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Full text of "Flora Malesiana"

UBRARY 

THE NEW YORK BOTANICAL GAROEW 
Bf?QH)^ NIW YORK 10499 



ijhji 



TAXONOMICAL REVISIONS 



REPUBLIK INDONESIA 
REPUBLIC OF INDONESIA 

LEMBAGA ILMU PENGETAHUAN INDONESIA (L.I.P.I.) 
INDONESIAN INSTITUTE OF SCIENCES 

FLORA MALESIANA 

BEING 

AN ILLUSTRATED SYSTEMATIC ACCOUNT OF THE MALESIAN FLORA / 

INCLUDING KEYS FOR DETERMINATION I DIAGNOSTIC DESCRIPTIONS I 

REFERENCES TO THE LITERATURE I S YN ON YM Y I AND DISTRIBUTION I 

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 ANINTERNATIONALCO-OPERATIVEBASIS 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 ^^VIlT^/^ • VOLUME 1 

PTERIDOPHYTA 



GENERAL EDITORS: 

Dr C. G. G. J. VAN STEENIS 

DIRECTOR OF THE FOUNDATION 'FLORA MALESIANA' 

&DrR. E. HOLTTUM 

PUBLISHED BY 

MARTINUS NIJHOFF / DR W. JUNK PUBLISHERS 
THE HAGUE / BOSTON / LONDON 

1959-1982 




COPYRIGHT 1982 ,^ \ 

All rights reserved, including the right to reproduce 

this book or parts thereof in any form 

ISBN 9024726530 



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 



Title-page (3) 

Contents (5) 

Dedication by R. E. Holttum (7) 



GENERAL CHAPTERS 

Introductory note by R. E. Holttum i 

List of Malesian Pteridophytes by K.'E..'Ho\i\\im ii 

The morphology of ferns by R. E. Holttum iii 

General key No. 1 to Pteropsida by R. E. Holttum ix 

General key No. 2 to Pteropsida by R. E. Holttum xii 

Keys to the genera of Pteropsida by R. E. Holttum xv 

Bibliography by R. E. Holttum xxii 



TAXONOMICAL REVISIONS 

in alphabetical sequence 

Cyatheaceae by R. E. Holttum 65 

Gleicheniaceae by R. E. Holttum 1 

Isoetaceae by A. H. G. Alstonf 62 

Lindsaea group by K. U. Kramer 177 

Lomariopsis group by R. E. Holttum & E. Hennipman 255 

Schizaeaceae by R. E. Holttum 37 

Thelypteridaceae by R. E. Holttum 331 



ADDENDA 
Addenda, corrigenda et emendanda by R. E. Holttum & K. U. Kramer 561, 565 



INDEX 

Irulex to scientific plant narnes hy Mrs. M. J. vdinS,iQQms-K.Tn%QVD.z.n 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 Diels) for Engler & Prantl's Pffanzenfamilien 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 Diels. 
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, etc., 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 I 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, Betty Molesworth 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.) Mirbel {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.) 
Copel., 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. 

Hooker 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 Presl (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. ScHOTT, at the imperial palace of Schonbrunn, 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. Fitch, 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 P*resl 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. Fitch 
(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 diff'ers from Blechnum only in 
the absence of indusia. He placed Brainea between Gymnogramme (which included the diverse 
genera Selligiiea 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 Mett.), 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 diff"erence in venation between Stenochlaena and Lomariop- 
sis, not noticed by Fee. But his scheme only diff'ers 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. Beddome 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 (1 875) 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 etc., but from observation of his plants he did learn much that 



(10) 



Dedication 

Hooker never understood. I will refer later to some of his insights in a discussion of the work of 
DiELS. Smith and Mettenius, 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 Polypodiiiin and transferred them to one including 
Aspidiiim. 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 son 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 leiizeana (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 Presl) 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. leuzeana. Hooker assumed that some specimens had 
lost the indusia which they originally possessed. Fee had specimens of the same collections from 
Cuming, and speculated (1852, p. 31 1) 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 Copeland 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. leuzeana), 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. 21 A 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 1 864. 
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 Hernionitis; 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 diflferent 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. Smith 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 Holttum (1947) in Grammitidaceae are separate from Polypodiaceae. 
Mettenius also found that the spores of the two groups differ (see below on Diels 1899). 

Hermann Christ (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 Hooker 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 Diels. 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 (scns.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 Gleichenia 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 5. 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. Diels 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 Holttum 1947 and 1955. Diels 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. 



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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 (1863 A) and on Hymeiwphyllaceae (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 
GoEBEL 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 Milde 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 Christ, 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 PhyUitis in a group separate from Asplenium, not knowing that natural hybrids between 
the two existed. He accepted Christ'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 diff"erent 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 Leptochihis) 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 Mettenius 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 diff"erences 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 diff"er 
from Ctenitis 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 

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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 \olumej 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 
\'erdoorn"s Manual of Pteridology. This contains many ne\N 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 Blume 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 oi 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 irihQ Phegoplerideae 
widely separated from Aspidieae; he revived Pleocnemia Presl 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 obser\ ations on spores ; many of his new species are still recognized 
in the present work. 

In Malaya H. N. Ridley made large new collections in the years 1888-191 1, 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 

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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 1 960. He was the first person 
to understand that Hooker'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 suff"ered 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.) Holttum and Pneumatopteris truncata 
(PoiR.) Holttum (as named in the present work). Yet in Hytnenophyllaceae 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. Posthumus, 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, 1 959) 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 
PiCHi Sermolli (1977) is premature, though it is much nearer to being natural than Copeland's 
of 1947. 



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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 diff'erent 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 Pteris, 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. Holttom 



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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. 

Bov^'ER, 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. 

Christ, H. 1897. Die Farnkrauter der Erde. G. Fischer, Jena. 

1899. Monographic 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. I.e. VIII, 6 (1): 1-132. 

— 1932. The Pteridophyta of Madagascar. Dansk Bot. Ark. 7: 1-253, pi. 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. HOLTTUM. 1934. The ferns of Mt Kinabalu. Card. Bull. Str. Settl. 7: 191-324, 



pi. 51-62. 

CoPELAND, E. B. 1933. Trichomanes. Philip. J. Sci. 51 : 119-280, t. 1-61. 

1937. Hymenophyllum. I.e. 64: 1-188, t. 1-189. 

1938. Genera Hymenophyllacearum. I.e. 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. 

Fee, a. L. A. 1852. Memoires sur la famille des Fougeres, V. Genera Filicum. Paris & Stras- 
bourg. 

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Dedication 

HoLTTUM, R, E. 1932. On Stenochlaena, Lomariopsis and Teratophyllum in the Malayan Region. 

Card. 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 Presl. 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. 
Klekowski 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. 
KuHN, M. 1869. Filices. Ann. Mus. Bot. Lugd.-Bat. 4: 276-300. 
KuNZE, G. 1840-1851. Die Farrnkrauter in kolorirten Abbildungen (2 volumes). E. Fleischer, 

Leipzig. 
Levis, J. D. 1977. Evolutionary patterns and processes in ferns. Adv. Bot. Res. 4: 229-415. 
LuGARDON, B. 1971. Contribution a la connaissance de la morphogenese et de la structure des 

parois sporales chez les filicinees isosporees. These, Toulouse, 257 pp., 51 tab. 
Manton, 1. 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?). 

1863 A. tJber den Bau von Angiopteris. Abhandl. math. phys. CI. 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; I.e. 1 : 222-241. 

1864B. iJber die Hymenophyllaceae. Abhandl. math. phys. CI. K. Sachs. Gew. Wiss. 7: 

403-504, t. i-v. 

1866. Filices; praesertim Indicae et Japonicae. Ann. Mus. Bot. Lugd.-Bat. 2: 219. 

MiLDE, J. 1866. Das Genus Athyrium. Bot. Zeit. 24: 373-376. 

1870. iJber Athyrium, Asplenium und Verwandte. Bot. Zeit. 28: 329-337, 345-354, 370- 

371. 
Moore, 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. Mai. Br. R. As. Soc. 4: 1-121. 
ScHKUHR, C. 1804-1809. Vier und zwanzigste Klasse der Linneischen Pflanzensystems oder 

Kryptogamische Gewachse, vol. 1-3. Wittenberg. 

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Flora Malesiana 



ScHOTT, H. 1834. Genera Filicum. Wallishauser, Vienna. 

Smith, James E. 1793. Tentamen botanicum de Filicum generibus dorsiferarum. Mem. Acad. 

Turin 5: 401^22. 
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. 



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1. INTRODUCTORY 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 pubhshed and dealing with Malaysian ferns are listed 
after the keys to genera. 

R. E. HOLTTUM 



2. LIST OF MALAYSIAN PTERIDOPHYTES 



PSILOTOPSIDA 

PSILOTALES 

Psilotaceae 

Psilotum 
Tmesipteridaceae 

Tmesipteris 

LYCOPSIDA 

Lycopodiales 
Lycopodiaceae 

Lycopodium 
Selaginellaceae 

Selaginella 
Isoetaceae 

Isoetes 

SPHENOPSIDA 

Equisetales 
Equisetaceae 
Equisetum 

PTEROPSIDA 

Ophioglossales 
Oph ioglossaceae 
Botrychium 
Helminthostachys 
Ophioglossum 

Marattiales 
Marattiaceae 
Angiopteris 
Christensenia 
Macroglossum 
Marattia 

FiLlCALES 

Osmundaceae 

Leptopteris 

Osmunda 
Schizaeaceae 

Lygodium 

Schizaea 
Gleicheniaceae 

Dicranopteris 

Gleichenia 
Hymenophyllaceae 

Hymenophyllum 

Trichomanes 
Matoniaceae 

Matonia 

Phanerosorus 
Cyatheaceae 

Cyathea 
Polypodiaceae (s.str.) 

Aglaomorpha 

Arthromeris 

Belvisia 

Cheiropleuria 

Christiopteris 



Colysis 
Crypsinus 
Dendroconche 
Dendroglossa 
Diblemma 
Dipteris 
Drymoglossum 
Drynaria 
Goniophlebium 
Grammatopteridium 
Holcosorus 
Holostachyum 
Lecanopteris 
Lemmaphyllum 
Lepisorus 
Leptochilus 
Loxogramme 
Merinthosorus 
Microsorium 
Myrmecophila 
Neocheiropteris 
Oleandropsis 
Paragramma 
Photinopteris 
Platycerium 
Polypodiopsis 
Pycnoloma 
Pyrrosia 
Selliguea 
Thayeria 
Thylacopteris 
Crammitidaceae 
Acrosorus 
Calymmodon 
Ctenopteris 
Grammitis 
Nematopteris 
Oreogrammitis 
Prosaptia 
Scleroglossum 
Xiphopteris 

Remaining Genera of Ferns 
(in Groups, alphabetically) 
Adiantum Group 

Adiantum 

Anogramma 

Ceratopteris 

Cerosora 

Cheilanthes 

Coniogramme 

Doryopteris 

Hemionitis 

Nothoiaena 

Onychium 

Pellaea 

Pityrogramma 

Syngramma 

Schizolepton 

Taenitis 
Asplenium Group 

Asplenium 

Diplora 



Loxoscaphe 
Athyrium Group 
Anisocampium 
Athyrium 
Callipteris 
Cornopteris 
Cystopteris 
Diplaziopsis 
Diplazium 
Dryoathyrium 

Blechnum Group 
Blechnum 
Brainea 
Doodia 
Woodwardia 

Davallia Group 

Araiostegia 

Davallia 

Davallodes 

Humata 

Leucostegia 

Parasorus 

Scyphularia 

Trogostolon 
Dennstaedtia Group 

Dennstaedtia 

Histiopteris 

Hypolepis 

Microlepia 

Monachosorum 

Orthiopteris 

Paesia 

Pteridium 

Dicksonia Group 

Cibotium 

Culcita 

Cystodium 

Dicksonia 
Dryopteris Group 

Acrophorus 

Currania 

Diacalpe 

Didymochlaena 

Dryopteris 

Gymnocarpium 

Peranema 

Polystichum 

Polystichopsis 

Stenolepia 
Lindsaea Group 

Isoloma 

Lindsaea 

Sphenomeris 

Tapeinidium 

Lomariopsis Group 
Arthrobotrya 
Bolbitis 
Egenolfia 
Elaphoglossum 
Lomagramma 
Lomariopsis 



Morphology of ferns 



Teratophyllum 

Thysanosoria 
Nephrolepis Group 

Arthropteris 

Nephrolepis 

Oleandra 
Plagiogyria Group 

Plagiogyria 
Pteris Group 

Acrostichum 

Hemipteris 

Lepidocaulon 

Pteris 

Schizostege 

Stenochlaena 
Tectaria Group 

Arcypteris 



Ctenitis 

Cyclopeltis 

Dryopolystichum 

Hemigramma 

Heterogonium 

Hypodematium 

Lastreopsis 

Luerssenia 

Pleocnemia 

Psomiocarpa 

Pteridrys 

Quercifilix 

Stenosemia 

Tectaria 

Thelypteris Group 
Ampelopteris 
Cyclosorus 



Dictyocline 
Sphaerostephanos 
Stegnogramma 
Thelypteris 
Vittaria Group 
Antrophyum 
Monogramma 
Vaginularia 
Vittaria 

Heterosporous Ferns 
Marsileaceae 

Marsilea 
Salviniaceae 

Azolla 

Salvinia 



3. THE MORPHOLOGY OF FERNS 

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 thegametophyte. 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. 



Ill 



Flora Malesiana [ser. II, vol. 1^ 



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. Apellate 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 leaf-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 oiPteris); 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 lamina. The lamina 
is usually divided into parts called leaflets; 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 J. 

(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 

IV 



Dec. 1959] 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 lobed 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 tripinnate, and the 
leaflets are of the third order; fronds may also be quadripinnate. Pinnae or rachis-branches are 
usually alternate 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 dichotorna. Dipt er is 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 



Flora Malesiana [ser. II, vol. P 



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 casta 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 areoles, 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 diff"erent from that of fronds of mature plants; such young plants may off"er useful 
diagnostic characters. Somie 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 



Dec. 1959] 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 (Pteridiiim) , 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 diff"er 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 diff'erence between the two. 

(c) The Acrostichoid 5/are.— 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 diff"erent 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 oblique 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- 

vn 



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 always 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 Phaseohis 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 tapetiim) 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 Salviniaceae 

2. Plants rooted in earth or on rocks. Leaves larger, more divided. 

3. Leaves 4-partite. Sporocarps attached to stipes 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 lowtr 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. Epiphytes. 
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 Vittaria Group 

7. Fronds branched, or if simple with a more complex venation. 

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 Polypodiaceae 

14. Veins not or slightly anastomosing. 

15. Frond and stipe ± hairy; spores trilete Grammitidaceae 

15. Frond and stipe not hairy; spores monolete Polypodiaceae 

13. Fronds pinnate, pinnae articulate to rachis. 

16. Pinnae entire Polypodiaceae 

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 

18. Veins free, fronds more or less hairy 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 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 Nephrolepis Group 

24. Pinnae (if any) not articulate to rachis. 

25. Rhizome protostelic Lindsaea Group 

25. Rhizome with more complex vascular system Davallia Group 

23. Sori otherwise. 
26. Fronds articulate to rhizome. 
27. Sori at ends of veins, near edge of lamina. 

28. Pinnae (if present) not jointed to rachis Davallia Group 

28. Pinnae jointed to rachis Nephrolepis Group 

27. Sori close to costa of the simple frond Nephrolepis Group 

26. Fronds not articulate to rhizome Nephrolepis Group 

6. Terrestrial plants, or climbers starting from the ground, 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) Pteris Group 

30. Rhizome dorsiventral; veins either free or much anastomosing . Lomariopsis Group 

29. Terrestrial or rock plants. 

IX 



Flora Malesiana [ser. II, vol. 1^ 



31. Caudex massive, erect; stipes succulent, with stipule- like outgrowths at their bases; bases of 
pinnae swollen 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 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 pinnae) Plagiogyria Group 

37. Aerophores lacking 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 Gleicheniaceae 

43. Fronds otherwise Adiantum Group 

41. Sori at ends of veins or on special appenoages. 
44. Sporangia on special appendages which are at ends of veins of leaflets or attached 

near apex of frond or of its branches 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. 

50. Fertile frond simply pinnate Blechnum Group 

50. Fertile frond bipinnate Tectaria Group 

49. Veins much anastomosing in sterile fronds Tectaria Group 

45. Sporangia not acrostichoid. 
51. Sorus along edge of lamina, continuous or nearly so. 
52. 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 

towards edge of lamina 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. 



Dec. 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 
hairs Dryopteris Group 

59. Rachis somewhat grooved, groove not open to admit groove of pinna; scales with 
superficial hairs Thelypteris Group 

58. Sorus asymmetric, or on one side of 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. Slender unicellular hairs lacking Adiantum Group 

62. Sori not spreading along all veins. 

64. Several vascular bundles in stipe Tectaria Group 

64. Two vascular bundles at base of stipe, uniting upwards. 

65. Waxy powder on lower surface of lamina Adiantum Group 

65. No waxy powder present 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 reflexed edge of the lamina. 

67. Sori each in the base of a cup Dennstaedtia Group 

67. Sori protected by indusia or by edge of lamina. 
68. Sori protected by indusia opening outwards. 
69. Pmnae articulate to rachis Nephrolepis Group 

69. Pinnae not articulate Lindsaea Group 

68. Sori protected by reflexed edge of lamina. 

70. An inner indusium also present; rachis grooved, 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 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 Polypodium Group 

73. Fronds always having some dormant apices Gleicheniaceae 

72. Sori indusiate. 

74. Fronds simple Nephrolepis Group 

74. Fronds pinnately branched 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 Cyatheaceae 

76. Not tree-ferns; annulus vertical, interrupted; \ascular bundles in a simple ring 
(except Pleocnemia). 
11. 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-r.hape .... 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 



XI 



5. GENERAL KEY No 2 to PTEROPSIDA 



1. Aquatic plants. 

2. Plants floating; leaves small, simple or bilobed Salviniaceae 

2. Plants rooted in earth or on rocks; leaves larger, more divided. 

3. Leaves 4-partite; sporocarps attached to stipes 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 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. 
1 1. Fronds simple. 
12. Sori near midrib of frond Nephrolepis Group 

12. Sori at ends of veins, solitary or joining many veins Davallia Group 

11. Fronds more or less branched. 

13. Pinnae jointed to rachis Nephrolepis Group 

1 3. Pinnae not jointed to rachis Davallia 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 containing woody tissue; no stipules at bases of stipes. 

18. Veins free 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 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 Polypodiaceae 

22. Fronds otherwise. 

23. Sori elongate, either close to midrib or to edge. 
24. Sori close to midrib Blechnum Group 

24. Sori close to edge Nephrolepis Group 

23. Sori not elongate near midrib or edge. 

25. Sporangia acrostichoid, edge of lamina in some cases reflexed. 
26. Veins of sterile leaflets free. 

27. Fronds bipinnate Tectaria Group 

27. Fronds simply pinnate. 

28. Rhizome creeping, dorsiventral Lomariopsis Group 

28. Rhizome erect, radially arranged Blechnum Group 

26. Veins of sterile leaflets anastomosing. 

29. Rhizome creeping, dorsiventral Lomariopsis Group 

29. Rhizome erect, not dorsiventral 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 Schizaeaceae 

34. Rachis trailing, not twining; several sporangia in a sorus, indusiate . 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. 

36. Filmy ferns; lamina one cell thick Hymenophyllaceae 

36. Not filmy, lamina thicker. 
37. Massive rootstock, prostrate or erect and trunk-like, densely covered with long hairs. 

Dicksonia Group 

37. Slender creeping rhizome; hairs short Dennstaedtia Group 

35. Sori otherwise. 
38. Bases of stipes bearing prominent aerophores; fronds dimorphous, fertile with narrow 

acrostichoid pinnae Plagiogyria 

38. Bases of stipes lacking aerophores; 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 Adiantum Group 

41. Sori small and round, or acrostichoid Polypodiaceae 

40. Sporangia arranged otherwise. 

42. Annulus almost apical; fronds simple or branched dichotomously; sporangia on special 
appendages 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 Athyrium Group 

48. Unicellular slender hairs on frond and on scales 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. 
54. Scales clathrate Vittaria Group 

54. Scales not clathrate Grammitidaceae 

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 



xm 



Flora Malesiana 



56. Rachis not grooved in this way Adiantum Group 

55. Sori not at ends of veins or not so orotected. 

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 from margin 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 



XIV 



6. KEYS 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 Helininthostachys 

2. Fertile segment of frond amply branched with spreading branches; sterile segment (in Malaysian 
species) pinnately branched with many small divisions Botrychium 

MARATTIACEAE 

1. Sporangia in each group along the veins near margins of leaflets; veins free. 

2. Sporangia in each group united laterally 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 

surface; veins anastomosing Christensenia 

OSMUNDACEAE 

1. Fertile pinnae quite diff"erent from sterile, lacking a green lamina; lamina of sterile pinnae not trans- 
lucent Osmunda 

1. Fertile pinnae not diff'erent 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 Schizaea 

1. Fronds of adult plants scandent (rachis twining) with very short primary rachis-branches bearing 
leafy secondary branches 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 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 and protruding from old sori Trichcmanes 

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. 41^5). 

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 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 \n 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. II, vol. 1^ 



5. Sori otherwise. 

6. Fronds small, fertile ones (or fertile parts) contracted Lemmaphyllum 

6. Fronds larger, not dimorphous Lepisorus and Neochelropteris 

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. 
10. Sori at the ends of free veins (veins conspicuous). 
11. All veins free Thylacopteris 

11. Some veins anastomosing Goniophlebium 

10. Sori not 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 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. 

17. Fronds pinnate, sterile pinnae jointed to rachis Photinopteris 

17. Fronds pinnatifid Merinthosorus 

16. Fertile fronds wholly fertile. 

18. Fronds strongly trilobed Christiopteris 

18. Fronds simple. 

19. Fronds thin, all veins visible. 
20. Fronds c. 10 cm long; terrestrial Dendroglossa 

20. Fronds c. 30 cm long; rock-plants or epiphytes Leptochilus 

19. Fronds coriaceous, at most main veins visible. 

21. Sterile fronds narrow Oleandropsis 

21. Sterile fronds circular to broadly lanceolate. 

22. Sterile fronds circular Pycnoloma 

22. Sterile fronds longer than wide, apex acute 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. Fronds on special branches of rhizome Thayeria 

26. Fronds not on special branches Aglaomorpha 

25. Bases of fronds not humus-collecting. 

27. Fronds dimorphous Holostachyum 

27. Fronds not dimorphous Arthromeris 

23. Fronds simple or pinnatifid, segments not jointed to rachis. 

28. Separate humus-collecting fronds present Dendroconche 

28. Separate humus-collecting fronds lacking. 
29. Sori elongate, oblique, parallel to main lateral veins. 

30. Fronds thin, all veins visible 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. 

32. Sori elongate, near margins of thin frond 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 

1. Fronds simple and entire or with somewhat undulate edges. 

2. Sori round or obliquely elliptical, separate Grammitis 

2. Sori elongate parallel to costa and edge, or confluent near apex of frond. 
3. Sori in groo\es between margin and costa. 

4. Fronds almost terete apart from widened apical fertile part Nematopteris 

4. Fronds of about uniform width throughout Scleroglossum 

3. Sori more or less confluent near apex, not in grooves Oreogrammitis 

1. Fronds pinnately lobed to bipinnately lobed. 

5. One vein (sometimes forked) and one sorus on each lamina-lobe. 
6. Fertile lobes fiat. 
7. Sori superficial Xiphopteris 

7. Sori deeply sunk in submarginal pockets Prosaptia 

6. Fertile lobes partly reflexed, protecting the sori. 

8. Lobes thin, only basiscopic margin refle.xed 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 . . Prosaptia 

REMAINING GENERA OF FERNS 
ADIANTUM Group 

I. Water plants; sporangia borne singly and protected by reflexed edges of the lamina. 

Ceratopteris 
L 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 Cerosora 

3. Fronds simple or simply pinnate, or palmate with entire leaflets. 
4. Veins anastomosing only near edge of lamina; fronds simple or palmate; paraphyses (if present) 

with distinctive apical cell 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 seiisii Copel.). 

Schizolepton 
2. Scales on rhizome and bases of stipes. 
6. Lower surface of fronds covered with white or yellow powder. 
7. Sori along whole length of veins on lower surface Pityrogramma 

7. Sori at ends of veins, protected by reflexed lobes of margin Cheilanthes 

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 by a narrow wing. 

10. Fertile lobes much broader than sterile, the whole of each margin bearing a broad thin reflexed 
indusium Onychium 

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 pinnatilid; sori quite continuous along edge Doryopteris 
11. Fronds branched with distinct stalked leaflets Pellaea 

8. Sori otherwise. 
12. Sori elongate along all or most of the veins. 
13. Fronds simple Hemionitis 

13. Fronds pinnate or bipinnate Coniogramme 

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 m.uch-divided fronds .... Loxoscaphe 
1. Sori distinctly elongate along the veins; segments of frond usually with more than one vein. 

xvn 



Flora Malesiana [ser. II, vol. 1^ 



2. Sori on adjacent veins opening towards each other, a raised line remaining between the sori when 
they are ripe Diplora 

2. Sori all facing the same way except in a few species which have simple fronds and then the raised 
line between adjacent sori lacking Asplenium 

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 Athyrium 

3. Sori not elongate along veins Cystopteris 

2. Groove of rachis not open to admit grooves of branches 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 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 Blechnum 

2. Sori not continuous throughout length of a leaflet; one sorus to each areole along the costa. 

3. Small ferns, simply pinnate; edces of pinnae sharply toothed Doodia 

3. Large ferns, at least bipinnatifid; edges of pinnae not sharply toothed . . Woodwardia 
1. 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 Scyphularia 

2. Fronds pinnately branched. 

5. Fronds more or less copiously persistently hairy; lamina thin; indusium small, of various shapes. 

Davallodes 
5. Fronds not hairy when mature. 

6. Indusium attached along base and whole of sides. 

7. Fronds trifoliate or simply pinnate with elongate narrow pinnae .... Scyphularia 

7. Fronds more copiously branched Davallia 

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 Araiostegia 

8. Fronds of various form, lamina coriaceous or fleshy. 

9. Scales with long acicular tips Trogostolon 

9. Scales lacking long acicular tips Humata 

DENNSTAEDTIA Group 

1. Sori in cups at ends of veins. 
2. Rhizome slender, creeping, hairy Demistaedtia 

2. Rhizome stout, erect, scaly 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 

xvm 



Dec. 1959] Keys to the genera of Pteropsida 

pocket-shaped (attached by base and sides), opening towards margin .... Microlepia 
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 indusium Histiopteris 

6. Veins free; inner indusium present Pteridium 

5. Sori singly at ends of veins, or uniting a few veins, not continuous along all edges of lamina. 

7. No inner indusium Hypoiepis 

7. Inner indusium present 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 Cystodium 

1. Fertile leaflets little different from sterile in size and shape. 

3. Pinnules deeply lobed almost at right angles to costa, not pinnate .... 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, difi"erent from 

the scales. 
2. Sorus when young spherical and quite enclosed by indusium which when ripe breaks to expose 

sporangia. 
3. Sori on distinct stalks Peranema 

3. Sori sessile Diacalpe 

2. Sorus otherwise; indusium attached on basiscopic side of sorus. 

4. Hairs sparse; scales broad and thin Acrophorus 

4. Hairs copious; scales thick, elongate Stenolepia 

1. Fronds scaly; no hairs as distinct from scales (some scales may be narrow and almost hair-like). 

5. 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 stiS" slender points. . Polystichum 
9. Indusium reniform; teeth not so produced Dryopteris 

6. Two vascular bundles in base of stipe 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 di-ided, 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 
marginal sorus Lindsaea 

LOMARIOPSIS Group 

1. Rhizome short-creeping with fronds close together; epiphytes or rock-plants, not high-climbing. 
2. Fronds always simple; stipes jointed to phyllopodia; veins free except near margin; usually epiphytes. 

Elaphoglossum 
2. Fronds usually pinnate (if simple, veins much anastomosing); stipes not jointed to rhizome; usually 
rock-plants. 

3. Veins free Egenolfia 

3. Veins anastomosing Bolbitis 

1. Rhizome high-climbing. 

4. Veins free. 

XIX 



Flora Malesiana [ser. II, vol. 1^ 



5. Pinnate or bipinnate, all pinnae and pinnules jointed at base (except on some bathyphyjls). 
6. Fronds of high-climbing part of plant simply pinnate (bathyphylls sometimes bipinnate); only 
two rows of fronds on rhizome (two meristeles in internodes) .... Teratophyllum 

6. Fronds of high-climbing part of plant usually bipinnate; more than two rows of fronds on mature 
rhizome (more than two meristeles in internodes) Arthrobotrya 

5. Pinnate; terminal leaflet not jointed at base. 

7. Fertile pinnae fully acrostichoid Lomariopsis 

7. Fertile pinnae bearing separate exindusiate sori on small lobes at ends of veins. 

Thysanosoria 

4. Veins anastomosing Lomagramma 

NEPHROLEPIS Group 

1. Fronds pinnate, pinnae jointed to rachis. 

2. Stipes jointed to rhizome or to outgrowths from rhizome Arthropteris 

2. Stipes not jointed to rhizome Nephrolepis 

1. Fronds simple Oleandra 

PLAGIOGYRIA Group 

Single genus Plagiogyria 

PTERIS Group 

1. Sori marginal or submarginal, more or less elongate, protected by a reflexed indusium. 

2. Rhizome scandent Lepidocaulon 

2. Rhizome not scandent. 

3. Sorus along one edge of ultimate lobes of lamina Hemipteris 

3. Sorus along both edges of ultimate lobes. 
4. Sori joining many veins, usually one sorus along each edge of a lobe of the lamina. 

Pteris 
4. Sori short, often more than one on each edge of a lobe; fertile margin thickened Schizostege 
1. Sori acrostichoid. 

5. Stock stout, erect; veins reticulate throughout; only distal pinnae fertile . . Acrostichum 
5. Rhizome slender, long-creeping or climbing; veins forming one series of narrow areoles by costa 

(seen near apex of leaflet); fertile fronds normally wholly fertile .... Stenochlaena 

TECTARIA Group 

1. A tooth present in each sinus between lobes of lamina, the tooth not in the plane of the lamina. 
2. Vascular bundles in stipes numerous, not in a simple ring; veins more or less anastomosing. 
3. Veins anastomosing in a single series of costal and costular areoles .... Pleocnemia 

3. Veins more copiously anastomosing Arcypteris 

2. Vascular bundles in stipes in a simple ring; veins free Pteridrys 

1. No tooth present in sinuses between lobes of lamina. 

4. Pinnae jointed to rachis Cyclopeltis 

4. Pinnae not jointed to rachis. 
5. Fertile leaflets acrostichoid and very much contracted as compared with sterile ones. 

6. Veins free; sterile frond much divided . . Psomiocarpa 

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 Heterogoniura 
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. 
1 1 . Veins free. 
12. Basal basiscopic vein of a vein-group springing from the costule. 

13. Indusia peltate 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 iobed, basal basiscopic lobe not largest; few free veins in areoles. 

Heterogonium 
17. Basal pinnae Iobed or not; if Iobed, basal basiscopic lobe largest; many free veins in areoles. 

Tectaria 

THELYPTERIS Group 

1. Fronds bearing many buds on rachis 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 anastomosing, often with additional enclosed areoles . Dictyocline 

4. Indusia present Sphaerostephanos 

3. Sori not elongate, or slightly so if without indusia {Abacopteris Fee, Haplodictyum Presl). 

Cyclosorus 

VITTARIA Group 

1. Frond very small, with one vein only; sorus near apex, along the vein . . Monogramma 
1. Frond with lateral veins as well as a main vein. 
2. Frond linear, sori one or more, on separate free branches, which run close to and parallel with 

main vein Vaginularia 

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 Vittaria 

3. Sori along veins, variously disposed; veins anastomosing copiously throughout the lamina. 

Antrophyum 

MARSILEACEAE 

Sole genus Marsilea 

SALVINIACEAE 

1. Leaves less than 1 mm long, bilobed, with one lobe submerged Azolla 

1. Leaves at least I cm long, simple, floating with upper surface fully exposed . . . Salvinia 



XXI 



7. BIBLIOGRAPHY 

A list of books and papers dealing with the taxonomy of Malaysian ferns, published 
subsequent to Christensen, Index Filicum, Suppl. 3 (\92>A) 



Alston, 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, pi. 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. 

Alston, A. H. G. & R. E. Holttum, 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. Posthumus, Varenflora voor 
Java. Buitenzorg, 1939. 

Ching, R. C, On the genus Onychium Kaulf. from 

the Far Orient. Lingn. Sc. 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 pi., 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. 
— , A revision of 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, I.e. 275-334, pi. 6-7. 
— , Ditto, 10. I.e. 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, XXXI. Aleurito- 

pterisFee. 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. I.e. 
33-52. 



— , Ditto, V. Descriptions of 36 new species of 

ferns. I.e. 53-73, pi. 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, pi. 11-20. 

Clausen, R. T., A Monograph of the Ophioglos- 
saceae. Mem. Torrey Bot. CI. 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, 

pi. 1-14. 
— , Additional ferns of Kinabalu. Philip. J. Sc. 56 

(1935) 471^83, pi. 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, pi. 1-23. 
— , Hymenophyllum. Philip. J. Sc. 64 (1937) 

1-188, pi. 1-89. 
— , Genera Hymenophyllacearum. Philip. J. Sc. 

67 (1938) 1-110, pi. 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, pi. 1-10. 
— , Notes on Hymenophyllaceae. Philip. J. Sc. 

73 (1941) 457-469, pi. 1-4. 
— , Holttumia, genus novum. Philip. J. Sc. 74 

(1941) 153-156, pi. 1. 
— , Gleicheniaceae of New Guinea. Philip. J. Sc. 

75 (1941) 347-361, pi. 1-6. 
— , Miscellaneous ferns of New Guinea. Philip. J. 

Sc. 76 (1941) 23-25. 
— , Filicum no varum C XX VIII diagnoses. Un. 

Cal. Publ. Bot. 18 (1942) 217-226. 
— , Ferns of the Second Archbold Expedition to 

New Guinea. J. Am. Arb. 24 (1943) 440-443. 
— , Genera Filicum. Waltham, Mass., 1947. 
— , Cyathea in New Guinea. Philip. J. Sc. 77 

(1947) 95-125, pi. 1-15. 
— , Pteridaceae of New Guinea. Philip. J. Sc. 78 

(1950) 5-41, pi. 1-6. 

— , Aspleniaceae and Blechnaceae of New Guinea. 

PhiUp. J. Sc. 78 (1950) 207-229, pi. 1-6. 
— , Aspidiaceae of New Guinea. Philip. J. Sc. 78 

(1951) 389^75, pi. 1-44. 

— , Grammitis. Philip. J. Sc. 80 (1952) 93-276, 

pi. 1-6. 
— , New Philippine ferns, IX. Philip. J. Sc. 81 

(1952) 1-47, pi. 1-26. 

— , Grammitidaceae of New Guinea. Philip. J. Sc. 

81 (1952) 81-119, pi. 1-10. 
— , New Philippine ferns, X. Philip. J. Sc. 83 

(1954) 97-101, pi. 1-5. 

— , New Philippine ferns, XI. Philip. J. Sc. 84 

(1955) 161-165, pi. 1-2. 



XXII 



Bibliography 



DoNK, M. A., Notes on Malesian ferns, I. On the 
genus Lemmaphyllum Presl. Reinwardtia 2 
(1954) 403-410. 

HOLTTUM, R. E., The tree-ferns of the Malay 

Peninsula. Card. Bull. Str. Settl. 8 (1935) 

293-320, pi. 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, pi. 8-16. 
— , A redefinition of the genus Teratophyllum. 

Gard. Bull. Str. Settl. 9 (1938) 355-362, pi. 

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. 
— , A new fern from Malaya. Gard. Bull. Sing. 14 

(1953) 8. 
— , A new Malayan Alhyrium. Kew Bull. 1953, 

545 (1954). 
— , A revised Flora of Malaya, 11. 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, 

41^5. 



— , 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. Need. 6 (1957) 599-601, 1 fig. 

Nakai, 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. Verb. Kon. Ak. 

Wet. A'dam sect. 2, 36, no 5 (1937) 1-67. 
— , Ditto, //. On the fern-flora of Australia, and 

its relation to that of neighbouring countries, 

especially of the Malay Archipelago. Verb. Kon. 

Ak. Wet. A'dam sect. 2, 37, no 5 (1938) 3-35. 
— , Ditto, HI. The ferns of the Lesser Sunda 

Islands. Ann. Jard. Bot. Btzg, hors sir. (1943) 

35-113. 

Rensch, I., Fame und Bdrlappe der Sunda-Ex- 
pedition Rensch. Hedwigia 74 (1934) 224-256, 
pi. 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, pi. 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, pi. 5-25. 



XXIII 



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. 5/?ore5monolete 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 siibg. Diplopterygium (Heer, Flora Fossilis Arctica, III) 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 

0) 



Flora Malesiana [ser. II, vol. 1^ 



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 off"er evidence hard to reconcile with 
Copeland's opinion that the family is of antarctic origin (Genera Filicum, p. 26). 

Ecology. All species 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 efl"ectively 
pinnate form of branching has occurred, seen at its full development in D. speciusa (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 Copel.); in others they are protected by the reflexed 
margins of the lamina and by outgrowths of tissue from the costa [G. vulcanica Bl., fig. Id-e). Nakai 
[I.e.), 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.e.). 
Both trilete and monolete spores occur in the family. Nakai [I.e.) proposed a basic division of the family 



Dec. 1959] Gleicheniaceae (Holttum) 



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. Stokey (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 Siokey'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 stibg. 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, pi. 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 Ihe 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) cunningliamii. 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, I.e.). 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. PI. 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 Diels (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 Presl 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 1. Gleichenia 

1. Young parts of plants protected by branched hairs of various form, scales lacking. Sori of 8-15 or 

more sporangia. Veins forked at least twice 2. Dicranopteris 



Flora Malesiana [ser. II, vol. P 



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 stipiilar 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 ; ultimate branches either simply pinnatifid 
or bipinnatifid; lamina 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 TO THE SPECIES 

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. vulcanica 
5. Costae of fully developed fronds usually quite glabrous; tissue adjacent to the costa much swollen. 

Sporangia 2 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 2-2^2 cm. Upper surface of the lamina not swollen between 

the veins. Stipular leaflets present 7. G. volubilis 

10. Pinnules commonly 6-9 by 0.9-1.3 cm. Upper surface of the lamina much swollen between 



Dec. 1959] Gleicheniaceae (Holttum) 



the veins. No stipular leaflets 8. G. bullata 

8. Rachis-branches not persistently quite covered with scales nor with a felt of stellate hairs on the 
lower surface. 
1 1. Very few persistent scales on the lower surface of rachises and costae. Pinnules to 22 by 3.5 cm. 

9. G. longissima 
1 1. 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 2^2 mm wide; scales of dormant apex uniformly dark 
brown 11. G. angustiluba 

14. Lamina thin; segments of lamina 3'/2 nim 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 3V2-4V2 mm apart. 14. G. sordida 
15. Pinna-rachises sparsely scaly, soon glabrous; costules S^jo-l mm apart. 12. G. eimeri 

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 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 

in line with the part of the rachis beyond it 16. G. matthewii 

17. Distal half of each rachis-branch not thus zig-zag. 

18. Pinnules c. 10 by 2 cm 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 persistent on the lower surface of veins and 

lamina 19. G. conversa 

1. Ultimate branches simply pinnatifid. (subg. Mertensia). 

20. 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 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 straight 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 24. G. oceanica 

25. Primary branch-systems twice forked; stellate hairs (or at least their bases) persistent on the 

lower surface of veins 25. G. erecta 

20. Stipule-like leaflets absent (sometimes present in G. loheri var. major). 
26. Margins of segments of lamina not distinctly toothed. 
27. Only the ultimate branches fully leafy (penultimate without lamina or with some scattered 
segments). 

28. Ultimate branches to 20 cm long 26. G. pulchra 

28. Ultimate branches 30-40 cm long. 
29. Segments of the lamina to 25 mm long, costules 57,-6 mm apart. Primary rachis-branches 

usually 3 times forked 27. G. pseudoscandens 

29. Segments of lamina to 17 mm long, costules 4 mm apart. Primary rachis-branches usually 

twice forked 28. G. alstonii 

27. At least the penultimate branches fully leafy, lower ones sometimes also. 



Flora Malesiana [ser. II, vol. 1^ 



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 1st and 2nd orders 3-4 

cm long. Scales on the main rachis-apex brown, more than 1 mm wide at base; scales on the 

costae rusty brown 29. G. vestita 

31. Segments 3-6 mm long, costules 3 mm apart. Rachis-branches of 1st and 2nd orders 1-2 cm 
long. Scales on the main rachis-apex very dark, narrower; scales on the costae dark at base, 
paler distally 30. G. bolanica 

30. Lamina-segments much more than twice as long 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 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 25^5 cm long, 3-4V2 cm wide; costules on ultimate branches 4-5 mm 

apart 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. Habellata 
35. Margins toothed only near apex of segments. Costules at wider angle to costae. 35. G. hirta 

1. Subgenus Gleichenia 
HoLTT. Reinwardtia 4 (1957) 262.~CaIymella Presl, Tent. Pterid. (1836) 48; 
Ching, Sunyatsenia 5 (1940) 287; Nakai, Bull. Nat. Sc. Mus. Tokyo u. 29 (1950) 
40.— Gleicheniastrum Presl, Abh. (K.) Bohm. Ges. Wiss. M.-N. CI. 5 (1848) 
338; Nakai, I.e. 42. — Gleichenia subg. Eugleichenia Diels in E. & P. Pfl. Fam. 
1, 4 (1900) 355.— Gleichenia; Copel. 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 shining, very dark brown with narrow pale edge; 

(1929) 292, t. 1. — G. circinnata var. bonieensis Bak. similar but smaller scales scattered on lower sur- 

J. Bot. 17 (1879) 37. — G. bonieensis C. Chr. faces of rachises and less abundantly on costae; 

Card. Bull. S. S. 7 (1934) 211. — Calymella tor- ultimate branches 20-30 cm long, bearing many 

neensis Ching, Sunyatsenia 5 (1940) 288. leaflets, costae of adjacent leaflets 4-6 mm apart; 

leaflets 2-5 cm long, 2'/o-4 mm wide; lobes c. 

var. peltophora. 1 '/t mm wide at base, gradually narrowed to 

Rhizome-scales ovate, dark, entire. Determinate rounded apex, edges slightly reflexed, lower sur- 

bipinnatifid //-o/z^jT to 40 cm high (including stipe), face slightly concave, glaucous, bearing scattered 

often fertile; branched fronds bearing 1 or few circular scales 0.2-0.3 mm diameter. Sori, if 

pairs of branches; primary branches bipinnatifid- present, each sunk in a circular depression in the 

determinate or once forked with a permanently substance of the lamina, the lower surface of the 

dormant apex in the fork, the primary branch lamina somewhat raised round the edge of the 

always bearing leaflets (where primary branches depression which occupies about half the width of 

are not forked, the main rachis may be leafy below the base of a lobe. 

junction with branches); scales on resting apex Type: Copeland s.n., 1 May 1917, Mt Matutum, 

of rachis to P/2 by 1 mm, ovate, entire, convex, Mindanao, 1600 m (H. Copel.). 



Dec. 1959] 



Gleicheniaceae (Holttum) 




Fig. 1. Gleichenia micropliylla 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, b. one lobe of lamina, 
cleared to show veins and position of sorus, x 1 3, c. a scale, < 50. — G. vulcanica Bl. d. Part of pinnule 
from below (scales removed), :■ 10, e. same cleared to show veins and position of sorus, X 13. — G. 
dicarpa R. Br.,/. Lower surface of part of pinnule, x 10, g. same cleared to show veins, X \7>,h. 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., /. 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. Diplopterygiiim), I. 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 Holtt. 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 2V2 crn long, sterile. 

Distr. Borneo (Sarawak: Mt Murud), 2500 m, 
once found. 



2. Gleichenia microphylla R.Br. Prod. (1810) 161; 
V. A. V. R. Handb. Suppl. (1917) 80, incl. var. 
semivestita v. A. v. R. — G. semivestita Labill. 
Sert. Austro-Cal. (1824) 8, t. W.—Calymella 
microphylla Presl, Tent. Pter. (1836) 49. — 
Gleicheniastrum microphyllum Presl, 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 
i?G. circinnata Sw.) Ching, Sunyatsenia 5 (1940) 
288. — Calymella semivestita Ching, I.e. — 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. II, vol. 1^ 




Fig. 2. Gleichenia microphvUa R. Br. on edge of dwarf forest on wet sandy soil near summit of Kedah Peak, 

Malaya, 1200 m (Holttum, 1953). 



Dec. 1959] 



Gleicheniaceae (Holttum) 



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. Scales 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'/2-4 mm 
wide; lobes of la/nina 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^ 
(rarely 5), surrounded by brown hairs not longer 
than the ripe sporangia. 

Type: Robert Brown, Port Jackson, N.S. 
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 Eraser'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 Leptospermiim 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. 

3. Gleichenia vulcanica Bl. En. PI. Jav. (1828) 
251; Racib. F1. Btzg 1 (1898) 10; v. A. v. R. 
Handb. (1909) 56; Backer & Posth. Varenfl. 
Java (1939) 353. — Calymella vulcanica Presl, 
Abb. (K.) Bohm. Ges. Wiss. M.-N. CI. 5 (1848) 
338; Ching, Sunyatsenia 5 (1940) 287; Nakai, 
Bull. Nat. Sc. Mus. Tokyo //. 29 (1950) 42.— G. 
dicarpa var. vulcanica Christ, Ann. Jard. Bot. 
Btzg 15 (1898) 75. — G. squamosissima Copel. 
Philip. J. Sc. 75 (1941) 348, pi. \.— Calymella 
squamosissima Nakai, I.e. — Fig. Id-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 (C squamosissima Copel.). 




Fig. 3. Gleichenia vulcanicaBL. 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. Chr. Ark. Bot. 9" (1910) 33.— Calymella 
dicarpa Presl, Abh. (K.) Bohm. Ges. Wiss. M.-N. 
CI. 5 (1848) 338.— Calymella circinnata (non G. 
circinnata Sw.) Ching, Sunyatsenia 5 (1940) 288. 
—Fig. If-h. 

Like G. vulcanica, but the leaflets 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. II, vol. 1^ 



Type: Robert Brown, Tasmania (BM; dupl. 
at K). 

Distr. Australia, New Caledonia, in Malaysia: 
New Guinea, Philippines (Mindanao). 

Ecol. On Normanby and Rossel Islands, SE. 



New Guinea, occurring at 750-850 m, in open 
forest or rocky places. 

Note. Though typical specimens of G. vulcanica 
in Java differ markedly from typical specimens of 
G. dicarpa in Australia, some specimens on moun- 




-evel 5/ 



Fig. 4. a. Diagram of branching in Gleichenia sitbg. Diplopterygium. — G. longissima Bl., b. Part of 
rachis-branch bearing two pinnules, X 2/3, c. segment of lamina, lower surface, showing veins and son, 
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,/. edge of scale, X 133, g. stellate 
hair, x 33. - G. converja v. A. v. R., /j. A single scale, x 8, /. edge of scale, x 1 33, y. stellate hair, x 33. 



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 Svs'artz at Stoci<holm (I.e.) 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. circiunata 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 C. micropliylla 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. CI. 34 (1907) 249, 
p.p. — Sticherus § Hicriopteris C. Chr. in Verdoorn, Man. Pterid. (1939) 530. — 
Hicriopteris {jwn Presl) Ching, Sunyatsenia 5 (1940) 277; Copel. Gen. Fil. 
(1947) 2S.—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 (iat. 70-7 1'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. 1 1 (1902) 268; v. A. v. R. Handb. 
(1909) 795; Suppl. (1917) S2.— 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, 2'/2-3 cm wide, costae 2'/S-3'/2 cm 



apart; lamina lobed almost to the costa, costules 
at about 45' to the costa and 3'/2^'/2 rnm apart; 
segments of lamina 2-21/2 rnm 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. 



12 



Flora Malesiana 



[ser. II, vol. P 




Fig. 5. Gleichenia longissima Bl. on edge of forest, Taiping Hills, Malaya, c. 800 m (Holttum, 1952J. 



Dec. 1959] 



Gleicheniaceae (Holttum) 



13 



6. Gleichenia palcacea (Copel.) Holtt. Rein- 

vvardtia 4 (1957) 265. — Hicriopteris paleacea 
Copel. Philip. J. Sc. 81 (1952) 3. 

Scales on main rachis-apex not seen; scales 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 rac/iis- 
branch more than 100 cm long (collector's note); 
costae IVi cm apart, at right angles to rachis; 
largest pinnules 15-16 cm long, ^'^-^ 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; lamina- 
segments thin but firm, hardly narrowed abo\e the 
base, edges slightly refiexed 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, 
K, Bo, L, 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. 



stellate hairs on costules and lower surface of 
veins rusty brown. Primary rachis-hranches 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 3'/2-4 mm apart, 
at right angles to costae; lamina-segments firm, 
slightly narrowed above the base, their edges 
refiexed 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, 1 Yi cm 
wide), with thinner lamina and veins hardly 
prominent beneath. 

var. peninsularis Holtt. 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. 



7. Gleichenia volubilis Jungh. Java 1 (1853) 592, 
664; V. A. v. R. Handb. Suppl. (1917) 83,/7./7.; 
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 Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 47; 
Racib. F1. Btzg 1 (1898) 11 ; v. A. v. R. Handb. 
(1909) 58 (arachnoidea); nan G. arachnoidea 
Cl'NN. 1844. — G. glaiica var. arachnoides C. Chr. 
Ind. Fil. (1905) ^20.— 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 1 1/2 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- 
rachises, 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 
raised points of attachment of former scales; 



8. Gleichenia bullata Moore, Ind. Fil. (1862) 374; 
C. Chr. Card. Bull. S.S. 7 (1934) 212.— G. 
volubilis (non Jungh.) v. A. v. R. Handb. Suppl. 
(1917) 83, p.p. — Hicriopteris 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 lamina (when dry) 
much sunken, the surface between them swollen. 

Type: Low, Mt Kinabalu, 7000 ft (K). 

Distr. Malaysia: Borneo (Sarawak: Mts Duht 
and Tibang; N. Borneo: Mt Kinabalu), New 
Guinea (Mt Dayman). 

Ecol. Forming thickets like G. volubilis, 
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. 

9. Gleichenia longissima Bl. En. PI. Jav. (1828) 
250; Racib. F1. Btzg 1 (1898) 10; Backer & 



14 



Flora Malesiana 



[ser. II, vol. H 



PosTH. Varenfl. Java (1939) 256; Holtt. Rev. Fl. 
Mai. 2 (1955) 67. — G. exceha 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- 
sinia Ching, Sunyatsenia 5 (1940) 280. — Diplop- 
terygiiim longissimum Nakai, Bull. Nat. Sc. Mus. 
Tokyo //. 29 (1950) 53.— Fig. 4b-g, 5. 

Scales on resting apex of rachis as in G. voliibilis 
JuNGH.; young expanding parts of fronds covered 
with loose indumentum of narrow rust-coloured 
scales and lax stellate hairs, a few such scales 
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, 2 1/2-3 '/2 cm wide, costae 
2.8-3 1/2 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.) Holtt. comb, 
now— Hicriopteris clemensiae Copel. Un. Cal. 
Publ. Bot. 18 (1942) 1\1.—G. papuana Holtt. 
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 1 Vi mm 
long and Vi 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'/2 cm apart, largest 12 cm long, 2 cm 
wide; costules 21/2-3 mm apart; lamina thin, 
veins slightly prominent on lower surface, not on 



upper; sori commonly 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 Holtt. Reinwardtia 4 
(1957) 263. 

Rhizome 7 mm diam., young parts covered with 
shining dark brown scales 5-7 by 1 V2 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 3-3 '/2 cm wide, 2-31/2 cm 
apart, the distal ones distinctly deflexed, the lower 
ones at right angles to rachis; all segments of the 
lamina c. IVi 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, Leafl. 
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) 



15 



longest 1-1 V^ mni 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, 3'/2^ cm wide, 4'/2-5 cm apart; 
costules 5 1/2-7 mm apart, at right angles to costa; 
several segments of lamina constricted at base, 
widely separated and joined by a wing '/2 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, 
US, F, L, P). 

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 defiexed, making 
angles of about 75"" with the rachis; costae on 
large fertile rachis-branches 4 1/2 cm apart, on 
smaller sterile branches IVi-'^Vi cm apart; fertile 
pinnules 16-20 cm long, widest part 3'/^-3.8 cm 
wide, costules 4—5 mm apart; lamina 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. 3'/2 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 
Brause, Bot. Jahrb. 56 (1920) 210. — Hicriopteris 
novoguineensis Copel. Philip. J. Sc. 75 (1941) 358. 



— Hicriopteris astrotricha Copel. I.e. — Diplo- 
pterygium novoguineense Nakai, Bull. Nat. Sc. 
Mus. Tokyo n. 29 (1950) 54. — G. sumatrana 
Holtt. 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 2'/2-4'/2 cm apart; largest pinnules 
15-20 cm long, 2V2-3 cm wide, slightly defiexed 
or not; costules iVi-^Vi rnm 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, IVi-^Vi 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. Chr. Card. Bull. S.S. 7 (1934) 212, incl. \ar. 
floccigera C. Chr.; Holtt. Rev. Fl. Mai. 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 
defiexed, distal pinnules more acutely defiexed 
than basal ones; lobes of lowest pinnules usually 
only slightly enlarged and not stipuliform, but 
stipule-like leafiets, with broad blunt lobes, 
sometimes produced; costules 4V2-5V2 rnm 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 



16 



Flora Malesiana 



[ser. II, vol. 1^ 




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) 



17 



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. lungissima; above 750 m the latter, which forms 
denser thickets because of its closer pinnules, is 
ihe 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 Holtt. Reinwardtia 4 
(1957) 265. 

Scales of main rachis-apex hardly 1/2 rnm wide, 
edges with their setae shining dark brown, middle 
part paler; costae bearing a few scales near the 
base, the frond otherwise glabrous. Rachis- 
brauches 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 
21/2 cm wide, the costae 4 cm apart; costules 
4'/2 Tim apart; lamina firm, glaucous beneath, 
lobed to \-\V2 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 Holtt. Reinwardtia 4 
(1957) 264. 

Scales on apex of rachis c. 5 mm long, hardly 
V2 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 3-3 '/2 rnm apart; lamina 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 3'/2 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 2'/^ m long. 

18. Gleichenia blotiana C. Chr. Bull. Mus. Hist. 
Nat. Paris II, 6 (1934) 103.— Hicriopteris blotiana 
Ching, Sunyatsenia 5 (1940) 279. — Diploptery- 
gium bloiiunum 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, 3 '/a 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 31/2—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: Petelot 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. longissima, at 1500 m. 

19. Gleichenia conversa v. A. v. R. Bull. Jard. 
Bot. Btzg 11, n. 20 (1915) 17; Handb. Suppl. 
(1917) 81; Backer & Posth. Varenfl. Java (1939) 
255. — Diplopterygium conversum Nakai, Bull. 
Nat. Sc. Mus. Tokyo //. 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. 



11 



Flora Malesiana 



[ser. II, vol. P 



3. Subgenus Mertensia 
Hook. Sp. Fil. 1 (1844) 6, pro § 2; Holtt. Reinwardtia 4 (1957) 266.— Mertensia 
WiLLD. Kongl. Vet. Ak. Nya Handl. 25 (1804) 163, p.p., non Roth 1797.— 
Gleichenia subg. Mertensia sect. Holopterygium Diels in E. & P. Pfl. Fam. 1, 4 
(1900) 353.— Dicranopteris {non Bernh.) Und. Bull. Torr. Bot. CI. 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. So. Mus. Tokyo n. 29 (1950) 7.— 
Sticherus § Eu-Sticherus C. Chr. in Verdoorn, Man. Pterid. (1938) 530. — 
Mesosorus Hassk. Fil. Jav. 1 (1856) 2, p.p. — Fig. 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 




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. 



Dec. 1959] 



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. 




revel '5^ 



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, /. same with scales and sporangia removed, g. a single scale from costa, 
X 22. - G. hirta Bl. h. Lower surface of part of frond, x 4, /. tip of a segment of the lamina, showing 
toothed edge, j. a scale from the costa, x 20. 



20 



Flora Malesiana 



[ser. II, vol. P 



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. Chr. Brittonia 2 
(1937) 27. — Sticherus reflexipinnula Copel. Philip. 
J. Sc. 75 (1941) 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 
3 1/^-4 cm wide (shorter if primary branch is 
twice forked), penultimate branches 31/2-9 cm 
long; scales on resting apex of main rachis 3 V2 mni 
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, 1/2 rnrn wide, thin, pale, long- 
fringed; scales on costules similar and smaller, 
abundant, ranging down to stellate hairs; no 
stellate hairs on veins; lamina lobed almost to 
the costa, middle and upper segments slightly 
deflexed, lower ones much deflexed, costules 
31/2 mm apart; segments gradually narrowed to 
apex (which is sometimes refuse), 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) 
356, pi. 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 1 1/2 mm long and more than 
14 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; Holtt. Reinwardtia 4 (1957) 
271. — See for further synonyms under the varieties. 
—Fig. 7a, 8a-b, 9. 

KEY TO THE VARIETIES 

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 

surface 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 li/^-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, 

1-1 1/2 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. PI. 5 
(1810) 75. — Sticherus laevigatus Presl, Tent. 
Pterid. (1836) 52.— C. laevigata Hook. Sp. Fil. 1 
(1844) 10; Racib. F1. Btzg 1 (1898) 11; v. A. v. R. 
Handb. (1908) 59; Backer & Posth. Varenfl. 
Java (1939) 255; Holtt. Rev. Fl. Mai. 2 (1955) 
71. — Sticherus truncatus Nakai, Bull. Nat. Sc. 
Mus. Tokyo n. 29 (1950) 20. — Sticherus myriapoda 
Nakai, I.e. 12, f. 1. 

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] 



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 1 1/2 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 
2-31/2 cm long (to 5 cm) and about 2'/2 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; sort 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. 

Stichenis myriapoda Nakai was described from 



a plant in the Riouw Archipelago differing only 
from typical \ar. 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) 
HOLTT. Reinwardtia 4 (1957) 271. — G. bracteata 
Bl. ex Hook. & Baker, Syn. Fil. (1865) 14 (as 
synonym but with descr.) — G. laevigata var. 
bracteata v. A. v. R. Handb. Suppl. (1917) 85. 

Lamina-segments commonly less than 2 mm 
wide, 10-18 mm long; additional stipular leaflets 
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) Holtt. Rein- 
wardtia 4 (1957) 272. — Mertensia plumaeformis 
Presl, Abh. (K.) Bohm. Ges. Wiss. M.-N. CI. V, 



22 



Flora Malesiana 



[ser. II, vol. 1^ 



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 1-1 '/2 mm long, 
0.7-1 mm wide, on lower surface. 

Type: Cuming 377, Malacca (Prc; dupl. at K, 
BM, P, L, F). 

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 1/2 mm long; 
edges of dried lamina-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; scales on lower surface 
of costae copious, acuminate, fringed, c. 1 '/2-2 
mm long, 0.3-0.6 mm wide. 

Distr. Malaysia: Central Celebes, altitude 
2600-3000 m (once collected). 

23. Gleichenia milnei Baker, Syn. Fil. (1874) 449; 
Hook. Ic. PI. t. 1602; Holtt. Reinwardtia 4 (1957) 
270.— G. kajewskii Copel. Philip. J. Sc. 60 (1936) 
102, pi. 6. — Sticherus milnei Ching, Sunyatsenia 
5 (1940) 284.Sticherits 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 2V2-3'/2 cm wide (lower ones to 4'/2 
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. A/(7/ov5m.- Celebes,? Phil ippines(?Luzon), 
Moluccas, New Guinea, Admiralty Islands, 
Solomon Islands, New Hebrides. 

Ecol. Altitude 100-1400 m. 



24. Gleichenia oceanica Kuhn, Verh. K. K. Zool.- 
Bot. Ges. Wien 19 (1869) 583; non Christ, 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) 3SS.—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 1 '/2 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 (I.e.) are G. 
hispida and G. hirta var. paleacea; I have not 
seen the Warburg specimen. 

25. Gleichenia erecta C. Chr. Brittonia 2 (1937) 
269; Holtt. Reinwardtia 4 (1957) 268.— Sticherus 
erectus Copel. Philip. J. Sc. 75 (1941) 353.— 
Sticherus habbemensis Copel. I.e. 355, pi. 3. 

Rhizome 2'/2-4 mm diameter, young parts 
covered with thin brown ciliate scales to 3 mm 
long and I 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 iobed leaflet 1 5 mm 
long and sometimes 1 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, 
\<iT^Q?,\. lamina-segments commor\\y 15-23 mm long, 
sometimes to 40 mm, rather coriaceous, entire, 
edges much reflexed when dry, abruptly narrowed 
just above the base, the rest 2-21/2 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 falcate 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, I '/2 
mm long, V2 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 459 1, 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] 



Gleicheniaceae (Holttum) 



23 



26. Gleichenia pulchra (Copel.) Holtt. Rein- 
wardtia 4 (1957) 271. — Slicherus pulcher Copel. 
Philip. J. Sc. 75 (1941) 355, pi. 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 21/2 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 '/2-I 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. — Stichenis 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|/2-6 mm apart, at right angles 
to the costa or nearly so; segments of lamina 
17-25 mm long, 3 mm wide above the base, lamina 
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 Holtt. 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 
cm long; ultimate branches 30-35 cm long, 
3'/2 cm wide, the basal 10-15 mm sometimes 



bare of lamina; costules 4 mm apart, slightly 
oblique; segments oi lamina 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, 1 V^ 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. 






■5 



;*^ 




Fig. 10. Gleichenia vestita Bl. on ridge with cricoid 

ridge forest from Poka Pindjang to Rante Mario 

in the Latimodjong Range, Central Celebes, 

c. 3000 m (1937). 



29. Gleichenia vestita Bl. En. PI. Jav. (1828) 249; 
Racib. F1. Btzg 1 (1898) 11 ; v. A. v. R. Handb. 
(1908) 61; Suppl. (19 17) 86; Backer & Posth. 
Varenfl. Java (1939) 255. — Sticherus vestitus 
Ching, Sunyatsenia 5 (1940) 285.— Fig. 7e, 
8c-d, 10. 

Rachis of a well-grown frond bearing several 



24 



Flora Malesiana 



[ser. II, vol. P 



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; lamina 
cut down to within 1 mm of costa, segments 
distinctly oblique, thin but veryrigid, 8-1 2mm 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 1 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. — Stichenis 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; lamina 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 0.5-0.7 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 Mett. ex Kuhn, Verb. 
K. K. Zool.-Bot. Ges. Wien 25 (1875) 600; 
Racib. F1. Btzg 1 (1898) 12; v. A. v. R. Handb. 
(1908) 61; Backer & Posth. Varenfl. Java (1939) 
255; HoLTT. Reinwardtia 40 (1957) 270.— G. 
kourdersii Christ, Ann. Jard. Bot. Btzg 15 
(1898) 76, pi. 13, f. 1. — Sticherus caudatus Copel. 
Philip. J. Sc. 75 (1941) 354, pi. 2.— Sticherus 
hispidus Copel. Gen. Fil. (1947) 27. — Sticherus 
pinnatus Copel. Philip. J. Sc. 83 (1954) 98, pi. 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; lateral 
branch-systems with forks to 2nd or 3rd order; 
branches of first order leafless or only partly 
leafy, rest fully leafy; first order branches 2'/^-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; lamina 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'/2 cm long, on 
ultimate branches sometimes only 1 cm long, 
effective width when dry l'/2-2'/2 rnm, costules 
3-4 mm apart; lower surface of lamina more or 
less glaucous, veins dark and slightly prominent, 
not raised on upper surface; soriofi 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. Chr. 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 4-4 '/2 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, 1 '/2-2 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, 
I/2-I 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; 
HoLTT. Reinwardtia 4 (1957) 272. — Sticherus 
loheri CoPEL. Gen. Fil. (1947) 21. ^Sticherus 
perpaleaceus CoPEL. 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 1 '/2-2.2 cm wide, costules 
3-31/2 rnm apart, at right angles to costa or very 
slightly oblique, lamina 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 Holtt. Reinwardtia 4 (1957) 272. 

Larger than typical form of species; ultimate 
branches 20-30 cm long, 2'/2-3'/2 cm wide, costules 
3-4 mm apart; 3-lobed stipular leaflets 15 mm long 
sometimes present. 

Type: Biinnemeijer 11965, Celebes (Bo; dupl. 
at L, K). 

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. SCH. 
& Laut. F1. Schutzgeb. (1901) \AA.— 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; lamina cut almost 
to the costa, segments 2'/2 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. & SB. Australia, New Caledonia, 
New Zealand, in Malaysia: E. New Guinea 
(Schumann & Lauterbach I.e.). 

35. Gleichenia hirta Bl. En. PI. Jav. (1828) 250; 
v. A. V. R. Handb. (1908) 60; Suppl. (1917) 85; 
Holtt. Reinwardtia 4 (1957) 268. 

— See for further synonyms under the varieties. — 
Fig. 7d, 8h-i. 

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"; lamina thin, cut almost down 
to the costa i:-.to 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, IV2-2 mm long, less 
than V2 iTini wide, thin and rather pale, not 



26 



Flora Malesiana 



[ser. II, vol. P 



lacking; sori with raised receptacle, conspicuous 
paraphyses present or not. 



conspicuously hair-pointed, edges fringed with 
slender hairs; smaller scales and stellate hairs also 
present on costae, costules and veins, or quite 

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 3 '/■: cm wide. 

2. var. amoena 
2. Scales wider. Ultimate branches rarely over 25 by 2'/2 cm. 

3. Costules 4-41/2 mm apart 3. var. paleacea 

3. Costules 3'/2^ mm apart 4. \ar. Candida 

1. 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 1 ', 2-2' ; cm wide: primary rachis-branches always proliferous beyond first fork. 

5. var. amboinensis 

5. Scales on costules with very few long marginal hairs 6. var. virescens 

4. Segments of lamina c. 2 mm wide except at the very base, separated by sinuses 1-1 Y2 mm wide. 

7. var. lanuginosa 



1. var. hirta. — Dicranopteris dolosa Copel. in 
Perk. Frag. Fl. Philip. (1905) 193, t. 4 f.c.—G. 
dolosa C. Chr. Ind. Fil. (1906) 664; v. A. v. R. 
Handb. (1908) 62; Suppl. (1917) SS.Srichenis 
hirtus Ching, Sunyatsenia 5 (1940) 203 (err. 
hirsutus). 

Lateral brancli-systems forked in 3 or 4 orders; 
ultimate branches 15-22 cm long, 2-2 '/2 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.) Holtt. Reinwardtia 
4 (1957) 269.— C. amoena v. A. v. R. Bull. Jard. 
Bot. Btzg II, n. 23 (1916) 12; Handb. Suppl. (1917) 
497 — Sticherus amoeiiiis Nakai, Bull. Nat. Sc. 
Mus. Tokyo /;. 29 (1950) 13. — G. peninsularis 
Copel. Un.'Cal. Publ. Bot. 12(1931) 387.—Stic/ierus 
peninsularis Ching. Sunyatsenia 5 (1940) 284. — 
G. hirta; Holtt. Rev. Fl. Mai. 2 (1955) 71. 

Lateral brancli-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'/2-3'/2 cm wide; 
costules 4-4'/2 rnm apart; lamina-segments 3-3 V2 
mm wide above the base, strongly glaucous 
beneath; scales 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: Teysmann 16628, Lingga (Bo: dupl. 
at P). 

Distr. Malaysia: Sumatra, Lingga .Archipelago, 
Malay Peninsula. 



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. Chr. Gard.Bull. 
S. S. 7 (1934) 212; Holtt. Reinwardtia 4 (1957) 
269. — G. vestita var. paleacea Bak. J. Bot. 17 
(1879) 38.— G. hallieri Christ. Ann. Jard. Bot. 
Btzg 20 (1905) 138;v. A. V. R. Handb. (1908) 61.— 
G. barbida C. Chr. Dansk Bot. Ark. 9, 3 (1937) 
67. — Sticherus hallieri Nakai, Bull. Nat. Sc. Mus. 
Tokyo /;. 29 (1950) 18.— Sticherus barbulus Nakai, 
I.e. 13. 

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 2-2 '/2 
cm wide: costules 4-4 '/2 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. Malavsia: Borneo, Lesser Sunda Islands 
(Bali), Celebes' (Menado), 1100-3000 m. 

Ecol. On Mt Kinabalu on open places in ridge 
forest. 

4. var. Candida (Rosenst.) Holtt. Reinwardtia 
4 (1957) 269.— G. Candida Rosenst. 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 3 'A—* mm 
apart, lamina of firmer texture. 

Tvpe: Werner 72, Mt Gelu (B: dupl. at BM, 
US, L). 

Distr. East Malaysia: New Guinea, Admiralty 
Islands, Solomon Islands. 



Dec. 1959] Gleicheniaceae (Holttum) 27 

Ecol. On edges of forest and in light shade, Type: Ledermann 9935 (B). 

sometimes forming dense thickets to 3 m high, Distr. Malaysia: E. New Guinea. 

700-2150 m. Note. Collector of type specimen stated that 

e ™k^-.,^„ov /■,, A ., T> \ o^. .,-,- !>„• lower surface of lamina was white (/.e. glaucous) 

5. var. ambouiensis (v. A. v. R.) Holtt. Rem- h f h \ o y 

wardtia 4 (1957) 269. — G. amboinensis v. A. v. R. 

Bull. Dep. Agr. Ind. Neerl. «. 18(1908)3; Handb. _ , • / a r, . t ^ 

/lOAov /:t c 1 Mnn. OS /^ , '• *^''- lanuginosa (v. A. V. R.) comb. nov. — G. 

(1908) 62 Suppl. (1917) 85 (as a synonym . ;• r> v / • o /ir,i-.> 

, •, / . r 1. . u u ornamentalis Rosenst. Nova Guinea 8 (1912) 

Like var. Inria in scaliness; ultimate branches _,, . n n jl o ■ ,,n,-,. or ^ 

,,, ,w ■. / ■ i, L 1 715; V. A. V. R. Handb. Suppl. (1917) 85. — G. 

^/i-~n cm wide primary rachis-branch always ,. / • » r. vr 

vc ' L. ^c.ri ju T ornamentalis var. lanuginosa v. A. v. R. Nova 

proliierous beyond first fork and bearing 2 or ^ . , . /,„-,., _,"_., , . „ 

• / , , , Guinea 14 (1924) 23.- — Siicherus lamianus Copel. 

more pairs of secondary branches. „,.,. , c- m \,nA,^ t^^ i ^ ^ i- 

-T- T A u /D 1 1 . I , Philip. J. Sc. 75 (1941) 356, pi. 6. — G. hirta var. 

Type: Teysmann, Amboyna (Bo; dupl. at L). ^ ;■ /r> \ u r. • j ■ -i 

T< . %i 1 c r- ; 1 /- 1 u X4 I ornamentalis (Rosenst.) Holtt. Reinwardtia 4 

Distr. Malaysia: S. Central Celebes, Moluccas /-i0S7i ">fiQ 

(Ambon, Buru, Waigeo Isl.). rii. / irij a 

r- 1 /-I 1-- .1 1 Lateral branch-systems commonly lorked to 4 

Ecol. Climbing to 4 m in open places in , ■ . .i j j i i 

, ■ r . ft- orders; costae persistently and densely scaly, 

mountain forest. , . ^,. \ , -, a L 

scales as in var. nirta; costules 3-4 mm apart, 

6. var. virescens (Hieron.) Holtt. Reinwardtia segments of lamina 2 mm wide, thus separated 
4 (1957) 270. — G. dolosa var. virescens Hieron. ex by rather wide sinuses. 

Brause, Bot. Jahrb. 56 (1920) 2Q9. —Sticherus Type: Lam 1945, Doormantop, 3480 m (U; 

hirtus var. virescens Nakai, Bull. Nat. Sc. Mus. dupi. at Bo, US. L). 

Tokyo n. 29 (1950) 19. Distr. Malaysia: New Guinea. 

Similar to var. hirta. but scales on costules with Ecol. Altitude 800-2500 m; specimens from the 

very few long marginal hairs. higher altitudes are the most densely scaly. 

2. DICRANOPTERIS 

Bernhardi in Schrad. Neu. J. Bot. 1, 2 (1806) 38: Underw. Bull. Torr. Bot. 
CI. 34 (1907) 244. p.p.; Ching. Sunyatsenia 5 (1940) 272; Copel. Gen. Fil. (1947) 
28; Nakai. Bull. Nat. Sc. Mus. Tokvo n. 29 (1950) 56; Holtt. Reinwardtia 4 
(1957) ll^.—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. Mertensla. § III and § IV. Hook. Sp. Fil. 1 (1844) 11, 12. — Hicriopteris Presl, 
Epim. Bot. (1851) 26, non Ching. nee Copel. — G/eiehenia subg. Mertensia sect. 
Heleropterygiiim DiELS in E. & P. Pfl. Earn. 1. 4 (1900) 355. — Gleiehenia subg. 
Mertensia sect. Dicranopteris v. A. \. R. Handb. (1908) 56, p.p. — Mesosorus 
Hassk. Fil. Jav. 1 (1856) 2, p.p.—Y\g, 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 lamina, 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 



28 



Flora Malesiana 



[ser. II, vol. 1^ 




Fig. II. Dicranopteris cunanii Copel. Harau gorge, near Pajakumbuh, W. Sumatra, c. 500 m (Meijer, 

Jan. 1958). 



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 specie? 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 suhgQWMS Dicranopteris is much more diversified in Malaysia than in any other part of the world. 
Asmallgroupofspeciesmay 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 




Fig. 12. Diagrams showing variations of branching-habit in Dicranopteris. a. Basic habit, found in 
D. linearis {BvRM.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 (Christ) Holtt., var. alternans 
<Mett.) Holtt., and var. subspeciosa Holtt., d. special development of c, found only in D. speciosa 

(Presl) Holtt. 



30 



Flora Malesiana 



[ser, II, vol. 1^ 



*!i.-^^i ''|l^-' 







%* 






?' ',?. 




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 Ilex 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 £>. 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 




Fig. 14. Hairs of Dicranopteris, all y 60. Except in fig. a and b the cell of attachment is shown, black. — 
D. ciirranii Copel. a. Hair from base of costa. — D. clemensiae Holtt. b. One of the rigid, shining 
hairs from lower surface of costa, c. thin-walled hair from costule. — D. puhlgera (Bl.) Nakai. d-e. 
Hairs from costule, lower surface. — D. linearis (Burm. f.) Und., hairs from lower surface of costules: 
/. var. subferruginea (Hieron.) Nakai, g. var. linearis, h. var. subspeciosa Holtt., /. var. alternans 

(Mett.) Holtt. 

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 Vi 

mm long) throughout 2. D. clemensiae 

4. Ultimate branches 15-25 cm long, 2.7-5 cm wide; costae with such hairs near base only. 

3. D. pubigera 

2. Lamina very thick, the veins grooved on the lower surface 3. D. pubigera 

1. Spores trilete. Branches at a {c^ m many cases unequal; ultimate branches to c. 6 cm wide. Veins 

never grooved on lower surface; tii^v '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. 

5. D. linearis 



1. Dicranopteris curranii Copel. Philip. J. Sc. 81 
(1952) 4; Holtt. Reinwardtia 4 (1957) 274.— G/e/- 
chenia hermannii var. venosa Bl. incl. also var. 
tenera Bl. En. PI. Jav. (1828) 249.— Gleichenia 
dichotoma var. tenera Mett. in Miq. Ann. Mus. 
Bot.Lugd. Bat. 1 (1863) 50.— Gleichenia dichotoma 
var. malayana Christ, Ann. Jard. Bot. Btzg 15 
(1898) 77. — Gleichenia linearis var. malayana 
v. A. V. R. Handb. (1908) 59; Suppl. (1917) 84; 
Holtt. Rev. Fl. Mai. 2 (1955) 10.— D. lessonii 
Nakai, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 
61, quoad 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. 



accessory branches at penultimate forks commonly 
20-30 cm long and 7-10 cm wide; costules 5-7 
mm apart; lamina-segments 3'/^-4'/2 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. P 



Note. Gleichenia weatherbyi Fosberg (Am. 
Fern J. 40, 1950, 140), from the Caroline Islands, 
is very near D. ciirranii, with similar spores, but 
is even larger, and much more hairy. 

2. Dlcranopteris clemensiae Holtt. 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 f^rks; 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; lamina-segments 3'/^ 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 (1-1 Vi mm long, with 
several basal branches) at first abundant all along 
lower surface of costae, at length deciduous leav- 
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 x 16/i. 

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 /;. 29 (1950) 68; Holtt. Rein- 
wardtia 4 (1957) 274. — Gleichenia hermannii var. 
pubigera Bl. En. PI. Jav. (1828) 249.— Gleichenia 
dichotoma var. pubigera Mett. in Miq. Ann. 
Mus. Bot. Lugd. Bat. 1 (1863) 50.~Gleichenia 
linearis var. pubigera v. A. v. R. Handb. Suppl. 
(1917) 85. — Mertensia spissa (non Fee) var. pu- 
bigera Nakai, Bot. Mag. Tokyo 39 (1925) ISO.— 
Fig. 12a, 14(i-e, 15a-b. 

Primary rachis-branches 2-4 times equally or 
subequally forked; ultimate branches 15-25 cm 
long, 3-8 cm wide; costules 4'/2-6 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 thinner 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, x li- — ^- speciosa (Presl) 
Holtt. c. Part of frond to show venation, x 2, 
d. main fork showing stipular leaflets, x 14. 



4. Dicranopteris speciosa (Presl) Holtt. Rein- 
wardtia 4 (1957) 273. — Hicriopteris speciosa Presl, 
Epim. Bot. (1851) 27. — Gleichenia opposita v. A. 
V. R. Bull. Jard. Bot. Btzg II, n. 11 (1913) 13; 
Holtt. Rev. Fl. Mai. 2 (1955) 70, f. 14 ¥.— Glei- 
chenia parallela RiDL. J. Mai. Br. R. As. Soc. 4 
(1926) 3.— D. opposita Nakai, Bull. Nat. Sc. Mus. 
Tokyo /;. 29 (1950) 68.— Fig. 12d, 15c-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 31/2 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; lamina-segments 3'/^-^ 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) 



33 



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 siibg. Diplopterygium and conse- 



quently used the generic name Hicriopteris in a 
wrong sense. 

5. Dicranopteris linearis (Burm. /.) Underw. 
Bull. Torr. Bot. CI. 34 (1907) 249; Holtt. Rein- 
wardtia 4 (1957) 275. — Polypodium dichutomum 
Thunb. FI. Jap. (1784) 338, t. 2>1 .—Gleichenia 
dichotoma Hook. Sp. Fil. 1 (1844) 6; Racib. 
FI. Btzg 1 (1898) 13.— Fig. 12, 14f-i. 



KEY TO THE VARIETIES 

1. Accessory branches not always present at ultimate forks. 

2. Branches at each fork of a lateral branch-system approximately equal; lateral branch-systems 
forked 1-3 times. 
3. Costules less than 5 mm apart, their lower surface with some persistent hairs. 

4. Hairs not persistent on lower surface of veins 1. 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 rigid 4. var. rigida 

6. Lamina thin 5. var. latiloba 

2. Branches at successive forks alternately unequal; lateral branch-systems on well-grown plants 

forked at least 3 times. 

7. Lower surface quite glabrous and persistently pale glaucous; 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 . 7. var. subspeciosa 

10. Angle of secondary and later forks more than a right angle ... 8. 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. ... 3. var. subferruginea 
12. Hairs pale on dried specimens, sparse on the veins; W. Malaysia . 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 commonly 2:1; costules c. 3 mm apart, segments of lamina 

21/2-3 mm wide 11. var. tetraphylla 

14. Ratio of length to width greater; costules more widely separated. 

15. Costulesonultimatebranches5-7mmapart; texture subcoriaceous; lower surface quite glabrous; 

accessory branches at ultimate forks almost opposite the fork ... 12. var. montana 

15. Costules on ultimate branches 3'/2-4'/2 mm apart; texture thin; some rusty hairs often on lower 

surface; accessory branches at ultimate forks 3-4 mm above the fork 13. var. altissima 



1. var. linearis. — Polypodium lineare Burm. /. 
FI. Ind. (1768) 235, t. 67 f. 1.— Gleichenia herman- 
niiR. Br. Prod. (1810) 161. — Gleichenia dichotoma 
var. normalis Mett. in Miq. Ann. Mus. Bot. 
Lugd. Bat. 1 (1863) 50; Racib. FI. Btzg 1 (1898) 
13. — Mertensia pteridifolia Presl, Abh. (K.) 
Bohm. Ges. Wiss. M.-N. CI. 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. 



(1917) 84, ditto; Holtt. Rev. FI. Mai. 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; lamina- 
segments 2V2-3 mm wide, separated by rather wide 
sinuses, texture very firm; lower surface slightly 



34 



Flora Malesiana 



[ser. II, vol. 1^ 



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 (Gj. 

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 
{I.e.) 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 Dieranopteris 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 \ar. 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. 

1. var. ferruginea (Bl.) Holtt. Reinwardtia 4 
(1957) nS.—Gleichenia ferruginea Bl. En. PI. 
Jav. (1828) 249; Mett. Ann. Mus. Bot. Lugd. 
Bat. 1 (1863) 50. — Gleichenia dichotoma var. 
ferruginea Racib. F1. Btzg 1 (1898) U.— Gleichenia 
linearis var. ferruginea v. A. v. R. Handb. (1908) 
59; Suppl. (1917) 85. — ''Dieranopteris 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. 2Vi mm wide above the 
base, lower surface covered with very fine floccose 
entangled pale rusty hairs (c. 10 fj, 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. subfemiginea (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 fi) 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.) Holtt. Reinwardtia 4 
(1957) 277. — Gleichenia hermannii var. rigida Bl. 
En. PI. Jav. (1828) 249.— Mertensia crassifolia 
Presl, Abh. (K.) Bohm. Ges. Wiss. M.-N. CI. 
V, 5 (1848) 339; Epim. Bot. (1851) 23, t. 13.— 
Gleichenia dichotoma var. rigida Mett. in Miq. 
Ann. Mus. Bot. Lugd. -Bat. 1 (1863) 5Q.— Glei- 
chenia warburgii Christ, Ann. Jard. Bot. Btzg 15 
(1898) 78. — Gleichenia linearis var. rigida v. A. 
v. R. Handb. (1908) 59; Suppl. (1917) 84.— Glei- 
chenia crassifolia Cqpel. 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, 3-51/2 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. Mettemus (and apparently also Raci- 
BORSKi) referred some specimens of D. pubigera 
from Java to this variety of D. linearis. 

5. var. lataoba Holtt. 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. 1 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 Dicranopreris on edge of forest, Singapore. D. curranii Copel. (large fronds) to left 
and right; D. linearis (Burm. /.) Und. var. linearis in centre; var. subpectinata (Chr.J Holtt. to right 

of centre, below (Holttum, 1925). 



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 (Christ) Holtt. Rein- 
wardtia 4 (1957) 277. — Gleichenia subpectinata 
Christ, Bot. Tidsskr. 24 (1901) \\\.— Gleichenia 
pteridifolia inon Mertensia pteridifolia Presl) 
RiDL. J. Mai. Br. R. As. Soc. 4 (1926) 4.— £). 
warburgii (non Gleichenia warburgii Christ) sensu 
Nakai, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 
70. — Gleichenia linearis var. alternans {non Mett.) 
Holtt. Rev. Fl. Mai. 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, 
2-3 '/2 cm wide (sterile ones sometimes to 18 by 
6 cm); lamina thin, lower surface pale glaucous 
and quite glabrous when mature; segments of 



lamina gradually and rather evenly tapering from 
base to apex, the sinuses thus narrowly triangular; 
costules 4-41/2 rnrn 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 Holtt. 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. 1\ Dec. 1959] 



Ecol. On edges of forest, in thickets with other 
varieties, 0-1400 m. 

8. var. inaequalis (Rosenst.) Holtt. Rein- 
wardtia 4 (1957) 278. — Gleichenia linearis var. 
inaequalis Rosenst. in Fedde, Rep. 13 (1915) 212; 
V. A. V. R. Handb. Suppl. (1917) 85. 

Shape and venation of lamina-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 an accessory 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. var. alternans (Mett.) Holtt. Reinwardtia 4 
(1957) 278. — Gleichenia dichotoma var. alternans 
Mett. in Miq. 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 Holtt. Reinwardtia 4 (1957) 
275. 

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) 61.— 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, lamina 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 Holtt. Reinwardtia 4 (1957) 
276. — Gleichenia linearis var. montana Holtt. 
Rev. Fl. Mai. 2 (1955) 69, descr. angl.— Fig. 12b. 

Lateral branch-systems several times equally 
forked; ultimate branches 15-25 cm long, 3'/2-6 
cm wide; costules (5-)6-7 mm apart; lamina 
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. linearis and with other 
members of the family; 1000-1600 m. 

13. var. altissima Holtt. Reinwardtia 4 (1957) 
276. — Gleichenia linearis var. altissima Holtt. 
Rev. Fl. Mai. 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 3'/2-4'/2 
mm apart; lamina 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 (meduUated 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). Anemiahas 
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 Klukia 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, I.e.; 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 oi 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 oi 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, 5. 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. II, vol. 1^ 




-1«».^.>IW|L^*« 









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. H, 1881). Diels doubted Prantl's 
observations (in Engier, 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, I.e. 170). Most observations of gametophytes 
have been of non- Malaysian species. 



De c. 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, I.e.); the same number was also found by Lovis (unpublished) for 5. dichoioma in New 
Zealand. In Ceylon Lovis found that 5. 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 {I.e.); later works are cited 
by Bower. The rhizome oi 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 meduUated 
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 oi 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, etc., 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, Diet. Econ. Prod. Mai. Pen. 1378, 1975; also 
Heyne, Nutt. PI. 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 L>'g^oJ/M/77) in magical ceremonies connected with house-building, 
rice culture, fishing, etc. 

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.flexuosum. In the years immediately following 
1 800, 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 MiCHX (1803) and Vallifilix Thouars (1809). 
The following names were also given to Schizaea: Lophidium Rich. (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. Diff"erent 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 
& Holttum, Reinwardtia 5 (1959) 11—22. 

The only good monograph of the whole family is that of Prantl {I.e., 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. Diels (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. 

Nakai 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. II, vol. 1^ 

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 Rich. Act. Soc. Hist. 
Nat. Paris 1 (1792) \U.—Actuwstachys Wall, ex Hook. Gen. Fil. (1842) t. Ill; 
Reed, Bol. Soc. Brot. 21 (1947) 130.— Microschizaea Reed, I.e. 133.— Fig. 1^. 
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 faUing 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. 

D i s t r. 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 oi^ Schizaea (or of a branch of 5. dicho- 
toma) as a sorophore; he called the lateral appendages laciniae. 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. Goteb. 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, 431^50). Fossil spores are also known (see remarks on 
fossils, supra). 

KEY TO THE SPECIES 

1. Fronds repeatedly dichotomous 1. S. dichotoma 

1. Fronds simple. 

2. Sorophores 2V2-5 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 in two 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 
(1793) 422, t. 9; Bl. En. PI. Jav. (1828) 255; 
Hook. & Grev. Ic. Fil. (1827) t. 17; Bedd. Ferns 
S. India (1863) t. 65; Handb. (1883) 452; 
Prantl, Unters. Morph. Gefasskr. 2 (1881) 138; 
Racib. Pterid. Buit. (1898) 6; v. A. v. R. Mai. 




Fig. 2. Schizaea dichotoma (L.) Sm. near Kepa- 
hiang, Bencoolen, S. Sumatra (De Voogd). 



Ferns (1908) 116; Merr. Int. Rumph. (1917) 69; 
Backer & Posth. Varenfl. Java (1939) 256, 
fig. 66; HoLTT. Ferns Mai. (1955) 50, fig. 6.— 
Acrostichiim dichotonium Li'N>iE,Sp. Pl.( 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.-5. forsteri Spr. Anieit. 3 (1804) 57.-5'. 
cristata Willd. Sp. PI. 5 (1810) 88.— 5. biroi 
Richter, Math. Termeszet. Ertesito 29 (1911) 



1074; Troll, Flora 128 (1933) 339, fig. 1.— S. 
copelandica Richter, I.e. — 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 1-1 '/2 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^ 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 {I.e.) 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 ijakar 
ajam, radja hantii, 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; Bl. En. PI. Jav. (1828) 255; 
Bedd. 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; Racib. Pterid. Buit. (1898) 
7; V. A. v. R. Mai. Ferns (1908) 116: Backer & 
Posth. Varenfl. Java (1939) 257, f. 66; Selling, 
Svensk. Bot. Tidskr. 41 (1947) 431-450; Holtt. 
Ferns Mai. (1955) 51, f. 7. — Aerostiehum digitatiim 
Linne, Sp. PI. (1753) \Q6^.—Aetinostachys digitata 
Wall, ex Reed, BoL Soc. Brot. 21 (1947) 130.— 
Fig. 3a-€. 



42 



Flora Malesiana 



[ser. II, vol. V 



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 2-4 mm, 




Fig. 3. Schizaea digitata (L.) Sw. a. Habit, 
X '/i> 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. /. Detail of sorophore, x 18, 
g. spore, X 300, h. flattened part of exospore, 
X 200, /. lower surface of leaf, showing stomata, 
X 5 (a, e For. Bot. Burma 7670, b-c Matthew 
s.n. (Sing),/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 2'/2-5 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 (I.e.). 
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; Holtt. Ferns Mai. 
(1955) 52. — Actinostachys inopinata Reed, Bol. 
Soc. Brot. 21 (1947) 130.— Fig. 3f-i. 

In habit like S. digitata, differing as follows: 
fronds to 2'/2 miTi 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). 1 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 on each 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 



Dec. 1959] 



Schizaeaceae (Holttum) 



43 




Fig. 4. Schizaea dichotoma (L.) Sm. a. Habit, X 2/3, b. rhizome, X 2/3, c. sorophores, X 7, (/. lower 
surface of leaf, x 14.— S. wagneri Selling, e. Habit, x Vs./- 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.-5. fistulosa Labill. j. Sorophores, x Vs. ^- detail of sorophore, x 7, /. lower surface of leaf, 
showing stomata, x 27 (a Hose 199, b Brass 8583, c-d FMS 199, e-gGRETHER & Wagner 4177, h-i 

SFN 1100, y-/ Clemens 10729). 



long, about 1 V2 mm wide, costa very slightly 
prominent on the lower surface, with a distinct 
flat wing on each side of it; stomata in one row 
on each side of the costa, rather widely spaced 
in the rows; sorophores 1-3, 4-8 mm long (some- 
times longer?), glabrous, the sporangia in two 



rows (sometimes apparently in 4 rows near the 
middle); spores as in S. digitata. 

