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LIBRARY
FHE NEW YORK BOTANICAL GARDEN.
BRONX, NEW YORK 10458
TAXONOMICAL REVISIONS
_ | 23. U Death ; ; Map NA
REPUBLIK INDONESIA
REPUBLIC OF INDONESIA
LEMBAGA ILMU PENGETAHUAN INDONESIA (L.LP.I.)
INDONESIAN INSTITUTE OF SCIENCES
FLORA MALESIANA
BEING
AN ILLUSTRATED SYSTEMATIC ACCOUNT OF THE MALESIAN FLORA /
INCLUDING KEYS FOR DETERMINATION | DIAGNOSTIC DESCRIPTIONS |
REFERENCES TO THE LITERATURE|/SYNONYMY/| AND DISTRIBUTION |
AND NOTES ON THE ECOLOGY OF
ITS WILD AND COMMONLY CULTIVATED PLANTS
PUBLISHED
UNDER THE AUSPICES OF LEMBAGA BIOLOGI NASIONAL
BOTANIC GARDENS OF INDONESIA / BOGOR / JAVA AND
OF THE RIJKSHERBARIUM / LEYDEN / NETHERLANDS
PREPARED
ON AN INTERNATIONAL CO-OPERATIVE BASIS UNDER THE SUPERVISION OF
SEVERAL DIRECTORS OF BOTANIC GARDENS/ KEEPERS OF HERBARIA
AND VARIOUS PROMINENT BOTANISTS
FOR THE PROMOTION OF
BOTANICAL SCIENCE AND THE CULTURAL ADVANCEMENT OF
THE PEOPLES OF SOUTH-EASTERN ASIA TO
THE SOUTHWEST PACIFIC REGION
SERIES II
PTERIDOPHYTA
VOLUME 1
GENERAL EDITORS:
Dr C. G. G. J. VAN STEENIS
DIRECTOR OF THE FOUNDATION ‘FLORA MALESIANA’
& Dr R. E.HOLTTUM
PUBLISHED BY
MARTINUS NIJHOFF / DR W. JUNK PUBLISHERS
THE HAGUE / BOSTON / LONDON
1959-1982
529
My
Foo
COPYRIGHT 1982 vA
All rights reserved, including the right to reproduce
this book or parts thereof in any form
ISBN 90 247 2653 0
PUBLICATION DATES
Part 1 10 Dec. 1959
Part 2 11 Nov. 1963
Part 3 20 April 1971
Part 4 15 Dec. 1978
Part 5 1 March 1982
Printed in the Netherlands
CONTENTS
A ETE SEL, REG ee eee: Gee Ree - ee a a ee me cei ee pee (3)
RECTED LS Rare rth Wak CA al WwS, a Wis Lak rset icchcceke yeh Kgsane eit aniickasuecie, Gees (5)
memenere yy Teil). EIOItin: fo). Steps is Milage net tia peapfanns ie LOS serve na aenee ee (7)
GENERAL CHAPTERS
muroduciony note: Dy iho HOKU. . ass) ee ee a eg eo see, Bee Sas. i
went a, Aaclesian Pieridophytes by R.E.Holttum.. . . 2. ¢ 2 ciel cs Sco Se eS ee ii
ite GO LUADGY Gai Re leas GlGiil —5 5 9 ecw oe nouc ou. Gonlmae cee o ono See ili
eVCLANKCYUNO:l.to)Picropsiad by, Re Be Holttum = 6 4 sess, « Se 2 et ee ix
GencralkeyeNo-2 to-Pteropsida by R-B- Holttummems = 5 - = = - = Ge 6s eee ee ee: Xii
iKeys to the, genera of -Pteropsida by R. E. Holttum = =... ......0+%2+4s5:24.-48 XV
PIDMOLTADLYIOY Rove SP ELOLUEUI: cer crac OS. ow 2s a ee eae eo ha) ore ee aeons Seals XXil
TAXONOMICAL REVISIONS
in alphabetical sequence
Gyatheaccae bys EE. -HOlttum Fa fines es es ee eee ee 65
GicicheniaccacibyoR. BHOlttuni = so sos, 2a ey oy A ee eo 1
WSOCTACCOCID Ys Ac EA Gry AIStOM ye wet to oe renee ance ek cede eee) oe, 62
Pamsacaoroup by KU, Kramer o's 22. es, & Soe Se ee ee ee ee 177
Lomariopsis group by R. E. Holttum & E.Hennipman .............2.2.4... 255
Selnzacaccde by RB. Holttum one. sb hs eee ee ee ee. 37
mueiypicridaceae by. RE: Holttum’., ... 2-2.) 252.6 a PS eee eee 331
ADDENDA
Addenda, corrigenda et emendanda by R. E. Holttum & K.U. Kramer ......... 561, 565
INDEX
Index to scientific plant names by Mrs. M. J. van Steenis-Kruseman ...........-. 567
Dedicated to the memory of
C.F. A. CHRISTENSEN
DEDICATION
CARL FREDRICK ALBERT CHRISTENSEN (1872-1942) was the founder of modern fern taxonomy.
To appreciate the scope of his work, it is necessary to understand the confusions of thought on
the subject which persisted through the 19th century and were still evident in the summary
prepared (by Diets) for ENGLER & PRANTL’s Pflanzenfamilien in 1899. CHRISTENSEN’S first great
work was his Index Filicum (1905-6) in which he listed all known fern binomials and also rele-
gated many to synonymy. In the main he adopted the classification and nomenclature of DiEts.
While preparing the Index he came to realize that many generic concepts accepted in the Index
were unnatural or confused. This was especially evident in the great complex of species which he
listed under the name Dryopteris. He next made a study of the tropical American species of that
complex, and in so doing discovered how to separate them into natural groups (1913, 1920).
At the time I first made contact with him (about 1925) he had begun to study ferns of the Old
World tropics. I maintained a regular correspondence with him from 1925 to 1940, and sent him
many specimens for identification. I also met him in Europe in 1930, 1934 and 1938 and had long
discussions with him. I benefited from his wisdom also indirectly through the publications of
R. C. CHING, who studied with CHRISTENSEN in 1929-1932 and applied CHRISTENSEN’S ideas to
Chinese and Indian ferns in an important series of papers in the 1930s. CHRISTENSEN’s identifica-
tions of my collections and his comments upon them were the basis on which my own work was
built; in the present Series of Flora Malesiana I have tried to extend his methods and his ideas
to a much wider range of species than he could have encountered. To him I am profoundly
grateful, and I am concerned also to acknowledge my debt, through him, to some perceptive
earlier workers, notably G. H. METTENIUS and JOHN SMITH.
The objectives of any scheme of biological classification are to show natural relationships and
to provide a means for the identification of individual organisms. It has sometimes been suggested
that only the latter objective is important, and that a ‘practical’ scheme is all that is needed. The
history of fern classification has shown that artificial schemes, made without thought as to
relationships, do not work; and distribution-maps based on such schemes are meaningless. Fern
classification as understood today should be based not only on gross-morphological characters
but also on microscopical characters pertaining to the fern’s anatomy, indument, spores, gameto-
phytes, efc., and on cytotaxonomy.
There can be no doubt that existing ferns have originated through a process of evolution. They
have therefore an inbuilt classification, and our object is to find it; the nearer we get to it, the
nearer we are to the practical aspect of taxonomy. Fossils provide no clear evidence of the
progress of the evolution of the great majority of leptosporangiate ferns. In most cases our only
evidence for this is the natural relationships shown by taxonomy. We now have reached the stage
at which most Malesian species can be allocated to definable natural groups which may have
generic rank; most genera can also be associated in groups which appear to be natural; but it is
often not yet clear how groups of genera are inter-related. For example, within the family
Thelypteridaceae 1 cannot see a definite pattern of inter-relationships between the groups of
genera which I have recognized. For a better understanding of this wider problem, genera
throughout the tropics need to be taken into consideration; such an undertaking is beyond the
scope of Flora Malesiana Series II, but I believe that this Series has presented a great deal of new
evidence on which wider considerations may be based.
Some botanists appear to think that a Flora is not the place for discussion of such questions.
I disagree with that idea. Floras and taxonomic monographs always appear to account for every-
thing, owing to the nature of their presentation. But in fact there are always gaps and uncertain-
ties, especially in tropical Floras; some indication of this should be given; no classification can be
final.
(7)
FLORA MALESIANA
Even within groups of ferns already dealt with in Flora Malesiana, much more information is
needed. For example, existing specimens do not show clearly the distinctions between species in
the genus Plesioneuron (Thelypteridaceae). There is a great need for new collections made by
persons who have specialist knowledge and are prepared to undertake prolonged critical search.
After the publication of my book on the ferns of Malaya, Betry MOLESworRTH ALLEN, by per-
sistent collecting, discovered nearly twenty additional species including representatives of three
additional genera.
The genera of LINNAEUS, which should be the basis of fern classification, are very crudely
defined, and are only useful through agreement as to their typification. He did not notice indusia,
upon which J. E. SMITH (1793) was the first to base new generic concepts, but SMITH also was not
a critical observer. Within a few years, several other authors extended his observations and
proposed new generic names, some not well distinguished, and in 1801 Swartz and BERNHARDI
noted (more exactly than SmiTH) the differences between annulate (or gyrate) and exannulate
sporangia. These observations were collated by Swartz in his Synopsis Filicum (1806) where he
separated the genera of Osmundaceae, Schizaeaceae and Gleicheniaceae as spuriously gyrate,
distinguishing the annulate genera (Polypodiaceae) solely by the form of indusia and the form and
position of sori where indusia were lacking. An extreme example of the artificial nature of some
genera proposed at this period is Belvisia MiRBEL (1802) which, in addition to the recognized type
B. spicata (L.) MirBeEL (Polypodiaceae) included species now allocated to Actiniopteris, Schizaea,
Asplenium and Ceratopteris.
Simultaneous with SwARTZ, SCHKUHR was producing the first good series of illustrations of
ferns (1804-1809). When one makes a drawing of a plant with the intention of accuracy, one
often notices hitherto neglected characters. This was true of SCHKUHR, who observed and com-
mented upon many details, especially of hairs, which have, since CHRISTENSEN, become recog-
nized as important key characters. He noticed the jointed (septate) hairs of Ctenitis villosa (L.)
CopPEL., and portrayed accurately the equally long unicellular hairs on the indusia of a species of
Christella, though he did not comment on the latter. In some cases he illustrated venation clearly
and accurately, in others indistinctly or casually. Some of SCHKUHR’s drawings were made from
living plants, but most were from dried specimens.
HOoKeErR and GREVILLE’s Icones Filicum (1827-1831) was the next illustrated work. The plates
were better executed (by GREVILLE) than SCHKUHR’s and one can also detect a gradual increase of
interest in detail as the series progressed. For example, plate 5 depicts Ceterach pedunculosum and
plate 6 Grammitis decurrens, but in neither case are any details of venation shown; both species
are now placed in the genus Colysis Prest (Polypodiaceae). Plate 125 shows Polypodium irioides,
with enlarged details of venation well observed. Simultaneous with Hooker and GREVILLE,
BLUME (1829-1830) was publishing the wonderful plates 1-65 of his Flora Javae, Filices, in which
details are, on the whole, even more carefully dealt with. His subdivision of Polypodium is
important.
H. ScHoTrT, at the imperial palace of Sch6nbrunn, had living fern plants in his care and pub-
lished (1834) a short series of excellent engravings illustrating new genera, showing much more
detail than Hooker and GREVILLE; some of these were certainly based on living plants, in parti-
cular his Nephrodium which shows very exactly capitate hairs and the elongate unicellular glands
which are present on the stalks of sporangia, noted by no-one else until I made drawings of them
in Singapore in 1943 (published 1971). Between 1840 and 1851 G. Kunze published a series of
illustrations which he regarded as a continuation of SCHKUHR’s. He was in charge of the Botanic
Garden at Leipzig, in which was the best collection of living ferns in Europe (soon to be sur-
passed by Kew). His successor at Leipzig, G. H. METTENIUS, inherited KUNze’s living collections
(8)
Dedication
and herbarium, and on their basis prepared a new system of classification of ferns (1856) with
plates often showing new details. He subsequently prepared monographs describing all known
species of several major genera, after which he began observations on the collections from
Malesia in the Rijksherbarium. He had previously misinterpreted some of BLUME’s species
through not having seen authentic specimens, and corrected some of them in the Ann. Mus. Bot.
Lugd.-Bat.; he also incorporated new basic observations on several genera. While engaged on the
latter work, he died of cholera in 1866 at the age of 42. Had he lived longer, he would have
changed the course of pteridological thinking; I will revert to him later.
C. B. PRESL was given the task of describing the collections made by HAENKE on the Malaspina
Expedition; these included many specimens of ferns from the Philippines (described in 1825).
As a result PRESL became interested in the classification of ferns, and realized that characters
other than those of sori needed to be taken into account. In 1836 he published Tentamen Pterido-
graphiae, comprising a new scheme of classification in which the arrangements of vascular tissue,
and of venation, had an important place. His work is illustrated by many small drawings showing
details of venation in relation to sori, in most cases quite accurately. His later publication (1848)
showing arrangement of vascular strands in the stipes of ferns is not so well observed. PRESL’s
emphasis on venation led him to associate together species of very diverse relationship, but it was
a beginning of new thought.
At the same time JOHN SMITH of Kew had been taking an interest in the cultivated tropical ferns
in his charge, many raised (as at Leipzig) from spores from herbarium specimens. He was in
touch with ROBERT BRowN, who had made some original observations on the venation of ferns
when describing his own Australian specimens and also some collected by HorSFIELD in Java.
SMITH was also in touch with FRANCIS BAUER, the Kew botanical artist, and supplied him with
living fern plants and herbarium specimens, from which BAUER prepared a beautiful set of forty
plates. These were submitted to W. J. HOOKER (then at Glasgow) who arranged for them to be
published and added more, prepared by W. H. Fitcu, 120 plates in all, finalized after HOOKER
came himself to Kew. Many of the genera are those of PRESL, but twenty were newly named and
described by JOHN SMITH. SMITH himself had prepared a new scheme of classification indepen-
dently of PresL, finding much agreement between their ideas when the Tentamen appeared; he
collated his nomenclature with PREsL’s and his scheme was published by Hooker in 1841-1843.
JOHN SMITH continued to study ferns, and to add to the collection of living plants at Kew. By
1865, when he was obliged through failing sight to retire, he had seen more than 1000 species of
ferns in a living condition, of which he published a list in 1866. The final summary of his ideas,
resulting from continued observation of living plants, appeared in 1875 and will be considered
later.
Having published Genera Filicum, HooKER planned Species Filicum, in which he proposed to
describe all known species of ferns. For this, he had to re-consider the question of classification,
and concluded that Pres had proposed too many genera; the result was that HOOKER’s genera in
Species Filicum are almost the same as those of SWARTZ. The work was published in five volumes
over a period of twenty years (1844-1864); Hooker planned to follow it with a summary in one
volume, to be called Synopsis Filicum. He died just as the first part of the latter was printed.
J. D. Hooker, who succeeded his father at Kew, engaged J. G. BAKER to continue the Synopsis
on the lines planned, and this was completed in 1868 (second edition, with many additions by
BAKER, in 1874). In 1891 BAKER published a summary of new ferns discovered since 1874, still
with the same set of genera.
HOOKER’s Species Filicum was illustrated by 304 excellent plates prepared by W. H. FitTcH
(often two species on one plate). These show clearly and accurately details of venation and
(9)
FLORA MALESIANA
indusia, but rarely any smaller structures. In his descriptions HOOKER rarely described details of
hairs or scales. He thought such details unimportant; his main objective (see Vol. 3, p. 3) was
‘to assist the tyro in the verification of genera and species .. . natural habit is often a safer guide
than minute microscopic characters’. He placed most exindusiate species of Thelypteroid,
Tectarioid and Dryopteroid ferns in the genus Polypodium, but some in Gymnogramme and
Meniscium; Dictyocline was merged with Hemionitis. He could not understand how JOHN SMITH
could believe Brainea to be closely related to Blechnum, though it differs from Blechnum only in
the absence of indusia. He placed Brainea between Gymnogramme (which included the diverse
genera Selliguea and Syngramma) and Meniscium. His refusal to examine details led him to
include in one species specimens which show great diversity in what are now considered to be
significant characters. He united most of FEE’s species of Lomariopsis (including also Terato-
phyllum MeEtv.), thus including several distinct Malesian species in Acrostichum sorbifolium L.,
of which the type came from the West Indies. His confusions in the synonymy of Thelypteroid
ferns are very numerous, and can only be understood by reference to his herbarium. BAKER’s
descriptions of ferns discovered after HOOKER’s death are even less satisfactory than HOOKER’s
and often do not serve to identify specimens with any certainty.
The remaining authors who proposed new schemes of classification were FEE (1852) and
T. Moore (1857). FEe’s works were all admirably illustrated and his numerous plates show many
significant details, but not always accurately. For example, in tab. XXI A, fig. 2 he was the first
to show a transverse section of the stipe of Pleocnemia (sensu HOLTTUM 1974), but the accom-
panying figure of venation in an allied species (fig. 1) is not accurate and fails to show the distinc-
tive sinus-teeth. Neither FEE nor any earlier author (so far as I have observed) shows the distinc-
tive row of four cells on one side of the sporangia of leptosporangiate ferns. FEE attempted to use
the number of cells in the annulus as a generic character, but this is rarely practicable. His scheme
is more elaborate than PRESL’s but is no nearer to a natural arrangement by present standards.
He has Phegopteris as a genus separate from Polypodium, but in the same group of genera, not
with its true allies, which are in other groups. Under the tribe Acrosticheae he has an astonishing
diversity of genera.
THOMAS Moore’s scheme is accompanied by good small drawings to show diagnostic
characters. For example, he shows the difference in venation between Stenochlaena and Lomariop-
sis, not noticed by FEE. But his scheme only differs in minor features from that of PRESL.
MILDE in 1866 made important observations of scales and stipe-anatomy showing a clear
distinction between Asplenium and Athyrium (including Diplazium); he elaborated these in 1870.
METTENIUs had noted that previous attempts to distinguish these genera were unsatisfactory, and
Hooker denied that any clear distinction was possible (and in 1928 Bower still copied HOOKER’s
statement). In my judgement (HoLTTUM 1947) Asplenium and Athyrium are not very closely
related.
R. H. Beppome did not propose a new scheme of classification, but during the years 1856-1882
he made a more intensive field study of ferns in a tropical region than any previous author. He
was critical of HooKEr’s genera and made some minor alterations in them for purposes of his
Handbook (1883, with Supplement 1892), though still accepting the main scheme (his preface
hinted that more needed changing). His work covered the Malay Peninsula and so is important
for Flora Malesiana (he also accepted HooKER’s misidentification of some Indian ferns with
species in Java).
JOHN SmiTH’s Historia Filicum (1875) proposed a new scheme based on much study of living
plants subsequent to his first one (1841). He did not use a microscope, and rarely refers to details
of structure of sporangia, scales efc., but from observation of his plants he did learn much that
(10)
Dedication
HOooKER never understood. I will refer later to some of his insights in a discussion of the work of
Diets. SMITH and MetTeENtus, both observers of living plants, were the only authors of their
period who (apparently independently) separated Phegopteris, Dictyopteris and other terrestrial
exindusiate ferns from the alliance of Polypodium and transferred them to one including
Aspidium. Both authors maintained separate genera for the exindusiate ferns, but JoHN SMITH
admitted that probably some species were placed in Phegopteris and Dictyopteris merely because
the only known specimens had old sori from which indusia might have fallen. Presumably he
still thought the idea of uniting indusiate and exindusiate species in one genus too revolutionary.
It should be noted that both METTENIUS and SmiTH had a mixture of Thelypteroid, Tectarioid
and Dryopteroid ferns in their genera, whether indusiate or not; and SmitH kept Meniscium
(Thelypteridaceae) far from his Aspidioid ferns. The major advance in thinking was that ind-
usiate and exindusiate species could be closely allied; this was something HOoKER refused to
consider.
In this connection, the history of Pleocnemia leuzeana (GAUD.) PRESL is instructive.
GAUDICHAUD described the species (from the Moluccas) in the genus Polypodium because its sori
were exindusiate. PREsL founded a new genus based on the combination of a particular vein-
pattern with circular exindusiate sori. Later CUMING collected specimens in Luzon which were
similar in venation and general aspect, but some of them had indusiate sori. HooKER, who had
illustrated the genus Pleocnemia as exindusiate (Gen. Fil. t. 70A, copied from Pres) published
drawings made from two of CuMING’s specimens, one sterile and one showing indusiate sori
(t. 97), and stated that this gave him an opportunity to correct his previous ‘error’ in reporting
that P. leuzeana was exindusiate. But CUMING made four different collections (all seen by HOOKER)
which are now known to represent three distinct species, two of them indusiate, one exindusiate,
all different from the type specimen of P. Jeuzeana. HOOKER assumed that some specimens had
lost the indusia which they originally possessed. F&E had specimens of the same collections from
CumING, and speculated (1852, p. 311) on the strange fact that different plants of the same species
could have, or lack, indusia. BEDDOME, examining plants of Pleocnemia from N.E. India which
are in fact exindusiate (as seen from young sori) and belong to a species different from all three
in the Philippines, thought that his Indian specimens must have lost their indusia and figured a
fertile leaflet from a Philippine specimen (Ferns Br. India, t. 134). COPELAND in 1960 (p. 310) still
only recognized one species in the Philippines, noting that the indusia are ‘sometimes fugacious’.
Recent collections from Mt Makiling, at the foot of which CoPpELAND spent several years, confirm
that CumING’s three species are distinct. The fronds are very large, so that only small parts can be
put on herbarium sheets, and the stipe-scales (usually absent from herbaria) are distinctive. But
herbarium specimens do show enough peculiar details if one knows what to look for, and the sum
of these characters is sufficient to indicate that these species (and some others) form a genus
distinct from Tectaria (to which CopELAND referred P. Jeuzeana), though the venation-pattern of
Pleocnemia does occur in some species of the former. The sinus-teeth, which project out of the
plane of the frond and are very conspicuous in living plants, were not noted by anyone except
GAUDICHAUD until I re-defined the genus (HoLTTUM 1951, 1974); there are also distinctive glands
(noted by METTENIUS but not by others). The petiolar vascular structure, also peculiar, was
figured by FEE (1852, t. 21A fig. 2) and mentioned by no-one else.
It was details such as the presence and nature of scales, hairs and glands on pinnae that
METTENIUs noted; these have subsequently been found to be significant diagnostically, and they
give METTENIUS’s specific descriptions a significance that is often lacking in BAKER’s. METTENIUS
maintained large genera, perhaps (like CHRISTENSEN at a later time) because he did not want to
publish new binomials until he was more sure of them; he subdivided his large genera much more
(11)
FLORA MALESIANA
intelligently than Hooker, and made improvements in subdivision in his works published in 1864.
DIELs erred in ignoring several important observations made by METTENIUS.
The situation near the end of the century was that in most cases clear distinctions between
groups of genera, and often even between genera now known not to be closely related, had not
been discovered. This was due to a failure to understand that similar structures, whether of
venation or sori, could have come into existence along different evolutionary lines. It is very clear
that this is true of a simple type of anastomosis, seen in such genera as Acrostichum (s.str.),
Pteris, Elaphoglossum, Lomagramma, Taenitis, Lindsaea and Hemionitis; in Pteris, Lindsaea and
Elaphoglossum most species have free veins. The vein-pattern in Tectaria (Aspidium of Ind.
Fil. 1905) and Microsorium (a segregate from Polypodium) is closely similar; in Malaya I found
that up to 1955 a species of Tectaria had been included by all authors in Polypodium. It is also
evident that acrostichoid ferns belong to several different alliances; and the acrostichoid condition
is not exactly definable, so that authors disagreed in assigning genera to it. The sori of Davallia
and Microlepia are very similar, but in other respects the plants are very different. An extreme
case is Heterogonium PRESL, which I believe to be a natural genus (HOLTTUM 1949); some species
have indusia, some not; some species have free veins, some have anastomoses; some have separate
indusiate sori, some are acrostichoid.
So the problem is to look for characters which may be a better guide to relationship than vein-
patterns and sori. MILDE had shown the way by distinguishing between Diplazium and Asplenium
on the basis of scales combined with vascular anatomy of the stipe. SmirH had noted that
Polypodiaceae (s.str.) and the Davallia group of genera have a creeping caudex with stipes jointed
to the dorsal surface of it; he coined the term Eremobrya for ferns of this habit; other ferns he
called Desmobrya. The two terms were first defined in 1855. By this standard the ferns included
by CHING (1940) and HoLtrum (1947) in Grammitidaceae are separate from Polypodiaceae.
METTENIUS also found that the spores of the two groups differ (see below on DiE.s 1899).
HERMANN CunrisT (1833-1933) was a lawyer who throughout a long life was actively interested
in plants. He began to publish papers on ferns in 1890, and in 1897 produced Die Farnkrduter der
Erde, an attempt to give a more balanced view of the more important species throughout the
world than HooKeEr and BAKER. He recognized the nature of the problem stated in the preceding
paragraph, but did not manage to do much towards solving it. He placed Aspidium and Phegop-
teris (still separate genera, on the model of METTENIUS) in a family Aspidiaceae, distinct from
Polypodiaceae, but under both Aspidium and Phegopteris had a great mixture of species not
closely allied. In Polypodiaceae, tribe Acrosticheae, he had much confusion, especially in the
genus Stenochlaena (see HOLTTUM 1978, pp. 261, 266); some of this was copied by DieELs. His
later work also showed lack of critical insight. In his monograph of Elaphoglossum (1899) he
tried to subdivide the genus on characters of venation, but did not examine the veins carefully
and the result is confusion; in his paper of 1907 on the Philippine species of Dryopteris (the com-
posite genus of Ind. Fil. 1905) he did not make good descriptions nor understand relationships
between species. He did not know of MILDe’s work.
The century closed with the volume of ENGLER & PRANTL’s Pflanzenfamilien covering vascular
cryptogams, in which DiELs dealt with almost all the ferns (1899-1900). His Polypodiaceae con-
sisted of nine tribes. He transferred several genera of the tribe Acrosticheae of some previous
authors to Aspidieae, but mixed together Aspidioid and Polypodioid species under Polybotrya. He
placed the Polypodioid genus Platycerium in Acrosticheae. He united Phegopteris with Aspidium
but had a great mixture of species in it; his treatment only adds more confusion to an already very
confused situation. He did understand MILDe’s work, but he failed to notice some important
observations made by METTENIUS and JoHN SmiTH, of which the following are three examples.
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Dedication
1. Gleicheniaceae. PRESL based his primary division of Gleichenia (sens.lat.) on the position of
the sori on the veins, stating that in Eu-Gleichenia the sori were terminal, in all other cases dorsal
on the veins. This division was copied by HOOKER, CHRIST and DIELS; but in 1856 METTENIUS
had stated that the sori are not terminal in Eu-Gleichenia, and had repeated this in 1863. In the
latter paper he divided G/eichenia into three subgenera, stating that two of them agreed in scales
and in sporangia, the third (Dicranopteris) differing in both these structures. This was ignored by
DiELs, who did not cite the paper of 1863 and mixed together in one subgenus species of Dicranop-
teris with those which differed both in scales and in sporangia. When preparing an account of the
family for Flora Malesiana (HOLTTUM 1959) I failed to notice METTENIUS 1863 and repeated his
observations, differing only in the recognition of Dicranopteris as a genus distinct from Gleichenia,
the latter having three subgenera; this is certainly the important division.
2. Stenochlaena and Lomariopsis. In 1875 (p. 140) JOHN SMITH stated the distinctions between
these two genera (he had united them in 1841 and subsequently discovered the difference through
observation of living plants). METTENIUS still included them in the same genus (1869, in a post-
humous paper edited by KuHN) but in separate sections, and he established a new genus Terato-
phyllum, distinct from both, with two species. DiELs united Stenochlaena, Lomariopsis and one
species of Teratophyllum in one genus (in the tribe Asplenieae) which he divided into two sections:
Eu-Stenochlaena comprising the whole of Lomariopsis and Teratophyllum aculeatum (BL.) METT.,
and Cafraria, which consisted only of S. tenuifolia; the latter differs from the type species
S. palustris (BURM.) BEDD. in having bipinnate fertile fronds and in little else. This is an absurdly
unnatural division. DieELs included the second species of Teratophyllum (T. articulatum (J.SM.)
MetT.) in Polybotrya (tribe Aspidieae).
3. Grammitidaceae. This family was recognized as distinct by CHING in 1940; for fuller details
see also HoLtrumM 1947 and 1955. Diets placed all species of the family in Polypodium sect. Eu-
Polypodium, mixing them indiscriminately with true Polypodium species, except Prosaptia PRESL
which he included in Davallia. BLUME in 1830, though retaining them in Polypodium, had already
distinguished these ferns as ‘spurious’ in that genus. METTENIUS (1866) distinguished them in
Polypodium as Div. 1, Sphaerosporeae, placing true Polypodium in Div. 2, Nephrosporeae; he
did not mention Prosaptia in this paper, but had previously placed it with Davallia. As above
noted, JoHN SMITH placed most Grammitoid ferns in his division Desmobrya, and thus separated
them from Polypodium, but somehow he included Prosaptia (with the closely related Cryptosorus)
in Eremobrya; he did however note that their sori were very different from those of Davallia. For
some reason which is not at present understood, Grammitoid ferns are difficult to maintain in
cultivation, and not one of them appears in JOHN SMITH’s list of species which he had seen alive;
this probably accounts for his mistake in placing Prosaptia with Polypodium. METTENIUS always
noted the peculiar hairs on plants of Grammitidaceae, and also the hairs on scales and setae on
sporangia, where these occur (true Polypodiaceae never have these characters). When he died in
1866 he was just beginning to see the significance of such structures.
Towards the middle of the 19th century academic botanists realized that taxonomic study, of
the limited and formal kind which still prevailed, did not deal with important aspects of the life
of plants. So they started ‘scientific botany’, but they made the mistake of thinking that taxonomy
was an out-dated activity; many such botanists still persist in that mistake. What was needed was
a better taxonomy, not its abandonment. This was especially true of tropical plants in general, and
most ferns are tropical; significant facts about these plants had often not been put on record,
or if recorded (such as the hairs figured by ScHKUHR) had not been understood. As ‘scientific’
botany diverged more and more from taxonomy, the shortcomings of the latter were less and less
understood. A factor in this process was, and still is, the binomial system of nomenclature.
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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 (1863A) and on Hymenophyllaceae (1864B).
Morphologists who have not a wide knowledge of taxonomy are apt to think that any species
is representative of the generic name it bears, and thus are liable to have erroneous ideas about
genera (especially where such genera are still not clearly defined), and may be misled into making
wrong comparisons or invalid statements about phylogeny. In view of the above discussion on the
history of taxonomic study of the leptosporangiate ferns, it is evident that most 19th century
taxonomy was an inadequate guide to morphologists. The most important morphologists were
GorBEL and Bower. Bower began his studies in the 1880s, mainly on the more primitive ferns.
When he came to his summary on the leptosporangiate ferns (1928) he quoted GOEBEL’s com-
parison of their study to wandering in a dark and trackless forest, but he did not know enough
about existing information which could have provided him with some guiding light. He did not
know of the work of MiLpE and discussed the possible evolution of the sorus of Asplenium by
reference to a species of Diplazium. He discussed Stenochlaena, which he interpreted according to
the confused statement by CurisT, and described the anatomy of the rhizome, but the material
he described belonged to a species of Lomariopsis, as he could have learned from JOHN SMITH.
He placed Phyllitis ina group separate from Asplenium, not knowing that natural hybrids between
the two existed. He accepted Curist’s comparison of Elaphoglossum with Syngramma, though
the resemblance between the two is very superficial. He accepted the idea that the sorus of
Microlepia was marginal in phyletic origin, but did not realize that this might also be true of
Cyathea and Dryopteris. He insisted that Deparia was a natural genus, though each of the species
included in it shows an alliance to a different group of ferns. He had not looked at CHRISTENSEN’S
dismemberment of ‘Dryopteris’ and accepted a phyletic sequence (fig. 663 on p. 132) which
derives the vein-pattern of Bolbitis (then still included in Leptochilus) from the condition of a
Thelypteroid fern. But he did have a better understanding of the Gymnogrammoid ferns.
GoeBEL had far more understanding than Bower, having spent at least two periods of study in
Java (BOWER never went to the tropics), but he did not keep in touch with CHRISTENSEN’s work.
As above noted, CHRISTENSEN made a systematic study of all the tropical American species
included in the comprehensive Dryopteris of Index Filicum. In so doing he followed the example
of METTENIUs in looking for details of dermal appendages, but more critically and more con-
sistently than MeTTENIUus had done; he had also a much wider range of species to examine. In this
process he discovered that the many species could be separated into groups according to the
nature of their hairs; he rightly insisted that groups distinguished in this way show also many
other differences of a less easily definable character. CHING (1936, p. 243) added the distinctive
character of vascular anatomy of the stipe of Thelypteroid ferns, in which they constantly differ
from Crenitis and Dryopteris (s.str.), as indeed METTENIUS had noted in his discussion of Aspidium
in 1864. CHRISTENSEN (1911) expressed the opinion that some of the groups he had distinguished
should be regarded as good genera, but he retained them in Dryopteris because he wished to
(14)
Dedication
examine species of the Old World before publishing new combinations. This work of
CHRISTENSEN’S was a turning-point in fern taxonomy. R. C. CHING applied CHRISTENSEN’s ideas
to ferns of southeast Asia, clearing up much previous confusion. But Thelypteridaceae are far
more abundant and more diversified in Malesia than in mainland Asia. When writing my book
on the ferns of Malaya (1955) I adopted CHING’s generic concepts but stated (p. 236) that the
resulting arrangement was not a natural one. I made new observations, especially of glands and
hairs on sporangia (some not then published) but could not see my way to a good re-arrangement
on the basis of the limited number of species in Malaya. It was only when I looked at all species
in Malesia, mainland Asia and the Pacific (and also many previously unnamed collections),
noting in detail structures not mentioned in earlier descriptions, that I was able to see how to
improve on CHING’s scheme, and to establish new genera peculiar to the Old World. CHING’s
work and mine (presented in the present volume) are built on CHRISTENSEN’s methods and on his
insights.
CHRISTENSEN subsequently identified a number of collections of ferns from Malesia and Asia
(including my Kinabalu ferns, 1934) and wrote a fern flora of Madagascar (1932); in so doing he
examined a large number of type specimens which had not been well described and published
new information about them. The nomenclature in my book of 1955 was largely dependent on
his observations on types. In 1939 he contributed a chapter on the classification of ferns to
VERDOORN’S Manual of Pteridology. This contains many new ideas, the result of his wide-ranging
studies; from it one can judge the progress made since 1905, largely due to his own work and
thought. His last work was a fern flora of Samoa, published after his death (1943). Owing to the
stress of the war situation, no adequate obituary notice was published.
Pteridology in Malesia. The first considerable field work was on the ferns of Java, summarized
by BLumeE in 1828 and elaborated with excellent illustrations in 1829-30 (additional plates were
published in 1847 and 1851). Little more was published until RACIBORSKI went to Bogor and
undertook new field studies in West Java, summarized in his book of 1898; his descriptions are
better than most of their time and his ecological information is valuable. A few years later, VAN
ALDERWERELT VAN ROSENBURGH began fern studies covering the whole of Malesia by collating
all existing descriptions, most of them too brief or too inaccurate to form a good basis for the keys
which he prepared. In the main he followed the nomenclature of Index Filicum, but he wanted
more clear-cut distinctions between genera as a better guide to identification, and so he adopted
an artificial system. He reverted to a comprehensive tribe Acrosticheae, and a tribe Phegopterideae
widely separated from Aspidieae; he revived Pleocnemia Pres and included in it some Thelyp-
teroid ferns. After completing his Handbook (1908) he continued a critical study of the specimens
in the herbarium at Bogor, including many new collections, also plants in cultivation. He pub-
lished descriptions of these in a series of papers, those up to 1917 being summarized in his
Supplement. These new and amplified descriptions show a careful examination of much detail not
previously recorded, including observations on spores ; many of his new species are still recognized
in the present work.
In Malaya H. N. RIpLey made large new collections in the years 1888-1911, but his published
work on them (1926) is very uncritical; his generic and specific descriptions are confused and
sometimes inaccurate; the names he wrote on herbarium specimens at Kew are often wrong. Thus
his statements on distribution of species are also often wrong. It is sometimes impossible to know
the basis of such statements because often he did not write names on herbarium sheets in
Singapore; I re-arranged the sheets without noting in which covers RIDLEY had placed them.
RIDLEY’s work on ferns is therefore usually ignored in Flora Malesiana except for his new names.
E. B. CopELAND (1873-1964) began a study of ferns in 1893 but soon specialized in plant
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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 1960. He was the first person
to understand that HooKker’s genera Cyathea, Hemitelia and Alsophila were unnatural, but his
revised scheme for Cyatheaceae in 1947 was little better because he did not examine the detailed
structure of scales. Similarly, in dealing with Thelypteridaceae (which he did not recognize as a
distinct family) he did not look carefully at hairs and glands; his descriptions of species are little
better than BAKER’s. His floristic work suffered also because he did not see the types of many of
the older species and misconstrued some of them; this however does not excuse his failure to
distinguish between Sphaerostephanos penniger (HooK.) HoLTtTuM and Pneumatopteris truncata
(Poir.) Ho_trum (as named in the present work). Yet in Hymenophyllaceae he did make very
careful detailed observations of structure, which were very fully illustrated (COPELAND 1933,
1937, 1938). His families Pteridaceae and Aspidiaceae of 1947 are both confused mixtures which
are still not fully disentangled. In general, CopELAND’s failure was due to not looking for signifi-
cant characters. His statement that the generic separation of Gymnocarpium dryopteris NEWM.
from Phegopteris connectilis (MiCHx) WATT was hardly possible is an illustration of this.
Simultaneously with CopELAND’s work in the Philippines, C. A. BACKER (from 1905) was
making large collections of ferns, as part of his general herbarium of the flora of Java. He
collaborated with O. PosrHumus, who had specialized on ferns and had made many collections
in several other islands also, in the production of a fern flora of Java (1939). The nomenclature
follows that of the third supplement of Index Filicum. In general, this is a considerable advance on
VAN ALDERWERELT, though the descriptions of species in complex genera are not as good as
VAN ALDERWERELT’S later ones. In Cyatheaceae the genera are not well distinguished and the
specific descriptions are very inadequate; the authors could have learned much from METTENIUS
(1863B). In Dryopteris no attempt is made to separate Thelypteroid species from the rest. The
Grammitoid ferns are not separated from Polypodioids. The citation of synonyms is often un-
critical. The work is of value mainly for its ecological information, but where species have been
confused (e.g. under Dryopteris uliginosa) ecological information is also confused.
The present situation. At the beginning of this volume (p. ii, 1959) I presented a tentative list of
genera, remarking that much new information would arise in the course of study in the produc-
tion of the present Flora, and that new ideas on classification would probably emerge. I refrained
from assigning the majority of genera of leptosporangiate ferns to families. In part 2 (1963) I
showed Dicksonia and Cyathea to be much more nearly allied than I had thought in 1959. The
inter-relations between genera there presented still appear sound, but the assignment of genera to
families is still uncertain. In part 3 (1971) K. U. KRAMER presented a major revision of the genera
of the Lindsaea group, clarifying distinctions between them and making a new subdivision of
Lindsaea. Part 4, on the Lomariopsis group (as delimited by me in 1947) included much new
information, especially on Elaphoglossum and Bolbitis, with a revised account of my earlier work
on the other genera. The present part attempts a new subdivision and conspectus of the Thelyp-
teris group of genera, which is so sharply distinct from other groups that I judge it to deserve
family status. This decision commits us in some measure to the ultimate recognition of other
families, but for this we still need more evidence. I think that the elaborate arrangement of
Picu1 SERMOLLI (1977) is premature, though it is much nearer to being natural than COPELAND’S
of 1947.
(16)
Dedication
We need more information about significant characters, and to be useful they must be avail-
able for all species; but no-one can tell in advance which characters will be significant. Since
MANTON’s book of 1950 the observation of chromosome numbers in a great range of ferns has
provided important new evidence; but chromosome number by itself, without evidence of con-
formity with some quite different characters, can be misleading, and the proportion with species
with known chromosome numbers is still relatively small in some genera and not easily aug-
mented. As a result of the work by MANTON and others it has become clear that phenomena like
hybridization and polyploidization — formerly regarded to be extremely rare in ferns — com-
monly occur also in tropical ferns. The impact of cytotaxonomic work on fern classification has
recently been thoughtfully dealt with by Lovis (1977) and WALKER (1979). KLEKOwSKI (1979) has
contributed much to our knowledge of the reproductive biology of the ferns also in relation to
polyploidy.
Morphologists can provide useful suggestions for characters of possible significance. Mor-
phology and taxonomy are interdependent; morphology without a good taxonomy may arrive
at wrong conclusions; taxonomy without the stimulus of morphology may miss important dis-
tinguishing characters. Features which have recently been shown to be of great significance for
fern classification are stomata (VAN COTTHEM, 1970) and spores (LUGARDON, 1971). Gameto-
phytes, formerly a neglected item, are important and often indicate relationships within the larger
groups of ferns.
In the end, a practical taxonomy must rest on a limited number of characters which are
observable without very elaborate equipment, which is one reason why uninformed academic
botanists regard it as unscientific. I hope and believe that this Flora is producing new contribu-
tions to that end.
Work now in progress for further instalments of Flora Malesiana Series II is as follows.
Dr E. HENNIPMAN, with collaborators, has begun a study of Polypodiaceae (s.str.); Prof. K.
IWATSUKI is making progress with Hymenophyllaceae; Mr G. J. DE JONCHEERE is working on the
Davallia group; Dr B. S. CROXALL is studying the complexities of Grammitidaceae; Prof. K. U.
KRAMER has started on Preris, which seems to me to be an isolated genus; Prof. T. C. CHAMBERS
has made a world monograph of the genus Blechnum and it is hoped that he will be able to deal
with the Blechnum group for Flora Malesiana; Mr A. C. JERMy is working on the complex genus
Selaginella.
Of the other genera, Dennstaedtia is of basic importance. It is more diversified in the Philip-
pines, New Guinea and the Pacific than in any other part of the world and, as the fronds are very
large, existing herbarium material often does not give full information about them; more field
work, by people who know what to look for, is needed. The most complex groups still not dealt
with are those of Tectaria and Athyrium, the latter being very difficult, with need of much new
observation, especially of scales. I have made studies of two genera of the Tectaria group and
propose to continue with that group if I am able to do so. The Adiantum group (as listed in 1959) is
complex, but not so well developed in Malesia as in drier climates, and is more dependent on
studies of plants in such climates than are most other groups. This is true also of the Dryopteris
group, which is mainly temperate in distribution.
December 1979 R. E. HoLtruM
(17)
FLORA MALESIANA
REFERENCES
ALDERWERELT VAN ROSENBURGH, C. W. R. K. VAN: 1908. Malayan Ferns: Handbook to the
determination of the ferns of the Malayan Islands. Landsdrukkerij, Batavia.
— 1917. Ibid. Supplement.
BACKER, C. A. & O. PostHuMus. 1939. Varenflora voor Java. ’s-Lands Plantentuin, Buitenzorg.
BAKER, J. G. 1891. A summary of new ferns discovered since 1874. Ann. Bot. 5: 181-221, 301-
332, 455-500.
BEDDOME, R. H. 1883. Handbook to the ferns of British India and the Malay Peninsula. Thacker,
Spink & Co., Calcutta.
— 1892. Ibid. Supplement.
BERNHARDI, J. J. 1801. Tentamen alterum filices in genera redigendi, in Schrader, Journ. Bot.
1800, 2: 121-130.
BLuME, C. L. 1828. Enumeratio Plantarum Javae. Van Leeuwen, Leiden.
— 1829-1830. Flora Javae; Filices, t. 1-65. H. Remy, Bruxelles.
Bower, F. O. 1928. The Ferns, vol. III. Cambridge University Press.
CHING, R. C. 1936. A revision of the Chinese and Sikkim-Himalayan Dryopteris, I. Thelypteris.
Bull. Fan Mem. Inst. Biol. Bot. 6: 237-350.
—— 1940. On natural classification in the family ‘Polypodiaceae’. Sunyatsenia 5: 201-268.
CurisT, H. 1897. Die Farnkréuter der Erde. G. Fischer, Jena.
—— 1899. Monographie des Genus Elaphoglossum. Neue Denkschr. allg. Schweiz. Ges.
Naturwiss. 36: 1-159.
—— 1907. The Philippine species of Dryopteris. Philip. J. Sci. Bot. 2: 189-217.
CHRISTENSEN, C. 1905-1906. Index Filicum. Hagerup, Copenhagen.
1911. On a natural classification of the species of Dryopteris, in Biologiske Arbejder,
tilegnede Eug. Warming, ed. L. K. Rosenvinge. Hagerup, Copenhagen.
— 1913. A monograph of the genus Dryopteris, Part 1. K. Dansk Vid. Selsk. Skr. VII, 10 (2):
55-282.
— 1920. Ibid. Part 2. /.c. VIII, 6 (1): 1-132.
—— 1932. The Pteridophyta of Madagascar. Dansk Bot. Ark. 7: 1-253, pl. 1-80.
—— 1939. Filicinae. Chapter XX in Fr. Verdoorn, Manual of Pteridology. M. Nijhoff, The
Hague.
— 1943. A revision of the Pteridophyta of Samoa. Bernice P. Bishop Museum, Bulletin 177.
Honolulu.
— &R. E. Horttum. 1934. The ferns of Mt Kinabalu. Gard. Bull. Str. Settl. 7: 191-324,
pl. 51-62.
COPELAND, E. B. 1933. Trichomanes. Philip. J. Sci. 51: 119-280, t. 1-61.
— 1937. Hymenophyllum. /.c. 64: 1-188, t. 1-189.
— 1938. Genera Hymenophyllacearum. /.c. 67: 1-110, t. 1-11.
—— 1947. Genera Filicum. Chronica Botanica, Waltham, Mass.
— 1958-1960. Fern Flora of the Philippines. Institute of Science and Technology, Monograph
6, Manila.
CoTTHEM, W. VAN. 1970. Comparative morphological study of the stomata in the Filicopsida.
Bull. Jard. Bot. Nat. Belg. 40: 81-239.
Fre, A. L. A. 1852. Mémoires sur la famille des Fougéres, V. Genera Filicum. Paris & Stras-
bourg.
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Dedication
Ho.ttvuM, R. E. 1932. On Stenochlaena, Lomariopsis and Teratophyllum in the Malayan Region.
Gard. Bull. Str. Settl. 5: 245-312.
—— 1947. A revised classification of Leptosporangiate Ferns. Journ. Linn. Soc. Bot. 53: 123-
158.
— 1951. The fern-genus Pleocnemia Pres]. Reinwardtia 1: 171-189.
— 1955. A Revised Flora of Malaya, vol. 2, Ferns of Malaya. Government Printer, Singapore.
— 1959. Flora Malesiana, Ser. II, vol. 1, pt 1, Introduction and Gleicheniaceae.
— 1963. Ibid. pt 2, Cyatheaceae.
— 1973. Copeland’s contribution to fern taxonomy. Philippine Agriculturist 57: 17-20.
— 1974. The fern-genus Pleocnemia. Kew Bull. 29: 341-357.
—— 1978. Flora Malesiana, Ser. II, vol. 1, pt 4, The Lomariopsis Group.
Hooker, W. J. 1838-1842. Genera Filicum. G. Bohn, London.
— 1844-1864. Species Filicum, vol. 1-4, W. Pamplin; vol. 5, Dulau. London.
— & J. G. BAKER. 1865-1868. Synopsis Filicum. R. Hardwicke, London.
—— & —— 1874. Ibid. ed. 2.
— &R. K. GREVILLE. 1827-1831. Icones Filicum; vol. 1, 1827-28; vol. 2, 1829-31. Treuttel &
Wiirtz, London.
KLeEKowskKI Jr, E. J. 1979. The genetics and reproductive biology of ferns, in Dyer, A. F. (ed.),
The experimental biology of ferns. Exp. Bot. 14: 133-170.
Kunn, M. 1869. Filices. Ann. Mus. Bot. Lugd.-Bat. 4: 276-300.
KunzZE, G. 1840-1851. Die Farrnkrauter in kolorirten Abbildungen (2 volumes). E. Fleischer,
Leipzig.
Lovis, J. D. 1977. Evolutionary patterns and processes in ferns. Adv. Bot. Res. 4: 229-415.
LuGARDON, B. 1971. Contribution a la connaissance de la morphogénése et de la structure des
parois sporales chez les filicinées isosporées. Thése, Toulouse, 257 pp., 51 tab.
MANTON, I. 1950. Problems of cytology and evolution in the Pteridophyta. Cambridge University
Press.
METTENIUS, G. 1856. Filices horti botanici Lipsiensis. L. Voss, Leipzig.
— 1860. Uber Seitenknospen bei Farnen (reprint at Kew; origin ?).
— 1863A. Uber den Bau von Angiopteris. Abhandl. math. phys. Cl. K. Sachs. Gew. Wiss. 6:
501-570, t. i-x.
—— 1863B. Filices, praesertim Indicae et Japonicae. Ann. Mus. Bot. Lugd.-Bat. 1: 46-58.
—— 1864A. Ibid., para altera; /.c. 1: 222-241.
— 1864B. Uber die Hymenophyllaceae. Abhandl. math. phys. Cl. K. Sachs. Gew. Wiss. 7:
403-504, t. i-v.
— 1866. Filices; praesertim Indicae et Japonicae. Ann. Mus. Bot. Lugd.-Bat. 2: 219.
MILpE, J. 1866. Das Genus Athyrium. Bot. Zeit. 24: 373-376.
— 1870. Uber Athyrium, Asplenium und Verwandte. Bot. Zeit. 28: 329-337, 345-354, 370-
S71.
Moorg, T. 1857. Index Filicum, I. Synopsis of the genera of ferns. W. Pamplin, London.
PICHI SERMOLLI, R. E. G. 1977. Tentamen Pteridophytorum genera in taxonomicum ordinem
redigendi. Webbia 31: 313-512.
PRESL, C. B. 1825. Pteridophyta, in Reliquiae Haenkeanae, fasc. 1: 14-84. Prague.
— 1836. Tentamen Pteridographiae seu Genera Filicacearum. Prague.
RIDLEY, H. N. 1926. The ferns of the Malay Peninsula. J. Mal. Br. R. As. Soc. 4: 1-121.
SCHKUHR, C. 1804-1809. Vier und zwanzigste Klasse der Linnéischen Pflanzensystems oder
Kryptogamische Gewichse, vol. 1-3. Wittenberg.
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FLORA MALESIANA
Scuott, H. 1834. Genera Filicum. Wallishauser, Vienna.
SmiTH, JAMES E. 1793. Tentamen botanicum de Filicum generibus dorsiferarum. Mem. Acad.
Turin 5: 401-422.
SMITH, JoHN. 1841-1843. An arrangement and definition of the genera of ferns, in Hook. J. Bot.
4: 38-70, 147-198; in Hook. Lond. J. Bot. 1: 419-438, 659-668; 2: 378-394.
— 1855. Filices, in Seemann, Botany of the voyage of H.M.S. Herald: 226-244. L. Reeve,
London.
—— 1866. Ferns, British & Foreign. Hardwicke, London.
—— 1875. Historia Filicum. Macmillan, London.
SWARTZ, O. 1801. Genera et Species Filicum ordine systematico redactarum, in Schrader, Journ.
Bot. 1800, 2: 1-110.
— 1806. Synopsis Filicum. Kiliae.
WALKER, T. G. 1979. The cytogenetics of ferns, in Dyer, A. F. (ed.), The experimental biology
of ferns. Exp. Bot. 14: 87-132.
(20)
ly ENTRODUC TORY ‘NOTE
The work of preparation of a new survey of all the Pteridophytes of Malaysia
will occupy a considerable period. It is proposed to publish this work in parts,
as studies of particular families or genera are completed, but it is impossible
to plan in advance the precise sequence of these studies.
It is anticipated that the new information to be recorded, and new ideas based
upon it, will throw a good deal of new light on the delimitation of genera, and
upon the inter-relationships of genera, especially among the ferns, which are by
far the largest of the major groups concerned. Therefore one cannot now predict
what final scheme of classification will emerge. But it is necessary to have some
sort of conspectus at the start, as a preliminary survey of the ground to be covered,
and as a guide for those who wish to consult the parts of the work as they appear.
I have therefore drawn up a list of the major groups, with the genera in each,
and also a series of keys to the genera of ferns. The nomenclature of the major
groups, the generic concepts, and the keys, must all be regarded as tentative.
As regards the major groups, there are differences of opinion as to the status
of each and as to the most appropriate names. I hope at least that the significance
of the names here used is clear. As regards the ferns, there is still no general
agreement as to the subdivision of the vast and heterogeneous assemblage formerly
known as Polypodiaceae, though the more primitive families are clearly defined.
In the conspectus which follows, I have arranged the genera of Polypodiaceae
sens. lat. in Groups which seem to me to be natural, but I have given no formal
names to the Groups, nor specified their status, except for Polypodiaceae sens.
strict. and Grammitidaceae, which (with the exception of a few genera) are also
accepted by COPELAND. The fern specimens in the Herbarium at Bogor are now
arranged according to these Groups. Probably the Groups will be modified in
any final re-assessment at the conclusion of their treatment in ‘Flora Malesiana’.
So far as possible, genera within a family or Group will be treated together.
There are two alternative General Keys, which lead to families or Groups of
genera. In the first key a principal distinction is between epiphytes and terrestrial
ferns; in the second key this distinction is not used.
The delimitation of some genera of ferns is still not clear, and the generic names
are to be taken as tentative. In general, they are those accepted in COPELAND’S
Genera Filicum (1947), but there are some changes.
Critical literature on fern taxonomy up to the year 1934 is very fully listed in
CHRISTENSEN’S Index Filicum and its three Supplements. Some important books
and papers subsequently published and dealing with Malaysian ferns are listed
after the keys to genera.
R. E. Horrrum
2. LIST OF MALAYSIAN, PTERIDOPHYTES
PSILOTOPSIDA Colysis Loxoscaphe
PSILOTALES Crypsinus Athyrium Group
Dendroconche Anisocampium
Psilotaceae Dendroglossa Athyrium
Est orm Diblemma Callipteris
Tmesipteridaceae Dipteris Cornopteris
Tmesipteris Drymoglossum Cystopteris
Drynaria Diplaziopsis
LYCOPSIDA Goniophlebium piplaziers
LYCOPODIALES Grammatopteridium Dryoathyrium
: Holcosorus
Lycopodiaceae Holostachyum Blechnum Group
Lycopodium Lecanopteris Blechnum
Selaginellaceae Lemmaphyllum Brainea
Selaginella Lepisorus Doodia
Isoetaceae Leptochilus Weed ee
Isoetes Loxogramme Davallia Group
Merinthosorus Araiostegia
SPHENOPSIDA Microsorium Davallia
EQUISETALES Myrmecophila Davallodes
j Neocheiropteris Humata
Equisetaceae : :
: Oleandropsis Leucostegia
Equisetum
Paragramma Parasorus
PTEROPSIDA Photinopteris Scyphularia
6 Platycerium Trogostolon
pianist Polypodiopsis Dennstaedtia Group
Ophioglossaceae Pycnoloma Dennstaedtia
Botrychium Pyrrosia Histiopteris
Helminthostachys Selliguea Hypolepis
Ophioglossum Thayeria Microlepia
MARATTIALES Thylacopteris Monachosorum
Warattinceae Grammitidaceae Orthiopteris
: ; Acrosorus Paesia
Angiopteris a
: . Calymmodon Pteridium
Christensenia i
Macroglossum Ctenopteris Dicksonia Group
Marattia Grammitis — Cibotium
Nematopteris Culcita
FILICALES Oreogrammitis Cystodium
Osmundaceae Prosaptia Dicksonia
Leptopteris Scleroglossum Dryopteris Group
Osmunda Xiphopteris Acrophorus
Schizaeaceae REMAINING GENERA OF FERNS Currania
Lygodium (in Groups, alphabetically) Diacalpe
Schizaea Adiantum Group Didymochlaena
Gleicheniaceae Adiantum Dryopteris _
Dicranopteris Anogramma Gymnocarpium
Gleichenia Ceratopteris Peranema
Hymenophyllaceae Cerosora Polystichum
Cheilanthes Polystichopsis
Hymenophyllum ; Stenolepia
‘ Coniogramme p
Trichomanes : =
‘ Doryopteris Lindsaea Group
Matoniaceae Hemionitis Isoloma
Matonia Notholaena Lindsaea
Phanerosorus Onychium Sphenomeris
Cyatheaceae Pellaea Tapeinidium
Cyathea Pityrogramma Lomariopsis Group
Polypodiaceae (s.str.) Syngramma Arthrobotrya
Aglaomorpha Schizolepton Bolbitis
Arthromeris Taenitis Egenolfia
Belvisia Asplenium Group Elaphoglossum
Cheiropleuria Asplenium Lomagramma
Christiopteris Diplora Lomariopsis
II
Morphology of ferns
Teratophyllum Ctenitis Dictyocline
Thysanosoria Cyclopeltis Sphaerostephanos
Nephrolepis Group Dryopolystichum Stegnogramma
Arthropteris Hemigramma Thelypteris
Nephrolepis Heterogonium Vittaria Group
Oleandra Hypodematium Antrophyum
Plagiogyria Group SESH Monogramma
Plagiogyria Luerssenia Vaginularia
: Pleocnemia Vittaria
Pteris Group Psomiocarpa
Acrostichum Pteridrys HETEROSPOROUS FERNS
Hemipteris Quercifilix Marsileaceae
= aia Stenosemia Marsilea
mEIS Tectaria +6;
Schizostege oak
Stenochlaena Thelypteris Group Salvinia
Tectaria Group Ampelopteris
Arcypteris Cyclosorus
THE MORP HOVMOGY-OFeFERNS
A fern plant consists of a stem, bearing leaves and roots. The leaves (or some of them) bear
dehiscent sporangia, each sporangium containing unicellular spores, which are in most cases
wind-dispersed. A spore germinates to produce a small green plant called a prothallus. The
prothallus bears sexual organs (archegonia and antheridia). After fertilization by an antherozoid,
the female cell in an archegonium grows to form a new fern plant. The life cycle of a fern thus
has two phases, asexual (the fern plant) and sexual (the prothallus). These phases are also called the
sporophyte and the gametophyte. The sporophyte is much longer-lived, larger and more diversified
than the gametophyte, and its characters are mainly used in taxonomy. The following statement
deals with the parts of the sporophyte in turn, with discussion of the kinds of modification of
each which occur, and of special terminology. Finally, a note on the gametophyte will be
given, including reference to the not infrequent condition in which the sexual process is omitted.
Stem. (a) Shape, size, and habit of growth——A fern stem may be long and creeping or
climbing, in which case it is usually called a rhizome, or it may be short and compact, in which
case it is often called a stock, rootstock or caudex. If it grows erect, as in tree-ferns, with a tuft
of leaves at its apex, it is called a trunk.
A creeping stem or rhizome may be dorsiventral or radial in construction. In the former case
the leaves (or their stalks) are borne on the upper surface (often in two alternate rows, some-
times more than two), the roots entirely or mainly on the lower surface; the internal vascular
structure corresponds to this external differentiation. In a radially constructed rhizome leaves
and roots are borne on all sides (though of course the leaf-stalks all grow upwards), and the
vascular system shows a corresponding radial symmetry.
A short stem or rootstock, bearing crowded leaves, is usually radial in construction. Such
a stem may be quite erect, but more often it is more or less decumbent; the presence of a per-
sistently erect rootstock is sometimes an important diagnostic character. Tree-ferns, and others
in which the erect rootstock grows to an appreciable height above the surface of the ground,
have numerous roots which may form a close covering on the lower part of the stem, thus giving
stability to the plant.
(b) Branching.—In a few cases this is by simple dichotomy (the apical growing-point dividing
into two equal parts). More frequently branches arise in association with the bases of leaf-
stalks, usually on the outside. The method of branching may be important taxonomically; it
has been too little studied.
(c) Hairs, Bristles and Scales.—The young parts of a stem are in almost all cases protected
either by scales or hairs; similar scales or hairs also cover the very young leaves before they
uncurl. Such scales and hairs are more or less persistent on older parts of stem and leaves.
HI
FLORA MALESIANA [ser. II, vol F
Hairs consist of a single cell or of a single row of cells, and in different ferns are of characteristic
length and thickness; the thicker ones are sometimes quite rigid and bristle-like. True bristles,
which are more than one cell thick at the base, but round in section, also occur in some cases.
Scales are flat plates of cells, one cell thick; the details of their structure (especially characters
of edges, base and apex, and the presence or absence of superficial outgrowths) are often very
important taxonomically and may need microscopic examination. A peltate scale is one attached
at some point on its surface, not on its edge. Where lateral cell-walls of a scale are thickened
so as to form a distinct lattice-work pattern, the scale is called clathrate; the upper and lower
walls may or may not be translucent.
(d) Internal Structure.-—The most important internal structure is the vascular tissue. The
vascular strands which pass into leaves and roots are connected to the vascular system of the
stem; the latter is called a stele, or if at any cross section it consists of more than one part,
the parts are called meristeles. The simplest kind of stele is a protostele; in this there is one
single solid strand of xylem with phloem around it, the whole surrounded by an endodermis.
In some cases the xylem of a protostele is not solid, but has a core of non-vascular tissue;
such a structure is called a medullated protostele. In some long-creeping rhizomes (e.g. Micro-
lepia) the stele is a hollow cylinder, with phloem and endodermis both inside and outside the
xylem; this is called a solenostele or siphonostele. In this case there is usually a gap in the stele
where the vascular supply to a leaf is attached; such a gap is called a /eaf-gap. Rhizomes of
this type are usually dorsiventral. In Davallia, the solenostele does not consist of a continuous
hollow cylinder, but the cylinder (not circular in section) is composed of a network, most of
the gaps in which are not leaf-gaps. This may be called a dissected solenostele; in the case of
Davallia it is dorsiventral, the ventral meristele being broader than the rest and bearing roots.
In stems of radial structure bearing many leaves close together, the stele usually forms a hollow
cylinder in which there are many leaf-gaps, these gaps being like the meshes in a network.
Such a stele is called a dictyostele. In a few ferns there are additional steles internal to the
principal stele (e.g. Matonia, some species of Preris); also in a few there is a cortical system of
stem-bundles outside the principal stele (e.g. Stenochlaena). Besides vascular tissue, most fern-
stems contain thick-walled tissues which serve to give mechanical strength; the presence (or
absence) and distribution of such tissue may be important.
Roots. Roots are all adventitious. The primary root of a fern-embryo, having no power
of increasing in thickness, is soon inadequate to supply the needs of the growing plant, and
further roots must grow from the stem (in some cases also from leaf-bases). Little use has
been made of root characters for taxonomic purposes, but there is considerable diversity of
root-structure among ferns as a whole. In some species of Hymenophyllaceae there appear
to be no roots, the rhizome being covered with hairs which function as root-hairs.
Leaves. VEGETATIVE CHARACTERS. Fern-leaves are usually called fronds; their stalks or
petioles are called stipes; the blade or flat green part of a leaf is called the /amina. The lamina
is usually divided into parts called /eaflets; the axes on which leaflets are borne are called
rachises (rhachis is the more correct spelling, but rachis is in general use in English). Fronds
may be sterile (lacking sporangia), or fertile (bearing sporangia); sterile and fertile fronds may
be alike in form, or dissimilar (dimorphous).
(a) Stipes —A stipe may be jointed or articulated at its junction with the stem, or in some
cases it is articulated to an outgrowth from the stem called a phyllopodium (e.g. Elaphoglossum,
Oleandra); but more commonly it is not articulated. Externally a stipe usually bears hairs or
scales like those of the rhizome or caudex, but often smaller; sometimes the scales are borne
at the ends of warts or thorns. In some cases where the rhizome bears only scales, the stipe
(and rachis) may also bear hairs. The character of the hairs, whether unicellular or multicellular
(with transverse septa), and whether terminating in a glandular cell, is always important. In a
few ferns the stipe has slime-glands, producing mucilage when the fronds are very young, and
elongate aerophores, the latter often in association with much-reduced leaflets. In any case,
there is usually a pale and more or less raised line along each side of the stipe. This line is also
nV’
Dec. 1959] 7 _ Morphology of ferns
an aerophore; there are stomata on it, and internally the tissue is thin-walled, with air-spaces
between the cells. The line is sometimes not continuous but broken, and sometimes doubled
(Cyathea). Internally the number and arrangement of the vascular bundles in a stipe is important;
often the number and arrangement change between the base and apex.
(b) Branching of fronds.—Fronds may be simple (consisting of a continuous lamina, which
may be entire or /obed in various ways), or they may consist of two to many leaflets which
are borne by the rachis or its branches. The commonest arrangement is for the main rachis
(continuing the line of the stipe) to be almost straight, bearing leaflets (stalked or not, articulate
or not) along its sides; such a frond is said to be pinnate, and the leaflets are called pinnae.
Like a simple frond, a pinna may be entire or lobed; if pinnately lobed, the leaflet is pinnatifid,
if palmately lobed, palmatifid. Where the rachis bears branches, these are usually arranged in a
pinnate manner; if each such primary branch bears leaflets, the frond is said to be bipinnate,
and the leaflets are called pinnules. If each primary branch bears secondary branches pinnately
arranged, and the secondary branches bear leaflets, the frond is said to be fripinnate, and the
leaflets are of the third order; fronds may also be quadripinnate. Pinnae or rachis-branches are
usually a/ternate on the two sides of a main rachis, but in some cases they are regularly almost
exactly opposite (e.g. Gleichenia). The edge of a leaflet on the side towards the apex of the
frond is called acroscopic, that towards the base of the frond is called basiscopic. The terms
upper and lower are used for the two surfaces of the leaflets; the upper surface is that facing
the stem-apex (also called the adaxial surface, the lower surface being abaxial).
There are some cases in which the stipe is not continued into a single straight rachis, but
forks into two equal branches at its apex; such a dichotomous branching is seen in fronds of
young plants of Lygodium. The dichotomy may be repeated, in which case a fan-shaped frond
results (Schizaea dichotoma, Dipteris lobbiana); or only the outer branch at each dichotomy
may fork again, in which case pedate branching results. The ultimate branching of large fronds
or of their veins is also often dichotomous, and one can trace a transition from dichotomous
to pinnate branching in descending a frond to its base; the pinnate condition can thus be
regarded as a development from the dichotomous one. Pseudo-dichotomy occurs in a few cases
(notably in Gleicheniaceae and in Lygodium); in these a rachis bears a pair of opposite branches
and then stops growing, its dormant apex remaining between the branches.
(c) Shape of rachises and of junctions between them and with leaflets——In addition to the
characters of superficial hairs and scales, and of internal vascular systems, rachises of ferns
provide characters of another kind which are useful taxonomically. These characters relate
to the way in which the structure of a smaller rachis is adjusted to that of a larger one where
the two join, and ultimately to the relationship of a lateral leaflet to the rachis which bears it.
Some rachises are rather deeply grooved on the upper surface, and where a smaller rachis
joins a larger one, the groove of the latter is opened to admit that of the former. In some such
cases (e.g. Dryopteris, Athyrium, Pteris) the midrib of a leaflet is also grooved, and a rachis-
groove is open to admit the midrib-groove of a lateral leaflet; the edge of the leaflet is then more
or less decurrent down the side of the rachis. In other cases the thickened basiscopic edge
of a leaflet is decurrent on the edge of the rachis-groove (e.g. Lindsaea). Other rachises are not
or only slightly grooved on the upper surface and there is no opening of the groove of a main
rachis to admit that of a smaller one (e.g. Thelypteris, Ctenitis). In such cases the base of the
lamina of a leaflet is decurrent on the side of the rachis, sometimes forming a small continuous
ridge or wing. Characters such as this are constant throughout a genus, or even throughout a
group of genera, and are often valuable diagnostically, especially in sterile fronds. I believe
also that they may provide valuable evidence for those who are seeking a natural classification
of ferns.
(d) Venation.—Veins mark the course of the vascular bundles in a leaf; they are usually
evident on the surface, as the surface tissue over a vein is different from that over the rest of
the lamina. The pattern of venation is always important taxonomically. In fronds which are
thick and fleshy or leathery the veins are sometimes quite invisible on the surface, or only the
Vv
FLORA MALESIANA [ser. II, vol. 11
larger ones may be prominent; the pattern of venation can then be seen by clearing the frond
(boiling it and then treating it with a bleaching agent; or boiling alone may be sufficient, due to
replacement of internal air by water).
In large much-branched fronds the ultimate leaflets are usually small, with few veins in each,
and these veins commonly end in lobes or teeth at the edge of the lamina. Such veins are free;
a vein may fork once or more times, but the branches do not unite again at any point. Where
leaflets are very small, the veins are often dichotomously branched, while larger leaflets usually
have a midrib or costa with lateral veins which are simple or forked.
In larger leaflets the vein-pattern may be more complex, still with free veins. The sides of a
leaflet may be deeply lobed (pinnatifid) with pinnate vein-groups in each lobe; the main vein
of a lobe is then called a costule (e.g. Thelypteris, Cyathea). In a large number of ferns with
large leaflets, the veins join neighbouring veins after branching, thus forming areas of lamina
surrounded by veins; such areas are called areo/es, and veins which unite with others are said
to anastomose. The anastomosis may consist only in the formation of a single series of areoles
along costae or costules (the rest of the veins ending freely) or it may form a more or less
elaborate network occupying almost the whole lamina. In the latter case there are several
possible patterns (e.g. Cyclosorus, Tectaria, Goniophlebium, Acrostichum). One useful character
is the presence or absence of small veins which end freely inside the areoles.
At the ends of some veins are water-excreting pores or hydathodes, often evident on the
upper surface of the lamina as distinctive spots (round or elongate) which in some cases may
ultimately become covered with white scales due to deposits of salts left after the water originally
holding them has evaporated. Such hydathodes may provide useful diagnostic characters
(e.g. Pyrrosia, Grammitis, Coniogramme).
(e) Surface characters of lamina.—The lower surface of the lamina may be glaucous (covered
with a pale blue-green waxy layer); this character often disappears if specimens are subject
to much heat in drying. The lower surface in other cases may be more or less completely covered
with a layer of white or yellow loose waxy powder; this is excreted from glandular hairs (e.g.
Pityrogramma). Hairs on the lamina are always important, and often need to be examined
with a microscope for the structure to be clearly evident. The nature and arrangement of
stomata may in some cases be significant (e.g. Schizaea). The patterns of thickenings on walls
of epidermal cells may be characteristic (or of the single cell-layer of filmy ferns); and in some
ferns there are narrow spicular cells containing silica (e.g. Vittaria). In some ferns there are
false veins, which are lines along which surface cells are more or less elongate and sometimes
devoid of stomata, simulating the surface appearance of veins but with no underlying vascular
tissue (e.g. Angiopteris, Trichomanes sect. Crepidomanes).
(f) Polymorphism of fronds.—In many ferns, the fronds of young plants have a distinctive
shape different from that of fronds of mature plants; such young plants may offer useful
diagnostic characters. Some high-climbing ferns of the Lomariopsis Group have leaves of
distinctive shape on those parts of the plant which are near the ground (always in moist shady
evergreen forest) on rocks or tree-trunks; such leaves are called bathyphylls, and the leaves of
high-climbing parts of the same plants are called acrophylls.
FERTILE LEAVES. (a) Sori.—In most ferns sporangia are borne in distinct groups called sori.
In some cases the sori spread along the veins to such an extent that they can hardly be called
groups of sporangia, and this leads to the acrostichoid state (see below). Many sori are protected
by indusia, which are thin outgrowths from the surface of a frond. The sori of other ferns are
exindusiate, but in some of these the sori are protected by being produced in depressions or
grooves, or by being covered when young by paraphyses (hairs of various forms, or scales,
borne among the sporangia). The position and shape of sori and of their indusia (if present)
are always important taxonomically. The older schemes of classification, and also that of
Hooker and BAKER (Synopsis Filicum, 1868) were based entirely on these characters. But species
closely related in every other respect may differ in presence or absence of an indusium, or in
shape of sori (especially if these lack indusia) and a natural classification must take such facts
VI
Bec. 1959] te Morphology of ferns —
into consideration. The Orders Ophioglossales and Marattiales are quite different from Filicales
in the form and arrangement of their sporangia (see statement on sporangia below); further
remarks here refer to Filicales.
A sorus may be more or less circular, or elongate. If circular, it may be at the end of a free
vein (at the edge of the lamina or not) or seated upon a free vein, or at the junction of veins
in a network, or at the end of a free vein enclosed in an areole surrounded by other veins.
In any of these cases it may be protected by an indusium, which may be pocket-shaped (at-
tached at base and sides), or kidney-shaped, or circular and attached by the edge or the centre,
or cup-shaped, or of other shapes, or it may have no indusium. If the sorus is at the end of
a vein, it may be protected by the thin reflexed edge of the lamina, or by two more or less equal
outgrowths from upper and lower surface (Dicksonia), the two outgrowths sometimes more or
less joined to form a protective funnel or cup (Trichomanes, Dennstaedtia). Sori which are
elongate may spread along free veins, or along veins which anastomose, or they may spread
along the margin, joining the ends of veins which in a sterile leaflet would be free (fusion-
sori or coenosori) or they may lie close to the costa of a leaflet (Blechnum). Marginal fusion-
sori are sometimes protected by an inner indusium (Lindsaea), sometimes also by the reflexed
margin (Preridium), but if the margin is reflexed, the inner indusium may be lacking (Pteris).
It is especially exindusiate sori which spread along veins away from the margin, often to a
different extent in closely related species. The above survey of soral form is not exhaustive,
but is intended as an indication of the possibilities and as a guide in using the keys which
follow. Details of individual soral forms will be given in the taxonomic treatment of the families
and genera.
(b) Dimorphism of leaves.—In many ferns the fertile fronds differ in shape from sterile ones.
In such cases the fertile fronds often have a lamina of reduced size; this reduction may be slight
or it may be so considerable that the sterile and fertile fronds are quite different in aspect.
Fertile fronds also often have longer stipes than sterile, and in some cases this may be the
chief difference between the two.
(c) The Acrostichoid state.—Where sori spread along all the veins of a fertile leaflet and the
leaflet is of reduced size as compared with a sterile one of the same species, the ripe sporangia
may be so close together that they entirely cover the lower surface of the fertile leaflet (they
may also grow from the surface of the lamina between the veins); this is called the acrostichoid
condition, the name being taken from the Acrostichum, in which only the upper leaflets are
fertile in this way. Formerly all acrostichoid ferns were included in the genus Acrostichum,
but the acrostichoid state has certainly arisen along several different evolutionary lines, and a
genus based on it alone is a very unnatural one. In some acrostichoid ferns there is an additional
vascular system close to the lower surface of the lamina, in addition to the normal system found
in sterile fronds of the same species.
Sporangia. (a) Ophioglossales. —Here the sporangia are large, spherical, opening by
slits, and are attached to spike-like or branched outgrowths from the base of the lamina; the
fertile part of the frond is thus not fern-like in aspect.
(b) Marattiales—In this Order the sporangia are also large, more or less laterally joined
together in linear or circular groups on the surface of the lamina (along veins or at vein-
junctions); they do not have an annulus comparable with that of members of the Order Filicales.
(c) Filicales—In the great majority of this order the sporangia have a basically similar
structural plan, in which dehiscence is caused by contraction on drying of a more or less com-
plete ring of cells (the annulus) which have inner walls thickened but outer walls thin; there is
also a particular place (stomium) where rupture occurs. The more primitive families (Osmunda-
ceae, Schizaeaceae, Gleicheniaceae) have a less specialized development of the annulus, that of
Osmundaceae being the least specialized (its annulus is not ring-shaped). In Hymenophyllaceae,
Cyatheaceae, Plagiogyria and some other genera the annulus is complete and ob/ique in its position
on the sporangium; in the great majority of genera the annulus is almost vertical and incom-
plete, being broken by the stalk, but even in these cases the structure of the sporangium is not per-
VII
FLORA MALESIANA
fectly symmetrical when divided along the plane of the annulus. In some members of the
Adiantum Group the cells of the annulus are broad and more or less uneven. A detailed study of
the development and structure of sporangia has been made in comparatively few fern-genera.
Spores. Fern spores are aiways produced in groups of four (tetrads), each tetrad normally
the result of the meiotic divisions of one spore-mother-cell. A spore may have either of two
distinct shapes, monolete (or bilateral), and trilete (or tetrahedral). Monolete spores are more or
less bean-shaped (like a Phaseolus seed), with an angle along the straight edge where the spore
is in contact with the similarly angled edge of another spore of the tetrad; in each tetrad there
are two such pairs. Trilete spores meet together on three faces, and at the angles between them,
all four spores meeting at the centre of a tetrad. Usually all species in a genus have spores of
the same shape, but there are certainly some genera in which both shapes of spores occur
(e.g. Dicranopteris), and I have seen evidence that even within a single species there may be
spores of both kinds.
The inner layer of cells in the wall of a sporangium (called the tapetum) breaks down during
the development of the tetrads of spores, its substance being absorbed by the spores during
their development. In some cases part of the substance of the tapetum forms an external covering
on each spore, known as a perispore. The perispore is usually more or less folded into rather
irregular wing-like structures, or sometimes into more regular spines. Other genera of ferns
lack a perispore, and then the wall of the spore itself may be variously sculptured into a more
or less complex pattern of warts or ridges. The presence or absence of a perispore, its structure
if present, or the wall-characters where there is no perispore, are always of taxonomic importance
as well as the actual size of the spores.
In the case of hybrids, where normal meiosis does not occur, there are often shrivelled
empty spores, and the presence of such is always significant. In the case of apogamous ferns
(see note on the gametophyte below) the spores are of at least two kinds, large functional ones
and smaller ones which are not functional (for a detailed statement, see MANTON, Problems of
Cytology and Evolution in the Pteridophyta, Cambridge, 1950).
Gametophyte. In Ophioglossales the gametophytes are subterranean and saprophytic, and
obtain their nutriment through the activity of an endophytic fungus. In all other ferns the
gametophytes are green, and in the vast majority of cases they are more or less heart-shaped,
with a growing-point in the sinus between the two lobes; they are thickened in a median area
which bears the rhizoids, antheridia and archegonia on the lower surface. Distinctive characters
are provided by shape of the whole prothallus (in some cases this is asymmetric or elongate),
presence of superficial hairs of different kinds, colour and septation of rhizoids, and especially
in details of structure of archegonia and antheridia. In a few cases the gametophyte is more or
less filamentous (Schizaea, Hymenophyllaceae), or irregularly lobed (Vittaria and allied genera).
In apogamous ferns, no sexual process occurs. The prothallus is developed from a diploid
spore; it bears antheridia but no archegonia, and gives rise to a new sporophyte by vegetative
budding. The diploid antherozoids of such a prothallus may fertilize haploid archegonia of a
sexual prothallus of an allied species, the result being a triploid hybrid sporophyte. Such a
hybrid is normally sterile, but may develop vegetative means of propagation not involving the
formation of a prothallus; and some such hybrids have become apogamous. Higher polyploid
plants are also not uncommon among ferns, and in such cases cytological evidence is of great
taxonomic value.
Heterosporous Ferns. The two families Salviniaceae and Marsileaceae are very different
from other ferns in many respects. They have spores of two kinds, large and small, in separate
sporangia. The small spores produce very small gametophytes which are male, and the large
spores produce larger female gametophytes. Both kinds of sporangia are formed inside closed
structures called sporocarps, borne by the leaves. These ferns are all aquatic, and they are
sometimes collectively called Hydropterideae or Hydropteridales, but the two families are not
closely related, and probably had quite different evolutionary histories.
VIII
4. GENERAL KEY No 1 to PTEROPSIDA
1. Aquatic plants.
2. Plants floating. Leaves small, simple or bilobed sev, Swryrelaee Salviniaceae
2. Plants rooted in earth or on rocks. Leaves larger, more divided.
3. Leaves 4-partite. Sporocarps attached to stipes. ee : Marsileaceae
3. Leaves not 4-partite. Sporangia singly or in sori on lower surface ‘of lamina.
4. Sporangia borne singly, protected by reflexed edges of narrow lamina . Adiantum
Group
4. Sporangia grouped in sori, on lower surface of lamina, not protected by reflexed edges.
5. Fern of stream-beds in deep shade. Fronds pinnatifid, sori without indusia Polypodiaceae
5. Fern of open swamps. Fronds bipinnatifid, sori indusiate. . . . . Thelypteris
1. Land plants or epiphytes.
6. Epiphytes.
Group
7. Fronds simple, not over 2 mm wide, with a single vein, or with a few simple lateral soriferous
veins close to the main vein . . FEM MORK Vittaria
7. Fronds branched, or if simple with a more * complex venation.
Group
8. Lamina one cell thick apart from midribs of segments. . . . . . Hymenophyllaceae
8. Lamina throughout more than one cell thick.
9. Sporangia embedded in slender cylindrical appendages attached to surface of frond.
Ophioglossaceae
9. Sporangia not so arranged.
10. Sori not indusiate (sometimes otherwise protected).
11. Sporangia not acrostichoid.
12. Sori superficial (not in pockets or grooves).
13. Fronds simple, pinnatifid or pinnate; if pinnate, pinnae not articulate to rachis.
14. Veins much anastomosing . . : Le, Sire Son. tee APolypodiaceac
14. Veins not or slightly anastomosings
15. Frond and stipe + hairy; spores trilete at Seat lct ae Grammitidaceae
15. Frond and stipe not hairy; spores monolete . . . . . . . Polypodiaceae
13. Fronds pinnate, pinnae articulate to rachis.
(6MRinnacventires 1) Mone 2). 32) Pe, ease a es ee BPolypodiaceae
16. Pinnae lobed . . : Nephrolepis Group
12. Sori in pockets or grooves ‘(which are sometimes marginal).
17. Sori in pockets or depressions, + circular.
18. Veins anastomosing; or, if free, fronds not hairy . . . . . . Polypodiaceae
LSsaVeins: tree, tronds more or less hairy =) 5: -senoneeeee Grammitidaceae
17. Sori elongate, in grooves.
19. Grooves all evenly oblique to costa . . . . . . Polypodiaceae
19. Grooves marginal or parallel to margin, or uneven in direction, sometimes anastomosing.
20. Scales entirely opaque, usually brown . . : Grammitidaceae
20. Scales nearly black, strongly clathrate, lumina of cells translucent Vittaria Group
11. Sporangia acrostichoid, covering entirely part or whole of a frond.
21. Veins much anastomosing; spores without perispore eos ae . . Polypodiaceae
21. Veins free or slightly anastomosing near edge; perispore present . Lomariopsis Group
10. Sori indusiate.
22. Sori elongate along veins . . . . . . . . +. +. +. +. ~~. ~~ Asplenium Group
22. Sori otherwise.
23. Sori elongate along edge of lamina.
24. Pinnae articulate to rachis . . SE a ERT eae Nephrolepis Group
24. Pinnae (if any) not articulate to rachis.
25. Rhizome protostelic . . - ATES te GRR ae Lindsaea Group
25. Rhizome with more complex vascular system , Oe BUTE Davallia Group
23. Sori otherwise.
26. Fronds articulate to rhizome.
27. Sori at ends of veins, near edge of lamina.
P8ePinnae! (if present) not joimted stomrachis.. 5) S80) -) eee Davallia Group
28. Pinnae jointed to rachis . . ys ORAS.» HOSE) PE Nephrolepis Group
27. Sori close to costa of the simple frond 22> .. he elvia omiie ..aNephrolepis*Group
26. Fronds not articulate to rhizome . . . Nephrolepis Group
6. Terrestrial plants, or climbers starting from the exons or rock- plants.
29. High-climbing rhizome starting from the ground.
30. Rhizome not dorsiventral; veins anastomosing in a narrow series of costal areoles (seen at
apex of pinna) Ae leo Peet od ce) LRAT OS Shy ee Pteris
30. Rhizome dorsiventral; veins either free or much anastomosing . . Lomariopsis
29. Terrestrial or rock plants.
Group
Group
Ix
FLORA MALESIANA [ser: II, vole
31. Caudex massive, erect; stipes succulent, with stipule-like outgrowths at their bases; bases of
pinnae swollen Pe eat eras ee ge Loo oe ee Marattiaceae
31. Not these characters.
32. Lamina one or two cells thick apart from midribs of segments of lamina; no stomata.
33. Sporangia attached to elongating slender receptacles in funnel-shaped pockets at ends of veins.
Hymenophyllaceae
33. Sporangia attached to surface of veins . Ole Vheile. Bees rs eee Osmundaceae
32. Lamina throughout more than two cells thick; stomata present.
34. Caudex or rhizome at apex, and bases of stipes, hairy or bristly (no flat scales) or apparently
naked.
35. Rootstock massive, erect (in a few cases tree-like) or more or less decumbent, radially or-
ganized, its apex above ground, bearing a close group of fronds.
36. Fronds simply pinnate; apex of caudex not densely hairy.
37. Aerophores at bases of stipes (sometimes also at bases of ae Plagiogyria Group
37. Aerophores lacking : fey Osmundaceae
36. Fronds more amply divided; apex of caudex densely ‘hairy : Dicksonia Group
35. Rootstock otherwise, usually entirely below ground.
38. Fertile part of frond not leaf-like, erect and attached to base of leafy part Ophioglossaceae
38. Fertile part of frond leaf-like, sometimes reduced in size as compared with sterile.
39. Fronds palmately divided; leaflets 3 or 5; veins anastomosing . . Marattiaceae
39. Fronds otherwise.
40. Veins much anastomosing, with free veins in the areoles. . . . Polypodiaceae
40. Veins in most cases free; where anastomosing, no free veins in areoles.
41. Sori quite superficial, on lower surface of lamina, or in a marginal groove.
42. Sori indusiate; fronds fan-shaped or slender and trailing . . . Matoniaceae
42. Sori not indusiate.
43. Fronds repeatedly pseudo-dichotomous, with a dormant apex between each pair of
branches = «0 =. 3) Se Boe ie. Cy Soe Boe © Gleicheniaceae
tae Fronds otherwise. . sa Sale: Law 2 Adiantum Group
Sori at ends of veins or on 1 special appendages.
Sporangia on special appendages which are at ends of veins of leaflets or attached
near apex of frond or of its branches. . . Sol ter Schizaeaceae
44. Sporangia in sori at ends of single veins or uniting ends of several veins, not on special
appendages . . : Dennstaedtia Group
34. Caudex or rhizome at apex, and ‘bases of stipes (at least when young) scaly.
45. Sporangia acrostichoid.
46. Rhizome dorsiventral, creeping on rocks.
47. Veins free, or if anastomosing the free veins almost all pointing outwards; spores with
perispore - : Lomariopsis Group
47. Veins much anastomosing ‘with free veins in areoles pointing all ways; no perispore.
Polypodiaceae
46. Rhizome not dorsiventral, often massive, bearing a tuft of fronds at its apex.
48. Only the upper pinnae fertile; veins much anastomosing, no free veins in areoles.
Pteris Group
48. Whole frond fertile; in sterile frond veins free or anastomosing otherwise.
49. Veins free in sterile fronds, or a single row of costal areoles present.
SOs Bertiestrond simplyapimnate. ) =e eee cc) ee Blechnum Group
50. Fertile frond bipinnate . . Ty tate Sink She Tectaria Group
49. Veins much anastomosing in sterile fronds 2 its Fa ae Tectaria Group
- Sporangia not acrostichoid.
. Sorus along edge of lamina, continuous or nearly so.
s Edge of lamina reflexed, protecting sori.
53. Rachis grooved on upper surface, groove open to admit groove of midrib of pinna.
Pteris Group
53. Rachis not so grooved (if grooved, edge of lamina may be decurrent on edge of groove).
Adiantum Group
52. Edge of lamina not reflexed; sorus protected by indusium attached below it, opening
fowaLrdsvedeesotslaminay 2, %) -irayl eens tee Meera eee Lindsaea Group
51. Sorus otherwise.
54. Sorus elongate, continuous along each side of costa of pinna ; Blechnum Group
54. Sorus otherwise.
55. Sporangia on surface of reflexed marginal lobes . . . . . Adiantum Group
55. Sporangia not on such lobes.
56. Sorus along veins (at least some of them).
57. Sorus indusiate.
Pec. 1959] General key No. 1 to Pteropsida
58. Sorus symmetrically divided by line of vein.
59. Rachis grooved, groove open to admit groove of branch; scales lacking superficial
Haws, = Dryopteris Group
59. Rachis somewhat srooved, groove not open to admit groove of pinna; scales with
superficial hairs . . CW Ridtysbor e% Thelypteris Group
58. Sorus asymmetric, or on one side of 1 vein.
60. Sori along outer veins of costular or costal areoles . . . Blechnum Group
60. Sori otherwise.
61. Two strands in stipe, uniting upwards to form a single X-shaped strand.
Asplenium Group
61. Two strands in stipe, uniting upwards to form a single U-shaped strand.
Athyrium Group
57. Sorus not indusiate.
62. Sori spreading along all veins of lower surface.
63. Slender unicellular hairs present on frond and on scales . Thelypteris Group
63:3 Slender: unicellular shairsvlacking eens 9 -e Ae e Adiantum Group
62. Sori not spreading along all veins.
64. Several vascular bundles in stipe. . . a be Tectaria Group
64. Two vascular bundles at base of stipe, uniting upwards.
65. Waxy powder on lower surface of lamina x (rime e Seer. Adiantum Group
G5s. No waxyepowder present... 225 ees ese ee) eee one Athyrium Group
56. Sori not along veins.
66. Sori at ends of veins, at or close to edge of lamina, each in the base of a cup, or protect-
ed by an indusium attached below it or by the refiexed edge of the lamina.
67. Sori each in the base of a cup. ey Dennstaedtia Group
67. Sori protected by indusia or by edge of lamina.
68. Sori protected by indusia opening outwards.
692, Pinnae,articulatestoprachis) gre), fs sb act see 2 Nephrolepis Group
69. Pinnae not articulate . . vy Tie gs Lindsaea Group
68. Sori protected by reflexed edge of lamina.
70. An inner indusium also present; rachis sess groove open to admit groove
of midrib of pinna. . : Pteris Group
70. An inner indusium lacking: rachis not grooved, or if grooved, edge of pinna decur-
rent on edge of groove . . a: Adiantum Group
66. Sori not at ends of veins, or if so not ‘close to edge of lamina.
71. Rhizome dorsiventral.
72. Sori without indusium.
73. Fronds lacking dormant apices . . . emi. ©; Polypodium Group
73. Fronds always having some dormant apices oil Sieeet @ scan Gleichemacese
72. Sori indusiate.
74. Fronds simple . . i de doocwl. sheet. er eee NephrolepisiGronp
74. Fronds pinnately branched wit). qailesl. see Se eee Davallia Group
71. Rhizome not dorsiventral.
75. Fronds simple and jointed at base, or pinnate with pinnae jointed to rachis.
Nephrolepis Group
75. Fronds otherwise.
76. Tree-ferns; sporangia with complete oblique annulus; many vascular bundles in
Stipe: 2: i Cyatheaceae
76. Not tree- ferns; annulus vertical, ‘interrupted; vascular bundles in a simple ring
(except Pleocnemia).
77. Rachis grooved, groove open to admit groove of branch-rachis or pinna.
78. Several vascular bundles in stipe. . . . . Dryopteris Group
78. Two bundles, joining to form one of U-shape . give: Athyrium Group
77. Rachis not grooved, or if grooved groove not open to admit groove of branch.
79. Hairs (if present) multicellular; scales lacking superficial hairs or glands.
80. Several vascular bundles in stipe . . . .-. . . .- Tectaria Group
80. Two bundles, joining to U-shape . . ; Athyrium Group
79. Hairs unicellular; scales bearing superficial hairs or glands Thelypteris Group
5. GENERALSREY Nor24oePRTEROPSI DA
1. Aquatic plants.
2. Plants floating; leaves small, simple or bilobed. . . 3) ep RE, Salviniaceae
2. Plants rooted in earth or on rocks; leaves larger, more divided.
3. Leaves 4-partite; sporocarps attached to stipes cae : Marsileaceae
3. Leaves not 4-partite; sporangia singly or in sori on lower surface of lamina.
4. Sporangia borne singly, protected by reflexed edges of narrow lamina . Adiantum Group
4. Sporangia grouped in sori, on lower surface of lamina, not protected by reflexed edges.
5. Fern of stream-beds in deep shade; fronds pinnatifid, sori without indusia Polypodiaceae
5. Fern of open swamps; fronds bipinnatifid, sori indusiate. . . . Thelypteris Group
1. Land plants or epiphytes.
6. Stipe containing 3 or more vascular bundles throughout.
7. Rhizome in most cases dorsiventral, stipes jointed to it or to phyllopodia borne by rhizome.
8. Sori of various form (in some cases acrostichoid), not indusiate.
9. Fronds pinnate; pinnae articulate to rachis, veins free wile Bn ES Nephrolepis Group
9. Fronds rarely pinnate; pinnae (if present) not articulate to rachis, or if so veins reticulate.
10. Acrostichoid; spores with perispore; veins free or uniting near margin Lomariopsis Group
10. Acrostichoid or not; no perispore; where acrostichoid, veins much anastomosing.
Polypodiaceae
8. Sori indusiate.
11. Fronds simple.
12. Sori near midrib of frond . . . Wale MEAL. Nephrolepis Group
pe Sori at ends of veins, solitary or joining many VEINS: MAL SOs We Davallia Group
Fronds more or less branched.
me Rimnaemointedatoprachisit |eciaaha stl coe Ghee cunt) eee ne Nephrolepis Group
13. Pinnae not jointed to rachis. . ileieg -£ Davaliia Group
7. Rhizome not dorsiventral, or if so the stipes not jointed to. it.
14. Stipes containing numerous bundles in a complex pattern, not a simple ring, in cross section.
15. Rootstock massive, short or tall; if creeping, subterranean.
16. Mangrove plants; distal pinnae acrostichoid, lower ones sterile . . . Pteris Group
16. Not mangrove plants; not acrostichoid.
17. Stock and stipes Se woody tissue; no stipules at bases of stipes.
Hiss WSIS JOS 5 aoe iy eC RAMS €thch (0eet a Og Es ag SORA eB A Cyatheaceae
18. Veins anastomosing . . . . Tectaria Group
17. Stock and stipes without woody tissue; stipules ‘present at bases of stipes.
Marattiaceae
15. Rootstock slender, long-creeping or high-climbing ATA died Be Pa: Pteris Group
14. Stipes containing a simple open ring of bundles.
19. Sporangia on a separate branch attached at base of lamina, or to surface of lamina; no annulus.
Ophioglossaceae
19. Sporangia not on a separate branch of frond; annulus present.
20. Fronds pinnate or bipinnate, leaflets jointed to rachis.
21. Fronds in close tufts, not spaced on a climbing rhizome; sori separate and indusiate.
Nephrolepis Group
21. Fronds spaced on a climbing rhizome; sori not indusiate, in most cases acrostichoid.
Lomariopsis Group
20. Leaflets (if present) not jointed to rachis.
22; Fronds simple, sori oblique, exindusiate . . . . . . . . =. . BPolypodiaceae
22. Fronds otherwise.
23. Sori elongate, either close to midrib or to edge.
24 eS OnMCLOSERtOMMIdnI Digan). rss ree ay Oe, Devi) ea Blechnum Group
PASSO closesto) edges.) =) - b pe bis Seley Ae Te Nephrolepis Group
23. Sori not elongate near midhibe or ‘edge.
25. Sporangia acrostichoid, edge of lamina in some cases reflexed.
26. Veins of sterile leaflets free.
Oi eEronds“bipinnate, ¢ sa. 7 2Gthetn anes tach Ieee Tectaria Group
27. Fronds simply pinnate.
28. Rhizome creeping, dorsiventral A slg ool poo ae Lomariopsis Group
At, Uinecomte One, walellhy gyacinyel 2995 6 4 5 6 6 4 & Blechnum Group
26. Veins of sterile leaflets anastomosing.
POmiNhizomercteepinge. Gorsiventraly 1) ee ba) ae Lomariopsis Group
PO ARNIZOMeZerech, enotaGorsiyentralie. se eebuey 2) se aa Tectaria Group
25. Sporangia not acrostichoid.
30. Rachis grooved, groove open to admit branches or midribs of leaflets.
Dryopteris Group
XII
General key No. 2 to Pteropsida
30. Rachis not grooved, its wings (if any) confluent with edges of lamina. Tectaria Group
6. Stipe containing one vascular bundle, or two at the base joining upwards to form a single strand of
various shape.
31. Rhizome-apex and bases of stipes hairy or apparently glabrous, not scaly.
32. Fronds repeatedly pseudo-dichotomous, a dormant apex between pairs of branches.
Gleicheniaceae
32. Fronds otherwise branched.
33. Apices of some or all primary rachis-branches dormant.
34. Rachis climbing and twining; sporangia solitary : 2 tS £ Schizaeaceae
34. Rachis trailing, not twining; several sporangia in a sorus, indusiate 3 0R Matoniaceae
33. Apices of primary rachis-branches not dormant (apex of main rachis sometimes periodically
dormant).
35. Sori at ends of veins, either in cups or protected by two equal or subequal flaps.
Roary serus;slamina one Cell thick sect ls ss eee Hymenophyllaceae
36. Not filmy, lamina thicker.
37. Massive rootstock, prostrate or erect and trunk-like, densely covered with long hairs.
Dicksonia Group
Bversiender ‘creepins rhizome; hairs, short) 794. #2 Ss) 2 SoS Dennstaedtia Group
35. Sori otherwise.
38. Bases of stipes bearing prominent aerophores; fronds dimorphous, fertile with narrow
acrostichoid pinnae . . = tee... <, .Plagiogyria
38. Bases of stipes lacking acrophores; fertile fronds otherwise:
39. Sporangia large, round, lacking an annulus, on deeply dissected pinnae Osmundaceae
39. Sporangia otherwise.
40. Sporangia variously arranged on lower surface of fronds, usually along the veins, or in a
submarginal groove, not indusiate.
41. Sori elongate . . . id re ns eee oe Adiantum Group
41. Sori small and round, or - acrostichoid. cqeg tk Reece oe ee ee Se OLY POCIAceae
40. Sporangia arranged otherwise.
42. Annulus almost apical; fronds simple or branched dichotomously; sporangia on special
appendages ts WON Ae CA NS eo eee Schizaeaceae
42. Annulus and frond otherwise.
43. Fronds pedately branched; sori superficial, indusiate. . . . . Matoniaceae
43. Fronds pinnately branched; sori at ends of veins, indusiate or ‘not.
Dennstaedtia Group
31. Rhizome-apex and bases of stipes scaly.
44. Sporangia on surface of reflexed marginal lobes . . . ... . Adiantum Group
44. Sporangia not on such lobes.
45. Sori elongate along the veins of the lower surface.
46. Sori indusiate.
47. Scales clathrate; the 2 vascular bundles at base of stipe uniting upwards to an X-shape.
Asplenium Group
47. Scales not clathrate; the 2 vascular bundles at base of stipe uniting upwards to a U-shape.
48. Unicellular slender hairs lacking . . . Ba ae Oh ee ee Athyrium Group
48. Unicellular slender hairs on frond and on scales A ee ee Thelypteris Group
46. Sori not indusiate.
49. Fronds simple; sori in grooves (or rarely superficial); epiphytes or rock-plants.
Vittaria Group
49. Fronds usually not simple; sori not in grooves; not epiphytes or rock-plants.
50. Sori not occupying whole length of veins.
51. Lower surface of lamina not covered with powder . . . . . Athyrium Group
51. Lower surface of lamina covered with powder . . .. . . Adiantum Group
50. Sori occupying whole length of veins.
52. Slender unicellular hairs lacking on lamina and scales . . . . Adiantum Group
52. Slender unicellular hairs present on lamina and scales . . . . Thelypteris Group
45. Sori not elongate along the veins of the lower surface (in some cases elongate along a sub-
marginal vein).
53. Sori in marginal or submarginal grooves, or superficial and forming continuous lines parallel
to midrib.
SA: aScalesuclathrate: oS p56 se ees see wc. eae eS) hee | te i ariagGroap
545,Scales: notuclathrate. 4. tim At. fee Pace ue lee, ta ee PG Lanne
53. Sori otherwise.
55. Sori at ends of veins (uniting several veins or not) protected by reflexed edge of lamina.
56. Rachis grooved on upper surface, groove opening to admit groove of midrib of pinna.
Pteris Group
XI
FLORA MALESIANA
XIV
56. Rachis not grooved in this way . . . oo. . . . . . . Adiantum Group
55. Sori not at ends of veins or not so protected.
57. Lower surface of frond more or less covered with white or yellow powder.
Adiantum Group
57. Lower surface of frond not so covered.
58. Sori submarginal, at ends of veins, in small projecting cups, or protected by an indusium
attached on the side remote from the margin.
59. Sori in’ cups: projecting sirom) margin) 9 ee Dennstaedtia Group
59. Sori not in such cups.
60. Sori always solitary at ends of single veins; scales of rhizome bearing papillae on marginal
cells; rhizome not protostelic. . . . . Davallia Group
60. Sori often uniting ends of several veins; ‘scales lacking papillae; rhizome protostelic.
Lindsaea Group
58. Sori otherwise.
61. Main rachis of fronds periodic in growth; thicket-forming ferns . Gleicheniaceae
61. Main rachis of fronds continuous in growth; not thicket-forming ferns.
62. Small epiphytes (occasionally on rocks); sori superficial or immersed in pockets in
substance of the lamina, not indusiate . . . . ... . Grammitidaceae
62. Not epiphytes; sori in most cases indusiate.
63. Hairs on fronds unicellular (in rare cases very long hairs may consist of more than
one cell).
64. Scales confined to swollen basal part of stipe; no superficial hairs or glands on scales.
Tectaria Group
64. Scales not so confined, bearing superficial unicellular hairs or glands.
Thelypteris Group
63. Hairs on fronds multicellular, cells short . . . . .. . Athyrium Group
ace YS TO THE*GENERA)» OF PTEROPSIDA
OPHIOGLOSSACEAE
1. Sporangia in two rows, embedded in an almost terete spike . . . . . . Ophioglossum
1. Sporangia on branches of the fertile segment of a frond.
2. Fertile segment of frond compact, with many short branches; sterile segment tripartite, each part
with few leaflets . . . . Helminthostachys
2. Fertile segment of frond amply branched with spreading branches: sterile segment (in Malaysian
species) pinnately branched with many small divisions Suchet? twice sine sboayciium
MARATTIACEAE
1. Sporangia in each group along the veins near margins of leaflets; veins free.
PEOraneia ih ‘cach eroup) united laterallysymereeres Smee) Se Rll ieee Marattia
2. Sporangia in each group almost free.
3. Fronds bipinnate; sporangia in each group commonly 8-12, less often to 20 . Angiopteris
3. Fronds simply pinnate; sporangia in each group much more numerous . . Macroglossum
1. Sporangia in each group united laterally to form a small circle, the circular groups scattered over the
Rieiacesnveins; anastomosing’ . 4. 5. pe ey ed ers Rea ee Christensenia
OSMUNDACEAE
1. Fertile pinnae quite different from sterile, lacking a green lamina; lamina‘of sterile pinnae not trans-
lucent BIG og . . Osmunda
1. Fertile pinnae not different from sterile in “shape; lamina very thin and ‘translucent Leptopteris
SCHIZAEACEAE
1. Fronds of adult plant dichotomously branched or simple, the fertile lobes at the end of a frond
or of its branches. . . so Sette
1. Fronds of adult plants scandent (rachis twining) with very short primary “rachis- branches bearing
featvasccondary, (branches 5 oe ace oe he sae! gee Be ee a re Lygodium
GLEICHENIACEAE
1. Rhizome and resting apices of fronds bearing multicellular hairs which are branched near the base;
veins always at least twice forked . . . ; Dicranopteris
1. Rhizome and resting apices of fronds bearing flat scales, rest of fronds usually also stellate hairs
(one cell to each ray); veins simple or forked once ®° oie beetle alee Gleichenia
HYMENOPHYLLACEAE*
1. Lips of indusium always well developed, broader and longer than the hollow basal part; receptacle
usually much shorter than the lips of the indusium (an elongate receptacle in Meringium PRESL).
Hymenophyllum
1. Indusium tubular or trumpet-shaped, sometimes with two-lipped mouth; receptacle usually elongating
considerably vand» protruding iromvold soni ere. - eeeecne ene ee Trichemanes
MATONIACEAE
1. Fronds erect, branching pedate-dichotomous . . . . Matonia
1. Fronds drooping, elongate, pinnately branched with apices “of some branches dormant.
Phanerosorus
CYATHAEACEAE
Note. Probably all species will be united in the genus Cyathea (see HoLTtuM, Kew Bulletin 1957,
pp. 41-45).
POLYPODIACEAE
1. Rhizome bristly or hairy, stipes not jointed to it; terrestrial.
2. Fronds dimorphous; fertile fronds narrow, acrostichoid . . .... . Cheiropleuria
2. Fronds uniform; sori small and round . a sprtiveast i, dead, 2a) cae Dipteris
1. Rhizome scaly, stipes usually jointed to it; mostly epiphytes.
3. Young sori protected by umbrella-shaped paraphyses.
4. Sporangia acrostichoid on a constricted apical part of the frond . . . . . Belvisia
4. Sporangia in separate sori, or whole fertile frond narrow and acrostichoid.
5. Sori somewhat elongate, close to and parallel with edge of lamina . . . Paragramma
* For a much fuller subdivision of this family, see COPELAND, Genera Filicum (1947) pp. 31-44.
XV
FLORA MALESIANA [ser. Il, vol} i
5. Sori otherwise.
6. Fronds small, fertile ones (or fertile parts) contracted. . . . . Lemmaphyllum
65 Fronds larger; not dimorphousis. 26 cae oo) ee eee _ Lepisorus and Neocheiropteris
3. Young sori not so protected.
7. Fronds bearing stellate hairs.
8. Fronds simple, entire.
9. Sori continuous along margin of narrow fertile frond or between margin and midrib.
Drymoglossum
9. Sori separate, round, variously distributed . . . Pyrrosia
8. Fronds branched dichotomously, sporangia acrostichoid. on - part of surface ‘of fertile frond.
Platycerium
7. Fronds not bearing stellate hairs.
. ier at the ends of free veins (veins conspicuous).
. All veins free ae 3S eye! 6 Ge le ano AOA heel > i ee Thylacopteris
. Some veins anastomosing badltis . . . . . Goniophlebium
1 ‘Sori nat at ends of free veins (veins often inconspicuous).
12. Veins anastomosing to form only one series of areoles along costae . . Polypodiopsis
12. Veins more copiously anastomosing.
13. Rhizome swollen and ant-inhabited.
14. Rhizome not scaly; sori on distinct small lobes Sot Wo ee ee Lecanopteris
14. Rhizome copiously scaly; sori not on distinct lobes . . . . . . + Myrmecophila
13. Rhizome not ant-inhabited.
15. Fertile fronds (or fertile parts) acrostichoid, usually much reduced as compared with sterile.
16. Fronds pinnate or pinnatifid, lower pinnae or lobes always sterile.
I75 Bronds pinnate; sterile pmnae jointed toyrachis 4 =< = = =. © « Photinopteris
17. Fronds pinnatifid . . eT ro cet en eee ee igniniiacaaTe
16. Fertile fronds wholly fertile.
i Teron Sinonyube (inl 5 75 6 6 = 6 « oa 6 6 o oa -<% Christiopteris
18. Fronds simple.
19. Fronds thin, all veins visible.
20. Fronds c. 10 cm long; terrestrial ‘ Sg eke. ee Dendroglossa
20. Fronds c. 30 cm long; rock- plants or epiphytes » « « « » « » SMeptochilus
19. Fronds coriaceous, at most main veins visible.
oh Sterile fronds narrow. NORIO?) Fee FF Oleandropsis
Sterile fronds circular to broadly lanceolate.
o Sterile fronds circular ; SEER SSW. (Eire . Pycnoloma
22. Sterile fronds longer than wide, apex acute eae Te te Grammatopteridium
15. Fertile fronds (or fertile parts) not acrostichoid.
23. Fronds pinnate or pinnatifid, pinnae or segments jointed to rachis.
24. Separate short humus-collecting fronds present . . . . . . . +. . Drynaria
24. Separate humus-collecting fronds lacking.
25. Bases of fronds broad, humus-collecting.
26; Eronds) on ‘special! branches of rhizomes. (2) 2:02) 93 41-05-55 Belhaven
26:) Kronds; not ionvspecial branches) = 95 9.) 4 = 4.5 4 © one Aglaomorpha
25. Bases of fronds not humus-collecting.
27. Fronds dimorphous En ey, Oe We he olga claniate | AR in oe Da Sule iS Holostachyum
27. Fronds not dimorphous . . Shoe Clee Arthromeris
23. Fronds simple or pinnatifid, segments not jointed ‘to rachis.
25. separate humus-collecting fronds present... = 2 9. 4 «4 ee Dendroconche
28. Separate humus-collecting fronds lacking.
29. Sori elongate, oblique, parallel to main lateral veins.
30. Fronds thin, all veins visible . . . 7 SS) eee Colysis
30. Fronds fleshy or leathery, at most main veins visible.
31. Main lateral veins distinct; scales not clathrate . . . . .. . Selliguea
31. Main lateral veins not distinct; scales clathrate . . . . . . . Loxogramme
29. Sori otherwise.
S25 Sorivelongate, near margins ol thim fronds eyes) re) | enenennne Diblemma
32. Sori otherwise.
33. Fronds very narrow, not dimorphous; sori elongate, close to costa . Holcosorus
33. Fronds otherwise; sori round.
34. Fronds always coriaceous; edges usually notched; scales not clathrate.
Crypsinus
34. Fronds not always coriaceous; edges not notched; scales clathrate Microsorium
XVI
Dec. 1959] ; Keys to the genera’ of Pteropsida
GRAMMITIDACEAE
Fronds simple and entire or with somewhat undulate rene
a Sori round or obliquely elliptical, separate . . anes Grammitis
2. Sori elongate parallel to costa and edge, or confluent _ near apex of frond.
3. Sori in grooves between margin and costa.
4. Fronds almost terete apart from widened apical fertile part . . . . . Nematopteris
4. Fronds of about uniform width throughout . . . . . . . . . +. ~ Seleroglossum
3. Sori more or less confluent near apex, not in grooves . . . . . . . QOreogrammitis
1. Fronds pinnately lobed to bipinnately lobed.
5. One vein (sometimes forked) and one sorus on each lamina-lobe.
6. Fertile lobes flat.
feesoOnl supericial . 7... Ee ee ets es ga ee a ONLCTAS
7. Sori deeply sunk in submarginal. pockets <0 Sea a ae ee ea ee Prosaptia
6. Fertile lobes partly reflexed, protecting the sori.
8. Lobes thin, only basiscopic margin reflexed . . . . . Calymmodon
8. Lobes coriaceous, both margins reflexed towards apex and. enclosing the sorus Acrosorus
5. Veins in lobes pinnately branched, each lobe with more than one sorus.
9. Sori superficial, or sunk in cavities perpendicular to the surface . . . . Ctenopteris
9. Sori deeply sunk in submarginal pockets not perpendicular to the surface Oe. Prosaptia
REMAINING GENERA OF FERNS
ADIANTUM Group
1. Water plants; sporangia borne singly and protected by reflexed edges of the lamina.
Ceratopteris
1. Land plants; sporangia in sori or more or less acrostichoid.
2. Rigid hairs or bristles on rhizome and bases of stipes.
3. Fronds bipinnate, leaflets more or less lobed . . . . Sp Ss senkiseks me lesa ECL OSOEA
3. Fronds simple or simply pinnate, or palmate with entire leaflets.
4. Veins anastomosing only near edge of lamina; fronds simple or ae paraphyses (if present)
with distinctive apical cell opr . . . Syngramma
4. Veins anastomosing throughout lamina; ‘fronds pinnate, trilobed or simple: paraphyses always
abundant, hair-like, of many cells, apical one not different from the rest.
5. Sori broad, elongate, parallel to costa and edge, or more or less spreading along the veins.
Taenitis
5. Sori in submarginal grooves, edges of grooves of equal thickness (Schizoloma sensu COPEL.).
Schizolepton
2. Scales on rhizome and bases of stipes.
€. Lower surface of fronds covered with white or yellow powder.
7. Sori along whole length of veins on lower surface . . Sa ty ohn oc , .ityRogramma
7. Sori at ends of veins, protected Dy reflexed lobes of margin sal ca Lesiceeis mip Cnelanees
6. Lower surface of fronds not covered with such powder.
8. Sori elongate (continuous or broken) along margins of lamina and protected by reflexed margin.
9. Fronds much dissected, the ultimate lamina-lobes small and connected bya narrow wing.
10. Fertile lobes much broader than sterile, the whole of each margin bearing a broad thin reflexed
indusium. . aes of Onyehrom
10. Fertile lobes not much broader than sterile, reflexed margin not ‘continuous along larger
fertile lobes nor very broad . . . . . Cheilanthes
9. Fronds, if much branched, having quite distinct leaflets not joined by a wing.
11. Fronds simple and lobed, or deeply pinnatifid; sori quite continuous along edge Doryopteris
11. Fronds branched with distinct stalked leaflets EAL tad tare eer teed te llaea
8. Sori otherwise.
12. Sori elongate along all or most of the veins.
13. Fronds simple . . a ee el Re pe Re ee een ee EICTNIONINEES
13. Fronds pinnate or bipinnate Re ee en oe ree a ee OF ey pe a OT rae ne
12. Sori otherwise.
14. Sori on lower surface, spreading a little along the veins . . . . . . Anogramma
14. Sori at ends of veins.
15. Sori on surface of reflexed marginal flaps . . . . Adiantum
15. Sori at ends of veins, sometimes more or less protected by reflexed marginal flaps Notholaena
ASPLENIUM Group
1. Sori short, on one-veined ultimate segments of much-divided fronds . » 0 aatLoxoscaphe
1. Sori distinctly elongate along the veins; segments of frond usually with more than one vein..
XVII
FLORA MALESIANA [ser. II, vol. 1
2. Sori on adjacent veins ae towards each other, a raised line remaining between the sori when
they are ripe . . Diplora
2. Sori all facing the s same way except | ina 1 few species which have simple fronds and then the raised
lines between, adjacentysonmlacking. "2-959 = ee
ATHYRIUM Group
1. Veins free.
2. Groove of rachis open to admit grooves of branch rachises and midribs of leaflets.
3. At least some sori elongate along veins.
4. Sori not indusiate; fleshy outgrowths present at bases of pinnae on upper surface.
Cornopteris
4. Sori indusiate; no such outgrowths.
5. Double sori (on both sides of a vein) not connected at their distal ends. . Diplazium
5. Double sori (at least some of them) connected at their distal ends; usually some reniform and
J-shaped sori present Se a ie eee Tinea AAA Eee 8 ITT
3. Sori not elongate along veins. . 5 os me | ae OUST OMLCEIS
2. Groove of rachis not open to admit grooves of branches pea enc” ae Dryoathyrium
1. Veins anastomosing.
6. Sori almost circular, indusium reniform . . . . . - . . . . . +. Anisocampium
6. Sori elongate along veins.
7. Veins anastomosing regularly, about as in Cyclosorus; sori on nearly all veins of fertile frond.
Callipteris
7. Veins anastomosing less regularly, the areoles adjacent to costae large; sori never so abundant.
8. Indusium rather firm, its edge reflexed at maturity . . oes Diplazium
8. Indusium thin, sausage-shaped, breaking at maturity, its edge not reflexed . . Diplaziopsis
BLECHNUM Group
1. Sori not acrostichoid, or if apparently so, a thin indusium present all along edge of fertile leaflets.
2. Sori continuous along each side of the costa of a leaflet, a sterile lamina present beyond the sorus
or not = -; Phebe Blechnum
2. Sori not continuous throughout length of a leaflet: one sorus ‘to ‘each areole along the costa.
3. Small ferns, simply pinnate; edges of pinnae sharply toothed . . 2 ee One
3. Large ferns, at least bipinnatifid; edges of pinnae not sharply toothed . . . Woodwardia
. Sori acrostichoid (no thin reflexed indusium on fertile leaflets) . . . . . . +. Brainea
DAVALLIA Group
1. Rhizome bearing hairs and scales; marginal cells of scales bearing papillose outgrowths Leucostegia
1. Rhizome bearing scales only; marginal cells of scales not bearing papillose outgrowths.
2. Fronds simple.
3. Sori elongate along the margin, protected by continuous indusia . . . . ._ . Parasorus
3. Sori singly at ends of veins.
4. Indusium attached at base only (or slightly above base) . . . . . . . +. Humata
4. Indusium attached along sides as well as base . . . . . . . +. +. ~~. + Seyphularia
2. Fronds pinnately branched.
5. Fronds more or less copiously persistently hairy; lamina thin; indusium small, of various shapes.
Dayallodes
5. Fronds not hairy when mature.
6. Indusium attacheé along base and whole of sides.
7. Fronds trifoliate or simply pinnate with elongate narrow pinnae . . . . Scyphularia
7. Fronds more copiously branched . . . . Dayvallia
6. Indusium attached at base only, or also a little above the base, not along whole length of both sides.
8. Fronds much-branched, thin. . EE LO ON ES REY AUTESS LCE
8. Fronds of various form, lamina coriaceous or fleshy.
8: Scales: with lone*acicular tips 21'S SP Pe ee eee
Be seales tacking lone acicular tips "3. 2 Sw
DENNSTAEDTIA Group
1. Sori in cups at ends of veins.
Paleai7Omemssicndcr creeping, Hairy” 2 ese ge) ve Ge ee ee a ee ts Dennstaedtia
2. Rhizome stout, erect, scaly. . . a ret a re ae Orthiopteris
1. Sori at ends of veins, variously protected, ‘not in cups.
3. Sori lacking protection; glandular hairs mixed with sporangia . . Monachosorum
3. Sori with indusium or covered when young by reflexed lobe of edge of lamina; no glandular hairs
with sporangia.
4. Sori distinctly superficial, each at the end of a single vein and protected by an indusium which is
XVIII
Dec. 1959] Keys to the genera of Pteropsida
pocket-shaped (attached by base and sides), opening towards margin. . . . M@éicrolepia
4. Sori more or less protected by reflexed edge (or lobes of edge) of lamina, with or without an inner
indusium also.
5. Sori almost continuous along edges of lamina.
6. Veins anastomosing; NO’ Inner Indust yaes a) Ga Coslce)l) a/oc3 Oe) cee aeeishiopteris
6. Veins free; inner indusium present es , ‘ Pteridium
5. Sori singly at ends of veins, or uniting a few v veins, not continuous along all edges of lamina.
PRINTING USIiINT fe O5s 5. Ge sie eee REE ise ole Soa 27) |S Se Hypolepis
REET CINOUSINM Pleseht, .. .. .o iP eMC eel a ye Gee) a ee Paesia
DICKSONIA Group
1. Fertile leaflets distinctly narrower than sterile.
2. Fertile pinnules deeply lobed; massive tree-ferns . . . . . . . «.. . Dicksonia
2. Fertile pinnules not lobed; stem short, creeping : SSVI Ona Wed FAal?: Cystodium
1. Fertile leaflets little different from sterile in size and shape.
3. Pinnules deeply lobed almost at right angles to costa, not pinnate ft Leste} Cibotium
3. Pinnules pinnate, tertiary leaflets small, very oblique and lobed. . . . . . . Culcita
DRYOPTERIS Group
1. Fronds hairy as well as scaly on costae, costules and rachises; hairs septate, coarse, different from
the scales.
2. Sorus when young spherical and quite enclosed by indusium which when ripe breaks to expose
sporangia.
BMESGOUIBONEGIStiINCL StalIkKS = \.) 3 oo « be Ce ee ee os ee ee | erAanema
BESCIMINSESSIGGe tt gee) Ecc ne roa 2 eee cy Oe eee Sy es sek aco Diacalpe
2. Sorus otherwise; indusium attached on basiscopic side of sorus.
4. Hairs sparse; seales broad’ and thin’ %.) *..'-.- “2.4 | a bee, ce. oe, ReIOp ONES
4. Hairs copious; scales thick, elongate . . salle Stenolepia
. Fronds scaly; no hairs as distinct from scales (some ‘scales may be narrow and almost hair-like).
a Pinnules jointed to rachis; sori longer than wide . . . . . . . +. . Didymochlaena
5. Pinnules not jointed to rachis; sori not elongate.
6. Several vascular bundles in stipe.
7. Basal acroscopic leaflet or lobe of middle pinnae distinctly nearer to main rachis than basal
basiscopic leaflet or lobe; fronds never simply pinnate.
8. Fronds elongate, basal pinnae not much enlarged on basiscopic side; rachis often bearing a bud.
Polystichum
8. Fronds broadly deltoid, basal pinnae much enlarged on basiscopic side; rachis lacking buds.
Polystichopsis
7. Basal acroscopic leaflet or lobe of middle pinnae not distinctly nearer to main rachis than basis-
copic leaflet or lobe; or fronds simply pinnate throughout.
9. Indusium peltate; teeth on edges of lamina often with stiff slender points. . Polystichum
9. Indusium reniform; teeth not so produced . . ... . . Dryopteris
faiwo vascular bundles in ibase of stipe) =) )6 "seh alesis. e- pee Gymnocarpium and Currania
LINDSAEA Group
1. Fronds simply pinnate, veins free, pinnae numerous, spreading, jointed to rachis. . Isoloma
1. Fronds otherwise.
2. Pinnae or lobes elongate, bearing many sori, each at the end of a single vein; lamina rather thick.
Tapeinidium
2. Pinnae or lobes, if elongate, bearing a sorus connecting the ends of several veins; lamina thinner.
3. Fronds finely divided, the ultimate divisions small and connected with each other by wings; sori
solitary at apices of ultimate divisions . . . . . Sphenomeris
3. Fronds variously branched; leaflets distinct, each with several sori or a more or less continuous
miieeinal SOruUS. ‘.. 5. .<<imwlt: wie doa bees viedhsg) slau: sheet: o-1le, Paisaes
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.
Beaermsireed .. i+ ce cal So» we) ip be ey we) Go es (dy 3) -- WS Pi eee
3 Veins anastomosine... . = G% 6 ce ss » . = ceteche Ss se WR EM ole
1. Rhizome high-climbing.
4. Veins free.
FLORA MALESIANA [ser. I, vos
5. Pinnate or bipinnate, all pinnae and pinnules jointed at base (except on some bathyphylls).
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 ey . . Lomariopsis
7. Fertile pinnae bearing separate exindusiate sori on small lobes at ends of veins.
Thysanosoria
4sVeins anastoniosing: Sey =e 2 eet ee oe ey, ee ee ee en ee Lomagramma
NEPHROLEPIS Group
1. Fronds pinnate, pinnae jointed to rachis.
2. Stipes jointed to rhizome or to outgrowths from rhizome . .. .. . Arthropteris
De Stipes not jomted ‘to crhizomes. Pkt bees 2. Bees Hor. per, Grow, IR Nephrolepis
ieebrondssimple:..) 2) sv". LRRD NG eae es Pe.” a? oe Oleandra
PLAGIOGYRIA Group
Single’ zeenustaiw hes. ae he eh RO Re ot) Se a? Boe oes
PTERIS Group
1. Sori marginal or submarginal, more or less elongate, protected by a reflexed indusium.
2. Rhizome scandent ee tee OM Te a OR SEE Thee RS Oe Lepidocaulon
2. Rhizome not scandent.
3. Sorus along one edge of ultimate lobes of lamina . . . . . . . +. +. + #WHemipteris
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 BF Nac 08, lentes LUO ae Arcypteris
cs Vascular bundles in stipes in a simple ring; veins free fe BESS (0 SoS ee Pteridrys
No tooth present in sinuses between lobes of lamina.
2 PinndewomtedatOmmdachis.. G8 & .. 5 “a s8ls 6 soe | oe) OV cloneinn
4. Pinnae not jointed to rachis.
5. Fertile leaflets acrostichoid and very much contracted as compared with sterile ones.
G> Veins: free=ssterile’ frond? much divided". 92). © 5A. 19.1. OG Oe eee
6. Veins anastomosing.
7. Fronds deltoid, the basal basiscopic lobe of basal pinnae largest . . . . Stenosemia
7. Fronds otherwise, basal basiscopic lobes of basal pinnae not largest.
8. Fronds small, trifoliate; apical leaflet largest, commonly 5 cm long . . . Quercifilix
8. Fronds larger and in most cases more divided.
9. Sterile fronds simple to deeply pinnatifid, not truly pinnate . . . . Hemigramma
9. Sterile fronds with at least one pair of free pinnae, usually several pairs Heterogonium
5. Fertile leaflets not acrostichoid.
10. Two vascular bundles in base of stipe, uniting upwards to form a single bundle; scales confined
to swollen bases of stipes . . . . . Hypodematium
10. More than two vascular bundles throughout ‘length of stipe; ‘scales not so confined.
11. Veins free.
12. Basal basiscopic vein of a vein-group springing from the costule.
13. Indusia peltate Paes : TED AEP ey ae ere! Bee Dryopolystichum
13. Indusia reniform or absent.
14. Fronds usually much longer than wide; basiscopic margin of lamina-lobes not thickened.
Ctenitis
XX
Dec. 1959] Keys to the genera of Pteropsida
14. Fronds usually about as long as wide; basiscopic margin of lamina-lobes thickened.
Lastreopsis
12. Basal basiscopic vein of a vein-group springing directly from the costa.
15. Basal pinnae with basal basiscopic lobes longest... . . Tectaria
15. Basal pinnae with basal basiscopic lobes or pinnules shorter ‘than middle ones.
Heterogonium
11. Veins anastomosing.
16. Sori large, indusiate, terminal on a free vein, the receptacle elongate, fertile fronds narrow.
Luerssenia
16. Sori various, indusiate or not, on free or netted veins; if indusiate, the receptacle not elongate,
or fertile fronds not contracted as compared with sterile.
17. Basal pinnae deeply lobed, basal basiscopic lobe not largest; few free veins in areoles.
Heterogonium
17. Basal pinnae lobed or not; if lobed, basal basiscopic lobe largest; many free veins in areoles.
Tectaria
THELYPTERIS Group
ieeEronassbearing, many buds On Tachis; 50 - eesoa ce Weiner e em scl Ampelopteris
1. Fronds lacking buds.
2. Veins free (in some cases basal veins of adjacent groups just meet at the sinus). Thelypteris
2. Veins anastomosing.
3. Sori elongate along veins.
4. No indusia.
5. Venation as in Cyclosorus; veins from adjacent costules uniting to form a single excurrent vein.
Stegnogramma
5. Venation irregularly SES COS: often with additional enclosed areoles . . Dictyocline
4. Indusia present. . : Sphaerostephanos
3. Sori not elongate, or slightly so if without indusia \(Abacepieris FéE, Haplodictyune PRESL).
Cyclosorus
VITTARIA Group
1. Frond very small, with one vein only; sorus near apex, along the vein ee Monogramma
1. Frond with lateral veins as well as a main vein.
2. Frond linear, sori one or more, on eas free branches, which run close to and parallel with
main vein . . olen el (EATS b.8Vaginularia
2. Fronds otherwise; veins anastomosing ‘at least near “the margin.
3. Sori linear, in marginal grooves or superficial and parallel to margin; veins forming one series of
areoles, anastomosing only near the margin aA . . Wittaria
3. Sori along veins, variously disposed; veins anastomosing “copiously throughout the lamina.
Antrophyum
MARSILEACEAE
Solemeculisay arise EN? PRAVEEN ee ans Gs Jee ee) oe ee oe ne cee Vearsilea
SALVINIACEAE
1. Leaves less than 1 mm long, bilobed, with one lobe submerged . . . ete Azolla
1. Leaves at least 1 cm long, simple, floating with upper surface fully exposed . . . Salvinia
7. BIBLIOGRAPHY
A list of books and papers dealing with the taxonomy of Malaysian ferns, published
subsequent to Christensen, Index Filicum, Suppl. 3 (1934)
Aston, A. H. G., Fern notes II (New Guinea
ferns). J. Bot. 77 (1939) 288-290.
—, Undescribed ferns from New Guinea. Nova
Guinea n.s. 4 (1940) 109-112, pl. 4-10; also
in J. Bot. 78 (1940) 225-229, no plates.
—, Some undescribed ferns from New Guinea and
Ambon. Nova Guinea n.s. 7 (1956) 1-3.
Aston, A. H. G. & R. E. Ho_trum, Notes on
taxonomy and nomenclature in the genus
Lygodium. Reinwardtia 5 (1959) 11-22.
BACKER, C. A., Fam. Hymenophyllaceae—Equise-
taceae. Beknopte Flora van Java (em.ed.)
pts 1-2 (1940) fam. I-XV.
BACKER, C. A. & O. PostHuUMUS, Varenflora voor
Java. Buitenzorg, 1939.
CHING, R. C., On the genus Onychium Kaulf. from
the Far Orient. Lingn. Se. J. 13 (1934)
493-501.
—, A revision of the compound leaved Polysticha
and other related species in the continental Asia
including Japan and Formosa. Sinensia 5
(1934) 23-91, 18 pl., 2 fig.
—, On the genus Hypodematium Kunze. Sunyat-
senia 3 (1935) 3-15.
—, On the genus Pyrrosia Mirbel. Bull. Chin.
Bot. Soc. 1 (1935) 36-72.
—, On the genera Stegnogramma Bl. and Lepto-
gramma J.Sm. Sinensia 7 (1936) 89-112.
—, Arevisionof the Chinese and Sikkim-Himalayan
Dryopteris with reference to some species of
neighbouring regions, 1. Bull. Fan Mem. Inst.
Biol. 6 (1936) 237-352.
—, Ditto, 2-5. Bull. Fan Mem. Inst. Biol. 8
(1938) 157-268.
—, Ditto, 6-9, l.c. 275-334, pl. 6-7.
—, Ditto, 10. l.c. 363-507.
—, The studies of Chinese Ferns XXXV. op.
cit. 11 (1941) 79-82.
—, On natural classification of the family “‘Poly-
podiaceae’’. Sunyatsenia 5 (1940) 201-268,
table.
—, On the genus Gleichenia Smith. Sunyatsenia
5 (1940) 269-288.
—, The Studies of Chinese Ferns, XX XI. Aleurito-
pteris Fée. Hong Kong Nat. 10 (1941) 194-204.
.
CHRISTENSEN, C., New and noteworthy Papuan ferns.
Brittonia 2 (1937) 265-317.
—, Taxonomic Fern-studies, III. Revision of the
genera and species of ferns described by
A. J. Cavanilles. Dansk Bot. Ark. 9, no 3
(1937) 1-32.
—, Ditto, IV. Revision of the Bornean and New
Guinean ferns collected by O. Beccari and
described by V. Cesati and J. G. Baker. l.c.
33-52.
XXII
—, Ditto, V. Descriptions of 36 new species of
ferns. l.c. 53-73, pl. 1-6.
—, Filicinae (Ch. XX) in Verdoorn, Manual of
Pteridology. The Hague, 1938.
CHRISTENSEN, C. & R. C. CHING, Pteridrys, a new
fern genus from tropical Asia. Bull. Fan Mem.
Inst. Biol. 5 (1934) 125-148, pl. 11-20.
CLAUSEN, R. T., A Monograph of the Ophioglos-
saceae. Mem. Torrey Bot. Cl. 19, no 2
(1938) 1-177, fig. 1-33.
COPELAND, E. B., New or interesting Philippine
ferns, VIII. Philip. J. Sc. 56 (1935) 97-110,
pl. 1-14.
—, Additional ferns of Kinabalu. Philip. J. Sc. 56
(1935) 471-483, pl. 1-10.
—, The Philippine ferns collected by J. B. Steere.
Philip. J. Sc. 60 (1936) 19-25.
—, Solomon Island ferns. Philip. J. Sc. 60 (1936)
99-117, pl. 1-23.
—, Hymenophyllum. Philip. J. Sc. 64 (1937)
1-188, pl. 1-89.
—, Genera Hymenophyllacearum. Philip. J. Sc.
67 (1938) 1-110, pl. 1-11.
—, Fern evolution in Antarctica. Philip. J. Sc.
70 (1939) 157-188, 2 maps.
—, Oleandrid ferns (Davalliaceae) of New Guinea.
Philip. J. Sc. 73 (1940) 345-357, pl. 1-10.
—, Notes on Hymenophyllaceae. Philip. J. Sc.
73 (1941) 457-469, pl. 1-4.
—, Holttumia, genus novum. Philip. J. Sc. 74
(1941) 153-156, pl. 1.
—, Gleicheniaceae of New Guinea. Philip. J. Sc.
75 (1941) 347-361, pl. 1-6.
—, Miscellaneous ferns of New Guinea. Philip. J.
Sc. 76 (1941) 23-25.
—, Filicum novarum CXXVIII diagnoses. Un.
Cal. Publ. Bot. 18 (1942) 217-226.
—, Ferns of the Second Archbold Expedition to
New Guinea. J. Arn. Arb. 24 (1943) 440-443.
—, Genera Filicum. Waltham, Mass., 1947.
—, Cyathea in New Guinea. Philip. J. Sc. 77
(1947) 95-125, pl. 1-15.
—, Pteridaceae of New Guinea. Philip. J. Sc. 78
(1950) 5-41, pl. 1-6.
—, Aspleniaceae and Blechnaceae of New Guinea.
Philip. J. Sc. 78 (1950) 207-229, pl. 1-6.
—, Aspidiaceae of New Guinea. Philip. J. Sc. 78
(1951) 389-475, pl. 1-44.
—, Grammitis. Philip. J. Sc. 80 (1952) 93-276,
pl. 1-6.
—, New Philippine ferns, IX. Philip. J. Sc. 81
(1952) 1-47, pl. 1-26.
—, Grammitidaceae of New Guinea. Philip. J. Sc.
81 (1952) 81-119, pl. 1-10.
—, New Philippine ferns, X. Philip. J. Sc. 83
(1954) 97-101, pl. 1-5.
—, New Philippine ferns, XI. Philip. J. Sc. 84
(1955) 161-165, pl. 1-2.
Bibliography
Donk, M. A., Notes on Malesian ferns, I. On the
genus Lemmaphyllum Presi. Reinwardtia 2
(1954) 403-410.
Ho.ttum, R. E., The tree-ferns of the Malay
Peninsula. Gard. Bull. Str. Settl. 8 (1935)
293-320, pl. 29-36.
—, Notes on Malayan Ferns, with descriptions of
five new species. Gard. Bull. Str. Settl. 9
(1937) 119-138.
—, The genus Lomagramma. Gard. Bull. Str.
Settl. 9 (1937) 190-221, pl. 8-16.
, A redefinition of the genus Teratophyllum.
Gard. Bull. Str. Settl. 9 (1938) 355-362, pl.
28-30.
—, The fern genus Diplazium in the Malay Penin-
sula. Gard. Bull. Str. Settl. 11 (1940) 74-108,
fig. 1-6.
—, New species of vascular plants from the Malay
Peninsula (Filicales). Gard. Bull. Str. Settl.
11 (1947) 267-274, fig. 1.
—, A revised classification of Leptosporangiate
ferns. J. Linn. Soc. Bot. 53 (1947) 123-158.
—, The classification of ferns. Biol. Reviews 24
(1949) 267-296, 1 fig.
—, The fern-genus Heterogonium Presl. Sarawak
Mus. J. 5 (1949) 156-166, 2 fig.
—, The selection of type-species of some old
genera of ferns. Gard. Bull. Sing. 12 (1949)
303-306.
, Further notes on the fern-genus Heterogonium
Presl. Reinwardtia 1 (1950) 27-31.
—. The fern-genus Pleocnemia Presl. Reinwardtia
1 (1951) 171-189, 20 fig.
—, The fern-genus Arcypteris Underwood. Rein-
wardtia 1 (1951) 191-196, 3 fig.
—, Anew fern from Malaya. Gard. Bull. Sing. 14
(1953) 8.
—, A new Malayan Athyrium. Kew Bull. 1953,
545 (1954).
—, Arevised Flora of Malaya, II. Ferns of Malaya.
Singapore, 1954 (1955).
—, Some additional species of Heterogonium.
Reinwardtia 3 (1955) 269-274.
—, On the taxonomic subdivision of the Gleichenia-
ceae, with descriptions of new Malaysian species
and varieties. Reinwardtia 4 (1957) 257-280.
—, Morphology, growth-habit and classification
in the family Gleicheniaceae. Phytomorpholo-
gy 7 (1957) 168-184, 6 fig.
—, The scales of Cyatheaceae. Kew Bull. 1957,
41-45.
—, Notes on Malaysian Ferns, with descriptions
of a new genus and new species. Kew Bull.
1958, no 3 (1959) 447-455.
KRAMER, K. U., A new genus of Lindsaeoid ferns.
Acta Bot. Neerl. 6 (1957) 599-601, 1 fig.
Nakal, T., New classification of Gleicheniales,
etc. Bull. Nat. Sc. Mus. Tokyo no 29 (1950)
1-71.
PICHI-SERMOLLI, R. E. G., The nomenclature of
some fern genera. Webbia 9 (1953) 387-454.
—, Names and types of fern-genera, 1. Hymeno-
phyllopsidaceae, Loxsomaceae, Schizaeaceae.
Webbia 12 (1956) 1-40.
—, Ditto, 2. Angiopteridaceae, Marattiaceae,
Danaeaceae, Kaulfussiaceae, Matoniaceae,
Parkeriaceae, Adiantaceae. Webbia 12 (1957)
339-373.
PostHuMus, O., On some Malayan ferns. Proc.
Kon. Ak. Wet. A’dam 39 (1936) 823-827.
—, Malayan Fern Studies, I. The synonymy and
distribution of the Java-ferns. Verh. Kon. Ak.
Wet. A’dam sect. 2, 36, no 5 (1937) 1-67.
—, Ditto, II. On the fern-flora of Australia, and
its relation to that of neighbouring countries,
especially of the Malay Archipelago. Verh. Kon.
Ak. Wet. A’dam sect. 2, 37, no 5 (1938) 3-35.
—, Ditto, II. The ferns of the Lesser Sunda
Islands. Ann. Jard. Bot. Btzg, hors sér. (1943)
35-113.
REnSCcH, I., Farne und Bdrlappe der Sunda-Ex-
pedition Rensch. Hedwigia 74 (1934) 224-256,
pl. VII.
SELLING, O. H., Studies in the recent and fossil
species of Schizaea, with particular reference
to their spore characters. Act. Hort. Gotob. 16
(1944) 1-112, pl. 1-5.
—, Two new species of Schizaea and their affi-
nities. Svensk Bot. Tidskr. 40 (1946) 273-283,
fig. 1-11.
—, Further studies in Schizaea. Svensk Bot.
Tidskr. 41 (1947) 431-450, fig. 1-14.
WAGNER, W. H. & D. F. GreTHER, The Pterido-
phytes of the Admiralty Islands. Un. Cal.
Publ. Bot. 23 (1948) 17-110, pl. 5-25.
XXIII
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GLEICHENIACEAE (R. E. Holttum, Kew)
Rhizome relatively slender, creeping, protostelic (solenostelic only in Dicranopteris
pectinata (WILLD.) UND. of tropical America), in Stromatopteris bearing erect
irregularly dichotomous branches which bear the fronds, in all other cases bearing
fronds directly; young parts covered with peltate fringed scales (scales otherwise
in Stromatopteris) or branched hairs. Fronds unbranched in Stromatopteris, in
all other cases branched in fully developed plants, the main rachis bearing a
series of pairs of branches, its apex periodically dormant while each successive
pair of branches develops; each primary branch often bearing a pair of secondary
branches and a permanently dormant apex between them, the process some-
times repeated several times; ultimate branches either bipinnatifid or pinnatifid,
the lamina (whether of an ultimate branch, or leaflet of an ultimate branch) cut
almost to the costa; veins in lamina-segments pinnate, branches simple or forked,
free (in some cases apparently joining a thickened non-vascular margin). Sori
of 2-15 or more sporangia, attached to a small receptacle on the surface of a vein
(except in Stromatopteris, where each sorus is spread along part of both branches
of a forked vein), never at the end of a vein, all sporangia in one sorus developing
simultaneously; branched hairs or scales often present with sporangia but no
indusium. Sporangia with complete oblique annulus, dehiscing vertically, containing
c. 200-800 or more spores. Spores monolete or trilete, smooth, translucent, colourless.
Gametophyte (not known in Stromatopteris) at first cordate, then ribbon-like
with heavy midrib, finally branching at apex; rhizoids stiff, abundant, usually
reddish-brown; two-celled glandular hairs developed by many species in as-
sociation with archegonia and also on margin; antheridia comparatively large
and complex in structure (some more so than others); archegonia with long necks
(longest in Gleichenia subg. Gleichenia) directed towards apex of prothallus; no
cases of apogamy observed.
Distribution. Throughout the wetter parts of the tropics and subtropics, and in south temperate
regions. Three genera: Stromatopteris (monotypic, in New Caledonia), Gleichenia (3 subgenera,
c. 150 species, in the tropics mainly on mountains, at lower altitudes in southern temperate regions), and
Dicranopteris (2 subgenera, c. 10 species, mainly tropical, at low and moderate altitudes). Dicranopteris is
much more polymorphic in Malaysia than in any other part of the tropics. (The genus Platyzoma is
excluded from the family; see HoLTTUM in Kew Bulletin no 3, 1956, p. 551).
Fossils. The fragmentary nature of the earlier fossils ascribed by various authors to the family makes
judgment upon them difficult. The form and arrangement of sporangia is the best criterion; on this
basis paleozoic fossils named Oligocarpia have been assigned to Gleicheniaceae, but they do not always
show details of structure of the individual sporangia clearly. Some have been found on fronds of the
Pecopteris type (not unlike Gleichenia of today), some on the rather different Sphenopteris, but none
show pseudo-dichotomy of the kind now universal in the family. It may be that some of these fossils
represent members of the family before periodic growth and pseudo-dichotomy developed.
Gleichenia gracilis ZiGNo, from the Jurassic of northern Italy, has been more confidently referred to
the family by paleobotanists; but the main branches of the frond are not opposite, and the irregular
forking of the smaller branches looks more like true dichotomy than pseudo-dichotomy (judging from
ZIGNO’s figure); there are no sori.
Wealden fossils from Belgium show anatomical structure of rhizome and petioles as in living Glei-
chenia. In somewhat later Cretaceous rocks of various parts of the world, but especially in West Green-
land, are fossils which show every character (except scales, which have not been seen) of existing members
of Gleichenia subg. Diplopterygium (HEER, Flora Fossilis Arctica, II[) and some are very like subg.
Gleichenia, with more or less transitional stages between the two conditions which are not shown by
living members of the family. Some of them show permanent dormancy of a lateral branch of a frond,
but none show the condition either of Gleichenia subg. Mertensia or of Dicranopteris, which have what
I regard as the most highly developed forms of branching. Some Greenland fossils show many sporangia
in a sorus, a character not shown by any living member of the family having similar leaf-form,
and TuTIN (Ann. Bot. Lond. 46, 1932, p. 503-508) has described Gleicheniopsis with numerous small
(1)
2 FLORA MALESIANA [ser. Il, yoru
sporangia each containing less than 32 spores; the latter were not attached to any forked axis.
The only fossil known to me which corresponds in leaf-form with Dicranopteris is Gleichenia han-
tonensis WANKLYN from Eocene beds at Bournemouth (southern England). This was the subject of a
reconstruction, copied by other authors, by GARDNER and ETTINGSHAUSEN (British Eocene Flora, I
fig. 28) which (on the same frond as typical Dicranopteris ultimate branches) incorporated accessory
branches as in living D. linearis (not shown in the fossils) and also some curious hook-bearing leaflets
which are associated with the Dicranopteris fossil leaflets but nowhere attached to them. Disregarding
these unwarranted additions, Gleichenia hantonensis agrees with Dicranopteris in branching, venation
and sori (position, shape and number of sporangia); if hairs could be found, agreement with Dicranop-
teris would be complete. The Greenland and Bournemouth fossils offer evidence hard to reconcile with
COPELAND’S opinion that the family is of antarctic origin (Genera Filicum, p. 26).
Ecology. Allspecies are sun-ferns, and most form thickets, to which they are adapted by their creeping
rhizomes, and by the indefinite growth in length of the fronds (at least in the wet tropics). Periodic
dormancy of the apex of the main rachis permits periodic upward growth of the rachis-apex, unen-
cumbered by branches; it may thus pass between any other leaves or branches which may be above it,
and then develop its new pair of leafy branches in a fully exposed position while the apex itself is again
dormant. The most efficient members of the family from this standpoint are species of Dicranopteris
and of Gleichenia subg. Mertensia, because by repeated forking their lateral branch-systems produce a
spreading fan-shaped arrangement, but it is notable that in Dicranopteris a reversion to an effectively
pinnate form of branching has occurred, seen at its full development in D. speciosa (PR.) HOLTT.
Most members of the family are pioneer plants, establishing themselves on bare ground, sometimes
in fully exposed places, talus, earthslides, precipices, rocks (fig. 3), steep ridges (fig. 2, 10), often on
poor rocky or leached soils. Prothalli (which are often very abundant) need a little shelter, but a young
sporophyte can quickly spread by means of its rhizome, and Dicranopteris linearis, for example, can
grow in places where few other plants can establish themselves. A Dicranopteris thicket, once formed,
can persist for a long time (fig. 11, 13), unless tree seedlings have established themselves in it at an early
stage (shade of trees weakens or kills Dicranopteris plants unless they can climb); such thickets have
been greatly encouraged by man-made clearings of forest. On the edge of forest, some of the larger
members of the family can climb to a considerable height.
At high altitudes on mountains, where slower-growing alpine scrub occurs, smaller species of the
family are often abundant and form lower thickets. They may grow in association with Sphagnum,
or in dry rocky places. An important character is that their demands of mineral substances are small.
Vegetative morphology. Stromatopteris is peculiar in its erect rhizome-branches, in its scales
and hairs, and in having sori spread along both branches of a vein. NAKAI (Bull. Nat. Sc. Mus. Tokyo
n. 29, 1950) has proposed its separation as a distinct subfamily, and I consider this separation fully
justified. The genus does not occur in Malaysia. It is probably to be regarded as a reduced relic of an
otherwise extinct group.
As above noted (under Ecology), the fronds of other members of the family are all branched (except
in young or stunted plants), the branching being pinnate in plan, with dormancy, either periodic or
permanent, of apices of various orders. Fig. 1, 7, 12. The branch-patterns are in some cases complex,
and are characteristic in the different genera and subgenera (for a comparative account, see HOLTTUM,
Phytomorphology 7, 1957, 168-184). FILARsKy attempted a detailed analysis of these branch-patterns
(Ann. Hist.-Nat. Mus. Nat. Hungarici 20, 1923, 1-23; ibid. 21, 1924, 163-170) but the results seem to
me confused. He did not distinguish clearly between Dicranopteris and Gleichenia subgen. Mertensia
of the present treatment, nor between Platyzoma and Gleichenia subgen. Gleichenia. The form of branching
shown in his fig. 12 and 24 is one I have never seen (shorter branches on the same side at successive
unequal forkings).
The major divisions of the family are also distinguished by their dermal appendages. Those of Stro-
matopteris are quite peculiar (scales, only on rhizome, not peltate, and long simple hairs also on rhizome).
The rest of the family have either fringed peltate scales and stellate hairs with unicellular rays (Gleichenia)
or branched hairs of complex structure, true scales being absent (Dicranopteris, fig. 14).
The venation in Gleichenia is pinnate in each segment of the lamina, lateral veins being forked (fig.
8c) except in subg. Gleichenia (where the area of the segment is very small). In Dicranopteris the lateral
veins are at least twice forked. Fig. 15a, c. The sori are always attached on the surface of a vein, the
sporangia attached to a small raised receptacle. In Gleichenia subg. Gleichenia the veins are very short,
and are not visible unless the lamina is cleared; in many taxonomic works the sori are said to be ter-
minal on the veins in this subgenus, but cleared specimens show that they are not. The arrangement
of sporangia in a sorus, in both genera, has been discussed and illustrated by Bower (The Ferns, vol. 2,
203-206, fig. 476, 486-489). In some species of Gleichenia subg. Gleichenia the sori are in depressions
in the substance of the lamina (e.g. G. peltophora CopPeEL.); in others they are protected by the reflexed
margins of the lamina and by outgrowths of tissue from the costa (G. vulcanica BL., fig. 1d—e). NAKAI
(/.c.), following PRESL, attempted to use these characters to establish separate genera, but the species
G. microphylla R.Br. is intermediate.
Sporangia and spores. The structure of sporangia has been fully described by Bower (i.c.).
Both trilete and monolete spores occur in the family. NAKAI (/.c.) proposed a basic division of the family
Dec. 1959] . Gleicheniaceae (Holttum) ; 3
on spore-form, but in so doing he supposed that only trilete spores occur in his restricted genus Di-
cranopteris (D. linearis and its near allies); in fact closely related species of this group have spores of
different forms, and a division of the family on spore-form is certainly unnatural.
Gametophyte. The most recent and most complete account of gametophytes in the family is by
A. G. Stroxey (Bull. Torrey Bot. Club 77, 1950, 323-339) and includes references to earlier accounts.
Species of all genera and subgenera here recognized were studied. Dr Stokey’s conclusion is that the
gametophyte in all cases shows many primitive characters, but there is little significant difference within
the family to indicate that one part is more primitive than the rest. Gleichenia subg. Gleichenia shows
specialized characters (notably the long neck of the archegonium) and on characters of the gametophyte
is judged to be further removed from the rest of the family than they are from each other. Two-celled
hairs of a peculiar nature have an origin similar to that of the larger hairs on prothalli of Cyatheaceae;
apart from these hairs, the prothalli of Gleicheniaceae most resemble those of Dipteris.
Cytology. MANTON & SLEDGE (Phil. Trans. Roy. Soc. B, 238, p. 143, pl. 4) report for Dicranopteris
linearis from Ceylon both n = 39 and n = 78; from Singapore, for what is now recognized as D. cur-
ranii COPEL., n = 39. MANTON reports verbally that a plant of D. linearis sent from Singapore and cul-
tivated at Kew is a Sterile triploid hybrid. MEHRA & SINGH also report n = 39 for D. linearis from
Northern India (Curr. Sc. 25, 1956, 168), and n = 56 for Hicriopteris glauca (probably Gleichenia
gigantea WALL., which is the common Himalayan species of this group), a species of Gleichenia subg.
Diplopterygium. BROWNLIE (Trans. R. Soc. N.Z. 85, 1958, 213-214) reports n = 20 for G. (subg. Glei-
chenia) microphylla R. Br. and n = 34 for G. (subg. Mertensia) cunninghamii. T. G. WALKER reports
(personal communication) n = 34 for two species of subg. Mertensia from Jamaica.
Anatomy. The rhizome has a simple protostelic structure except in Dicranopteris pectinata (see
Bower, /.c.). The rachis has a single C-shaped vascular strand; all the outer tissues in the rachis are
thick-walled and when mature form a very strong protection for the vascular strand. Such protection
is important in fronds which continue to grow in length for a long period.
Economic importance. HEYNE (Nutt. Pl. N.I. 1927, p. 97) records the following uses for parts of
plants of this family; the chief species used are Dicranopteris linearis and D. curranii. The rachises of
mature fronds are tied into bundles and the bundles used in making the fences of a certain kind of
fish trap; they will last two years when immersed in sea-water. Parts of the rachis of a large frond (the
largest are produced by D. curranii) when suitably split make excellent pens for writing Arabic characters
(I learned of this use also in Singapore). The vascular strands of stipe and rachis are separated and used
for special kinds of fine plaited work, being strong and pliable.
Dicranopteris thickets may be useful as preventing erosion, but they are troublesome to the forester
when they prevent regeneration of tree-seedlings in forest clearings. As the rhizomes are almost or quite
superficial, they are exposed when the thicket is cut down, and usually one such cutting is enough to
kill almost all of the plants. Fronds are sometimes cut — when in absence of other suitable material —
to provide light shade for transplanted seedlings.
Taxonomy. I published a statement on taxonomy, with discussion of the present arrangement,
in 1957 (Reinwardtia 4, p. 257-280). CopELAND (Gen. Fil. 1947) divided the Malaysian members into
four genera. It seems to me, however, that Dicranopteris is so different from the rest that a main division
should indicate this difference, and I therefore recognize two genera, Gleichenia and Dicranopteris,
the former with three subgenera which correspond to COPELAND’s genera. For one of the latter he used
the name Hicriopteris PRESL, but the type species of that genus is a Dicranopteris (described as D. speciosa
in the present work). I have therefore adopted the subgeneric name Diplopterygium, first proposed (as a
sectional name) for this group of ferns by DreLs (in ENGLER & PRANTL, Pfl. Fam. 1, 4, p. 350-356).
For the other subgenus the name Mertensia is available; it was first used in this rank by Hooker, though
with a larger content. This name was first proposed by WILLDENOw in 1804 as a generic name to cover all
known members of the family with larger divisions of the lamina than the original G. polypodioides
(THUNB.) SMITH; but as a generic name it was antedated by Mertensia RoTH (Boraginaceae) and so it
is illegitimate. In 1806 BERNHARDI published the generic name Dicranopteris, citing under it only one
species of Mertensia WILLD., viz Polypodium dichotomum THUNB., which thus becomes the type species
of Dicranopteris. I have typified Mertensia by the species M. truncata WILLD. (Reinwardtia 4, 1957,
261). Both Mertensia and Dicranopteris are used here in a more restricted sense than intended by some
earlier authors. COPELAND used the generic name Sticherus PRESL for subg. Mertensia of the present
account. This name was established by Pres for two species of which he had seen no specimens, with a
brief and confused description, and the lapse of the name is not to be regretted.
KEY TO THE GENERA
1. Young parts of plants protected by fringed peltate scales and stellate hairs. Sori of 2-5 sporangia.
Veins simple or once forked. . ai gate 1. Gleichenia
1. Young parts of plants protected by branched hairs of various s form, scales lacking. Sori of 8-15 or
more sporangia. Veins forked at least twice... = 9 -..4) ©) « «..« « 2. Dicranopteris.
+ FLORA MALESIANA [ser. II, vol. 12
1. GLEICHENIA
SMITH, Mem. Ac. Turin 5 (1793) 419, non NEcK. 1790, nom. cons. — For synonyms
see the subgenera.—Fig. 1-10.
Rhizome dichotomously branched, protostelic, near the apex protected by
peltate scales. Fronds of mature plants of indefinite growth in length (except
sometimes at high altitudes), bearing primary branches in pairs, the apex of the
main rachis dormant during the development of each pair of primary branches,
the dormant apex in some cases protected by a pair of stipule-like leaflets of
distinctive form (such stipular leaflets less often present in conjunction with
dormant apices of lateral branch-systems); primary branches often each bearing
a pair of secondary branches with a usually dormant apex between them, the
process sometimes repeated to produce ultimate branches of fourth or fifth orders;
dormant apices protected by peltate scales which (with two exceptions) are fringed
by outgrowths from the marginal cells; u/timate branches either simply pinnatifid
or bipinnatifid; /amina in all cases lobed almost to the costa, the veins in each
segment pinnately branched; lateral veins simple or once forked; costae, costules
and veins when young protected by small fringed peltate scales and by stellate
hairs, sometimes glabrescent when mature; sori one or several to each segment
of the lamina, upon the lateral veins, not terminal upon them (on the acroscopic
branch of a forked vein), exindusiate, each consisting of 2-5 large sporangia;
paraphyses, in the form of small stellate hairs or small scales with long marginal
hairs, often present with the sporangia; annulus complete and oblique, dehiscing
dorsally. Spores monolete or trilete, smooth and translucent, 256 or more in each
sporangium.
KEY LO DME SRE CIES
1. Ultimate branches (branches on each side of an ultimate dormant apex) bipinnatifid.
2. Segments of the lamina not much longer than wide; one sorus on each segment. (subg. Gleichenia).
3. Fully developed pinnule-lobes not deeply concave beneath (the edges only slightly revolute).
Tissues adjacent to costa not swollen. Sporangia when young in a circular depression in the lamina.
4. Edge of depression occupied by sorus distinctly raised. Small scales on lower surface of lamina.
1. G. peltophora
4. Edges of depression occupied by sorus not raised. No small scales on lower surface of lamina.
2. G. microphylla
3. Fully developed pinnule-lobes deeply concave beneath, the distal edges strongly revolute. Tissue
adjacent to the costa more or less swollen. Sporangia not in a depression in the lamina.
5. Costae persistently scaly; tissue adjacent to the costa slightly swollen. Sporangia often 3.
3. G. yulcanica
5. Costae of fully developed fronds usually quite glabrous; tissue agent to the costa much swollen.
Sporangia 2. . : F 4. G. dicarpa
2. Segments of the lamina elongate, the costule bearing several forked veins, each of which may bear
a sorus on its acroscopic branch. (subg. Diplopterygium).
6. Scales on dormant apex of rachis entire, to 10 by 3 mm. Segments of lamina at c. 45° to costae.
5. G. laevissima
6. Scales on dormant apex of rachis fringed with hairs or setae, or with a broad translucent margin.
Segments of lamina almost at right angles to costae.
7. Scales on dormant apex of rachis 1 mm wide or more, edges fringed with spreading hairs to
0.5 mm long, or with translucent edge bearing fine short hairs.
8. Rachis-branches persistently quite covered with scales, or with a mixture of scales and hairs,
on lower surface.
9. Rachis-branches covered almost entirely with thin brown scales 4-5 mm long and nearly 1 mm
wide, stellate hairs few. . . . 6. G. paleacea
9. Rachis-branches covered with a dense felt of stellate hairs, and also with + abundant scales.
10. Pinnules commonly 12-16 by 2-21/, cm. Upper surface of the lamina “not swollen between
the veins. Stipular leaflets present . . . . . . 7. G. yolubilis
10. Pinnules commonly 6-9 by 0.9-1.3 cm. Upper surface of the lamina much swollen between
Dec. 1959] Gleicheniaceae (Holttum) 5
the veins. No stipular leaflets .. 8. G. bullata
8. Rachis-branches not persistently quite covered with scales nor r with ¢ a felt of stellate hairs on the
lower surface.
11. Very few persistent scales on the lower surface of rachises and costae. Pinnules to 22 by 3.5 cm.
9. G. longissima
11. Many persistent dark scales on rachises and costae. Pinnules to 12 by 2 cm 10. G. clemensiae
7. Scales on dormant apex of rachis narrower, their edges bearing short stiff oblique setae.
12. Segments of the lamina near base of each pinnule constricted at the base (i.e. widening just a
little above the constriction), joined together by a very narrow wing of even width along the
costa.
13. Many such segments (20 or more pairs) on each pinnule.
14. Lamina rigid; segments of lamina ae mm wide; scales of dormant apex uniformly dark
DIOWDs = |. . . . 11. G. angustiloba
14. Lamina thin; segments of lamina 31), mm wide; scales of dormant apex pale with dark edges.
13. G. deflexa
13. Few (at most 7-8) such segments on each pinnule.
15. Pinna-rachises more or less persistently scaly; costules 31/,-41/, mm apart. 14. G. sordida
15. Pinna-rachises sparsely scaly, soon glabrous; costules 5!/,-7 mm apart. 12. G. elmeri
12. Segments of lamina near base of each pinnule not thus constricted (or not more than one
segment on the largest pinnules).
16. Stipular leaflets having lobes similar to pinnule-segments, not narrow and acuminate. Scales
on rachis-apex not dark-fringed. . . eye 15. G. norrisii
16. Stipular leaflets having very narrow acuminate lobes quite different from normal segments of
pinnules. Scales on rachis-apex dark-fringed.
17. Distal half of each rachis-branch strongly zig-zag, the pinnules deflexed so that each is
IimUINem with thempanrt. ofethesrachisebeyondeit sea) Hen t eee = 16. G. matthewii
17. Distal half of each rachis-branch not thus zig-zag.
I, Praimwes ClO x7 2 Cal, 4 ee ERC. eer Poel Ur, 17. G. brevipinnula
18. Pinnules commonly 20 by 3-4 cm.
19. Dark rigid stellate hairs abundantly persistent on the lower surface of veins and lamina.
18. G. blotiana
19. Stellate hairs light brown, not abundant nor Been on the lower surface of veins and
lamina. ee Ce eee 19. G. conversa
Ultimate branches simply, pinnatiide Gee Mertensia):
Sh Stipule-like leaflets, different from ordinary segments of the lamina, present at the branchings
of the main rachis (on fronds of mature size).
21. Segments of lamina at right angles to the costa, or more or less deflexed. Lower surface not con-
spicuously glaucous.
22. Segments of lamina deflexed (lower ones much so), thick, with edges much revolute when dry.
Main rachis-branches once or twice forked . . . rae es 20. G. reflexipinnula
22. Segments of lamina not or little deflexed. Main rachis- branches often more than twice forked.
23. Forkings of rachis-branches equal or nearly so, not alternately unequal with larger branches
at successive forks forming almost a straight line.
24. Scales on costules very abundant, spreading so as almost to cover the lower surface of the
lamina. Veins on lower surface strongly raised . . . 21. G. venosa
24. Scales on costules not persistently covering the lower surface of ‘the lamina. Veins not strongly
raised ‘ : 22. G. truncata
23. Forkings of rachis- branches unequal, ‘the larger branch at successive > forks alternately to left
and right, axes of these larger branches in almost siraight line. . . . 23. G. milnei
21. Segments of lamina distinctly oblique-ascending. Lower surfaces distinctly glaucous.
25. Primary branch-systems 5 times forked on large fronds, branches of all orders except the first
leafy; no stellate hairs on the lower surface of veins . . 8 24. G. oceanica
25. Primary branch- ~systems twice forked; stellate hairs (or at least their bases) persistent on the
lower surface of veins . . a: 25. G. erecta
20. Stipule-like leaflets absent (sometimes present in G. lohers var. aoe.
26. Margins of segments of lamina not distinctly toothed.
27. Only the ultimate branches fully leafy (penultimate without lamina or with some scattered
segments).
28s Witimate, branches: to20)cmylongieeie . 9 linea ecu ees Senn eeeee2O.8G. opulchra
28. Ultimate branches 30-40 cm long.
29. Segments of the lamina to 25 mm long, costules aye mm apart. Primary rachis-branches
usually 3 times forked ii ‘ 27. G. pseudoscandens
29. Segments of lamina to 17 mm long, ‘costules 4 mm apart. Primary rachis-branches usually
twice forked 5 2142-98285). G.walstonii
27. At least the penultimate ‘branches fully leafy, lower ones ‘sometimes also.
6 FLORA MALESIANA (ser. Ik volar
30. Lamina-segments rarely more than twice as long as the distance between bases of adjacent
costules.
31. Segments 8-12 mm long, costules 4-5 mm apart. Rachis-branches of Ist and 2nd orders 3-4
cm long. Scales on the main as ek oe brown, more than | mm wide at base; scales on the
costae rusty brown 3 aS 29. G. vestita
31. Segments 3-6 mm long, costules 3 mm apart. Rachis- branches of ist and 2nd orders 1—2 cm
long. Scales on the main peut very dark, narrower; scales on the costae dark at base,
paler distally 30. G. bolanica
30. Lamina-segments much 1 more than twice as Jong as the distance between adjacent costules.
32. Scales on the costae only a few cells wide at the base, about half their length consisting of a
hair, these hairs crisped and entangled » ids : 31. G. hispida
32. Scales on the costae with short hair-points, their bases wider and their apices not entangled.
33. Costules at 60°-70° to costae : : 25. G. erecta
33. Costules at almost a right angle to the costae.
34. Ultimate branches 25-45 cm long, eee cm wide; costules on ultimate branches 4-5 mm
aparte yee. . . 32. G. brassii
34. Ultimate branches smaller, ‘costules on ultimate branches 3-4 mm apart. 33. G. loheri
26. Margins of the segments of the lamina distinctly toothed, at least towards the apex.
35. Margins toothed almost to base of segments. Costules at c. 45° to costae. 34. G. flabellata
35. Margins toothed only near apex of segments. Costules at wider angle to costae. 35. G. hirta
1. Subgenus Gleichenia
Hott. Reinwardtia 4 (1957) 262.—Calymella Pres_, Tent. Pterid. (1836) 48;
CHING, Sunyatsenia 5 (1940) 287; Naxkal, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950)
40.—Gleicheniastrum Pres_, Abh. (K.) Bo6hm. Ges. Wiss. M.-N. Cl. 5 (1848)
338; NAKAI, /.c. 42.—Gleichenia subg. Eugleichenia Diets in E. & P. Pfl. Fam.
1, 4 (1900) 355.—Gleichenia; CopeEL. Gen. Fil. (1947) 26.—Fig. 1-3.
Young plants first producing determinate bipinnatifid fronds, the largest of
these as large as the branches of later fronds and sometimes fertile (plants in
exposed places at high altitudes sometimes producing only fronds of this kind);
then fronds bearing pairs of primary branches like the fronds of the first stage;
in some cases the primary branches bearing one or more pairs of determinate
bipinnatifid secondary branches, or the secondary branches leafless and bearing
pairs of tertiary branches, the ultimate branches always determinate and _ bipin-
natifid. Leaflets of bipinnatifid fronds or of ultimate branches lobed almost to
the costa, lobes hardly longer than wide; veins in each lobe pinnately branched
but without a conspicuous costule, lateral veins simple. One sorus on each lobe,
upon the basal acroscopic vein, superficial or sunk in the substance of the lamina.
Distr. About 10 spp. in tropical and southern Africa, the Mascarene Islands, Malaysia and Austra-
lasia (not in Ceylon and India).
Ecol. In Malaysia only on mountains, in exposed places, usually in sandy or acid, peaty soils, sometimes
in association with members of the other subgenera.
1. Gleichenia peltophora CopeL. Philip. J. Sc. 40
(1929) 292, t. 1.—G. circinnata var. borneensis BAK.
J. Bot. 17 (1879) 37.—G. borneensis C. Cur.
Gard. Bull. S. S. 7 (1934) 211.—Calymella bor-
neensis CHING, Sunyatsenia 5 (1940) 288.
var. peltophora.
Rhizome-scales ovate, dark, entire. Determinate
bipinnatifid fronds to 40 cm high (including stipe),
often fertile; branched fronds bearing 1 or few
pairs of branches; primary branches bipinnatifid-
determinate or once forked with a permanently
dormant apex in the fork, the primary branch
always bearing leaflets (where primary branches
are not forked, the main rachis may be leafy below
junction with branches); sca/es on resting apex
of rachis to 11/, by 1 mm, ovate, entire, convex,
shining, very dark brown with narrow pale edge;
similar but smaller scales scattered on lower sur-
faces of rachises and less abundantly on costae;
ultimate branches 20-30 cm long, bearing many
leaflets, costae of adjacent leaflets 4-6 mm apart;
leaflets 2-5 cm long, 2!/,-4 mm wide; lobes c.
114 mm wide at base, gradually narrowed to
rounded apex, edges slightly reflexed, lower sur-
face slightly concave, glaucous, bearing scattered
circular scales 0.2-0.3 mm diameter. Sori, if
present, each sunk in a circular depression in the
substance of the lamina, the lower surface of the
lamina somewhat raised round the edge of the
depression which occupies about half the width of
the base of a lobe.
Type: Copeland s.n., 1 May 1917, Mt Matutum,
Mindanao, 1600 m (H. Copel.).
Dec. 1959] Gleicheniaceae (Holttum) T
Fig. 1. Gleichenia microphylla R. Br. a. Lower surface of part of a pinnule, showing one mature sorus,
and two soral depressions from which sporangia have been removed, x 10, 5. one lobe of lamina,
cleared to show veins and position of sorus, 13, c. a scale, x 50. — G. vulcanica BL. d. Part of pinnule
_ from below (scales removed), x 10, e. same cleared to show veins and position of sorus, x 13. —G.
dicarpa R. Br., f. Lower surface of part of pinnule, « 10, g. same cleared to show veins, 13, A. upper
surface of cleared pinnule showing position of sori. — Diagrams of various stages of branching which
may occur during development of a single plant of G. vulcanica BL., i. First stage: plants in exposed
places or at high altitudes may never develop further, j. transition stage, k. primary branches simple
(corresponding to condition of subg. Diplopterygium), 1. primary branches once forked, m. primary
branches twice forked, n. primary branch proliferous beyond the first fork.
Distr. Malaysia: N. Borneo (Mt Kinabalu),
Philippines (Mindanao), South Central Celebes,
West New Guinea (several localities).
Ecol. In open places on mountain summits or
high ridges, 1500-2500 m; on Mt Kinabalu in
association with G. vulcanica and Dipteris novo-
guineensis POSTH.
var. schizolepis C. CHR. ex HoLtrT. Reinwardtia
4 (1957) 262.
Scales on rhizome short-fringed; scales on costae
bearing a few rigid dark brown marginal hairs;
scales on lower surface of lamina replaced by dark
red-brown stellate hairs; fronds of the only speci-
men unbranched, with leaflets 2!4 cm long, sterile.
Distr. Borneo (Sarawak: Mt Murud), 2500 m,
once found.
2. Gleichenia microphylla R.Br. Prod. (1810) 161;
y. A. v. R. Handb. Suppl. (1917) 80, incl. var.
semivestita v. A. v. R.—G. semivestita LABILL.
Sert. Austro-Cal. (1824) 8, t. 11.—Calymella
microphylla PResL, Tent. Pter. (1836) 49.—
Gleicheniastrum microphyllum Pres_, Abh. (K.)
Bohm. Ges. Wiss. M.-N. Cl. 5 (1848) 338; NAKAI,
Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 45, incl.
var. semivestitum NAKAI.—Calymella circinnata
(7G. circinnata Sw.) CHING, Sunyatsenia 5 (1940)
288.—Calymella semivestita CHING, I.c.—Glei-
cheniastrum lowei NAKAI, Bull. Nat. Sc. Mus.
Tokyo n. 29 (1950) 44.—Fig. la-c, 2.
Rhizome to 3 mm diameter, the young parts
covered with narrow dark brown rigid fringed
scales. Fronds on young plants _bipinnatifid,
determinate, rarely fertile, on old plants usually
FLORA MALESIANA [ser. Tl, volaet
Fig. 2. Gleichenia microphylla R. Br. on edge of dwarf forest on wet sandy soil near summit of Kedah Peak,
Malaya, 1200 m (HoLtrum, 1953).
Dec. 1959]
Gleicheniaceae (Holttum) i 9
branched, the primary branches sometimes
forked with leaflets below the fork, rarely proli-
ferous beyond the fork and then not bearing
leaflets below the fork; secondary branches
sometimes also forked. Sca/es on resting apices of
rachis narrow, very dark, with pale thin marginal
hairs towards the base and rigid dark setae near
apex; rachises and costae more or less densely
clothed with very small dark scales, their edges
bearing rigid concolorous setae. Ultimate branches
12—25 by 4-5 cm; leaflets 7-9 mm apart, 2!4-4 mm
wide; lobes of /amina 1—1.8 mm long, 0.7—1.5 mm
wide at the base, apex rounded, edges usually a
little reflexed; lower surface slightly concave, not
glaucous. Sorus (if present) at first embedded in a
circular cavity which occupies half the width of
the lobe, edges of cavity not raised; sporangia 3-4
(rarely 5), surrounded by brown hairs not longer
than the ripe sporangia.
Type: Robert Brown,
Wales (BM; dupl. at K).
Distr. From Annam through Malaysia to
Australia, New Caledonia, and New Zealand; in
Malaysia: Sumatra, Malay Peninsula, Lingga
Arch., Borneo, and Moluccas (Ambon).
Ecol. In open places in sandy or thin peaty soil
on summits or exposed ridges of mountains at
750-1800 m, especially on sandstone. At Fraser’s
Hill, Malay Peninsula, 1250 m, this species occurs
in open places on a quartzite ridge, not on the
neighbouring granite; it occurs on the summits of
some isolated granite mountains (Mt Ophir,
G. Belumut). It is abundant from 1000 m upwards
in open Leptospermum forest, with Sphagnum, in
thin peaty soil on the sandstone mountain G.
Jerai or Kedah Peak.
Note. For a note on the type specimen of
G. circinnata, see note under 4. G. dicarpa.
Port Jackson, N.S.
3. Gleichenia vuleanica BL. En. Pl. Jav. (1828)
251; Racis. Fl. Btzg 1 (1898) 10; v. A. v. R.
Handb. (1909) 56; BACKER & PosTH. Varenfl.
Java (1939) 353.—Calymella vulcanica PRESL,
Abh. (K.) B6hm. Ges. Wiss. M.-N. Cl. 5 (1848)
338; CHING, Sunyatsenia 5 (1940) 287; NAKAI,
Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 42.—G.
dicarpa var. vulcanica CHRIST, Ann. Jard. Bot.
Btzg 15 (1898) 75.—G. squamosissima COPEL.
Philip. J. Sc. 75 (1941) 348, pl. 1.—Calymella
Squamosissima NAKAI, |.c.—Fig. 1d—e, i-n, 3.
Similar in habit to G. microphylla, differing as
follows: leaflets commonly 4-5 mm apart; seg-
ments of lamina less than 1 mm long and wide;
lower surface glaucous, deeply concave, with
edges much reflexed and the surface adjacent to
the costa also swollen; many small thin scales,
with slender crisped marginal hairs, all along
costules; sporangia 2 or 3, not in a depression
in the lamina. At very high altitudes often only un-
branched fronds, very densely scaly, are produced.
Type: Blume, Java (L).
Distr. Malaysia: Sumatra, Malay Peninsula
(3 localities), W. Java, Borneo, Philippines
(Mindanao, Mindoro), Celebes, and New Guinea.
Ecol. Abundant in open scrub (on both dry
stony and wet ground) on volcanic mountains
throughout W. Java and Sumatra, 1800-3600 m,
at these altitudes apparently less abundant on
granite mountains in Borneo; densely scaly at
highest elevations both in N. Sumatra and in New
Guinea (G. sqguamosissima COPEL.).
Fig.3. Gleichenia vulcanica BL. in the high mountains
of Mt Goh Lembuh, Gajo Lands, N. Sumatra,
sheltered in rock crevices, c. 3000 m (1937).
4. Gleichenia dicarpa R.Br. Prod. (1810) 161;
C. Cur. Ark. Bot. 9% (1910) 33.—Calymella
dicarpa PRESL, Abh. (K.) BOhm. Ges. Wiss. M.-N.
Cl. 5 (1848) 338.—Calymella circinnata (non G.
circinnata Sw.) CHING, Sunyatsenia 5 (1940) 288.
—Fig. 1f-h.
Like G. vulcanica, but the /eaflets not copiously
scaly (costae almost or quite glabrous when
mature), the aperture of the concave lower surface
of each segment of the lamina reduced to much
less than half of the area enclosed by the outline
of the segment as seen from below, owing to much
swelling of the costal tissue; sporangia 2, filling
the aperture.
10 FLORA MALESIANA [ser. Il, voleaie
Type: Robert Brown, Tasmania (BM; dupl. New Guinea, occurring at 750-850 m, in open
at K). forest or rocky places.
Distr. Australia, New Caledonia, in Malaysia: Note. Though typical specimens of G. vulcanica
New Guinea, Philippines (Mindanao). in Java differ markedly from typical specimens of
Ecol. On Normanby and Rossel Islands, SE. _ G. dicarpa in Australia, some specimens on moun-
ale unnl
Fig. 4. a. Diagram of branching in Gleichenia subg. Diplopterygium. — G. longissima BL., 6. Part of
rachis-branch bearing two pinnules, x 2/3, c. segment of lamina, lower surface, showing veins and sori,
x 4, d. resting apex of main rachis and bases of branches, with stipular leaflets, nat. size. — Scales and
hairs. — G. longissima BL., e. A single scale from apex of rachis, x 8, f. edge of scale, x 133, g. stellate
hair, x 33.—G. conversav. A. v. R.,h. A single scale, x 8, i. edge of scale, x 133, j. stellate hair 3336
Dec. 1959]
Gleicheniaceae (Holttum) 11
tains in Mindanao and New Guinea seem some-
what intermediate. Ecological distinctions between
the two have not been studied.
CHRISTENSEN, in his notes on specimens in the
herbarium of Swartz at Stockholm (/.c.) stated
that the type specimen of G. circinnata was
identical with G. dicarpa R.Br. With his original
description SWARTZ gave no precise locality, but
in his ‘Synopsis Filicum’ he stated that the species
was from Botany Bay, near Sydney. The only
specimen labelled G. circinnata in SWARTZ’s
herbarium from that locality is certainly not G.
dicarpa, though it has only two sporangia on each
lamina-segment; the segments are not pouch-
shaped in the manner of G. dicarpa. One can
only conclude that CHRISTENSEN did not make a
careful examination of the specimen. However,
the Botany Bay specimen is not certainly the
original on which the species G. circinnata was
based (furthermore, the original description in-
cluded the phrase capsulis quaternis, not true of
the Botany Bay specimen). I therefore refrain
from reverting to CHRISTENSEN’S earlier identifica-
tion of G. microphylla R. BR. with G. circinnata Sw.
After the publication of CHRISTENSEN’s note,
some botanists used the name G. circinnata to
replace G. dicarpa, but it seems clear that this was
an error, and I| therefore restore R. BROWN’s name.
2. Subgenus Diplopterygium
HoLtt. Reinwardtia 4 (1957) 261.—Gleichenia subg. Mertensia sect. Diplopterygium
DiELs in E. & P. Pfl. Fam. 1, 4 (1900) 353.—Gleichenia subg. Mertensia § 1 HooK.
Sp. Fil. 1 (1844) 4.—Gleichenia subg. Mertensia sect. Dicranopteris v. A. v. R.
Handb. (1909) 57, p.p.—Dicranopteris UND. Bull. Torr. Bot. Cl. 34 (1907) 249,
p.p.—Sticherus § Hicriopteris C. CHR. in Verdoorn, Man. Pterid. (1939) 530.—
Hicriopteris (non PRESL) CHING, Sunyatsenia 5 (1940) 277; CopeL. Gen. Fil.
(1947) 28.—Mesosorus Hassk. Fil. Jav. 1 (1856) 2, p.p.—Diplopterygium NAKAI,
Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 47.—Fig. 4-6.
Fronds of a young plant producing pairs of bipinnatifid branches, with periodic
dormancy of the apex, from an early stage, the lateral branches of such fronds of
immature plants small and sterile; periodic dormancy of the apex of the main
rachis, and no other dormancy, occurring on fronds of mature plants; lowest
leaflets on the branches often deltoid and bipinnatifid (or with deltoid-bipin-
natifid lowest branchlets), forming stipule-like structures which protect the
dormant apex of the main rachis; remaining leaflets very deeply pinnatifid,
segments of the lamina oblong, each with a costa bearing on either side several
once-forked veins. Sori on the acroscopic branches of the veins, several to each
segment of the lamina.
Distr. More than 20 spp. in NE. India, Burma and Indo-China, China, Japan, Malaysia, Polynesia,
Hawaii, and tropical America (1 sp.). This subgenus is far more diversified in Malaysia than else-
where.
Ecol. All species occur on mountains, some descending to only 600 m. All may form dense thickets
on edges of forest or other open places; fronds can climb to a height of 6-7 m, if trees suitable for
support are present.
Note. Some fossils of Cretaceous age in West Greenland (lat. 70-71°N) have exactly the characters
of this subgenus (habit of branching, venation, position of sori, nature of sporangia).
5. Gleichenia laevissima CHRisT, Bull. Ac. Inst.
Geogr. Bot. III, 11 (1902) 268; v. A. v. R. Handb.
(1909) 795; Suppl. (1917) 82.—Hicriopteris
laevissima CHING, Sunyatsenia 5 (1940) 280.—
Diplopterygium laevissimum NAKAI, Bull. Nat. Sc.
Mus. Tokyo n. 29 (1950) 52.
Scales on rachis-apex 10 mm long, nearly 3 mm
wide, acuminate, entire; rest of frond quite
glabrous except for very short simple hairs on
lower surface of veins and lamina of young fronds
and soral hairs. Primary rachis-branches to at
least 70 cm long, lowest leaflets not stipule-like;
leaflets at an angle of about 45° to rachis, largest
17-20 cm long, 214-3 cm wide, costae 214-314 cm
apart; lamina lobed almost to the costa, costules
at about 45° to the costa and 314-414 mm apart;
segments of lamina 2-2!4 mm wide above the
base, narrowed gradually towards the apex;
lower surface not glaucous, veins only slightly
raised; upper surface not raised along margins of
costae, veins raised only near their bases. Sporangia
usually 4, pale, with very slender pale hairs.
Type: Bodinier 1295, Kouy-yang (P).
Distr. China (Kweichow, Chekiang, Yunnan,
Fukien), Formosa, and Malaysia: Philippines
(Luzon).
Ecol. In the mountains of Luzon, c. 2000 m.
[ser. Tly volute
\ ky
\ Nth {i tui
14,
a
<
Z.
<
n
ca]
4
<
<
a4
e)
4
fL,
Fig. 5. Gleichenia longissima BL. on edge of forest, Taiping Hills, Malaya, c. 800 m (HOLTTUM, 1952).
Dec. 1959]
Gleicheniaceae (Holttum) —
13
6. Gleichenia paleacea (CopeL.) Ho.ttr. Rein-
wardtia 4 (1957) 265.—Hicriopteris paleacea
CopeEL. Philip. J. Sc. 81 (1952) 3.
Scales on main rachis-apex not seen; sca/es on
primary branches and costae beneath very
copious, light brown, larger ones 4-5 mm long
and 1 mm wide, those on costae commonly 2 mm
long and more than 0.5 mm wide, thin, fringe fine,
close, concolorous; smaller scales narrower with
long fine fringe and hair-tips; no persistent stellate
hairs On veins except in sori. Primary rachis-
branch more than 100 cm long (collector’s note);
costae 214 cm apart, at right angles to rachis;
largest pinnules 15-16 cm long, -244—3 cm wide,
lobed almost to the costa, basal lobes slightly
reduced and very close to the rachis; costules
c. 4 mm apart, very slightly oblique; /amina-
segments thin but firm, hardly narrowed above the
base, edges slightly reflexed when dry so that
sinuses between segments are 1-2 mm wide;
lower surface glaucous, veins distinct and slightly
raised; lamina on each side of the costa on upper
surface raised as in G. gigantea WALL. and some
long hairs persistent in the groove, veins slightly
sunk in upper surface of lamina when dry. Sori
commonly of 3 sporangia, with numerous pale
hairs nearly 1 mm long.
Type: Elmer 9902 (Herb. Copel.; dupl. at BM,
Keebor is. -P):
Distr. Malaysia: Philippines (Negros: Cuernos
Mts), once collected.
Note. This is near to G. longissima BL., but
differs in persistent abundant scaliness. A collec-
tion of G. longissima from Canlaon Volcano,
Negros (MERRILL 607) shows very scaly young
parts of a frond, but the scales are much narrower
and the old parts are glabrous.
7. Gleichenia volubilis JUNGH. Java 1 (1853) 592,
664; v. A. v. R. Handb. Suppl. (1917) 83,p.p.;
BACKER & PostH. Varenfl. Java (1939) 256.—
Mertensia arachnoides Hassk. in Hook. J. Bot.
Kew Misc. 7 (1855) 332.—Mesosorus arachnoides
Hassk. Fil. Jav. 1 (1856) 6.—G. arachnoides METT.
in Mig. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 47;
Racis. Fl. Btzg 1 (1898) 11; v. A. v. R. Handb.
(1909) 58 (arachnoidea); non G. arachnoidea
CUNN. 1844.—G. glauca var. arachnoides C. CHR.
Ind. Fil. (1905) 320.—Hicriopteris volubilis CHING,
Sunyatsenia 5 (1940) 280.—Diplopterygium volu-
bile NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29
(1950) 55.
var. volubilis.
Scales on main rachis-apex nearly black, to at
least 6 mm long and 114 mm wide, gradually
narrowed towards apex, edges closely fringed
with fine spreading pale hairs (or with a broad,
thin translucent border bearing short hairs) ; similar
but smaller scales on lower surfaces of branch-
tachises, costae and costules (smallest scales not
dark), these surfaces also densely covered with
a persistent felt of pale stellate hairs each less
than 0.5 mm diameter, old costae showing the
Taised points of attachment of former scales;
stellate hairs on costules and lower surface of
veins rusty brown. Primary rachis-branches to
at least 150 cm long and 30 cm wide; stipule-like
leaflets present and deeply lobed; pinnules 12-16
cm long, 2—3 cm wide, costules 314-4 mm apart,
at right angles to costae; /amina-segments firm,
slightly narrowed above the base, their edges
reflexed when dry, lower surface not (or very
slightly?) glaucous, veins strongly raised; upper
surface of rachis covered with stellate hairs as
lower surface together with very narrow pale
ciliate scales; smaller pale scales and hairs at
first in groove of upper surface of costae (lamina
slightly raised on each side of the groove); veins
on upper surface paler than lamina and slightly
raised in dried specimens (lamina between veins
sometimes slightly raised but not swollen); sporan-
gia surrounded by a group of crisped red-brown
hairs.
Type: Junghuhn s.n., 1839, G. Gedeh (L).
Distr. Malaysia: Java, Sumatra, and Central
Celebes (one collection).
Ecol. Forming dense thickets in open places
on ridges, and on edges of forest, often very
abundant, 1800-3000 m.
Note. The Celebes specimen has smaller pin-
nules than normal in Java (10-12 cm long, 114 cm
wide), with thinner lamina and veins hardly
prominent beneath.
var. peninsularis HoLtr. Reinwardtia 4 (1957)
265.
Scales on lower surfaces of rachises and cosiae
rusty brown like the stellate hairs, not black.
Type: F. M. S. Museum s.n., June 1917 (K).
Distr. Malaysia: Malay Peninsula (G. Bintang
on Kedah-Perak boundary), once collected.
8. Gleichenia bullata Moore, Ind. Fil. (1862) 374;
C. Cur. Gard. Bull. S.S. 7 (1934) 212.—G.
volubilis (non JUNGH.) v. A. v. R. Handb. Suppl.
(1917) 83, p.p.—RHicriopteris bullata CHING,
Sunyatsenia 5 (1940) 279.
Differs from G. volubilis JUNGH. as follows:
scales on rachises, costae and costules much more
abundant, those on costules often pale with a
dark base; stipular leaflets lacking; primary
rachis-branches shorter, pinnules commonly 1.3—
1.5 cm apart, 6-9 cm long, 0.9-1.3 cm wide;
veins on upper surface of Jamina (when dry)
much sunken, the surface between them swollen.
Type: Low, Mt Kinabalu, 7000 ft (K).
Distr. Malaysia: Borneo (Sarawak: Mts Dulit
and Tibang; N. Borneo: Mt Kinabalu), New
Guinea (Mt Dayman).
Ecol. Forming thickets
1700-3000 m.
Note. The bullate character of the upper surface
of the lamina, and the scaliness, distinguish this
from dwarfed specimens of G. volubilis which may
have pinnules no larger than those normal in
G. bullata.
like G. volubilis,
9. Gleichenia longissima BL. En. Pl. Jav. (1828)
250; Racis. Fl. Btzg 1 (1898) 10; BACKER &
14 FLORA MALESIANA
[ser. rhe vol. |!
PostH. Varenfl. Java (1939) 256; HoLtrT. Rev. FI.
Mal. 2 (1955) 67.—G. excelsa J. SM. ex HOOK.
Sp. Fil. 1 (1844) 5, t. 4B.—G. glauca v. A. v. R.
Handb. (1908) 58 et al. p.p.—Hicriopteris longis-
sima CHING, Sunyatsenia 5 (1940) 280.—Diplop-
terygium longissimum NAKAI, Bull. Nat. Sc. Mus.
Tokyo n. 29 (1950) 53.—Fig. 4b-g, 5.
Scales on resting apex of rachis as in G. volubilis
JUNGH.; young expanding parts of fronds covered
with loose indumentum of narrow rust-coloured
scales and lax stellate hairs, a few such scaies
and hairs sometimes persistent on lower surface
of costae and costules, mature rachises quite
smooth and glabrous. Primary rachis-branches to
about 200 cm long and 40 cm wide; largest pin-
nules 15-22 cm long, 2144-314 cm wide, costae
2.8-3'4 cm apart; lamina thin, lobed almost to
costa, costules 4-5 mm apart, slightly oblique to
costa; segments of lamina slightly narrowed above
the base, edges slightly reflexed when dry; lower
surface glaucous, veins slender and slightly raised;
upper surface of rachis and costae glabrous,
lamina slightly raised on each side of costae; veins
on upper surface slightly raised near base, rarely
also distally, very slender; sori usually of 3
sporangia, surrounded by crisped brown hairs.
Type: Blume, Java (L; dupl. at K, P).
Distr. Southern China to Indo-China through
Malaysia to Melanesia (Fiji, Tahiti), in Malaysia:
not collected in the Lesser Sunda Islands E of
Flores, Celebes, the Moluccas, and New Guinea.
Ecol. Forming thickets in clearings and on
edges of forest, more or less persisting in open
secondary forest; 750-1800 m. RACIBORSKI states
that this species may climb very high on trees.
Note. This is the most widely distributed species
in the subgenus. In Java and Sumatra it is replaced
at higher altitudes by G. volubilis; a comparison
of the two species near the transition zone needs
to be made.
10. Gleichenia clemensiae (CoPEL.) HoLtTtT. comb.
nov.—Hicriopteris clemensiae CopEL. Un. Cal.
Publ. Bot. 18 (1942) 217.—G. papuana Hovtt.
Reinwardtia 4 (1957) 266.
var. clemensiae.
Scales on resting apex of main rachis dark
brown, shining, 4 mm long, more than 1 mm wide,
edges bearing spreading hairs to 0.5 mm long;
lower surface of rachises and costae bearing
numerous persistent scales like those on the resting
apex but smaller, dark and shining with pale edges
and fringe, those on the costae commonly 1144 mm
long and 14 mm wide, old costae minutely warty
from the raised bases of former scales; scales
on lower surface of costules abundant, very small,
laxly fringed; no hairs on veins, apart from sori;
upper surface of rachis rather persistently covered
with very narrow scales and long lax hairs, upper
surface of costae hairy near the base only. Rachis-
branches 120 cm long; pinnules spreading at right
angles, 2!4 cm apart, largest 12 cm long, 2 cm
wide; costules 2!4-3 mm apart; lamina thin,
veins slightly prominent on lower surface, not on
upper; soricommonly of 3 sporangia, with crisped
paraphyses.
Type: Clemens 41227, Morobe, Matap, 5000-
6000 ft (H. Copel.).
Distr. Malaysia: SE. New Guinea (Morobe,
and Milne Bay Distr.: Mt Dayman).
Ecol. Scrambling to 2-3 m, forming dense
tangles on edges of mossy forest, 1600-2250 m.
var. membranacea (HOLTT.) comb. nov.—G.
papuana var. membranacea HOLTT. Reinwardtia 4
(1957) 266.
Scales on rachis-apex pale and very thin near
the edges which are very shortly fringed; scales
on rachises and costae more abundant, dark at
the base only.
Type: Brass 24763 (BM).
Distr. Malaysia: New Guinea (Goodenough I.).
Ecol. Plentiful in openings in forest, 1600 m.
11. Gleichenia angustiloba HoLtr. Reinwardtia 4
(1957) 263.
Rhizome 7 mm diam., young parts covered with
shining dark brown scales 5-7 by 144 mm, old
parts warty from bases of former scales; stipes
6-7 mm diam., the base at first scaly like the
rhizome and then warty, upper part covered with
small scales like those of the rachises, later
asperulous. Primary rachis-branches 100 cm long
or more; stipular leaflets broadly deltoid, to 4 cm
long, pinnatifid, lowest lobes deeply and narrowly
lobed again; scales on apex of main rachis dark
brown, shining, narrow, 3-4 mm long, the edges
bearing concolorous oblique rigid setae; rachises
and costae, and lower surfaces of costules more or
less persistently covered with smaller scales of
various sizes and red-brown stellate hairs. Pin-
nules to 15 cm long and 3-314 cm wide, 2—3'4 cm
apart, the distal ones distinctly deflexed, the lower
ones at right angles to rachis; all segments of the
lamina c. 2% mm wide, separated by wide sinuses,
many of the lower segments being constricted at
the base and joined laterally by a very narrow
wing along the costa (wing about 0.2 mm wide);
lowest segments forming distinct separate leaflets;
costules 4 mm apart; veins prominent on both
surfaces; sori of 4 sporangia with long red-brown
crisped hairs, no other hairs on the veins of mature
fronds.
Type: Brass 4960, Mt Tafa (BM, dupl. at K,
Bo, US).
Distr. Malaysia: E. New Guinea (Mt Tafa,
and on Asaro-Mairi Divide, Goroka Subdistr.).
Ecol. Altitude 2400 m; ‘Common in native
rest clearings in ridge-crest forest; conspicuous,
rambling amongst forest fringe bushes and small
trees; fronds 4-5 m with 6-8 or more pairs of
large spreading pinnae”’ (BRAss).
12. Gleichenia elmeri CopeL. in Elmer, Leaf.
Philip. Bot. 3 (1910) 799.—Diplopterygium elmeri
NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 49.
