabrous apart from short hairs in groove; basal scales 8x 1mm, copiously setiferous; base of stipe to first large pinna 45 cm. Reduced pinnae c. 6 pairs, uppermost 5-6 mm long, lowest 2mm. Lamina to 45 cm long; pinnae to 9 pairs, widely spaced, basal ones longest; apical lamina almost pinna-like but with widened base; aerophores not elongate. Basal pinnae to 11.5X2.5cm, widest 1/3 from apex and gradually narrowed to narrowly cuneate base, in middle lobed half way to costa, apex 460 FLORA MALESIANA fser. II, vol. 1° short-acuminate; lobes slightly falcate, tapered from base to an obtuse tip; costules to 7mm apart; veins to 9 pairs, 15 pairs anastomosing, one vein or a pair to short sinus-membrane; hairs on lower surface of rachis and costae 0.2-0.3 mm long, antrorsely curved, shorter hairs on costules and veins, between veins minute erect hairs and copious glands; hairs on upper surface of rachis and costae to 0.5 mm, rest as lower surface. Upper pinnae gradually less narrowed at base but up- permost somewhat narrowed. Sori medial, basal ones a little divergent and a little elongate along veins; indusium with copious short hairs and some glands; no glands nor hairs on sporangia. Distr. Malesia: Philippines (Samar), only known from the type. Ecol. In primary forest on limestone and limestone-derived soil. Note. Young plants have fewer and short pinnae but almost equally wide, with few lobes and widely-spaced costules. 29. Sphaerostephanos nov. S. pennigero affinis, ab eo differt: fronde minore; pinnis redactis 3—S-jugatis, omnibus par- vis; pinnis normalibus 7-12-jugatis, fertilibus usque 15 X2.1 cm; venis 23-jugatis anastomosan- tibus; sporangiis nec setiferis nec glanduliferis. — Type: MOSELEY s.n. Challenger Exp., Aru Islands (K; BM). Caudex short-creeping; stipe 20-35cm long, basal scales 7 x 1 mm; base of stipe to first normal pinna 35-65 cm; reduced pinnae 3-5 pairs, lowest very small, uppermost c. 5mm long. Lamina to 50cm long; pinnae 7-12 pairs. Largest fertile pin- nae 15 x2.1cm, shaped as in S. penniger but with costules at a very wide angle to costa; veins 8 pairs, 2; pairs anastomosing, | pair to sinus-mem- brane; pubescence as in S. penniger but some short erect hairs in addition to glands between veins on both surfaces. Sori medial, lower ones divergent, those on basal veins from adjacent costules sometimes confluent; indusia copiously glandular; sporangia lacking glands or hairs dis- tally, glands at ends of hairs on sporangium-stalks conspicuous. Distr. Malesia: S.E. Moluccas (Aru Is.). Notes. In addition to the type are BUWALDA 4984 and 5169, both from Kobroor I., the former much like the type. Number 5169 (from forest in steep rocky limestone country) is sterile, with longer fronds than the type and pinnae to 24x 2.7 cm, with 13-14 pairs of veins but only 23 pairs anastomosing, 2 or 3 pairs passing to a long sinus- membrane. moseleyi HOLTTUM, sp. 30. Sphaerostephanos productus (KAULF.) HOLTTUM, Kalikasan 4 (1975) 59.— Aspidium productum KAULF. Enum. Fil. Chamisso (1824) 237; METT. Farngatt. IV (1858) 109. — Dryopteris producta (KAULF.) C. CHR. Ind. Fil (1905) 286; v.A.v.R. Handb. (1908) 230.— Cyclosorus productus (KAULF.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 248; COPEL. Fern FI. Philip. (1960) 348, nomen tantum. — Thelypteris producta (KAULF.) REED, Phytologia 17 (1968) 306.— Type: CHAMISSO, Manila (LE). Nephrodium philippinense BAK. Ann. Bot. 5 (1891) 327.— Dryopteris basilaris C. CHR. Ind. Fil. (1905) 254, nom. nov. superfl.; CHRIST, Philip. J. Sci. 2 (1907) Bot. 186; v.A.v.R. Handb. (1908) 820. — Dryopteris philippinensis (BAK.) COPEL. Philip. J. Sci. 56 (1935) 100. — Lectotype (HOLT- TUM 1975): CUMING 10, Luzon (K). Dryopteris luzonica var. puberula CHRIST, Philip. J. Sci. 2 (1907) Bot. 197.—Type: BOL- STER 175, Luzon (P). Dryopteris kotoensis HAYATA, Ic. Pl. Formosa 5 (1915) 279, f. 107.— Cyclosorus truncatus var. kotoensis H. Iro, Bot. Mag. Tokyo 51 (1937) 729. — Thelypteris kotoensis (HAYATA) K. IWATS. Acta Phytotax. Geobot. 21 (1964) 42; Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 189. — S. kotoensis (HAYATA) HOLTTUM in C. M. Kuo, FI. Taiwan 1 (1975) 436.— Type: KAWAKAMI & SASAKI S.n. July 1912, Taiwan, Kotosho I. (TI; seen by IWATSUKI). Dryopteris pseudohirsuta’ ROSENST. Meded. Rijksherb. n. 31 (1917) 7.— Thelypteris pseudo- hirsuta (ROSENST.) REED, Phytologia 17 (1968) 306. — Type: CUMING 82, partim (L). Cyclosorus serratus COPEL. Philip. J. Sci. 81 (1952) 36; Fern Fl. Philip. (1960) 365.— Type: COPELAND P. P. E. 19, Luzon, Lamao River (MICH; K). Cyclosorus weberi COPEL. Philip. J. Sci. 81 (1952) 36, t. 25; Fern Fl. Philip. (1960) 366. — Thelypteris weberi (COPEL.) REED, Phytologia 17 (1968) 324.—Type: WEBER 1174, Mindanao (US; K). Cyclosorus rigidus COPEL. Philip. J. Sci. 81 (1952) 27; Fern FI. Philip. (1960) 340, not C. rigi- dus (RIDL.) COPEL. 1951. — Thelypteris ugoensis REED, Phytologia 17 (1968) 322, nom. nov. — Type: RAMOS BS 5744, Luzon, Mt Ugo (MICH). Cyclosorus leucadenius COPEL. Philip. J. Sci. 81 (1952) 27; Fern Fl. Philip. (1960) 339.— Thelypteris leucadenia (COPEL.) REED, Phy- tologia 17 (1968) 287.—Type: ELMER 8999, Luzon, Baguio (MICH; K, E, L, BO). S. punctatus HOLTTUM, Kalikasan 4 (1975) 60.— Type: ELMER 16642, Luzon, Mt Bulusan (US; K, L, G, BO, NSW). Cyclosorus megaphyllus sensu COPEL. Fern FI. Philip. (1960) 367. Cyclosorus nitidulus sensu COPEL. Fern FI. Philip. (1960) 364, p.p. Caudex short-creeping, branched. Size of fronds very variable, with pinnae from 101 to 30 2.7cm. Stipe short; basal scales 10-20mm long, narrow. Reduced pinnae 6-12 pairs, not 1981] deflexed, broadly ovate to bluntly triangular, subentire, uppermost commonly I-1.5 cm long and wide, lowest 0.5cm. Normal pinnae crenate in smallest fronds, commonly lobed almost 1/3, in largest fronds 1/2 towards costa; costules always at about 45° to costa except in largest fronds; veins 4-11 pairs, in almost all cases 12 pairs anas- tomosing and | pair to sinus-membrane; hairs on lower surface of costae near base very short (rarely with some more than 0.5mm long), to 0.2mm or more long distally, very short and sparse on costules and veins, many glands and few hairs between veins; hairs on upper surface of costae to 0.5 mm long, on costules and veins all very short, glands between veins abundant. Hairs on lower surface of rachis variable, 0.2-0.5 mm or more long, on upper surface to Imm long. Sori medial, lower ones not or little divergent; indusia firm, bearing glands and sometimes a few short hairs; sporangia bearing 1-2 setae or glands on body and a spherical gland at end of hair on stalk; spores with many small wings. Distr. Malesia: Philippines (Luzon to Min- danao); Taiwan (Orchid Island only). Ecol. In forest at low and medium altitudes. Notes. In 1975 I distinguished S. punctatus, with large pinnae lobed fully half-way to costa and costules at a rather broad angle, but these do not appear to be sharply distinct; such specimens occur also on Orchid Island (S.E. of Taiwan) along with others indistinguishable from the type of Nephrodium philippinense BAK. The specimen originally described by KAULFUSS has pinnae 9x0.9cm; the type of Cyclosorus serratus CoPEL. 10x0.8cm, the type of Nephrodium philippinense 18 x 1.4cm; some other types come between these extremes. The name Nephrodium basilare PRESL (Epim. Bot. 258) was published as a substitute for N. caudiculatum J. SM. non PRESL (Hook. J. Bot. 3:411), but SMITH’s name was a nomen nudum, and the first valid name based on the specimens cited by SMITH was N. philippinense BAK. When transferring this to Dryopteris, CHRISTENSEN (1905) preferred the epithet basilaris because of the existence of Phegopteris philippinensis METT. METTENIUS evidently did not see _ the CHAMISSO specimen (he cited CUMING LI, which I cannot trace), but he gave a fairly good description, mentioning the glands on the lower surface which no other author noticed. HOOKER and BAKER made no reference to Aspidium productum KAULF. CHRISTENSEN, CHING, COPELAND and REED transferred the species to other genera without describing it. When describ- ing Dryopteris producta, VAN ALDERWERELT referred to METTENIUS, but did not copy MET- TENIUS’s description; he gave a better descrip- tion under D. basilaris. COPELAND misconstrued the species in his Fern Flora of the Philippines, placing it as a synonym of Nephrodium nitidulum THELYPTERIDACEAE (Holttum) 461 PRESL, which belongs to the genus Pneumatop- teris. 31. Sphaerostephanos penniger (HOOK.) HOLT- TUM in Nayar & Kaur, Comp. to Bedd. (1974) 209.— Aspidium pennigerum sensu BL. Enum. Pl. Jav. (1828) 153.— Nephrodium pennigerum Hook. Spec. Fil. 4 (1862) 82, nom. nov.; RACIB. Fl. Btzg 1 (1898) 190.—Type: BLUME, Java, Boerangrang (L). Aspidium megaphyllum METT. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 233; MIQUEL, ibid. 4 (1869) 159. — Dryopteris megaphylla (METT.) C. CHR. Ind. Fil. (1905) 277: v-A.v.R. Handb. (1908) 218: BACKER & POSTH. Varenfl. Java (1939) 50.— Cyclosorus megaphyllus (METT.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 225; HOoLT- TUM, Rev. Fl. Mal. 2 (1955) 268, f. 154.— Thelypteris megaphylla (METT.) K. IWATS. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 34.— Type: METTENIUS cited REINWARDT and ZIPPELIUS, Java (L). Aspidium multilineatum WALL., nom. nud. — Nephrodium pennigerum var. multilineatum CLARKE, Trans. Linn. Soc. Bot. | (1880) 532. — Nephrodium multilineatum BEDD. Handb. Suppl. (1892) 80 (not Aspidium multilineatum METT. 1858).— Dryopteris multilineata (BEDD.) O. KiTrZE; ‘Rev; Gen! (PL? (1891) she wAy Handb. (1908) 219.—Type: WALLICH 353, Penang (K). Aspidium abortivum sensu METT. Farngatt. IV (1858) 110, quoad ZOLLINGER 3525 tantum. Nephrodium abruptum sensu HOOK. Spec. Fil. 4 (1862) 78, p.p., t. 241B. — Fig. la-h. KEY TO THE VARIETIES 1. Lower pinnae not over 4cm wide, lobed not more than 1/3 towards costae. 2. Normal pinnae to 25 pairs, reduced pinnae 10-15 pairs : . a. var. penniger . Normal pinnae 7 pairs: reduced pinnae 5 pairs c. var. karoensis lobed to 3mm . b. var. excellens 1. Lower pinnae to 7cm wide, from costa a. var. penniger Caudex short-creeping. Stipe 8-10cm long; scales to 10x 1mm, closely setiferous; base of stipe to first large plant 80cm. Reduced pinnae 10-15 pairs, upper ones to 32cm, spreading, triangular with a not quite truncate symmetric base, blunt apex and subentire edges, lowest on a large plant 1x 1cm. Lamina to 100 cm long; pin- nae 25 pairs; basal pinnae not or slightly narrowed at base; apex not pinna-like; aerophores slightly swollen. Largest pinnae commonly 20 x 2.8 cm (to 30*x4cm), lobed to a depth of 3-Smm, lobes oblique, subdeltoid, broadly pointed; costules 5.5- 7mm apart, at 60°; veins 8-12 pairs, 3-33 pairs anastomosing to form a zig-zag excurrent vein, 462 1-2 pairs to sinus-membrane; hairs on lower sur- face of rachis and base of costae very short, distally on costa and costules 0.2-0.3 mm, some- what antrorse, between veins many glands; hairs on upper surface of costae 0.5 mm long, minute on costules, many glands between veins. Sori medial, lower ones not divergent; indusia glandular with a few short acicular hairs; sporangia with short setae and glands on body, hair on stalk often with a conspicuous terminal gland. Distr. Peninsular Thailand, in Malesia: Malaya, Sumatra, Borneo, Java, Lesser Sunda Is. (Bali, Lombok), S.W. and N. Celebes. Notes. On p. 78 of Species Filicum vol. 4 (which was printed off in fascicles of 16 pages) HOOKER cited his specimens of this species un- der Nephrodium abruptum and with them speci- mens belonging to six other species; his plate 241B certainly represents a specimen of S. pen- niger. In the next fascicle, on p. 82, he revised his opinion and cited all his specimens of the present species under N. pennigerum, based on Aspidium pennigerum BL. stating correctly that they agreed with a specimen so named by BLUME. HOOKER’s description is a translation of BLUME’s; his name, in Nephrodium, ranks as a new name. BLUME’s name was based on a_misinter- pretation of Polypodium pennigerum FORST. which represents a New Zealand species very different from the Java one, and in citing BLUME HOOKER excluded the FORSTER name; in his fifth volume HOOKER retained Polypodium pen- nigerum FORST. for the New Zealand fern, which is exindusiate. HOOKER’s confusions on p. 78 misled BED- DOME, who united an Indian species with the Malesian S. penniger under the name Nephrodium abruptum in Ferns of S. India, and then under N. pennigerum (Handb. p. 277). In the Supplement to his Handbook BEDDOME separated the two, adopting WALLICH’s name multilineatum for the Malesian species; this in turn led to confusions in the name of another Indian species. b. var. excellens (BL.) HOLTTUM, Stat. nov. — Aspidium excellens BL. Enum. Pl. Jav. (1828) 160; C. CHR. Ind. Fil. (1905) 73.— Proferea excellens (BL.) PRESL, Epim. Bot. (1851) 259; HOLTTUM, Novit. Bot. Univ. Carol. Prag. 1968 (1969) 48.— Pleocnemia excellens (BL.) v.A.v.R. Handb. (1908) 171.— Tectaria excellens C. CHR. Ind. Fil. Suppl. III (1834) 179. — Type: BLUME, Java (PRC; L, K). Upper part of frond indistinguishable from the typical form of S. penniger; lower pinnae greatly enlarged, lowest 30 7cm, lobed to 3mm from costa, costules 9mm apart, lobes falcate and tapered to an acute tip, venation in lobes pleoc- nemioid, in largest lobes aeroles of more than one series along costule. Distr. Malesia: Java, only one collection. FLORA MALESIANA [ser. II, vol. 1° Note. In the Rijksherbarium at Leiden are two sheets, one bearing the apical part of a frond, the other a lower part of the same frond, but not the base. The base of the frond appears to have been divided between HOOKER (one pair of pinnae) and PRESL who had the three lowest pairs of large pinnae and two pairs of reduced pinnae. c. var. karoensis HOLTTUM, var. nov. A typo Sspeciei differt: fronde minore; pinnis redactis S-jugatis; pinnis evolutis 7-jugatis, usque 13 x 2.7 cm, infimis basin versus valde angustatis ; sporangiis setiferis, non glanduliferis. —Type: MOLESWORTH ALLEN 2407, Sumatra, Prapat by Lake Toba (K). Distr. Malesia: N. Sumatra, only known from the type. 32. Sphaerostephanos warburgii (KUHN & CHRIST) HOLTTUM, Allertonia 1 (1977) 202.— Aspidium warburgii KUHN & CHRIST in War- burg, Monsunia 1 (1900) 81.— Dryopteris war- burgi (KUHN & CHRIST)» 9G io 3@ar: Ind. Fil. (1905) 98; v.A.v.R. Handb. (1908) 180. — Cyclosorus warburgii (KUHN & CHRIST) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 250; COPEL. Philip. J. Sci. 78 (1951) 441.— Thelypteris warburgii (KUHN & CHRIST) B. C. STONE, Micronesica 2 (1966) 3.— Type: WAR- BURG, N.E. New Guinea, Sattelberg (B). Dryopteris calcicola C. CHR. Bot. Jahrb. 66 (1933) 44.—Type: KJELLBERG 2611, S.E. Celebes, Kosali-Porema (S-PA; BM). Caudex short, erect. Stipe 1-3 cm long. Lamina 10-15 cm long; pinnae to 18 pairs, several lower pairs gradually smaller, lowest 3-5 mm long. Lar- gest pinnae 1.5—1.8cm long, 0.3-0.4 cm wide, dis- tinctly stalked, base unequally cuneate, edges entire or rarely crenulate, apex rounded or obtusely pointed; costules 1.5mm apart, at 45°, distal ones unbranched, rest forked once or twice, branches of neighbouring ones anastomosing; lower surface of costae bearing short hairs, some- times with scattered long ones, glands throughout between veins; upper surface glabrous apart from costae and costules, not glandular. Hairs on rachis to 1 mm long, both sides. Sori uniseriate on each side of the costa; indusia with a few short hairs; sporangia bearing glands, on specimens from Tenimbar Islands and Celebes also setae. Distr. Micronesia (Guam, Admiralty Is.), and Malesia: New Guinea, S.E. Moluccas (Tenimber Is.), and Central & S.E. Celebes. Note. Specimens from Guam only differ from those of New Guinea by their larger size (frond to 30cm long, pinnae to 3cm long), agreeing in glandular sporangia. The type of D. calcicola, from limestone, differs in fewer glands on lower surface of pinnae, densely hairy indusia and setose sporangia. Ecol. On rocky stream banks. 1981] 33. Sphaerostephanos tandikatensis (v.A.v.R.) HOLTTUM, comb. nov.— Dryopteris tandikaten- sis v.A.v.R. Bull. Jard. Bot. Btzg II, 11 (1913) 11; Handb. Suppl. (1917) 178, 503.— Type: MATTHEW 635, Sumatra, G. Tandikat (BO). Caudex short-creeping. Stipe 4-5 cm long, short- hairy; basal scales 3-4 = 1mm; base of stipe to first large pinna 35-45cm. Reduced pinnae 12- 15 pairs, broadly ovate, uppermost 10 x 8 mm with a small auricle on acroscopic base, lowest 3-4 mm long. Lamina to 45cm long; pinnae 3-4 pairs, lowest pinna usually a little reduced; apex of lamina pinna-like, larger than pinnae on fronds of young plants; aerophores slightly swollen. Pinnae of type to 11 x 4cm, of another specimen to 17 x 6.5cm, widest at or above middle, base abruptly broad-cuneate, of lower ones asymmetric; apex abruptly short-cuspidate; edges entire, or crenate distally; costules 3.5-4.5 mm apart, at 60° or rather less; veins 12-14 pairs, almost all anastomosing, combined excurrent veins almost straight, mostly continuous; lower surface of costae and costules densely covered with erect hairs 0.1mm long, veins and surface less densely, glands present on veins and surface between them: upper surface covered with appressed hairs 0.3mm long, no longer hairs present. Hairs on both sides of rachis also short. Sori all supramedial, those on veins from adjacent costules often coalescing; indusia small, short-hairy; sporangia with many setae. Distr. Malesia: South and Central Sumatra, 720-1400 m, several collections. 34. Sphaerostephanos carrii HOLTTUM, sp. nov. Pinnae redactae 4-jugatae, superiores 1.5cm longae; lamina usque 45cm longa; pinnae 6-7- jugatae, omnes oppositae, 10 x 2.7 cm vel majores, vix usque 1/3 costam versus lobatae; lamina inter venas subtus glandulosa, supra glabra eglan- dulosa; indusia tenuia, pilis brevibus glandulisque praedita; sporangia setifera. Type: C. E. CARR 14202, Papua, Boridi, 1500 m (K; BM, L, LAE). Caudex short-creeping or suberect. Stipe 12- 20cm long; basal scales 7-10 x 1.5-2 mm, thin, setiferous; base of stipe to first large pinna 40— 50cm. Reduced pinnae 4-5 pairs, lowest 4mm long, uppermost 1.5cm, deflexed and auricled. Lamina 304S5cm_ long; pinnae 6-7 pairs, opposite; lowest pinnae narrowed in basal 1/3 on basiscopic side, truncate but not auricled on acroscopic side; apical lamina almost pinna-like but larger and more deeply lobed at base; texture thin; aerophores not elongate. Suprabasal pinnae 10-17 x 2.7-3.5cm, base truncate, apex abruptly short-acuminate, edges lobed rather less than 1/3 to costa, lobes slightly falcate with rounded tips; costules 5-6 mm apart; veins 10-12 pairs, 23 pairs anastomosing, I-l> pairs passing to sinus mem- brane; lower surface of costae and costules bear- ing short spreading hairs, distally on costa to 0.5mm long, short erect hairs and sparse glands THELYPTERIDACEAE (Holttum) 463 between veins; hairs of upper surface of costae mixed in length, longest 1 mm or more, scattered similar long hairs on costules and veins, surface between veins glabrous. Hairs on lower surface of rachis rather sparse, 1-1.5mm long, on upper surface copious, mostly 0.5 mm, longest I-1.5 mm. Sori medial, basal ones not divergent; indusia bearing short hairs and a few glands: sporangia setiferous. Distr. Malesia: Eastern New Guinea, 3 col- lections. 35. Sphaerostephanos exindusiatus HOLTTUM, sp. nov. Pinnae redactae 15-jugatae, superiores vix 10mm longae; lamina 60cm longa; aerophora 2mm longa; pinnae maximae 12X1.5cm, 3/5 costam versus lobatae; venae 8-jugatae: costa subtus pilis erectis 0.5 mm longis praedita, lamina inter venas glandulosa; pagina superior omnino pilis appressis vestita; sori exindusiati, sporangia setifera.— Type: POSTHUMUS 3494, S.W. Celebes, near Patapang, 1200m, on river bank (BO). Caudex short, thick, suberect. Stipe 15-20cm long, densely scaly throughout when young: scales to 10 x 2 mm, thin; base of stipe to first large pinna 50cm. Reduced pinnae at least 15 pairs, 1.5-2.cm apart, lowest 4mm long, uppermost hardly 10 mm. Lamina 60cm long; pinnae 25 pairs or more: lower pinnae not or little narrowed at base; aero- phores 2mm long. Largest pinnae 12 1.5cm, base truncate and a little dilated both sides, apex narrowly acuminate, edges lobed about 3/5 to costa, lobes slightly falcate; costules 3-3.5mm apart; veins to 8 pairs, 1 pair anastomosing, next acroscopic vein to side of sinus-membrane; lower surface of costae and costules bearing rather sparse erect hairs 0.5mm long, surface between veins copiously glandular; hairs on upper surface of rachis and costae 1 mm long, scattered similar hairs on costules and veins, whole surface covered with slender appressed hairs. Sori in- framedial, exindusiate; young sporangia bearing several short setae. Distr. Malesia: S.W. Celebes, only known from the type. 36. Sphaerostephanos lucbanii HOLTTU™, sp. nov. Pinnae redactae 2-jugatae; lamina 25 cm longa: pinnae liberae 7-jugatae, tenues, maximae 6X 1.8 cm, 1/3-2/5 costam versus lobatae; venae 5—6- jugatae: costae subtus pilis usque 1 mm longis vestitae, pagina inter venas pilis erectis glan- dulisque praedita; pagina superior pilis appressis vestita: sori exindusiati; sporangia setifera. — Type: M. G. Price 2720A, Luzon, Mt Bana- haw, streamside at 1000 m (K). Caudex slender, short-creeping. Stipe 5cm long, densely short-hairy, scales 3 mm long; base of stipe to first large pinna 11 cm. Reduced pinnae 464 FLORA MALESIANA 2 pairs on largest frond, lower ones 63mm, upper one deflexed, 106mm, crenate, base asymmetric. Lamina 25cm long, thin, apex nar- rowly triangular, 13cm long, deeply lobed and grading to upper pinnae, venation in its basal lobes as in Haplodictyum; free pinnae 7 pairs, lowest distinctly stalked, somewhat narrowed at base on basiscopic side, aerophores swollen. Largest pin- nae 6X 1.8cm, base truncate, apex shortly poin- ted, edges lobed 1/3-2/5 to costa, lobes rounded, entire; costules 4.5mm apart, at 60°; veins 5-6 pairs, | pair anastomosing with long excurrent vein, next acroscopic vein to short sinus-mem- brane; hairs on lower surface of costae rather sparse, pale, slender, 0.5-1 mm long, on costules same but few, between veins slender erect hairs 0.2 mm long and glands; hairs on upper surface of costae 0.5 mm long, rest of upper surface covered with slender appressed hairs 0.2-0.3mm_ long. Hairs on both sides of rachis slender, to 1.5mm long. Sori medial, exindusiate; sporangia seti- ferous. Distr. Malesia: Philippines (Luzon), only known from the type. Note. Under the same number and near the same locality Mr PRICE also collected very much smaller plants with lamina to 7 cm long, pinnae 2 pairs, | cm long, subentire, but fully fertile. These small plants (separated as n. 2720B) agree in sori and in pubescence with the larger ones above described. They agree also closely in all respects with Dryopteris diminuta COPEL. from Mindanao, which I have transferred to Pronephrium. Mr PRICE calls my attention to the plants described by COPELAND as Dryopteris bakeri in Philip. J. Sci. 38 (1929) 135; one of them had much larger pinnae than the rest. It is thus possible that the whole collection 2720 represents different stages of plants on one species. 37. Sphaerostephanos major (COPEL.) HOLTTUM, comb. nov.— Haplodictyum majus COPEL. in Elmer, Leafl. Philip. Bot. 9 (1920) 3110; Fern FI. Philip. (1960) 378; HoLtTTuM, Kalikasan 2 (1973) 61.— Type: RAMOS BS 13992, Luzon, Mountain Prov., Apayao (MICH). Cyclosorus dimorphus COPEL. Philip. J. Sci. 83 (1954) 99, pl. 5, non (BRAUSE) COPEL. 1960. — Type: EDANO PNH = 17282, Luzon, Mountain Prov., Mt Magnas (MICH). ? Dryopteris bakeri sensu COPEL. Philip. J. Sci. 38 (1929) 135. Caudex short-creeping; fronds dimorphous. Sterile frond. Stipe to 4cm long; base of stipe to first large pinna 6-12cm; reduced pinnae 1-2 pairs, very small; apical lamina 12-20cm long, deeply lobed, 2.5—4 cm wide at base, veins in lobes mostly forked, the branches uniting to form a series of costular areoles, base of lamina grading into 2-4 pairs of pinnae closely placed; pinnae on largest frond 3.5 x 2.cm, broadly pointed, lobed to [ser. II, vol. 1° about 1/3, with long spreading hairs on lower surface of costae, short erect hairs and glands between veins, upper surface covered with short appressed hairs. Fertile frond. Stipe 13cm or more, base of stipe to first large pinna 20-27 cm; reduced pinnae as sterile but more widely spaced; apical lamina to 12 x2.cm, veins mostly not for- ked; pinnae to 2.5 1.4cm, more widely spaced and less deeply lobed than sterile; sori on both apical lamina and pinnae, basal ones sometimes a little elongate; indusium small with many hairs; sporangia with 2-3 long setae and sometimes also a gland. Distr. Malesia: northern Luzon). Philippines (mountains of 38. Sphaerostephanos polyotis (C. CHR.) HOLT- TUM, comb. nov. — Dryopteris polyotis C. CHR. Bot. Jahrb. 66 (1933) 46. — Thelypteris polyotis (C. CHR.) REED, Phytologia 17 (1968) 305.— Type: KJELLBERG 1718, S.W. Celebes, Todjamboe, below 1000 m (BO; BM). Caudex short, suberect. Stipe 5-10cm long; base of stipe to first normal pinna 35-70cm. Reduced pinnae 25-30 pairs, |cm apart on type (to 2.5cm on other specimens), lowest 4-5 mm long, uppermost 1.5-2.0cm long, deflexed with basal acroscopic lobe 7 mm long, basiscopic lobe shorter, edges incised, apex obtuse or rounded. Lamina of type 75cm long; normal pinnae 25 pairs, basal ones a little narrowed towards their bases, with enlarged basal lobes; aerophores not elongate. Lower surface of rachis densely short- hairy, hairs to 0.5mm long; upper surface with antrorse pale hairs. Largest pinnae of type 7 x 10cm (of WALKER 12187 14x2.0cm); base truncate; apex acuminate; edges lobed 1/2-2/3 towards costa; costules 3—3.5 mm apart, at 60° to costa; veins of type to 5 pairs, of WALKER 12187 10-12 pairs, in both cases | pair anastomosing and next acroscopic vein ending beside sinus-mem- brane; lower surface of costae densely covered with erect hairs of mixed length to 0.5 mm, more sparse hairs and sometimes glands on costules (not seen on type), surface between veins of WALKER 12187 bearing short erect hairs and a few glands; upper surface + closely covered with short appressed hairs. Sori medial; indusia bearing short hairs and sometimes a few glands; sporangia bearing glands and sometimes setae. Distr. Malesia: S.W. Central Celebes (Lati- modjong Mts). Notes. Additional collections are T.G. WALKER 12186-12188, 12270, 12273, 12282 (BM). Most of these are larger than the type, but 12188 has normal pinnae of the same size as the type with fewer reduced pinnae. WALKER’S specimens also have some glands on the lower surface of costules and between veins, but setae more abundant than glands on sporangia. This species and S. foliolosus are related to S. hispiduliformis 1981] THELYPTERIDACEAE (Holttum) 465 of New Guinea, but the transition from normal to reduced pinnae at the base of fronds of the latter is not abrupt. 39. Sphaerostephanos grandescens HOLTTUM, sp. nov. Pinnae redactae c. 6-jugatae, omnes parvae; lamina 45cm longa; pinnae 12-jugatae, 2/3 vel ultra costam versus lobatae; costulae 5 mm inter se distantes; pagina subtus inter venas sparsim glandulifera, supra omnino pilis appressis vestita; indusia pilis brevibus paucis praedita, sporangia setifera.— Type: MERRILL 6094, Mindoro, Mt Halcon (MICH). Stipe 7-14.cm long, glabrescent; base of stipe to first large pinna 45 cm or more. Reduced pinnae 6 pairs, 3rd from base 3—4mm long, upper ones apparently not much larger. Lamina 45 cm long; pinnae 12-13 pairs, basal pair widest above middle, narrowed to base which is 1.2cm wide, not auricled; aerophores elongate, less than | mm long. Largest pinnae 12.5 x 2.4 cm, base broadly cuneate, apex acuminate, edges lobed to 3.5mm from costa (more than 2/3), lobes oblique, slightly falcate, a little narrowed distally; costules 5mm apart, at less than 60°; veins to 9 pairs, 1 pair anastomosing, next acroscopic vein usually to side of sinus-membrane; hairs on lower surface of costae less than 0.5 mm long, antrorse distally, on costules same but sparse, sparse glands on cos- tules and on surface between veins; hairs on upper surface of costae 0.6mm, scattered similar hairs on costules and veins, whole surface covered with slender hairs 0.3-0.4mm long. Hairs on lower surface of rachis pale, curved, spreading, 0.6mm long, on upper surface similar but appressed. Sori medial, lower ones not divergent; indusia rather firm, with a few short hairs only; sporangia seti- ferous. Distr. Halcon). Note. This specimen was named Dryopteris bordenii (here placed as a variety of S. hetero- carpus) by CHRIST, but has much wider pinnae and setiferous sporangia and elongate aerophores; it is near S. magnus, but smaller. Malesia: Philippines (Mindoro: Mt 40. Sphaerostephanos magnus (COPEL.) HOLT- TUM, comb. nov. — Cyclosorus magnus COPEL. Philip. J. Sci. 81 (1952) 30; Fern FI. Philip. (1960) 346. — Thelypteris magnus (COPEL.) REED, Phy- tologia 17 (1968) 291.—Type: MERRILL 6952, Negros (MICH; B, P). Stipe 10cm, glabrescent; scales broadly ovate, thin, 2.5mm wide; base of stipe to first normal pinna to 85cm. Reduced pinnae many pairs, each consisting of an aerophore more than | mm long and a minute green blade. Lamina to 100 cm long; pinnae c. 30 pairs; basal pinnae apparently not narrowed at their bases. Largest pinnae 21x 2.5cm, base subtruncate, apex acuminate, edges lobed 3/5 to costa, lobes falcate, somewhat nar- rowed distally; costules 5mm apart, at more than 60°; veins to 9 pairs, basal pair anastomosing, next pair to sides of sinus-membrane; hairs on lower surface of rachis sparse, to | mm long, on costae and costules very short throughout with much longer ones also distally; many glands throughout lower surface; upper surface covered with slender appressed hairs, with scattered longer ones on costules and veins. Sori small, medial, lower ones not divergent; indusia bearing glands; sporangia setiferous, sometimes also a gland present. Distr. Malesia: Philippines (Negros). Known from two collections, the other being ELMER 9845 which consists in part of the present species, in part of Pneumatopteris nitidulus (PRESL) HOLTTUM. 41. Sphaerostephanos santomasii HOLTTUM, Kalikasan 4 (1975) 62. — Type: M. G. PRICE 1034, Luzon, Benguet Prov., Mt Santo Tomas, 2000 m (K; PNH). Caudex short, erect. Stipe 5-10 cm long. basal scales narrow, 8mm long; base of stipe to first large pinna 40-75cm. Reduced pinnae 34cm apart, upper ones 3mm long, lowest very small. Lamina to 55cm long; pinnae to 30 pairs; several pairs lower pinnae narrowed towards their bases, a basal short pair sometimes present; aerophores to 1mm long. Largest pinnae to 16x1.5cm (rarely to 2cm wide), base truncate, apex acu- minate, edges lobed 3/5—2/3 to costa, lobes slightly falcate; costules 34mm apart, at 60° or more; veins 7-10 pairs, basal pair anastomosing, next pair to edge or acroscopic one to sinus-membrane; lower surface of costae of sterile pinnae bearing pale spreading hairs 1mm long and more numerous shorter antrorse hairs, longer hairs on fertile pinnae less than | mm, similar hairs more sparse on costules, surface between veins bearing glands and a variable number of slender appressed hairs; hairs on upper surface of costae 1 mm long, scattered similar hairs on costules and veins, whole surface covered with slender appressed hairs 0.3-0.5 mm long. Sori medial, lower ones not divergent; indusia small, bearing a few short hairs; sporangia copiously setiferous. Distr. Malesia: Philippines (Northern Luzon: Mt Santo Tomas, Mt Nangaoto and Mt Data: ALCASID PNH 1750), at 2000 m. 42. Sphaerostephanos trichochlamys HOLTTUM, sp. nov. Caudex brevis, erectus; stipes 8cm longus; pinnae redactae usque 9-jugatae, superiores 3 mm longae; lamina 45cm longa; pinnae maximae 13 X 1.7 cm, dimidio costam versus lobatae; rachis costaeque subtus pilis multis usque 1.5 mm longis vestita, pagina inter venas pilis erectis glandu- lisque praedita; indusia pilosa; sporangia setifera. — Type: HOLTTUM 4, Mt Kinabalu (K). 466 FLORA MALESIANA [ser. II, volar Caudex erect, short. Stipe 8cm long, dark red- dish; base of stipe to first large pinna 50cm. Reduced pinnae to 9 pairs, uppermost 3 mm long, rest very small, with aerophores to 1 mm long. Lamina to 45cm long; pinnae to 20 pairs, red when young, lower ones narrowed towards their bases. Hairs on rachis, both surfaces, to more than 1 mm long, with shorter ones also on lower surface. Largest pinnae 13X1.7cm (sterile to 2cm wide), base (except lower ones) truncate, apex short-acuminate, edges lobed 2 way to costa; costules 3-4 mm apart, falcate distally; veins 7-8 pairs, thick and prominent below in sterile fronds, basal pair anastomosing, next acroscopic vein to sinus-membrane; lower surface of costae densely covered with spreading hairs 1-1.5mm long and much shorter ones, similar but more sparse hairs on costules and veins; surface between veins bearing short erect hairs and glands; upper surface of costae densely antrorse-hairy, sparse long hairs on costules and veins, surface between veins with a variable number of fine appressed hairs. Sori medial, lower ones not divergent; indusia firm, bearing many hairs 0.3mm _ long; sporangia copiously setiferous. Distr. Malesia: Borneo (Sabah: Mt Kinabalu, many specimens; Sarawak: G. Mulu). Ecol. In forest at c. 1500 m, not near streams; young fronds red, not mucilaginous. Notes. This is closely allied to S. hirsutus but is smaller and much more densely hairy, with copiously setose sporangia. A specimen from 2275 m on G. Mulu has few sori which have small glabrous indusia. 43. Sphaerostephanos adenostegius (COPEL.) HOLTTUM, comb. nov. — Dryopteris adenostegia COPEL. Univ. Cal. Publ. Bot. 18 (1942) 220. — Cyclosorus adenostegius (COPEL.) COPEL. Gen. Fil. (1947) 142; Philip. J. Sci. 78 (1951) 442, pl. 6B.— Thelypteris adenostegia (COPEL.) REED, Phytologia 17 (1968) 258.— Type: BRASS 10282, near Lake Habbema, New Guinea, 2800m, in forest (GH). Pronephrium nothofageti HoLTTUM, Blumea 20 (1972) 118. — Type: T. G. WALKER 8788, New Guinea, Finisterre Range, 2400 m, in Nothofagus forest (BM). Rhizome short-creeping, 3-4 mm diameter. Stipe commonly to 30cm long, minutely hairy, basal scales 5X 1mm. Lamina 20-30cm long; pinnae 8-10 pairs, lower ones with stalks 1-2 mm long, lowest 1-4 pairs + reduced, lowest sometimes 1.3 cm long; texture firm; aerophores not elongate; frond-apex deltoid, deeply lobed. Largest pinnae 5-8 cm long, 1.3-1.6cm wide, base subtruncate and sometimes slightly auricled, edges lobed about 1/3 to costa, lobes slightly dentate at ends of veins; costules 4-4)mm apart; veins to 5 pairs, one pair anastomosing, next acroscopic vein to side of sinus-membrane; hairs on lower surface of rachis pale, + curved, to 0.5 mm long, on costae 0.2-0.3 mm long, slightly antrorse, sparse glands usually present on costules and on surface be- tween veins; hairs on upper surface of rachis as lower, on costae short and sparse, scattered longer hairs on costules and veins, slender appressed hairs to 0.5mm long more or less abundant be- tween veins. Sori medial; indusia small, with a few short hairs and glands; sporangia bearing 1-3 slender setae; spores with a rather broad trans- lucent wing and cross-wings. Distr. Malesia: Eastern New Guinea, several localities in forest at 2300-2900 m. 44. Sphaerostephanos aquatilis (COPEL.) HOLT- TUM, comb. nov. — Dryopteris aquatilis COPEL. Philip. J. Sci. 6 (1911) Bot. 75.— Thelypteris aquatilis (COPEL.) REED, Phytologia 17 (1968) 260.— Type: C. KING 182, Papua (MICH, BO, NSW, P). Dryopteris caudiculata ROSENST. Repert. 9 (1911) 426, non v.A.v.R. Type: as above (BO). Caudex short, erect. Stipe to 18cm long, pale with dark base, short-hairy. Lamina to 45cm long, apical section narrowly triangular and deeply lobed; pinnae 20 pairs or more, well spaced, lower 3-4 pairs gradually smaller, lowest 0.8-1.3 cm long; in smaller fronds only | pair pinnae some- what reduced. Largest pinnae of type collection 8.5 x 1.0cm, of BRASS 6725 14 x 0.8 cm; base very narrowly cuneate on basiscopic side, at c. 45° on acroscopic; apex acuminate; edges very slightly crenate; costules little over 2mm apart, at 45°; veins of type 3 pairs, 2 pairs anastomosing; sinus- membrane hardly evident; hairs on lower surface of costae copious, antrorse, on costules sparse, rest of lower surface glabrous, glands present on costae and costules only of type (also between veins on BRASS 6725); upper surface bearing hairs on costae only, sparse distally. Sori medial to supramedial; indusia rather large, glabrous or sometimes bearing glands or a few short hairs; sporangia bearing glands. Distr. Malesia: S.E. Papua New Guinea; 2 collections (BRASS 6725, Fly River). Ecol. At low altitude by streams in flood zone. Note. Possibly not distinct from S. mutabilis. Fedde 1908. — 45. Sphaerostephanos _hispidifolius (v.A.v.R.) HOLTTUM, comb. nov.— Dryopteris hispidifolia v.A.v.R. Bull. Jard. Bot. Btzg II, 20 (1915) 15; Handb. Suppl. (1917) 186.— Thelypteris hispidi- folia (v.A.v.R.) REED, Phytologia 17 (1968) 283. — Lectotype (selected here): JAHERI 1124, Borneo (BO). Nephrodium hispidulum (non (DECNE) BAK.) CHRIST, Ann. Jard. Bot. Btzg 20 (1906) 107. — Fig. 12]-m. Caudex short-creeping, with a dense mass of roots. Stipe 6-10 cm long, slender, reddish, short- 1981] THELYPTERIDACEAE (Holttum) 467 hairy; basal scales 4mm long, narrow; base of Stipe to first large pinna 10-17 cm. Reduced pinnae usually 1-2 pairs, 1-2mm long, rarely lacking. Lamina 20-30cm long, apex not pinna-like; pin- nae 10-15 pairs, lower ones with stalks 1-2 mm long; texture firm, aerophores not enlarged. Lar- gest fertile pinnae 3.5-6cm long, 0.6-0.8 cm wide (sterile to 7.5 x 0.9cm), base narrowly cuneate on basiscopic side, more broadly on acroscopic, apex acuminate, edges lobed 2 way to costa (or more deeply in largest pinnae), lobes oblique; costules 3mm apart, at 45° to costa; veins 3-4 pairs, basal acroscopic vein ending exactly at base of sinus, basal basiscopic vein to edge very near base of sinus; rather sparse stiff hairs to 1mm long on lower surface of rachis, base of costae and mar- gins of lobes, costa distally glabrous, some glands present on lower surface between veins; upper surface of pinnae glabrous apart from slender hairs 0.5 mm long on costae. Sori medial; indusia firm, glabrous; sporangia bearing glands. Distr. Malesia: Borneo; collections several localities. Ecol. On rocks and earth of stream banks, in the flood zone. Note. v.A.v.R. in the Supplement to his Handbook describes the basal veins “‘uniting at or a little below the sinus”, but in the specimens examined by me they usually end separately, though near together, their tips sometimes touch- ing near base of pinnae. from 46. Sphaerostephanos_ uniauriculatus (COPEL.) HOLTTUM, comb. nov.—Dryopteris — uni- auriculata COPEL. Philip. J. Sci. 9 (1914) Bot. 3.— Thelypteris uniauriculata (COPEL.) REED, Phytologia 17 (1968) 322. — Type: C. KING 406, Papua (MICH; BM, P). Caudex short, suberect or creeping. Stipe 20- 27cm long, minutely hairy. Reduced pinnae one on each side of rachis, not opposite, c. 5 mm long. Lamina consisting of a large terminal leaflet and 2 pinnae (not opposite), sometimes a second pair also; aerophores slightly swollen. Apical lamina c. 18 cm long, fertile 3.4-3.7 cm wide, sterile 4.2 cm; base truncate or cordate, edges crenate; costules 5 mm apart along midrib; veins 8 pairs, almost all anastomosing; texture firm. Sterile pinnae to 8 x 3.2cm, widest 1/3 from apex, base broadly roun- ded to truncate, apex short-pointed, edges irre- gularly sinuous; costules 4-4.5 mm apart; veins 6 pairs, 3 distal veins reaching the margin, rest anastomosing; sinus-membrane hardly developed; hairs on lower surface of costae, costules and veins very short, somewhat antrorse, a few short hairs and some glands on surface between veins; upper surface glabrous apart from hairs on costae and costules. Fertile pinnae to 6.5 X2.5cm; sori exindusiate, distal ones medial, lower ones diver- gent, those on basal veins of adjacent costules sometimes confluent; sporangia bearing glands. Distr. Malesia: Papua, known only from type and KING 383 (P). Note. KING 383 includes a young plant which has a cordate-based apical lamina and one pair of pinnae 6 x 8mm, also an older plant with 2 pairs of pinnae and a single reduced pinna. COPELAND stated that there is a group of hairs in place of an indusium, but I did not see this. 47. Sphaerostephanos urdanetensis (COPEL.) HOLTTUM, comb. nov. — Dryopteris urdanetensis COPEL. in Elmer Leafl. Philip. Bot. 5 (1913) 1682. — Cyclosorus urdanetensis (COPEL.) COPEL. Gen. Fil. (1947) 143; Fern Fl. Philip. (1960) 355.— Thelypteris urdanetensis (COPEL.) REED, Phytologia 17 (1968) 322. —Type: ELMER 13762, Mindanao, Mt Urdaneta (MICH). Dryopteris matutumensis COPEL. Philip. J. Sci. 40 (1929) 299, pl. 3.— Cyclosorus matutumensis (COPEL.) COPEL. Gen. Fil. (1947) 143; Fern FI. Philip. (1960) 355.— Thelypteris matutumensis (COPEL.) REED, Phytologia 17 (1968) 291.— Type: COPELAND s.n. May 1917, Mindanao, Mt Matutum, 2000 m (MICH; UC). Caudex short, creeping; base of stipe to first normal pinna 3-5 cm (sterile), 10-18 cm (fertile); reduced pinnae 2-3 pairs, to 4-5 mm long. Lamina excluding reduced pinnae to 20 cm long, consisting of an apical lamina 12-15 cm long, to 3cm wide, deeply lobed and grading to pinnae at its base, with 3-5 pairs of free pinnae; basal pinnae nar- rowed towards their bases on basiscopic side, widest 1/3 from apex. Largest pinnae 1.5 x 0.8 cm (type of D. matutumensis); apex broadly pointed to rounded; edges crenate; costules 2mm apart; veins 2-3 pairs, one pair anastomosing near base of pinna, free elsewhere; lower surface of rachis bearing thick curved pale brown hairs 0.5mm long, hairs on costae shorter, yellow glands present throughout; upper surface of rachis as lower, of pinnae covered throughout with slender appressed hairs. Sori on apical lamina medial or supramedial, on pinnae near costae, one on each basal vein; indusia small, dark, firm, glabrous or with a few glands or short hairs; sporangia bearing glands. Distr. Malesia: Philippines (Mindanao). Note. The type of D. matutumensis has larger fronds than that of D. urdanetensis and has a few hairs on indusia, but agrees in other characters. 48. Sphaerostephanos batjanensis (ROSENST.) HOLTTUM, comb. nov. — Dryopteris batjanensis ROSENST. Meded. Rijksherb. n. 31 (1917) 5.— Thelypteris batjanensis (ROSENST.) REED, Phy- tologia 17 (1968) 263.— Type: DE VRIESE & TEYSMANN 589, Moluccas, Batjan (L). Aspidium canescens forma gymnogrammoides CHRIST, Ann. Jard. Bot. Btzg 15 (1898) J31.— Type: SARASIN 137, N. Celebes, G. Masarang (BAS). 468 Caudex short, erect. Base of stipe to first large pinna of sterile fronds 5-12 cm, of fertile to 30 cm. Reduced pinnae 1-2 pairs, very small, irregularly spaced. Lamina to 30 cm long; pinnae 12-15 pairs, basal 2-3 pairs deflexed and slightly reduced, lowest narrowed towards base on basiscopic side. Sterile pinnae to 3.5cm long, 1.2cm wide above base which ts a little dilated and sometimes auri- cled on acroscopic side; apex abruptly short- pointed; edges lobed to a depth of 1.5mm; cos- tules 3 mm apart; veins to 6 pairs, pale and prom- inent on lower surface, 13 pairs anastomosing, next pair to sinus-membrane; lower surface of rachis, costae and costules covered with ap- pressed pale hairs 0.3-0.4 mm long, copious glands on surface between veins; upper surface of pinnae covered throughout with slender appressed hairs 0.3 mm long, no glands; hairs on rachis more than 0.5mm long. Fertile pinnae a little smaller than sterile; sori almost covering lower surface; in- dusia firm, rather large, hairy; sporangia bearing glands. Distr. Malesia: N. Celebes & Moluccas (Bat- jan, Ceram). Note. The specimen of DE VRIESE & TEYS- MANN 589 at Kew is labelled Ceram, not Batjan: it agrees with the Leiden specimen. 49. Sphaerostephanos humilis HOLTTUM, sp. nov. Pinnae 5-jugatae, usque 1.6 = 1.1 cm, inferiores non decrescentes, subtus ubique glanduliferae, supra pilis adpressis vestitae; sori mediales;: in- dusia magna, glandulis pilisque brevibus praedita: sporangia copiose’ glandulifera. —Type: W. MEER 9826, Central Celebes, Mt Nokilalaki, 1500-2000 m (L). Caudex short-creeping; stipe of sterile fronds 4-5 cm, of fertile 6-13 cm, short-hairy throughout, basal scales thin, c. 3x 1.5mm. Sterile frond to c. 14cm long, consisting of an apical lamina 8.5 cm long, 1.2cm wide, lobed to a depth of 2-2.5 mm, and 5 pairs of pinnae, basal pinnae variably somewhat reduced, no very small reduced pinnae present; largest pinnae 1.6 1.1cm, subsessile, widening from the base to widest part near apex, irregularly shallowly lobed (most deeply towards apex) with 3-4 lobes on each side: costules 3- 5mm apart; veins to 4 pairs, basal pair anas- tomosing, next pair to margin; lower surface of rachis bearing rather sparse pale hairs 0.4mm long, costae covered sparsely with very short ap- pressed hairs, glands copious throughout lower surface which is not pustular; upper surface of rachis hairy as lower, whole upper surface of pinnae covered with appressed hairs 0.3 mm long. Fertile fronds with apical lamina a little narrower than sterile, edges of lobes dentate at ends of veins; pinnae to 1.4 0.9cm:; sori medial; indusia rather large, firm, with a few glands and usually a few short hairs; sporangia copiously glandular; spores spinulose. FLORA MALESIANA [ser. II, vol. 1° Distr. Malesia: Central Celebes. Only known from the type. 50. Sphaerostephanos lastreoides (PRESL) HOLT- TUM, Kalikasan 4 (1975) 54.— Pronephrium las- treoides PRESL, Epim. Bot. (1851) 259: Holttum, Novit. Bot. Univ. Carol. Prag. 1968 (1969) 49. Type: “Java (JUNGHUHN, comm. Vriese 1850)” (PRC). Cyclosorus duplosetosus COPEL. Philip. J. Sci. 81 (1952) 32; Fern Fl. Philip. (1960) 354. — Thelypteris duplosetosus (COPEL.) REED, Phy- tologia 17 (1968) 274.—Type: MERRILL 9530, Palawan, Mt Capoas (MICH). Caudex short, creeping. Stipe to 5cm (sterile), to 15cm (fertile) covered with short pale hairs; base of stipe to first large pinna 10-20cm. Reduced pinnae 1-2 pairs, small, irregularly spaced. Lamina to 15cm long; free pinnae c. 10 pairs, lower ones with short stalks and narrowed at base on basiscopic side; apical lamina short, acuminate; aerophores not elongate. Supramedial pinnae to 2.3x0.9cm, base truncate, apex ab- ruptly obtuse-pointed, lobed to about 3 way to costa; costules 3 mm apart; veins 3-4 pairs, basal pair anastomosing with excurrent vein to sinus, next pair to edge; sparse pale spreading hairs 1 mm long and many much shorter ones on lower surface of costae and costules, glands on surface between veins; scattered long hairs on upper sur- face of costae, costules and veins, whole surface covered with slender appressed hairs 0.2-0.3 mm long. Rachis also with spreading hairs 1 mm long and shorter ones on both surfaces. Sori medial; indusia with short hairs; sporangia bearing glands. Distr. Malesia: Philippines (Palawan, Luzon, Negros, Mindanao). Notes. Part of CUMING 251 from Luzon, in several herbaria, exactly matches the type of this species; nothing like it is otherwise known from Java. Probably the type was a CUMING specimen misplaced by PRESL. M. G. PRICE 2734, from Mindanao, Agusan del Sur Prov., is like the type in all essentials but a little larger; its fertile pinnae are the same size as sterile pinnae on the Kew specimen of CUMING 251, which has smaller fer- tile ones. M. G. PRICE 690, from Mt Halcon, Mindoro, differs from the above description only in larger size: lamina to 35 cm long, pinnae 20 pairs, largest 7x1.7cm. It may represent a distinct local variety. A. LOHER s.n. March 1915, from Rizal Pro- vince, Luzon, differs in having shallowly crenate pinnae, and slender appressed hairs on lower sur- face of costules. 51. Sphaerostephanos cataractorum (WAGN. & GRETH.) HOLTTUM, comb. nov. — Cyclosorus cataractorum WAGN. & GRETH. Un. Cal. Publ. Bot. 23 (1948) 50, pl. 16. — Thelypteris catarac- 1981] THELYPTERIDACEAE (Holttum) 469 torum (WAGN. & GRETH.) REED, Phytologia 17 (1968) 266. — Type: GRETHER & WAGNER 3971, Admiralty Islands, Manus I. (UC; NSW). Caudex short, erect. Stipe 10-20 cm long, pallid; basal scales narrow, 5mm long. Lamina to 30cm long; pinnae to 16 pairs; basal pinnae reduced, sometimes one or both to a very narrow leaflet less than |1cm long; apex of frond sometimes almost pinna-like; texture thin; aerophores not elongate. Largest pinnae to 10x 0.9cm, widest at middle, base very narrowly cuneate, apex acu- minate, edges lobed } way to costa, lobes very oblique, slightly falcate, acute; costules 44.5 mm apart, at 45°; veins 5-6 pairs, basal pair anas- tomosing with excurrent vein to sinus, next acroscopic vein to side of short sinus-membrane; lower surface of pinnae quite glabrous with resinous pale glands between veins; upper surface short-hairy on costae, otherwise glabrous. Rachis glabrous on lower surface or with very short hairs; hairs on upper surface thick, brown, curved, 0.3-0.4mm long. Sori medial; indusium firm, glabrous; sporangia bearing glands. Distr. Admiralty Islands, New _ Britain, Bougainville, in Malesia: Eastern New Guinea. Ecol. By streams in flood-zone at altitudes to 250 m. Note. This is very similar to S. hispidifolius of Borneo, but has veins always anastomosing and almost glabrous pinnae. The resinous glands on lower surface are usually flattened when dry, not spherical as normally in this genus. 52. Sphaerostephanos menadensis HOLTTUM, sp. nov. Pinnae redactae 2-jugatae, minutae; lamina 20 cm longa, pars apicalis angusta 10cm longa; pinnae 6-jugatae, infimae basin versus valde angustatae, mediales 5.5 x 1.6 cm, dimidio costam versus lobatae; costae subtus pilis minutis ves- titae; pagina inter venas subtus glandulifera, supra eglandulosa; indusia glabra, sporangia setifera.— Type: KOORDERS' 17133, Celebes, Menado, 1800-2000 m (L). Base of stipe to first large pinna 25 cm. Reduced pinnae 2 pairs, very small and widely spaced. Terminal lamina 10cm long, narrowly deltoid; free pinnae 6 pairs, lowest deflexed and gradually much narrowed towards their bases, widest above middle; aerophores not elongate. Middle pinnae to 5.5 x 1.6 cm, base truncate, apex abruptly caudate, edges lobed 3 way to costa, lobes falcate; costules 3.5mm apart, at more than 60°; veins 6 pairs (fertile) 7-8 pairs (sterile), 1 pair anastomosing, next pair of the acroscopic one only to sinus- membrane; long coarse hairs on lower surface of rachis, very short ones on costae and costules, surfaces between veins bearing glands only; hairs on upper surface of costules and veins all very short, neither hairs nor glands between veins. Sori medial; indusia glabrous; sporangia bearing glands. Distr. Malesia: from the type. N.E. Celebes; only known 53. Sphaerostephanos subcordatus HOLTTUM, sp. nov. Pinnae redactae unijugatae, parvae; frondes dimorphae; pinnae usque 8-jugatae, steriles 3 1.2cm, fertiles 1.7X0.9cm, usque 1/3 costam versus lobatae; costae subtus brevi-pilosae, pagina inter venas glabra glandulis paucis praedita; pagina superior inter venas glabra, eglandulosa. indusia parva, glabra; sporangia glandulosa — Type: M. G. PRICE & B. F. HER- NAEZ 61, Western Samar (K). Caudex short, creeping, 3mm diameter when dry. Stipe of sterile fronds 5-7 cm, of fertile 15- 18 cm to first large pinna below which are 2 much- reduced pinnae, not opposite. Lamina 15 cm long, apex broadly deltoid and deeply lobed, free pinnae to 8 pairs, basal ones slightly reduced, defiexed, with stalks 1mm long and asymmetric base; aerophores not elongate. Largest sterile pinnae 3 x 1.2cm, in some cases 1.5 cm wide near apex; base truncate to subcordate; apex abruptly blunt- pointed; edges lobed to a depth of 2mm, more deeply near apex if it is widened, lobes rounded; costules 4mm apart, at less than 60°; veins 4-5 pairs, basal pair anastomosing, next acroscopic vein to sinus-membrane; hairs on lower surface of costae 0.2mm _ long, antrorsely curved, more sparse and shorter on costules, surface between veins glabrous with a few glands; hairs on upper surface of costae sparse, 0.4mm long, similar or longer ones scattered on costules and veins; sur- face between veins glabrous. Hairs on lower sur- face of rachis 0.3-0.4 mm long, thick and curved, on upper surface rather sparse, | mm long. Sori medial, lower ones sometimes a little elongate; indusia small, glabrous; sporangia with glands. Distr. Malesia: Philippines (W. Samar), only known from the type. Ecol. On limestone-derived soils, in forest. 54. Sphaerostephanos novoguineensis (BRAUSE) HOLTTUM, comb. nov.—Dryopteris novo- guineensis BRAUSE, Bot. Jahrb. 49 (1912) 21; v.A.v.R. Handb. Suppl. (1917) 159.— Lastrea novoguineensis (BRAUSE) COPEL. Gen. Fil. (1947) 139; Philip. J. Sci. 78 (1951) 426. — Thelyp- teris novoguineensis (BRAUSE) REED, Phytologia 17 (1968) 297. — Type: SCHLECHTER 17719, N.E. New Guinea, Kani Mts, 1000 m (B; P). Dryopteris glaucescens BRAUSE, Bot. Jahrb. 56 (1920) 85; CoPEL. Philip. J. Sci. 78 (1951) 426. — Thelypteris glaucescens (BRAUSE) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 251. — Type: LEDERMANN 13034, N.E. New Guinea, Sepik Dist. 1400 m (B). Caudex short, creeping. Stipe 10-15 cm long; basal scales thin, 10 x 2.5 mm; base of stipe to first large pinna 20-45 cm. Reduced pinnae 3-8 pairs, 470 FLORA MALESIANA [ser. II, vol. 1° not more than 3mm long. Lamina to 80cm long; pinnae to 25 pairs, lower ones not narrowed at base; aerophores to more than I mm long. Pinnae somewhat dimorphous, sterile to 10 x 1.7 cm, fer- tile to 7.5 x 1.2 cm; base truncate, basal acroscopic lobe to 4mm longer than next; apex short-acu- minate; edges lobed to less than | mm from costa, lobes falcate; costules 3 mm apart (sterile), 2.5 mm (fertile); veins 10 pairs, basal acroscopic vein passing to side of sinus-membrane, basiscopic vein to edge above it; lower surface of costae bearing many short hairs and less abundant long ones to Imm (long hairs sometimes lacking), similar hairs on costules, glands sometimes few present on surface between veins; upper surface with hairs | mm long scattered on costae, costules and veins, surface between veins + closely covered with fine appressed hairs. Sori medial to supramedial; indusia large, firm, with short hairs and sometimes glands; sporangia bearing glands; spores finely spinulose. Distr. Malesia: Eastern New Guinea, several collections, in forest at 850-1400 m. Note. The type of D. glaucescens differs from that of D. novoguineensis in the absence of long hairs on lower surface of rachis and costae and the presence of glands on indusia. 55. Sphaerostephanos hispiduliformis (C. CHR.) HOLTTUM, comb. nov.— Dryopteris hispiduli- formis C. CHR. Ind. Fil. Suppl. III (1934) 88, new name for Dryopteris hispidula BRAUSE, Bot. Jahrb. 56 (1920) 102, non (Sw.) O. KTZE 1891. — Thelyp- teris hispiduliformis (C. CHR.) REED, Phytologia 17 (1968) 283.—Type: LEDERMANN 11758, N.E. New Guinea, Sepik Distr. 2070 m, in forest (B). KEY TO THE VARIETIES 1. Caudex to 150. cm tall; lower surface of pinnae lacking glands between veins a. var. hispiduliformis 1. Caudex short; lower surface of pinnae between veins copiously glandular. . Fronds to 90cm long; pinnae to 14 1.5 cm; hairs on lower surface of costae 0.1 mm long b. var. vinkii 2. Fronds to 40 cm long; pinnae to 5 x | cm; hairs on lower surface of costae 1 mm long c. var. brassii nN a. var. hispiduliformis Caudex erect, to 150cm tall; stipe 10-15cm long, basal scales broad, thin; base of stipe to pinnae of maximum size 60cm or more, whole frond to 150cm long; reduced pinnae 1.5-2.0cm apart, broadly triangular, c. 12 pairs gradually increasing upwards from very small basal ones to lcm, then 7-12 pairs grading to pinnae of maxi- mum size; aerophores more than | mm_ long. Largest pinnae 11cm long, to 1.7cm wide at dilated base, rather evenly tapered to apex, lobed c. 2/5 towards costa; costules to 3.5mm apart, at more than 60° to costa; veins 7-9 pairs, 12 pairs anastomosing, 1-15 pairs ending beside sinus- membrane; lower surface of costae with spreading pale hairs 1mm long and shorter ones, hairs on costules more sparse, a few short hairs between veins, glands confined to costules and veins, sometimes lacking; hairs on upper surface of cos- tae more than | mm long, scattered similar hairs on costules and veins, fine appressed hairs all over surface. Sori medial; indusia bearing hairs and glands, sporangia with neither; spores covered with very small wings. Distr. Malesia: Eastern mountain forest at c. 2000 m. New Guinea, in b. var. vinkii HOLTTUM, var. nov. A typo speciei differt: caudice breve; pinnis 1/3 costam versus lobatis; costis subtus pilis minutis erectis vestitis; pagina inferiore inter venas copiose glandulifera.— Type of variety: VINK 17605, N.E. New Guinea, W. Sepik Distr., in low secondary growth on limestone (L). Distr. Malesia: Papua New Guinea (Sepik). c. var. brassii HOLTTUM, var. nov. — Dryopteris strigosissima COPEL. Univ. Cal. Publ. Bot. 18 (1942) 221. — Cyclosorus strigosissimus (COPEL.) COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78 (1951) 456, pl. 36.— Thelypteris strigosissima (COPEL.) REED, Phytologia 17 (1968) 316.— Type: BRASS 11436, New Guinea, Bele River, 2200 m, on limestone cliff in shade (MICH). Differs from var. vinkii: fronds much smaller; reduced pinnae contiguous, lower ones wider than long; largest pinnae 5 x | cm, crenate to a depth of 1mm; long hairs present on lower surface of costae. Distr. Malesia: Papua New Guinea. Notes. The greater number of glands on the lower surface may be due to the limestone habitat of var. vinkii and var. brassii (plants of S. heterocarpus on limestone are densely glandular), also their short caudex. But there is no evidence that LEDERMANN’s type did not grow on limes- tone. When more information is available it may be better to regard var. vinkii and var. brassii as representing a distinct species, for which COPELAND’s epithet strigosissimus would be available. 56. Sphaerostephanos hastatopinnatus (BRAUSE) HOLTTUM, comb. nov. — Dryopteris hastatopin- nata BRAUSE, Bot. Jahrb. 56 (1920) 112.— Thelypteris hastatopinnata (BRAUSE) REED, Phytologia 17 (1968) 281. — Type: LEDERMANN 8237, N.E. New Guinea, Sepik Distr., 200m on rocks in stream-bed in forest (B; K). Plants varying much in size according to habi- tat. Caudex erect, short. Stipe 3-4cm on type, to 1981] THELYPTERIDACEAE (Holttum) 471 15cm on large plant. Lamina of type 45 cm long, of large plant 130cm; pinnae 28 pairs on type, 35 pairs on large plant; 10-20 pairs lower pinnae in all cases gradually smaller, lowest 5S-10mm_ long; apex of frond almost pinna-like; aerophores dis- tinctly swollen, to almost 1mm long in large fronds. Reduced pinnae broadly and symmettric- ally triangular, spreading, subentire. Largest pin- nae of type 10X1.3cm, of a large plant 20x 3.0cm; base truncate, dilated and more or less auricled both sides (upper ones auricled on acroscopic side only), apex short-acuminate, edges sinuous or slightly crenate; costules of small plants 2.5-3 mm apart, of a large one to 4.5 mm, at more than 60°; veins 3-7 pairs, almost all anas- tomosing; sinus-membrane hardly developed; lower surface of pinnae quite glabrous or with a few hairs on costae, glands sometimes present on and between veins; hairs on upper surface of costae slender, brown, to 0.5mm _ long, none elsewhere, a few glands rarely present. Lower surface of rachis glabrous or with sparse ap- pressed hairs, upper surface bearing copious red- brown hairs | mm long. Sori medial, lower ones + elongate and sometimes confluent; no indusia; sporangia bearing glands. Distr. Malesia: Central Celebes, Moluccas (Ceram) and widely distributed in New Guinea. Ecol. On river banks in forest at low altitudes, up to 650m. The type was probably stunted owing to its rocky habitat and perhaps exposed position. 57. Sphaerostephanos_ latebrosus (KUNZE ex METT.) HOLTTUM in Nayar & Kaur, Comp. to Bedd. (1974) 209. — Aspidium latebrosum KUNZE ex METT. Farngatt. IV (1858) 104; BAK. Syn. Fil. (1867) 294; MIQUEL, Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 159.— Dryopteris latebrosa (METT.) C. Cure. Ind. Fil. (1905) 274; v.A.v.R. Handb. (1908) 221. — Thelypteris latebrosa (METT.) REED, Phy- tologia 17 (1968) 287. — Type: ZOLLINGER 354 pt, Java (L, n. 908, 333-517 only). Nephrodium glaucostipes BEDD. Handb. Suppl. (1892) 80.— Dryopteris glaucostipes (BEDD.) C. Cure. Ind. Fil. (1905) 268; Gard. Bull. Str. Settl. 4 (1929) 389; v.A.v.R. Handb. (1908) 226. — Cyclo- sorus heterocarpus var. glaucostipes (BEDD.) HOLTTuM, Rev. Fl. Mal. 2 (1955) 271. — Thelyp- teris heterocarpus var. glaucostipes (BEDD.) REED, Phytologia 17 (1968) 282. — Type: KING’s Collector (KUNSTLER) 2046, Perak, Larut (K; CAL, SING). Dryopteris todayensis CHRIST, Philip. J. Sci. 2 (1907) Bot. 193; v.A.v.R. Handb. Suppl. (1917) 184. — Cyclosorus todayensis (CHRIST) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 249; COPEL. Fern FI. Philip. (1960) 344. — Thelypteris todayensis (CHRIST) REED, Phytologia 17 (1968) 319. — Sphaerostephanos todayensis (CHRIST) HOLTTUM, Kalikasan 4 (1975) 58.—Type: COPELAND 1463, Mindanao, Mt Apo 1220 m (P; WiSiB): Dryopteris angustipes COPEL. Philip. J. Sci. 7 (1912) Bot. 60; v.A.v.R. Handb. Suppl. (1917) 184.— Thelypteris angustipes (COPEL.) REED, Phytologia 17 (1968) 260.— Type: BROOKS 110, Sarawak (MICH; BM). Cyclosorus sagittifolioides COPEL. Philip. J. Sci. 81 (1952) 29, 30, pl. 21; Fern FI. Philip. (1960) 344. — Thelypteris sagittifolioides (COPEL.) REED, Phytologia 17 (1968) 311. —Type: EDANO BS 24832, Samar, Catubig River (US). Cyclosorus halconensis COPEL. Philip. J. Sci. 81 (1952) 29; Fern Fl. Philip. (1960) 345.— Thelypteris halconensis (COPEL.) REED, Phy- tologia 17 (1968) 305. — Type: EDANO PNH 3849, Mindoro, Mt Halcon (MICH; L). Cyclosorus polypterus COPEL. Philip. J. Sci. 84 (1955) 161.—Thelypteris polyptera (COPEL.) REED, Phytologia 17 (1968) 305. — Type: EDANO 7852 (PNH 21373), Negros, Canlaon Volcano 650 m (MICH). Caudex erect, not or little branched at base. Stipe 6-12cm long; base of stipe to first large pinna 45-70 cm. Reduced pinnae 10-15 pairs, 2.5— 4 cm apart, lowest commonly 5 mm long, gradually increasing upwards to 2.5-3.cm, larger ones nar- rowly triangular, somewhat deflexed, with acros- copic auricle, edges incised, apex broad-pointed; transition to large pinnae subabrupt; aerophores elongate, in some cases | mm or rarely more (type of C. sagittifolioides). Lamina above reduced pinnae to 60cm or more long; basal large pinnae not narrowed at base. Rachis lower surface commonly quite glabrous, in some specimens with sparse long hairs or some very short ones; hairs on upper surface to | mm long, usually appressed (spreading in some Philippine specimens). Largest pinnae 12 X 1.6 to 20 x 2.0 cm; base truncate, usu- ally a little dilated both sides; apex acuminate; edges lobed about half-way to costa or a little more in large pinnae; costules 3.5-4.5 mm apart; veins 7-8 pairs, basal pair anastomosing, one or both of next pair to sinus-membrane; lower sur- face of costae usually glabrous at base with very short antrorse hairs distally, in some cases very short hairs and a few somewhat longer throughout; costules as costae but with shorter hairs (if any); rest of lower surface usually quite glabrous with many glands; upper surface with long hairs on costae, similar hairs scattered on costules and veins, rest of surface bearing a vari- able number of fine appressed short hairs. Sori medial, lower ones not divergent; indusia glabrous or with a few short hairs, usually also glands; sporangia bearing glands, rarely a seta; spores with many very small wings. Distr. Malesia: Western Malesia, Philippines. Ecol. In forest, low altitudes to 800 m. Notes. In Malaya, Sumatra and Java all lower surfaces are almost glabrous, but in nearly all 472 cases there are short hairs on distal parts of costae; aerophores are little elongate. In Sabah (Mt Kinabalu) and the Philippines some hairs are nearly always present and aerophores are more developed. In the type of Cyclosorus sagit- tifolioides some aerophores are 2mm long, and short erect hairs are present on lower surface between veins. Some Philippine specimens seem rather inter- mediate between this species and S. hirsutus; they may be hybrids. Hybrids with S. heterocarpus in Sabah are also possible. The solitary erect caudex of S. latebrosus, large upper reduced pinnae and broad base of lowest normal pinnae seem dis- tinctive. 58. Sphaerostephanos porphyricola) (COPEL.) HOLTTUM, Kalikasan 4 (1975) 59.— Dryopteris porphyricola COPEL. Philip. J. Sci. 7 (1912) Bot. 60; v.A.v.R. Handb. Suppl. (1917) 186. — Thelyp- teris porphyricola (COPEL.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 6 (1936) 287, nomen tantum. — Cyclosorus porphyricola (COPEL.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 248; HOLTTUM, Rev. FI. Malaya 2 (1955) 271. — Type: BROOKS 112, Sarawak, Bau (MICH; BM). Dryopteris angustipes sensu. C. CHR. Gard. Bull. Str. Settl. 4 (1929) 389. Caudex erect, short. Stipe 10cm long; base of stipe to first large pinna 35 cm. Reduced pinnae to 12 pairs, 1.5cm apart, uppermost I-1.5 cm long, deflexed, narrowly triangular with auricle on acroscopic base, edges incised, lowest ones 3- 5mm long. Lamina 55 cm long; pinnae to 20 pairs or more, lowest ones narrowed a little at base on basiscopic side only; aerophores elongate, to less than 1mm long; young fronds covered with mucilage. Lower surface of rachis almost glabrous or with very short hairs, also many glands; upper surface copiously hairy. Largest pinnae 14x 2.5 cm (sterile to 15 x 3 cm), base sub- truncate, apex abruptly short-acuminate; edges lobed 2/5 to costa, lobes distinctly falcate with forward-pointing tips; costules 3-4.5mm_ apart; veins 7-9 pairs, | pair anastomosing, next pair to sinus-membrane; lower surface of costae, costules and veins covered with fine short closely ap- pressed hairs, also glands; surface between veins bearing copious glands and a variable number of short fine appressed hairs; hairs on upper surface of costae antrorse, pale, similar hairs sometimes scattered on costules and veins, surface between veins typically covered closely with slender ap- pressed hairs 0.5mm long, fewer such hairs on specimens from Malaya. Sori medial, lower ones not divergent but sometimes a little elongate; in- dusia bearing short hairs and glands; sporangia with glands. Distr. Malesia: Malaya and Borneo (Sarawak; Sulu Arch.: Tawi-Tawi). Ecol. In forest at low altitudes. FLORA MALESIANA [ser. II, vol. 1° Note. This is closely allied to S. latebrosus; more field observations are needed. 59. Sphaerostephanos caulescens HOLTTUM, sp. nov. — Dryopteris porphyricola (non COPEL.) C. Cur. Gard. Bull. Str. Settl. 7 (1934) 244. Caudex erectus; stipes Scm longus; pinnae redactae usque 8-jugatae, omnes minutae; aero- phora |mm longa; pinnae majores 13.5 Xx 1.5- 2.0cm, dimidio costam versus lobatae; rachis subtus fere glabra; pagina inter venas subtus glandulifera, supra pilis appressis vestita; indusia fere glabra; sporangia glandulifera. — Type: G. P. LEWIS 284, Sarawak, G. Mulu (K). Caudex erect, to at least 20cm tall. Stipe 5 cm long, base covered with large thin scales; base of stipe to first large pinna 40 cm. Reduced pinnae to 8 pairs, when young covered with mucilage, all very small, with aerophores |mm long. Lower surface of rachis often bearing narrow scales, otherwise almost glabrous, upper surface with hairs Imm long closely appressed, with some longer and spreading. Lamina to 75 cm long; pin- nae to more than 30 pairs, not opposite; basal pinnae (at least sterile ones) narrowed at their bases. Largest fertile pinnae 13.5 X 2.0 cm (sterile sometimes larger, to 20*3cm); base truncate; apex acuminate (sometimes with cauda 1.5 cm long); edges lobed about 3 way to costa; costules c. 3.5mm apart, at 60°; veins c. 7 pairs, 1 pair anastomosing, next pair to sides of sinus-mem- brane; lower surface of costae almost glabrous at base, with appressed hairs to 0.2 mm long distally, costules similar, surfaces between veins bearing copious glands; hairs on upper surface of costae to 1mm long, similar hairs scattered on costules and veins, short appressed hairs on surface be- tween veins. Sori medial, lower ones not diver- gent; indusia glabrous or with few very short hairs; sporangia bearing glands. Distr. Malesia: Borneo (N. Sarawak; Sabah), N. Celebes, and Moluccas (Batjan). Ecol. In forest near streams at 500-1500 m. Note. This is near S. magnus, but pinnae of S. caulescens (except largest sterile omnes) are generally smaller and less deeply lobed, with different pubescence and glandular sporangia. 60. Sphaerostephanos reconditus HOLTTUM, sp. nov. Pinnae redactae c. 6-jugatae, parvae; pinnae normales usque 7.2 X 1.5 cm, dimidio costam ver- sus lobatae; venae 5-6-jugatae, infimae solum anastomosantes; costulae subtus pilis adpressis vestitae; pagina inferior inter venas pilis adpressis glandulisque praedita; sori mediales; indusia dense pilosa; sporangia copiose glandulifera. — Type: B. S. PARRIS 6872, Sarawak, Gunong Mulu (CGE). Caudex short, erect or suberect; stipe Scm long; base of stipe to first normal pinna 30cm; 1981] THELYPTERIDACEAE (Holttum) 473 reduced pinnae all very small, 3-4cm apart. Lamina excluding reduced pinnae 30cm long; pinnae c. 12 pairs; basal pinnae much narrowed in basal third, above base 2cm wide; aerophores when dried barely 0.5mm long, on living plant “vellow’’. Lower surface of rachis densely hairy, hairs thick, curved, brown, | mm long with shorter pale ones (brown hairs more numerous on basal part); upper surface with thicker brown hairs and shorter pale ones. Suprabasal pinnae to 7.2 1.5cm; base truncate and slightly auricled; apex short-acuminate; edges lobed not quite half-way to costa; costules 3.5-4.5 mm apart, at c. 60°; veins 5-6 pairs, hardly prominent, | pair anastomosing, next acroscopic vein or pair passing to sinus- membrane; lower surface of costae bearing pale antrorse hairs which are appressed distally, 0.5mm long, also glands, costules covered with slender appressed hairs, between veins some ap- pressed hairs and rather sparse glands; upper sur- face of costae bearing pale hairs, no long hairs on costules or veins, between veins sparse appressed hairs. Sori medial or a little inframedial, lower ones not divergent; indusia densely hairy, some- times with a gland; sporangia bearing many glands. Distr. Malesia: Borneo (Sarawak: Mt Mulu), only known from type. Ecol. In forest clearing in Hidden Valley at 450 m; very young fronds covered with mucilage. 61. Sphaerostephanos cyrtocaulos (v.A.v.R.) HOLTTUM, comb. nov. — Dryopteris cyrtocaulos v.A.v.R. Bull. Jard. Bot. Btzg III, 5 (1922) 201. — Thelypteris cyrtocaulos (v.A.v.R.) REED, Phy- tologia 17 (1968) 270. — Lectotype (here selected): BUNNEMEIJJER 9922, Sumatra, G. Kerinci, 2100 m (BO; L). Caudex erect. Stipe 10-15cm long; base of stipe to first large pinna 40-75 cm. Reduced pinnae 8 pairs or more, 2-3 mm long; aerophores to 2mm long. Lamina to 75 cm long; pinnae more than 20 pairs, well spaced; lowest pinnae not narrowed at base, nor auricled. Hairs on both sides of rachis short. Largest pinnae of lectotype 12x2.5cm (possibly larger on sterile fronds); base truncate; apex acuminate with cauda to 1.5 cm; edges lobed to 2.5-3 mm from costa (about 3/4), lobes slightly falcate; costules 44.5 mm apart at more than 60°; veins 8-11 pairs, basal pair anastomosing, next pair both to edge above base of sinus; hairs on lower surface of costae and costules 0.2 mm long, appressed, a few also on veins, glands present on and between veins; hairs on upper surface of costae 1 mm long, antrorse, similar hairs scattered on costules and veins, pale appressed hairs over whole surface. Sori rather supramedial; indusia firm, rather large, with short hairs; sporangia bearing glands; spores densely short-spinulose. Distr. Malesia: Sumatra (on and near G. Kerinci), 1800-2300 m, in forest. Note. VAN ALDERWERELT cited three col- lections by BUNNEMEIJER without indicating a type; the largest one is here selected. One other collection is known from a neighbouring locality. 62. Sphaerostephanos baramensis (C. CHR.) HOLTTUM, comb. nov. — Dryopteris baramensis C. CHR. Gard. Bull. Str. Settl. 7 (1934) 246. — Thelypteris baramensis (C. CHR.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 6 (1936) 287. — Type: MJOBERG 5 partim, Sarawak, Baram Valley, 900— 1200 m (BM). — Fig. 13f-g. Caudex short, erect. Stipe 5-10 cm long, basal scales thin, to 2mm wide at base; base of stipe to first large pinna 40-60 cm. Reduced pinnae to 10 pairs, 4-6 cm apart, c. 2mm long, with aerophores c. 1mm long. Lamina to at least 70cm long; pinnae to 20 pairs, lower ones often opposite, upper ones usually not; 3-4 pairs lower pinnae much narrowed towards their bases; middle pin- nae often distinctly stalked; aerophores 1-2 mm long. Lower surface of rachis glabrous; hairs on upper surface less than | mm long, brown, ap- pressed. Largest pinnae commonly 15 x2.5cm, largest on type 24 4cm; apex of wider pinnae rather abruptly caudate-acuminate; edges lobed to 3mm from costa; lobes falcate; costules com- monly 4mm apart, on large fronds to 7 mm; veins 10-12 pairs, prominent on lower surface, tips of basal veins touching sides of short sinus-mem- brane or sometimes uniting just below the sinus to form a short excurrent vein; lower surface of costae and costules covered with antrorsely cur- ved hairs 0.2-0.3mm long, those on costules closely appressed; surface between veins bearing a variable number of small glands and short ap- pressed hairs; upper surface of costae bearing pale antrorse hairs less than 1 mm long, hairs on costules much shorter, surface between veins of type covered with short appressed hairs, of other specimens glabrous or with appressed hairs near costa only. Sori a little supramedial; indusia rather large, firm, glabrous or with short hairs; sporangia sometimes with a small gland, not setiferous. Distr. Malesia: Borneo (Northern Sarawak, E. Kalimantan and Sabah), in forest at 900-1500 m. 63. Sphaerostephanos batulantensis HOLTTUM, sp. nov. Caudex brevis, repens; stipes 10cm longus; pinnae redactae usque 10-jugatae, superiores 4— 5mm longae; pinnae maximae 111.5 cm, 2/3 costam versus lobatae; costulae subtus pilis bre- vibus appressis vestitae, pagina inter venas glan- dulifera; pagina superior omnino pilis appressis vestita; sporangia glandulifera.— Type: Kos- TERMANS 18850, W. Sumbawa, Mt Batulante, 900 m (L; BO, K). Caudex short, prostrate, c. 5 mm diameter. Stipe 10 cm long; base of stipe to first large pinna 40 cm or more. Reduced pinnae to 10 pairs, upper ones 474 4-5mm long. Lamina 45cm long; lowest pinnae narrowed to a base 5 mm wide; aerophores hardly 1mm long. Lower surface of rachis densely covered with pale erect hairs 0.3mm long, hairs on upper surface to |mm long, pale. Largest pinnae 11 x 1.5 cm; base subtruncate, not auricled; apex acuminate; edges lobed 2/3 to costa or a little more deeply, lobes falcate; costules 3-4 mm apart, at more than 60°; veins 8-9 pairs, basal pair anas- tomosing, next acroscopic vein sometimes to sinus-membrane; lower surface of costae hairy as rachis, hairs on distal part longer and antrorse; lower surface of costules covered with short ap- pressed hairs; surface between veins bearing glands and a few short erect hairs; upper surface covered entirely with fine appressed hairs, no long hairs on costules or veins. Sori a little inframedial, lower ones not divergent; indusia thin with many hairs 0.3 mm long; sporangia bearing glands. Distr. Malesia: Lesser Sunda Is. (W. Sum- bawa: Mt Batulante, 900 m); only known from the type. 64. Sphaerostephanos subalpinus (v.A.v.R.) HOLTTUM, comb. nov.— Dryopteris subalpina v.A.v.R. Bull. Jard. Bot. Btzg III, 5 (1922) 200. — Thelypteris subalpina REED, Phytologia 17 (1968) 317. Type: BEGUIN 1496, Ternate, N. Formad- jahi, 1200 m (BO; L). Polypodium acutum ROXB. Calc. J. Nat. Hist. 4 (1844) 492, non BURM. f.; MORTON, Contr. U.S. Nat. Herb. 38 (1974) 335.—Type: ‘““Amboina” (probably C. SMITH, Ceram). Aspidium hispidulum DECNE var. ternatense Miq. Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 160. — Type: TEYSMANN, Ternate (L). Caudex short, creeping. Stipe 10cm long; base of stipe to first large pinna 30-35cm. Reduced pinnae 1.5-2.0cm apart, c. 10 pairs, upper ones 1.5cm long, deflexed, narrowly triangular, deeply lobed, with basiscopic auricle to 10mm _ long, lowest 5mm long. Lamina 60-70 cm long; pinnae to 30 pairs, basal ones not narrowed at base; aerophores c. | mm long. Lower surface of rachis covered with very short hairs and sparse longer ones to | mm long; hairs on upper surface pale or brownish, to 1mm long. Largest pinnae to 15 x 2.0cm, base subtruncate and a little dilated both sides; apex caudate-acuminate; edges lobed to 2mm from costa, lobes falcate, narrowed above their bases; costules to 4mm apart; veins 8-10 pairs, basal pair anastomosing near base of pinna, near apex both touching sides of short sinus- membrane; hairs on lower surface of costae near base 0.2-0.3 mm long, suberect, distally to 0.7 mm, hairs on costules. similar, glands present throughout lower surface; hairs on upper surface of costa mostly less than 1 mm long, with scat- tered longer ones, similar longer hairs scattered on costules and veins; a few appressed hairs some- times present between veins. Sori a little FLORA MALESIANA [ser. II, vol. 1° supramedial, basal ones not divergent; indusia fairly large, with a few short hairs and sometimes glands; sporangia with 1-2 glands. Distr. Malesia: Moluccas (Ternate, collections; Halmahera; Ceram?). several 65. Sphaerostephanos lobatus (COPEL.) HOLT- TUM, Kalikasan 4 (1975) 64. — Cyclosorus lobatus (COPEL.) COPEL. Philip. J. Sci. 81 (1952) 33; Fern Fl. Philip. (1960) 356.— Thelypteris lobata (COPEL.) REED, Phytologia 17 (1968) 289.— Type: COPELAND 1966, Luzon, Baguio (MICH). Dryopteris canescens var. lobata CHRIST, Philip. J. Sci. 2 (1907) Bot. 198, quoad COPELAND 1866 (1966). Cyclosorus bordenii sensu COPEL. Fern FI. Philip. (1960) 345, p.p. Caudex short, creeping. Stipe 5-8 cm long; base of stipe to first large pinna 40 cm or more (shorter on sterile fronds). Reduced pinnae 4-8 pairs, all very small with thick aerophores Imm _ long. Lamina commonly to 35cm long; pinnae to c. 16 pairs; basal pinnae much narrowed towards their bases. Rachis bearing thick curved dark red hairs to 1 mm long on both surfaces. Largest suprabasal pinnae acuminate, to 15 x 1.7 cm (basal pinnae to 2cm wide) on larger plants, on type abruptly short-pointed, c. 8 x 1.7 cm, lobed 2/3-3/4 to costa; costules to 4mm apart; veins 7-9 pairs, basal pair anastomosing, next pair both to edge; hairs on lower surface of costae at base 0.1-0.2 mm, dis- tally to almost 1 mm long, not appressed; hairs on costules sparse; copious glands on surface be- tween veins; hairs on upper surface of costae less than | mm long, scattered similar hairs on costules and veins at least distally on pinna-lobes, surface between veins bearing a variable number of fine appressed hairs. Sori inframedial; indusia small, glabrous or with a few glands; sporangia bearing glands. Distr. Malesia: Philippines (Luzon, many col- lections), in forest at 700-1000 m. Note. On Mt Makiling a large form of this species, above described, is common. On moun- tains further north are plants (including the type) with shorter abruptly pointed pinnae but closely similar in pubescence and in reduced pinnae to the plants on Mt Makiling. COPELAND placed most of the Mt Makiling plants in Cyclosorus bordenii (here included in Sphaerostephanos heterocarpus). He failed to notice the reduced basal pinnae of the type of C. lobatus. The plants here named S. sessilipinna and S. urdanetensis are very similar to the type of S. lobatus except in size of pinnae. 