Type: Troll, Ambon (M). 

Distr. Micronesia (Ponape); in Malaysia: 
Moluccas (Ambon). 

Ecol. Growing in moss-cushions on trees. 



44 



Flora Malesiana 



[ser. II, vol. P 



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- 
phylla but differ in costa and stomata and in more 
crowded sporangia. 

5. Schizaea wagneri Selling, Svensk Bot. Tidskr. 
40 (1946) 278, f. 8-11; Holtt. Ferns Mai. (1955) 
52. — Actinostachys wagneri Reed, Bol. Soc. Brot. 
21 (1947) n\.—S. paucijuga Holtt. Card. Bull. 
Sing. 11 (1947) 267.— Fig. 4e-g. 

Rhizome short, apex clothed with slender 
brown hairs 1 Vi mm long. Fronds simple, 6-20 
cm long, base terete, upper part winged and in 
all '/2-% 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 Is!., 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 Labile. Nov. Holl. PI. 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. Mai. Ferns (1908) 116; 
C. Chr. & Holtt. Gard. Bull. S. S. 7 (1934) 210.— 
Microschizaea fistulosa Reed, BoI. Soc. Brot. 21 
(1947) 134. — 5. propinqua A. CuNN. in Hook. 
Comp. Bot. Mag. 2 (1836) 362.-5. australis 
Gaud. Ann. Sc. Nat. Bot. 5 (1825) 98.-5. 
chilensis Phil. Linnaea 30 (1859-60) 207.— Fig. 
4j-I. 

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 1 0-20 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: Labillardiere, 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 5. papuana Brause, which is here 
placed as a synonym of 5. malaccana. 

7. Schizaea malaccana Bak. Syn. Fil. (1868) 428; 
Beddome, Ferns Br. India (1870) t. 255; Handb. 
(1883) 452; Tansley & Chick, Ann. Bot. 17 
(1903) 493-510; v. A. v. R. Mai. Ferns (1908) 
116; Holtt. Ferns Mai. (1955) 52, fig. 8.-5. 
fistulosa var. malaccana Prantl, Unters. Morph. 
Gefasskr. 2 (1881) \36.—S. papuana Brause, Bot. 
Jahrb. 56 (1920) 211. — Microschizaea malaccana 
Reed, BoI. Soc. Brot. 21 (1947) 134.— Fig. 4h-i. 

var. malaccana. 

Habit of 5. fistulosa, differing as follows: hairs 
on rhizome pale hrown, fronds 6-15 cm long, less 
than 1 mm wide, apical fertile part of axis c. 5 mm 
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. Chr. Gard. Bull. S.S. 7 (1934) 
210.— 5. 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 etc.— 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, iiom. cons.; Prantl, 
Unters. Morph. Gefasskr. 2 (1881) 60; Copeland, Gen. Fil. (1947) 23; Pichi- 
Sermolli, Webbia 12 (1956) 10, preprint (\9 55). —Ramondia Mirbel, Bull. 
Soc. Philom. Paris 2 (Feb.-Mar. 1801) 179.— Ugena Cavan. Ic. Descr. PI. 6 (Oct. 
1801) 13.—Odontoptens Bernh. in Schrader, J. Bot. 1800 pt 2 (Nov.-Dec. 1801) 
127, t. 2 f. 4.—Ripidium Bernh. I.e. 127, t. 2 f. 3.—Gisopteris Bernh. I.e. 129, 



Dec. 1959] Schizaeaceae (Holttum) 45 

t. 2 f. 1. — Hvdroglossum Willd. Abh. Kurfurstl. Mainz. Ak. Niitzl. Wiss. Erfurt 
2, pt 4 (1802) 13, 20.— Hugona Cav. ex Roemer, Arch. Bot. 2 (1801-02) 486.— 
Cteisium MiCHX, Fl. Bor. Am. 2 (1803) llS.— Vallifilix Thouars, Gen. Nov. 
Madag. (1808) \.—Lvgodictvon J. Smith in Hook. Gen. Fil. (1842) t. 1 1 1 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 sUghtly raised and papillose ; 
all branch rachises and stalks of leaflets similarly winged (fig. 13d), the vdngs always 
interrupted to join with those of a lateral hiQ-noh: 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 leaflets (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. 

Dislr. 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 
ape.x 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 Lygodiiim 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. 

Tax on. 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. II, vol. 1^ 

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. salicifohum; L. borneense and L. auriciilatum; L. dimorphum and L. trifurcatum. The following 
species are very distinct: L. polystacnyum, L. micropftyllum, L. circinnatum, L. merrillii, andL. 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 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. 

2. L. microphyllum 

4. Secondary branches amply bipinnate. Leaflets not articulate at the base 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; leaflet-stalks thickened at 
their junction with the lamina 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 spreading 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 lower 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. longifoiium 

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 11. L. circinnatum 

2. Venation of sterile leaflets reticulate. 

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) . 12. L. merrillii 

13. Leaflets on much shorter stalks, simple or the apical ones forked . . 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 13. L. versteegii 

l.Lygodiumpolystachyum Wall. PA Moore, Gard. v. A. v. R. Mai. Ferns (1908) 113; Copel. Philip. 

Chron. (1859) 671; Hook. Sec. Cent. (1861) t. 76; J. Sc. Bot. 4 (1909) 19; Tard. & C. Chr. F1. 

Syn. Fil. (1868) 438; Beddome, Ferns Br. G6n. I.-C. 7 (1939) 40; Holtt. Ferns Mai. (1955) 

India (1868) t. 300; Handb. (1883) 458, f. 284; 56, f. 10; Alston & Holtt. Reinwardtia 5 (1959) 



Dec. 1959] 



Schizaeaceae (Holttum) 



47 



11. — Hvdroglossum pituuitifidum Willd. Abh. 
Kurt". Mainz. Ak. Wiss. Erfurt 2, pt 4 (1802) 21, 
p.p. — L. pinnatifidum {nun (Willd.) Sw.) Prantl, 
Unters. Morph. Gefiisskr. 2 (1881) 83, t. 1 
f. 11.— Fig. 5c, 8a-c. 






^g 



I 



CSC?, 







Fig. 5. Spores of Lygodium. a. L. longifolium 
(Willd.) Sw., b. L. bomeense v. A. v. R., c. L. 
polystachyum Wall, ex Moore, d. L. circinnatum 
(Burm. /.) Sw., e-f. L. microplnllum (Cav.) 
R. Br., lateral and upper view. All ■ 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, with short very slender hairs mixed with thick 
longer multicellular ones; ultimate branches leafy 
like the secondary rachis-branches of climbing 
fronds. Roc/iis of climbing fronds 2'/2 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 
3'/2-7'/2 cm long, 1 '/2-2 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 
stifl" 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 sol 
limestone provides a drier habitat than oti 
rocks, and on it occur a number of species whii. 
have their main distribution in the seasonal 
climate north of Malaya.) 

2. Lygodium microphyllum (Cav.) R. Br. Prod. 
Fl. Nov. Holl. (1810) 162; Bl. En. PI. Jav. (1828) 
253; Clarke, Trans. Linn. Soc. Bot. 1 (1880) 583; 
Bedd. Handb. (1883) 455, t. 282; Alston & 
HoLTT. Reinwardtia 5 (1959) 12. — Ugena micro- 
phylla Cav. Ic. Descr. PI. 6 (1801) 76, t. 595; 
C. Chr. 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; Hook. Syn. Fil. (1868) 437, p.p.; 
Prantl, Unters. Morph. Gefasskr. 2 (1881) 81, 
t. 6 f. 101; Christ, Farnkr. Erde (1897) 354, f. 
1116; Racib. Pterid. Buit. (1898) 8; v. A. v. R. 
Mai. Ferns (1908) 113; Philip. J. Sc. 11 (1916) 
Bot. 116; Heyne, Nutt. PI. (1927) 97; Back. 
Krakatoa (1929) 254; Ochse & Bakh. Veg. 
D.E.I. (1931) 657; Burk. Diet. 2 (1935) 1378; 
Backer & Posth. Varenfl. Java (1939) 258; 
Ogata, Ic. Fil. Jap. 7 (1936) t. 324; Holtt. 
Ferns Mai. (1955) 58, f. \2.—Ophioglossum 
filiforme Roxb. Calc. J. Nat. Hist. 4 (1844) 476, 
t. 26 f. 3. — L. scandens var. microphyllum (Cav.) 
LuERSS. J. Mus. Godefl'r. 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, 21/2 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). 
Racliis of climbing fronds glabrous, commonly 
2-3 m long, hardly 1 V2 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. II, vol. P 




Fig. 6. Lygodium microphyllum (Cav.) R. Br. and L. salicifolium Pr. (larger leaflets on left) on edge of a 
thicket in the Botanic Gardens, Singapore (Holttum). 



Dec. 1959] 



Schizaeaceae (Holttum) 



49 









Fig. 7. Lygodium microphyllum (Cav.) R. Br. a. Fertile leaf, x 3/4, 6. sterile leaf, X 3/4, c. rhizome, 
X 3/4, rf. 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 1-A mm long) and a similar or geminate 
terminal leaflet; leaflets 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 with a raised reticulum 
on the outer surface. 

Type: N6e, 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. P 













~<\ 




"^ 



a ^#^^ 




Fig. 8. Lygodium polystachyum Wall, ex Moore, a. Sterile leaf, x ^/j, 6. fertile leaflet, x ^1^, c. ditto, 

detail x 21/2- — L.japonicum (Thunb.; Sw. rf. Fertile leaf, x 3/^, g. sterile leaf, x ^j-^,f. rhizome, x ^/^ 

(a Balansa 168, b-c Matthew s.n., J, / 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. Rihu-ribu, M, selada, capey papua, capay 
aim, M, paku kawcir. Sum., paku liata bejas (bias), 
paku hata leutik, S, nitung-puli. Tag., paku rambat, 
Bali, sickey, Luhea, gomuha 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 j^ponicum (Thunb.) Sw. in Schrader, 
J. Bot. 1800, pt 2 (1801) 106; Bedd. Ferns S. India 
(1863) 21, t. 64; Clarke, Trans. Linn. Soc. Bot. 1 
(1880) 584; Prantl, Unters. Morph. Gefiisskr. 
2 (1881) 68, t. I f. 10, 15; Christ, Farnkr. Erde 

(1897) 355, 356, f. 1122; Ractb. Pterid. Buit. 

(1898) 8; Hope. J. Bomb. Nat. Hist. Soc. 15 
(1903) 106; V. A. v. R. Mai. Ferns (1908) 114; 
.Merr. F1. Manila (1912) 60; Domin, Bibl. Bot. 20, 
Heft 85(1914) 211, f. 50; Haines, Bot. Bih. & Or. 
6 (1924) 1210; Ogata, Ic. Fil. Jap. 7 (1936) t. 
322; Backer & Posth. Varenfl. Java (1939) 259; 
Alston & Holtt. Reinwardtia 5 (1959) 14. — 
Ophioglossum japonicum Thunb. F1. Jap. (1784) 
328. — Hvdroglossum japonicum (Thunb.) Willd. 
Abh. Kurf. Mainz. Ak. Wiss. Erfurt 2. pt 4 (1802) 
26. — L. dissect um Desv. Mag. Ges. Naturf. Fr. 
Berl. 5 (1811) 308. — L. microstachyum Desv. I.e.; 
Nakai, Bot. Mag. Tokyo 39 (1925) 182; Ogata, 
Ic. Fil. Jap. 7 (1936) t. 323.— L. pubeseens Kaulf. 
En. Fil. (1824) 47, t. 1 f. A.—L. ehaerophylloides 
Desv. Mem. Soc. Linn. Paris 6 (1827) 205.— L. 
cochinehinense Desv. ibid. 206. — L. tenue Bl. 
En. PI. Jav. (1828) 254.— L. microphyllum Link, 
Hort. Berol. 2 (1833) 141.— L. japonicum f. 
elongata v. A. v. R. Bull. Jard. Bot. Btzg 11, n. 1 
(1911) 10, t. 3; Mai. Ferns Suppl. (1917) 117.— 
L. japonicum var. microstaehya (Desv.) Tard. & 
C. Chr. F1. Gen. 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 leaflets 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 Isl.. 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 kembang, J, paku areuj, hata 
kawat, S, nito, nitong puti. Tag., madik silai, 
durhawa, babar, talsiga, Alor. 

4. Lygodium salicifolium Presl, Suppl. Pterid. 
(1845) 102, p.p. excl. pi. Wallich p.p. and syn. 
Rheede; Prantl, Unters. Morph. Gefiisskr. 2 
(1881) 79; V. A. v. R. Mai. Ferns (1908) 113; 
Bull. Jard. Bot. Btzg III, 5 (1922) 213; Backer 
& Posth. Varenfl. Java (1939) 258; Tard. &. C. 
Chr. FI. Gen. I.-C. 7, 2 (1939) 41; Alston & 
Holtt. Reinwardtia 5 (1959) 14.— L. kingii 
CoPEL. Philip. J. Sc. 6 (1911) Bot. 68; v. A. v. R. 
Mai. Ferns Suppl. (1917) 117. — L. pinnatifidum 
sensu Rac. FI. Buit. (1898) 7, p.p.— L. flexuosum 
sensK Holtt. Ferns Mai. (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 - 1 5 cm long, '/2-2cm wide, acute 



52 



Flora Malesiana 



[ser. II, vol. P 




Fig. 9. Lygodium dimorphum Copel. a. Part of fertile leaf, X %, b-c. parts of sterile leaves, X %, 

d. detail of a, x 3.—L. flexuosum (L.) Sw. ^. Sterile leaf, x %,/. fertile leaf, x % (a Labillardiere 

s.n., b-c Carr 12479, ^ Peekel 6, e Hose 5024, / Hallier s.n. from Java). 



Dec. 1959] 



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; Presl, Suppl. Pterid. (1845) 100 
Bedd. Ferns S. India (1863) t. 63; Clarke 
Trans. Linn. Soc. Bot. 1 (1880) 584; Prantl 
Unters. Morph. Gefasskr. 2 (1881) 72, p.p. 
Bedd. Handb. (1883) 457, f. 283; v. A. v. R. Mai 
Ferns (1908) 114; Merr. F1. 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. Diet. 2 (1935) 
1378; C. Chr. Dansk Bot. Ark. 9, pt 3 (1937) 
30; Backer & Posth. Varenfl. Java (1939) 259; 
HoLTT. Ferns Mai. (1955) 57, p.p.; Alston & 
HOLTT. Reinvvardtia 5 (1959) 15. — Ophioglossum 
flexuosum Linne, Sp. PI. (1753) 1061,.— Ophio- 
glossum scandens Linne, Sp. PI. (1753) 1063, p.p. 
— Ramondia flexuosa (L.) MiRB. Bull. Soc. Philom. 
Paris 2 (Feb.-Mar. 1800) 179, t. 12 f. 3.—Hydro- 
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. 
pimuitifidum Sw. in Schrader, J. Bot. 1801, pt 2 

(1803) 303; Hook. Syn. Fil. (1868) 438, p.p.— 
L. semibipimiatum R. Br. Prod. Fl. Nov. HoII. 
(1810) 162.— L. serrulatum Bl. En. PI. Jav. 
(1828) 254. — L. flexuosum var. setulosum Tard. & 
C. Chr. Fl. Gen. 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. II, vol. V 



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 besar, ikat 
sidin, M, paku ribu-ribu, Asahan, hata kembang, 
J, durhawa, Alor, nito. Tag., Vise, tatan, Orokawa 
Horata (N.G.), zangi, Orokawa Mumuni. 

Uses. For tying rice sheaves; in native medicine 
for skin diseases and fever. 

6. Lygodium dimorphum Copel. Philip. J. Sc. 6 

(1911) Bot. 67; Rosenst. in Fedde, Rep. 10 

(1912) 343; v. A. v. R. Mai. Ferns Suppl. (1917) 
116; Philip. J. Sc. 11 (1916) Bot. 116; Alston 
& HoLTT. Reinwardtia 5 (1959) 18.— L. flexuo- 
sum [non (L.) Sw.] Gaudich. in Freyc. Voy. Bot. 
(1826) 298. — L. circinnatum var. trifurcatum 
Christ, Monsunia 1 (1900) 93. — L. novoguineense 
Rosenst. in Fedde, Rep. 9 (1911) 427.— L. 
trifurcatum sensu v. A. v. R. Mai. Ferns (1908) 
112, 802, 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 (Mich, Bo). 

Distr. A/a/(7>'5/a.- 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, Moluan, 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. Mai. Ferns (1908) 112, 
802, p.p.; Wagner & Grether, Un. Cal. Publ. 
Bot. 23 (1948) 27, t. 8; Alston & Holtt. Rein- 
wardtia 5 (1959) 17. 

Scandentrflf/!« about 1 '/2 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, \V2-IV2 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. PI. 6 (1801) 74, t. 594, nomen. illegit., 
excl. svn. Reichard & Rumph.; C. B. Rob. 
Philip. J. Sc. 6 (1911) Bot. 97; C. Chr. Dansk 
Bot. Ark. 9, pt. 3 (1937) 29.— Hydroglossum 
auriculatum Willd. Sp. PI. 5 (1810) 84. — L. semi- 
hastatum Desv. M6m. Soc. Linn. Paris 6 (1827) 
203, nom. illeg.; Hook. Syn. Fil. (1868) 437; 
V. A. v. R. Mai. Ferns (1908) 111, excl. pi. bor- 
neensis; Mai. Ferns Suppl. (1917) 115; C. Chr. 
Ind. Fil. Suppl. I (1913) 119.— L. circinnatum 
var. semihastatum Fosb. 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) 



55 




^'"'^'^^p/^ifmmmmtf^^^^^ 



Fig. 11. Lygodium auricidatum (Willd.) Alston, a. Habit, x 2 3, 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 commonlyoncedichotomous, 
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. longifoliiim, in others 
with many smaller warts of variable size. 

Type: Nee, 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 Nee 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 II, n. 20 (1915) 29; Mai. Ferns Suppl. 
(1917) 115; CoPEL. Sarawak Mus. J. 2 (1917) 
303; HoLTT. J. R. As. Soc. Mai. Br. 6 (1928) 16 
with fig.; Ferns Mai. (1955) 56.— L. semi- 
hastatum sensu v. A. v. R. Mai. Ferns (1908) 111, 
p.p. quoad pi. borneenses. — L. borneense f. 
samarindae v. A. v. R. Bull. Jard. Bot. Btzg II, 
n. 20 (1915) 29; Mai. 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. II, vol. 1^ 







"^^ ^^^^ 










l«S 



.••::^ 



.'^ 



><i^^ 



>s^i^'' 






^ff/riy^.fm^ 



Fig. 12. Lygodium longifolium (Willd.) Sw. «. Habit, fertile, X ^/j, 6. sterile, x 2/3, c. leaf detail showing 
edge, ;■ 3 (a King's coll. 259, b Motley s.n., c Hose 5034). 



Dec. 1959] 



Schizaeaceae (Holttum) 



57 




Fig. 13. Lygodium salicifolium Pr. a. Habit, x ^j^, b. detail, x 10. — L. borneense v. A. v. R. c. Habit, 
X 2/3, d. detail of branching, x 2V2, e. ditto, x *l^ {a 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), 31/2-5 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), surophores 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 Imperata on 
sandy ground, has unusually small leaflets. 

Note. This species has a branching habit closely 
similar to that of L. auriculatiim. L. borneense 
usually diff"ers 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. II, vol. 1^ 




Fig. 14. Lygodium circinnatum (Burm. /.) S\v. a. Part of sterile leaf, x -'-i, b-c. parts of fertile leaves, 

showing different kinds of branching, x -/-i, 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] 



Schizaeaceae (Holttum) 



59 



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 & 
HoLTT. Reinwardtia 5 (1959) 19. — Hydroglossum 
longifolium Willd. 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. Mai. 
Ferns (1908) 112; Holtt. Ferns Mai. (1955) 55.— 
L. dichotomum {non (Cav.) Sw.) Bedd. Ferns S. 
India (1863) t. 62. — L. leysmannii v. A. v. R. Bull. 
D6p. Agr. Ind. Neerl. 18 (1908) 5; Mai. 
Ferns (1908) 111, 801; Mai. Ferns Suppl. 
(1917) 115. — L. circinnatum var. cristatum v. A. 
V. R. Bull. Dep. Agr. Ind. Neerl. w. 18 (1908) 5; 
Mai. 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 io 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-A 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 1-3 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 (BuRM. /.) Sw. Syn. 
Fil. (1806) 153; Bl. En. PI. Jav. (1828) 253; 
Prantl, Unters. Morph. Gefasskr. 2 (1881) 64; 
Bedd. Handb. (1883) 455; v. A. v. R. Mai. 
Ferns (1908) 111; C. B. Rob. 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. Mai. Ferns Suppl. (1917) 115; W. H. Brown, 
Bull. Bur. For. Philip, n. 19 (1919) t. 4; ibid. 



n. 22 (1920) 328, pi. IV; Heyne, Nutt. PI. (1927) 
96; Back. Krakatoa (1929) 253; Burk. Diet. 2 
(1935) 1378; Backer & Posth. Varenfl. Java 
(1939) 258, f. 67; Holtt. Ferns Mai. (1955)55, 
f. 9; Alston & Holtt. Reinwardtia 5 (1959) 20. 
— Ophioglossum circinnatum Burm. /. Fl. Ind. 
(1768) 228. — Ophioglossum pedatum Burm. /, 
ibid. 227, t. 66 f. 1. — Ugena dichotoma Cav. Ic. 
Descr. PI. 6 (1801) 74, t. 594 f. 2; C. Chr. Dansk 
Bot. Ark. 9, pt 3 (1937) 30.— L. pedatum (Burm. 
/.) Sw. Syn. Fil. (1806) 154; Merr. Philip. J. Sc. 
19 (1921) 336. — Hydroglossum circinnatum (Burm. 
/.) Willd. Abh. Kurf. Mainz. Ak. Wiss. Erfurt 
2, pt 4 (1802) 24. — Hydroglossum pedatum 
(Burm. /.) Willd. ibid. 25. — L. dichotomum 
(Cav.) Sw. Syn. Fil. (1806) 154; Hook. & Grev. 
Ic. Fil. 1 (1831) t. 55; Hook. & Bak. Syn. Fil. 
(1868) 437; Racib. Pterid. Buit. (1898) %.— Ophio- 
glossum furcatum RoxB. Calc. J. Nat. Hist. 4 
(1844) 478.— L. basilanicum Christ, Philip. J. Sc. 
2 (1907) Bot. 179; v. A. v. R. Mai. Ferns (1908) 
802. — L. circinnatum var. monstruosum v. A. v. R. 
Bull. Dep. Agr. Ind. Neerl. n. 18 (1908) 5; Mai. 
Ferns (1908) 112; Mai. 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'/2 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 3-5-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. II, vol. P 




Fig. 15. Lygodium merrillii CoPEL. a. Part of fertile leaf, X 1/2, b. part of sterile leaf, x '/2. c. detail 
of fertile leaf, x U^.—L. versteegii Christ, d. Part of fertile leaf, X 1/2. e. part of sterile leaf, x 1/2, 
/. detail, main rachis bearing a primary branch with its secondary branches, each ultimate branch 
bearing a single leaflet like those shown in d and e, x 2 (a, c Mabesa 26112, 6 Topping 1318, d-f Lam 

1386). 



Dec. 1959] 



Schizaeaceae (Holttum) 



61 



altitude, never where the ground becomes sea- 
sonally very dry. 

Vern. Ribu-ribu dudok, rihu-ribu bukit, paku 
jari nierali, akar sidin, kapai besar, reribu, M, 
tura, Nias, kapai gorita, Mol., paku hata, hata 
areuj, S, paku ata, paku ribu-ribu, Asahan, paku 
rambat, J, ata, Bali, babar, Alor, masem, Minah., 
raga-raga, Mak., tjiwang, Bug., mongodo, Tob., 
gomoho. Tern., gomongo, Tidore, tjito. Tag., Vise. 

Uses. Medicinal, and for plaiting. In the 
Philippines used for making hats and cigarette 
cases, in Ambon for adorning houses for marriage 
festivals, in Sumatra in rice ceremonies. Young 
leaves edible. 

12. Lygodium merrillii Copel. Philip. J. Sc. 2 
(Apr. 1907) Bot. 146, t. 4; ibid. 4 (1909) Bot. 
20, t. 12; V. A. V. R. Mai. Ferns (1908) 803; 
Mai. Ferns Suppl. (1917) 118.— L. matthewii 
Copel. Philip. J. Sc. 3 (1908) Bot. 36; ibid. 4 
(1909) Bot. 20; v. A. v. R. Mai. Ferns (1908) 
803.— Fig. 15a-c. 

Rhizome and juvenile fronds not seen. Rachis 
of scandent fronds to 5 mm diameter, minutely 
hairy (hairs slender and erect), near base also 
bearing long dark hairs like those on dormant 
apices; primary rachis-branches very short, 
dormant apices prominent, covered with long 
dark brown hairs; secondary rachis-branches 
unifoliate and sterile near the base of a frond, 
upper ones pinnate; unifoliate secondary branches 
6-9 cm long, leaflets c. 25 cm long, palmately 
5-6-lobed with acute sinuses to within 6 cm of the 
base, lobes 2Vi-4 cm wide, acute and acuminate, 
shallowly and irregularly crenate-serrate, margin 
not thickened, veins oblique, anastomosing, with 
about four rows of elongate areoles between costa 
and margin, lower surface of lamina and of veins 
minutely hairy or glabrous; largest upper sterile 
secondary rachis-branches pinnate, with 2-4 
lateral deeply bilobed or palmate leaflets (on stalks 
to 3 cm long) and a terminal one; fertile secondary 
rachis-branches pinnate (or the largest bipinnate 
at the base), in all 30 cm or more long, with 5-7 
leaflets which are 2-4-lobed; fertile leaflets 8-10 
cm long, lamina of each lobe to 1 1/2 cm wide, 
veins anastomosing, stalks 5-15 mm long; soro- 
phores 7-15 mm long, somewhat contracted at 
base, indusia glabrous or with few pale hairs; 
spores very coarsely and irregularly verrucose. 

Type : Merrill 6057, Mt Halcon, Mindoro (Mich). 

Distr. Tonkin and Kweichow; in Malaysia: 
Sumatra, Sarawak, Philippines (southern Luzon, 



Leyte and Mindoro). 

Ecol. In forest, to 600 m altitude. 

13. Lygodium versteegii Christ, Nova Guinea 8 
(1909) 161; V. A. v. R. Bull. Jard. Bot. Btzg II, 
n.\ (1911) 10;? Copel. Philip. J. Sc. 6 (1911) 
Bot. 68; ? ibid. 11 (1916) Bot. 41; v. A. v. R. 
Mai. Ferns Suppl. (1917) 118, 499; Alston & 
HOLTT. Reinwardtia 5 (1959) 22.— L. mosz- 
kowskii Brause, Bot. Jahrb. 49 (1912) 57; v. A. 
V. R. Mai. Ferns Suppl. (1917) 116.— Fig. 15d-f. 

Rhizome and juvenile fronds not seen. Rachis 
of scandent frond to 2 1/2 mm diameter, minutely 
hairy (hairs very slender, erect); primary rachis- 
branches very short, the sunken dormant apex 
bearing pale hairs; secondary rachis-branches also 
very short (1-3 mm long), so that the leaflets 
appear to be verticillate on the main rachis; 
leaflets 3 or 4 on each side of a dormant apex; 
sterile leaflets c. 20 cm long, 2-2 1/2 cm wide, 
entire (margin more or less thickened and car- 
tilaginous), gradually narrowed to the narrow 
truncate base, sometimes with one or two minute 
auricles forming separate small lateral leaflets, 
lamina covered with a close network of conspicuous 
anastomosing veins, with about 5 rows of areoles 
between costa and margin (outer areoles progres- 
sively smaller), ultimate free veins joining the 
thickened margin, surface glabrous with scattered 
warts when dry; fertile leaflets often longer than 
sterile (to 30 cm long) but narrower, usually 
reduced to a narrowly winged costa with a row of 
sorophores on each side, less commonly to a 
lamina 8 mm wide in which there is slight anasto- 
mosis of veins ; sorophores 3-4 mm long, contracted 
at base; indusia more or less hairy; spores coarsely 
verrucose. 

Type: Versteeg 1400, Noord River, W. New 
Guinea (P, Bo, G, U). 

Distr. Malaysia: New Guinea. 

Ecol. In open places in forest at altitudes up 
to 1200 m, climbing to a height of 8 m; also 
reported as an epiphyte in moss on a tall tree. 

14. Lygodium reticulatum Schkuhr, Farnkr. 
(1809) 139, t. 139; Kuhn in Forschungsr. S. M. S. 
Gazelle 4 Bot., pt 6 (1889) 14; Brause, Bot. Jahrb. 
56 (1920) 212. 

Similar to L. microphyllum in habit and in 
spores, dilTering: venation reticulate, lamina 
firmer, the leaflets usually more elongate. 

Distr. Queensland, Fiji to Tahiti, New Hebri- 
des, New Caledonia; reported by Kuhn (Ac.) from 
New Ireland but without citation of a specimen, 
not otherwise known in Malaysia. 
Excluded 
Cheilanthes fuscata Bl. En. PI. Jav. 2 (1828) 136 = Mohria caffrorum (L.) Desv. The type of this 
plant (L) was identified by Rosenstock and later verified by Posthumus. As Backer & Posthumus 
have pointed out (Varenflora Java p. 144, footnote) various ferns from S. Africa and Macaronesia 
have been described by Blume as native in Malaysia, e.g. Blechnum punctulatum Sw., Pellaea pteroides 
Prantl, Cheilanthes hirta Sw., Ch. multifida Sw., Asplenium adiantum-nigrum L., Hemitelia capensis 
R. Br., Todea barbara Moore. Really they had been collected during short stays en route to Java via 
the Cape of Good Hope and were later mixed up with collections made in Java. It is not improbable 
that they were collected by Kuhl & Van Hasselt as the herbarium materials collected by these young 
men and ardent collectors, who unfortunately fell untimely victims of tropical diseases in West Java, 
came into the hands of Blume. On the label there is no indication that the specimen originated from 
the Moluccas, as stated by Blume. 



62 



Flora Malesiana 





i^^MLMl . 




.r^', 




'^n^^S^^' 



/ 



Fig. I. Isoetes habbemensis Alston, tufted in marginal shallows of Habbema Lake, c. 3225 m, with 
Libocedrus papuana F. v. M., on ridge in background (Archbold Expeditions, Brass, 1938). 



ISOETACEAE (A. H. G. Alston ^ London) 

1. ISOETES 

LiNNE, Sp. PI. (1753) 1100; Gen. PI. ed. 5 (1754) 486.— Fig. 1. 

Herbaceous, perennial, submerged aquatics or marsh plants, usually with 
annual grass-like leaves arising in a tuft from a lobed, flattened, corm-like stock. 
Stock divided into stem and rhizophore, 2-4-lobed, with black dichotomous 
roots arising from the furrows between two lobes. Roots monarch, with the 
stele attached to one side of a central cavity, vascular system protostelic, 2-A- 
lobed at base. Leaves distichous, crowded, with overlapping bases, terete or 
flattened above, with a broad spoon-like base. Blades with a simple trace and 
median, unbranched vein, accessory peripheral strands often present; mesophyll 
chambered with four longitudinal cavities di\ided by transverse diaphragms, 
which give the leaf a muriform appearance when seen in transmitted light. Stomata 
present on one or both surfaces in some species and absent in others. Leaf-bases 
usually membranaceous and hyaline but in some species persistent as hard, 
brown. 2-lobed. horny structures. Ligule present near the base of the leaf above 
the sporangium, arising from a cavity called the ligular pit, cordate-triangular 
or subulate, 2-15 mm long, without chlorophyll or cuticle, secreting mucilage at 
least when young. All leaves potentially sporophyll with a sporangium seated in 
a pit (fovea) on the adaxial surface below the ligule. Megasporophylls normally 
arising below the microsporophylls; opening of fovea often wholly or partly 
covered by a membrane (velum) extending downwards from the apex. Sporangia 
large, 4-7 mm long, oblong, thin-walled (walls w ith 3-4 layers of cells), subdivided 
irregularly and incompletely by oblique sterile plates (trabeculae); of two kinds, 
megasporangia and microsporangia, sessile and broadly adnate. Sporangia with 
both megaspores and microspores have been reported and the megaspores often 
vary considerably in size. Megasporangium containing 50-300 trilete spores, 
250-900 // in diam., white, grey or black, smooth or with warts, spines, or ridges. 
Microspores monolete, elliptic, 20^5 u long, smooth or papillose, 150.000- 
1.000.000 in each sporangium. Annulus wanting, spores released by the decay of 
sporangial walls. Some species may be aposporous with young plants taking 
the place of the developing sporangia. Gametophytes dioecious. Female prothallus 
green, development starting within and the prothallus remaining attached to the 
wall of the megaspore. Archegonia one or more up to 30, deeply sunken. Rhizoids 
present, projecting beyond the spore wall. Male gametophyte arising within the 
microspore, consisting of only a single prothallial cell and an antheridium, with 
4 peripheral cells and 4 central cells, each giving rise to a single antherozoid with 
15 flagellae. 

Distribution. About 75 spp., in all parts of the world except the Pacific Islands (present in Tasmania 
and New Zealand), but mainly temperate, scarce in Asia, in Malaysia 3 spp., one in the hills and two 
in the high mountains. 

Ecology. The Malaysian species are submerged aquatics in hill and mountain lakes or streams. 
Outside Malaysia tropical species may also occur in temporary pools, rice-fields, and on damp ground 
at low altitude. The spores are sometimes dispersed by being carried by detached floating sporophylls; 
also earthworms have been reported as dispersing both megaspores and microspores in their excreta 
(DuTHiE, Ann. Bot. 43, 1929, 411-412). 

Morphology and taxonomy. The most recent comparative survey of the family Isoetaceae, 
regarded as sole family of an order Isoetales, is by Rauh & Falk (Sitz. Ber. Heidelb. Ak. Wiss. 1959, 
I, 1959, 3-160). Three genera are included: Isoetes (worldwide), Stylites (Peru), and Nathorstiana (fossil 



64 



Flora Malesiana [ser. II, vol. 1\ Dec. 1959] 



of lower Cretaceous, Germany). Stylites, with its elongate stem and unbranched roots, shows some 
resemblances to Nathorstiana, which has been considered to be a possible link between Isoetes and 
the Triassic fossil Pleuromeia. Isoetes has a very short stem of complex structure, and is generally regarded 
as reduced and specialized. The fossil gsnus Isoetites, differing from Isuetes in the relative size of megaspo- 
res and microspores, in the shape of leaf-tips and in having an unlobed stem, has been most recently 
discussed by R. W. Brown (J. Wash. Acad. Sci. 29, 1939, 261-269) who described two species from 
N. America, ranging from lower Cretaceous to early Tertiary; he considers that a Portuguese fossil 
from the lower Cretaceous probably belongs to the same genus, though the specimens are imperfect. 

There has been no recent monograph of the whole genus Isoetes, and estimates of the number of 
species \ ary. C. F. Reed has published a very full list of names, with bibliography, in Bol. Soc. Brot. 
27 (1953) 5-72. 

Cytology. Manton records chromosome numbers as follows for Isoetes: I.hystrix Durieu, n = 10; 
/. lacustris L., n = 54-56 (Problems of Cytology and Evolution in the Pteridophyta, 1950, 254-259). 
Rauh & Falk {I.e.) record 2n = c. 50 for Stylites gemmifera W. Rauh. 

Uses. Leaves of /. pliilippinensis are said to be eaten. 



KEY TO THE SPECIES 

1. Megaspores smooth on inner surfaces. 

2. Leaves up to 50 cm. Stock 3-4-lobed. Microspores minutely scabrous 
2. Leaves up to 14 cm. Stock 2-lobed. Microspores densely spinulose 



1. L philippinensis 
2. I. habbemensis 



1. Megaspores warted on inner surfaces. Stock 3^-lobed. Leaves up to 7,5 cm 3. I. neoguineensis 



1. Isoetes philippinensis Merrill & Perry, Am 
Fern Journ. 30 n940) 19, fig. 

Submerged aquatic. Stock apparently 3-4 
lobed. Leaves numerous, elongate, up to 50 cm 
slender, rather flaccid, 3 mm broad in the middle 
c. 1 mm broad at base, with membranaceous 
wings. Accessory peripheral strands wanting 
Foveae 3-4 cm long, narrow, 1 14-2 mm broad 
gradually narrowed upwards, with hyaline 
margins. Velum none. Ligule elongate, ovate 
triangular. Sporangia oblong, c. 9 by 3 mm, pale 
Megaspores above 420 ^ in diam. with a pro- 
minent triradiate marking, usually smooth on the 
inner surfaces, sometimes sparingly and minu- 
tely rugose; reticulate on the outer surface. 
Microspores 25-30 by c. 22 ^, very minutely 
scabrous. 

Distr. A/a/av.s/a.- Philippines (Mindanao :Lanao 
Prov. near Momungan, vicinity of Olangu). 

Ecol. Bottom of a stream, 400-500 m, once 
found. 

Vern. Kabatiingbauing, Lanao. 

2. Isoetes habbemensis Alston, J. Arn. Arb. 26 
(1945) 180.— Fig. 1. 

Submerged aquatic. Stock apparently 2-lobed, 
appressed-semiglobose c. 3'/4 by 1 V2 cm across, 
1 cm high, with numerous short, brownish-black 
roots 2 mm in diam. arising from the lower 
surface. Leaves numerous, up to 14 cm, stout, 
more or less recurved, c. 3 mm broad in the centre, 
semicircular in transverse section, rounded on the 
back and flattened above; central vascular strand 
rather prominent; margins slightly winged. Upper 
part of the leaves green, apices caducous. Lower 
part of leaves c. 3 cm long, pale reddish-brown, 



up to 1 cm broad and winged at base. Stomata 
none. Ligule deltoid. Velum none. Sporangia 
obovate-oblong, c. 1 cm by 4 mm, pale brown. 
Megaspores c. 575 n in diam., almost smooth, 
with a prominent triradiate marking, pale greyish- 
white when dry. Microspores c. 43 fx long, densely 
spinulose, brown when dry. 
Distr. Malaysia: West New Guinea (Mt Wilhel- 
mina; Lake Habbema), twice found. 

Ecol. Abundant in marginal shallows of Lake 
Habbema and also on Mt Wilhelmina, 3225- 
3660 m. 



3. Isoetes neoguineensis Baker [ex F. v. M» 
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. Card. 9(1922) 211. 

Submerged aquatic. Stock 3^-lobed. Leaves 
numerous, 5-71/2 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 ju 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) 

Caudcx 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. 3 If, 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 Culcita 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 {siibg. Culcita in Azores and tropical America; subg. Calochlaena in Malaysia and 
Australasia; in all c. 1 spp.); Cibotium (SE. Asia, Malaysia, Hawaii, Central America, c. \2spp.); 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 Kylikiptehs. Owing to the fragmentary nature of the fossils it is very diflficult 
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 Cror/zea-ancestor. Though no fossils with 
Cyathea-Uke 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) 



66 



Flora Malesiana 



[ser. II, vol. 12 




Fig. 1. Cyathea gleichenioides C. Chr. and C. atrox C. Chr. growing on steep slopes deforested by fire. 
Murray Pass, Wharton Range, Central Division, Papua, 2840 m (L. J. Brass, 1933). 



Dec. 1963] 



Cyatheaceae (Holttum) 



67 




Fig. 2. Cyathea gleichenioides C. Chr., same locality as fig. 1. Trunks have been blackened by a recent 

grass fire (L. J. Brass). 



T/ielypteris 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, pi. 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. Jacob 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 moliiccaiia, C. sqiiamulata and C. glabra 
occur only in quite shady forest and will not tolerate exposure; C latebrosa 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. Neerl. 
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 



[ser. II, vol. P 




Fig. 



3. Cyathea contaminans (Wall.) Copel. with branching of upper part of trunic. Mt Telemojo, 
Central Java (P. Arens). — Fig. 4. The branched trunic shown in iig. 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. Schoute (I.e.) 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 f^ronds was 
21 days (Mitteil. Naturf. Ges. SchafThausen 12, 1943, 211-217). It is remarkable that fronds thus ap- 



Dec. 1963] Cyatheaceae (Holttum) 69 

pearing singly leave ultimately the scars of their bases in regular alternate whorls. In New Guinea, HooG- 
LAND has observed that some species produce their fronds in whorls, those of one whorl being simul- 
taneous. More observations on phyllotuxis 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. cuistrdlis (R. Br.) Domin and Dicksunia antcirctica, which both have fronds in many 
orthostichies. Jaag observed three plants o^ Dicksunia bluinei 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 Cyothea (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, Ciilcita 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 all 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 difterence 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. II, vol, P 



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. 







e 





Fig. 5. Cyathea capensis (L./.) 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 contamimins (Wall.) Copel. Trunk showing scars after abscission of stipes. 
Mt Bukit Tungul, W. Java (L. van der Pijl). 

Gametophyte. Stokey gave a comparative account of gametophytes of all genera except Metaxya 
and Cystodiiim (Bot. Gaz. 90, 1930, 1^5). 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 Lophosuiia, 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 Holttum, Rev. 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 Cystodiiim. 
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 Mettenius, in the course of 
his study of Angiopteris (Abh. M.-Ph. Kl. K. Sachs. Ges. Wiss. 6, 1863, 525-531, t. V.) The most recent 
full account of the anatomy of members of Cvatheaceae 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 Holttum & Sen 



Dec. 1963] Cyatheaceae (Holttum) 71 

(Phytomorphology 11, 1961, 406-420). The vascular structure, with its accompanying sclerotic tissue, 
is very similar in ail Malaysian genera except Cihotiiim; it is most fully developed in Cyaihea, to which 
the followmg 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 Cihutiiim. 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. Ochse & Bakhuizen van den Brink reported the use of coiled young 
fronds of Cyathea contaminans and C. jiinghiihniatia (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-tern 
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.}m 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 w^hich the second was Cra'^'^^c"^^'^' which corresponded 
exactly with the present arrangement with the addition of Matonia (Mettenius used the name Balantium 
in place of Dicksonia). Christ (Farnkr. d. Erde, 1897), Diels (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 Siiperficiales, 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 Chris- 
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 (I.e. 304). In this he disagreed with Goebel (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.l. in a separate family 
Cyatheaceae. Holttum & Sen have published a discussion of the whole question (Phytomorphology 11, 
1961, 406^20), 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. 



72 Flora Malesiana [ser. II, vol. P 

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 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 (1 sp., tropical America) (Lophosoria) 

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 pinnules, or tripinnate; stem usually a thick erect trunk; cubical 

cells as in Cyatheae 2. Dicksonia 

Fronds bipinnate with simple pinnules; stem prostrate; condition of cubical 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 Cukiteae. 

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 Z)/fA-5o/;;a 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 
cells lacking; stomata with 3 subsidiary cells 4. Cibotium 

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 .s/)., tropical America) (Metaxya) 

ARTIFICIAL KEY TO MALAYSIAN GENERA 

1. Upper surface of costae of ultimate leaflets raised; fronds ~ elliptical, mostly bipinnate. 

2. Young parts of plant protected by scales and hairs 1. Cyathea 

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 

4. Pinna-rachis grooved on upper surface; fertile pinnules not lobed 3. Cystodium 

3. Outer indusium quite distinct from lamina of leaflet 4. Cibotium 

1. Upper surface of costae of ultimate leaflets grooved, the groove decurrent into the groove of the 
supporting rachis; fronds deltoid in outline, 3-4-pinnate 5. Culcita 

1. CYATHEA 

Smith, Mem. Ac. Turin 5 (1793) 416; Swartz, Syn. Fil. (1806) 139, 364; Kaulf. 
En. Fil. Chamisso (1824) 254; Presl, Tent. Pterid. (1836) 54; Hook. Gen. Fil. 
(1839) t. 23; Sp. Fil. 1 (1844) 14; Syn. Fil. (1865) 16; J. Smith, Lond. J. Bot. 1 
(1842) 659-668; Hist. Fil. (1875) 244; Christ, 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 orieiUalis (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). 



m.—Hemitelia R. Br. Prod. (1810) \5%, p.p. (excl. H. horrida).—Alsophila R. 
Br. I.e. — Clmoophora Kaulf. En. Fil. Chamisso (1824) 250. — Gymnosphaera Bl. 
En. PI. Jav. (1828) 242; Copel. Gen. Fil. (1947) 9S.—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 Presl, Abh. K. Bohm. Ges. Wiss. V, 5 (1848) 55.— Fourniera Bommer, 
Bull. Soc. Bot. France 20 (1873) xix.—Eatoniopsis Bommer, I.e. — Thysanobotrya 
V. A. V. R. Bull. Jard. Bot. Btzg 11, n. 28 (1918) 66.— Fig. 1-30. 

Trunk always erect, short or tall. Seales 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; hairs 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-raehis and costa raised; veins simple or branched, 



74 



Flora Malesiana 



[ser. II, vol. 12 



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, i 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 siibg. 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 setiferoiis (see Holttum, Kew Bull. 1957, 41-45; Holttum & 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 Copel. b. Part of flabelloid margin of scale from stipe, x 100. — C. squamulata (Bl.) Copel. c. 
Apical part of scale from costa of a pinnule, showing setiferous (not flabelloid) edge, x 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 







96:2 




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 Copel.; indusium attached on costular side of receptacle, covering part of base of 
mature sorus. — d. C. oinops Hassk.; 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, x 6; b-f, x 10; a De 
Vriese 323, c Elmer 11634, d Matthew s.n., e Koorders 37469, /Meijer 119). 



76 Flora Malesiana [ser. II, vol. 1^ 

of equal thickness (they are rarely so thick as in flabelloid scales); some marginal cells grow obliquely 
outwards at their distal ends to form straight or outcurved rigid, usually dark, setae on the edge of the 
scale (fig. 8a). 

Taxon. Early authors attempted to distinguish genera (within the genus Cyathea as here recognized) 
by characters of the indusium, whether cup-shaped, attached to one side of the receptacle, or absent. 
These three conditions do not cover all cases, and indusia have often been inadequately described. 
Species lacking indusia often resemble indusiate species more closely than other exindusiate ones. 
It thus appears that the exindusiate condition has arisen on more than one evolutionary line, and it does 
not give a natural subdivision of the genus. Presl attempted also to distinguish species in which the 
receptacle splits into two halves {Disphenici, Dichorexia), but this is not a significant characters. 

CoPELAND at first united all Malaysian species in the genus Cyathea (Philip. J. Sc, Bot. 3, 1909, 353; 
4, 1909, 28) but later attempted to distinguish the three genera Cyathea, Gymnosphaera and Schizocaena 
(Gen. Fil. 1947, 94-99), a division which I criticized (Kew Bull. 1957, 41-45). Most of the species of 
Copeland's Gymnosphaera sect. 3 appear to be closely related to the type species of Schizocaena. Elim- 
inating these, I have found it extremely difficult to make a clear-cut separation between Gymnosphaera 
and some species included by Copeland in Cyathea. Copeland's Cyathea is divisible, on the basis of 
scale-characters (see above) into two groups which I call siibg. Cyathea (flabelloid scales) and siibg. 
Sphaeropteris (setiferous scales). Gymnosphaera (excluded Copeland's sect. 3) is then a section of 
siibg. Cyathea, and Schizocaena a section of subg. Sphaeropteris. 

In Cyathea subg. Cyathea all possible conditions of the indusium occur, and it appears that species 
with cup-shaped indusia can be closely related to others with hemitelioid indusia; in some species, 
e.g. C.javanica Bl. (fig. 9b) and C. hymenodes Mett., intermediate conditions may occur on the same 
leaflet as typical indusia. Some species have very large hemitelioid indusia which cover the sorus almost 
to maturity, often breaking later (C. loheri Christ, C. oinops Hassk., fig. 9d); these have usually not been 
distinguished from species in which the indusium is at first quite complete {e.g. C. creniilata Bl., fig. 9f). 
Other species show various stages of reduction of the hemitelioid type of indusium; in many cases this is 
quite hidden by the mature sorus (fig. 18) and has been reported as lacking (the species thus being placed 
in Alsophila). In subg. Sphaeropteris the hemitelioid condition has not been found. Most species of this 
subgenus have a complete indusium, breaking at maturity (never truly cup-shaped) or none; two cases 
where partial indusia occur (C. alternans (Wall.) Pr. and C. discophora Holttum) appear to be inter- 
mediate between fully exindusiate and exindusiate species and are probably hybrids. 

Descriptions of species of Cyathea have rarely been satisfactory, and misidentifications have been 
frequent; hence names in collectors' lists, and distribution data based on them, are often unreliable. 
Eor clear distinction between species, detailed descriptions of scales, hairs and indusia are essential, and 
often these cannot be seen satisfactorily with a ■ 10 lens. The only authors who described such details 
adequately were Mettenius and Christensen. Because of inadequate early descriptions, many species 
have been named more than once, and the only way to know this is to examine type material. 1 have seen 
such material of almost all the 350 species described from Malaysia, and have examined also types of 
species from the mainland of Asia and from the Pacific. 

SUBDIVISION OF THE GENUS CYATHEA 

1. Stipe-scales flabelloid; hairs on lower surfaces, if present, crisped and appressed; pinnules in most 
cases deeply lobed, basal basiscopic vein rarely from costa; indusia in some cases hemitelioid. 

Subg. Cyathea. 
2. Indusiate (indusia in some cases very small) or if exindusiate hairy on lower surface of pinna-rachis; 
axes not very dark; little dimorphism between sterile and fertile pinnules. Spp. 1-103. 

1. Sect. Cyathea 

2. Exindusiate; axes very dark, not hairy beneath; fertile and sterile pinnules usually very dimorphous. 
Spp. 104-120 2. Sect. Gymnosphaera 

1. Stipe-scales setiferous; hairs on lower surfaces, if present, rather thick and spreading; where pinnules 
are shallowly lobed, basiscopic vein always from costa; indusia complete, or lacking, or formed of 
separate scales (in a few cases imperfect, and then not hemitelioid). Subg. Sphaeropteris. 

3. Costulesnot widely spaced (rarely over 4 mm apart in pinnules 10 cm long); pinnules usually 10 cm 
or more long, lobed almost or quite to costa throughout, or fully pinnate. Spp. 121-151. 

3. Sect. Sphaeropteris 

4. Free tertiary leaflets few; indusia present or absent; sori never covered with overlapping scales. 

Spp. 121-143 3a. Subsect. Sphaeropteris 

4. Eree tertiary leaflets many; no indusia; sori covered with overlapping scales. Spp. 144-151. 

3b. Subsect. Fourniera 

3. Costules widely spaced (at least 4 mm apart except where pinnules are under 4 cm long); pinnules 

mostly less than 10 cm long, not lobed to within 1 mm of costa except near base; basal basiscopic 

vein always from costa; indusiate or not. Spp. 152-191 4. Sect. Schizocaena 

5. Scales of stipe 1 cell thick throughout. 5/>p. 152-178 4a. 5'///w£'fA Schizocaena 

5. Scales of stipe thick and fleshy at base, tapering and flat distally (fig. 30). Spp. 179-191. 

4b. Subsect. Sacropholis 



Dec. 1963] Cyatheaceae (Holttum) 77 

Subgenus Cyathea 

1. Section Cyathea 

Distr. Pantropic— Fig. 1-2, 7-18. 

Taxon. This is by far the largest subdivision of the genus, and includes species with all forms of indu- 
sia and with none. I have not been able to distinguish sharply defined groups which could appear as 
main divisions of the key. The main key-characters are based on indusia, and allied species are some- 
times separated by this method. The species with hemitelioid indusia are the most difficult to characterize 
clearly; this group is particularly polymorphic in Sumatra and in the Philippines. There may be hybrids, 
but the remarkably uniform characters of C. Icitebrosa (Wall.) Copel., a common species in Malaya of 
which 1 have seen a large number of specimens, indicates that quite small differences can be constant. 
Size of pinnules is not a reliable character, unless one can see ample material; plants growing in exposed 
conditions may have much smaller pinnules than others of the same species in the shade. However, the 
very large pinnules of C. iiicisosenatci Copel., otherwise very near C. Icitebrosa, appear to be constant, 
and are exactly reproduced in new plants raised from spores at Kew. For local floras it will probably be 
possible to devise keys based on macroscopic characters easily observed in the field; in preparing keys to 
cover all Malaysian species, I have not been able to use such characters. 

I am not sure whether the hemitelioid species of tropical America should be placed in this section, or 
in a separate section; if the latter, the name Hemitelia is available for them (type-species H. midtiflora 
(Sm.) R. Br.). Some exindusiate tropical American species appear distinct in their dermal appendages. 
These have been referred to Alsophila, but do not appear to be nearly related to the type-species of 
Alsopliila, A. aiistraHs R. Br.; they need further study. 

Ecol. All species, except those of very high altitudes, are more or less shade-demanding. 

KEY TO THE SPECIES 

1. Pinnules not over 30 mm long; segments mostly constricted at base or the lower ones quite free. 
2. Lower surface of pinna-rachis bearing crisped hairs. 

3. Sori lacking indusia 1. C. lepidoclada 

3. Sori indusiate; indusium cup-shaped. 
4. Bullate scales present on pinna-rachis and/or costae; stipe-scales to 2 mm wide. 

5. Pinnules to c. 30 mm long 2. C. microphylloides 

5. Pinnules to c. 15 mm long 3. C. perpelvigera 

4. Bullate scales lacking from pinna-rachis and/or costae; stipe-scales to 3 1/2 rnrn wide. 

4. C. hunsteiniana 
2. Lower surface of pinna-rachis not hairy. 

6. Longest pinna 8V2 cm, pinnules to 11 mm long 5. C. arfakensis 

6. Longest pinna 22-26 cm, pinnules 23-30 mm long. 
7. Stipe 30 cm. Pinna-rachis glabrescent. Sori 4-5 to each tertiary leaflet .... 6. C. ledermannii 
7. Stipe 10 cm. Pinna-rachis covered with small bullate scales. One sorus at base of each tertiary 

leaflet 7. C. hooglandii 

\. Pinnules longer, or if under 30 mm long without free basal segments. 

8. Indusium an entire cup with even edge at maturity of sorus, if fragile sometimes breaking later. 
9. Lower surface of lamina-segments almost or quite covered with scales and hairs. 
10. Small scales abundant on lower surface of veins; costal scales mostly bearing dark setae. 

8. C. percrassa 

10. Crisped hairs on lower surface of veins; costal scales bearing many slender crisped marginal hairs. 

9. C. vandeusenii 
9. Lower surface of lamina-segments not so covered. 

11. Lower surface of lamina strongly glaucous 10. C. pruinosa 

11. Lower surface of lamina not glaucous. 

12. BuUate-based scales present on pinna-rachis and/or costae. 
13. No fine crisped hairs on lower surface of pinna-rachis nor on edges of its scales. 

11. C. pycnoneura 
13. Fine crisped hairs present on pinna-rachis or on its scales. 
14. Indusial cup shallow. Small scales on pinna-rachis bearing very fine hairs; no coarse crisped 

hairs; bullate scales only on distal part of pinna-rachis 12. C. rigens 

14. Indusial cup deep. Crisped hairs, not small fringed scales, also bullate scales, present on 

pinna-rachis 13. C. everta 

12. BuUate-based scales lacking on pinna-rachis and/or costae (at most convex ovate brown scales 
on costa). 

15. Lower surface of pinna-rachis covered with crisped hairs 14. C. cincinnata 

15. Lower surface of pinna-rachis lacking such hairs. 
16. Pinna-rachis almost glabrescent on lower surface. 

17. Larger pinnules bearing c. 6 pairs of free tertiary leaflets .... 15. C. subtripinnata 



78 Flora Malesiana [ser. II, vol. P 

17. Larger pinnules bearing at most one pair of free leaflets. 
18. Pinnules commonly to 100 mm long, basal lobes not free. Long brown scales lacking on 

costae 16. C. orientalis 

18. Pinnules commonly to 65 mm long, basal pair of lobes free. Long dark scales abundant on 

costae 17. C. apoensis 

16. Pinna-rachis persistently scaly on lower surface. 
19. Scales on pinna-rachis mostly narrow, 4-5 mm long, with long flexuous marginal setae. 

Stipe over 20 cm 18. C. costalisora 

19. Scales on pinna-rachis mostly very small, some or all bearing short setae. Stipe c. 10 cm. 

20. Scales on pinna-rachis dark, all setiferous 19. C. pallidipaleata 

20. Scales on pinna-rachis thin, short-fringed, mostly not setiferous, forming a continuous felt. 

20. C. coactilis 
8. Indusium otherwise or lacking. 
21. Indusium at maturity an almost flat disc, symmetric or not, in some cases hidden by mature sorus, 
in some cases the residual part of a complete indusium of which the thin apical parts falls away at 
maturity. 
22. Indusium a very narrow ring round base of receptacle. 
23. On lower surface of pinna-rachis and costae very small scales bearing long crisped hairs, or 
single crisped hairs. 

24. Scales on costae bullate-based to buUate 21. C. parva 

24. Scales on costae flat or lacking, not buUate. 

25. Broad flat pale scales with a few marginal setae on costae 22. C. wengiensis 

25. Broad flat setiferous scales lacking on costae 23. C. batjanensis 

23. On lower surface of pinna-rachis small scales with short fringe of hairs. 24. C. ternatea 
22. Indusium ultimately a regular or irregular disc, almost or quite as big as base of sorus; if ir- 
regular, usually the base of a formerly complete indusium. 
26. Indusium at maturity a disc with fairly even edge (except in C. patellifera). 
11. Pinna-rachis hairy on lower surface, at least distally. 
28. Pinna-rachis bearing copious bullate-based scales on lower surface . 25. C. albidosquamata 
28. Pinna-rachis lacking such scales. 
29. Pinnules to at least 85 by 20 mm, veins 8-10 pairs. 
30. Pinna-rachis and costae densely hairy on lower surface; costae glabrous or nearly so on 
upper surface 26. C. javanica 

30. Pinna-rachis hairy towards apex only; costae with few hairs on lower surface, many on 
upper surface 27. C. hymenodes 

29. Pinnules c. 30-70 by 12-15 mm, veins 4-6 pairs. 

31. Lower 3^ pairs of segments on each pinnule free and deeply lobed. Indusium irregularly 
lobed 28. C. patellifera 

31. Lowest segments not free, crenate. Indusium an entire disc. . 25. C. albidosquamata 
27. Pinna-rachis not hairy on lower surface. 
32. Distal part of pinna-rachis covered with scales. 

33. Lowest pinna c. 6 cm long, stipe short 29. C. negrosiana 

33. Lowest pinna much longer, stipe not very short. 

34. Scales on stipe dark; scales on pinna-rachis buUate 30. C. catillifera 

34. Scales on stipe pale; scales on pinna-rachis not bullate, larger ones having marginal setae. 

31. C. horridula 

32. Distalpartof pinna-rachis glabrescent, never covered with bullate scales. 43. C. bunnemeijerii 

26. Indusium at first covering sorus, apical part very thin and caducous, an irregular disc remaining 

on old sorus. 

35. Pinna-rachis hairy on lower surface, at least distally. 

36. Pinnules c. 50 by 15 mm, pinnae to 21 cm long 32. C. tenuicaulis 

36. Pinnules to 95 by 18 mm, pinnae to 50 cm long 33. C. sumatrana 

35. Pinna-rachis not hairy on lower surface. 

37. Frond simply pinnate, pinnae to 7 by 13/4 cm, lobed to 2 mm from costa. 34. C. klossii 
37. Frond bipinnate. 

38. Small pale fringed scales abundant at least on costae. 
39. Spines on stipe 2-3 mm long; rachises ± covered beneath with small fringed scales as on 
costae 35. C. trachypoda 

39. Spines on stipe less than 1 mm long; rachises not so covered .... 36. C. crenulata 
38. Small pale fringed scales lacking or very few. 

40. Stipe bearing spines 3-5 mm long. 

41. Larger scales on costae bearing dark setae 37. C. macropoda 

41. Larger scales on costae not bearing dark setae. 

42. Small scales on costae bearing long crisped hairs 38. C. saccata 

42. Small scales on costae lacking such hairs. 



Dec. 1963] Cyatheaceae (Holttum) 79 

43. Largest pinnules 150-175 by 30-40 mm 39. C. magnifolia 

43. Largest pinnules c. 90 by 18 mm 40. C. acanthophora 

40. Stipe not spiny, or spines under I mm long. 
44. Stipe-scales less than 1 mm wide above the base, edges when young setiferous; abundant 

narrow scales bearing long setae on pinna-rachis 4L C. rubiginosa 

44. Stipe-scales otherwise; pinna-rachis scales rarely setiferous. 

45. Stipe densely scaly throughout 42. C. apiculata 

45. Stipe persistently scaly near base only. 
46. Stipe long; lowest pinnae not greatly reduced. 
47. Stipe dark, scales firm, shining; bullate scales abundant on costules. 

43. C. biinnemeijerii 
47. Stipe green, scales thin, dull; no bullate scales on costules . . . 44. C. excavata 

46. Stipe short; lowest pinnae c. 5 cm long 45. C. christii 

21. Indusium otherwise or lacking. 
48. Indusium covering sorus to maturity, then breaking and persistent. 

49. Pinna-rachis conspicuously hairy and scaly on lower surface 46. C. geluensis 

49. Pinna-rachis not hairy, though sometimes scaly on lower surface. 
50. Lamina very rigid, the small tertiary leaflets with edges much reflexed so that the sori are almost 
enclosed. 
51. Largest tertiary leaflets lobed, each with 3-5 sori 47. C. macgregorii 

51. Tertiary leaflets not lobed, each with 1-2 sori 48. C. gleichenioides 

50. Lamina not very rigid with strongly reflexed edges. 

52. Pinnules to 21/2 cm long, less than 10 mm wide. 

53. Pinna-rachis and costae persistently brown-scaly; scales entire, smaller ones bullate. 

49. C. havilandii 

53. Pinna-rachis and costae glabrescent; scales small, not bullate . . 50. C. imbricata 
52. Pinnules to at least 45 by 10 mm, in most cases much larger. 

54. Largest pinnules more than 100 by 20 mm; costules 5 mm or more apart. 

55. Bullate scales present on costules. Pinnules distinctly stalked; several pairs of free segments 
on larger pinnules. 
56. Stipe long, slender, very spiny. Pinnules commonly with stalks 3-6 mm long, to 10 mm on 

lowest 51. C. longipes 

56. Stipe not known. Pinnule-stalks not over 4 mm 52. C. acuminata 

55. Bullate scales lacking on costules. Pinnules sessile or on stalks to 2 mm; at most 1-2 pairs 

of free segments 53. C. insulana 

54. Largest pinnules not over 100 by 20 mm; costules 3-41/2 mm apart. 
57. Pinnules c. 45 by 10 mm, very rigid. Stipe-scales rigid, 40 by 1 mm. 

54. C. pseudomuelleri 
57. Pinnules commonly more than 65 by 15 mm. Stipe-scales otherwise. 
58. Costae and costules densely scaly; scales mostly setiferous, not bullate; pinna-rachis 
persistently covered with very small setiferous scales. 
59. Veins bearing scales on lower surface. 

60. Stipe 50 cm or more 55. C. archboldii 

60. Stipe 5-15 cm, lowest pinna 5-12 cm. 
61. Larger scales on pinna-rachis pale. Veins dark and raised on lower surface. 

56. C. foersteri 
61. Larger scales on pinna-rachis with dark median band. Veins concolorous, not raised 

below 57. C. nigrolineata 

59. Veins lacking scales on lower surface. 

62. Large scales on stipe and rachis light red-brown 58. C. inquinans 

62. Large scales not red-brown. 

63. Larger scales on pinna-rachis pale 56. C. foersteri 

63. Larger scales on pinna-rachis with dark median band. . . . 57. C. nigrolineata 
58. Costae and costules bearing scales which are mostly not setiferous; pinna-rachis glabres- 
cent or its scales mostly not setiferous. 
64. Stipe-scales pale; pinna-rachis rather persistently covered with small scales beneath; 

veins bearing some scales 56. C. foersteri 

64. Stipe-scales dark; pinna-rachis glabrescent; veins not scaly. 
65. Costae and/or costules bearing bullate scales. 
66. Pinnules at base of larger pinnae distinctly stalked (stalks 2-4) mm; costules 4-41/2 

mm apart 59. C. ferruginea 

66. Pinnules sessile; costules usually not over 31/2 mm apart. 

67. Indusium firm, brown 60. C. oosora 

67. Indusium pale, fragile 45. C. christii 

65. Costae and/or costules lacking bullate scales 61. C. halconensis 



80 Flora Malesiana [ser. II, vol. P 

48. Indusium hemitelioid (sometimes almost covering sorus to maturity, sometimes very small) 
or lacking. 
68. Jndusium lacking. 
69. Fronds simply pinnate, or bipinnate with small pinnules lobed halfway to costa. 
70. Fronds simply pinnate 62. C. ascendens 

70. Fronds bipinnate. pinnules c. 30 mm long 63. C. recurvata 

69. Fronds bipinnate, pinnules more deeply lobed. 

71. Pinna-rachis densely covered with long tangled pale crisped hairs and narrow scales on lower 
surface 64. C. eriophora 

71. Pinna-rachis rather sparsely hairy on lower surface. 

72. Sori near costules. Dark hairs present on costae beneath 65. C. gregaria 

72. Sori medial or nearly so. No dark hairs on costae beneath. See sect. Gymnosphaera. 

116. C. macgillivrayi 
68. Indusium hemitelioid, of varying size, sometimes hidden by sorus. 
73. Pinna-rachis densely hairy throughout on lower surface. 
74. Bullate scales abundant on costae. 
75. Raised median part of upper surface of pinna-rachis and costae hairy 66. C. modesta 

75. Raised median part of upper surface of pinna-rachis and costae glabrous. 

67. C. doctersii 
74. Bullate scales absent, or a few distally on costae. 

76. Pinnules to 40 by 10 mm. Scales on costae setiferous 68. C. cucullifera 

76. Pinnules much larger. Scales on costae not setiferous 69. C. setulosa 

73. Pinna-rachis hairy at most on distal part. 
77. Indusium entirely brown and rather firm, quite covering sorus to maturity, breaking only 
when old. 
78. Stipe-scales 50 mm long, shining brown. Costae almost glabrous. Pinnules c. 50 mm long. 

70. C. muelleri 
78. Stipe-scales 20-35 mm long, pale. Costae very scaly. Pinnules to 100 mm long. 
79. Larger scales on costae uniformly brown, rather thin, edges with some setae. 
80. Bullate scales lacking. Lower pinnae not gradually reduced (sometimes a pair 5 cm long 

near base of stipe) 71. C. oinops 

80. Bullate scales on costules. Lower pinnae gradually reduced to 7 cm long. 
8 1 . Pinna-rachis densely covered with overlapping scales. Indusium firm, shining. 72. C. loheri 
81. Pinna-rachis sparsely covered with very small pale fringed scales. Indusium dull, thinner. 

73. C. cinerea 
79. Larger scales on costae with narrow very dark median band and broad pale edges, or en- 
tirely pale 74. C. pachyrrhachis 

77. Indusium in part pale and fragile, breaking at maturity, or not entirely covering sorus. 
82. Pinnules 20-40 mm wide; costules 5-6 mm apart; lowest basiscopic vein from costa or base 

of costule 75. C. latipinnula 

82. Pinnules rarely over 2 cm wide; costules not over 5 mm apart; basiscopic vein always from 
costule. 
83. Indusium visible as a scale backing the costule, not entirely hidden by sorus. 
84. Pinnules conspicuously stalked; stalks to 7 mm long. 
85. Stipe bearing spines 3^ mm long. No bullate scales on costules. 

76. C. masapilidensis 

85. Stipe not spiny. Bullate scales present on costules 77. C. loerzingii 

84. Pinnules not conspicuously stalked. 

86. Indusium more than a semicircle, concave towards sorus and sometimes covering part 
of sorus to maturity. 

87. Pinna-rachis densely and persistently scaly 78. C. rufopannosa 

87. Pinna-rachis not densely and persistently scaly. 

88. Indusium fragile, breaking and often in part disappearing 79. C. callosa 

88. Indusium firm throughout. 
89. Frond almost fully tripinnate; tertiary leaflets with strongly reflexed edges. 

80. C. dicksonioides 

89. Frond bipinnate with at most 1-2 pairs of tertiary leaflets; edges of segments not 

strongly reflexed. 

90. Pinnules c. 100 mm long. Segments of lamina mostly not constricted at base on 

acroscopic side; bullate or convex pale scales on costules 81. C. heterochlamydea 

90. Pinnules c. 60 mm long. Segments of lamina mostly constricted at base on acroscopic 

side; no bullate scales 82. C. edanoi 

86. Indusium not more than a semicircle, reflexed against costule at maturity. 
91. Bullate scales abundant and rather persistent on pinna-rachis (at least distally); pinna- 
rachis closely and finely warty after fall of scales. 



Dec. 1963] Cyatheaceae (Holttum) 81 

92. Stipe-scales dark with thin pale edges. Basal 1-2 segments of pinnules almost free. 

83. C. fuliginosa 

92. Stipe-scales (at least larger ones) pale. Basal 6 pairs of segments almost free, separately 
adnate to costa 84. C. semiamplectens 

91. Bullate scales lacking on pinna-rachis. 

93. Basal scales on costae bearing some marginal setae. 

94. Bullate scales abundant on costules. Stipe long, basal pinnae not much reduced. 
95. Costules AVi-SVj mm apart. Pinnules 100 mm or more long. 85. C. alleniae 
95. Costules i-^Vi mm apart. Pinnules to 60 mm long 86. C. costulisora 

94. Bullate scales absent. Stipe short, basal pinnae gradually reduced, lowest very short. 

87. C. caudata 

93. Basal scales on costae without setae. 
96. Bullate scales present (if at all) towards apex of costa; costal scales few. Pinnules to 
100 mm long. 
97. Scales on costules distinctly bullate; pinna-rachis slightly hairy towards apex on lower 

surface 88. C. borneensis 

97. Scales on costules mostly flat or convex, or distal ones bullate; pinna-rachis not 
hairy on lower surface. 
98. 1-2 pairs basal segments on larger pinnules almost or quite free. 89. C. fenicis 

98. Basal segments not free 90. C. junghuhniana 

96. Bullate scales abundant to base of costae and on costules. Pinnules to 65 mm long. 

91. C. raciborskii 
83. Indusium very small, hidden by sporangia. 
99. Pinnules commonly more than 20 mm wide. 

100. Pinnules cut to 3-4 mm from costa 92. C. glaberrima 

100. Pinnules cut almost to costa throughout. 

101. Lower pinnules distinctly stalked. Sinuses between segments narrow. 93. C. punctulata 
101. Lower pinnules sessile or nearly so. Sinuses between segments wide. 

94. C. incisoserrata 
99. Pinnules commonly not more than 20 mm wide. 
102. Pinna-rachis densely covered with bullate scales; pinnules c. 32 by 10 mm. 

95. C. physolepidota 
102. Pinna-rachis not so covered; pinnules larger. 
103. Scales on costae and costules all flat. 
104. Lower pinnules stalked. No hairs on pinna-rachis and costae. 
105. Lowest 1-2 pairs of segments quite free and articulate. Lamina thick, rigid. 

96. C. kanehirae 

105. Lower segments, several pairs, contracted at base but not free. Lamina not thick. 

97. C. nigropaleata 
104. Lower pinnules sessile. Hairs present on pinnae-rachis and costae. 

106. Sori near costules (Luzon) 98. C. microchlamys 

106. Sori medial (New Guinea) 22. C. wengiensis 

103. Scales on costules bullate; some bullate scales often on costae. 

107. Pinnules articulate to pinna-rachis 99. C. perpunctulata 

107. Pinnules not articulate. 
108. Paraphyses conspicuous, longer than sporangia, 2-3 cells wide at base. 
109. Stipe-spines 3^ mm long. Pinna-rachis hairy towards apex on lower surface. 

100. C. aldernereltii 
109. Stipe-spines l-2i 2 mm long. Pinna-rachis not hairy on lower surface. 
110. Lamina-segments strongly crenate-serrate. Indusium minute. Bullate scales 
throughout costae. 
111. Stipe over 50 cm; lowest pinnae little reduced. . . . 101. C. amboinensis 

111. Stipe 20-25 cm; lowest pinna 8-12 cm long 102. C. media 

110. Lamina segments shallowly crenate. Indusium distinct, 2-lobed; bullate scales 

only on distal part of costa 103. C. latebrosa 

108. Paraphyses shorter than sporangia, slender. 
112. Pinna-rachis hairy towards apex on lower surface. Lower pinnae much reduced and 

spaced 88. C. borneensis 

112. Pinna-rachis not hairy on lower surface. Lower pinnae not greatly reduced. 
113. One to two pairs of basal segments on larger pinnules almost or quite free. 

89. C. fenicis 
113. No free segments at base of pinnules. 

114. Stipe-thorns 5 mm or more long 40. C. acanthophora 

114. Stipe-thorns to 21/2 mm long 90. C. junghuhniana 



82 



Flora Malesiana 



[ser. II, vol. 12 



1. Cyathea lepidoclada (Christ) Domin, Acta Bot. 
Bohem. 9 (1930) 130; C. Chr. Brittonia 2 (1937) 
278;CoPEL. Philip. J. Sc. 77 (1947) 120.— Alsophila 
lepidoclada Christ in K. Sen. & Laut. Nachtr. 
(1905) 37; v. A. v. R. Handb. (1908) 37; Handb. 
Suppl. (1917) 62. 

Trunk slender, to 2 m; fronds to 150 cm. Stipe 
10-24 cm; scales to 20 by 4-5 mm, shining brown 
with dull thin edges; scales on upper part of stipe 
and on rachis paler. Lower pinnae gradually 
reduced, lowest 3-6 cm long, longest 16-22 cm. 
Pinnules to 20 by 8 mm, segments c. 6 pairs, 
distinctly oblique, entire or slightly crenate, lowest 
free or contracted on acroscopic base; costules 
little over 2 mm apart; veins 3-4 pairs, simple. 
Sori near costules, rather small, not indusiate, 
paraphyses not longer than sporangia. Scales and 
hairs: lower surface of main rachis densely covered 
with interlacing pale flexuous hairs which are 
branched at the base, with scattered flat elongate 
scales; pinna-rachis similarly clothed, the scales 
more abundant, to about 3 by % mm; pale hairs 
on costae more sparse, the scales bullate; hairs 
also scattered on lower surface of costules and 
veins. 

Type specimen: Schlechter 14417, Torricelli 
Mts, E. New Guinea (P; dupl. at K, BM, BO). 

Distr. Malaysia: Central and E. New Guinea 
(3 collections). 

Ecol. At 800-1000 m, 'frequent on slope in 
rain forest' (Brass). 

2. Cyathea microphylloides Ros. in Fedde, Rep. 12 
(1913) 164; V. A. v. R. Handb. Suppl. (1917) 38.— 
C. peranemiformis C. Chr. Brittonia 2 (1937) 277. 

Trunk slender, to 1 m; fronds less than 100 cm 
long. Stipe 3-10 cm; scales to 15 by 2 mm, median 
band shining, dark or sometimes partly or en- 
tirely pale, with broad dull fragile edges. Lower 
pinnae gradually reduced, lowest 3-5 cm long, 
longest to c. 20 cm. Pinnules to c. 30 by 10 mm, 
lobes almost all constricted at the base (connected 
by a very narrow wing along the costa) but only 
the lowest quite free, edges slightly crenate; 
costules to 3'/2 mrn apart; veins 4-5 pairs, the lower 
ones forked. Sori to 3 or 4 pairs on each segment 
of a pinnule; indusium ultimately a shallow light 
brown cup round base of sorus; receptacle prom- 
inent, paraphyses slender, short. Scales and 
hairs: main rachis bearing many long narrow 
crisped spreading brown scales, and minute fringed 
scales, on lower surface; lower surface of pinna- 
rachis bearing many bullate-based acuminate 
brown scales, the smaller ones hair-pointed, and 
also crisped brown hairs; costae bearing similar 
small bullate scales. 

Type specimen: Keysser B71, Bolan Mts, E. 
New Guinea (S-PA; dupl. at B). 

Distr. Malaysia: Eastern and Central New 
Guinea. 

Ecol. At c. 1800-3000 m; Brass notes of type 
of C. peranemiformis 'common in forests of slopes 
and valleys'. 



3. Cyathea perpelvigera v. A. v. R. Nova Guinea 
14(1924) 1 1 ; CoPEL. Philip. J. Sc. 77 (1947) 119, 120. 

Trunk to 2'/! m by 5 cm 0; leaf-bases persistent; 
fronds c. 10, spirally arranged. Stipe 15-20 cm, 
base bearing spines 2 mm long; scales shining 
brown with dull fragile edges, to 1 by almost 2 mm, 
narrower on distal part of stipe and on rachis; 
rachis bearing abundant crisped short brown hairs 
on lower surface and some residual very narrow 
long twisted scales. Lamina of frond 40-60 by 
20-25 cm, lower pinnae gradually reduced. Largest 
pinnae 10-15 cm long, lowest pinnae 4 cm. Pin- 
nules close together, more than 30 pairs on larger 
pinnae, to 15 by 5 mm, almost fully pinnate with c. 
6 pairs of tertiary leaflets and a lobed apex; 
tertiary leaflets to 3 by almost 2 mm (fertile ones 
1 mm wide), edges entire or crenate; veins 3^ 
pairs, lowest forked. Sori 1-2 (rarely 3) to a leaf- 
let; indusium forming a firm brown entire cup 
about half the height of the mature sorus. Scales 
and hairs: lower surface of pinna-rachis bearing 
copious crisped brown hairs, also brown bullate 
long-acuminate scales; similar scales on lower 
surface of costae, with a few hairs. 

Type specimen: Lam 1441, ridge near Doorman 
summit, W. New Guinea (BO; dupl. at L, K, S, 
US, UC). 

Distr. Malaysia: New Guinea, Moluccas 
(Ceram), N. Celebes (doubtful; plant young and 
sterile). 

Ecol. At 1200-1800m both in New Guinea and 
in Ceram; reported by Brass as abundant in rain- 
forest in absence of woody undergrowth and by 
ScHODDE in mixed Nothofagus forest. 

4. Cyathea hunsteiniana Bralse, Bot. Jahrb. 56 
(1920) 58 {incl. var. acuminata); Copel. Philip. 
J. Sc. 77 (1947) 119. 

Trunk 2 cm 0; fronds to 100 cm long. Stipe 
8-16 cm; scales to 13 by 3 '/2 mm, dark with rather 
broad flabelloid edges. Pinnae 25-30 pairs, lower 
ones gradually reduced, lowest 3 cm long, longest 
12 cm. Pinnules almost sessile, to 12 by 5 mm, 
segments to 7 pairs, lowest free and more or less 
lobed, middle ones constricted at base. Sori 1 or 2 
to each lamina-segment; indusium a deep firm 
brown cup with e\en rim. Scales and hairs: lower 
surface of main rachis and pinna-rachis densely 
covered with appressed shining brown flexuous 
hairs; lower surface of costae with few such hairs; 
no scales seen. 

Type specimen: Ledermann 11139, Hunstein- 
spitze, E. New Guinea. 

Distr. Malaysia: Eastern New Guinea. 

Ecol. At 1300-2000 m. 

Notes. This is very near C. perpelvigera, but 
appears to lack scales on the frond. Brause 
distinguished var. acuminata (at 2070 m), with 
fronds long-acuminate and sori always solitary; 
it looks like a less robust plant, possibly grown in 
a more shady place than usual. 

5. Cyathea arfakensis Gepp in Gibbs, Arfak (1917) 
69. — Hemitelia arfakensis v. A. v. R. Bull. Jard. 
Bot. Btzgll, n. 28 (1918)26. 



Dec. 1963] 



Cyatheaceae (Holttum) 



83 




Fig. 10. Cyathea hooglandii Holttum. a. Part of pinna, upper surface, ■ 2, b. same, lower surface, 
showing sori and bullate scales on pinna-rachis and costa, x 6. — C. dicksonioides Holttum. c. Part of 
pinna, upper surface, ■: 2, d. a single tertiary leaflet showing sori, x 8. — C. macgregorii F. v. M. e. Part 
of pinna, upper surface, >' 2,/. one tertiary leaflet showing sori, x 6, g. scale from costule, x 40 {a-b 

HOOGLAND 7203, C HOOGLAND & SCHODDE 1506, d ditto l\l\,e-g HOOGLAND & PULLEN 5745). 



Stipe 12 cm, dark, warty; scales 10 by 1 mm. 
Lamina c. 70 cm long, lowest pinnae 5 cm long, 
longest pinna 8 1/2 cm. Pinnules to 1 1 mm by 4 mm, 
apex rounded, largest with 1 or 2 basal segments 
free as almost circular tertiary leaflets, rest of 
pinnule crenate; lowest pinnules of upper pinnae 
deflexed and overlapping main rachis. Sori in one 
row on each side of costa of pinnule and close to it; 
indusium a broad open dark brown cup with firm 
even edge. Scales and hairs: some small bullate 
scales on costae; no hairs seen on lower surface. 

Type specimen: L. S. Gibbs 6008, Anggi Lakes, 
SW. Ridge, W. New Guinea (BM; dupl. at P, K). 

Distr. Malaysia: W.New Guinea (2 collections). 



Eco 1. In undergrowth of mossy forest at 2500 



m. 



6. Cyathea ledermannii Brause, Bot. Jahrb. 56 
(1920) 56 {incl. var. dilatata. I.e. 58); Copel. 
Philip. J. Sc. 77 (1947) 119. 

Trunk slender, to 21/2 m; fronds few, to 150 cm 
long. Stipe 30 cm, base bearing blunt spines to 
1 mm high and dark shining scales 10-15 by 
1 1/2-2 mm with pale edges. Largest pinnae 22 cm 
long. Pinnules to 23 by 8 mm, with c. 1 pairs of 
entire segments, lowest 2 segments free, next 2 
pairs contracted at base, rest decurrent basis- 
copically to form a wing along the costa; costules 
21/2 mm apart; veins 5-6 pairs, mostly simple. 



84 



Flora Malesiana 



[ser. II, vol. P 



Sori 4-5 on each segment, near costules, usually 
not on basal veins; indusium a very firm deep cup. 
Scales and hairs: pinna-rachis glabrescent on lower 
surface or with a few narrow dark scales; costae 
and costules bearing a few rather large brown buUate 
scales (costae may have narrow flat scales also). 

Type specimen: Ledermann 9651, Sepik region, 
E. New Guinea (B). 

Distr. Malaysia: Central and Eastern New 
Guinea. 

Ecol. At 200-1000 m, in rain-forest or mossy 
forest, locally abundant. 

7. Cyathea hooglandii Holttum, Kew Bull. 16 
(1962) 56.— Fig. 10a. 

Trunk to 3 m by 10 cm o, bearing 10 fronds in 
two whorls of 5 (Hoogland); fronds to 140 cm 
long. Stipe 10 cm, densely covered with scales, 
dull and warty after fall of scales; scales to 20 by 
1 V^ mm, lower ones very dark, shining, upper ones 
medium brown, all with rather broad paler fragile 
edges, not setiferous. Lower pinnae gradually 
reduced, lowest 2-3 cm long, longest 26 cm. 
Largest pinnules 20-30 by 7-8 mm, pinnate, with 
7-11 pairs of tertiary leaflets, the distal ones 
joined by a narrow wing to costa; tertiary leaflets 
ovate to elliptic, to 4 by 1 Y2 mm, edges of distal 
ones entire or sinuous, of basal ones more or less 
deeply lobed at the base, lobes 1-3, acute; veins 
4 pairs, simple. A single sorus at base of each 
tertiary leaflet, apparently seated on its costule or 
at base of lowest acroscopic vein; indusium a 
firm saucer nearly 1 mm wide with the receptacle 
at its centre; receptacle elongate and slightly swol- 
len; paraphyses slender, shorter than sporangia. 
Scales and hairs: lower surface of main rachis 
glabrescent, dull, minutely warty, with residual 
irregular small brown scales; lower surface of 
pinna-rachis rather closely covered with small 
brown, bullate scales, some hair-pointed but most 
not obviously so; similar smaller bullate scales 
on costae of pinnules, not on tertiary leaflets. 

Type specimen: Hoogland & Schodde 7203, 
Western Highlands, NE. New Guinea (L). 

Distr. Malaysia: NE. New Guinea (3 coll.). 

Ecol. In Nothofagus forest at 3000 m (type); 
'in heavily mossed forest', 3170 m (Brass 30216). 

Note. This species appears to be related to 
3. C. perpelvigera and 2. C. microphylloides, but 
is quite peculiar in having the sori singly at the 
bases of the tertiary leaflets, apparently seated 
directly on the costule, not on one of the lateral 
veins as in all other species of Cyathea here re- 
ported. Brass 30216 bears the note 'leaves 7, pale, 
flat-spreading'; possibly the number of leaves in 
a whorl varies from 5 to 7. 

8. Cyathea percrassa C. Chr. Brittonia 2 (1937) 
279; CoPEL. Philip. J. Sc. 77 (1947) 107. 

Trunk 2-A m by 8-12 cm 0, bearing 6-12 fronds 
125-200 cm long; lower pinnae gradually reduced, 
or one small pair near base and then a long gap 
to the next. Stipe not spiny, lower part covered 
with ascending rigid twisted scales 25-40 by 1-2 
mm, with shining median band (pale, streaked, or 



wholly brown) and a rather broad pale fragile 
edge; stipe and rachis also densely covered 
with small dull brown scales which have dark 
setiform apices; pneumathodes 4-6 mm long, 
well spaced, in a single row. Largest pinnae 
30-40 cm long. Largest pinnules of type 80 
by 22 mm, of another collection 55 by 13 mm, 
lowest segments free or nearly so, rest of pinnule 
lobed almost to the costa; costules 2'/2-3 mm 
apart; veins to 10 pairs, mostly forked; lamina- 
segments very firm, edges finely crenate and not 
reflexed, sinuses narrow. Sori near costules; 
indusium a firm light-brown cup, wider than 
deep, with even rim. Scales and hairs: lower 
surface of pinna-rachis densely covered with small 
scales which bear dark setae, also scattered narrow 
scales 3-5 mm long bearing a few dark setae on 
their fragile edges; costae copiously scaly, scales 
near base to 2 mm long, brown to pale with pale 
edges bearing dark setae, grading to small paler 
scales fringed with hairs; costular scales as those 
of costae but not over 1 mm long; lower surface 
of veins densely covered with very small scales, 
some setiferous. 

Type specimen: Brass 4375, Mt Albert Edward, 
E. New Guinea (A; dupl. at BM, MICH, BRI). 

Distr. Malaysia: Eastern New Guinea. 

Ecol. At 3000-3500 m, fairly common in valley 
forest (Brass); fairly common in mossy forest 
(Brass, Hoogland & Pullen) or on edge of mos- 
sy forest. 

9. Cyathea vandeusenii Holttum, Blumea 11 
(1962) 529. 

Trunk stout, to 2 m; fronds c. 10, spreading, 
80-100 cm long. Stipe 12-18 cm, completely cov- 
ered, as lower part of main rachis, with scales; 
scales pale, to 40 by 1-1 '/2 mni, firm and shining 
with narrow dull fragile edges, thick and dark at 
base only, with an under-coat of small thin brown 
scales bearing long flexuous setae. Lowest pinnae 
12-15 cm long, longest 23 cm. Pinnules to 50 mm 
long, 15 mm wide at base, tapering evenly from 
base to apex, 2-3 pairs of basal segments separately 
adnate to costa; costules 3 1/2 mrn apart; veins to 
7 pairs; lamina-segments very rigid, with edges 
reflexed when dry, edges rather deeply crenate 
where fertile. Sori almost completely embedded 
in scales; indusium a complete light-brown cup 
with smooth edge, as wide as deep. Scales and 
hairs: lower surface of all rachises, costae and 
costules covered with a close felt of thin flat 
brown scales bearing many slender crisped mar- 
ginal hairs and scattered flexuous brown setae; 
lower surface of veins bearing crisped pale hairs. 

Type specimen: Brass 29989, Eastern High- 
lands Distr., NE. New Guinea (L; dupl. at K, 
US). 

Distr. Malaysia: NE. New Guinea (one 
collection). 

Ecol. At 3700 m, 'frequent in edges of patches 
of subalpine forest'. 

10. Cyathea pruinosa Rosenst. in Fedde, Rep. 12 
(1913) 163; V. A. v. R. Handb. Suppl. (1917) 29; 
CoPEL. Philip. J. Sc. 77 (1947) 101. 



Dec. 1963] 



Cyatheaceae (Holttum) 



85 



Trunk 5 m; fronds 10-14. Stipe 15 cm or more, 
purplish with a fine glaucous covering, warty 
after fall of scales; scales near base of stipe to 
30 by 1 V2 mm, stiff and twisted, dark brown, 
shining, with paler fragile edges. Lowest pinnae 
15 cm long; largest pinnae 30-50 cm. Largest 
pinnules 50-75 by 15-17 mm, sessile, lobed nearly 
to costa throughout, lowest segment almost free, 
lower surface distinctly glaucous; costules 3 — 31/2 
mm apart; veins 9-10 pairs; lamina-segments rat- 
her rigid, segments crenate. most deeply so when 
fertile. Sori near costules; indusium at maturity 
a rather thin brown cup with entire rim, about as 
wide as deep. Scales and hairs: pinna-rachis rather 
pale, minutely warty on lower surface; scales on 
costae flat, broad, brown, bearing a few long dark 
setae; on costules imbricating fiat broad scales 
as on costae, with ovate convex or almost buUate 
scales distally; a few similar smaller scales on lower 
surface of veins. 

Type specimen: Keysser B44, Bolan Mts, 
E. New Guinea (S-PA; dupl. at B, DC). 

Distr. Malaysia: Eastern New Guinea. 

Ecol. At 2400-3000 m, on edge of forest and 
tree-fern grasslands; also (sterile) in undergrowth 
of forest. 

Note. The type material did not include the 
stipe, description of which is taken from Hoog- 
LAND & ScHODDE 7633, which also bears the 
information that there were 14 fronds in two 
whorls. 