Scales on rachis-apex not seen; scales on costae
sparse; scales on costules at first abundant,
Dec. 1959]
Gleicheniaceae (Holttum) = a 15
longest 1-1'14 mm long, narrow, rusty brown with
well-spaced rigid spreading marginal hairs; very
small scales and stellate hairs present on veins
and with sporangia, hair-branches stiff, c. 0.2 mm
long; dark brown short clavate hairs abundant
on costules and less so on sides of costae. Pinnae
150 cm long (fide COPELAND); largest pinnules
25 cm long, 344-4 cm wide, 4144-5 cm apart;
costules 514-7 mm apart, at right angles to costa;
several segments of lamina constricted at base,
widely separated and joined by a wing 14 mm
wide along the costa; lower surface pale green,
veins slightly raised; upper surface of lamina
along sides of costae wrinkled when dry, probably
somewhat raised in living fronds; veins not raised
on upper surface except near base; sporangia
commonly 3-5, less often 6 or 7, not always
completely covering lower surface (as reported
in original description); numerous pale hairs,
longer than sporangia, attached to the receptacle.
Type: Elmer 11423 (H. Copel.; dupl. at K,
WS2eb, LP):
Distr. Malaysia: Philippines (Mindanao: Mt
Apo), once collected.
13. Gleichenia deflexa HoLTT. Reinwardtia 4 (1957)
addendum p. 280.
Scales on apex of main rachis light brown with
darker obliquely setose edges; branch rachises and
lower parts of costae at first covered on both
surfaces with a close felt of small setose scales and
stellate hairs and also with numerous narrow
scales (those on rachis 5 mm long, 0.2 mm wide),
setae of scales and hairs of upper surface all rigid
and rather dark, of smaller scales and hairs on
lower surface pale and lax; lower surface of
costules scaly and hairy near their bases like the
costae; lower surface of veins bearing persistent
scattered stellate hairs with slender pale rays.
Rachis-branches 120 cm or more long, fertile
ones to 35 cm wide, sterile sometimes only 18 cm
wide, all pinnules distinctly deflexed, making
angles of about 75° with the rachis; costae on
large fertile rachis-branches 414 cm apart, on
smaller sterile branches 214-314 cm apart; fertile
pinnules 16-20 cm long, widest part 3!4-3.8 cm
wide, costules 4-5 mm apart; /amina thin, veins
raised near their bases on both surfaces when dry;
20 or more pairs of segments on each larger pin-
nule constricted at the base, connected by a very
narrow wing along the costa, c. 314 mm wide
above the base, lowest segment sometimes quite
free and shortly stalked; sori of 3-5 sporangia;
stipular leaflets to 4 cm long, bipinnatifid, segments
c. 1 mm wide, acuminate.
Type: Brass 27171, Ferguson Isl. (L).
Distr. Malaysia: SE. New Guinea (Normanby
and Fergusson Islands).
Ecol. Scrambling to 7 m high in rather open
forest, 800-850 m.
14. Gleichenia sordida CopeL. in Elmer, Leafl.
Philip. Bot. 3 (1910) 798.—G. novoguineensis
BrRAusE, Bot. Jahrb. 56 (1920) 210.—Hicriopteris
novoguineensis COPEL. Philip. J. Sc. 75 (1941) 358.
—Hicriopteris astrotricha Cope. /.c.—Diplo-
pterygium novoguineense NAKAI, Bull. Nat. Sc.
Mus. Tokyo n. 29 (1950) 54.—G. sumatrana
Ho.tr. Reinwardtia 4 (1957) 264.
Scales on dormant apex of rachis 8-10 mm long,
0.5 mm wide, light brown with dark edges bearing
oblique setae; rachises and costae more or less
persistently covered with a felt of stellate hairs
(all dark brown or some rusty brown) and some
narrow scales to 4 mm long; lower surfaces of
costules and veins more or less persistently covered
with stellate hairs. Rachis-branches 120-200 cm
long, costae 214-414 cm apart; largest pinnules
15-20 cm long, 2!4-3 cm wide, slightly deflexed
or not; costules 314-414 mm apart; 2-8 basal
segments of lamina on each pinnule constricted
at the base and connected by a very narrow wing
along the costa, rest separated by narrow sinuses;
lamina rather thin, not glaucous beneath; veins
raised on lower surface, sometimes not on upper
surface; sori of 3-5 sporangia; stipular leaflets
broadly deltoid, 214-414 cm long, their lobes c.
1 mm wide.
Type: Elmer 11423a, Mt Apo, Mindanao (H.
Copel.; dupl. at US, BM).
Distr. Malaysia: Sumatra, Malaya, Celebes,
Moluccas (Halmaheira, Batjan, Morotai), Philip-
pines (Mindanao), New Guinea, and Solomons.
Ecol. High-climbing in open forest and on
forest-edge, apparently not forming thickets,
1000-2000 m.
15. Gleichenia norrisii METT. in Kuhn, Linnaea
36 (1869) 165; v. A. v. R. Handb. (1908) 58;
C. Cur. Gard. Bull. S.S. 7 (1934) 212, incl. var.
floccigera C. CHR.; Hottrt. Rev. Fl. Mal. 2 (1955)
67.—Hicriopteris norrisii CHING, Sunyatsenia 5
(1940) 280.—Diplopterygium norrisii NAKAI, Bull.
Nat. Sc. Mus. Tokyo n. 29 (1950) 54.—Fig. 6.
Scales on apex of main rachis 2-3 mm long,
narrow, medium brown, edges bearing short
oblique concolorous setae; costae and costules
when mature quite glabrous or bearing a few
scales near the base; young rachises, costae and
costules bearing scattered very narrow brown
scales, small setose scales and stellate hairs with
rusty brown rays; stellate hairs at first present on
veins but not persistent. Rachis-branches 100 cm
or more long; largest pinnules to 20 cm long and
3 cm wide, the costae 4-5 cm apart, distinctly
deflexed, distal pinnules more acutely deflexed
than basal ones; lobes of lowest pinnules usually
only slightly enlarged and not stipuliform, but
stipule-like leaflets, with broad blunt lobes,
sometimes produced; costules 414-514 mm apart;
lamina light green, drying light olive green without
reddish tinge, thin, lower surface more or less
glaucous in young plants, often not appreciably
so in older ones; segments of lamina only slightly
narrowed above the base, separated by sinuses
not over 1 mm wide, apices broadly rounded,
lowest segments much reduced but not separated
from the rest; veins slender, distinctly raised on
lower surface, slightly on upper surface; sori
FLORA MALESIANA [ser. Ui, -volaaie
Fig. 6. Gleichenia norrisii METT. on edge of forest, Penang, c. 650 m; one pair of rachis-branches showing
widely spaced and deflexed pinnules (HOLTTUM, 1926).
Dec. 1959]
Gleicheniaceae (Holttum) Sed : bz
commonly of 3-5 sporangia, with paraphyses
consisting of small scales bearing long, pale,
marginal hairs.
Type: Norris, Malay Peninsula (B?; dupl. at
K, BM, P).
Distr. Malaysia: Malay Peninsula, Sumatra
(Bencoolen), and N. Borneo.
Ecol. In clearings and on edge of forest, 650—
1250 m. On Penang Hill this species is abundant
at 650-750 m, below the lower limit of altitude of
G. longissima; above 750 m the latter, which forms
denser thickets because of its closer pinnules, is
the common species.
Note. CHRISTENSEN’S var. floccigera was based
on immature fronds which had not lost their
indumentum. The development of stipule-like
leaflets varies, but they never have the narrow
caudate lobes found in most other species.
16. Gleichenia matthewii HoLtr. Reinwardtia 4
(1957) 265.
Scales of main rachis-apex hardly 14 mm wide,
edges with their setae shining dark brown, middle
part paler; costae bearing a few scales near the
base, the frond otherwise glabrous. Rachis-
branches 70 cm or more long, strongly flexuous
above the basal part; all pinnules deflexed, the
upper ones at an angle to 45° so that each is in
line with the portion of the rachis beyond it, the
uppermost grading into the spreading lobes of the
terminal lamina; largest pinnules 12 cm long and
2'4 cm wide, the costae 4 cm apart; costules
414 mm apart; /amina firm, glaucous beneath,
lobed to 1-114 mm from the costa, segments
oblong, contiguous, almost truncate at the apex
and often slightly retuse; no sori seen; stipular
leaflets deltoid, 2 cm long, narrowly lobed.
Type: Matthew s.n., 31 Jan. 1912, G. Sing-
galang (K).
Distr. Malaysia: Central Sumatra (Mt Sing-
galang), twice collected, altitude 1800 m.
17. Gleichenia brevipinnula HoL_tr. Reinwardtia 4
(1957) 264.
Scales on apex of rachis c. 5 mm long, hardly
4 mm wide, brown, with almost black shining
edges bearing oblique setae; rachises rather
persistently covered with very dark rigid stellate
hairs with a few narrow scales; scattered similar
hairs on costae and costules beneath, and scattered
somewhat paler stellate hairs on lower surface
of veins. Main rachis 5 mm diameter; branches
70 cm long; costae c. 2 cm apart, pinnules to
10 cm long and 2 cm wide, not or slightly deflexed;
costules 3-314 mm apart; /amina cut down to
1 mm from the costa, very firm, sinuses very
narrow, edges of segments slightly toothed near
apices at the ends of the veins, lower surface
strongly glaucous; veins raised on both surfaces;
stipular leaflets 31 cm long, broadly deltoid, all
lobes with very narrow caudate tips, lower lobes
deeply and very obliquely lobed again; segments
of sub-basal pinnules often with caudate tips
like the stipular leaflets; sori of 3 or 4 sporangia.
Type: Bell 2042, Pueh Range, Sarawak (BM).
Distr. Malaysia: Borneo (Sarawak & N.Borneo).
Ecol. On ridges and summits of mountains,
in dwarf forest or scrub, 1250-2500 m. Fronds are
reported up to 214 m long.
18. Gleichenia blotiana C. Cur. Bull. Mus. Hist.
Nat. Paris Il, 6 (1934) 103.—Hicriopteris blotiana
CHING, Sunyatsenia 5 (1940) 279.—Diploptery-
gium blotianum NAKAI, Bull. Nat. Sc. Mus. Tokyo
n. 29 (1950) 49.
Scales on dormant apex of rachis nearly black,
edges bearing oblique black setae; scales on
growing main rachis usually with a paler median
band; rachis-branches when young bearing many
dark stellate hairs with rigid rays (so close that
hairs touch each other but do not completely
obscure surface of rachis) and also very narrow
ciliate scales 2 mm long and some long lax multi-
cellular hairs; a similar indumentum on upper
surface of costae at first, later deciduous; in-
dumentum on lower surface of costae and
costules mostly of dark rigid stellate hairs, with
a few scales at bases of costae; dark stellate hairs
scattered abundantly on veins and lamina beneath,
persistent. Main rachis-branches c. 150 cm long,
the lowest pinnules stipule-like, 314 cm long,
deltoid, with very narrow deeply pinnatifid basal
lobes; pinnules commonly 20 cm long, 3-4 cm
wide, the base of the costa distinctly deflexed as
in G. norrisii; costae 4-5 cm apart, costules c.
5 mm apart; segments of lamina 3!4-4 mm wide,
rather thin, their apices broadly rounded and
sometimes retuse; veins distinctly raised on upper
surface and slightly so below; sporangia 3-4 in
each sorus.
Type: Pételot 3900, near Chapa, Indo-China
(BM; dupl. at P).
Distr. Indo-China and S. China, in Malaysia:
Malay Peninsula, once collected.
Ecol. In the Malay Peninsula on edge of forest
in a thicket with G. Jongissima, at 1500 m.
19. Gleichenia conversa v. A. v. R. Bull. Jard.
Bot. Btzg II, ». 20 (1915) 17; Handb. Suppl.
(1917) 81; BACKER & Postu. Varenfl. Java (1939)
255.—Diplopterygium conversum NAKAI, Bull.
Nat. Sc. Mus. Tokyo n. 29 (1950) 49.—Fig. 4h-j.
Differs from G. norrisii as follows: scales on
rachis-apex brown with almost black edges; some
rather light brown stellate hairs often persistent
on costules and sides of costae; largest pinnules
to 25 cm long and 4 cm wide; distal pinnules not
appreciably deflexed; segments of lamina separated
by sinuses about 2 mm wide; lamina and rachises
drying rather red-brown, lamina usually thicker
than in G. norrisii; veins usually not at all raised
on upper surface (slightly so in specimens with
thin lamina); stipular leaflets well developed, the
lobes narrow.
Type: Hasskarl, Kandang Badak, G. Gedeh
(Bo; dupl. at L).
Distr. South Malaysia: Java and Lesser Sunda
Islands (Flores).
Ecol. In open places in forest and on forest-
edges, 900—2400 m.
18 FLORA MALESIANA [ser. II, vol. 1?
3. Subgenus Mertensia
Hook. Sp. Fil. 1 (1844) 6, pro § 2; Hottr. Reinwardtia 4 (1957) 266.—Mertensia
WILLD. Kongl. Vet. Ak. Nya Handl. 25 (1804) 163, p.p., non RotH 1797.—
Gleichenia subg. Mertensia sect. Holopterygium Dies in E. & P. Pfl. Fam. 1, 4
(1900) 353.—Dicranopteris (non BERNH.) UND. Bull. Torr. Bot. Cl. 34 (1907) 249,
p.p.—-Gleichenia subg. Mertensia sect. Dicranopteris v. A. v. R. Handb. (1908) 56,
p.p.—Sticherus PRESL, Tent. Pterid. (1836) 51; CHING, Sunyatsenia 5 (1940) 281;
CopeL. Gen. Fil. (1947) 27; NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 7.—
Sticherus § Eu-Sticherus C. Cur. in Verdoorn, Man. Pterid. (1938) 530.—
Mesosorus Hassk. Fil. Jav. 1 (1856) 2, p.p.—F ig. 7-10.
Primary rachis-branches each ending in a dormant apex which lies in the angle
between a pair of secondary branches; secondary branches behaving similarly,
the process usually repeated to produce pseudo-dichotomous branching of several
orders; ultimate branches simply deeply pinnatifid, the venation of segments of the
lamina and sori as in subg. Diplopterygium; penultimate branches (often also
+
TTT
Fig. 7. Diagrams showing branching habit of species of Gleichenia subg. Mertensia. a. G. truncata
(WILLD.) Spr., b. G. milnei BAK., c. G. hispida METT., d. G. hirta BL., e. G. vestita BL.
Bect959] or) Gleicheniaceae (Holttum) 19
branches of lower orders) more or less completely provided with a deeply pinnatifid
lamina like that of the ultimate branches.
Notes. This subgenus includes far more species than any other major division of the family, in
all continents, and its distribution is mainly south of the equator. Most of the species are rather small
in size, as compared with representatives of subg. Diplopterygium, and could not compete as thicket-
forming ferns either with the latter or with Dicranopteris. The only Malaysian species which can so
compete is G. truncata, and it is also the only species of subg. Mertensia which occurs at sea-level in our
region. The others are mountain plants occurring in scrub and dwarf forest of high ridges or in open
grassy or rocky places, which are comparable with the habitats of related species in south temperate
regions.
As in other divisions of the family, the characters and distribution of scales and hairs are important
diagnostically in this subgenus. The number of times the lateral branch-systems are forked (the number
of orders of forking) is probably important, but shows considerable variation within a species ac-
cording to the age of the plant and to environmental conditions, and these cannot be fully judged
from dried specimens. A more important kind of character is the relative length of branches of the
first and ultimate orders. But in some species one frond will have branch-systems forked to 2 orders
with long ultimate branches, other fronds (even part of the same frond) may be forked to 3 orders with
much shorter ultimate branches. The glaucous character of the lower surface of the lamina may be
significant, but is easily destroyed by heat in drying. In all these characters, and in all species, more
field study is needed to establish distinctions between species more clearly.
Judging from anatomical characters, CHRYSLER (Am. J. Bot. 31, 1944, 483-491) has argued that subg.
(eer ‘oF
Fig. 8. Gleichenia truncata (WILLD.) Spr. a. Stipular leaflets at main branching, nat. size, b. lower
surface of part of frond, x 4. — G. vestita BL., c. Lower surface of a large segment of lamina, x 4,
d. a single scale from costa, x 8. — G. hispida METT., e. Lower surface of part of frond, showing sori
and hair-pointed scales, x 8, f. same with scales and sporangia removed, g. a single scale from costa,
xX 22. -G. hirta BL. h. Lower surface of part of frond, x 4, i. tip of a segment of the lamina, showing
toothed edge, j. a scale from the costa, x 20.
20 FLORA MALESIANA
[ser. IL -yvolaat
Mertensia is the most primitive division of the genus. But from the point of view of sporangia and
sori it is exactly in equality with subg. Diplopterygium, and in leaf-form it is clearly more highly or-
ganized (in having a branching-pattern dependent on a series of permanently dormant apices). It seems
to me significant that among fossils none are clearly referable to this subgenus, whereas there are abun-
dant fossils having a close resemblance to subg. Diplopterygium. An apparent exception is Gleichenites
gracilis ZIGNO. But the branching-pattern of this fossil is very irregular and there is no clear evidence
of pseudo-dichotomy; I think that its resemblance to subg. Mertensia is superficial.
20. Gleichenia reflexipinnula C. Cur. Brittonia 2
(1937) 27.—Sticherus reflexipinnula CoPEL. Philip.
Ie SeseaCl94)) 355%
Rhizome 5 mm diameter, covered when young
with dark brown shining acuminate scales hardly
1 mm wide at the base, when old closely warty;
stipe 30 cm or more long, scaly and then warty
like the rhizome near the base; main rachis bearing
2 or more pairs of primary branches, with deeply
pinnatifid stipular leaflets 2-4 cm long at bases of
branches; primary branches once or _ twice
forked, leafy down to the junction with the main
rachis, not proliferous beyond the first fork;
angle of ultimate forks rather less than a right
angle; ultimate branches to c. 20 cm long and
314-4 cm wide (shorter if primary branch is
twice forked), penultimate branches 314-9 cm
long; scales on resting apex of main rachis 314 mm
long, less than 1 mm wide, acuminate but not hair-
pointed, rather thin, medium brown with short
sparse fringe of paler hairs; scales on main rachis
persistent, abundant, thin, pale and long-fringed;
scales on lower surface of costae abundant,
largest 2 mm long, 14 mm wide, thin, pale, long-
fringed; scales on costules similar and smaller,
abundant, ranging down to stellate hairs; no
stellate hairs on veins; /Jamina lobed almost to
the costa, middle and upper segments slightly
deflexed, lower ones much deflexed, costules
3144 mm apart; segments gradually narrowed to
apex (which is sometimes retuse), coriaceous,
edges strongly revolute when dry, veins hardly
raised on either surface; sporangia usually 4, sori
close together.
Type: Brass 4847, Mt Tafa (BM; dupl. at Bo,
US).
Distr. Malaysia: E. New Guinea (Mt Tafa,
Mt Dayman).
Ecol. Forest-edge, subscandent to 2 m (Mt
Dayman), plentiful in summit clearing (Mt Tafa),
2230-2700 m.
21. Gleichenia venosa (CoPEL.) HOLTT., comb. nov.
—Sticherus venosus CoPEL. Philip. J. Sc. 75 (1941)
3565 ple 5.
Habit of G. truncata and of similar size (rachis-
branches of second order sometimes only partly
leafy); segments of lamina thicker, with edges
sometimes much reflexed when dried, the veins
on the lower surface strongly raised; scales of
apex of main rachis similar to those of G. truncata;
scales on costae abundant and persistent, dark
rusty-brown, to 1144 mm long and more than
4 mm wide, fringed; scales on costules similar
but smaller, very abundant, spreading and closely
overlapping so as completely to cover lower
surface of lamina; smaller scales also present
throughout on rachises, persistent as a continuous
rusty covering at and just below the forks.
Type: Brass 12348, Idenburg River (H. Copel.;
dupl. at Bo, BM, L).
Distr. Malaysia: New Guinea.
Ecol. Thickets in open places, scrambling to
2-3 m, 1600-2700 m.
22. Gleichenia truncata (WILLD.) Spr. Syst. Veg.
ed. 16, 4 (1827) 25; HoLtr. Reinwardtia 4 (1957)
271.—See for further synonyms under the varieties.
—Fig. 7a, 8a—b, 9.
KEY TO THE VARTERIESs
1. Additional stipular leaflets present about 1 cm
below main and first lateral forks of rachis, as
well as just above forks 2. var. bracteata
1. Such additional stipular leaflets lacking.
2. Costae persistently densely scaly on the upper
Giighes 5 So 6 5 5 4. var. involuta
2. Costae at most sparingly and not persistently
scaly on the upper surface.
3. Main rachis-branches several times forked.
Scales on costae beneath very small or nar-
row and acuminate.
4. Scales on costae sparse and very small.
1. var. truncata
4. Scales on costae 114-2 mm long, 0.3-0.6
mm wide, acuminate 5. var. celebica
3. Main rachis-branches 2 or 3 times forked.
Scales on costae beneath rather abundant,
1-114 mm long, 0.7—1 mm wide.
3. var. plumaeformis
1. var. truncata.—Mertensia truncata WILLD.
Kongl. Vet. Ak. Nya Handl. 25 (1804) 169,
t. V, f. A.—Mertensia laevigata WILLD. Sp. Pl. 5
(1810) 75.—Sticherus laevigatus PRESL, Tent.
Pterid. (1836) 52.—G. laevigata Hook. Sp. Fil. 1
(1844) 10; Racts. Fl. Btzg 1 (1898) 11; v. A. v. R.
Handb. (1908) 59; BAcKER & PostH. Varenfi.
Java (1939) 255; Hoxtr. Rev. Fl. Mal. 2 (1955)
71.—Sticherus truncatus NAKAI, Bull. Nat. Sc.
Mus. Tokyo n. 29 (1950) 20.—Sticherus myriapoda
NAKAI Zc) 125 f 12
Main rachis 8 mm or more diameter near the
base, often high-climbing with many pairs of
branches; primary rachis-branches several times
almost equally forked, the angle of forking about
a right angle, of ultimate forks sometimes more
than 90°; basal segment of primary rachis-branch
leafless (apart from stipular leaflets), branches of
second and later orders usually all leafy, length
of each branch from one fork to the next commonly
10—15(—20) cm; stipular leaflets usually present at
base of primary branch and at its first fork,
Dec. 1959]
a4 TS y
sae ie
’
Gleicheniaceae (Holttum) 21
Fig. 9. Gleichenia truncata (WILLD.) SPR. on Taiping Hills, Malaya, by roadside, c. 1200 m (HOLTTUM,
1952).
deltoid and deeply lobed (basal lobes again lobed),
at base of primary rachis-branch to 3 cm long,
at next fork 114 cm long; lamina cut almost to
the costa into lobes at right angles to the costa;
lobes rather irregular in length even on same
branch, longest on lowest branches, commonly
2-314 cm long (to 5 cm) and about 2!4 mm wide
(to 3 mm), costules 3-4 mm apart; lower surface
not glaucous, veins dark when dry, slightly raised;
veins on upper surface not raised when dry,
concolorous; sori of 3-5 sporangia, surrounded by
pale hairs. Scales on smaller dormant apices
mostly very small, often not more than 1 mm
long, rusty brown, of varying shape, fringed; on
larger apices also some very narrow scales; sca-
les on costae very small, long-fringed, at first
rather abundant; very short simple hairs, often
shrivelled and black when old, present on lower
surface of costules, veins and lamina.
Type: Herb. Willd. (B).
Distr. Throughout Malaysia, apparently most
abundant in the west.
Ecol. On edges of forest, often climbing to a
considerable height, 0-1600 m, growing with
varieties of Dicranopteris linearis, also in thickets
at higher altitudes.
Note. The original specimen of Mertensia
truncata WILLD. had incompletely expanded
lamina-lobes, which thus appeared to be truncate.
Sticherus myriapoda NAKAI was described from
a plant in the Riouw Archipelago differing only
from typical var. truncata in having a twining
main rachis. Such a condition has also been
observed in one plant in Johore, but is not general
in this variety. Experimental culture is desirable
to decide whether the twining condition is genetic-
ally controlled. The Johore plant had rachises
twining both to the left and to right, and the
length of rachis making one complete turn round
the support was about 20 cm. NAKAI described a
closer spiral.
2. var. bracteata (BL. ex Hook. & BAKER)
Ho.tr. Reinwardtia 4 (1957) 271.—G. bracteata
BL. ex Hoox. & BAKER, Syn. Fil. (1865) 14 (as
synonym but with descr.)—G. /aevigata var.
bracteata v. A. v. R. Handb. Suppl. (1917) 85.
Lamina-segments commonly less than 2 mm
wide, 10-18 mm long; additional stipular leafiets
present about 1 cm below main and first lateral
forks of rachis, on the outside.
Type: Blume, Java (L; dupl. at K).
Distr. Malaysia: Java, Flores, S. Celebes.
Ecol. On edges of forest, apparently abundant
at 1000-1600 m; extremes of altitude recorded
600 and 2000 m.
3. var. plumaeformis (PRESL) HOoLTr. Rein-
wardtia 4 (1957) 272.—Mertensia plumaeformis
PresL, Abh. (K.) B6hm. Ges. Wiss. M.-N. Cl. V,
22 FLORA MALESIANA
5 (1848) 338; Epim. Bot. (1851) 24, t. 15.—
Sticherus plumaeformis NAKAI, Bull. Nat. Soc. Mus.
Tokyo n. 29 (1950) 25.
Primary rachis-branches once or twice forked,
3-6 cm to first fork ‘this part almost or quite
leafless), ultimate branches up to 25-35 cm long;
costae usually bearing rather numerous, thin,
broad, almost entire scales 1-144 mm long,
0.7-1 mm wide, on lower surface.
Type: Cuming 377, Malacca (PRc; dupl. at K,
BM;)P) EE):
Distr. Malaysia: Sumatra, Malay Peninsula,
Borneo (?).
Ecol. In open places on mountains, 1200-1800
m. As seen by me in W. Sumatra this variety does
not form long climbing fronds.
4. var. involuta HoLTT. Reinwardtia 4 (1957) 272.
Primary rachis-branches 3 times forked; ultimate
branches sometimes much longer than lower ones;
ultimate forks forming an angle of rather less
than 90°; upper surfaces of costae densely scaly,
scales persistent, 1 mm long, pale-fringed; lower
surfaces of costae bearing scales ¥% mm long;
edges of dried Jamina-lobes much revolute.
Type: J. Winkler; Rosenst. Fil. Sum. Exsic. 182
(L; dupl. at BM, P).
Distr. Malaysia: N. Sumatra (Karo Plateau).
5. var. celebica HOLTT., var. nov.
Primary rachis-branches 3 times equally forked,
ultimate and penultimate branches leafy, angle of
ultimate forks less than a right angle, segments
of lamina c. 2 cm long; sca/es on lower surface
of costae copious, acuminate, fringed, c. 114-2
mm long, 0.3-0.6 mm wide.
Distr. Malaysia: Central
2600-3000 m (once collected).
Celebes, altitude
23. Gleichenia milnei BAKER, Syn. Fil. (1874) 449;
Hook. Ic. Pl. t. 1602; HoL_tr. Reinwardtia 4 (1957)
270.—G. kajewskii Core. Philip. J. Sc. 60 (1936)
102, pl. 6.—Sticherus milnei CHING, Sunyatsenia
5 (1940) 284.—Sticherus kajewskii CopEL. Gen.
Fil. (1947) 27.—Fig. 7b.
General aspect of frond as in G. truncata (in its
wide forkings and narrow lamina-segments at
right angles to the costa), but at each fork (apart
from forks of main rachis) one branch larger
than the other, larger branches alternately to left
and right and successive larger branches almost
in line with each other; ultimate branches, and
branches of 2—3 lower orders leafy; leafy branches
in general 214-314 cm wide (lower ones to 414
cm), lamina-segments 2-3 mm wide, costules
3-5 mm apart; scales as in G. truncata; additional
stipular leaflets often present outside the lower
forks, as in G. truncata var. bracteata but nearer
the forks.
Type: Milne 341, New Hebrides (K).
Distr. Malaysia: Celebes,? Philippines (?Luzon),
Moluccas, New Guinea, Admiralty Islands,
Solomon Islands, New Hebrides.
Ecol. Altitude 100-1400 m.
[ser. II, vol. 11
24. Gleichenia oceanica KUHN, Verh. K. K. Zool.-
Bot. Ges. Wien 19 (1869) 583; non CuHRistT, Ann.
Jard. Bot. Btzg 15 (1898) 76; v. A. v. R. Handb.
(1908) 60; Suppl. (1917) 85, 497; CopeL. Univ.
Cal. Publ. Bot. 12 (1931) 388.—Sticherus oceanicus
St JOHN, Occ. Pap. Bish. Mus. 17 (1942) 81.
Habit and scaliness of G. truncata, but the
forkings of the rachis-branches much less than a
right angle (ultimate forks often less than 45°),
the lamina glaucous beneath, its segments distinct-
ly oblique and commonly less than 2 cm long
(often not more than 114 cm); primary rachis-
branches of large fronds 5 times forked, with
deltoid stipular leaflets at their base, of smaller
fronds sometimes only 3 times forked and lacking
stipular leaflets.
Type: Herus 66, Aneityum, New Hebrides (B).
Distr. Melanesia (New Hebrides, Fiji, Samoa),
may occur in East Malaysia.
Note. The SARASIN specimens from Celebes
referred to this species by CHRisT (/.c.) are G.
hispida and G. hirta var. paleacea; 1 have not
seen the WARBURG specimen.
25. Gleichenia erecta C. Cur. Brittonia 2 (1937)
269; HoLtr. Reinwardtia 4 (1957) 268.—Stricherus
erectus COPEL. Philip. J. Sc. 75 (1941) 353.—
Sticherus habbemensis Cope. l.c. 355, pl. 3.
Rhizome 2!4-4 mm diameter, young parts
covered with thin brown ciliate scales to 3 mm
long and | mm wide, old parts closely warty;
stipe at first scaly, then warty, near the base, like
the rhizome, rest smooth, 12-40 cm long; frond
bearing one or more pairs of primary branches
which are simple or once or twice forked, sometimes
proliferous beyond the first pair of secondary
branches; if twice forked, the first-order branch
2-3 cm long, bearing a stipular lobed leaflet 15 mm
long and sometimes | or 2 other lamina-segments
but not fully leafy, the second-order branch 3-6
cm long, fully leafy; angle of ultimate forks less
than 45°; ultimate branches 12-30 cm _ long,
largest /amina-segments commonly 15-23 mm long,
sometimes to 40 mm, rather coriaceous, entire,
edges much reflexed when dry, abruptly narrowed
just above the base, the rest 2—-2'4 mm wide
when flattened, tapering to apex, lower surface
glaucous; veins not raised on either surface;
costules 3-4 mm apart (lowest ones on large
fronds to 5 mm apart), basal part at 60—70° to
the costa, distal part often faleate when dry;
scales on resting apices of main rachis like those
on base of stipe but more finely fringed; some
persistent scales on lower surface of costae, 1!
mm long, 14 mm wide, shortly hair-pointed, edges
fringed at least towards the apex (these scales
sometimes very pale with a darker base); scales
on costules very small and long-fringed; pale-
rayed stellate hairs on veins, their red-brown bases
sometimes persistent; sori of 3-4 sporangia.
Type: Brass 4591, Murray Pass, Wharton Range,
Papua (BM; dupl. at K, Bo, US).
Distr. Malaysia: New Guinea.
Ecol. In open places, in peaty grassland, and
on wet clay-soil of land-slips, 1000-3225 m.
Dec. 1959]
26. Gleichenia pulchra (CopeL.) Ho.ttr. Rein-
wardtia 4 (1957) 271.—Sticherus pulcher COPEL.
Philips. Sc: 75° (1941).355, pl. 4.
Main rachis slender, bearing several pairs of
branches, each primary branch bearing up to at
least 3 pairs of secondary branches, each secondary
branch simple or once forked; only ultimate
branches leafy, except sometimes where a second-
ary branch is forked, in which case the penul-
timate branch may bear a few lamina-lobes;
ultimate branches to 20 cm long and 2!4 cm wide
(to 3 cm wide if a simple secondary branch),
lamina lobed almost to the costa, segments very
slightly oblique, 3 mm wide above the base,
costules 4 mm apart; veins slightly raised on lower
surface, on upper surface only at the base of each
vein; lower surface glaucous when living (not
clearly so on dried specimens available); scales
on main dormant apices small, on lower surface
of costae 14-1 mm long, firm, dark brown with
a pale fringe; scales on lower surface of costules
abundant, small, long-fringed; many small rusty-
brown stellate hairs on lower surface of veins and
on lamina.
Type: Brass 12351, Idenburg River (H. Copel.;
dupl. at BM, L).
Distr. Malaysia: New Guinea (NW. part, once
collected, SE. part, doubtful).
Ecol. Scrambling in mossy forest, 1650 m.
27. Gleichenia pseudoscandens v. A. v. R. Nova
Guinea 14 (1924) 24.—Sticherus pseudoscandens
CopEL. Philip. J. Sc. 75 (1941) 356.
Fronds to 4 m long, with several pairs of
primary branches; primary branch-systems usually
with 3 orders of forking, the primary branch
sometimes proliferous beyond its first pairs of
branches, only the ultimate branches leafy; ulti-
mate branches to 40 cm long, lower ones 10—20
cm long; costules 5'4-6 mm apart, at right angles
to the costa or nearly so; segments of lamina
17-25 mm long, 3 mm wide above the base, /amina
firm, drying dark, veins not raised on either sur-
face; scales on resting apex of main rachis 4-5
mm long, rigid, dark, very narrow, with short
oblique marginal setae; scales on lower surface
of costae dark and small, almost entire, mostly
deciduous leaving the surface warty; very short
hairs copious on lower surface of lamina and on
costules; some hairs with the sporangia.
Type: Lam 1935, Doorman summit (U; dupl.
at L, Bo, BM).
Distr. Malaysia: NW. New Guinea, once
collected.
Ecol. Altitude 2480 m.
28. Gleichenia alstonii HoL_tr. Reinwardtia 4
(1957) 267.
Main rachis 4 mm diameter; primary branch-
systems usually with forking of two orders, only
the ultimate branches leafy; primary branches
rarely proliferous beyond the first fork; branches
of first order 4-5 cm long, of second order 6-7
em long; ultimate branches 30-35 cm _ long,
314 cm wide, the basal 10-15 mm sometimes
Gleicheniaceae (Holttum) 23
bare of lamina; costules 4 mm apart, slightly
oblique; segments of /amina c. 17 mm long, 3 mm
wide above the base, thin, apices entire or nearly
so, veins slightly prominent on both surfaces; sori
of 3-4 sporangia. Scales on dormant apex of
rachis 4 mm long, 0.7 mm wide, narrowly acumi-
nate, brown, with short spreading pale marginal
hairs near base, dark oblique sparse setae towards
apex; scales on lower surface of costae scattered,
dark, 1144 mm long, 0.2 mm wide, edges with
sparse oblique dark hairs; scales on lower surface
of costules pale, very narrow; scattered pale lax
stellate hairs on lower surface of veins.
Type: Alston 14981, Lae Pondom (BM).
Distr. Malaysia: N. Sumatra (Tapanuli), once
collected.
Fig. 10. Gleichenia vestita BL. on ridge with ericoid
ridge forest from Poka Pindjang to Rante Mario
in the Latimodjong Range, Central Celebes,
c. 3000 m (1937).
29. Gleichenia vestita BL. En. Pl. Jav. (1828) 249;
Racis. Fl. Btzg 1 (1898) 11; v. A. v. R. Handb.
(1908) 61; Suppl. (1917) 86; BACKER & PosTH.
Varenfl. Java (1939) 255.—Sticherus vestitus
CHING, Sunyatsenia 5 (1940) 285.—Fig. Te,
8c—d, 10.
Rachis of a well-grown frond bearing several
24 FLORA MALESIANA
[ser.: Il, ovolal
pairs of primary branches, usually without stipular
leaflets at their base; primary branch-systems
usually forked to 2 orders (rarely 3), primary
branch not proliferous beyond its first fork; all
branches leafy, down to junction with main
rachis; angle of forks about 45°; ultimate branches
15-25 cm long, lower ones 5—7 cm long; /amina
cut down to within 1 mm of costa, segments
distinctly oblique, thin but very rigid, 8-12 mm long,
not narrowed immediately above the base but
tapering very gradually, then abruptly to broadly
pointed or rounded entire apex, edges not usually
much reflexed, lower surface glaucous; costules
4-5 mm apart; veins slightly raised on both
surfaces; sori of 3-5 sporangia, without evident
paraphyses. Scales on dormant apex of main
rachis firm, rusty to rather dark brown, 4-5 mm
long, more than | mm wide, gradually narrowed
to hair point, edges rather sparsely obliquely
ciliate towards apex; scales on lower surface of
costae abundant and persistent, spreading, rusty
brown, 0.6-1.2 mm wide, 2—3 mm long, ciliate
towards hair-pointed apex; smaller scales few,
stellate hairs quite lacking; very short reddish
simple hairs frequent on costules and veins
beneath.
Type: Blume, Java (L; dupl. at K).
Distr. Malaysia: Sumatra, Java, S. Celebes.
Ecol. In open alpine forest and scrub, 2300-—
2800 m.
Note. At high elevations and in exposed places,
fronds may be smaller than above described, with
branches of first and second order only 2-3 cm
long.
30. Gleichenia bolanica RosENsT. in Fedde, Rep.
12 (1913) 162; v. A. v. R. Handb. Suppl. (1917)
86.—G. monticola RiDL. Trans. Linn. Soc. Bot.
9 (1916) 252.—G. subulata v. A. v. R. Nova
Guinea 14 (1924) 23.—Sticherus bolanicus COPEL.
Philip. J. Sc. 75 (1941) 352.—Sticherus monticola
NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 23.
Lateral branch-systems of well-grown fronds
with forks to 4th order; primary branches not
normally proliferous beyond first fork nor bearing
stipular leaflets at their base; ultimate branches
7-20 cm long, 6-10 mm wide, lower branches 1-2
cm long; usually only ultimate and penultimate
branches leafy, but lower ones sometimes partly
leafy; /amina cut almost down to the costa,
segments apparently triangular because of their
much-reflexed edges, 3 mm wide at the base,
rather thick and very rigid when dry, lower sur-
face glaucous with slightly raised veins; costules
slightly oblique; sori often very crowded, of 3-6
sporangia. Scales on dormant apex of main
rachis rigid, dark and shining, 3-4 mm long, to
0.7 mm wide, apex not hair-pointed, marginal
hairs short, oblique, rigid; scales on lower surface
of costae abundant, to 2 mm long, not hair-
pointed, base 0.50.7 mm wide, brown at base and
pale distally, edges bearing short hairs which are
sometimes very rigid and dark brown; small
similar scales or stellate hairs on costules and veins.
Type: Keysser B. 14, Mt Bolan, New Guinea (B).
Distr. Malaysia: New Guinea.
Ecol. In open scrub or rocky places, fronds
to 2 m or more long, 2800-3950 m.
31. Gleichenia hispida Metr. ex KUHN, Verh.
K. K. Zool.-Bot. Ges. Wien 25 (1875) 600;
RAcis. Fl. Btzg 1 (1898) 12; v. A. v. R. Handb.
(1908) 61; BACKER & PostH. Varenfl. Java (1939)
255; Hoxtr. Reinwardtia 40 (1957) 270.—G.
koordersii CuHrRist, Ann. Jard. Bot. Btzg 15
(1898) 76, pl. 13, f. 1.—Sticherus caudatus COPEL.
Philip. J. Sc. 75 (1941) 354, pl. 2.—Sticherus
hispidus Corer. Gen. Fil. (1947) 27.—Sticherus
pinnatus COPEL. Philip. J. Sc. 83 (1954) 98, pl. 3.—
Fig. 7c, 8e-g.
Well-grown fronds with several pairs of primary
branches which are not proliferous beyond their
first fork and have no stipular leaflets; \ateral
branch-systems with forks to 2nd or 3rd order;
branches of first order leafless or only partly
leafy, rest fully leafy; first order branches 214-6
cm long, second order branches 5-12 cm long;
from second-order fork to end of ultimate branches
(whether 2 or 3 forks present) 20-30 cm, the
third fork (if present) about the middle of this
length; /amina cut almost down to the costa, the
segments slightly oblique, coriaceous, stiff and
rigid when dry with much-recurved edges, longest
segments on lower branches to 3!4 cm long, on
ultimate branches sometimes only | cm long,
effective width when dry 114-214 mm, costules
3-4 mm apart; lower surface of lamina more or
less glaucous, veins dark and slightly prominent,
not raised on upper surface; soriof 3 or 4sporangia
with rather long crisped rusty hairs. Scales on
dormant apex of main rachis rusty brown, 5-7
mm long, 0.5 mm wide, narrowed above the base
and for the most part very narrow, hair-pointed;
scales on lower surface of costae only a few cells
wide at the base, the hair-point forming at least
half of the length, hair-points crisped and entangled
with those of other scales; scales on costules
similar and smaller, grading to stellate hairs on
veins and edges.
Type: Jagor 558, G. Galungung (B; not seen).
Syntype: De Vries, Ternate (dupl. at U).
Distr. Malaysia: Sumatra, Java, Bali, N. Ce-
lebes, Ternate, and Philippines (Negros, Luzon).
Ecol. In thickets in open places, 1000-2200 m.
Note. Isolated plants in exposed places may
have fertile fronds much smaller and less branched
than those described; Sticherus pinnatus COPEL.
appears to represent a condition of this species
with unbranched fertile fronds.
32. Gleichenia brassii C. Cur. Brittonia 2 (1937)
271.—Sticherus brassii NAKAI, Bull. Nat. Sc. Mus.
Tokyo n. 29 (1950) 15.
Rhizome 6 mm diameter, strongly warty from
bases of old scales, young parts scaly, scales 5 mm
long, 1 mm wide, ciliate; stipes to 100 cm long,
persistently scaly throughout but only warty
near the base, scales of all sizes with rather copious
pale fringe, the longest ones hair-pointed; main
rachis to about 200 cm long, bearing several pairs
Dec. 1959]
Gleicheniaceae (Holttum) 25
of primary branches which lack stipular leaflets;
lateral branch-systems with forks of 2 orders,
ultimate and penultimate branches fully leafy,
ultimate forks less than 90°; first and second order
branches 12-15 cm long, ultimate branches
30-40 cm long and 4-414 cm wide; costules about
5 mm apart, at right angles to costae; segments of
lamina 3 mm wide above the base, firm, edges
more or less revolute when dry, lower surface not
glaucous when dry; veins slightly prominent on
lower surface, not on upper surface except their
bases; sori of 3-4 sporangia without evident para-
physes. Scales on dormant apex of main rachis
like those on rhizome but thinner, edge shortly
ciliate, apex a short hair; scales on lower surface
of costae spreading, 114-2 mm long, 0.5 mm wide,
thin, brown, darker at the base, shortly hair-
pointed, closely fringed with spreading pale hairs;
scales on lower surface of costules abundant,
14-1 mm long, rusty brown with long pale fringe;
upper surface of costae raised and terete, bearing
persistent small pale long-fringed scales, rest of
upper surface glabrous; lower surface of veins
bearing many very short simple hairs.
Type: Brass 4922, Mt Tafa (BM; dupl. at Bo).
Distr. Malaysia: New Guinea (Mts Arfak and
Tafa).
Ecol. Scrambling in open places in forest,
1550-2400 m.
33. Gleichenia loheri CHRIST, Bull. Herb. Boiss. II,
6 (1906) 1009; v. A. v. R. Handb. (1908) 796;
Ho.ttr. Reinwardtia 4 (1957) 272.—Sticherus
loheri CoreL. Gen. Fil. (1947) 27.—Sticherus
perpaleaceus CopeEL. Philip. J. Sc. 81 (1952) 3.
var. loheri.
Lateral branch-systems with forks of 1, 2 or 3
orders, ultimate and penultimate branches fully
leafy, angle of ultimate fork much less than a right
angle; with forks of 1 or 2 orders, penultimate
branches 5—7 cm long, ultimate branches 17-23
cm long; with forks of 3 orders, ultimate and
penultimate branches together 17-20 cm long;
ultimate branches 114-2.2 cm wide, costules
3-314 mm apart, at right angles to costa or very
slightly oblique, /amina cut almost down to the
costa, segments not abruptly narrowed above the
base, edges when dry somewhat reflexed, apex
rounded and entire, texture firm but not coriaceous,
veins distinctly raised on lower surface, slightly
so above, lower surface not glaucous when dry;
sori of 3-4 sporangia. Scales on apex of main
rachis 5 mm long, 1 mm wide, thin, medium rusty
brown, hair-pointed, edges with oblique slender
hairs; scales on lower surface of costae copious
and spreading, the largest 2 mm long, 0.8 mm wide,
shortly hair-pointed; scales on costules very small,
long-fringed, pale, mixed with long-armed stellate
hairs; upper surface of costae raised and terete,
more or less persistently covered with small pale
long-fringed interlacing scales; very short simple
hairs abundant on lower surface of veins and
occasionally also on lamina.
Type: Loher s.n., 7 Feb. 1906, Mt Banaho (P).
Distr. Malaysia: Philippines (Luzon).
var. major HoLtr. Reinwardtia 4 (1957) 272.
Larger than typical form of species; ultimate
branches 20-30 cm long, 2'4—31/4 cm wide, costules
3-4 mm apart; 3-lobed stipular leaflets 15 mm long
sometimes present.
Type: Biinnemeijer 11965, Celebes (Bo; dupl.
Alea).
Distr. Malaysia: S. Celebes, N. Borneo, and
Philippines (Negros).
Ecol. Altitude 2000-2500 m.
34. Gleichenia flabellata R. Br. Prod. (1810) 161;
Hook. Sp. Fil. 1 (1844) 6; Domin, Bibl. Bot. 85
(1915) 204; v. A. v. R. Handb. (1908) 60; K. Scu.
& Laur. Fl. Schutzgeb. (1901) 144.—Sticherus
flabellatus H. St. JOHN, Occ. Pap. Bish. Mus. 17
(1942) 81.
Stipe to 60 cm long, scaly at the base; main
rachis bearing 2 or more pairs of branches;
lateral branch-systems compact, forked to 3
orders at short intervals (lower branches com-
monly 1 cm long, penultimate 2—3 cm long), ulti-
mate and penultimate branches leafy, angle of
forks less than 45°; ultimate branches 12-15 cm
long, 2—3(-4) cm wide; costules 4 mm apart, at
an angle of 45° to the costa; /amina cut almost
to the costa, segments 214 mm wide above the
base, edges toothed almost to the base, apex
broadly pointed, veins very oblique and raised
on both surfaces, lower surface not glaucous.
Scales on apex of main rachis 4 mm long, shining
medium brown, acuminate, fringe copious, long,
lax; scales on lower surface of costae few and
persistent only near base, narrow, hair-pointed
and long-fringed; smaller scales or long-armed
pale stellate hairs occasional on costules near base
of branch, surfaces otherwise glabrous.
Type: Robert Brown, Port Jackson, N. S.
Wales (BM; dupl. at K).
Distr. E. & SE. Australia, New Caledonia,
New Zealand, in Malaysia: E. New Guinea
(SCHUMANN & LAUTERBACH /.c.).
35. Gleichenia hirta BL. En. Pl. Jav. (1828) 250;
v. A. vy. R. Handb. (1908) 60; Suppl. (1917) 85;
Ho tr. Reinwardtia 4 (1957) 268.
—See for further synonyms under the varieties.—
Fig. 7d, 8h-j.
Main rachis bearing several pairs of branches,
not high-climbing; lateral branch-systems forked
in 2-4 orders (rarely 5), the first-order branches
in most varieties not normally proliferous beyond
their first fork; ultimate branches always much
longer than those of lower order, angle of ultimate
forks less than 45°; /amina thin, cut almost down
to the costa into thin oblique segments, edges
distinctly toothed towards apices of segments,
lower surface more or less glaucous (glaucous
character destroyed by heat in drying). Scales on
apex of main rachis to 5 mm long, less than 1 mm
wide, fringed with short hairs; scales on costae
abundant or not, spreading, 114-2 mm long, less
than 14 mm wide, thin and rather pale, not
26 FLORA MALESIANA
conspicuously hair-pointed, edges fringed with
slender hairs; smaller scales and stellate hairs also
present on costae, costules and veins, or quite
[ser. IE vor
lacking; sori with raised receptacle, conspicuous
paraphyses present or not.
KEY TO THE VARIETIES
1. Lower surface of costae of fully expanded ultimate branches scaly near the base only or more generally,
the scales all about equal in size with rather rigid fringing hairs; no scales on costules nor on veins
except when young.
2. Scales 3-4 cells wide at the base. Ultimate branches to 40 cm long and 344 cm wide.
2. Scales wider. Ultimate branches rarely over 25 by 214 cm.
3. Costules 4414 mm apart
3. Costules 314-4 mm apart
2. var. amoena
3. var. paleacea
4. var. candida
. Lower surface of costae persistently scaly almost throughout, the scales of various sizes, often with
lax fringe; smaller scales also persistent on costules and sometimes scales or hairs on veins.
4. Segments of the lamina not abruptly narrowed above the base, separated by narrow sinuses.
5. Scales on costules abundantly long-fringed.
6. Ultimate branches of well-grown fronds 2-3 cm wide; primary rachis-branches not always proli-
ferous beyond first fork
1. var. hirta
6. Ultimate branches 114-214 cm wide; primary rachis- branches always proliferous beyond first fork.
5. Scales on costules with very few long marginal hairs
5. var. amboinensis
6. var. virescens
4. Segments of lamina c. 2 mm wide except at the very base, separated by | sinuses 1-114 mm wide.
1. var. hirta.x—Dicranopteris dolosa COPEL. in
Perk. Frag. Fl. Philip. (1905) 193, 4. 4 fic.—G.
dolosa C. Cur. Ind. Fil. (1906) 664; v. A. v. R.
Handb. (1908) 62; Suppl. (1917) 85.—Sticherus
hirtus CHING, Sunyatsenia 5 (1940) 203 (err.
hirsutus).
Lateral branch-systems forked in 3 or 4 orders;
ultimate branches 15—22 cm long, 2-214 cm wide
(rarely to 3 cm); penultimate branches fully leafy
and often wider than ultimate ones; costules 3 mm
apart; scales on lower surface of costae copious,
persistent, of many sizes together, very long-
fringed; pale stellate hairs present on veins.
Type: Reinwardt, Tidore (L; dupl. at BM).
Distr. Malaysia: Moluccas and Philippines
(Luzon, Mindoro, Mindanao), 1200-1800 m.
2. var. amoena (v. A. v. R.) Hott. Reinwardtia
4 (1957) 269.—G. amoena v. A. v. R. Bull. Jard.
Bot. Btzg II, . 23 (1916) 12; Handb. Suppl. (1917)
497—Sticherus amoenus NAKAI, Bull. Nat. Sc.
Mus. Tokyo n. 29 (1950) .—G. peninsularis
CopeL. Un. Cal. Publ. Bot.12 (1931) 387.—Sticherus
peninsularis CHING, Sunyatsenia 5 (1940) 284.—
G: lurta; Hottrr. Rev. Fl. Mal. 2 (1955) 7-
Lateral branch-systems forked in 1-3 orders;
penultimate branches fully or partly leafy,
branches of lower order not leafy; ultimate
branches 25-40 cm long, 2!4-314 cm wide;
costules 4-414 mm apart; /amina-segments 3—-3\/
mm wide above the base, strongly glaucous
beneath; sca/es on lower surface of costae. narrow
(fringe of hairs longer than width of scale), very
sparse and near base of costae only, mature fronds
otherwise glabrous (stellate hairs on veins when
young).
Type:
ateP):
Distr. Malaysia: Sumatra, Lingga Archipelago,
Malay Peninsula.
Teysmann 16628, Lingga (Bo; dupl.
7. var. lanuginosa
Ecol. Altitude 700-1600 m; scrambling or
trailing in lightly shaded places or edge of forest,
not forming dense thickets; fronds usually not
over 200 cm long.
3. var. paleacea (BAK.) C. Cur. Gard.Bull.
S. S. 7 (1934) 212; Hottr. Reinwardtia 4 (1957)
269.—G. vestita var. paleacea BAK. J. Bot. 17
(1879) 38.—G. hallieri CuHrist, Ann. Jard. Bot.
Btzg 20 (1905) 138; v. A. v. R. Handb. (1908) 61.—
G. barbula C. Cur. Dansk Bot. Ark. 9, 3 (1937)
67.—Sticherus hallieri NAKAI, Bull. Nat. Sc. Mus.
Tokyo n. 29 (1950) 18.—Sticherus barbulus NAKAI,
byes, Mia}
Lateral branch-systems usually forked in 2 or 3
orders (to 5 orders in type specimen of G. barbula);
if in 2 orders penultimate branches incompletely
leafy and ultimate branches to 30 cm long, other-
wise ultimate branches 15—24 cm long and 2-214
cm wide; costules 4-4!4 mm apart; scales on
mature fronds confined to costae and all alike
(not mixed with smaller ones) with rather stiff
fringing hairs as long as width of scale.
Type: Burbidge, N. Borneo (K).
Distr. Malaysia: Borneo, Lesser Sunda Islands
(Bali), Celebes (Menado), 1100-3000 m.
Ecol. On Mt Kinabalu on open places in ridge
forest.
4. var. candida (RoseNstT.) Hott. Reinwardtia
4 (1957) 269.—G. candida RosENstT. in Fedde,
Rep. 5 (1908) 33; v. A. v. R. Handb. (1908) 796.—
Sticherus hirtus var. candidus NAKAI, Bull. Nat.
Sc. Mus. Tokyo n. 29 (1950) 19.
Similar to var. paleacea, but costules 314-4 mm
apart, lamina of firmer texture.
Type: Werner 72, Mt Gelu (B; dupl. at BM,
US, EE
Distr. East Malaysia: New Guinea, Admiralty
Islands, Solomon Islands.
Dec. 1959]
Ecol. On edges of forest and in light shade,
sometimes forming dense thickets to 3 m high,
700-2150 m.
5. var. amboinensis (v. A. v. R.) Hott. Rein-
wardtia 4 (1957) 269.—G. amboinensis v. A. v. R.
Bull. Dép. Agr. Ind. Néerl. 7. 18 (1908) 3; Handb.
(1908) 62; Suppl. (1917) 85 (as a synonym).
Like var. hirta in scaliness; ultimate branches
1144-2144 cm wide; primary rachis-branch always
proliferous beyond first fork and bearing 2 or
more pairs of secondary branches.
Type: Teysmann, Amboyna (Bo; dupl. at L).
Distr. Malaysia: S. Central Celebes, Moluccas
(Ambon, Buru, Waigeo Isl.).
Ecol. Climbing to 4 m in open places in
mountain forest.
6. var. virescens (HIERON.) HoLtr. Reinwardtia
4 (1957) 270.—G. dolosa var. virescens HIERON. ex
BRAUSE, Bot. Jahrb. 56 (1920) 209.—Sticherus
hirtus var. virescens NAKAI, Bull. Nat. Sc. Mus.
Tokyo n. 29 (1950) 19.
Similar to var. hirta, but scales on costules with
very few long marginal hairs.
Gleicheniaceae (Holttum) pe |
Type: Ledermann 9935 (B).
Distr. Malaysia: E. New Guinea.
Note. Collector of type specimen stated that
lower surface of lamina was white (i.e. glaucous)
when fresh.
7. var. lanuginosa (v. A. v. R.) comb. nov.—G.
ornamentalis ROSENST. Nova Guinea 8 (1912)
715; v. A. v. R. Handb. Suppl. (1917) 85.—G.
ornamentalis var. lanuginosa vy. A. v. R. Nova
Guinea 14 (1924) 23.—Sticherus lamianus COPEL.
Philip. J. Sc. 75 (1941) 356, pl. 6.—G. hirta var.
ornamentalis (ROSENST.) HoLtr. Reinwardtia 4
(1957) 269.
Lateral branch-systems commonly forked to 4
orders; costae persistently and densely scaly,
scales as in var. hirta; costules 3-4 mm apart,
segments of lamina 2 mm wide, thus separated
by rather wide sinuses.
Type: Lam 1945, Doormantop, 3480 m (U;
dupl. at Bo, US, L).
Distr. Malaysia: New Guinea.
Ecol. Altitude 800-2500 m; specimens from the
higher altitudes are the most densely scaly.
2. DICRANOPTERIS
BERNHARDI in Schrad. Neu. J. Bot. 1, 2 (1806) 38; UNDERW. Bull. Torr. Bot.
Cl. 34 (1907) 244, p.p.; CHING, Sunyatsenia 5 (1940) 272; CopeL. Gen. Fil. (1947)
28; NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950) 56; HoLtr. Reinwardtia 4
(1957) 273.—Mertensia WILLD. Kongl. Vet. Ak. Nya Handl. 25 (1804) 163,
p.p.; PRESL, Tent. Pterid. (1836) 50, p.p.; non RoTH, 1797 (Borag.).—Gleichenia
subg. Mertensia, § 1 and § IV, Hook. Sp. Fil. 1 (1844) 11, 12.—Hicriopteris PRESL,
Epim. Bot. (1851) 26, non CHING, nec COPEL.—Gleichenia subg. Mertensia sect.
Heteropterygium Diets in E. & P. Pfl. Fam. 1, 4 (1900) 355.—Gleichenia subg.
Mertensia sect. Dicranopteris v. A. v. R. Handb. (1908) 56, p.p.—Mesosorus
Hassk. Fil. Jav. 1 (1856) 2, p.p.—Fig. 11-16.
Indumentum on rhizome, dormant apices, and on other parts of fronds con-
sisting of multiseptate hairs which have outgrowths from cells near the base and
sometimes also from other cells. Apex of main rachis of fronds resting periodically
while each pair of lateral branch-systems develops; primary rachis-branches
repeatedly branched pseudo-dichotomously (forked), the apex between each pair
of branches usually permanently dormant, the members of a pair equal or unequal;
a short stipule-like lobed leaflet usually present at the base of each primary
rachis-branch, these leaflets growing upwards and protecting the temporarily
dormant apex of the main rachis (smaller stipule-like leaflets sometimes also
present at forks of the lateral branch-systems); a pair of accessory branches,
bearing a lamina like that of the ultimate branches, present at some of the forks
of the lateral branch-systems, on the outside of the fork (usually attached just
above it) and deflexed; apart from stipular leaflets and accessory branches,
only the ultimate branches leafy. Ultimate branches bearing throughout a deeply
pinnatifid /amina, the segments of the lamina usually entire, each with costule
bearing lateral veins which are forked at least twice; sori one to each vein-group
on an acroscopic branch (rarely also on a basiscopic branch), each of 8-15 or
. 500 m (MEIJER,
[ser. Il; voles
INN we
f
'
/ |
f
<
Z
<
DN |
sa)
4
<
~S
= SS
Z|
me |
eo)
4
ew
i
i]
Ly
Fig. 11. Dicranopteris curranii CopeL. Harau gorge, near Pajakumbuh, W. Sumatra, c
Dec. 1959] | Gleicheniaceae (Holttum) 29
more sporangia without paraphyses, the sporangia smaller than in Gleichenia;
spores monolete or trilete.
Distr. and Ecol. Thicket-forming ferns, abundant in open places throughout the wetter parts of
the tropics and subtropics, especially characteristic of equatorial lowlands (the species of Gleichenia
being almost exclusively mountain plants in Malaysia).
Notes. There are two well-marked subgenera, Dicranopteris (pantropic) and Acropterygium (one
species, in tropical America; no accessory branches, rhizome with solenostele).
The subgenus Dicranopteris is much more diversified in Malaysia than in any other part of the world.
Asmall group of species may be clearly distinguished by their monolete spores. Almost all other specimens
are at present regarded as belonging to a polymorphic species, D. linearis. Some of the varieties of
D. linearis are more clearly distinct than others, and probably should be recognized as species. They
/,
11K
Zz
vil
my
\
‘
.*
C d ©
Fig. 12. Diagrams showing variations of branching-habit in Dicranopteris. a. Basic habit, found in
D. linearis (BuRM. f.) UND. var. linearis, D. pubigera (BL.) NAKAI, and D. curranii COPEL., b. accessory
branches present at ultimate forks, as in D. linearis var. montana HOLtT. and var. tetraphylla (Ros.)
NAKAI, c. alternate unequal forking, as in D. linearis var. subpectinata (CurRIstT) HoLtt., var. alternans
(Me_tt.) Hottt., and var. subspeciosa Ho.tr., d. special development of c, found only in D. speciosa
(PRESL) HOLTT.
30 FLORA MALESIANA [ser. IT, volyatz
Fig. 13. Dicranopteris brake in an old crater swamp at c. 1200 m, Gajo Lands, N. Sumatra, surrounded
by pole wood forest consisting mostly of J/ex cymosa (1937).
need, however, to be more clearly characterized from field studies. The existence of a triploid hybrid
(see note below on cytology) indicates that it is probably impossible to refer every plant to a clearly
defined variety.
Four varieties of D. linearis have the common character of invariably producing accessory branches
at the bases of ultimate branches; in other characters they differ considerably from each other and do
not form a natural group. Another variety having this character occurs commonly in west tropical
Africa, but no such variety has been found in east Africa.
Cytology (observations by I. MANTON). Haploid chromosome number of D. curranii CopeL. (Singa-
pore) is 39. D. linearis (apparently the normal form of the species) in Ceylon is either diploid (n = 39)
or tetraploid (n = 78); a plant of D. linearis sent to Kew from Singapore is a sterile triploid.
KEY TO THE SPECIES
1. Spores monolete. The two branches at each fork equal; either lowland plants with ultimate branches
9-12 cm wide, or mountain plants. First fork of each vein at its very base.
2. Lamina not very thick nor with the veins grooved on the lower surface.
Dec. 1959]
Gleicheniaceae (Holttum) ¢ 31
60. Except in fig. a and b the cell of attachment is gS black.—
a. Hair from base of costa. — D. clemensiae Ho.tr. b. One of the rigid, shining
hairs from lower surface of costa, c. thin-walled hair from costule. — D. pubigera (BL.) NAKAI. d-e.
Hairs from costule, lower surface. — D. linearis (BURM. f.) UND., hairs from lower surface of costules:
Fig. 14. Hairs of Dicranopteris, all x
D. curranii COPEL.
f. var. subferruginea (H1IERON.) NAKAI, g. var. linearis, h. var.
sf
subspeciosa HOutTrt., i. var. alternans
(Metr.) HoLtr.
3. Ultimate branches commonly 40 cm long and 9-12 cm wide;
lower surface almost glabrous.
1. D. curranii
3. Ultimate branches not over 7 cm wide; lower surface of costae and costules rather persistently
hairy.
4. Ultimate branches 30-55 cm long, 5—7 cm wide; costae at first with many shining hairs (1-1)
mm long) throughout .
2451 DB). pee
4. Ultimate branches 15-25 cm long, D qe 5 cm wide: costae with such hairs near base only.
2. Lamina very thick, the veins grooved on the lower surface
3. D. pubigera
3. D. pubigera
1. Spores trilete. Branches at a fer’ in many cases unequal; ultimate branches to c. 6 cm wide. Veins
never grooved on lower surface; tis. ork of each vein distinctly above the base.
5. At each fork, except primary and ultimate ones, one branch leafy and without accessory branch,
the other branch not leafy and with an accessory branch
4. D. speciosa
5. At each fork, except (in most cases) an ultimate one, a pair of accessory branches present.
1. Dicranopteris curranii CopeL. Philip. J. Sc. 81
(1952) 4; Hottr. Reinwardtia 4 (1957) 274.—Glei-
chenia hermannii var. venosa BL. incl. also var.
tenera BL. En. Pl. Jav. (1828) 249.—Gleichenia
dichotoma var. tenera METT. in Mig. Ann. Mus.
Bot. Lugd. Bat. 1 (1863) 50.—Gleichenia dichotoma
var. malayana Curist, Ann. Jard. Bot. Btzg 15
(1898) 77.—Gleichenia linearis var. malayana
v. A. v. R. Handb. (1908) 59; Suppl. (1917) 84;
Hottr. Rev. Fl. Mal. 2 (1955) 70.—D. lessonii
NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950)
61, guoad specim., excl. basion. Mertensia lessonii
A. RicH.—Fig. 11, 12a, 14a, 16.
Fronds large, strongly erect but not greatly
elongate nor high-climbing; primary rachis-
branches few times equally forked; ultimate
branches commonly 40 cm long and 12 cm wide,
5. D. linearis
accessory branches at penultimate forks commonly
20-30 cm long and 7—10 cm wide; costules 5—7
mm apart; Jamina-segments 314-4144 mm wide
above the base, texture firm, lower surface
glaucous, usually quite glabrous apart from minute
simple hairs on the veins, often pinkish on costae
and lower parts of costules; veins slender, slightly
prominent on lower surface but not on upper;
stipular leaflets to 3 cm long at bases of primary
branches; spores monolete.
Type: Curran, For. Bur. 19265, Laguna Prov.,
Luzon (H. Copel.; dupl. at US, P).
Distr. Malaysia: Sumatra, Malay Peninsula,
W. Java, Lesser Sunda Islands (Flores), Borneo,
Celebes, Philippines (Luzon).
Ecol. In thickets with varieties of D. linearis,
from sealevel up to 1500 m.
32 FLORA MALESIANA
[ser. II, vol. 12
Note. Gleichenia weatherbyi FosBERG (Am.
Fern J. 40, 1950, 140), from the Caroline Islands,
is very near D. curranii, with similar spores, but
is even larger, and much more hairy.
2. Dicranopteris clemensiae Hott. Reinwardtia 4
(1957) 275.—Fig. 14b-c.
Main rachis 7 mm or more diameter; primary
branches twice equally forked, copious dark red
rigid hairs very persistent near the furks; ultimate
branches 30-55 cm long, 5—7 cm wide, the lowest
outer segments of the lamina sometimes enlarged
and more or less lobed; accessory branches at
penultimate forks 16-18 cm long, 5 cm wide;
stipular leaflets at base of primary branches 3 cm
long, lobes 3 mm wide; costules of ultimate
branches 5—6 mm apart; /amina-segments 314 mm
wide above the base, thin but firm; veins usually
3 times forked, first fork very close to the costule,
very slightly prominent on both surfaces: stout
dark red multicellular hairs (1-114 mm long, with
several basal branches) at first abundant all along
lower surface of costae, at length deciduous leay-
ing costae perceptibly rough; thinner and paler,
dull rusty-brown, hairs at first abundant on
costules, slightly crisped and with short unicellular
branches near base only; sori of 10-15 sporangia,
often two on a vein-group (on each of the outer-
most branches); spores monolete, c. 32 16 yu.
Type: Clemens 28745, Mt Kinabalu (BM;
dupl. at Bo, K, L, US).
Distr. Malaysia: N. Borneo (once collected,
1600 m).
3. Dicranopteris pubigera (BL.) NAKAI, Bull. Nat.
Sc. Mus. Tokyo n. 29 (1950) 68; Hott. Rein-
wardtia 4 (1957) 274.—Gleichenia hermannii var.
pubigera Bu. En. Pl. Jav. (1828) 249.—Gleichenia
dichotoma var. pubigera Metr. in Mig. Ann.
Mus. Bot. Lugd. Bat. 1 (1863) 50.—Gleichenia
linearis var. pubigera v. A. vy. R. Handb. Suppl.
(1917) 85.—Mertensia spissa (non FEE) var. pu-
bigera NaAKal, Bot. Mag. Tokyo 39 (1925) 180.—
Fig. 12a, 14d—e, 15a-b.
Primary rachis-branches 2-4 times equally or
subequally forked; ultimate branches 15-25 cm
long, 3-8 cm wide; costules 414-6 mm apart;
lamina-segments c. 4 mm wide when flattened,
thick and rigid when dry and usually with edges
reflexed; lower surface not glaucous, the costules
at first covered rather densely with coarse dull
rusty flexuous hairs and also some rigid shining
hairs; veins grooved on the lower surface when the
lamina is very thick, on thirner leaves hardly
grooved, pale but not raised on the upper surface;
spores monolete.
Type: Blume, Java (L; dupl. at K, P).
Distr. Malaysia: Sumatra, Java, and Lesser
Sunda Islands (Bali, Lombok, Flores).
Ecol. Abundant in open scrub and in clearings
in mountain forest, forming dense thickets,
1100-3000 m.
Note. The extreme forms of this species, one with
very thick lamina with veins grooved on the lower
surface, the other with thinner lamina and veins
not distinctly grooved, appear very different (the
former is larger as well as thicker) but there are
intermediates. The differences may in part be
connected with altitude and exposure; no ob-
servations have been made.
Fig. 15. Dicranopteris pubigera (BL.) NAKAI.
a. Lower surface of part of frond (veins grooved),
< 4, b. a main fork with stipular leaflets and ac-
cessory branches, * 1%. — D. speciosa (PRESL)
Ho itr. c. Part of frond to show venation, x 2,
d. main fork showing stipular leaflets, «x %.
4. Dicranopteris speciosa (PRESL) HoLtr. Rein-
wardtia 4 (1957) 273.—Hicriopteris speciosa PRESL,
Epim. Bot. (1851) 27.—Gleichenia opposita v. A.
v. R. Bull: Jard. Bot. Btzg I, 72 TGs ise
Ho tt. Rev. Fl. Mal. 2 (1955) 70, f. 14 F.—Glei-
chenia parallela RiDL. J. Mal. Br. R. As. Soc. 4
(1926) 3.—D. opposita NAKAI, Bull. Nat. Sc. Mus.
Tokyo n. 29 (1950) 68.—Fig. 12d, 15¢e—-d.
Primary rachis-branches several times very
unequally forked; at each fork the smaller branch
unbranched and leafy throughout, commonly
20-25 cm long and 4-5 cm wide, with no accessory
branch, the larger branch continuing almost the
same straight line as the larger branch from the
previous fork and bearing an accessory branch
12-15 cm long and 314 cm wide (this accessory
branch almost opposite the simple leafy branch
of the fork); ultimate fork formed by two normal
leafy branches without accessory branches;
irregularities from this basic scheme occasionally
present; lJamina-segments 31,4 mm wide, flat
and rather rigid, separated by very narrow sinuses,
apices often retuse; veins distinctly prominent on
the upper surface; lower surface more or less
glaucous, the costules when young rather densely
covered with fine pale entangled woolly hairs,
some similar hairs on veins; some dark redbrown,
rigid, shining hairs, like those on resting apices,
also at first present on costae and costules; sori
as in D. linearis; spores trilete.
Type: collector unknown, ‘“‘Pendschab”’ (PRc).
Perhaps from Penang.
Distr. Malaysia: Sumatra, Malay Peninsula,
Moluccas (Sula Isl.: P. Mangoli).
———————
Dec. 1959]
Gleicheniaceae (Holttum) 35
Ecol. Growing with D. linearis on the edges of
forest, 100-600 m.
Note. The remarkable pseudo-pinnate lateral
branch-systems of this species were described in
detail by PresL, but because he compared these
branches to Gleichenia glauca (THUNB.) HOOK.
later authors thought he was describing a species
of Gleichenia subg. Diplopterygium and conse-
quently used the generic name Hicriopteris in a
wrong sense.
5. Dicranopteris linearis (BURM. f.) UNDERW.
Bull. Torr. Bot. Cl. 34 (1907) 249; Ho.ttr. Rein-
wardtia 4 (1957) 275.—Polypodium dichotomum
THUNB. FI. Jap. (1784) 338, t. 37.—Gleichenia
dichotoma Hook. Sp. Fil. 1 (1844) 6; Racts.
Fl. Btzg 1 (1898) 13.—Fig. 12, 14f-i.
KEY 10, DHE VARIETIES
1. Accessory branches not always present at ultimate forks.
2. Branches at each fork of a lateral branch-system approximately equal;
forked 1-3 times.
lateral branch-systems
3. Costules less than 5 mm apart, their lower surface with some persistent hairs.
MeeLAIrS not persistent om lower surface OlevelnS -)ee 2 4) 6 4 8 6 6 Le
var. linearis
4. Hairs persistent along lower surface of veins.
5. Hairs long, very finely woolly and entangled; no distinct separate hairs present.
2. var. ferruginea
5. Hairs coarse, shorter, more or less entangled on costules, some distinctly separate on veins.
3. var. subferruginea
3. Costules 5-7 mm apart, their lower surface glabrous.
6. Lamina thick and gid
6. Lamina thin
2. Branches at successive forks alternately unequal;
forked at least 3 times.
7. Lower surface quite glabrous and persistently pale glaucous;
4. var. rigida
Sane 5. var. latiloba
lateral branch- -systems on well-grown plants
lamina thin 6. var. subpectinata
7. Lower surface more or less persistently hairy, less strongly glaucous (often losing glaucous character
on drying); lamina firm.
8. Hairs very slender, their long branches much crisped and entangled; no separate short hairs.
9. Hairs copious, at first covering whole lower surface, persistent along the veins, rusty when dry;
edges of lamina strongly reflexed when dry .
2. var. ferruginea
9. Hairs not covering whole lower surface, persistent only. on “costules, pale when dry; edges of
lamina not much reflexed when dry.
10. Angle of secondary and later forks not more than a right angle
10. Angle of secondary and later forks more than a right angle . . . 8.
7. var. subspeciosa
var. inaequalis
8. Hairs coarser and shorter, some more or less crisped and entangled but separate short hairs also
present.
11. Hairs confined to bases of some of the costules
1. var. linearis
11. Hairs more abundant on costules, and some also on veins.
12. Hairs persistently rust-coloured, copious all along veins. ns
12. Hairs pale on dried specimens, sparse on the veins; W. Malaysia
3. var. subferruginea
9. var. alternans
1. Accessory branches always present at ultimate forks.
13. Accessory branches, especially the lower ones, distinctly below the fork; branches at successive
forks alternately somewhat unequal
. 10. var. demota
13. Accessory branches opposite the fork or distinctly above it; branching usually equal at all forks.
14. Ratio of length to width of branches ee Die
214-3 mm wide
1; costules c. 3 mm apart, segments of lamina
11. var. tetraphylla
14. Ratio of length to width greater: costules more widely ‘separated.
15. Costules on ultimate branches 5-7 mm apart; texture subcoriaceous; lower surface quite glabrous;
accessory branches at ultimate forks almost opposite the fork . . . 12.
var. montana
15. Costules on ultimate branches 314-414 mm apart; texture thin; some rusty hairs often on lower
surface; accessory branches at ultimate forks 3-4 mm above the fork
1. var. linearis.—Polypodium lineare Burn. f.
Fl. Ind. (1768) 235, t. 67 f. 2.—Gleichenia herman-
nii R. Br. Prod. (1810) 161.—Gleichenia dichotoma
var. normalis MetT. in Mig. Ann. Mus. Bot.
Lugd. Bat. 1 (1863) 50; Racts. FI. Btzg 1 (1898)
13.—Mertensia pteridifolia PRest, Abh. (K.)
Bohm. Ges. Wiss. M.-N. Cl. V, 5 (1848) 339;
Epim. Bot. (1851) 23, t. 14.—Gleichenia linearis
CLARKE, Trans. Linn. Soc. Bot. 1 (1880) 428; v. A.
v. R. Handb. (1908) 59 pro var. normalis; Suppl.
13. var. altissima
(1917) 84, ditto; Hottr. Rev. Fl. Mal. 2 (1955)
70, ditto.—Fig. 12a, 14g, 16.
Primary rachis-branches commonly 2 or 3 times
forked, the two branches at all forks equal or
nearly so (not regularly alternately unequal);
stipular leaflets at bases of primary branches c. 1
cm long, lobed near the base only; ultimate
branches 15-25 cm long, 4-6 cm wide; /amina-
segments 214-3 mm wide, separated by rather wide
sinuses, texture very firm; lower surface slightly
34 FLORA MALESIANA
[ser. Il, volets
glaucous; veins slightly prominent on lower
surface and bearing very short simple hairs, not
prominent on upper surface; some persistent much-
branched rusty hairs on costules; spores trilete.
Type: Ceylon, Herb. Delessert (G).
Distr. Tropical and subtropical Africa, Asia,
Malaysia, Australasia, and Polynesia.
Ecol. Thicket-forming but not high-climbing,
from sealevel to 1400 m.
Note. Polypodium dichotomum THUNB. was
described from Japan, and as noted by NAKAI
(/.c.) is not quite identical with Polypodium lineare
BuRM. (BURMAN’S specimens, preserved at Ge-
neva, are from Ceylon). I have therefore ranked it
as a Dicranopteris linearis var. dichotoma (Rein-
wardtia 4, 1957, 277). The lamina of the fronds is
thinner than in var. linearis and the veins are
prominent on the upper surface. This may be
connected with the fact that (at least in part of
its range) the fronds of var. dichotoma only last
for one season, dying in the winter, whereas those
of var. linearis grow for an indefinite period which
is usually not climatically limited. Var. dichotoma
apparently occurs also in China and probably
in the Sikkim region; field studies in these regions
are needed to establish clear distinctions between
var. linearis and var. dichotoma.
2. var. ferruginea (BL.) Ho_tr. Reinwardtia 4
(1957) 278.—Gleichenia ferruginea Bu. En. Pl.
Jav. (1828) 249; Metr. Ann. Mus. Bot. Lugd.
Bat. 1 (1863) 50.—Gleichenia dichotoma_ var.
ferruginea Racis. Fl. Btzg 1 (1898) 13.—Gleichenia
linearis var. ferruginea v. A. v. R. Handb. (1908)
59; Suppl. (1917) 85.—‘Dicranopteris ferruginea’
CopeL. Philip. J. Sc. 75 (1941) 349, not Mertensia
ferruginea Desv. 1811.
Lateral branch-systems of small plants sym-
metrically forked twice, on larger plants forked
to 3 or more orders, branches at successive forks
alternately unequal; ultimate branches 15—20 cm
long, 3-5 cm wide; costules 4-5 mm apart; seg-
ments of lamina brittle when dry, with recurved
edges, when flattened c. 214 mm wide above the
base, lower surface covered with very fine floccose
entangled pale rusty hairs (c. 10 w diam.) which
are attached mainly along the veins, the veins
thus persistently hairy on old fronds; stiff hairs
present on dormant apices only; upper surface
smooth, the veins very distinct and slightly raised
when dry; accessory branches not normally
present at the ultimate forks, but the lowest basi-
scopic lobes of the ultimate branches usually
larger than the rest, their edges more or less lobed.
Type: Blume, Java (L).
Distr. Malaysia: Java, Celebes, Moluccas, New
Guinea.
Ecol. Altitude 1000-2500 m.
3. var. subferruginea (HIERON.) NAKAI, Bull.
Nat. Sc. Mus. Tokyo n. 29 (1950) 66.—Gleichenia
linearis var. subferruginea HIERON. ex BRAUSE,
Bot. Jahrb. 56 (1920) 209.—Fig. 14f.
Like var. linearis but perhaps more often with
unequal branches at the more distal forkings,
veins somewhat prominent on upper surface,
veins on the lower surface rather persistently
hairy, the hairs rusty, rather coarse (diam. c.
25 yw) and much-branched, not finely woolly nor
forming a continuous entangled web as in var.
ferruginea.
Type: Ledermann 6926, New Guinea (B).
Distr. Queensland, Polynesia (Fiji, Samoa),
and East Malaysia: Celebes, Moluccas, New Guinea.
Ecol. From sealevel to 1200 m.
Note. It is probable that this variety is much
more abundant than var. linearis in New Guinea
and regions further eastwards. The hairs on the
lower surface of the veins are of the same general
character as those of var. linearis, but much more
copious; they are more abundant on plants at
higher altitudes than near sealevel, and possibly
abundance of hairs also depends on degree of
exposure.
4. var. rigida (BL.) Hottr. Reinwardtia 4
(1957) 277.—Gleichenia hermannii var. rigida BL.
En. Pl. Jav. (1828) 249.—Mertensia crassifolia
Pres_, Abh. (K.) Béhm. Ges. Wiss. M.-N. Cl.
V, 5 (1848) 339; Epim. Bot. (1851) 23, t. 13.—
Gleichenia dichotoma var. rigida METT. in Miq.
Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 50.—Glei-
chenia warburgii Curist, Ann. Jard. Bot. Btzg 15
(4898) 78.—Gleichenia linearis var. rigida vy. A.
v. R. Handb. (1908) 59; Suppl. (1917) 84.—Glei-
chenia crassifolia CopeL. Philip. J. Sc. 1 (1906)
Suppl. I 257.—D. crassifolia NAKAI, Bull. Nat.
Sc. Mus. Tokyo n. 29 (1950) 57. (Non Mertensia
rigida Kze 1834, D. rigida NAKAI).
Lateral branch-systems once or twice forked,
members of all forks equal or nearly so; ultimate
branches 15-25 cm long, 3—5!4 cm wide; costules
5-6 mm apart; segments of lamina 4-5 mm wide
above the base, thick and very rigid when dry
with edges slightly reflexed, apices not or slightly
retuse; lower surface quite glabrous from an early
stage; veins broad, slightly prominent on lower
surface, distinctly so on upper surface; spores
trilete.
Type: Reinwardt, Tidore (L).
Distr. Malaysia: Moluccas (type from Tidore),
Celebes, and Philippines (Mindanao, Luzon,
Negros).
Ecol. Altitude 1500-3000 m.
Note. MEeETTENIUS (and apparently also RACI-
BORSKI) referred some specimens of D. pubigera
from Java to this variety of D. linearis.
5. var. latiloba HoLtr. Reinwardtia 4 (1957) 277.
Lateral branch-systems forked to about 4th
order, branches at all forks subequal; stipular
leaflets at bases of primary branches to 4 cm long,
deeply and broadly lobed; ultimate branches
c. 20 cm long, to 9 cm wide, the lowest segments
of the lamina somewhat enlarged and deflexed
but not pinnately lobed; costules c. 7 mm apart;
segments 4-5 mm wide above the base, firm but
not very thick, edges slightly sinuous, only
reflexed when dry; lower surface quite glabrous
apart from minute simple hairs; veins sometimes
Dec. 1959] ;
Gleicheniaceae (Holttum) 35
Fig. 16. Thicket of Dicranopteris on edge of forest, Singapore. D. curranii CopPeEL. (large fronds) to left
and right; D. linearis (BURM. f.) UND. var. linearis in centre; var. subpectinata (CHR.) HOLTT. to right
of centre, below (HoLtTrum,
pale and distinctly prominent on the upper surface,
concolorous and only slightly prominent on
lower surface.
Type: Merrill 975, Luzon (US; dupl. at P, U, F).
Distr. Malaysia: Philippines (Luzon, Negros,
Mindanao), Celebes (?).
Ecol. Altitude 1000-1600 m.
6. var. subpectinata (CHrRist) Ho.tr. Rein-
wardtia 4 (1957) 277.—Gleichenia subpectinata
CurisT, Bot. Tidsskr. 24 (1901) 111.—Gleichenia
Pteridifolia (non Mertensia pteridifolia PRESL)
RIpL. J. Mal. Br. R. As. Soc. 4 (1926) 4.—D.
warburgii (non Gleichenia warburgii CHRIST) sensu
NAKAI, Bull. Nat. Sc. Mus. Tokyo n. 29 (1950)
70.—Gleichenia linearis var. alternans (non METT.)
Ho tr. Rev. Fl. Mal. 2 (1955) 70 and frontispiece.
—Fig. 12c, 16.
Lateral branch-systems several times forked,
branches at successive forks alternately unequal,
successive larger branches forming almost a
Straight line; ultimate branches 9-15 cm long,
2-314 cm wide (sterile ones sometimes to 18 by
6 cm); /amina thin, lower surface pale glaucous
and quite glabrous when mature; segments of
1925).
lamina gradually and rather evenly tapering from
base to apex, the sinuses thus narrowly triangular;
costules 4414 mm apart; veins slightly raised on
both surfaces, concolorous with lamina; the
smaller accessory branches usually attached 3-5
mm above the forks (sometimes lacking or reduced
at penultimate forks).
Type: Schmidt, Koh Chang Isl. (P).
Distr. Lower Siam (type from Koh Chang
Island), and Malaysia: Sumatra, Malay Peninsula,
Lingga Archipelago, Banka, Borneo.
Ecol. On edges of forest, in thickets with var.
linearis and other varieties, climbing higher than
var. linearis, from sealevel up to 700 m.
7. var. subspeciosa Hott. Reinwardtia 4 (1957)
278.—Fig. 12c, 14h.
Resembling D. speciosa in shape, texture etc.
of ultimate branches and in hairiness, but lateral
branch-systems branched as in var. alternans.
Type: Topping 1516, Mt Kinabalu, Kiau (US;
dupl. at SING).
Distr. Malaysia: Sumatra, Malay Peninsula,
Borneo, and Philippines (Mindanao, Mindoro,
Luzon).
36 FLORA MALESIANA [ser. II, vol. 11, Dec. 1959]
Ecol. On edges of forest, in thickets with other
varieties, 0-1400 m.
8. var. inaequalis (ROSENST.) HoLTr. Rein-
wardtia 4 (1957) 278. — Gleichexia linearis var.
inaequalis ROSENST. in Fedde, Rep. 13 (1915) 212;
vy. A. v. R. Handb. Suppl. (1917) 85.
Shape and venation of /amina-segments, and
hairiness, as in var. subspeciosa, differing as follows
in branching: forking in lateral branch-systems
alternately very unequal, the angle of the forks
much more than a right angle (sometimes nearly
180°), the smaller branch at a fork simple or once
forked and sometimes lacking anaccessory branch,
the accessory branch on the larger branch at-
tached 5-10 mm above the base of that branch.
Type: J. Winkler 114, Batakerland, Sumatra
(H. Selim Birger, S).
Distr. Malaysia: Sumatra, Malay Peninsula
(three collections in all).
Ecol. Altitude c. 1400 m.
9. yar. alternans (MetTT.) HoLtr. Reinwardtia 4
(1957) 278.—Gleichenia dichotoma var. alternans
Me_etr. in Mig. Ann. Mus. Bot. Lugd. Bat. 1
(1863) 51.—Gleichenia linearis var. alternans
v. A. v. R. Handb. Suppl. (1917) 84.—Fig. 12c,
14i.
Very similar to var. linearis in texture, shape,
and hairiness of the ultimate branches, but lateral
branch-systems several times forked, the branches
at successive forks alternately unequal about as
in var. subpectinata, differing from the latter in
larger somewhat hairy ultimate branches of much
firmer texture.
Type: Korthals, Sumatra (L).
Distr. Malaysia: Sumatra, Malay Peninsula,
Banka, Borneo.
Ecol. Apparently not common, but needs field
study; perhaps only a growth-form of var.
linearis; from sealevel up to 500 m.
10. var. demota Ho.tr. Reinwardtia 4 (1957)
Did
Lateral branch-systems several times forked,
branches at successive forks alternately unequal;
accessory branches normally present at all ulti-
mate forks; ultimate branches to 18 cm long,
3-6 (rarely to 7) cm wide; costules 5 mm apart;
lamina thin, lower surface glabrous and probably
glaucous; veins distinctly prominent on upper
surface and slightly so on lower surface; accessory
branches at the lower forks attached distinctly
below the fork.
Type: Clemens 29535, Mt Kinabalu, 5600 ft
(K; dupl. at BM, Bo, L, US).
Distr. Malaysia: N. Borneo, Ceram (doubt-
ful), New Guinea.
Ecol. Altitude 1600-2250 m.
11. var. tetraphylla (ROsENST.) NAKAI, Bull.
Nat. Sc. Mus. Tokyo n. 29 (1950) 67.—Gleichenia
linearis var. tetraphylla ROSENST. in Fedde, Rep.
13 (1914) 213; v. A. v. R. Handb. Suppl. (1917) 84.
—Fig. 12b.
Lateral branch-systems several times forked,
branches at successive forks alternately somewhat
unequal; penultimate branches less than 1 mm
diam. when dry; accessory branches always present
at bases of ultimate branches; ratio of length to
width of ultimate and accessory branches about
2:1; ultimate branches to 12 cm long and 7 cm
wide, costules 3 mm apart, /amina very thin and
strongly glaucous on the lower surface, quite
glabrous; veins very slender, distinctly prominent
on both surfaces.
Type: J. Winkler 136, Batakerland, Sumatra
(H. Selim Birger, S).
Distr. Indo-China, Hainan, Kwangtung, in
Malaysia: Sumatra.
Ecol. Altitude 1250 m.
Note. The specimens from China have ultimate
branches proportionately rather narrower than
those from Sumatra, but are otherwise similar.
12. var. montana HoLtr. Reinwardtia 4 (1957)
276.—Gleichenia linearis var. montana HOLTT.
Rev. Fl. Mal. 2 (1955) 69, descr. angl.—Fig. 12b.
Lateral branch-systems several times equally
forked; ultimate branches 15-25 cm long, 314-6
cm wide; costules (5—)6-7 mm apart; /amina
coriaceous, glabrous and glaucous on the lower
surface, veins rather strongly prominent on both
surfaces; accessory branches, about half as long
as the ultimate branches, always present at the
bases of ultimate branches.
Type: Molesworth Allen 2720 (SING).
Distr. Ceylon, S. India, Sikkim, in Malaysia:
Sumatra, Malay Peninsula, Java, Borneo, and
Moluccas (Ternate).
Ecol. On the edges of forest and in clearings,
with other varieties of D. Jinearis and with other
members of the family; 1000-1600 m.
13. var. altissima Ho.Ltr. Reinwardtia 4 (1957)
276.—Gleichenia linearis var. altissima WHOLtr.
Rev. Fl. Mal. 2 (1955) 69, descr. angl.
Sometimes very high-climbing, the main rachis
to 10 mm diameter; lateral branch-systems
equally forked 4 or 5 times; ultimate branches
to 15 cm long and 3 cm wide; costules 314-414
mm apart; /amina thin, lower surface glaucous,
glabrous apart from scattered rusty hairs on lower
surface of costules; veins prominent on upper
surface, not on lower surface; accessory branches
always present at bases of ultimate branches.
Type: Corner 31447, Johore, Malay Peninsula
(SING; dupl. at K, L).
Distr. Malaysia: Malay Peninsula, Philip-
pines (Palawan, Luzon), Moluccas (Talaud), New
Guinea, and Solomon Islands.
Ecol. At low altitudes, climbing on edges of
forest.
SCHIZAEACEAE (R. E. Holttum, Kew)
Rhizome usually short-creeping with closely-placed fronds, less often wide-
creeping or somewhat erect, the young parts covered with thick septate hairs
(except Mohria, not Malaysian), structure dorsiventral or radial, vascular strand
in Malaysian genera a protostele (medullated in Schizaea). Fronds of very varied
structure, their branching showing varying gradations from dichotomous to
pinnate; veins usually free; sporangia borne on specialized segments of the fronds
(sorophores) except in the non-Malaysian Mohria. Sorophores at the ends of veins
of fertile leaflets (Lygodium), or in small pinnate groups at the apex of a frond
or of its branches (Schizaea), or confined to special branches of the frond
(Anemia, not Malaysian). Sporangia arising marginally but becoming superficial
due to subsequent extra-marginal growths, large, borne on short massive stalks
or sessile, with an almost apical annulus of a single row of elongate thickened
cells, dehiscing on a line from annulus to base. Spores trilete or monolete (Schizaea
only), without perispore, the surface usually sculptured. Gametophytes filamentous
in Schizaea, thalloid in other genera, symmetrical or not.
Distribution. The Malaysian genera Schizaea and Lygodium are pantropic with a few outlying
species of both in temperate regions (U.S.A., S. Africa, Chile, Japan, and New Zealand). Anemia has
its main distribution in tropical America, with a few species in Africa and one in southern India. Mohria
is confined to southern and eastern Africa and the Mascarene Islands.
Fossils. The older fossils belong to extinct genera, which are quite different in frond-form from
living members of the family; their relationship to these living members is shown only by their sporangia
and spores. The Upper Carboniferous genus Senftenbergia has been most recently and fully described
by RADFORTH (Trans. R. Soc. Edinb. 59, 1938, 385-396; ibid. 1939, 745-761). The fronds were bipinnate-
tripinnatifid, comparable in size with those of Cyathea of today, with sterile and fertile parts alike in
shape of lamina; the rhizome is not known. The sporangia, attached to the edge of the lamina by very
short stalks, were similar in shape to those of the living genus Anemia but differed in having an annulus
of 2-5 rows of cells, in at least one case not sharply differentiated from the rest of the sporangial wall.
The Jurassic genus K/ukia had fronds of similar form, but the sporangia had an annulus of a single
row of cells, much as in Anemia. C. F. REED has made the most recent summary of fossil forms in the
family (Bol. Soc. Brot. 21, 1947, 71-197); he includes also the genera Tempskya, Acrostichopteris,
Pelletiera, and Schizaeopsis, from Upper Jurassic and Cretaceous horizons, but sporangia of these
have not been seen. Fossils of the living genera Anemia, Lygodium, and Schizaea have been found in
late Cretaceous and early Tertiary rocks in Europe and North America (REED, /.c.; CHANDLER, Bull.
Br. Mus. Nat. Hist., Geol., 2, 1955, 291-314; SELLING, Act. Hort. Gotob. 16, 1944, 1-112); fossils of
Schizaea have also been found in Eocene of SE. Australia and Pliocene in New Guinea (CooKsoN,
Proc. R. Soc. Victoria n.s. 69, 1957, 41). Fossils of Anemia and Lygodium include good leaf-impressions
and sporangia; those of Schizaea are confined to spores. One species of Schizaea is also known from
fossils of Quaternary age in Hawaii.
Ecology. Lygodium plants are twining climbers, mainly in secondary vegetation (fig. 6, 10), producing
fertile leaflets on parts exposed to the brightest light (in most cases, to full sunlight). Fronds of the larger
species may climb to a height of 10 m, but others only to 2-3 m. In regions where there is a prolonged
dry season the plants may be confined to wet ground. L. microphyllum, L. polystachyum and L. salici-
folium have caducous leaflets which fall when old. Whether plants are ever quite bare of leaflets in dry
seasons is not recorded; certainly the rhizomes have no power of resisting considerable drought, and
most plants are evergreen. L. japonicum occurs outside Malaysia in decidedly seasonal climates, and in
Malaysia does not occur in the uniformly wet region of Sumatra, Malaya and Borneo. L. polystachyum,
native of the seasonal climate of the region from northern Malaya to Burma and Indochina, is only
found in shady forest; possibly this is also true of L. merrillii. All species of Lygodium in Malaysia
are lowland plants, the highest altitude record being little over 1000 m, apart from L. japonicum (to
2500 m).
Schizaea plants are all small, and occur in poor acid sandy or peaty soils where the vegetation is rather
open (fig. 1, 2), apart from S. digitata, which always occurs in light to moderate shade. S. fistulosa is a
high mountain plant, S. malaccana on exposed ridges at moderate elevations, the other species in low
country. S. inopinata occurs only on limestone. S. wagneri and S. spirophylla are reported as growing
in moss-cushions on trees.
Vegetative morphology. Both Lygodium and Schizaea are specialized, in quite different ways;
38 FLORA MALESIANA [ser. Il, volaae
Fig. 1. Unusually dense stand of Schizaea dichotoma (L.) SM. on very poor sandy podsol, under shrubs
near Pasir Pandjang, west coast of Borneo (DUNSELMAN, 1936).
as above indicated, they are adapted to peculiar habitat-conditions. It is probably significant that
neither is represented by fossils earlier than late Cretaceous or Eocene. The sorophores of both genera
(and of Anemia) are perhaps reduced modifications of the lobes of fertile leaflets of ancestral ferns with
fronds like those of Senftenbergia. The photosynthetic function of such lobes is transferred in both
genera to a lamina developed from the wing of the rachis which occurs throughout the family (see
further notes under the genera); such development is much more extensive in Lygodium than in Schizaea
(for a discussion of transference of function, see CORNER in Journ. Linn. Soc. Bot. 56, 1958, 33-48).
Fronds of Schizaea are either unbranched (the ultimate condition of reduction) or symmetrically
dichotomously branched. Fronds of young plants of Lygodium are dichotomously branched, but later
fronds develop an elongate sympodial climbing rachis by means of a series of alternate unequal dichoto-
mies. The short branch at each dichotomy of a climbing rachis has a pair of opposite secondary branches
beyond which it is dormant (HoLTTUM, Phytomorphology 7, 1957, 152). The dormant apices may grow
if there is injury to a distal part of the frond. Similar, but not identical, dormancy occurs in the families
Gleicheniaceae and Matoniaceae. Periodic dormancy of apices occurs in Dennstaedtia, Hypolepis and
Paesia, which are perhaps derivatives of Schizaeaceae.
PRANTL investigated the development of sorophores in Schizaeaceae, and stated that the sporangia
were marginal in origin (Untersuch. z. Morph. der Gefasskrypt. II, 1881). Diets doubted PRANTL’s
observations (in Engler, Pflanzenfam. 1, 4, 1900, 356-372), but later workers confirmed them (BOWER,
The Ferns 2, 1926, 163-165). The lamina of the sorophores, and the separate indusia for each of the
sporangia in Lygodium, are later extra-marginal developments.
Sporangia and spores. The sporangia are the most distinctive feature of the family; they are well
illustrated by PRANTL for all genera and his drawings have been frequently copied. The sporangia are
large, almost or quite sessile, and all have an almost apical annulus with a longitudinal line of dehiscence.
Those of Lygodium are peculiar in having an asymmetrically placed lateral attachment. The spores are
in most cases rather elaborately sculptured on the surface, and are often specifically diagnostic (see
further discussion under the genera).
Gametophyte. The gametophyte of Schizaea is filamentous, with antheridia and archegonia on
special short branches; some cells have an endophytic fungus. Other genera have more normal gameto-
phytes, sometimes with asymmetric growth; their antheridia are large, and they show some other
primitive features (for references up to 1926, see Bower, /.c. 170). Most observations of gametophytes
have been of non-Malaysian species.
Dec. 1959] | Schizaeaceae (Holttum) 39
Cytology. The only records are by MANTON & SLEDGE (Phil. Trans. R. Soc., B, 238, 1954, 142-143)
and Lovis (Nature 181, 1958, 1085). Lygodium scandens (L. microphyllum of present work) in Ceylon
had n = 30. L. circinnatum in Ceylon had n = 58, but a plant in cultivation at Kew had n = 29. A plant
of L. japonicum at Kew also had n = 58. The basic number in Anemia is 38; a naturalized plant in Ceylon
was tetraploid, a plant in cultivation at Kew diploid. Schizaea asperula WAkeF. in New Zealand had
n = 77 (Lovis, /.c.); the same number was also found by Lovis (unpublished) for S. dichotoma in New
Zealand. In Ceylon Lovis found that S. digitata had a very high number (n = 325 + 30). These figures
indicate that polyploidy is not uncommon in the family, and that the extreme reduction of plant-form
in Schizaea may be associated with high polyploidy.
Anatomy. The fullest account of anatomy in the family is by PRANTL (J/.c.); later works are cited
by Bower. The rhizome of Lygodium has a solid protostele; the rachis of the climbing frond has also a
compact vascular strand (not C-shaped), with very large tracheids in the xylem, no doubt in adaptation
to its habit (a slender twining rachis carrying many leaflets, which thus needs a vascular system of capacity
large in proportion to its area of cross-section). The rhizome of Schizaea has a more or less medullated
protostele, and the stipe has a small compact vascular strand in which the xylem is reduced, more or less
3-armed as seen in cross-section. In both genera there is considerable development of sclerenchyma.
The genera Anemia and Mohria have a more complex vascular anatomy. Schizaea shows specialization
in the arrangement of stomata, the details of this varying from species to species.
Economic importance. The tough slender climbing rachises of Lygodium find various uses, either
in their natural form, or prepared by splitting for finer purposes. They are used as a substitute for cord
(e.g. for tying sheaves of rice), for plaiting into hats, bracelets, erc., for fastening the rims of sieves, and
in other ways. There are records of the use of several species of Lygodium, and also of Schizaea dichotoma,
for a great variety of medicinal purposes (see BURKILL, Dict. Econ. Prod. Mal. Pen. 1378, 1975; also
HEYNE, Nutt. Pl. Ned. Ind. ed. 2, 96, 97), but no critical study of such uses has been made. Very young
leaves of Lygodium microphyllum and L. circinnatum are eaten in Java (OCHsE, Veg. D.E.I. 1931, 655—
657). The plants are also used (especially Lygodium) in magical ceremonies connected with house-building,
rice culture, fishing, efc.
Taxonomy. The early history of the taxonomy of the genus Lygodium is very complex, for various
reasons. LINNAEUS began badly by including references to three distinct species under his Ophioglossum
scandens (the basis of Lygodium scandens Sw.); and he was unfortunate in having a poor specimen of a
sterile frond of an immature plant on which to base his O. fexuosum. In the years immediately following
1800, several authors were independently studying specimens of Lygodium, and the following generic
names were given: Lygodium Sw., Ugena Cav., Ramondia MirBeL, Odontopteris BERNH., Gisopteris
BERNH. and Hydroglossum WILLD. (all in 1801), Cteisium Micux (1803) and Vallifilix THOUARS (1809).
The following names were also given to Schizaea: Lophidium Ricu. (1792) and Ripidium BERNH. (1801).
For a full bibliographic statement on these genera, and a discussion of their typification, see PICHI-
SERMOLLI, Webbia 12 (1955) 4-36.
In the case of Lygodium, there is so much variation in leaflet-form, due to (a) age of plant, (b) en-
vironmental conditions, (c) height above ground from which specimen is taken, (and probably also to
polyploidy and hybridization) that, even with ample material, it is not easy to define specific limits,
and there was much confusion in the use of names by earlier authors. Different forms of the same
species received different names, while in other cases two quite different species were confused under
one name. WILLDENOw based his Hydroglossum pinnatifidum on two specimens, one sterile and one
fertile, belonging to two quite distinct species, and his name, transferred to Lygodium, was subsequently
used by different authors for both these species. For a detailed discussion of this subject, see ALSTON
& Hoxrtrum, Reinwardtia 5 (1959) 11—22.
The only good monograph of the whole family is that of PRANTL (/.c., 1881), whose very thorough
morphological study (of material then available) laid a sound basis on which others, working on various
aspects of the family and with new material, could build. Diets (in Engler, Pflanzenfam.) followed
PRANTL with little alteration, as did also CHRISTENSEN (Index Filicum, 1905). As regards nomenclature,
PRANTL did not look fully into the typification of L. scandens (L.) Sw. and he did not follow some of
our modern rules. Also he did not have good material (in a few cases he had no material) of some of
the less common Malaysian species.
Nakal published a survey of the whole family in 1937 (J. Jap. Bot. 13, 139-154). He devided it into
the three families Schizaeaceae, Lygodiaceae and Anemiaceae (including Mohria in the last) and also
raised some infra-generic groups to generic rank. This process was carried further by C. F. REED (Bol.
Soc. Brot. 21, 1947, 71-197) who raised the rank of the whole group to that of an Order, Schizaeales,
with families for the fossil as well as living representatives. He separated Mohria as a family distinct
from Anemia, and raised further subdivisions of Lygodium and Schizaea to generic rank, but made no
critical contribution to the understanding of species, nor any new basic morphological study. COPELAND
(Gen. Fil. 1947) made little change from CHRISTENSEN’S arrangement.
KEY TO THE GENERA
1. Fronds simple and linear, or dichotomously branched with linear branches (which in some non-
Malaysian species are joined laterally), the sorophores borne laterally near the apex of the frond
40 FLORA MALESIANA [ser. Il; volo
or of its branches; no indusia. Spores monolete. . . . « 1. Schizaea
1. Fronds of young plants dichotomously branched, of older plants of indefinite growth with twining
rachis, the rachis-branches variously branched and bearing leaflets which have sorophores at the
ends of their veins; each sporangium protected by a separate indusium. Spores trilete 2. Lygodium
1. SCHIZAEA
Sm. Mem. Ac. Turin 5 (1793) 419, nom. cons.—Lophidium Ricu. Act. Soc. Hist.
Nat. Paris 1 (1792) 114.—Actinostachys WALL. ex Hook. Gen. Fil. (1842) t. 111;
REED, Bol. Soc. Brot. 21 (1947) 130.—Microschizaea REED, /.c. 133.—Fig. 1-4.
Rhizome creeping or suberect, the young parts, and bases of stipes, covered
with coarse septate hairs, vascular system a medullated protostele. Stipes erect,
slender, narrowly winged towards apex in most species. Frond simple or dicho-
tomously branched, lamina reduced to a narrow wing bearing a single (rarely
double) row of stomata on the lower surface; two-celled glandular hairs frequent
on the surface of fronds, the basal cells persistent and often forming small warts,
distal cells shrivelling or falling when old or dried. Sorophores pinnately arranged
at the apex of a frond or of its branches, each sorophore with a median ridge on
its lower (abaxial) surface, the sporangia attached to the sides of the ridge, the
reflexed edge of the lamina protecting them. Sporangia not quite symmetrically
ovoid or ellipsoid, sessile, with distal annulus of a single row of cells; spores pale,
monolete, the surface variously sculptured.
Distr. Pantropic, comprising c. 30 spp., widely distributed also in temperate regions of the southern
hemisphere (S. Africa, Chile, New Zealand, Tasmania) but only in N. America in the northern hemisphere.
Morph. PRANTL referred to the whole fertile apex of a frond of Schizaea (or of a branch of S. dicho-
toma) as a sorophore; he called the lateral appendages /aciniae. But it seems probable that each lacinia
is homologous with a sorophore of Lygodium, and the term sorophore is therefore here used for what
PRANTL called a lacinia. Photosynthetic tissue in Schizaea is reduced to the wings on each side of the
axis of a simple frond, or of the branches of S. dichotoma. The regular row of stomata associated with
this photosynthetic tissue (a double row in S. inopinata) corresponds to a similar less regular row along
each side of the stipe and rachis of many ferns. In Lygodium, there are irregularly scattered stomata on
both sides of the narrow rachis-wings.
Spores. SELLING published a very full account of the spores of all species known to him in 1944
(Medd. Géteb. Bot. Tradg. 16, 1-112, p. 1-5). He also discussed the later-known species of the group
of S. digitata (Svensk Bot. Tidskr. 41, 1947, 431-450). Fossil spores are also known (see remarks on
fossils, supra).
KEY TO THE SPECIES
ieeirondssrepeatediy dichotomous? ails) se) es) ee) eco tee atcle ele) Syne. ee 1. S. dichotoma
1. Fronds simple.
2. Sorophores 214-5 cm long, all attached close together at apex of frond; frond 2 mm or more wide.
3. Sporangia in four rows on the sorophores; one row of stomata on each wing of the frond.
2. S. digitata
3. Sporangia in two rows on sorophores; stomata in two rows on each wing of the frond.
3. S. inopinata
2. Sorophores much shorter, fronds narrower.
4. Edges of sorophores smooth and glabrous.
5. Fronds more than 1 mm wide, with distinct flat wing on each side of costa; no hairs with
sporangia.
6. Costa very prominent on lower surface; stomata in each row very close together 2. S. digitata
6. Costa hardly prominent on lower surface; stomata in each row rather widely spaced.
4. S. spirophylla
5. Fronds less than 1 mm wide, wing very slightly developed; hairs present with sporangia.
5. S. wagneri
4. Edges of sorophores irregularly lobed, the lobes bearing coarse hairs.
7. Sorophores attached along the distal 10-20 mm of the axis of the frond; upper surface of frond
deeply grooved, lower surface evenly rounded and bearing superficial rows of stomata.
6. S. fistulosa
7. Sorophores attached along the distal 5-10 mm of the axis of the frond; upper surface of frond
slightly grooved, stomata intwo slight grooves on the distinctly flattened lower surface 7.S. malaccana
Dec. 1959]
Schizaeaceae (Holttum) ~— 41
1. Schizaea dichotoma (L.) SM. Mem. Ac. Turin 5
(798) 422. t.09; Buc Ens) Pls Javs (1828) 2555
Hook. & GrReV. Ic. Fil. (1827) t. 17; BEDp. Ferns
S. India (1863) t. 65; Handb. (1883) 452;
PRANTL, Unters. Morph. Gefasskr. 2 (1881) 138;
Racis. Pterid. Buit. (1898) 6; v. A. v. R. Mal.
Fig. 2. Schizaea dichotoma (L.) SM. near Kepa-
hiang, Bencoolen, S. Sumatra (DE VooGD).
Ferns (1908) 116; Merr. Int. Rumph. (1917) 69;
BACKER & Postu. Varenfl. Java (1939) 256,
fig. 66; Hoxttr. Ferns Mal. (1955) 50, fig. 6.—
Acrostichum dichotomum LINNE, Sp. PI. (1753) 1068.
—Osmunda dichotoma Spr. in Schrader, J. Bot.
(1799) pt 2, 268.—Ripidium dichotomum BERNH.
in Schrader, J. Bot. 1800, pt 2 (1801) 127, t. 2,
f. 3.—S. forsteri Spr. Anleit. 3 (1804) 57.—S.
cristata WILLD. Sp. Pl. 5 (1810) 88.—S. biroi
RICHTER, Math. Termeszet. Ertesito 29 (1911)
1074; TROLL, Flora 128 (1933) 339, fig. 1.—S.
copelandica RicuTeER, /.c.—Fig. 1, 2, 4a-d.
Rhizome 3-6 cm below surface of ground,
creeping, sometimes to 6 cm or more long, densely
covered with coarse shining brown hairs 2-3 mm
long. Stipes commonly 15-30 cm long (extremes
10-50 cm), narrowly winged towards apex; frond
commonly 10-20 cm long and wide, dichoto-
mously branched 2-8 times, the basal branches
like the stipe, the distal ones gradually with wider
wings and 1-114 mm wide, lacking a prominent
costa on the lower surface; all parts with scattered
small projections which are the bases of glandular
hairs; sorophores occupying the distal 3-5 mm
of each branch of the frond, 5-10 pairs, the lowest
3-4 mm long, upper ones smaller, edges hairy;
sporangia in two rows, mixed with conspicuous
long brown hairs; spores smooth or minutely
granular.
Type: Petiver, Gaz. t. 70, f. 12 (drawing of a
specimen from Cochinchina).
Distr. Mascarene Isl., Ceylon and S. India,
Burma, Siam, Indochina, throughout Malaysia
except for East Java and Lesser Sunda Isl., to
Tahiti, Australia, and New Zealand.
Ecol. In lightly shaded places, or sometimes in
forest, often (always?) in sandy ground, sealevel
to 1000 m, rarely abundant.
Note. Small plants like those named S. biroi
and S. copelandica by RICHTER are not uncommon,
and have been found at many places near larger
plants. As pointed out by TROLL (/.c.) the small
little-branched fronds of these small plants are
usually fertile, whereas some fronds of much larger
plants are sometimes sterile. I have however
examined a very large number of specimens and
have failed to find any sharp distinction between
those with little-branched and much-branched
fronds. If one picks out individual specimens,
one can separate fronds with long and with short
ultimate branches; but in some cases fronds from
one collection may show ultimate branches of very
diverse length.
Vern. Tatagar payong, Kedayan, pirangas,
Murut, oemiar, biak, E. New Guinea, paku tjakar
ajam, radja hantu, Banka, rumput bulu merak,
Billiton, silaju, Sum.
Uses. HEYNE records medicinal use (in Billiton)
for coughs and affections of the throat and also
in childbirth.
2. Schizaea digitata (L.) Sw. Syn. Fil. (1806)
150, 380, t. 4 f. 1; Bu. En. Pl. Jav. (1828) 255;
Bepp. Ferns S. India (1864) t. 268; Handb.
(1883) 452; PRANTL, Unters. Morph. Gefasskr.
2 (1881) 133, t. 5 f. 83; CLARKE, Trans. Linn. Soc.
Bot. 1 (1880) 583; Racts. Pterid. Buit. (1898)
7: v. A. v. R. Mal. Ferns (1908) 116; BACKER &
PostH. Varenfl. Java (1939) 257, f. 66; SELLING,
Svensk. Bot. Tidskr. 41 (1947) 431-450; Ho tr.
Ferns Mal. (1955) 51, f. 7.—Acrostichum digitatum
LINneE, Sp. Pl. (1753) 1068.—Actinostachys digitata
WALL. ex REED, Bol. Soc. Brot. 21 (1947) 130.—
Fig. 3a-e.
42 FLORA MALESIANA
[ser. Il; vole
Rhizome very short, creeping or suberect, 3-4
cm below surface of ground, bearing many
crowded fronds, apex clothed with brown hairs
under 2 mm long. Fronds erect, unbranched,
grass-like, 20-35 cm long, base (stipe) slender and
triquetrous, rest winged, greatest width 24 mm,
r,
Fig. 3. Schizaea digitata (L.) Sw. a. Habit,
x \4, b. sorophores, nat. size, c. detail of soro-
phore, x 5, d. spore, x 300, e. detail of lower
surface showing stomata, x 5.—S. inopinata
SELLING. f. Detail of sorophore, x 18,
g. spore, x 300, A. flattened part of exospore,
x 200, i. lower surface of leaf, showing stomata,
x 5 (a, e For. Bot. Burma 7670, bB-c MATTHEW
s.n. (SING), f SYNGE S606, g SYMINGTON CF 37414,
h after SELLING 1946).
costa very prominent on lower surface of winged
portion and slightly grooved on upper surface, 2-
celled glandular hairs abundant on lower surface
of wing, stomata in a close even single row on each
side of costa. Sorophores all attached very close
together (apparently digitate), 5-18, all about
equal in length, commonly 214-5 cm long (on
stunted plants sometimes shorter), little over 1 mm
wide, edges thin, entire, glabrous; sporangia small,
apparently in four rows and completely covering
lower surface of sorophore; spores small, finely
and evenly obliquely striate.
Type: Herb. Hermann, Ceylon (BM).
Distr. Ceylon, NE. India, Siam, Indochina,
Micronesia; in Malaysia: throughout except for
East Java and Lesser Sunda Isl.
Ecol. In lightly shaded forest, rubber estates,
etc., sealevel to 1200 m, rarely very abundant.
Notes. The species was formerly credited with
a much wider distribution, due to confusion with
other species. SELLING has distinguished most of
the latter, and has given a comparative survey of
the group and of individual distributions (/.c.).
Actinostachys boninensis NAKAI (J. Jap. Bot. 13,
1937, 140) appears to differ from S. digitata only
in the greater number of sorophores (to 30)
which are shorter (8-40 mm long); I have seen no
specimens.
3. Schizaea inopinata SELLING, Svensk Bot.
Tidskr. 40 (1946) 274, f. 1-7; Hottr. Ferns Mal.
(1955) 52.—Actinostachys inopinata REED, Bol.
Soc. Brot. 21 (1947) 130.—Fig. 3f-i.
In habit like S. digitata, differing as follows:
fronds to 244 mm wide, with a double (occasional-
ly triple) row of stomata on each side of the costa
on the lower surface, the wings thicker and more
rigid and the edges reflexed on drying, the costa
not so strongly raised and rather variable;
sporangia much larger, in two rows; spores much
larger, with broad irregular longitudinal ridges.
Type: Henderson 19460, Gua Tipus, Chigar
Perah, Pahang (SING, K).
Distr. Micronesia, in Malaysia: Sumatra,
Malaya, Borneo, Philippines (Bohol), W. New
Guinea.
Ecol. On limestone crags, sealevel to 300 m.
Note. FosBerG, correctly reporting the oc-
currence of the species in Micronesia, considered
it conspecific with S. ponapensis HOSOKAWA (Am.
Fern J. 40, 1950, 145). I have examined the
isotype of S. ponapensis in the Arnold Arboretum
Herbarium, and find that, apart from its much
smaller size (fronds to 8 cm long, sorophores to
8 mm long) it differs in having a hardly raised
costa with a rather widely-spaced single row of
stomata oneach side of it. The specimen corresponds
well with the description of S. spirophylla TROLL.
and I have placed S. ponapensis as a synonym
of that species. I saw no spores.
4. Schizaea spirophylla TRoLL, Flora 128 (1937)
343, fig. 2-6.—S. ponapensis HOSOKAWA, Trans.
Nat. Hist. Soc. Formosa 31 (1941) 39.
Rhizome short, apex covered with dark hairs.
Fronds unbranched, sometimes twisted, 4-8 cm
mec; 1959] | Schizaeaceae (Holttum) 43
Fig. 4. Schizaea dichotoma (L.) SM. a. Habit, x ?/3, b. rhizome, x 2/3, ¢. sorophores, x 7, d. lower
surface of leaf, x 14.—S. wagneri SELLING. e. Habit, x 4/,, f. detail of sorophore, x 14, g. lower sur-
face of leaf, x 27.—S. malaccana BAKER. h. Sorophores, x */;, i. lower surface of leaf showing stomata,
x 27.—S. fistulosa LaBILL. j. Sorophores, X 4/3, k. detail of sorophore, x 7, /. lower surface of leaf,
showing stomata, x 27 (a Hose 199, b BRAss 8583, c-d FMS 199, e-g GRETHER & WAGNER 4177, h-i
SFN 1100, j-7 CLEMENS 10729).
long, about 114 mm wide, costa very slightly rows (sometimes apparently in 4 rows near the
prominent on the lower surface, with a distinct middle); spores as in S. digitata.
flat wing on each side of it; stomata in one row Type: Troll, Ambon (M).
on each side of the costa, rather widely spaced Distr. Micronesia (Ponape); in Malaysia:
in the rows; sorophores 1-3, 4-8 mm long (some- Moluccas (Ambon).
times longer?), glabrous, the sporangia in two Ecol. Growing in moss-cushions on trees.
44 FLORA MALESIANA
[ser. I, volosi4
Note. See note under S. inopinata. The twisting
of the fronds of S. spirophylla is probably not a
constant character; such twisting is also common,
but not universal, in S. digitata. Stunted plants
of the latter may superficially resemble S. spiro-
phylia but differ in costa and stomata and in more
crowded sporangia.
5. Schizaea wagneri SELLING, Svensk Bot. Tidskr.
40 (1946) 278, f. 8-11; Hottr. Ferns Mal. (1955)
52.—Actinostachys wagneri REED, Bol. Soc. Brot.
21 (1947) 131.—S. paucijuga Hoitr. Gard. Bull.
Sing. 11 (1947) 267.—Fig. 4e-g.
Rhizome short, apex clothed with slender
brown hairs 114 mm long. Fronds simple, 6—20
cm long, base terete, upper part winged and in
all 14-34 mm wide, with rather broad midrib
prominent on the lower surface, the stomata
rather widely spaced in one row close to each side
of the midrib. Sorophores 2-5, 7-15 mm long,
edges smooth and glabrous, sporangia in two rows,
mixed with brown hairs; spores finely verrucose.
Type: Grether & Wagner 4177, summit of
Mt Tjajiak, 600 m, Manus Isl., Admiralty Is.
(S-PA, dupl. at K).
Distr. Malaysia: Malaya (P. Rumbia in Perak,
Singapore) Borneo, Ambon, W. New Guinea, and
Admiralty Is.
Ecol. “Epiphytic in mosses on stumps and
bases of trees; abundant at one place, but ex-
ceedingly inconspicuous” (GRETHER & WAGNER).
In forest near sea (Western New Guinea and
Borneo).
6. Schizaea fistulosa LABILL. Novy. Holl. Pl. Spec.
2 (1806) 103, t. 250 f. 3; PRANTL, Unters. Morph.
Gefasskr. 2 (1881) 135, excl. var. malaccana and
var. robusta; v. A. v. R. Mal. Ferns (1908) 116;
C. Cur. & Hott. Gard. Bull. S. S. 7 (1934) 210.—
Microschizaea fistulosa REED, Bol. Soc. Brot. 21
(1947) 134.—S. propinqua A. CUNN. in Hook.
Comp. Bot. Mag. 2 (1836) 362.—S. australis
Gaupb. Ann. Sc. Nat. Bot. 5 (1825) 98.—S.
chilensis Putt. Linnaea 30 (1859-60) 207.—Fig.
4j-l.
Rhizome short-creeping, young parts clothed
with shining brown hairs 2-3 mm long; stipes very
crowded; fronds unbranched, 9-18 (rarely to 30)
cm long below the fertile part, width to about
1 mm, upper surface rather deeply grooved, lower
surface almost evenly rounded and bearing two
rows of stomata which are not in grooves; surfaces
bearing scattered glandular hairs the bases of
which are slightly prominent. Sorophores all about
equal, arranged in a distinctly pinnate manner
along the distal 10-20 mm of the axis of the frond,
8-20 pairs, lowest often forked, 4-6 mm long,
edges much reflexed and bearing many coarse
forward-pointing hairs; sporangia in two rows,
without hairs; spores smooth.
Type: Labillardiére, Australia (F?; not seen).
Distr. Madagascar, Australia, Tasmania, New
Zealand, Fiji, New Caledonia, Chile; in Malaysia:
Borneo (Mt Kinabalu), New Guinea.
Ecol. In alpine bogs at 2400-3750 m, and on
fine rock-screes.
Note. Some New Guinea specimens have been
distributed as S. papuana BRAUSE, which is here
placed as a synonym of S. malaccana.
7. Schizaea malaccana BAK. Syn. Fil. (1868) 428;
BEDDOME, Ferns Br. India (1870) t. 255; Handb.
(1883) 452; TANsLEY & Cuick, Ann. Bot. 17
(1903) 493-510; v. A. v. R. Mal. Ferns (1908)
116; Hoxttr. Ferns Mal. (1955) 52, fig. 8.—S.
fistulosa var. malaccana PRANTL, Unters. Morph.
Gefasskr. 2 (1881) 136.—S.papuana BRAUSE, Bot.
Jahrb. 56 (1920) 211.—Microschizaea malaccana
REED, Bol. Soc. Brot. 21 (1947) 134.—Fig. 4h-i.
var. malaccana.
Habit of S. fistulosa, differing as follows: hairs
on rhizome pale brown, fronds 6-15 cm long, less
than 1 mm wide, apical fertile part of axis c. 5mm
long, upper surface of frond nearly flat or shal-
lowly grooved, lower surface when dry with two
small grooves in which the stomata are situated;
sorophores 4-10 pairs, lowest 4-5 mm long, upper
ones shorter.
Type: Cuming 379, Mt Ophir, Malaya (K, BM).
Distr. Malaysia: Malaya, Borneo, Ambon, W.
New Guinea.
Ecol. In open mossy places on mountain
ridges, or in moss cushions on trees at 800-2000
m; also in swamp forest in Sarawak at lower
altitudes.
var. robustior C. Cur. Gard. Bull. S.S. 7 (1934)
210.—S. hallieri RicHTER, Med. Rijksherb. n. 28
(1916) 24, t. 1 f. 5, etc.; Math. Naturw. Ber.
Ungarn 31 (1916) 24, 28, t. 1 f. 5 etce.—Mi-
croschizaea hallieri REED, Bol. Soc. Brot. 21 (1947)
134.
Larger than var. malaccana; fronds to 25 cm
long, 1 mm or more wide; sorophores to 15 pairs,
spread along 10 mm of the axis, lowest sorophores
5-10 mm long.
Type: Clemens 10919, Mt Kinabalu, Borneo
(BM, Bo).
Distr. Malaysia: Malaya (Gunong Tahan),
Borneo, W. New Guinea.
2. LYGODIUM
Swartz in Schrader, J. Bot. 1800 pt 2 (Nov.—Dec. 1801) 106, nom. cons.; PRANTL,
Unters. Morph. Gefasskr. 2 (1881) 60; CopELAND, Gen. Fil. (1947) 23; PICHI-
SERMOLLI, Webbia 12 (1956) 10, preprint (1955).—Ramondia MIRBEL, Bull.
Soc. Philom. Paris 2 (Feb.—Mar. 1801) 179.—Ugena Cavan. Ic. Descr. Pl. 6 (Oct.
1801) 73.—Odontopteris BERNH. in Schrader, J. Bot. 1800 pt 2 (Nov.—Dec. 1801)
127, t. 2 f. 4.—Ripidium BERNH. /.c. 127, t. 2 f. 3.—Gisopteris BERNH. /.c. 129,
Dec. 1959] Schizaeaceae (Holttum) 45
t. 2 f. 1—Hydroglossum WitLp. Abh. Kurfiirstl. Mainz. Ak. Niitzl. Wiss. Erfurt
2, pt 4 (1802) 13, 20.—Hugona Cav. ex Roemer, Arch. Bot. 2 (1801-02) 486.—
Cteisium Micux, Fl. Bor. Am. 2 (1803) 275.—Vallifilix THouARS, Gen. Nov.
Madag. (1808) 1.—Lygodictyon J. SMitH in Hook. Gen. Fil. (1842) t. 111 B.—
Fig. 5-15.
Rhizome creeping, below ground surface, protostelic, short with fronds very
close together or longer with spaced fronds, young parts densely covered with
rather thick rigid multiseptate hairs, branching dichotomous; fronds borne in
two rows on upper surface of rhizome, roots mainly from lower surface. Fronds
of young plants erect, once or twice dichotomously branched and bearing usually
palmately lobed leaflets; fronds of older plants with slender elongate twining
rachises formed by a succession of very unequal dichotomies, at least the upper
part of the rachis (except in L. polystachyum) bearing two narrow wings towards
the adaxial side, the surface between the wings flat or slightly raised and papillose;
all branch rachises and stalks of leaflets similarly winged (fig. 13d), the wings always
interrupted to join with those of a lateral branch; primary rachis-branches always
short, usually hardly developed, ending in dormant apices which are covered with
hairs (such apices proliferous if the main rachis beyond them is injured), each
primary branch bearing a pair of secondary branches which bear the leaflets;
sterile /eaflets (or their lobes) with costa and oblique lateral veins which are 1-3
times forked (anastomosing in a few species), edges entire or serrate (pinnatifid
only in L. polystachyum); fertile leaflets often with contracted lamina, bearing
narrow sorophores spreading from the edges of the lamina at the ends of most
of the veins; edges of sorophores serrate, the main vein in each sorophore bearing
alternate short lateral veins each of which bears a single sporangium protected
by a separate indusium attached along the vein and opening forwards; sporangia
oblong-ovoid with a short lateral stalk, the annulus at the narrower end which
is directed away from the margin of the sorophore, splitting longitudinally when
ripe; spores trilete, pale, variously sculptured on the surface, lacking perispore.
Gametophyte thalloid, sometimes asymmetric; antheridia larger and more complex
in structure than in most leptosporangiate ferns.
Distr. Pantropic, comprising c. 40 spp., also extratropical southwards in New Zealand and S. Africa,
northwards in Japan and in eastern U.S.A. to Massachusetts.
Morph. Owing to the peculiar structure of the climbing leaves, it is difficult to apply the usual descrip-
tive terms pinna and pinnule to them, especially where the branching of the leafy parts is dichotomous.
The following terms are here used. The climbing rachis of the frond (sympodial in structure but for
convenience considered as a unit) bears alternate short primary branches, each ending in a dormant
apex and bearing a pair of apparently opposite secondary branches. The secondary branches may bear
leaflets or tertiary branches pinnately arranged, or they may be once or more times dichotomously
branched.
The narrow wings on rachis-branches of all orders, those of the ultimate branches joined to the edge
of the lamina, correspond with the wings which are the only lamina in fronds of Schizaea, but do not
have the very regular single rows of stomata found in Schizaea (there are irregularly scattered stomata
on both sides of a wing in Lygodium). If the sorophores of Lygodium are regarded as homologous with
the fertile leaflet-lobes of the paleozoic fossil genus Senftenbergia, the lamina of Lygodium may be con-
sidered as a specialized development of the rachis-wing consequent on the reduction of the original
lamina-lobes to sorophores.
The dormant apices of the primary rachis-branches are covered with septate hairs. In a group of
Malaysian species these hairs have swollen bases, each base formed of a mass of cells; these species are
L. borneense, L. longifolium, L. auriculatum, L. trifurcatum, and L. dimorphum. So far as I know, this
type of hair has not hitherto been reported in Lygodium.
Taxon. There are four pairs of species which in some measure intergrade. These need experimental
study in cultivation to discover how much variation is due to environmental conditions, and also cyto-
46 FLORA MALESIANA
[ser. Il, vole
logical study. It may be that natural hybridization occurs, and as tetraploids as well as diploids have
been already discovered in L. japonicum and L. circinnatum they may occur also in other species, leading
to the possibility of the formation of sterile triploids; apogamy has however not yet been discovered
in the genus. The pairs of species which intergrade are: L. flexuosum and L. japonicum; L. flexuosum
and L. salicifolium; L. borneense and L. auriculatum; L. dimorphum and L. trifurcatum. The following
species are very distinct: L. polystacnyum, L. microphyllum, L. circinnatum, L. merrillii, and L. versteegii.
KEY TO THE SPECIES
1. Secondary rachis-branches pinnate, leaflets 10-15 on each side. Sterile leaflets evenly pinnatifid
throughout
1. L. polystachyum
1. Secondary rachis- branches pinnate with fewer leaflets, or r dichotomous. Sterile leaflets simple, palmate,
or lobed near the base only.
2. Venation of sterile leaflets free.
3. Primary rachis-branches 4-10 mm long below the pair of secondary branches. Rhizome wide-
creeping, fronds distinctly spaced.
4. Secondary branches simply pinnate. Leaflets articulate at the base and ultimately deciduous.
4. Secondary branches amply bipinnate. Leaflets not articulate at the base
2. L. microphyllum
3. L. japonicum
3. Primary rachis-branches hardly elongated, the pair of secondary branches thus almost sessile
on the main rachis. Rhizome short, fronds close together.
5. Secondary rachis-branches regularly pinnate, normally with 3-5 leaflets on each side of the
axis (young or depauperate fronds may have fewer leaflets).
6. Leaflets all about equal and all stalked (terminal one sometimes geminate), not auricled or
branched at the base, or rarely with short spreading basal lobes;
their junction with the lamina .
leaflet-stalks thickened at
4. L. salicifolium
6. Lateral leaflets larger towards base of secondary branch, smaller distal ones sessile, basal ones
stalked and usually auricled or with obliquely genau lobes, or with subsidiary leaflets below
them;
leaflet-stalks not thickened at apex
. 5. L. flexuosum
5. Secondary branches simple or dichotomously branched one or more times (in some cases, by
alternate unequal dichotomy, subpinnate with 1-3 lateral leaflets). Leaflets simple, forked, or
palmately branched, never thickened at apex of stalk.
7. Leaflets strongly cordate on outer side at base.
8. Lamina of fertile leaflets reduced to a narrow wing along midrib and bases of veins, each vein
usually with an apical sorophore.
9. Fertile tertiary branches evenly deltoid in outline with quaternary leaflets of increasing length
towards the base. Sterile leaflets strongly auriculate-cordate at base .
6. L. dimorphum
9. Fertile leaflets not deltoid in outline, 10-15 mm wide (including sorophores), sometimes with
1 or 2 long branches at the base. Sterile leaflets more or less cordate at the base.
7. L. trifurcatum
8. Lamina of fertile leaflets not reduced to a narrow wing along midrib and veins.
8. L. auriculatum
7. Leaflets not strongly cordate at base on outer side.
10. Secondary branches once forked (rarely simple, or one branch forked again). Leaflets always
simple. Spores smooth
9. L. borneense
10. Secondary branches at least twice forked (or Tower branches bearing single 3—5-lobed leaflets),
sometimes sub-pinnate with 1—2 tertiary members on each side and a terminal leaflet or pair
of leaflets. Leaflets often deeply 2-lobed to palmate. Spores verrucose.
11. Margins of sterile leaflets serrate, not much thickened, veins ending in the teeth. Spores
coarsely and irregularly verrucose. Dormant apices of primary branches not sunk, hairs
with swollen base
10. L. longifolium
11. Margins of sterile leaflets entire, thickened but not ‘vascular, the raised veins joining the
thickened edge. Spores finely and evenly verrucose. Dormant apices sunk, hairs lacking swollen
bases :
2. Venation of sterile leaflets retienlate:
11. L. circinnatum
12. Secondary rachis-branches (at least the upper ones) elongate and pinnate.
13. Leaflets on stalks to 3 cm long, palmately lobed or equally bilobed (basal secondary branches
may bear only one palmately lobed leaflet on a longer stalk)
13. Leaflets on much shorter stalks, simple or the apical ones forked
12. L. merrillii
14. L. reticulatum
12. Secondary branches bearing 3 leaflets (lateral ones sometimes forked), all arising 1-3 mm from
dormant apex of primary rachis-branch
1. Lygodium polystachyum WALL. ex Moore, Gard.
Hook. Sec. Cent. (1861) t. 76;
Ferns Br:
India (1868) t. 300; Handb. (1883) 458, f. 284;
Chron. (1859) 671;
Syn. Fil. (1868) 438; BEDDOME,
13. L. versteegii
v. A. v. R. Mal. Ferns (1908) 113; Copev. Philip.
J. ‘Sc. Bot. 4 (1909) 19; Tarp. & Go Car El:
Gén. I.-C. 7 (1939) 40; Hottr. Ferns Mal. (1955)
56, f. 10; ALSTON & HoLtTrT. Reinwardtia 5 (1959)
Dec. 1959]
Schizaeaceae (Holttum) 47
11. — Hydroglossum pinnatifidum WiLLD. Abh.
Kurf. Mainz. Ak. Wiss. Erfurt 2, pt 4 (1802) 21,
p.p.—L. pinnatifidum (non (WILLD.) Sw.) PRANTL,
Unters. Morph. Gefasskr. 2 (1881) 83, t. 1
f. 11.—Fig. 5c, 8a-—c.
Fig. 5. Spores of Lygodium. a. L. longifolium
(WILLD.) Sw., b. L. borneense v. A. v. R., c. L.
polystachyum WALL. ex Moore, d. L. circinnatum
(Burm. f.) Sw., e-f. L. microphyllum (CaAv.)
R. Br., lateral and upper view. All x 330.
Rhizome short-creeping, 3-5 mm _ diameter,
densely clothed with spreading black hairs.
Juvenile fronds once or twice dichotomous;
stipes up to 30 cm long to the first dichotomy,
brown, withshort very slender hairs mixed with thick
longer multicellular ones; ultimate branches leafy
like the secondary rachis-branches of climbing
fronds. Rachis of climbing fronds 214 mm dia-
meter, shortly hairy, not winged; primary rachis-
branches very short, ending in a dormant apex
covered with brown hairs (the apices of lower
primary branches sometimes proliferous); second-
ary rachis-branches 20-30 cm long, not winged,
shortly hairy, bearing 10-15 leaflets on each side
and a similar terminal leaflet; sterile leaflets
3144-714 cm long, 114-2 cm wide, apex rather
abruptly narrowed and rounded, base truncate or
cordate, jointed to a hairy stalk 1-4 mm long,
sides lobed half-way to the costa or rather more,
lobes 4-5 mm wide, oblong with rounded apex,
each lobe with a sinuous costule bearing oblique
forked lateral veins, costa and costules bearing
scattered stiff hairs on both surfaces, the costa
also with shorter hairs; fertile leaflets like the
sterile but the distal half or more of each lobe
narrowed to about 2 mm wide, the narrow part
(sorophore) 4-10 mm long, bearing sporangia
on the under surface; indusia bearing scattered
stiff hairs; spores finely and evenly verrucose.
Type: Wallich 177, Penang (K).
Distr. Assam, Burma, Siam, Yunnan, Indo-
china; in Malaysia: northern half of Malay
Peninsula.
Ecol. In lowland forest, climbing trees to a
considerable height; in the Malay Peninsula
especially on limestone except in the extreme
north (Kedah) where it is locally common
apparently primary forest. (Further sor
limestone provides a drier habitat than ott
rocks, and on it occur a number of species whic
have their main distribution in the seasona.
climate north of Malaya.)
2. Lygodium microphyllum (CaAv.) R. Br. Prod.
FI. Nov. Holl. (1810) 162; Br. En. Pl. Jav. (1828)
253; CLARKE, Trans. Linn. Soc. Bot. 1 (1880) 583;
Bepp. Handb. (1883) 455, t. 282; ALSTON &
Hott. Reinwardtia 5 (1959) 12.—Ugena micro-
phyllatGAve ice Descra Elon sO) a76: sty 5955
C. Cur. Dansk Bot. Ark. 9, n. 3 (1937) 30.—L.
scandens Sw. in Schrader, J. Bot. 1800 pt 2 (1801)
106, p.p. excl. syn. LINN.; BEDD. Ferns S. India
(1863) t. 61; Hoox. Syn. Fil. (1868) 437, p.p.;
PRANTL, Unters. Morph. Gefasskr. 2 (1881) 81,
t. 6 f. 101; CHrRist, Farnkr. Erde (1897) 354, f.
1116; Racts. Pterid. Buit. (1898) 8; v. A. v. R.
Mal. Ferns (1908) 113; Philip. J. Sc. 11 (1916)
Bot. 116; HEyNne, Nutt. Pl. (1927) 97; BAck.
Krakatoa (1929) 254; OcHsE & BAKH. Veg.
DIE-E 9s) "6o7- Burke Dict. 2) C1935)" 1378:
BACKER & PostTH. Varenfl. Java (1939) 258;
OGatTaA, Ic. Fil. Jap. 7 (1936) t. 324; Hott.
Ferns Mal. (1955) 58, f. 12.—Ophioglossum
filiforme Roxs. Calc. J. Nat. Hist. 4 (1844) 476,
t. 26 f. 3.—L. scandens var. microphyllum (CAV.)
Luerss. J. Mus. Godeffr. 6 (1874) 4.—L. scandens
var. intermedium Ces. Att. Ac. Sc. Fis. Nat.
Napoli 7 (1876) 33.—Fig. 5e-f, 6, 7.
Rhizome wide-creeping, dichotomously branch-
ed, 2!14 mm diameter, densely clothed with short
spreading brownish-black hairs. Juvenile fronds
small, commonly once dichotomous (the stipe
distinctly winged below the dichotomy), each
branch bearing a 4-lobed leaflet not jointed at the
base, lobes 3—5 cm long and c. 5 mm wide, thin,
glabrous, edges crenately toothed (teeth larger
towards apex where veins are unbranched).
Rachis of climbing fronds glabrous, commonly
2-3 m long, hardly 114 mm diameter; primary
branches 4 mm or more long, ending in a dormant
apex covered with dark brown hairs; secondary
rachis-branches pinnate, in all to about 15 cm
48 FLORA MALESIANA [ser. IT, volxaes
Kee
RIT AES Ant are
Uy: eee
ts
Fig. 6. Lygodium microphyllum (Cav.) R. Br. and L. salicifolium PR. (larger leaflets on left) on edge ofa
thicket in the Botanic Gardens, Singapore (HOLTTUM).
Dec. 1959]
Fill
Q
Schizaeaceae (Holttum) 49
Fig. 7. Lygodium microphyllum (Cav.) R. Br. a. Fertile leaf, x %4, b. sterile leaf, x 34, c. rhizome,
x 34, d. detail of base of leaflet, x 7 (a FloyD NGF 5566, b-d cult. Kew from Ceylon).
long, with 3-6 stalked leaflets on each side
(stalks 2-4 mm long) and a similar or geminate
terminal leaflet; /eaflets quite glabrous, mostly
ovate (sterile leaflets often elongate with broader
base on young plants), 1-4 cm long (sterile ones
sometimes to 6 cm), 6-18 mm wide, edges of
sterile ones minutely crenate, a joint always
present at base of blade, where the wing which
in other species connects stalk and lamina is
constricted; fertile leaflets usually shorter than
sterile but with lamina hardly narrowed, soro-
phores 4-6 mm long; spores witha raised reticulum
on the outer surface.
Type: Née, Luzon (Ma).
Distr. Tropical Africa, SE. Asia (north to
Bengal and Hong Kong and the Riu Kiu Isl.),
Melanesia (Solomon Isl., New Caledonia), N. and
E. Australia south to N. S. Wales; in Malaysia:
throughout, but few records from the Lesser
Sunda Isl. (certainly to Flores).
50 FLORA MALESIANA [ser. II, vol. 1}
VV.
SS
Wd
me
We
Ny
SEA
t)
a
ERS
‘S
ot
ie
Seay
Y% :
ee, Wy :
ee NWMWn, *
ea UN BSNS Ak ae
LL py Wn
Zo Bigs
SSN
“any
Su
MONA Sy
Se NTN), Lys
Fig. 8. Lygodium polystachyum WALL. ex Moore. a. Sterile leaf, x #/;, b. fertile leaflet, x */;, c. ditto,
detail, x 214.—L. japonicum (THUNB.) Sw. d. Fertile leaf, x #/;, e. sterile leaf, x 3/,, f. rhizome, x 4/5
(a BALANSA 168, b-c MATTHEW s.n., d, f cult. Kew, e SAVINIERRE 71).
Dec. 1959]
Schizaeaceae (Holttum) . 51
Ecol. Edges of secondary forest, or climbing
woody plants in open places, sometimes as a weed;
in clay soil, or in swamps in regions subject to a
dry season; from the lowlands to c. 1300 m.
Vern. Ribu-ribu, M, sélada, capey papua, capay
alus, M, paku kawat, Sum., paku hata béjas (bias),
paku hata leutik, S, nitong-puti, Tag., paku rambat,
Bali, sickey, Luhea, gomoha papua, Ternate,
paku kawa, Ambon, paloge, N. Celebes.
Uses. Native medicine (leaves macerated and
mixed with lime for open wound), magic; young
leaves edible; rachises of old leaves used for string
and for plaiting.
3. Lygodium japonicum (THUNB.) Sw. in Schrader,
J. Bot. 1800, pt 2 (1801) 106; Bepp. Ferns S. India
(1863) 21, t. 64: CLARKE, Trans. Linn. Soc. Bot. |
(1880) 584; PRANTL, Unters. Morph. Gefasskr.
2 (1881) 68, t. 1 f. 10, 15; CHrist, Farnkr. Erde
(1897) 355, 356, f. 1122; Racis. Pterid. Buit.
(1898) 8; Hore, J. Bomb. Nat. Hist. Soc. 15
(1903) 106; v. A. v. R. Mal. Ferns (1908) 114;
Merk. FI. Manila (1912) 60; Doin, Bibl. Bot. 20,
Heft 85 (1914) 211, f. 50; HaAtnes, Bot. Bih. & Or.
6 (1924) 1210; OcarTa, Ic. Fil. Jap. 7 (1936) t.
322; BACKER & PostH. Varenfl. Java (1939) 259;
ALSTON & HoLtr. Reinwardtia 5 (1959) 14.—
Ophioglossum japonicum THUNB. Fl. Jap. (1784)
328.—Hydroglossum japonicum (THUNB.) WILLD.
Abh. Kurf. Mainz. Ak. Wiss. Erfurt 2, pt 4 (1802)
26.—L. dissectum Desv. Mag. Ges, Naturf. Fr.
Berl. 5 (1811) 308.—L. microstachyum Desvy. lI.c.;
NAKAI, Bot. Mag. Tokyo 39 (1925) 182; OGaTa,
Ic. Fil. Jap. 7 (1936) t. 323.—L. pubescens KAULF.
En. Fil. (1824) 47, t. 1 f. 4.—L. chaerophylloides
Desv. Mém. Soc. Linn. Paris 6 (1827) 205.—L.
cochinchinense Desy. ibid. 206.—L. tenue BL.
En. Pl. Jav. (1828) 254.—L. microphyllum Link,
Hort. Berol. 2 (1833) 141.—L. japonicum f.
elongata v. A. v. R. Bull. Jard. Bot. Btzg II, n. 1
(1911) 10, t. 3; Mal. Ferns Suppl. (1917) 117.—
L. japonicum var. microstachya (DESV.) TARD. &
C. Cur. Fl. Gén. I.-C. 7 (1939) 38.—L. mearnsii
CopEL. Philip. J. Sc. 3 (1908) Bot. 37.—Fig. 8d-f.
Rhizome wide-creeping, dichotomously branch-
ed, 2-5 mm diameter, densely clothed with
dark brown hairs, fronds commonly 5-10 mm
apart. Juvenile fronds erect, the first branching
an unequal dichotomy (always?), the two main
branches of large fronds bipinnate, deltoid in
outline, with palmatisect leaflets, their margins
doubly serrate. Rachis of climbing fronds hardly
2 mm diameter, glabrous apart from minute hairs
on the flattened adaxial surface between the
Narrow wings; primary rachis-branches 3-10 mm
long, the dormant apex covered with pale hairs;
secondary branches of fronds on young or stunted
plants pinnate, on well-grown fronds bipinnate or
tripinnate, deltoid in outline, commonly 12 cm
long and wide, rachises densely short-hairy on
the upper surface and bearing fewer longer hairs
elsewhere; sterile tertiary /eaflets of lower rachis-
branches palmate with 5—7 lobes, the middle lobe
much longer than the laterals, tertiary leaflets
higher up the leaf trilobed with an elongate
middle lobe or pinnate with small oblique and
often lobed quaternary leaflets and a usually
deltoid-pinnatisect terminal leaflet about 3 cm
long, edges acutely biserrate, apex obtuse or
subacute; stalks of leaflets to 3 mm long, never
articulate or thickened at the apex; costae
usually bearing long scattered hairs, veins and
surfaces usually glabrous but sometimes short-
hairy; fertile secondary branches tripinnate, the
leaflets smaller than sterile ones, sorophores 2-12
mm long; indusia glabrous or with a few hairs
if the lamina is hairy; spores finely verrucose.
Type: Herb. Thunberg, Japan (Ups).
Distr. Ceylon, from Himalayas (Kashmir
eastwards) to Chekiang in N. China, Korea,
Japan (Nagasaki) and southwards to Siam and
Indochina and southern China, naturalized in
Florida and Texas; in Malaysia: Banka, Central
and East Java, Celebes, Philippines, Moluccas
(Ambon, Ternate, Banda Is!., Sula Isl.), Lesser
Sunda Isl. (to Timor), and New Guinea.
Ecol. Climbing in secondary vegetation, at
altitudes up to 2550 m; only found native in
regions with a pronounced dry season, during
which fronds perhaps die (no records in Malaysia);
absent from Sumatra, Borneo and the Malay
Peninsula (except perhaps the extreme north).
Small forms are not always clearly distinct from
L. flexuosum.
Vern. Pakis kémbang, J, paku areuj, hata
kawat, S, nito, nitong puti, Tag., madik silai,
durhawa, babar, talsiga, Alor.
4. Lygodium salicifolium Pres_, Suppl. Pterid.
(1845) 102, p.p. excl. pl. WALLICH p.p. and syn.
RHEEDE; PRANTL, Unters. Morph. Gefadsskr. 2
(1881) 79; v. A. v. R. Mal. Ferns (1908) 113;
Bull. Jard. Bot. Btzg III, 5 (1922) 213; BACKER
& Postu. Varenfl. Java (1939) 258; TARD. &. C.
Cur. FI. Gén. I.-C. 7, 2 (1939) 41; ALSTON &
Ho.ttr. Reinwardtia 5 (1959) 14.—L. kingii
CopEL. Philip. J. Sc. 6 (1911) Bot. 68; v. A. v. R.
Mal. Ferns Suppl. (1917) 117.—L. pinnatifidum
sensu RAc. Fl. Buit. (1898) 7, p.p.—L. flexuosum
sensu Hott. Ferns Mal. (1955) 57, p.p.—Fig.
6, 10, 13a—b.
Rhizome and juvenile fronds as in L. flexuosum,
except that the rachis is thickened at its junction
with the midribs of the lobes of the leaflets. Rachis
of climbing fronds to 2 mm diameter, to 10 m
long; primary rachis-branches always very short
(hardly measurable), ending in a dormant apex
covered with brown hairs; secondary rachis-
branches normally pinnate, rarely somewhat
bipinnate and then the tertiary branches bearing
one or more pairs of short spreading lateral
leaflets (jointed at the base) and a large terminal
one; secondary branch-system usually consisting
of about 4 (rarely to 6) leaflets on each side, and
a terminal deeply bilobed leaflet (or a pair of
leaflets), all leaflets of about equal size and all
stalked, the stalks 2-10 mm long and thickened
at junction with lamina (old leaflets sometimes
deciduous but not regularly so as in L. micro-
phyllum); leaflets 4- 15 cmlong, 14-2 cm wide, acute
2 FLORA MALESIANA [ser. Tl, voliat=
Fig. 9. Lygodium dimorphum Cork. a. Part of fertile leaf, x %, b-c. parts of sterile leaves, x 3%,
d. detail of a, x 3.—L. flexuosum (L.) Sw. e. Sterile leaf, x %, f. fertile leaf, x % (@ LABILLARDIERE
s.n., b-c CARR 12479, d PEEKEL 6, e Hose 5024, f HALLER s.n. from Java).
Dec. 1959]
7 Schizaeaceae (Holttum) 53
and attenuate or subobtuse, edges of sterile
leaflets finely crenate-serrate, base truncate to
cordate, lamina thicker than in L. flexuosum;
upper surface of costae more or less hairy es-
pecially towards the base, lower surface often
glabrous, veins usually glabrous; sorophores 2-5
mm long, usually constricted at the base, often
with hairs on upper surface of midrib; indusia
glabrous; spores finely verrucose.
Fig. 10. ‘Columns’ of Lygodium salicifolium PR.
on scattered pole trees of mostly Ilex cymosa,
surrounded by a dense ground cover of ferns and
sedges, Nepenthes, and orchids, in an old crater
swamp in the Gajo Lands, N. Sumatra, c. 1200 m
(1937).
Type: Cuming 365, Singapore (W, K).
Distr. Assam, Siam, Indochina to Yunnan,
Formosa, south-east to New Guinea and Micro-
nesia; in Malaysia: Sumatra, Malay Peninsula,
Banka, W. Java, Borneo, and New Guinea.
Ecol. In open secondary vegetation, sometimes
in wet places, in the low country and to 1200 m;
reported in teak forest in West Java.
Note. There are specimens which are inter-
mediate between this species and L. flexuosum;
they may be hybrids. L. salicifolium occurs only
in regions with a short dry season, whereas L.
flexuosum will tolerate a longer dry season and
has a wider distribution. In Burma and Assam
very large forms of both species occur.
Vern. Hata, S, mintuh, Dayak, paku kawat,
Sum., akar sidin, M.
5. Lygodium flexuosum (L.) Sw. in Schrader, J.
Bot. 1800, pt 2 (1801) 106; ibid. 1801, pt 2
(1802) 304; Pres_, Suppl. Pterid. (1845) 100;
Bepp. Ferns S. India (1863) t. 63; CLARKE,
Trans. Linn. Soc. Bot. 1 (1880) 584; PRANTL,
Unters. Morph. Gefasskr. 2 (1881) 72, p.p.;
Bepp. Handb. (1883) 457, f. 283; v. A. v. R. Mal.
Ferns (1908) 114; Merr. Fl. Manila (1912) 61;
Domin, Bibl. Bot. 20, Heft 85 (1914) 209, f. 49;
Haines, Bot. Bih. & Or. 6 (1924) 1211; Back.
Onkr. Suiker 7 (1928) 1, t. 1; Burk. Dict. 2 (1935)
1378: "C."Cur: Dansk Bot. ‘Ark: 9, ‘pt 3 (1937)
30; BACKER & PosTH. Varenfl. Java (1939) 259;
Hott. Ferns Mal. (1955) 57, p.p.; ALSTON &
Ho.ttr. Reinwardtia 5 (1959) 15.—Ophioglossum
flexuosum LINNE, Sp. Pl. (1753) 1063.—Ophio-
glossum scandens LINNE, Sp. Pl. (1753) 1063, p.p.
—Ramondia flexuosa (L.) Mire. Bull. Soc. Philom.
Paris 2 (Feb.—Mar. 1800) 179, t. 12 f. 3.—Aydro-
glossum flexuosum (L.) WILLD. Abh. Kurf. Mainz.
Ak. Wiss. Erfurt 2 (1802) 23, t. 1 f. 3.—Hydro-
glossum pinnatifidum WiLLD. ibid. 21, p.p.—L.
pinnatifidum Sw. in Schrader, J. Bot. 1801, pt 2
(1803) 303; Hook. Syn. Fil. (1868) 438, p.p.—
L. semibipinnatum R. Br. Prod. Fl. Nov. Holl.
(1810) 162.—L. serrulatum Bi. En. Pl. Jav.
(1828) 254.—L. flexuosum var. setulosum TARD. &
C. Cur. Fl. Gén. I.-C. 7 (1939) 39.—Fig. 9e-f.
Rhizome short-creeping and densely covered
with roots, the stipes very close together; apex of
rhizome covered with dark brown to nearly black
hairs. Juvenile fronds once or twice dichotomous,
each branch bearing a single leaflet which is
deeply palmately 3-7-lobed, the lobes almost
equal, the base of the whole leaflet cordate, edges
serrate and sometimes crenately lobed. Rachis of
scandent fronds narrowly winged, flattened and
puberulous on the upper surface between the
wings; primary rachis-branches up to 3 mm long
(lower ones longest), dormant apex covered with
pale brown hairs; secondary rachis-branches
pinnate to somewhat bipinnate, narrowly ovate
to deltoid in outline, commonly about 15 cm long
and 8 cm wide; sterile leaflets of lower branches
palmate, often 5-lobed, base strongly cordate;
higher secondary branches bearing 3-5 (some-
times to 7) leaflets on each side and an apical
one, the apical and lower leaflets asymmetric or
more or less lobed at the base, the lowest often
with 2 or 3 (exceptionally to 6) separate quaternary
leaflets at its base; sterile leaflets 3-10 cm long,
8-15 mm wide above the lobed base, apex sub-
acute, edges serrate, lower leaflets stalked, upper
sessile, lamina rather thin; costae usually bearing
scattered long hairs, less often densely short-
hairy, veins often with scattered short hairs on
the lower surface, the lamina sometimes similarly
hairy; fertile leaflets smaller than sterile, soro-
phores 3-5 mm long (rarely up to 10 mm), at the
apices of small triangular lobes; indusia glabrous
or with a few hairs like those of the lower surface
of the lamina; spores finely verrucose.
54 FLORA MALESIANA
[ser. TL vole
Type: Hermann, Ceylon (BM).
Distr. Ceylon, from the Himalayas (Dehra
Dun eastwards) to southern China, Hong Kong,
Riu Kiu Isl., south and south-east to Melanesia
and northern Queensland, throughout Malaysia.
Ecol. In open places, climbing on shrubs, in
teak and bamboo forest, in low country and to
1000 m, not in shady evergreen forest. In very dry
or exposed places the veins and lamina are often
rather copiously hairy.
Vern. Ribu-ribu gajah, ribu-ribu bésar, ikat
sidin, M, paku ribu-ribu, Asahan, hata kémbang,
J, durhawa, Alor, nito, Tag., Visc., tatan, Orokawa
Horata (N.G.), zangi, Orokawa Mumuni.
Uses. For tying rice sheaves; in native medicine
for skin diseases and fever.
6. Lygodium dimorphum Cope-. Philip. J. Sc. 6
(1911) Bot. 67; RosENsT. in Fedde, Rep. 10
(1912) 343; v. A. v. R. Mal. Ferns Suppl. (1917)
116; Philip. J. Sc. 11 (1916) Bot. 116; ALSTON
& Hoitr. Reinwardtia 5 (1959) 18.—L. flexuo-
sum [non (L.) Sw.] GAuUDICH. in Freyc. Voy. Bot.
(1826) 298.—L. circinnatum var. trifurcatum
Curist, Monsunia | (1900) 93.—L. novoguineense
RosensTt. in Fedde, Rep. 9 (1911) 427.—L.
trifurcatum sensu v. A. v. R. Mal. Ferns (1908)
112, 802, p.p.—Fig. 9a-d.
Rhizome short-creeping, bearing fronds very
close together, apex densely covered with almost
black hairs. Juvenile fronds once dichotomous,
each branch bearing a deeply palmatisect leaflet,
lobes subequal, base cordate, edges rather ir-
regularly serrate and somewhat thickened. Rachis
of climbing fronds hardly 2 mm diameter, glabrous
or nearly so; primary rachis-branches very short,
ending in a somewhat projecting dormant bud
covered with light brown hairs having slightly
swollen bases; secondary rachis-branches bearing
sterile leaflets unbranched or more commonly once
dichotomous, those bearing fertile leaflets usually
sub-pinnate with a few tertiary branches; sterile
leaflets 10-18 cm long, simple or forked (less often
3-lobed), usually strongly cordate and auriculate
at the base on one side (sometimes with a separate
rounded leaflet replacing the cordate base), when
forked the lamina lobed to within 1 cm of the
base, leaflets or lobes 1-2 cm wide, tapering, acute,
irregularly doubly serrate, edge somewhat thicken-
ed, surfaces glabrous, sometimes sparingly
warty; fertile leaflets usually with the lamina
reduced to a narrow wing (0.2 mm wide) along
the costae and along each vein and its branches;
tertiary fertile branches deltoid in outline (5 cm
or more wide at the base), with quaternary leaf-
lets of increasing length below each terminal one;
sorophores 2-4 mm long, indusia glabrous; spores
minutely verrucose on an unevenly undulating
surface (always?).
Type: C. King 134, Papua (Micu, Bo).
Distr. Malaysia: Celebes(?), Moluccas(Ambon,
Rawak), New Guinea.
Ecol. “Climbing small trees to a height of
about 12 feet. Quite common in low wet places
along the coast’? (RUSSELL, on specimen from
E. New Guinea). Collections have also been made
inland, at altitudes up to 1000 m.
Vern. Cana, Motuan, gailei, Bragi, paku kawa,
Ambon.
Uses. Used for making arm- and leg-bands.
7. Lygodium trifurcatum BAK. in Hook. Syn. Fil.
(1868) 437; v. A. v. R. Mal. Ferns (1908) 112,
802, p.p.; WAGNER & GRETHER, Un. Cal. Publ.
Bot. 23 (1948) 27, t. 8; ALSTON & HoLtTrT. Rein-
wardtia 5 (1959) 17.
Scandent rachis about 114 mm diameter; primary
rachis-branches very short, hairs on the dormant
apex having swollen bases; sterile secondary
rachis-branches once or twice equally or sub-
equally dichotomous (12-20 mm to first dicho-
tomy), the leaflets simple or very deeply bilobed,
10-20 cm long, 1!4-2'4 cm wide, the outer base
of each more or less cordate (rarely strongly
auriculate), edges irregularly serrate and slightly
thickened; fertile secondary branches usually
sub-pinnate with flexuous axis bearing two
lateral and one terminal leaflets, sometimes twice
symmetrically dichotomous; fertile leaflets usually
deeply bilobed or geminate, one or both members
often having a shorter lobe commonly 9-15 cm
long and up to 13 mm wide including sorophores,
lamina reduced to a wing along the costa and
along each vein-group (sometimes two adjacent
vein-groups with a common lamina) the soro-
Phores thus in groups of 2—5; sorophores usually
2-3 mm long; spores coarsely and unevenly
verrucose, the warts often confluent.
Type: Milne 511, Solomon Isl. (K).
Distr. Melanesia (Solomon Isl. and New
Hebrides); in Malaysia: Admiralty Isl., Louisiades.
Ecol. ‘Climbing abundantly in brackish marsh”
(GRETHER & WAGNER 3997, Admiralty Islands).
Note. The specimen of GRETHER & WAGNER
has fertile branches more like those of L. dimor-
phum than is usual in L. trifurcatum, and has one
branch intermediate between sterile and fertile;
the fully sterile leaflets are smaller than normal
in L. dimorphum and only slightly cordate at the
base. The two species are very closely allied, and
it may be that in the Admiralty Islands, where
their areas of distribution overlap, there has been
hybridization.
8. Lygodium auriculatum (WILLD.) ALSTON, Rein-
wardtia 5 (1959) 16.—Ugena semihastata Cav.
Ic. Descr. Pl. 6 (1801) 74, t. 594, nomen. illegit.,
excl. syn. REICHARD & RuMPH.; C. B. Ros.
Philip. J. Sc. 6 (1911) Bot. 97; C. Cur. Dansk
Bot. Ark. 9, pt. 3 (1937) 29.—Hydroglossum
auriculatum WILLD. Sp. Pl. 5 (1810) 84.—L. semi-
hastatum Dresv. Mém. Soc. Linn. Paris 6 (1827)
203, nom. illeg.; Hoox. Syn. Fil. (1868) 437;
v. A. v. R. Mal. Ferns (1908) 111, excl. pl. bor-
neensis; Mal. Ferns Suppl. (1917) 115; C. Cur.
Ind. Fil. Suppl. I (1913) 119.—L. circinnatum
var. semihastatum Foss. Am. Fern J. 40 (1941)
142.—L. flexuosum [non (L.) Sw.] PRANTL,
Unters. Morph. Gefasskr. (1881) 73, p.p. quoad
syn. Cav. and WILLD.—Fig. 11.
Dec. 1959]
Schizaeaceae (Holttum) g 55
*s
oe
ay
Fig. 11. Lygodium auriculatum (WILLD.) ALSTON. a. Habit, x 24, b. hair, x 65 (a-b LE Roy TopPING
1287).
Rhizome short-creeping, bearing fronds close
together, its apex and bases of stipes densely
covered with dark hairs. Juvenile fronds once
dichotomous, each branch bearing a palmatisect
leaflet, usually 5-lobed, with truncate base, edges
closely and irregularly serrate. Rachis of climbing
fronds hardly 2 mm diameter, usually glabrous;
primary rachis-branches very short, dormant
apex covered with pale brown hairs having swollen
bases; secondary rachis-branches rarely bearing
a simple leaflet, most commonly once dichotomous,
one branch with a simple, one with a forked leaflet,
less often each branch with a simple leaflet; sterile
leaflets 12-20 cm long, 12-30 mm wide, subacute,
edges not thickened, very shallowly serrate, base
usually asymmetric and strongly cordate-auriculate
on the outer, rarely on both sides, costae glabrous
except near the base on upper surface; lamina of
fertile leaflets 12-20 mm (rarely to 30 mm) wide,
sorophores 3-9 mm long, constricted at the base,
at the apices of short triangular lobes of the lamina;
indusia glabrous; spores irregularly warty,
variable as between different specimens, in some
cases resembling those of L. /ongifolium, in others
with many smaller warts of variable size.
Type: Née, Luzon (Ma).
Distr. Indochina, Micronesia; in Malaysia:
E. Borneo, Philippines (Polillo, Luzon, Mindoro,
Samar. Mindanao), to 600 m altitude.
Note. The Micronesian specimens seen (from
Guam) are all smaller than Philippine ones, and
their secondary branches are regularly twice
dichotomous; they may constitute a distinct local
race (specimens of Née from Marianas not seen).
It may be that this species intergrades with L.
borneense in Borneo.
Vern. Nito, Tag.
Uses. Climbing rachises used for weaving,
making hats, and magic (contra-poison) bracelets.
9. Lygodium borneense v. A. v. R. Bull. Jard. Bot.
Btzg Il, m. 20 (1915) 29; Mal. Ferns Suppl.
(1917) 115; CopeL. Sarawak Mus. J. 2 (1917)
303; Hottr. J. R. As. Soc. Mal. Br. 6 (1928) 16
with fig.; Ferns Mal. (1955) 56.—L. semi-
hastatum sensu v. A. v. R. Mal. Ferns (1908) 111,
p.p. quoad pl. borneenses.—L. borneense if
samarindae v. A. v. R. Bull. Jard. Bot. Btzg II,
n. 20 (1915) 29; Mal. Ferns Suppl. (1917)
116.—Fig. 5b, 13c-e.
Rachis of scandent fronds glabrous, up to 2 mm
diameter; primary rachis-branches very short,
dormant apex covered with pale hairs having
swollen bases; secondary rachis-branches rarely
unbranched, normally once dichotomous (10-20
mm long below the dichotomy), each branch
56 FLORA MALESIANA [ser. Il, -voleat=
ALL Me aaa: ,
Z\ WTMIT EIS TT ry sO :
EVD wd
ey ww att
Oo ae
ei ay Pe * x) K ¢ rn)
SLL UTR
\ Drprea
Fig. 12. Lygodium longifolium (WILLD.) Sw. a. Habit, fertile, x 2/3, b. sterile, < 2/3, c. leaf detail showing
edge, x 3 (a KiNG's coll. 259, 6 MoTLEy s.n., c Hose 5034).
Dec. 1959]
Schizaeaceae (Holttum)
Fig. 13. Lygodium salicifolium PR. a. Habit, x 2/3, b. detail, « 10.—L. borneense v. A. v. R. c. Habit,
x #/,, d. detail of branching, x 21/,, e. ditto, x 4/3 (@ BERKHOUT s.n., b BRASS 5686, c SFN 18656, d-e
ENDERT 2057).
with a simple leaflet or rarely one leaflet double,
rarely the secondary branch-system sub-pinnate
with two lateral and one terminal leaflets; sterile
leaflets 20-35 cm long (on upper parts of frond
smaller), 314-5 cm wide, margin more or less
distinctly serrulate and slightly thickened, base
cuneate (rarely cordate on the outer base), surfaces
quite glabrous except the upper surface of the
costa towards its base, lamina not verrucose when
dry; fertile leaflets similar to sterile but slightly
smaller, 10-30 cm long, 12-40 mm wide (smallest
on upper branches), sorophores 4-10 mm long,
usually about 2 mm apart, at the apices of small
triangular projections of the lamina; spores
quite smooth.
Type: Teysmann, Borneo (Bo).
Distr. Malaysia: Malaya (SE. Johore only),
Sumatra (Mentawai Isl.), Borneo (many localities),
Talaud Isl.
Ecol. In light places in freshwater swamp-
forest; in Sarawak twice reported in the neigh-
bourhood of limestone hills but certainly occurring
also not in the vicinity of limestone; one specimen
from Sarawak, found growing with Jmperata on
sandy ground, has unusually small leaflets.
Note. This species has a branching habit closely
similar to that of ZL. auriculatum. L. borneense
usually differs from L. auriculatum in the larger
size and cuneate base of its leaflets, but some
Bornean specimens seem to be intermediate in
58 FLORA MALESIANA [ser. Il, volar
= er
x
ae
*
Vy
ey
Em
ZAI AY
q
i
DAIWA.
STII IIL SM
Fig. 14. Lygodium circinnatum (BurRM. f.) Sw. a. Part of sterile leaf, x 24, b-c. parts of fertile leaves,
showing different kinds of branching, x 24, d. veins of sterile leaf, x 3, e. ditto, leaf edge, x 9 (a,
d-e CLEMENS 9487, b ZOLLINGER 169, c Lady DALHOUSIE s.n.).
Dec. 1959]
the latter character. So far as observed, L. bor-
neense seems to be consistent in its smooth spores.
10. Lygodium longifolium (WILLD.) Sw. in Schra-
der, J. Bot. 1801, pt 2 (1803) 305; ALston &
Ho tr. Reinwardtia 5 (1959) 19.—Hydroglossum
longifolium WiLLp. Abh. Kurf. Mainz. Ak. Wiss.
Erfurt 2, pt 4 (1802) 22, t. 2.—L. digitatum PRESL,
Rel. Haenk. 1 (1825) 73 (?); v. A. v. R. Mal.
Ferns (1908) 112; Hoitr. Ferns Mal. (1955) 55.—
L. dichotomum (non (CAv.) Sw.) BEepDpb. Ferns S.
India (1863) t. 62.—L. teysmannii v. A. v. R. Bull.
Dép. Agr. Ind. Néerl. 18 (1908) 5; Mal.
Ferns (1908) 111, 801; Mal. Ferns Suppl.
(1917) 115.—L. circinnatum var. cristatum vy. A.
v. R. Bull. Dép. Agr. Ind. Néerl. 7. 18 (1908) 5;
Mal. Ferns (1908) 112, 802.—L. derivatum
v. A. v. R. Bull. Jard. Bot. Btzg III, 5 (1922) 213.—
Fig. 5a, 12.
Rhizome short-creeping, its apex and bases of
stipes covered with shining black hairs. Juvenile
fronds once or twice dichotomous, leaflets palma-
tely divided with 4~7 subequal lobes, base often
more or less cordate, lobes to about 18 cm long
and 18 mm wide, acuminate, edges shallowly
serrate, a vein ending in each tooth. Scandent
frond to about 4 m long, rachis to 2 mm diameter;
primary rachis-branches very short, with a dor-
mant apex covered with brown hairs having small
swollen bases; secondary rachis-branches 1-3
times dichotomous or sub-pinnate (lowest ones
sometimes unbranched and bearing large 6-lobed
leaflets); sterile leaflets composed of 2-4 subequal
lobes 15 cm or more long and c. 15 mm wide, the
sinuses between the lobes reaching to 15 mm
from the base of the leaflet, edges regularly ser-
rate, not or little thickened, base cuneate to
cordate, surfaces glabrous and usually not warty
when dried; fertile secondary branches \-3 times
dichotomous or (if the dichotomies are unequal)
more or less distinctly pinnate with two dicho-
tomous tertiary branches (tertiary branches may
rarely have three separate leaflets); fertile leaflets
simple or more usually consisting of two subequal
lobes united at the base, lamina 3-10 mm wide,
sorophores commonly 2—3 mm long, less often
to 6 mm; spores coarsely and irregularly verru-
cose.
Type: Herb. Willdenow, Malabar (B).
Distr. Southern India, Hainan; in Malaysia:
Malaya, Riouw and Lingga Islands, Sumatra,
Borneo, Luzon (doubtful).
Ecol. Edges of forest, probably in more ex-
posed places than L. circinnatum, and not at-
taining so large a size as that species.
11. Lygodium circinnatum (BurRM. f.) Sw. Syn.
Bil= (1806) 1533 Br. Eno Plt Jay. (1828) 253:
PRANTL, Unters. Morph. Gefasskr. 2 (1881) 64;
BEDD. Handb. (1883) 455; v. A. v. R. Mal.
Ferns (1908) 111; C. B. Ros. Philip. J. Sc. 6
(1911) Bot. 102; v. A. v. R. Philip. J. Sc. 11 (1916)
Bot. 116; Merr. Int. Rumph. (1917) 69; v. A. v.
R. Mal. Ferns Suppl. (1917) 115; W. H. Brown,
Bull. Bur. For. Philip. n. 19 (1919) t. 4; ibid.
Schizaeaceae (Holttum) 59
n. 22 (1920) 328, pl. IV; HEYNE, Nutt. Pl. (1927)
96; Back. Krakatoa (1929) 253; Burk. Dict. 2
(1935) 1378; BACKER & PostH. Varenfl. Java
(1939) 258, f. 67; HoLtr. Ferns Mal. (1955)55,
f.9; ALsTon & HoLtr. Reinwardtia 5 (1959) 20.
—Ophioglossum circinnatum Burm. f. Fl. Ind.
(1768) 228.—Ophioglossum pedatum Burm. f.
ibid. 227, t. 66 f. 1. —Ugena dichotoma Cav. Ic.
Descr. Pl. 6 (1801) 74, t. 594 f. 2; C. Cur. Dansk
Bot. Ark. 9, pt 3 (1937) 30.—L. pedatum (BuURM.
f.) Sw. Syn. Fil. (1806) 154; Merr. Philip. J. Sc.
19 (1921) 336.—Hydroglossum circinnatum (BURM.
f.) WiLtLp. Abh. Kurf. Mainz. Ak. Wiss. Erfurt
2, pt 4 (1802) 24.—Hydroglossum pedatum
(BurM. f.) WILLD. ibid. 25.—L. dichotomum
(Cav.) Sw. Syn. Fil. (1806) 154; Hook. & GREV.
omni 183i) Sd HOOK. ec BAK SYyt. bil
(1868) 437; Racts. Pterid. Buit. (1898) 8.—Ophio-
glossum furcatum Roxs. Calc. J. Nat. Hist. 4
(1844) 478.—L. basilanicum Curist, Philip. J. Sc.
2 (1907) Bot. 179; v. A. v. R. Mal. Ferns (1908)
802.—L. circinnatum var. monstruosum v. A. v. R.
Bull. Dép. Agr. Ind. Néerl. n. 18 (1908) 5; Mal.
Ferns (1908) 112; Mal. Ferns Suppl. (1917)
115.—Fig. 5d, 14.
Rhizome short-creeping, bearing stipes very
close together, its apex and bases of stipes densely
covered with black hairs. Juvenile fronds once
dichotomous, each branch bearing a _ pedato-
palmatisect leaflet, lobes usually 4 or 5, subequal,
to about 25 by 3!4 cm, the midrib of an outer lobe
arising near base of the next inner lobe, edges
entire, often somewhat crisped, pale and much
thickened (translucent when living), apices acute
to acuminate, surfaces glabrous but conspicuously
warty when dry (not when living), veins uniting
with the thickened margin. Rachis of climbing
frond to about 10 m long, 2-5 mm diameter,
glabrous; primary rachis-branches very short,
with sunken dormant apex covered with pale
hairs which are not thickened at the base; second-
ary rachis-branches unbranched and 2-6 cm
long, or once dichotomous with each branch
1-2 cm long beyond the fork; sterile leaflets
usually with 2-6 subequal diverging lobes which
are separate to within 2 cm from the base, entire,
margin pale and thickened, base cuneate or
truncate, surfaces nearly always warty when dry;
fertile secondary rachis-branches unbranched or
1—3 times dichotomous (rarely sub-pinnate); fertile
leaflets usually sessile in pairs at the ends of the
ultimate branches, or members of a pair partly
fused at the base, less often 3—5S-lobed (always so
if the secondary rachis is unbranched), lamina
more or less reduced and commonly 3-6 mm
wide, rarely less than 2 mm or approximating in
width to the sterile leaflet-lobes; sorophores 2-5
mm long, sessile; spores finely and evenly ver-
rucose.
Type: Java, herb. Burman (G, not seen).
Distr. Ceylon, NE. India to southern China,
Siam and Nicobar Isl. to Micronesia, the New
Hebrides and Solomons; throughout Malaysia.
Ecol. In lightly shaded places in primary or
secondary forest, in the lowlands and to 1500 m
60 FLORA MALESIANA [ser. IT, voleais
(Oe "58
Aha
—— <a
AMM AMIA SAIS A22
SV iA
RS ta
SNe:
a
Fig. 15. Lygodium merrillii Cope. a. Part of fertile leaf, x 14, b. part of sterile leaf, x 14, c. detail
of fertile leaf, « 114.—L. versteegii Curist. d. Part of fertile leaf, x 1/4, e. part of sterile leaf, x 4,
f. 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, b TopPING 1318, d—f LAM
1386).
Dec. 1959]
altitude, never where the ground becomes sea-
sonally very dry.
Vern. Ribu-ribu dudok, ribu-ribu bukit, paku
jari mérah, akar sidin, kapai bésar, réribu, M,
tura, Nias, kapai gorita, Mol., paku hata, hata
areuj, S, paku ata, paku ribu-ribu, Asahan, paku
rambat, J, ata, Bali, babar, Alor, masém, Minah.,
raga-raga, Mak., tjiwang, Bug., mongodo, Tob.,
gomoho, Tern., gomongo, Tidore, nito, Tag., Visc.
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. Mal. Ferns (1908) 803;
Mal. Ferns Suppl. (1917) 118.—L. matthewii
CopeL. Philip. J. Sc. 3 (1908) Bot. 36; ibid. 4
(1909) Bot. 20; v. A. v. R. Mal. 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 214-4 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 144 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: Merrill6057, Mt Halcon, Mindoro (Micw).
Distr. Tonkin and Kweichow; in Malaysia:
Sumatra, Sarawak, Philippines (southern Luzon,
Schizaeaceae (Holttum) 61
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,
re (19 11) 10%) 2 Corer, Philips. sco" (191)
Bot. 68; ? ibid. 11 (1916) Bot. 41; v. A. v. R.
Mal. Ferns Suppl. (1917) 118, 499; ALSTON &
Ho.tr. Reinwardtia 5 (1959) 22.—L. mosz-
kowskii BRAUSE, Bot. Jahrb. 49 (1912) 57; v. A.
v. R. Mal. Ferns Suppl. (1917) 116.—Fig. 15d-f.
Rhizome and juvenile fronds not seen. Rachis
of scandent frond to 2'4 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, 2-214 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 witha 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, differing: venation reticulate, lamina
firmer, the leaflets usually more elongate.
Distr. Queensland, Fiji to Tahiti, New Hebri-
des, New Caledonia; reported by KUHN (/.c.) from
New Ireland but without citation of a specimen.
not otherwise known in Malaysia.
Excluded
Cheilanthes fuscata BL. En. Pl. 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
Fig. 1. Isoetes habbemensis ALSTON, tufted in marginal shallows of Habbema Lake, c. 3225 m, with
Libocedrus papuana F. y. M., on ridge in background (Archbold Expeditions, Brass, 1938).
ISOETACEAE (A. H. G. Alston +, London)
1. ISOETES
LINNE, Sp. Pl. (1753) 1100; Gen. Pl. 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—-4-
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 divided 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 with 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, 20-45 « 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
(DutuHiz, 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: Jsoetes (worldwide), Stylites (Peru), and Nathorstiana (fossil
64 FLORA MALESIANA [ser. II, vol. 11, 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 Jsoetes and
the Triassic fossil Pleuromeia. Isoetes has a very short stem of complex structure, and is generally regarded
as reduced and specialized. The fossil genus /soetites, differing from Jsoetes 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 /soetes, and estimates of the number of
species vary. 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;
I. lacustris L., n = 54-56 (Problems of Cytology and Evolution in the Pteridophyta, 1950, 254-259).
RAUH & FALK (i/.c.) record 2n = c. 50 for Stylites gemmifera W. RAUH.
Uses. Leaves of J. philippinensis are said to be eaten.
KEY 10° THE SPECIES
1. Megaspores smooth on inner surfaces.
2. Leaves up to 50 cm. Stock 3-4-lobed. Microspores minutely scabrous 1. I. philippinensis
2. Leaves up to 14 cm. Stock 2-lobed. Microspores densely spinulose 2. I. habbemensis
1. Megaspores warted on inner surfaces. Stock 3-4-lobed. Leaves up to 7,5 cm 3. I. neoguineensis
1. Isoetes philippinensis MERRILL & PERRY, Am.
Fern Journ. 30 (1940) 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. 7 mm broad at base, with membranaceous
wings. Accessory peripheral strands wanting.
Foveae 3-4 cm long, narrow, 114-2 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 uw 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 mw, very minutely
scabrous.
Distr. Malaysia: Philippines (Mindanao: Lanao
Prov. near Momungan, vicinity of Olangu).
Ecol. Bottom of a stream, 400-500 m, once
found.
Vern. Kabauingbauing, Lanao.
2. Isoetes habbemensis Atston, J. Arn. Arb. 26
(1945) 180.—Fig. 1.
Submerged aquatic. Stock apparently 2-lobed,
appressed-semiglobose c. 314 by 114 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 yw in diam., almost smooth,
with a prominent triradiate marking, pale greyish-
white when dry. Microspores c. 43 uw 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. y. 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. Gard. 9(1922) 211.
Submerged aquatic. Stock 3—4-lobed. Leaves
numerous, 5-714 cm by 3 mm, recurved, terete
towards the apex, flattened lower down, abruptly
dilated at base. Dilated base hyaline, c. 1 cm long
and broad. Upper part of leaves dark green.
Stomata few. Ligule broadly cordate. Velum none.
Sporangia oblong, 6 by 4 mm, pale brown.
Megaspores c. 800 w in diam., deeply and ir-
regularly warted and reticulate on the outer
face, warted on the inner faces, with a strongly
marked triradiate marking.
Distr. Malaysia: E. New Guinea (Mts Scratch-
ley and Albert Edward), twice found.
Ecol. Shallows of an alpine lake, 3000-3680 m.
Note. BAKER’S statement that the megaspores
are smooth between the triradiate ridges is
incorrect.
CYATHEACEAE (R. E. Holttum, Kew)
Caudex massive, usually erect and unbranched, where prostrate not dorsiventral in
structure; fronds arranged on caudex in spiral series; vascular system of caudex
a hollow cylinder with gaps corresponding with leaf-bases, in some cases small
medullary bundles also present; a cylinder of very hard sclerenchyma, with gaps
at leaf-bases, present both inside and outside the vascular cylinder (but absent in
Cibotium), the surfaces of the sclerenchyma covered with cubical cells containing
silica; tangentially arranged sieve-tubes present in the phloem as well as longitudi-
nal ones. Stipes of Cyathea containing numerous small vascular strands arranged
in 3 series (fig. 6), these strands more or less united in smaller axes of Cyathea-
fronds and also in larger axes of other genera (fig. 31f, 33d); stipe-bases per-
sistent, or sooner or later caducous leaving a pattern of scars on the caudex;
pneumathodes present along each side of stipe, in a single discontinuous or almost
continuous row, or in 2—3 rows close together, the row joining upwards to a similar
row on the basiscopic side of the first pinna, a + circular pneumathode at the
base of the pinna beginning the row on the main rachis to the next pinna. Dermal
appendages on fronds: multiseptate hairs only, or both hairs and scales (Cyathea);
if both, the hairs often confined to the adaxial surface of the fronds. Fronds in
most cases bipinnate-tripinnatifid, with varying gradations to tripinnate, in a few
cases simply pinnate, in Cu/cita 3-4-pinnate; pinnules almost symmetrical at the
base except in Culcita; veins normally free except in Cyathea capitata and in the
genus Cnemidaria (trop. America). Sori either terminal on veins and protected
by an inner indusium as well as by the more or less reflexed edges of a small lobe
of the lamina (outer indusium), or apparently not terminal on veins and not near
the edge of the lamina, with indusia of various form or without indusia; receptacle
of various shape, in all cases containing vascular tissue which in the case of
Cyathea represents the termination of a short vein; stalks of sporangia short or
long, 4 or more cells in transverse section, annulus more or less oblique, with a
more or less clearly defined lateral stomium; spores trilete, surfaces smooth or
variously sculptured; multiseptate paraphyses, of a single row of cells (terminal
cell glandular or not) or scale-like at the base, present with sporangia.
Distribution. Throughout the wetter parts of the tropics, especially on mountains; a few species
just north of the tropics, more south of the tropics especially in Australasia. As here construed, 9 genera,
of which 5 are Malaysian: Cyathea (pantropic, at least 600 spp.); Cnemidaria (limited to species with
simply pinnate fronds, anastomosing veins and distinctive spores, tropical America, 10 spp.); Lophosoria
(tropical America, monotypic); Dicksonia (tropics and southern subtropics in Malaysia, Australasia,
America, St Helena, c. 25 spp.); Cystodium (Malaysia, monotypic); Thyrsopteris (Juan Fernandez,
monotypic); Culcita (subg. Culcita in Azores and tropical America; subg. Calochlaena in Malaysia and
Australasia; in all c. 7 spp.); Cibotium (SE. Asia, Malaysia, Hawaii, Central America, c. 12 spp.); Metaxya
(tropical S. America, monotypic).
Fossils. SEwARD gave a summary of knowledge to 1920 (Fossil Plants 2, 365-375). T. M. HARRIS
has recently published a fully illustrated account of some Jurassic frond-fossils which he includes in the
family Dicksoniaceae (The Yorkshire Jurassic Flora, 1, 1961, 140-181), referring them to the genera
Dicksonia, Coniopteris and Kylikipteris. Owing to the fragmentary nature of the fossils it is very difficult
to judge how they compare with existing ferns. In my judgement, the fossil most like living Dicksonia is
Coniopteris hymenophylloides (BRONGN.) SEWARD; C. murrayana BRONGN. is perhaps more like Culcita.
The Jurassic fossils most resembling Cyathea in form of sterile leaflets are placed in the genus Kylikip-
teris; their fertile leaflets have sori at the ends of veins on a reduced lamina, and seem more like those of
Thyrsopteris than Dicksonia. Kylikipteris looks like a possible Cyathea-ancestor. Though no fossils with
Cyathea-like sori have been found in the Yorkshire Jurassic, HARRIS described a genus Aspidistes which
has sori and sporangia resembling Dryopteris or Thelypteris, spores trilete (known in a few species of
(65)
[ser. II, vol. 1?
, Papua, 2840 m (L. J. Brass, 1933).
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FLORA MALESIANA
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athea gleichen
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Dec. 1963] iy ¥ CYATHEACEAE (Holttum) ne : 67
Fig. 2. Cyathea gleichenioides C. CHR., same locality as fig. 1. Trunks have been blackened by a recent
grass fire (L. J. Brass).
Thelypteris but not in Dryopteris) and spherical unicellular glands (common in Thelypteris). Aspidistes
looks like an early Thelypteris, and Thelypteris has several features in common with Cyathea; but one
would have expected Cyathea to have existed prior to Thelypteris. Fossil tree-fern trunks of Lower
Cretaceous have also been called Coniopteris; they have leaf-scars and vascular system comparable with
those of Cyathea and Dicksonia. OGURA described fossil tree-ferns from Upper Jurassic and Cretaceous
rocks of Japan and Korea (J. Fac. Sc. Univ. Tokyo III, 1, 1927, 351-380, pl. 2-8). BANCROFT, describing
a fossil Cyatheoid stem from the late Tertiary of East Africa (New Phytol. 31, 1932, 241-253) pointed
out that OGuRA’s fossils differed in some respects from existing Cyatheaceae. K. JAcos described impres-
sions of parts of a tree-fern stem from middle Jurassic of NE. India, but the pattern of vascular strands
in the leaf-scars is not clearly preserved. He gives references to other descriptions of fossils of presumed
Cyatheaceous affinity (Proc. Ind. Ac. Sc. 6, sect. B, 1938, 73-90).
Ecology. Most species are forest plants, with varying degrees of tolerance of exposure to direct
sunlight and to drying wind. In Western Malaysia Cyathea moluccana, C. squamulata and C. glabra
occur only in quite shady forest and will not tolerate exposure; C. /atebrosa is most vigorous where it
has more light and will tolerate almost full exposure of its crown to the sun; C. contaminans is only
vigorous where its crown is fully exposed, though its roots need shade. It is probably significant that
C. contaminans, flourishing in clearings in the forest, is more widely distributed than any other Ma-
laysian species. On high mountains in New Guinea a few species (notably C. macgregorii) can tolerate
the full exposure of open grassland and will tolerate periodic burning of the grass (fig. 1, 2).
Vegetative morphology. The majority of species are arborescent, and the habit of growth very
similar in all genera. The height to which the trunk will grow varies from species to species; full records
of this are not available. The lower part of the trunk has many adventitious roots, which become entangled
and form a rigid covering of increasing thickness, supporting the base of the trunk; the cover of roots at
the base of an old tree-fern is many times thicker than the original trunk. Some species produce branches,
either near the base of the trunk or higher up; in the former case there will be a small cluster of trunks,
in the latter the main trunk will have lateral crowns of small leaves upon it (fig. 3-4).
SCHOUTE reported an exceptional case in which 33 such lateral branches on one plant formed many
roots which coalesced with those of the main trunk, each lateral branch growing upwards and forming
a separate trunk; the result was a cluster of trunks all growing out from one great mass of roots, and the
nature of the branching could only be seen by removing the roots (Ann. Jard. Bot. Btzg 20, 1906, 198-207).
ScHoutE also published a detailed study of four cases in which trunks of tree-ferns appeared to branch
by bifurcation, remarking that this condition is rare and may be due to injury (Rec. Trav. Bot. Néerl.
11, 1914, 95-192, t. 5-21). He noted that at each bifurcation is an ‘Angularblatt’, as in ferns with dicho-
tomously branched rhizomes, and he suggested that perhaps there is no sharp distinction between such
bifurcation and lateral branching in which the branch occurs on one side of a leaf-base.
Some species of Cibotium and Culcita, and also Cystodium sorbifolium, have prostrate stems with
68 FLORA MALESIANA
Ch aS
Fig. 3. Cyathea contaminans (WALL.) CopEL. with branching of upper part of trunk. Mt Telemojo,
Central Java (P. ARENS). —Fig. 4. The branched trunk shown in fig. 3 with fronds removed.
indefinite apical growth; these prostrate stems are massive, with leaves close together, never long-creeping
which is a distinction from Dennstaedtia, a genus united to Dicksonia by Hooker. The indefinite horizon-
tal growth of these stems contrasts with the vertical growth of the arborescent species, which sooner
or later outgrow their mechanical strength if they do not earlier succumb to other injuries. Cyathea
biformis has a very slender trunk which supports itself by adventitious roots clinging to a tree-trunk;
its fronds are more widely spaced than in arborescent species.
The diameter of the trunk varies considerably, from the very slender C. biformis just mentioned to
the massive C. contaminans and allied species. The fronds usually break near the base when they are
old, their bases persisting for a longer or shorter period; in many cases they are ultimately shed, leaving
distinctive scars on the trunk (the scars sometimes later covered by roots). The shape and arrangement of
the scars depends on various factors, one being rate of growth of the trunk. A fast-growing trunk will
have rather widely spaced frond-scars which are vertically elongated; a slow-growing trunk will have
frond-scars closely placed and almost circular (fig. 6). Frond-scars seem always to be in vertical orthosti-
chies, and in several spiral parastichies. ScHourTe (/.c.) described the arrangement of leaf-scars on the
branching trunks he examined, and especially the way in which the pattern on the branches is related
to that on the parent trunk. In the uniform conditions of the forest of Western Malaysia, new fronds ap-
pear singly, but probably they are more abundant following wetter periods. At Tjibodas, West Java,
JAAG made observations on seven young plants of Cyathea contaminans (Alsophila glauca) and found
that the average time between development of successive fronds varied from 25 to 28 days, and the
life of a single frond from 165 to 200 days; the number of fronds on a single plant varied from 6 to 10,
and the time taken for a complete renewal of the whole crown of fronds from 182 to 243 days. An old
plant with trunk 10 m tall bore about 12 fronds and the mean time between unfolding of new fronds was
21 days (Mitteil. Naturf. Ges. Schaffhausen 12, 1943, 211-217). It is remarkable that fronds thus ap-
Dec. 1963] | o3 CYATHEACEAE (Holttum) iy 69
pearing singly leave ultimately the scars of their bases in regular alternate whorls. In New Guinea, Hooc-
LAND has observed that some species produce their fronds in whorls, those of one whorl being simul-
taneous. More observations on phyllotaxis and on rate of growth are needed. There is also the con-
sideration that plants of the same species growing under more or less favourable conditions may vary
considerably in the size of trunk and of fronds. Few Malaysian species have such massive trunks as the
Australian C. australis (R. Br.) Domin and Dicksonia antarctica, which both have fronds in many
orthostichies. JAAG observed three plants of Dicksonia blumei at Tjibodas. Each crown usually consisted
of 16-18 fronds, each new frond appearing at an average interval of 22—27 days, the life of a single frond
being 185-191 days.
The pneumathodes which occur along each side of the stipe and rachis often afford distinctive char-
acters in Cyathea (fig. 7) but are usually not seen in herbarium specimens, which shrink along the line
of thin-walled tissue. In young fronds the pneumathode has a continuous epidermis containing stomata;
later the epidermis ruptures and the cells of the underlying tissue become more or less separated from
each other, sometimes having peg-like outgrowths.
In shape of frond, shape of leaflets and external form of rachis-branches, Cu/cita and Thyrsopteris
differ from the other genera. These differences are summarized as follows. Frond-form: in Culcita and
Thyrsopteris broadly deltoid, 3—4-pinnate, stipe always long; in other genera elliptical, usually bipinnate-
tripinnatifid, lower pinnae always somewhat reduced, sometimes much so and then the stipe very short.
Shape of leaflets: in Culcita and Thyrsopteris very asymmetric at base (broad on acroscopic side), with a
gradual reduction from largest to smallest; in other genera nearly symmetrical at base, usually with
many leaflets (pinnules) on each pinna of approximately equal size. External form of rachis-branches:
in Culcita and Thyrsopteris upper surface grooved (fig. 34c), the groove open to admit grooves of smaller
branches and of midribs of leaflets (which are similarly grooved), edge of lamina separately decurrent
on side of rachis-branch; in other genera upper surfaces raised (or at most slightly grooved), midribs of
leaflets also raised (fig. 18b). It may be noted that in ali these characters there is more or less complete
agreement between Culcita and the Dryopteris-Athyrium group of genera, and between the other genera
and the Thelypteroid ferns.
Dermal appendages. In Dicksonia, Cystodium, Culcita and Cibotium the dermal appendages are
all simple septate hairs, the longest often quite thick at the base; the characters of the hairs (rigid or
flaccid, long or short, varied colour) especially on the stipe, are always important diagnostically. In
Cyathea there are always septate hairs (sometimes branched near the base) on the upper (adaxial) surface
of stipe and rachises, these hairs rather crisped and antrorse; in most species there are no hairs on the
lower surfaces. All species of Cyathea have scales on the lower surfaces, in size decreasing from those on
the base of the stipe to those on costules of pinnule-lobes. The genus may be divided into two subgenera,
subg. Cyathea having flabelloid, subg. Sphaeropteris setiferous scales (fig. 8). Where hairs occur on lower
surfaces in subg. Sphaeropteris they are rather thick and straight, much as in Dicksonia; in subg. Cyathea
they are crisped and more or less appressed. For further notes on scales, see Cyathea.
Sori. In Dicksonia, Cystodium, Culcita and Cibotium the sorus is at the end of a vein (or of the acros-
copic branch of a vein) near the margin; it is protected by a small reflexed marginal lobe (the outer in-
dusium) and by an inner indusium which shows varying degrees of difference from the outer indusium.
The inner indusium is more or less fused to the side of the receptacle remote from the margin (fig. 31b, c).
The surface of the receptacle is not very prominent, but spreads at right angles to the end of the vein.