66. Sphaerostephanos ellipticus (ROSENST.) HOLTTUM, Kalikasan 4 (1975) 66.— Dryopteris elliptica ROSENST. Meded. Rijksherb. Leiden n. 31 (1917) 6.—Cyclosorus ellipticus (ROSENST.) COPEL. Gen. Fil. (1947) 142; Fern Fl. Philip. (1960) 351, nomen tantum. — Thelypteris elliptica 1981] THELYPTERIDACEAE (Holttum) 475 (ROSENST.) REED, Phytologia 17 (1968) 274.— Type: ELMER 13976 partim, Mindanao, Mt Urdaneta (L, n. 914, 130-95; FI, G, K, U). KEY, LOPTHE VARIETIES 1. Basal veins free on distal part of pinnae; lower surface of rachis hairy . a. var. ellipticus 1. Basal veins not free in distal part of pinnae; lower surface of rachis almost glabrous b. var. glabrior a. var. ellipticus Caudex short, probably erect. Stipe 4-7cm long, covered with very short hairs; base of stipe to first large pinna 35cm. Reduced pinnae to 6 pairs, 34cm apart, uppermost 3-5 mm _ long, lowest 2mm or less. Lamina 45 cm long; pinnae to 18 pairs; basal pinnae slightly narrowed at base; aerophores hardly 1mm long. Lower surface of rachis covered with very short hairs and rather sparse ones to imm long; upper surface with uniform hairs 1mm. Largest pinnae 8.5 X 1.6cm, base truncate, apex short-acuminate, edges lobed 2/3 towards costa; costules 3.5 mm apart, at more than 60°; veins 8-9 pairs, basal pair near base of pinnae anastomosing with short excurrent vein to sinus, near apex of pinnae just meeting at the sinus-membrane; lower surface of costae hairy as rachis but longest hairs less than 1 mm, hairs on costules sparse, between veins very short erect hairs and glands; hairs on upper surface of costae 1mm long, similar or shorter hairs scattered on c>stules and veins, short appressed hairs copious to rather sparse between veins. Sori inframedial; indusium firm with short stiff hairs; sporangia sometimes with glands. Distr. Malesia: Philippines (Mindanao), only known from type. Note. ELMER’s n. 13976 included specimens of another species S. norrisii; COPELAND’s des- cription of 1960 applies to the latter. b. var. glabrior HOLTTUM, var. nov. A speciei typica differt: pinnis 3/5 costam ver- sus lobatis; venis semper anastomosantibus; rachidi subtus fere glabra; costis subtus pilis 0.1- 0.2mm longis praeditis. —Type: M. G. PRICE 2558, Negros Oriental, Amlan, near river, 600— 800 m (K). Distr. Malesia: Philippines (Negros Oriental), only known from the type. 67. Sphaerostephanos foxworthyi HOLTTUM, sp. nov. S. elliptico (Rosenst.) Holttum affinis, differt: pinnis redactis omnibus minutis; pinnis evolutis inferioribus basin versus valde angustatis. — Type: FOXwWworRTHY BS 2441, Luzon, Tayabas Prov. Mt Banajao (K). Caudex short, creeping. Stipe 12cm _ long, glabrescent; scales broad and thin; base of stipe to first large pinna 36cm. Reduced pinnae 5 pairs, each consisting of an aerophore without detect- able blade. Lamina 50 cm long; pinnae c. 20 pairs; lowest 2 pairs much narrowed towards their bases; aerophores thick, to 1mm long. Rachis bearing pale, erect hairs 0.2mm long on lower surface, not dense. Largest pinnae 13x 1.8cm; base subtruncate; apex acuminate, sometimes with cauda I-2cm long; edges lobed 2/3-3/4 to costa; costules 4-4.5mm apart, at 60°; veins 9 pairs, | pair anastomosing, next pair usually both to margin; lower surface of costae near base bearing erect hairs 0.1-0.2 mm long, distally hairs to 0.4 mm, antrorsely curved, not appressed, hairs on costules few, between veins many glands, not hairs; upper surface of costae bearing antrorse hairs less than I mm long, rest of surface covered with fine appressed hairs 0.2-0.3mm long. Sori medial, lower ones divergent; indusia thin with glands and a few short hairs; sporangia bearing glands. Distr. Malesia: Philippines (Luzon), only known from the type. 68. Sphaerostephanos indrapurae HOLTTUM, sp. nov. Pinnae redactae usque 10-jugatae, superiores 3mm _ longae, aerophoris elongatis praeditae; pinnae sSteriles usque 13X2.3cm, fertiles 9x 1.7 cm, c. 2/5 costam versus lobatae; rachis subtus glabra, costae costulaeque pilis appressis vestitae; pagina inter venas glandulifera; pagina superior inter venas glabra; indusia glabra, sporangia glandulifera. — Type: C. G. MATTHEW 655, Sumatra, G. Kerinci 1500 m (BO; K). Stipe 10cm long; base of stipe to first large pinna 50cm. Reduced pinnae 10 pairs, blade of largest 3mm long, with elongate aerophore. Lamina 50-60 cm long; pinnae c. 20 pairs, several lower pairs much narrowed towards their bases, only upper ones truncate to full width at base. Rachis glabrous on lower surface; hairs on upper surface slender, less than 1mm long. Largest pinnae 13X2.3cm (sterile), 9x 1.7cm (fertile), lobed c. 2/5 towards costa, lobes with blunt for- ward-pointing tips; apex abruptly — short- acuminate; costules 5mm (sterile) 4mm (fertile) apart; veins 6-8 pairs, basal pair anastomosing, one or both of next pair ending at sinus-mem- brane; lower surface of costae and costules covered with fine closely appressed hairs 0.2- 0.4mm long, surface between veins bearing glands; hairs on upper surface of costae 0.5mm long, sparse very short ones on costules, no others. Sori medial, lower ones a little divergent; indusia glabrous; sporangia bearing glands. Distr. Malesia: Central Sumatra (Mt Kerinci), 1500 m, only known from the type. (ROSENST.) 69. Sphaerostephanos batacorum 476 FLORA MALESIANA [ser. II, vol. 1° HOLTTUM, comb. nov.— Dryopteris batacorum ROSENST. Fedde Rep. 13 (1914) 217, excl. var. winkleri; v.A.v.R. Handb. Suppl. (1917) 185.— Thelypteris batacorum (ROSENST.) REED, Phy- tologia 17 (1968) 263.— Type: J. WINKLER 158, Sumatra, Batak Lands (S-PA). Dryopteris stipellata var. obtusata v.A.v.R. Bull. Jard. Bot. Btzg II, 2 (1920) 151. — Syntypes: BROOKS 277, Sumatra, Lebong Tandai; 249, Tambang Sawah (BO; BM). — Fig. 13h-k. Caudex short; erect (?). Stipe 5-10cm long; base of stipe to first large pinna 30-50cm. Reduced pinnae to 20 pairs, 1.5-2cm apart, deflexed, largest 1.5-2cm long, 8mm wide, with acroscopic auricle 8mm long, edges shallowly lobed, apex broadly obtuse. Lamina to 60cm long; pinnae to 30 pairs, basal ones sometimes narrowed a little at base on basiscopic side; aerophores thick, to 2mm long. Rachis beneath bearing short pale appressed or + spreading hairs 0.1-0.2 mm long, hairs on upper surface 1mm long. Largest pinnae 9X 1.6cm; base truncate; apex abruptly short-pointed; edges lobed 1/3-2/5 to costa; costules 3—3.5 mm apart; veins to 8 pairs, basal pair anastomosing, one or both of next pair to sinus-membrane; hairs on lower surface of costa and costules pale, closely appressed, 0.2- 0.3 mm long; surface between veins bearing glands which in some specimens are sparse (most abun- dant on reduced pinnae); hairs on upper surface of costae less than 1 mm long, sometimes no long hairs on costules, rest of upper surface glabrous. Sori medial; indusia rather large, glabrous, some- times with a few glands; sporangia sometimes with a gland. Distr. Malesia: Sumatra, several collections, at 850-1000 m. Note. The type is a small specimen; the above description is prepared partly from larger speci- mens collected on G. Singgalang and G. Kerinci which agree in shape and pubescence of frond and of reduced pinnae. 70. Sphaerostephanos angustibasis HOLTTUM, sp. nov. Caudex erectus, gracilis; pinnae redactae 6- jugatae, superiores 5mm _ longae, auriculatae; aerophora elongata; pinnae usque 9.5 X2.1 cm, inferiores basi angustatae, 2/3 costam versus lobatae; costae subtus patenti-pilosae; pagina in- ter venas subtus glandulosa; indusia sporangiaque glandulifera. — Type: ALSTON 16724, Tidore, G. Kiematuba (BM). Caudex slender, to 20 cm tall. Stipe 15 cm long, short-hairy; base of stipe to first large pinna 60cm. Reduced pinnae c. 6 pairs, subopposite, upper ones 5mm long, deflexed, narrow, with basal acroscopic auricle 4 mm long. Lamina 38 cm long; pinnae to 15 pairs, almost all opposite, lower 3-4 pairs narrowed towards their bases, basal pair 5-7 mm wide at base; aerophores 1mm _ long. Lower surface of rachis bearing curved brown hairs and very short ones, brown hairs on upper surface more uniform. Largest pinnae 9 x 2.1 cm, base truncate, apex short-acuminate, edges lobed up to 2/3 towards costa, lobes slightly falcate; costules 5mm apart, at more than 60°; veins 7-8 pairs, all at a wide angle to costule, pale and prominent both sides, basal pair anastomosing, next pair both to edge; most hairs on lower sur- face of costae minute, with some longer, erect, on costules all short, glands present throughout lower surface; upper surface of costae covered with brown hairs, similar hairs scattered on costules and veins, surface between veins glabrous. Sori medial; indusia small, with glands and a few hairs; sporangia with many glands, sometimes also with a seta; spores with many small wings. Distr. Malesia: Moluccas (Tidore), only known from the type. 71. Sphaerostephanos nudisorus HOLTTUM, sp. nov. Pinnae redactae 12-jugatae, superiores 3mm longae; aerophora 2mm longa; pinnae normales usque 15X1.8cm, profunde lobatae; rachis sub- tus glabra, costae subtus pilis minutis erectis praeditae; pagina inter venas subtus glandulifera, supra glabra; sori mediales, exindusiati; sporan- gia nec glandulis nec setis praedita. — Type: T. G. WALKER 12269, Central Celebes (BM). Caudex not known. Stipe 5cm long, glabrous; base of stipe to first large pinna 50cm. Reduced pinnae c. 12 pairs, upper ones 3mm long, with aerophores 2 mm long. Lamina 95 cm long; pinnae c. 40 pairs, lowest not narrowed at base; one pair of intermediate length between normal and reduced pinnae. Lower surface of rachis bearing glands and small scales only, upper surface covered with hairs 1mm long. Largest pinnae 15x 1.8cm; base truncate with basal acroscopic lobe 2 mm longer than next; apex acuminate with cauda 1.5-2cm long; edges lobed to 2mm from costa, lobes oblong, hardly falcate, with rounded ends; costules 4mm apart, at more than 60°; veins 8 pairs, basal pair anastomosing near base of pinna, near apex just meeting at the sinus; hairs on lower surface of costae very short, erect, glands abundant all over lower surface; hairs on upper surface of costae 1 mm long, similar hairs scattered on costules and veins, rest of upper surface glabrous. Sori medial, exindusiate; neither glands nor setae on sporangia; spores minutely spinulose. Distr. Malesia: Central Celebes, only known from the type. 72. Sphaerostephanos paripinnatus (COPEL.) HOLTTUM, comb. nov.— Dryopteris paripinnata COPEL. Univ. Cal. Publ. Bot. 18 (1942) 220.— Cyclosorus paripinnatus (COPEL.) COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78 (1951) 448, pl. 1981] THELYPTERIDACEAE (Holttum) 477 28.— Thelypteris paripinnata (COPEL.) REED, Phytologia 17 (1968) 302. — Type: BRASS 12435, New Guinea, Idenburg River (MICH; BO, L). Caudex probably erect. Stipe 10cm _ long, glabrous, basal scales narrow; base of stipe to first large pinna 60cm. Reduced pinnae to 10 pairs, each consisting of an aerophore 2 mm long and a minute blade; transition to normal pinnae abrupt. Lamina to 75cm long; apical section long and deeply lobed with gradual transition to pinnae; pinnae 20 pairs, their ends much upcurved, sub- coriaceous. Largest pinnae 21X2.5cm; base truncate to subcordate, apical 3-S cm gradually attenuate and crenate, middle part lobed more than >» way to costa, lobes somewhat narrowed, not falcate; costules to 6 mm apart, almost at right angles to costa; veins 11-12 pairs, at a very wide angle to costule, basal pair anastomosing, next pair both to sides of short sinus-membrane; lower surface of rachis, costae and costules with spreading hairs 0.4mm long, shorter erect hairs and glands between veins; hairs on upper surface of costa 1mm long, similar hairs scattered on costules and veins, no hairs nor glands between veins. Sori medial, basal ones divergent; no in- dusia; sporangia lacking hairs and glands on body, hairs on stalk with yellow gland, often 2; spores not seen. Distr. Malesia: W. New Guinea. Only known from the type. Ecol. At 1700 m, “a large clump-fern common in rain-forest gullies”’. 73. Sphaerostephanos novae-britanniae HOLT- TUM, Sp. nov. Caudex erectus; pinnae redactae 7-jugatae, superiores 7mm longae; pinnae normales opposi- tae, usque 8 X 1.2 cm, 2/3 costam versus lobatae; venae infimae raro anastomosantes; costae subtus pilis erectis praeditae; pagina inter venas subtus glandulifera, supra pilis brevibus suberectis praedita; indusia brevi-pilosa, sporangia glandulifera. — Type: STEVENS & LELEAN LAE 58644, New Britain, Subdistr. Pomio, 890m (K). Caudex erect, to 15cm tall. Stipe 4cm long, scales at base thin, 8x 1mm, closely setiferous; base of stipe to first large pinna 27cm. Reduced pinnae 3cm apart, c. 7 pairs, uppermost 7mm long, 5mm wide at slightly asymmetric base, edges lobed, apex acute. Lamina 40cm long; pinnae 22 pairs, almost all opposite, basal ones with 3 basal lobes gradually smaller, base 6mm wide, not auricled; aerophores not elongate. Hairs on lower surface of rachis 0.3-0.4 mm long, erect, on upper surface more than 0.5 mm long. Largest pinnae 8X1.2cm; base broadly cuneate, apex acuminate with cauda 10 x 2mm; edges lobed 2/3 to costa; costules 2.5 mm apart, at 60°; veins to 7 pairs, basal veins either meeting just below the short sinus-membrane or touching its sides, rarely joining to form an excurrent vein below the sinus; hairs on lower surface of costae as rachis, few on costules, surface between veins densely glandular with some short erect hairs; hairs on upper sur- face of costae pale, more than 0.5mm _ long, similar hairs scattered on costules and veins, sur- face between veins bearing suberect hairs 0.2 mm long and sometimes a few glands. Sori in- framedial, basal ones not divergent; indusia large, firm, short-hairy; sporangia sometimes bearing glands. Distr. Malesia: New Guinea (New only known from the type. Britain), 74. Sphaerostephanos convergens HOLTTUM, sp. nov. Pinnae redactae 4—S—jugatae, omnes parvae: aerophora non elongata; pinnae usque 6.5 X 1.3cm, 3/5 costam versus lobatae; venae infimae conniventes, non anastomosantes; costae subtus pilis erectis usque 1mm longis praeditae: pagina inter venas utrinque pilis brevibus erectis praedita, subtus etiam glandulifera; indusia pilosa; sporangia glandulifera. — Type: M. G. PRICE & B. F. HERNAEZ 164, Western Samar, 500m (K). Caudex short, suberect, bearing a close tuft of fronds. Stipe 4cm long; basal scales 7 x 0.8mm, thin; base of stipe to first large pinna 20cm. Reduced pinnae 4-S pairs, uppermost 2 mm long; aerophores swollen, hardly elongate. Lamina 28cm long; pinnae 10-12 pairs, lowest 1-2 pairs somewhat narrowed at their bases. Hairs on lower surface of rachis erect, pale, 0.3mm long, with some more than | mm long also; hairs on upper surface more than 1mm long. Largest pinnae 6.5 x 1.3. cm; base truncate; apex short-acuminate; edges lobed c. 3/5 to costa, lobes slightly falcate, separated by sinuses nearly | mm wide; costules 4mm apart, at 60°; veins to 6 pairs, basal pair both touching sides of short sinus-membrane or meet- ing just below it without fusing, next pair to edge; hairs on lower surface of costae and costules erect, as on rachis but shorter, surface between veins bearing many short erect hairs and glands; hairs on upper surface of costae 1mm long, similar hairs scattered on costules and veins, sur- face between veins bearing suberect hairs 0.2 mm long. Sori inframedial, basal ones not divergent; indusia bearing hairs 0.4 mm long; sporangia bear- ing glands. Distr. Malesia: Philippines (W. Samar), only known from the type. Ecol. In forest on limestone soils. 75. Sphaerostephanos unitus (L.) HOLTTUM, J. S. Afr. Bot. 40 (1974) 165; Kalikasan 4 (1975) 63. — Polypodium unitum L. Syst. Nat. ed. 10. 2 (1759) 1326, excl. syn.— Aspidium unitum (L.) Sw. in Schrad. J. Bot. 1800, 2 (1801) 32, nomen tantum; WILLD. Sp. Pl. ed. 4, 5 (1810) 241. — Dryopteris unita (L.) O. KTZE, Rev. Gen. Pl. 2 (1891) 811; BACKER & PosTH. Varenfl. Java (1939) 51.— 478 FLORA MALESIANA [ser. II, vol. 1° Cyclosorus unitus (L). CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 192; HOLTTUM, Rev. FI. Mal. 2 (1955) 260, f. 147; COPEL. Fern FI. Philip. (1960) 360.— Thelypteris unita (L.) MORTON, Amer. Fern J. 49 (1959) 113.—Type: without locality (LINN). Tectaria serrata CAV. Descr. (1802) 251; C. CHR. Dansk Bot. Ark. 9, n. 3 (1937) 16, t. 1, f. 10-11.— Nephrodium serratum (CAV.) PRESL, Rel. Haenk. (1825) 34.— Type: NEE, Marianas (MA). Nephrodium insculptum DeEsv. Mém. Soc. Linn. Paris 6 (1827) 254.—Type: Réunion, no collector named (P). Aspidium cucullatum BL. En. Pl. Jav. (1828) 151.— Nephrodium cucullatum (BL.) BAK. Syn. Fil. (1867) 290; BEDD. Handb. (1883) 270; RACIB. Fl. Btzg I (1898) 184. — Dryopteris cucullata (BL.) CHRIST, Philip. J. Sci. 2 (1907) Bot. 194; v.A.v.R. Handb. (1908) 213.— Type: BLUME, Java (L, n. 908, 337-5). Nephrodium haenkeanum PRESL, Epim. Bot. (1851) 46; HoLtTruM, Novit. Bot. Univ. Carol. Prag. 1968 (1969) 17.— Thelypteris haenkeana (PRESL) REED, Phytologia 17 (1968) 281. — Type: HAENKE, Marianas (PRC). KEY TO THE VARIETIES 1. Glands present on some part of lower surface of pinnae. 2. Rather large glossy brownish glands present on and between veins on lower surface a. var. unitus 2. Smaller dull yellow glands present on costules and veins only .b. var. mucronatus 1. Glands lacking on lower surface; cells between veins all papilliform . ¢. var. papilliferus a. var. unitus Rhizome long-creeping, commonly to 5mm diameter; stipe 10-20 cm long, base of stipe to first normal pinna 30-60cm; reduced pinnae 6-12 pairs, lower ones very small, upper ones triangular with symmetric base, to 10mm long. Lamina 30- 60cm long; pinnae closely placed, lowest not or little narrowed at base, aerophores not elongate. Hairs on both sides of rachis less than | mm long. Largest pinnae 10-15 x 0.8-1.5cm; base broadly cuneate; apex rather evenly attenuate; edges lobed 1/3 towards costa; lobes rounded with a slight point at falcate end of costule; costules 3-4mm apart; veins 8-10 pairs, 13 pairs anas- tomosing, next 2-4 pairs passing to sides of sinus- membrane which forms a ridge on the lower sur- face; hairs on lower surface of costae c. 0.5 mm long, slender, always antrorse but not closely ap- pressed, similar hairs on costules; rather large brownish glossy glands present on costules and veins, and between veins; upper surface of pinnae hairy on costae only. Sori supramedial; indusia firm, glabrous or with a few short hairs, some- times also glands; sporangia bearing glands near annulus, also copious glands at the ends of hairs on sporangium-stalks. Distr. E. Africa; Mascarene Islands; Ceylon & S. India; N.E. India, Burma & Thailand (few specimens); Malesia (Malaya, Sumatra, Java, S.W. Celebes, Lesser Sunda Is., few specimens from Luzon and Mindanao, | from W. New Guinea); Guam, N. Queensland. Ecol. At low altitudes, in open places, in moist (not swampy) ground. Notes. The identity of this species is fixed by the type. LINNAEUS’s description is not adequate, and his references to plants from Asia all indicate the species here named Cyclosorus interruptus (WILLD.) CHING. ROBERT BROWN and others interpreted Polypodium unitum according to the references, and BEDDOME followed this usage. It is now customary to accept the type as fixing the application of a name in such cases; it is to be noted that the specimen in the Willdenow Her- barium agrees with the Linnaean type. The next name, Tectaria serrata CAV., was based on a specimen collected by NEE in Guam. Specimens of the same species were also collected there by HAENKE, and PRESL at first (1825) identified them with T. serrata, but later (1851) thought this was wrong and published the new name Nephrodium haenkeanum for them. There is no doubt that the two collections were identical. b. var. mucronatus (CHRIST) HOLTTUM, comb. nov.— Dryopteris cucullata var. mucronata CHRIST, Philip. J. Sci. 2 (1907) Bot. 195; v.A.v.R. Handb. (1908) 819. — Type: CUMING 182, Luzon (CHRIST’s specimen not seen; isotypes FI-W, K, IE): Aspidium multilineatum METT. Farngatt. IV (1858) 108. — Syntypes: CUMING 182, 278, Luzon (duplicates of 278 at FI-W, G, K, LE, PRC). Nephrodium haenkeanum sensu BAK. Syn. Fil. (1867) 290. Pinnae 15-20 1.0-2.0cm; veins 10-14 pairs; glands present only on costules and veins of lower surface, small, dull, yellow; indusia always hairy. Distr. Polynesia (Samoa, Fiji, Tonga), Micro- nesia (Palau), and Malesia: New Guinea, Moluc- cas, Lesser Sunda Is. (Sumba, Wetar, Timor), Celebes, Philippines and Borneo. Notes. The characteristic glands on the lower surface of costules and veins were described by METTENIUS but ignored by all others. MET- TENIUS however made a confusion by adopting the name multilineatum from WALLICH and citing WALLICH 353 from Penang, which represents the species here named S. penniger, very different from the CUMING specimens which were clearly the basis of METTENIUS’s description. BEDDOME later adopted the WAL- LICH epithet for S. penniger. 1981] BAKER wrongly referred CUMING 182 to Nephrodium haenkeanum. FOURNIER also adop- ted this name in his work on the ferns of New Caledonia, but his citation of specimens shows that he confused it with other species. The New Caledonia specimens which he so named are S. invisus. The varietal name mucronatus was first pub- lished by J. SMITH in 1843, as a nomen nudum under which he cited specimens of three species; it was not validated until CHRIST’s description of 1907. Both var. unitus and var. mucronatus occur in the Lesser Sunda Islands; they may occupy different habitats; records are inadequate. c. var. papilliferus HOLTTUM, var. nov. A typo speciei differt: costulis venisque subtus eglandulosis; pagina subtus inter venas omnino minute papillifera.—Type: HOOGLAND 9084, N.E. New Guinea, Huon Peninsula, 1320m (K). Distr. Malesia: New Guinea, many specimens, at 1300-2000 m. Note. All cells on the lower surface of pinnae have small papilliform colourless outgrowths. Stomata presumably occur, but they are not detectable. The condition is probably an adap- tation to extreme exposure. 76. Sphaerostephanos sessilipinna (COPEL.) HOLTTUM, Kalikasan 4 (1975) 53.— Dryopteris sessilipinna COPEL. Philip. J. Sci. 6 (1911) Bot. 145.— Cyclosorus sessilipinna (COPEL.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 249; COPEL. Fern FI. Philip. (1960) 353. — Thelypteris sessilipinna REED, Phytologia 17 (1968) 313. — Type: MERRILL 6934, Negros, Mt Canlaon (MICH). Dryopteris canlaonensis COPEL. Philip. J. Sci. 40 (1929) 300.— Thelypteris canlaonensis (COPEL.) REED, Phytologia 17 (1968) 266. — Type: MERRILL 6934, Negros (MICH). Dryopteris austrophilippina COPEL. Philip. J. Sci. 40 (1929) 300. — Cyclosorus austrophilippinus (COPEL.) COPEL. Gen. Fil. (1947) 142; Fern FI. Philip. (1960) 354. — Thelypteris austrophilippina (COPEL.) REED, Phytologia 17 (1968) 262.— Type: COPELAND 1705, Mindanao, San Ramon (MICH). Caudex short, creeping. Stipe 5cm long; base of stipe to first large pinna 15-20cm. Reduced pinnae 4-6 pairs, all less than 2 mm long. Lamina to 20cm long; apical section 10-12cm long, deeply lobed, tapering very gradually to tip, at base with gradual transition to pinnae; free pinnae 10-12 pairs, very close; basal pinnae deflexed and narrowed towards their bases; aerophores not elongate. Rachis with thick curved brown hairs both sides. Middle pinnae 2.3x0.8cm, lobed about 1/3 to costa, apex obtuse; costules 2mm apart; veins 2-4 pairs, basal pair anastomosing in basal half of pinna, free near its tip; hairs on lower THELYPTERIDACEAE (Holttum) 479 surface of costae pale, antrorsely curved; glands present on surface between veins; upper surface of type bearing rather sparse appressed hairs be- tween veins, of other specimens many. Sori medial or inframedial; indusia glabrous or with a few short hairs; sporangia bearing glands. Distr. Malesia: Philippines (Negros, danao). Notes. The type of D. austrophilippina has a less tapering apical lamina than the other type, and paler hairs on lower surface of rachis. In 1975 I included D. urdanetensis COPEL. also as a synonym of the present species, but it lacks very small basal reduced pinnae. Apart from size, there is little to distinguish this from S. lobatus. Min- 77. Sphaerostephanos lobangensis (C. CHR.) HOLTTUM, comb. nov. — Dryopteris lobangensis C. CHR. Gard. Bull. Str. Settl. 7 (1934) 245.— Thelypteris lobangensis (C. CHR.) REED, Phy- tologia 17 (1968) 289.— Type: CLEMENS 10728, Mt Kinabalu, Pakka Cave to Lobang (MICH; BO). Caudex short, suberect. Stipe 12cm long; base of stipe to first large pinna 50cm. Reduced pinnae 10 pairs, upper ones 3 mm long. Lamina to 40 cm long; pinnae to 20 pairs; basal pinnae a little reduced and deflexed, narrowed towards their bases; aerophores not elongate. Lower surface of rachis glabrous. Largest pinnae 6X 1.4cm; base truncate; apex short-acuminate; edges lobed 1/3 to costa; costules 3 mm apart, at 60°; veins to 6 pairs, 1 pair anastomosing, next acroscopic vein to side of sinus-membrane; lower surface of pinnae quite glabrous, with yellow glands throughout; hairs on upper surface of costae short, on costules shorter, few on veins, no others. Sori inframedial, basal ones not divergent; indusia small, glandular; sporangia bearing glands. Distr. Malesia: Sabah (Mt Kinabalu), in forest at 1500-2000 m; two collections, the second being an unnumbered CLEMENS specimen from Marei Parei spur “on rocks in stream” (BO). Notes. The above description is prepared from the Bogor isotype; the type at MICH is somewhat smaller. CHRISTENSEN described the rhizome as creeping with widely-spaced stipes, but this is not true of the Bogor specimen. 78. Sphaerostephanos gymnorachis HOLTTUM, sp. nov. Caudex erectus; pinnae redactae usque 8-10- jugatae, superiores 3 mm longae; pinnae normales fertiles usque 16X1.9cm, 1/2-5/8 costam versus lobatae; rachis subtus glabra, costae costulaeque pilis brevibus antrorsis vestitae, pagina inter venas glandulosa; indusia glabra; sporangia glanduli- fera.—Type: HoLttTumM 34, Mt Kinabalu, Pinosok area, Nov. 1972 (K). Caudex erect, at least to 10cm tall. Stipe 10cm long, dull reddish; base of stipe to first large pinna 480 FLORA MALESIANA [ser. II, vol. 1° of largest fronds 50cm. Reduced pinnae 4-5 cm apart, 8-10 pairs, uppermost 3mm long, lowest with no evident blade. Lamina to 70cm long; pinnae more than 20 pairs, in most cases all opposite; basal pinnae gradually narrowed towards base in basal 4cm, next 2-3 pairs less narrowed; aerophores not elongate. Lower sur- face of rachis glabrous; hairs on upper surface 0.5mm long, brown. Largest fertile pinnae 16x 1.9cm (sterile to 2.3cm wide); base broadly cuneate; apex acuminate with cauda 1-1.5cm; edges lobed a little more than > way to costa (about 3 way on upper pinnae or those of small fronds); lobes falcate, distinctly tapered to a blunt tip; costules 3.5-4mm apart (to 5mm in sterile fronds); veins to 12 pairs, basal pair anastomosing with long excurrent vein to sinus, second pair both to sides of sinus-membrane; lower surface of cos- tae near base covered with hairs 0.1-0.2 mm long, near apex 0.3-0.4mm, antrorse but not closely appressed, on costules similar, sparser and shorter on veins, many glands on surface between veins (hairs on lower surface of sterile pinnae somewhat longer, distal ones on costae 0.5mm); hairs on upper surface of costae brown, 0.7 mm long, on costules 0.2 mm, pale, rarely with a solitary longer hair; surface between veins glabrous or with a few pale appressed hairs. Sori medial, basal ones somewhat divergent; indusia large, glabrous; sporangia bearing glands. Distr. Malesia: Sabah (Mt Kinabalu), in forest at 1500 m. Notes. In addition to the type collection, CLEMENS 28233 and 28382 (from Tenompok, 1500 m) have pinnae not opposite and upper sur- face between veins covered with many appressed hairs; CLEMENS 29050 (Silau Basin,.2100 m) is a smaller plant (pinnae to 9x 1.5cm) with almost sterile fronds, differing in scattered pale hairs 1mm long on lower surface of rachis. S. gym- norachis may be a fully developed form of S. lobangensis. 79. Sphaerostephanos pullenii HOLTTUM, sp. nov. Pinnae redactae 6-jugatae vel ultra, omnes parvae; pinnae normales 263.0cm, 3/4-4/5 costam versus lobatae; rachis costaeque subtus glabra vel pilis minutis paucis praedita; pagina subtus inter venas glandulosa, supra pilis erectis brevibus praedita; indusia parva, glandulifera, sporangia nec glandulis nec setis instructa. — Type: R. PULLEN 672, N.E. New Guinea, East- ern Highlands (BM; L, LAE). Caudex erect. Stipe 30cm long, glandular; base of stipe to first long pinna 85cm. Reduced pinnae 6 cm apart, at least 6 pairs, all very small. Lamina 85cm or more long; pinnae to 30 pairs; basal pinnae narrowed towards their bases, basal lobes 6-10mm long; aerophores not elongate. Lower surface of rachis glabrous or with sparse minute erect hairs; upper surface with pale hairs less than Imm long. Largest pinnae 26 x 2.0 cm; base trun- cate; apex caudate-acuminate (cauda to 3cm); edges lobed to 3-3.5 mm from costa, lobes slightly falcate; costules 5-7 mm apart, at more than 60°; veins to 12 pairs, basal pair at a wide angle to costule meeting to form a long excurrent vein, one or both of next pair passing to sinus-membrane; lower surface of costae near base with sparse minute hairs, somewhat longer hairs distally, cos- tules similar; copious glands between veins; hairs on upper surface of costae 0.5mm long, shorter on costules with a few long ones near apices of lobes, surface between veins bearing very short suberect hairs. Sori medial, lower ones divergent, lowest somewhat elongate; indusia small, thin, with marginal glands; sporangia bearing neither glands nor setae. Distr. Malesia: Papua New Guinea. Ecol. In Nothofagus forest at 2000 m. 80. Sphaerostephanos gregarius (COPEL.) HOLT- TUM, comb. nov. — Cyclosorus gregarius COPEL. J. Arn. Arb. 24 (1943) 440; Philip. J. Sci. 78 (1951) 451, pl. 31. — Thelypteris gregaria (COPEL.) REED, Phytologia 17 (1968) 280.— Type: BRASS 6759, Papua, Fly River (MICH; BM, BO, L). Caudex short; erect or suberect. Stipe 25-30 cm long, dull reddish; base of stipe to first large pinna 60cm. Reduced pinnae 7 pairs, lowest 3-4mm long, uppermost 2.2 cm long, broadly triangular, subentire, base slightly asymmetric. Lamina 60 cm long; apex on type pinna-like, on isotype at L not pinna-like; pinnae 16 pairs, lower ones 4cm apart, subopposite, not narrowed towards their bases, upper ones close and alternate; aerophores not elongate. Rachis glabrous on lower surface or with minute appressed hairs; hairs on upper sur- face not over 1mm long. Largest pinnae 15 x 1.5cm; base subequally broadly cuneate and a little dilated both sides; apex acuminate; edges lobed to a depth of 2-2.5mm (on BM and BO isotypes little more than | mm); costules 3.5-4 mm apart, at 60° or more; veins 6-7 pairs, 2-3 pairs anastomosing, | pair to short sinus-membrane; lower surface of costa glabrous or with minute appressed hairs, no other hairs on lower surface, sparse glands present on costules and on surface between veins; hairs on upper surface of costae 1mm long, rest of upper surface glabrous apart from short hairs on margin. Sori medial or in- framedial, lowest not divergent; indusia very small, thin, bearing a few very short hairs; sporangia bearing glands. Distr. Malesia: Eastern New Guinea, at al- titudes to 1000 m, in forest. 81. Sphaerostephanos_ tephrophyllus (COPEL.) HoOLtTTuM, Kalikasan 4 (1975) 58.— Dryopteris tephrophylla COPEL. Philip. J. Sci. 40 (1929) 296. — Cyclosorus tephrophyllus (COPEL.) CopEL. Gen. Fil. (1947) 143; Fern FI. Philip. 1981] (1960) 350.— Thelypteris tephrophylla (COPEL.) REED, Phytologia 17 (1968) 319.—Type: COPELAND s.n. 18 April 1905, Mindanao, Zam- boanga, San Ramon (MICH). Caudex suberect, short. Stipe 3-4 cm long; base of stipe to first large pinna 12-15cm. Reduced pinnae 3-5 pairs, lowest 3-5 mm long, uppermost to 1.5x0.8cm, entire, deflexed, auricled on acroscopic base; transition to normal pinnae not abrupt. Lamina to 25cm long; apical sec- tion to 12cm, deeply lobed and grading to upper pinnae; free pinnae 5-6 pairs, lower ones slightly reduced and sometimes a little auricled on acroscopic base; aerophores not elongate. Hairs on lower surface of rachis short, with sparse longer ones, on upper surface similar but thicker, the long ones more than 1mm. Largest pinnae 4.5 x 1.4cm (sterile), 3.5 x 1.2 cm (fertile): base truncate; apex short-pointed; edges crenate to a depth of less than 1 mm; costules 3-3.5mm apart; veins to 4 pairs, 1 pair anastomosing, next pair to sides of sinus-membrane; hairs on lower surface of costae and costules short, with a few longer ones, between veins short hairs and glands: short hairs on upper surface of costae, between veins very short suberect hairs. Sori medial; in- dusia large, with a few hairs; sporangia sometimes with a gland. Distr. Malesia: Philippines (Mindanao), 3 col- lections. Note. Allied to S. spenceri, differing in little but much smaller size. 82. Sphaerostephanos pilososquamatus (v.A.v.R.) HOLTTUM, comb. nov. — Dryopteris pilososqua- mata v.A.v.R. Bull. Dép. Agr. Ind. Néerl. 21 (1908) 4; Handb. (1908) 222: Handb. Suppl. (1917) 181, excl. var. obtusata.— Thelypteris piloso- squamata (v.A.v.R.) REED, Phytologia 17 (1968) 304.— Type: cult. Hort. Bog., origin W. New Guinea (BO; BM, L). Dryopteris paraphysata COPEL. Philip. J. Sci. 6 (1911) Bot. 74. — Thelypteris paraphysata REED, Phytologia 17 (1968) 301.— Type: C. KING 306, Papua (MICH; NSW, P). Dryopteris megaphylloides ROSENST. Fedde Repert. 12 (1913) 174.— Thelypteris megaphyl- loides (ROSENST.) REED, Phytologia 17 (1968) 292. Type: KEYSSER 120, N.E. New Guinea, Sattelberg (orig.?; UC). — Fig. 12i-k. Caudex short, creeping. Stipe on a well-grown plant 30-40cm long; base of stipe to first large pinna 45-SScm (sterile) 60-70cm (fertile). Reduced pinnae 2-3(-4) pairs, largest 5 mm long; an intermediate pair sometimes present. Lamina dimorphous; pinnae to 15 pairs, sometimes with stalks 1-2mm long; basal pinnae narrowed towards their bases; apical lamina almost pinna- like; aerophores not elongate. Hairs on rachis of sterile fronds to 1mm long both sides, rather sparse on lower surface. Largest sterile pinnae THELYPTERIDACEAE (Holttum) 481 20-27 x 3.5-4cm; base broadly cuneate; apex acuminate with cauda to 2 cm; edges lobed 1/4~-2/5 to costa, lobes falcate, narrowed to tips in more deeply lobed pinnae; costules 5—5.5 mm apart, at 60° or rather less; veins 12-15 pairs, 2+33 pairs anastomosing, 2-3 pairs to sides of sinus-mem- brane; hairs on lower surface of costae spreading, 0.5—1 mm long, not dense, on costules and veins similar but fewer, glands usually present on cos- tules and veins and between veins, not abundant: scattered hairs on upper surface of costules and veins like those on costae, appressed hairs be- tween veins sometimes present. Fertile pinnae commonly to 18x2cm, less deeply lobed than sterile and with closer costules, hairs on all parts shorter and more sparse; veins 7-9 pairs: sori medial, lower ones not or little divergent; indusia bearing glands and hairs; sporangia usually with a seta, sometimes also a gland, near annulus, and a gland at end of hair on stalk. Distr. Malesia: New Guinea, Archipelago, in forest, 0-1400 m. Note. One specimen assigned to this species appears to be quite eglandular. Another (NAK- AIKE 73) is small, lacks reduced pinnae and has no glands on sporangia. Bismarck 83. Sphaerostephanos angustifolius (PRESL) HOLTTwuM, Kalikasan 4 (1975) 66. — Nephrodium angustifolium PRESL, Epim. Bot. (1851) 48: Hook. Spec. Fil. 4 (1862) 69.— Cyclosorus angustifolius (PRESL) COPEL. Fern Fl. Philip. (1960) 349. — Thelypteris setulosa REED, Phytolo- gia 17 (1968) 313, nom. nov. — Type: CUMING 268, Luzon (PRC; B, BM, E, FI-W, G, K, LE, US). Dryopteris hispidula sensu C. CHR. Ind. Fil. (1905) 271, p.p.; v.A.v.R. Handb. (1908) 228. Caudex short, erect. Stipe 46cm long, dark reddish, densely short-hairy; base of stipe to first large pinna 15cm or more. Reduced pinnae to 5 pairs, highest 3mm long. Lamina to 35cm long: pinnae c. 17 pairs; lower 2-3 pairs somewhat narrowed towards their bases with basal acros- copic lobe a little elongate: aerophores not elon- gate. Lower surface of rachis covered with short erect hairs and scattered longer ones (some of them dark red), upper surface covered with pale hairs 0.5mm long. Largest pinnae 9.5 x 0.8cm; base truncate and not dilated: apex a subentire cauda 1.5-2.0 cm long; edges lobed 3 way to costa, lobes oblique; costules 3mm apart, at 60°; veins 4-5 pairs, 1 pair anastomosing, next pair to mar- gin; hairs on lower surface of costae not dense, very short, with scattered hairs to 0.8 mm long, on costules similar but more sparse, minute erect hairs present between veins, glands throughout lower surfaces; hairs on upper surface of costae 0.5mm long, similar hairs scattered on costules and veins, suberect hairs 0.1-0.2 mm long between veins. Sori medial; indusia short-hairy; sporangia bearing glands. 482 FLORA MALESIANA (ser. II, vol. 1° Distr. Malesia: Philippines (Luzon); a second collection is M. G. PRICE 2553, Zambales Prov., from exposed place by stream (small plants). Note. S. angustifolius is very near S. hetero- carpus; it is peculiar in its very narrow pinnae. 84. Sphaerostephanos efogensis HOLTTUM, sp. nov. S. conferto (Brause) affinis, differt: membranis sinuum brevibus; costis subtus pilis usque 1mm longis vestitis; costulis subtus glanduliferis; soris supramedialibus, sporangiis glanduliferis. — Type: J. R. Crorr LAE 61873, Papua, Central Distr., Port Moresby Subdistr. Efogi (K). Caudex short, decumbent (collector). Stipe 12- 20cm long, basal scales short and thick, ap- parently 1.5 1mm; base of stipe to first large pinna 40-60 cm. Reduced pinnae 12 pairs, lowest 2-4 mm long, middle ones 10 x 10mm, uppermost 1.7cm long, 1.3cm wide at truncate base, trian- gular, crenate. Lamina 43cm long; apex broadly deltoid and deeply lobed; pinnae 20 pairs, basal ones not narrowed at base; aerophores not elon- gate. Hairs on lower surface of rachis not dense, mostly 0.1-0.3mm long with some more than Imm, on upper surface long hairs more numerous, to 1.5mm long. Largest pinnae 8.5 x 1.7 cm: base subtruncate, almost symmetric; apex short-acuminate; edges lobed to a depth of 2mm, lobes oblique, subtriangular; costules to 3.5mm apart; veins 7-8 pairs, all except basal ones very oblique, 2-25 pairs anastomosing to form a zig-zag excurrent vein, | pair to short sinus-membrane; hairs on lower surface of costae as on rachis, on costules shorter, glands present on costules and veins, between veins sparse short erect hairs and sometimes a few glands; hairs on upper surface of costae mostly 0.3mm long with scattered long ones which occur also on costules and veins, surface between veins bearing appressed hairs 0.2-0.3 mm long. Sori a little supramedial; indusia thin, with a few hairs 0.5mm long; sporangia bearing glands. Distr. Malesia: Papua New Guinea. Ecol. At 1500 m in riverine hill forest. 85. Sphaerostephanos dichrotrichoides (v.A.v.R.) HOLTTUM, Kalikasan 4 (1975) 65.— Dryopteris dichrotricha COPEL. Philip. J. Sci. 7 (1912) Bot. 54, non COPEL. 1911.—Dryopteris di- chrotrichoides v.A.v.R. Handb. Suppl. (1917) Corr. 48, nom. nov. — D. weberi COPEL. Philip. J. Sci. 38 (1929) 135, nom. nov. superfl. — Lastrea dichrotrichoides (v.A.v.R.) COPEL. Fern FI. Philip. (1960) 329.— Thelypteris dichrotrichoides (v.A.v.R.) REED, Phytologia 17 (1968) 272.— Type: WEBER 1173, Mindanao, Mt Hilong-Hilong (MICH; E, G, K, NSW, P, US). Caudex short, creeping. Stipe 15cm long, pale, short-hairy; base of stipe to first large pinna 30cm. Reduced pinnae to 5 pairs, upper ones 4mm long, auricled, lowest very small. Lamina 35cm long; pinnae 18 pairs, basal pinnae a little narrowed towards their bases, strongly auricled on acroscopic base; aerophores swollen. Hairs on both surfaces of rachis 1 mm long, pale. Largest pinnae 7-12cm long, 1.5cm wide at broadly cuneate base from which they are gradually attenuate to apex; edges lobed to 2mm from costa, lobes oblique, slightly falcate; costules to 3mm apart; veins 8-9 pairs, basal pair anas- tomosing near base of pinna, passing to sides of a short sinus-membrane towards apex of pinna; coarse rather sparse hairs 1mm long on lower surface of costae and costules, shorter hairs on veins and surface between them, glands present on costules and veins, few between veins; hairs on upper surface of costae 1mm long, similar hairs scattered on costules and veins, whole upper sur- face covered with fine appressed hairs nearly 0.5mm long. Sori medial, basal ones not diver- gent; indusia with copious hairs 0.5mm _ long; sporangia often with 2-3 glands. Distr. Malesia: Philippines Panay”). (Mindanao: 86. Sphaerostephanos nakaikei HOLTTUM, sp. nov. Pinnae redactae S-jugatae, superiores 10mm longae, valde auriculatae; pinnae normales usque 14 x 2.3 cm, 2/3-3/4 costam versus lobatae; venae infimae anastomosantes vel ad basin sinus con- niventes; costae subtus pilis erectis brevibus ves- titae; pagina inter venas subtus glandulifera, supra pilis appressis vestita; indusia glandulifera, sporangia non glandulifera. — Type: T. NAKAIKE 358, E. New Guinea, Mt Wilhelm 2500-3500 m (K; TNS). Caudex ‘“‘massive or ascending” (NAKAIKE). Stipe 20cm long, short-hairy, basal scales broad and thin; base of stipe to first large pinna 45 cm. Reduced pinnae 5 pairs, lowest 2mm long, up- permost 10 mm with basal acroscopic auricle 5mm long; 1-2 pairs pinnae of intermediate length sometimes present. Lamina 60-70 cm long; pinnae 25 pairs; basal large pinnae not narrowed at base, slightly auricled; aerophores slightly elongate. Hairs on lower surface of rachis dense, brownish, erect, 0.3-0.5 mm long, on upper surface a little longer. Largest pinnae 13-14 cm long, 1.8-2.3cm wide; base truncate; apex acuminate; edges lobed 2/3-3/4 to costa, lobes falcate, separated by dis- tinct sinuses; costules 5mm apart; veins 8-10 pairs, basal pair anastomosing with a short excur- rent vein to sinus, less commonly both touching sides of short sinus-membrane; lower surface of costae densely covered with erect hairs 0.2mm long, hairs on costules sparse, glands present on costules and veins and on surface between veins; hairs on upper surface of costae 0.7mm long, similar hairs scattered on costules and veins, sur- face between veins rather sparsely covered with 1981] THELYPTERIDACEAE (Holttum) 483 appressed hairs. Sori medial; indusia bearing glands; sporangia with neither setae nor glands. Distr. Malesia: Papua New Guinea (Mt Wil- helm and Mt Giluwe: PARRIS & CROXALL 5900, at 2950 m). 87. Sphaerostephanos woitapensis HOLTTUM, sp. nov. S. nakaikeo affinis, differt: pinnis minoribus, brevi-stipitatis, inferioribus sensim decrescenti- bus; lobis pinnarum distaliter dentatis; venis infimis semper anastomosantibus. — Type: NAKAIKE 576, Papua, Central Distr., Woitape (TNS; K). Caudex thick, short-creeping or ascending. Stipe 25cm long, copiously short-hairy. Lamina to 75cm long: pinnae to more than 20 pairs, distinctly stalked: 5-6 lower pairs gradually decrescent and more widely spaced, lowest 8 mm long; aerophores slightly swollen. Hairs on lower surface of rachis suberect, 0.2-0.4mm long, on upper surface 0.7mm long. Largest pinnae 9.5 x 2.0cm; base truncate and in some cases slightly dilated both sides; apex short-acuminate: edges lobed c. 3/5 to costa, lobes oblique. slightly fal- cate, with distinct teeth at ends of some distal veins; costules 44.5mm apart at 60°; veins 6~7 pairs, pale and prominent both sides, basal pair always anastomosing, second pair both to margin: hairs on lower surface of costae near base 0.1 mm, distally 0.3 mm long, on costules and veins sparse and very short, glands present on surface between veins; upper surface of costae hairy as rachis, scattered similar hairs on costules and veins, ap- pressed hairs 0.2mm long between veins. Sori medial; indusia with short hairs and many glands: sporangia lacking glands and setae. Distr. Malesia: Eastern New Guinea at 2500— 2800 m. Note. Other collections have smaller pinnae than the type, on smallest fronds 5.0 1.2cm, rather rigid; these do not show teeth at ends of distal veins. 88. Sphaerostephanos ekutiensis HOLTTUM, sp. nov. Pinnae redactae 8-jugatae. usque 10mm lon- gae; pinnae normales usque 7.0*1.8cm, c. 3/5 costam versus lobatae: venae 7-jugatae, infimae solum anastomosantes: costae subtus pilis paucis praedita, pagina inferior pinnarum cetera glabra, glandulifera, glandulis inter venas paucis: sori in- framediales; indusia tenuia, glandulis multis omata: sporangia nec _ setis nec glandulis praedita.— Type: B. S. PARRIS & J. P. CROXALL 6022, N.E. New Guinea, Morobe Distr., Ekuti Range (CGE; K). Caudex not known; stipe 7 cm long, glabrous on abaxial surface, scales at base 7 x 1.5 mm; base of stipe to first normal pinna 30 cm; reduced pinnae 8 pairs, uppermost 104mm with an auricle 8mm long, lowest 5mm long, deflexed, narrowly trian- gular with acroscopic auricle 3 mm long; two pairs of pinnae of intermediate length present between reduced and normal pinnae. Lamina excluding reduced pinnae 37cm long; pinnae 18 pairs, all opposite, texture firm; basal pinnae not narrowed at their bases. Both surfaces of rachis bearing thick curved pale brown hairs | mm long. Largest pinnae 7.0 1.8cm, sessile; aerophores slightly swollen; base subtruncate, not auricled; apex ab- ruptly short-caudate; edges lobed 3/5 towards costa, lobes slightly falcate with rounded tips; costules 4mm apart. at more than 60° to costa: veins to 7 pairs, basal pair anastomosing, next pair to margin; lower surface of costae bearing rather sparse thick pale hairs 0.3mm long and glands, rather large glands and no hairs on costules and veins, rather sparse glands between veins: upper surface of costae covered with pale hairs less than 1 mm long, shorter hairs scattered on costules and veins, no other hairs. Sori inframedial; indusia thin, with many glands: sporangia bearing neither setae nor glands. Distr. Malesia: known from the type. Ecol. In Nothofagus forest at 2250 m. Papua New Guinea: only 89. Sphaerostephanos omatianus HOLTTUM, sp. nov. Stipes 2.5 cm longus; lamina 7-8 cm longa, pars apicalis profunde lobata 5.5cm longa, pinnae liberae 3-4-jugatae. crenatae. infimae leviter redactae; venae liberae: costae costulaeque subtus pilis longis glandulisque praeditae: sori exindusi- ati, sporangia nuda.— Type: WOMERSLEY & SIMMONDS 5076A, Papua. Gulf Div., Omati (BRI). Caudex apparently short-creeping. Stipe 2.5cm long, covered with pale spreading hairs 1 mm long. Lamina 7-8 cm long. apical 5.5 cm deeply lobed, grading to pinnae; free pinnae 34 pairs, lowest only reduced, 6-7 mm long, deflexed and a little narrowed on basiscopic side. Largest pinnae 1.1 x 0.45cm, almost sessile; base broadly cuneate: apex obtusely pointed: edges crenate near base, entire distally; costules near base of pinnae hardly 2mm apart. twice forked, distal ones forked or simple, all veins free: lower surface of rachis, costae and costules bearing hairs 1 mm long and some shorter ones, slender erect hairs on surface between veins, glands present on costae and cos- tules: slender hairs 1 mm long on upper surface of costae and costules. appressed hairs on surface generally. Sori in an uneven row on each side of costa, on acroscopic branches of forked costules; no indusia: neither glands nor setae on sporangia: spores minutely papillose. Distr. Malesia: Papua, known only from type collection. Ecol. At an altitude of 30m, in shallow soil over pinnacle limestone, in dense rain-forest. 484 FLORA MALESIANA [ser. II, vol. 1° 90. Sphaerostephanos alticola HOLTTUM, sp. nov. Caudex brevis, crassus; stipes 12cm longus; lamina 45-55 cm longa; pinnae 40-jugatae, 12-14- jugatae inferiores sensim decrescentes, maximae 3.3 cm longae, 7-8 mm latae, leviter lobatae, rigidae, lobis cucullatis; costae costulaeque subtus patenti-pilosae paulo glandulosae; indusia parva, glandulis pilisque praedita; sporangia setifera. — Type: HOOGLAND 9762, N.E. New Guinea, Huon Peninsula, Salawaket Range (LAE; L). Caudex “thick, + horizontal”. Stipe 12 cm long, base covered with glossy scales 10x 1.3-3 mm. Lamina 45-55 cm long; pinnae to 40 pairs, rigidly coriaceous, lower 12-14 pairs gradually smaller and more widely spaced, lowest 5mm long, 6mm wide at truncate base, 3-lobed, upper ones trian- gular, crenate; aerophores not elongate. Rachis covered with dense pale spreading hairs 0.7 mm long. Largest pinnae 3.3cm long, 7-8mm wide above base, base truncate and a little dilated; apex shortly obtuse; edges lobed to a depth of | mm or a little more deeply; lobes rounded, their edges much deflexed so that the lower surface is con- cave; costules little more than 2mm apart; veins 3-5 pairs, pale, prominent both sides, | pair anas- tomosing, next pair to short sinus-membrane; lower surface of costae covered with erect thick pale hairs to | mm long, similar hairs sparse on costules; sparse glands present on costules and veins; upper surface of costae densely hairy, similar hairs more sparse on costules and veins, some erect hairs between veins. Sori medial; in- dusia small, bearing glands and short hairs; sporangia setiferous; spores with many small wings. Distr. Malesia: Papua New Guinea (Salawaket Range); 2 collections. Ecol. “In low open forest on landslide, lime- stone’. 2500-3200 m. 91. Sphaerostephanos rigidus (RIDL.) HOLTTUM, comb. nov.—Goniopteris rigida RtIDL. Trans. Linn. Soc. Bot. 9 (1916) 258. — Phegopteris wol- lastonii v.A.v.R. Handb. Suppl. (1917) 515, nom. nov., not P. rigida (HOOK. & GREV.) METT. — Thelypteris rigida (RIDL.) REED, Phytologia 17 (1968) 309.— Type: C. B. KLOss s.n. 18 Feb. 1913, W. New Guinea, Mt Carstensz, Camp VIc, 1680 m (BM; K). Caudex lacking from type and stipe incomplete. Lamina to 40cm long; pinnae to 30 pairs; basal 3-4 pairs pinnae gradually decrescent, lowest 2.2 x 0.6 cm, below them 3 pairs reduced pinnae 3-4 mm long; aerophores c. 0.5mm long. Rachis bearing throughout stiff spreading brown hairs to 1.2mm long. Largest pinnae 6.0 1.3cm; base truncate and slightly dilated both sides; apex narrowed to a short obtuse or rounded tip; edges lobed 1/4-1/3 to costa, lobes rounded with edges strongly reflexed (lower surface thus concave); costules 2.5mm apart, pale and prominent on lower sur- face, grooved above; veins 4-5 pairs, prominent beneath, one pair anastomosing, next acroscopic vein to sinus-membrane; stiff brown hairs to 0.7mm long on lower surface of costae, more sparse on costules, veins and edge, glands present on costae and costules; upper surface of pinnae glabrous apart from sparse short hairs on costae. Sori medial; indusia very small with a few stiff hairs (sometimes absent?); sporangia with neither glands nor hairs on body, yellow glands some- times on their stalks. Distr. Malesia: Western New Guinea; 2 col- lections at 1700 m; on limestone? Note. The second collection, EYMA 4986 from Wissel Lake region, is small, with lamina of fronds 15cm long, pinnae to 2 cm long, 3-4 pairs lower ones gradually reduced, lowest 8 mm long; the caudex is short-creeping. It seems probable that the type was found on limestone. 92. Sphaerostephanos arfakianus (BAK.) HOLT- TUM, comb. nov.— Polypodium arfakianum BAK. in Beccari, Malesia 3 (1880) 45.— Dry- opteris arfakianus (BAK.) C. CHR. Ind. Fil. (1905) 253; Dansk Bot. Ark. 9, n. 3 (1937) 50. — Phegop- teris arfakiana (BAK.) v.A.v.R. Handb. (1908) 502.— Cyclosorus arfakianus (BAK.) COPEL. Gen. Fil. (1947) 142; Philip. J. Sci. 78 (1951) 451. — Thelypteris arfakiana (BAK.) REED, Phy- tologia 17 (1968) 260. — Type: BECCARI s.n. 1872, W. New Guinea, Mt Arfak at Putat (FI; K). Dryopteris sepikensis BRAUSE, Bot. Jahrb. 56 (1920) 101; COPEL. Philip. J. Sci. 78 (1951) 443. — Type: LEDERMANN 12053, N.E. New Guinea, Schraderberg (B). Dryopteris arborea v.A.v.R. Bull. Jard. Bot. Btzg II, 28 (1918) 24, non BRAUSE 1914. — Dry- opteris rosenburghii C. CHR. Ind. Fil. Suppl. III (1934) 96. — Type: RUTTEN 161, Ceram (BO). Caudex slender, erect, to at least 100cm tall. Stipe 10-20 cm long, dark, short-hairy. Lamina to 80cm long, + dimorphous, thin; pinnae to 40 pairs, lowest 6-10 pairs gradually smaller, lower ones broadly triangular with slightly asymmetric base, entire; aerophores not elongate. Hairs on both sides of rachis less than 0.5 mm_ long, brownish, + appressed. Largest sterile pinnae 12- 17x 2.2-4cm, fertile commonly to 10x 1.5cm, sometimes larger; base broadly cuneate to trun- cate, sometimes a little dilated both sides; apex acuminate; edges shallowly crenate or sinuous; costules to 4.5mm apart on large pinnae; veins 5-7 pairs, slender, concolorous, slightly prominent both sides, almost all anastomosing to form zig- zag excurrent veins between costules; hairs on lower surface of costae 0.1-0.2 mm long, slender, appressed, similar hairs on costules and veins, also sometimes a few glands; hairs on upper sur- face of costae 0.3-0.4 mm long, on costules shor- ter and sparse, usually a few slender appressed hairs on surface between veins at least near mar- 1981] gin. Sori inframedial; indusia small, thin, usually with a few hairs; sporangia bearing glands. Distr. Malesia: Moluccas (Ceram;? Amboina, BROOKS 18102) and widely in New Guinea. Ecol. In forest at altitudes to 2000 m. Note. The type of D. sepikensis and some others from an altitude of about 2000m are smaller than BECCARI’s type and other lowland specimens, with fewer reduced pinnae and dis- tinctly lobed sterile ones, but agree in details of pubescence and sori. 93. Sphaerostephanos archboldii (C. CHR.) HOLTTUM, comb. nov.— Dryopteris archboldii C. CHR. Brittonia 2 (1937) 297.— Thelypteris archboldii (C. CHR.) REED, Phytologia 17 (1968) 260. — Type: BRASS 4873, Papua, Mt Tafa, valley forest, 2400 m (BM; BRI). Dryopteris protecta COPEL. Univ. Cal. Publ. Bot. 18 (1942) 221.—Cyclosorus protectus (COPEL.) COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78 (1951) 453, pl. 33. — Thelypteris protecta (COPEL.) REED, Phytologia 17 (1968) 306. — Type: BRASS 10933, New Guinea, 9km N.E. of Lake Habbema, 2800m (MICH; BM, L). — Fig. 13d-e. Caudex slender, erect, to 100cm or more tall. Stipe 5-15cm long, minutely hairy, when young covered with rather firm brown scales 7 X 1mm. Lamina to 65cm long, texture very firm; lower 8-15 pairs pinnae gradually smaller, lowest 3-6 pairs of these more widely spaced with asym- metric base, lowest c. 1 mm long; aerophores not elongate. Hairs on lower surface of rachis erect, to 0.5mm long, on upper surface a little longer, also spreading. Largest pinnae of type to 6x 10cm (type of D. protecta 9x1.5cm); base truncate; apex evenly attenuate; edges lobed about 1/3 to costa; costules to 3mm apart; veins 4-6 pairs, prominent on lower surface, 1-1} pairs anastomosing, next acroscopic vein or pair to short sinus-membrane; hairs on lower surface of costae 0.1-0.3 mm long, antrorse distally, sparse glands and hairs on costules and veins (glands best seen on Sterile fronds); hairs on upper surface of costae short, sparse on costules, veins and be- tween veins. Sori near costules; indusia bearing hairs and glands; sporangia sometimes with a seta. Distr. Malesia: Eastern New Guinea at 1800— 2800 m, many collections. 94. Sphaerostephanos_ tibangensis (C. CHR.) HOLTTUM, comb. nov.— Dryopteris tibangensis CG Gur. Dansk Bot Ark 9) 13° (1937)66.— Pronephrium tibangense (C. CHR.) HOLTTUM, Blumea 20 (1972) 118.—Type: MJOBERG s.n. Oct.-Dec. 1925, E. Kalimantan, Mt Tibang, 1400- 1700 m (BM; S-PA). Caudex short, suberect; stipe to 20cm long, short-hairy. Lamina to 20cm long; no reduced basal pinnae; normal pinnae 8-10 pairs, lower THELYPTERIDACEAE (Holttum) 485 ones with stalks to 2mm long; basal pinnae slightly shorter than next pair, with 3 pairs of basal basiscopic lobes slightly reduced. Largest pinnae 3.5 1.4cm; base truncate and slightly auricled on the acroscopic side, narrowed rather evenly from base to acute apex; edges lobed 2/5 towards costa or rather more deeply; lobes sub- falcate, acute; costules 3mm apart, at more than 60° to costa; veins 6 pairs, basal pair anastomos- ing, second pair to sides of sinus-membrane, rest to margin; lower surface of costae and costules bearing pale spreading hairs 0.2-0.6mm _ long, short erect hairs present between veins, many glands on costules and veins; upper surface of costae copiously short-hairy, rest of surface bear- ing short suberect hairs. Sori supramedial; indusia small, densely short-hairy; sporangia not seti- ferous. Distr. Malesia: 1400-1700 m. Note. This is certainly very near S. norrisii. The type consists of one fertile and one sterile frond, the latter not quite fully expanded, with immature sori. The fertile frond is smaller than any specimen undoubtedly referable to S. norrisii, and the pinnae are of a different shape; mature sori might show further differences. East Borneo (Mt Tibang), 95. Sphaerostephanos norrisii (ROSENST.) HOLTTUM, comb. nov.— Dryopteris — norrisii ROSENST. Med. Rijksherb. n. 31 (1917) 8.— Type: W. NORRIS, Malaya (L; K). Nephrodium pennigerum var. malayense BEDD. Handb. Suppl. (1892) 74, excl. Parish, Tenasserim. — Dryopteris indica var. malayensis (BEDD.) v.A.v.R. Handb. (1908) 224. — Lectotype (selected here): KUNSTLER 2360, Perak (K). Dryopteris subfalcinella v.A.v.R. Bull. Jard. Bot. Btzg III, 2 (1920) 151. — Thelypteris subfal- cinella (v.A.v.R.) REED, Phytologia 17 (1968) 317. — Lectotype (selected here): LORZING 5338, Sumatra, Bandar-baroe (BO; L). Dryopteris elmerorum COPEL. Philip. J. Sci. 40 (1929) 295, pl. 2.—Cyclosorus elmerorum (COPEL.) COPEL. Gen. Fil. (1947) 142; Fern FI. Philip. (1960) 351.— Thelypteris elmerorum (COPEL.) REED, Phytologia 17 (1968) 274. —S. elmerorum (COPEL.) HOLTTUM, Kalikasan 4 (1975) 53.—Type: COPELAND s.n. Nov. 1911, Mindanao, San Ramon (MICH). Nephrodium indicum sensu RIDL. J. Mal. Br. R. As. Soc. 4 (1926) 73. Dryopteris toppingii sensu C. CHR. Gard. Bull. Str. Settl 4 (1929) 391.—Cyclosorus toppingii sensu HOLTTUM, Rev. FI. Mal. 2 (1955) 280, f. 161. Cyclosorus ellipticus sensu COPEL. Fern FI. Philip. (1960) 351. — Fig. 12f-h. Caudex short-creeping. Stipe 30-90cm long, dull reddish, minutely hairy; reduced pinnae 0-3 pairs, present only on largest fronds, irregularly spaced and very small. Lamina to 100cm long 486 (type), on type of D. elmerorum 30cm long; pin- nae 10-18 pairs, several pairs distinctly stalked; basal pinnae often somewhat reduced if small reduced pinnae absent, with stalks 1-3 mm long, narrowed towards their bases, not auricled; aero- phores not elongate. Hairs on lower surface of rachis dense, short, stiff, erect; on upper surface copious, appressed, less than 0.5 mm long. Largest pinnae of type 24x3.4cm, of type of D. elmerorum 6.5 X 1.8cm; base subtruncate on large fronds, subcordate on small ones; apex rather abruptly short-acuminate; edges lobed c. 2/5 towards costa (less deeply on small fronds), lobes falcate with broadly pointed or rounded forward- pointing tips; costules 5-6 mm apart (3.5-4 mm in type of D. elmerorum); veins to 12 pairs, 1-13 pairs anastomosing, 2-23 pairs passing to sides of sinus-membrane; hairs on lower surface of costae and costules dense, short, erect, some short hairs also between veins, glands present on costules at least near apex of pinna-lobes; hairs on upper surface of costae and costules short, some short appressed or suberect hairs on surface between veins. Sori supramedial, somewhat elongate, especially the lower ones; indusia large, densely short-hairy, sometimes with a _ few glands; sporangia sometimes with a gland or a short seta. Distr. Malesia: Malaya, Sumatra, Borneo, Philippines (Mindanao). Ecol. In forest at 1000-1500 m; apparently not common anywhere. Note. Though the type of D. elmerorum is small, other specimens from Mindanao have pin- nae to 15x3cm. No small fertile fronds have been found on plants in Sumatra and Malaya. Some specimens have very few glands on the lower surface of costules. Specimens from Suma- tra and Malaya have appressed hairs between the veins on the upper surface, specimens from Bor- neo and Mindanao have shorter suberect hairs. 96. Sphaerostephanos semimetralis HOLTTUM sp. nov. S. arfakiano affinis, differt: frondibus minori- bus; pinnis redactis 2-3-jugatis, minutis; pinnis fertilibus usque 5X1.2cm; costis costulisque subtus pilis sparsis suberectis instructis. — Type: PULLE 518, W. New Guinea, Perameles Mts, 1100 m, on limestone (L). Caudex erect, slender, 50cm tall. Stipe 15cm long, basal scales c. 8 x 1 mm; base of stipe to first large pinna 25-30cm. Reduced pinnae 2-3 pairs, all very small, with above them an apparently much larger intermediate pair (incomplete). Lamina 35 cm long; pinnae 10-12 pairs, very firm, dimorphous; basal pinnae not narrowed at base; aerophores less than 0.5 mm long. Hairs on lower surface of rachis thick, brown, 0.5mm long, on upper surface similar, to 1 mm long. Largest sterile pinnae 9X2.2cm; base truncate with a winged stalk 1mm long; apex abruptly short-acuminate; FLORA MALESIANA [ser. II, vol. 1° edges crenate to a depth of 1-2mm; costules 3.5-4mm apart, at 60°; veins 6-8 pairs, very oblique except basal ones, prominent both sides, 2 pairs anastomosing to form a zig-zag excurrent vein, | pair to short sinus-membrane; lower sur- face of costae bearing sparse erect hairs 0.1 mm long, similar hairs very sparse on costules and veins with a few glands, sparse erect hairs be- tween veins; hairs on upper surface of costae pale, 0.5 mm long, rest of surface glabrous. Fertile pinnae to 5 x 1.2. cm; veins 5 pairs; pubescence as sterile; sori near costules; indusia small with a few short hairs and glands; sporangia sometimes with a gland. Distr. Malesia: New Guinea. A specimen from N.E. New Guinea, Western Highlands, at Kompiai in the Jimmi valley at 1900m (MANNER & STREET 454), probably not from limestone, is similar to the type. Note. This differs from the mountain form of S. arfakianus in the very small reduced pinnae, and in pubescence of lower surface; but better specimens are needed to characterize it clearly. 97. Sphaerostephanos semicordatus HOLTTUM, sp. nov. Pinnae redactae 8-jugatae, omnes minutae; lamina 35 cm longa; pinnae 10-12-jugatae, omnes oppositae, subintegrae; costulae subtus glabrae, glanduliferae; sori exindusiati; sporangia Setifera; sporae alatae. —Type: PULLEN 1476, N.E. New Guinea, Sepik Distr., Prince Alexander Range, 900 m (CANB). Caudex short, creeping. Stipe 8cm long, red- dish, glabrescent, basal scales 3 x 1.5 mm; base of stipe to first large pinna 60 cm. Reduced pinnae 8 pairs or more, 5-6cm apart, all very small. Lamina 35cm long; pinnae 10-12 pairs, all opposite, basal pinnae not narrowed at their bases; aerophores slender, to 1 mm long. Hairs on rachis both sides 0.5mm long, on upper surface brown. Largest sterile pinnae 14x2.5cm, fertile to 10x 2.0cm; basiscopic base cordate and over- lapping rachis, acroscopic truncate or slightly cordate; apex acuminate; edges irregularly sinu- ous to slightly crenate; costules 4.5 mm (sterile) 3.5mm (fertile) apart, at more than 60°; veins 6-7 pairs, 32 pairs anastomosing, one vein to short sinus-membrane; lower surface of costae minutely hairy, rest of lower surface glabrous, glands present on costules; on upper surface base of costae covered with brown hairs as rachis, above base spare short hairs, rest of upper surface glabrous. Sori on distal veins medial, on lower ones supramedial, those on veins from adjacent costules often coalescent; no indusia; sporangia with 4-6 setae; spores with a + continuous trans- lucent wing and a few irregular other wings. Distr. Malesia: Papua New Guinea (Sepik). (v.A.v.R.) 98. Sphaerostephanos _ pterosporus 1981] THELYPTERIDACEAE (Holttum) 487 HOLTTUM, comb. nov. — Dryopteris pterospora v.A.v.R. Bull. Jard. Bot. Btzg III, 2 (1920) 148. — Thelypteris pterospora (v.A.v.R.) REED, Phy- tologia 17 (1968) 307.—Type: BRooKs 447, Sumatra, Benkoelen, Tambang Sawah (BO; BM). Dryopteris bungoensis C. CHR. Dansk Bot. Ark. 9, n. 3 (1937) 59. — Type: BROOKS 10, April 1909, Sarawak, Bungo Range (BM). KEYerO THE VARTIEERIES 1. Pinnae lobed at least 1/3 towards costa. 2. Pinnae lobed less than 1/2 towards costa; basal pair of veins often anastomosing a. var. pterosporus 2. Pinnae lobed 3/5—2/3 towards costa; basal veins rarely anastomosing . b. var. altilobus 1. Pinnae crenate . ¢. var. crenatus a. var. pterosporus Caudex short, erect. Stipe 3-10 cm long, basal scales 8 X 1 mm; base of stipe to first normal pinna 20-30 cm (sterile fronds) 30-45 cm (fertile fronds). Reduced pinnae 6-8 pairs, all less than 2 mm long. Lamina c. 40cm long; pinnae to 18 pairs, basal pinnae narrowed to a width of 6-8 mm or less at base; aerophores less than | mm long. Hairs on lower surface of rachis of sterile fronds thick, brown, curved, to | mm long, on fertile paler and shorter; hairs on upper surfaces as on lower sur- face of sterile. Largest sterile pinnae to 12 x2cm; base truncate; apex rather abruptly caudate-acu- minate; edges lobed less than 1/2 towards costa, lobes falcate with forward-pointing tips; costules 4.5-5 mm apart, at 60° to costa; veins 5-7 pairs, all very oblique, basal pair uniting to form an excur- rent vein to sinus or touching just below sinus, next acroscopic vein to side of sinus-membrane or to margin; hairs on lower surface of costae 0.3 mm long, pale, closely appressed, on costules and veins similar, sometimes also a few glands, on surface between veins some fine appressed hairs may be present but not glands; hairs on upper surface of costae 0.5 mm long with some to | mm, on costules and veins short and sparse, sometimes a few appressed hairs between veins. Fertile pin- nae to 7X 1.4cm, lobed a little more deeply than sterile; costules 3.5mm apart; basal veins more often connivent at the sinus without joining. Sori medial; indusia firm with short hairs; sporangia bearing glands; spores with many small separate wings. Distr. Malesia: Southern Malaya (several localities), S. Sumatra, W. Sarawak, in forest at 9-700 m. b. var. altilobus HOLTTUM, var. nov. A typo speciei differt: pinnis maximis sterilibus usque 14X2.6cm, fertilibus 10 1.8 cm, 3/5-2/3 costam versus lobatis; venis infimis plerisque ad sinum conniventibus, raro anastomosantibus. — Type: MOLESWORTH ALLEN 2736, Perak, Maxwell’s Hill, 730m, in forest, “deep green fronds, brown rachis, quite distinct” (K). Also from Taiping Hills: KUNSTLER 6345, at 750- 900 m, “‘pinnae dark green”. Distr. Malesia: Malaya. c. var. crenatus HOLTTUM, var. nov. A typo speciei differt: pinnis crenatis; venis 1 : = l5-jugatis anastomosantibus, vena sequente acroscopica ad membranam sinus terminata; pinnis fertilibus usque 101.5 cm.—Type: JERMY 13870, Sarawak, G. Mulu National Park, by Tapin River (BM). Also M. HoTTa 13809, Brunei, along Sungei Lacquan, 50-300 m (L). Distr. Malesia: Borneo (Sarawak; Brunei). 99. Sphaerostephanos sagittifolius (BL.) HOLT- TUM, comb. nov.— Aspidium sagittifolium BL. Enum. Pl. Jav. (1828) 153; MIQUEL, Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 150.— Nephrodium sagittifolium (BL.) Moore, Ind. Fil. (1858) 103; RACIB. FI. Btzg 1 (1898) 191. — Dryopteris sagit- tifolia (BL.) O. KTZE, Rev. Gen. Pl. 2 (1891) 813; v.A.v.R. Handb. (1908) 222; BACKER & POSTH. Varenfl. Java (1939) 52.— Thelypteris sagittifolia (BL.) REED, Phytologia 17 (1968) 311.— Type: BLUME, Java (L, n. 908, 336-24). Caudex short, massive, decumbent or suberect: young fronds covered with slime. Stipe 15cm long; scales (fide RACIBORSKI) 4 X 3 mm; base of stipe to first large pinna 50-80 cm. Reduced pinnae more than 30 pairs, 1.5-2 cm apart, deflexed, up- per ones 2.5-3.0cm long with basal auricle to 1.5cm, apex acuminate, edges crenate: lowest lcm long. Lamina 100cm or more long; several pairs lower pinnae with elongate basal acroscopic auricle in which veins are forked; aerophores thick but hardly 1 mm long. Rachis covered with thick spreading hairs more than | mm long, both sides. Largest pinnae to 20 2.3cm; base trun- cate, basal acroscopic lobe somewhat elongate: apex caudate-acuminate; edges lobed 2/5 to costa, lobes with falcate tips; costules 5-5.5 mm apart at more than 60°; veins to 10 pairs, pale and prom- inent on lower surface, 1:2 pairs anastomosing, 2-23 pairs to sides of sinus-membrane; hairs on lower surface of costae stiff, spreading, of varying length to 1 mm, on costules similar but sparse, a few glands on costules and veins; hairs on upper surface of costae 1mm long, scattered similar hairs on costules and veins, no others. Sori medial, basal ones divergent; indusia small with stiff hairs 0.3mm long; sporangia bearing glands or setae; spores with many small wings. Distr. Malesia: Sumatra, Java, and Lesser Sunda Is. (Lombok). Ecol. In forest at 1000-1500 m. Note. v.A.v.R. (Handb. Suppl. 182) reported specimens from New Guinea; I did not find these in Herb. Bog. 488 FLORA MALESIANA [ser. II, vol. P? 100. Sphaerostephanos foliolosus HOLTTUM, sp. nov.— Aspidium hirsutum and A. cucullatum sensu CHRIST, Ann Jard. Bot. Btzg 15 (1898) 134, quoad Sarasin 745 tantum. S. polyoti affinis, differt: pinnis c. 2/5 costam versus lobatis; venis inferioribus 23-3-jugatis vel anastomosantibus vel ad membranam sinus con- niventibus; aerophoris usque 1mm longis; pinnis redactis superioribus minoribus.— Type: P. & F. SARASIN 745, N. Celebes, G. Matinang 350m (BAS). Stipe to 15cm long; scales broad, thin; base of stipe to first large pinna 60cm. Reduced pinnae c. 30 pairs, 1-1.5 cm apart, upper ones to 12x 6mm, lowest 4-8mm long, shape as in S. polyotis, covered with mucilage when young. Lamina 65- 70cm long; pinnae 30 pairs, lower ones not nar- rowed at base; aerophores slender, to 1 mm long. Lower surface of rachis densely short-hairy (sometimes with longer hairs?); brown hairs to 1mm long on upper surface. Largest pinnae 15 x 1.8cm, base subtruncate, symmetric (more cuneate and less symmetric on upper pinnae); apex gradually attenuate, not caudate; edges lobed c. 2/5 to costa, lobes falcate with obtusely pointed tips; costules 3mm apart, at 60° or more; veins 8-10 pairs, 13 pairs anastomosing, 1+-2 pairs pass- ing to sides of sinus-membrane; hairs on lower surface of costae short, dense, mixed with long ones on sterile fronds, sparse hairs with glands on costules and veins, short erect hairs between veins; upper surface of costae bearing pale hairs 1 mm long, rest of surface + closely covered with slender appressed hairs. Sori medial; indusia small, pilose; sporangia usually with 1-2 glands, sometimes a seta. Distr. Malesia: N. Celebes, Moluccas (Talaud Islands: LAM 2786, 2788, on river bank at 80- 350 m). Note. LAM’s sterile specimens differ from the type (which is fertile) in the presence of hairs 1 mm long on lower surface of rachis and costae. 101. Sphaerostephanos nov. Pinnae redactae 6—7-jugatae, superiores 12mm longae; lamina 45cm longa; pinnae maximae 9.5 1.8cm, 3/4 costam versus lobatae; costae, costulae venaeque subtus glandulosae, pagina in- ter venas pilosa; pagina superior pilis appressis vestita; indusia glandulis et pilis brevibus praedita; sporangia nec glandulis nec setis praedita.— Type: A. C. JERMy 5248, N.E. New Guinea, Western Highlands, Keglsugl (BM). Caudex short, creeping. Stipe 15—20cm long, minutely hairy, basal scales broad, thin; base of stipe to first large pinna 45-50 cm. Reduced pinnae 6-7 pairs, 34cm apart, lowest 5-7 mm long, up- permost 12mm long, somewhat deflexed, nar- rowly triangular, with auricle 6-8mm long on acroscopic base. Lamina 45 cm long; pinnae to 24 alpinus HOLTTUM, sp. pairs; basal pinnae not or little narrowed at their bases; aerophores thick, hardly 1 mm long. Hairs on lower surface of rachis dense, short, spread- ing, on upper surface | mm long. Largest pinnae 9.5cm long, 1.8cm wide above slightly dilated base; apex acuminate with cauda 1.5cm; edges lobed almost or quite 3/4 to costa, lobes slightly falcate; costules 3.5-4mm apart, their bases al- most at right angles to costa; veins 9 pairs, at a wide angle, pale and prominent both sides, basal pair anastomosing, next pair usually both to mar- gin; lower surface of costae densely covered with short spreading hairs, hairs less dense, with glands, on costules, surface between veins with short erect hairs; hairs | mm long on upper sur- face of costae, similar hairs scattered on costules and veins, whole upper surface covered with ap- pressed hairs. Sori medial; indusia bearing glands and short hairs; sporangia with neither glands nor setae; spores with many small wings. Distr. Malesia: Papua New Guinea. Ecol. Type locality at 2500-2750m, and Mt Wilhelm at 3300-3500 m (NAKAIKE 274). 102. Sphaerostephanos _ polisianus HOLTTUM, Kalikasan 4 (1975) 65.— Type: M. G. PRICE 431, Luzon, Mountain Prov. Mt Polis (CAHUP; K). Caudex creeping, 5-6mm diameter, bearing a close series of fronds. Stipe 4-5 cm long, minutely hairy, basal scales short and firm; base of stipe to first large pinna 25-30 cm. Reduced pinnae 34cm apart, all very small. Lamina to 28 cm long; pin- nae 14 pairs, lowest narrowed at their bases; aerophores 1mm long. Lower surface of rachis covered with very short erect reddish hairs, upper surface with thick curved dark red hairs 0.5 mm long. Largest pinnae 6.5 x 1.3cm; base subtrun- cate; apex short-acuminate; edges lobed > way to costa, lobes oblique with forward-pointing obtuse tips; costules 3-3.5mm apart at 60°; veins 6-7 pairs, slender and prominent on lower surface, basal pair anastomosing, next acroscopic vein to side of short sinus-membrane; lower surface of costae covered with hairs 0.1 mm long or a little more, antrorse at least distally, few similar hairs on costules, glands on costules and veins; hairs on upper surface of costae 0.3 mm long, rest of upper surface glabrous. Sori medial; indusia small, glabrous; sporangia sometimes with a gland. Distr. Malesia: Philippines (Luzon); known from the type. Note. This should perhaps be regarded as a form of S. lobatus. only 103 Sphaerostephanos hoalensis HOLTTUM, sp. nov. — Dryopteris ceramica ROSENST. MS, non v.A.v.R. Pinnae redactae c. 13-jugatae, superiores 15 x 8mm, infimae 10mm longae; aerophora elon- gata; pinnae maximae 11 X2cm, dimidio costam versus lobatae; costae subtus pilis erectis usque 1981] THELYPTERIDACEAE (Holttum) 489 Imm longis vestitae, costulae pilis brevioribus glandulisque praeditae; pagina superior pilis ap- pressis vestita; indusia_ pilifera; sporangia glandulifera. — Type: STRESEMANN 110, Ceram, Central Mts, G. Hoale, 1000 m (L). Stipe lacking. Reduced pinnae 2.5 cm apart, c. 13 pairs, one near the base 10 mm long, uppermost 15Sx8&mm, deflexed, slightly auricled on acros- copic base, edges crenate, apex obtuse to roun- ded. Lamina c. 60cm long; lowest pinnae not narrowed at base; aerophores elongate. Lower surface of rachis covered with erect hairs 0.2- 0.3 mm long and some longer ones, hairs on upper surface | mm long. Largest pinnae 11 x 2. cm; base truncate; apex abruptly short-acuminate; edges lobed c. 3 way to costa, lobes oblique, slightly falcate; costules 3mm apart, at more than 60°; veins 9 pairs, 1> pairs anastomosing, next acros- copic vein to sinus-membrane; lower surface of costae hairy as rachis, costules and veins less densely covered with shorter hairs and some glands, surface between veins bearing short erect hairs; hairs on upper surface of costae 1 mm long, shorter ones on costules, appressed hairs all over surface. Sori small, medial to supramedial, lower ones a little divergent; indusia with many rather long hairs; sporangia bearing | or 2 glands. Distr. Malesia: Moluccas (Central Ceram, 2 collections, second one n. 57). Ecol. In the mountains, 1000 m. 104. Sphaerostephanos larutensis (BEDD.) C. CHR. Ind. Fil. Suppl. III (1934) 172.— Nephrodium larutense BEDD. Handb. Suppl. (1892) 73.— Mesochlaena larutensis (BEDD.) v.A.v.R. Handb. (1908) 232, incl. var. borneensis. — Cyclosorus larutensis (BEDD.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 245; HOLTTUM, Rev. FI. Mal. (1955) 284, f. 165.—Thelypteris larutensis (BEDD.) REED, Phytologia 17 (1968) 286. — Lec- totype (here selected): KUNSTLER 2398, Perak, Larut (K). Mesochlaena sumatrensis v.A.v.R. Bull. Jard. Bot. Btzg III, 2 (1920) 160.—S. sumatrensis (v.A.v.R.) C. CHR. Ind. Fil. Suppl. III (1934) 172. — Type: BROOKS 391, Sumatra, Benkoelen, Lebong Tandai (BO; BM). — Fig. 12d-e. Caudex short, creeping. Stipe 20-30cm long, above base covered with spreading hairs to 1 mm long; base of stipe to first large pinna 40-70 cm. Reduced pinnae 6-12 pairs, lower ones less than lcm long, upper ones c. 1.5 x 1.2 cm, spreading, triangular, edges crenate, apex acute; an inter- mediate pair sometimes present. Lamina 50- 70cm long; apex pinna-like; pinnae c. 12 pairs, well spaced; lower pinnae narrowed towards their bases; middle pinnae with truncate bases, upper ones broadly cuneate; aerophores not elongate. Rachis covered throughout with slender light brown hairs !-1.5mm long. Largest pinnae 20- 30 cm long, 2.5—-3.8 cm wide, acuminate with cauda 1-3 cm long; edges lobed 1/4~1/3 to costa, lobes falcate with forward-pointing obtuse tips; costules 4.5-6 mm apart; veins to 11 pairs, 13-2 pairs anas- tomosing, next 2 pairs to sides of long sinus- membrane; lower surface of costae covered with erect hairs 0.5-1mm long, shorter hairs on cos- tules, veins and between veins, some glands with the hairs on costules and veins; upper surface of costae covered with fine pale antrorse hairs, on costules and veins much shorter hairs with scat- tered long ones, surface between veins glabrous or with suberect hairs. Sori medial, about twice as long as wide; indusia large, thin, bearing slender hairs and glands; sporangia bearing neither glands nor setae on body, hair on stalk of several cells, terminal one acicular and thick-walled; spores with many small wings. Distr. Malesia: Malaya, Sumatra, Borneo. Ecol. In forest at 50-1000 m. Note. v.A.v.R. distinguished var. borneensis which has many glands and no hairs on indusia. His M. sumatrensis was based on a rather small specimen. Glands are apparently sometimes lack- ing on lower surface of costules. 105. Sphaerostephanos loherianus (CHRIST) HOLTTUM, Kalikasan 4 (1975) 61.— Aspidium loherianum CHRIST, Bull. Herb. Boiss. 6 (1893) 191.