11. Cyathea pycnoneura Holttum, Blumea 1 1 
(1962) 533. 

Fronds to 230 cm long. Stipe more than 30 cm, 
dark, spiny throughout, spines to nearly 3 mm 
long; scales many, dark, to 35 by 2 mm wide near 
base which is thick and hairy, fiabelloid edges 
narrow and mostly abraded with scattered long 
dark setae; very small dull scales, some with dark 
setae, also present. Lowest pinnae not seen, 
longest 65 cm long. Largest pinnules 75-105 by 
15-17 mm, sessile, lowest segment not free; 
costules 3-31/2 rnm apart; veins to 13 pairs; 
lamina-segments firm, falcate, crenate-serrate. 
Sori near costules; indusium a rather pale thin cup 
about as wide as deep; paraphyses shorter than 
sporangia. Scales and hairs: pinna-rachis pale 
beneath and covered sparsely with very small 
pale ± bullate scales which end in dark setae, 
scattered narrow dark scales with long marginal 
setae also present; scales on lower surface of 
costae rather broad, more or less buUate-based, 
acuminate, with scattered long marginal setae, 
also pale small bullate scales mostly lacking 
setae; costular scales pale, bullate, larger ones acu- 
minate and sometimes with setae; all scales on costae 
and costules leaving very short hair-like bases when 
they fall; veins bearing scattered small pale scales 
on lower surface and many very short hairs. 

Type specimen: Pullen 562, Upper Omahaiga 
valley Goroka District, Territory of New Guinea 
(CANB, type; BM, L). 

Distr. Malaysia: N.E. New Guinea (two col- 
lections). 



Ecol. At 2300-2500 m in Podocarpus-Lauraceae 
forest. 

12. Cyathea rigcns Rosenst. in Fedde, Rep. 12 
(1913) 163; V. A. v. R., Handb. Suppl. (1917) 33; 
Copel. Philip. J. Sc. 77 (1947) 103, 104. 

Trunk 3-5 m tail; fronds numerous, spreading, 
100-1 50(-200) cm long. Stipe 7-17 cm, near base 
bearing many spines to 2 or 3 mm long; scales pale 
or dark, shining, with dull edges which often have 
long dark setae near base, 20-50 by 3-4 mm. 
Lo-wer pinnae gradually reduced, or a gap between 
the small basal ones and the next; lowest 5-8 cm 
long, largest pinnae c. 30 cm. Largest pinnules 
40-60 by 12-1 4 mm, deeply lobed, lowest segment 
not free; costules 3-3 1/2 mm apart; veins to 8 pairs, 
mostly forked; lamina-segments crenate. Sori near 
costules; indusium a firm light brown shallow 
cup with even rim. Scales and hairs: pinna-rachis 
very scaly when young, larger scales narrow, flat, 
acuminate with a few dark marginal setae, these 
scales mostly caducous; towards apex of pinna- 
rachis some more persistent buUate-based scales; 
also on lower surface of pinna-rachis small scales 
bearing very fine crisped hairs; scales on costae 
near base narrow, flat, pale, acuminate with a 
few dark setae, distally bullate with long hair- 
point; on costules bullate hair-pointed scales 
grading to very fine pale hairs. 

Type specimen: Keysser B79, Bolan Mts, E. 
New Guinea (S-PA; dupl. at B, UC). 

Distr. Malaysia: Central and Eastern New 
Guinea, Goodenough Island. 

Ecol. At 2000-2800 m on the mainland of 
New Guinea; on Goodenough Island at 1570-1600 
m;inoak forest and mossy forest, locally abundant; 
also reported in undergrowth on edge of Notho- 
fagus forest, and on edge of grassland. 

13. Cyathea everta Copel. Un. Cal. Publ. Bot. 18 
(1942) 218; Philip. J. Sc. 77 (1947) 103, pi. 2.— 
C. globosora Copel. Un. Cal. Publ. Bot. 18 (1942) 
218; Philip. J. Sc. 77 (1947) 106, pi. 4. 

Trunk up to 8 mm, 5-6 cm 0, bearing fronds 
120-150 cm long. Stipe 20-30 cm, spiny through- 
out, spines 1-2 mm long; scales near base as in 
C rigens; upper part of stipe bearing a thin felt of 
irregular small pale fringed scales; sometimes a 
single isolated small pinna near base of stipe. 
Lowest pinnae 12-15 cm long, longest 30 cm. 
Pinnules 35-50(-75?) by 11-15 mm, lobed almost 
to costa, lower segments of larger pinnae separated 
by a narrow wing along costa, not truly free; 
costules 31/2 mm apart; veins to 8 pairs, mostly 
forked; lamina-segments crenate. Sori near cos- 
tules; indusium a firm brown cup when mature, 
about as wide as deep. Scales and hairs: pinna- 
rachis bearing more or less abundant crisped hairs, 
also buUate-based hair-pointed scales, the scales 
often deciduous; lower surface of costae bearing 
pale acuminate bullate scales and sometimes 
crisped hairs. 

Type specimen: Brass 10712, near Lake Hab- 
bema, W. New Guinea (A; dupl. at BO, UC). 

Distr. Malaysia: Western New Guinea. 



86 



Flora Malesiana 



[ser. II, vol. P 



Ecol. At 1400-2800 m, in oak forest and mossy 
forest. 

Note. Possibly this species should be united 
with C. rigens. It is sometimes difficult to distin- 
guish minute fringed scales from separate hairs, 
and perhaps there are transitions between the two. 

14. Cyathea cincinnata Brause, Bot. Jahrb. 56 
(1920) 52; Copel. Philip. J. Sc. 77 (1947) 104. 

Sripe 30 cm, scaly throughout; scales to 20 by 
2 mm, shining brown, sometimes with a black 
median band, the dull flabelloid edges bearing 
many long setae. Lowest pinnae 15 cm long, long- 
est pinnae 52 cm. Pinnules to 80 by 18 mm, 
sessile, lobed almost to the costa; costules 31/2 mm 
apart; veins to 9 pairs; lamina-segments slightly 
toothed. Sori near the costules; indusium a firm 
brown cup with even edge, somewhat wider than 
deep; paraphyses short and slender. Scales and 
hairs: lower surface of pinna-rachis covered evenly 
with short crisped hairs, with scattered very narrow 
flexuous dark scales which bear a few dark flexuous 
marginal setae; lower surface of costae of lower 
pinnules bearing some crisped hairs near base; 
no scales seen on costae and costules. 

Type specimen: Ledermann 11279, Sepik re- 
gion, E. New Guinea (B). 

Distr. Malaysia: E. New Guinea (one collec- 
tion). 

Ecol. At 1300 m. 

15. Cyathea subtripinnata Holttum, Blumea 11 
(1962) 534. 

Trunk lYj m, bearing 18 fronds which are not 
whorled. Stipe 12 cm, densely covered with scales 
throughout, warty after fall of scales; scales to 
40 by 1 1/2 rnm wide at base, dark brown, shining, 
twisted, fragile edges narrow except near base, 
also under-coat of irregular small brown scales 
which are mostly not setiferous. Rachis dull, 
brown, glabrescent, minutely warty. Lowest 
pinnae 17 cm long, longest 40 cm. Pinnules to 
65 by 17 mm, almost sessile, the larger ones with 
c. 6 pairs of quite free tertiary leaflets, remaining 
lamina-segments more or less broadly adnate to 
costa, distal ones connected by a narrow wing; 
costules (bases of tertiary leaflets) 4 mm apart; 
veins 7-8 pairs, mostly forked, middle ones some- 
times twice forked, hardly raised on upper sur- 
face, impressed on lower surface; lamina-segments 
or tertiary leaflets rigid when dry, fertile ones lobed 
about half way to costule. Suri near costules; 
indusium a firm brown cup wider than deep; 
sporangia very numerous; paraphyses not seen. 
Scales and hairs: pinna-rachis dull, brown, gla- 
brescent except for scattered small flat roundish 
entire pale or brown scales and a few very narrow 
dark brown scales, all lacking setae; costal scales 
like those of pinna-rachis but smaller ones some- 
times convex; costules usually glabrous beneath. 

Type specimen: Schodde 1763, Mt Giluwe, 
Southern Highlands Distr., Papua (CANB). 

Distr. Malaysia: East New Guinea. 

Ecol. Margin of alpine grassland and alpine 
shrubbery at 3120 m. 



16. Cyathea orientalis (Kunze) Moore, Ind. Fil. 
(1861) 272; Mett. Ann. Mus. Bot. Lugd.-Bat. 1 
(1863) 58; Hook. & Bak. Syn. Fil. (1865) 24; 
Racib. F1. Btzg 1 (1898) 37; v. A. v. R. Handb. 
(1908) 21; Suppl. (1917) 29; Backer & Posth. 
Varenfl. Java (1939) 24. — Disphenia orientalis 
KuNZE, Bot. Zeit. 6 (1848) 283 {excl. syn.).— 
C. arborea [non (L.) Sm.] var. pallida Hassk. Obs. 
Bot. Fil. 1 (1856) 15.— Fig. 7, 9e. 

Stipe dark, to at least 50 cm, bearing copious 
spines 1 Vi rnm long when old; scales abundant, 
dark, to 35 by 3 mm, pale edges narrow, bearing 
dark setae; basal part of scale bearing superficial 
outgrowths; pneumathodes 4-7 mm long, in a 
single row. Lower pinnae somewhat reduced, 
largest 65 cm long. Pinnules almost sessile, com- 
monly 100 by 18 mm, lobed almost to the costa, 
lowest 1-2 segments almost free, apex rather 
abruptly acuminate; costules 3-31/2 mm apart; 
veins 9-10 pairs; lamina-segments falcate, apex 
blunt, edges crenate towards apex. Sori near cos- 
tules; indusium a rather thin brown cup about 
1 mm 0, slightly constricted at mouth, about as 
wide as deep; apex of receptacle level with mouth 
of indusium; paraphyses slender, short. Scales and 
hairs: lower surface of main rachis shortly spiny in 
basal part to minutely warty distally, with residual 
very small scales some of which bear short dark 
setae; similar scales on lower surface of pinna- 
rachis; costae bearing very small brown short- 
fringed scales and flat ovate to elongate entire 
scales (mostly caducous); costules of sterile pin- 
nules bearing flat to convex ovate almost entire 
uniformly brown scales. 

Type specimen: Zollinger 2538, Tengger Mts, 
E. Java ( holotype not seen; dupl. at L, P, BM, BO). 

Distr. Malaysia: W.-E. Java, Lesser Sunda Is 
(Bali, Lombok, Flores). 

Ecol. In mountain forest, 1000-1800 m. 

17. Cyathea apoensis Copel. in Elmer, Leafl. 
Philip. Bot. 3 (1910) 802; v. A. v. R. Handb. 
Suppl. (1917) 26; Copel. Fern Fl. Philip. 2 (1960) 
213.— C. lobata Copel. Philip. J. Sc. 81 (1952) 
15, p!. 13; Fern Fl. Philip. 2 (1960) 214. 

Stipe unknown. Pinnae to 40 cm long. Pinnules 
commonly to 65(-85) by 14-18 mm, lowest 1-2 
segments quite free, rest of pinnule lobed almost 
to costa; costules 3-31/2 mm apart; veins 7-9 pairs, 
mostly forked; lamina-segments rather deeply 
crenate-serrate or basal ones lobed up to 1/2 way 
to costule, each lobe bidentate. Sori near costules; 
indusium a rather thin cup with even rim (often 
broken when old), about as wide as deep; para- 
physes slender, short. Scales and hairs: main rachis 
dark, minutely warty on lower surface, bearing 
residual very small irregular dull brown scales 
which sometimes bear a dark seta, also a few 
elongate dark flat scales with pale edges; pinna- 
rachis similarly scaly, often with very narrow brown 
scales 3-5 mm long; costae near base rather co- 
piously scaly, scales to 3 mm long and -'4 mm wide, 
shining brown, flat, tapering evenly from base, 
edges with a few dark setae, distal scales similar 
but shorter, more ovate; no scales seen on costu- 



Dec. 1963] 



Cyatheaceae (Holttum) 



87 



les, but may be present on costules of sterile pin- 
nules (none such seen). 

Type specimen: Elmer 11482, Mt Apo, Min- 
danao (Mich; dupl. at US, FI, K, BO, P, A, 
SYD, L, BM). 

Distr. Malaysia: Philippines (Mindanao, Ne- 
gros, southern Luzon). 

Ecol. 'In dense woods along Seriban creek, 
Mt Apo, alt. 1800 m' (Elmer). 

18. Cyathea costalisora Copel. Un. Cal. Publ. Bot. 
18 (1942) 218; Philip. J. Sc. 77 (1947) 101, pi. 1. 

Trunk to 4 m, branching near base. Stipe warty 
after fall of scales, c. 20 cm; scales to 25 by 1 mm, 
central band dark shining brown, pale dull edges 
bearing a few long dark setae; upper part of stipe 
and rachis bearing very narrow flexuous spreading 
scales to 10 mm long. Lowest pinnae c. 12 cm 
long, longest 20-30 cm. Largest pinnules 35^5 
by 8-10 mm, lobed nearly to costa, lowest 1-2 
segments sometimes a little constricted at base, 
not free, apex abruptly pointed; costules 2-2 1/2 
mm apart; veins 7-8 pairs, mostly forked; lamina- 
segments firm, crenate. Sori near costules (on 
type only on basal veins and so only near costa); 
indusium at maturity a firm dark cup with slightly 
contracted mouth, wider than deep, facing ob- 
liquely away from costule. Scales and hairs: pinna- 
rachis densely scaly on lower surface, scales 4-5 
mm long with dark median band and broad pale 
edges bearing a few long setae, also smaller pale 
scales with some setae; costae and costules of 
type almost glabrous on lower surface. 

Type specimen: Brass 9488, Lake Habbema, 
W. New Guinea (A; dupl. at BO, BM). 

Distr. Malaysia: Western New Guinea (2 col- 
lections). 

Ecol. In forest: near Lake Habbema (3225 m) 
in moist hollows; in Arfak Mts (1900 m) near 
Lake Giji. 

Note. The specimen from Arfak Mts differs 
from the type as follows: stipe-scales IV2 mm wide, 
pale or dark near apex; on lower surface of costae 
and costules are narrow pale scales bearing a 
few dark marginal setae; lamina-segments deeply 
serrate. 

19. Cyathea pallidipaleata Holttum, Kew Bull. 
16 (1962) 60. 

Stipe 10 cm, medium brown, densely covered 
with shining pale brown scales to 32 by 1 mm, 
their dull edges narrow; similar smaller scales 
at first all along rachis but caducous. Lowest 
pinna 13 cm long, longest pinna 25 cm; pinnules 
at 60' to pinna-rachis. Pinnules to 45 by 1 1 mm, 
abruptly acuminate, lobed nearly to costa, lowest 
segment nearly free; costules 2'/2 mm apart; 
lamina-segments thick and rigid, edges not re- 
flexed, deeply crenate, the larger crenations 
notched; veins 7-8 pairs. Sori near costules; 
indusium a complete firm dark open cup. Scales 
and hairs: on pinna-rachis many very small dark 
short-setiferous scales and some residual long 
dark scales; on costae shining dark brown flat ovate 
scales bearing a few setae; on costules many very 
small dark setiferous scales, these mostly caducous. 



Type specimen: Eyma 776, Tinabang, W. side 
of Mt Rante Mario, Subdistr. Enrekang, SW. 
Celebes (BO). 

Distr. Malaysia: SW. Celebes: Latimodjong 
Mts. 

Ecol. Mountain forest, 3000 m. 

20. Cyathea coactilis Holttum, Blumea II (1962) 

533. 

Trunk IVi m, bearing 10 fronds in 2 whorls; 
fronds c. 170 cm long. Stipe c. 10 cm, not spiny, 
densely covered with scales throughout; scales to 
35 by less than 1 mm wide at base, pale with darker 
narrow fragile edges; under-coat of small dull 
appressed fringed scales, some with apical seta. 
Rachis persistently covered on lower surface with 
a thin felt of small scales as stipe with a few long 
narrow pale scales. Lowest pinna c. 6 cm long, 
longest 30 cm. Largest pinnules sessile, abruptly 
acuminate, 37 by 12 mm, lowest pair of segments 
almost or quite free, then a few pairs joined by 
narrow wing along costa; costules 3 mm apart; 
veins 6-7 pairs; lamina-segments firm, crenate. 
Sori near costules; indusium a thin cup, wider 
than deep; paraphyses slender, dark. Scales and 
hairs: pinna-rachis covered completely with a felt 
of scales which are small, light brown, thin, 
copiously short-fringed, not bullate nor acuminate 
but sometimes with dark setiform apex, also a 
few long narrow pale scales with fragile non- 
setiferous edges; costae bearing larger dark flat 
scales with irregular marginal setae; costular 
scales flat, brown, usually not setiferous, distal 
ones very small; on lower surface of veins very 
short hairs (bases of scales?). 

Type specimen: Schodde 1887, Mt Giluwe, 
Southern Highlands Distr., Papua (CANB). 

Distr. Malaysia: E. New Guinea (once col- 
lected). 

Ecol. In alpine shrubbery at 3000 m. 

21. Cyathea parva Copel. Un. Cal. Publ. Bot. 
18 (1942) 219; Philip. J. Sc. 77 (1947) 120, pi. 
13. 

Trunk I'i m tall. 3V2 cm o, bearing 4 fronds 
90 cm long. Stipe c. 10 cm, copiously warty; 
scales to 15 by IV2 mm, pale or sometimes dark 
near apex, the thin dull edges bearing a few long 
dark setae. Lower pinnae gradually reduced, 
lowest 5-6 cm long, longest 17 cm, pinnules well 
spaced. Pinnules to 30 by 6 mm, not acuminate, 
lobed almost to costa, lowest segment not free; 
costules 1 '/2-2 mm apart; veins to 6 pairs, lower 
ones forked; lamina-segments firm, the fertile 
ones crenate, sterile almost entire. Sori near 
costules; indusium a very narrow dark red ring 
round base of receptacle. Scales and hairs: lower 
surface of main rachis almost completely covered 
with very small dull long-fringed scales, with 
scattered large pale scales; on lower surface of 
pinna-rachis many pale thin-edged scales 5 by I 
mm, long-acuminate, sometimes bearing long dark 
marginal setae, these scales bullate at base near 
apex of pinna, also many minute scales bearing 
long marginal hairs; costal scales pale, narrow, 



88 



Flora Malesiana 



[ser. II, vol. 12 



with bullate bases, grading to rather large pale 
buUate scales. 

Type specimen: Brass 12197, 15 km SW of 
Bernhard Camp, Idenburg River, W. New 
Guinea (UC; dupl. at MICH, BO, A, L). 

Distr. Malaysia: Western New Guinea (one 
collection). 

Ecol. In undergrowth of rain forest in gully, 
1700 m. 

22. Cyathea wengiensis (Brause) Domin, Pterid. 
(1929) 263; Copel. Philip. J. Sc. 77 (1947) 113.— 
Alsophila wengiensis Brause, Bot. Jahrb. 49 
(1912) 13, fig. Ic; v. A. v. R. Handb. Suppl. 
(1917J 68. — Alsophila hieronymi Brause, Bot. 
Jahrb. 49 (1912) 14; v. A. v. R. Handb. Suppl. 
(1917) 67.— C. braitseana Domin, Pterid. (1929) 
262. 

Stipe to 10 cm or more, bearing spines to 2 mm 
long; scales not seen. Lowest pinna 10 cm long, 
longest 48 cm. Largest pinnules 70 by 12-18 mm 
(sometimes larger; see note below); lobed to c. 1 
mm from costa, lowest segment not quite free; 
costules 3 1/2 rnm apart; veins 9-11 pairs; lamina- 
segments firm, edges more or less crenate, sinuses 
narrow. Sori medial, or distal ones near costule; 
indusium a dark ring less than 1/2 rnm o round 
base of receptacle, sometimes not quite encircling 
receptacle; paraphyses longer than sporangia. 
Scales and hairs: on lower surface of pinna - 
rachis crisped light brown hairs, or small scales 
bearing such hairs, throughout but more abundant 
distally. also dark scales to 3 mm long with broad 
pale edges bearing long dark setae; on costae 
similar hairs or hair-bearing scales and scattered 
broad scales bearing few setae; on costules flat 
pale scales to bullate scales. 

Type specimen: Schlechter 16100, in forest 
near Wengi, NE. New Guinea (B; dupl. at P, 
UC). 

Distr. Malaysia: Eastern New Guinea. 

Ecol. In forest or secondary growth after 
cultivation, up to 600 m. 

Notes. A specimen collected from 'tall garden 
re-growth' in Sepik District (Darbyshire & 
Hoogland 8042) agrees with the type except in 
larger size of pinnules, largest being 120 by 23 mm 
with costules 5-5 '/i mm apart and sterile segments 
very deeply lobed. This large size may be due to 
conditions of habitat. 

In position of sori C wengiensis agrees with 
C. macgillivrayi (Bak.) Do.min, but normally the 
latter has no indusium. Specimens with variable 
very small indusia which are mostly hemitelioid 
appear intermediate between the two species. 

23. Cyathea batjanensis (Christ) Copel. Philip. J. 
Sc. 4 (1909) Bot. 45. — Alsophila batjanensis 
Christ in Warb. Monsunia (1900) 90; v. A. v. R. 
Handb. (1908) 38. — Alsophila saparuensis v. A. 
V. R. Bull. Dep. Agr. Ind. Neerl. n. 18 (1908) 2; 
Handb. (1908) 38. — Alsophila straminea Gepp in 
Gibbs. Arfak (1917) 192.— C. saparuensis v. A. 
V. R. Bull. Jard. Bot. Btzg II, n. 28 (1918) 13.— 
C. straminea v. A. v. R. I.e. 14 (non Karst. 



1856). — C. geppiana Domin, Acta Bot. Bohem. 9 
(1930) 118.— Fig. 9a. 

Trunk slender. Stipe 15 cm or more long, 
bearing many warts or conical spines 1 mm 
high; scales dark brown with paler fragile edges. 
Lower pinnae often considerably reduced (to 
10 cm long or less), longest to 45 cm. Largest 
pinnules 70-90 by 15-18 mm, lobed almost to 
the costa; costules 3'/2-5 mm apart; veins to 10 
pairs; lamina-segments rather thin, edges rather 
strongly crenate towards apices. Sori medial; 
indusium a small ring round base of receptacle; 
paraphyses longer than sporangia, several cells 
wide at the base. Scales and hairs: lower surface of 
pinna-rachis and costae bearing flexuous pale 
hairs and scattered irregular small pale scales 
with long flexuous slender marginal hairs; no bul- 
late scales on costae or costules. 

Type specimen: Warburg 17844, Mt Sibela, 
Batjan, Moluccas (B; not seen at Paris). 

Distr. Malaysia: Moluccas (Batjan, Ceram, 
Tidore, Ternate, Buru, Saparua, Halmaheira); 
Western New Guinea. 

Ecol. At about 600 m. 

24. Cyathea ternatea v. A. v. R. Bull. Jard. Bot. 
Btzg III. 5 (1922) 191. 

Stipe bearing many spines to Ih'j mm long; 
scales to 25 by 3 mm, shining brown with narrow 
dull edges. Pinnae to 65 cm long. Pinnules to 150 
by 22 mm, long-acuminate, lowest on stalks 2-3 
mm long, lowest segment almost free; costules 
4V2 rnm apart; veins to 1 1 pairs, mostly forked 
(some twice forked); lamina-segments firm, 
rather strongly crenate throughout. Sori medial; 
indusium a small disc round base of receptacle; 
paraphyses long. Scales and hairs: pinna-rachis 
and costae bearing many very small irregular 
shortfringed scales, also on pinna-rachis a few 
broad thin scales having odd marginal setae; 
on costules sometimes brown bullate scales 
which are ciliate towards acuminate apices. 

Type specimen: Beguin 1126, Ternate (BO; 
dupl. at L, P). 

Distr. Malaysia: Moluccas (Ternate, 2 col- 
lections). 

Ecol. At 600-1300 m. 

25. Cyathea albidosquamata Rosenst. in Fedde, 
Rep. 12 (1913) 525; v. A. v. R. Handb. Suppl. 
(1917) 31; Copel. Philip. J. Sc. 77 (1947) 104, 
121.— C. pumilio v. A. v. R. Bull. Jard. Bol. 
Btzg II. n. 28 (1918) 14. 

Stipe to at least 20 cm, scaly throughout, scales 
firm, pale, shining, with dull fragile edges; on lower 
surface of rachis similar scales to 7 by 1 mm, 
bearing a few long dark setae, also a close cover of 
pale flexuous hairs. Pinnae to 35 cm long. Largest 
pinnules 32^0 by 12 mm, lobed almost to costa; 
costules 3-3 '/2 mrn apart; veins 4-6 pairs; lamina- 
segments firm, slightly crenate. Sori near costules; 
indusium an almost flat disc c. 1 mm wide, some- 
times slightly asymmetric; paraphyses not longer 
than sporangia. Scales and hairs: on lower sur- 
face of pinna-rachis a close cover of pale hairs. 



Dec. 1963] 



Cyatheaceae (Holttum) 



89 



also narrow pale acuminate scales, all except the 
largest bullate at base; on costae similar hairs 
and scales, distal scales bullate; on costules small 
pale bullate scales and pale crisped harirs. 

Type specimen: Keysser 177, Sattelberg, NE. 
New Guinea (S-PA; dupl. at B, UC). 

Distr. Malaysia: Moluccas (Ceram), New 
Guinea. 

Ecol. At 1200-1500 m. 

Note. The type of C. pumilio, from Ceram 
(RuTTEN 373) differs from the type of C. albi- 
dosquamata chiefly in having more sparse and 
hardly bullate scales on the pinna-rachis. 

26. Cyathea javanica Bl. En. PI. Jav. (1828) 245 
{incl. var. rigida, p.p.); Mett. Ann. Mus. Dot. 
Lugd.-Bat. 1 (1863) 56; Hook. Syn. Fil. (1865) 
23; Racib. F1. Btzg I (1898) 35; v. A. v. R. Handb. 
(1908) 23; Suppl. (1917) 32; Backer & Posth. 
Varenfl. Java (1939) 24. — Hemitelia caudipinnula 
V. A. V. R. Bull. Jard. Bot. Btzg II, n. 1 (1912) 
16; Handb. Suppl. (1917) 42,.— Hemitelia ba- 
risanica v. A. v. R. Bull. Jard. Bot. Btzg II. 
n. 20 (1915) 17; Handb. Suppl. (1917) 49.— 
Alsophila benciilensis v. A. v. R. Bull. Jard. Bot. 
Btzg II. n. 23 (1916) 2; Handb. Suppl. (1917) 493. 
— Alsophila palembanica v. A. v. R. Bull. Jard. 
Bot. Btzg II, //. 23 (1916) 4; Handb. Suppl. (1917) 
493. — C. benculensis v. A. v. R. Bull. Jard. Bot. 
Btzg II, «. 28 ( 1 9 1 8 ) 1 4.— C. palembanica v. A. v. R 
I.e. 13.— C. caudipinnula Domin, Pterid. (1929) 
263. — C barisanica Domin, I.e. — Fig. 9b. 

Trunk to 10 m. Stipe 10-30 cm, bearing many 
spines to I mm long, and dark fragile-edged 
scales to 15 by 1 mm. Lower pinnae gradually 
reduced, lowest 5-10 cm long where stipe is 10 cm, 
longer where stipe is longer; longest pinnae to 
70 cm. Largest pinnules sessile, caudate-acuminate, 
80-100 by 15-23 mm, lobed almost to costa, lowest 
segments not free; costules 4-5 mm apart; veins to 
10 pairs; lamina-segments firm, slightly crenate or 
the largest fertile ones sometimes deeply so, 
sinuses narrow except near base of largest fertile 
pinnules. Sori near costules except the basal ones; 
indusium a rather firm disc about as wide as base 
of mature sorus, sometimes excentric or almost 
entirely on costular side of receptacle; paraphyses 
slender, short. Scales and hairs: lower surface of ra- 
chis and pinna-rachis closely hairy throughout, 
hairs short, crisped, with some residual narrow 
dark scales; on lower surface of costae similar 
hairs, also narrow scales near base grading to 
buUate-acuminate scales distally; on costules small 
brown bullate scales; upper surface of pinna- 
rachis and costae usually glabrous, sometimes 
slightly hairy. 

Type specimen: Blume, West Java (L; dupl. K). 

Distr. Malaysia: Sumatra, Java. 

Ecol. In forest, 250-1500 m. 

Note. The type specimen of Hemitelia bari- 
sanica, from Sumatra, is unusually large, with 
pinnules to 120 by 28 mm, costules to 6 mm apart; 
the indusia are usually excentric and sometimes 
only on the costular side of the receptacle; hairs 
and scales agree with type. 



Some specimens at Kew of C. javanica var. 
rigida so named by Blume are C. crenulata Bl. 

27. Cyathea hymenodes Mett. Ann. Mus. Bot. 
Lugd.-Bat. 1 (1863) 57; Hook. Syn. Fil. (1865)24; 
V. A. V. R. Handb. (1908) 24.— C. korthalsii 
Mett. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 57; 
Hook. Syn. Fil. (1865) 25; v. A. v. R. Handb. 
(1908) 21; Suppl. (1917) 27, p.p. —C.amphieos- 
mioides v. A. v. R. Bull. Jard. Bot. Btzg III, 
2 (1920) 138.— C. arthropterygia v. A. v. R. 
ibid. Ill, 5 (1922) 188.— C. latebrosa var. indusiata 
Holttum, Gard. Bull. S.S. 8 (1935) 305; Rev. 
Fl. Mai. 1 (1954) 121. 

Stipe not spiny, sometimes with a pair of short 
pinnae near base; scales 20-35 by 1 '/2-4 mm, 
dark, fragile edges soon disappearing. Lower 
pinnae more or less reduced; longest 50 cm long. 
Largest pinnules sessile, 70-100 by 15-20 mm, 
lowest segment (rarely several pairs of segments) 
almost free and often separated from next by a 
narrow wing along costa, rest of pinnule lobed 
almost to costa; costules 3^ mm apart; veins 
8-10 pairs; lamina-segments rather thin, more or 
less deeply crenate (lowest sometimes deeply 
lobed). Sori near costules; indusium an almost 
circular brown disc about as wide as base of sorus, 
its edge entire, often somewhat asymmetric about 
the receptacle, sometimes only on the costular 
side; receptacle swollen, paraphyses short, apical 
ones sometimes flat at the base. Scales and hairs: 
short crisped hairs more or less abundant on distal 
part of lower surface of pinna-rachis; scales near 
bases of costae elongate, usually entire but some- 
times with a marginal seta, grading to bullate 
scales distally; bullate scales on costules, often 
deciduous from fertile pinnules; upper surface of 
costae always hairy. 

Type specimen: Korthals, Sumatra (L). 

Distr. Malaysia: Sumatra, Malay Peninsula. 

Ecol. In mountain forest at 900-2000 m. 

Note. Mettenius described the indusia of both 
C. hymenodes and C. korthalsii as breaking 
(fatiscens) but they are in almost all cases flat 
discs with an entire margin which gives no sign 
of having been formed by the breakdown of a 
complete indusium. In both types some of the 
indusia are hemitelioid, some are symmetrical 
about the receptacle as centre. Probably the 
hemitelioid indusia are more abundant in the 
type of C. korthalsii. 

28. Cyathea patellifera v. A. v. R. Bull. Jard. Bot. 
Btzg II, n. 16 (1914) 4; Handb. Suppl. (1917) 31. 

Stipe to 50 cm, dark at base, red-brown up- 
wards, spiny throughout, spines near base 2 mm 
long, distally less than 1 mm, scaly near base; 
scales to 20 by 2 mm, firm, medium brown, fragile 
edges early caducous. Middle pinnae 35^0 cm 
long. LsiTgesi pinnules 50-70 by 15 mm, abruptly 
pointed or short-acuminate, almost sessile, lowest 
pair of segments quite free, then 1 to several pairs 
constricted on acroscopic side at base, connected 
to rest by a narrow wing on costa; costules 3 mm 
apart; veins 6-8 pairs; lamina-segments firm, more 



90 



Flora Malesiana 



[ser. II, vol. P 



or less deeply crenate, free ones sometimes lobed 
half-way to costule. So/7 near costules; indusium 
a firm brown disc hardly as wide as base of ripe 
sorus, its edges rather irregularly lobed; paraphy- 
ses as long as sporangia, abundant, some of them 
2 cells wide at base. Scales and hairs: lower 
surface of pinna-rachis covered throughout with 
short crisped hairs, more copiously towards apex; 
scales near base of costa elongate, brown, entire, 
some with bullate base, grading to brown bullate 
scales; scattered short crisped hairs also on lower 
surface of costae; small entire brown bullate 
scales on costules. 

Type specimen: Matthew 667, Mt Singgalang, 
Sumatra (BO; dupl. at K). 

Distr. Malaysia: Central Sumatra (2 collec- 
tions). 

Ecol. In forest, 2200-2400 m. 

Notes. The second collection (Schiffner 
P223), from same locality, has longer pinnules 
than the type, with fewer free or sub-free basal 
segments, segments less deeply lobed, and scales 
paler and less rigid than those of the type. Though 
the indusium has in all cases an irregular margin, 
there is no indication that it represents the re- 
mains of a larger indusium that previously covered 
the sorus. 

29. Cyathea negrosiana Christ, Philip. J. Sc. 2 
(1907) Bot. 181; v. A. v. R. Handb. (1908) 786; 
COPEL. Fern Fl. Philip. 2 (1960) 214. 

Stipe c. 15 cm long, warty; scales to 20 by 1 Vi 
mm, dark brown, shining, with narrow dull con- 
colorous edges. Lowest pinna 5-6 cm long, longest 
pinna 35 cm. Largest pinnules 70 by 14 mm, 
sessile, acuminate, lowest 1-2 segments free or 
nearly so, rest of pinnule lobed nearly to costa; 
costules 3 mm apart; veins 8-10 pairs; lamina- 
segments rather thin, slightly crenate, sinuses 
narrow. Sori near costules; indusium ultimately a 
rather thin brown disc usually widest on the 
costular side, where it is conspicuous, sometimes 
with uneven edge and possibly the remains of a 
former complete indusium; receptacle very 
prominent; paraphyses shorter than sporangia. 
Scales and hairs: lower surface of pinna-rachis 
covered with very numerous small pale bullate 
scales, hairs lacking; lower surface of costae and 
costules bearing similar but smaller scales, some 
elongate almost flat entire scales also near base of 
costae. 

Type specimen: Whitford, F. B. 1536, Mt 
Silay, Negros (P; dupl. at MICH). 

Distr. Malaysia: Philippines (Negros, Leyte, 
Biliran). 

Ecol. At 1000 m. 

30. Cyathea catillifera Holttum, Kew Bull. 16 
(1962) 53. 

Trunk branching at base, 1 m by 10 cm 0, 
bearing about 6 fronds, stipe-bases persistent. 
Stipe 50 cm, base dark brown, paler upwards, 
closely spiny near base, minutely warty between 
spines, the latter rather slender, to 3 mm long; 
scales near base of stipe to 20 by 2-21/2 mm. 



shining dark brown with concolorous thinner 
edges; very small irregular brown scales on surface 
between thorns. Lowest pinna 23 cm long, longest 
pinna 29 cm. Largest pinnules 55 by 15 mm, 
lowest segment not free, apex acuminate; costules 
3-3'/^ mm apart; veins to 8 pairs; lamina-segments 
rather rigid, edges deeply crenate when fertile. 
Sori near costules; indusium a dark brown disc 
c. 1 mm wide, symmetric about the receptacle or 
slightly wider on costular side, with thin paler 
edges; paraphyses slender, as long as sporangia. 
Scales and hairs: lower surface of main rachis light 
brown, dull, minutely warty, glabrescent; lower 
surface of pinna-rachis similar near base but 
bearing scattered very narrow long entire brown 
scales, distal half more or less covered with small 
light brown entire more or less bullate scales; 
costae rather densely scaly throughout lower 
surface, scales mostly ovate in outline, larger ones 
bullate, shining light brown, smaller ones of all 
sizes; costules bearing similar bullate scales and 
sometimes very narrow entire scales; veins bearing 
neither scales nor hairs. 

Type specimen: Brass 4549, Murray Pass, 
Wharton Range, Central Division, Papua (BRI; 
dupl. at NY). 

Distr. Malaysia: Eastern New Guinea (one 
collection). 

Ecol. At 2840 m. 

31. Cyathea horridula Copel. Un. Cal. Publ. Bot. 
18 (1942) 219; Philip. J. Sc. 77 (1947) 111, pi. 8. 

Trunk 3 m, slender. Stipe 40 cm, bearing many 
short spines (to 1 mm long), scaly near base; 
scales to 15 by 1 V2 mm, pale, with distinct fragile 
edges bearing occasional long dark setae. Lowest 
pinna 25 cm long, longest 45 cm. Pinnules to 
55 by 13 mm, lowest 1-2 pairs of segments sepa- 
rately adnate to costa, rest of pinnule lobed 
nearly to costa; costules 3 mm apart; veins to 9 
pairs; lamina-segments rather thin. Sori near 
costules; indusium a very small dark brown disc, 
somewhat irregular in shape, receptacle usually 
excentric. Scales and hairs: lower surface of pinna- 
rachis bearing elongate pale scales having long 
dark setae, also many very small pale fringed 
scales; scales on costae elongate, pale, with some 
long setae, grading to pale bullate scales lacking 
setae distally and on costules, very small scales 
bearing long slender hairs also present. 

Type specimen: Brass 12043, near Idenburg 
River, W. New Guinea (UC; dupl. at MICH, A). 

Distr. Malaysia: Western New Guinea (one 
collection). 

Ecol. At 1700 m. 

32. Cyathea tenuicaulis Domin. Acta Bot. Bohem. 
9 (1930) 165; Copel. Philip. J. Sc. 77 (1947) 113. 
— Alsophila tenuis Brause, Bot. Jahrb. 56 (1920) 
71, non C. tenuis Brause, 1911. 

Trunk 1-2 m, 2 cm o, bearing fronds 75 cm long. 
Stipe 18-40 cm, dull purplish, warty; scales to 
8 by 1 mm, rigid, medium brown, fragile edges 
soon abraded. Lowest pinna 7-1 1 cm long, longest 
21 cm. Largesl pinnules 40-50 by 13-15 mm, lobed 



Dec. 1963 ] 



Cyatheaceae (Holttum) 



91 



to 1-2 mm from costa, no free basal segments; 
costules 3 mm apart (fully fertile pinnules) to 4 mm 
(sterile); veins 7-9 pairs; lamina-segments entire 
or slightly crenate. Sari near costules; indusium 
very thin and fragile, appearing as an irregular 
fragment on old sori; receptacle swollen, para- 
physes thin and short. Scales and hairs: lower 
surface of pinna-rachis bearing rather sparse short 
crisped hairs, also scattered elongate dark scales 
bearing marginal setae, and distally pale bullate 
scales; lower surface of costae bearing crisped 
hairs and sometimes long setiferous scales near 
base, also small pale bullate scales throughout; 
bullate scales also on costules. 

Type specimen: LEDERiVANN 7498, Sepik region, 
NE. New Guinea (B). 

Distr. Malaysia: Eastern New Guinea. 

Ecol. In forest 300-1500 m. 

Notes. The type specimen is from an old frond; 
no remnants of indusia have been seen on it. 
Specimens collected by Carr (13363, 14542), 
which agree in scaliness and form of pinnules with 
the type, show fragments of indusia on most sori. 
If the type is truly lacking in indusia, Carr's 
specimens should probably be regarded as re- 
presenting a distinct species. 

33. Cyathea sumatrana Bak. J. Bot. 18 (1880) 
209; V. A. V. R. Handb. (1908) 23; Suppl. (1917) 
32.— C. schizochlamys Bak. J. Bot. 18 (1880) 209; 
V. A. V. R. Handb. (1908) 25.— C. subuliformis 
V. A. V. R. Bull. Jard. Bot. Btzg II, /;. 1 1 (1913) 6; 
Handb. Suppl. (1917) 32. — C tubercitlata v. A. v. 
R. Bull. Jard. Bot. Btzg II, n. 28 (1918) 11. 

Stipe to 60 cm or more, warty, copiously scaly 
throughout; scales to 25 by 1-3 mm, dark to 
medium brown, shining, with paler fragile edges. 
Longest /7/«nae 50 cm long. Pinnules to 85 by 1 8 mm, 
lowest 1-2 segments of larger pinnules free and 
sometimes deeply lobed, rest of pinnule lobed 
almost to costa; costules 3-4 mm apart; veins 
9-12 pairs, those in lobes of free segments forked 
twice or more; lamina-segments firm, crenate. 
Sori near costules; indusium at first completely 
covering sorus, thin and fragile, breaking and 
partly caducous at maturity; receptacle swollen; 
paraphyses slender, shorter than sporangia. Scales 
and hairs: lower surface of pinna-rachis covered 
throughout with short flexuous brown hairs, also 
when young with narrow flexuous brown scales 
5-10 mm long often bearing long dark setae on 
their paler edges, those near apex of pinna with 
bullate base; lower surface of costae bearing bullate 
scales throughout, those near base of costae having 
long acuminate apices, some crisped hairs also 
present; bullate scales present on costules; upper 
surface of costae bearing dark antrorse hairs 
throughout (diff'erence from C. javanica). 

Type specimen: Beccari 438, Mt Singgalang, 
Sumatra (K; dupl. at FI). 

Distr. Malaysia: Sumatra, Malay Peninsula. 

Ecol. In forest, 500-1500 m. 

34. Cyathea klossii Ridl. Trans. Linn. Soc. II, 
Bot. 9 (1916) 251; v. A. v. R. Handb. Suppl. 
(1917J 489. 



Trunk 1 '/2 cm 0. Stipe 10 cm; scales dark, dull, 
thick, to 8 by hardly 1 mm, with a few long mar- 
ginal setae towards apex. Frond c. 60 cm long, 
simply pinnate; pinnae close and spreading 
throughout, several lower pairs gradually reduced, 
lowest 3 cm long, longest 70 by 18 mm, lobed to 
c. 1 '/2 iT>ni from costa; costules of pinna-lobes to 
3'/2 mm apart; veins to 6 pairs, usually simple; 
lamina-segments entire or slightly crenate, firm. 
Sori near costules; indusium at first completely 
covering sorus, rather thin and pale but not 
translucent, breaking irregularly and in part ca- 
ducous; receptacle somewhat swollen; paraphyses 
short, slender. Scales and hairs: on lower surface 
of main rachis throughout rather copious small 
pale bullate scales, with residual very narrow dark 
scales 3-4 mm long; costae of pinnae bearing scat- 
tered small pale bullate scales on lower surface, 
hairy on upper surface near base only; no scales 
seen on costules. 

Type specimen: Kloss, Camp III, Dec. 1912, 
Mt Carstensz, W. New Guinea (BM; dupl. at K). 

Distr. Malaysia: W. New Guinea. 

Ecol. At 750 m and lower. 

Note. Kloss made two collections, one labelled 
Camp III, one Camp III-IV; the former includes 
the trunk. As the trunk is well developed, this 
appears to be a species which always has simply 
pinnate fronds. It has no obvious near relatives 
among species with bipinnate fronds in New 
Guinea. 

35. Cyathea trachypoda v. A. v. R. Bull. Jard. Bot. 
Btzg III, 5 (1922) 19\. —Alsophila alpina v. A. v. R. 
ibid. II, n. 20 (1915) 4; Handb. Suppl. (1917) 62.— 
C. alpina v. A. v. R. Bull. Jard. Bot. Btzg II, n. 28 
(1918) 13 (not C. alpina Roth, 1800).— C. alpicola 
DoMiN, Acta Bot. Bohem. 9 (1930) 89. 

Stipe at least 50 cm, bearing scattered spines to 
3 mm long, more or less completely covered 
throughout (as also lower surface of rachis) with 
a thin pale woolly layer of small finely-fringed 
scales; larger scales scattered throughout, and also 
present on rachis, to 30 by 4 mm, dark, shining 
brown with rather broad paler fragile edges, those 
on main rachis commonly 10 by 1 mm. Pinnae to 
40 cm long, l.avgt'ii pinnules 100 by 18-20 mm, ses- 
sile, lowest segment free, then 1-2 pairs adnate 
and joined by a wing along costa; costules 3 Vi rnm 
apart; veins 10-11 pairs; lamina-segments thin, 
entire or slightly crenate. Sori near costules; in- 
dusium at first complete, the top web-like, breaking 
and mostly falling away, leaving an irregular disc 
with lacerate edges; receptacle elongate and some- 
what swollen; paraphyses short. Scales and hairs: 
lower surface of pinna-rachis at first covered with 
fine woolly scales as main rachis, these more or 
less caducous, with scattered narrow entire pale to 
light brown scales; lower surface of costae bearing 
small fringed scales and a few longer narrow ones; 
costules bearing ovate to elongate flat or convex 
pale scales, their edges fragile and lacerate, grading 
to very small scales. 

Type specimen: Bunnemeuer 10205, Mt Ke- 
rintji, Sumatra (BO; dupl. at K). 



92 



Flora Malesiana 



[ser. II, vol. 12 



Distr. Malaysia: Central Sumatra. 

Ecol. In forest, 2000-2750 m. 

Notes. The type of Alsophila alpina has rachises 
almost completely covered on lower surface with 
woolly scales, but others from type locality have 
only remnants of such a covering. This species is 
very near C. creniilata, but differs in its spiny stipe. 

A specimen from Harau-Canyon, near Pa- 
jakumbuh, at 800 m ( Meijer 5282) has a spiny stipe 
with fine woolly covering, but costae and costules 
almost glabrous apart from minute long-fringed 
scales on costae, like those forming the woolly 
covering on the stipe but smaller. 

36. Cyathea crenulata Bl. En. PI. Jav. (1828) 244; 
Hook. Sp. Fil. 1 (1844) 25; Syn. Fil. (1863) 23; 
Mett. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 57; 
Racib. F1. Btzg 1 (1898) 36; v. A. v. R. Handb. 
(1908) 21; Bull. Jard. Bot. Btzg II, n. 20 (1915) 9, 
10; Handb. Suppl. (1917) 27, 488, Corr. 63, 64 
{excl. f. hitissima, f. squamidosa, f. subspimdosa); 
Backer & Posth. Varenfl. Java (1939) 26.— 
C.javanica var. rigida Bl. En. PI. Jav. (1828) 245, 
p.p. — (?) C. polvcarpa Jungh. Nat. Geneesk. 
Arch. N.I. 2 (1845) 40; Flora 30 (1847) 522.— 
C. excelsa [iwn Sw.] Kunze, Bot. Zeit. 6 (1848) 
284. — C spinulosa var. muricidata Hassk. in 
Hook. J. Bot. Kew Misc. 7 (1855) 322; v. A. v. R. 




Fig. 11. Cyathea crenulata Bl., showing how old 
fronds persist and cover upper part of trunk. 
Kawah Baru, Papandajan, Java (van Steenis). 



Handb. (1908) 25;Suppl. (1917)36.— C./eMco/7Aoej 
Hassk. in Hook. J. Bot. Kew Misc. 7 (1855) 323; 
Obs. Fil. Jav. 1 (1856) 26; v. A. v. R. Handb. 
(1908) 24. — C. zollingeriana Mett. Ann. Mus. Bot. 
Lugd.-Bat. 1 (1863) 57 (not of v. A. v. R. Handb. 
(1908) 20, 785; Suppl. (1917) 26, Corr. 42, which 
is C. oiiiops Hassk.). — C. oinops (non Hassk. )Rac. 
El. Btzg 1 (1898) 63 {'sinops'); v. A. v. R. Handb. 
Suppl. (1917) Corr. 44. — C. distans Rosenst. Med. 
Rijksherb. 31 (1917) 2.— Fig. 9f, 11. 

Stipe warty, not spiny; scales to 25 by 3 mm, 
dark with paler fragile edges; pneumathodes in 
double row; main rachis rather pale when dry, 
finely warty, glabrescent. Lower pinnae not 
greatly reduced, longest 60 cm long. Largest 
pinnules commonly 70-100 by 17-20 mm, in some 
cases 25-30 mm wide (C sinops, C. distans), lobed 
almost to the costa, the wider ones with a few pairs 
of basal segments separately adnate to costa and 
joined by a narrow wing; costules 3 '/2-5 mm apart; 
veins to 12 pairs, mostly forked, middle ones often 
2-forked; lamina-segments firm, crenate, the basal 
ones where fertile often deeply lobed. Sori near 
costules; indusium at first complete, thin, trans- 
lucent, mostly caducous after breaking, leaving an 
irregular disc with pale thin edges; receptacle 
swollen, paraphyses short. Scales and hairs: pinna- 
rachis rather pale (green when living), finely warty 
on lower surface, residual scales small, pale, with 
short fringe of hairs; similar scales abundant on 
costae, especially near base, sometimes with flat 
elongate brown scales with pale edges which may 
bear dark setae; on costules of sterile pinnules pale 
brown convex to bullate scales. 

Type specimen: Blume, Java (L; dupl. at K). 

Distr. Malaysia: Java, Lesser Sunda Is (Flores). 

Ecol. In forest, 1700-2700 m. 

Notes. C. trachypoda, C. macropoda and C. 
magnifolia of Sumatra are closely allied, and clear 
distinctions need to be established; the warty, not 
spiny, stipe of C. crenulata seems to be distinctive. 
C. sinops and C. distans were described from 
specimens with unusually wide pinnules having 
several basal segments separately adnate to the 
costa. It may be that these should rank as a 
distinct species, as C. incisoserrata Copel. is here 
ranked as distinct from C. latebrosa. 

37. Cyathea macropoda Domin, Acta Bot. Bohem. 
9 (1930) 133.— C. longipes v. A. v. R. Bull. Jard. 
Bot. Btzg III, 5 (1922) 189 {non Copel. 1917). 
Stipe nearly 100 cm, dark at base and armed 
with spines to 5 mm long, paler distally with very 
short spines, pneumathodes in a single row, well 
spaced; scales not seen; rachis pale, lower surface 
glabrescent and somewhat warty. Lower pinnae 
slightly reduced, longest 50 cm long. Pinnules to 
70 by 17 mm, abruptly short-acuminate to cau- 
date, 1-2 basal segments free, rest of pinnule 
lobed almost to costa; costules 31/2-^ miri apart; 
veins 10 pairs; lamina-segments firm, very slightly 
crenate. Sori near costules; indusium at first 
complete, very thin over apex of sorus, breaking 
and falling, leaving an irregular disc. Scales and 
hairs: pinna-rachis beneath smooth, pale, gla- 



Dec. 1963] 



Cyatheaceae (Holttum) 



93 



brescent; scales on costae few, those near base 
narrow, dark with pale edges bearing long dark 
setae, grading to ovate fiat pale scales bearing a 
few short marginal hairs distally and on costules; 
no bullate scales seen. 

Type specimen: Bunnemeuer 9642, Mt Kerintji, 
Sumatra (BO; dupl. at L, U, US, A). 

Distr. Malaysia: Central Sumatra (3 collec- 
tions). 

Ecol. At 2000-2400 m. 

38. Cyathea saccata Christ, Ann. Jard. Bot. Btzg 
19 (1904) 42; v. A. v. R. Handb. (1908) 19. 

Stipe 25 cm long, medium brown, bearing slen- 
der spines 3 mm long; scales not seen. Pinnae to 
at least 43 cm long. Pinnules to 80 by 17 mm, 
lowest on stalks 2' 2 rnm long, basal segments not 
free, apex caudate-acuminate; costules 4-41/2 mm 
apart; veins 8-9 pairs; lamina-segments rather 
thin, light green, conspicuously crenate. Sori near 
costules; indusium complete, thin, translucent, 
breaking and caducous except for a brown disc 
at the base. Scales and hairs: pinna-rachis light 
brown, shortly spiny near base, glabrescent; on 
lower surface of costae some very small pale 
scales with long crisped pale fringing hairs, also 
elongate flat pale scales grading to rather sparse 
pale bullate scales on distal part of costae and on 
costules. 

Type specimen: P. & F. Sarasin 2045, Mt 
Topapu, Central Celebes (P; dupl. at BAS). 

Distr. Malaysia: Celebes. 

Ecol. At 13()0-1700m. 

39. Cyathea magnlfolia v. A. v. R. Bull. Jard. Bot. 
Btzg III, 2 (1920) 135.— C. acanthopodax. A. v. R. 
ibid. Ill, 5 (1922) 190. 

Stipe dark with spines to 5 mm long; scales not 
seen. Largest pinnae c. 100 cm long. Largest 
pinnules 150-175 by 30-40 mm, lobed almost to 
costa; costules 4-5 ',4 mm apart; veins c. 12(-14) 
pairs; lamina-segments firm, crenate. Sori near 
costules, at first covered with a thin complete 
indusium which breaks at maturity and is in part 
caducous, leaving an irregular disc with pale thin 
edges. Scales and hairs: pinna-rachis smooth and 
glabrous beneath; costae bearing a few small pale 
fringed scales; on costules a few entire bullate or 
strongly convex scales. 

Type specimen: Bunnemeuer 4175, Mt Ma- 
lintang, Sumatra (BO; dupl. at L). 

Distr. Malaysia: Sumatra. 

Ecol. At 1100-2000 m. 

Note. This species is near C. crenulata Bl., 
differing in large size, long spines on stipe and 
very slight scaliness. 

40. Cyathea acanthophora Holttum, Kew Bull. 1 6 
(1962) 51. 

Stipe to 80 cm; base dark with rather slender 
spines 4—7 mm long; scales early caducous, those 
on a young frond to 20 by 1 mm, medium shining 
brown with narrow dull concolorous edges lacking 
setae; pneumathodes 14-17 mm long, in a single 
row, rather widely spaced. Pinnae to 65 cm long. 



Pinnules more or less articulate to rachis, largest 
85-100 by 15-18 mm, lowest segment contracted 
at base but not free, rest of pinnule lobed almost 
to the costa; costules 4-4'/2 mm apart; veins 9-10 
pairs; lamina-segments rather thin, more or less 
crenate, separated by sinuses '/a width of segments. 
Sori near costules; indusium at first covering sorus, 
very thin and mostly caducous, leaving a thin 
basal disc which either has the receptacle at its 
centre or may be only on costular side of recep- 
tacle; paraphyses short. Scales and hairs: pinna- 
rachis smooth and glabrous, pale; lower surface 
of costae bearing some residual flat elongate 
entire brown scales, grading to small flat scales; 
pale bullate scales on costules. 

Type specimen: Clemens 34012, Mt Kinabalu, 
N. Borneo (BO; dupl. at US). 

Distr. Malaysia: N. Borneo. 

Ecol. In forest, at 1250-2000 m. 

41. Cyathea rubiginosa (Brause) Domin, Acta 
Bot. Bohem. 9 (1930) 154; Copel. Philip. J. Sc. 77 
(1947) 109. — Alsophila rubiginosa Brause, Bot. 
Jahrb. 56 (1920) 66. — Alsophila hunsteiniana 
Brause, ibid. 65. — C. albidula Domin, Acta Bot. 
Bohem. 9 (1930) 88. 

Stipe 60 cm, dark purplish with many small 
conical spines to 1 mm high; scales to 15 mm 
(5 cm?) by hardly 1 mm wide above the base, 
thick at the base, edges pale and thin, bearing 
many long flexuous setae, also apparently super- 
ficial setae, near base of scale; very small dull 
brown scales bearing short dark setae also pres- 
ent. Lowest pinna 25 cm long, longest 42 cm. 
Pinnules well-spaced, to 67 by 17 mm, lobed to 
c. 1 mm from costa, subsessile, acuminate; cos- 
tules 4-^'/2 mm apart; veins to 8 pairs; lamina- 
segments rigid, edges crenate. Sori on distal veins 
near costule. on basal veins further from it; in- 
dusium very thin, at first covering sorus, break- 
ing and mostly caducous, leaving an irregular 
disc round base of sorus. Scales and hairs: 
pinna-rachis smooth and more or less glabres- 
cent, usually with scattered narrow dull copiously 
setiferous scales 3-6 mm long and very small 
irregular non-setiferous dull brown scales; basal 
scales on costae elongate, dull, with irregular 
long setae, grading to pale bullate scales distally 
and on costules, some bullate scales bearing setae. 

Type specimen: Ledermann 12539, Sepik Re- 
gion, NE. New Guinea (B). 

Distr. Malaysia: New Guinea. 

Ecol. In rocky open forest or mossy forest, at 
1 100-2840 m. 

42. Cyathea apiculata (Rosenst.) Domin, Pterid. 
(1929) 262. — Alsophila apiculata Rosenst. in 
Fedde, Rep. 13 (1914) 213; v. A. v. R. Handb. 
Suppl. (1917) 73. — Alsophila indrapurae v. A. v. R. 
Bull. Jard. Bot. Btzg II, n. 20 (1915) 2; Handb. 
Suppl. (1917) 63. — C. crenulata f. subspinulosa 
V. A. V. R. Handb. Suppl. (1917) 28.— C. indra- 
purae V. A. V. R. Bull. Jard. Bot. Btzg II, n. 28 
(1918) 13.— C. paleata Copel. Un. Cal. Publ. Bot. 
14(1929)372,1.56. 



94 



Flora Malesiana 



[ser. II, vol. 12 



Stipe more than 30 cm, dark at base, paler up- 
wards, not spiny, scaly almost throughout; scales 
to 15 by 3 mm, dark with paler fragile edges. 
Pinnae to at least 40 cm long. Pinnules to 75 by 
20 mm, almost sessile, caudate-acuminate, lobed 
almost to costa; costules 4-41/2 mm apart; veins 
10 pairs; lamina-segments slightly crenate, sinuses 
narrow. 5or/ near costules; indusium very thin and 
fragile, breaking and mostly caducous; receptacle 
globular; paraphyses short, slender. Scales and 
hairs: pinna-rachis smooth and glabrescent be- 
neath, not hairy; costal scales few, those near base 
flat, ovate, bearing a few setae on pale edges; 
costular scales broad, thin, pale, not buUate. 

Type specimen: J. Winkler, Pea Radja, Ba- 
takerland, Sumatra (Rosenst., Fil. Sumatr. Exsic. 
197, S-PA; dupl. at BM, L, P, UC). 

Distr. Malaysia: Sumatra. 

Ecol. In forest, at 1800 m. 

Note. Under the original description of C. in- 
drapiirae, three separate collections, made by 
C. G. Matthew in different localities, were cited. 
In view of the name indrapurae. the specimen n. 
374, from Indrapoera (= Mt Kerintji) is selected 
as type, and appears to me to be not different from 
C apiculata; n. 696, from Mt Merapi, is C. 
hyme nodes Mett. 

43. Cyathea biinnemeijerii v. A. v. R. Bull. Jard. 
Bot. Btzg III, 5 (1922) 187. 

Stipe dark, at least 30 cm. near base bearing 
many conical spines hardly 1 mm long; scales 
dark with paler fragile edges, mostly caducous. 
Pinnae to 40 cm or more long. Pinnules to 70 by 
16 mm, lobed almost to costa, lowest segment 
almost free; costules 31/2-^ mm apart; veins to 10 
pairs; lamina-segments rather thin, drying very 
dark, edges crenate. Sori near costules; indusium 
thin, at first completely covering sorus, breaking 
and in part falling, the persistent part some- 
times cup-shaped; paraphyses longer than spo- 
rangia, in some cases 2 cells wide at base. Scales 
and hairs: pinna-rachis purplish beneath, slightly 
warty, glabrescent; scales at base of costa brown, 
flat, elongate, entire, grading to paler ovate or 
bullate scales distally and on costules. 

Type specimen: BIjnnemeijer 5839, Mt Ranai 
Bunguran, Natuna Is (BO; dupl. at L). 

Distr. Malaysia: Natuna Islands (NW. of 
Sarawak). 

Ecol. In open scrub on summits of two hills, 
at 600 m. 

44. Cyathea excavata Holttum, Gard. Bull. S. S. 
8 (1935) 306; Rev. Fl. Malaya 2 (1954) 121; 
Molesworth Allen, Gard. Bull. Sing. 17 (1959) 
255, photogr. facing p. 266. — Fig. 12. 

Trunk to 2 m, sometimes with lateral buds 
forming small crowns of fronds; fronds to 2 m long. 
Stipe at least 40 cm, smooth and green (pale when 
dry); basal scales few, dull, thin, soon disappearing; 
pneumathodes 15-20 mm long, in a single almost 
continuous row, at base of stipe shorter and deeply 
excavated. Pinnae to 60 cm long, lowest somewhat 
reduced. Largest pinnules commonly 80-100 by 




Fig. 12. Cyathea excavata Holttum. Trunk-apex, 

showing deeply excavated pneumathodes arranged 

as V below base of a stipe. Cameron Highlands, 

Malaya (R. E. Holttum). 

18-22 mm, sometimes to 130 by 27 mm, sessile, 
rather shortly acuminate, lobed almost to costa, 
lowest segment not free; costules 4-5 mm apart; 
veins 10-12(-14) pairs, often twice forked, pin- 
nately branched where segments are deeply lobed; 
lamina-segments thin, the larger ones usually 
lobed I3-V2 towards costule. Sori near costules, 
usually only on 1-3 basal pairs of veins; indusium 
pale, thin, covering young sorus completely but 
soon breaking, leaving an irregular persistent disc 
as large as base of sorus; receptacle swollen. 
often split when dry; paraphyses short. Hairs and 



Dec. 1963] 



Cyatheaceae (Holttum) 



95 



scales: pinna-rachis smooth and glabrous; costal 
scales sparse, brown, thin, entire, broadly and 
irregularly ovate, sometimes with a few dark setae; 
thinner similar scales on costules. distal ones pale, 
entire, fiat; on veins many conspicuous very small 
hairs (bases of former scales). 

Type specimen: Holttum 23538, Cameron 
Highlands, Malaya (S; dupl. at BO, K). 

Distr. Malaysia: Malay Peninsula. 

Ecol. Only known on Main Range in Ca- 
meron Highlands district at c. 1500 m; originally 
found in primary forest, in recent years on stream- 
sides in clearings and on forest edges, also in open 
grassy places ( Molesworth Allen, I.e.). Old 
fronds persist, hanging down and covering the 
trunk, as in C. orientalis (also in C. crenuUitci). 

45. Cyathea christii Copel. Philip. J. Sc. 1. Suppl. 
II (1906) 144; ibid. 4 (1909) Bot. 49; v. A. v. R. 
Handb. (1908) 785; Suppl. (1917) 29; Copel. 
Fern Fl. Philip. 2 {I960) 217. 

Stipe to 15 cm. hov-er pinnae gradually reduced 
(sometimes a gap between lowest and next ?), 
lowest 5 cm or more long, longest 50 cm. Pinnules 
to 70 by 15 mm, lobed nearly to costa; costules 
3-3 '/a rnm apart; veins to 8 pairs; lamina-segments 
rather thin, more or less crenate. Sori near cos- 
tules; indusium at first completely covering sorus, 
rather thin, breaking irregularly and sometimes in 
part caducous. Scales and hairs: pinna-rachis pale 
and glabrescent on lower surface, warty towards 
base, sometimes with residual narrow scales 
bearing a few setae; buUate scales present through- 
out lower surface of costae and on costules; 
sometimes narrow setiferous scales present also 
near base of costae. 

Type specimen: Copeland 1141, Mt Apo, 
Mindanao (MICH; dupl. at US). 

Distr. Malaysia: Philippines (Mindanao). 

Ecol. At 900-1800 m. 

46. Cyathea geluensis Rosenst. in Fedde, Rep. 5 
(1908) 371; ibid. 12 (1913) 525, incl. var. tomentosa 
RosENST.; V. A. V. R. Handb. Suppl. (1917) 30; 
Copel. Philip. J. Sc. 77 (1947) 102.— C. no\o- 
guineensis Brause, Bot. Jahrb. 49 (1912) 12, 
fig. IB; V. A. V. R. Handb. Suppl. (1917) 35.— 
C. sepikensis Brause, Bot. Jahrb. 56 (1920) 54; 
Copel. Philip. J. Sc. 77 (1947) 103.— C. subspathu- 
lata Brause, Bot. Jahrb. 56 (1920) 53. 

Trunk slender; fronds to c. 10, 100-230 cm long. 
Stipe variable in length, warty or shortly spiny, 
scaly near base; scales pale, or partly or wholly 
dark, with dull fragile edges, commonly to 15 by 
2 mm, in some specimens to 20 by 5 mm; stipe 
also covered with a more or less continuous layer 
of pale flexuous hairs or very small scales bearing 
such hairs. Lower pinnae reduced, sometimes con- 
tinuously to a small size (5-8 cm long), sometimes 
a single small pair near base and then a gap; 
longest pinnae 25^5 cm long. Largest pinnules 
commonly 40-60 by 12-15 mm, on some plants 
to 80 by 20 mm, almost sessile, acuminate, lobed 
almost to costa, lowest segment sometimes almost 
free; costules 31/2-4 mm apart; veins 7-9 pairs; 



lamina-segments rather thin, crenate to subentire. 
Sori near costules; indusium pale and thin but 
firm, at first covering sorus, breaking irregularly 
and persistent; paraphyses slender, as long as 
sporangia. Scales and hairs: lower surface of 
main rachis and pinna-rachis closely covered 
with entangled flexuous pale hairs, with more or 
less abundant pale scales mostly bullate at base; 
costae rather closely covered with pale bullate 
scales, sometimes near base also flexuous hairs 
and elongate pale flat scales; pale bullate scales on 
costules. 

Type specimen: Werner 80, Mt Gelu, NE. New 
Guinea (S-PA; dupl. at B). 

Distr. Malaysia: Central and Eastern New 
Guinea, Louisiade Archipelago. 

Ecol. In forest at 1000-2000 m on mainland; 
at 700-900 m on the islands, mostly in mossy forest. 

Note. Possibly more than one species should be 
recognized; specimens examined seem to show 
various combinations of characters, especially as 
regards length of stipe and degree of reduction of 
lower pinnae. 

47. Cyathea macgregorii F. v. M. Trans. R. Soc. 
Victoria 1 (1889) 40; Bak. J. Bot. 28 (1890) 104; 
V. A. V. R. Handb. (1908) 17; Ridl. Trans. Linn. 
Soc. II, Bot. 9 (1916) 251; C. Chr. Brittonia 2 
(1937) 280. — C keysseri Rosenst. in Fedde, Rep. 
12 (1913) 164; v. A. v. R. Handb. Suppl. (1917) 
23. — C. cheilanthoides Copel. Un. Cal. Publ. Bot. 
18 (1942) 219; Philip. J. Sc. 77 (1947) 121, pi. 14.— 
Fig. lOe, f, g, 13. 

Trunk to 3 m, to 24 cm 0, bearing up to 60 
fronds 70-100 cm long. Stipe 5-15 cm, its base 
covered with shining brown dull-edged scales to 
50 by 2-3 mm, warty and glaucous beneath after 
fall of scales; main rachis more or less persistently 
covered with small pale long-fringed scales, also 
narrow elongate thin pale scales, sometimes quite 
glabrescent. Lower pinnae gradually reduced, 
sometimes irregularly spaced, with a rather long 
gap between lowest pinnae (sometimes hidden by 
basal scales) and the next; lowest pinna 5-7 cm 
long; longest pinna 11-18 cm long. Largest 
pinnules 20-35 by 7-8 mm, bearing free but almost 
sessile very rigid tertiary leaflets, their bases 2-2V2 
mm apart. Largest tertiary leaflets commonly 3 by 
212 mm, ovate to deltoid, edges strongly reflexed 
and inroUed, more or less lobed near base; veins 
to 4 pairs, basal ones forked, strongly raised on 
lower surface but not on upper. Sori 4-6 on largest 
tertiary leaflets, fewer on smaller ones; indusium 
firm, brown, at first covering sorus, breaking ir- 
regularly and persistent; receptacle swollen, no 
persistent paraphyses. Scales and hairs: pinna- 
rachis and costae more or less persistently scaly on 
lower surface as main rachis, at length glab- 
rescent and finely warty; costules bearing small 
long-fringed pale or brown scales between the sori. 

Type specimen: W. McGregor 63, Mt Knuts- 
ford, E. New Guinea (MEL; dupl. at K, P). 

Distr. Malaysia: New Guinea. 

Ecol. In open peaty grassland or on edge of 
forest, sometimes gregarious, 3000-3700 m. 



96 



Flora Malesiana 



[ser. II, vol. P 




Fig. 13. Cyathea macgregorii F. v. M., Mt Wilhelm, Eastern Highlands, Territory of New Guinea, 

3450 m (R. D. Hoogland). 



48. Cyathea gleichenioides C. Chr. Brittonia 2 
(1937)281;CoPEL. Philip. J. Sc. 77 (1947) 123.— 
Fig. 1-2, 14. 

Differs from C macgregorii as follows: pinnae 
to 101/2 cm long; pinnules mostly 13-15 mm long, 
basal acroscopic pinnule to 17 mm; tertiary leaf- 
lets almost circular, c. 1 mm long and wide, bear- 
ing 1 or 2 sori. 

Type specimen: Brass 4595, Murray Pass, 
Wharton Range (BM; dupl. at A, BO, BRI). 

Distr. Malaysia: Eastern New Guinea. 

Ecol. 'A conspicuous feature of the open grass- 
lands' at 2840-3680 m (Brass). 

Note. Christensen stated that another dis- 
tinctive character of this species in the occurrence 
of short pinnae close to the base of the stipe, with 



a gap of 15 cm to the next pair; but such a con- 
dition has also been seen in fronds which have 
leaflets like typical C. macgregorii. It seems that 
there is a good deal of variation in the disposition 
of the lower pinnae in the latter species. It seems 
to me also possible that the distinction of size of 
tertiary leaflets is not a constant one, in which 
case C. gleichenioides should be united with C. 
macgregorii. 

49. Cyathea havilandii Bak. Trans. Linn. See. II, 
Bot. 4 (1894) 249; v. A. v. R. Handb. (1908) 22; 
C. Chr. Gard. Bull. S. S. 7 (1934) 221.— C. 
paleacea Corel. Philip. J. Sc. 12 (1917) Bot. 53.— 
C. rigida Corel. I.e. 

Trunk very short; fronds almost erect, to c. 100 



Dec. 1963] 



Cyatheaceae (Holttum) 



97 




Fig. 14. Cyathea gleichenioides C. Chr. (Brass 4265), top of trunk and bases of fronds, showing lowest 
pinnae partly covered by mass of scales. Mt Albert Edward, Papua, 3680 m (L. J. Brass). 



cm long. Stipe 30^0 cm, dark and warty where 
scales are removed, densely scaly throughout; 
scales near base to 15 by 1-2 mm, shining medium 
brown with very narrow concolorous fragile edges, 
size decreasing gradually upwards; upper part of 
stipe on abaxial surface densely covered also with 
much smaller scales of same colour, each ending 
in a long flexuous brown seta. Lower pinnae not 
much reduced; longest pinna 10-16 cm long. 
Pinnules to 25 by 7 mm, only a few near the base 
of larger pinnae free, rest separately adnate to 
pinna-rachis or connected by a narrow wing; 
largest pinnules, where fertile, lobed about half- 
way to costa, smaller and sterile ones slightly 
lobed; costules 2 mm apart; veins 2-3 pairs, 
strongly raised on lower surface; lamina very rigid, 
drying dark. Sori in a single row on each side of 
costa of pinnule, 1 or 2 to each vein-group; indu- 
sium very firm and dark, covering sorus to ma- 
turity, then gaping a little at the apex and breaking 
irregularly; paraphyses slender, short. Scales and 
hairs: main rachis densely and persistently scaly 
beneath throughout as upper part of stipe, some 
scales on distal part having bullate bases; pinna- 
rachis and costae of pinnules similarly scaly, scales 
smaller, costal ones mostly quite bullate, all 
ending in a long flexuous shining brown hair 
(hair-tip of costal scales often more than 1 mm 
long); upper surface of main rachis and pinna- 
rachis densely hairy, hairs long, darker than scales, 
antrorse; upper surface of costae glabrescent. 



Type specimen: Haviland 1485, Mt Kinabalu, 
N. Borneo (K). 

Distr. Malaysia: North Borneo. 

Ecol. Abundant in the low open Leptospermiim- 
Dacrydiiim forest of the ridges on Mt Kinabalu, 
2400-3000 m. 

50. Cyathea imbricata v. A. v. R. Nova Guinea 
14 (1924) 11; C. Chr. Brittonia 2 (1937) 282; 
Corel. Philip. J. Sc. 77 (1947) 123. 

Trunk to 2 m; fronds 60-70 cm long. Stipe 
71/2-10 cm, dark, bearing spines 1 mm long, scaly 
when young; largest scales 12-15 by 2-A mm, 
castaneous with paler fragile edges; small scales 
forming a woolly covering also present. Lower 
pinnae gradually reduced, lowest less than 5 cm 
long; middle pinnae close and imbricate, to 8 V2 cm. 
Pinnules to 18 by 8 mm, deeply lobed except 
towards apex, lobes close, 2-3 by 2 mm, thick and 
rigid but edges not reflexed; veins 4-5 pairs, simple 
or forked. Sori 1-4 to each lobe of lamina; indu- 
sium firm, at first quite covering sorus, breaking 
irregularly and persistent. Scales and hairs: pinna- 
rachis glabrescent beneath, residual scales firm, 
brown, with thin paler edges bearing a few short 
hairs and a hair-point; scales on costae similar 
but smaller, with long caudate tips, mostly 
caducous. 

Type specimen: Lam 1625, foot of Doorman 
summit, W. New Guinea (BO; dupl. at S, US, 
L, U, UC). 



98 



Flora Malesiana 



[ser. II, vol. 12 



Distr. Malaysia: Western New Guinea. 
Ecol. In open forest at 3240 m. 

51. CyathealongipesCoPEL. Philip. J. Sc. 12(1917) 
Bot. 54; C. Chr. Card. Bull. S. S. 7 (1934) 222. 

Stipe slender, to 200 cm, dark and copiously 
spiny near base; spines to 5 mm long; basal scales 
mostly caducous, rather broad; no persistent small 
scales. Pinnae to at least 70 cm long. Pinnules all 
stalked except distal ones, stalks of lowest on 
lower pinnae to 10 mm, on smaller pinnae 3-6 mm; 
largest pinnules 100-1 30 by 20-32 mm, acuminate, 
1-3 pairs of basal segments free or separately 
adnate and connected by a costal wing, rest of 
pinnule lobed almost to costa; costules 41/2-6 mm 
apart; veins 10 pairs; lamina-segments crenate or 
the lowest ones more deeply lobed. Sori near 
costules; indusium rather thin, at first completely 
covering sorus, breaking irregularly and mostly 
persistent; receptacle swollen; paraphyses slender, 
about as long as sporangia. Scales and hairs: 
main rachis and pinna-rachis quite glabrescent, 
rather pale, sparsely short-spiny; scales on lower 
surface of costae ovate-acuminate, thin, entire, 
brown, shorter and more or less buUate distally; 
bullate-acuminate scales on costules. 

Type specimen: Clemens 10915, Mt Kinabalu, 
North Borneo (MICH; dupl. at K, BO, UC, A). 

Distr. Malaysia: North Borneo. 

Ecol. In ridge forest at 1250-1500 m, only on 
Marai-Parai spur of Mt Kinabalu, locally abundant. 

52. Cyathea acuminata Copel. Philip. J. Sc. 81 
(1952) 15; Fern Fl. Philip. 2 (1960) 213. 

Stipe not known. Pinnae to 70 cm long. Largest 
pinnules 100-120 by 22 mm, long-acuminate, lower 
ones on stalks to 4 mm long; all segments separated 
by rather wide sinuses, several lower pairs sep- 
arately adnate to costa; costules 41/2-6 mm apart; 
veins to 12 pairs; lamina-segments rather thin, 
strongly crenate-serrate. Sori near costules; indu- 
sium at first quite covering sorus, rather firm, 
breaking irregularly and persisting; receptacle 
much swollen, bearing some small scales at its 
apex; paraphyses slender. Scales and hairs: pinna- 
rachis glabrescent beneath, bearing scattered small 
spines near base; scales near base of costae elon- 
gate, flat, dull brown, entire, grading to bullate 
ones distally and on costules. 

Type specimen: Ramos & Edano, BS 30900, 
Jamindan, Capiz Prov., Panay (UC; dupl. at S, 
US, K, P, BM). 

Distr. Malaysia: Philippines (Panay, Samar). 

Note. The type of this species and other spec- 
imens were distributed from Manila as C. spi- 
nulosa. 

53. Cyathea insulana Holttum, Kew Bull. 16 
(1962) 56. 

Trunk 8-10 m, 14 cm 0; fronds spreading, 3 m 
long including stipe. Stipe 100 cm, covered with 
many thick conical spines 2 mm long and through- 
out with a thin felt of very small pale-brown short- 
fringed or setiferous scales; large scales abundant 
along edges of grooves of adaxial surface of basal 



25 cm of stipe, these scales 35-45 mm long, shining 
castaneous, rigid, much twisted, with pale fragile 
edges, largest scales 1 '/2-3 mm wide at base; 
pneumathodes in a single row on each side of 
stipe, 7-1 1 mm long, those near base deeply 
excavated. Pinnae to at least 50 cm long. Largest 
pinnules 120-130 mm long, caudate-acuminate, 
lobed nearly to costa with basal 1-2 segments 
separately adnate; sterile pinnules 30 mm wide, 
fertile 20 mm, lowest ones with stalks 2 mm long; 
costules to 5 mm apart; veins 12-14 pairs; lamina- 
segments strongly crenate or lowest ones more 
deeply lobed. Sori near costules; indusium at 
first completely covering sorus, thin and pale, 
breaking irregularly. Scales and hairs: scales on 
pinna-rachis abundant, very small, light brown, 
mostly setiferous; on costae similar scales and 
sometimes also narrow setiferous ones; on cos- 
tules similar very small setiferous scales, also 
a few larger thin ovate flat or convex scales bearing 
a few slender setae; on veins many very small 
light brown almost circular somewhat bullate 
scales, not setiferous. 

Type specimen: Brass 24725, Goodenough L, 
Milne Bay District, Papua (A; dupl. at L). 

Distr. Malaysia: d'Entrecasteaux Is. (Goode- 
nough I., Normanby I., Fergusson I.). 

Ecol. In mossy forest or in ravines near rain- 
forest-mossy forest transition, 750-1600 m. 

Note. The specimens here included from Nor- 
manby and Fergusson Is are smaller than the type 
and from lower elevations; they have stipes less 
spiny, pinnules to 82 by 20 mm. They also show 
elongate narrow setiferous scales on costae; pos- 
sibly these were caducous on the type specimen. 

54. Cyathea pseudomuelleri Holttum, Kew Bull. 
16 (1962) 61.— Fig. 15. 

Trunk 4 m, bearing many fronds; stipe-bases 
persistent. Stipe 10 cm, above base dull, rather 
pale, finely warty; scales near base many, to 40 by 
hardly 1 mm, rigidly erect, twisted, dark with very 
narrow paler edges. Lower pinnae gradually re- 
duced, lowest 12 cm long, longest 24 cm. Largest 
pinnules 45 by 10 mm, very rigid, shape and tex- 
ture about as in C. muelleri, diff'ering in having 
veins prominent on both surfaces. Sori at first 
completely covered by firm indusia which break 
irregularly and persist. Scales and hairs: most 
scales on pinna-rachis and costae soon caducous, 
remaining larger ones light brown, flat, elongate, 
setiferous, also pale sub-buUate scales 1/2 mm 
long and wide; costules usually bare; veins 
bearing minute hairs on lower surface. 

Type specimen: Brass 9430, Mt Wilhelmina, 
W. New Guinea (A; dupl. at L, BO, MICH, 
UC). 

Distr. Malaysia: W. New Guinea (one collec- 
tion). 

Ecol. At 3200 m. 

55. Cyathea archboldii C. Chr. Brittonia 2 (1937) 
278; Copel. Philip. J. Sc. 77 (1947) 105.— C. bi- 
dentata Copel. Un. Cal. Publ. Bot. 18 (1942) 218; 
Philip. J. Sc. 77 (1947) 105, pi. 3. 



Dec. 1963] 



Cyatheaceae (Holttum) 



99 




Fig. 15. Cyathea pseiidomuelleri Holttum 
(Brass 9430), showing habit and persistent stipe- 
bases throughout trunk. Mt Wilhelmina, W. New 
Guinea, 3200 m (L. J. Brass). 

Fronds 200-300 cm long, about 10. Stipe c. 50 cm, 
base bearing short spines and covered with scales; 
largest scales 30^0 by 1 Vi-l mm, very pale to 



pale brownish with fragile edges which may be 
darker than the rest; also present a rather close 
thin layer of rusty brown scurf consisting of very 
small irregular scales mostly bearing 1 or more 
dark flexuous setae; rachis similarly covered with 
small scales, sometimes also with narrow scales 
to 10 mm long. Largest pinna 45 cm long. Largest 
pinnules 60-90(-l20) byl3-20(-25) mm, sessile, 
several basal pairs of segments separately adnate 
to costa; costules 3-5 mm apart; veins to 9 pairs; 
lamina-segments rather thick and rigid, crenate- 
serrate, basal ones lobcd up to '/2 way to costule, 
lobes retuse or bidentate. Sori near costules; 
indusium at first covering sorus, sometimes open- 
ing by a small apical circle, soon breaking ir- 
regularly, persistent. Scales and hairs: pinna- 
rachis, costa and costules covered with scales like 
the small ones on the stipe, those on costules with 
many long flexuous shining marginal setae, none 
buUate; small dark setiferous scales also present 
on lower surface of veins; narrow elongate scales 
sometimes also present on pinna-rachis and costae. 

Type specimen: Brass 4551, Murray Pass, 
Wharton Range, Papua (BM; dupl. at A, BO, 
BRl, MICH). 

Distr. Solomon Is (Bougainville), in Malaysia: 
New Guinea. 

Ecol. At 1950-2840 m,in forest, in New Guinea; 
at 1000 m on Bougainville I. The Bougainville 
specimen is less rigid and small scales generally 
are pale-fringed, not setiferous. A Papuan speci- 
men from 3000 m, in alpine shrubbery (Schodde 
1890) is smaller and much more densely scaly 
throughout than other specimens. 

var. horrida Holttum, var. nov. 

A typo speciei differ t: paleis s tip it is atrocastaneis, 
spinis ad 1^2 ''W longis, paleis venarum paiicioribus, 
nan vel raro setiferis. 

Type specimen: Hoogland & Pullen 5506, 
NE. New Guinea, at 2400 m, in mountain forest 
(K). 

56. Cyathea foersteri Rosenst. in Fedde, Rep. 10 
(1912) 321; v. A. v. R. Handb. Suppl. (1917) 28; 
C. Chr. Brittonia 2 (1937) 279; Copel. Philip. J. 
Sc. 77 (1947) 104. 

Trunk to 10 m, bearing 9 fronds 225 cm long 
(Brass). Stipe 5-15 cm, closely covered throughout 
with pale scales to 15 by little over 1 mm. Lower 
pinnae gradually reduced, lowest 8 cm long; 
longest 35 cm. Largest pinnules to 90 by 15 mm 
(on a small frond to 55 by 16 mm), lowest seg- 
ments (sometimes several pairs) free or distinctly 
separate and joined by a narrow wing; costules 
21/2-3 1/2 mm apart; veins 7-10 pairs, dark and raised 
on lower surface; lamina-segments rather thin, 
finely crenate-serrate, free basal ones sometimes 
deeply lobed, lobes bidentate. Sori near costules; 
indusium thin and firm, covering sorus to ma- 
turity, breaking and persisting; receptacle swollen; 
apparently no paraphyses. Scales and hairs: pinna- 
rachis more or less persistently covered with 
small pale scales which are usually setiferous, 
also sometimes scattered narrow scales 5 mm or 



100 



Flora Malesiana 



[ser. II, vol. P 



more long, often with long dark marginal setae; 
costal scales ovate to elongate, often closely over- 
lapping, edges usually setiferous but the smaller 
sometimes with a short pale fringe; costular scales 
as costal but smaller, in some cases smallest ones 
setiferous, in other cases fringed with short hairs; 
similar scales more or less abundant on lower 
surface of veins. 

Type specimen: Keysser 16, Sattelberg, NE. 
New Guinea (S-PA; dupl. at B). 

Distr. Malaysia: Eastern New Guinea. 

Ecol. In scrub on forest edge and in mossy 
forest, at 1600-2800 m. 

Note. This species is very near C. archboldii, 
but appears to be distinct in its short stipe with 
reduced lower pinnae, and shorter narrower 
stipe-scales; also possibly in thinner pinnule- 
segments. There is a good deal of variation in the 
abundance of dark setae on the scales of pinna- 
rachis, costae and costules. 

57. Cyathea nigrolineata Holttum, Kew Bull. 16 
(1962) 58. 

Fronds 210 cm or more long, in 1 or 2 whorls of 
5-8 each. Stipe to 12 cm, not spiny, densely cov- 
ered throughout with scales to 25 by less than 1 
mm, shining and almost white or more usually 
with a narrow median black line, all ending in a 
dark seta, fragile edges dull; also a thin under- 
cover of small dull brown scales, some setiferous. 
Lower pinnae gradually reduced, lowest 5 cm long, 
longest 35-45 cm (rarely to 60 cm). Largest 
pinnules of type 47 by 12 mm, of another collec- 
tion 90 by 20 mm, sessile, apex abruptly pointed, 
several lower pairs of segments partly free (con- 
stricted on acroscopic side at base), 1-2 pairs 
sometimes quite free; costules 21/2-3 mm apart; 
veins 8-10 pairs, concolorous and not raised on 
lower surface; lamina-segments very firm, edges 
entire or undulate, not crenate, or those on larger 
pinnules sometimes lobed where fertile, lobes 
sometimes bidentate. Sori near costules; indusium 
firm, brown, at first complete, breaking and 
persistent. Scales and hairs: pinna-rachis bearing 
copious very small brown scales with setiferous 
apex, also scattered scales 2-3 by '/2 mm, with 
dark shining mid-band and paler edges, and some- 
what smaller, narrower scales bearing marginal se- 
tae; scales on costae many, small, dark, shining,with 
many curved dark setae; on costules and veins 
smaller, those on veins scattered but abundant. 

Type specimen: Hoogland & Pullen 5495, 
Eastern Highlands, NE. New Guinea (K; dupl. at 
BO, US, SYD, BRI, L). 

Distr. Malaysia: Eastern New Guinea (4 
collections). 

Ecol. At 2300-2400 m in forest or secondary 
growth (specimen from secondary growth is 
largest). This species should perhaps be united 
with C foersteri Rosenst. Type material of the 
latter does not show stipe nor larger scales of 
pinna-rachis. 

58. Cyathea inquinans Christ, Verb. Nat. Ges. 
Basel 11 (1896) 422; Ann. Jard. Bot. Btzg 15 



(1898) 83, t. 13, f. 5; v. A. v. R. Handb. (1908) 23. 

Stipe 15 cm, densely scaly; larger scales bright 
red-brown, rather thin, to 30 by 2 mm, long- 
acuminate, apex a dark red seta, fragile edges nar- 
row; small scales very abundant, often with dark 
red setiform apex. Rachis similarly scaly on lower 
surface, larger scales to 10 by 1 mm, bearing 
rather many dark red setae more than Yi mm long. 
Pinnules to 60 by 15 mm, short-acuminate, lobed 
nearly to costa, close and more or less imbricate; 
costules 3 mm apart; veins 9 pairs; lamina-seg- 
ments where fertile rather deeply lobed. Sori 
near costules; indusium thin, brown, at first com- 
plete, breaking and persistent. Scales and hairs: 
pinna-rachis scaly as main rachis, small scales 
very abundant and mostly setiferous; costal scales 
light brown, all setiferous, grading from elongate 
acuminate to very small; costular scales as smaller 
ones on costae; no hairs on lower surface of 
pinnules. 

Type specimen: P. & F. Sarasin 1328, Mt 
Lompobattang (= Bonthain), SW. Celebes (BAS). 

Distr. Malaysia: SW. Celebes (2 collections), 
Moluccas: Ceram (?). 

Ecol. At 2000-2800 m. 

Note. A sterile specimen collected by Eyma in 
Ceram (//. 2371) agrees with C. inquinans in scali- 
ness, but is larger, apparently with a longer stipe; 
it has pinnae to 60 cm long, pinnules to 100 by 
20 mm, costules 4 mm apart, veins 15 pairs. If not 
C. inquinans, it probably represents a new species. 
The altitude given for this specimen is 40 m, but 
Eyma had just descended from the mountains and 
it seems possible that the specimen was collected 
there. 

59. Cyathea ferruginea Christ, Philip. J. Sc. 2 
(1907) Bot. 181; V. A. v. R. Handb. (1908) 784; 
COPEL. Fern Fl. Philip. 2 (1960) 215.— C. fer- 
rugineoides Copel. Philip. J. Sc. 81 (1952) 15; 
Fern Fl. Philip. 2 (1960) 215. 

Stipe 12 cm, bearing many spines 1 mm long; 
scales rather sparse, to 15 by 1 mm, dark with 
narrow pale fragile edges bearing setae. Lowest 
pinnae 5-6 cm long, possibly a gap between these 
and next pinnae; longest pinna 35 cm or more. 
Pinnules to 80 by 15-20 mm, lower ones on stalks 
2-4 mm long, acuminate, lobed nearly to costa; 
costules 3V2-4I/2 mm apart; veins to 9 pairs; 
lamina-segments thin to firm, slightly crenate; 
fertile pinnules narrower than sterile, with closer 
costules. Sori near costules; indusium at first com- 
plete, thin, pale, breaking irregularly and per- 
sistent; paraphyses many, as long as or longer 
than sporangia, not widened at base. Scales and 
hairs: pinna-rachis glabrescent on lower surface, 
residual scales mostly very narrow, on distal part 
some scales as costae and also slender crisped hairs; 
scales near base of costae narrow, flat, entire, 
grading to paler acuminate bullate scales; bullate 
scales on costules. 

Type specimen: Foxworthy BS 560, Palawan 
(P; dupl. at BO, K, US, MICH, A). 

Distr. Malaysia: Philippines (Palawan, Bala- 
bac, Negros). 



Dec. 1963] 



Cyatheaceae (Holttum) 



101 



Ecol. In forest, to 1 150 m, the type from mossy 
forest, near summit of Mt Pulgar, other specimens 
apparently from lower altitudes. 

60. Cyathea oosora Holttum, Kew Bull. 16 (1962) 
59. — C. assimilis [nan Hook.] Christ, Ann. Jard. 
Bot. Btzg 15 (1898) 82. 

Stipe 50 cm or more, warty near base; pneuma- 
thodes 14-20 mm long, in an almost continuous 
row; scales not seen. Longest pinnae 60 cm long. 
Largest pinnules 90 by 20 mm, sessile or nearly so, 
acuminate, lobed almost to costa. lowest segment 
not free;costules 3'/2-4 mm apart; veins 9-10 pairs; 
lamina-segments rigid, edges crenate. Sori near 
costules; indusium firm, shining brown when old, 
at first ovoid with a small apical aperture, breaking 
irregularly and persistent; paraphyses short, 
slender. Scales and hairs: pinna-rachis glabrescent 
on lower surface, residual scales 3-4 mm long, 
narrow, brown, crisped, not setiferous; costae 
densely scaly, scales uniformly brown, lower ones 
elongate-acuminate with slightly buUate base, 
grading to hair-pointed bullate scales distally and 
on costules; some of the larger costal and costular 
scales have marginal concolorous hairs, not dark 
setae. 