In Cyathea the sorus is usually seated at the fork of a vein, well away from the margin; or where the
veins are not branched, it is apparently in the middle of a vein. There is always vascular tissue in the
receptacle, and this is the end of a short branch-vein. The receptacle is prominent, more or less spherical
or clubshaped. The indusium is of very varied form, with also varying degrees of reduction, and is
sometimes lacking. The Cyathea sorus may be compared to Dicksonia by considering the form called
Hemitelia (fig. 9c), which has an indusium attached to the base of the receptacle on the side remote from
the margin (there is never an indusium attached only on the marginal side). The beginning of the de-
velopment of such a sorus and of a sorus of Dicksonia are identical; the receptacle appears to be on
the true leaf-margin, with an outgrowth on upper and lower sides. In Dicksonia the two outgrowths
develop almost equally, in Cyathea very unequally so that the sorus is ultimately far from the margin, and
new veins are needed to supply the additional marginal area of leaf-lamina. The derivation of the other
types of Cyathea sorus from this one is described under Cyathea.
In Dennstaedtia, formerly included in Dicksonia, the receptacle is more or less columnar and free,
at the end of the vein, and is surrounded by almost completely fused outer and inner indusia; it thus
differs from both Dicksonia and Cyathea.
Sporangia and spores. Sporangia are not very large, having in most cases 64 spores. The annulus
is in all cases more or less oblique and usually indurated at the base where it passes the stalk. For details,
see Bower, The Ferns 2 (1926) 266, 282, 301 and fig. 5. The stalk is in all cases rather massive, consist-
ing of 4-7 rows of cells. In genera with the Dicksonia type of sorus, where the receptacle is not prominent,
sporangia have rather long stalks; in Cyathea and Thyrsopteris, where the receptacle is prominent, spo-
rangia-stalks are short. Spores are in all cases trilete, and the sculpturing of the surface varies consid-
erably, though it is always slight in Cyathea. The spores of Cnemidaria (confined to C. horrida and re-
70 FLORA MALESIANA [ser.; Il, vole
lated species with simply pinnate fronds and anastomosing veins) have an almost spherical cavity in the
thickened lateral walls (alternating with the trilete ridges). This tropical American group have the most
distinctive spores in the family, and should rank as a separate genus.
ol
Fig. 5. Cyathea capensis (L. f.) SM. a. Sporangium, outer face showing complete annulus and short stalk,
b. same, lateral view to show stomium.—C. brownii DoMIN (Alsophila excelsa R. BR.). c. Sporangium,
inner face, d. same, lateral view, e. transverse section of stalk. All x 100 (after F. O. Bower, The Ferns,
2, fig. 563). —Fig. 6. Cyathea contaminans (WALL.) CopPEL. Trunk showing scars after abscission of stipes.
Mt Bukit Tungul, W. Java (L. VAN DER PUL).
Gametophyte. STOKEY gave a comparative account of gametophytes of all genera except Metaxya
and Cystodium (Bot. Gaz. 90, 1930, 1-45). Mature prothalli are longer, with more massive cushion, and
with greater tendency to fork, than prothalli of the majority of more specialized ferns. Multicellular hairs
of peculiar origin occur abundantly in Cyathea, rarely and late in Lophosoria, not in the other genera.
These hairs are in origin like those only of Gleichenia and Loxsoma (see STOKEY & ATKINSON, Phyto-
morphology 6, 1956, 260); they are longer in Cyathea than in Gleichenia and lack terminal glandular
cells. Antheridia in all cases are relatively primitive, with wall of 5 cells; those of Thyrsopteris are largest
and least symmetrical. Archegonia have rather long necks, longer in the Dicksonia group of genera than
in Cyathea.
Cytology. Chromosome numbers in the genera Dicksonia, Culcita and Cibotium have been recorded
by MANTON (J. Linn. Soc. Bot. 56, 1958, 84) and in Cyathea by MANTON & SLEDGE (Phil. Trans. R. Soc.
B, 238, 1953, 137) and by MANTON (Appendix to HoL_trum, Rey. Fl. Malaya 2, 1954, 623); Prof.
MANTON also permits me to report unpublished observations on Culcita and Dr T. G. WALKER on
Cnemidaria. No observations are yet available for Thyrsopteris, Lophosoria, Metaxya and Cystodium.
The numbers are: Dicksonia, n = 65 (3 spp.); Cibotium, n = 68 (2 spp.); Cyathea, n = 69 (several spp.);
Cnemidaria horrida, n = 69; Culcita macrocarpa, n = 66 approx.; Culcita dubia, n = 58.
Anatomy. The first critical account of anatomy in this family was by METTENTIUs, in the course of
his study of Angiopteris (Abh. M.—Ph. K1. K. Sachs. Ges. Wiss. 6, 1863, 525-531, t. V.) The most recent
full account of the anatomy of members of Cyatheaceae is by OGURA (J. Fac. Sc. Imp. Univ. Tokyo,
Bot. 1, 1927, 141-350).
U. SEN has recently completed a new anatomical study of the family, summarized in HoOLTTUM & SEN
Dec. 1963] ‘ved CY ATHEACEAE (Holttum ) a 2 71
(Phytomorphology 11, 1961, 406-420). The vascular structure, with its accompanying sclerotic tissue,
is very similar in all Malaysian genera except Cibotium; it is most fully developed in Cyathea, to which
the following notes apply. As seen in a transverse section of the trunk, there are several meristeles, with
gaps between them, together forming a hollow cylinder, the gaps corresponding to leaf-bases. On the
outer and inner sides of each meristele are plates of very hard sclerotic tissue. Small vascular strands
arise from the margins of the gaps and supply the leaves, and in Cyathea there are also small medullary
bundles which anastomose with each other and with the meristeles. Distinctive ‘cubical cells’ form a
more or less continuous layer surrounding each mass of sclerenchyma; their walls adjacent to each other
and to the sclerenchyma are much thickened, and they contain crystals which appear to be silica. The
sclerotic tissue, with its cubical cells, is lacking in Cibotium. In the phloem are tangentially elongated
cells, in structure like the longitudinal sieve-tubes; such cells are only otherwise known to occur in Os-
munda. The pattern of arrangement of the numerous vascular strands in the stipe of Cyathea is distinc-
tive; in other genera they are more or less joined. In the smaller axes of the frond the pattern is progres-
sively simplified. The stomata of Cibotium show more complex developmental stages than those of the
other genera.
Economic importance. OcCHSE & BAKHUIZEN VAN DEN BRINK reported the use of coiled young
fronds of Cyathea contaminans and C. junghuhniana (mis-named C. latebrosa) as food, also the pith of
young parts of the trunk of the former species (Vegetables D. E. I., 1931, 212-215). Other species (perhaps
all) are similarly edible; HOOGLAND notes this of some from the mountains of New Guinea. The pith of
trunks was formerly eaten by Maoris in New Zealand. The common name in Java and Sumatra for the
larger tree-ferns, Pakis (or Paku) tiang (tiyang, teehang), indicates the use of the trunks as posts; this name
does not seem to have been noted in the Malay Peninsula. The sclerenchyma of most tree-fern trunks is
exceedingly hard and durable, and provides nearly all the mechanical strength when they are used as
posts. It also provides an interesting pattern when cut in different ways, and this effect is used in the
construction of ornamental objects in various parts of the world. In North Borneo I noted old tree-fern
trunks, hollowed out, in use as bee-hives around Dusun houses. On Mt Patuha, W. Java, hollowed tree-
fern trunks are filled with carbide gas for making booms on New Year’s eve. The masses of adventitious
roots at the bases of Cyathea trunks are used in orchid culture, either as solid slabs (cut with a saw) or
broken, in potting mixtures.
Taxonomy. BERNHARDI (in Schrader, Neues J. Bot. 1, ii, 1806, 1-204) attempted a classification of
ferns according to the form and position of the annulus of a sporangium, proposing a division into
Helicogyratae (including Cyathea and Dicksonia), Cathetogyratae (majority of leptosporangiate ferns),
Pseudogyratae (including Gleichenia) and Agyratae. PRESL (Tentamen, 1836) varied this by associating
Gleichenia and Cyathea (sens. lat.)in Helicogyratae and placing Dicksonia (under the name Balantium)
in Cathetogyratae. HOOKER (Sp. Fil., 1844 and Syn. Fil., 1868) arranged all ferns in seven suborders, all
genera here treated being included in suborder Polypodiaceae; they are divided as tribe Cyatheae (Cyathea,
s.l.) and tribe Dicksoniae (all other genera, also some additional ones). METTENIUS (Fil. Hort. Bot.
Lips., 1856) arranged all ferns in eight orders, of which the second was Cyatheaceae, which corresponded
exactly with the present arrangement with the addition of Matonia (METTENIUS used the name Balantium
in place of Dicksonia). Curist (Farnkr. d. Erde, 1897), Diets (in Engl. & Prantl, Pflanzenfam. 1, Abt. 4,
1899, 113-139) and CHRISTENSEN (Ind. Fil., 1905) adopted a family Cyatheaceae with the same content
as the order Cyatheaceae of METTENIUS, with omission of Matonia. Bower, however, believed that
Cyathea and its near allies should be associated closely with Gleicheniaceae, as one of the more primitive
elements of the series Superficiales, while he placed Dicksonia and allies in the series Marginales, regarding
the separation of the two as ‘long overdue’ (The Ferns 2, 1926, 326). This idea was followed by CuHrIs-
TENSEN in 1938 (in Verdoorn, Man. Pterid., 532, 533), where he recognized two families, Dicksoniaceae
and Cyatheaceae. BowER’s arrangement involves the assumption that primitive Cyathea, like Gleichenia,
was exindusiate, so that indusia in his Cyatheaceae are a new development, not homologous with the
inner indusium of Dicksonia (l.c. 304). In this he disagreed with GogBeL (Flora 105, 1913, 45), who
regarded the indusium of Hemitelia (now included in Cyathea) as strictly homologous with the inner
indusium of Dicksonia. COPELAND (Gen. Fil., 1947) included Dicksonia and allies in a family Pteridaceae,
associating with them Lindsaea, Dennstaedtia, etc., while maintaining Cyathea s./. in a separate family
Cyatheaceae. HOLTTUM & SEN have published a discussion of the whole question (Phytomorphology 11,
1961, 406-420), with the conclusion that GoEBEL’s contention was correct; they give a new subdivision
of the family Cyatheaceae, as here constituted, based partly on new evidence. This subdivision is sum-
marized as follows.
CONSPECTUS OF THE FAMILY
Subfamily Cyatheoideae.
Fronds normally bipinnate with lower pinnae more or less reduced; pinnules almost symmetrical; upper
surfaces of costae and pinna-rachis raised (or, if grooved, the groove of a major axis not open to admit
that of a minor one borne upon it); sori terminal on veins or on lower surface of veins, indusiate or not;
dermal appendages hairs or scales or both; cubical cells present in association with sclerenchyma;
stomata with single subsidiary cell.
UZ FLORA MALESIANA [ser: I voliate
Tribe Cyatheae.
Scales and hairs present as dermal appendages; sori superficial, indusiate or not; cubical cells in
continuous layer on surfaces of sclerenchyma.
Fronds mostly bipinnate; veins almost always free; spores with thin walls of uniform thickness,
smooth or papillose; indusium various or lacking . aber . . 1. Cyathea
Fronds simply pinnate with anastomosing veins; spores with ‘wall much thickened, a spherical
hollow in the middle of each face; indusium hemitelioid (c. 10 spp., tropical America) (Cnemidaria)
Tribe Lophosorieae.
Hairs only as dermal appendages; sori superficial, no indusia; cubical cells singly in association with
sclerenchyma’ (lsp;.. tropical@Amenica)) Go °05 F750 SPR ie tee ae gant en tance ra ea
Tribe Dicksonieae.
Hairs only as dermal appendages; sori marginal, protected by slightly modified marginal lobe of
lamina (outer indusium) and a thinner inner indusium; receptacle of sorus fused to inner indusium.
Fronds bipinnate with deeply lobed ah a or ars stem usually a thick erect trunk; cubical
cellsasin Cyatheae .. . : . . . 2. Dicksonia
Fronds bipinnate with simple pinnules; stem prostrate; veondition of enbical cells not known.
3. Cystodium
Subfamily Thyrsopteridoideae.
Fronds 3—4-pinnate, lowest pinnae largest; leaflets asymmetric; upper surface of axes and of leaflet-
midribs grooved, grooves of major axes open to admit those of minor ones; sori at ends of veins; cubical
cells present; stomata with single subsidiary cell.
Tribe Thyrsopterideae.
Fertile and sterile parts of frond strongly dimophous (lamina much reduced in fertile part); receptacle
of sorus columnar with sporangia all round it, indusium ultimately a shallow uniform cup; stem
massive, erect; cubical cells scattered (1 sp., Juan Fernandez Is). . . .. . . . . (Thyrsopteris)
Tribe Culciteae.
Fertile and sterile parts of frond not greatly dimorphous; receptacle of sorus fused to inner indusium
(as in Dicksonia); inner indusium thinner than outer, the two slightly joined together at the base;
stem prostrate or erect; cubical cells in a continuous layer as in Dicksonia. . .. . . . 5. Culcita
Subfamily Cibotioideae.
Fronds normally bipinnate; pinnules almost symmetrical; upper surfaces of pinna-rachises and costae
raised; sori terminal on veins, shape much as in Dicksonia but with outer and inner indusia both unlike
the lamina of the frond, lacking chlorophyll and lacking intercellular spaces; sclerenchyma and cubical
celisilackinis:stomatal withi3isubsidiary, cells 34. 5+. =. -aem QueIey nc) ee suse Or DO tina
Subfamily Metaxyoideae.
Fronds simply pinnate, pinnae lobed on young plants only; upper surface of rachis and of midribs of
pinnae grooved, groove of rachis open to admit grooves of pinna-midribs; sori superficial on lower
surface of veins, usually more than one to a vein, no indusium; sclerenchyma and cubical cells lacking;
stomata with 3)subsidiary. cells) (1 sp:, tropical America)... «3% >...» « «.- + = = (Metaxga))
ARTIFICIAL KEY TO MALAYSIAN GENERA
1. Upper surface of costae of ultimate leaflets raised; fronds + elliptical, mostly bipinnate.
2 Youngspartsiot plant protected’ by scales‘and hairs}. .-. 2. .)- - = 1): o)e 9) eeeele@yatnes
2. Young parts of plant protected by hairs only.
3. Outer indusium not distinct from rest of lamina.
4. Pinna-rachis raised on upper surface; fertile pinnules deeply lobed. . . . . . . 2. Dicksonia
ae Pinna-rachis grooved on upper surface; fertile pinnules not lobed. . . . . . . 3. Cystodium
Outer indusium quite distinct from lamina of leaflet. . . . ;-.-. 4. @ibotum
ile are surface of costae of ultimate leaflets grooved, the groove decurrent into the groove of the
supporting rachis; fronds deltoid in outline, 3-4-pinnate ..........~... . #S. €uleita
1.CYATHEA
SMITH, Mem. Ac. Turin 5 (1793) 416; Swartz, Syn. Fil. (1806) 139, 364; KAULF.
En. Fil. Chamisso (1824) 254; Pres_, Tent. Pterid. (1836) 54; Hook. Gen. Fil.
(1839) t. 23; Sp. Fil. 1 (1844) 14; Syn. Fil. (1865) 16; J. Smiru, Lond. J. Bot. 1
(1842) 659-668; Hist. Fil. (1875) 244; Curist, Farnkr. Erde (1897) 10, 317;
DIELS in E. & P. Pfl. Fam. 1, 4 (1899) 123; CopEL. Philip. J. Sc. 3 (1909) Bot. 353;
Gen. Fil. (1947) 95.—Sphaeropteris BERNH. in Schrader, J. Bot. 1800, ii (1801)
Dec. 1963] CYATHEACEAE (Holttum ) 73
Fig. 7. On left Cyathea orientalis (KUNZE) Moore, young frond showing widely-spaced short pneuma-
thodes. On right C. incisoserrata CopEL., bases of stipes showing almost continuous and often double
rows of pneumathodes. Cult. R. B. G. Kew (R. VAN CREVEL, 1961).
122.—Hemitelia R. Br. Prod. (1810) 158, p.p. (excl. H. horrida).—Alsophila R.
Br. /.c.—Chnoophora KAULF. En. Fil. Chamisso (1824) 250.—Gymnosphaera BL.
En. Pl. Jav. (1828) 242; CopeL. Gen. Fil. (1947) 98.—Disphenia PResL, Tent.
Pterid. (1836) 55.—Schizocaena J. SM. in Hook. Gen. Fil (1838) t. 2; COPEL.
Gen. Fil. (1947) 99.—Amphicosmia GARDNER, Lond. J. Bot. 1 (1842) 441.—
Dichorexia PrReEsL, Abh. K. B6hm. Ges. Wiss. V, 5 (1848) 55.—Fourniera BOMMER,
Bull. Soc. Bot. France 20 (1873) xix.—Eatoniopsis BoMMER, /.c.—Thysanobotrya
v. A. v. R. Bull. Jard. Bot. Btzg II, n. 28 (1918) 66.—Fig. 1-30.
Trunk always erect, short or tall. Scales present on lower (abaxial) and lateral
surfaces of axes of frond, at least in early stages of growth, larger ones on stipe and
rachis sometimes borne on spine-like outgrowths; fairs always on upper surfaces
of all but smallest axes, antrorse, sometimes branched; hairs of various kinds some-
times on lower surfaces of axes and veins, rarely on lamina between veins. Pneu-
mathodes present in a discontinuous line (or 2-3 lines close together) along each
side of stipe and rachis, in subg. Cyathea converging downwards on each side of
base of stipe and there often deeply excavated at maturity (fig. 7, 12). Fronds +
elliptical, lower pinnae always smaller than middle ones, sometimes gradually
much reduced and then the stipe short; pinnae normally pinnate-bipinnatifid, in a
few cases simple, in a few cases fully bipinnate; pinnules almost symmetrical at
the base, many on each pinna subequal, distal ones more or less abruptly decreas-
ing; upper surface of pinna-rachis and costa raised; veins simple or branched,
74 FLORA MALESIANA (ser. Lip volume
lower ones usually once forked, sometimes pinnate where pinnule-segments are
deeply lobed. Sori usually at the fork of a vein, or seated on a simple vein, a branch
of the vein always entering the receptacle; indusium either attached all round base
of receptacle and covering young sorus, opening to form a firm-edged cup or open-
ing by irregular rupture, or attached on costular side of receptacle (hemitelioid)
and of varying size, in some cases quite hidden by mature sorus, or lacking;
receptacle erect, + club-shaped to spherical; sporangia many, always short-
stalked; paraphyses usually present as multicellular hairs, sometimes flat and
several cells wide at base; in some species of swhg. Sphaeropteris scales present
round base of receptacle, more or less covering young sporangia; spores thin-
walled, smooth or papillose.
Type-species: Cyathea arborea (L.) SM. (tropical America).
Distr. & Ecol. See under the sections.
Morph. Dermal appendages. Scales on the stipes of Cyathea are of two kinds, and these appear to
provide the best subdivision of the genus, at least in Malaysia. The two types of scale are called flabelloid
and setiferous (see Ho_ttruM, Kew Bull. 1957, 41-45; Hottrum & SEN, Phytomorphology 11, 1961,
406-420).
Flabelloid scales (fig. 8a, b) have a broad median portion consisting of longitudinally elongated cells
with all walls thickened, and edges, of varying width, consisting of thin-walled cells diverging fan-wise
outwards, with irregularly projecting marginal cells, some of them sometimes thick-walled and dark,
often flexuous (fig. 9a, b). The scales develop at the apex of more or less massive (multicellular) out-
growths from the surface of the stipe and at right angles to these outgrowths (thus parallel to the surface
of the stipe); the base of a scale is peltate, with a narrow part encircling the supporting outgrowth on the
Fig. 8. Cyathea oinops Hassk. a. Scale from costa of pinnule, showing median band of dark thick-walled
cells and flabelloid margins of thin-walled cells with a few very thick-walled setae, x 70.—C. incisoser-
rata CopgL. b. Part of flabelloid margin of scale from stipe, < 100.—C. squamulata (BL.) COPEL. c.
Apical part of scale from costa of a pinnule, showing setiferous (not flabelloid) edge, < 70, c’. two very
small scales from costa, x 70.
basiscopic side. The outgrowths become very large and spine-like in some species, in others they are
quite small. As one proceeds from the base of the stipe to the ultimate axes, the scales become progressi-
vely smaller, and their character changes in ways characteristic of individual species. These scales pro-
vide some of the most important diagnostic characters in Cyathea.
Setiferous scales (fig. 8c, c’) also develop at the apex of outgrowths from the surface of the stipe,
but not at right angles to the outgrowths; the base of a scale widens more or less abruptly from the apex
of the outgrowth, and in C. sangirensis may be seen transitions from stout erect hairs (like those of
Dicksonia) to setiferous scales. All cells in setiferous scales are longitudinally elongate and all have walls
Dec. 1963] CYATHEACEAE (Holttum) = 75
Fig. 9. Different types of indusia in subg. Cyathea. a. C. batjanensis (CHRIST) COPEL.; indusium a small
disc covered by mature sorus.—b. C. javanica BL.; indusium saucer-shaped, often asymmetric.—c. C.
heterochlamydea CoreL.; indusium attached on costular side of receptacle, covering part of base of
mature sorus.—d. C. oinops HASskK.; indusium covering sorus almost to maturity, but open on side remote
from costule (two indusia cut to show receptacle, costular scales also shown).—e. C. orientalis (KUNZE)
Moore; indusium a complete rather deep cup.—f. C. crenulata BL.; indusium very fragile, covering sorus
completely to maturity, then breaking irregularly and in part disappearing (a, < 6; b-f, <x 10; a DE
VRIESE 323, c ELMER 11634, d MATTHEW s5.n., € KOORDERS 37469, f MEIJER 119).
716 FLORA MALESIANA a (ser: Dk volume
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 (Disphenia, 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 CoPpELAND in Cyathea. COPELAND’s Cyathea is divisible, on the basis of
scale-characters (see above) into two groups which I call subg. Cyathea (flabelloid scales) and subg.
Sphaeropteris (setiferous scales). Gymnosphaera (excluded COPELAND’s sect. 3) is then a section of
subg. 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 MeEtt., 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. /oheri 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. crenulata 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.
For 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. I 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
. Exindusiate; axes very dark, not hairy beneath; fertile and sterile pinnules eta) very dimorphous.
spp» 104=120) 3 2 = Macrae Sect. Gymnosphaera
1. Stipe-scales setiferous; hairs on hlower surfaces, Ae present, rather thick and Re. 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. Costules not widely spaced (rarely over 4 mm apart in pinnules 10cm 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 Dae or absent; sori never covered with overlapping scales.
sya, WIEN 5 5 6a 6 . . . 3a. Subsect. Sphaeropteris
4. Free 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 | 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. Spp. 152-178. . . . . . 4a. Subsect. 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) i | Ta
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. latebrosa (WALL.) COPEL., a common species in Malaya of
which I 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. incisoserrata CoPEL., otherwise very near C. /atebrosa, 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. multiflora
(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
Alsophila, A. australis 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
ES Pinnules not over 30 mm long; segments mostly constricted at base or the lower ones quite free.
Lower surface of pinna-rachis bearing crisped hairs.
a Sori lacking indusia ... . nll ra yk tee aes se Pay Orel elepidoclada
3. Sori indusiate; indusium cup- -shaped.
4. Bullate scales present on pinna-rachis and/or costae; stipe-scales to 2 mm wide.
Seabimmnules tole: 300mm long.) 2-6 sos 6 ccs sete es ee os 25 Camicrophylloides
Se eImUles toc) >) mimi longs. 4. . .. 3. C. perpelvigera
4. Bullate scales lacking from pinna- “rachis and/or costae; Stipe. -scales to 3144 mm wide.
4. C. hunsteiniana
2. Lower surface of pinna-rachis not hairy.
Gu leongest pinna 814 cm, pinnules to 11 mmilong:. «=... 2)... 2. . 195. Cx 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
lepiiei 5 6 6c .... . 7. C. hooglandii
1. Pinnules longer, or if “ioe 30 mm Mone without free basal ccomenis!
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. ............. . 410. C. pruinosa
11. Lower surface of lamina not glaucous.
12. Bullate-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 . . . . | ee el 22 amigens
14. Indusial cup deep. See hairs, not small fringed scales, also bullate scales, present on
pinna-rachis . . olny each SSE veverta!
12. Bullate-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. #3
17. Larger pinnules bearing c. 6 pairs of free tertiary leaflets. . . . 15. C. subtripinnata
78 FLORA MALESIANA [ser. II, vol. 12
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
COStACi ete sane ee 16. C. orientalis
18. Pinnules commonly to 65 mm long, basal pai of lobes ee. itone dark scales abundant on
COStae ie anes see tei oaks). RC wapoensis
16. Pinna-rachis persistently scaly 0 on lower surface.
19. Scales on pinna-rachis mostly narrow, 4-5 mm bees 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>ascalesonicostaeibullate-baseditojbullates: = 9.62) si 1 came cei) oe eC mpanva
24. Scales on costae flat or lacking, not bullate.
25. Broad flat pale scales with a few marginal setae oncostae ..... . . 22. C. wengiensis
25. Broad flat setiferous scales lacking on costae. .. . 5 . . 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).
27. 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 one hairy on lower surface; costae glabrous or nearly so on
upper suniacey .-) - ... . 26. C. javanica
30. Pinna-rachis hairy towards ‘apex < only; ‘costae “with few hairs on lower surface, many on
upper surface. . oa Saw eo Ss DAG whymenodes
29. Pinnules c. 30-70 by 12- 15 mm, veins we 6 pairs.
31. Lower 3-4 pairs of segments on each pinnule free and deeply lobed. Indusium irregularly
lobed ame ... . . 28. C. patellifera
31. Lowest segments not free, érenate, 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 bullate .... . . 30. C. catillifera
34. Scales on stipe pale; scales on pinna-rachis not bullate, larger ones having marginal setae.
31. C. horridula
32. Distal part of 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 134 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 eas rachises + covered beneath with small fringed scales as on
COstaem eee See, 35. C. trachypoda
39. Spines on stipe less than ‘1 mm long: rachises. not s so “covered ... . 36. C. crenulata
38. Small pale fringed scales lacking or very few.
40. Stipe bearing spines 3—S 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
43m bargest pinnules: ci 90) bys samme rs | = 3. 2405 nacanthophora
40. Stipe not spiny, or spines under | 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 ...... . 41. C. rubiginosa
44. Stipe-scales otherwise; pinna-rachis scales rarely setiferous.
AS Supe mGensely,. Scalys throughouteeem emer en cs cnn ae le apicniata
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 2!4 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 ne geo with stalks 3—6 mm long, to 10 mm on
WOES ee : Sa, sh hl ob sub DC longipes
56. Stipe not orn Binnie: ‘stalks on over 4 mien 9% 96 ... . 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 .. . tee es 535.@sansulana
54. Largest pinnules not over 100 by 20n mm; costules 3-41, 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.
GOS Stipes 50cm!) on more). ee Sect oh. Soy hawenaaes siooa, Para SateeiS ai al AeCHbOlolt
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 pean with dark median band. Veins concolorous, not raised
belows en 5a tates pte e Loe Gil SL ne sare cae CO NIMCAGH
59. Veins lacking scales | on fewer Sun btes.
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 .. . eh earn dr iam Ca) LOCTSLCTA
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 gees covered with small ue beneath;
veins bearing some scales .... Ree Sab eas ets . 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 See stalked een aa mm; costules 4414
mimneapant wer : . . 59. C. ferruginea
66. Pinnules sessile; costules usually not over aM mm apart.
67. Indusium firm, brown. ........ ae 3 oe oe sek oat O08 © OOSOLA
67. Indusium pale, fragile . . an een, mmr RY i) So an
65. Costae and/or costules lacking bullate series 2 “) teistés 6... ey oe g 4 Olea Ce nalconensrs
80 FLORA MALESIANA [ser. II, vol. 1?
48. Indusium hemitelioid (sometimes almost covering sorus to maturity, sometimes very small)
or lacking.
68. Indusium lacking.
69. Fronds simply pinnate, or bipinnate with small Sees lobed halfway to costa.
70. Fronds simply pinnate ... os 2 oe sss 2. (OZ @ pascemens
70. Fronds bipinnate, pinnules c. 30. mm Mone we) otic, See east es 2) aw OS SEO RCCIEN AES
69. Fronds bipinnate, pinnules more deeply lobed.
71. Pinna-rachis densely covered with a tangled pale crisped hairs and narrow scales on lower
SuLtace = scene ota os 3 > ss 6 OSD MerOphons
71. Pinna-rachis rather ‘sparsely hairy on lowe: 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) ... os 2 = JCS omaps
80. Bullate scales on costules. Tower pinnae gradually reduced to 7 cm long.
81. 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 ee ss « « 74 Co pachyanimens
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
OlmVcostules cae) = Py 75. C. latipinnula
82. Pinnules rarely over O; cm awities “costules not over 5A mm n 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 3-4 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... . Prana, 72. Charen ts
86. Indusium not more than a semicircle, Geered 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 444-54 mm apart. Pinnules 100 mm or more long. 85. C. alleniae
95. Costules 3-414 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; ees aun ee towards apex on lower
Suriaces a ane : 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) . . : . =... ...: . 1. 98. ©. microchlamys
106. Sori medial (New Guinea) .. . sit steer eer Coe, WENSIENSIS
103. Scales on costules bullate; some bullate scales Ghea on 1 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 3-4 mm long. Pinna-rachis hairy towards apex on lower surface.
100. C. alderwereltii
109. Stipe-spines 1-214 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. . . bee. 2 ol02-7@z meta
110. Lamina segments shallowly crenate. Indusium Bee 2-lobed; bullate scales
only on distal part of costa .... het Ah ser 03 a@Slatebresa
108. Paraphyses shorter than sporangia, slender.
112. Pinna-rachis hairy towards apex on lower surface. Lower pinnae much reduced and
spaced . . . Meh Mea 88. C. borneensis
112. Pinna-rachis not hairy < on i lower surface. Lowes 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 2144 mm long. ....... .. .. 90. C. junghuhniana
82 FLORA MALESIANA
[ser: Il, volar
1. Cyathea lepidoclada (CHRIST) Domin, Acta Bot.
Bohem. 9 (1930) 130; C. Cur. Brittonia 2 (1937)
278; CopEL. Philip. J. Sc. 77 (1947) 120.—Alsophila
lepidoclada Curist in K. ScH. & Laur. 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 24 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—S 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 314 mm 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: KeryssER 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) 11; Cope. Philip. J. Sc. 77 (1947) 119, 120.
Trunk to 24 m by 5 cm g; 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 10 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 3-4
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,
WSU):
Distr. Malaysia: New Guinea, Moluccas
(Ceram), N. Celebes (doubtful; plant young and
sterile).
Ecol. At 1200-1800 m 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 BrAuse, Bot. Jahrb. 56
(1920) 58 (incl. var. acuminata); COPEL. Philip.
J-aSe. 77 (W947) 19:
Trunk 2 cm g; fronds to 100 cm long. Stipe
8-16 cm; scales to 13 by 314 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 even 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. Btzg II, n. 28 (1918) 26.
Dec. 1963]
CYATHEACEAE (Holttum) =_ . 83
Fig. 10. Cyathea hooglandii HoLtrum. a. Part of pinna, upper surface, x 2, b. same, lower surface,
showing sori and bullate scales on pinna-rachis and costa, * 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, f. one tertiary leaflet showing sori, < 6, g. scale from costule, x 40 (a—b
HOOGLAND 7203, c HOOGLAND & SCHODDE 7506, d ditto 7171, e-g HOOGLAND & PULLEN 5745).
Stipe 12 cm, dark, warty; scales 10 by | mm.
Lamina c. 70 cm long, lowest pinnae 5 cm long,
longest pinna 814 cm. Pinnules to 11 mm by 4mm,
apex rounded, largest with 1 or 2 basal segments
free as almost circular tertiary leaflets, rest of
pinnule crenate; lowest pinnules of upper pinnae
defiexed 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. Grpss 6008, Anggi Lakes,
SW. Ridge, W. New Guinea (BM; dupl. at P, K).
Distr. Malaysia: W. New Guinea (2 collections).
Ecol. In undergrowth of mossy forest at 2500 m.
6. Cyathea ledermannii BrAuse, Bot. Jahrb. 56
(1920) 56 (incl. var. dilatata, I.c. 58); COPEL.
Philip. J. Sc. 77 (1947) 119.
Trunk slender, to 214 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
114-2 mm with pale edges. Largest pinnae 22 cm
long. Pinnules to 23 by 8 mm, with c. 7 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
214 mm apart; veins 5-6 pairs, mostly simple.
84 FLORA MALESIANA
[ser. Il, volai2
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 bullate
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 HoLttrumM, Kew Bull. 16
(1962) 56.—Fig. 10a.
Trunk to 3 m by 10 cm g, 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
114 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 1144 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. Cur. Brittonia 2 (1937)
279; CopEL. Philip. J. Sc. 77 (1947) 107.
Trunk 2-4 m by 8-12 cm g, 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 214-3 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 Ho_trum, 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!4 mm, 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 fiexuous 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 314 mm 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.
collection).
Ecol. At 3700 m, ‘frequent in edges of patches
of subalpine forest’.
Malaysia: NE. New Guinea (one
10. Cyathea pruinosa Rosenst. in Fedde, Rep. 12
(1913) 163; v. A. v. R. Handb. Suppl. (1917) 29;
CopeEL. 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 114 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—314
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 flat broad scales
as on costae, with ovate convex or almost bullate
scales distally; a few similar smaller scales on lower
surface of veins.
Type specimen: KerysseErR B44, Bolan Mts,
E. New Guinea (S—PA; dupl. at B, UC).
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 Hooc-
LAND & SCHODDE 7633, which also bears the
information that there were 14 fronds in two
whorls.
11. Cyathea pycnoneura HoLttrum, Blumea 11
(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, flabelloid 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 3-314 mm 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 bullate-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 rigens Rosenst. in Fedde, Rep. 12
(1913) 163; v. A. v. R., Handb. Suppl. (1917) 33;
CopeEL. Philip. J. Sc. 77 (1947) 103, 104.
Trunk 3-5 m tall; fronds numerous, spreading,
100—150(—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.
Lower 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 12-14 mm, deeply lobed, lowest segment
not free; costules 3—3!4 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 bullate-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: Krysser 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, pl. 2.—
C. globosora CopEL. Un. Cal. Publ. Bot. 18 (1942)
218; Philip. J. Sc. 77 (1947) 106, pl. 4.
Trunk up to 8 mm, 5-6 cm @g, 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 314 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 bullate-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. Il; vom
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; CopeEL. Philip. J. Sc. 77 (1947) 104.
Stipe 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 314 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 HoLttrum, Blumea 11
(1962) 534.
Trunk 2\% 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 114 mm 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. Sori 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; Metr. Ann. Mus. Bot. Lugd.-Bat. 1
(1863) 58; Hook. & BAK. Syn. Fil. (1865) 24;
Racis. FI. Btzg 1 (1898) 37; v. A. v. R. Handb.
(1908) 21; Suppl. (1917) 29; BACKER & Postn.
Varenfi. Java (1939) 24.—Disphenia orientalis
KUNZE, Bot. Zeit. 6 (1848) 283 (excl. syn.).—
C. arborea [non (L.) SM.] var. pallida HassKk. Obs.
Bot. Fil. 1 (1856) 15.—Fig. 7, 9e.
Stipe dark, to at least 50 cm, bearing copious
spines 114 mm 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 3-3!14 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 9, 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; Cope. Fern Fl. Philip. 2 (1960)
213.—C. lobata Cope. 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 3-3!4 mm apart; veins 7—9 pairs,
mostly forked; lamina-segments rather deeply
crenate-serrate or basal ones lobed up to 14 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 34 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]
les, but may be present on costules of sterile pin-
nules (none such seen).
Type specimen: ELMer 11482, Mt Apo, Min-
danao (Micu; 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, pl. 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 35-45
by 8-10 mm, lobed nearly to costa, lowest 1-2
segments sometimes a little constricted at base,
not free, apex abruptly pointed; costules 2-214
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 214 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 Ho_ttrum, 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 11 mm,
abruptly acuminate, lobed nearly to costa, lowest
segment nearly free; costules 244 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.
CYATHEACEAE (Holttum ) 87
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 HoLttrum, Blumea 11 (1962)
333:
Trunk 214 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 Copet. Un. Cal. Publ. Bot.
18 (1942) 219; Philip. J. Sc. 77 (1947) 120, pl.
1B
Trunk 114 m tall, 314 cm g, bearing 4 fronds
90 cm long. Stipe c. 10 cm, copiously warty;
scales to 15 by 114 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 114-2 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 1
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. I; volpet
with bullate bases, grading to rather large pale
bullate 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.
(1917) 68.—Alsophila hieronymi BRAUusE, Bot.
Jahrb. 49 (1912) 14; v. A. v. R. Handb. Suppl.
(1917) 67.—C. brauseana Domin, Pterid. (1929)
262.
Stipe to 10 cm or more, bearing spines to 2 mm
long; scales not seen. Lowest pinna 10 cm Jong,
longest 48 cm. Largest pinnules 70 by 12-18 mm
(sometimes larger; see note below); lobed toc. 1
mm from costa, lowest segment not quite free;
costules 314 mm 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 144 mm @ 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,
We):
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 5-514 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.) DomIN, 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
Curist in Warb. Monsunia (1900) 90; v. A. v. R.
Handb. (1908) 38.—Alsophila saparuensis v. A.
v. R. Bull. Dép. Agr. Ind. Néerl. 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. Lc. 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 314-5 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 fiexuous 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 214 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
414 mm apart; veins to 11 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;
duplearj Je):
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. Bot.
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 32-40 by 12 mm, lobed almost to costa;
costules 3-314 mm 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]
CYATH EACEAE (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),
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.
New
26. Cyathea javanica BL. En. Pl. Jav. (1828) 245
(incl. var. rigida, p.p.); Mrtr. Ann. Mus. Bot.
Lugd.-Bat. 1 (1863) 56; Hook. Syn. Fil. (1865)
23; RaAcis. Fl. Btzg I (1898) 35; v. A. v. R. Handb.
(1908) 23; Suppl. (1917) 32; BACKER & POosTH.
Varenfl. Java (1939) 24.—Hemitelia caudipinnula
v. A. v. R. Bull. Jard. Bot. Btzg II, n. 7 (1912)
16; Handb. Suppl. (1917) 48.—Hemitelia ba-
risanica V. A. Vv. R. Bull. Jard. Bot. Btzg II,
n. 20 (1915) 17; Handb. Suppl. (1917) 49.—
Alsophila benculensis 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, n. 23 (1916) 4; Handb. Suppl. (1917)
493.— C. benculensis v. A. v.R. Bull. Jard. Bot.
Btzg II, n. 28 (1918) 14.—C. palembanicav. A.v.R
lic. 13.—C. caudipinnula Domin, Pterid. (1929)
263.—C. barisanica Domin, /.c.—Fig. 9b.
Trunk to 10 m. Stipe 10-30 cm, bearing many
spines to 1 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
bullate-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 Metr. Ann. Mus. Bot.
Lugd.-Bat. 1 (1863) 57; Hook. Syn. Fil. (1865)24;
v. A. v. R. Handb. (1908) 24.—C. korthalsii
Metr. 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. amphicos-
mioides v. A. v. R. Bull. Jard. Bot. Btzg III,
2 (1920) 138.—C. arthropterygia v. A. v. R.
ibid. Il, 5 (1922) 188.—C. latebrosa var. indusiata
Hotttum, Gard. Bull. S.S. 8 (1935) 305; Rev.
Fl. Mal. 1 (1954) 121.
Stipe not spiny, sometimes with a pair of short
pinnae near base; scales 20-35 by 114-4 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 3-4 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, m. 16 (1914) 4; Handb. Suppl. (1917) Bile
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 35-40 cm
long. Largest 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. 1?
or less deeply crenate, free ones sometimes lobed
half-way to costule. Sori 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 CuHrist, Philip. J. Sc. 2
(1907) Bot. 181; v. A. v. R. Handb. (1908) 786;
CopeEL. Fern FI. Philip. 2 (1960) 214.
Stipe c. 15 cm long, warty; scales to 20 by 114
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.
Silay, Negros (P; dupl. at MICH).
Distr. Malaysia: Philippines (Negros, Leyte,
Biliran).
Ecol. At 1000 m.
1536, Mt
30. Cyathea catillifera HoLtrum, Kew Bull. 16
(1962) 53.
Trunk branching at base, 1 m by 10 cm g,
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 2-214 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
3-314 mm apart; veins to 8 pairs; lamina-segments
rather rigid, edges deeply crenate when fertile.
Sori near costules; indusium a dark brown disc
c. | 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, pl. 8.
Trunk 3 m, slender. Stipe 40 cm, bearing many
short spines (to 1 mm long), scaly near base;
scales to 15 by 1144 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 g, 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—11 cm long, longest
21 cm. Largest pinnules 40-50 by 13-15 mm, lobed
Dec. 1963 ]
CYATHEACEAE (Holttum) : oe: 91
to 1-2 mm from costa, no free basal segments;
costules 3 mm apart (fully fertile pinnules) to 4mm
(sterile); veins 7-9 pairs; lamina-segments entire
or slightly crenate. Sori 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: LEDERMANN 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;
vy. A. v. R. Handb. (1908) 25.—C. subuliformis
v. A. v. R. Bull. Jard. Bot. Btzg II, n. 11 (1913) 6;
Handb. Suppl. (1917) 32.—C. tuberculata 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 pinnae 50cm long. Pinnules to 85 by 18 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 (difference from C. javanica).
Type specimen: BeccaAri 438, Mt Singgalang,
Sumatra (K; dupl. at FI).
Distr. Malaysia: Sumatra, Malay Peninsula.
Ecol. In forest, 500-1500 m.
34. Cyathea klossii Ripit. Trans. Linn. Soc. II,
Bot. 9 (1916) 251; v. A. v. R. Handb. Suppl.
(1917) 489.
Trunk 114 cm @. 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. 1144 mm from costa; costules of pinna-lobes to
314 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: Kioss, 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) 191.—Alsophila alpinay. A. v. R.
ibid. I, n. 20 (1915) 4; Handb. Suppl. (1917) 62.—
C. alpinay. 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. Largest pinnules 100 by 18-20 mm, ses-
sile, lowest segment free, then 1—2 pairs adnate
and joined by a wing along costa; costules 314 mm
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: BUNNEMEVER 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. crenulata, but differs in its spiny stipe.
A specimen from Harau-Canyon, near Pa-
jakumbuh, at 800 m ( MEER 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. Pl. Jav. (1828) 244;
Hoox. Sp. Fil. 1 (1844) 25; Syn. Fil. (1863) 23;
Metr. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 57;
Racin. Fl. 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. latissima, f. squamulosa, f. subspinulosa);
BAacKER & PostH. Varenfl. Java (1939) 26.—
C. javanica var. rigida Bu. En. Pl. Jav. (1828) 245,
p.p.—() C. polycarpa JUNGH. Nat. Geneesk.
Arch. N.I. 2 (1845) 40; Flora 30 (1847) 522.—
C. excelsa [non Sw.|] KUNZE, Bot. Zeit. 6 (1848)
284.—C. spinulosa var. muriculata HASsK. in
Hook. J. Bot. Kew Misc. 7 (1855) 322; v. A. v. R.
&
Fig. 11. Cyathea crenulata Bu., showing how old
fronds persist and cover upper part of trunk.
Kawah Baru, Papandajan, Java (VAN STEENIS).
Handb. (1908) 25; Suppl. (1917) 36.—C. leucophaes
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. oinops HASSK.).—C. oinops (non HASSK.)RAC.
Fl. 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!4—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: BLuMme, 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. /atebrosa.
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 314-4 mm 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 flat pale scales bearing a
few short marginal hairs distally and on costules;
no bullate scales seen.
Type specimen: BUNNEMELJER 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 CurRist, 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 214 mm long, basal segments not
free, apex caudate-acuminate; costules 4414 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 1300-1700 m.
39. Cyathea magnifolia v. A. v. R. Bull. Jard. Bot.
Btzg III, 2 (1920) 135.—C. acanthopodav. A. v. R.
ibid. II, 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 4-514 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: BUNNEMEIJER 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. 16
(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
Tow, 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 4-414 mm apart; veins 9-10
pairs; lamina-segments rather thin, more or less
crenate, separated by sinuses 14 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; Cope. 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 4-414 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
1100-2840 m.
42. Cyathea apiculata (RosENsT.) Domi, 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 Il, n. 20 (1915) 2; Handb.
Suppl. (1917) 63.—C. crenulata f. subspinulosa
v. A. v. R. Handb. Suppl. (1917) 28.—C. indra-
purae Vv. A. v. R. Bull. Jard. Bot. Btzg II, n. 28
(1918) 13.—C. paleata CopreL. Un. Cal. Publ. Bot.
14 (1929) 372, t. 56.
94 FLORA MALESIANA
[ser. Ty volte
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 4414 mm apart: veins
10 pairs; lamina-segments slightly crenate, sinuses
narrow. Sori 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 bullate.
Type specimen: J. WINKLER, Pea Radja, Ba-
takerland, Sumatra (RoseENsT., 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-
drapurae, 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.
hymenodes METT.
43. Cyathea biinnemeijerii v. A. vy. 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 314-4 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-
Trangia, 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: BUNNEMEIWER 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 HoLtrum, 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
we
be
Fig. 12. Cyathea excavata HoLTTUM. Trunk-apex,
showing deeply excavated pneumathodes arranged
as V below base of a stipe. Cameron Highlands,
Malaya (R. E. HoLtrum).
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 14-14 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, flat; on veins many conspicuous very small
hairs (bases of former scales).
Type specimen: HoLtrumM 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, /.c.). Old
fronds persist, hanging down and covering the
trunk, as in C. orientalis (also in C. crenulata).
45. Cyathea christii Copev. Philip. J. Sc. 1, Suppl.
II (1906) 144; ibid. 4 (1909) Bot. 49: v. A. v. R.
Handb. (1908) 785; Suppl. (1917) 29; CopeEL.
Fern FI. Philip. 2 (1960) 217.
Stipe to 15 cm. Lower 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
3-314 mm 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; bullate 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,
Mindanao (MICH; dupl. at US).
Distr. Malaysia: Philippines (Mindanao).
Ecol. At 900-1800 m.
Mt Apo,
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;
CopeEL. Philip. J. Sc. 77 (1947) 102.—C. novo-
guineensis BRAUSE, Bot. Jahrb. 49 (1912) 12,
fig. 1B; 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 25-45 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 314-4 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. vy. M. Trans. R. Soc.
Victoria 1 (1889) 40; BAK. J. Bot. 28 (1890) 104;
v. A. v. R. Handb. (1908) 17; RipL. Trans. Linn.
Soc. II, Bot. 9 (1916) 251; C. Cur. Brittonia 2
(1937) 280.—C. keysseri ROSENST. in Fedde, Rep.
12 (1913) 164; v. A. v. R. Handb. Suppl. (1917)
23.—C. cheilanthoides Cope. Un. Cal. Publ. Bot.
18 (1942) 219; Philip. J. Sc. 77 (1947) 121, pl. 14.—
Fig. 10e, f, g, 13.
Trunk to 3 m, to 24 cm g, 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 2-214
mm apart. Largest fertiary leaflets commonly 3 by
214 mm, ovate to deltoid, edges strongly reflexed
and inrolled, 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. McGrecor 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. Ly voat4
Fig. 13. Cyathea macgregorii F. v. M., Mt Wilhelm, Eastern Highlands, Territory of New Guinea,
3450 m (R. D. HOOGLAND).
48. Cyathea gleichenioides C. Cur. Brittonia 2
(1937) 281; CopeL. Philip. J. Sc. 77 (1947) 123.—
Fig. 1-2, 14.
Differs from C. macgregorii as follows: pinnae
to 1014 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 | 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. Soc. II,
Bot. 4 (1894) 249; v. A. v. R. Handb. (1908) 22;
C. Cur. Gard. Bull. S. S. 7 (1934) 221.—C.
paleacea CopPEL. Philip. J. Sc. 12 (1917) Bot. 53.—
C. rigida CopeEL. l.c.
Trunk very short; fronds almost erect, to c. 100
Dec. 1963)
CY ATHEACEAE (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 30-40 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 Leptospermum-
Dacrydium forest of the ridges on Mt Kinabalu,
2400-3000 m.
50. Cyathea imbricata v. A. v. R. Nova Guinea
14 (1924) 11; C. Cur. Brittonia 2 (1937) 282;
CopEL. Philip. J. Sc. 77 (1947) 123.
Trunk to 2 m; fronds 60-70 cm long. Stipe
714-10 cm, dark, bearing spines | mm long, scaly
when young; largest scales 12-15 by 24 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 82 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,
EUS WKS).
98 FLORA MALESIANA
[ser. II, vol. 1?
Distr. Malaysia: Western New Guinea.
Ecol. In open forest at 3240 m.
51. Cyathea longipes Cope-. Philip.J.Sc. 12 (1917)
Bot. 54; C. Cur. Gard. 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 100-130 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 444-6 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 bullate 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-Paraispur 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 414-6 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: RAmMos & EDANo, BS 30900,
Jamindan, Capiz Prov., Panay (UC; dupl. at S,
WiStaike P2s 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 HoL_trumM, Kew Bull. 16
(1962) 56.
Trunk 8-10 m, 14 cm 9; 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 114-3 mm wide at base;
pneumathodes in a single row on each side of
stipe, 7-11 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 I.,
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, differing 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-bullate scales % 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. Cur. Brittonia 2 (1937)
278; Cope. Philip. J. Sc. 77 (1947) 105.—C. bi-
dentata CopEL. Un. Cal. Publ. Bot. 18 (1942) 218;
Philipy Je Sc. 77194) LOSS splees:
[Eyre Ihe
(BRAss 9430), showing habit and persistent stipe-
bases throughout trunk. Mt Wilhelmina, W. New
Guinea, 3200 m (L. J. BRAss).
Cyathea pseudomuelleri 'HOLTTUM
Fronds 200-300 cm long, about 10. Stipe c. 50cm,
base bearing short spines and covered with scales;
largest scales 30-40 by 114-2 mm, very pale to
CYATHEACEAE (Holttum ) . 99
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(—120) by 13—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 lobed up to 14 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
bullate; 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,
BRI, 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 differt: paleis stipitis atrocastaneis,
spinis ad 214 mm longis, paleis venarum paucioribus,
non 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. Cur. Brittonia 2 (1937) 279; Cope. 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
214-314 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. 0G, vole
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 pinnuie-
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 214-3 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 14 mm, with
dark shining mid-band and paler edges, and some-
what smaller, narrower scales bearing marginal se-
tae; scales oncostae many, small, dark, shining,with
many curved dark setae; on costules and veins
smaller, those on veins scattered but abundant.
Type specimen: HOoOGLAND & PULLEN 5495,
Eastern Highlands, NE. New Guinea (K; dupl. at
BOMUS a Sab SB Ribels):
Distr. Malaysia: Eastern New
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.
Guinea (4
58. Cyathea inquinans CHrRist, Verh. 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 14 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 (n. 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
EymMa had just descended from the mountains and
it seems possible that the specimen was collected
there.
59. Cyathea ferruginea CuHRISsT, Philip. J. Sc. 2
(1907) Bot. 181; v. A. v. R. Handb. (1908) 784;
Cope. 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 314-414 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] _
Ecol. In forest, to 1150 m, the type from mossy
forest, near summit of Mt Pulgar, other specimens
apparently from lower altitudes.
60. Cyathea oosora HoLtrum, Kew Bull. 16 (1962)
59.—C. assimilis [mon Hoox.] Curist, 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 314-4 mmapart; 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 bullate 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 CuHrist, Philip. J. Sc. 3
(1908) Bot. 270; CopeL. Philip. J. Sc. 4 (1909)
Bot. Sls v. Ay v. R. Handb. Suppl: (1917)) 24;
CopeEL. Fern FI. Philip. 2 (1960) 216.—C. mearnsii
CopeEL. 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. melanophlebia Copev. 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 314-414 mm
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.
CYATHEACEAE (Holttum)
101
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 814 by
134 cm, almost sessile, pinnatifid nearly to costa;
costules of pinna-segments 414 mm apart (fertile)
to 514 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 3-4 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) Doin, Acta Bot.
Bohem. 9 (1930) 153.—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 114, 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 214 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: 1; vole
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 HoL_trumM, Kew Bull. 16
(1962) 55.
Trunk to 3m; 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 3-314 mm 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 314 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,
Region, NE. New Guinea (B).
Distr. Malaysia: Eastern New Guinea (one
collection).
Ecol. In forest, growing in groups, 100 m.
Sepik
66. Cyathea modesta (BAK.) CoPEL. Philip. J. Sc. 4
(1909) Bot. 48.—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, n. 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,
lc. 263.—C. subconfluens Domin, Acta Bot.
Bohem. 9 (1930) 162.
Stipe 35-SO cm or more; spines bluntly conical,
hardly 1 mm high; scales abundant throughout,
to 30 by 114 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 14mm, 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 A/sophila modesta was an
unusually small specimen, with pinnules to
40 by 11 mm.
67. Cyathea doctersii v. A. v. R. Bull. Jard. Bot.
Btzg III, 2 (1920) 136.
Stipe-base 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 314-4 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)
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 Ho_ttrum, 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 14 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; rachis 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), 2-214 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 bullate
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.
ai? Ibe, 1XOR WWISS SADE EIRIDE
Distr. Malaysia: E. New Guinea (4 collections).
Ecol. Common in mountain forest, at c. 2400 m.
69. Cyathea setulosa CopEL. 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 314 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;
103
no bullate scales seen on costae and costules.
Type specimen: ALcaAsip & 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 g, 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
114 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 3-314 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.I. 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
-- 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. I; volaaé
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 14
Or more towards costule, lobes bidentate. Sori
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. Malaysia: Sumatra, Java, Lesser Sunda Is
(Lombok), SW. Celebes.
Ecol. At 2000-2500 m, in forest (specimen from
Lombok in Casuarina forest).
Note. In Fl. Btzg 1 (1898) 36, RAcrBoRSKI
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. crenulata BL.,
which see for further references.
72. Cyathea loheri Curist, 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) 211.—C. fruc-
tuosa CoPEL. in Elmer, Leafl. Philip. Bot. 2 (1908)
419; v. A. v. R. Handb. Suppl. (1917) 37; Cope-.
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; Copet.
Fern Fl. Philip. 2 (1960) 211.—C. campbellii
CopeEL. Philip. J. Sc. 38 (1929) 132; Fern FI.
Philip. 2 (1960) 210.—C. korthalsii (non METT.)
C. Cur. Gard; Bull. S. S. 7°(1934) 222== Gare
dusiosa CopeEL. Philip. J. Sc. 81 (1952) 14; Fern
Fl. Philip. 2 (1960) 211.
Trunk to 10 m. Stipe short; scales to 25 by
114-3 mm, pale, firm, their fragile edges bearing
scattered long dark setae; pneumathodes to 11 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 314-4 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., 7 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, Leaf.
Philip. Bot. 5 (1913) 1681; v. A. v. R. Handb.
Suppl. (1917) 36; Cope. Fern FI. Philip. 2 (1960)
DUIS).
Trunk 5 m, 12 cm @. Stipe bearing spines 8 mm
Dec. 1963]
CYATHEACEAE (Holttum)
105
long (fide CopPEL., 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!4 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
14 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 few long dark setae, also very small scales
as on pinna-rachis; on costules (of sterile pinnules)
pale entire bullate-acuminate scales.
Type specimen: ELMer 13860, Mt Urdaneta,
Agusan Province, Mindanao (US; dupl. at MICH,
KeBOse A. UG, L, U, BM):
Distr. Malaysia: Philippines (Mindanao, one
collection).
Ecol. At 1050 m.
74. Cyathea pachyrrhachis Copet. Un. Cal. Publ.
Bot. 18 (1942) 218; Philip. J. Sc. 77 (1947) 107, pl. 5.
Trunk to 7 m, 714 cm g, 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,
3A):
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 (30316) 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 Cope. in Elmer, Leafl.
Philip. Bot. 4 (1911) 1149; Fern FI. Philip. 2 (1960)
226.—Hemitelia latipinnula vy. A. v. R. Handb.
Suppl. (1917) 52.
Trunk 120 cm, 10 cm g; 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 brownscales 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: Ermer 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 Cope. Philip. J. Sc. 81
(1952) 17; Fern FI. 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 314 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 Subproy., Luzon (MICH;
dupl. at US, BO).
Distr. Malaysia: Philippines (Luzon, 2 col-
lections).
77. Cyathea loerzingii HoLtTruM, Kew Bull. 16
(1962) 58.
Stipe at least 40 cm, rather sparsely warty near
base; persistent scales few, to 20 by 24% mm,
shining dark brown with rather broad paler fragile
106
FLORA MALESIANA
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. 1mm 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: L6rRzING 14904, Mt Sibajak,
Sumatra (L; dupl. at BO).
Distr. Malaysia: Sumatra (one collection).
Ecol. In forest at 1300-1400 m.
78. Cyathea rufopannosa CuristT, Philip. J. Sc. 2
(1907) Bot. 180; v. A. v. R., Handb. (1908) 784;
CopEL. Fern FI. Philip. 2 (1960) 212.
Trunk (dry) 4 cm 9. 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 37 cm long. Pinnules
to 65 by 13-15 mm, sessile, short-acuminate, sey-
eral pairs of lower segments contracted at base,
lowest 1—3 quite free; costules 214-3 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 CurisT, 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; Cope. Fern FI. 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
[ser. I; vole
less free, rest of pinnule lobed nearly to costa;
costules 4-414 mm 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 Ho_LttumM, Blumea 11
(1962) 529.—Fig. 10c, d, 17.
Trunk to 3 m, 20 cm g, 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)
Fig. 17. Cyathea dicksonioides HoLTTUM (small plant in centre) with taller C. atrox C. Cur., in tree-fern
grassland, N. slopes of Sugarloaf complex, Western Highlands, Territory of New Guinea, 2880 m
(R. D. HooGLAnpb).
108
FLORA MALESIANA
[ser. Il, vole
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
RoseEnstT., Med. Rijksherb. 31 (1917) 2.—C. cau-
diculata Domin, Acta Bot. Bohem. 9 (1930) 104.—
C. merrillii CopeEL. Philip. J. Sc. 46 (1931) 212;
Fern Fl. 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 120 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 314-4 mm apart; veins com-
monly 10-11 pairs, on largest pinnules to 14 pairs;
lamina-segments firm, crenate, basal free ones
sometimes deeply so. Sori 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 Ul is) 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 Cope-. Philip. J. Sc. 46 (1931)
211; Fern Fl. 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 3-314 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 FI. Philip. 2 (1960) 224.
—Alsophila fuliginosa CuristT, Bull. Herb. Boiss. 6
(1898) 138; v. A. v. R. Handb. (1908) 39.—A/so-
Phila mindanensis CuHRIst in Warb. Monsunia
(1900) 90, p.p. (fertile specimen; sterile is Dickso-
nia); Vv. A. v. R. Handb. (1908) 44.—C. loheri var.
tonglonensis CuRisT, Philip. J. Sc. 2 (1907) Bot.
180; v. A. v. R. Handb. (1908) 787.—C. lanaensis
CHRIST, Philip. J. Sc. 3 (1908) Bot. 271; v. A. v. R.
Handb. Suppl. (1917) 23; Cope. Fern Fl. Philip.
2 (1960) 219.—C. mindanensis CorPEL. 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 Cope. in Elmer, Leafl. Philip. Bot. 5
(1913) 1680; Fern Fl. Philip. 2 (1960) 226.—
Alsophila warihon C. Cnr. Ind. Fil. Suppl. 2 (1917)
4.—Hemitelia bicolor vy. A. v. R. Handb. Suppl.
(1917) 44.—Hemitelia warihon v. A. v. R. L.c. 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 FI. Philip. 2 (1960)
206.—C. lepidigera CopeEL. 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. 7 cm, closely warty
or with conical spines under | mm long; scales to
15 by 114 mm, dark, shining, with fragile concol-
orous edges. Lower pinnae gradually reduced,
lowest c. 7 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)
Ne
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 bullate scales
distally, all scales light brown and entire; old
costae conspicuously warty beneath; costules
glabrescent or bearing bullate scales.
Type specimen: LOHER 893, March 1897, Baguio,
Luzon (P; dupl. at K, US).
Distr. Malaysia: Philippines (Luzon, Min-
danao, Biliran).
Ecol. In forest at 640-2400 m.
Notes. Loner’s n. 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 HoL_ttum, 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 3-414 mm apart; veins
to 7 pairs, not prominent on either surface; lamina-
segments firm, sterile ones crenate, larger fertile
ones lobed 14-34 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 alleniae HoL_trum, Kew Bull. 16
(1962) 52.
Trunk to 4m, 15 cm 9; 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 114 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 12 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. Fronds 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) 108.—Hemitelia montana v. A. Vv. 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 114 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 14 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).
110
FLORA MALESIANA
[ser IT, voliaiZ
87. Cyathea caudata (J. SM.) Cope. Philip. J. Sc.
1, Suppl. Il (1906) 144; Fern Fl. Philip. 2 (1960)
222.—Alsophila caudata J. SM. ex Hook. [J. Bot. 3
(1841) 419, nomen] Sp. Fil. 1 (1844) 52, t. 20B;
Syn. Fil. (1866) 42; v. A. v. R. Handb. (1908) 37,
785.—Hemitelia manilensis Pr. Abh. K. Bohm.
Ges. Wiss. V, 5 (1848) 351; v. A. v. R. Handb.
Supplement (1917) 53.—Amphicosmia manilensis
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 CopPEL. Philip. J. Sc. 46
(1931) 211; Fern Fl. Philip. 2 (1960) 217.—C.
brevipes CoPEL. Philip. J. Sc. 81 (1952) 18; Fern FI.
Philip. 2 (1960) 232.—C. arborescens COPEL.
Philip. J. Sc. 84 (1955) 162.
Stipe short, copiously warty; scales narrow, dark,
shining, to 15 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 3-314 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 bullate.
Type specimen: CUMING 267, Luzon (K; dupl.
atelwlawA):
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 BeEvDD.
J. Bot. 31 (1893) 225; Rip. J. Mal. Br. R. As. Soc.
4 (1926) 9.—C. hemichlamydea Copet. 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;
Ho.Lttum, Gard. Bull. S. S. 8 (1935) 306, pl. 30;
Rev. Fl. Mal. 2 (1954) 121.
Trunk to 2 m or more. Stipe 5—25 cm, short-
spiny or warty; scales to 15 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 314-5 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 suffused 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 (/eg. KING’s Coll., Perak).
89. Cyathea fenicis Cope. Philip. J. Sc. 3 (1909)
Bot. 354; ibid. 4 (1909) 61; Fern FI. 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 314-444 mm
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. 58.—Alsophila extensa [non
(Forst.) Spr.] BL. En. Pl. Jav. (1828) 246 (p.p.?).
—Alsophila lunulata [non (Forst.) R. Br.] BL. /.c.
(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 Vrtese, 1.c.—Alsophila melanopus
Hassk. in Hook. J. Bot. Kew Misc. 7 (1855) 325;
Dec. 1963]
CYATHEACEAE (Holttum)
wan
Obs. Bot. Fil. 1 (1856) 42; v. A. v. R. Handb.(1908)
40; Suppl. (1917) 65(?).—Amphicosmia javanica
Moore, Ind. Fil. (1857) 60.—Hemitelia jung-
huhniana Metr. Fil. Lechl. 2 (1859) 31; Ann.
Mus. Bot. Lugd.-Bat. 1 (1863) 55; Racis. FI.
Btzg 1 (1898) 38, p.p., excl. var. dissoluta; v. A.
v. R. Handb. (1908) 28; Suppl. (1917) 45, Corr.
44; BACKER & PostH. Varenfl. Java (1939) 28.—
Hemitelia latebrosa (WALL.) Metr. Ann. Mus.
Bot. Lugd.-Bat. 1 (1863) 54, p.p. (pl. jay. tantum);
Racis. Fl. Btzg 1 (1898) 39; v. A. v. R. Handb.
(1908) 38, p.p.: BACKER & PosTtH. Varenfl. Java
(1939) 27. —C. melanopus CoPEL. Philip J. Sc. 4
(1909) Bot. 48. —Hemitelia glaucophylla v.A.v.R.
Bull. Jard. Bot. Btzg II n. 7 (1912) 16; Handb.
Suppl. (1917) 50.—Hemitelia alsophiliformis v. A.
v. R. Bull. Jard. Bot. Btzg II, n. 16 (1914) 15;
Handb. Suppl. (1917) 46.—Hemitelia merapiensis
v. A. v. R. Bull. Jard. Bot. Btzg II, n. 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. glaucophylla Domin, Pterid.
(1929) 264.—C. alsophiliformis Doin, /.c. 263.—
C. merapiensis Doin, I.c. 264.—C. fallax 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
114—2'44 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—115 by 14-21 mm, sessile, acuminate,
lobed almost to costa, lowest segment not free;
costules 314,—414 mm apart; veins 10-12 pairs;
lamina-segments firm, subentire to distinctly cre-
nate (the latter usually when fertile). Sori 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 bullate, or acuminate with bullate 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. /atebrosa
(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. 1 (1844) 36, p.p.; Syn. Fil.
(1865) 29, p.p.—Hemitelia crenulata Metr. Ann.
Mus. Bot. Lugd.-Bat. 1 (1863) 55; Racts. FI.
Btzg 1 (1898) 38; v. A. v. R. Handb. (1908) 27;
Suppl. (1917) 42; BAcKER & PosTH. Varenfl. Java
(1939) 27, non C. crenulata BL. 1828.—Alsophila
crenulata Hook. Syn. Fil. (1866) 44.—Hemitelia
junghuhniana var. dissoluta RAcis. Fl. Btzg (1898)
38.—Alsophila brevifoliolata vy. 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, n. 28 (1918) 13.
Trunk rarely to 2 m. Stipe 30-50 cm, warty near
base; scales little over 10 mm by 1 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
14(—20) mm, sessile, abruptly narrowed at apex or
short-acuminate, lobed almost to costa, lowest
segment not free; costules 314 mm (rarely to 414
mm) apart; veins to 8 pairs; 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 HoLtrum, 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 114 mm,
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 6-614 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 5-514 mm 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).
FLORA MALESIANA
[ser. Il} voltae
Fig. 18. Cyathea incisoserrata CopEL. a. Pinnules attached to pinna-rachis, upper surface,
< 10, d. indusium and base of receptacle,
< 20, e. scales from costule,
x 20, g. scale from stipe, < 6, h. paraphyses, < 50, i. section of stipe, <
Y 2/, b
an 35 °
pinna-rachis and base of pinnule, upper surface, < 4, c. as b, lower surface, showing sori and scales,
x 20. f. scale from pinna-rachis,
4/,, 7. 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.
Schizocaena.
93. Cyathea punctulata v. A. v. R. Bull. Jard. Bot.
Btzg II, n. 28 (1918) 13.—Alsophila punctulata
v. A. v. R. Bull. Jard. Bot. Btzg II, n. 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 4144 mm
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 1% 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 mumerous; 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 bullate scales, very small irregular short-fringed
scales also abundant; most scales on costules
bullate, 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; Hottrum, Gard. Bull. S. S. 8
(1935) 305.—Alsophila ornata var. sikkimensis (non
Ci. & BAK.) Bepp. J. Bot. 31 (1893) 225.—
Alsophila ornata (non Scott) Bepp. Kew Bull.
(1909) 423.—Alsophila incisoserrata C. Cur. Ind.
Fil. Suppl. (1913) 5; v. A. v. R. Handb. Suppl.
(1917) 72.—Alsophila latebrosa var. ornata RIDL.
J. Mal. Br. R. As. Soc. 4 (1926) 8.—Fig. 7, 8b, 18.
Trunk to 4 m, 12 cm @ including leaf-bases.
Stipe to 85 cm, warty or with conical spines to
114 mm 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 414-514 mm
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. Sori 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. Brooxs 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. Jatebrosa, 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. /atebrosa.
95. Cyathea physolepidota ALstoN, Nova Guinea
n.s. 7 (1956) 1.
Stipe 25 cm, strongly spiny, spines 3-5 mm long;
scales sparse, to 10 by 114 mm, brown with pale
fragile edges which have many slender hairs
irregularly directed (not all defiexed as originally
described). Main rachis smooth and rather pale,
glabrescent. Lower pinnae reduced (lowest not
seen); longest pinna 22 cm long. Largest pinnules 32
by 9 mm, sessile, short-acuminate, lobed to within
1 mm of costa, lowest segment not free; costules
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 HoL_trum, nom. noy.—
Alsophila arfakensis Gepp in Gibbs, Arfak (1917)
70 (non C. arfakensis GeEpp, I.c.).
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. Gipsps 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 Ho_trum, 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 2-4 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 3-414 mm 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. Il; vole
Ecol. In Nothofagus forest, 2000 m.
98. Cyathea microchlamys HoL_trum, 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 14mm
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
collection).
Note. This appears to be very near C. caudata,
but has a long stipe and small indusia.
(Luzon, one
99, Cyathea perpunctulata (v. A. v. R.) DomIN,
Acta Bot. Bohem. 9 (1930) 146.—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 314 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 CopPeL. Philip. J. Sc. 4
(1909) Bot. 50.—Hemitelia sumatrana vy. A. v. R.
Bull. Dép. Agr. Ind. Néerl. m. 18 (1908) 2; Handb.
(1908) 28; Suppl. (1917) 48.—Hemitelia horridipes
v. A. v. R. Bull. Jard. Bot. Btzg II, nm. 16 (1914) 16;
Handb. Suppl. (1917) 47.—Hemitelia salticola
v. A. v. R. Bull. Jard. Bot. Btzg II, n. 20 (1915) 18;
Handb. Suppl. (1917) 50.—Hemitelia paraphy-
sophora v. A. v. R. Bull. Jard. Bot. Btzg III, 2
(1920) 154.—Alsophila spinifera v. A. v. R. ibid.
Ill, 5 (1922) 182.—C. horridipes Domin, Pterid.
(1929) 264.—C. salticola Domin, I.c.—C. pa-
raphysophora Domin, Acta Bot. Bohem. 9 (1930)
145.—C. spinifera Domin, /.c. 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 314 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.
101. Cyathea amboinensis (v. A. v. R.) MERR.
Interpr. Rumph. Herb. Amb. (1917) 63.—
Alsophila latebrosa var. batjanensis CHRIST in
Warb. Monsunia (1900) 89.—A/sophila amboinen-
Sis’ V. Aw Ve Re Philip. J; Sc: 11) (SiG) eBota0s:
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 314-4 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)
LpS
from Menado collected by PosrHuUMuUs are referred
with some doubt to this species.
102. Cyathea media WAGN. &. GreTH. Un. Cal.
Publ. Bot. 23 (1948) 44, pl. 15.
Stipe 20-25 cm, dark, warty; scales to 20 by
114 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 314 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. Cur. Gard. Bull.
S. S. 7 (1934) 222; HoLttum, ibid. 8 (1935) 303,
pl. 29, p.p. max.; Rev. Fl. Mal. 2 (1954) 120.—
Polypodium latebrosum WALL. Cat. (1828) n. 318,
nomen.—Alsophila latebrosa WALL. ex HOoK. Sp.
Fil. 1 (1844) 37; Syn. Fil. (1866) 43, p.p.; BEepp.
Handb. (1883) 11, p.p.; v. A. v. R. Handb. (1908)
38, 789, p. p.—Dichorexia latebrosa PresL, Abh.
K. BG6hm. Ges. Wiss. V, 5 (1848) 344.—He-
mitelia latebrosa Metr. Fil. Hort. Lips. (1856)
111; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 54, p.p.;
v. A. v. R. Bull. Jard. Bot. Btzg II, n. 23 (1916) 13,
incl. var. paraphysata v. A. Vv. R.; Handb. Suppl.
(1917) 51, p.p., 489.—C. leucocarpa CoPEL. Philip.
J. Sc. 6 (1911) Bot. 362.—C. longipinna Copev.
l.c. 363.—Alsophila leucocarpa C. Cur. Ind. Fil.
Suppl. (1913) 5; v. A. v. R. Handb. Suppl. (1917)
66.—Alsophila longipinna C. Cur. Ind. Fil. Suppl.
(1913) 5; v. A. v. R. Handb. Suppl. (1917) 67.—
Alsophila lastreoides vy. A. v. R. Bull. Jard. Bot.
Btzg II, m. 23 (1916) 5; Handb. Suppl. (1917) 495.
—Hemitelia leptolepia v. A. v. R. Bull. Jard. Bot.
Btzg II, n. 23 (1916) 12; Handb. Suppl. (1917) 488.
—RHemitelia rudimentaris v. A. vy. R. Bull. Jard.
Bot. Btzg III, 5 (1922) 205.—C. lastreoides Domin,
Acta Bot. Bohem. 9 (1930) 128.—C. leptolepia
Domin, /.c. 130.—C. rudimentaris Domin, I.c. 154.
Trunk to c. 3 m, rather slender. Stipe 50 cm or
more, closely short-spiny (spines 1-214 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 3-314 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. nov.—Gymnosphaera BL. En. Pl. Jay. (1828) 242; CopEL.
Gen. Fil. (1947) 98, p.p. (excl. sect. 3).—Thysanobotrya v. A. Vv. R. Bull. Jard.
Bot. Btzg II, n. 28 (1918) 66, t. 10.—A/sophila sensu C. CHR. Dansk Bot. Ark. 7
(1932) 37.—Cyathea subg. Gymnosphaera TINDALE, Contr. N. S. W. Nat. Herb. 2
(i956) 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. | have included C. macgillivrayi in the keys to both sections.
116 FLORA MALESIANA [ser.! IL voleuie
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 of normal 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. ieee ate: 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.
Ne Segments of reduced basal pinnae all with broad lamina
106. C. atropurpurea
107. C. recommutata
. 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 .
6. Fertile pinnules only slightly lobed .
5. Fronds with normal apex.
7. Pinnules almost entire.
108. C. biformis
109. C. scandens
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.
7. Pinnules distinctly lobed.
9. Scales on lower surface of costules bullate.
111. C. glabra
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.
11. Pinnules to 65 mm long without free basal segments .
10. Bullate scales lacking on veins.
12. Axes very dark; scales on lower surface of costae bearing many setae .
C. hornei
113. C. dimorpha
114. C. lurida
12. Axes not very dark; scales on lower surface of costae sometimes with a few setae.
13. Pinnules to 110 by 30 mm, cut to 2 mm from costae; costules of sterile pinnules to 6 mm apart.
115. C. rubella
13. Pinnules to 70 by 20 mm, cut to within 1 mm from costa; costules of sterile pinnules to 4144 mm
apart.
9. Scales on lower paniaee of costules not Bulla’
14. Pinnules lobed almost to costa.
a Costal scales not setiferous :
. Costal scales bearing many setae .
116. C. macgillivrayi
117. C. acrostichoides
118. C. schlechteri
ie ‘Pinnules not lobed more than 24 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.
7h 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 of stipe; largest pinnules with stalks to4mm; 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.
104. Cyathea annae (v. A. v. R.) Domin, Acta
Bot. Bohem. 9 (1930) 90.—Alsophila annae v. A.
v. R. Bull. Jard. Bot. Btzg Il, n. 23 (1916) 3;
Handb. Suppl. (1917) 490.—Fig. 19c.
Stipe dark, slender; scales to 60 by 1 mm, dark
and shining with dull paler edges. Basal pinnae
about 4 on each side of the stipe, all within 11 cm
120. C. gigantea
from the base, lowest with very narrow lamina
on each side of veins and midribs of pinnules,
upper ones with pinnules to 20 by 7 mm with cre-
nate edges; largest pinnae 24 cm long, sterile and
fertile pinnules dimorphous. Sterile pinnules to
43 by 10 mm, almost sessile, base broad, apex
gradually narrowed, edges lobed to 14 towards
Dec. 1963]
CYATHEACEAE (Holttum)
/
Lf»,
/ &
bs _
/
/
2 Fs os
7 SS
]
=,
Ti}
SEZ SS CY
ALAS
Fig. 19. Cyathea recommutata CopEL. a. Part of pinna showing transition from fertile to sterile pinnules,
x 24, b. lower surface of fertile pinnule, x 4.—C. annae (v. A. v. R.) Doin. c. Reduced pinnae at
base of stipe, < 24 (a—-b KUNSTLER 7130, c cult. Hort. Bog.).
costa; costules 3-314 mm apart; veins 2-3 pairs,
simple. Fertile pinnules to 30 by 8 mm, lobed 34
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.) Cope-. Philip. J.
Sc. 4 (1909) Bot. 36; Sarawak Mus. J. 2 (1917)
346, 349; C. Cor. & HoLtrum, Gard. Bull. S.S.
7 (1934) 200, 220.—Alsophila ramispina HooK.
Syn. Fil. (1866) 42; v. A. v. R. Handb. (1908)
34; Curist, Ann. Jard. Bot. Btzg 20 (1906) 138.
—Alsophila burbidgei (non BAK.) CHRIST, Ann.
Jard. Bot. Btzg 20 (1906) 138, p.p.—Alsophila
hallieri v. A. v. R. Bull. Jard. Bot. Btzg II, 7. 28
(1918) 2 (non RoseEnst.).—Alsophila amaiambiten-
sis Vv. A. v. R. Le. 1.—Alsophila kenepaiana
v. A. v. R. ibid. III, 2 (1920) 129.—C. kenepaia-
na Domin, Acta Bot. Bohem. 9 (1930) 127.—
C. amaiambitensis Domtn, /.c. 90.—Gymnosphaera
ramispina CopeL. 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 114 mm, with narrow thin pale
edges. Basal pinnae 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. Tl; voles
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 4414 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: Lops, Sarawak (K).
Distr. Malaysia: Borneo.
Ecol. In rather exposed places on mountain
tidges 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. vy. 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, pl. 18.—
Alsophila atropurpurea C. Cur. Ind. Fil. Suppl.
(1913) 4.—Gymnosphaera atropurpurea COPEL.
Gen. Fil. (1947) 98; Fern FI. Philip. 2 (1960) 234.
Differs from C. ramispina 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. Malaysia: Philippines (Luzon, Mindoro,
Leyte, Mindanao).
Ecol. At altitudes of 1000 m and over.
107. Cyathea recommutata CopeL. Philip. J. Sc.
4 (1909) Bot. 36; C. Cur. Gard. Bull. S.S. 7 (1934)
220; Hottrum, Rev. Fl. Mal. 2 (1954) 125.—
Gymnosphaera squamulata (non BL.) J. SM. ex
Hook. Gen. Fil. (1842) t. 100.—Alsophila squa-
mulata Hook. Sp. Fil. 1 (1844) 51, p.p.; Bepp.
Ferns Br. Ind. (1867) t. 235.—Alsophila com-
mutata Metr. Ann. Mus. Bot. Lugd.-Bat. 1 (1863)
53; BeEpp. Handb. (1883) 14: v. A. v. R. Handb.
(1908) 34 (non C. commutata SPR.).—C. hewittii
CopeEL. Philip. J. Sc. 6 (1911) Bot. 134, t. 14.—
Alsophila heteromorpha v. A. v. R. Bull. Jard.
Bot. Btzg II, n. 16 (1914) 1; Handb. Suppl. (1917)
56; Bull. Jard. Bot. Btzg III, 2 (1920) 129, incl.
var. decomposita v. A. Vv. R.—Alsophila hewittii
v. A. v. R. Handb. Suppl. (1917) 55; C. Cure.
Gard. Bull. S.S. 7 (1934) 221.—C. toppingii
CopeL. Philip. J. Sc. 12 (1917) Bot. 51; C.. Cnr.
Gard. Bull. S.S. 7 (1934) 220.—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 CopPEL. I.c.
—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; /argest pinnae 40 cm. Pinnules dimorphous
(sometimes with intermediate conditions); sterile
pinnules commonly 60—-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 4414 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 3-4 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.) CopPEL. Philip.
J. Sc. 6 (1911) Bot. 364; ibid. 77 (1947) 117.—
Stenochlaena dubia v. A. v. R. Bull. Dép. Agr.
Ind. Néerl. n. 18 (1908) 26; Handb. (1908) 721;
Ho.ittum, Gard. Bull. S.S. 5 (1932) 250, non C.
dubia (BEDD.) DomINn, 1929.—Alsophila biformis
RosENST. in Fedde, Rep. 9 (1911) 423; v. A. v.
R. Handb. Suppl. (1917) 71; BRAusE, Hedwigia 61
(1920) 401.—Polybotrya arfakensis Gerp in Gibbs,
Arfak (1917) 71.—Thysanobotrya arfakensis Vv.
A. v. R. Bull. Jard. Bot. Btzg II, n. 28 (1918)
66, t. 10.—C. gibbsiae CopeL. Philip. J. Sc. 38
(1929) 129.—Gymnosphaera biformis CoPEL. Gen.
Fil. (1947) 99.—Fig. 20.
Stem 1-114 cm g, 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)
mae |
1 EU ULE
ih OS ORNY WPA Ma te,
rs
EEC,
one : AAS wren
SSN Ys ee } |
a
Fig. 20. Cyathea biformis (ROSENST.) COPEL. a. Fertile pinnules, =
part of frond, showing abortion of true apex,
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, base slightly 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 4-1!
towards costa; fertile pinnules stalked (3-5 mm),
c. 50 by 4-10 mm, lobed almost or quite to the
costa, costules 314 mm apart, lobes 1 14-2 mm wide,
entire with rounded tips, basiscopic edge of
a
< 24, c. apical
< 24, b. sterile pinnules,
x 2%, d. part of c, x 8 (a-b BRAss 6806, c-d BRASS 8947).
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 bullate scales seen on sterile leaflets.
Type specimen: CopLAND KING 57, Papua
(S-PA; dupl. at MICH, BO).
Distr. Malaysia: Moluccas (Ambon),
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 Nothofagus forest.
Note. The only specimen with rather deeply
lobed sterile pinnules is Grpss 5984, from 2200 m
(type of Polybotrya arfakensis GEPP).
New
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. Il, voli
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 bullate 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 (F. v. M.) Domin, Pterid.
(1929) 263; Acta Bot. Bohem. 9 (1930) 153;
TINDALE, Contr. N.S. W. Nat. Herb. 2 (1956) 334.
—Alsophila rebeccae F. v. M. Fragm. 5, xxxiil
(1865) 53, xxxvi (1886) 117.
Stipe very dark at base, finely warty after fall
of scales; scales to 15 by 1!4 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 24, broadly
blunt-serrate in distal 14; 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 bullate 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 214 m long.
111. Cyathea glabra (BL.) CopEL. Philip. J. Sc. 4
(1909) Bot. 35; Hottrum, Gard. Bull. S.S. 8
(1935) 316; Rev. Fl. Mal. 2 (1954) 127.—Gym-
nosphaera glabra Bui. En. Pl. Jav. (1828) 242;
CopeL. Gen. Fil. (1947) 98.—Alsophila glabra
Hook. Sp. Fil. 1 (1844) 51; Metr. Ann. Mus.
Bot. Lugd.-Bat. 1 (1863) 52, p.p. (?); BEDD.
Handb. (1883) 14, p.p.; RActBorskKI, Fl. Btzg 1
(1898) 34, p.p.—Alsophila vexans Crs. Atti Ac.
Napol. 7, n. 8 (1876) 4.—Alsophila dubia BEvp.
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) 1.—C. reducta Domin,
Acta Bot. Bohem. 9 (1930) 153.—C. vexans C.
Cur. Gard. Bull. S.S. 7 (1934) 218.—Gymno-
Sphaera vexans Cope. 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 2-4 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 HooKker’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.) Cope. Bull. Bern.
P. Bish. Mus. 59 (1929) 38; Philip. J. Sc. 77 (1947)
119.—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.c. 76.—
Alsophila olivacea BRause, l.c. 74.—Alsophila
melanocaulos v. A. v. R. Nova Guinea 14 (1924) 1.
—C. dissitifolia Domtn, Pterid. (1929) 262.—
C. brunnea Domin, Acta Bot. Bohem. 9 (1930) 101;
CopeEL. Philip. J. Sc. 77 (1947) 117, 119.—C. di-
morphophylla Domtin, Acta Bot. Bohem. 9 (1930)
111 (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. melanoclada Domin,
Acta Bot. Bohem. 9 (1930) 174 (new name for
Alsophila melanocaulos); CopreL. Philip. J. Sc. 77
(1947) 118.—Gymnosphaera hornei CopeL. Gen.
Fil. (1947) 99.—Gymnosphaera melanoclada Co-
PEL. l/.c.
Trunk rather slender (4 cm @ when dry), to
3 or 4 m tall; leaf-scars to 2 cm go. 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—SO
cm long (60 cm reported of A/sophila 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 24 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 314-4
mm apart; veins usually fewer than in sterile
pinnules, often forked; segments crenate, usually
separated by rather wide sinuses; sori 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.
ramispina and C. recommutata 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 (CuHrRIsT) CopeL. Philip.
J. Sc. 4 (1909) Bot. 34.—Alsophila dimorpha
Curist, Ann. Jard. Bot. Btzg 19 (1904) 41;
yv. 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. Srerile pinnules to 65 by 20 mm, on
stalks to 3 mm, near base lobed 34 towards costa,
for the most part lobed only 144 towards costa;
costules 3!4 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 214 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.) Cope. Philip. J. Sc. 4
(1901) Bot. 45.—Chnoophora lurida Bui. En. PI.
Jav. (1828) 244.—Alsophila lurida Hoox. 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 CopPeL. Philip. J. Sc.
4 (1909) Bot. 38.—C. kingii Cope. lc. 56;
Ho.trum, Gard. Bull. S.S. 8 (1935) 315, pl. 36;
Rey. Fl. Mal. 2 (1954) 126.—C. subdimorpha
CopeL. Philip. J. Sc. 8 (1913) Bot. 140, pl. 2.—
Alsophila heterophylla v. A. vy. R. Bull. Jard. Bot.
Btzg Il, n. 16 (1914) 2; Handb. Suppl. (1917) 60.
—Alsophila subdimorpha v. A. v. R. Bull. Jard.
Bot. Btzg Il, n. 16 (1914) 2.—C. heterophylla
Domin, Pterid. (1929) 262.—Gymnosphaera me-
lanorachis CopreL. Gen. Fil. (1947) 98; Fern FI.
Philip. 2 (1960) 234.—Gymnosphaera_ kingii
Cope. 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 114 mm, 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 314-444 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. I> volpa
Fig. 21. Cyathea lurida (BL.) CopeL. a. Base of a fertile pinna,
pinna, < 24, c. normal form of sterile pinna,
reduced pinnae,
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 Fraser’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. Jurida in Malaya. No specimens of
C. lurida have been found in Borneo, so that there
is a considerable gap in distribution.
115. Cyathea rubella Ho_trum, Kew Bull. 16
(1962) 61.
Trunk to 5 m; fronds many, to 200 cm long.
VV
\NN WN} AN NUNES oe
x V4 (a-c ERY SmitH 833, d after HoL_trum, Flora of Malaya
€
ex
Q
Za
Pa
Za gp fv
= 4 ij
of
==
Ox
=
x 24, b. base of unusually large sterile
< 24.—C. recommutata COPEL. d. Base of frond, showing
2, fig. 50).
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 414-6 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 distally,
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 I.
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 Core. Un. Cal. Publ.
Bot. 18 (1942) 219; Philip. J. Sc. 77 (1947) 118,
pl. 11.—Gymnosphaera 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. \t 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.) Domin,
Acta Bot. Bohem. 9 (1930) 88.—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 314-414 mm 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), W. 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 514 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) Domi, Pterid.
(1929) 263; CopeL. Philip. J. Sc. 77 (1947) 117,
119.—Alsophila schlechteri BRAUSE, Bot. Jahrb. 49
(1912) 15, f. 1D; v. A. v. R. Handb. Suppl. (1917)
61.—Gymnosphaera 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 4414 mm
apart, veins 10 pairs, mostly forked; fertile part
to 11 mm wide, fertile segments little over 2 mm
124
FLORA MALESIANA
[ser. Il, -volai@
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) 263.—Alsophila subdubia v. A. v. R.
Bull. Jard. Bot. Btzg II, n. 20 (1915) 3; Handb.
Suppl. (1917) 54.—Alsophila persquamulata vy. 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. Pinnae to 50cm long, lower ones smaller.
Largest pinnules 90-115 by 15-20 mm, almost
sessile, lobed 14-24 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.
Ecole Atic: 1500) m:
120. Cyathea gigantea (WALL.ex Hook.) HOLTTUM,
Gard. Bull. S. S. 8 (1935) 318; Rev. Fl. Mal. 2
(1954) 128.—Alsophila gigantea WALL. ex Hook.
Sp. Fil. 1 (1844) 53.— Alsophila polycampta
Kunze, Bot. Zeit. 4 (1846) 475.—Alsophila
helferiana Prest, Abh. K. Béhm. Ges. Wiss.
V, 5 (1848) 341.—Alsophila glabra [non BL.]
Bepp. Ferns S. India (1863) t. 60; Handb. (1883)
14; Hoox. & BAK. Syn. Fil. (1866) 43, p.p.;
J. Scott, Trans. Linn. Soc. 30 (1874) 38; CLARKE,
ibid. 11, Bot. 1 (1880) 433; RactporskI, FI. Btzg 1
(1898) 34; v. A. v. R. Handb. (1908) 32.—
Alsophila umbrosa WALL. ex RiDL. J. Mal. 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 114 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, 80-110 by 15-20 mm, evenly nar-
rowed to apex, edges lobed 14—24 towards costa;
costules 414-6 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,
ISO DD:
Type species: Cyathea medullaris (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 Sop., C. insignis EATON,
C. princeps (LINDEN) Meyer, and C. brunei CurRist.
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.
CYATHEACEAE (Holttum) 125
3a. 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 are 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. contaminans (WALL.) CoPEL. (fig. 22) which is the most widely
distributed Cyathea 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 inulifolium 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
COStUIES TY ian ee See ee wl, cut rots Gal ak e aildin enon g ean aloe alencotncha
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. 12
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 to45mmlong. . . . . . 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.
11. 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- n-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 .. . Beet . . 136. C. persquamulifera
18. Costae rather ares scaly, small pale-fringed scales lacking: bullate scales on costules
Sctilenousi yan .. . . 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
pli2. 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 1144 cm g. 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 44144 mm
apart; lamina thin, the fertile segments deeply
crenate or more usually lobed %,4 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 | 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 bullate at base, dark-setose to-
wards apex (some setae are superficial), these
grading to pale bullate 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 BraAuse, 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. pulcherrima 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 14 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 bullate 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,
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.
Sepik
ALE
ms
——¥
SSE
aie =
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We
BESSES
Ayer
Fig. 23. Cyathea procera BRAUSE. a. Segment of a
pinnule, upper surface, < 6, b. part of costa and
fertile segment, showing scales and sori, * 6,
c. spine from base of stipe, = 4, d. flattened tip
of stipe-spine, < 13, e. scale from pinna-rachis,
<x 13 (HooGLaAND & PULLEN 5502).
123. Cyathea leucotricha Curist, Ann. Jard. Bot.
Btzg 20 (1905) 135; v. A. v. R. Handb. (1908) 18.
—Alsophila cyclodonta Curist, Ann. Jard. Bot.
Btzg 20 (1905) 137.—C. cyclodonta v. A. Vv. R.
Bull. Dép. Agr. Ind. Néerl. . 18 (1908) 1; 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. IL, vobue
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
spaced localities).
Ecol. In forest, up to c. 700 m.
widely-
124. Cyathea strigosa CuHrist, Ann. Jard. Bot.
Btzg 15 (1898) 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 rachis 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. /eucotricha of the low-
lands of Borneo.
125. Cyathea tomentosissima CopeL. Un. Cal.
Publ. Bot. 18 (1942) 219; Philip. J. Sc. 77 (1948)
1235 p> LS:
Trunk to 24% m, 16 cm g, bearing up to 40
fronds 60-105 cm long. Stipe 20-30 cm, when
dry light red-brown and warty after fall of scales,
base densely covered with shining firm twisted
brown scales to 50 by 2 mm, edges bearing ir-
regular concolorous short setae; above the base
more or less densely covered with small scales as
rachis. Rachis covered beneath with a close felt
of very small pale scales which have fiexuous
marginal hairs, also with larger scales: (a) elongate
flat pale scales to 1 mm wide having many brown
setae near apices and often pale hairs near bases,
(b) very narrow pale thin flexuous scales with
slender marginal hairs; all scales more or less
caducous with age. Longest pinnae 10-16 cm.
Longest pinnules 30 by 8 mm, lobed almost or
quite to costa; costules 2 mm apart, veins 6-7
pairs; lamina-segments very rigid, their edges
much reflexed and inrolled, entire or slightly
crenate. Sori filling cavity of lower surface of a
segment; no indusium. Scales and hairs: lower
surface of pinna-rachis and costae as main rachis
but the flat scales smaller or lacking; on costules
small brown bullate scales with long crisped
marginal hairs; hairs on upper surface of costae
pale, with a few on costules.
Type specimen: Brass 9116, Lake Habbema,
West Central New Guinea (A; dupl. at MICH,
BOVE UE):
Distr. Malaysia: New Guinea.
Ecol. At 3225 m, ‘along streams of open
grassland, in drier shrubberies, never in forest’
(BRASs).
Note. This appears to be distinct in its narrow
stipe-scales and small size of frond; also in the
close felt of small woolly scales on lower surfaces
of all rachises, though this is more or less ca-
ducous. In other characters not clearly distin-
guishable from C. atrox var. inermis.
126. Cyathea atrox C. Cur. Brittonia 2 (1937)
215; CopeL. Philip. J. Sc) 77 9@S4i) eis —
Fig. 1, 17, 24, 25.
var. atrox.
Trunk to 6 or 7 m, bearing 20-30 fronds 125-175
cm long. Stipe 30-50 cm, spiny, spines dark,
shining, 3-5 mm; scales near base of stipe to
50 by 3-6 mm, shining brown, firm but not very
rigid, edges bearing irregular concolorous or pale
hairs or setae; distal part of stipe glabrescent,
light brown when dry, warty and thorny. Lower
pinnae slightly reduced; longest 30-45 cm. Largest
pinnules commonly 65-80 by 13-15 mm, excep-
tionally to 100 by 20 mm, almost all segments
separately adnate to costa (a few only on pinnules
of upper pinnae) and constricted at base, 1-2
basal ones quite free; costules commonly 3 mm
apart, on largest fronds to 4 mm; veins 8—10(—12)
pairs, pale and raised on lower surface; lamina-
segments rigid, edges reflexed and more or less
inrolled, on distal segments entire, on largest ones
rather deeply crenate, the basal free segments often
deeply lobed. Sori nearer to costules than edge;
no indusium; paraphyses short. Scales and hairs:
pinna-rachis rather pale, warty, glabrescent,
bearing residual scales as stipe but smaller; on
lower surface of costae some elongate flat brown
scales with dark marginal setae (these mostly
caducous) and very small pale scales bearing long
pale or dark marginal hairs; on costules many
small brown bullate scales bearing crisped pale or
dark marginal hairs; similar scales sometimes on
lower surface of veins.
Type specimen: Brass 4596, Murray Pass,
Wharton Range, Papua (A; dupl. at BM, BO,
BRI).
Distr. Malaysia: E. New Guinea.
Ecol. On edges of forest or in grassland,
2800-3600 m. 4
Notes. Most collections are reported as having
many fronds, but two as having fronds in distinct
age-groups, 6 or 7 (in a whorl) in each age-group.
One of these (HOOGLAND & SCHODDE 7165) is
unusually large, with pinnules 100 mm long.
Dec. 1963] CYATHEACEAE (Holttum) 129
Fig. 24. Cyathea atrox C. CHR. var. inermis HOLTTUM in peaty grassland, Eastern Highlands, Territory
of New Guinea, 3000 m (R. D. HOOGLAND).
130
FLORA MALESIANA
fser. Il, vou
var. inermis HoLtrum, var. nov.—Fig. 24, 25.
Stipes yverrucosus, vix spinosus; paleae Sstipitis
6-10 cm longae, ad 6 mm latae; pinnulae vulgo ad
45 & 12 mm; costulae 2\% mm inter se distantes;
rhaches pinnarum subtus paleis minutis ciliatis pal-
lidis vestitae.
Stipe with numerous conical warts 14 mm high;
stipe-scales 60-100 mm long, to 6 mm wide;
pinnules commonly to 45 by 12 mm, edges strongly
reflexed and inrolled, costules 21% mm apart;
pinna-rachis often with very small pale fringed scale.
Type specimen: HoOGLAND & SCHODDE 7457,
NE. New Guinea, Western Highlands (L).
Ecol. ““Commonest species in tree-fern grass-
land’’, at 2600-2900 m. A specimen from 3500 m
(BrRAss 31013) has darker, more rigid, twisted
stipe-scales to 60 mm long.
Note. Two collections are reported as bearing
22-30 fronds spirally arranged, one (SCHODDE
1762) as having fronds in two whorls of 6 or 7.
127. Cyathea sangirensis (CHRIST) CopeL. Philip.
J. Sc. 4 (1909) Bot. 37.—Alsophila concinna BAK.
Syn. Fil. ed. 2 (1874) 459, non C. concinna (BAK.)
Jenm. 1891.—Alsophila polyphlebia Bax. J. Linn.
Soc. Bot. 15 (1876) 104, non C. polyphlebia BAK.
1883.—Alsophila sangirensis CHRIST in Warb.
Monsunia 1 (1900) 90.—Alsophila_ scaberula
Curist in K. Sch. & Laut. Fl. Deut. Schutzgeb.
Siidsee (1901) 110; v. A. v R. Handb. (1908) 35.—
Fig. 25. Cyathea atrox C. Cur. var. inermis HoLttuM on the Sugarloaf complex, Western Highlands
(R. D. HOOGLAND).
Dec. 1963]
CYATHEACEAE (Holttum)
131
C. scabriseta CopPEL. Philip. J. Sc. 9 (1914) Bot. 2;
ibid. 77 (1947) 111.—Alsophila okiana v. A. Vv. R.
Bull. Jard. Bot. Btzg II, n. 23 (1916) 4; Handb.
Suppl. (1917) 494.—Alsophila rumphiana v. A. V.
R. Philip. J. Sc. 11 (1916) Bot. 104; Handb.
Suppl. (1917) 491.—Alsophila scabriseta v. A.V. R.
Handb. Suppl. (1917) 73.—C. rumphiana MERR.
Interpr. Rumph. Herb. Amb. (1917) 63.—
Alsophila buruensis ROSENST. Med. Rijksherb. n. 31
(1917) 1.—C. okiana v. A. v. R. Bull. Jard. Bot.
Btzg II, n. 28 (1918) 14.—Alsophila scaberulipes
v. A. v. R. Nova Guinea 14 (1924) 2.—C. eminens
Donn, Pterid. (1929) 262 (new name for Alsophila
concinna BAK.); CopeL. Philip. J. Sc. 77 (1947)
116.—C. aruensis Domin, Pterid. (1929) 262
(new name for A/sophila polyphlebia BaAK.); Co-
PEL. Philip. J. Sc. 77 (1947) 115.—C. scaberula
Domin, Pterid. (1929) 263; Cope. Philip. J. Sc.
77 (1947) 109.—C. brassii Cope. J. Arn. Arb. 10
(1929) 175; Philip. J. Sc. 77 (1947) 111.—C.
buruensis Domtin, Acta Bot. Bohem. 9 (1930) 102.
— C. scaberulipes Doin, I.c. 174.
Trunk stout, to at least 8 m. Stipe long (ap-
parently to 100 cm), pale, finely warty, persistently
scaly near base; scales pale, thin but firm, finely
acuminate, to c. 30 by 2 mm, edges closely set
with dark setae. Lamina to 300 cm or more long.
Main rachis beneath pale and finely warty, some-
times with residual spreading very narrow se-
tiferous scales or hairs. Pinnae: largest about 75
cm long, lowest not greatly reduced. Pinnules:
largest 120 by 25 mm, sessile, lowest 1 or 2 seg-
ments usually free, rest of pinnule lobed almost to
costa; costules 4-414 mm apart; veins commonly
15 pairs in largest segments; segments of lamina
thin, strongly crenate (the largest crenae, where a
vein is forked, slightly notched). Sori rather near
costules; no indusium; paraphyses slender, dark at
base and pale distally, alittle longer than sporangia;
a ring of very narrow ciliate pale scales round base
of sorus. Scales and hairs: pinna-rachis minutely
warty, glabrescent or with very small pale scales
which do not bear setae; costal scales small, pale,
appressed, more or less elongate, bearing pale
short marginal hairs; costular scales pale bullate;
a few stout pale hairs rarely present near apices of
costae and costules on lower surface, these
normally lacking; upper surface of costules and
veins bearing scattered stout pale hairs, and
similar hairs on edges of segments at apices of
some veins (these latter early caducous).
Type specimen: WaARBURG 16605, Sangir (=
Sangihe) I. (P?; dupl. at B).
Distr. Malaysia: Moluccas (Sangihe I. (NE
of Celebes), Buru, Ambon, Aru Is), throughout
New Guinea, and Louisiades (Sudest I.).
Ecol. A large lowland tree-fern, in secondary
growth and in open places in forest; often near
rivers. In Sudest I. common at junction of rain-
forest with mangrove (BRASS).
Note. MerrRILL (/.c.) has suggested that
Palmifilix alba RumMeH. (Herb. Amb. 6, 63) is to
be regarded as representing this species, but the
data given by RUMPHIUS are not adequate for a
certain identification.
128. Cyathea lunulata (Forst.) Copec. Bull. Bern.
P. Bish. Mus. 59 (1929) 37; C. Cur. ibid. 177
(1943) 29.—Polypodium lunulatum Forst. Prod.
(1786) 83, n. 456.—Alsophila lunulata R. Br.
Prod. (1810) 158; Hook. Sp. Fil 1 (1844) 51;
Syn. Fil. (1868) 41, p.p.; CARR. in Seem. FI. Vit.
(1873) 333.—Alsophila vitiensis CARR. in Seem.
Fl. Vit. (1873) 170.—Alsophila naumannii KUHN,
Forschungsr. Gazelle 4 (1889) Farne 13; v. A.
v. R. Handb. Suppl. (1917) 59.—C. naumannii
Domin, Pterid. (1929) 263.—C. vitiensis DOMIN,
Acta Bot. Bohem. 9 (1930) 170.
Stipe smooth or finely warty; scales to 30 by 114
mm, pale, thin, with a few dark marginal setae.
Pinnules to at least 120 by 20 mm; costules 314-414
mm apart; segments of lamina distinctly crenate-
serrate; veins to 15 pairs. Sori exindusiate; pa-
raphyses shorter than sporangia, usually dark
when dry. Scales and hairs: pinna-rachis pale,
beneath smooth or slightly prickly, quite glabres-
cent; costae bearing stout pale hairs distally on
lower surface, scales mostly deciduous except a
few very small ones; costules bearing similar
hairs and also scales which are bullate and more
or less fringed, or on lower pinnules may be
setiferous; minute hairs sometimes visible on lower
surface of veins; hairs on upper surface of costules
rare.
Type specimen: Forster, Pacific (Géttingen;
dupl. at BM).
Distr. Solomon Is, New Hebrides, Tonga, Fiji
Samoa, in Malaysia: Bismarck Arch.
Ecol. Apparently a large tree-fern of low
country.
Note. The type specimen is labelled “In
Societatis Ins.?”; but a distribution extending to
Tahiti seems improbable.
129. Cyathea elmeri Cope-. Philip. J. Sc. 4 (1909)
Bot. 54; Fern Fl. Philip. 2 (1960) 233.—Alsophila
comosa (non WALL.) CHRIST, Ann. Jard. Bot.
Btzg 15 (1898) 80.—Alsophila latebrosa (non
WALL.) var. major CurisT, Philip. J. Sc. 2 (1907)
Bot. 183.—Alsophila elmeri CopeL. in Elmer,
Leafi. Philip. Bot. 2 (1908) 419; v. A. v. R.
Handb. Suppl. (1917) 66.—Alsophila christii Vv. A.
v. R. Handb. (1908) 42 (non Sop. 1908); Suppl.
(1917) 69, 70.—C. dimorphotricha CopEL. in Elmer,
Leafl. Philip. Bot. 5 (1913) 1681; Fern Fi. Philip. 2
(1960) 230.—Alsophila subcomosa C. Cure. Ind.
Fil. Suppl. 1 (1913) 5.—Alsophila dimorphotricha
v. A. v. R. Handb. Suppl. (1917) 61.—C. subco-
mosa Domin, Pterid. (1929) 263.—Alsophila fe-
nicis [non (CoPEL.) C. Cur.] Post. in Holthuis &
Lam, Blumea 5 (1942) 153, p.p.—C. argyrolepis
CopeL. Philip. J. Sc. 81 (1952) 17; Fern Fl. Philip.
2 (1960) 227.
Stipe 30 cm or more, the base densely scaly,
closely warty when scales have fallen; scales
25 by 114 mm, medium brown, thin but firm,
shining, acuminate, edges throughout closely set
with short concolorous setae. Main rachis, lower
surface, pale, closely warty, glabrescent. Pinnae
to at least 60 cm long. Pinnules 90-130 by 13-25
mm, sessile, lowest 2 segments of the largest free,
132
FLORA MALESIANA
[ser. II, vol. 1?
next few segments separately adnate to costa,
rest (whole of smaller pinnules) lobed nearly to
costa: costules 3—414 mm apart; veins 9-12 pairs;
lamina-segments rather thin, strongly crenate-
serrate, sinuses narrow. Sori near costules; no
indusium; paraphyses dark, not usually longer
than sporangia. Scales and hairs: pinna-rachis
beneath pale, finely warty, glabrescent, the smallest
residual scales pale with a short pale fringe, larger
ones with setiferous edges; on costae near base
narrow scales with long dark marginal setae, on
distal half more or less abundant stout pale spread-
ing hairs; on costules bullate-based broad scales,
the larger bearing dark setae near apex, the distal
ones wholly bullate and lacking setae; hairs also
more or less abundant on costules; on upper
surface of costules no hairs, or in some cases a
few.
Type specimen: Ermer 9457, Cuernos Mts,
Negros (MICH; dupl. at US, BO, F, K, P, L,
U; SYD):
Distr. Malaysia: Philippines (Mindanao, Ley-
te, Negros, Biliran), Talaud Is, and N. Celebes.
Ecol. In forest at 500-1400 m.
Notes. The specimen from Talaud Is (LAM
3319), listed as Alsophila fenicis by HOLTHUIS &
Lam, appears to lack hairs on lower surface of
costae, but its setiferous scales are like those of
C. elmeri, not of C. sangirensis.
The type specimen of C. argyrolepis, from an
exposed place on Camiguin de Mindanao, has
small pinnules (to 90 by 11 mm) and is more
scaly than normal.
C. haenkei (PR.) MeErRR. from Guam is very
nearly allied.
130. Cyathea tomentosa (BL.) ZoLL. & Mor. in
Moritzi, Syst. Verz. (1846) 108.—Chnoophora
tomentosa Bui. En Pl. Jay. (1828) 244.—Chnoo-
Phora lanuginosa JUNGH. Tijd. Nat. Gesch. &
Physiol. 8 (1841) 372; Flora 30 (1847) 522.—
Alsophila tomentosa Hook. Sp. Fil. 1 (1844) 55;
Racis. Fl. Btzg 1 (1898) 32; v. A. v. R. Handb.
(1908) 43; Suppl. (1917) 493; BACKER & PosTuH.
Varenfl. Java (1939) 30.—Alsophila lanuginosa
Pr. Epim. Bot. (1851) 29.—Alsophila crinita (non
Hook.) v. A. v. R. Handb. (1908) 40.
Trunk to 15 m, older parts showing leaf-scars
4-414 cm g. Stipe to at least 50 cm, densely scaly
throughout, spines short; scales on stipe light
brown, shining, firm, 30-45 by 1-3 mm, edges
bearing close dark setae throughout. Lamina to
250 cm long; main rachis rather closely warty
with more or less persistent smaller scales. Pinnae:
largest to 70 cm or more long. Pinnules to 110 by
20 mm, of plants in exposed places sometimes
only 60 by 10 mm wide; basal 1 or 2 segments of
lowest pinnules quite free, then 1-2 pairs sep-
arately adnate to costa, rest of pinnule lobed
almost to costa, apex acuminate; costules 3-4 mm
apart; veins 10-12 pairs; lamina-segments very
firm, edges usually reflexed when dry, crenate.
Sori nearer costule than edge; no indusium; pa-
raphyses slender, not longer than sporangia.
Scales and hairs: pinna-rachis closely warty and
more or less persistently scaly on lower surface,
scales strongly setiferous, the larger ones long and
narrow; costae densely scaly throughout, scales
near base long and narrow, flat, strongly seti-
ferous (setae long and dark), grading to smaller
long-fringed scales towards the apex; costules
densely covered with pale elongate pale-fringed
scales, the hairs of the fringes crisped and entan-
gled, at least the distal scales witha bullate base;
stout spreading pale hairs present on lower sur-
face towards apices of costae and costules and
scattered on lower surface of veins; a few also on
upper surface of costules; hairs on upper surface
of pinna-rachis always pale.
Type specimen: BLumMeE, Mt Gedeh, Java (L).
Distr. Malaysia: Java, Lesser Sunda Is (Flores).
Ecol. At 2200 m and over, in ridge forest and
in Open swampy places in gullies (abundant near
hot springs at 2200 m on Mt Gedeh, West Java).
Note. C. crinita (HOOK.) CopEL., of Ceylon and
South India, is very closely related to C. tomentosa.
It occurs at lower elevations, is less scaly generally,
the pinnules are wider, the lamina less rigid, the
hairs on scales of costules straighter and less
entangled. Specimens from Java formerly recorded
as C. crinita are C. tomentosa; if the two species
should be united, C. tomentosa is the older name.
131. Cyathea magna CopeL. Un. Cal. Publ. Bot.
18 (1942) 218; Philip. J. Sc. 77 (1947) 110, pl. 7.—
Alsophila tomentosa var. novoguineensis ROSENST.
in Fedde, Rep. 5 (1908) 34.—C. ordinata COPEL.
Philip. J. Sc. 77 (1947) 109.
Trunk to 8 m; fronds 8-14, spirally arranged.
Stipe 40-90 cm, medium brown when dry, near
base more or less spiny, spines 3-5 mm long,
finely warty between the spines; large scales abun-
dant and persistent near base only, pale or in
part medium brown, firm and shining, to 50
by 114(-2) mm, edges closely set throughout
with short dark setae; rest of stipe + persistently
covered with smaller pale to brown scales, the
larger freely dark-setiferous, the smallest fringed
with pale hairs. Lamina to nearly 300 cm long;
main rachis on lower surface light brown when
dry, rather closely warty and scaly as upper part
of stipe. Pinnae: largest 90 cm long. Pinnules
mostly rather stiffly spreading almost at right
angles to pinna-rachis, largest 100-140 by 14-20
mm, sessile, acuminate, lobed almost to the costa
(lowest lobe sometimes just free but adnate by
its lamina); costules of large fronds 4 mm apart,
of smaller ones 3 mm; veins 10-12 pairs, pale and
prominent beneath, not raised above; segments
of lamina thick, stiffly spreading when dry, edges
often rather much reflexed on drying, finely
crenate to almost entire. Sori nearer costule than
edge; indusium lacking; paraphyses not longer than
sporangia, no scales round base of sorus. Scales
and hairs: pinna-rachis beneath pale, scaly as
main rachis; costae beneath densely scaly, scales
near base like the larger ones of pinna-rachis, flat
and elongate, strongly setiferous, smaller and
narrow scales bearing long flexuous dark marginal
setae present almost throughout (smallest have pale
CYATHEACEAE (Holttum)
133
Dec. 1963]
marginal hairs); costules densely scaly, scales
near base narrow flat and bearing long dark setae,
rest light brown with + bullate base and long
pale flexuous marginal hairs; erect hairs present
distally on costules and on lower surface of
veins; upper surface of costules not hairy; hairs on
upper surface of pinna-rachis dark except towards
apex of frond.
Type specimen: Brass 11278, Bele R., 18 km
NE of Lake Habbema, 2250 m, W. New Guinea
(A; dupl. at MICH, BO, K, BM, L, UC).
Distr. Malaysia: New Guinea.
Ecol. In open places in forest or in grassland,
1700-2750 m.
Note. This species is closely allied to C.
tomentosa of Java, differing in narrower and
firmer stipe scales, more rigid pinnules, and a
greater preponderance of small scales bearing
long dark setae, these giving a darker aspect; the
hairs on upper surface of pinna-rachises are also
dark.
132. Cyathea pilulifera CopeL. Un. Cal. Publ.
Bot. 18 (1942) 219; Philip. J. Sc. 77 (1947) 112,
folk, Ye
Stipe 50 cm, basal part copiously spiny, spines
dark, to 5 mm; scales pale, to 25 by 2 mm, edges
bearing sparse dark setae or concolorous hairs;
main rachis spiny, glabrescent; pinna-rachis
sparsely spiny or warty. Lamina c. 100 cm long;
longest pinnae 45 cm. Pinnules to 110 by 20 mm,
lowest segment just free, rest lobed almost to
costa; costules 4 mm apart, distinctly curved;
lamina-segments firm, close, edges crenate; veins
12-14 pairs, mostly forked, lowest basiscopic
vein sometimes from costa. Sori exindusiate;
paraphyses not longer than sporangia; scales
present round base of receptacle. Scales and
hairs on lower surface of costae, costules and
veins: flat pale scales with many long dark setae
present near bases of costae and smaller ones dis-
tally and on costules; pale bullate scales with
some setae present on distal part of costae and on
costules and veins; stout pale hairs abundant on
distal part of costae and on costules and veins;
upper surface of costules and veins glabrous.
Type specimen: BrAss 11492, 18 km NE. of Lake
Habbema, 2200 m, W. New Guinea (A; dupl. at
BM, BO, MICH).
Distr. Malaysia: Moluccas (Ceram),
Guinea, and Louisiades.
Ecol. At 1250-2750 m; at some lower elevations
reported in forest with tall trunk, at higher ele-
vations in open places (the type in young secondary
growth on old garden land, with short trunk).
New
133. Cyathea curranii Cope. Philip. J. Sc. 3
(1909) Bot. 356; ibid. 4 (1910) Bot. 52; Fern
Fl. Philip. 2 (1960) 231.— Alsophila curranii
Cy Grins Iie lel Soysylk We (MNS) Se awe ANS we TRE
Handb. Suppl. (1917) 70.
Trunk 3 m, 20 cm @; leaf-scars 3 cm o. Stipe
35 cm, pale, warty, bearing light brown shining
scales to 60 by 4 mm, their edges bearing rather
few concolorous setae. Lamina 100 cm long. Lower
pinnae long-stalked, reduced and deflexed, largest
35 cm. Pinnules to 90 by 13 mm, lowest segments
almost free; costules 3 mm apart; segments of
lamina rigid, nearly entire, glaucous beneath;
veins 8-10 pairs. Sori exindusiate, paraphyses
present. Scales and hairs: pinna-rachis beneath
closely warty (warts dark), with a few residual
narrow light brown scales, their edges bearing
concolorous setae; costae glabrescent; costules
and veins beneath bearing scattered pale bullate
scales and many very short hairs (probably bases
of fallen scales); long hairs absent.
Type specimen: CURRAN & MerritT FB 7925,
Mt Banajao, 2000 m, Luzon (MICH; dupl. at
PSUs):
Distr. Malaysia: Philippines (Luzon).
Note. Specimens seen are in poor condition
and do not include largest pinnae. The nearest
relationship seems to be to C. mertensiana (KUNZE)
CopeL. of the Bonin Is and C. aeneifolia (v. A.
v. R.) Domin, and allied species of New Guinea
and the Pacific.
134. Cyathea aeneifolia (v. A. v. R.) Domin, Acta
Bot. Bohem. 9 (1930) 174; CopeL. Philip. J. Sc.
77 (1947) 113.—Alsophila aeneifolia v. A. v. R.
Nova Guinea 14 (1924) 3.—Alsophila aeneifolia
var. subglauca v. A. Vv. R. l.c. 4.—C. curvipinnula
C. Cur. Brittonia 2 (1937) 276; CopeL. Philip.
J. Sc. 77 (1947) 116.
Trunk to 414 m. Fronds 9-12, 175-250 cm long.
Stipe 50-60 cm, strongly spiny and scaly; spines
dark and shining, to 6 mm; largest scales 30-60
by 3-5 mm, thin but firm and flat when young,
rather pale shining brown, edges bearing scattered
concolorous hairs or dark setae; rachis short-
spiny or warty, bearing similar but smaller scales
which are mostly caducous. Lower pinnae slightly
reduced and deflexed, largest pinnae 30-60 cm.
Largest pinnules 70-120 by 15-25 mm, sessile,
lobed nearly to costa, a few lower segments almost
or quite free; costules 3-414 mm apart; lamina-
segments very firm, almost entire, glaucous be-
neath; veins 10-12 pairs. Sori medial on veins, no
indusium; paraphyses numerous, as long as spo-
rangia; some narrow pale scales surrounding
base of receptacle. Scales and hairs: pinna-rachis
bearing -- caducous pale brown scales to 151
mm, also small pale fringed scales; throughout
lower surface of costae small fringed pale scales,
with some elongate setiferous scales near base and
bullate ones towards apex; on costules pale bullate
hair-tipped scales, often with crisped marginal
hairs or sometimes setae; on veins beneath small
pale bullate scales more or less abundant, with
lax hairs like tips of scales at apices of veins; no
hairs on upper surface of costules and veins.
Type specimen: Lam 1751, W. New Guinea, near
foot of Doormantop, in sunny ravine, 3240 m
(BO; dupl. at US, L, U).
Distr. Malaysia: New Guinea.
Ecol. On forest edges or in grass-land, 2840—
3240 m. LAM 1805, distinguished as var. subglauca
v. A. v. R., was collected in a shady valley at 3200
m, and has larger thinner fronds than the type but
134
FLORA MALESIANA
[ser. DL, youl
is otherwise similar. This species is nearly allied to
C. pilulifera CopEL., but the latter has abundant
long hairs on costae, costules and veins, and has
been found at lower altitudes. Perhaps the two
should be united.
var. melanacantha (CoPEL.) HOLTTUM, var. nov.—
C. melanacantha CopEL. Un. Cal. Publ. Bot. 18
(1942) 219; Philip. J. Sc. 77 (1948) 114, pl. 10.
Stipe-scales to 15 by 2 mm, dark brown.
Type specimen: Brass 9311, Lake Habbema,
W. New Guinea (A; dupl. at BM, BO, UC, L,
MICH).
Ecol. Occasional in forest-edge, at 3225 m.
var. macrophylla HOLTTUM, var. nov.
A typo speciei differt: frondibus ad 425 cm longis,
stipitibus 200 cm longis inclusis; paleis stipitis ad
50 < 4 mm; pinnis maximis 70 cm longis, pinnulis
maximis 135 * 21 mm; costulis 5 mm inter se
distantibus.
Type specimen: HOOGLAND & SCHODDE 7209,
Western Highlands, NE. New Guinea (L).
Ecol. In cloud-forest at 2900 m.
135. Cyathea tenggerensis (ROSENST.) DOMIN, Ac-
ta Bot. Bohem. 9 (1930) 165.—Alsophila haenkei
var. angustata HAssk. in Hook. J. Bot. Kew Misc.
7 (1855) 326.—Alsophila glauca (BL.) HOOK. p.p.:
v. A. v. R. Handb. (1908) 41; Suppl. (1917) 68;
BACKER & PostH. Varenfl. Java (1939) 28.—
Alsophila tenggerensis ROSENST. Med. Rijksherb.
n. 31 (1917) 1.
Stipe to at least 60 cm, warty, persistently scaly
near base; stipe-scales to 45 by 214-314 mm,
shining pale brown, firm, setiferous. Pinnae to at
least 50 cm long. Pinnules to 110 by 16 mm,
sessile, lobed almost to costa with | or 2 basal
segments almost free; costules 214-3 mm apart;
lamina-segments firm, edges -+ reflexed when dry,
lower surface probably glaucous; veins 10—12 pairs.
Sori exindusiate, nearer to costule than to edge,
pale paraphyses present. Scales and hairs: pinna-
rachis closely warty beneath, with some residual
pale scales to 7 by 14 mm, setiferous; costae with
similar shorter elongate scales near base, for the
most part covered with rather large pale brown
bullate scales which have a few concolorous
marginal hairs; near apex of costae a few long pale
hairs; costules bearing similar bullate scales and
occasionally hairs; no long hairs on veins but very
short appressed hairs (bases of former scales ?)
often conspicuous; hairs on upper surface of
costules rare.
Type specimen: HAsskKARL, Java (L?; not seen).
Cited by RosENsTOCK: ZOLLINGER 2541, Mt
Tengger, E. Java (L; dupl. at P).
Distr. Malaysia: East Java, Lesser Sunda Is
(Flores), South Celebes.
Ecol. In open places at 1500-2300 m; locally
abundant on Mt Tengger.
Note. This species was established by ROSEN-
STOCK by reference to the description of Also-
Phila haenkei var. angustata HASsK., without further
description; HASSKARL’s specimen, for which he
cites no locality, is therefore the type. The above
description is largely based on several later col-
lections from Mt Tengger.
136. Cyathea persquamulifera (v. A. v. R.) Domin,
Acta Bot. Bohem. 9 (1930) 146.—C. contaminans
var. persquamulifera v. A. v. R. Bull. Jard. Bot.
Btzg II, n.28 (1918) 13.—Alsophila persquamulifera
v. A. v. R. ibid. III, 2 (1920) 130.
Stipe scaly throughout, no long spines; basal
scales to 30 by 1 mm, medium brown, shining,
strongly setiferous (on a young frond). Main
rachis closely warty beneath. Pinnae to at least 60
cm long, lower ones rather long-stalked. Pinnules
to 135 by 25 mm, | or 2 basal segments free or
contacted at base, rest of pinnule cut nearly to
costa, sessile, caudate-acuminate; costules 314-4
mm apart; segments firm, edges often + reflexed
when dry, crenate; veins to at least 12 pairs, lowest
from costa or base of costule. Sori nearer costule
than margin; no indusium, thin paraphyses not
longer than sporangia. Scales and hairs: pinna-
rachis beneath bearing very small fringed pale
scales and copious narrow light brown strongly
setiferous ones up to 10 mm long; costae at base
as pinna-rachis, the larger scales 5-6 mm long,
grading to very narrow setiferous scales and some
bullate hair-pointed scales, also many very long
(2 mm) stout brown hairs except near base;
costules bearing copious similar hairs, sometimes
narrow pale scales 2-3 mm long with marginal
hairs or setae, always hair-pointed pale bullate
scales; veins with a few stout erect hairs on lower
surface; upper surface of costules and veins gla-
brous.
Type specimen: BUNNEMEIJER 961, Mt Talamau,
2300 m, Sumatra (BO; dupl. at K, L).
Distr. Malaysia: Central Sumatra, throughout
Java.
Ecol. On mountains at 1500-2500 m.
137. Cyathea sarasinorum Hoittum, Kew Bull.
16 (1962) 61.—Alsophila contaminans var. longe-
paleata Curist, Ann. Jard. Bot. Btzg 19 (1904) 42.
—Alsophila glauca var. longepaleata v. A. Vv. R.
Handb. (1908) 41; Suppl. (1917) 69.
Main rachis pale beneath, with dark shining
warts bearing dark shining setiferous scales to
15 by 1 mm. Pinnae to 67 cm long. Pinnules to
90 by 18 mm, lowest 1 or 2 lamina-segments
constricted at base, rest of pinnule lobed almost to
costa, apex rather shortly acuminate; costules 4
mm apart; segments of lamina thin, oblique, their
rounded ends curved forwards, edges crenulate;
veins 9-10 pairs. Sori near costules, exindusiate.
Scales on costae beneath near base very narrow,
dark shining brown, with many concolorous setae,
smaller distally; pale spreading hairs also present
towards apex of costae and on costules, on lower
surface; scales on costules dark narrow, setiferous,
also pale bullate scales bearing setae near their
tips.
"Type specimen: F. & P. SARASIN 2105, Si-
baronga-Riicken, Central Celebes (BAS).
Distr. Malaysia: Central Celebes (one collec-
tion).
Dec. 1963 ]
CYATHEACEAE (Holttum)
ge
138. Cyathea angiensis (GepP) DomIN, Acta Bot.
Bohem. 9 (1930) 90.—Alsophila angiensis GEPP
in Gibbs, Arfak (1917) 69.
Like C. contaminans (WALL.) COPEL., but rather
small; pinnules 100-150 by 16-20 mm; costules
314 mm apart, veins flat or impressed on lower
surface; lamina-segments very firm, edges almost
entire to crenate, glaucous beneath; more or less
abundant pale flat setiferous scales present on
lower surface of costae and costules; long pale
spreading hairs more or less abundant towards
apex of costae and on costules and veins on lower
surface, rarely on upper surface of costules; pa-
raphyses pale, longer than sporangia.
Type specimen: Gipps 5968, Angi Lakes, 7000 ft,
W. New Guinea (BM; K, P, BO).
Distr. Malaysia: Moluccas
Guinea.
Ecol. Mountains, 600-2200 m.
Note. A specimen from Sepik District, Ter-
ritory of New Guinea (DARBYSHIRE & HooG-
LAND 8191) bears the following information:
“fronds immediately fully deciduous, the leaf-
scars in distinct orthostiches (8) ...12 fronds
in 3 whorls of 4, the outer whorl old, the middle
whorl young fertile, the upper whorl not yet fully
expanded, within each whorl the fronds very
closely of the same age’’. DARBYSHIRE 384 has
similar information, but fronds in whorls of 5.
If I have correctly identified these specimens, the
following characters distinguish this species from
C. contaminans: fronds in whorls of 4 or 5, old
fronds immediately and fully caducous. The re-
maining specimens referred to C. angiensis do not
bear such information.
See also under 140. C. contaminans.
(Buru), New
139. Cyathea verrucosa HoLttuM, Kew Bull. 16
(1962) 63.
Stipe rather pale, closely warty throughout,
persistently scaly near base only; scales to 40 by
3 mm, thin, pale, edges strongly dark-setiferous;
main rachis, lower surface, glabrescent, closely
warty (warts to 1 mm high). Pinnae to at least 60
cm long, lower ones with stalks 6 cm long. Pinnules
to 110 by 20 mn, sessile or the lowest stalked,
caudate-acuminate, basal 1-2 segments free or
nearly so, rest cut almost to costa; costules 314-4
mm apart; lamina-segments rigid, edges minutely
crenate, sinuses narrow; veins to 12 pairs. Sori
at about 14 distance from costule to edge, marked
by a depression on the upper surface, exindusiate;
paraphyses pale, as long as sporangia, no scales
round base of sorus. Scales and hairs: pinna-
rachis beneath rather closely warty, bearing
scattered pale setiferous scales to 10 by 1144 mm
and very small pale setiferous or short-fringed
scales; costae at first bearing many narrow pale
setiferous scales to 2 mm long, these mostly
caducous, also much shorter pale scales which are
setiferous or the smallest pale-fringed, not bullate,
distal half of costae bearing copious long pale
spreading hairs with few scales; costules at first
bearing many narrow pale setiferous scales to
2 mm long, also flat ovate-acute setiferous or
fringed scales, and long pale spreading hairs;
veins beneath bearing stout erect hairs (2 or 3 on
a vein) and very short hairs which appear to be
bases of fallen scales; upper surface of costules
and veins glabrous.
Type specimen: MATTHEW s.n., 3.2.1912, Mt
Merapi, 5000 ft, Sumatra (K).
Distr. Malaysia: Central Sumatra (two col-
lections).
Ecol. In open places in forest at 1600-1900 m.
Note. Closely related to C. contaminans but
much more densely scaly, the stipe closely warty
instead of rather sparsely spiny.
140. Cyathea contaminans (WALL. ex Hook.)
CopeL. Philip. J. Sc. 4 (1909) Bot. 60; v. A. v. R.
Bull. Jard. Bot. Btzg II, n. 28 (1918) 13; Copet.
Philip. J. Sc. 77 (1949) 115; Hottrum, Rev. FI.
Mal. 2 (1954) 119; CopeL. Fern FI. Philip. 2
(1960) 230.—Polypodium contaminans WALL. Cat.
(1828) n. 320, nomen.—Chnoophora glauca BL.
En. Pl. Jav. (1828) 243 (non C. glauca Bory, 1804).
—Alsophila glauca J. Sm. J. Bot. 3 (1841) 419;
Bepp. Handb. (1883) 12; v. A. v. R. Handb.
(1908) 41 (incl. var. celebica, var. squamulata,
var. densa, var. setulosa, and var. microloba);
Koorp.-ScHum. Syst. Verz. 1, 2 (1912) 5; Ro-
SENST. Hedwigia 56 (1915) 349, incl. var. tricho-
carpa ROSENST.; Vv. A. v. R. Handb. Suppl. (1917)
69 (incl. var. squamulosa); Bull. Jard. Bot. Btzg
III, 2 (1920) 129; BACKER & PostH. Varenfl. Java
(1939) 28.—Alsophila contaminans WALL. ex
Hook. Sp. Fil. 1 (1844) 52, t. 18, f. 2; HaAssxk. in
Hook. J. Bot. Kew Misc. 7 (1855) 323 (incl. var.
robusta, var. squamulata, var. densa, var. microloba
and var. setulosa HAssk.); BEDD. Ferns Br. Ind.
(1865) pl. 85; Hook. Syn. Fil. (1866) 41; Scott,
Trans. Linn. Soc. 30 (1874) 35; CuHrist, Verh.
Nat. Ges. Basel 11 (1895) 199, incl. var. celebica
Curist; Farnkr. Erde (1897) 327; Ann. Jard. Bot.
Btzg 15 (1898) 79, pl. XIII f. 2; Diets in E. & P.
Pfi. Fam. 1, 4 (1899) 136, p.p.—Alsophila acuta
PresL, Abh. k. BOhm. Ges. Wiss. V, 5 (1848) 343.
—Alsophila smithiana PREsL, I.c. 342.—Alsophila
clementis Core. Philip. J. Sc. 1, Suppl. 2 (1906)
Bot. 143.—C. clementis Copet. ibid. 4 (1909)
Bot. 59; Fern Fl. Philip. 2 (1960) 230.—Fig. 3,
6, 22, 26.
Trunk often very tall and much thickened by
adventitious roots at base, only when old showing
leaf-scars in upper part. Stipe to 100 cm long,
glaucous, purplish towards the base, usually
strongly spiny, at first scaly throughout, per-
sistently so near the base; scales of all sizes up to
45 by 3 mm, pale brown, very thin, edges bearing
close dark short setae. Main rachis pale, spiny, at
first scaly as stipe, later + glabrescent. Pinnae:
lowest somewhat reduced and with stalks to 10 cm
long, largest 60 cm. Pinnules to c. 150 by 30 mm,
often smaller, lowest distinctly stalked, largest
with 1-2 pairs of basal segments more or less free,
rest of pinnule lobed almost to the costa; costules
commonly 4414 mm apart, rarely to 5 mm;
segments of lamina firm, glaucous beneath,
edges + crenate-serrate; veins commonly 12 pairs.
136
FLORA MALESIANA
{ser. Il, volte
Fig. 26. Cyathea contaminans (WALL.) CoPEL. in Tjibodas mountain garden (VAN STEENIS).
Sori near costules, lacking indusia; paraphyses
pale, not longer than sporangia. Scales and hairs:
pinna-rachis more or less spiny, pale, glabrescent;
lower surface of costae at first bearing scattered
pale setiferous scales to 3 by 144 mm, shorter
distally, these in most cases early caducous leaving
the costae glabrous; costular scales small, + ovate,
pale-fringed, mostly caducous, not bullate; a few
hairs normally present towards apex of pinnules
on both costae and costules, their abundance
varying much; in a few localities (notably Mt
Kinabalu, also in New Guinea) stout erect hairs,
in variable number, may be present on lower
surface of veins.
Type specimen: WALLICH 320, Penang (K).
Distr. Throughout Malaysia, in Peninsular
India as far N as Mergui. Specimens so named
from Hong Kong are C. /epifera (J. SM.) COPEL.
Ecol. In clearings and open places in forest,
especially near streams, 200 to 1600 m, often very
abundant.
Notes. This is the most widespread species
of Cyathea in Malaysia, having no close allies at
low elevations in Western Malaysia. In New
Guinea there are several allied species, of which
C. angiensis (GEPP) DOMIN appears to be nearest;
it may perhaps better be united to C. contaminans.
The several varieties of C. contaminans which have
been described appear mostly to be due to the
influence of environmental factors, or to be
Dec. 1963]
CYATHEACEAE (Holttum)
137
young, and therefore unusually scaly, at the time
of collection.
I have only seen one specimen of this species
from NE. India; it was collected by Scott, who
noted (/.c.) that typical A/sophila contaminans
occurred there only at low altitudes. He distin-
guished var. brunoniana (A. brunoniana HOoK.),
which is common in Sikkim at higher altitudes and
has smooth (not spiny) stipes and copiously scaly
costae and costules. I prefer to regard this as a
distinct species, which must be called Cyathea
brunoniana (HooK.) CLARKE & BAK., though this
name was actually used by CLARKE & BAKER for
a quite different species. I hope to publish else-
where descriptions of this and other species of the
mainland of Asia.
141. Cyathea lepifera (J. SM.) Cope. Philip. J.
Sc. 4 (1909) Bot. 40; ibid. 56 (1935) 98, pl. 3,
fig. 4-7; Fern FI. Philip. 2 (1960) 228.—Alsophila
lepifera J. SM. ex Hook. Sp. Fil. 1 (1844) 54;
CurisT, Bull. Herb. Boiss. 6 (1898) 137, incl. var.
congesta CHRIST; Vv. A. v. R. Handb. (1908) 39;
Suppl. (1917) 65.—Alsophila pustulosa CHRIST,
Bull. Herb. Boiss. Il, 1 (1901) 1019; Naxkat,
Bot. Mag. Tokyo 41 (1927) 73; Ito, Fil. Jap. Ill.
(1944) pl. 453.—Alsophila calocoma CurIsT,
Philip. J. Sc. 2 (1907) Bot. 182; v. A. v. R. Handb.
(1908) 789.—C. pustulosa CopeL. Philip. J. Sc. 4
(1909) Bot. 51.—C. calocoma CopPEL. /.c. 53; Fern
Fl. Philip. 2 (1960) 229.—C. umbrosa COPEL.
Philip. J. Sc. 56 (1935) 98, pl. 3, fig. 1-3.—(?)
C. pteridioides Cope. l.c. 98, pl. 4; Fern FI.
Philip. 2 (1960) 229.
Stipe 16 cm or more, almost wholly covered
with scales, not spiny (warty when scales have
fallen); scales pale, thin, to 40 by 24 mm wide at
base, apical part narrowly acuminate, edges
throughout closely setiferous, narrow apical part
sometimes entirely brown; smaller scales on stipe
very narrow (under 14 mm wide), brown, with
concolorous marginal setae. Rachis and pinna-
rachis closely conspicuously warty, pale, more or
less completely glabrescent, warts darker, to nearly
1 mm high. Pinnae: lowest somewhat reduced;
largest to at least 80 cm long. Pinnules 100-150 by
15-22 mm, caudate-acuminate, sessile, largest with
a few free or separately adnate basal segments,
the rest lobed almost to the costa; costules 3-314
mm apart; veins 12-14 pairs; lamina-segments
firm, entire or nearly so, glaucous beneath. Sori
near costules; no indusia; paraphyses pale, longer
than sporangia, some in form of narrow scales.
Scales and hairs: pinna-rachis bearing some
persistent appressed small pale fringed scales and
occasionally longer spreading setiferous scales;
costae beneath more or less densely scaly, basal
scales elongate, narrow, pale with dark setae,
distal ones smaller with pale marginal hairs,
all flat, also some very small pale fringed scales
throughout; at least the apical part of each costa
also bearing stout pale hairs beneath; costules
beneath bearing pale flat ovate to elongate scales
bearing short pale marginal hairs, also more or
less abundant stout spreading pale hairs; upper
surface of costules lacking hairs.
Type specimen: CuMING 180, Luzon (K; dupl.
at BM, F, A, BO).
Distr. Ryukyu Is, Formosa, Kwangtung, Hong
Kong, in Malaysia: Philippines (Luzon, Panay,
Mindoro, Babuyan).
Ecol. In the Philippines a mountain species,
but herbarium labels bear little ecological in-
formation. I have been unable to find the type
specimen of C. pteridioides CoPeL.; it is not in
COPELAND’s herbarium.
142. Cyathea atrospinosa HoL_trum, Kew Bull.
16 (1962) 52.
Trunk 6 m; fronds 350 cm long, in whorls of
4, 5 or 6, usually two whorls green at one time.
Stipe to 125 cm long, lower part armed with dark
spines to 8 mm long, also densely covered with
scales which are up to 30 by 5 mm, thin, pale or
pale brown, their edges bearing concolorous short
hairs or dark setae. Rachis pale and glabrescent
on lower surface, with scattered small thorns.
Pinnae to 80 cm long. Pinnules to 120 by 25-30
mm, lowest on stalks up to 5 mm, lobed almost to
the costa with the 1—2 lowest segments more or
less free; costules 314-4 mm apart; lamina-seg-
ments crenate, apices rounded, sinuses between
them about 1 mm wide; veins 13-17 pairs. Sori
exindusiate; paraphyses numerous, longer than
sporangia. Scales and hairs: pinna-rachises pale
beneath, glabrescent or with scattered setiferous
scales; scales near base of costae shining brown,
ovate, flat, long-setiferous, towards apices paler
fringed scales; costules bearing pale brown flat
fringed scales, and a few similar smaller ones also
on lower surface of veins.
Type specimen: HOoOGLAND & PULLEN 6090,
Western Highlands District, 2650 m, NE. New
Guinea (K).
Distr. Malaysia: E. New Guinea.
Ecol. Inmountain forest, on limestone (always? )
at c. 2400-2850 m.
143. Cyathea fugax v. A. v. R. Bull. Jard. Bot.
Btzg Il, m. 7 (1912) 8: Handb. Suppl. (1917) 34;
CopEL. Philip. J. Sc. 77 (1947) 116.
Trunk to 10 m; fronds up to 300 cm long;
leaf-scars 4 cm 9, in alternate whorls of 5 (BRAss
25585, Normanby I.). Stipe to at least 40 cm,
spiny and rather densely scaly for most of its
length; spines 2 mm or more long; scales pale,
thin, to 25 by 2 mm, bearing many dark marginal
setae near apices. Pinnae to 70 cm or more long.
Pinnules to 145 by 35 mm, lowest 1—2 segments
free, then some separately adnate, rest of pinnule
cut nearly to costa, apex caudate-acuminate;
costules 414 mm apart; lamina-segments rigid,
glaucous beneath, little over 3 mm wide, almost
entire; veins to 20 pairs, basiscopic basal vein
from costa. Sori nearer costules than edge, ex-
indusiate, with a ring of narrow fringed scales
round base of receptacle; long paraphyses also
present. Scales and hairs: pinna-rachis pale, bear-
ing scattered short thorns, glabrescent; on costae
138
FLORA MALESIANA
[ser. II, vol. 7
near base pale scales (apex dark or not) bearing
dark flexuous setae, grading to smaller pale fringed
scales without setae (or with a few) on costules
and on veins; small scales on veins with short
marginal hairs; very small hairs present on lower
surface between veins; no hairs on upper surface
of costules.
Type specimen: CoPpLAND KING 215, Papua
(BO; dupl. at MICH, SYD).
Distr. Malaysia: E. New Guinea and adjacent
islands.
Ecol. In wet ground, secondary forest and
open places in forest in low country up to 1400 m.
Note. Details of trunk and stipe are taken
from specimens other than type. Under type
number at Sydney and Brisbane are also specimens
of C. contaminans (WALL. ex HooKk.) COpPEL.,
which differ notably in scales from the type.
3b. Subsection Fourniera
(BOMMER) HOLTTUM, stat. nov.—Fourniera BOMMER, Bull. Soc. Bot. Fr. 20 (1873)
x1x.—Fig. 27.
Type species: Cyathea novae-caledoniae (METT.) COPEL.
Distr. The centre of distribution is New Guinea, with eastward and south extension to Samoa, New
Caledonia and NE. Australia, westward and north to W. Java, Celebes, North Borneo (also Pulau Tio-
man) and Philippines.
Taxon. A distinction between species with long stipes and those with short stipes (the latter having
gradually reduced lower pinnae) appears to be valid, but some collections do not include the stipe and
so are not easy to place. More field study is needed in Eastern New Guinea.
Ecol. These appear all to be forest species.
KEY TO THE SPECIES
1. Pinnules 30-40 mm long; lower pinnae gradually reduced.
2. Tertiary leaflets on stalks almost 1 mm long. Pinnae to 25 cm long. Bullate scales present on costules.
144. C. carrii
2. Tertiary leaflets almost sessile. Pinnae to 45 cm long. No bullate scales on costules.
145. C. womersleyi
1. Pinnules commonly at least 70 mm long, in most cases much longer.
3. Lower pinnae gradually reduced,
lowest 11
cm long near base of stipe.
146. C. auriculifera
3. Lower pinnae not thus reduced (sometimes 1-2 pairs of isolated small pinnae at base of stipe).
4. Bullate scales present on costae and costules.
5. Pinnules to 110 by 22 mm. Soral scales small, not covering sorus at maturity.
147. C. teysmannii
5. Pinnules to about 65 by 16 mm. Soral scales broad, covering sorus at maturity.
4. Bullate scales lacking.
148. C. aciculosa
6. Veins and lower surface of lamina bearing ee fringed ee scales which cover lower
surface of leaflets
149. C. celebica
6. Veins bearing at most small separate scales on lower surface.
7. Scales absent from lower surface of veins .
150. C. tripinnata
7. Scales present on lower surface of veins, not long- fringed 7 nor r covering lower surface of leaflets.
144. Cyathea carrii HoLttum, Kew Bull. 16 (1962)
53.—Fig. 27.
Stipe c. 7 cm, covered with a felt of very small
dull scales, the larger ones setiferous, surface
where exposed dark and smooth; larger scales
near base to 30 by hardly 1 mm, edges bearing
short dark setae, apex a long dark seta; lower
surface of rachis dark brown, smooth or finely war-
ty, glabrescent or with sparse minute dull scales.
Lowest pinnae c. 3 cm long, succeeding ones
gradually longer, up to 25 cm long. Pinnules to
32 by 7 mm, fully pinnate; tertiary leaflets to
c. 12 pairs, distinctly stalked (stalks to almost
1 mm long and 2 mm apart), to 4 by a little more
than 1 mm, edges sinuous, lowest sometimes
151. C. macrophylla
deeply lobed at the base; veins 4-5 pairs. Sori up
to 6 on each tertiary leaflet, covered to ripeness
by broad pale lacerate overlapping scales which
give the appearance of a complete indusium.
Scales and hairs: pinna-rachis beneath dark,
bearing many very small pale dull scales, some
small bullate scales and scattered long narrow
pale or partly dark closely setiferous scales;
scales of pinnule-rachis (costa) small, brown,
shining, mostly bullate and ending in a seta, some
nearly flat and slightly elongate, rarely with lat-
eral setae; scales of midribs of leaflets shining
dark brown, bullate, acuminate, similar scales
sometimes also on lower surface of veins; upper
surface of tertiary leaflets glabrous.
Dec. 1963] ,
CYATHEACEAE (Holttum)
a
Fig. 27. Cyathea carrii HOLTTUM.a. Single pinnule,
upper surface, x 2, b. part of pinnule, lower
surface, showing scales and sori, 6, c. sorus,
showing covering of overlapping scales, x 20,
d. scales from costule, < 30 (CARR 13526).
Type specimen: CARR 13526, in forest, Boridi,
5000 ft, Papua (K; dupl. at BM, L).
Distr. Malaysia: E. New Guinea (one col-
lection).
145. Cyathea womersleyi Ho_ttrum, Kew Bull.
16 (1962) 63.
Trunk to 8 m, fronds to 10, suberect, to 300 cm
long, with pinnae gradually reduced to the base.
Stipe 8-20 cm, sparsely spiny (spines to 214 mm),
densely and persistently covered with scales which
are pale, rather dull and soft, with a dark red
seta at apex and no others, 10-20 by less than 1
mm, smaller ones 4-5 by !/; mm; main rachis
persistently scaly beneath, smaller scales all with
setiform apex, some elongate ones with pale
edges bearing dark setae. Pinnae: lowest 8 cm long,
longest 45 cm. Pinnules to 40 by 10 mm, almost
fully pinnate; costules (midribs of tertiary leaf-
lets) 2 mm apart; tertiary leaflets mostly 4-5 mm
long, hardly stalked, slightly oblique, entire,
apex rounded, base unequal, veins 6-7 pairs,
lowest leaflets on largest pinnules to 7 mm long,
bearing at base 1 or 2 pairs of 4th order leaflets
which are round and entire, 1 mm long. Sori
covered by overlapping fringed scales. Scales and
hairs: lower surface of pinna-rachis densely scaly,
some scales dark,shining,elongate with pale sparsely
fringed or setiferous edges, some smaller, light
brown and sub-bullate with setiform apex; near
base of costae dark shining ovate to elongate
scales with pale edges bearing some dark setae,
then light brown nearly flat roundish scales with
short setiform apex to pale hair-tip; on costules
a few scales which usually lack setae but may
bear marginal hairs; small scales sometimes on
lower surface of veins.
Type specimen: WoOMERSLEY & MILLAR NGF
8470, Skindewai, 5400 ft, Morobe District, NE.
New Guinea (K; dupl. at L, BO, A, SYD).
Distr. Malaysia: E. New Guinea (3 collec-
tions).
Ecol. At 1700-2400 m; “‘common through the
rain-forest’’ (BRASS), in mixed Nothofagus forest
(SCHODDE).
146. Cyathea auriculifera Cope. Philip. J. Sc. 6
(1911) Bot. 364; ibid. 77 (1947) 107.
Trunk to 3 m; fronds 220 cm long, lower pinnae
gradually reduced. Stipe 10 cm, copiously spiny
(spines to 3 mm long), also densely covered with
pale scales 10-55 mm long, larger ones twisted
and rather straight, edges closely set with short
dark setae; main rachis beneath rather closely
spiny throughout, covered with a dark brown
felt of very small mostly setiferous scales and at
first with elongate pale scales attached to the
thorns. Pinnae: lowest 11 cm long, largest 54 cm.
Pinnules slightly overlapping, to 70 by 15 mm,
fully pinnate, bases of tertiary leaflets 3 mm
apart; rertiary leaflets mostly stalked, largest 9 by
214 mm, several pairs near base of pinnule having
1 or 2 pairs of free quaternary leaflets, acroscopic
basal tertiary leaflets deflexed across pinna-rachis;
veins to 9 pairs, those in quaternary leaflets pin-
nately branched, rest mostly forked. Sori nearer
to costules than to edge, covered with overlapping
fringed scales until nearly ripe, fringe of these
scales longer than on costal scales. Scales and
hairs: pinna-rachis scaly beneath as main rachis;
costae copiously scaly, larger scales 1 mm long,
ovate-acute, dark and shining with pale edges
bearing dark setae, smaller ones entirely pale,
either with setae or fringing hairs, no bullate
scales; costules and veins beneath bearing similar
smaller scales; upper surface of costules glabrous.
Type specimen: C. KING 227, Goodenough
Bay, 1200 m, Papua (MICH; fragm. at BM).
Distr. Malaysia: New Guinea and Louisiades.
Ecol. In mountain forest or transition to
mossy forest, on mainland 1200-2600 m, on the
islands 750-1000 m.
147. Cyathea teysmannii Cope. Philip. J. Sc. 4
(1909) Bot. 51 (new name for C. celebica v. A.
v. R.); v. A. v. R. Handb. Suppl. (1917) 39.—
Hemitelia truncata (non BRACK.) CHRIST, Ann.
Jard. Bot. Btzg 15 (1898) 81; in Warburg, Mon-
sunia (1900) 91.—C. celebica v. A. Vv. R. Bull.
Dép. Agr. Ind. Néerl. 18 (1908) 2, non BL. 1828;
Handb. (1908) 26.
Differs from C. tripinnata in the presence of
distinctly bullate pale scales on the costules, and
in thinner and smaller soral scales which do not
140
FLORA MALESIANA
[ser. I, yolue
cover the fully developed sporangia. The speci-
mens do not give information about characters
of the stipe and its scales, which may also be
distinctive.
Type specimen: TEYSMANN 13681 (BO).
Distr. Malaysia: SW. Celebes.
Ecol. At c. 1000 m altitude.
148. Cyathea aciculosa CopeL. Philip. J. Sc. 60
(1936) 104, pl. 9.—C. arachnoidea (non Hoox.)
GRETHER & WAGNER, Un. Cal. Publ. Bot. 23
(1948) 43.
Stipe 18 cm long, dark, smooth, covered with
minute scales, also near base with pale scales
25 by less than 1 mm, bearing dark marginal setae
near apices. Pinnae: lowest 10 cm long, largest
50 cm. Pinnules to 65 by 16 mm, fully pinnate;
tertiary leaflets to 20 pairs, to 7 by almost 2 mm,
larger ones crenate; veins 7 pairs. Sori covered
with overlapping pale scales; no true indusium.
Scales and hairs: pinna-rachis warty, shining,
bearing many very small pale short-fringed scales
and a few long narrow setiferous ones, these latter
also on bases of costae; most scales on costae
pale, bullate, also very small pale scales.
Type specimen: Brass 2887, San Christoval,
900 m, Solomon Is (MICH; dupl. at L, BRI).
Distr. Solomon Is, in Malaysia: Admiralty Is.
Ecol. In forest, to c. 1000 m.
Note. This species is very near C. truncata
BRACK.) CopPeEL. of Fiji and Samoa, differing in the
much larger indusial scales.
149. Cyathea celebica BL. En. Pl. Jav. (1828) 245
(not v. A. v. R. 1908); TINDALE, Contr. N.S.W.
Nat. Herb. 2 (1956) 338, p.p.—Alsophila celebica
Metr. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 53;
v. A. v. R. Handb. (1908) 42.—C. arachnoidea
Hook. Syn. Fil (1865) 24; v. A. v. R. Handb.
(1908) 26; Suppl. (1917) 38.—Alsophila truncata
var. Sagittata CHRIST, Bull. Herb. Boiss. II, 1
(1901) 458.—Alsophila truncata var. nivea CHRIST,
K. Sco. & Laut. Nachtr. (1905) 36; v. A. v. R.
Handb. (1908) 42.—C. quadripinnatifida CopeEL.
Un. Cal. Publ. Bot. 18 (1942) 218; Philip. J. Sc.
77 (1947) 108, pl. 6.
Stipe to 100 cm long, dark, bearing slender dark
spines to 3 mm long, near base bearing persistent
light brown scales 20-40 mm long, mostly not over
1 mm wide, firm and shining, edges bearing close
short oblique concolorous setae, rest of stipe
covered closely with small interlacing setiferous
or fringed scales. Pinnae: small pinnae (5-8 cm
long) sometimes present near base of stipe (seen
only in two specimens); largest pinnae to 70 cm
long. Pinnules 90-140 by 17-28 mm, fully pinnate,
the lower tertiary leafiets stalked; costules (stalks
of tertiary leaflets) 3-5 mm apart; tertiary leaflets
to 15 by 214-5 mm, sometimes dilated at the base,
the largest deeply lobed near the base, the basal
lobes sometimes forming free quaternary leaflets.
Sori medial, protected when young by overlapping
very thin finely fringed scales. Scales and hairs:
pinna-rachis and costae beneath almost covered
with very small irregular pale scales bearing short
marginal hairs and some larger scales bearing
setae; costules at first bearing small ovate scales
bearing setae, lower surface of costules and veins
covered throughout with small pale fringed scales
which interlace and completely cover lower
surface of lamina.
Type specimen: REINWARDT, Ternate (L; dupl.
at BM).
Distr. Queensland, in Malaysia: New Guinea,
Moluccas (Ternate, Ambon), ? Celebes.
Ecol. At 100-1750 m.
Note. The type of C. quadripinnatifida was in
open forest in a ravine and had 4 fronds 600 cm
long (including stipe of 100 cm); these do not show
reduced basal pinnae.
150. Cyathea tripinnata Cope. Philip. J. Sc. 1,
Suppl. 4 (1906) Bot. 251; ibid. 4 (1909) Bot. 40;
v. A. v. R. Handb. (1908) 788; Suppl. (1917) 39;
Ho.Lttum, Rev. FI. Mal. 2 (1954) 120; Copet.
Fern Fl. Philip. 1 (1960) 208.—C. densisora v. A.
v. R. Bull. Jard. Bot. Btzg III,2 (1920) 138; CopeE-.
Fern Fl. Philip. 2 (1960) 208. —C. leucostegia
CopeEL. Philip. J. Sc. 38 (1929) 130; Fern FI.
Philip. 2 (1960) 209.—C. leytensis CopEL. Philip
J. Sc. 38 (1929) 131; Fern FI. Philip. 2 (1960) 209.
—C. arachnoidea (non HooK.) BACKER & POSTH.
Varenfl. Java (1939) 25.
Trunk to 4or 5m. Stipe to at least 40 cm, dark,
bearing scattered sharp spines 1-3 mm long,
covered almost throughout by a felt of very small
setiferous scales; basal scales to 25 by 1 mm,
thin and soft, matted together, edges with some
dark setae, apex a dark seta. Pinnae: lowest 20-30
cm long, largest to 60 cm. Pinnules 90-140 by 17—25
mm, fully pinnate, lower tertiary leaflets distinctly
stalked; costules 4-6 mm apart; tertiary leaflets
to 15 by 3!4mm, the larger ones deeply lobed at
the base, edges crenate; veins to 9 pairs, those in
basal lobes pinnate. Sori near costules, covered to
maturity by overlapping pale thin scales. Scales
and hairs: lower surface of pinna-rachis covered
with minute pale fringed scales; costae bearing
similar scales with some longer narrow ones bearing
a fringe of hairs or dark setae; on costules ovate
flat brown to pale scales, setiferous or fringed;
on veins no scales.
Type specimen: CoPELAND 2068, Mt Mariveles,
in extinct crater, 900 m, Luzon (MICH; dupl. at
LUIS), 12, WK, SYAD. SHAY)
Distr. Malaysia: West Java, Pulau Tioman (E
off Malaya), N. Borneo, Philippines (Luzon to
Mindanao), Moluccas (Ambon).
Ecol. In forest, 250-1700 m; the smaller forms,
represented by C. densisora, C. leytensis and
C. leucostegia probably in more exposed places.
151. Cyathea macrophylla Domin, Acta Bot.
Bohem. 9 (1930) 133 (mew name for Hemitelia
ledermannii BRAUSE); CopEL. Philip. J. Sc. 77
(1947) 108.—Hemitelia ledermannii BRAUSE, Bot.
Jahrb. 56 (1920) 60.
Differs from C. tripinnata in firmer texture of
lamina and in presence of more or less abundant
small scales on lower surface of veins, these scales
Dec. 1963]
CYATHEACEAE (Holttum)
141
bearing short dark setae or short pale hairs, never
long-fringed nor covering lower surface of lamina.
foliolis tertiariis maximis 18 mm longis, omnino
profunde lobatis, segmentis infimis interdum liberis
Type specimen: LEDERMANN 12533, Sepik et stipitulatis.
Region, E. New Guinea (B). Type specimen: WomerRSLEY NGF 13959, Mo-
Distr. Malaysia: New Guinea. robe Distr., NE. New Guinea (K).
Ecol. In forest, from sea-level to 1500 m. Ecol. “Short tree-fern in under-storey of the
forest. Trunk not more than 2 feet tall.
5 feet’, 2000 m.
ay Fronds
var. quadripinnata HOLTTUM, var. nov.
A typo speciei differt: stipitibus c. 10 cm longis;
4. Section Schizocaena
(J. SM.) HOLTTUM, stat. nov.—Schizocaena J. SM. in Hook. Gen. Fil. (1838) t. 2;
Lond. J. Bot. 1 (1842) 661, p.p.; CopEL. Gen. Fil. (1947) 99, p.p.—Fig. 8c, 28-30.
Type species: Schizocaena brunonis J. SM. = Cyathea moluccana R. Br.
Distr. Malaysia and Polynesia.
Taxon. The division into two subsections is perhaps not sharp, but extreme examples of subsect.
Sarcopholis are strikingly different from species of subsect. Schizocaena from Western Malaysia. The
characteristic feature of the section is the position of the basal basiscopic vein of each vein group; this
vein springs from the costa, not from the costule (or in C. moluccana from the midrib of the pinna).
This condition is correlated with the relatively shallow lobing of the pinnules; similar lobing in subg.
Cyathea sect. Gymnosphaera is usually not associated with this vein-character.
The species C. sinuata Hook. and C. hookeri THw., of Ceylon, included here by J. SmitH and Cope-
LAND, have flabelloid scales; in my opinion their nearest relatives are in the group of C. borbonica DEsv.
of Madagascar.
4a. Subsection Schizocaena
Distr. Malaysia, except Moluccas and New Guinea.—Fig. 8c, 28-29.
Taxon. In Borneo and the Philippines species of this subsection are often difficult to characterize
learly, appearing to vary in size and in distribution of scales and hairs on axes.
KEY TO THE SPECIES
1. Fronds simply pinnate, pinnae entire or at most serrate-crenate.
2. Apex of frond a deltoid deeply lobed lamina. Outer veins of each group joining to form a single
excurrent vein. Pinnae sessile, base truncate or subcordate . aa 2 1S22 CC eapiata
2. Apex of frond a pinna of same shape as other pinnae. No anastomosis. Pinnae usually stalked.
3. Pinnae not over 15 mm wide, base narrowly cuneate, stalkes to 12 mm long. Sori in 1-2 rows on
each side of costa, fully indusiate : si 153. C. angustipinna
3. Pinnae 2-4 cm wide, base rather broadly cuneate, ‘stalked o or not. Sori i in fully fertile fronds in more
— 2 rows, indusiate or not.
. Pinnae not long-acuminate, upper usually sessile .
- Pinnae long-acuminate, all stalked .
1. Fronds simply pinnate with deeply lobed pinnae, or bipinnate.
5. Fronds simply pinnate with deeply lobed pinnae, the largest sometimes with free pinnules at their base.
6. Lower surface of pinna-midribs covered near base with long hairs; no free pinnules.
156. C. deminuens
6. Lower surface of pinna-midrib lacking hairs; some free pinnules on largest pinnae.
7. Indusium present (sometimes hidden by mature sorus).
8. Lower pinnae little narrowed at base, free leaflets as zone as lobes, apex not long-acuminate;
pinnae commonly 252 Cmielon tar. w. = = ce 157-2@>altermans
8. Lower pinnae narrowed to base so that free leaflets a are , very small, apex long-acuminate; pinnae
to 18 cm long : 158. C. binuangensis
7. Indusium lacking.
9. Pinnae commonly 25 cm long; no long pale hairs on rachis .
9. Pinnae much shorter; main rachis bearing long pale hairs .
5. Fronds amply bipinnate.
10. Pinnules entire or with crenate edges.
11. Indusia present; pinnules sessile
11. Indusia lacking; pinnules stalked
10. Pinnules distinctly lobed.
12. Sori indusiate.
13. Pinnules of larger pinnae not lobed more than 34, to costa throughout, no free basal pinnules,
Or sometimes one on lowest pinnule of lower pinnae.
154. C. moluccana
155. C. arthropoda
157. C. alternans
159. C. elliptica
157. C. alternans
160. C. obliqua
142
FLORA MALESIANA
[ser.. TI; vols
14. Bullate scales lacking on costules .
14. Bullate scales present on costules.
161. C. integra
15. Basal pinnules of middle pinnae with stalks to at least 4 mm long; pinnules lobed less than
1, way to costa
162. C. stipitipinnula
15. Basal pinnules sessile or nearly SO; ypiinules lobed more than half way to costa.
16. Segments of pinnules acute, falcate; largest pinnules 20-30 mm wide.
163. C. zamboangana
16. Segments of pinnules rounded; largest pinnules not over 20 mm wide.
17. Indusium a disc hidden by sorus .
17. Indusium complete, breaking and persistent at ‘maturity.
18. Lower surface of costae densely scaly .
18. Lower surface of costae not densely scaly . 5
13. Pinnules of larger pinnae always with one or more free basal segments.
164. C. discophora
165. C. megalosora
166. C. suluensis
19. Copious long hairs on lower surface of veins as well as costules.
20. About half the tertiary segments free or separately adnate to costa on largest pinnules. Pin-
nules to 100 by 20 mm. Costae not densely scaly. . . i to, MULGHE
C. robinsonii
20. One or two pairs tertiary segments free on largest pinnules. Pinnules to 80 by 12 mm. Costae
ieee scaly.
Costal scales not setiferous. Pinnules sessile .
5 Costal scales strongly setiferous.
19. No hairs on lower surface of veins.
165. C. megalosora
Lower pinnules with stalks 2-3 mm long.
168. C. senex
22. Free basal segments of largest pinnules deeply lobed. Rachis and pinna-rachis densely
persistently scaly, scales small,
sub-bullate, setiferous .
169. C. sibuyanensis
22. Free basal segments entire. Rachises glabrescent .
23. Pinnules sessile, to c. 65 by 15 mm, often smaller. . .
23. Pinnules stalked (stalks to 8 mm), to 100 by 25 mm .
12. Sori without indusia.
170. C. philippinensis
171. C. assimilis
24. Long-spreading hairs abundant on lower surface of rachis and/or pinna-rachis, often also on
costae.
25. Costules and veins bearing hairs like those of costae on lower surface.
26. Pinnules to 110 mm long, cut 24-34, to costa; costules 5 mm apart .
172. C. trichodesma
26. Pinnules to 65 mm long, cut to within 1 mm of costa; costules 3-314 mm apart.
173. C. wallacei
25. Costules and veins lacking hairs on lower surface.
27. Hairs of lower surface confined, or almost confined, to main rachis .
159. C. elliptica
27. Hairs of lower surface present on pinna-rachis and costae, few or none on main rachis.
174. C. trichophora
24. Long spreading hairs lacking on lower surface of rachis and pinna-rachis.
28. Largest pinnules with a free segment at the base; pinnules on stalks to 4 mm or more long;
texture firm .
28. Largest pinnules lacking. a free segment at the base, ‘almost sessile.
175. C. polypoda
29. Sori on 3-4 pairs of basal veins in each group only, not on distal veins, at maturity confluent.
176. C. obscura
29. Sori on almost all veins, not confluent at maturity.
30. Bullate scales lacking on costae and costules. Pinnules cut to less than 1 mm from costa;
stout erect hairs abundant on upper surface of costules and veins .
177. C. agatheti
30. Bullate scales present on costae and costules. Pinnules less deeply lobed; hairs on upper
surface of costules few
152. Cyathea capitata Corer. Philip. J. Sc. 12
(1917) Bot. 49; C. Cur. & HoLtrum, Gard. Bull.
S. S. 7 (1934) 199, 218.—Schizocaena capitata
CopEL. Gen. Fil. (1947) 99.—Fig. 28f-h.
Trunk \—3 m, bearing c. 12 fronds. Stipe dark,
smooth, at least 40 cm; basal scales pale brown,
firm, to 25 by 3-4 mm, rather thick at the base,
edges bearing rather irregular concolorous setae;
pneumathodes 9-15 mm long, rather widely
spaced. Rachis dark to medium brown, smooth;
lamina 100 cm or more long, simply pinnate, apex
of frond not like a pinna but broadly deltoid and
deeply lobed, lobes grading to upper pinnae.
Pinnae c. 40 pairs, sessile, jointed to rachis, largest
15-19 by 2-3 cm wide at base, lower ones some-
what smaller, edges entire except near apex, base
.178. C. squamulata
truncate to cordate, in the latter case more or less
auricled on both sides, apex short-acuminate,
crenate. Veins pinnate in each group, usually with
3 pairs of veinlets, outer veinlets of each group
always anastomosing with outer veinlets of ad-
jacent groups to form a single excurrent vein;
basal basiscopic vein of each group springing
separately from the costa. Sori usually in 2 rows
on each side of the costa (3-4 sori on each vein-
group) or in narrower pinnae only one row;
indusium thin and translucent, at first covering
sorus, later breaking irregularly and more or less
persistent.
Type specimen: CLEMENS 11033, Mt Kinabalu,
N. Borneo (MICH; dupl. at A, UC, K).
Distr. Malaysia: Borneo (Sarawak: Mt
Dec. 1963]
Murud; N. Borneo: Mt Kinabalu).
Ecol. Near a waterfall and in wet ground near
a stream, in forest, 1400-2100 m.
153. Cyathea angustipinna Ho_trumM, Kew Bull.
16 (1962) 52.
Stipe 30 cm, smooth, scaly near base, scales pale,
firm and shining, to c. 20 by 114 mm, edges bearing
copious short dark setae. Frond 70 cm long,
simply pinnate; pinnae about 18 pairs, jointed to
rachis; terminal pinna like the rest, usually with a
small rudiment beside it. Pinnae stalked to 12 mm
or more (lowest longest), to 12 cm long, fertile
ones 1—1.2 cm wide, sterile to 1.6 cm, bases narrow-
ly cuneate (of lower ones somewhat asymmetric)
apices shortly acuminate, edges entire except to-
wards apices where they are crenate; veins common-
ly in groups of three, middle one sometimes forked,
basiscopic one separately attached to costa. Sori
2 or 3 on each vein-group (middle vein not always
soriferous), in two rather uneven rows on each
side of the costa; indusium thin and pale, covering
young sorus and breaking later, more or less
persistent as a disc around base of old sorus;
costae usually quite glabrous on both surfaces.
Type specimen: RICHARDS 1675, Mt Dulit,
1200 m, Sarawak (K).
Distr. Malaysia:
collections).
Ecol. Sandy bank of a stream, near waterfall,
in shade; trunk 50 cm.
Borneo (Sarawak, two
154. Cyathea moluccana R. Br. in Desv. Mém.
Soc. Linn. Paris 6 (1827) 322; v. A. v. R. Handb.
(1908) 15; CopeL. Philip. J. Sc. 4 (1909) Bot.
32; v. A. v. R. Bull. Jard. Bot. Btzg I, n. 28 (1918)
12.—Schizocaena brunonis J. SM. ex Hook. Gen.
Fil. (1838) t. 2; J. Sm. Hist. Fil. (1875) 243.—
C. brunonis WALL. ex Hook. Sp. Fil. 1 (1844)
15; Syn. Fil. (1865) 16; Bepp. Ferns Br. Ind.
(1865) pl. 87; Handb. (1883) 5; CuHrist, Farnkr.
Erde (1897) 318; Diets in E. & P. Pfl. Fam. 1, 4
(1899) 127; CopeL. Sarawak Mus. J. 2 (1917) 345,
347; HoL_trum, Rev. Fl. Mal. 2 (1954) 117.—
C. pinnata Roxs. Calc. J. Nat. Hist. 4 (1844) 517;
Fl. Ind. ed. Clarke (1874) 762.—Schizocaena
gaudichaudii Fre, Gen. Fil. (1852) 354.—C.
pseudobrunonis CoreL. Philip. J. Sc. 12 (1917)
Bot. 50.—C. fuscopaleata CopeL. I.c. 50.—C.
kinabaluensis Cope. /.c. 51; C. Cur. Gard. Bull.
S.S. 7 (1934) 218.—Schizocaena moluccana Co-
PEL. Gen. Fil. (1947) 99.—Schizocaena kinaba-
luensis COPEL. I.c. 99.—Fig. 28a-e.
Trunk usually not over 50 cm tall. Stipe common-
ly 20-30 cm, dark, scaly near base and finely
warty when scales have fallen; scales medium
brown, firm, 15-30 by 14-3 mm, edges bearing
setae little darker than scale. Lamina to 150 cm
or more long (largest frond reported, stipe with
lamina 308 cm), simply pinnate, apical pinna
usually like the rest (sometimes with a rudiment at
its base), all pinnae articulate to rachis. Pinnae
stalked or the upper ones sessile (stalks variable
in length, the lowest 5-10 mm), 12—28 by 2-4 cm,
edges parallel for most of their length, base asym-
CYATHEACEAE (Holttum)
ze hi
metric (rounded on acroscopic side, broadly
cuneate on basiscopic), apex shortly acuminate
and crenate, edges otherwise entire, rarely slightly
lobed with one lobe to each vein-group; veins
in groups of 3 from the costa (basiscopic one
usually separate from the other two), the median
one forked once or twice to give a group of 3-6
(rarely to 10) veins at the edge, veins all free and
all ending close to the margin, or sometimes the
acroscopic vein ending in a sorus or joining with
another vein. Sori in 1—3 (rarely more) rows on
each side of the midrib, commonly 4—6 on each
vein-group, exceptionally to 10 (in such cases the
edge usually lobed), covered when young by a
thin translucent indusium which breaks at ma-
turity and is more or less persistent, or in some
cases the indusium forming a disc which is hidden
by the mature sporangia; lower surfaces of rachis,
costae and veins usuaily glabrous, occasionally a
few persistent small ciliate or setiferous scales
present.
Type specimen: C. SmitH, Moluccas (BM).
Distr. Malaysia: Central Sumatra, Malay
Peninsula, Lingga, Borneo (excluding south and
south-west), South & Central (?) Celebes, Mo-
luccas (Ceram, Ambon).
Ecol. In forests, 0-900 m.
Notes. Fronds bearing imperfect indusia
appear to be most common in Borneo, but
occur also in the Malay Peninsula. COPELAND
described the three species C. pseudobrunonis,
C. fuscopaleata and C. kinabaluensis as lacking
indusia, but I have not found a specimen totally
devoid of indusia, though in old specimens only
small fragments remain (VAN ALDERWERELT
made the same observation, 1918 /.c.). COPELAND
thought that species could also be distinguished by
size and colour of stipe-scales, but, after seeing a
very large number of specimens, I cannot see any
clear distinction into groups based on such charac-
ters. COPELAND’s fourth species, C. arthropoda, is
here regarded as distinct in shape of pinnae.
R. Brown described this species, but did not
name it, in 1810 (Prod. Fl. N. Holl. 158); DEsvAUXx
supplied a name, copying the information provided
by Brown. The type must be the specimen seen
from the Moluccas by Brown, though it is not
named by him.
155. Cyathea arthropoda Copev. Philip. J. Sc. 6
(1911) Bot. 134, t. 13; v. A. v. R. Handb. Suppl.
(1917) 22.—Schizocaena arthropoda Copel. Gen.
Fil. (1947) 99.
Fronds simply pinnate; lamina to c. 70 cm long,
apical pinna like the rest. Pinnae jointed to rachis,
always stalked (stalks of upper pinnae c. 5 mm,
of lower ones 12-15 mm), 12-20 cm long, fertile
to 214 cm wide, sterile to 314 cm, base almost
equally cuneate, apex caudate-acuminate (cauda
to 4.cm long), sides for the most part not parallel.
Sori in 1-3 irregular rows on each side of the
costa; indusium a narrow irregular ring, hidden by
sporangia, or lacking.
Type specimen: Brooxs 8, Bungo Range,
Sarawak (MICH; dupl. at BM).
144 FLORA MALESIANA [ser Ik voles
Fig. 28. Cyathea moluccana R. Br. a. Middle part of rachis with pinnae, * 14, b. apex of frond, x 4,
c. part of pinna, showing venation and sori, * 114, d. scale from stipe, < 4, e. edge of stipe-scale, * 40.
—C. capitata Core. f. Middle part of rachis with pinnae, < 14, g. apex of frond, 14, A. part of pinna
showing venation and sori, * 114 (a—e CUMING 378, f CLEMENS 27959, g JAcoBs 5790, h CLEMENS 11033).
Dec. 1963]
CYATHEACEAE (Holttum)
145
Distr. Malaysia: Sarawak (neighbourhood of
Kuching).
Ecol. In lowland forest.
Note. This seems to be the most distinct of the
““exindusiate’” species described by CopELAND
from Sarawak (see C. moluccana). Field study is
necessary to decide whether or not it is connected
by intermediate with typical C. moluccana, which
occurs in the same district.
156. Cyathea deminuens HOLTTUM, nom. nov.—
Alsophila parvifolia Hotttum, J. Mal. Br. R. As.
Soc. 6 (1928) 19.
Trunk to 75 cm. Stipe to 15 cm; scales at base
pale brown, rather firm, 10-15 mm by 114 mm,
edges closely set with dark setae. Lamina to 65 cm
long, simply pinnate, pinnae pinnatifid, lower ones
gradually reduced, lowest 314 cm long. Rachis
densely covered beneath throughout with stout
pale spreading hairs to 3 mm long. Largest pinnae
9 by 2 cm, lobed 24 towards the costa; costules
5 mm apart; veins 6—7 pairs, basal basiscopic vein
from costa; lower surface of costae hairy towards
base, and a few hairs on lower costules, no bullate
scales seen; a few long hairs present on upper
surface of costules and veins. Sori at about one
third distance from costule to edge; no indusia;
long paraphyses, with dark walls between the
cells, present.
Type specimen: BoDEN KLoss 14579, Siberut,
Mentawai Is (S; dupl. at BO; K).
Distr. Malaysia: Sumatra (two collections, the
second from S. Sumatra).
Ecol. In lowland forest.
157. Cyathea alternans (WALL. ex HooK.) PRESL,
Abh. K. B6hm. Ges. Wiss. V, 5 (1848) 347;
Bepp. Handb. Suppl. (1892) 2; v. A. v. R. Handb.
(1908) 17; Rip. J. Mal. Br. R. As. Soc. 4 (1926)
5, incl. var. serrata RIDL.; Domin, Acta Bot.
Bohem. 9 (1930) 90, incl. var. sarawakensis (HOOK.)
Domin and var. lobbiana (HooK.) DoMIN; HOLTTUM
Rey. FI. Mal. 2 (1954) 119.—Polypodium alternans
WALL. Cat. (1829) n. 329, nom. nud.—Hemitelia
alternans Hook. Ic. Pl. (1844) t. 622; Sp. Fil. 1
(1844) 29.— Amphicosmia alternans Moore,
Ind. Fil. (1857) civ; Bepp. WHandb. (1883)
10.—C. lobbiana Hoox. Syn. Fil. (1865)
24.—C. sarawakensis Hoox. lc. 23; Hose,
J. Str. Br. R. As. Soc. 32 (1899) 35.—Alsophila
alternans Hook. Syn. Fil. (1866) 41; Bepp. Ferns
Br. Ind. (1867) t. 236.—Schizocaena alternans
J. Sm. Hist. Fil. (1875) 244; Coper. Gen. Fil.
(1947) 99.—Alsophila janseniana v. A. v. R. Bull.
Jard. Bot. Btzg III, 5 (1922) 179.—C. janseniana
Domin, Acta Bot. Bohem. 9 (1930) 126.
Trunk usually less than 2 m. Stipe to 60 cm,
dark, persistently scaly near base, otherwise
glabrescent and smooth; scales medium brown,
firm, shining, to 30 by 2 mm, edges closely seti-
ferous. Pinnae articulate to rachis, lowest some-
what reduced, largest commonly 25 by 4-5 cm,
sometimes to 40 by 9 cm (in type of C. sarawaken-
sis 50 by 16 cm), deeply lobed throughout or with
few to many of the lobes separately joined by the
more or less contracted base of their lamina to the
axis of the pinna, rarely the lowest one or two
forming true pinnules with lamina quite free;
costules of lobed pinnae 7-9 mm apart, of pinnae
in which the lobes are free up to 15 mm apart;
lobes which are not free usually entire and rounded
at apex, free lobes acute at the apex or with margins
sometimes broadly crenate, one crenation to each
vein-group. Veins in the narrower, lobed, pinnae
to c. 10 pairs in each lobe, individual veins forked
Once or twice; in free lobes the veins in small
lateral pinnate groups. Sori usually in one row
on each side of the costule of a lobe of a pinna, in
the largest free lobes occasionally an incomplete
second row; indusium varying greatly, in a mi-
nority of cases completely covering the young
sorus and persistent, in most cases forming a disc
of irregular width covered by the mature sorus,
sometimes only detectable as small fragments;
long pale paraphyses present. Scales on lower
surface of pinna-midribs, costules and veins usual-
ly rather sparse, elongate, narrow and strongly
setiferous, pale to medium brown, in some cases
also bullate scales on costules; in some cases stout
pale hairs present on distal parts of costae and
costules and a few on veins.
Type specimen: WALLICH 329, Penang (K).
Distr. Malaysia: Sumatra (Karo plateau,
Benkulen Distr.), Malay Peninsula (Penang, Pe-
rak, Negri Sembilan, Pahang, Trengganu, Kelan-
ton), Sarawak and North Borneo.
Ecol. In forest,often near streams,at 300-1300 m.
Note. This species is usually found growing in
association with C. moluccana R. Br. and always
within the range of C. moluccana. The variable
C. alternans has the appearance of being a series
of hybrids between C. moluccana and either C.
squamulata or C. ridleyi (which are bipinnate and
exindusiate) with many possible combinations of
characters of pinna-shape and of indusia and
scales and hairs. The type specimen of C. alternans
has pinnae to about 614 cm wide, with 3 pairs of
free adnate lobes, a few lobes longer than adjacent
ones, as in the type of Alsophila janseniana v. A.
v. R.; the sori have disc-shaped indusia, as shown
in Hooker, Ic. Pl. t. 622, though Hooker stated
later (Syn. Fil. 41) that ‘“‘a depression formed by
the sorus on the lobe was mistaken by the artist
for an involucre”’. Hairs occur on the lower sur-
face of costules in some Peninsular specimens
(especially Mou. Nur 11269); I have not seen any
on Bornean ones. Bullate scales appear more
common in Peninsular than in Bornean specimens.
158. Cyathea binuangensis v. A. v. R. Bull.
Jard. Bot. Btzg III, 2 (1920) 136; CopeL. Fern
Fl. Philip. 2 (1960) 206.
Stipe 20 cm, densely scaly at base only; scales
pale, thin, to 20 by 2 mm, edges bearing concol-
orous hairs. Lamina 50 cm long, simply pinnate.
Pinnae: lowest slightly reduced; largest 18 by
4 cm, lobed to 3-4 mm from costa except at base
where one segment is usually almost free; bases of
pinnae conspicuously narrowed (free segment
always smaller than rest), apex rather long-acu-
minate; lobes falcate, almost entire, their costules
146
FLORA MALESIANA
[ser. II, vol: 12
to 9 mm apart; veins to about 10 pairs, mostly
forked. Sori medial; indusium thin, breaking and
persistent; paraphyses very dark. Scales on costae
of pinnae very few, very narrow, pale, bearing
sparse short concolorous hairs; no hairs seen on
lower surface of pinnae, on upper surface present
only near base of costae.
Type specimen: RamMos & EDANOo, BS 28779,
Mt Binuang, Luzon (BO; dupl. at US, MICH, K).
Distr. Malaysia: Philippines (Luzon, one
collection).
Note. This specimen has the aspect of a pre-
maturely fertile plant of C. integra J. SM. ex HOOK.,
but it lacks the characteristic scales of that species.
It has the same degree of division of the frond as
C. alternans (WALL.) PRESL, but is very different in
shape of pinnae, and in paraphyses. It might be a
hybrid of C. integra, but I cannot suggest what
other parent is likely.
159. Cyathea elliptica Cope. Philip. J. Sc. 12
(1917) Bot. 51.—C. subbipinnata Cope . ibid. 56
(1935) 471, pl. 1.—C. holttumii Corer. I.c. 472,
pl. 2.—Gymnosphaera subbipinnata CopeL. Gen.
Fil. (1947) 99.—Gymnosphaera holttumii COPEL.
es 99:
Stipe scaly throughout, also bearing on abaxial
surface some long spreading hairs, the latter some-
times very abundant and to 10 mm long; scales
medium to light brown, shining, edges setiferous;
main rachis similarly scaly and hairy, scales some-
times deciduous. Pinnae: lower ones not greatly
reduced, largest 30-45 cm long. Largest pinnules
4.5-6.5 cm long, 10-14 mm wide, sessile, lobed half
way to costa; costules 3!4-4 mm apart; veins to
5 pairs; lamina-segments rather thin, entire, ends
rounded. Sori medial; no indusia; paraphyses not
longer than sporangia. Scales and hairs: lower
surface of pinna-rachis lacking hairs or a few
present near the base; costae bearing sparse narrow
pale scales and sometimes a few hairs; costules
bearing pale hair-pointed bullate scales.
Type specimen: CLEMENS 10859, Mt Kinabalu,
N. Borneo, (MICH; dupl. at UC).
Distr. Malaysia: North Borneo.
Ecol. In forest, 900-1800 m.
Note. The type-collection of C. holttumii has
much more abundant hairs on all parts of the
frond than the other specimens referred to this
species. The type of C. subbipinnata was collected
at 1800 m, and is small, with largest pinnae only
10 cm long, having only one or two pairs of small
free pinnules; the specimen is old and has lost most
of its scales and hairs. It seems doubtful whether
C. elliptica should be maintained as distinct from
174. C. trichophora COPEL.
160. Cyathea obliqua CopeL. in Elmer, Leafl.
Philip. Bot. 4 (1911) 1150.—Al/sophila obliqua C.
Cur. Ind. Fil. Suppl. 1 (1913) 5; v. A. v. R.
Handb. Suppl. (1917) 54.—Gymnosphaera obliqua
CopreL. Gen. Fil. (1947) 98; Fern Fl. Philip. 2
(1960) 236.
Stipe unknown. Pinnae to 30 cm long. Pinnules
to 72 by 10 mm, the lowest on stalks to 5 mm long,
articulate to rachis, base very unequally cuneate
(narrow on basiscopic side), apex caudate-acu-
minate, edges broadly crenate, lamina rather thin;
veins in pinnate groups each with up to 4 pairs,
lowest basiscopic vein of each group springing
from the costa. Sori 1-3 on each vein-group; no
indusia; paraphyses as long as sporangia. Scales
and hairs: pinna-rachis glabrous on lower surface;
costal scales rather sparse, flat and ovate-acute to
bullate and acuminate, light brown, most with a
short fringe of pale hairs, some bearing dark setae.
Type specimen: ELMER 12354, Sibuyan L.,
Philippines (MICH; dupl. at K, A, BO, US, FI,
S-PAQ SYD se):
Distr. Malaysia: Philippines
once collected), at 600 m.
(Sibuyan LI.,
161. Cyathea integra J. Sm. ex Hook. Ic. PI.
(1844) t. 638, incl. also var. petiolata HooK. lI.c.
t. 638, fig. 2; Sp. Fil. 1 (1844) 26; Syn. Fil. (1865)
23; v. A. v. R. Handb. (1908) 20 (not Suppl. 25,
which is C. urdanetensis COPEL.); COPEL. Philip.
J. Sc. 4 (1909) Bot. 35; Fern Fl. Philip. 2 (1960)
200.—Trichopteris falcata LLANOS, Fragm. PI.
Filip. (1851) 111 (fide MERRILL, Sp. Blanc. 1918,
41).—C. hypocrateriformis v. A. v. R. Bull. Jard.
Bot. Btzg II, n. 7 (1912) 9; Handb. Suppl. (1917)
39; CopEL. in Elmer, Leafl. Philip. Bot. 5 (1913)
1680.—C. bulusanensis CopeEL. in Elmer, Leafl.
Philip. Bot. 9 (1920) 3109; Fern FI. Philip. 2
(1960) 201.—C. arguta CopeL. Philip. J. Sc. 38
(1929) 133; Fern Fl. Philip. 2 (1960) 203.—
C. breviloba Cope. Philip. J. Sc. 81 (1952) 13;
Fern FI. Philip. 2 (1960) 201.—Fig. 29a, b.
Stipe 20-40 cm, base with spines 2 mm long;
scales at base of stipe thin, pale, edges closely
setiferous, to 25 by 114 mm; also above base a
more or less persistent cover of very small pale
fringed or setiferous scales. Main rachis medium
brown when dry, in basal part bearing scattered
short spines, otherwise smooth and glabrescent.
Pinnae: lowest c. 15 cm long, largest 60 cm.
Pinnules on larger pinnae 80-120 by 15-25 mm,
sessile or lowest stalked to 2 mm (rarely to 4 mm),
apex acuminate (not caudate), in lowest pinnules
on lower pinnae the basal segment sometimes just
free, otherwise whole pinnule lobed to 2-3 mm
from costa; costules 6 mm apart; veins 6—8 pairs;
lamina-segments firm, edges almost entire or
distinctly crenate towards apices, apex bluntly to
acutely pointed at end of falcate costule. Sori
medial; indusium at first complete, thin and trans-
lucent but firm, breaking and persisting at ma-
turity; paraphyses abundant, as long as sporangia,
pale. Scales and hairs: pinna-rachis beneath gla-
brescent, smooth or sparsely warty, residual
scales small, pale, fringed or some with setae;
on costae near base narrow pale scales with dark
marginal setae, smaller ones with irregular fringe
of pale hairs; on costules scales like smaller ones
on costae, not bullate; stout spreading hairs rarely
present on lower surface of costae and costules near
apex of pinnule, not on upper surface of costules.
Type specimen: CumING 120, Luzon (K; dupl.
at US).
Dec. 1963]
CYATHEACEAE (Holttum)
Fig. 29. Cyathea integra J. SM. a. Part of pinnule, upper surface,
x 2, b. lower surface showing scales
and sori, X 6.—C. squamulata (BL.) CopeL. c. Part of sterile pinnule, lower surface, = 2, d. part of
fertile pinnule showing sori and scales, < 6 (a—b SINCLAIR 9539, c SINCLAIR 10336, d KIAH 32179).
Distr. Malaysia: Philippines (Luzon, Mindoro,
Panay, Samar, Catanduanes, Basilan, Biliran,
Mindanao).
Ecol. In forest at low and medium altitudes;
range reported 500-1200 m, but few specimens
bear relevant information.
Notes. With the original description of this
species, HooKER cited first a specimen from
Ambon, then CuminG 120 from Luzon; his figure
was certainly prepared from the Luzon specimen,
which I therefore regard as the type. The Ambon
specimen is quite distinct in shape of lamina-
segments and details of scales; it belongs to
C. tripinnatifida Roxs.
There is considerable variation in size, shape
and marginal teeth of the segments of the pinnules
in C. integra; there is also variation in the presence
of hairs on the lower surface of costae and in
length of stalks of lower pinnules. I have not ob-
served clear correlation of such characters, and
do not think that var. petiolata Hook. can be
regarded as a distinct variety.
162. Cyathea stipitipinnula HoLtrum, Kew Bull.
16 (1962) 62.
Stipe to more than 30 cm, medium dull brown,
rather persistently scaly, warty where scales have
fallen; larger scales to 25 by 3 mm, shining
brown with paler edges bearing many dark setae,
also minute pale irregularly fringed scales; main
rachis smooth, glabrescent. Pinnae to 45 cm long.
Pinnules almost at right angles to rachis, to 65 by
12 mm, lobed less than half way to costa, lobes
rounded and entire, texture coriaceous, the lowest
pinnules with cordate bases and on stalks to
4 mm long; costules 314-4 mm apart; veins 3-4
pairs, thick. Sori usually 3 to each lobe, medial;
indusia pale, firm, complete, breaking irregularly
and persistent; paraphyses dark. Scales and hairs:
scales near base of costae ovate-acute, flat, light
brown, with numerous crisped marginal hairs or
the larger with some setae, grading to light brown
bullate scales (often fringed near apices) on
costules and on veins.
Type specimen: CLEMENS 33156, Mt Kinabalu,
N. Borneo (K; dupl. at BO, A, UC, L).
Distr. Malaysia: N. Borneo (Mt Kinabalu,
several collections).
Ecol. In open places in forest, 1200-1500 m.
163. Cyathea zamboangana CopeL. Philip. do. SE
30 (1926) 325; Fern FI. Philip. 2 (1960) 201.—
C. urdanetensis CopeL. Philip. J. Sc. 38 (1929) 132;
Fern Fl. Philip. 2 (1960) 203.—C. integra (non
148 FLORA MALESIANA [ser. II, vole
J. Sm. ex Hook.) v. A. v. R. Handb. Suppl. of C. megalosora CopeEL., and is intermediate in
(ET), 23 soral characters between C. megalosora and the
Trunk: \eaf-scars elliptic, 244 cm wide, in al-
ternate whorls of 3. Stipe 30 cm, base spiny (spines
slender, dark, abundant, 1-3 mm) and densely
covered with scales; scales to c. 10 by 1 mm,
mostly smaller, edges with close dark setae. Main
rachis spiny, pale, glabrescent. Pinnae: lowest
somewhat reduced, largest to 60 cm or more long.
Pinnules on larger pinnae 100-130 by 25-35 mm,
lower ones on stalks 3-4 mm long, one lowest
segment sometimes just free, rest of pinnule
lobed to about 214 mm from costa; costules 6 mm
apart; veins to 8 pairs, mostly forked, a few with
acroscopic braoch again forked; lamina-segments
thin but firm, apices falcate, acute, edge some-
times serrate near apex. Sori almost medial;
indusium complete, globose, breaking and per-
sistent at maturity. Scales and hairs: pinna-rachis
pale, smooth, with some residual narrow pale
setiferous scales; on costae scattered very small
pale short-fringed scales and near base a few
narrow pale setiferous scales, sometimes a few
stout pale hairs near apex of pinnule; on costules
pale bullate scales, sometimes with setae near
apex, also sometimes stout pale hatrs near apices
of segments; upper surface of costules lacking
hairs.
Type specimen: COPELAND 1646, near San
Ramon, Mindanao (MICH).
Distr. Malaysia: Philippines (Mindanao).
Ecol. In forest, 500-800 m.
164. Cyathea discophora HOoLTTUM, Kew Bull. 16
(1962) 54.
Stipe finely warty throughout, persistently scaly
near base; scales pale, to 25 by 2 mm, edges
closely set with short dark setae; rachis light
brown, glabrescent, sparsely and finely warty.
Pinnae to 50 cm long, pinnules rather widely
spaced. Pinnules sessile, to 8 by 114 cm, lobed
24, towards the costa except at the very base,
lowest segment not free; costules 4-414 mm
apart; lamina-segments rather thin, edges cre-
nulate, apex broad, sinuses narrow; veins 6—7
pairs. Sori medial; indusium at length a thin
brown disc of irregular shape, sometimes excentric,
completely covered by mature sorus; paraphyses
dark, a little longer than sporangia. Scales and
hairs: pinna-rachis glabrescent beneath; costae
rather densely scaly, scales at base pale, flat,
elongate with short rather stiff pale marginal
hairs, grading through similar scales bullate at
base to rather large pale bullate scales with or
without marginal hairs near their acuminate
apices; a few long pale hairs on costa near apex of
pinnule; costules bearing large pale bullate acu-
minate scales and few hairs; upper surface of
costules glabrous.
Type specimen: CLEMENS 31698, Mt Kinabalu,
N. Borneo (B; dupl. at US, BO).
Distr. Malaysia: N. Borneo (Mt Kinabalu,
once collected).
Ecol. In open place in forest, 2400 m.
Note. This occurs within the altitudinal range
exindusiate C. squamulata (BL.) COPEL. which
occurs at lower altitudes.
165. Cyathea megalosora CopeL. Philip. J. Sc. 12
(1917) Bot. 54; C. Cur. Gard. Bull. S.S. 7 (1934)
221.
Trunk to at least 2 m. Stipe c. 30 cm, pale to
medium brown when dry, densely scaly, finely
warty where scales have fallen; scales thin, pale,
somewhat crisped, to c. 25 by 1144 mm, edges
bearing rather sparse dark setae towards apex.
Pinnae: lowest 20 cm long, largest 35 cm. Pinnules
to 60 by 12 mm, almost sessile; lowest 1-2 seg-
ments free, rest of pinnule lobed nearly to costa;
costules 4-5 mm apart; veins 5—7 pairs; lamina-
segments very firm, edges crenate, apices rounded,
sinuses narrow. Sori medial; indusium firm, brown
-translucent, quite covering sorus to maturity,
breaking irregularly and persistent; paraphyses
as long as sporangia. Scales and hairs: pinna-
rachis persistently densely scaly on lower surface,
scales long, pale, entire or nearly so, bases of
smaller ones bullate; costae densely scaly on lower
surface, scales elongate, pale, almost entire, not
setiferous, on distal part of costae many long
spreading hairs; costules bearing similar hairs, and
scattered hairs on veins also; upper surface of
costules and veins bearing long pale hairs.
Type specimen: ToppinGc 1759, Mt Kinabalu,
N. Borneo (US; dupl. at A, K, SING, S-PA).
Distr. Malaysia: N. Borneo (Mt Kinabalu).
Ecol. In mossy forest on ridges, 2200-2900 m.
At the highest altitudes the lamina of fronds may
be only 40 cm long, pinnae to 1014 by 214 cm,
with about 5 pairs of free pinnules which are
lobed half-way to the costa.
166. Cyathea suluensis BAK. J. Bot. 17 (1879) 65;
v. A. v. R. Handb. (1908) 18; Cope. Philip. J.
Sc. 4 (1909) Bot. 35; Fern Fl. Philip. 2 (1960) 202.
—C. sessilipinnula Cope. Philip. J. Sc. 38 (1929)
134; Fern Fl. Philip. 2 (1960) 202.
Stipe to more than 30 cm, minutely spiny to the
base; no scales seen. Pinnae: lowest about 10 cm
long, largest 30 cm. Pinnules on larger pinnae
50-70 by 13-18 mm, sessile, short-acuminate,
lobed 14-34 towards the costa, no free basal
segments; costules 4-5 mm apart; veins 4~7 pairs,
strongly oblique; lamina-segments thin, slightly
crenate near rounded apices, sinuses narrow.