— Dryopteris loheriana (CHRIST) C. CHR. Ind) Fil 1@905), 275: CurismePhilipa ws Scie 2 (1907) Bot. 207; v.A.v.R. Handb. (1908) 221, 820. — Cyclosorus loherianus (CHRIST) COPEL. Gen. Fil. (1947) 142; Fern FI. Philip. (1960) 342. — Thelypteris loheriana (CHRIST) REED, Phytologia 17 (1968) 289. — Type: LOHER 900, Luzon, Mon- talban, Oct. 1890 (P; K). Caudex creeping, sometimes with well-spaced fronds. Stipe 5-30cm long, reddish, glabrescent; base of stipe to first large pinna 40-50 cm. Reduced pinnae 5-15 pairs, defiexed, strongly auricled, uppermost 6-10mm long, lowest 2- 5mm. Lamina to 60cm or more long; pinnae well spaced, several lower pairs narrowed towards their bases; aerophores slightly swollen. Lower surface of rachis densely covered with short erect hairs, those on upper surface to 1mm _ long. Suprabasal pinnae of type 11 x 1.8 cm, of largest specimen 303.2cm; basal lobes somewhat reduced; apex acuminate, sometimes with cauda to 2cm; edges lobed to 2mm from costa, lobes slightly falcate, narrowed gradually from base; costules 44.5 mm apart, almost at right angles to costa; veins of type to 14 pairs, of largest speci- men 24 pairs, basal veins near base of pinna often anastomosing to form a short excurrent vein, those near apex of pinna connivent at the sinus without uniting; lower surface of costae densely covered with erect hairs 0.3-0.5 mm long, on some specimens to | mm, on costules and veins shorter hairs and some glands, short erect hairs between veins; hairs on upper surface of costae to 1mm 490 long, scattered similar hairs on costules and veins, between veins a variable number of short hairs. Sori all close to margin; indusia small, covered with short hairs and sometimes a few glands; sporangia setiferous, rarely with a gland. Distr. Malesia: Philippines (N. Luzon). Ecol. In mountain forest, 1500-2000 m; smaller specimens apparently from streamsides in open places. 106. Sphaerostephanos erectus (COPEL.) HOLT- TUM, Kalikasan 4 (1975) 64. — Cyclosorus erectus COPEL. Philip. J. Sci. 81 (1952) 30, t. 22; Fern FI. Philip. (1960) 346. — Thelypteris erecta (COPEL.) REED, Phytologia 17 (1968) 275. — Type: RAMOS BS 41550, Leyte, Cabalian (US; B). Caudex slender, erect, to 100cm tall. Stipe 10cm long, reddish, minutely hairy; base of stipe to first large pinna 45cm. Reduced pinnae to at least 6 pairs, uppermost 5mm long. Lamina to 50cm long; pinnae 15 pairs, lower ones narrowed on basiscopic side near base; apex of frond nar- rowly triangular and deeply lobed with abrupt transition to pinnae; aerophores not elongate. Hairs on rachis, both sides, to | mm long, thick, brown. Suprabasal pinnae to 14x2.8cm; base truncate; apex rather abruptly caudate-acuminate; edges lobed 3/5-2/3 to costa, lobes falcate; cos- tules 4-5.5mm apart, at more than 60°; veins 10-12 pairs, prominent on lower surface, basal pair anastomosing, second pair touching sides of sinus-membrane; hairs on lower surface of costae spreading, of varied length to 1mm, on costules shorter with glands; hairs on upper surface of costae 1mm long, on costules short, sometimes with a few glands, rest of upper surface glabrous. Sori medial; indusium firm, large, short-hairy; sporangia bearing glands. Distr. Malesia: Philippines (Leyte, 2 collec- tions; Biliran). Ecol. In mossy forest at 1000 m (M. G. PRICE 3278). Note. M. G. PRICE informs me that SULIT PNH 20306 from Biliran, which he identifies with this species, has a few glands on upper surface of costules; I have seen none in this position on the specimens from Leyte. 107. Sphaerostephanos stresemannii HOLTTUM, sp. nov. Basis frondis imperfecta; pinnae infimae 12 cm longae, medio 3.2 cm latae, basin versus angus- tatae; pinnae suprabasales usque 10X2.7cm, oppositae, 3/4 costam versus lobatae; costae sub- tus pilis erectis usque 0.5 mm longis vestitae, cos- tulae venaeque pilis sparsis glandulisque praedi- tae; pagina superior inter venas pilis brevibus suberectis instructa; sori exindusiati; sporangia setifera.— Type: STRESEMANN 416, Moluccas, Buru, G. Fogha 1400 m (L). Caudex and stipe lacking. Reduced pinnae at FLORA MALESIANA [ser. II, vol. 1° least 2 pairs, 3-4mm long; above them are an intermediate unequal pair 1.0 and 2.0cm long. Lamina 42cm long; apex narrowly deltoid with gradual transition to upper pinnae; pinnae 15 pairs, almost opposite; basal pinnae largest, 12 x 3.2cm, widest in middle and narrowed to base both sides, basiscopic lobes longer than acros- copic; aerophores swollen, not elongate. Hairs on lower surface of rachis dense, short, erect, with scattered longer ones to 0.7mm, hairs on upper, surface longer and appressed. Suprabasal pinnae to 10x3.0cm; base truncate; apex short-acu- minate; edges lobed fully 3/4 to costa, lobes hardly falcate, slightly tapered; costules 4.5mm apart at more than 60°; veins 10 pairs, slender, pale both sides, | pair anastomosing, next acros- copic veins to side of short sinus-membrane; hairs on lower surface of costae as on rachis, on cos- tules and veins more sparse and shorter spreading hairs with glands; short erect hairs on surface between veins; hairs on upper surface of costae more than 0.5 mm long, similar hairs scattered on costules and veins, between veins short suberect hairs. Sori medial; indusia lacking; sporangia copiously setose; spores with many small wings. Distr. Malesia: Moluccas (Buru), 1400 m. 108. Sphaerostephanos simplicifolius (J. SM. ex HoOoK.) HOLTTUM, Kalikasan 4 (1975) 61.— Aspidium simplicifolium J. SM, ex HOoK. Ic. Pl. 10 (1854) t. 919. — Polypodium simplicifolium (J. SM.) HOOK. Sp. Fil. 5 (1863) 2, excl. SEEMANN 736 (Fiji). — Dryopteris canescens var. C. CHR. Ind. Fil. (1905) 256. — Dryopteris simplicifolia (J. SM.) CHRIST, Philip. J. Sci. 2 (1907) Bot. 206. — Phegopteris simplicifolia (J. SM.) v.A.v.R. Handb. (1908) 500. — Cyclosorus simplicifolius (J. SM.) COPEL. Gen. Fil. (1947) 143; Fern FI. Philip. (1960) 371.— Thelypteris simplicifolia (J. SM.) REED, Phytologia 17 (1968) 314. — Type: CUM- ING 315, Samar (K; E, G, L). Caudex short-creeping; stipe to 20cm long, pale; basal scales 7 x | mm; base of stipe to first normal pinna 40-50cm; reduced pinnae to 6-7 pairs, lowest 3mm long, uppermost 10 mm, trian- gular with asymmetric base. Lamina to 50cm long; apex pinna-like, pinnae to 8 pairs, lowest somewhat narrowed at base; aerophores not elongate. Hairs on both sides of rachis pale, longest more than | mm rather sparse, with shor- ter ones. Largest pinnae to 164cm; base broadly cuneate, not auricled; apex abruptly short-acuminate; edges crenate to subentire; cos- tules 4-4.5 mm apart, at 60°; veins to 10 pairs, pale and prominent on lower surface, almost all anas- tomosing to produce a zig-zag excurrent vein; copious stiff short erect hairs on lower surface of costae with scattered long ones, on costules and veins fewer short hairs and also glands; hairs on upper surface of costae and costules short with scattered long ones. Sori near apices of veins, 1981] basal ones on veins from adjacent costules some- times confluent; indusia with many short hairs; sporangia with many glands; spores with many broad translucent wings which anastomose. Distr. Malesia: Philippines (Samar, Biliran). Note. The original description was based on a partially fertile small plant with apical lamina much larger than the pinnae. The above descrip- tion is based on fully grown plants from other collections. Both the type and larger specimens have glands on sporangia, not setae as stated by COPELAND. In Sp. Fil. vol. S HOOKER misidentified a specimen of the exindusiate Pronephrium bec- carianum (from Fiji) with the present species, for which reason he transferred the latter to Poly- podium. Leyte, 109. Sphaerostephanos spenceri (CHRIST) HOLT- TUM, Kalikasan 4 (1975) 58. — Dryopteris spenceri CHRIST, Philip. J. Sci. 2 (1907) Bot. 206; v.A.v.R. Handb. Suppl. (1917) 188.— Phegopteris spen- ceri (CHRIST) v.A.v.R. Handb. (1908) 508. — Cyclosorus spenceri (CHRIST) COPEL. Gen. Fil. (1947) 143; Fern Fl. Philip. (1960) 350. — Thelyp- teris spenceri (CHRIST) REED, Phytologia 17 (1968) 314. Type: COPELAND 1464, Mindanao, Davao Distr., Todaya (P; B, US). Caudex short, suberect or creeping. Stipe S- 10cm long, densely short-hairy; base of stipe to first large pinna 35 cm. Reduced pinnae to at least 10 pairs, lowest S-10 mm long, uppermost to 2cm long, 1.5 cm wide above auricled acroscopic base, triangular, crenate. Lamina 45 cm long; apex nar- rowly triangular, deeply lobed at base; pinnae to at least 10 pairs; basal pinnae not narrowed at base; aerophores not elongate. Hairs on both sides of rachis dense, c. 0.3mm long, with some less abundant | mm or more long. Largest pinnae 8.5 x 2.0cm; base broadly rounded to subcordate on basiscopic side, truncate to broadly cuneate on acroscopic; apex short-acuminate; edges crenate to a depth of 1mm or a little more; costules 4-5 mm apart, at 60°; veins 7-8 pairs, prominent on lower surface, 2-23 pairs anastomosing to form a Zig-zag excurrent vein, 2-2) pairs passing to sides of sinus-membrane; lower surface of costae hairy as rachis, hairs on costules shorter with some glands, short erect hairs on surface between veins; hairs on upper surface of costae 0.3-0.4 mm long, on costules shorter. Sori medial, distinctly elongate; indusia with short hairs; sporangia bear- ing glands. Distr. Malesia: Philippines (E. & S.E. Min- danao). Note. CHRIST failed to see the indusia and thus at first misled v.A.v.R. The short stipe and deeply lobed apical lamina appear to be distinctive and are shown by M. G. PRICE 2790 from Agusan del Sur Province, which is a rather small sterile plant differing from the type in scattered long THELYPTERIDACEAE (Holttum) 491 hairs on the upper surface of costules and veins and in abundant glands on lower surface of cos- tules and veins. 110. Sphaerostephanos mengenianus HOLTTUM, sp. nov. Caudex 40cm longus, erectus; pinnae omnes oppositae, basi late cuneatae; pinnae redactae 8- jugatae, maximae 22cm; pinnae normales 13.5X2.0cm, crenatae; pili paginae inferioris omnes breves; costulae subtus glanduliferae; in- dusia_ pilifera; sporangia glandulifera? (non setifera). —Type: STEVENS & LELEAN LAE 58680, E. New Britain, in Castanopsis forest, 900 m (K; L). Caudex erect, 50cm tall (collectors). Stipe 15cm long, minutely hairy, dull reddish; base of stipe to first large pinna 60cm. Reduced pinnae opposite, S-6cm apart, lowest 7mm long, up- permost 22cm, almost symmetrically rhombic with basal angle more than 90°, apical angle less than 90°, subentire; one pair intermediate pinnae 4cm long also present. Lamina 55cm long; apex not pinna-like; pinnae 20 pairs, all opposite; basal pinnae not narrowed at base; aerophores not elongate. Hairs on lower surface of rachis 0.2 mm long, slender, appressed, on upper surface thicker, antrorsely curved, 0.5mm long. Largest pinnae 13.5 2.0cm:; base broadly cuneate and slightly dilated both sides; apex acuminate, not caudate; edges crenate to a depth of 1.5 mm or little more; costules 4.5-5 mm apart, at less than 60°; veins 7-8 pairs, 2-23 pairs anastomosing to form a zig-zag excurrent vein, 1>2 pairs passing to rather long sinus-membrane; lower surface of costae covered with slender appressed hairs 0.2 mm long, similar sparse hairs with glands on costules, surface be- tween veins glabrous, eglandular; hairs on upper surface of costae 0.5 mm long, rest of upper sur- face glabrous. Sori medial; indusia rather small, thin, with short hairs; sporangia probably with glands, not setiferous. Distr. Malesia: Papua New Guinea (New Bri- tain), only the type. Ecol. In Castanopsis forest, 900 m. 111. Sphaerostephanos confertus (BRAUSE) HoOLTTUM, Webbia 30 (1976) 194. — Dryopteris conferta BRAUSE, Bot. Jahrb. 49 (1912) 22, f. IF; v.A.v.R. Handb. Suppl. (1917) 175.— Cyclo- sorus confertus (BRAUSE) COPEL. Gen. Fil. (1947) 142; HoLtTuM & Roy, Blumea 13 (1965) 134. — Thelypteris conferta (BRAUSE) REED, Phyto- logia 17 (1968) 268.—Type: SCHLECHTER 17864, N.E. New Guinea, Rani Mts 700m (B; K, EYP Ue): Dryopteris terrestris COPEL. Univ. Cal. Publ. Bot. 18 (1942) 221.—Cyclosorus terrestris (COPEL.) COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78 (1951) 454, pl. 34. — Thelypteris terrestris (COPEL.) REED, Phytologia 17 (1968) 319.— 492 Type: BRASS 13660, New Guinea, Idenburg River, in forest at 700 m (MICH; BO, L). Caudex short, creeping, branched, apices of branches with tufts of fronds. Stipe 5-15 cm long, minutely hairy; basal scales narrow, firm; base of stipe to first large pinna 15-30 cm. Reduced pinnae 5-6 pairs, uppermost 6-12mm long and wide, broadly triangular, lowest 3-5 mm long. Lamina 30-40 cm long; apex deltoid, deeply lobed; pinnae to 15 pairs, basal ones slightly narrowed at base which in some cases is slightly auricled; aero- phores not elongate. Lower surface of rachis covered with short erect pale hairs; upper surface with long curved brown hairs and shorter ones in the groove. Largest pinnae 5-10cm long, 1.5- 2.0 cm wide, in some cases widest 1/3 from apex; base truncate, in some cases a little dilated both sides or only on acroscopic side; apex abruptly short-acuminate, with or without a short cauda; edges crenate throughout (near apex only in small pinnae) to a depth of 1-2 mm; costules commonly 4mm apart, at 60°; veins 5-6 pairs, 2 pairs anas- tomosing to form a zig-zag excurrent vein, 1-2 pairs to rather long sinus-membrane; lower sur- face of costae covered with antrorse (not closely appressed) hairs 0.1-0.3 mm long, hairs on cos- tules and veins sparse, sometimes with a few glands, short erect hairs on surface between veins; hairs on upper surface of costae 0.3-0.4 mm long, a few minute hairs on costules, between veins a variable number of short appressed hairs. Sori medial; indusia with many short stiff hairs; sporangia bearing several setae, sometimes also a gland; spores minutely papillose. Distr. Malesia: New Guinea and Waigeo; at low altitudes in forest. Note. Many collections from eastern New Guinea, none from west except Waigeo. 112. Sphaerostephanos multiauriculatus (COPEL.) HOLTTUM, comb. nov.— Dryopteris multiauri- culata COPEL. Univ. Cal. Publ. Bot. 18 (1942) 221.—Cyclosorus multiauriculatus (COPEL.) COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78 (1951) 453, pl. 32.— Thelypteris multiauriculata (COPEL.) REED, Phytologia 17 (1968) 294. — Type: BRASS 12850, New Guinea, Idenburg River, 1150 m, in gullies in forest (MICH; BM, L). Caudex erect, to 40cm tall, 1.5cm diameter. Stipe 5-10 cm long, reddish, minutely hairy, with many narrow scales 10mm long. Lamina 150 cm long, somewhat dimorphous; pinnae to 45 pairs, lower 12-15 pairs gradually reduced, lowest 1 cm long, suprabasal ones spreading, broadly trian- gular with symmetric base; aerophores elongate barely to 1 mm. Lower surface of rachis of fertile frond densely covered with short appressed hairs, hairs on upper surface similar but with some thick brown hairs 1.5mm long; similar long thick hairs sometimes present on lower surface of sterile fronds. Largest sterile pinnae to 15 x 1.8cm; base FLORA MALESIANA (ser. II, vol. 1° subtruncate; apex short-acuminate, not caudate; edges crenate to a depth of 1mm; costules 3 mm apart, at 60°; veins to 7 pairs, slender and prom- inent both sides, 3-32 pairs anastomosing to form a zig-zag excurrent vein, | pair to very short sinus-membrane; lower surface of costae, costules and veins covered with appressed hairs 0.1- 0.3mm long, a few similar hairs on surface be- tween veins, glands lacking; upper surface of costae covered with hairs less than 0.5 mm long, rest of surface glabrous. Fertile pinnae to 9x 1.0cm; sori medial, often spreading a little along veins; no indusia; sporangia sometimes bearing a few glands, not setae. Distr. Malesia: Eastern New Guinea at 750- 1220 m. 113. Sphaerostephanos stipellatus (BL.) HOLT- TUM, comb. nov.— Aspidium stipellatum BL. Enum. Pl. Jav. (1828) 152; MIQUEL, Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 160.— Nephrodium stipellatum (BL.) Moore, Ind. Fil. (1858) 105; RACcIB. FI. Btzg 1 (1898) 188. — Dryopteris stipel- lata (BL.) O. KTZE, Rev. Gen. Pl. 2 (1891) 813; v.A.v.R. Handb. (1908) 227; BACKER & POSTH. Varenfl. Java (1939) 49.— Thelypteris _ stipel- lata (BL.) K. Iwats. Acta Phytotax. Geobot. 21 (1965) 168.—Type: BLUME, Java (L, n. 908, 337-123). Caudex thick, short-creeping, apex and very young fronds covered with slime. Stipe 10cm long, glabrescent; scales very thin; base of stipe to first large pinna 65cm. Reduced pinnae 2-3cm apart, to 15 pairs or more, lowest 6mm long, uppermost 12 mm long, deflexed, narrowly trian- gular and strongly auricled. Lamina to 70 cm long; pinnae to 30 pairs, lower ones not narrowed at their bases; aerophores thick, to 2 mm long. Hairs on lower surface of rachis of fertile fronds 0.3mm long, closely appressed, on sterile ones sparse longer hairs also; on upper surface pale, spreading, | mm long. Largest pinnae commonly 12x 1.8cm; base subtruncate; apex acuminate, apical 1.5cm entire; edges lobed c. 1/3 to costa (always less than 3), lobes of fertile fronds dis- tinctly falcate; costules 3.5-4.5mm apart, at a wide angle; veins 6-8 pairs, | pair anastomosing, next acroscopic vein to side of short sinus-mem- brane; hairs on lower surface of costae and cos- tules short, closely appressed, with a few longer ones distally on costae, few or none on veins and surface between them, no glands; hairs on upper surface of costae 1mm long, scattered similar hairs on costules and veins. Sori medial; indusia firm, with a few glands; sporangia bearing glands; spores finely spinulose. Distr. Malesia: Java, Sumatra. Ecol. In forest at 1000-1700 m. 114. HOLTTUM, Sphaerostephanos _plurifolius (v.A.v.R.) comb. nov.— Dryopteris plurifolia 1981] THELYPTERIDACEAE (Holttum) 493 v.A.v.R. Bull. Jard. Bot. Btzg III, 5 (1922) 201. — Thelypteris plurifolia (v.A.v.R.) REED, Phy- tologia 17 (1968) 305.—Type: LORZING 6393, Sumatra, Deli, Bandar-baroe, 800 m (BO). Caudex erect, 20-30cm tall, apex and young fronds covered with mucilage; stipe 10-l1Scm long; base of stipe to first normal pinna 40-50 cm; reduced pinnae c. 4cm apart, to 8 pairs, lowest very small, upper ones 6mm long. Lamina 70cm or more long; pinnae to 25 pairs, 3 or more pairs of lower ones distinctly narrowed towards their bases, basal ones much narrowed; aerophores to 2mm long. Hairs on lower surface of rachis very short, erect, on upper surface 0.5 mm long, or to 1 mm on sterile fronds. Largest suprabasal pinnae commonly 15 x2 cm (sterile to 25 x 3.2 cm): base truncate; apex acuminate; edges lobed 2/3-3/4 towards costa, lobes falcate; costules 3.5-S mm apart, at more than 60° to costa; veins 10-13 pairs, basal pair anastomosing, next pair both passing to margin or the acroscopic vein to sinus-mem- brane; lower surface of costae, costules and veins covered with short pale appressed hairs, without glands, area between veins glabrous; hairs on upper surface of costae short, with some longer ones, similar long hairs scattered on costules and veins. Sori medial; indusia large, firm, with a few hairs; sporangia usually bearing glands. Distr. Malesia: Sumatra. Note. The type has distinctly dimorphous fronds, but a specimen from Gunong Kerinci (ROBINSON & KLOSS s.n. at BM) has fertile pinnae to 3.2 cm wide. 115. Sphaerostephanos hendersonii HOLTTUM, sp. nov. Pinnae redactae c. 8-jugatae, usque 6mm lon- gae; pinnae normales usque 14 Xx 2.3 cm, 3/5 cos- tam versus lobatae, basi aerophoris longis praedi- tae, subtus eglandulosae; venae 10-jugatae, infimae solum anastomosantes; rachis, costae costulaeque subtus pilis adpressis vestitae; sori mediales; indusia parva, tenuia, sparsim pilosa; sporangia glandulifera. —Type: HENDERSGN 23352, Malaya, Cameron Highlands, 1500m (K; SING). Caudex short, erect or suberect; stipe 10-15 cm long; base of stipe to first normal pinna 40-50 cm; reduced pinnae c. 4cm apart, c. 8 pairs, uppermost 6mm long, rest very small. Lamina excluding reduced pinnae 70cm long; pinnae more than 20 pairs; basal pinnae narrowed towards their bases which are 1.2 cm wide; aerophores to almost 2mm long. Largest pinnae (fertile) 14x 2.3cm; base truncate; apex abruptly short-acuminate with nar- row entire cauda 1l-2cm long; edges lobed to 4mm from costa; costules 4-5 mm apart, at more than 60°; veins to 10 pairs, basal pair anas- tomosing, next acroscopic vein passing to the sinus-membrane, sometimes a basiscopic vein also; lower surface of rachis bearing slender ap- pressed pale hairs 0.2mm _ long (sometimes sparse), costae and costules bearing many ap- pressed hairs 0.3-0.4mm long, on and between veins neither hairs nor glands; upper surface of rachis bearing pale curved hairs 0.5 mm long, hairs on costae to !mm or more, similar hairs rare on costules and veins, surface between veins bearing a variable number of appressed hairs. Sori medial, basal ones not divergent; indusia small, thin, with a few hairs; sporangia bearing glands. Distr. Malesia: Malaya (Cameron Highlands, 3 collections); N. Sumatra at 1700 m. Note. The Sumatran specimens (SURBECK 747, 1201) are somewhat smaller than those from Malaya, with pinnae lobed a little less deeply, and have a seta on some sporangia. 116. Sphaerostephanos posthumii HOLTTUM, sp. nov. Pinnae redactae c. 10-jugatae, usque 4 mm lon- gae; pinnae normales usque 12.5 1.5 cm, basi aerophoris 1mm longis praeditae, profunde loba- tae; venae infimae prope sinum conniventes, vel liberae vel anastomesantes; pagina inferior pin- narum eglandulosa, pagina superior pilis adpressis vestita; indusia parva, brevi-pilosa; sporangia glandulifera. — Type: POSTHUMUS 3157, Flores, near Sita, 600-700 m (BO). Caudex not known; stipe 24cm long, basal scales broad, thin, with rather long superficial hairs; base of stipe to first normal pinna 80cm; reduced pinnae c. 5cm apart, uppermost 4mm long. Lamina 60cm long, excluding reduced pin- nae; pinnae 20 pairs; lowest large pinnae much narrowed towards their bases, several successive pairs gradually less narrowed; aerophores 1mm long. Rachis on both surfaces covered with pale appressed hairs c. 0.6mm long. Largest pinnae (fertile) 12.5x1.5cm, sessile; apex gradually attenuate, not caudate; edges lobed to 1.5-2mm from costa, lobes slightly falcate; costules 3- 35mm apart, at 60°; veins 10-12 pairs, basal pair either both touching sides of a short sinus-mem- brane or meeting just below it to form a short excurrent vein; lower surface of costae and cos- tules closely covered with pale appressed hairs, between veins some shorter appressed hairs, glands lacking; upper surface throughout covered with pale appressed hairs 0.3-0.4 mm long. Sori a little inframedial, basal ones not much divergent; indusia rather small, thin, with many hairs 0.2 mm long; sporangia often with a gland. Distr. Malesia: Lesser Sunda Is. (Flores), only known from type. 117. Sphaerostephanos inconspicuus (COPEL.) HOLTTUM, comb. nov. — Dryopteris inconspicua CoPEL. Philip. J. Sci. 12 (1917) Bot. 55; C. CHR. Gard. Bull. Str. Settl. 7 (1934) 242. — Thelypteris inconspicua (COPEL.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 252. — Type: TOPPING 494 1543, Sabah, Mt Kinabalu, Kiau (isotype NY). — Fig. 12n-p. Caudex short, creeping. Stipe 4-8cm long, minutely hairy; base of stipe to first large pinna 15-30cm. Reduced pinnae 4-7 pairs, all very small, not opposite. Lamina 25-40 cm long; pin- nae to 25 pairs, basal pinnae narrowed at base; aerophores not elongate. Hairs on lower surface of rachis variable, sometimes very few on fertile fronds, on sterile often many, thick, curved, 0.5 mm or more long; hairs on upper surface 0.8- 1.0mm long. Largest pinnae 3.5x0.8 to 6.0 1.3cm; base subtruncate; basal acroscopic lobe sometimes elongate; apex acuminate, sometimes caudate; edges lobed to 1-1.5mm from costa, lobes oblique; costules 3-3.5 mm apart, usually at less than 60°; veins 4-5 pairs, slender, basal acroscopic vein touching short sinus-membrane, basal basiscopic vein to edge above base of sinus; hairs on lower surface of costae 0.2-0.3 mm long, pale, appressed, fewer on costules, sometimes with rudimentary scales which have a glandular tip, sessile spherical glands usually lacking; hairs on upper surface of costae less than 0.5 mm long, no others. Sori medial; indusia rather large with a few short hairs or glabrous; sporangia bearing a gland or a seta; spores not seen. Distr. Malesia: Borneo; several localities, in forest at 1200-1800 m. 118. Sphaerostephanos subulifolius (v.A.v.R.) HOLTTUM, comb. nov.— Dryopteris subulifolia v.A.v.R. Bull. Jard. Bot. Btzg II, 28 (1918) 22.— Thelypteris subulifolia CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 254.— Type: BROOKS 334 partim, Sumatra, Benkoelen, Lebong Simpang (BO; BM). Closely allied to S. inconspicuus, differing as follows: reduced pinnae 2-3 pairs, all very small; normal pinnae to 30 or more pairs; pinnae ab- ruptly short-acuminate, lobed to 1 mm from costa; hairs on lower surface of costae 0.5 mm long, not closely appressed. Distr. Malesia: Southern Sumatra, 3 collec- tions. Notes. The differences between this and S. inconspicuus need checking from further collec- tions from Sumatra. BROOKS 334 at BM includes specimens of Coryphopteris hirsutipes (CLARKE) HOLTTUM. 119. Sphaerostephanos mjobergii HOLTTUM, sp. nov. S. inconspicuo (Copel.) Holttum affinis, differt: pinnis evolutis 27-jugatis, usque 3.6*1.0cm metientibus, ad alam 0.5 mm latam lobatis; venis infimis omnibus ad marginem supra basin sinuum terminatis; costis subtus pilis patentibus vestitis. — Type: E. MJOBERG s.n. Sarawak, Mt Poi, 1370-1520 m (BM). Caudex and stipe missing. Reduced pinnae all FLORA MALESIANA [ser. II, vol. 1° very small, 2-3 pairs on specimen. Lamina 30cm long; pinnae 27 pairs; basal pinnae apparently narrowed on basiscopic side only; aerophores not elongate. Hairs on lower surface of rachis 0.2- 0.3 mm long, copious. Largest pinnae 3.6 X 1.0 cm; base symmetric, subtruncate; apex rather abruptly acute; edges lobed to 0.5mm from costa, lobes hardly falcate; costules to 2.5 mm apart; veins 3-4 pairs, basal ones both passing to margin above base of sinus: lower surface of costae covered with somewhat antrorse hairs 0.2-0.3mm long, longer hairs scattered on costules; hairs on upper surface of costae short, with longer ones scattered on distal parts of costae and on costules. Sori inframedial; indusia sparsely hairy; sporangia bearing glands; spores with a narrow erose wing, surface otherwise bearing scattered small wings or papillae. Distr. Malesia: Borneo (Sarawak), only known from the type. 120. Sphaerostephanos mutabilis (BRAUSE) HOLTTUM, comb. nov.— Dryopteris mutabilis BRAUSE, Bot. Jahrb. 56 (1920) 97. — Cyclosorus mutabilis (BRAUSE) COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78 (1951) 443. — Thelypteris mutabilis (BRAUSE) REED, Phytologia 17 (1968) 295.— Type: LEDERMANN 9745, N.E. New Guinea, Sepik Distr., on rocks, 200-400 m (B; UC). Caudex short, erect. Stipe 5-8 cm long, reddish, glabrous; scales c. 2% 1mm. Lamina to 20cm long; apex pinna-like; pinnae 10 pairs, lower 3-4 pairs gradually smaller, lowest S-10mm_ long; aerophores not elongate. Largest pinnae com- monly 60.3cm (to 8.5x0.5cm, fide BRAUSE); base and apex narrowly attenuate, edges entire; costules 1.5mm apart, very oblique, forked or simple, without anastomosis; lower surface quite glabrous; upper surface hairy in groove of rachis only. Sori in one row on each side of costa, often on both branches of a costule; indusia large, firm, with a few glands; sporangia sometimes bearing glands; spores minutely spinulose. Distr. Malesia: New Guinea. Known only from type and BRASS 13047 from Idenburg River. Ecol. In forest, “massed in semi-shade on steep flood-swept bank of river’ (BRASS), 200- 850 m. 121. Sphaerostephanos uaniensis HOLTTUM, Sp. nov. Stipes 5cm longus; pinnae redactae 3-4-juga- tae: lamina 33cm longa, pinnae normales 12- jugatae, superiores plures non liberae; pinnae liberae subintegrae, basi auriculatae; costae sub- tus pilis erectis 0.1-0.2 mm longis vestita, pagina inferior cetera pilis brevioribus praedita, eglan- dulosa; sori elongati; indusia parva (vel nulla?); sporangia interdum glandula una_praedita. — Type: T. G. WALKER 10132, cult. Kew, origin New Britain, Uani River (BM). 1981] THELYPTERIDACEAE (Holttum) 495 Stipe 5cm long; lamina in all 33cm long, con- from the type; ‘““common” at the type locality. sisting of 3 pairs of reduced pinnae (basal one 7x5mm with stalk 1.5mm, uppermost 1.2 123. Sphaerostephanos peltochlamys (C. CHR.) 0.6cm with auricled base), 1 intermediate pair (unequal) 2.0 and 2.7 cm long, and 12 pairs normal pinnae of which several upper ones are adnate to the rachis. Largest pinnae 4.2cm long, 1.4cm wide at the auricled base, 1.1 cm wide above base; apex rather abruptly obtuse; edges irregularly sinuous; costules 2.5—3 mm apart, at c. 45°; veins 3 pairs, 2 pairs anastomosing, slender, concolorous; lower surface of rachis bearing sparse stiff curved pale hairs 0.2-0.3mm long, costal hairs 0.1- 0.2 mm, erect, very short hairs on costules and on surface between veins, no glands; upper surface hairy only on costae and rachis. Sori all con- siderably elongate, those on basal veins of ad- jacent costules almost meeting, distal ones shorter and near costules; indusia very small with 2 or 3 short hairs, apparently sometimes _ lacking; sporangia not setiferous, sometimes with a gland. Distr. Malesia: New Guinea (New Britain), only known from type. 122. Sphaerostephanos_ echinosporus (v.A.v.R.) HOLTTUM, comb. nov. — Dryopteris echinospora v.A.v.R. Bull. Jard. Bot. Btzg III, 2 (1920) 149. — Thelypteris echinospora (v.A.v.R.) REED, Phytologia 17 (1968) 274.—Type: BROOKS 451, Sumatra, Seblat River, Benkoelen (BO; BM). Caudex not known; stipe 5cm long; base of stipe to largest pinna 80cm; reduced pinnae c. 15 pairs, uppermost (v.A.v.R.) 61cm, a middle one seen 4.5 cm long, 7 mm wide in the middle and gradually narrowed towards auricled base, lobed half-way to costa, one near base 1.8cm long, 2mm wide apart from a narrow basal auricle. Lamina excluding reduced pinnae 100cm long; basal pinnae narrowed towards their bases which have a narrow auricle 6mm long, upper pinnae truncate to full width at base; aerophores to 2mm long. Largest pinnae 16x 2.1cm; apex narrowly caudate and entire; edges lobed to 2mm from costa; lobes slightly falcate, separated by wide sinuses; costules 3.5-4mm apart; veins to 11 pairs, basal pair, at a wide angle to costule, anas- tomosing to form a short excurrent vein, next pair both to margin above base of sinus; lower surface of costae densely covered with hairs 0.2-0.3 mm long, hairs on costules more sparse and somewhat antrorse, sparse minute erect hairs present be- tween veins, no glands; upper surface of costae covered with antrorse hairs 0.6mm long, rest of surface covered with slender appressed hairs 0.3mm long. Sori medial, covering lower surface of pinna-lobes at maturity; indusia large, firm, with sparse short hairs; a small proportion of sporangia bearing a single seta or (less com- monly?) a gland. Distr. Malesia: South Sumatra. Known only HOLTTUM, comb. nov.— Dryopteris peltoch- lamys C. CHR. Dansk Bot. Ark. 9, 3 (1937) 65; BACKER & POSTH. Varenfl. Java (1939) 338, excl. syn.— Abacopteris peltochlamys (C. CHR.) HOLTTUM, Rev. Fl. Mal. 2 (1955) 295, f. 171. — Thelypteris peltochlamys (C. CHR.) REED, Phy- tologia 17 (1968) 303.— Type: C. W. FRANCK s.n. E. Java, Tanggore (BM). Nephrodium urophyllum sensu RAciB. FI. Btzg 1 (1898) 184. Caudex a short or long-creeping rhizome. Stipe 20-30cm long, glabrous, stramineous; scales broad, thin; base of stipe to first large pinna 30-60cm (longest on fertile fronds). Reduced pinnae 2-4 pairs (lacking on young plants), less than 5mm long. Lamina to 60cm long; apex pinna-like or with one basal lobe; pinnae 6-10 pairs, lower ones sometimes a little reduced, al- ways with a narrowed base; aerophores not swollen. Lower surface of rachis glabrous; upper surface with short hairs in groove. Largest pinnae of type 21 x 3.5cm; sterile pinnae to 4.5 cm wide, fertile ones on small plants 14 x 2.8 cm; suprabasal pinnae widest at or above the middle, slightly and gradually narrowed to an abruptly contracted broadly cuneate base; apex abruptly short cau- date-acuminate; edges entire to crenate; costules 4mm apart, at 60°; veins 7-10 pairs, almost all anastomosing to form a zig-zag excurrent vein: sinus-membrane only distinct where margin is crenate; lower surface quite glabrous; between veins + verrucose when dry; upper surface of costae with slender hairs 0.3 mm long. Sori medial or somewhat inframedial; indusia large, thin, glabrous, rarely with hairs or a few glands; sporangia often bearing glands; spores with a continuous translucent wing and several other separate small wings. Distr. Burma, in Malesia: Malaya (one collec- tion in Kelantan), Sumatra and Java (many locali- ties), in lowland forest. Note. Apart from the reduced pinnae, this is very similar to Pronephrium lineatum (BL.) PRESL. It seems uniform over a wide area, and may be a tetraploid of hybrid origin. 124. Sphaerostephanos irayensis (COPEL.) HOLT- TUM, comb. nov. — Cyclosorus irayensis COPEL. Philip. J. Sci. 81 (1952) 28; Fern FI. Philip. (1960) 340.— Thelypteris irayensis (COPEL.) REED, Phytologia 17 (1968) 285. — Plesioneuron irayense (COPEL.) HOLTTUM, Blumea 22 (1975) 239. — Type: RAMOS BS 80344, Batanes Islands, Mt Iraya (MICH; SING). Sphaerostephanos fenixii HOLTTUM, Kalikasan 4 (1975) 55.— Type: FENIX BS 3780, Batanes Islands (P; US). Caudex short-creeping; stipe 35-70cm long, 496 glabrescent, reddish. Lamina 35-40 cm long; pin- nae to 15 pairs, of firm texture; 1 pair reduced basal pinnae 1.5cm long sometimes present; lowest pinnae variably narrowed towards base with stalk 1 mm long, to 2.5 cm wide at the middle; aerophores not elongate. Lower surface of rachis densely short-hairy. Suprabasal fertile pinnae to 12 x 2.2 cm (sterile to 2.8cm wide); base broadly cuneate to full width; apex acuminate; edges lobed more than 1/2 way to costa, in basal sterile pinnae to more than 2/3; lobes falcate, those of basal pinnae attenuate; costules 5 mm apart, at 60° or more; veins 10-12 pairs, pale and prominent on both surfaces, basal pair always connivent below sinus, either anastomosing or touching sides of sinus-membrane without fusing; lower surface of costae hairy as rachis but less densely, of costules and veins sparsely, hairs longer on sterile than on fertile pinnae; upper surface densely short-hairy on costae, sparsely on costules. Sori medial, im- pressed on upper surface when dry, lower ones distinctly elongate; indusia small, thin, with short hairs; sporangia bearing 2-3 spherical yellow glands; spores with a narrow erose translucent wing and a few cross-wings. Distr. Malesia: Philippines (Batanes Islands, 3 collections). Note. This very interesting and isolated spe- cies needs further study. The type of C. irayensis resembles species of Plesioneuron in aspect and texture, but its glands (on sporangia), and venation and sinus-characters are unlike that genus. 125. Sphaerostephanos lithophyllus = (COPEL.) HOLTTUM, comb. nov.— Dryopteris lithophylla CoPEL. Philip. J. Sci. 12 (1917) Bot. 57; C. CHR. Gard. Bull. Str. Settl. 7 (1934) 206, 245. — Thelyp- teris lithophylla (COPEL.) REED, Phytologia 17 (1968) 289.—Type: TOPPING 1850, Sabah, Mt Kinabalu (AFS; US). Caudex short, suberect; apex covered with slime. Stipe 1-2cm long, dark; base of stipe to first large pinna 20-35cm. Reduced pinnae near base 1 cm apart, distally 2-3 cm, in all to at least 10 pairs, all with rigid aerophore, lower ones without a perceptible blade, upper ones 2 mm long and wide. Lamina to 30cm long, red when young; pinnae 12-18 pairs, fleshy when living, thick and rigid when dried, 2-3 basal pairs slightly narrowed at base; aerophores elongate. Rachis flushed with red; both surfaces bearing a varied number of short pale to dark red erect hairs; upper surface deeply grooved. Largest pinnae 5-6.5 x 1.1cm; base truncate; apex short-acuminate; edges lobed about 1/3 to costa, lobes rounded, their margins revolute on drying; costules to 3mm apart; each with a small swelling at the base; veins to 8 pairs, prominent on the upper surface and not below, basal pair anastomosing, 2 pairs to sinus-mem- brane which is very prominent on the lower sur- face; lower surface of costae almost glabrous or FLORA MALESIANA (ser. II, vol. 1° with pale short spreading hairs, some short hairs present on surface between veins; sparse hairs present on edges of groove of costa on upper surface, no others. Sori supramedial; indusia with a few short hairs and glands; sporangia bearing glands, often several; spores copiously minutely spinulose. Distr. Malesia: Sabah (Mt Kinabalu). Ecol. In wet ground in fully exposed places, often among rocks, at 1500-2000 m. 126. Sphaerostephanos hamiferus (v.A.v.R.) HOLTTUM, comb. nov.— Dryopteris hamifera v.A.v.R. Bull. Jard. Bot. Btzg II, 16 (1914) 12; Handb. Suppl. (1917) 181.— Thelypteris hamifera (v.A.v.R.) REED, Phytologia 17 (1968) 281. — Type: MATTHEW 5.n. Sumatra, G. Singgalang (BO). Caudex erect. Stipe 12cm long; basal scales thin, narrow; base of stipe to first large pinna 50cm. Reduced pinnae at least 10 pairs, lowest 5 mm long, upper ones 1.0-1.5 cm long, spreading, narrowly triangular with auricle 5mm long on acroscopic side, short on basiscopic; on type 1 pair intermediate pinnae also present. Lamina 70 cm long; pinnae 30 pairs; lowest pinnae slightly narrowed and slightly auricled at base; aerophores 1-2 mm long (at least on reduced pinnae). Hairs on lower surface of rachis dense, erect, to 0.5mm long on fertile frond, mixed with many hairs 1 mm long on sterile; hairs on upper surface 0.5-1 mm long, less difference between fertile and sterile. Largest fertile pinnae of type 10x 1.5cm; base truncate; apex acuminate, not caudate; edges lobed more than 3 way to costa (on a larger specimen to 2/3), lobes slightly falcate; costules 3.5-4 mm apart, at more than 60°; veins 7-8(-10) pairs, basal pair anastomosing, next pair both to margin or acroscopic one to the short sinus- membrane; lower surface of costae hairy as rachis but. hairs a little shorter, hairs on costules sparse, on fertile fronds antrorse but not closely ap- pressed, glands absent or rare; hairs on upper surface of costae to 1 mm long, scattered similar hairs on costules and veins, surface between veins + closely covered with slender appressed hairs. Sori medial; indusia large, firm, with short hairs; sporangia sometimes with a gland, not setiferous. Distr. Malesia: Central Sumatra, at 1500 m or less. Note. There is variability in size of fertile pinnae from 6.4 0.9cm to 12 x 2.0cm, the latter lobed fully 2/3. 127. Sphaerostephanos HOLTTUM, sp. nov. Caudex brevis, repens; pinnae redactae 20-30- jugatae, aerophoris 3 mm longis constitutis; pin- nae 10-jugatae, fertiles usque 305.5 cm; venae 18-20-jugatae; pagina inferior pinnarum glabra; sori exindusiati; sporangia nec setis nec glandulis praedita.—Type: JERMY 4038, N.E. New plurivenosus 1981] THELYPTERIDACEAE (Holttum) 497 Guinea, Madang Distr., gorge below Moro (BM). Caudex massive, prostrate. Stipe c. 5cm long; base of stipe to first true pinna 30-50 cm. Reduced pinnae consisting of aerophores 3mm long with no evident blade, 10-15 mm apart, 20-30 pairs. Lamina 100cm or more long; apex pinna-like; pinnae c. 10 pairs, opposite, basal ones not or little reduced, lower ones 10cm apart. Rachis reddish, glossy, quite glabrous apart from small hairs in groove of upper surface. Largest fertile pinna 30 x 3.5 cm (sterile to 35 X 6.5 cm) with stalk 2mm long and stiff aerophores | mm; base rounded to broadly cuneate; apex acuminate with slender cauda to 3cm long; edges parallel for most of their length, entire or slightly crenate; costules 445mm apart, at almost 90°, falcate distally; veins 18-20 pairs, close, at a wide angle to cos- tules (more oblique in sterile pinnae), excurrent veins mostly free, 2-3 veins from each costule passing to margin; both surfaces quite glabrous, eglandular. Sori supramedial, those on veins from adjacent costules touching or fusing, exindusiate, lower ones elongate; sporangia bearing neither glands nor setae; spores with a continuous wing of irregular width and some _ smaller’ wings. Chromosomes: n= 36 (T. G. WALKER). Distr. Malesia: Papua New Guinea; Madang and Morobe Districts at 1300-1700 m; additional specimens are CLEMENS 4841, Sattelberg; HOOGLAND 9094, Mt Rawlinson. Ecol. HOOGLAND’s specimen was from a shaded vertical limestone cliff near a creek. 128. Sphaerostephanos squamatellus HOLTTUM, sp. nov. Pinnae redactae 8-jugatae, superiores 3mm longae; pinnae normales 2/5 costam versus loba- tae; aerophora elongata; costae costulaeque sub- tus pilis appressis vestitae, eglandulosae; pagina superior pinnarum pilis appressis vestita; indusia pilosa; sporangia setifera. —Type: HOLTTUM 56, Sabah, Mt Kinabalu, near waterfall in forest, 1900 m (K). Caudex not preserved. Stipe 12cm _ long, minutely hairy with many small scales; base of stipe to first large pinna 35 cm. Reduced pinnae 8 pairs, uppermost 3mm long, with elongate aero- phores. Lamina 40cm long; pinnae 16 pairs, all almost opposite; basal pair of pinnae somewhat shortened and a little narrowed at base which is 0.8cm wide. Rachis stramineous; lower surface covered at base with very short erect hairs and small appressed scales, distally with longer hairs and few scales; upper surface bearing brown hairs 1mm long. Largest pinnae 10.5 1.7cm; base broadly cuneate; apex rather abruptly short-acu- minate; edges lobed about 2/5 to costa, lobes slightly falcate; costules 44.5 mm apart, at 60° or more; veins 7 pairs, basal pair anastomosing, 1-13 pairs passing to sinus-membrane which is not prominent on the lower surface; lower surface of costae covered with slender appressed hairs 0.3mm long, costules and veins the same with minute uniseriate scales, appressed hairs 0.3 mm long scattered on surface between veins; hairs on upper surface of costae 0.7 mm long, rest of upper surface bearing slender appressed hairs 0.3 mm long. Sori medial; indusia firm with a few short hairs; sporangia all with 2-4 short setae. Distr. Malesia: Sabah (Mt Kinabalu) (also CLEMENS 32548A at 2450 m, BO, L). 129. Sphaerostephanos canescens (BL.) HOLTTUM, comb. nov. — Polypodium canescens BL. Enum. Pl. Jav. (1828) 133.— Gymnogramme canescens (BL.) BL. FI. Jav. Fil. (1829) 93, t. 40. — Phegop- teris canescens (BL.) METT. Ann. Mus. Bot. Lugd.- Bat. 1 (1864) 223 (non MEeETT. 1858); v.A.v.R. Handb. (1908) 507, excl. var. omn.— Dryopteris canescens (BL.) C. CHR. Ind. Fil. (1905) 250, quoad typus tantum; BACKER & POSTH. Varenfl. Java (1939) 56.— Haplodictyum canescens (BL.) CHING, Sunyatsenia 5 (1940) 251.— Type: REINWARDT, Moluccas, Tidore (L, n. 908, 300-133). Caudex short, creeping. Stipe 9-12cm long, densely short-hairy in groove; base of stipe to first large pinna 19-24cm. Reduced pinnae to 5S pairs, 2-3mm long, 3cm apart. Lamina 27cm long consisting of a deeply lobed apical section 18 cm long, 2.7 cm wide at base, with gradual transition to pinnae, and 5 pairs free pinnae, lowest longest, all widest near their apices; aerophores elongate. Rachis bearing hairs 1mm long both sides, less abundant on lower surface with many shorter hairs. Largest pinnae 4—4.5 cm long, 1.6cm wide at 1/3 from apex; base truncate; apex very ab- ruptly short-pointed; edges lobed 1/3 to costa; costules to 3 mm apart at more than 60°; veins 7-8 pairs, | pair anastomosing and 2-2; pairs passing to long sinus-membrane which is prominent on the lower surface; hairs on lower surface of costae pale, spreading, 0.4-0.7mm long, on costules similar, erect veins present on surface between veins; hairs on upper surface of costae more than 0.5mm long, similar hairs scattered on costules and veins, whole surface covered with fine pale appressed hairs. Sori medial, exindusiate; sporangia bearing several setae; spores with many small wings. Distr. Malesia: West Java (Mt Gedeh), N. Celebes, Moluccas (Tidore, Halmahera). Notes. BLUME (1829) stated that the name was given by REINWARDT to the specimen col- lected by REINWARDT in Tidore, which may have been the model for BLUME’s plate; BLUME stated that he had also specimens from G. Gedeh (n. 908, 337-391) and these are certainly like REIN- WARDT'’S. This species was construed in a very broad sense by CHRIST to include specimens which are here referred to several other species, and in part 498 FLORA MALESIANA (ser. II, vol. 1° he was copied by v.A.v.R. The veins on some of the lower lobes of the apical lamina are sometimes forked as in Haplodictyum heterophyllum PRESL, for which reason CHING transferred this species to Haplodictyum. 130. Sphaerostephanos suboppositus HOLTTUM, sp. nov.— Aspidium procurrens sensu CHRIST, Ann. Jard. Bot. Btzg 15 (1898) 130, quoad SARASIN 128.— Aspidium heterocarpus sensu CHRIST, l.c. 133, quoad SARASIN 1326. Pinnae redactae 9-jugatae, superiores 5mm longae; pinnae normales usque 11 cm longae, ad basin auriculatam 2.5 cm latae, suboppositae, 3/5— 2/3 costam versus lobatae, infimae basi leviter angustatae; costae costulaeque subtus pilis antrorsis non appressis vestitae, eglandulosae; pagina superior pilis appressis vestita; sporangia setifera. — Type: ALSTON 15744, Celebes, Mina- hassa, G. Lokon (BM). Caudex short, suberect. Stipe 8cm_ long, glabrescent; scales thin, 5 X 1.5mm; base of stipe to first large pinna 30-45 cm. Reduced pinnae 9 pairs, subopposite, lowest very small, uppermost 5mm long. Lamina 50-70 cm long; pinnae 18-21 pairs, subopposite in basal half of frond, basal 2-3 pairs somewhat narrowed at base; aerophores slender, to 1mm long. Hairs on lower surface of rachis dense, short, pale; on upper surface pale, less than 0.5mm long. Middle pinnae to 11cm long, to 2.5cm wide at auricled base (which is sometimes a little dilated also on basiscopic side), 2.1cm wide above base; apex acuminate; edges lobed 3/5—2/3 to costa, lobes slightly falcate with obtuse tips; costules 4.5mm apart, at more than 60°; veins slender, 9-11 pairs, basal pair anas- tomosing, next acroscopic vein or pair to sinus- membrane; lower surface of costae with dense somewhat antrorse short hairs, on sterile fronds scattered long ones also present, hairs on costules and veins short, antrorse, not appressed, short erect hairs on surface between veins; upper sur- face of costae covered with pale hairs more than 0.5mm long, similar hairs scattered on costules and veins, surface generally covered with pale appressed hairs. Sori medial, lower ones a little divergent; indusia small, thin, with a few hairs; sporangia bearing copious short setae; spores short-spinulose. Distr. Malesia: N. Celebes at 1100-1400 m. 131. Sphaerostephanos makassaricus HOLTTUM, sp. nov. Pinnae redactae c. 10-jugatae, usque 2 mm lon- gae; pinnae normales usque 17 X 2.8 cm, profunde lobatae; aerophori elongati; venae 12-jugatae, infimae tantum anastomosantes; costae costu- laeque subtus pilis 1mm longis brevioribus in- termixtis vestitae, eglandulosae; indusia parva, tenuia, brevi-pilosa; sporangia setifera.— Type: POSTHUMUS 2727, S.W. Celebes, G. Bonthain, S.W. slope, 1500 m, above Karoenglowe (BO). Caudex short, prostrate; stipe 8cm long; base of stipe to first normal pinna 50cm or more; reduced pinnae c. 10 pairs, uppermost 2 mm long, lower ones very small. Lamina excluding reduced pinnae more than 50 cm long; 3 pairs lower pinnae somewhat narrowed at their bases which are not auricled; upper pinnae with almost symmetric bases. Lower surface of rachis bearing copious erect hairs 0.2mm long and scattered pale hairs 1mm long; upper surface with slender pale hairs 0.7mm long. Largest pinnae 17 X2.8cm, almost sessile; aerophores more than Imm long; base broadly cuneate; apex gradually attenuate with cauda to 2cm long; edges lobed to 3mm from costa, lobes falcate and somewhat tapered to a blunt apex; costules 4.5-5.5mm apart, at 60° to costa; veins to 12 pairs, | pair anastomosing, next pair both ending at margin; lower surface of costae bearing copious erect hairs 0.2-0.4 mm long with many to 1 mm or more, on both sterile and fertile pinnae, hairs on costules similar but fewer, few hairs on veins, no hairs and no glands between veins; upper surface of costae with pale hairs 1mm long, shorter hairs scattered on cos- tules and veins, appressed hairs 0.3—-0.4 mm long on surface between veins. Sori medial, lower ones not divergent; indusia small, thin, with many short hairs; sporangia bearing 3-4 slender setae. Distr. Malesia: S.W. Celebes; a second spe- cimen is POSTHUMUS 3476, from Patapang. 132. Sphaerostephanos sarasinorum HOLTTUM, sp. nov. — ASpidium longipes sensu CHRIST, Ann. Jard. Bot. Btzg 15 (1898) 133. Caudex erectus; stipes 7cm longus; pinnae redactae c. 9-jugatae, omnes parvae; lamina 100 cm longa; pinnae inferiores plurimae basin versus valde angustatae; pinnae omnes profunde lobatae, apicem versus abrupte caudatae; rachis costaeque subtus pilis brevibus vestitae; pagina inter venas utrinque glabra; sporangia setifera. — Type: JERMyY 7458, Central Celebes, Masim- bolong River, 1700 m (BM). Caudex short, erect, Stipe 7cm long; basal scales 10 x 1.5 cm, acuminate; base of stipe to first large pinna 50cm or more. Reduced pinnae c. 9 pairs, all very small with swollen aerophores. Rachis short-hairy on both surfaces. Lamina to 100 cm long; pinnae in basal half of frond all much narrowed towards their bases which in lower ones are c. 5mm wide; upper pinnae with basal lobes only reduced. Largest pinnae of type 16 x 3 cm (of SARASIN 996 21%X3.5cm); apex abruptly nar- rowed to a subentire cauda 3 cm long; edges lobed fully 3/4 towards costa, lobes falcate; costules 4.5-6.5mm apart; veins 12-16 pairs, basal ones anastomosing with a short excurrent vein to sinus, next acroscopic vein sometimes to sinus-mem- brane; hairs on lower surface of costae short, 1981] THELYPTERIDACEAE (Holttum) 499 distal ones and those on costules appressed, sur- face between veins glabrous, eglandular; hairs on upper surface of costae | mm long, similar hairs scattered on costules and veins, surface between veins glabrous or sometimes a few hairs present near tips of pinna-lobes. Sori medial, lower ones slightly divergent; indusia sparsely short-hairy; sporangia setiferous. Distr. Malesia: Central Celebes; specimens are SARASIN 993, 996. Ecol. In partial shade on very steep hillside. Note. The SARASIN specimens lack base of fronds, for which reason JERMY’s more perfect specimen is chosen as type. Very young fronds are covered with mucilage (JERMY). additional 133. Sphaerostephanos muluensis HOLTTUM, sp. nov. Pinnae redactae 8-jugatae, omnes parvae; aerophori leviter elongati; pinnae normales usque 15 x 2.5 cm, profunde lobatae; venae 11-12-juga- tae, infimae non anastomosantes; pagina inferior pinnarum eglandulosa; costulae et pagina inter venas subtus pilis adpressis vestita; sori supramediales; indusia pilosa; sporangia inter- dum _setifera.—Type: B. S. PARRIS 6729, Sarawak, Gunong Mulu, 1680 m (CGE). Caudex short-creeping; stipe 15cm long, basal scales 10mm long, narrow; base of stipe to first large pinna 50 cm; reduced pinnae 8 pairs, all very small. Lamina 70 cm long; pinnae 22 pairs, middle ones 3-4cm apart, basal pair much narrowed towards their bases (several successive ones gradually less narrowed); aerophores (when dried) less than 1mm long. Lower surface of rachis rather densely covered with erect stiff brown hairs 0.3-0.4 mm long; upper surface bearing similar but longer hairs, longest distally. Largest pinnae 15 x 2.5 cm, sometimes with a stalk to 1 mm long; apex rather abruptly acuminate with subentire cauda 1.5-2cm long; edges lobed to 1.5-2mm from costa; lobes almost at right angles to costa, slightly narrowed distally, their tips obtuse; cos- tules 5mm apart; veins 11-12 pairs, slender, slightly prominent both sides, basal ones usually both touching sides of the sinus-membrane, less often both ending just above base of sinus; lower surface of costae covered with antrorse but not closely appressed pale hairs 0.6 mm long, hairs on costules a little shorter, closely appressed, slender appressed hairs 0.2-0.3mm long on surface be- tween veins; upper surface bearing hairs about the same length as on lower surface but thicker, no long hairs on costules and veins, surface between veins bearing appressed hairs 0.3 mm long. Sori distinctly supramedial; indusia rather thin, covered with hairs 0.3 mm long; some sporangia bearing a short seta. Distr. Malesia: Borneo (Sarawak), 3 collec- tions. Ecol. In forest at 1680-1800 m. 134. Sphaerostephanos telefominicus HOLTTUM, sp. nov. S. roemeriano (Rosenst.) Holttum affinis, ab ea differt: caudice tenue, erecto vel scandente; lamina longiore, pinnis 20-jugatis constata; pinnis dis- taliter solum crenatis; pagina supra et subtus pilis adpressis vestita. — Type: W. R. BARKER & J. R. CROFT LAE 67649, N.E. New Guinea, W. Sepik Distr., Telefomin Subdistr. (K; LAE). Caudex slender, erect, sometimes creeping up small shrubs (collectors); stipe 5—-12cm long, copiously scaly, scales 5 mm long, less than | mm wide, firm, hairy. Lamina to 25cm long; pinnae more than 20 pairs, isomorphous; basal 2-3 pairs slightly and gradually reduced, smallest 1.2 cm long, subsessile, base unequally cuneate, not auricled; apical lamina not pinna-like. Largest pinnae 3.00.8 cm, with stalk 0.5 mm long; base broadly cuneate to subtruncate, sometimes slightly dilated both sides; apex short-acuminate; edges subentire or slightly crenate distally; cos- tules 2mm apart or a little more; veins to 4 pairs, 2 pairs anastomosing, next acroscopic vein pass- ing to short sinus-membrane; lower surface of rachis and costae bearing copious erect hairs 0.2 mm long (on sterile pinnae 0.3 mm), costules and veins sparsely hairy, surface between veins of sterile pinnae bearing many slender appressed hairs; upper surface of rachis bearing hairs 0.5mm long with scattered ones |_mm, shorter hairs present on costae and a few on costules and veins, between veins slender appressed hairs 0.3 mm long. Sori medial; indusia thin, fairly large, with short hairs; sporangia of type immature, not setiferous, no glands seen on them. Distr. Malesia: Papua New Guinea (Tele- fomin), only known from the type. Ecol. In disturbed montane forest near river, at 1500 m. 135. Sphaerostephanos roemerianus (ROSENST.) HOLTTUM, comb. nov. — Dryopteris roemeriana ROSENST. Nova Guinea 8 (1912) 723; v.A.v.R. Handb. Suppl. (1917) 179. — Thelypteris roemeri- ana (ROSENST.) REED, Phytologia 17 (1968) 310. — Type: VON ROEMER 1023, W. New Guinea, 1350 m (BO). Caudex short, erect. Stipe 3cm long, covered with short pale hairs; scales narrow. Lamina to 12cm long; apex pinna-like; pinnae 10-12 pairs, lower 2-3 pairs gradually smaller, lowest 3-7 mm long. Lower surface of rachis covered with pale spreading hairs 0.3mm long. Largest pinnae to 3.0 x 0.8 cm; base unequally cuneate, sometimes a little auricled on acroscopic side; apex slightly tapered to a rounded tip; edges crenate; costules 2.5mm apart, at 45°; veins to 3 pairs, basal pair anastomosing; lower surface of costae, costules and veins bearing rather sparse short hairs; hairs present on upper surface of costae, scattered on costules and veins. Sori inframedial; indusia large, 500 FLORA MALESIANA [ser. II, vole glabrous; sporangia not setose, no glands seen. Distr. Malesia: W. New Guinea, only known from the type. 136. Sphaerostephanos hellwigensis HOLTTUM, sp. nov. Caudex erectus, usque 30cm altus; pinnae redactae 4—S-jugatae, usque 8-9 mm longae; pin- nae normales usque 4.5 x 0.9 cm, crenatae, eglan- dulosae, costae costulaeque subtus pilis erectis brunneis 0.5-0.6 mm longis vestitae; sori mediales, indusia parva, pilis brevibus praedita; sporangia non setifera, ut videtur non glandulosa. — Type: PULLE 884, W. New Guinea, Mt Hellwig, 2600 m (BO). Caudex erect, to 30cm tall (collector); stipe 5-10 cm long, scales rather firm, 5 x 1 mm; base of stipe to first normal pinna 20cm; reduced pinnae 4-5 pairs, lowest 1-2 mm long, uppermost 8-9 mm long with an acroscopic auricle 4mm long, basis- copic base narrower, edges almost entire. Lamina excluding reduced pinnae 40 cm long; pinnae to c. 30 pairs, texture rigid; basal normal pinnae not narrowed at their bases. Largest pinnae 4.5 x 0.9cm; base truncate and slightly dilated; aero- phores c. 0.5mm long; apex short-acute; edges crenate to a depth of 1 mm or a little more deeply; costules 2.5 mm apart; veins to 4 pairs, prominent both sides, 1 pair anastomosing, next acroscopic vein or pair to the sinus-membrane; lower surface of rachis bearing thick erect brown hairs 1mm long, of costae similar hairs 0.5-0.6mm_ long, fewer on costules, no hairs between veins, no glands; upper surface of rachis covered with antrorse brown hairs, brownish antrorse hairs less than 0.5 mm long on costae, a few short hairs on costules distally. Sori medial; indusia small, thin, with a few short hairs; sporangia not setiferous, no glands seen on them. Distr. Malesia: W. New Guinea (Mt Hellwig), 2600 m. 137. Sphaerostephanos benoitianus (GAUD.) HOLTTUM, comb. nov. — Polystichum benoiti- anum GAUD. in Freyc. Voy. Uran. Phys. Bot. (1827) 331. — Aspidium benoitianum GAUD. ibid. t. 11.— Dryopteris benoitiana (GAUD.) v.A.v.R. Handb. (1908) 225. —Type: GAUDICHAUD, New Guinea, Rawak Island (G; BM, FI-W). Caudex short, creeping. Stipe 7-9cm long, short-hairy; basal scales dark, firm, 4mm long. Lamina 27cm long; apical 10-12cm narrowly triangular and deeply lobed, grading to upper pin- nae; free pinnae 10-12 pairs; basal 1-2 pairs somewhat reduced, narrowed at base on basis- copic side, not on acroscopic, in middle wider on basiscopic than on acroscopic side of costa. Lower surface of rachis bearing short erect hairs and scattered pale ones 1 mm long, hairs on its upper surface thick, brown, spreading. Largest pinnae 5.5 x 1.4cm; base subtruncate, symmetric; apex short-acuminate; edges lobed a little more than 3 way to costa (to 2/3 on fertile fronds), lobes slightly falcate; costules 3mm apart; veins 6-8 pairs, basal pair anastomosing, next acroscopic vein to sinus-membrane or to margin; lower sur- face of costae and costules hairy as rachis; hairs on upper surface of costae sparse, short and long, rest of surface usually glabrous. Sori submarginal; indusium small, bearing short hairs; sporangia with neither glands nor setae; spores dark, minutely papillose. Distr. Malesia: W. New Guinea (Rawak I.), only known from the type. Note. This appears to be an isolated species, possibly related to S. invisus. 138. Sphaerostephanos neotoppingii HOLTTUM, nom. nov. — Dryopteris toppingii COPEL. Philip. J. Sci. 12 (1917) Bot. 56; C. CHR. Gard. Bull. Str. Settl 7 (1934) 246.— Thelypteris toppingii (COPEL.) K. IwaTs. Acta Phytotax. Geobot. 21 (1965) 168, not Sphaerostephanos _ toppingii (COPEL.) C. CHR. 1934. — Type: TOPPING 1766, Sabah, Mt Kinabalu, Lobang (AFS, not seen). Caudex short, creeping. Stipe 50-90 cm long, dull reddish, minutely hairy; scales narrow, to 10mm long. Reduced pinnae lacking. Lamina 50- 80cm long; pinnae 10-15 pairs, lowest not or little reduced; several pairs lower pinnae much nar- rowed towards their bases and stalked 34mm, distal ones broadly cuneate to subtruncate, almost sessile; aerophores not elongate. Rachis densely short-hairy (0.2-0.3 mm) on both surfaces. Largest pinnae commonly 15-20 x 2.0-2.5cm (maximum 24x 3.5cm); base cuneate; apex acuminate with subentire cauda 1.5—2.5 cm; edges lobed 1/3-2/5 to costa, lobes distinctly falcate with narrow blunt forward-pointing tips; costules 5-6.5 mm apart at more than 60°; veins to 12 pairs on acroscopic side of costule, to 14 on basiscopic side, prominent on lower surface, 13-23 pairs anastomosing, 1-2 pairs to sides of sinus-membrane; hairs on lower sur- face of costae antrorse but not closely appressed, 0.2-0.3 mm long with some 0.5mm, similar and more sparse hairs on costules and veins, on sur- face between veins a variable number of short suberect hairs, glands lacking; hairs on upper sur- face of costae 0.3mm long, shorter on costules, between veins (especially on young fronds) some short appressed hairs. Sori supramedial, those on lowest veins from adjacent costules sometimes touching; indusia firm, glabrous, sometimes a little elongate and asymmetric at base; sporangia lack- ing glands and setae, hair on sporangia-stalk end- ing in a red gland. Distr. Malesia: Borneo (Sabah and Kaliman- tan), in forest at 1200-1500 m. 139. Sphaerostephanos oosorus (BAK.) HOLTTUM, comb. nov.— Nephrodium oosorum BAK. Kew Bull. (1896) 41.— Dryopteris oosora (BAK.) C. 1981] THELYPTERIDACEAE (Holttum) 501 Cur. Ind. Fil. (1905) 280; Gard. Bull. Str. Settl. 7 (1934) 245. — Type: G. F. HOSE 334, Sabah, Pulo Gaya (K). Caudex short, creeping. Stipe 25-40cm long, minutely hairy; base of stipe to first large pinna 40-55 cm. Reduced pinnae 2-4(—6) pairs, upper- most 4mm long. Lamina 60 cm long; pinnae to 20 pairs, sessile; 2-3 pairs lower pinnae narrowed at their bases; aerophores not elongate. Lower sur- face of rachis covered with short erect pale hairs 0.2-0.4mm long, in some specimens mixed with long ones to 1mm; upper surface with uniform pale hairs 0.5 mm long. Largest pinnae commonly 12-15 x 1.8-2.0 cm (largest seen 21 x 2.4 cm); apex acuminate with cauda to 1.5cm; edges lobed a little less than 2 way to costa, lobes oblique, nar- rowed to obtuse tips; costules 44.5 mm apart, at 60°; veins to 8 pairs, basal 12 pairs anastomosing, 1-1 pairs to sinus-membrane; lower surface of costae covered with erect hairs varying from 0.2 to 0.5 mm, hairs on costules and veins similar but shorter, surface between veins bearing many short erect hairs; hairs on upper surface of costae 0.5mm long, shorter on costules, abundant short suberect hairs between veins. Sori medial, elon- gate, 1.5 mm long; indusia large, pale, firm, bearing many short hairs and rarely a gland; neither glands nor setae on sporangia; spores minutely papillose. Distr. Malesia: Borneo (Sabah & P. Balam- bangan). Ecol. In forest at low altitudes. 140. Sphaerostephanos rudis (RIDL.) HOLTTUM, comb. nov.—Goniopteris rudis RiDL. Trans. Linn. Soc. Bot. 9 (1916) 259.— Phegopteris ridleyana v.A.v.R. Handb. Suppl. (1917) 515, nom. nov.— Dryopteris rufopilosa BRAUSE, Bot. Jahrb. 56 (1920) 106, nom. nov. (not D. rudis (KUNZE) C. CHR. nor D. ridleyana BRAUSE). — Type: KLoss s.n. W. New Guinea, Mt Carstensz, Camp I-III, 150-750 m (BM; K). KEY TO THE VARIETIES 1. Lower surface of rachis and costae bearing thick reddish hairs : . a. var. rudis 1. Lower surface of rachis and costae glabrous b. var. micans a. var. rudis Caudex short-creeping; stipe 10cm long, dark reddish, dark-hairy in groove, scales 7 x 1mm, thin; base of stipe to first normal pinna 35cm; reduced pinnae 5-6 pairs, all very small, with short aerophores. Lamina 50cm long; apex pinna-like; pinnae 4 pairs, opposite, all widest at or above middle and gradually narrowed to an abruptly cuneate base. Lower surface of rachis covered with thick spreading dark red hairs 1 mm long. Largest pinnae 20x4.5cm; apex abruptly acu- minate with cauda 2-2.5cm long; edges entire; costules 5mm apart; veins 8 pairs, most of them united to form a zig-zag excurrent vein; lower surface of costae bearing thick erect reddish hairs 0.5 mm long which are more sparse distally, hairs on costules short and sparse, none on surface between veins; upper surface glabrous apart from a few hairs in groove of costa. Sori medial, exin- dusiate; sporangia bearing neither glands nor setae. Distr. Malesia: W. New Guinea and Japen Island, at low altitudes. b. var. micans (BRAUSE) HOLTTUM, Stat. nov. — Dryopteris micans BRAUSE, Bot. Jahrb. 56 (1920) 98.—Cyclosorus micans (BRAUSE) COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78 (1951) 458.— Thelypteris micans (BRAUSE) REED, Phytologia 17 (1968) 293.—Type: LEDERMANN 12468, N.E. New Guinea, Sepik Distr. 1400-1500 m (B). Differs from var. rudis in quite glabrous lower surface of rachis and costae; all specimens also are larger, with up to 10 pairs of reduced pinnae and 6 pairs of normal ones which are not opposite, largest 27 x 5.5 cm. Distr. Malesia: New Guinea, from Idenburg River eastwards, at 850-1750. 141. Sphaerostephanos dimorphus (BRAUSE) HOLTTUM, comb. nov.— Dryopteris dimorpha BRAUSE, Bot. Jahrb. 56 (1920) 100. — Cyclosorus dimorphus (BRAUSE) COPEL. Philip. J. Sci. 78 (1951) 442.— Thelypteris dimorpha (BRAUSE) REED, Phytologia 17 (1968) 273.— Type: LEDERMANN 12622, N.E. New Guinea, Sepik Distr., 1400-1500 m (B). Dryopteris morobensis COPEL. Univ. Cal. Publ. Bot. 18 (1942) 221.—Cyclosorus morobensis (COPEL.) COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78 (1951) 459, pl. 38. — Thelypteris moroben- sis (COPEL.) REED, Phytologia 17 (1968) 294. — Type: CLEMENS s.n. N.E. New Guinea, Morobe Distr., Finugan, 1400 m (MICH). Caudex erect, 100-150cm tall. Stipe 10-20cm long, dull red, glabrous, with many scales to 15 x 2mm. Lamina to 140cm long; apex pinna-like; pinnae to 40 pairs, dimorphous, lower 10-12 pairs gradually reduced, lower ones narrowly triangular with + hastate base, lowest 7-15 mm long; aero- phores swollen, not or little elongate. Rachis glabrous apart from dark red hairs in groove of upper surface, most abundant distally. Largest sterile pinnae commonly 20 2.0cm (on type to 36 x 1.8cm); base asymmetric, rounded on basis- copic side, cuneate on acroscopic; apex narrowly acuminate; edges slightly undulate; costules 4— 5mm apart, at a wide angle to costa; veins 4-6 pairs, all very oblique, pale and prominent on lower surface, 2-23 pairs anastomosing and 1 pair to sides of sinus-membrane; both surfaces of pinnae quite glabrous. Fertile pinnae 7-14 cm long, 502 0.7-1.2 cm wide; edges entire as sterile, or crenate (rather strongly crenate in type of D. morobensis); veins 4 pairs; sori confluent at maturity; indusia glabrous or with a few short hairs; sporangia lacking glands and setae; spores minutely papil- lose. Distr. Malesia: Eastern New Guinea, in forest at 1200-2000 m, many collections. 142. Sphaerostephanos potamios HOLTTUM, sp. nov. Caudex erectus, c. 20cm altus; frondes dimor- phae, omnes pinnis redactis 6-8-jugatis praeditae; pinnae Steriles 15*1.6cm, fertiles usque 8.0 0.8 cm, non ultra 1/4 costam versus lobatae, subtus eglandulosae; rachis pilis brunneis vestita; sori mediales; indusia parva, glabra vel pauci-pilosa; sporangia nec setis nec glandulis praedita. — Type: J. CROFT & Y. LELEAN NGF 34913, S.E. New Guinea, Port Moresby Subdistr. near Lake Myola (I): Caudex erect, 20cm tall; stipe 8 cm long, basal scales 7-8 X 1 mm, covered with many short hairs: base of stipe to first large pinna of sterile frond 25cm, of fertile 35-42 cm; reduced pinnae of fer- tile frond 6-8 pairs, not opposite, uppermost 1.5cm long with basal acroscopic auricle 8mm long, apex acute, lowest reduced pinna 3-4 mm long. Lamina 63 cm long (sterile); pinnae 30 pairs, basal pinnae with slightly dilated truncate bases; texture firm; apex of frond not pinna-like; aero- phores slightly swollen. Sterile pinnae to 15x 1.6 cm; apex narrowly acuminate; edges lobed to a depth of 1-2.5mm, lobes with falcate tips and slight projections at ends of veins; costules 3.5- 4.5 mm apart; veins 6-7 on basiscopic side, 5-6 on acroscopic, pale and prominent on lower surface, reddish on upper, basal 1-15 pairs anastomosing, 13 pairs passing to the rather long sinus-mem- brane; lower surface of rachis bearing rather sparse thick brown hairs 0.7 mm long, hairs on costae sparse, pale, erect, 0.3-0.4mm long, hairs on costules shorter, + antrorse, not appressed, surface between veins hairless, slightly pustular; upper surface of rachis with brown hairs to 1 mm long, of costae with pale hairs 0.50.7 mm, no other hairs. Fertile pinnae to 8.0x0.8cm (ap- parently not quite fully expanded); veins 3-4 pairs; hairs on both surfaces shorter and more sparse than on sterile pinnae; sori medial; indusia small, thin, glabrous or with a few short hairs; sporangia not setiferous, no glands seen on them. Distr. Malesia: S.E. New Guinea; only known from the type. Ecol. In forest close to stream-bank, at 1900 m. 143. Sphaerostephanos invisus (FORST. f.) HOLT- TUM, Webbia 30 (1976) 195; Allertonia 1 (1977) 211, f. 7B. — Polypodium invisum ForstT. f. FI. Ins. Austr. Prodr. (1786) 81.— Nephrodium in- visum (ForRST. f.) CARR. in Seem. FI. Vit. (1873) FLORA MALESIANA [ser. II, vol. 1° 362. — Dryopteris invisa (FORST. f.)O. KTZE, Rev. Gen. Pl. 2 (1891) 813; v.A.v.R. Handb. (1908) 212; COPEL. Bishop Mus. Bull. 59 (1929) 45; 93 (1932) 38. — Cyclosorus invisus (FORST. f.) COPEL. Gen. Fil. (1947) 142; BROWNLIE in Aubrév. FI. Nouv. Cal. 3 (1969) 215, excl. syn. Nephrodium haen- keanum PRESL.— Thelypteris forsteri MORTON, Contr. U.S. Nat. Herb. 38 (1967) 60, nom. nov. (not T. invisa (DESV.) PROCTOR). — Type: FORSTER, locality unrecorded (BM; K, LE). Aspidium dissectum sensu METT. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 232, excl. synon. Caudex a long-creeping rhizome to 7mm diameter. Stipe 25-45cm or more long. Lamina commonly 40-75 cm long (to 120 cm); basal 2 pairs of pinnae gradually smaller and irregularly spaced, lowest sometimes 2cm long, usually longer; aerophores not elongate. Largest pinnae com- monly 15 x 1.3cm (to 24x 1.8cm), narrowly acu- minate, lobed 1/3-2/5 to costa, lobes falcate, dis- tinctly pointed, oblique; costules commonly 3- 3.5mm apart; veins 7-8 pairs, close, very oblique except basal pair, 1-1) pairs anastomosing, 1-2 pairs to sinus-membrane; short spreading hairs on all part of lower surface, a little antrorse on cos- tules, erect (sometimes lacking) between veins; upper surface almost glabrous apart from costae. Sori supramedial; indusia thin, hairy; sporangia bearing a variable number of setae; spores dark, with irregular + continuous brownish wing and a few cross-wings. Distr. Polynesia (many islands in the Pacific eastward to Tahiti and Hawaii), New Caledonia, Queensland; in Malesia: New Guinea (lowlands of north and east) and Moluccas (Kei & Tenimber Is., Amboina). Ecol. In open or lightly shaded places, some- times in wet ground, at low altitudes. 144. Sphaerostephanos austerus (BRAUSE) HOLTTUM, comb. nov.—Dryopteris austera BRAUSE, Bot. Jahrb. 56 (1920) 108. — Cyclosorus austerus (BRAUSE) COPEL. Philip. J. Sci. 78 (1951) 446.— Thelypteris austera (BRAUSE) REED, Phytologia 17 (1968) 262.—Type: LEDERMANN 11750, N.E. New Guinea, Sepik Distr., 2070 m (B). Caudex slender, scandent. Stipe 11-18 cm long. Lamina 65-100cm long; 5-6 pairs lower pinnae gradually reduced, all with an acroscopic auricle, lowest 1.3cm long, subtriangular with rounded apex; aerophores not elongate. Hairs on lower surface of rachis dense, short; on upper surface a little longer. Largest pinnae 9-13cm long, 2.2- 3.0cm wide; base subtruncate; apex abruptly short-acuminate; edges lobed 3 way to costa or a little more deeply, lobes falcate, blunt-pointed; costules 5—S.5mm apart, at 60°; veins 7 pairs, slender, basal pair anastomosing, next pair usually both to margin; lower surface of costae bearing dense short pale somewhat antrorse hairs of uni- 1981] THELYPTERIDACEAE (Holttum) 503 form length, similar hairs sparse on costules, scattered on surface between veins; hairs on up- per surface of costae short, scattered longer ones present on costules and veins, surface between veins bearing scattered short appressed hairs. Sori slightly inframedial, exindusiate; sporangia with a few setae; spores minutely spinulose. Distr. Malesia: Papua New Guinea (Sepik), only known from type. 145. Sphaerostephanos mundus (ROSENST.) HOLTTUM, comb. nov.— Dryopteris munda ROSENST. Meded. Rijksherb. n. 31 (1917) 5.— Thelypteris munda (ROSENST.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 252. — Type: ATASRIP s.n. 1903, W. New Guinea (L). Dryopteris oblonga BRAUSE, Bot. Jahrb. 56 (1920) 109.—Cyclosorus oblongus (BRAUSE) CoPEL. Philip. J. Sci. 78 (1951) 446. — Thelypteris oblonga (BRAUSE) REED, Phytologia 17 (1968) 297.— Type: LEDERMANN 10100, N.E. New Guinea, Sepik Distr., 1000 m (B). Dryopteris farinosa BRAUSE, Bot. Jahrb. 56 (1920) 111. — Cyclosorus farinosus (BRAUSE) Co- PEL. Philip. J. Sci. 78 (1951) 446. — Thelypteris far- inosa (BRAUSE) REED, Phytologia 17 (1968) 276. — Type: LEDERMANN 12103, Sepik Distr., 2070 m (B). Caudex a scandent rhizome 7-10 mm diameter, or sometimes free, slender, erect. Stipe 5-10cm long, dark; base of stipe to first large pinna 25- 35 cm. Reduced pinnae 5-7 pairs, smallest 3-4 mm long, uppermost 10 mm, spreading, triangular with base auricled both sides (more on acroscopic side). Lamina 60-100 cm long; pinnae c. 25 pairs, not dimorphous; lower pinnae a little narrowed to symmetric base; aerophores not elongate. Lower surface of rachis sparsely short-hairy; upper sur- face with hairs to 1mm long. Largest pinnae commonly 9-12 cm long, 2.0-2.5 cm wide (on type of D. oblonga 18 x 2.6cm); base broadly cuneate; apex rather abruptly acuminate with cauda 1.5— 3cm long; edges lobed 2/3-3/4 to costa, lobes falcate with blunt forward-pointing tips; costules 4-5 mm apart (to 6.5mm on type of D. oblonga); veins 7-9 pairs, basal pair anastomosing to form a short excurrent vein to sinus, next pair both to margin; lower surface of costae bearing rather sparse short erect hairs (dense with some longer ones on type of D. oblonga), sparse short hairs on costules, veins and surface between veins; hairs on upper surface of costae less than 1 mm long, sparse short hairs on costules and veins, a variable number of fine appressed hairs between veins (in some cases very sparse). Sori inframedial, lowest not or little divergent; indusia absent or very small with a few hairs; sporangia setiferous; spores closely papillose. Distr. Malesia: New Guinea, from West to East. Ecol. Usually climbing on trees in forest, at 1000-2000 m. Note. BRAUSE distinguished D. farinosa by a grey-yellow deposit on the lower surface; this consists of a fungus. In the type of D. munda and others from Western New Guinea, basal veins sometimes meet at the sinus without anas- tomosing. 146. Sphaerostephanos_ obtusifolius (ROSENST.) HOLTTUM, comb. nov. — Dryopteris obtusifolia ROSENST. Fedde Repert. 10 (1912) 336. — Phe- gopteris obtusifolia (ROSENST.) v.A.v.R. Handb. Suppl. (1917) 315.—Cyclosorus obtusifolius (ROSENST.) COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78 (1951) 442.— Thelypteris obtusifolia (ROSENST.) REED, Phytologia 17 (1968) 297. — Type: Frau BAMLER 43, N.E. New Guinea, Sat- telberg (not seen). Caudex short, creeping. Stipe 2-8cm long. Lamina 8 cm long; apex not pinna-like; pinnae c. 8 pairs, | or 2 basal pairs reduced, lowest 4-6 x 3- 5 mm; aerophores slightly swollen. Hairs on lower surface of rachis minute, on upper surface 0.4 mm long, brown. Largest pinnae 1.5—2.0 cm long, 0.8- 10cm wide; base subtruncate, slightly asym- metric; apex broadly rounded, edges sinuous with very slight development of sinus-membranes; costules 2.5mm apart; veins 2-3 pairs, 15 pairs anastomosing; all hairs on lower surfaces minute; on upper surface short hairs on costae only or a few between veins near margin. Sori exindusiate, those on basal veins from adjacent costules sometimes fusing; sporangia setiferous; spores pale, with a moderate number of wings of varying extent. Distr. Malesia: New Guinea several collections; Manus Island). Notes. At BM is a specimen labelled KEyYS- SER 43, from the type locality, named D. obtusi- folia by ROSENSTOCK;; this is probably an error for BAMLER 43 (ROSENSTOCK was describing specimens from both collectors). Also from Sattelberg is Rosenst. Fil. Novoguin. exsicc. 115, coll. BAMLER (B, BM, W). M. SANDS 2682 (K) from limestone on Manus Island is closely similar to the Sattelberg specimens but less hairy on both surfaces of pinnae. (Sattelberg, 147. Sphaerostephanos alcasidii HOLTTUM, sp. nov. Pinnae redactae plurimae, 1.5 cm inter se dis- tantes, aerophoris tumidis tantum constitutae; lamina 80cm longa; pinnae 35-jugatae, maximae 15 1.5cm, 3/5 costam versus lobatae, eglan- dulosae; pagina utroque latere inter venas pilis adpressis vestita; indusia parva brevi-pilosa; sporangia setifera.— Type: ALCASID PNH 1747, Luzon, Mountain Prov., Mt Data (MICH). Caudex not known. Stipe 10cm long; base of stipe to first large pinna 50cm. Reduced pinnae many pairs, 1.5 cm apart, all except the very small upper ones consisting of an aerophore without 504 perceptible blade. Lamina 80cm long; pinnae 35 pairs, lowest narrowed to 8mm wide at the base. Lower surface of rachis near base covered with appressed hairs 0.5 mm long, distally with longer spreading hairs; hairs on upper surface similar. Largest pinnae 15x 1.5cm; base truncate; apex acuminate with entire cauda 1-2cm long; edges lobed a little more than > way to costa, lobes falcate; costules 3.5mm apart at more than 60°; veins 8-9 pairs, basal pair anastomosing, next acroscopic vein to sinus-membrane; hairs on lower surface of costae near base appressed, 0.5 mm long, stally and on costules longer and spread- ing, appressed hairs on lower surface between veins; upper surface of costae bearing hairs 1 mm long, similar hairs scattered on costules and veins, copious appressed hairs 0.3mm long on sur- face between veins. Sori medial; indusia small with short hairs; sporangia sometimes with 1-2 setae. Distr. Malesia: Philippines (Luzon: Mt Data), only known from the type. 148. Sphaerostephanos pilosissimus HOLTTUM, sp. nov. Pinnae redactae 12-jugatae, infimae 0.7. cm longae, superiores 2.5cm, anguste triangulares, crenatae; pinnae evolutae usque 16.5 X2.3 cm, dimidio costam versus lobatae; costae costulaeque subtus pilis erectis vestitae, eglandulosae; indusia pilifera; sporangia setifera.— Type: HOLTTUM 20222, Sumatra, G. Kerinci, 1800-2100 m (SING; BO). Caudex erect, short. Stipe 15 cm long, glabrous; base of stipe to first large pinna 40cm. Reduced pinnae at least 12 pairs, deflexed, basal one 7 mm long, uppermost 2.5 cm long, narrowly triangular with acute apex, crenate edges and slight basal acroscopic auricle. Lamina 65 cm long; pinnae 25 pairs; basal pinnae narrowed and less deeply lobed to base which is 1.3cm wide, basal basiscopic lobes of several successively higher pinnae shor- tened; aerophores swollen, not elongate. Hairs on lower surface of rachis dense, erect, 0.3 mm long, on upper surface more than 0.5mm long, pale. Largest pinnae 16.5 X 2.3 cm; base truncate; apex acuminate with subentire cauda to 2.5cm long; edges lobed about 3 way to costa, lobes slightly oblique with rounded tips; costules 4.5 mm apart; veins 10-14 pairs, basal 2 pairs anastomosing, | pair to sinus-membrane; lower surface of costae and costules densely covered with erect hairs of varied length to 0.5 mm, less dense hairs on veins and surface more than 0.5 mm long; hairs on upper surface of costae 0.7 mm long, costules, veins and surface between them covered with slender ap- pressed hairs 0.3mm long. Sori a little in- framedial, lower ones not divergent; indusia fringed with short hairs and a few on surface; sporangia bearing 2-3 short setae. Distr. Malesia: West Central Sumatra (G. FLORA MALESIANA [ser. II, vol. 1° Kerinci), 2 collections (ALSTON 1800-2100 m. 14185, BM), 149. Sphaerostephanos — erwinii nov. Frondes parvae, lamina apicali 7-8 cm longa, pinnis evolutis 6-jugatis pinnisque redactis 4- jugatis constitutae; pinnae redactae usque 8mm longae; pinnae evolutae usque 3.21.3 cm, subintegrae; indusia parva, pilosa; sporangia setifera. — Type: STRESEMANN 43, Ceram, Cen- tralgeb., G. Hoale, 1000 m (L). Caudex short, suberect. Stipe 7cm long, slen- der, short-hairy; base of stipe to first large pinna 17cm. Reduced pinnae 4 pairs, ovate, uppermost 8 mm, lowest 3-4 mm long. Apical lamina 7-8 cm long, 2.5 cm wide at base, deeply lobed and grad- ing to pinnae; free pinnae 6 pairs. Lower surface of rachis bearing thick brown hairs and shorter pale ones. Largest pinnae 3.2 x 1.3cm, widest a little above middle; base truncate, sometimes with a slight acroscopic auricle; apex abruptly nar- rowed to a rounded or obtuse tip; edges shallowly crenate; costules 3.5 mm apart, at little more than 45°: veins 3 pairs, 1 pairs anastomosing; hairs on lower surface of costae short, erect, with some longer, sparse hairs on costules and veins, short erect hairs on surface between veins; scattered long hairs on upper surface of costules and veins like those on costae, appressed hairs on sur- face between veins. Sori medial; indusia small with hairs more than 0.5 mm long; sporangia seti- ferous. Distr. Malesia: Moluccas (Ceram) and West New Guinea. Note. The plant from W. New Guinea (Kos- TERMANS 2209a, Angi-gita Lake, at 2000 m, BO) is smaller than the type, with pinnae to 2.2x0.8cm, and only 2 pair of reduced basal pinnae. HOLTTUM, | sp. 150. Sphaerostephanos atasripii (ROSENST.) HOLTTUM, Webbia 30 (1976) 194.