Type specimen: Clemens 51188, Mt Kinabalu, 
N. Borneo (K; dupl. at A, L, MICH, UC). 

Distr. Malaysia: N. Borneo, N. Celebes. 

Ecol. At 2200-3000 m in ridge-forest, on Mt 
Kinabalu; at 1500-2000 m in Celebes. 

Note. The Celebes specimens differ from those 
on Mt Kinabalu in having paler scales, and one of 
them (Sarasin 933) has very few scales on the 
pinnules. 

61. Cyathea halconensis Christ, Philip. J. Sc. 3 

(1908) Bot. 270; Copel. Philip. J. Sc. 4 (1909) 
Bot. 51; V. A. V. R. Handb. Suppl. (1917) 24; 
Copel. Fern Fl. Philip. 2 (1960) 216.— C. meanisii 
Copel. Philip. J. Sc. 3 (1909) Bot. 356; ibid. 4 

(1909) Bot. 57; v. A. v. R., Handb. Suppl. (1917) 
24; Copel. Fern Fl. Philip. 2 (1960) 214.— 
C. melaiwphlebia Copel. Philip. J. Sc. 38 (1929) 
131; Fern Fl. Philip. 2 (1960) 220. 

Stipe commonly 5 cm, spines copious, conical, 
to 1 mm long; scales to 30 by 2 mm, dark brown, 
hardly shining, with distinct concolorous fragile 
edges bearing a few dark setae. Lower pinnae 
gradually reduced, lowest 7-12 cm long, sometimes 
bipinnate; longest pinna 50 cm. Largest pinnules 
commonly 70-100 by 20 mm wide at base, sessile, 
caudate-acuminate (cauda to 15 mm), lowest 1-3 
pairs of segments free or nearly so, rest of pinnule 
lobed almost to costa; costules 3'/2^V2 rnm 
apart; veins to 12 pairs; lamina-segments thin but 
firm, crenate. Sori near costules, quite covered 
when young by rather thin pale indusia which 
break and persist; receptacle swollen; paraphyses 
short, some apical ones with a broad base. Scales 
and hairs: pinna-rachis rather pale, glabrescent, 
sparsely warty, residual scales narrow, bearing a 
few setae; scales on costae and costules few, thin, 
brown, ovate or narrower, entire, flat; minute hairs 
on veins abundant and often conspicuous. 



Type specimen: Merrill 6055, Mt Halcon, 
Mindoro, Philippines (P; dupl. at US, MICH, A). 

Distr. Malaysia: Philippines (Mindoro, Luzon). 

Ecol. Probably in forest, at 1200-1700 m. 

Note. The wide pinnules are a character of this 
species; on the type collection are pinnules only 
65 by 20 mm. Copeland reported pinnules 29 mm 
wide on the type of C mearnsii, but I found none 
larger than 110 by 23 mm. 

62. Cyathea ascendens Domin, Acta Bot. Bohem. 
9 (1930) 94. — Alsophila rosenstockii Brause, Bot. 
Jahrb. 56 (1920) 63; Copel. Philip. J. Sc. 77 
(1947) 116. 

Trunk slender; fronds to 100 cm long, well 
spaced, with long-decurrent bases. Stipe 8-12 cm; 
scales on stipe and lower part of rachis to 8 by 
1 mm, dark and shining with fragile pale edges 
bearing long flexuous dark setae. Pinnae 35-40 
pairs, lower ones gradually reduced, longest 8 1/2 by 
1 3,4 cm, almost sessile, pinnatifid nearly to costa; 
costules of pinna-segments 4'/2 mm apart (fertile) 
to 5Y2 mm (sterile); veins to 10 pairs, simple or 
forked. Sori median; no indusia. Scales and hairs: 
main rachis beneath covered with copious crisped 
dark hairs, with a few scales attached to wart- 
like bases, scales very narrow, to 3^ mm long, 
brown with irregular long flexuous marginal setae; 
costae of pinnae similarly hairy and scaly beneath; 
costules of pinna-segments bearing paler and 
sparser hairs on lower surface and a few small 
scales; no bullate scales seen. 

Type specimen: Ledermann 9963, Sepik Region, 
NE. New Guinea (B). 

Distr. Malaysia: NE. New Guinea (2 col- 
lections). 

Ecol. In forest, at 800-1000 m. 

Notes. The collector wrote of the trunk 
'1-2 m lang, krumm'. The herbarium specimen 
shows the apex of the trunk, 14 cm long, straight, 
lacking roots. There is no evidence of climbing 
habit; it seems more likely that the slender trunk 
had fallen, or partly fallen, and that the apical 
part grew erect after the fall, the whole being thus 
crooked. This species seems nearest to C. gregaria, 
but, apart from the simply pinnate condition, the 
rachises are far more hairy on the lower surfaces. 

63. Cyathea recurvata (Brause) Domin, Acta Bot. 
Bohem. 9 (1930) \53.— Alsophila recurvata Brau- 
se, Bot. Jahrb. 56 (1920) 61. 

Trunk to 5 m; fronds of type not over 150 cm 
long {sec. coll. 200 cm). Stipe 10 cm; scales medium 
brown, to 12 by II/2. their fragile edges bearing 
dark setae; pneumathodes 4 mm long, well- 
spaced. Lower pinnae gradually reduced, lowest 
4 cm long, longest 20 cm. Pinnules to 28 by 8 mm 
wide at base, rather suddenly contracted to 5-6 
mm wide, lobed to about half-way to costa; cos- 
tules 21/2 mm apart; veins to 5 pairs in basal seg- 
ments, in others 3 pairs, simple except in basal 
segments. Sori near costules; no indusium; re- 
ceptacle rather elongate; paraphyses short. Scales 
and hairs: lower surface of pinna-rachis bearing 
narrow setiferous scales to 2 mm long and crisped 



102 



Flora Malesiana 



[ser. II, vol. P 



dark hairs; costae similar; no scales seen on cos- 
tules (all pinnules of type are fully fertile). 

Type specimen: Ledermann 9264, Sepik Re- 
gion, NE. New Guinea (B). 

Distr. Malaysia: NE. New Guinea (one col- 
lection). 

Ecol. At 850 m. 

64. Cyathea eriophora Holttum, Kew Bull. 16 
(1962) 55. 

Trunk to 3 m; fronds few, to 225 cm long. 

Stipe 15 cm, dark, with spines to 1 mm long, 
covered throughout with a close felt of very small 
pale scales, the larger with dark setiform apex; 
large scales abundant on stipe and base of rachis, 
to 20 by 2 mm, narrowed to twisted tip, shining 
dark brown with narrow fragile edges bearing 
many long dark setae. Lower surface of main 
rachis smooth, medium brown, with dense felt of 
pale crisped hairs and scattered long narrow dark 
setiferous scales. Lower pinnae gradually reduced, 
lowest 5-8 cm long, longest 30-42 cm. Pinnules to 
75 by 18 mm, sessile, short-acuminate, lowest 
segment almost free, rest of pinnule lobed nearly 
to costa; costules 3-3 '/2 rnm apart; veins 8-9 pairs; 
lamina-segments rather thin, almost entire or 
lowest ones on lower pinnules deeply crenately 
lobed. Sori near costules, without indusia; re- 
ceptacle swollen; paraphyses short. Scales and 
hairs: lower surface of pinna-rachis densely covered 
with long tangled pale crisped hairs, also many 
narrow dark scales with pale edges bearing long 
setae; costae similar; costules bearing pale bullate 
scales, usually hair-pointed; upper surface of pinna- 
rachis and costae covered with dark hairs, also on 
pinna-rachis some scales as on lower surface. 

Type specimen: Carr 14439, Boridi, Papua (K; 
dupl. at BM, L). 

Distr. Malaysia: Eastern New Guinea (3 col- 
lections). 

Ecol. In wet ravine in forest, locally common, 
at 1400-1950 m. 



65. Cyathea gregaria (Brause) Domin, Acta Bot. 
Bohem. 9 (1930) 120. — Alsophila gregaria Brause, 
Bot. Jahrb. 56 (1920) 68. 

Trunk 4-5 m, 'arm-thick'. Stipe 40 cm; spines to 
1 mm long; scales few, to 15 by 1 mm, medium 
brown with pale edges bearing long flexuous 
setae. Lowest pinna 17 cm long, longest 32 cm. 
Pinnules to 60 by 15 mm, lobed almost to costa; 
costules 3'/2 mm apart; veins to 9 pairs; lamina- 
segments crenate, sinuses rather wide. Sori near 
costules, without indusia; receptacle rather high. 
Scales and hairs: lower surface of pinna-rachis 
bearing rather sparse dark crisped hairs; on lower 
surface of costae dark crisped hairs and some dark 
narrow scales with scattered marginal setae; on 
costules similar hairs and a few scales. 

Type specimen: Ledermann 8596, Sepik 
Region, NE. New Guinea (B). 

Distr. Malaysia: Eastern New Guinea (one 
collection). 

Ecol. In forest, growing in groups, 100 m. 



66. Cyathea modesta (Bak.) Corel. Philip. J. Sc. 4 
(1909) Bot. A^.— Alsophila modesta Bak. J. Bot. 18 
(1880) 210; V. A. v. R. Handb. (1908) 37.— 
Hemitelia singalanensis v. A. v. R. Bull. Jard. 
Bot. Btzg II, n. 16 (1914) 15; Handb. Suppl. (1917) 
43. — Hemitelia confluens v. A. v. R. Bull. Jard. 
Bot. Btzg II, //. 16(1914) 14; Handb. Suppl. (1917) 
49. — Hemitelia subconfluens v. A. v. R. Bull. Jard. 
Bot. Btzg II, n. 28 (1918) 25.— C. singalanensis 
Domin, Pterid. (1929) 264.— C. confluens Domin, 
I.e. 263. — C. subconfluens Domin, Acta Bot. 
Bohem. 9 (1930) 162. 

Stipe 35-50 cm or more; spines bluntly conical, 
hardly 1 mm high; scales abundant throughout, 
to 30 by 1 1/2 mm, medium brown with narrow con- 
colorous fragile edges; also more or less abundant 
hairs at rachis. Main rachis covered with a close 
felt of crisped pale hairs, also with more or less 
caducous very narrow entire crisped brown scales 
5-10 mm long. Pinnae to 40 cm or more long. 
Pinnules to 60 by 14 mm, sessile, abruptly pointed, 
one to several basal segments almost free; costules 
3-4 mm apart; veins 7-10 pairs; lamina-segments 
firm, crenate, or the basal ones more deeply lobed, 
sinuses (apart from basal ones) narrow. Sori near 
costules; indusium hemitelioid, firm, brown, some- 
times semicircular, or smaller and a little bilobed, 
hidden or almost hidden by ripe sorus; paraphyses 
as long as sporangia, sometimes widened at base. 
Scales and hairs: pinna-rachis densely covered 
throughout with crisped hairs, very narrow crisped 
entire brown scales at first abundant, also distally 
smaller scales bullate at base; at base of costae 
narrow acuminate entire scales, flat or bullate at 
base, grading to bullate-acuminate scales, some of 
which occur throughout costa; scales on costules 
bullate, often hair-pointed. 

Type specimen: Beccari 434, G. Singgalang, 
Sumatra (K; dupl. at FI). 

Distr. Malaysia: Sumatra. 

Ecol. In forest at 1800-2400 m. 

Note. The type of Alsophila modesta was an 
unusually small specimen, with pinnules to 
40 by 1 1 mm. 

67. Cyathea doctersii v. A. v. R. Bull. Jard. Bot. 
Btzg III, 2 (1920) 136. 

Stipe-h'dse not seen. Main rachis and pinna- 
rachis glabrous on the median raised part of the 
upper surface (some hairs present in the small 
groove on each side of this), hairy throughout on 
lower surface. Pinnae to 45 cm long. Pinnules to 
75 by 17 mm, lobed almost to the costa, almost 
sessile, shortly pointed; costules 31/2-4 mm apart; 
veins to 8 pairs; lamina-segments almost entire. 
Sori near costules; indusium a thin brown scale of 
varied shape and size on costular side, often 
2-lobed, sometimes encircling the base of the recep- 
tacle. Scales and hairs: scales near base of costae 
elongate, entire, flat, grading to similar scales 
bullate at the base and to bullate-acuminate; hairs 
also present on lower surface of costae, not on up- 
per surface. 

Type specimen: Docters van Leeuwen 3265, 
Deli, Sumatra (BO; dupl. at L). 



Dec. 1963] 



Cyatheaceae (Holttum) 



103 



Distr. Malaysia: Sumatra (one collection). 

Ecol. Probably in forest, at 150 m. 

Note. This is near C. javanica in character and 
distribution of hairs and scales, but almost all 
indusia are hemitelioid; it is possibly a hybrid 
between C. javanica and one of the species of the 
C. latebrosa alliance. 

68. Cyathea cucullifera Holttum, Kew Bull. 16 
(1962) 54. 

Fronds 150-175 cm long, in 2 whorls of 4-6 
each. Stipe 15 cm, warty, densely scaly throughout; 
larger scales all round base of stipe, along sides 
above base, to 20 by 1/2 mm, dark, shining, with 
narrow pale edge bearing scattered long flexuous 
dark setae; small scales forming a dense felt over 
whole abaxial surface of stipe, dark brown, larger 
ones setiferous; rac/iis pale brown, finely warty, 
with sparse covering of very small pale brown 
scales. Lower pinnae gradually reduced, lowest less 
than 5 cm long, largest 30 cm. Pinnules to 40 mm 
long and 10 mm wide (sterile), 6-8 mm wide 
(fertile), sessile, short-acuminate, lobed nearly to 
costa, lowest segment not quite free; costules 3 mm 
apart (sterile), 2-2 Vi mm (fertile); veins 8-9 pairs 
(sterile), 6 pairs (fertile); lamina-segments firm, 
close, nearly entire. Sori close to costule; indusium 
a pale brown scale backing the costule, concave 
towards sorus, when flattened usually more than a 
semicircle (rarely spreading round base of recep- 
tacle); paraphyses slender, short. Scales and hairs: 
lower surface of pinna-rachis covered with in- 
terlacing crisped hairs and very small scales with a 
crisped hair-tip, also scattered elongate flat scales 
bearing a few long marginal setae; at base of 
costae some (usually deciduous) narrow setiferous 
scales, also very small pale scales, some with long 
flexuous hair-tips, grading to crisped hairs and to 
very small scales, distally to small pale buUate 
scales; on costules very small pale hair-tipped 
scales and pale bullate scales; on veins abundant 
very short appressed hairs (bases of scales?). 

Type specimen: Hoogland & Pullen 5497, 
Eastern Highlands, NE. New Guinea (K; dupl. 
at L, BO, US, SYD, BRI). 

Distr. Malaysia: E. New Guinea (4 collections). 

Ecol. Common in mountain forest, at c. 2400 m. 

69. CyatheasetulosaCoPEL. Philip. J. Sc. 81 (1952) 
14; Fern Fl. Philip. 2 (1960) 212. 

Stipe not known. Pinnae 45 cm long. Largest 
pinnules 90 by 17 mm, sessile, shortly caudate, 
basal 1-2 segments free, then 1-2 pairs separately 
adnate to costa, rest of pinnule lobed nearly to 
costa; costules 3 'A mm apart; veins 9-11 pairs; 
lamina-segments almost entire except the lower 
fertile ones which are sometimes deeply crenate 
near base. Sori near costules; indusium dark brown 
hemitelioid, more than half covering mature sorus, 
at length reflexed and backing the costule; para- 
physes short. Scales and hairs: lower surface of 
pinna-rachis densely covered with dark shining 
crisped hairs, with a few dark very narrow entire 
scales; costae hairy beneath as pinna-rachis 
except near apex, scales few, small, not setiferous; 



no bullate scales seen on costae and costules. 

Type specimen: Alcasid & Edano, PNH 5068, 
Mt Camatis, Quezon Province, Luzon (UC). 

Distr. Malaysia: Philippines (Luzon). 

70. Cyathea muelleri Bak. J. Bot. 28 (1890) 104; 
V. A. V. R. Handb. (1908) 25; Copel. Philip. J. Sc. 
77 (1947) 105.— C. longipaleata Alston, J. Bot. 78 
(1940) 226; Nova Guinea n.s. 4 (1940) 110, t. 4, 
f. 2, t. 5, f. 3.— Fig. 16. 

Trunk to 10 m, 15-20 cm 0, bearing fronds in 
whorls of 10-12, usually 2 whorls present; fronds 
c. 100 cm long, apices upcurved. Stipe 10-12 cm; 
spines to 1 mm long; scales rigid, :- twisted, 50 by 
1 Y2 mm, shining brown, fragile edges very narrow. 
Lower pinnae gradually reduced, lowest 4-5 cm 
long; longest pinna 20 cm. Largest pinnules 
40-50 by 10-16 mm, sessile, shortly pointed, 
lowest 1-2 segments free, then c. 3 pairs con- 
stricted at base on acroscopic side and decurrent 
on basiscopic side; costules 3-3 '/2 mm apart; 
veins 7 pairs, flat or slightly grooved on both 
surfaces; lamina-segments very rigid, edges slightly 
reflexed and minutely crenate, or lower ones 
distinctly lobed if fertile. Sori usually to 6 pairs on 
a segment; indusium very firm, brown, almost 
covering sorus to maturity but open on side remote 
from costule; receptacle swollen, sporangia very 
numerous; paraphyses short, slender. Scales and 
hairs: pinna-rachis glabrescent and finely warty; 
scales on costae and costules early caducous, some 
residual scales on costae broad and flat, some very 
narrow and setiferous; costules of sterile pinnules 
sometimes bearing dark brown entire ovate con- 
vex to almost bullate scales. 

Type specimen: W. McGregor 62, Mt Knuts- 
ford, Papua (K; dupl. at BM, MEL). 

Distr. Malaysia: New Guinea. 

Ecol. At 3500-3600 m, 'common in marginal 
scrub of subalpine forest and drier more sheltered 
grass slopes; one or several stems from a common 
base' (Brass). 

71. Cyathea oinops Hassk. in Hook. J. Bot. Kew 

Misc. 7 (1855) 322; Obs. Fil. Jav. 1 (1856) 23; 
V. A. V. R. Handb. (1908) 25; Suppl. (1917) Corr. 
43; Backer & Posth. Varenfl. Java (1939) 25, 
p.p. — C. oligocarpia Jungh. Nat. Geneesk. Arch. 
N.L 2 (1845) 39 {non Kunze, 1834).— C. zollinge- 
riana {non Mett.) v. A. v. R. Handb. (1908) 20, 
785; Suppl. (1917) 26, Corr. 42.— C. crenulata f. 
squamulosa v. A. v. R. Bull. Jard. Bot. Btzg II, 
n. 20 (1915) 10; Handb. Suppl. (1917) 27, Corr. 63. 
— C crenulata f. latissima v. A. v. R. Bull. Jard. 
Bot. Btzg II, n. 28 (1918) 11; Handb. SuppL 
(1917) Corr. 64. — C.faberiana DoMiN, Acta Bot. 
Bohem. 9 (1930) 114.— Fig. 8a, 9d. 

Stipe c. 35-60 cm, sometimes with a pair of 
small pinnae near base, dark, warty, base covered 
with firm pale scales to 35 by 3 mm, rest with 
rt abundant very small scurfy scales or glabres- 
cent. Lower pinnae more or less reduced, longest 
40-55 cm long. Largest pinnules 70-100 by 15-20 
mm wide just above base, sessile, shortly acumi- 
nate, basal segments largest, 2-3 pairs often 



104 



Flora Malesiana 



[ser. II, vol. P 




Fig. 16. Cyathea muelleri Bak. (Hoogland 5707). 

Mt Wilhelm, Eastern Highlands, E. New Guinea, 

3200 m (R. D. Hoogland). 

separately adnate to costa; costules 3-4 mm apart; 
veins 10-12 pairs; lamina-segments firm, where 
sterile finely crenate-serrate, where fertile lobed I3 
or more towards costule, lobes bidentate. Sort 
near costules; indusium firm, brown, covering the 
sorus almost to maturity but open on side remote 
from costule; paraphyses slender, shorter than 
sporangia. Scales and hairs: pinna-rachis more or 
less glabrescent, finely warty, often with a persis- 
tent covering of very small, dull, pale brown 
irregularly short-fringed scales, with some scattered 
larger elongate scales; costal scales abundant, 
often closely overlapping, elongate, brown with 
pale thinner edges bearing scattered dark setae 
especially near apices, also smaller thin pale fringed 
scales; on costules thin brown short-fringed 
scales, ovate to narrower and acuminate, almost 
flat to convex, sometimes with a few setae. 

Type specimen: Hasskarl, Mt Gedeh, W. Java 
(BO; dupl. at L). 



Distr. Ma/aji/a: Sumatra, Java, Lesser Sunda Is 
(Lombok), SW. Celebes. 

E col. At 2000-2500 m, in forest (specimen from 
Lombok in Casuarina forest). 

Note. In Fl. Btzg 1 (1898) 36, Raciborski 
described a species under the name Cyathea sinops 
Hassk., which appears to have been a misprint for 
oinops. Raciborski's specimen was, however, 
from an unusually large frond of C cremdata Bl., 
which see for further references. 

72. Cyathea loheri Christ, Bull. Herb. Boiss. II 
6 (1906) 1007; v. A. v. R. Handb. (1908) 787 
Tagawa, Act. Phytotax. Geobot. 14 (1951) 94 
COPEL. Fern Fl. Philip. 2 (1960) 2\\.—C.fnic- 
tuosa CoPEL. in Elmer, Leafl. Philip. Bot. 2 (1908) 
419; V. A. v. R. Handb. Suppl. (1917) 37; Copel. 
Fern Fl. Philip. 2 (1960) 207.— C. mitrata Copel. 
Philip. J. Sc. 3 (1909) Bot. 354; ibid. 4 (1909) Bot. 
42; v. A. V. R. Handb. Suppl. (1917) 37; Copel. 
Fern Fl. Philip. 2 (1960) 211.— C. campbellii 
Copel. Philip. J. Sc. 38 (1929) 132; Fern Fl. 
Philip. 2 (1960) 210.— C. korthalsii (non Mett.) 
C. Chr. Card. Bull. S. S. 7 (1934) 222.— C. in- 
dusiosa Copel. Philip. J. Sc. 81 (1952) 14; Fern 
Fl. Philip. 2 (1960) 211. 

Trunk to 10 m. Slipe short; scales to 25 by 
l'/4-3 mm, pale, firm, their fragile edges bearing 
scattered long dark setae; pneumathodes to 1 1 mm 
long, in one row. Lower pinnae gradually reduced, 
lowest c. 7 cm long; longest pinnae to 40 cm or 
more. Largest pinnules 75-95 by 15-19 mm, 
sessile, short-acuminate, lowest 1-4 segments more 
or less contracted at base, rest of pinnule lobed 
nearly to costa; costules 3'/2^ mm apart; veins 
10-12 pairs; lamina-segments firm, crenate or the 
larger ones more deeply lobed where fertile. Sori 
near costules; indusium firm, shining brown to 
purplish, almost covering sorus to maturity but 
open on side remote from costule, more or less 
breaking when old. Scales and hairs: pinna-rachis 
finely warty, at first densely scaly, many scales 
usually persistent; small ones rusty, dull, short- 
fringed, larger ones pale with some dark setae; 
on costae many scales, lower ones rather pale 
brown, acuminate with setae or hairs on edges, 
grading to numerous bullate scales; bullate scales, 
sometimes with dark setae, present on costules. 

Type specimen: Loher s.n., 1 Jan. 1906, Mt 
Banajao, Luzon (not found at P; dupl. at S-PA). 

Distr. Formosa, in Malaysia: Philippines 
(Luzon, Negros, Mindanao), N. Borneo. 

Ecol. In forests, 600-2500 m. 

Notes. This species is closely allied to C. oinops, 
but differs in bullate scales and apparently in the 
lower pinnae always gradually reduced. Young 
plants (as seen by me on Mt Kinabalu) have long- 
stalked fronds. Possibly C. oinops and C. loheri 
should be united; C. oinops is the older name. 

73. Cyathea cinerea Copel. in Elmer, Leafl. 
Philip. Bot. 5 (1913) 1681; v. A. v. R. Handb. 
Suppl. (1917) 36; Copel. Fern Fl. Philip. 2 (1960) 
215. 

Trunk 5 m, 1 2 cm 0. Stipe bearing spines 8 mm 



Dec. 1963] 



Cyatheaceae (Holttum) 



105 



long {fide CoPEL., not seen); length of stipe not 
recorded. Main rachis spiny near base, spines to 
3 mm long. Pinnae to 55 cm long. Largest sterile 
pinnules 90 by 18 mm, fertile 13 mm wide, sessile, 
acuminate, lobed almost to costa, basal segment 
not free; costules 3-3 '/a mm apart; veins 12 pairs; 
lamina-segments firm, crenate-serrate, sterile ones 
close, fertile narrower and separated by sinuses 
1 mm wide. Sori near costules; indusium almost 
covering sorus to maturity but open on side remote 
from costule; paraphyses short, slender. Scales and 
hairs: pinna-rachis pale, with a few slender spines 
1/2 mm long, persistently but sparsely covered with 
irregular pale scales less than 1 mm long and a 
few narrow entire brown scales to 3 mm long; 
scales on costae dull brown, flat, elongate, rarely 
with a tew long dark setae, also very small scales 
as on pinna-rachis; on costules (of sterile pinnules) 
pale entire buUate-acuminate scales. 

Type specimen: Elmer 13860, Mt Urdaneta, 
Agusan Province, Mindanao (US; dupl. at MICH, 
K, BO, P, A, UC, L, U, BM). 

Distr. Malaysia: Philippines (Mindanao, one 
collection). 

Ecol. At 1050 m. 

74. Cyathea pachyrrhachis Copel. Un. Cal. Publ. 
Hot. 18 (1942) 218; Philip. J. Sc. 77 (1947) 107, pi. 5. 

Trunk to 7 m, 7'/2 cm 0, stipe-bases not per- 
sistent, scars in alternate whorls of 5; fronds 
10 (or 5), 150-300 cm long. Stipe 30-40 cm, 
copiously warty; scales pale or partly dark, to 
20 by 1 mm, with narrow fragile edges, not setifer- 
ous, also very small dull pale irregular scales. 
Lowest pinnae 20-25 cm long, longest 40-60 cm. 
Largest pinnules 60-100 by 15-20 mm, sessile, 
acuminate, 1-2 pairs basal segments free, rest of 
pinnule lobed nearly to costa; costules 3-4 mm 
apart; veins to 12 pairs (sterile), 9 pairs (fertile); 
lamina-segments very firm, sterile ones crenate, 
fertile rather deeply lobed, lobes bifid; sterile 
segments contiguous, fertile separated by sinuses. 
Sori near costules; indusium firm, covering sorus 
to maturity but open on side remote from costule, 
breaking somewhat when old. Scales and hairs: 
pinna-rachis more or less glabrescent, residual 
scales setiferous, very small with a few elongate 
narrow dark ones; costae rather densely scaly 
near base, some scales to 3 mm long, narrow, 
very dark, shining, with thin pale edges bearing a 
few setae near apices, grading to entirely pale 
setiferous scales, smaller ones all flat, ovate to 
nearly circular; similar scales on costules and 
rarely also on veins. 

Type specimen: Brass 12118, Idenburg River, 
W. New Guinea (MICH; dupl. at BO, BM, UC, 
L, A). 

Distr. Malaysia: New Guinea (several col- 
lections), d'Entrecasteaux Is (Goodenough I.). 

Ecol. In forest, 1000-2850 m. A specimen from 
secondary Nothofagus forest at 2060 m (Brass 
29674) has small fronds (pinnules 60 mm long), 
the smaller scales all strongly dark-setiferous, in- 
cluding those on veins, the latter being abundant, 
another (303 16j from mossy forest at 2770 m, had 



fronds of similar size, scales on veins rarely se- 
tiferous; Brass 30668, from 2850 m, has the 
broadest pinnules, with several pairs of basal 
segments almost free. 

75. Cyathea latipinnula Copel. in Elmer, Leafl. 
Philip. Bot. 4 (1911) 1149; Fern Fl. Philip. 2 (1960) 
226. — Hemitelia latipinnula v. A. v. R. Handb. 
Suppl. (1917) 52. 

Trunk 120 cm. 10 cm o; fronds 180 cm long. 
Stipe 60 cm long (fide Copel.), scales not seen; 
main rachis glabrescent, bearing numerous spines 
less than 1 mm long. Pinnae to 40 cm long. Pinnules 
to 120 by 40 mm, caudate-acuminate, lower ones 
somewhat shorter and on stalks to 7 mm long, 
lowest 1-2 segments of larger pinnules quite free, 
next 1-2 pairs constricted at base; costules 5-6 mm 
apart; veins to 12 or more pairs, basal basiscopic 
vein of each group attached at very base of costule; 
lamina-segments very firm, tapering and crenate 
towards apices, free basal ones sometimes deeply 
lobed. Sori near costules; indusium hemitelioid, 
small, dark, outer edge uneven, not reaching cos- 
tule and hidden by mature sorus; receptacle 
swollen; paraphyses as long as sporangia, some 
several cells wide at base. Scales and hairs: pinna- 
rachis minutely spiny, also bearing short crisped 
hairs and a few residual narrow brown scales which 
may be setiferous; a few narrow scales at base of 
costae, most being ovate-acute, grading to bullate; 
entire bullate scales on costules. 

Type specimen: Elmer 12512, Sibuyan Island 
(MICH; dupl. at US, K, FI, P, A, SYD. BO. BM). 

Distr. Malaysia: Philippines (Sibuyan Island, 
one collection). 

Ecol. On windy ridge at 1400 m. 

76. Cyathea masapilidensis Copel. Philip. J. Sc. 81 
(1952) 17; Fern Fl. Philip. 2 (1960) 227. 

Stipe rather slender, bearing close sharp slender 
spines to 5 mm long; scales not seen. Lower 
pinnae unknown; middle pinnae 50 cm long. 
Largest pinnules 65-80 by 16-18 mm, acuminate, 
on stalks to 6 mm long, lobed almost to costa, 
lowest segment sometimes free; costules 3'/2 mm 
apart; veins 10 pairs; lamina-segments very firm, 
nearly entire. Sori near costules; indusium thin, 
dull brown, covering about half of sorus at ma- 
turity, hemitelioid. Scales and hairs: pinna-rachis 
glabrescent, residual scales very narrow, dark, 
with long marginal setae; scales on costae sparse, 
flat, brown with pale margins bearing some setae, 
grading to very small flat scales; no costular scales 
seen, and none bullate. 

Type specimen: Ramos & Edano, BS 37858, 
Mt Masapilid, Bontoc Subprov., Luzon (MICH; 
dupl. at US, BO). 

Distr. Malaysia: Philippines (Luzon, 2 col- 
lections). 

77. Cyathea loerzingii Holttum, Kew Bull. 16 
(1962) 58. 

Stipe at least 40 cm, rather sparsely warty near 
base; persistent scales few, to 20 by 2V2 mm, 
shining dark brown with rather broad paler fragile 



106 



Flora Malesiana 



[ser. II, vol. P 



edges; pneumathodes 10-15 mm long, in an irregu- 
lar double row. Pinnae probably to 50 cm long 
(only upper ones seen). Pinnules to 100 by 18 mm, 
lowest on stalks to 7 mm long, apex acuminate, 
lowest 1-2 segments not free but on larger pinnules 
separated by a narrow wing from the rest; costules 
4 mm apart; veins 10-11 pairs; lamina-segments 
firm, drying very dark on upper surface, edges 
finely crenate, lowest ones not deeply lobed. Sori 
near costules; indusium at maturity firm, brown, 
semicircular, reflexed against costule, c. 1 mm wide; 
receptacle swollen, bearing at its apex a small 
group of scales 4-5 cells wide (bases of paraphyses?), 
other paraphyses short, slender. Scales and hairs: 
lower surface of pinna-rachis smooth, glabrescent; 
scales near bases of costae elongate, shining brown 
with a few hairs or dark setae on margins, grading 
to acuminate bullate-based scales distally; on cos- 
tules brown bullate scales, often acuminate. 

Type specimen: Lorzing 14904, Mt Sibajak, 
Sumatra (L; dupl. at BO). 

Distr. Malaysia: Sumatra (one collection). 

Ecol. In forest at 1300-1400 m. 

78. Cyathea rufopannosa Christ, Philip. J. Sc. 2 
(1907) Bot. 180; v. A. v. R., Handb. (1908) 784; 
CoPEL. Fern Fl. Philip. 2 (1960) 212. 

Trunk (dry) 4 cm o. Stipe 40-50 cm, sometimes 
with a pair of small pinnae near base; base warty; 
scales 15 by 2 mm, castaneous or paler, fragile 
edges narrow. Largest pinnae 'il cm long. Pinnules 
to 65 by 13-15 mm, sessile, short-acuminate, sev- 
eral pairs of lower segments contracted at base, 
lowest 1-3 quite free; costules 21/2-3 mm apart; 
veins 8 pairs; lamina-segments firm, crenate. Sori 
near costules; indusium hemitelioid, more than a 
semicircle, covering costular side of sorus at 
maturity; receptacle rather slender; paraphyses 
long, often broad at the base. Scales and hairs: 
pinna-rachis densely and persistently scaly beneath; 
scales of all sizes, largest 6 by 1 mm, flat, entire, 
light brown, smaller ones bullate at base; at bases 
of costae many elongate flat light brown entire 
scales, grading to bullate (some bullate to base of 
costa); bullate scales on costules. 

Type specimen: Copeland 1730, San Ramon, 
Mindanao (P; dupl. at MICH); also from same 
locality Copeland 1735 (P, US, SYD, S-PA). 

Distr. Malaysia: Philippines (Mindanao). 

Ecol. Probably in forest, 1200 m. 

79. Cyathea callosa Christ, Bull. Herb. Boiss. II, 
6 (1906) 1008; v. A. v. R. Handb. (1908) 787; 
CoPEL. Fern Fl. Philip. 2 (1960) 222, excl. syn. 
Hemitelia caudiculata Rosenst. — C. foxworthyi 
CoPEL. Philip. J. Sc. 3 (1909) Bot. 355; v. A. v. R. 
Handb. Suppl. (1917) 35; Copel. Fern Fl. Philip. 
2 (1960) 220. — C camaguinensis Copel. Philip. J. 
Sc. 81 (1952) 16; Fern Fl. Philip. 2 ( 1960) 223. 

Stipe to c. 15 cm, copiously short-spiny (spines 
c. 2 mm); scales 15-20 by 1 mm, dark with pale 
fragile edges. Lower pinnae gradually reduced, 
lowest commonly less than 10 cm long; largest 
pinna 40 cm long or more. Pinnules to 100 by 24 
mm, sessile, acuminate, lowest segment more or 



less free, rest of pinnule lobed nearly to costa; 
costules 4^'/2 miTi apart; veins to 12 pairs; lamina- 
segments firm, crenate, lowest ones sometimes 
deeply so. Sori near costules; indusium thin and 
pale except near receptacle, almost covering sorus 
to maturity but not closed on side remote from 
costule, at maturity breaking and the thinner parts 
sometimes caducous, remnant then reflexed against 
costule; paraphyses not longer than sporangia. 
Scales and hairs: pinna-rachis glabrescent, rather 
pale, bearing scattered small slender spines or 
warts (lower pinnae most spiny); costal scales 
sparse, dull brown, rather broad, sometimes with 
a dark seta near apex; costular scales few, ovate, 
flat, pale, not bullate but sometimes convex; on 
lower surface of veins many very short appressed 
but conspicuous hairs. 

Type specimen: Loher, Mt Maquiling, Luzon, 
April 1906 (not seen at P; dupl. at M). 

Distr. Malaysia: Philippines (Luzon). 

Ecol. In mid-mountain forest; few records of 
altitude in Luzon. 

Note. This species is very near C. spinulosa 
Wall, which is widely distributed from the NE. 
Himalayas to S. China and Formosa (syn. C. 
austrosinica Christ and C taiwaniana Nakai), 
but appears to differ in short less spiny stipe and 
in gradually reduced lower pinnae. 

80. Cyathea dicksonioides Holttum, Blumea 11 
(1962) 529.— Fig. 10c, d, 17. 

Trunk to 3 m, 20 cm o, bearing fronds in 2 whorls 
of 10-12 each; fronds to 90 cm long, those of outer 
whorl almost straight, those of inner whorl bent 
downwards near the base. Stipe to 10 cm, not 
spiny, covered with scales; scales 45 mm by 2 mm 
wide at base, shining, castaneous with narrow 
paler fragile edges, straight, the finely acumi- 
nate apical part much twisted; sparse small 
scales beneath the larger ones. Lowest pinnae 
rather abruptly reduced, 8-10 cm long, longest 
pinnae 17-20 cm. Pinnules close, largest 35 mm 
long, sessile, 5 mm wide above the widened base, 
pinnate almost throughout; midribs of tertiary 
leaflets 2 mm apart. Tertiary leaflets contiguous, 
almost triangular with rounded tip, edges strongly 
reflexed, basal leaflets to almost 4 mm long and 
distinctly lobed; veins to 4 pairs, lower 1-2 pairs 
forked, not raised on upper surface, pale and strong- 
ly raised on lower surface. Sori 2-4 on each leaflet; 
indusium pale, firm, attached on costular side and 
forming a hood which partly covers the mature 
sorus, in shape very much like the inner indusium 
of Dicksonia; receptacle swollen; sporangia many; 
no paraphyses seen. Scales and hairs: all rachises 
and costae minutely warty and glabrescent on 
lower surface, antrorse hairs on upper surface pale; 
residual scales on rachises long, pale brown, very 
narrow, entire; on lower surface of midribs of 
tertiary leaflets a few spreading light-brown hair- 
tipped scales. 

Type specimen: Hoogland & Schodde 7171, 
Western Highlands, NE. New Guinea (L). 

Distr. Malaysia: NE. New Guinea (2 col- 
lections). 



Dec. 1963] 



Cyatheaceae (Holttum) 



107 




Fig. 17. Cyathea dicksonioides Holttum (small plant in centre) with taller C. atrox C. Chr., in tree-fern 
grassland, N. slopes of Sugarloaf complex, Western Highlands, Territory of New Guinea, 2880 m 

(R. D. Hoogland). 



108 



Flora Malesiana 



[ser. II, vol. 12 



Ecol. In tree-fern grassland, infrequent, 2600- 
2900 m. 

Note. In form of leaflets and in basal scales this 
is very like C. macgregorii, but it is quite different 
in indusium and costular scales, and in habit of 
growth, the fronds being in two whorls, those of 
the inner whorl bent downwards at the base. 

81. Cyathea heterochlamydea Copel. in Elmer, 
Leafl. Philip. Bot. 2 (1908) 418; Fern Fl. Philip. 2 
(1960) 218. — Hemitelia heterochlamydea v. A.v. R. 
Handb. Suppl. (1917) 53. — Hemitelia caudiculata 
RosENST., Med. Rijksherb. 31 (1917) l.—C. cau- 
diculata DoMiN, Acta Bot. Bohem. 9 (1930) 104.— 
C. merrillii Copel. Philip. J. Sc. 46 (1931) 212; 
Fern FI. Philip. 2 (1960) 231.— Fig. 9c. 

Stipe short, warty or short-spiny; pneumathodes 
in 2-3 irregular rows; scales dark, shining, with 
narrow (often abraded) paler fragile edges. Lower 
pinnae gradually reduced; longest to 60 cm long. 
Pinnules commonly to 100 by 18 mm, exceptionally 
to 1 20 by 23 mm, sessile, strongly acuminate, lowest 
segment often free, on largest pinnules 2-3 pairs 
segments constricted on acroscopic base, de- 
current basiscopically, most pinnules lobed almost 
to costa throughout with narrow sinuses between 
segments; costules 3'/2-4 mm apart; veins com- 
monly 10-1 1 pairs, on largest pinnules to 14 pairs; 
lamina-segments firm, crenate, basal free ones 
sometimes deeply so. Sort near costules; indusium 
firm, brown, overarching costular side of sorus at 
maturity, its edge firm, open on side remote from 
costule; receptacle rather tall, slightly swollen; 
paraphyses short. Scales and hairs: lower surface 
of pinna-rachis finely warty, glabrescent or bearing 
very small irregular short-fringed pale brown scales 
and sometimes sparse crisped hairs; lower surface 
of costae usually with numerous very small ir- 
regular short-fringed scales and sometimes also 
short crisped hairs, larger scales usually all de- 
ciduous, residual ones flat, dull brown, with seti- 
form apex and sometimes 1-2 other setae; 
costules of fertile segments usually without scales, 
on sterile ones a few ovate thin convex or just 
bullate scales. 

Type specimen: Elmer 9742, Cuernos Mts, 
Negros (MICH; dupl. at US, FI, BO, K, P, A, 
SYD, U, L, BM). 

Distr. Malaysia: Philippines (Luzon, Negros, 
Panay, Mindanao). 

Ecol. Little information; apparently in mid- 
mountain forest. 

Note. Apart from size of indusium and larger 
pinnules, there seems little distinction between 
this and C. caudata (J. Sm.) Copel. 

82. Cyathea edanoi Copel. Philip. J. Sc. 46 (1931) 
211; Fern FI. Philip. 2 (1960) 219. 

Stipe c. 5 cm; scales dark, rather narrow with 
narrow fragile edges. Lower pinnae gradually 
reduced and close together, lowest 5 cm long; 
longest pinnae 40 cm. Pinnules to 60 by 14 mm, 
almost sessile, shortly caudate-acuminate, lowest 
2 segments almost free, most of remaining seg- 
ments constricted at base on acroscopic side, de- 



current basiscopically; costules 3-3 1/2 mm apart; 
veins 8 pairs; lamina-segments rather thin, edges 
crenate. Sori near costules; indusium large, he- 
mitelioid, covering at least half sorus on costular 
side at maturity, rather firm and dark. Scales and 
hairs: lower surface of pinna-rachis glabrescent; 
lower surface of costae bearing few flat light- 
brown scales, sometimes with a long seta; on 
costule no scales seen (all fertile). 

Type specimen: Edano BS 78709, summit of 
Mt Cagua, Luzon (MICH; dupl. at BO). 

Distr. Malaysia: Philippines (Luzon, 3 col- 
lections). 

Ecol. At 1300 m. 

Note. This is very closely related to C hete- 
rochlamydea, having similar sori and scales, and 
may perhaps be a small form of that species due to 
habitat conditions at or near a mountain-summit. 

83. Cyathea fuliginosa (Christ) Copel. Philip. J. 
Sc. 4 (1909) Bot. 43; Fern Fl. Philip. 2 (1960) 224. 
— Alsophila fuliginosa Christ, Bull. Herb. Boiss. 6 
(1898) 138; V. A. v. R. Handb. (1908) 39.— Also- 
phila mindanensis Christ in Warb. Monsunia 
(1900) 90, p.p. (fertile specimen; sterile is Dickso- 
nia); v. A. v. R. Handb. (1908) 44.— C. loheri var. 
tonglonensis Christ, Philip. J. Sc. 2 (1907) Bot. 
180; v. A. V. R. Handb. (1908) 1%1.—C. lanaensis 
Christ, Philip. J. Sc. 3 (1908) Bot. 271; v. A. v. R. 
Handb. Suppl. (1917) 23; Copel. Fern Fl. Philip. 
2 (1960) 219.— C. mindanensis Copel. Philip. J. Sc. 
4 (1909) Bot. 34; Fern Fl. Philip. 2 (1960) 223.— 
C. bicolor Copel. in Elmer, Leafl. Philip. Bot. 3 
(1910) 804; Fern Fl. Philip. 2 (1960) 225.— 
Hemitelia tonglonensis v. A. v. R. Bull. Jard. Bot. 
Btzg II, n. 7 (1912) 14; Handb. Suppl. (1917) 42. 
— C warihon Copel. in Elmer, Leafl. Philip. Bot. 5 
(1913) 1680; Fern Fl. Philip. 2 (1960) 226.— 
Alsophila warihon C. Chr. Ind. Fil. Suppl. 2(1917) 
4. — Hemitelia bicolor v. A. v. R. Handb. Suppl. 
(1917) 44. — Hemitelia warihon v. A. v. R. I.e. 43. 
— C. tonglonensis Domin, Pterid. (1930) 264.— 
C. squamicosta Copel. Philip. J. Sc. 46 (1931) 212; 
Fern Fl. Philip. 2 (1960) 209.— C. dura Copel. 
Philip. J. Sc. 81 (1952) 13; Fern Fl. Philip. 2 (1960) 
206.— C. lepidigera Copel. Philip. J. Sc. 81 (1952) 
16; Fern Fl. Philip. 2 (1960) 224.— C. sulitii 
Copel. Philip. J. Sc. 81 (1952) 18; Fern Fl. Philip. 
2 (1960) 225.— C. biliranensis Copel. Philip. J. Sc. 
84(1955) 162. 

Trunk to at least 5 m. Stipe c. 1 cm, closely warty 
or with conical spines under 1 mm long; scales to 
15 by 1 '/2 mm. dark, shining, with fragile concol- 
orous edges. Lower pinnae gradually reduced, 
lowest c. 1 cm long; longest pinna 40-50 cm. 
Largest pinnules commonly 60-75 by 13-18 mm 
(exceptionally to 90 by 21 mm), sessile, short- 
acuminate, basal basiscopic segment longest, 
basal 1-2 segments free, rest of pinnule lobed 
nearly to costa; costules 3-4 mm apart; veins 8-9 
pairs; lamina-segments very firm, crenate. fertile 
ones more deeply so than sterile, on largest 
pinnules the basal segments sometimes deeply 
lobed, several pairs being separately adnate to 
costa. Sori near costules; indusium hemitelioid. 



Dec. 1963] 



Cyatheaceae (Holttum) 



109 



firm, dark, with thinner uneven edge at maturity, 
on type specimen about a quadrant of a circle, on 
some others more nearly a semicircle; paraphyses 
not longer than sporangia, some near apex of 
receptacle flat at base. Scales and hairs: upper 
surface of main rachis and pinna-rachis bearing 
spreading very narrow crisped scales to 7 mm long 
in addition to usual antrorse hairs; lower surface of 
main rachis closely warty, glabrescent; lower 
surface of pinna-rachis closely warty, pale, bearing 
more or less deciduous pale bullate-based scales, 
these more abundant distally; costae at first co- 
piously scaly, scales near base acuminate, slightly 
bullate at base, grading to small buUate scales 
distally, all scales light brown and entire; old 
costae conspicuously warty beneath; costules 
glabrescent or bearing bullate scales. 

Type specimen: Loher893, March 1897, Baguio, 
Luzon (P; dupl. at K, US). 

Distr. Malaysia: Philippines (Luzon, Min- 
danao, Biliran). 

Ecol. In forest at 640-2400 m. 

Notes. Loher's /;. 893 was not cited with the 
original description by Christ, but appears on the 
original label of the specimens at Paris, Kew and 
Washington. The degree of scaliness of lower 
surfaces of pinna-rachis and costae varies consid- 
erably, and also the abundance of warts; warts and 
scales are more abundant on the lower pinnae, and 
possibly also on plants in more exposed conditions 
of habitat. 

84. Cyathea semiamplectens Holttum, Kew Bull. 
16 (1962) 62. 

Trunk 150-200 cm tall, stout, bearing numerous 
fronds to 175 cm long. Stipe 5-15 cm, not spiny, 
covered with scales; scales 20-30 by 1-3 mm, 
narrower ones often with dark median band, wider 
ones mostly entirely pale except the apex, setae on 
the fragile edges rare. Lower pinnae gradually 
reduced, lowest 4-12 cm long, longest 40 cm. 
Largest pinnules 50-75 by 15-20 mm, sessile, 
hardly acuminate, with 6-10 pairs of separately 
adnate segments; costules 3^'/2 mm apart; veins 
to 7 pairs, not prominent on either surface; lamina- 
segments firm, sterile ones crenate, larger fertile 
ones lobed 1/2-^4 to costule. Sori near costules; 
indusium hemitelioid, nearly 1 mm wide, covering 
costular side of ripe sorus; no paraphyses seen. 
Scales and hairs: pinna-rachis rather persistently 
covered beneath with very small light brown 
bullate scales, sometimes with setiform apex, and 
very narrow entire scales; scales on costae small, 
pale brown, bullate, entire, with some broader 
flat elongate ones; costular scales mostly decid- 
uous. 

Type specimen: Womersley 11500, Eastern 
Highlands, NE. New Guinea (BRI; dupl. at L). 

Distr. Malaysia: E. New Guinea (two col- 
lections). 

Ecol. At 3300-3560 m, 'in broken subalpine 
shrubbery" (Womersley), exposed to sun; 
and 'in subalpine forest' (Brass). The latter 
collection has larger fronds, with pinnules more 
dissected. 



85. Cyathea alleniac Holttum, Kew Bull. 16 
(1962) 52. 

Trunk to 4 m, 1 5 cm 0; several smaller crowns of 
fronds sometimes produced by branches from the 
trunk. Stipe at least 30 cm, near base with many 
thick conical spines to 2 mm long; scales dark, 
shining, to 20 by 1 Vi mm, fragile edges mostly 
abraded; pneumathodes to 25 mm long, in a single 
row. Pinnae to 70 cm long. Pinnules sessile, rather 
easily detached when dry, short-acuminate, largest 
140-150 by 30 mm, more than half the segments 
separated by wide sinuses and constricted at base, 
connected by a narrow costal wing, only the 
lowest segments sometimes quite free; costules 5-6 
mm apart; veins to 1 2 pairs; lamina-segments deep- 
ly crenate, the larger ones lobed halfway to cos- 
tule, veins in such lobes pinnate. Sori near cos- 
tules; indusium a firm brown scale of rather 
irregular shape on costular side, often nearly 
circular, sometimes unevenly lobed, rarely ex- 
tending round base of receptacle on side remote 
from costule; paraphyses shorter than sporangia, 
some near apex of receptacle broad and scale- 
like at base. Scales and hairs: pinna-rachis near 
base glabrescent and warty, distal half at least 
bearing crisped hairs on lower surface; costae 
bearing some hairs as pinna-rachis on lower 
surface, also firm brown elongate scales, the 
larger ones with a dark setiform apex and some- 
times dark marginal setae; scales on costules 
bullate, brown. Frotids of branch-crowns: pinnae 
to 18 cm long, pinnules to 65 by 13 mm, lowest 
1-2 pairs of segments deeply lobed, lowest 4-6 
pairs of segments constricted at base on acroscopic 
side; veins to 9 pairs; sori and scales as fronds of 
main crown. 

Type specimen: B. E. G. Molesworth-Allen 
4127, Kuala Terla, Pahang (K; dupl. at US). 

Distr. Malaysia: Malay Peninsula. 

Ecol. On forest edge, steep ground, at 1200 m. 

86. Cyathea costulisora Domin, Acta Bot. Bohem. 
9 (1930) \Q%.—-Hemitelia montana v. A. v. R. 
Bull. Jard. Bot. Btzg III, 2 (1920) 153, non C. 
montana Sm. 1793. 

Stipe 70 cm, warty almost throughout; many 
persistent scales to 20 by 1 1/2 mm, on lower 30 cm; 
main rachis glabrescent, pale, smooth or moder- 
ately warty. Pinnae to 50 cm long. Pinnules to 
60 by 16 mm, sessile, abruptly pointed, lobed 
almost to costa, no free segments; costules 3 mm 
apart; veins 7-8 pairs; lamina-segments thin, 
subentire, sinuses '/2 mm wide. Sori near costules; 
indusium firm, brown, large, backing the costule, 
more than a semicircle; receptacle swollen; 
paraphyses not evident on old sori. Scales and 
hairs: lower surface of pinna-rachis distally bearing 
a thin covering of pale crisped hairs and some very 
small scales; scales near base of costae narrow, flat, 
dark, with some dark setae near apices, also very 
small scales; many small pale bullate scales on 
costules. 

Type specimen: Bunnemeijer 4606, Mt Merapi, 
Sumatra (BO; dupl. at L). 

Distr. Malaysia: Sumatra (one collection). 



no 



Flora Malesiana 



[ser. II, vol. P 



87. Cyathea caudata (J. Sm.) Copel. Philip. J. Sc. 
1, Suppl. II (1906) 144; Fern Fl. Philip. 2 (1960) 
222. — Alsophila caudata J. Sm. ex Hook. [J. Bot. 3 
(1841) 419. iwmen] Sp. Fil. 1 (1844) 52, t. 20B; 
Syn. Fil. (1866) 42; v. A. v. R. Handb. (1908) 37, 
785. — Hemitelia mauilensis Pr. Abh. K. Bohm. 
Ges. Wiss. V, 5 (1848) 351; v. A. v. R. Handb. 
Supplement (1917) 53. — Amphicosmia matiilensis 
Moore, Ind. Fil. (1857) 61. — Hemitelia caudata 
Mett. Fil. Lechl. 2 (1859) 30; v. A. v. R. Handb. 
Suppl. (1917) 52. — C. manilensis Domin, Pterid. 
(1929) 264.— C. dupaxensis Copel. Philip. J. Sc. 46 
(1931) 211; Fern Fl. Philip. 2 (1960) 217.— C. 
brevipes Copel. Philip. J. Sc. 81 (1952) 18; Fern Fl. 
Philip. 2 (1960) 232.— C. arborescem Copel. 
Philip. J. Sc. 84 (1955) 162. 

Stipe short, copiously warty; scales narrow, dark, 
shining, to 1 5 mm long. Lower pinnae gradually 
reduced, lowest 5-8 cm long; longest pinnae 
40-50 cm. Pinnules commonly to 85 by 16-18 mm, 
almost sessile, more or less caudate-acuminate 
(Cauda on type 25 mm); costules 3-3 '/^ mm 
apart; veins to 10 pairs; lamina-segments firm, 
rather strongly crenate-serrate. Sori near costules; 
indusium hemitelioid, in shape a quadrant of a 
circle to a semicircle, firm, reflexed against costule; 
receptacle swollen; paraphyses short, some apical 
ones broad at base. Scales and hairs: lower surface 
of pinna-rachis glabrescent, sometimes bearing 
crisped hairs towards apex; lower surface of costae 
bearing sparse flat brown thin-edged scales, some 
with a few marginal setae, also very small scales 
and sometimes a few crisped hairs; scales on 
costules flat or somewhat convex, not buUate. 

Type specimen: Cuming 267, Luzon (K; dupl. 
at P, L, A). 

Distr. Malaysia: Philippines (Luzon, Mindoro). 

Ecol. In mid-mountain forest; few records of 
altitude. 

Note. Apart from size of indusium and of 
pinnules, there seems no clear distinction between 
this species and C. heterochlamydea Copel. 

88. Cyathea borneensis Copel. Philip. J. Sc. 6 
(1911) Bot. 135; V. A. v. R. Handb. Suppl. (1917) 
33. — Alsophila latebrosa var. denudata Bedd. 
J. Bot. 31 (1893) 225; Ridl. J. Mai. Br. R. As. Soc. 
4 (1926) 9.— C. hemichlamydea Copel. Philip. J. 
Sc. 6 (1911) Bot. 361. — Hemitelia hemichlamydea 
V. A. V. R. Handb. Suppl. (1917) 47, 488.— C. ob- 
tusata Rosenst. Med. Rijksherb. n. 31 (1917) 1; 
HoLTTUM, Card. Bull. S. S. 8 (1935) 306, pi. 30; 
Rev. Fl. Mai. 2 (1954) 121. 

Trunk to 2 m or more. Stipe 5-25 cm, short- 
spiny or warty; scales to 1 5 by 1 mm, dark, shining, 
fragile edges narrow and often abraded; pneumath- 
odes 12-18 mm long, in a single row, with gaps 
between them. Lower pinnae rather irregularly 
reduced and variable in size, lowest 4-10 cm long, 
longest where stipe is longest; longest pinnae 
60 cm. Largest pinnules 80-100 by 17-22 mm, 
almost sessile, acuminate, lobed almost to costa, 
basal segment not free; costules 31/2-5 mm apart; 
veins to 10 pairs; lamina-segments thin but firm, 
almost entire, ends rounded, sinuses narrow. Sori 



near costules; indusium rather thin, on costular 
side, variable in shape and size, reaching the cos- 
tule and usually visible at maturity of sorus; 
receptacle swollen; paraphyses short. Scales and 
hairs: pinna-rachis pale to purplish, bearing some 
crisped hairs distally and sometimes throughout, 
also some residual very narrow dark spreading 
flexuous entire scales; scales on costae dark, 
entire, acuminate, flat or bullate-based, grading to 
bullate scales distally; rather dark bullate scales on 
costules. 

Type specimen: C. J. Brooks 58, Mt Penrissen, 
Sarawak (MICH; dupl. at BM). 

Distr. S. Siam northwards to Mergui, in Ma- 
laysia: Malay Peninsula, Borneo. 

Ecol. In forest, from lowland to 1100 m. 

Notes. Bornean specimens seem on the whole 
larger than those from Malaya, and are also more 
often suff"used with purple on the rachis which in 
Malayan specimens is usually green (pale when 
dry). Rosenstock wrongly cited the number of 
the type collection of his C obtusata as 1148; it 
should be 7148 {leg. King's Coll., Perak). 

89. Cyathea fenicis Copel. Philip. J. Sc. 3 (1909) 
Bot. 354; ibid. 4 (1909) 61; Fern Fl. Philip. 2 
(1960) 232.— Alsophila fenicis C. Chr. Ind. Fil. 
Suppl. (1913) 5; V. A. v. R. Handb. Suppl. (1917) 
66. — Alsophila fujiiana Nakai, Bot. Mag. Tokyo 
41 (1927) 72. 

Stipe 60 cm; spines 1 mm long; scales 15 mm 
long, narrow, dark; one pair of pinnae 6 cm long 
near base of stipe (fide Copeland). Largest pinnae 
40 cm long. Largest pinnules 80-100 by 13-21 mm, 
sessile, acuminate, lowest 1-2 pairs of segments 
constricted at base and nearly free, rest of pinnule 
lobed nearly to costa; costules 3V2-4V2 rnm 
apart; veins 10-11 pairs; lamina-segments firm, 
crenate. Sori near costules; indusium small, brown, 
on costular side. Scales and hairs: pinna-rachis 
beneath glabrescent; costal scales few, flat, rather 
broad, entire, paler distally; on costules pale entire 
scales, ovate, grading to bullate. 

Type specimen: Fenix BS 3797, Batan Islands 
(US; dupl. at P). 

Distr. Taito I. (near Formosa), in Malaysia: 
Philippines (N. Luzon: Batan Is., several collec- 
tions). 

Notes. The type was presumably in the Manila 
herbarium, destroyed during the war. Specimens 
of the same collection at US and P do not show the 
stipe; all other details given above are taken from 
them. Copeland wrongly reported this species as 
exindusiate. 

90. Cyathea junghuhniana (Kunze) Copel. Philip. 
J. Sc. 4 (1909) Bot. 5^.— Alsophila extensa [tjon 
(FoRST.) Spr.] Bl. En. PI. Jav. (1828) 246 (/?./?.?). 
— Alsophila lunulata [non (Forst.) R. Br.] Bl. I.e. 
{p.p.?}. — Alsophila junghuhniana Kunze, Bot. 
Zeit. 6 (1848) 284.— Hemitelia javanica Presl, 
Epim. Bot. (1851) 34. — Alsophila robusta De 
Vriese in Jungh. Java 1 (1852) 310, 476.— Also- 
phila debilis De Vriese, I.e. — Alsophila melanopus 
Hassk. in Hook. J. Bot. Kew Misc. 7 (1855) 325; 



Dec. 1963] 



Cyatheaceae (Holttum) 



111 



Obs. Bot. Fil. 1 (1856)42;v. A.v. R. Handb.(1908) 
40; Suppl. (1917) 65(?). — Amphicosinia javcinica 
Moore, Ind. Fil. (1857) 60. — Hemitelia jiing- 
huhniana Mett. Fil. Lechi. 2 (1859) 31; Ann. 
Mus. Bot. Lugd.-Bat. 1 (1863) 55; Racib. F1. 
Btzg 1 (1898) 38, p.p., c.xcl. var. dissoluta; v. A. 
V. R. Handb. (1908) 28; Suppl. (1917) 45, Corr. 
44; Backer & Posth. Varenli. Java (1939) 28.— 
Hemitelia latebrosci (Wall.) Mett. Ann. Mus. 
Bot. Lugd.-Bat. 1 (1863) 54, p.p. (pl.jav. tantiim); 
Racib. F1. Btzg 1 (1898) 39; v. A. v. R. Handb. 

(1908) 38, p.p.: Backer & Posth. Varenfl. Java 
(1939) 27. — C. melanopus Copel. Philip J. Sc. 4 

( 1909) Bot. 48. — Hemitelia glaitcophvlla v. A. v. R. 
Bull. Jard. Bot. Btzg II //. 7 (1912) 16; Handb. 
Suppl. (1917) 50. — Hemitelia alsophiliformis v. A. 
V. R. Bull. Jard. Bot. Btzg II, /;. 16 (1914) 15; 
Handb. Suppl. (1917) 46. — Hemitelia merapiensis 
V. A. V. R. Bull. Jard. Bot. Btzg II, /;. 16 (1914) 16; 
Handb. Suppl. (1917) 45.— Hemitelia fallax v. A. 
V. R. Bull. Jard. Bot. Btzg III, 2 (1920) 153, incl. 
var. major. — C. glaiicoplnila DoMiN, Pterid. 
(1929) 264.— C. alsophiliformis Domin, I.e. 263.— 
C. merapiensis Domin, I.e. 264. — C.falla.x Domin, 
Acta Bot. Bohem. 9 (1930) 115. 

Trunk to 2 m or more; leaf-bases persistent. 
Stipe 30-50 cm or more, base bearing spines 
1 lA — 2'/2 mm long; scales to 30 by 2 mm, dark, 
shining; pneumathodes 5-14 mm long, in a close 
double or triple row. Lower pinnae somewhat 
reduced; longest pinnae 55-70 cm long. Largest 
pinnules 80 — 1 15 by 14-21 mm, sessile, acuminate, 
lobed almost to costa, lowest segment not free; 
costules 31/2 — 41/2 mm apart; veins 10-12 pairs; 
lamina-segments firm, subentire to distinctly cre- 
nate (the latter usually when fertile). Sari near 
costules; indusium hemitelioid, variable in size and 
shape, rather thin, when largest semicircular in 
shape and distinctly visible on costular side of 
mature sorus; receptacle swollen; paraphyses short, 
slender. Scales and hairs: pinna-rachis smooth, 
glabrescent, sometimes with minute fringed scales; 
scales on costae elongate, flat, entire, of varying size; 
on costules buUate, or acuminate with buUate base. 

Type specimen: Junghuhn, Java (Herb. 
Schlechtendal, not seen; dupl. at L, K). 

Distr. Malaysia: South and Central Sumatra, 
Java. 

Ecol. In forest at 1000-2000 m, very abundant 
above Tjibodas on Mt Gedeh in West Java. 

Note. All authors from Mettenius to Backer 
& PosTHUMUS tried to distinguish both C. latebrosa 
(Wall.) Copel. and C. junghuhniana in West Java, 
but the former does not occur there. For con- 
fusion of C. junghuhniana with C. raciborskii, see 
note under latter species. 

91. Cyathea raciborskii Copel. Philip. J. Sc. 4 
(1909) Bot. 45. — Hemitelia capensis [non {L. f.) 
R. Br.] Hook. Sp. Fil. I (1844) 36, p.p.; Syn. Fil. 
(1865) 29, p.p. — Hemitelia crenulata Mett. Ann. 
Mus. Bot. Lugd.-Bat. I (1863) 55; Racib. F1. 
Btzg I (1898) 38; v. A. v. R. Handb. (1908) 27; 
Suppl. (1917) 42; Backer & Posth. Varenfl. Java 
(1939) 27, non C. crenulata Bl. 182S. —Alsophila 



crenulata Hook. Syn. Fil. (1866) 44. — Hemitelia 
junghuhniana var. dissoluta Racib. F1. Btzg (1898) 
38. — Alsophila brevifoliolata V. A. v. R. Bull. 
Jard. Bot. Btzg II, n. 20 (1915) 3; Handb. Suppl. 
(1917) 64.— C. brevifoliolata v. A. v. R. Bull. 
Jard. Bot. Btzg II, «. 28 (1918) 13. 

Trunk rarely to 2 m. Stipe 30-50 cm, warty near 
base; scales little over 10 mm by I mm, dark, 
shining; pneumathodes in a single row (in two 
rows on largest fronds), 2-5 mm long. Lowest 
pinnae varying in size, smallest seen 22 cm long; 
largest pinnae 40-50 cm. Largest pinnules 65 by 
I4(-20) mm, sessile, abruptly narrowed at apex or 
short-acuminate, lobed almost to costa, lowest 
segment not free; costules 3'/2 mm (rarely to 41/2 
mm) apart; veins to 8pairs; lamina-segments rather 
thin, usually almost entire, lobed in very wide 
pinnules. Sori near costules, usually only on lowest 
2-3 pairs of veins; indusium on costular side of 
sorus, of variable size (in Sumatran specimens 
sometimes encircling base of receptacle); paraphy- 
ses not longer than sporangia, some several cells 
wide at base. Scales and hairs: lower surface of 
pinna-rachis bearing many persistent very small 
fringed scales and sometimes sparse crisped hairs 
on distal part; on costae and costules many small 
pale bullate scales throughout, with some darker 
elongate scales near base of costae. 

Type specimen: Blume, Mt Boerangrang, W. 
Java (L; dupl. at BO). 

Distr. Malaysia: S. Sumatra, W. Java. 

Ecol. In forest at 1200-1600 m, abundant above 
Tjibodas on Mt Gedeh. 

Note. This species was confused with C. 
junghuhniana by v. A. v. R. and others, as indicated 
by their determination of specimens in herbaria. 
The two species grow side by side in the forest 
above Tjibodas, and in my experience are quite 
distinct in size, shape of pinnules, scales on costae, 
and in pneumathodes of stipe. 

92. Cyathea glaberrima Holttum, Kew Bull. 16 
(1962) 55. 

Trunk slender, to 2 m, bearing many fronds to 
190 cm long. Stipe c. 8 cm, finely warty; scales 
many, light castaneous, shining, to 15 by 1 1/2 rnrn> 
fragile edges narrow, not setiferous. Lower pinnae 
gradually reduced, lowest 3-4 cm long; longest 
pinna 50 cm. Pinnules distinctly dimorphous; 
sterile pinnules to 150 by 25 mm, lowest on stalks 
to 8 mm long, apex long-acuminate, lobed to 
3-4 mm from costa, costules 6-6 V2 mm apart, 
veins 9 pairs, lowest from costa; fertile pinnules 
to 120 by 18 mm, on stalks to 6 mm long, lobed as 
sterile, costules 5-5 1/2 rnm apart. Lower sori 
medial, distal ones nearer to costule; indusium a 
small dark brown scale on costular side; recep- 
tacle large; paraphyses shorter than sporangia. 
Scales and hairs: pinna-rachis smooth, pale, gla- 
brous except at bases of costae; at bases of costae 
on lower surface and adjacent parts of rachis a 
few rather thick crisped hairs and bullate scales; 
on costules very few bullate scales. 

Type specimen: Brass 27092, Fergusson I. 
(K; dupl. at US). 



112 



Flora Malesiana 



[ser. II, vol. P 




Fig. 18. Cyathea incisoserrata Copel. a. Pinnules attached to pinna-rachis, upper surface, x %, b. 
pinna-rachis and base of pinnule, upper surface, x 4, c. as b, lower surface, showing sori and scales, 
X 10, rf. indusium and base of receptacle, ■ 20, e. scales from costule, x 20./. scale from pinna-rachis, 
X 20, g. scale from stipe, x 6, /;. paraphyses, x 50, /. section of stipe, X *j^, j. section of pinna-rachis, 
X 6, (cult. R. B. G. Kew, origin Malay Peninsula). 



Distr. Malaysia: D'Entrecasteaux Islands: 
Fergusson and Goodenough Is. 

Ecol. In mossy oak forest, at 900-1400 m. 

Note. The shallowly lobed pinnules are unusual 
in this group of species; the position of the 
basiscopic vein, springing from the costa, 
is associated with this condition, as in sect. 
Scliizocaena. 

93. Cyathea punctulata v. A. v. R. Bull. Jard. Bot. 
Btzg II, /;. 28 (1918) U.—Alsophila punctiilara 
V. A. V. R. Bull. Jard. Bot. Btzg II, /;. 20 (1915) 
5; Handb. Suppl. (1917) 64. 

Stipe dark, warty, 40 cm or more, near base 
persistently scaly; scales to 40 by 2-3 mm, cas- 
taneous, shining, with narrow fragile edges. Pinnae 
to 70 cm long. Largest pinnules 125 by 22 mm, 
caudate-acuminate, lowest 6 pairs on larger pinnae 
distinctly stalked (stalk of lowest 3-5 mm), rest 
sessile, lobed nearly to costa; costules 4'/i rnm 



apart; veins to 12 pairs; lamina-segments very 
firm, rather strongly crenate-serrate, lowest one 
on larger pinnules reduced and almost free, 
sinuses narrow. Sori at I3 distance from costule 
to edge; indusium very small, usually entirely on 
costular side, dark near its attachment to base of 
receptacle, outer edge pale, uneven; receptacle tall, 
sporangia very numerous; paraphyses short, 
slender. Scales and hairs: pinna-rachis closely and 
finely warty on lower surface, rather pale, gla- 
brescent, residual scales small, pale, short-fringed; 
scales near base of costae broad, thin, ovate-acute, 
flat, entire or with short pale hairs, grading distally 
to buUate scales, very small irregular short-fringed 
scales also abundant; most scales on costules 
buUate, pale brown. 

Type specimen: Matthew 679, Mt Korinchi, 
Sumatra (BO; dupl. at K). 

Distr. Malaysia: Sumatra (one collection). 

Ecol. In forest, at 2400 m. 



Dec. 1963] 



Cyatheaceae (Holttum) 



113 



94. Cyathea incisoserrata Copel. Philip. J. Sc. 6 
(1911) Bot. 361; Holttum, Card. Bull. S. S. 8 
(1935) 305. — Alsophila ornata var. sikkimensis (non 
Cl. & Bak.) Bedd. J. Bot. 31 (1893) 225.— 
Alsophila ornata {non Scott) Bedd. Kew Bull. 
(1909) 423. — Alsophila incisoserrata C. Chr. Ind. 
Fil. Suppl. (1913) 5; v. A. v. R. Handb. Suppl. 
(1917) 72. — Alsophila tatebrosa var. ornata Ridl. 
J. Mai. Br. R. As. Soc. 4 (1926) 8.— Fig. 7, 8b, 18. 

Trunk to 4 m, 12 cm o including leaf-bases. 
Stipe to 85 cm, warty or with conical spines to 
1 1/2 Tini long on abaxial surface, rather persistently 
but sparsely scaly almost throughout, scales hardly 
more than 10 by 1 mm; pneumathodes in a con- 
tinuous double row. almost coalescent. Lower 
pinnae slightly reduced, longest 70 cm. Pinnules 
commonly to 100 by 25 mm, largest sometimes 
120 by 35 mm, sessile, acuminate, several pairs of 
segments near base distinctly separate, connected 
by a narrow costal wing; costules AVi-'^Vi rnm 
apart; veins commonly 12 pairs. 14-15 pairs in 
largest segments, middle ones in largest segments 
pinnately branched; lamina-segments rather thin, 
strongly crenate, lower ones of largest pinnules 
often deeply lobed, almost all separated by wide 
sinuses except on smaller pinnae. Sari near cos- 
tules; indusium hemitelioid, very small, often 
bilobed, hidden by sporangia; receptacle not 
swollen; paraphyses longer than sporangia, some 
of them 2-3 cells wide at base. Scales and hairs: 
lower surface of pinna-rachis almost smooth, 
rather pale (green when living), glabrescent or 
with some very small fringed scales; scales near 
base of costae elongate, flat, entire, or with short 
marginal hairs, brown, grading to bullate scales; 
bullate scales abundant on costules. 

Type specimen: C. J. Brooks 105, Mt Singie, 
Sarawak (MICH; dupl. at BM). 

Distr. Malaysia: Sarawak, Malay Peninsula. 

Ecol. In forest or on edge of forest, from the 
lowland to 1250 m. 

Note. This is very near C. latebrosa, agreeing 
in sori and scales on pinnules, but the size and 
shape of pinnules is characteristic. Young plants 
grown from spores at Kew agree exactly with the 
parent plant in these characters. The scales per- 
sistent throughout the stipe also show a difference 
from those of C. latebrosa. 

95. Cyathea physolepidota Alston, Nova Guinea 
n.s. 7 (1956) I. 

Stipe 25 cm, strongly spiny, spines 3-5 mm long; 
scales sparse, to 10 by \V^ mm, brown with pale 
fragile edges which have many slender hairs 
irregularly directed (not all defle.xed as originally 
described). Main rachis smooth and rather pale, 
glabrescent. Lower pinnae reduced (lowest not 
seen); longest pinna 22 cm long. Largest p/V;/7«/e5 32 
by 9 mm, sessile, short-acuminate, lobed to within 
1 mm of costa, lowest segment not free; costules 
21/2 mm apart; veins 6 pairs, usually simple; 
lamina-segments firm, edges slightly sinuous or 
crenulate. Sori near costules; indusium very small, 
hemitelioid, usually bilobed; receptacle slender, 
rather tall; paraphyses short. Scales and hairs: 



lower surface of pinna-rachis and costae bearing 
more or less abundant pale brown bullate scales, 
on bases of costae also sometimes flat scales with 
setiform apex. 

Type specimen: Carr 13871, above the Gap, 
Papua (BM; dupl. at L, SING). 

Distr. Malaysia: E. New Guinea (2 collections). 

Ecol. In forest, 2200-2500 m. 

96. Cyathea kanehirae Holttum, nom. now — 
Alsophila arfakensis Gepp in Gibbs, Arfak (1917) 
70 (non C. arfakensis Gepp, I.e.). 

Stipe not known; rachis smooth and glabrescent 
on lower surface. Pinnae to 50 cm long; pinnules 
widely spaced and jointed to rachis. Pinnules to 
80 by 20 mm. lower ones on stalks 6-7 mm long; 
basal 1-2 pairs of segments quite free and articu- 
late to costa, next 2-3 pairs constricted at base, 
rest of pinnule lobed nearly to costa; costules 6 mm 
apart; veins 7-8 pairs, mostly forked, lowest often 
with each branch again forked, not prominent on 
either surface; lamina-segments thick and rigid, 
edges slightly crenate, apices rounded. Sori near 
costules (except lowest); indusium very small, 
hemitelioid, sometimes bilobed. Scales and hairs: 
scales on costae and costules few, flat, entire. 

Type specimen: L. S. Gibbs 5990, Arfak Mts, 
W. New Guinea (BM; dupl. at K, P). 

Distr. Malaysia: W. New Guinea. 

Ecol. In mossy forest, 1600-2700 m. 

Note. The original specimens are fragmentary. 
The above description is drawn mainly from much 
better ones collected near the original locality by 
Kanehira & Hatusima (n. 13499, at BO, A). 

97. Cyathea nigropaleata Holttum, Kew Bull. 16 
(1962) 59. 

Trunk of type 1.2 m, bearing 12 mature fronds 
and 8 young ones; fronds 200 cm long. Stipe warty, 
very dark at base, paler upwards; scales near base 
20 by 2 mm. nearly black, shining, with narrow 
dull brown edges bearing scattered long dark setae. 
One pair of pinnae 6 cm long 20 cm from base of 
stipe, next pair much larger and higher; longest 
pinna 45 cm long (collector). Largest pinnules 
55-80 by 12-17 mm, lowest with stalks 2^ mm 
long, lowest 2 or more segments constricted at 
base and separated by a costal wing from the rest, 
rest of pinnule lobed to within 1 mm of costa; 
costules 3-4 '/2 rnm apart; veins 6-8 pairs; lamina- 
segments firm, rather pale, strongly crenate where 
fertile. Sori near costules; indusium a small lobed 
dark scale on costular side, hidden by sporangia; 
paraphyses a little longer than sporangia, some 2 
cells wide at base. Scales and hairs: pinna-rachis 
pale, minutely warty and glabrescent on lower 
surface, dark-hairy above; costal scales rather 
sparse, flat, ovate to elongate, the larger with 
dark centre and pale edges sometimes with 1 or 2 
setae; costular scales thin, light brown, flat, not 
fringed, setae rare. 

Type specimen: Pullen 666, Eastern High- 
lands, NE. New Guinea, (L). 

Distr. Malaysia: Eastern New Guinea (2 
collections). 



114 



Flora Malesiana 



[ser. II, vol. P 



Ecol. In Nothofagus forest, 2000 m. 

98. Cyathea microchlamys Holttum, Kew Bull. 
16 (1962) 58. 

Stipe 45 cm, base dark, paler upwards, finely 
warty; scales near base to 15 by 1 mm, dark, 
shining, with pale edges bearing dark setae. Lowest 
pinna 24 cm long, longest 27 cm. Largest pinnules 
65 by 13-15 mm, sessile, short-acuminate, lobed 
almost to costa, lowest segment not free; costules 
4 mm apart; veins to 9 pairs; lamina-segments 
thin, minutely crenate, apices obtuse, sinuses 
narrow. Sori near costules; indusium a scale '/amm 
wide on costular side of receptacle; paraphyses as 
long as sporangia. Scales and hairs: lower surface 
of pinna-rachis bearing scattered crisped hairs, 
also scattered flat narrow entire brown scales; 
costal scales sparse, flat, brown, entire; some 
scattered pale crisped hairs also on costae beneath; 
no costular scales seen (specimen is entirely fertile). 

Type specimen: Ramos BS 30475, Catanduanes, 
Luzon (US; dupl. at P). 

Distr. Malaysia: Philippines (Luzon, one 
collection). 

Note. This appears to be very near C. caudata, 
but has a long stipe and small indusia. 

99. Cyathea perpunctulata (v. A. v. R.) Domin, 
Acta Bot. Bohem. 9 (1930) \46.—Hemitelia 
perpunctulata v. A. v. R. Bull. Jard. Bot. Btzg II, 

n. 28 (1918) 25. 

Stipe unknown. Rachis near base copiously 
finely warty on lower surface, smooth and gla- 
brescent distally. Pinnae articulate to rachis and 
rather easily detached on drying, longest 47 cm 
long. Pinnules articulate to pinna-rachis, largest 
90 by 17 mm, sessile, acuminate, lobed nearly to 
costa, basal segment not free; costules 3'/2 mm 
apart; veins 10 pairs; lamina-segments rather thin, 
crenate, apices rounded, sinuses narrow. Sori 
near costules; indusium a small brown scale on 
costular side of receptacle, covered by sporangia; 
paraphyses not seen, certainly no long ones present. 
Scales and hairs: lower surface of pinna-rachis 
glabrescent, residual scales few, very narrow, dark, 
entire; scales on lower surface of costae near base 
numerous, elongate, firm, brown with pale edge, 
entire, grading to similar scales bullate at base and 
distally to many pale bullate acuminate entire 
scales; scales on costules bullate. 

Type specimen: Bunnemeijer 1219, Bt Kabung, 
Lubu Sikaping, Sumatra (BO; dupl. at L). 

Distr. Malaysia: Sumatra (2 collections). 

Ecol. In forest, 650 m. 

100. Cyathea alderwereltii Copel. Philip. J. Sc. 4 
(1909) Bot. 50. — Hemitelia sumatrana v. A. v. R. 
Bull. Dep. Agr. Ind. Neerl. n. 18 (1908) 2; Handb. 
(1908) 28; Suppl. (1917) 4S.— Hemitelia horridipes 
V. A. v. R. Bull. Jard. Bot. Btzg II, //. 16 (1914) 16; 
Handb. Suppl. (1917) Al .—Hemitelia salticola 
V. A. V. R. Bull. Jard. Bot. Btzg II, /;. 20 (1915) 18; 
Handb. Suppl. (1917) 50. — Hemitelia paraphy- 
sophora v. A. v. R. Bull. Jard. Bot. Btzg III, 2 
(1920) \5A.—Alsophila spinifera v. A. v. R. ibid. 



Ill, 5 (1922) 182.— C. horridipes Domin, Pterid. 
(1929) 264.— C. salticola Domin, l.c.—C. pa- 
raphysophora Domin, Acta Bot. Bohem. 9 (1930) 
145. — C spinifera Domin, I.e. 160. 

Stipe 60 cm or more, copiously spiny at base, 
spines 3-4 mm long; scales to 18 by 2 mm, dark, 
shining, with narrow fragile edges; pneumathodes 
in a close double row. Lower pinnae somewhat 
reduced, longest 60 cm or more long. Pinnules 
to 90 by 18 mm, sessile, rather strongly acuminate, 
lobed almost to costa, basal 1-2 segments, rarely 
to 6 pairs, constricted at base; costules 3|/2 mm 
apart; veins 10-12 pairs; lamina-segments firm, 
rather strongly crenate, sinuses narrow. Sori 
near costules; indusium a small brown scale on 
costular side of receptacle, hidden by sporangia; 
paraphyses many, longer than sporangia, some 
of them 2-3 cells wide at base. Scales and hairs: 
lower surface of pinna-rachis sparsely warty, 
almost glabrescent, at length bearing scattered 
very small fringed scales and a few narrow dark 
ones, towards apex also some pale crisped hairs; 
lower surface of costae densely scaly towards 
base, basal scales 2 mm long, narrow, acuminate, 
firm, brown, entire, grading to thinner shorter 
ovate scales and to pale bullate scales distally; 
pale bullate scales abundant on costules. 

Type specimen: Teysmann 2436, Talang Solok, 
Sumatra (BO; dupl. at L, K, US, U). 

Distr. Malaysia: Central Sumatra. 

Ecol. In forest at 1000-1500 m, very abundant 
on Mt Sago (Mt Malintang) where were collected 
the types of Hemitelia horridipes and H. para- 
physophora. The type of H. salticola only differs 
from the others in having several basal pairs of 
segments on each pinnule constricted at the base, 
the pinnules to 22 mm wide. 

lOL Cyathea amboinensis (v. A. v. R.) Merr. 

Interpr. Rumph. Herb. Amb. (1917) 63.— 
Alsophila latebrosa var. batjanensis Christ in 
Warb. Monsunia (1900) 89. — Alsophila amboinen- 
sis v. A. V. R. Philip. J. Sc. 11 (1916) Bot. 103; 
Handb. Suppl. (1917) 492. 

Stipe more than 50 cm, warty near base. Lower 
pinnae not greatly reduced, longest 50 cm long. 
Largest pinnules to 85 by 16-18 mm, sessile, 
acuminate, lobed nearly to costa, lowest segment 
not free; costules iV2-4 mm apart; veins 10 pairs; 
lamina-segments firm, rather strongly crenate. 
Sori near costules; indusium a very small dark 
scale on costular side of receptacle; paraphyses 
abundant, longer than sporangia, some of them 
2 cells or more wide at base. Scales and hairs: 
pinna-rachis smooth and glabrescent beneath; 
costae bearing many pale bullate scales almost 
to base, at base some flat ovate-acuminate scales; 
many pale bullate scales on costules. 

Type specimen: C. B. Robinson 464, Ambon 
(BO; dupl. at K, A, L, BM). 

Distr. Malaysia: Moluccas (Ambon, Ceram, 
Batjan), Central and North Celebes (?). 

Ecol. In forest at low elevations. Kjellberg 
2088, from Malili, Central Celebes, was collected 
in swamp-forest at sea-level; this and a specimen 



Dec. 1963] 



Cyatheaceae (Holttum) 



15 



from Menado collected by Posthumus are referred 
with some doubt to this species. 

102. Cyathea media Wagn. &. Greth. Un. Cal. 
Publ. Bot. 23 (1948) 44, pi. 15. 

Stipe 20-25 cm, dark, warty; scales to 20 by 
1 '/2 mm, dark to medium brown, fragile edges 
narrow and usually eroded. Lower pinnae reduced, 
lowest 8-12 cm long; longest pinnae 48 cm long. 
Pinnules to 85 by 15 mm, almost sessile, lobed 
almost to costa, 1-2 basal segments constricted 
at base and almost or quite free; costules 31/2 mm 
apart; veins 8-10 pairs; lamina-segments distinctly 
oblique, edges crenate, apices acute or rounded. 
Sori inframedial; indusium a minute scale on 
costular side of receptacle, hidden by sporangia; 
paraphyses long. Scales and hairs: near base of 
costae some elongate flat brown scales, bullate 
scales also throughout; bullate scales present on 
costules. 

Type specimen: Grether & Wagner 4162, 
Manus I., Admiralty Is (UC; dupl. at MICH). 

Distr. Malaysia: Islands to NE. of New Guinea. 

Ecol. In forest, to 1600 m. 

Note. I have included here specimens collected 
by Brass on Goodenough Island, in mossy oak 
forest at 1600 m, which are smaller than the type 
(pinnules to 50 by 11 mm) but agree in other 
respects. 

103. Cyathea latebrosa (Wall, ex Hook.) Copel. 
Philip. J. Sc. 4 (1909) Bot. 52; C. Chr. Gard. Bull. 
S. S. 7 (1934) 222; Holttum, ibid. 8 (1935) 303, 
pi. 29, p.p. max.; Rev. Fl. Mai. 2 (1954) 120.— 
Polypodium latebrosum Wall. Cat. (1828) n. 318, 
nomen. — Alsophila latebrosa Wall, ex Hook. Sp. 
Fil. I (1844) 37; Syn. Fil. (1866) 43, p.p.: Bedd. 
Handb. (1883) l\,p.p.; v. A. v. R. Handb. (1908) 
38, 789, p. p. — Dichorexia latebrosa Presl, Abh. 
K. Bohm. Ges. Wiss. V, 5 (1848) 344.— //e- 
mitelia latebrosa Mett. Fil. Hort. Lips. (1856) 
111; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 54, p.p.; 
V. A. V. R. Bull. Jard. Bot. Btzg II, /;. 23 (1916) 13, 
incl. var. paraphysata v. A. v. R.; Handb. Suppl. 
(1917) 5\,p.p., 489.— C. leiicocarpa Copel. Philip. 



J. Sc. 6 (191 1) Bot. 362.— C. longipinna Copel. 
I.e. 363. — Alsophila leucucarpa C. Chr. Ind. Fil. 
Suppl. (1913) 5; V. A. v. R. Handb. Suppl. (1917) 
66. — Alsophila longipinna C. Chr. Ind. Fil. Suppl. 
(1913) 5; v. A. v. R. Handb. Suppl. (1917) 67.— 
Alsophila lastreoides v. A. v. R. Bull. Jard. Bot. 
Btzg II, n. 23 (1916) 5; Handb. Suppl. (1917) 495. 
— Hemitelia leptolepia v. A. v. R. Bull. Jard. Bot. 
Btzg 11,//. 23 (1916) 12; Handb. Suppl. (1917)488. 
— Hemitelia rudimentaris v. A. v. R. Bull. Jard. 
Bot. Btzg III, 5 (1922) 205.— C. lastreoides Domin, 
Acta Bot. Bohem. 9 (1930) 128.— C. leptolepia 
Domin, I.e. 130. — C. rudimentaris Domin, I.e. 154. 

Trunk to c. 3 m, rather slender. Stipe 50 cm or 
more, closely short-spiny (spines l-2'/2 mm long); 
few persistent scales, near base only, to 15 by 
little over 1 mm, dark, shining, fragile edges soon 
abraded; pneumathodes in an almost continuous 
row. Lower pinnae somewhat reduced; longest 
c. 60 cm long. Largest pinnules 80-100 by 12-18 
(-20) mm, sessile, acuminate, lobed almost to 
costa, lowest segment sometimes almost free; 
costules 3-31/2 mm (rarely 4 mm) apart; veins 
c. 10 pairs; lamina-segments rather thin, more or 
less crenate, almost touching or more often sep- 
arated by distinct sinuses. Sori near costules; 
indusium a small often bilobed scale on costular 
side of receptacle, variable in size, hidden by ma- 
ture sorus; paraphyses usually longer than spo- 
rangia, some of them 2-4 cells wide at base. 
Scales and hairs: pinna-rachis sparsely warty 
beneath, glabrescent; costae not densely scaly, 
scales near base elongate, flat, brown, entire, 
grading to bullate scales; bullate scales on costules 
(most abundant on sterile pinnules). 

Type specimen: Wallich 318, Penang (K; 
dupl. at L, US, A). 

Distr. Hainan, Indo-China, and Thailand, 
southwards to Malaysia: Sumatra, Malay Pe- 
ninsula, Borneo. 

Ecol. In forest or on edge of forest from low- 
lands to 1500 m. 

Note. The South Indian plants formerly in- 
cluded here have larger thinner indusia, and should 
rank as a separate species not yet described. 



2. Section Gymnosphaera 

(Bl.) Holttum, stat. iiov. — Gymnosphaera Bl. En. PI. Jav. (1828) 242; Copel. 
Gen. Fil. (1947) 98, p.p. (e.xcl. sect. 3).—Thysanobotrya v. A. v. R. Bull. Jard. 
Bot. Btzg II, n. 28 (1918) 66, t. 10.— Alsophila sensu C. Chr. Dansk Bot. Ark. 7 
(1932) 37. — Cyathea subg. Gymnosphaera Tindale, Contr. N. S. W. Nat. Herb. 2 
(1956) 331. 
Type species: Cyathea glabra (Bl.) Copel. — Fig. 19-21. 

Distr. Madagascar; India and Ceylon, eastwards to southern China and Formosa; throughout 
Malaysia, eastwards to Fiji, and in NE. Australia. Alsophila salvinii Hook, of Guatemala has dark axes, 
reduced fertile pinnules and no indusia, but it may not be closely related to Malaysian species. 

Taxon. There is no doubt that C. glabra and its immediate allies are a natural group of species, but 
none of the characters by which they are separated from sect. Cyathea is sharply definable, and a few 
species (notably C. macgillivrayi) appear to be intermediate. To the distinguishing characters given by 
Copeland, I have added hairiness of the lower surfaces of rachises; using this as a character which may 
occur in sect. Cyathea, almost all species which are exindusiate and have hairless lower surfaces belong to 
Gymnosphaera. I have included C. macgillivrayi in the keys to both sections. 



116 Flora Malesiana [ser. II, vol. P 

Ecol. The species of this section appear all to be ferns of shady forest (some even of swamp forest), 
not of the open. Several, as here delimited, are rather widely distributed and also variable in the degree 
of division of pinnules, especially of fertile pinnules. Experimental cultivation might establish how much 
of this variation is due to edaphic and other environmental conditions, and how much is of genetic origin. 
C. biformis (Rosenst.) Copel. is peculiar in the genus by its scandent habit. 