Sori medial; indusium complete, thin, pale,
breaking irregularly and persistent. Scales and
hairs: pinna-rachis beneath glabrescent, residual
scales small, pale, short-fringed; on costae near base
narrow flat pale dark-setiferous scales, throughout
brown bullate scales, often with setae near apex,
distally some stout pale hairs; on costules bullate
scales, pale brown, and a few hairs; no hairs on
upper surface of costules.
Type specimen: BuRBIDGE s.n., 1877-78, Sulu
Is (K).
Distr. Malaysia: Philippines (Mindanao, Ba-
silan, Sulu Is), Moluccas (Ternate ?).
Dec. 1963]
CYATHEACEAE (Holttum)
uae
Ecol. At c. 600 m.
Note. Specimens from Ternate placed tenta-
tively in this species are young and sterile; they
agree in form of pinnules, venation and scales,
but differ in having rather numerous stout hairs
on the upper surface of costules.
167. Cyathea robinsonii Copet. Philip. J. Sc. 6
(1911) Bot. 145; v. A. v. R. Handb. Suppl. (1917)
30; CopeL. Fern Fl. Philip. 2 (1960) 203.—C.
pseudoalbizzia Cope. Philip. J. Sc. 38 (1929) 135;
Fern FI. Philip. 2 (1960) 203.
Stipe to at least 40 cm, warty where scales have
fallen; scales pale to brownish, shining, to 20 mm
long, edges setiferous; main rachis finely warty,
bearing some hairs throughout, densely hairy
towards apex of frond, hairs mixed with small
pale scales. Pinnae to 40 cm long. Pinnules to
100 by 20 mm, the lowest with stalks 5-10 mm
long; at base of larger pinnules 1-2 pairs of free
tertiary leaflets, then several pairs of segments
separately adnate by narrow bases to costa, rest of
pinnule lobed nearly to costa; costules on larger
pinnules 5-7 mm apart; lamina-segments firm,
dark on upper surface when dry, edges almost
entire; veins 5—6 pairs. Sori medial; indusium at
first complete, rather firm, breaking irregularly
and persistent. Scales and hairs: pinna-rachis and
costae densely hairy and scaly on lower surface,
hairs spreading, pale, 2 mm long, scales narrow,
pale, sometimes setiferous; on lower surface of
costules and veins abundant pale spreading hairs,
no scales.
Type specimen: ROBINSON BS 9394, Mt Binuang,
Luzon (MICH; dupl. at K, US, P, UC).
Distr. Malaysia: Philippines (Luzon).
Ecol. At 875-1150 m.
Note. The type of C. pseudoalbizzia is small,
with pinnules to 70 by 12 mm.
168. Cyathea senex v. A. v. R. Bull. Jard. Bot.
Btzg II, n. 16 (1914) 4; Handb. Suppl. (1917)
34.
Stipe densely scaly throughout, scales to 30 mm
long, mostly not over 1 mm wide, pale, edges
closely set with dark setae; main rachis similarly
scaly near base only. Pinnae to 40 cm long; lowest
pinnae somewhat reduced. Pinnules to 80 by 12
mm, lowest 1 or 2 segments almost free, rest of
pinnule lobed to within 1 mm of costa; costules
314 mm apart; lamina-segments firm, dark above
when dry, edges crenate, sinuses narrow; veins to
6 or 7 pairs. Sori medial, indusium at first thin and
complete, breaking and persistent; paraphyses
pale. Scales and hairs: pinna-rachis bearing pale
spreading hairs and many small pale setiferous
scales; scales at base of costae narrow, pale, seti-
ferous, grading to acuminate bullate scales at
apex, spreading hairs present throughout; a few
bullate scales present on costules, with many
hairs; erect pale hairs also present on lower surface
of veins; thick curved hairs scattered on upper
surface of costules and veins.
Type specimen: MATTHEW 526-A, Mt Singga-
lang, Sumatra (BO; dupl. at K).
Distr. Malaysia: Central Sumatra.
Ecol. At 1500-1800 m.
Note. MATTHEW gave the number 526 also to
specimens of C. sumatrana BAK., which are at Kew,
from the same locality.
169. Cyathea sibuyanensis Cope. in Elmer, Leafl.
Philip. Bot. 4 (1911) 1150; v. A. v. R. Handb.
Suppl. (1917) 38; Copev. Fern Fl. Philip. 2 (1960)
204.
Stipe not known. Main rachis closely warty on
lower surface, bearing many very small dull
scales. Pinnae to 35 cm long. Pinnules to 90 by
mostly to 20 mm (on a lower pinna to 25 mm),
lowest 1-2 pairs of segments quite free (lowest
sometimes stalked) and deeply lobed at the base,
then several segments contracted at the base and
separately adnate to the costa, rest of pinnule
lobed almost to the costa; costules 5-6 mm apart,
decidedly oblique; lamina-segments firm, edges
almost entire to crenate; veins 6—7 pairs, the lowest
in lobes of basal segments pinnately branched.
Sori medial; indusium pale and translucent, at
first completely covering sorus, breaking and
persistent. Scales and hairs: pinna-rachis closely
warty and covered with very small dull scales, the
largest setiferous, hairs also mixed with the
scales, at least towards apex of pinna; costae
scaly near base, hairy in apical half, scales mostly
ovate-acute, hardly 1 mm long, brown, edges
with a few setae or short hairs near tip, none
bullate; costules bearing hairs on lower surface,
but no hairs on veins; costules glabrous on upper
surface.
Type specimen: Evmer 12513, Mt Géiting-
Giting, Sibuyan I. (MICH; dupl. at K, A, FI, US,
BOR Siw sla):
Distr. Malaysia: Philippines (Sibuyan, one
collection).
Ecol. Altitude 1450 m.
170. Cyathea philippinensis BAk. Ann. Bot. 5
(1891) 186; v. A. v. R. Handb. (1908) 16, 783;
CopeL. Philip. J. Sc. 4 (1909) Bot. 111, incl. var.
nuda CopEL.; Vv. A. v. R. Handb. Suppl. (1917) 23;
Cope-. Fern FI. Philip. 2 (1960) 205.—C. bicolana
CopeL. in Elmer, Leafl. Philip. Bot. 9 (Mar. 1920)
3108; Fern Fl. Philip. 2 (1960) 204.—C. ramosiana
v. A. v. R. Bull. Jard. Bot. Btzg III, 2 (June 1920)
137; CopeL. Fern Fl. Philip. 2 (1960) 206.—C.
heteroloba Copret. Philip. J. Sc. 38 (1929) 134;
Fern Fl. Philip. 2 (1960) 204.—C. bontocensis
Cope. Philip. J. Sc. 46 (1931) 209; Fern FI.
Philip. 2 (1960) 205.
Stipe 12-35 cm, base dark and warty; scales
pale to brownish, to 25 by 2 mm, edges bearing
concolorous setae. Lowest pinnae 4-10 cm long,
largest to 30 cm. Pinnules to 65 by 15 mm, lowest
2-3 segments of larger pinnules free or nearly so,
rest of pinnule lobed 14-34 towards costa; cos-
tules 314-514 mm apart; lamina-segments entire,
apex broadly rounded; veins 4-6 pairs. Sori
nearer to costule than edge; indusium complete,
thin, translucent, breaking and persistent. Scales
and hairs: on lower surface of pinna-rachis narrow
150
FLORA MALESIANA
[ser.- Tl) vel
pale setiferous scales; on costae at base elongate
flat scales with pale fringe or dark setae, grading
to pale bullate scales, pale thick hairs also usually
present near apex of costa; on costules bullate
scales, some fringed near apices, and usually also
a few hairs; no hairs on upper surface of costules
and veins.
Type specimen: ex Hort. Veitch, cult. Kew,
origin Philippines, Feb. 1878 (K).
Distr. Malaysia: Philippines (Luzon, Min-
doro).
Ecol. In mountain forest (only altitude records
are 1400-1500 m).
Note. The type specimen is a frond of a small
cultivated plant, bearing pinnules only 20 by 6 mm,
but in general shape and in scales and sori it re-
sembles the type specimens of the other species
above cited, from which the present description is
prepared.
171. Cyathea assimilis Hoox. Syn. Fil. (1865) 24;
v. A. v. R. Handb. (1908) 20; CopeL. Philip. J. Sc.
4 (1909) Bot. 49.—C. beccariana CESATI, Atti Ac.
Napoli 7® (1876) 3.—C. dulitensis BAK. Kew Bull.
(1896) 40; v. A. v. R. Handb. (1908) 16; CopeE-.
Philip. J. Sc. 4 (1909) 33.—C. ampla CopPeEL.
Philip. J. Sc. 6 (1911) Bot. 361; v. A. v. R. Handb.
Suppl. (1917) 26; non HoLttrum, Rev. FI. Mal. 2
(1954) 135.—C. stipitulata CopeEL. Philip. J. Sc. 6
(1911) Bot. 362; v. A. v. R. Handb. Suppl. (1917)
29.
Stipe to 65 cm, medium to dark brown, finely
warty, persistently scaly near base; scales medium
brown, shining, firm, 15-20 by 1-2 mm, edges
closely setiferous. Lamina to almost 200 cm long;
largest pinnae to 55 cm long, more or less distinctly
articulate to rachis, the lower pinnae with stalks
to 4cm long. Largest pinnules 80-90 by 25 mm, the
lowest with stalks 4-8 mm, basal 1-2 lamina-
segments quite free, next 1-2 pairs scmetimes free
with adnate base, rest of pinnule lobed to 1-2 mm
from costa, apex evenly attenuate; in somewhat
smaller pinnules only the basal basiscopic seg-
ment free, or no free segments; costules S-7 mm
apart; lamina-segments firm, edges more or less
crenate, apices rounded; veins 8—10 pairs. Sori
medial; indusium at first quite covering sorus,
pale and thin, breaking irregularly and persistent.
Scales and hairs: pinna-rachis beneath glabrous
or with a few small setiferous scales; costae be-
neath near base bearing narrow strongly setiferous
scales grading to small bullate setiferous scales
distally and on costules; no hairs on lower surfaces
of pinna-rachis, costae and costules.
Type specimen: Loss s.n., 1857, hills, Sarawak
K).
Distr. Malaysia: S. Sumatra (Mt Dempo),
Borneo (Sarawak).
Ecol. Forests, 300—2000 m, the more coriaceous
specimens in ridge-forest on sandstone.
Notes. The type collection of C. dulitensis
was a small plant with largest pinnae only just
pinnate at base; the later collection of RICHARDS
from the same locality has amply bipinnate fronds,
agreeing in other characters, and probably re-
presents the full development of the species. All
specimens from Mt Dulit are much darker in all
parts than those from elsewhere in Sarawak (e.g.,
the type of C. assimilis) and have a thicker lamina,
but do not differ in other characters. The specimen
from Mt Dempo in Sumatra is dark like those
from Dulit.
The type collection of C. stipitulata is an unusu-
ally small specimen, with pinnules 70 by 17 mm,
costules 4144 mm apart and veins 6—7 pairs; it
was found on Mt Matang, near Kuching, at 300
m.
In HoLtrum, Rev. FI. Mal. 2 (1954) 135, the
name C. ampla is wrongly given to a specimen of
C. polypoda which has unusually wide pinnules;
this latter species is quite exindusiate.
C. assimilis is closely related to C. philippinensis
Bak. and appears to differ chiefly in the consistent-
ly larger size of all parts of the frond.
172. Cyathea trichodesma (ScortT.) CopeL. Philip.
J. Sc. 4 (1909) Bot. 55.—Alsophila trichodesma
Scort. in Bedd. J. Bot. 25 (1887) 321; BeEpp.
Handb. Suppl. (1892) 3; v. A. v. R. Handb.
(1908) 35.—Alsophila margarethae SCHROET. ex
Curist, Ann. Jard. Bot. Btzg 20 (1905) 136;
v. A. v. R. Handb. (1908) 33.—C. margarethae
Cope-. Philip. J. Sc. 4 (1909) Bot. 38; C. Cur.
Gard. Bull. S.S. 7 (1934) 220.—C. burbidgei [non
(BAK.) CopeL.] Ho_ttum, Rev. Fl. Mal.2 (1954) 124.
Trunk slender, to 414 m. Stipe fairly long, near
base densely scaly (not hairy), finely warty when
scales have fallen; scales medium to light brown,
shining, firm, rather dark brown when dried, to
about 25 by 2 mm, edges closely set with short
dark setae. Frond including stipe 2-314 m long.
Pinnae: lowest slightly reduced, largest about
60 cm long. Pinnules commonly to 90 by 15 mm,
largest seen 110 by 20 mm, nearly sessile, shortly
acuminate, lobed to about 2 mm from costa, no
free basal segments; costules 444-5 mm apart;
veins 6-8 pairs; lamina-segments thin, crenate,
sinuses narrow. Sori medial, often confluent at
maturity; no indusium; paraphyses a little longer
than sporangia. Scales and hairs: pinna-rachis,
costae, costules and veins on lower surface bearing
many pale spreading hairs 1-2 mm long; scales on
costae and costules sparse, pale, some narrow and
flat, some bullate, most bearing dark setae; hairs
present on upper surface of costules and veins.
Type specimen: SCORTECHINI s.n., Perak (BM).
Distr. Malaysia: Malay Peninsula (central
part), Borneo (Sarawak, N. Borneo).
Ecol. Lowland forest, sometimes by rivers, to
1500 min N. Borneo.
Notes. Bornean specimens are all smaller than
those from the Malay Peninsula, the largest having
pinnules to 70 by 15 mm; they also lack hairs on
the upper surface of costules. In size, the Bornean
specimens are nearer to C. trichophora, but the
distribution of hairs is different.
In the Malay Peninsula has been found a speci-
men rather intermediate between C. trichodesma
and C. alternans, both of which species were
growing near it.
Dec. 1963]
CYATHEACEAE (Holttum)
151
173. Cyathea wallacei (MeTT. in KUHN) COPEL.
Philip. J. Sc. 4 (1909) Bot. 48.—Alsophila wallacei
Metrt. in Kuhn, Linnaea 36 (1869) 153; v. A. v. R.
Handb. (1908) 36.—Alsophila burbidgei Bax. J.
Bot. 17 (1879) 38; v. A. v. R. Handb. (1908) 33.—
C. burbidgei CopeL. Philip. J. Sc. 4 (1909) Bot. 55;
non HOLTTuM, Rev. Fl. Mal. 2 (1954) 124, which
is C. trichodesma.—Gymnosphaera burbidgei Co-
PEL. Gen. Fil. (1947) 99.
Stipe 30 cm or more, pale and smooth above the
base; scales at base light brown, firm, to 15 by 2
mm, setiferous; main rachis finely hairy on
abaxial surface in apical part. Largest pinna seen
38 cm long. Pinnules to 65 by 13 mm, sessile, apex
abruptly narrowed, lobed to within 1 mm of costa,
lowest segment almost free; costules 3-314 mm
apart; lamina-segments thin, slightly crenate, si-
nuses narrow; veins 4-6 pairs, mostly simple.
Sori medial, lacking indusia. Scales and hairs:
lower surface of pinna-rachis, costae, costules
and veins bearing pale spreading hairs 1 mm long;
pale bullate scales present on costae and costules;
upper surface of costules and veins bearing
scattered long spreading hairs.
Type specimen: WALLACE s.n., 1857, Borneo
(original lost ?; dupl. at Kew).
Distr. Malaysia: Borneo (Sarawak; N. Borneo).
Ecol. In lowland forest, at least sometimes on
poor sandstone soil.
Notes. The pinnules of this species are more
deeply cut than those of other species of sect.
Schizocaena, and have closer costules; they have
the aspect of sect. Sphaeropteris, but are much
smaller than normal in that section, and have
hairiness like that of some members of sect.
Schizocaena.
Alsophila wallacei was credited to METTENIUS by
CHRISTENSEN, Ind. Fil., but it was described without
author’s name in the original paper.
174. Cyathea trichophora CopeL. Philip. J. Sc. 6
(1911) Bot. 363.—C. poiensis CopEL. I.c. 362.—
Alsophila poiensis v. A. v. R. Handb. Suppl. (1917)
56.—Alsophila trichophora v. A. v. R. L.c. 72.—
C. mollis Corey. Philip. J. Sc. 12 (1917) Bot. 52;
C. Cur. Gard. Bull. S.S. (1934) 220.—C. ramosii
CopeL. Philip. J. Sc. 30 (1926) 325.—Alsophila
ramosii C. Cur. Ind. Fil. Suppl. 3 (1934) 23.—
C. bipinnatifida CoPeEL. Philip. J. Sc. 56 (1935) 97,
pl. 2 (not C. bipinnatifida (BAK.) Domtn, 1929).—
Gymnosphaera bipinnatifida CopEL. Gen. Fil. (1947)
99.—Gymnosphaera mollis CoPEL. l.c. 99.—Gymno-
sphaera trichophora Cope. I.c. 99; Fern Fl. Philip.
2 (1960) 236.
Trunk to 50 cm. Stipe 25-50 cm, at least the
basal part persistently scaly, scales to 20 by 3 mm,
light brown, shining, edges setiferous; main rachis
bearing more or less abundant narrow pale seti-
fercus scales and also spreading hairs 2 mm long.
Pinnae: lowest reduced and deflexed, largest 25-30
cm long. Pinnules 30-55 by 10-14 mm, lobed half-
way to costa; costules 314-4 mm apart; veins 3-5
pairs; lamina-segments thin, entire, ends rounded.
Sori medial; no indusia; paraphyses not longer
than sporangia. Hairs rather abundant on lower
surface of pinna-rachis and costae, sometimes on
costules; some pale bullate scales present on cos-
tules.
Type specimen: Ramos 949, Prov. Laguna,
Luzon (MICH; dupl. at FI, UC).
Distr. Malaysia: Philippines (throughout),
Borneo (Sarawak and N. Borneo).
Ecol. Apparently in low country forest, highest
record 1200 m.
Notes. The type of C. bipinnatifida (from
Basilan I.) is from a young plant, with simply
pinnate frond bearing few sori; the upper pinnae
are closely similar to pinnules of other specimens
referable to C. trichophora. It may be that C. tri-
chophora and C. elliptica should be united; the
latter has a different distribution of hairs and has
apparently only been found at higher altitudes.
175. Cyathea polypoda BAK. Trans. Linn. Soc. II,
Bot. 4 (1894) 250; v. A. v. R. Handb. (1908) 18;
C. Cur. Gard. Bull. S.S. 7 (1934) 219; HoLttrum
Rev. Fl. Mal. 2 (1954) 122.—C. kemberangana
CopeEL. Philip. J. Sc. 12 (1917) Bot. 52; C. Cur.
Gard. Bull. S.S. 7 (1934) 219.—Alsophila kem-
berangana C. Cnr. Ind. Fil. Suppl. 3 (1934) 22.—
Gymnosphaera dinagatensis CopeL. Philip. J. Sc.
81 (1952) 19, pl. 14; Fern FI. Philip. 2 (1960) 235.
—C. ampla (non CopeEL.) HoLttum, Rev. Fl. Mal.
2 (1954) 125.—Gymnosphaera glabra (non BL.)
CopeL. Fern FI. Philip. 2 (1960) 235.
Trunk to 3 m, covered with persistent leaf-bases;
small branches often borne on lower part of trunk.
Stipe to 80 cm, pale (green when living) to rather
dark, densely scaly near base, finely warty after
scales have fallen; scales shining medium brown,
firm, to 30 by 2 mm, edges bearing close con-
colorous setae; rachis and pinna-rachis glabrescent
on lower surface. Pinnae to 60 cm long, lower ones
long-stalked, not greatly reduced. Pinnules com-
monly 85 by 20 mm, sometimes to 110 by 27 mm,
all stalked, stalks of lowest to 9 mm; basal 1-2
segments of lowest pinnules of largest pinnae
quite free, then sometimes a pair with lamina
adnate at base but free, rest of pinnule (whole of
smaller pinnules) lobed to 1-2 mm from costa;
costules 414-514 (sometimes to 644) mm apart;
veins 7-9 pairs, forked, acroscopic branch some-
times forked again; lamina-segments rather thick
and rigid when dry, edges crenate, apices rounded,
sinuses narrow except near base of largest pin-
nules. Sori nearer to costule than to edge; no in-
dusium; paraphyses a little longer than sporangia.
Scales and hairs: small dark to medium brown
setiferous scales near base of costae; bullate scales,
often setiferous, on costules, all scales often early
caducous.
Type specimen: HAVILAND 1479, Mt Kinabalu,
N. Borneo (K).
Distr. Malaysia: Malay Peninsula, Borneo
(Sarawak, N. Borneo), Philippines (Panay, Minda-
nao).
soe In open places on ridge-crests and sum-
mits, 600-2200 m; specimens from the higher
elevations are decidedly coriaceous.
Note. BAKER described this species as indusiate;
152
he had a young frond on which bullate scales in
some cases partly cover the immature sori.
176. Cyathea obscura (ScortT.) CopeL. Philip. J.
Sc. 4 (1909) Bot. 37; Ho_trum, Rev. Fl. Mal. 2
(1954) 124.—Alsophila obscura Scort. in BEpp. J.
Bot. 25 (1887) 321, t. 278, fig. 2; Handb. Suppl.
(1892) 3; v. A. v. R. Handb. (1908) 34; Suppl.
(1917) 57.—Alsophila subobscura v. A. v. R. Bull.
Jard. Bot. Btzg II, n. 20 (1915) 1, t. 1; Handb.
Suppl. (1917) 57.—C. bartlettii CopeL. Un. Cal.
Publ. Bot. 14 (1929) 371.—C. pulchra CopEL.
I.c. 372.—C. subobscura Domin, Pterid. (1929)
263.—Alsophila bartlettii C. Cur. Ind. Fil. Suppl.
3 (1934) 20.—Alsophila pulchra C. Cure. I.c. 23.—
Gymnosphaera pulchra CopPeEL. Gen. Fil. (1947) 99.
Stipe dark to medium brown, densely scaly
towards the base, finely warty where scales have
fallen; scales 20-40 by 2-314 mm, shining, pale
brown, edges closely setiferous. Pinnae: lowest
somewhat reduced (sometimes only 10 cm long),
largest 50 cm long. Pinnules: largest commonly
60-70 by 12-13 mm, largest seen 80 by 15 mm, on
stalks 1-2 mm long, apex shortly acuminate,
edges lobed 14-24 distance to costa; costules
3144-4 mm apart; veins about 6 pairs; lamina-
segments firm, edges almost entire, apices bluntly
pointed and asymmetric. Sori medial, on about
3 pairs of basal veins only, becoming quite con-
fluent at maturity; no indusium; paraphyses co-
pious, pale, much longer than sporangia. Scales
and hairs: main rachis and pinna-rachis minutely
warty and glabrescent beneath or bearing small
setiferous scales; costae near base bearing very
narrow dark-setiferous scales; costules bearing
pale bullate scales often with dark setae near their
apices; no hairs on lower surface; no hairs on
upper surface of costules and veins.
Type specimen: SCORTECHINI S.7., Perak (BM;
dupl. at K, SING).
Distr. Malaysia: Sumatraand Malay Peninsula.
Ecol. In forest, 900-1400 m.
177. Cyathea agatheti Ho_trum, Kew Bull. 16
(1962) 51.
Trunk hardly 5 cm high. Stipe 35-75 cm, dark
towards base which is covered with scales, slightly
warty where scales have fallen, distal part and
rachis pale (green when living), smooth and gla-
brous; scales on base of stipe to 10 by 2-214 mm,
light brown with somewhat paler edges which
bear many rather long dark setae. Lamina 50-60 cm
long; pinnae distinctly articulate to rachis, lowest
somewhat reduced, largest 18-25 cm long. Pin-
nules to 35 by 10 mm, abruptly narrowed at apex,
lobed to within 1 mm of costa, lowest on stalks
1 mm long; costules 3 mm apart; lamina-segments
thin, almost entire, sinuses narrow; veins 4—5 pairs,
simple. Sori medial on veins; no indusia; para-
physes dark, shorter than sporangia. Scales and
hairs: lower surface of costae bearing scattered
spreading pale hairs, and a very few hairs on
costules; scales very few, only seen on young frond,
narrow, pale, ciliate; upper surface of costae,
costules and veins bearing scattered long spreading
FLORA MALESIANA
[ser. ly vole
hairs in addition to the usual antrorse hairs on
costae.
Type specimen: KosTERMANS 12870, W. Kutai,
E. Borneo (BO; dupl. at K, L).
Distr. Malaysia: E. Borneo (one collection).
Ecol. In Agathis-forest on water-logged white
acid sand, 600 m.
178. Cyathea squamulata (BL.) Cope. Philip. J.
Sc. 4 (1909) Bot. 37; HoLttrum, Rev. Fl. Mal. 2
(1954) 122, fig. 49.—Gymnosphaera squamulata
BL. En. Pl. Jav. (1828) 243; CopreL. Gen. Fil.
(1947) 99; Fern FI. Philip. 2 (1960) 235.—AI-
sophila squamulata Hook. Sp. Fil. 1 (1844) 51,
p.p.; Metr. Ann. Mus. Bot. Lugd.-Bat. 1 (1863)
52; Racis. Fl. Btzg 1 (1898) 33; v. A. v. R.
Handb. (1908) 33; Suppl. (1917) 56.—Al/sophila
comosa WALL. ex Hook. Sp. Fil. 1 (1844) 53;
Syn. Fil. (1865) 41; Bepp. Ferns Br. Ind. (1865)
pl. 84; Handb. (1883) 13.—Alsophila laeta KUNZE,
Bot. Zeit. 4 (1846) 476, p.p.—Alsophila oligosora
Mia. Verh. Kon. Ned. Inst. Wet. 3, pt 4 (1851) 43.
—Alsophila ridleyi BAk. Ann. Bot. 8 (1894) 122;
v. A. v. R. Handb. (1908) 32.—C. ridleyi COPEL.
Philip. J. Sc. 4 (1909) Bot. 36.—C. brooksii
CopeL. ibid. 6 (1911) Bot. 135, pl. 16, not C.
brooksii MAXON, 1904.—C. paraphysata COPEL.
I.c. 135, pl. 15.—Alsophila sarawakensis C. CHR.
Ind. Fil. Suppl. (1913) 5.—Alsophila xantholepia
v. A. v. R. Bull. Jard. Bot. Btzg II, n. 23 (1916)
1; Handb. Suppl. (1917) 489, not Alsophila
xantholepis CHRIST, 1899.—Alsophila paraphysata
v. A. v. R. Handb. Suppl. (1917) 58.—Alsophila
allocota v. A. v. R. Bull. Jard. Bot. Btzg III, 5
(1922) 180.—Alsophila glabrescens v. A. Vv. R. Le.
181.—C. deuterobrooksii Cope. Philip. J. Sc. 38
(1929) 131.—C. allocota Domin, Acta Bot. Bohem.
9 (1930) 89.—C. glabrescens Domin, I.c. 119.—
C. xanthina Domin, /.c. 172.—Alsophila xanthina
C. Cure. Ind. Fil. Suppl. 3 (1934) 24.—Gymno-
sphaera sarawakensis CopeL. Gen. Fil. (1947) 99.
—Fig. 8c, 29c-d.
Trunk to c. 2 m. Stipe 40-60 cm, densely and
persistently scaiy for most of its length (main
rachis sometimes also persistently scaly); scales
firm, medium brown, largest 30 by 2-3 mm
(rarely to 40 by 4 mm), edges closely set with
dark setae. Frond to c. 150 cm long. Pinnae:
lowest somewhat reduced, variable; largest 50 cm
long. Pinnules commonly to 80 by 15 mm, largest
seen 100 by 20 mm, lower ones on stalks 1-2 mm
long, apex shortly acuminate, edges lobed 14-74
distance towards costa, no free basal segments
but pinnules on exceptionally large pinnae some-
times lobed nearly to costa at base; costules 314—
414 mm apart; veins 6-9 pairs, of smaller pinnules
mostly simple, of larger ones mostly forked;
lamina-segments rather thin, edges almost entire,
apices rounded and asymmetric. Sori a little
nearer to costule than to edge; no indusia; pale
paraphyses usually longer than sporangia. Scales
and hairs: pinna-rachis glabrescent beneath, finely
warty, sometimes with residual small setiferous
scales; on costae, near base, usually narrow flat
brown strongly setiferous scales, grading to bul-
Dec; 1963]
CYATHEACEAE (Holttum)
153
late ones distally; on costules pale bullate scales,
the larger ones acuminate and setiferous near
apices; a few stout hairs on upper surface of
costules.
Type specimen: KUHL & VAN HASSELT, Pasir
Ipis, W. Java (L).
Distr. Malaysia: Sumatra, Malay Peninsula,
Java, Borneo, S. Philippines (Sulu Arch.).
Ecol. A small tree-fern of forest, not in open
places, in lowlands and to c. 1500 m (rarely above
1000 m in the Peninsula); specimens from 2500 m
have been found on Mt Kinabalu, N. Borneo, and
are small, with exceptionally scaly rachis, but do
not appear to differ significantly in other ways.
Note. The type specimens of Alsophila comosa
and A. ridleyi are both from Singapore I. The
latter is only distinct in smaller size and less deeply
lobed pinnules. Similar differences characterize
the other species reduced to synonymy. The type
collection of C. sqguamulata includes a small frond
only 60 cm long in all, with pinnae to 8 cm long
and very few free pinnules, but fertile.
4b. Subsection Sarcopholis
Ho.ttum, subsect. nov.—Fig. 30.
A subsectione Schizocaena differt: paleis stipitis carnosis ascendentibus, apices
versus planis setiferisque.
Type species: Cyathea rosenstockii BRAUSE.
Distr. Malaysia: Moluccas and New Guinea to Polynesia.
Taxon. Fleshy upcurved bases of stipe-scales (fig. 30) are the distinctive character of this subsection;
distally these fleshy bases are more or less abruptly flattened to scales which have setiferous margins. The
flat distal parts are often eroded from herbarium specimens; a careful morphological and developmental
study from good fresh material is needed. Some species have the fleshy bases less developed, and I am
not sure that there is a sharp distinction from subsect. Schizocaena. The scale-bases in C. pulcherrima
CopEL. (subsect. Sphaeropteris) are comparable developments, but they are slender, rigid, and spread at
right angles to the surface of the stipe.
KEY TO THE SPECIES
1. Pinna-rachis hairy on the lower surface.
2. Scales on costae setiferous.
3. Few or no bullate scales. Pinnules lobed to 2 mm from costa .
3. Bullate scales present. Pinnules lobed to within 1 mm from costa .
2. Scales on costae not setiferous.
1. Pinna-rachis not hairy on lower surface.
4. Indusium quite lacking.
5. Pinnules almost sessile, to 28 mm wide .
5. Pinnules long-stalked, to 45 mm wide .
4. Indusium present.
6. Pinnules on stalks to 6 mm long. Stipe to 60 cm . ‘
6. Pinnules sessile or on much shorter stalks. Stipe much shorter.
7. Pinnules to 35 mm wide, larger ones with lowest 1-2 lamina-segments free. No bullate scales.
185. C. inaequalis
7. Pinnules rarely over 25 mm wide, lowest segments not free. Bullate scales usually present on cos-
tules.
8. Pinnules lobed only in basal half .
8. Pinnules deeply lobed throughout.
9. Pinnules not over 20 mm wide, lobed to c. 3 mm from costa. . eae 37-7 © werner
9. Pinnules on largest pinnae in most cases over 20 mm wide, lobed to 1-2 mm from costa.
10. Scales on costae and costules bullate, mostly entire. ...... . 188. C. insularum
10. Scales on costae mostly small and fringed or setiferous; bullate scales on costae and costules
mostly fringed.
11. Pinnules of largest pinnae 22-28 mm wide.
12. Segments of lamina narrowed rather evenly from base to pointed apex, sinuses thus triangu-
lar. Scales on costae not setiferous; bullate scales few. Lowest pinnae not greatly reduced.
189. C. tripinnatifida
12. Segments of lamina with rounded apex. Scales on costae often setiferous, bullate scales
usually present. Lowest pinnae small, stipe very short . 190. C. runensis
11. Pinnules of largest pinnae less than 20 mm wide . 191. C. moseleyi
179. C. fusca
. 180. C. setifera
181. C. rosenstockii
182. C. marginata
183. C. mesosora
184. C. papuana
186. C. parvipinna
154
FLORA MALESIANA
[ser. II, vol. 1?
179. Cyathea fusca BAK. in Beccari, Malesia 3
(1886) 31; v. A. v. R. Handb. (1908) 19; Suppl.
(1917) 25; CopeEL. Philip. J. Sc. 77 (1947) 100.
Trunk to 2 or 3 m; fronds to 150 cm long.
Stipe to 15 cm, bearing thick fleshy scales which
have dark setae on their edges (at least when
young); rachis bearing similar scales near base,
also short hairs on lower surface. Pinnae: lowest
c. 8 cm long, largest 40 cm. Pinnules to c. 90 by
20 mm, lobed to 2-3 mm from costa, lowest seg-
ment not free; costules 5 mm apart; lamina-
segments firm, almost entire, apices rounded;
veins c. 6 pairs. Sori medial, indusiate; indusium
complete, pale, breaking and largely persistent.
Scales and hairs: pinna-rachis densely hairy on
lower surface, hairs spreading, c. 14 mm long;
some similar hairs on bases of costae; scales on
costae very small, bearing long dark setae; a few
similar scales on costules, none bullate; no hairs
on upper surface of costules and veins.
Type specimen: D’ALBERTIS s.n., Fly R., Papua
(K).
Distr. Malaysia: Eastern New Guinea.
Ecol. Casual in undergrowth of forest near
river (BRASS).
Note. This species is very near C. werneri
ROsENST., but appears to differ from it in presence
of abundant hairs on lower surface of pinna-rachis
and in absence of bullate scales.
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PERRIER
eo 4
Fig. 30. Cyathea rosenstockii BRAusE. Base of
stipe, showing fleshy scales; scale in cm (PULLE
509, Mt Perameles, 1100 m, BM).
180. Cyathea setifera Hottrum, Kew Bull. 16
(1962) 62.
Stipe 10 cm, covered with thick dark pale-edged
scales 20 by 1144 mm, their edges (at least near
apex) bearing dark setae; also covered, between
the large scales, with a thin dark felt of small
scales of irregular shape, some with dark setae;
scales of very young fronds very thick at their
bases which have internal air-spaces. Rachis
bearing scattered narrow dark scales 10 mm long
and also thick hairs, and the remains of a felt of
small scales as on the stipe. Pinnae: lowest about
4cm long, largest 40 cm. Pinnules to 70 by 16 mm,
sessile, very shortly acuminate, lobed nearly to
the costa; basal 1-2 segments contracted at base;
costules 4-414 mm apart, at 60° to the costa;
lamina-segments rigid but not very thick, entire
or slightly crenulate, ends rounded; veins 6—7
pairs, strongly raised on lower surface, forked
rather far from costule. Sori nearer to edge than to
costules; indusium thin and pale, breaking and
persistent; paraphyses dark, longer than sporangia.
Scales and hairs: upper part of main rachis and
pinna-rachis bearing thick pale hairs on iower
surface, also scattered dark strongly setiferous
scales of various sizes, to 5 mm long; costae
densely covered beneath with elongate brown
setiferous scales, grading to brown bullate scales
setiferous near their apices; bullate scales distally
on costae and on costules brown, pale-fringed at
apices; hairs on upper surface of costules few
(not more than | on a costule).
Type specimen: MAIN & ADEN 1306, Morotai,
N. Moluccas (BO; dupl. at K, L).
Distr. Malaysia: Moluccas (Morotai),
collection.
Ecol. At 1000 m.
one
181. Cyathea rosenstockii BRAUSE, Bot. Jahrb. 56
(1920) 49; Cope. Philip. J. Sc. 77 (1947) 101.—
Fig. 30.
Trunk 114-2 m, bearing 6-8 fronds. Stipe
10-15 cm, covered with thick ascending fleshy
scales to 45 by 4 mm wide at base; main rachis
almost smooth, glabrescent, pale. Pinnae: lowest
3-5 cm long, increasing upwards, largest 30 cm.
Pinnules to 85 by 23 mm, sessile, lowest segment a
little constricted at base, rest of pinnule lobed to
1 mm from costa; costules 5-544 mm apart,
distinctly oblique; lamina-segments rigid, edges
slightly crenate, sinuses wider in fertile than in
sterile pinnules; veins 7-10 pairs, much raised on
upper surface, less so beneath. Sori nearer costule
than edge; indusium thin, at first complete, later
breaking and sometimes almost disappearing.
Scales and hairs: lower surface of pinna-rachis
throughout bearing many pale thick crisped hairs;
scales on lower surface of costae and costules dark,
bullate, sometimes very few.
Type specimen: LEDERMANN 11264, Sepik area,
E. New Guinea (B).
Distr. Malaysia: New Guinea.
Ecol. In mossy forest, or rain forest, at 1300—
1750 m.
Dec. 1963]
182. Cyathea marginata (BRAUSE) Domin, Acta
Bot. Bohem. 9 (1930) 134.—Al/sophila marginata
BraAuse, Bot. Jahrb. 56 (1920) 63.
Trunk to 3 m tall; fronds 150-200 cm long.
Stipe to at least 35 cm, warty, with one pair of
reduced pinnae near base; stipe-scales not seen.
Pinnae: lowest 18 cm long, largest 42 cm. Pinnules
to 125 by 28 mm, sessile, acuminate, lobed to 2-3
mm from costa, lowest segment not free; costules
6-7 mm apart; lamina-segments rather thick and
rigid, crenulate, apices rounded, sinuses narrow;
veins 11-12 pairs. Sori medial; no indusium; re-
ceptacle large. Scales and hairs: pinna-rachis
beneath light brown, smooth, with a few persistent
pale thin scales bearing irregular long flexuous
setae; costal scales few, thin, ovate, bearing long
dark setae; no scales seen on costules.
Type specimen: LEDERMANN 12586, Sepik region,
E. New Guinea (B).
Distr. Malaysia: E. New Guinea (one collec-
tion).
Ecol. In mountain forest at 1400-1500 m.
183. Cyathea mesosora Ho_truM, Kew Bull. 16
(1962) 57.
Trunk slender, to 3 m. Fronds few, 150-220 cm
long. Stipe to 10 cm, dull, spines to 1 mm, rather
abundant; scales to c. 15 by 114 mm, slightly
thickened at base, dark, edges bearing irregular
flexuous setae or hairs. Pinnae: lowest 3—5 cm
long, simply pinnate (sometimes widely separated
from next); largest 42 cm long. Pinnules widely
spaced, to 110 by 45 mm, lowest on stalks to 8 mm
long, lobed throughout to 3-4 mm from costa;
costules 714-9 mm apart; lamina-segments very
firm, edges crenate (more strongly towards apex)
and thickened, apex rounded; veins 10—12 pairs on
largest pinnules, mostly rather narrowly forked,
strongly raised on lower surface. Sori medial,
usually at the fork of a vein, sometimes one sorus
in middle of each branch of lowest vein; no
indusium; receptacle large, often very broad;
paraphyses short, pale, slender. Scales and hairs:
pinna-rachis beneath smooth and glabrous; no
scales seen on lower surface of costae and costules;
minute hairs (bases of former scales ?) sometimes
abundant on lower surface of veins; no hairs on
upper surface of costules and veins.
Type specimen: CARR 15720, Lala River, Papua
(BM; dupl. at K, L, MICH).
Distr. Malaysia: Eastern New Guinea (three
collections).
Ecol. In forest at 1400-1750 m.
184. Cyathea papuana (RIDL.) v. A. v. R. Handb.
Suppl. (1917) 487; Copev. Philip. J. Sc. 77 (1947)
121.—Alsophila papuana Riv. Trans. Linn. Soc.
Il, Bot. (1916) 252.—Gymnosphaera papuana
CopeL. Gen. Fil. (1947) 98.
Stipe to 60 cm, reduced basal pinnae lacking;
spines and scales as C. mesosora. Pinnae to more
than 40 cm long. Pinnules to 90 by 22 mm, lobed
to 3 mm from costa, lowest on stalks to 6 mm long;
costules 5-7 mm apart; lamina-segments firm,
edges thick, crenate; veins to 8 or 9 pairs, thick
CYATHEACEAE (Holttum)
1655
and raised on lower surface. Sori medial, indusiate;
indusium thin and translucent, at first complete,
breaking and in part persistent; paraphyses short,
thin, pale. Scales and hairs: residual scales on
pinna-rachis and costae few, narrow with irregular
long dark setae; no hairs on upper surface of
costules and veins.
Type specimen: BopEN KLoss, Mt Carstensz,
Camp III, Jan. 1913, W. New Guinea (BM; dupl.
at K).
Distr. Malaysia:
collections).
Ecol. At 700-1100 m.
West New Guinea (two
185. Cyathea inaequalis Ho_tTumM, Kew Bull. 16
(1962) 56.
Trunk 4-6 m, 5-6 cm @g; leaf-scars in alternate
whorls of 5. Stipe 24 cm, copiously thorny and
scaly near the base; thorns | mm, acute; scales
ascending, fleshy at their bases and thinner distally,
to 20 by 114 mm, dark and shining except for the
thin pale edges of the distal part which bear dark
setae; scales above base of stipe very small, brown,
the larger ones setiferous. Pinnae: lowest 8 cm
long, pinnatifid, largest 50 cm long. Pinnules to
130 by 35 mm, lowest with stalks 3 mm long,
apex acuminate, base very unequal (acroscopic
segment much larger than basiscopic), 1-2 basal
segments of largest pinnules free or nearly so, rest
of pinnule lobed to 2 mm from costa; costules
71-814 mm apart; lamina-segments thin, almost
entire, apex bluntly pointed and slightly falcate;
veins to 9 pairs. Sori medial; indusia complete,
thin, breaking and persistent. Scales and hairs:
pinna-rachis minutely warty beneath, glabrescent;
near base of costae a few small flat elongate seti-
ferous scales; no scales seen on costules; no hairs
on lower surface of pinnules, nor on upper surface
of costules.
Type specimen: Brass 23547, Mt Dayman,
Milne Bay Distr., Papua (L; dupl. at A).
Distr. Malaysia: E. New Guinea
collected).
Ecol. In rain forest ravine, at 700 m.
(once
186. Cyathea parvipinna HOLTTUM, Kew Bull. 16
(1962) 60.
Trunk 1 m tall. Stipe 20 cm; spines on stipe and
basal part of rachis many, sharp, 1 mm; scales on
stipe sparse, to 10 by 2 mm, thick at base, nar-
rowed and thinner distally with paler edges bearing
dark setae; also on stipe very small brown scales,
the larger setiferous. Pinnae: lowest 31 by 1 cm,
on stalks 8 mm long, lamina simple; largest 40 cm
long. Pinnules to 90 by 18 mm, sessile, acuminate,
apical half subentire, basal half gradually more
deeply lobed, at base lobed more than half-way to
costa, lobes almost entire, thin; costules 5 mm
apart; veins in basal lobes 6-7 pairs. Sori medial;
indusia thin, translucent, breaking and in part
caducous; paraphyses pale. Scales and hairs:
pinna-rachis glabrescent beneath, residual scales
small, brown, sparingly setiferous; a few rigid
brown setiferous scales at bases of costae; on
distal part of costae small dark thick bullate scales;
no scales seen on costules; no hairs on lower
156
FLORA MALESIANA
[ser. Ik yole
surface of costae and none on upper surface of
costules.
Type specimen: BrAss 25837, Normanby I.,
Papua (L; dupl. at K, US).
Distr. Malaysia: E. New Guinea (once
collected).
Ecol. On banks of stream in forest at 270 m.
187. Cyathea werneri ROSENST. in Fedde, Rep. 5
(1908) 34; v. A. v. R. Handb. (1908) 786; CopeL.
Philip. J. Sc. 77 (1947) 101.—C. kingii ROSENsT.
in Fedde, Rep. 9 (1911) 422, non (CLARKE) Co-
PEL. 1909; v. A. v. R. Handb. Suppl. (1917) 25;
CopeEL. Philip. J. Sc. 77 (1947) 100.
Trunk to 314 m, bearing c. 8 fronds to 2144 m
long. Stipe 8-25 cm; scales to 20 by 144 mm,
thick at base, pale and thin distally with setae on
the edges. Pinnae: lowest 3-8 cm long, longest to
at least 50 cm. Pinnules to 100 by 20 mm, sessile,
acuminate, lobed to c. 3 mm from costa; costules
514-614 mm apart; lamina-segments almost entire,
distinctly narrowed from the base so that sinuses
are triangular. Sori medial; indusium complete,
thin, breaking and persistent; paraphyses dark.
Scales and hairs: pinna-rachis glabrescent beneath
or with small residual scales, not hairs; scales on
costae near base small, bearing some dark setae,
usually grading to bullate scales distally and on
costules; a few thick hairs sometimes near apex of
costae on lower surface.
Type specimen: WERNER 66, Damun, NE. New
Guinea (S-PA; dupl. at P).
Distr. Malaysia: S. & E. New Guinea.
Ecol. In forest at 200-1200 m.
Notes. WAKEFIELD has noted that plants are
often fertile when juvenile (i.e. bearing only
simply pinnate fronds). Some specimens lack
bullate scales, and are then intermediate between
this species and C. fusca; possibly the two should
be united.
188. Cyathea insularum HoL_trum, Kew Bull. 16
(1962) 57.
Trunk 3-5 m, 314-6 cm @ after decay of leaf-
bases; leaf-scars 18-25 mm g, in 3 rather steep
spirals. Fronds numerous, 165—200 cm long. Stipe
10-15 cm, near base covered with scales; scales
10-15 by 1 mm, dark and thick at the base, distal
part dark with pale edges bearing dark setae;
rest of stipe sparsely covered with very small pale
short-fringed scales. Lowest pinnae less than 5 cm
long, rest gradually larger, largest 45 cm. Largest
pinnules 65-90 by 18-22 mm, sessile, acuminate,
lobed to 1-2 mm from costa; costules 5-6 mm
apart; lamina-segments slightly crenate towards
obtusely pointed apex; veins 8—9 pairs. Sori medial;
indusium complete, thin, pale, breaking and
persistent. Scales and hairs: pinna-rachis below
bearing small thin pale finely fringed scales; scales
on costae throughout pale, bullate, rather large,
mostly entire; similar scales on costules; a few
thick hairs towards apex of costa on lower surface,
no hairs on upper surface of costules.
Type specimen: Brass 27419, Misima Island,
Louisiades (L).
Distr. Malaysia: Louisiade Arch. (on 4 islands).
Ecol. In forest, near streams, at 100-350 m.
189. Cyathea tripinnatifida Roxs. Calc. J. Nat.
Hist. 4 (1844) 518.—C. integra J. SM. ex Hook.
Ic. Pl. (1844) t. 638, p.p. excl. ic.—C. nigrospinulosa
v. A. v. R. Bull. Jard. Bot. Btzg II, n. 28 (1918) 15.
Stipe to 25 cm (sometimes much shorter ?),
bearing scattered thick scales to 20 by 114 mm,
setiferous near their apices; stipe also covered with
very small dull fringed scales. Pinnae to at least
55 cm long. Largest pinnules 90-110 by 20-28 mm,
lobed to 114-2 mm from costa, sessile, short-
acuminate; costules 6-7 mm apart; lamina-
segments thin but firm, narrowed from the base to
broadly pointed apex and so separated by tri-
angular sinuses, edges crenate to almost entire;
veins to 8 or 9 pairs. Sori nearer to costule than to
edge; indusia rather thin, at first complete, break-
ing and persistent; paraphyses dark, as long as
sporangia. Scales and hairs: \ower surface of
pinna-rachis bearing small pale fringed scales,
hairs lacking; scales on costae small, fringed with
pale hairs, not bullate; a few bullate scales on
costules, sometimes caducous; a few thick hairs
sometimes present on lower surface of costae and
costules near apex of pinnule.
Type specimen: Herb. Wallich n. 7076, Mo-
luccas (CAL ?; dupl. at K, BM).
Distr. Malaysia: Moluccas (Ambon).
Ecol. In lowland forest.
Notes. Hooker cited a specimen of this species,
from Ambon, with his original description of
C. integra, but his illustration was prepared from
a Philippine specimen, which is thus taken as type
of C. integra.
Most specimens of C. tripinnatifida lack the
stipe. One from Ambon, collected by REINWARDT
(L) has a stipe at least 25 cm long, and does not
show lower pinnae.
190. Cyathea runensis v. A. v. R. Bull. Dép. Agr.
Ind. Néerl. 7. 18 (1908) 1; Handb. (1908) 22.—
C. versteegii CHRIst, Nova Guinea 8 (1909) 161;
v. A. v. R. Handb. Suppl. (1917) 25.
Stipe short, warty; scales dull, dark, more than
1 cell thick at base not fleshy as in C. rosenstockii,
Lowest pinnae short, gradually increasing upwards.
largest at least 50 cm long. Pinnules to 120 by 25
mm (rarely to 30 mm), lowest segment of largest
pinnules free or nearly so, rest lobed to 2 mm
from costa; costules 6-7 mm apart; segments of
lamina almost entire or slightly crenate, apices
rounded, separated by sinuses 1-2 mm wide;
veins to 8 pairs. Sori medial; indusium complete,
thin, persistent, breaking irregularly; paraphyses
dark, as long as sporangia. Scales and hairs:
pinna-rachis glabrescent on lower surface, hairs
on upper surface short and dark; scales on
lower surface of costae small, brown, of rather
irregular shape, edges set closely with short
spreading rather stiff concolorous hairs or darker
setae; bullate scales on costules few, small.
Type specimen: TEYSMANN, Pulu Roon, W. New
Guinea (BO; dupl. at L).
Dec. 1963]
CYATHEACEAE (Holttum)
iS7
Distr.Malaysia:W.New Guinea, Bismarck Arch.
Ecol. In lowland forest.
Note. This species is very close to C. tripinna-
tifida, and I am not sure of a clear distinction. It
seems probable however that a short stipe is in-
variable in C. runensis, that of C. tripinnatifida
being longer, but very few specimens show this
character.
191. Cyathea moseleyi BAK. J. Linn. Soc. Bot. 15
(1876) 104.
Stipe short; stipe and lower part of rachis
bearing scales to 20 by 114 mm, dark and some-
what thickened in the middle with pale edges
bearing dark setae. Pinnae: lowest gradually
reduced, largest to at least 45 cm long. Largest
pinnules to 80 by 18 mm, lobed to c. 1 mm from
costa, sessile, acuminate; costules 5-6 mm
apart; lamina-segments rather thin, edges more
or less crenate, apices bluntly pointed; veins 8—10
pairs. Sori medial; indusium pale, thin, at first
complete, breaking and persistent; paraphyses dark.
Scales and hairs: pinna-rachis beneath smooth,
usually with some residual very small dull brown
fringed scales; small fringed scales present on
lower surface of costae, with small fringed bullate
scales, the latter also abundant on lower surface of
costules; a few thick hairs present on upper sur-
face of costules but not on lower surface.
Type specimen: MoseLey, Admiralty Is (K).
Distr. Malaysia: Admiralty Is, Bismarck Arch.
Ecol. In lowland forest.
Note. C. brackenridgei Metrt., of the Solomon
Is, appears to be closely related to this species,
but to differ in wider pinnules (to 24 mm wide) and
in very abundant scales throughout the stipe and
lower part of rachis.
Doubtful species
Alsophila hallieri RosENst. Med. Rijksherb. 7. 31
(1917) 2.—C. hallieri Domin, Acta Bot. Bohem.
9 (1930) 120.
The specimens of HALLIER (735, 737, 738, 4726,
4727) cited with the description have not been
found in the Rijksherbarium at Leiden. The brief
description does not permit the species to be
identified with certainty.
Alsophila matthewii Curist, J. Linn. Soc. Bot. 39
(1909) 213.—C. matthewii Doin, Pterid. (1929)
263.
I have only seen the specimen from the type
collection at Kew; it is a small sterile frond,
evidently from an immature plant, said to have
been borne on a slender trunk 60 cm high. As the
specimen was found in a much-collected locality
(Mt Maquiling, Luzon, at 1100 m), it probably
belongs to a species already known, possibly
C. heterochlamydea CopeL., but I am very uncer-
tain of this.
Alsophila philippinense Hort. Veitch, Gard. Chron.
II, 4 (1875) 179, nomen.—C. veitchiana Domin,
Acta Bot. Bohem. 9 (1930) 168.
A cultivated plant, described as follows: ‘“‘a
handsome new fern, with regularly crimped pinnae
from the Philippine Islands”. There is no specimen
in Kew Herbarium, and no reference to the plant
in Hortus Veitchii.
Alsophila speciosa [non (MEYEN) PRESL] GOLD-
MANN, Nova Acta 19, Suppl. I (1843) 465.
The type of MEYEN’s species came from Brazil:
GOLDMANN’s brief Latin description of a Phi-
lippine fern is inadequate for identification.
Cyathea adenochlamys CurisT, Bull. Herb. Boiss.
II, 6 (1906) 1008; v. A. v. R. Handb. (1908) 785.
I have not found the type specimen at Paris.
Curist described the indusia as persistent, rigid,
green, covered densely with pale sessile glands,
opening transversely; I have not seen any Cyathea
with such indusia.
Cyathea grata Domtn, Acta Bot. Bohem. 9 (1930)
120, nomen.
Under this name is cited C. frondosa ROSENST.
in Fedde, Rep. 12 (1913) 163, but no such species
is there described, nor have I found any other
reference to C. frondosa ROSENST. Provenance New
Guinea.
Species not occurring in Malaysia
Alsophila truncata BRACK. in Wilkes, U.S. Expl.
Exped. 16 (1854) 289; v. A. v. R. Handb. (1908)
42.
Malaysian specimens referred to this species are
C. celebica BL. or an allied species.
Cyathea aneitensis Hook. Syn. Fil. (1865) 26;
v. A. v. R. Handb. (1908) 16.
After citing specimens from Aneitijum, HOOKER
added “‘Ternate, Herb. Hort. Calc.’ There is no
Ternate specimen at Kew, and I have seen no Ma-
laysian specimens referable to this species.
Cyathea rumphii Desy. Mém. Soc. Linn. Paris 6
(1827) 323.—Polypodium arboreum Lour. FI.
Coch. (1790) 831, non LINN.
DESVAUX published a new name for LOUREIRO’s
species, without description; like LouRErIRo, he
also cited a plate by RUMPuHius. LouREIRO described
a fern from Indo-China, not Ambon; neither
his description, nor RuMputus’s plate is clearly
identifiable. Therefore I cannot agree with MER-
RILL’s suggestion that the name C. rumphii DEsv.
should replace C. amboinensis (v. A. v. R.) MERR.
(see Trans. Am. Phil. Soc. Philad. n.s. 24, 1935,
Sif):
Excluded from Cyathea
Alsophila dielsii BRAUSE, Bot. Jahrb. 56 (1920) 67.
—C. dielsii Domtn, Acta Bot. Bohem. 9 (1930) 111.
The type specimen belongs to the genus Di-
plazium.
158
FLORA MALESIANA
[ser. Il, volume
Alsophila dryopteroidea BRrAuseE, Bot. Jahrb. 56
(1920) 70.—C. atrispora Domin, Acta _ Bot.
Bohem. 9 (1930) 95.—Dryopteris atrispora C.
Cur. Brittonia 2 (1937) 296.—Lastrea dryopteroi-
dea CopEL. Gen. Fil. (1947) 138.
The type is a Thelypteroid fern; generic limits
in this group need to be clarified.
Alsophila gazellae KUHN, Forschungsr. Gazelle 4
(1889) 13.—C. gazellae Domin, Pterid. (1929) 262.
I have seen the type specimen, and identified it
as Pleocnemia cumingiana PRESL (see Reinwardtia
1, 1951, 188).
Alsophila tristis (BL.) BL. ex Moore, Ind. Fil.
(1857) 58, based on Aspidium triste BL. En. Pl.
Alsophila warburgii Curist, Ann. Jard. Bot. Btzg
15 (1897) 80, from Celebes = Dryopteris sp.
Alsophila xantholepis Curist ex Digs in E. & P.
Pfl. Fam. 1, 4(1899) 138; v. A. v. R. Handb. (1908)
44.—C. xantholepis Domin, Pterid. (1929) 263;
Acta Bot. Bohem. 9 (1930) 172 = Dryopteris sp.
Cyathea aspidioides (BL.) ZoLt_. & Mor. Syst.
Verz. (1846) 108 = Diacalpe aspidioides BL.
Cyathea woodlarkensis CopeL. Philip. J. Sc. 9
(1914) Bot. 1; ibid. 77 (1947) 124.—Alsophila
woodlarkensis C. Cur. Ind. Fil. Suppl. 2 (1917) 4.
This is a large species of The/ypteris, apparently
Jav. (1828) 169 = Stenolepia tristis (BL.) v. A.
near 7. immersa (BL.) CHING. There are speci-
v. R. Bull. Dép. Agr. Ind. Néerl. . 27 (1909) 45.
mens in Herb. Copel. (MICH) and at Paris.
2. DICKSONIA
L’HErRIT. Sert. Angl. (1788) 30, p.p.; Curist, Farnkr. Erde (1897) 313, p.p.;
DieLs in E. & P. Pfl. Fam. 1, 4 (1899) 119; UNDERW. Mem. Torr. Bot. Cl. 6 (1899)
259, 278; C. Cur. Ind. Fil. (1905) xvi, 220; MAxon, J. Wash. Ac. Sc. 12 (1922)
454; Copet. Gen. Fil. (1947) 48.—Balantium KAuLr. En. Fil. Chamisso (1824)
288, p.p. typ.; PRESL, Tent. Pterid. (1836) 134 (excl. Balantium brownianum).—
Dicksonia subg. Balantium Hook. Sp. Fil. 1 (1844) 66, p.p.—Dicksonia § Eudickso-
nia HooK. & BAK. Syn. Fil. (1868) 50, p.p.—Fig. 31.
Stem arborescent, bearing numerous fronds in several spiral series, the vascu-
lar tissue supported both internally and externally with strong bands of scleren-
chyma; leaf-bases usually persistent; young leaves, and at least the bases of
stipes, persistently covered with long hairs; stipes usually short, vascular system
complex, in two continuous or divided transverse arcs concave adaxially; pinnae
numerous, the lower ones gradually reduced; pinnules usually bearing free deeply
lobed tertiary leaflets which are dimorphous, sterile and fertile; upper surfaces of
rachises and axes of leaflets raised, bearing rather stiff antrorse hairs, lower sur-
faces variously hairy; fertile leaflets more deeply lobed than sterile and with re-
duced lamina, each lobe bearing one sorus at the end of the vein or of the basal
acroscopic branch if the vein is branched; sorieach protected by a reflexed marginal
lobe of the lamina and by a somewhat thinner inner indusium joined to the re-
ceptacle on its basiscopic side (fig. 31c); free surface of receptacle slightly elevated
and distinctly elongated transverse to the end of the vein; paraphyses numerous,
hair-like, multiseptate, each with a red-brown terminal cell; sporangia distinctly
stalked, with complete oblique annulus indurated round the base, the stomium
lateral; spores trilete, surfaces variously sculptured or almost smooth, exospore
more or less thickened at the angles.
Type species: Dicksonia arborescens HERItT. (St Helena).
Distr. About 25 spp. St Helena; S. America and north to Mexico; New Zealand, E. Australia and
Tasmania, New Caledonia, Samoa, Fiji; in Malaysia: New Guinea, Celebes, Philippines, N. Borneo,
Java, and Sumatra.
Morph. The trunk of Dicksonia is closely similar in form and anatomy to that of Cyathea, from which
Dicksonia differs by the indument consisting entirely of hairs. For comment on the possible relationships
between the very different soral forms in the two genera, see p. 69.
Cytol. Chromosome counts have been made by BROWNLIE for two New Zealand species; both showed
159
Dec. 1963]
n = 65 (New Phytol. 56, 1957, 207). MANTON found the same number for D. arborescens (type species
of genus) in cultivation at Kew (J. Linn. Soc. Bot. 56, 1958, 84).
Taxon. The history of the varied use of the generic name Dicksonia is summarized by MAxon, /.c.
Earlier authors confused not only Dicksonia, Culcita and Cibotium, but also Dennstaedtia and other
genera now recognized as distinct. Within the genus, the discrimination of species is difficult because the
soral form is so constant and the variation in shape of tertiary leaflets as between those on larger or
smaller pinnules in one species may be considerable. The character of the hairs on the stipe appears to be
one of the clearest distinctions between species, as are scale-characters in Cyathea; but the differences
between one hair and another are less easily defined than those between different scales. The species in
New Guinea are particularly difficult to discriminate, and the present account can only be regarded as
CYATHEACEAE (Holttum)
tentative; more field observation is needed.
KEY TO THE SPECIES
1. Base of stipe clothed with dark red spreading hairs 30-50 mm long; hairs of under-coat pale, slender or
flaccid, much shorter.
2. Hairs of under-coat of stipe-base, and most hairs on lower surface of pinna-rachis and costae, pale,
slender, rigid, spreading, not flaccid. Spores smooth .
1. D. blumei
2. Hairs of under-coat of stipe-base, and smaller hairs on lower eure of pinna- -rachis (in some cases
also of costa) short, pale, flaccid, their lateral walls collapsed. Spores verrucose.
|
a=
3. Hairs on costae almost all flaccid, pale,
antrorse, matted . 2. D. mollis
3. Hairs on costae mostly rigid, each cell evenly cylindrical.
4. Hairs on costae: some flaccid, some spreading, mostly pale .
4. Hairs on costae ascending, mostly dark red .
. 3. D. sciurus
4. D. archboldii
1. Base of stipe clothed mainly with soft matted red-brown hairs, “with or r without a smaller number of
stout rigid dark red hairs.
5. Some stout rigid spreading dark red hairs present near base of stipe, and scattered also on pinna-
rachis; pinna-rachis mainly clothed with a close layer of pale matted flaccid hairs.
5. D. hieronymi
5. No stout rigid spreading dark red hairs mixed with softer hairs on stipe-base.
6. Pinna-rachis rather dark and + glabrescent; costae bearing flaccid pale hairs + mixed with thicker
dark hairs with pale bases
6. D. grandis
6. Pinna-rachis brown, persistently hairy beneath; costae bearing entirely pale flaccid hairs.
1. Dicksonia blumei (KUNZE) Moore, Ind. Fil.
(1860) 190; v. A. v. R. Handb. (1908) 47; BACKER
& PostH. Varenfl. Java (1939) 22.—Balantium
blumei Kunze, Bot. Zeit. 6 (1848) 214.—Ba-
jantium chrysotrichum Hassk. Obs. Fil. Jay. 1
(1856) 53.—D. chrysotricha Moore, Ind. Fil.
(1860) 190; Hook. & BAK. Syn. Fil. (1868) 50;
Ractis. Fl. Btzg 1 (1898) 121; Diets in E. & P.
Pfl. Fam. 1, 4 (1899) 121; Curist, Ann. Jard. Bot.
Btzg 19 (1904) 41.—Fig. 31a—c.
Trunk to 6 m; stipes to 60 cm or more, clothed
at the base with spreading red-brown shining
hairs 30-50 mm long, with an under-coat of much
finer short pale hairs; upper part of stipe and main
rachis dark, finely and closely warty, bearing
sparse short pale hairs; lamina to 300 cm long;
pinnae to 70 cm long, pinna-rachis bearing pale
spreading hairs on lower surface, with a few red
ones; pinnules to 100 by 15-20 mm; costules of
tertiary leaflets 4 mm apart; largest fertile tertiary
leaflets lobed throughout almost to the costule,
with 4-5 pairs of soriferous lobes, the lowest
lobes usually bilobulate with forked vein, sterile
lobule not longer than fertile; largest sterile ter-
tiary leaflets less deeply lobed, veins in lowest lobe
pinnate, in rest forked or simple; hairs on lower
7. D. lanigera
surface of costae and costules pale, firm (cell-walls
not collapsed), spreading; spores almost smooth.
Type specimen: ZOLLINGER 1894, Java (B?; dupl.
at P, L); also cited SPORLEDER, Java.
Distr. Malaysia: Sumatra (north to Karo
Plateau), Java, Central Celebes (SARASIN 2030).
Ecol. In mountain forest, 1500-2500 m.
2. Dicksonia mollis HoLTTUM, Kew Bull. 16 (1962)
64.—D. blumei [non (KUNZE) Moore] C. CHR.
Gard. Bull. S.S. 7 (1934) 223; CopeL. Fern FI.
Philip. 1 (1958) 84.—Fig. 31d-e.
Differs from D. blumei as follows: hairs of un-
der-coat on the stipe shorter, thicker, but flaccid
with walls collapsed when dry; hairs on lower
surface of pinna-rachis and costae more numerous,
appressed, flaccid, mostly ascending; spores bear-
ing conspicuous more or less coalescent warts.
Type specimen: ELMER 9874, Dumaguete
(Cuernos Mts) Negros Oriental, Negros (K; dupl.
at L, P, MICH, US).
Distr. Malaysia: NE. Borneo, Central Celebes
(2), Philippines (Mindanao to S. Luzon).
Ecol. Mountain forests. 1500-2000 m. The
Celebes specimen, much smaller than those from
FLORA MALESIANA
[ser. Il; vole
SS
Fig. 31. Dicksonia blumei (KUNZE) Moore. a. Part of pinna, showing sterile and fertile pinnules, x 7/,
b. part of pinnule, lower surface, showing sori and rigid hairs, < 6, c. section of sorus, * 8.—D. mollis
Ho.ttuM. d. Part of sterile pinnule, lower surface, showing flaccid hairs, < 6, e. hairs from lower surface,
< 30.—D. antarctica LABILL. f. Section of stipe (hairs omitted), < 2, g. section of pinna-rachis, < 4
(a DE VRIESE S.n., b—-c MATTHEW S.n , d SINCLAIR 9005, e ELMER 9874, f-g cult. R. B. G. Kew).
Borneo and the Philippines, was found at 2900 m,
in dry open vegetation (EYMA 959); it lacks a
stipe. It agrees better with D. mollis than with
New Guinea species (it is certainly not D. blumei),
but without more complete material its status is
doubtful.
3. Dicksonia sciurus C. Cur. Brittonia 2 (1937)
283; CopeEL. Philip. J. Sc. 78 (1949) 6.
Stipe 35 cm, the base covered with spreading
red-brown hairs 25-40 mm long, with an under-
coat of short pale flaccid hairs; upper part of stipe
and lower surface of main rachis more sparsely
clothed with similar but shorter red-brown hairs
and short flaccid pale ones; longest pinnae 60 cm
long, rachis dark brown, its lower surface bearing
rather sparse spreading reddish hairs and a more
or less abraded cover of short flaccid pale hairs;
pinnules to 150 mm long (on type not over 80 mm),
costules of tertiary leaflets 4-5 mm apart, hairs on
lower surface of costae and costules mostly pale,
some flaccid and some spreading; tertiary leaflets
to 20 mm long, fertile ones lobed almost to cos-
tule, basal lobes of largest fertile leaflets having a
bluntly-toothed sterile lobule exceeding the fertile
one, sterile tertiary leaflets less deeply lobed than
fertile, lobes mostly with pinnately arranged veins
and bluntly toothed edges.
Type specimen: Brass 4991, Mt Tafa, Papua
(BM; dupl. at BO, MICH, BRI, UC).
Distr. Malaysia: E. New Guinea (several
collections).
Ecol. Plentiful in ridge forest, at 1800-2400 m.
4. Dicksonia archboldii CoreL. Philip. J. Sc. 78
(1949) 6, pl. 1.
Dec. 1963]
Stipe 30-40 cm, base clothed with spreading
rather soft red-brown hairs 25 mm or more long,
with short flaccid pale hairs also; leafy part of
frond to c. 300 cm long; lower pinnae gradually
reduced, lowest 15 cm long, longest 65 cm, pinna-
rachis beneath dark, clothed rather sparsely with
dark red rigid hairs, the smallest ones with a pale
inflated basal cell, also short pale flaccid hairs;
pinnules to 140 mm long, costa covered beneath
with rigid flexuous antrorse dark red hairs, some
with a pale base, and on distal part a few pale
flaccid ones; costules of tertiary leaflets 5-6 mm
apart; sterile fertiary leaflets to 20 mm long,
lobed to costule at base, lobes to 6 pairs, veins
pinnate in each, edges toothed; fertile tertiary leaf-
lets lobed throughout to costule, lobes to 5 pairs,
each with a sorus, the basal ones also with an
acute sterile tooth.
Type specimen: Brass 10970, Lake Habbema,
2750 m, near Mt Wilhelmina, W. New Guinea
(MICH; dupl. at L).
Distr. Malaysia: West New Guinea, only
known from the type collection and a much
smaller one from Mt Arfak, Anggi Lakes (SLEv-
MER & VINK BW 14193).
Ecol. “Abundant in forest of lower slopes,
stem 4 m high, 13 cm o under the leaves and
thickened downwards; leaves 10, spreading’.
Note. This is near D. sciurus C. Cur. (an
earlier name) and the two should perhaps be united.
The type of D. archboldii is larger than that of
D. sciurus in all parts of the frond, and the hairs
on the costae are mostly red and ascending; the
fertile tertiary leaflets of D. sciurus have in some
cases a larger sterile portion, but this varies ac-
cording to position on the frond, and whole fronds
of each type are not available for comparison.
5. Dicksonia hieronymi BRAUSE, Bot. Jahrb. 56
(1920) 48; C. Cur. Brittonia 2 (1937) 282;
CopeEL. Philip. J. Sc. 78 (1949) 6.
Stipe 20-30 cm, covered near the base with soft
dull brown matted hairs with a few longer rigid
shining dark red-brown ones; upper part of stipe
and main rachis dark, rather persistently covered
with appressed soft brown matted hairs with a
few rigid shining ones; largest pinnae 30-60 cm
long; pinna-rachis dark brown, hairy on under
surface as main rachis; largest pinnules 70-120 mm
long, costa pale distally, dark near base, covered
beneath with flaccid + appressed light brown hairs,
some with rigid dark apical portion; largest
tertiary leaflets 10-20 mm long, fertile lobed to
the costule, fertile lobes 3—5 pairs, the larger with
forked vein and a sterile lobule which is rarely
larger than the fertile one; sterile tertiary leaflets
deeply lobed at base, less deeply upwards, larger
lobes containing pinnately branched veins, edges
of lobes almost entire.
Type specimen: LEDERMANN 12851, Sepik re-
gion, 1400-1500 m, E. New Guinea (B).
Distr. Malaysia: New Guinea (several locali-
ties) including Louisiade Arch.
Ecol. In mossy forest, at 1400-3000 m; fronds
usually few.
CYATHEACEAE (Holttum)
161
6. Dicksonia grandis ROsENsT. in Fedde, Rep. 5
(1908) 34; CopeL. Philip. J. Sc. 78 (1949) 5.—
D. schlechteri Brause, Bot. Jahrb. 49 (1912) 11:
CopeL. Philip. J. Sc. 78 (1949) 6.—D. ledermannii
BrAuse, Bot. Jahrb. 56 (1920) 46; CopeL. Philip.
J. Sc. 78 (1949) 5.
Stipe at least 30 cm, nearly black, lower part
covered with fine soft light brown hairs, longest
25 mm, all of same texture; pinnae to 100 cm long,
pinna-rachis beneath very dark, minutely warty,
more or less glabrescent, with vestiges of short
appressed hairs which are mostly lax and pale
brown, with some firm and reddish (the latter
sometimes with pale bases); pinnules to 140 mm
long, costa hairy beneath as pinna-rachis, dark at
base and paler distally; tertiary leaflets to 20 mm
long, fertile ones deeply lobed, lobes to 5 pairs,
each lobe of largest leaflets with a pinnate vein
bearing a sorus on its basal acroscopic branch, of
smaller leaflets sometimes with simple vein bearing
a sorus; sterile tertiary leaflets lobed almost to
costule at base, less deeply upwards, each lobe of
larger ones containing a pinnately branched vein,
edges of lobes almost entire.
Type specimen: WERNER 79, in forest, 1000 m,
Mt Gelu, E. New Guinea (also distributed as
RosEnsT. Fil. Novoguin. exsic. 7. 50 (B; dupl. at
ES eSUsSs Ue):
Distr. Malaysia: New Guinea (several localities).
Ecol. In forest, most specimens at 1000-1800 m;
one (PULLE 939, Mt Hellwig) from 2500 m, a
small plant but agreeing in hairiness.
7. Dicksonia lanigera HottrumM, Kew Bull. 16
(1962) 64.
Stipe to at least 30 cm, densely hairy throughout,
hairs uniform, soft, medium brown, to 15 mm,
the shorter hairs of same texture, no thick firm red
hairs present; /amina c. 150 cm long; rachis
beneath red-brown when dry, covered with matted
short hairs like those of the stipe; pinnae to 40 cm
long, pinna-rachis persistently hairy as rachis on
lower surface; largest pinnules 80 mm long, cos-
tules 514 mm apart; costa beneath brown-hairy
almost to apex; fertiary leaflets to 15 mm long,
sterile ones deeply lobed near the base and less
deeply upwards, larger lobes each with a pinnate
vein, edges toothed; fertile tertiary leaflets deeply
lobed throughout, vein in larger lobes forked,
acroscopic branch bearing a sorus, sterile part of
lobe not longer than fertile.
Type specimen: PuLLte 1010, Mt Wichman,
3000 m, SW. New Guinea (L; dupl. at BM).
Distr. Malaysia: W. & E. New Guinea.
Ecol. In forest, at 2500-3000 m.
Excluded from Dicksonia
Dicksonia ampla Bax. J. Linn. Soc. Bot. 22
(1886) 223 = Dennstaedtia ampla (BAK.) BEDD.
Dicksonia cuneata Hook. Sp. Fil. 1 (1844)
80 = Dennstaedtia cuneata (HOOK.) Moore.
Dicksonia deltoidea Hook. Sp. Fil. 1 (1844) 80
= Dennstaedtia scabra (WALL.) Moore.
162 FLORA MALESIANA [ser. II, vol. 12
Dicksonia erythrorachis Curist, Ann. Jard. Bot. Dicksonia neglecta Fe, Gen. Fil. (1850-52)
Btzg 15 (1897) 86 = Dennstaedtia erythrorachis 335 = Dennstaedtia smithii (HooK.) Moore.
(CHRIST) DIELS.
Dicksonia_ nephrolepioides Curist, Verh. Nat.
Dicksonia flaccida (Forst.) Sw. in Schrader, J. Ges. Basel 11 (1895) 241 = Nephrolepis dicksonioi-
Bot. 1800, ii (1801) 90 = Dennstaedtia flaccida des CHRIST.
(Forst.) BERNH.
Dicksonia remota Curist, Verh. Nat. Ges.
Dicksonia glabrata Ces. Rend. Ac. Napoli 16 Basel 11 (1896) 423 = Dennstaedtia remota
(1877) 24, 28 = Dennstaedtia glabrata (CEs.) (CurRisT) DIELs.
C. CHR.
Dicksonia rhombifolia Bax. J. Bot. 28 (1890)
Dicksonia gomphophylla Bax. J. Linn. Soc. Bot. 105 = Dennstaedtia rhombifolia (Bak.) C. CHR.
22 (1886) 223 = Dennstaedtia gomphophylla : ;
(BAK.) C. Cur. Dicksonia scabra WALL. ex Hook. Sp. Fil. 1
(1844) 80 = Dennstaedtia scabra (WALL.) MOorRE.
Dicksonia japonica Sw. in Schrader, J. Bot. 1800,
il (1801) 92 = Méicrolepia strigosa (THUNB.)
PRESL.
Dicksonia scandens Bui. En. Pl. Jav. (1828)
240 = Dennstaedtia scandens (BL.) Moore.
Dicksonia smithii Hook. Sp. Fil. 1 (1844) 80 =
Dicksonia kingii BEpp. Handb. Suppl. (1892) Dennstaedueaemi iit (HOO) Noort
6 = Orthiopteris kingii (BEDD.) HOLTTUM.
Dicksonia strigosa THUNB. Trans. Linn. Soc. 2
Dicksonia linearis Cav. Descr. (1802) 274 = (4794) 341 = Microlepia strigosa (THUNB.)
Tapeinidium pinnatum (CAv.) C. CHR. PRESL.
Dicksonia moluccana BL. En. Pl. Jay. (1828) Dicksonia zippeliana KUNzE, Bot. Zeit. 3 (1845)
239 = Dennstaedtia moluccana (BL.) Moore. 838 = Dennstaedtia scandens (BL.) Moore.
3. CYSTODIUM
J. Sm. in Hook. Gen. Fil. (1841) t. 96; Hist. Fil. (1875) 258.—Fig. 32.
Stem massive, usually creeping, covered with long hairs. Fronds tufted, bipin-
nate; stipe grooved on adaxial surface, vascular tissue arranged as in Dicksonia;
upper surface of pinna-rachis antrorsely hairy and grooved, the groove not open
to admit the grooves of midribs of pinnules; pinnules slightly dimorphous, sterile
ones serrate, fertile ones with slightly narrower lamina and teeth enlarged to
protect sori; veins in pinnules pinnately arranged, oblique, often forked in larger
pinnules, simple in smaller ones except for the basal pair which may be pinnate;
sori terminal on veins, receptacle slightly raised, round in section, tooth of lamina
reflexed and enlarged, forming an outer indusium, inner indusium smaller and
thinner, attached only at the base of the sorus; paraphyses abundant, as simple
multiseptate hairs lacking a terminal glandular cell; sporangia stalked, stalk as
long as body of the sporangium, annulus complete and slightly oblique but not
indurated at base where it passes the stalk; spores trilete.
Distr. Monotypic, Malaysia: from NE. Borneo to New Britain & Louisiades.
Notes. The single species agrees with Dicksonia in hairs and their distribution, in arrangement of
vascular tissue in the frond, and in form of sori; it differs from Dicksonia (1) in normally prostrate
position of stem, (2) in grooved upper surface of rachis, pinna-rachis and costules, (3) in shape of pin-
nules, which are crenate or serrate (not deeply lobed as in Dicksonia), (4) in circular transverse section
of receptacle of sorus, (5) in smaller sporangia on longer stalks, and (6) almost vertical annulus, not
indurated at the base.
JOHN SMITH (/.c. 1875) stated that the pinnules are jointed to the rachis (a character to which he at-
tached great importance), but they are not so.
Dec. 1963]
CYATHEACEAE (Holttum)
C
i2
Kee
\Qiaz
IS
.
tt,
EWS
mS
—
—
hd ty
Sy
Fig. 32. Cystodium sorbifolium (SM.) J. SM. a. Middle part of rachis with pinnae, x 24, b. lower surface
of part of pinna, near its base; sporangia removed from one sorus to show inner indusium, x 6, c.
section of stipe, x 4, d. section of pinna-rachis, <x 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. Cur. Gard. Bull. S.S. 7 (1934)
239; Ind. Fil. Suppl. 3 (1934) 66; CopeL. Philip.
J. Sc. 78 (1950) 7.—Dicksonia sorbifolia Sm. in
Rees, Cyclop. 11 (1808) n.p.; Hoox. Sp. Fil. 1
(1844) 72, t. 25A; Hook. & BAK. Syn. Fil. (1868)
52; Curist, Ann. Jard. Bot. Btzg 15 (1898) 87;
v. A. v. R. Handb. Suppl. (1917) 74.—Dicksonia
moluccana Roxs. Calc. J. Nat. Hist. 4 (1844) 517.
—Dicksonia papuana F. v. M. Descr. Not. 4
(1876) 76; CopEL. Philip. J. Sc. 6 (1911) Bot. 69.—
Saccoloma sorbifolia CHRisT, Farnkr. Erde (1897)
309; C. Cur. Ind. Fil. (1905) 612; v. A. v. R.
Handb. (1908) 282.—Fig. 32.
Stem usually creeping (once reported erect,
14 m high), densely covered like the bases of the
stipes with soft golden brown shining hairs c. 3 cm
164
FLORA MALESIANA
[ser. II, vol. 12
long. Stipes to 75 cm, glabrescent and smooth
above the base; /amina 100-250 cm long, lowest
pinnae not greatly reduced; /argest pinnae 20-30
by 5-9 cm, bearing free pinnules for 14-74 of
their length, apical part deeply lobed; sterile
pinnules 6-8 mm wide, fertile 5—6 mm wide, sep-
arated by gaps of c. 2 mm; pinnules slightly
oblique, their costules falcate, asymmetric at the
base (rounded to cordate on basiscopic side,
broadly truncate and sometimes auricled on acro-
abundant spreading hairs | mm long. Sori 114-2
mm apart, at ends of all veins on fertile pinnules
(on acroscopic branch if vein is forked). Spores
bearing on their surface a fine irregular slightly
raised reticulum.
Type specimen: C. SmitH, Honimoa, Ceram
(Herb. J. E. Smith; dupl. at K).
Distr. Malaysia: N. Borneo, N. Celebes, Mo-
luccas (Ceram), New Guinea (incl. New Britain,
Admiralty Is and Louisiade Arch.).
Ecol. In lowland forest, to 400 m, often near
rivers.
scopic side), edges almost parallel except towards
apex; lower surface of pinna-rachis, costules of
pinnae and sometimes veins bearing more or less
4. CIBOTIUM
KaAuLFuss, 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; Diets in E. & P. Pal. Fam. 1, 4 (1899) 121; ©. Gemiiaes
Fil. (1905) xvi, 183; Maxon, Contr. U. S. Nat. Herb. 16 (1912) 54; CopEL. Gen.
Fil. (1947) 49.—Pinonia GaupicH. 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 1i5= 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 . ater: 23 Cercumingi
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
CYATHEACEAE (Holttum)
Dec. 1963]
1. Cibotium barometz (L.) J. SM. Lond. J. Bot. 1
(1842) 437; Bepp. Handb. (1883) 24; Suppl.
(1892) 6; Diets in E. & P. Pfl. Fam. 1, 4 (1899)
121; Curist, Philip. J. Sc. 2 (1907) Bot. 117,
incl. var. sumatranum CHRIST, /.c. 118; v. A. v. R.
Handb. (1908) 48, 792, p.p.; Suppl. (1917) 77;
HeyneE, Nutt. Pl. (1927) 103; Burk. Dict. (1935)
536; BACKER & PostH. Varenfl. Java (1938) 23,
f, 7; LARD:-BLor & C. Cur. Fl. Gén. I.-C. 7; 2
(1939) 78, f. 10; HoLtrum, Rev. Fl. Mal. 2 (1954)
114, f. 45; A. F. Tryon, Miss. Bot. Gard. Bull. 43
(1955) n. 2; Am. Fern J. 47 (1957) 1; Cuine, FI.
Rep. Pop. Sin. 2 (1959) 197.—Polypodium ba-
rometz LINNE, Sp. Pl. (1753) 1092; Lour. FI.
Cochin. (1790) 675.—Aspidium barometz WILLD.
Sp. Pl. 5 (1810) 268.—Balantium glaucescens
Link, Fil. Sp. Hort. Berol. (1841) 40.—Dicksonia
baranetz LINK, /.c. 166; Hook. & BAK. Syn. Fil.
(1868) 49 (barometz).—C. glaucescens KUNZE,
Harnkr. 1 (1841) 63, t. 31; Hook. Sp: Fil. 1
(1844) 82.—C. assamicum Hook. Sp. Fil. 1 (1844)
83, t. 29B; Curist, Philip. J. Sc. 2 (1907) Bot. 117.
—Dicksonia assamicum GrirF. Notul. 2 (1849)
607; Ic. Pl. As. 2 (1849) t. 136, f. 2.—C. djambia-
me
num Hassk. Obs. Fil. Jav. 1 (1856) 61.—C.
glaucum [non (SM.) Hook. & ARN.] BEDD. Ferns
Br. Ind. (1865) t. 83.—Fig. 33a-c.
Caudex 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 314-414 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 2-4 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.
® 1962
Fig. 33. Cibotium barometz (L.) J. SM. a. Part of pinna-rachis with pinnules, x 24, 6. part of pinnule,
upper surface, showing venation (unbranched basal veins lead to sori) * 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. 12
Type specimen: none in Linnean Herbarium.
LINNAEUS stated that 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 Fraser’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 from
the story of the vegetable lamb of Tartary (see
A. F. Tryon, /.c.); 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 Pénawar (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, Farrnkr. 1 (1841)
64, 65; CurRisT, Philip. J. Sc. 2 (1907) Bot. 118;
CopeL. Fern Fl. Philip. 1 (1958) 84.—C. glaucum
[non (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. 7. 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.) HOLTTuUM,
comb. noy.—C. cumingii var. arachnoideum C.
Cur. Gard. 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: HoLtruM 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 min 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. Cur. Ind. Fil. Suppl. 3 (1934) 5, 57; CopEL. Gen.
Fil. (1947) 49.—Balantium Kau.r. En. Fil. (1824) 228, p.p.; J. Sm. Hist. Fil.
(1875) 257; Diets in E. & P. Pfl. Fam. 1, 4 (1899) 119; C. Cur. Ind. Fil. (1905)
xvi, 148; CopeL. Philip. J. Sc. 3 (1908) Bot. 301.—Dicksonia subg. Balantium
Hook. Sp. Fil. 1 (1844) 66, p.p.—Dicksonia § Eudicksonia Hook. & BAK. Syn.
Fil. (1868) 50, p.p.—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 ona
Dec. 1963] Le | _CYATHEACEAE _(Holttum) | . 167
larger axis open to admit grooves of smaller ones borne upon it t (fig. 340). 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 L’ HERIT.).
Distr. Subg. Culcita: Madeira and Azores, tropical America. Subg. 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 Dennstaedtia, as originally placed by CurisT.
Nomencl. The genus Balantium KAULF. was founded on two species, B. auricomum (a synonym of
Dicksonia arborescens, 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 KAULFUSsS. MAXON (l/.c.
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
PRrESL, 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, subg.
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’sS Index Filicum
(1905).
The genus Culcita differs 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 differs 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 Crenitis 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. n. 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. villosa
1. Inner indusium at maturity larger than outer, pouch- shaped and not reflexed; “midribs of quaternary
leafletss Sparsely( hairy. Lowland) plants+.. «i. -./(.i)eh%%e & dass Bie cbliee ee See Sa
168
FLORA MALESIANA
[ser: TI, vole
Fig. 34. Culcita javanica (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,
x 2, e. section of pinna-rachis, < 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. Varenfi.
Java (1939) 22.—Dicksonia javanica BL. En. PI.
Jav. (1828) 240; Hook. Sp. Fil. 1 (1844) 79; C.
Cure. Ind. Fil. (1905) 222.—Dennstaedtia javanica
Curist, Bull. Herb. Boiss. II, 4 (1904) 617;
v. A. v. R. Handb. (1908) 143.—Dicksonia cope-
landii Curist, Philip. J. Sc. 2 (1907) Bot. 183.—
Balantium copelandii CHRist in Copel. Philip. J.
Sc. 3 (1908) Bot. 301; ibid. 4 (1909) 62, t. 19;
Cope. in Elmer, Leafl. Philip. Bot. 2 (1908) 395;
yv. A. v. R. Handb. Suppl. (1917) 76.—Balantium
javanicum CopeEL. Philip. J. Sc. 4 (1909) Bot. 62;
v. A. v. R. Handb. Suppl. (1917) 75.—Balantium
Pilosum CopeEL. J. Str. Br. R. As. Soc. n. 63 (1912)
7: Sarawak Mus: J. 2 (1917)! 335sveeAe wees
Handb. Suppl. (1917) 76.—Dennstaedtia paraphy-
satay. A.V. R. Bull. Jard. Bot. Btzg II, 7. 16 (1914)
7; Handb. Suppl. (1917) 128.—Dennstaedtia
multifida v. A. v. R. Bull. Jard. Bot. Btzg II, 1. 20
(1915) 10; Handb. Suppl. (1917) 129.—C. cope-
landii Maxon, J. Wash. Ac. Sc. 12 (1922) 457;
C. Cur. Ind. Fil. Suppl. 3 (1934) 57; Gard. Bull.
S.S. 7 (1934) 224: Cope. Fern FI. Philip. 1 (1958)
86.—C. pilosa C. Cur. Ind. Fil. Suppl. 3 (1934)
57.—Fig. 34a-c.
Caudex 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].
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. 150 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 14 mm @,
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: BLumeg, 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 1500—2000 m in Benguet
Proy., 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
Balantium pilosum.
2. Culcita villosa C. Cur. Brittonia 2 (1937) 283;
CopEL. Philip. J. Sc. 78 (1949) 7.
Caudex ‘‘short, woody” (BRASs). Stipes to 80cm,
dark at base and covered with shining brown
CYATHEACEAE (Holttum)
169
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 FI. Philip. 1
(1958) 86.—Dicksonia straminea LABILL. Sert.
Austr. Cal. (1824) 7, t. 10. —Sitolobium stramineum
BrACcK. in Wilkes, U.S. Expl. Exped. 16 (1854)
273.—Dicksonia torreyana BRACK. l.c. 278, t. 58,
f. 2.—Dennstaedtia straminea J. SM. Hist. Fil.
(1875) 265.
Caudex 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 vy. A. v. R. 123
acuta Pr. 135
aeneifolia v. A. v. R. 133
var. subglauca v. A. v. R. 133
allocota v. A. v. R. 152
alpina v. A. v. R. 91, 92
alternans Hook. 145
amaiambitensis v. A. vy. R. 117, 118
amboinensis v. A. v. R. 114
(Alsophila) angiensis Gepp 135
annae v. A. v. R. 116
apiculata Rosenst. 93
arfakensis Gepp 113
atropurpurea C. Chr. 118
australis R. Br. 77
bakeri Zeill. 121
bartlettii (Copel.) C. Chr. 152
batjanensis Christ 88
benculensis v. A. v. R. 89
biformis Rosenst. 118
brevifoliolata vy. A. v. R. 111
170 FLORA MALESIANA
(Alsophila) brunnea Brause 120
brunoniana Hook. 137
burbidgei Bak. 151
burbidgei (non Bak.) Christ 117
buruensis Rosenst. 131
calocoma Christ 137
caudata J. Sm. ex Hook. 110
celebica (Bl.) Mett. 140
christii v. A. v. R., non Sod. 131
clementis 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 134
var. microloba Hassk. 135
var. robusta Hassk. 135
var. setulosa Hassk. 135
var. squamulata Hassk. 135
crenulata (Mett.) Hook. 111
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
dimorphotricha (Copel.) v. A. v. R. 131
dissitifolia Bak. 120
dryopteroidea Brause 158
dubia Bedd. 120
elmeri Copel. 131
extensa [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.) Hook 120
glabra (non BI.) 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. vy. R. 135
var. longepaleata y. A. v. R. 134
var. microloba y. A. v. R. 135
var. setulosa v. A. vy. R. 135
var. squamulata yv. A. y. R. 135
var. squamulosa v. A. vy. R. 135
var. trichocarpa Rosenst. 135
gregaria Brause 102
haenkei Pr.
var. angustata Hassk. 134
hallieri Rosenst. 157
hallieri vy. A. v. R. 117
helferiana Pr. 124
heteromorpha vy. A. y. R. 118
var. decomposita v. A. v. R. 118
heterophylla vy. A. v. R. 121
hewittii (Copel.) v. A. v. R. 118
(Alsophila) hieronymi Brause 88
hornei Bak. 120
hunsteiniana Brause 93
incisoserrata C. Chr. 113
indrapurae v. A. v. R. 93
janseniana v. A. v. R. 145
junghuhniana Kze 110
kemberangana (Copel.) C. Chr. 151
kenepaiana v. A. vy. R. 117
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. 113
latebrosa (non Wall.)
var. major Christ 131
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 (B\.) Hook. 121
macgillivrayi Bak. 123
margarethae Schroet. ex Christ 150
marginata Brause 155
matthewii Christ 157
melanocaulos vy. A. v. R. 120, 121
melanopus Hassk. 110
melanorachis Copel. 121, 122
mindanensis Christ 108
modesta Bak. 102
naumannii Kuhn 131
obliqua (Copel.) C. Chr. 146
obscura Scort. 152
okiana vy. A. v. R. 131
oligosora Mig. 152
olivacea Brause 120
ornata (non Scott) Bedd. 113
var. sikkimensis (non Clarke & Bak.) Bedd.
113
palembanica vy. A. vy. R. 89
papuana Ridl. 155
paraphysata v. A. vy. R. 152
parvifolia Holtt. 145
persquamulata y. A. vy. R. 124
persquamulifera v. A. v. R. 134
Dhilippinense Hort. Veitch. 157
poiensis (Copel.) v. A. v. R. 151
polycampta Kze 124
polyphlebia 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
recuryvata Brause 101
reducta v. A. v. R. 120
ridleyi Bak. 152, 153
[ser. IT, volag2
Dec. 1963]
Index to Cyatheaceae
(Alsophila) robusta De Vr. 110
rosenstockii Brause 101
rubiginosa Brause 93
rumphiana v. A. v. R. 131
salyinii Hook. 115
sangirensis Christ 130
saparuensis v. A. v. R. 88
sarawakensis C. Chr. 152
scaberula Christ 130
scaberulipes v. A. v. R. 131
scabriseta (Copel.) v. A. v. R. 131
scandens Brause 119
schlechteri Brause 123
smithiana Pr. 135
speciosa [non (Meyen) Pr.] Goldm. 157
spinifera v. A. v. R. 114
squamulata (Bl.) Hook. 152
squamulata (non Bl.) Hook. 118
straminea Gepp 88
subcomosa C. Chr. 131
subdimorpha (Copel.) v. A. v. R. 121
subdubia v. A. v. R. 124
subobscura v. A. vy. R. 152
subulata v. A. v. R. 118
tenggerensis Rosenst. 134
tenuis Brause 90
tomentosa (Bl.) Hook. 132
var. novoguineensis Rosenst. 132
trichodesma Scort. 150
trichophora (Copel.) v. A. v. R. 151
tristis (Bl.) Bl. ex Moore 158
truncata Brack. 157
var. nivea Christ 140
var. sagittata Christ 140
umbrosa Wall. ex Ridl. 124
vexans Ces. 120
vitiensis Carr 131
wallacei Mett. 151
warburgii Christ 158
warihon (Copel.) C. Chr. 108
wengiensis Brause 88
woodlarkensis (Copel.) C. Chr. 158
xanthina (Domin) C. Chr. 152
xantholepia v. A. vy. R. 152
xantholepis Christ ex Diels 158
Amphicosmia Gardn. 73
alternans Moore 145
Javanica (Pr.) Moore 111
manilensis (Pr.) Moore 110
Aspidistes Harris 65, 67
Aspidium barometz Willd. 165
triste Bl. 158
Athyrium 69
Balantium Kaulf. 71, 158, 166, 167
auricomum 167
blumei Kze 159
chrysotrichum Hassk. 159
copelandii Christ 168
culcita 167
glaucescens Link 165
javanicum Copel. 168
pilosum Copel. 168, 169
Cathetogyratae 71
Chnoophora Kaulf. 73
glauca Bl. 135
(Chnoophora) lanuginosa Jungh. 132
lurida Bl. 121
tomentosa Bl. 132
17
Cibotium Kaulf. 65, 67, 69, 70, 71, 72, 159, 164-
166, 169
arachnoideum (C. Chr.) Holtt. 164, 166
assamicum Hook. 165
barometz (L.) J. Sm. 164, 165*
var. cumingii v. A. v. R. 166
var. sumatranum Christ 165
chamissoi Kaulf. 164
crassinerva Rosenst. 166
cumingli 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
villosa C. Chr. 167, 169
Cyathea Smith 65, 67, 69, 70, 71, 72-158, 159
acanthophora Holtt. 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. vy. R.) Domin 152
alpicola Domin 91
alpina v. A. v. R. 91
alsophiliformis (v. A. v. R.) Domin 111
alternans (Wall. ex Hook.) Pr. 76, 141, 145, 150
var. lobbiana (Hook.) Domin 145
var. sarawakensis (Hook.) Domin 145
var. serrata Ridl. 145
amaiambitensis (vy. 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
2 ___ FLORA MALESIANA [ser. IT, vole
(Cyathea) aneitensis Hook. 157 (Cyathea) burbidgei [non (Bak.) Copel.] Holtt. 150
angiensis (Gepp) Domin 126, 135, 136 buruensis (Rosenst.) Domin 131
angustipinna Holtt. 141, 143 callosa Christ 80, 106
annae (vy. A. v. R.) Domin 116, 117* calocoma (Christ) Copel. 137
apiculata (Rosenst.) Domin 79, 93, 94 camaguinensis Copel. 196
apoensis Copel. 78, 86 campbellii Copel. 104
arachnoidea Hook. 140 capensis (L. f.) Sm. 70*
arachnoidea (non Hook.) Grether & Wagner 140 capitata Copel. 65, 141, 142, 144*
arachnoidea (non Hook.) Back. & Posth. 140 carrii Holtt. 138, 139*
arborea (L.) Sm. 74 catillifera Holtt. 78, 90
arborea [non (L.) Sm.] caudata (J. Sm. ex Hook.) Copel. 81, 108, 110,
var. pallida Hassk. 86 114
arborescens Copel. 110 caudiculata (Rosenst.) Domin 108
archboldii C. Chr. 79, 98 caudipinnula (v. A. v. R.) Domin 89
var. horrida Holtt. 99 celebica Bl. 138, 140, 157
arfakensis Gepp 77, 82 celebica vy. A. v. R. (non BI.) 139
arguta Copel. 146 cheilanthoides Copel. 95
argyrolepis Copel. 131 christii Copel. 79, 95
arthropoda Copel. 141, 143 cincinnata Brause 77, 86
arthropterygia v. A. v. R. 89 cinerea Copel. 80, 104
aruensis Domin 131 clementis (Copel.) Copel. 135
ascendens Domin 80, 101 coactilis Holtt. 78, 87
aspidioides (Bl.) Zoll. & Mor. 158 confluens (v. A. v. R.) Domin 102
assimilis Hook. 142, 150 contaminans (Wall. ex Hook.) Copel. 67, 68*,
assimilis (non Hook.) Christ 101 102. 71a 252126 135318670138
atrispora Domin 158 var. persquamulifera v. A. v. R. 134
atropurpurea Copel. 116, 118 costalisora Copel. 78, 87
atrospinosa Holtt. 126, 137 costulisora Domin 81, 109
atrox C. Chr. 66*, 107*, 126, 128 crassipes Sod. 124
var. atrox 128 crenulata Bl. 76, 78, 89, 92*, 93, 95, 104
var. inermis Holtt. 126, 128, 129*, 130* f. Jatissima v. A. v. R. 103
auriculifera Copel. 138, 139 f. squamulosa vy. A. v. R. 103
australis (R. Br.) Domin 69 f. subspinulosa v. A. v. R. 93
austrosinica Christ 106 crinita (Hook.) Copel. 132
barisanica (v. A. v. R.) Domin 89 cucullifera Holtt. 80, 103
bartlettii Copel. 152 curranii Copel. 126, 133
batjanensis (Christ) Copel. 75*, 78, 88 curvipinnula C. Chr. 133
beccariana Ces. 150 cyclodonta (Christ) v. A. v. R. 127
benculensis (v. A. v. R.) v. A. v. R. 89 deminuens Holtt. 141, 145
bicolana Copel. 149 densisora v. A. v. R. 140
bicolor Copel. 108 deuterobrooksii Copel. 152
bidentata Copel. 98 dicksonioides Holtt. 80, 83*, 106, 107*
biformis (Rosenst.) Copel. 68, 116, 118, 119*, dielsii (Brause) Domin 157
120 dimorpha (Christ) Copel. 116, 121
biliranensis Copel. 108 dimorphophylla Domin 120
binuangensis v. A. v. R. 141, 145 dimorphotricha Copel. 131
bipinnatifida Copel. 151 discophora Holtt. 76, 142, 148
bontocensis Copel. 149 distans Rosenst. 92
borbonica Desy. 141 dissitifolia Domin 120
borneensis Copel. 81, 110 doctersii v. A. v. R. 80, 102
brackenridgei Mett. 157 dubia Bedd. 120
brassii Copel. 131 dulitensis Bak. 150
brauseana Domin 88 dupaxensis Copel. 110
brevifoliolata (v. A. vy. R.) v. A. v. R. 111 dura Copel. 108
breviloba Copel. 146 edanoi Copel. 80, 108
brevipes Copel. 110 elliptica Copel. 141, 142, 146, 151
brooksii Copel. (non Maxon) 152 elmeri (Copel.) Copel. 126, 131, 132
brownii Domin 70* eminens Domin 131
brunei Christ 124 eriophora Holtt. 80, 102
brunnea (Brause) Domin 120 everta Copel. 77, 85
brunoniana (Hook.) Clarke & Bak. 137 excavata Holtt. 79, 94*
brunonis (J. Sm.) Wall. ex Hook. 143 excelsa sensu Kze (non Sw.) 92
bulusanensis Copel. 146 faberiana Domin 103
biinnemeijerii v. A. v. R. 78, 79, 94 fallax (vy. A. v. R.) Domin 111
burbidgei (Bak.) Copel. 151 fenicis (C. Chr.) Copel. 81, 110
Dec. 1963] Index to Cyatheaceae 7 173
(Cyathea) ferruginea Christ 79, 100 (Cyathea) kemberangana Copel. 151
ferrugineoides Copel. 100
foersteri Rosenst. 79, 99, 100
foxworthyi Copel. 106
frondosa Rosenst. 157
fructuosa Copel. 104
fugax v. A. v. R. 126, 137
fuliginosa (Christ) Copel. 81, 108
fusca Bak. 153, 154, 156
fuscopaleata Copel. 143
gazellae (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, 111
glabra (BI.) Copel. 67, 115, 116, 120
glabrescens (vy. A. vy. 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
hemichlamydea Copel. 110
heterochlamydea Copel. 75*, 80, 108, 110, 157
heteroloba Copel. 149
heteromorpha (vy. A. vy. R.) Domin 118
heterophylla (v. A. vy. R.) Domin 121
hewittii Copel. 118
holttumii Copel. 146
hooglandii Holtt. 77, 83*, 84
hookeri Thw. 141
hornei (Bak.) Copel. 116, 120
horridipes (v. A. v. R.) Domin 114
horridula Copel. 78, 90
hunsteiniana Brause 77, 82
var. acuminata Brause 82
hymenodes Mett. 76, 78, 89, 94
hypocrateriformis 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 (vy. 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
var. petiolata Hook. 146
integra (non J. Sm. ex Hook.) v. A. v. R. 147
janseniana (y. 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
kenepaiana (v. A. v. R.) Domin 117
keysseri Rosenst. 95
kinabaluensis Copel. 143
kingii (Clarke) Copel. 121
kingii Rosenst. 156
klossii Ridl. 78, 91
korthalsii Mett. 89
korthalsii (non Mett.) C. Chr. 104
lanaensis Christ 108
lastreoides (vy. 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. 115
leucophaes Hassk. 92
leucostegia Copel. 140
leucotricha Christ 125, 127, 128
leytensis Copel. 140
lobata Copel. 86
lobbiana Hook. 145
loerzingii 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
melanopus (Hassk.) Copel. 111
melanorachis (Copel.) Copel. 121
merapiensis (vy. 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. Il, vole
(Cyathea) mitrata Copel. 104 (Cyathea) raciborskii Copel. 81, 111
modesta (Bak.) Copel. 80, 102 ramispina (Hook.) Copel. 116, 117, 118, 121
mollis Copel. 151 ramosiana v. A. yv. R. 149
moluccana R. Br. 67, 141, 143, 144*, 145 ramosii Copel. 151
montana Sm. 109 rebeccae (F. v. M.) Domin 116, 120
moseleyi Bak. 153, 157 recommutata Copel. 117*, 118, 121, 122*
muelleri Bak. 80, 98, 103, 104* recurvata (Brause) Domin 80, 101
naumannii (Kuhn) Domin 131 reducta (vy. A. v. R.) Domin 120
negrosiana Christ 78, 90 ridleyi (Bak.) Copel. 145, 152
nigrolineata Holtt. 79, 100 rigens Rosenst. 77, 85, 86
nigropaleata Holtt. 81, 113 rigida Copel. 96
nigrospinulosa v. A. v. R. 156 robinsonii Copel. 142, 149
novae-caledoniae (Mett.) Copel. 138 rosenstockii Brause 153, 154*, 156
novoguineensis Brause 95 rubella Holtt. 116, 122
obliqua Copel. 141, 146 rubiginosa (Brause) Domin 79, 93
obscura (Scort.) Copel. 142, 152 rudimentaris (v. A. v. R.) Domin 115
obtusata Rosenst. 110 rufopannosa Christ 80, 106
oinops Hassk. 74*, 75*, 76, 80, 92, 103, 104 rumphiana (vy. A. v. R.) Merr. 131
oinops (non Hassk.) Racib. 92 rumphii Desy. 157
okiana (v. A. v. R.) v. A. v. R. 131 runensis v. A. v. R. 153, 156
oligocarpia Jungh. 103 saccata Christ 78, 93
olivacea (Brause) Domin 121 salticola (v. A. v. R.) Domin 114
oosora Holtt. 79, 101 sangirensis (Christ) Copel. 74, 126, 130, 132
ordinata Copel. 132 saparuensis (v. A. v. R.) v. A..v. R. 88
orientalis (Kze) Moore 73*, 75*, 78, 86, 95 sarasinorum Holtt. 126, 134
pachyrrhachis Copel. 80, 105 sarawakensis Hook. 145
paleacea Copel. 96 scaberula (Christ) Domin 131
paleata Copel. 93 scaberulipes (v. A. v. R.) Domin 131
palembanica (vy. A. v. R.) v. A. v. R. 89 scabriseta Copel. 131
pallidipaleata Holtt. 78, 87 scandens (Brause) Domin 116, 119
papuana (Ridl.) v. A. v. R. 153, 155 schizochlamys Bak. 91
paraphysata Copel. 152 schlechteri (Brause) Domin 116, 123
paraphysophora (vy. A. v. R.) Domin 114 semiamplectens Holtt. 81, 109
parva Copel. 78, 87 senex v. A. v. R. 142, 149
parvipinna Holtt. 153, 155 sepikensis Brause 95
patellifera v. A. v. R. 78, 89 sessilipinnula Copel. 148
peranemiformis C. Chr. 82 setifera Holtt. 153, 154
percrassa C. Chr. 77, 84 setulosa Copel. 80, 103
perpelvigera v. A. v. R. 77, 82, 84 sibuyanensis Copel. 142, 149
perpunctulata (v. A. v. R.) Domin 81, 114 singalanensis (vy. A. v. R.) Domin 102
persquamulata (vy. A. v. R.) Domin 124 sinops (sphalma), cf. oinops 92, 104
persquamulifera (v. A. v. R.) Domin 126, 134 sinuata Hook. 141
philippinensis Bak. 142, 149, 150 spinifera (v. A. y. R.) Domin 114
var. nuda Copel. 149 spinulosa Wall. ex Hook. 98, 106
physolepidota Alston 81, 113 var. muriculata Hassk. 92
pilulifera Copel. 126, 133, 134 squamicosta Copel. 108
pinnata Roxb. 143 squamulata (BI.) Copel. 67, 74*, 142, 145, 4g
podophylla (Hook.) Copel. 120 148, 152
poiensis Copel. 151 stipitipinnula Holtt. 142, 147
polycarpa Jungh. 92 stipitulata Copel. 150
polypoda Bak. 142, 150, 151 straminea (Gepp) v. A. v. R., non Karst. 88
princeps (Linden) Meyer 124 strigosa Christ 125, 128
procera Brause 125, 127* subbipinnata Copel. 146
pruinosa Rosenst. 77, 84 subcomosa (C. Chr.) Domin 131
pseudoalbizzia Copel. 149 subconfluens (v. A. v. R.) Domin 102
pseudobrunonis Copel. 143 subdimorpha Copel. 121
pseudomuelleri Holtt. 79, 98, 99* subdubia (v. A. v. R.) Domin 116, 120, 124
pteridioides Copel. 137 subg. Cyathea 69, 73, 75*, 76, 77-124
pulcherrima Copel. 125, 126, 127, 153 sect. Cyathea 76, 77-115, 123
pulchra Copel. 152 sect. Gymnosphaera (BI.) Holtt. 76, 80,
pumilio vy. A. v. R. 88, 89 115-124, 141
punctulata (v. A. v. R.) v. A. v. R. 81, 112 subg. Gymnosphaera Tindale 115
pustulosa (Christ) Copel. 137 subg. Sphaeropteris (Bernh.) Holtt. 69, 74, 76,
pycnoneura Holtt. 77, 85 124-158
quadripinnatifida Copel. 140 sect. Schizocaena (J. Sm.) Holtt. 76, 124, 141
Dec. 1963]
Index to Cyatheaceae_
(Cyathea)
subsect. Sarcopholis Holtt. 76, 141, 153-158
subsect. Schizocaena 76, 112, 141-153
sect. Sphaeropteris 69, 76, 124-141, 151
subsect. Fourniera (Bommer) Holtt. 76,
124, 138-141
subsect. Sphaeropteris 76, 124, 125-138, 153
subobscura (v. A. v. R.) Domin 152
subspathulata Brause 95
subtripinnata Holtt. 77, 86
subulata (vy. A. v. R) Domin 118
subuliformis vy. A. v. R. 91
sulitii Copel. 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.) Zoll. & 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
tuberculata vy. 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
zollingeriana 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
175
(Dennstaedtia) ampla (Bak.) Bedd. 161
cuneata (Hook.) Moore 161
erythrorachis (Christ) Diels 162
flaccida (Forst.) Bernh. 162
glabrata (Ces.) C. Chr. 162
gomphophylla (Bak.) C. Chr. 162
javanica Christ 168
moluccana (BI.) Moore 162
multifida v. A. v. R. 168
paraphysata v. A. v. R. 168
remota (Christ) Diels 162
rhombifolia (Bak.) C. Chr. 162
scabra (Wall.) Moore 161, 162
scandens (Bl.) Moore 162
smithii (Hook.) Moore 162
straminea J. Sm. 169
Diacalpe aspidioides Bl. 158
Dichorexia Pr. 73, 76
latebrosa Pr. 115
Dicksonia L’Hérit. 65, 67, 68, 69, 70, 71, 72, 74,
108, 158-162, 167
ampla Bak. 161
antarctica Labill. 69, 160*
arborescens L’Hérit. 158, 159, 167
archboldii Copel. 159, 160
assamicum Griff. 165
baranetz Link 165
blumei (Kze) Moore 69, 159, 160*
blumei [non (Kze) Moore] C. Chr. 159
chrysotricha (Hassk.) Moore 159
copelandii Christ 168
culcita L’ Hérit. 167
cuneata Hook. 161
deltoidea Hook. 161
erythrorachis Christ 162
flaccida (Forst.) Sw. 162
glabrata 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 BI. 163
neglecta Fée 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 Thunb. 162
subg. Balantium Hook. 158, 166
torreyana Brack. 169
176 : FLORA MALESIANA
(Dicksonia) zippeliana Kze 162
Dicksoniaceae 65, 71
Diplazium 157
Disphenia Pr. 73, 76
orientalis Kze 86
Dryopteris 65, 67, 69, 158, 167
atrispora C. Chr. 158
Eatoniopsis Bommer 73
Fourniera Bommer 73, 138
Gleichenia 70, 71, 169
Gleicheniaceae 71
series Marginales 71
series Superficiales 71
Gymnosphaera Bl. 73, 76, 115
atropurpurea (Copel.) Copel. 118
biformis (Rosenst.) Copel. 118
bipinnatifida Copel. 151
burbidgei (Bak.) Copel. 151
dinagatensis Copel. 151
glabra Bl. 120
glabra (non BI.) Copel. 151
gracillima Copel. 123
hewittii Copel. 118
holttumii Copel. 146
hornei Copel. 121
kingii (Copel.) Copel. 121
melanoclada Copel. 121
melanorachis Copel. 121
mollis (Copel.) Copel. 151
obliqua (Copel.) Copel. 146
papuana (Ridl.) Copel. 155
pulchra Copel. 152
ramispina (Hook.) Copel. 117
recommutata Copel. 118
sarawakensis (C. Chr.) Copel. 152
schlechteri (Brause) Copel. 123
squamulata Bl. 152
squamulata (non Bl.) J. Sm. ex Hook. 118
subbipinnata Copel. 146
trichophora Copel. 151
vexans (Ces.) Copel. 120
Helicogyratae 71
Hemitelia R. Br. 69, 71, 73, 77
alsophiliformis v. A. v. R. 111
alternans Hook. 145
arfakensis (Gepp) v. A. v. R. 82
barisanica v. A. v. R. 89
bicolor (Copel.) v. A. vy. R. 108
capensis Hook. 111
caudata (J. Sm.) Mett. 110
caudiculata Rosenst. 106, 108
caudipinnula y. A. v. R. 89
confluens vy. A. vy. R. 102
crenulata Mett. 111
fallax v. A. vy. R. 111
var. major v. A. v. R. 111
glaucophylla y. A. v. R. 111
hemichlamydea (Copel.) v. A. v. R. 110
heterochlamydea vy. A. v. R. 108
horrida 73
horridipes v. A. v. R. 114
javanica Pr. 110
jJunghuhniana (Kze) Mett. 111
var. dissoluta Racib. 111
var. paraphysata v. A. v. R. 115
latipinnula v. A. v. R. 105
ledermannii Brause 140
leptolepia v. A. v. R. 115
manilensis Pr. 110
merapiensis v. A. v. R. 111
montana y. A. v. R. 109
multiflora (Sm.) R. Br. 77
paraphysophora v. A. vy. R. 114
perpunctulata v. A. v. R. 114
rudimentaris v. A. v. R. 115
salticola y. A. vy. R. 114
singalanensis v. A. v. R. 102
subconfluens v. A. v. R. 102
sumatrana v. A. vy. R. 114
tonglonensis v. A. vy. R. 108
truncata (non Brack.) Christ 139
warihon (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 kingit (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
lunulatum Forst. 131
Pseudogyratae 71
Pteridaceae 71
Saccoloma sorbifolia Christ 163
Schizocaena J. Sm. 73, 76, 141
alternans J. Sm. 145
arthropoda (Copel.) Copel. 143
brunonis J. Sm. ex Hook. 141, 143
capitata Copel. 142
gaudichaudii Fée 143
kinabaluensis (Copel.) Copel. 143
moluccana (R. Br.) Copel. 143
Sitolobium stramineum Brack. 169
Sphaeropteris Bernh. 124
Stenochlaena dubia v. A. v. R. 118
Stenolepia tristis (Bl.) v. A. v. R. 158
Tapeinidium pinnatum (Cay.) C. Chr. 162
Thelypteris 65, 66, 158
immersa (BI.) Ching 158
Thyrsopteris 65, 69, 70, 72, 167
Thysanobotrya v. A. v. R. 73, 115
arfakensis (Gepp) v. A. v. R. 118
Trichopteris falcata Llanos 146
[ser. Il, vol. 17, Dect 19Ga]
(Hemitelia) latebrosa (Wall.) Mett. 111, 115
LINDSAEA-GROUP (KK. 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. Jndument of the rhizome of scales, these non-
peltate, non-clathrate (in Mal. 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 -+ 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
+- 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., 2 African spp.), Ormoloma (2 spp., neotropical), Tapeinidium (17 spp.,
SE. Asia to Samoa), Sphenomeris (11 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 /soloma, 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. I; vole
most species of Lindsaea; in the latter genus the taxonomic value of the character is slight. In Sphenomeris
it is also variable but apparently well-marked and constant for each species (KRAMER, 1957, /.c. 106).
Cytotaxonomy. 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. orbiculata 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. chienii (KurRITA). 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. Odontoloma there is a count of n = c. 44 for L. repens var.
sessilis (WALKER, in litt.) and 44 or 45 for L. pulchella var. blanda (WALKER, in litt.), in sect. Lindsaea
n = 88 in L. portoricensis (neotropical) (WALKER) and c. 84in L. arcuata (neotropical) (MICKEL, WAGNER
& Lim CHEN). Other numbers may or may not be related, e.g. L. ‘nitida’ (= integra?) (MANTON in
KRAMER) and L. ‘scandens var. terrestris’ (= L. doryphora) (MANTON in KRAMER), both c. 47. The
number n = 47 has, however, been found unequivocally in certain species, e.g. L. ‘macraeana’ (= L.
repens var.) (WAGNER), L. ‘concinna’ (Australia; L. brachypoda?) (MANTON in Kramer) and L. parallelo-
gramma (MANTON in KRAMER). The numbers c. 50 and c. 100 reported for L. decomposita (= L. obtusa?)
(MANTON in Hotttum), 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 Hotttum, Rev. FI. Mal. 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 (/.c.), although he noted the great differences
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 (HoLtTTuM, FI. Mal. 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, /.c.), 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-
loma, 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, /.c.; 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
Ultimate free or nearly free divisions of a linear- « or cuneate- divaricate type, with the sorus (sori)
on their ance margin; a al 2- or 3-celled, not usually found; spores monolete or tri-
fete! ee. : 1. Sphenomeris
1. Sorion many vein- 1-ends, « or, if on 8 or fewer, the sides of the indusium free, or r the pinnules dimidiate,
or the veins anastomosing, or these characters combined.
3. Ultimate free divisions lanceolate, equal-sided or the base apo e. auricled; veins free; spores
N
monolete. §2 ~ 25 Xyropteris
3. Ultimate free divisions not lanceolate and equal- sided, or, if so, the Spores trilete: veins free or
anastomosing. “SA et eli) Oe “Re Fae eee See EGE, 4. Lindsaea
1SPHENOMERIS
MAxon, J. Wash. Ac. Sc. 3 (1913) 144, nom. cons.; COPELAND, Gen. Fil. (1947) 54;
Philip. J. Sc. 78 (1949) 24; Ho_trrum, Rev. Fl. Mal. 2 (1954) 340; KRAmer, Act.
Bot. Neerl. 6 (1957) 152; COPELAND, Fern FI. Philip. 1 (1958) 115.—Davallia
J. E. SmirH, Mém. Ac. Turin 5 (1793) 414; Hooker, Sp. Fil. 1 (1845) 151; and
of many other authors; all in part.—Stenoloma Fer, Gen. Fil. (1852) 330, p.p.
min.; CHING, Fl. Reip. Pop. Sin. 2 (1959) 275, in part; and of other authors.—
Odontosoria F&E, Gen. Fil. (1852) 325; J. Smitu, Hist. Fil. (1875) 263; Dies in
E. & 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. a/utacea). 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 11 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 FE£E£ 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 of Sphenomeris,
and certainly not in Sphenomeris clavata. The issue should be reconsidered.
KEY TO THE SPECIES
1. Rhizome 5-20 mm 4g, 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 sae segments on (2—)5-8 vein-ends,
occupying their whole apical margin (fig.5). . . : 1. S. retusa
1. Rhizome 2-4 mm g, with a lindsaeoid protostele; scales. to 4 mm long, to ‘4 seriate at the base (wider
in S. biflora); spores monolete; sori of larger segments on 1—2(—4) vein-ends, not occupying their
whole apical margin.
180
FLORA MALESIANA
[ser. II, vol. 13
2. Scales up to 3-seriate at the base; sori strictly uninerval; ultimate divisions 4-34 mm 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-
mate; crenate = 1)
2. S. biflora
2. Scales 1—3-seriate (or to 4-seriate at the suddenly broadened base); sori 1—-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 + bipinnatifid, tripinnate + pinnat-
ifid, or more dissected .
1. Sphenomeris retusa (CAv.) MAxon, J. Wash.
Ac. Sc. 3 (1913) 144; COPELAND, Philip. J. Sc. 78
(1949) 24: Fern Fl. Philip. 1 (1958) 116.—Da-
vallia retusa Cav. Descr. (1802) 278.— Stenoloma
retusum (CAV.) FEE, Gen. Fil. (1852) 330.—Lind-
saea retusa (CAv.) Metr. Fil. Lips. (1856)
105.— Odontosoria retusa (CAv.) J. SMITH, Bot.
Voy. Herald (1857) 430.—Schizoloma retusum
(CAv.) KUHN, Chaetopt. (1882) 346. — Type:
NEE s.n., Mt Isarrog, Luzon (MA, n.v.; phot. U).
Lindsaea cuneifolia Pres_, Rel. Haenk. 1 (1825)
60.—? Saccoloma cuneifolium PRESL, Tent. Pterid.
(1836) 126. — Acrophorus cuneifolius (PRESL)
Moors, Ind. Fil. (1857) 41. — Type: HAENKE
s.n., Luzon (n.v.).
? Adiantum falcatum BLANCO, FI. Filip. (1837)
833. — Type: BLANCO s.n., ‘Mandalogon’ (n.yv.;
perhaps this species acc. to C. Cure. Ind. Fil.
Suppl. 1, 1913, corr. 89).
Davyallia 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 (FI, 2 sh.).
Odontosoria lindsayae v.A.v.R. Bull. Dép. Agr.
Ind. Néerl. 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 14-2 cm g, 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. 24-1 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
3. S. chinensis
triangular, shortly petiolulate, acuminate, c. 5-10
cm long, 24%-6 cm wide, pinnate + 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, 1'%-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, 244-3 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. + 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 w.
Distr. Malesia: Celebes, Philippines (almost
throughout), Moluccas (Morotai, Halmahera,
Tidore, Ambon, Ceram), New Guinea, Admiralty
Is., Bismarck Arch.; Solomon Is.,?7New 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. chinensis?), but this is scarcely the
case with all.
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. Cur. Pinna, x 14% (CLEMENS
50992). — F‘g. 5. S. retusa (CAv.) MAxon. Pinnule, x 114 (KoorDERS 17033).
182
FLORA MALESIANA
[ser. JF vole
2. Sphenomeris biflora (KAULFUSS) TAGAWA, J.
Jap. Bot. 33 (1958) 203; KRAMER, Blumea 15
(1968) 573.—Davallia biflora KAULFUss, Enum.
(1824) 221.— Microlepia biflora (KAULFUSS) METT.
Fil. Lips. (1856) 104. — Odontosoria biflora
(KAULFuss) C. Cur. Ind. Fil. (1906) 464. —
Stenoloma biflorum (KAULFUSS) CHING, Sinensia 3
(1933) 338; TaGawa, J. Jap. Bot. 22 (1948)
160.—Type: CHAmMisso s.n., Manila, Luzon (B).
Davallia tenuifolia SWARTZ var. lata HOOKER
ex Moore, Ind. Fil. 2 (1861) 301, based on B
(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 (n.y.).
Stenoloma littorale TAGAWA, Act. Phytotax.
Geobot. 6 (1937) 225. — Stenoloma chusanum
(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. Ill. Jap.
Pterid. 1959, 256, n.v.) — Type: TAsiro s.n.,
Oshima J., Shikoku (KYO, z7.¥.).
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 9g; scales gold-
en to medium brown, occasionally castaneous,
to 314 mm long, in the larger ones the apical half
or less umiseriate, 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
14% 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,
4-114 mm long (at right angles to vein). Spores
monolete, ellipsoid, medium brown, smooth,
c. 32-35 by 40-45 wu.
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.
Cur. 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. Pl. 2 (1753) 1099.—
Adiantum chinense (L.) BURMAN, FI. Ind. (1768)
236 (prob.).—Davallia chinensis (L.) J. E. Smit,
Mém. Ac. Turin 5 (1793) 414. — Microlepia
chinensis (L.) Metr. 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 Kunn, Fil. Afr. (1868) 67, non CHING (1929).—
Stenoloma chinense (L.) Bepp. Handb. Ferns
Br. Ind. (1883) 70. — Type: OsBECK s.n., China
(S-PA).
Adiantum chusanum L. Sp. Pl. 2 (1753) 1095.—
Davallia chusana (L.) WILLD. Sp. Pl. 5 (1810)
475. — Sphenomeris chusana (L.) COPELAND,
Bull. Bish. Mus. 59 (1929) 69; Philip. J. Sc. 78
(1949) 24; Hottrum, Rev. FI. Mal. 2 (1954)
341; COPELAND, Fern FI. Philip. 1 (1958) 115. —
Odontosoria chusana (L.) MASAM. Mem. Fac.
Sci. Agr. Taihoku Imp. Univ. 11 (1934) 67.—
Type: coll.?, China (n.v.).
Adiantum tenuifolium LAMK, Encycl. 1 (1783)
44. — Dayallia tenuifolia (LAMK) SWARTZ in
Schrader, J. Bot. 18002 (1801) 88. — Stenoloma
tenuifolium (LAMK) FEE, Gen. Fil. (1852) 330.—
Microlepia tenuifolia (LAMK) Mertr. Fil. Lips.
LINDSAEA-GROUP (Kramer)
183
April 1971]
(1856) 104, pl. 27, f. 1-4, non Presi (1851).—
Odontosoria tenuifolia (LAMK) J. SMITH, Cat.
Cult. Ferns (1857) 67. — Odontosoria chinensis
(L.) J. SmituH var. tenuifolia (LA)MK MATSUM.
Ind. Pl. Jap. (1904) 330.—Sphenomeris chinensis
(L.) Maxon var. tenuifolia (LAMK) C. CHR.
Dansk Bot. Ark. 7 (1932) 78. — Odontosoria
chusana (L.) MAsAM. var. tenuifolia (LAMK)
(err. “MAK.’) MASAM. Mem. Fac. Sci. Agr.
Taihoku Imp. Univ. 11 (1934) 67.— Sphenomeris
chusana (L.) COPELAND var. tenuifolia (LAMK)
HOLTTUM (incorr. ascr. to C. Cur.), Rev. Fl. Mal.
2 (1954) 341, as to type only. — Type: SONNERAT
S73 inde: (P):
Davallia didyma HeEpwic, Fil. Gen. & Sp.
(1803) pl. 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).
Hymenophyllum ramosissimum HAM. ex D.
Don, Prod. Fl. Nepal. (1825) 12. — Type:
HAMILTON s.n., Nilkantha, Nepal (n.v.; identity
teste C. Cue. Ind. Fil.).
Trichomanes malayanum RoxsB. 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 g; 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
144-3 mm g in the middle, c. 12-60 cm long,
2,-1 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 %—)3-10 cm wide, 2-41% 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 shallowly 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, 44-2 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-(pinnula-)apex. 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, + 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, Icss often
on two (mostly connivent) vein-ends; indusium
often 4% by 4% mm, with convex base. Spores as
in var. chinensis, c. 44-48 yw 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) 572.—Odontosoria chinensis (L.) J.
SMITH var. divaricata CHRIST, Journ. de Bot.
sér. 2, II (1909) 23.—Sphenomeris chusana (L.)
COPELAND var. divaricata (CHRIST) TARDIEU-
Biot, Fl. Madag. Com. Se fam. I (1958) 29.—
Type: CHEVALIER 14309, Sao Tomé (P).
Sphenomeris chusana (L.) COPELAND var.
tenuifolia of HoL_trum, Rey. Fl. Mal. 2 (1954)
341, and of other authors; not Adiantum tenui-
folium LamMK.— 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-
184
FLORA MALESIANA
[ser. Ty volar
iculate; at the base often 144 mm wide, slightly
broadened to the apex, 1-144 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 yw long.
Distr. Sao Tomé; 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).
Ecol. Asthenext variety, but mostly above 700m.
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 34-114 mm long. Spores mostly
42-48 uw 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, efc., 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) 39.—Stenoloma veitchii (BAKER)
C. Cur. 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 g;
scales dark castaneous, acicular, the upper half
uniseriate, the base bi- to triseriate, to 24% mm
long. Leaves %-1 cm apart; petioles slender,
14%-2 mm g, fuscous or upward stramineous,
abaxially terete, adaxially broadly and shallowly
grooved, 30-40 cm long. Lamina narrowly
oblong, c. 30 by 5 to 70 by 15 cm, bipinnate +
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. 44-34 mm wide,
4 mm long (at right angles to its vein), + 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 yw.
Distr. Malesia: confined to Mt Kinabalu,
Sabah, Borneo (3 coll.).
Ecol. In mountain forests, 2000-2500 m.
2. TAPEINIDIUM
(PRESL) C. Cure. Ind. Fil. (1906) 631; CopELAND, Gen. Fil. (1947) 53; HoLTTUM,
Rev. Fl. Mal. 2 (1954) 338; KRAMER, Blumea 15 (1968) 545.—Microlepia sect.
Tapeinidium PresL, Epimel. Bot. (1851) 968.—Protolindsaya COPELAND, Philip.
J. Sc. 5 (1910) Bot. 283.—Wéibelia auct. non BERNHARDI; Fee, Gen. Fil. (1852)
April 1971] | LINDSAEA-GROUP (Kramer) 185
331; Diets, in E. & P. Nat. Pfl. Fam. I, 4 (1902) 216. ie nadaniA 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
Smmlong: . . . . 15. T. longipinnulum
2. Sori intramarginal, almost ‘always uninerval: rachis abaxially. sharply bi-angular or sulcate, basally
dark and pale-margined; scalesto24%mmlong . . . . . 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 sorinot 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. Laminato 12cmlong;petioleslender,lessthaniImmge. . . . . . 410. T. oligophlebium
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. pinnatum
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.
2 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
bases am 2)8),. <- Pig 3.) eee? (2 0or.2. C.-C es Pe ee eee
* For intermediates see 12a. T. biserratum (BLUME) v.A.v.R
186
FLORA MALESIANA
[ser. TS vole
11. Secondary rachises abaxially various, but not black with pale ridges; lamina often bipinnate +
pinnatifid.
12. Primary rachis Ae indusia black; ultimate lobes abaxially with very broad and prominent
veins occupying 14-14 of their width (fig. 9)
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 Os the apical sorus of the
segment usually denticulate; texture herbaceous
1. T. denhamii
13. Indusia longer, or, if only 4% 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. Atleast the larger secondary rachises abaxially carinate; basiscopic pinnules of basal pinnae
rarely enlarged and more dissected (fig. 11)
1. Tapeinidium denhamii (HOOKER) C. Cnr.
Ind. Fil. (1906) 631; KRAMER, Act. Bot. Neerl.
15 (1967) 583. — Davallia denhami HooKer,
Second Cent. Ferns (1861) pl. 47.— Microlepia
denhami (HooKeR) Moore, Ind. Fil. 2 (1861)
292. — Lindsaea denhami (HOOKER) METT. ex
KuHN, Verh. Zool. Bot. Ges. 19 (1869) 573.—
Wibelia denhami (HOOKER) KUHN, Chaetopt.
(1882) 346. — Type: MILNE 116, Viti Levu,
Fiji (K).
T. tenuius COPELAND, Philip. J. Sc. 60 (1936)
110, pl. 17. — Type: Brass 3025, San Cristéval,
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, |-2 mm 4g; scales
reddish brown, acicular, to 14% 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-angular
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 1% 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, 144-1 cm wide,
11. T. luzonicum
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, efc. 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 uw.
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
sori is very characteristic, no other species regular-
ly having sori on the terminal lobes.
2. Tapeinidium buniifolium KRAMER, Blumea 15
(1968) 549.—T. 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% mm 9g; scales reddish brown, to c. 1 mm (or
more?) long, to + 4-seriate at base, with long,
acicular, uni- or pauciseriate apex. Petioles stout,
rather lustrous, medium brown, crushed but appar-
ently abaxially upward obtusely bi-angular, c.60cm
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 144-3 cm by
4-5 mm, deeply pinnatifid at base, at the acute
apex like the smaller segments dentato-lobate.
Ultimate lobes \anceolate-ligulate, the larger ones
up to 4 by 134 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; aN? b Sk :
indusium brownish, pouch-shaped, narrowed at NAY 3 Mi Soee Ze Lu
base, often with irregular free edge, 0.3-0.5 mm aN? Gh Se Pa
long and broad. Spores brownish, subellipsoidal, ane? «ei SS. ay
smooth, c. 35 by 26 wu. RNS, APN ee SS
Distr. Only known from the type collection. A YF ANY SSS VA
Ecol. In mountainside woods, c. 700 m. Qs \Y Gy, Ny) Ro tN
@ NVA aay / 7 we SS LL
3. Tapeinidium amboynense (HOOKER) C. Cur. Ss > \\t?, his Wa er
Ind. Fil. (1906) 631; KRAMER, Blumea 15 (1968) Ww 3 WON
549. — Davallia amboynensis Hooker, Sp. Fil. WaT aAMwe e SR 7
1 (1845) 178, pl. 56 C. — Lindsaea amboynensis SN i a\h te) Se Gea
(Hooker) Mett. ex KUHN in Mig. Ann. Mus. A Wie ei Sal Foe
Bot. Lugd.-Bat. 4 (1869) 279. — Wibelia amboynen- 7 AN al Xs. Sra
sis (HOOKER) KUHN, Chaetopt. (1882) 346.— RNY Ae ae
Type: Cur. SMITH s.n., Ambon (K). Qi Ee de
Davallia stenoloba BAKER in Becc. Malesia 3 ' & i rk ee
(1886) 35. — T. moluccanum (BLUME) C. CHR. e Q WP = Pee
var. stenolobum (BAKER) C. Cnr. Ind. Fil. Suppl. 3 5
(1934) 176.—T. stenolobum (BAKER) WAGNER & Wo a WA ;
GRETHER, Un. Cal. Publ. Bot. 23 (1948) 36.— Nee NN {i a fs ;
Type: BeccaRI s.n., Mt Salhutu, Ambon (FI; al / WU \
fragm. in K). eames iL
T. amplum COPELAND, Occ. Pap. Bish. Mus. 15 | es va f 8
(1939) 82, f. 3. — Type: TAKAMATSU 1572, Gara-
sumao, Palau Is. (MICH; dupl. in BISH, K, US).
T. moluccanum C. Cur. Ind. Fil. Suppl. 3
Fig. 6. Tapeinidium amboynense (Hook.) C. CHR. Two pinnules from different parts of the lamina,
x 1% (DE VrigsE 358). — Fig. 7. T. denhamii (Hoox.) C. Cur. Pinnule, x 11% (Brass 3335). —
Fig. 8. T. atratum KRAMER. Pinna, x 34(H. J. Lam 1556). — Fig. 9. T. obtusatum y.A.v.R. Pinna,
x 34 (H. J. Lam 1857).
188
(1934) 176, and of later authors; not Davallia
moluccana BLUME. — Fig. 6.
Rhizome short- to moderately long-creeping,
3-5 mm g; scales castaneous, narrowly triangular,
or with ovate base and acicular apex, to 244 mm
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, abaxially rounded or mostly
upward obtusely or sometimes sharply bi-angular,
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. 25-40 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 abaxially
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,
abaxially terete, often marginate; secondary and
ultimate divisions ascending, asymmetric, with
cuneate base, + decurrent, elongate-ovate, lan-
ceolate, or linear, obtuse to subacute, often 1-3 cm
long and 144-5 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, abaxially 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
tvo) 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,
(G5 &)3) los" 25) [Vip
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.
FLORA MALESIANA
[ser. Il; vole
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 g; 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, abaxially terete, c. 25-75 cm long, slightly
shorter than to twice as long as the lamina. Lamina
bipinnate, or bipinnate ++ pinnatifid, at the base
often tripinnate, elongate-triangular or subpentag-
onal-triangular, less often oblong, c. 20-55 cm
long; primary rachis stramineous to pale brown,
abaxially 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; /Jarger 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, abaxially 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 3-4 cm long and 3-4 mm wide, crenate to
pinnatifid to the middle (or beyond in the basal pin-
nae), With abaxially prominulous, pale costa.
Veins simple in the lobes (crenations) or forked
in the larger ones, somewhat prominulous. Lobes
oblong, or + 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 wu, exceptionally larger.
Distr. Malesia: Japen I., New Guinea (all
Div.); Solomon Is.
Ecol. In forests and thickets, 200-2100 m, often
said to be locally common.
5. Tapeinidium stenocarpum y.A.v.R. Nova Guinea
14 (1924) 52; KRAMER, Blumea 15 (1968) 550.—
Type: H. J. Lam 1442, mountain ridge near
April 1971]
Idenburg R., W. New Guinea (BO; dupl. in K,
L, SING, U; fragm. in US).
Rhizome not very short-creeping, 2-214 mm g;
scales fuscous, elongate-triangular, to c. 3mm 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-
angular 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, + 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 314-4 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 24 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 14-1 times its
width. Spores brownish, ellipsoidal, smooth, c.
35 by 25 uw (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 9;
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
LINDSAEA-GROUP (Kramer)
189
base tripinnate + pinnatifid, elsewhere bipinnate
+ 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 + pinnatifid pinnule,
otherwise the pinnae once pinnate, with the larger
pinnules pinnatifid or pinnatilobate. Secondary
axes dark or, especially the upper ones, pale,
abaxially marrowed-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 44 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, 44-1 mm
long, 1% mm wide, not quite reaching the margin.
Spores medium brown, ellipsoidal, smooth, 60
by 45 wu.
Distr. Only known by the type collection.
Ecol. Terrestrial in mossy forest, c. 2500 m.
7. Tapeinidium atratum KRAmMeR, Blumea 15 (1968)
551.~Type: H. J. LAM 1556, mountain ridge near
Doormantop, W. New Guinea (BO, 2 sh.; dupl.
in L, SING, U).—Fig. 8.
Rhizome not short-creeping, c. 2 mm g; 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
2%-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. Il, vole
attached at the sides, not reaching the margin by
little less than its width. Spores brownish, ellip-
soidal, smooth, c. 35 by 25 yp.
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 (i842) 427. — Wibelia gracilis (BLUME)
Curist, 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 g; scales reddish brown, narrowly tri-
angular, to 24% mm long, to c. 6-seriate at base,
with a long uniseriate apex. Leaves close to c.
1 cm apart; petioles slender, 23-1 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 114 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,
(%4-)1%-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 24% 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; indusium
pouch-shaped, + semi-elliptic, with straight or
slightly lobed edge, 4%—’% 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 wu.
Distr. E. Annam; in Malesia: W. Java, Bali,
Sarawak, Brunei, Celebes, Ceram, Philippines.
Fa ~
Vaya |
\ eRe ie
Wirsoa
L S> QZ : SS
Ce, NN |
Map 1. Distribution of Tapeinidium gracile
(BL.) v.A.v.R.
The only species with a notably disjunct area.
Map lI.
Ecol. Terrestrial and epilithic, in forests, 500—
1300 m.
9. Tapeinidium calomelanos 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 9g; 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 + deeply
pinnatifid, or at the base bipinnate (+ pinnat-
ifid), subcoriaceous or coriaceous, olivaceous-
brown when dry. Primary rachis atropurpureous
or dark castaneous, abaxially terete. Pinnae
ascending, c. 12-20 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 24% cm;
major pinnules (segments) to 18 to a side, lan-
ceolate, ascending, 30 by 244 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 pw.
April 1971]
LINDSAEA-GROUP (Kramer)
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. — Dayvallia oligophlebia BAKER, J. Bot. 26
(1888) 323. — Wibelia oligophlebia (BAKER) CHRIST,
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. Cur. Ind. Fil. Suppl. 3 (1934) 176.—Type:
Brooks 47, G. Bengkaim, Sarawak (SAR,
holotype?; dupl. in BM).
Rhizome short-creeping, 24-2 mm 9g; scales light
brown, lanceolate to elongate-triangular, to 1%
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, 244-12 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 keel. 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 4% mm broad, %4—% mm 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 p.
Distr. Malesia: Borneo (Sarawak, Kaliman-
tan).
Ecol. Terrestrial, in forests and on shaded
cliffs, 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, pl. 60 B f. 2, 3,
5. — Wibelia bipinnata FEE, Gen. Fil. (1852) 331,
nom. superfl. —Type: CuMING 139, p.p., Luzon
(dupl. in B, GH, L).
Lindsaea pinnata (CAv.) METT. ex KUHN var.
191
bipinnata Metr. ex KUHN in Miq. Ann. Mus.
Bot. Lugd.-Bat. 4 (1869) 279. — Type: ZoL-
LINGER 1305, Java (dupl. in HBG, L).
Dayallia philippinnesis HARRINGTON, J. Linn.
Soc. Lond. 16 (1877) 27. — Microlepia philippi-
nensis (HARRINGTON) COPELAND, Polypod. Philip.
(1905) 56. — T. philippinense (HARRINGTON) C.
Cur. Ind. Fil. Suppl. 3 (1934) 176. — Type:
STEERE s.n., Mt Mahayhay, Luzon (K).
Dayallia hosei BAKER, J. Bot. 26 (1888) 323. —
Type: Hose 219, Lambur, Sarawak (K).
T. sumatranum vy.A.v.R. Bull. Jard. Bot.
Btzg III, 2 (1920) 174. — Type: Brooxs 332/S,
Bencoolen, Sumatra (BM; fragm. in BO).
T. biserratum auct. non (BLUME) v.A.v.R.;
Ho.trum, Rev. Fl. Mal. 2 (1954) 339, f. 197;
and of other authors. — Fig. i1.
Rhizome usually short-creeping, to 4 mm 9;
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 + 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 w.
KEY TO THE VARIETIES
1. Petiole less than 1 mm @ at base of lamina;
lamina subtripinnate at the base
1. var. leptophyllym
1. Petiole over 1 mm g 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 2% of their width
3. var. thelypteridoides
1. var. leptophyllum KRAMER, Blumea 15 (1968)
553.—Type: ELMER 14103, Mt Urdaneta, 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
(ser. 1) voln@
FLORA MALESIANA
192
ESET
ema a
ee
“A
sr onrrpr >
AEEDET aM
Ye
Dy oe & ¥)
+ x 7 te fala WY Me WN
epee neers i Sng Tatty 22a Ws”
1247 pl a tle “aa aaa AAG ‘ eA
rhor LAS Ay, eS hhe Nhe ip
\
\
JaEVE
espcreoet 3
EIS EPI _ a2 DIPDLO UTA
SS Ei Pe
AES
ere eeaey,
ete tat |
on~
rr
eet
1s,
Fig. 11. T. Juzonicum (HooK.) KRAMER var. luzonicum. Basal part
le, X 2, portion of rachis, x 34% (SAN 24071).
Fig. 10. Tapeinidium novoguineense KRAMER. Basal part of lamina, x #/;, pinnule, x 2, portion of rach
of lamina, x 2/;, pinnu
x 3% (v. ROYEN & SLEUMER 6394). —
April 1971]
LINDSAEA-GROUP (Kramer)
193
calomelanos (L.) LinK; also resembling TJ. den-
hamii, from which it differs, i.a., 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 %4-2 mm
9 at the base of the lamina. Lamina 18—S0 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 TJ. novoguineense).
Ultimate free (or almost free) divisions ascending,
variable in size and shape, often 14-24% 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. Sori often slightly immersed; indusium
dark, rather rigid, pouch-shaped, 13-4 mm wide,
1,34, 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. biserratum.
3. var. thelypteridoides KRAMER, Blumea 15
(1968) 553.—Type: Brooke 8190, Mt Santu-
bong, Sarawak (L; dup]. in SING, US).
Rhizome short-creeping, with relatively narrow
scales; petioles clustered, 10-27 cm long, not
rarely less than 1 mm g 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 24, the upper ones gradually less,
serrate, then subentire; segments oblong-ligulate,
ascending, the largest c. 12 by 3 mm, obtuse,
shallowly 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.
(4 coll.).
Ecol. On rocks in forest, 400-1300 m.
Malesia: Borneo (Sarawak, Sabah)
12. Tapeinidium pinnatum (Cav.) C. Cur. Ind.
Fil. (1906) 631; Ho_trum, Rev. Fl. Mal. 2 (1954)
339, f. 196; COPELAND, Fern FI. Philip. 1 (1958)
114; KRAMER, Blumea 15 (1968) 553.— Davallia
pinnata Cay. 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 (CAvV.) BERNHARDI ex FEE,
Gen. Fil. (1852) 331, pl. 27 bis B.—Lindsaea
pinnata (CAV.) METT. ex KUHN in Mig. Ann.
Mus. Bot. Lugd.-Bat. 4 (1869) 279.—Type:
NEE s.n., Philippines (also incorr. cited ‘Chile’)
(MA, n.v.; phot. seen).
Divan flagellifera HOOKER & Geena
Ic. Fil. (1831) pl. 183.—Type: WALLICH s.n.,
Pulu Penang (n.v.).
Davallia serrata RoxsB. ex GRIFF. Calc. J. Nat.
Hist. 4 (1844) 514, non WILLp. (1810).—Type:
ROXBURGH s.n., Pulu Penang (n.yv.).
Wibelia javae F&e, Gen. Fil. (1852) 331.—Type:
KOLLMANN 5S.n., Java (n.yv.).
Dayallia firmula BAKER, Ann. Bot. 8 (1894)
123.—Microlepia firmula (BAKER) C. Cur. Ind.
Fil. (1906) 426.—Tapeinidium firmulum (BAKER)
C. Cur. Ind. Fil. Suppl. 3 (1934) 176.—Type:
HANCOCK 72, Barisan Range, Sumatra (K).
Rhizome short- to rather short-creeping, 2-3
mm @; 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 @g at base of lamina,
abaxially at least in the upper half obtusely
or more often acutely bi-angular, c. 10-35 cm
long, 4-24 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
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 yw.
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 7. 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 Hottrum, Rev. Fl. Mal. 2
(1954) 339.—Davallia biserrata BLUME, En.
Pl. 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, /.c. 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 @g;
FLORA MALESIANA
[ser. Il, volwie
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, 24-1'% times the length of the
lamina. Lamina simply pinnate, oblong, with c.
10-18 free pinnae to a side and a pinnatifid,
basally 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, 44-1 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
+ 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, %-1 mm long, % mm wide,
almost reaching the margin. Spores light brown,
ellipsoidal, smooth, c. 36 by 28 yw. :
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: EscritTor 21173, Palanan Bay,
Luzon (L; dupl. in BO, BRI, GH, MICH, SING,
US). —Fig. 14.
Rhizome rather short-creeping, 2-3 mm @;
scales castaneous, lanceolate, long-acuminate,
to 24% mm 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, 44 mm wide, 44-*4 mm
long, with convex free edge and + convex base,
l¥, mm 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. Cur. Basal part of lamina, x 1, portion of pinna, x 3h
portion of rachis, x 5 (BAMLER Ros. 115).— Fig. 13. T. prionoides KRAMER. Basal part of lamina, x %,
portion of pinna, x 3, portion of rachis, x 5 (BUNNEMEIJER 1910).
196 FLORA MALESIANA [ser. II} voli
-)
she |
| E.V.'66
Fig. 14. Tapeinidium acuminatum Kramer. Upper part of lamina, x 1%, portion of pinna, Xx 3, portion
of rachis, x 5 (EscritTor 21173).
ee
April 1971]
197
15. Tapeinidium longipinnulum (CEsATI) C. CHr.
Ind. Fil. Suppl. 3 (1934) 176; CopELAND, Philip.
J. Sc. 78 (1949) 23: KRAMER, Blumea 15 (1968)
555.—Davallia longipinnula Crsati, Rerdic. R.
Accad. Sci. Fis. Mat. Napoli 16 (1877) 26;
Beccari, Malesia 3 (1886) 35.—Type: BECCARI
s.n., Ramoi, New Guinea (FI).
Davyallia intramarginalis CESATI, Rendic. R.
Accad. Sci. Fis. Mat. Napoli 16 (1877) 29.—
Type: Beccari s.n., Mt Arfak, New Guinea (FI).
T. marginale 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 @;
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, %-2, most often 24-1 mm
long, c. % mm wide, very nearly equaling the
margin, often strongly bulging at maturity.
Spores brownish yellow, oblong, smooth, 30-35
by 24-26 wu.
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, pl. 60 = Xyropteris
stortii (v.A.v.R.) KRAMER.
Tapeinidium moluccanum (BLUME) C. CHR.
Ind. Fil. Suppl. 3 (1934) 176 = Saccoloma sp.
(see under 7. amboynense).
Tapeinidium tenue (BRACKENR.) COPELAND,
Bull. Bish. Mus. 59 (1929) 69 = Saccoloma
sp. (see under 7. 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 fig.—Schizoloma
Stortii v.A.v.R. Bull. Jard. Bot. Btzg II, 16
(1914) 36; Handb. Suppl. 1 (1917) 214. —Type:
AMDJAH (vy. 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, pl. 60.—Type: BaArRT-
LETT 673la, near waterfall Si Saliang, Asahan,
Sumatra (dupl. in GH, L, MICH).
Rhizome rather short-creeping, in full-grown
plants to 144,cm@; scales golden-brown, elongate-
triangular, acuminate, to 5 by 144 mm, to c. 18-
seriate at base. Petioles close, to a few mm apart,
to 40 cm long, to 14% cm g 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, 142-3 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 1 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
outward, mostly 2 or 3 times forked, free; auricles
of pinnae with a costule. Upper pinnae in large
leaves 4-10 cm long; terminal pinna free, to c.
15 cm long, unequally and deeply trilobate, with
long-acuminate lobes, the central shank much
longer than the lateral ones. Sori plurinerval but
interrupted in young, continuous in adult plants,
absent only from the extreme base and apex of
FLORA MALESIANA
[ser. II, vol. 14
paraphyses present among the sporangia. Spores
medium brown, monolete, bean-shaped, smooth,
c. 50-60 by 30-35 wu.
Distr. Malesia: Sumatra (3 coll.), Borneo
(Kalimantan) (type coll.).
Ecol. In ravines by streams, c. 300-500 m;
presumably a rheophyte.
Note. An excellent series of specimens from
Sumatra (SURBECK 49, L, U), including young
as well as full-grown plants, shows that Tapei-
nidium bartlettii COPELAND is the juvenile form
of X. stortii.
the pinnae and the apex of the auricle; indusium
brown, entire or wavy, %4-’3 mm wide, not
reaching the margin by about its own width,
+ reflexed at maturity. Multicellular filiform
4. LINDSAEA
DRYANDER in J. E. Smith, Mém. Ac. Turin 5 (1793) 401; Trans. Linn. Soc. 3
(1797) 39; Hooker, Sp. Fil. 1 (1846) 203; J. Smiru, Hist. Fil. (1875) 267; Diets
in E. & P. Nat. Pfl. Fam. I, 4 (1902) 219; v.A.v.R. Handb. (1908) 260; Suppl.
(1917) 202; Hottrum, 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;
Ho.LtTtuMm, Rev. FI. Mal. 2 (1954) 321; KRAmegr, Act. Bot. Neerl. 6 (1957) 97;
CHING, FI. Reip. Pop. Sin. 2 (1959) 257; KRAMER, Blumea 15 (1968) 557.—Davallia
J. E. SMITH sensu Hooker, Sp. Fil. 1 (1845) 151, p.p.—Schizoloma GAupb. 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) 218, p.p.; v.A.v.R. Handb. (1908) 275; Suppl]. (1917)
214; HoLttum, Rev. FI. Mal. 2 (1954) 342; CuinG, FI. Reip. Pop. Sin. 2 (1959)
272.—Isoloma J. SmitH, Hook. J. Bot. 3 (1841) 414; Hist. Fil. (1875) 227;
COPELAND, Gen. Fil. (1947) 55; Philip. J. Sc. 78 (1949) 24; HoLtrum, Rev. FI.
Mal. 2 (1954) 336; COPELAND, Fern FI. Philip. 1 (1958) 100.—Odontoloma J.
SmitH, Hook. J. Bot. 3 (1841) 415, nom. subnud.; in Hooker & Bauer, Gen. Fil.
(1842) pl. 114 B; Fée, Gen. Fil. (1852) 329; J. Smiru, Hist. Fil. (1875) 269.—
Synaphlebium J. SMitH in Hooker & Bauer, Gen. Fil. (1842) pl. 101; Hist. Fil.
(1875) 268.—Lindsaenium (or Lindsaynium) FEE, Mém. 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., 24 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 of ) 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 Synaphlebium, 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, Blumea
15, 1968, 557 seg.). 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). Spp. yp ariel We . . . 1. Sect. 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 + sclerotic.
4. An articulation at the base of each pinnule. Spp.36-41 . . . . . . 8. Sect. Isoloma
4. No articulations at the pinnule-bases. Sp.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. Veins anastomosing, sometimes only irregularly. Spp.15-30. . . . 3. Sect. Synaphlebium
6. Veins free.
7. Spores monolete; lamina simply pinnate, with abaxially rounded rachis and (except in small
forms) interrupted sori. Sp.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. Spp. 42-43 : = oe 9. Sect. 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. Spps 31-32 Be A, ee, Be, ns Seer amdsaes
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 114, 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 i ina 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. Sp.52. . . .- . . . Il. Sect. Pseudolancea
12. Pinnules incised, with interrupted sori, or with only one short sorus near the apex, otherwise
sterile, Sppo.53=59s iri ax Reh ee en slate lhe eprint seer Lindsaenium
200 FLORA MALESIANA [ser. II} volst=
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) igs ' 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 TE SOT Oe ie eae i 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 . . ee ee Cr! Group E
5. Pinnules continuous with the rachis (sect. Psammolindsaea) J) Biel TM Ee 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.
fi Meinsiirees
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... ee 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% 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 as sterile leaves not usually present, their pin-
nules not sharply dentate. . mh oS teccheal cpl te ictal ae ee 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 seer. Trepidolindvace) 34. L. adiantoides
11. Simply pinnate, with abaxially bi-angular rachis; or bi-(subtri-)pinnate; spores trilete.
12. Simply pinnate; pinnules up to 2% times as long as wide; lamina basally mostly at least
somewhat reduced. . . . es Group F
12. Simply pinnate; at least some pinnules over 2 2¥, Ramee as long; as wide; lamina truncate at the
LEC Ale as i i a ee eee GD CH
IOrmBipinnatem eles As OR OS an Se Oe eG oe ee Groupsbs
(AV eINnSkanastOmOsin Core ots hy of es 4 015) eee call aa 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 . . . ‘amt 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-angular; 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 poke lanceolate undivided
apical portion . . pe 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 TTT: be the creel es De See IS ensiiona
Group B (sect. Temnolindsaea)
1. Petiole and rachises abaxially keeled .. Sail ee 14. L. tenuifolia
1. Petiole and rachises abaxially bi-angular, rarely only obscurely so.
PaeBasall(primary)) pinnae stronglyeduced) (fies 25)) mes ee Oe king
2. Basal pinnae hardly or not reduced.
3. Pinnules incised beyond 34 of their width, with capillary segments 0.2-0.4 mm wide, wider at the
sorus, the wing connecting them as wide (fig. 28); soriuninerval . . . . 13. L. polyctena
3. Pinnules at the base incised beyond the middle, the segments not capillary, 0.3-0.8(—1) mm wide,
connected by a wing of 44-1 mm; 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 e of the width of the pinnules: Jobes broader than long.
Ones lammiapart = =. Se) 0 AP ce Ao ce ee a ace Lok enarumae
6. Veins 4% mm or less apart Seek? Meese tance. on Pee ea See 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, 2-242 mm wide; veins
adaxially impressed, abaxially prominulous; larger pinnules rarely entirely free-veined; sterile
pinnules sinuate-dentate; petioles et pale, usually brown, abaxially obtusely to acutely bi-
angular shoes Se 27. 1. crispa
2. Sori continuous or interrupted, the i incisions of the pinnules not reaching iy mm deep in fully fertile
pinnules; pinnules very little narrowed close to the apex, 10-12 mm long, 3-4 mm wide, often 344-4
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, + 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 21-3 times as long as wide; petiole stramineous. . oe os 2S. cultrata
4. Upper pinnules 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 2seriesofareoles . . . 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 24% to over 3 times as long as wide.
8. Larger pinnuléss6—20smnvilong « (4Ge. i os ae Oe eee ee, Pee ape
8. Larger pinnules to 12 mm long. . 2 bya beepers Bee GSE napaea
3. Sori of larger pinnules interrupted by incisions of the margin.
202 FLORA MALESIANA iser: IE. vole?
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 2-314 cm long, 3-314 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 5mm long... . os » = = « 6, See phewittn
13. Pinnules more elongate, or, if not, at least 7 mm long.
14. Pinnae (if any) strongly ascending; pinnules 7-10 by 344-5 mm, 114-2% 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 2%
times as long as wide, or the indusium closer to the margin.
15. At least the inner incisons reaching to 4% or 1% of the width of the pinnules; pinnules
translucent; all lobes and pe Cre convex; indusium reaching the margin
or very nearly so. . . Mace: 21. L. lobata
15. Pinnules more shallowly incised, or, if the i inner incisions s reach 1 1,, 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 ‘ pee 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 usa ae een SO ee Oe oe eee OInmapaes
Group D (sect. Lindsaea)
1. Pinnules to 11 mm long, to 5mm wide. . . ee See to Pee on OS 31. L. borneensis
1. Pinnules to 20(-35) mm long, to 6(-13) mm wide 5. be ap ee eke si auch on (SZ leony pHOra
Group E (sect. Isoloma)
1. Pinnules 5 times or more as ont as broad, 3-4 mm wide oper from basal auricles), with con-
tiguous bases. : 41. L. divergens
1. Pinnules relatively much shorter, | or, if up to 5 times as silones as wide, 8 mm n 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 5 pote 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, haa keeled to the base; upper Cees little or not reduced,
terminal division free . . i Ge 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 244 times as long as wide . 36. L. gueriniana
4. Rachis lustrous, reddish to dark brown; es CBee. the ees ones 5—8 mm long, less than
114% 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,
tomatimesiastlongiasiwidess ann men ae e 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. . . . . . . . . . . . . 43.1. bakeri
April 1971] | LINDSAEA-GROUP (Kramer) 203
Group G (subg. Odontoloma)
1. Rhizome wiry, not over 1144 mm g, 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 114-3 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.. . . ne \icteg ag COOXOUp iG
2. Veins free, or, if anastomosing, the largest pinnules less than 144 cm long; upper pinnules gradu-
ally and strongly reduced.
4. Pinnules deeply pinnatifid, the segments connected by wings 144-14 mm wide (fig. 23)
62. L. roemeriana
4. Pinnules shallowly incised, or, if more deeply incised, the lobes much more broadly connected.
60. L. pulchella
1. Rhizome not wiry, 14-2 mm g, 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 corsiderably beyond the receptacle.
7. Veins anastomosing rer, 5) ee” ae tee) |e ce Cr a Poser 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 44 mm 9g; 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); soriuninerval. . 51. L. capillacea
11. Rhizome 34-2 mm g; ultimate pinnule-lobes 4,-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 44 mm width (fig. 48)... . . 49. L. carvifolia*
12. 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%4 or 44, the lobes evenly narrowed from
base to apex (fig. 49). . . . . 44, L. glandulifera
15. Larger pinnules 15 mm or more long, with truncate or apically rounded lobes, or, if shorter,
much less deeply incised, or the lobes + parallel-sided. _ SI, PAG aE repels =
5. Lamina of all, most, or at least some leaves bipinnate.
KGa eSormcontinuous 6s sO See eke a Be nr een, wey ge Eee io 52. L. parasitica
16. Sori interrupted.
17. Secondary rachises abaxially rounded. . . . . . . . +. +. +. +‘54,L. monocarpa
17. Secondary rachises abaxially carinate . . eee 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. rigida
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. versteegii 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. Il, vole
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
- « » Sdallmicidac
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
, 56. L. regularis
Adaxial groove of the primary rachis narrow, occupying much less than half its width; most
1,—3 times its
width; not rarely simply pinnate leaves present alongside the bipinnate ones 57. L. microstegia
59. L. versteegii
20.
the margin by less than its width; bipinnate .
20.
width; bipinnate
20.
pinnules entirely fertile; veins free; indusium not reaching the margin by 1}
19. Pinnules incised considerably beyond the middle.
21. Segments capillary, c. ‘ls a; mm wide, all forked.
21. Segments not capillary 2
—2 mm wide, the outer ones often mot forked 58. L. rosenstockii
Subgenus Lindsaea
1. Section Schizoloma
(GAUD.) KRAMER, Act. Bot. Neerl. 15 (1967) 571.—Schizoloma Gaupb. Ann. Sc. Nat. 3 (1824)
507.
Type species: Schizoloma billardieri 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. orbiculata (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. Davalliastrum. 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
(1968) 563.—Type: HALLIER 3244, Mt Amai
Ambit, Borneo, Kalimantan (BO).—Fig. 19.
Rhizome short-creeping, c. 0.6 mm g; scales
reddish brown, c. 144 cm long, narrowly lanceo-
late, biseriate at base, the apical uniseriate part
consisting of 1-3 cells. Leaves close; petioles
2-4 cm long, somewhat shorter than the lamina,
adaxially pale, channelled, abaxially dark brown,
sharply bi-angular with narrowly paler edges
and flat faces, subterete at base. Lamina herbace-
ous, oOlivaceous when dry, ovate with truncate
base, to 5-6 cm long, 3-414 cm wide, pinnate +
deeply pinnatifid, without conform terminal pinna.
Major pinnae c. 10 to a side, spreading or slightly
ascending, less than their width apart, the larger
ones 20 by 7 mm, acuminate, inequilateral,
deeply pinnatifid. Segments cuneate to ligular,
2 or in larger pinnae 3 on the acroscopic side,
3-4 by 2 mm,’with only a very narrow wing on
the opposite, basiscopic side; upper segments of
acroscopic and all (or all but one) of basiscopic
side narrowly cuneate, all narrowed at base and
connected by wings of leaf-tissue; uppermost
pinnae and segments reduced, confluent. Apex
of segments rounded-truncate, entire or sinuate;
sterile segments often subacute. Largest segments
often bilobed. Costae evident, stramineous;
veins immersed, evident, single or in larger
lobes paired, one sometimes forked. Sori uni-
to trinerval; indusium greyish, 15-144 mm long,
0.4 mm wide, suborbicular to oblong, free at the
narrowed sides, subentire, not reaching the margin
by a little less than its width, not reflexed at
maturity. Spores medium brown, trilete, smooth,
Cel Z yu:
Distr. Only known from type collection. No
ecological data.
2. Lindsaea bouillodii CHRist, Not. Syst. 1 (1909)
59.—Type: BouILLop 48, Cam-chay Mts, Cam-
bodia (P).
L. orbiculata (LAMK) METT. ex KUHN var.
odontosorioides COPELAND, Philip. J. Sc. 6 (1911)
Bot. 138.—Type: Brooxs 19, Tringos, Sarawak
(MICH).
L. orbiculata (LAMK) METTY. ex KUHN var.
sumatrana ROSENSTOCK in Fedde, Rep. 13 (1914)
214.—Type: WINKLER 55 (Ros.-exs. 117), Batak
Lands, Sumatra (S-PA; dupl. in BM, L, P).
L. cambodgensis auct. non CHRIST; KRAMER,
Blumea 15 (1968) 563.
L. tenera auct. non DRYAND. (or Schizoloma
tenerum) of other authors, e.g., HoLttruM, Gard.
Bull. S. S. 5 (1930) 64; Rev. Fl. Mal. 2 (1954)
348, f. 201.—Fig. 21.
Rhizome moderately to very short-creeping,
1144-2 mm 9; scales ferrugineous, very narrowly
triangular, to 24% mm long, to 5-seriate at base,
with long uniseriate apex. Leaves close to cluster-
* Epiphytic specimens with unusually long rhizomes of species of sect. Synaphlebium (Group C) will
run to this heading; they have regularly anastomosing veins.
April 1971] LINDSAEA-GROUP (Kramer) 205
AW?
(
ye if
Qh
f sy sy
AY
{
(
iG
iS
}
AW)
(\
NWS
(
t
Fig. 15. Lindsaea orbiculata (LAMK) METT. ex KUHN var. orbiculata. Plant, « %%, pinnule, x 1%
(TANAKA & SHIMADA 13522).—Fig. 16. L. gomphophylla BAKER. Lamina, x 1 (HOSE? s.n., SAR).— Fig.
17-18. L. heterophylla DrYAND.; fig. 17. Lamina, x 1% (BAKHUIZEN VAN DEN BRINK 7153); fig. 18.
Sterile pinnule, nat. size (PETERSEN s.n., L).—Fig. 19. L. hemiacroscopica KRAMER. Basal pinna, 2
(HALLIER 3244).
206
FLORA MALESIANA
[ser. II, volaté
ed; petioles 8-40 cm long, to 14% 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,
134-2% 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. 7-12 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, | 4-2 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 | or 2, the outer with | incision, sometimes the
lobes shallowly incised again; incisions 44-14% mm
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, 4-24 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 wu.
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 Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1869)
279; Hottrum, Gard. Bull. S. S. 5 (1930) 64;
TaGAwa, Act. Phytotax. Geobot. 6 (1937) 33,
f. 3 D-G; CopeLAND, Fern FI. Philip. 1 (1958)
112, p.p.— Adiantum orbiculatum LAMK, Encycl. 1
(1783) 41.—Schizoloma orbiculatum (LAMK)
KuuHN, Chaetopt. (1882) 346; Hortrum, Rev.
Fl. Mal. 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. flabeliulata. DryANbD. Trans. Linn. Soc. 3
(1797) 41, pl. 8 f. 2.—Lectotype: NELSON s.n.,
Macao (BM).
L. polymorpha WALL. ex HOOKER & GREVILLE,
Ic. Fil. (1828) pl. 75.—L. flabellulata DRYAND.
var. polymorpha (Hook. & GREvV.) HOOKER, Sp.
Fil. 1 (1846) 211.—L. orbiculata (LAMK) METT. ex
KUHN var. polymorpha (Hook. & GREy.) v.A.v.R.
Handb. (1908) 270.—Type: a specimen without
data from Herb. HOOKER (K).
L. montana CoPpELAND in Perkins, Fragm. FI.
Philip. (1904) 182, non FEE (1866).—L. covelandi
C. Cure. Ind. Fil. (1906) 392.—Type: CoPpELAND
230, Mt Mariveles, Luzon (MICH; dupl. in B).
L. bonii Curist, Not. Syst. 1 (1909) 187.—
Type: Bon 8, S. Tonkin (P).
L. longipes C. Cur. & TARDIEU-BLoT, Not.
Syst. 5 (1936) 263; Fl. Gén. I.-C. 7 (1939) 125,
f. 15 3-4.—Type: PoILANE 8208, Nhatrang,
Annam (P).
Rhizome short-creeping, 1-14 mm g; 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, -- gradually passing into the non-
conform terminal pinna. Terminal pinnules (seg-
ments) + rhombic. Sori continuous or interrupted;
indusium minutely to strongly and irregularly
erose, 4% 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 mu.
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
INO 6 og o « 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; /amina
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 \4-elliptic, often 24-1 by % 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%
cm long and %4-34 cm wide, obtuse, less often
acute, cuneate-flabellate, 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 sterile 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 Jarger
pinnate pinnae, very asymmetrically flabellate-
suborbicular, with broadly rounded upper/
outer and + concave, descending lower margin,
then often 1-14 cm long and broad; in sub-
bipinnate laminas some of the lower (but not
necessarily the lowermost) pinnae pinnatifid
or pinnate, with some cuneate-flabellate or 14-
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,
noy.—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,
n.v.).
? L. montana COPELAND.—L. copelandi C. Cure.
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).—Fig. 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. Il; voleue
Petiole and rachis dark, abaxially terete or the
rachis obtusely bi-angular; secondary rachises
abruptly pale. Sterile margin less acutely dentate.
Spores c. 30 pw.
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-
biculata.
5. Lindsaea javanensis BLUME, En. Pl. Jav. (1828)
219.—Schizoloma javanense (BLUME) HOLTTUM,
Rev. Fl. Mal. 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, pl. 63 c.—L. orbiculata
(LAMK) METT. ex KUHN var. gigantea (HOOKER)
Mett. ex KUHN in Mig. Ann. Mus. Bot. Lugd.-
Bat. 4 (1869) 279.—L. tenera DRYAND. var.
gigantea (HOOKER) Hoittum, Gard. Bull. S. S.
5 (1930) 65.—L. gigantea (HOOKER) C. Cure.
Bot. Jahrb. 66 (1933) 53.—Lectotype: GRIFFITH
s.n., Khasya and Assam (K).
Rhizome short-creeping, 1-2 mm g; scales
medium brown, lanceolate, long-acuminate, to
144 mm 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 Jamina, 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% to (in large
leaves) 13/4, 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 py.
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. Holl.
(1810) 156.—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. subtripinnata 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, 1-14, mm 9,
rather thinly and deciduously scaly; scales
yellow, ovate-triangular, to c. | mm long, to
6-seriate at base, a short apical portion uni-
seriate. Leaves to 4 cm apart; petioles stramineous
to fawn-coloured, 10-40 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 toa side, most or
all subopposite, the largest, basal ones 244-10 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
3>
| /
Fig. 20. Lindsaea media R.Br. Lamina, x 2/, (BRASS 8491).— Fig. 21. L. bouillodii Curist. Lamina,
x ?/; (CUMING 399). — Fig. 22. L. bakeri (C. Cur.) C. Cur. var. bakeri. Lamina, x 2/; (BRASS 13651). —
Fig. 23. L. roemeriana ROSENSTOCK. Leaf, x 2/; (v. ROMER 1137).
210
FLORA MALESIANA
[ser. II, vol. 13
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 3% 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. % 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 wu.
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, pl. 8 f. 1—Adiantum hetero-
phyllum (DRYAND.) PotretT, Encycl. Suppl. 1
(1810) 139, non CoLENSO (1888).—Schizoloma
heterophyllum (DRYAND.) J. SmiTH, Hook. J.
Bot. 3 (1841) 414; Hottrum, Rev. Fl. Mal. 2
(1954) 345.—L. ensifolia Swartz var. heterophylla
(DRYAND.) BENTH. FI. Austr. 7 (1878) 722.—
Schizoloma ensifolium (SwArtTzZ) J. SMITH var.
heterophyllum (DRYAND.) DomIn, Bibl. Bot. 20
(1915) 77.—Schizolegnia heterophylla (DRYAND.)
ALSTON, Bol. Soc. Brot. II, 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, pl. 52.—Type: MILLETT
s.n., Macao (n.v.).—Fig. 17, 18.
Rhizome rather to very short-creeping, 114-3
mm g; scales medium brown, very narrowly
triangular, to 24% mm long, to 7-seriate at base,
long-acuminate, the apical uniseriate part rather
long. Leaves close; petioles (3—)10-50 cm long,
142 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, 34-14% cm
wide, 114-8 times as long as wide; larger pinnate
primary pinnae 8-18 cm long, (34-)1%-6 cm
wide, 2-9 times as long as wide, triangular to
linear, with 1-15 pinnules, these %—5 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 % 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 %-1 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
ee
Map 3. Distribution of Lindsaea heterophylla
DRYAND.
April 1971]
LINDSAEA-GROUP (Kramer)
214
pale yellowish, trilete, smooth, c. 30 «4 (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), Philippines (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 FI. 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 & PoOsTHUMUS, Varenfl. Java (1939) 110,
f. 20; TARDIEU-BLoT & C. Cur. FI. Gén. I.-C. 7
(1939) 129, f. 15 1-2; Hottrum, Rev. FI. Mal.
2 (1954) 346, f. 200.—Schizolegnia ensifolia
(Swartz) ALSTON, Bol. Soc. Brot. II, 30 (1956)
24.—Type: coll. unknown, Mauritius (S-PA).
Pteris stricta POIRET in Lamk, Encycl. 5 (1804)
713.—L. pteroides DESvAUxX, Prod. (1827) 312.
Type: COMMERSON s.n., Mauritius (‘Ile de France’)
(P).
L. lanceolata LaxBitt. Nov. Holl. Pl. Sp. 2
(1806) 98, pl. 248 f. 1.—Adiantum lanceolatum
(LABILL.) PorreT, Encycl. Suppl. 1 (1810) 134,
non FEE (1852).—Schizoloma lanceolatum (La-
BILL.) PRESL, Tent. Pterid. (1836) 132.—Schizo-
loma ensifolium (SWARTZ) J. SMITH var. lanceolata
(LABILL.) R. BONAPARTE, Notes Ptérid. 13 (1921)
259.—Schizoloma billardieri GAaup. Ann. Sc.
Nat. 3 (1824) 508. nom. superfl.—L. billardieri
(GAUD.) CARRUTHERS ex SEEMANN, FI. Vit.
(1873) 337.—Type: LABILLARDIERE s.n., Cape
Van Diemen, Australia (P; identity of specimen
uncertain).
L. erecta MIRBEL in Lamk & Mirbel, Hist.
Nat. Vég. 5 (1803) 126, non METTENIUS (1861 ).—
Type: coll.?, Réunion (7.¥.).
Pteris angulata PRreEsL, Rel. Haenk. 1 (1825)
54, nom. illeg., incl. L. lanceolata LABILL.—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 KuNzeE, Linnaea 18 (1844) 121.—
Type: CUMING 369, Malacca (B; dupl. in GH, L,
SING, W).
L. griffithiana Hooker, Sp. Fil. 1 (1846) 219,
pl. 68B.—Schizoloma_ griffithianum (HOOKER)
FEE, Gen. Fil. (1852) 108. —Type: GRIFFITH s.n.,
Mergui, Burma (K).
L. pentaphylla Hooxer, Sp. Fil. 1 (1846) 219,
pl. 67A.—Schizoloma pentaphyllum (HOOKER)
FEE, Gen. Fil. (1852) 108.—Type: BYNOE s.n.,
Australia (“New Holland’) (K).
L. oligoptera KUNZE, Bot. Zeit. (1846) 445.—
Type: ZoOLLINGER 1513 (‘& 1515’), Java (B;
dupl. in HBG, L, Z).
Schizoloma javae Fre, Gen. Fil. (1852) 109,
pl. 29 f. 1.—Type: ZoLLINGER 1504, Java (n.y.).
L. schizoloma EttincsH. Farnkr. 3 (1865)
213, pl. 145 f. 4, pl. 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
f. typicum Domin, type: C. B. CLARKE s.n., several
coll. from India; f. pteroides DomIn, types: GRIF-
FITH 173, Mergui, Burma, and WALLICH s.n.,
Singapore; f. praelongum DOMIN, types: KUNST-
LER 1881, Singapore, and coll. ?, Mauritius (7.v.).
Schizoloma ensifolium (Swartz) J. SMITH var.
borneense DomiIn, I/.c.—Type: coll. ?, Borneo
(n.v.).
Schizoloma ensifolium (SWARTZ) J. SMITH var.
clarkeanum Domin, l.c. 76.—Type: CLARKE S.n.,
N. India (n.y.).
Schizoloma ensifolium (SwARTZ) J. SMITH var.
longipinnum Domin, l.c. 76, with f. typicum
DomIN, type: GOMEZ s.n., Javoy, India; f. sub-
simplex DOMIN, types: GRIFFITH s.n., Malacca,
and WALLICH 92, Ceylon; f. griffithianum Domin,
l.c. 77, type: GRIFFITH s.n., Mergui, Burma (7.y.).
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) 579.—Schizoloma agatii BRAC-
KENR. U.S. Expl. Exp. (1854) 216, pl. 30 f. 1.—
Type: U.S. Expl. Exp. s.n., Fiji (dupl. in K).
Schizoloma ensifolium (SwARTZ) J. SMITH var.
intercedens Doin, Bibl. Bot. 20 (1915) 80, pl.
14 f. 8, pl. 15 f. 3.—Type: DomIn s.n., Yarraba,
N. Queensland (v.v.).
Rhizome not very shortly creeping, 14%. mm
thick; scales as in the next subspecies. Leaves
PND
FLORA MALESIANA
fser. 1 vol
%4-1 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 subauriculate
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 1% 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 ML.
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 has often been confused
with L. heterophylla, e.g. by DomiIn, /.c. Bipinnate
forms may closely resemble that species but may
be told apart by relatively wider pinnules, notably
the upper ones, and more regularly anastomosing
veins. Ssp. ensifolia and ssp. agatii overlap geo-
graphically and are connected by intermediates
whereas L. heterophylla overlaps only with ssp.
ensifolia from which it is sharply distinct.
2. ssp. ensifolia.
Rhizome sometimes short-creeping, (1—)1%4—2
(24%) mm g; scales light reddish brown, narrowly
triangular, to 2 mm long, to 5-seriate at the base,
about the apical 14 uniseriate. Leaves to 2 cm
apart. Petioles c. 10-35 cm long, 4-1 times as
long as, rarely longer than the lamina, strami-
neous to reddish brown, rarely darker, abaxially
at least upward bi-angular and sometimes also
sulcate, if dark not or hardly pale-margined.
Lamina very variable, c. 15-45 cm long, mostly
once pinnate, rarely simple, very rarely subbipin-
nate; if simple lanceolate, c. 10 by 14-3 cm, or
linear, c. 10 cm by 3-10 mm. Pinnate lamina with
the rachis like the upper part of the petiole,
abaxially sharply bi-angular and mostly also
sulcate. Lateral pinnae one odd one to 12 toa
side, most often in 2-8 pairs, not contiguous,
spreading to strongly ascending, the larger ones
usually subpetiolulate, lanceolate to linear, +t
evenly narrowed from base to apex, subacute
to acuminate, 10-22 cm long, 4-25 mm wide, 4 to
over 25 times as long as wide (the great vari-
ability at least in part due to the presence of
juvenile yet fertile plants), the base broadly to
narrowly cuneate, the basiscopic side usually
slightly longer and narrower. Texture herbaceous
to chartaceous, rarely thicker; colour dark green
or olivaceous when dry. Sterile leaves (not com-
mon) with fewer, relatively broader pinnae;
sterile margin (in fertile pinnae often present
at the apex) serrate, less often subentire. Upper
pinnae little reduced, in large leaves c. 13 the size
of the lower ones; terminal pinna conform, with
asymmetric base, of the size of the larger lateral
ones, free or slightly connected with 1 or 2 not
lobe-like upper pinnae. Costa stramineous, not
carinate. Areoles of veins 14—-1144(-2) mm wide.
Indusium entire, 0.4-0.5 mm wide, strongly re-
flexed and concealed at maturity. Spores light
yellow, c. 25-28 wu.
Distr. Uncommon near the coasts of W. and
E. Africa; Madagascar and Mascarenes, Seychel-
les; S. and E. India, Ceylon, Ryu Kyu Is., SE.
China to Malesia: throughout, including some of
the Lesser Sunda Is., but rare and local in the
Philippines, Celebes, and most of New Guinea;
Micronesia, tropical Australia, to the Solomon
Is. and New Caledonia, Hawaii.
Ecol. Terrestrial or epilithic, in swampy to
moderately dry places, exposed or in light shade,
from sea-level up to 1000 (rarely up to 1750) m;
common in most parts of its Asiatic area.
Note. Various authors, notably Domin, /.c.,
tried to classify the confusing array of forms in
this subspecies. These are, however, connected
by all kinds of intermediates and defy description.
The lamina of simple forms probably represents
the terminal pinna.
3. ssp. coriacea (v.A.v.R.) KRAMER, Blumea 15
(1968) 564.—Schizoloma coriaceum v.A.v.R. Bull.
Dép. Agr. Ind. Neéerl. 18 (1908) 10.—Type:
HALLIER 1934, between S. and G. Kenepai,
Kalimantan, Borneo (BO).
Rhizome 2-3 mm g, mostly not short-creeping;
scales to 2144 mm long, to 8-seriate at the base,
with a short uniseriate apex. Petioles atropur-
pureous, shining, abaxially terete or upwards
obtusely carinate. Lamina rarely simple, mostly
once pinnate, with 1-5 pinnae to a side and a
conform terminal one. Rachis like the petiole,
abaxially terete or obtusely carinate, rarely ob-
tusely bi-angular. Pinnae lanceolate, acute or
acuminate, chartaceous or subcoriaceous, 10—25
cm long, 1-2'% cm wide, 6-15 times as long as
wide, widest above the scarcely unequal base,
not rarely sterile, then entire; terminal pinna the
largest. Costa stout, stramineous to dark brown.
Areoles in 2 or 3 rows, often to 2 mm wide. Jn-
dusium entire, usually not over 0.3 mm wide.
Spores yellowish brown, trilete, smooth, c. 20 yw.
Distr. Malesia: Johore, Singapore, Riouw
April 1971]
LINDSAEA-GROUP (Kramer)
213
Is., Sumatra, Borneo (Sarawak, Kalimantan);
a single collection doubtfully from Java.
Ecol. In swamp forests, on acid soil.
Note. It is feasible that the criterium of the
rachis structure used here for distinguishing ssp.
coriacea from the exceedingly variable ssp. ensi-
folia is not entirely reliable. There are a few col-
lections that match ssp. coriacea in other characters
but have abaxially -- pronouncedly bi-angular
axes.
2. Section Temnolindsaea
KRAMER, Act. Bot. Neerl. 6 (1957) 176.
Type species: Lindsaea klotzschiana Moritz.
Distr. Neotropical, and 7 spp. in Malesia and New Caledonia.
Taxon. The short-creeping rhizome, bipinnate lamina with conform terminal pinna, dimidiate
free-veined pinnules, interrupted sori, and trilete spores are characters of all species. Otherwise the
paleotropical species do not have very much in common with the neotropical ones, and it may well
be that they are more closely related to sect. Synaphlebium.
9. Lindsaea kingii COPELAND, Philip. J. Sc. 6
(1911) Bot. 83; ibid. 78 (1949) 19.—Type: KiNG
241, Papua (MICH; fragm. in BM).—Fig. 25.
Rhizome short-creeping, 2144-3 mm 4g; scales
fawn-coloured, narrowly triangular, to c. 3 mm
long, to 18-seriate at base, with a very short
apical uniseriate portion. Leaves clustered;
petioles quadrangular with channelled sides,
stramineous to olivaceous brown, their lateral
faces with a series of median elongate pale pustu-
les, 10-25 cm long, 4%4—% as long as the lamina.
Lamina to c. 70 cm long, with 5-11 major lateral
pinnae to a side, a similar terminal one, and 1-5
strongly reduced basal ones. Primary rachis like
the petiole, sometimes also with pustules, the
adaxial groove with thick pale ridges. Pinnae
alternate, ascending, subsessile, linear, the major
ones 10-20 by 144-2 cm, broadest at base or in
the basal third, evenly narrowed to the subacute
to shortly acuminate apex. Secondary rachises
abaxially brownish in the lower, pale in the upper
part or throughout, abaxially bi-angular except
for a short basal portion. Pinnules 25-35 to a
side, alternate, subcontiguous to half their width
apart, subsessile, herbaceous, medium or dark
green when dry, slightly ascending, subtrapezoidal
to semi-ovate, the upper edge convex and without
an outer edge, or the upper edge straight and
with a distinct outer edge; larger pinnules 9-10
by 4-5 mm. Upper/outer margin with about 4
incisions to 2 mm deep, with very narrow si-
nus; sterile pinnules with alternating deeper and
shallower incisions; margin otherwise entire.
Upper pinnules strongly reduced, denticuliform.
Reduced basal pinnae with a few almost or quite
sterile pinnules. Veins immersed, evident, once or
twice forked, free. Sori interrupted by the incisi-
ons of the margin, usually binerval (uni- to quadri-
nerval); receptacle laterally exceeding the soral
veins. Indusium pale, entire or nearly so, with
straight or slightly convex base, often c. 1 mm
long, 0.3 mm wide, narrowed at the ends, not
reaching the edge by about its own width, little
reflexed at maturity. Spores pale brown, smooth,
trilete, c. 30 pw.
Distr. Malesia: Moluccas (Morotai, Ceram),
New Guinea (all Div.), Schouten I., Waigeu,
d’Entrecasteaux Is., Admiralty Is. (Manus),
Solomon Is.
Ecol. In forests, terrestrial and on logs, 30-
1000 m.
Note. The reduced basal pinnae of bipinnate
leaves are unique in the genus.
10. Lindsaea multisora v.A.v.R. Bull. Jard. Bot.
Btzg II, 16 (1914) 21.—Type: RACHMAT (exp.
VAN VUUREN) 669, Mt Lambolo, Celebes (BO;
dupl. in L).
L. kjellbergii C. Cur. Bot. Jahrb. 66 (1933)
53.—Type: KJELLBERG 3615, Malili, Celebes
(S-PA; dupl. in BM, BO).—Fig. 27.
Rhizome short-creeping, c. 2 mm g; scales
reddish brown, narrowly triangular, to 2 mm long,
to c. 8-seriate at the sometimes rather suddenly
broadened base, the apical 14 or 1% uniseriate.
Leaves close; petioles stramineous, sharply
quadrangular and shallowly sulcate, 10-30 cm
long, about as long as the lamina. Lamina oblong,
15-30 cm long, bipinnate (in juvenile yet partly
fertile plants simply pinnate), with 1-4 pinnae to
a side and a conform terminal one. Primary
rachis similar to the petiole. Pinnae 10-15 cm
long, 9-16 mm wide, obliquely ascending, sub-
opposite, sessile, widest in the lower half, gradual-
ly and strongly acuminate. Secondary rachises
abaxially sharply bi-angular almost to the base,
shallowly sulcate. Pinnules c. 25-40 to a side,
close, subcontiguous, subsessile, -- ascending,
especially the upper ones, herbaceous, dark green
or olivaceous when dry, approximately semi-ovate
in outline, the larger ones 7-8 by 2144-3 mm,
without a distinct outer margin, the lower margin
usually convex, the upper margin with mostly
3 major incisions reaching down to the middle or
a little less, the basal lobes often, the other ones
occasionally again incised; lobes, especially the
outer ones, often oblique, mostly 34-1 mm wide,
approximately parallel-sided, truncate or faintly
convex and slightly erose if fertile, narrowed-
rounded if sterile, the wing joining the lobes c.
1 mm wide. Upper pinnules gradually and strongly
reduced, confluent into the narrow pinna-apex.
Veins immersed, evident, 2 or 3 in the larger lobes,
single in the smaller ones. Sori usually bi- or
trinerval (uni- to quadrinerval); indusium thin,
pale, entire, 0.3-0.4 mm wide, not quite reaching
214
the margin, + reflexed at maturity. Spores pale
yellowish, trilete, smooth, c. 22 uw.
Distr. Malesia: Celebes (2 coll.).
Ecol. Terrestrial in rain-forest, 100 m.
11. Lindsaea natunae BAKER, Kew Bull. (1896)
40.—Type: Hose 315, Natuna Is. (K; dupl.?
in E, SAR).
L. canaliculatipes v.A.v.R. Bull. Jard. Bot.
Btzg III, 5 (1922) 211.—Lectotype: BUNNE-
MEVJER 5835, Riouw (BO; dupl. in K, L, P,
SING, U; fragm. in US).—Fig. 26.
Rhizome rather short-creeping, 2 mm @g; scales
reddish brown, narrowly triangular, acuminate,
to 24% mm long, to c. 10-seriate at base, the apical
uniseriate part relatively short. Leaves rather close;
petioles 8-35 cm long, about equaling the lamina,
stramineous to fawn-coloured, quadrangular,
with paler edges, -- sulcate. Lamina bipinnate
(rarely subtripinnate), with 3—S pinnae to a side
and a conform terminal one. Primary rachis
similar to the petiole, abaxially narrowly channelled.
Pinnae obliquely ascending, not close, subsessile,
10-15 cm long, 144-2% cm wide, narrowed in
the upper 13, the upper ones little shortened;
basal pinnae rarely with 1 or 2 basal pinnate
secondary pinnae. Secondary rachises abaxially
bi-angular. Pinnules c. 20-30 to a side, mostly
not contiguous, herbaceous, dark green when dry,
semi-ovate to subtrapeziform, subsessile, 9 by 3
to 12 by 44% mm, 214-3 times as long as wide,
the lower margin + straight, the upper straight
or slightly convex, a distinct outer margin not or
scarcely developed. Upper pinnules gradually
and strongly reduced, several denticuliform
ones connected with the narrow terminal segment.
Upper margin of pinnules mostly with 3 oblique
incisions which scarcely reach farther than 14
but get slightly deeper towards the apex of the
pinnule; outer margin without or with one
incision. Larger pinnule lobes with a straight or
sinuate outer margin, laterally somewhat convex;
sinus acute. Veins immersed, evident, mostly
once forked. Sori one per lobe, mostly binerval,
laterally exceeding the vein-ends; indusium entire,
y4—-l, mm wide, almost or quite reaching the
margin, scarcely reflexed at maturity. Spores
pale brown, trilete, smooth, c. 28 wu.
Distr. Malesia: Natuna Is. (3 coll.), Riouw
(2 coll.).
Ecol. Terrestrial in forests, 500-1000 m.
12. Lindsaea tetragona KRAMER, Blumea 15
(1968) 564.—L. tenuifolia auct. non BLUME, with
syn., of other authors, in part.—Type: BINNEN-
DIJK 160, Ambon (U).
Rhizome short-creeping, 14-2 mm @; scales
reddish or yellowish brown, narrowly triangular,
to 134 mm long, to c. 4-seriate, the uniseriate
apex short. Leaves close; petioles stramineous
or mottled with age, quadrangular and sulcate,
10-30 cm long, about as long as the lamina.
Lamina c. 10-40 cm long, oblong, bipinnate, with
3-9 pinnae to a side and a conform terminal
one; primary rachis like the petiole. Pinnae
FLORA MALESIANA
[ser. Il; volaste
sessile, obliquely ascending, not close, 8-15 cm
long, 17-25 mm wide, gradually to rather suddenly
narrowed at apex. Secondary rachises stramineous,
abaxially bi-angular, with a flat channel. Pinnules
c. 20-45 to a side, herbaceous, mostly pale green
when dry, close but usually not contiguous, some-
what ascending; lower margin often concave.
Upper/outer margin deeply incised, at least the
outer incisions reaching the middle of the pinnule,
very oblique. Lobes linear, parallel-sided, the
larger ones often forked. Sterile lobes narrowed-
rounded at apex, fertile ones often slightly
broadened at the sorus, truncate or slightly con-
vex. Upper pinnules gradually and strongly re-
duced, confluent with the pinnatifid, narrow pinna-
apex. Veins immersed, evident. Sori uni- or bi-
nerval. Spores very pale brown, trilete, smooth,
CHD:
1. var. tetragona.
Lobes of pinnules 13-34 (rarely to 1) mm wide;
pinnules incised to 24 or 34 (in sterile ones less),
semi-ovate to asymmetrically triangular; veins
rarely two per lobe; sori rarely binerval; indusium
pale, entire or sinuate, 44—'% (to almost 1) mm
long, 4% mm wide, not reaching the margin by
14-1, mm, strongly reflexed and concealed at
full maturity.
Distr. Malesia: Celebes, Philippines (Min-
danao), Moluccas (Talaud, Ceram, Ambon),
Louisiades; Solomon Is., Fiji, Samoa, Tahiti.
Ecol. Terrestrial in forests, often by water-
courses, to c. 600 m.
2. var. brassiana KRAMER, Blumea 15 (1968) 565.—
Type: Brass 27919, Sudest I., Louisiades (L;
dupl. in GH).
Pinnules incised to 13 or 14, only at the apex
to 4%, semi-ovate to subrhomboid; lobes 34-1 mm
wide, with -- erose outer edge, often binerval;
apices of pinnules often protracted into a caudi-
form, lobed segment; indusium erose, often not
reaching the edge of the pinnule by more than its
width.
Distr. Louisiades (2 coll.).
Ecol. In rain-forest, 300 m.
13. Lindsaea polyctena KRAMER. Blumea 15 (1968)
565.—Davallia blumeana of HooKer, Sp. Fil.
1 (1845) pl. 54 A.—Type: CuminG 309, Philip-
pines, Leyte (US; dupl. in B, BM, HBG, K, L,
MICH, P, W)—Fig. 28.
Rhizome short-creeping, c. 2 mm 9; scales light
reddish brown, elongate- to very narrowly
triangular, to c. 2 mm long, up to 8-seriate at
base, the uniseriate apex short. Leaves close;
petioles stramineous, quadrangular, at least
adaxially sulcate, 10-25 cm long, mostly somewhat
shorter than the lamina. Lamina bipinnate (rarely
subtripinnate), oblong, 10-30 cm long, with 3-11
pinnae to a side and a conform terminal one;
primary rachis like the petiole. Pinnae ascending,
sessile, 8-12 cm long, 14%—2 cm wide, the upper
ones hardly or not shortened. Secondary rachises
abaxially sharply bi-angular, the angles sometimes
April 1971] -_ LINDSAEA-GROUP (Kramer) 215
Fig. 24. Lindsaea adiantoides J. SmitH. Lower part of lamina, x 1% (EDANo 40514).—Fig. 25. L. kingi
CopeEL. Basal part of lamina, x 1% (BRaAss 25622).—Fig. 26. L. natunae BAKER. Pinna, nat. size (BUNNE-
MEIJER 5840).— Fig. 27. L. multisora v.A.v.R. Part of a pinna, x 3% (KJELLBERG 3515).—Fig. 28. L.
polyctena KRAMER. Part of a pinna, * 6 (CUMING 309).
216
FLORA MALESIANA
[ser. Il, vol
irregular and slightly wing-like, the face between
them scarcely concave. Pinnules c. 20-40 to a side,
mostly slightly ascending and with slightly
decurved upper part, not very close, curved-
narrowly triangular in outline, the largest 7-12
by 2-3 mm, thinly herbaceous, mostly dark green
when dry, even the lower margin at base only
very little sclerotic, deeply incised from the upper
margin, mostly with 6 primary lobes, these
capillary, 0.2-0.4 mm wide at base, twice as wide
at apex, connected by a wing of 0.2-0.4 mm,
towards the apex more oblique, in small pinnules
the basal one(s) forked, in large ones all except
a few apical ones forked, the basal one often
twice forked and then its basal branch sometimes
sterile. Segments slightly broadened from the base,
then suddenly broadened at the sorus, the outer
edge truncate-convex and often irregularly erose;
sterile segments subacute. Veins immersed, evi-
dent, single in the segments. Upper pinnules grad-
ually reduced, confluent into a pinnatifid pinna-
apex. Sori strictly uninerval; indusium pale, sub-
entire to erose, 44-4 mm long, c. 0.2 mm wide, not
reaching the apical margin by 1-2 times its width,
reaching the lateral margin or not, if longer with
concave’ base, free at the sides, + reflexed at
maturity. Spores pale brown, trilete, smooth.
cy 20).