— Dryopteris atasripii ROSENST. Meded. Rijksherb. n. 31 (1917) 6; C. CHR. Brittonia 2 (1937) 298.— Thelypteris atasripii (ROSENST.) REED, Phy- tologia 17 (1968) 261.— Type: ATASRIP 211, W. New Guinea (L). Cyclosorus diminuens HOLTTUM, Blumea 13 (1965) 135.—Type: BRASS 32106, N.E. New Guinea, Eastern Highlands, Arau, 1400 m (K). Caudex massive, erect. Stipe 13cm long; base of stipe to first large pinna 75 cm. Reduced pinnae 15 pairs, spreading, broadly triangular, lowest 6— 7mm long and 10mm wide at base, uppermost 4-5 x 2.5-3.0cm. Lamina to more than 100cm long; apex not pinna-like; lower pinnae not nar- rowed at base; aerophores slightly swollen. Lower surface of rachis densely covered with erect hairs 0.1-0.2 mm long; upper surface covered with ap- pressed pale hairs 0.5mm long with thicker 1981] spreading brown hairs | mm long on each side of the groove. Largest fertile pinnae 17*2.5cm, sterile to 22 x 3.1 cm; base truncate and somewhat dilated both sides; apex short-acuminate; edges lobed to a depth of 2-3 mm; costules 44.5 mm (fertile), to 6mm (sterile) apart; veins 9-11 pairs, 233 pairs anastomosing, 2-3 pairs to sides of sinus-membrane; lower surface of costae densely covered with short hairs as rachis, hairs on cos- tules and veins more sparse, on sterile pinnae longer than those on costae, copious short erect hairs on surface between veins; hairs on upper surface of costae 0.7 mm long, a few similar hairs on costules and veins, surface between veins + densely covered with fine appressed hairs. Sori medial, lower ones not divergent; indusia covered with hairs 0.3mm _ long; sporangia setiferous, sometimes with an acicular hair on the stalk; spores closely and minutely _ spinulose. Chromosomes: n= 36 (T. G. WALKER). Distr. Malesia: throughout New Guinea, in forest at 700-1400 m. Notes. This species is very similar to S. con- fertus in all characters except the massive erect caudex and much larger fronds which never bear glands. In cultivation the two remain quite distinct in size and habit. 151. Sphaerostephanos albosetosus (COPEL.) HOLTTUM, comb. nov.— Dryopteris albosetosa CopEL. Univ. Cal. Publ. Bot. 18 (1942) 221.— Cyclosorus albosetosus (COPEL.) COPEL. Gen. Fil. (1947) 142; Philip. J. Sci. 78 (1951) 455, pl. 35.— Thelypteris albosetosa (COPEL.) REED, Phytologia 17 (1968) 259.— Type: BRAss 11435, W. New Guinea, 18km north of Lake Habbema, on dry face of a limestone wall, in shade (MICH: BM, BO, L). Caudex short, creeping, 4-Smm_ diameter, bearing close but seriate stipes; stipe 6cm long, basal scales ovate, 1 mm long; base of stipe to first normal pinna 12-24cm; reduced pinnae 6 pairs, lowest 4mm long, others gradually longer; tran- sition to normal pinnae subabrupt, an intermediate pair 1.5cm long, 6mm wide above auricled base, apex broadly obtuse. Lamina 21 cm long; pinnae 10 pairs, lower ones not auricled at their bases; aerophores not elongate. Lower surface of rachis bearing short pale erect hairs with some to 1mm long; hairs 1 mm long also on upper surface. Lar- gest pinnae 4.0 1.0cm; base truncate on acros- copic side, rounded on basiscopic; apex abruptly narrowed to obtuse tip: edges lobed to a depth of 1.5-2 mm, lobes strongly concave beneath (spe- cimen at BO); costules 3.5mm apart; veins 4-6 pairs, thick and prominent on lower surface, basal pair anastomosing, second pair passing to sinus- membrane or margin; lower surface of costae bearing short erect hairs mixed with longer ones to 1mm, more sparse similar hairs on costules, THELYPTERIDACEAE (Holttum) 505 short slender erect hairs on and between veins; upper surface of costae bearing hairs less than 1 mm long, scattered similar hairs on costules and veins, surface between veins covered with fine appressed hairs. Sori medial, basal ones much divergent; indusia small with copious long hairs; sporangia setiferous. Distr. Malesia: W. New Guinea (near Lake Habbema), only known from the type. Ecol. On dry face of limestone wall, in shade. 152. Sphaerostephanos wauensis HOLTTUM, sp. nov. Pinnae redactae usque 9-jugatae, infimae 5 x 4mm, superiores 1.4 * 1.0 cm, triangulares; pinnae evolutae usque 12*2.0cm, 2/5 costam versus lobatae; rachis utrinque pilis patentibus brunneis 1mm vel ultra longis pilisque brevioribus vestita; costae costulae venaeque subtus pilis pallidis 1mm longis praeditae, eglandulosae; indusia magna, tenuia, pilis brevibus instructa; sporangia setifera.— Type: T. NAKAIKE 96, N.E. New Guinea, Morobe Distr., Wau, Mt Missim 1300- 1600 m (K; TNS). Caudex massive, suberect. Stipe 17cm long; basal scales narrow, 7mm or more long; base of stipe to first large pinna 5S cm. Reduced pinnae to 9 pairs, alternate; basal ones 5 x 4mm, uppermost 1.5x1.0cm, triangular with symmetric base, crenate; one pair intermediate pinnae present. Lamina 60 cm long: pinnae 21 pairs; lower pinnae slightly narrowed towards base which may be slightly auricled both sides; aerophores not elon- gate. Rachis bearing on both sides spreading thick brown hairs more than 1mm long (sometimes lacking on lower surface), on lower surface also short erect pale hairs, on upper surface short brown hairs. Largest pinnae 12 x2.0cm, sessile; base broadly cuneate to subtruncate, not dilated; apex short-acuminate; edges lobed rather less than > way to costa; lobes oblique and slightly falcate; costules 4-5mm apart, at 60° or a little less; veins to 8 pairs, 1> pairs anastomosing and 1 pair to sinus-membrane; lower surface of costae bear- ing many spreading pale hairs more than 1mm long with a varied range of shorter ones, hairs on costules and veins sparse, to 1mm long, short erect hairs between veins; upper surface of costae bearing slender pale hairs 1 mm long, similar hairs scattered on costules and veins, short appressed hairs on surface between veins not abundant. Sori inframedial, lower ones not divergent; indusia rather large, thin, bearing hairs 0.3mm _ long; sporangia setiferous. Distr. Malesia: Papua New Guinea (Morobe District), at 1300-1800 m, several collections. Note. A smaller plant has fertile pinnae 6.8 x 1.3 cm which are slightly stalked (WOMERSLEY & MILLAR 8309, from 1800m), otherwise agreeing with the type. [ser. II, vol. 1° FLORA MALESIANA 506 “CN hae Nast ‘ as Mi Gy kam ry ay — Bho 7 yay ay ala Spin ppp als b. venation and position of sori in apical 2 35 lamina, x 3.— P. heterophyllum (PR.) HOLTTUM. c. Fertile and sterile fronds, x of apical lamina, x 3.— P. micropinnatum HOLTTUM. e. Frond, x Fig. 14. Pronephrium womersleyi HOLTTUM. a. Frond, xX 3; d. venation of part 2 3 2 3 3; f. part of apical lamina, x 4. — P. h. venation and position of sori, x 2; i. one sorus, X< 16 (a 2 33 NAKAIKE 69, b BRASS 31670, c-d PRICE & HERNAEZ 162, e-f cult. Kew, g-i HENDERSON 21863). asperum (PR.) HOLTTUM. g. One pinna, xX 1981] THELYPTERIDACEAE (Holttum) 507 18. PRONEPHRIUM PRESL, Epim. Bot. (1851) 258, excl. P. lastreoides PRESL; HOLTTUM, Novit. Bot. Inst. Bot. Univ. Carol. Prag. 1968 (1969) 48; Blumea 19 (1971) 34; Blumea 20 (1972) 105—126.— Type species: Pronephrium lineatum (BL.) PRESL, l.c. (designated by HOLTTUM, I.c. 1969). Haplodictyum PRESL, l.c. 50; CHING, Sunyatsenia 5 (1940) 251, p.p.; COPEL. Fern Fl. Philip. (1960) 377; HoLTTUM, Blumea 19 (1971) 37; Kalikasan 2 (1973) 59, excl. H. majus. — Thelypteris subg. Haplodictyum K. IWATS. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 32.—Type species: Haplodictyum heterophyllum PRESL, l.c. Abacopteris FEE, Gen. Fil. (1852) 309; CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 230; Acta Phytotax. Sinica 8 (1963) 331; HOLTTUM, Rev. FI. Malaya 2 (1955) 285, excl. A. peltochlamys (C. CHR.) HOLTTUM. — Dry- opteris sect. Abacopteris C. CHR. Gard. Bull. Str. Settl. 7 (1934) 247. — Thelypteris subg. Abacopteris K. IWATS. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 34.— Type species: Abacopteris philippinarum FEE, l.c. (desig- nated by HOLTTUM, I.c. 1969) = Pronephrium asperum (PRESL) HOLTTUM. Dimorphopteris TAGAWA & K. IWATS. Acta Phytotax. Geobot. 19 (1961) 8.— Type species: Dimorphopteris moniliformis TAGAWA & K. IWATS. l.c. — Fig. In-o, 14-16. Caudex creeping or suberect (never truly erect); fronds simple or simply pinnate, basal pinnae not reduced but often narrowed at base on basiscopic side, less often on acroscopic; apical lamina pinna-like or widened towards its base and then merging with the upper pinnae; pinnae crenate to entire; veins several pairs (except in a few species with small pinnae), almost all anastomosing, the excurrent veinlets arising from their union in some species free, more often joining to form zig-zag composite veins alternating with the costules; sinus-membranes short or (in species with entire pinnae) lacking; lower surface of old fronds usually pustular when dried; acicular hairs usually present on lower surfaces, hooked in sect. Grypothrix; sessile spherical yellow glands present on lower surface, indusia and/or sporangia in some species; sori in some species exindusiate and then often spreading along the veins; sporangia often bearing short setae, less often spherical glands or both glands and setae; spores of most species with a + continuous longitudinal wing and cross-wings, three species with many small separate wings or papillae. Distr. India & Ceylon; from S. China southwards throughout Malesia; North Queensland; Solomon Islands, New Hebrides, Fiji; c. 68 species, of which 12 not Malesian. Ecol. Most species are terrestrial in forest or on stream-banks in shade (in several cases on rocks only); two are limestone ferns in more exposed positions (P. simillimum, P. scopulorum). Cytol. Chromosome number 36; 10 species investigated, of which P. pentaphyllum, P. peltatum var. peninsulare, P. womersleyi and P. triphyllum are tetraploid. Taxon. CHRISTENSEN (Ind. Fil. 1905) accepted Abacopteris FEE (which he included in Dryopteris) as published in 1843, but that publication consisted of the generic name only with no description. CHING gave generic status to Abacopteris, and cited Aspidium lineatum BL. as type, but FEE did not mention that species when the generic name was validly published by him in 1852; he then listed Pronephrium PRESL as a genus not adopted, as he had seen none of the species (p. 358). 508 FLORA MALESIANA [ser. II, vol. 1° cS = ig NE TST ee Tt Gi & : Fig. 15. Pronephrium amboinense (WILLD.) HOLTTUM. a. Lower pinnae, <3. — P. celebicum (BAK.) HOLTTUM. b. Sterile pinna, x 1.5; c. fertile pinna, x 1.5; d. two indusia, x 100. — P. hewittii (COPEL.) HOLTTuo. e. Middle sterile pinna, x 1.5; f. basal sterile pinna, x 1.5; g. fertile pinna, x 1.5; h. venation of sterile pinna, x 4; i. sorus, x 16.— P. rhombeum (CHRIST) HOLTTUM. j. Fertile frond, x3; k. ster- ile frond, <3; i. sorus, X 16.— P. granulosum (PR.) HOLTTUM. m. Fertile frond, <3; n. sorus, X24 (a from photo of type, b DRANSFIELD 3776, c-d CURTIS 431, e-i MOLESWORTH ALLEN 3048, j—n cult. Kew). 1981] THELYPTERIDACEAE (Holttum) 509 Pronephrium PRESL (1851) comprised four species, which I discussed in 1969, citing P. lineatum (BL.) PRESL as type and accepting the statement by METTENIUS and CHRISTENSEN that the second species P. affine (BL.) PRESL was not distinct from the first. Subsequently (HOLTTUM 1971) I examined the types of both of BLUME’s species and found them to be quite distinct from each other. The third of PRESL’S species is here named P. rhombeum; the fourth (P. lastreoides PRESL) is here transferred to Sphaerostephanos. PRESL’s most distinctive character is dimorphism of fertile and sterile fronds; this is doubtfully true of P. lineatum (the fertile frond of the type is not fully expanded) but is true of species 2 and 3. FEE distinguished Abacopteris mainly by subentire pinnae with meniscioid venation but the excurrent veinlets not free, and indusiate sori. CHING (1938) extended it to admit species in mainland Asia with exindusiate sori and typical meniscioid venation; he also cited absence of sinus-membranes as a generic character, but short membranes are often present (the line of distinction is not a sharp one). CHRISTENSEN (1934) had already added species from Malesia as Dryopteris sect. Abacopteris. In 1971 and 1972, under the earlier name Pronephrium, 1 accepted the generic concepts of CHING and CHRISTENSEN, adding the character absence of reduced basal pinnae, and divided the whole into three sections: Pronephrium, Dimorphopteris and Grypothrix. But some of the species then placed in sect. Pronephrium have resemblances to species in sect. Dimorphopteris and some to species in sect. Grypothrix. I here attempt to adjust this situation by re-arranging the species in two subgenera Pronephrium and Menisciopsis. Haplodictyum PRESL (which I treated as a separate genus in 1971) is included in sect. Pronephrium. But though sections Dimorphopteris and Grypothrix, as re-arranged, appear to be natural groups, the other two sections are probably not; problems of relationships are discussed under the sections. KEY TO THE SUBGENERA AND SECTIONS . Veins in pinnae less than 10 pairs, or if more than 10, sori indusiate; yellow spherical glands often ene at least on sporangia sigh: . . . . 1. SUBG. PRONEPHRIUM 2. Apical lamina pinna-like or much larger than pinnae: cainnce not auricled on acroscopic base; fronds of young plants simple, in P. beccarianum all fronds simple . . . . . .41. Sect. Pronephrium . Apical lamina not ae or if so pinnae auricled on acroscopic base; fronds of young plants rarely Simpleia ee . . . . 2. Sect. Dimorphopteris 1. Veins in pinnae 10 pairs or more, or if Tease excurrent veinlets all free; sori exindusiate; spherical glands never present a Fe 7) ee 22 SUBGS MENTISCIOPSIS 3. No hooked hairs on any part of plant; sori rarely ‘coalescent . . . . . . 3, Sect. Menisciopsis 3. Hooked hairs present on sporangia or on some other part of plant; sori often elongate or coalescent 4. Sect. Grypothrix 1. Subgenus Pronephrium 1. Section Pronephrium Haplodictyum PRESL, Epim. Bot. (1851) 50. The following species, included here in 1972, are now transferred to other sections: the non-Malesian P. lakhimpurense and allied species, also the Malesian P. nitidum, P. repandum and P. acanthocarpum, to sect. Menisciopsis; P. aquatiloides and P. menisciicarpon to sect. Dimorphopteris. The species here included belong to three groups: 1. Species which appear to be related to the type, P. lineatum. Of these, P. euryphyllum has glands on the lower surface and appressed hairs on the upper surface as in P. glandulosum of sect. Dimor- phopteris. P. asperum and P. gymnopteridifrons lack glands but have the same frond-form as the type. None of these have the reddish tinge shown by species of subg. Menisciopsis. But they show resemblances to Sphaerostephanos peltochlamys, S. simplicifolius and S. spenceri, which differ only in the presence of several pairs of much-reduced basal pinnae. 2. Species which have an elongate apical lamina and a few much smaller pinnae, and sori which lack indusia. These are specialized forms the relationship of which it is difficult to suggest. Three of them have glands on their sporangia as in P. lineatum. The most aberrant species, lacking glands, is P. micropinnatum; it may belong elsewhere. 3. Species included in Haplodictyum in my arrangement of the genera in 1971, but excluding H. majus COPEL. which is now placed in Sphaerostephanos because it has much-reduced basal pinnae in addition to normal ones. These species have glands on the lower surface of pinnae and indusiate sori. 510 FLORA MALESIANA [ser. II, vol. 1° I NG 2 Fig. 16. Pronephrium rubicundum (v.A.v.R.) HOLTTUM. a. Complete frond, <3; b. venation of sterile frond, x 3; c. venation of fertile frond and position of sori, x 4; d. costa and costule showing hooked hairs, x 12.— P. ramosii (CHRIST) HOLTTUM. e. Upper part of frond, x3; f. bud at base of a pinna, x 8.— Sporangia, x35, and spores, 235: g—h. P. nitidum HOLTTUM; i-j. P. repandum (FEE) HOLTTuM; k-l. P. triphyllum (Sw.) HOLTTUM (a-d MATTHEW s.n., e-f ELMER 17851, g-I after HOLTTuUM, Blumea 19: 24, fig. 8-9 and SEM photos). 1981] THELYPTERIDACEAE (Holttum) ei Two of them have the peculiar venation in their terminal lamina which caused PRESL to establish the genus Haplodictyum: the veins in the lower part of the apical lamina are forked, their branches anastomosing to form a series of areoles along the costules, also sometimes additional areoles below the sinuses. This venation resembles that of Pleocnemia, and on that account CHING placed Haplodictyum in his family Aspidiaceae, though it is unquestionably thelypteroid. But the third species (P. bakeri) does not have the veins thus forked, though in every other character it resembles the other two. The Haplodictyum type of venation occurs to a small extent at the base of the apical lamina in several species of other genera of Thelypteridaceae where the transition from apical lamina to pinnae is not sharply defined. The peculiar venation is like that of pinnae, of which the upper ones are more or less adherent to the base of the apical lamina. There is a North American fossil named Goniopteris claiborniana BERRY (Bull. Torr. Bot. Cl. 44, 1917, 331, t. 22) which shows the Haplodictyum venation. KEY TO LAE SPECIES 1. Pinnae of fronds of adult plants about equal in size to apical lamina. 2. Glands present on lower surface of pinnae 2. Glands lacking on lower surface of pinnae. 3. Pinnae 2-3 pairs; glands present on indusia or sporangia or both .1. P. euryphyllum . 2. P. lineatum 3. Pinnae to at least 7 pairs; no glands on indusia and sporangia. 4. Pinnae to 30 x 5 cm; spores with a median wing and cross-wings 4. Pinnae to 15 x 2.5 cm; spores with many separate small wings F . 3. P. asperum . 4. P. gymnopteridifrons 1. Apical lamina much larger than pinnae, or fronds simple. 5. Sori exindusiate; glands usually lacking on lower surface between veins. 6. No free pinnae; sometimes one small pair broadly adnate 6. Free pinnae present on adult plants. 7. Sporangia bearing glands near annulus. .5. P. beccarianum 8. Apical lamina to 20 x 3.5 cm; midrib of apical lamina and rachis glabrous on lower surface 6. P. womersleyi 8. Apical lamina to 6X 1.7cm; hairs present on lower surface of midrib of apical lamina and rachis 7. P. melanophlebium 7. Sporangia bearing setae, “not glands, near annulus. 9. Terminal lamina lobed 1/4—1/3 towards midrib; pinnae to at least 1 cm long. 10. Pinnae 4-8 cm long SF Pinnae not over 2 cm long. . No glands on lower surface Glands present on lower surface 9: eee lamina crenate; pinnae 2-S mm long. 3 . Sori indusiate; glands always present on lower surface between veins. 8. P. pentaphyllum . 9. P. brauseanum . 10. P. diminutum 11. P. micropinnatum 5 Veins in basal part of apical lamina forked, their branches anastomosing. 13. Apical lamina widest near its base; pinnae 1 pair 12. P. bulusanicum 13. Apical lamina not widest near its base; pinnae 3 pairs, decreasing downwards 12. Veins in apical lamina not forked 1. Pronephrium euryphyllum (ROSENST.) HOLT- TUM, Blumea 20 (1972) 112.— Dryopteris eury- phylla ROSENST. Meded. Rijksherb. n. 31 (1917) 7.— Thelypteris euryphylla (ROSENST.) REED, Phytologia 17 (1968) 276.— Type: KORTHALS, Sumatra (L; B). Dryopteris urophylla var. teysmannii v.A.v.R. Bull. Jard. Bot. Btzg II, 28 (1918) 24.— Type: TEYSMANN, Sumatra, Loeboe Alang (BO?; not found). Sterile fronds unknown. Fertile fronds consis- ting of terminal lamina to 21 x 5.5 cm and at least 3 pairs of pinnae. Pinnae to 18 X 6cm, base broadly cuneate, sides parallel for most of their length, crenate to depth of 1-1.5 mm, apex abruptly nar- rowed to a cusp 1-2 cm long; costules 44.5 mm apart, at more than 60° to costa, slightly falcate; 13. P. heterophyllum 14. P. bakeri veins c. 15 pairs, almost all anastomosing, excur- rent veinlets not free; lower surface of rachis, costae and costules covered densely with short erect hairs, similar shorter hairs with many glands on surface between veins; upper surface densely covered throughout with short appressed hairs. Sori supramedial, spreading a little along veins; indusia hairy, persistent but shrivelling; sporangia bearing several short setae; spores pale, with con- tinuous wing and cross-wings. Distr. Malesia: Sumatra; the two types cited. Notes. The KORTHALS collection at L con- sists of 5 sheets (2 also at B); on one of the smaller fronds the basal pinnae are somewhat reduced and narrowed at their base, but the lowest (of three pairs) on the largest specimen are not, so that there may have been more pairs of pinnae on il the latter. VAN ALDERWERELT’Ss short descrip- tion of TEYSMANN’s specimen is adequate for identification; the number of pairs of pinnae is not specified. 2. Pronephrium lineatum (BL.) PRESL, Epim. Bot. (1851) 259; Ho_truM, Blumea 19 (1971) 34; 20 (1972) 112.— Aspidium lineatum BL. Enum. PI. Jav. (1828) 144; METT. Farngatt. IV (1858) 264, p.p. — Meniscium lineatum (BL.) KUNZE, Bot. Zeit. 6 (1848) 259.—Nephrodium lineatum (BL.) PRESL, Epim. Bot. (1851) 48; HooK. Spec. Fil. 4 (1862) 74, p.p. — Cyclosorus lineatus (BL.) TARD. & C. CHR. Notul. Syst. 7 (1938) 74, nomen tantum. — Abacopteris lineata (BL.) HOLTTUM, Rev. Fl. Malaya 2 (1955) 292, nomen tantum. — Thelypteris lineata (BL.) K. IwaTs. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 34.— Type: BLUME, Java (L. n. 909, 27-60; isotypes P, PRC). Alsophila fragilis ZOLL. & MOoORITZI, Nat. Geneesk. Arch. Ned. Ind. 1 (1844) 400. — Menis- cium sp. Morirzi, Syst. Verz. (1846) 116.— Meniscium fragile (ZOLL. & MOR.) KUNZE, Bot. Zeit. 6 (1848) 259.—Type: ZOLLINGER 1019, Java, Tjikoya river (L; FI). Dryopteris verruculosa v.A.v.R. Bull. Jard. Bot. Btzg II, 11 (1915) 12; Handb. Suppl. (1917) 177. — Thelypteris verruculosa (v.A.v.R.) REED, Phy- tologia 17 (1968) 323.—Type: BACKER 3954, Java, Pasuruan (BO). Dryopteris menisciicarpa (BL.) POSTH. in Backer & Posth. Varenfl. Java (1939) 61, p.p. Caudex short-creeping or suberect. Stipe of sterile frond 20cm, of fertile 40cm long, pale, glabrescent except for hairs in groove. Sterile frond. Apical lamina to 15 x 4 cm; pinnae 2-3 pairs, to 9.5 3.0cm, widest 1/3 from apex, gradually narrowed to abruptly broad-cuneate base, apex rather abruptly short-cuspidate, edges crenate distally, sinuous towards base; costules 3mm apart; veins 7-8 pairs, almost all anastomosing, excurrent veinlets not free; lower surface with rather sparse erect hairs on costae, much shorter on costules, surface between veins glabrous and finely pustular, no glands present; upper surface hairy on costae only. Fertile pinnae of type not fully expanded, largest possibly 3.0 1.5cm, of ZOLLINGER 1019 9X2.5cm, of BACKER 3954 7 x 2.2 cm; sori inframedial; indusia small, bearing a few short hairs and glands; sporangia with glands, or both glands and setae; spores with translucent wing and cross-wings. Distr. Malesia: Central & East Java (3 collec- tions) and Sabah (Bettotan, KLOss SFN 19083); Sulu. Archipelago, Tawi-Tawi (ALCASID & CELESTINO PNH 7507). Notes. METTENIUS (1858) included Aspidium - affine BL. as a synonym, and his description is composite; HOOKER (1862) copied him; later authors mostly described A. affine under the name lineatum, not noticing that BLUME’s descriptions FLORA MALESIANA {ser. II, vol2 1? of the two species indicate fronds of quite different form. BLUME gave Noesa Kambangan as type local- ity, but the only specimen named in his hand is unlocalized; it agrees well with his description. The published description of Alsophila fragilis ZOLL. & Mor. is doubtfully valid, but the spe- cimens of ZOLLINGER 1019 certainly agree with BLUME’s. The sporangia of the type have both glands and setae; those of ZOLLINGER 1019 glands only. 3. Pronephrium asperum (PRESL) HOLTTUM, Blumea 20 (1972) 112.— Polypodium asperum PRESL, Rel. Haenk. (1825) 24, t. 3, f. 4, non LINN.; HOLTTUM, Novit. Bot. Inst. Bot. Univ. Carol. Prag. 1968 (1969) 18.— Goniopteris aspera PRESL, Tent. Pterid. (1836) 183, nom. nov. — Aspidium asperum (PRESL) MEtTT. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 225, excl. syn. Neph- rodium latifollum PRESL.— Dryopteris presliana CHING in C. CHR. Ind. Fil. Suppl. 3 (1934) 95, nom. nov. superfl.— Abacopteris presliana (CHING) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 248, p.p.— Abacopteris aspera CHING, Acta Phytotax. Sinica 8 (1963) 332. — Thelypteris aspera REED, Phytologia 17 (1968) 261. — Type: HAENKE, Luzon (PRC). Abacopteris philippinarum FEE, Gen. Fil. (1852) 310, t. 18C f. 1. — Cyclosorus philippinarum (FEE) COPEL., Fern Fl. Philip. (1960) 372.— Nephrodium latifolium PRESL, Epim. Bot. (1851) 45, p.p.— Type: CUMING 16, Luzon (isotypes BM, G, K, LE). Meniscium kennedyi F.v.M. Fragm. 4 (1864) 165.— Goniopteris kennedyi (F.v.M.) BAILEY, Handb. Queensl. Ferns (1874) 41, f. 31.— Type: DALLACHY & KENNEDY, Rockingham Bay, Queensland (MEL; K). Dryopteris urophylla (WALL. ex HOOK.) C. CHR. var. novoguineensis ROSENST. in Fedde Rep. 10 (1912) 336; v.A.v.R. Handb. Suppl. (1917) 177.— Type: G. Bamler 80, Sattelberg, New Guinea (orig. not seen; probable isotypes BM, L, W, as Rosenst. fil. novoguin. exsic. 174). Abacopteris multilineata (WALL. ex HOOK.) CHING var. malayensis HOLTTUM, Rev. FI. Malaya 2 (1955) 297, f. 173, nom. illeg. — Thelyp- teris multilineata (WALL. ex HOOK.) CHING var. malayensis (HOLTTUM) REED, Phytologia 17 (1968) 294 (type not cited). — Lectotype: HOLT- TUM SFN_ 15303, Trengganu, Kuala Telumong (SING; K). Aspidium repandum sensu BL. Enum. PI. Jav. (1828) 144, excl. syn. Nephrodium glandulosum sensu J. SM. in Hook. J. Bot. 3 (1841) 411 (misidentification of CUMING 16); DIELS in E. & P. Nat. Pfl. Fam. I Abt. 4 fig. 92G “nach Fée”’ (inaccurate copy of FEE, Gen. Fil. t. 18C, f. 1).— Aspidium glandulosum sensu KUNZE, Bot. Zeit. 6 (1848) 260, quoad Cuming 16 1981] THELYPTERIDACEAE (Holttum) S13 tantum; METT. Farngatt. IV (1858) 111, quoad syn. Abacopteris philippinarum FEE tantum. Polypodium urophyllum sensu. BENTH. FI. Austral. 7 (1878) 766, excl. BEDD. Ferns Brit. Ind. t. 3.—Nephrodium urophyllum sensu. BEDD. Handb. (1883) 274, p.p.; sensu RAcIB. FI. Btzg 1 (1898) 184. — Dryopteris urophylla sensu v.A.v.R. Handb. (1908) 216, p.p.; sensu BACKER & PosTH. Varenfl. Java (1939) 64, p.p. — Fig. 14g-i. Caudex short-creeping. Stipe to 70cm or more long, slightly flushed red, glabrescent, base at first covered with narrow thin scales. Lamina to 60 cm or more long; pinnae 6-8 pairs, lower ones + narrowed at base, lowest sometimes reduced, tex- ture thin; apical lamina more strongly crenate than pinnae. Largest pinnae to 37 x 6cm, sessile, base abruptly broad-cuneate, apex short-acuminate, edges subentire to distinctly crenate, parallel for most of their length; costules 5—5.5 mm apart, at 60° to costa; veins 12-15 pairs, at c. 45°, nearly all anastomosing, excurrent veinlets usually not free; sinus-membrane short but distinct; lower surface of costae, costules and veins bearing short hairs, surface between veins glabrous or rarely with short erect hairs, finely verrucose when dry; upper surface hairy only on costa. Sori medial, lowest ones supramedial and + coalescent; indusia small, with or without short hairs; sporangia of type lacking setae, a hair of 2 cells on the stalk; spores dark with a continuous wing and cross-wings. Distr. Polynesia (Fiji), Solomon Is. (Bougain- ville), North Queensland and throughout Malesia. Ecol. In lowland forest. Notes. Until the work of CHING (1938) this species and others were much confused with Polypodium urophyllum WALL. (= Pronephrium repandum (FEE) HOLTTUM), the present species also with Nephrodium latifollum PRESL (= Pronephrium menisciicarpon) and Aspidium glandulosum BL. (Pronephrium glandulosum); under the last-named METTENIUS also cited spe- cimens of Sphaerostephanos peltochlamys (C. CHR.) HOLTTUM (ZOLLINGER 2608, 2920). Spe- cimens from New Guinea and Queensland often have 1-2 short setae on sporangia. 4. Pronephrium gymnopteridifrons (HAYATA) HOLTTuUM, Blumea 20 (1972) 112; C. M. Kuo, FI. Taiwan 1 (1975) 429, pl. 150; EprE, Ferns Hong Kong (1978) 151, f. 73.— Dryopteris gymnop- teridifrons HAYATA, Ic. Pl. Formos. 8 (1919) 148, fig. 75, 76.—Abacopteris gymnopteridifrons (HAYATA) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 251.—Type: T. SOMA s.n. 1912, Taiwan (isotypes TAI). Polypodium urophyllum var. uniseriale HOOK. Spec. Fil. 5 (1863) 10, new name for Polypodium granulosum sensu BENTH. Fl. Hongkong (1861) 459. — Lectotype (HOLTTUM Ke 1972): URQUHART, Hong Kong (K). Cyclosorus pustulosus COPEL. Philip. J. Sci. 81 (1952) 37; Fern Fl. Philip. (1960) 374. — Thelyp- teris pustulosa (COPEL.) REED, Phytologia 17 (1968) 307. — Type: Copeland 218, Lamao River, Luzon (MICH). Dryopteris glandulosa sensu CHR. Philip. J. Sci. 2 (1907) Bot. 205, p.p. Abacopteris presliana sensu CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 249, p.p. Differs from P. asperum as follows. Caudex (at least in some cases) long-creeping; frond smaller, pinnae commonly to 15 x 2.5 cm (exceptionally to 25x 4cm); sori more often coalescent (in some cases many pairs); sporangia often setiferous; spores with many small wings. Distr. Southern China and Malesia: Philip- pines (Luzon). 5. Pronephrium beccarianum (CESATI) HOLT- TUM, Blumea 20 (1972) 107; BROWNLIE, Pterid. Fiji (1977) 256, pl. 26, f. 2.— Meniscium bec- carianum CESATI, Rendic. Acad. Napoli 16 (1877) 27, 30.— Dryopteris cesatiana C. CHR. Ind. Fil. (1905) 257, nom. nov. (not D. beccariana (CESATI) C. CHR.); COPEL. Bishop Mus. Bull. 59 (1929) 48. — Phegopteris beccariana (CESATI) v.A.v.R. Handb. (1908) 509.—Cyclosorus beccarianus (CESATI) COPEL. Gen. Fil. (1947) 142; Philip. J. Sci. 78 (1951) 460.— Thelypteris cesatiana (C. CHR.) REED, Phytologia 17 (1968) 267 (not T. beccariana (CESATI) REED).—Type: Beccari, Andai, W. New Guinea (FI, Herb. Becc. 12727). Goniopteris simplicifolia (J. SM. ex HOOK.) CARR. var. vitiensis CARR. in Seem. FI. Vitiensis (1893) 366. — Type: SEEMANN 736, Fiji (K). Dryopteris oblanceolata COPEL. Philip. J. Sci. 9 (1914) Bot. 3.—Phegopteris oblanceolata (COPEL.) v.A.v.R. Handb. Suppl. (1917) 320.— Type: C. KING 394, Taupota, Papua (MICH; SING). Phegopteris rutteniana v.A.v.R. Bull. Jard. Bot. Btzg II, 28 (1918) 36.— Type: KORNAssI 725, Ceram (BO). Caudex short-creeping. Stipe 2-10cm_ long, usually longest in fertile fronds, near base minutely hairy, above base hairy in groove. Lamina simple, to 35cm long, sterile to 5cm wide, fertile usually narrower, widest 1/3 from short-acuminate apex, very gradually narrowed to base, edges entire or sinuous, base sometimes dilated or with a pair of small broadly adnate separate leaflets; main veins 3.5-4mm apart, at broad angle to midrib, distally upcurved; veins 9-10 pairs, almost all anastomosing, excurrent veinlets mostly not free; lower surface between veins pustular, sparse short hairs present on main veins and veins, sometimes also rudimentary scales with red end-cell; upper surface short-hairy on midrib, sparsely on main veins. Sori exin- dusiate, spreading along veins, those on connivent veins often joining, sori rarely extending a little along excurrent veinlets; sporangia bearing small 514 FLORA MALESIANA {ser. II, vol. 1° glands and sometimes a short seta; spores pale with rather broad minutely erose translucent wing and a few cross-wings. Distr. Polynesia (Fiji), Solomon Is., in Malesia: New Guinea and Moluccas (Ceram). Ecol. In forest, usually near streams, to 1000 m or a little higher. Notes. In Fiji the basal part of the lamina is more often strongly sinuous and there may be 3 pairs of broadly adnate basal leaflets; glands on sporangia are larger than in New Guinea. 6. Pronephrium womersleyi HOLTTUM, Blumea 20 (1972) 108.— Cyclosorus beccarianus sensu HOLTTUM & Roy, Blumea 13 (1965) 134.— Type: FLOYD & WOMERSLEY NGF 6308, N.E. New Guinea, Eastern Highlands, Goroka Subdistr. 2200 m (LAE; BM). — Fig. 14a-b. Differs from P. beccarianum as follows. Adult plants always with several (to 10) pairs of free + orbicular pinnae 7-10mm long, 7-8mm_ wide; veins in pinnae forked, the branches joining to form a series of areoles; apical lamina smallest on fronds with most pinnae; lower surface of midrib and veins glabrous or nearly so; sori (on apical lamina only, or rarely on a pinna) on lower 1-3 pairs of veins coalescent, more distal sori on basal part of each vein; sporangia bearing small glands, not setae. Distr. Solomon Is. (Guadalcanal), in Malesia: Papua New Guinea, 1400-2400 m. Note. A plant cultivated at Kew was found to be tetraploid (HOLTTUM & Roy l.c.). 7. Pronephrium melanophlebium (COPEL.) HOLTTuM, Blumea 20 (1972) 108.— Dryopteris melanophlebia COPEL. Philip. J. Sci. 6 (1911) Bot. 147. — Phegopteris —melanophlebia (COPEL.) v.A.v.R. Handb. Suppl. (1917) 319. — Cyclosorus melanophlebius (COPEL.) COPEL. Fern FI. Philip. (1960) 359. — Thelypteris melanophlebia (COPEL.) REED, Phytologia 17 (1968) 292. — Type: MER- RILL 6959, Canlaon Volcano, Negros (isotypes B, BO, E, K, L, NSW). Dryopteris canescens’ var. subsimplicifolia CHRIST, Philip. J. Sci. 2 (1907) Bot. 199. — Type: WHITFORD 784, Luzon (not seen). In habit similar to P. womersleyi but much smaller. Stipe 2-3cm (sterile), to 8cm (fertile), covered with hairs more than 0.5 mm long. Apical lamina of sterile fronds to 5 x 1.8 cm, of fertile to 6X 1.4cm; pinnae 3 pairs, sterile 6 x 5 mm, fertile 5x4mm and more widely spaced than sterile, oblong-ovate with midrib and 4 pairs of forked veins, the branches anastomosing; lower surface of costa with copious spreading pale hairs to 1 mm long, sparse shorter hairs on veins; on upper sur- face a few short appressed hairs between veins, especially near margins; sori from base of veins, + elongate, exindusiate, those on connivent veins sometimes meeting; sporangia bearing glands. Distr. Malesia: Philippines (Negros, Mindanao (ELMER 10784, Mt Apo), (?) Luzon). Note. ELMER’s specimen from Mindanao was distributed under the name Dryopteris canescens var. subsimplicifolia; it agrees with CHRIST’s brief description of that variety, but ELMER 10278, also so named, is Pronephrium granulosum. 8. Pronephrium pentaphyllum (ROSENST.) HoLtTtruM, Blumea 20 (1972) 108.— Dryopteris pentaphylla ROSENST. in Fedde Rep. 12 (1913) 529. — Phegopteris —_ pentaphylla (ROSENST.) v.A.v.R. Handb. Suppl. (1917) 317. — Thelypteris pentaphylla (ROSENST.) REED, Phytologia 17 (1968) 303.— Type: KEYSSER 186, N.E. New Guinea, Sattelberg 1400 m (S-PA; B, BM). Caudex long-creeping, 2-3 mm diameter when dry, with fronds 1-2cm apart. Stipe of sterile frond c. 10cm long, of fertile to 20cm, bearing pale spreading hairs more than 0.5mm _ long; fronds consisting of apical lamina and 1-2 pairs of pinnae. Apical lamina 10-15 x 1.5-3.0cm, lobed 1/3 towards midrib, narrowed slightly to + trun- cate base, apex acuminate; main veins c. 5mm apart, veins to 10 pairs, 2 pairs anastomosing, 13-2 pairs passing to long sinus-membrane; lower sur- face of midrib and veins bearing pale spreading hairs | mm long, rather sparse slender erect hairs between veins, no glands (but see note below); upper surface glabrous except for midrib. Pinnae opposite, 4-8 cm long, 1-2.5 cm wide (sterile wider than fertile), widest above middle, lobed 1/4-1/3 towards costa; veins 4-6 pairs; pubescence as apical lamina. Sori inframedial, spreading a little along the veins, basal ones only sometimes confluent; sporangia bearing 2-6 setae, on stalk a hair of 2 cells; spores pale, opaque, with rather broad translucent wing and _ cross-wings. Chromosomes: n= 72 (T. G. WALKER). Distr. Malesia: N.E. New Guinea (also New Ireland). Ecol. In forests, 1200-2000 m. Notes. One specimen from Mt Oga, Western Highlands (B. S. CROXALL 4362) differs from all others seen in having scattered yellow glands in- stead of hairs between veins on lower surface of sterile fronds, and a yellow spherical gland at tip of hair on stalk of sporangium; also in the presence of fairly abundant short suberect hairs between veins of upper surface. 9. Pronephrium brauseanum HOLTTUM, Blumea 20 (1972) 107. — Dryopteris canescens var. novo- guineensis BRAUSE, Bot. Jahrb. 49 (1912) 22; CoPEL. Philip. J. Sci. 78 (1951) 460.— Type: SCHLECHTER 18844, N.E. New Guinea, Bismarck Mts 1100 m (B; K, L, BISH). Caudex short-creeping; fronds dimorphous. Sterile fronds. Stipe 1.5-3 cm long; frond consis- ting of terminal lamina and 1-2 pairs of pinnae close below it; terminal lamina 5-6 x 1.7-2.1 cm, 1981] THELYPTERIDACEAE (Holttum) 515 lobed 1/4 towards midrib, main veins 4—4.5 mm apart, veins 5-6 pairs, 15 pairs anastomosing, 1 pair to sinus-membrane, strongly prominent on lower surface and bearing stiff hairs to 1 mm long, a few glands and short erect hairs present on surface between veins; upper surface bearing copious appressed hairs 0.3-0.4 mm long between veins; pinnae 1.0-2.0cm long, 0.5-1.1cm wide, sessile, edges crenate, pubescence as apical lamina. Fertile frond. Stipe to 12cm long; pinnae more widely spaced than sterile, not opposite, to 1.2 x 0.6cm with stalks 1 mm long, subentire with rounded apex, lowest pinna longest; sori (mainly on the terminal lamina) a little elongate, exin- dusiate; sporangia copiously setose. Distr. Malesia: Eastern New Guinea at 1100- 1700 m, in forest (5 collections). 10. Pronephrium diminutum (COPEL.) HOLT- TUM, Blumea 20 (1972) 115.— Dryopteris diminuta COPEL. Philip. J. Sci. 40 (1929) 298. — Cyclosorus diminutus (COPEL.) COPEL. Gen. Fil. (1947) 142; Fern FI. Philip. (1960) 359. — Type: COPELAND s.n. Nov. 1911, Mindanao, San Ramon 1200 m (MICH). Caudex creeping, 1mm diameter; scales 2mm long. Stipe of sterile fronds 1-3 cm long, of fertile to 6cm, covered with short pale erect hairs; frond consisting of apical lamina and 1-2 pairs of pinnae (not opposite). Sterile fronds. Apical lamina 2- 4cm long, 1.5 cm wide, lobed 1/4 towards midrib; main veins 4-4.5 mm apart, veins 3-4 pairs, 1-2 pairs anastomosing, prominent on lower surface; lower surface of midrib bearing stiff pale hairs to 1mm long, shorter hairs on veins with a few glands, slender erect hairs and a few glands on surface between veins; upper surface covered with slender appressed hairs; pinnae broadly ovate, and slightly lobed, to 8 mm long, on stalks more than 1mm long. Fertile fronds somewhat smaller than sterile, with more widely-spaced pinnae; sori exindusiate, those on lower veins elongate and coalescing, on distal veins round; sporangia bearing several short setae distally and sometimes a yellow gland on the stalk. Distr. Malesia: Philippines (Luzon, Mindanao). Note. COPELAND made 2 collections at San Ramon, one earlier than the type (1907). The only other collection is by M. G. PRICE (n. 2720B) from Mt Banahaw, Luzon, at 1000m “on rocky slope 10m above stream”’; it only differs from the type in slightly larger fronds with fertile pinnae to 1.1 x 0.8 cm. 11. Pronephrium micropinnatum HOLTTUM, Blumea 20 (1972) 108. — Type: DARBYSHIRE & HOOGLAND 8014, N.E. New Guinea, Sepik Distr. (CANB; BM, L, LAE). — Fig. 14g-f. Caudex short, suberect. Stipe to 6cm long, minutely hairy on abaxial side, hairs in adaxial groove to 1mm long. Fronds uniform, consisting of terminal lamina and 2-3 pairs of pinnae 1- 1.5cm apart. Terminal lamina to 18 x 1.4m, nar- rowed both to apex and base, edges crenate; main veins 4-5 mm apart, each with 4-5 pairs of veins of which 1) pairs anastomose, | pair passing to sides of short sinus-membrane; lower surface of midrib covered with pale erect hairs almost 1 mm long, similar hairs sparse on main veins, short erect hairs between veins; upper surface glabrous apart from midrib; sori all medial, not elongate, exindusiate; sporangia bearing several setae nearly as long as the body; spores with many small wings. Pinnae of type c. 1mm long, 2mm wide; of plants cultivated at Kew to 53mm; veins free; sori present on largest pinnae. Distr. Malesia: N.E. New Guinea (Sepik and Madang Districts, 3 collections). Ecol. On shaded river bank (type) and sand- stone walls of river gorge, at 30-200 m. 12. Pronephrium bulusanicum (HOLTTUM) HOLTTUM, comb. nov. — Haplodictyum bulus- anicum HOLTTUM, Kalikasan 2 (1973) 61.— Type: ELMER 16585, Mt Bulusan, Luzon (K; BO, PGE): Stipe of sterile frond 5-6 cm, of fertile 9-14 cm long. Lamina to 12cm long, 2.2-3.0cm wide, short-acuminate, at the middle lobed 2/5 towards midrib, more deeply lobed at the base which is not narrowed, the basal lobes almost or quite free; veins in lobes near apex of frond 7-8 pairs, not forked, 13-2 pairs anastomosing, next pair to sides of sinus-membrane; veins in lower lobes forked, forming areoles on each side of costule and ad- ditional irregular ones on each side of sinus- membrane and below it; veins and costules prom- inent on both surfaces; on lower surface of midrib stiff pale hairs 1 mm long and shorter ones abun- dant, shorter hairs present on costules and veins, yellow glands and fine erect hairs between veins; upper surface covered with short appressed hairs, longer ones also on midrib and costules. Pinnae 0.9-1.5 x 0.5-0.8 cm, crenate, narrowed towards base on basiscopic side. Sori inframedial; indusia rather large, bearing a variable number of short hairs and a few glands; sporangia glandular; spores with a wing and cross-wings. Distr. Malesia: Philippines (Luzon, Sorsogon Province, 2 collections from type locality, | from Mt Pulog). 13. Pronephrium heterophyllum (PRESL) HOLT- TUM, comb. nov. — Haplodictyum heterophyllum PRESL, Epim. Bot. (1851) 51; FEE, Gen. Fil. (1852) 309, t. 18C, f. 2; COPEL. Fern Fl Philip. (1960) 378; HoLtTruM, Kalikasan 2 (1973) 62.— Aspidium heterophyllum (PRESL) HOOK. Ic. PI. 10 (1854) t. 920; COPEL. Polypod. Philip. (1905) 39. — Pleocnemia heterophylla (PRESL) v.A.v.R. Handb. (1908) 171.— Type: CUMING 322, Samar (PREABRES KEE): 516 FLORA MALESIANA [ser. II, vol. 1° Aspidium blumei KUNZE ex METT. Fil. Hort. Lips. (1856) 94, t. 22, f. 5. — Aspidium blumei var. subpinnata METT. Farngatt. IV (1858) 98, excl. syn. Polypodium canescens BL. et CUMING 251. — Type: CUMING 322 (formerly at LZ, now lost). Aspidium canescens (BL.) CHRIST var.; CHRIST, Farnkr. der Erde (1897) 244.— Dry- opteris canescens (BL.) C. CHR. var.; C. CHR. Ind. Fil. (1905) 256. — Fig. 14e-d. Caudex short-creeping; fronds dimorphous. Sterile fronds. Stipe 1-3 cm long, with spreading pale hairs 1mm long; fronds consisting of ter- minal lamina and 1-3 pairs of pinnae. Terminal lamina to 17 X2.8cm, apex rather abruptly poin- ted, base somewhat narrowed, middle part lobed not quite half-way to midrib, main veins 5-6 mm apart, veins in lobes 6-7 pairs, forked and anas- tomosing as in lower part of lamina of P. bulus- anicum; base of lamina more deeply lobed, with transition to 2-3 pairs of pinnae close below it: hairs and glands as in P. bulusanicum. Fertile fronds. Stipe to 7cm long; terminal lamina 13 x 1.6cm, acuminate, lobed less deeply than sterile, veins closer and fewer than in sterile; pinnae 2 pairs, rather irregularly spaced, lowest 3-4mm long; sori in terminal lamina medial or inframedial on the veins, not elongate; indusia with short hairs; sporangia usually with 1 gland; spores with translucent wing and cross-wings. Distr. Malesia: Philippines (Samar, Southern Luzon, Mindanao). Ecol. On rocks in stream-beds at 200-350 m (M. G. PRICE). 14. Pronephrium bakeri (HARR.) HOLTTUM, comb. nov. — Nephrodium bakeri HARR. J. Linn. Soc. Bot. 16 (1877) 29; BAk. in Hook. Ic. Pl. 17 CoPEL. Philip. J. Sci. 2 (1907) Bot. 405; v.A.v.R. Handb. (1908) 208.— Haplodictyum _ bakeri (HARR.) CHING, Sunyatsenia 5 (1940) 251; HOLTTUM, Kalikasan 2 (1973) 63.— Cyclosorus bakeri (HARR.) COPEL. Gen. Fil. (1947) 142; Fern Fl. Philip. (1960) 358.— Thelypteris bakeri (HARR.) REED, Phytologia 17 (1968) 262. — Type: STEERE, Panay (MICH, K). Aspidium canescens (BL.) CHRIST var. sensu CHRIST, Farnkr. der Erde (1897) 244.— Dry- opteris canescens (BL.) C. CHR. var. sensu C. Cur. Ind. Fil. (1905) 256. Caudex short-creeping. Stipe of sterile frond 5 cm, of fertile to 11 cm long; fronds consisting of terminal lamina and 1 pair (rarely 2 pairs) of pinnae; terminal lamina to 13 x2.1cm, apex ab- ruptly short-pointed, middle part lobed 1/3 towards midrib, somewhat narrowed towards base which is sometimes dilated; main veins to 5mm apart, veins to 7 pairs, not forked except some- times in basal lobes of largest fronds; lower sur- face with pubescence and glands as in P. bulus- anicum; pinnae opposite or not, to 107mm, apex rounded, veins anastomosing. Sori medial on veins of terminal lamina, sometimes also on pin- nae, not elongate; indusia hairy; sporangia some- times with a gland; spores with a wing and cross- wings. Distr. Malesia: Philippines (Luzon; Panay; Negros Oriental; Mindanao, cf. COPELAND, not seen). Ecol. In forest at 500-800 m, near river (M. G. PRICE). Notes. CLEMENS 16530, from Isabella Prov., Luzon, has dimorphic fronds; sterile fronds have an apical lamina to 2.5cm wide, fertile to 1.4 cm. Some fronds also have a pair of very small pinnae some distance below the normal ones. (1886) t. 1664.— Dryopteris bakeri (HARR.) 2. Section Dimorphopteris (TAGAWA & K. Iwats.) HoLtTTuM, Blumea 19 (1971) 36; Blumea 20 (1972) 113-121. — Dimorphopteris TAGAWA & K. Iwarts. Acta Phytotax. Geobot. 19 (1961) 8. — Thelypteris subg. Dimorphopteris K. wats. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 35. Type species: Pronephrium moniliforme (TAGAWA & K. IwaTs.) HOLTTUM. Plants of moderate size, always with a relatively short deltoid apical lamina on fronds of adult plants, young plants rarely (as in P. menisciicarpon) having simple fronds 10 cm or more long; sterile and fertile fronds often dimorphous, the latter with longer stipes and smaller pinnae than the former; pinnae in most cases distinctly auricled on the acroscopic base. Distr. 45 species, all Malesian (P. beccarianum extending to Fiji) except P. articulatum (HOULST. & Moore) HOLTTvuM in Ceylon, India to S. China, and P. palauense (HOSOKAWA) HOLTTUM in Palau island. Notes. I suggest that a probable prototype for this section is shown by the Indian species P. 1981] THELYPTERIDACEAE (Holttum) 547 articulatum (see HOLTTUM 1972: 116); the Malesian P. glandulosum differs from it in smaller size and greater abundance of glands. The other species are mostly still smaller, with a pronounced tendency to dimorphism, the most extremely dimorphic being the type species P. moniliforme. This section agrees with Sphaerostephanos in the presence of spherical yellow glands, on either the lower surface of pinnae or on indusia or sporangia, in a majority of species; it differs from Sphaeroste- phanos in the absence of much-reduced basal pinnae and (in most species) in the pustular nature of the lower surface of dried fronds. But there appears to be an evolutionary trend within Sphaerostephanos towards reduction in the number of such small basal pinnae, and there are species in which some fronds may have 1-2 pairs of reduced pinnae and others none, on the same plant. I see no sharp distinction between such species and others which have no reduced basal pinnae. Thus on this character there appears to be no clear separation of this section from Sphaerostephanos. The section does however give the impression of being natural if one excludes from it species with deeply lobed non-pustular normal pinnae (e.g. Nephrodium diversilobum PRESL); such species are here placed in Sphaerostephanos (in some cases they have obvious relatives in that genus). Even when this is done, the present section may still include species of two different ancestral origins: those derived from ancestors with, and without, reduced basal pinnae. The whole problem of recognizing affinities is complex; the present arrangement is one of convenience pending further investigation of an experimental nature. The present arrangement includes some species placed in sect. Pronephrium in 1972; P. buwaldae HOLTTUM, included here in 1972, is now transferred to Christella. KEY 1O THE SPECIES 1. Sori exindusiate. 2. Pinnae not more than 3 cm long. 35 Upper surface short-appressed-hairy; som elongate << . . 2. = . . . =. . 15. PB: millarae 3. Upper surface glabrous; sori not elongate «Lee 8 4 OE ee PE OLGA: peramelense 2. Pinnae longer on mature plants. 4 RRS. pMMTAO MATING. 5 0 216 5 obo oie go ow 6 6 6 6 oo no JU einen 4. Fertile pinnae to at least 1 cm wide. 5. Surfaces between veins glabrous. G, SoOminGR SoG 5 66 456 o coon is 6 6 4616 4 5p o o o olen IE Amini: 6. Sporangia not setiferous. Veeeinnae.2—sicm wide: 22h". 75 “2 Slo. 30 bie PU eee ee AO ee fiemulom 7. Pinnae not over | cm wide eer Beet ce. 1 Set ile Cees Ge) Ree OE faquatiloides 5. Surfaces both short-hairy between veins. SambinnactwGwlSipairseus’ “koe cake ie eee | ee 28S fee ee ee ye ee ee simillimum Seekinnae 2=3 pairs: Wes Ae GI Are i ees eis B22 PS oiluwense 1. Sori indusiate. 9. Fronds simple, 10-20 cm long with cordate base away ee ees] Ae 7- 4 Pmenisciicarpon 9. Fronds pinnate or, if simple, smaller and sterile. 10. Lower surface between and/or on veins bearing sessile spherical glands. 11. Veins free aie aren). (oe ee), Saar Pee ae 23 SP akielibercii 11. Veins anastomosing. 12. Free pinnae not over 7 pairs, not over 2.0 cm wide. 13. Hairs on lower surface of costules and on upper surface between veins not appressed 24. P. amphitrichum 13. Hairs on lower surface of costules and on upper surface ate 14. Indusium bearing glands is ate 3 a aL, . . . . 25. P. clemensiae 14. Indusium bearing hairs . . . Bry... Sigh nn 2a) +: 25b. P. clemensiae var. degenerum 12. Free pinnae more numerous or wider. 15. Lower pinnae with stalks 1-2mmlong . ..... . . . . . . 26. P. glandulosum 15. Lower pinnae sessile. 16. Hairs on lower surface of costae and costules erect . . ea. See 7 Bdebile 16. Hairs on lower surface of costae and costules slender, appressed. 17. Sterile and fertile pinnae both 8-9cmlong . . . . . 25b. P. clemensiae var. degenerum 17. Sterile and fertile pinnae dimorphous, smaller. 18. Sterile pinnae c. 4.5 x 1.4 cm; rachis almost glabrous on abaxial surface 28. P. minahassae 18. Sterile pinnae c. 2.0 x 0.8 cm; rachis bearing copious erect hairs on abaxial surface 29. P. solsonicum 10. Lower surface between and on veins lacking sessile spherical glands. 19. Glands present on sporangia, in some cases setae also. 518 20. Fertile pinnae 3 FLORA MALESIANA —4mm wide; sterile to 10 mm wide [ser. II, vol. 1° 30. P. celebicum 20. Fertile pinnae 10mm or more wide; little dimorphism. 21. Pinnae 6-10 pairs; sporangia bearing small glands only. 22. Pinnae 2-3 cm wide ses Pinnae not over 1 cm wide . Pinnae 3-4 pairs; sporangia often with both ‘glands and setae 19. P. firmulum 20. P. aquatiloides 31. P. granulosum a ae lacking glands; setae present in most cases. 23. Fertile pinnae less than 1.5 cm long; sori in | row on each side of costa . 32. P. samarense 23. Fertile pinnae to at least 2cm long; sori in several rows. 24. Pinnae and/or aa on both young and adult plants + dilated and irregularly lobed distally 24. Pinnae not thus dilated nor irregularly lobed. 33. P. X xiphioides 25. Basal pinnae narrowly cuneate at base both sides, not auricled; fertile pinnae not or little narrower than sterile. .18. P. amboinense 25. Basal pinnae otherwise, usually auricled; fertile pinnae in ) most cases narrower than sterile. 26. Indusia not very small, bearing many short hairs. 27. Upper surface of pinnae bearing short suberect hairs between veins. 28. Pinnae 7 pairs; indusia thin with some superficial hairs 28. Pinnae 15 pairs; indusia firm with stiff marginal hairs 27. Upper surface glabrous between veins. 29. Sporangia setiferous. 30. Sterile pinnae barely 1 cm wide . 34. P. trachyphyllum . 35. P. thysanoides . 36. P. hosei 30. Sterile pinnae commonly 2 cm or more wide: 31. Sterile pinnae entire, to 13 x 4cm 31. Sterile pinnae crenate, to 11 x 2.5cm 29. Sporangia lacking setae. 32. Indusia firm, almost circular, with hairs in the middle 37. P. menisciicarpon .31. P. granulosum . 38. P. hewittii 32. Indusia thin, shrivelled when dry, hairy throughout. 33. Sterile pinnae crenate, at least distally 33. Sterile pinnae entire 26. Indusia small, with dark red cell- walls; “marginal hairs few | or none. 34. Fertile pinnae 1.5 cm or more wide 34. Fertile pinnae not more than | cm wide. 39. P. affine 37. P. menisciicarpon 40. P. borneense 35. Sterile pinnae not or little more than 1 cm wide, fertile 3-6 mm wide. 36. Pinnae thick; lower surface glabrous with thick pale prominent veins 41. P. exsculptum 36. Pinnae thin; hairs present on lower surface of costae; veins concolorous, not prominent. 37. Pinnae to 15 pairs, sterile ones 4-6 cm long, narrowly acuminate 37. Pinnae not over 10 pairs, sterile ones shorter, abruptly pointed 35. Sterile pinnae 1.5 cm or more wide, fertile 5-10 mm wide 15. Pronephrium millarae HOLTTUM, Blumea 20 (1972) 115.—Type: WOMERSLEY & MILLAR NGF 8500, N.E. New Guinea, Morobe Distr., Wau-Salamaua Road 1600 m (LAE). Caudex short-creeping; fronds dimorphous. Sterile fronds. Stipe 7-8 cm long, covered densely with hairs 0.1-0.2 mm long. Lamina 14cm long; pinnae 7-8 pairs, all with stalks 1-1.5 mm long; basal pinnae slightly reduced and deflexed, basis- copic base rounded, acroscopic truncate and slightly auricled. Largest pinnae 2.5-3.5 x 1cm; base subtruncate, apex rounded to bluntly poin- ted, edges slightly crenate near base, more dis- tinctly so distally; costules 2.5-3.5mm apart, at little more than 45° to costa; veins slender and slightly prominent beneath, 3-4 pairs, 12 pairs anastomosing, sinus-membrane not distinct; lower surface of rachis and costa bearing stiff erect hairs 0.2-0.4 mm long, hairs on costules and veins shorter and somewhat antrorse, on surface be- . 42. P. merrillii . 43. P. rhombeum 44. P. peltatum tween veins many appressed hairs 0.1—0.2 mm long, no glands; upper surface throughout bearing stiff suberect hairs 0.2-0.3 mm long. Fertile fronds. Pinnae 2.0-2.3cm long, 0.8cm wide, lower ones rather widely spaced, shape and pubescence as sterile; sori exindusiate, occupying whole of basal veins and sometimes the base of an excurrent vein, on middle part of distal veins; sporangia sometimes with a gland, not setiferous. Distr. Malesia: Eastern New Guinea, Goodenough Island. Ecol. On Goodenough Island moist gully” (BRASS 24636). “common in a 16. Pronephrium peramelense HOLTTUM, Blumea 20 (1972) 115. — Type: PULLE 415, Western New Guinea, Perameles bivouac, 1000 m (BM; L). Caudex horizontal, 2-2.5 mm diameter, bearing stipes 5mm or more apart; fronds not dimor- phous. Stipe 10-20cm long, dark, short-hairy; 1981] THELYPTERIDACEAE (Holttum) oye) basal scales 3-4 mm long, narrow, rather firm with few hairs. Lamina to iScm long; apex broadly triangular, pinnae to 8 pairs, basal ones a little deflexed with somewhat asymmetric and some- times slightly auricled base. Middle pinnae to 3.0 x 1.3cm; base truncate, apex rather abruptly nar- rowed to a rounded tip, edges irregularly slightly crenate to subentire; costules to 4 mm apart; veins 2 pairs, 15 pairs anastomosing; sinus-membrane not evident; lower surface of rachis covered with dense spreading dark rigid curved hairs 0.3 mm long, rest glabrous; upper surface of costa hairy near base only, rest of surface glabrous. Sori inframedial, on all veins, not elongate, exindusiate; sporangia without glands or setae. Distr. Malesia: Western New Guinea; only known from the type. 17. Pronephrium moniliforme (TAGAWA & K. IWATS.) HOLTTUM, Blumea 20 (1972) 115.— Dimorphopteris moniliformis TAGAWA & K. IwaTs. Acta Phytotax. Geobot. 19 (1961) 8, f. 14-16. — Thelypteris moniliformis (TAGAWA & K. Iwats.) K. Iwats. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 36.— Type: HARADA s.n. 30 June 1944, Halmahera (KYO). Caudex short-creeping, 3-3.5mm_ diameter; fronds dimorphous. Sterile fronds. Stipe 7-10 cm long, pale, minutely hairy; lamina 15-20 cm long; pinnae c. 15 pairs; basal pinnae deflexed, not reduced; middle pinnae 41cm, base truncate with acroscopic auricle, apex acute, edges ser- rulate; costules 2.5mm apart; veins 4-5 pairs, 2 pairs anastomosing; lower surface minutely hairy. Fertile frond. Stipe 20-24cm long; lamina 20- 24 cm long; pinnae more than 20 pairs, 4cm long, 2mm _ wide, entirely covered beneath with sporangia (which spread to the surface between veins); veins forming a single row of narrow areoles with a short free vein in each crenature of margin; sporangia bearing glands; spores with longitudinal wing and cross-wings. Distr. Malesia: Moluccas (Halmahera; Batjan: ALSTON 16908, BM). 18. Pronephrium amboinense (WILLD.) HOLT- TUM, Blumea 20 (1972) 120.— Aspidium am- boinense WILLD. Sp. Pl. ed. 4, 5 (1810) 228. — Thelypteris amboinensis (WILLD.) REED, Phy- tologia 17 (1968) 259. — Type: VENTENAT, Am- boina (B, Herb. Willd. 19751). Aspidium canescens forma nephrodiiformis CHRIST, Ann. Jard. Bot. Btzg 15 (1898) 131, quoad SARASIN 975, Celebes, Palopo (BAS). Dryopteris subconformis C. CHR. Bot. Jahrb. 66 (1933) 47.—Syntypes: KJELLBERG 1999, Celebes, Lamasie, Palopo (BO); KJELLBERG 1360, S.E. Celebes, Labibia (BO). Pronephrium palopense HOLTTUM, Blumea 20 (1972) 116.—Type: KJELLBERG 1999, Celebes (BO). — Fig. 15a. Caudex short, suberect; fronds slightly dimor- phous. Stipe 20-40cm long (longest on fertile fronds), minutely hairy; lamina c. 20cm long, apex acuminate, deeply lobed with gradual tran- sition to pinnae; free pinnae 10-15 pairs, basal pair not reduced, narrowed to base both sides. Middle pinnae c. 5.0X1.3cm (sterile), to 6.0 1.1cm (fertile); base truncate and slightly auricled on acroscopic side, apex short-acuminate (more ab- ruptly pointed on pinnae of smaller fronds), edges crenate to a depth of 1-2 mm; costules to 3mm apart; veins 4-5 pairs, I-13 pairs anastomosing, >] pairs to sides of short sinus-membrane; lower surface of rachis, costae and costules bearing rather sparse minute hairs, surface between veins slightly pustular; hairs on upper surface of costae less than 0.5mm, sparse and shorter hairs on costules, a few minute hairs sometimes between veins near margin. Sori medial; indusia small, short-hairy (in some cases absent?); sporangia with several setae; spores pale with translucent wing and a few cross-wings. Distr. Malesia: S.E. & Central Moluccas (Amboina). Ecol. At low altitudes, in forest, on stream- bank (KJELLBERG 1360). Notes. Four collections known. The type from Amboina is a small frond (pinnae to 3.0 x 0.8 cm) of which the sori are in poor condition; a few sporangia are certainly setiferous but no indusia are present. SARASIN 975 from Palopo agrees closely with the type in shape of frond and pinnae (to 4.5 1.2cm) and certainly has small indusia. One of KJELLBERG’s specimens on_ which CHRISTENSEN based Dryopteris subconformis (n. 1999), also from Palopo, is somewhat larger than SARASIN’S and apparently lacks indusia; KJELL- BERG’S second specimen (n. 1360) from S.E. Celebes has indusia. In 1972 I confusedly cited KJELLBERG 1999 both as type of D. subcon- formis and of Pronephrium palopensis; I had in- tended to cite 1360 as type of the former. BLUME misinterpreted Aspidium amboinense WILLD. and described under that name small specimens of Christella subpubescens which had many club-shaped glands on them. This error was copied by later authors and all subsequent cita- tions of WILLDENOW’s species are erroneous. Celebes, 19. Pronephrium firmulum (BAK.) HOLTTUM, Blumea 20 (1972) 116.— Polypodium firmulum BAK. Kew Bull. (1893) 211. — Dryopteris firmula (BAK.) C. Cue. Ind. Fil. (1905) 266; Gard. Bull. Str. Settl. 7 (1934) 249.— Phegopteris firmula (BAK.) v.A.v.R. Handb. (1908) 501.— Type: C. HOSE 295, Sarawak, Mt Dulit (K). Caudex long-creeping, 3-5 mm diameter, bear- ing fronds 1-1.5cm apart; fronds not or little dimorphous. Stipe to 45 cm long, glabrous; lamina to 35cm long, apical section subentire with widened base, small on largest fronds, pinnae to 520 FLORA MALESIANA [ser. II, vol. 1° 12 pairs, of firm texture; basal pinnae narrowed both sides towards the base, acroscopic base sometimes with a slight auricle. Middle pinnae from Mt Dulit to 11 x 2.5 cm, from Mt Kinabalu to 15 x 3.3cm, base subtruncate and distinctly auri- cled (more on fertile than sterile pinnae), apex acuminate, edges irregularly slightly sinuous; cos- tules 3.5—4.5 mm apart, at less than 60°; veins 5S—7 pairs, prominent on both surfaces, almost all anas- tomosing with continuous zig-zag excurrent veins; lower surface of rachis covered with thick curved hairs barely 0.5mm long, costae of Dulit speci- mens bearing sparse short antrorse (not ap- pressed) hairs, many such hairs on Kinabalu spe- cimens, rest glabrous, surface between veins pus- tular; upper surface of rachis as lower, of costae bearing rather sparse hairs less than 0.5 mm long, no others. Sort somewhat inframedial; indusia none or very small, glabrous; sporangia many, with rather small red or yellow glands, no setae; spores not seen. Distr. Malesia: Borneo (Sarawak; Sabah). Ecol. On rocks by streams in forest at 250+ 1000 m. Note. Kinabalu specimens have broader and somewhat thinner pinnae than those from Dulit; indusia have only been seen on some sori of the former. 20. Pronephrium aquatiloides (COPEL.) HOLT- TUM, Blumea 20 (1972) 108.— Dryopteris aqua- tiloides COPEL. Philip. J. Sci. 7 (1912) Bot. 59; v.A.v.R. Handb. Suppl. (1917) 177. — Thelypteris aquatiloides (COPEL.) REED, Phytologia 17 (1968) 260. — Type: BROOKS 9, Sarawak, Bongo Range (MICH; BM, L). Cyclosorus jacobsii HOLTTUM, Blumea 11 (1962) 530.— Thelypteris jacobsii (HOLTTUM) REED, Phytologia 17 (1968) 285. — Type: JACOBS 5086, Sarawak, Mt Penrissen (L; K). Caudex long-creeping, 2-3 mm diameter; fronds c. 5mm apart. Stipe 15-30cm long, minutely hairy; lamina to 27cm long, apex pinna-like, pin- nae 6-8 pairs; basal pinnae narrowed towards base both sides, with stalks to 4mm long. Middle pin- nae 9-11x1.0cm; base auricled on acroscopic side; apex acuminate with rounded tip 2 mm wide; edges almost entire; costules 3mm apart; veins 3-4 pairs, anastomosing to form a zig-zag excur- rent vein, slightly prominent; hairs on lower sur- face of rachis thick, brown, curved, 0.5 mm long, on costae same length, slender, antrorsely ap- pressed, no others hairs; surface between veins pustular; upper surface of rachis hairy as lower, costal hairs shorter, no others hairs. Sori medial, basal ones sometimes coalescent; indusia very small, glabrous or glandular; sporangia bearing red glands; spores not seen. Distr. Malesia: Borneo (W. Sarawak). Ecol. On rocks by streams in forest at 300- 1000 m. 21. Pronephrium simillimum (C. CHR.) HOLT- TUM, Blumea 20 (1972) 116.— Dryopteris simil- lima C. CHR. Ind. Fil. (1905) 292, new name for Nephrodium simulans BAK. J. Bot. 26 (1888) 325, non BAK. 1874; v.A.v.R. Handb. (1908) 223. — Thelypteris simillima (C. CHR.) K. IwaTs. Acta Phytotax. Geobot. 21 (1965) 169.— Type: G. F. HOSE 231, Sarawak, limestone hills (K). Caudex short-creeping to suberect; fronds hardly dimorphous. Stipe 12-25(—45)cm_ long, minutely hairy, basal scales dark, narrow, rigid, to 8mm long with stiff pale hairs 0.2mm _ long; lamina very firm, 25-40 cm long, apex deltoid and deeply lobed, pinnae 12-16 pairs; basal pinnae slightly reduced, narrowed to base on basiscopic side. Largest pinnae of type 4.7 x 1.6cm (width above base), largest seen 6.0 x 1.7 cm; base trun- cate and distinctly auricled on acroscopic side; apex abruptly narrowed, rounded or broadly pointed; edges lobed to depth of 1.5-2mm (on young plants almost entire), lobes oblique, roun- ded, more or less distinctly dentate at ends of veins; costules to 4mm apart, at c. 50° to costa; veins 5-6 pairs, slightly prominent both sides, 13-2 pairs anastomosing, | pair to sides of sinus-mem- brane; lower surface of rachis covered with stiffly erect pale hairs 0.2-0.4mm long, similar hairs 0.1-0.2 mm long on costae, costules, veins and on surface between veins; upper surface similarly hairy, hairs between veins more dense than on lower surface. Sori small, medial, exindusiate; sporangia with 6-8 slender setae; spores with rather broad finely erose translucent wing and cross-wings. Distr. Malesia: Borneo; S.W. Celebes (BROOKS 16853, Maros). Ecol. On limestone, 0-1700 m. Notes. One specimen from a shaded place at entrance to a cave has thinner pinnae more deeply lobed than above described, to 2.2 cm wide, cos- tules 5mm apart, pinna-lobes distinctly dentate at all vein-ends, veins often forked, only 1 pair anastomosing. A specimen from 1700m on G. Mulu is small, with 5 pairs of free pinnae, largest 3.0 x 1.3. cm, hairs on lower surface of rachis and costae more than 0.5 mm long. 22. Pronephrium giluwense HOLTTUM, sp. nov. Pinnae paucijugatae, inferiores leviter redactae auriculatae stipitataeque, pinnae majores 5 cm longae, 1.5 cm latae, vix dimorphae; pagina inter venas utraque pilis brevibus vestita; sori exin- dusiati, sporangia setis 3-6 praedita.— Type: WOODHAMS 138, Papua, Mt Giluwe, 2400 m, cult. Hort. Kew. (K). Caudex short-creeping; stipe to 14cm long, short-hairy; lamina thin, to 16 cm long consisting of apical section 7x2.4cm lobed as pinnae and 3 pairs of pinnae; basal pinnae 2.5 x 1.4cm, rather strongly auricled, with stalks 1.5 mm long. Largest pinnae 5.0 x 1.5 cm with stalks 1 mm; base broadly 1981] THELYPTERIDACEAE (Holttum) 324 cuneate and slightly auricled on acroscopic side, apex abruptly short-acuminate; edges lobed 1/3 towards costa; costules 4mm apart at 45° to costa; veins slender, slightly prominent, 5-6 pairs, | pair anastomosing, 1-1} pairs to sides of sinus-mem- brane; lower surface of rachis and costae bearing spreading hairs of varied length to 0.8 mm, shorter slender hairs on costules, veins and surface be- tween them; upper surface covered with suberect hairs 0.2-0.3mm long, longer hairs present on costae and scattered on costules and veins. Sori medial, lowest supramedial, exindusiate; sporan- gia bearing 3-6 rather long setae; spores with a longitudinal wing and cross-wings. Distr. Malesia: Papua New Guinea. The only other collection is CLEMENS 12110 D, from a branch of the Buso River, Morobe District, at 1500-1800 m (MICH); this is a less mature plant with longer apical lamina than the type; it differs also in having a few glands on the lower surface of pinnae. Note. The type plant perhaps did not attain its full size before dying. Possibly this species should be transferred to Sphaerostephanos but it does not match any known species in that genus. 23. Pronephrium kjellbergii HoLTrumM, Blumea 20 (1972) 117.—KJELLBERG 2638, Central Celebes, Mt Porema 1400 m (BO). Caudex short-creeping; fronds subdimorphous. Stipe of sterile frond 3-4 cm long, of fertile 10cm, short-hairy, basal scales 4x 1mm bearing short hairs; lamina 8cm long, thin; pinnae 7 pairs, lower sterile ones sometimes a little reduced, fer- tile not; apical lamina deltoid and deeply lobed. Largest pinnae 1.5<0.6cm (sterile) 1.0 x 0.6cm (fertile); base truncate and a little dilated on acroscopic side; apex blunt; edges crenate; cos- tules 2.5mm apart; veins 2 pairs (sometimes 3 pairs in basal acroscopic lobe), all free; lower surface of rachis, costae and costules bearing sparse hairs 0.5mm long, many glands on and between veins; upper surface throughout bearing short antrorse hairs with longer ones scattered on costules and veins. Sori near bases of veins; indusia bearing a few hairs and glands; sporangia some- times with a gland. Distr. Malesia: Central Celebes, 1400 m. Only known from the type. Note. This specimen was named Dryopteris urdanetensis COPEL. by CHRISTENSEN (Bot. Jahrb. 66, 1933, 48) but differs from that species in having a much shorter apical lamina and no much- reduced basal pinnae. 24. Pronephrium amphitrichum HOLTTUM, sp. nov. P. clemensiae (Copel.) Holttum affinis, differt: pinnis multo minoribus; pilis costarum costu- larumque subtus patentibus, pilis inter venas paginae superioris non appressis; soris venarum inferiorum elongatis. —Type: EDANO BS 75790, Luzon, Camarines Sur, Mt Potianay (MICH; NY). Caudex suberect; fronds subdimorphous. Stipe 3 cm long (sterile) 6-7 cm (fertile), short-hairy, basal scales 3 mm long, narrow; lamina 5 cm long; pinnae 4-5 pairs; basal pinnae slightly reduced, much narrowed towards base on basiscopic side. Largest pinnae to 1.8 x 0.9 cm (sterile) 1.5 x 0.8 cm (fertile); base truncate and slightly auricled on acroscopic side, narrowed and rounded on basiscopic; apex abruptly blunt-pointed; edges crenate to depth of 1mm; costules to 3mm apart at little over 45° to costa; veins 3 pairs, | pair anastomosing; lower surface of rachis bearing thick pale spreading hairs 0.5 mm long, hairs on costae and costules similar but shorter, abundant slender erect hairs and a few glands on surface between veins; hairs on upper surface of rachis thick, curved, to | mm long, on costae 0.2-0.3 mm, on costules and veins scattered longer hairs, between veins copious suberect hairs 0.2 mm long. Sori inframedial to medial, lower ones elongate; indusium small with short stiff hairs; sporangia sometimes with a gland. Distr. Malesia: Philippines known from the type. (Luzon). Only 25. Pronephrium clemensiae (COPEL.) HOLTTUM, Blumea 20 (1972) 118.— Dryopteris clemensiae CoPEL. Philip. J. Sci. 46 (1931) 213. — Cyclosorus clemensiae (COPEL.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 243; COPEL. Fern FI. Philip. (1960) 369.— Thelypteris clemensiae (COPEL.) REED, Phytologia 17 (1968) 268.— Type: CLEMENS 16490, Luzon, Isabella Prov., Mt Moises (UC; US). a. var. clemensiae Caudex short-creeping; fronds subdimorphous. Stipe 8-10cm long (sterile), 16cm (fertile), hairy as rachis distally, basal scales to 3 x 1 mm, firm; lamina 13cm long, texture firm, apex broadly triangular and deeply lobed, pinnae 5 pairs, basal pinnae not reduced, narrowed towards base of basiscopic side. Largest pinnae 4.3 1.8cm (sterile) 3.2x1.5cm (fertile); base subequally broadly cuneate; apex abruptly narrowed to acute tip; edges lobed to depth of 2mm; costules 3.5— 4mm apart, at 45° to costa; veins 4-5 pairs, prom- inent both sides, 1-1; pairs anastomosing; lower surface of rachis covered with coarse erect hairs, hairs on costae and costules shorter and antror- sely appressed, glands abundant on lower surface generally; hairs on upper surface of costae, cos- tules and veins antrorse as lower surface, between veins rather sparse appressed hairs. Sori medial; indusia rigid, dark, with yellow glands and some- times a few hairs; sporangia often with a yellow gland. Distr. Malesia: known from the type. Philippines (Luzon); only 522 b. var. degenerum (CHRIST) HOLTTUM, comb. nov.— Dryopteris canescens var. degenera CHRIST, Philip. J. Sci. 2 (1907) Bot. 199. — Phe- gopteris canescens var. degenera (CHRIST) v.A.v.R. Handb. (1908) 507. — Cyclosorus degener (CHRIST) CopeEL. Fern FI. Philip. (1960) 356. — Thelypteris degenera (CHRIST) REED, Phytologia 17 (1968) 271. — Lectotype (HOLTTUM 1972): LOHER s.n. March 1906, Luzon, Montalban (P). Differs from var. clemensiae: indusia densely setose; sporangia bearing glands or setae. Spores have a complete translucent wing. Distr. Malesia: Philippines, Luzon (C. B. ROBINSON BS 9455, Tayabas Prov. Infanta; RAMOS BS 1791, Rizal Prov.; M. G. PRICE 2824, Zambales Prov., Mts above Palauig at 1000 m). Notes. PRICE’s specimen is much larger than the others (pinnae to 8.5 x 2.0cm) and has rather sparse glands on lower surface but agrees in other respects. I designate the LOHER collection from Montalban as lectotype because I did not find the other syntype, also collected by LOHER, at Paris. 26. Pronephrium glandulosum (BL.) HOLTTUM, Blumea 20 (1972) 118.— Aspidium glandulosum BL. En. Pl. Jav. (1828) 144; Metr. Farngatt. IV (1858) 111, quoad descr. tantum.— Nephrodium glandulosum (BL.) J. SM. in Hook. J. Bot. 3 (1841) 411, nomen tantum; PRESL, Epim. Bot. (1851) 45, excl. pl. Zoll.; HOOK. Spec. Fil. 4 (1862) 76, excl. syn. omn. praeter BI.; RActiB. Fl. Btzg 1 (1898) 185.— Abacopteris glandulosa (BL.) FEE, Gen. Fil. (1852) 310. — Dryopteris glandulosa (BL.) O. KTZE, Rev. Gen. Pl. 2 (1891) 812; v.A.v.R. Handb. (1908) 215.— Dryopteris malayensis C. CHR. Dansk Vid. Selsk. Skr. VII, 10 (1913) 171, nom. nov. illegit.; BACKER & POSTH. Varenfl. Java (1939) 59.—Cyclosorus glandulosus (BL.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 227; HOLTTUM, Rev. FI. Malaya 2 (1955) 278, f. 160. — Thelypteris malayensis (C. CHR.) REED, Phytologia 17 (1968) 291.—Type: BLUME s.n., W. Java (L, n. 908, 337-89). Dryopteris iridescens v.A.v.R. Bull. Jard. Bot. Btzg II, 11 (1913) 11; Handb. Suppl. (1917) 176. — Thelypteris iridescens (v.A.v.R.) REED, Phy- tologia 17 (1968) 285.—Type: MATTHEW S517, Sumatra, Padang Panjang (BO). Dryopteris excrescens COPEL. Univ. Cal. Publ. Bot. 14 (1929) 374. — Type: RAHMAT SI TOROES 142, Sumatra (UC; L). Dryopteris bartlettii COPEL. ibid. l.c.— Thelypteris bartlettii (COPEL.) REED, Phytologia 17 (1968) 263. — Type: BARTLETT 6692, Sumatra, Asahan (UC). Nephrodium lineatum sensu. BEDD. Handb. Suppl. (1892) 71, excl. syn. Aspidium affine BL. Caudex short-creeping; fronds dimorphous. Sterile fronds. Stipe 20-30cm long, minutely hairy; lamina 30-40cm long; pinnae 12 pairs; basal pinnae narrowed at base, more on basiscopic FLORA MALESIANA [ser. II, vol. 1° than acroscopic side, with stalks 2mm long. Lar- gest pinnae commonly to 12 x 2.5 cm; base trun- cate, auricled on acroscopic side, apex abruptly short-acuminate; edges obliquely lobed to a depth of 1-2 mm or sometimes more deeply; costules to 5 mm apart; veins to 8 pairs, 2-4 pairs anastomos- ing (according to depth of lobing), 15-2 pairs to sides of sinus-membrane; lower surface of rachis covered with thick curved hairs more than 0.5 mm long, similar but shorter hairs on costae, very short and sparse on costules and veins, surface between veins pustular, bearing many glands and sometimes very short erect hairs; upper surface covered throughout more or less closely with fine appressed hairs 0.2-0.3mm long. Fertile fronds. Stipe to 50cm long; pinnae more widely spaced than sterile, commonly to 7 x 1.5 cm, edges shal- lowly crenate; sori medial, lower ones at least somewhat elongate along veins; indusia thin bear- ing many very short hairs; sporangia with glands on body and a hair with glandular tip on stalk; spores with a complete wing and cross-wings. Distr. Malesia: Java, Sumatra, Malaya. Ecol. Lowland, up to 800m, in forest near streams. Notes. The lobing of pinnae is variable. One Java specimen at Kew (coll. LOBB) has both ster- ile and fertile pinnae almost entire. A specimen collected by KURZ on G. Salak (PR) has fertile pinnae 7 x 1.7 cm, lobed fully half-way to costa. The name Aspidium glandulosum BL. was variously misinterpreted by J. SMITH, KUNZE and HOOKER, who cited specimens of other spe- cies; these citations were copied by METTENIUS and PRESL who both gave good descriptions. Apart from the presence of copious glands on lower surface and appressed hairs on the upper, this species is very similar to A. affine BL., with which BEDDOME confused it under the name Nephrodium lineatum. 27. Pronephrium debile (BAK.) HOLTTUM, Blu- mea 20 (1972) 118. — Nephrodium debile BAK. J. Bot. 18 (1880) 212. — Dryopteris pseudoreptans C. Cure. Ind. Fil. (1905) 286, nom. nov. (not D. debilis (METT.) C. CHR.); v.A.v.R. Handb. (1908) 223. — Thelypteris pseudoreptans (C. CHR.) REED, Phy- tologia 17 (1968) 306.— Type: BECCARI 433, Sumatra, G. Singgalang 1700 m (K). Nephrodium pilosiusculum RAciB. FI. Btzg 1 (1898) 189. — Dryopteris pilosiuscula (RACIB.) C. Cur. Ind. Fil. (1905) 284; v.A.v.R. Handb. (1908) 215; BACKER & POSTH. Varenfl. Java (1939) 65, excl. syn. Aspidium pilosiusculum METT. — Thelypteris pilosiuscula (RACIB.) REED, Phy- tologia 17 (1968) 304. — Type: RACIBORSKI, Java, G. Salak, Tjiapoes (BO; K, L, P). Caudex short-creeping; fronds slightly dimor- phous. Stipe 15-20 cm long, glabrous except base which is covered with minute hairs and firm seti- ferous scales 2x 1mm; lamina 15-25cm long, 1981] THELYPTERIDACEAE (Holttum) 323 firm in texture; apical lamina narrowly acuminate; pinnae 10-15 pairs; basal pinnae slightly narrowed at base on basiscopic side, sessile. Largest pinnae 4.5x1.5cm (sterile) 3.0 1.0cm (fertile); base truncate and slightly auricled; apex abruptly short- acuminate with rounded tip; edges crenate or in the largest sterile pinnae lobed to a depth of 2-3 mm; costules 3-4 mm apart; veins 3-4 pairs, | pair anastomosing, next pair to sides of sinus- membrane; lower surface of rachis covered with thick spreading brown hairs 0.7 mm or more long and shorter pale ones, hairs on costae and costules dense, pale, 0.4mm long, somewhat antrorse, hairs on veins shorter with some yellow glands, surface between veins bearing many short erect hairs and a few glands (especially distal on the lobes); upper surface of rachis bearing erect thick brown hairs Imm long, pinnae covered closely with appressed hairs 0.5mm long. Sori medial; indusia covered with short hairs, 1-2 glands also sometimes present; sporangia bearing glands. Distr. Malesia: Sumatra, Java, Flores. Ecol. On wet rocks by streams at 1400-1800 m. Note. BECCARI’s specimen from G. Singgalang has more deeply lobed pinnae than RACIBORSKI’s from Tjiapoes but differs little in other respects. RACIBORSKI cited a specimen of ZIPPELIUS (to whom he attributed the name) as well as his own; I have not found the former and take RACIBOR- SKI’s own, which he described, to be the type. Confusion has been caused because METTENIUS independently published the name Aspidium pilosiusculum Zipp. as a new name for Gym- nogramme appendiculata BL. (here transferred to Sphaerostephanos). Both species were collected by RACIBORSKI at Tjiapoes (Mt Salak, W. Java). 28. Pronephrium minahassae HOLTTUM, Blumea 20 (1972) 119.—Type: ALSTON 16410, N. Celebes, G. Tetawiran (BM). Caudex short, suberect; fronds dimorphous. Sterile fronds. Stipe to 15 cm long, minutely hairy, basal scales small. Lamina to 25 cm long, texture thin; apical lamina pinna-like, to 10cm long; pin- nae to 10 pairs, basal pair more remote and nar- rowed to base on basiscopic side, widest in middle. Largest pinnae 4.5cm long, 1.5cm wide above base; base truncate, slightly auricled espe- cially on lower pinnae; apex abruptly short-poin- ted, edges lobed to a depth of 1.5-2 mm; costules 3.5-4mm apart; veins 6-7 pairs, 1) pairs anas- tomosing, | pair to sides of sinus-membrane; lower surface of rachis glabrous or sparsely hairy, of costae, costules and veins covered with slender appressed hairs, glands abundant throughout; upper surface covered with slender appressed hairs. Fertile fronds. Pinnae 2.8 cm long, 0.7-0.9 cm wide above base, lobed as sterile and similarly hairy and glandular; sori at maturity covering whole surface; indusia hairy; sporangia bearing glands; spores with a translucent wing and cross-wings. Distr. Malesia: N. Celebes; only known from the type. 29. Pronephrium solsonicum HOLTTUM, Kalik- asan 3 (1974) 197.