KEY TO THE SPECIES 

1. Reduced pinnae present at base of stipe, separated from normal pinnae. 
2. Sterile pinnules of normal pinnae c. 40 by 10 mm, fertile c. 30 by 8 mm; veins 4 pairs. 

104. C. annae 

2. Sterile pinnules ofnormal pinnae 60-100 by 15-20 mm, fertile 40-100 by 11-15 mm; veins 6-10 pairs. 

3. Segments of reduced basal pinnae all with very narrow lamina, forming a wing along each side of the 

veins. 

4. Lower sori medial on veins, distal ones close to costule. Paraphyses slender at base, widening to 

scale-like apex 105, C. ramispina 

4. All sori close to costules. Paraphyses thick and dark at base, tapering to apex. 

106. C. atropurpurea 

3. Segments of reduced basal pinnae all with broad lamina 107. C. recommutata 

1. Reduced pinnae, separate from the rest, lacking; lower pinnae in some cases reduced gradually almost 

to base of stipe, the lowest sometimes with narrow segments. 
5. Fronds simply pinnate with entire pinnae (sterile) or bipinnate with few pinnae and almost entire 
sterile pinnules; in either case the true frond-apex short and abortive, evident above attachment of 
uppermost pinna. 
6. Fertile pinnules lobed almost or quite to costa 108. C. biformis 

6. Fertile pinnules only slightly lobed 109. C. scandens 

5. Fronds with normal apex. 

7. Pinnules almost entire. 

8. Sterile pinnules to 50 by 10 mm, on stalks to 1 mm long; bullate scales present on costules. 

110. C. rebeccae 

8. Sterile pinnules to 120 by 20 mm, on stalks to 4 mm long; no bullate scales on costules. 

111. C. glabra 
7. Pinnules distinctly lobed. 

9. Scales on lower surface of costules bullate. 

10. Bullate scales present on lower surface of veins of sterile pinnules. 
11. Pinnules to 100 mm long with several pairs of free segments at base . . . 112. C. hornei 

11. Pinnules to 65 mm long without free basal segments 113. C. dimorpha 

10. Bullate scales lacking on veins. 

12. Axes very dark; scales on lower surface of costae bearing many setae . . . 114. C. lurida 
12. Axes not very dark; scales on lower surface of costae sometimes with a few setae. 

13. Pinnules to 1 10 by 30 mm, cut to 2 mm from costae; costules of sterile pinnules to 6 mm apart. 

115. C. rubeUa 
13. Pinnules to 70 by 20 mm, cut to within 1 mm from costa; costulesof sterile pinnules to 4'/2 rnm 

apart 116. C. macgillivrayi 

9. Scales on lower surface of costules not bullate. 
14. Pinnules lobed almost to costa. 
15. Costal scales not setiferous 117. C. acrostichoides 

15. Costal scales bearing many setae 118. C. schlechteri 

14. Pinnules not lobed more than 2/, towards costa. 

16. Scales on costae bearing lateral setae. Segments or lobes of pinnules rounded and subentire. 
Sori usually not converging towards apices of segments. Basal basiscopic vein from costule. 

17. Stipe and basal part of rachis persistently scaly; largest pinnules with stalks at most 2 mm; 

veins to 6 pairs 119. C. subdubia 

17. Persistent large scales only at base ofstipe; largest pinnules with stalks to 4 mm; veins 3-5 pairs 

111. C. glabra 
16. Scales on costae lacking lateral setae. Segments of lamina deltoid and distinctly toothed. 
Sori converging towards apices of segments. Basal basiscopic vein often from costa. 

120. C. gigantea 

104. Cyathea annae (v. A. v. R.) Domin, Acta from the base, lowest with very narrow lamina 

Bot. Bohem. 9 (1930) 90.— Alsopliila annae v. A. on each side of veins and midribs of pinnules, 

V. R. Bull. Jard. Bot. Btzg II, n. 23 (1916) 3; upper ones with pinnules to 20 by 7 mm with cre- 

Handb. Suppl. (1917) 490.— Fig. 19c. nate edges; largest pinnae 24 cm long, sterile and 

Stipe dark, slender; scales to 60 by 1 mm, dark fertile pinnules dimorphous. Sterile pinnules to 

and shining with dull paler edges. Basal pinnae 43 by 10 mm, almost sessile, base broad, apex 

about 4 on each side of the stipe, all within 11 cm gradually narrowed, edges lobed to Ys towards 



Dec. 1963] 



Cyatheaceae (Holttum) 



117 




Fig. 19. Cyathea recommutata Copel. a. Part of pinna showing transition from fertile to sterile pinnules, 

X 2/3, b. lower surface of fertile pinnule, x 4. — C. annae (v. A. v. R.) Domin. c. Reduced pinnae at 

base of stipe, ■■' % {a-b Kunstler 7130, c cult. Hort. Bog.). 



costa; costules 3-31/2 mm apart; veins 2-3 pairs, 
simple. Fertile pinnules to 30 by 8 mm, lobed ^^ 
towards costa, veins commonly 2 pairs, all sori- 
ferous; sori without indusia. Scales and hairs: 
scales on lower surface of costae near base narrow, 
dark with pale edges not setiferous, grading to 
pale narrow scales and to dull brown bullate 
scales distally and on costules. 

Type specimen: Cult. Hort. Bog. II-K-XIII-10; 
origin Ambon, J. J. Smith (BO; dupl. at L). 

Distr. Malaysia: Moluccas (Ambon, two 
collections). 

Ecol. At 650 m. 

Note. The original description contains no 
reference to the fact that the species was described 
from a cultivated plant, the citation being 'Am- 
boina, J. J. Smith'; the type specimens in Herb. 
Bog. bear no reference to J. J. Smith, but only the 
location of the plant in the garden. J. J. Smith 
went to Ambon in 1900 (with Boerlage), and 
brought back many plants for cultivation at Bogor. 
The name annae commemorates Mrs Smith. 

105. Cyathea ramispina (Hook.) Corel. Philip. J. 
Sc. 4 (1909) Bot. 36; Sarawak Mus. J. 2 (1917) 
346, 349; C. Chr. & Holttum, Card. Bull. S.S. 



7 (1934) 200, 220.— Alsophila ramispina Hook. 
Syn. Fii. (1866) 42; v. A. v. R. Handb. (1908) 
34; Christ. Ann. Jard. Bot. Btzg 20 (1906) 138. 
— Alsophila biirbidgei {non Bak.) Christ, Ann. 
Jard. Bot. Btzg 20 (1906) 138, p.p.— Alsophila 
hallieri v. A. v. R. Bull. Jard. Bot. Btzg II, /;. 28 
(1918) 2 {non Rosenst.). — Alsophila amaiambiten- 
sis V. A. V. R. I.e. 1. — Alsophila kenepaiana 
V. A. V. R. ibid. Ill, 2 (1920) 129.— C. kenepaia- 
na Domin, Acta Bot. Bohem. 9 (1930) 127.— 
C. amaiambitensis Domin, I.e. 90. — Gymnosphaera 
ramispina Corel. Gen. Fil. (1947) 98. 

Trunk rather slender, persistently covered with 
the finely-divided basal pinnae attached to the 
persistent leaf-bases. Stipe dark, almost covered 
with small dull brown scales; larger scales very dark, 
shining, to 10 by 1 '4 rnm, with narrow thin pale 
edges. Basal pimiae several pairs, all with lamina 
reduced to a narrow wing along veins and costae, 
to c. 6 by 4 cm, with c. 6 pairs of pinnules; seg- 
ments of pinnules to 8 mm long; rest of stipe and 
rachis dark and shining, glabrescent on lower 
surface, pinna-rachises sometimes paler and 
distinctly reddish. Normal pinnae to 45 cm long. 
Pinnules slightly dimorphous (fertile smaller), 
lowest with stalks 2-3 mm long, largest 70-90 by 



118 



Flora Malesiana 



[ser. II, vol. 12 



12-19 mm, lowest 1-2 pairs of segments some- 
times almost or quite free, rest of pinnule lobed to 
about 2 mm from costa; costules 4—4y2 mm 
apart; veins to 8 pairs, usually all simple; lamina 
firm, segments slightly crenate with rounded ends. 
Sori exindusiate, distal ones close to costule. basal 
ones more distant from it; paraphyses slender at 
base, widening abruptly to a small flat apex. 
Scales and hairs: scales on costae and costules 
narrow, dark, shining with pale edges which oc- 
casionally bear a dark seta, grading to pale bullate 
scales (sometimes with setiform apex). 

Type specimen: Lobb, Sarawak (K). 

Distr. Malaysia: Borneo. 

Ecol. In rather exposed places on mountain 
ridges at 1800-2500 m; abundant on the main 
ridge of Mt Kinabalu. Also recorded at 100-170 m 
on sandstone hill-side in Tawau R. For. Res. 
Young plants do not bear the small pinnae at the 
bases of stipes. 

Note. The type specimen of Alsophila amaiam- 
bitensis v. A. v. R. has no stipe, and thus does 
not show the reduced basal pinnae, but agrees in 
other characters except that all scales seen on 
costules are narrow with dark setae, no bullate 
scales being present. 

106. Cyathea atropurpurea Copel. Philip. J. Sc. 3 
(1909) Bot. 354; ibid. 4 (1909) Bot. 36, pi. 18.— 
Alsophila atropurpurea C. Chr. Ind. Fil. Suppl. 
(1913) 4. — Gvmnosphaera atropurpurea Copel. 
Gen. Fil. (1947) 98; Fern Fl. Philip. 2 (1960) 234. 

Differs from C. ratnispina in somewhat smaller 
size of pinnae (largest seen 30 cm long) and pin- 
nules (to 80 by 17 mm), veins to 6 pairs, segments 
of lamina more strongly crenate, sori all close to 
the costa, paraphyses thick and dark at the base, 
tapering and paler distally. 

Type specimen: Merrill 6056, Mt Halcon, 
Mindoro (MICH; dupl. at A). 

Distr. 7V/a/av.s/a: Philippines (Luzon, Mindoro, 
Leyte, Mindanao). 

Ecol. At altitudes of 1000 m and o\er. 

107. Cyathea recommutata Copel. Philip. J. Sc. 
4 (1909) Bot. 36; C. Chr. Card. Bull. S.S. 7 (1934) 
220; Holttum, Rev. FI. Mai. 2 (1954) 125.— 
Gymnosphaera squamulata {nan Bl.) J. Sm. ex 
Hook. Gen. Fil. (1842) t. 100.— Alsophila squa- 
mulata Hook. Sp. Fil. I (1844) 51, p.p.; Bedd. 
Ferns Br. Ind. (1867) t. 235.— Alsophila com- 
mutata Mett. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 
53; Bedd. Handb. (1883) 14; v. A. v. R. Handb. 
(1908) 34 iuoii C. commutata Spr.). — C. hewittii 
Copel. Philip. J. Sc. 6 (1911) Bot. 134, t. 14.— 
Alsophila heteromorpha v. A. v. R. Bull. Jard. 
Bot. Btzg II, «. 16 (1914) 1; Handb. Suppl. (1917) 
56; Bull. Jard. Bot. Btzg III, 2 (1920) 129, incl. 
var. decomposita v. A. v. R. — Alsophila hewittii 
V. A. V. R. Handb. Suppl. (1917) 55; C. Chr. 
Gard. Bull. S.S. 7 (1934) 221.— C. toppingii 
Copel. Philip. J. Sc. 12 (1917) Bot. 51; C. Chr. 
Gard. Bull. S.S. 7 (1934) 110.— Alsophila subulata 
V. A. V. R. Bull. Jard. Bot. Btzg II, n. 28 (1918) 1. 
— C heteromorpha Domin, Pterid. (1929) 262. — 



C. subulata Domin, Acta Bot. Bohem. 9 (1930) 
164. — Gymnosphaera recommutata Copel. Gen. 
Fil. (1947) 98. — Gymnosphaera hewittii Copel. I.e. 
—Fig. 19a, b, 21d. 

Trunk rather slender, commonly not over 3 m. 
Stipes very dark; basal scales dark, shining, with 
thin fragile margins, to c. 20 by 2 mm; small 
pale dull scales also abundant. Reduced pinnae, 
several (rarely to 9) pairs attached to lower part 
of stipe, largest to 10 cm long, on old fronds often 
reduced to their stout spine-like bases, their 
pinnules simple and entire or the largest slightly 
lobed; largest pinnae 40 cm. Pinnules dimorphous 
(sometimes with intermediate conditions); sterile 
pinnules commonly 6O-70(-90) by 16 mm, 
lowest on stalks 2-3 mm long, one basal segment 
sometimes almost free, rest lobed about halfway 
to costa, apex acuminate; costules 4-^'/2 mm 
apart; veins to c. 7 pairs, simple; lamina-segments 
rigid, dark on upper surface, apices rounded and 
slightly crenate; fertile pinnules 6-12 mm wide, 
costules 3^ mm apart, sori close to costules, no 
indusia; paraphyses dark, not longer than sporan- 
gia. Scales and hairs: pinna-rachis smooth and gla- 
brescent on lower surface; scales near bases of 
costae narrow, dark and shining with pale edges, 
not setiferous, grading to brown bullate scales 
distally and on costules. 

Type specimen: Cuming 396, Mt Ophir, Malay 
Peninsula (original lost; dupl. at K, BM, FI, GH). 

Distr. Malaysia: Sumatra (central and south), 
Malay Peninsula, Borneo. 

Ecol. Commonly at 600-1500 m, in acid peaty 
or sandy soil, in forest (not in exposed places); 
in Borneo also at 0-60 m, in swamp forest on sandy 
ground. 

Note. The type collection of C. hewittii 
(Brooks & Hewitt 21, Bongo Mt), both in Herb. 
Copel. (MICH) and at BM, is certainly referable 
to the present species, but another collection of 
Brooks so named at Kew (s.n., Jan. 1908) is a 
mixture of C. recommutata and C. ramispina. 

108. Cyathea biformis (Rosenst. ) Copel. Philip. 
J. Sc. 6 (1911) Bot. 364; ibid. 11 (1947) 117.— 
Stenochlaena dubia v. A. v. R. Bull. Dep. Agr. 
Ind. Neerl. n. 18 (1908) 26; Handb. (1908) 721; 
Holttum, Gard. Bull. S.S. 5 (1932) 250, non C. 
dubia (Bedd.) Domin, 1929. — Alsophila biformis 
Rosenst. in Fedde, Rep. 9 (1911) 423; v. A. v. 
R. Handb. Suppl. (1917) 71; Brause, Hedwigia6I 
(1920) 401. — Polybotrya arfakensis Gepp in Gibbs, 
Arfak (1917) 71. — Thvsanobotrva arfakensis v. 
A. V. R. Bull. Jard. Bot. Btzg 11, n. 28 (1918) 
66, t. 10.— C. gibbsiae Copel. Philip. J. Sc. 38 
(1929) 129. — Gvmnosphaera biformis Copel. Gen. 
Fil. (1947) 99.— Fig. 20. 

Stem 1-1 '/2 cm 0, climbing (clinging to support- 
ing tree by its roots), apex and bases of stipes 
covered with shining very dark scales to 20 by 
less than 1 mm wide, long-acuminate, edges pale. 
Stipe nearly black, slightly rough and scaly near 
base only, rest smooth and shining. Fronds of two 
kinds, simply pinnate (always sterile) and bi- 
pinnate, the true frond-apex always short and abor- 



Dec. 1963] 



Cyatheaceae (Holttum) 



119 



x^-- 






s^^^^i^^mn 



a 





Gz 



Fig. 20. Cvathea biformis (Rosenst.) Copel. a. Fertile pinnules, X 2/3, b. sterile pinnules, x 2/3, c. apical 
part of frond, showing abortion of true apex, x 2/3^ d. part of c, x 8 {q-b Brass 6806, c-f/ Brass 8947). 



tive, evident above the attachment of the upper- 
most pinna, the true pinna-apex of bipinnate 
fronds similarly abortive. Simply pinnate fronds 
to 45 cm long (excluding stipe) with up to 17 pairs 
of pinnae and a false apical one; pinnae to c. 
60 by 18 mm, edges deeply crenate, apex shortly 
caudate-acuminate,baseslightly unequal, narrower 
and rounded acroscopically, broadly cuneate basi- 
scopically, stalks 2 mm long, lamina firm, veins in 
pinnate groups, none separately from costa. 
Bipinnate fronds: stipe to 21 cm, sometimes bearing 
1-2 small pinnae near the base, pneumathodes 
commonly 3-7 mm long, rather widely spaced; 
lamina to at least 100 cm long with pinnae 10-13 
cm apart on each side of the rachis; pinnae to 
28 cm long, lowest pinnules reduced, middle 
sterile pinnules stalked (2 mm), 60-90 by 15-18 
mm, shaped as pinnae of simply pinnate fronds, 
occasionally (at high elevations) lobed V^-Yi 
towards costa; fertile pinnules stalked (3-5 mm), 
c. 50 by 4-10 mm, lobed almost or quite to the 
costa, costules 3 Vi mm apart, lobes 1 V2-2 mm wide, 
entire with rounded tips, basiscopic edge of 



each lobe decurrent to join the lobe below it; 
sori to 4 pairs on each lobe, no indusia. Scales on 
frond: lower surfaces usually glabrous, in one case 
a few dark buUate scales seen on sterile leaflets. 

Type specimen: Copland King 57, Papua 
(S-PA; dupl. at MICH, BO). 

Distr. Malaysia: Moluccas (Ambon), New 
Guinea. 

Ecol. Climbing to 2-3 m or more, on trees in 
forest, at 300-2200 m, reported by Brass to be 
common in mossy forest, rainforest and Agathis 
forest; Pullen reported 'very common ground 
fern which often ascends trees to 8 feet', at 850 m, 
in Notlwfagus forest. 

Note. The only specimen with rather deeply 
lobed sterile pinnules is Gibbs 5984, from 2200 m 
(type of Polybotrya arfakensis Gepp). 

109. Cyathea scandens (Brause) Domin, Acta 
Bot. Bohem. 9 (1930) 156. —Alsophila scandens 
Brause, Bot. Jahrb. 56 (1920) 77. 

Habit of C. biformis, agreeing in abortive apices 
of frond and pinnae, differing: stipe 17 cm, 



120 



Flora Malesiana 



[ser. II, vol. P 



lowest pinnae 4 cm long, largest pinna 22 cm long, 
fertile pinnules c. 5 mm wide, shallowly lobed; 
elongate dark shining scales with narrow pale 
edges on costae, small dark buUate scales on veins 
of sterile pinnules. 

Type specimen: Ledermann 9885, E. New 
Guinea, Sepik Region (B). 

Distr. Malaysia: Eastern New Guinea (one 
collection). 

Ecol. Climbing fern, at 1000 m. 

Note. The type collection consists of two in- 
complete fronds, on two sheets. It is possible that 
this species should be united with C. biformis, 
but the shape of the fertile pinnules appears dis- 
tinctive; the type collection bears no wholly sterile 
pinnules (some are sterile at the base, contracted 
and fertile towards apex). 



110. Cyathea rebeccae (P. v. M.) Domin, Pterid. 
(1929) 263; Acta Bot. Bohem. 9 (1930) 153; 
TiNDALE, Contr. N. S. W. Nat. Herb. 2 (1956) 334. 
— Alsop/iila rebeccae F. v. M. Fragm. 5, xxxiii 
(1865) 53, xxxvi (1886) 117. 

Stipe very dark at base, finely warty after fall 
of scales; scales to 15 by 1 1/2 mm, median band 
shining black to brown, edges thin and dull; 
upper part of stipe smooth purplish; pneumathodes 
4-5 mm long, rather widely spaced. Lower pinnae 
gradually reduced, the lowest 10 cm or less long; 
several lower pairs soon caducous, leaving an 
apparently long stipe; largest pinnae 30 cm long 
(45 cm in Queensland). Sterile pinnules to 50 by 
10 mm, on stalks to 1 mm long, edges entire or 
slightly unevenly crenate in basal 2/3, broadly 
blunt-serrate in distal y^; base rounded on acro- 
scopic side, broadly cuneate on basiscopic side; 
lamina firm; each main lateral vein (which would 
be a costule in a lobed pinnule) forked alternately 
3 or 4 times, the first fork at or very close to the 
costa. Fertile pinnules slightly smaller than sterile; 
sori 2-4 to a vein-group; no indusia. Scales and 
hairs: near base of costae on lower surface dark 
flat elongate scales with pale edges; distally on 
costae and on veins of sterile pinnules a few small 
dark buUate scales; no hairs on upper surface of 
costae. 

Type specimen: Dallachy, Rockingham Bay, 
Queensland (MEL; dupl. at K). 

Distr. NE. Queensland, in Malaysia: Lesser 
Sunda Is (Flores, 3 collections). 

Ecol. In Flores found at 1300-1700 m in forest; 
a tree fern with short trunk. In Queensland re- 
ported from sea level to 1400 m, locally abundant, 
a small tree-fern with trunk up to c. 3 m high, 
fronds to 21/2 m long. 

111. Cyathea glabra (Bl.) Copel. Philip. J. Sc. 4 
(1909) Bot. 35; Holttum, Card. Bull. S.S. 8 
(1935) 316; Rev. Fl. Mai. 2 (1954) 127.— Crw- 
nosphaera glabra Bl. En. PI. Jav. (1828) 242; 
Copel. Gen. Fil. (1947) 98.—Alsophila glabra 
Hook. Sp. Fil. 1 (1844) 51; Mett. Ann. Mus. 
Bot. Lugd.-Bat. 1 (1863) 52, p.p. (?); Bedd. 
Handb. (1883) 14, p.p.; Raciborski, Fl. Btzg I 



(1898) 34, p.p. — Alsophila vexans Ces. Atti Ac. 
Napol. 7, n. 8 (1876) 4.— Alsophila diibia Bedd. 
J. Bot. 25 (1883) 1, t. 279a; Handb. Suppl. (1892) 
4; V. A. V. R. Handb. (1908) 31; Suppl. (1917)489. 
— Alsophila reducta v. A. v. R. Bull. Jard. Bot. 
Btzg II, n. 28 (1918) I.— C. reducta Domin, 
Acta Bot. Bohem. 9 (1930) 153.— C. vexans C. 
Chr. Card. Bull. S.S. 7 (1934) m.—Gymno- 
sphaera vexans Copel. Gen. Fil. (1947) 98. 

Trunk rather slender. Stipe very dark, base 
rough after scales have fallen; scales dark, shining, 
with fragile pale edges; rachis dark to purplish, 
smooth and glabrescent on lower surface. Lowest 
pinnae sometimes much reduced, especially on 
young lowland plants; largest pinnae commonly 
45 cm long, to 55 cm. Largest pinnules 90-120 by 
15-20 mm, lowest ones on stalks 1-A mm long, 
base broadly rounded, apex short-acuminate, 
edges crenate (one crenature to each vein-group) 
to slightly lobed, in some cases lobed half-way to 
the costa; costules 4-5 mm apart; veins 3-5 pairs, 
usually all simple. Sori 1-3 (rarely 4) pairs, on 
each vein-group; no indusia; paraphyses slender, 
shorter than sporangia. Scales and hairs: scales on 
costae few, narrow, dark with pale edges which 
often bear a few dark setae, on costules similar 
but smaller, no bullate scales. 

Type specimen: van Hasselt, Mt Karang, 
W. Java (L; fragment at K). 

Distr. Malaysia: Sumatra, Malay Peninsula, 
Borneo, W. Java. 

Ecol. In lowland swamp forest and in mountain 
forest to 1500 m. 

Note. This species is very near C. gigantea, 
C. subdubia and C. podophylla (Hook.) Copel. 
In Hooker's herbarium is a small fragment of 
true C. glabra from Blume, with a good specimen 
of C. gigantea, also from Java, on the same sheet. 
Probably because of this, Beddome and others 
gave the name C. glabra to ferns from India and 
Ceylon which are C gigantea, and when Beddome 
received the true C glabra from the Malay 
Peninsula he re-named it C. dubia. C. podophylla, 
from Indo-China and S. China, has pinnules 
always almost entire and almost sessile, the basal 
basiscopic vein usually from the costa, veins of 
adjacent groups often slightly anastomosing, 
and in well-grown specimens several pairs of sori 
to each vein-group. Hooker published an ex- 
cellent figure of C podophylla (2nd Cent. Ferns, 
t. 66, 1861). 

112. Cyathea hornei (Bak.) Copel. Bull. Bern. 
P. Bish. Mus. 59 (1929) 38; Philip. J. Sc. 77 (1947) 
\\9.— Alsophila hornei Bak. J. Bot. 17 (1879) 293. 
—Alsophila dissitifolia Bak. ibid. 24 (1886) 182.— 
Alsophila brunnea Brause, Bot. Jahrb. 56 (1920) 
73. — Alsophila ledermannii Brause, I.e. 76. — 
Alsophila olivacea Brause, I.e. lA. — Alsophila 
melanocaulos v. A. v. R. Nova Guinea 14 (1924) 1. 
— C. dissitifolia Domin, Pterid. (1929) 262.— 
C. /)/-«//;?£'« Domin, Acta Bot. Bohem. 9 (1930) 101; 
Copel. Philip. J. Sc. 77 (1947) 117, 119.— C. di- 
morphophylla Domin, Acta Bot. Bohem. 9 (1930) 
III (new name for Alsophila ledermannii); Copel. 



Dec. 1963 ] 



Cyatheaceae (Holttum) 



121 



Philip. J. Sc. 77 (1947) 117.— C. olivacea Domin, 
Acta Bot. Bohem. 9 (1930) 143; Copel. Philip. J. 
Sc. 77 (1947) 118, 119.— C. nwlanuclada Domin, 
Acta Bot. Bohem. 9 (1930) 174 (new name for 
Alsophila melanocaulos); Copel. Philip. J. Sc. 77 
(1947) 118. — Gymnospliaera hornei Copel. Gen. 
Fil. (1947) 99. — Gymnosphaera melanoclada Co- 
pel. I.e. 

Trunk rather slender (4 cm o when dry), to 
3 or 4 m tall; leaf-scars to 2 cm o. Stipe to 25 cm 
but often much shorter, very dark, the basal part 
covered with dark shining pale-edged scales to 
15 by 2 mm; pneumathodes 7-13 mm long. Lower 
pinnae gradually reduced, lowest commonly 10 cm 
long, sometimes with lamina reduced to a narrow 
wing along veins and costa; largest pinnae 40-50 
cm long (60 cm reported of Alsophila olivacea), 
sterile and fertile pinnules strongly dimorphous. 
Largest sterile pinnules to 100 by 25-30 mm wide 
near the base, the lower ones with stalks to 3 mm, 
basal 2-^ pairs of segments quite free, then several 
pairs separately adnate to costa; costules 5 mm 
apart; veins to 10 pairs, mostly forked, middle ones 
in free segments twice forked; lamina-segments 
strongly crenate or the larger free ones somewhat 
lobed. Fertile pinnules to 50-60 by 11-17 mm, 
with basal free segments as sterile; costules 3'/^-4 
mm apart; veins usually fewer than in sterile 
pinnules, often forked; segments crenate, usually 
separated by rather wide sinuses; 50/7 almost 
covering lower surface of fertile segments, ex- 
indusiate. Scales and hairs: pinna-rachis dark, 
glabrescent or bearing narrow dark pale-edged 
scales; scales near bases of costae elongate, dark, 
shining with pale edges, grading to light brown 
bullate scales distally and on costules; similar 
bullate scales also abundant on veins of sterile 
pinnules. 

Type specimen: J. Horne 620, Fiji (K). 

Distr. Fiji, Louisiade Arch., in Malaysia: 
Eastern New Guinea. 

Ecol. On the mainland of New Guinea at 
850-2000 m, in forest; on the Louisiade islands at 
c. 700 m in stunted or mossy forest of ridge-crests. 
Small plants have pinnules almost sessile, even on 
the largest pinnae. 

Note. This species is here broadly interpreted, 
but I cannot see any clear differences in characters 
of the larger pinnae which would warrant its sub- 
division. There are differences in the small basal 
pinnae, which in some cases have very narrow 
ultimate divisions, in others broad divisions, 
comparable with the difference between C. 
raniispina and C. reconunutata in Borneo. Many 
specimens however lack these basal pinnae, and 
it is at present impossible to estimate whether 
the differences in basal pinnae are in any way 
correlated with differences in the larger pinnae. 

113. Cyathea dimorpha (Christ) Copel. Philip. 
J. Sc. 4 (1909) Bot. U.~Alsophila dimorpha 
Christ, Ann. Jard. Bot. Btzg 19 (1904) 41; 
V. A. V. R. Handb. (1908) 36. 

Stipe short. Lower pinnae gradually reduced, 
lowest less than 9 cm long; largest pinnae seen 



30 cm long; fertile and sterile pinnules strongly 
dimorphous. Sterile pinnules to 65 by 20 mm, on 
stalks to 3 mm, near base lobed % towards costa, 
for the most part lobed only % towards costa; 
costules 31/2 mm apart; veins to 6 pairs, simple; 
lamina-segments very firm, almost truncate, 
slightly crenate. Fertile pinnules to c. 35 by 7 mm, 
with stalks to 2 mm, lobed to within 1 mm of 
costa; costules 2 1/2 mm apart; veins to 4 pairs; 
sori without indusia, paraphyses short. Scales and 
hairs: on lower surface of costae narrow shining 
dark scales with pale edges, not setiferous, grading 
to brown bullate scales on costules and veins of 
sterile pinnules. 

Type specimen : Sarasin 2031, Bohaa Mts Cele- 
bes (BAS; dupl. at P). 

Distr. Malaysia: Central and SE. Celebes (2 
collections). 

Ecol. At 125-645 m in SE. Celebes, at 1500- 
1700 m in Central Celebes. 

114. Cyathea lurida (Bl.) Copel. Philip. J. Sc. 4 
(1901) Bot. 45.— Chnoophora lurida Bl. En. PI. 
Jav. (1828) 244.— Alsophila lurida Hook. Sp. Fil. 
1 (1844) 55; v. A. v. R. Handb. (1908) 44; Suppl. 
(1917) 70. — Alsophila kingii Clarke in Bedd. 
Handb. (1883) 475; v. A. v. R. Handb. (1908) 
36. — Alsophila bakeri Zeiller, Bull. Soc. Bot. Fr. 
32 (1885) 72. — Alsophila melanorachis Copel. 
Philip. J. Sc. 2 (1907) Bot. 146; v. A. v. R. Handb. 
(1908) 791.— C. melanorachis Copel. Philip. J. Sc. 
4 (1909) Bot. 38.— C. kingii Copel. I.e. 56; 
Holttum, Card. Bull. S.S. 8 (1935) 315, pi. 36; 
Rev. Fl. Mai. 2 (1954) 126.— C. subdimorpha 
Copel. Philip. J. Sc. 8 (1913) Bot. 140, pi. 2.— 
Alsophila heteroplnlla v. A. v. R. Bull. Jard. Bot. 
Btzg II, n. 16 (1914) 2; Handb. Suppl. (1917) 60. 
— Alsophila subdimorpha v. A. v. R. Bull. Jard. 
Bot. Btzg II, n. 16 (1914) 2.— C. heterophylla 
Domin, Pterid. (1929) 262. — Gymnosphaera me- 
lanorachis Copel. Gen. Fil. (1947) 98; Fern Fl. 
Philip. 2 (1960) 234.— Gymnosphaera kingii 
Copel. Gen. Fil. (1947) 99.— Fig. 21a-c. 

Trunk short. Stipe long (no reduced basal 
pinnae), very dark, rough near base after fall of 
scales; scales to 10 by 1 '/2 mrri' dark with pale 
edges; pneumathodes rather widely spaced, 6-9 
mm long. Pinnae commonly to 50 cm long, 
rarely to 75 cm; pinnules strongly dimorphous, 
with occasional intermediate conditions. Largest 
sterile pinnules 75-110 by 16-25 mm, on largest 
fronds with several free basal segments, on all 
fronds lobed almost to the costa; stalks of lower 
pinnules 3-5 mm; costules 3i/2-^V2 mm apart; 
veins to 10 pairs, mostly forked; lamina-segments 
firm, in the larger pinnules strongly crenate. 
Fertile pinnules 60-90 by 6-12(-17) mm wide, 
on stalks to 3 mm, the largest with free segments at 
base; costules commonly 3 mm apart, on largest 
fronds occasionally to 6 mm, in the latter case the 
segments separated by wide sinuses; sori ex- 
indusiate, almost covering lower surface of 
segments; receptacle much elongate; paraphyses 
shorter than sporangia. Scales and hairs: near 
bases of costae on lower surface narrow dark 



122 



Flora Malesiana 



[ser. II, vol. 12 




Fig. 21. Cyathea luhda (Bl.) Copel. a. Base of a fertile pinna, x 2/3, b. base of unusually large sterile 

pinna, x 2/3, c. normal form of sterile pinna, x 2/3. — c. recommutata Copel. d. Base of frond, showing 

reduced pinnae, x y^ {a-c Eryl Smith 833, d after Holttum, Flora of Malaya 2, fig. 50). 



scales with pale edges sometimes bearing dark setae, 
grading to bullate scales on costules of sterile 
pinnules. 

Type specimen: Java, herb. Blume (L). 

Distr. Malaysia: Sumatra, Malay Peninsula, 
W. Java, Philippines (Mindoro). 

Ecol. In ridge forest (not in exposed places) 
at 1250-1800 m. In the Malay Peninsula especially 
abundant on a quartzite ridge at Eraser's Hill, but 
absent from forest on the neighbouring granite. 
On granite, it occurs only on the crests of steep 
ridges, often in moss-forest. 

Note. The type of Alsophila melanorachis 
Copel. from Mindoro agrees in all essential char- 
acters with C. liirida in Malaya. No specimens of 
C. lurida have been found in Borneo, so that there 
is a considerable gap in distribution. 

115. Cyathea rubella Holttum, Kew Bull. 16 
(1962) 61. 

Trunk to 5 m; fronds many, to 200 cm long. 



Stipe 12-20 cm, the base dark purple-brown 
when dry, paler upwards, throughout bearing 
scattered slender spines 2 mm long; scales abun- 
dant near base and throughout on each side of the 
band of hairs on upper surface, to 25 by hardly 
2 mm, dark with narrow paler edges bearing many 
irregular setae; small irregular dull pale scales 
also scattered over surface of stipe. Lower pinnae 
gradually reduced, lowest 6-7 cm long; longest 
pinnae 45 cm long. Pinnules to 110 by 30 mm 
(more commonly narrower), the lowest stalked 
2 mm, lobed throughout to 1-2 mm from costa 
(lowest 1-2 segments free only on largest pinnules), 
apex caudate-acuminate; costules 4'/2-6 mm apart 
(a little more widely spaced in sterile than in fertile 
pinnules); veins 8-10(-12) pairs, mostly forked, 
the basal basiscopic vein attached well above base 
of costule; lamina-segments thin, somewhat 
tapering so that sinuses are triangular, edges 
crenate-serrate towards apex. Sori medial, at forks 
of veins or on one or both branches above the 



Dec. 1963] 



Cyatheaceae (Holttum) 



123 



fork in the case of basal veins; no indusia; para- 
physes slender, shorter than sporangia. Scales 
and hairs: pinna-rachis reddish, paler dislally, 
smooth apart from a few small spines, with 
scattered very small pale fringed scales; costae 
pale or somewhat suffused with red, sometimes 
with very small scales as pinna-rachis, larger scales 
sparse, flat, rather pale, apex setiform and some- 
times one or more setae on the thin edges; costules 
bearing rather large pale bullate scales; no hairs 
on upper surface of costules. 

Type specimen: Hoogland 4487, Papua, 
Northern District, Tufi Subdistrict (K; dupl. at 
BM, L, A). 

Distr. Malaysia: East New Guinea and d' 
Entrecasteaux Is. 

Ecol. At 650-900 m, 'common in fairly dense 
fairly low forest' (type); in rain-mossy forest 
transition in Normanby I. and in oak-rain forest 
transition in Goodenough 1. 

116. Cyathea macgillivrayi (Bak.) Domin. Pterid. 
(1929) 263; Copel. Philip. J. Sc. 77 (1947) 109.— 
Alsophila macgillivrayi Bak. Syn. Fil. ed. 2 
(1874) 458.— C. gracillima Copel. Un. Cal. Publ. 
Bot. 18 (1942) 219; Philip. J. Sc. 77 (1947) 118, 
pi. 11. — Gvmnosphaera gracillima Copel. Gen. 
Fil. (1947)99. 

Trunk slender, to 4 m, often with bulbils growing 
from it or extra trunks at the base, bearing rather 
few^ fronds 100-150 cm long. Stipe usually 20-30 
cm, in some cases only 10 cm, dark brown near 
base only, closely warty; scales to 15 by 1 mm, 
brown with narrow fragile edges. Lower pinnae 
gradually reduced, lowest 5 cm long when stipe 
is very short; longest pinnae to 40 cm. Pinnules 
commonly to 60 mm long, sometimes to 100 mm, 
fertile and sterile somewhat dimorphous (dif- 
ference between them variable); sterile pinnules 
to 18 mm wide, costules 4-5 mm apart, veins 10 
pairs, lamina-segments crenate with rounded 
apices, sinuses narrow; fertile pinnules 12-14 mm 
wide, costules about as in sterile pinnules but 
lamina-segments narrower so that sinuses are 
rather wide. Sori — medial (at or sometimes above 
forks of veins, rarely on both branches of a vein), 
the distal ones sometimes nearer the costules; 
no indusium but sometimes 1 or 2 narrow scales 
on costular side at base of sorus; paraphyses about 
as long as sporangia. Scales and hairs: pinna- 
rachis pale, smooth, glabrescent, residual scales 
pale, largest with an occasional dark seta, smallest 
bearing long slender hairs, sometimes separate 
slender crisped hairs also present; on costae 
pale flat scales, elongate near base of costa, ovate 
and somewhat convex or bullate distally; on 
costules pale ovate convex or bullate. 

Type specimen: Macgillivray, Louisiades (K). 

Distr. Malaysia: New Guinea (except extreme 
west), Louisiade Archipelago. 

Ecol. In forest, on the mainland on mountain 
ridges to 2000 m but apparently most abundant 
at lower elevations, on the islands mostly below 
300 m, also on ridges, locally abundant. 

Note. This species is on the border-line between 



sect. Cyathea and sect. Gymnosphaera. It appears 
to be near C. gregaria, which differs in position of 
sori and in having dark crisped hairs and dark 
scales on costae. 

117. Cyathea acrostichoides (v. A. v. R.) Do.min, 
Acta Bot. Bohem. 9 (1930) SS.— Alsophila acrosti- 
choides V. A. V. R. Bull. Jard. Bot. Btzg II, n. 28 

(1918) 2. 

Stipe slender, copiously spiny throughout, 
spines 1-2 mm; rachis similarly spiny near base, 
slightly rough near apex; no scales seen on stipe 
and rachis. Frond (excluding stipe) 90 cm long; 
lowest pinnae somewhat reduced and more wide- 
ly spaced; longest pinnae 35 cm. Pinnules somewhat 
dimorphous; largest 50-75 mm long, sterile 14—18 
mm wide, fertile 9-15 mm wide, almost sessile, 
lowest 1-2 segments free or nearly so (except on 
smaller fronds), rest lobed almost to costa, apex 
shortly acuminate; costules 3 ',2^1 2 rnm apart; 
veins to 8 or 9 pairs, lower ones forked; lamina- 
segments crenate. or the lowest distinctly lobed, 
sterile separated by sinuses to 1 mm wide, fertile 
by sinuses to 2 mm wide (as wide as segments). 
Sori covering whole surface of segments of fertile 
pinnules; receptacle much raised and irregularly 
enlarged above the base; no indusium; paraphyses 
slender, shorter than sporangia. Scales and hairs: 
scales on costae flat, mostly ovate, medium brown, 
with some short marginal hairs; on costules smaller 
similar scales, more or less convex. 

Type specimen: Kornassi 541, Kaniki, Ceram 
(BO: dupl. at L). 

Distr. Malaysia: Moluccas (Ceram). \V. New- 
Guinea (Japen I.). 

Ecol. On Japen I. found at 650-1100 m, in 
forest, abundant, trunk 1-3 m tall; "one com- 
munity above a landslip which has opened the 
forest" (Cheesman). 

Note. Four collections from Japen Island agree 
well in size and all other characters with the type 
from Ceram. One of them includes a small frond, 
fully fertile, with largest pinna 12 cm and pinnules 
22 mm long. Another collection (Cheesman 1431, 
BM) consists of a much larger frond, with pinnae 
to 75 cm long, pinnules to 135 by 32 mm (all 
sterile), costules 5' 2 mm apart, veins to 14 pairs; 
it agrees with other specimens in shape of pinnules, 
in scales, and in spiny stipe. 

118. Cyathea schlechteri (Brause) Domin, Pterid. 

(1929) 263; Copel. Philip. J. Sc. 77 (1947) 117, 
\\9.— Alsophila schlechteri Brause, Bot. Jahrb. 49 
(1912) 15, f. ID; V. A. v. R. Handb. Suppl. (1917) 
61.— Gvmnosphaera schlechteri Copel. Gen. Fil. 
(1947) '99. 

Stipe not known. Pinnae to 48 cm long. Sterile 
and fertile parts of pinnules strongly dimorphous 
(no completely sterile pinnules seen); largest 
pinnules 85 mm long (fertile in basal part, sterile 
distally), largest completely fertile pinnules 60 mm 
long; sterile part of pinnule 16 mm wide, lobed 
nearly to costa at the base, costules 4-412 rnm 
apart, veins 10 pairs, mostly forked; fertile part 
to 1 1 mm wide, fertile segments little over 2 mm 



124 



Flora Malesiana 



[ser. II, vol. 12 



wide. Sori exindusiate, receptacle much raised. 
Scales and hairs: pinna-rachis smooth and gla- 
brescent on lower surface, paler than main rachis; 
scales on costae sparse, the larger flat, ovate to 
elongate, pale with many long flexuous dark setae, 
also very small irregular pale scales; on costules 
setiferous scales as on costae but smaller, none 
bullate. 

Type specimen: Schlechter 17140, Kani Mts, 
E. New Guinea (B; dupl. at P, BM, UC). 

Distr. Malaysia: East New Guinea (one col- 
lection). 

Ecol. At 1000 m. 

119. Cyathea subdubia (v. A. v. R.) Domin, 
Pterid. (1929) 26i. —Alsophila subdubia v. A. v. R. 
Bull. Jard. Bot. Btzg II, /;. 20 (1915) 3; Handb. 
Suppl. (1917) 54. — Alsoplnla persquamulata v. A. 
V. R. Bull. Jard. Bot. Btzg II, n. 28 (1918) 1.— 
C. persquamulata Domin, Acta Bot. Bohem. 9 
(1930) 146. 

Stipe dark, slightly warty near base after fall 
of scales; stipe and lower part of rachis persistently 
scaly on each side of the hairy median band of the 
upper surface; scales to 2 mm wide, median band 
dark and shining, edges pale, bearing irregular 
dark setae. F/V/^ae to 50 cm long, lower ones smaller. 
Largest pinnules 90-115 by 15-20 mm, almost 
sessile, lobed Vj-Vi towards costa, apex acu- 
minate; costules 5-6 mm apart; veins 5-6 pairs, 
simple, lowest basiscopic vein not from costa; 
lamina-segments rather thin, lobes broadly round- 
ed and slightly crenate. Sori near costules, usually 
3 pairs, exindusiate. Scales and hairs: on lower 
surfaces of pinna-rachis and costae smaller scales 
like those of main rachis (scales on costae 3 mm 
long); no scales seen on costules. 

Type specimen: Matthew 672, Indrapura 
(G. Kerintji), Sumatra (BO; dupl. at K). 

Distr. Malaysia: Central Sumatra, W. Java. 

Ecol. At c. 1500 m. 

120. Cyathea gigantea (Wall. ^a: Hook.) Holttum, 
Gard. Bull. S. S. 8 (1935) 318; Rev. Fl. Mai. 2 



(1954) 128. — Alsophila gigantea Wall, ex Hook. 
Sp. Fil. 1 (1844) 53. — Alsophila polvcampta 
KuNZE, Bot. Zeit. 4 (1846) Al 5. —Alsophila 
helferiana Presl, Abh. K. Bohm. Ges. Wiss. 
V, 5 (1848) 34\.— Alsophila glabra [non Bl.] 
Bedd. Ferns S. India (1863) t. 60; Handb. (1883) 
14; Hook. & Bak. Syn. Fil. (1866) 43, p.p.; 
J. Scott, Trans. Linn. Soc. 30 (1874) 38; Clarke, 
ibid. II, Bot. 1 (1880) 433; Raciborski, Fl. Btzg 1 
(1898) 34; v. A. v. R. Handb. (1908) 32.— 
Alsophila umbrosa Wall, ex Ridl. J. Mai. Br. R. 
As. Soc. 4 (1926) 6, p.p. 

Stipe c. 50 cm, black or very dark, slightly rough 
after fall of scales; scales to 10 by 1 '/2 mm, dark 
brown, shining, with narrow pale fragile edges, 
not setiferous; very small dull brown scales also 
present ;pneumathodes well-spaced, 7-15 mm long. 
Pinnae commonly to 45 cm long. Largest pinnules 
short-stalked, 80-1 10 by 15-20 mm, evenly nar- 
rowed to apex, edges lobed Vj-Vi towards costa; 
costules 4'/2-6 mm apart; veins 5 or 6 pairs, simple, 
basal basiscopic vein of each group usually attached 
direct to costa; lamina-segments thin, tapering 
rather evenly from base so that sinuses are triangu- 
lar, edges more or less strongly crenate. Sori 
exindusiate, those on basal veins widely separated 
from costule, on highest veins close to costule, so 
that each group forms an inverted V; paraphyses 
dark, attenuate from a thick base, shorter than 
sporangia. Scales and hairs: pinna-rachis dark 
purplish, smooth and glabrescent or with residual 
small scales like those on the stipe; scales near 
bases of costae firm, dark, elongate, brown with 
paler edges, apex setiform but no marginal setae; 
distally and on costules similar smaller and paler 
scales, none bullate. 

Type specimen: Wallich 321, Nepal, Sylhet 
(K). 

Distr. Ceylon and S. India, NE. India south- 
wards to Burma, Siam, Indochina; in Malaysia: 
Central Sumatra, N. part of Malay Peninsula, 
and W. Java. 

Ecol. In rather open places from low elevation 
to c. 600 m. 



Subgenus Sphaeropteris 

(Bernh.) Holttum, Stat. nov. — Sphaeropteris Bernh. in Schrader, J. Bot. 1800, 
ii (1801) 122. 
Type species: Cyathea meduUaris (Forst.) Sw. 

Distr. Throughout Asia, Malaysia, Australasia and Polynesia. A few tropical American species have 
similar scales and possibly should be included in the subgenus, e.g. C. crassipes Sod., C. insignis Eaton, 
C. princeps (Linden) Meyer, and C. brunei Christ. 

Taxon. The distinction between sect. Sphaeropteris and sect. Schizocaena is on the whole quite sharp, 
but there are small species of sect. Schizocaena in Borneo which have deeply divided pinnules which, if 
larger, would be very like sect. Sphaeropteris. 

3. Section Sphaeropteris 

Distr. Throughout the whole range of the subgenus. — Fig. 1, 3, 4, 6, 17, 22-27. 

Taxon. The division into two subsections is quite sharp, subsect. Fourniera being distinct in having a 
combination of the following characters: sori lacking indusia but covered with scales like other scales on 
lower surface of pinnules, and fronds fully tripinnate. Several exindusiate species of subsect. Sphaerop- 
teris, however, have narrow scales spreading round the base of the receptacle of a sorus, so that the dis- 
tinction on this character alone is not absolute. 



Dec. 1963] 



Cyatheaceae (Holttum) 



125 



7>a. Subsection Sphaeropteris 

Distr. Throughout the range of the subgenus. — Fig. 1, 3, 4, 6, 17, 22-26. 

Taxon. The species of this subsection in New Guinea are difficult to characterize clearly. Sterile 
pinnules show the characteristic scales better than fertile ones, and arc not always present on herbarium 
specimens. The distribution of stout pale hairs on the lower surface of pinnules is usually a distinctive 
character. Most species are exindusiate. 

Ecol. Mostly large tree-ferns which early become established in secondary growth and will tolerate 
full exposure of their fronds, notably C contaminuns (Wall.) Copel. (fig. 22) which is the most widely 
distributed Cyuthea in Malaysia. 




Fig. 22. Cyathea contaminans (Wall.) Copel. in secondary growth on abandoned tea plantations, 
Tjibodas, with trees of Trema amboinensis and undergrowth of Eupatorium inulifoliuin giving ground 

cover (W. M. Docters van Leeuwen). 



KEY TO THE SPECIES 

1. Indusium present. 

2. Stipe bearing many bristles 10-20 mm long, spreading at right angles, surface of bristles covered with 
dark setae. 
3. Slender hairs 1 mm long abundant on costae, costules and veins beneath . 121. C. pulcherrima 

3. Such hairs lacking; at most much shorter hairs present towards apices of pinnules. 

122. C. procera 
2. Stipe lacking such bristles. 

4. Largest pinnules 90-120 by 16-20 mm; costae not densely scaly; no bullate scales on costae and 
costules 123. C. leucotricha 

4. Largest pinnules 50 by 10 mm; costae densely scaly; bullate scales present on costules. 

124. C. strigosa 
1. Indusium lacking. 

5. Scales of stipe-base to 50 mm or more long, shining medium brown, rigid, edges bearing sparse rather 
long concolorous setae. Lamina very rigid, edges strongly reflexed and inrolled. 



126 Flora Malesiana [ser. II, vol. P 

6. Pinnules to 30 mm long. Scales on stipe-base 2 mm wide; pinna-rachis rather persistently covered 
with a felt of small pale fringed scales; scales on costules bearing pale crisped marginal hairs. 

125. C. tomentosissima 
6. Pinnules 45-80 mm long. Scales on stipe-base 3-5 mm wide; pinna-rachis not persistently or 
densely so covered; scales on costules often bearing long dark setae. 
7. Stipe bearing many dark shining spines 3-6 mm long. Pinnules to 80 mm long. 126. C. atrox 

7. Stipe lacking spines. Pinnules commonly to 45 mm long 126. C. atrox var. inermis 

5. Scales on stipe-base mostly shorter, often not rigid and pale, edges mostly bearing close dark setae. 
Edges of lamina rarely much reflexed and then not inrolled. 
8. Upper part of stipe and all rachises finely and very closely warty; bullate scales always present on 

costules. 
9. Long pale hairs normally lacking on lower surface of costae and costules, present on upper sur- 
face of costules; no setiferous scales on costae 127. C. sangirensis 

9. Long pale hairs always rather abundant distally on lower surface of costae and costules, lacking 
or very few on upper surface of costules; setiferous scales present at least near base of costae. 
10. Veins commonly 15 pairs; setiferous scales on costae of lower pinnules only. 

128. C. lunulata 

10. Veins 9-12 pairs; setiferous scales always abundant on costae 129. C. elmeri 

8. Upper part of stipe and rachises thorny or conspicuously warty, warts not very fine and close; 

bullate scales present or not. 

1 1. Scales on costules abundant, copiously fringed with long crisped hairs which become entangled 

and form a woolly covering. 

12. Scales on costules not setiferous; lamina-segments much curved forwards when dry; hairs on 

upper surface of pinna-rachis pale 130. C. tomentosa 

12. Scales at bases of costules bearing many long dark setae; lamina-segments spreading at right 
angles to costa; hairs on upper surface of pinna-rachis dark 131. C. magna 

11. Scales on costules not thus fringed. 

13. Bullate scales present on lower surface of costae and or costules. 
14. Bullate scales present on lower surface of veins. 

15. Copious long hairs also on lower surface of veins 132. C. pilulifera 

15. Long hairs lacking on lower surface of veins, or only at vein-tips. 
16. Pinna-rachis closely warty, warts dark; pinnules to 13 mm wide. Luzon. 133. C. curranii 
16. Pinna-rachis sparsely thorny or warty; pinnules 15-25 mm wide. New Guinea. 

134. C. aeneifolia 
14. Bullate scales lacking on lower surface of veins. 
17. Stipe conspicuously spiny; pale hairs on lower surface of costae few or lacking. 

135. C. tenggerensis 
17. Stipe warty; pale hairs abundant on lower surface of distal half of costae and on costules. 
18. Costae copiously scaly throughout, with small pale-fringed scales as well as elongate se- 
tiferous ones; bullate scales not setiferous 136. C. persquamulifera 

18. Costae rather sparsely scaly, small pale-fringed scales lacking; bullate scales on costules 

setiferous 137. C. sarasinorum 

13. Bullate scales lacking on lower surface of costae and costules. 
19. Pale erect hairs abundant on lower surface of veins. 
20. Costae bearing many much-setiferous scales; pinnules to 20 mm wide. 
21. Veins flat or impressed beneath; costular scales with long brown setae. 138. C. angiensis 
21. Veins much raised beneath; costular scales with short dark setae . . 139. C. verrucosa 
20. Costae bearing very few scales; pinnules commonly 20-30 mm wide. 140. C. contaminans 
19. Pale erect hairs absent or rare (near apices of segments) on lower surface of veins. 
22. Stipe-scales pale, 40-50 mm long, 4-5 mm wide at base, distal half very narrow; costular 

scales broad, pale, flat, short-setiferous or short-fringed 141. C. lepifera 

22. Stipe-scales otherwise; costular scales (if present) narrower, bearing long setae. 

23. Setiferous scales abundant on costae; small scales present on lower surface of veins; no 

stout hairs on lower surface of costae. 

24. Stipe-scales 5 mm wide. Pinnules to 25 mm wide. Veins 13-15 pairs. 142. C. atrospinosa 

24. Stipe-scales 2 mm wide. Pinnules to 35 mm wide. Veins to 20 pairs. . . 143. C. fugax 

23. Setiferous scales lacking or rare on lower surface of costae of mature fronds; no scales on 

lower surface of veins; stout pale hairs present near apices on lower surface of costae. 

140. C. contaminans 

121. Cyathea pulcherrima Copel. Un. Cal. Publ. 60-100 cm, covered closely with rigid spreading 

Bot. 18 (1942) 219; Philip. J. Sc. 77 (1947) 119, dark brown bristles 2 cm long, their surfaces 

pi. 12. covered with short dark setae, some bristles flat 

Trunk to 10 m, 3-5 cm below completely at apex and bearing marginal setae; lower surface 

caducous fronds; leaf-scars 1 '/2 cm 0. Stipe of rachis bearing similar shorter bristles mixed 



Dec. 1963] 



Cyatheaceae (Holttum) 



127 



with stout spreading reddish to pale hairs; similar 
hairs on lower surface of pinna-rachis. Lamina 
100-190 cm long; longest pinnae 55-70 cm long. 
Pinnules to 100 by 20-24 mm, lobed to a narrow 
wing between the segments which are separated by 
wide sinuses; at least half the segments constricted 
on acroscopic side at base; costules 4-4 '/2 mm 
apart; lamina thin, the fertile segments deeply 
crenate or more usually lobed '^j towards the 
costule, each lobule with 2 or 3 teeth; veins to 
8 pairs, forked once or twice. Sori one to each lobe, 
indusiate; indusium thin, translucent, covering the 
young sorus, breaking irregularly and persistent; 
paraphyses as long as sporangia. Pale slender 
spreading hairs 1 mm long abundant on both 
surfaces of costae, costules and veins; scales on 
lower surface of costae near base small, flat with 
copious dark marginal setae, most scales on 
costae pale and buUate at base, dark-setose to- 
wards apex (some setae are superficial), these 
grading to pale buUate hair-pointed scales distally 
and on costules. 

Type specimen: Brass 8940, east slopes of Cy- 
clops Mts, 575 m, W. New Guinea (UC; dupl. at 
K, BM). 

Distr. Malaysia: New Guinea, both west and 
east, and Admiralty Is. 

Ecol. In forest, 100-1100 m. 



122. Cyathea procera Brause, Bot. Jahrb. 56 
(1920) 50; CoPEL. Philip. J. Sc. 77 (1947) 104.— 
Fig. 23. 

Trunk up to 20 m (Pulle), bearing fronds in 
whorls of 6; fronds 250-350 cm long. Stipe 
densely covered with spreading bristles as in 
C. pulchenima but these sometimes pale above the 
base; bristles in the adaxial groove at base of stipe 
flattened from near the base to form flat dark 
scales 1/3 mm wide with setiferous margins; very 
small strongly setiferous scales also on surface of 
stipe between bristles; upper part of stipe pale, 
copiously warty from bases of abraded bristles. 
Rachis finely warty on lower surface, glabrescent 
except for some very small scales. Pinnae to 
65 cm long. Largest pinnules 85-120 by 18-23 mm 
wide, shape and sori as in C. pulcherrima. Pinna- 
rachis beneath as main rachis, upper surface bear- 
ing narrow pale dark-setiferous scales to 5 mm 
long with the usual antrorse hairs; costae beneath 
copiously scaly, scales small, basal ones entirely 
dark and shining with setae on edges, grading to 
distal ones and those on costules which are pale 
and buUate at base, apex with dark setae, or the 
smallest (these sometimes also on veins) with a 
pale fringe of fine hairs; in some cases thick pale 
hairs distally on lower surface of costae and cos- 
tules. 

Type specimen: Ledermann 11856, Sepik 
Region, 2070 m, NE. New Guinea (B). 

Distr. Malaysia: New Guinea. 

Ecol. In forest, at 1800-2400 m. Pulle noted 
on Mt Hellwig that this was the largest tree-fern 
he had seen. C. procera is the high-mountain 
counterpart of C pulcherrima. 




Fig. 23. Cyathea procera Brause. a. Segment of a 
pinnule, upper surface, • 6, b. part of costa and 
fertile segment, showing scales and sori, x 6, 
c. spine from base of stipe, x 4, d. flattened tip 
of stipe-spine, x 13, £>. scale from pinna-rachis, 

X 13 (HOOGLAND & PULLEN 5502). 

123. Cyathea leucotricha Christ, Ann. Jard. Bot. 
Btzg 20 (1905) 135; v. A. v. R. Handb. (1908) 18. 
— Alsophila cyclodonta Christ, Ann. Jard. Bot. 
Btzg 20 (1905) 137.— C. cyclodonta v. A. v. R. 
Bull. Dep. Agr. Ind. Neerl. //. 18 (1908) I; Handb. 
(1908) 19. 

Stipe 50 cm, minutely warty; scales mostly 
caducous, dark brown with concolorous marginal 
setae. Pinnae: lowest hardly reduced, longest 60 
cm. Pinnules: largest 90-120 by 16-20 mm, sessile 
or nearly so, apex acuminate, 1-3 basal segments 
free or nearly so (lowest may be stalked), next 
one or more pairs of segments contracted at base 
on acroscopic side, rest of pinnule lobed nearly 
to costa; costules 4-5 mm apart; veins 10-12 pairs; 
segments firm, edges crenate. Sori nearer costule 
than edge, indusiate; indusium pale and thin, 



128 



Flora Malesiana 



[ser. II, vol. F 



covering young sorus completely, breaking ir- 
regularly and persistent. Scales and hairs: pinna- 
rachis glabrescent; costae bearing a few small 
flat elongate dark-setiferous pale scales near base 
(some caducous) and throughout many stiff 
spreading pale hairs; costules bearing similar 
hairs, and a few on veins also; a few hairs present 
on upper surface of costules and veins. 

Type specimen: Hallier 2302, Borneo, Mt 
Klam (BO; dupl. at P, L, S). 

Distr. Malaysia: Borneo (several widely- 
spaced localities). 

Ecol. In forest, up to c. 700 m. 

124. Cyathea strigosa Christ, Ann. Jard. Bot. 
Btzg 15 (1898J 84, t. 13, f. 6; v. A. v. R. Handb. 
(1908) 24. 

Stipe 45 cm long, bearing spines 3 mm long and 
pale setiferous scales; main roc/iis pale beneath 
with many spines under 1 mm long, scales mostly 
caducous. Pinnae to 20 cm long. Pinnules: largest 
50 by 10 mm, lobed nearly to costa, sessile; cos- 
tules 3 mm apart; veins 8-9 pairs. Sori near 
costules, indusiate; indusium pale, at first covering 
sorus, breaking irregularly and persistent. Scales 
and hairs: pinna-rachis sparsely warty, glabrescent; 
costae densely scaly beneath, scales mostly bullate- 
acuminate, pale-fringed near apex which is a 
long hair, grading to long pale hairs distally, near 
base also some narrow pale dark-setiferous 
scales; costules bearing similar bullate scales and 
hairs; no hairs on lower surface of veins; upper 
surface of costules bearing a few long pale hairs. 

Type specimen: Warburg 16793, summit of 
Mt Wawo-Karaeng (= Bonthain), 2800 m, 
SW. Celebes (P?; dupl. at B). 

Distr. Malaysia: SW. Celebes (one collection). 

Note. Described from duplicate specimen at 
Berlin, which is sterile. This species appears to be 
most nearly related to C. leucotricha of the low- 
lands of Borneo. 

125. Cyathea tomentosissima Copel. Un. Cal. 
Publ. Bot. 18 (1942) 219; Philip. J. Sc. 77 (1948) 
123, p. 15. 

Trunk to 21/2 m, 16 cm o, 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 flexuous 
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 inroUed, 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, 
BO, L, UC). 

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. Chr. Brittonia 2 (1937) 
275; Copel. Philip. J. Sc. 77 (1947) 115.— 
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 30^5 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. 

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. \ar. imrmis Holttum in peaty grassland, Eastern Highlands, Territory 
of New Guinea, 3000 m (R. D. Hoogland). 



130 



Flora Malesiana 



[ser. II, vol. 1' 



var. inermis Holttum, var. nov. — Fig, 24, 25. 

Stipes verrucosus, vix spinosus; pcileae stipitis 
6-10 cm longae, ad 6 mm latae; pinnulae vulgo ad 
45 X 12 mm; costulae lyj '"'" ""'^'' ^^ distantes; 
rhaches pinnarum subtus paleis mimitis ciliatis pal- 
lidis vestitae. 

Stipe with numerous conical warts Vi 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 2'/2 rnm 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 



(Brass 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. il.-'Alsophila concinna Bak. 
Syn. Fil. ed. 2 (1874) 459, non C. concinna (Bak.) 
Jenm. 1891. — Alsophila pohphlebia Bak. J. Linn. 
Soc. Bot. 15 (1876) 104, non C. pohphlebia Bak. 
1883. — Alsophila sangirensis Christ in Warb. 
Monsunia 1 (1900) 90. — Alsophila scaberula 
Christ in K. Sch. & Laut. Fl. Deut. Schutzgeb. 
Sudsee(1901) 110; v. A. v R. Handb. (1908) 35.— 




Fig. 25. Cyathea atrox C. Chr. var. inermis Holttum on the Sugarloaf complex, Western Highlands 

(R. D. Hoogland). 



Dec. 1963] 



Cyatheaceae (Holttum) 



131 



C. scabriseta Copel. Philip. J. Sc. 9 (1914) Bot. 2; 
ibid. 11 (1947) HI. — Alsophi/a okiana v. A. v. R. 
Bull. Jard. Bot. Btzg 11, //. 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) \A.— Alsophila scaberulipes 
V. A. V. R. Nova Guinea 14 (1924) 2. — C. eminens 
DOMiN. Pterid. (1929) 262 (new name for Alsophila 
concinna Bak.); Copel. Philip. J. Sc. 77 (1947) 
116.— C. ariiensis Domin, Pterid. (1929) 262 
(new name for Alsophila polyphlebia Bak.); Co- 
pel. Philip. J. Sc. 77 (1947) 115.— C. scaberula 
DoMiN, Pterid. (1929) 263; Copel. Philip. J. Sc. 
77 (1947) 109.— C. brassii Copel. J. Am. Arb. 10 
(1929) 175; Philip. J. Sc. 77 (1947) 111.— C. 
buruensis Domin, Acta Bot. Bohem. 9 (1930) 102. 
— C. scaberulipes Domin, I.e. 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 4-41/2 rnm 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, a little 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 buUate; 
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: Warburg 16605, Sangir (= 
Sangihe)I. (P?;dupl. atB). 

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. Merrill (I.e.) has suggested that 
Palmifilix alba Rumph. (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.) Copel. Bull. Bern. 
P. Bish. Mus. 59 (1929) 37; C. Chr. ibid. 177 
(1943) 29. — Polypodium lunula turn Forst. Prod. 
(1786) 83, /;. 456.— Alsophila lunulata R. Br. 
Prod. (1810) 158; Hook. Sp. Fil 1 (1844) 51; 
Syn. Fil. (1868) 41, p.p.; Carr. in Seem. Fl. Vit. 
(1873) 333. — Alsophila vitiensis Carr. in Seem. 
Fl. Vit. (1873) 170. — Alsophila naumannii KuHN, 
Forschungsr. Gazelle 4 (1889) Fame 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 I V2 
mm, pale, thin, with a few dark marginal setae. 
Pinnules to at least 1 20 by 20 mm; costules 3 Vi-^Vi 
mm apart; segments of lamina distinctly crenate- 
serrate; veins to 15 pairs. Sori e.xindusiate; 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 (Gottingen; 
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. Cvathea elmeri Copel. 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 inon 
Wall.) var. major Christ, Philip. J. Sc. 2 (1907) 
Bot. \S3.— Alsophila elmeri Copel. in Elmer, 
Leafl. Philip. Bot. 2 (1908) 419; v. A. v. R. 
Handb. Suppl. (1917) 66.— Alsophila christiiv. A. 
V. R. Handb. (1908) 42 {non Sod. 1908); Suppl. 
(1917) 69. 70.— C. dimorphotricha Copel. in Elmer, 
Leafl. Philip. Bot. 5 (1913) 1681; Fern Fl. Philip. 2 
(1960) 230. — Alsophila subcomosa C. Chr. Ind. 
Fil Suppl. 1 (1913) 5. — Alsophila dimorphotricha 
V A V R. Handb. Suppl. (1917) 61.— C. subco- 
mosa Domin, Pterid. (1929) 163.— Alsophila fe- 
nicis [non (Copel.) C. Chr.] Posth. in Holthuis & 
Lam. Blumea 5 (1942) 153, p.p.—C. argyrolepis 
Copel. Philip. J. Sc. 81 (1952) 17; Fera Fl. Philip. 
2 (1960) 227. 

Stipe 30 cm or more, the base densely scaly, 
closely warty when scales have fallen; scales 
25 by \Vi rrirn. 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. 12 



next few segments separately adnate to costa, 
rest (whole of smaller pinnules) lobed nearly to 
costa; costules 3 — 4'/2 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 buUate-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: Elmer 9457. Cuernos Mts, 
Negros (MICH; dupl. at US, BO, F. K, P, L, 
U, SYD). 

Distr. Malaysia: Philippines (Mindanao, Ley- 
te, Negros, Biliranj, 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 Holthlis & 
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.) Merr. from Guam is very 
nearly allied. 

130. Cyathea tomentosa (Bl.) Zoll. & Mor. in 
Moritzi, Syst. Verz. (1846) lOS.—Chnoophora 
tomentosa Bl. En Pl. Jav. (1828) 244.— C/;/;oc;- 
phora lanuginosa Jungh. Tijd. Nat. Gesch. & 
Physiol. 8 (1841) 372; Flora 30 (1847) 522.— 
Alsophila tomentosa Hook. Sp. Fil. 1 (1844) 55; 
Racib. F1. Btzg 1 (1898) 32; v. A. v. R. Handb. 
(1908) 43; Suppl. (1917) 493; Backer & Posth. 
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 
4-4 V^ cm 0. 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 1 10 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-^ 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 with a 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: Blume, Mt Gedeh. Java (L). 

Distr. Malaysia: Ja\a.. Lesser Sunda Is (Flores). 

Ecol. .M 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.) Corel., 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. 

13L Cyathea magna Corel. Ln. 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 Corel. 
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 l'/2'-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 



Dec. 1963] 



Cyatheaceae (Holttum) 



133 



marginal hairs); costules densely scaly, scales 
near base narrow flat and bearing long dark setae, 
rest light brown with -Az biiUate base and long 
pale flexuous marginal hairs; erect hairs present 
distaliy on costules and on lower surface of 
veins; upper surface of costules not hairy; hairs on 
upper surface of pinna-rachis darlc 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, 
pi. 9. 

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 mni, 
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- 
taliy 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), New 
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). 

133. Cyathea curranii Copel. Philip. J. Sc. 3 
(1909) Bot. 356; ibid. 4 (1910) Bot. 52; Fern 
Fl. Philip. 2 (1960) 231.— Alsophila curranii 
C. Chr. Ind. Fil. Suppl. I (1913) 5; v. A. v. R. 
Handb. Suppl. (1917) 70. 

Trunk 3 m, 20 cm 0; leaf-scars 3 cm 0. 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 
P, US). 

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. v. R. I.e. 4. — C. curvipinmda 
C. Chr. Brittonia 2 (1937) 276; Copel. Philip. 
J. Sc. 77 (1947) 116. 

Trunk to 41/2 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 3-41/2 mni 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 15 -1 
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 1 75 1 , 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. II, vol. P 



is otherwise similar. This species is nearly allied to 
C piluUfera 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, pi. 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 diff'ert: frondibus ad 425 cm longis, 
stipitibus 200 cm longis inchisis; paleis stipitis ad 
50 X 4 mm; pinnis maximis 70 cm longis, pinnulis 
maximis 135 X 21 mm; costidis 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) \65.~Alsophila luienkei 
var. angiistata Hassk. in Hook. J. Bot. Kew Misc. 
7 (1855) 326. — Alsophila glaiica (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 2'/2-3i/2 mrn, 
shining pale brown, firm, setiferous. Pinnae to at 
least 50 cm long. Pinnules to 110 by 16 mm, 
sessile, lobed almost to costa with 1 or 2 basal 
segments almost free; costules 2V2-3 mm apart; 
lamina-segments firm, edges i reflexed when dry, 
lower surface probably glaucous; veins 10-12pairs. 
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 1/2 mrn, 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: Hasskarl, 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. Ill, 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, 1 or 2 basal segments free or 
contacted at base, rest of pinnule cut nearly to 
costa, sessile, caudate-acuminate; costules 3'/2-4 
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, MtTalamau, 
2300 m, Sumatra (BO; dupl. at K, L). 

Distr. Malaysia: Central Sumatra, throughout 
Java. 

Ecol. On mountains at 1500-2500 m. 

137. Cyathea sarasinorum Holttum, Kew Bull. 
16 (1962) 61. — Alsophila contaminans var. longe- 
paleata Christ, Ann. Jard. Bot. Btzg 19 (1904) 42. 
— Alsophila glauca var. longepaleata V. A. v. 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-Rucken, Central Celebes (BAS). 

Distr. Malaysia: Central Celebes (one collec- 
tion). 



Dec. 1963 ] 



Cyatheaceae (Holttum) 



135 



138. Cyathea angiensis (Gepp) Domin, Acta Bot. 
Bohem. 9 (1930) 90. — Alsopliila oiifdensis Gepp 
in Gibbs, Arfak (1917) 69. 

Like C. contamincins (Wall.) Copel., but rather 
small; pinnules 100-150 by 16-20 mm; costules 
31/2 nim apart, veins flat or impressed on lower 
surface; lamina-segments very tirm, 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: Gibbs 5968, Angi Lakes, 7000 ft, 
W. New Guinea (BM; K, P, BO). 

Distr. Malaysia: Moluccas (Buru), New 
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. contaminaiis: 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. coutaminans. 

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 I mm high). Pinnae to at least 60 
cm long, lower ones with stalks 6 cm long. Pinnules 
to 110 by 20 mm, sessile or the lowest stalked, 
caudate-acuminate, basal 1-2 segments free or 
nearly so, rest cut almost to costa; costules 31/2-4 
mm apart; lamina-segments rigid, edges minutely 
crenate, sinuses narrow; veins to 12 pairs. Sori 
at about 1 3 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 1 1/2 rnni 
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 buUate, 
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, /;. 28 (1918) 13; Copel. 
Philip. J. Sc. 77 (1949) 115; Holttum. Rev. FL 
Mai. 2 (1954) 119; Copel. Fern Fl. Philip. 2 
(1960) 230. — Polypodium contaminans Wall. Cat. 
(1828) /;. 320, nomen. — Chnoophora glauca Bl. 
En. PI. Jav. (1828) 243 {non C. glauca Bory, 1804). 
—Alsophila glauca J. Sm. J. Bot. 3 (1841) 419; 
Bedd. Handb. (1883) 12; v. A. v. R. Handb. 
(1908) 41 {incl. var. celebica, var. squamiilata, 
var. densa, var. setulosa, and var. microloba); 
Koord.-Schum. Syst. Verz. 1, 2 (1912) 5; Ro- 
senst. Hedwigia 56 (1915) 349, ittcl. var. tricho- 
carpa Rosenst.; v. 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; Hassk. 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) pi. 85; Hook. Syn. Fil. (1866) 41; Scott, 
Trans. Linn. Soc. 30 (1874) 35; Christ, Verh. 
Nat. Ges. Basel 11 (1895) 199, incl. var. celebica 
Christ; Farnkr. Erde (1897) 327; Ann. Jard. Bot. 
Btzg 15 (1898) 79, pi. XIII f. 2; Diels in E. & P. 
Pfl. Fam. 1, 4 (1899) 136, p.p.— Alsophila acuta 
Presl, Abh. k. Bohm. Ges. Wiss. V, 5 (1848) 343. 
— Alsophila smithiana Presl, I.e. 'iAl. — Alsophila 
dementis Copel. Philip. J. Sc. 1, Suppl. 2 (1906) 
Bot. 143.— C. dementis Copel. 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 4—4 V2 mrn apart, rarely to 5 mm; 
segments of lamina firm, glaucous beneath, 
edges ±: crenate-serrate; veins commonly 12 pairs. 



136 



Flora Malesiana 



[ser. II, vol. P 




Fig. 26. Cyathea contaminant (Wall.) Copel. in Tjibodas mountain garden (van Steems). 



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 V2 iTim» shorter 
distally, these in most cases early caducous leaving 
the costae glabrous; costular scales small, i 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. lepifera (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 (I.e.) that typical Alsuphila coiuaminans 
occurred there only at low altitudes. He distin- 
guished var. brununiana {A. hnuwiiiana HoOK.), 
which is common in Sikkim at higher altitudes and 
has smooth (not spiny) stipes and copiously scaly 
costae and costules. 1 prefer to regard this as a 
distinct species, which must be called Cyathea 
brunoniaiui (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.) Copel. Philip. J. 
Sc. 4 (1909) Bot. 40; ibid. 56 (1935) 98, pi. 3, 
fig. 4-7; Fern Fl. Philip. 2 (1960) 22S.—Alsophila 
lepifera J. Sm. e.x Hook. Sp. Fil. 1 (1844) 54; 
Christ, Bull. Herb. Boiss. 6 (1898) 137, incl. var. 
congesta Christ; v. A. v. R. Handb. (1908) 39; 
Suppl. (1917) 65. — Ahophila pustulosa Christ. 
Bull. Herb. Boiss. II, 1 (1901) 1019; Nakai. 
Bot. Mag. Tokyo 41 (1927) 73; Ito, Fil. Jap. 111. 
(1944) pi. 453. — Alsophila calocoma Christ, 
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 Copel. I.e. 53; Fern 
Fl. Philip. 2 (I960) 229.— C. umbrosa Copel. 
Philip. J. Sc. 56 (1935) 98, pi. 3, fig. 1-3.— (?) 
C. pteridioides Copel. I.e. 98, pi. 4; Fern Fl. 
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 2-4 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 1/2 mrri wide), brown, with 
concolorous marginal setae. Raeliis 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 3-3 1/2 
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: Cl.ming 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 Holttum, 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 3 '2-4 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: Hoogland & Pullen 6090, 
Western Highlands District, 2650 m, NE. New 
Guinea (K). 

Distr. Malaysia: E. New Guinea. 

Ecol. In mountain forest, on limestone (always?) 
at c. 2400-2850 m. 

143. Cyathea fugax v. A. v. R. Bull. Jard. Bot. 
Btzg II, /;. 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 0, 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 41/2 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. P 



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: Copland 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 Hook.) Copel., 
which differ notably in scales from the type. 



2)b. Subsection Fourniera 

(Bommer) Holttum. slat. nov. — Fourniera Bommer, Bull. Soc. Bot. Fr. 20 (1873) 
xix. — Fig. 27. 
Type species: Cyathea novae-caJedoniae (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. 

148. C. aciculosa 
4. Bullate scales lacking. 

6. Veins and lower surface of lamina bearing finely fringed interlacing 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 nor covering lower surface of leaflets. 

151. C. macrophylla 



144. Cyathea carrii Holttum, Kew Bull. 16 (1962) 
53.— Fig. 27. 

Stipe c. 1 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 rac/iis 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 



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) 



139 




'62. 







Fig. 27. Cyathea carrii Holttum. a. Single pinnule, 
upper surface, y. 2, b. part of pinnule, lower 
surface, showing scales and sori, x 6, c. sorus, 
showing covering of overlapping scales, x 20, 
d. scales from costule, x 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 woraersleyi Holttum, 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 21/2 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 1/5 mm; main rac/iis 
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-buUate 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: Womersley & Millar NGF 
8470, Skindewai, 5400 ft, Morobc 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 Nuthufagus forest 

(SCHODDE). 

146. Cyathea auriculifera Corel. Philip. J. Sc. 6 
(1911) Bot. 364; ibid. 11 (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 1 1 cm long, largest 54 cm. 
Pinnules slightly overlapping, to 70 by 15 mm, 
fully pinnate, bases of tertiary leaflets 3 mm 
apart; tertiary leaflets mostly stalked, largest 9 by 
21/2 rnm. 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 Corel. 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. v. R. Bull. 
Dep. Agr. Ind. Neerl. 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. II, vol. P 



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, pi. 9. — C arachnoidea {non Hook.) 
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. triincata 
Brack. ) Copel. of Fiji and Samoa, differing in the 
much larger indusial scales. 

149. Cyathea celebica Bl. En. PI. Jav. (1828) 245 
(not V. A. v. R. 1908); Tindale, Contr. N.S.W. 
Nat. Herb. 2 (1956) 338, p.p.—Alsophila celebica 
Mett. 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) 3S.—Alsophila triincata 
var. sagittata Christ, Bull. Herb. Boiss. II, 1 
(1901 ) 458. — Alsophila triincata var. nivea Christ, 
K. ScH. & Laut. Nachtr. (1905) 36; v. A. v. R. 
Handb. (1908) 42.— C. quadripinnatifida Copel. 
Un. Cal. Publ. Bot. 18 (1942) 218; Philip. J. Sc. 
77 (1947) 108, pi. 6. 

Stipe to 100 cm long, dark, bearing slender dark 
spines to 3 mm long, near base bearing persistent 
light brown scales 20^0 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 leaflets stalked; costules (stalks 
of tertiary leaflets) 3-5 mm apart; tertiary leaflets 
to 15 by 21/2-5 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. 
atBM). 

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 Copel. Philip. J. Sc. 1, 
Suppl. 4 (1906) Bot. 251; ibid. 4 (1909) Bot. 40; 
v. A. v. R. Handb. (1908) 788; Suppl. (1917) 39; 
HoLTTUM, Rev. Fl. Mai. 2 (1954) 120; Copel. 
Fern Fl. Philip. 1 (1960) 208.— C. densisora v. A. 
V. R. Bull. Jard. Bot. Blzg III. 2 (1920) 138; Copel. 
Fern Fl. Philip. 2 (1960) 208. — C. leucostegia 
Copel. Philip. J. Sc. 38 (1929) 130; Fern Fl. 
Philip. 2 (1960) 209.— C. levtensis Copel. Philip 
J. Sc. 38 (1929) 131; Fern Fl. Philip. 2 (1960) 209. 
— C. arachnoidea (non Hook.) Backer & Posth. 
Varenfl. Java (1939) 25. 

Trunk to 4 or 5 m. 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 1 7-25 
mm, fully pinnate, lower tertiary leaflets distinctly 
stalked; costules 4-6 mm apart; tertiary leaflets 
to 15 by 3'/2mrni 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 
US, P, UC, SYD, S-PA). 

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 (new name for Hemitelia 
ledermaniiii 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 foliolis tertiariis maximis 18 mm longis, omnino 

long-fringed nor covering lower surface of lamina. prufunde lohatis, segmentis infimis interdum liheris 

Type specimen: Ledermann 12533, Sepik et stipitulatis. 

Region, E. New Guinea (B). Type specimen: Womersley NGF 13959, Mo- 

Distr. Malaysia: New Guinea. robe Distr., NE. New Guinea (K). 

Ecol. In forest, from sea-le\el to 1500 m. Ecol. '"Short tree-fern in under-storey of the 

forest. Trunk not more than 2 feet tall. Fronds 

var. quadripinnata Holttum, var. now 5 fegf 2000 m. 

A typo speciei differ t: stipitibits c. 10 cm longis; 

4. Section Schizocaena 

(J. Sm.) Holttum. stat. now — Schizocaena J. Sm. in Hook. Gen. Fil. (1838) t. 2; 
Lond. J. Bot. 1 (1842) 66\, 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 152. C. capitata 

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 153. C. angustipiiina 

3. Pinnae 2-4 cm wide, base rather broadly cuneate, stalked or not. Sori in fully fertile fronds in more 
than 2 rows, indusiate or not. 

4. Pinnae not long-acuminate, upper usually sessile 154. C. moluccana 

4. Pinnae long-acuminate, all stalked 155. C. arthropoda 

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 long as lobes, apex not long-acuminate; 
pinnae commonly 25 cm long 157. C. alternans 

8. Lower pinnae narrowed to base so that free leaflets 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 157. C. alternans 

9. Pinnae much shorter; main rachis bearing long pale hairs 159. C. elliptica 

5. Fronds amply bipinnate. 
10. Pinnules entire or with crenate edges. 
11. Indusia present; pinnules sessile 157. C. alternans 

11. Indusia lacking; pmnules stalked 160. C. obliqua 

10. Pinnules distinctly lobed. 

12. Sori indusiate. 

13. Pinnules of larger pinnae not lobed more than 3^ to costa throughout, no free basal pinnules, 
or sometimes one on lowest pinnule of lower pinnae. 



142 Flora Malesiana [ser. II, vol. P 

14. Bullate scales lacking on costules 161. C. Integra 

14. Bullate scales present on costules. 
15. Basal pinnules of middle pinnae with stalks to at least 4 mm long; pinnules lobed less than 

1/2 way to costa 162. C. stipitipinnula 

15. Basal pinnules sessile or nearly so; pinnules lobed more than half way to costa. 
16. Segmentsof 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 164. C. discophora 

17. Indusium complete, breaking and persistent at maturity. 

18. Lower surface of costae densely scaly 165. C. megaiosora 

18. Lower surface of costae not densely scaly 166. C. suluensis 

13. Pinnules of larger pinnae always with one or more free basal segments. 
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 167. C. robinsonii 

20. One or two pairs tertiary segments free on largest pinnules. Pinnules to 80 by 12 mm. Costae 
densely scaly. 

21. Costal scales not setiferous. Pinnules sessile 165. C. megaiosora 

21. Costal scales strongly setiferous. Lower pinnules with stalks 2-3 mm long. 

168. C. senex 
19. No hairs on lower surface of veins. 
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 f. 65 by 15 mm, often smaller 170. C. philippinensis 

23. Pinnules stalked (stalks to 8 mm), to 100 by 25 mm 171. C. assimilis 

12. Sori without indusia. 

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 %-% to costa; costules 5 mm apart . . 172. C. trichodesma 

26. Pinnules to 65 mm long, cut to within 1 mm of costa; costules 3-3 '/2 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 175. C. polypoda 

28. Largest pinnules lacking a free segment at the base, almost sessile. 
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 178. C. squamulata 

152. Cyathea capitata Copel. Philip. J. Sc. 12 truncate to cordate, in the latter case more or less 
(1917) Bot. 49; C. Chr. & Holttum, Gard. Bull. auricled on both sides, apex short-acuminate, 
S. S. 7 (1934) 199, 218. — Scluzocaena capitata crenate. Ke///5 pinnate in each group, usually with 
Copel. Gen. Fil. (1947) 99. — Fig. 28f-h. 3 pairs of veinlets, outer veinlets of each group 
Trunk 1-3 m, bearing c. 12 fronds. Stipe dark, always anastomosing with outer veinlets of ad- 
smooth, at least 40 cm; basal scales pale brown, jacent groups to form a single excurrent vein; 
firm, to 25 by 3-4 mm, rather thick at the base, basal basiscopic vein of each group springing 
edges bearing rather irregular concolorous setae; separately from the costa. Sori usually in 2 rows 
pneumathodes 9-15 mm long, rather widely on each side of the costa (3-4 sori on each vein- 
spaced. Rachis dark to medium brown, smooth; group) or in narrower pinnae only one row; 
lamina 100 cm or more long, simply pinnate, apex indusium thin and translucent, at first covering 
of frond not like a pinna but broadly deltoid and sorus, later breaking irregularly and more or less 
deeply lobed, lobes grading to upper pinnae. persistent. 

Pinnae c. 40 pairs, sessile, jointed to rachis, largest Type specimen: Clemens 11033, Mt Kinabalu, 

15-19 by 2-3 cm wide at base, lower ones some- N. Borneo (MICH; dupl. at A, UC, K). 

what smaller, edges entire except near apex, base Distr. Malaysia: Borneo (Sarawak: Mt 



Dec. 1963] 



Cyatheaceae (Holttum) 



143 



Murud; N. Borneo: Mt Kinabalu). 

Ecol. Near a waterfall and in wet ground near 
a stream, in forest, 1400-2100 m. 

153. Cyathea angustipinna Holttum, Kew Bull. 
16 (1962) 52. 

Stipe 30 cm, smooth, scaly near base, scales pale, 
firm and shining, to c. 20 by 1 Vz 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; vc'//w 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: Borneo (Sarawak, two 
collections). 

Ecol. Sandy bank of a stream, near waterfall, 
in shade; trunk 50 cm. 

154. Cyathea moluccana R. Br. in Desv. Mem. 
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 II, n. 28 (1918) 
12. — Schizocaena bninonis 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; Bedd. Ferns Br. Ind. 
(1865) pi. 87; Handb. (1883) 5; Christ, Farnkr. 
Erde (1897) 318; Diels in E. & P. Pfl. Fam. 1, 4 
(1899) 127; Copel. Sarawak Mus. J. 2 (1917) 345, 
347; Holttum, Rev. Fl. Mai. 2 (1954) 117.— 
C. pimiata Roxb. Calc. J. Nat. Hist. 4 (1844) 517; 
Fl. Ind. ed. Clarke (1874) 762.—Schizocaena 
gaudichaudii Fee, Gen. Fil. (1852) 354.— C. 
pseudobriinouis Copel. Philip. J. Sc. 12 (191 7) 
Bot. 50. — C. fuscopaleata Copel. I.e. 50. — C. 
kinabaliiensis Copel. I.e. 51; C. Chr. Gard. Bull. 
S.S. 7 (1934) 218. — Schizocaena moluccana Co- 
pel. Gen. Fil. (1947) 99. — Schizocaena kinaba- 
liiensis Copel. I.e. 99. — Fig. 28a-e. 

Trunk usually not over 50 cm tall. 5///Je common- 
ly 20-30 cm, dark, scaly near base and finely 
warty when scales have fallen; scales medium 
brown, firm, 15-30 by '/2-3 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- 



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 usually 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 I.e.). 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. HoU. 158); Desvaux 
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 Copel. 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 21/2 cm wide, sterile to 31/2 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: Brooks 8, Bungo Range, 
Sarawak (MICH; dupl. at BM). 



144 



Flora Malesiana 



[ser. II, vol. P 




Fig. 28. Cyathea moliiccana R. Br. a. Middle part of rachis with pinnae, ■' Vi, b. apex of frond, X Vi, 
c. part of pinna, showing venation and sori, x 1 1/2 , ^- scale from stipe, x 4, e. edge of stipe-scale, ■' 40. 
— C. cfl/j/raro CoPEL./. Middle part of rachis with pinnae, X Vi, g'- apex of frond, x 1/2? /'• part of pinna 
showing venation and sori, x 1 1/2 {a-e Cuming 378, /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, nam. nov. — 
Alsophila parvifolia Holttum, J. Mai. 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 1 '/2 mm, 
edges closely set with dark setae. Lamina to 65 cm 
long, simply pinnate, pinnae pinnatifid. lower ones 
gradually reduced, lowest 3' 2 cm long. Rachis 
densely covered beneath throughout with stout 
pale spreading hairs to 3 mm long. Largest pinnae 
9 by 2 cm, lobed 2/3 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. Bohm. Ges. Wiss. V, 5 (1848) 347; 
Bedd. Handb. Suppl. (1892) 2; v. A. v. R. Handb. 
(1908) 17; RiDL. J. Mai. Br. R. As. Soc. 4 (1926) 
5, incl. var. serrata Ridl.; Domin, Acta Bot. 
Bohem. 9 (1930)90, incl. var. sarawakensis (Hook.) 
DoMiN and \ar. lobbiana (Hook. ) Domin; Holttum 
Rev. Fl. Mai. 2 (1954) \\9.^Polypodium alternans 
Wall. Cat. (1829) n. 119, nom. nud. — Hemitelia 
alternans Hook. Ic. PL (1844) t. 622; Sp. Fil. 1 
(1844) 29. — Amphicosmia alternans Moore, 
Ind. Fil. (1857) civ; Bedd. Handb. (1883) 
10.— C. lobbiana Hook. Syn. Fil. (1865) 
24. — C. sarawakensis Hook. I.e. 23; Hose, 
J. Str. Br. R. As. Soc. 32 (1899) 35.— Alsophila 
alternans Hook. Syn. Fil. (1866) 41; Bedd. Ferns 
Br. Ind. (1867) t. 236. — Schizocaena alternans 
J. Sm. Hist. Fil. (1875) 244; Copel. Gen. Fil. 
(1947) 99.— Alsophila janseniana v. A. v. R. Bull. 
Jard. Bot. Btzg III, 5 (1922) 119.— 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 300-1 300 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 61/2 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. PI. 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 Moh. Nur 1 1269); 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. P 



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: Ramos & Edano, 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 Copel. Philip. J. Sc. 12 
(1917) Bot. 51. — C. subbipitmata Copel. ibid. 56 
(1935) 471, pi. 1.— C. Iwlttumii Copel. I.e. All, 
pi. 2. — Gymnosphaera subbipinnata Copel. Gen. 
Fil. (1947) 99. — Gymnosphaera Iwlttumii Copel. 
I.e. 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 racliis similarly scaly and hairy, scales some- 
times deciduous. Pinnae: lower ones not greatly 
reduced, largest 30^5 cm long. Largest pinnules 
4.5-6.5 cm long, 10-14 mm wide, sessile, lobed half 
way to costa; costules 31/2-4 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, Leafi. 
Philip. Bot. 4 (1911) \ \ 50.— Alsophila obliqua C. 
Chr. Ind. Fil. Suppl. 1 (1913) 5; v. A. v. R. 
Handb. Suppl. (1917) 54. — Gymnosphaera obliqua 
Copel. 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 I., 
Philippines (MICH; dupl. at K, A, BO, US, FI, 
S-PA, SYD, L). 