Distr. Malesia: Philippines (Samar, Leyte,
Mindanao; 8 coll.). Map 4.
oot ie ee a. sneha
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Map 4. Distribution of Lindsaea polyctena
KRAMER (squares) and L. tenuifolia BL. (dots).
Ecol. Terrestrial in forest; one record 60 m,
one 800 m.
14. Lindsaea tenuifolia BLUME, En. Pl. Jav. (1828)
219; COPELAND, Philip. J. Sc. 78 (1949) 17; not
of COPELAND, Fern FI. Philip. 1 (1958) 110.—
Odontoloma tenuifolium (BLUME) J. SMITH,
Hook. J. Bot. 3 (1841) 415.—Davallia blumeana
Hooker, Sp. Fil. 1 (1845) 177 (not of pl. 54 A);
non Dayallia tenuifolia SWARTz.—Stenoloma blu-
meanum (HOOKER) FEE, Gen. Fil. (1852) 330,
pl. 27bis A f. 2.—Odontoloma blumeanum
(Hooker) Mett. Fil. Lips. (1856) 104.—Davallia
triquetra BAKER, Syn. Fil. ed. 1 (1867) 93.—L.
blumeana (HOOKER) KUHN in Mig. Ann. Mus.
Bot. Lugd.-Bat. 4 (1869) 277, nom. superfl._—
Odontoloma triquetrum (BAKER) J. SmitH, Hist.
Fil. (1875) 269 (‘triquetra’), nom. superfl.—L.
triquetra (BAKER) CHRIST, Farnkr. d. Erde (1897)
294, nom. superfl.—Type: BLUME s.n., Java (L).
Rhizome short-creeping, c. 2 mm g; scales
reddish brown, narrowly triangular, to 24% mm
long, to c. 10-seriate at base, with a short uni-
seriate apex. Leaves close; petioles stramineous,
triangular, especially the abaxial keel pronounced,
acute, (S5—)15-30 cm long, mostly somewhat
shorter than the lamina. Lamina oblong, mostly
olivaceous or dark green when dry, thinly herb-
aceous, 10-35 cm long, bipinnate (once pin-
nate in juvenile but nearly always sterile plants),
with 3-9 pinnae to a side and a conform terminal
one. Primary rachis similar to the petiole. Pinnae
obliquely ascending, mostly not close, sessile,
7 by 1 to 15 by 2 cm, acuminate; secondary
rachises abaxially sharply carinate virtually to
the base; upper pinnae little or not shortened.
Pinnules c. 20-45 to a side, mostly rather close,
often subcontiguous, spreading, slightly ascending,
or occasionally somewhat falcately decurved,
semi-ovate in outline, subsessile, 5-14 mm long,
24%-4 mm wide, 2-3% times as long as wide;
lower edge straight or -+- concave, upper edge
especially towards the apex convex, a distinct
outer edge scarcely developed. Pinnules deeply
incised from the upper margin into 4-6 major
segments, the larger ones nearly always forked, the
apical one often protracted and pinnatilobate;
ultimate lobes linear, parallel-sided to near the
slightly broadened apex if fertile, rounded or
subacute if sterile, the outer ones progressively
shorter and more oblique, rounded or erose at
apex, 144-34 mm wide, the wing joining them
%—(at the base of the pinnule)l mm wide. Upper
pinnules gradually and strongly reduced, confluent
into a narrow, pinnatifid pinna-apex. Veins
single or less often paired in the ultimate lobes,
immersed, evident. Sori uni- or less often binerval;
indusium pale, entire or subentire, 0.4-1 mm
long, 0.2 mm wide, if short with flat or convex,
if longer with concave base, reaching the margin
to falling short of it by almost its width. Spores
pale brown, trilete, smooth, c. 20 uw.
Distr. Malesia: Celebes, Moluccas (Ceram,
Halmahera. Ternate, Morotai), Waigeu, New
Guinea (West to East); a few collections from
S. Sumatra, Sipora (Mentawei Is.), West and East
Java. Map 4.
Ecol. Terrestrial and on rocks and tree bases,
in forest and by watercourses, to 1000 m, mostly
at lower altitude.
3. Section Synaphlebium
(J. SmirH) Diets in E. & P. Nat. Pfl. Fam. I, 4 (1902) 221.—Synaphlebium J. Smitty, Hook.
J. Bot. 3 (1841) 415, nom. nud.; in Hooker & Bauer, Gen. Fil. (1842) pl. 101.
April 1971]
LINDSAEA-GROUP (Kramer)
yi |
Type species: Synaphlebium recurvatum HOOKER = Lindsaea cultrata (WILLD.) SWARTZ.
Distr. Paleotropical, from India to Taiwan, the Marquesas, and NE. Australia.
Taxon. Lamina bipinnate or subtripinnate, with conform terminal pinna, or simply pinnate; pin-
nules dimidiate, reticulate-veined; spores trilete. The rhizome is short-creeping, but it may be more
long-creeping in exceptionally epiphytic specimens from moist montane habitats. Species with anastomo-
sing veins from other sections have in the past been incorrectly assigned to this section. A few species:
L. crispa BAKER, L. malayensis HOLTTUM, L. napaea v.A.v.R., usually have irregularly, i.e. incompletely
anastomosing veins, and the boundary of the section is not entirely sharp. It is particularly close to sect.
Temnolindsaea. The delimitation of the species meets with unusual difficulties, and the present treatment
should by no means be regarded as final.
15. Lindsaea malayensis Ho_ttrum, Gard. Bull.
S. S. 5 (1930) 69, f. 8; Rev. Fl. Mal. 2 (1954) 335,
f. 194.—L. subalpina auct. nonv.A.v.R.; HOLTTuM,
Gard. Bull. S. S. 5 (1930) 71.—Type: Md. HANIFF
4032, G. Kerbau, Perak (SING).
Rhizome short-creeping, 1-2 mm 9; scales me-
dium brown, very narrowly triangular, to 134 mm
long, to c. 5-seriate at base, with a long uniseriate
apex. Leaves clustered; petioles stramineous,
quadrangular, abaxially broadly and shallowly
sulcate or flat below, (6—)10—30 cm long, about as
long as the lamina, or in small leaves shorter.
Lamina simply pinnate or more often paucijugate-
bipinnate, with one odd lateral pinna to 2 pairs and
a conform terminal one; primary rachis like the
petiole, abaxially often more sulcate. Lamina if
simply pinnate c. 10-20 cm long and 3144-4 cm
wide, if bipinnate 18-30 cm long. Primary pinnae
(if any) not strongly ascending, not close, 8-20 cm
long, 2-4, often 24% cm wide, mostly rather ab-
Tuptly acuminate. Pinnule-bearing rachises like
the primary, or abaxially more sulcate. Pinnules
herbaceous to chartaceous, pale to dark olivaceous
when dry, rounded-subtrapeziform-ligulate or
less often rounded-subrectangular, spreading or
+ ascending, 12 by 4 to 20 by 6 mm, 2% to
almost 34% times as long as wide; lower margin
+ straight, upper margin often outward convex,
outer margin rounded into the upper or meeting
both lower and upper at approximately right
angles, the pinnules then scarcely narrowed to
the subtruncate apex. Upper margin with 3-5
Narrow incisions to 1 mm, occasionally to 2 mm
(43 of the width) deep, the lobes between them
flattish, the largest rarely shallowly incised again;
outer margin with 1 or without any incision;
fertile margin often in addition minutely erose;
sterile margin with the major incisions equally
developed, the lobes crenate. Upper pinnules
gradually or mostly rather abruptly reduced,
some denticuliform ones confiuent with the pin-
natifid terminal segment; /ower pinnules of simply
pinnate leaves often more remote. Veins immersed
or adaxially slightly prominulous, evident, once or
twice forked, 144-1 mm apart, connivent, here
and there anastomosing, or sometimes quite free,
but most leaves at least with a few anastomoses,
the row of areoles rarely complete in one pinnule.
Sori interrupted by the incisions, 14 mm long,
the one on the outer margin usually continuous
with the outermost one of the upper; indusium
greenish or brownish, subentire, 0.2-0.3 mm
wide, not reaching the margin by an equal or
larger distance, reflexed and often quite concealed
at maturity. Spores very pale brown, trilete,
smooth, c. 22-25 w.
Distr. S. Thailand, in Malesia: Malay Penin-
sula (Pahang, Perak). Incorrectly reported from
Borneo (Act. Phytotax. Geobot. 22, 1966, 90).
Ecol. In mountain forests, c. 1200-2000 m.
16. Lindsaea napaea v.A.v.R. Bull. Jard. Bot-
Btzg II, 20 (1915) 19, pl. 3; Hottrum, Gard. Bull-
S. S. 5 (1930) 66; Rev. FI. Mal. 2 (1954) 330, f.
189.—Type: TEIJSMANN 16616, Mt Dai, Lingga
Is. (BO).
Rhizome short-creeping, 115-2 mm g; scales
reddish brown, narrowly triangular or linear,
to slightly over 1 mm long, to 6-seriate at base
but often narrower, with a short uniseriate apex.
Leaves close; petioles stramineous or darker with
age, abaxially flat or convex and laterally bi-
angular, with obtuse, downward gradually evanes-
cing ridges, c. 10-30 cm long, about as long as the
lamina. Lamina broadly ovate, c. 10-25 cm
long, bipinnate (rarely simply pinnate and fertile,
or subtripinnate), with 1-4 pinnae to a side and a
conform terminal one; primary rachis abaxially
slightly concave or upward channelled, bi-angular.
Pinnae sessile, ascending, not close, 10-22 cm
long, 134-24 cm wide, gradually narrowed from
the lower third or the middle but more strongly
so near the apex; upper pinnae slightly or not
shortened. Secondary rachises abaxially bi-angular,
between the angles flat or convex. Pinnules c.
20-45 to a side, herbaceous, olivaceous or brown-
ish when dry, spreading or a little ascending or
slightly decurved, often subcontiguous, evenly
spaced, 10-12 by 344 mam, ligular or trapezi-
form, truncate-rounded or narrowed-rounded
at apex; upper margin very faintly sinuate or
more often with 3 or 4 oblique incisions, the inner
very shallow, the outer a little deeper but not
reaching to 14 of the width; outer margin mostly
not incised; sterile pinnules crenate-sinuate on
the upper and outer margin. Upper pinnules
strongly reduced, some denticuliform ones con-
fluent with the caudate pinna-apex. Veins im-
mersed, not evident, mostly twice forked, 2-74
mm apart, very oblique, free or less often conniv-
ent or more freely anastomosing. Sori rarely
continuous, mostly interrupted by the incisions,
the sorus on the outer margin mostly continuous
with the outermost one of the upper margin.
Sori of completely fertile pinnules straight or
the outer ones slightly convex, mostly on 2-6
218
FLORA MALESIANA
[ser. Il; yoke
veins. Indusium entire, 1/,—1/; mm wide, not
teaching the margin by less than its width, re-
flexed and + concealed at maturity. Spores pale
brown, trilete, smooth, c. 17-19 yp.
Distr. S. Peninsular Thailand, in Malesia:
Malay Peninsula (Perak, Pahang, Penang, Selan-
gor, Kelantan, Negri Sembilan, Kedah, Malacca),
Lingga Is., W. Sumatra, Siberut.
Ecol. Terrestrial and epilithic, in mountain
forests, c. 1000-1300 m, apparently of local
occurrence.
Note. The three collections from Sumatra and
Siberut have more regularly anastomosing veins
than the others. but are otherwise not aberrant.
17. Lindsaea subalpina v.A.v.R. Bull. Jard. Bot.
Btzg II, 23 (1916) 15; not of Hottrum, Gard.
Bull. S. S. 5 (1930) 71.—Type: AsorB 388, Rimbo
Pengadang, Bencoolen, Sumatra (BO; dupl. in
L).—_Fig. 33.
Rhizome short-creeping, c. 2 mm g; scales
(few seen) reddish brown, narrowly triangular,
2mm long, to 4-seriate at base, with a very short
uniseriate apex. Leaves close; petioles stramineous
to pale reddish brown, stout, quadrangular with
shallowly sulcate sides, c. 15-30 cm long, about
as long as the lamina. Lamina simply pinnate,
linear, c. 15-40 cm long, 44%4-5% cm wide; rachis
similar to the petiole. Pinnules c. 12—30 to a side,
spreading or slightly ascending, firmly herbaceous,
mostly dark green when dry, usually not con-
tiguous, elongate-parallelogram-shaped to ligular,
2 by 0.6 to 3% by 1 cm, 3-31 times as long as
wide, subsessile, little narrowed to the apex,
this rounded or occasionally caudate-protracted;
upper margin with 5—7 major incisions, these
rather oblique, reaching 44-1 down, narrow,
the largest lobes sometimes more shallowly in-
cised again; lobes slightly convex, often erose.
Sterile pinnules not seen. Relatively few upper
pinnules reduced, mostly one of c. 34 cm adnate
to the narrow, lanceolate, lobed terminal segment.
Veins immersed but evident, especially above,
very oblique, regularly anastomosing, often with
2 series of areoles between the two margins, al-
most 1 mm apart at the vein-bases. Sori single
in the lobes, usually on 4-8 veins, the one on the
outer margin often continuous with the outermost
of the upper; indusium pale, erose, /,—1/4, mm wide,
not reaching the margin by about the same dis-
tance or less, reflexed and + concealed at maturity.
Spores pale brown, trilete, smooth, c. 19-22 w.
Distr. Malesia: Sumatra (Bencoolen, 5 coll.,
West Java (1 coll.).
Ecol. Terrestrial in forest, 600-1000 m.
18. Lindsaea obtusa J. SmitH in Hooker, Sp. Fil.
1 (1846) 224.—Type: CumING 394, Malacca
(K; dup]. in B, E,-GH, P. W).
? L. tripartita BLUME, En. Pl. Jav. (1828) 219.—
Type: BLUME s.n., Java (L).
? L. ambigens CEsatTi, Rendic. R. Accad. Sci.
Fis. Mat. Napoli 16 (1877) 25, nom. subnud.—
Type: Beccari s.n., Andai, Papua (n.yv.).
L. schultzei BRAUSE, Bot. Jahrb. 49 (1912) 29.—
Type: SCHULTZE 3049, Sepik R., Terr. of New
Guinea (B).
L. decomposita WiLLp. f. minor v.A.v.R. Bull.
Jard. Bot. Btzg II, 7 (1912) 21.—Type: KiNG 48
‘p.p., Papua (BO).
L. ceramica v.A.v.R. Bull. Jard. Bot. Btzg II,
28 (1918) 32.—Type: Kornassi 707, Ceram (BO).
? L. sinuato-crenata v.A.v.R. Nova Guinea 14
(1924) 30; CopELAND, Philip. J. Sc. 78 (1949)
20.—Type: H. J. LAM 1888, mountain ridge near
Doormantop, W. New Guinea (L).
L. decomposita WILLD. f. longipinnula v.A.y.R.
Nova Guinea 14 (1924) 30.—Type: H. J. Lam
1163, near Prauwenbivak, Mamberamo R., W.
New Guinea (BO; dupl. in L).
L. furcata COPELAND, Un. Cal. Publ. Bot. 18
(1942) 218; Philip. J. Sc. 78 (1949) 21, pl. 4.—
Type: Brass 13229, Bernhard Camp, Idenburg
R., W. New Guinea (MICH; dupl. in GH, L).
L. decomposita auct. non WiLLD.; COPELAND,
Philip. J. Sc. 78 (1949) 20.
L. davallioides auct. non BLUME; HOLTTUM,
Gard. Bull. S. S. 5 (1930) 69, f. 7; Rev. Fl. Mal. 2
(1954) 332, f. 190.—Fig. 31.
Rhizome short-creeping, 1-24% mm, usually
14% mm @; scales medium brown, narrowly
triangular, to 1144 mm long, to 4-seriate at base,
with a rather short uniseriate apex. Leaves close
(less so in epiphytic plants); petioles stramineous
or in mature specimens mostly medium to dark
brown or blackish, sometimes pale-margined or
mottled, -- sharply quadrangular and mostly
somewhat sulcate, c. 10-30 cm long, % as long
as to equaling the lamina. Lamina simply pinnate
or in full-grown plants mostly bipinnate, with 1
or 2, less often with 3 pairs of pinnae (very
rarely the basal pinnae forked) and a conform
terminal one; primary rachis like the petiole.
Pinnae mostly subopposite, obliquely ascending,
10-20 cm long, 144-3 cm wide (simply pinnate
laminas may be wider), widest in the lower half,
gradually and strongly narrowed to the usually
long-acuminate apex; secondary rachises abaxially
bi-angular, mostly distinctly sulcate, usually
pale. Pinnules herbaceous to chartaceous, mostly
rather dark when dry, c. 20-35 to a side, mostly
distinctly ascending, close but hardly contigu-
ous, the basal ones of simply pinnate leaves often
more remote, ligulate to subtrapeziform, the lar-
ger ones 10-16 mm long and 5—7 mm wide, nearly
always slightly over twice as long as wide, nar-
rowed from the base to the broadly rounded
or subtruncate apex, less often of almost equal
width from base to apex, the upper margin
straight at the base, outward convex. Upper
pinnules gradually and strongly reduced, mostly
some denticuliform ones present below the cre-
nate-pinnatilobate pinna-apex. Upper and outer
margins of pinnules incised, mostly with 3-5
narrow, oblique incisions usually less than 1 mm
(but in extreme cases to 2 mm) deep; margin
otherwise minutely but distinctly crispate and/or
erose. Lobes of fertile pinnules, especially the
inner ones, flat, truncate, not convex. Veins
immersed and often obscure, mostly twice
April 1971]
LINDSAEA-GROUP (Kramer)
219
forked, regularly anastomosing, forming one or
sometimes towards the apex of the pinnule two
series of areoles; larger areoles to | mm wide.
Sori interrupted by the incisions, variable in size,
the innermost one often quadrinerval, the outer
ones bi- or trinerval, but often on more or fewer
vein-ends. Indusium subentire or, if broader,
often erose, very veriable in width, 0.2-0.7 mm
wide, not reaching the margin by 1-1 times its
width, bulging but hardly reflexed at maturity.
Spores pale yellowish, trilete, smooth, c. 24 wu.
Distr. Taiwan; throughout Malesia (but
absent from the Lesser Sunda Is. except Bali),
east to the Solomon Is. and the Bismarck Archi-
pelago; Micronesia, Queensland.
Ecol. In primary forest, often on banks, on
rocks, by or on tree bases, from sea-level up
to 2000 m, mostly below 1500 m.
Notes. As defined here this is one of the most
common and widespread species of SE. Asia.
It is exceedingly variable, and the extremes look
quite distinct. In E. Malesia and W. Melanesia
a form occurs with indusia 0.2-0.35 mm wide,
less than half their width removed from the edge,
pinnule lobes not rarely concave and then some-
times laterally subcorniculate, thinner texture,
and truncate, subrectangular pinnules; it may
represent a distinct infraspecific taxon. It occurs,
however, together with the more widespread form,
with large numbers of intermediates, and classi-
fication on the basis of herbarium material alone
seems at present impossible.
The type of L. tripartita BLUME is largely sterile,
and it is impossible to ascertain which species it
really represents. The type of L. sinuato-crenata
v.A.v.R. is a diseased or for another reason
malformed !eaf; it probably belongs to the present
species. The type of L. schultzei BRAUSE is a
specimen with extremely narrow pinnules (4 by
18 mm); it may represent a distinct taxon.
19. Lindsaea longifolia COPELAND, Philip. J. Sc.
38 (1929) 145, pl. 3; Fern Fl. Philip. 1 (1958)
109.—Type: ReEILLo B. Sc. 16227, Basilan, Phi-
lippines (MICH; dupl. in B, P).—Fig. 30.
Rhizome short-creeping, 2 mm g; scales rather
dark brown, narrowly triangular, to 144 mm long,
to 8-seriate at base, with a short uniseriate apex.
Leaves crowded; petioles c. 10-40 cm long,
4-2 times as long as the lamina, stramineous
or reddish brown or darker with age, at least
in the upper part obtusely to sharply bi-angular,
often also sulcate. Lamina bipinnate (rarely
simply pinnate and fertile, or subtripinnate),
12-35 cm long, with 1-5 pinnae to a side and a
conform but often longer terminal one; primary
rachis abaxially bi-angular, sulcate. Pinnae
strongly ascending, 2144 cm distant, linear, 10-25
by 0.7-2 cm, long-acuminate; secondary rachises
abaxially bi-angular to shallowly sulcate. Pinnules
c. 30-40 to a side, herbaceous, mostly dark green
when dry, close, subcontiguous to overlapping,
ascending or the lower ones spreading, 7-10 by
314-5 mm (rarely smaller, 5 by 242 mm), 1 2-2
times as long as wide, subovate to dimidiate-
subligulate, very obtuse, little narrowed to the
rounded apex; upper/outer margin with 1-3
incisions, these in large pinnules 4%4—'4 mm deep,
in small ones sometimes deeper; margin otherwise
entire; sterile margin shallowly crenate. Upper pin-
nules + entire, gradually and very strongly re-
duced, several denticuliform ones confluent with
the small, narrow pinna-apex. Veins immersed,
not evident, mostly twice forked, forming a
(sometimes incomplete) series of areoles 1-24
mm wide. Sori interrupted by the incisions,
mostly tri- or quadrinerval, longer in upper,
entire pinnules. Indusium pale, subentire, 0.3
mm wide, not reaching the margin by 0.2 mm,
little reflexed at maturity. Spores pale yellowish,
trilete, smooth, c. 25 yu.
Distr. Malesia: Philippines
danao, Polillo, Samar, Leyte,
Panay, Biliran, Basilan).
Ecol. Terrestrial, in mountain forests (always?).
(Luzon, Min-
Catanduanes,
20. Lindsaea ramosii COPELAND, Philip. J. Sc. 38
(1929) 144, pl. 2; Fern Fl. Philip. 1 (1958) 110.—
Type: Ramos B. Sc. 7652, Cagayan Prov.,
Luzon (MICH).
Very similar to L. longifolia; pinnules small,
7 by 4 to 5 by 3 mm, more rounded, not incised.
Distr. Malesia: Philippines (Luzon, 4 coll.).
Note. Very doubtfully distinct from L. longi-
folia.
21. Lindsaea lobata PorreT in Lamk, Encycl.
Suppl. 3 (1813) 448.—Adiantum lobatum (PotRET)
PorRET ex STEUDEL, Nomenkl. 2 (1824) 275,
nom. invalid. in syn.; non PRESL (1825), nec KUNZE
ex KuHN (1881).—Davallia lobata (POIRET)
DesvAux, Prod. (1827) 315.—Schizoloma loba-
tum (PotrET) BepD. Ferns S. Ind. & Br. Ind.
Suppl. (1876) 6, quoad typum.—Type: COMMERSON
s.n., Java (P).
L. davallioides BLUME, En. PI. Jay. (1828) 218;
Hooker, Sp. Fil. 1 (1846) 224, pl. 68 A; DIELs
in E. & P. Nat. Pfl. Fam. I, 4 (1902) 221, f. 119
G; BACKER & PostHumMus, Varenfl. Java (1939)
115, f. 22; not of Hottrum, Rev. Fl. Mal. 2
(1954) 332.—Synaphlebium davallioides (BLUME)
J. Smitu, Lond. J. Bot. 1 (1842) 424.—Davallia
kunzeana Hooker, Sp. Fil. 1 (1845) 177.—Schi-
zoloma davallioides (BLUME) Moore, Ind. Fil.
(1857) 35.—L. decomposita WILLD. var. davyalli-
oides (BLUME) Doin, Bibl. Bot. 20 (1915) 84.—
Type: BLUME s.n., Java (L; dupl. in P).
L. lobata PotretT var. incisa METT. ex KUHN in
Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 277.—
Lectotype: ZOLLINGER 1087, Java (L; dupl. in
B, HBG) (the other coll. cited, ZOLLINGER II
381, is L. obtusa).
Rhizome short-creeping, 1144-2 mm 9g; scales
dark reddish brown, narrowly triangular, to
2 mm long, to c. 10-seriate at base, with a short
uniseriate apex. Leaves clustered; petioles stram-
ineous to pale brown, quadrangular, occasion-
ally with paler angles, abaxially flat to shallowly
sulcate, at the base often dark-verruculose, 8-45
cm long, equaling to over twice as long as the
220
FLORA MALESIANA
lamina. Lamina bipinnate (very rarely once
pinnate and fertile), with 1—4, most often 2
or 3 pinnae to a side and a conform terminal
one; primary rachis like the petiole, often
more distinctly pale-margined, abaxially usually
sulcate. Pinnae ascending, subsessile, linear,
their width apart to contiguous, 10-20 cm long,
1.7-2.5 cm wide, widest in the lower third or the
middle, gradually and strongly tapering to the
often long-acuminate apex; terminal pinna often
the longest and widest; secondary rachises abaxi-
ally broadly and + shallowly sulcate, sometimes
pale-margined. Pinnules c. 25-40 to a side, half
their width apart to contiguous, spreading or
mostly ascending, herbaceous, medium to dark
or blackish green when dry, translucent, 14-
elliptic, narrowed and obtuse to subacute, 7-15
mm long, 2!4—-5 (often 4) mm wide, 214-3 times
as long as wide, almost evenly narrowed through-
out, the upper margin outward increasingly
convex, with 3 or 4 major incisions, the inner
ones reaching to % or '%, or rarely to %, the
outer ones to ¥% or slightly beyond, with acute
sinus; at least the broader lobes with convex
outer margin, the lateral margins almost parallel,
the major ones often shallowly incised again,
otherwise entire, a distinct outer margin not
developed. Upper pinnules strongly reduced, some
denticuliform ones confluent with the narrow,
+ caudiform terminal segment. Veins immersed,
evident, mostly twice forked, regularly anasto-
mosing, with one series of areoles, one under each
incision, c. %4 mm wide, the lobes containing
2-5 vein-ends. Sori joining all vein-ends of a lobe;
receptacle at least in larger lobes distinctly
convex; indusium pale, entire, 0.3-0.4 mm wide,
reaching the margin or very nearly so, + reflexed
at maturity; sporangia often spreading beyond
the pinnule margin. Spores rather pale brown,
trilete, smooth or almost so, c. 21-23 yu.
Distr. Malesia: Sumatra, West and Central
Java, Lesser Sunda Ils. (Bali, Sumba, Flores),
Borneo, Celebes, Philippines (rare and local),
New Guinea (rare), Admiralty Is.; a few doubtful
collections from the Malay Peninsula; Caroline
Is. Distinct varieties in Hainan and Indo-China.
Records from elsewhere are due to confusion with
related species.
Ecol. Terrestrial or on tree bases in primary
forest, 50-1800, mostly 600-1500 m.
Note. In West Java, where this species is
common, it is very homogeneous. Specimens
from elsewhere, where it is much rarer, are often
somewhat aberrant. There may be some hybridiz-
ation with related species.
22. Lindsaea parallelogramma y.A.v.R. Bull.
Jard. Bot. Btzg III, 5 (1922) 212; HoLtrum,
Gard. Bull. S. S. 5-(1930) 70, f. 9; Rev. Fl. Mal.
2 (1954) 335, f. 193.—Type: BUNNEMEIJER 7359,
Mt Tjikalu, P. Singkep, Lingga Is. (BO; dupl.
in L).
L. dayallioides BLUME f. parallelogrammoides
v.A.v.R. Nova Guinea 14 (1929) 31.—Type:
H. J. LAM 859, near Prauwenbivak, W. New
[ser. II, vol. 1%
Guinea (BO; dupl. in L).—Fig. 29.
Rhizome short-creeping, 114,-2 mm @; scales
fawn-coloured, narrowly triangular, to 2 mm
long, up to c. 10-seriate at base, with a rather
short uniseriate apex. Leaves close; petioles
characteristically olivaceous-brown, dull, sharply
quadrangular, adaxially channelled and pale-
margined, c. 15—30 cm long, 114-2 times as long
as the lamina. Lamina bipinnate, broadly oblong,
sometimes wider than long (rarely simply pinnate
and fertile, then linear), c. 10-20 cm long, with
1-4 pairs of lateral pinnae and a conform terminal
one; primary rachis similar to the petiole,
abaxially channelled. Pinnae patent or ascending,
often contiguous, sessile, c. 8-18 cm long, 2-3%
cm wide, the upper ones little or not shortened;
secondary rachises similar to the primary,
abaxially sulcate and pale-edged. Pinnules c.
15-30 to a side, mostly somewhat ascending,
scarcely to. slightly contiguous, herbaceous,
mostly drying dark brownish green, not translu-
cent, parallelogram-shaped, the larger 9-14 mm
long, 3-5 mm wide, 21% to nearly 3 times as long
as wide (except when the apex is caudiform),
very little or not narrowed to the obliquely
truncate apex. Upper and outer margin obliquely
incised, the upper usually with 4, the outer with
1 incision, those of the upper margin reaching
Y4-1,(-¥,), the apical ones often %4(—24); lobes
parallel-sided, convex, entire or minutely erose,
the basal, broadest often with a shallow incision;
sinus narrow, acute. Some pinnules occasionally
caudate-protracted at the apex. Upper pinnules
rather suddenly strongly reduced, several denti-
culiform ones confluent with the caudate, linear
pinna-apex. Veins immersed or slightly raised,
mostly evident, regularly anastomosing, forming
a series of areoles 145 mm wide below the level
of the incisions, rarely the larger lobes with
another series. Sori one per lobe, up to c. 2 mm
long, bi- to quadri-, mostly trinerval; indusium
1/.1/, mm wide, entire, not reaching the margin
by less than its width to about twice its width,
reflexed at maturity. Spores very pale brown,
trilete, smooth, c. 17-20 w.
Distr. Malesia: Peninsular Thailand, Malay
Peninsula, Singapore, Lingga Is., Banka, Sumatra,
Java (very rare, no recent collections), Borneo
(apparently quite common), New Guinea (a
few collections from W. New Guinea and the
Territory of New Guinea). Map 5.
0
oe eos
o 8 o>
Map 5. Distribution of Lindsaea parallelogramma
v.A.v.R. Several localities, mainly in Borneo,
could not be located.
—
221
LINDSAEA-GROUP (Kramer)
April 1971]
OOKER. Part of
x 228 Wear,
* 2 (BROOKE 9482).—Fig. 30. L.
* 11% (BAKHUIZEN VAN DEN BRINK 3360).—
Fig. 31. L. obtusa J. SMITH in H
L. integra Hoitt. A. Pinnule,
(AET & IDJAN 297).
\,, pinnule,
Fig. 34. L. modesta KRAMER. Leaf, nat. size
Fig. 29. Lindsaea parallelogramma v.A.v.R. Leaf,
longifolia CoPEL. Part of a pinna, x 2% (WENZEL 585).—
a pinna, X 2 (BAKHUIZEN VAN DEN BRINK 6630). — Fig. 32.
«x V, (Wray 3674). — Fig. 33. L. subalpina v.A.v.R. Pinnule,
222
FLORA MALESIANA
[ser. II, vol. 13
Ecol. Terrestrial in forests, on clay soil and
rocks, from sea-level up to c. 1200 m.
23. Lindsaea cultrata (WILLD.) Swartz, Syn:
Fil. (1806) 119; KRAMER, Blumea 15 (1968)
565; not of other authors.—Adiantum cultratum
WILLD. Phytogr. (1794) 14, pl. 10 f. 2.—Type:
coll.?, “Malabaria’ (B, herb. WILLDENow).
L. decomposita WILLD. Sp. Pl. 5 (1810) 425;
Hoitrum, Gard. Bull. S. S. 5 (1930) 66, f. 5;
Rev. Fl. Mal. 2 (1954) 333, f. 192; CopELAND,
Fern Fl. Philip. 1 (1958) 111, p.p.—Type: coll.?,
‘India’ (B, herb. WILLDENOW).
L. nitens BLUME, En. Pl. Jav. (1828) 217.—Schi-
zoloma nitens (BLUME) BEDb. Ferns S. Ind. ed.
2, corr. (1873).—Synaphlebium nitens (BLUME)
J. SmitH, Hist. Fil. (1875) 268.—Type: BLUME
S.n., Java (L).
Synaphlebium recurvatum Hooxer in Hooker
& Bauer, Gen. Fil. (1842) pl. 101; J. Smitn,
Hist. Fil. (1875) 268, pl. 18 c.—L. recurvata
(HOOKER) Hooker, Sp. Fil. 1 (1846) 220, pl.
70 A; HoLttum, Gard. Bull. S. S. 5 (1930) 66.—
Schizoloma recurvatum (HOOKER) Moore, Ind.
Fil. (1857) 35.—Type: a plant without data,
prob. coll. by WALLICH, must be the type (K).
L. intermedia Hooker, Sp. Fil. 1 (1846) 222,
pl. 67 B.—Type: CuminG 404 [err. (?) cited as
*464’] ‘Philippines’, more probably Malay Pen-
insula (K; dupl. in BM).
L. intermedia HOoKer var. minor HOOKER,
Sp. Fil. 1 (1846) 222, nom. subnud.—Type:
CUMING 392, Malacca (K; dupl. in B, BM, E,
GH, L, SING, W).
L. sarasinorum Cuyrist, Verh. Naturf. Ges.
Basel 11 (1897) 429; Ann. Jard. Bot. Btzg 15
(1897) 101, pl. 14 f. 13 (poor). —Type: SARASIN
s.n., Ussu, Celebes (P).
L. nitida COPELAND, Philip. J. Sc. 6 (1911)
Bot. 138, pl. 21; not of HoLtrum, Rev. FI. Mal. 2
(1954) 333. — Type: Brooxs 12, Mt Penrissen,
Sarawak (MICH).
L. trapezoidea COPELAND, Un. Cal. Publ. Bot.
14 (1929) 376, pl. 61.—Type: BARTLETT 7029,
near Aek Kanopan, Kualu, Sumatra (UC, n.y.;
dupl. in MICH, US).
Rhizome short-creeping, 14%-2 mm g; scales
reddish brown, narrowly triangular, to 14% mm
long, to 8-seriate at base, with a short uniseriate
apex. Leaves close; petioles stramineous, abaxially
at least in the upper part obtusely or usually
acutely bi-angular and often sulcate, 10-40 cm
long, %-1'% times as long as the lamina. Lamina
simply pinnate or in full-grown plants (probably
always) bipinnate, with 1-2 pinnae to a side and
a conform but sometimes larger terminal one,
10-30 cm long; primary rachis of bipinnate leaves
like the upper part of the petiole. Pinnae obliquely
patent, narrowly oblong, like simply pinnate
laminas, 10-20 cm long, 2-4 cm wide, acute or
shortly, rarely more long-acuminate; pinnule-
bearing rachises abaxially sharply bi-angular or
shallowly sulcate, the edges sometimes slightly
wing-like. Pinnules c. 12-30 to a side, less than
their width apart to contiguous, spreading to
somewhat ascending or, especially in smaller,
simply pinnate laminas the basal ones decurved,
herbaceous, medium to dark green when dry,
trapeziform to rhombic, or in larger leaves often
ligulate or truncate-ligulate, not very rarely
with a slightly protracted apex, the larger ones
12 by 3 to 19 by 7 mm, almost 2% to 3 times as
long as wide, usually little narrowed; upper
margin straight or convex outward, a separate
outer margin present, joining the upper at right
angles or more often at an acute angle, less often
rounded into the upper margin; upper margin of
larger fertile pinnules with 1-3 narrow oblique
incisions to | mm deep, the outer margin occa-
sionally with one incision; smaller pinnules often
entire; sterile margin shallowly crenate; fertile
margin not erose; lobes flat. A few upper pinnules
strongly reduced, or less often many, | or 2 or
a few connected with the narrow, acute, asym-
metrically lanceolate terminal segment. Veins
immersed, evident or not, regularly anastomosing,
forming one (very rarely parts of a second)
series of areoles, the larger ones 34-1 Y, mm wide.
Sori interrupted by the incisions, the sorus on
the outer margin mostly continuous with the
outermost one of the upper. Indusium pale or
greenish, entire or almost so, 0.2—0.3 mm wide,
not reaching the margin by an equal distance
or usually less, little reflexed and scarcely concealed
at maturity. Spores pale brown, trilete, smooth,
C20) E:.
Distr. S. India?, Ceylon, Botel Tobago, S.
Thailand; in Malesia: Malay Peninsula, Singa-
pore, Riouw Is., S. China Sea Is., Sumatra,
Mentawei Is., Banka, West Java, Borneo, Celebes
(a few collections), Philippines (Palawan,
Sibuyan). In the Solomon Is. an aberrant, perhaps
distinct form. Map 6.
Cc, @ \
Len, ©
‘ \ Oe “Se, . 5
N : we. (se
e@\ ° ae :
eo ©
8 een
L 0
Map 6. Distribution of Lindsaea_ cultrata
(WILLD.) Sw. in Malesia; further known from
Ceylon, S. India, and the Solomons (a distinct?
form).
Ecol. Terrestrial in forests, up to c. 1300 m;
often locally frequent.
Note. The name L. decomposita, under which
this species was known for a long time, has been
greatly misused for various species of sect. Syna-
April 1971]
LINDSAEA-GROUP (Kramer)
225
phlebium. The combination of little reduced
upper pinnules, sori interrupted by shallow in-
cisions of the margin, and the sorus on the outer
margin continuous with the outermost one on
the upper, is distinctive, but it is not rare to
find plants which otherwise agree with L. cul-
trata but in which one or the other of these
characters is missing.
24. Lindsaea papuana COPELAND, Philip. J. Sc.
7 (1912) Bot. 68; ibid. 78 (1949) 21.—Type:
KING 358, Lakekamu, Papua (MICH).
Rhizome short-creeping, c. 14, mm @g; scales
very narrowly triangular to acicular, to 3 mm
long, to 7-seriate at base but often the apical half
uniseriate, the basal half biseriate almost to base.
Leaves close; petioles stramineous, abaxially
below obtusely, upward acutely bi-angular and
sulcate, c. 12-30 cm long, 1-114 times as long as
the lamina. Lamina 15-22 cm long, simply
pinnate or with one pair of pinnae to 15 cm long
and a conform terminal one; rachises like the
upper part of the petiole. Pinnules c. 20-30 to
a side, dark olivaceous when dry, herbaceous,
spreading, the upper ones subcontiguous, the
lower ones their width apart; larger pinnules
asymmetrically elongate-triangular to subligulate,
18 by 6 to 30 by 9 mm, 3-314 times as long as
wide, not much narrowed to the narrowed-
rounded apex, a distinct outer margin not devel-
oped; both margins almost straight at base and
convex outward. Upper pinnules gradually and
strongly reduced, 1 or 2 of a few mm long con-
nected with the narrowly lanceolate acuminate
terminal segment. Margins entire, or the upper
remotely and shallowly crenate; sterile margin not
seen. Veins immersed, + evident, regularly
anastomosing, with | or not rarely 2 rows of
areoles of fluctuating width. Sori continuous, or
interrupted by the crenations of the upper margin
(probably quite continuous in fully fertile pin-
nules). Indusium entire, greenish, 44 mm wide or
less, not reaching the margin by its width or
more, scarcely reflexed at maturity. Spores
light brown, trilete, smooth, c. 23 w.
Distr. Only known from the type collection.
Ecol. No data.
Note. In habit this species rather closely
resembles certain forms of the neotropical L.
arcuata KUNZE.
25. Lindsaea integra HoL_trum, Gard. Bull. S. S.
5 (1930) 67, f. 6.—L. nitida auct. non COPELAND;
Ho.Lttum, Rev. FI. Mal. 2 (1954) 333, f. 191.—
Type: HoL_ttum 20934, Tahan R., Pahang (SING;
dup!. in BM, BO, K, US).—Fig. 32.
Rhizome short-creeping, 1144-2 mm g; scales
medium brown, narrowly to very narrowly
triangular, to 1144 mm long, to 6-seriate at base,
with a long uniseriate apex. Leaves clustered;
petioles sharply quadrangular almost throughout,
reddish, abaxially in the upper part mostly pale-
and somewhat wing-angled, rarely stramineous,
2-20 cm long, much shorter than to about
equaling the lamina. Lamina linear, 8-22 cm
long, 1-24% cm wide (wider if bipinnate), simply
pinnate, or rarely with one pair of pinnae;
rachis similar to the upper part of the petiole,
abaxially nearly always distinctly wing-angled.
Pinnules c. 6-30 to a side, ascending or the lower
Ones spreading, their width apart to contiguous,
chartaceous, mostly olivaceous when dry, sub-
trapezoidal to semi-ovate, the larger ones 7-12
(-15) by 3%4-6(-7) mm, twice as long as wide,
rounded or subtruncate at apex, narrowed from
the base to the rounded apex, or, especially if
subtruncate, little narrowed, the outer margin
often meeting the upper at an angle of less
than 90°. Lower pinnules sometimes slightly
remote; a few (more in larger leaves) upper
pinnules rather suddenly but not strongly re-
duced, rarely less than half as long as the larger
ones, the terminal segment narrowly rhombic
to lanceolate, c. %-1 cm long, free or almost so.
Veins immersed, little evident, not very regularly
anastomosing, with one (very rarely two) series
of areoles, the outer veins often (in small
pinnules sometimes all veins) free; areoles
)—34, mm wide. Sori continuous, uniting all vein-
ends; indusium subentire, brownish, 0.3-0.4 mm
wide, almost or quite reaching the margin, re-
flexed at maturity. Spores pale yellowish, trilete
smooth, c. 22 mu.
Distr. S. Peninsular Thailand; Malesia: Malay
Peninsula (Pahang, Perak, Malacca, Selangor), Su-
matra (1 coll.), Borneo (Sabah, Sarawak, Brunei).
Ecol. Terrestrial in forest, often (always?)
by streams on rocks; from c. 100-1200 m; few
ecological data. Reported to be a rheophyte by
ASHTON, Sarawak.
26. Lindsaea azurea CHRIST, Verh. Naturf. Ges.
Basel 11 (1897) 429; Ann. Jard. Bot. Btzg 15
(1897) 101, pl. 14 f. 12; CopELAND, Philip. J. Sc.
78 (1949) 21.—Type: SARASIN s.n., Ussu, Ce-
lebes (P).
L. azurea CHRIST var. mambae v.A.v.R. Bull.
Jard. Bot. Btzg Hl, 7 (1912) 21.—Type: KING
48 ‘p.p.’, Mamba, Papua (BO).
Rhizome short-creeping, 2-3 mm 9g; scales red-
dish brown, narrowly triangular, to 2 mm long,
to c. 8-seriate at base, with a short uniseriate
apex. Leaves close; petioles stramineous to pale
brown, or darker with age, abaxially subterete,
upwards obtusely or less often acutely bi-angular,
20-35 cm long, about equaling the lamina. Lamina
oblong, c. 15-35 cm long, with 2-4 pinnae to a
side and a conform terminal one (rarely simply
pinnate but fertile); primary rachis similar to the
upper part of the petiole, rarely shallowly
sulcate. Pinnae obliquely ascending, sessile, 12—25
cm long, 144-2% cm wide, widest in the lower
third or in the middle, gradually and strongly taper-
ing at the tip. Secondary rachises abaxially terete
at the base, upward gradually bi-angular, or bi-
angular almost to the base, usually little sulcate.
Pinnules c. 25-40 to a side, spreading or little
ascending, mostly subcontiguous, rounded-
trapeziform, the larger ones 9-15 by 4-7 mm.
about twice as long as wide; apex rounded, but
224
FLORA MALESIANA
[ser. II, vol. 13
the outer margin distinct, especially from the
lower, forming an angle of about 90° with the
slightly convex upper margin. Margins entire,
or shallowly crenate if sterile; texture firmly
herbaceous, dark green and + glossy above when
dry, sometimes described as bluish in the living
state. Upper pinnules strongly reduced, several
denticuliform ones connected with the small,
narrow terminal segment. Veins immersed, evi-
dent, not very oblique, regularly anastomosing,
with one, occasionally part of a second series of
areoles; larger areoles %-1 mm wide. Sori in
completely fertile pinnules continuous on upper
and outer margin; indusium entire, 1/,-'%4 mm
wide, not reaching the margin by an equal or
slightly larger distance, reflexed at maturity.
Spores medium brown, trilete, smooth, c. 22 p.
Distr. Malesia: New Guinea; a few collections
from Celebes and Borneo.
Ecol. Terrestrial in rain-forests, 80-1500 m.
Note. In habit this species is rather like the
South American L. guianensis (AUBL.) DRYAND.
ssp. lanceastrum KRAMER.
27. Lindsaea crispa BAKER, J. Bot. n.s. 8 (1879)
39; Hooker, Ic. Pl. 17 (1886) pl. 1627; C.
Cur. & Hortrum, Gard. Bull. S. S. 7 (1934)
238.—Type: BURBIDGE s.n., Sabah (K).
L. impressa Curist, Ann. Jard. Bot. Btzg II,
5 (1905) 132. —Type: HALLIER 3137, Amai Ambit,
Kalimantan, Borneo (L; fragm. in BO, P).
L. kinabaluensis Ho_ttrum, Gard. Bull. S. S. 7
(1934) 237.—Type: CLEMENS 25433, Tenompok
to Lumulumu, Mt Kinabalu, Sabah (SING;
dupl. in BM).
Rhizome short-creeping, 1-14, mm g; scales
medium brown, elongate-triangular, to c. 144 mm
long, to c. 10-seriate at base, with a short uniseriate
apex. Leaves clustered; petioles medium to dark
brown (rarely pale), in large leaves longer than the
lamina and abaxially mostly obtusely bi-angular,
in small leaves shorter than the lamina and
abaxially sharply bi-angular, pale-edged or not,
6-45 cm long. Lamina simply pinnate or bipin-
nate (very rarely subtripinnate), if bipinnate
with up to 3 pinnae to a side and a conform
terminal one, c. 15-20 cm long, if once pinnate,
c. 8-15 by 1%-2 cm. Primary rachis (if any)
like the petiole. Pinnae 10 by 11% to 18 by 3 cm;
pinnule-bearing rachises stramineous to dark
brown, abaxially sharply bi-angular, pale-edged
or not, + sulcate. Pinnules c. 25-30 to a side,
spreading, ascending, or falcately decurved, their
width apart to slightly overlapping, 1-elliptic
to falciform, 5 by 2 to 15 by 5 mm, 214-3 times
as long as wide, dark greén to blackish when dry,
chartaceous; apex of pinnules narrowed-rounded
to very obtuse, a distinct outer margin not de-
veloped. Upper pinnules gradually and strongly
reduced, several denticuliform ones confluent
into a narrow pinnatifid leaf- or pinna-apex.
Veins slightly impressed on the adaxial and pro-
minulous on the abaxial side, once or twice
forked, anastomosing to form an incomplete
series of areoles 144-1 mm wide, the outer veins
often free, in small (rarely in larger ones) pinnules
the veins may be quite free. Sterile margin sinuate-
dentate, fertile margin mostly crispate. Sori
continuous in fully fertile pinnules; indusium
delicate, pale, subentire, sometimes crispate,
0.3-0.5 mm wide, not reaching the margin by
%—-1 times its width, reflexed and often con-
cealed at maturity. the fertile margin sometimes
revolute over part of the sorus. Spores pale brown,
trilete, smooth, 20-22 wu.
Distr. Malesia: Borneo (Kalimantan, Sara-
wak, more frequent on Mt Kinabalu, Sabah).
Ecol. Terrestrial in forests, c. 1000-1600 m.
Note. The series of plants assigned to this
species is rather variable. If it were not for the
limited geographical distribution they might be
taken for depauperate specimens of one or several
other species.
28. Lindsaea hewittii COPELAND, Philip. J. Sc. 3
(1909) Bot. 346, pl. 5.—Type: Hewitt 36, Mt
Poi, Sarawak (MICH; dupl. in BM, P, SAR).
Rhizome short-creeping, 1-2 mm g; scales
castaneous, narrowly triangular, to c. 1 mm long,
to c. 7-seriate at the base, with a short uniseriate
apex. Leaves close; petioles rather dark brown,
faintly pale-margined, the greater part acutely,
the base obtusely quadrangular, abaxially -+-
sulcate, c. 22-30 cm long, twice as long as the
lamina. Lamina approximately rhombic in out-
line, 12-15 cm long, bipinnate, with 2 or 3 ascend-
ing pinnae to a side and a conform terminal
one; rachis like the petiole, pale to medium
brown. Pinnae linear, 9-12 cm long, 1-1%4 cm
wide, widest above the base, long-acuminate,
with c. 25-35 pinnules to a side; secondary
rachises reddish brown, abaxially sharply bi-angu-
lar, stramineous-margined, hardly sulcate. Pinnules
slightly ascending, not contiguous, chartaceous,
olivaceous when dry, ligulate or more often
¥,-elliptic, the larger ones 5 by 24% mm, twice
as long as broad; upper margin with 1 or 2
oblique incisions 4% mm deep or less; lobes with
a slightly convex or the inner ones with a
straight outer margin. Upper pinnules gradually
and very strongly reduced, several minute, denticu-
liform ones confluent with the linear, very narrow
pinna-apex. Veins immersed, + evident, very
close, regularly anastomosing, forming one series
of areoles c. ¥3 mm wide. Sori interrupted by the
incisions on the outer, sometimes also on the inner
lobes (the material probably incompletely fertile),
2- to 8-nerval; indusium brownish, subentire,
0.3-0.4 mm wide, not reaching the margin by an
approximately equal distance, scarcely reflexed
at maturity. Spores yellowish brown, trilete,
smooth, c. 20 mu.
Distr. Beside the type known from two other
collections from the same area.
Ecol. No data.
Note. This is perhaps only an extreme form of
the very variable L. obtusa.
29. Lindsaea obscura BRAUSE, Bot. Jahrb. 56
(1920) 132; CopELAND, Philip. J. Sc. 78 (1949)
April 1971]
20.—Type: LEDERMANN 8969, Etappenberg, Sepik
R. region, Terr. of New Guinea (B).
Rhizome rather short-creeping, | mm gQ;
scales not seen. Leaves close; petioles quadran-
gular, sulcate, blackish, narrowly pale-margined,
c. 5cm long, almost as long as the lamina. Lamina
simply pinnate, c. 5-7 % cm long; rachis similar to
the petiole. Pinnules c. 4-8 to a side, almost
contiguous at their bases, spreading, subtrapezi-
form to 44-elliptic-subligulate, 1-2 cm _ long,
5-8 mm wide, 2—2'% times as long as wide, dark
green when dry, herbaceous; upper margin out-
ward increasingly convex, a distinct outer margin
hardly or not developed. Margin of smaller
pinnules entire, in larger ones the upper/outer
margin with 1-3 oblique rather broad incisions
to 1 mm deep. Upper pinnules very little re-
duced; terminal pinnule free, large, flabellate with
cuneate base, almost symmetric, with convex
upper margin, incised like the lateral ones, c. 1
cm long and 1% cm wide. Veins immersed,
not evident, mostly twice forked, regularly anasto-
mosing, with one or towards the apex of the
pinnules two series of areoles 1-144 mm wide.
Sori interrupted by the incisions of the margin,
if any, also on the outer margin of the terminal
pinnule, 4- to plurinerval; indusium greenish,
subentire, 44 mm wide, falling short of the margin
by 14-1 times its width. Spores very pale brown,
trilete, smooth, c. 20 yw.
Distr. Beside the type only known from an-
other collection from the same area (LEDER-
MANN 9163a, B).
Ecol. Epiphytic in mountain forest, 850 m.
30. Lindsaea modesta KRAMER, Blumea 15 (1968)
LINDSAEA-GROUP (Kramer)
225
565.—Type: Aet & IDJAN 297, Waimiami near
Serui, Japen I. (BO).—Fig. 34.
Rhizome rather long-creeping, wiry, 0.4 mm @,
fuscous, dull; scales rather persistent, dark brown,
scarcely | mm long, acicular, biseriate in the
lower, uniseriate in the upper half. Petioles
Y%-%/, cm apart, to 7 cm long, a little longer
than the lamina, stramineous and quadrangular
almost to the base, slender, 44-14 mm @ at the
apex. Lamina c. 4-6 cm long, 3 cm wide, oblong,
simply pinnate, with 4-5 pinnules to a side;
rachis like the petiole. Pinnules thinly herbaceous,
olivaceous when dry, translucent, somewhat
ascending, their width or a little less apart,
obliquely triangular, the base rather strongly
cuneate, somewhat stalk-like, the apex subacute,
the largest 15 by 4-5 mm, almost evenly narrowed
from base to apex; upper margin with mostly 3
very oblique major incisions to 1 mm deep,
the lobes with slightly to strongly and irregularly
erose outer edge. Upper pinnules \ittle reduced.
Terminal pinnule free, flabellate-cuneate, 1-2 cm
long, usually once deeply and once or twice
shallowly bifid. Veins immersed, evident, very
oblique, once or twice forked, forming one series
of areoles, the outer veins not rarely free. Sori
mostly bi- or trinerval, in the larger lobes often
interrupted by the gashes in the margin, basally
straight or with slightly concave ends; indusium
0.3-0.4 mm wide, subentire or usually erose, not
reaching the margin by less than its width to
reaching it, -+_ reflexed at maturity. Spores pale
brownish, trilete, smooth, c. 20 yw.
Distr. Only known from the type collection.
Ecol. No data on the label, but probably
epiphytic, as the rhizomes are rather long and
interwoven with mosses and Hymenophyllaceae.
4. Section Lindsaea
Type species: Lindsaea trapeziformis DRYAND. [= L. lancea (L.) BEDD.].
Distr. Tropical America, SE. Asia.
Taxon. The two paleotropical species share with the neotropical ones a short-creeping rhizome,
a bipinnate lamina with conform terminal pinna, or a simply pinnate lamina, dimidiate pinnules, free
veins, and continuous sori. The spores are trilete in most neotropical and both paleotropical species.
Otherwise the resemblance is not close, in spite of much habitual similarity between L. borneensis HOOKER
and L. guianensis (AUBL.) DRYAND., and between L. doryphora KRAMER and L. lancea (L.) BEDD. It is
not at all certain that the paleotropical species are of the same derivation as the neotropical ones.
31. Lindsaea borneensis HOOKER ex BAKER,
Syn. Fil. ed. 1 (1867) 107; Hoxttrum, Gard.
Bull. S. S. 5 (1930) 65; Rev. Fl. Mal. 2 (1954)
331.—Lectotype: Lops 175, Labuan I., Borneo
(K).
Rhizome short- to long-creeping, 2-4 mm g;
scales castaneous, narrowly triangular, to 24% mm
long, to c. 8-seriate at base, with acicular, uni-
seriate apex. Leaves close; petioles stramineous,
abaxially terete, to 60 cm long, to 2 times as long
as the lamina. Lamina oblong, to c. 40 by 25 cm,
bipinnate, with up to 12 pinnae to a side and a
conform terminal one; primary rachis like the
petiole. Pinnae mostly subopposite, about their
width apart, linear, long-acuminate, spreading
or ascending, 8-25 cm long, 1144-2 cm wide,
the upper ones little or not shortened, the terminal
often the longest; secondary rachises abaxially
terete at base, upward gradually bi-angular, sul-
cate only in the upper part. Pinnules to c. 50 to
a side, dark green when dry, firmly herbaceous,
spreading, mostly slightly overlapping, 14—elliptic
or shortly ligulate, 7 to 11 mm long, 34-5 mm
wide; margin crenate if sterile, otherwise entire.
Upper pinnules gradually and strongly reduced,
denticuliform below the pinnatifid, sometimes
caudate pinna-apex. Veins immersed or more often
slightly prominulous, mostly twice forked, close,
free. Sori in fully fertile pinnules occupying all
vein-ends but often only on the inner ones, con-
226 FLORA MALESIANA [ser. Il voleale
JSG
Fig. 35. Lindsaea doryphora KRAMER. A. Simply pinnate lamina, x #/; (KORTHALS 298); B. Bipinnate
lamina, <x 4% (KORTHALS 296).
April 1971]
LINDSAEA-GROUP (Kramer)
Ze)
tinuous; indusium brownish, entire, c. !/,—¥, mm
wide, falling short of the margin by its own width
or less, reflexed and concealed at maturity.
Spores pale brown, trilete, smooth, c. 22 wu.
Distr. Malesia: Malay Peninsula (Selangor,
Johore, Perak), Singapore (according to HOLTTUM
perhaps extinct), Sumatra (few coll.), Borneo
(apparently frequent throughout).
Ecol. In forests, on shaded rocks, mostly on
poor, moist soil, to c. 700 m.
Note. In appearance rather like the neotropi-
cal L. guianensis (AUBL.) DRYAND. ssp. guianensis,
and perhaps related to it.
32. Lindsaea doryphora KRAMER, Blumea 15
(1968) 566.—L. scandens HOOKER var. terrestris
Hoitrum, Rev. Fl. Mal. 2 (1954) 327, nom.
invalid. (not typified).—L. lancea or L. trapezi-
formis auctt. quoad specimina asiatica.—Type:
ALSTON 13358, Permantang, S. of Kwala Kwajan,
Kalimantan, Borneo (U; dupl. in BM).—Fig. 35.
Rhizome rather short-creeping, 14.-2 mm g;
scales medium brown, very narrowly triangular, to
1144 mm long, to c. 12-seriate at base, with a short
uniseriate apex. Leaves close; petioles stramineous,
abaxially rounded or rarely upward obtusely bi-
angular, to 70 cm long, 1—2 times as long as the
lamina. Lamina 15—35 cm long, bipinnate or (only
when not yet full-grown?) simply pinnate, with
1-6 pinnae to a side and a conform terminal
One; primary rachis similar to the petiole. Pin-
nae (and simply pinnate lamina) to 25 cm long,
2%-5(-7) cm wide, ascending, half their width
apart to contiguous, widest below or in the middle
or sometimes scarcely narrowed close to the apex;
secondary rachises similar to the primary. Pin-
nules usually 20-25 to a side, subcontiguous,
chartaceous, dark green when dry, described by
many collectors as having a metallic sheen, some-
what variable in shape, if large ligular, spreading
or slightly decurved, not rarely with concave
lower margin, 2144-3 times as long as wide; if
smaller similar or rounded-trapeziform, 2-2'%
times as long as wide; largest pinnules seen 35 by
13 mm, but more often about 15-20 by 6-8 mm.
Upper margin outward increasingly convex, a
distinct outer margin hardly developed. Sterile
margin shallowly crenate towards the apex of the
pinnule. Upper pinnules in simply pinnate and
paucijugate-bipinnate leaves mostly little reduced,
few or none denticuliform and confluent with
the comparatively large, obliquely hastate, obtuse
to acuminate terminal pinnule; in larger, more
amply bipinnate leaves not rarely the upper pin-
nules more strongly reduced, several confluent
with the small, basally lobed, lanceolate terminal
segment. Veins immersed, evident, rather close,
free, mostly twice forked. Sori occupying all
vein-ends of a pinnule or not rarely only the inner
ones, continuous; indusium pale to dark, entire,
Y4,—1/, mm wide, falling short of the margin by
1-2 times its width, reflexed at maturity.
Spores pale brownish, trilete, smooth, c. 17 pu.
Distr. Tenasserim, S. Thailand, in Malesia:
Malay Peninsula, Singapore, Sumatra, Sipora,
Natuna Is., Lingga Is., West Java (rare), Bor-
neo, Philippines (Negros, Leyte, Mindanao).
Map 7.
Map 7. Distribution of Lindsaea doryphora
KRAMER.
Ecol. Terrestrial in moist to swampy forests,
from sea-level to 1300 m, apparently frequent in
large parts of its area.
Note. This species was described by HoLTTUM
as a variety of L. scandens (= L. parasitica);
in the present treatment the two species are placed
in different subgenera. Yet the similarity between
them is striking, and detached leaves, without
the rhizome, are sometimes difficult to determine
as the one or the other. The following additional
characters may help in determining such incom-
plete specimens:
L. doryphora: lamina of simply pinnate leaves
hardly or not narrowed at base; sterile pinnule-
margin crenate, especially near the apex; groove
of rachis (adaxial) narrow, quite concave; in-
dusium falling short of the margin by its width
or more; colour of dry pinnules dark green or oli-
vaceous.
L. parasitica: \jamina of simply pinnate leaves
gradually narrowed to base; sterile pinnule-margin
entire or faintly sinuate; groove of rachis (adaxial)
broad, flat- or convex-bottomed; indusium falling
short of the margin by its width or less; colour
of dry pinnules often blackish.
5. Section Osmolindsaea
KRAMER, Blumea 15 (1968) 560.
Type species: Lindsaea odorata Roxs. [L. cultrata auct. non (WILLD.) SWARTZ].
Distr. Rhodesia, Madagascar; N. India and China to Japan, Malesia and the Solomon Is.
Taxon. Well-marked by a short-creeping rhizome, simply pinnate lamina with abaxially rounded
axes and reduced upper pinnules, dimidiate free-veined pinnules with (except in reduced forms) broken
228
FLORA MALESIANA
[ser: I voli
sori, and monolete spores; the last character is not found in any other Malesian section. A group of rather
obscure affinity, perhaps related to the next.
33. Lindsaea odorata Roxs. Calc. J. Nat. Hist. 4
(1844) 511; KRAMER, Blumea 15 (1968) 567.—
Type: a plant from the Garrow Hills, India;
no specimen extant; pl. 2578 of Icones Rox-
burghianae to be regarded as type (K).
L. apiculata KuNze, Farrnkr. 1 (1846) 206,
pl. 4 A.—Type: SONDER 16, Java (n.v.).
L. cultrata (WILLD.) SWARTZ var. attenuata
Hooker, Sp. Fil. 1 (1846) 204.—Lectotype:
GRIFFITH s.n., Khasia, Assam (K).
L. cultrata (WILLD.) SWARTZ var. pallens
Hooker, /.c.—Type: WALLICH s.n., Nepal (K).
L. calomelanos KuNze, Bot. Zeit. 6 (1848)
214.—Type: ZOLLINGER 1892, Java (dupl. in B,
HBG EZ):
L. loheriana Curist, Bull. Herb. Boiss. I, 6
(1898) 44, pl. 4 f. 6.—Type: LoHER s.n., Atoc,
Luzon (P).
L. cultrata (WILLD.) SWARTZ var. varia COPE-
LAND in Perkins, Fragm. FI. Philip. (1905) 181;
Fern FI. Philip. 1 (1958) 108.—Type: ELMER 6003,
Baguio, Luzon (MICH; dupl. in B, BO, US).
L. crassipes ROSENSTOCK in Fedde, Rep. 5
(1908) 36; COPELAND, Philip. J. Sc. 78 (1949)
19.—Type: WERNER 74, Upper Mojo R., Terr.
of New Guinea (B; dup]. in L, US).
L. bullata v.A.v.R. Bull. Jard. Bot. Btzg II,
16 (1914) 20.—Type: DocTERS vAN LEEUWEN
1104, Mt Telemojo, Java (BO; dupl. in L.)
L. plumula Rip. J. Mal. Br. R. As. Soc. 4
(1926) 22; Ho_trum, Gard. Bull. S. S. 5 (1930)
62, f. 1-2.—L. cultrata (WILLD.) SWARTZ var.
plumula (RIDL.) HOLtTTuM, Rev. FI. Mal. 2 (1954)
329.—Type: RIDLEY 15997, G. Tahan, Pahang
(SING; dupl. in BM, K).
L. cultrata (WILLD.) SWARTZ var. parvula
Ho.ttum, Gard. Bull. S. S. 5 (1930) 61, f. 3.—
Type: Md. Nur 7297, Sibolangit, Sumatra (SING;
dupl. in K).
L. cultrata auct. non (WILLD.) Swartz of all
later authors, e.g. TAGAWA, Act. Phytotax.
Geobot. 6 (1937) 26, f. 1 A-C; BACKER & Post-
HUMUS, Varenfl. Java (1939) 118, f. 23; HoLtrum,
Rev. Fl. Mal. 2 (1954) 328, f. 188; COPELAND,
Fern Fl. Philip. 1 (1958) 107.
Rhizome short-creeping, dark, 14%-2% mm @,
occasionally more slender and long-creeping in
epilithic specimens; scales reddish brown, very
narrow, to 21% mm long, to 4-seriate at base,
Y%4—Y uniseriate. Leaves close (or occasionally
more remote); petioles (3-)6—-25 cm long, 1/3;—*/3,
rarely as long as the lamina, stramineous with
darker base or more often pale reddish brown to
atropurpureous throughout, hardly lustrous,
abaxially rounded or upward narrowed-rounded,
the adaxial groove often pale-edged. Lamina
narrowly oblong or narrowly lanceolate, 8-30 cm
long, 2-4% cm wide, simply pinnate, with 15-30
(rarely fewer) pinnules to a side, these subpetiolu-
late, alternate except the basal ones, spreading
or slightly ascending, less than their width apart
to contiguous, in the lower 144 or 4% more remote,
the basal ones usually remote and slightly or
occasionally more strongly reduced and sometimes
decurved. Rachis abaxially rounded or narrowed-
rounded, stramineous or dark at base (rarely
throughout), the pinnule-bases abruptly pale.
Larger pinnules 8-21 mm long, 3-8 mm wide,
2%, rarely 3 times as long as wide, ¥4-elliptic or
rarely asymmetrically oblong, subacute, less often
acute or obtuse, the truncate base sometimes
overlying the rachis, firmly herbaceous to sub-
coriaceous, occasionally coriaceous, mostly
light green or olivaceous when dry; lower margin
convex, upper margin straight or outward faintly
convex; rarely both margins evenly convex;
a distinct outer margin never developed. Upper
margin with 1-6 but little oblique, narrow incis-
ions, 44-1, rarely to 2 mm deep, reaching beyond
the receptacle, with acute sinus, the lobes with
flat or faintly convex or not rarely slightly concave
edge, sometimes with small, pointed, horn-like
extremities. Lobes of sterile pinnules crenate.
Upper pinnules gradually reduced, with few or no
incisions, some denticuliform ones confluent
with the small, narrow leaf-apex. Veins immersed,
evident or hidden, free, simple to twice forked,
4-1 mm apart. Sori interrupted by the incisions
except in reduced pinnules, (1—)2—6 per pinnule,
on 2 to 6 vein-ends, 1-6 mm long; receptacle
straight or in short sori often convex. Indusium
pale, entire to slightly erose, % mm wide (narrow-
erin small forms), not quite reaching the margin
to equaling it, with narrow, adnate ends, not re-
flexed but bulging at maturity. Spores medium
brown, monolete, smooth, c. 55 by 38 wu.
Distr. Rhodesia, Mascarenes?, Madagascar,
S. India, Ceylon, E. Himalaya, South and Central
China, Japan, to Malesia: Malay Peninsula (Pa-
hang, Perak), Sumatra, Java, Borneo, Philip-
pines (Luzon, Mindanao, Negros, Panay, Min-
doro), Talaud, Flores, New Guinea; Solomon Is.
So far not collected in Celebes or the S. Mo-
luccas.
Ecol. In open or slightly shaded places, often
on escarpments and earth banks, apparently
never in very dry places, from sea-level up to
over 2000 m. Apparently common in most parts
of its area, but rare east of Borneo and Java.
Notes. Freshly dried leaves have a distinct
coumarin-like odour which may persist for several
months in the herbarium.
In spite of its large area and rather diverse
habitats L. odorata is only moderately variable.
Only in extreme habitats aberrant forms occur,
e.g. in exposed places in the mountains where
the plants are dwarfed, with rigid pinnules and
very dark axes (‘L. crassipes’), or on moss-
covered tree trunks and rocks where they are
small but not rigid (‘L. loheriana’. ‘L. bullata’,
°L. plumula’, etc.).
Some small plants from the Philippines, e.g.
CuMING 65, MANGUBAT B. Sc. 1354, MERRILL
5930, approach a small form that occurs in China
April 1971]
LINDSAEA-GROUP (Kramer)
229
and Japan and is recently treated as a species,
L. japonica (BAKER) DIELS, but is probably not
rupted sori, and are more probably extreme forms
of L. odorata proper.
more than a variety. They have. however, inter-
6. Section Tropidolindsaea
KRAMER, Act. Bot. Neerl. 6 (1957) 267.
Type species: Lindsaea seemannii J. SMITH.
Distr. Very discontinuous; the neotropical species for which the section was described occur in
Costa Rica, Panama, W. Colombia, and Hispaniola; the sole paleotropical species occurs in the eastern
Philippines.
Taxon. The combination of short-creeping rhizome, once-pinnate lamina that tapers basally as well
as apically, and sclerotic, abaxially narrowed-rounded or carinate rachis, is sufficient for character-
ization. The pinnules are dimidiate and free-veined. The section is rather isolated in the genus; similar-
ities with sect. Osmolindsaea, and with the Madagascan group segregated as Sambirania by TARDIEU-
BLot, may be due to true relationship or to convergence.
34. Lindsaea adiantoides J. SmitH in Hooker,
Sp. Fil. 1 (1846) 204, pl. 61 C; CopELAND, Fern
Fl. Philip. 1 (1958) 108; KRAMER, Blumea 15
(1968) 560; non (BLUME) KUHN (1869).—L.
humilis KUHN in Mig. Ann. Mus. Bot. Lugd.-
Bat. 4 (1869) 278.—Type: CuMING 176, S. Cama-
rines, Luzon (K; dupl. in B, BM, E, GH, P, W).
L. tropidorachis v.A.v.R. Bull. Jard. Bot. Btzg
III, 5 (1922) 211; CopELANp, Fern FI. Philip. 1
(1958) 108.—Type: RAmMos & EDANO B. Sc.
33778, Camarines, Luzon (BO; dupl. in B, BM, P,
WS):
L. trimarginata C. CHR. Dansk Bot. Ark. 9
(1937) 68.—Type: RAMos & EDANO B. Sc. 33778,
Camarines, Luzon (B; dupl. in BM, BO, P, US).—
Fig. 24.
Rhizome very short-creeping or ascending, c.
14%, mm g, with densely scaly apex; scales
reddish brown, to almost 2 mm long, narrowly
triangular, up to S-seriate at base, biseriate below
the glandular top-cell. Leaves very close; petioles
blackish brown to atropurpureous, hardly shining,
sharply triangular to base, with somewhat paler
edges, the lateral faces much wider than the adaxial
one, 3-10 cm long, much shorter than the lamina.
Lamina simply pinnate, linear, ¢. 15-50 by 2-3
cm, widest mostly somewhat above the middle,
with c. 25-40 pinnules to a side; rachis similar
to the petiole, dark throughout, the edges often
irregular and more pronouncedly pale. Pinnules
firmly herbaceous, mostly olivacecus when dry,
subopposite almost throughout, spreading, close
to overlapping in the upper part, gradually more
remote in the lower part of the lamina, sessile;
larger pinnules obliquely triangular, often sub-
acute, 9 by 4% to 15 by 9 mm; lower margin
straight or slightly convex, ascending, upper
margin straight or somewhat convex to the apex,
with 1-3 narrow, unequal incisions that may reach
down to the middle but are usually much shallow-
er; a distinct outer margin not developed: bases
of pinnules abruptly non-sclerotic, suggesting
an articulation. Lower pinnules gradually very
remote, reduced, auriculiform, incised; upper pin-
nules gradually and strongly reduced, little or not
incised, 1 or 2 confluent with the lobed, non-
caudate terminal segment. Veins slightly impress-
ed above and prominulous beneath, giving the
pinnules a striate appearance, free, once or twice
forked, 144-34 mm apart. Sori one per lobe, on
4-10 vein-ends; receptacle straight, with convex
ends; indusium pale, herbaceous, erose, adnate at
the sides, 0.6-0.9 mm wide, mostly not quite
reaching the margin, scarcely reflexed at maturity.
Spores yellowish, trilete, almost smooth, c. 38-41 yu.
Distr. Malesia: Philippines (Luzon, Samar,
Dinagat, 9 coll.).
Ecol. Very few data; once reported from an
open bank.
Note. This species is taxonomically isolated in
Asia; its closest relatives are in tropical America.
7. Section Psammolindsaea
KRAMER, Blumea 15 (1968) 560.
Type and sole species the following, which has been included in Schizoloma and in Isoloma,
but is not closely related to either.
35. Lindsaea walkerae HooKER, Sp. Fil. 1 (1846)
209, pl. 69 A.—JIsoloma walkerae (HOOKER)
PREsL, Epimel]l. Bot. (1851) 101.—Schizoloma
walkerae (HOOKER) KUHN, Chaetopt. (1882)
346; Diets in E. & P. Nat. Pfl. Fam. I, 4 (1902)
218; HoL_trum, Rev. FI. Mal. 2 (1954) 344.—
Schizolegnia walkerae (HOOKER) ALSTON, Bol.
Soc. Brot. II, 30 (1956) 25.—Type: Mrs WALKER,
Ceylon (K; dupl. in B).
Rhizome rather short- to long-creeping, 114-2
mm @; scales reddish brown to castaneous, al-
most linear, to 2 mm, to 4-seriate at base, there
usually with laterally projecting cell partitions,
the apex uniseriate, paler. Leaves rather close to
4 cm apart; petioles dark castaneous to black,
lustrous, abaxially rounded, adaxially flattened
or broadly sulcate, c. 10-45 cm long, much shorter
to longer than the lamina. Lamina narrowly
230
FLORA MALESIANA
[ser. II, vol. 13
oblong, 15-70 cm long, 144-20 cm wide, simply
pinnate, with 3-17 pinnules to a side and a free
terminal one; rachis like the upper part of the
petiole. Pinnules chartaceous to rigidly coriaceous,
mostly olivaceous or dark brown when dry,
paler beneath, subsessile, obliquely to very strong-
ly ascending, a few cm apart, opposite or sub-
opposite throughout, linear, 24%-15 cm long,
4-8 mm wide, 6—20 times as long as wide; lower
pinnules often more remote and sometimes
slightly shortened; upper pinnules little or not
reduced. Base of pinnules slightly unequally
cuneate, the basiscopic side narrower, the dark
colour of the rachis ending rather abruptly in the
stalk-like base, but without an articulation;
margin entire, somewhat revolute; upper half of
pinnule narrowed, obtuse, if acuminate the tip
still obtuse. Terminal pinnule conform, symmetric,
occasionally joined to an upper lateral one or
lobed at base, usually soriferous. Costa distinct,
abaxially elevated, almost percurrent. Veins ele-
vated on both sides, very oblique, less so towards
their apices, 2-3 times forked, close, 4%4—-% mm
apart, free. Sori continuous, extending around the
apices of the pinnules, the vein-ends below the
receptacle thickened; indusium rigid, yellow or
brown, entire, 0.4 mm wide, almost reaching the
margin, somewhat reflexed at maturity. Spores
dark brown, trilete, smooth, 25-30 y.
Distr. Ceylon, Indo-China, in Malesia: Malay
Peninsula (Pahang, Trengganu, Johore, Malacca,
Kedah), Singapore, Lingga Is., Banka, Borneo
(Sarawak, Brunei, Kalimantan), W. New Guinea,
Queensland; Carolines (Palau, Yap, Truk,
Ponape). Rare in relation to its large area. Map 8.
i |
(
\
Map 8. Distribution of Lindsaea walkerae HOOK.
{
|
|
|
yi |
J
Ecol. In moist, open places, often on poor,
acid soil, in swamps and by streams, at low ele-
vation, but in Malaya also at 1000-1200 m.
8. Section Isoloma
(J. SMITH) KRAMER, Blumea 15 (1968) 560.—Jsoloma J. SmitH, Hook. J. Bot. 3 (1841) 414;
hist Pill (1875) 227.
Type species: Lindsaea divergens HOOKER & GREVILLE.
Distr. Throughout Malesia to western Polynesia. All species occur in the Flora Malesiana area.
Taxon. One of the groups treated most persistently as a distinct genus, mainly on account of the
non-dimidiate, basally articulate pinnules. Other characters are the short-creeping rhizome, sclerotic
axes, continuous sori, free veins, and trilete spores. The species of this section were in the past often
placed in the genus Schizoloma, together with the species lacking dimidiate pinnules of sect. Schizoloma
as treated here; but the resemblance is only a negative one.
36. Lindsaea gueriniana (GAUD.) DEsvAUx, Prod.
(1827) 312.—Schizoloma guerinianum GauD.
Ann. Sc. Nat. 3 (1824) 508; Freyc. Voy. Bot.
(1829) 380, pl. 18.—Jsoloma guerinianum (GAUD.)
Fre, Gen. Fil. (1852) 108; Tarprieu-BLot, Not.
Syst. 14 (1952) 332, comb. superfl.; COPELAND,
Fern FI. Philip. 1 (1958) 118.—Guerinia articulata
J. SmitH, Hist. Fil. (1875) 272.—Type: GaAupI-
CHAUD s.n., Rawak, Papua Is. (‘Moluccas’)
(Ridupleins Bike 1):
L. indurata BAKER, J. Bot. 26 (1888) 324.—
Schizoloma induratum (BAKER) C. Cnr. Ind. Fil.
(1906) 618.—Isoloma induratum (BAKER) TARDIEU-
BLoT, Not. Syst. 14 (1952) 332; TaGawa, Act.
Phytotax. Geobot. 16 (1956) 174, comb. superfl.
—Type: Hose 221, Niah, Sarawak (K; dupl.?
in BM).
Schizoloma fuligineum COPELAND, Philip. J.
Sc. 1 (1906) Suppl. IV, 252, pl. 1 A.—Isoloma fuli-
gineum (COPELAND) COPELAND, Philip. J. Sc. 78
(1949) 24.—Type: BoLsTtER 276, Surigao, Min-
danao (MICH; dupl. in P, US).
Nephrolepis schizolomae y.A.v.R. Bull. Jard.
Bot. Btzg II, 7 (1912) 22.—Schizoloma schizolo-
mae (y.A.v.R.) v.A.v.R. Handb. Suppl. (1917) 214,
—Isoloma schizolomae (v.A.v.R.) TAGAWA, Act
Phytotax.Geobot.16 (1956) 174.—Type: GJELLERUP
1000, Mt Cycloop, W. New Guinea (BO).—Fig. 36.
Rhizome short- to moderately long-creeping,
1144-2% mm g; scales reddish to dark brown,
lustrous, acicular, to 14 mm long, uni- or
biseriate or sometimes triseriate at the base.
Leaves close or not; petioles reddish brown to
fuscous, rarely blackish, rather dull, abaxially
terete, adaxially upwards (sometimes pale)
bimarginate, 5-25 cm long, much shorter than
to about equaling the lamina. Lamina linear,
10-30 cm long, 1%-3% cm wide, with up to c.
30 pinnules to a side; rachis medium to dark red-
dish brown, rarely blackish, dull, abaxially terete
at the base, upwards gradually carinate, occasion-
ally the keel extending to the base or present only
near the apex, sometimes pale, adaxial side +
distinctly pale-margined. Pinnules chartaceous to
coriaceous, usually drying brown, spreading or
slightly upcurved, 14-1 times their width apart
or less often contiguous, the lower ones remote,
not reduced, the upper ones gradually reduced,
April 1971]
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) —
lsh ‘
fc
}
‘ }
Fig. 36. Lindsaea gueriniana (GAUD.) DEsv.
Leaf, x ?/;, and portion of the rachis, x 4
(ZIPPELIUS s.n., L).—Fig. 37. L. lucida BL. ssp.
brevipes (COPEL.) KRAMER. Lamina, x 2/,
(BRASS 28253).
LINDSAEA-GROUP (Kramer)
231
the smallest c. 1 or 2 mm long, connected with the
small lanceolate terminal segment. Larger pin-
nules 7-18 mm long, 344-8 mm wide (excluding
basal auricles), asymmetrically ovate, 2-214 times
as long as wide, obtuse to broadly rounded or
rarely subacute, the acroscopic side of the base
mostly with a small but distinct auricle often touch-
ing the rachis; articulation at the bases of the
pinnules distinct. Veins immersed, obscure, once
or twice forked, free; costa evident, not percurrent.
Margin entire in fertile, crenate in sterile pinnules.
Sori continuous, or absent from the pinnule-
apices, always absent from the base of the lower
margin; indusium rigid, subentire, 4,—14 mm wide,
almost or quite reaching the margin, scarcely
reflexed at maturity. Spores medium brown,
trilete, almost smooth, c. 25-30 yp.
Distr. Malesia: Borneo (Sarawak), Celebes,
Philippines (Sibuyan, Mindanao), Moluccas
(Talaud, Halmahera), New Guinea (Japen, Wai-
geu, and other small islands; W. New Guinea,
Papua); Solomon Is., Tahiti. Many collections
are from small islands.
Ecol. In forests and open, drier places, terres-
trial, sometimes on decaying wood; from sea-level
up to 750 m.
Note. Some specimens, e.g. the type of L.
indurata and TEYSMANN 11822 from Celebes,
are atypical in having short, rounded pinnules.
L. gueriniana is the most variable species of sect.
Isoloma.
37. Lindsaea ovata J. SMITH in Hooker, Sp. Fil.
1 (1846) 204, pl. 64 A.—Isoloma ovatum (J.
SMITH) PRESL, Epimel. Bot. (1851) 101; HoLtrum,
Rev. FI. Mal. 2 (1954) 338; CoPpELAND, Fern FI.
Philip. 1 (1958) 117.—Schizoloma ovatum (J.
SMITH) COPELAND, Philip. J. Sc. 1 (1906) Suppl. IV,
252 (‘ovata’).—Type: CuMING 175, S. Camarines,
Luzon (K; dupl. in B, BM, E, W).
Schizoloma divergens (HOOKER & GREVILLE)
KURN var. auriculata v.A.v.R. Handb. (1908) 278.
—Schizoloma auriculatum (y.A.v.R.) v.A.v.R.
Bull. Jard. Bot. Btzg III, 5 (1922) 224.—TIsoloma
auriculatum (v.A.v.R.) TAGAWA, Act. Phytotax.
Geobot. 16 (1956) 174.—Type: coll. not cited,
‘Malaya’ (specimens in herb. BO so annotated
by v.A.v.R. are this species, but none is from
Malaya).
L. vrieseana ROSENSTOCK, Meded. Rijksherb.
31 (1917) 4, nom. subnud.—Type: DE VRIESE
52, Borneo, sine loc. (L).
Rhizome short-creeping, 2 mm g; scales dark
castaneous, to 1144 mm long, almost acicular, up
to 3-seriate at base, the apex long, uniseriate.
Leaves clustered; petioles dark brown or black,
lustrous, adaxially flattened and with two pale
lateral ridges, abaxially usually keeled in the upper
part, 6-18 cm long, 1% as long as to equaling the
lamina. Lamina very narrowly oblong, 8-20 cm
long, (1—-)2 %4—4 cm wide, scarcely or not narrowed
at the base, very shortly narrowed at the apex,
with up to c. 30 pinnules to a side; rachis abax-
ially keeled or rarely basally subterete, the keel
at least upward pale, otherwise the rachis like the
232
FLORA MALESIANA
[ser. II, vol. 13
petiole. Pinnules chartaceous, mostly drying
green, spreading or the basal ones deflexed,
asymmetrically ovate to lanceolate, (5—)10-20 mm
long, (3-)5-7 mm wide, 2-314 times as long as
wide, broadly rounded to subacute; the base at
the upper side sharply rectangular to shortly
auriculate. Lower pinnules not, a few upper ones
slightly, or no upper pinnules reduced, the termi-
nal pinnule free, rhombic to asymmetrically
triangular. Articulation at pinnule-bases distinct.
Fertile pinnules entire, sterile ones crenate-
sinuate or rarely lobed. Veins immersed but abax-
ially mostly evident, mostly twice forked; costa
weak, diagonal, not percurrent. Sori unbroken,
occupying the upper and outer or also part of the
lower margin; indusium yellowish to green, en-
tire, 0.3-0.4 mm wide, equaling the margin, +
reflexed at maturity. Spores rather pale brown,
trilete, verruculose, c. 22 yu.
Distr. Malesia: Malay Peninsula (Johore),
Singapore, Anambas Is., Lingga Is., Banka,
Borneo, Philippines (Luzon, Mindanao, no recent
coll.). Apparently common only in Sarawak.
Ecol. Terrestrial and on rocks, often on poor
soil and in moist places, from sea-level up to c.
1000 m.
38. Lindsaea pellaeiformis Curist, Verh. Naturf.
Ges. Basel 11 (1897) 430.—Schizoloma pellaeifor-
me (CHRIST) C. Cur. Ind. Fil. (1906) 619.—Iso-
loma pellaeiforme (CHRIST) TARDIEU-BLOT, Not.
Syst. 14 (1952) 332; TaGawa, Act. Phytotax.
Geobot. 16 (1956) 174, comb. superfl.—Type:
SARASIN 5.n., SE. Celebes (P).
Rhizome short-creeping, 2 mm g; scales medium
brown, to 24% mm long, acicular, the lower part
biseriate, the upper half uniseriate. Leaves
close; petioles dark castaneous to blackish, little
lustrous to dull, abaxially terete, to c. 20 cm long.
Lamina linear, to c. 30 cm long, to 544 cm wide,
widest a little above the base, with up to c. 35
pinnules to a side; rachis dark brown, dull, adaxi-
ally scarcely pale-margined, abaxially terete,
the upper half + pronouncedly carinate, the
keel pale. Pinnules chartaceous to coriaceous,
brown when dry, not contiguous, the basal ones
remote, spreading or slightly falcately ascending,
asymmetrically lanceolate, actue or rarely sub-
acute, the larger ones 18-30 by 8 mm, 1% to
almost 4 times as long as wide (disregarding the
auricle); base unequal, the basiscopic side narrow-
er, the acroscopic side with a short (to 2 mm)
subacute auricle. Margin entire, or notched in
sterile pinnules, somewhat revolute. Basal pin-
nules slightly or not reduced; upper pinnules
more strongly and gradually reduced, one or two
small ones connected with the narrow, asymmetri-
cally lanceolate terminal segment. Veins hidden,
once or twice forked, oblique; costa almost percur-
tent. Sori continuous; indusium brown, rigid,
entire, 4% mm wide, reaching the margin or
nearly so, little reflexed at maturity. Spores
medium brown, smooth, trilete, c. 27 yw.
Distr. Malesia: Celebes (3 coll.; beside the type
KJELLBERG 2059 and 3561, both BO and S-PA).
Ecol. Terrestrial in rain-forest, 0-300 m.
Note. A little known species, close to L.
gueriniana and perhaps only an extreme form of
that variable species. The upper pinnules are less
gradually reduced than in L. gueriniana but
much more strongly so than in L. ovata.
39. Lindsaea philippinensis KRAMER, nom. noy.—
Schizoloma angustum COPELAND, Philip. J. Sc. 1
(1906) Suppl. IV, 252, pl. 1 B.—Isoloma angustum
(COPELAND) TARDIEU-BLOoT, Not. Syst. 14 (1952)
332; TAGAWA, Act. Phytotax. Geobot. 16 (1956)
174, comb. superfl.; COPELAND, Fern FI. Philip. 1
(1958) 117, comb. superfl.; non L. angusta COPE-
LAND (1952).—Type:? FoxworTuHy 875, Palawan
(MICH).
Rhizome short-creeping, 144-2 mm g; scales
castaneous, lustrous, narrowly triangular to
almost acicular, up to S-seriate at base, the uni-
seriate apex relatively short. Leaves close;
petioles brown to blackish, often + persistently
fibrillose-scaly, abaxially terete or upward +
distinctly keeled, c. 3-15 cm long, mostly much
shorter than the lamina. Lamina linear, c. 10-30
cm long, 8-15 mm wide, with up to c. 35 pinnules
to a side; rachis abaxially (in small leaves) en-
tirely or (in larger ones) in the upper part keeled,
the keel evanescent downward, adaxially some-
times pale-margined, otherwise dark brown to
blackish, lustrous. Pinnules chartaceous or less
often coriaceous, not or hardly contiguous,
spreading, shortly and asymmetrically ovate,
5 by 4 to 8 by 6 mm, especially if large the base
acroscopically distinctly angular and often
slightly auriculate; upper pinnules gradually and
+ strongly reduced, the terminal pinnule asym-
metrically rhombic, free or connected with one
upper pinnule; lower pinnules more remote,
hardly or not reduced. Veins hidden, once forked
or simple, a costa scarcely developed. Sori oc-
cupying the whole margin except the inner;
indusium rigid, entire, c. 0.6-0.8 mm wide, not
reaching the margin by half its width or less,
little reflexed at maturity. Spores medium to dark
brown, trilete, subglobose, smooth, c. 30-35 yw.
Distr. Malesia: Philippines (Luzon, Sibuyan,
Palawan; 4 coll.).
Ecol. Among boulders on shaded riverbank,
800 m (one record).
Note. Doubtfully distinct from L. jamesoni-
oides, the variability insufficiently known.
40. Lindsaea jamesonioides BAKER, J. Bot. 17
(1879) 39; Hooker, Ic. Pl. 17 (1886) pl. 1626.—
Schizoloma jamesonioides (BAKER) COPELAND,
Philip. J. Sc. 1 (1906) Suppl. IV, 252.—J/soloma
jamesonioides (BAKER) TAGAWA, Act. Phytotax.
Geobot. 16 (1956) 174; TarpiEU-BLoT, Not.
Syst. 14 (1952) 332, with incorr. citation of
author of basionym.—Type: BURBIDGE S.n.,
Mt Kinabalu, Sabah (K; dupl. in BM).
Rhizome short-creeping, 144-2 mm g; scales
castaneous, narrowly triangular, to 3 mm long,
up to 8-seriate at the base, with a short uniseriate
apex. Leaves clustered; petioles black, dull, abaxi-
April 1971]
LINDSAEA-GROUP (Kramer)
23
ally rounded, adaxially with a flat groove, often
bearing some scattered dark narrow scales,
2-15 cm long, mostly considerably shorter than
the lamina. Lamina linear, c. 6-25 cm long, 8-18
mm wide, with up to c. 50 pinnules to a side;
rachis dark, not pale-margined, abaxially terete
at base, upwards (starting at the lower to upper
1) with a keel that becomes progressively more
acute above. Pinnules thickly coriaceous, some-
times wrinkled when dry, dark, half their width
apart to (above) overlapping, spreading or slight-
ly decurved, suborbicular to very shortly and
asymmetrically ovate, the smaller ones 4 by 4
to 5 by 6 mm, the larger ones 8 by 7 mm; base
unequal, basiscopically narrower, acroscopically
broadly rounded to subangular; margin entire,
also in sterile pinnules, thinner in texture and
paler, not rarely somewhat revolute. Lower
pinnules slightly or not remote and reduced,
upper pinnules gradually and more strongly
reduced but not confluent, the terminal pinnule
free or nearly so, suborbicular or asymmetrically
rhombic, occasionally lobed. Veins and_ sori
as in the preceding species. Spores rather dark
brown, trilete, smooth, c. 31-34 wu.
Distr. Malesia: Borneo (only known from Mt
Kinabalu, Sabah; 11 coll.).
Ecol. On moist rocks, in forest and open
places, c. 1500-3000 m.
Note. Young leaves described as bright red.
See also the note after the preceding species.
41. Lindsaea divergens HOOKER & GREVILLE, Ic.
Fil. (1831) pl. 226.— Vittaria divergens (HOOKER &
GREVILLE) ROXBURGH ex GRIFFITH, Calc. J. Nat.
Hist. 4 (1844) 510.—Schizoloma divergens
(Hooker & GREVILLE) KUHN, Chaetopt. (1882)
346; Diets in E. & P. Nat. Pfl. Fam. I, 4 (1902)
219, comb. superfl.—Isoloma divergens (HOOKER &
GREVILLE) J. SmitH, Hook. J. Bot. 3 (1841) 414;
HoLtrum, Rev. FI. Mal. 2 (1954) 337, f. 195;
COPELAND, Fern FI. Philip. 1 (1958) 117.—Type:
probably a specimen without data from herb.
Hooker, said to be a WALLICH coll. from Rox-
BURGH’s herbarium (K).
Rhizome short-creeping, 144-2 mm g; scales
very dark brown, to c. 1 mm long, almost acicular,
to 4-seriate at base, many scales largely or en-
tirely uniseriate. Leaves clustered; petioles black,
lustrous, abaxially terete, adaxially flattened and
with 2 sharp, occasionally paler lateral ridges
but hardly sulcate, to c. 15 cm long, mostly
about 144 as long as the lamina. Lamina narrowly
oblong, 15-50 cm long, 344-6 cm wide, narrowed
at both ends but more gradually at the base,
widest about the middle; rachis like the petiole,
abaxially keeled only near the apex. Pinnules
spreading or the upper ones ascending, the basal
ones deflexed, contiguous with their broadened
bases, elongate-ligulate, chartaceous to sub-
coriaceous, dark green or brown when dry, the
lower side markedly paler, often somewhat
glaucous; larger pinnules 134-3 cm long, 3-4 mm
wide (not counting the broadened base), little
narrowed to the obtuse apex; acroscopic side
of base with an obtuse or subacute auricle that
does not touch the rachis; basiscopic side less
broadened. Lowermost, reflexed pinnules mostly
several mm long, scarcely auriculiform; upper-
most pinnules more strongly reduced, a few
confluent with the lobate-hastate, lanceolate
terminal segment, or a free, scarcely lobed,
lanceolate pinnule present. Margin entire, often
revolute when dry, then the pinnules seemingly
acute. Veins immersed, obscure, lax, very oblique,
mostly once or twice forked. Sori continuous,
absent from the basal auricle and sometimes,
from the apex; indusium 0.2-0.3 mm wide, pale,
entire, not reaching the margin by less than its
width. Spores medium brown, trilete, smooth, c.
17-19 pw.
Distr. S. Thailand; Malesia: Malay Peninsula,
Singapore, Riouw Is., Lingga Is., Sumatra,
Banka, Borneo, Philippines (Palawan).
Ecol. Terrestrial in forest, from sea-level up
to c. 800 m.
9. Section Stenolindsaea
Kramer, Blumea 15 (1968) 561.
Type species: Lindsaea lucida BLUME.
Distr. A small section of only three species, occurring from China and Southern Japan to Fiji.
Taxon. Except for its constantly unipinnate leaves and basally tapering lamina (not always distinct
in L. Jucida BLUME) this section agrees in technical characters with sect. Temnolindsaea, but the groups
are probably not closely related. The axes are abaxially bi-angular.
42. Lindsaea lucida BLUME, En. PI. Jav. (1828)
216; Hottrum, Rev. Fl. Mal. 2 (1954) 328, f.
187; KRAMER, Blumea 15 (1968) 567.—Type:
BLUME s.n., Java (L).
L. gracilis BLUME, En. Pl. Jav. (1828) 217, non
Kiorzscu (1844); not or only in part of later
authors.—Type: BLUME s.n., Java (L).
L. concinna J. SmitH, Hook. J. Bot. 3 (1841)
415, nom. subnud.; HooKer, Sp. Fil. 1 (1846)
205, pl. 61 B.—L. cultrata (WILLD.) SWARTZ var.
concinna (J. SMITH) Domin, Bibl. Bot. 20 (1915)
82.—Type: CumING 198, Luzon (K; dupl. in B,
BNGHSHBG) Ey ESSING US iW):
L. securifolia PRESL ex GOLDMANN, Nova Acta
Ac. Caes. Leop.-Car. Suppl. I ad 16 (1843) 464;
Kunze, Farrnkr. 1 (1846) 204, pl. 85 f. 1; CopE-
LAND, Fern FI. Philip. 1 (1958) 108.—L. cultrata
(WILLD.) SWARTZ var. securifolia (PRESL) BAKER
in Beccari, Malesia 3 (1886) 36.—Type: MEYEN
s.n., Manila, Luzon (dupl. in B, HBG).
234
FLORA MALESIANA
[ser. IT, vol. 1
L. colobodes KUNZE, Bot. Zeit. (1846) 446.—
Type: ZOLLINGER 1507, Java (dupl. in B, HBG,
W, Z).
L. lobbiana Hooker, Sp. Fil. 1 (1846) 205,
pl. 62 C; Hottrum, Gard. Bull. S. S. 5 (1930)
61.—L. cultrata (WILLD.) Swartz var. lobbiana
(HooKER) Beccari, Malesia 3 (1886) 36.—
Type: Loss s.n., Java (K).
L. crenulata Fre, Gen. Fil. (1852) 105, pl.
28 f. 2.—Type: GRIFFITH s.n., ‘Malacca?’ (n.v.).
L. gracilis BLUME var. major METT. ex KUHN
in Miq. Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 276.—
Lectotype: HASSKARL s.n., Java (L).
L. gracillima COPELAND in Perkins, Fragm.
Fl. Philip. (1905) 181.—Type: MERRILL 287,
Caraballo Sur, Luzon (dupl. in B, BO, US).
L. propria v.A.v.R. Bull. Jard. Bot. Btzg II,
16 (1914) 20.—Type: BACKER 6084, Mt Hand-
jawung near Bogor, Java (BO).
Rhizome short- or very short-creeping, c.
1 mm @g; scales rather dark brown, to 1 mm long,
acicular, to 6-seriate at the base, the greater part
uni- or biseriate. Leaves very close; petioles
stramineous to reddish brown, quadrangular,
adaxially hardly sulcate. Lamina linear, c. 10-40
cm long, c. 10-25 times as long as wide, simply
pinnate; rachis like the petiole. Pinnules spreading
or slightly ascending or decurved, the upper ones
subcontiguous to slightly overlapping, the lower
ones gradually more widely spaced; larger pin-
nules 1144-2% times as long as wide, firmly
herbaceous, medium to dark green or olivaceous
when dry. Upper pinnules reduced; larger pin-
nules mostly incised. Veins immersed, evident,
once or twice forked or the outer ones simple,
free; a costa scarcely developed. /ndusium pale,
entire; spores trilete.
KEY TO THE SUBSPECIES
1. Petiole 3-15 cm long, 1/;—% of the length of
the lamina; indusium 0.3-0.4 mm wide, at
least in the inner sori almost or quite reaching
(iveimeigain, ¢ oo 5 6% 1. ssp. lucida
1. Petiole 1-2(-4) cm long, +/,;—1/25 of the length
of the lamina; indusium 0.2 mm wide, more
strongly intramarginal. 2. ssp. brevipes
1. ssp. lucida.
Petioles c. 3-15 cm long, 1/;—¥ as long as the
lamina. Lamina 1-3 cm wide, with c. 25-60 pin-
nules to a side. Larger pinnules 5 by 3 ta 15 by
6 mm, almost 2—21% times as long as wide, rather
variable in shape, 14-elliptic or subtriangular if
small, ligulate to subtrapeziform or elongate-
Y,-elliptic if larger, the apex subacute to broadly
rounded, the lower margin + straight, the upper
margin especially in small pinnules evenly con-
vex and no separate outer margin developed,
or outward increasingly convex, the outer margin
+ distinct. Upper pinnules gradually and suddenly
rather strongly reduced, some denticuliform ones
connected with the narrow, small, lobed, usually
obtuse and often fertile leaf-apex. Lower pinnules
not reduced or more often slightly to strongly
and gradually reduced, then often somewhat
decurved, in extreme cases only 2 mm long but
usually larger, the petiole below them well
developed. Sterile pinnules crenate; fertile pin-
nules if very small entire or with 1 or 2 incisions
on the outer/upper margin to 4% mm deep, in
larger forms rarely entire, mostly at least with
one incision of %—-l1 mm between upper and
outer margin, most commonly the upper margin
with 1-3 incisions of c. 1 mm (but up to 2 mm),
the outer entire. Sori continuous except as inter-
rupted by the incisions, on 2-—7(—10) vein-ends,
the outermost usually longer; upper, smaller
pinnules entire, fertile; lower, reduced ones, if
any, partly or entirely sterile. Receptacle straight,
or concave in the outer sorus. Indusium 0.3—0.4
mm wide, little narrowed at the edges, almost or
quite reaching the margin, or more strongly
intramarginal in the outer sori, reflexed or not at
maturity. Spores yellowish brown, smooth, c.
25-28 pw.
Distr. Central Himalaya, S. China, and Ryu
Kyu Is. to Malesia: throughout Malesia (except
Lesser Sunda Is.) but rare in New Guinea; Palau
Is.
Ecol. Terrestrial or on rocks, rarely epiphytic,
in primary forests, from 400-1600 m, sporadically
to sea-level.
2. ssp. brevipes (COPELAND) KRAMER, Blumea 15
(1968) 567.—L. brevipes COPELAND, Philip. J.
Sc. 6 (1911) Bot. 83; ibid. 78 (1949) 19.—Type:
KING 237, Papua (MICH; dupl. in BO).—
Fig. 37.
Petioles 1-2(—-4) cm long, 1/,;—'/o) of the length
of the lamina. Lamina 14%-2 cm wide, widest
above the middle, very gradually long-tapering
at the base, rather shortly acuminate at the apex,
with 25-80 pinnules to a side. Larger pinnules
6-9 mm long, 3-4 mm wide, mostly 4-elliptic,
occasionally subligulate, mostly with straight
lower and outward increasingly convex upper
margin and + distinctly angular at the junction
of outer and upper margins. Incisions various,
occasionally none, usually 1-3 per pinnule, 1-1,
sometimes to 2 mm deep and then reaching to or
the outer ones beyond the middle. Upper pin-
nules as in ssp. lucida. Lower pinnules from above
the middle of the lamina downward gradually
and strongly reduced, many lower ones auriculi-
form, 1-2 mm long, sterile, crenate to cleft.
Indusium 0.2 mm wide, not reaching the margin,
usually reflexed and + concealed at maturity.
Spores c. 20 uw, otherwise as in ssp. lucida.
Distr. Malesia: S. Moluccas (Ceram, Am-
bon), New Guinea (Waigeu, W. New Guinea,
Terr. of New Guinea, Papua); Admiralty Is.
(Los Negros), Solomon Is.
Ecol. Terrestrial and on rocks and tree-bases,
in forests, from sea-level up to c. 1000 m.
Note. This is a somewhat variable species.
The extremes certainly look different, e.g., the
form described from (and particularly common
in) the Philippines as L. concinna with short
pinnules and basally rather strongly tapering
April 1971]
lamina; but they all grade into each other. Ssp.
brevipes alone seems reasonably distinct, although
there are a few intermediates, notably in New
Guinea.
43. Lindsaea bakeri (C. Cur.) C. Cur. Ind. Fil.
Suppl. 3 (1934) 121; CopELAND, Philip. J. Sc.
78 (1949) 17.—Davallia lanceolata BAKER, Kew
Bull. (1899) 119; non Asplenium lanceolatum
Hupson (1762), nec L. lanceolata LABILLARDIERE
(1806).—Asplenium bakeri C. Cur. Ind. Fil.
(1906) 102.—Type: GIULIANETTI & ENGLISH
s.n., Vanape valley, Papua (K).
L. trichophylla COPELAND, Philip. J. Sc. 6
(1911) Bot. 83.—Type: KING 262, Papua (MICH;
dupl. in BRI, ?P).
L. schlechteri BRAUSE, Bot. Jahrb. 49 (1912) 28,
f. 1 J; CopELAND, Philip. J. Sc. 78 (1949) 17.
—Type: SCHLECHTER 19395, Pema, Terr. of New
Guinea (B).
L. ledermannii BRause, Bot. Jahrb. 56 (1920)
130.—Lectotype: LEDERMANN 10017, Lordberg,
Sepik R. region, Terr. of New Guinea (B; dupl.
in S-PA).—Fig. 22.
Rhizome short-creeping, 44-24 mm 4g; scales
not seen. Leaves clustered; petioles abaxially
obtusely to sharply bi-angular, 1-10 cm long,
shorter than to about equaling the lamina.
Lamina linear, once pinnate + deeply uni- to
tripinnatifid, narrowed on both sides or truncate
at the base, with c. 20-50 pinnules to a side;
rachis abaxially sharply bi-angular, hardly
sulcate. Pinnulés subcontiguous to slightly over-
lapping, the basal ones farther apart, spreading,
slightly ascending, or somewhat falcately decurved,
thinly herbaceous, medium to dark green when
dry, 5 by 24% mm to 3 by 1% cm, semi-ovate to
suboblong in outline, 2-2% times as long as
wide, very deeply incised from the upper margin,
with 3-9 primary segments (in one plant at the
apex also incised from the lower edge), these
rarely all simple, mostly at least the basal ones
once or twice forked, or in very large pinnules
acroscopically-dimidiately or subequally pinna-
tifid, then some of the secondary segments again
bifid; ultimate lobes + divergent, linear, often
with revolute margin, gradually but slightly
broadened upward, then suddenly broadened at
the sorus, the wing connecting them about as
wide as the segments. Veins immersed, single or
LINDSAEA-GROUP (Kramer)
235
very rarely paired in the lobes. Upper pinnules
gradually reduced, confluent into a pinnatifid
leaf-apex; basal pinnules -+- reduced. Sori uni-
or binerval; indusium pale or greenish, delicate,
subentire, laterally free, 14 mm wide, not reaching
the margin by its width or a little less, not
strongly reflexed at maturity but the sporangia
often spreading beyond the margin. Spores pale
brown, trilete, smooth, c. 20 uw.
KEY TO THE VARIETIES
1. Axes stramineous to medium brown; ultimate
lobes 0.3—0.5 mm wide, at the sorus 4-1 mm
wide; sori hardly ever binerval 1. var. bakeri
1. Axes medium to dark brown; ultimate lobes
0.6—1 mm wide, 0.8—1.2 mm wide at the sorus;
sori not rarely binerval. 2. var. pycnophylla
1. var. bakeri.—Fig. 22.
Petioles stramineous to medium brown, then
not rarely pale-angled. Lamina 4-30 cm long,
1-6 cm wide; rachis stramineous or basally pale
brown. Ultimate lobes of the pinnules divergent,
linear-capillary, 0.3-0.5 mm wide, at the sorus
0.5—1 mm wide, their apices erose, rounded or
subtruncate, not rarely laterally bicorniculate,
rounded-subacute when sterile. Veins evident.
Sori very rarely binerval, 0.3-0.8 mm long; base
of indusium + straight.
Distr. Malesia: Moluccas (Morotai),
Guinea (all Div.).
Ecol. Terrestrial, on banks and mounds, and
on tree-bases, in forests, from c. 80-1000 m.
New
2. var. pycnophylla KRAMER, Blumea 15 (1968)
567.—Type: Brass 6838, Fly R. region, Papua
(GH).
Petioles and rachises medium to dark brown,
pale-margined. Lamina to 12 by 3 cm, little
narrowed at base. Pinnule lobes not capillary,
0.6-1 mm wide, at the sorus 0.8-1.2 mm wide,
little divergent, apically hardly or not erose.
Veins concealed. Sori not rarely binerval, to 1 mm
long. Base of indusium not rarely strongly
concave.
Distr. Malesia: Papua (2 coll.; beside the type
also Brass 6902, BM, GH, from the same area).
Ecol. 80-100 m.
Subgenus Odontoloma
(Hooker) Kramer, Blumea 15 (1968) 561.—Davallia subg. Odontoloma Hooker, Sp. Fil. 1
(1845) 174.—Odontoloma J. SmitH, Hook. J. Bot. 3 (1841) 415, nom. subnud.; in Hooker &
Bauer, Gen. Fil. (1842) pl. 114 B; non H.B.K. (1820) (Compositae).
Type species: Dicksonia repens Bory = Lindsaea repens (BORY) THWAITES.
10. Section Odontoloma
Type species: As the subgenus.
Distr. Mascarene Islands, Ceylon, NE. India, Ryu Kyu Is. to Hawaii, and NE. Australia.
Taxon. A group of closely related species with closed xylem strand, + persistently scaly rhizome, and
236
FLORA MALESIANA
[ser. Il, voleere
unipinnate leaves. The differences between most taxa are slight, and the present species and varieties
are not quite satisfactory.
44. Lindsaea glandulifera v.A.v.R. Bull. Jard.
Bot. Btzg II, 1 (1911) 9.—Type: KoorpeErs
15415, Besuki, Java (BO; dupl. in K, L).
L. repens (BoRY) THWAITES var. laciniata METT.
ex KUHN in Miq. Ann. Mus. Bot. Lugd.-Bat. 4
(1869) 277.—Lectotype: ZOLLINGER II 3712, Mt
Idjen, Java (L; dupl. in B, BM, P, W).—Fig. 49.
Rhizome long-scandent, brown, 1-2 mm g,
dorsally + persistently scaly; scales honey-
coloured to pale golden-brown, narrowly trian-
gular, to c. 25-seriate at base, to 4 by 1 mm, the
uniseriate apex of up to 10 cells. Leaves 1-3 cm
apart, issuing under an acute angle; petiole al-
most wanting to c. 2 cm long, stramineous, darker
and verruculose at base, sharply quadrangular
almost to the base, laterally and sometimes also
adaxially broadly and shallowly sulcate. Lamina
linear, 20-35 cm long, 144-2'% cm wide, simply
pinnate, with c. 40-60 pinnules to a side; rachis
stramineous, abaxially flat, sharply bi-angular,
adaxially broadly sulcate, laterally + sulcate.
Pinnules regularly spaced except at the base, not
contiguous, spreading or mostly somewhat ascend-
ing, at least the basal ones inserted below the level
of the adaxial-lateral ridges, asymmetrically
elongate-triangular, herbaceous, bright green
to olivaceous when dry, the largest 10-12 by
4-5 mm, + evenly narrowed from base to apex,
cuneate at base, narrowed-rounded at apex;
lower margin mostly somewhat convex; upper
margin outward convex, incised, with mostly
3 major, oblique incisions reaching down to 14 or
'% or occasionally to 34, the lobes not parallel-
sided but narrowed, rounded at apex, the largest
lobes shallowly incised again. Pinnules of juvenile
plants and sterile ones not more incised. Lower
pinnules gradually remote and reduced, largely
sterile; upper pinnules gradually and strongly
reduced, confluent into a pinnatifid leaf-apex.
Veins immersed, evident, simple or once forked.
Sori one per lobe, on the ends of 1 or 2 connivent
veins; indusium transversely elongate to hippo-
crepiform, with -+- concave base, free at the
narrowed sides, 43-1 mm long, c. 4% mm wide,
not reaching the margin by its width or a little
less, reflexed and concealed at maturity. Spores
pale brown, trilete, nearly smooth, c. 22 w.
Distr. Ceylon; in Malesia: East Java, Lesser
Sunda Islands (Lombok, Sumbawa). Map 10.
Ecol. Epiphytic in mountain forests, c. 1000 m.
Apparently rare.
45. Lindsaea oblanceolata v.A.v.R. Bull. Jard.
Bot. Btzg II, 23 (1916) 15.—Type: AsoeB 100,
Rimbo Pengadang, Bencoolen, Sumatra (BO;
dupl. in L).
L. subsemilunularis v.A.v.R. Bull. Jard. Bot.
Btzg II, 28 (1918) 31.—Type: HaALuier 591,
Mt Damus, Borneo (BO).
? L. lunulata v.A.v.R. Bull. Jard. Bot. Btzg
II, 11 (1913) 15.—Type: Raap 634, Batu Islands
(BO). .
L. pectinata auct. non BLUME; HOoLtrum,
Gard. Bull. S. S. 5 (1930) 62; Rev. Fl. Mal. 2
(1954) 324.
Rhizome l\ong-scandent, dark castaneous, 1-2
mm @; scales when mature dark brown, narrowly
triangular or lanceolate, to 24% by 34 mm, up
to c. 15-seriate at the base, the uniseriate apex
very short. Leaves remote, 1-5 cm apart, not
rarely issuing almost at right angles; petioles
abaxially bi-angular, mostly brown and _ pale-
angled but sometimes quite stramineous except
at the base, 2-10 cm long (exceptionally longer),
much shorter than the lamina, at the most half as
long. Lamina linear, 15-50 cm long (rarely shorter
but fertile), 2-414 cm wide, 5-15 times as long as
wide, in juvenile plants usually broadest just below
the apex, in full-grown ones narrowed to both
ends, simply pinnate, with c. 20-60 pinnules to
a side; rachis pale brown, stramineous-margined,
or stramineous throughout, at the base sharply
quadrangular with flat or slightly convex faces,
the pinnules in the basal part inserted below the
adaxial face, upward sulcate. Pinnules herba-
ceous or chartaceous, dark green or blackish when
dry, sometimes described as glossy, less than their
width apart to slightly overlapping, spreading, the
lower ones not rarely deflexed and the upper
ones somewhat ascending, dimidiate-ligulate or
trapezoidal, broadly rounded or narrowed-
rounded at the apex; larger pinnules 10 by 5
to 22 by 9 mm, 2-2'% times as long as wide,
entire; sterile or incompletely fertile pinnules
crenate. Lower pinnules gradually reduced but
few or none so strongly as to be auriculiform;
upper pinnules not reduced in juvenile plants,
increasingly so in older ones, in the largest leaves
several very small ones connected with the small
lanceolate terminal segment; small leaves with a
large, free, very obtuse, transversely elongate,
asymmetrically lance-shaped or flabellate or broad-
ly lanceolate terminal pinnule, with all intermedi-
ates between the two extremes possible. Veins
immersed, + evident, once or twice forked, free,
34-14% mm apart. Sori continuous, occupying
all vein-ends, interrupted only in incompletely
fertile pinnules; indusium 0.3—0.5 mm wide, pale,
subentire, not reaching the margin by 15-1 times
its width. Spores yellowish brown, trilete, almost
smooth, c. 20 yu.
Distr. Annam, Peninsular Thailand; in Ma-
lesia: Malay Peninsula, Sumatra, West Java,
Borneo, Philippines (Luzon, Leyte, Mindanao),
Moluccas (Ceram, Ambon). Not yet found on
Celebes.
Ecol. On tree trunks, or occasionally terrestrial,
800-1800 m, rarely lower. Apparently uncommon
in most parts of its area.
Notes. Incompletely fertile leaves without
rhizome cannot always be distinguished from L.
repens. Juvenile plants lack the deeply incised pin-
nules of that species.
The type of L. lunulata y.A.v.R. is a juvenile,
April 1971]
nearly quite sterile plant of uncertain identity.
46. Lindsaea repens (Bory) THwaites, En. PI.
Zeyl. (1864) 388; Bepp. Ferns S. Ind. (1864?)
pl. 209; Curist, Geogr. d. Farne (1910) f. 46,
47; Hotttum, Gard. Bull. S. S. 5 (1930) 63, f. 4.
—Dicksonia repens Bory, Voy. 2 (1804) 323.—
Davallia repens (Bory) DEsvAUx, Prod. (1827)
314, non KUHN _ (1867).—Odontoloma repens
(Bory) PRESL, Epimel. Bot. (1851) 97.—Acro-
phorus repens (Bory) Moore, Ind. Fil. (1857) 91.
—Davallia boryana PRESL, Rel. Haenk. (1825) 66,
as to type only.—Saccoloma boryanum (PRESL)
Pres_, Tent. Pterid. (1836) 126.—Odontoloma
boryanum (PRESL) J. SmitH, Hook. J. Bot. 3
(1841) 415 (‘boryana’).—L. boryana (PRESL)
BrAUSE, Bot. Jahrb. 56 (1920) 129.—Type: Bory
s.n., Bourbon (= Réunion) (P; dupl. in B, BM).
L. bantamensis BLUME, En. PI. Jav. (1828) 218.—
Type: BLUME s.n., Bantam, Java (L).
L. hymenophylloides BLUME, I.c.; not or in part
only of later authors.—Acrophorus hymenophyl-
loides (BLUME) Moore, Ind. Fil. (1857) 2.—Da-
vallia hymenophylloides (BLUME) BAKER, Syn. Fil.
ed. 1 (1867) 93.—Odontoloma hymenophylloides
(BLUME) J. SMITH, Hist. Fil. (1875) 269.—Type:
BLUME s.n., Java (L).
L. pectinata BLuMe, En. Pl. Jav. (1828) 217;
not of Ho_trum, Rey. Fl. Mal. 2 (1954) 324, nor
of COPELAND, Fern FI. Philip. 1 (1958) 104.—
Type: BLUME s.n., Java (L).
Davallia hemiptera Bory, Bél. Voy. Bot. 2
(1833) 73, pl. 7 f. 2.—Saccoloma hemipterum
(Bory) Pres_, Tent. Pterid. (1836) 126.—Odon-
toloma hemipterum (Bory) PREsL, Epimel. Bot.
(1851) 98.—Acrophorus hemipterus (Bory)
Moore, Ind. Fil. (1861) 295.—L. repens (Bory)
THWAITES var. hemiptera (Bory) v.A.v.R. Handb.
(1908) 261; not L. hemiptera KRAMER (1957).—
Type: BELANGER s.n., Java (P).
L. oblongifolia REINW. ex HooKeEr, Sp. Fil. 1
(1846) 206, at least in part.—Type: CUMING 186,
S. Camarines, Luzon (K; dupl. in B, GH, L,
SING, US, W).
Davallia delicatula CHRIst, Verh. Naturt. Ges.
Basel 11 (1895) 224, pl. 3, 1-3.—L. delicatula
Curist, /.c., nom. altern. illeg—Type: SARASIN
987, Borau, Celebes (P).
L. sessilis COPELAND, Philip. J. Sc. 6 (1911)
Bot. 82; ibid. 78 (1949) 16.—Type: KING 244,
Papua (MICH).
L. repens (BoRY) THWAITES var. intermedia
Curist, Ann. Cons. Jard. Bot. Genéve 15/16
(1912) 197.—Type: HoOcCHREUTINER 782, Mt
Salak, Java (n.v.).
L. foersteri ROSENSTOCK in Fedde, Rep. 12
(1913) 527; CopELAND, Philip. J. Sc. 78 (1949)
17.—Type: Keysser 193, Mt Sattelberg, Terr. of
New Guinea (B; dupl. in MICH).
L. pectinata (Bory) TuHwatites f. dimorpha
ROSENSTOCK in Fedde, Rep. 13 (1914) 213.—Type:
GRASHOFF, 59, W. Sumatra (S-PA).
L. repens (Bory) TuHwaites f. truncatiloba
ROosENSTOCK, /.c. 214.—Type: GRASHOFF 60,
W. Sumatra (S-PA).
LINDSAEA-GROUP (Kramer)
237
L. longa COPELAND, Philip. J. Sc. 46 (1931) 216;
Fern Fl. Philip. 1 (1958) 103.—Type: EpDANo
B.Sc. 77978, Mt Balabag, Palawan (MICH;
dupl. in GH).
L. sagincola WAGNER & GRETHER, Un. Cal.
Publ. Bot. 23 (1948) 34, pl. 9.—Type: GRETHER
& WAGNER 4010, Los Negros, Admiralty Is.
(dupl. in MICH, US).
L. cultripinna COPELAND, Philip. J. Sc. 81
(1952) 6; Fern Fl. Philip. 1 (1958) 103.—Type:
SuLit P.N.H. 8718, Mt Katanglad, Mindanao
(MICH).
? L. roxasii COPELAND, Philip. J. Sc. 81 (1952)
6; Fern Fl. Philip. 1 (1958) 102.—Type: Coprg-
LAND P.P.E. 259, Mindanao (MICH).—Fig.
43-45.
Rhizome long-scandent, castaneous; scales tri-
angular, narrowly triangular, or lanceolate, with
a very short uniseriate apex, the larger ones 2%4—
3144, mm long. Leaves one to a few cm apart,
issuing under a small angle; petioles stramineous
to pale brown with dark base, quadrangular, the
faces not sulcate. Lamina linear, simply pinnate,
narrowed at both ends, or sometimes little narrow-
ed at the base; rachis stramineous, quadrangular,
adaxially at the base + convex, the pinnules
inserted below the edges, upwards sulcate, bearing
the pinnules at its edges. Pinnules numerous,
herbaceous, elongate-triangular, ligulate, or
\4-elliptic, the upper ones gradually and strongly
reduced, a few confluent into a pinnatifid leaf-
apex. Upper margin shallowly incised. Juvenile
plants with deeply (bi)pinnatifid, thin pinnules,
sterile; but in transitional cases leaves may be
fertile in the upper part and bear sterile pinnules
of the juvenile shape in the basal part. Veins
immersed free, once or twice forked. Sori inter-
rupted. Spores pale brown, trilete, smooth, c.
22-26 wu.
Distr. Mascarenes, Ceylon, Assam,
China to Malesia, Queensland, and Hawaii.
Indo-
KEY TO THE VARIETIES
1. Rhizome delicate, less than 1 mm @
2. var. delicatula
1. Rhizome of full-grown plants (1 4.—-)2-3 mm og.
2. Indusium with strongly concave base, sub-
hippocrepiform; most sori uninerval and
round, occasionally some binerval and
crescent-shaped; margin regularly incised,
the incisions reaching down almost twice the
distance from receptacle to margin; most
lobes rounded: sporangia at full maturity
strongly laterally spreading; petiole very short
1. var. sessilis
2. Indusium with weakly concave, straight, or
somewhat convex base, or, if more strongly
concave, the sori almost marginal; sori uni-
to plurinerval.
3. At least the broader basal lobes of fully
fertile pinnules truncate; most or all
sori straight and plurinerval (except in
incomplete fertile pinnules); most incisions
not reaching the level of the receptacle.
238 FLORA MALESIANA [ser. IL, volwie
eRe 2 © 2
ee)
Fig. 38. Lindsaea werneri ROSENSTOCK. Leaf, = 4% (SURBECK s.n., L).—Fig. 39. L. versteegii (CHRIST)
v.A.v.R. Pinnule and part of rachis, x 24% (VERSTEEG 1222).— Fig. 40. L. rosenstockii BRAUSE. Part of a
pinna, x 1% (WomersLEy 8346).—Fig. 41. L. microstegia CopEL. Basal part of lamina, x 1%, pinnule,
x 1% (DARBYSHIRE 275).—Fig. 42. L. merrillii CopeL. ssp. merrillii. Pinnule, < 21% (ELMER 13488).—
Fig. 43-45. L. repens (Bory) THWaAITEs.— Fig. 43. var. pectinata (BL.) METT. ex KuHN. Pinnule, x 1%
(BACKER 530).— Fig. 44. var. submarginalis KRAMER. Pinnule, x 2 (SURBECK 946).— Fig. 45. var. sessilis
(CopEL.) KRAMER. Pinnules of two forms, above, x 3 (NGF 8493), below, x 2 (BRASs 23394). — Fig. 46.
L. capillacea Curist. Part of a lamina, * 4 (ELMER 17635).—Fig. 47. L. fissa CopEL. Part of a lamina,
x 2% (MERRILL 9527).
April 1971]
LINDSAEA-GROUP (Kramer)
239
4. Sori very close to the margin, the sporangia
at full maturity often adaxially visible;
pinnules 3-4 times as long as wide
4. var. submarginalis
4. Sori more distinctly intramarginal, the
sporangia never adaxially visible; pinnules
21-3 times as long as wide. 3. var. pectinata
3. Lobes narrowed-rounded, irregular, with
nearly marginal, mostly uni- or binerval
+ round sori; incisions mostly reaching 3
or more times the distance from margin to
receptacle; petiole often well-developed.
5. var. pseudohemiptera
1. var. sessilis (COPELAND) KRAMER, Blumea 15
(1968) 568.—L. sessilis COPELAND.—L. foersteri
Rosenstock.—L. longa COPELAND.—L. cultri-
pinna COPELAND.—Fig. 45.
Rhizome c. 2 mm @; scales honey-coloured to
medium brown, to over 20-seriate at the broadened
base but usually narrower, to 34% by 1 mm.
Petioles to 5 cm long but usually much shorter,
less than 1 cm. Lamina 20 by 2 to 70 by 6 cm,
with c. 40-80 pinnules to a side, rather suddenly,
shortly, and strongly narrowed at both ends.
Pinnules sessile, spreading, or slightly ascending
or the basal ones somewhat falcately decurved,
15 by 4 to 28 by 8 mm, 3-4 times as long as
wide, rarely less. Margins little convex except
if pinnules falcate, the outer margin rounded,
subtruncate, or virtually absent. Colour mostly
dark green when dry. Incisions of upper/outer
margin 44 to 144 mm deep, occasionally deeper,
mostly progressively deeper from base to apex,
reaching almost to the level of the receptacle
to considerably beyond; lobes regular, rounded
or narrowed-rounded, often 1 mm wide. Basal
pinnules reduced, often decurved, sterile, not rarely
deeply pinnatifid. Veins single or rarely paired
in the lobes. Sori on one, or on two connivent
vein-ends (rarely a few on two more divergent
vein-ends, then more elongate and the lobe sub-
truncate), roundish, distinctly intramarginal even
in more deeply incised pinnules; indusium with
concave base, reniform or subhippocrepiform,
pale, entire, 0.4-0.8 mm long, 0.2-0.3 mm wide,
not reaching the extremity of its lobe by its own
width or more.
Distr. Malesia: Borneo (Sabah: Mt Kinabalu),
Philippines (Palawan), Celebes, S. Moluccas, Aru
and KeiIs., New Guinea, Admiralty Is., Solomon
Is., Santa Cruz Is.; Western Polynesia.
Ecol. On trees, rarely epilithic, in moist for-
ests, from sea-level up to c. 1800 m.
Note. The specimens from Mt Kinabalu have
relatively shorter, less incised pinnules. In the W.
part of its range there are some transitions to
var. pectinata; there is probably some hybridi-
zation.
2. var. delicatula (CHRIST) KRAMER, Blumea 15
(1968) 569.— Davallia delicatula CHRIst.—L. sagin-
cola WAGNER & GRETHER.
Rhizome slender, c. 14—3/, mm g; scales honey-
coloured, to 4 by % mm, to c. 16-seriate at the
base. Petioles to 14% cm long, brownish and pale-
angled. Lamina c. 12-25 by 2-3 cm, with up to c.
27 pinnules to a side; rachis adaxially shallowly
grooved. Pinnules spreading or slightly ascending,
mostly pale green when dry, the larger ones 12 by
5 to 15 by 7 mm, 2-3 times as long as wide, narrow-
ed from the base to the narrowed-rounded or
subacute apex; upper and lower margins almost
straight, a distinct outer margin not developed.
Upper margin crenately incised, the lobes round-
ed, often with irregular margin, mostly broader
than long; incisions 5-6 per pinnule, 44-1 mm
deep, rarely deeper, reaching to the level of the
receptacle or beyond. Veins lax, once forked or
simple. Sori uni- or binerval, roundish to oblong;
indusium pale, entire, 44-244 mm long, c. 0.4mm
wide, not reaching the margin by its width or
more.
Distr. Malesia: Celebes, Territory of New
Guinea, Admiralty Is. (Manus, Los Negros),
New Ireland; 1 coll. from each locality, 2 from New
Guinea.
Ecol. Epiphytic on sago palms, in swamps at
sea-level.
3. var. pectinata (BLUME) MeTT. ex KUHN in
Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 277.—
L. pectinata BLUME.—L. hymenophylloides BLUME.
—L. bantamensis BLUuME.—L. boryana (PRESL)
BRAUSE.—L. pectinata BLUME f. dimorpha ROSEN-
sTOCK.—L. repens BLUME f. truncatiloba ROSEN-
sTOCK.—L. repens BLUME var. hemiptera (BORY)
y.A.v.R.—L. repens BLUME var. intermedia
Curist.—L. macraeana (HOOKER & ARNOTT)
CopELAND of Ho.ttrum, Rev. FI. Mal. 2 (1954)
324; COPELAND, Fern FI. Philip. 1 (1958) 101,
etc., at least in part.—Fig. 43.
Rhizome 1%-2%, mostly 2 mm g, eventually
dark brown and devoid of scales; scales medium
brown, to 34% by % mm, to 16-seriate at the
base. Petiole usually 2-5, sometimes up to 8
cm long. Lamina c. 30-80 by 244-5 cm; rachis
usually adaxially sulcate only near the apex.
Pinnules c. 40-100 to a side, spreading or, especi-
ally the basal ones, falcately decurved, less often
somewhat ascending, variable in shape, subtrape-
zoidal, decurved-ligulate, or 14-elliptic, the apex
broadly rounded, subtruncate, narrowed-rounded,
or subacute, usually the lower margin almost
straight and the lower outward increasingly
convex, but sometimes the opposite, occasion-
ally chartaceous; larger pinnules 12 by 5 to 27 by
9 mm, 214-3 times as long as wide, but mostly
less than 3 times. Upper/outer margin with up
to 12 incisions, these to 1 mm deep, the inner
ones shallow, reaching at the most to the level
of the receptacle, the outer ones deeper; or some-
times the pinnules with only vestigial or rarely
without any incisions. Lobes truncate, even most
or all small ones, their margin sometimes a little
irregular. Sori straight, bi- to plurinerval, inter-
rupted by the incisions of the margin, even inter-
rupted if these do not reach their level, or in entire
pinnules, but seemingly confluent with age if
the interruptions are short. Indusium entire or
240
sinuate, 0.3-0.5 mm wide, not reaching the
margin by a little more than its width to almost
reaching it, strongly reflexed and often quite
concealed at maturity.
Distr. Assam, Ceylon, Thailand, Indo-China,
in Malesia: Malay Peninsula, widespread and
often common on the Greater Sunda Is., Men-
tawei Is., Bawean, Philippines.
Ecol. Epiphytic in moist forests, 600-1500 m,
sporadically to sea-level.
Note. In Mindanao, Luzon, and Leyte an
aberrant form is found, with little or not incised
pinnules 9 by 5 to 15 by 8 mm, 2 to less than 114
times as long as wide. It may be distinguished as
f. angusta (COPELAND) KRAMER, Blumea 15
(1968) 568 (L. angusta COPELAND).
4. var. submarginalis KRAMER, Blumea 15 (1968)
569.—Type: BARTLETT 6603, Karo Plateau, Su-
matra (L; dupl. in GH, MICH).—Fig. 44.
Rhizome scales to 3% by % mm, medium brown,
to c. 16-seriate at base. Lamina often basally
little reduced, with a well-developed petiole of
several cm. Pinnules often ascending and falcately
upcurved, narrowed-rounded or subacute at apex,
without an outer margin, the largest 20 by 5 to
26 by 6 mm, c. 3% to over 4 times as long as wide,
mostly dark when dry. Upper margin shallowly
and regularly incised, the incisions to % mm
deep, up to twice the distance from margin to
receptacle; lobes truncate, the smaller ones trun-
cate-rounded. Sori bi- to quadrinerval, straight
with concave extremities or the shorter ones
basally quite concave. Indusium entire, c. 0.3
mm wide, practically equaling the margin, strong-
ly reflexed and quite concealed at maturity.
Distr. Malesia: Sumatra (7 coll.; some doubt-
ful ones).
Ecol. Epiphytic in mountain forests, c. 1200—
1800 m.
Note. Not quite sharply distinct from var.
pectinata.
5. var. pseudohemiptera v.A.v.R. Bull. Jard. Bot.
Btzg III, 2 (1920) 157, f. C.—Lectotype: BUN-
NEMEWER 5419, Mt Merapi, Sumatra (BO; dupl.
inves SING. WL US):
Rhizome dark, 14%-2 mm g; scales medium to
rather dark brown, to 24% mm long, to c. 13-
seriate at base. Petioles a few cm or up to 10
(exceptionally to 15) cm long. Lamina c. 30-50
by 3-4 cm, with c. 40-70 pinnules to a side.
Pinnules spreading, slightly ascending or slightly
decurved, often trapezoidal, narrowed to the apex
or not, there subacute to truncate, mostly very
dark when dry, the larger ones 20 by 5 to 30 by
6 mm, 34 to over 5 times as long as wide; incisi-
ons of upper/outer margin rather regular, shallow,
c. Y% mm deep, or, especially towards the apex
of the pinnules, deeper, oblique, to 4 mm deep
and then irregular and reaching to the middle
of the pinnule, the pinnule-apex then sometimes
protracted, with 1 or more pinnatifid prolonga-
tions. Lobes ligulate, narrowed-rounded, uni- to
FLORA MALESIANA
[ser. II, vol. 1%
quadrinerval, with rather broad sinuses. Veins
rather irregularly spaced. Sori uni- to quadriner-
val, often on strongly connivent vein-ends, nearly
marginal; indusium c. 4%—2 mm long, 0.2 mm wide,
pale, subentire, with concave base, not reaching
the margin by its width or less, strongly reflexed
and usually entirely concealed at maturity by
the sporangia that spread in all directions, mostly
somewhat beyond the edges of the soral lobes.
Distr. Malesia: Sumatra, Borneo (Sabah:
Mt Kinabalu, efc.).
Ecol. Epiphytic in mountain forests,
1800 m.
Notes. Several juvenile plants from N. Borneo
with rigid bipinnatifid pinnules may belong to
this variety.
The very irregularly and deeply cleft pinnules, as
described and figured by VAN ALDERWERELT VAN
ROSENBURGH, Occur in the two syntypes and some
other specimens, but usually the incisions are
much less deep and more regular.
This variety may be a distinct species; the
scales are darker than in the other varicties.
1500-
47. Lindsaea apoensis COPELAND in Perkins,
Fragm. Fl. Philip. (1904) 182, pl. 4 f. A; Fern
F]. Philip. 1 (1958) 103.—Type: CoPELAND 1181,
Mt Apo, Mindanao (n.v.); paratype: DE VoRE &
Hoover 365 (dupl. in K, S-PA).
L. havicei COPELAND, Philip. J. Sc. 1 (1906)
Suppl. I1,149; Fern Fl. Philip. 1 (1958) 103.—Type:
COPELAND 1758, Mt Balabac, Mindanao (MICH;
dupl. in B, P).—Fig. 50.
Rhizome \ong-scandent, brown, 1-14% mm g;
scales deciduous, dorsally more persistent, go!den-
brown, to c. 24% mm long, elongate-ovate, to c.
20-seriate at base, with a very short uniseriate
apex. Leaves 1-3 cm apart, issuing at a small
angle but usually curved to right angles with the
rhizome; petioles stramineous to pale brown,
rarely darker and then pale-margined, quadrangu-
lar, laterally + sulcate, 5-20 cm long, shorter
than the lamina. Lamina linear, 20-40 cm long,
3-6 1% cm wide, shortly to very shortly acuminate,
shortly narrowed or more often abrupt at the base,
with 20-50 pinnules to a side; rachis stramineous,
adaxially shallowly sulcate, or flat at the base.
Some basal pinnules inserted below the edges
of the adaxial face of the rachis, spreading or
slightly ascending, rather variable in shape,
narrowly triangular to subtrapezoidal, narrowed
from base to apex and acute or less narrowed
and with truncate apex, the apex occasionally
caudate-protracted; texture herbaceous, colour
medium to dark green when dry. Larger pinnules
2-3 cm long, 4-7 mm wide, 4-5 times as long as
wide; lower margin mostly at least at the base
concave, upper margin straight or + convex,
with c. 6-8 major incisions, these slightly oblique,
almost parallel, towards the apex of the pinnule
progressively deeper, the inner ones 1-2 mm deep,
reaching to 1/,-'’% (rarely deeper and reaching
beyond the middle), the outer ones at least 2 mm
deep, reaching to the middle or far beyond;
inner lobes often more shallowly incised again;
April 1971]
lobes little marrowed, almost parallel-sided,
rounded at apex, uni- or binerval. Pinnule-apex
sometimes protracted into a pinnatifid cauda.
Lower pinnules not remote and reduced or very
few more remote and/or slightly reduced; some
upper pinnules gradually reduced, confluent into
a pinnatifid leaf-apex. Juvenile plants without the
deeply incised pinnules found in L. repens.
Veins immersed, evident, once or twice forked, or
the outer ones simple. Sori bi- or more often
uninerval; indusium entire, if binerval elongate
and with concave base, if uninerval reniform to
hippocrepiform, 44-1 mm long, 0.6 mm wide,
falling short of the margin by 14-1 times its width,
strongly reflexed, often shrivelled and not visible
at maturity. Spores pale yellowish, hyaline, tri-
lete, smooth, c. 23 wu.
Distr. Malesia: Philippines (Luzon, Negros,
Leyte, Mindanao).
Ecol. Epiphytic in moist mountain forests,
1200-2100 m.
Note. The basally truncate or almost truncate
lamina and the pinnules incised to the middle at
least near the apex are distinctive.
48. Lindsaea merrillii COPELAND in Perkins, Fragm.
Fl, Philip. (1904) 181; TaAGAwa, Act. Phytotax.
Geobot. 6 (1937) 33, f. 2 E, F; CopELAND, Fern
Fl. Philip. 1 (1958) 102.—Type: MerrILv 1774,
Baco R., Mindoro (MICH; dupl. in B, GH, K,
US).
Odontoloma repens auct. non (BORY) PRESL;
J. SmitH, Hist. Fil. (1875) pl. 18 d.—Fig. 42.
Rhizome \ong-scandent, dark brown, 1-2 mm 9;
scales deciduous, dorsally more persistent, Jemon-
coloured to light brown, elongate-triangular,
to 3% by 1 mm, up to c. 20-seriate at base, with
a very short uniseriate apex. Leaves 14 cm
apart; petioles virtually absent to 8 cm long,
stramineous with dark base or brown and then
often pale-angled, quadrangular, only if very
short sometimes subterete, very much shorter than
the lamina. Lamina linear, 25-50 cm long, 2.5-4.2
cm wide, moderately to strongly narrowed at base,
acuminate, with c. 35-60 pinnules to a side;
rachis stramineous, sharply quadrangular, adax-
ially flat or shallowly sulcate, abaxially usually flat.
Pinnules spreading or somewhat ascending,
elongate-triangular or less often semi-ovate,
herbaceous, medium to dark green when dry,
the larger ones 11-22 mm long, 4-6 mm wide,
2-41, times as long as wide, narrowed from base
to apex, the apex subacute or less often narrowed-
rounded or rounded; lower margin straight, out-
ward convex; upper margin approximately
straight, regularly dentate if sterile, more irregular-
ly crenate or dentate if fertile, the incisions 44-1 mm
deep, not reaching down to the middle except
sometimes near the apex; teeth narrowed from
base to apex, acuteif sterile, obtuse and erose to
acute and often very irregularly erose if fertile,
the middle more protruding. Basal pinnules
reduced and remote, often auriculiform and a
few mm long, sterile; upper pinnules gradually and
LINDSAEA-GROUP (Kramer)
241
strongly reduced, confluent into a _ pinnatifid
leaf-apex. Veins immersed, evident, once forked,
less often twice forked or simple, 1 or 2 running
to each lobe, their ends c. 1 mm apart. Sori uni-
or binerval, single on the teeth; indusium pale,
subentire, semicircular or reniform to oblong,
144-1 mm long, c. 0.3 mm wide, with convex to
slightly concave base, not reaching the margin by
about its width or more, + reflexed and some-
times concealed at maturity. Spores pale brown,
trilete, smooth, c. 22 w. Juvenile plants have deeply
divided pinnules, but less regularly so than in L.
repens.
Distr. Malesia: Philippines (Samar, Polillo,
Batanes, Leyte, Camiguin, Mindanao, Mindoro,
Luzon); these plants belong to ssp. merrillii.
Another subspecies in the Ryu Kyu Is. and Tai-
wan.
Ecol. Epiphytic on palms and other trees,
from sea-level up to 700 m.
Note. Davallia boryana PRESL was described
from a specimen from Sorzogon belonging to
this species; but PRESL cited Dicksonia repens asa
synonym, and D. boryana must therefore be re-
garded as a new name for that species.
49. Lindsaea carvifolia KRAMER, Blumea 15 (1968)
569.—Type: BEGUIN 1116, Ternate (BO; dupl.
in L).—Fig. 48.
Rhizome long-scandent, brown, 1-2 mm 4g,
deciduously scaly; scales honey- to fawn-coloured,
narrowly triangular, to 2 mm long, to c. 16-
seriate at the base, scarcely uniseriate at the apex.
Leaves remote, c. 1-5 cm apart; petioles strami-
neous, or with brown base, adaxially flattened or
sulcate, abaxially convex, rounded to obtusely
bi-angular, extremely short to 4 cm long. Lamina
linear, 20-50 cm long, 1144-4 cm wide, tapering at
both ends, with c. 50-80 pinnules to a side;
rachis adaxially flattened or sulcate, abaxially
bi-angular to broadly and shallowly sulcate.
Pinnules herbaceous to chartaceous, dark green
when dry, spreading or somewhat ascending, at
least in the basal part of the lamina inserted
somewhat below the level of the adaxial rachis
face, often more remote in the lower part of the
lamina, the larger ones 6 by 2 to 20 by 6 mm, 3
times as long as wide, almost evenly narrowed
from base to apex, subtriangular; upper margin
deeply incised, the primary segments 5-7, all
but the smallest once, the largest twice bifid;
lobes little divergent, linear to capillary, (0.3—)
0.5-1 mm wide, often seemingly narrower when
dry because of the revolute margins, sometimes
somewhat broadened at the sorus, the apex round-
ed; wing connecting the primary segments c.
14 mm wide at the narrowest points. Veins im-
mersed, evident, single, rarely paired in larger,
scarcely bifid lobes. Sori uninerval or very rarely
binerval, one per ultimate lobe, mostly roundish;
indusium pale, with straight or concave base,
flattened-elliptic to hippocrepiform, 0.3—-0.5(0.7)
mm long, c. 0.2 mm wide, almost reaching the
margin to not reaching it by more than its width,
reflexed and + concealed at maturity, the spo-
242 FLORA MALESIANA [ser. II, vol. 1
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Fig. 48. Lindsaea carvifolia KRAMER. Two laminas, < 2/; (ANDERSON S 20179).— Fig. 49. L. glandulifera
v.A.v.R. Leaf, x 2/; (VAN HEURN 196).— Fig. 50. L. apoensis CopeL. Lamina, x ?/, (EDANO 5229).
April 1971]
rangia often spreading beyond the edges of the
segment. Spores pale brown, trilete, smooth, c.
PIS) [Ve
Distr. Malesia: Borneo, Celebes (?), Moluccas
(Morotai, Halmahera, Ternate).
Ecol. Epiphytic or rarely terrestrial, in moun-
tain forests, 500-1700 m.
Note. The single collection from Celebes
(WARBURG 16527, B) may belong to L. rosen-
stockii, but the occurrence of that species in Ce-
lebes is unlikely.
50. Lindsaea fissa COPELAND, Philip. J. Sc. 38
(1929) 143, pl. 1; Fern Fl. Philip. 1 (1958) 105.—
Type: MERRILL 9527, Mt Capoas, Palawan
(MICH; dupl. in BM, BRI, GH, L, P, US).
L. hymenophylloides auct. non BLUME; COPE-
LAND, Fern FI. Philip. 1 (1958) 105.—Fig. 47.
Rhizome long-scandent, 34-2 mm g, dark
brown, deciduously scaly; scales dorsally more
persistent, golden brown to fawn-coloured, tri-
angular, to 3 by 1 mm, to c. 22-seriate at the
base, there the cell partitions sometimes bulging
beyond the margin, the apex very shortly uni-
seriate. Leaves rather remote, %—4 cm apart;
petioles dark or reddish brown, or paler above,
abaxially terete at the base, upward gradually
bi-angular, laterally often sulcate, adaxially flat-
tened to shallowly sulcate, 1—S(—10) cm long,
much shorter than the lamina. Lamina linear,
tapering at both ends, (10—-)15-40 cm long,
(1 %—-)2-3 cm wide, with c. 20-60 pinnules to a
side; rachis stramineous, quadrangular, adaxially
at least upward sulcate. Pinnules thinly herbaceous,
dark green to olivaceous or blackish when dry,
spreading or falcately deflexed, rarely ascending,
rather close or contiguous, approximately %-
elliptic in outline, the larger ones (7—)10—-15
by 3-5 mm, 2-3 times as long as wide; upper
margin outward gradually more convex, deeply
incised, larger pinnules consisting of 4-5 primary
segments, the inner, larger ones usually forked
(occasionally twice forked); ultimate segments
divergent, linear-cuneate, %—2 mm wide at the
apex, narrowed just below the sorus, little or not
narrowed to the base, there connected by wings
of %4-% mm; apex of segments truncate-sinuate
or -erose, not rarely with two slightly larger
lateral protuberances, sometimes also with 1-2
smaller ones between them. Few upper pinnules
reduced and confluent into a pinnatifid leaf-
apex; some to many basal ones remote and grad-
ually reduced, usually inserted just below the
lateral ridges of the adaxial rachis face. Veins
immersed, evident, single or paired in the ulti-
mate lobes. Sori uni- or not rarely binerval;
indusium thin, pale to brown, subentire, %4—-2 mm
long, 0.4 mm wide, at least laterally convex at
the base, not reaching the margin by about its
width to almost reaching it, not reflexed at matu-
LINDSAEA-GROUP (Kramer)
243
rity. Spores very pale brown, trilete, smooth, c.
PUSS [ie
Distr. Malesia: Philippines (Luzon, Palawan,
Panay, Negros, Mindoro, Biliran, Mindanao).
Ecol. Epiphytic in humid mountain forests,
860-1400 m; few ecological notes given.
51. Lindsaea capillacea Curist, Bull. Herb.
Boiss. I, 6 (1898) 144, pl. 4 f. 7; COPELAND,
Fern Fl. Philip. 1 (1958) 106.—Type: Loner s.n.,
Mt Maqguiling, Luzon (P; dupl. in B, K,
S-PA).—Fig. 46.
Rhizome long-scandent, dark brown, % mm @,
deciduously scaly; scales golden brown, narrowly
triangular, to 144 mm long, up to 9-seriate at
base, scarcely uniseriate at apex. Leaves 1-2 cm
apart; petioles dark brown, lustrous, adaxially
flattened or sulcate, abaxially terete at base,
obtusely or acutely bi-angular above, 1—5 cm
long, much shorter than the lamina. Lamina
linear, 4-15 cm long, 1-2% cm wide, narrowed
at both ends; rachis brown at the base, upward
stramineous, adaxially often quite stramineous,
at least above sulcate, abaxially bi-angular.
Pinnules c. 15-60 to a side, thinly herbaceous,
dark green when dry, spreading or slightly
ascending or the basal ones deflexed, the larger
ones 4-15 mm long, 2-4 mm wide, 2 to almost 4
times as long as wide, %-elliptic to almost tri-
angular in outline, the upper margin straight or
more often outward increasingly convex, deeply
incised; larger pinnules with 2—5 primary lobes,
the largest one or two forked; segments 0.4—0.8
mm wide near the apex, narrowed to the base,
there about half as wide, joined by wings of 0.1—
0.3 mm, + divergent; fertile segments at the
apex bi-apiculate, with two lateral horn-like
projections of 0.3-0.4 mm, the margin between
them subentire, erose, or with similar but shorter
projections. Upper pinnules reduced, confluent into
a pinnatifid leaf-apex; lower pinnules more re-
mote, scarcely to rather strongly reduced. Veins
immersed, evident, single in the lobes. Sori
strictly uninerval; indusium pale, brownish,
subentire, %4-1 mm long, % mm wide, with
straight or'slightly concave, laterally convex base,
not quite reaching the margin, scarcely reflexed
at maturity. Spores pale brown, trilete, smooth,
(Drs) [Pe
Distr. Malesia: Philippines (Luzon, c. 25
coll.).
Ecol. Epiphytic or occasionally terrestrial,
in moist mountain forests, 600—2200 m.
Note. This might be taken for a small form of
L. fissa, and small specimens of that species may
resemble L. capillacea, but they lack the well-
developed latero-apical horns of the segments.
The fact that in Luzon the two species occur
together shows that they are not geographical
variants.
244 FLORA MALESIANA [ser. II, volar?
11. Section Pseudolancea
KRAMER, Blumea 15 (1968) 563.
Type and sole species: Vittaria parasitica ROXB. ex GRIFF. = L. parasitica (ROXB. HIERON.
L. parasitica is probably related to L. oblanceolata in sect. Odontoloma. But L. parasitica has
bipinnate fronds on fully developed plants, whereas the fronds of all species of sect. Odontoloma
are simply pinnate; the separation of L. parasitica thus leaves a more clearly natural sect.
Odontoloma. It seems probable that the bipinnate condition has arisen on more than one
evolutionary line within subg. Odontoloma; this subject will be further discussed elsewhere.
52. Lindsaea parasitica (ROXBURGH ex GRIFFITH)
HIERON. Hedwigia 62 (1920) 14; KRAMER, Blu-
mea 15 (1968) 570.—Vittaria parasitica ROXBURGH
ex GRIFFITH, Calc. J. Nat. Hist. 4 (1844) 510.—
Type: ROXBURGH s.n., Prince of Wales’ Island
(Penang) (n.¥.).
L. scandens HOOKER, Sp. Fil. 1 (1846) 205, pl.
63 B; Bepp. Ferns Brit. Ind. 2 (1868) pl. 298;
Ho.ttum, Rev. Fl. Mal. 2 (1954) 325, f. 186;
COPELAND, Fern FI. Philip. 1 (1958) 105.—Lecto-
type: CUMING 405, Luzon (K).
L. lancea auct. non (L.) BEDD. of various authors,
as to Asiatic specimens, in part.
Rhizome \ong-scandent, often knotted, fuscous,
2-3 mm g; scales castaneous, to 4 by 1 mm, tri-
angular, to c. 25-seriate at base, the uniseriate apex
very short. Leaves one to many cm apart; petioles
stramineous or with dark base, stout, up to 3 mm
@ at base, adaxially with a very narrow groove
across which its borders often touch, abaxially
terete, c. 7-30 cm long in bipinnate leaves, much
shorter than the lamina, almost wanting to c.
5 cm long in simply pinnate leaves. Lamina sim-
ply pinnate or bipinnate; if simply pinnate c.
20-35 cm long, 244-5 cm wide, very narrowly
oblong, gradually narrowed at base, shortly and
abruptly narrowed at apex; if bipinnate c. 35-50
cm long, c. 20-35 cm wide, oblong, c. 1% times as
long as wide, not narrowed at base, with 1-6
primary pinnae to a side and a conform terminal
one; primary rachis of bipinnate leaves similar to
the petiole but the adaxial groove mostly more
open, with flat or convex bottom. Primary pinnae
ascending or spreading, usually alternate, remote,
up to 8 cm apart, subsessile, 15-25 by 2-5 cm,
acute to shortly acuminate, with c. 20-35 pinnules
to a side; simply pinnate laminas similar but
narrowed at the base, usually longer and less
narrowed at the apex. Pinnule-bearing rachises
abaxially terete, adaxially laterally sharply mar-
ginate, with a convex-bottomed groove. Pinnules
firmly herbaceous, drying medium to dark green
or blackish, described as glossy by some col-
lectors, close, regularly spaced, usually contiguous,
spreading or slightly ascending, the basal ones
sometimes deflexed, dimidiate-elliptic to ligulate
or slightly subfalcately decurved, little or not
narrowed to the apex, apically + rounded, a
separate outer margin not developed, sometimes
an angle between lower and outer part of upper
margin; larger pinnules 11 by 6 to 20 by 8 mm,
2-2% times as long as wide. Margins entire,
shallowly crenate in sterile pinnules, more deeply
crenate in juvenile plants. Very few upper pinnules
reduced in simply pinnate leaves, more numerous
and more strongly reduced in bipinnate ones,
none or | or 2 connected with the asymmetrically
triangular to broadly lanceolate terminal pinnule
(segment). Veins immersed, usually evident, often
visible as striations, free, mostly twice forked.
Sori continuous, occupying all vein-ends; in-
dusium pale, entire, 0.3-—0.4 mm wide, not
reaching the margin by less than its width to
nearly reaching it, strongly reflexed and often
concealed at maturity. Spores yellowish, trilete,
smooth, c. 22 wu.
Distr. Peninsular Thailand; Malesia: Malay
Peninsula, Singapore, Sumatra, Borneo.
Ecol. Epiphytic or occasionally terrestrial,
from sea-level up to c. 1100 m.
Notes. The lectotype of L. scandens, CUMING
405, was said to have come from Luzon; a syn-
type is labelled ‘Leyte’. The identity of the latter
is doubtful; and as L. parasitica is not otherwise
known from the Philippines it seems more likely
that CUMING’s plant, like others of his collection,
was mislabelled and came actually from the Malay
Peninsula.
Simply pinnate specimens of L. parasitica may
be very similar to L. oblanceolata; the two are
probably related. In L. oblanceolata the basal
pinnules are inserted below the adaxial face of
the rachis, in L. parasitica they are borne on the
edges of the rachis groove.
Another species with strikingly similar foliage
but different rhizome and scales is 32. L. dory-
Phora. For additional characters serving to distin-
guish detached leaves see under that species.
12. Section Lindsaenium
(FEE) KrAMerR, Blumea 15 (1968) 563.—Lindsaenium (or Lindsaynium) FEE, Mém. Soc. Mus.
Hist. Nat. Strasb. 4 (1850) 201; Gen. Fil. (1852) 333.
Type species: Lindsaea rigida J. SmMiTH in Hooker.
Distr. Malay Peninsula to Micronesia and Tahiti; all species occur in Malesia.
Taxon. A very natural group of epiphytic species with long-creeping rhizomes, at least some bi-
pinnate leaves, and interrupted sori. The first four and the last three species are more closely inter-
related.
April 1971]
LINDSAEA-GROUP (Kramer)
245
53. Lindsaea rigida J. SMITH in Hooker, Sp. Fil.
1 (1846) 217, pl. 63 A; Hottrum, Gard. Bull.
S. S. 5 (1930) 65; COPELAND, Philip. J. Sc. 78
(1949) 19; Hottrum, Rev. Fl. Mal. 2 (1954)
330; KRAMER, Blumea 15 (1968) 570.—Lina-
saenium rigidum (J. SMITH) FEE, Mém. Soc. Mus.
Hist. Nat. Strasb. 4 (1850) 201; Gen. Fil. (1852)
333, pl. 27 bis.—Lectotype: Loss s.n., Mt Ophir,
Johore (‘Malacca’) (K, 2 sh.; dupl. in E).
L. longissima Curist, Ann. Jard. Bot. Btzg
II, 5 (1905) 131.—Type: JAHERI (exp. NIEUWEN-
HUIS) 958, Lelibulan Teputsy, Kalimantan,
Borneo (P; dupl. in BO).
L. monosora COPELAND in Elmer, Leafl. Philip.
Bot. 2 (1908) 398; Fern FI. Philip. 1 (1958) 109;
non ROSENSTOCK (1912).—Type: ELMER 10077,
Cuernos, Negros (MICH; dupl. in B, BM, BO,
BRIaE GH BG KL aPs UsUS:) WwW. 2):
L. diplosora v.A.v.R. Bull. Jard. Bot. Btzg II,
16 (1914) 21.—Type: MatTTHEw, 523, Mt Sing-
galang, Sumatra (BO; dupl. in K).
L. triplosora y.A.v.R. Bull. Jard. Bot. Btzg
II, 16 (1914) 21.—Type: MATTHEW 686, Mt Sago,
Sumatra (BO; dupl. in K).
L. rigida J. SmitH f. acutata v.A.v.R. Handb.
Suppl. (1917) 210.—Type: not designated; no
specimen so annotated seen from BO.
L. sepikensis BRAUSE, Bot. Jahrb. 56 (1920) 131;
COPELAND, Philip. J. Sc. 78 (1949) 22.—Type:
LEDERMANN 9423 p.p., Etappenberg, Sepik R.
region, Terr. of New Guinea (B; dupl. in BM).
L. diplosora y.A.v.R. var. acrosora C. CHR. Gard.
Bull. S. S. 7 (1934) 235.—Type: MJOBERG 94,
Mt Tibang, Sarawak (BM; dupl. in S-PA).
Rhizome long-scandent, 144-3 mm g; scales
golden brown, narrowly triangular, to 3 by 4% mm,
to c. 10-seriate at base, with a short uniseriate
apex. Leaves usually 24% cm or more apart,
rarely closer; petioles issuing almost at right angles
from the rhizome, olivaceous- to dark brown,
often with elongate, paler protuberances, abaxially
terete, adaxially at least above flattened to sul-
cate, 10-40 cm long, about equaling to much
longer than the lamina. Lamina oblong, 20-50
cm long, bipinnate, with 1-7 pairs of usually sub-
opposite pinnae and a conform terminal one;
primary rachis abaxially terete or upward obtusely
bi-angular, adaxially broadly and_ shallowly
sulcate, with obtuse lateral ridges. Pinnae not
close, often 6-8 cm apart, ascending, linear, sessile,
10-25 cm long, 10-20 mm wide, gradually taper-
ing from the lower third or the middle upward;
secondary rachises brown, abaxially sharply bi-
angular except for a short terete basal portion,
the angles often paler, the face between them flat
or concave. Upper pinnae of plurijugate leaves
slightly shortened. Pinnules herbaceous to rigidly
coriaceous, mostly dark olivaceous to blackish
when dry, usually spreading, c. 25-45 to a side,
their width apart or less but not contiguous,
subtrapeziform, obliquely ovate, or 14-elliptic,
the larger ones 7 by 3 to 12 by 6 mm, mostly
twice (to 2% times) as long as wide, narrowed
(sometimes very little) from base to apex; margins
pale-sclerotic, especially in rigid pinnules; lower
margin straight or distally convex, upper margin
distally gradually convex, a distinct outer margin
usually not developed. Upper margin of sterile
pinnules (bi)crenate to dentate, of fertile pinnules
crenate with incisions to 1 mm deep, less in rigid
pinnules, the lobes subtruncate. Veins immersed
and evident in herbaceous, + elevated on both
sides in rigid pinnules, simple or once, rarely
twice, forked, free, connivent, or with one anasto-
mosis between upper and lower margin, the anas-
tomoses irregular, often involving only part of
the veins and mostly not found in all pinnules.
Upper pinnules very gradually and strongly re-
duced, some denticuliform ones confluent into a
narrow, lobed pinna-apex. Sori 1-4 per pinnule,
1 per lobe, on the outer lobes; larger pinnules
only partly soriferous, the inner part of the upper
margin sterile; very often only the outermost
sorus present, on 2-6 veins, to 5 mm long; inner
sori shorter. Receptacle straight or with convex
ends; indusium mostly rigid, brownish, subentire
to erose, narrowed at the adnate ends, 15-34 mm
wide, not quite reaching the margin, reflexed or
not at maturity. Spores dark brown, trilete, slightly
pustulate, c. 25 yw.
Distr. Malesia: Malay Peninsula, Sumatra,
Borneo, Celebes, Philippines (Palawan, Negros,
?Luzon), Moluccas (Batjan), Japen, New Guinea;
Ponape; Solomon Is.; Tahiti. ;
Specimens labelled ‘New Hebrides’ possibly,
‘Singapore’ and ‘Java’ probably from elsewhere.
Ecol. Epiphytic or epilithic, or terrestrial,
among mosses, not high above the ground (ac-
cording to HoLttuM); 1000-2000 m, rarely lower.
The most rigid specimens are from higher altitude
in Malaya, Sumatra, and New Guinea.
Note. In spite of a certain variablity this is
a distinctive species, and it is surprising that it
has comparatively many synonyms and is often
misidentified.
54. Lindsaea monocarpa ROSENSTOCK in C. Chr.
Ind. Fil. Suppl. 1 (1913) 49, based on: L. mono-
sora ROSENSTOCK, Nova Guinea 8 (1912) 720,
non COPELAND (1908).—Lectotype: VON ROMER
785, Hellwig Mts, W. New Guinea (L; dupl.
in BO).
In most respects like the rigid, little incised form
of L. rigida. Petiole, primary, and secondary
rachises dark castaneous, abaxially quite terete.
Pinnules subtrapezoidal, the larger ones 10 by 4
mm, very shallowly crenate; veins almost hidden,
occasionally connivent or anastomosing; pinnules
with a single apical sorus.
Distr. Both syntypes, VON ROMER 785 and
1163a, from the same locality.
Ecol. No data.
Note. Perhaps only a form of L. rigida; but
that species is so constant in the characters of
its axes that in the absence of transitions L.
monocarpa is better retained as a species.
55. Lindsaea sarawakensis KRAMER, Blumea 15
(1968) 570.—Type: Ms6BerRG 9, Mt Murud,
Sarawak (P, 3 sh.).
246
FLORA MALESIANA
[ser. Il; vole
Rhizome unknown. Petioles dark brown, shin-
ing, abaxially rounded or upward obtusely bi-
angular. Lamina c. 35-40 cm long, oblong,
bipinnate, with 4-5 pinnae to a side and a con-
form terminal one; primary rachis dark brown,
abaxially rounded or narrowed-rounded. Pinnae
linear, 15-30 cm by 12-15 mm, acute, with c.
55-95 pinnules to a side; secondary rachises
brown at base, otherwise stramineous, abaxially
carinate almost to base. Pinnules herbaceous,
subcontiguous, asymmetrically ligulate, narrowed-
rounded, at apex, the larger ones about 7 by 2 to
2% mm, the upper ones gradually and + strongly
reduced, a few confluent with the small lanceolate
terminal segment. Upper margin of larger pinnules
with 2 or 3 very oblique, acute incisions reaching
down to !/;; lobes rounded. Veins immersed,
not evident, free, simple or once forked. Sori only
in the outer part of the upper margin (in fully
fertile pinnules possibly more extensive), bi- or
rarely uninerval; indusium brownish, c. 0.3
mm wide, not quite reaching the margin. Spores
pale brown, trilete, smooth, c. 20 pw.
Distr. Only known from the type collection.
Ecol. No data.
Note. In the absence of arhizome the taxonomic
position of this species is not quite certain, but
it resembles L. rigida in several characters.
56. Lindsaea regularis ROSENSTOCK, Meded.
Riksherb. 14 (1912) 31.—Type: ELBERT 1789,
Mt Pussuk, Lombok, Lesser Sunda Is. (L; dupl.
in BO, K, SING).—Fig. 52.
Rhizome \ong-scandent, 1144-214 mm 9g; scales
deciduous, fawn-coloured, triangular-acuminate,
to 4 by 2 mm, the base broadened, over 20-
seriate, with often laterally projecting cell-walls,
the apex shortly uniseriate. Leaves c. 2-10 cm
apart; petioles medium brown, often scaly at
the base, terete, or adaxially in the upper part
flattened or subsulcate, 10-25 cm long, about as
long as the lamina. Lamina oblong, bipinnate
(occasionally sterile simply pinnate ones also
present), c. 15-30 cm long, with 3-9 pinnae to
a side and a conform terminal one; primary
rachis terete, adaxially with a distinct but shallow
and very narrow groove that is usually not
clearly continuous with the grooves of the lower
pinnae and occasionally extends to the petiole.
Pinnae ascending or upcurved, sessile, the lower
Ones several times their width apart, the upper
ones closer, linear, c. 10-20 cm long, 11-16 mm
wide, often widest near the middle, gradually and
strongly narrowed in the upper 14 or 13; secondary
rachises abaxially terete at the base, upward
gradually bi-angular, then sulcate, or occasionally
bi-angular to base, sometimes pale-angled.
Pinnules c. 20-50 to a side, herbaceous, mostly
dark olivaceous when dry, regularly spaced,
mostly not contiguous, asymmetrically ovate
to shortly ligulate, 6-10 by 3-4 mm, 2-2'% times
as long as wide, slightly ascending, the lower
margin almost straight, the upper margin distally
convex, a distinct outer margin not developed,
the pinnule-apex rounded or narrowed-rounded.
Upper pinnules gradually and strongly reduced,
several denticuliform ones connected with the
narrow, caudiform pinna-apex. Margins scarcely
sclerotic; upper margin shallowly incised, with
3-5 incisions 4%4—'% mm or even less deep, the
lobes slightly convex to flat; sterile pinnules cre-
nate to subentire. Veins immersed, evident, simple
or once, less often twice forked, free. Sori inter-
rupted, mostly 1 per lobe, even though most
incisions do not reach to the receptacle, shorter
than the lobes, often 3-5 per pinnule, uni- to
quadrinerval; indusium pale, stiff, entire or nearly
so, with straight base and narrowed adnate sides,
mm wide, almost or quite reaching the margin,
little reflexed at maturity. Spores hyaline, trilete,
smooth, c. 22-25 wu.
Distr. Malesia: East Java (2 coll.), Lesser
Sunda Is. (Bali, Lombok, Flores, Timor).
Ecol. Epiphytic in primary forest, 600-1700 m,
on trees and tree-ferns. Apparently uncommon.
57. Lindsaea microstegia COPELAND, Philip. J.
Sc. 6 (1911) Bot. 83; ibid. 78 (1949) 20.—Type:
KING 242, Papua (MICH).
L. pectinata BLUME var. brevipinnula ROSEN-
stock, Hedwigia 56 (1915) 351.—Type: BAMLER
126, Mt Sattelberg, Terr. of New Guinea (B;
dupl. if P).—Fig. 41, 51.
Rhizome long-scandent, brown, I-2 mm @,
the older parts loosely scaly; scales honey-
coloured, triangular to lanceolate with concave
base, there with + protruding cell-walls, to 2 mm
long, to c. 20-seriate at base, the apex scarcely or
not uniseriate. Leaves not close, to c. 10 cm apart;
petioles stramineous to medium brown, adaxially
grooved, abaxially terete, or above obtusely bi-
angular, 8-20 cm long, mostly shorter than the
lamina. Lamina bipinnate, occasionally simply
pinnate fertile leaves also present, rarely the whole
plant with simply pinnate leaves, broadly oblong,
15-60 cm long, with 1-7 pinnae to a side and a
conform, often very large terminal one; primary
rachis stramineous, abaxially often -+- pronounced-
ly bi-angular. Pinnae mostly little ascending,
alternate or the basal ones subopposite, sessile,
not close, 10-20 cm long, 134-3 cm wide, acumi-
nate; secondary rachises abaxially bi-angular and
flat or shallowly sulcate almost to base. Pinnules
c. 20-25 to a side, herbaceous, medium to dark
or olivaceous green when dry, regularly spaced,
close, spreading, approximately trapezoidal or
rarely subligulate or semi-ovate, the larger ones
10-15 by 5-7 mm, 2-2% times as long as wide,
mostly narrowed from base to apex, the apex
narrowed-rounded or subtruncate, outer margin
distinct or broadly rounded into the upper margin;
upper/outer margin sinuate to shallowly crenate,
the incisions less than 1 mm deep, not reaching
to the level of the receptacle (rarely slightly
beyond), the lobes very broadly rounded, 1 4-2
mm wide. One or two basal pinnules slightly
reduced; upper pinnules gradually and strongly
reduced, some denticuliform ones confluent with
the narrow, caudiform terminal segment, or rarely
much less reduced. Pinnules of juvenile plants
April 1971] LINDSAEA-GROUP (Kramer) 247
Baik
Fig. 51. Lindsaea microstegia CopEL. Apex of plant, x 1 (Brass 25846).—Fig. 52. L. regularis ROSEN-
STOCK. Leaf, x 4% (PosTHUMUS 3202).
248
deeply incised. Veins immersed, evident, mostly
once forked, free except if loop-connected by
the receptacle. Sori 4-12 per pinnule, on (1-)2
(-4) vein-ends; indusium ovate, transversely
oblong to linear, 0.15-0.4 mm wide, not reaching
the margin by 114-3 times its width, reflexed,
concealed, often evanescent at maturity. Spores
very pale, trilete, smooth, c. 25 w.
Distr. Malesia: New Guinea (all
d’Entrecasteaux Is.
Ecol. Epiphytic or rarely terrestrial, in moist
forests, 80-2100 m.
Note. See after L. versteegii.
Div.);
58. Lindsaea rosenstockii BRAUSE, Bot. Jahrb.
56 (1920) 128.—Lectotype: LEDERMANN 10030,
Lordberg, Sepik R. region, Terr. of New Guinea
(B, 3 sh.).—Fig. 40.
Rhizome \ong-scandent, brown, 1144-2 mm g,
+ persistently scaly; scales honey-coloured,
narrowly triangular or triangular-ovate, up to
3 by 14% mm, up to c. 25-seriate at base, the
apex scarcely or not uniseriate. Leaves remote,
1144-15 cm apart, issuing at about right angles;
petioles stramineous to pale brown or mottled,
adaxially narrowly sulcate, abaxially terete or
faintly bi-angular, short in simply pinnate leaves,
6-20 cm long in bipinnate ones, shorter than the
lamina. Lamina simply pinnate or bipinnate; if
bipinnate oblong, 15-50 cm long, with 2-13
(often 6) pinnae to a side and a conform terminal
one; primary rachis abaxially subterete or mostly
bi-angular and convex. Pinnae little ascending,
their width apart or a little closer, sessile, linear,
8-15 cm long, 144-2144 cm wide, shortly acumi-
nate; simply pinnate laminas similar but mostly
larger. Secondary rachises adaxially broadly
sulcate, abaxially bi-angular almost to the base,
the greater part narrowly sulcate and green-
winged. Pinnules herbaceous, dark green when
dry, c. 15-25 to a side, close, spreading, slightly
ascending, or often slightly falcately decurved,
¥,-ovate in outline, the larger 1-134 cm by 4-7
mm, mostly about 2% times as long as wide;
lower margin straight or concave, upper margin
convex, a distinct outer margin not developed.
Pinnules deeply incised from the upper margin,
with 4 or 5 segments, the inner 1 or 2 bifid, or
rarely twice bifid; lobes 24-2 mm wide, at the sorus
broadened, to 144 mm wide, parallel-sided or
slightly narrowed at the base, joined by a wing of
Y, mm, subacute if sterile, rounded or sinuate-
rounded if fertile. Upper pinnules rather suddenly
reduced, confluent into a pinnatisect pinna-
apex. Veins immersed, evident, 1 or 2 per lobe.
Sori uni- or on the inner lobes binerval; indu-
sium pale, subentire, 0.6-1144 mm long, 0.3 mm
wide, with approximately straight base, not reach-
ing the margin by about twice its width, strongly
teflexed and often concealed at full maturity.
Spores pale brown, trilete, smooth, c. 28 yu.
Distr. Malesia: New Guinea (all 3 Divisions).
Ecol. Epiphytic in moist montane forest,
1000-1800 m.
Note. It is possible that in this species adult
FLORA MALESIANA
[ser. II, vol. 14
plants may produce simply pinnate leaves.
59. Lindsaea versteegii (CHRIST) v.A.v.R. Handb.
Suppl. (1917) 206; CopELAND, Philip. J. Sc. 78
(1949) 18.—Odontosoria versteegii CHRIST, Nova
Guinea 8 (1909) 157.—Type: VERSTEEG 1222,
Noord-Rivier, W. New Guinea (L; dupl. in B,
BO, K, U; fragm. in US).—Fig. 39.
Rhizome long-scandent, 1-2 mm g, + per-
manently scaly; scales honey-coloured, triangular,
to 2 mm long, up to c. 20-seriate, the apex scarcely
uniseriate. Leaves 1%-4 cm apart; petioles
stramineous to medium brown, + lustrous,
abaxially terete, adaxially narrowly sulcate, in
simply pinnate leaves almost wanting, in bi-
pinnate ones 10-15 cm long, shorter than the
lamina. Lamina simply pinnate or bipinnate;
if bipinnate oblong, c. 30 cm long, with c. 6 pinnae
to a side and a conform terminal one; rachis like
the petiole. Pinnae not strongly ascending,
sessile, their width apart or slightly closer, linear,
10-15 cm long, 2-214 cm wide, shortly acuminate;
upper pinnae somewhat shortened. Secondary
rachises slender, stramineous, abaxially + distinct-
ly bi-angular and sulcate. Pinnules thinly herba-
ceous, dark olivaceous when dry, spreading, close
but scarcely contiguous, c. 25 to a side; larger
pinnules arcuate-ligulate in outline, 10 to 12 by
4 to 5mm, c. 2% times as long as wide, the lower
margin concave, the upper margin deeply in-
cised; primary segments 4-6, the basal ones twice,
the upper ones mostly once forked; ultimate
lobes linear, divergent, 0.3-0.4 mm wide, almost
parallel-sided to the spathulate soriferous apex,
subacute if sterile, usually all lobes fertile, the
fertile segments suddenly broadened at the sorus,
there 15-34 mm wide, the apex rounded or nar-
rowed-rounded, subentire to erose; wings con-
necting the lobes about as wide as the lobes.
Upper pinnules gradually and strongly reduced,
confluent into the pinnatisect pinna-apex. Veins
immersed, evident, single in the lobes. Simply
pinnate leaves like the pinnae of bipinnate ones.
Sori uninerval; indusium pale, delicate, subentire
to erose, 4-34 mm long, % mm wide, if long
sometimes with concave base, otherwise semi-
elliptic, with free sides, not reaching the margin
by its own width to almost reaching it, some-
what reflexed at maturity. Spores pale brownish,
trilete, smooth, 22-25 yp.
Distr. Beside the type known from one other
collection from W. New Guinea (DOCTERS VAN
LEEUWEN 10276, BO, K, L).
Ecol. Epiphytic in forest; one reord 250 m.
Notes. It seems that adult plants may produce
once and twice pinnate leaves side by side.
Curist (Geogr. d. Farne, 1910, 233) made the
cryptic statement about this plant ‘eine prachtige
dimorphe Art mit halben Wasserblattern’.
The last three species, L. microstegia, L. rosen-
stockii, and L. versteegii, are very closely related
and are perhaps not specifically distinct.
April 1971]
LINDSAEA-GROUP (Kramer)
249
13. Section Penna-arborea
KRAMER, Blumea 15 (1968) 563.
Type species: Lindsaea pulchella (J. SMITH) METT. ex KUHN.
Distr. Malesia (except Malay Peninsula) to Polynesia and NE. Australia; most diversified in New
Guinea. Epiphytes of moist montane forest.
Taxon. The affinity of the species included in this section was not before recognized; L. pulchella
(J. SmirH) Metr. ex KUHN was usually compared, and often confused, with L. lucida BLUME (mostly
as L. concinna BLUME), L. werneri ROSENSTOCK Was associated with sect. Synaphlebium, because of its
anastomosing veins, efc. The wiry, long-creeping, eventually ++ scaleless and polished rhizome with
open xylem strand is quite distinctive. The lamina is unipinnate and the sori are interrupted.
60. Lindsaea pulchella (J. SMITH) METT. ex KUHN,
Linnaea 36 (1869) 81; KRAMER, Blumea 15 (1968)
570.—Odontoloma pulchellum J. SMiTH in Fielding
& Gardner, Sert. Pl. (1844) pl. 51.—Davallia
pulchella (J. SMiTH) Hooker, Sp. Fil. 1 (1845)
175, pl. 53 B.—Type: CuminG 217, Luzon (K;
dupl. in B, BM, E, L, MICH, P, US, W).
Aspidium adiantoides BLume, En. Pl. Jav.
(1828) 145.—Saccoloma adiantoides (BLUME)
PresL, Tent. Pterid. (1836) 126.—Dayvallia adi-
antifolia Hooker, Sp. Fil. 1 (1845) 176, non
Davallia adiantoides | Swartz. — Odontoloma
adiantoides (BLUME) PRESL, Epimel. Bot. (1851)
97.—Acrophorus adiantoides (BLUME) Moore,
Ind. Fil. (1857) 91.—L. adiantoides (BLUME)
KuuN in Mig. Ann. Mus. Bot. Lugd.-Bat. 4
(1869) 278, non J. SmitH in Hooker (1846).—
L. adiantifolia (HOOKER) COPELAND, Fern FI.
Philip. 1 (1958) 108.—Type: BLUME s.n., ‘Java’
(but probably from elsewhere).
L. cyathicola COPELAND, Philip. J. Sc. 1 (1906)
Suppl. II, 149, pl. 5; Fern Fl. Philip. 1 (1958)
106.—Type: CoPpELAND 1938, Mt Bulusan, Luzon
(MICH; dupl. in B, SING).
L. marginata BRAuUSsE, Bot. Jahrb. 56 (1920)
126.—Lectotype: LEDERMANN 11384, Hunstein-
spitze, Sepik R. region, Terr. of New Guinea (B).
L. alvestris v.A.v.R. Bull. Jard. Bot. Btzg III,
5 (1922) 210.—Type: BUNNEMEIJER 9907, Mt
Kerintji, Sumatra (BO; dupl. in K, L, U; fragm.
in BM, US).
L. binervata C. Cur. Bot. Jahrb. 66 (1933)
52.—Type: KJELLBERG 3632, Preho, Celebes
(BM; dupl. in BO, S-PA).—Fig. 53-56.
Rhizome long-scandent, dark reddish brown
to black, lustrous, wiry, %-14% mm g; scales
deciduous, more persistent at the petiole-bases,
elongate-triangular, to c. 8-10-seriate at base.
Leaves one to a few cm apart, issuing at about
right angles; petioles slender, wiry, less than 1 mm
@. Lamina much longer than the petiole, linear,
simply pinnate; pinnules asymmetrically ovate,
semi-elliptic, ligulate, or rounded-trapezoidal,
2-3 times as long as wide; upper margin incised.
Upper pinnules gradually and strongly reduced,
basal pinnules farther apart and often + reduced.
Sori usually uni- or binerval. Spores pale yellow,
trilete, smooth, c. 20-25 yw.
KEY TO THE VARIETIES
ils Uninerval sori with straight or convex base;
indusium falling short of the margin by its own
width or less; veins free or anastomosing;
scales reddish brown, with a well-developed
uniserate apex. . . 4. var. lomatosora
1. Uninerval sori mostly with very concave base;
indusium falling short of the margin by its
own width or more (except sometimes at
incisions); veins free; scales golden brown,
apically very shortly uniseriate.
2. Pinnules herbaceous, mostly green when dry,
41%-8 mm long, c. twice as long as wide,
the major incisions less than 4% mm deep;
adaxial and abaxial sides of petiole not
discolorous . Ac 1. var. pulchella
2. Pinnules herbaceous, mostly brown when
dry, the larger ones 7-12 mm long, up to
twice as long as wide, the major incisions
1-3 mm deep; petiole faces not discolorous.
2. var. blanda
Pinnules herbaceous to subcoriaceous, 10-16
mm long, 214-3 times as long as wide; in-
cisions to 1 (sometimes to 2) mm deep;
adaxial face of petiole much paler than abaxi-
al side . ‘ 3. var. falcata
i)
1. var. pulchellaa—KRAmeER, Blumea 15 (1968)
571.—Aspidium adiantoides BLUME.—L. binervata
C. Cur. —Fig. 53.
Rhizome ¥%—24 mm 9; scales narrowly trian-
gular, golden-brown, to 24% mm long, to c. 8-
seriate at base, with a short uniseriate apex. Peti-
oles 1-7 cm long, c. %-13 mm 9g, stramineous
with dark base, or brown, abaxially rounded and
bi-angular above or less often largely bi-angular,
adaxially often pale-margined. Lamina 10-30 cm
long, 0.8-14% cm wide; rachis stramineous,
quadrangular, often shallowly sulcate, Wiry.
Pinnules herbaceous, mostly pale to medium green
when dry, in larger specimens c. 25-70 to a side,
close to contiguous, alternate, spreading, asym-
metrically ovate to semi-elliptic, the larger ones
41%-8(-9) mm long, 2-4(-5) mm wide, c. twice
as long as wide, mostly narrowed-rounded or
subacute at the apex; margin not sclerotic, the
upper margin with 1-3 very shallow incisions less
than % mm deep, the lobes truncate-rounded
or the outer ones with a small tooth; upper pin-
nules gradually and strongly reduced, several
denticuliform ones confluent into a short, narrow,
caudiform, pinnatifid leaf-apex. Veines immersed,
evident, simple or once forked, ending well
within the margin, mostly 1 mm apart. Sori
uni- or binerval, often absent from part of the
lamina; indusium elongate and often, especially
250 FLORA MALESIANA [ser. Il; volal@
r).
6
PBL
S
IPP
SYVVSVs
ihe
_LbL£
Fig. 53-56. Lindsaea pulchella (J. SM.) Metr. ex KuHN.—Fig. 53. var. pulchella. Part of a plant,
x Ye, pinnule, x 3 (FORSTEN s.n., L).—Fig. 54. var. falcata (BRAUSE) KRAMER. Part of a lamina,
x 2 (BRASS 13494).— Fig. 55. var. lomatosora KRAMER. Part ofa lamina, x 21% (BRASs 27911).— Fig.56.
var. blanda (METT. ex KUHN) KRAmer. Part of a lamina, * 3 (BUNNEMEIJER 9907).
April 1971]
LINDSAEA-GROUP (Kramer)
25k
in outer sori, with concave base, or if uninerval
ovate to subreniform and very obtuse to acute,
variable in width, usually 4%,—'% mm wide, often
with irregular margin, not reaching the margin by
less than its width to reaching it, laterally free,
scarcely reflexed at maturity.
Distr. Malesia: Philippines (Mindanao,
Negros, Mindoro, Luzon, Panay, Leyte), Mo-
luccas (Halmahera, Ternate, Tidore, Buru),
Celebes, Flores, New Guinea (1 coll.). Map 9.
Map 9. Distribution of Lindsaea pulchella (J.
Sm.) Metr. ex KUuHN.—var. pulchella (lined),
var. blanda (METT. ex KUHN) KRAMER (inter-
rupted line, arrows!), var. falcata (BRAUSE)
KRAMER (stippled line, arrow!), var. lomatosora
KRAMER (squares).
Ecol. Epiphytic in moist forests, c. 600-2200
m, on trees and tree-ferns.
2. var. blanda (METT. ex KUHN) KRAMER, Blumea
15 (1968) 571.—L. blanda Mett. ex KunNn,
Linnaea 36 (1869) 80.—Type: WICHURA s.n.,
Java (B).
L. alpestris v.A.v.R.—L. cyathicola COPELAND.—
Fig. 56.
Rhizome (*4-)34-1% mm g; scales often a little
longer and broader than in var. pulchella. Petioles
1-10 cm long, %4-%3 mm g, dark reddish brown
or upward paler, hardly pale-margined, adaxially
flattened, upward sulcate, abaxially bi-angular
except at the rounded base. Lamina 5-30 cm long,
1-24% cm wide; rachis quadrangular, at least
adaxially sulcate, stramineous or with darker base,
wiry. Pinnules herbaceous, mostly olivaceous-
brown when dry, c. 20-50 to a side, usually a
little ascending, mostly not contiguous, asym-
metrically ovate to 14,-elliptic, 7-12 mm long,
3144-6 mm wide, twice as long as wide or slightly
less; margins not or little sclerotic, a distinct
outer margin usually not or scarcely developed,
the upper margin with 1-3 oblique major incisions
1-3 mm deep, reaching 13 to 1% (rarely to 7)
down, sometimes with some shallower additional
incisions; lobes convex, not rarely erose, -+
divergent. Basal pinnules farther apart but scarce-
ly reduced; upper pinnules gradually reduced,
as in var. pulchella. Veins immersed, not evident,
simple or once forked, free, ending well within
the margin, %4—-1 mm apart. Sori uni- or binerval
or less often to 5-nerval, %-—2(-4) mm long, in
longer sori the base + concave. Indusium pale
to brownish, delicate, subentire, with -+_ convex
free edge, narrowed at the free sides, 0.3-0.5 mm
wide, nearly always strongly intramarginal.
Distr. Malesia: Sumatra, West and Central
Java, Philippines (Luzon, Mindanao), Celebes, ?
Ternate, New Guinea (all Div.); Queensland;
Solomon Is. Map 9.
Ecol. Epiphytic, often among mosses, on trees
and tree-ferns, very rarely terrestrial, from c.
1500 up to 2750 m.
3. var. falcata (BRAUSE) KRAMER, Blumea 15
(1968) 571.—L. marginata BRAUSE var. marginata
et var. falcata BRAUSE, Bot. Jahrb. 56 (1920)
126-127.—Type: LEDERMANN 11384, Hunstein-
spitze (var. marginata) (B), LEDERMANN 12864,
Felsspitze (var. falcata), Sepik R. region, Terr.
of New Guinea (B; dupl. in BM).
L. rhombifoliolata v.A.v.R. Nova Guinea 14
(1929) 29.—Type: H. J. Lam 1892a, crest to
Doormantop, W. New Guinea (L).—Fig. 54.
Rhizome 1-1% mm @g; scales to 24% mm long,
otherwise as in var. pulchella. Petioles c. 10-20
cm long, 4% mm g, adaxially flattened or shallow-
ly sulcate, pale at least above, abaxially dark,
rounded, bi-angular at least near the apex. Lamina
c. 20-35 cm long, 2-3 cm wide, with c. 25-50
pinnules to a side; rachis abaxially dark, bi-angu-
lar, sometimes pale-angled, upward usually
pale and sulcate, adaxially pale, sulcate. Pinnules
herbaceous or usually subcoriaceous, dark brown-
ish when dry, usually everywhere sclerotic-
margined, somewhat ascending, not contiguous,
ligulate, the larger ones 10-16 by 4-6 mm (2-)
24-3 times as long as wide; upper margin with
2-4 narrow major incisions to 2 mm but usually
only 1 mm or less deep, sometimes with some
smaller additional ones, the lobes rounded,
often with one or more teeth. Basal pinnules
often more remote but scarcely reduced, upper
pinnules gradually and strongly reduced, as in
var. pulchella. Veins immersed, not evident,
simple or once forked, lax, 1-2 mm apart. Sori
uni- or binerval, usually with distinctly concave
base, 34-2 mm long; indusium pale, rather rigid,
subentire to dentate or lobed, subhippocrepi-
form to crescent-shaped, narrowed at the free
sides, usually strongly intramarginal but some-
times with protruding lobes, 0.3-0.5 mm wide,
often reflexed at maturity.
Distr. Malesia: New Guinea (all Div.), Japen;
New Hebrides (Aneityum). Map 9.
Ecol. Epiphytic, or terrestrial
1300-1800 m.
on mosses,
4. var. lomatosora KRAMER, Blumea 15 (1968)
571.—Type: Brass 27911, Mt Riu, Sudest I.,
Louisiades (L; dupl. in GH, K).—Fig. 55.
Rhizome scales reddish brown, to 134 mm long,
with a well-developed uniseriate apex. Leaves up
to 30 by 2 cm; faces of the petiole not or hardly
discolorous; pinnules green when dry, spreading
Pay
FLORA MALESIANA
[ser. I, volza4
or slightly ascending, mostly ligulate or rounded-
trapezoidal, scarcely narrowed at the apex;
veins sometimes connivent or anastomosing,
the series of areoles usually incomplete. Margin
scarcely sclerotic, its incisions 1 mm deep or less;
sori mostly binerval, basally scarcely concave or
straight, if short more concave; indusium pale,
delicate, 0.3 mm wide, falling short of the margin
by less than its width to almost reaching it, often
strongly bulging at maturity and then seemingly
more strongly intramarginal. Otherwise much
like var. falcata.
Distr. Louisiades, d’Entrecasteaux Is.; one
less typical collection from the Territory of New
Guinea. Map 9.
Ecol. Epiphytic on trees and tree-ferns, 250—
900 m.
Note. The scales suggest a distinct species;
but it is in other respects very much like the
other varieties of L. pulchella.
61. Lindsaea werneri ROSENSTOCK in Fedde, Rep.
5 (1908) 37; COPELAND, Philip. J. Sc. 78 (1949)
21.—Type: WERNER (ROSENSTOCK-exs. 19), Mt
Gelu, Terr. of New Guinea (B; dupl. in E, L,
P, S-PA, U).—Fig. 38.
Rhizome long-scandent, fawn-coloured, 0.6-1
mm 9g, deciduously scaly; scales light golden-
brown, narrowly triangular, to 2 mm long, to
10-seriate at base, with a very short or virtually
lacking uniseriate apex. Leaves 2-10 cm apart,
issuing at about right angles; petioles quadrangular
and sulcate almost to base, stramineous to reddish
brown, often with paler angles, 4-15 cm long,
somewhat shorter than the lamina. Lamina narrow-
ly oblong, simply pinnate, 7-32 cm long, 3-6 cm
wide, with 6-20 pinnules to a side, shortly acumi-
nate; rachis stramineous, quadrangular, sometimes
also quadrisulcate. Pinnules herbaceous, oliva-
ceous when dry, spreading, alternate except the
basal ones, their width apart to slightly over-
lapping, sessile, subtrapezoidal to semi-ovate-
ligulate, 144-3 cm long, 6-12 mm wide, 2 to
almost 3 times as long as wide; upper margin
straight or outward convex, a distinct outer
margin sometimes developed, then forming an
angle of 90° or less with the upper; upper (and
outer) margin shallowly crenate, if fertile with c. 5
incisions and ++ flat lobes, if sterile with more and
closer incisions and rounded lobes, the deepest
incisions to c. 1 mm, mostly not reaching the
level of the receptacle. Few or very few upper
pinnules suddenly reduced, one or two connected
with the narrow, often caudate, to 4 cm long
terminal segment. Veins immersed, evident, lax,
anastomosing, forming an often incomplete series
of areoles of varying length and 14-2 mm
maximum width between the margins. Sori single
in the lobes, most often binerval, sometimes tri-
or uninerval, 1-3 mm long, with straight or some-
what convex receptacle; indusium pale, delicate,
subentire, free at the sides, 0.3-0.4 mm wide,
not reaching the margin by 1 44-2 times its width,
reflexed and + concealed at maturity. Spores
pale brown, trilete, smooth, c. 22 m.
Distr. Malesia: Sumatra (one coll., SURBECK
s.n., L), Moluccas (Halmahera, Ceram), New
Guinea (W. New Guinea, Terr. of New Guinea).
Map 10.
Map 10. Distribution of Lindsaea werneri ROSEN-
stock (dots) and L. glandulifera v.A.v.R.
(crosses).
Ecol. Epiphytic, or occasionally terrestrial,
in dense, moist forests, 800-1800 m.
62. Lindsaea roemeriana ROSENSTOCK, Nova
Guinea 8 (1912) 719; CopPELAND, Philip. J. Sc.
78 (1949) 17.—Lectotype: VON ROMER 731, Hell-
wig Mts, W. New Guinea (L; dupl. in BO).
L. wollastonii v.A.v.R. Handb. Suppl. (1917)
505, based on: Odontosoria tenera RIDLEY, Trans.
Linn. Soc. II, Bot. 9 (1916) 254; non L. tenera
DRrYAND.—Type: B. KLoss (WOLLASTON Exp.)
s.n., Carstensz Peak, W. New Guinea (K; dupl.
in BM).—Fig. 23.
Rhizome long-scandent, wiry, %-34 mm 4g,
atropurpureous to blackish, lustrous, soon devoid
of scales; scales honey-coloured, narrowly tri-
angular, to 2 mm long, to 8-seriate at base, the
uniseriate apex of one cell or wanting. Leaves
remote, 144-5 cm apart; petioles stramineous
except for the dark base, quadrangular almost to
base, at least adaxially sulcate, 4-15 cm long,
shorter than the lamina. Lamina narrowly oblong,
8-25 cm long, 2—3 % cm wide, with 15-30 pinnules
to a side; rachis similar to the petiole, upward
narrowly green-margined. Pinnules herbaceous,
olivaceous when dry, spreading, their width
apart to subcontiguous, mostly alternate except
the basal ones; larger pinnules semi-ovate in
outline, 10-18 mm long, 5-8 mm wide, 2-2%
times as long as wide, deeply pinnatifid from the
upper margin, with 3—4 primary segments, the
inner one or two forked; segments 0.3-0.6(-1)
mm wide at the base, 0.5—1.5(-2) mm wide at the
sorus, gradually widened below, more suddenly
at the sorus, connected by a wing %43—'% mm wide,
rounded and erose to irregularly corniculate at
apex. Sterile segments not broadened at apex,
acute or subacute. Basal pinnules more remote and
sometimes slightly reduced; upper pinnules (in
the upper 144 or 1%) reduced, the uppermost
confluent into a small pinnatifid leaf-apex. Veins
immersed, evident, solitary in the lobes. Sori
April 1971]
solitary near the apices of the lobes, uninerval;
indusium delicate, ovate to hippocrepiform, sub-
entire, free at the sides, 0.3-0.5 mm wide, 0.5-1
mm long, not reaching the margin by 1-2 times
its width, scarcely reflexed at maturity. Spores
very pale, trilete, smooth, c. 22 yu.
Distr. Malesia: New Guinea (W. New Guinea,
LINDSAEA-GROUP (Kramer)
253
Terr. of New Guinea, 12 coll.).
Ecol. Epiphytic in mountain forests, c. 1200-
2500 m.
Note. There are two extreme forms, one with
more rounded lobes and sori with concave base,
the other with erose lobes and ovate sori, but
they pass into each other.
Excluded from Lindsaea
(referred to Jsoloma, Lindsaea, Schizolegnia, or Schizoloma)
Isoloma dicksonioides (CHRIST) TARDIEU-BLOT, Not. Syst. 14 (1952) 331 = Nephrolepis dicksonioides
CHRIST.
Tsoloma lanuginosum J. SMiTH in Hooker & Bauer, Gen. Fil. (1842) pl. 102 = Nephrolepis acutifolia
(DESVAUX) CHRIST.
Isoloma lindsayae (CHRIST) TARDIEU-BLOT, Not. Syst. 14 (1952) 331 = Nephrolepis spec.
Lindsaea acutifolia DESVAUXx, Prod. (1827) 312 = Nephrolepis acutifolia (DESVAUX) CHRIST.
Lindsaea amboynensis (HOOKER) METT. ex KUHN in Miq. Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 279 =
Tapeinidium amboynense (HOOKER) C. CHR.
Lindsaea chinensis (L.) METT. ex KUHN, Fil. Afr. (1868) 67 (non CHING, 1929) = Sphenomeris chi-
nensis (L.) MAXON.
Lindsaea cordata (GAUD.) DEsvVAUx, Prod. (1827) 312 = Schizolepton cordatum (GAuUD.) FEE =
Taenitis cordata (GAUD.) HOLTTUM.