—Type: M. G. PRICE 2903, Luzon, Ilocos Norte, Solsona, 1100 m, in forest (K; PNH). Caudex short-creeping; fronds dimorphous. Sterile fronds. Stipe 5-6 cm long, short-hairy near base, basal scales thin, 3mm long. Lamina to 15 cm long; apex 8 cm long, deeply lobed; pinnae 9 pairs, basal pair slightly reduced and narrowed to base on basiscopic side. Largest pinnae 2.1 0.8 cm; base truncate and slightly auricled; apex obtuse; edges lobed to a depth of 1 mm; costules 2.5mm apart; veins slender, 3 pairs, basal pair anastomosing, next pair to edge; lower surface of rachis bearing coarse spreading pale hairs | mm long, hairs on costae and costules much shorter, slender, appressed, many glands on surface be- tween veins; upper surface of rachis as lower, hairs on costae 0.5mm long, similar hairs scat- tered on costules and veins, no others. Fertile fronds. Stipe 12-20cm long; pinnae to 1.4 0.7 cm, edges subentire or slightly crenate; sori near costules, indusia large, glabrous; sporangia bearing glands. Distr. Malesia: known from the type. Philippines (Luzon); only 30. Pronephrium celebicum (BAK.) HOLTTUM, Blumea 20 (1972) 119.— Acrostichum celebicum BAK. Kew Bull. (1901) 741.— Leptochilus cele- bicus (BAK.) C. CHR. Ind. Fil. (1905) 384; v.A.v.R. Handb. (1908) 741. — Dryopteris celebica (BAK.) COPEL. Philip. J. Sci. 37 (1928) 410.— Thelypteris celebica (BAK.) REED, Phytologia 17 (1968) 267. — Lectotype (HOLTTUM 1972): DE LA SAVINIERRE 61, N. Celebes (K; P). Aspidium canescens forma _ acrostichoides CHRIST, Ann. Jard. Bot. Btzg 15 (1898) 132.— Dryopteris acrostichoides (CHRIST) v.A.v.R. Handb. Suppl. (1917) Corr. 49, non O. KTZE 1891 excl. var. rhombea & var. lanceola. — Lectotype (HOLTTUM 1972): KOORDERS 17153, N. Celebes (BOs): (2?) Meniscium hosei var. sumbensis v.A.v.R. Bull. Dép. Agr. Ind. Néerl. 21 (1908) 7. — Phegop- teris hosei var. sumbensis v.A.v.R. Handb. (1908) 510.— Type: TEYSMANN 10693, Sumba (BO?; not seen). — Fig. 15b-—d. Caudex short-creeping; fronds dimorphous. Sterile fronds. Stipe 6-15 cm long, pale, minutely hairy or glabrescent, basal scales thin, narrow, 3mm long. Lamina thin, 15-18cm long; apex acuminate and deeply lobed; pinnae 8-10 pairs, basal pinnae narrowed to base on basiscopic side, sessile or with a very short stalk. Largest pinnae 4.2 1.2cm (width above base); base truncate, strongly auricled on acroscopic side; apex short- acuminate; edges serrate-crenate to a depth of 524 FLORA MALESIANA 1-1.5mm; costules 2.5mm apart; veins slender and little prominent, 4-5 pairs, 2 pairs anastomos- ing, sinus-membrane hardly evident; lower surface of rachis bearing hairs 0.3 mm long, pale, sometimes sparse, on costae and costules hairs much shorter and always sparse, surfaces between veins pus- tular; hairs on upper surface of rachis thick, to 0.5 mm long, on costae 0.2 mm, rather sparse, no others. Fertile fronds. Stipe 20-30 cm long; lamina to 25cm; pinnae widely spaced, 2.5-4.0 cm long, 2-4 mm wide above auricled base, edges crenate; veins 3 pairs, basal pair only anastomosing; lower surface covered with sori (also upper surface near apex?); indusia very small with a few hairs; sporangia of type bearing both glands and short setae, of Curtis specimen (also cited by Baker) glands only; spores light brown, with a translucent wing and cross-wings. Distr. Malesia: Celebes, Moluccas (Ceram, Tenimber Is.), West New Guinea (Vogelkop), and ? Lesser Sunda Is. (Sumba). Notes. SARASIN 1022, from Central Celebes, cited by CHRIST with those of KOORDERS, has sterile pinnae to 6 x 1.5 cm and sporangia strongly setiferous. v.A.v.R.’s description of D. acrosti- choides was based on KOORDERS’S specimens at Bogor named by CHRIST; these were not cited by CHRIST in 1898; I have selected KOORDERS 17153 as the type of v.A.v.R.’s name. 31. Pronephrium granulosum (PRESL) HOLTTUM, Blumea 20 (1972) 119. — Polypodium granulosum PRESL, Rel. Haenk. (1825) 24, pl. 4, f. 2; HOLT- TUM, Novit. Bot. Inst. Bot. Univ. Carol. Prag. 1968 (1969) 19. — Dryopteris granulosa (PRESL) C. Cure. Ind. Fil. (1905) 269; CHRIST, Philip. J. Sci. 2 (1907) Bot. 217. — Phegopteris granulosa (PRESL) v.A.v.R. Handb. (1908) 503, p.p.— Cyclosorus granulosus (PRESL) COPEL. Fern FI. Philip. (1960) 373, p.p. — Thelypteris granulosa (PRESL) REED, Phytologia 17 (1968) 280.—Type: HAENKE, Luzon (PRC; specimen at W is marked HAENKE 98). Dryopteris chamaeotaria CHRIST, Philip. J. Sci. 2 (1907) Bot. 203.— Phegopteris chamaeotaria (CHRIST) v.A.v.R. Handb. (1908) 505.— Thelyp- teris chamaeotaria (CHRIST) REED, Phytologia 17 (1968) 267.—Lectotype (COPELAND 1960): WHITFORD 1369, Luzon, Mt Mariveles (not found at MICH, P, US); Neotype (HOLTTUM 1972): ELMER 6970, same locality (K). Dryopteris maquilingensis COPEL. Philip. J. Sci. 56 (1935) 103, pl. 8. — Type: COPELAND s.n. Nov. 1932, Luzon, Mt Makiling (formerly in PNH); Neotype (HOLTTUM 1972): ELMER 18169, same locality (K; BISH, G, L). Cyclosorus subdimorphus COPEL. Philip. J. Sci. 81 (1952) 38; Fern Fl. Philip. (1960) 374.— Thelypteris palawanensis REED, Phytologia 17 (1968) 300, nom. nov.—Type: EDANO PNH 13925, Palawan (MICH). — Fig. 15m-n. [ser. II, vol. 1° Caudex short to long-creeping, 3 mm diameter; fronds close or 1l-2cm apart, subdimorphous. Stipe 15-20cm long (sterile) 30-45 cm (fertile); lamina of mature plants 15-24cm long, thin but firm, consisting of apical section 8-15cm long, 1.5-3.0cm wide, and 2-4 pairs of pinnae; lowest pinnae stalked 1-2mm, widest at about the middle, base asymmetric and + auricled. Middle pinnae to 11 x2.5cm (sterile; fertile smaller, to 9x2.2cm); base truncate and slightly auricled; apex acuminate (narrowly on longest pinnae); edges crenate to depth of 1 mm (entire on pinnae of young plants); costules to 4mm part; veins 5-6(—8) pairs, slender and slightly prominent, 2-5 pairs anastomosing (according to width of pinnae); lower surface of rachis bearing a variable number of coarse pale hairs to 0.8mm long; hairs on costae sparse and much shorter, surface between veins pustular and sometimes with very short erect hairs; upper surface of rachis covered with thick pale hairs 1mm long, costal hairs rather sparse, 0.3mm long, no other hairs. Sori inframedial; indusia small, soon caducous, dark red with short hairs on surface and margin; sporangia bearing glands and also short setae; spores with trans- lucent wing and cross-wings. Distr. Malesia: Philippines (Luzon, Palawan, Balabac, Panay, Mindanao). Ecol. In forest at low altitudes, to 500 m. Notes. One frond in HAENKE’s collection at Prague has two pinnae widened and irregularly deeply lobed distally; PRESL named this var. lobata. This feature may indicate hybridity; it needs experimental investigation. Young plants have much smaller pinnae than above described; the smallest fertile pinna seen measures 4.5 1.2cm. A specimen from Balabac (J. B. STEERE) has no glands on sporangia, and pinnae crenate to a depth of 2 mm; otherwise it is very near the type. A specimen from Mindanao has very narrow pinnae, the largest fertile one 5.0 x 0.8 cm. 32. Pronephrium samarense (COPEL.) HOLTTUM, Blumea 20 (1972) 119.—Cyclosorus samarensis CoPEL. Philip. J. Sci. 81 (1952) 35. — Thelypteris samarensis (COPEL.) REED, Phytologia 17 (1968) 311.— Type: GACHALIAN PNH _ 15240, Samar (MICH; 15220 at SING). Caudex short-creeping, 2mm diameter; fronds dimorphous. Sterile fronds. Stipe 3-5 cm long, basal scales to 2mm long, pale stiff hairs 0.7 mm long in the groove. Lamina 5-8cm long; apex narrowly triangular, 2.5cm long, lobed at base only; pinnae to 5 pairs, stalked 1mm, lowest a little reduced and narrowed to base on basiscopic side. Largest pinnae 2-3 cm long, 0.8-1.0 cm wide; base subtruncate, slightly auricled on acroscopic side, rounded on basiscopic; apex obtusely poin- ted to rounded; edges entire; costules 2 mm apart; veins mostly 3 on basiscopic side of costule, 2 on 1981] THELYPTERIDACEAE (Holttum) D2 acroscopic, anastomosing to form one series of areoles with irregular additional ones; hairs on lower surface of rachis 0.3 mm long, on costae and costules shorter and sparse, no other hairs, sur- face between veins pustular; upper surface of rachis and costae similarly hairy, a few minute hairs present near margin. Fertile fronds. Stipe 8-18 cm long; lower pinnae widely spaced, largest pinnae to 1.2x0.5cm, shaped and stalked as sterile; costules simple or forked; where forked the branches anastomosing near margin; sori in one row on each side of costa with additional ones present in basal acroscopic lobe; indusia small, dark red, bearing short hairs; sporangia with 1 or 2 short setae. Distr. Malesia: Philippines (Samar, 3 collec- tions). Ecol. Streambanks at c. 100 m. 33. Pronephrium x xiphioides (CHRIST) HOLT- TUM, Blumea 20 (1972) 119.— Dryopteris xiphioides CHRIST, Philip. J. Sci. 2 (1907) Bot. 201. — Phegopteris xiphioides (CHRIST) v.A.v.R. Handb. (1908) 501.—Cyclosorus xiphioides (CHRIST) COPEL. Gen. Fil. (1947) 143; Fern FI. Philip. (1960) 349.— Thelypteris xiphioides (CHRIST) REED, Phytologia 17 (1968) 324. — Type: COPELAND s.n. April 1905, Mindanao, San Ramon (MICH). Cyclosorus edanyoi COPEL. Philip. J. Sci. 81 (1952) 37; Fern Fl. Philip. (1960) 370. — Thelyp- teris edanyoi (COPEL.) REED, Phytologia 17 (1968) 274. — Type: Edano BS 46065, Panay (UC). Dryopteris diversiloba sensu CHRIST, Philip. J. Sci. 2 (1907) Bot. 199, excl. var. acrostichoides. Dryopteris rhombea sensu COPEL. Philip. J. Sci. 56 (1935) 102, pl. 6, p.p. — Cyclosorus rhom- beus sensu COPEL. Fern FI. Philip. (1960) 357, p.p. Plants with a frond-form intermediate between P. granulosum and P. rhombeum, with pinnae of irregular shape; young plants with lamina 4-5 cm long often with 2-3 pairs of pinnae; adult plants mostly with 4-6 pairs of pinnae (rarely 8 pairs); in almost all cases some or all pinnae, often also the apical lamina of the frond, irregularly dilated and irregularly + deeply lobed distally (see COPELAND 1935, line drawings on pl. 6); at least the lower pinnae + stalked as in P. granulosum; indusia small with a few short hairs; sporangia always setiferous. Distr. Malesia: Philippines, widely. Notes. These plants look like a hybrid swarm, which needs experimental cytotaxonomic in- vestigation. The type collection of P. granulosum includes a frond with one pinna dilated and deeply lobed at the apex, indicating perhaps a slight degree of hybridity. Both parent species have very similar indusia; the sporangia of P. granulosum have glands on sporangia, but the hybrids do not. I have regarded this as an important distinction. The type of D. xiphioides is the only specimen seen in which no pinnae are dilated distally. 34. Pronephrium trachyphyllum HOLTTUM, sp. nov. Stipes frondis fertilis 23cm longus; lamina 14 cm longa pinnis 7-jugatis laminaque 5 cm longa constituta; pinnae usque 3.00.8 cm, crenatae, eglandulosae, supra inter venas pilis minutis suberectis multis vestitae; sori inframediales, non elongati; indusia parva tenuia atrorubra, pilis multis 0.2-0.3 mm longis vestita; sporangia setis pluribus praedita.— Type: A. B. COLINA 772, Mindanao, Surigao (K ex CAHP). Caudex short-creeping, bearing small sterile and much larger fertile fronds. Sterile fronds. Stipe 1.5cm long; lamina 4cm long, apex deltoid and lobed, pinnae 3-4 pairs, to 1.3 x 0.5 cm, subentire. Fertile fronds. Stipe 10-23cm long, minutely hairy; basal scales thin, narrow, to 3mm long. Lamina 8-14cm long; apex 3—4cm long, rather deeply lobed throughout with some _ subapical lobes irregularly elongate; pinnae to 7 pairs; basal pinnae of largest frond not reduced but more widely spaced, not opposite, 3.0cm long, 0.8 cm wide above base, somewhat dilated distally with a few lobes irregularly slightly elongate. Suprabasal pinnae 2.7cm long, 0.8cm wide above truncate and auricled base; apex abruptly obtuse; edges lobed to a depth of 1mm; costules 2.5 mm apart; veins to 4 pairs, 1: pairs anastomosing; lower surface of rachis covered with pale hairs 0.2- 0.3 mm long, costae bearing pale spreading hairs of mixed length to 0.6mm, sparse short hairs on costules and veins and between veins; hairs on upper surface of rachis 0.5 mm, on costae 0.2 mm, very short suberect hairs present on surface be- tween veins. Sori inframedial, not elongate; in- dusia small, thin, dark red, with many stiff pale hairs 0.2-0.3mm; sporangia with several short setae; spores pale with translucent wing and cross-wings. Distr. Malesia: Philippines (Mindanao), only known from type. Notes. It is possible that larger sterile fronds might be produced on other plants. The fertile fronds differ from those of P. amphitrichum (in which the upper surface is hairy in a similar way) in having larger pinnae, sori not elongate and setose sporangia. The smaller fertile fronds of the type have shallowly crenate pinnae, but agree with the largest in having irregular long lobes on the apical lamina. Such irregular lobes are sometimes a sign of hybridity, but there are abundant good spores. 35. Pronephrium thysanoides HOLTTUM, sp. nov. P. simillimo (C. Chr.) Holttum affinis, ab ea differt: frondibus minoribus; pinnis non auri- culatis; rachide costisque subtus glabris; soris indusiatis, indusiis parvis, rigidis, setis brevibus 526 FLORA MALESIANA (ser. Il, voleate ciliatis. — Type: JERMY 14049, Sarawak, Gunong Mulu, Ulu Sungei Air Jernih, north wall (BM). Caudex short-creeping; stipe 15-20cm long, glabrous, basal scales narrow, firm, c. 3mm long with few hairs; fronds of type all fertile. Lamina to 24cm long; pinnae 15 pairs, lower ones only subopposite; apical lamina 4 cm long, deeply lobed at base and grading to upper pinnae; basal pinnae slightly narrowed towards their bases, more on basiscopic than on acroscopic side, not auricled. Largest pinnae 3.5 xX 1.1cm; base truncate; apex abruptly acute, + upcurved; edges crenate to a depth of less than 1 mm; costules 3 mm apart, at c. 60° to costa; veins 4 pairs, slender, | or 13 pairs anastomosing, next vein passing to the short sinus-membrane; lower surface of rachis and cos- tae glabrous, of costules, veins and surface be- tween veins bearing rather sparse very short hairs which may be abraded from old fronds; upper surface of rachis and costae bearing rather sparse pale hairs 0.3mm long, rather sparse very short hairs between veins. Sori medial, small; indusia small but firm, with many short stiff marginal hairs; sporangia with several short setae; spores with many small wings. Distr. Malesia: Borneo (Sarawak: Mt Mulu), only known from the type. Ecol. “In clay in crevice of open rock-face’’, on limestone at 700 m. 36. Pronephrium hosei (BAK.) HOLTTUM, Blumea 20 (1972) 120.— Meniscium hosei BAK. J. Linn. Soc. Bot. 22 (1886) 142.—Dryopteris hosei (BAK)! Gy 1GHrs Inds) Fill (1905) 92712. COREL. Philip. J. Sci. 10 (1915) Bot. 146. — Phegopteris hosei (BAK.) v.A.v.R. Bull. Dép. Agr. Ind. Néerl. 21 (1908) 7; Handb. (1908) 510.— Cyclosorus hosei (BAK.) COPEL. Gen. Fil. (1947) 142; Fern Fl. Philip. (1960) 352. — Thelypteris hosei (BAK.) K. IwatTs. Acta Phytotax. Geobot. 21 (1965) 170. — Type: G. F. HOSE 160, Sarawak (K). Meniscium proliferum sensu. HOOK. 2 Cent. Ferns (1861) t. 15, quoad plantam totam et fig. 2 tantum. Caudex short-creeping; fronds dimorphous. Sterile fronds. Stipe 5-15 cm long, minutely hairy at base, scales c. 2X 1mm, caducous. Lamina 12-15 cm long, firm; apex to 7 cm long, narrowly triangular, lobed near base only; pinnae 6-8 pairs; lower pinnae not or little reduced, in most cases narrowed slightly near base on basiscopic side, edges sometimes crenate. Middle pinnae to Scm long, 0.9cm wide above base; base truncate, al- ways auricled on acroscopic side, often slightly also on basiscopic; distal half of pinna evenly attenuate to narrow rounded apex; edges entire or + crenate distally; costules 2mm apart; veins 4-5 on basiscopic side of costules, 3-4 on acroscopic, to 7 pairs in basal auricle, 2 pairs anastomosing; sinus-membrane not evident; lower surface of tachis bearing curved hairs 0.5 mm long, hairs on costae 0.1-0.2 mm, no other hairs except on mar- gin, surface between veins very pustular; hairs of upper surface of rachis to 1mm long, on costae copious, 0.3mm long, some longer hairs present near edge. Fertile fronds. Stipe to 25cm long; lamina 10-12 cm long; apex 2.5 cm long; pinnae to 9 pairs, to 3.0cm long, 0.4 cm wide above auricled base; pubescence as sterile but hairs on rachis shorter; sori near costules, spreading somewhat long veins; indusia persistent, bearing many hairs 0.1mm long; sporangia with 1-2 short setae; spores pale, with longitudinal wing and cross- wings. Chromosomes: n= 36 (T. G. WALKER). Distr. Malesia: Borneo; Philippines (Min- danao: Zamboanga Prov., SANTOS 4114). Ecol. On rocky stream-banks in forest, at low altitudes. 37. Pronephrium menisciicarpon (BL.) HOLTTUM, Blumea 20 (1972) 111. — Aspidium menisciicarpon BL. En. Pl. Jav. (1828) 142. — Dryopteris menisci- icarpa (BL.) PosTH. Bull. Jard. Bot. Btzg III, 13 (1933) 93; BACKER & POSTH. Varenfl. Java (1939) 61, excl. syn. Dryopteris verruculosa v.A.v.R. — Abacopteris menisciicarpa (BL.) HOLTTUM, Rev. Fl. Malaya 2 (1955) 290, f. 168.— Thelypteris menisciicarpa (BL.) K. IwATs. Acta Phytotax. Geobot. 21 (1965) 171.— Type: BLUME, W. Java (L, n. 908, 333-724). Nephrodium latifolium PRESL, Epim. Bot. (1851) 45, excl. CUMING 16; HOLTTUM, Novit. Bot. Inst. Bot. Univ. Carol. Prag. 1968 (1969) 40. — Cyclosorus latifolius (PRESL) COPEL. Fern Fl. Philip. (1960) 370.— Thelypteris latifolia (PRESL) REED, Phytologia 17 (1968) 287. — Lec- totype (HOLTTUM 1969): CUMING 298, Leyte (PRC). Abacopteris truncata FEE, Gen. Fil. (1852) 310. — Type: CUMING 298 (not seen). Dryopteris holophylla C. CHR. Ind. Fil. (1905) 271, nom nov. for Polypodium holophyllum BAK. J. Bot. 26 (1888) 325, non BAK. 1879. — Phegop- teris holophylla (C. CHR.) v.A.v.R. Handb. (1908) 500. — Type: C. HOSE 242, Sarawak, Niah (K; E). Dryopteris cordifolia v.A.v.R. Bull. Jard. Bot. Btzg II, 11 (1913) 19, pl. 5; Handb. Suppl. (1917) 320. — Type: AMDJAH 322 p.p. N. Borneo (BO; Kee): Dryopteris mirabilis COPEL. Philip. J. Sci. 6 (1911) Bot. 137, pl. 19; v.A.v.R. Handb. Suppl. (1917) 174; C. CHR. Gard. Bull. Str. Settl. 4 (1929) 391; 7 (1934) 249. — Type: BROOKS |6a, Sarawak, Bidi (MICH; BM). Dryopteris _korthalsii ROSENST. Meded. Rijksherb. n. 31 (1917) 5.—Type: KORTHALS, Sumatra (L; K). Dryopteris verruculosa var. sumatrana v.A.v.R. Bull. Jard. Bot. Btzg III, 2 (1920) 151.— Type: Brooks 157/S, Sumatra, Lebong Tandai (BO; BM). Dryopteris urophylla var. peraspera v.A.v.R. 1981] THELYPTERIDACEAE (Holttum) “yal Bull. Jard. Bot. Btzg III, 5 (1922) 303.— Type: LAM 666, W. New Guinea (BO; L). Nephrodium glandulosum sensu BEDD. Handb. Suppl. (1892) 70, p.p. Caudex short-creeping; fronds tufted, dimor- phous. Sterile fronds. Stipe to 20cm_ long, minutely hairy; basal scales thin, broad, soon shrivelling. Lamina 30-40 cm long, consisting of 4-6(-8) pairs of pinnae and an apical lamina which is longest on fronds with few pinnae; young plants in Borneo (D. holophylla) have simple fronds to 25x8cm with cordate base. Pinnae to 13.5 x 4.5cm, usually widest at or above the middle; base truncate with a slight acroscopic auricle (especially on lower ones); apex abruptly short- pointed; edges + crenate distally; costules to 4mm apart; veins to 10 pairs, anastomosing to form a zig-zag excurrent vein; both surfaces of rachis covered with coarse pale hairs, those on costae much shorter, sparse on costules, surface between veins strongly pustular. Fertile fronds. Stipe to 45cm long; pinnae more widely spaced than sterile ones, commonly 7 x 1.5 to 10 x 2.5 cm (on some plants much smaller), shape as sterile but edges almost entire; sori medial to suprame- dial, slightly elongate; indusia with many short hairs; sporangia bearing 0-2 setae; spores with longitudinal wing and cross-wings. Distr. Malesia: Sumatra, Malaya, Borneo, W. Java; Philippines (Samar, Leyte); W. New Guinea, Papua, New Britain. Ecol. In forest near streams at low altitudes. Notes. In Malaya (Pahang and Perak) plants grow on earthen banks of streams; in such places they are swept away by floods before they have attained their full size, but they commonly produce fertile fronds with pinnae to c. 3.0 0.6cm. Young plants with very large simple fronds like the type of Dryopteris holophylla have not been found in Malaya, but the largest adult specimens from Malaya are not different from those from Borneo. In the Philippines the largest simple fronds seen are 18 x 6cm (sterile) and 13 x 4cm (fertile). The type of Dryopteris korthalsii has up to 6 pairs of rather small fertile pinnae (4x 1.3 cm); there are similar fronds from Borneo with pinnae 10 x 2 cm. Under the name Aspidium menisciicarpon BL. METTENIUS described a specimen of a species of Tectaria from the Philippines; his description was copied by HOOKER who transferred the name to Polypodium. BEDDOME’s description of Neph- rodium glandulosum (Handb. Suppl. 70) was based mainly on specimens cited which belong to the present species, but in part on the true Aspi- dium glandulosum BL. CHRISTENSEN (Ind. Fil. 1905) confused the present species with A. linea- tum BL. and A. affine BL. 38. Pronephrium hewittii (COPEL.) HOLTTUM, comb. nov. — Dryopteris hewittii COPEL. Philip. J. Sci. 3 (1909) Bot. 344; v.A.v.R. Handb. Suppl. (1917) 189. — Thelypteris hewittii (COPEL.) REED, Phytologia 17 (1968) 282.—Type: Brooks & HEWITT s.n. Feb. 1908, Sarawak, Bongo Range (MICH). Dryopteris compacta COPEL. Philip. J. Sci. 6 (1911) Bot. 137, pl. 18; v.A.v.R. Handb. Suppl. (1917) 176.— Thelypteris compacta (COPEL.) REED, Phytologia 17 (1968) 268.— Type: BROOKS 4, Sarawak, Bongo Range (MICH; BM). — Fig. 15e-i. Caudex short-creeping; fronds subdimorphous. Sterile fronds. Stipe 10-15cm long, glabrous; basal scales thin, to 31mm. Lamina to 25cm long; apex narrowly acuminate, to 5 x | cm, lobed at base; pinnae to 20 pairs, close, spreading at right angles to rachis; lower pinnae deflexed, nar- rowed to base on both sides, with a small acros- copic auricle. Middle pinnae 4-6cm long, 0.9- 1.1 cm wide above the truncate and strongly auri- cled base; apex acuminate; edges crenate throughout, distally to a depth of 1mm; costules 2.5-3 mm apart, at 45° to costa; veins to 3 pairs, 15-2 pairs anastomosing, sinus-membrane distinct but very short; lower surface of rachis glabrous, of costae bearing scattered hairs 0.5mm long, a few shorter hairs on costules and veins, surface between veins slightly pustular; upper surface of rachis bearing thick pale hairs | mm long, costae covered with hairs 0.2-0.3 mm long, a few longer ones near ends of veins. Fertile frond. Stipe to 25 cm long; pinnae to 4cm long, 0.5—0.9 mm wide, auricled as sterile but usually less deeply crenate; sori near costules; indusia firm, dark, almost cir- cular with hairs in the middle; sporangia not seti- ferous. Distr. Malesia: Borneo (Western Sarawak, 4 collections). Ecol. “On small rock well above stream” (MOLESWORTH ALLEN 3048, foot of Gunong Santubong). Notes. The type of D. hewittii is a small plant with dimorphic fronds; that of D. compacta much larger with fertile pinnae nearly as wide as sterile; two later collections have narrower fertile pinnae. In 1972 I united this species with P. merrillii but the latter is quite distinct in venation and sori. 39. Pronephrium affine (BL.) PRESL, Epim. Bot. (1851) 259; HoLTTUM, Blumea 20 (1972) 121, excl. syn. praeter BL. et Gymnogramme macrotis KUNZE. — Aspidium affine BL. En. PI. Jav. (1828) 148.— Type: BLUME, W. Java (L, n. 908, 333- 740). Gymnogramme macrotis KUNZE, Bot. Zeit. 6 (1848) 114.—Dryopteris oxyotis ROSENST. Meded. Rijksherb. n. 31 (1917) 5, nom. nov. (not D. macrotis (HOOK.) O. KTZE).—Type: ZOL- LINGER 324z, Java (L). Aspidium lineatum sensu METT. Farngatt. IV (1858) 110, quoad syn. A. affine BL. et G. macrotis 528 FLORA MALESIANA (ser. II, vol. 1° KUNZE tantum.— Nephrodium lineatum sensu HooK. Spec. Fil. 4 (1862) 75, p.p.; RACIB. FI. Btzg 1 (1898) 186.— Dryopteris lineata sensu C. Cur. Ind. Fil. (1905) 63, 275, p.p.; sensu v.A.v.R. Handb. (1908) 209; sensu BACKER & POSTH. Varenfl. Java (1939) 60. Caudex short-creeping or suberect; fronds dimorphous. Sterile fronds. Stipe 15-20cm long, glabrous except in groove, basal scales very thin, to 4x2mm. Lamina to 25cm long, thin; apex small and deeply lobed; pinnae to 12 pairs; basal pinnae somewhat narrowed towards base both sides with acroscopic auricle. Middle pinnae of type 4x 1.7 cm (width above base), of a Sumatran specimen 7X2.0cm; base truncate with acros- copic auricle 2-4 mm long; apex of type abruptly short-pointed, of Sumatran specimen short-acu- minate; edges crenate throughout in smaller plants, distally on larger ones; costules 3-3.5 mm apart, at less than 60°; veins 5-6 pairs, 3-33 pairs anastomosing, sinus-membrane very short; lower surface of rachis bearing coarse hairs to 0.7 mm long, hairs on costae 0.1mm, sometimes with a few longer ones, similar hairs distally on costules and veins, surface between veins pustular; hairs on upper surface of rachis to 0.5mm long, very short on costae and scattered distally on veins, none between veins. Fertile fronds. Stipe to 35 cm long; pinnae of type 3.0x0.7cm, of others to 4.0 x 0.9 cm, base strongly auricled (sometimes on both sides), edges almost entire; veins to 4 pairs; sori medial, slightly elongate; indusia thin, bearing copious acicular hairs 0.1-0.2mm or sometimes short capitate hairs; sporangia lacking setae; spores pale with a rather broad continuous wing and cross-wings. Distr. Peninsular Thailand, in Malesia: Central & S. Sumatra, West Java. Ecol. By streams in forest, low altitudes. Notes. The description by METTENIUS of A. lineatum was evidently based on specimens of several distinct species, doubtfully including the type of A. affine; it was copied by HOOKER who varied the list of synonyms, and the status of A. affine as a synonym of A. lineatum was accepted by CHRISTENSEN, who cannot have read BLUME’s descriptions. But the descriptions of RACIBORSKI, VAN ALDERWERELT and BACKER & POSTHUMUS clearly apply to the present spe- cies only. BLUME’s type matches specimens from a low altitude in Sumatra which, like BLUME’s, were found on limestone rocks by streams. A specimen from Pattani in Thailand is similar. 40. Pronephrium borneense (HOOK.) HOLTTUM, comb. nov.— Polypodium borneense HOOK. Spec. Fil. 5 (1863) 11. — Dryopteris labuanensis C. Cure. Ind. Fil. (1905) 273, nom. nov. (not D. bor- neensis (HOOK.) O. KTZE).— Phegopteris bor- neensis (HOOK.) v.A.v.R. Handb. (1908) 500. — Thelypteris borneensis (HOOK.) REED, Phytolo- gia 17 (1968) 264. — Type: T. Loss, Labuan (K). Caudex short-creeping; fronds not or little dimorphous. Stipe to 12cm long, pale, minutely hairy; basal scales thin, c. 2 1mm. Lamina thin, to 18cm long; apex 6-9 cm long, triangular, lobed near base which is 3-4 cm wide; pinnae 4—S pairs, not opposite; basal pinnae narrowed towards base on basiscopic side, slightly so to auricled acros- copic base. Middle pinnae 3.5 x 1.5 to 5.0 x 2.2 cm; base truncate with auricle 2-3 mm long; apex ab- ruptly short-pointed; edges shallowly crenate throughout; costules 2.5-3mm apart; veins 6 pairs, slender, at a broad angle to costules, almost all anastomosing to form a zig-zag excurrent vein (rarely with gaps in it), sinus-membrane not evident; lower surface of rachis and costae spar- sely short-hairy, rest glabrous and slightly pus- tular; upper surface of rachis with copious hairs 0.2-0.7 mm long, costal hairs 0.3 mm, a few short hairs also present near margin. Sori small, not elongate, mostly medial; indusia very small (sometimes lacking?), red when young, variable in shape, with a few mostly marginal hairs 0.1 mm long; sporangia with 4-5 short setae; spores with a narrow wing and a few cross-wings. Distr. Malesia: Borneo (Sarawak, Sabah). Ecol. ‘Hill forest, on clay bank, 420m” (JERMY 13072, G. Mulu); ‘on wet rocks, 300 m” (BROOKS l6c, Bongo Range). Notes. The type is a single fertile frond. Living plants of JERMY 13072, cultivated at Kew, resemble the type closely and have sterile and fertile fronds of similar size; the above description is based on the type and cultivated plants. Dried specimens of JERMY 13078, also from G. Mulu, are larger, with stipes 30cm long and dimorphous fronds to 30 cm long with 6 pairs of pinnae; sterile pinnae to 8.5 2.8cm (veins 7 pairs), fertile to 4.5 x 1.6cm. 41. Pronephrium exsculptum (BAK.) HOLTTUM, Blumea 20 (1972) 117. — Acrostichum exsculptum BAK. J. Bot. 26 (1888) 326.— Leptochilus exsculptus (BAK.) C. CHR. Ind. Fil. (1905) 9, 385; v.A.v.R. Handb. (1908) 740.— Dryopteris exsculpta (BAK.) COPEL. Philip. J. Sci. 37 (1928) 410; C. CHR. Gard. Bull. Str. Settl. 7 (1934) 248, p.p.— Thelypteris exsculpta (BAK.) K. IWATS. Acta Phytotax. Geobot. 21 (1965) 170. — Type: C. HOSE 244, Sarawak, Niah (K). Meniscium stenophyllum BAK. J. Bot. 29 (1891) 108. — Phegopteris stenophylla (BAK.) v.A.v.R. Handb. (1908) 510.— Thelypteris stenophylla (BAK.) REED, Phytologia 17 (1968) 315. — Type: G. F. HOSE 20, Sarawak (K). Caudex short-creeping; fronds dimorphous. Sterile fronds: Stipe 10-17cm long, glabrous except distally; basal scales to 51mm, dark, firm. Lamina 25cm long, very firm; apex small, triangular, deeply lobed; pinnae 20-25 pairs; basal 1981] THELYPTERIDACEAE (Holttum) 529 pinnae slightly narrowed both sides towards auri- cled base. Middle pinnae 3.5 X 1.0cm; base trun- cate, slightly auricled; apex short-acuminate (pin- nae of type of M. stenophyllum to 2.0 x 0.8 cm, apex shortly obtuse); edges crenate to a depth of 0.5mm; costules 2—2.5mm apart; veins 3 pairs, thick, pale and prominent on lower surface, 2 pairs anastomosing, sinus-membrane not evident; lower surface of rachis bearing thick curved hairs less than 0.5mm long, a few minute hairs on costae, rest of surface glabrous, not or hardly pustular; upper surface of rachis as lower, sparse short hairs on costa. Fertile fronds: Stipe 25 cm or more long; pinnae to 2.0cm long (type of M. stenophyllum 1.0 cm), 0.3—-0.4 cm wide above base which is strongly auricled (sometimes both sides); veins 2 pairs; sori near costules, at maturity filling lower surface; indusia small with a few very short marginal hairs; sporangia not setiferous; spores not seen. Distr. Malesia: Borneo (Sarawak). Ecol. Probably on limestone. Note. CHRISTENSEN (l.c. 1934) referred here specimens with thinner pustular pinnae and seti- ferous sporangia; these are now placed in P. rhombeum and P. peltatum. 42. Pronephrium merrillii (CHRIST) HOLTTUM, Blumea 20 (1972) 117.—Dryopteris merrillii CHRIST, Philip. J. Sci. 2 (1907) Bot. 201. — Phe- gopteris merrillii (CHRIST) v.A.v.R. Handb. Suppl. (1917) 505.—Cyclosorus merrillii (CHRIST) CopEL. Gen. Fil. (1947) 143; Fern Fl. Philip. (1960) 352.— Thelypteris merrillii (CHRIST) REED, Phytologia 17 (1968) 292.— Type: Fox- WORTHY 742, Palawan (P). Differs from 38. P. hewittii as follows: fertile pinnae to 2.7 x 0.4 cm, entire or nearly so; indusia very small, asymmetric, with a few very short marginal hairs. Distr. Malesia: Philippines (Palawan, 2 collec- tions; second is ELMER 13031). 43. Pronephrium rhombeum (CHRIST) HOLTTUM, Blumea 20 (1972) 120.— Dryopteris diversiloba var. acrostichoides subvar. rhombea CHRIST, Philip. J. Sci. 2 (1907) Bot. 200.— Dryopteris acrostichoides var. rhombea (CHRIST) v.A.v.R. Handb. Suppl. (1917) Corr. 49.— Dryopteris rhombea (CHRIST) COPEL. Philip. J. Sci. 56 (1935) 102, pl. 6 quoad icon. photogr. tantum. — Cyclosorus rhombeus (CHRIST) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 192; COPEL. Fern Fl. Philip. (1960) 357, p.p.— Thelypteris rhombea (CHRIST) REED, Phytologia 17 (1968) 309. — Lectotype (COPELAND 1935): CUMING 149, Luzon (orig. PNH destroyed; isotype K). Dryopteris diversiloba var. acrostichoides sub- var. lanceola CHRIST, lI.c.— Dryopteris acrosti- choides var. lanceola (CHRIST) v.A.v.R., l.c.— Dryopteris lanceola (CHRIST) COPEL. Philip. J. Sci. 56 (1935) 102, pl. 7. — Cyclosorus lanceolus (CHRIST) COPEL. Gen. Fil. (1947) 142; Fern FI. Philip. (1960) 353.— Thelypteris —_lanceola (CHRIST) REED, Phytologia 17 (1968) 286.— Lectotype (COPELAND 1935): COPELAND 250, Luzon, Mt Mariveles (orig. PNH destroyed; iso- types K, MICH, US). — Fig. 15j-1. Caudex short-creeping; fronds dimorphous. Sterile fronds. Stipe 5-8cm long with pale hairs 1 mm long throughout. Lamina to 15 cm long, thin but firm; apex narrowly triangular, longer than pinnae; pinnae 8(-10) pairs; basal pinnae not or little narrowed towards base. Middle pinnae 2.0- 3.5 cm long, 0.7—1.0 cm wide above auricled base; apex abruptly acute or obtuse, sometimes a little dilated; edges crenate, more deeply towards apex; costules 2mm apart, at little more than 45° to costa; veins mostly 3 pairs, concolorous and slightly prominent, 2 pairs anastomosing, sinus- membrane not evident; lower surface of rachis bearing pale hairs 0.7 mm long, a few similar hairs with shorter ones present on costae, sparse short hairs on costules and veins, surface between veins pustular; upper surface of rachis hairy as lower, hairs on costae 0.2-0.3 mm long, a few short hairs near margin. Fertile fronds. Stipe to 25cm long; lamina 10-20cm long, in the latter case pinnae more widely spaced; pinnae 2.0-3.0 cm long, 0.4— 0.6cm wide, base auricled, edges almost entire; sori near costules, not elongate; indusia small, red, with marginal hairs 0.2-0.3 mm long; sporan- gia mostly with 2-3 short setae (in some cases none); spores with translucent wing and cross- wings. Distr. Malesia: Philippines (Luzon to Min- danao), N. Borneo, Celebes. Notes. The Kew isotypes of the lectotypes cited do not differ in the ways indicated in COPELAND’s keys and descriptions of 1960, and among many specimens seen I cannot see two distinct forms. The Kew specimen of COPELAND 250 has pinnae more dilated distally than that of CUMING 149, though COPELAND indicates the contrary. The pinnae with irregular large distal lobes figured (as line drawings) with the pho- tograph of the type of D. rhombea by COPELAND (1935) are representative of a group of forms, almost always with 4-5 pairs of pinnae, which appear to be hybrids and are here treated as P. x xiphioides. 44. Pronephrium peltatum (v.A.v.R.) HOLTTUM, comb. nov.— Dryopteris peltata v.A.v.R. Bull. Jard. Bot. Btzg II, 16 (1914) 12; Handb. Suppl. (1917) 190. — Thelypteris peltata (v.A.v.R.) REED, Phytologia 17 (1968) 303. — Type: MATTHEW 632, Sumatra, Gunong Tandikat (BO). Dryopteris zippelii ROSENST. Meded. Rijksherb. n. 31 (1917) 6.—Type: ZIPPEL, Java (L) Dryopteris lineata var. subacrostichoides 530 v.A.v.R. Bull. Jard. Bot. Btzg II, 28 (1918) 24. — Type: CRAMER 37, S. Sumatra, Kota Agoeng (BO). BEY TO THE VARTEGIES 1. Indusia glabrous; sporangia not setiferous. 2. Sterile pinnae thin, margin deeply serrate dis- tally nee Bess . a. var. peltatum . Sterile pinnae rigid when dry, entire b. var. tenompokense 1. Indusia with a few short hairs; sporangia seti- ferous or not. 3. Sporangia not setiferous 3. Sporangia setiferous. 4. Sporangia with several setae; basal pinnae not reduced d. var. persetiferum 4. Sporangia with 0-2 setae; reduced basal pinnae present on most specimens e. var. aberrans in) c. var. peninsulare a. var. peltatum Stipe of sterile frond 14cm long, of fertile 20cm. Lamina to 21cm long; pinnae c. 10 pairs. Sterile pinnae to 6cm long, base rounded on basiscopic side and auricled on acroscopic, 1.5 cm wide above base, apex short-acuminate, edges crenate throughout or in distal part only; costules 2.5mm apart; veins 3-4 pairs, almost all anas- tomosing; lower surface glabrous, pustular when dry; upper surface hairy on costae only. Fertile pinnae to 4X 1 cm; sori medial; indusia firm, dark, lacking acicular hairs; sporangia not setiferous. Distr. Malesia: Central & S. Sumatra, W. Java, at 900-1500 m. Note. Specimens from Tjibodas in W. Java have sterile pinnae to 8x 1.6cm. One specimen from Sumatra has lower sterile pinnae auricled on both sides at the base. b. var. tenompokense (C. CHR.) HOLTTUM, Stat. nov. — Dryopteris tenompokensis C. CHR. Gard. Bull. Str. Settl. 7 (1934) 248. — Type: HoLTTUM 25388, Sabah, Mt Kinabalu, Tenompok (BM; K, SING). Differs from var. peltatum as follows: texture of lamina firm; edges of pinnae almost entire; FLORA MALESIANA [ser. II, vol. 1° lower surface of costae glabrous or sparsely minutely hairy. Distr. Malesia: Sabah (Mt Kinabalu). Ecol. In forest, on rocks by streams or near streams, at 1500 m. c. var. peninsulare HOLTTUM, var. nov.— Abacopteris lineata sensu HOLTTUM, Rev. FI. Mal. 2 (1955) 292, excl. syn., f. 169. — Type: A. G. PIGGOTT 1668, Malaya, Negri Sembilan, G. Telapak Burok, 900 m (K). A var. tenompokense differt indusiis semper breviter pilosis. Distr. Malesia: Malaya, on Main Range and Taiping Hills. Ecol. In forest, not (or not always) by streams, 900-1500 m. Note. A plant of this variety from Taiping Hills, cultivated at Kew, was found by MANTON to be tetraploid (HOLTTUM I.c. p. 625). d. var. persetiferum HOLTTUM, var. nov. — Dryopteris exsculpta sensu C. CHR. Gard. Bull. Str. Settl. 7 (1934) 248, p.p.— Type: HOLTTUM s.n. 1931, Mt Kinabalu, crest of Penibukan Ridge, 1200 m (SING; BM). A var. tenompokense_ differt pinnis crenatis, sporangiis setis pluribus praeditis. Distr. Malesia: Borneo. Besides type, known from G. Mulu, Sarawak, in stream-bed at 1760 m. leviter e. var. aberrans HOLTTUM, var. nov. A var. tenompokense differt: frondibus pinnis inferioribus redactis usque 12-jugatis, aequaliter bilobatis (lobis usque 1.2cm longis) praeditis; pinnis normalibus sterilibus usque 122.5 cm, distaliter crenatis, fertilibus usque 7X 1.7cm; sporangiis 0-2 setis praeditis. —Type: CLEMENS 33702 & 32020’, Sabah. Mt Kinabalu, Colombon Basin, 1400-1600 m (K; G, MICH). Distr. Malesia: Sabah. Note. These specimens have the frond-form of Sphaerostephanos, but the reduced basal pinnae are not like those of that genus; apart from size, the normal pinnae agree with the present species. Spores of a specimen at MICH are normal. 2. Subgenus Menisciopsis HOLTTUM, subg. nov. Pinnae subintegrae, plurivenosae, venis plerisque more generis Meniscii Schreb. anastomosantibus; sori exindusiati, saepe elongati, interdum coalescentes; sporangia vel nudis vel pilis rectis vel hamatis praedita. Type species: Pronephrium lakhimpurense (RosENST.) HOLTTUM, Blumea 20 (1972) 110. Taxon. The type species, P. lakhimpurense, is distributed from N.E. India to southern China and southwards to northern Thailand and Vietnam. It has large entire pinnae with many pairs of anastomos- ing veins, the excurrent veinlets all free; the sori of large plants spread all along the veins, the sporangia 1981] THELYPTERIDACEAE (Holttum) 531 lacking setae. In these characters it is very close to the species of Meniscium of tropical America, and differs from them most conspicuously in the red colour of dried fronds. Another species of the same distribution in Asia, P. penangianum (HOOK.) HOLTTUM, has narrower pinnae which are strongly crenate-serrate and distal excurrent veinlets not free. In Malesia, P. rubrinerve, P. rubidum and P. scopulorum are clearly related to the two species of mainland Asia, but their sori are not more than slightly elongate; only in P. rubidum are excurrent veinlets all free. Agreeing with the species mentioned above in the reddish colour of dried fronds and also in elongate sori are species of another Malesian group which are distinguished by having hooked hairs on frond-axes and usually also on sporangia; formerly these species were all included in Meniscium. Several of them have buds at the bases of pinnae. They are here included in section Grypothrix. There remain three species which appear to be somewhat intermediate between the two groups above mentioned. Of these, P. acanthocarpum has buds at the bases of its pinnae as in sect. Grypothrix but sori not elongate and straight setae on its sporangia. The other two, P. nitidum and P. repandum, have similar sori and sporangia but no buds on the frond; they are very similar to each other in general aspect but differ in sporangia and spores. They are possibly a connecting link between subgenera Pronephrium and Menisciopsis. They are here placed in section Menisciopsis. The earlier-described species of this group were much confused by HOOKER, and it was not until the work of CHING in 1938 that their taxonomy and nomenclature were clarified. CHING however did not notice the hooked hairs which are the distinctive feature of sect. Grypothrix. 3. Section Menisciopsis KEY TO THE SPECIES 1. Body of sporangium lacking setae. 2. Pinnae 4-6 cm wide: edges distinctly serrate throughout 2. Pinnae 2.5—3.5 cm wide; edges entire or nearly so. 3. Excurrent veinlets mostly free; sori close to costules = Excurrent veinlets mostly not free; sori not close to costules . Body of sporangium bearing setae. m Pinnae one pair, a bud at the base of each 4. Pinnae several pairs on adult plant; no buds present. 5. Caudex long-creeping: veins very oblique; several acicular hairs on stalk of sporangium 49. P. nitidum 5. Caudex short-creeping; veins at a broad angle to costules; a hair of 2-3 cells, not acicular on stalk of sporangium 50. P. repandum . 45. P. rubrinerve .46. P. rubidum .47. P. scopulorum 48. P. acanthocarpum 45. Pronephrium rubrinerve (METT.) HOLTTUM, _ pinna-like; basal pinnae distinctly stalked, nar- Blumea 20 (1972) 110; Allertonia 1 (1977) 213; BROWNLIE, Pterid. Fiji (1977) 258. — Phegopteris rubrinervis METT. in Kuhn, Linnaea 36 (1869) 116; KUHN, Verh. Zool. Bot. Ges. Wien 19 (1869) 576; v.A.v.R. Handb. Suppl. (1917) 316. — Goniopteris rubrinervis (METT.) CARR. in Seem. FI. Vit. (1873) 366.— Polypodium rubrinerve (METT.) BAK. Syn. Fil. ed. 2 (1874) 315.— Cyclosorus rubrinervis (METT.) COPEL. Gen. Fil. (1947) 143; J. Arn. Arb. 30 (1949) 438.— Thelypteris rubrinervis (METT.) K. IWATS. Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 195.— Lectotype (HOLTTUM 1972): TURNER, New Ireland (B). Caudex massive, short-creeping to suberect; fronds closely tufted; scales small, thin, broad- ovate, glabrous or nearly so. Stipe to 145 cm long, glossy, glabrous apart from a few very short aci- cular hairs near base (often lost), drying light reddish; lamina to 170cm long, thin but firm, drying brown-olivaceous with red costae and in some cases costules; pinnae to 24 pairs, upper ones on large fronds gradually decrescent; apex rowly cuneate at base, the lamina on basiscopic side ending 1.5-2.0 cm from rachis. Largest pinnae to 436cm; base broadly cuneate: apical half gradually narrowed to short-acuminate tip: edges crenate to a depth of 1mm, cartilaginous margin thick, pale to reddish; costules 4-5 mm apart, at 60° to costa, falcate distally; veins 15-20 pairs, at 45° to costules, almost straight, fine and slightly prominent on lower surface, almost all anas- tomosing, excurrent veinlets, except basal ones, rarely free: lower surface of rachis and pinnae quite glabrous (young fronds may have thin red- dish scales on rachis and costae), surface between veins slightly pustular; upper surface of both rachis and costae glabrous. Sori medial or supramedial, basal ones sometimes meeting and slightly elongate; no indusia; sporangia lacking setae, a hyaline hair of 2 cells on the sporangium- stalk; spores red-brown, with a rather broad translucent wing and cross-wings. Distr. Polynesia (Fiji), Melanesia (New Heb- rides) and Malesia: New Guinea (New Ireland). 532 Ecol. On river banks in forest at low altitude (to 900 m in Fiji). Notes. In 1977 I reported the caudex of this species to attain a height of 30cm, on the evi- dence of a herbarium label; but BRAITHWAITE reports it to be prostrate, and this is also true of a plant he sent from the New Hebrides, cultivated at Kew. 46. Pronephrium rubidum (HOOK.) HOLTTUM, Blumea 20 (1972) 111.— Polypodium rubidum HooK. Spec. Fil. 5 (1863) 12, excl. DE VRIESE 41.— Dryopteris rubida (HOOK.) C. CHR. Ind. Fil. (1905) 289; CHRIST, Philip. J. Sci. 2 (1907) Bot. 207, excl. WHITFORD 272. — Phegopteris rubida (HOOK.) v.A.v.R. Handb. (1908) 502. — Cyclosorus rubidus (HOOK.) COPEL. Gen. Fil. (1947) 143; Fern Fl. Philip. (1960) 375. — Thelyp- teris rubida (HOOK.) K. IwWATS. Mem. Coll. Univ. Kyoto B, 31 (1965) 195.— Type: CUMING 415, Euzoni(K; E; FIEW,G, LE; P).: Caudex short, suberect. Stipe dark reddish, glabrous except for minute hairs near base, to 100cm long; basal scales not seen, reported by CHRIST to be 2cm long. Lamina to 120cm long; apex pinna-like; pinnae to 20 pairs, subopposite, drying dark reddish; basal pair not or little reduced, with stalks 1-2 mm long. Largest pinnae of type 21 x 2.7cm, largest seen 28 x 3.8 cm; base cuneate (angle c. 45° each side); apex caudate- acuminate (cauda 2.5-3.0cm long); edges irre- gularly slightly sinuous to slightly crenate, car- tilaginous margin rather broad and red; costules 3-5 mm apart, at 60°; veins 9-13 pairs, at 45° to costules, slightly curved, dark and not prominent on lower surface, excurrent veinlets all free; lower surface of rachis and pinnae quite glabrous; upper surface of rachis with a few short hairs in the groove, hairs on costae sparse, slender. Sori very close to costules except basal basiscopic ones on large pinnae, sporangia few; no indusia; no setae on body of sporangia; hair on sporangium-stalk 3-celled, distal cell not much enlarged. Distr. Malesia: Philippines (Luzon; Palawan, 4 collections). Ecol. “Streamside, in open pine 1700 m” (PRICE 2831; no other record). Notes. The largest specimen is FOxWORTHY 684 from Palawan (MICH). HOOKER described var. B, but he cited no specimen and I have not found one so marked. DE VRIESE 41, wrongly cited by HOOKER, is P. ramosii, as also WHIT- FORD 272 wrongly cited by CHRIST. forest, 47. Pronephrium scopulorum HOLTTUM, sp. nov. A P. rubido differt: pinnis omnibus basi anguste inaequaliter cuneatis, pinnis basalibus non stipi- tatis; venulis excurrentibus plerisque non liberis; soris costulas non _ contingentibus. — Type: BUWALDA 5168, Aru Is., P. Kabroor, on lime- stone (K; BO). FLORA MALESIANA [ser. II, vol. 1° Caudex not seen. Stipe probably to 50 cm long, dark reddish, glabrous; basal scales not seen. Lamina c. 50 cm long; apex pinna-like; pinnae to 8 pairs, firm, drying dull reddish, subopposite; bases of all pinnae narrowly unequally cuneate, lowest pinnae with narrowest base. Middle pinnae 18 x 2.8-3.5 cm, widest 1/3 from base; acroscopic base narrowed to attachment to rachis, lamina on basiscopic side ending 3-4 mm from rachis; apex narrowly caudate; edges entire or slightly sinuous distally; costules 3-4mm apart at 60° to costa; veins to 10 pairs, at c. 45° to costules, almost straight, almost all anastomosing to form zig-zag excurrent veins between costules, basal basiscopic vein arising from costa; both surfaces quite glabrous (upper surface of rachis and costae probably have some hairs when pinnae are young). Sori near costules but not touching them, basal ones sometimes supramedial; no indusia; sporangia lacking setae. Distr. Malesia: S.E. Moluccas (Aru Is.: P. Kabroor) and West New Guinea (Waigeu L: CHEESMAN 1240-1243, BM). Ecol. On steep limestone which carries little other vegetation. 48. Pronephrium acanthocarpum (COPEL.) HOLTTUM, Blumea 20 (1972) 107.— Dryopteris acanthocarpa COPEL. Philip. J. Sci. 6 (1911) Bot. 136, pl. 17. — Phegopteris acanthocarpa (COPEL.) v.A.v.R. Handb. Suppl. (1917) 315. — Thelypteris acanthocarpa (COPEL.) REED, Phytologia 17 (1968) 257.—Type: BRooKs 54, Sarawak, Mt Penrissen (MICH; BM). Caudex short-creeping; bases of stipes less than lcm apart. Stipe to 50cm long, dark at base with small dark scales and sparse minute hairs, rest glabrous, dull reddish-brown. Lamina consisting of one pair of opposite pinnae with a bud at the base of each and an apical section larger than pinnae, the whole dull reddish when dry. Apical lamina to 23 x 10 cm, widest near broadly rounded and symmetric base, apex short-acuminate, edges irregularly slightly sinuous; main veins 6mm apart, at wide angle to midrib, falcate distally; veins more than 20 pairs, not prominent, at a wide angle to main veins, almost all anastomosing with occasional irregularities, excurrent veinlets free; lower surface of midrib copiously minutely hairy (less than 0.1mm), similar hairs sparse on main veins, surface between veins not pustular; sori medial or inframedial, those on basal veins some- times supramedial, in many cases slightly elon- gate; sporangia bearing several short straight setae; spores not seen. Pinnae to 17 X7cm; base broadly cuneate and very asymmetric, the basis- copic side 4.5 cm wide at 1/3 length of pinna, acro- scopic side 3cm; veins to 18 pairs on basiscopic side; pubescence and sori as terminal lamina. Distr. Malesia: Borneo (Western Sarawak, two localities; the second is Mt Merinjak). 1981] THELYPTERIDACEAE (Holttum) 533 Ecol. On rocks near streams at 900 m. 49. Pronephrium nitidum HOLTTUM, Blumea 20 (1972) 109.— Dryopteris urophylla (METT.) C. Cur. var. nitida HOLTTUM, Gard. Bull. Str. Settl. 7 (1934) 249, 251 (fig.).— Thelypteris urophylla var. nitida (HOLTTUM) K. IWATs. Acta Phytotax. Geobot. 22 (1966) 94.— Type: HoLTTUM SFN 25592, Sabah, Mt Kinabalu, Menetendok (SING; BO, K). Cyclosorus urophyllus sensu COPEL. Fern FI. Philip. (1960) 373, excl. loc. Malacca. — Fig. 16g—h. Caudex long-creeping, 5-8mm diameter. Stipe commonly 70-100 cm long; base dark with dark rigid scales c. 5X 1mm bearing many short stiff hairs, above base brownish to slightly reddish, glabrous. Lamina 50cm long, firm; apex pinna- like, often with almost symmetrical base; pinnae 6-7 pairs, subopposite, lowest pinnae usually with a Stalk to 4mm long, narrowly and asymmetrically cuneate at base, upper pinnae sessile with broader and more symmetric bases (very variable). Middle pinnae commonly 20 x 4-5 cm, narrowly elliptic, + abruptly narrowed to cuneate base and to caudate apex (cauda commonly 2 cm long); edges crenate- serrate with a thick pale cartilaginous margin; costules 3.5-4 mm apart, at a broad angle, falcate distally; veins 18-20 pairs, slender and slightly prominent both sides, very oblique and slightly S-curved, excurrent veinlets almost all free; lower surface of rachis and costae covered with pale thin hairs 0.2-0.3 mm long (antrorse but not ap- pressed on costae), similar and sometimes also longer hairs on costules, surface between veins glabrous or with some very short erect hairs, at most slightly pustular; upper surface of rachis and costae glabrous or with sparse short hairs, surface between veins smooth and glossy. Sori supra- medial, those on connivent veins sometimes + confluent; receptacle bearing many short acicular hairs with the sporangia; sporangia short-stalked with several acicular hairs on stalks, also 8-12 short acicular hairs distally on body; spores closely and minutely papillose. Distr. Malesia: Banka, Borneo, N. & Central Celebes, Philippines (Sulu Archipelago, Min- danao). Ecol. At 0-1200 m, usually in secondary forest, often on edges of forest or rather open places on stream-banks. Note. COPELAND (1960) reports from Palawan, but I have seen no specimen thence. Both sporangia and spores are markedly different from those of P. repandum. 50. Pronephrium repandum (FEE) HOLTTUM, Blumea 20 (1972) 109.—Goniopteris repanda FEE, Gen. Fil. (1852) 251.—Type: GAvuDI- CHAUD, Penang (FI-W, isotype). Polypodium cuspidatum ROxB. Calc. J. Nat. Hist. 4 (1844) 491 (not Pronephrium cuspidatum (BL.) HOLTTUM).— Type: Penang (BR; K). Goniopteris dalhousiana FEE, 8e Mém. (1857) 92.— Type: Lady DALHOUSIE, “Indes Orien- tales” (Herb. Graham, not seen; probable isotype, from Penang, K). Phegopteris urophylla MetTT. Farngatt. IV (1858) 26, excl. var. aspera (PRESL).— Poly- podium urophyllum (METT.) HOOK. Spec. Fil. 5 (1863) 9, excl. syn. et var.— Nephrodium urophyl- lum (METT.) KEYSERL. Pol. et Cyath. Herb. Bung. (1873) 49; BEpp. Handb. (1883) 274, p.p. — Dryopteris urophylla (METT.) C. CHR. Ind. Fil. (1905) 299, p.p.; v.A.v.R. Handb. (1908) 216, p.p.; BACKER & PosTH. Varenfl. Java (1939) 64, p.p.— Abacopteris urophylla (METT.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 251; HOLT TUM, Rev. Fl. Malaya 2 (1955) 296, f. 172. — Cyclosorus urophyllus (METT.) TARD. Notul. Syst. 7 (1938) 77; TARD. & C. CHR. FI. Gén. I.-C. 7, pt. 2 (1941) 391, p.p. — Thelypteris urophylla (METT.) K. IWATS. S.E. Asian Studies 3 (1965) 81; Acta Phytotax. Geobot. 22 (1966) 94.— Type: WALLICH 299, Penang (B?; isotype K). Polypodium pinwillii BAK. Ann. Bot. 5 (1891) 460.— Nephrodium uropyllum_ var. _ pinwillii BEDD. Handb. Suppl. (1892) 72.—Type: PIN- WILL, Malacca (K). — Fig. In, 16i-j. Caudex short-creeping, fronds tufted at its apex. Stipe to 80cm long, slightly flushed with red, densely short-hairy with narrow very hairy scales at base, glabrescent above base. Lamina to 60cm long, firm, drying green but rachis, costae and costules + flushed with red; apex pinna-like with + asymmetric cuneate base; pinnae 3-6 pairs; lowest pinnae usually slightly smaller than next, always more narrowed at base than the rest, which are very variable in width in relation to length. Middle pinnae commonly c. 22 *4.0cm, largest 30x 7cm, widest at, or more commonly above, the middle, gradually narrowed to a rather abruptly cuneate base, apex abruptly caudate- acuminate, cauda 3-4cm long; edges + deeply crenate-serrate with very narrow _ pale cartilaginous margin; costules 4-5 mm apart, at 45-60°, slightly curved; veins 15-18 pairs, slender, slightly prominent both sides, slightly curved; at 45° to costule, excurrent veinlets often but not always free (if free, extending almost to the junction of the next pair of connivent veins); lower surface of rachis and costae + densely covered with erect pale hairs 0.2-0.3 mm long, similar hairs more sparse on costules, surface between veins finely pustular, bearing a variable number of short erect hairs some of which may be hooked; upper surface of rachis and costae hairy as lower surface, rest glabrous, surface between veins minutely pustular. Sori medial or supramedial, not elongate, exin- dusiate; sporangia bearing 10 or more short setae distally, stalk longer than in P. nitidum witha hair of WALLICH- 299, 534 FLORA MALESIANA [ser. II, vol. 1° one cell which is in some cases almost spherical; Moulmein southwards to Malesia: Malaya, spores with a continuous wing and cross- Sumatra, Riouw, Lingga & Anambas Is., and West wings. Java. Distr. South Thailand & Vietnam, from Ecol. In lowland forest, not in exposed places. 4. Section Grypothrix HOLTTUM, Blumea 19 (1971) 36. Type species: Pronephrium cuspidatum (BL.) HOLTTUM. Taxon. See discussion under subg. Menisciopsis. The earlier-known species of this section were placed in the genus Meniscium SCHREB. by SWARTZ, BLUME, PRESL, HOOKER, BEDDOME and RACIBORSKI, and in Phegopteris sect. Meniscium by METTENIUS and VAN ALDERWERELT. CHRIS- TENSEN (Monogr. Dryopteris 1, 1913, 248) was the first author to regard them as belonging to a group distinct from the type of Meniscium. CHING (1938) clarified the taxonomy and nomenclature of the species of Mainland Asia but did not note the hooked hairs by which these species are distinguished from those here placed in sect. Menisciopsis. P. triphyllum is distributed throughout the range of the genus Pronephrium; the others are more local in distribution. There are plants which appear to be hybrids of which one parent is P. triphyllum. In Malesia P. x parishii is so designated; there are others, more irregular in frond-form, in Sri Lanka and the Ryukyu Islands. FEE described a species Abacopteris elegans (Gen. Fil. 1852, 310), based on a specimen from Cochinchina which I have not seen; he stated that it had hooked hairs on sporangia, indusiate sori, and excurrent veinlets not free, a combination of characters which I have never seen, and I doubt the accuracy of his observations. KEY LO THESSRECIES uuCauder long- creeping with well-spaced fronds. . Pinnae one pair, opposite, always asymmetric : wh Di Pinnae 2-6 pairs, distal ones small and adnate, rest with variable asymmetry Caudex short-creeping, fronds close together. a Pinnae all narrowly cuneate at base, fertile ones ee lcm wide; hooked hairs lacking on lamina and sporangia 3 . . . . 53. P. salicifolium 3. Pinnae not all narrowly cuneate at base, fertile ones always much 1 more than | cm wide; hooked hairs present on some parts of lamina and usually on sporangia. 4. Pinnae 1-3 pairs; no buds at their bases. . Pinnae always opposite; fertile pinnae with undulate margins; hooked hairs present on basal scales and on sporangia 3 54. P. rubicundum 5. Pinnae not always opposite; fertile pinnae sharply serrate- crenate distally; no hooked hairs on scales nor on sporangia : 55. P. sulawesiense 4. Pinnae of well-grown plants more than 3 pairs; a 1 bud always present at base of uppermost pinna. 6. Bud confined to base of uppermost pinna; pinnae widest in basal half; hydathodes on upper surface conspicuous : : . 56. P. cuspidatum 6. Buds present at bases of several pinnae: pinnae “usually widest above middle; hydathodes not conspicuous. . Sat ®.. ene tatty ee ee . 2) J See ramos 7. Hooked hairs present on sporangia 57a. P. ramosii var. ramosii 7. Hooked hairs lacking on sporangia . 57b. P. ramosii var. minahassae . 51. P. triphyllum 52. P. Xx parishii 51. Pronephrium triphyllum (Sw.) HoLtTTum, Blumea 20 (1972) 122; Allertonia 1 (1977) 214; BROWNLIE, Pterid. Fiji (1977) 257. — Meniscium triphyllum Sw. in Schrad. J. Bot. 1800, 2 (1801) triphylla (Sw.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 241; Ho_truM, Rev. FI. Malaya 2 (1955) 287, f. 166.— Cyclosorus triphyllus (Sw.) TARD. Notul. Syst. 7 (1938) 78; TARD. & C. CHR. 16; HooK. & GREV. Ic. Fil. (1820) t. 120; BEDD. Ferns S. India (1863) t. 56; Handb. (1883) 397: Racis. Fl. Btzg 1 (1898) 66.— Phegopteris triphylla (Sw.) Metr. Fil. Lechl. (1859) 21; v.A.v.R. Handb. (1908) 509. — Dryopteris triphylla (Sw.) C. CHr. Ind. Fil. (1905) 298; BACKER & PosTH. Varenfl. Java (1939) 59.— Abacopteris Fl. Gén. I.-C. 7, pt. 2 (1941) 386; COPEL. Fern FI. Philip. (1960) 371. — Thelypteris triphylla (Sw.) K. IwaATs. Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 190. — Type: China (S-PA). Meniscium cumingii FEE, Gen. Fil. 222.— Type: CUMING s.n., Philippines seen). — Fig. 16k-I. (1852) (not 1981] THELYPTERIDACEAE (Holttum) 535 Caudex long-creeping, 2-3mm_ diameter, covered with short pale hooked hairs; scales c. 3mm long, narrow, bearing many short straight hairs. Stipe of sterile fronds 7-20cm long, of fertile 20-50cm, + tinged with dull red, rather sparsely hairy, hairs as caudex. Lamina consisting of an apical section and one pair of opposite pinnae attached 0.5-2.0 cm below it. Sterile apical lamina 10-18cm long, 2.5-4.0cm wide; base al- most symmetrical, broadly cuneate to rounded, rarely somewhat cordate; apex short-acuminate; edges entire or slightly sinuous; main veins 4— 4.5 mm apart along midrib; veins 10-12 pairs, pale, slender, slightly prominent on lower surface, at a broad angle to main veins, excurrent veinlets free or meeting the junction of the next pair of con- nivent veins; lower surface of midrib and main veins bearing pale hooked hairs 0.3-0.4 mm long, rest of surface glabrous, or with a few short erect hooked hairs, slightly pustular between veins; upper surface of midrib bearing copious short hooked hairs, hairs on main veins sparse. Fertile apical lamina 1.0-2.5cm wide; veins to 7 pairs, almost at right angles to main veins; excurrent veinlets almost always short and free; sori extending all along each vein so that sporangia are distributed in a crescent-shaped row along each pair of connivent veins; sporangia bearing hooked hairs distally and sometimes a hair of 2 cells on the stalk; spores bearing many separate but not crowded small wings. Sterile pinnae 5-10 cm long, 1.5-3.0 cm wide, with stalks 1-3 mm long, shaped as apical lamina but asymmetric, the lamina wider on basiscopic side of costa than on acroscopic; veins to c. 8 pairs on basiscopic side. Fertile pinnae 1.0-2.5 cm wide, sori as apical lamina. Distr. Tropical and subtropical mainland S.E. Asia, Taiwan, Ryukyu Islands; throughout Malesia; N. Queensland, Fiji (only 1 record from Fiji). Ecol. Low altitudes in light shade, sometimes abundant under fruit trees or palms in villages; few collections from Philippines and New Guinea. 52. Pronephrium x parishii (BEDD.) HOLTTUM, Blumea 20 (1972) 123.— Meniscium parishii BEDD. Ferns Brit. India (1866) t. 184. — Menis- cium triphyllum var. parishii BEDD. Handb. (1883) 399. — Abacopteris triphylla var. parishii (BEDD.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 241; HOLTTUM, Rev. FI. Malaya 2 (1955) 287. — Thelypteris triphylla var. parishii (BEDD.) K. IwaTs. Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 191. — Thelypteris parishii (BEDD.) PANIGRAHI, Phytologia 31 (1975) 372.— Type: PARISH 135, Burma, Moulmein (K). Caudex long-creeping, to 4mm diameter, as P. triphyllum in hairs and scales (hairs on scales of some Indian specimens not hooked). Lamina variable; apical section about as in P. triphyllum but often with 1 or 2 narrow lobes at the base, or 1-2 small narrow adnate free pinnae just below it, no buds at the bases of upper pinnae; pinnae to 6 pairs, usually opposite but sometimes not, usually increasing in size from apex to base of frond, upper ones always + adnate to rachis at basis- copic base; largest 15 x 2-3cm, sometimes with lamina symmetrically divided by the costa, some- times the two halves unequal in width as in P. triphyllum; pubescence as in P. triphyllum but more numerous erect hooked hairs often present on lower surface between veins. Sori as in P. triphyllum; sporangia sometimes with longer’ hooked hairs; spores not seen. Distr. Assam, Burma, Thailand, Vietnam; in Malesia: Malaya (Perak and Pahang). Ecol. In Malaya along stream banks in rather open forest, at low altitudes. Notes. The specimens referred here are vari- able, and in their small adnate upper pinnae agree with Meniscium thwaitesii HOOK. of Ceylon, but the pinnae of the latter are more irregular and some of its hairs are not hooked. Variability among the present specimens suggests hybridity, one parent certainly being P. triphyllum; it is possible that plants in different areas have different genetic origins. 53. Pronephrium salicifolium (WALL. ex HOOK.) HOLTTUM, Blumea 20 (1972) 123.— Meniscium salicifollum WALL. ex Hook. Ic. Pl. 10 (1854) t. 990; BEDD. Ferns Brit. India (1866) t. 207; Handb. (1883) 399.— Phegopteris salicifolia (WALL. ex Hook.) METT. Fil. Lechl. 2 (1859) 22; v.A.v.R. Handb. (1908) 511.—Abacopteris salicifolia (WALL. ex HooK.) HOLTTUM, Rev. FI. Malaya 2 (1955) 288, f. 167.— Thelypteris — salicifolia (WALL. ex HOOK.) REED, Phytologia 17 (1968) 311.— Type: WALLICH 63, Penang (K). Caudex short-creeping, 7-8 mm diameter, bear- ing sparse hooked hairs. Stipe 20-60 cm long; near base a few hooked hairs and rigid dark scales to 10 x 1 mm bearing pale hooked hairs, rest of stipe glabrous, glossy, + castaneous. Lamina 30-80 cm long, very firm, somewhat dimorphous, slightly flushed with red when dry; apex pinna-like, not or little larger than distal pinnae; pinnae to 18 pairs, a bud sometimes present at base of uppermost pinna; basal pinnae asymmetric at base, the blade decurrent to the attachment to rachis on acros- copic side, ending 5mm or more from rachis on basiscopic side. Middle pinnae commonly 10— 15cm long, fertile ones 0.6-1.2cm wide, sterile sometimes wider, largest sterile pinnae 20 2.5cm, fertile rarely to 2cm wide; base always very narrowly cuneate; apex narrowly acuminate but not caudate; edges quite entire; costules 3- 4mm apart, at 45° to costa; veins 3-10 pairs (according to width of pinna), almost all anas- tomosing, not prominent on either side, excurrent veins all free except distal ones; both surfaces quite glabrous. Sori supramedial, elongate along 536 FLORA MALESIANA [ser. II, vol. 1° veins, those on connivent veins usually meeting; no hairs on body of sporangia and none seen on stalk; spores with a + continuous wing and cross- wings. Distr. Malesia: Malaya, Sumatra, Borneo. Ecol. On rocks in and beside streams, in forest, in flood zone, at 0-500m; plants in the more exposed places on rocks in a stream-bed have smallish fronds with narrow ‘pinnae. The very firm narrow pinnae are probably an adap- tation to dry conditions when the stream is low and also to survival in the periodic rush of flood- water; in form they closely resemble the pinnae of Dipteris lobbiana (HOOK.) MOORE, which grows in similar habitats. 54. Pronephrium rubicundum (v.A.v.R.) HOLT- TUM, Blumea 20 (1972) 123. — Phegopteris rubi- cunda v.A.v.R. Bull. Jard. Bot. Btzg III, 2 (1920) 162. — Abacopteris rubicunda (v.A.v.R.) HOLT- TUM, Rev. Fl. Malaya 2 (1955) 292, f. 170.— Thelypteris rubicunda (v.A.v.R.) K. IWATS. Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 195. — Type: BROOKS 232/S, Sumatra, Lebong Tandai (BO; BM). — Fig. lo, 16a-—d. Caudex short-creeping, to 8 mm diameter, bear- ing hooked hairs. Young plants bearing simple fronds up to 30x 12cm, sometimes with fertile ones 20 x 5 cm; later fronds on these plants having lateral pinnae, the apical lamina progressively smaller. Stipe 20-30 cm long on sterile fronds, to 70cm on largest fertile ones, base bearing thin narrow scales to c. 8X1mm with a variable number of hooked hairs on them, above base + flushed with dull red and bearing sparse pale short hooked hairs. Lamina 20-35 cm long, firm, drying dull reddish, consisting of an apical segment and 1-3 pairs of opposite pinnae; no buds at bases of pinnae; all pinnae asym- metric, broader on basiscopic side of costa than on acroscopic; basal pinnae short-stalked. Apical lamina of fronds with 3 pairs of pinnae 15-20 x 5-8 cm (proportion of length to width very variable), base broadly rounded to broadly cuneate, apex short-acuminate, edges entire to rather strongly undulate. Largest pinnae usually at a wide angle to rachis, 10-24 x3-7cm, widest about the middle or sometimes above the middle, base cuneate to rounded, apex abruptly caudate (cauda commonly to 2cm long), edges entire (sterile) or undulate (most fertile ones); costules 3.5mm apart (fertile) or 4.5mm (sterile); veins 12-15 pairs, slightly prominent on lower surface, where Sterile at 45° to costules, where fertile closer and almost at right angles to costules, excurrent veinlets all free; lower surface of rachis and costae sparsely to rather copiously hairy, most abundantly on young plants, hairs all hooked, similar hairs sometimes present on sur- face between veins; upper surface of rachis and costae bearing + abundant pale hooked hairs less than 0.5mm long. Sori medial, + elongate, those on connivent veins often uniting; sporangia usu- ally with 2 hooked hairs distally, hairs on stalk not seen; spores with rather narrow wing and a few cross-wings. Distr. Peninsular Thailand, in Malesia: Suma- tra, Malaya. Ecol. In primary forest at 0-800m, not on stream-banks. 55. Pronephrium = sulawesiense (K. IWATS.) HOLTTUM, sp. nov.—P. rubicundum _ ssp. sulawesiense K. IWATS. Acta Phytotax. Geobot. 28 (1977) 162, f. 2.— Type: K. SOMA et al. 218, 249, Central Celebes, inland from Polo, Kelawi Distr. 1000-1700 m (TUS, not seen). Differs from P. rubicundum as follows: scales on rhizome 4X 2.5mm, on stipe 5 x 1.8 mm, lack- ing hooked hairs; pinnae not always opposite, to 24 x 4.5 cm including apical cauda 4 cm long, more oblique to rachis, distal ones longest, apparently almost symmetric in shape, widest 1/3 from apex, edges of fertile pinnae strongly crenate-serrate distally, of sterile ones less strongly; sori medial, those on connivent veins not meeting; sporangia lacking hooked hairs. Distr. Malesia: Central Celebes, only known from the type. Notes. This is geographically remote from P. rubicundum and differs in scales, proportionately narrower more oblique more symmetrical pinnae with different edges. The figure shows a fertile frond with distinctly alternate pinnae. In frond- form this is nearer to P. ramosii of the Philippines but differs in other characters. 56. Pronephrium cuspidatum (BL.) HOLTTUM, Blumea 20 (1972) 123, excl. syn. praeter Bl. — Meniscium cuspidatum BL. En. Pl. Jav. (1828) 114; Fl. Jav. Fil. (1828) 102, t. 45; RActB. Fl. Btzg 1 (1898) 65. — Phegopteris cuspidata (BL.) METT. Farngatt. [TV (1858) 25; v.A.v.R. Handb. (1908) 511. — Dryopteris cuspidata (BL.) CHRIST, Philip. J. Sci. 2 (1907) Bot. 205, p.p.; BACKER & POSTH. Varenfl. Java (1939) 63, f. 10. — Abacopteris cus- pidata (BL.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 245.— Cyclosorus cuspidatus (BL.) CopEL. Gen. Fil. (1947) 142; Fern Fl. Philip. (1960) 375, nomen tantum.— Thelypteris cuspi- data (BL.) K. IwaTs. Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 192.—Type: BLUME, Java (L). Dryopteris amaiensis ROSENST. Meded. Rijksherb. n. 31 (1917) 6. — Type: HALLIER 3157, Java (L). Caudex short-creeping, 7-l10mm_ diameter, bearing hooked hairs. Stipe to 50 cm or more long, dark at base with sparse hooked hairs and firm dark brown scales c. 101mm bearing a few hooked hairs, above base light-castaneous, glabrous. Lamina commonly 30-5S0cm long (to 1981] THELYPTERIDACEAE (Holttum) 537 120 cm, fide BACKER & POSTHUMUS), firm, dry- ing dull reddish; apical segment of similar size to upper pinnae, base cuneate and slightly asym- metric; pinnae commonly to 6 pairs (10 pairs on type), lowest usually opposite, the rest often not so, a bud always at base of highest pinna only of old fronds; basal pinnae with stalks to 5 mm long, slightly asymmetric on either side of costa, otherwise like the rest. Pinnae commonly to 16 x 3.5cm, sometimes dimorphous, smallest fertile ones 9X 2cm, largest sterile 20 x 6cm, symmetric on either side of costa, usually widest 1/4-1/3 from base, base narrowly to broadly cuneate, slightly asymmetric with the acroscopic edge decurrent to junction with rachis, the basiscopic edge less decurrent; apex caudate-acuminate to abruptly cuspidate (narrow tip 1.5—2.5cm long); edges entire or nearly so; costules 3.5—-4 mm apart, at 45° to costa, grooved above when dry and slightly prominent beneath; veins 10-12 pairs, not prominent either side, dark on lower surface, at a narrower angle to costules in sterile than in fertile fronds, excurrent veinlets mostly free and ending in a conspicuous hydathode on upper surface (with white incrustation on old fronds); lower surface glabrous throughout or (especially in Sumatra) with a few hooked hairs on costae and costules; upper surface quite glabrous, even in groove of rachis. Sori near costules, somewhat elongate, those: on connivent veins near costa uniting, distal ones sometimes not; sporangia lacking hooked hairs on Java specimens, such hairs present on specimens from Sumatra. Distr. Malesia: Sumatra, Borneo, Java and Lesser Sunda Is. (Flores). Ecol. On rocks or steep valley-sides in forest at altitudes to 1200 m. Note. There may be some introgressive hybri- dization between this species and P. rubicundum in Sumatra. 57. Pronephrium ramosii (CHRIST) HOLTTUM, comb. nov. — Dryopteris ramosii CHRIST, Philip. J. Sci. 2 (1907) Bot. 203.— Lectotype (selected here): RAMOS BS 1792, Luzon, Rizal Prov. (P; FI, K). Cyclosorus cuspidatus sensu COPEL. Fern FI. Philip. (1960) 375, excl. syn. Meniscium cuspida- tum BL. — Fig. 16e-f. a. var. ramosii. Differs from P. cuspidatum as follows: scales at base of stipe to 4 x 1 mm, lacking hooked hairs; pinnae always alternate, including basal ones, in almost all cases widest above the middle and narrowly cuneate at base (exceptions are speci- mens from Mindoro and Sibuyan), commonly c. 11 x2.5cm, buds present at the bases of several pinnae (to 4 on each side of rachis); veins 6-8 pairs, no conspicuous hydathodes present at apices of excurrent veinlets; hooked hairs usually more abundant on lower surface of costae and costules, often present between veins on lower surface, present also in groove of upper surface of costae and in groove of rachis at base of a costa; sori medial; sporangia always bearing hooked hairs. Distr. Malesia: Philippines (Luzon to Min- danao), Moluccas (Halmahera), N.E. New Guinea. Ecol. “Common at 600-800 m along streams in damp gulches” (COPELAND). b. var. minahassae HOLTTUM, var. nov. A var. ramosii differt: pinnis usque 23x4cm (steriles), 18 X 3.5 cm (fertiles); pilis hamatis sub- tus inter venas carentibus ; sporangiis pilis hamatis non praeditis. —Type: ALSTON 16551, N. Celebes, G. Manembo-nembo (BM). Distr. Malesia: N. Celebes. Note. Meniscium liukiuense CHRIST, which I listed in the synonymy of P. cuspidatum in 1972, should also be included in P. ramosii, perhaps as a distinct variety; its excurrent veins are mostly not free. In the Solomon Islands is another possibly distinct variety which has no hooked hairs on sporangia and lower surface; they occur only at the base of costae on the upper surface. 19. NANNOTHELYPTERIS HOLTTUM, Blumea 19 (1971) 38; Kalikasan 2 (1973) 64. — Fig. 17. Caudex short-creeping; fronds somewhat dimorphous, sterile ones with shorter stipes and larger pinnae than fertile; pinnae 16-35 pairs, not more than 2.5 cm long, close together except in some fertile fronds, lower pinnae more widely spaced but hardly reduced; veins free or the basal ones anastomosing; lower surface of pinnae between veins + pustular when dry, not glandular, in some cases with minute acicular hairs; upper surface between veins glabrous or with minute erect acicular hairs; sori indusiate or exindusiate; indusia bearing hairs or glands; sporangia bearing spherical 538 FLORA MALESIANA [ser. II, vol. 1° yellow glands, as in Sphaerostephanos; spores with a + some Cross-wings. complete wing and Type species: Nannothelypteris aoristisora (HARR.) HOLTTUM. Distr. Malesia: Philippines (Luzon, Panay, Samar, Mindanao). Ecol. In forest, in some cases on rocks. Cytol. n= 36 (M. G. PRICE, N. camarinensis). Taxon. A natural group of species closely related to Pronephrium sect. Dimorphopteris, with which it might be united. No transference is here made to Pronephrium because of the uncertain status of the latter. COPELAND’s treatment of these species in his Fern Flora of the Philippines is confused because he did not examine details. Some earlier confusion was caused by the inclusion by CUMING of some specimens of Sphaerostephanos lastreoides in his n. 251, nothelypteris philippina. together with the type collection of Nan- KEY *tO THE SPECIES 1. One pair of veins regularly anastomosing, at least near bases of larger pinnae; short hairs present between veins on upper surface. 2. Sori exindusiate; pinnae less than 2 cm long 2. Sori indusiate; pinnae to 3 cm long 1. N. aoristisora . 2. N. camarinensis Veins all free, or with casual slight anastomosis; upper surface between veins glabrous, or nearly so. S Pinnae to 1.3cm long, almost entire; costules mostly forked, sometimes anastomosing the branches of adjacent ones 3. N. nervosa 3. Pinnae (at least sterile ones) to 2cm or more e long, distinctly lobed at least o on acroscopic side; veins all free. 4. Sterile pinnae to 2.5 x 1.0cm, lobed equally both sides half-way to costa or more deeply, not conspicuously auricled; indusia setose 4. N. philippina 4. Sterile pinnae usually not over 2 cm long, 5-6 mm n wide above the auricled base, lobed more deeply on acroscopic than on basiscopic side; indusia glandular 1. Nannothelypteris aoristisora (HARR.) HOLTTUM, Blumea 19 (1971) 38; Kalikasan 2 (1973) 65.— Polypodium aoristisorum HARR. J. Linn. Soc. Bot. 16 (1897) 30.— Phegopteris aoristisora (HARR.) v.A.v.R. Handb. (1908) 501.— Cyclo- sorus aoristisorus (HARR.) COPEL. Gen. Fil. (1947) 142; Fern Fl. Philip. (1960) 355, p.p.— Thelypteris aoristisora (HARR.) REED, Phytolo- gia 17 (1968) 260.—Type: STEERE, Panay (MICH; K). Stipe 3-12 cm long (longest in fertile fronds), short-hairy; lamina to 20cm long; pinnae to 25 pairs. Sterile pinnae to 1.7 X0.6cm; base slightly to distinctly auricled on acroscopic side, rounded on basiscopic; apex obtusely pointed to rounded; edges crenate to lobed, more deeply on acroscopic than on basiscopic side; costules 2.5mm apart; veins 2-3 pairs, basal pair anastomosing except near apex of pinna; lower surface of rachis den- sely short-hairy, hairs more sparse on costules and veins; upper surface of rachis with thicker and more sparse hairs, surface of pinnae between veins bearing scattered very short erect hairs. Fertile pinnae 1.1 x 0.35 cm; sori at maturity filling lower surface, exindusiate. Distr. Malesia: Philippines (Panay; Luzon: Mt Makiling). 2. Nannothelypteris camarinensis HOLTTUM, Kal- 5. N. inaequilobata ikasan 5 (1976) 119.— Type: M. G. Price 3101, Luzon, Camarines Norte, Basud, Cone Mt (PNH; K). — Fig. 17e-f. Sterile fronds of young plants: stipe 3-6cm long; lamina 15cm long with 10-16 pairs of pin- nae; pinnae to 2.0cm long, 0.4cm wide above a rounded basiscopic auricle, edges subentire, apex rounded. Larger fronds, partially fertile: stipe S— 10cm long; pinnaé to 2.5 x 0.7 cm, basal auricles smaller; apex more narrowly obtuse; edges stron- gly crenate; costules 3mm apart at 45° to costa; veins to 4 pairs, basal pair anastomosing; lower surface of rachis bearing copious pale hairs 0.3mm long, shorter hairs on costae with scat- tered thick ones 0.5mm long, minute hairs on surface between veins; upper surface of rachis with more sparse and thicker hairs, whole surface of pinna bearing abundant short erect hairs. Fully fertile fronds: stipe slender, to 22 cm long; pinnae to 1.5x0.5cm; sori medial, slightly elongate on the veins, with short-hairy indusia and glandular sporangia. Distr. Malesia: known from the type. Ecol. On forested rocky slope at 250-350 m. Philippines (Luzon), only 3. Nannothelypteris nervosa (FEE) HOLTTUM, Kalikasan 2 (1973) 66.— Phegopteris nervosa FEE, Gen. Fil. (1852) 244; 6e Mém. (1853) 13, t. 2, 1981] THELYPTERIDACEAE (Holttum) 539 NAA sin SS \y AM TARR AN VN ae oe EID) Fig. 17. Nannothelypteris nervosa (FEE) HOLT- TUM. a. Frond, <3; b. pinna, x 3.—N. philippina (PR.) HOLTTUM. c. Pinna, x 2.—N. inaequilo- bata HoxLttum. d. Sterile pinna, x2.—N. camarinensis HOLTTUM. e. Pinna, x2; f. sorus, x 24 (a-b PRICE & HERNAEZ 71, c isotype K, d holotype, e-f holotype). f. 4.— Aspidium exiguum KUNZE ex METT. Farngatt. IV (1858) 76, var. a. — Lastrea nervosa (FEE) COPEL. Philip. J. Sci. 81 (1952) 32: Fern FI. Philip. (1960) 323, nomen tantum. — Type: Cum- ING S.n., Philippines (FEE’s specimen not seen: specimens at B, BM, G, K agree with FEe’s figure and are perhaps isotypes). Polystichum —auriculatum var. — nervosum CHRIST, Bull. Herb. Boiss. 