Distr. Malaysia: Philippines (Sibuyan I., 
once collected), at 600 m. 

161. Cyathea integra J. Sm. e.x Hook. Ic. PI. 
(1844) t. 638, incl. also var. petiolata Hook. I.e. 
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 il, n. 7 (1912) 9; Handb. Suppl. (1917) 
39; Copel. in Elmer, Leafl. Philip. Bot. 5 (1913) 
1680. — C. bulusanensis Copel. in Elmer, Leafl. 
Philip. Bot. 9 (1920) 3109; Fern Fl. Philip. 2 
(1960) 201.— C. arguta Copel. Philip. J. Sc. 38 
(1929) 133; Fern Fl. Philip. 2 (1960) 203.— 
C. breviloba Copel. Philip. J. Sc. 81 (1952) 13; 
Fern Fl. Philip. 2 (1960) 201.— Fig. 29a, b. 

Stipe 10-AO cm, base with spines 2 mm long; 
scales at base of stipe thin, pale, edges closely 
setiferous, to 25 by 1 '/2 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) 



147 




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, X 2, d. part of 
fertile pinnule showing sori and scales, x 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. tripiiuiatifida RoxB. 

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 Holttum, 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 3'/2^ 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 
buUate 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. J. Sc. 
30 (1926) 325; Fern Fl. 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, vol. P 



J. Sm. ex Hook.) v. A. v. R. Handb. Suppl. 
(1917) 25. 

Trunk: leaf-scars elliptic, 21/2 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 
racliis 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^ mm long, one lowest 
segment sometimes just free, rest of pinnule 
lobed to about IV2 mm from costa; costules 6 mm 
apart; veins to 8 pairs, mostly forked, a few with 
acroscopic braiich again forked; lamina-segments 
thin but firm, apices falcate, acu*e, 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 hairs 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 Holttum, 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; racliis light 
brown, glabrescent, sparsely and finely warty. 
Pinnae to 50 cm long, pinnules rather widely 
spaced. Pinnules sessile, to 8 by 1 '/2 cm, lobed 
2/3 towards the costa except at the very base, 
lowest segment not free; costules 4-4'/2 rnm 
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 



of C. megalosora Copel., and is intermediate in 
soral characters between C. megalosora and the 
exindusiate C. sc/uamulata (Bl.) Copel. which 
occurs at lower altitudes. 

165. Cyathea megalosora Copel. Philip. J. Sc. 12 
(1917) Bot. 54; C. Chr. Card. 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 1 V2 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: Topping 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 IOV2 by 2I/2 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; Copel. Philip. J. 
Sc. 4 (1909) Bot. 35; Fern Fl. Philip. 2 (1960) 202. 
—C. sessilipinnula Copel. 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 V2-% 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) 



149 



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 Copel. Philip. J. Sc. 6 
(191 1 ) Bot. 145; v. A. v. R. Handb. Suppl. (1917) 
30; Copel. Fern Fl. Philip. 2 (1960) 203.— C. 
pseudoalbizzia Copel. Philip. J. Sc. 38 (1929) 135; 
Fern Fl. 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; mciin 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 
3'/2 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 buUate 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 Copel. in Elmer, Leafl. 
Philip. Bot. 4 (1911) 1150; v. A. v. R. Handb. 
Suppl. (1917) 38; Copel. 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: Elmer 12513, Mt Giting- 
Giting, Sibuyan I. (MICH; dupl. at K, A, Fl, US, 
BO, P, SYD, U, L). 

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. Ill, inch var. 
mida Copel.; v. A. v. R. Handb. Suppl. (1917) 23; 
Copel. Fern Fl. 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 Copel. Philip. J. Sc. 38 (1929) 134; 
Fern Fl. Philip. 2 (1960) 204.— C. bontocensis 
Copel. Philip. J. Sc. 46 (1931) 209; Fern Fl. 
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 Vi-'^.i towards costa; cos- 
tules 31/2-51/2 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. II, vol. P 



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. 



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 ha\e 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 4'/2 rnm apart and veins 6-7 pairs; it 
was found on Mt Matang, near Kuching, at 300 
m. 

In HoLTTUM, Rev. Fl. Mai. 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. 



171. Cyathea assimilis Hook. Syn. Fil. (1865) 24; 
V. A. V. R. Handb. (1908) 20; Copel. Philip. J. Sc. 
4 (1909) Bot. 49. — C. beccahana Cesati, Atti Ac. 
Napoli 78 (1876) 3.— C. dulitensis Bak. Kew Bull. 
(1896) 40; v. A. v. R. Handb. (1908) 16; Copel. 
Philip. J. Sc. 4 (1909) 33.— C. ampla Copel. 
Philip. J. Sc. 6 ( 191 1 ) Bot. 361 ; v. A. v. R. Handb. 
Suppl. (1917) 26; non Holttum, Rev. Fl. Mai. 2 
(1954) 135.— C. stipitulata Copel. 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 4 cm 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 sometimes 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 5-7 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: Lobb 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- 



172. Cyathea trichodesma (Scort.) Copel. Philip. 
J. Sc. 4 (1909) Bot. 55. — Alsophila trichodesma 
Scort. in Bedd. J. Bot. 25 (1887) 321; Bedd. 
Handb. Suppl. (1892) 3; v. A. v. R. Handb. 
(1908) 35. — Alsophila margarethae Schroet. ex 
Christ, Ann. Jard. Bot. Btzg 20 (1905) 136; 
V. A. V. R. Handb. (1908) 33.— C. margarethae 
Corel. Philip. J. Sc. 4 (1909) Bot. 38; C. Chr. 
Card. Bull. S.S. 7 (1934) 220.— C. hurbidgei [non 
(Bak.) Copel.] Holttum, Rev.Fl.Mal.2( 1954) 124. 

Trunk slender, to 4 '2 rn. 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 2-31/2 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 4V2-5 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 m in 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 (Holtlum) 



51 



173. Cyathea wallacei (Mett. in Kuhn) Copel. 
Philip. J. Sc. 4 (1909) Bot. 4S.—AlsopliUa wallacei 
Mett. in Kuhn, Linnaea 36(1869) 153; v. A. v. R. 
Handb. (1908) 36.^Alsopliila hurhidgei Bak. J. 
Bot. 17 (1879)38; V. A. v. R. Handb. (1908)33.— 
C. biirbidgei Copel. Philip. J. Sc. 4 (1909) Bot. 55; 
non HoLTTUM, Rev. Fl. Mai. 2 (1954) 124, which 
is C trichodesma. — Gymnosphaera biirbidgei 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; costuies 3-3 '/i rnrn 
apart; lamina-segments thin, slightly crenate, si- 
nuses narrow; veins 4-6 pairs, mostly simple. 
Suri medial, lacking indusia. Scales and hairs: 
lower surface of pinna-rachis, costae, costuies 
and veins bearing pale spreading hairs 1 mm long; 
pale bullate scales present on costae and costuies; 
upper surface of costuies 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 costuies; 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 Mettemus 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.e. 362.— 
Alsophila poiensisw. A. v. R. Handb. Suppl. (1917) 
56. — Alsophila trichophora v. A. v. R. i.e. 72. — 
C. mollis Copel. Philip. J. Sc. 12 (1917) Bot. 52; 
C. Chr. Card. Bull. S.S. (1934) 220.— C. ramosii 
Copel. Philip. J. Sc. 30 (1926) 7,25.— Alsophila 
ramosii C. Chr. Ind. Fil. Suppl. 3 (1934) 23.— 
C. bipinnatifida Copel. Philip. J. Sc. 56 (1935) 97, 
pi. 2 (not C. bipinnatifida (Bak.) Domin, 1929). — 
Gymnosphaera bipinnatifida Copel. Gen. Fil. (1947) 
99. — Gymnosphaera mollis Copel. I.e. 99. — Gymno- 
sphaera trichophora Copel. I.e. 99; Fern FI. 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- 
ferous 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; costuies 3 '2^ 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 

costuies; some pale bullate scales present on cos- 
tuies. 

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. Chr. Card. Bull. S.S. 7 (1934) 219; Holttum 
Rev. Fl. Mai. 2 (1954) 122.— C. kemberangana 
Copel. Philip. J. Sc. 12 (1917) Bot. 52; C. Chr. 
Gard. Bull. S.S. 7 (1934) 219.— Alsophila kem- 
berangana C. Chr. Ind. Fil. Suppl. 3 (1934) 22. — 
Gymnosphaera dinagatensis Copel. Philip. J. Sc. 
81 (1952) 19, pi. 14; Fern Fl. Philip. 2 (1960) 235. 
— C. ampla {non Copel.) Holttum, Rev. Fl. Mai. 
2 (1954) 125. — Gymnosphaera glabra (non Bl. ) 
Copel. Fern Fl. 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 glabrescsnt 
on lower surface. Pinnae to 60 cm long, lower ones 
long-stalked, not greatly reduced. Pinnules com- 
monly 85 by 20 mm. sometimes to 1 10 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; 
costuies 4 1/2-5 '/2 (sometimes to 6 1/2) 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 costuies, 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). 

Ecol. 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 



Flora Malesiana 



[ser. II, vol. P 



he had a young frond on which bullate scales in 
some cases partly cover the immature sori. 

176. Cyathea obscura (Scort.) Copel. Philip. J. 
Sc. 4 (1909) Bot. 37; Holttum, Rev. Fl. Mai. 2 
(1954) 124. — Alsophila obscura Scort. in Bedd. J. 
Bot. 25 (1887) 321, t. 278, fig. 2; Handb. Suppl. 
(1892) 3; V. A. v. R. Handb. (1908) 34; Suppl. 
(1917) 57. — Alsophila subohscura v. A. v. R. Bull. 
Jard. Bot. Btzg II, /;. 20 (1915) 1, t. 1; Handb. 
Suppl. (1917) 57.— C. bartlettii Copel. Un. Cat. 
Publ. Bot. 14 (1929) 371.— C. pulchra Copel. 
I.e. 372.— C. subobscura Domin, Pterid. (1929) 
263. — Alsophila bartlettii C. Chr. Ind. Fil. Suppl. 
3 (1934) 20.— Alsophila pitlehra C. Chr. I.e. 23.— 
Gymnosphaera pulehra Copel. Gen. Fil. (1947) 99. 

Stipe dark to medium brown, densely scaly 
towards the base, finely warty where scales have 
fallen; scales 20-40 by 2-31/2 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 1/2-% distance to costa; costules 
31/2-4 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.n., Perak (BM; 
dupl. at K, SING). 

Distr. M(^//rtr5z'o: Sumatra and Malay Peninsula. 

Ecol. In forest, 900-1400 m. 

177. Cyathea agatheti Holttum, 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 2-21/2 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 



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-iovQsi on water-logged white 
acid sand, 600 m. 

178. Cyathea squamulata (Bl.) Copel. Philip. J. 
Sc. 4 (1909) Bot. 37; Holttum, Rev. Fl. Mai. 2 
(1954) 122, fig. 49. — Gvmnosphaera squamulata 
Bl. En. PI. Jav. (1828) 243; Copel. Gen. Fil. 
(1947) 99; Fern Fl. Philip. 2 (1960) 21,5.— Al- 
sophila squamulata Hook. Sp. Fil. 1 (1844) 51, 
p.p.; Mett. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 
52; Racib. Fl. Btzg 1 (1898) 33; v. A. v. R. 
Handb. (1908) 33; Suppl. (1917) 56.— Alsophila 
comosa Wall, ex Hook. Sp. Fil. 1 (1844) 53; 
Syn. Fil. (1865) 41; Bedd. Ferns Br. Ind. (1865) 
pi. 84; Handb. (1883) \3.— Alsophila laeta Kunze, 
Bot. Zeit. 4 (1846) 476, p.p. — Alsophila oligosora 
MiQ. Verb. 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, pi. 16, not C. 
brooksii Maxon, 1904. — C paraphysata Copel. 
I.e. 135, pi. 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) 5^.— Alsophila 
alhcota v. A. v. R. Bull. Jard. Bot. Btzg III, 5 
(1922) 180. — Alsophila glabrescens v. A. v. R. I.e. 
181. — C. deuterobrooksii Copel. Philip. J. Sc. 38 
(1929) 131.— C. alhcota Domin, Acta Bot. Bohem. 
9 (1930) 89.— C. glabrescens Domin, I.e. 119.— 
C. xanthina Domin, I.e. 172. — Alsophila xanthina 
C. Chr. 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 scaly 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 1/2--/3 
distance towards costa, no free basal segments 
but pinnules on exceptionally large pinnae some- 
times lobed nearly to costa at base; costules 31/2- 
41/2 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, have been found on Mt Kinabalu, N. Borneo, and 

the larger ones acuminate and setiferous near are small, with exceptionally scaly rachis, but do 

apices; a few stout hairs on upper surface of not appear to differ significantly in other ways, 
costules. Note. The type specimens of Alsophila comosa 

Type specimen: Kuhl & van Hasselt, Pasir and A. ridleyi are both from Singapore I. The 

Ipis, W. Java (L). latter is only distinct in smaller size and less deeply 

Distr. Malaysia: Sumatra. Malay Peninsula, lobed pinnules. Similar differences characterize 

Java, Borneo. S. Philippines (Sulu Arch.). the other species reduced to synonymy. The type 

Ecol. A small tree-fern of forest, not in open collection of C. squamulata includes a small frond 

places, in lowlands and to c. 1500 m (rarely above only 60 cm long in all, with pinnae to 8 cm long 

1000 m in the Peninsula); specimens from 2500 m and very few free pinnules, but fertile. 

Ab. Subsection Sarcopholis 

Holttum, subsect. now — Fig. 30. 

A subsectione Schizocaena differ t\ 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 t>om 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. piilcherrima 
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 179. C. fusca 

3. Bullate scales present. Pinnules lobed to within 1 mm from costa 180. C. setifera 

2. Scales on costae not setiferous 181. C. rosenstockii 

1. Pinna-rachis not hairy on lower surface. 
4. Indusium quite lacking. 
5. Pinnules almost sessile, to 28 mm wide 182. C. marginata 

5. Pinnules long-stalked, to 45 mm wide 183. C. mesosora 

4. Indusium present. 

6. Pinnules on stalks to 6 mm long. Stipe to 60 cm 184. C. papuana 

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 186. C. parvipinna 

8. Pinnules deeply lobed throughout. 

9. Pinnules not over 20 mm wide, lobed to c. 3 mm from costa 187. C. werneri 

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 tYinged 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 



154 



Flora Malesiana 



[ser. 11, vol. 12 



179. Cyathea fusca Bak. in Beccari, Malesia 3 
(1886) 31; V. A. v. R. Handb. (1908) 19; Suppl. 
(1917) 25; CoPEL. 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); rac/iis 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. V2 rnm 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. 




Fig. 30. Cyathea rosenstockii Brause. Base of 

stipe, showing fleshy scales; scale in cm (Pulle 

509, Mt Perameles, 1100 m, BM). 



180. Cyathea setifera Holttum, Kew Bull. 16 
(1962) 62. 

Stipe 10 cm, covered with thick dark pale-edged 
scales 20 by 1 1/2 rnrn, 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 
4 cm 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 4-4' 2 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 lower 
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 I on a costule). 

Type specimen: Main & Aden 1306, Morotai, 
N. Moluccas (BO; dupl. at K, L). 

Distr. Malaysia: Moluccas (Morotai), one 
collection. 

Ecol. At 1000 m. 

181. Cyathea rosenstockii Brause, Bot. Jahrb. 56 
(1920) 49; Copel. Philip. J. Sc. 77 (1947) 101.— 
Fig. 30. 

Trunk 1 i/i-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 5-51/2 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] 



Cyatheaceae (Holttum) 



155 



182. Cyathea marginata (Brausu) Domin, Acta 
Bot. Bohem. 9 (1930) \?>4.^A/soplii/(i marginata 
Brausf., Bot. Jahrb. 56 (1920) 63. 

Trunk to 3 m tail; 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. Sort 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 Holttum, 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 1 '^ 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 1 10 by 45 mm, lowest on stalks to 8 mm 
long, lobed throughout to ?>-4- mm from costa; 
costules 71/2-9 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; Copel. Philip. J. Sc. 77 (1947) 
121. — Alsophila papuana Ridl. Trans. Linn. Soc. 
II, 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 



and raised on lower surface. 5o/-/ 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: Boden Kloss, Mt Carstensz, 
Camp III, Jan. 1913, W. New Guinea (BM; dupl. 
at K). 

Distr. Malaysia: West New Guinea (two 
collections). 

Ecol. At 700-1100 m. 

185. Cyathea inaequalis Holttum, Kew Bull. 16 
(1962) 56. 

Trunk 4-6 m, 5-6 cm 0; leaf-scars in alternate 
whorls of 5. Stipe 24 cm, copiously thorny and 
scaly near the base; thorns 1 mm, acute; scales 
ascending, fleshy at their bases and thinner distally, 
to 20 by 1 1/2 iTiiTi> 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 
71/2-8 1/2 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 (once 
collected). 

Ecol. In rain forest ravine, at 700 m. 

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 31/2 by 1 cm, 
on stalks 8 mm long, lamina simple; largest 40 cm 
long. Pinnules to 90 by 1 8 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. II, vol. 12 



surface of costae and none on upper surface of 

costules. 

Type specimen: Brass 25837, Normanby I., 
Papua (L; dupl. at K, USj. 

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; 
Copel. Philip. J. Sc. 77 (1947) 100. 

Trunk to 3^/2 m, bearing c. 8 fronds to 2'/2 m 
long. Stipe 8-25 cm; scales to 20 by 1 1/2 rnm, 
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 
5'/2-6V2 rnrn 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 (/.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 Holttum, Kew Bull. 16 

(1962) 57. 

Trunk 3-5 m, 3'/2-6 cm after decay of leaf- 
bases; leaf-scars 18-25 mm 0, 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 Roxb. Calc. J. Nat. 
Hist. 4 (1844) 518.— C. Integra J. Sm. ex Hook. 
Ic. PI. (1844) t. 638, p. /5. excl. ic. — C. nigrospinulosa 
V. A. V. R. Bull. Jard. Bot. Btzgll,/?. 28 (1918) 15. 

Stipe to 25 cm (sometimes much shorter ?), 
bearing scattered thick scales to 20 by 1 V2 mm, 
setiferous near their apices; stipe also covered with 
very small dull fringed scales. Pinnae to at least 
55 cm long. Largest pinnules 90-1 10 by 20-28 mm, 
lobed to 1 '/i-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: lower 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 /;. 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. Dep. Agr. 

Ind. Neerl. /;. 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) 



157 



Dist r.A/a/a>'j/a: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 1 It rnm, 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 buUate 
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 Mett., 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. n. 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 Christ, J. Linn. Soc. Bot. 39 
(1909) 213.— C. matthewii Domin, 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 DoNUN, 
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 Christ, Bull. Herb. Boiss. 
II, 6 (1906) 1008; v. A. v. R. Handb. (1908) 785. 
I have not found the type specimen at Paris. 
Christ 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 Domin, 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 tntncata 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 Desv. Mem. Soc. Linn. Paris 6 
(1827) 323. — Polvpodiiim arboreum Lour. F1. 
Coch. (1790) 831, non Linn. 

Desvaux published a new name for Loureiro's 
species, without description; like Loureiro, he 
also cited a plate by Rumphius. Loureiro described 
a fern from Indo-China, not Ambon; neither 
his description, nor Rumphius'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, 
57). 

Excluded from Cyathea 

Alsophila dielsii Brause. Bot. Jahrb. 56 (1920) 67. 
— C. dielsii Domin, Acta Bot. Bohem. 9 (1930) 111. 
The type specimen belongs to the genus Di- 
plazium. 



158 Flora Malesiana [ser. II, vol. P 

Alsophila dryopteroidea Brause, Bot. Jahrb. 56 Alsophila warburgii Christ, Ann. Jard. Bot. Btzg 

(1920) 70.— C. atrispora Domin, Acta Bot. 15 (1897) 80, from Celebes = Dryopteris sp. 
Bohem. 9 (1930) 95. — Dryopteris atrispora C. 

Chr. Brittonia 2 (1937) 296.— Lastrea dryopteroi- Alsophila xantlwlepis Christ ex Diels in E & P 

dea COPEL. Gen. Fil. (1947) 138. Pfl. Earn. 1, 4( 1899) 138; v. A. v. R. Handb. (1908) 

The type is a Thelypteroid fern; generic limits u.—C. xantlwlepis Domin, Pterid. (1929) 263- 

m this group need to be clarified. Acta Bot. Bohem. 9 (1930) 172 = Dryopteris sp. 

Alsophila gazellae KuHN, Forschungsr. Gazelle 4 ^ , •.- ., /r, . r, „ ., 

(1889) 13.-C.^«ze'//a^ Domin, Pterid. (1929) 262. ,? " «'if ! ^^nt' ^^^;,' ^°'-'- ^ '^or. Syst. 

I have seen the type specimen, and identified it ^^'^- ^'^^^^ ^^^ = ^'^'^^'^P^ asp,d,oides Bl. 
as Fleocnemia cumingiana Presl (see Reinwardtia 

1, 1951, 188). Cyathea woodlarkensis Copel. Philip. J. Sc. 9 

(1914) Bot. 1; ibid. 77 (1947) MA.— Alsophila 

Alsophila tristis (Bl.) Bl. ex Moore, Ind. Eil. woodlarkensis C. Chr. Ind. Eil. Suppl. 2 (1917) 4. 

(1857) 58, based on Aspidium triste Bl. En. PI. This is a large species of r/?e/v/i/m5, apparently 

Jav. (1828) 169 = Stenolepia tristis (Bl.) v. A. near T. immersa (Bl.) Ching. There are speci- 

V. R. Bull. Dep. Agr. Ind. Neerl. n. 11 (1909) 45. mens in Herb. Copel. (MICH) and at Paris. 

2. DICKSONIA 

l'Herit. Sert. Angl. (1788) 30, p.p.; Christ, Farnkr. Erde (1897) 313, p.p.; 
DiELS in E. & P. Pfl. Fam. 1, 4 (1899) 1 19; Underw. Mem. Torr. Bot. CI. 6 (1899) 
259, 278; C. Chr. Ind. Fil. (1905) xvi, 220; Maxon, J. Wash. Ac. Sc. 12 (1922) 
454; Copel. Gen. Fil. (1947) 4S.—Ba/antium Kaulf. En. Fil. Chamisso (1824) 
2SS, p.p. typ.; Presl, Tent. Pterid. (1836) 134 (e.xcl. Balantium brownianum).— 
Dicksonia subg. Balantium Hook. Sp. Fil. 1 (1844) 66,p.p.—Dicksonia § Eudickso- 
nia Hook. & Bak. Syn. Fil. (1868) 50, /7./7.— 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; 50/7 each 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 l'Herit. (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 



Dec. 1963] 



Cyatheaceae (Holttum) 



159 



n = 65 (New Phytol. 56, 1957, 207). Manton found the same number for D. arhuresceiis (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, I.e. 
Earlier authors confused not only Dicksonia, Culcita and Cibutiitnu but also Deniistaedtia 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 Cyat/iea; but the differences 
between one hair and another are less easily defined than those between different scales. The species in 
New Guinea are particularly difTicult to discriminate, and the present account can only be regarded as 
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 surface of pinna-rachis (in some cases 
also of costa) short, pale, flaccid, their lateral walls collapsed. Spores verrucose. 

3. Hairs on costae almost all flaccid, pale, zr 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 3. D. sciurus 

4. Hairs on costae ascending, mostly dark red 4. D. archboldil 

1. Base of stipe clothed mainly with soft matted red-brown hairs, with or 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 ^* ^' S''^"'*'^ 

6. Pinna-rachis brown, persistently hairy beneath; costae bearing entirely pale flaccid hairs. 

7. D. lanigera 



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- 
lanriiim chrysotriehum Hassk. Obs. Fil. Jav. 1 
(1856) 53. — D. ehnsotrieha Moore, Ind. Fil. 
(1860) 190; Hook. & Bak. Syn. Fil. (1868) 50; 
Racib. F1. Btzg 1 (1898) 121; Diets in E. & P. 
Pfl. Fam. 1,4 (1899) 121; Christ, 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 



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.— £). blumei [non (Kunze) Moore] C. Chr. 
Card. Bull. S.S. 7 (1934) 223; Copel. Fern Fl. 
Philip. 1 (1958) 84.— Fig. 31d-€. 

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 
(?), Philippines (Mindanao to S. Luzon). 

Ecol. Mountain forests, 1500-2000 m. The 
Celebes specimen, much smaller than those from 



160 



Flora Malesiana 



[ser. II, vol. P 




Fig. 31. Dicksonia bliimei (Kunze) Moore, a. Part of pinna, showing sterile and fertile pinnules, X 2/3, 
b. part of pinnule, lower surface, showing sori and rigid hairs, >: 6, c. section of sorus, x 8. — D. mollis 
HoLTTUM. d. Part of sterile pinnule, lower surface, showing flaccid hairs, < 6, e. hairs from lower surface, 
X 30. — D. antarctica Labill. /. Section of stipe (hairs omitted), x 2, g. section of pinna-rachis, X 4 
{a DE Vriese s.n., b-c Matthew s.n , c/ Sinclair 9005, e Elmer 9874, /-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. Chr. Brittonia 2 (1937) 
283; CoPEL. 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 1 50 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 Copel. 
(1949) 6, pi. 1. 



Philip. J. Sc. 78 



Dec. 1963] 



Cyatheaceae (Holttum) 



161 



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 cni 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 tertiary 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 (Sleu- 
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. sciiiriis C. Chr. (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. Chr. Brittonia 2 (1937) 282; 
CoPEL. 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. 



6. Dicksonia grandis Rosenst. in Fedde, Rep. 5 
(1908) 34; Corel. 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. n. 50 (B; dupl. at 
L, P, US, UC). 

Distr. Malaysia: NewGuinea (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 Holttum, 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; lamina 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 51/2 rnm apart; costa beneath brown-hairy 
almost to apex; tertiary 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: Pulle 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 Bak. J. Linn. Soc. Bot. 22 
(1886) 223 = Dennstaedtia ampla (Bak.) Bedd. 

Dicksonia cuneata Hook. Sp. Fil. I (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 Christ, Ann. Jard. Bot. 
Btzg 15 (1897) 86 = Demistaedtia erythrorachis 
(Christ) Diels. 

Dicksonia flaccida (Forst.) Sw. in Schrader, J. 
Bot. 1800, ii (1801) 90 -= Demistaedtia flaccida 
(Forst.) Bernh. 

Dicksonia glabra ta Ces. Rend. Ac. Napoli 16 
(1877) 24, 28 = Dennstaedtia glabrata (Ces.) 
C. Chr. 

Dicksonia gomphophyUa Bak. J. Linn. Soc. Bot. 
22 (1886) 223 = Dennstaedtia gomphophyUa 
(Bak.) C. Chr. 

Dicksonia japonica Sw. in Schrader, J. Bot. 1 800, 
ii (1801) 92 = Microlepia strigosa (Thunb.) 
Presl. 

Dicksonia kingii Bedd. Handb. Suppl. (1892) 
6 = Orthiopteris kingii (Bedd.) Holttum. 

Dicksonia linearis Cav. Descr. (1802) 274 = 
Tapeinidium pimiatum (Cav.) C. Chr. 

Dicksonia moluccana Bl. En. Pi. Jav. (1828) 
239 = Dennstaedtia moluccana (Bl.) Moore. 



Dicksonia neglecta Fee, Gen. Fil. (1850-52) 
335 = Dennstaedtia smithii (Hook.) Moore. 

Dicksonia nephrolepioides Christ, Verh. Nat. 
Ges. Basel 11 (1895)241 = Nephrolepis dicksonioi- 
des Christ. 

Dicksonia remota Christ, Verh. Nat. Ges. 
Basel 11 (1896) 423 = Dennstaedtia remota 
(Christ) Diels. 

Dicksonia rhombifolia Bak. J. Bot. 28 (1890) 
105 = Dennstaedtia rhombifolia (Bak.) C. Chr. 

Dicksonia scabra Wall, ex Hook. Sp. Fil. 1 
(1844) 80 = Dennstaedtia scabra (Wall.) Moore. 

Dicksonia scandens Bl. En. PI. Jav. (1828) 
240 = Dennstaedtia scandens (Bl.) Moore. 

Dicksonia smithii Hook. Sp. Fil. 1 (1844) 80 = 
Dennstaedtia smithii (Hook.) Moore. 

Dicksonia strigosa Thunb. Trans. Linn. Soc. 2 
(1794) 341 = Microlepia strigosa (Thunb.) 
Presl. 

Dicksonia zippeliana KuNZE, Bot. Zeit. 3 (1845) 
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 />//7««-rachis antrorsely hairy and grooved, the groove not open 
to admit the grooves of midribs of pinnules; pinnules shghtly 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 {I.e. 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) 



163 




Fig. 32. Cystodium sorbifoliiim (Sm.) J. Sm. a. Middle part of rachis with pinnae, X y^, 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, ■: 4, d. section of pinna-rachis, < 4 (C. Hose 217). 



1. Cystodium sorbifolium (Sm.) J. Sm. in Hook. 
Gen. Fil. (1841) t. 96; Copel. Sarawak Mus. J. 2 
(1917) 344; C. Chr. Card. Bull. S.S. 7 (1934) 
239; Ind. Fil. Suppl. 3 (1934) 66; Copel. Philip. 
J. Sc. 78 (1950) l.—Dicksonia sorbifolia Sm. in 
Rees, Cyclop. 11 (1808) n.p.; Hook. Sp. Fil. 1 
(1844) 72, t. 25A; Hook. & Bak. Syn. Fil. (1868) 
52; Christ, Ann. Jard. Bot. Btzg 15 (1898) 87; 
V. A. V. R. Handb. Suppl. (1917) lA.—Dicksonia 



mohtccana RoxB. Calc. J. Nat. Hist. 4 (1844) 517. 
— Dicksonia papuana F. v. M. Descr. Not. 4 
(1876) 76; Copel. Philip. J. Sc. 6 (191 1 ) Bot. 69.— 
Saccoloma sorbifolia CHKiST.YsiTnkx. Erde (1897) 
309; C. Chr. Ind. Fil. (1905) 612; v. A. v. R. 
Handb. (1908) 282.— Fig. 32. 

Stem usually creeping (once reported erect, 
'/2 ni high), densely covered like the bases of the 
stipes with soft golden brown shining hairs c. 3 cm 



164 Flora Malesiana [ser. II, vol. 1^ 

long. Stipes to 75 cm, glabrescent and smooth abundant spreading hairs 1 mm long. Sort 1 1/2-2 

above the base: lamina 100-250 cm long, lowest mm apart, at ends of all veins on fertile pinnules 

pinnae not greatly reduced; largest pinnae 20-30 (on acroscopic branch if vein is forked). Spores 

by 5-9 cm, bearing free pinnules for Vi-Vi of bearing on their surface a fine irregular slightly 

their length, apical part deeply lobed; sterile raised reticulum. 

pinnules 6-8 mm wide, fertile 5-6 mm wide, sep- Type specimen: C. Smith, Honimoa, Ceram 
arated by gaps of c. 2 mm: pinnules slightly (Herb. J. E. Smith; dupl. at K). 
oblique, their costules falcate, asymmetric at the Distr. Malaysia: N. Borneo, N. Celebes, Mo- 
base (rounded to cordate on basiscopic side, luccas (Ceram), New Guinea (incl. New Britain, 
broadly truncate and sometimes auricled on aero- Admiralty Is and Louisiade Arch.), 
scopic side), edges almost parallel except towards Ecol. In lowland forest, to 400 m, often near 
apex; lower surface of pinna-rachis, costules of rivers, 
pinnae and sometimes veins bearing more or less 

4. CIBOTIUM 

Kaulfuss, Berl. Jahrb. Pharm. 21 (1820) 53; En. Fil. Chamisso (1824) 229; 
Presl, Tent. Pterid. (1836) 67, excl. syn. Deparia; Hook. Gen. Fil. (1839) t. 25; 
Sp. Fil. 1 (1844) 82; Diels in E. & P. Pfl. Earn. 1, 4 (1899) 121; C. Chr. Ind. 
Fil. (1905) xvi, 183; Maxon, Contr. U. S. Nat. Herb. 16 (1912) 54; Corel. Gen. 
Fil. (1947) 49.— Pinonia Gaudich. Ann. Sc. Nat. 3 (1824) 507.— Dicksonia § 
Cibotium Hook. & Bak. Syn. Fil. (1868) 49.— Fig. 33. 

Caudex massive, erect or prostrate (Malaysian species mostly the latter), the 
apex protected by a thick cover of long slender hairs. Stipes always long, smooth, 
covered with hairs at the base or throughout. Fronds large, bipinnate, lowest 
pinnae not greatly reduced, all axes more or less hairy, or in some cases glabres- 
cent; pinnules deeply pinnatifid throughout, the lowest lobes sometimes free as 
tertiary leaflets, lobes each with a costule bearing simple or forked lateral veins; 
costae of pinnules raised (not grooved) on upper surface; fertile pinnules not 
different in shape or size from sterile. Sori at the ends of veins, protected by 
two indusia which are alike in texture and different from the green lamina of the 
lobes on which they are borne, the outer indusium deflexed so that the sorus ap- 
pears to be on the under side of the lobe, the inner indusium at maturity bending 
back towards the costule and elongating, usually becoming oblong, the two indusia 
joined together for a short distance at the base, thus forming a small cup round 
the receptacle of the sorus; receptacle somewhat prominent, elongate obliquely 
to the end of the vein which bears it, bearing numerous sporangia and long 
paraphyses; sporangia similar to those of Dicksonia; spores trilete, bearing few 
strongly raised ridges on the outer surface. 

Type species: Cibotium chamissoi Kaulf., Hawaii. 

Distr. About 12 spp., distributed in Central America and Mexico, Hawaii, Assam to southern China, 
southwards to Western Malaysia and Philippines. 

KEY TO THE SPECIES 

1. Sori 2 or more pairs on each pinnule-lobe of larger fronds. Largest pinnules 20-35 mm wide; pinnules 
on the two sides of a pinna not greatly different in length. Hairs on lower surface of costae and costules 

almost always thin and flaccid, never spreading 1. C. barometz 

1. Sori never more than 2 pairs on each pinnule-lobe. Largest pinnules 15-25 mm wide; pinnules on 
basiscopic side of lower pinnae much shorter than those on acroscopic side. Some rigid hairs usually 
present with flaccid ones on lower surface of costae, the rigid hairs sometimes spreading. 
2. Always one pair of sori. Spreading hairs usually present on lower surface of costae; hairs normally 

absent on lower surface of lamina between veins 2. C. cumingii 

2. Always two pairs of sori on large fronds. Spreading hairs lacking, but rigid (often red) appressed 
hairs always present and sometimes abundant; small flaccid hairs present on lower surface of lamina 
between veins 3. C. arachnoideum 



Dec. 1963] 



Cyatheaceae (Holttum) 



165 



1. Cibotium barometz (L.) J. Sm. Lond. J. Bot. 1 

(1842) 437; Bhdd. Handb. (1883) 24; Suppl. 
(1892) 6; DiELS in E. & P. Pfl. Fam. 1, 4 (1899) 
121; Christ, Philip. J. Sc. 2 (1907) Bot. 117, 
incl. var. sumatramiin Christ, I.e. I 18; v. A. v. R. 
Handb. (1908) 48, 792, p.p.; Suppl. (1917) 77; 
Heyne, Nutt. PI. (1927) 103; Burk. Diet. (1935) 
536; Backer & Posth. Varenfl. Java (1938) 23, 
f. 7; Tard.-Blot & C. Chr. F1. Gen. I.-C. 7, 2 
(1939) 78, f. 10; Holttum, Rev. Fl. Mai. 2 (1954) 
1 14, f. 45; A. F. Tryon, Miss. Bot. Card. Bull. 43 
(1955) /;. 2; Am. Fern J. 47 (1957) 1; Ching, Fl. 
Rep. Pop. Sin. 2 (1959) \91 .—Polypodium ba- 
rometz LiNNE, Sp. PI. (1753) 1092; Lour. Fl. 
Cochin. (1790) 675. — Aspidiiim barometz Willd. 
Sp. PI. 5 (1810) 268.^Ba/antii<m gkiueeseens 
Link, Fil. Sp. Hort. Berol. (1841) 40.— Dieksonia 
baranetz Link, I.e. 166; Hook. & Bak. Syn. Fil. 
(1868) 49 (barometz). — C. gkiueeseens KuNZE, 
Farrnkr. 1 (1841) 63, t. 31; Hook. Sp. Fil. 1 
(1844) 82.— C. assamicum Hook. Sp. Fil. 1 (1844) 
83, t. 29B; Christ, Philip. J. Sc. 2 (1907) Bot. 117. 
— Dieksonia assamieiim Griff. Notul. 2 (1849) 
607; Ic. PI. As. 2 (1849) t. 136, f. 2.— C. djambia- 



mtm Hassk. Obs. Fil. Jav. 1 (1856) 61.— C. 
gkiuctim [noil (Sm.) Hook. & Arn.] Bedd. Ferns 
Br. Ind. (1865) t. 83.— Fig. 33a-c. 

Caiidex usually prostrate; hairs near apex 
shining, brown, to 4 cm or more long. Stipes to at 
least 120 cm, base densely hairy, rest smooth when 
old, softly short-hairy when young. Lamina to 
200 cm long; longest pinnae 80 cm long. Largest 
pinnules 150 by 20-35 mm, lobed almost to the 
costa, often with 1-2 pairs of free tertiary leaflets at 
the base; costules ^Vi-^Vi mm apart; segments of 
lamina glaucous beneath, edges where sterile 
crenate-serrate; lower surface of costae and cos- 
tules more or less densely covered with pale 
entangled flaccid appressed hairs (young plants 
softly hairy throughout); veins oblique, usually 
forked, in largest pinnules sometimes twice forked. 
Sori 2-A or more pairs on largest fronds, at the 
ends of lower veins on each lobe of the lamina, 
soriferous veins usually unbranched; receptacle 
rather oblique to the end of the vein so that it is 
parallel to the edge of the lobe; outer indusium 
permanently round, inner elongating at maturity 
and more or less oblong. 




Fig. 33. Cibotium barometz (L.) J. Sm. a. Part of pinna-rachis with pinnules, x 2/3, b. part of pinnule, 
upper surface, showing venation (unbranched basal veins lead to sori) x 6, c. part of pinnule, lower 
surface, showing sori; 2 sori covered by indusia, 2 with inner indusium reflexed and extended, < 6. — 
C. regale Linden, d. Section of stipe, nat. size, e. section of pinna-rachis, nat. size {a-c Curtis 3103, 

d-e cult. R. B. G. Kew). 



166 



Flora Malesiana 



[ser. II, vol. P 



Type specimen: none in Linnean Herbarium. 
Linnaeus stated tiiat he had a specimen from 
China, and the identity is not in doubt. 

Distr. NE. India to S. China and Formosa, 
southwards to Malaysia, in Malaysia: Sumatra, 
Malay Peninsula, and Java. 

Ecol. Amongst non-calcareous rocks and on 
steep ground in mountain forest, from near sea- 
level to 1600 m, in the Malay Peninsula and Su- 
matra; in Java only known from Mt Slamat at 
1600 m. I have seen young plants of this species 
growing on a bare earth bank in a rather open 
place, above a path recently cut through forest on 
a mountain slope; perhaps in nature the species 
chiefly spreads by the establishment of new plants 
on land-slides. A specimen from Eraser's Hill 
(E. Smith 802) bears the note "frond 4 m, [incl.] 
stipe 2 m". Young plants are soft-hairy all over. 

Note. The name barometz was derived t>om 
the story of the vegetable lamb of Tartary (see 
A. E. Tryon, I.e.); but whatever plant gave rise to 
the strange fable of the vegetable lamb, it was not 
Cibotium. Link, Kunze, and some later botanists 
have noted that baranetz (from the Russian baran, 
a lamb; baranets, a diminutive form) is a more 
correct spelling of the name of the vegetable lamb 
than barometz; but the latter spelling was that 
adopted by Linnaeus. 

Uses. The hairs on the rhizome of this species 
of Cibotium (probably also of other species) have 
long been used in China as a styptic for bleeding 
wounds, and they have also been so used by va- 
rious peoples in Malaysia; for details, see Heyne 
and BuRKiLL. The common trade-name for these 
hairs seems to be Fenawar (D)jambi. Cibotium 
hairs have also been used for stuffing cushions 
and for upholstery, but it is reported that they are 
not very satisfactory for this purpose. 

2. Cibotium cumingii Kunze, Earrnkr. 1 (1841) 
64, 65; Christ, Philip. J. Sc. 2 (1907) Bot. 118; 
CoPEL. Eern El. Philip. 1 (1958) 84.— C. glauciim 



[nan (Sm.) Hook. & Arn.] J. Sm. in Hook. J. Bot. 

3 (1841 ) 418. — C. baranetz var. cumingii v. A. v. R. 
Handb. Suppl. (1917) 77. — C crassinerve Rosen- 
ST. Med. Rijksherb. /;. 31 (1917) 4. 

Habit as in C. barometz, but fronds probably 
never quite so large; pinnules to 180 mm long, 
rarely over 20 mm wide, those on basiscopic 
side towards base of a pinna much shorter than 
those on acroscopic side (often less than half as 
long), distal pinnules and those of upper pinnae 
more equal; sori always one pair at the base of 
each fertile segment of a pinnule; hairiness of 
lower surface of costae and costules variable, 
usually some stiff" spreading hairs (pale or reddish) 
present along with appressed flaccid hairs, in 
some cases either the one or the other kind pre- 
dominating. 

Type specimen: Cuming 123, Luzon (K; dupl. 
at P, US). 

Distr. Malaysia: Philippines (Luzon, Mindoro, 
Mindanao). 

3. Cibotium arachnoideum (C. Chr.) Holttum, 
comb, now — C. cumingii var. arachnoideum C. 
Chr. Card. Bull. S.S. 7 (1934) 224. 

Like C. cumingii in asymmetry of the pinnae, but 
larger; pinnules to 200 by 25 mm; sori one or two 
pairs to each lamina-segment; many appressed 
antrorse hairs on lower surface of costae, some of 
them rigid and usually reddish but never spreading; 
pale flaccid hairs also on costules and veins, and 
often on the surface of the lamina between veins. 

Type specimen: Holttum 25378, Mt Kinabalu, 
N. Borneo (K; dupl. at BM, SING). 

Distr. Malaysia: Central and South Sumatra, 
Sarawak, and N. Borneo. 

Ecol. On Mt Kinabalu abundant in secondary 
forest at c. 1000 m, on steep slopes which are 
periodically cleared by burning for cultivation, 
apparently surviving the burning; fronds were 
reported as up to 4 or 5 m in total height. Sumatran 
specimens were found at 1200-2000 m. 



5. CULCITA 



Presl, Tent. Pterid. (1836) 135; Hook. Gen. Fil. (1840) t. 60A; Maxon, J. Wash. 
Ac. Sc. 12 (1922) 454; C. Chr. Ind. Fil. Suppl. 3 (1934) 5, 57; Copel. Gen. 
Fil. (1947) 49.— Balantium Kaule. En. Fil. (1824) 228, p.p.; J. Sm. Hist. Fil. 
(1875) 257; Diels in E. & P. Pfl. Fam. 1, 4 (1899) 119; C. Chr. Ind. Fil. (1905) 
xvi, 148; Copel. Philip. J. Sc. 3 (1908) Bot. 30i. —Dicksonia subg. Balantium 
Hook. Sp. Fil. 1 (1844) 66, p.p.— Dicksonia § Eudicksonia Hook. & Bak. Syn. 
Fil. (1868) 50, ;?./?.— Fig. 34. 

Caudex massive, prostrate or erect, the young parts covered with hairs. Stipes 
long in proportion to lamina, more or less hairy throughout; pneumathodes 
prominent and separate {subg. Culcita) or forming a not quite continuous line 
along each side (subg. Calochlaena, fig. 34d). Lamina of large fronds quadripin- 
nate, deltoid, pinnae and pinnules also deltoid, all branches of the frond asymmet- 
ric at the base, acroscopic sub-branch longer and at a broader angle than basi- 
scopic; all axes and midribs of leaflets grooved on adaxial surface, the groove on a 



Dec. 1963] Cyatheaceae (Holttum) 167 

larger axis open to admit grooves of smaller ones borne upon it (fig. 34c). Sori at 
the ends of veins, receptacle elongate at right angles to the vein-end, protected by 
a small reflexed lobe of the lamina (the outer indusium) and by a thinner inner 
indusium attached to the base of the receptacle; sporangia about as in Dick- 
sonia, accompanied by many hair-like paraphyses; spores trilete, outer surface 
verrucose (Malaysian species). 
Type species: Culcita macrocarpa Presl (based on Dicksonia culcita CHerit.). 

Distr. Subg. Culcita: Madeira and Azores, tropical America. Suhg. Calochlaena: Australia, New 
Caledonia, New Hebrides, Fiji, Samoa, Solomon Islands, Malaysia; in Malaysia: Java, Lesser Sunda Is, 
Borneo, N. Celebes, Philippines, New Guinea. The species C. formosae (Chr.) Maxon, of Formosa, 
appears to be a Deiuistaedtia, as originally placed by Christ. 

Nomencl. The genus Balantium Kaulf. was founded on two species, B. aiiricomiim (a synonym of 
Dicksonia arhorescens, type species of Dicksonia) and B. culcita. The description was almost entirely based 
on the former, which was figured, and the sori of the latter were not seen by Kaulfuss. Maxon (I.e. 
1922) therefore argued that Balantium Kaulf. should be typified by B. auricomum and so regarded as a 
synonym of Dicksonia. The generic distinction between the two species of Kaulfuss was recognized by 
Presl, who founded a new genus Culcita for B. culcita, but left B. auricomum in Balantium, which he 
construed almost as we now construe Dicksonia; under Dicksonia he placed mostly species now included 
in Dennstaedtia. 

Taxon. The genus Culcita, as first fully assembled by Maxon, consists of two distinct parts, siibg. 
Culcita and subg. Calochlaena Maxon. The former includes only two very closely related species, C. 
macrocarpa Pr. and C. coniifolia (Hook.) Maxon; subg. Calochlaena includes several species in a quite 
different geographic area which includes part of Malaysia. The species of this subgenus have much 
smaller sori than those of subg. Culcita, and were not regarded as part of the genus until Diels (1899) 
included C. straminea (Labill.) and Copeland (1908, 1909) C. dubia (R. Br.) (fig. 34d-e) and C. 
javanica (Bl.) in it under the name of Balantium. John Smith (1875), who recognized Balantium as 
distinct from Dicksonia, had placed C. straminea in Dennstaedtia. C. dubia remained doubtfully in its 
original genus Davallia, and C. javanica doubtfully in Dennstaedtia, in Christensen's Index Filicum 
(1905). 

The genus Culcita diff'ers strikingly from Dicksonia in the shape of the frond, which always has a long 
stipe and is more finely divided, with all major parts triangular in outline and asymmetric at the base. 
It diff'ers also from Dicksonia in the grooves of the upper surface of all axes, including midribs of leaflets, 
being open to receive grooves of minor axes. In both these characters, Culcita agrees with Thyrsopteris. 
The difference in rachis-characters between Dicksonia and Culcita is exactly that between Ctenitis and 
Dryopteris. 

Culcita differs from Dennstaedtia in its more massive rhizome which is sometimes erect and when creep- 
ing has a radially symmetrical vascular system (dorsiventral in Dennstaedtia) with overlapping leaf- 
gaps; also in the inner indusium not being appreciably joined to the outer along its sides (the two are 
partly or almost wholly united in Dennstaedtia, forming a cup or funnel) but joined to the receptacle 
of the sorus on its basiscopic side as in Dicksonia (the receptacle is free or columnar in Dennstaedtia). 
As regards rachis-characters, species like both Dicksonia and Culcita are at present placed in Dennstaedtia 
(see Tryon, Contr. Gray Herb. //. 187, 1960, who makes this distinction the main division in his key to 
American species of Dennstaedtia). 

Subg. Calochlaena, to which Malaysian species belong, might possibly rank as a separate genus, but 
it appears to be much more nearly related to subg. Culcita than to any other ferns. Apart from size of 
sorus, subg. Culcita differs from subg. Calochlaena in the shape and size of pneumathodes, and in the shape 
of the vascular strands as seen in transverse section near the base of the stipe; in the latter character 
subg. Calochlaena conforms closely to the general scheme of Dicksonia and Cyathea, whereas in subg. 
Culcita the vascular strands on the adaxial side diverge instead of converging and curving inwards. 

Cytol. Manton reported n = 66-68 for C. macrocarpa (J. Linn. Soc. Bot. 56, 1958, 84), and has 
found n = approx. 55 for C. dubia in cultivation at Kew (unpublished). 

KEY to the species 

1. Inner indusium smaller than outer; midribs of quaternary leaflets rather closely hairy on lower surface. 
Mountain plants. 

2. Largest quinary lobes commonly with only one sorus 1. C. javanica 

2. Largest quinary lobes commonly with 3 or 2 sori 2. C. vUlosa 

1. Inner indusium at maturity larger than outer, pouch-shaped and not reflexed; midribs of quaternary 
leaflets sparsely hairy. Lowland plants 3. C. straminea 



168 



Flora Malesiana 



[ser. II, vol. P 




Fig. 34. Ciilcita jaxanica (Bl.) Maxon. a. Part of pinna-rachis and complete pinnule, upper surface, < 2, 

b. lower surface of leaflet showing sori, c. upper surface of part of pinnule, showing shape of rachis- 

branches, x 6. — C dubia (R. Br.) Maxon. d. Section of stipe, ■: 2, e. section of pinna-rachis, X 4 

{a-c Matthew 617, d-e cult. R. B. G. Kew). 



1. Culcita javanica (Bl.) Maxon, J. Wash. Ac. 
Sc. 12 (1922) 456; Backer & Posth. Varenfl. 
Java (1939) 22. — Dicksonia javanica Bl. En. PI. 
Jav. (1828) 240; Hook. Sp. Eil. 1 (1844) 79; C. 
Chr. Ind. Fil. (1905) 222. — Dennstaedtia javanica 
Christ, Bull. Herb. Boiss. II, 4 (1904) 617; 
V. A. V. R. Handb. (1908) \43.— Dicksonia cope- 
landii Christ, Philip. J. Sc. 2 (1907) Bot. 183.— 
Balantium cope/andii Christ in Copel. Philip. J. 
Sc. 3 (1908) Bot. 301; ibid. 4 (1909) 62, t. 19; 
Copel. in Elmer, Leafl. Philip. Bot. 2 (1908) 395; 
V. A. V. R. Handb. Suppl. (1917) 16.— Balantium 
javanicum Copel. Philip. J.Sc. 4 (1909) Bot. 62; 
V. A. V. R. Handb. Suppl. (1917) 15.— Balantium 
pilosum Copel. J. Str. Br. R. As. Soc. n. 63 (1912) 



71; Sarawak Mus. J. 2 (1917) 335; v. A. v. R. 
Handb. Suppl. (1917) 76. — Dennstaedtia paraphy- 
satav.A.w. R. Bull. Jard. Bot. Btzgll,//. 16(1914) 
7; Handb. Suppl. (1917) \2S.— Dennstaedtia 
tnultifida v. A. v. R. Bull. Jard. Bot. Btzg II, n. 20 
(1915) 10; Handb. Suppl. (1917) 129.— C. cope- 
landii Maxon, J. Wash. Ac. Sc. 12 (1922) 457; 
C. Chr. Ind. Fil. Suppl. 3 (1934) 57; Card. Bull. 
S.S. 7 (1934) 224; Copel. Fern Fl. Philip. 1 (1958) 
86.— C. pilosa C. Chr. Ind. Fil. Suppl. 3 (1934) 
57. — Fig. 34a~c. 

Caude.x ascending or erect (to 60 cm high, 
fide CoPELAND), apex covered with rather stiff 
dark brown hairs to 10 mm long. Stipes to 100 cm 
or more, dark and densely hairy like the rhizome 



Dec. 1963] 



Cyatheaceae (Holttum) 



169 



at the base, pale and more sparsely hairy above 
the base; main rachis covered beneath with lax 
pale hairs 2 mm long and scattered dark thicker 
ones 3-4 mm long, upper surface with dense pale 
hairs in the groove. Lamina c. 1 50 cm long, basal 
pinnae to at least 50 cm; largest pinnules about 
180 by 100 mm; tertiary leaflets to 50-60 by 30 
mm; several free quaternary leaflets on larger 
pinnules, largest to about 12 by 4 mm, deeply 
lobed; quinary lobes up to 5 or 6; veins in largest 
quinary lobes pinnate, in smaller ones forked or 
simple. Sori usually one to each quinary lobe, at 
the end of the basal acroscopic vein if the vein in 
the lobe is branched; sori little over 1/2 rnm o, 
when young quite enclosed by the two indusia 
which appear almost equal, the inner more or less 
reflexed at maturity and then seen to have a thinner 
edge. Hairs c. 1 mm, pale and spreading, rather 
numerous on lower surface of pinnule-rachis and 
axes of tertiary leaflets, more sparse on quaternary 
leaflets and veins; hairs of the same kind more 
sparse on upper surface of axes. 

Type specimen: Blume, Java (L). 

Distr. Malaysia: Java, Lesser Sunda Is (Lom- 
bok, Flores), Sarawak and North Borneo, 
Philippines (Luzon, Negros). 

Ecol. In Java at 1500-2300 m, in wet shady 
forest (Backer & Posthumus; but some speci- 
mens so named by Posthumus are Microlepia); in 
the Philippines common at 1 500-2000 m in Benguet 
Prov., N. Luzon and found at 1200 m in Negros; 
in Sarawak and North Borneo found at 1250-1600 
m, on a landslide and on a ridge-top. Probably 
plants establish themselves in open places in forest 
and on landslides and persist when shadier con- 
ditions develop, as in the case of Cibotium. 
In exposed places, plants with small fronds may 
be fertile; one such was apparently the type of 
Balautium pilosum. 

2. Culcita villosa C. Chr. Brittonia 2(1937) 283; 
COPEL. Philip. J. Sc. 78 (1949) 7. 

CflMc/eA- "short, woody" (Brass). Stipes to 80 cm, 
dark at base and covered with shining brown 



hairs 10 mm long with thick bases, pale and gla- 
brescent above base. Pinnae to at least 50 cm 
long; pinnules to 150 by 70 mm; tertiary leaflets 
to 50 mm long; quaternary leaflets or segments to 
12 pairs, largest 13 mm long, mostly about 10 by 
3 mm with 5 pairs on quinary lobes; basal acro- 
scopic quinary lobe bearing 3 sori, quite filling the 
lobe, 2 sori on next lobe, one on each remain- 
ing acroscopic and on all basiscopic lobes; inner 
indusium very delicate, fringed on the edge, 
pushed back and hidden by ripe sporangia. Hairs 
on lower surface of all axes and veins very co- 
pious. 

Type specimen: Brass 4791, Vanapa Valley, 
1900 m, Papua (BM). 

Distr. Malaysia: New Guinea, west to east; 
N. Celebes (G. Tampusu, coll. Hutton). 

Ecol. Secondary growth (on old garden land) 
and in grassland, at 1600-2230 m. 

3. Culcita straminea (Labill.) Maxon, J. Wash. 
Ac. Sc. 12 (1922) 457; Copel. Fern Fl. Philip. 1 
(1958) 86. — Dicksunia straminea Labill. Sert. 
Austr. Cal. (1824)7, t. 10. — Sitolobium stramineuni 
Brack, in Wilkes, U.S. Expl. Exped. 16 (1854) 
273. — Dicksonia torreyana Brack. I.e. 278, t. 58, 
f. 2. — Dennstaedtia straminea i. Sm. Hist. Fil. 
(1875) 265. 

Caiidex erect, up to 3 m (Brackenridge). 
Fronds including stipes to 300 cm long, stipes to 
120 cm; lamina similar in size and dissection to 
that of C javanica but less hairy; inner indusia 
large, at maturity broader than outer, always 
evident and not deflexed at maturity. 

Type specimen: Labillardiere, New Caledonia 
(P; not seen). 

Distr. Polynesia (Samoa, Fiji), Melanesia 
(New Caledonia, New Hebrides, Solomon Is), 
and E. Malaysia: Louisiades, Admiralty Is, and 
Philippines (Mindanao). 

Ecol. In lowland country, up to 900 m (most 
specimens below 250 m), growing with Gleichenia 
on deforested slopes, in secondary forest and on 
river banks. 



INDEX TO CYATHEACEAE 

All suprageneric and infrageneric epithets have been entered under the name to which they belong 
preceded by the indication of their rank. 

New epithets and new combinations have been printed in bold type, synonyms in italics. 

An asterisk behind a page number marks the presence of a figure of the concerned taxon. 

Page numbers in bold type denote main treatment. 



Agyratae 71 

Alsophila sensu C. Chr. 115 

Alsophila R. Br. 73, 76 

acrostichoides v. A. v. R. 123 

acuta Pr. 135 

aeneifolia v. A. v. R. 133 

var. siibglauca v. A. v. R. 133 

allocota V. A. v. R. 152 

alpina v. A. v. R. 91, 92 

alternans Hook. 145 

amaiambitensis v. A. v. R. 1 1 7, 1 1 8 

amboinensis v. A. v. R. 114 



(Alsophila) angiensis Gepp 135 
annae v. A. v. R. 1 16 
apiculata Rosenst. 93 
arfakensis Gepp 1 1 3 
atropurpiirea C. Chr. 1 1 8 
aus trails R. Br. 77 
bakeri ZeiW. 121 
bartlettii (Copel.) C. Chr. 152 
batjanensis Christ 88 
benculensis v. A. v. R. 89 
biformis Rosenst. 118 
brevifoliolata v. A. v. R. Ill 



170 



Flora Malesiana 



[ser. II, vol. P 



(Alsophila) brimnea Brause 120 
brunoniana Hook. 137 
burbidgei Bak. 151 
bitrbidgei (non Bak.) Christ 117 
buniensis Rosenst. 131 
calocoma Christ 137 
Cauda fa J. Sm. ex Hook. 110 
celebica (Bl.) Mett. 140 
christii v. A. v. R., non Sod. 131 
dementis Copel. 135 
commutata Mett. 118 
comosa Wall, ex Hook. 152, 153 
comosa (non Wall, ex Hook.) Christ 131 
concinna Bak. 130, 131 
contaminans Wall, ex Hook. 135. 137 

var. brunoniana Scott 137 

var. celebica Christ 135 

var. densa Hassk. 135 

var. longepaleata Christ 1 34 

var. microloba Hassk. 135 

var. robusla Hassk. 135 

var. setulosa Hassk. 135 

var. squamulata Hassk. 135 
crenulata (Mett.) Hook. 1 1 1 
crinita (non Hook.) v. A. v. R. 132 
curranii (Copel.) C. Chr. 133 
cyclodonta Christ 127 
debilis De Vr. 110 
dielsii Brause 157 
dimorpha Christ 121 
dimorphotiicha (Copel.) v. A. v. R. 131 
dissirifolia Bak. 120 
dryopteroidea Brause 158 
di'ibia Bedd. 120 
elmeri Copel. 131 

externa [non (Forst.) R. Br.] Bl. 110 
fenicis (Copel.) C. Chr. 110 
fenicis [non (Copel.) C. Chr.] Posth. 131, 132 
fujiiana Nakai 110 
fuliginosa Christ 108 
gazellae Kuhn 158 
gigantea Wall, ex Hook. 124 
glabra (Bl.j Hook 120 
glabra (non Bl.) Bedd. 124 
glabrescens v. A. v. R. 152 
glauca (Bl.) Hook. 134 
glauca (Bl.) J. Sm. 68, 135 

var. celebica v. A. v. R. 135 

var. densa v. A. v. R. 135 

var. longepaleata v. A. v. R. 1 34 

var. microloba v. A. v. R. 135 

var. setulosa v. A. v. R. 135 

var. squamulata v. A. v. R. 135 

var. squamulosa v. A. v. R. 135 

var. trichocarpa Rosenst. 135 
gregaria Brause 102 
haenkei Pr. 

var. angustata Hassk. 1 34 
hallieri Rosenst. 157 
hallieri v. A. v. R. 1 1 7 
helferiana Pr. 124 
heteromorpha v. A. v. R. 1 1 8 

var. decomposita v. A. v. R. 118 
heterophylla v. A. v. R. 121 
hewittii (Copel.) v. A. v. R. 118 



(Alsophila) hieronymi Brause 88 
hornei Bak. 120 
hunsteiniana Brause 93 
incisoserrata C. Chr. 1 1 3 
indrapurae v. A. v. R. 93 
janseniana v. A. v. R. 145 
junghuhniana Kze 110 
kemberangana (Copel.) C. Chr. 151 
kenepaiana v. A. v. R. 1 1 7 
kingii Clarke 121 
laeta Kze 152 
lanuginosa (Jungh.) Pr. 132 
lastreoides v. A. v. R. 115 
latebrosa Wall, ex Hook. 115 

var. batjanensis Christ 114 

var. denudata Bedd. 110 

var. ornata Ridl. 1 1 3 
latebrosa (non Wall.) 

var. major Christ 1 3 1 
ledermannii Brause 120 
lepidoclada Christ 82 
lepifera J. Sm. ex Hook. 137 

var. congesta Christ 137 
leucocarpa (Copel.) C. Chr. 115 
longipinna (Copel.) C. Chr. 115 
lunulata (Forst.) R. Br. 131 
lunulata [non (Forst.) R. Br.] Bl. 110 
lurida (Bl.) Hook. 121 
macgillivrayi Bak. 123 
margarethae Schroet. ex Christ 150 
marginata Brause 155 
matthewii Christ 157 
melanocaulos v. A. v. R. 120, 121 
melanopus Hassk. 110 
melanoraclus Copel. 121, 122 
mindanensis Christ 108 
modes ta Bak. 102 
naumannii Kuhn 131 
obliqua (Copel.) C. Chr. 146 
obscura Scort. 152 
okiana v. A. v. R. 131 
oligosora Miq. 152 
olivacea Brause 120 
ornata (non Scott) Bedd. 113 

var. sikkimensis (non Clarke & Bak.) Bedd. 
113 
palembanica v. A. v. R. 89 
papuana Ridl. 155 
paraphysata v. A. v. R. 152 
parvifolia Holtt. 145 
persquamulata v. A. v. R. 124 
persquamulifera v. A. v. R. 134 
philippinense Hort. Veitch. 157 
poiensis (Copel.) v. A. v. R. 151 
polvcampta Kze 124 
poivphlebia Bak. (1876) 130, 131 
pulchra (Copel.) C. Chr. 152 
punctulata v. A. v. R. 112 
pustulosa Christ 137 
ramispina Hook. 117 
ramosii (Copel.) C. Chr. 151 
rebeccae F. v. M. 120 
recurvata Brause 101 
reducta v. A. v. R. 120 
ridleyi Bak. 152, 153 



Dec. 1963] 



Index to Cyatheaceae 



171 



(Alsophila) robusta De Vr. 110 

rosenstockii Brause 101 

nihiginosci Brause 93 

nimpliianci v. A. v. R. 131 

sal villi i Hook. 115 

sangirensis Christ 1 30 

saparuensis v. A. v. R. 88 

sarawakensis C. Chr. 152 

scaberiila Christ 130 

scaheruUpes v. A. v. R. 131 

scahrisetci (Copel.) v. A. v. R. 131 

scandeiis Brause 119 

schk'clueri Brause 123 

smitliiana Pr. 135 

speciosa [non (Meyen) Pr.] Goldm. 157 

spinifera v. A. v. R. 114 

squaniulata (Bl.) Hook. 152 

sqiiamulata (non Bl.) Hook. 118 

straminea Gepp 88 

subcomosa C. Chr. 131 

subdimorpha (Copel.) v. A. v. R. 121 

siibdubia v. A. v. R. 124 

subobscura v. A. v. R. 152 

subulata v. A. v. R. 1 18 

tenggerensis Rosenst. 134 

tenuis Brause 90 

tomeutosa (Bl.) Hook. 132 

var. novogiiineensis Rosenst. 132 

tiichodesma Scort. 150 

trichophora (Copel.) v. A. v. R. 151 

tristis (Bl.) Bl. ex Moore 158 

truiicata Brack. 157 
var. nivea Christ 140 
var. sagittata Christ 140 

umbrosa Wall, ex Ridl. 124 

vexans Ces. 120 

vitieiisis Carr 131 

wallacei Mett. 151 

warbiirgii Christ 158 

warilwn (Copel.) C. Chr. 108 

wengiensis Brause 88 

woodlarkensis (Copel.) C. Chr. 158 

xanthina (Domin) C. Chr. 152 

xautlwlepia v. A. v. R. 152 

xantholepis Christ ex Diels 158 
Amphicosmia Gardn. 73 

alternaiis Moore 145 

Javanica (Pr.) Moore 111 

mamlensis (Pr.) Moore 110 
Aspidistes Harris 65, 67 
Aspidium barometz Willd. 165 

triste Bl. 158 
Athyrium 69 
BalarUium Kaulf. 71, 158, 166, 167 

auricomiim 167 

bluniei Kze 159 

chrysotrichiim Hassk. 159 

copelandii Christ 168 

ciilcita 167 

glaucescens Link 165 

javamciim Copel. 168 

pilosum Copel. 168, 169 
Cathetogyratae 71 
Chnoophora Kaulf. 73 

glaiica Bl. 135 



(Chnoophora) lanuginosa Jungh. 132 

liirida Bl. 121 

tumentusa Bl. 132 
Cibotium Kaulf. 65, 67, 69, 70, 71, 72, 159, 164- 

166, 169 

arachnoideum (C. Chr.) Holtt. 164, 166 

assumicum Hook. 165 

barometz (L.) J. Sm. 164, 165* 
var. cumingii v. A. v. R. 166 
var. siimcitrainim Christ 165 

chamissoi Kaulf. 164 

crassinerva Rosenst. 166 

cumingii Kze 164, 166 

var. arachnoideum C. Chr. 166 

djambianum Hassk. 165 

glaucescens Kze 165 

glaucum [non (Sm.) Hook. & Arn.] Bedd. 165 

glaucum [non (Sm.) Hook. & Arn.] J. Sm. 166 

regale Linden 165* 
Cnemidaria 65, 69, 70, 72 

horrida 69, 70 
Coniopteris 65, 67 

hymenophylloides (Brongn.) Seward 65 

murrayana Brongn. 65 
Ctenitis 167 
Culcita Pr. 65, 67, 69, 70, 72, 159, 166-169 

coniifolia (Hook.) Maxon 167 

copelandii Maxon 168 

dubia (R. Br.) Maxon 70, 167, 168* 

formosae (Christ) Maxon 167 

javanica (Bl.) Maxon 167, 168* 

macrocarpa Pr. 70, 167 

pilosa Christ 168 

straminea (Labill.) Maxon 167, 169 

subg. Calochlaena 65, 166, 167 

subg. Culcita 65, 166, 167 

villosaC. Chr. 167, 169 
Cyathea Smith 65, 67, 69, 70, 71, 72-158, 159 

acanthophora HoUt. 79, 81, 93 

acanthopoda v. A. v. R. 93 

aciculosa Copel. 138, 140 

acrostichoides (v. A. v. R.) Domin 116, 123 

acuminata Copel. 79, 98 

adenochlamys Christ 157 

aeneifolia (v. A. v. R.) Domin 126, 133 
var. macrophylla Holtt. 134 
var. melanacantha (Copel.) Holtt. 134 

agatheti Holtt. 142, 152 

albidosquamata Rosenst. 78, 88 

albidula Domin 93 

alderwereltii Copel. 81, 114 

alleniae Holtt. 81, 109 

allocota (v. A. v. R.) Domin 152 

alpicola Domin 91 

alpina v. A. v. R. 91 

alsophiliformis (v. A. v. R. j Domin 1 1 1 

alternans (Wall, ex Hook.) Pr. 76, 141, 145, 150 
var. lobbiana (Hook.) Domin 145 
var. sarawakensis (Hook.) Domin 145 
var. serrata Ridl. 145 

amaiambitensis (v. A. v. R.) Domin 117 

amboinensis (v. A. v. R.) Merr. 81, 114, 157 

amphicosmioides v. A. v. R. 89 

ampla Copel. 150 

ampla (non Copel.) Holtt. 151 



172 



Flora Malesiana 



[ser. II, vol. 12 



(Cyathea) aneitensis Hook. 157 

angiensis (Gepp) Domin 126, 135, 136 

angustipinna Holtt. 141, 143 

annae (v. A. v. R.) Domin 116, 117* 

apiculata (Rosenst.) Domin 79, 93, 94 

apoensis Copel. 78, 86 

arachnoidea Hook. 140 

aiacluwidea (non Hook.) Grether & Wagner 140 

arachnoidea (non Hook.) Back. & Posth. 140 

arborea (L.) Sm. 74 

arborea [non (L.) Sm.] 

var. pallida Hassk. 86 
arborescens Cox)Q\- 110 
archboldii C. Chr. 79, 98 

var. horrida Holtt. 99 
arfakensis Gepp 77, 82 
argil ta Copel. 146 
argyrolepis Copel. 131 
arthropoda Copel. 141, 143 
arthropterygia v. A. v. R. 89 
anieiisis Domin 131 
ascendens Domin 80, 101 
aspidioides (Bl.) ZoU. & Mor. 158 
assimilis Hook. 142, 150 
assimilis (non Hook.) Christ 101 
atrispora Domin 158 
atropurpurea Copel. 116, 118 
atrospinosa Holtt. 126, 137 
atrox C. Chr. 66*, 107*, 126, 128 

var. atrox 128 

var. inermis Holtt. 126, 128, 129*, 130* 
auriculifera Copel. 138, 139 
australis (R. Br.) Domin 69 
aiistrosinica Christ 106 
barisanica (v. A. v. R.) Domin 89 
bartlettii Copel. 152 
batjanensis (Christ) Copel. 75*, 78, 88 
beccariana Ces. 150 
benculensis (v. A. v. R.) v. A. v. R. 89 
bicolana Copel. 149 
bicolor Copel. 108 
bideiitata Copel. 98 
biformis (Rosenst.) Copel. 68, 116,118,119*, 

120 
biliranensis Copel. 108 
binuangensis v. A. v. R. 141, 145 
bipiimatifida Copel. 1 5 1 
bontocensis Copel. 149 
borbonica Desv. 141 
borneensis Copel. 81, 110 
brackenridgei Mett. 157 
brassii Copel. 131 
braiiseana Domin 88 

brevifoliolata (v. A. v. R.) v. A. v. R. Ill 
breviloba Copel. 146 
brevipes Copel. 110 
brooksii Copel. (non Maxon) 152 
brownii Domin 70* 
brunei Christ 124 
briinnea (Brause) Domin 120 
brunoniana (Hook.) Clarke & Bak. 137 
brimonis (J. Sm.) Wall, ex Hook. 143 
buhisanensis Copel. 146 
bunnemeijerii v. A. v. R. 78, 79, 94 
burbidgei (Bak.) Copel. 151 



(Cyathea) burbidgei [non (Bak.) Copel.] Holtt. 150 

buriiensis (Rosenst.) Domin 131 

callosa Christ 80, 106 

calocoma (Christ) Copel. 137 

camagiiinensis Copel. 106 

campbellii Copel. 104 

capensis (L. f. ) Sm. 70* 

capitata Copel. 65, 141, 142, 144* 

carrii Holtt. 138, 139* 

catillifera Holtt. 78, 90 

caudata (J. Sm. ex Hook.) Copel. 81, 108, 110, 

114 
caiidicidata (Rosenst.) Domin 108 
caiidipifunila (v. A. v. R.) Domin 89 
celebica Bl. 138, 140, 157 
celebica v. A. v. R. (non Bl.) 139 
cheilanlhoides Copel. 95 
christii Copel. 79, 95 
cincinnata Brause 77, 86 
cinerea Copel. 80, 104 
dementis (Copel.) Copel. 135 
coactilis Holtt. 78, 87 
confliiens (v. A. v. R.) Domin 102 
contaminans (Wall, ex Hook.) Copel. 67. 68*, 

70*, 71, 125*, 126,135, 136*, 138 

var. persquamulifera v. A. v. R. 134 
costalisora Copel. 78, 87 
costulisora Domin 81, 109 
crassipes Sod. 124 
crenulata Bl. 76, 78, 89, 92*, 93, 95, 104 

f. latissima v. A. v. R. 103 

f. sqiiamitlosa v. A. v. R. 103 

f. subspiniilosa v. A. v. R. 93 
crinita (Hook.) Copel. 132 
cucLillifera Holtt. 80, 103 
curranii Copel. 126, 133 
ciirvipinnida C. Chr. 133 
cyclodonta (Christ) v. A. v. R. 127 
deminuens Holtt. 141, 145 
deiisisora v. A. v. R. 140 
deuterobrooksii Cope\. 152 
dicksonioides Holtt. 80, 83*, 106, 107* 
dielsii (Brause) Domin 157 
dimorpha (Christ) Copel. 116, 121 
dimorphophylla Domin 120 
dimorplwtricha Copel. 131 
discophora Holtt. 76, 142, 148 
distans Rosenst. 92 
dissitifolia Domin 120 
doctersii v. A. v. R. 80, 102 
diibia Bedd. 120 
diilitensis Bak. 150 
diipaxensis Copel. 1 10 
dura Copel. 108 
edanoi Copel. 80, 108 
elliptica Copel. 141, 142, 146, 151 
elmeri (Copel.) Copel. 126, 131, 132 
eminens Domin 131 
eriophora Holtt. 80, 102 
everta Copel. 77, 85 
excavata Holtt. 79, 94* 
excelsa sensu Kze (non Sw.) 92 
faberiana Domin 103 
fallax (v. A. v. R.) Domin 111 
fenicis (C. Chr.) Copel. 81, 110 



Dec. 1963] 



Index to Cyatheaceae 



173 



(Cyathea) ferruginea Christ 79, 100 
ferrugineoides Copel. 100 
foersteri Rosenst. 79, 99, 100 
foxwurthyi Copel. 106 
frondosa Rosenst. 157 
fnictiiosa Copel. 104 
fugax V. A. V. R. 126, 137 
fuliginosa (Christ) Copel. 81, 108 
fusca Bak. 153, 154, 156 
fiiscopaleata Copel. 143 
gazelkie (Kuhn) Domin 158 
geluensis Rosenst. 79, 95 

var. tomentosa Rosenst. 95 
geppiana Domin 88 
gibbsiae Copel. 118 

gigantea (Wall, ex Hook.) Holtt. 116, 120, 124 
glaberrima Holtt. 81, HI 
glabra (Bl.) Copel. 67, 115, 116, 120 
glabrescens (v. A. v. R.) Domin 152 
glauca Bory 135 

glaucophylla (v. A. v. R.) Domin 111 
glaucum [non (Sm.) Hook. & Arn.] J. Sm. 166 
gleichenioides C. Chr. 66*, 67*, 79, 96, 97* 
globosora Copel. 85 
gracillima Copel. 123 
grata Domin 157 

gregaria (Brause) Domin 80, 101, 102, 123 
haenkei (Pr. ) Merr. 132 
halconensis Christ 79, 101 
hallieri (Rosenst.) Domin 157 
havilandii Bak. 79, 96 
hemichlamxdea Copel. 110 
heterochlamydea Copel. 75*, 80, 108, 110, 157 
heteroloba Copel. 149 
heteiomorpha (v. A. v. R.) Domin 118 
heterophylla (v. A. v. R.j Domin 121 
hewittii Copel. 118 
holttumii Copel. 146 
hooglandii Holtt. 77, 83*, 84 
hookeri Thw. 141 
hornei (Bak.) Copel. 116, 120 
honidipes (v. A. v. R.) Domin 114 
horridula Copel. 78, 90 
hunsteiniana Brause 77, 82 

var. acuminata Brause 82 
hymenodes Mett. 76, 78, 89, 94 
hypocraterijormis v. A. v. R. 146 
imbricata v. A. v. R. 79, 97 
inaequalis Holtt. 153, 155 
incisoserrata Copel. 73*, 74*, 77, 81, 92, 112*, 

113 
indrapurae (v. A. v. R.) v. A. v. R. 93, 94 
indusiosa Copel. 104 
inquinans Christ 79, 100 
insignis Eaton 124 
insulana Holtt. 79, 98 
insularum Holtt. 153, 156 
Integra J. Sm. ex Hook. 142, 146, 147*, 156 

\ar. petiolata Hook. 146 
Integra (non J. Sm. ex Hook.) v. A. v. R. 147 
janseniana (v. A. v. R.) Domin 145 
javanica Bl. 75*, 76, 78, 89, 103 

var. rigida Bl. 89, 92 
junghuhniana (Kze) Copel. 71, 81, 110, 111 
kanehirae Holtt. 81, 113 



(Cyathea) kemberangana Copel. 151 
kenepakina (v. A. v. R.) Domin 117 
keysseri Rosenst. 95 
kinahaliiensis Copel. 143 
kingii (Clarke) Copel. 121 
kingii Rosenst. 156 
klossii Ridl. 78, 91 
kort/ialsii Mett. 89 
kurt/uilsii (non Mett.) C. Chr. 104 
lanaensis Christ 108 
lastreoides (v. A. v. R.) Domin 115 
latebrosa (Wall.) Copel. 67, 71, 77, 81, 92, 103, 

111, 113, 115 

var. indusiata Holtt. 89 
latipinnula Copel. 80, 105 
ledermannii Brause 77, 83 

var. dilatata Brause 83 
lepidigera Copel. 108 
lepidoclada (Christ) Domin 77, 82 
lepifera (J. Sm.) Copel. 126, 136, 137 
leptolepia (v. A. v. R.) Domin 115 
leucocarpa Copel. 1 1 5 
leucophaes Hassk. 92 
leiicostegia Copel. 140 
leucotricha Christ 125, 127, 128 
leytensis Copel. 140 
lohata Copel. 86 
lobbiana Hook. 145 
ioerzingii Holtt. 80, 105 
loheri Christ 76, 80, 104 

var. tonglonensis Christ 108 
longipaleata Alston 103 
longipes Copel. 79, 98 
longipes v. A. v. R. 92 
longipinna Copel. 115 
lunulata (Forst.) Copel. 126, 131 
lurida (Bl.) Copel. 116, 121, 122* 
macgillivrayi (Bak.) Domin 80, 88, 115, 116, 123 
macgregorii F. v. M. 67, 79, 83*, 95, 96*, 108 
macrophylla Domin 138, 140 

var. quadripinnata Holtt. 141 
macropoda Domin 78, 92 
magna Copel. 126, 132 
magnifolia v. A. v. R. 79, 92, 93 
manilensis (Pr.) Domin 110 
margarethae (Schroet. ex Christ) Copel. 150 
marginata (Brause) Domin 153, 155 
masapilidensis Copel. 80, 105 
matthewii (Christ) Domin 157 
mearnsii Copel. 101 
media Wagner & Grether 81, 115 
medullaris (Forst.) Sw. 124 
megalosora Copel. 142, 148 
melanacantha Copel. 134 
melanoclada Domin 121 
melanophlebia Copel. 101 
mekuwpiis (Hassk.) Copel. Ill 
melanorachis (Copel.) Copel. 121 
merapiensis (v. A. v. R.) Domin 111 
merrillii Copel. 108 
mertensiana (Kze) Copel. 133 
mesosora Holtt. 153, 155 
microchlamys Holtt. 81, 114 
microphylloides Rosenst. 77, 82, 84 
mindanensis (Christ) Copel. 108 



174 



Flora Malesiana 



[ser. II, vol. 12 



(Cyathea) mitrata Copel. 104 
modesta (Bak.) Copel. 80, 102 
mollis Copel. 151 

moluccana R. Br. 67, 141, 143, 144*, 145 
montana Sm. 109 
moseleyi Bak. 153, 157 
muelleri Bak. 80, 98. 103, 104* 
naiimanini (Kuhn) Domin 131 
negrosiana Christ 78, 90 
nigrolineata Holtt. 79, 100 
nigropaleata Holtt. 81, 113 
nigrospiiudosa v. A. v. R. 156 
novae-caledoniae (Mett.) Copel. 138 
novogiiineensis Brause 95 
obliqua Copel. 141, 146 
obscura (Scort.) Copel. 142, 152 
obtusata Rosenst. 110 

oinops Hassk. 74*, 75*, 76, 80, 92, 103, 104 
oinops (non Hassk.) Racib. 92 
okiana (v. A. v. R.) v. A. v. R. 131 
oligocarpia Jungh. 103 
olivacea (Brause) Domin 121 
oosora Holtt. 79. 101 
ordinata Copel. 132 

orientaiis (Kze) Moore 73*, 75*, 78, 86, 95 
pachyrrhachis Copel. 80, 105 
paleaceci Copel. 96 
paleata Copel. 93 

palembanica (v. A. v. R.) v. A. v. R. 89 
pallidipaleata Holtt. 78, 87 
papuana (Ridl.) v. A. v. R. 153, 155 
paraphysatci Copel. 152 
paraphysophora (v. A. v. R.J Domin 114 
parva Copel. 78, 87 
parvipinna Holtt. 153, 155 
pateliifera v. A. v. R. 78, 89 
peranemiformis C. Chr. 82 
percrassa C. Chr. 77, 84 
perpelvigera v. A. v. R. 77, 82, 84 
perpunctulata (v. A. v. R.) Domin 81, 114 
persquamulata (v. A. v. R.) Domin 124 
persquamulifera (v. A. v. R.) Domin 126, 134 
philippinensis Bak. 142, 149, 150 

var. Hilda Copel. 149 
physolepidota Alston 81, 113 
pilulifera Copel. 126, 133, 134 
pinna ta Roxb. 143 
podophylla (Hook.) Copel. 120 
poiensis Copel. 151 
polycarpa Jungh. 92 
polypoda Bak. 142, 150, 151 
princeps (Linden) Meyer 124 
procera Brause 125, 127* 
pruinosa Rosenst. 77, 84 
pseudoalbizzia Copel. 149 
pseiidobninonis Copel. 143 
pseudomuelleri Holtt. 79, 98, 99* 
pteridioides Copel. 137 
pulcherrima Copel. 125, 126, 127, 153 
pidchra Copel. 152 
pumilio V. A. v. R. 88, 89 
punctulata (v. A. v. R.) v. A. v. R. 81, 112 
pustulosa (Christ) Copel. 137 
pycnoneura Holtt. 77, 85 
quadripinnatifida Copel. 140 



(Cyathea) raciborskii Copel. 81, 111 

ramispina (Hook.) Copel. 116, 117, 118, 121 

ramosiana v. A. v. R. 149 

ramosii Copel. 151 

rebeccae (F. v. M.) Domin 116, 120 

recommutata Copel. 117*, 118, 121, 122* 

recurvata (Brause) Domin 80, 101 

reducta (v. A. v. R.) Domin 120 

ridleyi (Bak.) Copel. 145, 152 

rigens Rosenst. 77, 85, 86 

rigida Copel. 96 

robinsonii Copel. 142, 149 

rosenstockii Brause 153, 154*, 156 

rubella Holtt. 116, 122 

rubiginosa (Brause) Domin 79, 93 

nidimentaris (v. A. v. R.) Domin 115 

rufopannosa Christ 80, 106 

riimphiana (v. A. v. R.) Merr. 131 

rumphii Desv. 157 

runensis v. A. v. R. 153, 156 

saccata Christ 78, 93 

salticola (v. A. v. R.) Domin 114 

sangirensis (Christ) Copel. 74, 126, 130, 132 

sapaniensis (v. A. v. R.) v. A .v. R. 88 

sarasinorum Holtt. 126, 134 

sarawakensis Hook. 145 

scaberida (Christ) Domin 131 

scabendipes (v. A. v. R.) Domin 131 

scabriseJa Copel. 131 

scandens (Brause) Domin 116, 119 

schizochlamys Bak. 91 

schlechteri (Brause) Domin 116, 123 

semiamplectens Holtt. 81, 109 

senex v. A. v. R. 142, 149 

sepikensis Brause 95 

sessilipinnida Copel. 148 

setifera Holtt. 153, 154 

setulosa Copel. 80. 103 

sibuyanensis Copel. 142, 149 

singalanensis (v. A. v. R.) Domin 102 

sinops (sphalma), of. oinops 92, 104 

sinuata Hook. 141 

spinifera (v. A. v. R.) Domin 114 

spinulosa Wall, ex Hook. 98, 106 

var. miiricidata Hassk. 92 
squamicosta Copel. 108 
squamulata (Bl.) Copel. 67, 74*, 142, 145, 147*, 

148. 152 
stipitipinnula Holtt. 142, 147 
stipititlata Copel. 150 

straminea (Gepp) v. A. v. R., non Karst. 88 
strigosa Christ 125, 128 
subbipinnata Copel. 146 
subcumosa (C. Chr.) Domin 131 
siibconjiuens (v. A. v. R.) Domin 102 
subdimoipha Copel. 121 
subdubia (v. A. v. R.) Domin 116, 120, 124 
subg. Cyathea 69, 73, 75*. 76, 77-124 

sect. Cyathea 76, 77-115, 123 

sect. Gymnosphaera (Bl.) Holtt. 76, 80, 
115-124, 141 
subg. Gymnosphaera Tindale 115 
subg. Sphaeropteris (Bernh.) Holtt. 69, 74, 76, 

124-158 

sect. Schizocaena (J. Sm.) Holtt. 76, 124, 141 



Dec. 1963] 



Index to Cyatheaceae 



175 



(Cyathea) 

subsect. Sarcopholis Holtt. 76, 141, 153-158 
subsect. Schizocaena 76, 112, 141-153 
sect. Sphaeropteris 69, 76, 124-141, 151 
subsect. Fourniera (Bommer) Hollt. 76, 

124, 138-141 
subsect. Sphaeropteris 76, 124, 125-138, 153 
subobscura (v. A. v. R.) Domin 152 
subspaihulata Brause 95 
subtripinnata Holtt. 77, 86 
siibulata (v. A. v. R) Domin 118 
subuliformis v. A. v. R. 91 
sulitii Co pel. 108 
suluensis Bak. 142, 148 
sumatrana Bak. 78, 91, 149 
taiwaniana Nakai 106 
tenggerensis (Rosenst.) Domin 126, 134 
tenuicaulis Domin 78, 90 
tenuis Brause 90 
ternatea v. A. v. R. 78, 88 
teysmannii Copel. 138, 139 
tomentosa (Bl.) ZoU. & Mor. 126, 132, 133 
tomentosissima Copel. 126, 128 
tonglonensis (v. A. v. R.) Domin 108 
toppingii Copel. 118 
trachypoda v. A. v. R. 78, 91, 92 
trichodesma (Scort.) Copel. 142, 150, 151 
trichophora Copel. 142, 146. 150, 151 
tripinnata Copel. 138, 139, 140 
tripinnatifida Roxb. 147, 153, 156, 157 
truncata (Brack.; Copel. 140 
tuber culata v. A. v. R. 91 
umbrosa Copel. 137 
urdanetensis Copel. 146, 147 
vandeusenii Holtt. 77, 84 
veitchiani Domin 157 
verrucosa Holtt. 126, 135 
versteegii Christ 156 
vexans C. Chr. 120 
vitiensis (Carr.) Domin 131 
wallacei (Mett.) Copel. 142, 151 
warihon Copel. 108 

wengiensis (Brause) Domin 78, 81, 88 
werneri Rosenst. 153, 154, 156 
womersleyi Holtt. 138, 139 
woodlarkensis Copel. 158 
xanthina Domin 152 
xantholepis (Christ ex Diels) Domin 158 
zamboangana Copel. 142, 147 
zoUingeriana Mett. 92 
zollingeriana (non Mett.) v. A. v. R. 103 
Cyatheaceae 65-169 

subfamily Cibotioideae 72 
subfamily Cyatheoideae 71 

tribe Cyatheae 71, 72 

tribe Dicksonieae 71, 72 

tribe Lophosorieae 72 
subfamily Metaxyoideae 72 
subfamily Thyrsopteridoideae 72 

tribe Culciteae 72 

tribe Thyrsopterideae 72 
Cystodium J. Sm. 65, 69, 72, 162-164 

sorbifolium (Sm.) J. Sm. 67, 163* 
Davallia 167 
Dennstaedtia 68, 69. 71, 159, 167 



(Dennstaedtia) ampla (Bak.) Bedd. 161 
cuneata (Hook.; Moore 161 
erythrorachis (Christ) Diels 162 
flaccida (Forst.) Bernh. 162 
giabrata (Ces.) C. Chr. 162 
gomphophylla (Bak.) C. Chr. 162 
javanica Christ 168 
moluccana (Bl.) .Moore 162 
midtifida v. A. v. R. 168 
paraphysata v. A. v. R. 168 
remota (Christ) Diels 162 
rhombifolia (Bak.) C. Chr. 162 

scabra (Wall.) Moore 161, 162 

scandens (Bl.) Moore 162 

smithii (Hook.) Moore 162 

straminea i. Sm. 169 
Diacalpe aspidioides Bl. 158 
Dichorexia Pr. 73, 76 

latebrosa Pr. 115 
Dicksonia L'Herit. 65, 67, 68, 69, 70, 71, 72, 74, 

108, 158-162, 167 

ampla Bak. 161 

antarctica Labill. 69, 160* 

arborescens L'Herit. 158, 159, 167 

archboldii Copel. 159, 160 

assamicum Griff. 165 

baranetz Link 165 

blumei (Kzej Moore 69, 159, 160* 

blumei [non (Kze) Moore] C. Chr. 159 

chrysotricha (Hassk.) Moore 159 

copelandii Christ 168 

cidcita L'Herit. 167 

cuneata Hook. 161 

deltoidea Hook. 161 

erythrorachis Christ 162 

flaccida (Forst.) Sw. 162 

giabrata Ces. 162 

gomphophylla Bak. 162 

grandis Rosenst. 159, 161 

hieronymi Brause 159, 161 

japonica Sw. 162 

javanica Bl. 168 

kingii Bedd. 162 

lanigera Holtt. 159, 161 

ledermannii Brause 161 

linearis Cav. 162 

mollis Holtt. 159, 160* 

moluccana Bl. 162 

moluccana Roxb., non Bl. 163 

neglecta Fee 162 

nephrolepioides Christ 162 

papuana F. v. M. 163 

remota Christ 162 

rhombifolia Bak. 162 

scabra Wall, ex Hook. 162 

scandens Bl. 162 

schlechteri Brause 161 

sciurus C. Chr. 159, 160, 161 

sect. Eudicksonia Hook. & Bak. 158, 166 

smithii Hook. 162 

sorbifolia Sm. 163 

straminea Labill. 169 

strigosa Ihunb. 162 

subg. Balantium Hook. 158, 166 

torrevana Brack. 169 



176 



Flora Malesiana [ser. II, vol. P, Dec. 1963] 



(Dicksonia) zippeliana Kze 162 
Dicksoniaceae 65, 71 
Diplazium 157 
Disphenia Pr. 73, 76 

or lent alls Kze 86 
Dryopteris 65, 67, 69, 158, 167 

atrispora C. Chr. 158 
Eatoniopsis Bommer 73 
Foiirniera Bommer 73, 138 
Gleichenia 70, 71, 169 
Gleicheniaceae 71 

series Marginales 71 

series Siiperficiales 71 
Gymnosphaera Bl. 73, 76, 115 

atropiirpurea (Copel.) Copel. 118 

biformis (Rosenst.) Copel. 118 

bipinnatifida Copel. 151 

burbidgei (Bak.) Copel. 151 

dinagatensis Copel. 151 

glabra Bl. 120 

glabra (non Bl.) Copel. 151 

gracillima Copel. 123 

hewittii Copel. 1 18 

holttumii Copel. 146 

hornei Copel. 121 

kingii (Copel.) Copel. 121 

melaiwclada Copel. 121 

melanorachis Copel. 121 

mollis (Copel.) Copel. 151 

obliqiia (Copel.) Copel. 146 

papiiana (Ridl.) Copel. 155 

piilchra Copel. 152 

ramispimi (Hook.) Copel. 117 

recommutata Copel. 118 

sarawakensis (C. Chr.) Copel. 152 

schleclueri (Brause) Copel. 123 

sqitamiikita Bl. 152 

sqiiatnulata (non Bl.) J. Sm. ex Hook. 118 

siibbipiiuuita Copel. 146 

trichophora Copel. 151 

vexans (Ces. ) Copel. 120 
Helicogyratae 71 
Hemitelia R. Br. 69, 71, 73, 77 

alsopluliformis v. A. v. R. Ill 

alternans Hook. 145 

arfakensis (Gepp) v. A. v. R. 82 

barisanica v. A. v. R. 89 

bicolor (Copel.) v. A. v. R. 108 

capensis Hook. 1 1 1 

caudata (J. Sm.) Mett. 110 

caiidiciilata Rosenst. 106, 108 

caiidipinnula v. A. v. R. 89 

coiijiuens v. A. v. R. 102 

crenulata Mett. 1 1 1 

fallax V. A. V. R. Ill 

var. major v. A. v. R. Ill 

glaucophylla v. A. v. R. Ill 

hemichlamydea (Copel.) v. A. v. R. 110 

heterochlamydea v. A. v. R. 108 

horrida 73 

horridipes v. A. v. R. 1 14 

javanica Pr. 110 

junghiihnkina (Kze) Mett. Ill 
var. dissoluta Racib. 1 1 1 



(Hemitelia) latebrosa (Wall.) Mett. Ill, 115 
var. paraphysata v. A. v. R. 115 

latipiivutla v. A. v. R. 105 

ledermannii Brause 140 

leptolepia v. A. v. R. 115 

manilensis Pr. 110 

merapiensis v. A. v. R. 1 1 1 

montana v. A. v. R. 109 

midtiflora (Sm.) R. Br. 77 

paraphysophora v. A. v. R. 1 14 

perpunctiilata v. A. v. R. 114 

riidimentaris v. A. v. R. 115 

salticola v. A. v. R. 1 14 

singalanensis v. A. v. R. 102 

subconfliiens v. A. v. R. 102 

sumatraua v. A. v. R. 114 

tonglonensis v. A. v. R. 108 

truncata (non Brack.) Christ 139 

war ikon (Copel.) v. A. v. R. 108 
Kylikipteris 65 

Lastrea dryopteroidea Copel. 158 
Lindsaea 71 
Lophosoria 65, 70, 72 
Loxsoma 70 
Matonia 71 
Metaxya 65, 72 
Microlepia 168 

strigosa (Thunb.) Pr. 162 
Nephrolepis dicksonioides Christ 162 
Orthiopteris kingii (Bedd.) Holtt. 162 
Osmunda 71 

Palmifilix alba Rumph. 131 
Pleocnemia cumingiana Pr. 158 
Polybotrya arfakensis Gepp 118, 119 
Polypodiaceae 71 
Polypodium 

alternans Wall. 145 

arboreum Lour. 157 

barometz L. 165 

contaminans Wall. 135 

latebrosum Wall. 115 

liiniilatiim Forst. 131 
Pseudogyratae 71 
Pteridaceae 71 

Saccoloma sorbifolia Christ 163 
Schizocaena J. Sm. 73, 76, 141 

alternans J. Sm. 145 

arthropuda (Copel.) Copel. 143 

brunonis J. Sm. ex Hook. 141, 143 

capita ta Copel. 142 

gaudichaudii Fee 143 

kinabaliiensis (Copel.) Copel. 143 

moluccana (R. Br.) Copel. 143 
Sitolobiiim stramineum Brack. 169 
Sphaeropteris Bernh. 124 
Stenochlaena diibia v. A. v. R. 118 
Stenolepia tristis (Bl.) v. A. v. R. 158 
Tapeinidium pinnatum (Cav.) C. Chr. 162 
Thelypteris 65, 66, 158 

immersa (Bl.) Ching 158 
Thyrsopteris 65, 69, 70, 72, 167 
Thysanobotrya v. A. v. R. 73, 115 

arfakensis (Gepp) v. A. v. R. 118 
Trichopteris /a/cara Llanos 146 



LINDSAEA-GROUP (K. U. Kramer, Utrecht) 

Small to medium-sized, rarely large, terrestrial or epiphytic ferns. Rhizome 
creeping, terrestrial and radially symmetric or nearly so, solenostelic or more 
often with a special type of protostele with internal phloem but without internal 
endodermis and medulla; or epiphytic and with a similar but strongly dorsiventral 
protostele with the internal phloem close to the dorsal side of the xylem, or in 
some small species the xylem strand open and U-shaped. Petioles with a single 
U- or V-shaped vascular bundle. Indument of the rhizome of scales, these non- 
peltate, non-clathrate (in Mai. spp.), glabrous, entire, or with weakly developed 
teeth of two protruding cell-ends; terminal cell of scale glandular. Juvenile leaves 
with similar but narrower, caducous scales. In some species some or even all 
scales are entirely uniseriate but not true hairs. Laminal parts with scattered 
microscopical two-celled hairs, hardly ever with macroscopically visible hairs. 
Axes of leaves adaxially with a single groove bordered by ridges, both mostly 
continuous with those on axes of different order. Lamina once pinnate to decom- 
pound (rarely simple in a single Old World sp.), anadromous. Ultimate divisions 
various, often dimidiate. Veins free, or reticulate without free included veinlets, 
not reaching the margin. Sori terminal on one to many veins, often on a com- 
missure i; parallel to the margin, submarginal, indusiate; indusium attached at 
its base, the sides free or adnate, the free edge next to the leaf-margin and often 
rt equaling it. Sporangia ± long-stalked, with a triseriate stalk; bow of annulus 
interrupted; stomium well differentiated or not. Spores trilete or less often mono- 
lete, without perisporium, smooth or with little sculpture. Paraphyses mostly, 
perhaps always, present, filiform, 2- to many-celled, often early disappearing. 
Gametophyte known in very few species, cordate. 