Lindsaea cuneifolia PRESL, Rel. Haenk. 1 (1825) 60 = Sphenomeris retusa (CAV.) MAXON.
Lindsaea denhamii (HOOKER) METT. ex KUHN, Verh. Zool. Bot. Ges. 19 (1869) 573 = Tapeinidium
denhamii (HOOKER) C. CHR.
Lindsaea grandifolia J. E. SmMitH in Rees, Cyclop. 21 (1812) no 12 = Taenitis blechnoides (WILLD.)
SWARTZ feste ALSTON, Philip. J. Sc. 50 (1933) 180.
Lindsaea hosei C. Cur. Ind. Fil. (1906) 394, based on: L. trilobata BAKER, J. Bot. 29 (1891) 107 (non
COoLENSO, 1884) = Taenitis trilobata HoLTTUM, Blumea 16 (1968) 93.
Lindsaea lanuginosa WALL. ex HOOKER, Sp. Fil. 1 (1846) 210, pl. 69 B = Nephrolepis acutifolia
(DESVAUX) CHRIST.
Lindsaea lowei hort. = Arthropteris obliterata (R. BROWN) J. SMITH teste C. Cure. Ind. Fil. (1906)
395.
Lindsaea parishii BAKER, Syn. Fil. ed. 1 (1867) 109 = Stenochlaena sorbifolia (L.) J. SMITH teste C.
Cnr. Ind. Fil. (1906) 625 = ? Teratophyllum.
Lindsaea pinnata (CaAv.) METT. ex KUHN in Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 279 = Tapei-
nidium pinnatum (CAv.) C. CHR.
Lindsaea pinnata (CAv.) METT. ex KUHN var. bipinnata METT. ex KUHN in Mig. Ann. Mus. Bot.
Lugd.-Bat. 4 (1869) 279 = Tapeinidium luzonicum (HOOKER) KRAMER.
Lindsaea retusa (CAv.) MeEtTT. Fil. Lips. (1856) 105 = Sphenomeris retusa (CAy.) MAXON.
Lindsaea trilobata BAKER, J. Bot. (1891) 107 = see under L. hosei C. CHR.
Lindsaea vittata ZOLL. & Mor. Nat. Geneesk. Arch. N. I. 1 (1844) 400 = Preris ?
Schizolegnia stortii (v.A.v.R.) ALSTON, Bol. Soc. Brot. I, 30 (1956) 24 = Xyropteris stortii (v.A.v.R.)
KRAMER.
Schizoloma cordatum Gaup. Ann. Sc. Nat. 3 (1824) 507 = see under Lindsaea cordata (GAUD.)
DESVAUX.
Schizoloma ferulaceum (MOORE) KUHN, Chaetopt. (1882) 346 = Davallia spec.
Schizoloma hosei (C. CHR.) COPELAND, Sarawak Mus. J. 2 (1917) 327 = see under Lindsaea hosei
C. Cure.
Schizoloma retusum (CAvV.) KUHN, Chaetopt. (1882) 346 = Sphenomeris retusa (CAV.) MAXON.
Schizoloma stortii v.A.v.R. Bull. Jard. Bot. Btzg II, 16 (1914) 36 = Xyropteris stortii (v.A.v.R.)
KRAMER.
Doubtful species and varieties
Lindsaea bipinnata RoxBURGH, Calc. J. Nat. Hist. 4 (1844) 511.—Type: RoxBuRGH s.n., Prince of
Wales’ Island (Pulau Penang) (n.v.).
Judging from the description this is a large form of L. parasitica, with which RoxBURGH compared
it; or it might be L. doryphora.
Lindsaea decrescens COPELAND, Philip. J. Sc. 81 (1952) 6, pl. 5.—Type: Loner 13621, Umiray,
Quezon Proy., Luzon (dupl. in MICH).
As stated by CopELAND, this is intermediate between L. repens (‘L. macraeana, L. longa’) and L.
254 FLORA MALESIANA [ser. I vou
fissa. The isotype in MICH from CopELAND’s herbarium is rather scrappy. I doubt whether it represents a
distinct species.
Lindsaea longa COPELAND, Philip. J. Sc. 46 (1931) 216.—Type: EDANo B.Sc. 77978, Mt Balagbag,
Palawan (MICH; dupl. in GH).
This is probably only an unusually deeply incised form of L. repens (BoRY) THWAITES var. sessilis
(COPELAND) KRAMER.
Lindsaea orbiculata (LAMK) METT. ex KUHN var. integra METT. ex KUHN in Mig. Ann. Mus. Bot.
Lugd.-Bat. 4 (1869) 279.—Type: Loss s.n., Java; also “Ceylania’.
I have seen no specimen that can with certainty be regarded as the type. It is not impossible that it
represents var. commixta for which it would then be an older name.
Lindsaea striata BLUME, En. Pl. Jay. (1828) 220.—Type: BLUME s.n., ‘in Javae montibus altissimis’
(n.¥v.).
The type of this species could not be found at L and is presumably lost. CHRISTENSEN (Ind. Fil. 1906)
referred it to L. orbiculata; judging from the description and the provenience this is probably correct.
Lindsaea tricrenata BAKER, J. Bot. 28 (1890) 106.—Type: MCGREGOR 24, Mt Musgrave, New
Guinea (K).
Compared with L. rigida and L. borneensis by its author, two species not closely related in the present
author’s opinion. The type is a detached, very poorly preserved, nearly sterile leaf of uncertain identity,
and one can only marvel at BAKER’s temerity to describe this as a new species.
Vittaria interrupta ROXBURGH ex GRIFFITH, Calc. J. Nat. Hist. 4 (1844) 511.—Type: RoxBuRGH s.n.,
Prince of Wales’ Island (Pulau Penang) (n.y.).
Referred to L. orbiculata by CHRISTENSEN (Ind. Fil.), to L. tenera by BEDDOME (Ferns Brit. Ind. 1,
1866). Judging from the description the latter opinion seems better; it would then be the oldest name for
what is here called L. bouillodii CHRIST.
Vittaria lunulata ROXBURGH ex GRIFFITH, Calc. J. Nat. Hist. 4 (1844) 510.—Type: coll.?, Prince of
Wales’ Island (Pulau Penang) (n.y.).
Perhaps a simply pinnate form of L. parasitica; it is not stated whether the plant was terrestrial or
epiphytic, with long or short rhizome, and its identity remains obscure.
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, it —v
LOMARIOPSIS GROUP (R. E. Holttum, Kew)!
Aspidiaceae sensu CHING, Sunyatsenia 5 (1940) 247-253, p.p.; Aspidiaceae sensu
CopeL. Genera Filicum (1947) 100-154, p.p. — Dennstaedtiaceae subfam. Lomariop-
sidoideae HoLtTuM, J. Linn. Soc. Bot. 53 (1947) 146-149. — Lomariopsidaceae
ALSTON, Taxon 5 (1956) 25. — Elaphoglossaceae PicHi SERM. Webbia 23 (1968)
209-217.
Rhizome creeping or low-climbing (Bolbitis) or climbing (Lomariopsis, etc.) or
epiphytic (Elaphoglossum), dorsiventral, with a broad ventral vascular strand which
supplies the roots and one or more dorsal strands (the fronds in two or more
longitudinal rows, according to the number of strands); stipes jointed to rhizome
(Teratophyllum, Elaphoglossum) or not, containing several separate vascular
strands; scales peltate or pseudopeltate, clathrate or not; no elongate unicellular
hairs. Rhizomes of young plants always with one dorsal meristele, this condition
persisting to the adult plant in Teratophyllum and many species of Elaphoglossum.
Fronds simple (Elaphoglossum, Bolbitis spp.), pinnate (all but Elaphoglossum) or
bipinnate (Teratophyllum and Lomagramma spp.), the pinnae on fronds of Loma-
riopsis, Teratophyllum and Lomagramma jointed to the rachis, terminal unjointed
lamina present in Lomariopsis; distinctive bathyphylls, usually more dissected than
acrophylls, present in genera with climbing rhizomes (least distinctive in Lomariop-
sis); veins free (Teratophyllum, Lomariopsis, most Elaphoglossum, some Bolbitis) or
uniting near the margin (Elaphoglossum spp.) or in several series of areoles with
(most species of Bolbitis) or without (Lomagramma; Bolbitis p.p.) free veins in the
areoles. Fertile fronds with reduced lamina, covered beneath (rarely also above)
with sporangia (except 7hysanosoria, where sori are at ends of veins only), a special
vascular supply for the sporangia variously developed or not; spores with peris-
pore (except Lomagramma).
Genera. Bolbitis ScHott, Lomariopsis FEE, Lomagramma J.SM., Teratophyllum Metr., Thysanosoria
Gepp, Elaphoglossum J.SM.; also Peltapteris LINK (Rhipidopteris FEE ex ScHott) and Microstaphyla
PRESL, small genera of tropical America and St Helena, allied to Elaphoglossum and not dealt with in the
present work.
Taxonomy. In earlier systems species of all genera (except Thysanosoria) were included in Acrostichum
because fertile leaflets are covered beneath with sporangia, without any distinction of separate sori. When
Pres and FEE distinguished separate genera among such ferns, they depended mainly on venation, and
so for example associated Lomagramma with the very different genus Chrysodium (now known as Acro-
stichum s.str.) because of similarity of venation, and Egenolfia and Bolbitis (here united as one genus) were
placed wide apart because one had free veins and the other anastomosis. JOHN SMITH, who knew a large
number of ferns from the living plants he cultivated at Kew, introduced habit of growth, and especially
articulation of fronds to rhizome and pinnae to rachis, as additional characters, and thereby made
further progress towards a natural system (Historia Filicum 1875).
In CHRISTENSEN’S Index Filicum (1905-06), based largely on the system of DieELs in Engler’s Pflanzen-
familien (I, Abt. 4, 1899) the genera here included were ranked as follows: Bo/lbitis (excluding free-veined
species, i.e. Egenolfia) and Lomagramma (excluding L. polyphylla) were treated as sections of the genus
Leptochilus in tribe Aspidieae; Egenolfia was treated as a section of the genus Po/ybotrya (also in Aspidieae)
and Teratophyllum articulatum with Lomagramma polyphylla were placed in section Arthrobotrya of the
same genus; Lomariopsis and Teratophyllum (apart from T. articulatum) were merged with the genus
Stenochlaena (not even distinguished as sections) in Blechninae, a subtribe of Asp/enieae; the single species
of Thysanosoria (a name not then established) was included in the genus Notho/aena in Pterideae; Elapho-
glossum was placed with Acrostichum in the tribe Acrosticheae.
In CHRISTENSEN’s first Supplement to his Index (1913) he recognized Lomagramma as a distinct genus,
following CoPELAND’s observations of Philippine spp. In 1931 he recognized that Bo/bitis (under the name
Campium) and Egenolfia are so closely related that they might well be united, and regarded them as
‘acrostichoid derivatives from the Dryopterideae’ (Contr. U.S. Nat. Herb. 26: 291). In 1932 I distinguished
Teratophyllum and Lomariopsis from Stenochlaena (Gard. Bull. S. S. 5: 245-312) and demonstrated that
the latter is peculiar in growth-habit, venation, anatomy and spores, so that it should belong to another
(1) The treatment of Bolbitis is by E. Hennipman, Leyden.
(255)
256 FLORA MALESIANA [ser. I, vokw@
group of genera. In discussing the possible relationships of Teratophyllum and Lomariopsis I remarked on
the similarity of their rhizome-structure and spores to those of Bolbitis and Egenolfia, suggesting that the
four genera, and also Lomagramma, formed a natural group (/.c. 307-309). In 1938 this conclusion was
accepted by CHRISTENSEN (in Verdoorn, Man. Pterid. 545) who regarded all the genera (with the addition
of Thysanosoria) as acrostichoid derivatives allied to Dryopteris. In the same work (p. 549) CHRISTENSEN
placed Elaphoglossum in a distinct subfamily (of Polypodiaceae) of doubtful relationship. CHING empha-
sized the isolation of Elaphoglossum by establishing a distinct family for it (Sunyatsenia 5, 1940, 265).
When preparing my fern-flora of the Malay Peninsula, I was struck by resemblances between Elapho-
glossum and Lomariopsis, and added Elaphoglossum to the group, which I called subfam. Lomariopsi-
doideae because Lomariopsis seemed to be a central genus (J. Linn. Soc. Bot. 53, 1947, 146-149). I placed
this subfamily in a family Dennstaedtiaceae, not for nomenclatural reasons but because I wished to empha-
size the idea that these ferns, and may be others, were likely to have evolved from something like Denn-
staedtia and that their relationship to Polypodium is much more remote, so that the family name Poly-
podiaceae is quite inappropriate for them. In this broader aspect of classification my ideas have now
changed somewhat (Brit. Fern Gaz. 9, pt 6, 1965, 205-212) but I still think that these genera form a
natural group and that they should not be included in a family named Polypodiaceae. ALSTON (l.c.)
proposed for them the family name Lomariopsidaceae, but included formally in his family only the genera
found in West tropical Africa. COPELAND (1947) included all genera in his family Aspidiaceae, though
regarding Elaphoglossum as not closely related to the others.
In 1949 I placed subfamily Lomariopsidoideae near Davallioideae (Bot. Rev. 24: 275, 290). I still think
these two groups may be rather closely related, but would not consider them to have had a common
origin from ferns like Microlepia. Some comments on other possible relationships are given below. The
chromosome number 41, found in Bolbitis, Lomariopsis, Teratophyllum, Lomagramma and Elaphoglossum
might indicate an association with Davallia or Ctenitis or Dryopteris (ROY & MANTON, J. Linn. Soc. Bot.
59, 1966, 343-347; T. G. WALKER, Trans. R. Soc. Edinb. 69, 1966, 178).
PICHI SERMOLLI (/.c. 1968) has argued that E/aphoglossum is not closely related to the other genera, and
excluded it from a family Lomariopsidaceae. But he based this opinion mainly on a comparison between
Elaphoglossum and Bolbitis, which admittedly are not closely allied. In my original proposal for its present
assignment (J. Linn. Soc. Bot. 53, 1947, 149) I expressly compared Elaphoglossum with Lomariopsis, which
I regarded as the central genus of the group. Young plants of most species of Lomariopsis (and adult plants
of some African species), which have simple fronds, only differ conspicuously from Elaphoglossum in lack
of articulation of fronds to rhizome. The range of spore-structure in the two genera is similar and I do not
think it affords distinguishing characters. However, gametophytes of Elaphoglossum do appear to be
sharply distinct from those of Lomariopsi§ (though the latter have not as yet been raised to maturity). The
two genera have certainly had a long separate history and both are pantropic. Differences in both sporo-
phyte and gametophyte may be due to adaptations to differences of habitat, to which gametophytes are
probably more sensitive than sporophytes.
gkeole gy. The genera may be divided into three groups, according to growth-habits and ecological
adaptations.
(1) Bolbitis (in Malesia) has short-creeping rhizome (except sometimes in young plants) usually growing
on rocks or stream-banks in shady forest, sometimes climbing short distances up tree-trunks but never
high-climbing. Pinnae are never jointed to the rachis, there being no need for adaptation to a dry season,
but probably fertile fronds are produced in response to somewhat drier conditions.
(2) Lomariopsis, Thysanosoria, Teratophyllum and Lomagramma. Young plants start their lives on rocky
or earthy banks of streams in tall evergreen forest, or on the exposed roots of trees, always in very humid
conditions (some in fresh-water swamp forest), much as Bolbitis, but the rhizomes ultimately climb a
considerable height up the trunks of trees (never into fully exposed positions), bearing spreading or
drooping fronds. In all cases the pinnae are jointed to the rachis and shed when old; this is an adaptation
to the high-climbing conditions in which the fronds are in air drier than near the ground (epiphytic orchids
all have jointed leaves). In Teratophyllum the stipes are rather imperfectly jointed to the rhizome, leaving
round scars when shed; in Lomariopsis and Lomagramma the stipe-bases are decurrent on the rhizome
without a joint. In Lomariopsis the early fronds are simple and entire (dissected only in L. variabilis
(WILLD.) FEE of Mauritius) and are followed by pinnate fronds of increasing size, without any abrupt
transition. In Teratophyllum the rhizome of a young plant climbing a tree-trunk near the ground has much-
divided fronds, here called bathyphylls, of different form in different species, and there is a fairly abrupt
transition to the simply pinnate adult condition. In Lomagramma young plants creep for some distance on
stream-banks (often on rocks) with erect simply pinnate fronds of a distinctive kind, and show a rather
abrupt transition to the adult condition, both as regards size of rhizome and of fronds, when they start to
climb a tree. In all these genera the plants are rooted in the ground and draw their principal water-supply
from the ground. Thysanosoria is exactly like Lomariopsis in its growth-form as an adult plant and probably
resembles Lomariopsis in its young stages, though these have not yet been observed.
(3) Elaphoglossum species are all normally epiphytic (in Malesia). They have short-creeping rhizomes,
to which the fronds are jointed; the joint is rather an imperfect one, between an outgrowth from the
rhizome (phyllopodium) and the base of a frond (some American species have no joint). Fronds are thick
and fleshly in many cases, especially those growing in more exposed positions in the crowns of trees;
those growing near the ground in shade (e.g. E. melanostictum) have thinner fronds. The shedding of whole
fronds, and their fleshy nature, are adaptations to the epiphytic condition (Elaphoglossum fronds are often
much like leaves of orchids which grow with them).
The dorsiventral rhizome. The dorsal half of a creeping rhizome of Bo/bitis has a vascular struc-
1978] LOMARIOPSIS GROUP (Holttum) 257
ture similar to that of Dryopteris or Tectaria but the ventral half of the Bo/bitis rhizome is quite different,
being specialized to root-bearing functions and carrying no fronds. There is no doubt that this structure
is an advantage to ferns growing on stream-banks and subject to periodic submersion in swift-flowing
flood-water. A similar structure is also well-adapted to the high-climbing condition, and to epiphytes
which need to be firmly attached to tree-branches; it is found throughout the family Pol/ypodiaceae (s.str.),
the principal group of epiphytic ferns, though Polypodiaceae cannot be considered closely allied to the
present group of genera.
As indicated below I suggest that the Lomariopsis group of genera are related to the Tectaria group; if
so, they presumably evolved from ancestors with radially organized suberect stems. P. R. BELL has shown
that some tropical American species of Elaphoglossum have a radially organized stem, though most are
dorsiventral, and has found one species in which the change from dorsiventral to radial occurs in the
development of a single plant (Ann. Bot. n.s. 19, 1955, 178-180). If one regards all genera of the present
group as originating from ancestors with radially-organized rhizomes, BELL’s plant indicates a reversion
to the ancestral condition; whether all those tropical American species of Elaphoglossum which have
radially organized rhizomes in the adult plant represent a similar reversion is an open question. It is also
an open question whether the dorsiventral condition can have originated on more than one evolutionary
line within the group of genera.
NAYAR makes a suggestion as to the origin of a dorsiventral rhizome from a radially symmetrical one,
based on “‘some species of Tectaria in which some of the leaves on the ventral side of the procumbent or
creeping rhizome are partially suppressed, and are associated with markedly smaller leaf-gaps in the stelar
cylinder of the rhizome’’ (New Phyt. 65, 1966, 237). If such smaller leaf-gaps were completely suppressed a
dorsiventral rhizome would result.
Inter-relationships. Assuming that this group of genera had a common ancestor, it was probably a
terrestrial fern with dorsiventral creeping rhizome, bipinnate fronds with free veins and a tendency to
contraction of fertile leaflets. Changes from this condition were: fronds in most cases to simply pinnate or
simple; anastomosis of veins; a high-climbing rhizome and in conjunction with it the articulation of
pinnae; loss of indusia and spreading of sporangia to cover the lower surface of reduced fertile leaflets.
Some of these changes certainly occurred on more than one evolutionary line. Bolbitis represents the
evolutionary line (or lines?) in which plants remained terrestrial and pinnae did not become articulated.
It is more diversified than any of the other genera, especially in venation; those species which have free
veins are primitive so far as that condition is concerned but not necessarily so in other ways.
Rhizome-scales indicate a division of the genera into two groups, those with clathrate scales and those
with concolorous scales in which lateral cell-walls are not thickened. (It is notable that Polypodiaceae s.str.
can also be thus subdivided). Bo/bitis (in Malesia) and Lomagramma have (sub)clathrate scales; fronds of
young plants in the two genera also show considerable resemblances. Lomariopsis, Teratophyllum and
Elaphoglossum have non-clathrate scales. The bipinnate fronds of Teratophyllum sect. Polyseriatae may
represent a primitive frond-form. Characters indicating possible cross-relationships between the groups
of genera as defined by scales are: fronds of young plants of Lomariopsis sorbifolia and some allies in
tropical America have much resemblance to those of young plants of Lomagramma; and paraphyses in
Lomagramma guianensis which is of doubtful generic identity are like hairs on the margins of scales of
Lomariopsis. Elaphoglossum is related only to Lomariopsis. I regard the development of a joint at the base
of the lamina as a separate evolutionary development in this genus; it does not occur in all species (one
may compare the orchid genera Liparis and Oberonia, in which some species have jointed leaves and some
have not, the differences being related to the epiphytic condition).
CHRISTENSEN considered the whole group allied to his subfamily Dryopteridoideae (in Verdoorn, Man.
Pterid. 1938, 545); within the subfamily he suggested a possible affinity of Bo/bitis with Tectaria and its
allies. I believe that Pteridrys is an ally of Ctenitis and Tectaria, wrongly placed in the Thelypteris group of
genera by CHRISTENSEN (/.c. 544). The rachis structure and form of attachment of pinnae to rachis are
closely similar in Bolbitis and Pteridrys (HoLTTUM, Rev. Fl. Mal. 2, 1954, 451, 529, fig. 263, 311). The
sinus-teeth of Pteridrys resemble those of free-veined species of Bolbitis (HOLTTUM, /.c. fig. 270, 312); and
B. sinensis (BAKER) K.Iwats. has a frond-form like that of Pleocnemia (sensu proprio; see HOLTTUM,
Reinwardtia 1, 1951, 171-189) which I believe to be closely related to Pteridrys. The rachis structure and
frond-form of Dryopteris are different, and a close relationship of Bolbitis to Dryopteris is unlikely. A closer
study of the whole of CHRISTENSEN’s Dryopteridoideae and the inter-relations of its parts is still needed.
As regards fertile fronds, the acrostichoid condition is clearly derivative, and within the Lomariopsis
group (even within Teratophyllum) there are various ways in which the vascular system is adapted to it.
Kaur (Bot. J. Linn. Soc. 68, 1974, 153) reported that the fertile segments of all lomariopsidoid ferns are
provided with a diplodesmic venation; such a pattern is not present in Bo/bitis (HENNIPMAN, Leid. Bot.
Ser. 2, 1977, 35). ;
The question presents itself: what kind of discrete sori had the ancestors of the group? The little-known
genus Thysanosoria perhaps gives an indication of the answer; it is so like Lomariopsis that the two could
not be distinguished in the absence of fertile fronds. It has narrow fertile pinnae with separate sori at the
vein-ends, each of which enters a small lobe on the pinna-margin, much as in some species of Nephrolepis.
Thysanosoria sori however lack indusia and the sporangia are distributed a little distance from the end of
the vein; a similar spreading has occurred in exindusiate species of many diverse genera which are nor-
mally indusiate (e.g. Tectaria, Stegnogramma). One might therefore postulate an indusiate sorus at a vein-
ending as the original fertile state of the group, but not necessarily a lobed margin with sori in the lobes.
It seems possible that the genus Arthropteris is more closely related to the Lomariopsis group of genera
than is Nephrolepis. Arthropteris resembles Teratophyllum in slender climbing rhizome with dorsiventral
258 FLORA MALESIANA [ser. IT, yoke
structure and two longitudinal rows of fronds on the dorsal surface, also in having the stipes jointed to the
rhizome and pinnae to rachis. The jointing of stipe to rhizome is somewhat more definite than in Terato-
phyllum; there are phyllopodia projecting from the rhizome as in Elaphoglossum and Oleandra. The sori of
Arthropteris are either indusiate (indusia reniform) or not, for which reason Hooker placed one species in
Nephrolepis, one in Nephrodium and one in Polypodium. The apex of the frond is in some species a termi-
nal lobed lamina continuous with the rachis, in others a jointed pinna as in Teratophyllum. The spores
have a perispore. The species of Arthropteris which most nearly resembles Teratophyllum is A. tenella
(Forst.) J.SM. of New Zealand and Australia; young plants of this species have been reported as having
bipinnate fronds (CARsE, Trans. New Zeal. Inst. 47, 1911, 85).
Arthropteris differs from Teratophyllum in bearing abundant short multiseptate hairs, much as in
Ctenitis and Tectaria. Such hairs are also abundant in Davallodes, a genus allied to Davallia, though the
latter has almost glabrous adult fronds in all species (SEN, SEN & HoLttum, Kew Bull. 27, 1972, 217).
Thus Arthropteris shows possible relationships to both Teratophyllum and Davallia; in frond-form of some
species it comes near to Nephrolepis, in others to Crenitis. | conclude that Arthropteris looks like a separate
offshoot from the Crenitis stock, not nearly related to Teratophyllum, and that, with Davallodes, it may
indicate relationships between Crenitis and Davallia.
Spores of Lomariopsis and Teratophyllum were first figured by HoLtrum in 1932 (Gard. Bull. S.S. 5:
252). NAYAR and Kaur have described and illustrated spores of species from all genera in the group
(Pollen et Spores 5, 1963, 87-94; J. Palyn. 1, 1965, 10-26). HENNIPMAN (Acta Bot. Neerl. 19, 1970, 671—
680; Leid. Bot. Ser. 2, 1977, 38-46) studied several species of Bo/bitis including some formerly referred to
Egenolfia with both the transmission and scanning electron microscope. All Malesian species except those
of Lomagramma have large folded perispore; but see note on species of Lomariopsis in Africa and America
under that genus. Representative types of perispores are illustrated in fig. 26.
Gametophytes. NAYAR and KAurR (Bot. Rev. 37, 1971, 345-346) summarize published information on
gametophytes of Bolbitis, Egenolfia and Elaphoglossum. Dr. L. R. ATKINSON (in Jermy et a/., Bot. J. Linn.
Soc., Suppl. 1, 1973, 81) has made observations on gametophytes of four species of Lomariopsis (1
Malesian, 3 African) and one of Lomagramma, also of Bolbitis spp. including the American B. cladorrhizans
(Spr.) CHING (a synonym of B. portoricensis (SPR.) HENNIPMAN). Gametophytes of Elaphoglossum differ
from the other genera in their ribbon-like form with marginal rhizoids; for further comment, see Elapho-
glossum. Gametophytes of Lomariopsis and Lomagramma have not yet been raised to maturity. In all cases
spathulate young gametophytes are formed with a meristem along the anterior margin, no single apical
cell being evident. Gametophytes of Bolbitis portoricensis and of B. (Egenolfia) hookeriana wats. (a
synonym of B. appendiculata (WILLD.) Iwats.) are strap-like, those of the Malesian B. heteroclita (PRESL)
CuinG and B. repanda (BL.) SCHOTT are broadly cordate and have curved multicellular hairs at the notch
and along the anterior margins of the wings; such hairs are lacking in the other species. The sex organs of
the lomariopsidoid ferns are of the so-called advanced type. Data on gametophytes, though not offering
strong supporting evidence, do not contradict the idea of an alliance between the genera Lomariopsis,
Lomagramma and Bolbitis. Gametophytes of Teratophyllum have not yet been studied.
KEY TO THE GENERA
1. Fronds of adult plants simply pinnate or bipinnate.
2. Fronds of adult plants simply pinnate.
3. Veins all free.
4. Pinnae jointed to rachis; rhizome high-climbing with widely spaced fronds.
5. Terminal pinna not jointed to rachis at its base.
6. Fertile pinnae covered beneath with sporangia ............... 1. Lomariopsis
GESOniscpardaterat chUsiOlsvelnS-ntw ole) cael saa- fee Toul 8-8 ctw nia . 2. Thysanosoria
56 emuainalepinna- jointed) at ats. base’ sj p< «eo ss ams a e nay! Bate apes | + yO ee nee aa
AS Pininae-notyointed: thizome SHOT... .- yes). 6 sees cas +o do d,s ee ee ee
3. Veins anastomosing.
7. Pinnae jointed to rachis; high-climbing plants ............ . . 4. Lomagramma
7. Pinnae not jointed; short-creeping plants ................ . ._. 6, Bolbitis
2aBronds ohadult plants, bDipinnate \.c-sea s +s, wy.3ehenies denis ieue>s ori te. 2. se) lb Oee ee De
1. Fronds of adult plants simple.
8. Veins all free or united at margin only; fronds jointed to phyllopodia . . . . . 5. Elaphoglossum
8. Veins freely anastomosing; no joint between frond and rhizome ........ . . 6. Bolbitis
1. LOMARIOPSIS
Fé, Hist. Acrost. (1845) 10, 66-71, p.p.; J.Sm. Hist. Fil. (1875) 139; CHrisT,
Farnkr. Erde (1897) 39, p.p.; HoLtrum, Gard. Bull. S. S. 5 (1932) 264-277; Not.
Syst. 8 (1939) 48-62; Kew Bull. 1939, n. 10 (1940) 613-628; TarD.-BL. & C.CHR.
Fl. Gén. I.-C. 7, 2 (1939) 427; BACKER & Post. Varenfl. Java (1939) 150; COPEL.
1978] LOMARIOPSIS GROUP (Holttum) 259
Gen. Fil. (1947) 117; Philip. J. Sc. 78 (1949) 400; Fern FI. Philip. (1960) 267;
HOLtTTuM, Rev. FI. Mal. 2 (1954) 476; Blumea 14 (1966) 218. — Acrostichum sect.
Lomariopsis HooK. Spec. Fil. 5 (1864) 241, p.p. — Lomariopsis sect. Eulomariopsis
METT. ex KUHN, Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 294. — Stenochlaena sect.
Eustenochlaena Digts in E. & P. Nat. Pfl. Fam. 1, 4 (1899) 251, p.p. — Stenochlaena
sect. Lomariopsis UNDERW. Bull. Torr. Bot. Cl. 33 (1906) 37, 44—-50; v.A.v.R.
Handb. (1908) 718, p.p.; Suppl. (1917) 427, p.p. — Fig. 1, 26d-e.
Rhizome climbing, broad, rooting on ventral surface only and bearing several
rows (to 5 or 6) of fronds on upper surface, densely scaly on younger parts; scales
thin, brown, to 10 by 3 mm, base peltate (or cordate?), edges + fringed with hairs,
lateral cell-walls not thickened; vascular system dorsiventric, showing in transverse
section a broadly U-shaped ventral strand and above this a half-ring of wedge-
shaped bundles with narrow leaf-gaps between them (fig. lc). Stipe gradually
decurrent at base to a ridge on rhizome, free part containing an open ring of c. 10
vascular strands, surface scaly when young; fronds simply pinnate, pinnae entire,
jointed to rachis, terminal lamina pinna-like but not jointed; veins free, usually
uniting with the (non-vascular) cartilaginous margin; surface when young bearing
scattered minute fimbriate scales. Pinnae of fertile fronds much narrower than
sterile, their lower surface completely covered with sporangia and small scales as on
sterile pinnae; spores large, with copious folded perispore. Young plants: rhizome
slender, bearing fronds in 2 rows; in Malesian spp. fronds simple and usually
entire, successively larger to about 30 cm long, later fronds with smaller apical
lamina and an increasing number of pinnae; fertile fronds usually not produced
until the rhizome is of adult size and has climbed 2 m or more above ground.
Type species: Acrostichum sorbifolium L.
Distribution. Throughout the wetter parts of the tropics (America 15 spp., W. Africa & Uganda
10 spp., Islands of Indian Ocean 9 spp., Asia, Malesia, Queensland, and Pacific 10 spp.).
Ecology. Plants of primary evergreen forest; prothalli growing on the ground or on exposed roots of
trees, the slender rhizome creeping until it meets a tree-trunk, up which it climbs to 5—10 m, retaining a
root-system in the ground. As the rhizome grows upwards successive fronds are borne in stronger light
and less humid air. Fertile fronds are produced as a response to drier conditions according to local
climatic change. Pinnae are deciduous but not whole fronds as Teratophyllum. In Luzon M. G. PrRIcE has
found stunted fertile plants creeping on stones in a stream-bed in a semi-exposed position (see L. /ineata).
Morphology. The broad + flattened dorsiventral rhizome, ridged on the dorsal surface with decur-
rent bases of fronds and bearing roots on the ventral surface is exactly as in Lomagramma. Teratophyllum
sect. Polyseriatae differs only in having the bases of stipes a little constricted, not decurrent, and at length
deciduous leaving round scars. Young plants of some species of Lomariopsis in other geographic regions
differ in having fronds which are fully pinnate from an early stage, the terminal lamina never much larger
than the lateral pinnae (e.g. L. sorbifolia and allied species in West Indies; see Ho_trum, 1940). In West
Africa is one species which has simple fronds throughout its life (L. palustris (HooK.) MetTT.). In Mauri-
tius L. variabilis (WILLD.) FEE has the early simple fronds more or less deeply dissected (they are compar-
able to mature sterile fronds of Pe/tapteris). In all cases there is a gradual transition from the frond-form
of young plants to that of adult plants, without sharp distinction between bathyphylls and acrophylls.
Gametophytes have not yet been raised to maturity; for available information see p. 258.
Cytology. The only Malesian species investigated is L. /ineata in cultivation at Kew, from root-tips
(2n = 164, tetraploid with base 41; Roy & MANTON, J. Linn. Soc. Bot. 59, 1966, 343). Plants from West
Africa, also at Kew, gave various different results, and two species also showed unevenness in size of
chromosomes.
Spores. In Malesian species the perispore is + elaborately folded and in some cases is very large
(Nayar, New Phyt. 65, 1966, 235-236); in some species of tropical Africa and America it is not folded, or
is produced into numerous small flattened appendages or spines which need to be examined by modern
techniques. See fig. 26d-—e.
Taxonomy. The genus was founded by FEE in 1845; he included in it both Lomariopsis and Terato-
phyllum of the present treatment (except T. articulatum). JOHN SMITH cited Acrostichum sorbifolium L. as
type species (Hist. Fil. 140, 1875); the choice of L. cochinchinensis FEz by HOLTTUM in 1932, copied by
COPELAND (1947) was therefore illegitimate. Most other authors, if they did not follow Hooker in retaining
260 FLORA MALESIANA {ser. II, vol. 14
Fig. 1. Lomariopsis intermedia (CopeL.) HoLtrum. a. Apex of sterile frond, x 2/,, b. apex of fertile frond,
x ?/3,¢. CS of rhizome, x 2, d. scale from base of stipe, x 4. —L. spectabilis (KUNZE) METT. e. Rhizome
and decurrent bases of stipes, « ?/; (a-d T. G. WALKER 10064, e ENDERT 3785).
1978] LOMARIOPSIS GROUP (Holttum) 261
Fée’s genus in Acrostichum, united it with Stenochlaena J.SM. (1841). Hooker (Spec. Fil. vol. 5) would not
recognize most of FEE’s species as distinct and included nearly all of them (including those now separated
as Teratophyllum) in the single species A. sorbifolium. In Farnkrauter der Erde (1897) Curist included also
in this same species (as Lomariopsis sorbifolia) some Asplenioid ferns which have finely dissected fronds in
their young stages and simply pinnate in adult condition; in this treatment he was uncritically followed by
Bower (The Ferns 3, 1928, 175-176). In Index Filicum (1906, 625-626) CHRISTENSEN attempted to distin-
guish twenty “‘subspecies (vel species?)’’ within Stenochlaena sorbifolia. UNDERWOOD was the first subse-
quent author to attempt to characterize the species (Bull. Torr. Bot. Cl. 33, 1906, 35-50); he recognized
Lomariopsis and Teratophyllum (excl. T. articulatum) of the present work as sections of Stenochlaena.
Ho.ttuM (/.c. 1932) pointed out distinctions between Stenochlaena and the other two genera in spores,
scales, anatomy and venation, and later made observations on Lomariopsis in the islands of the Indian
Ocean (1939) and tropical America and Africa (1940).
KEY TO THE SPECIES
1. Sterile pinnae very coriaceous, drying brownish; fertile pinnae 10-20 mm wide; spores 90-110 um long
excluding perispore.
2. Stalks of largest pinnae 5-20 mm long; apices of sterile pinnae rather abruptly short-acuminate; veins
in sterile pinnae widely spaced and at c. 60° to costa ‘ : 1. L. intermedia
2. Stalks of lowest pinnae always short; apices of sterile pinnae gradually attenuate; veins of sterile pinnae
less conspicuous, closer, at their bases almost at right angles to costa. . . 2. L. subtrifoliata
1. Sterile pinnae not very coriaceous, drying dark olive-green; fertile pinnae rarely to 10 mm wide; spores
c. 45-65 um long excluding perispore.
3. Upper surface of fertile pinnae not over 2mm wide; wing of apere narrow, lacking reticulate
thickening . ; hale aie ee 3. L. kingii
3. Upper surface of fertile pinnae wider; wing ‘of spore ‘otherwise.
4. Sterile pinnae 3-5 cm wide, on young plants at least abruptly contracted to a short-acuminate apex;
ReneS Witt ample 1OIGCd PeriSPOle, 2. 8 pk scp ae Soe ndhcensh Oe dee: op 4. L. lineata
4. Sterile pinnae less than 3 cm wide (usually less than 2 a apex always gradually attenuate; spores
bearing a perispore of many small wings . 5. L. spectabilis
1. Lomariopsis intermedia (CopeL.) HoLtrum, _ ing as follows: pinnae never long-stalked even on
Gard. Bull. S. S. 5 (1932) 270; CopeL. Philip. J.
Sc. 78 (1949) 400. — Stenochlaena intermedia
CopeEL. Philip. J. Sc. 7 (1912) Bot. 67; v.A.v.R.
Handb. Suppl. (1917) 429. — Type: C. K1nc 370,
Ambasi, Papua (MICH; dupl. in BM). — Fig.
la-d, 26d.
Rhizome 15 mm or more wide, densely scaly near
apex; scales to 10 by 3 mm. Stipes to 30 cm long;
frond to 70 cm, pinnae to 10 pairs. Sterile pinnae
18-35 by 21/,-5 cm, rather abruptly short-acumi-
nate, conspicuously stalked (stalks of lowest pinnae
5-20 mm long), thick, rigid and brownish when dry,
veins to 2 mm apart, simple or once forked, at
c. 60° to costa. Fertile pinnae as long as sterile and
similarly stalked, commonly 10-15 mm wide (to
20 mm); spores c. 100 um long excluding perispore
which is very wide, elaborately folded, without
conspicuous reticulate thickening.
Distr. Malesia: eastern half of New Guinea,
Louisiade Archipelago.
Ecol. In forest, at 120-1950 m.
Note. This species is near L. oleandrifolia
(BRACK.) MeETT. of Fiji. The latter has pinna-stalks
21/,-3'/, cm long, and sterile pinnae very abruptly
contracted below the short narrow apex.
2. Lomariopsis subtrifoliata (CopeL.) HoLTTuM,
Gard. Bull. S. S. 5 (1932) 274; Cope. Philip. J.
Sc. 78 (1949) 400; Fern FI. Philip. (1960) 269. —
Stenochlaena subtrifoliata CopeL. Philip. J. Sc. 1
(1906) Suppl. 152; v.A.v.R. Handb. (1908) 721.
(Not Gymnogramma_ subtrifoliata Hook. 1864
which is Lomariopsis brackenridgei CARR.). —
Type: CopELAND 1749, San Ramon, Mindanao
(MICH).
Similar to L. intermedia in rigid sterile pinnae
drying brownish and in wide fertile pinnae, differ-
large fronds; apices of pinnae gradually attenuate;
veins of sterile pinnae closer, starting almost at
right angles to costa.
Distr. Malesia: Philippines (southern Luzon,
Leyte, Samar, Biliran, Mindanao).
EcoL. In forest near streams, at c. 800 m.
Norte. One specimen from Mt Bulusan has
fertile pinnae 8 mm wide, spores 83 um long.
3. Lomariopsis kingii (CoPpEL.) HoLtrum, Gard.
Bull. S. S. 5 (1932) 273; Cope. Philip. J. Sc. 78
(1949) 400. — Stenochlaena kingii CopeL. Philip.
J. Sc. 6 (1911) Bot. 80; v.A.v.R. Handb. Suppl.
(1917) 429. — Type: C. KiNG 285, Ambasi, Papua
(MICH). — Fig. 26e.
Rhizome to 12 mm wide; leaf-gaps in vascular
system up to 6; young parts and bases of stipes of
young fronds densely scaly; scales thin, medium
brown, to 10 by 3 mm, edges irregularly hairy when
young. Sterile pinnae to 20 by 21/.-3 cm, lower ones
on stalks to 5 mm long, base unequally cuneate,
widest in basal half, narrowed gradually to acumi-
nate apex (sometimes a rather abrupt narrowing
near apex), texture firm. Fertile pinnae to 20 cm
long, upper surface 2 mm wide (sporangia spread-
ing beyond edges of lamina may produce an
apparent width of 3 mm or more); spores 50-65 um
long excluding perispore which is not very wide and
has few folds.
Distr. Malesia: Philippines (Mindanao to
S. Luzon), New Guinea; Queensland.
Ecol. In forest, at 600-2000 m.
Note. This species is very near L. brackenridgei
Carr. (from which I cannot clearly distinguish
L. setchellii (MAxon) HoLttum) distributed from
Fiji to Tahiti, which has even narrower and longer
fertile pinnae.
262
FLORA MALESIANA
[ser. IT, vole
4. Lomariopsis lineata (PRESL) HoLTruM, Novit.
Bot. Inst. Prag. 1968 (1969) 9. — Olfersia lineata
Pres_, Tent. Pterid. (1836) 235. — Acrostichum
speciosum PRESL, Rel. Haenk. (1825) 16, non WILLD.
— Type: HAENKE s.n., Philippines (PRC).
L. cochinchinensis FEE, Hist. Acrost. (1845) 66,
t. 26; HoLtrum, Gard. Bull. S. S. 5 (1932) 266;
TARD.-BL. & C.Cur. Fl. Gén. I.-C. 7, 2 (1939) 428;
CopeL. Philip. J. Sc. 78 (1949) 400; HoLtrum, Rev.
Fl. Mal. 2 (1954) 476, f. 279. — Stenochlaena
cochinchinensis (FEE) UNDERW. Bull. Torr. Bot.
Cl. 33 (1906) 46. — Type: GAUDICHAUD s.n. 1827,
Tourane, Cochinchina (P; dupl. in B, K).
L. smithii FEE, Hist. Acrost. (1845) 71, t. 33 f. II
(excl. t. s. rhizome), t. 53 f. Il; HoLtrum, Gard.
Bull. S. S. 5 (1932) 269; CopeL. Fern FI. Philip.
(1960) 268. — Stenochlaena smithii UNDERW. Bull.
Torr. Bot. Cl. 33 (1906) 50. — Type: CuminG 143,
Luzon (P; dupl. in B, BM, K, L, MICH, US).
Stenochlaena abrupta v.A.v.R. Bull. Jard. Bot.
Btzg II, 20 (1915) 24; Handb. Suppl. (1917) 429. —
Lectotype: AMDJAH 119, Pladjoe, Borneo (BO;
dupl. in L, UC).
L. papyracea CoPEL. Philip. J. Sc. 56 (1935) 104,
pl. 1 f. 3; Fern Fl. Philip. (1960) 268. — Type:
COPELAND s.n. 30—8-1932, Mindanao (MICH).
Stenochlaena sorbifolia [non (L.) J.SM.] BEDD.
Ferns Br. India (1866) t. 192; Handb. (1883) 423
puoad f. 254 tantum; v.A.v.R. Handb. (1908) 720,
q-p.
Acrostichum laurifolium [non (PRESL) Hook.]
CuristT, Ann. Jard. Bot. Btzg 15 (1898) 177.
Simple fronds of young plants to 30 by 6 cm,
with stipe to 15 cm; lamina abruptly contracted to
a narrow apex 2 cm long. Fronds of mature plant,
including stipe, to 100 cm long, pinnae to 20 pairs.
Sterile pinnae to 20 by 5 cm, lowest on stalks
5-15 mm long, base narrowly cuneate on _basi-
scopic side, broadly on acroscopic, edges almost
parallel for most of length, -- suddenly contracted
to a narrow caudate apex 2-3 cm long, edges entire,
texture subcoriaceous; upper pinnae often less
abruptly contracted near apex. Fertile pinnae
8-15 cm long, 3—6(-10) mm wide; spores 43-50 um
long excluding perispore which is wide with reticu-
late thickening.
Distr. Tenasserim, S. Thailand, S. Vietnam;
throughout Malesia except E. New Guinea.
Ecol. In wet lowland forest and to c. 1200 m;
stunted fertile plants also observed in Luzon
creeping on stones in seasonally dry stream-bed.
Notes. FEe’s figures of L. smithii (based on
CuMING 143, Luzon) show both fertile and sterile
pinnae with very long stalks, but the specimen of
this collection at Kew has stalks 3-5 mm long. In
Malaya I have found plants growing near together,
one with typically broad sterile pinnae, the other
with pinnae only half as wide. The type specimen of
L. papyracea has a normal sterile frond, with a
contorted and incomplete fertile one having an
apical lamina 16 mm wide; it seems to me im-
probable that this is a normal frond of a distinct
species.
Specimens from Taiwan and Hainan which have
been named L. cochinchinensis should perhaps be
ranked as a distinct species; they are near L. kingii,
having sterile pinnae 1'/, cm wide, not abruptly
contracted towards apex, and very narrow fertile
pinnae.
A specimen consisting entirely of simple fronds,
one of them fertile, was collected by M. G. Pricr
(463) growing on stones in a seasonally dry stream-
bed in a semi-exposed position, in Camarines Norte
Province, southern Luzon. This is the only example
of a simple fertile frond of this species seen by me;
failure to produce pinnate fronds was doubtless a
reaction to the exposed habitat.
5. Lomariopsis spectabilis (KUNZE) Metr. Fil.
Hort. Bot. Lips. (1856) 22; Ann. Mus. Bot. Lugd.-
Bat. 4 (1869) 294, excl. spec. Zippelii; HOLTTUM,
Gard. Bull. S.S. 9 (1937) 141; Tarp.-BL. & C.CHr.
Fl. Gén. I.-C. 7, 2 (1939) 428; Corer. Fern FI.
Philip. (1960) 269 (p.p.?). — Lomaria spectabilis
KuNzZE, Bot. Zeit. 6 (1848) 144. — Acrostichum
spectabile (KUNZE) RaciB. Fl. Btzg 1 (1898) 54,
non ZOLL. — Stenochlaena sorbifolia var. specta-
bilis v.A.v.R. Handb. (1908) 721. — Type: ZoL-
LINGER 395, W. Java (L).
Acrostichum smithii RactB. Bull. Int. Acad. Sci.
Lett. Cracovie, Cl, Math. & Nat. B (1902) 59, non
BAKER. — Stenochlaena raciborskii C.Cur. Ind.
Fil. (1905) 18, 625 (new name for A. smithii
RactiB.); v.A.v.R. Handb. Suppl. (1917) 428. —
Stenochlaena smithii (RactB.) v.A.v.R. Handb.
(1908) 720. — L. raciborskii (C.CHR.) HoLTTuM,
Gard. Bull. S. S. 5 (1932) 272. — Type: cult. Hort.
Bog. origin Moluccas, J. J. SmitH, II.K(V) 23;
II.KCXIV) 41 (BO).
Lomaria variabilis (non WILLD.) BL. En. Pl. Jav.
(1828) 263.
Stenochlaena leptocarpa (non FEE) UNDERW. Bull.
Torr. Bot. Cl. 33 (1906) 47 quoad syn. tantum;
v.A.v.R. Handb. Suppl. (1917) 428, p.p.
L. leptocarpa (non FEE) HoLttrum, Gard. Bull.
S.S. 5 (1932) 270. — Fig. le.
Lamina of simple fronds of young plants to 30 by
1.2 cm, narrowed gradually to apex; pinnae of first
pinnate fronds sessile. Fronds of mature plants
70 cm or more long, pinnae c. 15 pairs. Sterile
pinnae of largest fronds commonly 20 by 1'/, cm,
rarely to 2!/, or 3 cm wide, lowest with stalks to
10 mm long, apices acuminate (not abruptly
narrowed), when dry rigid and brittle, dark olive-
green. Fertile pinnae to 25 cm long, 4-5 mm wide,
stalked as sterile; spores about same size as those of
L. lineata, perispore consisting of many small wings.
Distr. Malesia: Central & South Sumatra, Java,
Bali, Borneo, Moluccas; Celebes?; Philippines?.
Ecol. In forest, at 1250-1500 m.
Notes. Acrostichum smithii RActB. was des-
cribed from a plant brought from the Moluccas
(exact locality unrecorded) by J. J. SmitH. The
fronds of the type had few pinnae, sterile ones to
3 cm wide; a later-collected frond, apparently from
the same plant, has 6 pairs of pinnae less than 2 cm
wide. No other collections are known from the
Moluccas.
Specimens from the Philippines formerly referred
to this species all appear to have narrower fertile
pinnae and I believe they must be regarded as
L. kingii. Bornean specimens referred here are few
and have broad fertile pinnae; their sterile pinnae
are thinner and somewhat broader than those of
specimens from Java but are narrowly acuminate,
not abruptly narrowed near apex as in L. /ineata.
The spores of Kinabalu specimens are somewhat
intermediate between those of L. /ineata and of
1978] LOMARIOPSIS GROUP (Holttum) 263
typical L. spectabilis in form of perispore; but
those of ENDERT 3785 from 1200 m in W. Kutai are
very like spores of Java specimens.
Excluded
Lomariopsis hiigelii PresL, Epim. Bot. (1851)
263 = Blechnum filiforme (A.CUNN.) ETTINGs.
Type: HuGEL, New Zealand in Herb. PRESL, seen;
duplicate in W. (Not Stenochlaena hiigelii UNDERW.
Bull. Torr. Bot. Cl. 33 (1906) 46 which is Terato-
phyllum brightiae (F.v.M.) HoLttrum; see Blumea
14 (1966) 218).
2. THYSANOSORIA
GepP in Gibbs, Dutch N.W. New Guinea (1917) 193, pl. 4; CopeL. Gen. Fil. (1947)
117. — Fig. 2.
Like Lomariopsis Fée in habit and in form of sterile fronds, differing in fertile
pinnae which have a small rounded marginal lobe at each vein-ending, the
sporangia in separate sori, one at the end of each vein and spreading a little back-
wards along the vein, without indusia.
Distribution. One species, known only from two collections from neighbouring localities in NW.
New Guinea, one in 1875, the other in 1913.
Notes. The second collection, as illustrated in Gepp’s plate above cited, shows rhizome and frond-
characters exactly matching Lomariopsis, and I agree with CHRISTENSEN in regarding the two genera as
closely related. Possibly the sori of Thysanosoria show a stage in the evolution of the acrostichoid condi-
tion of Lomariopsis. Alternatively, Dr HENNIPMAN suggests that the two specimens named Thysanosoria
may represent an abnormal condition of a species of Lomariopsis; but the collection of identical material
after an interval of nearly 40 years is against this. The similarity of the fertile pinnae of Thysanosoria to
Nephrolepis was noted by GeppP; but in vegetative habit and in spores the two genera are very different.
1. Thysanosoria pteridiformis (CESATI) C.Cur. Ind.
Fil. Suppl. 3 (1934) 187; Dansk Bot. Ark. 9, 3
(1937) 51; PicHt SERMOLLI, Webbia 32 (1977) 91. —
Gymnogramme pteridiformis CESATI, Rend. Acad.
Napoli 16 (1877) 30. — Notholaena pteridiformis
(CESATI) BAKER, Malesia 3 (1886) 49; v.A.v.R.
Handb. (1908) 484. — Type: Beccari s.n., Andai
(FI, Herb. Beccari 12704; dupl. in K).
T. dimorphophylla Gepr in Gibbs, Dutch N.W.
New Guinea (1917) 193, pl. 4. — Type: L. S. Grpss
6162 (BM). — Fig. 2.
Rhizome as Lomariopsis; young parts covered
with peltate scales. Stipe of sterile frond 7-9 cm
long; pinnae to 8 pairs, jointed to rachis, terminal
lamina pinna-like but not jointed at its base; pinnae
sessile, c. 15 cm long, 1.8—2 cm wide, entire, rather
thin; veins simple or forked near costa, joining to
edge which is pale, slightly thickened and slightly
crisped when dry. Fertile pinnae 10-15 cm long,
6 mm wide when mature (BECCARI’s specimen),
edge with a small rounded projection at the end of
each vein; sori on distal part of each vein, c. 2 mm
apart, hemispherical when young. Spores with
broad winged perispore (width of wing = half
diameter of spore) with a few irregular folds.
Distr. Malesia: NW. New Guinea.
Ecol. “Common, climbing in karang forest’’
(Gipss 6162). Karang is a belt of coral limestone
near the coast.
Notes. The two collections differ in the fol-
lowing ways. BECCARI’s sterile fronds have only
3 pairs of pinnae, those of GipBs 6-8 pairs. BEC-
CARI’s fertile fronds are old, with pinnae 6 mm wide
(Kew specimen; PICHI SERMOLLI reports 7-12 mm),
those of GrpBs young and not over 3 mm wide; the
latter would have been wider when old, but perhaps
not to 6 mm. Sterile plants would look very much
like Lomariopsis kingii, and it is possible that some
Fig. 2. Thysanosoria pteridiformis (CESAT1) C. CHR.
a. Young fertile pinna, < 2/3, b. part of a, upper
surface, showing reflexed sorus-bearing lobes,
x 6, c. old fertile pinna, x 2/3, d. part of c, slightly
enlarged (a—b Gipps 6162, c-d BECCARI s.n. 1872).
FLORA MALESIANA [ser. II, vol. 14
oe vi di ‘iis
Fig. 3. Teratophyllum aculeatum (BL.) METT. ex KUHN. Normal bathyphylls on tree-trunk, Fraser’s Hill,
Malaya (Photogr. R. E. HoLtrum).
1978]
sterile specimens credited to that species are
Thysanosoria. The spores of Thysanosoria are very
much like those of L. kingii.
PicHi SERMOLLI (/.c.) regarded the collections of
BECCARI and GIBBS as representing two distinct
species. The differences he cited include number of
pinnae, but probably young plants of Thysanosoria,
like those of Lomariopsis, have simple fronds, the
number of pinnae in subsequent fronds gradually
LOMARIOPSIS GROUP (Holttum)
265
increasing with decrease in size of the apical lamina;
in some cases fertile fronds with few pinnae are
produced. As regards differences he described in
the fertile pinnae, he failed to note that those on
the GiBBS specimen are young and not fully ex-
panded; the apical part of a Kew pinna from BEc-
CARI is not greatly different from the Grpss speci-
men in width, and the sori on the BECCARI specimen
are not all wholly intramarginal.
3. TERATOPHYLLUM
METT. ex KuHN, Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 296; HoLtrum, Gard. Bull.
S. S. 5 (1932) 277-304; ibid. 9 (1938) 355. — Stenochlaena sect. Teratophyllum
UNDERW. Bull. Torr. Bot. Cl. 33 (1906) 37. — Fig. 3-8.
Rhizome scandent, dorsiventral, bearing on the ventral surface roots only and on
the dorsal surface fronds alternately in 2 rows (sect. Teratophyllum) or at maturity
in several rows (sect. Polyseriatae); bases of stipes swollen and + articulate to
rhizome; rhizome-scales small, peltate, appressed, fugacious, not clathrate. Fronds
of young plants, or on branches of old plants near ground level (bathyphylls) of a
distinctive form characteristic of each species, often bipinnate, with winged rachis
and pinnae jointed to it; fronds of high-climbing rhizomes (acrophylls) larger, simply
pinnate (sect. Teratophyllum) or sometimes bipinnate (sect. Polyseriatae), all pinnae
(including the apparently apical one) subequal and jointed to rachis; veins in
sterile pinnae or pinnules free, simple or forked; scales small, peltate, with marginal
hairs ending in glandular cells; fertile pinnae or pinnules much narrower than
sterile, their lower surfaces covered with sporangia, the lamina thicker than that of
sterile leaflets and containing an additional vascular system close to the surface
which bears the sporangia, the strands of this vascular system often anastomosing;
paraphyses like the small scales on sterile frond but with a stalk of 5-8 cells; spores
with folded perispore.
Distribution. Peninsular Thailand and Lower Burma; throughout Malesia to N. Queensland, east-
wards to Tahiti and south to New Caledonia.
KEY TO THE SECTIONS
1. Fronds always in 2 rows on dorsal! surface of rhizome; all acrophylls simply pinnate. Spp. 1-9
1. Sect. Teratophyllum
1. Fronds of fully developed rhizome in more than 2 rows; acrophylls bipinnate or pinnate (in Malesian
sp. 10 bipinnate) 2. Sect. Polyseriatae
1. Section Teratophyllum
Ho.trum, Gard. Bull. S. S. 9 (1938) 355. — Stenochlaena J.Sm. in Hook. J. Bot. 3
(1841) 401; ibid. 4 (1841) 149, p.p.; PRESL, Epim. Bot. (1851) 165, p.p.; J.SM. Hist.
Fil. (1875) 312, p.p.; BEpp. Handb. Ferns Br. India (1883) 423, p.p. — Stenochlaena
sect. Eustenochlaena Diets in E. & P. Nat. Pfl. Fam. 1, 4 (1899) 251, p.p. —
Stenochlaena sect. Teratophyllum UNDERW. Bull. Torr. Bot. Cl. 33 (1906) 37, 40, 41;
v.A.v.R. Handb. (1908) 722; Suppl. (1917) 428. — Lomariopsis FEE, Hist. Acrost.
(1845) 71, p.p.; J.SM. Hist. Fil. (1875) 139, p.p.; CHRIsT, Farnkr. Erde (1897) 39,
p.p. — Acrostichum sect. Lomariopsis Hook. Spec. Fil. 5 (1864) 242, p.p.; sect.
Stenochlaena I.c. 249, p.p. — Acrostichum L. in Racib. Fl. Btzg 1 (1898) 53, p.p. —
Teratophyllum METT. sensu HOLTTUM, Gard. Bull. S. S. 5 (1932) 277; ibid. 9 (1937)
266 FLORA MALESIANA [ser. Tl, vole?
142; Rev. Fl. Mal. 2 (1954) 470; C.Cur. Ind. Fil. Suppl. 3 (1934) 186; BACKER &
PostH. Varenfl. Java (1939) 151; CopeL. Gen. Fil. (1947) 117; Fern Fl. Philip.
(1960) 269.
Type species: Lomaria aculeata BL.
Distribution. Mergui and Peninsular Thailand; throughout Malesia, except Lesser Sunda Is.; 9 spp.
Ecology. Confined to primary forest. Prothalli grow on exposed tree-roots or bases of buttresses;
young sporophytes develop a slender rhizome which usually grows upwards and may branch (in T. /udens
it produces long trailing branches which pass along the ground from one tree to another) bearing
successively larger and more complex bathyphylls, the earlier ones often asymmetric, with lamina more
fully developed on lower side, with a more or less abrupt transition to acrophylls at c. 2m above ground
level. Fertile fronds produced at 3-5 m or more above ground, seasonally (probably in response to short
periods of drier weather, which in many parts of Malesia are of irregular occurrence). A copious root-
system develops on the lower parts of the climbing stem, spreading in the soil, and I believe this to provide
most of the water needed by the plant. KARSTEN, believing Teratophyllum plants to lack such a soil-
penetrating root-system, suggested that the chief function of bathyphylls is water-absorption (Ann. Btzg
12, 1895, 143-150). Bathyphylls can absorb some water, but not enough to be of much importance; and
they do not always lie in close contact with the bark of the supporting tree, as described by KARSTEN
(see fig. 3).
Vegetative morphology. See Hottru, Gard. Bull. S. S. 5 (1932) 277-283. Bathyphylls were for-
merly considered to be abnormal growths, but they are quite normal, and are distinctive in each species,
showing a series of forms from the simple lamina of the first fronds to the pinnate (often bipinnatifid)
condition of normal bathyphylls. In the pinnate condition, the lower pinnae are jointed to the rachis but
the distal ones are not, becoming merged in an apical lamina. By contrast, all pinnae of acrophylls are
jointed at their bases and are caducous; the apparently apical pinna is probably a lateral one which takesa
terminal position, the true apex being suppressed. In some cases transition bathyphylls are found, with
pinnae much as in normal bathyphylls but all pinnae jointed; I have not seen intermediate stages between
these and normal bathyphylls. It seems possible that transition bathyphylls are borne on strong new
branches arising near the base of an old plant.
The adult rhizome has always a gutter-shaped root-bearing ventral vascular strand, and a smaller
dorsal one (fig. 6j); fronds are borne on the dorsal side, alternately to right and left of the dorsal strand.
The arrangement of vascular strands in the stipe is like that on Dryopteris and Tectaria (fig. 6k). The very
narrow fertile pinnae are fleshy, with a special development of vascular strands near the convex sporangia-
bearing surface (HOLTTUM, /.c.; NAYAR, New Phyt. 65, 1966, 221-239); in the narrower pinnae of Terato-
phyllum aculeatum and other species these strands anastomose rather irregularly, but in the broad fertile
pinnae of 7. rotundifoliatum they do not. Probably such independent vascular supply for the sporangia
has evolved independently in various unrelated genera as a necessary adjunct to the acrostichoid state. The
fertile pinnae show no indication of an indusium-like thin edge, such as occurs in Stenochlaena.
Cytology. The only observation is from roots of a plant of 7. /udens in cultivation at Kew, showing
212
Taxonomy. The two species native in Java were first described by BLUME (1828) in the genus Lomaria;
he saw bathyphylls and thought they were fertile fronds. JoHN SMITH, when enumerating CUMING’s
Philippine ferns (1841) established a new genus Stenochlaena, and stated, on the evidence of CUMING’s
specimens, that Stenochlaena scandens (now known as S. palustris (BURM. f.) BEDD.), the type-species of
Stenochlaena, sometimes produced abnormal bipinnate fronds. These fronds were bathyphylls of Terato-
phyllum aculeatum, which were associated with normal sterile and fertile fronds of Stenochlaena palustris
by CuMING under his n. 347, as shown by specimens in JOHN SmiTH’s herbarium (BM), one annotated by
him. This idea persisted with SMITH and is repeated in his last book (Hist. Fil. 1875); it was also accepted
by Hooker (Spec. Fil. 5: 250) and by BEDDoME (Handb.) who so named a specimen of T. aculeatum bearing
bathyphylls collected by WALLICH in Penang in 1822. WALLIcH had given the name Davallia achillaeifolia
to this specimen, and the name was formally published, with a figure, by Hooker (Spec. Fil. 1: 195), with
doubts expressed as to the affinity of the fern and a reference to its resemblance to Lomaria aculeata BL.
BAKER named similar ferns from Moulmein Lindsaea parishii in 1867. Even as late as 1929 COPELAND
wrote of “the polymorphism of the fronds of immature plants’’ of Stenochlaena palustris (Univ. Cal.
Publ. Bot. 16: 75). Sporeling plants of S. palustris are very rare in Malaya, though mature plants are
abundant and frequently fertile. I have however raised young plants from spores. Their first fronds were
simple, then simply pinnate, showing from the first the leaflet-form and venation of Stenochlaena, quite
unlike Teratophyllum. I can confidently assert that no true Stenochlaena produces fronds which could be
confused with the bathyphylls of Teratophyllum. It should be noted that young plants of Asplenium
epiphyticum Cope. also produce bathyphylls. These were confused with those of species of Teratophyllum
by Curist, who concluded that “‘Stenochlaena’”’ was an acrostichoid derivative of Asplenioid origin;
CoPELAND accepted this derivation as “absolutely clear’’ in 1929. (See CurisT, Philip. J. Sc. 2, 1907, Bot.
166; also his earlier composite fig. 96, p. 40, under Lomariopsis sorbifolia, which included fronds of
Lomariopsis, Teratophyllum and Asplenium, in Farnkr. der Erde, a figure copied without question by
Bower in The Ferns 3, fig. 697).
In his monograph of the acrostichoid ferns (1845) Fér did not mention Stenochlaena palustris. He
included the species of Teratophyllum sect. Teratophyllum known to him in his new genus Lomariopsis; the
1978] - LOMARIOPSIS GROUP (Holttum) 267
species were L. spinescens (Lomaria aculeata BL.), L. leptocarpa (based on CUMING 132 from Luzon) and
L. ludens (based on a WALLICH specimen from Singapore). Of these, the first was known only from bathy-
phylls and sterile acrophylls, the second from sterile and fertile acrophylls, the third only from bathyphylls.
Hooker later included all these, and all true Lomariopsis specimens known to him, in Acrostichum
sorbifolium L. (Spec. Fil. 5, 1864, 242), a species now regarded as confined to tropical America.
METTENIUs was the first to recognize Teratophyllum as a genus (1869; posthumous work edited by
Kunn); he included in it two species, one in each of the sections here recognized (he included Stenochlaena,
as a section, in Lomariopsis). He included all specimens of sect. Teratophyllum in the species T. aculeatum.
UNDERWOOD clearly distinguished for the first time between Stenochlaena, Lomariopsis and Teratophyl-
/um, ranking all as sections of Stenochlaena (Bull. Torr. Bot. Cl. 33, 1906). But he failed to recognize the
great differences between Stenochlaena proper and the other sections, as he did not examine spores, scales,
or vascular anatomy, and failed to notice the ‘‘glands’’ at the bases of pinnae (including basal reduced ones)
in Stenochlaena; he also failed to notice that Teratophyllum differs from Lomariopsis in having all pinnae
jointed to the rachis, including the apparently terminal one. UNDERWooD did not recognize the close
relationship between sect. Teratophyllum and sect. Polyseriatae, remarking only that the latter (as Arthro-
botrya J.SM.) appeared to be a valid distinct genus.
The present account is based on that of Hottrum in Gard. Bull. S. S. 5 (1932) 277-304, with some
additional material, especially that recently collected by M. G. Price in the Philippines.
KEY TO THE SPECIES
Based partly on bathyphylls
1. Pinnae of normal bathyphylls (excluding youngest stages) deeply lobed.
2. All pinnae of sterile acrophylls almost sessile.
3. Bathyphyll pinnae to 5 mm wide, pinnules with few segments . . . . 1. T. aculeatum
3. Bathyphyll pinnae to 15 mm wide, pinnules with several segments on each side . . 2. T. gracile
2. Lower pinnae of sterile acrophylls with stalks at least 6 mm long, often longer.
4. Pinnae of bathyphylls deeply dissected to linear lobes.
5. Ultimate lobes 1/;mm wide .. . hts Pans Phra ee Se leptocarpum
5. Ultimate lobes much wider. . . 6. T. arthropteroides
4. Pinnae of bathyphylls less deeply dissected, ultimate lobes almost circular.
6. Fertile pinnae 3 mm wide... . . PRS Re A wer.. eee 24: 6 Elazonicum
6. Fertile pinnae 10 mm wide . . . 5. T. rotundifoliatum
1. Pinnae of bathyphylls (excluding youngest stages) crenate, ‘not deeply lobed.
7. Most pinnae of sterile acrophylls, and of bathyphylls, very asymmetric at base.
8. Largest sterile acrophyll pinnae to 9 by 2 cm; basal acrophyll pinnae with almost symmetric base,
almost all other pinnae strongly asymmetric; fertile pinnae 4*/,-5'/, cm long _6. T. arthropteroides
8. Largest sterile acrophyll pinnae to 15 by 2.8 cm; basal acrophyll Ene strongly async at base,
distal ones much less so; fertile pinnae 20-24 cm long . ae 7. T. koordersii
7. All pinnae almost symmetric at base, both acrophylls and bathyphylls..
9. Early stages with lamina on one side of midrib only; small transition bathyphylls with all pinnae on
one side of rachis common; fertile pinnae to 20 cm by 3 mm, stalks to 5 mmlong . . 8. T. ludens
9. Earliest stages with lamina on both sides of midrib; normal and transition bathyphylls all with
pinnae on both sides of rachis; fertile pinnae 5-7 cm by 5 mm, stalks 10-15 mm long
9. T. clemensiae
KEY TO THE SPECIES
Based on acrophyll characters only
1. Pinnae of sterile acrophylls sessile or nearly so.
PSteilespinnace2 24/7 CIV WIGewn "meats ree. sree. Ee . 1. T. aculeatum
Derstenlespinnac 11/4, CM Wide + sae Vomn kn .. STEN CE oe cee 1. T. aculeatum var. montanum
2. T. gracile
1. Pinnae of sterile acrophylls, at least lower ones, stalked.
3. Bases of some sterile pinnae conspicuously asymmetric.
4. Stalks of sterile pinnae 3-5 mm long.
5. Basal pinnae with almost symmetric base, base of others strongly asymmetric; fertile pinnae
41/,-5'/,cm by 5-7 mm . . . 6. T. arthropteroides
5. Basal pinnae strongly asymmetric at ‘base, bases of F upper pinnae I less sO; fertile pinnae 20-24 cm
by 2!/,-3 mm. . ah 4 . . . 7. T. koordersii
4. Stalks of sterile pinnae 5-10 mm long ae fats) mene dae dees eS ig elemearase
3. Bases of sterile acrophyll pinnae symmetric or nearly so.
6. Pinnae of sterile acrophylls to 20 by2cm .. . . 3. T. leptocarpum
6. Pinnae of sterile acrophylls not over 16 cm long, proportionately ‘wider.
268 FLORA MALESIANA [ser. Il, vou?
(@revaleisce
Fig. 4. Teratophyllum aculeatum (BL.) METT. ex KUHN. a. Apex of sterile frond, x 2/3, b. pinnae of fertile
frond, x 2/3, c. scale from rhizome, x 4, d. scale from bathyphyll, x 80, e. earliest stage, f. dimidiate
normal bathyphyll, both nat. size, f’. part of f, x 2, g. later stage but pinnae less deeply lobed, A. mature
normal bathyphyll, both nat. size, h’. part of h, x 2. — T. gracile (BL.) HoLtrum. i. Apex of transition
bathyphyll, nat. size, i’. part of i, x 2 (a-b KING’s Collector 10005, c-d DoNK s.n., e KOSTERMANS 6163,
f Donk 73, g MATTHEW s5.n., 28 Jan. 1908, h LORZING 12257, i WINCKEL 1713).
1978]
7. Stalks of sterile pinnae to 5mm long .
LOMARIOPSIS GROUP (Holttum)
269
8. T. ludens
7. Stalks of sterile pinnae to at least 10 mm long.
8. Sides of sterile pinnae parallel except in distal /,; fertile pinnae 3 mm wide .
8. Sides of sterile pinnae not parallel; fertile pinnae c. 10 mm wide .
1. Teratophyllum aculeatum (BL.) METT. ex KUHN,
Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 296, excl. syn.
and var. inermis; Ho_tTruM, Gard. Bull. S. S. 5
(1932) 284, f. 32, 38-40, 45, pl. 2, 3; ibid. 9 (1937)
144; Rev. Fl. Mal. 2 (1954) 473, f. 276; BACKER &
PostH. Varenfl. Java (1939) 151, f. 29; CopEL.
Philip. J. Sc. 78 (1949) 400; Fern Fl. Philip. (1960)
270. — Lomaria aculeata BL. Enum. Pl. Jav. (1828)
205. — Stenochlaena aculeata (BL.) KUNZE, Bot.
Zeit. 6 (1848) 142; KaArsTEN, Ann. Jard. Bot. Btzg
12 (1895) 143, t. 14, 15; Curist, Philip. J. Sc. 2
(1907) Bot. 166, p.p.; v.A.v.R. Handb. (1908) 722,
p.p.; Suppl. (1917) 430, excl. var. crassior. —
Olfersia aculeata BL. Fi. Jav. Fil. Suppl. (1883?)
t. 96. — Acrostichum aculeatum (BL.) RActsB. FI.
Btzg 1 (1898) 53, non LINN., nec Desv. — Type:
BLUME s.n., W. Java (L).
Lomaria polymorpha ZoLv. & Mor. apud Zou.
Nat. Geneesk. Arch. N. I. 2 (1845) 204; Hassk.
Flora 30 (1847) 319. — Type: ZOLLINGER 2303,
Java (BM, BO, L, P).
Davallia achilleifolia WALL. ex Hook. Spec. Fil.
1 (1846) 195, t. 56D; PReEsL, Epim. Bot. (1851) 263.
— Acrostichum (Stenochlaena) scandens Hook.
Spec. Fil. 5 (1864) 250 quoad WALLICH 248 tantum.
— Type: WALLICH 248, Penang (K).
Lindsaea parishii BAKER, Syn. Fil. (1867) 109;
BepD. Ferns Br. India (1866) t. 209. — Stenochlaena
palustris BEDD. Handb. Ferns Br. India (1883) 422
quoad F.B.I. t. 209 tantum. — Type: ParisH 196,
Tenasserim (K). — Fig. 3, 4a—-h.
1. var. aculeatum.
Fronds on young plants with lamina wholly on
one side of midrib, lobed towards the base, later
fronds still wholly dimidiate but with lower lobes
(to c. 20) separate as + lobed articulated pinnae to
9 by 6 mm (fig. 4f), basal one deflexed across
rhizome. Normal bathyphylls sessile, glabrous apart
from minute scales when young, commonly c. 9 by
2 cm (to 12 by 5 cm); pinnae to 5 mm wide, in most
cases deeply lobed to a winged costa, basal one
deflexed and overlapping rhizome, all jointed to
rachis except distal ones which merge with apical
lamina. Transition bathyphylls variable, some with
deeply lobed pinnae (lobes linear), pinnae of others
grading to an almost or quite entire condition
12-16 by 5-8 mm with acute or rounded apices.
Adult rhizomes 4-6 mm diameter, glabrescent
except near apex, bearing scattered spines 1-3 mm
long; scales near apex abundant, to 2 mm long,
narrow, medium brown, base peltate, edges bearing
scattered hairs. Stipes of acrophylls 3-7 cm long,
lamina c. 40 cm long with 15 pairs pinnae; sterile
pinnae sometimes drying reddish (on midrib or
throughout), almost sessile (in Luzon lowest pin-
nae sometimes stalked 3-5 mm), 9-10(-15) cm
long, 2—21/,(—3) cm wide, base almost symmetric, of
lower pinnae sometimes subtruncate, of others
broadly cuneate, edges parallel in basal half, apical
half tapering, edges minutely sinuous, texture thin,
veins distinct, those near pinna-apex ending freely
before reaching margin and bearing on both sur-
faces very small brown fringed scales (abundant on
. 4. T. luzonicum
5. T. rotundifoliatum
young fronds). Fertile pinnae to 15 cm long, hardly
2 mm wide.
Distr. N. to Mergui, throughout Malesia except
the Lesser Sunda Is.
Ecol. Lowland forest, in moist places; young
plants on exposed tree-roots or bases of trunks,
climbing to 5 m or more.
2. var. montanum HoLtrum, Gard. Bull. S. S. 5
(1932) 289, pl. 1, 4; Rev. Fl. Mal. 2 (1954) 473. —
Type: HoL_ttum 23364, Malaya (K, S).
Differs from var. aculeatum as follows: normal
bathyphylls often larger (to 16 cm long) with pinnae
to 25 by 6 mm, pinnules to 31/, by 2!/, mm with
forked apex and 2-3 narrow lateral lobes; transi-
tion bathyphylls more common and larger; sterile
pinnae 8-15 mm wide, sometimes drying reddish.
Distr. Malesia: Central & North Sumatra,
Malay Peninsula.
Ecol. In forest, in valleys at c. 1200-1500 m.
Notes. This is somewhat intermediate between
T. aculeatum and T. gracile. 1 am not sure that a
sharp line can be drawn between var. aculeatum
and var. montanum, but extreme forms of the latter
are quite distinct. Most specimens from Taiping
Hills have pinnae with narrowly cuneate bases. A
specimen from North Sumatra (VAN STEENIS 9724,
Mt Kemiri, 1400 m) has bathyphylls intermediate
between var. montanum and T. gracile, and sterile
acrophylls like var. aculeatum, not narrow as in
T. gracile. ¢
Aberrant large normal bathyphylls of two
distinct types are borne by a few specimens. (a)
Fronds 17 cm long, larger pinnae 31/, by 1 cm,
segments partly webbed as in many transition
bathyphylls, upper pinnae gradually smaller and
apical lamina not jointed to rachis (KEHDING 3245,
Sumatra, P; ZOLLINGER 2303, Java, L, P). (b)
Fronds to 14 cm long, largest pinnae 4 by 1.7 cm,
pinnules bipinnatifid (AHERN’s collector 2695,
Luzon, P, US; WARBURG s.n., Java, P).
2. Teratophyllum gracile (BL.) Hottrum, Gard.
Bull. S. S. 5 (1932) 291, pl. 5; BACKER & POSTH.
Varenfl. Java (1939) 152. — Lomaria gracilis BL.
En. Pl. Jav. (1828) 205; Hook. Spec. Fil. 3 (1860)
36. — Stenochlaena gracilis KZE, Bot. Zeit. 6 (1848)
142; Fée, Gen. Fil. (1852) 78; UNDERW. Bull. Torr.
Bot. Cl. 33 (1906) 41; v.A.v.R. Handb. Suppl.
(1917) 430. — T. aculeatum METT. ex KUHN, Ann.
Mus. Bot. Lugd.-Bat. 4 (1869) 296, p.p. — Olfersia
gracilis BL. Fl. Jav. Fil. Suppl. (1883?) t. 96. —
Acrostichum gracile Racts. Fl. Btzg 1 (1898) 53. —
Stenochlaena aculeata sensu v.A.v.R. Handb.
(1908) 722, p.p. — Type: BLuME, W. Java (L). —
Fig. 4i.
Normal bathyphylls to ce. 12 by 4 cm; pinnae to
12 pairs, to 21/, by 1 cm, pinnate, tapering to apex;
largest pinnules 5 mm long, consisting of 2-3 pairs
of very narrow lobes (sometimes forked) joined by
a very narrow wing. Transition bathyphylls to 18 cm
long; pinnae 12-15 pairs, to 4 by 1?/, cm, with 12
pairs of pinnules; pinnules pinnatisect, largest with
4 pairs of linear lateral lobes 2 mm long, some
270
FLORA MALESIANA
[ser. II, vol. 14
forked, ultimate divisions 0.3 mm wide. Sterile
acrophylls as T. aculeatum but pinnae 1-1'/, cm
wide, dark red when dry. Fertile pinnae to 10 cm
long, 1-2 mm wide.
Distr. Malesia: West Java.
Ecol. In mountain forest, 1000-1500 m.
3. Teratophyllum leptocarpum (FEE) HoOLTTuUM,
Gard. Bull. S. S. 9 (1937) 143; CopeL. Fern FI.
Philip. (1960) 271. — Lomariopsis leptocarpa FEE,
Hist. Acrost. (1845) t. 29. — Stenochlaena sorbi-
folia ssp 5 C.Cur. Ind. Fil. (1906) 625, p.p. — Type:
CumING 132, Philippines (orig.?; dupl. in B, BM,
US).
Stenochlaena williamsii UNDERW. Bull. Torr.
Bot. Cl. 33 (1906) 41; Curist, Philip. J. Sc. 2 (1907)
Bot. 167. — Stenochlaena gracilis var. williamsii
v.A.v.R. Handb. Suppl. (1917) 431. — T. williamsii
Hottrum, Gard. Bull. S. S. 5 (1932) 292, pl. 6. —
Type: WILLIAMS 684, Mt Mariveles, Luzon (MICH,
(NY).
Smallest bathyphylls seen 2 cm long, pinnate as
normal bathyphylls (not wholly dimidiate as in
T. aculeatum). Normal bathyphylls to 6 cm long,
pinnate to base on lower side of rachis, not quite
to base on upper side; pinnae to 2 by 1.8 cm, with
4-5 pairs of oblique lateral lobes 3-5 mm long, each
lobe pinnatisect with 2-4 linear divisions less than
1/, mm wide; basal pinnae deflexed, overlapping
rhizome. Transition bathyphylls to 12 cm long;
pinnae to 4 by 11/, cm, lobed as normal bathyphylls
but with larger lobes which are more widely spaced;
other specimens with lobes of pinnae + coalescent,
pinnae of the extreme form almost sessile with
finely crenate edges and broad bases. Adult rhizome
4-5 mm @, finely thorny. Sterile acrophylls with
stipes to 20 cm long, lamina 45 cm; pinnae all
stalked (stalks of lower ones 6-9 mm), 10-20 cm
long, 1-2 cm wide, base slightly unequal, of lower
pinnae narrowly cuneate, of upper ones wider,
apex long-acuminate, texture rather thin, all parts
turning reddish on drying. Fertile pinnae to 25 cm
long, c. 2: mm wide.
Distr. Malesia: Philippines (Luzon, Sibuyan,
Samar, Panay, Basilan).
Ecol. In forest, especially near rivers, 100-
600 m.
Notes. Curist (/.c. 1907) refers to specimens
having ‘“‘secondary leaves’’ with a tendency ‘“‘to
present auricles at the anterior base of the pin-
nules’’; these are young plants of an Asplenium,
probably A. epiphyticum CoPeL.
The specimen of CUMING 132 at Paris is Loma-
riopsis lineata and is certainly not the one figured
by FEE.
4. Teratophyllum luzonicum Ho.ttum, Gard. Bull.
S. S. 5 (1932) 297, pl. 9; ibid. 9 (1937) 142; CoPEL.
Fern Fl. Philip. (1960) 270. — Type: F. Fenix BS
28272, Apayao Prov., Luzon (BO; dupl. in US).
Bathyphylls very like those of T. rotundifoliatum
(fronds 11/, cm long not dimidiate); transition
bathyphylls bipinnate, larger pinnules with one
rounded or slightly bifid lobe on acroscopic side.
Adult rhizome c. 4 mm @, somewhat spiny.
Sterile acrophylls to 60 cm long including stipe,
with 8 or more pairs of pinnae; pinnae to 15 by
3 cm, on stalks to 12 mm long, base broadly and
slightly unequally cuneate, sides nearly parallel
except in distal third, apex acuminate, texture firm,
drying green, veins near midrib 11/,-2 mm apart.
Fertile acrophylls with pinnae to 10 cm long and
3 mm wide, on stalks to 5 mm long.
Distr. Malesia: Philippines (Luzon, Palawan,
Mindanao).
Ecol. At 600-700 m (JAcoss 7944, Sierra Madre
Mts; sterile and fertile acrophylls).
Note. No bathyphylls were collected in associa-
tion with the original specimens. I subsequently
described bathyphylls from Palawan, some of
which were associated with sterile acrophylls.
There are also transition bathyphylls which in 1937
I thought represented 7. clemensiae, but now I
think it more likely that they also belong to T. /uzoni-
cum. A final proof can only come from a collection
of all stages from the same place. The fertile fronds
of T. luzonicum and T. clemensiae are very different.
There are also bathyphylls, without acrophylls, ina
collection by LoHER from Mt Mariveles; these
agree well with Palawan specimens.
5. Teratophyllum rotundifoliatum (R.BONAPARTE)
Ho.ttum, Gard. Bull. S. S. 5 (1932) 294, pl. 7, 8,
f. 36, 37, 44; Rev. Fl. Mal. 2 (1954) 474, f. 277. —
Stenochlaena rotundifoliata R.BONAP. Notes Ptérid.
14 (1923) 58. — Type: HoLtTuM 9384, G. Lambak,
Johore (P; dupl. in SING).
Stenochlaena aculeata var. crassior V.A.V.R.Bull.
Jard. Bot. Btzg II, 23 (1916) 20; Handb. Suppl.
(1917) 430. — Type: AJoEB 387, Bencoolen, S.
Sumatra (BO). — Fig. 6f-k.
Earliest stage as 7. clemensiae (not dimidiate as
in T. aculeatum), the entire fronds soon succeeded
by bipinnatifid normal bathyphylls up to 8 by
31/, cm, with closely-placed pinnae to 15 by 4 mm
with almost circular pinnules, texture firm, usually
drying light green. Transition bathyphylls to 15 by
6 cm, with up to 20 pairs of pinnae more widely
separated than in normal bathyphylls, pinnae to
30 by 4mm, coriaceous, deeply lobed, lobes c. 2mm
wide, apices rounded, retuse or bilobed. Adult
rhizome to 8 mm @, strongly aculeate, when young
densely covered (also young fronds) with small
red-brown scales, largest scales seen 7 by | mm.
Sterile acrophylls to 60 cm long including stipe,
with 12 pairs of pinnae; all pinnae stalked (stalks
of lowest 10-15 mm), commonly to 15 by 3 cm
(largest seen 16 by 6 cm), widest below middle,
gradually tapering to apex, base rather broadly and
almost equally cuneate, coriaceous, drying light
green. Fertile pinnae to 10 by 1 cm, on stalks to
18 mm long; tips of veins not connected by a sub-
marginal vein.
Distr. Malesia: Central & South Sumatra,
Malay Peninsula, Borneo.
Ecol. In forest on hillsides (not in swamp-forest),
at 150-1000 m.
Note. The only known fertile specimens were
collected by WRAY (679, K, SING) in Perak; they
are young, with immature sporangia. This species
differs from T. clemensiae in the broader fertile
pinnae without marginal vein, coriaceous sterile
pinnae, and in invariably bipinnatifid bathyphylls
after the earliest stages. Spores have not been seen.
6. Teratophyllum arthropteroides (CHRIST) HOLT-
TuM, Gard. Bull. S. S. 5 (1932) 303, pl. 12; CoPEL.
Fern Fl. Philip. (1960) 271. — Stenochlaena
1978]
LOMARIOPSIS GROUP (Holttum)
271
arthropteroides CuHrRist, Bull. Herb. Boiss. II, 6
(1906) 998; Philip. J. Sc. 2 (1907) Bot. 167;
v.A.v.R. Handb. (1908) 719. — Type: LoHeErR s.n.
April 1906, Rio Ampailit, Mt Makiling, Laguna
Prov., Luzon (P). — Fig. 5a-c.
On all bathyphylls seen (smallest 4 cm long) all
pinnae articulate; largest seen 12 cm long with 6
pairs pinnae and stipe nearly 2 cm long; pinnae 10
by 5 to 30 by 12 mm, base narrowly cuneate on
basiscopic side, broadly on acroscopic, edges
crenate, apex blunt; on one collection most pinnae
deeply lobed. Adult rhizome smooth. Sterile
acrophylls to 40 cm long; rachis usually reddish
when dry; pinnae stalked (stalk of lowest 3-4 mm),
5-9 cm long, 1'/,-2 cm wide, basal pinnae with
symmetric base, rest with base very unequal
(basiscopic narrowly cuneate, acroscopic sub-
truncate), tapered gradually from widest part near
base to acuminate apex, texture thin, edges in most
cases distinctly crenulate (one crena to each vein-
end). Fertile fronds shorter than sterile, pinnae
41/,-51/, cm long, 5—7 mm wide.
Distr. Malesia: Philippines (Luzon, Mindanao).
Ecol. In forest, at 250-400 m.
Note. I have not seen the youngest stage of this
species. Small transition bathyphylls are very like
those of T. koordersii but thinner, and those seen
have a distinct stipe so that basal pinnae do not
overlap the rhizome.
7. Teratophyllum koordersii HoLTrum, Gard. Bull.
S. S. 5 (1932) 301, f. 48, 49, pl. 11. — Type:
Koorpers 17065, Minahassa, N. Celebes (BO). —
Fig. 5d.
Youngest stage: fronds up to 3 cm long dimidiate
with 3 free pinnae all on lower side of rachis (rarely
1 leaflet or lobe on upper side); fronds 3 cm long
and larger dimidiate with jointed apical pinna,
pinnae 4, to 10 by 3-5 mm, base asymmetric, edges
crenate to deeply lobed; /ater bathyphylls with
pinnae both sides, on a frond 7'/, cm long apical
pinna 20 by 8 mm, lateral pinnae 6 pairs, crenate,
decreasing downwards, lowest 13 by 6 mm, base
narrowly cuneate on basiscopic side, broadly on
acroscopic. Intermediate fronds 14 cm long, pinnae
8-9 pairs, apical one 3.8 by 1.4 cm, edges almost
entire. Adult rhizome 4-5 mm @, aculeate. Sterile
acrophylls: stipe 5'/, cm, lamina 45 cm, pinnae
11-12(-20?) pairs, to 15 by 2.8 cm, lowest stalked
5 mm, basiscopic base narrowly cuneate, acro-
scopic broadly rounded, upper pinnae with more
symmetric bases. Fertile fronds: pinnae 20-24 cm
long, 2'/.-3 mm wide, stalks to 5 mm long.
Distr. Malesia: N. Celebes, Philippines (Luzon).
Ecol. In moist forest, low altitude.
Note. The above description is taken in part
from specimens collected in Luzon by M. G. PRICE
(nos 927, 928, 940, 958, 979, 981). These agree
closely with the Celebes type and two other speci-
Fig. 5. Teratophyllum arthropteroides (CHRIst) HoLTTuM. a. Upper pinnae of sterile acrophyll, b. bathy-
phylls, c. fertile pinnae. — T. koordersii HoLTTUM. d. Bathyphylls. All < 2/; (a ELMER 18353, b—c CoPE-
LAND PPE 251, d Koorpers 17064).
212 FLORA MALESIANA [ser. II, vol. 14
J
te
t
px
4
oN
t
if
Fig. 6. Teratophyllum ludens (FEE) HoLTTum. a. Pinnae of sterile acrophyll, b. bathyphyll, early stage,
c. bathyphyll from creeping rhizome, d. bathyphyll from climbing rhizome, all 2/3, e. scale from base of
stipe, x 40. — T. rotundifoliatum (R. BONAP.) HoLTTuM. f. Earliest stage, g. second stage, both nat. size,
g’. part of g, pinnae not jointed to rachis, x 2, A. mature bathyphyll, nat. size, A’. part of a pinna from A,
x 2,7. rhizome and stipe-bases, 2/3, 7. CS of rhizome, k. CS of stipe, both x 4(a HoLttum s.n., 13 Oct.
1929, b PoLak 301, d HoL_trum 24632, f-g ORE h-i WRay 679, e, 7, k from Gard. Bull. S. S. 5,
1932, 278, 279).
1978] _ LOMARIOPSIS GROUP (Holttum) 273
Fig. 7. Teratophyllum clemensiae HOLTTUM. a. Earliest stage, b. second stage, c. mature bathyphylls,
d. pinnae of sterile acrophyll, e. pinnae of fertile acrophyll, all x 2/3 (a CLEMENS 30890, b CLEMENS 50592,
c CLEMENS 40560, d—e CLEMENS 31346).
274 FLORA MALESIANA [ser. II, vol. 14
Fig. 8. Teratophyllum articulatum (J. SM. ex FEE) MetTT. ex KunN. a. Sterile acrophyll and rhizome, x 1/3,
b. one pinna of sterile acrophyll, c. fertile pinnules, d. frond of young plant, second stage, basal pinna
pinnate, all x +/,, e. rhizome with base of stipe, x 1/3, f. CS of rhizome, 2, g. bullate scales from rachis
of acrophyll, x 16 (a BAMLER in Rosenst. Fil. Novog. exsicc. 122, b, e-g BRAss 12202, c CLEMENS 1073,
d EDANO 15165).
1978]
LOMARIOPSIS GROUP (Holttum)
275
mens from the original locality in details of bathy-
phylls and sterile acrophylls. The fertile specimen
(PRICE 927) was found on a high-climbing plant on
the edge of forest, not associated with bathyphylls;
‘no fertile fronds are known from Celebes.
8. Teratophyllum ludens (FEE) Ho_trum, Gard.
Bull. S. S. 5 (1932) 298, pl. 10, f. 35, 46, 47; Rev.
FI. Mal. 2 (1954) 474, f. 278; TAGAWA & IWATSUKI,
Act. Phytotax. Geobot. 24 (1970) 62. — Lomaria
ludens FEE, Hist. Acrost. (1845) 70, t. 30. — Type:
GAUDICHAUD s.n. 1836-37, Singapore (P).
Without name, Bepp. Ferns Br. India (1866)
t. 210. — Stenochlaena sorbifolia [non (L.) J.SM.]
Bepp. Handb. (1883) 423, quoad F.B.I. t. 210
tantum. — Type: PARISH s.n. 1863, Mergui (K).
T. aculeatum var. inermis METT. ex KUHN, Ann.
Mus. Bot. Lugd.-Bat. 4 (1869) 297. — Fig. 6a-e.
Earliest fronds almost as T. aculeatum, with
triangular lamina, deeply lobed or with free pinnae
at base, on one side of midrib only. Transition
bathyphylls on climbing stems 21/,-7'/, cm long;
pinnae 5-9, apical one largest, all on side of rachis
towards ground and standing away from support-
ing tree to which rachis is closely appressed;
pinnae 5-30 mm long, broadly elliptic or ovate,
apices rounded, edges crenulate distally; later
fronds of this type may have 1-2 pinnae on upper
side, towards apex; fronds on stems which creep on
the ground very variable, bearing (on both sides of
rachis) few subequal larger pinnae which are in
some cases widest near base, tapering gradually, no
basal pinnae overlapping rhizome. Rhizome of
adult plant c. 3 mm @, not aculeate. Sterile acro-
phylls to 50 cm long including stipe; pinnae to 10
pairs, lower ones with stalks to 5 mm long, upper
sessile, 7-12 cm long, 2—4!/, cm wide, firm, drying
light olive green, almost elliptic or with edges
parallel in middle part, narrowed about equally to
broadly cuneate base and abruptly short-acuminate
apex; veins near midrib 1—1'/, mm apart; edges
slightly sinuate, narrowly cartilaginous. Fertile
fronds as long as sterile; pinnae to 20 cm long,
3 mm wide, on stalks to 5 mm long.
Distr. Peninsular Thailand and Tenasserim (N.
to Mergui); in Malesia: Malay Peninsula, Borneo
(Sarawak).
Ecol. In fresh-water swamp forest.
9. Teratophyllum clemensiae HoLtruM, Gard. Bull.
S. S. 7 (1934) 262, f. 1-9 (not ibid. 9, 1937, 142,
which refers to 7. /uzonicum). — Type: CLEMENS
31614, N. Borneo, Mt Kinabalu, Penibukan
(SING). — Fig. 7.
Simple fronds on very young plants to 3 by
1'/, cm, base on lower side broadly cordate and
overlapping rhizome, on upper side cuneate, edges
sinuate, apex broadly rounded. Smallest normal
bathyphylls narrowly deltoid, 4'/, by 2 cm, with
3 or 4 pairs of jointed pinnae below deeply lobed
triangular apex, pinnae elliptic, entire, apex
rounded, base unequally cuneate; largest fronds to
9 by 4 cm with 9 pairs of free pinnae 2.2 by 1 cm
somewhat narrowed to rounded apex. Transition
bathyphylls of various types, all with entire pinnae
shaped nearly as acrophylls; pinnae in some cases
all on one side of rachis, usually on both sides,
largest fronds to 18 cm long with 10 pairs of pinnae
on stalks to 3 mm long, lowest pinna always
deflexed, overlapping rhizome. Rhizome of adult
plant 5 mm @ with short spines. Sterile acrophylls
40-70 cm long including stipe 5-15 cm; pinnae
c. 10 pairs, stalks 5-10 mm long, blade 8-15 cm by
11/,-2!/, cm, broadest near unequally cuneate base,
tapering gradually to acuminate apex, texture thin
but firm, drying rather light green, veins c. 2 mm
apart near midrib, margin narrowly cartilaginous
and regular sinuous (prominences at vein-ends).
Fertile fronds with 10-12 pairs of pinnae on stalks
10-15 mm long; each pinna 5-7 cm long, 5 mm
wide, veins uniting in a submarginal vein.
Distr. Malesia: Borneo (Sarawak & Sabah).
Ecol. In forest, at 600-1500 m.
Norte. In 1937 I suggested that transition bathy-
phylls from Palawan (MERRILL 862, US) were
referable to this species. I now think that they are
more likely to belong to 7. /uzonicum, as their
pinnae have a more oblong shape and veins closer
together than in bathyphylls of this size in
T. clemensiae.
2. Section Polyseriatae
Ho.trum, Gard. Bull. S. S. 9 (1938) 356; Blumea 14 (1966) 216-218. — Polybotrya
sensu FEE, Hist. Acrost. (1845) 12, 72, p.p. — Acrostichum sect. Polybotrya HOOK.
Spec. Fil. 5 (1864) 247. — Teratophyllum METT. ex KUHN, Ann. Mus. Bot. Lugd.-
Bat. 4 (1869) 297, p.p. — Arthrobotrya J.SM. Hist. Fil. (1875) 141; UNDERw. Bull.
Torr. Bot. Cl. 33 (1906) 40; Cope. Gen. Fil. (1947) 118; Fern Fl. Philip. (1960)
272. — Polybotrya sect. Teratophyllum CurisT, Farnkr. Erde (1897) 42; DIELs in
E. & P. Nat. Pfl. Fam. 1, 4 (1899) 198. — Polybotrya sect. Arthrobotrya v.A.V.R.
Handb. (1908) 725. — Lomagramma sensu CoPEL. Philip. J. Sc. 3 (1908) Bot. 32,
p.p.; C.Cur. Ind. Fil. Suppl. 1 (1913); v.A.v.R. Handb. Suppl. (1917) 437.
Type species: Polybotrya articulata J.SM. ex FEE.
Distribution. Malesia: Celebes, Moluccas, Philippines, New Guinea; eastwards to Tahiti, southeast-
wards to Queensland and New Caledonia; 3 spp.
Vegetative morphology. See Hottrum, /.c. Acrophylls of the Malesian species T. articulatum are
always bipinnate; in 7. wilkesianum (BRACK.) HoLTTUM (New Caledonia to Tahiti) some acrophylls (both
276 FLORA MALESIANA [ser. Il, volar
sterile and fertile) are simply pinnate, and not infrequently one frond is partly pinnate, partly bipinnate;
in T. brightiae (F.v.M.) HottruM (N. Queensland) all acrophylls are simply pinnate. Detached simply
pinnate acrophylls, whether sterile or fertile, are indistinguishable from such fronds of sect. Teratophyllum.
The youngest stages of bathyphylls are inadequately known; one specimen of a young plant of T. articu-
latum has bipinnate fronds with defiexed basal pinna, not differing in any significant character from sect.
Teratophyllum. The bathyphylls of T. brightiae are always simply pinnate (HOLTTUM, 1938, pl. 28) but the
earliest stage is not known. Vascular anatomy of adult rhizomes is very like that in Lomariopsis, but the
bases of stipes are somewhat swollen and rather imperfectly jointed, not gradually decurrent as in
Lomariopsis. Fertile leaflets have an additional vascular supply for the sporangia, as in sect. Teratophyllum.
Taxonomy. As indicated in the synonymy, T. articulatum was first included in the genus Polybotrya
Humps. & BONPL. and transferred to the new genus Teratophyllum by KUHN in 1869. JOHN SMiTH
established a new genus Arthrobotrya for it in 1875, but CHrist and DIiELs retained it in Polybotrya sect.
Teratophyllum, in which they did not include the species here placed in Teratophyllum sect. Teratophyllum.
The second species of the section, T. wilkesianum, was described in Polybotrya by BRACKENRIDGE in 1854,
the third, T. brightiae, by F. voN MUELLER in Acrostichum in 1870. UNDERWOOD places T. brightiae (as
Stenochlaena hiigelii) in Stenochlaena sect. Lomariopsis, remarking that T. articulatum belonged to a
distinct genus Arthrobotrya. In his early work on Philippine ferns, COPELAND transferred T. articulatum
to the genus Lomagramma because of its similarity to L. polyphylla (with which it had been associated by
Diets, who placed the simple pinnate Lomagramma species in Gymnopteris). In Genera Filicum (1947)
COPELAND revived the genus Arthrobotrya; but he avoided consideration of the species T. brightiae, which
has always simple acrophylls. The only clearly definable character distinguishing the two sections of
Teratophyllum, as here presented, is the polyseriate or biseriate arrangement of fronds on the upper
surface of the rhizome, with accompanying difference of vascular structure; this does not appear to me to
warrant generic separation of the two.
small widely-spaced lobes of a narrow apical
10. Teratophyllum articulatum (J.SM. ex FEE) METT.
ex Kuun, Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 297;
Ho.trum, Gard, Bull. S. S. 9 (1938) 356. — Poly-
botryaarticulata J.SM. ex FEE, Hist. Acrost. (1845)
74, t. 37; Curist, Farnkr. Erde (1897) 42, f. 101;
Diets in E. & P. Nat. Pfl. Fam. 1, 4 (1899) 198;
CopEL. Polypod. Philip. (1905) 40; Hottrum,
Blumea 14 (1966) 217. — Acrostichum (sect. Poly-
botrya) articulatum Hook. Spec. Fil. 5 (1864) 247.
— Arthrobotrya articulata J.SM. Hist. Fil. (1875)
141; CopeL. Gen. Fil. (1947) 118; Philip. J.
Sc. 78 (1949) 401; Fern FI. Philip. (1960) 272. —
Lomagramma articulata CopeL. Philip. J. Sc. 3
(1908) Bot. 32; v.A.v.R. Handb. Suppl. (1917) 437.
— Type: CuMING 296, Luzon (BM, GH, K, P).
Lomagramma bipinnata Cope. Philip. J. Sc. 11
(1916) Bot. 41; v.A.v.R. Handb. Suppl. (1917) 438.
Fi ila Ramos BS 17515, Samar (MICH). —
ig. 8.
Youngest stage with simply pinnate fronds to
9 cm long with deeply crenate articulate pinnae 8
by 4—5 mm, apex of frond a narrow lobed lamina
not articulate; next stage with basal pinnae pinnate
and in some cases deflexed across rhizome, upper
simple pinnae to 22 by 11 mm, pinnules of basal
pinnae to 6 by 4 mm; later fully bipinnate fronds
2-ranked on slender rhizome are like acrophylls but
smaller. Adult rhizome 7-8 mm or more wide,
smooth. Sterile acrophylls with stipes 10-20 cm,
lamina to 60 cm long, bipinnate; pinnae 15—24 cm
long, articulate to rachis, pinnate with to 20 or more
pairs of articulate pinnules which grade into the
lamina; pinna-rachis winged throughout; pinnules
sessile or nearly so, base very asymmetric, narrowly
cuneate basiscopically, very broadly cuneate and
usually with a well-developed auricle acroscopi-
cally, edges crenate, apex rounded, largest pinnules
15-25 mm long, 6-8 mm wide above auricle;
brown bullate scales on lower surface of costa.
Fertile fronds somewhat smaller than sterile; pin-
nules distinctly stalked, 7-15 mm long, c. 2 mm
wide when dry, distinctly auricled.
Distr. Malesia: Celebes (SW. & NE.), Moluccas
(Ceram, Halmahera), Philippines (Luzon, Leyte,
Samar, Mindanao), New Guinea; Solomon Islands.
Notes. This species is distinguished from T. wil-
kesianum (New Caledonia, Fiji, Samoa, Tahiti) by
the auricled acroscopic base of its pinnules, and by
the complete absence of simply pinnate acrophylls;
also apparently by the brown bullate scales of the
lower surface of pinnules.
Two SARASIN specimens from N. Celebes match
T. wilkesianum from New Caledonia very closely
in shape of leaflets, but they have the scales charac-
teristic of T. articulatum. It seems possible that
these specimens represent a distinct local form
of the species. COPELAND (1960: 273) refers to a
specimen of his own from Mindanao which
resembles 7. wilkesianum; this has rather small
fronds with deeply incised small leaflets which are
very asymmetric at the base and seem to me much
less like T. wilkesianum than the Celebes specimens.
4. LOMAGRAMMA
J.SM. in Hook. J. Bot. 3 (1841) 402, nom. nud.; in Hook. Gen. Fil. (1842) t. 98;
Bepp. Handb. Ferns Br. India Suppl. (1892) 105; C.Cur. Ind. Fil. Suppl. 1 (1913)
49 (p.p.), 118; v.A.v.R. Handb. Suppl. (1917) 436, excl. L. articulata et bipinnata;
CopeL. Univ. Cal. Publ. Bot. 16 (1929) 76, excl. L. articulata; HOLTTUM, Gard.
Bull. S. S. 9 (1937) 190-221; Rev. Fl. Mal. 2 (1954) 477; Cope. Fern FI. Philip.
1978] LOMARIOPSIS GROUP (Holttum) 277
(1960) 273-275; HoLtrum, Blumea 14 (1966) 221-223. — Leptochilus KAULF. p.p.
BL. En. Pl. Jav. (1828) 206; C.Cur. Bot. Tidsskr. 26 (1904) 283; Ind. Fil. (1905)
xxvi; V.A.Vv.R. Handb. (1908) 746. — Cheilolepton Fée, Hist. Acrost. (1845) 19. —
Neurocallis FEE p.p. PRESL, Epim. Bot. (1851) 177. — Chorizopteris Moore, Gard.
Chron. Agr. Gaz. (1855) 854. — Neurocallis sect. Cheilolepton Moore, Ind. Fil.
(1857) xix. — Acrostichum sect. Chrysodium (FEE) Hook. Spec. Fil. 5 (1864) 268,
p.p.; BAKER, Syn. Fil. (1868) 423. — Polybotrya HumsB. & BONPL. sect. Loma-
gramma KuHN, Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 295. — Gymnopteris BERNH.
p.p. Diets in E. & P. Nat. Pfl. Fam. 1, 4 (1899) 198. — Fig. 9-14.
Rhizome of adult plant climbing tree-trunks, broad, bearing 3-5 rows of fronds
on dorsal surface (except L. brassii which has 2 rows), 3 or more fronds often
attached close together, such groups rather widely separated; young parts of
rhizome and young fronds densely scaly; scales peltate, smaller ones always
clathrate, larger ones often with a central or basal area of uniformly thin-walled
cells with darker contents; vascular system as in Lomariopsis. Stipes gradually
decurrent at their bases to ridges on rhizome, the pale linear aerophore on each
side of a stipe decurrent also. Young plants with slender rhizome bearing only 2
longitudinal rows of fronds, creeping on rocks or on the ground; fronds widely
separated, of distinctive form (bathyphylls). Fronds simply pinnate (bipinnate in
L. polyphylla BRACK., New Hebrides to Fiji, also recently discovered in L. cordi-
pinna HOLTTvUM in Fiji) with all pinnae, apical one included, jointed to rachis; scales
on fronds clathrate, smaller ones bullate at base; veins forming a uniform network
of 3 or more rows of oblique areoles without included veinlets and without main
veins; edges of sterile pinnae entire or crenate. Fertile fronds with pinnae narrower
than sterile, covered beneath with sporangia, venation as sterile but areoles fewer;
slender-stalked paraphyses present, about as long as sporangia, their apices dilated,
irregular in shape, formed of 8-10 cells with thick lateral walls as small scales on
other parts of plant. Spores lacking perispore.
Type species: L. pteroides J.SM.
Distribution. Assam to S. China and Thailand; throughout Malesia except the eastern Lesser Sunda
Is.; Solomon and New Hebrides to Tahiti; about 18 spp. (one species in tropical America has been
included by CHING, but its status is doubtful,; see p. 278, 289).
Ecology. In all cases where young plants have been observed, they grow on wet rocky banks of small
streams in high forest, at altitudes from sea level to 1500 m. Their slender rhizomes, bearing erect fronds,
are wide-creeping, extending into the forest away from the stream until they meet tree-trunks, up which
they climb vertically, attached by roots, to 10 m or more (fig. 9), developing a much thicker rhizome and
larger horizontal or drooping fronds. Fertile fronds are produced on the upper parts of climbing rhizomes,
probably in response to drier conditions; the only occasion on which I have seen fertile fronds on L. suma-
trana in Malaya was on a plant which had recently been exposed by felling of neighbouring trees. L. sinuata
in New Guinea tolerates more exposed conditions than L. sumatrana and can continue to grow after par-
tial clearing of forest. Spores are probably short-lived; this may limit their dispersal range.
Vegetative morphology. In morphology and anatomy of rhizome Lomagramma is closely similar
to Lomariopsis, but scales are wholly or partly clathrate. Young plants of all species have simply pinnate
fronds with jointed pinnae and a lobed apical lamina continuous with the rachis. Sooner or later fronds are
produced in which this lobed apex is aborted (a rudiment can sometimes be seen) and its place taken by a
pinna which is jointed at its base. In some species this change takes place only when the plant is large
enough to produce fronds with many pinnae (see special key below); in others the apical lamina is lacking
from a much earlier stage when fronds have few pairs of pinnae. I have observed this distinction con-
stantly between the two species in Malaya which I have seen many times in their native habitat. Unfor-
tunately young stages have not been well collected for most species, or collections have consisted only of
young stages without association of the adult stage. The venation of sterile acrophyll pinnae is very
constant in all cases; that of fertile pinnae has small areoles, in number varying with width of pinnae.
There is not a great range in shape and size of pinnae, whether sterile or fertile, and I have not found it
possible to construct a complete key based only on sterile fronds (which are commonest in herbaria).
278 FLORA MALESIANA [ser. II, vol. 14
Gametophyte. See p. 258 supra.
Cytology. The only observations are by Roy and MANTON on root-tips of plants of L. sinuata and
L. melanolepis sent by me from New Guinea and cultivated at Kew; both cases showed 2n = 82.
Taxonomy. JOHN SMITH based the genus on the peculiar condition of the fertile pinnae of L. pteroides,
which have the sporangia confined to a marginal band with a narrow sterile area between this and the
midrib, at least near the base of a pinna. In 1845 FEe based the genus Cheilolepton on L. lomarioides (of
which he published a good figure) but did not refer to L. pteroides, presumably because he thought it not
to be acrostichoid, though J. SmirH had noted its possible identity with L. Jomarioides (which he had not
seen). Moore also, in his scheme of classification (1857) placed Lomagramma (limited to L. pteroides) ina
group of genera quite distinct from Cheilolepton, which latter he united with Neurocallis (a near ally of
Acrostichum s.str.) because of similarity of venation. HOOKER (1864) stated that L. pteroides was only an
abnormal form of L. /omarioides, and regarded specimens from Samoa (L. cordipinna HOLTTUM) as
belonging to the same species. He placed L. /omarioides near Acrostichum s.str. and Neurocallis on account
of their common characters of acrostichoid fertile pinnae and reticulate venation. KUHN (1869) placed
Lomagramma as a section of Polybotrya. In his last work BEDDOME (1892) recognized Lomagramma once
more as a distinct genus, and described a new species (L. perakensis); but DIELs (1899) relegated it again
to a confused mixture of acrostichoid ferns of very diverse affinity in Gymnopteris. CHRISTENSEN (1904)
placed Lomagramma as a section of Leptochilus (another section being Bolbitis) but later recognized it as a
distinct genus.
As noted in the introductory statement on this group of genera, I regard Lomagramma as closely related
to Bolbitis. Lomagramma differs from Bolbitis constantly in Malesia in the following characters: high-
climbing rhizome, articulate pinnae, reticulate venation without main veins or included free veinlets and
thin-walled spores lacking perispore. In tropical America there is a species of Bolbitis which is very near
Lomagramma in venation, B. serratifolia (KAULF.) CHING. The species originally named Polypodium
guianense AUBL. has been placed in Lomagramma by CHING and in Bolbitis by KRAMER. It has
Lomagramma-like bathyphylls, a high-climbing rhizome and articulated pinnae. In my view it is not closely
related to Malesian Lomagramma (see p. 289). There is considerable variation in the pattern of venation
in Bolbitis, and in Malesia also are species which have a reticulate venation lacking free included veinlets,
though none lack main lateral veins. Bathyphyll pinnae of Lomagramma have a toothed margin much as
some Bolbitis spp. If one has a small detached bathyphyll from a young plant of Lomagramma, the only
character by which one can be sure it is not Bolbitis is the jointed pinnae.
COPELAND wrote (Fern FI. Philip. 1960, 275) ‘‘at present it does not seem to me quite impossible that
our specimens (of Lomagramma) represent a single widely variable species’’. I am sure that distinct species
exist, but admit that in New Guinea I have found their delimitation difficult. More field work is necessary
before a better arrangement can be established; information on all stages of development, gained sepa-
rately in individual localities, is needed. It is possible that hybrids exist, as in Bolbitis.
Keys. As noted above, the most characteristic parts of a Lomagramma species are often the bathyphylls
and the fertile fronds. I find it impossible to construct a satisfactory key which does not include these
characters, and they are used in the main key which follows. Incomplete keys, based only on one of the
three different kinds of fronds, are also given, as a help to the identification of incomplete specimens.
KEY TO THE SPECIES
1. Mature plants with broad rhizomes bearing 3 or more rows of fronds; sterile acrophyll pinnae not
toothed throughout.
2. Sterile acrophyll pinnae very firm, veins indistinct, not prominent on upper surface.
3. Sterile acrophyll and bathyphyll pinnae entire or at most sinuate towards apices; fertile pinnae to
20 cm long.
4. Sterile acrophyll pinnae to 12'/, by 1’/, cm, base broadly cuneate to truncate and sometimes slightly
auricled on acroscopic side; fertile pinnae to 2mm wide. . . 1. L. lomarioides
4. Sterile acrophyll pinnae to 25 by 21/, cm, base narrowly cuneate; fertile pinnae 3-5 mm wide
2. L. perakensis
3. Sterile acrophyll pinnae distinctly toothed towards apex; fertile ras to 10 cm long.
5. Fertile pinnae 3 mm wide, stalks to 5 mm or SiN ee : POLS . 3. L. merrillii
5. Fertile pinnae 8 mm wide, sessile . . . ae. ae novoguineensis
2. Sterile acrophyll pinnae with veins distinct and + . prominent on upper ‘surface.
6. Fertile pinnae 4!/, by 0.8 cm with rounded apices; sterile pinnae 1.3 cm wide, toothed towards
Apices =) 2: . 4, L. novoguineensis
6. Fertile pinnae narrower or much longer: sterile wider or with entire edges.
7. Sterile acrophyll pinnae to 8 mm wide, entire. . . Se angustipinna
7. Sterile acrophyll pinnae more than 1 cm wide, narrower ones often toothed towards apices.
8. Bathyphylls with narrow lobed apical lamina coutinuous with rachis on fronds with 20 pairs of
pinnae.
9. Sterile acrophyll pinnae commonly 1.2-1.8 cm wide, broad and almost symmetrical at base;
fertile pinnae dilated and partly sterile at base eae Cea pteroides
9. Sterile acrophyll pinnae commonly 2 cm or more wide, asymmetric at ‘base; fertile pinnae not
dilated at base . . . . 7, L. sumatrana
8. Bathyphylls with jointed apical pinna on fronds with few pairs of pinnae.
1978] ; LOMARIOPSIS GROUP (Holttum) 279
10. Base of lower sterile acrophyll pinnae broadly rounded or subcordate on both sides.
11. Pinnae firm and opaque, veins hardly prominent; scales on rhizome distinctly clathrate
8. L. brooksii
11. Pinnae thin, veins prominent; scales on rhizome not clathrate except near their tips
9. L. leucolepsis
10. Base of lower sterile acrophyll pinnae distinctly asymmetric, narrower on basiscopic side,
acroscopic side usually broadly cuneate.
12. Fertile pinnae 5-8 mm wide; sterile pinnae 2-4 cm wide. . . . . . 10. L. sinuata
12. Fertile pinnae not over 4 mm wide, sterile mostly not over 2 cm wide.
13. Suprabasal sterile acrophyll pinnae rounded on basiscopic side at base 11. L. copelandii
13. Suprabasal acrophyll pinnae narrowly cuneate on basiscopic side at base
12. L. melanolepis
1. Mature plants with slender rhizome bearing 2 rows of fronds; sterile pinnae with edges strongly toothed
eetabtipniet. eS NN oe PES ee rs
KEY BASED ON STERILE ACROPHYLLS
1. Pinnae very firm and opaque, veins not or slightly ag amc
2. Pinnae to 26 cm long, base cuneate both sides .... . ote ner i cee oe bee, AC DETAKENSIS
2. Pinnae not over 15 cm long.
3. Pinnae entire.
4. Lower pinnae almost equally subcordate at base 8. L. brooksii
4. Lower pinnae unequal at base, acroscopically broadly truncate or slightly auricled
. L. lomarioides
Seebitmaec distinctly toothed towards:apices . ~. 2. 5. 2.5... :.: s:- . . 3. L. merrillii
4, L. novoguineensis
1. Pinnae thinner, usually translucent when dry, veins always distinctly prominent.
5. Largest pinnae not over 1'/, cm wide.
6. Largest pinnae 10 by 0.8 cm, entire . . A 5 ea Gy) ee, oe Ss IG angustipmna
6. Largest pinnae usually wider, strongly toothed . A Be i hire ie cate wl) A) LA el stT
5. Largest pinnae wider.
7. Lower pinnae with basiscopic base broadly rounded to subcordate. . ... . . 6. L. pteroides
8. L. brooksii
9. L. leucolepis
7. Lower pinnae with basiscopic base cuneate or narrowly rounded.
8. Basiscopic base narrowly cuneate.
Seinnae 2—4.cm) wide, to 20;cmilonge” 2. 2s a es ee ae eo ee . 10. L. sinuata
Srebinnae rarely.over.2 cnt wide shorter: =... @ 2 8s) = sss elec. eee) on Dm! B melanolepis
SEE ASISCODICIDASE LOUNGE gms ene) ea al) coy i Se ee 7. L. sumatrana
“10. L. sinuata f. papuana
11. L. copelandii
KEY BASED ON BATHYPHYLLS ONLY
1. Apical lamina lobed and continuous with rachis on fronds with 15-20 pairs of pinnae.
2. Pinnae entire or with slightly sinuateedges ....... SS Sy tee ee. & cual ale loniariouies
2. Pinnae distinctly toothed, at least towards apices.
3. Pinnae blunt-tipped, bluntly toothed, drying reddish . . . . . 6, L. pteroides
3. Pinnae acute (except youngest stages), acutely toothed, not drying reddish.
4. Pinnae falcate, toothed near apices only ......... Ree Oe aye AO DE OOKSIE
4. Pinnae toothed throughout.
5. Pinnae to 5 by 1.2 cm ee Mae eer ris hae ee ee We a . 7. L. sumatrana
5. Pinnae shorter and narrower ........ 4, L. novoguineensis
1. Apical lamina replaced by a pinna jointed to rachis on fronds with few pairs of pinnae.
6. Pinnae entire or with slightly sinuateedges ........... . . . 2. L. perakensis
Gr Paniae distinctly:toothed . is. 2 a a 2 ee ie
10. L. sinuata
11. L. copelandii
12. L. melanolepis
13. L. brassii
Bathyphyilicharacters notsknowns (4-2) cee ae ee eae 5. L. angustipinna
9. L. leucolepis
1. Lomagramma lomarioides (BL.) J.SM. Hist. Fil. En. Pl. Jav. (1828) 206; v.A.v.R. Handb. (1908)
(1875) 143; Bepp. Handb. Suppl. (1892) 106, p.p.; 746, p.p. — Cheilolepton blumeanum FE, Hist.
v.A.v.R. Handb. Suppl. (1917) 438, p.p.; HoLtrum, Acrost. (1845) 89, t. 51. — Acrostichum blumeanum
Gard. Bull. S. S. 9 (1937) 204, pl. 12,13; BACKER & (FEE) Hook. Spec. Fil. 5 (1864) 268, p.p.; BAKER,
Postu. Varenfl. Java (1939) 153; CopeL. Philip. J. Syn. Fil. (1868) 423, p.p.; RaciB. Fl. Btzg 1 (1898)
Sc. 78 (1949) 401. — Leptochilus lomarioides BL. 55. — Polybotrya lomarioides (BL.) KUHN, Ann.
- ’ , Pt by, pS: tit
ERS ao Roget BT nF
[ser. II, vol. 14
"
4 fal 4
wep at ee
(Photogr. R. E. HoLTTum).
<
Zz
=
n
a]
4
s
<
ia
oe)
|
,
Fig. 9. Lomagramma perakensis Bepp. Acrophylls, on tree in forest at Cameron Highlands, Malaya
1978] LOMARIOPSIS
Mus. Bot. Lugd.-Bat. 4 (1869) 295. — Type:
BLuME s.n., Salak, Java (L).
L. abscondita v.A.vV.R. Bull. Jard. Bot. Btzg I,
11 (1913) 16; Handb. Suppl. (1917) 439; Bull. Jard.
Bot. Btzg III, 2 (1920) 159. — Type: RActBoRSKI
g.n., G. Salak (BO).
Rhizome scales thin, to 2 mm wide, strongly
clathrate with brown cell-walls. Bathyphylls with
pinnae to 25 pairs and apical lamina continuous
with rachis; largest pinnae 5 by 1 cm, acroscopic
base broadly cuneate to subcordate, basiscopic
narrowly rounded, edges quite entire or slightly
sinuate, apex acuminate, falcate, texture firm, veins
hardly visible on upper surface, slightly prominent
beneath. Sterile acrophylls: stipe to 20 cm, frond to
100 cm long, middle pinnae largest, lowest gradu-
ally reduced and more widely spaced; largest
pinnae 12'/, by 1'/, cm, acroscopic base broadly
cuneate to truncate and sometimes slightly auricled,
basiscopic narrowly rounded; texture of old fronds
very firm, veins hardly visible on upper surface,
somewhat prominent below but rather broad;
edges entire, often inrolled when dried, apex acumi-
nate, falcate. Fertile fronds: pinnae sessile, to at
least 18 cm long, 1-2 mm wide.
Distr. Malesia: Java, Lesser Sunda Is. (Bali).
Ecol. In forest at 750-1500 m.
Note. Bathyphylls have not often been collected;
a good example is PALMER & BRYANT 1190,
Tjibodas (US).
2. Lomagramma perakensis BEDD. Handb. Suppl.
(1892) 107; v.A.v.R. Handb. Suppl. (1917) 439;
Ho.trum, Gard. Bull. S. S. 9 (1937) 210, pl. 16;
Rev. Fl. Mal. 2 (1954) 478, f. 281. — Leptochilus
perakensis (BEDD.) C.CuHrR. Ind. Fil. (1906) 387;
v.A.v.R. Handb. (1908) 747. — Type: J. Day s.n.,
Perak (K). — Fig. 9, 11f-g.
Rhizome scales to 10 by 11/,; mm, dull brown,
distinctly clathrate. Bathyphylls as small as 5 cm
long with apical pinna jointed to rachis; stipes of
largest bathyphylls 25 cm long, lamina c. 30 by
14cm, pinnae 10-15 pairs oblique to rachis, sessile,
edges entire or somewhat sinuous, not toothed,
largest 7 by 1.2 cm, texture firm, veins not promi-
nent. Sterile acrophylls: stipe to 40 cm, fronds to
125 cm long; pinnae oblique, largest 26 by 2!/, cm,
subsessile, base rather narrowly unequally cuneate,
edges slightly undulate, not toothed, apex falcate
acuminate, texture very firm, veins hardly promi-
nent on either surface. Fertile fronds: pinnae com-
monly to 20 cm long (largest seen 40 cm), 3-5 mm
wide, on stalks 3-7 mm long.
Distr. Malesia: Sumatra, Malay Peninsula
(north to Pattani in S. peninsular Thailand).
ae In valleys in mountain forest, at 600—
m.
3. Lomagramma merrillii HoLtTruM, Gard. Bull.
S. S. 9 (1937) 208, pl. 14; Cope. Fern FI. Philip.
(1960) 274. — Type: MERRILL 8282, Mindanao,
Zamboanga District (M, destroyed).
Bathyphylls: pinnae to c. 10 pairs, to 3 by 1.3 cm,
acroscopic bases broadly, basiscopic narrowly,
crenate, edges acutely toothed, apex acute; apical
pinna largest and articulate, c. 6 by 1.4 cm. Sterile
acrophylls: stipe to 15 cm, frond to 45 cm long;
upper pinnae largest, c. 9 by 1.7 cm, falcate, base
unequally cuneate, edges finely and irregularly
GROUP (Holttum)
281
toothed towards apex, subcoriaceous, veins not
conspicuous. Fertile pinnae (seen only in immature
condition) to 9 cm long, 3 mm wide, apex acute (?),
stalks to 5 mm long.
Distr. Malesia:
Negros).
Ecol. In forest, at 1500 m.
Note. In 1937 I saw the type specimen from the
Manila herbarium; this and the CLEMENS specimen
also cited with the original description are now lost,
and I do not know of any duplicates. The only post-
war collections consist of bathyphylls.
Philippines (Mindanao,
4. Lomagramma novoguineensis (BRAUSE) C.CHR.
Ind. Fil. Suppl. 3 (1934) 124; Hottrum, Gard.
Bull. S. S. 9 (1937) 208, pl. 15; Copev. Philip. J. Sc.
78 (1949) 401. — Leptochilus novoguineensis
BrAusE, Bot. Jahrb. 56 (1920) 117. — Type:
LEDERMANN 9524, NE. New Guinea, Sepik River,
Etappenberg, 850 m (B). — Fig. 10a.
Bathyphylls with apex continuous with rachis on
fronds with many pinnae. Sterile acrophylls: pinnae
to 11 by 1.3 cm, base subequally rounded to trun-
cate (lowest pinnae with base narrower on basi-
scopic side), edges serrate towards apex, texture
very firm, veins at most slightly prominent; bullate
acuminate scales abundant on lower surface of
costae and veins; apex of frond sometimes a
narrow lobed lamina continuous with rachis. Fer-
tile pinnae sessile, to 41/, by 0.8 cm, apex rounded.
Distr. Malesia: New Guinea (W.-E.).
Note. Only two collections of acrophylls and
fertile fronds are known, the second being from
50 m alt. in western New Guinea. A third collec-
tion, of bathyphylls only, may also represent this
species (WOMERSLEY & MILLAR NGF 8571,
Western Highlands of NE. New Guinea, 700 m).
The fronds of this have up to 40 pairs of closely-
placed pinnae below the narrow lobed apex; the
larger pinnae resemble the upper pinnae of the type
of L. novoguineensis. A poor specimen collected by
CLEMENS (8083A, Sattelberg, 1000 m, B) may also
represent this species, but fertile fronds are young,
not fully expanded, so that the mature size of the
pinnae cannot be judged.
5. Lomagramma angustipinna CopeL. Univ. Cal.
Publ. Bot. 18 (1942) 222; Philip. J. Sc. 78 (1950)
402, pl. 6A. — Type: Brass 13446, Idenburg River,
W. New Guinea, 750 m (GH; dupl. in MICH).
Rhizome 7 mm wide, containing 4 meristeles
besides the root-bearing ventral one; rhizome-
scales 1-2 mm long, cell-walls very dark. Sterile
acrophylls: stipe 14 cm; lamina 50 cm long with
c. 30 pairs of pinnae, lower ones decreasing; middle
pinnae 10'/, by 0.8 cm, falcate-acuminate, sub-
sessile, base very narrowly cuneate on basiscopic
side, broader on acroscopic, edges entire, texture
thin, veins fine, distinct on both surfaces. Fertile
pinnae to 9 cm by 3 mm, on stalks to 2 mm long.
Distr. Malesia: W. New Guinea. Known only
from type collection.
6. Lomagramma pteroides J.SM. in Hook. J. Bot. 3
(1841) 402; ibid. 4 (1841) 152; in Hook. Gen. Fil.
(1842) t. 98; CopeL. Philip. J. Sc. 1 (1906) Suppl.
166; v.A.v.R. Handb. Suppl. (1917) 438; HoLtTrum,
Gard. Bull. S. S. 9 (1937) 213, pl. 13; CopeL. Fern
Fl. Philip. (1960) 273. — Acrostichum blumeanum
282 FLORA MALESIANA [ser. II, vol. 14
Natok:
@1970 Q 2
Fig. 10. Lomagramma novoguineensis (BRAUSE) C. Cur. a. Fertile pinnae, x 2/;. — L. sinuata C. Cur.
Sf. papuana C. Cur. b. Sterile acrophyll, c. apex of bathyphyll, d. rhizome of young plant, all x 2/,;, —
L. sinuata C. Cur. Typical form, e. sterile acrophyll, f. apex of bathyphyll, both x 2/,. — L. melanolepis
V.A.v.R. g. Apex of bathyphyll, x 2/3, A. scale from rhizome, < 20 (a DoctEers VAN LEEUWEN 9616, b-c
NGF 17699, d, A ccult. Kew, origin Lae, e KooRDERS e204: J BAKHUIZEN VAN DEN BRINK Jr 3664, g NGF
15892).
1978]
(FEE) Hook. Spec. Fil. 5 (1864) 268, p.p.; BAKER,
Syn. Fil. (1869) 423, p.p. — Polybotrya pteroides
(J.SM.) KUHN, Ann. Mus. Bot. Lugd.-Bat. 4 (1869)
295. — Leptochilus lomarioides BL. var. pteroides
v.A.v.R. Handb. (1908) 746. — Type: CUMING 223,
Luzon (BM; dupl. in K).
L. pteroides var. subcoriacea CopPeL. Philip. J.
Sc. 3 (1908) Bot. 32; v.A.v.R. Handb. Suppl.
(1917) 439. — L. subcoriacea Cope. Philip. J. Sc.
40 (1929) 308; Fern FI. Philip. (1960) 274. — Type:
COPELAND 1736, Mindanao (MICH; dupl. in B,
BM, P).
L. pedicellata CoPEL. Philip. J. Sc. 81 (1952) 23;
Fern FI. Philip. (1960) 275. — Type: EDANo PNH
531, Palawan (MICH).
L. cordata Cope. Philip. J. Sc. 84 (1955) 162,
t. 2. — Type: SuLir PNH 20277, Biliran (MICH).
— Fig. 1la-d.
LOMARIOPSIS GROUP (Holttum)
283
Bathyphylls with up to 28 pairs pinnae have
lobed terminal lamina continuous with rachis;
lower pinnae gradually reduced, lowest c. 7 by
3 mm, largest pinna 2!/, by 0.7 cm, base unequally
cuneate (more narrowly on basiscopic side), edges
crenulate with teeth 2-3 mm apart, apex often not
acute, bases and apices of lower pinnae more
rounded; stipe, rachis and costae rather densely
scaly, scales dark; pinnae reddish when dry; veins
strongly raised on both surfaces. Sterile acrophylls:
stipes 20-30 cm, pinnae commonly 1.2—1.8 cm wide,
12-15 cm long (largest seen 22 by 2.2 cm), base
slightly unequally subtruncate to subcordate,
edges subentire and parallel, apex acuminate and
somewhat falcate, irregularly sinuate or somewhat
toothed, uppermost pinnae gradually smaller with
less truncate base and more toothed apex, lowest
pinnae not much reduced but often with cordate
Fig. 11. Lomagramma pteroides J. SM. a. Middle pinnae of bathyphyll, b. apex of bathyphyll, c. sterile
acrophyll, d. fertile pinna, all x 2/,. — L. sumatranav.A.v.R. e. Apex of bathyphyll, x 2/;.— L. perakensis
Bebb. f. Apex of bathyphyll, g. base of pinna of acrophyll, both x 2/3; (a-b EDANo 476, c-d ELMER 9068,
e MATTHEW s.n., Jan. 1907, fg HOLTTUM 24994).
284 FLORA MALESIANA [ser. II, vol. 14
Fig. 12. Lomagramma sinuata C. Cur. Transition from bathyphylls (centre) to acrophylls (base, left &
right), Cult. R.B.G. Kew, origin NE. New Guinea (Photogr. R. VAN CREVEL).
1978]
base; texture thin, colour often reddish when dry;
veins slightly prominent, areoles rather small, dark
scales rather abundant. Fertile fronds: pinnae
commonly to 20 cm long, base unequally cuneate,
widening suddenly from a winged stalk to 10 mm
long; base of pinna 7-10 mm wide, only the edges
fertile, upper part gradually narrowed, distal half
sometimes covered with sporangia but often with a
narrow sterile band each side of midrib.
Distr. Malesia: Philippines (Luzon, Palawan,
Negros, Panay, Mindoro, Mindanao).
Ecol. Apparently in lowland forest and to
c. 1000 m (records few).
Notes. I have not seen the type of L. cordata
CopeL. The photographic illustration shows a
complete frond sterile at the base, fertile in distal
parts. The sterile pinnae are only 7 mm wide, but
on such a frond would probably not be normal.
The fertile pinnae are rather long-stalked even to
the apex of the frond (stalks are evidently 4-6 mm
long, not 4-6 cm as in the printed description)
where stalks are usually shorter.
M. G. Price has collected on Mt Makiling
bathyphylls of this species in which the lower pin-
nae are gradually decrescent, but the lowest are
elongate and deeply lobed so as to be almost pin-
nate.
7. Lomagramma sumatrana v.A.v.R. Bull. Jard.
Bot. Btzg III, 2 (1920) 158; HoLtrum, Gard. Bull.
S. S. 9 (1937) 217, pl. 16; Rev. Fl. Mal. 2 (1954)
477, f. 280. — Type: LOrzING 5564, Sumatra,
Sibolangit (BO; dupl. in P, SING). — Fig. 11e.
Bathyphylls: stipes 3-10 cm long; lamina to 40
by 9 cm; pinnae 20-30 pairs, lowest rather reduced
and deflexed, upper gradually smaller, uppermost
grading into lobes of frond-apex which is not
pinna-like not jointed; middle pinnae to 5 by
1.2 cm, at right angles to rachis, acroscopic base
broadly subtruncate to subauriculate, basiscopic
narrower, rounded, sides almost parallel for 2/; of
length, then tapering to acute apex; edges shallowly
crenate with 1-2 teeth at distal end of each crena-
tion; texture thin, veins slender, prominent.
Sterile acrophylls: stipes to 20 cm, frond to at least
90 cm long; lowest pinnae somewhat reduced and
more distant, uppermost gradually smaller; middle
pinnae to 18 by 21/, cm, sessile or subsessile,
acroscopic base broadly cuneate, basiscopic
narrower and rounded to cuneate; edges usually
with a few irregular teeth towards acuminate apex;
texture thin, veins conspicuous. Fertile pinnae to
15 cm by 5 mm, on stalks to 2 mm, upper ones
gradually reduced, uppermost c. 4 cm long.
Distr. Malesia: N. Sumatra, Malay Peninsula.
Ecol. Near rocky streams in forest, 0-500 m.
Note. Acrophyll pinnae of this species are very
similar in shape to those of L. sinuata, though never
so large; bathyphylls show a constant difference in
all cases observed.
8. Lomagramma brooksii Copev. Philip. J. Sc. 3
(1908) Bot. 345; ibid. 7 (1912) Bot. 60; v.A.v.R.
Handb. Suppl. (1917) 439; HoLtrum, Gard. Bull.
S. S. 9 (1937) 199, pl. 8 (by error as L. borneensis).
— Type: Hewitt & Brooks, Bongo Mt, Sarawak
(MICH).
Bathyphylls on young plants, with apical lamina
continuous with rachis, to 20 cm long with 12-15
LOMARIOPSIS GROUP (Holttum)
285
pairs pinnae; larger bathyphylls with apical pinna
articulate; pinnae of youngest plants with rounded
apex, of larger ones acute; acroscopic base broadly
subtruncate and sometimes slightly auricled,
basiscopic much narrower, rounded; edges toothed
only towards apices; texture thin, veins distinct and
slightly raised on both surfaces. Sterile acrophylls:
stipe short (often only 4 cm); fronds to 75 cm long;
middle pinnae at right angles to rachis, sessile, to
15 by 2 cm, basiscopic base rounded to subcordate,
a little narrower than subtruncate acroscopic base;
edges entire, sometimes sinuate towards apices;
texture thin but very firm, veins slightly raised;
lower pinnae slightly stalked, their bases sym-
metrical and subcordate. Fertile pinnae to 15 cm by
4 mm, more commonly smaller, on winged stalks
to 2 mm.
Distr. Malesia: N. Borneo (Sarawak), North &
Central Celebes, Philippines (Luzon, Palawan).
Ecol. Near streams in forest, at 300-1500 m.
Note. The North Celebes specimen (PostHUMUS
2335) has bathyphylls with 20 pairs of pinnae and
still a narrow lobed apical lamina continuous with
the rachis. Central Celebes specimens (SARASIN
953) have fronds not completely expanded.
9. Lomagramma leucolepis HoLTruM, Blumea 14
(1966) 224. — Type: Brass 12950, W. New Guinea,
Idenburg River (L; dupl. in BM, BO, GH, MICH).
Rhizome 12 mm @, containing 3 meristeles
besides ventral root-bearing one; scales rather
light brown, not clathrate, to 1!/, mm wide. Sterile
acrophylls: stipes 12 cm long, sparsely covered with
small pale brown scales, only the smallest clathrate;
frond 75 cm long, rachis bearing scales with bullate
bases; pinnae to 14 by 2 cm, sessile, basiscopic base
subcordate or rounded, acroscopic broader and
rounded in lower pinnae, broadly cuneate in upper,
apex acuminate, edges very broadly crenate in
distal 1/3; veins fine, distinctly raised on both
surfaces; scales rather abundant on midrib and
veins of lower surface, bullate-acuminate. Fertile
pinnae of type mostly broken, largest intact one
(near base of frond) 7 cm long (in herb. MICH to
101/, cm), 3-5 mm wide, subsessile, base rounded,
slightly narrowed towards blunt apex.
Distr. Malesia: W. New Guinea. Only known
from the type.
Ecol. Steep slope in rain-forest, at 1200 m,
locally common.
10. Lomagramma sinuata C.CuHr. Svensk Bot.
Tidskr. 16 (1922) 98, f. 5; HoL_trum, Gard. Bull.
S.S. 9 (1937) 215; BACKER & PostH. Varenfl. Java
(1939) 153; CopeL. Philip. J. Sc. 78 (1949) 401;
Ho.trum, Blumea 14 (1966) 222. — L. sinuata f.
papuana C.Cnrr. Brittonia 2 (1937) 302. — Type:
KAUDERN 73, N. Celebes (BM; dupl. in BO).
Leptochilus cuneatus R.BONAP. Notes Pterid. 14
(1923) 453. — Type: Brooxs 467, Bencoolen,
Sumatra (P; dupl. in BM).
Lomagramma sp. HoLttum, Gard. Bull. S. S. 9
(1937) 219. — Fig. 10b-f, 12.
Rhizome often bearing 3-5 fronds close together
at wide intervals; scales on rhizome-apex distinctly
clathrate with very dark cell-walls, edges paler and
sometimes fringed. Bathyphylls: fronds with 10-12
pairs of pinnae have apical pinnae jointed to rachis;
rachis distinctly winged; middle pinnae to 9 by
286 FLORA MALESIANA [ser. II, voli
Fig. 13. Lomagramma melanolepis v.A.v.R. Young plant, bathyphylls stage. Cult. R.B.G. Kew, origin
NE. New Guinea (Photogr. R. VAN CREVEL).
1978]
LOMARIOPSIS GROUP (Holttum)
287
2 cm with broadly cuneate acroscopic base and
narrowly cuneate or slightly rounded basiscopic,
edges shallowly crenately lobed (more deeply
towards pinna-apex) with irregular small teeth on
the crenations, apex rounded to bluntly pointed.
Sterile acrophylls to 100 cm long with many
pinnae; middle pinnae 11 by 2 cm to 20 by 4 cm,
more or less stalked, acroscopic base broadly
cuneate, its edge forming a distinct S-curve, basi-
scopic base narrowly cuneate (sometimes rather
rounded in New Guinea), edges almost entire on
larger fronds, slightly and irregularly toothed
towards apex on smaller ones, texture thin, veins
fine and distinctly raised on both surfaces. Fertile
fronds: pinnae 10-25 cm long, 5—8 mm wide, on
stalks 2-8 mm long.
Distr. Malesia: S. Sumatra, Java, Lesser Sunda
Is. (Bali), Borneo, Celebes, Moluccas (Batjan),
New Guinea; Solomon Is.
Ecol. In forest near streams, up to 750 m.
Notes. The type, and some other specimens
from Celebes and Java, have pinna-stalks 5 mm or
more long, narrowly winged on the acroscopic
side (fig. 10e). Most specimens from Java and New
Guinea have pinnae almost sessile but do not differ
significantly in other ways, though some in New
Guinea (f. papuana, fig. 10b) are more rounded
basiscopically.
In 1937 (/.c., p. 219) I thought that most speci-
mens now ascribed to this species from Java
probably represented a distinct species, but the
specimens in the Bogor herbarium on which I
based this opinion did not include sterile acrophylls
from fully mature plants nor fertile fronds. In the
Rijksherbarium at Leiden are excellent specimens
of large sterile acrophylls with quite entire pinnae
3 cm wide, and fertile fronds with pinnae 6-7 mm
wide; these are indistinguishable from typical
L. sinuata except that the sterile pinnae are shorter-
stalked. Bathyphylls from West Java have more
deeply crenate pinnae with a more acute apex than
pinnae of fronds of a comparable size attributed to
L. sinuata in East Java; this may be due to environ-
mental factors. The species is probably now rare in
Java owing to destruction of lowland forest.
11. Lomagramma copelandii HoLttum, Gard. Bull.
S. S. 9 (1937) 201, pl. 9, 10; CopeL. Fern FI. Philip.
(1960) 274. — Type: RAMos & EDANo BS 28826,
Mt Binuang, Prov. Tayabas, Luzon (BO; original
at MAN lost).
L. pteroides var. negrosensis COPEL. in Elmer,
Leafl. Philip. Bot. 2 (1908) 393; v.A.v.R. Handb.
Suppl. (1917) 438. — Type: ELmer 10175, Negros
(MICH).
Bathyphylls to 15 cm long with apical lamina
continuous with rachis, larger ones with jointed
small terminal pinna; pinnae to c. 15 pairs, largest
61/, by 1.8 cm, acroscopic side broadly cuneate at
base, basiscopic rounded and narrower, edges
crenately lobed (lobes c. 5 mm) each lobe with 2-4
short teeth, largest pinnae toothed only towards
acute apex, on smaller fronds apex blunt. Sterile
acrophylls: stipe 15-25 cm; pinnae to 18 by 3 cm,
sessile, acroscopic base of middle pinnae broadly
cuneate to subtruncate, basiscopic rounded, edges
slightly undulate, not toothed, apex acuminate,
texture thin, veins conspicuous. Fertile fronds:
pinnae to 20 cm long and 4 mm wide, sessile, base
not dilated, whole lower surface except costa
soriferous, areoles long and narrow.
Distr. Malesia: Philippines (Luzon, Catan-
duanes, Samar, Bohol, Negros, Mindanao).
Ecol. Apparently in lowland forest, altitudes
not recorded.
Note. ELmer distributed, with at leact some
specimens of his collection no 16919, fertile pinnae
of Lomariopsis subtrifoliata with sterile fronds of
Lomagramma copelandii.
12. Lomagramma melanolepis v.A.v.R. Bull. Jard.
Bot. Btzg III, 5 (1922) 212; HoLtrum, Gard. Bull.
S. S. 9 (1937) 208. — Type: BeGuINn 1114, Ternate
(BO). — Fig. 10g-h, 13, 14.
Rhizome-scales 4-5 mm long, narrow, dark-
clathrate with paler edges bearing some hairs;
fronds often 3 close together. Bathyphylls (New
Guinea) with closely-placed strongly toothed
pinnae which are commonly c. 20 by 6 mm; apical
pinna jointed to rachis on fronds with 12 pairs of
pinnae. Sterile acrophylls: middle pinnae subsessile,
to 12 by 2 cm, rather thin with distinctly raised
veins, base rather broadly cuneate on acroscopic
side, narrower and cuneate to narrowly rounded
on basiscopic, edges entire or somewhat toothed
towards apex. Fertile fronds: pinnae to 9 cm long
and 2-3 mm wide, usually stalked.
Distr. Malesia: Moluccas (Ternate, Halma-
heira), New Guinea, d’Entrecasteaux Is.
Ecol. In lowland forest, to 900 m.
Note. This is very like L. sinuata, but has con-
sistently smaller pinnae of both bathyphylls and
acrophylls and narrower fertile pinnae. I have
included numerous New Guinea collections which
differ from the type in the very narrowly cuneate
basiscopic base of sterile pinnae. If further collec-
tions from the Moluccas should indicate that the
New Guinea plants are distinct, the latter will need
a new name.
13. Lomagramma brassii HoLtrumM, Blumea 14
(1966) 224. — Type: Brass 24947, Goodenough I.
(A).
Rhizome of adult plant 3-4 mm @, carrying
fronds in two longitudinal rows; scales to 3 by
1/, mm, clathrate with brown cell-walls. Sterile
fronds: stipe 3-6 cm long; lamina to 25 cm long
with 12-15 pairs of pinnae, apical one largest and
jointed to rachis; a few basal pinnae gradually
smaller and with rounded apices; middle pinnae of
type to 4'/, by 0.9 cm, of another collection 8 by
1.1 cm, sessile, basiscopic base narrowly cuneate,
acroscopic broadly so, apex tapered and acute,
edges with an acute falcate tooth corresponding to
each costal areole and irregular smaller inter-
mediate teeth; veins forming one series of costal
areoles and in the largest pinnae an irregular second
series; small dark bullate-acuminate strongly
clathrate scales abundant on lower surface of costa
and a few on veins. Fertile fronds: lamina c. 15 cm
long; pinnae 2 mm wide, largest 2'/,-51/, cm long,
apex rounded, stalks 1 mm long.
Distr. Malesia: New Guinea (Goodenough I.).
Two collections.
Ecol. In transitional oak-rain-forest, 900 m,
climbing to 2-3 m.
Note. This is very near L. melanolepis, but
smaller, and the only clear difference is in the
FLORA MALESIANA [ser. II, vol. 14
Fig. 14. Lomagramma melanolepis v.A.v.R. Part of plant shown in fig. 13; bathyphyll with winged rachis
and toothed pinnae which are all articulate (Photogr. R. VAN CREVEL).
1978]
slender rhizome of the adult stage of the plant. In
view of the clear difference between L. sinuata and
L. melanolepis, which is mainly one of size (and
which is maintained by plants growing side by side
in cultivation at Kew) I think it probable that
L. brassii is a distinct species.
A plant in cultivation at Kew (accession n. 020/
74-00297, J. R. WoopHAms), collected in the
vicinity of Sogere in Papua climbing on the trunk
of a dead Cyathea, agrees well with the above
description except that the pinnae are smaller:
sterile to 3!/, by 1.1 cm, fertile c. 1'/, by 0.3 cm. The
small size may be due to conditions of cultivation;
at least the plant confirms that the fertile condition
can be attained by quite small plants, a condition
otherwise unknown in the genus.
Doubtful species
Lomagramma guianensis (AUBL.) CHING, Amer.
Fern J. 22 (1932) 17. — Polypodium guianense
AUBL. Hist. Pl. Guian. 2 (1775) 962.
This species occurs in the Greater Antilles and in
South America from Guiana to southern Brazil.
In habit, frond-form, venation and articulated
pinnae it agrees closely with species of Loma-
gramma in Malesia and the Pacific. It differs in
the following characters: acrophyll frond-apex
deltoid, lobed, not pinna-like (as in bathyphylls of
Malesian species); small scales not bullate; para-
physes hair-like with a glandular apical cell; spores
with folded perispore as in Bolbitis and Lomariop-
sis. The paraphyses are much like the hairs on the
margins of small scales of Lomariopsis, but scales
on rhizome and frond are clathrate as in Bo/bitis
and Lomagramma. CHING included this species in
Lomagramma, but its differences from Malesian
LOMARIOPSIS GROUP (Holttum)
289
species are such that I doubt its genetic unity with
them; an independent origin in South America
seems to me more probable, in which case a new
genus might be established for it.
Lomagramma sorbifolia (WiLLD.) CHING, Lingn.
Sc. J. 12 (1933) 566; Ho_trum, Gard. Bull. S. S. 9
(1937) 220. — Aspidium sorbifolium WILLD. Spec.
PI. 5 (1810) 23. — Nephrodium sorbifolium (WILLD.)
Pres_, Rel. Haenk. (1825) 31, nomen tantum;
Ho.ttum, Novit. Inst. Bot. Uniy. Carol. Prag.
1968 (1969) 17.
WILLDENOW’s type is a detached bathyphyll; the
only locality given is “Ind. or.’’ When CHING
regarded WILLDENOW’s specimen as conspecific
with L. lomarioides (BL.) J.SM. he took a very broad
view of the latter species; certainly WILLDENOW’s
specimen is quite unlike bathyphylls of L. /oma-
rioides as described in the present work. It seems
to me probable that WILLDENOW’s specimen re-
presents the same species as the type of L. matthewii
(CHING) HoLtTruM, known from Assam, Tonkin
and Thailand, but this cannot be regarded as
certain, and I prefer to regard WILLDENOW’s name
as of doubtful application.
PRESL gave the name Nephrodium sorbifolium to
two bathyphylls, representing two distinct species,
collected by HAENKE in Luzon; I identify the
specimens as L. copelandii and L. pteroides.
Excluded
Lomagramma _ praestantissimuyn (BORY) GRIES.
Fl. Brit. W. Ind. (1864) 678 = Neurocallis praestan-
tissimum (Bory) FEE.
Lomagramma wilkesiana (BRACK.) COPEL. Philip.
J. Sc.3 (1908) Bot.32 = Teratophyllum wilkesianum
(BRACK.) HOLTTUM.
5. ELAPHOGLOSSUM
J.Sm. in Hook. J. Bot. 4 (1841) 148, nom. cons.; in Hook. Gen. Fil. (1842) t. 105A;
Moore, Ind. Fil. (1857) xvi; ibid. (1862) 351; J.SM. Hist. Fil. (1875) 125; Bepp.
Handb. Ferns Br. India (1883) 416; Curist, Farnkr. Erde (1897) 33; Neue
Denkschr. Allg. Schweiz. Ges. Naturw. 36 (1899) 1-159; Diets in E. & P. Nat. Pf.
Fam. 1, 4 (1899) 331; C.Cur. Ind. Fil. (1905) lii, 302; CopeL. Gen. Fil. (1947) 119;
Morton, Amer. Fern J. 45 (1955) 11; W. R. ANDERSON, Regn. Veget. 40 (1965) 18;
Ho.ttum, Blumea 14 (1966) 317-326. — Olfersia PREsL, Tent. Pterid. (1836) 232,
p.p. max., non RADDI. — Aconiopteris PRESL, Tent. Pterid. (1836) 236. — Acrosti-
chum (non L.) FEE, Hist. Acrost. (1845) 8, 27. — Dictyoglossum J.SM. Bot. Mag. 72
(1846) Comp. 18. — Acrostichum sect. Elaphoglossum Hook. Spec. Fil. 5 (1864)
195-241; Hook. & BAKER, Syn. Fil. (1868) 400. — Fig. 15-24.
Rhizome creeping, in Malesian spp. usually dorsiventral and bearing 2-ranked
fronds with a branch-bud at the base of each (some tropical American spp. with
fronds in more than 2 ranks and a few with radially organized rhizome); young
parts protected by scales which are usually cordate (often strongly) at the base, with
edges bearing short teeth by projection of the wall between 2 adjacent cells, or hairs
which may be of one or several cells, some cells usually glandular; outgrowth from
the rhizome (in all Malesian spp.) forming similarly scaly terete phyllopodia to
290 FLORA MALESIANA [ser. II, vol. 14
which fronds are + distinctly jointed (when dry, phyllopodia are usually darker
than stipes). Fronds simple, entire, dimorphous, stipitate or sessile, usually +
coriaceous, often with a colourless cartilaginous thinner edge, costa usually some-
what prominent and grooved on upper surface, rounded and + prominent on
lower; veins conspicuous or not, forked once or twice, all branches almost reaching
the margin, their tips in most spp. free and thickened, in a few spp. the vein-tips
joining each other in a series of arcs just within the margin; small scales + abun-
dant, persistent or not, on both surfaces, in a few species peltate. Fertile fronds with
smaller lamina than sterile, sometimes of different and distinctive shape, often with
longer stipes than sterile, the lower surface quite covered with sporangia except
for the thin decoloured margin and (in some species) a narrow decurrent part of
the base. Spores with well-developed folded perispore.
Type species: Acrostichum conforme Sw.
Distribution. More than 400 spp., throughout wetter parts of tropics and subtropics, with greatest
diversity on the Andes.
Ecology. Almost all Malesian species are epiphytes, though some will grow on mossy rocks in low
forest at high altitudes; one is reported growing on wet rocks in a stream-bed (E. resiniferum); few if any
grow in full exposure to the sun. They usually grow in association with other ferns and orchids on heavily-
laden tree-branches; when seen at a distance their simple entire fronds often resemble orchid leaves and
so may be overlooked. Few species occur in lowland forest, most at 1000-2500 m. The fronds of most are
somewhat fleshy, with thick cuticle, and some are very rigid when dried; those with thinnest fronds occur
in sheltered places, some on tree-trunks not far above ground level (E. melanostictum). The surface often
shows a bluish hue. Fronds are shed when old by breaking at the joint between phyllopodium and stipe,
where there is an internal change of structure though no true absciss-layer (see BELL, infra, 1951) and I
believe that most fronds probably persist for more than a year, some possibly 2 years or more (old ones
may bear a considerable growth of epiphyllous bryophytes). Fertile fronds are produced periodically,
probably in response to dry weather, as in other genera of this group. I had a plant of E. amblyphyllum in
cultivation in a hanging pot in Singapore for more than 20 years and only once saw a fertile frond on it,
perhaps because it was always watered on rainless days (the species was native locally on old trees in
mangrove but such trees are fast disappearing. R. M. LLoyp has discussed some aspects of the ecology of
tropical American species (Amer. Fern J. 60, 1970, 73-82); at high altitudes only about 50% are epiphytic.
Some species lack a joint at the base of a stipe; these were mostly epiphytic, not terrestrial.
Vegetative morphology and anatomy. P. R. BELL has published “Studies in the genus Elapho-
glossum’’ in five papers, based largely on his own observations of tropical American species (Ann. Bot.
n.s. 14, 1950, 545-555; ibid. 15, 1951, 333-346, 347-357; ibid. 19, 1955, 173-199; ibid. 20, 1956, 69-88). He
dealt with stelar structure in relation to habit, vascular supply to roots and branches in relation to bases of
fronds, the anatomy of fronds and structure of scales and hairs. He proceeded to arrange in series obser-
vations on 87 spp. relating to (a) stelar structure (dorsiventral with fronds in 2 or more ranks, or radially
symmetrical), (b) development of the joint, and aerenchyma of the phyllopodium, (c) scales on the frond;
and works out the frequencies of various combinations of characters, thereby indicating that the species of
Malaya (which alone of Malesian spp. he compared with American spp.) show predominant a condition
he regards as primitive, having a combination of dorsiventral rhizome with 2 ranks of fronds, phyllo-
podium with joint at its tip and aerenchyma near its base, and flat scales with basal attachment. So far as
I have been able to observe, the other Malesian species agree in the 2-ranked arrangement of the fronds,
which resemble that of young plants in the other genera of the group. I have not examined the aerenchyma
of phyllopodia; the external characters associated with this are difficult to see clearly on dried specimens
(and to examine them one needs to remove all scales) and should be studied by someone having access to
abundant living plants.
Scales are important diagnostically, both on rhizome and surface of fronds; size, shape and colour are
distinctive, also the nature of marginal hairs and the position of glandular cells. It appears that glandular
cells of many species can produce a resinous exudation which may persist after the scale has fallen or be-
come disintegrated. This also needs examination on living plants. A further comment on scales is given in
the section on Taxonomy below.
Venation is often not easy to distinguish, especially the form of the ends of the veins which can usually
only be seen if the frond is cleared with chloral hydrate. In a few species the ends of the veins anastomose
just within the margin but these do not constitute a natural group and PREsL’s genus Aconiopteris, based
on this character, cannot be maintained; probably not all cases of this kind have yet been recognized,
because of opacity of fronds.
Gametophyte. A. G. Stoxey and L. R. ATKINSON have published a study of gametophytes of 19
tropical American spp. (Phytomorphology 7, 1957, 275-292); these include one(E. gayanum (FEE) Moore)
belonging to the group of E. conforme, which shows no peculiar characters (see infra on subdivision of
1978] ; LOMARIOPSIS GROUP (Holttum) 291
genus). Prothalli are slow-growing and long-lived, ribbon-like with crisped wings, marginal rhizoids and
abundant short hairs with waxy caps; these wax-bearing hairs may be compared with glandular hairs on
scales of the sporophytes. Marginal rhizoids are reported by SroKEYy and ATKINSON as occurring also in
Selliguea Bory and in Grammitoid ferns; their presence is perhaps an adaptation to epiphytic growth
among mosses, and cannot be an indication of any close relationship between such very diverse genera as
Elaphoglossum, Selliguea and Grammitis. Archegonia and antheridia “‘conform in type to those of the
higher ferns’’. Gametophytes of Elaphoglossum are thus distinct from those of all other genera in the
present group. It may be relevant that NAYAR found narrow ribbon-like gametophytes in Microsorium
pteropus (BL.) CHING. (Polypodiaceae) whereas the type species M. punctatum (L.) Cope. has normal
cordate ones. NAYAR separated M. pteropus as type of a distinct genus Kaulinia, but there is no reason to
think that the two genera are not closely related (Taxon 13, 1964, 67-69).
Taxonomy. The generic name Elaphoglossum was proposed by ScHortrt in 1834 for certain specified
species of the composite genus Acrostichum, but he gave no description, and therefore his name, though
long accepted, does not comply with the present conditions of valid publication (Morton, /.c. 1955).
When PRESL attempted a more complete survey of the Acrostichoid ferns (1836) he adopted the name
Olfersia Rapp! for the species listed as Elaphoglossum by ScHoTT; but RADDI’s name was given originally
to the single species O. corcovadensis, which is now included in the earlier Po/ybotrya H. & B. PRESL also
established a new genus Aconiopteris for Acrostichum subdiaphanum Hook. & GREV., the only distinctive
character of which is that the veins join in a series of arcs along the margin. This character is not now
regarded as significant for generic separation, and the type species of Aconiopteris is now included in the
same genus as most of those listed as Olfersia by PRESL. Thus, according to a strict interpretation of the
Code, Aconiopteris is the correct name for the species which have been commonly called Elaphoglossum
for more than a century. The first valid publication of the name Elaphoglossum was by JOHN SMITH in
1841. A proposal was therefore made (ANDERSON 1965) for the conservation of Elaphoglossum J.SM. as
against Aconiopteris and this has been approved.
In 1845 FEE published his elaborate and finely illustrated work on the Acrostichoid ferns, in which he
criticized PRESL’s arrangement. He restricted the genus Ol/fersia PRESL to two species and redefined
Acrostichum to include the bulk of Prest’s Olfersia; for Acrostichum as restricted by PREsL (but excluding
a few species) he proposed the new name Chrysodium. He maintained Aconiopteris as a separate genus.
In 1846 JOHN SMITH published the name Dictyoglossum for tropical American species having anastomosing
veins, but a generic separation on this character is now regarded as unnatural. In 1857 Moore adopted
the name Elaphoglossum in the modern sense and transferred many names to it, but Hooker (1864)
reverted to a comprehensive Acrostichum with Elaphoglossum as a section. BEDDOME (1865 onwards)
followed Moore, and the name Elaphoglossum came into quite general use after the publications of
Crist (1897, 1899) and DieELs (1899).
Subsequent treatment of the genus in relation to other genera in systems of classification is dealt with in
the introduction to the present group of genera (supra p. 257). One aberrant suggestion should however be
mentioned. In his Monograph (1899, p. 17) CHrist recognized the isolation of Elaphoglossum among
Acrostichoid ferns, but pointed out similarities between the genera Elaphoglossum and Syngramma,
noting however the great difference between the bristle-like hairs which clothe the rhizome of Syngramma
and the scales of Elaphoglossum. Bower accepted a relationship between the two genera, and placed them
as simple-fronded derivatives of the same stock as Metaxya (The Ferns 1, 1928, 233-238). But the super-
ficial resemblance between Elaphoglossum and Syngramma is due to the kind of convergent evolution that
has occurred many times among ferns. It is clear that one general evolutionary trend which has occurred
along several separate lines is reduction from a branched frond to a simple one; Syngramma and Elapho-
glossum appear to be the ends of two quite different such lines. They differ greatly in spores as well as in
scales; probably a study of sporangia would show further differences. The association of Syngramma (an
exclusively Old World genus) and Elaphoglossum with Metaxya, which is an isolated monotypic south
American genus placed with some doubt in a separate subfamily of Cyatheaceae (Fl. Males. II, 1, 1963, 72)
seems to me highly improbable. I know of no other author who has seriously upheld Bower’s ideas on
these genera.
Subdivision of the genus. PrEsL (1836: 233-235) divided the species of his genus Olfersia into two
‘phalanges’: frons herbacea and frons coriacea. This is not a practicable arrangement. FEE (1845) divided
the species of his Acrostichum into two groups, Oligolepideae and Polylepideae. Though this also is not
satisfactory, it seems to me to be in the right direction, namely to use the scales on the fronds as a basis for
subdivision. The type species of the genus, E. conforme, comes into FEE’s Oligolepideae. If now we use
characters of the scales, and not their abundance, to distinguish E. conforme and its allies from the
species of FEE’s Polylepideae, we can have a clear-cut division; as almost all species of a re-defined
Polylepideae have in fact conspicuously scaly fronds, the name is not inappropriate. It appears to me that
this second division may be again subdivided on scale characters, as indicated below.
CurisT (1899) divided the genus into two ‘ordines’, Stenoneura and Condyloneura, each ordo being
divided into sections and subsections. The species of Stenoneura were said to have veins running to the
margin without thickened tips, whereas in Condyloneura the veins are described as having thickened tips.
In fact most species placed by CHRIST in Stenoneura are seen to have thickened vein-tips if the fronds are
cares with chloral hydrate, so that his main division is unreal; the minor subdivisions also are not more
useful.
The following conspectus is tentative, for which reason I have not formally proposed any new names
for subdivisions of the genus. The conspectus differs from that published by me in 1966 (Blumea 14: 319)
owing to comments received from W. R. ANDERSON, University of Michigan, to whom I express my
292 FLORA MALESIANA [ser. II, vol. 14
thanks; he has made a far more thorough study of tropical American species than I could attempt. No
doubt the present conspectus is over-simplified, and is not adequate to cover all tropical American species;
it is intended only as a step towards more light on a complex problem.
1. Scales on frond bearing some marginal cells which are swollen and glandular, usually also bearing
marginal hairs which consist of several cells, the terminal cell being glandular
group of E. conforme (Sw.) J.SM.
1. Scales on frond not bearing swollen glandular marginal cells except sometimes near the base; marginal
hairs, if present, each consisting of a single cell which is usually acicular and thin-walled.
2. Scales on frond flat, with marginal hairs. . .... .. . group of E. muscosum (Sw.) Moore
2. Scales on frond narrow, with reflexed edges (at least in basal part), thus bristle-like, lacking marginal
hairs hts OI, eA As Re Ae Me i Ara a group of E. spathulatum (BoRY) MoorRE
Malesian species. The earliest attempt at a comparative account of Malesian species was by BLUME,
who published full descriptions and excellent plates of those he knew in Java (Fl. Jav. Fil. 1829). But
BLUME adopted some earlier names, originally given to plants from other parts of the world which are
distinct from the Java species (e.g. Acrostichum decurrens, A. gorgoneum, A. viscosum). Some other early
names, notably E. conforme (described originally from St Helena) have also been used in too broad a
sense. The result has been considerable confusion in the use of such names by past authors, and without
reference to specimens one cannot always be sure of the sense in which such names have been used.
A few Malesian species are common in mountain forests, and have been frequently collected, especially
E. callifolium, E.angulatum and E. blumeanum. Most other species have been collected too little for a really
good assessment of their full range of variation, and of their geographical distribution. This is especially
the case in New Guinea, where (in Malesia) the genus has its greatest diversity, and several species are
known only from single collections, so that their descriptions may need subsequent modification. Further
collections of good sterile specimens would help considerably. It appears that the most widely distri-
buted Malesian species are E. angulatum (mountains of E. tropical Africa, Mascarene Is., Ceylon and
S. India, throughout Malesia) and E. callifolium (throughout Malesia and eastwards to Fiji).
Almost all Malesian species belong to the group of E. conforme in the conspectus. A few belong to the
group of E. muscosum, none to that of E. spathulatum, the nearest members of which are in Polynesia
(E. samoense BRACK., E. rapaense CopeL., E. societarum Cope.) and in Africa (about 6 spp.), a curious
distribution.
Key to Malesian species. Apart from separation of the few representatives of the group of E. mus-
cosum (nos 45-48) the key which follows is not an attempt at a natural arrangement, but only an attempt
to provide a means of identification, and is usable only for mature plants. Young plants often have fronds
different in shape from older ones; in general, fronds on young plants have a lamina more gradually
decurrent at the base and a broader apex than those on mature plants, and plants of immature size rarely
have fertile fronds. The key should serve to identify most mature plants, whether or not they have fertile
fronds. Fertile fronds are almost always narrower than sterile; in each species widths of the two are ina
fairly constant ratio. A distinctive feature of fertile fronds in some species is that their stipes are much
longer than those of sterile fronds, but this is not always a constant character. However, I believe that in
general there is a good contrast between species in which the stipes of the two kinds of fronds are of similar
length, and those in which the fertile stipes are twice (or more times) as long as the sterile ones.
Shape of fronds is always important, particularly the shape of the apex; there seems to be more varia-
tion in the base in some species. Size is also significant, but one must remember that plants of the same
species in situations differing in altitude or in exposure may differ considerably in size, also in texture.
Texture, and the degree of distinctness of veins, can be significant, especially extreme conditions, but many
species are in an intermediate state between very thick and rather thin. The development of the colourless
margin is certainly significant, though I think that a narrow margin of thick-walled colourless cells is
always present. An anatomical study to show the relationship between internal structure and external
form would be of interest, but is beyond the scope of the present work (an indication of the kind of
anatomical structure which can occur at the margin of a frond is given by BELL, Kew Bull. 14, 1960, 81).
KEY TO THE SPECIES
1. Small scales on frond stellate or elongate, their marginal hairs glandular or with a glandular apical cell,
thick-walled unicellular marginal hairs lacking; rhizome-scales various.
2. Lamina of sterile fronds rarely over 7 cm long; rhizome 1-2 mm @, fronds usually well-spaced on it.
3. Rhizome-scales light brown.
4. Sterile lamina elliptic, commonly 4—7 cm long, base decurrent as a narrow wing.
5. Fertile lamina spathulate, abruptly contracted at base and then decurrent as a narrow wing 2 cm
long; rhizome-scales tol mm wide ..........2.2........ _ 1. E. bolanicum
5. Fertile lamina narrowly elliptic, little decurrent; rhizome-scales 2 mm wide 2. E. habbemense
4. Sterile lamina ovate, almost same shape and size as fertile, c.2cmlong . . . . 3. E. pumilum
3. Rhizome-scales dark. . . .. . 4. E. hellwigianum
2. Lamina of sterile fronds usually more than 7 cm long; rhizome thicker.
6. Rhizome long-creeping with well-spaced fronds.
7. Apex of sterile fronds acute or short-acuminate.
8" Scales dark glossy, 1 mm/iwides.). 9) 292) ee ee. 5c eeu
8. Scales thin, medium brown, 2-3 mm wide ............... 6, E, angulatum
1978] LOMARIOPSIS GROUP (Holttum) 293
7. Apex of sterile frond rounded.
9. Rhizome-scales light brown; scales on frond red-brown . . . . 7. E. pullenii
9. Rhizome-scales dark brown; scales on frond dark, glossy (not known in no 0 9).
10. Rhizome-scales to 5 by 1’), mm; stipe of sterile fronds 4-15 cm long. . 8. E. sclerophyllum
10. Rhizome-scales shorter, less than 1 mm wide; stipe of sterile fronds 3-S cm. . 9. E. repens
6. Rhizome short, with closely- placed fronds.
11. Lamina of sterile frond decurrent to joint with phyllopodium or within 2 cm of joint.
12. Edge of sterile lamina conspicuously pale and thin, '/, mm or more wide.
13. Sterile lamina twice as wide as fertile.
14. Apex of sterile frond bluntly pointed or rounded, base cuneate and then decurrent for 4-7 cm
as a wing 1-2 mm wide on each side of costa . . . . 10. E. ophioglossoides
14. Apex of sterile frond short-acuminate, base narrowed evenly almost to joint with phyllo-
podium. . a. Peay ie a os . Il. E. latemarginatum
13. Sterile lamina little wider than fertile . . MEANS BE Say 08 pellucido-marginatum
12. Edge of sterile lamina not conspicuously pale ‘and thin.
15. Sterile frond acuminate or apiculate.
16. Rhizome-scales 10-15 by 3-4 mm, light brown, thin; lamina widest above middle
13. E. thamnopteris
16. Rhizome-scales much smaller, dark; lamina widest at middle.
17. Sterile frond c.22 by2cm . . stic © 01054927400, f 2. 143K idenburgensis
17. Sterile frond much larger, to 4Y) cm wide.
18. Apex of sterile frond acuminate.
19:, Sterne frond, 35-50 by: 3*/-—9 cm. 2°: 2 es as SP 1S: Exarchboldii
19. Sterile frond to 35 by2.3cm ... . . 27. E. angustifrons
18. Apex of sterile frond broad with a narrow apiculus 5 mm long . . . 16. E. apiculatum
15. Sterile frond not acuminate nor apiculate.
20. Sterile frond widest '/, from apex, narrowed rather abruptly to broad-pointed apex.
21. Rhizome-scales 15-20 mm long, 2 mm wide; fronds thick, veins obscure; costa broad and
little prominent on lower surface. . . . 17, E. annamense
21. Rhizome-scales not over 10 mm long; fronds rather thin, ‘veins distinct: costa slender and
prominent on lower surface.
22. Scales on surface of frond pale, bearing slender hairs 1 mm long; stipe of fertile frond
10-15cm . . 18. E. melanostictum
22. Scales on frond dark, bearing very short hairs; stipe of fertile frond 25-30 cm
19. E. heterostipes
20. Sterile frond widest about middle, or if above middle narrowed gradually to a rounded apex.
23. Sterile frond widest at middle, apex acute; fertile frond to 2 cm wide 20. E. calanasanicum
23. Sterile frond widest a little above middle, gradually narrowed to rounded apex; fertile fronds
HOU OVER! SICMWIGe 05 seo z : we, eae: Pee Ey. eel OrTishl
11. Lamina of sterile frond not thus decurrent.
24. Scales on rhizome dark, rigid, glossy.
25. Small scales on upper surface of frond dark, glossy, elongate, with dark rigid cylindrical margi-
nal hairs each consisting of one cell with a small thin-walled cell at its apex.
26. Scales on lamina 1-2 mm long; fronds with broadly rounded apex .. . . 22. E. apoense
26. Scales on lamina much smaller; apex of fronds not broadly rounded . . 23. E. vepriferum
25. Small scales on upper surface otherwise, at least some of their marginal hairs multicellular with
glandular apical cell.
27. Apex of sterile frond rounded.
28. Phyllopodia */,-1 cm long; lamina of sterile frond always c. 10 times as long as wide.
29. Scales on costa and stipe appressed, on lamina very small ....... 24. E. nigripes
29. Scales on stipe and costa spreading, on lamina 1-2 mm long, abundant
25. E. melanochlamys
28. Phyllopodia 11/,-2 cm long; lamina of sterile frond often proportionately broader
8. E. sclerophyllum
27. Apex of sterile fronds acute or acuminate.
30. Lamina of sterile frond c. 10 times as long as wide.
31. Rhizome-scales less than 1 mm wide, somewhat crisped; stipe of fertile frond not or little
longer than: that ‘of sterile’. i 2 >. see ee Dee . . 26. E. nesioticum
31. Rhizome-scales more than 1 mm wide, flat; stipe of fertile frond usually twice as long as
sterile.
327 Sterile frond,37/5—5 cm wide: 35-50 COIONS Pianmesiase . home . 15. E. archboldii
32. Sterile frond to 2.3 cm wide, 35cmlong .......... ... 27. E. angustifrons
30. Lamina of sterile frond c. 5 times as long as wide. :
33. Rhizome bearing fronds 5-10 mm apart; scales 6 by 1 mm; sterile lamina widest below
middle, rather abruptly contracted at base, apex short-pointed . ... 5. E. brunneum
33. Rhizome bearing fronds close together; scales 10 by 11/, mm; sterile lamina widest at middle,
base rather narrowly cuneate, apex acuminate ........... 28. E. pallescens
24. Scales on rhizome medium to light brown, mostly rather thin.
294 FLORA MALESIANA [ser. II, vol. 14
34. Lower surface of sterile lamina covered with a felt of narrow scales 2-3 mm long
29. E. arachnoideum
34. Lower surface of lamina bearing smaller, usually appressed, scales.
35. Fronds of adult plants widest above middle, with rounded apices and decurrent bases.
36. Fronds lacking a fringe of spreading scales.
37... Rhizome-scales less than sl Cmimewide oi. sg ke ee ee re 30. E. stenolepis
37. Rhizome-scales 2mm or more wide . - .-...... - . . . 31. E. amblyphyllum
36. Fronds bearing a fringe of spreading scales. ............ 32. E. luzonicum
35. Fronds of adult plants, if widest above middle, rather much narrowed upwards, apex not
broadly rounded.
38. Sterile fronds with distinct thin pale edge which bears abundant scales 1—1'/, mm long, per-
sistent on young plants, mostly lost on old ones ........ . . 32. E. luzonicum
38. Sterile fronds without a fringe of scales.
39. Sterile lamina twice as long as wide (to 12 by 6 cm); stipe of fertile frond as long as sterile
frond WithtitsiStipess tec ee ee SPER: cchcenseg ats : 33. E. brevifolium
39. Not this combination of characters.
40. Apex of sterile frond distinctly pointed, point broad or acute.
41. Sterile frond widest in distal half, base very gradually and evenly narrowed, stipe to 5 cm
ORES eT te gee ee ne re aaah eee . 19. E. heterostipes
41. Sterile frond widest at or below middle, base otherwise, stipe in most cases longer.
42. Pale margin of sterile frond thin, flat, '/; mm wide or nearly so.
43. Fronds widest below middle, narrowed gradually to apex; veins distinct
34. E. negrosensis
43. Fronds widest at middle, apex obtuse-angled; veins not distinct 35. E. planicosta
42. Pale margin of sterile frond very narrow, usually reflexed.
44. Scales to 1!/, mm wide, not flat.
45. Scales straight and rather stiff with inrolled edges; frond commonly 35 cm or more
lone narrowly elliptic ©. 0 oe ee gE rE
45. Scales thin, crisped throughout, with long twisted hair-points; frond rarely 30 cm long,
broadly elliptic .. . - - ie) en perme mae 5-13, COMMA
44. Scales wider, flat or little crisped, edges not inrolled.
46. Stipe of fertile frond not much longer than stipe of sterile.
47. Scales commonly 20-25 by 2!/,--3 mm .......... . 38. E. malayense
47. Scales not over 10 mm long.
48. Rhizome-scales with many spreading marginal hairs; scales on lamina to 1 mm long,
TIATEOW tne ere ok a eee ee, ee a
48. Rhizome-scales with few marginal hairs; scales on lamina stellate, under '/, mm long.
49. Costa broad and not prominent beneath; rhizome-scales firm 41. E. novoguineense
49. Costa prominent beneath; rhizome-scales thin, with isodiametric cells
43. E. favigerum
46. Stipe of fertile frond at least twice as long as stipe of sterile.
50. Scales 6 by 2!/, mm, sterile frond short-acuminate .. . . 39. E. recommutatum
50. Scales to 15 by 2 mm; sterile frond broadly pointed ... . . 42. E. robinsonii
40. Apex of sterile frond distinctly rounded.
51. Rhizome-scales 3-5 mm long; fertile frondc.8 by3.3cm ... . 7. E. pullenii
51. Rhizome-scales 10-15 mm long; fertile frond at least 6 times as long as wide.
52. Spores 52-59 um long with broad perispore of few folds; stipe of fertile frond twice as
long as that of sterile; edge of lamina thick, deflexed; lamina greenish when dry.
53. Lamina of sterile frond to 30 by 5 cm, stipe 5-8 cm long. . . . 44. E. sumatranum
53. Lamina of sterile frond 35-55 cm long, stipe 14-18 cm long . 45. E. indrapurae
52. Spores c. 37 um long with narrow much-folded perispore; stipe of fertile fronds not much
longer than of sterile; pale edge of lamina thin; fronds brownish when dry, thick, rigid
46. E. spongophyllum
1. Small scales on frond (except near edge) + orbicular, pale, with marginal teeth or hairs each of one
thick-walled cell, not glandular; rhizome-scales always narrow, rigid, dark, glossy.
53. Scales on upper surface conspicuously stellate; sterile fronds often to 3 cm wide.
54. Scales on lower surface very small, each with 3-5 marginal hairs 1/,-1 mm long spreading away
ELOUITTONG-SHTACCIIEE ELA te uals soe ab-S sunsets pcg, tae oe eee 47. E. blumeanum
54. Scales on lower surface with a distinct lamina (flat or not) bearing more slender and shorter
53. Scales on upper surface with very short marginal hairs; sterile fronds c. 1 cm wide
50. E. resiniferum
1978]
LOMARIOPSIS GROUP (Holttum)
295
1. Elaphoglossum bolanicum RosensT. in Fedde,
Rep. 12 (1913) 180; v.A.v.R. Handb. Suppl.
(1917) 427; Copev. Philip. J. Sc. 78 (1949) 403. —
Type: Krysser B.62, Bolan Mts, NE. New Guinea
(S-PA). — Fig. 15.
Rhizome \ong-creeping, 1—-1'/, mm @; fronds
1-2 cm apart; scales thin, light brown, to 3 by
1 mm, edges with few short hairs; phyllopodia
6-8 mm long. Sterile frond: stipe 2—3'/,cm; lamina
4-7 cm by 9-13 mm, widest at or below middle,
gradually narrowed distally, apex acute, towards
base similarly narrowed and then decurrent as a
narrow wing for 10-15 mm; pale thin edge distinct ;
veins visible but not prominent; scales on both
surfaces minute, dark brown, with short spreading
hairs. Fertile frond: stipe 4-6 cm; lamina 21/,-3 cm
by 9-12 mm, widest !/; from base, narrowed to
blunt apex and more abruptly to base, with nar-
row basal wing as sterile.
Distr. Malesia: NE. New Guinea (3 collections).
Ecol. At 2400-3000 m.
Note. The type specimen has only one sterile
frond, which is smaller than the fertile ones; sterile
fronds are described above from the other collec-
tions, which agree exactly in rhizome-scales and in
fertile fronds with the type.
Fig. 15. Elaphoglossum bolanicum ROseENST. a.
Habit, 2 sterile fronds, 1 fertile, x 2/3, b. scale from
rhizome, x 13, c. scale from surface of frond, <x 27
(HOOGLAND & SCHODDE 7699).
2. Elaphoglossum habbemense CopeL. Univ. Cal.
Publ. Bot. 18 (1942) 226; Philip. J. Sc. 78 (1949)
403, pl. 7. — Type: Brass 9083, W. New Guinea,
Lake Habbema (GH).
Rhizome long-creeping, c. 2 mm @, fronds
1-2 cm apart; scales light brown, translucent,
closely imbricating, 3 by 2 mm, ovate, acute,
subentire; phyllopodia to 8 mm long. Sterile frond:
stipe 2~3 cm; lamina 3'/,-6 cm by 8-15 mm, thinly
coriaceous to rigid, elliptic, apex bluntly pointed,
base narrowly cuneate and then decurrent as a
wing 10-15 mm; thin edge distinct, narrow; veins
visible or not; costa slightly prominent beneath;
scales on costa beneath rather persistent, narrowly
acuminate from a broad base, '/,-1'/, mm long;
scales on lower surface of lamina stellate, smaller.
Fertile frond: stipe 1*/,-3'/, cm; lamina 21/,-
4'/, cm by 7-10 mm, shape as sterile.
Distr. Malesia: New Guinea (3 collections).
Ecol. Ina cushion of hepatics on exposed tree-
branch, 3225 m.
3. Elaphoglossum pumilum LAM & VERHEY,
Blumea 5 (1945) 559, f. 2. — Type: MoNnop DE
FROIDEVILLE 253, Lompobatang, Bonthain, S.
Celebes (L; dupl. in BO).
Rhizome slender, creeping, bearing fronds
/,-1 cm apart; scales brown, thin, entire, ovate-
acute, to 2'/, mm long; phyllopodia 5 mm long.
Sterile frond: stipe 2 cm; lamina to 2 by 1 cm,
thinly coriaceous, ovate, base shortly decurrent as a
narrow wing, apex blunt, rounded; thin pale edge
narrow, + reflexed; veins slightly prominent be-
neath; costa hardly prominent on lower surface;
no superficial scales seen. Fertile frond: stipe
3'/,-6 cm; lamina to 20 by 9 mm, shape as sterile.
Distr. Malesia: S. Celebes. Only known from
type collection.
Ecol. At 2500 m.
4. Elaphoglossum hellwigianum Rosenst. Nova
Guinea 8 (1912) 731; v.A.v.R. Handb. Suppl.
(1917) 423; CopEL. Philip. J. Sc. 78 (1949) 403. —
Type: VON ROMER 1273, Hellwig Mts, W. New
Guinea (S-PA; dupl. in BO).
Rhizome 2 mm @, fronds c. 1 cm apart; scales
rigid, dark, glossy, narrow, acuminate, 2-3 mm
long, edges with short teeth or sparse short hairs;
phyllopodia not distinct. Sterile frond: stipe 1-
1'/, cm; lamina rigid-coriaceous, 3 by 1!/, cm,
almost elliptical, base slightly decurrent, apex
broadly pointed (fronds on a young plant obovate
with much-decurrent bases); thin pale edge distinct;
veins not visible; costa slightly prominent near
base beneath; many scales on lower surface of
young frond (mostly caducous), dark, elongate,
with some marginal hairs. Fertile frond: stipe 3 cm;
lamina 11/, by 1 cm, elliptic, base hardly decurrent;
small dark glossy scales abundant among sporan-
gia.
Distr. Malesia: W. New Guinea. Only known
from type collection.
Ecol. Probably at 3000 m or more.
5. Elaphoglossum brunneum CopeL. Univ. Cal.
Publ. Bot. 18 (1942) 226; Philip. J. Sc. 78 (1949)
405, pl. 10. — Type: Brass 12808, Idenburg
River, W. New Guinea (GH).
Rhizome bearing fronds 5-10 mm apart; scales
dark brown, glossy, rigid, 6 by 1 mm, narrowed toa
hair-tip; phyllopodia 10 mm long. Sterile frond:
stipe 8-10 cm; lamina rigid, to 19 by 4 cm, widest
below middle, + abruptly narrowed at base and
296 FLORA MALESIANA [ser. I, voleue
ii
ll
Ii
/]
/
ae
Fig. 16. Elaphoglossum commutatum (METT. ex KUHN) v.A.v.R. a. Habit, x 1/,. — E. angulatum (BL.)
Moore. b, Habit, x 4/,, c. margin of sterile frond, x 5 (a SURBECK 277, b-c L 908.331-949).
1978]
LOMARIOPSIS GROUP (Holttum)
297
then slightly decurrent, more gradually to acute
but not acuminate apex; pale edge c. '/, mm wide;
veins just distinct; costa prominent and rounded
beneath; scales on both surfaces appressed, dark,
less than 1 mm long, elongate with a few short
marginal hairs. Fertile frond: stipe 18 cm; lamina
12 by 2.2 cm, shape as sterile.
Distr. Malesia: W. New Guinea. Only known
from type collection.
Ecol. Low epiphyte in rain-forest at 1200 m.
6. Elaphoglossum angulatum (BL.) Moore, Ind.
Fil. (1857) 5; v.A.v.R. Handb. (1908) 713; Suppl.
(1917) 423; C.Cur. Gard. Bull. S. S. 7 (1934) 290;
BACKER & PosTH; Varenfl. Java (1939) 250;
TARD.-BL. & C.Cur. Fl. Gén. I.-C. 7, 2 (1941) 541;
CopEL. Philip. J. Sc. 78 (1949) 405; Fern FI. Philip.
(1960) 278; SLEDGE, Bull. Brit. Mus. (Nat. Hist.)
4 (1967) 83. — Acrostichum angulatum BL. En. Pl.
Jav. (1828) 101; Fl. Jav. Fil. (1828) 25, t. 6; Fee,
Hist. Acrost. (1845) 32; Racts. Fl. Btzg 1 (1898)
47. — Olfersia angulata PrRest, Tent. Pterid.
(1836) 234. — Type: BLUME, Java (L).
E. minahassae v.A.v.R. Handb. Suppl. (1917)
527; Bull. Jard. Bot. Btzg II, 28 (1918) 24. —
Acrostichum conforme (non Sw.) Curist, Ann.
Jard. Bot. Btzg 15 (1898) 174, p.p. — Type:
Koorpbers 17097, Menado, Celebes (BO).
E. dolichocaulon v.A.v.R. Bull. Jard. Bot. Btzg
Ill, 5 (1922) 203; BACKER & PosTH. Varenfl. Java
(1939) 250. — Type: JEswiET 348, Mt Jang (BO).
E. ogatai C.Cur. Dansk Bot. Ark. 9, 3 (1937) 67.
— Type: OGATA 59, Taiwan (BM).
E. alstonii TARD.-BL. Not. Syst. Paris 15 (1959)
433. — Acrostichum laurifolium (non THOUARS)
Fee, Hist. Acrost. (1845) 36, t. 7, f. 1. — Type:
SIEBER, Mauritius (P).
E. laurifolium [non (THOuARS) Moore] BEDD.
Ferns S. India (1866) t. 200. — E. latifolium [non
(Sw.) J.SM.] Bepp. Handb. Ferns Br. India (1883)
416, p.p. incl. f. 248. — Fig. 16b, c.
Rhizome l\ong-creeping, bearing fronds 1-3 cm
apart; scales thin, light brown, 4 by 2-3 mm,
triangular, tip blunt to acute, edges subentire;
phyllopodia 1-2 cm long. Sterile frond: stipe 5-
15 cm, when young scaly as rhizome; lamina thin
but firm, 11 by 3 to 28 by 5 cm, narrowly elliptic
(largest with sides parallel in middle part), apex
short-acuminate, base slightly decurrent; thin edge
nearly 1 mm wide, translucent; veins at a rather
wide angle, distinct on lower surface, their ends
joining a thickened submarginal band in which they
bifurcate and form a + continuous vascular com-
missure; costa not very prominent beneath;
mature fronds glabrous except for a few scales on
or near costa beneath, as those on stipe but smaller.
Fertile frond: stipe to 18 cm or more long; lamina
13 by 2.2 to 22 by 3/, cm, base more abruptly
contracted than sterile, thin edge somewhat nar-
rower.
Distr. E. tropical Africa, Madagascar, Réunion,
Ceylon, S. India, Tonkin, Taiwan; throughout
Malesia on higher mountains (not in the Malay
Peninsula); New Hebrides.
Ecol. Epiphyte in low forest and sometimes on
mossy rocks in + sheltered places, at 2000-3500 m.
Note. The sterile frond of the Bogor type speci-
men of E. minahassae has a damaged apex which
has a falsely rounded appearance; a Leiden speci-
men of the same number has fronds typical of
E. angulatum. This is the most widely distributed
Malesian species.
7. Elaphoglossum pullenii Ho_trum, Blumea 14
(1966) 324. — Type: PULLEN 5034, partim, NE.
New Guinea (L).
Rhizome creeping, fronds in each rank 1-1"/, cm
apart; scales light brown, firm, ovate-acuminate,
5 by 1'/, mm, edges with few hairs; phyllopodia
1—-1'/, cm long. Sterile frond: stipe slender, 13-14 cm
long, glabrescent; lamina thick, rigid, to 12 by
3 cm, almost evenly elliptical, base shortly decur-
rent, apex narrowly rounded; thin pale edge
c. '/, mm wide; veins + distinct, not prominent;
costa beneath broad, pale, hardly prominent;
scales on lower surface red-brown, to 1 mm long,
narrow with long marginal hairs near base, smaller
ones stellate. Fertile frond: stipe 14 cm; lamina 6 by
2 to 8 by 3.3 cm, widest below middle, base
abruptly narrowed and then decurrent 1 cm, apex
broadly rounded.
Distr. Malesia: E. & NE. New Guinea.
Ecol. Epiphyte, low on trees in mossy forest at
3140-3300 m, growing with FE. angulatum, a
specimen of which was collected with the type.
8. Elaphoglossum sclerophyllum v.A.v.R. Nova
Guinea 14 (1924) 22; CopEL. Philip. J. Sc. 78 (1949)
405. — Type: Lam 1793, W. New Guinea (BO;
dupl. in L, U).
E. fuscum CopeLt. Univ. Cal. Publ. Bot. 18
(1942) 226; Philip. J. Sc. 78 (1949) 406, pl. 12. —
Type: Brass 9088, W. New Guinea (MICH; dupl.
in BM, BO, GH, K, L).
E. laticuneatum Cope. Univ. Cal. Publ. Bot. 18
(1942) 226; Philip. J. Sc. 78 (1949) 405, pl. 9. —
Type: Brass & MEeEtER-DReEES 10032, W. New
Guinea (GH).
Rhizome creeping, fronds 1-2 cm apart; scales
rigid, dull brown to glossy dark brown, 4-6 by
1-11/, mm, apex acuminate, edges with teeth or
stiff hairs; phyllopodia 11/,-2 cm long. Sterile
fronds: stipe 4-15 cm long, pale, rather persistently
scaly as rhizome; lamina thick, rigid, variable in
shape, commonly 10-13 by 2!/,-3'/, cm, on some
plants 8-10 by 4 cm, widest about middle or above
it, apex narrowed to rounded tip, base cuneate and
+ decurrent (more so in small than in large fronds);
thin edge pale, !/, mm wide; veins distinct or not;
costa broad and slightly prominent beneath; scales
on both surfaces at first abundant, to 1 mm long,
dark, glossy, appressed, usually with some
spreading marginal hairs. Fertile frond: stipe
15-30 cm; lamina commonly 9-14 cm long, 2-
2'/, cm wide, of one specimen 8 by 4 cm.
Distr. Malesia: New Guinea.
Ecol. On terrestrial moss-cushions or epiphytic
in open thickets at c. 3000 m (several collections).
Note. A specimen at Leiden from W. New
Guinea (VERSTEEGH BW 12609, partim, Arfak
Mts, 1750 m) has apparently a short thick rhizome
with close fronds which are larger than those of the
type (sterile to 18 by 4, fertile to 20 by 2 cm) but in
shape and scaliness very similar. Brass collections
from near Lake Habbema at 2000 m (11038,
13040) are similar. These may constitute a distinct
species which occurs at about 2000 m.
298
FLORA MALESIANA
[ser. Il, voku?
9. Elaphoglossum repens CopeL. Univ. Cal. Publ.
Bot. 18 (1942) 226; Philip. J. Sc. 78 (1949) 406,
pl. 11. — Type: Brass 12124, W. New Guinea,
near Idenburg River (MICH; dupl. in GH, L).
Rhizome 2 mm @, creeping; fronds !/,-2 cm
apart; scales dark brown, glossy, rigid, entire, to
3 by 2/,; mm, narrowly acuminate; phyllopodia
1 cm long. Sterile frond: stipe slender, pale,
3-8 cm long; lamina thinly coriaceous, 12 by
2!/,-3 cm, widest at middle, base + decurrent,
apex gradually narrowed to a + broadly rounded
tip; pale thin edge distinct; veins not prominent,
sometimes distinct; costa slightly prominent
beneath; surfaces glabrescent. Fertile frond: stipe
9-14 cm; lamina probably to 9 by 1*/, cm (larger
frond broken), apex narrowly rounded, base a
little decurrent.
Distr. Malesia: W. New Guinea. Only known
from type collection.
Ecol. Frequent low epiphyte in mossy forest,
1800 m.
10. Elaphoglossum ophioglossoides (GoLDM.) HoL-
TruM, Kalikasan, Philip. J. Biol. 3 (1974) 197. —
Acrostichum ophioglossoides GOLDM. Nova Acta
19, Suppl. 1 (1843) 451. — Type: MEYEN s.n.,
Manila (B).
E. merrillii Curist, Philip. J. Sc. 3 (1908) Bot.
275; v.A.v.R. Handb. Suppl. (1917) 424; Copev.
Fern Fl. Philip. (1960) 279. — Type: MERRILL
5853, Mt Halcon, Mindoro (P; dupl. in GH).
Rhizome short, fronds very close; scales medium
brown, rather thin and crisped, hair-pointed, to
15 by 1 mm, edges with a few spreading hairs;
phyllopodia less than 1 cm. Sterile frond: stipe
3-8 cm long with a wing c. 1 mm wide throughout,
edge of wing pale cartilaginous as edge of lamina;
lamina coriaceous, to 26 cm long, largest fronds
31/,-5 cm wide, usually widest above. middle,
narrowed gradually to base and more abruptly to
broadly pointed or almost rounded apex; thin pale
edge c. +/, mm wide; costa beneath slightly promi-
nent near base, distally flat; veins faintly visible;
lower surface glabrescent, residual scales stellate,
small. Fertile frond: stipe to 8 cm; lamina c. 10 by
11/,-2 cm.
Distr. Malesia: Philippines (Luzon, Mindoro,
Samar).
Ecol. In forest, at 500 m, and probably higher.