6 (1898) 192: v.A.v.R. Handb. (1908) 165. Cyclosorus aoristisorus sensu COPEL. Fern. FI. Philip. (1960) 355, p.p. — Fig. 17a—b. Fronds almost uniform, but stipe of sterile ones 7-10 cm long, fertile 15 cm. Lamina to 24 cm long; pinnae 30-40 pairs. Largest pinnae 1.3cm long, 3-3.5mm _ wide above the dilated base which is 44.5 mm wide (base more auricled on acroscopic side in fertile than in sterile pinnae); apex obtuse; edges entire or slightly crenate distally; costules 1.5mm apart, simply forked or with one branch forked again, in basal acroscopic lobe pinnate with 2-3 pairs of veins; veins free or with casual anas- tomosis; lower surface of rachis densely short- hairy, hairs sparse on costae and costules; upper surface of rachis with more sparse and shorter hairs than lower, sparse minute hairs on costae. Sori near margins of pinnae, on acroscopic branch of each costule; indusia small, hairy. Distr. Malesia: Philippines Samar), at low altitudes. Notes. The name Lastrea exigua J. SM., on which Aspidium exiguum KUNZE ex METT. was based, was published (without description) with citation of CUMING 251 and 272, types of N. philippina and N. inaequilobata; Mettenius cited P. nervosa FEE under his var. a. If transferred to Thelypteris, N. nervosa will need another specific epithet, as T. nervosa (KLOTZSCH) TRYON represents another species. (Mindanao, 4. Nannothelypteris philippina (PRESL) HoLt- TUM, Kalikasan 2 (1973) 66.— Physematium philippinum PRESL, Epim. Bot. (1851) 192: HOLTTUM, Novit. Bot. Univ. Carol. Prag. 1968 (1969) 44, excl. syn. Phegopteris nervosa FEE. — Thelypteris philippina (PRESL) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 283. — Type: CUMING 251, Luzon, S. Ilocos (PRC; G, K, P, US). Dryopteris confusa CopEL. Philip. J. Sci. 8 (1911) Bot. 146. — Lastrea confusa COPEL. Fern Fl. Philip. (1960) 324.— Thelypteris confusa (COPEL.) REED, Phytologia 17 (1968) 268.— Type: CUMING 251 partim (M, destroyed); neo- type (HOLTTUM 1973): RAMOs BS 8271 (MICH; US). Aspidium exiguum KUNZE ex METT. Farngatt. IV (1858) 76, var. b, p.p.— Dryopteris exigua sensu v.A.v.R. Handb. (1908) 190. — Fig. 17c. Fertile and sterile fronds almost alike. Stipe c. 20 cm long, pale, copiously short-hairy. Lamina to 540 30cm long; pinnae to 20 pairs, lower ones more widely spaced and slightly reduced. Largest pin- nae 2.5 1.0cm, gradually narrowed from trun- cate base to abruptly obtuse or rounded apex, lobed equally on both sides 1/3-1/2 towards costa; lobes oblique, basal acroscopic one often slightly enlarged; costules 2.5—-3 mm apart; veins 3-4 pairs, basal acroscopic one ending at base of sinus, basiscopic one passing to margin above base of sinus; lower surface of rachis bearing coarse pale hairs 0.3-0.5 mm long, hairs on costae very short with a few longer ones, sparse on costules; upper surface of rachis as lower, hairs on costae minute. Sori inframedial; indusia small, setiferous: sporangia bearing glands. Distr. Malesia: Philippines (Luzon). Note. In some herbaria specimens of CUMING 251 and 272 are mounted without distinction on the same sheet. The number 251 here refers to the specimens at Prague and elsewhere as indicated, but at BM the specimen of the present species bears the number 272. 5. Nannothelypteris inaequilobata HOLTTUM, Kalikasan 2 (1973) 67.—N. nervosa sensu HOLTTuUM, Blumea 19 (1971) 38, p.p.— Type: FLORA MALESIANA [ser. II, vol. 1° CUMING 272, Luzon (K; B, E, G, L, P). Aspidium exiguum KUNZE ex METT. Farngatt. IV (1858) 76, var. b, p.p. — Fig. 17d. Fronds distinctly dimorphous; pubescence as N. philippina. Sterile fronds: stipe 5S-9cm_ long; lamina to 20cm long, pinnae 25 pairs or more, lower 3-4 pairs not reduced but more widely spaced, lowest with free basal acroscopic lobe; largest pinnae 1.5-2.0cm long, 0.5—0.6cm wide above base, base rather narrowly rounded, on basiscopic side, auricled on acroscopic, basiscopic margin slightly crenate distally, acroscopic margin deeply crenate; costules 2 mm apart, very oblique, veins 2 pairs (3-4 pairs on basal auricle), basal acroscopic vein ending at base of sinus. Fertile fronds: stipe to 15 cm long; lamina 20-30 cm long, pinnae to 30 pairs, lower ones more widely spaced than in sterile fronds; largest pinnae 1.0-1.3cm long, to 0.4mm wide above base, on acroscopic side lobed 1/2 way to costa, costules less than 2mm apart; sori inframedial, indusia bearing glands, not hairs. Distr. Malesia: Philippines (Luzon: Mt Makil- ing and Nueva Vizcaya Province). Ecol. On Mt Makiling at 300m, near mud- springs in open place in forest. 20. STEGNOGRAMMA BLUME, En. Pl. Jav. (1828) 172; CHING, Sinensia 7 (1936) 90; Acta Phy- totax. Sinica 8 (1963) 329; COPEL. Gen. Fil. (1947) 144; K. IwaTs. Acta Phytotax. Geobot. 19 (1963) 112; Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 18-21; Amer. Fern J. 54 (1964) 141; HoLTrumM, Blumea 19 (1971) 38. — Leptogramma J. Sm. in Hook. J. Bot. 4 (1841) 51; Hist. Fil. (1875) 231; CHING, Sinensia 7 (1936) 96. — Fig. 18. Caudex short-creeping or erect; stipes densely hairy, hairs unicellular (pluricellular in sect. Haplogramma, not Malesian); scales bearing uni- cellular acicular hairs on edges and surface; fronds simply pinnate; apical lamina always deltoid and deeply lobed with basal basiscopic veins of some of the middle or lower lobes springing directly from the rachis; pinnae subentire to deeply lobed, upper ones always + adnate to rachis, in some species only the lowest fully free; lowest pinnae not or little reduced; spherical glands lacking; hairs between veins on upper surface few, rather long, not appressed; veins free or anastomosing; sori exindusiate, extending along the veins (where veins anastomose, sori also along the intermediate excurrent veins between costules); sporangia setiferous; spores with many small wings. Type species: Stegnogramma aspidioides BL. Distr. Pantropic (excluding New Guinea and the Pacific) and subtropic, c. 18 spp.; in Malesia on mountains in Sumatra, Java, Borneo, Celebes, Luzon, Mindanao. Cytol. Chromosome number 36 in sect. Leptogramma (diploid in Mexico, Ceylon and Japan; tetraploid in Madeira): in other sections unknown. Taxon. IWATSUKI’s arrangement of 1963 is here adopted. In it he recognized four sections, two of 1981] THELYPTERIDACEAE (Holttum) 541 Fig. 18. Stegnogramma aspidioides BL. a. Pinna, <1; b. part of fertile pinna showing extent of sori, x 3. — S. subcalcarata (v.A.v.R.) HOLTTUM. c. Pinna, x 1; d. venation and sori, * 6; e. part of sorus, x 24 (a—b T. LOBB s.n., c-e isotype K). which (Haplogramma and Dictyocline) do not occur in Malesia. The New World species (see IWATSUKI 1964) are not yet well studied. In Malesia, the widely-distributed species S. pozoi is represented by a variety in Java; the other plants of sect. Leptogramma, from four widely-separated small areas, are here regarded as representing separate species, but more material is needed to characterize them clearly; only S. dissitifolia has been well collected. Stegnogramma is related to those species of Sphaerostephanos which have no spherical glands, but is distinct in the combination of characters mentioned in the generic description. In the 19th century, species of this genus were usually placed in Gymnogramme. The peculiar Hawatian species Pneumatopteris sandwicensis (BRACK.) HOLTTUM shares with Stegnogramma the characters of venation of the apical lamina and few thick hairs between veins on the upper surface of pinnae, and has somewhat elongate sori, but differs in sporangia and spores. KEY TO THE SPECIES 1. Veins anastomosing (sect. Stegnogramma) .......... .. . . . 41.8. aspidioides 1. Veins free (sect. Leptogramma). 2. Pinnae to 7.5 1.6cm or yeti lower surface of costae bearing many hairs, short ones more abundant than longer . . . .2. 8. pozoi var. petiolata 2. Pinnae, if over 4cm long, not more than if cm . wide: lower surface of costae bearing rather sparse hairs 0.5 mm or more long and a few short ones. 3. Pinnae not over 1.5 cm long, only basal ones lobed; sporangia sparsely setose 3. S. gymnocarpa 3. Pinnae longer, several pairs lobed; sporangia all freely setose. 4. Pinnae to 5.0 x 0.8 cm with narrowly triangular apex 1|O0mmlong .. . . . 4.8. subcalcarata 4. Pinnae proportionately wider with short entire apex. 5. Fronds to 30 cm long; basal pinnae lobed more than half-way towards costa .5. S. dissitifolia 5. Fronds c. 16cm long; basal pinnae less deeply lobed 6. S. celebica 1. Stegnogramma aspidioides BL. En. Pl. Jav. KAULF. 1824 nec DESv. 1827 nec BL. 1828; Ic. (1828) 173.— Gymnogramme stegnogramma BL. PI. 10 (1854) t. 950; Spec. Fil. 5 (1864) 150, excel. pl. Fl. Jav. Fil. (1829) 98, t. 44.—Gymnogramme Khasya & Ceylon; RACIB. FI. Btzg | (1898) 70. — aspidioides HooK. Gen. Fil. (1841) t. 120B non Phegopteris stegnogramma METT. Fil. Hort. Lips. 542 (1856) 84; v.A.v.R. Handb. (1908) 508. — Dry- opteris stegnogramma (BL.) C. CHR. Ind. Fil. (1905) 294, p.p.; Gard. Bull. S. S. 7 (1934) 250; BACKER & POSTH. Varenfl. Java (1939) 63.— Thelypteris stegnogramma (BL.) REED, Phy- tologia 17 (1968) 466. — Type: BLUME, Java. Phegopteris stegnogramma var. meniscioides v.A.v.R. Bull. Jard. Bot. Btzg II, 16 (1914) 27; Handb. Suppl. (1917) 319.—Type: MATTHEW 584a, G. Singgalang, Sumatra (BO). — Fig. 18a—b. Stipe to 40cm long, dark, bearing long pale hairs mixed with short ones; basal scales c. 6 Imm; hairs on rachis to 2mm long. Lamina commonly 55cm long; pinnae 8-10 pairs, adnate to rachis except basal 2-3 pairs; lowest pinnae slightly reduced, narrowed to base on basiscopic side and slightly auricled on acroscopic, veins in auricle forked and anastomosing; texture thin, dark when dried. Largest pinnae commonly 10 x 3cm (to 133.5cm); aerophores swollen; base truncate to subcordate; apex short-acuminate; edges crenate; costules 5mm apart, at a wide angle to costa; veins 8 pairs, 3 basal pairs anas- tomosing with zig-zag excurrent veins, next 2-3 pairs to sides of a long sinus-membrane; lower surface of costa bearing copious long spreading hairs mixed with shorter ones, costules and veins with shorter spreading hairs; rather sparse short erect hairs on surface between veins; copious antrorse hairs on upper surface of costae, shorter spreading hairs on costules and veins, sparse erect hairs between veins. Sori spread all along lower veins and also on the excurrent intermediate veins arising from them, usually on basal part only of distal veins. Distr. Malesia: West and Central Java; Suma- tra (G. Kemiri, G. Singgalang, G. Kerinci); Sabah (Mt Kinabalu). Wrongly recorded from Khasya and Ceylon by HOOKER and BEDDOME. 2. Stegnogramma pozoi (LAGASCA) K. IWATsS. Acta Phytotax. Geobot. 19 (1963) 124, var. petiolata (CHING) HOLTTUM, comb. nov. — Lep- togramma petiolata CHING, Acta Phytotax. Sinica 8 (1963) 319.—Type: G. WALL (“WAL- LICH’’), Ceylon (PE). Gymnogramme aspidioides BL. En. Pl. Jav. (1828) 112, non KAULF. 1824 nec DESv. 1827. — Gymnogramme totta sensu BL. Fl. Jav. Fil. (1829) 90, t. 38.— Grammitis blumeana PRESL, Tent. Pterid. (1836) 209, nom. nov. — Phegopteris totta sensu METT. Farngatt. IV (1858) 18, quoad pl. javan. tantum:; v.A.v.R. Handb. (1908) 497, p-p.; Handb. Suppl. (1917) 515, excl. var. subcalcarata. — Leptogramma totta sensu BEDD. Handb. (1883) 377, quoad pl. zeyl. tantum. — Dryopteris africana (DESV.) C. CHR. Ind. Fil. (1905) 251, p.p.; BACKER & PosTH. Varenfl. Java (1939) 36.— Type: BLUME, Waterfall Tjikundal, G. Gedeh, Java (L). Stipe 15-30cm long; basal scales c. 5X 1mm. FLORA MALESIANA [ser. Il, volawe Lamina 25-40cm long; pinnae 12-15 pairs of which 4-5 pairs are free, the rest + adnate to the rachis; basal 1-2 pairs in smaller fronds somewhat reduced, at least the lowest with stalks | mm long and reduced basal basiscopic lobe. Largest pinnae seen 7 1.6cm; base subequally truncate; apex entire, triangular, 8 x 3.5mm; edges lobed almost half-way to costa, lobes oblique, hardly falcate, entire; costules 4.5-S mm apart; veins 6 pairs (or 7 veins on basiscopic side of costule), basal veins from adjacent costules usually both touching sides of sinus-membrane which may be decurrent as a ridge between them; hairs on lower surface of costae abundant, spreading, of varied length, longest 0.6-0.7 mm, costules and veins with shor- ter hairs, copious erect short hairs between veins; hairs on upper surface of costae mostly not over 0.5mm long with some to 1mm, sparse shorter hairs on costules and veins, a few thick hairs 0.5mm long between veins. Sori on basal veins from costule along more than half length of vein, on distal veins shorter, medial. Distr. Ceylon; in Malesia: Java. Ecol. In Java on higher mountains, on steep earth banks in forest (BACKER & POSTHUMUS); in Ceylon at 1800 m. Notes. The plants of Java and Ceylon here included are closely related to S. pozoi, the type of which was collected in northern Spain. Similar plants found in Madeira are tetraploid (MANTON). The latter differ from var. petiolata (diploid) in having all pinnae except the lowest adnate to the rachis, hairs on lower surfaces, confined to costae and costules, uniformly | mm long, and basal sori not extending to the bases of veins. Plants named S. pozoi occur throughout Africa; they are vari- able and no chromosome counts have been made. BLUME’s type has pinnae to 4.0 x 1.3 cm, and all pinnae above the basal ones are adnate; BACKER & POSTHUMUS report plants with fronds to 50 x 25 cm but I have not seen any so large. Prof. CHING (in lit.) informs me that the type of Leptogramma petiolata bears the collector's name WALL, which he interpreted as WALLICH, but it was surely G. WALL, who collected ferns in Cey- lon, not WALLICH. 3. Stegnogramma gymnocarpa (COPEL.) K. IwatTs. Acta Phytotax. Geobot. 19 (1963) 122. — Dryopteris gymnocarpa COPEL. in Elmer, Leafi. Philip. Bot. 3 (1910) 807.— Phegopteris gym- nocarpa (COPEL.) v.A.v.R. Handb. Suppl. (1917) 313.— Lastrea gymnocarpa COPEL. Gen. Fil. (1947) 139; Fern Fl. Philip. (1960) 325, quoad typ. tantum. — Leptogramma gymnocarpa (COPEL.) CHING, Acta Phytotax. Sinica 8 (1963) 318.— Thelypteris gymnocarpa (COPEL.) MORTON, Amer. Fern J. 56 (1966) 179.—Type: ELMER 11508, Mt Apo, Falls of Cati Creek, 1750 m, Min- danao (MICH; BO, K, L etc.). Stipe 2-3cm long; hairs 1mm long rather 1981] THELYPTERIDACEAE (Holttum) 543 sparse, many much shorter; scales little over 1 mm long. Lamina 9-15 cm long (COPEL.), distal half or more pinnatifid, free or adnate pinnae 3-4 pairs, lowest with stalks 1mm long; basal pinnae 1.0 x 0.7 cm, distinctly lobed; other pinnae, and lobes of terminal lamina, entire. Veins in lobes of basal pinnae 2-3 pairs, in lobes of terminal lamina sim- ple or forked; lower surface of rachis bearing many hairs 1mm long and few short ones, on costae and costules more sparse similar hairs, no others; on upper surface. of rachis many short hairs and fewer | mm long, hairs on costae, cos- tules and between veins sparse, 0.5 mm long. Sori on basal acroscopic veins on pinnae, sometimes on both branches of a forked vein on apical lamina; a majority of sporangia lacking setae. Distr. Malesia: Philippines (Mindanao), only known from type. Note. COPELAND included the Luzon plants here named S. dissitifolia. 4. Stegnogramma subcalcarata (v.A.v.R.) HOLT- TUM, comb. nov. — Phegopteris totta var. sub- calcarata v.A.v.R. Handb. Suppl. (1917) 515.— Type: BECCARI 430, G. Singgalang, Sumatra, 1700 m (BO; FI, K, MEL). — Fig. 18c-e. Stipe 10-12cm long; hairs 1mm long sparse, with shorter ones; scales small, narrow. Lamina to 18cm long; free pinnae 4-5 pairs, above these 4-5 pairs gradually more broadly adnate to rachis; basal pinnae on some fronds reduced, in all cases narrowed towards base on both sides (more so on basiscopic), basal acroscopic lobe 2-3.5 mm long, almost free, stalks 1.5-2 mm long. Largest pinnae 3.5-5cm long, 6.5-8mm wide; base broadly cuneate; apex entire, triangular, 7-12 mm _ long, 2.5mm wide at base; edges lobed half-way to costa, lobes subtriangular; costules 3 mm apart, at less than 60° to costa; veins 4-5 pairs, acroscopic basal vein passing to side of short sinus-mem- brane, basal basiscopic vein to edge above base of sinus; hairs on lower surface of costa sparse, 0.7 mm long with a few shorter ones, very few on costules and veins, none between veins; on upper surface hairs few apart from costa. Sori on basal acroscopic vein almost from its base, medial on other veins. Distr. Malesia: Sumatra. Apart from type, known from one poor specimen from 2000m on G. Kerinci (BUNNEMEIJER 10480). 5. Stegnogramma dissitifolia HOLTTUM, - sp. nov. — Lastrea gymnocarpa sensu COPEL. Fern Fl. Philip. (1960) 325, p.p. Stipes usque 18 cm longus; lamina usque 30 cm longa; pinnae liberae usque 12-jugatae, inferiores 6-jugatae stipitatae, maximae 3.5 cm longae, basi dilatata usque 1.2cm lata, supra basin 0.9 cm latae, dimidio costam lobatae; pinnae infimae valde dissitae. —Type: M. G. PRICE 1668, Baguio City, Luzon (K). Stipes 10-18cm long, bearing sparse hairs I- 1.5mm long and shorter ones; basal scales to 5x1lmm. Lamina to 30cm long; pinnae c. 12 pairs below deeply lobed apical lamina, c. 6 pairs of lower pinnae distinctly stalked, stalks to 1.5mm; basal pinnae 2-3cm from next pair (in another collection 5 cm), wider on basiscopic side of costa than on acroscopic and lobed more than half-way to costa, much narrowed to base on basiscopic side. Largest pinnae 3.5cm_ long, commonly to 0.9cm wide above dilated base which is 1.0-1.2cm wide; base subtruncate to broadly cuneate; apex abruptly short-pointed; edges lobed about half-way to costa; costules 3-3.5(-4) mm apart; veins 3—4 pairs; on lower sur- face of rachis sparse hairs 1mm long and many much shorter, on costae sparse hairs 0.5 mm long and a few short ones, sparse slender erect hairs between veins; hairs on upper surface of rachis more than 1 mm long with many short ones, costal hairs 0.3-0.5mm, on costules fewer, scattered thick hairs 0.3-0.5 mm long between veins. Sori on basal half of basal veins, in middle of distal veins; sporangia all with several setae. Distr. Malesia: Philippines; mountains of middle-north of Luzon at 1400-2700 m (7 collec- tions). Notes. Young plants of these Luzon coliec- tions differ from the small plants of comparable size from Mindanao which constitute the type of S. gymnocarpa; the former have more pairs of free pinnae, which are all lobed, and sporangia all have more than one seta. Leptogramma amabilis TAGAWA, based on small plants from Okinawa (Ryukyu Is.) cannot be effectively compared until better known. Plants described as S. pozoi on Orchid Island (near the south of Taiwan) have pinnae 5 cm long. 6. Stegnogramma celebica (CHING) HOLTTUM, comb. nov.—Leptogramma celebica CHING, Sinensia 7 (1936) 99, t. 5.—S. gymnocarpa (COPEL.) K. IWATS. ssp. celebica K. IWATs. Acta Phytotax. Geobot. 19 (1963) 123. — Thelyp- teris bunnemeijeri REED, Phytologia 17 (1968) 265.— Type: BUNNEMEJJER 12073, G. Bonthain, Celebes (NY; BO, L). Stipe to 7cm long; hairs hardly 0.5 mm long. Lamina to 16cm long; free pinnae 5 pairs, 5 pairs adnate to rachis; lowest pinnae slightly reduced, smallest seen 1.3cm long, with stalks 1 mm long, narrowed to base on basiscopic side. Largest pinnae 3 cm long, 7-8 mm wide; base broadly and rather unequally cuneate; edges lobed, almost to apex, less than half-way to costa, lobes small, subtriangular; costules 2.5 mm apart, at little more than 45°; veins 3 pairs, basal acroscopic veins passing to side of sinus-membrane, basiscopic vein to edge above base of sinus (in sterile fronds, veins sometimes uniting just below sinus); hairs on lower surface of costae rather sparse, on fertile 544 pinnae hardly 0.5mm long, on sterile pinnae longer, few hairs on costules, none between veins on fertile pinnae (a few on sterile ones); a few hairs at least 0.5 mm long between veins on upper surface. Sori along basal half of basal veins, FLORA MALESIANA [ser. II, vol. 1° sometimes also on second veins. Distr. Malesia: S.W. Celebes. Known from two collections from about 2000m on G. Bon- thain; the second is EVERETT s.n. Oct. 1895 (K, SING). 21. AMPHINEURON HOLTTUM, Blumea 19 (1971) 45; Blumea 23 (1977) 205. — Fig. 19. Caudex erect, or short- or long-creeping; scales narrow, setiferous; stipe minutely hairy, scaly at the base only except in A. distinctum; lamina often very large, pinnate, pinnae in most species deeply lobed; basal pinnae much narrowed at their bases; in a few species 1-2 (rarely 3) pairs of irregularly- placed and -shaped small pinnae sometimes present below the normal ones; aerophores at the bases of pinnae usually narrowly elliptic and (when dry) discoloured, not swollen; veins pinnate in the pinna-lobes, simple, basal veins either free and passing to the margin separately, or connivent at the sinus-membrane, or anastomosing to form an excurrent vein, these con- ditions sometimes not constant in a single frond; sinus-membrane usually ending in a prominent tooth; short acicular hairs always present on some part of the lower surface, also glandular hairs of varying size and shape, spherical to pyriform or club-shaped, the larger ones sometimes collapsing in drying to form resinous spots which may be faint and hardly detectable; sori in most species medial or supramedial; indusia usually present, bearing glands and/or hairs, apparently lacking in A. distinctum; glandular cells often present on the stalks of sporangia, directly attached to the stalk or at the end of a short hair, capitate hairs seen on the body of sporangia only in A. subattenuatum; spores usually dark, irregularly rugose or with irregular thick + branched ridges. Type species: Amphineuron opulentum (KAULF.) HOLTTUM. Distr. S.E. Asia; Malesia; Queensland; in the Pacific to Tahiti; East Africa; in all c. 12 species. Cytol. Base chromosome number 36; A. opulentum (Singapore) and A. terminans (Ceylon) both tetraploid; no experimental work reported. Taxon. This genus resembles Christella in its spores and in its mainly elongate glandular hairs; the latter are more varied than in Christella. The name Amphineuron is intended to indicate the fact that in several species the anastomosis of veins is inconstant. The two most widely-distributed species, A. opulentum and A. terminans, are both variable, and both have a complex taxonomic and nomenclatural history. Hybrids between them probably exist, at least in Thailand. As indicated in the key, the species may be divided into two groups. Rather large glands, of varied form, occur on most species of the second group and are undoubtedly distinctive, but they are often not well preserved on herbarium specimens and are sometimes not detectable, so that such specimens are difficult to name with certainty, and I have found it impossible to be sure how many species can be recognized. The present arrangement is tentative. Specimens dried without heat retain their glands perfectly, as I have discovered when dealing with plants in cultivation at Kew. It may be that alcohol, sometimes used to preserve specimens temporarily, also has an effect on glands in this genus. KEY TO THE SPECIES 1. Indusia rather large, persistent; lamina thin, veins slender, slightly prominent on both sides. 2. Caudex long-creeping; pinnae lobed half-way to costa or less deeply; apex of frond + pinna-like 1. A. terminans 1981] THELYPTERIDACEAE (Holttum) 545 2. Caudex short-creeping or erect; pinna-like. pinnae lobed more than half-way to costa; apex of frond not 3. Pinnae lobed to | mm from costa or more deeply; basal veins free or uniting to form very short excurrent veins. 4. Basal veins not meeting; sori in slight depressions 2. A. immersum 4. Basal veins meeting just below the sinus, sometimes forming. a short excurrent vein; sori not immersed 3. A. subattenuatum 3. Pinnae lobed less deeply: ‘basal veins ; anastomosing to form an excurrent vein on most parts of a frond 4. A. opulentum . Indusia small, almost hidden by sporangia at maturity of s sorus, or “apparently lacking; lamina firm, veins not prominent on either surface. . Hairs more than 0.5 mm long present on both sides of costa; stipe and abaxial surface of rachis bearing scales throughout 5. All hairs on costa very short; scales confined to base of stipe. 5. A. distinctum 6. Lower surface between veins (sometimes in part) bearing very short erect acicular hairs; short capitate hairs also in this position on the upper surface 6. Both surfaces lacking hairs between veins. 6. A. attenuatum 7. Pinnae lobed to c. 2mm from costa, basal veins often anastomosing. 8. Many glandular hairs on lower surface between veins, some spherical, some + elongate; similar hairs also on indusia 7. A. ceramicum 8. Glandular hairs less abundant, always spherical tending to collapse and form resinous patches or to disappear on drying 8. A. pseudostenobasis 7. Pinnae lobed more deeply; basal veins not PAS eoroSite! 9. Pinnae thin, to 3cm wide; glands on lower surface of pinnae and on indusia elongate 9. A. paraphysophorum 9. Pinnae firm, to 1.5 cm wide; glands on lower surface glabular to pyriform, on indusia spherical, resinous 1. Amphineuron terminans (HOOK.) HOLTTUM, Amer. Fern J. 62 (1973) 82; Blumea 23 (1977) 207. — Nephrodium terminans HOOK. Spec. Fil. 4 (1862) 73, excl. syn. N. conioneuron FEE & Las- trea malaccensis PRESL. — Thelypteris terminans (HOOK.) TAGAWA & K. IwatTs. Acta Phytotax. Geobot. 26 (1975) 169.—Type: WALLICH 386, Burma, Kamoun (= Kumon) (K). Nephrodium oreopteris FEE, Gen. Fil. (1852) 306, non (EHRH.) DESv. 1827. — Type: CUMING 48, Luzon (holo?; isotypes G, K, LE, W). Thelypteris wagneri FosB. & SACHET, Smiths. Contr. Bot. 8 (1972) 6, excl. syn. Polypodium pteroides RETZ.— Type: RACIBORSKI, Java (US). Nephrodium pteroides sensu BEDD. Handb. (1883) 269; RActB. FI. Btzg 1 (1898) 183. — Dry- opteris pteroides sensu v.A.v.R. Handb. (1908) 209. Dryopteris interrupta sensu BACKER & POSTH. Varenfl. Java (1939) 56. — Cyclosorus interruptus sensu HOLTTUM, Rev. FI. Malaya 2 (1955) 262, f. 149; CopEL. Fern Fl. Philip. (1960) 361. — Fig. 19a. Caudex long-creeping, c. 5mm diameter (dry); stipe commonly to 50cm long, + flushed dull reddish, glabrescent abaxially, basal scales c. 8mm long. Lamina to 50cm long; pinnae to 25 pairs, basal pair somewhat reduced, rarely very small, always distinctly narrowed at their bases; apex of frond usually pinna-like but variable. Largest pinnae commonly 17-20 x 1.7-2.0cm, if longer not more than 2cm wide; base of middle . 10. A. kiauense pinnae broadly cuneate to truncate; apex acu- minate; edges lobed 1/3 towards costa or less deeply, lobes as wide as long (or wider) with broad asymmetric apex and forward-pointing tip; costules 4-5 mm apart, usually at less than 60° to costa; veins 6-9 pairs, basal pairs spreading at a broad angle to their costules and uniting to form a rather long excurrent vein to the sinus, next veins very oblique, 1 or 2 ending beside the sinus- membrane; lower surface of rachis, costae, cos- tules and veins bearing short acicular hairs, longer hairs usually lacking, subsessile almost spherical rather pale glandular hairs abundant on distal veins, usually few and smaller on lower veins, very short acicular hairs often present between veins; upper surface of costae bearing antrorse pale acicular hairs, similar hairs scattered on cos- tules and veins. Sori close to margins of lobes, not on lower veins; indusia large, thin, often with some short acicular hairs and a few small glan- dular hairs which are not marginal. Distr. Ceylon & S. India; Burma to Hainan and Macao; throughout Malesia; Queensland (to 18° S); one specimen from Central Africa and one from Fernando Poo. Ecol. In Malesia only abundant in areas with a distinct dry season, in rather open but not too dry places, spreading by long rhizomes. Notes. The complex nomenclatural and tax- onomic history of this species is set forth and discussed in HOLTTUM 1977. It is probable that in Thailand this species has become hybridized with A. opulentum but I have not seen intermediates 546 FLORA MALESIANA [ser. II, vol. 1° f Nn / Ne % Se PS y i A fy \ Se ee ane = 3 t n Fe SO TTY ES SEA poe Femnhsesunscteunere eaubneeh fubina tet asl ene) JCD Fig. 19. Amphineuron terminans (HOOK.) HOLTTUM. a. Venation and sori, xX 4.— A. opulentum (KAULF.) HOLTTUM. b. Two pinna-lobes, showing differences in course of basal veins, x 3; c. sorus, x 18.— A. subattenuatum (ROSENST.) HOLTTUM. d. Venation and sori, X 3.— A. immersum (BL.) HOLTTUM. e. Venation and sori, x 3.— A. pseudostenobasis (COPEL.) HOLTTUM. f. Base of basal pinna, X 1; g. two pinna-lobes, = 4; h. upper surface of costa with capitate hairs, x 16 (a FORMAN 145, b-c SEEMANN S.n., d WOMERSLEY & HOLTTUM 17692, e MOUSSET 39, f-h HOOGLAND & CRAVEN 10122). 1981] THELYPTERIDACEAE (Holttum) 547 from Malesia. FOSBERG and SACHET used the pinna-like apex of fronds as the main distinguishing character, ignoring the evidence of venation, hairs and glands which appear to me more significant. This is the only species of Amphineuron in which anastomosis of veins is invariable. 2. Amphineuron immersum (BL.) HOLTTUM in Nayar & Kaur, Comp. to Bedd. (1974) 203; Blu- mea 23 (1977) 211.— Aspidium immersum BL. En. Pl. Jav. (1828) 156; RActrB. FI. Btzg 1 (1898) 169. — Lastrea immersa (BL.) MOorRE, Ind. Fil. (1857) Ixxxix; BEDD. Ferns Br. India (1867) t. 252; Handb. (1883) 234; COPEL. Fern FI. Philip. (1960) 327. — Dryopteris immersa (BL.) O. KTZE, Rev. Gen. Pl. 2 (1891) 813; v.A.v.R. Handb. (1908) 188; BACKER & POSTH. Varenfl. Java (1939) 39.— Thelypteris immersa (BL.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 6 (1936) 306; HOLTTUM, Rev. Fl. Malaya 2 (1955) 243. — Parathelypteris immersa (BL.) CHING, Acta Phytotax. Sinica 9 (1963) 303. — Type: BLUME, Java (L, n. 908, 335- 404). Lastrea caudiculata PRESL, Epim. Bot. (1851) 36; HoLTTUM, Novit. Bot. Univ. Carol. Prag. 1968 (1969) 35. —Type: CUMING s.n. Philippines (PRC). Lastrea verrucosa PRESL, Epim. Bot. (1851) 36; CoPEL. Fern FI. Philip. (1960) 327. — Thelypteris verrucosa (PRESL) CHING, Bull. Fan Mem. Inst. Biol. Bot. 6 (1936) 308.— Type: CUMING 72, Luzon (PRC). ?Dryopteris diversifolia v.A.v.R. Bull. Dép. Agr. Ind. Néerl. 18 (1908) 7; Handb. (1908) 189. — Type: RAAP 27, Sumatra, Batu Isl. (BO). Dryopteris besukiensis v.A.v.R. Bull. Jard. Bot. Btzg II, 1 (1911) 7; Handb. Suppl. (1917) 156. — Lectotype (HOLTTUM 1977): KOORDERS 15436, Java, Besoeki (BO; L). Thelypteris subimmersa CHING, Bull. Fan Mem. Inst. Biol. Bot. 6 (1936) 306. — Parathelyp- teris subimmersa CHING, Acta Phytotax. Sinica 8 (1963) 303. — Type: S. K. LAu 1395, Hainan (PE, not seen). — Fig. 19e. Caudex short, erect; stipe to 70cm or more long, green when living; basal scales thin, to 15 mm long. Lamina to at least 120 cm long (often fertile at a much smaller size); pinnae close, tex- ture thin, drying pale-olivaceous; 1-2 pairs of irregularly spaced small pinnae sometimes present at base of frond. Largest pinnae of type 25 x 3. cm, largest seen-45 x 5cm, lobed to 1 mm from costa or more deeply; lobes, except distally, almost at right angles to costa, separated by wide sinuses; costules 3.5-6mm apart; veins 14-20 pairs, basal acroscopic vein ending beside the short sinus- membrane, basiscopic one passing to margin above base of sinus; lower surface of costae and costules of type lacking acicular hairs except near apex of pinna, of other specimens bearing a vari- able number of hairs up to 1 mm long, costules and veins usually bearing small pale yellow glands (abundant on the type), between veins sometimes short erect acicular and capitate hairs; upper sur- face with antrorse hairs on costae and scattered hairs on costules and veins. Sori supramedial, in depressions in the lamina (forming convexities on the upper surface); indusia thin with a variable fringe of short yellow capitate hairs. Distr. Assam; Hainan; southern Thailand; throughout Malesia; Queensland (to 16°S); New Hebrides, New Caledonia. Ecol. At low altitudes, in sheltered places but not in deep shade, common on limestone in Malaya, found also on stream banks. Notes. All the living plants observed by me have an erect caudex, but BACKER & POST- HUMUS state ‘‘wortelstok kruipend”. ERYL SMITH 2432, from Timor, is a small plant with definitely creeping caudex; it might be a hybrid with A. opulentum. The types of Dryopteris diversifolia v.A.v.R. and D. besukiensis v.A.v.R. are small plants, perhaps stunted owing to exposure. There is much variation in_ the development of acicular hairs and of small yel- lowish glandular hairs on the lower surface of pinnae. The types of both Lastrea caudiculata PRESL and L. verrucosa PRESL have acicular hairs, that of Thelypteris subimmersa CHING has none, but the published description gives no other character which would distinguish it from the type of Aspidium immersum BL. I know of no other record of this species in China. 3. Amphineuron subattenuatum (ROSENST.) HOLTTuM, Blumea 23 (1977) 412.— Dryopteris subattenuata ROSENST. in Fedde, Rep. 10 (1912) 332.— Thelypteris subattenuata (ROSENST.) REED, Phytologia 17 (1968) 317.— Type: BAm- LER 37, N.E. New Guinea, Logaueng, 300 m (S- PA?; BM). — Fig. 19d. Caudex short, erect; stipe to at least 50 cm long, pale, basal scales c. 10 1.5mm. Lamina 150cm or more long; pinnae well-spaced, the lowest with very narrow bases bearing auricles 5 mm long on both sides; 1-3 pairs of much-reduced pinnae, similar auricled, present below normal ones. Lar- gest pinnae 35cm long, 2.5-3.5 cm wide, lobed to 1-1.5mm from costa, lobes separated by wide sinuses and almost at right angles to the costa, not falcate, tips broadly rounded; costules to 6mm apart; veins to 20 pairs, basal veins spreading at a wide angle, their tips usually touching the sides of a short sinus-membrane or sometimes uniting to form a very short excurrent vein; lower surface bearing a variable number of very small colourless spherical glands, sometimes also very short aci- cular hairs; upper surface of costae bearing pale acicular hairs 0.6 mm long, few hairs on costules, between veins a variable number of short acicular and capitate hairs. Sori a little supramedial, not impressed; indusia bearing very small glandular 548 hairs; sporangia sometimes with a small capitate hair; spores pale, with highly prominent thick ridges. Distr. Malesia: Eastern New Guinea (6 collec- tions). Ecol. In somewhat exposed places near streams in forest. 4. Amphineuron opulentum (KAULF.) HOLTTUM, Blumea 19 (1971) 45; Blumea 23 (1977) 212. — Aspidium opulentum KAULF. Enum. Fil. Chamisso (1824) 238.— Thelypteris opulenta (KAULF.) FOSBERG, Smiths. Contr. Bot. 8 (1972) 3, excl. syn. Aspidium terminans WALL. — Type: CHAMISSO, Guam (LE). Nephrodium impressum DESv. Mém. Soc. Linn. Paris 6 (1827) 259.— Dryopteris impressa (DESvV.) POSTH. Verh. K. Akad. Wet. Amst. 36, 5 (1937) 14; BACKER & POSTH. Varenfl. Java (1939) 57.— Thelypteris impressa (DESV.) REED, Phy- tologia 17 (1968) 284. — Type: collector not cited, Timor (P). Aspidium extensum BL. En. Pl. Jav. (1828) 156. — Nephrodium extensum (BL.) MOORE, Ind. Fil. (1858) 91; BEDpb. Handb. (1883) 269. — Dry- opteris extensa (BL.) O. KTZE. Rev. Gen. Pl. 2 (1891) 812; v.A.v.R. Handb. (1908) 210. — Cyclo- sorus extensus (BL.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 182; HoLTTuM, Rev. FI. Malaya 2 (1955) 264, f. 150; COPEL. Fern FI. Philip. (1960) 368.— Thelypteris extensa (BL.) Morton, Amer. Fern J. 49 (1959) 113. — Type: no collector cited, Pulu Pinang (L). Lastrea malaccensis PRESL, Epim. Bot. (1851) 35. — Type: CUMING 391, Malacca (PRC; K). Nephrodium conioneuron FEE, Gen. Fil. (1852) 308. — Aspidium conioneuron (FEE) METT. Farn- gatt. IV (1858) 102.— Type: GARDNER, Ceylon (Isotype W). Aspidium nephrodioides Hook. Spec. Fil. 4 (1862) 162, t. 235, non KLOTZSCH 1847. — Aspi- dium hookeri BAK. Syn. Fil. (1867) 257, nom. nov. non KLOTZSCH 1847 — Dryopteris orbicularis C. Cure. Ind. Fil. (1905) 281, nom. nov. — Thelypteris orbicularis (C. CHR.) REED, Phytologia 17 (1968) 299.— Type: SEEMANN, Indian Archipelago (K). — Fig. 19b-c. Caudex short-creeping; stipe to 70cm long, rufescent, basal scales 10mm long, hardly 1mm wide. Lamina to 80 cm long; pinnae to 25 pairs or more; a pair of much-reduced basal pinnae some- times present; apex of frond narrowly acuminate, deeply lobed in its basal part and grading into the upper pinnae. Largest pinnae commonly to 25 x 2.5cm, largest seen 403.5cm, lobed 3/5-3/4 towards costa; lobes slightly oblique, slightly fal- cate; costules commonly 4mm apart, on large sterile fronds to 6 mm; veins 8-10(-15) pairs, basal pair both touching sinus-membrane or meeting below it at a varying angle to produce an excur- rent vein; lower surface of rachis, costae and FLORA MALESIANA (ser. II, vol. 1° costules bearing very short acicular hairs with scattered longer ones (to 0.5mm) on costules, veins, sinus-membranes and margin, small pale yellowish glands variously distributed along cos- tules and veins, surface between veins usually bearing some short erect acicular hairs and small colourless to yellowish capitate hairs; upper sur- face of costae covered with pale acicular hairs, similar hairs scattered on costules and veins. Sori confined to lobes of pinnae, supramedial, in slight depressions; indusia thin, shrivelled when old, bearing marginal yellow glandular hairs and sometimes a few acicular hairs which are not marginal; an elongate gland sometimes present on stalks of sporangia. Distr. East Africa; Seychelles; S. India and Ceylon; Burma, Thailand; Malesia; N. Queensland; islands of the Pacific to Tahiti; naturalized at various places in tropical America. Ecol. Few records; apparently adapted to semi-exposed situations among rocks, especially in areas with a dry season; in S.E. New Guinea (dry season area) found in secondary swamp- forest. Notes. A fuller synonymy, and commentary on it, is given in HOLTTUM 1977. METTENIUS pub- lished the first good description (as Aspidium conioneuron) with a note on the diversity of venation. The species seems not to be common in most parts of Malesia (doubtfully native in Java, few specimens from Borneo and the Philippines). Plants have long been cultivated in Singapore (origin unrecorded) and occur spontaneously on the edges of drains and elsewhere near the Botanic Garden; these plants are tetraploid. The center of distribution of the species is uncertain; possibly southern India. 5. Amphineuron distinctum (COPEL.) HOLTTUM, Blumea 23 (1977) 215.—Dryopteris distincta CoPEL. Univ. Cal. Publ. Bot. 18 (1942) 220.— Cyclosorus distinctus (COPEL.) COPEL. Gen. Fil. (1947) 142; Philip. J. Sci. 78 (1951) 444, pl. 26. — Thelypteris distincta (COPEL.) REED, Phytologia 17 (1968) 273. — Type: BAMLER W.11, N.E. New Guinea, Wareo, 150m (UC). Dryopteris longissima _ var. ROSENST. Hedwigia 56 (1915) BAMLER 132, same locality (B). Caudex unknown; stipe incomplete, dark at basé, distally reddish, minutely hairy, bearing thin narrow scales throughout, basal ones 12 mm long; abaxial surface of rachis also bearing similar scales or their small wart-like bases. Size of lamina not known; basal pinnae narrowed in their basal 4cm, base 4mm wide. Largest pinnae 25 x 3cm, rather short-acuminate, lobed to 1.5-2 mm from costa, lobes slightly oblique and slightly fal- cate; costules 44.5 mm apart; veins to 18 pairs, basal pair anastomosing to produce an excurrent vein to the sinus or passing to sides of sinus- novoguineensis 351.— Type: 1981] THELYPTERIDACEAE (Holttum) 549 membrane; lower surface of rachis sparsely short- hairy, of costae bearing copious erect hairs more than 0.5mm long, fewer such hairs on costules and veins, between veins copious slender erect acicular hairs and small + yellowish capitate hairs; upper surface of costae bearing thick acicular hairs and small capitate ones, shorter hairs of both kinds present between veins. Sori inframedial, exindusiate; hairs on stalks of sporangia consis- ting of 3 cells, distal ones club-shaped, orange; spores dark with irregular thick ridges. Distr. Malesia: Papua New Guinea; only known from the type. 6. Amphineuron attenuatum (O. KTZE) HOLT- TUM, Blumea 23 (1977) 215. — Aspidium attenua- tum KUNZE ex METT. Farngatt. IV (1858) 96, non Sw. 1801.— Nephrodium attenuatum BAK. Syn. Fil. (1867) 263, non T. MOORE 1858. — Dryopteris attenuata O. KTZE, Rev. Gen. PI. 2 (1891) 812, nom. nov.; v.A.v.R. Handb. (1908) 184.— Dry- opteris stenobasis C. CHR. Ind. Fil. (1905) 294, nom. nov. superfl.— Thelypteris stenobasis (C. CHR.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 254. — Mesoneuron attenuatum (O. KTZE) CHING, Acta Phytotax. Sinica 8 (1963) 326.— Thelypteris attenuata (O. KTZE) MORTON, Contr. U.S. Nat. Herb. 38 (1967) 35.— Type: CUMING 327, Samar (B; BRI, E, G, K, L, LE, SING, US). Dryopteris superficialis v.A.v.R. Bull. Jard. Bot. Btzg II, 20 (1915) 12; Handb. Suppl. (1917) 155. — Type: SA-ANAM 125, Obi I. (BO; L). Dryopteris erubescens sensu CHRIST, Philip. J. Sci. 2 (1907) Bot. 210. — Lastrea erubescens sensu CoPEL. Fern FI. Philip. (1960) 329. Cyclosorus alatellus sensu COPEL. Fern FI. Philip. (1960) 341, excl. syn. Caudex short, erect or suberect; stipe to at least 60 cm long, glabrous, rufescent, basal scales thin, to c. 10x 1mm. Lamina to 80cm or more long; lower pinnae much narrowed towards their bases, the narrowed part several cm long, distinctly lobed throughout; no reduced pinnae seen. Lar- gest pinnae 30 x 2 cm, lobed to 1.5 mm from costa; apex gradually attenuate to a cauda 3-5 cm long; lobes slightly oblique and slightly falcate; cos- tules 4-4.5 mm apart; veins to 18 pairs, basal pair with upcurved tips passing to sides of sinus- membrane or rarely joining just below the mem- brane; lower surface of costae and costules bear- ing scattered minute capitate hairs, between veins slender short acicular hairs variably present on different parts of the same frond, also short capi- tate hairs and almost spherical red resinous glands; upper surface of costa bearing very short capitate hairs, also acicular hairs distally, some capitate hairs present between veins. Sori some- what inframedial, lower ones divergent; indusia small with copious marginal spherical red resinous glands; spores dark, rugose. Distr. Malesia: Philippines (Luzon, Samar, Mindanao), Moluccas (Obi Island), N. Celebes? Ecol. ‘Edge of forest, petrophytic and ter- restrial’’ on limestone and limestone-derived soils (M. G. PRICE, on Samar), but probably not confined to limestone. Note. The hairs on the lower surface of pinnae are inconstant. I am not sure of a clear distinction between this species and A. ceramicum. A spe- cimen collected by H. H. BARTLETT (n. 7693) from Asahan, Sumatra has hairs on the lower surface as in the type of A. attenuatum, but glands are not well preserved. ENDERT 4165 from E. Kalimantan is similar. 7. Amphineuron ceramicum (v.A.v.R.) HOLTTUM, Blumea 23 (1977) 217.— Phegopteris ceramica v.A.v.R. Bull. Dép. Agr. Ind. Néerl. 18 (1908) 15; Handb. (1908) 506.— Type: TREUB s.n. Ceram (BO). Polypodium erubescens sensu HOOK. Spec. Fil. 4 (1862) 236, quoad pl. Molucc. tantum. — Poly- podium erubescens var. amboinense BAK. Syn. Fil. (1867) 306.— Type: collector not cited, Am- boina, ex Herb. Webb (K; FI, P). Dryopteris logavensis ROSENST. in Fedde Rep. 10 (1912) 232.—Lectotype (HOLTTUM 1977): BAMLER L34 (S-PA; BM). Phegopteris mamberamensis v.A.v.R. Bull. Jard. Bot. Btzg II, 24 (1917) 3; Handb. Suppl. (1917) 516.—Type: THOMSON 645, W. New Guinea, Mamberamo River (BO; L). Dryopteris moluccana C. CHR. Dansk Bot. Ark. 9 (1937) 64.—Type: FORBES 3273, Amboina (BM; B). Differing uncertainly from A. attenuatum as follows: basal pinnae gradually contracted at their bases to a narrow wing along the costa; no short acicular hairs on lower surface of pinnae between veins, no capitate hairs in that position on the upper surface; glands between veins on the lower surface varying from spherical to pyriform; glands on indusia spherical, resinous. Distr. Malesia: Moluccas (Amboina, Halma- hera, Ceram), New Guinea. Note. The type of Phegopteris ceramica v.A.v.R. does not show glands on the lower sur- face; it has pinnae to 22x 1.6cm. HOOKER’s Amboina specimen which he wrongly included in Polypodium erubescens is larger, with pinnae to 30x 3cm, the glands on the lower surface well preserved, those near sinuses distinctly elongate. A specimen of DE VRIESE from Ceram at Kew is similar but sterile. 8. Amphineuron pseudostenobasis (COPEL.) HOLTTUM, comb. nov.— Dryopteris pseudo- stenobasis COPEL. J. Arn. Arb. 10 (1929) 176. — Thelypteris pseudostenobasis (COPEL.) REED, Phytologia 17 (1968) 306.— Type: BRAss 1000, S.E. New Guinea, Vailala River (A; BRI, UC). 550 FLORA MALESIANA [ser. II, vol. 1° Cyclosorus alatellus sensu COPEL. Philip. J. Sci. 78 (1951) 445, p.p. — Fig. 19f-h. Differing from A. attenuatum as follows: no acicular hairs on lower surface between veins, no capitate hairs in this position on upper surface; glands between veins on lower surface fewer, apparently spherical, tending to collapse and form irregular thin patches of resinous substance or to disappear on drying of specimens; indusia very small, none seen bearing glands (which are pos- sibly present on living fronds). Distr. Solomon Islands and Malesia: S.E. New Guinea. Ecol. In riverine forest “in large masses” (BRASS). Notes. The type has pinnae to 26x 1.8cm, veins 11 pairs; pinnae on BRAITHWAITE’Ss spe- cimens from the Solomon Islands are up to 40 x 4cm (n. 4505) with veins to 25 pairs. Two of BRAITHWAITE’S specimens (4505 from Kolom- bangara and 4014 from Guadalcanal) certainly have a creeping caudex; his n. 4188 from San Cristobal has the note “rhizome short, erect’’, but otherwise is little different from the others. Spreading resinous glands are only observable on n. 4505. 9. Amphineuron paraphysophorum = (v.A.v.R.) HOLTTUM, Blumea 23 (1977) 217, excl. syn. Dry- opteris kiauensis C. CHR. — Dryopteris paraphy- sophora v.A.v.R. Bull. Jard. Bot. Btzg III, 2 (1920) 143.—Type: C. J. BrRooKxs 250S, Sumatra, Lebong Tandai (BO; BM). Caudex short, erect or suberect; stipe to 80cm long, basal part (up to 20cm in large fronds) copiously scaly, scales thin, narrow, 7-8 mm long. Lamina to 100 cm long (but plants of smaller size may be fertile), texture rather thin; pinnae to 28 pairs; lower pinnae not reduced but 8-10 pairs gradually narrowed towards their bases, lobes on basal 2cm of lowest pinnae 1-2 mm long, then gradually increasing to a maximum at 7cm from base. Largest pinnae 25-35cm long, 2.0-3.5 cm wide (widest on sterile fronds), apex acuminate but not long-caudate; edges lobed to 1mm from costa or more deeply; costules 3-4mm_ apart; lobes hardly falcate, at more than 60° to costa: veins to 20 pairs, basal acroscopic vein spreading at a wide angle and abruptly upcurved near its tip which touches the sinus-membrane or ends just above it, basal basiscopic vein more often ending above base of sinus; lower surface of rachis and costae bearing very short capitate hairs only, on costules, veins and between veins many pyriform to almost cylindrical yellow glands; upper surface of rachis bearing capitate hairs only, basal parts of costae the same, acicular hairs up to 0.3 mm long present on distal parts, very short ones also on costules. Sori medial, lower ones divergent; in- dusia small, lacerate, with elongate yellow glands on the margin; sporangia sometimes with similar glands on their stalks; spores with few large thick protuberances of varied shape. Distr. Malesia: S. Sumatra (Sarawak). Ecol. In Sarawak, in open places in forest at 90 m alt. Notes. Part of the above description is based on a plant from Gunong Mulu in Sarawak cul- tivated at Kew. This shows the abundance of glands of a distinctive shape on the lower surface of pinnae and on indusia. The glands on a dried specimen from the same locality are so shrivelled as to be hardly distinguishable. and Borneo 10. Amphineuron kiauense (C. CHR.) HOLTTUM, comb. nov. — Dryopteris kiauensis C. CHR. Dansk Bot. Ark. 9, 3 (1937) 64. — Type: ENDERT 4632 (wrongly cited as 4433), E. Kalimantan, Kutai, Kiau, 700 m (BO; L, SING). Description of type: stipe to 100cm_ long; lamina to 100cm long, texture firm; pinnae (fer- tile) to 18 x 1.5 cm, lobed to I-1.5 mm from costa; basal 2cm of basal pinnae consisting of a narrow wing above which is a gradual transition to full width of the pinna; veins 8-10 pairs, basal pair touching sides of sinus-membrane without anas- tomosis; short capitate hairs present on lower surface of costae, no glands detectable between veins; sori inframedial; indusia small, lacerate. Specimens from Sabah (KAKAWA & HOTTA 1291, distributed as Thelypteris erubescens) are very similar but have many spherical to pyriform glands on the lower surface, especially near the sinuses, and spherical resinous reddish glands on indusia. It seems probable that these represent the same species as the type of A. kiauense; there is a similar specimen (CLEMENS 29765) from Mt Kinabalu, 1200 m. Distr. Malesia: Borneo. 22. CHRISTELLA LEVEILLE, FI. de Kouy-tchéou (1915) 472, emend. Ho_trum, Taxon 20 (1971) 533, Blumea 19 (1971) 43, Kew Bull. 31 (1976) 293. — Nephrodium SCHOTT, Gen. Fil. (1834) t. 10 et sp. N. molle tantum, non RICHARD 1801. — Thelypteris subg. Cyclosoriopsis K. IWATS. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 28, p.p. — Cyclosorus sensu auctt. plur. p.p. — Fig. 1i, p, 20. 1981] THELYPTERIDACEAE (Holttum) 551 Caudex erect, suberect or creeping, in some species slender and wide- creeping; scales almost always narrow with many superficial hairs. Lamina in almost all species with I—5 (rarely to 10) pairs of lower pinnae gradually decrescent, the lowest usually not less than 2cm long and in almost all cases auricled on the acroscopic base, aerophores at the bases of pinnae not swollen; largest pinnae shallowly to deeply lobed, bearing erect acicular hairs (in some species minute) on all parts of the lower surface, also in almost all species between veins on the upper surface (these hairs never appressed), small capitate hairs sometimes also present; thick orange-red glandular hairs (not erect) present in some species on costules and veins on lower surface, sessile spherical glands lacking; veins free in a few species, in most species at least the basal veins from adjacent costules anastomos- ing, in some cases several pairs; sori indusiate (except in C. nana and C. buwaldae); sporangia lacking hairs or glands distally (except in some specimens of C. subpubescens) but bearing unicellular elongate glandular hairs on their stalks (except in spp. 1-4); spores dark, variously tuberculate or ridged, lacking thin wings. Type species: Christella parasitica (L.) LEv. Distr. About 50 spp.; throughout the warmer parts of the Old World; one species (C. hispidula) also in the neotropics; in addition c. 15S—20 spp. in Africa and the neotropics the status of which still needs to be established. Ecol. In Malesia, almost all are ferns of open places; species confined to forest occur in the region from N.E. India and Burma to S. China. Cytol. Base chromosome number 36. C. dentata, C. parasitica and C. subpubescens are tetraploid, C. hispidula diploid. Experimental hybridization of these species was undertaken at Leeds (see HOLTTUM 1976, p. 295); C. dentata and C. parasitica were shown to be allotetraploids with C. hispidula as one parent of each. C. arida has been shown to be diploid in northern India but has not been experimentally hybridized with tetraploids. It is evident that natural hybrids between some of the commoner species have also developed, but it is difficult to assign a parentage to them. Taxon. LEVEILLE characterized his genus as follows: pinnae lobed, with simple veins pinnately arranged in the lobes, sori with reniform indusia, seated on the veins in a row on each side of the costules. No type species was indicated. The characters apply to most species of Thelypteridaceae, and LEVEILLE’s list included representatives of four genera as arranged in the present treatment, also two which do not belong to the family. Three of his species were transferred to other genera by CHING; several others do not strictly conform to LEVEILLE’s own generic definition. Three of his species are closely allied and belong to a group recognized by me in my preliminary studies of the family, for which I was seeking an appropriate generic name; I therefore chose a species from this group as type. In 1964 IWATSUKI had cited an allied species, C. dentata, as type of Thelypteris subg. Cyclosoriopsis, but his definition of the subgenus would include many species which seem to me not nearly allied. In the earlier literature the species of this genus were not clearly distinguished, so that there is much confusion in the use of names. I have not attempted to assign meanings to all names cited; e.g. Aspidium patens, A. nymphale and A. parasiticum in BLUME’s ‘Enumeratio’ of 1828 are not clearly distinguished and I have not found the particular specimens to which he gave those names, which were copied from earlier works by others, who had described them very briefly. The single most distinctive character is the presence of an elongate unicellular gland on the stalks of sporangia (shown in SCHOTT’s figure of 1834 but mentioned by no-one else); similar glands are also present on the lower surface of pinnae in some species. This character is associated with others less precisely definable, among them the rather thick protuberances or ridges of the perispore, a character shared by Amphineuron. But the elongate gland on the sporangium-stalk is lacking in a group of four species in New Guinea, for which I propose a new section as follows: Leptochristella HOLTTUM, sect. nov. — Plantae parvae, calcicolae; pinnae 2-4.5 cm longae, tenues, subtus omnino piliferae; venae liberae vel inferiores anastomosantes; sori indusiati vel exindusiati; pedicelli sporangiorum glandulis destituti, interdum pilis acicularibus praediti. 552 FLORA MALESIANA [ser. Il, volauty SR PAAR OS BIR fA SD SE AGEITES AAO EAA Bees Ny QA LAW Rs atten SESE MC RL AAV eee Ge MS SCD Fig. 20. Christella dentata (FORSSK.) BROWNSEY & JERMY. a. Two pinna-lobes, x 4.— C. hispidula (DECNE) HOLTTum. b. Two pinna-lobes, x 4: c. upper surface of costa and base of costule, showing acicular and capitate hairs, x 18.—C, papilio (HOPE) HoLTTUmM. d. A reduced basal pinna, X 1; e. venation and sori, showing minute hairs on costa, x 6. — C. parasitica (L.) LEv. f. Pinna-lobe, showing hairs and glands, x 6.—C. subpubescens (BL.) HOLTTUM. g. Venation and sori, x 4.—C. harveyi (METT.) HOLTTUM. h. Two pinna-lobes, X 4; i. tip of a pinna-lobe showing sori, glands and hairs, x 16 (a PIGGOTT 1479, b-c YAPP 201, d~-e MOLESWORTH ALLEN 4949, f HOLTTUM s.n., g WALLICH 354, h-i BRYCE 5). 1981] THELYPTERIDACEAE (Holttum) 553 Type species: Christella perpubescens (ALSTON) HOLTTUM. The plants of this group of species, small in stature and consequently simplified in structure, are difficult to characterize and need more study. They have the frond-form of Christella with lower pinnae gradually and slightly reduced (shown most notably by C. perpubescens), abundant slender acicular hairs (fewest in C. nana) and spores of the Christella type. | see no other genus to which they could be considered allied. In Africa also are species with free veins and lacking (so far as I have observed) glandular hairs on the stalks of sporangia. For them I proposed the sectional name Pelazoneuron (J. S. Afr. Bot. 40, 1974, 144) but they appear to be related to a group of tropical American species in which glandular hairs of the Christella type do occur on the stalks of sporangia of at least two species (A. R. Smith, Univ. Cal. Publ. Bot. 29: 15, 66, 79) and apart from their free veins I see no clear distinction between these American species and C. hispidula. A few species in the Western Pacific (including C. harveyi in the present work) also have free veins; I suggest that they are derived from Malesian species with anastomosing veins. KEY TO THE SPECIES 1. Limestone plants; pinnae not over 4.5cm long; no elongate glands on stalks of sporangia (sect. Leptochristella). 2. Indusia distinctly present. 3. Pinnae to 27 pairs, S—7 lower ones gradually reduced .... . . . . I. C. perpubescens 3. Pinnae c. 6 pairs, lowest only slightly reduced . ....... 2... =. =. 2 C.minima 2. Indusia lacking or not clearly distinguishable. 4. Pinnae lobed almost to costae; veins all free nie: POOR eE Mee S33 MC Anana we Pinnae lobed less deeply; veins near bases of pinnae anastomosing . . . . «4. C. buwaldae . Plants not confined to limestone; pinnae commonly 10cm or more long; elongate glands present on stalks of sporangia (sect. Christella). 5. Veins all free. 6. Red ellipsoid glands present on lower surface of pinnae sep Re De ee en OS Caiharveyi 6. Such glands lacking et AT a, PESTA Be aie i ee OE CS neckelil 5. At least the basal veins anastomosing. 7. At least 3 pairs of lower pinnae gradually reduced. 8. Sinus-membrane prominent on lower surface, 3 Sea of veins ending beside it; reduced pinnae hardly-auricled’ 54s): JU eA: arida 8. Sinus-membrane not prominent, at most ‘3 pairs of veins s ending beside it: reduced pinnae clearly auricled. 9. Caudex strictly erect. 10. Reduced pinnae c. 8 pairs; hairs on lower surface of costae c.0.1mmlong ._ 8. C. papilio 10. Reduced pinnae 3—4 pairs; some hairs on lower surface of costae 0.3 mm or more long. 11. Pinnae lobed less than half-way to costa; hairs on lower surface of costae all less than 0.5 mm long : . . . 9, C. adenopelta . Pinnae lobed n more than half- -way to costa: hairs « on lower surface of costae mostly more than 0.5Smmilong .. . Sate & 2 Fe seo e0h Cahispidula 9. Caudex short-creeping, or ‘at most suberect distally. 12. Pinnae lobed at least half-way to costa; only 1 pair of veins truly anastomosing; some hairs 0.3 mm long present on lower surface of costae st04 . . . . 11. C. dentata 12. Pinnae lobed 1/4-1/3 towards costa; at least 13 pairs of veins ; truly anastomosing; all hairs on lower surface of costae c.0.1mmlong . . fo ES RI ee 123 | subpubescens 7. At most 3 pairs of lower pinnae gradually reduced. 13. Hairs on lower surface of costae all very short. 14. Pinnae lobed c. 1/4; minute capitate hairs present in addition to acicular hairs on lower surface of pinnae : . . . . . . . 12. C. subpubescens 14. Pinnae lobed c. 2/5: no capitate hairs on lower surface Ate . . . . 13. C. subdentata 13. Hairs on lower surface of costae always conspicuous, to | mm long. 15. Caudex long-creeping. 16. Pinnae lobed more than 1/2; apices of pinna-lobes not apiculate. 17. Basal 1-15 pairs of veins anastomosing, 1 pair passing to sinus-membrane 14. C. timorensis 17. Basal 1 pair of veins ea Sea at most the next acroscopic vein meeting the sinus- membrane : Loy Ree ree S='Caparasitica 16. Pinnae lobed less than 1/2; apices of pinna- lobes apiculate 2002. 216") acuminata 15. Caudex short-creeping or erect. 554 FLORA MALESIANA [ser. II, vol. 1° 18. Pinnae lobed much more than 1/3; some hairs more than 0.5 mm long present on lower surface. 19. Caudex erect; 2 or more pairs of lower pinnae reduced 19. Caudex not erect; basal pinnae not reduced 10. C. hispidula 15. C. parasitica 18. Pinnae lobed little more than 1/2; hairs on lower surface all less than 0. 5 mm long 1. Christella perpubescens (ALSTON) HOLTTUM, Kew Bull. 31 (1976) 304.— Dryopteris per- pubescens ALSTON, J. Bot. 78 (1940) 227; Nova Guinea n.s. 4 (1940) 111, pl. 8, f. 9, 10. — Thelyp- teris perpubescens (ALSTON) REED, Phytologia 17 (1968) 303. — Type: CLEMENS 7902C, N.E. New Guinea, Morobe Distr., Kalasa (B; BM). Caudex short-creeping; stipe 3-5 cm long, den- sely short-hairy throughout, basal scales 5mm long, narrow, soft, covered with minute hairs. Lamina 22cm long; texture thin; pinnae to 27 pairs, c. 7 pairs of lower ones gradually decrescent, basal pinnae 3-5 mm long; apex of frond also gradually attenuate. Largest pinnae of type 2.8x0.9cm (sterile), 2.2x0.8cm (fertile), sessile, short-acuminate, lobed to 0.5mm from costa, basal pair of lobes sometimes separately adnate to the costa, the acroscopic lobe not elon- gate but sometimes with a sinuous margin; cos- tules 2.5-3 mm apart; veins to 4 pairs, free, basal acroscopic one sometimes touching side of sinus- membrane, basiscopic one to margin above base of sinus; lower surface of rachis and costae den- sely covered with short hairs, scattered hairs more than 0.5 mm long also present, costules, veins and surface between veins bearing many short erect acicular hairs, also pale capitate hairs; upper sur- face throughout bearing slender erect acicular hairs, also some capitate hairs. Sori medial or a little supramedial; indusia firm, hairy; no hairs seen on sporangium-stalks; spores dark, closely and irregularly tuberculate. Distr. Solomon Islands; Malesia: New Guinea. Ecol. On limestone cliffs near sea-level (Wai- geo Island and Solomons); in eastern New Guinea reported to occur in rock crevices (type and BRASS 23774) at 240-500 m. Note. BRASS 23774, from Milne Bay District, lacks capitate hairs, and also BRAITHWAITE 4841 from the Solomon Islands; the latter also differs in longer stipes and less reduced basal pinnae. L. E. CHEESMAN 1227 from Waigeo Island has pinnae to 4.5 cm long, and a few capitate hairs. 2. Christella minima HOLTTUM, Kew Bull. 31 (1976) 304. — Type: JERMY 7875, New Ireland, on limestone (BM). Caudex short, apparently erect; stipe to 4.5 cm long, slender, short-hairy, basal scales c. 3mm long. Lamina to 9cm long, texture very thin; pinnae 6 pairs; basal pinnae a little reduced, with enlarged basal acroscopic lobes; apex of frond gradually attenuate. Largest pinnae 1.8 x0.9cm, lobed a little more than halfway to costa; apex obtuse, lobes rounded, entire; costules 3mm 11. C. dentata apart; veins free, to 4 pairs in the largest lobes, basal acroscopic vein ending at base of sinus, basiscopic one above base of sinus; lower surface covered throughout with slender short hairs, some longer ones present on rachis and costae; upper surface covered with short slender hairs, some longer ones also present on costules and veins. Sori supramedial, on basal acroscopic veins, a few on the second vein; indusia bearing many slender hairs; spores very dark with irregular blister-like protuberances. Distr. Malesia: New Guinea (New Ireland), only known from the type. Note. This may be an immature state of C. perpubescens. 3. Christella nana HOLTTUM, Kew Bull. 31 (1976) 304. — Type: MCKEE 1938, W. New Guinea, Biak Island (L). Caudex short-creeping; stipe 4-Scm_ long, glabrous, basal scales 1.5mm_ long, narrow. Lamina dimorphous, c. 7cm long, thin; pinnae 6 pairs, wider on basiscopic than on acroscopic side of costa, basal pair a little reduced. Sterile pinnae to 3.0 1.0cm, with stalks 1mm long, lobed al- most to the costa near base; lobes oblique, entire, with rounded tips, basal acroscopic lobe a little enlarged; apex of pinnae obtuse; costules to 3 mm apart; veins to 4 pairs, free, both basal ones passing to margin above base of sinus; lower surface of rachis covered with stiff erect hairs of varying length, costae and costules bearing very short somewhat antrorse hairs, rest of surface glabrous; upper surface glabrous apart from the costae. Fertile pinnae to 2.0 0.6cm; sinuses be- tween lobes wider than in sterile pinnae; veins in largest lobes 3 pairs, costules in upper pinnae once forked; sori confined to the pinna-lobes, on upper pinnae | or 2 in each lobe; no indusia; spores as in S. minima. Distr. Malesia: known from type. Ecol. Beneath overhanging coral rock just above beach. New Guinea (Biak I.), only 4. Christella buwaldae (HOLTTUM) HOLTTUM, comb. nov. — Pronephrium buwaldae HOLTTUM, Blumea 20 (1972) 115.— Type: BUWALDA 4979, Aru Islands, P. Kobro6r (L; K). Caudex short-creeping; stipe to 8cm _ long, slender, covered with short pale erect hairs, basal scales to 5mm long, narrow, with superficial short hairs. Lamina of type 16cm long consisting of an apical section 13cm long, widening downwards and deeply lobed, the lobes veined as pinnae, with 1981] THELYPTERIDACEAE (Holttum) 555 2-3 pairs of free pinnae; texture very thin, trans- lucent; basal pinnae somewhat reduced. Largest pinnae 2.8 1.0cm, sessile or with very short stalks; base symmetrically broadly cuneate; apex abruptly obtuse; edges lobed 1/3 towards costa, lobes subtriangular; costules 3mm apart; veins 3-4 pairs, basal pair anastomosing except near apex of pinna; lower surface of rachis, costae and costules covered with erect hairs of varying length, longest almost 1 mm long, surface between veins bearing very slender erect hairs; upper sur- face similarly hairy. Sori small, on pinnae in- framedial, on lobes of apical lamina supramedial; no indusia but acicular hairs present on the recep- tacle (or a very small hairy indusium?); sporangia lacking glands or setae; spores light brown. Distr. Malesia: Moluccas (Buru & Aru Is.). Ecol. On P. Kobro6r “in forest” (the island is mainly limestone); on Buru “limestone, 1200 m” (TOXOPEUS s.n. 4 Sept. 1921, BO). Note. The Buru specimen is larger than the type, with an apical lamina 8cm long and c. 6 pairs of free pinnae, largest pinnae 3.2 x 1.3.cm, lobed more deeply than those of the type; in other respects it agrees. An acicular hair was observed on the stalk of a sporangium of the Buru speci- men, and the pubescence of the frond is very like that of C. perpubescens, not like that of any spe- cies of Pronephrium, from which genus the spe- cies is here transferred. But the spores of the type need a careful re-examination; sporangia of the Kew isotype are immature. 5. Christella harveyi (METT.) HOLTTUM, Kew Bull. 31 (1976) 306; Allertonia 1 (1977) 219, f. 8 A-D. — Aspidium harveyi METT. in Kuhn, Lin- naea 36 (1869) 115.— Dryopteris harveyi (METT.) O. KTZE, Rev. Gen. Pl. 2 (1891) 812; C. CHR. Bishop Mus. Bull. 177 (1943) 83.— Thelypteris harveyi (METT.) PROCTOR ex K. IWATS. Amer. Fern J. 53 (1963) 133. — Type: HARVEY, Fiji (B; K). Dryopteris euaensis COPEL. Univ. Cal. Publ. Bot. 12 (1931) 391.— Type: PARKS, Tonga (UC; BM, K). Thelypteris novae-hiberniae HOLTTUM, Dansk Bot. Ark. 25, 2 (1967) 50.— Type: KOIE 1848, New Ireland (C). Nephrodium patens sensu HOOK. Spec. Fil. 4 (1862) 95, p.p.— Dryopteris patens sensu v.A.v.R. Handb. (1908) 189, p.p. — Fig. 20h-i. Caudex long-creeping, 5-7 mm diameter when dry; stipe 20-30cm long, hairy in groove, basal scales to 8X 1.5mm. Lamina 50-70 cm long, tex- ture thin; pinnae 20-25 pairs; lower pinnae variously reduced, lowest sometimes only 3cm long; reduced pinnae more widely spaced, some- times with basal acroscopic lobe enlarged and lobulate. Largest pinnae commonly 15-20 x 1.5- 2.2cm, rarely to 3cm wide; base not auricled; apex caudate-acuminate; edges lobed to 1-1.5 mm from costa; lobes oblique, slightly falcate, entire, with rounded apices; costules 4-4.5mm_ apart; veins 10-12 pairs, basal acroscopic one passing to side of the short sinus-membrane, basiscopic one to margin above base of sinus; lower surface of rachis, costae and costules bearing copious short hairs and a variable number of longer ones, some thick orange glandular hairs present on costules and veins, between veins a variable number of slender short erect acicular hairs and glandular hairs; upper surface of costae densely hairy, scat- tered long hairs present on costules and veins, between veins a variable number of short suberect hairs and sometimes glandular hairs. Sori near margin; indusia firm, glabrous or with a few hairs; an orange glandular hair on the stalks of some sporangia. Distr. Solomon Islands, New Hebrides, Fiji, Samoa, Wallis Isiand, Tonga, in Malesia: New Guinea (Admiralty Islands, Bismarck Archipelago). Ecol. In open places in forest at 0-1000 m. 6. Christella peekelii (v.A.v.R.) HOLTTUM, Kew Bull. 31 (1976) 306. — Dryopteris peekelii v.A.v.R. Bull. Dép. Agr. Ind. Néerl. 18 (1908) 7; Handb. (1908) 188.— Type: PEEKEL 44, “New Guinea” (BO). Caudex probably long-creeping; stipe probably to 20cm long, copiously short-hairy, basal scales not seen. Lamine 3040cm long; pinnae 18-22 pairs; basal pinnae slightly reduced, their basal pair of lobes almost free, the acroscopic one enlarged and deeply dentate. Largest pinnae 10 1.6cm (sterile), 9*1.3cm (fertile), acuminate, lobed to less than 1mm from costa, lobes entire and slightly falcate; costules of sterile pinnae 4mm apart, of fertile pinnae 3 mm; veins 8-10 pairs, arranged as in C. harveyi; lower sur- face bearing very short hairs throughout, no glandular hairs present; upper surface bearing scattered long hairs on costules and veins and short suberect hairs between veins. Sori near margins of lobes; indusia short-hairy; orange glands present on stalks of sporangia. Distr. Malesia: Papua New Guinea (New Bri- tain). Note. PEEKEL collected in the Bismarck Archipelago; the type was probably found in New Ireland. It is closely related to C. prolixa (WILLD.) HoL_trumM (Aspidium obliquatum METT.) of New Caledonia, which also lacks glandular hairs on the lower surface of pinnae. 7. Christella arida (D. DON) HOLTTUM in Nayar & Kaur, Comp. to Bedd. (1974) 206; Kew Bull. 31 (1976) 320; Allertonia 1 (1977) 172, 225, f. 9A. — Aspidium aridum D. Don, Prodr. Fl. Nepal (1825) 4. — Nephrodium aridum (D. Don) J. SM. in Hook. J. Bot. 4 (1841) 188; BEDpD. Handb. (1883) 272.—Dryopteris arida (D. DON) O. 556 FLORA MALESIANA [ser. II, vol. 1° KTZE, Rev. Gen. Pl. 2 (1891) 812; v.A.v.R. Handb. (1908) 212; BACKER & POSTH. Varenfl. Java (1939) 50.—Cyclosorus aridus (D. DON) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 194; HOLTTUM, Rev. FI. Malaya 2 (1955) 259, f. 146; COPEL. Fern FI. Philip. (1960) 362.— Thelypteris arida (D. DON) MORTON, Amer. Fern J. 49 (1959) 113. — Type: WALLICH, Nepal (not found at BM; possibly at BR, see MORTON 1974, infra). Aspidium obscurum BL. En. Pl. Jav. (1828) 150. — Nephrodium obscurum (BL.) T. MOORE, Ind, Fil. (1858) 98; Racrs. FI. Btzg 1 (1898) 125. — Type: BLUME, Java (L). Polypodium acuminatum ROXxB. Calc. J. Nat. Hist. 4 (1844) 490, non HoutTtT. 1786; MORTON, Contr. U.S. Nat. Herb. 38 (1974) 335.— Type: ROXBURGH, “Ind. Or.” (BR; G). Dryopteris arida var. ebeneorachis COPEL. in Elmer, Leafl. Philip. Bot. 2 (1908) 390. — Type: ELMER 1044, Negros (MICH; BO, G). — Fig. 1i. Caudex long-creeping, 4-5 mm diameter; stipe 15-30cm long, glabrous except in groove, basal scales 5 mm long, narrow. Lamina to 150 cm long; pinnae to 30 pairs or more; lower 3-5 pairs gradually or subabruptly decrescent and more widely spaced, not or little auricled, lowest com- monly 5S-10mm long; apex of frond almost pin- na-like; texture very firm. Largest pinnae to 16x 1.8cm (width above the somewhat dilated base), acuminate, lobed c. 1/4 towards costa. lobes with a short stiff point; costules 3-4 mm apart; veins to 10 pairs, prominent beneath, 15 pairs, at a broad angle to costule, anastomosing, next 3 pairs very oblique and passing to the sinus-membrane which is prominent on the lower surface; lower surface of costae bearing rather sparse stiff erect hairs 0.2mm long and a few very narrow scales, cos- tules and veins with scattered short acicular hairs and thick yellow glandular hairs, short erect hairs sometimes present between veins; upper surface of costae covered with short antrorse hairs, costules, veins and surface between veins almost or quite glabrous. Sori medial, lower ones diver- gent; indusia glabrous or with some glandular and short acicular hairs. Distr. N.W. India to southern China, Thailand and Vietnam; throughout Malesia: N. Queens- land, New Hebrides, New Caledonia, Fiji, Samoa. Ecol. In open places in low country, often among tall grasses, less commonly at higher al- titudes (once in Malaya at 1800 m). Plants growing in tall grass have a greater number of reduced basal pinnae than those in more exposed places. Note. Reports on the distribution of this spe- cies are often erroneous because it has been con- fused with Sphaerostephanos invisus and S. unitus which have a similar habit (see Holttum in Al- lertonia, 1977). LoyAL found a plant in North India to be diploid; some specimens are possibly hybrids with C. subpubescens. 8. Christella papilio (HOPE) HOLTTUM in Nayar & Kaur, Comp. to Bedd. (1974) 208; Kew Bull. 31 (1976) 321.—Nephrodium papilio Hope, J. Bombay Nat. Hist. Soc. 12 (1899) 625, t. 12.— Cyclosorus papilio (HOPE) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 214; MOLESWORTH ALLEN, Gard. Bull. Sing. 22 (1967) 180, 185. — Thelypteris papilio (HOPE) K. IwaTs. Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 175.— Lectotype (HOLTTUM 1976): LEVIGNE s.n. 1880, N.E. India, below Darjeeling 1200 m (K). Nephrodium molle var. major BEDD. Handb. Suppl. (1892) 76, p.p. — Fig. 20d-e. Caudex massive, erect; stipe 10-30cm long, minutely hairy, basal scales thin, c. 7X 1mm. Lamina to 150cm or more long; c. 10 pairs of lower pinnae gradually decrescent, broadly trian- gular, strongly auricled on acroscopic base, broadly truncate on basiscopic, margins above base shallowly lobed, apex acuminate; lowest pinnae I|-3cm long. Largest pinnae to 17 x2cm; base subtruncate; apex acuminate with short cauda; edges lobed 1/4-1/3 towards costa (most deeply in fertile pinnae), lobes slightly falcate and rounded at their tips; costules 4-4.5mm apart; veins 7-9 pairs, 1) pairs anastomosing, 1-2 pairs ending beside the sinus-membrane; lower surface of all parts bearing very short erect hairs (less than 0.1 mm long on rachis and bases of costae, a little longer on distal parts of pinnae), some thick orange glandular hairs sometimes present on veins in pinna-lobes; upper surface of costae covered with hairs 0.3-0.5 mm long, scattered shorter hairs on costules, minute erect hairs on surface between veins. Sori medial; indusia rather large, thin, with short hairs as lamina. Distr. Southern India & Ceylon; N.E. India, Thailand; Malesia: to northern Malaya. Ecol. In Malaya, in forest on sloping ground at 600-900 m. Notes. The Indian specimens lack orange glandular hairs on the lower surface, but such glands are present on all specimens from Thailand and Malaya. MANTON found a Ceylon plant to be tetraploid, LOYAL found plants in N. India to be diploid. 9. Christella adenopelta HOLTTUM, Kew Bull. 31 (1976) 322. — Type: W. A. SLEDGE 1698, Samoa (K). Near C. papilio, with the same erect caudex and similar pubescence, but only 3-4 pairs of lower pinnae gradually decrescent, the lowest 3.5 cm long; largest pinna 12x 1.8cm; hairs between veins on the lower surface a little longer than in C. papilio and many short capitate hairs also present with them, some scattered hairs up to 0.5 mm long present distally on costae and costules; indusia large, bearing glandular hairs like those on the lower surface of veins in addition to many very short acicular hairs. 1981] THELYPTERIDACEAE (Holttum) 55 i] Distr. Samoa and Malesia: S.E. New Guinea (Brown River, near Port Moresby). Ecol. In secondary swamp forest, low altitude. 10. Christella hispidula (DECNE) HOLTTUM, Kew Bull. 31 (1976) 312.—Aspidium hispidulum DECNE, Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834) 346.— Dryopteris hispidula (DECNE) O. KTZE, Rev. Gen. Pl. 2 (1891) 813; C. Cur. Ind. Fil. (1905) 271, excl. syn. Nephrodium angusti- follum PRESL & N. smithianum PRESL. — Thelypteris hispidula (DECNE) REED, Phytologia 17 (1968) 283.— Type: GUICHENOT, Timor (P). Dryopteris —_ contigua ROSENST. Meded. Rijksherb. n. 31 (1917) 8; C. CHR. Gard. Bull. Str. Settl 7 (1934) 244.—Cyclosorus contiguus (ROSENST.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 243; HoLTTum, Rev. FI. Mal. 2 (1955) 282, f. 163.— Thelypteris contigua (ROSENST.) REED, Phytologia 17 (1968) 269. — Type: TEUSCHER, Borneo (L). Dryopteris hirtopilosa ROSENST. Meded. Rijksherb. n. 31 (1917) 7; M. G. PRICE, Kalikasan 2 (1973) 112. — Thelypteris hirtopilosa (ROSENST.) REED, Phytologia 17 (1968) 283. — Type: MERRILL 7671, Luzon (L; K). Dryopteris repandula v.A.v.R. Nova Guinea 14 (1924) 20.—Cyclosorus repandulus (v.A.v.R.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 248; PANIGRAHI & MANTON, J. Linn. Soc. Bot. 55 (1958) 729-743.— Thelypteris repandula (v.A.v.R.) REED, Phytologia 17 (1968) 308. — Type: H. J. LAM 1058, W. New Guinea, Mam- beramo River (BO; L). Dryopteris parasitica var. falcatula CHRIST, Philip. J. Sci. 2 (1907) Bot. 197.— Cyclosorus falcatulus (CHRIST) COPEL. Fern FI. Philip. (1960) 339.— Thelypteris falcatula (CHRIST) REED, Phytologia 17 (1968) 276.—Type: COPELAND 1677, Mindanao (MICH). — Fig. 20b-c. Caudex erect; stipe 15-25cm long bearing slender pale hairs, basal scales 7-10mm long. Lamina 30-50 cm long, texture thin; pinnae 20-30 pairs, 2-4(—6) lower pairs gradually decrescent with dentate acroscopic auricles, lowest com- monly 3 cm long. Largest pinnae 8-12 (—15) x 1.2- 1.5(—2.0) cm; base truncate with basal acroscopic lobe a little elongate; apex short-acuminate; edges lobed 3/4 towards costa, lobes slightly oblique; costules 3-4mm apart; veins 7-9 pairs, lowest pair, at a wide angle to the costules, uniting to form a short excurrent vein to the sinus, next pair to margin; lower surface of rachis bearing rather sparse slender pale hairs 1-1.5 mm long, shorter hairs on costae, costules and veins, short slender hairs on surface between veins, some short capi- tate hairs usually present on costules and veins, no thick glandular hairs; upper surface of rachis and costae hairy as lower surface, scattered long hairs present on costules and veins, short capitate hairs sometimes present between veins. Sori medial or a little supramedial; indusia thin, rather small, bearing hairs of varying length. Distr. Tropical America and wetter parts of tropical Africa; Ceylon & S. India; Khasya Hills southwards to Malaya; throughout Malesia: Caroline Islands. Ecol. Less common than C. parasitica and in more sheltered places, low altitudes to 1500 m. Notes. Plants from Florida, Ascension Island, Ghana, Nigeria, Ceylon and Sarawak have proved to be diploid. For cytotaxonomic studies involving this species, see PANIGRAHI & MANTON 1955 (under Cyclosorus repandulus). Only synonyms based on Malesian specimens are cited above; see HOLTTUM 1976 for a fuller synonymy. The latest information on plants in tropical America is by A. R. SMITH, Univ. Cal. Publ. Bot. 59 (1971) 64, under the name Thelyp- teris quadrangularis (FEE) SCHELPE. Christella siamensis (TAGAWA & K. IWATS.) HOLTTUM, in Thailand and further north, differs only in less deeply lobed pinnae, and a distinction from T. hispidula seems doubtful; study of plants in cul- tivation is desirable. 11. Christella dentata (FORSSK.) BROWNSEY & JERMY, Brit. Fern Gaz. 10 (1973) 338; HOLTTUM, Kew Bull. 31 (1976) 314. — Polypodium dentatum Forssk. Fl. Aegypt. Arab. (1773) 185.— Dry- opteris dentata (FORSSK.) C. CHR. Vid. Selsk. Skr. VIII, 6 (1920) 24; BACKER & POSTH. Varenfi. Java (1939) 58, excl. syn. Aspidium parasiticum & A. patens.— Thelypteris dentata (FORSSK.) E. St. JOHN, Amer. Fern J. 26 (1936) 44. — Cyclo- sorus dentatus (FORSSK.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 206; COPEL. Fern Fl. Philip. (1960) 337, p.p.— Type: FORSSKAL, Arabia (C). Polypodium nymphale G. Forst. FI. Ins. Austr. Prodr. (1786) 81.—Aspidium nymphale (G. FORST.) SCHKUHR, Kr. Gew. 1 (1806) 36, t. 34; BLUME, En. Pl. Jav. (1828) 157.— Thelypteris nymphalis (G. ForsT.) REED, Phytologia 17 (1968) 297.—C. nymphalis (G. FORST.) PICHI SERMOLLI, Webbia 31 (1977) 252.— Type: G. FORSTER, New Zealand (BM). Polypodium molle JAca. Collect. Bot. 3 (1789) 188, non SCHREB. 1771. — Aspidium molle Sw. in Schrad. J. Bot. 1800, 2 (1801) 34, nom. nov. — Nephrodium molle (Sw.) R. BR. Prodr. Fl. N. Holl. (1810) 149; Racip. FI. Btzg 1 (1898) 188, p.p.?— Dryopteris mollis (Sw.) HTlERON. Hed- wigia 46 (1907) 348; v.A.v.R. Handb. Suppl. (1917) 183, p.p.? — Type: Cult. Vienna (W). Dryopteris mindanaensis CHRIST, Philip. J. Sci. 2 (1907) Bot. 194.