Distribution. Pantropic, extending considerably beyond the tropics in Japan, Australia, South 
Africa, and eastern South America; comparatively weakly represented in continental Africa. Six genera: 
Odontosoria (10 American spp., 1 African spp.), Ormoloma (2 spp., neotropical), Tapeinidium (17 spp., 
SE. Asia to Samoa), Sphenomeris (1 1 spp., pantropic-subtropic), Xyropteris (monotypic, Malesian), and 
Lindsaea (c. 150 spp., pantropic-subtropic). 

The group is much more diversified in the Old than in the New World, but species here regarded as 
primitive occur in both hemispheres, and the origin and early history of the group cannot be traced; 
there are no fossils that can be positively attributed to one of its genera. 

Ecology. Most Lindsaeoid ferns are forest plants, but some of them occur often or mostly by and 
in beds of watercourses. A number of species grow on rocks in locally moist situations, e.g. by the coast 
(Lindsaea orbiculata, Sphenomeris biflora). A few prefer open, exposed situations, on banks, in natural 
and artificial grassland, and may then become somewhat weedy (Lindsaea ensifolia). None of the species 
goes beyond an altitude of c. 3000 m; in subtropical regions the altitudinal limit is, of course, much lower. 
Above c. 1500 m some species that are otherwise strictly terrestrial are sometimes found on moss- 
covered tree trunks etc. and are then often described on labels as epiphytes which they are not in the true 
sense of the word. 

Morphology & anatomy. For an account of the morphology and anatomy of the group see 
Perez Arbelaez (Bot. Abh. Goebel 14, 1928), Wagner (Un. Cal. Publ. Bot. 26, 1, 1952), and Kramer 
(Act. Bot. Neerl. 6, 1957, 97-134; ibid. 15, 1967, 562-584; Blumea 15, 1968, 557-561). To this may be 
added the following notes on the sporangium. The annulus has a bow of c. 8-23 thickened cells. In Lind- 
saea very many species have 10, 11, or 12 bow cells, 11 being a particularly common number. Some 
species have, however, consistently larger numbers, especially in the sections Isoloma, Osmolindsaea, 
and Tropidolindsaea, where numbers between 14 and 20 are common. Here the sporangial head is also 
somewhat larger. This holds for Tapeinidium, Xyropteris, and Sphenomeris, too, where the annulus has 
15-23, usually between 16 and 18 indurated cells. 

In most species the stomium is morphologically not or scarcely differentiated from the non-indurated 
part of the annulus. Two to four well-marked lip cells occur, however, in the sections Isoloma, Trop- 
idolindsaea, and nearly all species of sect. Schizoloma. The character appears to fluctuate, with many 
transitional cases, in Tapeinidium and Sphenomeris. 

The bow of the annulus reaches up to or slightly beyond the insertion of the stalk in Tapeinidium and 

(177) 



178 Flora Malesiana [ser. II, vol. P 

most species o( Lindsaea; in the latter genus the taxonomic value of the character is slight. In Sphenotneris 
it is also variable but apparently well-marked and constant for each species (Kramer, 1957, I.e. 106). 

Cy totaxonomy. Relatively very few chromosome numbers of Lindsaeoid ferns are known at present. 
Moreover, several of them are only approximately known, and in some cases the identity of the plants 
is uncertain, as can be concluded from the names under which they have been reported. 

In sect. Schizoloma there are counts of n = 88 for Lindsaea ensifolia ssp. ensifolia (Manton & Sledge) 
and for L. 'tenera (prob. L. orbicidata var. commixta) (Manton), c. 88 for L. viridis (Brownlie), 44 or 
45 for L. ensifolia ssp. coriacea (Manton ex Holttum; see Kramer, 1968), c. 42 for L. trichomanoides 
(as 'cuneata) (Brownlie), c. 40 for L. prolongata (Brownlie), and c. 47 for L. vieillardii (Brownlie) 
and L. chietui (Kurita). It seems likely that most, if not all, of these numbers are 44 or twice as many. 
This was also found in other sections; in sect. Odontoloxia there is a count of n = c. 44 for L. repens var. 
sessilis (Walker, in litt.) and 44 or 45 for L. pidchella var. blanda (Walker, in litt.), in sect. Lindsaea 
n = 88 in L.portoricensis (neotropical) (Walker) and c. 84 inZ,. arcuata (neotropical) (Mickel, Wagner 
& LiM Chen). Other numbers may or may not be related, e.g. L. 'nitida" (= integral) (Manton in 
Kramer) and L. 'scandens var. terrestris' (= L. doryphora) (Manton in Kramer), both c. Al. The 
number n = 47 has, however, been found unequivocally in certain species, e.g. L. 'macraeana'' (= L. 
repens var.) (Wagner), L. 'concinna" (Australia; L. brachypodal) (Manton in Kramer) and L. parallelo- 
gramma (Manton in Kramer). The numbers c. 50 and c. 100 reported for L. decomposita (= L. obtusal) 
(Manton in Holttum), from the same section as L. parallelogramma, may have been 47 and 94, re- 
spectively. L. 'pectinata' (prob. L. oblanceolata) was also found to have c. 50 chromosomes (Manton 
in Holttum), but this species is closely related to L. repens (c. 44, see above). In L. odorata, placed in 
sect. Osmolindsaea, divergent in its monolete spores but not otherwise very distinct, a polyploid series 
was found, ranging from n = 150 (Mehra & Khanna) or 150-152 (Walker, in litt.), to c. 220 (Walker, 
in litt.); the report of 82 (Manton in Kramer) from Ceylon may be due to misidentification of the plant. 
A basic number of 50 or 51 for this species seems possible. L. linearis, a member of the distinct sect. 
Paralindsaea, was found to have n = 34 (Brownlie). 

The picture is equally confusing in the related but much smaller genus Sphenomeris. In S. chinensis, 
by far the most widespread species, there are reports of n = 94 (Mehra & Khanna; Kurita & Nishida), 
c. 100 (Bir; Manton & Sledge), and 145, 146, 147 (Manton & Sledge). Its close relative S. biflora 
was counted as n = 48 (Kurita & Nishida). S. retusa had n = 88 and c. 88 (Walker, in litt.); another 
specimen, apparently of hybrid origin, with abortive spores, had 162-164 chromosomes, with univalents 
(Walker, in litt.). Two counts of n = 38 and 39, respectively, for the New World S. clavata (Walker; 
Wagner) are again divergent. It has been suggested that Sphenomeris is not a natural genus (Wagner, 
Am. Fern J. 53, 1963, 4), but morphological data do not seem to support this. 

Two species of Odontosoria from Jamaica have been counted as c. 96 (Walker). No counts for Tapeini- 
dium or Xyropteris have been found in the literature. 

It seems that the numbers 44 and 47 are widespread in the group, and that some counts of approxi- 
mately one of these numbers are equal to them or have been derived from them. It is also certain that 
one or more divergent basic numbers occur besides. Differences in basic number are, however, not 
necessarily connected with considerable morphological ones. It may be hoped that a clearer picture 
emerges when more data are available and that then some more light may be shed on the affinities in the 
group. 

Principal sources of data: Bir, Curr. Sci. 31 (1926) 248; Brownlie, Trans. R. Soc. New Zeal. 85 
(1958) 213; ibid. Bot. 1 (1961) 1; Pac. Sci. 19 (1965) 4; Kurita & Nishida, J. Jap. Bot. 38 (1963) 4; 
Manton in Holttum, Rev. Fl. Mai. 2 (1954) 623; Manton in Kramer, Act. Bot. Neerl. 6 (1957) 108; 
Manton & Sledge, Phil. Trans. R. Soc. Lond. B 238 (1954) 127; Mehra & Khanna, J. Genet. 56 (1959) 
296; Mickel, Wagner & Lim Chen, Caryologia 19 (1966) 95; Walker, Trans. R. Soc. Edinb. 66 
(1966) 169. 

Taxonomy. In the older literature the Lindsaea-group is usually associated with, or included in, the 
Davallioid ferns, even as late as 1928 by Perez Arbelaez {I.e.), although he noted the great difterences 
in scale and rhizome structure, spore morphology, etc., between the two groups. More recent authors 
have tended to emphasize the differences between them and have placed the Lindsaeoids in a separate 
family (Ching, Sunyatsenia 5, 1940, 216), associated them with the very broadly defined Pteris-group 
(Copeland, Gen. Fil. 1947), or placed them in Dennstaedtiaceae. In the present Flora, the question of 
formal delimitation of families has been left open (Holttum, Fl. Mai. II, 1, 1959, I-II); therefore no 
formal status is here proposed for the Lindsaea group of genera. I would, however, express the opinion 
that, though the group is a very natural one, its separation as a distinct family does not seem warranted, 
in view of its many similarities to Dennstaedtia and allied genera. 

As expounded in the revision of the American Lindsaeoids (Kramer, 1957, I.e.), and in the chapter on 
the classification of the Malesian representatives (Kramer, Blumea 15, 1968, 557 seq.), the leaf pattern, 
greatly and excessively used in the past, is by itself insufficient as a basis for generic classification. Such 
genera as have been based entirely on characters of leaf architecture and venation: Schizoloma, Iso- 
lonia, Synaphlebium, are here merged with Lindsaea. Comments on the circumscription of Sphenomeris 
and Tapeinidium can be found in the above-cited papers (Kramer, 1957, 1968, I.e.; Act. Bot. Neerl. 15, 
1967, 562). 



April 1971] LiNDSAEA-GROUP (Kramer) 179 

KEY TO THE GENERA 

1. Sori on 1-8 vein-ends; indusium laterally entirely or largely adnate to the lamina; ultimate divisons 

never dimidiate; veins free. 
2. Ultimate free divisions not of a linear- or cuneate-divaricate type, subentire to pinnatifid; sori on 
the lateral margin of the divisions or in their lobes; pluricellular filiform paraphyses usually (al- 
ways?) present; spores monolete 2. Tapeinidium 

2. Ultimate free or nearly free divisions of a linear- or cuneate-divaricate type, with the sorus (sori) 
on their apical margin; paraphyses 2- or 3-celled, not usually found; spores monolete or tri- 
lete 1. Sphenomeris 

1. Sori on many vein-ends, or, if on 8 or fewer, the sides of the indusium free, or the pinnules dimidiate, 
or the veins anastomosing, or these characters combined. 

3. Ultimate free divisions lanceolate, equal-sided or the base anteriorly auricied; veins free; spores 
monolete 3. Xyropteris 

3. Ultimate free divisions not lanceolate and equal-sided, or, if so, the spores trilete; veins free or 
anastomosing 4. Lindsaea 

1. SPHENOMERIS 

MAXON,J.Wash.Ac.Sc.3(1913)144, /?om. cons.; Copeland, Gen. Fil. (1947) 54; 
Philip. J. Sc. 78 (1949) 24; Holttum, Rev. Fl. Mai. 2 (1954) 340; Kramer, Act. 
Bot. Neerl. 6 (1957) 152; Copeland, Fern Fl. Philip. 1 (1958) \\5.—Da\allia 
J. E. Smith, Mem. Ac. Turin 5 (1793) 414; Hooker, Sp. Fil. 1 (1845) 151; and 
of many other authors; all in part. — Stenoloma Fee, Gen. Fil. (1852) 330, p.p. 
min.; Ching, Fl. Reip. Pop, Sin. 2 (1959) 275, in part; and of other authors. — 
Odontosoria Fee, Gen. Fil. (1852) 325; J. Smith, Hist. Fil. (1875) 263; Diels in 
E. 8l p. Nat. Pfl. Fam. I, 4 (1902) 215; v.A.v.R. Handb. (1908) 258; Suppl. (1917) 
202; and of many other authors; all in part. 

Terrestrial ferns with a short- to moderately long-creeping rhizome, if stout 
with a solenostele with a sclerotic medullary strand, if more slender mostly with a 
lindsaeoid protostele. Rhizome scales elongate-triangular to acicular, in the 
smaller species some scales wholly uniseriate and therefore the scales grading 
into hairs. Petioles abaxially terete, adaxially upward sulcate. Lamina much 
dissected, decompound, strongly anadromic, without a conform terminal pinna, 
the ultimate divisions confluent near the pinna-apices, not free and conform 
(except in the New Caledonian S. alutacea). Veins free, simple or forked in the 
ultimate divisions. Sori uni-to paucinerval, on the apical margin of the segments; 
indusium attached at the base and the sides. Paraphyses 2- or 3-celled, observed 
in one species {S. chinensis), presumably present in all and fugacious. Spores 
monolete or trilete. Gametophyte undescribed. 

Type species: Sphenomeris clavata (L.) Maxon (tropical America). 

Distr. In the tropics and in the northern subtropical regions of both hemispheres 1 1 spp., 6 of them 
with very small areas. 

Note. Sphenomeris Maxon was conserved against Stenoloma Fee, a name of somewhat controversial 
application. In the Code of Nomenclature Stenoloma dumosum Fee was at first designated as type species, 
but afterwards, on Morton's suggestion (Taxon 8, 1959, 29), this was changed to Stenoloma clavatum 
(L.) Fee, as this was said to agree much better with the original description of the genus. However, 
Fee explicitly mentioned as an essential character of Stenoloma, setting it apart from Odontosoria, the 
only basally attached but laterally free indusium. This is not found in any true species oi Sphenomeris, 
and certainly not in Sphenomeris clavata. The issue should be reconsidered. 

key to the species 
1. Rhizome 5-20 mm 0, solenostelic, with an internal sclerotic strand; rhizome scales to 7 mm long, the 

larger ones to c. 20-seriate at the base; spores trilete; sori of larger segments on (2-)5-8 vein-ends, 

occupying their whole apical margin (fig. 5) 1. S. retusa 

1. Rhizome 2-4 mm 0, with a lindsaeoid protostele; scales to 4 mm long, to 4-seriate at the base (wider 

in S. biflord); spores monolete; sori of larger segments on l-2(-4) vein-ends, not occupying their 

whole apical margin. 



180 



Flora Malesiana 



[ser. II, vol. 1^ 



2. Scales up to 3-seriate at the base; sori strictly uninerval; ultimate divisions %-% nim wide, or the 
fertile gradually widened to the sorus and there 1 mm wide; texture coriaceous . . 4. S. veitchii 

2. Scales to 5-6-seriate at the gradually widened base; sori uni- or binerval; larger free ultimate 
divisions c. 2 mm wide; lamina subcoriaceous or coriaceous, usually bipinnate — pinnatifid or tripin- 
nate -p crenate 2. S. biflora 

2. Scales 1-3-seriate (or to 4-seriate at the suddenly broadened base); sori l-3(-4)-nerval; larger free 
ultimate divisions 1 mm or more wide, or, if narrower, spathulately widened at the sorus; lamina 
herbaceous to subcoriaceous, in full-grown plants bipinnate -i- bipinnatifid, tripinnate — pinnat- 
ifid, or more dissected 3. S. chinensis 



1. Sphenomeris retusa (Cav.) Maxon, J. Wash. 
Ac. Sc. 3 (1913) 144; Copeland, Philip. J. Sc. 78 
(1949) 24; Fern Fl. Philip. 1 (1958) \\6.-Da- 
vallia retusa Cav. Descr. (1802) 278. — Stenoloma 
retusiun (Cav.) Fee, Gen. Fil. (1852) 330. -Lind- 
saea retusa (Cav.) Mett. Fil. Lips. (1856) 
\05. — Odontosoria retusa (Cav.) J. Smith, Bot. 
Voy. Herald (1857) 430. — Schizoloina retusum 
(Cav.) Kuhn, Chaetopt. (1882) 346. - Type: 
Nee s.n., Mt Isarrog, Luzon (MA, n.v.; phot. U). 

Lindsaea cuneifolia Presl, Rel. Haenk. 1 (1825) 
60. — ? Saccoloma cuneifoUum Presl, Tent. Pterid. 
(1836) 126. — Acrophorus cuneifolius (Presl) 
Moore, Ind. Fil. (1857) 41. - Type: Haenke 
s.n., Luzon iri.v.). 

? Adiantum falcatum Blanco, Fl. Filip. (1837) 
833. — Type: Blanco s.n., 'Mandalogon' («.v.; 
perhaps this species ace. to C. Chr. Ind. Fil. 
Suppl. 1, 1913, corr. 89). 

Davallia decipiens Cesati, Rendic. R. Accad. 
Sci. Fis. Mat. Napoli 16 (1877) 29. -Odonto- 
soria decipiens (Cesati) Christ, Nova Guinea 8 
(1909) 158. — Type: Beccari s.n., Mt Arfak at 
Putat, W. New Guinea (Fl, 2 sh.). 

Odontosoria lindsayae v.A.v.R. Bull. Dep. Agr. 
Ind. Neerl. 21 (1908) 4.-Type: not cited; 3 sh. 
so annotated in BO; on the ground of the date 
chosen as lectotype: Versteeg 1467, s.w. W. 
New Guinea, prob. back of Sabang (BO; dupl. 
in B, K, L, U). - Fig. 5. 

Rhizome probably short-creeping (only short 
pieces seen), enveloped in a dense mass of roots, 
stout, in full-grown plants ^2-2 cm 0, with an 
internal sclerotic strand; scales dark reddish 
brown, elongate-triangular, to 7 mm long, to c. 
20-seriate at base, a short apical portion uniseri- 
ate. Petioles stramineous or darker with age, 
the base often slightly verrucose from scale-bases, 
2-15 mm at base, 20-70 cm long (on labels 
said to attain 1.6 m), c. 73-I times as long as the 
lamina; all axes abaxially rounded, marginate near 
the ultimate divisions. Lamina oblong-trian- 
gular, to 3 m long, mostly brown or olivaceous 
when dry, chartaceous to coriaceous, sub- 
tripinnate to tripinnate + pinnatifid, or less 
often quadripinnate. Major pinnae 5-12 to a 
side, alternate or the lower ones subopposite, 
obliquely ascending, the lower ones with a 
stalk of 1-2 cm, the upper ones gradually sub- 
sessile, elongate-triangular, to 30 by 12 cm, 
caudate-acuminate, equal-sided but the larger 
pinnae basiscopically in their basal part often 
slightly narrower. Lower pinnae their width apart 
or more, the upper ones usually somewhat over- 
lapping. Larger secondary pinnae narrowly 



triangular, shortly petiolulate, acuminate, c. 5-10 
cm long, 25/2-6 cm wide, pinnate -i- pinnatifid 
or less often bipinnate, basiscopically narrower, 
with often 3 or 4 free pinnules to a side, the 
larger ones pinnatifid or less often pinnate, the 
upper ones cuneate, confluent; upper primary 
and distal secondary pinnae rhombic, subdi- 
midiate, basiscopically more narrowly cuneate. 
Apices of primary and secondary, sometimes 
also of tertiary pinnae narrow, serrate, ± cau- 
diform. Ultimate pinnules, except if transitional 
between pinnate and non-pinnate ones, subrhom- 
bic and with a few very shallow incisions, or 
cuneate and entire, often 7-15 mm long and 
5-10 mm wide, P2-2 times as long as wide 
(upper, smaller ones relatively narrower), evenly 
cuneate from the base, widest at the truncate 
or just below the triangular apex, with straight 
or slightly convex, sometimes subrevolute lateral 
edges. The quadripinnate form with narrower 
pinnules, c. 5-6 by 2 mm, 2^4-3 times as long 
as wide. Sterile pinnules (hardly found in full- 
grown plants) crenate-cleft. Veins adaxially 
impressed, abaxially prominulous, usually twice 
forked; close, c. i mm apart. Sori continuous 
across the whole apical margin of cuneate, interrup- 
ted in subrhombic pinnules and then continuous 
across the separate straight portions of their apical 
margin, on 5-8 vein-ends and to 5 mm long, 
in smaller segments or lobes shorter, in the 
quadripinnate form often bi- to quadrinerval. 
Indusium pale to brown, entire, the slightly 
narrowed sides adnate, rather thin, 0.4—0.6 mm 
wide, almost reaching the margin, bulging but 
scarcely reflexed at maturity. Spores medium 
brown, trilete, almost smooth, c. 38-40 it. 

Distr. Malesia: Celebes, Philippines (almost 
throughout), Moluccas (Morotai, Halmahera, 
Tidore, Ambon, Ceram), New Guinea, Admiralty 
Is., Bismarck Arch.; Solomon Is.,?New Hebrides. 

Ecol. On banks, in open places, often among 
rocks by rivers, less often in forests, from sea 
level to c. 2000 m. 

Note. A quadripinnate form with narrower 
pinnules and shorter sori, as described above, 
has been collected several times in New Guinea 
and also in Manus. It is not sharply distinct from 
the broader, less dissected form. Its status is 
uncertain; one specimen had an irregular meiosis 
(Walker, pers. comm.) and proved to have 
abortive spores; it is almost certainly a hybrid 
(with S. chinensisl), but this is scarcely the 
case with all. 



April 1971] 



LiNDSAEA-GROUP (Kramer) 



181 




Fig. 1-3. Sphenomeris chinensis (L.) Maxon. — Fig. 1. var. divaricata (Christ) Kramer. Pinnule, X 4 
(Rant 21). — Fig. 2. var. rheophila Kramer. Pinnule, x 4 (Surbeck 315). — Fig. 3. var. chinensis. Pin- 
nule, X 4 (Bartlett & DE LA RuE 77). -Fig. 4. S. veitchii (Baker) C. Chr. Pinna, x 11/4 (Clemens 
50992). -F'g. 5. S. retusa (Cav.) Maxon. Pinnule, X \% (Koorders 17033). 



182 



Flora Malesiana 



[ser. II, vol. P 



2. Sphenomeris biflora (Kaulfuss) Tagawa, J. 
Jap. Bot. 33 (1958) 203; Kramer, Blumea 15 
(1968) 573. — Davallia biflora Kaulfuss, Enum. 
(1824) 22\. — Microlepia biflora (Kaulfuss) Mett. 
Fil. Lips. (1856) 104. — Odontosoria biflora 
(Kaulfuss) C. Chr. Ind. Fil. (1906) 464. - 
Stenoloma biflorum (Kaulfuss) Ching, Sinensia 3 
(1933) 338; Tagawa, J. Jap. Bot. 22 (1948) 
160. — Type: Chamisso s.n., Manila, Luzon (B). 

Davallia tenuifolia Swartz var. lata Hooker 
ex Moore, Ind. Fil. 2 (1861) 301, based on /9 
(unnamed) of Hooker, Sp. Fil. 1 (1845) 186.- 
Lectotype: Exp. Acad. Petersb. 44, Bonin (K). 

Odontosoria tsoongii Ching, Bull. Fan Mem. 
Inst. Biol. 1 (1930) 149. - Lectotype: Tsoong 
1423, Hailin Is., Kwangtung, China («.v.). 

Stenoloma littorale Tagawa, Act. Phytotax. 
Geobot. 6 (1937) 225. — Stenoloma cliusanum 
(L.) Ching var. littorale (Tagawa) Ito, Bot. 
Mag. Tokyo 52 (1938) 6. — Sphenomeris chusana 
(L.) CoPELAND var. littoralis (Tagawa) H. Ito 
ex MizusHiMA, Misc. Rep. Inst. Natur. Res. 38 
(1955) 115 (quoted by Tagawa, Col. III. Jap. 
Pterid. 1959, 256, n.v.) — Type: Tasiro s.n., 
Oshima I., Shikoku (KYO, n.v.). 

Sphenomeris chinensis or chusana, etc., of 
various authors, in part, e.g. Copeland, Fern 
Fl. Philip. 1 (1958) 115. 

Rhizome short-creeping, 2-3 mm 0; scales gold- 
en to medium brown, occasionally castaneous, 
to 3^2 mm long, in the larger ones the apical half 
or less uniseriate, acicular, the lower part 
gradually broadened to the 5-6-seriate (excep- 
tionally even broader) base. Leaves clustered; 
petioles stramineous to pale brown, dull, abaxially 
terete, or flattened and obtusely bi-angular, 
adaxially sulcate, c. 10-25 cm long, almost 
equaling to about half as long as the lamina. 
Lamina oblong or less often triangular or sub- 
pentagonal, c. 15-35 cm long, subcoriaceous or 
more often coriaceous, olivaceous when dry, 
the upper side often blackish, bipinnate + pinnat- 
ifid or tripinnate + pinnatifid, with 7-12 major 
pinnae to a side; primary rachis abaxially sub- 
terete or usually upward ± flattened and 
gradually marginate. Primary pinnae elongate- 
triangular or narrowly triangular, obliquely 
ascending, or in small leaves spreading, with 
a petiolule of a few mm, subacute to acuminate, 
the lower ones about their width apart, 
the upper ones closer, the larger ones c. 5 by 
1 V2 to 13 by 5 cm, with 3-6 pinnate secondary 
pinnae, and some subentire to pinnatifid upper 
ones, to a side, the apex pinnatifid-serrate; 
secondary rachises abaxially rounded, at least 
upward marginate. Upper pinnae gradually 
reduced, sometimes the basal pair slightly shorter 
than the next. Lower pinnae in their basal part 
usually more compound than the rest of the 
lamina, the basal basiscopic secondary pinnae 
sometimes larger than all the others. Axes of 
higher order often ± flexuous. Smaller pinnules 
rhombic, pinnatifid + pinnatisect or forked, the 
smallest entire, obtriangular-cuneate. Largest 
free or nearly free non-dissected pinnules 



(segments) often 5 by 2 mm, asymmetric, 
decurrent, with straight or more often upward 
rounded sides, usually widest just below the 
apex, the apical margin subtruncate, erose to 
shallowly crenate, or subentire, mostly with two 
small latero-apical projections, the thickened 
lateral margins often ± revolute. Veins immersed 
or usually slightly prominulous on both sides, 
forked, the ultimate segments, except the smallest, 
with two to several veins. Sori one or not rarely 
two per segment, or, if more, separated by inci- 
sions, uni- or occasionally binerval; indusium 
elliptic, very convex at the base, adnate at the 
± convex sides, rigid, the free margin rounded, 
subtruncate, ± equaling the margin, or not rarely 
with lobes exceeding the margin, 1 mm wide, 
/i-l Vi rrirri long (at right angles to vein). Spores 
monolete, ellipsoid, medium brown, smooth, 
c. 32-35 by 40^5 /i. 

Distr. S. Japan, Bonin Is., Guam, Hong 
Kong and other islands on the Chinese S. coast; 
in Malesia: only in Luzon and the Batanes Is. 

Ecol. Very few data from Malesia; elsewhere 
in ± exposed places, often by the sea, never in 
forests; at lower and middle elevations, to c. 
1300 m. 

Note. The more dissected forms of S. biflora 
can be safely distinguished from the larger, 
broader forms of S. chinensis var. chinensis 
only by the rhizome scales. 

3. Sphenomeris chinensis (L.) Maxon, J. Wash. 
Ac. Sc. 3 (1913) 144; Contr. U.S. Nat. Herb. 
17 (1913) 159 (in both places the basionym 
incorrectly cited as Adiantum chinense L.); C. 
Chr. Ind. Fil. Suppl. 2 (1917) 31; Kramer, 
Act. Bot. Neerl. 15 (1967) 565; Blumea 15 
(1968) 572; Fosberg, Taxon 18 (1969) 596.- 
Trichomanes chinense L. Sp. PI. 2 (1753) 1099.- 
Adiantum chinense (L.) Burman, Fl. Ind. (1768) 
236 (pxoh.). — Davallia chinensis (L ) J. E. Smith, 
Mem. Ac. Turin 5 (1793) 414. — Microlepia 
chinensis (L.) Mett. Fil. Lips. (1856) 104. - 
Odontosoria chinensis (L.) J. Smith, Bot. Voy. 
Herald (1857) 430.- Davallia tenuifolia (Lamk) 
Swartz var. chinensis (L.) Moore, Ind. Fil. 
2 (1861) 302. — Lindsaea chinensis (L.) Mett. 
ex KuHN, Fil. Afr. (1868) 67, non Ching (1929).- 
Stenoloma chinense (L.) Bedd. Handb. Ferns 
Br. Ind. (1883) 70. — Type: Osbeck s.n., China 
(S-PA). 

Adiantum chusanum L. Sp. PI. 2 (1753) 1095.- 
Davallia chusana (L.) Willd. Sp. PI. 5 (1810) 
475. — Sphenomeris chusana (L.) Copeland, 
Bull. Bish. Mus. 59 (1929) 69; Philip. J. Sc. 78 
(1949) 24; Holttum, Rev. Fl. Mai. 2 (1954) 
341; Copeland, Fern Fl. Philip. 1 (1958) 115. - 
Odontosoria chusana (L.) Masam. Mem. Fac. 
Sci. Agr. Taihoku Imp. Univ. II (1934) 67.- 
Type: coll.?, China {n.v.). 

Adiantum tenuifolium Lamk, Encycl. 1 (1783) 
44. — Davallia tenuifolia (Lamk) Swartz in 
Schrader, J. Bot. 1800^ (1801) 88. - Stenoloma 
tenuifolium (Lamk) Fee, Gen. Fil. (1852) 330.- 
Microlepia tenuifolia (Lamk) Mett. Fil. Lips. 



April 1971 



LiNDSAEA-GROUP (Kramer) 



183 



(1856) 104, pi. 27, f. 1^, non Presl (1851).- 
Odontosoria tenuifolia (Lamk) J. Smith, Cat. 
Cult. Ferns (1857) 67. — Odontosoria chinensis 
(L.) J. Smith var. tenuifolia (La)mk Matsum. 
Ind. PI. Jap. (1904) 230. — Sphenunieris chinensis 
(L.) Ma.xon var. tenuifolia (Lamk) C. Chr. 
Dansk Bot. Ark. 7 (1932) 78. - Odontosoria 
chusana (L.) Masam. var. tenuifolia (Lamk) 
(err. 'Mar.') Masam. Mem. Fac. Sci. Agr. 
Taihoku Imp. Univ. 11 (1934) 67. — Sphenomeris 
chusana (L.) Copeland var. tenuifolia (Lamk) 
HoLTTUM (incorr. ascr. to C. Chr.), Rev. Fl. Mai. 
2 (1954) 341, as to type only. — Type: Sonnerat 
s.n., 'Inde' (P). 

Davallia didyma Hedwig, Fil. Gen. & Sp. 
(1803) pi. 22, ex icon. — Type: not cited. 

Davallia microcarpa J. E. Smith in Rees, 
Cyclop. 11 (1808) sine pag. — Type: Chr. Smith 
s.n., Amboina (LINN). 

Hvmenophyllum ramosissimum Ham. ex D. 
Don, Prod. Fl. Nepal. (1825) 12. - Type: 
Hamilton s.n., Nilkantha, Nepal {n.v.; identity 
teste C. Chr. Ind. Fil.). 

Trichomanes malavanum RoxB. ex Griff. 
Calc. J. Nat. Hist. 4 (1844) 519. -Type: coll.? 
'native of the Malay Islands' {n.v.; identity 
uncertain). — Fig. 1-3. 

Rhizome short-creeping, 2-4 mm 0; scales 
reddish brown to castaneous, acicular, entirely 
uniseriate, or the base often biseriate, less often 
the extreme base tri- or quadriseriate, to 4 mm 
long. Leaves clustered; petioles stramineous with 
darker base, or darker with age, abaxially terete, 
adaxially upward gradually sulcate, the groove 
broad and flat; petioles of full-grown plants 
1^/2-3 mm in the middle, c. 12-60 cm long, 
73-I times as long as the lamina, shorter and 
more slender in juvenile but fertile plants. Lamina 
oblong, elongate-ovate, or narrowly triangular, 
15-85 cm long (rarely smaller in fertile plants), 
usually olivaceous, medium or dark brown to 
blackish when dry, herbaceous to chartaceous or 
occasionally subcoriaceous, if fertile at the base at 
least bipinnate — pinnatifid, usually bipinnate — 
bipinnatifid or tripinnate — pinnatifid, or in large 
specimens up to quadripinnate - pinnatifid. 
Primary rachis and axes of higher order stramine- 
ous, abaxially terete, upward gradually marginate. 
Larger leaves with c. 6-10 major primary pinnae 
to a side; pinnae rather strongly spreading to 
strongly ascending, elongate-triangular or -rhom- 
bic, with a stalk of a few mm to 3 cm, the base 
usually inequilateral, the anterior side broader 
as the pinna is strongly anadromic and the 
anterior side has longer and/or less ascending 
pinnules, the apex acuminate; larger pinnae (5-)- 
10-20 cm long, (1 U-)3-10 cm wide, 2^^,z times 
as long as wide, the lower ones usually subopposite 
and several cm apart, the upper ones gradually 
alternate, smaller, and closer. Secondary pinnae 
triangular or rhombic, acute or acuminate, as- 
cending, shortly petiolulate, alternate, often c. 
6-8 major ones to a side, often twice as long 
as wide, size and dissection depending upon 
size and dissection of the lamina and their place 



in it. Largest ultimate free divisions rhombic, 
asymmetric, pinnatisect on both sides, smaller 
ones cuneate, unequally and shallowjy bifid, 
or, if deeply bifid, usually once again bifid, linear- 
spathulate to cuneate (depending on the variety; 
see below). Veins immersed, in dry material usu- 
ally little or not evident. Ultimate lobes uni- 
or binerval, or in broader forms occasionally 
to quadrinerval, Yo-l mm wide at the apex, the 
larger undivided ones 2-3 mm long, the ones 
below them incised, the ones above them reduced, 
denticuliform, confluent into a pinnatifid, often 
caudate-acuminate pinna-fpinnula-Japex. Sterile 
lobes apically subacute, or, if broader, often 
denticulate. Fertile lobes with straight or slightly 
convex apical margin, broader ones not rarely 
erose-denticulate. Sori on 1 or 2, less often on 
3, exceptionally on 4 vein-ends, not quite reaching 
the apico-lateral extremities of the lobes; indusi- 
um brownish and chartaceous when dry, with 
— straight to convex base, adnate, convex sides, 
and straight, slightly convex, or sometimes erose- 
denticulate free margin, :r equaling or, if denti- 
culate, sometimes slightly exceeding the margin, 
never reflexed at maturity. Spores monolete, 
ellipsoid, smooth, medium brown. 

Distr. Throughout the tropical and subtropical 
parts of the Old World, but wanting in continental 
Africa. 

1. var. rheophila Kramer, Blumea 15 (1968) 573. — 
Type: Bartlett 6718a, Sumatra, Asahan, water- 
fall of Asahan R. (L; dupl. in GH, MICH, 
S-PA, US). -Fig. 2. 

Lamina 15-20 cm long, tripinnate ~ pinnatifid 
or less often bipinnate — bipinnatifid at the base; 
segments rigid, often with — revolute margin, 
narrowly cuneate, gradually broadened to base, 
most of them monosorous and 4-5 times as long 
as wide, often 4-5 by 1 mm. Outer edge of seg- 
ments entire or sinuate. Sori on one, kss often 
on two (mostly connivent) vein-ends; indusium 
often ^2 by I2 irini, with convex base. Spores as 
in var. chinensis, c. 44-48 fi long, but more 
elongate, bean-shaped, — twice as long as broad. 

Distr. Malesia: Pahang and Central Sumatra 
(9 coll.). 

Ecol. By torrents and waterfalls, on river- 
banks, 100-500 m. 

2. var. divaricata (Christ) Kramer, Blumea 15 
(1968) 511. — Odontosoria chinensis (L.) J. 
Smith var. divaricata Christ, Journ. de Bot. 
ser. 2, II (1909) 23. — Sphenomeris chusana (L.) 
Copeland var. divaricata (Christ) Tardieu- 
Blot, Fl. Madag. Com. 5e fam. I (1958) 29.- 
Type: Chevalier 14309, Sao Tom6 (P). 

Sphenomeris chusana (L.) Copeland var. 
tenuifolia of HoLTTU.M, Rev. Fl. Mai. 2 (1954) 
341, and of other authors; not Adiantum tenui- 
folium Lamk. — Fig. 1. 

Lamina usually over 20 cm long, at the base 
often tripinnate — bipinnatifid; segments cuneate, 
suddenly spathulate-broadened at the sorus, 
slightly narrowed at the rounded apex, the apical 
margin not rarely erose, the sides often corn- 



84 



Flora Malesiana 



[ser. II, vol. F 



iculate; at the base often ^2 rnrn wide, slightly 
broadened to the apex, 1-1^4 mm wide at the 
sorus, of varying length; sori not rarely two to- 
gether in a segment, mostly uninerval, if binerval 
mostly on two connivent vein-ends; indusium even 
in binerval sori with distinctly convex base. 
Spores mostly 55-60 f^i long. 

Distr. Sao Tome; sporadically from Sikkim 
and S. China to Malesia: throughout Malesia, 
common in Sumatra, Java, and New Guinea, 
rare in Borneo (Sarawak only) and local in the 
Philippines (Luzon, Mindanao). 

E c o 1. As the next variety, but mostly above 700 m. 

Note. See the note at the end of the species. 

3. var. chinensis. — In so far as known most or 
all of the synonyms enumerated under the 
species apply to this variety. — Fig. 3. 

Lamina of variable length, at the base often 
tripinnate + pinnatifid; segments cuneate, grad- 
ually broadened from the base, rarely with one 
sorus only, except the upper, reduced ones, usu- 
ally with several sori and shallow incisions 
between them (not entire and with one sorus 
across the apical margin, as in S. retusa), often 
— twice as long as wide, the apical margin not 
or scarcely erose; sori not rarely uninerval, most 
often bi- or tri-, rarely to quadrinerval, to 2 mm 
long, most often %-l ^o rnrn long. Spores mostly 
42-48 1^1 long. 

Distr. As the species. 

Ecol. Terrestrial and on rocks, in thickets 
and open forests, in exposed or lightly shaded 
places, often by hollow roads, on slopes, escarp- 
ments, by streams, etc., 100-2400 m, apparently 
most common from c. 800-1500 m, often said to 
be locally numerous. 

Notes. The three varieties of S. chinensis, 
notably the last two, are not quite sharply distinct; 
this is one reason for treating them as varieties. 
Intermediates between var. chinensis and var. 
divaricata occur not rarely, but are much less 
numerous than typical specimens. The two 
varieties overlap throughout Malesia, only in 
New Guinea the former is quite rare and the latter 
relatively frequent. On the continent var. divari- 
cata becomes increasingly rare to the North and 
West; it is absent from Japan and nearly so from 
China. In the Pacific it has not been found so far. 
However, on many islands a form occurs with 
narrower, mostly monosoral segments and 
prevailingly uninerval sori. It lacks the spathulate 
segments of var. divaricata and is therefore 
regarded as an aberrant form of var. chinensis. 
Similar specimens occur here and there in Malesia, 
too, notably in Celebes. Their taxonomic status 



cannot be elucidated with the help of dried 
material only. A form with exceptionally broad 
segments occurs in Sumatra. It has been confused 
with S. retusa and S. biflora, but must be assigned 
to S. chinensis because of its monolete spores 
and its narrow rhizome scales, and some other 
characters, too. 

4. Sphenomeris veitchii (Baker) C. Chr. Gard. 

Bull. S. S. 7 (1934) 234.~-Davallia veitchii Baker, 
J. Bot. 17 (1879) 29.Stenoloma veitchii (Baker) 
C. Chr. Ind. Fil. Suppl. 3 (1934) 174.— Type: 
Burbidge 49 or s.n., Mt Kinabalu, Sabah (K, 
2 sh.; dupl. in BM). — Fig. 4. 

Rhizome rather long-creeping, c. 4 mm 0; 
scales dark castaneous, acicular, the upper half 
uniseriate, the base bi- to triseriate, to 2V2 mm 
long. Leaves ^2-1 cm apart; petioles slender, 
1 V2-2 mm 0, fuscous or upward stramineous, 
abaxially terete, adaxially broadly and shallowly 
grooved, 30^0 cm long. Lamina narrowly 
oblong, c. 30 by 5 to 70 by 15 cm, bipinnate -r 
bipinnatifid or tripinnate — bipinnatifid, with c. 
10-20 primary pinnae to a side; primary rachis 
like the petiole, upward paler and sometimes very 
obtusely bi-angular. Basal pinnae remote, to 
12 cm apart, the upper ones gradually closer, 
subcontiguous, all ascending, elongate-triangular, 
acuminate, with a stalk of up to 1 cm, pronounc- 
edly anadromic, the larger ones c. 10-12 by 4-5 
cm, the upper ones gradually smaller; major se- 
condary pinnae c. 6-8 to a side in larger pinnae, 
rhombic, acute. Axes of secondary and higher 
order stramineous, adaxially narrowly and 
deeply sulcate, abaxially rounded or narrowed- 
rounded, upward gradually somewhat flexuous 
and marginate, therefore in the lamina the pinnate 
grading into the pinnatifid condition. Ultimate 
segments rigid, coriaceous, olivaceous when dry, 
basally often with adaxially raised, thick edges, 
linear, often 4—5 by 0.5-0.7 mm, the fertile 
ones gradually to 1 mm wide at the sorus, sub- 
obtuse to erose-truncate, the sterile ones acute, 
the larger ones forked. Veins strictly single and 
undivided in the lobes, immersed, scarcely 
visible. Sori uninerval; indusium rigid, yellowish, 
subelliptic, with adnate sides, not reaching the 
lateral margins of its segment, c. ^4-% mm wide, 
V2 mm long (at right angles to its vein), 4- equal- 
ing the often laterally shortly bicorniculate outer 
margin of the segment, subentire to shallowly 
erose. Spores monolete, subellipsoid, light brown, 
smooth, c. 45 by 38 ^. 

Distr. Malesia: confined to Mt Kinabalu, 
Sabah, Borneo (3 coll.). 

Ecol. In mountain forests, 2000-2500 m. 



2. TAPEINIDIUM 



(Presl) C. Chr. Ind. Fil. (1906) 631; Copeland, Gen. Fil. (1947) 53; Holttum, 
Rev. Fl. Mai. 2 (1954) 338; Kramer, Blumea 15 (1968) 545.— Microlepia sect. 
Tapeinidium Presl, Epimel. Bot. (1851) 96S. —Protolindsaya Copeland, Philip. 
J. Sc. 5 (1910) Bot. 2S3.— Wibelia auct. non Bernhardi; Fee, Gen. Fil. (1852) 



April 1971] LiNDSAEA-GROUP (Kramer) 185 

331; DiELS, in E. & P. Nat. Pfl. Fam. I, 4 (1902) 2l6.—Davallia or Microlepia 
auctt. plur., p.p. 

Small to medium-sized terrestrial ferns with very short to moderately long- 
creeping rhizome with at least in the larger species a true solenostele with external 
and internal endodermis and a medullary strand of sclerenchyma. Scales long 
and narrow, glabrous, non-clathrate. Lamina up to the last divisions pinnately 
compound, at least once pinnate; ultimate divisions not dichotomously divaricate. 
Veins free. Sori terminal on the veins, uni- or less often binerval (rarely trinerval), 
mostly close to the margin. Indusium rigid, attached at the base and at least the 
greater part of the sides. Pluricellular uniseriate filiform paraphyses present 
(probably in all spp.). Spores monolete. Gametophyte unknown. 

Type species: Tapeinidium pinnatum (Cav.) C. Chr. 

Distr. 17 spp., from S. India, the Malay Peninsula, and the Ryu Kyu Is. to Melanesia and Samoa; 
absent from Australia, probably also from New Caledonia. 

Ecol. In forests, mostly at lower and middle altitudes, to c. 2500 m. 

KEY TO THE SPECIES 

1. Lamina simply pinnate and with a conform terminal pinna (fig. 14). 

2. Sori submarginal, most often binerval; petiole and rachis abaxially sharply bi-angular; scales to 
5 mm long 15. T. longipinnulum 

2. Sori intramarginal, almost always uninerval; rachis abaxially sharply bi-angular or sulcate, basally 
dark and pale-margined; scales to 2' 2 mm long 14. T. acuminatum 

1. Lamina more strongly dissected, or, if simply pinnate, the upper pinnae reduced and confluent into a 
pinnatifid leaf-apex, or at least the terminal division strongly lobed at its base. 

3. Lamina simply pinnate, or, if more dissected, the primary rachis abaxially sharply carinate; at least 
a considerable upper portion of the petiole abaxially bi-angular. 

4. Petiole, at least in the upper part, and rachis dark, pale-angled; lamina pinnate + pinnatifid or 
bipinnate. 
5. Larger pinnae of full-grown plants 20-25 mm wide at the widest point; texture subcoriaceous; 
margin often reflexed in dry leaves; lobes of larger pinnae sinuate, the sori not on lobes 8. T. gracile 

5. Larger pinnae of full-grown plants 10-12 mm wide at the widest point; texture herbaceous; 
margin not reflexed; lobes of larger pinnatifid or basally subpinnate pinnae lobed, each sorus on 
a lobe: smaller forms of 1- T. denhamii 

4. Petiole pale, or, if occasionally darker, the rachis not also dark and pale-angled; or lamina simply 
pinnate. 

6. Lamina to 12 cm long; petiole slender, less than 1 mm 10. T. oligophlebiura 

6. Lamina larger; petiole stouter. 

7. Petiole abaxially obtusely bi-angular, dark and dull, ± pale-angled; rachis abaxially mostly 
narrowed-rounded; sori submarginal, on saw-teeth; lamina simply pinnate . 13. T. prionoides 
7. Petiole abaxially sharply bi-angular at least near the apex, nearly always pale; rachis abaxially 
carinate or bi-angular; sori intramarginal; lamina variously dissected. 

8. Lamina pinnate + pinnatifid or more incised* 11. T. luzonicum 

8. Lamina simply pinnate or in large leaves the basal pinnae with very few basal lobes* 

12. T. pimiatum 
3. Lamina at least pinnate + pinnatifid; primary rachis abaxially terete or bi-angular, or, if obtusely 
carinate, the petiole abaxially not (or only at the apex) sharply bi-angular. 
9. Primary rachis atropurpureous; secondary rachises (except sometimes the basal ones) abruptly 
pale; pinnae pinnatifid, with crenate segments, only the basal pinnae occasionally with some pinnat- 
ifid basiscopic pinnules; most sori with their greatest extension at right angles to their vein. 

9. T. calomelanos 

9. Primary rachis at least at the base dark; secondary rachises pale; pinnae pinnate + pinnatifid or 

subbipinnate; sori with their greatest extension in the prolongation of the vein 5. T. stenocarpum 

9. Primary rachis pale, or, if dark, the secondary rachises not abruptly pale; lamina often more incised. 

10. All axes, except the primary, green-margined to base or almost so, i.e., lamina only once fully 

pinnate, then pinnatifid; secondary axes abaxially rounded 2. T. buniifolium 

10. Lamina mostly fully bipinnate; secondary rachises, if marginate, abaxially carinate. 
11. Secondary rachises abaxially black, with two pale lateral or one pale median ridge; lamina 
bipinnate or almost so, with superficially crenato-lobate pinnules; pinnae not enlarged at the 
base 7. T. atratum 



For intermediates see 12a. T. biserratum (Blume) v.A.v.R. 



186 



Flora Malesiana 



[ser. II, vol. 1 



1 1. Secondary rachises abaxially various, but not black with pale ridges; lamina often bipinnate + 
pinnatifid. 
12. Primary rachis and indusia black; ultimate lobes abaxially with very broad and prominent 

veins occupying 1/4-1/3 of their width (fig. 9j 6. T. obtusatum 

12. Primary rachis and indusia pale to dark brown; ultimate lobes with immersed, if slightly 
prominent, relatively much narrower veins. 
13. Indusia about twice as broad as long*, 0.3 mm long; margin bordering the apical sorus of the 

segment usually denticulate; texture herbaceous 1. T. denhamii 

13. Indusia longer, or, if only I3 mm long, — isodiametric or longer than broad*; margin 
bordering the apical sorus entire, or no apical sorus; texture firmer. 
14. Larger segments pinnatifid, each lobe with a sorus overtopped by part of the lobe (fig. 6). 

3. T. amboynense 
14. Larger segments (except for the basal pinnae) crenate, each lobe with a terminal or sub- 
terminal sorus. 
15. Secondary rachises abaxially terete in a considerable basal portion; basiscopic pinnules of 
basal pinnae usually enlarged and more dissected than the others (fig. 10) 

4. T. novoguineense 

15. At least the larger secondary rachises abaxially carinate; basiscopic pinnules of basal pinnae 

rarely enlarged and more dissected (fig. 11) 11. T. luzonicura 



1. Tapeinidium denhamii (Hooker) C. Chr. 
Ind. Fil. (1906) 631; Kramer, Act. Bot. Neerl. 
15 (1967) 583. — Davallia denhami Hooker, 
Second Cent. Ferns (1861) pi. 47. — Microlepia 
denhami (Hooker) Moore, Ind. Fil. 2 (1861) 
292. — Lindsaea denhami (Hooker) Mett. ex 
KuHN, Verb. Zool. Bot. Ges. 19 (1869) 573.- 
Wibelia denhami (Hooker) Kuhn, Chaetopt. 
(1882) 346. - Type: Milne 116, Viti Levu, 
Fiji (K). 

T. teniiiiis Copeland, Philip. J. Sc. 60 (1936) 
110, pi. 17. — Type: Brass 3025, San Cristoval, 
Solomon is. (MICH, dupl. in BISH, GH, L). 

T. tenue auct. non (Brackenr.) Copeland, 
Bull. Bish. Mus. 59 (1929) 69; Kramer, Blumea 
15 (1968) 548; not Microlepia tenuis Brackenr. 
U.S. Expl. Exp. (1854) 236. -Fig. 7. 

Rhizome short-creeping, 1-2 mm 0; scales 
reddish brown, acicular, to I l-i mm long, up to 
4-seriate at base, the greater part uni- or biseriate. 
Leaves clustered to moderately close; petioles 
stramineous to reddish or less often dark brown, 
abaxially obtusely or upward sharply bi-angu!ar 
and then mostly pale-angled, 5-30 cm long, less 
than half to about as long as the lamina. Lamina 
oblong to triangular or subpentagonal, 10-35 cm 
long, 5-20 cm wide, in small specimens pinnate — 
pinnatipartite or mostly bipinnate, in larger ones 
tripinnate — pinnatipartite at base; primary 
rachis like the upper part of the petiole or abaxi- 
ally subterete. Primary pinnae c. 12-20 to a side, 
laxly ascending, subsessile or the basal short- 
petiolutate, if small and once pinnate, linear 
and up to c. 6 by P 2 cm, if larger and more 
compound, to 15 by 10 cm; secondary rachises 
stramineous, abaxially rounded or narrowed- 
rounded, or almost keeled. Secondary pinnae 
of at least twice pinnate leaves c. up to 20 to a 
side, pinnate or pinnate — pinnatifid, the basal 
ones of the basal primary pinnae often prolonged 
and then the leaf approximately pentagonal; 
ultimate pinnules 1-5 cm long, '^-l cm wide. 



lanceolate to linear, acuminate, pinnatifid to 
bipinnatifid. Ultimate segments herbaceous, dark 
green or olivaceous when dry, oblique, ovate or 
oblong to lanceolate, decurrent and connected, 
obtuse or subacute, dentate to pinnatifid, size 
and shape depending greatly on the degree of 
dissection of and the place in the lamina, larger 
and broader in less dissected pinnules, somewhat 
asymmetric, without thickened margin, with evi- 
dent, flexuous costa giving off single (or forked 
in larger lateral lobes), evident veins. Upper 
pinnae, secondary pinnae, and pinnules gradually 
reduced, confluent into pinnatifid, acute or mostly 
acuminate apices of lamina, pinnae, etc. Sori 
single or rarely paired on the teeth of the lobes, 
in most cases also on the apical one, uninerval. 
the soriferous vein distinctly broadened below the 
indusium, the margin opposite the indusium 
occasionally denticulate; indusium brownish, 
delicate, suborbicular to elongate in prolongation 
of its vein, 0.3-0.7 mm long, 0.4-0.8 mm wide, 
not quite reaching the margin and with rounded 
edge or with a small protracted lobe that often 
slightly exceeds the margin, sometimes ruptured 
at maturity, attached at the base and sides. 
Spores brownish, subellipsoidal, smooth, c. 36 
by 28 n- 

Distr. Admiralty Is. (Manus), Bismarck Arch. 
(New Ireland, New Hannover), Solomon Is., 
New Hebrides, Fiji. 

Ecol. Terrestrial in forests, 100-900 m, often 
said to be locally common. 

Note. Quite variable in the degree of dissec- 
tion, but the shape and especially the place of the 
son is very characteristic, no other species regular- 
ly having sori on the terminal lobes. 

2. Tapeinidium buniifolium Kramer, Blumea 15 
(1968) 549.-7". moluccanum auct. non (Blume) 
C. Chr.; Wagner & Grether, Un. Cal. Publ. 
Bot. 23 (1948) 36. -Type: Grether & Wagner 
4188, Tjajiak Mts, Mt Dremsl region, Manus, 



* For the sake of consistency with the terminology employed in Lindsaea the length of an indusium 
is measured at right angles to its vein, the width in prolongation of its vein. 



April 1971; 



LiNDSAEA-GROUP (Kramer) 



187 



Admiralty Is. (MICH; dupl. in BISH, US). 

Rhizome only known from very short pieces, 
c. 3^2 rnm o; scales reddish brown, to c. 1 mm (or 
more?) long, to n: 4-seriate at base, with long, 
acicuiar, uni- or pauciseriate apex. Petioles stout, 
rather lustrous, medium brown, crushed but appar- 
ently abaxially upward obtusely bi-angular, c. 60 cm 
long. Lamina probably 30 cm or more long (no 
complete leaf seen), pinnate + quadripinnatifid 
or subbipinnate + tripinnatifid at the base, 
probably deltoid; primary pinnae at least 4 to a 
side, the lower ones remote. Primary rachis stram- 
ineous to medium brown, abaxially apparently 
bi-angular. Largest primary pinnae oblong, 
with a petiolule of a few mm, c. 18 by 5 cm, 
with c. 12-15 major secondary pinnae to a side, 
these somewhat ascending; axes of higher than the 
first order abaxially stramineous, rounded, nar- 
rowly green-margined to base, only the basal 
ones unmargined at the extreme base. Larger 
secondary pinnae c. 8-12 by 3-6 cm, elongate- 
triangular, with c. 10-15 segments to a side, these 
chartaceous, obliquely ascending, not close, 
lanceolate to linear, the largest 1 ' 2-3 cm by 
4—5 mm, deeply pinnatifid at base, at the acute 
apex like the smaller segments dentato-lobate. 
Ultimate lobes lanceolate-ligulate, the larger ones 
up to 4 by 1^4 mm, subacute, often somewhat 
falcately ascending, asymmetric, with a bulge 
on the anterior margin where the sorus is situated, 
smaller lobes triangular, with subterminal sorus. 
Apical lobes of segments not soriferous. Veins 
abaxially evident at base but higher up evanescing, 
simple, forked in larger lobes. Sori uninerval; 
indusium brownish, pouch-shaped, narrowed at 
base, often with irregular free edge, 0.3-0.5 mm 
long and broad. Spores brownish, subellipsoidal, 
smooth, c. 35 by 26 //. 

Distr. Only known from the type collection. 

Ecol. In mountainside woods, c. 700 m. 

3. Tapeinidium amboynense (Hooker) C. Chr. 
Ind. Fil. (1906) 631; Kramer, Blumea 15 (1968) 
549. — Davallia amboynensis Hooker, Sp. Fil. 
1 (1845) 178, pi. 56 C. — Lindsaea amboynensis 
(Hooker) Mett. ex Kuhn in Miq. Ann. Mus. 
Bot. Lugd.-Bat. 4 (1869) 279. - Wibelia amboynen- 
sis (Hooker) Kuhn, Chaetopt. (1882) 346.- 
Type: Chr. Smith s.n., Ambon (K). 

Davallia stenoloba Baker in Becc. Malesia 3 
(1886) 35. — T. moluccaniim (Blume) C. Chr. 
var. stenolobiim (Baker) C. Chr. Ind. Fil. Suppl. 3 
(1934) \16.-T. stenolobum (Baker) Wagner & 
Grether, Un. Cal. Publ. Bot. 23 (1948) 36.- 
Type: Beccari s.n., Mt Salhutu, Ambon (FI; 
fragm. in K). 

T.amplum Copeland, Occ. Pap. Bish. Mus. 15 
(1939) 82, f. 3. - Type: Takamatsu 1572, Gara- 
sumao, Palau Is. (MICH; dupl. in BISH, K, US). 

T. moluccaniim C. Chr. Ind. Fil. Suppl. 3 

Fig. 6. Tapeinidium amboynense (Hook.) C. Chr. Two pinnules from different parts of the lamina, 
X P/4 (De Vriese 358). - Fig. 7. T. denhamii (Hook.) C. Chr. Pinnule, x 1 1/4 (Brass 3335). - 
Fig. 8. T. atratum Kramer. Pinna, ■ ^(H. J. Lam 1556). - Fig. 9. T. obtusatum v.A.v.R. Pinna, 
X 34 (H. J. Lam 1857). 




188 



Flora Malesiana 



[ser. II, vol. 1 ^ 



(1934) 176, and of later authors; not Davallia 
moluccana Blume. — Fig. 6. 

Rhizome short- to moderately long-creeping, 
3-5 mm 0; scales castaneous, narrowly triangular, 
or with ovate base and acicular apex, to 2^4 irirn 
long, 8- to 10-seriate at base, with rather long 
uniseriate apex. Leaves close to somewhat re- 
mote; petioles medium to olivaceous brown or 
darker with age, abaxiaily rounded or mostly 
upward obtusely or sometimes sharply bi-anguiar, 
then also sulcate and occasionally pale-angled, 
25-50 cm long, as long as to twice as long as the 
lamina. Lamina oblong or elongate-deltoid, some- 
times subpentagonal, c. 25^0 cm long, at the 
base bipinnate + pinnatifid or + bipinnatifid, 
sometimes pinnate + bipinnatifid, with c. 10-15 
primary pinnae to a side and some strongly re- 
duced upper ones; primary rachis abaxiaily 
rounded to obtusely or less often sharply bi- 
angular, not rarely sulcate. Pinnae ascending or the 
basal ones almost spreading, oblong or elongate- 
deltoid or the basal ones subrhombic, scarcely 
asymmetric but the basal basiscopic secondary 
pinnae often somewhat reduced, except in the 
basal pair where one or a few pairs of secondary 
pinnae are often considerably produced; largest, 
basal pinnae up to 15 by 10 cm, the upper ones 
relatively much narrower, all acuminate or cau- 
date; upper pinnae ± gradually reduced, con- 
fluent into a bipinnatifid-pinnatifid leaf-apex. 
Secondary rachises stramineous or pale brown, 
abaxiaily terete, often marginate; secondary and 
ultimate divisions ascending, asymmetric, with 
cuneate base, i decurrent, elongate-ovate, lan- 
ceolate, or linear, obtuse to subacute, often 1-3 cm 
long and 1 ^2-5 mm wide, crenate to pinnatifid, 
or larger, more strongly incised, and more acute; 
number and size very variable, depending on the 
size of and the place in the lamina of the pinna; 
upper ones reduced, confluent. Texture charta- 
ceous or subcoriaceous, sometimes coriaceous; 
colour olivaceous or brownish when dry. Costa 
pale, abaxiaily prominulous, obtuse. Veins very 
oblique, immersed or slightly elevated, mostly ± 
evident, simple or once, or in larger lobes 2 or 3 
times forked; lobes of ultimate segments rounded, 
subacute or acute, each bearing one (or the largest 
f vo) sorus, which except in the very small upper- 
most lobes is distinctly overtopped by part of the 
lobe which, if large enough, receives a branch 
of the vein; apex of segment not soriferous. 
Sori uninerval; indusium brown, pouch-shaped, 
with convex or cuneate base, attached at the base 
and the sides, on the marginal side sometimes 
protracted as a slight ridge, c. 0.4 mm long and 
broad, its somewhat convex free edge not reaching 
the margin, falling short of it by a variable 
distance. Spores yellowish, ellipsoidal, smooth, 
c. 35 by 25 i^i. 

Distr. Malesia: Celebes, Moluccas (Talaud, 
Morotai, Halmahera, Ternate, Ceram, Ambon), 
Kei Is., Waigeo, Biak, W. New Guinea, Palau 
Is.; doubtfully Borneo. 

Ecol. Terrestrial in forests, from sea level to 
800 m. 



4. Tapeinidium novoguineense Kramer, Blumea 15 
(1968) 550. - Type: Schlechter 14319, Torri- 
celli Mts, Terr, of New Guinea (B; dupl. in BM, 
BO, K, P).-Fig. 10. 

Rhizome moderately long- to rather short- 
creeping, 2-4 mm 0; scales medium brown, 
elongate-triangular, to 3 mm long, to c. 12-seriate 
at base, the uniseriate apex relatively short. Leaves 
rather close; petioles stout, the base often verrucu- 
lose from scale bases or persistently scaly, dull, 
stramineous with darker base or darker through- 
out, abaxiaily terete, c. 25-75 cm long, slightly 
shorter than to twice as long as the lamina. Lamina 
bipinnate, or bipinnate -r pinnatifid, at tne base 
often tripinnate, elongate-triangular or subpentag- 
onal-triangular, less often oblong, c. 20-55 cm 
long; primary rachis stramineous to pale brown, 
abaxiaily rounded or narrowed-rounded, keeled 
only near the apex. Pinnae 15-35 to a side (not 
counting the confluent upper ones), laxly ascend- 
ing, lanceolate, or the basal ones broader at the 
base, often acroscopically wider at the base, 
shortly acuminate; larger pinnate pinnae 12-16 
cm long, 3-5 cm wide, with c. 20-30 pinnules to 
a side, the basal pinnae usually with a strongly 
protracted basal basiscopic secondary pinna (smal- 
ler but similar ones may be present next to and 
opposite it, and on the second pair of pinnae) 
similar in shape to a middle primary pinna. 
Secondary (and, if any, tertiary) rachises stram- 
ineous, abaxiaily rounded in the lower, carinate 
in the upper part. Ultimate free pinnules c. 20-30 
to a side, ascending, closely and regularly spaced 
but not contiguous, subcoriaceous or coriaceous, 
mostly olivaceous or brownish when dry, 
lanceolate, mostly obtuse or subacute and with 
narrowed, cuneate-decurrent base, the larger ones 
usually 3^ cm long and 3-4 mm wide, crenate to 
pinnatifid to the middle (or beyond in the basal pin- 
nae), with abaxiaily prominulous, pale costa. 
Veins simple in the lobes (crenations) or forked 
in the larger ones, somewhat prominulous. Lobes 
oblong, or it triangular if small, obtuse. Upper 
pinnae rather suddenly reduced, pinatifid, then 
crenate, gradually simpler, confluent into a pinnat- 
ifid leaf-apex; upper pinnules (segments) also 
reduced and confluent. Sori single in the lobes, or 
paired or a few together in exceptionally large 
ones, uninerval, their vein often conspicuously 
broadened at the end; indusium brownish, pouch- 
shaped to almost triangular, entire, 0.4-0.7 mm 
long, 0.3-0.5 mm wide, not reaching the margin 
by its width or less, about equally remote from 
both the lateral and the apical edges of its 
lobe. Spores brownish, ellipsoidal, smooth, c. 
35 by 24//, exceptionally larger. 

Distr. Malesia: Japen I., New Guinea (all 
Div.); Solomon Is. 

Ecol. In forests and thickets, 200-2 100 m, often 
said to be locally common. 



5. Tapeinidium stenocarpum v.A.v.R. Nova Guinea 
14 (1924) 52; Kramer, Blumea 15 (1968) 550.- 
Type: H. J. Lam 1442, mountain ridge near 



April 1971 



LiNDSAEA-GROUP (Kramer) 



189 



Idenburg R., W. New Guinea (BO; dupl. in K, 
L, SING, U; fragm. in US). 

Rhizome not very short-creeping, 2-2^2 rnm 0; 
scales fuscous, elongate-triangular, to r. 3 mm long, 
to c. 10-seriate at base, with well-developed 
uniseriate apex. Leaves not close; petioles atro- 
purpureous to blackish, dull, abaxially terete in the 
lower part, upward obtusely to subacutely bi- 
anguiar and somewhat pale-angled, c. 15-20 cm 
long, half as long as to as long as the lamina. 
Lamina oblong to elongate-rhombic, bipinnate — 
deeply pinnatipartite at the base (almost tripin- 
nate), c. 20-35 cm long; primary rachis abaxially 
obtusely bi-angular at the base, i distinctly 
pale-angled and often mottled, upward pale, 
narrowed-rounded, subcarinate near the apex. 
Primary pinnae c. 15-25 to a side, ascending, 
often somewhat overlapping, the basal ones c. 
10-15 by 3I2-4 cm, the upper ones gradually, 
then abruptly more strongly reduced, confluent 
into an acuminate pinnatifid leaf-apex. Secondary 
rachises abaxially pale, terete at the base, upward 
narrowed-rounded to subcarinate. Pinnules to 
15-20 to a side, ascending, acute to acuminate, not 
contiguous, the basal basiscopic ones sometimes 
subpinnate — pinnatilobate; average pinnules 
deeply pinnatisect, with c. 10-12 ascending linear 
lobes to a side, these up to 3 by % mm, obtuse, 
connected by narrow, upward broader wings, the 
upper ones denticuliform, broadly connected 
and forming a lobed pinnule-apex; upper pinnules 
of pinnae, and pinnules of upper pinnae, less 
incised, lobed to entire. Pinnule costules abaxially 
prominent, pale, obtuse; lobes often with revolute 
margin, chartaceous, brown when dry, the vein 
immersed but evident, simple or slightly branched 
in the largest ones. Sori single in the ultimate 
lobes, absent from the terminal ones, uninerval, 
terminal in small lobes, on an acroscopic lateral 
bulge in larger ones; indusium narrowly pouch- 
shaped, 0.2-0.4 mm long, 0.3-0.7 mm wide, dark, 
entire, not reaching the margin by U-1 times its 
width. Spores brownish, ellipsoidal, smooth, c. 
35 by 25 /n (very few seen). 

Distr. Malesia: W. New Guinea (2 coll.). 

Ecol. Terrestrial in mossy forests, c. 1400- 
1800 m. 

6. Tapeinidium obtusatum v.A.v.R. Nova Guinea 
14 (1924) 52; Kramer, Blumea 15 (1968) 550.- 
Type: H. J. Lam 1857, mountain ridge near Door- 
mantop, W. New Guinea (BO; dupl. in L, SING, 
U).-Fig. 9. 

Rhizome moderately long-creeping, 3-4 mm 0; 
scales light castaneous, narrowly triangular, to 
5 mm long, to c. 10-seriate at base, there often 
with laterally protruding cell-walls, with a rel- 
atively short uniseriate apex. Leaves not close; 
petioles black, dull, somewhat verruculose at 
base, abaxially terete or very obtusely and in- 
distinctly bi-angular near the apex, c. 25 cm long, 
usually longer than the lamina. Lamina oblong 
or elongate-triangular, c. 15-25 cm long, describ- 
ed as lustrous on both sides when fresh, rigidly 
coriaceous, brown to blackish when dry, at the 



base tripinnate -f- pinnatifid, elsewhere bipinnate 
-f pinnatifid, with c. 10 major pinnae to a side 
and some reduced upper ones; primary rachis 
brown to black, dull, abaxially like the petiole, 
near the apex subcarinate. Pinnae ascending, the 
basal ones triangular, c. 8 by 5 cm, the other 
ones oblong, c. 5-6 by 2 cm; basal pinnae with 
a basiscopic bipinnate -f pinnatifid pinnule, 
otherwise the pinnae once pinnate, with the larger 
pinnules pinnatifid or pinnatilobate. Secondary 
axes dark or, especially the upper ones, pale, 
abaxially narrowed-rounded to subcarinate. 
Larger secondary pinnae with 5-8 segments on a 
flexuous costa, the segments alternate, linear- 
subspathulate, oblique, not close, decurrent, 
joined by narrow wings, the margin subrevolute, 
often 2-4 by 1 mm, with a broad, abaxially much 
elevated, obtuse, stramineous costule that occupies 
about V3 of its width and is often conspicuously 
broadened under the sorus. Apex of lobes obtuse, 
in the larger ones slightly oblique. Sori single 
(rarely paired) and apical in the lobes, uninerval; 
indusium dark, rigid, in larger lobes somewhat 
oblique on the vein, with — straight base and 
convex free edge, adnate at the sides, ^o-' n^m 
long, U mm wide, not quite reaching the margin. 
Spores medium brown, ellipsoidal, smooth, 60 
by 45 //. 

Distr. Only known by the type collection. 

Ecol. Terrestrial in mossy forest, c. 2500 m. 

7. Tapeinidium atratum Kramer, Blumea 15 (1968) 
551. — Type: H. J. Lam 1556, mountain ridge near 
Dooimantop, W. New Guinea (BO, 2 sh.; dupl. 
in L, SING, U).-Fig. 8. 

Rhizome not short-creeping, c. 2 mm 0; scales 
reddish brown, narrowly triangular, to 2 mm long, 
to c. 8-seriate at base, with a long uni-biseriate 
apex. Petioles stout, 4-5 mm at base, almost 
black, sublustrous, basally verruculose, abaxially 
sharply bi-angular above, downward gradually 
rounded, not pale-margined, to 60 cm long, 
about as long as the lamina. Lamina oblong, 
bipinnate, with c. 15-20 remote but ascending and 
often ± touching pinnae to a side; primary 
rachis blackish, abaxially sharply bi-angular, 
upward pale-margined and ± sulcate. Pinnae 
narrowly deltoid, shortly acuminate, c. 20 by 
21/0-3 cm, basiscopically slightly narrowed at the 
base, pinnate, upward pinnatifid; secondary 
rachises dark, abaxially with one median or two 
lateral pale ridges. Upper pinnae gradually re- 
duced, confluent into a pinnatifid leaf-apex. 
Pinnules up to c. 35 to a side, rigidly coriaceous, 
brown when dry, about twice their width apart, 
somewhat ascending, very narrowly lanceolate, 
the largerst c. 17 by 4 mm, pinnatilobate, with c. 
9 lobes to a side, these broadly rounded, to 1 by 
1 mm; apex obtuse; base cuneate, decurrent. 
Costa pale, abaxially prominent, obtuse, flexuous 
towards the apex; veins hidden, simple or once 
forked. Upper pinnules reduced, confluent. Sori 1, 
less often 2 or 3 per lobe, near the anterior margin, 
just inside the apex, uninerval; indusium blackish, 
pouch-shaped, 0.3-0.5 mm long, 0.3-0.4 mm wide, 



190 



Flora Malesiana 



[ser. II, vol. 1 ^ 



attached at the sides, not reaching the margin by 
little less than its width. Spores brownish, ellip- 
soidal, smooth, c. 35 by 25 //, 

Distr. Only known by the type collection. 

Ecol. Collected at 1420 m. 



8. Tapeinidium gracile (Blume) v.A.v.R. Handb. 
(1908) 315; Kramer, Blumea 15 (1968) 551.- 
Davallia gracilis Blume, En. PI. Jav. (1828) 233.- 
Microlepia gracilis (Blume) J. Smith, Lond. J. 
Bot. 1 (1842) 427. - Wibelia gracilis (Blume) 
Christ, Ann. Jard. Bot. Btzg II, 5 (1905) 134.- 
Type: Blume 1731 or s.n., Java (L). 

? Dicksonia linearis Cav. Descr. (1802) 274.— 
Type: Nee s.n., Philippines (MA, n.v.). 

Rhizome rather short- to rather long-creeping, 
c. 2 mm 0; scales reddish brown, narrowly tri- 
angular, to 2^/2 mm long, to c. 6-seriate at base, 
with a long uniseriate apex. Leaves close to c. 
1 cm apart; petioles slender, ^^-i mm at apex, 
reddish brown to atropurpureous, at least upward 
pale-margined, abaxially sharply bi-angular, the 
base usually subterete, c. 10-35 cm long. Lamina 
oblong, c. 12-30 cm long, c. 2-3 times as long as 
wide, to 1 ^2 times as long as the petiole, rarely 
shorter, pinnate + pinnatifid, less often at the base 
pinnate + pinnatilobate; rachis abaxially at the 
base dark and pale-angled, the two abaxial angles 
of the petiole merging into one near the basal 
pinnae to form a sharp keel on the rachis, the 
upper part quite pale. Larger pinnae c. 10-15 to 
a side, somewhat ascending, about their width apart, 
lanceolate to linear, the largest c. 8-15 cm long, 
(V2-)l/4-3 cm wide, subsessile, acuminate; 
lowest pinnae the longest, the upper ones gradual- 
ly and strongly reduced, confluent into a pinnat- 
ifid leaf-apex. Texture chartaceous or subcori- 
aceous or sometimes coriaceous, colour medium 
green or brownish when dry. Lower pinnae 
pinnatilobate to the middle (rarely less) to deeply 
pinnatifid to a narrow costal wing, the lobes 
asymmetrically triangular and serrate or crenate 
if short, linear, obtuse, subentire to crenate if 
larger, ascending, occasionally somewhat falcate, 
often 12-15 major ones to a side in the larger 
pinnae, to 252 by 3 mm; basal basiscopic segments 
of most pinnae reduced, those opposite them often 
somewhat prolongate. Costae of pinnae pale, 
abaxially acute, of segments pale, flat, both 
abaxially prominulous. Upper segments strongly 
reduced, most pinnae with a long pinnatilobate- 
crenate apex. Veins hidden, ascending, simple or 
rarely once forked. Sori uninerval, on the lateral 
and not rarely also on the terminal veinlets of 
the segments, separated by crenations; indusiuni 
pouch-shaped, ± semi-elliptic, with straight or 
slightly lobed edge, V3-/2 mm long and broad, 
mostly falling short of the apex of its lobule by its 
width or more, the margin sometimes revolute 
and touching it when dry. Spores pale brownish, 
subellipsoidal to bean-shaped, smooth, 34-36 
by 24-28 /a.. 

Distr. E. Annam; in Malesia: W. Java, Bali, 
Sarawak, Brunei, Celebes, Ceram, Philippines. 




Map 1. Distribution of Tapeinidium gracile 
(Bl.) v.A.v.R. 

The only species with a notably disjunct area. 
Map 1. 

Ecol. Terrestrial and epilithic, in forests, 500- 
1300 m. 



9. Tapeinidium calomeianos Kramer, Blumea 15 
(1968) 551.— Type: Korthals s.n., G. Sakum- 
bang, SE. Borneo (L, 2 sh.). 

Rhizome short- to somewhat more long- 
creeping, 2-3 mm 0; scales castaneous, elongate- 
triangular, to c. 2 mm long, to c. 6-seriate at 
base, long-acuminate. Leaves close to 1 cm 
apart; petioles very dark purplish brown, dull 
or shining, 1-2 mm at apex, abaxially terete, 
10-35 cm long, mostly shorter than the lamina. 
Lamina elongate-triangular to oblong or sub- 
pentagonal, 10-35 cm long, pinnate -f deeply 
pinnatifid, or at the base bipinnate (-f pinnat- 
ifid), subcoriaceous or coriaceous, olivaceous- 
brown when dry. Primary rachis atropurpureous 
or dark castaneous, abaxially terete. Pinnae 
ascending, c. 1 2-20 major ones to a side, lanceolate, 
the lower ones usually triangular, subsessile, often 
somewhat overlapping, acuminate, the larger 
ones deeply pinnatifid or occasionally pinnate 
at the base and then sometimes the largest 
basiscopic pinnules pinnatifid, to c. 15 by 2^2 cm; 
major pinnules (segments) to 18 to a side, lan- 
ceolate, ascending, 30 by 2^/2 cm to 8 by 7 mm, 
or larger if incised, otherwise serrate or crenate, 
unequally cuneate at the base, obtuse or subacute, 
rapidly decreasing in size to the pinna-apex; 
secondary (and, if any, tertiary) rachises ab- 
ruptly pale (except sometimes those of the 
lowest pinnae), abaxially terete. Upper pinnae 
crenate, confluent into a pinnatifid leaf-apex. 
Veins immersed, hidden, or slightly elevated, 
simple, or forked in the larger lobes. Sori uni- 
nerval, slightly immersed, single in the lobes of 
the ultimate segments, the margin opposite the 
indusium sometimes notched; indusium dark, 
0.3-0.6 mm long, 0.3 mm wide, mostly longer 
than wide, not reaching the margin by its width 
or more. Spores brownish, ellipsoidal, smooth, 
c. 35 by 26 fi. 



April 1971 



LiNDSAEA-GROUP (Kramer) 



191 



Distr. Malesia: Sumatra (?), Borneo, Celebes, 
Philippines (Luzon). 

Ecol. One record from moist shady forest, c. 
140 m. 

10. Tapeinidium oligophlebium (Baker) C. Chr, 
Ind. Fil. (1906) 631; Kramer, Blumea 15 (1968) 
552. — Davallia oligophlebia Baker, J. Bot. 26 
(1888)323. — Wibelia oligophlebia {^.\kek)Chkisj, 
Ann. Jard. Bot. Btzg II, 5 (1905) 134. -Type: 
Hose 220, Laupi, Sarawak (K). 

Protolindsaya brooksii Copeland, Philip. J. 
Sc. 5 (1910) Bot. 283. — T. brooksii (Copeland) 
C. Chr. Ind. Fil. Suppl. 3 (1934) 176.— Type: 
Brooks 47, G. Bengkaim, Sarawak (SAR, 
holotype?; dupl. in BM). 

Rhizome short-creeping, -'i-l mm o; scales light 
brown, lanceolate to elongate-triangular, to l^i 
mm long, to c. 6-seriate at base, the uniseriate 
apex comparatively short. Leaves close; petioles 
slender, stramineous with darker base or brown 
throughout, abaxially bi-angular, 2-10 cm long. 
Lamina narrowly lanceolate to triangular, about 
as long as to 5 times as long as wide, 2^2^12 cm 
long, with c. 6-20 pinnae to a side; rachis 
abaxially bi-angular at the base, the angles fusing, 
mostly between the two basal pairs of pinnae, 
to form one kQQ\. Pinnae slightly ascending or the 
basal ones spreading, ovate and crenate to linear 
and then usually pinnatilobate to pinnatifid, 
or the basal ones fully pinnate, with crenate 
pinnules; size and shape of pinnules strongly 
dependent on the degree of dissection of and the 
place in the lamina; larger pinnules often c. 
2 mm wide, mostly obtuse. Upper pinnae and 
pinnules gradually reduced, confluent. Texture 
herbaceous to subcoriaceous, colour usually 
brownish or olivaceous when dry. Secondary 
rachises (costae) abaxially carinate, costae of 
smaller divisions usually rounded and obsolescent 
above the base. Veins immersed, abaxially — 
evident. Sori uninerval, single or a few together 
on the lateral lobes of the larger segments, in 
larger lobes sometimes overtopped by a sterile 
part of the lobe, otherwise subterminal; in- 
dusium \i mm broad, Vi-Vz rnm long, brownish, 
subentire, pouch-shaped, not reaching the mar- 
gin by its width or more, sometimes ruptured 
at maturity. Spores brownish, ellipsoidal, smooth, 
c. 35 by 25 fx. 

Distr. Malesia: Borneo (Sarawak, Kaliman- 
tan). 

Ecol. Terrestrial, in forests and on shaded 
cliff's, c. 700-1000 m. 

Note. The specific distinctness of this taxon 
is doubtful. It may be a reduced variety or 
form of another species, e.g. T. luzonicum. 

11. Tapeinidium luzonicum (Hooker) Kramer, 
Blumea 15 (1968) 552. - Davallia luzonica 
Hooker, Sp. Fil. 1 (1845) 174, pi. 60 B f. 2, 3, 
5.— mbelia bipinnata Fee, Gen. Fil. (1852) 331, 
nom. super fl. — Type: Cuming 139, p.p., Luzon 
(dupl. in B, GH, L). 

Lindsaea pinnata (Cav.) Mett. ex Kuhn var. 



bipinnata Mett. ex Kuhn in Miq. Ann. Mus. 
Bot. Lugd.-Bat. 4 (1869) 279. - Type: Zol- 
linger 1305, Java (dupl. in HBG, L). 

Davallia philippinnesis Harrington, J. Linn. 
Soc. Lond. 16 (1877) 27. — Microlepia philippi- 
nensis (Harrington) Copeland, Polypod. Philip. 
(1905) 56. — T. philippinense (Harrington) C. 
Chr. Ind. Fil. Suppl. 3 (1934) 176. - Type: 
Steere s.n., Mt Mahayhay, Luzon (K). 

Davallia hosei Baker, J. Bot. 26 (1888) 323. - 
Type: Hose 219, Lambur, Sarawak (K). 

T. sumatraniiin v.A.v.R. Bull. Jard. Bot. 
Btzg III, 2 (1920) 174. - Type: Brooks 332/S, 
Bencoolen, Sumatra (BM; fragm. in BO). 

T. biserratum auct. non (Blume) v.A.v.R.; 
Holttum, Rev. Fl. Mai. 2 (1954) 339, f. 197; 
and of other authors. — Fig. 11. 

Rhizome usually short-creeping, to 4 mm 0; 
scales golden brown, narrowly triangular, long- 
acuminate, to 4 mm long. Petioles about as 
long as to twice as long as the lamina, abaxially 
terete at the base, in the upper half or less often 
only near the apex obtusely to acutely bi-angular, 
flat or usually sulcate. Lamina oblong, narrowly 
oblong, triangular, or occasionally subpentagonal, 
at least once pinnate -f pinnatifid. Primary 
rachis stramineous or pale brown, abaxially 
sharply carinate. Secondary rachises (costae) 
abaxially elevated, sharply carinate, pale. Upper 
pinnae and pinnules (segments) gradually reduced, 
confluent. Sori uninerval. Spores pale brown, 
ellipsoidal, smooth, c. 40 by 28 /<. 

KEY TO the varieties 

1. Petiole less than 1 mm o at base of lamina; 
lamina subtripinnate at the base 

1. var. leptophyllym 

1. Petiole over 1 mm at base of lamina; lamina 

pinnate + deeply pinnatifid or bipinnate 

2. var. luzonicum 

1. Petiole sometimes less than 1 mm at base 

of lamina; lamina pinnate, the pinnae not 

incised beyond % of their width 

3. var. thelypteridoides 

1. var. leptophyllum Kramer, Blumea 15 (1968) 
553. -Type: Elmer 14103, MtUrdaneta, Mindanao 
(L; dupl. in BM, BO, HBG, MICH). 

Rhizome scales to 7-seriate at base. Petioles 
close, slender, at the most 1 mm at the base 
of the lamina, often medium brown. Lamina 
almost tripinnate, i.e. the tertiary divisions 
there almost free; ultimate free or nearly free 
divisons linear, obtuse, 1-2 mm wide, mostly 
not over 1 cm long; texture chartaceous, colour 
olivaceous or brownish when dry. Veins ± evi- 
dent, mostly simple in the lobes, these usually 
regularly rounded; indusium c. 0,3 mm long 
and broad. Otherwise like var. luzonicum. 

Distr. Malesia: Philippines (Luzon, Negros, 
Mindanao, Panay, Leyte). 

Ecol. No data. 

Note. In appearance not unlike Pityrogramma 



192 



Flora Malesiana 



[ser. II, vol. r 










is^ul ui 





Fig. 10. Tapeinidium novoguineense Kramer. Basal part of lamina, X ^j^, pinnule, X 2, portion of racnis, 

X 3^4 (v. ROYEN &SLEUMER 6394). — Fig. 11. r. luzonicum (Hook.) Kramer var. luzonicum. Basal part 

of lamina, x 2/^, pinnule, X 2, portion of rachis, X 3^4 (SAN 24071). 



April 1971 



LiNDSAEA-GROUP (Kramer) 



193 



calomelanos (L.) Link; also resembling T. den- 
hamii, from which it differs, /.«., by the rounded, 
not dentate, soriferous lobes, the structure of the 
axes, and the firmer texture. 