Note. In ‘Reise um der Erde’ 2 (1855) 264-270,
MEYEN described his ascent of Mt Sembrano above
Laguna (480-510 m) from Manila; he appears not
to have climbed higher mountains. The above
description is taken mainly from the type collection
of E. merrillii (P, GH); MEYEN’s type is smaller but
certainly represents the same species.
11. Elaphoglossum latemarginatum HOoLtTum,
Blumea 14 (1966) 322. — Type: Brass 24896,
Papua, Goodenough I. (A).
Rhizome short; scales 8-10 by 11/, mm, dull
medium brown, rather thin, acuminate, apices
sometimes a little twisted, edges with scattered
hairs; phyllopodia 5 mm long. Sterile frond: wing-
less stipe pale, c. 1 cm long; lamina thinly coria-
ceous, 19-23 by 2.6—-3.3 cm, widest 4/, from apex,
narrowed very gradually to base which forms a
gradually narrowing wing so that junction with
stipe is indistinct, apex shortly acuminate; thin
pale edge '/, mm wide; veins distinct on lower
surface; costa pale, not prominent on lower sur-
face; scales on surface appressed, + stellate with
several short arms, mostly less than 1 mm @
including arms. Fertile frond: stipe 4 cm; lamina
17 by 1.6 cm, shape as sterile, thin pale margin
1 mm wide.
Distr. Malesia: E. New Guinea (Goodenough
I.). Only known from type collection.
Ecol. Common low epiphyte on mossy trees,
1600 m.
12. Elaphoglossum pellucido-marginatum (CHRIST)
C.Cur. Bot. Jahrb. 66 (1933) 65. — Acrostichum
pellucido-marginatum CHRIST, Verh. Naturf. Ges.
Basel 11 (1895) 255. — Acrostichum gorgoneum
(non KAULF.) CuHRist, Ann. Jard. Bot. Btzg 15
(1898) 175, p.p.; v.A.v.R. Handb. (1908) 712, p.p.
— Type: SARASIN 947, Celebes (BAS).
Rhizome-scales 5 by 1 mm, medium brown,
ovate-acute; phyllopodia 5-10 mm long. Sterile
frond: stipe (unwinged) to 2!/, cm; lamina of
largest fronds 17 by 11/2 to 35 by 4 cm, firm but
not thick, widest in upper third, gradually nar-
rowed to decurrent base and more abruptly to
narrowly rounded or broadly pointed apex; trans-
lucent edge c. 3/, mm wide; veins not very distinct,
their free apices thickened; scales on lower surface
very small, stellate. Fertile frond: stipe 31/.-10 cm;
lamina almost as large as sterile, apex sometimes
more distinctly rounded.
Distr. Malesia: North to SE. and SW. Celebes.
Ecol. Epiphyte, at 1200-2400 m.
Note. The species E. gorgoneum (KAULF.)
BRACK. is confined to Hawaii (see p. 313).
13. Elaphoglossum thamnopteris HoLTTuM, Blumea
14 (1966) 326. — Type: Brass 22990, Papua, Mt
Dayman, 1650 m (A).
Rhizome short; scales 10-15 by 3-4 mm, rather
thin, light brown, subentire, not acuminate; phyl-
lopodia c. 1 cm long. Sterile frond: lamina rather
thin, of specimens seen to 57 by 51/, cm (to 80 cm
long, fide BRASS), widest above middle, very gradu-
ally narrowed to joint with phyllopodium, apex
acuminate; thinner edge hardly decoloured; veins
distinct and slightly prominent on both surfaces,
their apices joining in a series of submarginal arcs
as in Asplenium nidus; costa prominent beneath
throughout; scales on surfaces scattered, mostly
under 4/, mm, with a few short marginal hairs and
spherical cells near point of attachment. Fertile
frond: stipe 15 cm; lamina 25 by 2 cm, base long-
decurrent, apex shortly blunt-pointed.
Distr. Malesia: New Guinea, Ceram.
Ecol. Epiphyte (type in Nothofagus forest), at
1500-1700 m.
Note. A specimen from 2900 m in NE. New
Guinea (T. G. WALKER 7571) has fronds to 35 by
4 cm, thicker than those of the type, but appears to
belong to this species.
14. Elaphoglossum idenburgensis HOLTTUM,
Blumea 14 (1966) 321. — Type: Brass 12283,
W. New Guinea (GH; dupl. in MICH). _
Rhizome short; scales dark, glossy, rigid, to 4
by 2 mm; phyllopodia 12-17 mm long. Sterile
frond: lamina rigid, decurrent to joint with
phyllopodium, 211/,-1.8 cm, widest about middle,
1978]
LOMARIOPSIS GROUP (Holttum)
299
base rather narrowly decurrent, apex acuminate;
thin margin not decoloured, narrow, reflexed;
veins rather obscure; costa prominent beneath;
scales on both surfaces sparse, dark, appressed,
usually entire. Fertile frond: stipe 6 cm; lamina 12
by 1.1 cm, base decurrent, apex bluntly pointed.
Distr. Malesia: W. New Guinea. Only known
from type collection.
Ecol. In mossy forest on old log, 1800 m.
15. Elaphoglossum archboldii CopeL. Univ. Cal.
Publ. Bot. 18 (1942) 226; Philip. J. Sc. 78 (1949)
404, pl. 8. — Type: Brass 13220, W. New Guinea
(GH).
Rhizome short; scales dark, glossy, rigid, flat,
ovate, 4-5 by 1—-11/, mm; phyllopodia 2-4 cm long.
Sterile frond: unwinged stipe 1—3 (to 8?) cm long,
above which is a narrowly winged part 4-20 cm
long below lamina; lamina thinly coriaceous,
25-50 by 2'/,-5 cm, widest about middle, gradually
narrowed to narrowly cuneate base, apex acumi-
nate; thin edge not distinct; veins slightly promi-
nent, their ends joining in submarginal arcs (at
least near base of frond); costa prominent beneath;
scales on lower surface not persistent, very small,
stellate with arms of 2-3 cells. Fertile frond: stipe
10-30 cm long; lamina 20-30 cm long including a
narrow sterile decurrent base, 2.2 cm wide.
Distr. Malesia: W. New Guinea.
Ecol. Epiphytic at 850-2000 m (type, at 850 m, a
low epiphyte; specimen from 2000 m an epiphyte in
moss cushions).
16. Elaphoglossum apiculatum HoL_trum, Blumea
14 (1966) 320. — Type: Brass 13624, W. New
Guinea (GH).
Rhizome short; scales dark, glossy, rather broad
(no complete ones seen); phyllopodia 11/,-2 cm
long. Sterile frond: unwinged stipe less than 2 cm
long; lamina (including basal wing) to 30 by 4.7 cm,
thinly coriaceous, the main part (18 cm long)
elliptic with a narrow triangular apiculus 5 mm
long, at the base rather abruptly narrowed to the
basal wing; thin edge not distinct; veins slightly
prominent on both surfaces, sometimes anasto-
mosing at their tips; costa strongly prominent
beneath; scales on lower surface of lamina not
seen, on lower surface of costa small, stellate, also
a few larger appressed ovate dark glossy scales
hardly 1 mm long. No complete fertile frond seen;
one frond fertile in distal 8 cm, this fertile part
contracted, 2.3 cm wide, basal part shaped as
sterile but narrower.
Distr. Malesia: W. New Guinea. Only known
from type collection.
Ecol. Epiphyte in rain-forest, at 900 m.
Note. This specimen is near FE. archboldii, but
differs in the shape of both apex and base of
sterile fronds, characters which are usually dis-
tinctive in this genus.
17. Elaphoglossum annamense C.CHR. & TARD.-
BL. Not. Syst. Paris 8 (1939) 209. — Type:
POILANE 23783, S. Vietnam (P).
E. decurrens var. crassum C.Cur. Gard. Bull.
S.S. 7 (1934) 290, p.p. — Fig. 17.
Rhizome short; scales dense, to 20 mm long,
2 mm wide at base, distally very narrow, hair-
pointed, somewhat red-brown, older ones darker
Fig. 17. Elaphoglossum annamense C. CHR. &
TARD.-BL. Three sterile fronds, one fertile, < +/,
(KOSTERMANS 2412).
300 FLORA MALESIANA [ser. II, vol. 14
Fig. 18. Elaphoglossum norrisii (HooK.) BEDD. a. Plant with 1 fertile and 2 sterile fronds, x 1/3, b—-c. scales
from lower surface of frond, x 40. — E. melanostictum (BL.) Moore. d. Plant with 1 fertile and 3 sterile
fronds, x 1/2, e. scale from lower surface of frond, x 32 (a BACKER 10998, d RAAP 236).
1978] LOMARIOPSIS
and dull, with some slender marginal hairs;
phyllopodia 1-1'/, cm long. Sterile frond: stipe to
14/, cm long; lamina rather thick, rigid when dry,
to 30 by 5-6 cm, widest above middle, tapering
gradually to decurrent base and rather abruptly to
broadly pointed or slightly rounded apex; thin
pale edge very narrow; veins not distinct; costa
beneath broad, slightly prominent; scales on lower
surface sparse, stellate, under | mm @ including
rays, with some spherical cells near point of attach-
ment. Fertile frond: fertile part of lamina to 18 by
2'/, cm, gradually narrowed to base which con-
tinues as a sterile wing 10 cm long almost to phyl-
lopodium, apex abruptly narrowed and rounded.
Distr. S. Vietnam; in Malesia: Borneo (Sarawak,
Sabah), Moluccas (Ceram) (?), W. New Guinea.
Ecol. Epiphyte at 1800-2100 m (1000-1800 m
in Vietnam).
Note. Some specimens from S. Vietnam have
sterile fronds to 40 cm long and fertile to 20 by
4 cm; I have seen none so large in Malesia, and the
above description is mainly from Malesian speci-
mens.
18. Elaphoglossum melanostictum (BL.) Moore,
Ind. Fil. (1862) 361; v.A.v.R. Handb. (1908)
716; Suppl. (1917) 427; BACKER & PosTH. Varenfl.
Java (1939) 249. — Acrostichum melanostictum
BL. Fl. Jav. Fil. (1829) 26, t. 7. — Type: vAN
HASSELT, Java (L).
Acrostichum lessonii METT. in Kuhn, Linnaea 36
(1969) 60, excl. pl. LEsson. — Lectotype: KORTHALS
s.n., Sumatra (B).
Acrostichum beccarianum BAKER, Malesia 3
(1886) 27. — E. beccarianum C.Cur. Ind. Fil.
(1905) 303; v.A.v.R. Handb. (1908) 714. — Type:
BECCARI s.n. Nov. 1865, Kuching, Sarawak
(FI; dupl. in K).
E. basilanicum Cope. Philip. J. Sc. 11 (1916)
Bot. 41; v.A.v.R. Handb. Suppl. (1917) 425;
CopeL. Fern Fl. Philip. (1960) 280. — Type:
REILLO BS 16232, Basilan (MICH; dupl. in P).
E. peninsulare Ho_trum, Gard. Bull. S. S. 11
(1947) 270; Rev. Fl. Mal. 2 (1954) 456, f. 266. —
Type: ErYL SMITH 2021, Patani, Thailand (SING;
dupl. in K).
Acrostichum apodum (non KAULF.) FEE, Hist.
Acrost. (1845) 42, pl. jav. tantum.
Acrostichum norrisii (non Hook.) CESATI, Atti
Acad. Napoli 7, pt 8 (1876) 31. — Fig. 18d, e.
Rhizome short; scales dark brown, to 10 mm
long (often shorter), 1 mm wide, acuminate, edges
with short stiff hairs; phyllopodia 1-2 cm long.
Sterile frond: stipe 0-3 cm; lamina thinly coria-
ceous, to 40 by 6-7 cm, widest '/,; from apex,
narrowed gradually to long-decurrent base, apex
broadly but distinctly pointed; thinner edge nar-
row, not decoloured; veins usually distinct; costa
rounded and prominent beneath; scales on lower
surface light brown, irregularly stellate with slender
arms to 1-1'/, mm long, lacking conspicuous
spherical cells near point of attachment. Fertile
frond: stipe 7-15 cm; lamina 11-30 cm long,
1.3-3.7 cm wide, shape as sterile.
Distr. Peninsular Thailand and Malesia:
Sumatra, Malay Peninsula, Java, Borneo, Philip-
pines (Basilan, Samar, Negros, Mindanao).
Ecol. Epiphytic on tree-trunks in forest at
100-1700 m.
GROUP (Holttum)
301
Notes. E. basilanicum was described from rather
small sterile plants, but in shape of frond and
scales agrees closely with other specimens. The
species Acrostichum lessonii was based in part on
specimens collected by LkEssoN on Vanicoro
(Santa Cruz I.) and partly on a specimen from
Sumatra, coll. KorTHALS. The latter alone has a
fertile frond, and its sterile ones are much larger
than those from Vanicoro. The description agrees
with the Sumatran plant, which should thus be the
type; the Vanicoro plant is different and I cannot
identify it.
A collection of GRABOwsKY (s.n. Jan. 1882; B,
BM) from SE. Borneo, is rather intermediate both
in scales and frond-form between E. melanostictum
and E. norrisii; sterile fronds to 20 by 3 cm, fertile
with long stipe as melanostictum but only 1 cm
wide and very long-decurrent. I regard these
specimens as small F. melanostictum.
19. Elaphoglossum heterostipes HoLTruM, Blumea
14 (1966) 321. — Type: ENDERT 4424, Borneo,
W. Kutai, Mt Kemul (BO).
Rhizome short; scales medium brown, thin, 5—7
by 11/,-2 mm, subentire, apex narrowed but not
acuminate; phyllopodia 1 cm long. Sterile frond:
unwinged stipe 0-5 cm; lamina thinly coriaceous,
to 38 by 4.8 cm, widest above middle, base nar-
rowly long-decurrent, apex narrowed to acute
(not acuminate) tip; thin edge distinct but not
decoloured; costa beneath distinctly prominent;
veins slightly prominent on both surfaces, their
swollen ends distinct; scales on lower surface
numerous, very small, dark, lacking long hairs.
Fertile frond: stipe 25-30 cm; lamina 25 by 2 cm,
shape as sterile.
Distr. Malesia: E. Borneo. Only known from
type collection.
Ecol. In forest at 1800 m.
Note. This species has sterile fronds closely
similar to those of E. me/anostictum in shape, but
differs in scales on rhizome and fronds and in the
very long stipe of fertile frond. ENDERT collected
E. melanostictum on the same mountain at 1600 m.
20. Elaphoglossum calanasanicum HoLttuM, Kali-
kasan, Philip. J. Biol. 3 (1974) 196. — Type:
M. G. Price 2933, Luzon, Kalinga-Apayao
Prov., Calanasan (K, PNH).
Rhizome short; scales to 15 by 1 mm, brown,
glossy, crisped, with few marginal hairs; phyllo-
podia 1 cm long. Sterile frond: stipe to 2 cm; lamina
to 34 by 2.7 cm, widest about middle, gradually
narrowed to base and more abruptly to acute apex;
costa almost flat both sides; veins faintly visible,
not prominent; cartilaginous edge hardly '/, mm
wide. Fertile frond: unwinged stipe to 2 cm long,
then a narrowly winged portion 7-8 cm long,
gradually widening upwards, lamina 12-15 by
2 cm, widest at middle, narrowed about equally to
acute apex and to base, scales on upper surface to
1/, mm long, stellate, end cells of arms ellipsoid.
Distr. Malesia: Philippines (Luzon), only
known from the type.
Ecol. At 1400 m, abundant, growing with
E. ophioglossoides.
21. Elaphoglossum norrisii (HooK.) BEDD. Ferns
Br. India pt 23 (1870) addendum; Handb. Ferns
302
Br. India (1883) 418; Suppl. (1892) 104; v.A.v.R.
Handb. (1908) 716. — Acrostichum norrisii HooK.
Spec. Fil. 5 (1864) 215. — E. melanostictum [non
(BL.) Moore] Hottrum, Rev. Fl. Mal. 2 (1954)
455, f. 265. — Type: Norris, Penang (K). —
Fig. 18a-c.
Rhizome short; scales to 10 by 11/, mm, rather
dark red-brown, thin, tapering to a fine point,
edges with irregular teeth and hairs; phyllopodia
short. Sterile frond: no stipe; lamina 20-40 by
2-4 cm, thinly coriaceous, usually widest above
middle, very gradually narrowed to base, more
shortly to rather narrow but rounded apex; thin
pale edge very narrow; veins faintly visible; costa
beneath broad and little prominent; scales on lower
surface stellate with short arms and conspicuous
spherical cells near point of attachment. Fertile
frond: stipe commonly to 2 cm, sometimes longer;
lamina commonly to 20 by 1 cm, widest seen
pS oan.
Distr. Malesia: Sumatra, Malay Peninsula,
Borneo, W. Java, W. New Guinea.
Ecol. In forest, on mossy tree-trunks and rocks,
at 60-1500 m.
22. Elaphoglossum apoense HoL_trum, Blumea 14
(1966) 320. — Type: EDANo PNH 710, Mt Apo,
Mindanao (MICH). — Fig. 20d.
Rhizome short; scales dark, rigid, glossy, some-
what contorted, 5 by ?/, mm, hair-pointed with
some lateral stiff hairs; phyllopodia 1 cm long.
Sterile frond: stipe 22 cm long, pale, bearing scales
of various sizes from very small to 5 mm long, the
latter as those on rhizome but narrower; lamina
thinly coriaceous, 191/, by 3 cm, widest 1/; from
apex, tapered gradually towards base which is not
decurrent, towards apex narrowed a little and then
rounded; thin pale edge very narrow; veins dis-
tinct; costa prominent and rounded beneath; scales
on both surfaces abundant, dark brown, glossy,
very narrow, mostly 1-2 mm long, with spreading
rigid marginal hairs each consisting of one dark
cylindrical cell with a small pale cell at its apex (not
seen at < 10 magnification); many scales 1 mm
long attached close to edge of lamina and spread-
ing beyond it. Fertile frond not seen.
Distr. Malesia: Philippines (Mindanao), Papua
New Guinea (Mt Albert Edward, T. NAIKAKE 552).
Ecol. In mossy forest, at 2100 m.
23. Elaphoglossum vepriferum HoL_trum, Blumea
14 (1966) 326. — Type: CLEMENS 7417, NE. New
Guinea, Morobe District, Sambanga (B; dupl. in
GH sterile).
Rhizome creeping, bearing fronds to 8 mm apart
in each rank; scales as in E. apoense; phyllopodia
10-15 mm long. Sterile frond: stipe 15-25 cm,
rather persistently scaly with narrow dark glossy
scales to 1 mm long with rigid short marginal hairs;
lamina to at least 20 by 2.2 cm, thinly coriaceous,
widest above the middle, apex narrowed slightly
and rounded, base rather narrowly cuneate, not
long-decurrent; pale edge firm, distinct; veins just
visible; costa slightly prominent beneath; scales on
lower surface of costa dark, very narrow, */,-1 mm
long, with stiff marginal hairs as in E. apoense;
smaller paler scales with similar rigid hairs at
first abundant on both surfaces, those near edge
1/, mm long and forming a short tangled fringe
FLORA MALESIANA
[ser. TI; voller?
spreading beyond edge of lamina. Fertile frond:
stipe 17 cm; lamina 14 by 1.4 cm, shape as sterile.
Distr. Malesia: E. New Guinea. Only known
from type collection.
Ecol. In forest, at 1500-1800 m.
24. Elaphoglossum nigripes HoLTTUM, Blumea 14
(1966) 323. — Type: PULLE 493, W. New Guinea,
Mt Parameles (BM; dupl. in BO, L, UV).
Rhizome creeping, rather slender, fronds close;
scales dark, glossy, apparently 3 by 1 mm, not
slender-tipped; phyllopodia 5 mm long. Sterile
frond: stipe 5 cm, covered when young with
appressed small dark entire scales; lamina thinly
coriaceous, 25-40 by 1.8-2!/, cm, widest above
middle, very gradually narrowed to decurrent base,
also gradually to narrowly rounded apex; thin
edge very narrow, reflexed when dry; veins +
distinct; costa slender and prominent beneath;
scales on costa beneath small, appressed, ovate-
acute, dark, glossy; scales on lamina few, small,
dark, appressed, with a few spreading hairs 2-3
cells long. Fertile frond: stipe 10 cm; lamina to 25
by 1.3 cm, shape as sterile, with rather abruptly
narrowed blunt apex.
Distr. Malesia: W. New Guinea. Only known
from two collections.
Ecol. In forest, at 1100-2500 m.
25. Elaphoglossum melanochlamys HOLTTUM,
Blumea 14 (1966) 322. — Type: EymMa 5408, W.
New Guinea, Wissel Lakes (BO; dupl. in L).
Rhizome short; scales mostly black, rigid, glossy,
5 mm long, less than 1 mm wide, acuminate, edges
short-toothed or with some stiff hairs; phyllo-
podia to 1 cm long. Sterile frond: stipe 3-5 cm long,
pale, rather persistently scaly, scales as rhizome,
narrow, spreading, with more conspicuous mar-
ginal hairs; lamina rigid, drying rather pale, 15-21
by 1.6 cm, sides parallel for most of length, nar-
rowed rather abruptly to non-decurrent base and
to narrowly rounded apex; pale edge distinct,
1/, mm wide; veins obscure; costa beneath broad,
pale, slightly prominent; scales on lower surface
abundant, as those on stipe but smaller, mostly
1-3 mm long. Fertile frond: stipe 5 cm; lamina 15
by 1.4 cm; abundant scales among sporangia.
Distr. Malesia: W. New Guinea. Only known
from type collection.
Ecol. Probably at over 2000 m.
26. Elaphoglossum nesioticum Ho_truM, Blumea
14 (1966) 323. — Type: Brass 24882, Goodenough
I. (A).
Rhizome short; scales dark, glossy, rigid, c. 7 mm
long, hardly 1 mm wide, somewhat crisped, mar-
ginal hairs few; phyllopodia 1 cm long. Sterile
frond: stipe 6-8 cm; lamina thin, 20-37 by 1-3.8 cm,
widest about middle, narrowed gradually to decur-
rent base and to narrow slightly acuminate apex;
thin edge very narrow, deflexed; veins slightly
prominent, apices sometimes anastomosing; costa
beneath slightly prominent; surfaces glabrescent.
Fertile frond: stipe 8 cm; lamina 19 by 1'/, cm,
shape as sterile.
Distr. Malesia: E. New Guinea (Goodenough
I.) (4 collections).
Ecol. Epiphyte in mossy forest, at 1500-1600 m.
1978]
LOMARIOPSIS GROUP (Holttum)
303
27. Elaphoglossum angustifrons HoLtTrumM, Blumea
14 (1966) 319. — Type: WomersLEY NGF 11272,
NE. New Guinea, Minj (LAE; dupl. in A, K).
Rhizome short; scales dark, glossy, flat, to 6 by
2 mm, subentire, not long-acuminate; phyllopodia
2-3 cm long. Sterile frond: stipe 5-10 cm, rather
pale, at first with spreading scales as rhizome,
persistent scales very small, dark brown, entire or
with a few hairs; lamina thinly coriaceous, drying
dark, 25-35 by 1*/,-2.3 cm, sides parallel for most
of their length, base very narrowly cuneate and +
decurrent, apex short-acuminate; thin edge very
narrow, not pale; veins distinct, mostly with free
thickened tips, occasionally joining near margin;
costa pale and prominent beneath; scales persistent
on lower surface, very small, dark, with a few short
hairs which seem often abraded. Fertile frond: stipe
10-15 cm; lamina 16-20 by 1.2-11/, cm.
Distr. Malesia: E. New Guinea. Only known
from type collection.
Ecol. At 2100-2400 m.
Note. This has the aspect of E. heterolepium,
but very different scales, and appears to be nearly
related to E. archboldii.
28. Elaphoglossum pallescens HoLTTuM, Blumea 14
(1966) 323. — Type: Brass 22879, Papua, Mt
Dayman (A).
Rhizome short; scales dark brown, glossy, c. 10
by 11/, mm, rigid, flat, hair-pointed, with short
marginal teeth and slender marginal hairs; phyl-
lopodia 1 cm long. Sterile frond: stipe 9-12 cm,
pale; lamina rigid, drying pale, to at least 21 cm
long, 2.8-4 cm wide, edges parallel in middle part,
tapering about equally to slightly acuminate apex
and slightly decurrent base; decoloured edge very
narrow, reflexed; veins obscure; costa broad and
slightly prominent beneath; scales on lower sur-
face dark, appressed, + stellate with arms of
several cells, in all to !/, mm wide; upper surface
with scattered small pits in which are shrivelled
scales. Fertile frond: stipe 15 cm; lamina 15 by
21/, cm, base rather abruptly narrowed and slightly
decurrent.
Distr. Malesia: E. New Guinea (2 collections).
Ecol. Low on mossy tree in forest, 1800-2230 m.
29. Elaphoglossum arachnoideum HoLttTuM, Blumea
: "GA = 320. — Type: Brass 24541, Goodenough
Rhizome short; scales 10 by 11/, mm, acuminate,
rather dull medium brown, stiff, subentire, some-
what twisted; phyllopodia c. 1 cm long. Sterile
frond: stipe 20 cm, rather persistently scaly as
rhizome but scales thinner; lamina firm but rather
thin, 21 by 5.8 cm, widest in middle, almost ellipti-
cal, base slightly decurrent, apex rounded but not
broadly; thin edge not sharply distinct; veins just
distinct; costa beneath slightly prominent only
near base; lower surface rather persistently scaly,
many scales 2-3 mm long, very narrow, pale brown
with a few marginal hairs, also abundant much
smaller scales; scales near margin a little wider and
projecting 2-3 mm, mostly abraded. Fertile frond:
stipe 22 cm; lamina 12 by 3 cm, shape as sterile.
Distr. Malesia: E. New Guinea (Goodenough
I.). Only known from type collection.
Ecol. High on trees in oak forest, at 1600 m.
30. Elaphoglossum stenolepis BELL ex HoLttum,
Blumea 14 (1966) 325. — E. decurrens (non Desv.)
C.Cure. Gard. Bull. S. S. 7 (1934) 290, p.p. — Type:
CLEMENS 28019, Mt Kinabalu (US).
Rhizome thick, short; scales c. 10 mm long, less
than 1 mm wide, crisped, red-brown, entire, nar-
rowed to a slender tip; phyllopodia c. 1 cm long.
Sterile frond: stipe 7 cm long, the upper 2 cm
narrowly winged; lamina 17-20 by 31/,-4 cm, rigid,
rather thick, drying brown and wrinkled, widest
above middle, narrowed gradually to wing at base
and to broadly pointed or slightly rounded apex;
thin pale edge very narrow; veins obscure; costa
beneath broad, pale, little prominent; scales very
small, stellate with 3-6 arms of several cells with
small glandular cells laterally on arms. Fertile
frond: stipe 11 cm; lamina 11 by 2.2 cm, rather
abruptly narrowed at base and at rounded apex.
Distr. Malesia: N. Borneo (Mt Kinabalu),
several collections.
Ecol. Epiphyte at 1500-2150 m.
Note. This species grows with E. annamense,
and CLEMENS 27060 is a mixture of the two species.
The scales of E. stenolepis are very distinctive, also
the longer stipe.
31. Elaphoglossum amblyphyllum BELL, nom. nov.
— E. obtusifolium BELL, Kew Bull. 14 (1960) 83,
non BRACK. — Acrostichum decurrens (non DESV.)
BL. En. Pl. Jav. (1828) 102; Fl. Jav. Fil. (1829) 32,
t. 10; RactB. Fl. Btzg 1 (1898) 47. — E. decurrens
(non Desv.) v.A.v.R. Handb. (1908) 713, p.p.;
Suppl. (1917) 424, p.p.; BACKER & PosTH. Varenfi.
Java (1939) 249; HoLttum, Rev. FI. Mal. 2 (1954)
458, f. 268. — Type: BLUME, Java (L).
Acrostichum obtusifolium (non WILLD.) BL. En.
Pl. Jav. (1828) 102; Fl. Jav. Fil. (1829) 31; Hook.
Spec. Fil. 5 (1864) 204, p.p. — Olfersia blumeana
PRESL, Tent. Pterid. (1836) 235. — Type: BLUME,
Java (L). — Fig. 19.
Rhizome short; scales pale brown, 10-12 mm
long, to 2 mm wide at base, acuminate, edges -+
ciliate; phyllopodia 1 cm long. Sterile frond: stipe
7-15 cm long; lamina coriaceous, to 29 by 8!/, cm,
widest above the middle, apex broadly rounded,
base cuneate and shortly decurrent; pale edge
distinct, narrow; veins obscure; costa beneath
broad and little prominent; scales small, appressed,
stellate, with some small spherical cells near point
of attachment. Fertile frond: stipe 20 cm; lamina to
20 by 4 cm.
Distr. Malesia: Sumatra, Malay Peninsula, Java,
Borneo, Moluccas (Ambon).
Ecol. Epiphyte on old mangrove trees and by
rivers, at 0-800 m.
Notes. The type is the specimen illustrated by
BLUME as Acrostichum decurrens. As shown by
BELL (/.c. 1960) it is certainly different from the type
of A. decurrens DEsv. The latter is a poor specimen,
and I have been unable to identify any Malesian
species with it. The specimen named Acrostichum
obtusifolium WILLD. by BLUME is certainly not
conspecific with WILLDENOW’s type, which is
probably a young plant of Polypodium scolopendria
Burm. f. (photograph seen, showing venation).
The name Elaphoglossum obtusifolium was pub-
lished by BRACKENRIDGE with a description of a
Fiji specimen, and citation of Acrostichum obtusi-
folium BL. (non WILLD.) asasynonym. RACIBORSKI’S
304
FLORA MALESIANA
[ser. Il, volea?
description of Acrostichum decurrens was based on
a specimen from Ambon (leg. TEYSMANN) which I
have seen.
Fig. 19. Elaphoglossum amblyphyllum BELL. a.
Habit, 1 fertile and 2 sterile fronds, x 1/2, b-c.
scales from surface of frond, x 75 (L 908.332-621,
908.352-637).
32. Elaphoglossum luzonicum CopeEL. in Elmer,
Leafl. Philip. Bot. 1 (1907) 235; ibid. 2 (1908) 416;
v.A.v.R. Handb. (1908) 714; CopeL. Fern FI.
Philip. (1960) 278. — Type: ELMER 9036 (MICH,
teste CopEL. MS; dupl. in BO, K, L, P, US).
Acrostichum decurrens var. ornatum FEE, Hist.
Acrost. (1845) 34. — Acrostichum decurrens (non
Desv.) Hook. Spec. Fil. 5 (1864) 203. — Type:
CUMING 144, Luzon (BM, K, L, MICH, P, US).
Acrostichum cumingii (non FEE) BAKER, Syn.
Fil. (1868) 407. — Acrostichum decurrens var. major
MetTT. ex KUHN, Ann. Mus. Bot. Lugd.-Bat. 4
(1869) 292. — E. decurrens var. cumingii V.A.V.R.
Handb. (1908) 713. — Type: Cumine 193, Luzon
(BM, K, P).
E. elmeri Copev. in Elmer, Leafl. Philip. Bot. 3
(1910) 849; v.A.v.R. Handb. Suppl. (1917) 424;
CopeL. Fern Fl. Philip. (1960) 279. — Type:
ELMER 11658, Mindanao (not seen).
E. lepidopodum C.Cur. ex OGATA, J. Jap. Bot.
13 (1937) 121. — Type: M. OcaTa 60, Taiwan,
Prov. Taito (BM).
E. cumingii var. papuanum C.Cur. Brittonia 2
(1937) 317. — Type: Brass 4112, Papua (NY). —
Fig. 20a-c.
Rhizome short; scales thin, light brown, 5-7 by
2-3 mm, edges with short hairs; phyllopodia to
2 cm long. Sterile frond: on young plants widest
above middle with -- broadly rounded apex and
gradually narrowed base, with a rather persistent
fringe of spreading scales 11/,-2 mm long all along
edge of lamina; lamina to c. 20 by 4cm; on mature
plants with stipe 10-15 cm, persistently scaly as
rhizome, lamina rigidly coriaceous, to 25 by
3.2-4!/, cm, widest at or a little above middle,
sides parallel for most of their length, narrowed
rather abruptly at both ends, base a little decur-
rent, apex bluntly pointed or rounded; pale thin
edge distinct; veins obscure; costa slightly raised
on lower surface; small scales on lamina light
brown, to 1 mm long, narrow with hairs near base,
near edge of lamina on both surfaces broader
scales 1 mm long + persistent. Fertile frond: stipe
17 cm; lamina to 26 by 31/, cm.
Distr. Taiwan; in Malesia: Philippines (Luzon,
Negros, Panay, Mindanao), E. New Guinea.
Ecol. In mossy forest, at 2000-2300 m (few
altitudes recorded).
Notes. The type of E. /uzonicum cited with the
original description was ELMER 8190; but CopE-
LAND wrote ‘“‘TyPeE’’ on the specimen of no 9036 in
his own herbarium (MICH) without explaining the
discrepancy. I have not seen any specimen bearing
the number ELMER 8190.
The treatment of this species (and, with it, that
of E. amblyphyllum) has been very confused, both
taxonomically and nomenclaturally. In his enu-
meration of CUMING’s Philippine ferns, JOHN SMITH
cited CumiNG 144 and 193 under the name E. obtu-
sifolium, based on Acrostichum obtusifolium
WILLD. citing A. decurrens BL. as synonym. But as
the generic name Elaphoglossum had not then been
validly published, the binomial E. obtusifolium
J.SM. was illegitimate (see p. 314). Further, the
type of Acrostichum obtusifolium WILLD. is a Poly-
podium-ally; and the specimen described by BLUME
as A. decurrens does correspond with DEsvAUx’s
type of that species; both epithets will also be found
confusedly in the synonymy of E. amblyphyllum.
1978] LOMARIOPSIS GROUP (Holttum) 305
> 9S Ee ir a eee
y
Vv
oe.
AwesS
SS; ==
<
Fig. 20. Elaphoglossum luzonicum Cope. a. Habit, 1/,, b. scales on margin of frond, 10, c. scale from
margin, x 25. — E. apoense Ho.trum. d. Scale from surface of frond, x 75 (a—b BS 35595 & a part of
ELMER 17489, d EDANo PNH 710).
306
W
li
FLORA MALESIANA
===
(oh
Pad
i}
I!
Ib, 4m
[ser. II, vol. 14
oO
revel Fl
Fig. 21. Elaphoglossum callifolium (BL.) Moore. a. Habit, 1 sterile and 1 fertile frond, x 1/3, b. margin of
sterile frond, x 3,c.scale from surface of frond, x 50,d-e. scales from base of stipe, x 2 (a-b,d L 908.331-
933).
1978]
LOMARIOPSIS GROUP (Holttum)
307
Fée described his specimen of CUMING 144 as
Acrostichum decurrens var. ornatum. He cited
CUMING 193 as type of A. cumingii, but, as noted
by COPELAND, his description does not agree with
specimens of that collection in other herbaria, and
his name is therefore here regarded as doubtful
(see p. 314). HooKeR included A. cumingii FEE as
one of many synonyms of A. conforme Sw. (Spec.
Fil. 5: 199) citing CUMING 193; but BAKER, in Syn.
Fil., recognized A. cumingii as a distinct species,
basing his description on the specimen of CUMING
193 in HooKErR’s herbarium.
In 1960 CopELAND doubted whether his E. e/meri
and £. luzonicum were distinct species. I have
examined many specimens of the type collection of
E. luzonicum and of CumtNnG 144 and 193. I have
come to the conclusion that the type of E. luzonicum
and CuMING 193 are conspecific. CUMING 144
consists entirely of sterile plants, and I regard these
as + immature stages of the same species as 193;
some other collections show intermediate con-
ditions. As in some other species of this genus,
young plants have fronds with more broadly
rounded apices and more decurrent bases than
fronds of mature plants. Fronds of young plants
also have a more persistent fringe of scales which
are larger than on mature plants; old fronds of
some mature plants lack such a fringe, but the scars
of attachment of the scales can be seen.
33. Elaphoglossum brevifolium Hoitrum, Gard.
Bull. Sing. 11 (1947) 270; Rev. Fl. Mal. 2 (1954)
458, f. 267. — Type: Ho_trum 20750, G. Tahan,
Malaya (SING; dupl. in K).
Rhizome short; scales rather light brown, thin,
c. 10 by 11/, mm, tapered to a slender apex, entire;
phyllopodia 1 cm long. Sterile frond: stipe 4-12 cm
long, pale, about half of it with a very narrow
wing, bearing some residual scales as rhizome;
lamina thick and rigid when dry, 8-12 by 3'/,-6 cm,
almost elliptical but base decurrent, apex broadly
bluntly pointed; thin pale edge '/, mm wide, some-
times deflexed on drying; veins obscure; costa be-
neath slightly prominent in basal half; scales on
surfaces at first abundant, very small, with a few
marginal hairs of 2-3 cells. Fertile frond: stipe 20-
25 cm; lamina 8-10 by 2-3 cm, shape as sterile.
Distr. Malesia: Malay Peninsula (G. Tahan and
G. Batu Puteh).
Ecol. At 1400-1850 m; on G. Tahan a low
epiphyte in mossy forest.
34. Elaphoglossum negrosensis HoLttuM, Blumea
14 (1966) 323. — E. conforme [non (SW.) J.SM.]
CopPEL. in Elmer, Leafl. Philip. Bot. 2 (1908) 417. —
E. angulatum [non (BL.) Moore] Copev. Fern FI.
Philip. (1960) 278, p.p. — Type: ELMER 9885,
Philippines, Negros (MICH; BO).
Rhizome short; scales rather light brown, to 7
by 2!/, mm, acuminate but not hair-pointed, with
few marginal hairs; phyllopodia 1 cm long. Sterile
frond: stipe 2-8 cm, scaly when young as rhizome;
lamina thinly coriaceous, 11-17 by 2.7-3.3 cm,
widest at or below middle, apex evenly narrowed
to a very narrowly rounded tip, hardly acuminate,
base rather abruptly narrowed and then decurrent
as a wing 2 cm long on the stipe; thin pale edge
distinct, less than 1/, mm wide; veins slender,
distinct on both surfaces, rather widely spaced,
their apices thickened and not joining; costa hardly
prominent on lower surface; no scales seen on
lamina. Fertile frond: stipe 10-12 cm; lamina 8-11
by 1.3—11/, cm, shape as sterile.
Distr. Malesia: Philippines (Negros Oriental).
Only known from type collection.
Ecol. In forest, at 1400 m.
Note. The type was originally named E. con-
forme by COPELAND, and later re-named E. angula-
tum, from which it differs in rhizome and vein
characters.
35. Elaphoglossum planicosta Ho_trum, Blumea
14 (1966) 324. — Type: JERMy 4216, NE. New
Guinea, Madang Distr., Finisterre Mts (BM).
Rhizome short; scales 10 by 2-2!/, mm, nar-
rowed evenly to apex, light brown, thin, with
slender marginal hairs; phyllopodia to 1 cm long.
Sterile frond: stipe pale or slightly rufous, 6-9
(-13) cm long, upper 2-3 cm with narrow wing
decurrent from lamina; lamina thick and rigid
when dry, 11-15 by 2.7-3.4 cm, widest about
middle, rather abruptly narrowed to a + obtuse-
angled apex and more gradually to a narrowly
cuneate base; thin pale edge more than '/, mm wide;
veins obscure except on young fronds, their
apices thickened, sometimes forked, the branches
then sometimes meeting those of adjacent veins;
costa broad, pale, almost flat on lower surface;
scales on both surfaces of young fronds appressed,
stellate, mostly c. !/, mm @ including arms. Fertile
frond: stipe 15 cm; lamina 11 by 3 cm, shape as
sterile.
Distr. Malesia: NE. New Guinea (two collec-
tions).
Ecol. In degenerate cloud-forest, epiphytic, at
2750 m.
Note. A fixation from the type collection gave a
chromosome count n = ec. 82 (tetraploid).
36. Elaphoglossum callifolium (BL.) Moore, Ind.
Fil. (1857) 7; Curist, Monogr. Elaph. (1890) 34;
v.A.v.R. Handb. (1908) 714; Suppl. (1917) 425;
C.Cur. Gard. Bull. S. S. 7 (1934) 289; BACKER &
Postu. Varenfi. Java (1939) 251; Ho_trum, Rev.
Fl. Mal. 2 (1954) 459, p.p.; CoPEL. Fern FI. Philip.
(1960) 277. — Acrostichum callifolium BL. En. Pl.
Jav. (1828) 100; Fl. Jav. Fil. (1829) 22, t. 4; FEE,
Hist. Acrost. (1845) 28; Racts. Fl. Btzg 1 (1898)
47. — Olfersia callifolia Pres, Tent. Pterid. (1836)
234. — Type: BLumE, Java (L).
Acrostichum junghuhnianum KunNzZE, Bot. Zeit. 6
(1848) 101. — E. junghuhnianum Moore, Ind. Fil.
(1857) 10.— Type: JUNGHUHN, Mt Lawu (not seen).
E. reineckei HIERON. & LAUTERB. Bot. Jahrb. 41
(1908) 220. — Type: VAUPEL 452, Samoa (B).
E. permutatum var. mutatum v.A.V.R. Bull. Jard.
Bot. Btzg II, 16 (1914) 13; Handb. Suppl. (1917)
426. — Type: RACHMAT 604, Mt Boesoe, Celebes
(BO).
E. macgregorii Cope. Philip. J. Sc. 11 (1916)
Bot. 40; v.A.v.R. Handb. Suppl. (1917) 424. —
Type: R. C. McGrecor BS 19780, Luzon (MICH).
E. laurifolium [non (THouaRS] Moore)
v.A.v.R. Handb. (1908) 714, p.p.; BACKER &
PostH. Varenfl. Java (1939) 251, p.p.
E. commutatum [non (METTt.) v.A.V.R.] v.A.V.R.
Handb. Suppl. (1917) 427, p.p. — Fig. 21, 26f.
308
FLORA MALESIANA
[ser. II, vol. 14
Rhizome massive, fronds close; scales c. 10 by
1 mm, rather stiff and straight, often with inflexed
edges but not crisped even near apex, entire or
nearly so, dull brown; phyllopodia 2-3 cm long.
Sterile frond: stipe 10-18 cm, rather thick, grooved
deeply on upper surface only; lamina coriaceous,
commonly to 35 by 5-6 cm, to 48 by 8 cm or more
(80 by 10 cm reported in Java), acuminate, base
rather narrowly cuneate and a little decurrent; thin
edge pale, narrow, usually reflexed when dry; veins
distinct but little prominent; costa rounded and
prominent beneath; scales on lamina scattered,
very small, stellate, brown. Fertile frond: stipe
usually longer than sterile, to 20 cm or more;
lamina commonly to 25 by 4 cm, base rather
abruptly narrowed, apex short-acuminate.
Distr. S. Vietnam, throughout Malesia; east-
wards to Samoa.
Ecol. In mountain forest, sometimes in a dense
growth of other epiphytes, at 1000-2400 m.
Notes. There are a few quite small, but fertile,
specimens from Java, Sumatra and Borneo which
agree so closely with typical E. callifolium that I
place them here. At least some such have been
called E. laurifolium (the type of which, from
Tristan d’Acunha, is very different in scales) and
E. commutatum, which see for further comment.
But I do not think that the specimen originally
named E. conforme by BLUME belongs here (see
E. recommutatum). The Malay Peninsula specimens
here described as E. malayense have also been
called E. callifolium, but are very distinct in their
scales; I have seen only one Peninsula specimen
which seems to be true E. callifolium, with a frond
50 by 9 cm.
37. Elaphoglossum commutatum (METT. ex KUHN)
v.A.v.R. Handb. Suppl. (1917) 427; SLEDGE, Bull.
Brit. Mus. (Nat. Hist.) 4 (1967) 90. — Acrostichum
commutatum MEeEtTT. ex KUHN, Ann. Mus. Bot.
Lugd.-Bat. 4 (1869) 292, excl. spec. Blum. —
Acrostichum laurifolium (non THouaRs) THW. En.
Pl. Zeyl. (1864) 380. — Type: THwaites 1310 (B;
dupl. in K).
E. permutatum v.A.V.R. Bull. Jard. Bot. Btzg II.
16 (1914) 13; Handb. Suppl. (1917) 425, excl. var.
mutatum. — Type: MATTHEW 696, Mt Sago,
Sumatra (BO). — Fig. 16a.
Rhizome usually short with close fronds; scales
to 10 by 1!/, mm, medium brown, rather thin, +
crisped throughout but especially towards hair-
pointed apices which are often tangled, edges with
some spreading hairs; phyllopodia 1—1'/, cm long.
Sterile frond: stipe rather slender, 5-15 cm long,
rather deeply 3-sulcate almost to base; lamina
thinly coriaceous, commonly to 25 by 6 cm,
exceptionally to 30 by 9 cm, distinctly elliptic,
rather shortly acuminate, cuneate and slightly
decurrent at base; thin edge very narrow, often
defiexed ; veins slender, prominent on both surfaces,
not joining at tips; costa prominent on both
surfaces, rather deeply grooved on upper; scales
brown, stellate, mostly under 1/, mm in total @.
Fertile frond: stipe sometimes longer than sterile;
lamina 12-17 by 1.8-2!/, cm.
Distr. Ceylon & S. India; in Malesia: Sumatra,
W. Java, Borneo, New Guinea.
Ecol. In forest, in two cases recorded as terres-
trial (MATTHEW reported of the type of E. permu-
tatum “‘in a patch of moist ground in forest’’), at
900-1700 m.
Notes. THwaites 1310 from Ceylon is taken as
type, as it is apparent that KUHN’s description was
made chiefly from it. The BLUME specimen cited is
not a good one and the scales are mostly broken;
in my opinion it represents a distinct species
(E. recommutatum).
Below the decurrent base of the lamina of a
frond, a distinct groove develops on each side of
the median groove; these lateral grooves are on the
lines of the decurrent edges of the lamina (v.A.v.R.
ears the stipe of E. permutatum as plurisul-
cate).
I have wondered whether E. commutatum could
be an ecologic form of E. callifolium, growing in
unusually moist shady conditions. But the Malesian
specimens here described agree so closely (except
that fronds are somewhat larger) with those from
Ceylon and S. India, where typical E. callifolium is
lacking, that I think it right to maintain E. com-
mutatum as a distinct species. MATTHEW, who
knew E. callifolium well in the field, wrote a MS
note on his Sumatran specimen of E. permutatum
‘a most distinct species, fully 500 feet below the
level of E. callifolium’’.
38. Elaphoglossum malayense HoLTruM, Blumea
14 (1966) 322. — E. callifolium (non BL.) HOLTTUM,
Rev. Fl. Mal. 2 (1954) 459, p.p. max. — Type:
HENDERSON 17765, Cameron Highlands, Malaya
(K; dupl. in SING).
Rhizome short, thick, scales to 25 by 3 mm,
rather light red-brown, flat, firm, acuminate,
marginal hairs rare; phyllopodia to 11/, cm long.
Sterile frond: stipe on immature plants short, on
mature plants 5-10 cm; lamina rather thinly
coriaceous, to 40 by 5 cm, apex rather shortly
acuminate, narrowed more gradually to base and
then decurrent as a narrow wing 5 cm or more long;
thin edge very narrow, reflexed; veins distinct at
least on upper surface, hardly prominent; costa
prominent on both surfaces; superficial scales not
persistent, very small, stellate. Fertile frond: stipe
8-15 cm; lamina to 24 by 2.2 cm, base decurrent
or not.
Distr. Malesia: Malay Peninsula, Borneo.
Ecol. Epiphyte, at 1050-1400 m.
Notes. This species has been called E. calli-
folium but differs strikingly from the typical
E. callifolium of Java in its much larger flat rhizome-
scales and proportionately narrower, more rigid
fronds. Fronds on smallish plants mostly have
rather short stipes and much-decurrent lamina,
and such plants may bear fertile fronds. In the type
collection one specimen has a fertile frond with
stipe 8 cm and lamina 18 by 2 cm, base narrowly
decurrent, the other has stipe 15 cm, lamina 24 by
2.2 cm, base rather abruptly narrowed. The latter
also has costa deeply grooved on upper surface and
very prominent below, the former has costa shal-
lowly grooved above and not very prominent
beneath.
39. Elaphoglossum recommutatum HOLTTUM, nom.
nov. — Acrostichum conforme (non Sw.) BL. FI.
Jav. Fil. (1829) 23, t. 5. — Acrostichum aemulum
(non KAULF.) BL. En. PI. Jav. (1828) 101. — Type:
BLuMmE, Java (L; dupl. in P).
1978]
Rhizome short; scales c. 6 by 2!/, mm, not long-
acuminate, dull medium brown; _ phyllopodia
1 cm long. Sterile frond: stipe 2 cm (type) to 10 cm
long; lamina of type 15 by 2'/,; cm, of other
specimens to 27 by 4'/, cm, thinly coriaceous, base
cuneate and slightly decurrent, apex shortly
acuminate; pale edge narrow, deflexed; veins just
distinct; costa somewhat prominent beneath;
scales small, stellate, appressed. Fertile frond: stipe
of type 5 cm, of other specimens to 22 cm; lamina
of type 12 by 1.8 cm, of another Java specimen
28 by 2.2 cm, of a Celebes specimen 20 by 3'/, cm.
Distr. Malesia: Sumatra, Java, SW. Celebes.
Notes. KUHN cited BLUME’s specimen under
Acrostichum commutatum Metrt., but he evidently
described the scales of A. commutatum from the
Ceylon specimen of THWAITES, as /onge acuminatis,
adding parenthetically (apice denique delapsis) an
apparent reference to the BLUME specimen, the
scales of which are mostly broken. But I judge from
BLuMe’s drawings of the scales, and from another
Java specimen (LosBB s.n., K) that the scales are
very different from those of the THWAITES speci-
men; see further comment under £. commutatum.
E. recommutatum appears to be the Malesian
species nearest to E. conforme (Sw.) Moore, type
species of the genus. It differs from typical E. con-
forme in the short-acuminate frond-apices and in
lacking the resin-spots which occur abundantly on
the lower surface of that species.
40. Elaphoglossum sordidum Curist, Nova Guinea
8 (1909) 156; v.A.v.R. Handb. Suppl. (1917) 425;
CopeEL. Philip. J. Sc. 78 (1949) 404. — Type:
VERSTEEG 1432, W. New Guinea (BO). — Fig. 22.
Rhizome short; scales medium brown, thin, to 6
by 2 mm, acuminate, with many spreading mar-
ginal hairs; phyllopodia 1 cm long. Sterile fronds:
stipe pale, 10-12 cm long, rather persistently scaly,
scales narrow; lamina thinly coriaceous, 20-30 by
31/,-41/, cm, narrowly elliptical, base gradually
long-decurrent, apex more shortly acuminate; thin
pale edge narrow, deflexed; veins distinct, slightly
prominent; costa prominent, slender and terete on
lower surface; scales on lower surface appressed,
narrow, to | mm long, with long marginal hairs.
Fertile fronds: stipe 12-18 cm; lamina 13-25 by
1-3 cm.
Distr. Malesia: W. New Guinea (4 collections).
Ecol. Epiphyte, at 250-780 m.
41. Elaphoglossum novoguineense ROSENST. in
Fedde, Rep. 10 (1912) 341; v.A.v.R. Handb.
Suppl. (1917) 423; Copev. Philip. J. Sc. 78 (1949)
404. — Type: BAMLER S.67, NE. New Guinea,
Sattelberg (not seen).
E. brassii C.Cur. Brittonia 2 (1937) 316. —
Type: Brass 5558, Mafulu, Papua (NY).
Rhizome short; scales rather thick, dull medium
brown, c. 5-10 by 2!/, mm, subentire or sparsely
hairy, tapering evenly from base to acute apex;
phyllopodia 1—11/, cm long. Sterile frond: stipe
4-11 cm long; lamina thinly coriaceous, 20-33 by
2.8-4 cm, widest about middle, base gradually
narrowed and decurrent for 2-3 cm, apex acute to
acuminate; pale edge narrow, reflexed; veins often
slightly prominent; costa rather broad and pale
beneath, only slightly prominent; scales small,
LOMARIOPSIS GROUP (Holttum)
309
Fig. 22. Elaphoglossum sordidum CurisT.a. Habit, 2
sterile fronds, 1 fertile frond, =< 1/3, b-c. scales
from base of stipe, < 3 (a fertile, b Brass 23000,
a Sterile, c BrAss 11498).
310
FLORA MALESIANA
[ser. Il, vole
stellate. Fertile frond: stipe 10-16 cm; frond 15-20
by 1.2—2 cm.
Distr. Malesia: New Guinea.
Ecol. Epiphyte, at 1200-2400 m.
Notes. I have examined a KEYSSER specimen
(S-PA) from the type locality named and distri-
buted by RosENsTocK. The type of E. brassii is
smaller than above described, with sterile lamina
15 by 3 cm, fertile 7 by 1 cm, but agrees in scaliness.
One specimen collected in NE. New Guinea by
T. G. WALKER (8155) gave a chromosome count
n = c. 82 (tetraploid).
42. Elaphoglossum robinsonii HOLTTUM, sp. nov.
Paleis pallidis tenuibus integris stipitibusque longis
frondium fertilium cum E. brevifolio HOLTTUM
(sp. 33) congruens, ab eo differt frondibus multo
majoribus, fertilibus sterilibus aequilatis, sporisque
multialatis.
Rhizome short; scales to 15 by 2 mm, light
brown, thin, flat, margins entire, tapering to apex
but not hair-pointed; phyllopodia 11/, cm long.
Sterile frond: stipe to 12 cm long, scaly when
young; lamina to 28 by 4!/, cm, widest at middle
and gradually narrowed to both ends, apex broadly
pointed (less than 90°), base narrower and slightly
decurrent; midrib slightly prominent on lower
surface near base, distally almost flat, more
prominent and grooved on upper surface; cartila-
ginous edge less than 1/, mm wide, reflexed when
dry, lacking scales; scales on lower surface of
lamina stellate, to c. 1/, mm diameter, with a few
short rays. Fertile frond: stipe to 24 cm long;
lamina to 25 by 4'/, cm, sides almost parallel for
much of their length, base rather broadly cuneate
and slightly decurrent, apex broadly pointed;
spores bearing numerous small translucent wings.
Type: H. C. Rosinson, Jan. 1913, Gunong
Mengkuang, Selangor, Malaya (K).
Distr. Only known from the type and A. G.
Piccotr 1093, 1094, from the neighbouring moun-
tain, G. Ulu Kali, at 1500-1800 m.
Note. This species differs from E. malayense
HoLttum, the common species on the Main Range
in Malaya, in thinner paler scales, less decurrent
base of sterile frond and longer stipe of fertile fronds.
43. Elaphoglossum favigerum HoLttum, Blumea 14
(1966) 321. — Type: Brass 13440, W. New Guinea,
near Idenburg River (GH; MICH).
Rhizome short; scales dull dark brown, to c. 8
by 13/, mm, rather thin, not acuminate, cells +
hexagonal, isodiametric, edges with a few long
hairs; phyllopodia c. 11/, cm long. Sterile frond:
stipe 10-16 cm; lamina thinly coriaceous, to 39 by
4'/, cm, widest at middle or below it, base cuneate
and slightly decurrent, apex gradually narrowed,
acute; thin edge narrow, reflexed, hardly de-
coloured; veins distinct on upper surface; costa
beneath prominent; scales on lower surface minute,
slightly stellate or not. Fertile frond: stipe 14-16cm;
lamina to 22 by 21/, cm, base rather abruptly
narrowed, more gradually to acute apex.
Distr. Malesia: W. New Guinea (3 collections).
Ecol. Occasional low epiphyte, 1200-1500 m.
44. Elaphoglossum sumatranum HoLttum, Blumea
14 (1966) 325. — Type: MATTHEW s.n., 21 Jan.
1913, Sumatra, G. Tandikat (K).
Rhizome short; scales pale brown, rather thin,
to 10 by 2'/, mm, tapering to apex, subentire;
phyllopodia 1 cm long. Sterile frond: stipe 5—8 cm,
pale, bearing scales as rhizome but smaller; lamina
20-30 by 3'/,-5 cm, thick, rigid, drying green,
widest at or a little above middle, base gradually or
rather abruptly narrowed and slightly decurrent,
distally narrowed rather abruptly to rounded apex;
edge thick and pale; veins obscure; costa pale,
slightly prominent beneath; scales on lower surface
small, stellate. Fertile frond: stipe 15-20 cm;
lamina 14-17 by 1.8-3.3 cm; spores 52-59 by
33-37 um, with broad perispore having very few
folds in it.
Distr. Malesia: Central Sumatra (two collec-
tions).
Ecol. Epiphyte, at 1500 m.
45. Elaphoglossum indrapurae HoL_trum, Blumea
14 (1966) 321. — Type: ALsTon 14275, Central
Sumatra, near G. Kerintji, Sg. Tandok, Kayu Aro
Estate (BM).
Similar to E. sumatranum in rhizome-scales and
spores, but with thinner, much longer fronds.
Sterile frond: stipe 14-18 cm; lamina 35-55 by
3-5 cm, thinly coriaceous, base cuneate and slightly
decurrent, gradually narrowed distally to narrowly
rounded apex; pale edge thin, narrow, deflexed;
veins distinct and slightly prominent on upper sur-
face. Fertile frond: stipe 38 cm; lamina 33 by 3 cm,
base decurrent to a narrow wing 3 cm long.
Distr. Malesia: Central West Sumatra. Only
known from type collection.
Ecol. In forest, at 1500 m.
Note. This is closely related to E. sumatranum,
and was collected in the same region, but is so much
larger that I think it should stand as a distinct
species. It should be noted that ALSTON also collec-
ted typical E. sumatranum, and no intermediates
between the two have been found.
46. Elaphoglossum spongophyllum BELL ex HOoLrtT-
TUM, Blumea 14 (1966) 325. — Type: CLEMENS
31869, N. Borneo, Mt Kinabalu, Upper Kinataki
R. (BO).
Rhizome short; scales to 15 by 2!/, mm, acumi-
nate, rather thin, flat, pale when young, later red-
brown, edges with a few hairs; phyllopodia
11/,-2 cm long. Sterile frond: stipe 8-15 cm long,
pale; lamina thick, rigid, light brown when dry, to
33 by 7 cm, widest at or above middle, base rather
narrowly cuneate, little decurrent, towards apex +
abruptly narrowed to a rounded tip; pale edge
thin, very narrow; veins obscure; costa beneath
broad, pale, little prominent; scales on lower
surface with very small dark centre and 2-4
slender radiating arms. Fertile frond: stipe 15 cm;
lamina 20-25 by 3'/,-4 cm, widest above middle,
base rather narrowly cuneate, apex abruptly
narrowed to a narrowly rounded tip; spores
c. 37 um long, with much-folded perispore.
Distr. Hainan; in Malesia: Malay Peninsula,
N. Borneo (Mt Kinabalu).
Ecol. Epiphyte, at 2150-2920 m in N. Borneo,
1550-1850 m in Malaya.
Notes. The only Malayan specimen is from
G. Tahan. The Hainan specimen (MCCLURE
20066) was distributed as E. austrosinicum MATT.
& Cur., which has acuminate fronds much more
1978]
LOMARIOPSIS GROUP (Holttum)
311
decurrent at the base. E. spongophyllum has some-
what the same shape of frond as E. sumatranum,
but it always dries brownish, and the spores are
very different.
47. Elaphoglossum blumeanum (FEE) J.SmM. Ferns
Brit. & For. (1866) 106. — Acrostichum blumeanum
Fee, Hist. Acrost. (1845) 62, excl. syn. Olfersia
blumeana PRESL. — Type: CuminG 194, Luzon
(P; dupl. in US).
E. copelandii Curist, Philip. J. Sc. 2 (1907) Bot.
176; v.A.v.R. Handb. (1908) 717; CopeL. Fern FI.
Philip. (1960) 277. — Type: CopeLaNnp 1541,
Mindanao (P; dupl. in B, US).
Acrostichum viscosum (non Sw.) BL. FI. Jav. Fil.
(1829) 27; Racis. Fl. Btzg 1 (1898) 46.
E. petiolatum [non (Sw.) UrRB.] v.A.v.R. Handb.
(1908) 717; Suppl. (1917) Corr. 29; BACKER &
PostH. Varenfl. Java (1939) 250.
E. yunnanense [non (BAKER) C.CHR.] HOLTTUM,
Rev. Fl. Mal. 2 (1954) 455, f. 264. — Fig. 23a-f.
var. blumeanum.
Rhizome creeping, 3-5 mm @ when dry, bearing
fronds close together or to 1 cm apart in each of two
ranks; scales c. 5 mm long, 1/, mm wide, dark
brown, glossy, rigid, acuminate, + contorted with
inrolled edges which bear a few teeth or stiff hairs;
phyllopodia to 8 mm long. Sterile frond: stipe
12-30 cm long, near base with dark spreading
scales as rhizome but often with numerous stiff
marginal hairs, rest of stipe + persistently covered
with small stellate scales; lamina to 50 by 31/, cm,
rather thin, base rather narrowly cuneate, apex
acuminate; thin pale edge very narrow; veins
slightly prominent on lower surface; costa beneath
rounded, prominent; scales on upper surface soon
abraded, thin, pale, appressed, + circular with
rather long slender marginal hairs appressed to
surface; scales on lower surface rather persistent,
very small but with 3-5 stiff marginal hairs to al-
most 1 mm long spreading away from surface, on
old fronds often abraded but leaving resin-dots.at
points of attachment. Fertile frond: stipe 20-35 cm;
lamina to 30 by 1.4 cm.
Distr. Malesia: Sumatra, Malaya, Java, Borneo,
Celebes, Philippines.
Ecol. Epiphyte at 700-2000 m, fronds pendu-
ous.
var. philippinense Curist MS, var. nov. — Type:
ELMER 6509, Baguio, Luzon (P; dupl. in B, MICH).
Cum varietate typica squamulis rhizomatis
frondisque congruente, differt: frondibus multo
minoribus; stipite frondis sterilis 4-6 cm longo,
lamina ad 15 x 1.5 cm; stipite frondis fertilis
10-18 cm longo, lamina 5-12 cm X< 6-8 mm.
Distr. Malesia: Philippines (Luzon). Several
collections.
Notes. The name E. blumeanum was published
by JoHN SMITH in his list of CumMING’s Philippine
ferns, with a reference to Acrostichum viscosum
BL. non Sw.; but as the generic name Elaphoglos-
sum had not then been validly published JoHN
SMITH’s new name was illegitimate. FEE was the
next author to publish the epithet b/umeanum for
this species; he referred to J. SMITH (the illegitimate
publication of 1841) but not to BLume, and
described and cited CumiNG 194 only, for which
reason the CUMING specimen must be the type of
Acrostichum blumeanum FEE.
This species is nearly allied to E. petiolatum
(Sw.) Urs. of the West Indies. There are also
closely allied forms in Africa and the Mascarene
Is. (E. salicifolium (WILLD. ex KAULF.) ALSTON)
and India (Acrostichum stelligerum WALL. ex
BAKER), and it would be possible to regard all as
varieties of one species. E. blumeanum is however
larger than the others, and appears to have a greater
difference between scales of the upper and lower
surfaces, the latter being especially distinctive. The
name E. yunnanense (BAKER) C.Cur. is probably
to be regarded as synonymous with Acrostichum
stelligerum, the latter epithet has priority but has
not yet been formally transferred to Elaphoglossum.
48. Elaphoglossum miniatum (CHRIST) CurisT,
Monogr. Elaph. (1899) 72, f. 30; v.A.v.R. Handb.
(1908) 716. — Acrostichum miniatum Curis,
Verh. Nat. Ges. Basel 11 (1895) 254, t. 3, f. 25-27;
Ann. Jard. Bot. Btzg 15 (1898) 175. — Type:
SARASIN 954, Central Celebes (BAS). — Fig. 23g-j.
Rhizome and scales as E. blumeanum. Sterile
frond: stipe 7 cm or more, bearing spreading scales
2-3 mm long, paler than on rhizome and with
many stiff marginal hairs, also many appressed
orbicular ciliate scales; lamina to 25 by 2!/, cm or
more, shape as E. blumeanum; scales on upper
surface very pale, flat, appressed, + circular with
fringe of pale hairs shorter than width of scale,
close to edge of upper surface a fringe of similar
but elongate scales (1 mm long) spreading at right
angles to edge of lamina and beyond it; scales on
lower surface red-brown, a little smaller than those
on upper surface, with inflexed edges bearing
thicker much longer stiff hairs (to 1/, mm long).
Fertile frond: stipe 15 cm or more; lamina to 30 cm
long, hardly 1 cm wide.
Distr. Malesia: Celebes (two collections); New
Guinea (?).
Ecol. At c. 1000 m altitude.
Note. Some New Guinea specimens seem some-
what intermediate between this species and
E. heterolepium as regards scales on the lower
surface, but the types of the two species (both
from Celebes) are very distinct.
49. Elaphoglossum heterolepium v.A.v.R. Bull.
Jard. Bot. Btzg II, 16 (1914) 13; Handb. Suppl.
(1917) 426. — E. petiolatum (Sw.) URBAN, var.,
C.Cur. Gard. Bull. S. S. 7 (1934) 291. — Type:
RACHMAT 522, partim, Central Celebes (BO). —
Fig. 23k—m, 24.
Rhizome and scales as E. blumeanum. Sterile
frond: stipe to 20 cm or more long; lamina light
green when fresh, drying dark, thin, 25-45 by
2-2.7 cm, apex acuminate, base more abruptly
narrowed and slightly decurrent; thin edge narrow,
reflexed; veins distinct; costa rounded and promi-
nent beneath, bearing some dark appressed scales;
scales on upper surface pale, thin, flat, circular,
short-ciliate, those near edge of lamina sometimes
spreading a little beyond the edge; scales on lower
surface often (always?) peltate, similar in shape to
those of upper surface but firmer and more per-
sistent, when dry with edges turned away from
lamina-surface, thus + cup-shaped. Fertile frond:
312 FLORA MALESIANA [ser. II, vol. 14
i
Li
[4
ie
MU
lee
ide
L
hi
ry
UE
L
[be
Fig. 23. Elaphoglossum blumeanum (FE£) J. SM. a. Habit, < 4/3, b-c. scales of upper surface of frond,
x 20, d. scales of lower surface , x 20, e. scale from stipe, < 7, f. edge of sterile frond. — E. miniatum
(CHRIST) CurRIsT. g. Scale from upper surface of frond, < 20, A. marginal cells of g, « 65, i. scale from
edge of upper surface, < 20, 7. scale from lower surface, « 20. — E. heterolepium v.A.v.R. k. Habit,
x 1/3, /. scale from upper surface of frond, < 30, m. scale from lower surface, « 30 (a—f L 908.329-719,
g-j SARASIN 954, k NGF 19082, /-m CLEMENS 29064).
LOMARIOPSIS GROUP (Holttum)
313
Fig. 24. Elaphoglossum heterolepium v.A.v.R. Lower surface of frond, showing scales and cells of epider-
mis, <7175¢
stipe somewhat longer than sterile; lamina 15-
30 cm long, 9-15 mm wide.
Distr. Malesia: N. Borneo, Celebes,
Guinea.
Ecol. Epiphyte or on rocks, at 1000-2300 m.
Note. The type does not include a fertile frond;
with it in the same collection are two fronds of
E. blumeanum, which appears usually to occur at
lower altitudes than E. heterolepium in Borneo and
Celebes.
New
50. Elaphoglossum resiniferum Ho_truM, Blumea
14 (1966) 324. — Type: WAKEFIELD 1466, Papua,
Central Division, Astrolabe Range (BM).
Rhizome creeping, 4-5 mm @, bearing fronds
close together; scales dark brown, glossy, 2-3 mm
long, not acuminate, somewhat crisped; phyllo-
podia less than 5 mm long. Sterile frond: stipe
5-7 cm long, pale, almost covered with appressed
broadly ovate subentire brown scales less than
1 mm long; lamina thinly coriaceous, 20-26 cm
long, 8-9 mm wide, widest about middle, narrowed
gradually to decurrent base and caudate apex; edge
reflexed and inrolled when dry; veins distinct on
lower surface; costa rounded and raised beneath;
scales on upper surface of costa as stipe but with
short marginal projections of one cell; on upper
surface of lamina few persistent scales, same shape
as on costa but thin, pale, with short marginal
unicellular hairs; on lower surface many small spots
of dark reddish resin, apparently on sites of former
scales. Fertile frond: stipe 11 cm; lamina 18 cm by
7 mm, widest above middle, base gradually and
narrowly decurrent, apex not caudate.
Distr. Malesia: E. New Guinea. Only known
from type specimen.
Ecol. On wet rocks in creek bed, at 2400 m.
Species not occurring in Malesia
Elaphoglossum conforme (Sw.) J. SMITH in
Hook. J. Bot. 4 (1841) 148. — Acrostichum con-
forme Sw. Syn. Fil. (1806) 10, 192.
This species is confined to St Helena (type) and
South Africa. The name has been used confusedly
for species in various other parts of the world,
including Malesia; Hooker cited Acrostichum
angulatum BL. as a synonym of A. conforme. See
E. recommutatum, no 38, supra.
Elaphoglossum gorgoneum (KAULF.) BRACK. in
Wilkes, U.S. Expl. Exp. 16 (1854) 74. — Acrosti-
chum gorgoneum KAuLF. En. Fil. Chamisso (1824)
63
This species is confined to Hawaii. The specimen
described and figured as A. gorgoneum by BLUME
(Fl. Jav. Fil. 28, t. 8) is E. angustatum (SCHRAD.)
Hieron. of South Africa. BLUME’s specimen (L)
was presumably collected at the Cape by someone
travelling to Java, and later mixed with others
collected in Java (other examples of this are cited
by BACKER & PostHuMus, Varenfi. Java p. 144).
314
BLUME’s specimen was subsequently re-named
Acrostichum conforme var.javanicum by METTENIUS
(Ann. Mus. Bot. Lugd.-Bat. 4, (1869, 292). The
references to E. gorgoneum by v.A.v.R. (Handb.
712, Suppl. 423) are based partly on BLUME, partly
on E. pellucido-marginatum CHRIST.
Elaphoglossum hirtum (Sw.) C.Cure. Ind. Fil.
(1905). — Acrostichum hirtum Sw. in Schrad. J.
Bot. 1800/2 (1801) 10. — Acrostichum squamosum
Sw. non CAvy.
A West Indian species. BEDDOME gave the name
E. squamosum to plants from S. India and Ceylon,
and included Sumatra in the distribution of the
species (BEDD. Handb. 420); this statement was
copied, under the name E. hirtum, by v.A.v.R.
(Handb. 717), but I have seen no specimens of the
Indian species from Sumatra and do not know the
origin of BEDDOME’s statement.
Elaphoglossum latifolium (Sw.) J.Sm. Lond. J.
Bot. (1842) 197. — Acrostichum latifolium Sw.
Prodr. (1788) 128.
A tropical American species, described originally
from Jamaica. BEDDOME (Handb. 416) so named
plants of E. angulatum (BL.) Moore in Ceylon and
S. India which he had previously called E. /auri-
folium.
Elaphoglossum laurifolium (THOUARS) Moore,
Ind. Fil. (1857) xvi. — Acrostichum laurifolium
THouaRS, FI. Tristan d’Ac. (1804) 31.
Confined to Tristan d’Acunha. The use of this
name for species in other parts of the world has
caused much confusion; in Malesia the species
E. angulatum and E. commutatum have been so
named. BACKER & PostHuMus (Varenfi. Java
p. 251) have cited Acrostichum gorgoneum BL. and
several other species as synonyms of E. /aurifolium.
Elaphoglossum pellucidum Gaupb. in Vaillant,
Voy. Bonite Bot. Atlas (1844) t. 79, f. 5; MoRTON,
Contr. U.S. Nat. Herb. 38 (1967) 44. — Acrosti-
chum micradenium Fre, Hist. Acrost. (1845) 43,
t. 8, f. 1. — E. microphyllum v.A.v.R. Bull. Dép.
Agr. Ind. Néerl. 18 (1908) 35; BACKER & POSTH.
Varenfl. Java (1939) 249.
E. pellucidum is a Hawaiian species. The type of
E. microphyllum is a specimen (now in Herb. BO)
from the herbarium of Dr PLoEM, who lived in
Java for thirty years. The specimen bears no origi-
nal label; the label written at the time of acquisition
at Bogor bears the locality Java, probably on the
assumption that all PLOEM’s specimens were collec-
ted in Java. Other specimens from PLOEM have
also been shown to bear incorrect localities (see
Cyclopaedia of Collectors, Fl. Males. I, 1, 1950,
XXViii, 409).
FLORA MALESIANA
[ser. II, vol. 14
Doubtful species
Acrostichum cumingii Fée, Hist. Acrost. (1845)
4
FEE cited CUMING 193 as type, but his description
differs considerably from all specimens of CUMING
193 which I have seen (BM, K, P, US) in scales and
in size and shape of fronds; as COPELAND has stated
(Fern FI. Philip. p. 278) FEe’s description would
better fit E. callifolium. The specimen from which
FEE prepared his description has not been found at
Paris. I therefore regard FEE’s name as of doubtful
application, and have referred all specimens of
CuMING 193 which I have seen to E. luzonicum
COPEL.
Acrostichum decurrens Desv. Berl. Mag. 5 (1811)
310.
The type of this species is at Paris. BELL reported
upon it in some detail (see E. amblyphyllum supra,
no 30) and showed that it is not conspecific with
the Java species so named by BLume. I have also
examined the specimen, which consists of small
detached fronds, lacking rhizome. Its origin is
described as Ind. or. I cannot identify it with any
of the Malesian species of Elaphoglossum here
described, and the specimen is so unsatisfactory
that I would prefer to regard the name E. decurrens
DEsy. as a nomen dubium.
Excluded
Elaphoglossum borneense (BURCK) C.Cur. Ind.
Fil. (1905) 303. — Acrostichum borneense BURCK,
Ann. Jard. Bot. Btzg 4 (1884) 99 = Syngramma
borneensis (HOOK.) J.SM. (Gymnogramma borneen-
sis HOOK.).
It is to be noted that the species of BURCK and
HOooKER were based on different types, but they
are undoubtedly conspecific.
Names which should not be recognized
JOHN SMITH regarded the generic name Elapho-
glossum as having been published by ScuHotr in
1834, but as SCHOTT gave no description the pub-
lication has no status according to the present
Code. The first publication of the name with a
description was by SMITH himself, in Aug. 1841.
But a few months previously JOHN SMITH had
published two new binomials, transferring species
from the genus Acrostichum to Elaphoglossum.
These two binomials are therefore now regarded
as nomina nuda, but they have been recognized by
some authors, owing to the fact that the invalidity
of ScHoTT’s name has only recently been noticed
(it was considered valid by CHRISTENSEN, in Index
Filicum). These names are therefore cited below,
to indicate their status.
Elaphoglossum blumeanum J.SM. in Hook. J. Bot.
3 (May 1841) 401, nom. nud.
Elaphoglossum obtusifolium J.SmM. l.c., nom. nud.
6. BOLBITIS'
ScHoTT, Gen. Fil. (1835) ad t. 13; C.Cur. Ind. Fil. Suppl. 3 (1934) 102; BACKER &
PostH. Varenfl. Java (1939) 80; CopeL. Gen. Fil. (1947) 115; HoL_trum, Ferns
(1) Treatment by E. Hennipman, Leyden.
1978] LOMARIOPSIS GROUP (Hennipman) 315
Malaya (1954) 461; CopeL. Fern Fl. Philip. (1960) 254; HENNIPMAN, Leid. Bot.
Ser. 2 (1977) 123, with full synonymy.
Egenolfia SCHOTT, Gen. Fil. (1835) ad t. 16; BACKER & PosTHu. Nat. Tijd. N. I. 93
(1933) 152; Varenfl. Java (1939) 84; HoL_trum, Ferns Malaya (1954) 459; CopeEL.
Fern FI. Philip. (1960) 265. — Polybotrya sect. Egenolfia DiEts in E. & P. Nat. Pfl.
Fam. 1 (1900) 195; v.A.v.R. Handb. Mal. Ferns (1908) 722.
Campium PRESL, Tent. Pterid. (1836) 238, pl. 10 (22-23); CopeEL. Philip. J. Sc. 37
(1928) 341, p.p.; BACKER & PosTH. Nat. Tijd. N. I. 93 (1933) 157, p.p.
Edanyoa Copet-. Philip. J. Sc. 81 (1952) 22, pl. 17. — Fig. 25, 26a—c, 27-33.
Rhizome creeping or low-climbing, unbranched or with accessory branches
dorsally (and laterally), with 2-6 rows of fronds, ventrally (and laterally) with roots;
scales -- appressed, pseudo-peltate, triangular from a perfoliate or subcordate base,
up to 15 mm long, usually subclathrate, sometimes (in B. sinuata) + opaque, with
sparsely set, slender, uniseriate, thin-walled glandular hairs; vascular system
dorsiventric, with a broad gutter-shaped ventral bundle and 1-3 (in B. heteroclita
rarely up to 4) dorsal bundles (fig. 25d). Fronds usually close together, in B. hetero-
clita sometimes spaced; petiole (long-) decurrent on the rhizome, near the lamina
base with 1(—3) median and on either side a lateral longitudinal ridge (fig. 25e),
aerophores linear, pale, present laterally on either side and especially conspicuous
in the basal part of young fronds, in cross-section near the base with a +
U-shaped arrangement of 3-16 vascular bundles of which the two anterior ones
are largest. Sterile fronds simple or (bi)pinnate with the pinnae alternate or +
opposite and usually continuous with the rachis, in some species + subarticulate,
the two lowermost pinnae usually conform to the other pinnae, sometimes deltoid,
sessile or shortly petiolulate, usually herbaceous, sometimes coriaceous, with
usually one, rarely more subterminal or terminal bulbils, surface when young apart
from scales densely set with uniseriate glandular hairs; venation pattern: veins free
(ser. Egenolfianae) or variously anastomosed and with or without included free
veins in the areoles. Fertile fronds of similar shape as the sterile ones though with a
proportionally larger petiole and a smaller lamina, usually completely acrostichoid,
sometimes pteridoid or moniliform; spores with a thin exospore and a variously
shaped perispore (fig. 26a—c). — Chromosomes n = 41, 82; 2n = 82, 123.
Distr. Pantropic. In my monograph (/.c.) I recognized 44 species arranged in 10 series (of which only
ser. Bolbitianae is pantropic): Africa incl. Madagascar: 9 spp., America: 14 spp., Asia and the Pacific:
21 spp. of which 12 in Malesia, Australia (Queensland): 3 spp.
Fossils. No fossils can be attributed to the genus. B. coloradica R. W. BROWN from the Cretaceous has
now been recognized by REVEAL et al. (Bringham Young Univ. Geol. Stud. 14, 1967, 239) as a drynarioid
fern, and transferred by them to Astralopteris.
Ecol. The species are all forest ferns; most of them grow in seasonally dry forest, others in everwet
habitats. They mostly occur at low altitude, only a few species being sometimes found above 1500 m. The
greater part of the species occur on rocks and especially in stream-beds, a naturally disturbed habitat.
Some otherwise terrestrial species are sometimes also found as low-climbers (or as low-epiphytes). Some
species are not rarely reported to form pure stands of many closely aggregated plants in forests (B. sinen-
sis). Of B. heteroclita several forms occur which may cover (rocks of) stream-banks completely (HOLTTUM,
Ferns Malaya, 1954, 463), one of these being an autotriploid.
Morph. The morphology, in particular of the Indian species, has been studied by NAYAR c.s. (for
references see KAur, J. Linn. Soc. Bot. 68, 1974, 153-162). In my monograph an elaborate treatment of
the morphology and anatomy of all species is included to which the reader is referred for details.
The rhizome of some species bears + conspicuous buds situated on the posterior side of the leaf-bases ;
they may develop into accessory branches. The stele is a dorsiventral dictyostele with wide overlapping
leaf-gaps. The traces running into the buds or the accessory branch traces are situated at the posterior
316 FLORA MALESIANA [ser. II, vol. 14
Fig. 25. Bolbitis virens (Hook. & Grev.) ScHoTT. a. Juvenile plant, x 4/,, b-c. pinnae, nat. size. —
B. heteroclita (PRESL) CHING. d. CS of rhizome, <x 4; db = dorsal bundle, It = leaf trace, r = root, rt =
root trace, vb = ventral bundle. — B. virens. e. Adaxial view on junction of rachis with pinnae, and CS
of lower part of rachis, x 7 (a-c HENNIPMAN 3320, d MANN 5.n., e HENNIPMAN 4047).
side of the two lateral leaf-gaps only, and are formed in association with root traces and leaf traces in an
obviously characteristic arrangement.
Aerophores are part of the fronds but are continuous on the rhizome for some length. On the rhizomes
of living material they show much variation in shape and size.
The venation pattern provides important characteristics for the discrimination of taxa. In all the species
the secondary veins run parallel; typical differences are therefore expressed by the tertiary efc. veins only.
See fig. 27, 31g, j, k. The venation in the pinnate fronds is anadromic.
Within the genus different types of venation occur. Veins are free in ser. Egenolfianae. In ser. Hetero-
clitae and ser. Quoyanae the pattern is sagenioid; the veins anastomosing in a reticulate pattern, with +
isodiametric or elongate, angulate areoles, and generally without recurrent included free veins. In ser.
Bolbitianae the veins anastomose to form a costal areole and one to many smaller distal ones, the veins
near the margin running + parallel, the areoles with or without excurrent included free veins.
A so-called irregular venation, i.e. a venation in which the arrangements of the veins in the areas in-
cluded by the secondary veins are markedly different, is often (not always!) present in alloploids, hybrids,
and in crossings between cytotypes of one species.
The morphological evidence as given in my monograph indicates that divergent evolution in ferns with
a sagenioid venation — the condition which I regard to represent the original condition in the genus —may
lead to species having either a free venation or a venation with several types of included free, ex- and/or
recurrent veins.
All species have bulbils subterminally (or terminally) on the sterile (and fertile) fronds. Bulbils are +
globular, persistent structures, situated adaxially; they are covered with scales similar to those on the
rhizome. Development into mature plants occurs when the apex of the mother leaf strikes the ground;
under humid conditions bulbils may develop into small plants on the erect lamina. Plants grown from
bulbils stay connected with the mother plant for some time (‘walking ferns’).
The shape of the fertile segments is usually about the same (though much contracted) as that of the sterile
ones. The margin of the segments often lack the prominent marginal projections found in the sterile
material; bulbils are less prominent or even absent. Fronds that are in part fertile and in part sterile —
so-called intermediate fronds — do not rarely occur and show much variation as to the sporangial
arrangement, also in one species.
The venation pattern of the fertile segments is similar to that in the sterile ones; free included veins and
small areoles are however less frequent or even absent. Nayar c.s. (see KAurR, /.c.) reported a special
kind of venation pattern occurring in the fertile fronds of the present genus; this is incorrect.
1978] _ LOMARIOPSIS GROUP (Hennipman) 317
Fig. 26. Scanning electron micrographs of spores of Lomariopsidoid ferns. a. Bolbitis sinensis, < 1000
(HENNIPMAN 3229); b. B. appendiculata ssp. appendiculata, x 1000 (BUNAK 743); c. B. angustipinna,
< 1000 (HENNIPMAN 3536); d. Lomariopsis intermedia, x 500 (BRASS 27982); e. L. kingii, x 750 (BRASS
32384); f. Elaphoglossum callifolium, x 1000 (LORZING 13523)5
318 FLORA MALESIANA [ser. II, vol. 14
Insertion of the sporangia is variable; in some species the sporangia are situated all over the lower
surface, in other species they are present on the veins only; also both conditions may be present in one
species. Sporangia are formed in dense masses. In spite of the basipetal development of the fertile frond,
sporangia of all parts of the frond reach maturity at the same time.
Spores as seen with the light microscope are monolete, biconvex, and provided with a variously shaped
brown perispore. The exospore is thin, structurally not differentiated, and shows a short leasura. The
perispore as seen from cross-sections studied with the electron microscope (HENNIPMAN, Acta Bot.
Neerl. 19, 1970, 671-680) is composed of two more or less distinct elements, of which the outer perispore
can also be recognized with the light microscope. Using properties of the (outer) perispore, three types of
spores can be recognized with the light microscope. In B. appendiculata the perispore is reticulate and
cristate. A smooth and undulate perispore is a characteristic of ser. Bolbitianae. All other species have a
smooth, cristate-undulate or cristate perispore. Although some are characterized by either a cristate or a
cristate-undulate perispore, the spores of several other species display both types of perispore as wellas the
intermediates.
Karyology. The haploid chromosome number of a considerable number of taxa as listed in my
monograph was either 41 or 82. One species (B. sinuata) showed a weak indication of aneuploidy. In ser.
Bolbitianae and ser. Egenolfianae only diploids were found. Of 20 specimens belonging to the 4 Malesian
species of ser. Heteroclitae and ser. Quoyanae, 12 were diploids and 8 (auto)triploids; this obscured the
delimitation of species of these two series in the past. Thus far apogamy has not been reported for the genus.
Gametophytes. Cordate(-elongate) and elongate gametophytes have been reported by NAYAR &
Kaur (Bot. Rev. 37, 1971, 295-396), ATKINSON (in Jermy c.s. J. Linn. Soc. Bot., Suppl. 1, 1973, 81) and
HENNIPMAN (/.c.). The gametophytes are either naked or may bear uniseriate glandular hairs or small
glandular scales (the diploid prothallus of B. repanda).
Juvenile fronds. The ontogenetic frond stages of several species are surprisingly diverse. During
ontogenesis features of the frond may become gradually or + abruptly more complex when subsequent
fronds on a single rhizome are compared. Abrupt changes were observed for instance in the shapes of the
terminal segment and the pinnae, and the venation pattern (fig. 25a-c).
Also, comparable frond stages of different juvenile plants of one species may show variation. For in-
stance in B. virens juvenile fronds were found with a triangular terminal segment (the less complex condi-
tion) and a rather intricate venation, as well as fronds showing a terminal pinna (the complex condition)
but with a rather simple type of venation. This kind of variation — though often less clearly expressed —
is also found in the adults. The variation in the morphology of the fronds of adult plants can to a certain
extent be predicted when the morphology of the juvenile fronds is known.
Speciation. From the comparative study of the juvenile fronds it may be deduced that neotenous
processes may have played an important role. Quite a number of dwarfs formerly given distinct status
(e.g. Edanyoa difformis) could be referred to the B. heteroclita complex after it was found out that the
mature fronds of these dwarfs were similar to both the ontogenetic frond stages of well-developed plants
and the juvenile fronds grown from bulbils attached to such plants. The juvenile fronds grown from bulbils
are generally more complex than the juvenile fronds of the same size grown from sporelings.
The mature fronds of B. appendiculata ssp. appendiculata (ser. Egenolfianae) are surprisingly similar
to the juvenile fronds with free veins as found in several Asian species of ser. Bolbitianae. The idea that
B. appendiculata arose by retention of certain juvenile characters from ser. Bolbitianae is supported by the
occurrence of several hybrids between species belonging to the two series.
The recognition of precocious fructification in relation to the occurrence of morphologically different
ontogenetic frond stages has been of great importance for the circumscription and phylogenetic considera-
tions of the taxa.
Specific delimitation. Hybridisation between different species and between cytotypes of one
species, auto- and alloploidisation is supposed to be a common phenomenon in the genus. Precocious
fructification further adds to the surprising morphological variation found in several species(-aggregates).
Hybrids have aborted spores or aborted spore-mother-cells, and may multiply vegetatively (a property
of all Malesian taxa).
Taxonomy. The genus Bolbitis was founded by Scuott (Gen. Fil. 1835, ad t. 13) for a part of Acro-
stichum with a creeping rhizome and anastomosing veins. At the same time (/.c. ad t. 16) he accommodated
the Asian species with free veins in Egenolfia, keeping the American free-veined species separate in Poly-
botrya. Pres (Tent. Pterid. 1836) and Fr&e (Hist. Acrost. 1845) referred the species to several different
genera whereas Hooker (Spec. Fil. 5, 1864) and BAKER (in Hooker.& Baker, Syn. Fil. 1865-1868) merged
all acrostichoids again in Acrostichum. In spite of J. SmirH (Gen. Fil. 1875) who reinstated Egenolfia,
Curist (Farnkrauter der Erde, 1897) and Diets (in E. & P. Nat. Pfl. Fam. 1, 4, 1899) referred all the
free-veined species again to Polybotrya, those with anastomosing veins to the heterogeneous Gymnopteris.
CHRISTENSEN (Ind. Fil. 1906) recognized Egenolfia, including the other species in an assemblage he called
Leptochilus. COPELAND (Philip. J. Sc. 37, 1928, 333-416) attempted to clear up the heterogeneities called
Gymnopteris by Diets and Leptochilus by CHRISTENSEN. He referred the Old World species to Campium
in which he included a number of unrelated ferns as well. CHING (Bull. Fan Mem. Inst. Biol. 2, 1931,
297-317) monographed Egenolfia. He later (in Christensen, Ind. Fil. Suppl. 3, 1934) reinstated Bolbitis
and largely delimited the genus as presented in my book. IwaTsuki (Acta Phytotax. Geobot. 18, 1959,
44-59) studied the Japanese species and was the first to unite Egenolfia and Bolbitis. The emended genus
he divided into 4 sections. In my monograph most of the species are accommodated in 10 series (4 in Asia
all except ser. Bolbitianae endemic), whereas several species of hybrid origin are separately ranked as
species incertae sedis.
1978] LOMARIOPSIS GROUP (Hennipman) 319
Notes. The treatment of the 12 recognized species is followed by the record of 5 hybrids and 2 dubious
species. Of the latter two categories only those have been inserted in the key which were collected in more
than one locality. Their numbers are preceded by H and D respectively.
The synonymy has been restricted to those names which were used for Malesian taxa. No types are
cited because they seem not useful for botanists consulting this Flora. For full synonymy and types see my
monograph.
So-called intermediate fronds, dwarfs and aberrant specimens are generally not considered in the des-
criptions.
The term segment is used for a portion of the lamina that has an axis and is not a pinna. Of the (central)
segments the index (length/width ratio) is generally given.
The number of pinnae given refers to the total number of pinnae to a frond.
Caudate apices of pinnae are not included in the measures given for the length of the pinnae.
Primordia of bulbils can be traced when the sterile fronds are examined in transmitted light. They
appear as small knobs terminally or subterminally.
The costa of the pinna and the simple lamina are termed the primary vein; the pinnately arranged
lateral veins are designated as the secondary veins.
Chromosome numbers given are taken from the list supplied in my monograph.
The key to the species is based on characters of the sterile fronds, and sometimes on those of the spores.
Identification will be possible for most of the material using a good hand lens and preferably also a
source of transmitted light to study the venation pattern.
KEY TO THE SPECIES AND HYBRIDS!
1. Plants small (dwarfed) and/or fronds simple.
2. Bulbil + terminal. Pinnae 16-30 . . . 22 ee eee eee a eB novocuineensis
2. Bulbils subterminal (or absent). Pinnae less.
3. Veins irregularly anastomosing, usually with many ex- and recurrent free veins. Fronds entire or
(bi)trifid ... . H.5. Leptochilus < trifidus
3. Veins regularly anastomosing, without free veins or with few ex- and recurrent free veins. Fronds
entire.
4. Fronds coriaceous.
5. Areoles (except the costal one) all of about the same shape, decreasing in size towards the margin
10. B. rivularis
5. Areoles of different size and shape, not decreasing in size towards the margin . . 8. B. sinuata
4. Fronds herbaceous (to subcoriaceous).
6. Fronds pinnate. Venation of terminal segment as in fig.31j,k...... . . 9. B.quoyana
6. Fronds entire or if pinnate with a venation as in fig. 31g or simpler . .. . 7. B. heteroclita
1. Plants not dwarfed. Fronds pinnate.
7. Veins free.
8. Bulbil + terminal on the lamina. Spines on the margin of the pinnae + flattened at their bases.
Base of f pinnae symmetrical. . . . . 5. _B. rhizophylla
8. Bulbil subterminal on the lamina. Spines on the margin not flattened at their bases. Base of pinnae
symmetrical or asymmetrical.
9. Perispore smooth. Base of pinnae symmetrical, margin lobed 1/,—2/,(—3/,) towards the costa
6. B. sinensis
9. Perispore reticulate. Base of pinnae either symmetrical or mein if symmetrical, margin of
the pinnae entire or lobed to 1/,("/,) towards the costa ..... . . . 4, B. appendiculata
7. Veins anastomosing.
10. Fronds + coriaceous.
11. Terminal segment conform to the pinnae, ud ee. ie Jk Bea Sen Os bse SINTEALA
11. Terminal segment triangular ...... 5, oe een SAO Bo rivolaris
10. Fronds herbaceous.
12. Fronds drying reddish. Perispore undulate . . . . . 2. B. scalpturata
12. Fronds drying greenish. Perispore undulate, cristate- undulate, or cristate.
13. Terminal segment conform to the pinnae or composed of 2 or 3 lobes.
14. Venation pattern with many excurrent included free veins.
iSssberispore undulate 1f) Sos else See A eee oe wirens War Compacts
15. Perispore cristate . . . . D.1. B. interlineata
14. Venation pattern without or with only few excurrent (and recurrent) included free veins.
16. Pinnae 2-10(-15) toafrond .. . , Py dae ies FS, oO 3) 4] Bo neterocita
16: Pimnge more tian’ lS¢o6'a lear ss SE, ee fe ee Bee
13. Terminal segment triangular.
17. ‘Bulbil + terminal. Small plant... «. . 202 « . 3 </:.+ » »./. » 21.- Bi movogainecnas
17. Bulbil subterminal.
18. Terminal segment narrowly triangular. Perispore undulate_. . . . 1. B. angustipinna
18. Terminal segment triangular. Perispore cristate or cristate-undulate.
(1) Only hybrids known from more than one locality are included.
[ser. II, vol
FLORA MALESIANA
320
—
Qs
Ny
et MO
SMU ENS
UU MAMI \
XS
rS>
Y
> XN
TI
ANS
A
()
! MAA Ant yAe:
WEIL VAL USA
to
9A); i.
lecta (VAN
12); I. Leptochilus x
4/;. a. Bolbitis angustipinna
(SF 29084), d. B. appen-
B. appendiculata ssp. vivipara var. neg.
h. B. sinensis (HENNIPMAN 322
ROSENBURGH s.n., L); m. B. sinuosa nm. foxii (PNH 8904); n. B. x
singaporensis (HOLTTUM s.n., SING).
B. repanda (SF 25585); j. B. sinuata (JOHNSON s.n., L); k. B. interlineata (BROOKS
(HENNIPMAN 3637), b. B. scalpturata (MOUSSET 562), c. B. virens var. compacta
trifidus (VAN ALDERWERELT VAN
Fig. 27. Venation patterns of sterile (a—b, d-n) and fertile (c) pinnae, x
diculata ssp. appendiculata (HOLTTUM s.n., SING); e-f.
BorssuM WAALKES 603); g. B. rhizophylla (LEROY ToPPING 655);
1978]
19. Venation pattern irregular. See fig. 271
19. Venation pattern regular.
LOMARIOPSIS GROUP (Hennipman)
321
12. B. repanda
20. Base of pinnae narrow-acute or -cuneate. Pinnae 7'/,-12 My, 1- aa cm H.4. B. x sinuosa
20. Base of pinnae wider. Pinnae usually larger
1. Series Bolbitianae
HENNIPMAN, Leid. Bot. Ser. 2 (1977) 147. —
Bolbitis ScHotr, Gen. Fil. (1835) ad t. 13, typo incl.,
pro genere.
Sterile fronds pinnate; lamina with a (primor-
dium of a) subterminal bulbil; pinnae 3—50, the
margin sometimes with inconspicuous spines in the
sinuses; terminal segment usually conform to the
pinnae, sometimes narrowly triangular; venation
pattern: veins variously anastomosing, always
with a costal areole, with or without excurrent
included free veins. Spores with a smooth, undu-
late perispore. — Chromosomes n = 41, 2n = (c.)
82
Distr. Pantropical; most diversified in Asia
(7 spp., of which 3 in Malesia).
Note. A very homogeneous series. B. angusti-
pinna, the most widespread Asian representative,
is most closely related to the species from America
and Africa.
1. Bolbitis angustipinna (HAyYATA) Ito, J. Jap. Bot.
14 (1938) 443; HENNIPMAN, Leid. Bot. Ser. 2 (1977)
152, f. 40a—f. — Leptochilus angustipinnus HAYATA,
Ic. Pl. Form. 5 (1915) 297, f. 119. — Campium
ae rnin CopPEL. Philip. J. Sc. 37 (1928) 381,
a8
Leptochilus cuspidatus (PRESL) C.CHR. var.
crenatus ROSENST. Hedwigia 56 (1915) 348.
[Acrostichum contaminans WALL. Cat. (1829)
n. 22, nomen. — Poecilopteris contaminans MOORE,
Ind. Fil. (1857) 8, nomen.] — Acrostichum crispatu-
lum CLARKE var. contaminans CLARKE, Trans.
Linn. Soc. Bot. 1 (1880) 580, pl. 84: f. 2A, 2C. —
B. contaminans CHING in C.Chr. Ind. Fil. Suppl. 3
(1934) 47; Iwatsuxk1, Acta Phytotax. Geobot. 18
(1959) 53, f. 9; DEVoL & Kuo, Fl. Taiwan 1 (1975)
348. — Fig. 26c, 27a.
Sterile fronds pinnate, 55-150 cm long; lamina
index 1-3, 30-90 by 20-45 cm, terminal segment
8-25 cm long, (firm) herbaceous, usually light to
dark green, sometimes with a purple tinge; pinnae
20-50, index 5—12, the central part usually with
parallel margins, 11-30 by 2-3(-5) cm, base +
symmetrical, acute, broadly attenuate or truncate,
margin usually slightly serrate-crenate, sometimes
lobed to 1/3 to the costa and with a short or incon-
spicuous spine in each sinus; terminal segment
usually narrowly triangular, sometimes + conform
to the central pinnae; venation pattern: veins
forming a costal areole and one to several rows of
distal areoles, included free veins absent, the veins
in the marginal strip excurrent and parallel; see
fig. 27a. Fertile fronds 55-160 cm long; pinnae
index (6-)9-20, 5-23 by 0.4-1.7 cm. Sporangia
situated usually all over the lower surface, some-
times with a sterile strip along the costa. Spores
smooth, undulate.
Distr. Ceylon, Himalayas eastwards to S. China
and Taiwan, southwards to N. Thailand; in
Malesia: Philippines (Luzon), one collection.
Ecol. On rocks in monsoon and evergreen
forest, 250-1500 m.
9. B. quoyana
2. Bolbitis scalpturata (FEE) CHING in C.Chr. Ind.
Fil. Suppl. 3 (1934) 50; BACKER & PosTH. Varenfl.
Java (1939) 82, p.p.; PostH. Ann. Bot. Gard. Btzg
vol. hors série (1944) 62; HENNIPMAN, Leid. Bot.
Ser. 2 (1977) 163, f. 43a—d. — Heteroneuron scalp-
turatum FEE, Hist. Acrost. (1845) 95, pl. 56, p.p. —
Leptochilus scalpturatus C.Cur. Ind. Fil. (1906) 387,
p.p.; V.A.V.R. Handb. Mal. Ferns (1908) 743, p.p.
— Campium scalpturatum Cope . Philip. J. Sc. 37
(1928) 383, f. 35; BACKER & PosTH. Nat. Tijd.
N. I. 93 (1933) 162.
Leptochilus reimersii RENSCH, Hedwigia 74 (1934)
249, pl. 7: f. 2. — Fig. 27b.
Sterile fronds pinnate, 25-90 cm long; lamina
index 1-3, 20-55 by 7-30 cm, terminal segment
9-20 cm long, herbaceous to subcoriaceous,
purplish or purplish-brown; pinnae 4-24, index
3-7, 5.5-17.5 by 1.5-4 cm, base + symmetrical,
narrowly to broadly cuneate, margin + entire or
finely serrate-crenate, without spines, apex acute to
(long-)acuminate; terminal segment usually con-
form to the central pinnae, sometimes narrowly
triangular and/or somewhat prolonged; venation
pattern: veins forming a network of a costal areole
and one to few transverse rows of smaller distal
ones, part of the areoles with few to several, mostly
excurrent included free veins; see fig. 27b. Fertile
fronds 30-70 cm long; pinnae index 3-8, 3-9 by
0.6—1.5 cm. Sporangia inserted mostly on and near
the veins, either all over the lower surface, arranged
acrostichoid, or along the margin only, arranged
pteridoid. Spores smooth, undulate.
Distr. E. Burma to Indo-China; in Malesia:
S. Sumatra (Lampongs), E. Java (Mt Tengger),
Lesser Sunda Is. (Bali, Sumbawa, Flores), S
Celebes (also Saleyer Is.), Philippines (Palawan,
Luzon).
Ecol. Showing a preference for a seasonal
climate, 0-1200 m.
Note. A somewhat critical species. The venation
pattern shows considerable variation; it may come
near to that of B. angustipinna or B. virens.
3. Bolbitis virens (HooK. & GrREvV.) SCHOTT, Gen.
Fil. (1835) ad t. 13; Hottrum, Ferns Malaya
(1954) 468, f. 275, p.p. — Campium virens PRESL,
Tent. Pterid. (1836) 239; Cope. Philip. J. Sc. 37
(1928) 388; HENNIPMAN, Leid. Bot. Ser. 2 (1977)
180, f. 47 & 48. — Leptochilus virens C.Cur. Ind.
Fil. (1906) 388, p.p.; v-A.v.R. Handb. Mal. Ferns
(1908) 741, p.p.; ibid. Suppl. (1917) 435, p.p.
var. compacta HENNIPMAN, Blumea 18 (1970) 149;
Leid. Bot. Ser. 2 (1977) 184, f. 48i, |. — Fig. 25a—c,
e, 27c.
Sterile fronds pinnate, 40-115 cm long; lamina
index 1-3, 20-75 by 15-50 cm, terminal segment
14-40 cm long, firm herbaceous; pinnae 6-22,
index 3-7, 10-30 by 2.5-6.5 cm, base + sym-
metrical, usually narrowly, sometimes broadly
attenuate or cuneate, margin usually + entire,
sometimes (bi)serrate or (bi)serrate-crenate; fer-
minal segment conform to the pinnae or somewhat
prolonged; venation pattern: veins forming a costal
322
areole and several to many equally large distal ones,
the arches with two or more excurrent free veins.
Fertile fronds 40-125 cm long; central pinnae index
3-8, 4-11.5 by 0.8-2 cm. Sporangia inserted all
over the lower surface. Spores smooth, undulate.
Distr. ?India (Nicobar Is.), S. Vietnam, Penin-
sular Thailand; in Malesia: Malay Peninsula
(Kedah, Selangor, Penang, Langkawi Is.). Fig. 28.
Ecol. Usually on rocks and often near streams in
(dry) evergreen forest, 0O-400(—700) m.
Bolbitis virens
-10 @ Var. virens
A var. compacta
var. deltigera
S
Fig. 28. Range of Bolbitis virens (Hook. & GREV.)
SCHOTT.
2. Series Egenolfianae
HENNIPMAN, Leid. Bot. Ser. 2 (1977) 185. —
Egenolfia ScHoTt, Gen. Fil. (1835 or 1836) ad t. 16,
typo incl., pro genere.
Sterile fronds pinnate; lamina with one, usually
subterminal, in one species (B. rhizophylla) +-
terminal bulbil; pinnae 14-90, the margin with
spines corresponding to the most distal acroscopic
tertiary vein of each secondary vein; terminal
segment triangular; venation: veins free. Spores
with either a smooth, cristate or cristate-undulate
perispore, or with a reticulate + cristate perispore.
— Chromosomes n = 41, 2n = 82.
Distr. S. India and Ceylon to W. Malesia,
northwards to S. Japan.
4. Bolbitis appendiculata (WILLD.) IWATSUKI, Acta
Phytotax. Geobot. 18 (1959) 48; HENNIPMAN,
Blumea 18 (1970) 147; Leid. Bot. Ser. 2 (1977) 185,
f. 49. — Acrostichum appendiculatum WILLD. Sp.
Pl. 5 (1810) 114; Hoox. Exot. Fl. 2 (1825) 108,
pl. 108; Sp. Fil. 5 (1864) 251, p.p.; BAKER in Hook.
& Baker, Syn. Fil. (1868) 415. — Polybotrya
appendiculata J. SmMitH, Hook. J. Bot. 4 (1841) 150;
v.A.v.R. Handb. Mal. Ferns (1908) 724, p.p. —
Egenolfia appendiculata J. SmiTH, Ferns Br. For.
(1866) 111, fig.; BACKER & PostH. Nat. Tijd. N. I.
FLORA MALESIANA
[ser. Il; vole
93 (1933) 153; Varenfl. Java (1939) 84, f. 14;
HottruMm, Ferns Malaya (1954) 459, f. 270; CoPEL.
Fern FI. Philip. (1960) 266; DEVoL & Kuo, FI.
Taiwan 1 (1975) 350, pl. 123.
Polybotrya vivipara HAM. ex Hook. Exot. Fl. 2
(1825) 107, pl. 107. — Egenolfia vivipara C.CHR.
Ind. Fil. Suppl. 3 (1934) 102. — B. hookeriana
IwATSUKI, Acta Phytotax. Geobot. 18 (1959) 49. —
B. appendiculata (WILLD.) IWATSUKI ssp. vivipara
HENNIPMAN, Blumea 18 (1970) 147; Leid. Bot. Ser.
ZAI TM AIS N50) Sie
Polybotrya marginata BL. En. PI. Jav. (1828) 100,
nom. superfl.; Fl. Java Filices (1829) 18, pl. 3; FEE,
Hist. Acrost. (1845) 75. — Polybotrya appendicu-
lata (WILLD.) J. SMITH var. marginata C.CuR. Bot.
Tidsskr. 32 (1916) 343.
[Acrostichum hamiltonianum WALL. Cat. (1829)
n. 28, nomen.| — Polybotrya hamiltoniana [PRESL,
Tent. Pterid. (1836) 236, nomen;] FEE, Hist. Acrost.
(1845) 78, nom. superfl. — Egenolfia hamiltoniana
FEE, Genres Polyp. (1852) 48, non ScHoTt, 1835 or
1836 (= B. appendiculata ssp. appendiculata). —
Acrostichum appendiculatum WILLD. var. hamil-
tonianum BAKER in Hook. & Baker, Syn. Fil.
(1868) 415. — Polybotrya appendiculata (WiLLD.)
Bolbitis appendiculata
A SSP. vivipara var. neglecta
OF. 6:4 " var. vivipara
Fig. 29. Range of Bolbitis appendiculata (WILLD.)
IWATSUKI.
1978]
LOMARIOPSIS GROUP (Hennipman)
323
J. SMITH var. hamiltoniana BEDD. Handb. Ferns Br.
India (1883) 426, f. 256, p.p.; V.-A.v.R. Handb. Mal.
Ferns (1908) 724, guoad nomen solum; ibid. Suppl.
(1917) 431, ditto.
B. appendiculata (WILLD.) IWATSUKI var. neglecta
HENNIPMAN, Blumea 18 (1970) 147; Leid. Bot.
Ser. 2 (1977) 197, f. 50i-n, 51. — Fig. 26b, 27d-f.
Sterile fronds pinnate, 15-100 cm long; lamina
index 2—6(—8), 10-80 by 2.5—20 cm, terminal seg-
ment 1-10 cm long, herbaceous, usually olivaceous;
pinnae (20-)25-60(-85), index 2-S(-7), 1.5-
10(-15) by 0.5-2.5 cm, base symmetrical to
strongly oblique, margin + entire, finely serrate,
or lobed to 3/3; (*/,) towards the costa; terminal
segment triangular, apex sometimes somewhat
prolonged; venation pattern: secondary veins on
either side with 2 or 3 tertiary veins; see fig. 27d-f.
Fertile fronds 15-90 cm long; pinnae index 1-6,
0.2-11 by 0.15-1.7 cm. Sporangia inserted all over
the lower surface or + restricted to the veins.
Spores with a reticulate cristate perispore. —
Chromosomes n = 41, 2n = 82.
Distr. S. India and Ceylon, NE. India eastwards
through SE.-E. continental Asia to S. Japan; in
Malesia: Malaya, Java, Philippines. Fig. 29.
Ecol. On rocks or in soil in deciduous and ever-
green forest, 0-1500 m.
Note. The reticulate perispore is unique in the
genus.
KEY TO INFRASPECIFIC TAXA
1. Fertile pinnae with a lamina. Sporangia situated
on the lower surface of the frond
a. ssp. appendiculata
1. Fertile pinnae with a narrow strip of lamina
present along the costa only. Sporangia facing
to all directions __b. ssp. vivipara var. neglecta
a. ssp. appendiculata — Acrostichum appendicu-
latum WiLLpD. — Polybotrya marginata BL. —
Fig. 27d.
Sterile fronds 15—60(—80) cm long, the rachis
with a narrow wing; pinna base oblique. Fertile
fronds: pinna index 1-6, 0.25-4 by 0.2-0.7 cm.
Sporangia inserted mainly on the veins or all over
the lower surface.
Distr. Ceylon, S. India, Himalayas eastwards to
S. Japan and Indo-China; in Malesia: Sumatra
(West Coast, Bencoolen, East Coast), Malay
Peninsula, Java (West, Central). Fig. 29.
Ecol. See the species.
b. ssp. vivipara (Hook.) HENNIPMAN, Leid. Bot.
Ser. 2 (1977) 195. — Polybotrya vivipara HAM. ex
Hook. — B. appendiculata (WILLD.) IWATSUKI
var. neglecta HENNIPMAN.
var. neglecta HENNIPMAN, Blumea 18 (1970) 147;
Leid. Bot. Ser. 2 (1977) 197, f. 50i—n, 51.
Polybotrya appendiculata (WILLD.) J. SMITH var.
rhizophylla auct. non (KAULF.) KUHN: KUHN, Ann.
Mus. Bot. Lugd.-Bat. 4 (1869) 225.
Polybotrya appendiculata (WILLD.) J. SMITH var.
hamiltoniana auct. non (FEE) BEDD.: v.A.v.R.
Handb. Mal. Ferns, Suppl. (1917) 431. — Fig. 27e, f.
Sterile fronds 30-50 cm long, the rachis with a
narrow or inconspicuous wing; pinna base -++ sym-
metrical or oblique. Fertile fronds: pinnae +
moniliform, index 5—15, 0.7-3 cm long, with a
narrow strip of lamina along the costa. Sporangia
mainly inserted at the endings of the lateral veins.
Distr. India (Andaman and Nicobar Is.); in
Malesia: West Java. Fig. 29.
Ecol. On stone, 70 m (type).
5. Bolbitis rhizophylla (KAULF.) HENNIPMAN,
Blumea 18 (1970) 148; Leid. Bot. Ser. 2 (1977) 199,
f. 52d—r, 53. — Gymnogramma rhizophylla KAULF.
En. Fil. (1824) 78. — Polybotrya rhizophylla
PRESL, Tent. Pterid. (1836) 231; Fee, Hist. Acrost.
(1845) 77; Metr. Fil. Hort. Lips. (1856) 24. —
Egenolfia rhizophylla F&e, Genres Polyp. (1852) 48;
CopEL. Fern FI. Philip. (1960) 266; DEVoL & Kuo,
Fl. Taiwan 1 (1975) 352.
Polybotrya intermedia [J. SMitH, Hook. J. Bot. 3
(1841) 72, nomen;] FEE, Hist. Acrost. (1845) 76,
pl. 40: f. 1. — Egenolfia intermedia FEE, Genres
Polyp. (1852) 48; CHING, Bull. Fan Mem. Inst.
Biol. 2 (1931) 308; CopeL. Fern FI. Philip. (1960)
266. — B. intermedia \waTsuKI, Acta Phytotax.
Geobot. 18 (1959) 49.
Polybotrya neglecta FEE, Hist. Acrost. (1845) 76,
pesos tent
Polybotrya serrulata [J. SmitH, Hook. J. Bot. 3
(1841) 401, nomen;] FEE, Hist. Acrost. (1845) 76,
pl. 39: f. ii*. — Egenolfia serrulata FEE, Genres
Polyp. (1852) 358. — B. serrulata IWATSUKI, Acta
Phytotax. Geobot. 18 (1959) 49.
Polybotrya exaltata BRACKENR. in Wilkes, U.S.
Expl. Exp. 16 (1854) 78.
Polybotrya duplicato-serrata HaAyaTA, Ic. PI.
Form. (1915) 305, f. 123A.
Egenolfia fluviatilis Cope. Philip. J. Sc. 38 (1929)
152, pl. 5; Fern Fl. Philip. (1960) 267. — B. cope-
landii lwatTsuxk1, Acta Phytotax. Geobot. 18 (1959)
49, nom. illeg., non CHING ex TARDIEU-BLOT &
C.Cur. 1938. — Fig. 27g.
Sterile fronds pinnate, 20-90 cm long; lamina
index 3-9, 18-80 by 4.5-15 cm, terminal segment
1-6 cm long, herbaceous, green to blackish, the
petiole and lower side of the basal part of the rachis
usually with small, blackish, spreading, -_ perma-
nent scales; rachis with a narrow wing except for
the lowest part; pinnae 35-90, up to 35 mm apart,
index 2-6, 2.5-8 by 0.8—-2 cm, base usually sym-
metrical, (narrowly) cuneate to subcordate, margin
Bolbitis sinensis
rhizophylla
Fig. 30. Range of Bolbitis sinensis (BAKER) IWATSUKI
and B. rhizophylla (KAULF.) HENNIPMAN.
324 FLORA MALESIANA [ser. II, vol. 14
we
Fig. 31. Bolbitis heteroclita (PRESL) CHING. a-f. Habits, x '/,, g. venation pattern of sterile pinna, x */,. —
B. quoyana (GAUDICH.) CHING. h-i. Habits, x 4/;, /-k. venation patterns of sterile pinnae, < */,.—B.novo-
guineensis HENNIPMAN. /. Habit, < 1/; (a PRICE 2518A, b Price 351, c BS 28843, d PNH 8811, e JAcoBs
7950, f HENNIPMAN 4069, g HENNIPMAN 3836, A WALKER T 10052, i SCHLECHTER 16163, 7 WALKER T 9604,
k ELMER 13468a, / BRAss 28008).
1978]
finely serrate, biserrate, or serrately lobed to '/;
towards the costa, the spines often tooth-like;
terminal segment narrowly triangular; venation
pattern: secondary veins on either side with 1-3
tertiary veins, see fig. 27g. Fertile fronds 20-80 cm
long; pinnae index 1-6, 0.4-3(-4) by 0.3-0.7 cm.
Sporangia inserted on and near the veins. Spores
with a smooth cristate-undulate perispore. —
Chromosomes n = 41.
Distr. Taiwan; in Malesia: Philippines (Balabac
I., Palawan, Mindoro, common in Luzon). Fig. 30.
Ecol. On rocks in moist forests and often near
riverbanks, 0—-1000 m.
6. Bolbitis sinensis (BAKER) IwAtTsuKi, Acta
Phytotax. Geobot. 18 (1959) 49; HENNIPMAN, Leid.
Bot. Ser. 2 (1977) 202, f. 53, 54. — Acrostichum
sinense BAKER, Kew Bull. (1906) 14. — Egenolfia
sinensis MAxoONn, Proc. Biol. Soc. Wash. 36 (1923)
173. — Campium sinense C.CuHR. Contr. U.S. Nat.
Herb. 26 (1931) 292.
Acrostichum appendiculatum WILLD. var. costu-
latum Hook. Sp. Fil. 5 (1864) 252. — Polybotrya
appendiculata (WILLD.) J. SMITH var. costulata
Bepp. Ferns Br. India (1865) 110, pl. 110. —
Egenolfia bipinnatifida J. SmMitH, Hist. Fil. (1875)
132; BACKER & PosTH. Varenfl. Java (1939) 85. —
B. sinensis (BAKER) IWATSUKI var. costulata
TAGAWA & IWATSUKI, Acta Phytotax. Geobot. 22
(1967) 102. — Fig. 27h.
Sterile fronds pinnate, 35-145 cm long; lamina
index 1—4(—S), 20-115 by 10-35 cm, terminal seg-
ments 8-35 cm long, herbaceous, dark green, +
glabrescent; rachis usually with a narrow wing
except for the lowest part; pinnae 14—40, index 3-7,
5.5-17.5(-21) by 1.5—5 cm, base + symmetrical,
broadly cuneate to auricled, margin lobed 1/,—
2/,(—3/,) towards the costa, lobes spaced to partly
overlapping, c. 6-8 mm wide at the base; terminal
segment narrowly triangular, the apex usually
either prolonged or flagelloid; venation pattern:
secondary veins on either side with 4-6 tertiary
veins, see fig. 27h. Fertile fronds 18-85 cm long;
pinnae index 2-8, 1.3-6 by 0.4-1.2 cm. Sporangia
usually mainly inserted on and near the veins.
Spores with a smooth cristate-undulate perispore.
— Chromosomes n = 41, 2n = 82.
Distr. S. China, E. Himalayas to Indo-China;
in Malesia: E. Java (once), Lesser Sunda Is. (Bali,
Sumbawa). Fig. 30.
Ecol. Usually terrestrial, creeping in soil or on
rocks, sometimes low-epiphytic (up to 1 m), in
(hill) evergreen (monsoon) forest, obviously a rare
plant, 0-1900 m.
3. Series Heteroclitae
HENNIPMAN, Leid. Bot. Ser. 2 (1977) 220.
Sterile fronds simple or pinnate; petiole with
3-14 vascular bundles; lamina with usually one
subterminal bulbil; pinnae 2—11(—15), the margin
without teeth or spines; terminal segment usually
conform to the pinnae, sometimes (in dwarfs)
triangular; venation pattern: veins usually com-
pletely anastomosing into a regular network,
areoles varying in size and shape. Spores with a
smooth, cristate or cristate-undulate perispore. —
Chromosomes n = 41, 2n = 82, c. 123.
Distr. NE. India to the Pacific, northwards to
LOMARIOPSIS GROUP (Hennipman)
325
S. Japan; throughout Malesia, except for the Lesser
Sunda Is.
7. Bolbitis heteroclita (PRESL) CHING in C.Chr. Ind.
Fil. Suppl. 3 (1934) 48; BACKER & PostH. Varenfi.
Java (1939) 83, f. 13; Hotrrum, Ferns Malaya
(1954) 462, f. 271; CopeL. Fern FI. Philip. (1960)
257; CHING, FI. Hain. (1964) 165, f. 75; DEVoL &
Kuo, FI. Taiwan 1 (1975) 348, pl. 122; HENNIPMAN,
Leid. Bot. Ser. 2 (1977) 221, f. 60. — Acrostichum
heteroclitum PresL, Rel. Haenk. (1825) 15, pl. 2:
f. 2. — Heteroneuron heteroclitum Fé, Hist.
Acrost. (1845) 92. — Chrysodium heteroclitum
KuHN, Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 294. —
Leptochilus heteroclitus C.CuHr. Ind. Fil. (1906)
385; v.A.v.R. Handb. Mal. Ferns (1908) 739. —
Campium heteroclitum CopreL. Philip. J. Sc. 37
(1928) 359, f. 13; BACKER & PosTH. Nat. Tijd.
N. I. 93 (1933) 163.
Nephrodium cuspidatum Pres_, Rel. Haenk.
(1825) 31. — Polystichum cuspidatum PRESL, Tent.
Pterid. (1836) 82. — Heteroneuron cuspidatum
PrRESL, Epim. Bot. (1851) 169. — Chrysodium
cuspidatum KUHN, Ann. Mus. Bot. Lugd.-Bat. 4
(1869) 293, p.p. — Leptochilus cuspidatus C.CHR.
Ind. Fil. (1906) 384, quoad nomen solum. —
Campium cuspidatum CoPEL. Philip. J. Sc. 37 (1928)
365, f. 19. — B. cuspidata CHING in C.Chr. Ind.
Fil. Suppl. 3 (1934) 48; CopeL. Fern FI. Philip.
(1960) 259.
Acrostichum flagelliferum WALL. ex Hook. &
GreV. Ic. Fil. (1827) pl. 23, p.p.; BL. En. Pl. Jav.
(1828) 104; Fl. Java Fil. (1829) 37, pl. 13, p.p. —
B. flagellifera Scuott, Gen. Fil. (1835) ad t. 13. —
Gymnopteris flagellifera BEDD. Ferns Br. India,
Suppl. (1876) 27; Cope. Polyp. Philip. (1905) 42,
p.p.
Acrostichum proliferum BL. En. Pl. Jav. (1828)
104, non Hook. 1844 (= B. subcrenata).
Leptochilus linnaeanus FEE, Hist. Acrost. (1845)
87, pl. 47: f. 2, excl. syn.; v.A.v.R. Handb. Mal.
Ferns (1908) 735; Suppl. Corr. (1917) 60. — Dendro-
glossa linnaeana FEE, Gen. Polyp. (1852) 81. —
Gymnopteris linnaeana CHRIST, J. de Bot. 19 (1905)
125; CopEL. Polyp. Philip. (1905) 4. — Leptochilus
heteroclitus (PRESL) C.CurR. var. linnaeanus CHRIST,
Philip. J. Sc. 2 (1907) Bot. 160. — Campium lin-
naeanum CopPEL. Philip. J. Sc. 37 (1928) 343, quoad
nomen solum; BACKER & PosTH. Nat. Tijd. N. I. 93
(1933) 159, excl. syn. — B. linnaeana C.Cur. Ind.
Fil. Suppl. 3 (1934) 198; BACKER & POosTH.
Varenfl. Java (1939) 80.
Cyrtogonium acuminatum BRACKENR. in Wilkes,
U.S. Expl. Exp. 16 (1854) 86.
Poecilopteris stenophylla Kurz ex T. & B. Nat.
Tijd. N. I. 27 (1864) 15; Copev. Philip. J. Sc. 37
(1928) 393. — B. stenophylla CuinG in C.Chr. Ind.
Fil. Suppl. 3 (1934) 50.
Chrysodium heteroclitum (PRESL) KUHN _ var.
subcrenatum KuuN, Ann. Mus. Bot. Lugd.-Bat. 4
(1869) 294, p.p.
Acrostichum modestum BAKER, J. Linn. Soc. 22
(1886) 231; C.Cur. in Copel. Philip. J. Sc. 37 (1928)
411. — Leptochilus modestus C.Cur. Ind. Fil.
(1906) 386. — Campium modestum CoPEL. Brit-
tonia 1 (1931) 76, f. 1. — B. modesta CHING in
C.Chr. Ind. Fil. Suppl. 3 (1934) 49.
Gymnopteris inconstans CoPeL. in Perk. Fragm.
FI. Philip. (1905) 177; Polyp. Philip. (1905) 43. —
326
Leptochilus inconstans CHRIST, Bull. Herb. Boiss. I,
6 (1906) 1005. — Leptochilus heteroclitus (PRESL)
C.CuHR. var. inconstans CHRIST, Philip. J. Sc. 2
(1907) Bot. 160. — B. inconstans CHING in C.Chr.
Ind. Fil. Suppl. 3 (1934) 48.
Leptochilus heteroclitus (PRESL) C.CHR. var.
eurybasis CHRIST, Philip. J. Sc. 2 (1907) Bot. 159. —
Campium heteroclitum (PRESL) COPEL. var. eury-
basis CopPEL. Philip. J. Sc. 37 (1928) 361.
Leptochilus heteroclitus (PRESL) C.CHR. var.
foxworthyi CuristT, Philip. J. Sc. 2 (1907) Bot.
160.
Leptochilus sumatranus v.A.V.R. Bull. Jard. Bot.
Btzg II, 23 (1916) 15, pl. 2: f. la-c; Handb. Mal.
Ferns, Suppl. 1 (1917) 436; ibid. Corrections (1917)
60.
Campium nigrum CoPEL. Philip. J. Sc. 37 (1928)
361, f. 14, pl. 10. — B. nigra CHING in C.Chr. Ind.
Fil. Suppl. 3 (1934) 49.
Campium pseudoscalpturatum Cope. Philip.
J. Sc. 37 (1928) 363, f. 16, pl. 11. — B. pseudoscalp-
turata CHING in C.Chr. Ind. Fil. Suppl. 3 (1934)
49; CopeEL. Fern FI. Philip. (1960) 261.
Campium foxworthyi CopPeL, Philip. J. Sc. 37
(1928) 364, f. 17, pl. 12. — B. foxworthyi CHING in
C.Chr. Ind. Fil. Suppl. 3 (1934) 48; CopeL. Fern
Fl. Philip. (1960) 258.
Campium tenuissimum CopPeL. Philip. J. Sc. 37
(1928) 364, f. 18, pl. 13. — B. tenuissima COPEL.
Fern FI. Philip. (1960) 259.
Leptochilus simplicifolius HoLtruM, Gard. Bull.
S. S. 4 (1929) 409. — B. simplicifolia CHING in
C.Chr. Ind. Fil. Suppl. 3 (1934) 50; HoLtrum,
Ferns Malaya (1954) 464, f. 272.
Campium membranaceum CopPEL. Philip. J. Sc. 40
(1929) 307, pl. 7. — B. membranacea C.Cnr. Ind.
Fil. Suppl. 3 (1934) 198; Cope. Fern FI. Philip.
(1960) 261.
B. edanyoi Core. Philip. J. Sc. 81 (1952) 22,
pl. 15; Fern FI. Philip. (1960) 258.
Edanyoa difformis CoPEL. Philip. J. Sc. 81 (1952)
22, pl. 17; Fern Fl. Philip. (1960) 265. — Fig. 25d,
3la-g.
Sterile fronds simple or pinnate, 15-100 cm long.
Pinnate fronds 25-100 cm long; lamina index of
non-flagelloid leaves 1-5, that of flagelloid leaves
sometimes more, 10-> 75 by 5-30 cm, terminal
segment to > 75 cm long, herbaceous, usually
greenish, sometimes purplish; pinnae 2-10(-15),
index 2—5S(—7), 1-26 by 0.7-6.5 cm, base attenuate
to truncate, margin usually + entire, sometimes
sinuate or sinuate-serrate or finely repand with-
out teeth of spines; terminal segment usually +
conform to the pinnae, sometimes flagelloid;
venation pattern: veins anastomosing into a +
regular pattern of variously large areoles, see
fig. 31g. Simple fronds 15-45 cm long; lamina
index 3-7(-17), 10-30 by (1-)2-7 cm, otherwise
similar to the terminal segment of pinnate fronds.
— Fertile fronds 14-75 cm long. Pinnate fronds
30-75 cm long; lamina index 1-4, pinnae index
2-7, 1-13 by 0.5-2.5(-4) cm. Simple fronds 14-40
(-> 45) cm long; lamina index 3-10, 4-13 by
0.5—2 cm. Sporangia inserted all over the lower
surface. Spores with a smooth, cristate or cristate-
undulate perispore. — Chromosomes n = 41,
2 — 626. 12st
Distr. E. India to S. Japan, Formosa, and Indo-
China; in Malesia: throughout, except for the
FLORA MALESIANA
[ser. Il, vow
Lesser Sunda Is.; also in Micronesia (Caroline Is.)
and Melanesia (Solomon Is.).
Ecol. On rocks, in soil or on bases of trees in
moist places in rain-forest (often near streams),
sometimes rheophytic, 0-1500(—1750) m.
Note. The morphological variation is outstand-
ing. Throughout its area forms with simple fronds
occur, e.g. B. simplicifolia from higher elevations
in Malaya. In the Philippines several distinct
vegetatively propagating dwarfs occur, e.g.
Edanyoa difformis and B. cuspidata. Autoploidisa-
tion and hybridisation are common. Field studies
are necessary to further unravel this aggregate.
8. Bolbitis sinuata (PRESL) HENNIPMAN, Blumea 18
(1970) 148; Leid. Bot. Ser. 2 (1977) 232, f. 61, 62.
— Polypodium ? sinuatum PRESL, Rel. Haenk. 1
(1825) 21.
Acrostichum diversifolium Bv. En. Pl. Jav. (1828)
103; Fl. Jav. Fil. (1829) 36, pl. 12. — B. diversifolia
ScHott, Gen. Fil. (1835) ad t. 13; BACKER &
PostH. Varenfl. Java (1939) 84; HoLtrum, Ferns
Malaya (1954) 465; Cope. Fern FI. Philip. (1960)
258. — Leptochilus diversifolius C.CHR. Ind. Fil.
(1906) 385; Curist, Philip. J. Sc. 2 (1907) Bot. 160,
quoad nomen solum; v.A.vV.R. Handb. Mal. Ferns
(1908) 740. — Campium diversifolium CopPeEL.
Philip. J. Sc. 37 (1928) 362, f. 15; BACKER & PosTH.
Nat. Tijd. N. I. 93 (1933) 164.
Gymnopteris subsimplex Fre, Hist. Acrost.
(1845) 83, pl. 40: f. 3. — Campium subsimplex
Cope. Philip. J. Sc. 37 (1928) 356, f. 11, pl. 8;
BACKER & PostTH. Nat. Tijd. N. I. 93 (1933) 159. —
B. subsimplex CuHiNG in C.Chr. Ind. Fil. Suppl. 3
(1934) 50; BACKER & PostTH. Varenfl. Java (1939)
81; CopeL. Fern FI. Philip. (1960) 265.
[Gymnopteris subrepanda J. SmitH, Hook. J.
Bot. 3 (1841) 403, nomen.] — Poecilopteris sub-
repanda PRESL, Epim. Bot. (1851) 171.
Anapausia zollingeri PRESL, Epim. Bot. (1851)
187. — Leptochilus zollingeri FEE, Gen. Polyp.
(1852) 55; v.A.v.R. Handb. Mal. Ferns (1908) 742,
p.p.; ibid. Suppl. (1917) 436. — Acrostichum
zollingeri KUNZE, Bot. Zeit. (1864) 419; CopeEL.
Philip. J. Sc. 37 (1928) 357. — Chrysodium
zollingeri KUHN, Ann. Mus. Bot. Lugd.-Bat. 4
(1869) 293.
Acrostichum variabile Hook. var. rasamalae
Racis. Pterid. Fl. Btzg (1898) 50.
Leptochilus hydrophyllus Copev. Philip. J. Sc. 1
(1906) Suppl. 146. — Campium hydrophyllum
CopEL. Philip. J. Sc. 37 (1928) 358, f. 12, pl. 9. —
B. hydrophylla CutnG in C.Chr. Ind. Fil. Suppl. 3
(1934) 48.
Leptochilus malaccensis C.CHR. Gard. Bull. S.
S. 4 (1929) 394. — B. malaccensis CHING in C.Chr.
Ind. Fil. Suppl. 3 (1934) 49; HoLttrum, Ferns
Malaya (1954) 470.
B. nitens HoLTTuM, Kew Bull. 13 (1958) 453;
Ferns Malaya ed. 2 (1966) 635. — Fig. 27j.
Sterile fronds simple or pinnate, (8—)20-100 cm
long. Pinnate fronds 35-100 cm long; lamina index
1-3, 25-60 by 8-40 cm, terminal segment 15-50 cm
long, usually (sub)coriaceous, sometimes sub-
carnose, olivaceous or brownish; pinnae 2-11,
index 3-10(-20), 5-35 by (0.5—-)1-7 cm, base
attenuate to truncate, margin entire or irregularly
sinuate especially in the lower half, without teeth
or spines throughout; terminal segment + conform
1978]
LOMARIOPSIS GROUP (Hennipman)
327
to the central pinnae; venation pattern: veins
anastomosing into a regular network of variously
sized areoles, see fig. 27j. Simple fronds (8—)20-
75 cm long; lamina index 3-10, (7-)20—-60 by
(1—-)3—-15 cm, otherwise like the terminal segment of
pinnate fronds. — Fertile fronds (8—)20—90 cm long.
Pinnate fronds 30-90 cm long; lamina index 1-4,
pinnae index 4~-10(—16), 5-17 by (0.5—)0.7-3 cm.
Simple fronds (8—)20-60 cm long; lamina index
5-18, (4-)8-35 by (0.3-)0.5-4 cm. Sporangia in-
serted all over the lower surface. Spores with a
smooth cristate perispore. — Chromosomes
n = 41, 2n = 122 + fragm.
Distr. India (Nicobar Is.) and Peninsular Thai-
land; in Malesia: Sumatra, Malay Peninsula,
W. Java, Borneo, Philippines, New Guinea (in W.
and NE. each one collection). Fig. 32.
Ecol. Terrestrial and low-epiphytic in evergreen
forest, 0O—1200(—1800) m.
Note. In the Malay Peninsula HoLTTUM recog-
nized two quite distinct forms, B. malaccensis and
B. nitens.
Fig. 32. Range of Bolbitis sinuata (PRESL) HENNIP-
MAN.
4. Series Quoyanae
HENNIPMAN, Leid. Bot. Ser. 2 (1977) 250.
Sterile fronds usually pinnate, rarely simple;
petiole with 3-16 vascular bundles; lamina with
one subterminal bulbil; pinnae up to 52, the margin
with or without spines; terminal segment triangu-
lar; venation pattern: veins regularly reticulate,
areoles angulate, + isodiametric or elongate,
decreasing in size towards the margin. Spores with
a smooth, cristate or cristate-undulate perispore. —
Chromosomes n = 41, 2n = 82, 123.
Distr. E. Malesia, Pacific, Japan (Bonin Is.).
9. Bolbitis quoyana (GAUDICH.) CHING in C.Chr.
Ind. Fil. Suppl. 3 (1934) 49; BAcKER & PosTH.
Varenfil. Java (1939) 81, p.p.; CopeL. Fern FI.
Philip. (1960) 264; HENNIPMAN, Leid. Bot. Ser.
2 (1977) 250, f. 69, 70. — Acrostichum quoyanum
GaAUDICH. in Freyc. Voy. Uranie (1827) 306, pl. 3;
Hook. Sp. Fil. 5 (1864) 259. — Gymnopteris
repanda (BL.) CHRIST var. guoyana (GAUDICH.)
Drets in K.Sch. & Laut. Fl. Schutzgeb. (1900) 117.
— Leptochilus cuspidatus (PRESL) C.CHR. var.
gquoyanus C.CHR. ex V.A.vV.R. Handb. Mal. Ferns
(1908) 742, p.p.; BRAUSE, Bot. Jahrb. 56 (1920) 117.
— Campium quoyanum CopeL. Philip. J. Sc. 37
(1928) 366, f. 20a, p.p.; BACKER & PosTH. Nat.
Tijd. N. I. 93 (1933) 161, p.p.
Heteroneuron naumannii KUHN, Forsch. Reise
Gazelle 4, Farne (1889) 5, pl. 1. — Gymnopteris
naumannii Diets in K.Sch. & Laut. Fl. Schutzgeb.
(1900) 117. — Leptochilus naumannii C.Cur. Ind.
Fil. (1906) 386; v.A.v.R. Handb. Mal. Ferns
(1908) 742; Brause, Bot. Jahrb. 56 (1920) 118. —
B. naumannii CuiNG in C.Chr. Ind. Fil. Suppl. 3
(1934) 49.
Leptochilus cuspidatus (PRESL) C.CHR. var.
marginalis ROSENST. in Fedde, Rep. 9 (1911) 426;
v.A.vV.R. Handb. Mal. Ferns Suppl. (1917) 435;
Brause, Bot. Jahrb. 56 (1920) 117.
Aspidium novo-pommeranicum BRAUSE ex RECH.
Denkschr. K. Ak. Wiss. M.-N. KI. Wien 89 (1914)
471, pl. 3: f. 8b.
Campium validum CoPEL. Philip. J. Sc. 37 (1928)
3690822. pb'5:
Campium parvum CopPEL. Philip. J. Sc. 37 (1928)
375, f. 28, pl. 21. — B. parva CutnG in C.Chr. Ind.
Fil. Suppl. 3 (1934) 49.
Campium enorme CoPEL. Philip. J. Sc. 40 (1929)
307, pl. 8. — B. enormis C.Cnr. Ind. Fil. Suppl. 3
(1934) 197; CopeEL. Fern FI. Philip. (1960) 264.
Campium viviparum KJELLB. in C.Chr. Bot.
Jahrb. 66 (1933) 50. — B. vivipara C.Cur. Ind. Fil.
Suppl. 3 (1934) 51.
Stenosemia dimorpha Copet. Philip. J. Sc. 84
(1955) 161, pl. 1.
[Cyrtogonium laciniatum J. SMitH, Hook. J. Bot.
3 (1841) 403, nomen.] — Fig. 31h-k.
Sterile fronds pinnate, up to 130 cm long;
lamina index 1—3(-4), up to 80 by 40 cm, terminal
segment 9-26 cm long, herbaceous to subcoria-
ceous, green or brown (or blackish); pinnae 13-52,
index 2-7, 7-22 by 1.3-3.5(-4.5) cm, base sym-
metrical, (narrowly to) broadly cuneate to cordate,
margin usually lobed +/,;*/, towards the costa,
sometimes either + entire or lobed to */; towards
the costa, with a usually rather inconspicuous spine
in the sinuses, apex acute to acuminate, lobes close
together to spaced, straight or subfalcate, margin
entire or crenate-serrate; terminal segment narrowly
triangular, shorter than the remaining part of the
lamina; venation pattern: veins forming a regular
network, areoles angulate isodiametric or elongate
decreasing in size towards the margin, see fig. 31j—k.
Fertile fronds up to 120 cm long; lamina index 3-6,
pinnae index 5—15, 3-15 by 0.4-1.7 cm. Sporangia
usually inserted all over the lower surface, some-
times situated along the margin only, the arrange-
ment usually acrostichoid, sometimes + pteridoid.
Spores with a smooth cristate perispore. —
Chromosomes n = 41, 2n = 82, 123.
Distr. Queensland, Polynesia (Samoa, Fiji),
Melanesia (Solomons), Micronesia (Bonin Is.); in
Malesia: New Guinea (common, also in the Bis-
marcks), Moluccas (P. Pisang, Halmaheira, Ter-
nate, Morotai), Philippines, Celebes, Central &
West Java. Fig. 33.
Ecol. On rocks and in soil in rain-forest, mostly
near streams: several times reported from lime-
stone; 0—1200(—1700) m.
Note. The species has been confused with
B. repanda. It is closest to B. rivularis, its coriaceous
counterpart (and also showing a tendency to
dwarfing), and to B. taylorii, an endemic from
Queensland.
328 FLORA MALESIANA [ser. IT, voloe
ase or 2046
; — “~
e - f as ' a
Sel é igh Pate |
“2 ss St ee & mr
con a <= ae 7A
s Bes * eS M3
eres =
ees =e
<r.
e Bolbitis quoyana
o B. rivularis
gees
wv
“2
Fig. 33. Range of Bolbitis quoyana (GAUDICH.) CHING and B. rivularis (BRACKENR.) CHING.
10. Bolbitis rivularis (BRACKENR.) CHING in C.Chr.
Ind. Fil. Suppl. 3 (1934) 50; BAcKER & POosTH.
Varenfl. Java (1939) 82 (‘B. ? rivulare’); HENNIP-
MAN, Leid. Bot. Ser. 2 (1977) 255, f. 70, 71. —
Cyrtogonium rivulare BRACKENR. in Wilkes, U.S.
Expl. Exp. 16 (1854) 85, pl. 11: f. 2. — Leptochilus
rivularis C.CuHR. Ind. Fil. (1906) 387. — Campium
rivulare CoPEL. Philip. J. Sc. 37 (1928) 373, f. 27,
pl. 20; BACKER & PosTH. Nat. Tijd. N. I. 93 (1933)
161.
Campium kajewskii CopeL. Philip. J. Sc. 60
(1936) 112, pl. 22.
Sterile fronds usually pinnate, rarely simple, 10—
70 cm long. Pinnate fronds 10-70 cm long; lamina
index 1—3(—5S), 7-40 by 3-25 cm, terminal segment
5-30 cm long, fleshy-coriaceous, olivaceous to
brownish; pinnae 2-11, index 1-5, 1.5-16 by
0.8-4.5 cm, base ++ symmetrical, cuneate to sub-
cordate, margin + entire to lobed to 1/,(—/s)
towards the costa, without or with but an incon-
spicuous tooth or spine in the sinuses; terminal
segment triangular, as long as to longer than the
remaining part of the lamina; venation pattern:
veins forming a regular network of angulate,
isodiametric or elongate areoles which decrease in
size towards the margin. Simple fronds 15-30 cm
long, lamina index 4~7, 12-23 by 2.5-4.5 cm, base
cuneate to subcordate, margin + entire to lobed to
?/; towards the costa, otherwise similar to the
terminal segment of pinnate fronds. — Fertile
fronds pinnate, 20-75 cm long; lamina index 1-4,
pinnae index 1-6, 1.2-9 by 0.3-1.5 cm. Sporangia
inserted all over the lower surface. Spores with a
suipen cristate perispore. —- Chromosomes
hi— ie
Distr. Polynesia (Fiji), Melanesia (Solomons,
New Hebrides); in Malesia: New Guinea (West:
2 collections; East). Fig. 33.
Ecol. Terrestrial and on rocks in moist places in
rain-forest; several times reported to grow near or
in streams, 0—-2000 m.
Note. The species is not rarely found fertile
with small leaves composed of a large terminal
segment and two small pinnae. Dwarfs grow
massed on rocks in rivers in New Guinea.
Species incertae sedis
11. Bolbitis novoguineensis HENNIPMAN, Leid. Bot.
Ser. 2 (1977) 270, f. 74k—n. — Fig. 311.
Sterile fronds pinnate, 9-21 cm long; lamina
index 3-5, 7-17 by 2-4 cm, terminal segment
1.5-4 cm long, firm-herbaceous (to subcarnose?),
olivaceous, with a + terminal bulbil; rachis with
a narrow wing throughout or in the upper half
only; pinnae > 16 to 30, index 1-3, 1-2.4 by
0.7-0.8 cm, base usually symmetrical, cuneate,
sometimes somewhat asymmetrical, its basiscopic
side cuneate, the acroscopic side either broadly
attenuate or with a basal acroscopic lobe, margin
(bi)crenate-serrate with distinct spines in the
sinuses; terminal segment usually narrowly trian-
gular, much shorter than the remaining part of the
lamina, the basal half with few lobes, tapering
towards the acute or short-flagelloid apex, some-
times the whole terminal segment flagelloid;
venation pattern: veins usually forming a costal
areole, sometimes locally free. Fertile fronds 15-
> 20 cm long; lamina index c. 10, pinnae index
4-6, 0.6-0.8 by c. 0.2 cm. Sporangia inserted all
over the lower surface. Spores with a smooth
cristate perispore.
Distr. Malesia: E. New Guinea (D’Entre-
casteaux Is.), 3 collections.
Ecol. Creeping on rocky banks of streams in
rain-forest, 250 and 900 m.
1978]
Note. A small thickening presumably represent-
ing a primordium of a bulbil is found terminally on
the costae.
12. Bolbitis repanda (BL.) ScHotr, Gen. Fil. (1835)
ad t. 13; HENNIPMAN, Leid. Bot. Ser. 2 (1977) 274,
f. 78. — Acrostichum repandum BL. En. PI. Jav.
(1828) 104; Fl. Jav. Fil. (1829) 39, pl. 14, 15, p.p. —
Campium repandum PRESL, Tent. Pterid. (1836) 239.
— Gymnopteris repanda CurRIsT, Farnkr. Erde
(1897) 50, p.p.; Diets in E. & P. Nat. Pfi. Fam. 1, 4
(1901) 201, p.p.; CopeL. Publ. Bur. Gov. Lab. 28
(1905) 43, p.p. — Fig. 27i.
Sterile fronds pinnate, 35—100(—120) cm long;
lamina index of non-flagelloid fronds 1-3, of
flagelloid fronds up to 7, 15—70(—100) by 6-30 cm,
terminal segment 8—60(—70) cm long, herbaceous-
pergamentaceous, green to blackish, with a
(primordium of a) spherical subterminal bulbil;
pinnae 8—20(—24), index (2-)3-6(-8), 4.5-22 by
1.5-4.5 cm, base + symmetrical, angustate to
broadly cuneate, margin lobed +/,—'/, towards the
costa, usually with a distinct tooth in each sinus,
lobes usually finely serrate-crenate, sometimes
entire; the two lowermost pinnae + conform to
the pinnae; terminal segment triangular; venation
pattern irregular, veins forming areoles varying in
shape and size, some of which with usually one,
mostly excurrent included free vein, see fig. 27i.
Fertile fronds 35-85 cm long; lamina index 2-5,
pinnae index 3-9, 1.5—7 by 0.4-1.5(—2) cm. Sporan-
gia inserted all over the lower surface. Spores with
a smooth cristate-undulate perispore. — Chromo-
somes n = 82, 2n = c. 120.
Distr. Micronesia (one record from the Maria-
nas); in Malesia: Philippines (Luzon, Mindanao),
Borneo (Sabah, Sarawak), Celebes, Lesser Sunda
Is. (Bali to Flores).
Ecol. Usually terrestrial,
epiphytic, in forest, 125-1650 m.
Note. Presumably of hybrid origin. The irregu-
lar venation pattern shows considerable variation
and includes all intermediates between a venation
pattern as found in B. sinensis or B. rhizophylla
(both ser. Egenolfianae) and one as found in
B. heteroclita (ser. Heteroclitae).
Hybrids
H.1. Bolbitis « arguta (FEE) CHING in C.Chr. Ind.
Fil. Suppl. 3 (1934) 47; Cope. Fern FI. Philip.
(1960) 263, p.p., p. spec.; HENNIPMAN, Leid. Bot.
Ser. 2 (1977) 288, f. 83i, j. — Heteroneuron
argutum FEE, Hist. Acrost. (1845) 96, pl. 25: f. 2,
p.p.; PRESL, Epim. Bot. (1851) 169. — Campium
argutum CopPEL. Philip. J. Sc. 37 (1928) 376, p.p.
Fronds small, pinnate throughout or the basal
part bipinnate, with a triangular terminal segment.
Sterile fronds: lamina with a subterminal bulbil;
pinnae irregularly and variously lobed, pinnae of
one pair sometimes much different, odd pinnae
present in part of the material. Fertile fronds:
sporangia inserted all over the lower surface.
Spores abnormal or sporangia with aborted spore-
mother-cells.
Distr. Malesia: Philippines (Luzon), known
from the type collection only.
Parentage. I have doubts whether this fern
warrants a separate treatment as a hybrid; it may
be close to Edanyoa difformis (= B. heteroclita).
sometimes low-
LOMARIOPSIS GROUP (Hennipman)
329
H. 2. Bolbitis heteroclita » rhizophylla HENNIP-
MAN, Leid. Bot. Ser. 2 (1977) 291, f. 83l-o.
Sterile fronds pinnate, with a subterminal bulbil;
rachis with or without narrow wing; pinnae 16~24,
index 3-5, 4.5-6 by 1.2-1.6 cm, base truncate to
cuneate, margin usually + entire, sometimes ir-
regularly and finely serrate or with a few incisions
about halfway towards the costa, with distinct
spines; terminal segment triangular, deeply lobed
near the base; venation pattern irregular, veins
forming a costal areole (rarely lacking), sometimes
also one or a few smaller distal areoles. Fertile
fronds: pinnae index 2-5, 1.64 by 0.6-0.8 cm.
Sporangia inserted mainly on the veins. Spores ab-
normal, or with aborted spore-mother-cells. —
Chromosomes 2n = 82; at meiosis univalents only.
Distr. Malesia: Philippines (Luzon, Mt Maquil-
ing, 2 collections).
Ecol. A shady place in forest, at 350-400 m.
H. 3. Bolbitis »~ singaporensis HoLtrum, Gard.
Bull. S. S. 11 (1947) 271; Ferns Malaya (1954) 467,
f. 274; HENNIPMAN, Leid. Bot. Ser. 2 (1977) 296,
f. 85a—d.
B. quoyana auct.
Ho.ttum, Gard. Bull. S.
Fig. 27n.
Sterile fronds pinnate, 35-70 cm long; lamina
index 1-3, 22-45 by 10-25 cm, terminal segment
10-19 cm long, firm-herbaceous, bright green to
olivaceous, with a subterminal bulbil; rachis not
winged; pinnae 14-27, index 3-5, 5.5-12.5 by
1.5—3 cm, base varying from + symmetrical, sub-
cordate, cuneate or angustate, to (strongly)
oblique with its acroscopic side much better
developed and provided with a distinct basal
acroscopic lobe (or auricle), margin entire to lobed
to 1/(/,) towards the costa, with a more or less
distinct spine in (some of) the sinuses; terminal
segment triangular; venation pattern very irregular:
veins forming a costal areole (rarely lacking), with
or without few to several smaller distal areoles
varying in size and shape, the areoles with or
without usually one excurrent included free vein,
see fig. 27n. Fertile fronds 50-70 cm long; pinnae
index 4-8, 2-6.5 by 0.4-1.2 cm. Sporangia inserted
mainly on the veins. Spores abnormal, or sporangia
with aborted spore-mother-cells. — Chromosomes
2n = 82; at meiosis univalents only.
Distr. Malesia: Malay Peninsula (Singapore,
Fern Valley on Bt Timah).
Ecol. On granite rocks in stream-bed, in the
shade of primitive forest.
Parentage. MANTON (in Holttum, 1954)
correctly suggested this to be a hybrid between
Egenolfia appendiculata (= B. appendiculata ssp.
appendiculata) and B. diversifolia (= B. sinuata).
These two species and the hybrid grow inter-
mingled in the Fern Valley on Bt Timah.
non (GAUDICH.) C.CHR.:
S. 9 (1937) 122. —
H. 4. Bolbitis * sinuosa (FEE) Cope. Fern FI.
Philip. (1960) 262; HENNIPMAN, Leid. Bot. Ser. 2
(1977) 298, f. 85e-p. — Heteroneuron sinuosum
FéeE, Hist. Acrost. (1845) 95; CopeL. Philip. J.
Sc. 37 (1928) 368, f. 21.
B. X foxii CopeL. Philip. J. Sc. 81 (1952) 22;
Fern FI. Philip. (1960) 263. — Fig. 27m.
Distr. Malesia: Philippines (Luzon, Mindanao,
Central Polillo).
330
FLORA MALESIANA
[ser. Tiyvalrui?
Parentage. B. quoyana and possibly B. rhizo-
phylla or B. heteroclita.
Two nothomorphs are distinguished.
a. nm. sinuosa HENNIPMAN, Leid. Bot. Ser. 2 (1977)
299, f. 85e, f.
Sterile fronds pinnate, up to 50 cm long; lamina
index 1-2, 16-32 by 10-18 cm, terminal segment
8-12 cm long, herbaceous, dark olivaceous or
brown, with or without a spherical subterminal
bulbil; pinnae 13-20, index 5-7, 7.5-12 by 1.1-
1.8 cm, base symmetrical, narrowly cuneate,
margin shallowly and oblique lobed or finely
crenate-serrate, with small teeth in the sinuses;
terminal segment triangular; venation pattern: veins
forming a costal areole and several smaller distal
ones, the areoles without included free veins.
Fertile fronds: pinnae c. 5 by 0.7-0.9 cm. Sporangia
inserted all over the lower surface. Spores ab-
normal or sporangia with aborted spore-mother-
cells.
Distr. Malesia: Philippines (Luzon, 3 localities).
b. nm. foxii (COPEL.) HENNIPMAN, Leid. Bot. Ser. 2
(1977) 299, f. 85g-p. — B. x foxii CopEL. Philip.
J. Sc. 81 (1952) 22.
If compared with nm. sinuosa the plants are of
the same size or smaller, with smaller and generally
also narrower pinnae with a somewhat simpler
venation pattern, see fig. 27m. — Chromosomes
2n = c. 80.
Distr. Malesia: Philippines (3 collections from
Polillo I., Luzon, and Mindanao).
Hybridae incertae sedis
H. 5. Leptochilus < trifidus v.A.v.R. Bull. Dép.
Agr. Ind. Neéerl. 18 (1908) 26; HENNIPMAN, Leid.
Bot. Ser. 2 (1977) 300, f. 85q—w.
Hemigramma latifolia auct. non COPEL.: COPEL.
Philip. J. Sc. 37 (1928) 404, p.p. — Fig. 271.
Sterile fronds simple, either entire or trifid,
30-60 cm long, firm-herbaceous to subcoriaceous,
brownish, with a small but conspicuous subter-
minal bulbil, base narrowly cuneate or angustate,
gradually or abruptly decurrent on the petiole,
margin entire or slightly sinuate; venation pattern
rather irregular, see fig. 271. Fertile fronds 25-50 cm
long. Sporangia inserted all over the lower surface.
Spores usually shrivelled, normally shaped spores
with a smooth, (imperfectly developed) cristate-
undulate or cristate perispore.
Distr. Malesia: Sumatra (3 localities in West
Coast, East Coast, and Bencoolen).
Ecol. Forest; reported from rocks either in
streams or on stream-banks, (one record) 450-
500 m.
Parentage. B. sinuata and possibly Leptochilus
decurrens. A very interesting hybrid which needs
experimental study.
Species dubiae
D. 1. Bolbitis interlineata (CopeL.) CHING in
C.Chr. Ind. Fil. Suppl. 3 (1934) 48; Ito, J. Jap.
Bot. 14 (1938) 439, quoad nomen solum; HENNIP-
MAN, Leid. Bot. Ser. 2 (1977) 306, f. 86g, h. —
Campium interlineatum CopeL. Philip. J. Sc. 37
(1928) 370, f. 24, pl. 17. — Fig. 27k.
Sterile fronds pinnate, 60-75 cm long; lamina
index 1-2, 30-40 by 20-24 cm, terminal segment
12-20 cm long, herbaceous, with a spherical sub-
terminal bulbil; pinnae 8-16, index 3-5, 13.5-15
by 3-4 cm, base either + symmetrical, broadly
cuneate to subcordate, or asymmetrical with its
basiscopic side longer and/or wider than its
acroscopic side, margin entire or (in part) slightly
sinuate; terminal segment + conform to the pinnae
or triangular with 1-2 basal lobes; venation
pattern very intricate, reminiscent of that of
B. heteroclita but part of the areoles with usually
one excurrent included free vein, see fig. 27k.
Fertile fronds 60-85 cm long; lamina index 2-4,
pinnae index 5-8, 5-6 by 0.7-1 cm. Sporangia all
over the lower surface. Spores with a smooth cris-
tate perispore.
Distr. Malesia: Borneo (Sarawak: Bungo
Range, Mt Matang, and Mt Penrissen).
Ecol. Forest; on rocks in streams, 300-690 m.
Note. Possibly of hybrid origin and related to
B. heteroclita (venation pattern!) and to either
B. scalpturata or B. repanda.
D. 2. Leptochilus stolonifer CuHrist, Bull. Herb.
Boiss. II, 6 (1906) 1004; v.A.v.R. Handb. Mal.
Ferns (1908) 739.
Distr. Malesia: Philippines (Angilog), type only.
Note. Possibly synonymous with B. heteroclita
but the venation pattern as given by CurisT is
deviating. Type not traced.
Excluded
Bolbitis ovata (COPEL.) CHING in C.Chr. Ind. Fil.
Suppl. 3 (1934) 49. — Leptochilus ovatus COPEL.
Philip. J. Sc. 9 (1914) Bot. 229. — Campium
ovatum CoPEL. Philip. J. Sc. 28 (1937) 354, f. 9, pl. 6.
— Paraleptochilus ovatus CopEL. Gen. Fil. (1947)
198. — Type: C. J. Brooks 155 S, Sumatra, Ben-
coolen, Lebong Tandai, ii-1913 (iso in BM) =
Colysis cf. pedunculata (Hook. & GREvV.) CHING.
Bolbitis subcrenata (Hook. & GREV.) CHING;
HENNIPMAN, Leid. Bot. Ser. 2 (1977) 176, f. 46, 47.
BACKER & PosTHUMUS (Varenfl, Java, 1939, 80;
Nat. Tijd. N. I. 93, 1933, 158, pro Campium sub-
crenatum) reported this fern from Java. The
material traced belongs to B. subcrenata var.
prolifera (endemic in Ceylon) and is presumably
cultivated.
THELYPTERIDACEAE (R. E. Holttum, Kew)
CHING, Sunyatsenia 5 (1940) 237-240, excl. Hypodematium KUNZE &
Parapolystichum CHING; Acta Phytotax. Sinica 8 (1963) 289-335, excl.
Hypodematium KUNZE; HorLttuM, J. Linn. Soc. Bot. 53 (1947) 130;
Blumea 19 (1971) 17-52; K. IwaTs. Mem. Coll. Sci. Univ. Kyoto B, 30
(1929) 21-51; 31 (1964) 1-40, (1965) 125-197; PICHI SERMOLLI, Webbia 24
(1970) 709 (first Latin description). — Fig. 1-20.
Caudex erect, short-creeping or long-creeping, rarely scandent; vascular
structure in all cases a radially symmetrical dictyostele; scales usually thin,
not peltate, in almost all cases bearing both marginal and superficial
unicellular hairs which are either acicular or glandular. Vascular strands at
base of stipe 2, linear in section (rarely with an additional pair of small
ones), uniting upwards to a U-shape; a linear aerophore with stomata
continuous along each side of stipe and rachis. Fronds usually pinnate with
crenate or lobed pinnae, in a few cases simple or bipinnate, never with
basiscopically enlarged basal pinnae; apical lamina usually triangular and
lobed, grading into upper pinnae, in some cases pinna-like; lower pinnae in
many cases gradually much reduced or with abrupt transition to a series of
small rudiments; a small aerophore, sometimes swollen or elongate, present
at the base of each pinna; a translucent membrane present in the base of
each sinus between adjacent pinna-lobes; venation in each pinna consisting
of a costa bearing costules, each costule bearing pinnately-arranged veins in
a pinna-lobe; veins free in deeply lobed pinnae, or basal veins in adjacent
lobes anastomosing to form an excurrent vein, which may be joined by
other veins, terminating at the base of a sinus-membrane, successive veins
passing to the sides of the sinus-membrane where this is elongate. In-
dument: scales always present at base of stipe, gradually smaller upwards,
minute (often consisting of a single row of cells) on the distal parts of
fronds, often nearly all caducous; adaxial surface of rachis and costae
always bearing antrorsely curved acicular unicellular hairs, in a few cases
also septate acicular hairs; abaxial surface of rachis and costae usually
bearing a different indument consisting of more slender unicellular acicular
and/or glandular hairs or sessile glands of various forms (forked hairs in
Ampelopteris only); surface of lamina between veins either quite glabrous
or more often with a distinctive complement of hairs and glands different
adaxially and abaxially. Sori borne on abaxial surface of veins, orbicular or
sometimes elongate, indusiate or not; indusia reniform, glabrous or bearing
hairs and/or glands, in some cases very small, athyrioid in some species of
Coryphopteris; sporangia sometimes bearing glands or short acicular hairs
(setae) near annulus, often with a hair of distinctive form on the sporan-
gium-stalk; spores in almost all cases monolete, with perispore of varied
form, in Trigonospora trilete. Gametophyte in all cases symmetrical-cor-
date, with unicellular chlorophyllous hairs on all parts, these hairs with
+ swollen rounded tips which become wax-encrusted; in most cases, usu-
ally as a late development, unicellular acicular hairs, comparable with those
on the sporophyte, may occur; other types of hair may be distinctive of
some genera.
(331)
5
332 FLORA MALESIANA [ser. II, vol. 1
—
—
\ Sh
Fig. 1. Sphaerostephanos penniger (HOOK.) HOLTTUM. a. Whole frond, x 4; b. areduced pinna showing
linear and round aerophores, = 2; c. CS base of stipe, x 3; d. CS rachis, x 3; e. scale, x 6; f. rachis and
base of pinna, upper surface, x 13; g. part of pinna, upper surface, showing unicellular hairs and glands,
x 12.—Venation patterns in relation to sinus-membrane: h. Sphaerostephanos penniger (HOOK.)
Ho.ittuM, X2; i. Christella arida (D. DON) HoLTTUM, 6: j. Plesioneuron fulgens (BRAUSE)
HoLttuM, x3; k. Mesophlebion crassifolium (BL.) HOLTTUM, 4; |. Coryphopteris viscosa (BAK.)
HoLTTUM, X4; m. Metathelypteris dayi (BEDD.) HOLTTUM, x 3.— Sporangia, x 33: n. Pronephrium
repandum (FEE) HOLTTUM; o. P. rubicundum (v.A.v.R.) HOLTTUM; p. Christella dentata (FORSSK.)
BROWNSEY & JERMY; q. Sphaerostephanos heterocarpus (BL.) HOLTTUM; r. Cyclosorus interruptus
(WILLD.) H. ITo.
1981] THELYPTERIDACEAE (Holttum) 333
Distribution. Throughout the tropics, especially in wetter areas; species few in tem-
perate regions (5 in Europe), almost 1000 in all. The majority are terrestrial ferns of forest,
but a few (especially in Christella and Macrothelypteris) occur in open places only, and a
few (Cyclosorus, Thelypteris) in open swamps; some are adapted to grow on rocks by
streams; very few are scandent; a few are casually epiphytic.
Fossils. Fossil ferns with veins anastomosing as in this family have been found in Lower
Tertiary strata in both Europe and America, but in the absence of information concerning
scales, hairs, glands and spores the affinity of such fossils to any existing groups cannot be
definitely assessed; the oldest fossil which probably belongs to this family is Aspidistes
thomasii HARRIS (Yorkshire Jurassic Flora 1, 1951, 181) which has bipinnate or tripinnate
fronds with free veins, abundant superficial glands much as in Coryphopteris and trilete
spores resembling those of Trigonospora.
Vegetative morphology. A combination of two characters is distinctive of the family:
the arrangement of vascular strands in the stipe (Fig. 1c—d) and the presence of unicellular
acicular hairs on the adaxial surface of rachis and costae (Fig. 1g). The only other fern
with this combination is Hypodematium, which differs greatly in frond-form and in
scales. Also distinctive of Thelypteridaceae is the frond-form of most species: simply
pinnate fronds with most pinnae almost equal in length, each pinna symmetrical on either
side of the costa with veins symmetrically arranged in the pinna-lobes. The venation in its
relation to sinus-membranes is also unique among ferns.
Distinctions between species are provided by details of venation and of the distribution of
hairs and glands (more rarely of scales) on the two surfaces, the abaxial surface in general
providing the greater diversity. Maximum size of fronds is also distinctive in many cases, but
often plants may be fertile at an immature size.
Dimorphism. In some species sterile fronds have much larger (both longer and broader)
pinnae than fertile ones, and in such species transitional forms may occur. In other species
there is some irregular dimorphism. In general, sterile fronds often have longer hairs than
fertile ones on the lower surface, sometimes much longer ones.
In general the number of veins anastomosing is dependent on the depth of lobing of
pinnae, but this situation is complicated by the relative length of the sinus-membrane (Fig.
lh-m). Where the membrane is very short and pinnae shallowly lobed, anastomosing veins
from adjacent costules form a zig-zag excurrent vein and only one vein may end in contact
with the membrane, but if the membrane is long (forming a prominent fold on the lower
surface) only one pair of veins may be united, successive ones passing to the sides of the
membrane as if to the edge of a pinna-lobe. These details need to be carefully observed. In
rare cases a long membrane may be present with no veins actually united. In other cases
(especially on Pneumatopteris) a vein passing to the side of a membrane may be continued
to join with the vein next above it, thus forming a continuous vein on one side of the
membrane and sometimes extending into the pinna-lobe. CHING and IWATSUKI both state
that a character of Abacopteris FEE (Pronephrium of the present work) is the absence of a
sinus-membrane, but such a condition is not invariable even in pinnae which have almost
entire margins.
The species with bipinnate fronds have all, or most, of the pinnules adnate to a
pinna-rachis (Fig. 2h); this is a contrast to Cyathea, with which genus Thelypteridaceae have
some features in common, and I suggest that it is a secondary development (see below on
relationships with other groups). These bipinnate species (in Macrothelypteris and Pseudo-
phegopteris) also have veins terminating within the margin (Fig. Im) and adaxial surface of
rachises and costae not grooved, both conditions shared by Metathelypteris and Phegopteris;
in all other genera costae are grooved (Fig. 1f-g) and all free veins terminate at the margin.
In some species of several genera the aerophores at pinna-bases are swollen or elongate.
CHRISTENSEN united several such tropical American species in a subgenus Glaphyropteris,
and some species of Asia were also so placed by ITO in Nakai & Honda, Nov. FI. Japonica,
part 4 (1939), but CHING (1963) recognized that the species so named by ITO were neither
closely related to each other nor to the American species of CHRISTENSEN.
In most species (apart from Pronephrium) the apical lamina of a frond is narrowly to
334 FLORA MALESIANA (ser. II, vol. 1°
broadly triangular and +deeply lobed, with a gradual transition at its base to the upper
pinnae. The distal part of such an apical lamina has veins arranged in its lobes as in the lobes
of a pinna, but near the base there is a transition from lobes to pinnae, with a transition also
in venation. This transition in venation (especially when the terminal lamina is long and
narrow) sometimes results in a condition comparable to that of normal pinnae or pinnules of
Pleocnemia (a genus allied to Tectaria). In a few cases this peculiar transitional type of
venation also occurs above the base of the apical lamina, and such a case was made the basis
of a genus Haplodictyum by PRESL; some later authors, including CHING, thought that this
venation indicated a real relationship of Haplodictyum to Tectaria, but certainly the former
is a true thelypteroid fern. A fossil from America has also been reported with venation of
Haplodictyum type (Goniopteris claiborniana BERRY, Bull. Torr. Bot. Club 44, 1917, 331,
t222)
Much-reduced pinnae at the base of a frond, usually with an abrupt transition to normal
pinnae, occur in the large genera Sphaerostephanos (Fig. la—b) and Pneumatopteris; such a
condition is rare in American Thelypteridaceae, though in Amauropelta lower pinnae are in
most cases gradually much reduced. In young living fronds such reduced pinnae are seen to
have more or less swollen white aerophores at their bases; in some cases the aerophore is
larger than the lamina of the reduced pinna. The number, shape, and spacing of reduced
pinnae provide useful distinguishing characters, but unfortunately some dried specimens do
not show them fully and in other cases the frond-base is quite lacking. Where very young
fronds are covered with mucilage the aerophores are sufficiently elongate for their tips to be
free from the mucilage. Swollen aerophores (rarely much elongate) occur also in some
genera where basal pinnae are not reduced.
Sori, sporangia and spores. The position of sori on veins is characteristic in each
species; in Chingia sori are always near costules but within most genera there is much
variation. More or less elongate sori occur in several genera, especially in Stegnogramma (all
species) and Sphaerostephanos, but within the latter genus the species with elongate sori do
not form a natural group. In some species where indusia are normally very small they are
sometimes quite lacking, but in general size and indument of indusia are characteristic. In
Coryphopteris distal sori sometimes have athyrioid indusia, basal ones rarely.
The form of sporangia and the presence of glands or hairs on them are always important
diagnostically (Fig. In-r) but these characters are often unrecorded in original descriptions,
even of recent publication. I have tried to obtain exact information from type specimens. In
Christella, Coryphopteris and Metathelypteris there are neither glands nor hairs near the
annulus (but see note under Christella subpubescens). Hooked hairs occur on sporangia in
Pronephrium sect. Grypothrix (Fig. 1o) and in Cyclogramma. The sporangia of Coryphopteris
are almost sessile but in most other members of the family they have slender stalks on some
of which are hairs; the nature of these hairs is also distinctive (Fig. 1p-r) but I have not been
able to report it for all species.
Spores of all species are covered with a perispore which may have many different external
forms (see C. C. WOOD, J. Linn. Soc. Bot. 67, Suppl. I, 1973, 191-202, pl. 1-4); this has only
been well observed since the introduction of the scanning electron microscope. The inner
structure of spore-walls can only be seen by sectioning and has still been little investigated.
LUGARDON has made sections of a representative set of spores; a report on this work will
be published separately. It gives some support to the present arrangement but leaves many
questions unanswered.
Gametophyte. Apart from the development in some species of acicular hairs, the
general characters of gametophytes do not distinguish this family from others; the glandular
hairs universally present occur also in other families. Reports have been published by L. R.
ATKINSON on species of various genera in the Old World, summarizing also those of earlier
authors (Phytomorphology 25, 1975, 38-54) and on Chingia (Fern Gaz. 11, 1975, 87-93).
Gametophytes of Mesophlebion crassifolium have distinctive branched hairs and also
spherical red glands resembling those on the stalks of sporangia of sporophytes. Gameto-
phytes of Chingia sakayensis have distinctive multicellular hairs. Gametophytes of other
genera show differences in the colour of glandular hairs or other less easily specified
1981] THELYPTERIDACEAE (Holttum) 335
characters. Gametophyte characters support the separation of the genera Plesioneuron and
Mesophlebion which I formerly united (HOLTTUM 1971), but in general do not offer much
important evidence as to the separation of, or inter-relations between, genera, partly because
few species in most genera (as here recognized) have been examined.
Cytology. The first reliable reports on chromosomes were by I. MANTON in her book of
1950. Subsequently many reports have been published, as summarized by LOVE, LOVE and
PICHI SERMOLLI in their Cytotaxonomical Atlas of the Pteridophyta (1977). Haploid
numbers range from 27 to 36; several species are tetraploid and of some both diploid and
tetraploid forms have been reported, in the wide-ranging species Macrothelypteris torresiana
tetraploid and hexaploid. Only a small proportion of species have so far been examined, so
that generalizations cannot usefully be made. In all recorded cases species belonging to one
genus, as here recognized, have the same chromosome number except that both n= 35 and
n= 36 occur in the genus Pseudocyclosorus in India (there is only one Malesian species).
Hybridization experiments have only been reported by MANTON and her co-workers in the
genus Christella q.v. for details.
Taxonomy. The distinguishing characters of this family were not recognized until
CHRISTENSEN’s study of tropical American species in 1913 (see below); in the 19th century
there was much confusion in the use of generic names which were based mainly on soral
characters.
LINNAEUS placed the species known to him in Polypodium or Acrostichum; he did not use
characters of indusia in distinguishing genera. The name Thelypteris was published by
SCHMIDEL in 1763; it has been conserved against Thelypteris ADANSON (= Pteris) which
may have been published earlier in the same year. In 1791 SCHREBER proposed the name
Meniscium for a species from tropical America having elongate exindusiate sori resembling
those of the Malesian species here named Pronephrium triphyllum, which was transferred to
Meniscium by SWARTZ in 1801; other authors followed this assignment but CHRISTENSEN
stated his belief that the Malesian and American species in question are not closely related;
see further discussion under Pronephrium.
In 1800 ROTH published the name Polystichum for ferns with either peltate or reniform
indusia; these included some Thelypteridaceae in addition to Polystichum aculeatum
(generic type) and species of Dryopteris and Gymnocarpium. GAUDICHAUD later included
some tropical Thelypteridaceae in Polystichum.
In 1801 SWARTZ published Aspidium, with almost the same diagnosis as Polystichum
ROTH, and included in it, besides Thelypteridaceae of several genera here recognized,
species now placed in Tectaria, Oleandra, Nephrolepis, Didymochlaena, Polystichum, Dry-
opteris, Cystopteris and Athyrium. Also in 1801 CAVANILLES published the name Tectaria
with a single species (now recognized as type of a non-Thelypteroid genus) but he later
included in it a mixture similar to that in Aspidium Sw. In 1803 MICHAUX published
Nephrodium, which also included species of many genera now regarded as distinct; see
below for confused later usage of this name.
In 1824 BORY published the name Lastrea for “la plus grande partie des Polypodes a
feuilles bipinnatifides ou bipinnées”’, citing a few species only; he did not refer to Aspidium
or Nephrodium, though the species he cited could have been placed in either. In 1833 LINK
published Cyclosorus for the single species which he named C. gongylodes; this had
previously been included in Aspidium, Nephrodium and Polystichum. BLUME in 1828
included all indusiate Thelypteridaceae in Aspidium, exindusiate ones in Meniscium, Gym-
nogramme and Polypodium; he established Stegnogramma for a species with elongate
exindusiate sori and anastomosing veins.
SCHOTT’s ‘Genera Filicum’, published in 1834, contained twenty beautifully engraved
plates showing details of as many genera; these included a plate illustrating Nephrodium
which showed for the first time with great exactness the unicellular acicular hairs charac-
teristic of all Thelypteridaceae (also capitate hairs) and other details now considered
significant; he also mentioned the vascular strands in the stipe. His description of the genus
and list of species shows that he restricted it to thelypteroid ferns having anastomosing veins
and round indusiate sori, none of which were in the original list published by MICHAUX (to
336 FLORA MALESIANA [ser. II, vol. 1°
which he did not refer). SCHOTT also recognized Thelypteris SCHMIDEL as an allied genus
with free veins.
In 1836 appeared PRESL’s ‘Tentamen Pteridographiae’, containing a completely new
system of fern genera. He adopted SCHOTT’s definition of Nephrodium and included most
free-veined thelypteroids in Lastrea. But in Lastrea he also placed species of Dryopteris,
Ctenitis and several other genera now recognized. He retained exindusiate free-veined
Thelypteridaceae in Polypodium sect. Phegopteris (with much admixture of non-thelyp-
teroids), those with anastomosing veins of the Nephrodium pattern in Goniopteris and those
with elongate sori in Grammitis. In ‘Epimeliae Botanicae’ (1851) PRESL proposed the new
genera Haplodictyum, Physematium, Proferea and Pronephrium for thelypteroid ferns from
the Philippines and Java.
W. J. HOOKER devoted the last twenty years of his life to producing ‘Species Filicum’
(five volumes, 1844-1864). He united all indusiate Thelypteridaceae in Nephrodium, and with
them Dryopteris, Ctenitis and Pleocnemia; he followed PRESL in placing exindusiate species
in Polypodium, but those with elongate sori in Gymnogramme and Grammitis.
Simultaneously with HOOKER’s work, A. L. A. FEE produced his ‘Genera Filicum’ (1852),
in which his arrangement is more elaborate. He placed the majority of Thelypteridaceae in
Polypodiaceae, tribe Aspidieae. Free-veined indusiate species, mixed with others now
allocated to Dryopteris and Ctenitis, are placed in Aspidium; species with anastomosing
veins and indusiate sori in Nephrodium, Haplodictyum and Abacopteris ; species with round
exindusiate sori in tribe Polypodieae, genera Phegopteris and Goniopteris; species with
elongate sori in tribe Meniscieae and tribe Leptogrammeae (with admixture of species now
referred to very different groups).
G. METTENIUS, in his ‘Fil. Hort. Bot. Lipsiensis’ (1856) included both indusiate and
exindusiate thelypteroids with free or anastomosing veins in his “tribe” Aspidiaceae,
separating all exindusiate species as Phegopteris, the rest in Aspidium. In his monograph of
Phegopteris and Aspidium (1858) he had many non-thelypteroid species in both genera; but
in his descriptions he noted characters of hairs, glands and scales with much more care than
any previous author except SCHOTT.
JOHN SMITH, who observed more than 1000 species of ferns as cultivated plants,
published another classification in ‘Historia Filicum’ (1875) and in a long introduction
commented on previous schemes. He placed almost all thelypteroids in his tribe Phegop-
teridiae, separating indusiate and exindusiate species (the former as Lastrea and Neph-
rodium) in much the same way as METTENIUS, but noting that indusia are sometimes very
small or fugacious so that separating on this character was often doubtful. Among free-
veined ferns named Lastrea he had representatives of other genera (as now recognized) but
he subdivided Lastrea to show some of these differences, as METTENIUS had not done.
R. H. BEDDOME, studying ferns in the field in southern India from 1856 to 1882, used
HOOKER’S classification but recognized some of its unsatisfactory features. In his Hand-
book (1883) and its Supplement (1892) he included also species of the Malay Peninsula, and
owing to a lack of critical study of Malesian ferns he sometimes misidentified Indian species
with those of Java.
Field work on ferns in Java was undertaken by M. RACIBORSKI in 1897-1898, and the
results were published in ‘Flore de Buitenzorg, I, Pteridophytes’ (1898). This includes much
previously unrecorded ecological observation. RACIBORSKI placed free-veined Thelyp-
teridaceae in Aspidium (with species of Dryopteris and Ctenitis) and those with anastomos-
ing veins in Nephrodium.
H. CHRIST in 1897 attempted a new survey of all ferns (‘Die Farnkrauter der Erde’). He
followed METTENIUS in placing almost all thelypteroid ferns in Aspidium and Phegopteris.
Subsequently he adopted CHRISTENSEN’s concept of Dryopteris (see below) and in 1907
his survey of Dryopteris in the Philippines was published; his work on the present family
was uncritical.
L. DIELS compiled a summary of Polypodiaceae for Engler & Prantl’s ‘Natiirlichen
Pflanzenfamilien’ (I, Abt. 4, 1899). He included all thelypteroid ferns in his tribe Aspidieae,
almost all of them in Nephrodium, uniting indusiate and exindusiate species in one genus for
1981] THELYPTERIDACEAE (Holttum) B57
the first time. He made an elaborate subdivision of Nephrodium, with free or anastomosing
veins aS a main distinction (vascular anatomy, scales, hairs and glands had no part in his
system) and did not clearly separate thelypteroid ferns from the rest.
CARL CHRISTENSEN based his ‘Index Filicum’ (1905) mainly on DIELS but (following
OTTO KUNTZE) he adopted the older name Dryopteris ADANSON (1763) in place of
Nephrodium; under it he had the mixture as before. VAN ALDERWERELT VAN ROSEN-
BURGH made a compilation of all recorded taxonomic information on Malesian ferns, with
new descriptive data for many species, adopting CHRISTENSEN’s scheme (except that he
reverted to Phegopteris for exindusiate thelypteroids); this was published at Bogor in 1908.
In subsequent years he published new descriptions of many species, his later observations
being more detailed and critical, but in many cases he misapplied older names. C. A.
BACKER & O. POSTHUMUS, in ‘Varenflora voor Java’ (1939) also adopted CHRISTENSEN’S
comprehensive Dryopteris, without attempting to separate thelypteroid species. Their
work includes much new field observation but their citation of synonymy was often
uncritical.
Having completed his Index, CHRISTENSEN realized that there was much mixture of
unrelated species in his Dryopteris and he proceeded to make a detailed study of tropical
American species. In the course of this work he distinguished thelypteroid ferns from the
rest for the first time (Monograph of the genus Dryopteris, Dansk Selsk. Skr. VII, 10, 1913,
55-282; VIII, 6, 1920, 1-132), but he retained all in Dryopteris, placing the thelypteroids in
subgenera, because he had not studied the Old World species and could not foresee how to
fit them into his scheme. In subsequent years CHRISTENSEN made many studies of ferns of
the Old World, adding to his knowledge, and in 1929-1932 R. C. CHING worked with him,
specializing on the ferns of China and India. CHING wrote an important series of papers in
the decade 1930-1940, including monographic treatment of Thelypteridaceae in mainland
Asia, in the genera Thelypteris, Cyclosorus, Abacopteris and Leptogramma. In 1940 he
recognized for the first time a family Thelypteridaceae. | accepted CHING’s scheme, with
reservations, in my book ‘Ferns of Malaya’ (1955).
COPELAND’s work on Philippine ferns began in 1904, soon became extended to cover
those of neighbouring regions, and culminated in his ‘Genera Filicum’ (1947) and ‘Fern Flora
of the Philippines’ (1960). In the main, he accepted CHING’s genera, but substituted Lastrea
for Thelypteris which he regarded as illegitimate; he did not recognize a family Thelyp-
teridaceae, and separated Thelypteris from Cyclosorus solely on the character of free or
anastomosing venation; he regarded Lastrea as closely related to Athyrium.
In 1963 CHING elaborated his scheme of classification of Thelypteridaceae in mainland
Asia, recognizing some new genera. K. IWATSUKI made a detailed study of morphology in
the family, mainly in Japan and China, and published a new taxonomic survey in 1964-1965,
recognizing the three genera Stegnogramma, Thelypteris (with many subdivisions) and
Meniscium.
I began a study of all Old World species of the family in 1967, examining the types of
almost all species and the complete collections in several major herbaria. I discovered that
Malesian species are far more varied than those of mainland Asia, and I thus had a wider
field of study than CHING and IWATSUKI. I devised a new scheme of genera which was
published in 1971, and subsequently monographs of all Old World species of the major genera
except Sphaerostephanos which is almost entirely Malesian and is here treated fully for the
first time.
My first conclusion (at which I had already hinted in my book of 1955 on the ferns of
Malaya) was that a division between Thelypteris and Cyclosorus based on free or united
veins was not a natural one. I also concluded that the nature and distribution of glands and
hairs provided important evidence which had never been well recorded. In such a perspec-
tive Thelypteris and Cyclosorus, restricted to their type species and near allies, are small and
distinctive groups not closely allied to most of the species associated with them by previous
authors. There thus appeared to be two alternatives: to include all species of the family in
one genus Thelypteris, or to recognize a number of separate genera. I chose the latter
because there is such a great diversity in Malesia that within a single genus, to be intelligible,
338 FLORA MALESIANA [ser. II, vol. 1°
one would always have to specify a subgeneric or sectional name when citing a species; a
specific name alone would not give sufficient information.
In the New World are some other distinctive groups, one of which (Amauropelta) extends
across Africa to the Mascarene Islands (Holttum, J. S. Afr. Bot. 40, 1974, 130), doubtfully to
Ceylon and not to Malesia. The New World genus Meniscium is mentioned under the genus
Pronephrium in the present work. The species of islands in the Pacific Ocean eastwards
from New Guinea (about 100) are almost all common to Malesia or related to Malesian
species (Allertonia 1, pt 3, 1977).
The result of this long history of confusion is that all names published in the 19th century
have been transferred, by one author or another, to Aspidium, Nephrodium and Dryopteris,
in some cases also to Goniopteris, Lastrea, Meniscium or Phegopteris; and in the 20th
century most names have been transferred to Thelypteris or Cyclosorus or both. In view of
the preference by some people for a comprehensive genus Thelypteris, I have cited names in
that genus if such have been published, and I have tried to avoid the use of new specific
epithets where such would have to be changed on transfer to Thelypteris.
Few species were originally described in terms which distinguish them clearly from others,
with the result that names were often mis-used or new ones needlessly created. Few
collectors understood how to distinguish between species of this family and the need for
careful preservation of the base of a frond; much herbarium material is therefore in some
measure unsatisfactory and new collections by specialists are still needed. Too many species
(my own included) are based on a single collection which may not adequately show possible
variation; but in many other cases repeated collections show a degree of uniformity which
confirms their status. It cannot be doubted that the number of species in Malesia is very
large, and that probably more remain to be discovered.
Relationship to other groups. In Journ. Linn. Soc. Bot. 53 (1947) 130-133 I commented on
resemblances between Thelypteridaceae and Cyathea, and listed them more fully in Blumea
19 (1971) 18-19: shape of fronds; shape of leaflets; abundant hairs on adaxial surfaces of
rachis quite distinct in nature from scales; arrangement of aerating tissue with a separate
small area at the base of each pinna (this area sometimes swollen both in Cyathea and in
Thelypteridaceae); relationship of sori to veins. As Cyathea has invariably deeply lobed
leaflets and free veins, there is not scope for development of a sinus-membrane, but there is
a rudiment of it, especially in Cyathea subg. Sphaeropteris sect. Schizocaena of my
arrangement in Flora Malesiana. Cyathea has a base chromosome number 69, about double
that of members of Thelypteridaceae; Cyathea has also a much more complex vascular
system. In some species of Cyathea the lower pinnae are gradually reduced, in others there
is a long stipe and the lowest pinnae are not much reduced; in Thelypteridaceae also both
these conditions occur.
Most species of Cyathea have amply bipinnate fronds, but young plants of all of them
have simply pinnate fronds. I suggest that the evolution of Thelypteridaceae began by the
persistence of the simply pinnate form in an adult condition (neoteny). I further suggest that
those extant Thelypteridaceae which have bipinnate fronds are a secondary development
from simply pinnate ancestors. Their pinnules are always adnate to the pinna-rachis in a way
not found in Cyathea.
Cyathea fronds have free veins, and it is likely that this was a character also of early
Thelypteridaceae. But it is probable that the distinctive pattern of anastomosis of veins, with
its precise relationship to sinus-membranes (a pattern found in no other ferns) originated
early. In some of the genera here recognized it seems probable that the species with free
veins are not primitive in their own genera; for example, in Sphaerostephanos (the largest
genus in Malesia) the great majority of species have anastomosing veins, the few species
which have free veins not forming a closely allied group nor limited to one area. The same is
true in Pneumatopteris and Christella. But there are other genera which perhaps evolved
directly from free-veined ancestors (e.g. Coryphopteris and Trigonospora).
There is another group of terrestrial ferns which agree with Thelypteridaceae in having
abundant distinctive hairs, not homologous with scales, on the adaxial surface of frond-axes,
namely Ctenitis, Tectaria and allied genera. The latter differ from Thelypteridaceae in
1981] THELYPTERIDACEAE (Holttum) 339
vascular anatomy of stipes, shape of fronds (pinnae often asymmetric at their bases, basal
pinnae enlarged basiscopically), have a different kind of anastomosis of veins, lack sinus-
membranes, and the hairs on the adaxial surface of frond-axes are always multicellular with
short cells, not acicular in form. I see no close alliance between Thelypteridaceae and other
groups, but point out the similarity of sori to those of Tectaria, Dryopteris and other genera,
which seems to me to imply a common ancestry from ancient tree-ferns (see Journ. Linn.
Soc. Bot. 67, Suppl., 1973, 5).
In his recent scheme of classification of ferns (Webbia 31, 1977, 313-512), PICHI
SERMOLLI places Thelypteridaceae in Aspidiales, an Order which he regards as related
more nearly to Dennstaedtiales than to Dicksoniales. Within Aspidiales he regards Thelyp-
teridaceae as nearest to Aspleniaceae, but in my view these two families have little in
common except the chromosome number 36 in a majority of species and should not be
placed in the same Order. PICHI SERMOLLI regards a creeping rhizome as_ probably
primitive in Thelypteridaceae (p. 441) and (ignoring the genus Trigonospora) states that
difference in spores is an argument against a relationship to Cyathea (p. 437); he does not
mention the various similarities to Cyathea indicated above. After reading his statement, I
cannot agree with his arrangement, either in reference to Thelypteridaceae as a whole or as
to the interrelations of genera within it. As regards the latter, he offers no justification for the
subdivision proposed on pp. 440-445; for example, he places Thelypteris and Cyclosorus
wide apart, without explanation, whereas they seem to me to be rather nearly related.
I assume an erect caudex to be primitive in Thelypteridaceae partly because I regard a
common origin with Cyathea as probable, and partly because the caudex in all species has
the radial structure characteristic of an erect habit; in none of the species with a creeping
rhizome has a dorsiventral structure (of the kind occurring in Dennstaedtia) developed. All
species of Thelypteridaceae which have long creeping rhizomes appear to me to be more
specialized than those with an erect caudex; nearly all of the former are plants of open
places, whereas the great majority of members of the family are adapted to a forest habitat.
Cyclosorus and Thelypteris (in the strict sense here adopted) are ferns of freshwater swamps
in sunny places. Sphaerostephanos unitus and S. invisus also grow in the open, usually in
less wet places, and are among the very few species in that large and varied genus which
have a long-creeping rhizome.
I agree that, in evolution, the megaphyll of ferns may have been derived from a
branch-system arising from a creeping rhizome, but if so that stage occurred very early,
before any groups now known originated. In 1949 the idea was in my mind that living
Gleicheniaceae, with their long-creeping rhizomes, might preserve an original plant-form, and
that, as suggested by BOWER, Cyathea might have originated from a Gleichenia-like
ancestor (Biol. Reviews 24: 268). But after making a survey of the tree-ferns in the present
Series (vol. 1, part 2) I concluded that the erect habit was probably primitive for Cyathea,
and that the plant-form (though not the form of sporangia) of modern Gleicheniaceae is
probably derivative and not primitive (there is no evidence of its existence before the
Cretaceous); I have discussed this theme in Phytomorphology 14 (1965) 477-480.
CONSPECTUS OF MALESIAN GENERA
1. Upper surface of costae not grooved; veins not reaching margin (except distal ones in Phegopteris)
Group of PHEGOPTERIS
2. Fronds bipinnate; n= 31.
3. Scales thin; perispore forming a reticulum of few meshes separated by low broad ridges
1. Pseudophegopteris
3. Scales thickened at base; perispore minutely complex . .. . . . . . 2.™Macrothelypteris
2. Fronds simply pinnate; n= 30 or 35.
4. Basal scales with short superficial setae; pinnae free; n=35 . . . . . .3.Metathelypteris
4. Basal scales flat with marginal hairs; pinnae with semicircular rachis- -wings between them; n= 30
4. Phegopteris
1. Upper surface of costae grooved; veins all reaching margin.
5. n= 33 or less; sporangia sessile, lacking glands or hairs . . . . . Group of CORYPHOPTERIS
340 FLORA MALESIANA [ser. II, vol. 1°
6. Caudex erect; spores with thin translucent wing; plants of mountain ridges in Malesia
5. Coryphopteris
6. Caudex slender, creeping; spores various; a mixture still unresolved . . . . 6. Parathelypteris
5. n= 36 or 35; sporangia stalked, in many cases bearing glands or setae.
7. No reduced pinnae at base of frond; subentire pinnae with goniopteroid or meniscioid venation
rarely present.
8. No glands or hairs on body of sporangia except in Thelypteris.
9. Caudex short, erect; spores trilete with discontinuous perispore forming minute papillae; no
scales on costae . . . . . . . . Group of TRIGONOSPORA
10. On rocks by streams, S: E. ‘Asia and W. Malesia sonst Oat. See Oe Se OOS Ona
10. (Related genus in Africa: Menisorus)
9. Caudex creeping; spores monolete with ample sata scales present on lower surface of
costael ». . . Group of THELYPTERIS
11. No large red glands o on hairs ¢ on sporangium- -stalks nor on Jower surface of costae
8. Thelypteris
Large red glands on hairs of sporangium-stalks, sometimes on lower surface of pinnae also.
Eh Veins free; forest plants of Malesia ee - . . . . . . 9. Mesophiebion
12. Veins anastomosing; plants of wet ground in the open.
13. Fronds not proliferous; no forked hairs present oe ee Ue oe ONGC yclosonus
13. Fronds proliferous; forked hairs present . . . . . . =. +. +. +. +. 411. Ampelopteris
8. Glands or setae usually present on body of sporangia >. 2 e . « « GroipoherinGir
14. Veins anastomosing, with long sinus-membrane .. . .- 2) Gee Chines
14. Veins free; sinus-membrane short; decurrent obliquely towards costa . . 13. Plesioneuron
7. Reduced pinnae usually present at base of frond; species without such pinnae mostly having broad
subentire normal pinnae.
15. Hooked hairs on frond; veins free. . . . . . 14, Isolated genus Cyclogramma
15. Hooked hairs present only in Pronephrium seek. Grypothrix which has much anastomosis of
veins.
16. Spores with many small wings or a + continuous translucent wing with cross-wings; aerophores
+ swollen eee . . Group of SPHAEROSTEPHANOS
17. Broad thin scales, lacking superficial hairs, « on base of stipe: capitate hairs, rarely setae, or neither,
on sporangia.
18. Veins free; lamina between them not pustular when dry . . . . . 15. Pseudocyclosorus
18. Veins in most species anastomosing; lamina between them pustular when dry
16. Pneumatopteris
17. Scales on base of stipe narrow, hairy on surface; spherical glands on setae usually present on
sporangia.
19. Reduced pinnae normally present . .... . . +... +. . . 417.Sphaerostephanos
19. Reduced pinnae lacking.
20. Sori in most species round or nearly so; pinnae mostly free.
a Pinnae rarely to 20 pairs; veins anastomosing ... . . . . . . 18.Pronephrium
. Pinnae to 30 pairs, not over2.5cmlong . . . . . . 19, Nannothelypteris
it ‘Sori always elongate along veins; pinnae mostly adnate; sporangia always setiferous
20. Stegnogramma
16. Spores coarsely tuberculate or ridged; Snr not or rarely swollen; elongate glands often
MHS. <3 2 : . . . .Group of CHRISTELLA
22. Basal large minnae much narrowed at base: pairs of reduced basal pinnae inconstantly
present; venation in several species eee, ; ._. 21. Amphineuron
22. Basal pinnae not much narrowed at base, usually auricled: a few pairs of lower pinnae
praduallyneduced lowest notavery small) 272 2 92> 2 2 ee eee inastella
The above conspectus does not attempt to state all characters of the various divisions, but
is an attempt to show inter-relations as I see them at present.
Pseudophegopteris and Macrothelypteris have bipinnate fronds approaching those of
Cyathea in size but differ in their mainly adnate pinnules; as above noted, I doubt whether
they are to be regarded as primitive in the family. Other genera have more definitely
primitive characters, e.g. the miniature arborescent habit of Coryphopteris, the trilete spores
of Trigonospora, but the latter has 36 chromosomes and may be related to Pseudocyclo-
sorus. Coryphopteris plants are confined to acid peaty soil on mountain ridges, where they
grow with Plagiogyria which is certainly a primitive relic. In the New World (and to a small
1981] THELYPTERIDACEAE (Holttum) 341
extent in Africa) Amauropelta, in habit mainly similar to Coryphopteris, is a comparable
genus (of 200 species) but differs in having lower pinnae almost always gradually reduced
whereas that condition is rare in Coryphopteris.