—Cyclosorus mindanaensis (CHRIST) COPEL. Gen. Fil. (1947) 143; Fern FI. Philip. (1960) 363.— Thelypteris mindanaensis (CHRIST) REED, Phytologia 17 (1968) 293.— Type: COPELAND 607, Mindanao (MICH). 558 FLORA MALESIANA [ser. II, vol. 1° Dryopteris submollis v.A.v.R. Bull. Jard. Bot. Btzg III, 2 (1920) 152.—Type: LORZING 6040, Sumatra, Karo Plateau (BO; L). Cyclosorus subpubescens sensu. HOLTTUM, Rey. FI. Malaya 2 (1955) 273, f. 157. — Fig. 1p, 20a. Caudex short-creeping; stipe very variable, to 50cm long, short-hairy, basal scales c. 8 mm long. Lamina to 90cm long; pinnae 15-25 pairs; lower 2-4 pairs of pinnae gradually decrescent, lowest commonly 4-5 cm long, sometimes less, all stron- gly auricled on the acroscopic base, the auricles lobed with forked veins in the lobes. Largest pinnae commonly 8-10 x 1.5-1.8 cm (largest seen 23x2.2cm, type of Dryopteris mindanaensis); apex acuminate; edges lobed 1/2-2/3 towards costa, lobes slightly oblique with rounded tips; costules commonly 4mm apart; veins 8-9 pairs, basal ones anastomosing with excurrent vein to the sinus, next acroscopic vein (basiscopic some- times also) passing to side of the short sinus- membrane; lower surface of rachis bearing slen- der pale hairs 0.2-0.4 mm long, hairs on costae and costules mostly 0.2mm long with some longer ones, rarely to 0.5mm long, distally on pinnae short hairs present on surface between veins; hairs on upper surface of costae to 0.5mm or more long, scattered similar hairs on costules and veins, very short hairs between veins. Sori medial apart from lowest ones which sometimes touch those on veins from neighbouring costules; in- dusia thin, copiously short-hairy. Distr. Throughout tropics and subtropics of the Old World, since 1930 adventive in various places in the Americas. Ecol. In open or lightly shaded places, at 0- 1500 m. Notes. For a fuller synonymy, see HOLTTUM 1976. This is a very variable species which has greatly multiplied with the clearing of forest in the past 100 years. Plants examined from various sources have all been tetraploid; some experi- mental hybridization has been effected with C. hispidula (q.v.) and C. parasitica. It is probable that natural hybrids also occur; these are difficult to discriminate. Earlier authors did not distinguish between this species and its near allies, so that statements in literature are unreliable. Much more local study is needed. 12. Christella subpubescens (BL.) HoOLtTTuM, Webbia 30 (1976) 193; Kew Bull. 31 (1976) 323. — Aspidium subpubescens BL. En. Pl. Jav. (1828) 149. — Dryopteris subpubescens (BL.) C. CHR. Gard. Bull. Str. Settl. 4 (1929) 390; BACKER & PosTH. Varenfl. Java (1939) 65.— Thelypteris subpubescens (BL.) K. IwWATS. Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 173, excl. syn. Aspidium jaculosum CHRIST. — Type: BLUME, Java, river bank at Tanjong Krukot, Batavia (L, n. 910, 327- 113, large sheet collection). Aspidium subpubescens BL. var. B. BL. En. PI. Jav. (1828) 149.—Type: BLUME, Java, Noesa Kambangan (L, n. 922, 220-249, large sheet collec- tion). Aspidium subpubescens BL. var. C BL. L.c.— Type: BLUME, W. Java, Kolleket (L). Aspidium molle var. latipinna BENTH. FI. Hongkong. (1861) 455.— Nephrodium latipinna Hook. Syn. Fil. (1867) 292.— Dryopteris latipinna (BENTH.) O. KTZE, Rev. Gen. Pl. 2 (1891) 813; v.A.v.R. Handb. (1908) 217. — Cyclo- sorus latipinna (BENTH.) TARD. Notul. Syst. 7 (1938) 73; HOLTTUM, Rev. FI. Malaya 2 (1955) 276, f. 159. — Thelypteris latipinna (BENTH.) K. IWATS. Acta Phytotax. Geobot. 21 (1865) 166; MorRTON, Contr. U.S. Nat. Herb. 38 (1974) 361. — Lectotype (MORTON 1974): HANCE 135, Hong Kong (K). Dryopteris sumatrana v.A.v.R. Handb. (1908) 227. — Nephrodium molle var. major BEDD. Handb. Suppl. (1892) 76, quoad pl. Sumatr. tantum. — Dryopteris subpubescens var. major C. CHR. Gard. Bull. Str. Settl. 4 (1929) 390.— Cyclosorus sumatranus (v.A.v.R.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 249; HoLr- TUM, Rev. Fl. Malaya 2 (1955) 275, f. 158.— Thelypteris sumatrana (v.A.v.R.) TAGAWA & K. IwatTs. Acta Phytotax. Geobot. 22 (1967) 101. — Lectotype (HOLTTUM 1955): C. MILLER s5.n. 1778, Sumatra (BM). Dryopteris — pseudoamboinensis ROSENST. Meded. Rijksherb. n. 31 (1917) 7.— Thelypteris pseudoamboinensis (ROSENST.) PANIGRAHI, Phytologia 30 (1975) 410, pl. III]. — Lectotype (PANIGRAHI 1975): KORTHALS 270, Sumatra (L). Dryopteris acuminata ROSENST. Meded. Rijksherb. n. 31 (1917) 7, non Watts 1916. — Thelypteris blumei PANIGRAHI, Phytologia 30 (1975) 409, nom. nov.; op. cit. 31 (1975) 369. — Type: ZOLLINGER 735, Java, Ufer des Tjidurians (L, n. 908, 333-35). Aspidium amboinense sensu BL. En. Pl. Jav. (1828) 148; KUNZE, Bot. Zeit. 6 (1848) 261; METT. Farngatt. IV (1858) 105. — Nephrodium molle var. amboinense BEDD. Handb. (1883) 278, p.p. — Fig. 20g. Caudex short-creeping or suberect; stipe short, minutely hairy. Lamina varying much in size; several pairs of lower pinnae _ gradually decrescent, their acroscopic auricles rather small, entire or slightly crenate; largest pinnae lobed less than 1/2 towards costa (in small plants 1/4); in small plants at least 13 pairs of veins anastomosing and the next pair ending beside the sinus-mem- brane, in large plants 2-23 pairs of veins anas- tomosing to form a zig-zag excurrent vein and +1 pair ending beside the sinus-membrane; lower surface of rachis, costae and costules bearing minute hairs (0.1 mm long or less), in some spe- cimens also orange glandular hairs on costules and veins, surfaces between veins minutely hairy, usually with some very short capitate hairs; upper 1981] surface of costae bearing hairs c. 0.3mm long, scattered longer hairs sometimes present on cos- tules and veins, minute suberect hairs usually present between veins. Sori medial; indusia rather large, thin, glabrous or minutely hairy; in a few cases, capitate hairs present on sporangia. Distr. N.E. India to S.W. China; Burma, Thailand, Vietnam; throughout Malesia; N. Queensland, New Hebrides, Fiji. Ecol. On stream-banks in moderate shade in forest, also in lightly shaded places elsewhere, in lowlands. Notes. It would be possible to make a key distinguishing the types of the basionyms above cited, but I find it impossible to fit all other spe- cimens into such a key. More field study, and experimental study of cultivated plants, are needed. The types of Aspidium subpubescens BL. var. C and Nephrodium iatipinna HOoOK., also the specimens of BLUME and ZOLLINGER on which the misinterpretations of the name Aspidium am- boinense WILLD. were based, were small fertile plants. I have seen such plants on the banks of small streams in the forest in Pahang, where the streams are subject to periodic flooding with swiftly-flowing water after a sudden storm. In such places, plants are washed away before they attain their full size; a plant from the Tahan river in Malaya, grown at Kew, is much larger than any on the river banks, and is very like BLUME’s var. B, though less rigid in texture. Plants found grow- ing spontaneously in and near the Botanic Garden in Singapore (also others growing similarly at Bogor) are very like the type of A. subpubescens BL. and differ from the fully-grown plants of stream-bank origin in having somewhat broader pinnae. I have no evidence that the Singapore and Bogor plants are fertile from an early age, and think that they are probably genetically different from stream-bank plants, but I cannot point to clear distinctions. Thick orange glandular hairs are present on the lower surface of pinnae of most of the stream- bank plants from Malaya, and of the Java plants named amboinense by BLUME, KUNZE, MET- TENIUS and HOOKER. Such glands are absent from the type of Nephrodium latipinna HooK. and other specimens from Hong Kong, also from the type of A. subpubescens BL. and similar spe- cimens from Singapore and Bogor. But other specimens very similar to typical A. subpubescens have a few such glandular hairs, while on the other hand some stream-bank plants from north- ern Malaya are quite glandless like the type of N. latipinna from Hong Kong. Some stream-bank plants from Malaya, also from Luzon, have small capitate hairs on some of their sporangia; I have not observed this character in any other specimens of Christella. The capitate hairs appear to be just like those on the lower surface of pinnae between veins. THELYPTERIDACEAE (Holttum) 559 13. Christella subdentata HOLTTUM, Kew Bull. 31 (1976) 335. — Type: VILLAFLORES 57, Mindoro, Lubang Island (MICH). Caudex short, erect or suberect; stipe 25cm long, minutely hairy, basal scales c. 10 1mm. Lamina 50cm long; pinnae 15 pairs; lowest 1-2 pairs of pinnae somewhat reduced (basal pinnae 5cm long), not auricled on the acroscopic side. Largest pinnae 9.5 * 1.5cm; base truncate; apex acuminate with cauda 2cm long; edges lobed 2/5 towards costa; costules 3.5mm apart; veins to 8 pairs, lowest pair anastomosing, second acros- copic veins passing to side of sinus-membrane; lower surface of rachis, costae and costules bear- ing hairs 0.1mm long, similar hairs present be- tween veins; upper surface of rachis covered with hairs 0.5 mm long, shortet hairs present on costae, sparse minute hairs between veins. Sori medial: indusia short-hairy. Distr. Malesia: Philippines (Mindoro), only known from the type. Note. This may be a hybrid. 14. Christella timorensis HOLTTUM, Kew Bull. 31 (1976) 316. — Type: BLOEMBERGEN 3364, Timor, Mt Timan (L; BO, K). Caudex long-creeping, 5mm diameter, bearing fronds 2-3 cm apart; stipe short-hairy, basal scales c. 6X 1.5mm, rigid. Lamina of type incomplete, probably 70cm or more long with pinnae more than 20 pairs; 4-5 pairs of lower pinnae decrescent, with enlarged entire basal acroscopic lobes (to 2.0cm long), lowest pinna seen 7.5cm long. Largest pinnae 15cm long, 2.2cm wide above base; apex narrowly acuminate but not caudate; edges lobed 3/5—2/3 towards costa; lobes oblique, a little falcate, broadly pointed; costules 4.5-5 mm apart; veins to 12 pairs, basal 1-15 pairs anastomosing, next acroscopic vein, or next pair, passing to side of sinus-membrane; lower surface of rachis bearing rather sparse hairs 0.7-1.0mm long, costal hairs erect, to 0.7 mm, shorter hairs on costules and veins, sometimes also a few thick glandular hairs, slender erect hairs present be- tween veins; hairs on upper surface of costae to 0.5mm long, similar hairs scattered on costules and veins, no hairs between veins. Sori suprame- dial; indusia with short stiff hairs. Distr. Malesia: Lesser Sunda Is. (Timor), only known from type. Ecol. In grassland along stream at 1300 m. 15. Christella parasitica (L.) Lev. FI. Kouy- tchéou (1915) 475: HoLTTUM, Kew Bull. 31 (1976) 309. — Polypodium parasiticum L. Sp. Pl. (1753) 1090. — Dryopteris parasitica (L.) O. KTZE, Rev. Gen. Pl. 2 (1891) 811; @ Cur Ark: Bots 9)! 11 (1910) 26, f. 4.—Cyclosorus parasiticus (L.) FARW. Amer. Midl. Nat. 12 (1929) 259; HoLt- TUM, Rev. Fl. Malaya 2 (1955) 281, f. 162.— Thelypteris parasitica (L.) K. Iwats. Mem. Coll. 560 Sci. Univ. Kyoto B, 31 (1965) 172.—Type: OSBECK, Canton (S-PA in Herb. Swartz). Aspidium procurrens METT. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 231.— Dryopteris procurrens (METT.) O. KTZE, Rev. Gen. Pl. 2 (1891) 813; v.A.v.R. Handb. (1908) 211, 819, p.p.— Cyclo- sorus procurrens (METT.) COPEL. Fern FI. Philip. (1960) 340, nomen tantum. — Thelypteris procurrens (METT.) REED, Phytologia 17 (1968) 306. — Type: ZIPPELIUS, Java (L, n. 908, 335-152). Nephrodium didymosorum PARISH ex BEDD. Ferns Br. India (1866) t. 200.—N. molle var. didymosorum BEDD. Handb. (1883) 279. — Dry- opteris didymosora (BEDD.) C. CHR. Ind. Fil. (1905) 262; v.A.v.R. Handb. (1908) 225.— Type: PARISH, Burma, Moulmein (not found in Herb. Bedd. at K). Nephrodium tectum BEDD. Handb. Suppl. (1892) 79, excl. KING 8757.— Lectotype (HOLT- TUM 1976): WALLICH 394, Singapore (K-W). Dryopteris albociliata COPEL. J. Arn. Arb. 10 (1929) 177.— Thelypteris albociliata (COPEL.) REED, Phytologia 17 (1968) 259. — Type: BRASS 566, E. New Guinea, Basiatibu (GH; UC). Cyclosorus benguetensis COPEL. Philip. J. Sci. 81 (1952) 28; Fern Fl. Philip. (1960) 341.— Thelypteris benguetensis (COPEL.) REED, Phy- tologia 17 (1968) 263.—Type: MERRILL 7679, Luzon, Benguet Subprov. (MICH) — Fig. 20f. Caudex short- to long-creeping; stipe to 40cm long, copiously hairy. Lamina to 40 cm long, tex- ture thin; pinnae c. 20 pairs, closely placed; basal pinnae (as dried for herbaria) deflexed, not widely spaced, not or little reduced, auricled at the acroscopic base, auricle curved towards the rachis, sometimes dentate. Largest pinnae com- monly 162cm, short-acuminate, lobed 2/3-3/4 towards costa, lobes slightly oblique; costules c. 4mm apart; veins 8-10 pairs, basal pair anas- tomosing, next pair passing to margin above base of sinus or sometimes the second acroscopic vein touching the sinus-membrane; lower surface typically covered with slender erect hairs to 1 mm long, thick orange glandular hairs usually present on veins in the lobes; hairs on upper surface of costae thicker than on lower, similar hairs scat- tered on costules and veins, very short hairs be- tween veins. Sori medial, lower ones divergent; indusia hairy. Distr. Wetter parts of tropics and subtropics of Asia; throughout Malesia; Queensland; in the Pacific to Tahiti and Hawaii; E. Africa (Uganda, Kenya); St Helena (introduced ?). Ecol. In open places, mainly at low altitudes but also to 1500 m; apparently nowhere abundant. Notes. The type was discovered by CHRIS- TENSEN (1910); previously the name was used confusedly. In Kwangtung (source of the type) almost all plants have a rather short-creeping caudex and many glandular hairs on veins, but some have a longer rhizomatous caudex and few FLORA MALESIANA [ser. II, vol. 1° or no glands. The type of Aspidium procurrens METT. lacks glands. BEDDOME at first used the name procurrens for a mixture of species which had in common a creeping rhizome; v.A.v.R. also used the name confusedly. Under Cyclosorus procurrens COPELAND described a fern with narrow pinnae lobed half-way to the costa; I have not seen specimens. Some specimens have the frond-form of this species but short hairs; some agree in hairs and glands but have some reduced basal pinnae (type of Cyclosorus benguetensis); some such specimens are probably hybrids. MANTON found that C. parasitica in Ceylon is tetraploid and her co-workers produced hybrids with the diploid C. hispidula and with other tetraploids. In breeding, the absence of glandular hairs is recessive to their presence. 16. Christella acuminata (HoutTtT.) Lev. FI. Kouy-tchéou (1915) 476; HOLTTUM, Kew Bull. 31 (1976) 333.— Polypodium acuminatum HourTt. Nat. Hist. 14 (1783) 181, pl. 99, f. 2. — Thelypteris acuminata (HOUTT.) MORTON, Amer. Fern J. 48 (1958) 139; K. Iwatrs. Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 186.— Type: THUNBERG, Japan (UPS). Polypodium unitum THUNB. FI. Jap. (1784) 336, non LINN.— Polypodium sophoroides THUNB. Tr. Linn. Soc. 2 (1794) 341, nom. nov. — Neph- rodium sophoroides (THUNB.) DESV. Mém. Soc. Linn. Paris 6 (1827) 256; BAK. Syn. Fil. (1867) 289. — Type: as for P. acuminatum HOUTT. Caudex long-creeping, slender; stipe 25cm or more long, sparsely hairy. Lamina to 60cm long, texture firm; lower pinnae more widely spaced than the rest but not or little reduced. Largest pinnae to 9X 1.5cm (fertile), 15 x 2cm (sterile), lobed a little less than half-way to costa; lobes acute, basal acroscopic lobe enlarged, + curved towards the rachis, with some forked veins; cos- tules 3.5—4.5 mm apart; veins commonly 7 pairs (to 10), pale and prominent on both surfaces, basal pair anastomosing, next pair ending beside sinus- membrane; lower surface of costae and costules bearing a variable number of short erect hairs and a few longer ones, sparse short hairs present on and between veins, glandular hairs lacking; upper surface of costae bearing hairs to 0.5mm long, similar hairs often scattered on costules and veins, a variable number of very short hairs between veins. Sori medial or a little supramedial; indusia large, copiously short-hairy. Distr. Southern China and southern Japan, in Malesia: Philippines (Babuyan Is. and N. Luzon). Notes. For a fuller synonymy, see CHING 1938 and IWATSUKI 1965. The specific epithet sophoroides is used in most of the older lit- erature. In general aspect this species is similar to C. arida but lacks reduced basal pinnae and glandular hairs; the larger pinnae of C. acuminata are also rather strongly auricled. ADDENDA, CORRIGENDA ET EMENDANDA As has been done in Series I, Flowering Plants, it seems useful to complete the volume with worthwhile additions and corrections. Page numbers are provided with either a or b denoting the left and right columns respectively. 13b 14a 20a 20b 22a 22b 34a Gleicheniaceae Gleichenia volubilis var. peninsularis. Replace by the following: 7a. Gleichenia gigantea WALL. ex HOOK. Spec. Fil. 2 (1844) 5, t. 3A; BEDD. Ferns Brit. India (1865) t. 30.— Diplopterygium giganteum (HOOK.) NAKAI, Bull. Nat. Sci. Mus. Tokyo n. 29 (1950) 50.— Type: WALLICH 157, Nepal (K). G. glauca auct. non (THUNB.) HOOK.; BEDD. Handb. (1883) 2, quoad pl. Ind. Differs from G. volubilis JUNGH. of Java and Sumatra as follows: lower surface of lamina distinctly glaucous; stipular leaflets branched and more deeply lobed; scales on lower surface of rachis and costae rusty brown like the felt of stellate hairs. Distr. N.E. India at c. 1500 m; Vietnam; N. Thailand; in Malesia still only known from the collection cited from the Malay Peninsula. Note. Apart from the copious scales and hairs on rachis and costae this differs little from G. longissima BL. Further field study on the higher mountains of Malaya is desirable. Gleichenia clemensiae TUM. Add the following: Stipular leaflets lacking (thus resembling G. bullata, p. 13). Gleichenia reflexipinnula C. CHR. Add the following: BRASS 30382 from Mt Wilhelm, 2900 m, has ultimate branches to 42 by 4.5cm with costules 5 mm apart and veins distinctly prominent. Gleichenia truncata (WILLD.) SPR. var. truncata. Add the following synonym: G. bifurcata BL. En. Pl. Jav. (1828) 250. Gleichenia milnei BAKER. Add the following synonym: Sticherus kajewskii St JOHN, Occ. Pap. Bishop Mus. 17 (1942) 81. Add to Distr.: Mindanao (OLSEN 986). (COPEL.) HOLT- Insert the following after Gleichenia erecta: 25a. Gleichenia hooglandii HOLTTUM, Blumea 14 (1967) 327. — Type: HOOGLAND & SCHODDE 7692, W. High- lands of Papua New Guinea (CANB). Intermediate between G. erecta and G. bolanica (p. 24). Differs from G. erecta as follows: largest lamina-segments on the ultimate branches 8-15 mm long, costules 3-4 mm apart. Dicranopteris linearis (BURM. f.) UNDERW. 34a 35a 37 39 40 44b var. ferruginea (BL.) HOLTTUM. Correction for synonymy: The com- bination D. ferruginea was first published by HosoKAWaA, Trans. Nat. Hist. Soc. Formosa 25 (1935) 435. Additional synonym: D. tomentosa COPEL. Univ. Cal. Publ. Bot. 18 (1942) 217.—Type: CLEMENS 41228, N.E. New Guinea (UC). Dicranopteris linearis var. subferruginea (HIERON.) NAKAI. Add the following synonym: Gleichenia caudata COPEL. Bishop Mus. Bull. 59 (1929) 9.— Type: GILLESPIE 4389, Fiji (UC). Dicranopteris linearis var. subpectinata (CHRIST) HOLTTUM. Add to Distr.: Palawan. Additional varieties of Dicranopteris linearis which need study: var. bidentata v.A.v.R. Bull. Jard. Bot. Btzg III, 5 (1922) 204, Karimon I. var. crassifrons v.A.v.R. Ibid., Ternate. var. stipulosa CHRIST, Bull. Herb. Boiss. II, 6 (1906) 1010, Luzon. Schizaeaceae Schizaeaceae: Fossils. Add the following: JENNINGS & EGGERT, Amer. J. Bot. 59 (1972) 66 state that the anatomy of Senftenbergia is like that of Ankyropteris and conclude that Senften- bergia is not related to Schizaeaceae. But the sporangia and spores of Senftenbergia are very similar to those of some living Schizaeaceae; the latter are certainly much reduced and specialized vegetatively. The subject needs a much more comprehensive study. Cytology of Lygodium. Add the following: L. longifolium from Singapore: n=58, tetraploid; L. salici- folium from Perak: n= 28?, diploid. (In- formation from I. MANTON, in litt.). Schizaea SM. Add the generic synonym: Ripidium BERNH. in Schrader, J. Bot. 1800, pt 2 (1801) 127. Schizaea malaccana BAK. var. robustior C. CHR. Add the synonym: S. robusta BAK. Syn. Fil. (1868) 429.—Type: HILLEBRAND, Hawaii (K). Lygodium SWARTZ. Delete the generic BERNH. synonym Ripidium (561) 562 FLORA MALESIANA 53b 59a 61b 63 64 TA 76 Lygodium flexuosum (L.) Sw. Add the synonym: Ugena polymorpha CAV. Teones61(1801)) 75; t. S95, te 1: Lygodium longifolium (WILLD.) Sw. Add to Distr.: Palawan. Lygodium versteegii CHRIST. Add to Distr.: Philippines: Luzon. Mindanao, Isoetaceae Tsoetes LINNE. See revision of the genus in New Guinea by J. R. CROFT, Blumea 26 (1980) 177-190, with key, description of new species I. hopei and I. stevensii, and under I. neo- guineensis BAKER a new variety rheo- phila; also SEM photographs of spores of all species and much new information on vegetative morphology and distribution. Isoetes sp. has been found in Central West Sumatra, in addition to Mindanao and New Guinea the third island where the genus is hitherto discovered in Malesia. Cf. FI. Males. Bull. 30 (1977) 2767 and J. R. FLENLEY & R. J. MORLEY, J. Biogeogr. 5 (1978) 57-58. It is assumed to be an un- described species, differing from the Philip- pine one in the megaspores. The exact local- ity is on the westside of Mt Kerintji, midway the lakes Sati and Landak Panjang, at c. 2080 m altitude, in small, shallow, muddy depressions in swampy forest, 1°42’S and 101°11'E. Cyatheaceae Conspectus of the family Cyatheaceae: new comments: I would now raise the subfamilies to the rank of family, but see no reason for other changes. PICHI SERMOLLI (Webbia 31, 1977, 333-334, 423-427) includes all in the order Dicksoniales, with suborders and families thus: Thyrsopteridineae: (Thyrsopteris ). Culcitineae: Culcitaceae (Culcita). Dicksoniineae: Dicksoniaceae (including Cibotium and Cystodium), Lophosoriaceae (Lophosoria). Cyatheineae: Cyatheaceae (Sphaeropteris, Alsophila, Nephelea, Trichopteris, Cyathea, Cnemidaria). Metaxyineae: Metaxyaceae (Metaxya). Subdivision of the genus Cyathea: recent proposals: R. M. TRYON (Contr. Gray Herb. 200, 1970, 1-53) has recognized Sphaeropteris BERNH. as a distinct genus, in which he includes the tropical American species Thyrsopteridaceae 80 82b [ser. II, vol. 1° mentioned on p. 124 of the present work, and also several others which disagree in scales and sori from the specification on p. 65; in my opinion the latter should be excluded. TRYON limits the genus Cyathea to some tropical American species, dis- tinguishing them from all Malesian species here included in Cyathea subg. Cyathea solely on the lack of a seta at the apex of stipe-scales. He has transferred all Malesian species of Cyathea subg. Cyathea sensu HOLTTUM 1963 to the genus AI- sophila (type species A. australis R. BR.). I agree that subg. Sphaeropteris as defined on p. 76 is clearly distinct from all other members of the family, but I cannot agree that the remaining species, including those of tropical America, are divisible into natural groups on such clearly defined characters. CONANT has’ shown _ that hybrids exist between species of different genera as recognized by TRYON, and in one case such hybrids have good spores (Rhodora 77, 1975, 441-455). All species so far examined, of several different genera as recognized by TRYON (including Sphaeropteris) have the chromosome number 69. This is a very strong indication that Cyathea in the broad sense adopted in Flora Malesiana is a phy- letic unity, and I adhere to my recognition of it as a single genus. As Sphaeropteris is the only sub-group separable on well- defined characters, I still place all the rest in a subgenus Cyathea, the further sub- division of which seems to me still un- certain. The species C. australis (R. BR.) DOMIN (type of Alsophila) has spores which differ considerably from those of the majority of Malesian species transferred to Alsophila by TRYON (see GASTONY, Amer. J. Bot. 63, 1976, 754, f. 110 and compare with figures of Malesian species on p. 751). Taxonomy of Malesian species of Cyathea: Since 1963 many new collections have been made, especially in New Guinea. I have not been able to examine all of these. The following data are based on specimens which have come to my attention; probably more new species remain to be recognized, and new information about many species remains unrecorded. In Key to the species, after lead 77, add: 77a. Pinnules to 22 mm long, bearing free tertiary leaflets. 69a. C. nothofagorum 77a. Pinnules to at least 50 mm long, lack- ing free tertiary leaflets. Cyathea hunsteiniana BRAUSE. Add to description: Scales on pinna-rachis 1981] Addenda, corrigenda et emendanda 563 10la 103b 118b 121b 130b 13S5a 138 140b 140b elongate, thin, dull brown with a few mar- ginal setae (WOMERSLEY & VANDEN- BERG 37293, Western Highlands, 1500 m). Cyathea oosora HOLTTUM. Add to Distr.: Pulau Tioman (KLU 19781). The Celebes specimens here included should probably rank as a distinct species; more material is needed. Insert additional species: 69a. Cyathea nothofagorum HOLTTUM, Blumea 14 (1966) 327.— Type: PULLEN 5358, N.E. New Guinea, W. Highlands, Kubur Range, in Nothofagus forest at 2775 m (K). Aspect of C. microphylloides and C. perpelvigera (p. 82), having small fronds and pinnate pinnules, differing in (large) hemitelioid indusia and lack of hairs on lower surface of pinna-rachis and costae, also in scales not bullate. Cyathea biformis (ROSENST.) COPEL. Add the following: PULLEN 7295 from the Upper Fly River at 100m has 1-2 pairs on small pinnae (the pinnules of which do not have an incised lamina) near the base of stipes, and unusually large pinnules; sterile pinnules to 9 by 2cm, fertile to 10 by 12cm: Cyathea lurida (BL.) COPEL. Add the following: A sterile specimen from a low altitude on limestone near Kuala Lumpur (J. BoEY 4, KLU) closely resem- bles this species but differs in pale brown stipe and rachis. Typical C. lurida occurs only on high mountain ridges. Cyathea sangirensis (CHRIST) COPEL. Reduce to a synonym of the following: Cyathea felina (ROXB.) MORTON, Contr. U.S. Nat. Herb. 38 (1974) 344. — Polypodi- um felinum ROXB. Calc. J. Nat. Hist. 4 (1844) 496.— Lectotype: from Amboina (G); others at BR and in Herb. J.E. Smith. Cyathea angiensis (GEPP) DOMIN. Add the following note: Stipes always spiny, length of spines 2-7 mm. Key to the species of subsection Fourniera: alter as follows: 7. Scales absent from lower surface of veins. 8. Spines on stipe scattered, 1-3 mm long; soral scales many, covering sorus to maturity 150. C. tripinnata 8. Spines on stipe copious, to 5 mm long; soral scales not covering sorus at maturity .150a. C. jacobsii Cyathea tripinnata COPEL. Delete synonym C. arachnoidea (non HOOK.) BACKER & POSTH. After Cyathea tripinnata, insert the fol- lowing: 150a. Cyathea jacobsii HOLTTUM, Rein- wardtia 8 (1974) 499. — Type: M. JACOBS 141 166a 167 214a 8113, S. Sumatra at 1200-1300 m (L). This species, occurring in South Sumatra and West Java, differs from C. tripinnata in the characters shown in the above modified key; it also has a different dis- tribution. Specimens named C. arach- noidea in W. Java by BACKER & POSsT- HUMUS (see above) belong to C. jacobsii. In Key to the species of subsection Schizocaena, lead 13: alter the word ‘pin- nules’ at end of line 2 from bottom to ‘segments’. Cyathea rosenstockii BRAUSE. Add the following: NAKAIKE 717, Central District, Papua, Woitape, 1500 m, has flat pale scales to 4mm long and dark glossy thick hairs on lower surface of pinna- rachis; costal scales narrow, pale with dark bullate bases, smaller distally; on costules very few small scales. Cibotium barometz (L.) J. SM. Add to Distr.: N.E. New Guinea (B. S. PARRIS in Fern Gaz. 11, 1979, 428). Culcita PRESL. Correction in Cytol.: C. macrocarpa n= 66; C. villosa 2n = 232 (tetraploid with base number 58); from unpublished observations by G. VIDA on plants in cultivation at Kew. Lindsaea-group (KRAMER) Sphenomeris TAGAWA. Mapped in Pac. Pl. Areas 3 (1975) 344-345. Sphenomeris chinensis (L.) MAXON. Mapped in Pac. PI. Areas 3 (1975) 346-347. Tapeinidium denhamii (HOOKER) C. CHR. Mapped in Pac. PI. Areas 3 (1975) 348-349. Lindsaea bouillodii CHRIST. For this species MORTON (Contr. U.S. Nat. Herb. 38, 1974, 385) made the new combination Lindsaea interrupta (ROXB.) MorRTON, based on Vittaria interrupta ROXBURGH, referring L. cambodgensis CHRIST to its synonymy (following my earlier misinterpretation of this name: KRAMER, Blumea 15, 1967, 563). In view of the gross incompleteness of ROx- BURGH’s original description, and of the complete lack of indication of provenance on the label of the specimen regarded as type by MorTON, I think it unwise to adopt ROXBURGH’s (and MORTON’s) names which I prefer to regard as names of uncertain application. The same holds in my opinion for the interpretation of Vit- taria lunulata ROXBURGH, despite MorR- TON’s assertions to the contrary (I.c. 386). Lindsaea tetragona KRAMER. This East Malesian—Pacific species, map- ped in Pac. Pl. Areas 3 (1975) 340, was biflora (KAULFUSS) 564 FLORA MALESIANA (ser. II, vol. 1° recently collected as far west as the Nico- bar Is. (Great Nicobar), and may thus be expected in western Malesia, too. Lindsaea polyctena KRAMER. Mapped in Pac. Pl. Areas 3 (1975) 340-341. Lindsaea tenuifolia BLUME. Mapped in Pac. PI. Areas 3 (1975) 340-341. Lindsaea adiantoides J. SMITH. Mapped in Pac. PI. Areas 3 (1975) 342-343. Lindsaea gueriniana (GAUD.) DESVAUX. Mapped in Pac. Pl. Areas 3 (1975) 332-333. For the record from Fiji see BROWNLIE, Pterid. Fl. Fiji (1977) 134. Lindsaea repens (BORY) THWAITES, with varieties. Mapped in Pac. Pl. Areas 3 (1975) 336-337. Lindsaea carvifolia KRAMER. This was placed in sect. Odontoloma of subg. Odontoloma. Its occurrence in Celebes was reported with a ?. A good series of specimens from that island (HENNIPMAN .S. 5264, L) confirms its presence; furthermore, it shows that sim- ply pinnate as well as bipinnate leaves occur side by side, and that the species should be transferred to sect. Lindsaenium, where it must be placed between L. rosenstockii BRAUSE and L. versteegii (CHRIST) v.A.v.R. In appearance and width of the pinnule segments it is more 245a 245b like the latter but differs in less divergent and usually spathulately broadened seg- ments. It is undoubtedly distinct from both; the simply pinnate leaves are indis- tinguishable from specimens previously examined. The distinction between sect. Odontoloma and sect. Lindsaenium may require reconsideration. Lindsaea rigida J. SMITH. Mapped in Pac. PI. Areas 3 (1975) 334-335; for the record from Fiji see BROWNLIE, Pterid. Fl. Fiji (1977) 139. Lindsaea sarawakensis KRAMER. This was described from a single, incom- plete collection and was only provisionally placed next to L. rigida. Two additional collections from Sarawak, NIELSEN 815 (AAU) and JERMyY 14334 (BM, Z), both from Mt Mulu, confirm the taxonomic position assigned to it and provide ad- ditional data. The rhizome anatomy and morphology agree with that of Lindsaea subg. Odontoloma sect. Lindsaenium, and the difference from L. rigida, as given in the key (p. 203), proves to be constant. Further additional data: petiole to c. 35 cm long; rhizome 1-1.5mm_ diam., long- creeping, castaneous; scales very much like those of L. rigida. Fertile pinnae al- ways with a short apical sorus only. 81 113a 265 266 269b ADDENDA, CORRIGENDA ET EMENDANDA 2 At couplet 101 in key delete °94, C. in- cisoserrata’, and insert afterwards: 10la. Apex of stipe-scales setiform; bullate scales on costules dull brown, smooth 94. C. incisoserrata 10la. Apex of stipe-scales not setiform; bullate scales on costules almost white, clathrate. 94a. C. decurrens After 94. C. incisoserrata, insert: 94a. Cyathea decurrens (HOOK.) COPEL. Univ. Cal. Publ. Bot. 14 (1929) 356; HoLt- TUM, Blumea 12 (1964) 248.—Alsophila decurrens HOOK. Spec. Fil. 1 (1844) 51.— Type: NIGHTINGALE s.n., Pacific (K). Stipe-scales broad, castaneous with broad paler margins, apex broadly rounded or + attenuate, not setiform; pinnules-lobes oblique, almost all free but decurrent, all deeply lobulate; costules covered beneath with almost white clathrate bullate scales; indusia very small, hidden by sporangia. Distr. Solomon Is., New Hebrides, Fiji, Samoa, Cook I.; in Malesia: New Ireland. Note. Collected in New Ireland at 2300 m (CROFT 301); on other islands found at much lower altitudes. CROFT’s specimen differs from others in its firmer bullate scales and in lacking the thick white hairs which are usually + abundant on the lower surface of costae in this species; it may represent a distinct variety. C. decurrens and allied Pacific species have stipe-scales like those of the Neotropic species to which TRYON limits the genus Cyathea; no other known Paleotropic species have this character. Teratophyllum METT. ex KUHN. Add to Distr.: Ceylon. Teratophyllum, section Teratophyllum. Add to Distr.: Ceylon. Teratophyllum aculeatum (BL.) METT. ex KUHN. Add to Distr.: Ceylon; rare, first collected by THWAITES, recently recollected in Sin- haraja Forest, KOSTERMANS 27853, Oct. 1979 (L). 325a- Bolbitis, series Heteroclitae. 326b 361b 393b 452a 494b 502b HENNIPMAN has reduced some taxa which I regard as species, notably B. sim- plicifolia (HOLTTUM) CHING, B. malac- censis (C. CHR.) CHING and B. nitens HOLTTUM. See my paper on Malayan Bolbitis, Mal. Nat. J. 33 (1980) 79-82 for an alternative opinion. Coryphopteris gymnopoda (BAK.) HOLT- TUM var. bintangensis HOLTTUM. Add to the description: The type has many such hairs but recent collections by A. G. PIGGOTT on Gunong Ulu Kali (Genting Highlands) at 1830 m show much variation. Specimens from Borneo are all quite glabrous between veins. Line 7 from bottom: Dryopteris penangiana (HOOK.) C. CHR. was based on Polypodium penangianum HOOK. The type of this was a WALLICH specimen which bore the local- ity Penang, but the species is otherwise only known from northern India, so I am sure it was wrongly localized. I did not know any- thing about this species when I wrote my book on Malayan ferns. Dryopteris perakensis was based on Aspi- dium perakense BEDD.,a species I could not account for and omitted from my book on Malayan ferns. I could not find a type in Kew herbarium, but some years later it was found, among other BEDDOME specimens, in the basement. It is based on an immature plant of Sphaerostephanos polycarpus mixed with a small fragment of a fertile pinna from ano- ther plant, an aberrant Christella. VAN ALDERWERELT VAN ROSENBURGH of course misinterpreted it. Sphaerostephanos_ mutabilis HOLTTuo. Add: Note. This is near S. aquatilis (no. 44), differing in narrower pinnae, veins free, and lower surface quite glabrous; the two should perhaps be united. Sphaerostephanos HOLTTUM. Add to Ecol.: less commonly to 1000 m. (BRAUSE) invisus (FORST. f.) (565) F 2 ae o 7 | C?.3 ea te ary INDEX TO SCIENTIFIC PLANT NAMES compiled by M. J. VAN STEENIS-KRUSEMAN Epithets or taxa above generic rank have been entered under their name. Infrageneric epithets have been entered immediately under the generic name to which they belong, preceded by the indication of their rank (subgenera, sections, etc.). If possible sections are arranged under the subgenera to which they belong. Epithets of wrongly identified species precede the correct ones. Infraspecific epithets have been entered under the specific name to which they belong, preceded by the indication of their rank (subspecies, varieties, formas, etc.). Epithets of new names and new combinations have been printed in bold type, synonyms in italics. An asterisk behind a page number denotes the presence of an illustration of the concerned taxon; ‘map’ printed behind a page number denotes that a map of the concerned taxon is present on that page. Page numbers in bold type denote main treatment. Some minor printing errors in plant names have been corrected. Of synonyms with a double authority, the latter has not always been cited in full. The full authority can easily be derived from the text. Scientific plant names mentioned in the Dedication have not been indexed. (Acrostichum) crispatulum Clarke Acrosorus il, XVii Acrosticheae 255 Abacopteris Fée xxi, 333, 336, 337, 507, 509 aspera Ching 512 Acrostichopteris 37 var. contaminans Clarke cuspidata (BI.) Ching 536 Acrostichum sensu Fée 289 321 elegans Fée 534 Acrostichum L. ili, vi, vii, xx, cumingii sensu Baker 304, glandulosa (BI. ) Fée 522 gymnopteridifrons (Hayata) Ching 513 lineata (Bl.) Holtt. 512, 530 menisciicarpa (BI.) Holtt. 526 multilineata (Wall. ex Hook.) Ching var. malayensis Holtt. 512 peltochlamys (C.Chr.) Holtt. 495, 507 philippinarum Fée 507, 512, 513 presliana sensu Ching 513 presliana (Ching) Ching 512 rubicunda (v.A.v.R.) Holtt. 536 salicifolia (Wall. ex Hook.) Holtt. 535 triphylla (Sw.) Ching 534 var. parishii (Bedd.) Ching 535 truncata Fée 526 urophylla (Mett.) Ching 533 Aconiopteris Pres] 289, 290, 291 Acrophorus ti, xix adiantoides (Bl.) Moore 249 cuneifolius (Presl) Moore 180 hemipterus (Bory) Moore 237 hymenophylloides (BI.) Moore 237 repens (Bory) Moore 237 PSS, OI; AOS, QO, Pik 291, 314, 318, 335 sect. Chrysodium (Fée) Hook. 277 sect. Elaphoglossum Hook. 289 sect. Lomariopsis Hook. 259, 265 sect. Polybotrya Hook. 275 sect. Stenochlaena 265 aculeatum (BI.) Racib. 269 aemulum sensu BI. 308 angulatum BI. 297, 313 apodum sensu Fée 301 appendiculatum Willd. 323 var. costulatum Hook. 325 var. hamiltonianum Baker 322 articulatum Hook. 276 beccarianum Baker 301 blumeanum Fée 311 blumeanum (Fée) Hook. 279, 281 borneense Burck 314 callifolium Bl. 307 celebicum Baker 523 commutatum Mett. ex Kuhn 308, 309 conforme sensu BI. 297, 308 var. Javanicum Mett. 314 conforme Sw. 290, 313 contaminans Wall. 321 307 cumingii Fée 307, 314 decurrens sensu BI. 303, 304 decurrens sensu Hook. 304 decurrens Desv. 292, 303, 314 var. cumingii v.A.v.R. 304 var. major Mett. ex Kuhn 304 var. ornatum Fée 304, 307 dichotomum L. 41 digitatum L. 41 diversifolium Bl. 326 exsculptum Baker 528 flagelliferum Wall. ex Hook. 325 gorgoneum sensu BI. 314 gorgoneum sensu Christ 298 gorgoneum Kaulf. 292, 313 gracile Racib. 269 hamiltonianum Wall. 322 heteroclitum Presl 325 hirtum Sw. 314 junghuhnianum Kunze 307 latifolium Sw. 314 laurifolium sensu Christ 262 laurifolium sensu Fée 297 laurifolium sensu Thw. 308 laurifolium Thouars 314 lessonii Mett. 301 melanostictum Bl. 301 micradenium Fée 314 miniatum Christ 311 (567) 568 (Acrostichum) modestum Baker 325 norrisii sensu Cesati 301 norrisii Hook. 302 obtusifolium sensu BI. 303 obtusifolium Willd. 304 oligodictyon Baker 380 ophioglossoides Goldm. 298 pellucido-marginatum Christ 298 proliferum Bl. 325 quoyanum Gaud. 327 repandum BI. 329 scandens Hook. 269 sinense Baker 325 smithii Racib. 262 sorbifolium L. 259, 261, 263 speciosum Presl 262 spectabile (Kunze) 262 squamosum sensu Sw. 314 stelligerum Wall. ex Baker 311 subdiaphanum Hook. & Grev. 291 variabile Hook. var. rasamalae Racib. 326 viscosum sensu BI. 311 viscosum 292 zollingeri Kunze 326 Actinostachys Wali. ex Hook. 40 boninensis Nakai 42 digitata Wall. ex Reed 41 inopinata Reed 42 wagneri Reed 44 Racib. Adiantum ii, xvii chinense (L.) Burm. 182 chinense L. 182 chusanum L. 182 cultratum Willd. 222 ensifolium (Sw.) Poir. 211 falcatum Blanco 180 heterophyllum (Dryand.) Poir., non Colenso 210 lanceolatum (Labill.) Poir. 211 lobatum (Poir.) Poir. ex Steud. 219 orbiculatum Lamk 206 tenuifolium Lamk 182 Adiantum group ii, ix, x, xi, xii, XIll, XIV, XVil Aglaomorpha ii, xvi Agyratae 71 Alsophila sensu C.Chr. 115 Alsophila R.Br.—See index p. 169-171 and p. 562. Sub- sequent references are: australis R.Br. 562 decurrens Hook. 565 FLORA MALESIANA (Alsophila) dryopteroidea Brause 401 fragilis Zoll. & Mor. 512 polypodioides Hook. 348, 349 Amauropelta 334, 338, 341 Ampelopteris Kunze ili, xxi, 340, 341, 342, 375, 378, 386, 387 elegans Kunze 387 prolifera (Retz.) Copel. 385*, 387 Amphicosmia Gardn. 73 alternans Moore 145 javanica (Pr.) Moore 111 manilensis (Pr.) Moore 110 Amphineuron Holtt. 340, 341, 342, 343, 437, 544, 551 attenuatum (O.Ktze) Holtt. 451, 545, 549, 550 ceramicum (v.A.v.R.) Holtt. 401, 545, 549 distinctum (Copel.) Holtt. 544, 545, 548 immersum (BI.) Holtt. 370, 401, 432, 545, 546*, 547 kiauense (C.Chr.) Holtt. 545, 550 opulentum (Kaulf.) Holtt. 544, 545, 546, 547*, 548 paraphysophorum (v.A.v.R.) Holtt. 545, 550 pseudostenobasis (Copel.) Holtt. 451, 545, 546*, 549 subattenuatum (Rosenst.) Holtt. 544, 545, 546*, 547 terminans (Hook.) Holtt. 387, 544, 545, 546* Anapausia zollingeri Presl 326 Anemia 37, 38, 39 Anemiaceae 39 Angiopteris il, vi, xv, 70 Anisocampium ii, xviii Ankyropteris 561 Anogramma Il, xVil Antrophyum iil, xxi Araiostegia ii, Xvill Arcypteris ili, xx Arthrobotrya J.Sm. ii, xx, 267, 275, 276 articulata J.Sm. 276 Arthromeris ti, xvi Arthropteris tii, xx, 257, 258 obliterata (R.Br.) J.Sm. 253 tenella (Forst.) J.Sm. 258 Aspidiaceae 255, 511 tribe Aspidieae 255 Aspidiales 339 Aspidistes Harris 65, 67 thomasii Harris 333 Aspidium Sw. 335, 336, 338 abortivum sensu Mett. 461 [ser. II, vol. 1° (Aspidium) abortivum BI. 429 var. B (var. C?) 429 abruptum Bl. 429 adiantoides Bl. 249 affine Bl. 512, 522, 527, 528 alatellum (Christ) Christ 451 amboinense sensu BI. 558, 559 amboinense Willd. 519, 559 arbuscula sensu Christ 456 aridum D.Don 555 asperum (Presl) Mett. 512 attenuatum Kunze ex Mett. non Sw. 549 barometz Willd. 165 benoitianum Gaud. 500 blumei Kunze ex Mett. 516 var. subpinnata Mett. 516 brackenridgei Mett. 408 calcaratum BI. 374 callosum Bl. 419 var. B 420 canescens (BI.) Christ 516 f. acrostichoides Christ 523 f. gymnogrammoides Christ 467 f. nephrodiiformis Christ 519 var. sensu Christ 516 ciliatum Benth. 375 conioneuron (Fée) Mett. 548 crassifolium Bl. 384 crassifolium Mett. 384 cucullatum sensu Christ 488 cucullatum BI. 478 dayi Bedd. 352 decursive-pinnatum (van Hall) Kunze 354 dimorphum Kunze 458 dissectum sensu Mett. 502 echinatum sensu Christ 379 echinatum Mett. 379 eusorum Thw. 429 excellens (Bl.) Holtt. 462 exiguum Kunze ex Mett. 539 var. b 538, 540 extensum BI. 548 fasciculatum Fourn. 367 ferox Bl. 391 flaccidum BI. 351 glaberrimum Richard 436 glanduligerum Kunze 351, 373 glandulosum sensu. Kunze 512 glandulosum BI. 513, 522, 527 gracilescens BI. 351 grammitoides Christ 373 hallieri Christ 379 harveyi Mett. 555 1981] (Aspidium) heterocarpon BI. 457 var. B Blume 458 heterocarpus sensu 498 heterodon BI. 449 heterophyllum (Presl) Hook. 515 hirsutum sensu Christ 488 hirsutum Kunze ex Mett. 455 hispidulum Decne 459, 557 var. B Mett. 459 var. solutum Mig. 459 var. ternatense Miq. 474 hookeri Baker 548 immersum BI. 370, 547 jaculosum Christ 558 javanicum Mett. 449 koordersii Christ 375 laeve Mett. 424 latebrosum Kunze ex Mett. 471 latum Kunze ex Mett. 384 lineatum sensu Mett. 527, 528 lineatum BI. 507, 512, 527 loherianum Christ 489 longipes sensu Christ 498 longipes Bl. 423 megaphyllum Mett. 461 var. hirsutum Mett. 452 menisciicarpon Bl. 526, 527 molle Sw. 557 var. latipinna Benth. 558 multijugum Christ 419, 420 multilineatum Mett. 461, 478 multilineatum Wall. 461, 462 nephrodioides Hook. 548 novo-pommeranicum Brause ex Rech. 327 nymphale (G.Forst.) Schkuhr Spills Ey / obliquatum Mett. 555 obscurum BI. 556 obtusatum Sw. 386, 387 ochthodes 414 opulentum Kaulf. 548 paludosum BI. 347 parasiticum 551, 557 patens sensu BI. 551, 557 pennigerum sensu BI. 462 perakense Bedd. 449, 565 pilosiusculum Mett. 452, 522 pilosiusculum Zipp. 523 polycarpon Bl. 448 procurrens sensu Christ 498 procurrens Mett. 560 productum Kaulf. 456, 460, 461 prolixum Willd. 413 Christ 461, Index to scientific plant names (Aspidium) reinwardtianum Kunze ex Mett. 374 repandum sensu BI. 512 sagittifolium Bl. 487 setigerum sensu Racib. 348 simplicifolium J.Sm. ex Hook. 490 simulatum Davenp. 371 solutum BI. 459 sorbifolium Willd. 289 squamigerum Fée 377 stipellatum Bl. 492 subpubescens Bl. 558, 559 var. C BI. 558, 559 terminans Wall. 548 thelypteris var. squamigerum Schlechtend. 377 triste Bl. 158 truncatum sensu Christ 425 truncatum Gaud. 430 var. celebicum Miq. 425 tylodes Kunze 413, 414 uliginosum Kunze 348 unitum sensu Mett. 386 unitum sensu Schkuhr 386 unitum (L.) Sw. 477 venulosum BI. 386 vile sensu Racib. 349 warburgii Kuhn & Christ 462 Aspleniaceae 339 Asplenieae subtribe Blechninae 255 Asplenium ii, xviii, 266 adiantum-nigrum L. 61 bakeri C.Chr. 235 epiphyticum Copel. 266, 270 erectum Willd. 354 Asplenium group ii, ix, Xi, Xiii, XVii Astralopteris 315 Athyrium ii, v, xviii, 69, 335, 337, 355 benguetense Christ 351 horizontale Rosenst. 366 hyalostegium Copel. 373 ledermannii Hieron. 368 Athyrium group ii, Xi, Xiil, xiv, XVIil Azolla iii, xxi Balantium Kaulf. 71, 158, 166, 167 auricomum 167 blumei Kunze 159 chrysotrichum Hassk. 159 copelandii Christ 168 culcita 167 glaucescens Link 165 javanicum Copel. 168 569 (Balantium) pilosum Copel. 168, 169 Belvisia ii, xv Blechnum ii, vii, xviii filiforme (A.Cunn.) Ettingsh. 263 punctulatum Sw. 61 Blechnum group ii, x, Xi, Xi, XViil Bolbitis Schott ti, xix, 255, 256, 257, 258, 278, 289, 314 ser. Bolbitianae Hennipm. 315, 316, 318, 321 ser. Egenolfianae Hennipm. 3155318, '322;329 ser. Heteroclitae Hennipm. 316, 318, 325, 329, 565 ser. Quoyanae Hennipm. 316, 318, 327 angustipinna (Hayata) Ito S172 3195320*3321 appendiculata (Willd.) Iwats. 258, 318, 319, 322 map ssp. appendiculata 317*, 318, 320%, 322; 323, 329 ssp. vivipara (Hook.) Hennipm. 322 map, 323 var. neglecta Hennipm. 320*, 322 map, 323 var. vivipara 322 map x arguta (Fée) Ching 329 cladorrhizans (Spr.) Ching 258 coloradica R.W.Brown 315 contaminans Ching 321 copelandii Iwats. 323 cuspidata Ching 325, 326 diversifolia (Bl.) Schott 326, 329 edanyoi Copel. 326 enormis C.Chr. 327 flagellifera Schott 325 x foxii Copel. 329 foxworthyi Ching 326 heteroclita (Presl) Ching 258, 315, 316*, 318, 319, 324%, 325, 329, 330 heteroclita x rhizophylla Hennipm. 329 hookeriana Iwats. 258, 322 hydrophylla Ching 326 inconstans Ching 326 interlineata (Copel.) Ching 319, 320*, 330 intermedia Iwats. 323 linnaeana C.Chr. 325 malaccensis (C.Chr.) Ching 326, 327, 565 membranacea C.Chr. 326 modesta Ching 325 naumannii Ching 327 570 (Bolbitis) nigra Ching 326 nitens Holtt. 326, 327, 565 novoguineensis Hennipm. 319, 320*, 324*, 328 ovata (Copel.) Ching 330 parva Ching 327 portoricensis (Spr.) Hennipm. 258 pseudoscalpturata Ching 326 quoyana auct. 329 quoyana (Gaud.) Ching 319, 321, 324*, 327, 328 map, 330 repanda (BI.) Schott 258, S85 3202) 321327, 329: 330 rhizophylla (Kaulf.) Hen- nipm. 319, 320*, 322, 323 map, 329, 330 rivularis (Brack.) Ching 319, 327, 328 map scalpturata (Fée) Ching 319, 320*, 321, 330 serratifolia (Kaulf.) Ching 278 serrulata Iwats. 323 simplicifolia (Holtt.) Ching 326, 565 sinensis (Baker) Iwats. 257, SIS, SiR SIs spAlies sys} map, 325, 329 var. costulata Tagawa & Iwats. 325 x singaporensis Holtt. 320%, 329 sinuata (Presl) Hennipm. 315, 318, 319% 320%, 326; 327 map, 329, 330 x sinuosa (Fée) Copel. 321, 329 nm. foxii (Copel.) Hen- nipm. 320*, 330 nm. sinuosa Hennipm. 330 stenophylla Ching 325 subcrenata (Hook. & Grev.) Ching 325, 330 var. prolifera 330 subsimplex Ching 326 taylorii 327 tenuissima Copel. 326 virens (Hook. & Grev.) Schott 316*, 3118, 321) 322 map var. compacta Hennipm. S19 5205509 2p S22 emap var. deltigera 322 map var. virens 322 map vivipara C.Chr. 327 Botrychium ii, xv Brainea ii, xviii FLORA MALESIANA Callipteris ii, xviii Calymella Presl 6 borneensis Ching 6 circinnata Ching 7, 9 dicarpa Presl 9 microphylla Presl 7 squamosissima Nakai 9 vulcanica Presl 9 Calymmodon il, xvii Campium Presl 255, 315, 318 angustipinnum Copel. 321 argutum Copel. 329 cuspidatum Copel. 325 diversifolium Copel. 326 enorme Copel. 327 foxworthyi Copel. 326 heteroclitum (Presl) Copel. 325 var. eurybasis Copel. 326 hydrophyllum Copel. 326 interlineatum Copel. 330 kajewskii Copel. 328 linnaeanum Copel. 325 membranaceum Copel. 326 modestum Copel. 325 nigrum Copel. 326 ovatum Copel. 330 parvum Copel. 327 pseudoscalpturatum Copel. 326 quoyanum Copel. 327 repandum Pres] 329 rivulare Copel. 328 scalpturatum Copel. 321 sinense C.Chr. 325 subcrenatum sensu Back. & Posth. 330 subsimplex Copel. 326 tenuissimum Copel. 326 validum Copel. 327 virens Pres] 321 viviparum Kjellb. 327 Cathetogyratae 71 Ceratopteris 1, Xvii Cerosora ll, XVil Cheilanthes 11, xvii fuscata Bl. 61 gigantea Cesati 349 hirta Sw. 61 multifida Sw. 61 setigera Bl. 349 stenophylla Kunze 349 Cheilolepton Fée 277, 278 blumeanum Fée 279 Cheiropleuria ii, Xv Chingia Holtt. 334, 340, 342, 378, 389, 397 acutidens Holtt. 389, 390 atrospinosa (C.Chr.) Holtt. 389, 390 bewaniensis Holtt. 390, 393 [ser. Il, vol@ay (Chingia) christii (Copel.) Holtt. 388%, 390, 394 clavipilosa Holtt. 390, 393 var. Clavipilosa 390, 393 var. javanica Holtt. 388%, 390, 393 ferox (Bl.) Holtt. 388%, 389, 390, 391, 392, 393 horridipes (v.A.v.R.) Holtt. 390, 394 imponens (Ces.) Holtt. 390, 391 longissima (Brack.) Holtt. 421 lorzingii Holtt. 390, 393 muricata (Brause) Holtt. 389, 391 paucipaleata Holtt. 390, 394 perrigida (v.A.v.R.) Holtt. 390, 394 pricei Holtt. 388*, 389, 390, 392 pseudoferox Holtt. 392 sakayensis (Zeiller) Holtt. 334, 388%, 390, 392 sambasensis Holtt. 390, 393 supraspinigera (Rosenst.) Holtt. 390, 395, 402 tenerior Holtt. 390, 395 urens Holtt. 389, 390, 395 Chnoophora Kaulf. 73 glauca BI. 135 lanuginosa Jungh. 132 lurida Bl. 121 tomentosa BI. 132 Chorizopteris Moore 277 Christella Lév. 333, 334, 335, 338, 340, 341, 342, 343, 413, 429, 439, 449, 517, 550, 559 sect. Leptochristella Holtt. 551 sect. Pelazoneuron Holtt. 553 acuminata (Houtt.) Lév. 553, 560 adenopelta Holtt. 553, 556 arida (D.Don) Holtt. 332*, 551, 553, 555, 560 buwaldae (Holtt.) Holtt. 551, 553, 554 crinipes (Hook.) Holtt. 378 dentata (Forssk.) Brownsey & Jermy 332*, 551, 552*, 553, 554, 557 harveyi (Mett.) Holtt. 552*, 55555 hispidula (Decne) Holtt. 459, SSilk 55275 S53 oS 4s Sue 558, 560 minima Holtt. 553, 554 nana Holtt. 551, 553, 554 1981] (Christella) nymphalis (G.Forst.) Pichi Sermolli 557 papilio (Hope) Holtt. 431, 552515535556 parasitica (L.) Lév. 551, 552*, S)3}, Sse Spi, Sey Seb 560 peekelii (v.A.v.R.) Holtt. 553, 555 perpubescens (Alston) Holtt. S524 559,554,555 prolixa (Willd.) Holtt. 555 siamensis (Tagawa & Iwats.) Holtt. 557 subdentata Holtt. 553, 559 subpubescens (BI.) Holtt. 3B4E Sil, Sil, Seve, Sis 556, 558 timorensis Holtt. 553, 559 Christella group 340 Christensenia ii, Xv Christiopteris ii, xvi Chrysodium 255, 291 cuspidatum Kuhn 325 heteroclitum (Presl) Kuhn 325 var. subcrenatum Kuhn 325 zollingeri Kuhn 326 Cibotium Kaulf. ti, xix, 65, 67, 69; 70, 71, 72, 159; 164, 169, 562 arachnoideum (C.Chr.) Holtt. 164, 166 assamicum Hook. 165 barometz (L.) J.Sm._ 164, 165*, 563 var. cumingii v.A.v.R. 166 var. sumatranum Christ 165 chamissoi Kaulf. 164 crassinerva Rosenst. 166 cumingii Kunze 164, 166 var. arachnoideum C.Chr. 166 djambianum Hassk. 165 glaucescens Kunze 165 glaucum sensu Bedd. 165 glaucum sensu J.Sm. 166 regale Linden 165* Cnemidaria 65, 69, 70, 72, 562 horrida 69, 70 Colysis ti, xvi cf. pedunculata (Hook. & Grev.) Ching 330 Condyloneura Christ 291 Coniogramme ii, vi, xvii Coniopteris 65, 67 hymenophylloides (Brongn.) Seward 65 (Coniopteris) murrayana Brongn. 65 Cornopteris ii, Xviii Coryphopteris Holtt. 331, 333, 334, 338, 340, 341, 342, 343, 354, 370, 437 andersonii Holtt. 357, 358, 364 andreae Holtt. 358, 364 arthrotricha Holtt. 355, 357, 363 athyriocarpa (Copel.) Holtt. 358, 365 athyrioides Holtt. 357, 363 atjehensis Holtt. 357, 362 badia (v.A.v.R.) Holtt. 355, 356*, 359, 369 borealis Holtt. 358, 364 brevipilosa Holtt. 358, 368 coriacea (Brause) Holtt. 358, 368 diaphana (Brause) Holtt. 357, 362 dura (Copel.) Holtt. 359, 368 engleriana (Brause) Holtt. 358, 366 fasciculata (Fourn.) Holtt. 354, 358, 367 gymnopoda (Baker) Holtt. 361 var. bintangensis Holtt. 357, 361, 565 var. gymnopoda 357, 361 var. humilis Holtt. 357, 361 habbemensis (Copel.) Holtt. 355, 356*, 359, 362, 366 hirsutipes (Clarke) Holtt. 355, 356*, 357, 359, 364, 494 horizontalis (Rosenst.) Holtt. 358, 366 hubrechtensis Holtt. 358, 367 inopinata Holtt. 354, 359, 370 iwatsukii Holtt. 357, 362 klossii (Ridl.) Holtt. 356%, 357, 361 lauterbachii (Brause) Holtt. 358, 367 ledermannii (Hieron.) Holtt. 358, 368 meiobasis Holtt. 358, 364 microlepigera Holtt. 358, 366 multisora (C.Chr.) Holtt. 355, 356*, 357, 359, 363 obtusata (v.A.v.R.) Holtt. 357, 362 oligolepia (v.A.v.R.) Holtt. 358, 368 pectiniformis (C.Chr.) Holtt. 355, 357, 363, 364 var. hirsuta Holtt. 357, 363 var. minor Holtt. 357, 364 Index to scientific plant names 57] (Coryphopteris pectiniformis) var. pectiniformis 357, 363 platyptera (Copel.) Holtt. 359, 369 plumosa (C.Chr.) Holtt. 357, 360 propria (v.A.v.R.) Holtt. 358, 367 pubirachis (Baker) Holtt. 365 var. major Holtt. 358, 365 var. philippinensis Holtt. 358, 365 var. pubirachis 358, 365 var. sulawesica Holtt. 358, 365 squamipes_ (Copel.) 354, 358, 364, 365 stereophylla (v.A.v.R.) Holtt. 355, 358, 363, 366 subnigra (Brause) Holtt. 359, 369 tahanensis Holtt. 355, 357, 363 tanggamensis Holtt. 358, 365 unidentata (Bedd.) Holtt. 355, 356*, 357, 359 viscosa sensu Holtt. 363 viscosa (Baker) Holtt. 332*, 355, 356*, 360, 365 var. borneensis Holtt. 358, 361 var. poiensis Holtt. 357, 361 var. viscosa 357, 360 Coryphopteris group 339 Crypsinus ii, xvi Cteisium Michx 39, 45 Ctenitismi} veo 1672 256; 2575 258, 336, 338 habbemensis (Copel.) Copel. 362 Ctenopteris ii, xvii Culcita Presl ii, xix, 65, 67, 69, 70, 72, 159, 166, 562, 563 subg. Calochlaena 65, 166, 167 subg. Culcita 65, 166, 167 coniifolia (Hook.) Maxon 167 copelandii Maxon 168 dubia (R.Br.) Maxon 70, 167, 168* formosae (Christ) Maxon 167 javanica (BI.) Maxon 167, 168* macrocarpa Presl 70, 167, 563 pilosa Christ 168 straminea (Labill.) 167, 169 villosa C.Chr. 167, 169, 563 Culcitaceae 562 Culcitineae 562 Holtt. Maxon 572 FLORA MALESIANA [ser. Il, volivee (Cyclosorus) ecallosus Holtt. 425 edanyoi Copel. 525 Currania ii, Xix Cyclosorus sensu Copel. 414, Cyathea Smith—See index p. 436 171-175. Subsequent Cyclosorus sensu Holtt. 414, references are: Ii, iv, Vi, 436 ellipticus sensu Copel. 485 VIGEX VS 75 3935598539) Cyclosorus Link iii, vi, xxi, 333, matutumensis (Copel.) 439, 562, and: 335, 337, 338, 339, 340, 474 subg. Cyathea sensu Holtt. 341, 342, 375, 378, 385, elmerorum (Copel.) Copel. 562 387, 389, 437 485 subg. Sphaeropteris (Bernh.) adenophorus (C.Chr.) Ching erectus Copel. 490 Holtt. 455 extensus (BI.) Ching 548 sect. Schizocaena (J.Sm.) Holtt. 338 subsect. Fourniera (Bommer) Holtt. 563 subsect. Schizocaena 563 angiensis (Gepp) Domin 563 arachnoidea sensu Back. & Posth. 563 atrispora Domin 401 biformis (Rosenst.) Copel. 563 decurrens (Hook.) Copel. 565 felina (Roxb.) Morton 563 hunsteiniana Brause 562 incisoserrata Copel. 565 jacobsii Holtt. 563 lurida (BI.) Copel. 563 microphylloides Rosenst. 563 nothofagorum Holtt. 562, 563 oosora Holtt. 563 perpelvigera v.A.v.R. 563 rosenstockii Brause 563 sangirensis (Christ) Copel. 563 tripinnata Copel. 563 woodlarkensis Copel. 400 Cyatheaceae ii, xi, xii, xv, 3, 65- 169, 291, 562 subfam. Cibotioideae 72 subfam. Cyatheoideae 71 tribe Cyatheae 71, 72 tribe Dicksonieae 71, 72 tribe Lophosorieae 72 subfam. Metaxyoideae 72 subfam. Thyrsopteridoideae 72 tribe Culciteae 72 tribe Thyrsopterideae 72 Cyatheineae 562 Cyclogramma Tagawa 334, 340, 341, 343, 411 auriculata (J.Sm.) Ching 412* himalayensis Tagawa 412 omeiensis 412 simulans (Ching) Tagawa 412 Cyclopeltis tii, xx Cyclosorus sensu auctt. plur. 550 Cyclosorus sensu Ching 414, 436 adenostegius (Copel.) Copel. 466 alatellus sensu Copel. 549, 550 alatellus (Christ) Copel. 451 albosetosus (Copel.) Copel. 505 angustifolius (Presl) Copel. 481 aoristisorus sensu Copel. 539 aoristisorus (Harr.) Copel. 538 arfakianus (Baker) Copel. 484 aridus (D.Don) Ching 556 austerus (Brause) Copel. 502 austrophilippinus (Copel.) Copel. 479 bakeri (Harr.) Copel. 516 beccarianus sensu Holtt. & Roy 514 beccarianus (Cesati) Copel. 513 benguetensis Copel. 560 boholensis Copel. 456 bordenii sensu Copel. 474 bordenii (Christ) Ching 457 callosus (Bl.) Ching 419 cataractorum Wagn. & Greth. 468 christii Copel. 394 clemensiae (Copel.) Ching 521 confertus (Brause) Copel. 49] contiguus (Rosenst.) Ching 557 cuspidatus sensu Copel. 537 cuspidatus (BI.) Copel. 536 debilis (Mett.) Copel. 448 degener Copel. 522 deltipterus Copel. 420 dentatus (Forssk.) Ching 557 dicranogramma 419 diminuens Holtt. 504 diminuta (Copel.) Copel. 515 dimorphus (Brause) Copel. 464, S01 distinctus (Copel.) Copel. 548 diversilobus Ching 452 duplosetosus Copel. 468 falcatulus (Christ) Copel. 557 farinosus (Brause) Copel. 503 ferox sensu Holtt. 392 ferox (Bl.) Ching 391 glaber Copel. 418 glandulosus (Bl.) Ching 522 gongylodes sensu Ching 386 gongylodes (Schkuhr) Link 335, 386 granulosus (Presl) Copel. 524 gregarius Copel. 480 halconensis Copel. 471 heterocarpus sensu Holtt. & Roy 451 heterocarpus (Bl.) Ching 457 var. glaucostipes (Bedd.) Holtt. 471 hispidula sensu Copel. 456 hosei (Baker) Copel. 526 imponens sensu Copel. 391 imponens (Ces.) Copel. 391 interruptus sensu Holtt. 545 interruptus (Willd.) H.Ito 332*, 385*, 386, 478 invisus (Forst.f.) Copel 502 irayensis Copel. 495 jacobsii Holtt. 520 lanceolus (Christ) Copel. 529 larutensis (Bedd.) Ching 489 latifolius (Presl) Copel. 526 latipinna (Benth.) Tard. 558 lepidopodus C.Chr. & Tard. 429 leucadenius Copel. 460 lineatus (Bl.) Tard. & C.Chr. 512 lobatus (Copel.) Copel. 474 loherianus (Christ) Copel. 489 magnus Copel. 465 matutumensis (Copel.) Copel. 467 megaphyllus sensu Copel. 460 megaphyllus (Mett.) Ching 461 melanophlebius (Copel.) Copel. 514 merrillii (Christ) Copel. 529 micans (Brause) Copel. 501 1981] Index to scientific plant names 573 (Cyclosorus) (Cyclosorus) (Davallia) microlonchus sensu Copel. 424 microlonchus (Christ) Copel. 423 mindanaensis (Christ) Copel. 557 morobensis (Copel.) Copel. 501 multiauriculatus (Copel.) Copel. 492 mutabilis (Brause) Copel. 494 nitidulus sensu Copel. 424, 460 nitidulus (Presl) Copel. 430 oblongus (Brause) Copel. 503 obtusifolius (Rosenst.) Copel. 503 oligodictyus (Baker) Holtt. 380 papilio 556 parasiticus (L.) Farw. 559 paripinnatus (Copel.) Copel. 476 philippinarum (Fée) Copel. 512 polycarpus (Bl.) Holtt. 449 polypterus Copel. 471 porphyricola (Copel.) Ching 472 procurrens (Mett.) Copel. 560 productus sensu Copel. 456 productus (Kaulf.) Ching 460 proliferus (Retz.) Tard. 387 protectus (Copel.) Copel. 485 pustulosus Copel. 513 reederi Copel. 456 repandulus (v.A.v.R.) Ching 557 rhombeus sensu Copel. 525 rhombeus (Christ) Ching 529 rigidus (Ridl.) Copel. 460 riparis (Copel.) Copel. 456 rotumaensis St.John 426 rubidus (Hook.) Copel. 532 rubrinervis (Mett.) Copel. 531 sagittifolioides Copel. 471, 472 samarensis Copel. 524 serratus Copel. 460, 461 sessilipinna (Copel.) Ching 479 simplicifolius (J.Sm.) Copel. 490 sogerensis (Gepp) Copel. 428 spenceri (Christ) Copel. 491 stenodontus Copel. 450 strigosissimus (Copel.) Copel. 470 subappendiculatus (Copel.) Copel. 420 subdimorphus Copel. 524 subpubescens sensu Holtt. 558 sumatranus (v.A.v.R.) Ching 558 superbus (Brause) Ching 420 supraspinigerus (Rosenst.) Copel. 395 suprastrigosus (Rosenst.) Copel. 457 tephrophyllus (Copel.) Copel. 480 terrestris (Copel.) Copel. 491 todayensis (Christ) Ching 471 toppingii sensu Copel. 485 triphyllus (Sw.) Tard. 534 truncatus (Poir.) Farw. 429 var. kotoensis H.Ito 460 unitus (L.) Ching 478 urdanetensis (Copel.) Copel. 467 urophyllus sensu Copel. 533 urophyllus (Mett.) Tard. 533 vestigiatus (Copel.) Copel. 453 viridis Copel. 418, 419 warburgii (Kuhn & Christ) Ching 462 weberi Copel. 460 xiphioides (Christ) Copel. 525 Cyrtogonium acuminatum Brack. 325 laciniatum J.Sm. 327 rivulare Brack. 328 Cystodium J.Sm ii, xix, 65, 69, 72, 162, 562 sorbifolium (Sm.) J.Sm. 67, 163*, 405 Cystopteris il, xvili, 335 Davallia J.Sm. ii, iv, xviii, 167, 179, 185, 198, 256, 258 subg. Odontoloma Hook. 235 achillaeifolia Wall. 266 achilleifolia Wall. ex Hook. 269 adiantifolia Hook. 249 amboynensis Hook. 187 biflora Kaulf. 182 biserrata Bl. 194 blumeana Hook. 214, 216 boryana Presl 237, 241 chinensis (L.) J.Sm. 182 chusana (L.) Willd. 182 decipiens Cesati 180 delicatula Christ 237, 239 denhami Hook. 186 didyma Hedwig 183 firmula Baker 193 flagellifera Hook. & Grev. 193 gracilis Bl. 190 hemiptera Bory 237 hosei Baker 191 intramarginalis Cesati 197 kunzeana Hook. 219 lanceolata Baker 235 lobata (Poir.) Desv. 219 longipinnula Cesati 197 luzonica Hook. 191 microcarpa J.Sm 183 oligophlebia Baker 191 philippinensis Harr. 191 pinnata Cav., non Mett. ex Kuhn 193 pulchella (J.Sm.) Hook. 249 repens (Bory) Desv. 237 retusa Cav. 180 serrata Roxb. ex Griff. 193 spec. 253 stenoloba Baker 187 tenuifolia (Lamk) Swartz 182 var. chinensis (L.) Moore 182 var. lata Hook. 182 triquetra Baker 216 veitchii Baker 184 Davallia group Ul, 1x, Xi, Xil, xiv, XViil Davallioideae 256 Davallodes ii, xviii, 258 Dendroconche ii, xvi Dendroglossa il, xvi linneana Fée 325 Dennstaedtia Ii, vil, xviii, 38, 68, 69, 71, 159, 167, 178, 256, 339 ampla (Baker) Bedd. 161 cuneata (Hook.) Moore 161 erythrorachis (Christ) Diels 162 flaccida (Forst.) Bernh. 162 glabrata (Ces.) C.Chr. 162 gomphophylla (Baker) C.Chr. 162 javanica Christ 168 moluccana (BI.) Moore 162 multifida v.A.v.R. 168 paraphysata v.A.v.R. 168 remota (Christ) Diels 162 thombifolia (Baker) C.Chr. 162 scabra (Wall.) Moore 161, 162 scandens (BI.) Moore 162 smithii (Hook.) Moore 162 straminea J.Sm. 169 Dennstaedtia group ii, x, Xi, xiii, XiV Dennstaedtiaceae 178 574 FLORA MALESIANA [ser. II, vol. 1? (Dennstaedtiaceae) subfam. Lomariopsidoideae Holtt. 255, 256 Dennstaedtiales 339 Diacalpe ti, xix aspidioides BI. 158 Diblemma ii, xvi Dichorexia Presl 73, 76 latebrosa Presl 115 Dicksonia L’Hérit. ii, vil, xix, 65, 67, 68, 69, 70, 71, 72, 74, 108, 158, 167 sect. Eudicksonia Hook. & Baker 158, 166 subg. Balantium Hook. 158, 166 ampla Baker 161 antarctica Labill. 69, 160* arborescens L’Hérit. 158, 159, 167 archboldii Copel. 159, 160 assamicum Griff. 165 baranetz Link 165 blumei sensu C.Chr. 159 blumei (Kunze) Moore 69, 159, 160* chrysotricha (Hassk.) Moore 159 copelandii Christ 168 culcita L’Hérit. 167 cuneata Hook. 161 deltoidea Hook. 161 erythrorachis Christ 162 flaccida (Forst.) Sw. 162 glabrata Cesati 162 gomphophylla Baker 162 grandis Rosenst. 159, 161 hieronymi Brause 159, 161 japonica Sw. 162 javanica BI. 168 kingii Bedd. 162 lanigera Holtt, 159, 161 ledermannii Brause 161 linearis Cav. 162, 190 mollis Holtt. 159, 160* moluccana BI. 162 moluccana Roxb., non BI. 163 neglecta Fée 162 nephrolepioides Christ 162 papuana F.v.M. 163 remota Christ 162 repens Bory 235, 237, 241 rhombifolia Baker 162 scabra Wall. ex Hook. 162 scandens Bl. 162 schlechteri Brause 161 sciurus C.Chr. 159, 160, 161 smithii Hook. 162 sorbifolia Sm. 163 straminea Labill. 169 (Dicksonia ) strigosa Thunb. 162 torreyana Brack. 169 zippeliana Kunze 162 Dicksonia group Il, x, Xili, xix Dicksoniaceae 65, 71, 562 Dicksoniales 339, 562 Dicksoniineae 562 Dicranopteris sensu Und. 11, 18 Dicranopteris Bernh. ii, viii, xv, le 5 3B IO 4 SUS Silks S59 subg. Acropterygium 29 subg. Dicranopteris 29 clemensiae Holtt. 31*, 32 crassifolia Nakai 34 curranii Copel. 3, 28*, 29*, B0N3IEI357 dolosa Copel. 26 ferruginea Copel. 34 ferruginea Hosok. 561 lessonii Nakai 31 linearis (Burm. f.) Und. 2, 3, DIE 2963 0S 332533 var. alternans (Mett.) Holtt. 29*, 31*, 33, 35, 36 var. altissima Holtt. 33, 36 var. bidentata v.A.v.R. 561 var. crassifrons v.A.v.R. 561 var. demota Holtt. 33, 36 var. ferruginea (BI.) Holtt. 33, 34, 561 var. inaequalis (Rosenst.) Holtt. 33, 36 var. latiloba Holtt. 33, 34 var. linearis 29*, 31*, 33, 34, 35*, 36 var. montana Holtt. 29*, 33, 36 var. rigida (BI.) Holtt. 33, 34 var. stipulosa Christ 561 var. subferruginea (Hieron.) Nakai 31*, 33, 34, 561 var. subpectinata (Christ) FOltt 297 33555556; 561 var. subspeciosa BS Bik, BB), BG var. tetraphylla (Rosenst.) Nakai 29*, 33, 36 opposita Nakai 32 pectinata (Willd.) Und. 1, 3 pubigera (BI.) Nakai 29%, 31*, 32*, 34 speciosa (Presl) Holtt. 2, 3, PE). Bil, 2944, 3S) tomentosa Copel. 561 Holtt. (Dicranopteris) warburgii sensu Nakai 35 Dictyocline ili, xxi Dictyoglossum J.Sm 289, 291 Didymochlaena li, xix, 335 polycarpa (Bl.) Baker 448 Dimorphopteria Tagawa & Iwats. 507, 516 moniliformis Tagawa & Iwats. 507, 519 Diplaziopsis ti, xviii, 157 Diplazium 11, xviti, 157 Diplopterygium Nakai 11 blotianum Nakai 17 conversum Nakai 17 elmeri Nakai 14 giganteum (Hook.) Nakai 561 laevissimum Nakai 11 longissimum Nakai 14 norrisii Nakai 15 novoguineense Nakai 15 volubile Nakai 13 Diplora il, xviii Dipteris ti, xv, 3 lobbiana (Hook.) Moore v, 536 novoguineensis Posth. 7 Disphenia Pres 73, 76 orientalis Kunze 86 Doodia ii, xviii Doryopteris ii, xvil Drymoglossum ii, xvi Drynaria ti, Xvi Dryoathyrium 11, xviii Dryopolystichum iti, xx Dryopterideae 255 Dryopteridoideae C.Chr. 257 Dryopteris Adans. ii, v, xix, 65, 67, 69, 158, 167, 256, 257, 266; 335; 53365e5Si-eSSSs 339, 507 sect. Abacopteris C.Chr. 507, 509 subg. Cyclosorus C.Chr. 386 subg. Steiropteris C.Chr. 378, 398 abortiva (Bl.) O.Ktze 429 acanthocarpa Copel. 532 acromanes Christ 452, 453 acrostichoides (Christ) v.A.v.R. 523, 524 var. lanceola (Christ) v.A.v.R. 523, 529 var. rhombea _ (Christ) v.A.v.R. 523, 529 acuminata Rosenst. 558 adenophora C.Chr. 455 adenostegia Copel. 466 africana (Desv.) C.Chr. 542 alatella (Christ) C.Chr. 451 albociliata Copel. 560 1981] (Dryopteris) albosetosa Copel. 505 alta Brause 404 amaiensis Rosenst. 536 angustata Copel. 456 angustipes sensu C.Chr. 472 angustipes Copel. 471 appendiculata (BI.) C.Chr. 452 aquatilis Copel. 466 aquatiloides Copel. 520 arborea v.A.v.R. 484 arbuscula sensu Kjellb. & C.Chr. 454 archboldii C.Chr. 485 arfakianus (Baker) C.Chr. 484 arida (D.Don) O.Ktze 555 var. ebeneorachis Copel. 556 arisanensis Rosenst. 351 armata Rosenst. 392 atasripii Rosenst. 504 athyriocarpa Copel. 365 atrispora (Domin) C.Chr. 158, 401 var. varievestita C.Chr. 401 atrospinosa C.Chr. 390 attenuata O.Ktze 549 aureoviridis Rosenst. 353 auriculata (J.Sm.) Ching 412 auriculifera v.A.v.R. 383 aurita (Hook.) C.Chr. 345 austera Brause 502 austrophilippina Copel. 479 backeri v.A.v.R. 349 badia v.A.v.R. 369 bakeri (Harr.) Copel. 464, 516 baramensis C.Chr. 473 bartlettii Copel. 522 basilaris C.Chr. 460 basisora Copel. 409 batacorum Rosenst. 476 var. winkleri Rosenst. 429 batjanensis Rosenst. 467 beccariana (Cesati) C.Chr. 383, 513 beddomei (Baker) O.Ktze 371 var. nadiwonensis v.A.v.R. 371 belensis Copel. 403 benoitiana (Gaud.) v.A.v.R. 500 berastagiensis C.Chr. 423 besukiensis v.A.v.R. 547 bipinnata Copel. 405 bordenii Christ 457, 458, 465 borneensis (Hook.) O.Ktze 528 brachyodus sensu v.A.v.R. 382 Index to scientific plant names (Dryopteris) brassii C.Chr. 371 brooksii Copel. 423 brunnea (Wall.) C.Chr. 347 brunneo-villosa C.Chr. 349 brunnescens C.Chr. 452 bungoensis C.Chr. 487 calcarata (Bl.) O.Ktze 374 calcicola C.Chr. 462 callosa (Bl.) C.Chr. 419 var. sumatrana v.A.v.R. 420 calva Copel. 351 canescens (Bl.) C.Chr. 497 var. sensu C.Chr. 490, 516 var. degenera Christ 522 var. lobata Christ 474 var. novoguineensis Brause 514 var. subsimplicifolia Christ 514 canlaonensis Copel. 479 caudiculata v.A.v.R. 423 caudiculata Rosenst. 466 celebica (Baker) Copel. 523 ceramica Rosenst. non v.A.v.R. 488 cesatiana C.Chr. 513 chamaeotaria Christ 524 chlamydophora Rosenst. 384 christopherseni Christ 421 ciliata (Benth.) C.Chr. ex Wu 375 clemensiae Copel. 521 compacta Copel. 527 conferta Brause 491 confusa Copel. 539 conterminoides C.Chr. 367 contigua Rosenst. 557 cordifolia v.A.v.R. 526 coriacea Brause 368 var. elata Brause 368 costata (Brack.) Maxon 421 crassa Copel. 408 crassifolia (Bl.) O.Ktze 377, 384 var. motleyana (Hook.) v.A.v.R. 381, 384 var. purpureo-lilacina C.Chr. 384 crassinervia C.Chr. 414 creaghii (Baker) C.Chr. 353 cucullata (Bl.) Christ 478 var. mucronata Christ 478 cuspidata (Bl.) Christ 536 cyrtocaulos v.A.v.R. 473 dayi (Bedd.) C.Chr. 353 debilis (Mett.) C.Chr. 448, 522 decursive-pinnata (van Hall) O.Ktze 354 575 (Dryopteris) deltiptera Copel. 420 dentata (Forssk.) C.Chr. 557 diaphana Brause 362 dichrotricha Copel. 453, 482 dichrotrichoides v.A.v.R. 482 dicranogramma v.A.v.R. 419 didymosora (Bedd.) C.Chr. 560 diminuta Copel. 464, 515 dimorpha Brause 501 distans sensu v.A.v.R. 347 distincta Copel. 353, 548 divergens Rosenst. 384 diversifolia v.A.v.R. 547 diversiloba (Presl) Christ 452, 453 var. acrostichoides (J.Sm.) Christ 525 subvar. lanceola Christ 529 subvar. rhombea Christ 529 diversivenosa v.A.v.R. 353 dura Copel. 368 echinata (Mett.) O.Ktze 379 echinospora v.A.v.R. 495 elliptica Rosenst. 474 elmerorum Copel. 485, 486 eminens (Baker) C.Chr. 449 endertii C.Chr. 381 engleriana Brause 366 var. hirta C.Chr. 367 ensipinna Brause 404 erubescens sensu Christ 549 euaensis Copel. 555 euryphylla Rosenst. 511 var. teysmannii v.A.v.R. 511 excrescens Copel. 522 exigua sensu v.A.v.R. 539 exsculpta sensu C.Chr. 530 exsculpta (Baker) Copel. 528 extensa (Bl.) O.Ktze 397, 548 falcatipinnula Copel. 406 farinosa Brause 503 ferox (Bl.) O.Ktze 391 var. calvescens sensu v.A.v.R. 394, 395 var. calvescens Christ 391, 394, 395 var. mitis Rosenst. 395 finisterrae Brause 432 firmula (Baker) C.Chr. 519 flaccida (Bl.) O.Ktze 352 flavo-virens Rosenst. 353 foxii Christ 431 fulgens Brause 403 glanduligera (Kunze) Christ 373 glandulosa sensu Christ 513 576 FLORA MALESIANA [ser. II, vol Rit? (Dryopteris) glandulosa (Bl.) O.Ktze 522 glaucescens Brause 469, 470 glaucostipes (Bedd.) C.Chr. 471 gongylodes sensu v.A.v.R. 386 gracilescens (Bl.) O.Ktze 351 var. calva (Copel.) C.Chr. 351 var. chinensis Christ 359 gracilis Copel. 371 granulosa (Presl) C.Chr. 524 gymnocarpa Copel. 542 gymnopoda (Baker) C.Chr. 361 gymnopteridifrons Hayata 513 habbemensis Copel. 362 hallieri (Christ) C.Chr. 379 hamifera v.A.v.R. 496 harveyi (Mett.) O.Ktze 555 hastatopinnata Brause 470 heterocarpa (BI.) O.Ktze 457 hewittii Copel. 527 himalayensis C.Chr. 412 hirsuta (Kunze ex Mett.) O.Ktze 455 hirtisquamata Hayata 352 hirtopilosa Rosenst. 557 hispidifolia v.A.v.R. 466 hispidula sensu Brause 470 hispidula sensu C.Chr. 481 hispidula (Decne) O.Ktze 557 hispiduliformis C.Chr. 470 holophylla C.Chr. 526, 527 horizontalis (Rosenst.) v.A.v.R. 366 horridipes v.A.v.R. 394 hosei (Baker) C.Chr. 526 hunsteiniana Brause 403 immersa var. ligulata Christ 431 imponens (Ces.) C.Chr. 391 impressa (Desv.) Posth. 548 inclusa Copel. 423 ‘inconspicua Copel. 493 indica var. malayensis (Bedd.) v.A.v.R. 485 indochinensis Christ 359, 360 interrupta sensu. Back. & Posth. 545 invisa (Forst. f.) O.Ktze 502 iridescens v.A.v.R. 522 japonica var. stricta sensu Rosenst. 364 keysseriana Rosenst. 432 kiauensis C.Chr. 550 kinabaluensis Copel. 361 (Dryopteris) klossii (Ridl.) v.A.v.R. 361 koordersii (Christ) C.Chr. 375 korthalsii Rosenst. 526, 527 kotoensis Hayata 460 labuanensis C.Chr. 528 laevis (Mett.) C.Chr. 424 lanceola (Christ) Copel. 529 larutense 437 latebrosa (Mett.) C.Chr. 471 latipinna (Benth.) O.Ktze 558 lauterbachii Brause 367 leucolepis (Presl) Maxon 349 ligulata (Presl) O.Ktze 431 linearis Copel. 369 lineata sensu C.Chr. 528 var. subacrostichoides v.A.v.R. 529 lithophylla Copel. 496 lobangensis C.Chr. 479 logavensis Rosenst. 451, 549 loheriana (Christ) C.Chr. 489 longipes (BI.) O.Ktze 424 longissima var. novoguineensis Rosenst. 548 luerssenii sensu Christ 414 luerssenii (Harr.) C.Chr. 431 luzonica Christ 421, 424 var. puberula Christ 424, 460 macgregorit (Baker) C.Chr. 367 macrotis (Hook.) O.Ktze 527 malayensis C.Chr. 522 maquilingensis Copel. 524 marattioides Alston 402 marthae v.A.v.R. 374 mataanae Brause 365 matutumensis Copel. 467 media v.A.v.R. 352 megalocarpa v.A.v.R. 359, 360 megaphylla (Mett.) C.Chr. 461 megaphylloides v.A.v.R. 452 megaphylloides Rosenst. 481 melanophlebia Copel. 514 menisciicarpa (BI.) Posth. $12, 526 merrillii Christ 529 micans Brause 501 microcarpa v.A.v.R. 371 microloncha Christ 423 mindanaensis Christ 557, 558 mirabilis Copel. 526 mixta Rosenst. 457, 458 mollis (Sw.) Hieron. 557 moluccana C.Chr. 549 monodonta C.Chr. 359 morobensis Copel. 501 (Dryopteris) motleyana (Hook.) C.Chr. 381 var. dulitensis C.Chr. 382 moussetii Rosenst. 346 multiauriculata Copel. 492 multilineata (Bedd.) O.Ktze 461 multiseta (Baker) C.Chr. 349 multisora C.Chr. 359 muricata Brause 39] var. marginata Brause 391, 392 var. obscura Brause 391, 392 mutabilis Brause 494 nephrolepioides C.Chr. 431 normalis (Sod.) C.Chr. 413 norrisii Rosenst. 485 notabilis Brause 402 novoguineensis Brause 469, 470 oblanceolata Copel. 513 oblonga Brause 503 obtusata v.A.v.R. 362 obtusifolia Rosenst. 503 oligodictya (Baker) C.Chr. 380 oligolepia v.A.v.R. 368 oosora (Baker ) C.Chr. 500 ophiura Copel. 409 orbicularis C.Chr. 548 oxyotis Rosenst. 527 paleata Copel. 380 pallescens Brause 383 paraphysata Copel. 481 paraphysophora v.A.v.R. 550 parasitica (L.) O.Ktze 559 var. falcatula Christ 557 paripinnata Copel. 476 patens sensu v.A.v.R. 555 patens (Sw.) O.Ktze 413 pectiniformis C.Chr. 363 peekelii v.A.v.R. 555 peltochlamys C.Chr. 495 penangiana (Hook.) C.Chr. 565 var. calvescens sensu Copel. 393 pennigera (Forst. f.) C.Chr. 421 perakensis 565 var. sumatranus v.A.v.R. 452 perglandulifera v.A.v.R. 454 var. firmior v.A.v.R. 454 perpilifera v.A.v.R. 456, 457 perpubescens Alston 554 persquamifera v.A.v.R. 379 petrophila Copel. 435 1981] (Dryopteris) philippinensis (Baker) Copel. 460 pilosiuscula (Racib.) C.Chr. 522 pilososquamata v.A.v.R. 481 var. obtusata v.A.v.R. 449, 481 pinnata Copel. 375 platyptera Copel. 369 plumosa C.Chr. 360 plurifolia v.A.v.R. 492 polycarpa (BI.) Christ 449 polyotis C.Chr. 464 polytricha v.A.v.R. 380 porphyricola’ sensu C.Chr. 472 presliana Ching 512 procurrens (Mett.) O.Ktze 560 producta (Kaulf.) C.Chr. 460, 461 prolifera (Retz.) C.Chr. 387 propria v.A.v.R. 367 protecta Copel. 485 pseudoamboinensis Rosenst. 558 pseudoarbuscula v.A.v.R. 456 pseudohirsuta Rosenst. 460 pseudomegaphylla v.A.v.R. 452 pseudoreptans C.Chr. 522 pseudostenobasis Copel. 451, 549 pteroides sensu v.A.v.R. 545 pterospora v.A.v.R. 487 pubirachis (Baker) C.Chr. 365 quadriaurita Christ 404 quadriquetra v.A.v.R. 406 ramosii Christ 537 regis Copel. 434 repandula v.A.v.R. 557 repentula Christ 373 rhombea sensu Copel. 525 rhombea (Christ) Copel. 529 ridleyana Brause 501 ridleyi (Bedd.) C.Chr. 361 riedleana (Gaud.) v.A.v.R. 456 rigidifolia v.A.v.R. 369 riparia Copel. 456 robinsonii (Ridl.) C.Chr. 361 roemeriana Rosenst. 499 rosenburghii C.Chr. 484 rubidum (Hook.) C.Chr. 532 rudis (Kunze) C.Chr. 501 rufopilosa Brause 501 sagittifolia (B!.) O.Ktze 487 savaiensis (Baker) C.Chr. 404 (Dryopteris) schizophylla v.A.v.R. 407 schlechteri Brause 407 var. djammuense Brause 407 schultzei Brause 432, 433 sepikensis Brause 484, 485 septempedalis Alston 402 sessilipinna Copel. 479 setigera sensu C.Chr. 348 setigera (Bl.) O.Ktze 349 var. pallida v.A.v.R. 348, 349 simillimum C.Chr. 520 simplicifolia (J.Sm.) Christ 490 singalanensis (Baker) C.Chr. 352 sogerensis Gepp 428 spenceri Christ 491 squamaestipes sensu Hayata 413 squamipes Copel. 364 stegnogramma (BI.) C.Chr. 542 stenobasis C.Chr. 451, 549 stereophylla v.A.v.R. 366 stipellata (Bl.) O.Ktze 492 var. obtusata v.A.v.R. 476 strigosissima Copel. 470 subalpina v.A.v.R. 474 subappendiculata Copel. 420 subattenuata Rosenst. 547 subconformis C.Chr. 519 subdimorpha Copel. 384 subfalcinella v.A.v.R. 485 sublaxa Hayata 351 submollis v.A.v.R. 558 subnigra Brause 369 subpubescens_ (BI.) 558 var. major C.Chr. 558 subulifolia v.A.v.R. 494 subviscosa v.A.v.R. 361, 364 subviscosa Rosenst. 364 sumatrana sensu Troll 430 sumatrana v.A.v.R. 415, 558 sumbawensis C.Chr. 432 superba Brause 420 superficialis v.A.v.R. 549 supraspinigera Rosenst. 395 suprastrigosa Rosenst. 457, 458 supravillosa C.Chr. 362 tandikatensis v.A.v.R. 463 tenompokensis C.Chr. 530 tephrophylla Copel. 480 terrestris Copel. 491 teuscheri v.A.v.R. 381 thelypteris var. squamigera C.Chr. 377 @Chr: Index to scientific plant names BI) (Dryopteris) tibangensis C.Chr. 485 todayensis Christ 419, 471 toppingii sensu C.Chr. 485 toppingii Copel. 500 transversaria (Brack.) C.Chr. 421 var. rodigasiana 426 trichodes Rosenst. 348 trichopoda C.Chr. 380 triphylla (Sw.) C.Chr. 534 truncata (Poir.) C.Chr. 429 tuberculata (Ces.) C.Chr. 407 tylodes (Kunze) Christ 414 uliginosa sensu. Back. & Posth. 349 uliginosa (Kunze) C.Chr. 348 uniauriculata Copel. 467 unita (L.) O.Ktze 477 uraiensis Rosenst. 352 urdanetensis Copel. 467, 479, 521 urophylla sensu v.A.v.R. 513 urophylla (Mett.) C.Chr. 532 var. nitida Holtt. 532 var. novoguineensis Rosenst. 512 var. peraspera v.A.v.R. 526 verruculosa v.A.v.R. 512, 526 var. sumatrana v.A.v.R. 526 vestigiata Copel. 453 villosipes Gepp 369 vinosicarpa v.A.v.R. 381, 382 viscosa (Baker) O.Ktze 360 var. kamborangana C.Chr. 361 wantotensis Copel. 410 warburgii (Kuhn & Christ) C.Chr. 462 wariensis Copel. 403 weberi Copel. 482 williamsii Holtt. 450 xiphioides Christ 525 zippelii Rosenst. 529 Dryopteris group ii, Xi, xii, xix @CChr Eatoniopsis Bommer 73 Edanyoa Copel. 315 difformis Copel. 318, 326, 329 Egenolfia Schott ii, xix, 255, 256, 258, 315, 318, 322 appendiculata J.Sm. 322, 329 bipinnatifida J.Sm. 325 fluviatilis Copel. 323 hamiltoniana Fée 322 intermedia Fée 323 rhizophylla Fée 323 serrulata Fée 323 sinensis Maxon 325 578 FLORA MALESIANA [ser. II, vol. 1° (Egenolfia) vivipara C.Chr. 322 Elaphoglossaceae_ Pichi Ser- molli 255 Elaphoglossum J.Sm. ii, iv, xix, 255250; 25>, 25852289, 314 alstonii Tard.-Blot 297 amblyphyllum Bell 290, 294, 303, 304* angulatum sensu Copel. 307 angulatum (BI.) Moore 292, 296*, 297, 314 angustatum (Schrad.) Hieron. 313 angustifrons Holtt. 293, 303 annamense C.Chr. & Tard.- Blot 293, 299%, 303 apiculatum Holtt. 293, 299 apoense Holtt. 293, 302, 305* arachnoideum Holtt. 294, 303 archboldii Copel. 293, 299, 303 austrosinicum Matt. & Chr. 310 basilanicum Copel. 301 beccarianum C.Chr. 301 blumeanum (Fée) J.Sm. 292, 294, 311, 312*, 313, 314 var. blumeanum 311 var. philippense Christ MS 311 bolanicum Rosenst. 292, 295* borneense (Burck) C.Chr. 314 brassii C.Chr. 309 brevifolium Holtt. 294, 307 brunneum Copel. 292, 293, 295 calanasanicum Holtt. 293, 301 callifolium sensu Holtt. 308 callifolium (BI.) Moore 292, 294, 306*, 307, 308, 314, S17 commutatum sensu v.A.v.R. 307 commutatum (Mett. ex Kuhn) v.A.v.R. 294, 296*, 308, 314 conforme sensu Bl. 308 conforme sensu Copel. 307 conforme (Sw.) J.Sm. 290, 291, 292, 309, 313 copelandii Christ 311 cumingii var. papuanum C.Chr. 304 decurrens sensu v.A.v.R. 303 decurrens sensu C.Chr. 303 decurrens Desv. 314 var. crassum C.Chr. 299 dolichocaulon v.A.v.R. 297 elmeri Copel. 304, 307 (Elaphoglossum) fuscum Copel. 297 gayanum (Fée) Moore 290 gorgoneum (Kaulf.) Brack. 313 habbemense Copel. 292, 295 hellwigianum Rosenst. 292, 295 heterolepium v.A.v.R. 303, S11F 312+, 313" heterostipes Holtt. 293, 294, 301 hirtum (Sw.) C.Chr. 314 idenburgensis Holtt. 293, 298 indrapurae Holtt. 294, 310 junghuhnianum Moore 307 latemarginatum Holtt. 293, 298 laticuneatum Copel. 297 latifolium sensu Bedd. 297 latifolium (Sw.) J.Sm. 314 laurifolium sensu_ v.A.v.R. 307 laurifolium sensu Bedd. 297 laurifolium (Thouars) Moore 308, 314 lepidopodum C.Chr. ex Ogata 304 luzonicum Copel. 294, 304, 305*, 314 macgregorii Copel. 307 malayense Holtt. 294, 308, 310 melanochlamys Holtt. 293, 302 melanostictum sensu Holtt. 