2. var. luzonicum. — Fig. 11. 

Rhizome often not short-creeping, the petioles 
up to a few cm apart, stramineous to pale sordid 
brown, concolorous, c. 20-65 cm long, 1 ^ 2-~ rnrn 
at the base of the lamina. Lamina 18-50 cm long, 
8-30 cm wide, up to 4 times as long as wide, 
but mostly shorter, coriaceous, less often chart- 
aceous, dark brownish or olivaceous when dry, 
pinnate — deeply pinnatifid, or at the base 
bipinnate — pinnatifid. Major pinnae c. 6-12 to 
a side, spreading or ascending, 10-20 cm long, 
their shape and width very variable, depending 
on the degree of dissection, linear and 1-2 cm 
wide if pinnatifid, triangular and to 12 cm wide 
if pinnate — pinnatisect, acuminate, with long 
serrato-crenate apex. Basal basiscopic divisions 
of pinnae mostly somewhat reduced, those opposite 
them not distinctly enlarged, if more dissected than 
pinnate — pinnatifid, only rarely with one basi- 
scopic or a pair of basal enlarged, more incised 
secondary pinnae (as in T. novoguineense). 
Ultimate free (or almost free) divisions ascending, 
variable in size and shape, often 1^9-2^2 cm 
long, 1-4 mm wide, mostly obtuse or subacute, 
or the longest acute or shortly acuminate, shallow- 
ly crenate to pinnatifid, margin often — revolute 
when dry; small segments with only the anterior 
margin crenate. Veins immersed, not evident, 
simple, or in the largest segments with a few 
branches. Sari often slightly immersed; indusium 
dark, rather rigid, pouch-shaped, ^-'^ rnrri wide, 
/2-'4 mm long, the free edge entire, convex, not 
reaching the margin by about its width, some- 
times the exterior lateral margin protracted as a 
ridge on the leaf-tissue. 

Distr. Thailand, in Malesia: Malay Penin- 
sula, Natuna and Lingga Is., Banka, Sumatra, 
W. Java, Borneo, Celebes, Philippines (Luzon, 
Mindanao, Polillo). 

Ecol. Terrestrial in moist forests, mostly 
600-1500 m, occasionally to 2200 and down to 
100 m. Often said to be locally common, but 
rare in Java. 

Note. See under 12a. T. biserratuin. 

3. var. thelypteridoides Kramer, Blumea 15 
(1968) 553. -Type: Brooke 8190, Mt Santu- 
bong, Sarawak (L; dupl. in SING, US). 

Rhizome short-creeping, with relatively narrow 
scales; petioles clustered, 10-27 cm long, not 
rarely less than 1 mm o at base of lamina. Lamina 
to c. 40 by 20 cm, chartaceous, rather pale green 
when dry, pinnate — pinnatifid; pinnae to 15 cm 
long and 12 mm wide, very regularly pinnatifid, 
the lowest to %, the upper ones gradually less, 
serrate, then subentire; segments oblong-ligulate, 
ascending, the largest c. 12 by 3 mm, obtuse, 
shallow ly crenate-serrate; pinnae basally on the 
posterior margin shortly narrowed. Veins evi- 
dent. Sori often more distinctly intramarginal. 



Otherwise like less compound forms of var. 
luzonicum. 

Distr. Malesia: Borneo (Sarawak, Sabah) 
(4 coll.). 

Ecol. On rocks in forest, 400-1300 m. 

12. Tapeinidium pinnatum (Cav.) C. Chr. Ind. 
Fil. (1906) 631; Holttum, Rev. Fl. Mai. 2 (1954) 
339, f. 196; Copeland, Fern Fl. Philip. 1 (1958) 
114; Kramer, Blumea 15 (1968) 553. -Davallia 
pinna ta Cav. Descr. (1802) 277, non Mett. ex 
KuHN (1869). — Saccoloma pinnatum (Cav.) Presl, 
Tent. Pterid. (1836) 126. — Microlepia pinnata 
(Cav.) J. Smith, Hook. J. Bot. 3 (1841) 416.- 
Wibelia pinnata (Cav.) Bernhardi ex Fee, 
Gen. Fil. (1852) 331, pi. 27 bis B.—Lindsaea 
pinnata (Cav.) Mett. ex Kuhn in Miq. Ann. 
Mus. Bot. Lugd.-Bat. 4 (1869) 279.— Type: 
Nee s.n., Philippines (also incorr. cited 'Chile') 
(MA, n.v.; phot. seen). 

Davallia flagellifera Hooker & Greville, 
Ic. Fil. (1831) pi. 183. -Type: Wallich s.n., 
Pulu Penang (n.v.). 

Davallia serrata RoxB. ex Griff. Calc. J. Nat. 
Hist. 4 (1844) 514, non Willd. (1810). -Type: 
Roxburgh s.n., Pulu Penang (n.v.). 

Wibeliajavae Fee, Gen. Fil. (1852) 331. -Type: 
Kollmann s.n., Java (n.v.). 

Davallia firmula Baker, Ann. Bot. 8 (1894) 
123. — Microlepia firmula (Baker) C. Chr. Ind. 
Fil. (1906) 426. — Tapeinidium firmulum (Baker) 
C. Chr. Ind. Fil. Suppl. 3 (1934) 176.— Type: 
Hancock 72, Barisan Range, Sumatra (K). 

Rhizome short- to rather short-creeping, 2-3 
mm 0; scales castaneous, with an oval basal and 
a long acicular uni-biseriate apical part, to c. 
10-seriate at base, to 4 mm long. Leaves clustered 
to 1 cm apart; petioles stramineous to medium 
brown, mostly darker at base, or rarely quite 
dark, in adult plants 1-2 mm at base of lamina, 
abaxially at least in the upper half obtusely 
or more often acutely bi-angular, c, 10-35 cm 
long, 1-2-73 the length of the lamina. Lamina 
oblong, acuminate, simply pinnate, c. 15-75 cm 
long, with c. 12-30 major pinnae to a side; rachis 
stramineous to pale brown, abaxially sharply 
carinate, the keel starting at the base through 
coalescence of the two angles on the petiole. 
Pinnae coriaceous, mostly olivaceous-brown 
when dry, ascending, sessile, narrowly lanceolate 
to linear, shortly and somewhat unequally cuneate 
at the base, subacute to acuminate, the basal 
ones remote, slightly or not reduced, the upper 
ones gradually closer but usually at least their 
width apart, gradually, then just below the 
pinnatifid leaf-apex suddenly more strongly 
reduced, the terminal lobe often caudate. Major 
pinnae 7-22 cm long, 0.3-0.8 cm wide, shallowly 
serrate or more often crenate or bicrenate when 
fertile, serrate or biserrate when sterile, with 
strongly ascending teeth; costa percurrent, 
abaxially elevated, stramineous, acute. Veins im- 
mersed, mostly hidden, once or twice forked, 
oblique. Sori uninerval (rarely on two adjacent 
vein-ends), on the acroscopic vein-branch and then 



194 



Flora Malesiana 



[ser. II, vol. P 



single, or on both branches, and then paired in the 
lobes, scarcely or not embossed. Indusium dark, 
subentire, pouch-shaped, ovate to transversely 
elongate-semi-elliptic, 0.3-1 mm long, 0.3-0.5 mm 
wide, falling short of the margin by its width or 
more. Spores medium brown, subellipsoidal to 
bean-shaped, smooth, c. 35 by 25 /i. 

Distr. S. India, Thailand, Ryu Kyu Is., Tai- 
wan; in Malesia: Malaya, Singapore, Riouw and 
Lingga Is., Banka, Sumatra, W. Java, Borneo, 
Celebes, Philippines. Reports from elsewhere 
due to confusion with T. longipinnulum and T. 
melanesicum. Map 2. 




Map. 2 Distribution of Tapeinidium pinnatum 
(Cav.) C. Chr. 

Ecol. In moist forests, often by or on rocks in 
streams, 50-2700 m, mostly between 500 and 
1000 m. Apparently a facultative rheophyte. 

Notes. Juvenile plants have relatively broader, 
serrate pinnae, but apart from their size they are 
similar to adult ones. 

In Luzon there is a form with dark, often 
pale-margined petiole and abaxially narrowed- 
rounded rather than carinate rachis; the pinnae 
are smaller and narrower than in the typical 
form which is much more common on the island. 
A fairly uniform series of this form, Jacobs 
7852 (L), would seem to indicate that it is more 
than an accidental, phenotypic form; but it is 
neither very strongly nor very sharply distinct from 
typical T. pinnatum and therefore left unnamed. 

12a. Tapeinidium biserratum (Blume) v.A.v.R. 
Handb. Suppl. (1917) 509; Kramer, Blumea 15 
(1968) 554; not of Holttum, Rev. Fl. Mai. 2 
(1954) 339. — Davallia biserrata Blume, En. 
PI. Jav. (1828) 232.- Microlepia biserrata (Blume) 
Presl, Epimel. Bot. (1851) 97.-Type: Blume 
s.n., Java (L.). 

The type of this 'species' is intermediate between 
T. luzonicum and T. pinnatum; there are 25-30 
other intermediates. Their status is not clear; 
see Kramer, I.e. They occur throughout the com- 
mon range of the two species. 

13. Tapeinidium prionoides Kramer, Blumea 15 
(1968) 554. -Type: Bunnemeijer 1910, G. Siang, 
Banka (L; dupl. in BO). -Fig. 13. 

Rhizome rather short-creeping, c. 3-4 mm 0; 



scales fuscous, elongate-triangular, long-acuminate, 
to 3 mm long, to c. 8-seriate at base, the uniseriate 
apex rather short. Leaves close; petioles dark 
stramineous to castaneous, dull, abaxially (mostly 
obtusely) bi-angular, upward usually pale-edged, 
c. 10-45 cm long, 73-1 1/; times the length of the 
lamina. Lamina simply pinnate, oblong, with c. 
10-18 free pinnae to a side and a pinnatifid, 
basaliy hastate leaf-apex; rachis stramineous to 
dark, abaxially narrowed-rounded or sometimes 
carinate, at the base often shortly bi-angular. 
Pinnae coriaceous, dark olivaceous above when 
dry, ascending, linear, subacute to acuminate, 
c. 10-20 cm long, 4-6 mm wide, unequally 
cuneate at the base, the margin serrate; teeth 
ascending, 1/2-1 mm long, acute when sterile, 
± obtuse if soriferous, the basal acroscopic 
tooth often larger and slightly auricle-like. Costa 
stout, percurrent, mostly pale, abaxially strongly 
elevated, rounded to subacute; veins abaxially 
rt prominulous, close, very oblique, once forked, 
the anterior branches running to the marginal 
teeth, or when fertile often both branches bearing 
a sorus on their connivent ends. Upper pinnae 
rather suddenly reduced. Sori placed in the 
teeth, uni- or occasionally binerval; indusium 
darkish, entire, Y^-l mm long, V2 rnm wide, 
almost reaching the margin. Spores light brown, 
ellipsoidal, smooth, c. 36 by 28 ^i. 

Distr. Malesia: Anambas, Riouw, and Lingga 
Is., Banka. 

Ecol. In forests, sometimes on rocks in streams, 
200-300 m. 

14. Tapeinidium acuminatum Kramer, Blumea 15 
(1968) 554.— Type: Escritor 21 173, Palanan Bay, 
Luzon (L; dupl. in BO, BRl, GH, MICH, SING, 
US). -Fig. 14. 

Rhizome rather short-creeping, 2-3 mm 0; 
scales castaneous, lanceolate, long-acuminate, 
to 2^2 mrn long, to c. 10-seriate at base. Leaves 
close; petioles dark castaneous or atropurpureous, 
abaxially at least upward sharply bi-angular and 
pale-angled, c. 25 cm long, about as long as the 
lamina. Lamina oblong, simply pinnate, with 8-12 
pinnae to a side and a conform terminal one; 
rachis like the upper part of the petiole or paler. 
Pinnae coriaceous, brown when dry, ascending, 
very narrowly lanceolate, acuminate, 10-15 cm 
long, 6-7 mm wide, unequally cuneate at the base, 
the margin shallowly and evenly crenate or some- 
times bicrenate. Upper pinnae somewhat reduced; 
terminal pinna conform, smaller than the larger 
lateral ones. Costa prominent on the abaxial 
side, pale, flattish; veins immersed, not evident, 
close, very oblique, once forked, sometimes the 
posterior branch forked again. Sori uninerval, 
laminal or partly extending onto the largest lobes; 
indusium brown, subentire, y^ mm wide, ' 2-^4 tuti 
long, with convex free edge and ± convex base, 
V4 rnm or farther from the margin. Sporangia 
strongly protruding at maturity. Spores abortive. 

Distr. Beside the type two doubtful collections 
from Sabah. 

Ecol. No data. 



April 1971] 



LiNDSAEA-GROUP (Kramer) 



195 




Fig. 12. Tapeinidium longipinnulum (Cesati) C. Chr. Basal part of lamina, X 1/2, portion of pinna, X 3, 

portion of rachis, X 5 (Bamler Ros. 115). — Fig. 13. T. prionoides Kramer. Basal part of lamina, x 1/2. 

portion of pinna, x 3, portion of rachis, x 5 (Bunnemeijer 1910). 



196 



Flora Malesiana 



[ser. II, vol. P 




I'^l' 


tM 


W'i 


'h% 




i'i 


W:^''J 


'■m 


W,;t;'-' 


Mm 



Fig. 14. Tapeinidium acuminatum Kramer. Upper part of lamina, X 1/2. portion of pinna, X 3, portion 

ofrachis, x 5 (Escritor 21173). 



April 1971] 



LiNDSAEA-GROUP (Kramer) 



197 



15. Tapcinidium longipinnulum (Cesati) C. Chr. 
Ind. Fil. Suppl. 3 (1934) 176; Copeland, Philip. 
J. Sc. 78 (1949) 23: Kramer, Blumea 15 (1968) 
555. — Davcillici longipiniuila Cesati, Rer.dic. R. 
Accad. Sci. Fis. Mat. Napoli 16 (1877) 26; 
Beccari, Malesia 3 (1886) 35. — Type: Beccari 
s.n., Ramoi, New Guinea (FI). 

Davallia intnimarginolis Cesati, Rendic. R. 
Accad. Sci. Fis. Mat. Napoli 16 (1877) 29.- 
Type: Beccari s.n., Mt Arfak, New Guinea (FI). 

T. margincile Copeland, Philip. J. Sc. 6 (1911) 
Bot. 82.— Type: King 283, Papua (MICH; 
dupl. in P).— Fig. 12. 

Rhizome rather short-creeping, 4-6 mm o; 
scales dark castaneous, elongate-triangular, to 
5 mm long, to c. 16-seriate at base, the uniseriate 
apex very short. Leaves close to more remote; 
petioles dark castaneous to blackish, dull, quadran- 
gular, upward sharply so and often sulcate, ± 
distinctly pale-angled, 10-50 cm long, shorter than 
the lamina. Lamina oblong, simply pinnate, 30-65 
cm long, 15-25 cm wide, with 8-20 pinnae to a 
side and a conform terminal one; rachis similar 
to the petiole, sulcate, upward paler and narrowly 
marginate. Pinnae alternate or the lower ones op- 
posite, ascending, coriaceous, olivaceous to 
fuscous when dry, subsessile or the lower ones 
with a short stalk-like base, linear, the largest 
10-25 cm long, 7-10 mm wide, widest a little 
above the base, long-acuminate, unequally 
cuneate at the base; lower pinnae not rarely 
slightly reduced, the terminal pinna conform, 
with unequal base, occasionally with 1 or 2 



reduced pinnae at its base. Margin of sterile 
pinnae serrate or biserrate, of fertile ones serrate, 
the teeth strongly ascending, broadly truncate, 
posteriorly rounded, each bearing a sorus; basal 
acroscopic tooth not rarely somewhat larger. 
Costa percurrent, abaxially elevated and rounded, 
stramineous. Veins often hidden, rather oblique, 
once forked or the anterior branch forked again, 
c. 1 mm apart, almost reaching the margin. 
Sori uni- or binerval (rarely trinerval), the upper 
ones most often binerval; indusium brownish, 
entire, semi-elliptic, y->-2, most often 73-I mm 
long, c. Ys mm wide, very nearly equaling the 
margin, often strongly bulging at maturity. 
Spores brownish yellow, oblong, smooth, 30-35 
by 24-26 /«. 

Distr. Malesia: Ceram, Japen, New Guinea 
(all Div.), Rossell I. 

Ecol. Terrestrial in rain-forest, from sea level 
to c. 1500 m; often described as locally frequent. 



Excluded 

Tapeinidium bartlettii Copeland, Un. Cal. 
Publ. Bot. 14 (1929) 376, pi. 60 = Xyropteris 
stortii (v.A.v.R.) Kramer. 

Tapeinidium moluccanum (Blume) C. Chr. 
Ind. Fil. Suppl. 3 (1934) 176 = Saccoloma sp. 
(see under T. aniboynense). 

Tapeinidium tenue (Brackenr.) Copeland, 
Bull. Bish. Mus. 59 (1929) 69 = Saccoloma 
sp. (see under T. denhamii). 



3. XYROPTERIS 

Kramer, Act. Bot. Neerl. 6 (1957) 599. 

In most respects similar to the larger species of Tapeinidium. Lamina simply 
pinnate. Pinnae of full-grown plants on the acroscopic side of the base sharply 
auriculate. Sori plurinerval, in full-grown plants quite continuous and occupying 
all vein-ends. 

Distr. Monotypic. 

Note. This is undoubtedly a close relative of Tapeinidium. That genus is, however, so homogeneous 
in its short sori that the present species is better excluded. 



1. Xyropteris stortii (v.A.v.R.) Kramer, Act. 
Bot. Neerl. 6 (1957) 599, with fi%.~~Sclnzoloma 
stortii v.A.v.R. Bull. Jard. Bot. Btzg II, 16 
(1914) 36; Handb. Suppl. 1 (1917) 214. —Type: 
Amdjah (v. Genderen Stort's coll.) 711, Mt 
Djempanga, Kalimantan, Borneo (BO; dupl. 
in K, L; fragm. in BM). 

Tapeinidium bartlettii Copeland, Un. Cal. 
Publ. Bot. 14 (1929) 376, pi. 60.— Type: Bart- 
lett 6731a, near waterfall Si Saliang, Asahan, 
Sumatra (dupl. in GH, L, MICH). 

Rhizome rather short-creeping, in full-grown 
plants to P/4cm0; scales golden-brown, elongate- 
triangular, acuminate, to 5 by lYi mrn> to c. 18- 
seriate at base. Petioles close, to a few mm apart, 
to 40 cm long, to 1 V2 cm at base, more slender 
upward, stramineous to fawn-coloured, abaxially 
terete, or in young plants somewhat angular, 



adaxially flattened to sulcate. Lamina 70 to 150 
cm long, but already fertile in young plants and 
then c. 30 cm long, oblong, with up to 18 pinnae 
to a side; rachis like the petiole, adaxially narrow- 
ly sulcate. Pinnae ascending, alternate, very 
narrowly lanceolate, 15-30 cm long, 1^2-3 cm 
wide, the upper ones shortened, 4-10 cm long; 
texture chartaceous to subcoriaceous, colour 
olivaceous when dry; base of (especially larger) 
pinnae stalk-like, to I cm long. Margin obliquely 
and distantly serrate in young plants, entire with 
only a few teeth near the apex in full-grown 
ones. Lamina of pinnae unequal at the base, the 
basiscopic side narrower, the acroscopic side with 
a large acute auricle to 7 cm long, this wanting 
in juvenile plants. Costa median, percurrent, 
pale, prominent on both sides, adaxially sulcate; 
veins immersed, evident, very oblique, less so 



198 Flora Malesiana [ser. II, vol. P 

outward, mostly 2 or 3 times forked, free; auricles paraphyses present among the sporangia. Spores 

of pinnae with a costule. Upper pinnae in large medium brown, monolete, bean-shaped, smooth, 

leaves 4-10 cm long; terminal pinna free, to c. c. 50-60 by 30-35 //. 

15 cm long, unequally and deeply trilobate, with Distr. Malesia: Sumatra (3 coll.), Borneo 

long-acuminate lobes, the central shank much (Kalimantan) (type coll.). 

longer than the lateral ones. Sori plurinerval but Ecol. In ravines by streams, c. 300-500 m; 

interrupted in young, continuous in adult plants, presumably a rheophyte. 

absent only from the extreme base and apex of Note. An excellent series of specimens from 

the pinnae and the apex of the auricle; indusium Sumatra (Surbeck 49, L, U), including young 

brown, entire or wavy, V4-I3 mm wide, not as well as full-grown plants, shows that Tapei- 

reaching the margin by about its own width, nidium bartlettii Copeland is the juvenile form 

± reflexed at maturity. Multicellular filiform of X. stortii. 

4. LINDSAEA 

Dryander in J. E. Smith, Mem. Ac. Turin 5 (1793) 401; Trans. Linn. Soc. 3 
(1797) 39; Hooker, Sp. Fil. 1 (1846) 203; J. Smith, Hist. Fil. (1875) 267; Diels 
in E. & P. Nat. Pfl. Fam. I, 4 (1902) 219; v.A.v.R. Handb. (1908) 260; Suppl. 
(1917) 202; Holttum, Gard. Bull. S. S. 5 (1930) 58; Tagawa, Act. Phytotax. 
Geobot. 6 (1937) 24; Copeland, Gen. Fil. (1947) 52; Philip. J. Sc. 78 (1949) 15; 
Holttum, Rev. Fl. Mai. 2 (1954) 321; Kramer, Act. Bot. Neerl. 6 (1957) 97; 
Ching, Fl. Reip. Pop. Sin. 2 (1959) 257; Kramer, Blumea 15 (1968) 551 .—Davallia 
J. E. Smith sensu Hooker, Sp. Fil. 1 (1845) 151, p.p. — Schizoloma Gaud. Ann. 
Sc. Nat. 3 (1824) 507, p.p.; J. Smith, Hist. Fil. (1875) 270, p.p.; Diels in E. & P. 
Nat. Pfl. Fam. I, 4 (1902) 21^, p.p.; v.A.v.R. Handb. (1908) 275; Suppl. (1917) 
214; Holttum, Rev. Fl Mai. 2 (1954) 342; Ching, Fl. Reip. Pop. Sin. 2 (1959) 
lll.—Isoloma J. Smith, Hook. J. Bot. 3 (1841) 414; Hist. Fil. (1875) 227; 
Copeland, Gen. Fil. (1947) 55; Philip. J. Sc. 78 (1949) 24; Holttum, Rev. Fl. 
Mai. 2 (1954) 336; Copeland, Fern Fl. Philip. 1 (1958) 100.— Odontoloma J. 
Smith, Hook. J. Bot. 3 (1841) 415, nom. submid.; in Hooker & Bauer, Gen. Fil. 
(1842) pi. 114 B; Fee, Gen. Fil. (1852) 329; J. Smith, Hist. Fil. (1875) 269.— 
SynaphJebium J. Smith in Hooker & Bauer, Gen. Fil. (1842) pi. 101; Hist. Fil. 
(1875) 268. — Lindsaenium (or Lindsayniiiin) Fee, Mem. Soc. Mus. Hist. Nat. 
Strasb. 4 (1850) 201; Gen. Fil. (1852) 333. 

The name is often misspelled 'Lindsaya\ 

Small to medium-sized, terrestrial, epilithic, scandent, or epiphytic ferns 
with a Lindsaeoid protostele, the xylem with an internal phloem strand, or in some 
small epiphytes open. Scales variable in shape, mostly entire. Lamina rarely 
simple, mostly once or twice pinnate, sometimes more dissected, to decompound, 
anadromous; ultimate divisions various, most often dimidiate, sometimes partly 
or entirely equal-sided, rarely cuneate and dichotomously divaricate. Veins free, 
connivent, or anastomosing without free included veinlets. Sori terminal on the 
veins, bi- to plurinerval, less often uninerval, mostly very close to the margin. 
Indusium short, roundish, ovate, or hippocrepiform and then free at the sides, 
or more elongate, and laterally free or adnate, rarely fugacious. Bicellular filiform 
paraphyses present in some, probably in all species. Spores trilete or (very rarely 
in the Old World species) monolete. 

Type species: Lindsaea trapeziformis Dryander (neotropical). 

Distr. About 150 spp., 73 in the Old World, but few in continental Africa; extending north to Japan, 
south to Tasmania, east to the Marquesas. 

Descriptive terms. In accordance with the terminology used earlier in the Lindsaea group (Kramer, 
1957) the term 'pinnule' is always used for an ultimate free division, regardless of the degree of dissection 



April 1971] LiNDSAEA-GROUP (Kramer) 199 

of the lamina which bears it, except in some species of sect. Schizoloma, in which there are transitions 
from simple entire primary pinnae to pinnate ones. If the division is not entirely free it is called a segment. 
The length of a sorus is always measured at right angles to the vein(s) bearing it, its width parallel to 
(in prolongation ot ) the vein(s). 

SUBDIVISION OF THE GENUS 

The subgenera of Lindsaea, as proposed in the past, proved as a whole unsatisfactory. Too much value 
was attached to such characters as the shape of the pinnules and their venation and the length of the 
sori. Some of the subgenera defined on such characters were even raised to generic rank. It seems that 
these features are useful, together with several others, for distinguishing species, or sections at the most. 
It can be demonstrated, for instance, that anastomosing veins have independently arisen in four groups 
of species, here treated as sections Synaphlehiunu Schizoloma, Lindsaenium, and Penna-arborea, respec- 
tively. A feature of much more fundamental importance seems to be the morphology and anatomy of 
the rhizome. This led to the distinction of two clear-cut groups, treated as subgenera (Kramer, Biumea 
15, 1968, 557 seq.). Apparently natural, but not necessarily entirely sharply defined groups of species 
are treated as sections under these two subgenera. The number of characters available at present for 
defining these groups is relatively small in the Lindsaea group, these rather primitive ferns being of 
simpler structure than most other leptosporangiate ferns. As in some species, notably in sect. Schizoloma 
and sect. Synaphlebium, not all specimens show the sectional characters very clearly, the key to the species 
has not primarily been constructed on the basis of the sections, although in many instances most or all 
species of a section will be found together. 

SYNOPSIS OF MALESIAN SUBGENERA AND SECTIONS 

1. Rhizome essentially terrestrial, short- to moderately long-creeping, the stele radially symmetric or 

nearly so. Subg. Lindsaea. 
2. Lamina bi-(tri-)pinnate, with the upper pinnae gradually reduced, a conform terminal pinna wanting; 
or simply pinnate (rarely simple), with equal-sided pinnules and anastomosing veins (one species 

with combinations of both possibilities). 5/1/1. 1-8 1. Seer. Schizoloma 

2. Lamina bipinnate, with a conform terminal pinna; or simply pinnate, with free veins, or, if the veins 
are anastomosing, with dimidiate pinnules. 
3. Lamina simply pinnate, with — equal-sided pinnules; rachis ^i sclerotic. 
4. An articulation at the base of each pinnule. 5p/7. 36-41 8. Seer. Isoloma 

4. No articulations at the pinnule-bases. 5/?. 35 7. Sect. Psammolindsaea 

3. Lamina bipinnate, or, if simply pinnate, with dimidiate pinnules; rachis various. 

5. Lamina simply pinnate, the rachis strongly sclerotic, abaxially sharply carinate. Sp. 34. 

6. Sect. Tropidolindsaea 

5. Lamina bipinnate, or, if simply pinnate, the rachis not simultaneously sclerotic and abaxially 

carinate. 

6. Veinsanastomosing, sometimes only irregularly. 5/7/7. 15-30 . . . . 3. Sec^ Synaphlebium 

6. Veins free. 

7. Spores monolete; lamina simply pinnate, with abaxially rounded rachis and (e.xcept in small 

forms) interrupted sori. 5/7. 33 5. Sect. Osmolindsaea 

7. Spores trilete; if lamina simply pinnate and rachis abaxially rounded, the sori continuous. 

8. Lamina simply pinnate, basally distinctly reduced and/or with more widely spaced pinnules; 

petiole and rachis abaxially bi-angular; pinnules not more than twice as long as broad, or, 

if longer, deeply incised. 5/7/7. 42^3 9. 5ecf. Stenolindsaea 

8. Lamina bipinnate, or, if simply pinnate, basally not reduced nor with remote pinnules and the 
axes abaxially terete. 

9. Sori continuous; pinnules entire. 5/7/7. 31-32 4. Sect. Lindsaea 

9. Sori interrupted, pinnules - incised. Spp. 9-14 2. Sect. Temnolindsaea 

1. Rhizome epiphytic, long-creeping, the stele strongly dorsiventral. Subg. Odontoloma. 
10. Rhizome wiry, not over 1 14 mm thick, with an open xylem strand, deciduously scaly, more per- 
sistently so only near the petiole bases, lustrous when naked. Leaves simply pinnate. Veins free 

or anastomosing. Spp. 60-62 13. Sect. Penna-arborea 

10. Rhizome 1-2 mm or more thick (except in a few small species), with a closed xylem strand, more 
persistently scaly, not lustrous when naked. Leaves simply pinnate, free-veined, or bipinnate, 
free- or reticulate-veined. 
11. Lamina simply pinnate, the rachis on the adaxial side not grooved to the base (exc. often in L. 

capillacea). Spp. 44-51 10- Sect. Odontoloma 

11. Lamina bipinnate, in some species occasionally also simply pinnate leaves present, these with the 
rachis on the adaxial side grooved to the base. 

12. Pinnules entire, with uninterrupted sori. 5/7. 52 11. 5ecf. Pseudolancea 

12. Pinnules incised, with interrupted sori, or with only one short sorus near ihe apex, otherwise 
sterile. Spp. 53-59 12, Sect. Lindsaenium 



200 Flora Malesiana [ser. II, vol. P 

KEY TO THE SPECIES 

1. Lamina pinnate + deeply pinnatifid, bipinnate, or more compound, without conform terminal pinna, 

the upper (primary) pinnae gradually reduced (fig. 20, 21) Group A 

1. Lamina simple, simply pinnate, or, if bipinnate or subtripinnate, with a conform terminal pinna 

sharply set off from the upper lateral pinnae. 
2. Ultimate free divisions not, or only a very short basal portion, dimidiate. 

3. Veins anastomosing, sometimes irregularly so Group A 

3. Veins quite free. 
4. Leaf-apex and pinna-apices or the entire pinnae triangular or rhombic-triangular, acuminate 

5. L. javanensis 
4. Leaf- and pinna-apices otherwise. 

5. Pinnules articulate with the rachis Group E 

5. Pinnules continuous with the rachis {sect. Psammolindsaea) 35. L. walkerae 

2. Ultimate pinnules dimidiate. 
6. Rhizome long-creeping, epiphytic or scandent, with remote leaves; stele dorsiventrally symmetric 

Group G 
6. Rhizome short-creeping, terrestrial, or exceptionally longer and sometimes epiphytic (in case of 
doubt both choices will lead to the correct species), with radially or nearly radially symmetric 
stele. 
7. Veins free. 
8. Sori continuous in fully fertile pinnules. 
9. Petiole pale, abaxially terete; or, if upward obtusely bi-angular, the upper pinnules little re- 
duced, the terminal segment (pinnule) large, hastate, and the larger pinnules 2 cm or more 

long and over twice as long as wide Group D 

9. Petiole pale or dark, abaxially bi-angular; upper pinnules much reduced, the terminal segment 
small, not hastate; or the larger pinnules smaller and not over 1 Yi times as long as wide. 
10. Larger, basal pinnules asymmetrically suborbicular, only the basal half dimidiate (fig. 15); 
simply pinnate sterile leaves with sharply dentate pinnules often present 3. L. orbiculata 
10. Pinnules entirely dimidiate, not suborbicular; sterile leaves not usually present, their pin- 
nules not sharply dentate Group C 

8. Sori interrupted in fully fertile pinnules. 
11. Simply pinnate; rachis abaxially rounded or narrowed-rounded; spores monolete {sect. 

Osmolindsaea) 33. L. odorata 

11. Simply pinnate; rachis abaxially keeled; spores trilete (^^T/. rrop/V/o/zW^aea) 34. L. adiantoides 
11. Simply pinnate, with abaxially bi-angular rachis; or bi-(subtri-)pinnate; spores trilete. 
12. Simply pinnate; pinnules up to 2^2 times as long as wide; lamina basally mostly at least 

somewhat reduced Group F 

12. Simply pinnate; at least some pinnules over 2 Yi times as long as wide; lamina truncate at the 

base Group C* 

12. Bipinnate Group B* 

7. Veins anastomosing Group C 

Group A (sect. Schizoloma) 

1. Veins free. 
2. Pinnae in the basal part with segments only on the acroscopic side, in the apical part on both sides 

(fig. 19); lamina pinnate + deeply pinnatifid 1. L. hemiacroscopica 

2. Pinnae equal-sided, or basiscopically more compound than acroscopically; simply pinnate with 
entire pinnules, bipinnate, or more compound. 
3. Terminal segment of lateral pinnae narrow, acute, small in comparison to the pinnules next to it 
(fig. 21); indusium not or scarcely erose, falling short of the margin by less than half its width 

2. L. bouillodii 
3. Terminal segment of lateral pinnae broad, rhombic, obtuse, or sometimes acuminate-caudate, 
large in comparison to the pinnules next to it; or lamina simply pinnate; indusium falling short 
of the margin by more than half its width (except often in L. javanensis). 
4. Terminal segment of lateral pinnae (if any) as large as or often much larger than the basal pin- 
nules of that pinna, rhombic to caudate-rhombic; no simply pinnate leaves with sharply dentate 
pinnules present beside the fertile ones; texture herbaceous to chartaceous; petiole dark, abaxially 

at least upward obtusely or usually acutely bi-angular 5. L. javanensis 

4. Terminal segment of lateral pinnae about as large as the larger basal pinnules of that pinna, 
flabellate, suborbiculate (fig. 16); texture subcoriaceous to coriaceous; petiole dark, abaxially 
mostly rounded; sterile leaves with crenate pinnules often present. . . 4. L. gomphophylla 



* In case of doubt both ways will lead to the correct species. 



April 1971] LiNDSAEA-GROUP (Kramer) 201 

4. Terminal segment of lateral pinnae (if any) smaller than the larger basal pinnules of that pinna, 
rhombic, obtuse; texture herbaceous to chartaceous; petiole pale to dark, abaxially at least 
upward sharply bi-anguiar; unipinnate sterile leaves with sharply dentate pinnules often present 

3. L. orbiculata 
1. Veins anastomosing, sometimes irregularly so and then some pinnules quite free-veined. 
5. Larger secondary pinnules dimidiate, trapezoidal; pinnae without a large lanceolate undivided 

apical portion 6. L. media 

5. Free secondary pinnules (if present) flabellate, not dimidiate; pinnae entire or with a large lanceolate 
undivided apical portion. 
6. Veins even in larger pinnae (pinnules) irregularly anastomosing; upper (primary) free pinnules 

relatively wide, c. 1-3 times as long as wide 7. L. heterophylla 

6. Veins regularly anastomosing, except in small secondary pinnules (if present); upper (primary) 
pinnules mostly relatively narrower 8. L. ensifolia 

Group B (sect. Teinnolindsaea) 

1. Petiole and rachises abaxially keeled 14. L. tenuifolia 

1. Petiole and rachises abaxially bi-angular, rarely only obscurely so. 

2. Basal (primary) pinnae strongly reduced (fig. 25) 9. L. kingii 

2. Basal pinnae hardly or not reduced. 

3. Pinnules incised beyond ^,\ of their width, with capillary segments 0.2-0.4 mm wide, wider at the 

sorus, the wing connecting them as wide (fig. 28); sori uninerval .... 13. L. polyctena 

3. Pinnules at the base incised beyond the middle, the segments not capillary, 0.3-0. 8(-l) mm wide, 

connected by a wing of ^2-1 rnni; sori very predominantly uninerval ... 12. L. tetragona 

3. Pinnules at the base not incised to the middle, the segments 0.8 mm wide or more; sori usually on 
more than one vein (except in L. tetragona). 

4. Outer incisions of the pinnules reaching to the middle; lobes longer than broad. 
5. Sori uni- or binerval; largest pinnules 10-12 mm long 12. L. tetragona 

5. Sori bi- or trinerval; largest pinnules 7-8 mm long 10. L. multisora 

4. No incisions reaching beyond ^,4 of the width of the pinnules; lobes broader than long. 

6. Veins 1 mm apart 11. L. natunae 

6. Veins U rnm or less apart see Group C 

Group C {sect. Synaphlebium) 

1. Veins of larger fertile pinnules irregularly anastomosing, sometimes almost or even quite free (see 

also 3. L. orbiculata). 
2. Sori continuous; pinnules narrowed from base to apex, 5-15 mm long, 2-2^,2 irirn wide; veins 
adaxially impressed, abaxially prominulous; larger pinnules rarely entirely free-veined; sterile 
pinnules sinuate-dentate; petioles rarely pale, usually brown, abaxially obtusely to acutely bi- 
angular 27. L. crispa 

2. Sori continuous or interrupted, the incisions of the pinnules not reaching 1 mm deep in fully fertile 
pinnules; pinnules very little narrowed close to the apex, 10-12 mm long, 3^ mm wide, often 3^2-4- 
times as long as wide; veins immersed; pinnules often quite free-veined; petiole pale, abaxially flat 
or convex, bi-angular, the angles evanescing downward; sterile pinnules broadly crenate-sinuate 

16. L. napaea 

2. Sori interrupted; pinnules scarcely narrowed to the apex, 10-20 mm long, 4-6 mm wide, their 
incisions to 1 mm deep; veins immersed; pinnules often without any anastomoses; petiole pale, 
abaxially angular to the base, at least near the apex sulcate; sterile pinnules bicrenate. 

15. L. malayensis 
1. Veins of larger fertile pinnules regularly anastomosing, at least in the basal half of the pinnules. 

3. Sori of larger, fully fertile pinnules continuous. 

4. Upper pinnules little reduced, ir half as long as the larger ones; terminal segment comparatively 
large, lanceolate, free or nearly so (fig. 32b). 

5. Pinnules twice as long as wide; petiole usually reddish brown 25. L. Integra 

5. Pinnules 2^,2-3 times as long as wide; petiole stramineous 23. L. cultrata 

4. Upper pinnuFes strongly reduced, some denticuliform ones connected with the narrow pinna-apex. 

6. Pinnules not over twice as long as wide. 

7. Larger pinules 5-7 mm long, 3-4 mm wide; pinnae strongly ascending; outer veins, in small 
pinnules sometimes all or nearly all veins free; never 2 series of areoles . . . 20. L. ramosii 

7. Larger pinnules 9-15 mm long, 4-7 mm wide; pinnae laxly ascending; veins regularly anastom- 
osing; not rarely two series of areoles between lower and upper margin . . . 26. L. azurea 

6. Pinnules 2V2 to over 3 times as long as wide. 

8. Larger pinnules 16-20 mm long 24. L. papuana 

8. Larger pinnules to 12 mm long 16. L. napaea 

3. Sori of larger pinnules interrupted by incisions of the margin. 



202 Flora Malesiana [ser. II, vol. 1 



9. Terminal pinnule free, cuneate or cuneate-flabellate (fig. 34); rhizome not very short-creeping. 
10. Sori bi- to trinerval; pinnules translucent; terminal pinnule narrowly cuneate or with 2 narrow- 
ly cuneate divisions 30. L. modesta 

10. Sori quadri- to plurinerval; pinnules opaque; terminal pinnule flabellate . . 29. L. obscura 
9. Terminal segment narrow, connected with some denticuliform upper pinnules, or, if ± free, 

triangular, acute. 

11. Pinnules 2-3^2 cm long, 3-3^2 times as long as wide; simply pinnate . 17. L. subalpina 
11. Pinnules under 2 cm long, not over 3 times as long as wide; lamina usually bipinnate. 

12. Pinnules opaque, hardly narrowed to the obliquely truncate apex; outer margin distinct, with 
an incision; pinnae rather suddenly narrowed below the — caudate, pinnatifid apex (fig. 29); 
pinnule-bearing rachises abaxially brown, sulcate and pale-margined. 22. L. paralelogramma 
12. Pinnules with subacute to rounded apex, and/or distinctly narrowed to apex; or, if truncate, the 
outer margin not incised; pinnules often translucent; pinnae more gradually narrowed; pinnule- 
bearing rachises various. 

13. Pinnules twice as long as broad, to 5 mm long 28. L. hewittii 

13. Pinnules more elongate, or, if not, at least 7 mm long. 

14. Pinnae (if any) strongly ascending; pinnules 7-10 by 31/2-5 mm, 1^2-272 times as long as 

wide; indusium not reaching the margin by more than half its width . . 19. L. longifolia 

14. Pinnae (if any) not strongly ascending; pinnules larger, or, if 10 mm or less long, over iy2 

times as long as wide, or the indusium closer to the margin. 

15. At least the inner incisons reaching to V3 or 1,2 of the width of the pinnules; pinnules 

translucent; all lobes and receptacles distinctly convex; indusium reaching the margin 

or very nearly so 21. L. lobata 

15. Pinnules more shallowly incised, or, if the inner incisions reach I3, the pinnules not trans- 
lucent, or the inner or all lobes and receptacles straight. 
16. Pinnules usually with the upper margin convex, the lower concave, i.e., slightly falcately 
decurved; a distinct outer margin present, joining the upper at an angle of less than 90", 
its sorus mostly continuous with the outermost one of the upper margin; at least the inner 
incisions reaching considerably beyond the receptacle .... 23. L. cultrata 

16. Pinnules usually as above; a distinct outer margin wanting, rounded into the upper; 
most or all incisions reaching considerably beyond the receptacle . . 18. L. obtusa 
16. Pinnules usually with both margins straight or faintly convex; no distinct outer margin 
developed, rounded into the upper; incisions reaching to the level of the receptacle or 
shallower 16. L. napaea 

Group D {sect. Lindsaea) 

1. Pinnules to 11 mm long, to 5 mm wide 31. L. borneensis 

1. Pinnules to 20(-35) mm long, to 6(-13) mm wide 32. L. doryphora 

Group E {sect. Isoloma) 

1. Pinnules 5 times or more as long as broad, 3-4 mm wide (apart from basal auricles), with con- 
tiguous bases 41. L. divergens 

1. Pinnules relatively much shorter, or, if up to 5 times as long as wide, 8 mm wide (apart from basal 

auricles) and at least in the basal half of the lamina not contiguous. 
2. Pinnules suborbicular, not more than 1 mm longer than broad, coriaceous, not auricled; rachis 

black, lustrous 40. L. jamesonioides 

2. Pinnules ovate to lanceolate, the difference between length and width greater; at least some pinnules 
distinctly auricled; rachis brown to black, lustrous or not. 
3. Rachis blackish, lustrous, abaxially keeled to the base; upper pinnules little or not reduced, 

terminal division free 37. L. ovata 

3. Rachis not blackish and lustrous, or, if so, not keeled to the base; upper pinnules reduced, some 

confluent with the terminal segment. 

4. Rachis dull, medium to dark reddish brown, rarely darker or lustrous; pinnules obtuse or rarely 

subacute, the larger ones 7-18 mm long, up to 2^/2 times as long as wide . 36. L. gueriniana 

4. Rachis lustrous, reddish to dark brown; pinnules obtuse, the larger ones 5-8 mm long, less than 

1 1/2 times as long as wide 39. L. philippinensis 

4. Rachis dull, dark brown; pinnules acute or rarely subacute, the larger ones 18-30 mm long, 
to 4 times as long as wide 38. L. pellaeiformis 

Group F {sect. Stenolindsaea) 

1. Pinnules entire, or, if incised, the inner incisions of larger pinnules not going beyond the middle 

42. L. lucida 
1. Pinnules incised far beyond the middle 43. L. bakeri 



April 1971] LiNDSAEA-GROUP (Kramer) 203 

Group G (subg. Odontoloma) 

1. Rhizome wiry, not over P/4 titi 0, deciduously scaly, eventually naked, dark brown to blackish, 

polished, with open xylem strand; lamina simply pinnate but the pinnules sometimes incised (sect. 

Penna-arborea). 

2. Veins anastomosing; larger pinnules p2-3 cm long; few or no upper pinnules strongly reduced. 

3. Terminal segment narrow, lanceolate or caudate (fig. 38) 61. L. werneri 

3. Terminal pinnule broad, obtuse, cuneate or flabellate Group C 

2. Veins free, or, if anastomosing, the largest pinnules less than 1 |o cm long; upper pinnules gradu- 
ally and strongly reduced. 

4. Pinnules deeply pinnatitid. the segments connected by wings 14^^ 2 nrn wide ffig. 23j 

62. L. roeraeriana 

4. Pinnules shallowly incised, or, if more deeply incised, the lobes much more broadly connected. 

60. L. pulchella 
1. Rhizome not wiry, Xlo-- nim o, or, if thinner, not polished when naked, with closed, strongly 

dorsiventral xylem strand. 
5. Lamina always simply pinnate (species where this character fluctuates can be keyed out both ways). 
6. Lamina truncate at the base, with a well-developed petiole and few or no reduced basal pinnules; 
pinnules incised considerably beyond the receptacle. 

7. Veins anastomosing Group C 

7. Veins free. 
8. Pinnules very regularly incised (fig. 50); sori distinctly intramarginal, mostly uni-, some binerval. 

47. L. apoensis 
8. Pinnules irregularly incised; sori uni- to quadrinerval, very close to the margin, the sporangia 

often spreading beyond it at full maturity 46. L. repens 

6. Lamina gradually and strongly narrowed at the base, the petiole often very short or virtually 
absent; pinnules subentire to variously incised. 
9. Pinnules incised far beyond the middle; sori uni- or binerval. 
10. Pinnule-lobes truncate, their outer margin erose to corniculate (fig. 46, 47). 
11. Rhizome ^2 mm 0; ultimate pinnule-lobes 0.4-0.8 mm wide near the apex, there laterally 
with two horn-like projections 0.3-0.4 mm long (fig. 46); sori uninerval. . 51. L. capillacea 

11. Rhizome 3^-2 mm 0; ultimate pinnule-lobes ^2-2 mm wide near the apex, there laterally 
without, or with rudimentary horn-like projections (fig. 47); sori not rarely binerval 50. L. fissa 

10. Pinnule-lobes rounded or narrowed to subacute, not truncate. 

12. Pinnule-lobes 0.3-1 mm wide, nearly all of them once or twice bifid, joined by wings of less 
than 1. mm width (fig. 48) 49. L. carvifolia* 

1 2. Pinnule-lobes 1 mm or more wide, joined by wings of 1 mm or more (fig. 49) 44. L. glandulifera 
9. Pinnules subentire to incised to the middle; sori uni- to plurinerval. 
13. Sori continuous (shortly interrupted in very shallowly incised, incompletely fertile pinnules); 

rhizome scales chocolate brown 45. L. oblanceolata 

13. Sori interrupted (beware of old, abundantly fertile pinnules where the sori may be confluent); 

scales golden brown (darker in some varieties of L. repens with more deeply incised pinnules). 

14. At least some of the fertile lobes denticulate or erose; sterile lobes acute, tooth-like (fig. 42). 

48. L. merrillii 
14. Fertile lobes not denticulate, erose, or tooth-like, but truncate, rounded, or narrowed- 
rounded. 
15. Larger pinnules 10-12 mm long; pinnules incised to 1^3 or V2. the lobes evenly narrowed from 

base to apex (fig. 49) 44. L. glandulifera 

1 5. Larger pinnules 1 5 mm or more long, with truncate or apically rounded lobes, or, if shorter, 
much less deeply incised, or the lobes n parallel-sided 46. L. repens** 

5. Lamina of all, most, or at least some leaves bipinnate. 

16. Sori continuous 52. L. parasitica 

16. Sori interrupted. 

17. Secondary rachises abaxially rounded 54. L. monocarpa 

17. Secondary rachises abaxially carinate 55. L. sarawakensis 

17. Secondary rachises abaxially bi-angular (and/or sulcate) at least in the upper half. 

18. Pinnules coriaceous; veins sometimes anastomosing 53. L. ngida 

18. Pinnules herbaceous or chartaceous. 
19. Pinnules shallowly incised, at the most to the receptacle or slightly beyond, never to the 
middle. 



* Simply pinnate forms of L. rosenstockii and L. versteegli will also run to this heading. The former 
has only the basal pinnule-lobes bifid, the latter has more divergent lobes (see fig. 39 and 40). 

** Simply pinnate forms of L. microstegia will also run here; their basal pinnules are inserted on the 
edge of the adaxial face of the rachis, not below it, as in L. repens. 



204 Flora Malesiana [ser. II, vol, 1 ^ 

20. Adaxial groove of the primary rachis broad, occupying half of its width or more; most 
pinnules fertile only near the apex; veins sometimes anastomosing; indusium not reaching 
the margin by less than its width; bipinnate 53. L. rigida* 

20. Adaxial groove of the primary rachis narrow, occupying much less than half its width; 
most pinnules entirely fertile; veins free; indusium not reaching the margin by less than its 
width; bipinnate 56. L. regularis 

20. Adaxial groove of the primary rachis narrow, occupying much less than half its width; most 
pinnules entirely fertile; veins free; indusium not reaching the margin by 1 Vo-3 times its 
width; not rarely simply pinnate leaves present alongside the bipinnate ones 57. L. microstegia 

19. Pinnules incised considerably beyond the middle. 

21. Segments capillary, c. ^,'5-^4 mm wide, all forked 59. L. versteegii 

21. Segments not capillary %-2 mm wide, the outer ones often not forked 58. L. rosenstockii 

Subgenus Lindsaea 
1. Section Schizoloma 

(Gaud.) Kramer, Act. Bot. Neerl. 15 (1967) 51\.—Schi:oIomci Gaud. Ann. Sc. Nat. 3 (1824) 
507. 

Type species: Schizoloma hillardicri Gaud. (= Lindsaea ensifolia Swartz). 

Distr. Pantropic, extending far into the temperate zone in Japan, Australia, and New Zealand. Many 
species concentrated in Madagascar. 

Taxon. The short-creeping rhizome and the lamina lacking a conform terminal pinna characterize 
nearly all species. L. orbiciilata (Lamk) Mett. ex Kuhn is sometimes only simply pinnate, and L. 
ensifolia Swartz ssp. ensifolia is simply pinnate with a conform non-dimidiate terminal pinna, but 
otherwise the section is quite clear-cut. It seems to be the most primitive in the genus, with the 
possible exception of the New Caledonian sect. Davalliastruni. Several other subdivisions of the genus 
comprise one or a few species that show definite affinity with sect. Schizoloma: among the most in- 
teresting are some Madagascan species with the rhizome of subg. Odontoloma but a Schizoloma-like 
leaf architecture. 

1. Lindsaea hemiacroscopica Kramer, Blumea 15 to trinerval; indusium greyish, '2-1^2 mTi long, 

(1968) 563. — Type: Hallier 3244, Mt Amai 0.4 mm wide, suborbicular to oblong, free at the 

Ambit, Borneo, Kalimantan (BO). — Fig. 19. narrowed sides, subentire, not reaching the margin 

Rhizome short-creeping, c. 0.6 mm 0; scales by a little less than its width, not reflexed at 

reddish brown, c. Vs cm long, narrowly lanceo- maturity. Spores medium brown, trilete, smooth, 

late, biseriate at base, the apical uniseriate part c. 22 //. 

consisting of 1-3 cells. Leaves close; petioles Distr. Only known from type collection. No 

2-4 cm long, somewhat shorter than the lamina, ecological data, 
adaxially pale, channelled, abaxially dark brown, 

sharply bi-angular with narrowly paler edges 2. Lindsaea bouillodii Christ, Not. Syst. 1 (1909) 

and flat faces, subterete at base. Lamina herbace- 59. — Type: Bouillod 48, Cam-chay Mts, Cam- 

ous, olivaceous when dry, ovate with truncate bodia (P). 

base, to 5-6 cm long, 3-4^2 cm wide, pinnate — L. orbiculata (Lamk) .Mett. ex Kuhn var. 

deeply pinnatifid, without conform terminal pinna. odontosorioides Copeland, Philip. J. Sc. 6 (1911) 

Mayor /^m/iae c. 10 to a side, spreading or slightly Bot. 138. — Type: Brooks 19, Tringos, Sarawak 

ascending, less than their width apart, the larger (MICH). 

ones 20 by 7 mm, acuminate, inequilateral, L. orbiculata (Lamk) Mett. ex Kuhn var. 

deeply pinnatifid. Segments cuneate to ligular, sumatrana Rosenstock in Fedde, Rep. 13 (1914) 

2 or in larger pinnae 3 on the acroscopic side, 214. --Type: Winkler 55 (Ros.-exs. 117), Batak 

3-4 by 2 mm, with only a very narrow wing on Lands, Sumatra (S-PA; dupl. in BM, L, P). 

the opposite, basiscopic side; upper segments of L. cambodgensis and. nan Christ; Kramer, 

acroscopic and all (or all but one) of basiscopic Blumea 15 (1968) 563. 

side narrowly cuneate, all narrowed at base and L. tenera auct. non Dryand. (or Schizoloma 

connected by wings of leaf-tissue; uppermost tenerum) of other authors, e.g., Holttum, Gard. 

pinnae and segments reduced, confluent. Apex Bull. S. S. 5 (1930) 64; Rev. Fl. Mai. 2 (1954) 

of segments rounded-truncate, entire or sinuate; 348, f. 201. — Fig. 21. 

sterile segments often subacute. Largest segments Rhizome moderately to very short-creeping, 

often bilobed. Costae evident, stramineous; 1 '2-2 mm o; scales ferrugineous, very narrowly 

veins immersed, evident, single or in larger triangular, to 2^2 mm long, to 5-seriate at base, 

lobes paired, one sometimes forked. Sori uni- with long uniseriate apex. Leaves close to cluster- 



* Epiphytic specimens with unusually long rhizomes of species o^ sect. Synaphlebium (Group C) will 
run to this heading; they have regularly anastomosing veins. 



April 1971 



LlNDSAEA-GROUP (KramcTj 



205 




Fig. 15. Lindsaea orbiculata CLamk) Mett. ex KuHN var. orbiculata. Plant, x I/3, pinnule, X 134 
(Tanaka & Shimada 13522). — Fig. 16. L. gomphophyila Baker. Lamina, x 1/3 (Hose? s.n., Sar). — Fig. 
17-18. L. heterophylla Dry.and.; fig. 17. Lamina, x 1/3 CBakhuizen van den Brink 7153); fig. 18. 
Sterile pinnule, nat. size (Petersen s.n., L). — Fig. 19. L. hemiacroscopica Kramer. Basal pinna, x 2 

(Hallier 3244). 



206 



Flora Malesiana 



[ser. II, vol. 1 



ed; petioles 8^0 cm long, to 1 V^ times as long 
as the lamina, sharply quadrangular, dark reddish 
brown, atropurpureous, or blackish, ± pale- 
margined. Lamina ovate or triangular, 12-30 cm 
long, bipinnate to (at the base) tripinnate + pinna- 
tifid. Primary rachis dark, pale-margined ± sul- 
cate. Major primary pinnae 3-8 to a side, spreading 
or (mostly not strongly) ascending, subsessile, 
narrowly oblong, the largest 8-11 cm long, 
ly^-lYi cm wide, the lower ones up to their 
width apart, the upper ones closer; upper pinnae 
gradually reduced, rather abruptly passing into 
what is almost a conform terminal pinna. Sec- 
ondary rachises abruptly pale, often ± green- 
margined. Lower primary pinnae not rarely at 
the base with some pinnate secondary pinnae to 
c. 2 cm long, rarely the greater part of the lamina 
with pinnate secondary pinnae with green-mar- 
gined rachises. Pinnules c. 1-\1 to a side, herbace- 
ous, dark green when dry, spreading or slightly 
ascending, not contiguous, subsessile, dimidiate- 
ovate or less often subtrapeziform, the larger ones 
10-13 mm long, 5-7 mm wide, 1 Y2.-2 times as long 
as wide (the basal pinnules of the terminal pinna 
up to 15 by 10 mm), the upper (and, if present, 
the outer) margin incised, mostly the upper margin 
with 1 or 2, the outer with 1 incision, sometimes the 
lobes shallowly incised again; incisions Y-y-^ V2 rmri 
deep, rarely more, the sinus acute; pinnules rarely 
pinnatifid, with cuneate lobes. Fertile lobes some- 
what convex, usually slightly erose; sterile pinnules 
(not rare) with more and narrower, longer, 
subacute or acute lobes. Upper pinnules reduced, 
a few confluent with the narrow, acuminate or 
subcaudiform terminal segment; transitions 
between pinnate and non-pinnate pinnae in the 
leaf-apex few. Veins immersed, mostly little 
evident, free, Y2-% mm apart, once or twice 
forked. Sori interrupted by the incisions of the 
margin, convex but scarcely extending onto the 
sides of the lobes, often 2-4 mm long and 2- to 
4-nerval, rarely uninerval; indusium greyish or 
brownish, subentire or mostly slightly erose, 0.3- 
0.4 mm wide, almost or quite reaching the margin, 
not reflexed at maturity. Spores pale brown, 
trilete, smooth, c. 22 /<. 

Distr. Thailand and Tonkin to Malesia: 
Malay Peninsula, Sumatra, West Java, Borneo, 
Banka, and Natuna Is. 

Ecol. Terrestrial in (usually moist) forests, 
300-1400 m. 

Note. The tripinnate form has been described 
as L. orbiculata var. odontosorioides, but it does 
not seem to merit taxonomic recognition. 

3. Lindsaea orbiculata (Lamk) Mett. ex Kuhn 
in Miq. Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 
279; HoLTTUM, Gard. Bull. S. S. 5 (1930) 64; 
Tagawa, Act. Phytotax. Geobot. 6 (1937) 33, 
f. 3 D-G; CoPELAND, Fern Fl. Philip. 1 (1958) 
112, p.p. — Adiantum orbiculatum Lamk, Encycl. 1 
(1783) 41.— Schizoloma orbiculatum (Lamk) 
Kuhn, Chaetopt. (1882) 346; Holttum, Rev. 
Fl. Mai. 2 (1954) 344, f. 199. -Schizolegnia 
orbiculata (Lamk) Alston, Bol. Soc. Brot. II, 



30 (1956) 24. — Type: Sonnerat s.n., near Ma- 
lacca (P). 

L. flabellulata Dryand. Trans. Linn. Soc. 3 
(1797) 41, pi. 8 f. 2.-Lectotype: Nelson s.n., 
Macao (BM). 

L. polvmorpha Wall, ex Hooker & Greville, 
Ic. Fil. (1828) pi. 75. -L. flabellulata Drvand. 
var. polymorp/ia (Hook. & Grey.) Hooker, Sp. 
Fil. 1 (1846)'211.-L. orbiculata (Lamk) Mett. ex 
Kuhn var. polvmorpha (Hook. & Grey.) v.A.v.R. 
Handb. (1908) 270. —Type: a specimen without 
data from Herb. Hooker (K). 

L. montana Copeland in Perkins, Fragm. Fl. 
Philip. (1904) 182, non Fee (1866). -L. cooelandi 
C. Chr. Ind. Fil. (1906) 392. -Type: Copeland 
230, Mt Mariveles, Luzon (MICH; dupl. in B). 

L. bonii Christ, Not. Syst. 1 (1909) 187.— 
Type: Bon 8, S. Tonkin (P). 

L. longipes C. Chr. & Tardieu-Blot, Not. 
Syst. 5 (1936) 263; Fl. Gen. I.-C. 7 (1939) 125, 
f. 15 3-4.— Type: Poilane 8208, Nhatrang, 
Annam (P). 

Rhizome short-creeping, 1-1 Y2 mm 0; scales 
reddish brown, very narrowly triangular, to c. 
2 mm long, to 4-seriate at base, a considerable 
apical portion uniseriate. Leaves close; petioles 
stramineous with dark base or dark to blackish 
brown throughout, abaxially at least in the upper 
part sharply bi-angular and if dark pale-mar- 
gined. Lamina simply pinnate or bipinnate, her- 
baceous or chartaceous; pinnules dimidiate, very 
variable in shape, erose, free-veined. Upper pinnae 
of bipinnate leaves gradually to rather abruptly 
shortened, i gradually passing into the non- 
conform terminal pinna. Terminal pinnules (seg- 
ments) ± rhombic. S'on continuous or interrupted; 
indusium minutely to strongly and irregularly 
erose, Yi mm wide, usually falling short of the 
margin by half its width, rarely more strongly 
intramarginal or almost reaching the margin. 
Spores yellow to light brown, trilete, almost 
smooth, c. 25-30 //. 

KEY TO the varieties 

1. Plants usually (always?) with simply pinnate 
sterile leaves beside the fertile ones; fertile 
leaves simply pinnate; or, if bipinnate, at 
least some pinnules at the base of the terminal 
pinna, above the uppermost pinnate pinnae, 
suborbicular; the lamina not gradually passing 
from the bipinnate condition at base to the 
simply pinnate apex but with a rather abrupt 
transition .... 1. var. orbiculata 

1. Sterile simply pinnate leaves mostly wanting; 
no pinnules suborbicular; larger laminas up- 
ward gradually of simpler structure, with a 
gradual transition from the bipinnate base 
to the simply pinnate apex 2. var. commixta 

1. var. orbiculata. — Fig. 15. 

Sterile, simply pinnate leaves mostly, perhaps 
in nature always, present, together with fertile 
ones. Petioles of sterile leaves slender, stramineous 
to reddish brown or rarely darker, 2-8 cm long, 



April 1971] 



LiNDSAEA-GROUP (Kramer) 



207 



mostly about half as long as the lamina; lamina 
linear, c. 5-10 cm long with 6-12 pinnules to a 
side (larger ones mostly fertile in the upper part), 
slightly narrowed from base to apex, there sudden- 
ly narrowed. Rachis pale, abaxially bi-angular 
or sulcate. Pinnules mostly olivaceous or dark 
green when dry, chartaceous, asymmetrically 
ovate or V^-elliptic, often %-l by Vi; cm, often 
subcontiguous, spreading or the basal ones de- 
curved, the outer and upper margin sharply 
dentate, the teeth up to 1 mm long; terminal 
pinnule (segment) free or nearly so, often 1-1 Va 
cm long and Yo-% cm wide, obtuse, less often 
acute, cuneate-flabeliate, toothed, the pinnules 
just below it mostly not strongly reduced. Fertile 
leaves more numerous, simply pinnate or bipinnate, 
or very often subbipinnate; intermediates between 
sterile and fertile leaves not rare. Petioles of fertile 
leaves stouter than those of sttrile ones, c. 10-30 
cm long, as long as the lamina or in very large 
leaves sometimes much shorter. Lamina linear if 
once pinnate or subbipinnate, otherwise very 
variable in shape, depending on the number 
and size of the pinnae, c. 12-50 cm long. Pinnules 
herbaceous or more often chartaceous, olivaceous 
or dark green when dry, very variable in number, 
shape, and size, the lower ones often remote, 
the upper ones gradually closer, subcontiguous; 
rachis basally sometimes dark, otherwise stramine- 
ous, abaxially bi-angular, upward sulcate; second- 
ary rachises, if any, similar. Fertile pinnules like the 
sterile ones or at least the lower ones of simply 
pinnate leaves, simply pinnate leaf-apices, or larger 
pinnate pinnae, very asymmetrically flabellate- 
suborbicular, with broadly rounded upper/ 
outer and ± concave, descending lower margin, 
then often 1-1 V2 cm long and broad; in sub- 
bipinnate laminas some of the lower (but not 
necessarily the lowermost) pinnae pinnatifid 
or pinnate, with some cuneate-flabeliate or Y^- 
elliptic pinnules on both sides and a large, rhombic, 
very obtuse, often very asymmetric terminal 
pinnule. Fully bipinnate leaves with up to c. 8 
well-developed pinnae to a side, these spreading 
almost at right angles, with up to c. 10 pinnules 
to a side, the terminal pinnule smaller than in 
subbipinnate leaves but not much smaller than the 
lateral pinnules of the same pinna, obtuse. 
Transitions between pinnate pinnae and lobed 
pinnules nearly always present at the base of the 
relatively very long simply pinnate terminal 
portion of bipinnate leaves, occasionally also 
between and even below fully pinnate pinnae. 
Edge of pinnules erose-crispate, rarely dentate. 
Veins immersed, evident, close, 1-3 times forked, 
flabellate, a costa not or scarcely developed. 
Sori continuous except as interrupted by incisions 
of transitional pinnules. 

Distr. SE. China, S. Japan, to Malesia: 
Malay Peninsula, Singapore, Sumatra, Banka, 
Java, Bawean, Sarawak, Sulu Archipelago, Philip- 
pines (Luzon); incorrectly reported from Mada- 
gascar. 

Ecol. In thickets and open forests, on banks 
and rock faces, often in locally dry situations 



in ravines or by rivers or near the coast, to 500 
(rarely to 1000) m. 

2. var. commixta (Tagawa) Kramer, comb. 
nov.—L. commixta Tagawa, Act. Phytotax. 
Geobot. 6 (1937) 37, f. 3 H-J.-L. tenera Dryand. 
var. commixta (Tagawa) Iwatsuki, Act. Phyto- 
tax. Geobot. 19 (1961) 6. -Type: Tasiro s.n., 
I. Tane-ga-shima, Osumi Prov., Kyushu (KYO, 

rt.V.). 

? L. montana Copeland. — L. copelandi C. Chr. 

Differing from var. orbiculata in the following 
characters: Sterile simply pinnate leaves mostly 
wanting. Fertile leaves never simply pinnate; 
suborbiculate pinnules none, even at the base of the 
simply pinnate leaf-apex, i.e., all pinnules distinctly 
dimidiate; terminal segments larger and/or more 
acute. Larger laminas fully bipinnate, more evenly 
narrowed from base to apex, the terminal pinna 
hardly set off from the rest of the lamina, the 
simply pinnate apical portion relatively shorter; 
sterile pinnules more deeply incised; lowest non- 
subpinnate pinnules of terminal part of lamina 
shallowly incised, with interrupted sori. Rhizome 
scales and spores as in var. orbiculata. 

Distr. SE. China, India, Ceylon, Indo-China, 
S. Japan, Thailand, to Malesia: Malay Penin- 
sula, Sumatra, Philippines (Luzon), Celebes, 
Sumba, Sarawak. 

Ecol. As var. orbiculata. 

Notes. L. orbiculata is one of the most variable 
species of the genus, resulting in a comparatively 
extensive synonymy. The two varieties distinguish- 
ed here are not entirely sharply distinct, partly 
because depauperate plants from rock fissures etc. 
often cannot be assigned with certainty to one of 
them, e.g. the type of L. montana Copeland, 
partly because the differences between them are 
of a gradual rather than fundamental order. 
Nevertheless the majority of the specimens can be 
assigned to one of them. Var. commixta is more 
common to the North of Malesia, particularly 
in Japan, where it is fairly sharply distinct from 
var. orbiculata but is not entirely sharply 
distinguishable from L. chienii Ching which I 
prefer to treat as a species but which may also 
prove to be only a variety of L. orbiculata. Many 
more field and also cytotaxonomic observations 
are needed before the status of the species of this 
group can be ascertained. 

4. Lindsaea gomphophylla Baker, Ann. Bot. 5 
(1891) 204.— Type: H. Low s.n., Borneo, prob. 
Sarawak (K).-Flg. 16. 

In most respects like a subbipinnate form of 
L. orbiculata, characterized by the following 
combination of characters: 

Rhizome scales castaneous, very narrowly 
triangular, to 3 mm long, to 6-seriate at base, 
a considerable apical part uni- and biseriate. 
Lamina subcoriaceous to coriaceous; at least 
some leaves of each plant subbipinnate, with a 
few paucijugate pinnae at base, rarely the basal 
pinnae more fully pinnate; terminal pinnules of 
lateral pinnae large, flabellate-suborbicular. 



208 



Flora Malesiana 



[ser. II, vol. 1 ' 



Petiole and rachis dark, abaxially terete or the 
rachis obtusely bi-angular; secondary rachises 
abruptly pale. Sterile margin less acutely dentate. 
Spores c. 30 //. 

Distr. Malesia: Borneo (Sarawak, Brunei), 
Sumatra (5 or 6 coll. in all). 

Ecol. One record 300-400 m. 

Note. This may be another variety of L. or- 
bicidata. 

5. Lindsaea javanensis Blume, En. PI. Jav. (1828) 
2\9.—Schizoloma javanense (Blume) Holttum, 
Rev. Fl. Mai. 2 (1954) 349, f. 202.- Schizolegnia 
javanensis (Blume) Alston, Bol. Soc. Brot. 
II, 30 (1956) 24. -Type: Blume s.n., Java (L). 

L. flabellulata Dryand. var. gigantea Hooker, 
Sp. Fil. 1 (1846) 211, pi. 63 c.-L. orbicidata 
(Lamk) Mett. ex Kuhn var. gigantea (Hooker) 
Mett. ex Kuhn in Miq. Ann. Mus. Bot. Lugd.- 
Bat. 4 (1869) 219.— L. tenera Dryand. var. 
gigantea (Hooker) Holttum, Gard. Bull. S. S. 
5 (1930) 65.— £. gigantea (Hooker) C. Chr. 
Bot. Jahrb. 66 (1933) 53. — Lectotype: Griffith 
s.n., Khasya and Assam (K). 

Rhizome short-creeping, 1-2 mm 0; scales 
medium brown, lanceolate, long-acuminate, to 
1 ^2 nim long, to 4-seriate at base, with a long 
uniseriate apex. Leaves clustered; petioles 10-30 
cm long, especially in large leaves much longer 
than the lamina, dark reddish brown to atro- 
purpureous, ± lustrous, abaxially terete at base, 
upward gradually obtusely, at apex mostly acutely 
bi-angular, the angles and especially the borders 
of the adaxial groove often pale. Lamina her- 
baceous to chartaceous, dark green or olivaceous 
when dry, very variable, triangular or oblong or 
in small leaves sometimes transversely triangular, 
7-20 cm long, 6-15 cm wide, 1 V4 to (in large 
leaves) 1 ^,4 times as long as wide, simply pinnate 
to amply bipinnate, in the first case with up to 
as little as 2 pairs of lateral pinnules and a distinct 
but not conform terminal one, in the second case 
with up to 5 (pinnate) pinnae to a side and several 
simple ones above, which are gradually reduced 
to and confluent (or not) with the terminal pinnule 
(segment). Primary rachis dark, adaxially with a 
narrow, pale-edged groove, abaxially flattened, 
pale-angled. Primary pinnae their width apart to 
contiguous, spreading or in plurijugate bipinnate 
leaves ascending. Pinnules of simply pinnate or 
subbipinnate leaves rhombic, very often with 
long-acuminate to caudate apex, with very un- 
equal base, basiscopically much more cut away, 
to c. 8 by 2 cm; apex of lamina similar but ± 
symmetric at base; apices of fully pinnate pinnae 
similar but smaller, progressively smaller and less 
pointed as there are more secondary pinnules; 
larger transitional pinnules between pinnate pin- 
nae and the lamina apex often rhombic-caudate, 
smaller ones subtrapeziform or subflabellate, 
very obtuse; pinnules of fully pinnate pinnae up 
to 8 to a side, mostly not contiguous, sub- 
trapeziform, rounded-rhombic or subflabellate. 
Secondary rachises abaxially flattened or slightly 
sulcate, abruptly pale at their insertion on the 



dark primary rachis, usually distinctly green- 
margined. Fertile pinnule-margin subentire or 
minutely erose, in larger leaf segments here and 
there incised by shallow crenations; sterile margin, 
especially near the segment bases, sharply serrate 
or dentate, with deeper incisions. Lobes of pointed 
pinnules often slightly concave. Veins immersed 
but ± evident, free, mostly twice forked, c. 1 mm 
apart; larger pointed pinnules and terminal divi- 
sions with a percurrent costa. Sori continuous 
in small pinnules, progressively more interrupted 
in larger ones, bi- to plurinerval. Indusium pale, 
subentire to erose, 0.3-0.4 mm wide, almost 
reaching the margin to falling short of it by 
more than its width, little reflexed at maturity. 
Spores light brown, trilete, smooth, c. 25 ji. 

Distr. Assam, SE. China, and S. Japan to 
Malesia: Malay Peninsula, Sumatra, West Java, 
Borneo, Celebes (?), Philippines (Sibuyan, Min- 
doro). Apparently nowhere common. Incorrectly 
reported from Madagascar. 

Ecol. Terrestrial in forests, 80-1400 m, mostly 
above 800 m. 

Notes. The only collection from Celebes, 
Kjellberg 3527a (BO, S-PA) is not well developed 
and also resembles L. orbiculata var. commixta. 

The specimens from the continent North of the 
Malay Peninsula have more intramarginal indusia 
and are in several respects close to L. chienii 
Ching; there is probably some introgression be- 
tween the two species. 

6. Lindsaea media R.Br. Prod. Fl. Nov. Roll. 

(1810) \56. —Schizoloma medium (R.Br.) Kuhn, 
Chaetopt. (1882) 346.— Schizoloma ensifolium 
(Swartz) J. Smith var. medium (R. Br.) Domin, 
Bibl. Bot. 20 (1915) 78. -Type: R. Brown s.n., 
N. coast of Australia (K; several other authentic, 
possibly isotype coll. in K, P, U). 

L.subtripinnatu CoPELAND, J. Arn. Arb. 24 (1943) 
441. -Type: Brass 8491, Tarara, W. Div., Papua 
(MICH; dupl. in BO, GH, L).-Fig. 20. 

Rhizome rather long-creeping, 1-1 ^2 mrn 0, 
rather thinly and deciduously scaly; scales 
yellow, ovate-triangular, to c. 1 mm long, to 
6-seriate at base, a short apical portion uni- 
seriate. Leaves to V2 cm apart; petioles stramineous 
to fawn-coloured, 10^0 cm long, mostly longer 
than the lamina, abaxially terete below, upward 
gradually obtusely or acutely bi-angular and 
flattened. Lamina herbaceous or chartaceous, 
10-30 cm long, 4-17 cm wide, 2-3 times as long 
as wide, triangular or oblong, bipinnate or bi- 
pinnate + pinnatilobate or bipinnate -|- pinnat- 
ifid, rarely tripinnate at base; leaf apex gradu- 
ally of simpler structure. Primary rachis abaxially 
flattened, bi-angular. Pinnae spreading or slightly 
ascending, the major ones c. 4— 10 to a side, most or 
all subopposite, the largest, basal ones 2V2-IO cm 
long and 12-18 mm wide, not narrowed at the 
base, rather evenly narrowed in the upper half or 
throughout; secondary rachises abaxially flat- 
tened, bi-angular, green-margined to the base or 
almost so. Basal pinnules on both sides of lower 
pinnae of large leaves usually pinnatilobate to 



April 1971 



LiNDSAEA-GROUP (Kramer) 



209 




^t^ 



^M 




;4*»«**^^ 




Fig. 20. Lindsaea media R.Br. Lamina, X V5 (Brass 8491). -Fig. 21. L. bouillodii Christ. Lamina, 

X 2/5 (Cuming 399). -Fig. 22. L. bakeri (C. Chr.) C. Chr. vw. ftaAe/-/. Lamina, x ^U (Brass 13651). - 

Fig. 23. L. roemeriana RosENSTOCK. Leaf, X ^/^ (v. Romer 1137). 



210 



Flora Malesiana 



[ser. II, vol. 1 ^ 



pinnatifid, rarely fully pinnate, then with few 
pinnules. Ultimate pinnules variable in size and 
shape, largely depending on the degree of dissec- 
tion of and their place in the lamina, but distinctly 
dimidiate-subflabellate; larger ones trapezoidal, 
subquadratic, subsessile, the larger, not dissected 
ones up to 5 by 31/2 to 10 by 6 mm, if dissected 
the smaller ones with incisions only on the 
acroscopic side, the larger ones on both sides. 
Upper pinnules reduced, not strongly so in pau- 
cijugate pinnae, the terminal segment then 
obliquely rhombic, obtuse, free or almost so, 
to 5 mm long, more strongly reduced in plurijugate 
pinnae, the upper pinnules denticuliform, con- 
fluent into a narrow, pinnatifid pinna-apex. 
Margin of sterile pinnules sharply dentate, of 
fertile ones obscurely or mostly distinctly erose. 
"Veins immersed, usually not evident, 1-3 times 
forked, c. Yi mm apart, free, connivent, or spo- 
radically and irregularly anastomosing; leaves of 
adult plants hardly ever without any anastomoses, 
but often many pinnules, especially smaller ones, 
quite free-veined. Sori continuous except as 
interrupted by incisions of the margin; indusium 
pale, erose to gashed, almost reaching to slightly 
exceeding the margin, 0.3-0.5 mm wide, not 
reflexed at maturity. Spores medium brown, 
trilete, smooth, c. 25 //. 

Distr. N. Queensland; Malesia: Papua (one 
coll., type of L. subtripinnata). 

Ecol. Bank of gully in rain forest (Papua); 
grassy places and forests, 0-425 m (Australia). 

7. Lindsaea heterophylla Dryand. Trans. Linn. 
Soc. 3 (1797) 41, pi. 8 f. l.—Adiantum hetero- 
phyllum (Dryand.) Poiret, Encycl. Suppl. 1 
(1810) 139, non Colenso (1888).— ^r/z/zo/oma 
heterophyllum (Dryand.) J. Smith, Hook. J. 
Bot. 3 (1841) 414; Holttum, Rev. Fl. Mai. 2 
(1954) 345. — L. ensifolia Swartz var. heterophylla 
(Dryand.) Benth. Fl. Austr. 7 (1878) 722.— 
Schizoloma ensifolium (Swartz) J. Smith var. 
heterophyllum (Dryand.) Domin, Bibl. Bot. 20 
(1915) 77. — Schizolegnia heterophylla (Dryand.) 
Alston, Bol. Soc. Brot. H, 30 (1956) 24.— non 
L. heterophylla Prentice (1873). — Type: Robert- 
son s.n., Malacca (BM). 

L. variabilis Hooker & Walker Arnott, Bot. 
Beech. Voy. (1838) 257, pi. 52.— Type: Millett 
s.n., Macao {n.v.). — Fig. 17, 18. 

Rhizome rather to very short-creeping, 1 1/2-3 
mm 0; scales medium brown, very narrowly 
triangular, to lYz mrn long, to 7-seriate at base, 
long-acuminate, the apical uniseriate part rather 
long. Leaves close; petioles (3-) 10-50 cm long, 
1/2-2 times as long as the lamina, stramineous to 
dark brown, adaxially often paler, abaxially 
bi-angular, sharply so above, sometimes pale- 
angled. Lamina simply pinnate to bipinnate or 
rarely to subtripinnate, narrowly oblong or oblong 
or, especially if bipinnate, deltoid, (6-) 10-45 cm 
long, (3-)6-20 cm wide, 2-6 times as long as wide, 
with 4-25 primary divisions to a side. Primary 
rachis mostly stramineous, abaxially sharply 
bi-angular or broadly and shallowly sulcate. 



Primary pinnae rather remote, the upper ones 
closer, spreading or, if pinnate, ascending, 
herbaceous to chartaceous, medium green or 
olivaceous when dry. Simple primary pinnae 
lanceolate or elongate-triangular or the smaller 
ones rhombic, the base unequal, basiscopically 
narrower, acroscopically in extreme cases cordu- 
late, the larger ones 2-10 cm long, %-iy2 cm 
wide, 1 y^-S times as long as wide; larger pinnate 
primary pinnae 8-18 cm long, (%-)l/2-6 cm 
wide, 2-9 times as long as wide, triangular to 
linear, with 1-15 pinnules, these Yo-S cm long, 
cuneate-subflabellate to suborbicular or of the 
same shape as undivided primary pinnae, spread- 
ing or somewhat ascending; secondary rachises 
abaxially subterete to bi-angular, stramineous, or 
reddish and pale-margined, at least upward green- 
margined. Upper pinnae of lamina gradually 
reduced, the upper ones usually rhombic, subacute 
or obtuse, less often suborbicular or flabellate, 
usually not less than Yz cm long, a few connected 
with the mostly comparatively large, lanceolate 
or elongate-triangular, obtuse to acuminate ter- 
minal segment; apices of pinnate pinnae similar. 
Pinnate pinnae, if any, occurring basally in the 
lamina, but sometimes above pinnae of simpler 
structure, but not as irregularly arranged as 
sometimes in L. orbiculata. Transitions between 
pinnate and simple pinnae usually paucijugate- 
pinnate with large terminal segment rather than 
lobed to pinnatifid. Margin of sterile pinnules (very 
rare in adult plants) crenate-dentate to subentire; 
fertile margin subentire or mostly ± erose, little 
sclerotic. Ultimate divisions, if elongate, with a 
percurrent, distinct, stramineous, abaxially pro- 
minulous costa; veins oblique, less so towards the 
margin, 1-3 times forked, immersed but evident, 
irregularly anastomosing, forming (except in the 
bases of large pinnae) an interrupted series of 
areoles V2-I mm wide (very rarely a second 
series), or free, but scarcely a leaf without any 
anastomoses. Sori continuous except as interrupt- 
ed by the incisions, in pinna-apices sometimes 
interrupted by serrations, in lobed pinnae often 
extending close to or around the bottom of the 
sinus. Indusium pale, entire to erose, 0.3-0.5 mm 
wide, falling short of the margin by half its width 
to reaching it, little reflexed at maturity. Spores 




Map 3. 



Distribution of Lindsaea heterophylla 
Dryand. 



April 1971] 



LiNDSAEA-GROUP (Kramer) 



211 



pale yellowish, trilete, smooth, c. 30 i-i (sometimes 
irregular and apparently abortive, which might be 
connected with hybrid origin). 

Distr. Mascarene Is., Madagascar; S. India, 
Ceylon, Ryu Kyu Is., SE. China, to Malesia: 
Malay Peninsula, Sumatra, West Java, Bawean, 
Borneo (Sarawak, Sabah), PhiHppincs (Luzon, 
Mindanao), Ambon; much less common than the 
closely related L. ensifolia. Map 3. 

Ecol. In open to somewhat shaded, mostly 
moist places, up to 1100 m, mostly at lower al- 
titude. 

8. Lindsaea ensifolia Swartz in Schrader, J. Bot. 
18002 (1801) 77; Copeland, Fern Fl. Philip. 1 
(1958) 113; Kramer, Blumea 15 (1968) 564.— 
Adiantum ensifolium (Swartz) Poiret, Encycl. 
Suppl. 1 (1810) 139. — Schizoloma ensifolium 
(Swartz) J. Smith, Hook. J. Bot. 3 (1841) 414; 
Backer & Posthumus, Varenfl. Java (1939) 110, 
f. 20; Tardieu-Blot & C. Chr. Fl. G^n. I.-C. 7 
(1939) 129, f. 15 1-2; Holttum, Rev. Fl. Mai. 
2 (1954) 346, f. lOO.—Schizolegnia ensifolia 
(Swartz) Alston, BoI. Soc. Brot. II, 30 (1956) 
24. — Type: coll. unknown, Mauritius (S-PA). 

Pteris stricta Poiret in Lamk, Encycl. 5 (1804) 
713.— L. pteroides Desvaux, Prod. (1827) 312.— 
Type: Commerson s.n., Mauritius (Tie de France') 
(P). 

L. lanceolata Labill. Nov. Holl. PI. Sp. 2 
(1806) 98, pi. 248 f. \.-~Adiantum lanceolatum 
(Labile.) Poiret, Encycl. Suppl. 1 (1810) 134, 
non Fee (1852). — Schizoloma lanceolatum (La- 
bill.) Presl, Tent. Pterid. (1836) Ul.Schizo- 
loma ensifolium (Swartz) J. Smith var. lanceolata 
(Labile.) R. Bonaparte, Notes Pterid. 13 (1921) 
259. — Schizoloma billardieri Gaud. Ann. Sc. 
Nat. 3 (1824) 508. nom. superfl.—L. billardieri 
(Gaud.) Carruthers ex Seemann, Fl. Vit. 
(1873) 337. — Type: Labillardiere s.n.. Cape 
Van Diemen, Australia (P; identity of specimen 
uncertain). 

L. erecta Mirbel in Lamk & Mirbel, Hist. 
Nat. Veg. 5 (1803) 126, non Mettenius (1861).— 
Type: coll.?, Reunion {n.v.). 

Pteris angulata Presl, Rel. Haenk. 1 (1825) 
54, nom. illeg., incl. L. lanceolata Labile. — Type: 
coll. ?, Marianas (n.v.). 

L. membranacea Kunze, Linnaea 18 (1844) 
121. — Type: Gueinzius s.n.. Port Natal, S. Africa 
(B; dupl. in BM, HBG, L, P, W). 

L. sublobata Kunze, Linnaea 18 (1844) 121. — 
Type: Cuming 369, Malacca (B; dupl. in GH, L, 
SING, W). 

L. griff ithiana Hooker, Sp. Fil. 1 (1846) 219, 
pi. 68B. — Schizoloma griff ithianum (Hooker) 
Fee, Gen. Fil. (1852) 108. —Type: Griffith s.n., 
Mergui, Burma (K). 

L. pentaphylla Hooker, Sp. Fil. 1 (1846) 219, 
pi. 67 A. — Schizoloma pentaphyllum (Hooker) 
Fee, Gen. Fil. (1852) 108.— Type: Bynoe s.n., 
Australia ('New Holland') (K). 

L. oligoptera Kunze, Bot. Zeit. (1846) 445.— 
Type: Zollinger 1513 ('& 1515'), Java (B; 
dupl. in HBG, L, Z). 



Schizoloma javae FtE, Gen. Fil. (1852) 109, 
pi. 29 f. 1. — Type: Zollinger 1504, Java (n.v.). 

L. schizoloma Ettingsh. Farnkr. 3 (1865) 
213, pi. 145 f. 4, pi. 146 f. 6.— Type: not cited; 
perhaps a new name for Pteris stricta Poiret 
(vide supra). 

Schizoloma heterophyllum (Dryand.) J. Smith 
var. speluncae Copeland, Philip. J. Sc. 5 (1910) 
Bot. 284. — Type: Foxworthy 578, Sandakan 
(MICH). 

Schizoloma ensifolium (Swartz) J. Smith var. 
attenuatum Domin, Bibl. Bot. 20 (1915) 77, with 
/. typicuin Domin, type: C. B. Clarke s.n., several 
coll. from India;/, pteroides Domin, types: Grif- 
fith 173, Mergui, Burma, and Wallich s.n., 
Singapore; /. praelongum Domin, types: Kunst- 
LER 1881, Singapore, and coll. ?, Mauritius (n.v.). 

Schizoloma ensifolium (Swartz) J. Smith var. 
borneense Domin, I.e. — Type: coll. ?, Borneo 
(n.v.). 

Schizoloma ensifolium (Swartz) J. Smith var. 
clarkeanum Domin, I.e. 76. — Type: Clarke s.n., 
N. India (n.v.). 

Schizoloma ensifolium (Swartz) J. Smith var. 
longipinnum Domin, I.e. 76, with /. typicum 
Domin, type: Gomez s.n., Javoy, India; /. sub- 
simplex Domin, types: Griffith s.n., Malacca, 
and Wallich 92, Ceylon;/, griffithianum Domin, 
I.e. 11, type: Griffith s.n., Mergui, Burma (n.v.). 

Rhizome short- to mostly not very short- 
creeping. Lamina mostly simply pinnate, with 
up to c. 15 pinnae to a side. Pinnae lanceolate 
to linear, with almost symmetric base, non- 
dimidiate, with percurrent costa; veins immersed, 
± evident, very oblique near the costa, less so 
outward, regularly anastomosing even in narrow 
pinnae, with 1 or 2, rarely 3-5 rows of areoles 
between costa and margin. Sori continuous; 
indusium linear, pale brownish, 0.3-0.5 mm wide, 
almost reaching the margin. Spores trilete, smooth. 

KEY TO the subspecies 

1. Upper pinnae gradually and strongly narrow- 
ed, gradually confluent into or at least some 
small ones connected with the terminal seg- 
ment; sometimes bipinnate. . 1. ssp. agatii 
1. Upper pinnae little reduced; lamina with a 

free, conform terminal pinna. 
2. Rachis abaxially sharply bi-angular; sterile 
margin serrate, or rarely subentire 

2. ssp. ensifolia 

2. Rachis abaxially not sharply bi-angular; 

sterile margin entire ... 3. ssp. coriacea 

1. ssp. agatii (Brackenr.) Kramer, Act. Bot. 
Neerl. 15 (1967) 519. Schizoloma agatii Brac- 
kenr. U.S. Expl. Exp. (1854) 216, pi. 30 f. 1.— 
Type: U.S. Expl. Exp. s.n., Fiji (dupl. in K). 

Schizoloma ensifolium (Swartz) J. Smith var. 
intercedens Domin, Bibl. Bot. 20 (1915) 80, pi. 
14 f. 8, pi. 15 f. 3.— Type: Domin s.n., Yarraba, 
N. Queensland (n.v.). 

Rhizome not very shortly creeping, 1 1/2 rnm 
thick; scales as in the next subspecies. Leaves 



212 



Flora Malesiana 



[ser. II, vol. P 



1/2-1 cm apart. Petioles stramineous to reddish 
brown, quadrangular, often sulcate. Lamina 
often lanceolate, with c. 8-15 pinnae to a side, 
sometimes subbipinnate or fully bipinnate. Pinnae 
often rather strongly ascending, the major ones 
c. 5-10 cm by 4-7 mm, 10-15 times as long as 
wide, the lower ones sometimes subauricuiate 
at base, chartaceous or firmly herbaceous, acute 
or subacute, not rarely some lower (but not 
necessarily the lowermost) pinnatifid or pinnate, 
their segments usually rhombic or obovate, rarely 
prolongate-rhombic to lanceolate, up to c. 12 
to a side, decurrent and often wing-connected, 
the basal ones often broader. Apices of pinnatifid 
or pinnate pinnae with a long undivided segment. 
Upper primary pinnae gradually and strongly 
reduced, the uppermost ones less than ^''3 the 
size of the lower ones, terminal segment con- 
fluent with some reduced upper pinnae or lobed 
at the base. Veins in smaller secondary pinnules 
irregularly anastomosing; often only one row of 
areoles present. Sterile margin serrate. Sori 
continuous except as interrupted by incisions of 
the pinnae, in small pinnules of bipinnate leaves 
occupying only their outer margin. Indusium often 
with an irregular edge, occasionally slightly 
exceeding the margin. Spores light brown, c. 
26 n. 

Distr. Malesia: Ambon, Timor and New Gui- 
nea; 2 doubtful collections from Sabah. North- 
and eastward to Micronesia, Queensland, New 
Caledonia, the Tonga Is., and Samoa. 

Ecol. Terrestrial, very euryoecious, but not in 
swamps; to 1200 m. 

Notes. In regions where this and the following 
subspecies occur together intermediates are not 
very rare. 

L. ensifolia ssp. agatii h