302 melanostictum (BI.) Moore 256, 290, 293, 301 merrillii Christ 298 microphyllum v.A.v.R. 314 minahassae v.A.v.R. 297 miniatum (Christ) Christ 294, 311, 312* muscosum 292 negrosensis Holtt. 294, 307 nesioticum Holtt. 293, 302 nigripes Holtt. 293, 302 norrisii (Hook.) Bedd. 293, 300*, 301 novoguineense Rosenst. 294, 309 obtusifolium Bell 303 obtusifolium Brack. 303 obtusifolium J.Sm. 304, 314 ogatai C.Chr. 297 ophioglossoides (Goldm.) Holtt. 293, 298, 301 pallescens Holtt. 293, 303 pellucido-marginatum (Christ) C.Chr. 293, 298, 314 (Elaphoglossum) pellucidum Gaud. 314 peninsulare Holtt. 301 permutatum v.A.v.R. 308 var. mutatum v.A.v.R. 307, 308 petiolatum sensu v.A.v.R. 311 petiolatum (Sw.) Urban 311 planicosta Holtt. 294, 307 pullenii Holtt. 293, 294, 297 pumilum Lam & Verhey 292, 295 rapaense Copel. 292 recommutatum Holtt. 294, 308, 313 reineckei Hieron. & Laut. 307 repens Copel. 293, 298 resiniferum Holtt. 290, 294, 313 robinsonii Holtt. 294, 310 salicifolium (Willd. ex Kaulf.) Alston 311 samoense Brack. 292 sclerophyllum v.A.v.R. 293, 297 societarum Copel. 292 sordidum Christ 294, 309* spathulatum (Bory) Moore 292 spongophyllum Bell ex Holtt. 294, 310 stenolepis Bell ex Holtt. 294, 303 sumatranum Holtt. 294, 310, 311 thamnopteris Holtt. 293, 298 vepriferum Holtt. 293, 302 yunnanense sensu Holtt. 311 yunnanense (Baker) C.Chr. 311 Equisetaceae ii Equisetales 1 Equisetum ii Filicales 11, vii Fourniera Bommer 73, 138 Gisopteris Bernh. 39, 44 Glaphyropteridopsis Ching 389 erubescens (Hook.) Ching 389 Glaphyropteris sect. Cyclogramma H.Ito 411 Gleichenia Smith ti, v, xv, 1, 2, 3, 4, 6, 29, 70, 71, 169 subg. Diplopterygium Holtt. 1, 35:45. 7, 10% 1a Sao» 20, 33 subg. Eugleichenia Diels 6 subg. Gleichenia 1, 2, 3, 4, 6 1981] (Gleichenia) subg. Mertensia Hook. 1, 2, 3, 5, 18, 19, 20 sect. Dicranopteris v.A.v.R. 11, 18, 27 sect. Diplopterygium Diels 11 sect. Heteropterygium Diels 27 sect. Holopterygium Diels 18 sect. I Hook. 11 sect. IJ Hook. 18 sect. III Hook. 27 sect. IV Hook. 27 alstonii Holtt. 5, 23 amboinensis v.A.v.R. 27 amoena v.A.v.R. 20, 26 angustiloba Holtt. 5, 14 arachnoidea (Hassk.) v.A.v.R. 13 arachnoidea Cunn. 13 arachnoides Mett. 13 articulata J.Sm. 230 barbula C.Chr. 26 bifurcata Bl. 561 blotiana C.Chr. 5, 17 bolanica Rosenst. 6, 24, 561 borneensis C.Chr. 6 bracteata Bl. ex Hook. & Baker 21 brassii C.Chr. 6, 24 brevipinnula Holtt. 5, 17 bullata Moore 5, 13, 561 candida Rosenst. 26 caudata Copel. 561 circinnata Sw. 7, 11 var. borneensis Baker 6 clemensiae (Copel.) Holtt. 5, 14, 561 var. clemensiae 14 var. membranacea Holtt. 14 conversa v.A.v.R. 5, 10*, 17 crassifolia Copel. 34 cunninghamii 3 deflexa Holtt. 5, 15 dicarpa R.Br. 4, 7*, 9, 10, 11 var. volcanica Christ 9 dichotoma Hook. 33 var. alternans Mett. 36 var. ferruginea Racib. 34 var. malayana v.A.v.R. 31 var. normalis Mett. 33 var. pubigera Mett. 32 var. rigida Mett. 34 var. tenera Mett. 31 dolosa C.Chr. 26 var. virescens Hieron. ex Brause 27 elmeri Copel. 5, 14 erecta C.Chr. 5, 6, 22, 561 Index to scientific plant names (Gleichenia) excelsa J.Sm. ex Hook. 14 ferruginea Bl. 34 flabellata R.Br. 6, 25 gigantea Wall. ex Hook. 3, 561 glauca auct. 561 glauca v.A.v.R. 14 glauca (Thunb.) Hook. 33 var. arachnoides C.Chr. 13 hallieri Christ 26 hantonensis Wanklyn 2 hermannii R.Br. 33 var. pubigera BI. 32 var. rigida Bl. 34 var. tenera BI. 31 var. venosa BI. 31 hirta BI. 6, 18*, 19*, 25 var. amboinensis (v.A.v.R.) Holtt. 27 var. amoena (v.A.v.R.) Holtt. 26 var. candida (Rosenst.) Holtt. 26 var. hirta 26, 27 var. lanuginosa (v.A.v.R.) Holtt. 27 var. ornamentalis (Rosenst.) Holtt. 27 var. paleacea (Baker) @iGhis22226 var. virescens (Hieron.) Holtt. 27 hispida Mett. ex Kuhn 6, 18*, 19*, 22, 24, 349 hooglandii Holtt. 561 kajewski Copel. 22 koordersii Christ 24 laevigata Hook. 20 var. bracteata v.A.v.R. 21 laevissima Christ 4, 11 linearis (Burm. f.) Clarke 33 var. alternans (non Mett.) Holtt. 35, 36 var. altissima Holtt. 36 var. ferruginea v.A.v.R. 34 var. inaequalis Rosenst. 36 var. montana Holtt. 36 var. normalis v.A.v.R. 33 var. pubigera v.A.v.R. 32 var. rigida v.A.v.R. 34 var. subferruginea Hieron. ex Brause 34 var. tetraphylla Rosenst. 36 loheri Christ 6, 25 var. loheri 25 var. major Holtt. 5, 25 longissima BI. 5, 10*, 12*, 13, 17, 561 matthewii Holtt. 5, 17 3/9 (Gleichenia) microphylla R.Br. 2, 3, 4, 7*, 8*) 9; 14 var. semivestita v.A.v.R. 7 milnei Baker 5, 22, 561 monticola Ridl. 24 norrisii Mett. 5, 15, 16*, 17 var. floccigera C.Chr. 15, 17 novoguineensis Brause 15 oceanica Kuhn 5, 22 opposita v.A.v.R. 32 ornamentalis Rosenst. 27 var. lanuginosa v.A.v.R. 27 paleacea (Copel.) Holtt. 4, 13 papuana Holtt. 14 var. membranacea Holtt. 14 parallela Ridl. 32 peltophora Copel. 2, 4, 6 var. peltophora 6 var. schizolepis C.Chr. ex Holtt. 7 peninsularis Copel. 26 polypodioides (Thunb.) Smith 3 pseudoscandens v.A.v.R. 5, 23 pteridifolia Ridl. 35 pulchra (Copel.) Holtt. 5, 23 reflexipinnula C.Chr. 5, 20, 561 sordida Copel. 5, 15 squamosissima Copel. 9 subpectinata Christ 35 subulata v.A.v.R. 24 sumatrana Holtt. 15 truncata (Willd.) Spr. 5, 18%, 19*, 20, 22 var. bracteata (BI. ex Hook. & Baker) Holtt. 20521522 var. celebica Holtt. 20, 22 var. involuta Holtt. 20, 22 var. plumaeformis (Presl) Holtt. 20, 21 var. truncata 20, 21, 561 venosa (Copel.) Holtt. 5, 20 vestita BI. 6, 19*, 23* var. paleacea Baker 26 volubilis sensu v.A.v.R. 13 volubilis Jungh. 4, 13, 14, 561 var. peninsularis Holtt. 13, 561 var. volubilis 13 vulcanica BI. 2, 4, 7*, 8*, 9, 10 warburgii Christ 34, 35 weatherbyi Fosb. 32 Gleicheniaceae iii, v, vii, X, Xi, Xiii, Xiv, XV, 1-36, 38, 71, 339, 561 580 (Gleicheniaceae) ser. Marginales 71 ser. Superficiales 71 Gleicheniastrum Presl 6 lowei Nakai 7 microphyllum Pres] 7 var. semivestitum Nakai 7 Gleicheniopsis | Gleichenites gracilis Zigno 1, 20 Goniophlebium ii, vi, Xvi Goniopteris 336, 338, 341, 387 appendiculata (Bl.) Presl 452 aspera Presl 512 asymmetrica Fée 452 barbata Fée 391 claiborniana Berry 334, 511 costata Brack. 421 dalhousiana Fée 533 kennedyi (F.v.M.) Bail. 512 lobbiana Fée 419 meniscioides Fée 387 prolifera (Retz.) Presl 387 repanda Fée 533 rigida Ridl. 484 rubrinervis (Mett.) Carr. 531 rudis Ridl. 501 simplicifolia (J.Sm. ex Hook.) Carr. var. vitiensis Carr. 513 Grammatopteridium ii, xvi Grammitidaceae i, ii, ix, Xiii, XIV, XVil Grammitis ti, vi, xvii, 291, 336 blumeana Presl 542 Gymnocarpium ii, xix, 335 Gymnogramme 335, 336, 541 appendiculata Bl. 452, 523 aspidioides Bl. 542 aspidioides Hook. 541 aurita Hook. 345 borneensis Hook. 314 canescens (Bl.) BI. 497 macrotis Kunze 527 pteridiformis Cesati 263 rhizophylla Kaulf. 323 stegnogramma BI. 541 subtrifoliata Hook. 261 totta sensu Bl. 542 Gymnopteris Christ 276, 277, 278, 318 flagellifera Bedd. 325 inconstans Copel. 325 linnaeana Christ 325 naumannii Diels 327 repanda Christ 329 var. quoyana (Gaud.) Diels 327 subrepanda J.Sm. 326 subsimplex Fée 326 Gymnosphaera BI. 73, 76, 115 FLORA MALESIANA (Gymnosphaera) atropurpurea (Copel.) Copel. 118 biformis (Rosenst.) Copel. 118 bipinnatifida Copel. 151 burbidgei (Baker) Copel. 151 dinagatensis Copel. 151 glabra sensu Copel. 151 glabra Bl. 120 gracillima Copel. 123 hewittii Copel. 118 holttumii Copel. 146 hornei Copel. 121 kingii (Copel.) Copel. 121 melanoclada Copel. 121 melanorachis Copel. 121 mollis (Copel.) Copel. 151 obliqua (Copel.) Copel. 146 papuana (Ridl.) Copel. 155 pulchra Copel. 152 ramispina (Hook.) Copel. 117 recommutata Copel. 118 sarawakensis (C.Chr.) Copel. 152 schlechteri (Brause) Copel. 123 squamulata sensu J.Sm. ex Hook. 118 squamulata B1. 152 subbipinnata Copel. 146 trichophora Copel. 151 vexans (Ces.) Copel. 120 Haplodictyum Presl xxi, 334, 335, 336, 507, 509, 511 bakeri (Harr.) Ching 516 bulusanicum Holtt. 515 canescens (BI.) Ching 497 heterophyllum Presl 498, 507, 515 majus Copel. 464, 507, 509 Helicogyratae 71 Helminthostachys ii, xv Hemigramma iii, xx latifolia auct. non Copel. 330 Hemionitis ii, xvii prolifera Retz. 387 Hemipteris iii, xx Hemitelia R.Br. 69, 71, 73, 77 alsophiliformis v.A.v.R. 111 alternans Hook. 145 arfakensis (Gepp) v.A.v.R. barisanica v.A.v.R. 89 bicolor (Copel.) v.A.v.R. 108 capensis sensu Hook. 111 capensis (L. f.) R.Br. 61 caudata (J.Sm.) Mett. 110 caudiculata Rosenst. 106, 108 [ser. II, vol. 1° (Hemitelia) caudipinnula v.A.v.R. 89 confluens v.A.v.R. 102 crenulata Mett. 111 fallax v.A.v.R. 111 var. major v.A.v.R. 111 glaucophylla v.A.v.R. 111 hemichlamydea (Copel.) v.A.v.R. 110 heterochlamydea v.A.v.R. 108 horrida 73 horridipes v.A.v.R. 114 javanica Pres] 110 junghuhniana (Kunze) Mett. 111 var. dissoluta Racib. 111 latebrosa (Wall.) Mett. 111, 115 var. paraphysata v.A.v.R. 115 latipinnula v.A.v.R. 105 ledermannii Brause 140 leptolepia v.A.v.R. 115 manilensis Pres] 110 merapiensis v.A.v.R. 111 montana v.A.v.R. 109 multiflora (Sm.) R.Br. 77 paraphysophora v.A.v.R. 114 perpunctulata v.A.v.R. 114 rudimentaris v.A.v.R. 115 salticola v.A.v.R. 114 singalanensis v.A.v.R. 102 subconfluens v.A.v.R. 102 sumatrana v.A.v.R. 114 tonglonensis v.A.v.R. 108 truncata sensu Christ 139 warthon (Copel.) v.A.v.R. 108 Heterogonium ili, xx, Xxi Heteroneuron argutum Fée 329 cuspidata Presl 325 heteroclitum Fée 325 naumannii Kuhn 327 scalpturatum Fée 321 sinuosum Fée 329 Hicriopteris sensu Ching 11 Hicriopteris Presl 3, 27, 33 astrotricha Copel. 15 blotiana Ching 17 bullata Ching 13 clemensiae Copel. 14 glauca 3 laevissima Ching 11 longissima Ching 14 norrisii Ching 15 novoguineensis Copel. 15 paleacea Copel. 13 speciosa Presl 32 volubilis Ching 13 Histiopteris ti, xix 1981] Index to scientific plant names 581 Holcosorus ii, Xvi Holostachyum ti, xvi Hugona Cav. ex Roem. 451 Humata ii, Xvili Hydroglossum Willd. 39, 45 auriculatum Willd. 54 circinnatum (Burm. f.) Willd. 59 flexuosum (L.) Willd. 53 japonicum (Thunb.) Willd. 51 longifolium Willd. 59 pedatum (Burm. f.) Willd. 59 pinnatifidum Willd. 39, 47, 53 Hydropteridales viii Hydropterideae vili Hymenophyllaceae ii, iv, vii, viii, iX, X, XIll, XV Hymenophyllum ti, xv ramosissimum Ham. ex D.Don 183 Hypodematium Kunze ili, xx, 331, 333 Hypolepis ii, xix, 38 Hypopeltis marginifera Bory 386 propinquoides Bory 386 Isoetaceae 11, 63-64, 562 Isoetales 63 Isoetes L. ti, 63, 562 habbemensis Alston 62*, 64 hopei Croft 562 hystrix Durieu 64 lacustris L. 64 neoguineensis Baker 64 var. rheophila Croft 562 philippinensis Merr. & Perry 64 sp. 562 stevensii Croft 562 Isoetites 64 Isoloma J.Sm. ii, xix, 178, 198, 229, 230 angustum (Copel.) Tard.-Blot 232 auriculatum (v.A.v.R.) Tagawa 231 dicksonioides (Christ) Tard.- Blot 253 divergens (Hook. & Grev.) J.Sm. 233 fuligineum (Copel.) Copel. 230 querinianum (Gaud.) Fée 230 induratum (Baker) Tard.-Blot 230 jamesonioides (Baker) Tagawa 232 lanuginosum J.Sm. 253 lindsayae (Christ) Tard.-Blot 253 (Isoloma) ovatum (J.Sm.) Pres] 231 pellaeiforme (Christ) Tard.- Blot 232 schizolomae (v.A.v.R.) Tagawa 230 walkerae (Hook.) Pres] 229 Kaulinia Nayar 291 Klukia 37 Kylikipteris 65 Lastrea sensu Copel. 414 Eastrea Bory? 335," 336;- 337, 338, 341, 386, 397 alta (Brause) Copel. 404 attenuata Brack. 408 beddomei (Baker) Bedd. 371 belensis Copel. 403 bipinnata Copel. 405 brassii (C.Chr.) Copel. 373 calcarata var. ciliata Bedd. 375 calva (Copel.) Copel. 351 caudiculata Pres] 547 cavitensis Copel. 421 ciliata Hook. 375 confusa Copel. 539 conterminoides (C.Chr.) Copel. 367 coriacea (Brause) Copel. 368 costulisora Copel. 409 crassa Copel. 408 crassifolia (Bl.) Moore 384 dayi (Bedd.) Bedd. 353 decurrens J.Sm. 354 dichrotrichoides (v.A.v.R.) Copel. 482 dryopteroidea Copel. 158 dura (Copel.) Copel. 368 erubescens sensu Copel. 549 eugracilis Copel. 371 exigua J.Sm. 539 fairbankii Bedd. 377 falcatipinnula (Copel.) Copel. 406 finisterrae (Brause) Copel. 432 flaccida (Bl.) Moore 352 flavo-virens (Rosenst.) Copel. 353 foxii (Christ) Copel. 431 gracilescens Bedd. 359, 371 var. glanduligera Bedd. 373 gracilescens (Bl.) Moore 351 grammitoides (Christ) Copel. 373 gymnocarpa Copel. 542, 543 hirsutipes (Clarke) Bedd. 359, 360 immersa (Bl.) Moore 547 (Lastrea) keysseriana (Rosenst.) Copel. 432 klossii Holtt. 361 leucolepis Pres] 349 ligulata Pres] 431 macgregorii (Baker) Ridl. 367 malaccensis Presl 545, 548 microchlamys de Vriese 449 mingendensis Gilli 433 mixta (Rosenst.) Copel. 457 nephrodioides Bedd. 384 nervosa (Fée) Copel. 539 notabilis (Brause) Copel. 402 novoguineensis (Brause) Copel. 469 ophiura Copel. 409 petrophila Copel. 435 philippina Presl 431 platyptera (Copel.) Copel. 369 pyrrhorhachis sensu Copel. 347 quadriaurita (Christ) Copel. 404 regis Copel. 434 ridleyi Bedd. 361 robinsonii Ridl. 361 singalanensis (Baker) Bedd. 352 squamipes (Copel.) Copel. 364 subdimorpha (Copel.) Copel. 384 subnigra (Brause) Copel. 369 tenericaulis (Hook.) Moore 348 thelypteris var. squamigera Bedd. 377 tylodes (Kunze) Moore 413 unidentata Bedd. 359 uraiensis (Rosenst.) Tagawa 352 varievestita (C.Chr.) Copel. 401 verrucosa Presl 547 viscosa (Baker) Bedd. 360 viscosa J.Sm. 360 wantotensis Copel. 410 wariensis Copel. 403 williamsii Copel. 450 Lastreopsis ili, xxi Lecanopteris ii, xvi Lemmaphyllum ii, xvi Lepidocaulon ili, xx Lepisorus Ii, xvi Leptochilus Kaulf. ti, xvi, 255, 277, 318 sect. Bolbitis Christ 255 sect. Lomagramma_ Christ 255 582 (Leptochilus) angustipinnus Hayata 321 celebicus (Baker) C.Chr. 523 cuneatus R.Bonap. 285 cuspidatus (Presl) 3255327) var. crenatus Rosenst. 321 var. marginalis Rosenst. 327 var. quoyanus C.Chr. ex v.A.v.R. 327 decurrens 330 diversifolius C.Chr. 326 exsculptus (Baker) C.Chr. 528 heteroclitus C.Chr. 325 var. eurybasis Christ 326 var. foxworthyi Christ 326 var. inconstans Christ 326 var. linnaeanus Christ 325 hydrophyllus Copel. 326 inconstans Christ 326 linnaeanus Fée 325 lomarioides Bl. 279 var. pteroides v.A.v.R. 283 malaccensis C.Chr. 326 modestus C.Chr. 325 naumannii C.Chr. 327 novoguineensis Brause 281 oligodictyus (Baker) C.Chr. 380 ovatus Copel. 330 perakensis (Bedd.) C.Chr. 281 reimersii Rensch 321 rivularis C.Chr. 328 scalpturatus C.Chr. 321 simplicifolius Holtt. 326 stolonifer Christ 330 sumatranus v.A.v.R. 326 x trifidus v.A.v.R. 319, 320*, 330 virens C.Chr. 321 zollingeri Fée 326 Leptogramma J.Sm. 337, 540 amabilis Tagawa 543 aurita (Hook.) Bedd. 345 celebica Ching 543 decursive-pinnata (van Hall) J.Sm. 354 gymnocarpa (Copel.) Ching $42 petiolata Ching 542 totta sensu Bedd. 542 Leptopteris ii, xv Leucostegia ii, xviii Lindsaea Dryander ii, v, vii, xix, 177, 178, 179, 198 group A 200 group B 200, 201 group C 200, 201, 203 ©@Chr- FLORA MALESIANA (Lindsaea) group D 200, 202 group E 200, 202 group F 200, 202 group G 200, 203 sect. Paralindsaea 178 sect. Sambirania Tard.-Blot 229 subg. Lindsaea 204 sect. Davalliastrum 204 sect. Isoloma (J.Sm.) Kramer 177, 199, 202, 230 sect. Lindsaea 178, 199, 202, 225 sect. Osmolindsaea Kramens tla liss 199; 200, 227, 229 sect. Psammolindsaea Kramer 199, 200, 229 sect. Schizoloma (Gaud.) Kramer 177, 178, 199, 200, 204, 230 sect. Stenolindsaea Kramer 199, 202, 233 sect. Synaphlebium (J.Sm.) Diels 199, 201, 204, 213, 216 sect. Temnolindsaea Kramer 199, 201, 213, Dilfer 233 sect. Tropidolindsaea Kramen 17759199" 200, 229 subg. Odontoloma (Hook.) Kramer 203, 204, 235 sect. Lindsaenium (Fée) Kramer 199, 244, 564 sect. Odontoloma 199, 235, 244, 564 sect. Penna-arborea Kramer 199, 203, 249 sect. Pseudolancea Kramer 199, 244 acutifolia Desv. 253 adiantifolia (Hook.) Copel. 249 adiantoides (Bl.) Kuhn 249 adiantoides J.Sm. 200, 215*, 229, 564 alpestris v.A.v.R. 249, 251 ambigens Cesati 218 amboynensis (Hook.) Mett. ex Kuhn 187, 253 angusta Copel. 240 apiculata Kunze 228 apoensis Copel. 203, 240, 242* arcuata Kunze 178, 223 azurea Christ 201, 223 var. mambae v.A.v.R. 223 [ser. Il, volar (Lindsaea) baker (C:Chr.)§@iehraw202. 235 var. bakeri 209*, 235 var. pycnophylla Kramer 235 bantamensis Bl. 237, 239 billardieri (Gaud.) Carruth. ex Seemann 211 binervata C.Chr. 249 bipinnata Roxb. 253 blanda Mett. ex Kuhn 251 blumeana (Hook.) Kuhn 216 bonii Christ 206 borneensis Hook. ex Baker 202, 225, 254 boryana (Presl) Brause 239 bouillodii Christ 200, 204, 209*, 254, 563 brachypoda 178 brevipes Copel. 234 bullata v.A.v.R. 228 calomelanos Kunze 228 cambodgensis auct. 204 cambodgensis Christ 563 canaliculatipes v.A.v.R. 214 capillacea Christ 203, 238%, 243 carvifolia Kramer 203, 241, 242*, 564 ceramica v.A.v.R. 218 chienii Ching 178, 207, 208 chinensis (L.) Mett. ex Kuhn 182, 253 colobodes Kunze 234 commixta Tagawa 207 ‘concinna’ 178 concinna J.Sm. 233 copelandi C.Chr. 206 cordata (Gaud.) Desv. 253 crassipes Rosenst. 228 crenulata Fée 234 crispa Baker 201, 217, 224 cultrata auct. 227, 228 cultrata (Willd.) Swartz 201, 202, 217, 222 map var. concinna (J.Sm.) Domin 233 var. lobbiana (Hook.) Becc. 234 var. pallens Hook. 228 var. parvula Holtt. 228 var. plumula (Ridl.) Holtt. 228 var. securifolia (Presl) Baker 233 var. varia Copel. 228 cultripinna Copel. 237, 239 cuneifolia Pres] 180, 253 cyathicola Copel. 249, 251 davallioides auct. 218 1981] (Lindsaea) davallioides Bl. 219 f. parallelogrammoides v.A.v.R. 220 decomposita auct. 218 decomposita Willd. 178, 222 f. longipinnula” v.A.v.R. 218 f. minor v.A.v.R. 218 var. davallioides (Bl.) Domin 219 decrescens Copel. 253 delicatula Christ 237 denhami (Hook.) Mett. ex Kuhn 186, 253 diplosora v.A.v.R. 245 var. acrosora C.Chr. 245 divergens Hook. & Grev. 202, 230, 233 doryphora Kramer 178, 202, 226*, 227 map, 244, 253 ensifolia Swartz 177, 201, 204, 211 ssp. agatii (Brackenr.) Kramer 211 ssp. coriacea (v.A.v.R.) Kramer 178, 211, 212 ssp. ensifolia 178, 204, 211, 73 VAI) var. heterophylla (Dryand.) Benth. 210 erecta Mirbel 211 fissa Copel. 203, 238*, 243, 254 flabellulata Dryand. 206 var. gigantea Hook. 208 var. polymorpha (Hook. & Grev.) Hook. 206 foersteri Rosenst. 237, 239 furcata Copel. 218 gigantea (Hook.) C.Chr. 208 glandulifera v.A.v.R. 203, 236, 241*, 252 map gomphophylla Baker 200, 205*, 207 gracilis B1. 233 var. major Mett. ex Kuhn 234 gracillima Copel. 234 grandifolia J.Sm. 253 griffithiana Hook. 211 gueriniana (Gaud.) Desv. 202, 230, 231*, 564 guianensis (Aubl.) Dryand. ssp. lanceastrum Kramer 224 havicei Copel. 240 hemiacroscopica Kramer 200, 204, 205* heterophylla Dryand. 201, 205*, 210 map, 212 Index to scientific plant names (Lindsaea) heterophylla Prentice 210 hewittii Copel. 202, 224 hosei C.Chr. 253 humilis Kuhn 229 hymenophylloides auct. 243 hymenophylloides Bl. 237, 239 impressa Christ 224 indurata Baker 230 integra Holtt. 178, 201, 221*, 223 intermedia Hook. 222 var. minor Hook. 222 interrupta (Roxb.) Morton 563 jamesonioides Baker 202, 232 japonica (Baker) Diels 229 javanensis BI. 200, 208 kinabaluensis Holtt. 224 kingii Copel. 201, 213, 215* kjellbergii C.Chr. 213 klotzschiana Moritz. 213 lancea auct. 244 lancea auct. spec. Asiat. 227 lancea (L.) Bedd. 225 lanceolata Labill. 211 lanuginosa Wall. ex Hook. 253 ledermannii Brause 235 linearis 178 lobata Poir. 202, 219 var. incisa Mett. ex Kuhn 219 lobbiana Hook. 234 loheriana Christ 228 longa Copel. 237, 239, 253, 254 longifolia Copel. 202, 219, Dies longipes C.Chr. & Tard.-Blot 206 longissima Christ 245 lowei hort. 253 lucida BI. 202, 231*, 233 ssp. brevipes (Copel.) Kramer 234 ssp. lucida 234 lunulata v.A.v.R. 236 macraeana (Hook. & Arn.) Copel. 178, 239, 253 malayensis Holtt. 201, 217 marginata Brause 249 var. falcata Brause 251 var. marginata 251 media R.Br. 201, 208, 209* membranacea Kunze 211 merrillii Copel. 203, 241 ssp. merrillii 238*, 241 microstegia Copel. 203, 204, 238*, 246, 247*, 248 583 (Lindsaea) modesta Kramer 202, 221*, 225 monocarpa Rosenst. 203, 245 monosora sensu Rosenst. 245 monosora Copel. 245 montana Copel. 206, 207 multisora v.A.v.R. 201, 213, 215* napaea v.A.v.R. 201, 202, 217 natunae Baker 201, 214, 215* nitens Bl. 222 ‘nitida’ 178 nitida sensu Holtt. 222, 223 nitida Copel. 222 oblanceolata v.A.v.R. 178, 203, 236, 244 oblongifolia Reinw. ex Hook. 23H] obscura Brause 202, 224 obtusa J.Sm. 178, 202, 218, DIO; 22224 odorata Roxb. 178, 200, 227, 228 oligoptera Kunze 211 orbiculata (Lamk) Mett. ex Kuhn 177, 200, 201, 204, 206, 207, 254 var. commixta (Tagawa) Kramer 206, 207, 208, 254 var. gigantea (Hook.) Mett. ex Kuhn 208 var. integra Mett. ex Kuhn 254 var. odontosorioides Copel. 204, 206 var. orbiculata 205*, 206, 207 var. polymorpha (Hook. & Grev.) v.A.v.R. 206 var. sumatrana_ Rosenst. 204 ovata J.Sm. 202, 231 papuana Copel. 201, 223 parallelogramma v.A.v.R. 178, 202, 220 map, 221* parasitica (Roxb.) Hieron. 203, 227, 244, 253, 254 parishii Baker 253, 266, 269 ‘pectinata’ 178 pectinata auct. 236 pectinata Bl. 237, 239 f. dimorpha Rosenst. 239 var. brevipinnula Rosenst. 246 pectinata (Bory) Thw. 237 pellaeiformis Christ 202, 231 pentaphylla Hook. 211 philippinensis Kramer 202, 232 584 FLORA MALESIANA [ser. II, vol. 1° (Lindsaea) (Lindsaea) (Lindsaea) pinnata (Cav.) Mett. ex roemeriana’ Rosenst. 203, vieillardii 178 Kuhn 193, 253 209*, 252 viridis 178 var. bipinnata Mett. ex Kuhn 191, 253 plumula Ridl. 228 polyctena Kramer 201, 214, 215*, 216 map, 564 polymorpha Wall. ex Hook. & Grev. 206 portoricensis 178 prolongata 178 propria v.A.v.R. 234 pteroides Desv. 211 pulchella (J.Sm.) Mett. ex Kuhn 203, 249, 250*, 251 map var. blanda (Mett. ex Kuhn) Kramer 178, 249, 250*, 251 map var. falcata (Brause) Kramer 249, 250, 251 map var. lomatosora Kramer 249, 250*, 251 map var. pulchella 249, 250*, 251 map ramosii Copel. 201, 219 recurvata (Hook.) Hook. 222 regularis Rosenst. 204, 246, 247* repens (Bory) Thw. 178, 203, 239; 237,238". 241, 253, 564 f. angusta (Copel.) Kramer 240 f. truncatiloba Rosenst. 237, 239 var. delicatula Kramer 237, 239 var. hemiptera v.A.v.R. 237, 239 var. intermedia Christ 237, 239 var. laciniata Mett. ex Kuhn 236 var. pectinata (Bl.) Mett. ex Kuhn 238*, 239, 240 var. pseudohemiptera v.A.v.R. 239, 240 var. sessilis (Copel.) Kramer 178, 237, 238%, 239, 254 var. submarginalis Kramer 238*, 239, 240 retusa (Cav.) Merr. 180, 253 rhombifoliolata v.A.v.R. 251 rigida J.Sm. 203, 204, 244, 245, 246, 254, 564 f. acutata v.A.v.R. 245 (Christ) (Bory) rosenstockii Brause 203, 204, 238*, 248, 564 roxasii Copel. 237 sagincola Wagner & Grether 237), 239 sarasinorum Christ 222 sarawakensis Kramer 203, 245, 564 scandens Hook. 244 var. terrestris Holtt. 178, 227 schizoloma Ettingsh. 211 schlechteri Brause 235 schultzei Brause 218, 219 securifolia Presl ex Gold- mann 233 seemannii J.Sm. 229 sepikensis Brause 245 sessilis Copel. 237, 239 sinuato-crenata v.A.v.R. 218, 219 striata Bl. 254 subalpina auct. 217 subalpina v.A.v.R. 202, 218, DOA sublobata Kunze 211 subsemilunularis 236 subtripinnata Copel. 208 tenera auct. 204 tenera Dryand. 207, 254 var. commixta (Tagawa) Iwats. 207 var. gigantea Holtt. 208 tenuifolia auct. 214 tenuifolia Bl. 201, 216 map, 564 tetragona Kramer 201, 214, 563 var. brassiana Kramer 214 var. tetragona 214 trapeziformis auct. 227 trapeziformis Dryand. 198, 225 trapezoidea Copel. 222 trichophylla Copel. 235 tricrenata Baker 254 trilobata Baker 253 trimarginata C.Chr. 229 tripartita Bl. 218, 219 triplosora v.A.v.R. 245 triquetra (Baker) Christ 216 tropidorachis v.A.v.R. 229 variabilis Hook. & Walker Arnott 210 versteegii (Christ) v.A.v.R. 203, 204, 238*, 248, 564 v.A.v.R. (Hook.) vittata Zoll. & Mor. 253 vrieseana Rosenst. 231 walkerae Hook. 200, 229, 230 map werneri Rosenst. 203, 238%, 252 map wollastonii v.A.v.R. 252 Lindsaea group ii, ix, X, Xi, XIV, xix, 177-254, 563 Lindsaenium Fée 198, 244 rigidum (J.Sm.) Fée 245 Lindsaya (sphalm.) 198 Lindsaynium Fée 198, 244 Lomagramma sensu Copel. 275 Lomagramma J.Sm. ii, xx, 255, 256, 257, 2585325952765 278 angustipinna Copel. 278, 279, 281 articulata Copel. 276 bipinnata Copel. 276 borneensis (erron.) 285 brassii Holtt. 277, 279, 287, 289 brooksii Copel. 279, 285 copelandii Holtt. 279, 287, 289 cordata Copel. 283, 285 cordipinna Holtt. 277, 278 guianensis (Aubl.) Ching 257, 289 leucolepis Holtt. 279, 285 lomarioides (Bl.) J.Sm. 278, 279, 289 matthewii (Ching) Holtt. 289 melanolepis v.A.v.R. 278, 279, 282250 2862875 288*, 289 merrillii Holtt. 278, 279, 281 novoguineensis (Brause) C.Chr:. 2783279 5aest 282* pedicellata Copel. 283 perakensis Bedd. 278, 279, 280*, 281, 283* polyphylla Brack. 255, 277 praestantissimum (Bory) Gries. 289 pteroides J.Sm. 277, 278, 279, 281, 283*, 289 var. negrosensis Copel. 287 var. subcoriacea Copel. 283 sinuata C.Chr. 277, 278, 279, 282*, 284*, 285, 289 f. papuana C.Chr. 279, 285, 287 sorbifolia (Willd.) Ching 289 1981] (Lomagramma) sp. 285 subcoriacea Copel. 283 sumatrana v.A.v.R. 277, 278, 279, 283*, 285 Index to scientific plant names Lycopodium ti Lycopsida ii Lygodiaceae 39 Lygodictyon J.Sm. 45 Lygodium Swartz ii, v, xv, 37, wilkesiana (Brack.) Copel. 289 Lomaria 266 aculeata Bl. 266, 267, 269 gracilis Bl. 269 ludens Fée 275 polymorpha Zoll. & Mor. 269 spectabilis Kunze 262 variabilis sensu BI. 262 Lomariopsidaceae Alston 255, 256 Lomariopsis Fée ii, xx, 255, 256) 25715 2585 259; 261, 263, 265, 266, 267, 276, PINT, Pass) sect. Eulomariopsis Mett. ex Kuhn 259 sect. Stenochlaena Mett. 267 brackenridgei Carr. 261 cochinchinensis Fée 259, 262 hiigelii Presl 263 intermedia (Copel.) Holtt. 260*, 261, 317* kingii (Copel.) Holtt. 261, 262226355177 leptocarpa sensu Holtt. 262 leptocarpa Fée 267, 270 lineata (Presl) Holtt. 259, 261, 262, 270 ludens Fée 267 oleandrifolia (Brack.) Mett. 261 palustris (Hook.) Mett. 259 papyracea Copel. 262 raciborskii (C.Chr.) Holtt. 262 setchellii (Maxon) Holtt. 261 smithii Fée 262 sorbifolia Christ 257, 259, 261, 266 spectabilis (Kunze) Mett. 260*, 262 spinescens Fée 267 subtrifoliata (Copel.) Holtt. 261, 287 variabilis (Willd.) Fée 256, 259 Lomariopsis group ii, vi, ix, x, Xxli, x1x, 255-330, 564 Lophidium Rich. 39, 40 Lophosoria 562 Lophosoriaceae 562 Loxogramme Il, xvi Loxoscaphe li, xvii Luerssenia ili, xxi Lycopodiaceae ii Lycopodiales ti 38, 39, 40, 44, 561 auriculatum (Willd.) Alston 45, 46, 54, 55*, 57 basilanicum Christ 59 borneense v.A.v.R. 45, 46, AW, SRY Sif f. samarindae v.A.v.R. 55 chaerophylloides Desv. 51 circinnatum (Burm. f.) Sw. 39, 46, 47*, 58, 59 var. cristatum v.A.v.R. 59 var. monstruosum v.A.v.R. 59 var. semihastatum Fosb. 54 var. trifurcatum Christ 54 cochinchinense Desv. 51 derivatum v.A.v.R. 59 dichotomum sensu Bedd. 59 dichotomum (Cav.) Sw. 59 digitatum Pres] 59 dimorphum Copel. 45, 46, 52*, 54 dissectum Desv. 51 flexuosum sensu Gaud. 54 flexuosum sensu Holtt. 51 flexuosum sensu Prantl 54 flexuosum (L.) Sw. 46, 51, 5255 535562 var. setulosum Tard. & @Chi2s3 japonicum (Thunb.) Sw. 37, 39, 46, 50*, 51 f. elongata v.A.v.R. 51 f. microstachya (Desv.) Marda cae: Chri kingii Copel. 51 longifolium (Willd.) Sw. 46, A]z S659; SOLE SOL matthewii Copel. 61 mearnsii Copel. 51 merrillii Copel. 46, 60*, 61 microphyllum Link 51 microphyllum (Cav.) R.Br. 37, 39, 46, 47*, 48*, 49*, 61 microstachyum Desv. 51 moszkowskii Brause 61 novoguineense Rosenst. 54 pedatum (Burm. f.) Sw. 59 pinnatifidum sensu Prantl 47 pinnatifidum sensu Racib. 51 pinnatifidum Sw. 53 polystachyum Wall. ex Moore 37, 45, 46, 47*, 50* pubescens Kaulf. 51 585 (Lygodium) reticulatum Schkuhr 46, 61 salicifolium Pres] 37, 46, 51, S37 0 scandens (L.) Sw. 39, 47 var. intermedium Ces. 47 var. microphyllum (Cav.) Luerss. 47 semibipinnatum R.Br. 53 semihastatum sensu v.A.v.R. 55 semihastatum Desv. 54 serrulatum Bl. 53 tenue Bl. 51 teysmannii v.A.v.R. 59 trifurcatum sensu v.A.v.R. 54 trifurcatum Baker 45, 46, 54 versteegii Christ 46, 61, 562 Macroglossum ii, xv Macrothelypteris sensu Pichi Sermolli 343 Macrothelypteris (H.Ito) Ching 324, 333, 339, 340, 345, 347, 374 leucolepis (Presl) Ching 349 multiseta (Baker) Ching 348, 349 oligophlebia (Baker) Ching 347 ornata 348 paludosa (Bl.) Love & Love 347 polypodioides (Hook.) Holtt. 344*, 345, 348, 349 setigera (BI.) Ching 344*, 348, 349 torresiana (Gaud.) Ching 335, 344*, 345, 348, 349 uraiensis (Rosenst.) Love & Love 352 viridifrons 348 Marattia ll, xv Marattiaceae Ii, X, Xll, XV Marattiales ii, vil Marsilea iii, Xx1 Marsileaceae ili, vill, ix, Xii, XXi Matonia ii, iv, xv, 71 Matoniaceae il, X, Xilil, xv, 38 Meniscium Schreb. 335, 337, 338, 531, 534 sect. Ampelopteris Iwats. 387 beccarianum Cesati 513 cumingii Fée 534 cuspidatum Bl. 536, 537 fragile (Z. & M.) Kunze 512 hosei Baker 526 var. sumbensis v.A.v.R. 523 kennedyi F.v.M. 512 586 FLORA MALESIANA [ser. II, vol. 1° (Meniscium) lineatum (BI.) Ktze 512 liukiuense Christ 537 parishii Bedd. 535 proliferum sensu Hook. 526 proliferum (Retz.) Sw. 387 pseudoarfakianum (Hosok.) Hosok. 454 salicifolium Wall. ex Hook. 535 sp. Moritzi 512 stenophyllum Baker 528, 529 thwaitesii Hook. 535 triphyllum Sw. 534 var. parishii Bedd. 535 Menisorus Alston 340, 374 Meringium xv Merinthosorus il, Xvi Mertensia Roth 3 Mertensia Willd. 3, 18, 27 arachnoides Hassk. 13 crassifolia Pres] 34 laevigata Willd. 20 lessonii A.Rich. 31 plumdeformis Presl 21 pteridifolia Presl 33 spissa (non Fée) var. pubigera Nakai 32 truncata Willd. 3, 20, 21 Mesochlaena R.Br. 436, 437 asplenioides J.Sm. 449 javanica R.Br. ex Mett. 437, 449 larutensis 489 var. borneensis 489 polycarpa 449 sumatrensis v.A.v.R. 489 talamauensis v.A.v.R. 449 toppingii Copel. 449 Mesoneuron Ching 377, 378 (Bedd.) v.A.v.R. v.A.v.R. (BI.) Bedd. 437, attenuatum (O.Ktze) Ching 377, 549 chlamydophorum (C.Chr.) Ching 384 crassifolium (Bl.) Ching 384 echinatum (Mett.) Ching 379 hallieri (Christ) Ching. 379 paleatum (Copel.) Ching 380 persquamiferum (v.A.v.R.) Ching 379 teuscheri (v.A.v.R.) Ching 381 trichopodum (C.Chr.) Ching 380 wantotense sensu Holtt. 403 Mesophlebion Holtt. 335, 340, BAIENS 425s 528 S771.) 386, 397, 398 (Mesophlebion) subg. Plesioneuron Holtt. 377, 378, 397 arenicola Holtt. 378, 379, 381, 383 auriculiferum (v.A.v.R.) Holtt. 379, 383 beccarianum (Ces.) Holtt. 376*, 378, 379, 381, 383, 385 caroli Holtt. 378, 380 chlamydophorum (C.Chr.) Holtt. 341, 378, 379, 383, 384 crassifolium (BI.) Holtt. 332*, 334, 378, 379, 382, 384 dulitense Holtt. 379, 382 echinatum (Mett.) Holtt. 378, 379 endertii (C.Chr.) Holtt. 376*, 377, 379, 381 falcatilobum Holtt. 378, 379, 382 var. apiculatum Holtt. 382 var. falcatilobum 382 hallieri (Christ) Holtt. 378, 379 motleyanum (Hook.) Holtt. 377, 378, 379, 381, 385 oligodictyon (Baker) Holtt. 376*, 377, 378, 380 pallescens (Brause) 383 persquamiferum Holtt. 378, 379 rufescens Holtt. 379, 383 teuscheri (v.A.v.R.) Holtt. 378, 379, 381, 383 trichopodum (C.Chr.) Holtt. 376*, 378, 379, 380 vinosicarpum (v.A.v.R.) Holtt. 381 Mesosorus Hassk. 11, 18, 27 arachnoides Hassk. 13 Metathelypteris (H.Ito) Ching 333, 334, 339, 342, 350 dayi (Bedd.) Holtt. 332%, 3505351352 decipiens (Clarke) Ching 351 flaccida (BI.) Ching 350*, 351 gracilescens (BI.) Ching 350*, 351, 352 singalanensis (Baker) Ching 350*, 351, 352 var. singalanensis 352 var. surbeckii Holtt. 351, 352 uraiensis (Rosenst.) 350*, 351, 352 Metaxya 65, 72, 291, 562 Metaxyaceae 562 Holtt. (v.A.v.R.) Ching Metaxyineae 562 Mezonevron Desf., non Unger 378 Microlepia ii, iv, xix, 169, 185, 256 sect. Tapeinidium Presl 184 biflora (Kaulf.) Mett. 182 biserrata (Bl.) Pres] 194 chinensis (L.) Mett. 182 denhami (Hook.) Moore 186 gracilis (Bl.) J.Sm. 190 philippinensis (Harr.) Copel. 191 pinnata (Cav.) J.Sm. 193 strigosa (Thunb.) Pres] 162 tenuifolia (Lamk) Mett. 182 tenuis Brack. 186 Microschizaea Reed 40 fistulosa Reed 44 hallieri Reed 44 malaccana Reed 44 Microsorium li, Xvi pteropus (BI.) Ching 291 punctatum (L.) Copel. 291 Microstaphyla Pres] 255 Mohria 37, 39 caffrorum (L.) Desv. 61 Monachosorum ii, xviii Monogramma iii, XX1 Myrmecophila ti, xvi Nannothelypteris Holtt. 340, 343, 537 camarinensis Holtt. 538, 539* inaequilobata Holtt. 538, 539*, 540 nervosa sensu Holtt. 540 nervosa (Fée) Holtt. 538, 539* aoristisora (Harr.) Holtt. 538 philippina (Presl) Holtt. 538, 539* Nathorstiana 63, 64 Nematopteris tl, xvii Neocheiropteris ti, Xvi Nephelea 562 Nephrodium Schott 258, 335, 336; 337; 3383935459437. 550 abruptum sensu Hook. 461 abruptum Bedd. 462 acrostichoides Desv. 456 alatellum Christ 451 amboinense var. subglandulosum Baker 454 angustifolium Presl 481, 557 appendiculatum (Bl.) Racib. 452 aridum (D.Don) J.Sm. 555 arutense Bedd. 489 1981] (Nephrodium) attenuatum Baker 549 bakeri Harr. 516 basilare Pres] 461 beccarianum Cesati 383 beddomei Baker 371 brachyodes sensu Baker 382 calcaratum (BI.) Hook. 360, 374 callosum (BI.) Keys. 419 caudiculatum J.Sm. 423, 461 conioneuron Fée 545, 548 creaghii Baker 353 crinipes sensu Ridl. 380 cucullatum (BI.) Baker 478 cuspidatum Pres|l 325 debile Baker 522 decursive-pinnatum (van Hall) Hook. 354 didymosorum Parish ex Bedd. 560 diversilobum Pres] 452, 517 eminens Baker 449 eusorum (Thw.) Bedd. 429 extensum (BI.) Moore 548 falcilobum Hook. 374 fasciculatum (Fourn.) Baker 367 ferox (Bl.) Moore 391 flaccidum (BI.) Hook. 352 glandulosum sensu Bedd. 527 glandulosum sensu J.Sm. 512 glandulosum (BI.) J.Sm. 522 glaucostipes Bedd. 471 gracilescens (Bl.) Hook. 351 var. hirsutipes Clarke 359 griseum Baker 375 gymnopodum Baker 361 haenkeanum sensu Baker 478, 479 haenkeanum Presl 478, 502 heterocarpum_ (BI.) Moore 457 hirsutum Presl 455 hispidulum sensu Christ 466 impressum Desv. 548 indicum sensu Ridl. 485 insculptum Desv. 478 invisum (Forst. f.) Carr. 502 javanicum (Mett.) Hook. 437, 449 larutense Bedd. 437 latifolium Pres] 512, 513, 526 latipinna Hook. 558, 559 lineatum sensu Bedd. 522 lineatum sensu Hook. 528 lineatum (BI.) Pres] 512 longipes (Bl.) Moore 424 luerssenii Harr. 431 macgregorii Baker 367 microchlamys Baker 449 (Nephrodium) molle (Sw.) R.Br. 557 var. amboinense Bedd. 558 var. didymosorum Bedd. 560 var. major Bedd. 431, 556, 558 motleyanum Hook. 381 multijugum Baker 420 multilineatum Bedd. 461 multisetum Baker 349 nitidulum Presl 430, 461 obscurum (BI.) T.Moore 556 ochthodes var. tylodes Bedd. 414 oosorum Baker 500 oreopteris Fée 545 papilio Hope 556 patens sensu Hook. 555 pennigerum Hook. 461, 462 var. malayense Bedd. 485 var. multilineatum Clarke 461 perakense (Bedd.) Baker 449 philippinense Baker 460, 461 pilosiusculum Racib. 522 polycarpum (BI.) Keys. 448 polytrichum Baker 380 prolixum sensu Baker 414 var. tylodes 414 prolixum Clarke 413 propinquum R.Br. 386 pteroides sensu Bedd. 545 pubescens Brack. 457 pubirachis Baker 365 rodigasianum T.Moore 426 sagittifolium (Bl.) Moore 487 sakayense Zeiller 392, 393 savaiense Baker 404 serratum (Cav.) Pres] 478 setigerum sensu. Hook. & Baker 348 simulans Baker 367, 520 singalanense Baker 352 smithianum Pres] 456, 557 sophoroides (Thunb.) Desv. 560 sorbifolium 289 squamulosum Hook. f. 377 stipellatum (Bl.) Moore 492 tectum Bedd. 560 (Willd.) Pres! tenericaula (Hook.) Hook. 348 terminans Hook. 545 thelypteris var. squamulosum Hook. 377 truncatum sensu Racib. 429 tuberculatum Cesati 407 tylodes (Kunze) Hope 414 Index to scientific plant names 587 (Nephrodium) unitum sensu R.Br. et auct. 386 urophyllum sensu Racib. 495 urophyllum (Mett.) Keys. 533 var. pinwillii Bedd. 533 viscosum Baker 360 vulcanicum Baker 351 Nephrolepis iii, xx, 257, 258, 263, 335 acutifolia (Desv.) Christ 253 dicksonioides Christ 162, 253 schizolomae v.A.v.R. 230 Nephrolepis group Ill, ix, Xi, Xil, ROK Neurocallis Fée 277, 278 sect. Cheilolepton Moore 277 praestantissimum (Bory) Fée 289 Notholaena ii, xvii, 255 pteridiformis (Ces.) 263 Baker Odontoloma J.Sm. 198, 235 adiantoides (Bl.) Pres] 249 blumeanum (Hook.) Mett. 216 boryana (erron.) 237 boryanum (Presl) J.Sm. 237 hymenophylloides (BI.) J.Sm. 237 pulchellum J.Sm. 249 repens sensu J.Sm. 241 repens (Bory) Pres] 237 tenuifolium (BI.) J.Sm. 216 triquetra (erron.) 216 triquetrum (Baker) J.Sm. 216 Odontopteris Bernh. 39, 44 Odontosoria Fée 177, 178, 179 biflora (Kaulf.) C.Chr. 182 chinensis (L.) J.Sm. 182 var. divaricata Christ 183 var. tenuifolia (Lamk) Matsum. 183 chusana (L.) Masam. 182 var. tenuifolia (Lamk) Masam. 183 decipiens (Ces.) Christ 180 lindsayae v.A.v.R. 180 retusa (Cav.) J.Sm. 180 tenera Ridl. 252 tenuifolia (Lamk) J.Sm. 182 tsoongii Ching 182 versteegii Christ 248 Oleandra ii, iv, xx, 258, 335 Oleandropsis ti, xvi Olfersia Pres] 289, 291 aculeata Bl. 269 angulata Pres] 297 blumeana Pres] 303, 311 callifolia Pres] 307 588 (Olfersia) corcovadensis 291 gracilis Bl. 269 lineata Pres] 262 Olfersia Raddi 291 Oligocarpia 1 Oligolepideae Fée 291 Onychium ii, xvii Ophioglossaceae ii, ix, xX, Xi, XV Ophioglossales ii, vil, viii Ophioglossum ii, xv circinnatum Burm. f. 59 filiforme Roxb. 47 flexuosum L. 39, 53 furcatum Roxb. 59 japonicum Thunb. 51 pedatum Burm. f. 59 scandens L. 39, 53 Oreogrammites ii, xvii Ormoloma 177 Orthiopteris ti, xviii kingii (Bedd.) Holtt. 162 Osmunda ii, xv, 71 dichotoma Spr. 41 Osmundaceae ii, vii, X, Xili, Xv Paesia ii, xix, 38 Palmifilix alba Rumph. 131 Paragramma ii, xv Paraleptochilus ovatus Copel. 330 Parapolystichum Ching 331 Parasorus ii, xviii Parathelypteris (H.Ito) Ching 340, 342, 343, 370 FLORA MALESIANA (Parathelypteris) pectiniformis (C.Chr.) Ching 363 simozawae Ching 360 simulata (Davenp.) Nieuwl. 370 subimmersa Ching 547 viscosa (Baker) Ching 360 Pecopteris 1 Pellaea ii, xvii pteroides Prantl 61 Pelletiera 37 Peltapteris Link 255, 259 Peranema Ii, xix Phanerosorus li, xv Phegopteris sensu Tagawa 343 Phegopteris Fée 333, 336, 337, 338, 339, 342, 353 sect. Lastrella H.Ito 343 sect. Meniscium 534 acanthocarpa (Copel.) v.A.v.R. 532 acromanes (Christ) v.A.v.R. 452 aoristisora (Harr.) v.A.v.R. 538 arfakiana (Baker) v.A.v.R. 484 armata (Rosenst.) v.A.v.R. 392 auriculata J.Sm. 412 aurita (Hook.) J.Sm. 345 barbata (Fée) Mett. 391 beccariana (Ces.) v.A.v.R. $13 borneensis (Hook.) v.A.v.R. [ser. II, vol. 1° (Phegopteris) holophylla (C.Chr.) v.A.v.R. 526 hosei (Baker) v.A.v.R. 526 var. sumbensis v.A.v.R. 523 imponens (Ces.) v.A.v.R. 391 luxurians (Kunze) Mett. 387 mamberamensis v.A.v.R. 549 melanophlebia (Copel.) v.A.v.R. 514 merrillii (Christ) v.A.v.R. 529 moussetii (Rosenst.) v.A.v.R. 346 nervosa Fée 538, 539 oblanceolata (Copel.) v.A.v.R. 513 obtusifolia (Rosenst.) v.A.v.R. 503 oppositipinna v.A.v.R. 346 pentaphylla (Rosenst.) v.A.v.R. 514 perrigida v.A.v.R. 394 philippinensis Mett. 461 polypodioides Fée 353 prolifera (Retz.) Kuhn 387 pseudoarfakiana Hosok. 454 ridleyana v.A.v.R. 501 rigida (Hook. & Grev.) Mett. 484 rubicunda v.A.v.R. 536 rubida (Hook.) v.A.v.R. 532 rubrinervis Mett. 531 rutteniana v.A.v.R. 513 salicifolia (Wall. ex Hook.) Mett. 535 sect. Melanostipes Ching 354 angulariloba (Ching) Ching 360 angustifrons (Miq.) Ching 373 beddomei (Baker) Ching 370, S7i 32% var. beddomei 371, 373 var. brassii (C.Chr.) Holtt. 371 var. eugracilis Holtt. 371 cystopteroides (Eaton) Ching 370, 373 glanduligera (Kunze) Ching 370, 373 grammitoides (Christ) Holtt. 373 hirsutipes (Clarke) Ching 359 immersa (BI.) Ching 547 indochinensis (Christ) Ching 359 nipponica (Fr. & Sav.) Ching 371 noveboracensis (L.) Ching 370, 371 (Copel.) 528 canescens (BI.) Mett. 497 var. degenera (Christ) v.A.v.R. 522 ceramica v.A.v.R. 549 chamaeotaria (Christ) v.A.v.R. 524 cheilanthoides (Baker) v.A.v.R. 349 connectilis 354 cuspidata (Bl.) Mett. 536 debilis Mett. 448 decursive-pinnata (van Hall) Fée 343, 354 diversiloba (Presl) v.A.v.R. 452 engleriana (Brause) v.A.v.R. 366 ferox (Bl.) Mett. 391 firmula (Baker) v.A.v.R. 519 granulosa (Presl) v.A.v.R. 524 gymnocarpa (Copel.) v.A.v.R. 542 hexagonoptera 354 simplicifolia (J.Sm.) v.A.v.R. 490 smithii v.A.v.R. 371 spenceri (Christ) v.A.v.R. 491 stegnogramma Mett. 541 var. meniscioides v.A.v.R. 542 stenophylla (Bak.) v.A.v.R. 528 totta sensu Mett. 542 var. subcalcarata v.A.v.R. $42, 543 triphylla (Sw.) Mett. 534 urophylla Mett. 533 wollastonii v.A.v.R. 484 xiphioides (Christ) v.A.v.R. 525 Photinopteris ii, xvi Physematium Pres| 336 philippinum Pres! 539 Pityrogramma Ii, vi, XVii Plagiogyria iii, vii, xx, 340 Plagiogyria group iii, x, xiii, Xx Platycerium il, xvi Platyzoma 1, 2 1981] Pleocnemia ili, xx, 257, 334, 336, S11 cumingiana Presl 158 excellens (BI.) v.A.v.R. 462 heterophylla (Presl) v.A.v.R. 515 Plesioneuron Holtt. 335, 340, 342, 378, 389, 397 altum (Brause) Holtt. 398, 404 angiensis Holtt. 399, 406 archboldii (Copel.) Holtt. 406 attenuatum (Brack.) Holtt. 399, 407, 409 costulisorum (Copel.) Holtt. 399, 409, 410, 411 crassum (Copel.) Holtt. 399, 408 croftii Holtt. 399, 411 ctenolobum Holtt. 408 cystodioides Holtt. 398, 405 belense (Copel.) Holtt. 398, 403 bipinnatum (Copel.) Holtt. 398, 405 doctersii Holtt. 399, 410 dryas Holtt. 399, 411 dryopteroideum (Brause) Holtt. 398, 401, 402 var. buruense Holtt. 401 var. dryopteroideum 401 var. pilosum Holtt. 401 falcatipinnulum (Copel.) Holtt. 399, 406 fuchsii Holtt. 398, 400 fulgens (Brause) Holtt. 332*, 397, 398, 403 irayense (Copel.) Holtt. 495 kostermansii Holtt. 399, 406 kundipense Holtt. 399, 410 marattioides (Alston) Holtt. 396*, 398, 402 medusella Holtt. 398, 400 notabile (Brause) Holtt. 398, 402 ophiura (Copel.) Holtt. 397, 399, 409, 410 phanerophlebium (Baker) Holtt. 406 platylobum Holtt. 399, 408 pullei Holtt. 398, 399 quadriquetrum (v.A.v.R.) Holtt. 398, 406 rigidilobum Holtt. 399, 410 royenii Holtt. 398, 404 sandsii Holtt. 399, 407 savaiense 396*, 398, 404 septempedale (Alston) Holtt. 398, 402 stenura Holtt. 399, 408 (Baker) Holtt. Index to scientific plant names (Plesioneuron) stenura Holtt. 399, 408 subterminale Holtt. 398, 405, 406 tuberculatum (Ces.) Holtt. 396*, 397, 399, 404, 405, 407 varievestitum (C.Chr.) Holtt. 398, 401 var. obtusipilum Holtt. 401, 402 var. varievestitum 401 wantotense (Copel.) Holtt. 399, 410 wariense (Copel.) Holtt. 398, 403 woodlarkense (Copel.) Holtt. 398, 400 Pleuromeia 64 Pneumatopteris sensu Holtt. 413 Pneumatopteris Nakai 333, 334, 338, 340, 343, 413, 414, 437, 439, 461 angusticauda Holtt. 417, 422 auctipinna Holtt. 417, 425 basicurtata Holtt. 417, 424 boridensis Holtt. 418, 434 brooksii (Copel.) Holtt. 417, 423 callosa (BI.) Nakai 415, 416*, 419, 420 caudata (Holtt.) Holtt. 418, 433 cheesmaniae Holtt. 417, 422 christelloides Holtt. 429 costata (Brack.) Holtt. 414, 415, 417, 418, 420, 421, 432, 439 var. hispida Holtt. 421 deficiens Holtt. 418, 433 dicranogramma (v.A.v.R.) Holtt. 415, 419 eburnea Holtt. 418, 434 ecallosa (Holtt.) Holtt. 417, 425 egenolfioides Holtt. 417, 421 excisa (Holtt.) Holtt. 418, 434 finisterrae (Brause) Holtt. 418, 432 glaberrima sensu Holtt. 430 glabra (Copel.) Holtt. 415, 418 imbricata Holtt. 418, 435 incisa Holtt. 417, 427 inclusa (Copel.) Holtt. 417, 423 japenensis Holtt. 417, 426 jermyi Holtt. 417, 422, 426 kerintjiensis Holtt. 415, 418, 430 589 (Pneumatopteris) keysseriana (Rosenst.) Holtt. 416*, 418, 432, 433 laevis (Mett.) Holtt. 416*, 417, 424 laticuneata Holtt. 428 latisquamata Holtt. 415, 421, 435 lawakii Holtt. 418, 430 ligulata (Presl) Holtt. 418, 431 lithophila Holtt. 417, 422 longipes (BI.) Holtt. 417, 423, 425, 427 michaelis Holtt. 418, 430 microauriculata Holtt. 417, 425 microloncha (Christ) Holtt. 417, 423 micropaleata Holtt. 418, 428 mingendensis (Gilli) Holtt. 418, 433 nephrolepioides (C.Chr.) Holtt. 418, 431 nitidula (Presl) Holtt. 418, 430, 465 obliqua Holtt. 417, 424 papuana Holtt. 417, 426 patentipinna Holtt. 417, 422 pennigera (G.Forst.) Holtt. 415, 421 pergamacea Holtt. 417, 427 petrophila (Copel.) Holtt. 418, 435 psilophylla Holtt. 417, 427 regis (Copel.) Holtt. 418, 434 rodigasiana (T.Moore) Holtt. 417, 426 sandwicensis (Brack.) Holtt. 541 sibelana Holtt. 415, 419 sogerensis (Gepp) Holtt. 417, 428, 436 subappendiculata (Copel.) Holtt. 415, 420 sumbawensis (C.Chr.) Holtt. 418, 432 superba (Brause) Holtt. 415, 420 tobaica Holtt. 417, 427 truncata (Poir.) Holtt. 415, 416*, 418, 427, 429, 430 versteeghii Holtt. 418, 432 walkeri Holtt. 418, 435 Poecilopteris contaminans Moore 321 stenophylla Kurz ex T. & B. 325 subrepanda Presl 326 Polybotrya sensu Fée 275 Polybotrya Humb. & Bonpl. 276; 277-291 318 590 FLORA MALESIANA [ser. II, vol. 1° (Polybotrya) sect. Arthrobotrya v.A.v.R. 2555215) sect. Egenolfia Diels 255, 315 sect. Lomagramma Kuhn 277, 278 sect. Teratophyllum Christ 275, 276 appendiculata (Willd.) J.Sm. 322 var. costulata Bedd. 325 var. hamiltoniana auct. 323 var. hamiltoniana Bedd. 323 var. marginata C.Chr. 322 var. rhizophylla auct. 323 arfakensis Gepp 118, 119 articulata J.Sm. ex Fée 275, 276 duplicato-serrata Hayata 323 exaltata Brack. 323 hamiltoniana (Presl) Fée 322 lomarioides (BI.) Kuhn 279 marginata Bl. 322, 323 neglecta Fée 323 pteroides (J.Sm.) Kuhn 283 rhizophylla Pres] 323 serrulata (J.Sm.) Fée 323 vivipara Ham. ex Hook. 322, 323 Polylepideae Presl 291 Polypodiaceae 1, ix, x, Xii, Xiil, xv, 71, 256, 257, 291 tribe Aspidieae 336 tribe Leptogrammeae 336 tribe Meniscieae 336 tribe Phegopteridiae 336 tribe Polypodieae 336 Polypodiopsis ti, xvi Polypodium 256, 258, 304, 335, 336, 527 sect. Phegopteris Pres] 336, 353 acuminatum Houtt. 560 acuminatum Roxb. 556 acutum Roxb. 474 alternans Wall. 145 aoristisorum Harr. 538 appendiculatum Bedd. 412 var. squamaestipes Clarke 413 arboreum Lour. 157 arfakianum Baker 484 asperum Presl, non L. 512 auriculatum Wall. ex Hook. 412 auritum (Hook.) Lowe 345 barbatum (Fée) Hook. 391 barometz L. 165 borneense Hook. 528 canescens Bl. 497 (Polypodium) cheilanthoides Baker 349 contaminans Wall. 135 cuspidatum Roxb. 533 decursive-pinnatum van Hall 354 dentatum Forssk. 557 dichotomum Thunb. 3, 33, 34 distans sensu Racib. 347 distans Don var. minor Clarke 346 erubescens sensu Hook. 549 var. amboinense Baker 549 felinum Roxb. 563 firmulum Baker 519 fragile Baker 348 granulosum sensu Benth. 513 granulosum Presl 524 var. lobata Pres] 524 guianense Aubl. 278, 289 holophyllum Baker 526 imponens Ces. 391 intermedia (J.Sm.) Fée 323 invisum Forst. f. 502 latebrosum Wall. 115 lineare Burm. 33, 34 longifolium Roxb. 457 lunulatum Forst. 131 luxurians Kunze 387 molle Jacq. 557 nymphale G.Forst. 557 paludosum BI. 347 parasiticum L. 559 penangianum Hook. 565 pennigerum Forst. 462 phegopteris L. 353 pinwillii Baker 533 pteroides Retz. 545 rectangulare Zoll. 346 rubidum Hook. 532 rubrinerve (Mett.) Baker 531 scabrum Roxb. 391 scolopendria Burm. f. 303 simplicifolium (J.Sm.) Hook. 490 sinuatum Presl 326 sophoroides Thunb. 560 subvillosum Moore 412 tenericaule Hook. 348 tottum Thunb. 386, 387 trichodes Houlst. & Moore 348 truncatum Poir. 429 unitum L. 386, 477, 478 unitum Thunb. 560 urophyllum sensu Benth. 513 urophyllum sensu Racib. 513 urophyllum (Mett.) Hook. 533 var. uniseriale Hook. 513 urophyllum Wall. 513 Polypodium group xi Polystichopsis ii, xix Polystichum Roth i, xix, 335 aculeatum 335 auriculatum var. nervosum Christ 539 benoitianum Gaud. 500 cuspidatum Pres] 325 riedleanum Gaud. 456 torresianum Gaud. 348 truncatum Gaud. 429, 430, 436 Proferea Presl 336, 436 excellens (Bl.) Pres] 462 Pronephrium Pres] 333, 335, 336, 338, 340, 341, 343, 437, 464, 507, 555 subg. Menisciopsis Holtt. 509, 530 sect. Grypothrix Holtt. 334, 340, 341, 507, 509, 531, 534 sect. Menisciopsis 509, 531, 534 subg. Pronephrium 509 sect. Dimorphopteris (Tagawa & Iwats.) Holtt. 509, 516, 538 sect. Pronephrium 509, 517 acanthocarpum (Copel.) Holtt. 509, 531, 532 affine (BI.) Pres! 509, 518, 527 amboinense (Willd.) Holtt. 508*, 517, 519 amphitrichum Holtt. 517, §21, 525 aquatiloides (Copel.) Holtt. 509, 517, 520 articulatum (Houlst. & Moore) Holtt. 516, 517 asperum (Presl) Holtt. 506*, 507, 509, 511, 512, 513 bakeri (Harr.) Holtt. 511, 516 beccarianum (Ces.) Holtt. 491, 511, 513, 514, 516 borneense (Hook.) Holtt. 518, 528 brauseanum Holtt. 511, 514 bulusanicum (Holtt.) Holtt. 511, 515 buwaldae Holtt. 517, 554 celebicum (Baker) Holtt. 508*, 518, 523 clemensiae (Copel.) Holtt. 517, 521 var. clemensiae 521, 522 var. degenerum (Christ) Holtt. 517, 522 cuspidatum (BI.) Holtt. 533, 534, 536, 537 debile (Baker) Holtt. 517, 522 1981] Index to scientific plant names 591 (Pronephrium) (Pronephrium) Pseudophegopteris Ching 333, diminutum (Copel.) Holtt. pentaphyllum (Rosenst.) 339, 340, 342, 343, 345 $11, 515 Holtt. 507, 511, 514 aurita (Hook.) Ching 344*, euryphyllum (Rosenst.) peramelense Holtt. 517, 518 345 Holtt. 509, 511 ramosii (Christ) Holtt. 510*, cyclocarpa Holtt. 345, 346 exsculptum (Baker) Holtt. 532, 534, 536, 537 kinabaluensis Holtt. 345, 346, 518, 528 var. minahassae Holtt. 534, 347 firmulum (Baker) Holtt. 517, 537 oppositipinna (v.A.v.R.) 518, 519 var. ramosii 534, 537 Ching 346 giluwense Holtt. 517, 520 repandum (Fée) Holtt. 332*, paludosa (BI.) Ching 344%, glandulosum (BI.) Holtt. 509, 5095, S10*; 5135531, 533 345, 347 S17 522 rhombeum (Christ) Holtt. pyrrhorhachis (Kunze) Ching granulosum (Presl) Holtt. 508*, 509, 518, 525, 529 345, 347 508*, 514, 518, 524, 525 rubicundum (v.A.v.R.) Holtt. rectangularis (Zoll.) Holtt. gymnopteridifrons (Hayata) 33275105534. 536, 537 344*, 345, 346 Holtt. 509, 511, 513 heterophyllum (Presl) Holtt. 506*, 511, 515 hewittii (Copel.) Holtt. 508%, 518, 527, 529 hosei (Baker) Holtt. 389, 518, 526 kjellbergii Holtt. 517, 521 lakhimpurense (Rosenst.) Holtt. 509, 530 lastreoides Pres] 468, 507, 509 lineatum (BI.) Pres] 495, 507, 509, 511, 512 melanophlebium (Copel.) Holtt. 511, 514 menisclicarpon (BI.) Holtt. 432, 509, 513, 516, 517, 518, 526 merrillii (Christ) Holtt. 518, 527, 529 micropinnatum Holtt. 506%, 509, 511, 515 millarae Holtt. 517, 518 minahassae Holtt. 517, 523 moniliforme (Tagawa & K.I wats.) Holtt. 516, 517, 519 nitidum Holtt. 509, 510*, 531, 533 nothofageti Holtt. 466 palauense (Hosok.) Holtt. 516 palopense Holtt. 519 x parishii (Bedd.) Holtt. 534, 535 peltatum (v.A.v.R.) Holtt. 518, 529 var. aberrans Holtt. 530 var. peltatum 530 var. peninsulare Holtt. 507, 530 var. persetiferum Holtt. 530 var. tenompokense (C.Chr.) Holtt. 530 penangianum (Hook.) Holtt. 531 ssp. sulawiense Iwats. 536 tubidum (Hook.) Holtt. 531 532 rubrinerve (Mett.) Holtt. 531 salicifolium (Wall. ex Hook.) Holtt. 534, 535 samarense (Copel.) 518, 524 scopulorum Holtt. 507, 531, 532 simillimum (C.Chr.) 507, 517, 520, 525 solsonicum Holtt. 517, 523 sulawiense (Iwats.) Holtt. 534, 536 thysanoides Holtt. 518, 525 tibangense (C.Chr.) Holtt. 485 trachyphyllum Holtt. 518, 525 triphyllum (Sw.) Holtt. 335, 507, 510*, 534, 535 womersleyi Holtt. 506*, 507, 511, 514 x xiphioides (Christ) Holtt. 518, 525, 529 Prosaptia il, XVii Protolindsaya Copel. 184 brooksii Copel. 191 Pseudocyclosorus Ching 335, 340, 341, 343, 373, 374, 377, 413, 414 caudatus Holtt. 433 caudipinna Ching 413 ciliatus (Benth.) Ching 375, 413 esquirolii 413 excisus Holtt. 434 falcilobus Ching 374 ochthodes 413 Holtt. Holtt. petrophilus (Copel.) Holtt. 435 repens 413 tylodes (Kunze) Ching 412*, 413 xylodes 413 Pseudogyratae 71 sumatrana Holtt. 345, 346 tenggerensis Holtt. 345, 346 Psilotaceae ii Psilotales ii Psilotopsida ti Psilotum ti Psomiocarpa Ili, xx Pteridaceae 71 Pterideae 255 Pteridium 11, vii, xix Pteridrys ili, xx, 257 Pteris ili, iv, v, yli, xx, 253, 335 angulata Pres] 211 confluens Thunb. 377 interrupta Willd. 386, 387 polypodioides Poir. 387 stricta Poir. 211 tottum Thunb. 387 Pteris group ili, ix, xX, Xi, Xil, xiii, xx, 178 Pteropsida ii, ix, xil Pycnoloma ii, xvi Pyrrosia il, vi, XV1 Quercifilix iii, xx Ramondia Mirbel 39, 44 flexuosa (L.) Mirbel 53 Ripidium Bernh. 39, 44, 561 dichotomum Bernh. 41 Rhipidopteris Fée ex Schott 255 Saccoloma adiantoides (BI.) Pres] 249 boryanum (Presl) Pres! 237 cuneifolium Presl 180 hemipterum (Bory) Pres] 237 pinnatum (Cav.) Pres] 193 sorbifolia Christ 163 sp. 197 Salvinia iii, xxi Salviniaceae iii, viii, ix, xii, xxi Schizaea Sm. ii, vi, viii, xv, 37, 38, 39, 40, 45, 561 asperula Wakef. 39 592 (Schizaea) australis Gaud. 44 biroi Richter 41 chilensis Phil. 44 copelandica Richter 41 dichotoma (L.) Sm. v, 38%, 39, 40, 41*, 43 digitata (L.) Sw. 37, 40, 41, 42*, 44 fistulosa Labill. 37, 40, 43%, 44 var. malaccana Prantl 44 forsteri Spr. 41 hallieri Richter 44 inopinata Selling 37, 40, 42* malaccana Baker 37, 40, 43*, 44 var. malaccana 44 var. robustior C.Chr. 44, 561 papuana Brause 44 paucijuga Holtt. 44 ponapensis Hosok. 42 propinqua A.Cunn. 44 robusta Baker 561 spirophylla Troll 37, 40, 42 wagneri Selling 37, 40, 43*, 44 Schizaeaceae ii, vii, X, Xill, Xv, 37-61, 561 Schizaeales 39 Schizaeopsis 37 Schizocaena J.Sm. 73, 76, 141 alternans J.Sm. 145 arthropoda (Copel.) Copel. 143 brunonis J.Sm. ex Hook. 141, 143 capitata Copel. 142 gaudichaudii Fée 143 kinabaluensis (Copel.) Copel. 143 moluccana (R.Br.) Copel. 143 Schizolegnia ensifolia (Swartz) Alston 211 heterophylla (Dryand.) Al- ston 210 javanensis (Bl.) Alston 208 orbiculata (Lamk) Alston 206 stortii (v.A.v.R.) Alston 253 walkerae (Hook.) Alston 229 Schizolepton ii, xvii cordatum (Gaud.) Fée 253 Schizoloma Gaud. 198, 204, 229, 230 agatii Brack. 211 angustum Copel. 232 auriculatum (v.A.v.R.) v.A.v.R. 231 billardieri Gaud. 204, 211 cordatum Gaud. 253 FLORA MALESIANA (Schizoloma) coriaceum v.A.v.R. 212 davallioides (Bl.) Moore 219 divergens (Hook. & Grev.) Kuhn 233 var. auriculata v.A.v.R. 231 ensifolium (Swartz) J.Sm. 211 f. grifithianum Domin 211 f. praelongum Domin 211 f. pteroides Domin 211 f. subsimplex Domin 211 f. typicum Domin 211 var. attenuatum Domin 211 var. borneense Domin 211 var. clarkeanum Domin 211 var. heterophyllum and.) Domin 210 var. intercedens Domin 211 var. lanceolata (Labill.) R.Bonaparte 211 var. longipinnum 211 var. medium (R.Br.) Domin 208 ferulaceum (Moore) Kuhn 253 fuligineum Copel. 230 griffithianum (Hook.) Fée 211 guerinianum Gaud. 230 heterophyllum (Dryand.) J.Sm. 210 var. speluncae Copel. 211 hosei (C.Chr.) Copel. 253 (Dry- Domin induratum (Baker) C.Chr. 230 jamesonioides (Baker) Copel. japy javae Fée 211 jJavanense (BI.) Holtt. 208 lanceolatum (Labill.) Presl 211 lobatum (Poir.) Bedd. 219 medium (R.Br.) Kuhn 208 nitens (Bl.) Bedd. 222 orbiculatum (Lamk) Kuhn 206 ovata (erron.) 231 ovatum (J.Sm.) Copel. 231 pellaeiforme (Christ) C.Chr. 232 pentaphyllum (Hook.) Fée 211 recurvatum (Hook.) Moore 222 retusum (Cav.) Kuhn 180, 253 schizolomae (v.A.v.R.) v.A.v.R. 230 [ser. Il, vole (Schizoloma) stortii v.A.v.R. 197, 253 tenerum auct. 204 walkerae (Hook.) Kuhn 229 Schizostege iii, xx Scleroglossum ii, xvii Scyphularia ii, xviii Selaginella ii Selaginellaceae ii Selliguea Bory ii, xvi, 291 Senftenbergia 37, 38, 45, 561 Sitolobium stramineum Brack. 169 Sphaeropteris Bernh. 124, 562 Sphaerostephanos J.Sm. iii, xxi, 334, 337, 338, 340, 341, 343, 415, 436, 509, 517, 521,523,530) 58855411 acrostichoides (Desv.) Holtt. 441, 456 adenostegius (Copel.) Holtt. 442, 444, 447, 466 alatellus (Christ) Holtt. 440*, 441, 451 albosetosus 448, 505 alcasidii Holtt. 448, 503 alpinus Holtt. 445, 488 alticola Holtt. 444, 484 angustibasis Holtt. 443, 476 (Copel.) Holtt. angustifolius (Presl) Holtt. 444, 481 appendiculatus (BI.) Holtt. 441, 442, 452 aquatilis (Copel.) Holtt. 442, 444, 466 arbuscula (Willd.) Holtt. 439, 44] archboldii (C.Chr.) Holtt. 440*, 444, 485 arfakianus (Baker) Holtt. 439, 444, 446, 484, 486 asplenioides J.Sm. 437, 449 atasripii (Rosenst.) Holtt. 448, 504 austerus (Brause) Holtt. 447, 502 baramensis (C.Chr.) Holtt. 440*, 443, 447, 473 batacorum (Rosenst.) Holtt. 440*, 443, 475 batjanensis (Rosenst.) Holtt. 442, 467 batulantensis Holtt. 433, 473 benoitianus (Gaud.) Holtt. 447, 500 canescens (BI.) Holtt. 447, 497 carrii Holtt. 442, 463 cataractorum (Wagn. & Greth.) Holtt. 442, 468 caulescens Holtt. 443, 472 1981] Index to scientific plant names 593 (Sphaerostephanos) (Sphaerostephanos) (Sphaerostephanos) confertus (Brause) Holtt. 446, 448, 482, 491, 50S convergens Holtt. 443, 477 cyrtocaulos (v.A.v.R.) Holtt. 443, 473 daymanianus Holtt. 441, 451 debilis (Mett.) Holtt. 439, 448 dichrotrichoides (v.A.v.R.) Holtt. 444, 482 dichrotrichus (Copel.) Holtt. 441, 442, 453 dimidiolobatus Holtt. 441, 459 dimorphus (Brause) Holtt. 447, 501 diversilobus (Presl) Holtt. 441, 445, 452 echinosporus (v.A.v.R.) Holtt. 446, 495 efogensis Holtt. 444, 445, 482 ekutiensis Holtt. 444, 483 ellipticus (Rosenst.) Holtt. 443, 444, 474, 475 var. ellipticus 475 var. glabrior Holtt. 475 eminens (Baker) Holtt. 439, 449 erectus (Copel.) Holtt. 445, 490 erwinii Holtt. 448, 504 exindusiatus Holtt. 442, 463 fenixii Holtt. 495 flavoviridis Holtt. 439, 450 foliolosus Holtt. 445, 464, 488 foxworthyi Holtt. 443, 475 grandescens Holtt. 442, 465 gregarius (Copel.) Holtt. 443, 445, 447, 480 gymnorachis Holtt. 443, 479 hamiferus (v.A.v.R.) Holtt. 446, 447, 496 hastatopinnatus (Brause) Holtt. 439, 442, 444, 446, 470 hellwigensis Holtt. 447, 500 hendersonii Holtt. 446, 493 hernaezii Holtt. 441, 459 heterocarpus (BI.) Holtt. 332*, 441, 444, 457, 465, 470, 472, 474, 482 var. bordenii Holtt. 457, 465 hirsutus (Kunze ex Mett.) Holtt. 441, 443, 455, 466, 472 var. celebicus Holtt. 455, 456 var. hirsutus 455 hirtisorus (C.Chr.) Holtt. 439 hispidifolius (v.A.v.R.) Holtt. 438*, 442, 466, 469 hispiduliformis (C.Chr.) Holtt. 439, 442, 443, 447, 464, 470 var. brassii Holtt. 470 var. hispiduliformis 470 var. vinkii Holtt. 470 hoalensis Holtt. 445, 488 humilis Holtt. 442, 468 inconspicuus (Copel.) Holtt. 438*, 439, 446, 493, 494 indrapurae Holtt. 443, 475 invisus (Forst. f.) Holtt. 339, 439, 447, 479, 500, 502, 556, 565 irayensis (Copel.) Holtt. 446, 495 isomorphus Holtt. 441, 458 kalkmanii Holtt. 441, 444, 445, 454 kotoensis (Hayata) Holtt. 460 lamii Holtt. 441, 444, 454, 455 larutensis (Bedd.) 438*, 445, 447, 489 lastreoides (Presl) Holtt. 442, 468, 538 latebrosus (Kunze ex Mett.) Holtt. 443, 444, 471, 472 lithophyllus (Copel.) Holtt. 446, 496 lobangensis (C.Chr.) Holtt. 443, 479, 480 lobatus (Copel.) Holtt. 443, 458, 474, 479, 488 loherianus (Christ) Holtt. 445, 489 lucbanii Holtt. 442, 463 magnus (Copel.) Holtt. 442, 465, 472 major (Copel.) Holtt. 442, 464 makassaricus Holtt. 447, 498 melanorachis Holtt. 441, 457 menadensis Holtt. 442, 469 mengenianus Holtt. 446, 491 mjobergii Holtt. 446, 494 morotaiensis Holtt. 441, 453 moseleyi Holtt. 441, 460 multiauriculatus (Copel.) Holtt. 446, 447, 492 muluensis Holtt. 447, 499 mundus (Rosenst.) Holtt. 447, 448, 503 mutabilis (Brause) Holtt. 446, 466, 494, 565 nakaikei Holtt. 444, 482, 483 neotoppingii Holtt. 447, 500 norrisii (Rosenst.) Holtt. 437, 438*, 445, 446, 475, 485 novae-britanniae Holtt. 443, 477 ©@Chr: novoguineensis (Brause) Holtt. 439, 442, 445, 446, 469 nudisorus Holtt. 443, 476 obtusifolius (Rosenst.) Holtt. 447, 503 omatianus Holtt. 444, 483 oosorus (Baker) Holtt. 447, 500 paripinnatus (Copel.) Holtt. 443, 476 peltochlamys (C.Chr.) Holtt. 446, 495, 509, 513 penniger (Hook.) Holtt. 332*, 439, 441, 450, 452, 454, 460, 461, 478 var. excellens (BI.) Holtt. 461, 462 var. karoensis Holtt. 461, 462 var. penniger 461 perglanduliferus Holtt. 441, 454 pilosissimus Holtt. 448, 504 pilososquamatus _(v.A.v.R.) Holtt. 438*, 444, 448, 481 plurifolius (v.A.v.R.) Holtt. 446, 492 plurivenosus Holtt, 447, 496 polisianus Holtt. 445, 488 polycarpus (BI.) Copel. 437, 438*, 439, 444, 448, 565 polyotis (C.Chr.) Holtt. 442, 445, 446, 464, 488 porphyricola (Copel.) Holtt. 443, 444, 472 posthumii Holtt. 446, 493 potamios Holtt. 447, 502 productus (Kaulf.) Holtt. 423, 441, 460 pseudomegaphyllus (v.A.v.R.) Holtt. 441, 452 pterosporus (v.A.v.R.) Holtt. 445, 446, 486 var. altibus Holtt. 487 var. crenatus Holtt. 487 var. pterosporus 487 pullenii Holtt. 443, 480 punctatus Holtt. 460, 461 pycnosorus 439 reconditus Holtt. 443, 472 rigidus (Ridl.) Holtt. 444, 445, 484 roemerianus (Rosenst.) Holtt. 447, 499 rudis (Ridl.) Holtt. 447, 501 var. micans (Brause) Holtt. 501 var. rudis 501 sagittifolius (Bl.) Holtt. 445, 446, 487 (v.A.v.R.) 594 FLORA MALESIANA [ser. II, volo a (Sphaerostephanos) (Sphaerostephanos) (Stegnogramma) sakayensis (Zeiller) Holtt. vestigiatus (Copel.) Holtt. 441, subcalcarata (v.A.v.R.) Holtt. 392 453 541*, 543 santomasii Holtt. 442, 465 sarasinorum Holtt. 447, 498 semicordatus Holtt. 445, 486 semimetralis Holtt. 445, 486 sessilipinna (Copel.) Holltt. 443, 474, 479 simplicifolius (J.Sm. ex Hook.) Holtt. 445, 490, 509 smithianus (Presl) Holtt. 456 solutus Holtt. 441, 459 spenceri (Christ) Holtt. 445, 446, 481, 491, 509 squamatellus Holtt. 447, 497 stenodontus (Copel.) Holtt. 439, 450 stipellatus (Bl.) Holtt. 439, 446, 492 stresemannii Holtt. 445, 490 subalpinus (v.A.v.R.) Holtt. 443, 474 subcordatus Holtt. 442, 469 suboppositus Holtt. 447, 498 subulifolius (v.A.v.R.) Holtt. 446, 494 sudesticus Holtt. 439, 448 sumatrensis (v.A.v.R.) C.Chr. 489 talamauensis (v.A.v.R.) C.Chr. 449, 450 tandikatensis (v.A.v.R.) Holtt. 442, 463 telefominicus Holtt. 447, 499 tephrophyllus (Copel.) Holtt. 444, 480 tibangensis (C.Chr.) Holtt. 445, 485 todayensis (Christ) Holtt. 456, 471 toppingii (Copel.) C.Chr. 449, 500 trichochlamys Holtt. 442, 456, 465 trimetralis Holtt. 439, 451 uaniensis Holtt. 446, 494 uniauriculatus (Copel.) Holtt. 442, 467 unijugus Copel. 449 unitus (L.) Holtt. 339, 439, 443, 445, 446, 477, 556 var. mucronatus (Christ) Holtt. 478 var. papilliferus Holtt. 478, 479 var. unitus 478 urdanetensis (Copel.) Holtt. 442, 467, 474 veitchii Holtt. 454 warburgii (Kuhn & Christ) Holtt. 441, 462 wauensis Holtt. 448, 505 williamsii (Copel.) Holtt. 439, 450 woitapensis Holtt. 444, 483 Sphaerostephanos group 340, 341 Sphenomeris Maxon ii, xix, 177, 178, 179 alutacea 179 biflora (Kaulf.) Tagawa 177, 178, 179, 180, 182, 184, 563 chinensis (L.) Maxon 178, 563 var. chinensis 181*, 182, 184 var. divaricata (Christ) Kramer 181*, 183, 184 var. rheophila Kramer 181*, 183 var. tenuifolia (Lamk) C.Chr. 183 chusana (L.) Copel. 182 var. divaricata (Christ) Tard.-Blot 183 var. littoralis (Tagawa) H.Ito 182 var. tenuifolia (Lamk) Holtt. 183 clavata (L.) Maxon 178, 179 retusa (Cav.) Maxon 178, 179, 180, 181*, 184, 253 veitchii (Baker) C.Chr. 180 181*, 184 Sphenopsida ii Sphenopteris 1 Stegnogramma BI. ili, xxi, 257, 334, 335, 337, 340, 341, 342, 540 sect. Dictyocline Iwats. 541 sect. Haplogramma _ Iwats. 540 sect. Leptogramma_ Iwats. 540 aspidioides Bl]. 540, 541* asplenioides (J.Sm.) Fée 449 celebica (Ching) Holtt. 541, 543 dissitifolia Holtt. 541, 543 gymnocarpa (Copel.) Iwats. 541, 542, 543 ssp. celebica Iwats. 543 pozoi (Lagasca) Iwats. 542, 543 var. petiolata (Ching) Holtt. 541, 542 Stenochlaena J.Sm. ili, iv, xx, 255, 261, 265, 266, 267 sect. Eustenochlaena Diels 259, 265 sect. Lomariopsis 259, 261, 267, 276 sect. Stenochlaena Und. 267 sect. Teratophyllum Und. 261, 263, 265, 267 abrupta v.A.v.R. 262 aculeata sensu v.A.v.R. 269 aculeata (Bl.) Kunze 269 var. crassior v.A.v.R. 269, 270 arthropteroides Christ 270 cochinchinensis (Fée) Und. 262 dubia v.A.v.R. 118 gracilis Kunze 269 var. williamsii v.A.v.R. 270 hiigelii Und. 263, 276 intermedia Copel. 261 kingii Copel. 261 leptocarpa (non Fée) Und. 262 palustris (Burm. 266, 269 raciborskii C.Chr. 262 rotundifoliata R.Bonap. 270 scandens J.Sm. 266, 269 smithii (Racib.) v.A.v.R. 262 smithii Und. 262 sorbifolia sensu Bedd. 261, 262, 275 sorbifolia (L.) J.Sm. 253 ssp.) C:Chr. 270 var. spectabilis 262 subtrifoliata Copel. 261 williamsii Und. 270 Stenolepia li, xix tristis (BI.) v.A.v.R. 158 Stenoloma Fée 179 biflorum (Kaulf.) Ching 182 blumeanum (Hook.) Fée 216 chinense (L.) Bedd. 182 chusanum (L.) Ching var. littorala (Tagawa) Ito 182 clavatum (L.) Fée 179 dumosum Fée 179 littorala Tagawa 182 retusum (Cav.) Fée 180 tenuifolium (Lamk) Fée 182 veitchii (Baker) C.Chr. 184 Stenoneura Christ 291 Stenosemia ili, xx dimorpha Copel. 327 f.) Bedd. v.A.v.R. 1981] Index to scientific plant names 595 Sticherus Pres] 3, 18 sect. Eu-Sticherus C.Chr. 18 sect. Hicriopteris C.Chr. 11 amoenus Nakai 26 barbulus Nakai 26 bolanicus Copel. 24 brassii Nakai 24 caudatus Copel. 24 erectus Copel. 22 flabellatus H.St.John 25 habbemensis Copel. 22 hallieri Nakai 26 hirsutus (erron.) 26 hirtus Ching 26 var. candidus Nakai 26 var. virescens Nakai 27 hispidus Copel. 24 kajewskii Copel. 22 kajewskii St.John 561 laevigatus Pres] 20 lamianus Copel. 27 loheri Copel. 25 milnei Ching 22 monticola Nakai 24 myriapoda Nakai 20, 21 oceanicus St.John 22 peninsularis Ching 26 perpaleaceus Copel. 25 pinnatus Copel. 24 plumaeformis Nakai 22 pseudoscandens Copel. 23 pulcher Copel. 23 reflexipinnula Copel. 20 truncatus Nakai 20 venosus Copel. 20 vestitus Ching 23 Stromatopteris 1, 2 Stylites 63 gemmifera W.Rauh 64 Synaphlebium J.Sm. 179, 198 davallioides (Bl.) J.Sm, 219 nitens (Bl.) J.Sm. 222 recurvatum Hook. 217, 222 Syngramma ii, xvii, 291 angusta Copel. 380 borneensis (Hook.) J.Sm. 314 Taenitis ii, xvii blechnoides (Willd.) Swartz 253 cordata (Gaud.) Holtt. 253 trilobata Holtt. 253 Tapeinidium (Presl) C.Chr. ii, xix, 177, 178, 179, 184, 197 acuminatum Kramer 185, 194, 196* amboynense (Hook.) C.Chr. 186, 187*, 253 amplum Copel. 187 (Tapeinidium) atratum Kramer 185, 187*, 189 bartlettii Copel. 197, 198 biserratum sensu. v.A.v.R. 191 biserratum (BI.) v.A.v.R. 185, 194 brooksii (Copel.) C.Chr. 191 buntifolium Kramer 185, 186 calomelanos Kramer 185, 190 denhamii (Hook.) C.Chr. 185, 186, 187*, 253, 563 firmulum (Baker) C.Chr. 193 gracile (BI.) v.A.v.R. 185, 190 map longipinnulum (Ces.) C.Chr. 185, 194, 195*, 197 luzonicum (Hook.) Kramer 185, 186, 191, 194, 253 var. leptophyllum Kramer 191 var. luzonicum 191, 192%, 193 var. thelypteridoides Kramer 191, 193 marginale Copel. 197 melanesicum 194 moluccanum sensu Wagn. & Teratophyllum Mett. ex Kuhn LU PRX See ee ONT 258, 259, 261, 263, 266, 275, 565 sect. Polyseriatae Holtt. 257, 259, 265, 275 sect. Teratophyllum Holtt. 265, 276, 565 aculeatum (BI.) Mett. ex Kuhn 264*, 266, 267, 268*, 269, 275, 565 var. aculeatum 269 var. inermis Mett. ex Kuhn 269, 275 var. montanum Holtt. 267, 269 arthropteroides (Christ) Holtt. 267, 270, 271* articulatum (J.Sm. ex Fée) Mett. ex Kuhn 255, 259, DOE 27475255276 brightiae (F.v.M.) Holtt. 263, 276 clemensiae Holtt. 267, 270, VIB 275 gracile (BI.) Holtt. 267, 268%, 269 koordersii Holtt. 267, 271* leptocarpum (Fée) Holtt. 267, 270 ludens (Fée) Holtt. 266, 267, 269) 2727) 275 luzonicum Holtt. 267, 269, 270, 275 rotundifoliatum (R.Bonap.) Holtt. 266, 267, 269, 270, D125 wilkesianum (Brack.) Holtt. 275, 276, 289 Greth. 186 moluccanum (BI.) C.Chr. 187, 197 novoguineense Kramer 186, 188, 192* obtusatum v.A.v.R. 186, 187*, 189 oligophlebium (Baker) C.Chr. 185, 191 philippinense (Harr.) C.Chr. 191 pinnatum (Cav.) C.Chr. 162, 185, 193, 194 map, 253 prionoides Kramer 185, 194, Thayeria li, xvi Thelypteridaceae 331-560 Thelypteris Adans. 335 Thelypteris sensu Ching 386 Thelypteris Schmidel i, v, vi, 1954 stenocarpum v.A.v.R. 185, 188 stenolobum (Baker) Wagner 187 sumatranum v.A.v.R. 191 tenue sensu Kramer 186 tenue (Brack.) Copel. 197 tenuius Copel. 186 Megan (Cay, ii, VW, xox Wail 258, 266, 334, 335, 338, 339, 527 excellens C.Chr. 462 serrata Cav. 478 Tectaria group ill, x, Xi, Xi, Xili, Gh, 0. ZAI Tempskya 37 Teratophyllum sensu Holtt. 265 KX1,655)1 6681583395559. 336, 337, 338, 339, 340, 341, 342, 375, 378, 386, 539 sect. Macrothelypteris H.Ito 347 sect. Parathelypteris H.Ito 370 subg. Abacopteris Iwats. 507 subg. Cyclogramma_ Iwats. 411 subg. Cyclosoriopsis Iwats. 550, 551 subg. Dimorphopteris Iwats. 516 subg. Glaphyropteridopsis sect. Mesoneuron Iwats. 377 596 (Thelypteris) sect. Neocyclosorus Iwats. 436 subg. Glaphyropteris Christ 333 subg. Haplodictyum Iwats. 507 subg. Meniscium sect. Ampelopteris Reed 387 subg. Phegopteris sect. Lastrella Iwats. 343, 353 sect. Phegopteris Iwats. 353 subg. Pneumatopteris Iwats. 414 sect. Macrocyclosorus Iwats. 414, 436 subg. Sphaerostephanos Iwats. 436 subg. Thelypteris sect. Metathelypteris Iwats. 347, 348, 350 sect. Thelypteris Iwats. 370 acanthocarpa (Copel.) Reed 532 acrostichoides (Michx) Newl. 456 acuminata (Houtt.) Morton 560 adenophora (C.Chr.) Reed 455 adenostegia (Copel.) Reed 466 albociliata (Copel.) Reed 560 albosetosa (Copel.) Reed 505 alta (Brause) Reed 404 amboinensis (Willd.) Reed 519 angulariloba Ching 359, 360 angustata (Copel.) Reed 456 angustipes (Copel.) Reed 471 aoristisorus (Harr.) Reed 538 appendiculata (Bl.) Reed 452 aquatilis (Copel.) Reed 466 aquatiloides (Copel.) Reed 520 archboldii (C.Chr.) Reed 485 arfakiana (Baker) Reed 484 arida (D.Don) Morton 556 aspera Reed 512 asymmetrica (Fée) Reed 452 atasripii (Rosenst.) Reed 504 atrospinosa (C.Chr.) Reed 390 attenuata (O.Ktze) Morton 549 aureoviridis (Rosenst.) Reed 353 auriculata (J.Sm.) Iwats. 412 FLORA MALESIANA (Thelypteris) auriculifera (v.A.v.R.) Ching 383 aurita (Hook.) Ching 345 austera (Brause) Reed 502 austrophilippina (Copel.) Reed 479 badia (v.A.v.R.) Ching 369 bakeri (Harr.) Reed 516 baramensis (C.Chr.) Ching 473 bartlettii (Copel.) Reed 522 batacorum (Rosenst.) Reed 476 batjanensis (Rosenst.) Reed 467 beccariana (Ces.) Reed 383, 513 beddomei (Baker) Ching 371 belensis (Copel.) Reed 403 benguetensis (Copel.) Reed 560 bipinnata (Copel.) Reed 405 blumei Panigrahi 558 boholensis (Copel.) Reed 456 bordenii (Christ) Reed 457 borneensis (Hook.) Reed 528 brackenridgei_ (Mett.) Reed 408 brassii (C.Chr.) Reed 373 brooksii (Copel.) Reed 423 brunnea sensu Ching 347 brunnescens Reed 452 bunnemeijeri Reed 543 calcarata (BI.) Ching 374 callosa (Bl.) Iwats. 419 canlaonensis (Copel.) Reed 479 cataractorum (Wagn. & Greth.) Reed 469 cavitensis (Copel.) Reed 421 celebica (Baker) Reed 523 cesariana (C.Chr.) Reed 513 chamaeotaria (Christ) Reed 524 chlamydophora (C.Chr.) Ching 384 christii (C.Chr.) Reed 394 ciliata (Benth.) Ching 375 clemensiae (Copel.) Reed 521 compacta (Copel.) Reed 527 conferta (Brause) Reed 491 confluens (Thunb.) Morton BSI confusa (Copel.) Reed 539 conterminoides (C.Chr.) Reed 367 contigua (Rosenst.) Reed 557 coriacea (Brause) Ching 368 costata (Brack.) Reed 421 costulisora (Copel.) Reed 409 [ser. II, vol. (Thelypteris) crassa (Copel.) Reed 408 crassifolia (Bl.) Ching 377, 384 var. motleyana Ching 381 cuspidata (BI.) Iwats. 536 cyrtocaulos (v.A.v.R.) Reed 473 dayi (Bedd.) Nayar & Kaur 353 debilis (Mett.) Reed 448 decursive-pinnata (van Hall) Ching 343, 354 degenera (Christ) Reed 522 deltiptera (Copel.) Reed 420 dentata (Forssk.) E.St.John 557 diaphana (Brause) Ching 362 dichrotricha (Copel.) Reed 453 dichrotrichoides (v.A.v.R.) Reed 482 dicranogramma (v.A.v.R.) Reed 419 dimorpha (Brause) Reed 501 distincta (Copel.) Reed 548 divergens (Rosenst.) Reed 384 diversiloba (Presl) Reed 452 diversivenosa (v.A.v.R.) Ching 353 dryopteroidea (Brause) Reed 401 duplosetosus (Copel.) Reed 468 dura (Copel.) Reed 368 ecallosa (Holtt.) Reed 425 echinata (Mett.) Reed 379 echinospora (v.A.v.R.) Reed 495 edanyoi (Copel.) Reed 525 elliptica (Rosenst.) Reed 474 elmerorum (Copel.) Reed 485 engleriana (Brause) Reed 366 ensipinna (Brause) Ching 404 erectus (Copel.) Reed 490 erubescens 550 eugracilis (Copel.) Reed 371 euryphylla (Rosenst.) Reed S11 exsculpta (Baker) Iwats. 528 extensa (BI.) Morton 548 falcatipinnula (Copel.) Reed 406 falcatula (Christ) Reed 557 farinosa (Brause) Reed 503 fasciculata (Fourn.) Ching 367 ferox (Bl.) Tagawa & Iwats. 391 finisterrae (Brause) Reed 432 1981] Index to scientific plant names 597 (Thelypteris) (Thelypteris) (Thelypteris) flaccida (Bl.) Ching 352 flavo-virens (Rosenst.) Reed 353 forsteri Morton 502 foxii (Christ) Reed 431 fulgens (Brause) Ching 403 glanduligera (Kunze) Ching 370, 373 glaucescens (Brause) Ching 469 gracilescens (Bl.) Ching 351 grammitoides (Christ) Ching 373 granulosa (Presl) Reed 524 gregaria (Copel.) Reed 480 grisea (Baker) Ching 375 gymnocarpa (Copel.) Morton 542 haenkeana (Presl) Reed 478 halconensis (Copel.) Reed 471 hallieri (Christ) Reed 379 hamifera (v.A.v.R.) Reed 496 harveyi (Mett.) Proctor ex Iwats. 555 hastatopinnata (Brause) Reed 470 herbacea Holtt. 360 heterocarpa (BI.) Morton 457 var. glaucostipes (Bedd.) Reed 471 hewittii (Copel.) Reed 527 hirsutipes (Clarke) Ching 359, 370 hirtopilosa (Rosenst.) Reed 557 hispidifolia (v.A.v.R.) Reed 466 hispidula (Decne) Reed 557 hispiduliformis (C.Chr.) Reed 470 horridipes (v.A.v.R.) Reed 394 hosei (Baker) Iwats. 526 hunsteiniana (Brause) Reed 403 immersa (BI.) Ching 158, 547 imponens (Ces.) Reed 391 impressa (Desv.) Reed 548 inclusa (Copel.) Reed 423 inconspicua (Copel.) Ching 493 indochinensis (Christ) Ching 359 interrupta (Willd.) Iwats. 386 irayensis (Copel.) Reed 495 iridescens (v.A.v.R.) Reed 522 jacobsii (Holtt.) Reed 520 japonica 370 keysseriana (Rosenst.) Reed 432 khamptorum Holtt. 374 klossii (Ridl.) Ching 361 kotoensis (Hayata) Iwats. 460 laevis (Mett.) Reed 424 lanceola (Christ) Reed 529 larutensis (Bedd.) Reed 489 latebrosa (Mett.) Reed 471 latifolia (Presl) Reed 526 latipinna (Benth.) Iwats. 558 lauterbachii (Brause) Reed 367 lepidopoda (C.Chr. & Tard.) Reed 429 leucadenia (Copel.) Reed 460 leucolepis (Presl) Ching 349 ligulata (Presl) Ching 431 lineata (BI.) Iwats. 512 lithophylla (Copel.) Reed 496 lobangensis (C.Chr.) Reed 479 lobata (Copel.) Reed 474 loheriana (Christ) Reed 489 longipes (BI.) Reed 424 magnus (Copel.) Reed 465 malayensis (C.Chr.) Reed 522 matutumensis (Copel.) Reed 467 megalocarpa (v.A.v.R.) Ching 359 megaphylla (Mett.) Iwats. 461 megaphylloides Reed 481 melanophlebia (Copel.) Reed $14 menisciicarpa_ (BI.) Iwats. 526 merrillii (Christ) Reed 529 micans (Brause) Reed 501 microcarpa (v.A.v.R.) Ching 371 microloncha (Christ) Reed 423 mindanaensis (Christ) Reed 557 mixta (Rosenst.) Reed 457 moniliformis (Tagawa & Iwats.) Iwats. 519 morobensis (Copel.) Reed 501 motleyana (Hook.) Holtt. 381 multiauriculata (Copel.) Reed 492 multilineata (Wall. ex Hook.) Ching var. malayensis 512 multiseta (Baker) Ching 349 multisora (C.Chr.) Reed 359 (Rosenst.) munda (Rosenst.) Ching 503 muricata (Brause) Reed 391 mutabilis (Brause) Reed 494 nephrolepioides (C.Chr.) Reed 43] nervosa (Klotzsch) Tryon 539 nitidula (Presl) Reed 430 notabilis (Brause) Ching 402 novae-hiberniae Holtt. 555 novoguineensis (Brause) Reed 469 nymphalis (G.Forst.) Reed Spy// oblonga (Brause) Reed 503 obtusata (v.A.v.R.) Ching 362 obtusifolia (Rosenst.) Reed 503 oligodictua (Baker) Reed 380 oligolepia (v.A.v.R.) Ching 368 ophiura (Copel.) Reed 409 oppoOsitipinna (v.A.v.R.) Ching 346 opulenta (Kaulf.) Fosb. 548 orbicularis (C.Chr.) Reed 548 palawanensis Reed 524 paleata (Copel.) Holtt. 380 pallescens (Brause) Ching 383 palustris Schott 375 var. squamigera Weath. 377 papilio (Hope) Iwats. 556 paraphysata Reed 481 parasitica (L.) Iwats. 559 paripinnata (Copel.) Reed 477 parishii (Bedd.) Panigrahi 535 pectiniformis (C.Chr.) Ching 355, 363 peltata (v.A.v.R.) Reed 529 peltochlamys (C.Chr.) Reed 495 pentaphylla (Rosenst.) Reed 514 perakensis (Bedd.) Reed 449 perglandulifera (v.A.v.R.) Reed 454 perpilifera (v.A.v.R.) Reed 456 perpubescens (Alston) Reed 554 petrophila (Copel.) Reed 435 philippina (Presl) Ching 539 pilosiuscula (Racib.) Reed S22 pilososquamata Reed 481 (v.A.v.R.) 598 (Thelypteris) pinnata (Copel.) Ching 375 platyptera (Copel.) Reed 369 plumosa (C.Chr.) Reed 360 plurifolia (v.A.v.R.) Reed 493 polycarpa (BI.) Iwats. 449 polyotis (C.Chr.) Reed 464 polyptera (Copel.) Reed 471 porphyricola (Copel.) Ching 472 procurrens (Mett.) Reed 560 producta (Kaulf.) Reed 460 prolifera (Retz.) Reed 387 protecta (Copel.) Reed 485 pseudoamboinensis (Rosenst.) Panigrahi 558 pseudohirsuta (Rosenst.) Reed 460 pseudoreptans (C.Chr.) Reed 522 pseudostenobasis (Copel.) Reed 549 pterospora (v.A.v.R.) Reed 487 pubirachis Reed 365 pustulosa (Copel.) Reed 513 quadrangularis (Fée) Schelpe 557 quadriaurita 404 quadriquetra (v.A.v.R.) Ching 406 rectangularis (Zoll.) Nayar & Kaur 346 reederi (Copel.) Reed 456 regis (Copel.) Reed 434 repandula_ (v.A.v.R.) Reed Si, rhombea (Christ) Reed 529 rigida (Ridl.) Reed 484 rigidifolia (v.A.v.R.) Ching 369 riparia (Copel.) Reed 456 robinsonii (Ridl.) Ching 361 rodigasiana (T.Moore) Reed (Christ) Reed 426 roemeriana (Rosenst.) Reed 499 rubicunda (v.A.v.R.) Iwats. 536 rubida (Hook.) Iwats. 532 rubrinervis (Mett.) Iwats. 531 sagittifolia (Bl.) Reed 487 Sagittifolioides (Copel.) Reed 471 sakayensis (Zeiller) Reed 392 salacifolia (Wall. ex Hook.) Reed 535 samarensis 524 savaiensis (Baker) Reed 404 (Copel.) Reed FLORA MALESIANA (Thelypteris) sessilipinna Reed 479 setigera Ching 349 setulosa Reed 481 sevillana Reed 418 simillima (C.Chr.) Iwats. 520 simozawae Tagawa 360, 370 simplicifolia (J.Sm.) Reed 490 simulans Ching 413 singalanensis sensu Holtt. 353 singalanensis (Baker) Ching 352 sogerensis (Gepp) Reed 428 spenceri (Christ) Reed 491 squamigera Ching 377 squamipes (Copel.) Reed 364 squamulosa (erron.) 377 stegnogramma (BI.) Reed 542 stenobasis (C.Chr.) Ching 549 stenodonta Reed 450 stenophylia (Baker) Reed 528 stereophylla (v.A.v.R.) Ching 366 stipellata (Bl.) Iwats. 492 strigosissima (Copel.) Reed 470 subalpina Reed 474 subappendiculata Reed 420 subattenuata (Rosenst.) Reed 547 subdimorpha (Copel.) Reed 384 subfalcinella (v.A.v.R.) Reed 485 subglandulifera Ching 363 subimmersa Ching 547 subnigra (Brause) Ching 369 subpubescens (BI.) Iwats. 558 subulifolia Ching 494 subvillosa Ching 412 sumatrana (v.A.v.R.) Tagawa & Iwats. 558 superba (Brause) Reed 420 supraspinigera (Rosenst.) Reed 395 suprastrigosa (Rosenst.) Reed 457, 458 tephrophylla (Copel.) Reed 481 terminans (Hook.) Tagawa & Iwats. 545 terrestris (Copel.) Reed 491 teuscheri (v.A.v.R.) Reed 381 todayensis (Christ) Reed 471 toppingii (Copel.) Iwats. 500 torresiana (Gaud.) Alston 348 triphylla (Sw.) Iwats. 534 (Copel.) [ser. Il, ‘volar (Thelypteris) var. parishii (Bedd.) Iwats. 535 truncata (Poir.) Iwats. 429 tuberculata (Ces.) Ching 407 tylodes (Kunze) Ching 414 ugoensis Reed 460 uliginosa (Kunze) Ching 348 uniauriculata (Copel.) Reed 467 unidentata (Bedd.) Holtt. 359 unita (L.) Morton 478 uraiensis (Rosenst.) Ching 352 urdanetensis (Copel.) Reed 467 urophylla (Mett.) Iwats. 533 var. nitida (Holtt.) Iwats. 533 varievestita 401 verrucosa (Presl) Ching 547 verruculosa (v.A.v.R.) Reed 512 vestigiata (Copel.) Reed 453 vinosicarpa (v.A.v.R.) Ching 381 viridis (Copel.) Reed 418 viscosa sensu Holtt. 361 viscosa (Baker) Ching 355, 360 vulcanica (Baker) Reed 351 wagneri Fosb. & Sachet 545 wantotensis (Copel.) Reed 410 warburgii (Kuhn & Christ) B.C.Stone 462 wariensis (Copel.) Reed 403 weberi (Copel.) Reed 460 williamsii Ching 450 xiphioides (Christ) Reed 525 zamboangana Reed 394 Thelypteris group ili, ix, xi, xii, Xili, XiV, XX1, 257, 340 group 2 Ching 370 group 4 Ching 343 group 10 Ching 437 Thylacopteris tl, xvi Thyrsopteridaceae 562 Thyrsopteridineae 562 Thyrsopteris 65, 69, 70, 72, 167, 562 Thysanobotrya v.A.v.R. 73, 115 arfakensis (Gepp) v.A.v.R. 118 Thysanosoria Gepp iii, xx, 255, 256, 258, 263 dimorphophylla Gepp 263 pteridiformis (Cesati) C.Chr. 263* Tmesipteridaceae ti (C.Chr.) Reed 1981] Tmesipteris ii Todea barbara Moore 61 Toppingia Deg. 343 Trichomanes ii, vii, Xv sect. Crepidomanes vi chinense L. 182 malayanum Roxb. ex Griff. 183 Trichopteris 562 falcata Llanos 146 Trigonospora Holtt. 331, 333, 338, 339, 340, 342, 373 calcarata (BI.) Holtt. 372*, 374 ciliata (Benth.) Holtt. 372*, 374, 375, 377 koordersii (Christ) Holtt. 374, 375 zeylanica Ching 374 Trigonospora group 340 Trogostolon ii, xviii Index to scientific plant names Ugena Cav. 39, 44 dichotoma Cav. 59 microphylla Cav. 47 polymorpha Cav. 562 semihastata Cav. 54 Vaginularia ili, XXi Vallifilix Thouars 39, 45 Vittaria ili, vi, vill, Xxi divergens (Hook. & Grev.) Roxb. ex Griff. 233 interrupta Roxb. ex Griff. 254, 563 lunulata Roxb. ex Griff. 254, 563 parasitica Roxb. ex Griff. 244 Vittaria group Ill, ix, Xili, Xxi Wagneriopteris Love & Love 370, 371 aoe (Wagneriopteris) angulariloba (Ching) Love & Love 360 ogasawarensis 370 Wibelia auct. 184 Wibelia amboynensis Kuhn 187 bipinnata Fée 191 denhami (Hook.) Kuhn 186 gracilis (BI.) Christ 190 javae Fée 193 oligophlebia (Baker) Christ 191 pinnata (Cav.) Bernh. ex Fée 193 Woodwardia il, Xvili (Hook.) Xiphopteris ii, xvii Xyropteris Kramer 177, 178, 179, 197 stortil (v.A.v.R.) Kramer 197, 253 _ : >; een 2. | (wee fwarg <4 p den Library WLC TEATATY Se a : i ‘ Stl et a ee a. oe we oe a re ee ee ts Ye eatin ae np ae Wot i ened MM ean Las - oN OL SR as eh Ry te Ne ind ne ee AVE Oe ~* men OS eRe ee SS Sn be aS Peat ae . X . NID heen “Ye ies EES peed eS pea as Ny Gn Ge OL BS re oe Seana ae SNe. ~ . . pO hs - eee P ma \ SS : ° oe ‘ pee Pinon hs ; . x me ~s a 0 ‘ > See ON aed ae Zs sg Te nee ean tS: ee ee