The genera Thelypteris, Mesophlebion, Cyclosorus and Ampelopteris seem to be a natural
group with peculiar glands, scaly costae, creeping rhizome and unreduced basal pinnae.
Mesophlebion looks like the primitive element, adapted to a forest habitat; the other genera
have become adapted to open swampy ground and, like many other swamp-plants, are
widely distributed (it should be noted that Mesophlebion chlamydophorum grows in
swamp-forest). In my judgement, most species which have been included in Cyclosorus are
not at all nearly related to this group of genera. Ampelopteris is peculiar in having forked
hairs on the rachis similar to those of the American genus Goniopteris (in the strict sense of
CHRISTENSEN’s monograph) but is not otherwise much like Goniopteris; if they have a
common ancestor it must be rather far back.
Cyclogramma is a peculiarly isolated genus of a few species in mainland Asia (one of
them reaching northern Luzon). I doubt whether its hooked hairs indicate a relationship to
Pronephrium sect. Grypothrix; it should be noted that slender hooked hairs occur also in
some species of Amauropelta which is not nearly related to either Cyclogramma or
Pronephrium.
The series of genera from Pseudocyclosorus to Stegnogramma comprise the great
majority of Malesian species of the family, and I judge that this is a natural group; the
genera within it are not easy to characterize clearly, though typical species in each are
distinct enough. Most species of the group have anastomosing veins, but in most genera
there are some species with free veins, which again indicates the unnatural state of
COPELAND’s distinction between Lastrea and Cyclosorus by the character of free or
anastomosing veins. Most of these ferns have reduced basal pinnae, but there is much
variation, and I can find no sharp division between Pronephrium and Sphaerostephanos; |
maintain these two genera partly for convenience and partly because many members of
Pronephrium do form a group distinct from typical Sphaerostephanos. The whole question
needs much further study; it will not be solved by examining a few species.
The final group of two genera, Christella and Amphineuron is perhaps also related to the
Sphaerostephanos group, but appears to be distinct in its spores and glands.
Unicellular glands of one sort or another are characteristic of most genera. In many cases
these glands are distinctive when seen at a magnification of not less than x 25, but they are
in some measure modified in the process of preparing specimens for the herbarium, and their
differences are not easy to describe. They need a microchemical study.
The great range of variation in detail within almost all genera appears to me to indicate
that the family is still in an active state of evolution. I cannot agree with LOVE, LOVE and
PICHI SERMOLLI that “most of the recognized species of pteridophytes are evidently old
and well-established taxa” (p. xiv) nor that “the species of ferns are usually clearly distinct
and established through their substantial age” (p. xi), having an existence of “millions of
generations” (p. xiii). It seems to me evident that genera and species in Thelypteridaceae are
in most cases not older than those in many families of Angiosperms.
Citation of literature and synonyms. An attempt has been made to include all binomials
based on Malesian species, with citation of types, all of which have been examined except
where otherwise indicated. Binomials based on extra-Malesian species which are regarded as
synonyms are in most cases cited, or reference is made to their treatment in other recent
literature. Descriptions published in works on Malesian ferns are in all cases cited, though
some of them are of doubtful value. BEDDOME’s work is also cited, as he included
information on ferns of Malaya, and information about the occurrence of Malesian species in
India; similarly the Flore Générale de I’Indochine is often also cited. Some of this in-
formation occurs also in my precursory papers, but in some of these the information is
incomplete, and in all cases the work has been revised for the present Flora.
Descriptive details. So far as possible, these are arranged in the same sequence for all
species. The distance between costules is measured along the costa from which they arise;
this can be more exactly stated than the width of pinna-lobes. The nature and distribution of
342 FLORA MALESIANA [ser. II, vol. 1°
glands and hairs is separately indicated for the lower (abaxial) and upper (adaxial) surfaces;
those on the lower surface are usually more distinctive, but often both need to be observed
to provide clear distinctions between allied species. Failure to observe differences in these
characters has caused much confusion among earlier authors, and more information still
remains to be recorded. Spores have only been approximately described. I hope that a
separate publication on spores in the family may later be possible.
The key to genera which follows is artificial (the numbers of the genera are those of the
Conspectus) and is designed to lead as simply as possible to the right genus for any species
in Malesia. It will not serve a similar purpose for all species in the Old World. I have
compiled taxonomic studies of all species in Africa and adjacent islands (J. S. Afr. Bot. 40,
1974, 123-168) and in Australasia and the Pacific (Allertonia 1, part 3, 1977). Most Indian
species are covered in my precursory papers in Blumea or elsewhere, as cited under the
genera.
KEY TO THE GENERA IN MALESIA
1. Costae not grooved on upper surface; veins not reaching margin.
2. Fronds bipinnate.
. Scales on lower surface of frond, if present, consisting of a row of cells with red cross-walls
acicular hairs all unicellular. . . . . . . 1. Pseudophegopteri:,
. Scales on axes of frond thickened at base,, or replaced by long. slender septate acicular hairs
2. Macrothelypteris
2. Fronds simply pinnate.
4. Strpe-scales bearing superficial hairs; rachis-wing, if present, narrow and of even width.
Dar SOTMIMONSIATE Ih ae ohh ha phases iis Oe cell Dceedie a tip ay Oe WO: acc eal CEE Een EGIaS
5. Soriexindusiate .. . . . . . 1. Pseudophegopteris
4. Stipe-scales with marginal hairs, only: pinnae connected by wings along the rachis forming
semicircular lobes between pinnae . . oo. . . « « «4s Phegopteris
Costae grooved on upper surface; free veins all reaching margin.
2 Fronds simple, or pinnate with lower pinnae not decrescent (small basal pinnae inconstantly present
in Amphineuron).
7. Spores trilete; caudex short, erect; on rocks by streams , f bP a & & 6 Se7eDrivonespora
7. Spores normally monolete; caudex and habitat various.
8. Rhizome long-creeping in wet ground; broad flat scales present on lower surface of costae.
9. Veins free Gb cere ame ee ee ee eee ee eee es OTR
9. Veins anastomosing.
10: Fronds not proliferous; soriindusiate . ....... =. =... . . 105€yclosorus
10. Fronds proliferous; soriexindusiate . . . .. . . =. =. +... +... . 11. Ampelopteris
8. Not this combination of characters.
11. Caudex erect; sporangia unstalked, lacking hairs and glands; some sessile resinous glands on
lower surface in most species; plants of mountain ridges . . . . . . . 5.Coryphopteris
11. Not this combination of characters.
12. Caudex massive, erect; scales narrow, rigid, brittle; sori close to costules, exindusiate or with very
smallindusia . . . ee en eer mer re 1, Cinta
12. Not this combination of characters.
13. Sori elongate along veins, exindusiate; sporangia bearing slender straight setae
20. Stegnogramma
13. Sori round, or if elongate and exindusiate the sporangia bearing hooked hairs or none.
14. Veins free.
15. Basal basiscopic vein of each group arising from costa below the attachment of its costule;
a red gland at the end of hairs on stalks of sporangia. . spaviat o), Mesophlebion
15. Basal basiscopic vein of each group not always thus arising: ‘hairs on sporangium-stalk
otherwise.
16. Rhizome long-creeping; no glands nor hairs on sporangia.
17. Rhizome slender; pinnae rarely over6cmlong . . . . . . . . 6.Parathelypteris
17. Rhizome 5-7 mm diameter; pinnae 10cm or more long . . 22. Christella
16. Rhizome not long- -creeping; sporangia in some cases with glands | or hairs.
18. Bases of lower pinnae not greatly narrowed.
19. Pinnae rigid, commonly at least 10 x 1 cm MV fel 4 ee ee ae ml SS Rlesionenkon
19. Pinnae thin, to 3 cm long, rarely 1 cm wide.
1981] THELYPTERIDACEAE (Holttum) 343
20. Sporangia bearing glands.
a Pinnae 20-30 pairs, no glands on their lower surface . . . . 19. Nannothelypteris
- Pinnae)c..7 pairs; glands\presenty "=. >... . . 3» x. 18. Pronephrium
& ‘Sporangia lacking glands . . Vue. sos. ee a © 25 Olmistelia
18. Bases of lower pinnae much nz rowed Biicst ne) ty Seen VAI DHinennon
a Veins anastomosing.
Spores coarsely tuberculate or ridged; sporangia lacking aan
"93. Basal pinnae much narrowed at their bases oh . . . . 21. Amphineuron
23. Basal pinnae not much narrowed at their bases which are in most cases auricled on
acroscopic side. . . 72 = 222\Chnistella
22. Spores with a thin wing and c Ccross- wings. or many small thin wings.
24. Pinnae 20 pairs or more, 2-3cmlong . .... . . +. . . . 19, Nannothelypteris
24. Pinnae much longer, or fewer.
25. Pinnae or simple fronds subentire; lower surface between veins often pustular
18. Pronephrium
25. Pinnae deeply lobed; surface between veins not pustular . . 17. Sphaerostephanos
6. Lower pinnae gradually reduced, or an abrupt change to small pinnae at base of frond.
26. Hooked hairs present on lower surface of frond lisa 4 5 = « oo el4siCyclogramma
26. Hooked hairs lacking.
27. Rhizome slender, long-creeping; many lower pinnae gradually reduced; some septate acicular
hairs on lower surface. . . er ie ee a 4) Get, 268 Parathelyptenis
27. Not this combination of characters.
28. Caudex erect; pinnae to c. 5cm long, lower ones gradually reduced; sporangia sessile, lacking
glands or hairs; plants of high mountain ridges. . . . . . . . . +. +. +5. Coryphopteris
28. Not this combination of characters.
29. Basal scales on stipe in almost all cases broad and thin; lamina between veins on upper surface
almost always smooth and glabrous, lower surface of costae never densely hairy; glandular hairs
on lower surface, if present, minute and colourless.
30. Veins free; reduced basal pinnae consisting of an aerophore with a minute lamina; pinnae not
pustular between veins on lower surface when dried . . oflatre 15. Pseudocyclosorus
30. Veins anastomosing, or if free the reduced basal pinnae with a distinct green lamina; lower
surface of pinnae between veins +pustular when dried . . . . . 16. Pneumatopteris
29. Basal scales narrow; upper surface between veins often bearing hairs and/or glands, lower surface
of costae usually with copious hairs; sessile spherical or elongate glands often present.
31. Body of sporangium lacking hairs or glands; an elongate unicellular glandular hair on
sporangium-stalk, similar hairs sometimes also on lower surface of veins; spores coarsely
tuberculate or ridged a - fe 2 22-3Christella
31. Body of sporangium usually with sessile spherical glands or setae: hairs on sporangium- -stalk
consisting of several cells; spores with many small thin wings or a continuous wing with
cross-wings; sessile spherical glands present on lower surface in many species
17. Sphaerostephanos
1. PSPEUDOPHEGOPTERIS
CHING, Acta Phytotax. Sinica 8 (1963) 313; HoLTTUM, Blumea 17 (1969)
12. — Thelypteris group 4 CHING, Bull. Fan Mem. Inst. Biol. Bot. 6 (1936)
246. — Phegopteris sensu TAGAWA, Acta Phytotax. Geobot. 7 (1938) 73,
p.p. — Phegopteris sect. Lastrella H. Ito in Nakai & Honda, Nov. FI. Jap.
n. 4 (1939) 152, excl. P. decursive-pinnata. — Thelypteris subg. Phegopteris
sect. Lastrella K. IwAtTs. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 25,
(1965) 137, excl. T. decursive-pinnata. — Toppingia DEG., DEG. & A. R.
SMITH in Deg. & Deg. Fl. Hawaii (1968) fam. 17b. — Macrothelypteris sensu
PICHI SERMOLLI, Webbia 24 (1970) 715, p.p. — Fig. 2a-g.
Caudex erect or prostrate, in P. aurita long-creeping; scales thin with
short hairs on surface; stipe and rachis glossy, in most cases castaneous.
Lamina in most species bipinnate with pinnules adnate to pinna-rachis,
344 FLORA MALESIANA
[ser. II, vol. 1°
Fig. 2. Pseudophegopteris aurita (HOOK.) CHING. a. Rhizome and base of frond, x 6; b. lobe of a pinna,
x 3; c. spore, x 333; d. scale from stipe, x6.— P. rectangularis (ZOLL.) HOLTTUM. e. One pinna,
x 2/3; f. sporangium, x 33.— P. paludosa (BL.) CHING. g. Two pinna-lobes, x 2/3. — Macrothelypteris
torresiana (GAUD.) CHING. h. One pinnule, x 2; i. base of pinna-lobe (sori outlined), < 24; j. scale from
stipe, x 4.— M. polypodioides (HOOK.) HOLTTUM. k. Pinnule-lobes, <2; |. base of pinnule-lobe,
x 32; m. scales on pinna-rachis, x2; n. scale from stipe, x 4. — M. setigera (BL.) CHING. o. Part of
pinnule, showing hairs and scales, x 12 (a-d NAKAIKE 401, e~-g MATTHEW s.n., h-n cult. Kew, o
MOUSSET 718).
1981] THELYPTERIDACEAE (Holttum) 345
often connected by a narrow wing, in two species simply pinnate with lobed
pinnae; pinnae opposite or nearly so, lower ones usually somewhat reduced
and more widely spaced; tips of veins thickened, not running to margin;
scales on lower surface of rachis and pinna-rachis few at maturity of frond,
those on distal axes reduced to a single row of short cells with reddish
cross-walls; hairs on lower surfaces unicellular, acicular or capitate or both.
Sori always exindusiate, globose or + elliptic, in a few cases elongate along
veins; sporangia bearing acicular hairs or not; spores usually pale, with a
slightly-raised reticulum forming broad meshes on the surface.
Type species: Pseudophegopteris pyrrhorhachis (KUNZE) CHING.
Distr. St Helena, S. Tomé & Fernando Poo; tropical Africa; Mascarene Islands; tropical and subtropical
Mainland Asia; Malesia; Samoa, Hawaii; about 20 spp.
Ecol. In Malesia, only on mountains at 1200-2800 m, often near streams, in open places; most species
appear to be local, and have been little collected.
Cytol. Base chromosome number 31; P. pyrrhorhachis diploid, tetraploid and hexaploid (India, Ceylon);
P. aurita diploid (N. India), tetraploid (New Guinea); P. cyclocarpa tetraploid; P. rectangularis tetraploid
(N. India).
This genus has been united by PICHI SERMOLLI with Macrothelypteris, but the two are very distinct in
scales and spores. Pseudophegopteris differs also in its invariable mountain habitat; this is seen strikingly in
the Pacific where Macrothelypteris torresiana and M. polypodioides are widely dispersed at low altitudes,
Pseudophegopteris represented only by two isolated mountain species in Samoa and Hawaii.
KEY TO THE SPECIES
. Rhizome slender, wide- -creeping; some sori elongate
1. P. aurita
: Rhizome erect, or if creeping thick with fronds near together: sori not ‘elongate.
2. Pinnae to c. 6cm long, nearly all adnate to rachis, lobed not more than ; to costa
2. P. rectangularis
2. Pinnae much longer, mostly not adnate to rachis and with adnate pinnules.
3. Basal basiscopic pinnules conspicuously longer than next.
4. Copious short capitate hairs present on lower surface of pinna-rachis and costae of pinnules
4. All hairs on lower surfaces acicular
3. P. tenggerensis
4. P. cyclocarpa
3. Basal basiscopic pinnules not conspicuously longer. than next.
5. Pinnules on largest pinnae almost free, lobed almost to costa.
6. Lower surface of costae hairy; lobes of largest pinnules lobed almost to costule
6. Lower surface of costae glabrous; lobes of largest pinnules almost entire
5. P. kinabaluensis
6. P. sumatrana
5. Pinnules on largest pinnae all broadly adnate to pinna-rachis, lobed not more than half-way to
costa
1. Pseudophegopteris aurita (HOOK.) CHING, Acta
Phytotax. Sinica 8 (1963) 314: HOLTTUM & Roy,
Blumea 13 (1965) 131; Ho_trumM, Blumea 17
(1969) 13. — Gymnogramme aurita HOOK. Ic. PI.
10 (1854) t. 974, 989; Spec. Fil. 5 (1864) 141;
BEDD. Ferns Brit. Ind. Suppl. (1876) 24. — Phe-
gopteris aurita (HOOK.) J. SM. Cat. Cult. Ferns
(1857) 17; Hist. Fil. (1875) 234; METT. Farngatt.
IV (1858) 15.— Polypodium auritum (HOOK.)
Lowe, Ferns Brit. & Exot. 2 (1858) t. 51.—
Leptogramma aurita (HooK.) BEDD. Handb.
(1883) 377, f. 216. — Dryopteris aurita (HOOK.) C.
CHR. Ind. Fil. (1905) 253.— Thelypteris aurita
(HOOK.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 6
(1936) 266.—Type: GRIFFITH, Khasya Hills
(K). — Fig. 2a-d.
Rhizome 2-3 mm diam. (dry), scales near apex
7. P. paludosa
3-4 mm long. Stipe dark purplish, glossy, to 60cm
long. Lamina 30-70 cm long; pinnae well-spaced,
commonly to 15cm long, upper ones adnate to
rachis merging with broadly deltoid apical lamina,
lowest 1-2 pairs reduced; pinnae of largest fronds
lobed almost to costa, lobes rounded to acute,
basal lobes conspicuously longer than next, basal
basiscopic lobes of largest pinnae 3-5 cm long,
lobed up to half-way to costule; veins pinnate in
the pinna-lobes, simple or forked; rachis short-
hairy on upper surface, glabrous beneath; costae
densely short-hairy on both surfaces, rest
glabrous. Sori elongate to globose along middle to
distal part of veins; sporangia setose.
Distr. N. E. India to S. W. China, Tonkin; in
Malesia: Sabah (Mt Kinabalu, CLEMENS 40306,
MOLESWORTH ALLEN 3234), Philippines (Luzon,
346
FLORA MALESIANA
[ser. II, vol. 1°
PRICE 836, Mt San Cristobal), Papua New Guinea
(several coll.).
Ecol. In Malesia at 1400-2200 m, in open places.
2. Pseudophegopteris rectangularis (ZOLL.)
HOLTTUM, Blumea 17 (1969) 19.— Polypodium
rectangulare ZOLL. Syst. Verz. (1854) 37, 48. —
Thelypteris rectangularis (ZOLL.) NAYAR &
KAUR, Comp. to Bedd. (1974) 72.— Type: ZOL-
LINGER 1802, Tjiapoes, Java (G; L, LE, P).
Polypodium distans DON var. minor CLARKE,
Tre linn. .Socs Te Bots 1 ds80)P S455 te 79% ot
1.— Type: CLARKE, Sikkim (K).
Dryopteris moussetii ROSENST. Fedde Repert.
8 (1910) 278. — Phegopteris moussetii (ROSENST.)
v.A.v.R. Handb. Suppl. (1917) 306.— Type:
MOUSSET s.n. Tengger Mts, Java (isotype FI).
Phegopteris oppositipinna v.A.v.R. Bull. Jard.
Bot. Btzg II, 16 (1914) 24; Handb. Suppl. (1917)
307. — Thelypteris oppositipinna (v.A.v.R.)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 6 (1936)
268: HOoLTTUM, Rev. Fl. Malaya 2 (1955)
239, f. 137.— Pseudophegopteris oppositipinna
(v.A.v.R.) CHING, Acta Phytotax. Sinica 8 (1963)
315.— Type: MATTHEW, G. Singgalan, Sumatra
(BO; K). — Fig. 2e-f.
Caudex short, suberect. Stipe to 25cm long,
dark purplish, bearing short sparse spreading
hairs, scaly near base. Lamina to 30 12 cm, nar-
rowed to both ends, several pairs lower pinnae
more widely spaced; free pinnae few, rest with
base of lamina +adnate to rachis; largest pinnae
6x 1.5cm, narrowed evenly from base to apex,
lobed up to ; to costa at base, lobes entire, roun-
ded; veins in basal pinna-lobes 5 or more pairs;
rachis, costae and costules beneath with hairs
0.5mm long, some capitate hairs between veins.
Sori +globose, subterminal on veins, sporangia
with slender setae.
DiisitreeN: YES India,
Malaya, Borneo, Java.
Ecol. On mountains at 1000-1500 m, in open
sandy places by streams.
W. Malesia: Sumatra,
3. Pseudophegopteris tenggerensis HOLTTUM,
Reinwardtia 8 (1974) 500. — Type: BUYSMAN 40,
Tengger Mts, Nongko, 1200 m, 1906 (U).
Caudex unknown. Stipe castaneous, glossy.
Largest pinnae 33cm long, subpinnate; pinnules
lobed, all broadly adnate and connected by a
narrow wing on pinna-rachis, basal basiscopic
pinnule 7*1.6cm, conspicuously longer than
next; lower surface of pinna-rachis and of costae
of pinnae bearing copious short capitate hairs and
a few acicular hairs, upper surface bearing
spreading brown acicular hairs only. Sori almost
globose, those on lower veins ellipsoid; sporangia
lacking setae.
Distr. Malesia: East Java. Known only from
type and two other specimens at Utrecht, possibly
all from same source.
4. Pseudophegopteris cyclocarpa HOLTTUM,
Blumea 13 (1965) 131; Blumea 17 (1969) 15.—
Type: cult. Hort. Bot. Kew. 578/63 n. 57, origin
near Mt Hagen, Western Highlands, Papua New
Guinea, 2000 m, leg. HOLTTUM (K).
Caudex branching; branches short, suberect.
Stipe to 60cm long, dark purplish brown, scaly
and hairy near base, scales 6x 1mm. Lamina to
80 cm or more long, basal pinnae reduced; largest
pinnae to 20 cm long, narrowly deltoid; pinnules all
broadly adnate to pinna-rachis, lowest pair almost
free, basiscopic one to 4.5 x 1.4m, acroscopic to
3.5 1.2cm, lobed 3 towards costa; next pair of
pinnules c. 4 and 2.5 cm long, rest of pinna evenly
narrowed towards apex, middle pinnules 7-8 mm
wide; veins pinnate in lobes of pinnules; acicular
hairs present on lower surface of pinna-rachis at
least near its apex, also on costae of pinnules and
near margin. Sori medial on veins, globose or nearly
sO; sporangia copiously setose.
Distr. Malesia: Papua New Guinea. Known
from type and two other collections from Western
Highlands at 2000-2800 m, also one from Morobe
District at 2000 m.
5. Pseudophegopteris kinabaluensis HOLTTUM,
Blumea 17 (1969) 16. — Type: H. P. FUCHS 21475,
Mt Kinabalu, 2700 m (SRR; K, L).
Caudex not seen. Stipe to 150cm long, pale
brownish, with rachis bearing scattered narrow
scales on lower surface. Lamina c. 150 cm long,
lowest pinnae not or little reduced; largest pinnae
40 cm long, lower pinnules 2.5-3 cm apart, adnate
slightly to rachis on acroscopic side; largest pin-
nules 10*3cm, at base lobed almost to costa;
costules to 6mm apart, lobes incised up to half-
way to costule; lower surface of costae of pin-
nules bearing slender spreading hairs and small
uniseriate scales, hairs on upper surface more
abundant and thicker. Sori almost globose;
sporangia not setiferous.
Distr. Malesia: Sabah (Mt Kinabalu). Only
known from type and one other collection, from
wet inundated ground in narrow Goking’s valley
at 2700-2800 m.
6. Pseudophegopteris sumatrana HOLTTUM,
Blumea 17 (1969) 22.— Type: HOLTTUM 26211,
Kerinci Peak, 2000 m, Sumatra (K; BO, SING).
Caudex of type not recorded; of New Guinea
specimens erect. Stipe 100cm long, reddish.
Lamina 150cm long; lower pinnae distinctly
reduced, largest c. 30cm long; pinnules on basis-
copic side longer than on acroscopic; largest pin-
nules 7.5 X 1.7 cm, acuminate, partly adnate at base
to pinna-rachis, lobed 4 towards costae, lobes
subentire with rounded apices, costules 3-6mm
apart, veins mostly forked; lower surface of rachis,
costae and costules glabrous. Sori above the fork on
a vein, sometimes on both branches, almost glo-
bose; sporangia not setiferous.
1981]
THELYPTERIDACEAE (Holttum)
347
Distr. Malesia: Central West Sumatra (Mt
Kerinci). In addition to the type, several collec-
tions from Papua New Guinea at 1700-2300m
appear to be referable to this species.
Ecol. Open place by small stream.
Note. P. sumatrana is related to P.
kinabaluensis but smaller, and glabrous on lower
surface. New Guinea specimens resemble the
Sumatran type, not the plants from Kinabalu, but
it is possible that the two should be united.
7. Pseudophegopteris paludosa (BL.) CHING, Acta
Phytotax. Sinica 8 (1963) 315; HOLTTUM & Roy,
Blumea 13 (1965) 131; HoLTTUM, Blumea 17
(1969) 23.— Aspidium paludosum BL. Enum. PI.
Jav. (1828) 168, non RApDDr 1825.— Polypodium
paludosum BL. FI. Jav. Fil. (1851) 192, t. 90;
HOOK. Spec. Fil. 4 (1862) 214. — Macrothelypteris
paludosa (BL.) LOVE & LOVE, Taxon 26 (1977)
325. — Type: BLUME, Java (L, n. 908,335-309).
Polypodium distans sensu RACIB. FI. Buitenz. 1
(1898) 96.— Dryopteris distans sensu v.A.v.R.
Handb. (1908) 496, p.p.—Dryopteris brunnea
(WALL.) C. CHR. Ind. Fil. (1905) 255, nom. nud.:
BACKER & POSTH. Varenfl. Java (1939) 46,
p.p.— Thelypteris brunnea sensu CHING, Bull.
Fan Mem. Inst. Biol. Bot. 6 (1936) 74, p.p.;
HOLTTUM, Rev. FI. Malaya 2 (1955) 240 p.p. —
Lastrea pyrrhorhachis sensu. COPEL. Fern FI.
Philip. (1960) 330 p.p. — Fig. 2g.
Caudex massive, erect; stipe and rachis dark
reddish. Frond with stipe to more than 200cm
long; 2-4 pairs lower pinnae gradually reduced;
largest pinnae 20-35 cm long, bearing broadly ad-
nate segments or pinnules 2.5—-4 cm long which are
contiguous except in largest pinnae; costules 10-
12(-15)mm_ apart; basal basiscopic pinnules lar-
gest but not much longer than next; lower surface
of pinna-rachis, costae, costules and veins bearing
slender acicular hairs almost | mm long. Sori glo-
bose or nearly so; sporangia usually not setose.
Distr. Apparently throughout Malesia, to
Eastern New Guinea, on mountains at 1200-
2500 m, but specimens are few.
Ecol. BLUME’s type was from an open swamp.
The first collection from Malaya was in a small
forest clearing at Cameron Highlands; now where
much of the forest is cleared plants are abundant
along streams at 1500m. At Fraser’s Hill quite
small plants, on drier ground, have fertile fronds.
Notes. The specific epithet brunnea, taken up
by CHRISTENSEN in Index Filicum and sub-
sequently used by several authors, was never val-
idated. This Malesian species differs from the
Indian P. pyrrhorhachis (KUNZE) CHING in its
massive erect caudex and larger size. I cannot
decide, from herbarium specimens, how many
species of this alliance occur in India; there, as in
Malesia, more field work is needed.
2. MACROTHELYPTERIS
(H. ITo) CHING, Acta Phytotax. Sinica 8 (1963) 308; HOLTTUM, Blumea 17
(1969) 25; Allertonia 1 (1977) 177. — Thelypteris group 10 CHING, Bull. Fan
Mem. Inst. Biol. Bot. 6 (1936) 248. — Thelypteris sect. Macrothelypteris H.
ITO in Nakai & Honda, Nov. FI. Jap. n. 4 (1939) 141; K. IwaTs. Acta
Phytotax. Geobot. 18 (1960) 155.— Thelypteris subg. Thelypteris sect.
Metathelypteris K. IWATS. Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 145,
p.p. — Fig. 2h-o.
Caudex short, creeping or suberect; scales (also on base of stipe) narrow,
+ thickened at least near base, with marginal and superficial acicular and/or
capitate hairs. Lamina _ bipinnate-tripinnatifid with +adnate pinnules;
lowest pinnae little reduced; scales on rachis and pinna-rachis narrow,
pallid, thickened at base and sometimes wholly terete, with or without
marginal hairs; hairs on surface of frond slender and acicular or short and
capitate, some long multicellular hairs always present. Sori always small,
usually with a small but persistent indusium often hidden by mature
sporangia; sporangia bearing capitate hairs; spores with a very fine surface
reticulum not resolvable by light microscope, and slight wings.
Type species: Macrothelypteris oligophlebia (BAK.) CHING.
Distr. Mascarene Islands; warmer parts of mainland Asia; Malesia; Queensland; islands of the
Pacific (including Hawaii) south to Kermadec Island; 10 species.
348
FLORA MALESIANA
[ser. II, vol. 1°
As pointed out by CHING, the type selected by ITO is a little-known species from central China. In
1969 I placed it as a variety of M. torresiana, but | am now doubtful about this, and have not seen
enough material to make a good judgement. In 1965 IWATSUKI included this genus in Thelypteris subg.
Thelypteris sect. Metathelypteris but the scales of the latter (here ranked as a genus) are different, also
the chromosome number.
Cytol. Base chromosome number 31; M. ornata diploid (N. India); M. torresiana tetraploid (India,
Ceylon, Singapore) and hexaploid (Ceylon); M. viridifrons tetraploid (Japan).
KEY LO LAE SPECirES
1. Rachises and lower surface of axes of frond lacking scales
1. M. torresiana
1. Rachis, pinna-rachises and axes of pinnules scaly on lower surface.
Nh
ciliate.
3. Basal scales thin; scales on rest of stipe and rachis flat above the terete base
. Scales on main rachis sparse; scales on pinna-rachis copiously ciliate
. Scales on main rachis abundant, with strongly thickened bases; scales on pinna-rachis not or little
2. M. polypodioides
3. M. setigera
3. Basal scales rigid with inflexed edges; scales on rest of stipe and rachis almost wholly terete
1. Macrothelypteris torresiana (GAUD.) CHING,
Acta Phytotax. Sinica 8 (1963) 310; HOLTTUM,
Blumea 17 (1969) 27, excl. syn. Polypodium fragile
BAK. — Polystichum torresianum GAUD. in
Freyc. Voy. Uran. Physic. Bot. (1828) 333.—
Thelypteris torresiana (GAUD.) ALSTON, Lilloa
30 (1960) 111; K. IwATs. Mem. Coll. Sci. Univ.
Kyoto B, 31 (1965) 151, 153.— Type: GAUDI-
CHAUD, Mariana Is. (P; G).
Aspidium uliginosum KUNZE, Linnaea 20
(1847) 6.—Dryopteris uliginosa (KUNZE) C.
Cure. Ind. Fil. Suppl. II (1934) 100; BACKER &
POSTH. Varenfl. Java (1939) 42, p.p. — Thelypteris
uliginosa (KUNZE) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 6 (1936) 342; HoL_truM, Rev. FI.
Malaya 2 (1955) 241; BROWNLIE in Aubrév. FI.
Nouv. Caléd. 3 (1969) 209; MORTON, Contr. U.S.
Nat. Herb. 38 (1973) 219. — Type: cult. Hort. Bot.
Leipzig, origin Java (BR).
Polypodium trichodes HOULST. & MOORE,
Gard. Mag. Bot. 3 (1851) 18; MORTON, Contr. U.S.
Nat. Herb. 38 (1973) 264. — Type: cult., origin “East
Indies” (BM).
Polypodium tenericaule HOOK. in J. Bot. Kew
Misc. 9 (1857) 353.—Lastrea tenericaulis
(HOOK.) Moore, Ind. Fil. (1858) 99; BEDD.
Handb. (1883) 266; TAGAWA, Index Pterid. Jap.
(1959) 223.— Nephrodium tenericaule (HOOK.)
HOOK. Spec. Fil. 4 (1862) 142, p.p. excl. t. 269. —
Type: ALEXANDER, China (K).
Dryopteris setigera var. pallida v.A.v.R. Handb.
(1908) 203; Suppl. (1917) 169.— Type: origin not
specified (BO).
Dryopteris _trichodes ROSENST. Meded.
Rijksherb. Leiden 31 (1917) 6, p.p.; MORTON,
Contr. U.S. Nat. Herb. 38 (1973) 265. — Lectotype
(MORTON): ZOLLINGER 354, Java (L).
Nephrodium setigerum sensu HOOK. & BAK.
Syn. Fil. (1867) 284, p.p.— Aspidium setigerum
sensu RACIB. Fl. Buitenz. 1 (1898) 178, p.p.? —
Dryopteris setigera sensu C. CHR., Ind. Fil. (1905)
292, p.p.; v-A.v.R. Handb. (1908) 202, p.p.; C.
4. M. multiseta
CHR. Gard. Bull. Str. Settl. 7 (1934) 243. — Fig.
2h-j.
Caudex short-creeping. Stipe to 50cm long,
glaucous when young, persistent base swollen and
fleshy covered with many narrow dark brown
scales bearing both acicular and capitate hairs,
rest of stipe and rachis smooth. Lamina to 70x
50cm, deeply tripinnatifid, pinnae 12-15 pairs,
sub-basal ones longest; largest pinnae 20*9cm,
deltoid, pinnate with all pinnules but lowest ad-
nate to a narrowly green-winged pinna-rachis;
pinnules oblique to pinna-rachis, largest 5-8 x 1.5-
2.5 cm, acuminate, cut almost to costa into oblique
dentate to deeply lobed segments 2.5 —4 mm wide,
largest to 12 mm long; costae and costules bearing
scattered pale slender hairs on lower surface,
some of them multicellular and over 1 mm long;
whole lower surface of lamina bearing short capi-
tate hairs. Sori globose; indusia very small bearing
a few capitate hairs; sporangia bearing 2-3 short
capitate hairs.
Distr. Mascarene Islands; warmer parts of
mainland Asia and Japan; Malesia; Queensland,
Polynesia, Hawaii; adventive at various places in
the New World from Florida southwards.
Ecol. In Malesia in open or lightly shaded
places in low country, especially in areas with a
dry season.
Note.GAUDICHAUD described this species very
carefully, but his description was overlooked, and
there is much confusion in the literature between
this species, M. setigera and M. polypodioides.
The first author to distinguish it clearly was
CHING, under the name Thelypteris uliginosa
(1936) but BACKER & POSTHUMUS (1939)
ignored the distinction between T. uliginosa and
T. setigera. See also note under M. polypodioides.
2. Macrothelypteris polypodioides (HOOK.)
HOLTTUM, Blumea 17 (1969) 29; Allertonia 1
(1977) 179. — Alsophila polypodioides HOOK. in
Nightingale, Oceanic Sketches (1835) 131.—
1981]
THELYPTERIDACEAE (Holttum)
349
Type: NIGHTINGALE, “South Sea Islands” (K).
Lastrea leucolepis PRESL, Epim. Bot. (1851) 39;
COPEL. Fern Fl. Philip. (1960) 332. — Dryopteris
leucolepis (PRESL) MAXON, Proc. Biol. Soc.
Wash. 36 (1923) 172.— Thelypteris leucolepis
(PRESL) CHING, Bull. Fan Mem. Inst. Biol. Bot. 6
(1936) 345. — Macrothelypteris leucolepis (PRESL)
CHING, Acta Phytotax. Sinica 8 (1963) 309. —
Type: CUMING 114, Luzon (PRC; BM, K).
Cheilanthes gigantea CESATI, Rendic. Ac. Sci.
Nap. 16 (1877) 25, 29.— Polypodium cheilan-
thoides BAK. in Beccari, Malesia 3 (1866) 45,
nom. nov. (not P. gigantea DESv.).— Dryopteris
brunneo-villosa C. CHR. Ind. Fil. (1905) 255, nom.
nov. — Phegopteris cheilanthoides (BAK.)
v.A.v.R. Handb. (1908) 494.— Type: BECCARI,
Mt Arfak, W. New Guinea (FI; K). — Fig. 2k-n.
Caudex short, prostrate; stipe to 80cm long,
pale, basal half at least bearing copious acicular
and capitate hairs and scales; scales thin, pale, to
20 mm long, hardly | mm wide at thickened base,
bearing acicular hairs on surface near base, dis-
tally on edges, old stipe covered with wart-like
bases of fallen scales. Lamina to 80cm long;
lower pinnae to 35 X 15 cm, rest gradually smaller;
lower surface of rachis (sparsely) and pinna-rachis
(copiously) bearing narrow pale scales to 3mm
long, 0.3 mm wide with thickened bases and slen-
der patent marginal hairs; largest pinnules 10x
2.5 cm, almost at right angles to pinna-rachis, with
tertiary leaflets connected by a very narrow wing
on costa; upper surface of costa densely hairy, of
costules sparsely, very narrow hair-pointed and
ciliate scales on lower surface of costa grading
distally to slender septate hairs; tertiary leaflets at
right angles to costa, larger ones deeply lobed,
capitate hairs on lower surface. Sori small, on
acroscopic branch of a vein; indusia small, with
capitate hairs; sporangia with capitate hairs.
Distr. S.E. Asia (Thailand, Taiwan), through
Malesia (Philippines, New Guinea) to N.E. Aus-
tralia (Queensland) and the Pacific (Cook Is., Fiji,
Samoa, Tahiti, Rapa, Austral Is.).
Ecol. At low altitudes, to 700m, on edge of
forest.
Note. When writing Species Filicum, HOOKER
suppressed his earlier name Alsophila_poly-
podioides (although citing the specimen) and this
was not listed by CHRISTENSEN in Index Filicum.
This species has been much confused with the
preceding and with M. setigera. In 1923 MAXON
pointed out the distinctions between M. poly-
podioides (which he named Dryopteris leucolepis)
and M. torresiana (which he named D. setigera).
3. Macrothelypteris setigera (BL.) CHING, Acta
Phytotax. Sinica 8 (1963) 309; HOLTTUM, Blumea
17 (1969) 31.— Cheilanthes setigera BL. Enum.
Pl. Jav. (1828) 138. — Dryopteris setigera (BL.) O.
KTZEs Rev, Gent) PIF 289i) MSisse veAuveRe
Handb. (1908) 202, p.p.; Suppl. (1917) 169, excl.
var. pallida; BACKER & POSTH. Varenfl. Java
(1939) 337, p.p.— Thelypteris setigera CHING,
Bull. Fan Mem. Inst. Biol. Bot. 6 (1936) 345. —
Type: BLUME, Java (L, n. 908, 337-1168).
Cheilanthes stenophylla KUNZE, Bot. Zeit. 6
(1848) 212. — Type: ZOLLINGER 2675, Java (L).
Dryopteris backeri v.A.v.R. Bull. Dép. Agr. Ind.
Néerl. 18 (1908) 8; ibid. 21 (1908) 3; Handb. (1908)
817; Suppl. (1917) 169, 171.—Type: BACKER,
Krakatau (BO, not seen).
Aspidium vile (non KUNZE) RAcIB. FI.
Buitenz. 1 (1898) 173.—Dryopteris uliginosa
sensu BACKER & POSTH. Varenfl. Java (1939) 42,
p.p. — Fig. 20.
Similar to M. polypodioides in size and in
divisions of lamina, differing in abundant scales
with thickened terete bases and entire (rarely cili-
ate) margins on rachis and pinna-rachis. A com-
parison of living plants from Java and the Philip-
pines needs to be made.
Distr. Malesia: Sumatra, Java, Lombok,
Flores, Timor, S.W. Celebes, Ternate.
Ecol. In open places at medium altitudes;
reported by RACIBORSKI to form extensive
thickets with Gleichenia hispida on G. Guntur in
Java. Small plants may be fertile; one such was
type of Cheilanthes stenophylla KUNZE.
4. Macrothelypteris multiseta (BAK.) CHING, Acta
Phytotax. Sinica 8 (1963) 309; HoLTTUM, Blumea
17 (1969) 31. — Nephrodium multisetum BAK. J.
Linn. Soc. Bot. 22 (1886) 226.— Dryopteris
multiseta (BAK.) C. CHR. Ind. Fil. (1905) 279;
v.A.v.R. Handb. (1908) 203; Suppl. (1917) 171; C.
CHR. Gard. Bull. Str. Settl 7 (1934) 243.—
Thelypteris multiseta (BAK.) CHING, Bull. Fan
Mem. Inst. Biol. Bot. 6 (1936) 347. — Type: G. F.
HOSE, G. Matang, Sarawak, 610 m (K).
Stipe at least 70cm long, reddish when old;
basal scales rigid, 10 x 1 mm, base thickened and
edges inrolled; scales above base copious, per-
sistent, dark, hair-pointed, entire, 5-7 mm long,
hardly 0.5 mm wide, basal part terete; main rachis
similarly and persistently scaly. Pinnae commonly
35cm long, maximum 50cm; pinna-rachis pale
with darker spreading rigid entire scales which are
almost wholly terete; pinnules commonly 8 x
1.8cm, bearing tertiary leaflets connected by a
narrow wing, axis of pinnule scaly as pinna-rachis
but scales smaller, hair-pointed; tertiary leaflets
lobed half-way to costule; lower surface of lamina
bearing short capitate hairs. Sori one to each lobe
of a tertiary leaflet; indusium small, with capitate
hairs; sporangia with similar hairs.
Distr. Malesia: Borneo (North Borneo,
Sarawak), Sumatra.
Ecol. In rather open places at 500-1250 m,
sometimes abundant, but very local; only found at
one place on Mt Kinabalu.
350 FLORA MALESIANA [ser. II, vol. 1°
3. METATHELYPTERIS
(H. Iro) CHING, Acta Phytotax. Sinica 8 (1963) 304; HOLTTUM, Blumea 19
(1971) 26; J.S. Afr. Bot. 40 (1974) 127; Kalikasan 5 (1976) 115. — Thelypteris
sect. Metathelypteris H. Iro in Nakai & Honda, Nov. FI. Jap. n. 4 (1939)
137; K. IwaTs. Acta Phytotax. Geobot. 18 (1960) 147; Mem. Coll. Sci. Univ.
Kyoto B, 31 (1965) 145, p.p. — Fig. 1m, 3.
Caudex short, erect or decumbent with tufted fronds; stipes green when
living, scaly at base only, scales less than 10mm long, narrow with a few
short acicular hairs; lamina simply pinnate (in M. flaccida almost bipin-
nate) with deeply lobed pinnae, basal pinnae not or little reduced; veins
free, often forked, with thickened ends not reaching margin; upper surface
of costae prominent, not grooved; acicular and/or capitate hairs, not
spherical glands, present on lower surfaces; scales on lower surface of costae,
if present, uniseriate, about as in Pseudophegopteris; sori indusiate, indusia
thin; sporangia sometimes with a hair of several cells on the stalk, no hairs nor
glands on body; spores with irregular ridges variously united.
Type species: Metathelypteris gracilescens (BL.) CHING.
Fig. 3. Metathelypteris gracilescens (BL.) CHING. a. One pinna, X a De pinna-lobes, lower surface, X 6; c.
pinna-lobes, upper surface, x 6.— M. dayi (BEDD.) HOLTTUM. d. Pinna-lobe, Xx 4. — M. flaccida (BL.)
CHING. e. Pinna-lobe, upper surface, x 4; f. lower surface, x 12. — M. singalanensis (BAK.) CHING. g.
Base of a pinna-lobe, <6.— M. uraiensis (ROSENST.) CHING. h. Sorus, xX 16 (a-c HOLTTUM 53; d
DAY s.n.; e~-f MATTHEW s.n.; h M. G. PRICE 3368).
3)3)|
1981] THELYPTERIDACEAE (Holttum)
Distr. Fernando Poo & S. Tomé; Nigeria (an undescribed species); Madagascar, Ceylon & India, S.
China to Japan; Malesia; about 12 species.
Ecol. In Malesia all species occur on mountains at 1000-2000 m, usually on steep earth slopes or
rocks. The only species which are both widely distributed and locally common are the tetraploid M. dayi
and the (probably tetraploid) form of M. flaccida which has long hairs on the lower surface; these occur
in somewhat open places, especially where paths have been made in the forest. Of the rest, M.
singalanensis is a forest fern of restricted distribution in Sumatra, M. gracilescens is widely distributed
but rarely abundant.
Cytol. Chromosome number 35 (M. dayi tetraploid; M. flaccida diploid and tetraploid in Ceylon).
KEY LO tHE SPECIES
1. Veins always simple except sometimes in basal acroscopic lobes of pinnae . 1. M. gracilescens
1. Veins normally forked except distal ones.
. Pinnae with some hairs on lower surface between veins; lobes of larger pinnae lobed more than half
way to costule
2. M. flaccida
2. Pinnae not hairy between veins on lower surface: pinna: lobes entire to crenate, shallowly lobed only
on largest fronds.
3. Pinnae rarely over 6 cm long; indusia bearing acicular hairs
. 3. M. uraiensis
3. Pinnae of well-grown plants 12 cm or more long; indusia bearing capitate hairs.
4. Lower surface of costae and costules bearing copious capitate hairs.
5. Texture thin, drying dark green; veins conspicuously branched
4. M. singalanensis
5. Texture firm, drying yellow-green; veins forked near tips, branches little diverging
4. Lower surfaces of costae and costules bearing sparse acicular hairs
1. Metathelypteris gracilescens (BL.) CHING, Acta
Phytotax. Sinica 8 (1963) 305; HOLTTUM, Kalik-
asan 5 (1976) 116.— Aspidium gracilescens BL.
Enum. PI. Jav. (1828) 155; RAciB. FI. Buitenz. 1
(1898) 170. — Lastrea gracilescens (BL.) MOORE,
Ind. Fil. (1858) 93; BEDD. Handb. (1883) 234, p.p.;
Suppl. (1892) 51, p.p.; COPEL. Fern FI. Philip.
(1960) 322; TAGAWA, Col. Illus. Jap. Pterid. (1962)
f. 240.— Nephrodium gracilescens (BL.) HOOK.
Spec. Fil. 4 (1862) 93, p.p.— Dryopteris
gracilescens (BL.) O. KTZE, Rev. Gen. PI. 2 (1891)
812; v.A.v.R. Handb. (1908) 186 excl. syn. Aspi-
dium glanduligerum; BACKER & POSTH. Varenfl.
Java (1939) 38.— Thelypteris gracilescens (BL.)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 6 (1936)
327; K. IwaTs. Mem. Coll. Sci. Univ. Kyoto B,
31 (1965) 147. — Type: BLUME, Java (L).
Nephrodium vulcanicum BAK. Ann. Bot. 5
(1891) 127. — Thelypteris vulcanica (BAK.) REED,
Phytologia 17 (1968) 324. — Type: HANCOCK, G.
Pangerango, Java (K).
Athyrium benguetense CHRIST, Philip. J. Sci. 2
(1907) Bot. 161; COPEL. Philip. J. Sci. 3 (1908)
Bot. 300. — Type: LOHER, Mt Data, Luzon, Feb.
1894 (not found at P).
Dryopteris calva COPEL. in Elmer Leafl. Philip.
Bot. 3 (1910) 805.— Dryopteris gracilescens var.
calva (COPEL.) C. CHR. Ind. Fil. Suppl. III (1934)
82.— Lastrea calva (COPEL.) COPEL. Gen. Fil.
(1947) 138, Fern Fl. Philip. (1960) 323. — Type:
ELMER 11485, Mt Apo, Mindanao (MICH; BM,
EK):
Dryopteris sublaxa HAYATA, Icon. Pl. Formos.
4 (1914) 183, f. 122. — Type: HAYATA, Mt Arisan,
Taiwan, Jan. 1912 (TI, seen by IWATSUKI]).
4b. M. singalanensis var. surbeckii
5. M. dayi
Dryopteris arisanensis ROSENST. Hedwigia 56
(1915) 340.— Type: FAURIE 389, Mt Arisan,
Taiwan (isotype KY seen by IWATSUKI). — Fig.
3a-c.
Caudex short-creeping; stipe 18-25cm long,
pale, scales 2mm long. Lamina 25-30cm long;
pinnae 15-18 pairs, rather close; basal pinnae nar-
rowed at basiscopic base. Largest pinnae of type
7.5X1.4cm, of type of Nephrodium vulcanicum
9x2.0cm, short-acuminate, lobed to 1mm from
costa or more deeply, lobes entire, separated by
narrow sinuses, basal acroscopic lobe of middle
pinnae somewhat elongate; costules commonly
3 mm apart; veins 6-7 pairs, simple or rarely for-
ked in basal acroscopic pinna-lobes; lower surface
of rachis and costae with varied complement of
short acicular and capitate hairs, sometimes al-
most glabrous. Sori medial or a little supramedial,
near tips of veins; indusia small, thin, glabrous.
Distr. S. Japan, Taiwan; Darjeeling; in
Malesia: Malaya, Sumatra, W. Java, N. Borneo,
Luzon, Mindoro, Mindanao, New Guinea.
Ecol. In Malesia at 1500-2200 m, in forest, in
two cases on rocks near waterfall; only one col-
lection from Malaya, from 1800 m at Cameron
Highlands.
Note. M. decipiens (CLARKE) CHING, des-
cribed from Darjeeling, differs in having shorter
fronds with basal pinnae largest, veins mostly
forked, at least in the crenate lobes of basal
pinnae.
2. Metathelypteris flaccida (BL.) CHING, Acta
Phytotax. Sinica 8 (1963) 306.— Aspidium flac-
cidum BL. Enum. Pl. Jav. (1828) 161; RACIB. FI.
S52
Buitenz. 1 (1898) 176.— Lastrea flaccida (BL.)
Moore, Ind. Fil. (1858) 92; BEDD. Ferns S. India
(1864) t. 250; Handb. (1883) 244. — Nephrodium
flaccidum (BL.) HOOK. Spec. Fil. 4 (1862) 133, t.
263. — Dryopteris flaccida (BL.) O. KTZE, Rev.
Gen. PI. 2 (1891) 812; v.A.v.R. Handb. (1908) 195;
BACKER & POSTH. Varenfl. Java (1939) 41.—
Thelypteris flaccida (BL.) CHING, Bull. Fan
Mem. Inst. Biol. Bot. 6 (1936) 336.— Type:
BLUME, Boerangrang, Java (L, n. 908,332-
1093). — Fig. 3e-f.
Caudex short, erect. Stipe 25-45 cm long, pale,
hairy in groove only. Lamina 25-45 cm long; pin-
nae to 15 pairs, well-spaced; lowest pinnae nar-
rowed a little on basiscopic side. Largest pinnae
8.5—12 x 1.8-3.5 cm, acuminate, lobed to a narrow
wing on costa; lobes of largest pinnae lobed more
than half way to costule, their tips blunt-pointed;
costules 4-6mm apart; veins pinnate in larger
lobules of pinna-lobes, forked in smaller ones;
lower surface bearing slender erect acicular hairs
throughout, those on costae and costules 1 mm
long in typical form (some plants in Ceylon and
Malaya have hairs less than 0.5mm). Sori 1-2 in
each lobule of a pinna-lobe; indusia thin, pale,
bearing short acicular hairs.
Distr. Ceylon & S. India, N.E. India to Yun-
nan, Thailand; in Malesia: Java, Sumatra (?),
Malaya, Borneo.
Ecol. In W. Java abundant at 1500m. In
Malaya not found until in recent years plants have
appeared on earth banks beside roads where
forest has been cleared at Cameron Highlands and
on Taiping Hills; these plants have short hairs on
the lower surface of costae as in some plants
found by MANTON in Ceylon.
3. Metathelypteris uraiensis (ROSENST.) CHING,
Acta Phytotax. Sinica 8 (1963) 306; C. M. Kuo, FI.
Taiwan | (1975) 419; HOLTTUM, Kalikasan 5 (1976)
117. — Dryopteris uraiensis ROSENST. Hedwigia
56 (July 1915) 341.— Thelypteris uraiensis
(ROSENST.) CHING, Bull. Fan Mem. Inst. Biol. Bot.
6 (1936) 336; K. IwaTs. Mem. Coll. Sci. Univ.
Kyoto B, 31 (1965) 148, 152. — Lastrea uraiensis
(ROSENST.) TAGAWA, Col. Ill. Jap. Pterid. (1962)
223.— Macrothelypteris uraiensis (ROSENST.)
LOVE & LOVE, Taxon 26 (1977) 325.— Type:
FAURIE 22, Urai, Taiwan (isotypes G, P).
Dryopteris hirtisquamata HAYATA, Icon. PI.
Formos. 5 (Nov. 1915) 277.—Type: T. Iro &
Fusu, Taiwan (TI, seen by IWATSUKI). — Fig. 3h.
Caudex short-creeping; stipe c. 20 cm long, pale.
Lamina c. 18 X 12cm; pinnae to 18 pairs; lowest
pinnae short-stalked and narrowed at base. Lar-
gest pinnae 4.5-6.0cm long, 1.4-1.8cm wide,
lobed to 1 mm from costa; costules 3-4 mm apart;
veins to 6 pairs, mostly forked; lower surface of
costae, costules and veins bearing acicular hairs,
hairs between veins sparse. Sori medial to
supramedial, near tips of veins; indusia thin with
FLORA MALESIANA
[ser. II, vol. 1°
short acicular hairs on edge.
Distr. S. Japan, Ryukyu Islands, Taiwan,
Kwangtung, Yunnan; in Malesia: Philippines (N.
Luzon: M. G. PRICE 2915, Solsona; 3368, Mt
Data, Ilocos Norte Prov.).
Note. This species is very near M. gracilescens
but appears to be distinct in the characters men-
tioned in the key.
4. Metathelypteris singalanensis (BAK.) CHING,
Acta Phytotax. Sinica 8 (1963) 306. — Nephrodium
singalanense BAK. J. Bot. 18 (1880) 212. — Las-
trea singalanensis (BAK.) BEDD. Handb. Suppl.
(1892) 54.— Dryopteris singalanensis (BAK.) C.
Cure. Ind. Fil. (1905) 293; v.A.v.R. Handb. (1908)
194. — Thelypteris singalanensis (BAK.) CHING,
Bull. Fan Mem. Inst. Biol. Bot. 6 (1936) 334. —
Type: BECCARI 471, G. Singgalang 1700 m, Suma-
tra (K; FI n. 12963 in Herb. Becc.).
Dryopteris media v.A.v.R. Bull. Jard. Bot. Btzg
II, 11 (1913) 9; Handb. Suppl. (1917) 162. — Type:
MATTHEW 514, G. Singgalang, Sumatra (BO;
K). — Fig. 3g.
a. var. singalanensis
Caudex short, erect; stipe 30-50 cm long, pale;
scales c. 7mm long, narrow above widened base
which is fully 1 mm wide. Lamina 50-70 cm long;
pinnae to 25 pairs, often almost opposite, drying
dark green, thin; basal pinnae not reduced. Lar-
gest pinnae of type 9.3x2.3cm, of other speci-
mens to 12X3.0cm (fertile), 14x 4cm (sterile),
lobed almost to costa; lobes of almost all pinnae
crenate, of largest ones distinctly lobed; veins
10-12 pairs, nearly all forked, in largest lobes
twice forked; lower surface of rachis, costae,
costules and veins bearing copious very short
capitate hairs and sometimes a few acicular hairs,
some capitate hairs also on surface between veins;
upper surface of rachis and costae bearing
copious acicular hairs 0.2 mm long, hairs on cos-
tules minute. Sori on acroscopic branches of
veins; indusia bearing many short capitate hairs.
Distr. Malesia: Central W. Sumatra (G. Sing-
galang).
Ecol. ‘ton bank by path in forest” (MAT-
THEW); two collections from 1500 and 2100 m.
b. var. surbeckii HOLTTUM, var. nov.
A typo speciei differt: pinnis in sicco brunneo-
virentibus, firmioribus; venis saepe simplicibus,
versus apicem tantum breviter furcatis; pagina
inferiore inter venas pilis capitatis destituta. —
Type: SURBECK 1193, Sibuatan-Zuid, Sumatra,
1700 m (L). Also ALSTON 14974, Sidikalang (BM).
Distr. Malesia: Central Sumatra, c. 350km
north of G. Singgalang.
5. Metathelypteris dayi (BEDD.) HOLTTUM in
Nayar & Kaur, Comp. to Bedd. (1974) 205; Kal-
ikasan 5 (1976) 117.— Aspidium dayi BEDD. J.
1981]
Bot. 26 (1888) 4.— Lastrea dayi (BEDD.) BEDD.
Handb. Suppl. (1892) 54.—Dryopteris dayi
(BEDD.) C. CHR. Ind. Fil. (1905) 260; v.A.v.R.
Handb. (1908) 188.— Thelypteris dayi (BEDD.)
NAYAR & KAuR, Comp. to Bedd. (1974) 59. —
Type: J. DAy, Perak, March 1887 (K).
Nephrodium creaghii BAK. Kew Bull. (1898)
230.— Dryopteris creaghii (BAK.) C. CHR. Ind.
Fil. (1905) 258; v.A.v.R. Handb. (1908) 185; C.
Cure. Gard. Bull. Str. Settl. 7 (1934) 240. — Type:
CREAGH, N. Borneo (K).
Dryopteris flavo-virens ROSENST. Fed. Rep. 10
(1912) 334; v.A.v.R. Handb. Suppl. (1917) 165.—
Lastrea flavo-virens (ROSENST.) COPEL. Gen.
Fil. (1947) 138: Philip. J. Sci. 78 (1951) 423.—
Thelypteris flavo-virens (ROSENST.) REED, Phy-
tologia 17 (1968) 277.—Type: BAMLER W.11
partim, Wareo, 600m, Papua New Guinea, 11
June 1909 (S-PA).
Dryopteris aureoviridis ROSENST. Fed. Rep. 13
(1914) 216; v.A.v.R. Handb. Suppl. (1917) 161.—
Thelypteris aureoviridis (ROSENST.) REED, Phy-
tologia 17 (1968) 262. — Type: J. WINKLER 179, in
terra Batacorum, Sumatra, 1911 (not seen; Ros. Fil.
Sumatr. exsicc. 141 from same locality at K, L).
Dryopteris diversivenosa v.A.v.R. Bull. Jard.
Bot. Btzg II, 28 (1918) 23.— Thelypteris diver-
sivenosa (v.A.v.R.) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 251.— Type: BUNNEMEIJER
1104, Tanang Taloe, 1000 m, Ophir Distr. Sumatra
(BO).
Thelypteris singalanensis sensu HOLTTUM,
Rev. Fl. Malaya 2 (1955) 243, p.p., f. 138; M. G.
THELYPTERIDACEAE (Holttum)
353
PRICE, Brit. Fern Gaz. 10 (1973) 255. — Fig. 1m,
3d.
Caudex short, erect; plants fertile from a small
size. Stipe 20-45 cm long, pale; basal scales very
narrow, 5-7mm long, not widened at base.
Lamina to 50cm long, firm, drying yellow-green;
free pinnae c. 15 pairs, at least lower ones
opposite; basal pinnae sometimes a little
reduced, always +narrowed at base. Largest
pinnae to 16x3cm, lobed to 1-2 mm from costa;
lobes entire on small fronds, crenate to lobulate
on larger ones; costules almost at right angles to
costae, on larger pinnae S-6 mm apart; veins to 10
pairs, almost all forked, basiscopic branch some-
times forked again; lower surface of rachis and
base of costae glabrous, distally on costae and
costules sparse acicular hairs 0.2 mm long, rarely a
few short capitate hairs; hairs on upper surface of
rachis 0.2 mm long, on costae 0.3 mm, on costules
very short. Sori on acroscopic branches of veins;
indusia thin with capitate hairs on edge.
Distr. Tonkin; Thailand; Malesia: Sumatra,
Malaya, W. Java, Borneo, Luzon, Mindanao, New
Guinea.
Ecol. In Malaya locally abundant at hill sta-
tions on earth banks beside paths in open or
partially shaded places; in exposed places conspic-
uously yellow-green. One plant from Malaya
cultivated at Kew was tetraploid.
Note. Another specimen labelled W. 11 by
BAMLER (collected in 1914) was type of Dry-
opteris distincta COPEL.
4. PHEGOPTERIS
FEE, Gen. Fil. (1852) 242 emend. CHING, Acta Phytotax. Sinica 8 (1963)
312: Ho_trum, Blumea 17 (1969) 9.— Polypodium § Phegopteris PRESL,
Tent. Pterid. (1836) 179, p.p. — Thelypteris subg. Phegopteris sect. Phegop-
teris K. Iwats. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 25 and sect.
Lastrella p.p.
Caudex wide-creeping or short and suberect; fronds pinnate with deeply
lobed pinnae which are connected with each other by a wing along the
rachis, the wing forming, between one pinna and the next, a + semicircular
lobe containing a branched vein arising directly from the rachis; frond-form
either deltoid or (in sole Malesian species) lanceolate with lower pinnae
gradually reduced; veins in pinna-lobes simple or branched, tips of distal
ones only reaching margin; sori subterminal on veins or their branches,
exindusiate or with a very small indusium; sporangia often bearing short
acicular or capitate hairs; lower surface of rachis and costae of pinnae
copiously scaly; scales pale, thin, narrow, with slender spreading marginal
hairs and a hair-tip; larger, darker and often less hairy scales present at
base of stipe; surfaces of frond bearing acicular and capitate hairs.
Type species: Polypodium phegopteris L. (Phegopteris polypodioides FEE).
(ser. II, vol. 1°
354 FLORA MALESIANA
Distr. Three species, one widely distributed in north temperate regions and south to Himalayas, one
in E. North America, one in S.E. Asia.
Cytol. Base chromosome number 30: P. connectilis triploid; P. hexagonoptera diploid; P. decursive-
pinnata diploid and tetraploid.
Notes. FEE and other authors of the 19th century included in this genus free-veined species
resembling Nephrodium but lacking indusia; METTENIUS admitted also species with anastomosing
veins. The result in either case was an unnatural mixture. CHING was the first author to restrict the
genus in the sense here accepted. The species described below differs from the type in elongate fronds
on a suberect caudex.
1. Phegopteris decursive-pinnata (VAN HALL)
FEE, Gen. Fil. (1852) 242; CHING, Acta Phytotax.
Sinica 8 (1963) 312; Mirul, J. Jap. Bot. 40 (1965)
119,124; HoL_truM, Blumea 17 (1969) 11; C. M.
Kuo, Fl. Taiwan 1 (1975) 427, f. 148.— Poly-
podium decursive-pinnatum VAN HALL, Nieuwe
Verh. Kon. Ned. Inst. Wet. 5 (1836) 204; HOoK.
2nd Cent. Ferns (1861) t. 49.— Aspidium decur-
sive-pinnatum (VAN HALL) KUNZE, Bot. Zeit. 6
(1848) 555; MetTr. Fil. Hort. Lips. (1856) 89. —
Nephrodium decursive-pinnatum (VAN HALL)
Hook. in Blakiston, Five Months on the Yangtse
(1862) 365; DIELS in E. & P. Nat. Pfl. Fam. I, 4
(1899) 171.—Leptogramma_ decursive-pinnata
(VAN HALL) J. SM. Hist. Fil. (1875) 232. — Dry-
opteris decursive-pinnata (VAN HALL) O. KTZE
Rev. Gen. PI. 2 (1891) 812; v.A.v.R. Handb. Suppl.
(1917) Corr. 48; Bull. Jard. Bot. Btzg II, 28 (1918)
23. — Thelypteris decursive-pinnata (VAN HALL)
CHING, Bull. Fan Mem. Inst. Bot. 6 (1939) 275;
K. IwaTts. Mem. Coll. Sci. Univ. Kyoto B, 31
(1965) 137.—Type: South China ex Herb. VAN
HALL (L).
Lastrea decurrens J. SM. in Curt. Bot. Mag. 72
(1846) Comp. 32.—Type: CAMERON, China
(BM).
Caudex short, suberect; stipes tufted, 3-10cm
long, closely scaly and hairy throughout; scales at
base firm, brown with copious stiff marginal hairs;
lamina to 50cm long, narrowed gradually to apex
and base; lowest pinnae 10-15 mm long, middle
pinnae 3.5-8cm long, 6-10mm wide, lobed +3
towards costa; veins in lobes pinnate, veinlets
simple; sori subterminal on veinlets, to 8 on larger
lobes; indusium very small, bearing long stiff uni-
cellular hairs; sporangia bearing either acicular or
capitate hairs.
Distr. Central & S.E. China, Tonkin, Taiwan,
Korea; in Malesia recorded from Celebes and
East Java.
Ecol. In Taiwan, at low and median altitudes,
usually on rocks (C. M. KUO).
Notes. At Paris is a specimen of ZOLLINGER
1442 named Asplenium erectum WILLD., ex Herb.
DRAKE, from East Java, ‘““ad pedem Mt Lamon-
gan 1500’, Jan. 1845”. v.A.v.R. records a cul-
tivated plant at Bogor originating from Celebes,
collected there by KOORDERS. These are the only
known records of the occurrence of the species in
Malesia.
A small indusium was first described by J.
SMITH in Lastrea decurrens. CHING and IWAT-
SUKI have stated that the receptacle bears a tuft
of hairs, not an indusium; but there is a distinct
flat structure with hairs on its margin which I
would not call part of the receptacle.
MITUI has recorded diploid and tetraploid con-
ditions in the species, but no-one has distinguished
the two forms from other characters, and I cannot
see clear distinctions among herbarium specimens.
For some years a plant flourished in cultivation
in the Botanic Garden at Penang but did not
reproduce itself from spores. Plants grow well in
the Temperate Fern-House at Kew.
5. CORYPHOPTERIS
Hotttum, Blumea 19 (1971) 33; Blumea 23 (1976) 18-47; Allertonia 1
(1977) 195.— Parathelypteris sect. Melanostipes CHING, Acta Phytotax.
Sinica 8 (1963) 301, p.p. — Fig. ll, 4.
Caudex erect, to c. 30cm tall except in C. inopinata; stipes dark
throughout or paler distally, in a few cases bearing spreading septate hairs
near base, always with rather broad scales which in most species lack
acicular hairs. Lamina commonly 20-40cm long, rarely to 80cm, apex
never pinna-like; lowest pinnae not or little reduced except in C. fasciculata
and C. squamipes, narrowed towards their bases and often wider at their
middle than other pinnae; basal acroscopic lobe often enlarged and dentate
1981] THELYPTERIDACEAE (Holttum) 355
with some forked veins; aerophores at bases of pinnae somewhat swollen;
pinnae always deeply lobed, one or more basal lobes sometimes free; veins
in lobes simple or rarely once forked, both basal veins to edge above base
of sinus; lower surface of rachis, costae, costules and veins always bearing
some reduced scales (smallest filiform); spreading acicular hairs usually
present on lower surface of axes of frond (sometimes replaced by short
capitate hairs), in some cases longer hairs which may be septate; sessile
resinous glands (which may collapse on drying) present on lower surfaces
of many species, in some also on upper surface; costa always grooved on
upper surface; hairs on upper surface of rachis and costae always acicular,
normally unicellular, in a few species septate. Sori usually with rather large
indusia which may be glandular or hairy, distal sori (rarely all) sometimes
asymmetric as in Athyrium; sporangia short-stalked, never with glands or
hairs on body, sometimes with a sessile gland on the stalk; spores usually
pale, translucent, with a + continuous wing and a few cross-wings, in at
least one species with many small wings.
Type species: Coryphopteris viscosa (BAKER) HOLTTUM.
Distr. N.E. India to S. China and Japan; Malesia (except Java and Lesser Sunda Islands), Solomon
Islands, New Caledonia, Fiji, Samoa, Tahiti, Rapa, Marquesas; in all about 47 species.
Cytol. In HoLtTTuM, Ferns of Malaya (1955) p. 624, MANTON recorded n= 32 for Thelypteris
pectiniformis (fig. 8) and 2n=c. 70 for T. viscosa; the latter record was probably made from a plant of
the species later named Coryphopteris arthrotricha HOLTTUM. In June 1980 J. W. GRIMES examined a
plant of C. arthrotricha, from Cameron Highlands, Malaya, in cultivation at Kew, and found n= 66,
indicating a tetraploid with base number 33. This number had not previously been recorded for any
species in the family. There are no other records of observations on the chromosomes of this genus. It
seems possible that the count of n= 32 for C. pectiniformis might have been an error for 33, as the
photograph in fig. 8 shows some overlapping of chromosomes. Counts from other species are needed to
confirm that n = 33 is characteristic of the genus.
Ecol. Plants of this genus occur only in + mossy forest on mountain ridges where the soil is leached
and often peaty; Mrs A. G. PIGGOTT reports a pH of 4.0 for soil on G. Ulu Kali in Malaya (Fern Gaz.
11, 1978, 428). In this habitat there are usually no other thelypteroid ferns.
Notes. The greatest number of species (20) occur in New Guinea, where there is the greatest and
most varied development of suitable mountainous country.
Critical new field work is needed everywhere for a better understanding of this genus; more new
species may remain to be discovered, and some here described need to be more clearly characterized.
The presence and distribution of glands is always a significant character, as in other members of the
family. The glands of Coryphopteris are resinous and sometimes collapse when dried by artificial heat or
when treated with alcohol before drying; in the collapsed state they are often difficult to see.
Among species lacking glands, the only one widely distributed is C. badia. This appears to be peculiar
in growing always in moss-cushions, sometimes on branches of trees. The bases of stipes are covered
with slender hairs like those on roots and probably have the same function.
Species 1-3 have hairs of a different type at the bases of stipes; these are rigid and spreading,
consisting of many cells. Habitat conditions for these species have not been well recorded. One of them
is C. unidentata which has larger fronds than any other in the genus and is only known from three
mountains in the north of Malaya. C. arthrotricha, the common species on the Main Range of Malaya,
has septate hairs on the upper surface of costae, not at the base of stipes; C. tahanensis, only known
from Gunung Tahan in Malaya, is intermediate between C. multisora of Borneo and C. arthrotricha.
The almost invariable miniature arborescent habit of well-grown plants of all species of this genus,
and the fact that they are confined to ridge-tops where other thelypteroid ferns will not grow, suggests
that Coryphopteris may be the relic of a primitive section of the family. The Cyathea-like septate hairs
on upper surface of rachis and costae in C. unidentata, C. hirsutipes, C. multisora, C. tahanensis and C.
arthrotricha may also be a relic of a primitive condition. The only species which show a partially
bipinnate condition are C. habbemensis and C. stereophylla, both much smaller in stature than C.
unidentata.
356 FLORA MALESIANA [ser. II, vol. 1°
Fig. 4. Coryphopteris viscosa (BAK.) HOLTTUM. a. Lower and upper surface of pinna-lobes, showing
glands, hairs and uniseriate scales, x 8.— C. unidentata (BEDD.) HOLTTUM. b. Middle lobes of basal
pinna, x 3. —C. hirsutipes (CLARKE) HOLTTUM. c. Base of stipe, X 8 and part of one hair, x 32. — C.
multisora (C. CHR.) HOLTTUmM. d. Rachis and base of pinna, lower surface, showing septate hairs,
x 8.— C. habbemensis (COPEL.) HOLTTUM. e. One pinna, X 1.5.—C. klossii (RIDL.) HOLTTUM. if
One pinna-lobe, x 6.—C. badia (v.A.v.R.) HOLTTUM. g. Basal pinna, x 1.5: h. costa and part of
costule, showing acicular hairs and narrow scales, x 40 (a PIGGotr 1896, b KUNSTLER 7434, c-d
MOYSEY s.n., e PULLEN 5248, f VINK 17207, g-h MATTHEW s.n.).
1981] THELYPTERIDACEAE (Holttum) 357
KEY TO THE SPECIES
1. Base of stipe bearing pale, firmly cylindrical multiseptate hairs.
2. Stipe and abaxial surface of rachis densely covered with spreading hairs many of which are septate
(seeralsoin. 14))) 29.) ¢ . . . .1.C. unidentata
2. Stipe above base and abaxial surface of rachis less densely hairy with few septate hairs.
3. Indusium glandular; scales at base of stipeto10X*3mm_.._.. WEN! FEA P25 @ multisora
vat Indusia bearing acicular hairs; scales at base of stipe much smaller Hg 1s Ss Co hirsntipes
. Base of stipe not bearing such hairs.
er Sessile glands and/or acicular hairs abundant between veins on upper surface.
5. Scales on lower surface of costae at most 2 cells wide at base.
6. Stipe-scales thin, less than 1 mm wide above base; lower surface of costae bearing many acicular
hairs.
jambiimnae tor-5.cmilong, nearly allidefiexed) 2/2) 25 a an = 2) 24, Co plumosa
7. Pinnae to 5.5 cm long, a few basal ones deflexed.
8. Pinnae to 25 pairs or more; pinna-lobes mostly entire . . . . 5a. C. viscosa var. viscosa
8. Pinnae 12-18 pairs; pinna-lobes mostly crenate . . . . . . .5b.C. viscosa var. poiensis
6. Stipe-scales firm, 1mm or more wide; lower surface of costae bearing short capitate hairs and
glands, acicular hairs lacking or very few.
9. Upper surface of pinnae lacking acicular hairs between veins 6a. C. gymnopoda var. gymnopoda
9. Upper surface of pinnae bearing acicular hairs between veins.
10. Lower surface between veins, and indusia, lacking acicular hairs
6b. C. gymnopoda var. bintangensis
10. Lower surface bearing copious short erect acicular hairs between veins
6c. C. gymnopoda var. humilis
5. Scales on lower surface of costae widened at base.
11. Upper surface of pinnae lacking acicular hairs between veins.
12. Scales on lower surface of costae broadly ovate .. . we EN) SAG oC: Klossik
12. Scales on lower surface of costae linear with a + widened base.
13. Indusia bearing copious acicular hairs; stipe-scales 3x 0.8mm, firm, with many spherical
outgrowths on dorsal surface . . . . 8. C. iwatsukii
- Indusia lacking acicular hairs; stipe- -scales to 7» x1. 5 mm, _ thin, ‘smooth dorsally 9. C. atjehensis
Upper surface of pinnae bearing acicular hairs between veins . . Sf © s105@2 obtusata
4. eae glands and acicular hairs lacking or rare between veins on upper ‘surface of pinnae.
14. Sessile glands present on lower surface generally, or at least on costules.
Pee Som distinctly supramedial’’! 2°, 3 58 5 2 ee et Cadiaphana
15. Sori medial to inframedial.
16. Basal half of lower pinnae bearing free or separately adnate pinnules . . 12. C.habbemensis
16. At most one free pinnule present at base of largest pinnae.
17. Many septate hairs present on upper surface of rachis and costae.
18. Septate hairs on upper surface of rachis and costae less than 0.5 mm long; stipe-scales 3 mm
long met > 2: “135. arthrotricha
18. Septate hairs on upper surface of rachis and costae 1 mm or more long; stipe-scales much
longer.
19. Stipe-scales to 103mm, thin; hairs on upper surface of rachis and costae commonly
1-1.5mmlong . . y Gra, 2. C. multisora
19. Stipe-scales to 7 X 1 mm, tigi Gai pointed: hairs ¢ on upper surface of rachis and costae not
overlImmlong. . . . . . . 14. C. tahanensis
17. Septate hairs absent or rare on upper surface of rachis and costae.
20. Sori mainly along one side of a vein, rarely reniform Ee, ae errors Le 5. C. atnyriomles
20. Sori mainly reniform, distal ones sometimes + athyrioid.
21. All pinna-lobes entire.
22. Pinnae thin, copiously hairy on costae beneath; costal scales few.
23. Hairs on lower surface of costae unicellular opel . . 16. C. pectiniformis
24. Pinnae c. 1.6cm. wide; veins 7-8 pairs; hairs on lower surface of costae over 0.5mm
long es, ¢70- WRN H 16a. C. pectiniformis var. pectiniformis
24. Pinnae c.0.8 cm wide, veins 34 pairs: hairs on lower surface of costae c. 0.1 mm long
16c. C. pectiniformis var. minor
23. Hairs on lower surface of costae to 1.5mm long, septate 16b. C. pectiniformis var. hirsuta
22. Pinnae rigid; costae sparsely hairy beneath and bearing many small scales
17. C. andersonii
358 FLORA MALESIANA [ser. II, vol. 1°
21. At least basal acroscopic lobes of middle pinnae dentate.
25. Lamina to 15 cm long; largest pinnae 3.0x08cm .. . . . . . 18. C. andreae
25. Lamina always much longer; pinnae in most cases to at least 53 x lcm.
26. Lower pinnae, at least 2 pairs, reduced; basal pinnae not wider than next above them.
27. Six or more pairs lower pinnae gradually reduced; lowest 1.5-2 cm long; stipe commonly
to 1Scmlong . . . . . 19. C. squamipes
27. 2-4 pairs lower pinnae reduced, lowest larger: stipe 20-25 ¢ cm.
28. Fertile pinnae to 4.5 X 1.2 cm; stipe-scales to2.5mm wide . . . . . 20.C. borealis
28. Fertile pinnae to 8.5 x 1.7 cm, stipe-scales 1 mm wide . wl. FO PC ametohasis
26. Basal pinnae wider and not shorter than those next above them.
29. Basal acroscopic lobe of basal pinnae free.
30. Glands on lower surface usually confined to costules and veins; no hairs between veins
on upper surface.
31. Lamina to 25cm long; pinnae 12-15 pairs . . . 22a. C. pubirachis var. pubirachis
31. Lamina to 40cm long; pinnae to 20 pairs . . . . . 22b.C. pubirachis var. major
30. Glands present between veins on lower surface and hairs between veins on upper
surface.
32. Basal pinnae to 9 x 2cm; most pinna-lobes almost entire
22c. C. pubirachis var. philippinensis
32. Basal pinnae to 6 x 1.2 cm; most pinna-lobes dentate or crenate
22d. C. pubirachis var. sulawesica
29. Basal acroscopic lobe of basal pinnae not free.
33. Stipe-scales less than 1 mm wide above dilated base.
34. Pinnae to 18 pairs; lower surface of rachis bearing capitate hairs only; many capitate
hairs between veins on upper surface ae . . . . 23. C. athyriocarpa
34. Pinnae to 25 pairs; lower surface of pinna- rachis bearing acicular hairs; no hairs
between veins on upper surface at: . . . . . 5e.C. viscosa var. borneensis
33. Stipe scales at least 1 mm wide above base.
35. Pinna-lobes, except basal one, entire; glands present between veins on lower surface
17. C. andersonii
35. Pinna-lobes all distinctly dentate; no glands between veins on lower surface
24. C. tanggamensis
14. Sessile glands lacking on lower surface except in C. fasciculata which may have a few glands on
costae.
365 Indusiajlacking: “aieiee awe 40 So ais ewes a ea ee ee, ees Crenslenana
36. Indusia present.
37. Several pairs of free pinnules on lower pinnae 7 5 2 « wow & ve 3 | a26Rn@Sstereophylla
37. At most basal lobes free.
38. Upper surface bearing hairs between veins.
39. Hairs on upper surface between veins acicular.
40 uStipeibearineicopiousshairssl mmilong)) 4) a 29682) See 27 ahonizontalis
40. Stipe bearing hairs 0.5mm long in groove only . . . . . . . . 28.C.microlepigera
39. Hairs on upper surface between veins capitate cule aeihe 2. Wee". Zoe propria
38. Upper surface between veins normally glabrous.
41. Lower surface of rachis bearing acicular hairs, costae usually also.
42. Some hairs on upper surface of costae septate oss Sw eee ORCA lantenhachht
42. Hairs on upper surface of costae unicellular.
43. Several pairs lower pinnae gradually reduced, longest 5.5 cm, lowest 1-2 cm
31. C. fasciculata
43. Lower pinnae not or little reduced.
44. Texture rigid; pinnae to 3 cm long; hairs on lower surface of rachis 0.5 mm long
32. C. hubrechtensis
44. Texture thin; pinnae to 6.5 cm long; hairs on lower surface of rachis 0.2 mm long
33. C. brevipilosa
41. Lower surface of rachis and costae lacking (or almost lacking) acicular hairs.
45. Short capitate hairs present on lower surface of costae and costules and on indusia
34. C. oligolepia
45. Capitate hairs lacking or rare on lower surface and indusia.
46. Sori supramedial; stipe-scales 8-10mm long . . eee Ge PES S54C> conacea
46. Sori medial or inframedial; stipe-scales not over 5mm long.
47. Sori mostly athyrioid; pinnae to 2.5 cm long fee PEO ee 36: Caledenmannnr
1981]
THELYPTERIDACEAE (Holttum)
359
47. Sori mostly not athyrioid; pinnae longer.
48. Caudex erect.
49. Pinnae thin; no tangled hairs at base of stipe.
50. Veins 3-4 pairs; pinna-lobes entire except basal acroscopic ones
37. C. dura
50. Veins c. 6 pairs; pinna-lobes mostly crenate.
51. Pinnae caudate-acuminate
51. Pinnae short-acuminate
49. Pinnae thick; tangled hairs often present at base of stipe
48. Caudex slender, long-creeping
1. Coryphopteris unidentata (BEDD.) HOLTTUM,
Blumea 23 (1976) 26. — Lastrea unidentata BEDD.
Handb. Suppl. (1892) 53. — Dryopteris monodonta
C. CHR. Ind. Fil. (1905) 278, nom. nov.; C. CHR.
Gard. Bull. Str. Settl. 4 (1929) 388. — Thelypteris
unidentata (BEDD.) HOLTTUM, Rev. FI. Malaya 2
(1955) 251.—Type: KUNSTLER 7434, Gunung
Bubu, Perak (K). — Fig. 4b.
Stipe to 60 cm long, bearing throughout spread-
ing septate hairs | mm or more long; basal scales
10x 1.5mm. Lamina to 80cm long; pinnae c. 25
pairs; basal pinnae narrowed at base, basal acros-
copic lobe enlarged, dentate, free, some other
lobes with a single tooth at basiscopic base; lar-
gest pinnae 18.5 x2.6cm, sessile, lobed to 1mm
from costa, lobes entire except basal ones; cos-
tules to 5mm apart; veins to 10 pairs; lower
surface of rachis and costae bearing septate hairs;
sessile glands present on costae, costules and
veins, fewer between veins, narrow scales on
costae and costules; upper surface of costae bear-
ing septate hairs, few on costules, no glands. Sori
medial; indusia bearing short capitate hairs or
glands.
Distr. Malesia: Malaya. Only known from G.
Bubu, G. Bintang and G. Inas in Perak.
Note. The presence of a single large tooth at
the basiscopic base of lobes of basal pinnae,
which is denoted by the name unidentata, occurs
only on the type collection. The others are smaller
(the smallest frond has pinnae to 12 x 1.8cm) but
otherwise do not differ. The septate hairs at base
of stipe are shorter than those on C. hirsutipes.
2. Coryphopteris multisora (C. CHR.) HOLTTUM,
Blumea 23 (1976) 26.— Dryopteris multisora C.
Cur. Gard. Bull. Str. Settl. 7 (1934) 241. — Thelyp-
teris multisora (C. CHR.) REED, Phytologia 17
(1968) 295. — Type: HOLTTUM 25523, Sabah, Mt
Kinabalu 2100 m (BM; BO, SING). — Fig. 4d.
Stipe to 45cm, dark bearing sessile glands
throughout and long septate hairs near base, also
distally in the groove; scales thin, to 103mm.
Lamina to 55cm long; pinnae to 25 pairs; basal
pinnae to 2.1 cm wide in middle, narrowed to base
with basal acroscopic lobe enlarged, dentate to
deeply lobed, almost free, rarely to 15 mm long.
Pinnae above base to 11X1.8cm, acuminate;
lobes not falcate, entire or nearly so; costules
3.5-4 mm apart; veins 6—7 pairs; glands present on
38. C. platyptera
. 39. C. subnigra
. 40. C. badia
41. C. inopinata
lower surface of rachis, costae and costules,
sparse septate hairs on rachis and costae (some-
times absent), many very narrow scales on costae
and costules; upper surface of rachis bearing sep-
tate hairs 1-1.5 mm long, similar but shorter hairs
on costae and costules; sometimes a few glands
on costae and costules. Sori large, somewhat in-
framedial, filling lower surface at maturity; indusia
glandular.
Distr. Malesia: Sabah (Mt Kinabalu), Sarawak
(G. Mulu), 1350-3000 m, several collections.
Note. Long septate hairs are not present at the
base of stipes in all specimens; the species there-
fore appears in two places in the key. CHRIS-
TENSEN wrongly stated that glands are absent
from lower surface of rachis and costae.
3. Coryphopteris hirsutipes (CLARKE) HOLTTUM
in Nayar & Kaur, Comp. to Bedd. (1974) 203;
Blumea 23 (1976) 27. — Nephrodium gracilescens
var. hirsutipes CLARKE, Trans. Linn. Soc. II Bot.
1 (1880) 514, t. 67, f. 1.—Lastrea gracilescens
sensu BEDD. Handb. (1883) 234, p.p.— Lastrea
hirsutipes (CLARKE) BEDD. Handb. Suppl. (1892)
52, excl. var. didymochlaenoides. — Thelypteris
hirsutipes (CLARKE) CHING, Bull. Fan Mem.
Inst. Biol. Bot. 6 (1936) 314.— Parathelypteris
hirsutipes (CLARKE) CHING, Acta Phytotax.
Sinica 8 (1963) 303.— Lectotype (HOLTTUM
1976): CLARKE 18968, Assam, Khasya Hills
1400 m (K).
Dryopteris indochinensis CHRIST in Morot J.
Bot. 21 (1908) 263.— Thelypteris indochinensis
(CHRIST) CHING, Bull. Fan Mem. Inst. Biol. Bot.
6 (1936) 327; TARD. & C. CHR. FI. Gén. I.-C 7, pt.
2 (1941) 361, f. 43, 1-2.— Parathelypteris in-
dochinensis (CHRIST) CHING, Acta Phytotax.
Sinica 8 (1963) 304.— Type: EBERHARDT, Ton-
kin, Massif du Tam Dao, 900 m (P).
Dryopteris gracilescens (Bl.) O. KTZE_ var.
chinensis CHRIST, Not. Syst. 1 (1909) 40. — Type:
HENRY 10111, Yunnan (P; K).
Dryopteris megalocarpa v.A.v.R. Bull. Jard.
Bot. Btzg III, 5 (1922) 199. — Thelypteris megalo-
carpa (v.A.v.R.) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 252. —- Type: LORZING 7134,
Sumatra, Patjoer-batoe near Lake Toba (BO).
Thelypteris angulariloba CHING, Bull. Fan
Mem. Inst. Biol. Bot. 6 (1936) 323; K. IWATs.
Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 160. —
360
Parathelypteris angulariloba (CHING) CHING,
Acta Phytotax. Sinica 8 (1963) 304; C. M. Kuo,
Fl. Taiwan 1 (1975) 421, pl. 145. — Wagneriopteris
angulariloba (CHING) LOVE & LOVE, Taxon 26
(1977) 325. — Type: N. K. CHUN 42644, Kwang-
tung (PE, not seen).
Thelypteris simozawae TAGAWA, Acta Phy-
totax. Geobot. 6 (1937) 157.— Parathelypteris
simozawae CHING, Acta Phytotax. Sinica 8 (1963)
304.— Type: SIMOZAWA, Taiwan, 17 Oct. 1936
(KYO, not seen).
Thelypteris herbacea HOLTTUM, Gard. Bull.
Sing. 11 (1947) 268; Rev. Fl. Mal. 2 (1955) 254, f.
145.— Type: HoLTruM 20571, Malaya, Gunung
Tahan, 915 m (SING; K). — Fig. 4c.
Stipe 20-30(-45)cm long, dark at base only,
distal part and rachis pale reddish, at base spread-
ing septate hairs 1-3mm long; scales small,
setose. Lamina 25-35 cm long, pinnae 15-20 pairs;
basal pinnae narrowed near base, basal acroscopic
lobe almost free but not elongate; texture thin.
Largest pinnae commonly 7x1.5cm (to 10x
2cm), acuminate, lobes oblong, entire or some-
times slightly toothed at ends of distal veins;
costulés 3.5-4.5 mm apart; veins 4-6 pairs; lower
surface of rachis and costae bearing copious aci-
cular hairs which are sometimes unicellular (types
of L. hirsutipes, D. megalocarpa, T. herbacea) but
sometimes few or many are elongate and septate;
glands on lower surface absent or few (types of L.
hirsutipes and T. herbacea), rarely abundant;
some hairs on upper surface of rachis and costae
always septate, slender unicellular hairs often
present between veins. Sori medial or inframedial;
indusia hairy, hairs sometimes septate.
Distr. S. Japan, Ryukyu Islands, Taiwan, S.
China to N.E. India, Thailand; in Malesia:
Malaya, Sumatra.
Notes. IWATSUKI discussed the variability of
this species under T. angulariloba. I have not seen
CHING’s type of this, but at Kew are two speci-
mens from Hong Kong cited by him; these and
others from Hong Kong and Kwangtung show
much variability in abundance of long septate
hairs on lower surface, though these hairs are
always less abundant than in the type of D. in-
dochinensis. CHING stated that the type of T.
angulariloba lacked glands, but some Hong Kong
specimens (including one cited by CHING) have a
few. Sumatran specimens are varied both in
presence of glands and of septate hairs on lower
surface.
4. Coryphopteris plumosa (C. CHR.) HOLTTUM,
Blumea 23 (1976) 28.— Dryopteris plumosa C.
CHR. Dansk Bot. Ark. 9, 3 (1937) 65.— Thelyp-
teris plumosa (C. CHR.) REED, Phytologia 17
(1968) 305.— Type: MJOBERG 7, Sarawak, Mt
Murud 2700 m (BM).
Stipe 6-10cm, base very dark with copious
reddish firm scales 7x 0.7mm, upper part paler
FLORA MALESIANA
(ser. II, vol. 1°
and finely short-hairy; rachis almost stramineous.
Lamina 20cm long, pinnae nearly 30 pairs, nearly
all deflexed, middle ones largest; basal pinnae
2cm long, slightly narrowed on basiscopic base,
basal acroscopic lobe a little enlarged, dentate and
free, texture firm. Largest pinnae 2.5 0.8cm,
lobes entire; costules 2mm apart; veins 3-4 pairs,
pale and prominent both sides; lower surface of
rachis and costae bearing pale acicular hairs more
than 0.5 mm long, small capitate hairs present on
costae, costules and veins, some glands between
veins, a few very narrow scales on costae; upper
surface of costae hairy as lower, veins and cos-
tules with small capitate hairs, glands throughout.
Sori supramedial; indusia small, thin, with capi-
tate hairs.
Distr. Malesia: Borneo (Sarawak), only known
from the type.
5. Coryphopteris viscosa (BAK.) HOLTTUM,
Blumea 19 (1971) 33; Blumea 23 (1976) 29.—
Lastrea viscosa J. SM. in Hook. J. Bot. 3 (1841)
412, nom. nud. — Nephrodium calcaratum (BL.)
HooK. Spec. Fil. 4 (1862) 93, var. B tantum. —
Nephrodium viscosum BAK. Syn. Fil. (1867) 264,
excl. plant. Philip.—Lastrea viscosa (BAK.)
BEDD. Ferns Br. India (1870) t. 334; Handb. (1883)
238; RIDL. J. Mal. Br. R. As. Soc. 4 (1926) 65,
p.p.; COPEL. Fern FI. Philip (1960) 324, excl. plant.
Philip. — Dryopteris viscosa (BAK.) O. KTZE,
Rev. Gen. Pl. 2 (1891) 814; v.A.v.R. Handb. (1908)
186, p.p.; C. CHR. Gard. Bull. Str. Settl. 4 (1929)
380, p.p.; ibid. 7 (1934) 240, p.p.— Thelypteris
viscosa (BAK.) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 215; HOLTTUM, Rev. FI. Mal.
2 (1955) 252, p.p. — Parathelypteris viscosa (BAK.)
CHING, Acta Phytotax. Sinica 8 (1963) 304.—
Type: CUMING 401, Malacca, Mt Ophir (K;
BM). — Fig. ll, 4a.
a. var. viscosa
Stipe 15-20cm long, dark at base, paler up-
wards, basal scales 40.5 mm, thin; rachis dull
reddish throughout. Lamina to 30cm long, taper-
ing very gradually distally, thin, pinnae 25 pairs
or more, closely placed, a few lower ones
deflexed; basal pinnae to 1.4 cm wide, narrowed to
base, basal acroscopic lobe little or not dentate,
not free. Largest pinnae 5.5 X 1.2 cm, sessile; base
truncate with acroscopic lobe sometimes a little
elongate; apex short-pointed, obtuse; lobes mostly
entire; costules 2.5mm apart; veins to 6 pairs;
lower surface of rachis and costae bearing copious
acicular hairs 0.5mm long with some short capi-
tate hairs and glands; scales on costae 1-2 cells
wide; glands abundant between veins; glands
throughout upper surface. Sori medial, distal ones
+ athyrioid; indusia thin, with a few glands.
Distr. Malesia: Malaya (Mt Ophir), Borneo
(Sarawak: Mt Poi).
1981]
b. var. poiensis HOLTTUM, Blumea 23 (1976)
29. — Type: BURTT & WOODS 2828, Poi Range (K).
Pinnae 12-18 pairs, pinna-lobes mostly crenate.
Distr. Malesia: Borneo (Sarawak: Poi Ra.),
several collections.
c. var. borneensis HOLTTUM, Blumea 23 (1976)
29. — Type: RICHARDS 1702, Sarawak, Mt Dulit
(K).
Upper surface of pinnae lacking glands.
Distr. Malesia: Borneo (Sarawak: Mt Dulit,
several collections; G. Mulu; Mt Penrissen; Kal-
imantan: Mt Bengkaram).
6. Coryphopteris gymnopoda (BAK.) HOLTTUM,
Blumea 23 (1976) 29. — Nephrodium gymnopodum
BAK. Trans. Linn. Soc. Bot. 4 (1894) 252. — Dry-
opteris gymnopoda (BAK.) C. CHR. Ind. Fil. (1905)
269; Gard. Bull. Str. Settl. 7 (1934) 240. — Type:
HAVILAND 1486, Mt Kinabalu 3200 m (K).
Lastrea ridleyi BEDD. Kew Bull. (1909) 423. —
Dryopteris ridleyi (BEDD.) C. CHR. Ind. Fil.
Suppl. (1913) 38. — Type: RIDLEY 7849, Selangor,
Bukit Hitam, 1000 m (K).
Dryopteris subviscosa v.A.v.R. Bull. Jard. Bot.
Btzg II, 26 (1915) 14; Handb. Suppl. (1917) 153. —
Type: BECCARI 429, Sumatra, G. Singgalang
1700 m (BO; FI, K).
Dryopteris kinabaluensis COPEL. Philip. J. Sci.
12 (1917) Bot. 55.—Type: ToPpPING 1719, Mt
Kinabalu (Am. Fern Soc. Ann Arbor, not seen).
Lastrea robinsonii RIDL. J. Fed. Mal. St. Mus.
10 (1920) 156; J. Mal. Br. R. As. Soc. 4 (1926) 65,
p.p.— Dryopteris robinsonii (RIDL.) C. CHR.
Gard. Bull. Str. Settl. 4 (1929) 381. — Thelypteris
robinsonii (RIDL.) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 254. Type: F. M. Mus. Col-
lector, Perak, G. Kerbau (K).
Dryopteris viscosa var. kamborangana C. CHR.
Gard. Bull. Str. Settl. 7 (1934) 240. — HoLTTUM
25472, Mt Kinabalu 2130 m (BM; BO, K, SING).
Thelypteris viscosa sensu HOLTTUM, Rev. FI.
Mal. 2 (1955) 252, p.p.
a. var. gymnopoda
Stipe 15-20cm long; scales 6-8 x 1-1.5 mm,
dark, firm, bases not dilated. Lamina 25-30 cm
long; pinnae 15-18 (rarely to 25) pairs, texture
firm, well spaced; lowest pinnae wider than next,
in largest fronds more than 2 cm wide with almost
all lobes dentate, in smaller fronds lobes crenate.
Suprabasal pinnae commonly 6X 1.5 cm, on lar-
gest fronds 102cm, acuminate to subcaudate,
lobes mostly crenate; costules 3-3.5mm apart;
veins 6—7(—9) pairs; lower surface of rachis bear-
ing some acicular hairs, of costae bearing many
glands and capitate hairs, sometimes a few aci-
cular hairs, scales all linear, 1-2 cells wide, glands
present throughout lower surface; acicular hairs
on upper surface of rachis and costae 0.5mm
long, few on costules and veins distally; abundant
THELYPTERIDACEAE (Holttum)
361
glands throughout. Sori medial, basal ones some-
times a little elongate and asymmetric, distal ones
rarely athyrioid; indusia bearing short capitate
hairs and glands.
Distr. Peninsular Thailand, in Malesia: Sabah
(Mt Kinabalu, many collections), Sarawak; in
Malaya on Gunung Tahan and at scattered locali-
ties on the Main Range, perhaps only in rather
open places; Sumatra, Peninsular Thailand.
Notes. The type is a single small frond which
has lost almost all its scales but owing to the many
other collections its identity is not in doubt.
Peninsular specimens have narrower scales than
Bornean ones. Lastrea ridleyi and L. robinsonii
appear to differ only in their small size.
b. var. bintangensis HOLTTUM, Blumea 23 (1976)
30. — Type: C. B. KLoss, G. Bintang on Kedah-
Perak boundary, June 1917 (K).
Upper surface between veins bearing a variable
number of appressed acicular hairs 0.3-0.5 mm
long.
c. var. humilis HOLTTUM, Blumea 23 (1976) 30. —
Type: MOLESWORTH ALLEN 1026, Pahang,
Cameron Highlands, Gunung Perdah, 2130 m (K).
Fronds small; upper and lower surfaces both
covered with acicular hairs between veins.
Note. The type and another collection have
pinnae 2.5 X0.6cm; small size and hairiness may
be due to exposed position.
7. Coryphopteris klossii (RIDL.) HOLTTUM, Blu-
mea 23 (1976) 31. — Lastrea klossii RIDL. Trans.
Linn. Soc. Bot. 9 (1916) 257; COPEL. Philip. J.
Sci. 78 (1951) 428.— Dryopteris klossii (RIDL.)
v.A.v.R. Handb. Suppl. (1917) 501. — Thelypteris
klossii (RIDL.) CHING, Bull. Fan Mem. Inst. Biol.
Bot. 10 (1941) 252. — Type: C. B. KLoss, W. New
Guinea, Wollaston Exped. to Mt Carstensz, Camp
Vic, 1800 m (BM; K). — Fig. 4f.
Stipe 20-30cm long, dark, glandular but not
hairy except in the groove; scales 3-4 x 1-1.5 mm.
Lamina to 30cm or more long; pinnae c. 20 pairs
separated by half their width; basal pinnae a little
reduced or not, narrowed in basal third, basal
acroscopic lobe a little enlarged and strongly
crenate; texture rather rigid when dry. Largest
pinnae of type 3.3 x 1.0cm, of another specimen
6 X 1.3. cm; lobes slightly falcate, edges sinuate to
toothed at ends of distal veins; costules 2.5-
3.5mm apart; veins 5-7 pairs, thick and slightly
prominent; lower surface of rachis and costae
bearing rather sparse acicular hairs, many ovate
scales to 1 mm long on costae, smaller scales on
costules, glands generally; upper surface of rachis
and costae bearing short acicular hairs, glands and
capitate hairs on surface between veins. Sori
medial; indusia large, thin, bearing capitate hairs
and glands, usually all symmetric.
Distr. Malesia: Throughout New Guinea at
1800-3200 m.
362
8. Coryphopteris iwatsukii HOLTTUM, Blumea 23
(1976) 31.—Type: K. IWATSUKI et al. S. 1012,
Sumatra, Atjeh (K).
Caudex slender; stipe to 20cm long, very dark
at base, minutely hairy; scales rigid, 3 mm long,
0.8 mm wide at base, dorsally with small spherical
outgrowths. Lamina 25cm long, texture thin;
pinnae 15 pairs; lowest pinnae narrowed at base,
basal acroscopic lobe free, dentate, not elongate.
Suprabasal pinnae to 6.0 1.4cm, acuminate;
lobes at right angles to costa, almost all dentate,
separated by sinuses 1.5mm _ wide; costules
3.5mm apart; veins 5-6 pairs; lower surface of
rachis with hairs 0.5mm long, of costae with
sparse hairs and many glands, larger scales on
costae dilated at base, lamina between veins bear-
ing glands and minute erect hairs; upper surface of
rachis and costae bearing unicellular acicular
hairs, few on costules and veins, surface between
veins with many glands. Sori medial; indusia thin,
bearing very short acicular hairs.
Distr. Malesia: N. Sumatra (Gajolands: Mt
Kemiri), only known from the type.
Ecol. Mossy forest, 1800-2500 m.
9. Coryphopteris atjehensis HOLTTUM, Blumea 23
(1976) 32.— Type: K. IWATSUKI et al. S. 834,
Sumatra, Atjeh, G. Kemiri, 900-1600 m, in ever-
green forest (K).
Differs from C. iwatsukii as follows: scales on
stipe 7 x 1.5mm, smooth dorsally; pinnae to 9.5 x
1.9cm, pinna-lobes mostly entire; costules 4—
4.5mm apart; indusia glabrous or with minute
capitate hairs.
Distr. Malesia: N. Sumatra (Gajolands, Mt
Kemiri), known only from the type and DE
WILDE 13051.
10. Coryphopteris obtusata (v.A.v.R.) HOLTTUM,
Blumea 23 (1976) 30.—Dryopteris obtusata
v.A.v.R. Bull. Jard. Bot. Btzg I, 28 (1918) 22. —
Thelypteris obtusata (v.A.v.R.j CHING, Bull. Fan
Mem. Inst. Biol. Bot. 10 (1941) 253. — Type: C. J.
BRooKS 339 S, Sumatra, Benkoelen, Lebong
Simpang (BO; BM).
Dryopteris supravillosa C. CHR. Gard. Bull. Str.
Settl. 7 (1934) 241. — Type: HOLTTUM 25471, Mt
Kinabalu, 1800 m (BM; K, SING).
Stipe 15-25 cm long, dark at base, short-hairy,
upper part and rachis dull reddish, scales to 5 x
1.5mm. Lamina 20-35cm long, pinnae 15-18
pairs; basal pinnae somewhat reduced on smaller
fronds, narrowed at base, basal acroscopic lobe
dentate, not free. Largest pinnae to 7.5 x 1.5cm,
short-acuminate; basal lobes dentate, others
sometimes with slight teeth at ends of distal veins;
costules 3-4 mm apart; veins 5-8 pairs; lower sur-
face of rachis densely hairy, of costae less so,
glands present on all parts of lower surface, those
between veins sometimes small (type) or replaced
by capitate hairs; scales on lower surface of cos-
FLORA MALESIANA
{ser. II, vol. 1°
tae widened at base; upper surface densely
covered with short acicular hairs, some capitate
hairs or glands also present. Sori medial to in-
framedial, distal ones sometimes asymmetric
where small; indusia bearing glands and capitate
hairs.
Distr. Malesia: Sumatra, Borneo, New Guinea.
Note. There is variability in Sumatran speci-
mens in abundance of glands or capitate hairs on
lower surface of pinnae; a specimen from W. New
Guinea is very like the type. Specimens from
eastern New Guinea have rather few glands and
capitate hairs. The type of D. supravillosa is small
(lamina 20 cm long) and has capitate hairs between
veins on lower surface.
11. Coryphopteris diaphana (BRAUSE) HOLT-
TUM, Blumea 23 (1976) 32. — Dryopteris diaphana
BRAUSE, Bot. Jahrb. 56 (1920) 80. — Thelypteris
diaphana (BRAUSE) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 251.— Type: LEDERMANN
8903, N.E. New Guinea, Sepik Distr. (B; BM).
Stipe 10-15cm long, dark, glossy, sparsely
hairy; scales 2-3 x 0.8 mm, dark, rigid. Lamina to
25cm long; pinnae 15-18 pairs; basal pinnae not
or little reduced, a little narrowed at base, basal
acroscopic lobe almost free, dentate, not elongate.
Suprabasal pinnae to 4.5 1.2cm, short-acu-
minate; lobes oblique, basal ones strongly dentate,
rest + toothed at ends of veins; costules 2.5-3 mm
apart; veins 5-6 pairs; lower surface of rachis and
costae with short spreading acicular hairs, glands
on costae, costules and veins, a few dark narrow
scales on costae; upper surface of costules and
veins with scattered hairs, no glands nor hairs
between veins. Sori supramedial; indusia
glabrous. :
Distr. Malesia: New Guinea. Known only
from type collection and BRASS 13294 from
Idenburg River.
Ecol. Type from 850m, in moss cushions;
BRASS 13294 from 900 m, in Agathis forest.
12. Coryphopteris habbemensis (COPEL.) HOLT-
TUM, Blumea 23 (1976) 32.— Dryopteris hab-
bemensis COPEL. Un. Cal. Publ. Bot. 18 (1942)
216.— Ctenitis habbemensis (COPEL.) COPEL.
Gen. Fil. (1947) 124; Philip. J. Sci. 78 (1951) 411,
pl. 15.— Type: BRASS 9304, N.W. New Guinea,
Lake Habbema 3225 m (MICH; BM). — Fig. 4e.
Stipe 25-30cm long, base dark with short
brown hairs, reddish distally; scales thin, 5x
1.5mm. Lamina to 30cm long; pinnae 18 pairs;
basal pinnae not much narrowed at base. Largest
pinnae 72cm, pinnate in basal half; 2-3 pairs
pinnules quite free, rest + adnate to costa, lobes
of distal half of pinna connected by a very narrow
wing; pinnules 7-12 mm long, 2-3 mm wide, larger
ones deeply lobed near base and crenate distally;
costules 3.5-4 mm apart; veins 5-8 pairs, forked in
basal lobes of larger pinnules; lower surface of
1981]
THELYPTERIDACEAE (Holttum)
363
rachis and costae copiously short-hairy, also on
costae thin clathrate scales, the larger ones ovate;
some sessile glands and smaller scales on cos-
tules; acicular hairs on upper surface of costae
and costules, no glands. Sori medial; indusia thin,
glabrous.
Distr. Malesia: Middle to east of New Guinea
at 1800-3200 m.
Note. See note on 26. C. stereophylla.
13. Coryphopteris arthrotricha HOLTTUM, Blu-
mea 23 (1976) 33.—Type: HOLTTUM 23345,
Pahang, Cameron Highlands (K).
C. viscosa sensu HOLTTUM, Rev. Fl. Malaya 2
(1955) 252, p.p.
Stipe to 30cm long, dark at base, paler distally,
hairy only in groove, basal scales 3 mm long, thin.
Lamina 25-45cm long; pinnae 20 pairs, well
spaced, lowest much narrowed at base with basal
acroscopic lobe almost free but not elongate.
Suprabasal pinnae commonly 8 x 1.6cm, shortly
caudate-acuminate; lobes entire to dentate; cos-
tules 3.5-4 mm apart; veins 6-10 pairs; lower sur-
face of rachis and costae bearing minute acicular
and capitate hairs, on costae also glands and linear
scales, few glands between veins; upper surface
of rachis and costae bearing hairs hardly 0.5 mm
long which have 1-3 septa, surface between veins
glabrous or with a few short acicular and capitate
hairs. Sori medial; indusia small, bearing short
capitate hairs and sometimes glands.
Distr. Malesia: Malaya, Sumatra, in ridge
forest at 1220-1520 m. This is the common species
on the Main Range in Malaya.
14. Coryphopteris tahanensis HOLTTUM, Blumea
23 (1976) 33. — Type: HOLTTUM 20694, Pahang,
Gunung Tahan (K; SING).
Differs from C. arthrotricha: scales at base of
stipe 7-8 x | mm, rigid, acuminate; lobes of pin-
nae, except basal ones, almost entire; rachis and
costae of apical part of frond sometimes bearing
multicellular hairs 1.5mm long on lower surface,
on upper surface septate hairs always 1 mm long;
glands on costae in some cases rare.
Distr. Malesia: Malaya. Three collections from
1800 m on G. Tahan and two from Main Range.
Note. This is intermediate between C. arth-
rotricha and C. multisora. One specimen from G.
Tahan has the lower surface of rachis and costae
densely covered with hairs almost 2 mm long, and
a few on costules and veins.
15. Coryphopteris athyrioides HOLTTUM, Blumea
23 (1976) 33.—Type: BRASS 24722, Papua,
Goodenough Island, on mossy rock (BM; L,
LAE).
Stipe to 20cm long, black at base, reddish up-
wards, short-hairy throughout, basal scales thin,
ovate, 3-4 1.5mm. Lamina 30cm long; pinnae
18-20 pairs; lowest pinnae sometimes a little
reduced, basal acroscopic lobe not free. Largest
pinnae 6.7 X 1.7 cm, short-acuminate; lobes falcate
and slightly toothed; costules to 4mm apart; veins
7-8 pairs, prominent on lower surface; rachis and
costae on lower surface densely covered with
erect acicular hairs and filiform scales, on rest of
surface red glands; no glands on upper surface.
Sori asplenioid, occupying almost whole length of
all veins, some of them hooked as in Athyrium,
rarely reniform; indusia bearing glands and some-
times a few short acicular hairs.
Distr. Malesia: Papua New Guinea
(Goodenough I.), only known from the type.
16. Coryphopteris pectiniformis (C. CHR.) HOLT-
TUM, Webbia 30 (1976) 20; Blumea 23 (1976)
34. — Dryopteris pectiniformis C. CHR. Gard.
Bull. Str. Settl. 4 (1929) 379. — Thelypteris pec-
tiniformis (C. CHR.) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 253; HOLTTUM, Rev. FI. Mal.
2 (1955) 253, f. 144, 624, pl. 1 f. 8. — Parathelyp-
teris pectiniformis (C. CHR.) CHING, Acta Phy-
totax. Sinica 8 (1963) 304.— Type: G. F. HOSE
293™Peraka (P= sKs):
Thelypteris subglanduligera CHING, Bull. Fan
Mem. Inst. Biol. Bot. 6 (1936) 323. — Type: WRAY
367, Perak, Gunung Batu Puteh (US; CAL, L).
a. var. pectiniformis
Stipe dark at base with thin setiferous scales to
7x1mm,_ distally stramineous, throughout
covered with short unicellular hairs. Lamina 40-
45 cm long, texture thin; pinnae 15-20 pairs, well-
spaced; basal pinnae narrowed near base, basal
acroscopic lobe not enlarged, sometimes a little
dentate. Suprabasal pinnae to 10x 1.6cm, acu-
minate; lobes entire, slightly falcate; costules to
3mm apart; veins 7-8 pairs, very oblique; lower
surface of rachis, costae and costules bearing
short pale acicular hairs, glands and short erect
acicular and capitate hairs on surface between
veins; upper surface of costae bearing copious
pale acicular hairs more than 0.5mm long, few
hairs on costules and veins, no other hairs. Sori
medial; indusia with abundant capitate hairs,
sometimes a few acicular hairs.
Distr. Malesia: Malaya (Taiping Hills, a few
records from Main Range, G. Padang in Treng-
ganu).
b. var. hirsuta HOLTTUM, Blumea 23 (1976) 34. —
Type: HoLtruM 21547, Pahang, Pine-tree hill,
near Fraser’s Hill (K; SING).
Differs from typical variety: smaller (pinnae to
6.5cm long); lower surface of costules and veins
bearing acicular hairs 1.5mm long which are
septate; indusia bearing acicular hairs 0.5mm
long.
Distr. Malesia: Malaya (only known from
Pine-tree hill and the ridge connecting it to
Fraser’s Hill).
364
c. var. minor HOLTTUM, var. nov.
A typo speciei differt: pinnis minoribus (usque
4.2*0.8 cm); venis 3-jugatis; rhachidi et costis
subtus pilis 0.1mm longis vestitis; paleis stipitis
3 mm longis, 0.8 mm latis. Type: J. WINKLER S.n.
1912, Sumatra, Pea Radja, in Rosenst. Fil. sumatr.
exsicc. n. 143 (P).
Distr. Malesia: Sumatra.
Note. This was distributed as Dryopteris sub-
viscosa ROSENST., a name not published (not D.
subviscosa v.A.v.R.). Another specimen (n. 154a)
from the same locality, distributed as D. japonica
var. stricta, has somewhat larger pinnae (MICH).
The short pinna-lobes have broad tips, so that the
pinnae are much like those of C. hirsutipes in
aspect, but in other characters they agree with C.
pectiniformis.
17. Coryphopteris andersonii HOLTTUM, Blumea
23 (1976) 38.— Type: J. A. R. ANDERSON 4535,
Sarawak, Baram Distr., G. Mulu 2000 m (L; K).
Caudex to 30cm tall; stipes 20cm long, dark
castaneous at base, paler distally, short-hairy,
basal scales 4-5 1.5mm; rachis stramineous.
Lamina to 25cm long, subcoriaceous, rigid when
dry; pinnae 15 pairs, lowest not reduced but
slightly narrowed at base with basal acroscopic
lobe sometimes dentate, not free. Suprabasal
pinnae to 5X1.2cm, short-acuminate; lobes
oblique, hardly falcate, with rounded tips, basal
ones at most slightly dentate; costules 2mm or
little more apart; veins 6 pairs, very oblique, pale
and prominent on both surfaces; lower surface of
rachis covered with hairs 0.3mm long; hairs on
costae few or none, scales on costae narrowly
linear or with dilated bases, glands present on and
between veins; hairs on upper surface of costae
0.5 mm long, no other hairs nor glands on upper
surface. Sori medial, symmetric; indusia thin,
glabrous.
Distr. Malesia: Borneo (Sarawak: Mt Mulu),
only known from type.
18. Coryphopteris andreae HOLTTUM, Blumea 23
(1976) 35.—Type: MILLAR & HOLTTUM NGF
15768, N.E. New Guinea, Morobe Distr., Wau
Subdistr., Otibanda Creek 2150 m (LAE; K).
Stipe 5-6 cm long, black at base, paler upwards,
short-hairy throughout, basal scales thin, 2
1 mm, not setiferous. Lamina 14 cm long; pinnae
12-15 pairs, lower 2-3 pairs deflexed and nar-
rowed at base, their basal acroscopic lobes slightly
enlarged and dentate, almost free. Largest pinnae
3.0 x 0.9 cm, apex abruptly acute, lobed almost to
costa; lobes entire or slightly dentate; costules
2mm apart; veins 4 pairs; lower surface of rachis
and costae densely covered with short erect hairs,
very narrow scales and sometimes small ovate
ones also present; glands present on all lower
surfaces; upper surface with hairs on costa, the
rest glabrous or sometimes with a few hairs be-
FLORA MALESIANA
[ser. II, vol. 1°
tween veins. Sori medial; indusia large, thin,
sometimes bearing short capitate hairs.
Distr. Malesia: Papua New Guinea (neigh-
bourhood of Mt Kaindi, several collections; Ekuti
Range 2200 m).
19. Coryphopteris squamipes (COPEL.) HOLT-
TUM, Blumea 23 (1976) 35.— Dryopteris
squamipes COPEL. Philip. J. Sci. 56 (1935) 99, pl.
5.— Lastrea squamipes (COPEL.) COPEL. Fern
Fl. Philip. (1960) 325.— Thelypteris squamipes
(COPEL.) REED, Phytologia 17 (1968) 315.—
Type: RAMOS & EDANO BS 38525, Mindanao,
Bukidnon Prov., Mt Lipa, 2000m (MICH; BO,
US).
Stipe 5-15 cm long, dark at base, paler distally;
scales thin, to 5mm long, to 2.5 mm wide at base.
Lamina to 28 cm long; pinnae more than 20 pairs,
6-8 lower pairs gradually smaller, lowest 1.5—2 cm
long. Largest pinnae commonly 3.5*0.9cm,
short-acuminate; lobes oblong, mostly crenate,
basal ones dentate; costules 2-2.5 mm apart; veins
to 5 pairs; lower surface of rachis and costae
bearing stiff unicellular hairs to 1mm long, on
costae many small scales sometimes widened at
their bases, glands present on costules, veins and
surface between veins; upper surface sometimes
with a few glands on and near veins. Sori medial,
distal ones mostly symmetric; indusia glabrous or
with a few glands.
Distr. Malesia: Philippines (Mindanao, several
coll.) and Papua New Guinea (New Ireland, J. R.
CROFT 288, pinnae to 6 by 1.4cm).
20. Coryphopteris borealis HOLTTUM, Blumea 23
(1976) 35.—Type: M. JACOBS 7588, Northern
Luzon, Mt Tayaboc, 2300 m, on ridge in shade (K;
1s)!
Caudex to 30cm tall. Stipe 20-25 cm long, very
dark at base, dark reddish distally; scales to 4x
2.5mm. Lamina to 45 cm long; pinnae 20 pairs or
more; lower pinnae 2-4 pairs somewhat reduced,
lowest 2.5-3 cm long. Largest pinnae 4.5 x 1.2cm
(fertile), 5.0 1.3. cm (sterile), abruptly narrowed
at apex; lobes slightly oblique, subtruncate,
toothed at ends of veins, lowest lobes of fertile
pinnae strongly dentate; costules 3—3.5 mm apart;
veins 4 pairs; lower surface of rachis and costae
bearing hairs almost 1 mm long, scales on costae
all very narrow, costules and veins bearing small
glands; upper surface of pinnae glabrous apart
from hairs on costae. Sori inframedial; indusia
large, glabrous.
Distr. Malesia: Philippines (Luzon), type spe-
cimen only.
21. Coryphopteris meiobasis HOLTTUM, Blumea
23 (1976) 36. — Type: T. G. WALKER 8730, N.E.
New Guinea, Morobe Distr., trail from Sewe to
Freyburg Pass, 2300-2450 m (BM).
Stipe to 25cm long, covered with pale slender
1981]
hairs, basal scales little more than 1mm wide.
Lamina to 40cm long; pinnae 20 pairs, lower 3
pairs gradually smaller, lowest 3.5-4.5 x 1.1-
1.3.cm, their basal acroscopic lobes not enlarged.
Largest pinnae 9X1.9cm (sterile), 8.5 1.7 cm
(fertile), acuminate; lobes oblique, strongly den-
tate; costules 4.5-5.5 mm apart; veins to 7 pairs;
lower surface of rachis with many hairs | mm
long, similar but shorter hairs and very narrow
scales on costae, glands on costules and veins,
few glands between veins; upper surface glabrous
apart from hairs on costae and a few on costules
and veins, no glands. Sori inframedial, round;
indusia thin, pale, with a few hairs.
Distr. Malesia: Papua New Guinea; known
from type only.
22. Coryphopteris pubirachis (BAK.) HOLTTUM,
Blumea 23 (1976) 37.— Nephrodium pubirachis
BAK. J. Bot. 14 (1876) 344.— Dryopteris
pubirachis (BAK.) C. CHR. Ind. Fil. (1905) 287;
Bishop Mus. Bull. 177 (1943) 82.— Thelypteris
pubirachis REED, Phytologia 17 (1968) 307.—
Type: WHITMEE 202, Samoa (K).
Dryopteris mataanae BRAUSE, Notizbl. Bot.
Gart. Berlin 8 (1922) 1939. Type: VAUPEL 460,
Samoa (B; BM).
a. var. pubirachis
Stipe 15-20cm long, very dark throughout;
scales to 5X 1.5mm, thin, lacking acicular hairs:
rachis dark to light reddish. Lamina to 25cm
long; pinnae 12-15 pairs, well spaced; basal pinnae
wider than next pair, narrowed to base on basis-
copic side, basal acroscopic lobe free, strongly
dentate, not elongate. Suprabasal pinnae to 7.5 x
1.8cm, short-acuminate, lobes slightly oblique,
almost all dentate at ends of veins, basal acros-
copic lobe usually longer than rest; costules 4mm
apart; veins 6—7 pairs; lower surface of rachis and
costae with slender acicular hairs less than 0.5 mm
long, glands present on costae, costules and veins,
none or few between veins, a few narrow scales
widened at base present on costae; hairs on upper
surface of rachis and costae longer and thicker
than on lower surface, a few on costules and
veins, not elsewhere. Sori inframedial, basal ones
a little divergent; distal sori smallest, sometimes a
little asymmetric; indusia bearing glands, a few
hairs sometimes also present.
Distr. Polynesia: Samoa, Tahiti.
b. var. major HOLTTUM, Blumea 23 (1976) 37. —
Type: BRAITHWAITE 4378, Solomon Islands,
New Georgia Group, Kolombangara, 1650 m (K).
Differs from typical form: larger, with lamina to
40 cm long; pinnae to 20 pairs; largest suprabasal
pinnae to 10x 1.8cm; hairs on lower surface of
rachis and costae sometimes few.
Distr. Solomon Islands, Bougainville, in
Malesia: New Ireland, New Guinea.
THELYPTERIDACEAE (Holttum)
365
c. var. philippinensis HOLTTUM, Blumea 23 (1976)
37. — Type: RAMOS & EDANO BS 37959, Luzon,
Mt Masapilid, Bontoc Subprov. (K).
Differs from type: larger, with basal pinnae to
9x2cm; lower surface of pinnae between veins
glandular, of upper surface bearing some acicular
hairs; lobes of pinnae, apart from basal ones,
almost entire.
Distr. Malesia:
mountains.
Note. Young plants appear to have lower pin-
nae somewhat smaller than next and have some
resemblance to C. squamipes, but well-grown
plants of the two, as collected in Mindanao, are
distinct.
throughout Philippines, on
d. var. sulawesica HOLTTUM, Blumea 23 (1976)
38. — Type: T. G. WALKER 12354, Celebes, on
ridge above river Pasir, 2000-2200 m (BM).
Differs from type: largest pinnae 1.2cm wide;
lower surface between veins bearing glands; upper
surface between veins bearing acicular and capi-
tate hairs.
Distr. Malesia: Celebes and Borneo (Sarawak:
Mt Mulu), 1800-2200 m; two collections known.
23. Coryphopteris athyriocarpa (COPEL.) HOLT-
TUM, Blumea 23 (1976) 38. — Dryopteris athyrio-
carpa COPEL. Philip. J. Sci. 3 (1908) Bot. 344. —
Type: BRooKs & Hewitt 2, Bongo Range,
Sarawak (MICH).
Stipe to 21cm long, base very dark, becoming
reddish distally, rachis paler; basal scales to 5mm
long, base dilated with isodiametric cells, above
base less than 1mm wide with elongate cells.
Lamina 20 cm long; free pinnae 18 pairs; basal
pinnae largest, 5.5 1.8cm, their lobes lobulate,
3-4 pairs basal lobes gradually — shorter.
Suprabasal pinnae to 4 1.2 cm, short-acuminate;
basal pair of lobes lobulate, rest crenate; costules
3mm apart; veins 4-5 pairs, very oblique; lower
surface of rachis bearing short capitate hairs, of
costae capitate hairs, glands and very narrow
scales, acicular hairs lacking; upper surface of
rachis bearing acicular hairs 0.5 mm long, shorter
ones on costae and scattered on costules and
veins, between veins many capitate hairs and a
few short acicular hairs, no glands. Sori medial,
distal ones athyrioid; indusia with a few glands,
not hairs.
Distr. Malesia: Borneo (Sarawak; W. Kali-
mantan, Pontianak R., HANS WINKLER 534, in
BM); only two collections known.
Note. This has the narrow scales of C. viscosa
but smaller fronds and a different distribution of
glands and hairs.
24. Coryphopteris tanggamensis HOLTTUM, Blu-
mea 23 (1976) 38.— Type: M. JAcoss 8255, S.
Sumatra, G. Tanggamus, 2000 m (L; K).
Caudex to 30cm tall. Stipe 30cm _ long,
366
glabrous, dark purplish at base, distally dull red-
dish, basal scales 5X 1.2mm. Lamina to 35cm
long; pinnae 22 pairs, widely spaced; two pairs
lower pinnae deflexed but not reduced; lowest pair
somewhat narrowed at base, basal acroscopic lobe
elongate, dentate, not free. Suprabasal pinnae to
7x*1.5cm, acuminate; lobes oblique, falcate,
margins at base dentate, apex rounded and entire;
costules 3.5mm apart; veins 6-7 pairs; lower sur-
face of rachis bearing many capitate and fewer
acicular hairs, of costae bearing capitate hairs or
small glands and linear scales, no glands seen
between veins; upper surface of rachis bearing
hairs 0.7mm long, those on costae shorter, cos-
tules and veins with sparse hairs, none between
veins. Sori inframedial, distal ones mostly sym-
metric; indusia bearing capitate hairs.
Distr. Malesia: S. Sumatra (Mt Tanggamus),
known from type only.
25. Coryphopteris engleriana (BRAUSE) HOLT-
TUM, Blumea 23 (1976) 40.— Dryopteris engleri-
ana BRAUSE, Bot. Jahrb. 49 (1912) 19. — Phe-
gopteris engleriana (BRAUSE) v.A.v.R. Handb.
Suppl. (1917) 309.— Thelypteris engleriana
(BRAUSE) REED, Phytologia 17 (1968) 274.—
Type: L. SCHULTZE 330, N.E. New Guinea,
Sepik Distr. (B).
Stipe and rachis dark; base of stipe not seen.
Lamina to 50cm long; pinnae 18 pairs, widely
spaced: basal pinnae not reduced, narrowed
gradually in basal half, basal pair of lobes free, not
enlarged, not dentate. Suprabasal pinnae to 10.8 x
1.6cm with stalks 1mm long, apex narrowly
acuminate; middle lobes somewhat falcate with
rounded ends, separated by rather wide sinuses,
edges slightly crenate; costules 4.5 mm apart; veins
7 pairs; lower surface lacking acicular hairs, cos-
tae bearing narrow scales (often uniseriate), no
glands; upper surface with coarse brown hairs on
rachis and costae, no others. Sori inframedial
except basal ones, exindusiate; spores with many
small wings.
Distr. Malesia: Papua New Guinea. Known
only from type and one other collection from
same locality.
26. Coryphopteris stereophylla (v.A.v.R.) HOLT-
TUM, Blumea 23 (1976) 40.— Dryopteris stereo-
phylla v.A.v.R. Nova Guinea 14 (1924) 17.—
Thelypteris stereophylla (v.A.v.R.) CHING, Bull.
Fan Mem. Inst. Biol. Bot. 10 (1941) 254. — Type:
H. J. LAM 1785, W. New Guinea, Doormantop,
3200 m (L).
Stipe to 30cm, base dark, rest flushed with red;
hairs very short, thick, brownish; scales 3 x 1 mm,
dark. Lamina to 25cm long with 15-20 pairs
well-spaced pinnae, texture thick, rigid when dry;
lowest pinnae not reduced, narrowed a little at
basiscopic base. Suprabasal pinnae to 7.5
1.5 cm, with 4-8 pairs of free or separately adnate
FLORA MALESIANA
[ser. II, vol. 1°
pinnules in basal part, apical part lobed almost to
costa; pinnules and lobes 1-2 mm wide, almost all
crenate, separated by wide sinuses; costules 3 mm
apart on type, to 4mm on another specimen; veins
4-5 pairs, grooved on upper surface; lower sur-
face of rachis covered with brown hairs 0.3 mm
long and thin scales, costae with similar hairs and
ovate-acute to lanceolate clathrate scales 1mm
long, no glands; upper surface bearing very short
erect hairs on edges of grooved costa. Sori
medial; indusia small, thin, glabrous.
Distr. Malesia: Papua New Guinea. Only
known from the type and PULLE 883 from Mt
Hellwig, 2000 m (L).
Note. This species is close to C. habbemensis,
differing from the latter in more coriaceous tex-
ture, narrower pinnules and lack of glands on
lower surface.
27. Coryphopteris horizontalis (ROSENST.)
HOLTTUM, Blumea 23 (1976) 40.— Athyrium
horizontale ROSENST. Nova Guinea 8 (1912)
722. — Dryopteris horizontalis (ROSENST.)
v.A.v.R. Bull. Jard. Bot. Btzg II, 11 (1913) 10;
Handb. Suppl. (1917) 151. —Type: VON ROEMER
1136, W. New Guinea, Hellwig Mts, 1350-1600 m
(S-PA: BO).
Stipe 15cm long, dark, densely covered with
hairs Imm long and thin scales 3x 1.5mm.
Lamina to 40cm long; pinnae 18-20 pairs; lowest
pinnae a little reduced and narrowed to base.
Suprabasal pinnae to 7*1.5cm, sessile, short-
acuminate; lobes oblong, serrate-crenate, basal
lobes most strongly so; costules to 4mm apart;
veins 6-7 pairs: rachis beneath bearing acicular
hairs | mm long, shorter hairs on costae, sparse on
costules, no glands; linear scales, sometimes
dilated at base, on costules; hairs on upper surface
of rachis as lower, on costae shorter, copious
short hairs on surface between veins. Sori in-
framedial, mostly not athyrioid; indusia_ thin,
short-hairy.
Distr. Malesia: Papua New Guinea. Known
only from type and PULLE 633, Mt Dromedaris
1250 m (L).
28. Coryphopteris microlepigera HOLTTUM,
Blumea 23 (1976) 41.— Type: PULLE 1078, W.
New Guinea, Mt Treub, 2300 m (L; BM).
Stipe to 15cm long, very dark; hairs in groove
0.5 mm long, basal scales 2-3 mm long, thin; rachis
dark reddish. Lamina to 28cm long; pinnae 20
pairs; lower pinnae 1-3 pairs somewhat reduced.
Largest pinnae 5.5x1.l1cm, short-acuminate;
basal lobes conspicuously dentate, rest with +
sinuous margins; costules 3mm apart; veins 4-5
pairs; lower surface of rachis covered with brown
hairs 0.5-0.8mm long, acicular hairs on costae
shorter, costules and veins bearing very short
capitate hairs, scales on costae lanceolate, at base
5-10 cells wide; upper surface of rachis with hairs
1981]
THELYPTERIDACEAE (Holttum)
367
as lower surface, hairs on costae shorter, surface
between veins covered with short acicular hairs.
Sori a little supramedial, sometimes slightly athy-
rioid; indusia glabrous.
Distr. Malesia: Western New Guinea and
Moluccas (Amboina).
Note. A second New Guinea collection, from
Mt Nettoti at 1800m (VAN ROYEN & SLEUMER
8226) differs in shorter narrower pinnae with few
hairs between veins on upper surface; a specimen
collected by TEYSMANN on Mt Toena, Amboina
(BO) is similar.
29. Coryphopteris propria (v.A.v.R.) HOLTTUM,
Blumea 23 (1976) 41.—Dryopteris propria
v.A.v.R. Bull. Jard. Bot. Btzg 16 (1914) 10; Handb.
Suppi. (1917) 152. — Type: RACHMAT 496, Cen-
tral Celebes, Tondo-Tondo (BO; L).
Stipe to 13 cm long, dark; scales 3 x 1 mm, thin.
Lamina to 14cm long; pinnae 10 pairs; basal
pinnae largest, to 4x 1.3.cm, a little narrowed at
base, basal acroscopic lobe enlarged, dentate,
free. Suprabasal pinnae to 3.5 1.1cm, short
acuminate, lobes entire or with slight teeth at ends
of distal veins: costules to 3mm apart; veins 4
pairs; abundant very short capitate hairs on lower
surface of rachis, costae, costules, veins and sur-
face between veins; sparse short acicular and
many short capitate hairs on upper surface of
costae, very short acicular hairs on costules and
veins, short capitate hairs between veins. Sori
medial, basal ones often somewhat athyrioid; in-
dusia bearing many capitate hairs and sometimes a
few acicular ones.
Distr. Malesia: Central Celebes; known from
type collection only.
30. Coryphopteris lauterbachii (BRAUSE) HOLT-
TUM, Blumea 23 (1976) 41.— Dryopteris lauter-
bachii BRAUSE, Bot. Jahrb. 49 (1912) 18; v.A.v.R.
Handb. Suppl. (1917) 150.— Thelypteris lauter-
bachii (BRAUSE) REED, Phytologia 17 (1968)
i/-— hype, Lo SCHULTZE, 2735 (N-E: New
Guinea, Sepik Distr. (B).
Stipe to 25cm, dark, upper part and rachis dull
reddish; soft pale hairs on abaxial surface; scales
3-4 x 1 mm, thin. Lamina to 43 cm long; pinnae to
24 pairs; basal pinnae a little reduced, slightly
narrowed at base, basal lobe not free. Suprabasal
pinnae 7.5 1.7cm, sessile, base truncate and a
little dilated both sides, apex acuminate; lobes
slightly oblique, slightly crenate; costules 3.5mm
apart; veins to 8 pairs; rachis and costae beneath
bearing short acicular hairs, few on costules, very
small capitate hairs on surface between veins,
many small scales on costae and costules; hairs on
upper surface of costae longer, some consisting of
2 cells, sparse hairs on costules and veins. Sori
medial; indusia large, thin, short-hairy.
Distr. Malesia: Papua New Guinea, known
from type only.
31. Coryphopteris fasciculata (FOURN.) HOLT-
TUM, Blumea 23 (1976) 42.—Aspidium fasci-
culatum FOURN. Ann. Sci. Nat. V, 18 (1873)
295. — Nephrodium fasciculatum (FOURN.) BAK.
Ann. Bot. 5 (1891) 320. — Thelypteris fasciculata
(FOURN.) CHING, Bull. Fan Mem. Inst. Biol. Bot.
10 (1941) 251; BROWNLIE in Aubrév. FI. Nouv.
Cal. 3 (1969) 210, pl. xxvii.— Type: BALANSA
3568, New Caledonia, Mt Humboldt (P).
Nephrodium macgregorii BAK. Ann. Bot. 5
(1891) 320, new name for N. simulans BAK. J.
Bot. 28 (1890) 106, non BAK. 1874 nec BAK.
1888. — Dryopteris conterminoides C. CHR. Ind.
Fil. 258, nom. nov. superfl. — Lastrea macgregorii
(BAK.) RIDL. Trans. Linn. Soc. Bot. 9 (1916)
257. — Dryopteris macgregorii (BAK.) C. CHR.
Ind. Fil. Suppl. III (1934) 90.— Lastrea conter-
minoides (C. CHR.) COPEL. Philip. J. Sci. 78
(1951) 424.— Thelypteris conterminoides (C.
CHR.) REED, Phytologia 17 (1968) 269.— Type:
W. MCGREGOR 18, Papua, Mt Knutsford (K).
Dryopteris engleriana var. hirta C. CHR. Brit-
tonia 2 (1937) 296. — Type: BRASS 5032, Papua,
Mt Tafa, 2400 m (BM; K, NY).
Stipe 12-20 cm long, dark at base only, reddish
upwards, bearing hairs | mm long; scales 3 x 1 mm
or somewhat larger. Lamina 30-40 cm long; pin-
nae 25-30 pairs; up to 8 pairs lower pinnae
deflexed and gradually reduced, lowest 1-2 cm
long. Largest pinnae 4-S5.5 x 0.8-1.3cm, short
acuminate; lobes + dentate at vein-ends, basal
lobes most strongly; costules 2.5-3mm_ apart;
veins 3-4 pairs; lower surface of rachis and costae
bearing rather thick acicular hairs 1mm _ long,
linear scales present on costae and costules, rarely
a few glands on costae only, short erect hairs
sometimes present between veins; copious aci-
cular hairs on upper surface of rachis and costae,
few on costules and veins, sometimes short capi-
tate hairs present between veins. Sori inframedial;
indusia thin with short capitate hairs and few to
many short acicular hairs.
Distr. New Caledonia, East Malesia:
Guinea, Celebes, at 1800-3000 m.
Note. The type of N. macgregorii BAK. is a
poor specimen with basal pinnae lacking; it agrees
well with other specimens in pubescence. Some
specimens from both east and west New Guinea
have a few glands on lower surface of costae,
others agree with the type in having none. All
Celebes specimens have some short hairs between
veins on both sides.
New
32. Coryphopteris hubrechtensis HOLTTUM,
Blumea 23 (1976) 42.— Type: VERSTEEG 2433,
W. New Guinea, Mt Hubrecht 3000 m (BM; L).
Stipe to 35cm long, very dark, glossy, glabrous
except near apex which is dull reddish. Lamina
20cm long; pinnae 18 pairs, rigid, several lower
ones deflexed, lowest 1-2 pairs slightly reduced.
Largest pinnae 3cm long, to 1cm wide, apex
368
FLORA MALESIANA
(ser. II, vol. 1°
obtuse; lobes almost entire, basal acroscopic lobe
slightly enlarged, almost free and slightly dentate;
costules 2.5mm apart; veins 4-5 pairs, sometimes
forked in the basal acroscopic lobe; lower surface
of rachis bearing copious acicular hairs 0.5mm
long, of costae bearing narrow scales 0.5-0.8 mm
long, minute uniseriate scales also on costules and
veins, no glands; upper surface glabrous apart
from costae. Sori near costules; indusia rather
large, glabrous.
Distr. Malesia: West New Guinea, c. 3000 m.
Besides the type, 2 collections by BRASS from
Lake Habbema.
33. Coryphopteris brevipilosa HOLTTUM, Blumea
23 (1976) 43.—Type: PULLE 532, W. New
Guinea, Mt Perameles, 900m, on limestone (L).
Stipe 16-24cm long, very dark at base, paler
upwards, covered with hairs 0.3 mm long; scales
3 x 1mm, thin. Lamina to 30cm long, thin; pinnae
15-18 pairs; lowest pinnae not or little shorter
than next, 1.8 cm wide, with acroscopic basal lobe
free and strongly dentate. Suprabasal pinnae to
6.5 1.5cm, with winged stalk 1mm long; apex
acuminate, lobes almost all dentate and separated
by distinct sinuses; costules 3.5 mm apart; veins 6
pairs; rachis and costae beneath covered with
hairs 0.2 mm long, costae bearing also subsessile
glandular hairs and narrow scales, rest of lower
surface glabrous; upper surface of rachis with
hairs 0.5mm long, those on costae 0.2 mm long.
Sori medial, upper ones always athyrioid; indusia
thin, glabrous.
Distr. Malesia: West New Guinea
Perameles, 900 m), known from type only.
(Mt
34. Coryphopteris oligolepia (v.A.v.R.) HOLTTUM,
Blumea 23 (1976) 43.—Dryopteris oligolepia
v.A.v.R. Nova Guinea 14 (1924) 17. — Thelypteris
oligolepia (v.A.v.R.) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 253.— Type: LAm 1977, W.
New Guinea, Doormantop 2520 m (L).
Stipe 12-20cm long, dark glossy, scales 2-3 x
1mm. Lamina to 25cm long; pinnae 20 pairs;
lowest pinnae deflexed and reduced, their basal
lobes free and dentate. Largest pinnae 4.0
1.3cm, short acuminate, with stalks hardly | mm
long; lobes oblique, crenulate; costules 3-4mm
apart; veins 3-5 pairs; lower surface of rachis
castaneous, glabrous, all other lower surfaces
bearing scattered very small capitate hairs, some
linear scales on costae; upper surface hairy on
edges of groove of rachis and costae, few hairs on
costules. Sori near costules, sometimes asym-
metric; indusia small with some capitate hairs.
Distr. Malesia: Western New Guinea, several
collections.
Ecol. Twice recorded as an epiphyte; at 1800-
2500 m.
35. Coryphopteris coriacea (BRAUSE) HOLTTUM,
Blumea 23 (1976) 43.— Dryopteris coriacea
BRAUSE, Bot. Jahrb. 56 (1920) 63. — Thelypteris
coriacea (BRAUSE) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 251.—Lastrea coriacea
(BRAUSE) COPEL. Philip. J. Sci. 78 (1951) 428. —
Type: LEDERMANN 10965, N.E. New Guinea,
Sepik Distr., Hunsteinspitze 1300 m (B).
Dryopteris coriacea var. elata BRAUSE, l.c. —
Type: LEDERMANN 11291, same locality.
Stipe dark, glossy, 8-12 cm long, hairs on adax-
ial side short; scales 8-10 x 1.5-2 mm; rachis red-
dish, green towards apex. Lamina to 26cm long;
pinnae 12 pairs; lower 3-5 pairs pinnae deflexed
and short-stalked, narrowed at base on both sides.
Largest pinnae 5.5 1.5cm (of var. elata 8.5
2.0cm), short-acuminate; lobes oblique (except
basal ones) and slightly falcate, almost or quite
entire; costules 3.5mm apart; veins to 6 pairs,
grooved above and flat beneath; lower surface
quite glabrous, some small dark scales present on
costae; upper surface hairy on rachis and costae,
scattered hairs on margins of lobes. Sori near
margin; indusia thin, glabrous.
Distr. Malesia: Papua New Guinea. Known
only from the two LEDERMANN collections.
36. Coryphopteris ledermannii (HIERON.) HOLT-
TUM, Blumea 23 (1976) 44.— Athyrium leder-
mannii HIERON. Bot. Jahrb. 56 (1920) 133.—
Type: LEDERMANN 11906, N.E. New Guinea,
Schraderberg, 2070 m (B).
Stipe dark, glossy, hairy in groove only, 15cm
long. Lamina to 20cm long; pinnae well spaced,
to 13 pairs, texture thin; basal pinnae short-stal-
ked, somewhat reduced. Largest pinnae 2.5 x
0.8cm, apex obtuse, in the middle lobed about
half-way to costa, towards base more deeply,
lobes entire or slightly dentate; costules to 3mm
apart; veins 3-4 pairs in basal lobe, 2 pairs in
middle lobes; lower surfaces quite glabrous, no
scales seen; upper surface with short hairs on
rachis and costae only. Sori medial, almost all
athyrioid; indusia glabrous.
Distr. Malesia: Papua New Guinea, only
known from the type.
Ecol. Epiphyte in moss-forest.
37. Coryphopteris dura (COPEL.) HOLTTUM,
Blumea 23 (1976) 44.— Dryopteris dura COPEL.
Leafl. Philip. Bot. 3 (1910) 805; v.A.v.R. Handb.
Suppl. (1917) 148.—Lastrea dura (COPEL.)
COPEL. Gen. Fil. (1947) 135; Fern FI. Philip. (1960)
323. — Thelypteris dura (COPEL.) REED, Phy-
tologia 17 (1968) 274. — Type: ELMER 11674, Min-
danao, Mt Apo, 2600 m (MICH; BM, E, L).
Stipe dark near base, paler distally, 20-30 cm
long, glabrous except for groove of upper part;
scales 5X 1-2 mm, acuminate. Lamina to 25cm
long; pinnae 18 pairs, rather thick; basal pinnae
sessile, sometimes slightly reduced, a little nar-
rowed at base, basal acroscopic lobe free, orbi-
1981]
THELYPTERIDACEAE (Holttum)
369
cular, a little dentate. Largest pinnae 4.5 x 1.1 cm,
apex blunt, lobed at base to | mm from costa, less
deeply towards apex, lobes rounded, entire except
basal acroscopic ones; costules 3—3.5 mm apart;
veins 3-4 pairs, slightly prominent on both sides;
lower surfaces hairless apart from edges of lobes,
scales on costae 2-3 cells or more wide at base;
short thick hairs on upper surface of rachis and
costae. Sori near costules; indusia glabrous,
sometimes a little athyrioid.
Distr. Malesia: Philippines (Mindanao: Mt
Apo), 2600 m.
38. Coryphopteris platyptera (COPEL.) HOLT-
TUM, Blumea 23 (1976) 45.— Dryopteris platyp-
tera COPEL. Univ. Cal. Publ. Bot. 18 (1942)
219. — Lastrea platyptera (COPEL.) COPEL. Gen.
Fil. (1947) 139; Philip. J. Sci. 78 (1951) 433, pl.
17.— Thelypteris platyptera (COPEL.) REED,
Phytologia 17 (1968) 304.— Type: BRASS 11328,
N. New Guinea, Bele River (MICH; L).
Stipe 20-25cm, dark and glossy throughout;
scales ovate, cordate, 21mm, thin. Lamina
30 cm long; pinnae to 18 pairs, well spaced, many
distinctly stalked; basal pinnae slightly reduced,
stalked 1 mm, basal acroscopic lobe quite free, a
little dentate. Largest pinnae 5.5-7 cm long, 1.3-
2cm wide, caudate-acuminate (cauda 7-15 mm
long, entire), deeply lobed throughout, lobes in
basal part of pinna + crenate-dentate; costules
3.5-4mm apart; veins to 6 pairs; lower surface
hairless apart from edges of lobes, ovate-acute
scales present on costae, very small filamentous
scales on costules and veins; stiff dark hairs on
upper surface of rachis and costae. Sori in-
framedial; indusia small, glabrous.
Distr. Malesia: New Guinea.
from type collection.
Ecol. On rocky banks of stream at 2200 m.
Known only
39. Coryphopteris subnigra (BRAUSE) HOLTTUM,
Blumea 23 (1976) 45.—Dryopteris subnigra
BRAUSE, Bot. Jahrb. 56 (1920) 82. — Thelypteris
subnigra (BRAUSE) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 254.—Lastrea subnigra
(BRAUSE) COPEL. Gen. Fil. (1947) 140; Philip. J.
Sci. 78 (1951) 427. — Type: LEDERMANN 11962,
N.E. New Guinea, Sepik Distr. (B).
Stipe to 24cm long, very dark, glabrous apart
from hairs in groove; scales 3-4 x 1 mm. Lamina
to 25 cm long; pinnae 12 pairs, well spaced; lower
pinnae stalked hardly | mm, narrowed to base on
basiscopic side, basal acroscopic lobe free (also
on all other free pinnae) and dentate. Largest
pinnae 5-6X1.4cm, _ short-acuminate, where
sterile lobed to 1mm from costa, where fertile
more deeply; lobes + dentate, most strongly
where fertile, oblique; costules 3-3.5 mm apart;
veins to 6 pairs, prominent and slender on both
surfaces; lower surfaces lacking acicular hairs, a
few short capitate hairs sometimes on costae,
scales with widened base present on costae, uni-
seriate scales on costules; short acicular hairs on
upper surface of rachis and costae. Sori near
costules; indusia small, glabrous, sporangia often
with a red glandular cell on stalk.
Distr. Malesia: Papua New Guinea. Many col-
lections, 1500-3000 m. Misima I., 1000 m.
Ecol. The type grew as an epiphyte in moss
cushions; no others so reported.
40. Coryphopteris badia (v.A.v.R.) HOLTTUM,
Blumea 23 (1976) 44. — Dryopteris badia v.A.v.R.
Bull. Jard. Bot. Btzg II, 16 (1914) 9; Handb. Suppl.
(1917) 149. — Thelypteris badia (v.A.v.R.) CHING,
Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 250. —
Type: MATTHEW 674, Sumatra, Mt Tandikat
(BO; E).
Dryopteris linearis COPEL. Philip. J. Sci. 12
(1917) Bot. 56.—Type: CLEMENS 11069, Mt
Kinabalu, Marei Parei ridge (MICH; BM, BO,
K).
Dryopteris villosipes GEPP in Gibbs, Dutch
N.W. New Guinea (1917) 70. — Type: L. S. GIBBS
5627, W. New Guinea, Arfak Mts (BM).
Dryopteris rigidifolia v.A.v.R. Nova Guinea 14
(1924) 18.— Thelypteris rigidifolia (v.A.v.R.)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941)
254.— Lectotype (HOLTTUM 1976): LAM 1562,
W. New Guinea, Doormantop (BO). — Fig. 4g—h.
Stipe varying much in length according to habi-
tat (extremes 10cm, 70cm), dark, glossy, with
hairs in groove only, at base often bearing a
tangled mass of slender hairs which are glossy
golden brown when dry; scales narrow, varying
with size of frond; rachis also dark throughout.
Lamina varying from 10cm long (type of D.
villosipes) to 65 cm (specimen from Atjeh); pinnae
15 pairs on small plants, to 30 pairs on large ones,
texture always thick, rigid when dry; nearly all
pinnae distinctly stalked; basal pinnae sometimes
a little reduced, little narrowed at base, basal
acroscopic lobe free or nearly so. Largest pinnae
commonly 5-6 x 1.2-1.5cm (extremes 1.70.5,
8 x2cm), lobed to about 1 mm from costa; lobes
entire or nearly so, deciduously ciliate on edges;
costules less than 2mm apart on small plants,
3-3.5 mm on large ones; veins commonly 4-S pairs
(extremes 2 and 6 pairs), sometimes grooved on
both sides; lower surfaces usually quite glabrous
apart from hair-like scales on costae and costules
(a few acicular hairs on costae of type of D.
linearis, Fig. 4h); upper surface of rachis and
costae bearing rigid dark brown hairs. Sori near
costules; indusia glabrous, sporangia sometimes
bearing 2-3 slender non-glandular hairs on their
Stalks.
Distr. Malesia: Sumatra, Malaya, Sarawak,
Sabah, Celebes, New Guinea.
Ecol. In mossy forest at 1400-2500 m, usually
in moss cushions, several times reported in moss
cushions on tree-branches. The largest specimen
370
seen, from Atjeh, was an epiphyte. The slender
hairs at base of stipes resemble root-hairs, and
perhaps have the same function, absorbing water
from moss-cushions.
41. Coryphopteris inopinata HOLTTUM, sp. nov.
Caudex gracilis longe-repens; lamina usque
11.5cm longa; pinnae usque 2.1x0.6cm, lobis
dentatis; costae subtus pilis acicularibus destitu-
tae, eglandulosae, paleis linearibus praeditae;
pagina superior pinnarum inter venas glabra, in-
dusia glabra. — Type: J. R. CROFT, LAE 65849,
Telefomin Subdistr. West Sepik Distr., N.E. New
Guinea, 2700 m (LAE; K, NSW).
Caudex a slender black rhizome 1.5-2.0mm
diameter, bearing fronds 1-3cm apart; scales c.
4x 1mm; stipe 12-20 cm long, dark at base, paler
distally, slender hairs present on adaxial side only.
Lamina to 11.5cm long; pinnae 12 pairs; basal
FLORA MALESIANA
(ser. II, vol. 1°
basiscopic lobe smaller with decurrent base; fifth
pair of pinnae with almost symmetric base and
winged stalk 0.5mm long. Largest pinnae 2.3 x
0.7 cm; apex subabruptly narrowed and rounded;
edges lobed to |mm from costa, lobes obliquely
quadrate with teeth at ends of veins; costules
2.5mm apart; veins 3 on acroscopic side of cos-
tule, 2 on basiscopic side; lower surface of rachis
and pinnae lacking hairs of any kind, glands also
absent, scales on costae narrowly linear (2-4 cells
wide at base); upper surface lacking hairs between
veins. Sori near base of veins; indusia small,
glabrous.
Distr. Malesia: Papua New Guinea (Tele-
fomin, 2700 m).
Note. Apart from the rhizome, this is near C.
subnigra but has much smaller pinnae. No other
known Coryphopteris has this habit. The NSW
isotype has fronds closer together on the caudex,
which appears to be decumbent; it has a branch
near its base.
pinnae narrowed towards their bases, basal
acroscopic lobe 2 mm long, free or nearly so, basal
6. PARATHELYPTERIS
(H. Iro) CHING, Acta Phytotax. Sinica 8 (1963) 300, p.p. — Thelypteris sect.
Parathelypteris H. Iro in Nakai & Honda, Nov. FI. Jap. n. 4 (1939) 127,
excl. T. hirsutipes et T. simozawae. — Thelypteris group 2 CHING, Bull. Fan
Mem. Inst. Biol. Bot. 6 (1936) 246, excl. T. hirsutipes. — Thelypteris subg.
Thelypteris sect. Thelypteris K. IwATS. Mem. Coll. Sci. Univ. Kyoto B, 31
(1965) 157, groups of T. glanduligera and T. japonica p.p.— Wagneriop-
teris LOvE & LOvE, Taxon 26 (1977) 325 excl. W. ogasawarensis; HOLT-
TUM, Acta Phytotax. Geobot. 29 (1978) 16. — Fig. 5a—b.
Rather small ferns; caudex slender, creeping, sometimes much elongate;
scales bearing superficial unicellular hairs, either spherical, capitate or aci-
cular; basal pinnae in most species not or little reduced, in P. beddomei
several pairs progressively smaller; pinnae deeply lobed, basal one or more
lobes sometimes free; basal acroscopic lobe or leaflet often somewhat
enlarged and dentate; veins free, basal ones both reaching margin above
base of sinuses between lobes; some acicular hairs on lower surface of
costae of P. beddomei septate, on upper surface always unicellular; scales
on lower surface of costae never large or abundant; sessile glands almost
always present on lower surfaces; sori indusiate; sporangia short-stalked,
never with glands or setae near annulus, a sessile gland sometimes present
on the stalk; spores opaque with a narrow wing as seen by the light
microscope, covered with a finely reticulate perispore as seen with S.E.M.
Type species: Parathelypteris glanduligera (KUNZE) CHING.
Distr. c. 15 species; tropical and subtropical S.E. Asia and Malesia, N. America.
Cytol. n=27 (P. cystopteroides (EATON) CHING, P. noveboracensis (L.) CHING); n= 31 (P. bed-
domei); n= 32 or 31 (P. simulata (DAVENP.) NIEUWL.).
Taxon. In his original list of species, CHING included several which I have separated as Cory-
phopteris, also Aspidium immersum BL. (Amphineuron immersum HOLTTUM). In addition to species
1981] THELYPTERIDACEAE (Holttum) 371
from Asia, he added P. noveboracensis from North America which in habit is similar to P. beddomei of
Malesia. He did not include another North American species, originally named Aspidium simulatum
DAVENP. which is very similar to the type species P. glanduligera and has been made the type of a new
genus Wagneriopteris by LOVE and LOVE, who reported a chromosome number 31 for it, though two
previous authors had reported 32. In any event, the species here brought together are certainly not
uniform in chromosome number, and it seems probable that the genus should be subdivided, but a new
comprehensive study of all characters (including gametophytes) is necessary before this can be
satisfactorily effected.
KEY FO THE SPECIES
1. Basal pinnae, several pairs, gradually reduced, lowest 3-5 mm long . 1. P. beddomei
1. Basal pinnae, at most 2 pairs, slightly reduced.
2. Pinnae to 7.5 cm long; basal pinnae much narrowed at base
2. Pinnae to 2.0 cm long; basal pinnae not narrowed at base
1. Parathelypteris beddomei (BAK.) CHING, Acta
Phytotax. Sinica 8 (1963) 302; C. M. Kuo, FI.
Taiwan 1 (1975) 421, pl. 146.— Lastrea
gracilescens sensu BEDD. Ferns S. India (1863) t.
110.— Nephrodium beddomei BAK. Syn. Fil.
(1867) 267, nom. nov. — Lastrea beddomei (BAK.)
BEDD. Ferns Brit. India corr. (1870) 2; Handb.
(1883) 239; COPEL. Fern FI. Philip. (1960) 321. —
Dryopteris beddomei (BAK.) O. KTZE, Rev. Gen.
Pl. 2 (1891) 812; v.A.v.R. Handb. (1908) 191;
BACKER & POSTH. Varenfl. Java (1939) 37.—
Thelypteris beddomei (BAK.) CHING, Bull. Fan
Mem. Inst. Biol. Bot. 6 (1936) 308; HOLTTUM,
Rev. Fl. Mal. 2 (1955) 240; MANTON & SLEDGE,
Phil. Tr. R. Soc. B, 238 (1954) 137; K. IwarTs.
Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 166. —
Type: BEDDOME s.n. Nilgiri Mts, S. India (K).
Phegopteris smithii v.A.v.R. Bull. Dép. Agr.
Ind. Néerl. 18 (1908) 15; Handb. (1908) 490. —
Type: Java, Gedeh, Herb. Bog. 4170 (BO).
Dryopteris beddomei var. nadiwonensis v.A.v.R.
Bull. Jard. Bot. Btzg IH, 7 (1912) 13.—Type:
VERMOESEN, Java, Ngadiwono (BO).
Dryopteris microcarpa v.A.v.R. Bull. Jard. Bot.
Btzg III, 2 (1920) 147.— Thelypteris microcarpa
(v.A.v.R.) CHING, Bull. Fan Mem. Inst. Biol. Bot.
10 (1941) 252.—Type: BUNNEMEIJER 5636,
Sumatra, Padang Highlands (BO). — Fig. 5a—b.
KEY TO THE VARIETIES
1. Rhizome long-creeping; septate hairs many on
lower surface.
2. Pinnae commonly 3-5 cm long
a. var. beddomei
2. Pinnae not over 2cm long. Db. var. eugracilis
1. Rhizome usually short-creeping; few septate
hairs kw A .¢c. var. brassii
a. var. beddomei — Fig. 5a—b.
Rhizome long-creeping, 2.5 mm diameter, bear-
ing fronds 2-4 cm apart; scales broad, entire, with
superficial capitate hairs only. Stipe 5-10 cm long,
green; rachis with capitate hairs on lower surface,
. 2. P. glanduligera
3. P. grammitoides
usually also acicular hairs. Lamina commonly
20-25 cm long; pinnae many pairs, several pairs
towards base gradually smaller, lowest less than
1 cm long. Largest pinnae 3-5 cmlong, 6-8(— 10) mm
wide, incised almost to costa into oblique =
dentate lobes 1.5-2.0mm wide (basal lobes of
largest pinnae sometimes almost free and rather
deeply lobed, as in var. nadiwonensis v.A.v.R.):
lower surface of costa, costules and veins bearing
erect slender hairs 1 mm long, many consisting of
several cells, and scattered very short capitate
hairs or larger sessile yellow glands; upper surface
sometimes also with sessile glands between veins:
veins pinnate in the pinna-lobes, very oblique,
simple or (in largest lobes) forked. Sori small,
close to margin; indusia small bearing short capi-
tate hairs.
Distr. Southern India & Ceylon:
throughout Malesia.
Ecol. On mountains at 1500-2000 m, in rather
open places, on sloping ground where there is a
seepage of water; becoming more abundant with
increased felling of mountain forest.
Note. Specimens from Japan and China named
T. beddomei in Herb. Kew. appear to me to be P.
nipponica (Fr. & SAV.) CHING; they have pinna-
lobes which are flat when dried and at maturity
are covered beneath with sori, and their septate
hairs are shorter, consisting of at most 3 cells.
Taiwan:
b. var. eugracilis (COPEL.) HOLTTUM, comb.
nov. — Dryopteris gracilis COPEL. Philip. J. Sci.
40 (1929) 294, non DomiIn, 1929.— Lastrea
eugracilis COPEL. Gen. Fil. (1947) 138, nom. nov.;
Fern Fl. Philip. (1960) 321.— Thelypteris
eugracilis (COPEL.) REED, Phytologia 17 (1968)
275.— Type: ELMER 11520, Mindanao, Mt Apo
(MICH: K, L).
Differs from var. beddomei: pinnae 1.4-1.6cm
long, 4-6 mm wide near base.
Distr. Malesia: Philippines (Mindanao), 2 col-
lections, 1700-1800 m.
c. var. brassii (C. CHR.) HOLTTUM, comb. nov. —
Dryopteris brassii C. CHR. Brittonia 2 (1937)
372 FLORA MALESIANA [ser. II, vol. 1°
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INCA:
ele.
Fig. 5. Parathelypteris beddomei (BAK.) CHING. a. Rhizome and frond, x}; b. pinna-lobe, x 12.—
Trigonospora ciliata (BENTH.) HOLTTUM. c. One pinna, x 1.5; d. 2. pinna-lobes, X 8; e. spore, X 650. —
T. calcarata (BL.) HOLTTuM. f. One pinna, x 2.— Thelypteris confluens (THUNB.) MORTON. g. One
pinna, x 2; h. part of fertile pinna, x 8 (a—b HOLTTUM 26217, c-e HOLTTUM 31306, f Java in Herb. Hook.,
g—-h IWATSUKI et al. 155).
passat
1981]
295. — Lastrea brassii (C. CHR.) COPEL. Philip. J.
Sci. 78 (1951) 424. — Thelypteris brassii (C. CHR.)
REED, Phytologia 17 (1968) 265. — Type: BRASS
4937, Papua, Mt Tafa, 2400 m (BM).
Differs from var. beddomei: rhizome usually
short, with fronds + tufted; septate hairs on
lower surface few and shorter.
Distr. Malesia: Eastern New Guinea, at 2000—
3500 m.
Note. Some specimens have a partly creeping
rhizome but in almost all cases some fronds are in
rather dense tufts. It is possible that this variety
represents an adaptation to conditions at higher
altitudes and in more exposed places than the
normal habitats for var. beddomei in Western
Malesia. CHRISTENSEN wrongly stated that the
type is exindusiate.
2. Parathelypteris glanduligera (KUNZE) CHING,
Acta Phytotax. Sinica 8 (1963) 303; HOLTTUM,
Kalikasan 5 (1976) 113. — Aspidium glanduligerum
KUNZE, Analecta Pterid. (1837) 44.— Lastrea
gracilescens var. glanduligera BEDD. Handb.
Suppl. (1892) 51.—Dryopteris glanduligera
(KUNZE) CHRIST, J. de Bot. 21 (1908) 231.—
Thelypteris glanduligera (KUNZE) CHING, Bull.
Fan Mem. Inst. Biol. Bot. 6 (1936) 320; K. IWATs.
Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 167. —
Type: C. W. PETERSEN, near Canton (Herb. Leh-
mann et propria’; not found among KUNZE spe-
cimens at B).
Dryopteris repentula CHRIST, Notul. Syst. 1
(1909) 29.—Lectotype (CHING, l.c. 1936):
HENRY 13077, Yunnan (K).
Rhizome to 3 mm diameter. Stipe to 30 cm long,
slender, with many capitate and sometimes aci-
cular hairs. Lamina 20-30 cm long; pinnae 15-20
pairs, well spaced; basal pinnae not reduced, dis-
tinctly narrowed towards their bases, basal
acroscopic lobe often + elongate and sometimes
dentate. Largest pinnae 6-8.5 cm long, 1.3-1.5 cm
wide, acuminate, lobed to 0.5mm from costa;
lobes entire, subacute, falcate; costules 2-3 mm
apart; veins to 8 pairs, usually simple; lower sur-
face of rachis and costae densely hairy, hairs
erect, of varied length to 1 mm; hairs on costules
and veins more sparse, also some small scales
with swollen red terminal cell; large orange-red
glands present on and between veins; upper sur-
face lacking hairs and glands between veins. Sori
near apices of veins, rather small, distal ones
sometimes asymmetric; indusia small, bearing
glands (often many) and short hairs.
THELYPTERIDACEAE (Holttum)
373
Distr. Nepal eastwards to S. China, Korea and
Japan, and south to Thailand and N. Malesia:
Philippines (N. Luzon).
Note. The only known Malesian specimens are
from Baguio at 1200m (M. G. Price 1583) and
Zambales Province, above Palauig (PRICE 2828,
2850). In 1976 I wrongly referred the Zambales
specimens to P. grammitoides.
3. Parathelypteris grammitoides (CHRIST) HOLT-
TUM, Kalikasan Philip. J. Biol. 5 (1976) 114. —
Aspidium grammitoides CHRIST, Bull. Herb.
Boiss. 6 (1898) 193.— Thelypteris grammitoides
(CHRIST) CHING, Bull. Fan Mem. Inst. Biol. Bot.
6 (1936) 317.— Lastrea grammitoides (CHRIST)
CopEL. Fern Fl. Philip. (1960) 320.— Type:
LOHER 890, Nov. 1893, Luzon, Mt Mariveles (P:
K).
Athyrium hyalostegium COPEL. Philip. J. Sci. |
(1906) Suppl. 253. — Lectotype: COPELAND 2033,
Luzon, Mt Mariveles (MICH; P, US).
Rhizome to 2mm diameter. Stipes not widely-
spaced, 3-10 cm long, with acicular hairs 0.3 mm
long throughout; basal scales narrow, 1.5 mm long,
bearing small glands, distal ones bearing also short
acicular hairs. Lamina 5-8cm long; pinnae 6-8
pairs; lowest pinnae not or little smaller than next
pair, not narrowed at base, basal acroscopic lobe
always free and = crenate, basal basiscopic lobe
usually also free; apex of lamina gradually
attenuate. Largest pinnae 2.0x0.8cm, lobed al-
most to costa; basal acroscopic lobe almost at
right angles to costa, basiscopic and other acros-
copic lobes at c. 45°, with sinuous edges; apex of
pinna evenly attenuate; costules to 3 mm apart:
veins 4-5 pairs in largest lobes; lower surface of
rachis and costae bearing scattered slender hairs
of mixed length to | mm long, glands present on
and between veins; hairs on upper surface of
costae 0.5mm long, few hairs on costules. Sori
supramedial to medial; indusia small, thin, with
rather long slender hairs and a few glands.
Distr. Malesia: Philippines (Luzon).
Ecol. At 900-1400m; on Mt San Cristobal
found ‘‘at edge of crater lake, in shade with mos-
ses” (M. G. PRICE).
Note. This species is closely allied to P.
angustifrons (MIQ.) CHING from the Ryukyu
Islands and Taiwan, but the Philippine plants ap-
pear always to be much smaller than those from
Taiwan. P. crystopteroides (EATON) CHING in
Japan and the northern Ryukyu Islands is also
closely related.
7. TRIGONOSPORA
HOLTTUM, Blumea 19 (1971) 29; Reinwardtia 8 (1974) 503. — Pseudocyclo-
sorus CHING, Acta Phytotax. Sinica 8 (1963) 322, p.p. — Fig. 5c-f.
Caudex short, erect; lamina of fronds commonly to 20cm long, lacking
FLORA MALESIANA [ser. II, vol. 1°
374
reduced basal pinnae; veins in Malesian species free, unbranched, lowest
acroscopic vein ending beside short sinus-membrane, basal basiscopic vein
to edge above base of sinus; acicular unicellular hairs variously developed
on lower surface; sori indusiate; glands and hairs lacking on body of
sporangium, on its stalk usually a hair of several cells ending in a gland;
spores trilete, minutely papillose.
Type species: Trigonospora ciliata (BENTH.) HOLTTUM.
Distr. Ceylon, India except north-west, Burma to Kwangtung and southwards to Malesia; in
Malesia: Malaya northwards from 4°N, N. Sumatra, S. Sumatra and Java, N. Celebes. About 8 species,
not yet all described.
Ecol. On rocks in and beside streams in mountain forest.
Cytol. Base chromosome number 36: T. ciliata diploid in Malaya; species of uncertain identity
diploid and tetraploid in Ceylon and S. India.
Notes. This is the only group of species in the family in which trilete spores are normal, though such
have been observed as occasional in Macrothelypteris (P. CHANDRA, Amer. Fern J. 63, 1973, 9). In their
shape and in their minutely papillose perispore they are closely similar to the spores of the monotypic
African genus Menisorus ALSTON which are either spherical or monolete with a short laesura. Plants of
Menisorus also have a short erect caudex and grow on rocks in streams; I believe that the two genera
are related. An unnamed species in Ceylon has short capitate hairs, like those of Pseudocyclosorus, on
the lower surface and on indusia.
In India and Ceylon this genus is much diversified in shape and size of fronds and of their pinnae, and
in pubescence, but (apart from T. zeylanica CHING) individual species have not yet been clearly
distinguished, probably because intermediates bridge some of the gaps between them. The existence of a
tetraploid in S. India indicates hybridization; the habitat on wet rocks would facilitate this. One Ceylon
specimen is very like T. calcarata of Java in shape of pinnae but much larger; others in Ceylon are very
different. Hooker included most Indian specimens in Nephrodium calcaratum; BEDDOME added
N. falcilobum HOOK. (Pseudocyclosorus falcilobus CHING) as a variety, though it differs in its reduced
basal pinnae. HOOKER cited the type of T. zeylanica as a variety under N. falcilobum and expressed
uncertainty as to a distinction between N. calcaratum and N. falcilobum.
Thelypteris knamptorum HOLTTUM (Kew Bull. 26, 1971, 82) has crenate pinnae and anastomosing
veins, but agrees in spores with Trigonospora; two collections are known from Upper Burma.
KEY TO THE SPECIES
1. Edges of pinnae lobed to less than 1 mm from costa, lobes narrow with wide sinuses between them;
indusia glabrous.
2. Pinnae 3-4cm long, 10-12 mm wide above auricled base; lobes (except basal acroscopic one) not
narrowed towards their bases
. 1. T. calcarata
2. Pinnae to 8x 2cm, basal acroscopic lobe not much elongate: pinna- lobes widest 1/3 from their
apices
2. T. koordersii
. Edges of pinnae lobed to ie Be 2mm from costa; lobes wider than sinuses between them; indusia
hairy
1. Trigonospora calcarata (BL.) HOLTTUM, Rein-
wardtia 8 (1974) 506. — Aspidium calcaratum BL.
Enum. Pl. Jav. (1828) 159; RAciB. Fl. Btzg 1
(1898) 170.—Nephrodium calcaratum (BL.)
HOOK. Spec. Fil. 4 (1862) 93, p.p.— Dryopteris
calcarata (BL.) O. KTZE, Rev. Gen. Pl. 2 (1891)
812; v.A.v.R. Handb. (1908) 185, p.p.; BACKER &
POsTH. Varenfl. Java (1939) 38.— Thelypteris
calcarata (BL.) CHING, Bull. Fan Mem. Inst. Biol.
Bot. 6 (1936) 288. — Type: BLUME, Java (L).
Aspidium reinwardtianum KUNZE ex METT.
Farngatt. IV (1858) 86.— Lectotype (HOLTTUM
1974): ZOLLINGER 1600 (B; FI, G).
Dryopteris marthae v.A.v.R. Bull. Jard. Bot.
3. T. ciliata
Btzg II, 1 (1911) 7.—Type:
Slamat, 900 m (BO). — Fig. 5f.
Stipe 8-20cm long, dark at base, glabrescent
except in groove. Lamina 10—20cm long; pinnae
to 12 pairs. Largest pinnae 3-4 cm long, 10-12 mm
wide above base, lobed to less than 1mm from
costa; lobes c. 8 pairs, narrow, very oblique
except basal acroscopic one which is elongate,
close to rachis and almost free; veins c. 6 pairs;
rachis, costae and costules sparsely hairy on lower
surface. Sori near costules; indusia dark, firm,
glabrous; sporangia lacking gland-bearing hairs on
their stalks.
Distr. Ceylon and Malesia: throughout Java at
BACKER 262, G.
1981]
100-1000 m, southern Sumatra and north to Padang
Highlands.
2. Trigonospora koordersii (CHRIST) HOLTTUM,
Reinwardtia 8 (1974) 506.— Aspidium koordersii
CHRIST, Ann. Jard. Bot. Btzg 15 (1897) 128, t.15,
f.17.— Dryopteris koordersii (CHRIST) C. CHR.
Ind. Fil. (1905) 273; v.A.v.R. Handb. (1908) 181. —
Type: KOORDERS 1700, Celebes, near Karawutu,
50m (P; BO, L).
Stipe 15cm, glabrous. Lamina to 24cm long;
pinnae 8 pairs. Largest pinnae 8 X 2.cm, distinctly
stalked, lobed almost to costa; lobes at 45°, to
15mm long, 1-2 mm wide, widest 1/3 from apex
and narrowed towards base, basal acroscopic lobe
not much longer than next; sparse rather thick
hairs on lower surface of costules only; veins 8-12
pairs. Sori medial; indusia firm, dark, glabrous;
sporangia short-stalked, lacking a glandular hair
on the stalk.
Distr. Malesia: N.E. Celebes: only known
from the type.
3. Trigonospora ciliata (BENTH.) HOLTTUM,
Blumea 19 (1971) 29. — Lastrea ciliata HOOK. J.
Bot. Kew Misc. 9 (1857) 338, non LIEBM. —
Aspidium ciliatum BENTH. FI. Hongkong. (1861)
455.—Lastrea calcarata var. ciliata BEDD.
THELYPTERIDACEAE (Holttum)
375
Handb. (1883) 235, p.p.—Dryopteris ciliata
(BENTH.) C. CHR. ex WU et al. Bull. Dept. Biol.
Sunyatsen Univ. n. 3 (1922) 30, pl. 6.— Thelyp-
teris ciliata (BENTH.) CHING, Bull. Fan Mem.
Inst. Biol. Bot. 6 (1936) 289; C.C. Woop, J. Linn.
Soc. Bot. 67, Suppl. 1 (1973) pl. 2C.— Pseudo-
cyclosorus ciliatus (BENTH.) CHING, Acta Phy-
totax. Sinica 8 (1963) 324.— Lectotype (HOLT-
TUM 1974): “BOWMAN” (BOWRING), Hong Kong
(K).
Dryopteris pinnata COPEL. Univ. Cal. Publ.
Bot. 14 (1929) 373. — Thelypteris pinnata (COPEL.)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941)
253.— Type: BARTLETT 6641, Sumatra, Asahan
river (UC; L). — Fig. 5c-e.
Stipe of sterile fronds 10-15cm, of fertile to
25cm long. Lamina to 20cm long; pinnae c. 14
pairs. Largest pinnae 4-5 cm long, 1.0-1.2 cm wide
above the base, lobed to 1.5-2mm from costa;
lobes oblique, rather triangular, basal acroscopic
lobe a little elongate; veins 5-6 pairs; lower sur-
face of rachis and costae bearing copious soft pale
spreading hairs 1 mm long, shorter ones on cos-
tules and veins. Sori near costules; indusia large,
very hairy; stalk of sporangium bearing a hair of 3
cells, end cell glandular.
Distr. N. India and S. China southwards to
Malesia: N. Malaya and Sumatra.
8. THELYPTERIS
SCHMIDEL, Icon. Pl. ed. Keller (1763) 45, t. 11, 13, nom. cons.; FERNALD,
Rhodora 31 (1929) 27-36, pl. 179-180; CHING, Acta Phytotax. Sinica 8
(1963) 297 excl. T. grisea (BAK.) CHING; HOLTTUM, Taxon 17 (1968) 330;
Blumea 19 (1971) 28; A.F. TRYON, Rhodora 73 (1971) 444. — Fig. 5g—h.
Rhizome long-creeping, in wet ground; fronds simply pinnate with deeply
lobed pinnae, basal ones not reduced; veins free, usually forked, all
reaching margin; flat thin scales present on lower surface of costae; sori
indusiate; sporangia bearing short capitate hairs; perispore of spores varied
(see A.F. TRYON lI.c.).
Type species: Thelypteris palustris SCHOTT.
Distr. North temperate Eurasia, in Asia south to Himalayas; eastern temperate and subtropical N.
America; Africa south of the Sahara; S. India; N. Thailand; in Malesia: Sumatra and New Guinea; New
Zealand. Two species, northern and southern, the former with geographically distinct subspecies.
Cytol. Base chromosome number 35; many observations in north temperate regions, all diploid.
Notes. This genus, as restricted by CHING but excluding his T. grisea, is the temperate counterpart of
Cyclosorus (in the restricted sense here adopted). Both are rhizomatous ferns growing in open fresh-water
swamps. They also agree in having fronds with unreduced basal pinnae and in bearing flat scales on lower
surface of costae, a character uncommon in the family. In the present work, Cyclosorus is associated with
Ampelopteris and Mesophlebion; from these three Thelypteris differs in (a) capitate hairs on sporangia, (b)
lack of red glands at the ends of hairs on sporangium-stalk, (c) chromosome number 35, (d) basiscopic vein
always from costule, not from costa. But I suggest that Thelypteris is nearest to this group of genera.
CHING included Nephrodium griseum BAK. (see BEDD. Ferns Brit. India t.335, Handb. 241) as a
species of his restricted Thelypteris. The type of this is a specimen at Kew from Cochin which
resembles T. palustris in its forked veins but in little else. BEDDOME wrote a note on it in his own
Herbarium (now at Kew) and suggested that it was probably an abnormal form of Lastrea calcarata var.
376 FLORA MALESIANA [ser. II, vol. 1°
/\| \
WAV MIN
Fig. 6. Mesophlebion oligodictyon (BAK.) HOLTTUM. a. Whole plant, x 33 b. venation of sterile frond,
x 4. — M. trichopodum (C. CHR.) HOLTTUM. c. Rachis and basal pinna, x3; d. two pinna-lobes, hairs
omitted, x4; e. costa, costule and sorus, x 24.— M. beccarianum (CESATI) HOLTTUM. f. Suprabasal
pinna, X3; g. venation and sori, < 8.— M. endertii (C. CHR.) HOLTTUM. h. One pinna, x3 (a JACOBS
5600, b C. HOSE 210, c-e MATTHEW s.n., f-g LEE & MAHMUD UL37, h ENDERT 4433).
1981]
THELYPTERIDACEAE (Holttum)
og)
ciliata (Trigonospora ciliata of the present work) but in my opinion it is an abnormal form of a species
of Pseudocyclosorus.
The best account of the genus in the northern hemisphere is by FERNALD, l.c.
1. Thelypteris confluens (THUNB.) MORTON,
Contr. U.S. Nat. Herb 38 (1967) 71; SCHELPE, FI.
Zambes. Pterid. (1970) 190, t. 5SE; HOLTTUM,
J.S. Afr. Bot. 40 (1974) 150; Allertonia 1 (1977)
180. — Pteris confluens THUNB. Prodr. Fl. Cap.
(1800) 171. — Type: THUNBERG, S. Africa (UPS).
Aspidium thelypteris var. squamigerum
SCHLECHTEND. Adumb. Fil. Prom. B. Spei
(1825) 23, t.l1l.— Aspidium squamigerum FEE,
Mém. Fam. Foug. 8 (1857) 104. — Lastrea thelyp-
teris var. squamigera BEDD. Handb. Suppl. (1892)
54.— Dryopteris thelypteris var. squamigera C.
CHR. Ind. Fil. (1905) 297.—T. palustris var.
squamigera WEATH. Contr. Gray Herb. n.s. 73
(1924) 40. — T. squamigera (err. typ. squamulosa)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 6 (1936)
329; TARD. in Humbert, Fl. Madag. Se Fam. |
(1958) 282.—Type: SCHLECHTENDAL, Cape
Peninsula (HAL).
Nephrodium squamulosum HOoK. f. Fl. New
Zeal. 2 (1855) 39.— Nephrodium thelypteris var.
squamulosum Hook. Spec. Fil. 4 (1862) 88.—
Type: COLENSO, New Zealand (K).
Lastrea fairbankii BEDD. Ferns Br. Ind. (1867)
t.254; Handb. (1883) 240.— Type: BEDDOME, S.
India, Pulney Hills (K). — Fig. 5g-h.
Rhizome 2mm _ diameter (dry). Stipe
stramineous, 15-S0cm long. Lamina 25-50cm
long; pinnae to 20 pairs or more, dimorphous.
Sterile pinnae commonly 4-6 1.5cm, fertile 6-
9mm wide; lowest pinnae slightly reduced, some-
times with a free basal acroscopic lobe; middle
pinnae not dilated at base, lobed to less than 1 mm
from costa; veins in sterile fronds mostly forked,
in fertile often simple; lower surface of costae
bearing slender hairs and ovate-orbicular flat cili-
ate scales 0.5—1 mm long and wide; lower surface
between veins bearing a variable number of slen-
der acicular and short capitate hairs; hairs on
upper surface of costa short. Sori near costules;
indusia with short marginal hairs.
Distr. Africa south of the equator and to 7°N
in Ethiopia and Sudan; Madagascar; S. India; N.
Thailand and Laos; in Malesia: N. Sumatra in
Atjeh, near Lake Toba and near G. Kerinci;
Western New Guinea on Arfak Mts, Eastern New
Guinea in W. Highlands; New Zealand (North
Island).
Ecol. In N. Sumatra at 1180-1330m, in W.
New Guinea at 1750-1900 m, in E. New Guinea at
2500 m; in all cases in swampy open ground near
lakes.
9. MESOPHLEBION
HOLTTUM, Blumea 19 (1971) 29 (new name for Mesoneuron CHING, non
DEsF.), excl. subg. Plesioneuron. — Group of Dryopteris crassifolia (BL.) O.
KTZE, C. CHR. Gard. Bull. Str. Settl. 4 (1929) 381; of Thelypteris crassifolia
(BL.) CHING, HOLTTUM, Rev. FI. Mal. 2 (1955) 245-249, f. 139-141. —
Mesoneuron CHING, Acta Phytotax. Sinica 8 (1963) 325, excl. M.
attenuatum. — Thelypteris subg. Glaphyropteridopsis sect. Mesoneuron K.
IWATS. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 30, excl. Pseudocyclo-
sorus CHING. — Fig. 1k, 6.
Caudex creeping, sometimes elongate; scales narrow, rather rigid but
never spine-like, bearing short acicular hairs; similar scales at base of stipe,
in a few species throughout stipe and basal part of abaxial surface of rachis.
Fronds simply pinnate (in M. oligodictyon simple, in M. endertii partly
bipinnate), sometimes dimorphic; basal pinnae not reduced but + narrowed
towards their asymmetric bases; aerophores often distinctly swollen on
living fronds, collapsing on drying; all pinnae lobed, in some cases deeply;
veins all simple and free (except casually in M. oligodictyon and M.
motleyanum), basal basiscopic vein arising from costa near the base of the
costule to which it belongs; basal veins from adjacent costules usually both
passing to base of sinus between their lobes, touching sides of the sinus-
membrane, which may be slightly decurrent between them as a hairy ridge;
lower surface of costae, costules and veins always bearing many narrow
378 FLORA MALESIANA [ser. Il, vole?
scales in various degrees of reduction (smallest of 2 or 3 cells) in addition to
a varied complement of acicular unicellular (sometimes also capitate) hairs
and in some cases sessile spherical orange glands; upper surface hairy on
costa only; sori in almost all cases indusiate, the indusia in most cases thin
and sometimes small; young sporangia often pale violet-purple; large
spherical orange-red glands attached directly or by a hair to stalks of
sporangia, no glands nor hairs on body of sporangia; spores in most species
with a thin continuous wing and a few cross-wings, papillose without a wing
in M. beccarianum and M. chlamydophorum.
Type species: Mesophlebion crassifolium (BL.) HOLTTUM.
Distr. Peninsular Thailand and southern Burma, Malesia: except E. Java and Lesser Sunda Islands
(greatest diversity in Borneo), few specimens from New Guinea.
Ecol. In forest, apart from M. teuscheri and M. arenicola (on wet sandstone in exposed places) and
M. oligodictyon (in rocky stream-beds), at 0-1500 m.
Cytol. Base chromosome number 36; M. crassifolium and M. trichopodum both tetraploid in Malaya
(MANTON in Holttum 1955), M. falcatilobum diploid in Sarawak (T. G. WALKER, new obs.)
Taxon. Species of this genus were first recognized as forming a distinct group by CHRISTENSEN
(1929). CHING transferred two of them to Thelypteris in 1936, and the species of Malaya were treated
under that genus by HOLTTUM in 1955. In 1963 CHING established the genus Mesoneuron for them, but
as Mezonevron DESF. has also been spelled Mesoneuron I proposed the new name Mesophlebion in
1971; in doing so I included Plesioneuron as a subgenus.
Mesophlebion agrees with Cyclosorus (in the strict sense of the present account) and Ampelopteris in
the large red glands which occur at the ends of hairs on stalks of sporangia and sometimes on the lower
surface of veins, also in the presence of scales on the lower surface of costae. In my judgement, these
two genera and Thelypteris are the only near relatives of Mesophlebion.
In tropical America occur two species which CHRISTENSEN (Monogr. Dryopteris 1, 1913, 170-173)
placed with doubt in Dryopteris subg. Steiropteris; BAKER confused Mesophlebion motleyanum with
one of these and was copied by other authors. The American species differ from Mesophlebion in
having a long sinus-membrane and long aerophores, and in the absence of red glands.
Several species of Mesophlebion are variable, and it is possible that a complex of diploids, triploids
and tetraploids may exist. Experimental studies are needed to clarify this situation. I judge that the
species here recognized are reasonably distinct, though probably not all of equal status, and hybrids may
occur. Several plants of both M. motleyanum and M. falcatilobum, from Gunong Mulu in Sarawak, have
shown consistently distinct characters in cultivation at Kew.
It is notable that, as in Plesioneuron and Chingia, a few species of this genus have abundant scales
throughout the stipe and at least the basal part of the rachis. This development in the three genera
appears to be a case of parallel evolution; a similar condition occurs in one non-Malesian species of
Christella (C. crinipes (HOOK.) HOLTTUM) to which RIDLEY referred specimens of M. trichopodum.
KEY, HO MGEES SPE EHES
1. Stipe and lower part of rachis bearing numerous stiff spreading scales.
2. Pinnae to 12 cm long.
3. Stipe-scales dark, to | mm wide; veins 7-10 pairs ; 7 oe ee Ue le I eMivechinaton
3. Stipe-scales medium brown, to 2mm wide; veins 12-17 pairs . . . . 2, M. persquamiferum
2. Pinnae of well-grown plants much longer.
4. Costae, costules and veins beneath bearing short hairs only; stipe-scales to 6 mm long.
5. Lower pinnae not auricled; no glands on lower surface of costules . . . . . 3. M. hallieri
5. Lower pinnae strongly auricled; copious glands on lower surface of costules . . 4. M. caroli
4. Costae, costules and veins beneath bearing hairs | mm or more long, at least on sterile fronds;
stipe-scales 10mm or more long . . . So oe 4. w » oe oy SoMentrichopodum
Upper part of stipe and rachis acl large jealtss.
es Fronds simple . . SHAME ELOY SEERA LOG eMeolizodictyon
6. Fronds pinnate or partly bipinnate.
7. Pinnae not more than 3 cm long.
8. Pinnae lobed less than half-way to costa; acroscopic bases of pinnae auricled 7. M. teuscheri
8. Pinnae lobed more than half-way to costa, not auricled . . . . . . . . 8. M. arenicola
1981]
THELYPTERIDACEAE (Holttum)
379
7. Pinnae much longer.
9. Largest pinnae with several pairs of pinnules
9. M. endertii
9. Largest pinnae with at most one free basal lobe.
10. Sterile pinnae lobed not more than half-way to costa, except sometimes basal pinnae.
11. Pinnae 6—8(-10) pairs; young fronds red
11. Pinnae 14-18 pairs; young fronds not red
10. M. motleyanum
11. M. faleatilobum
10. Sterile pinnae lobed more than half-way to costa.
12. Pinnae to 6.5 cm wide, lower ones with stalks 3-4 cm long :
12. Pinnae not more than 4cm wide, lower ones with stalks to 1.5 cm long.
12. M. dulitense
13. Hairs on lower surface of costae and costules c. | mm long.
14. All pinnae distinctly stalked; a short hairs and glands between veins on lower
surface
. 13. M. auriculiferum
14. Pinnae sessile; epese erect nee 0. 5 mm lone aa no verenids between veins on lower surface
5. M. trichopodum
13. Hairs on lower surface of costae and costules not over 0.3 mm long.
15. Indusia firm, glabrous, covering the sorus almost to maturity; fertile ee lobed to 1-15 mm
from costa
. M. beccarianum
15. Indusia small, not covering developing sori, or + hairy: fertile pinnae “git less deeply.
16. Fertile pinnae lobed to 2 mm from costa; indusia small, firm, glabrous
15. M. rufescens
16. Fertile pinnae lobed less deeply; indusia usually with some hairs, thin and almost obscured
by sporangia if small.
17. Fronds rarely dimorphic; pinnae thin,
conspicuous, firm, bearing a variable number of short hairs
rarely more than 2cm wide; indusia always
16. M. chlamydophorum
17. Fronds usually dimorphic; pinnae rigid, fertile to 3 cm, sterile to 4cm wide; indusia varied
in size, thin and shrivelling when dried
1. Mesophlebion echinatum (METT.) HOLTTUM,
Blumea 22 (1975) 226.— Aspidium echinatum
METT. Ann. Mus. Bot. Lugd.-Bat. | (1864) 230;
MIQUEL, ibid. 4 (1868) 157. — Dryopteris echinata
(MemmnO. Kitz": “Revs Gen: Pl. 2) (1891) «812:
v.A.v.R. Handb. (1908) 182; C. CHR. Gard. Bull.
Str. Settl. 4 (1929) 386. — Mesoneuron echinatum
(METT.) CHING, Acta Phytotax. Sinica 8 (1963)
326. — Thelypteris echinata (METT.) REED, Phy-
tologia 17 (1968) 274. — Type: KORTHALS 498, G.
Prarawin, near Bandjermasin, Kalimantan (L, n.
908, 335-575).
Caudex short-creeping; stipe to 30cm long,
bearing throughout (also basal part of rachis and
near bases of costae on lower surface) spreading
dark scales to 10mm long, those of stipe 1mm
wide at base, on rachis narrower. Lamina of type
30cm long (of another specimen 70cm); basal
pinnae slightly narrowed at base with stalks 2mm
long. Largest pinnae 11-15cm long, 1.5-2.0cm
wide, acuminate with subentire cauda 10-15 mm
long, lobed to less than 2 mm from costa; costules
3.5 mm apart, at more than 60° to costa; veins 7-10
pairs, basal basiscopic vein arising from costa
near base of its costule; rachis and costae densely
short-hairy on lower surface, hairs sparse on cos-
tules and veins, large red glands present on cos-
tules. Sori inframedial; indusia small, firm,
caducous, with a few short hairs or glabrous.
Distr. Malesia: Borneo (Kalimantan,
Sarawak), N. Sumatra.
Ecol. Lowland forest.
Note. The Sumatran specimen (DE WILDE
12635) has the basal basiscopic vein arising from
the costa rather far from the base of its costule.
17. M. crassifolium
2. Mesophlebion persquamiferum (v.A.v.R.)
HOLtTtTuM, Blumea 22 (1975) 226.— Dryopteris
persquamifera v.A.v.R. Bull. Jard. Bot. Btzg II, 16
(1914) 10; Handb. Suppl. (1917) 149.—
Mesoneuron persquamiferum (v.A.v.R.) CHING,
Acta Phytotax. Sinica 8 (1963) 326.— Type:
RACHMAT 489, Somalilah, Celebes (BO; L).
Aspidium echinatum sensu CHRIST, Ann. Jard.
Bot. Btzg 15 (1898) 128.
Stipe 35-40cm long, densely scaly; scales
medium brown, to 8 x 2 mm, with thick persistent
bases; similar scales on rachis. Lamina 50cm
long; pinnae c. 20 pairs; basal pinnae slightly
narrowed at base, short-stalked. Largest pinnae
12.5 x 2.0-2.5 cm; base slightly dilated, apex short-
acuminate; edges lobed to 2 mm from costa, lobes
falcate; costules to 4mm apart, at more than 60°;
veins 12-17 pairs, basal basiscopic vein from costa
distant from its costule; rachis scaly beneath al-
most throughout; costae and costules almost hair-
less beneath with many narrow scales and sessile
glands, some glands also on surface between
veins. Sori inframedial; indusia small, short-hairy,
often caducous.
Distr. Malesia: Central Celebes, 2 collections
(SARASIN 952, Takale Kadjo, 900 m; BAS).
3. Mesophlebion hallieri (CHRIST) HOLTTUM,
Blumea 22 (1975) 227.—Aspidium _ hallieri
CHRIST, Ann. Jard. Bot. Btzg 20 (1905) 106. —
Dryopteris hallieri (CHRIST) C. CHR. Ind. Fil.
(1905) 269; Gard. Bull. Str. Settl. 4 (1929) 387;
v.A.v.R. Handb. (1908) 184. — Mesoneuron hallieri
(CHRIST) CHING, Acta Phytotax. Sinica 8 (1963)
326. — Thelypteris hallieri (CHRIST) REED, Phy-
380
FLORA MALESIANA
tologia 17 (1968) 281.— Type: HALLIER 3204,
Amai Ambit, W. Kalimantan (P; BO, L).
Stipe densely covered with short reddish hairs,
also scaly throughout; scales to 6 x 1mm. Lamina
slightly dimorphous, to 100cm long; pinnae to 25
pairs; lowest pinnae not seen. Largest pinnae
23 cm long, sterile ones to 3cm wide, fertile nar-
rower; base of middle pinnae truncate, somewhat
dilated both sides; apex acuminate with cauda to
2cm long; edges lobed to 3 mm from costa, lobes
falcate at tips; costules 4.5-5 mm apart, at more
than 60°; veins 12-15 pairs, pale and very prom-
inent beneath, basal basiscopic vein arising far
from its costule; rachis densely short-hairy on
lower surface, with scales as stipe, costae short-
hairy with smaller scales, some short hairs on
veins and surface. Sori inframedial; indusia small,
thin, hairy.
Distr. Malesia: West Borneo.
4. Mesophlebion caroli HOLTTUM, Blumea 22
(1975) 227. — Type: C. HOSE 712, Baram district,
Sarawak (K; BM).
Caudex not seen; scales on upper part of stipe
and rachis medium brown, firm, 6 x 0.5 mm. Basal
pinnae much narrowed towards base both sides,
with enlarged almost free basal acroscopic lobe
9mm long, basal basiscopic lobe attached 3 mm
from base of pinna; suprabasal pinnae with basal
acroscopic lobe to 1.8 cm long, basiscopic 0.9 cm;
upper pinnae with slightly elongate basal acros-
copic lobes, basiscopic ones not reduced. Largest
pinnae 25 x 2.5 cm, caudate-acuminate with cauda
3.0cm long, 5mm wide; edges lobed to 2.5mm
from costa; lobes slightly falcate with rounded
tips; costules 5-5.5mm apart; veins to 13 pairs,
basal veins touching sides of short sinus-mem-
brane, basal basiscopic vein arising far from its
costule; lower surface of rachis and costae bear-
ing narrow scales 24mm long and sparse short
spreading hairs, many large red glands present on
costae and costules, much-reduced scales of
various sizes on costules and veins. Sori in-
framedial; indusia firm, of medium size, glabrous
or rarely with a short hair.
Distr. Malesia: Borneo. Two collections, the
second near Sambas in N.W. Kalimantan.
5. Mesophlebion trichopodum (C. CHR.) HOLT-
TUM, Blumea 22 (1975) 226. — Dryopteris tricho-
poda C. Cur. Ind. Fil. (1905) 298, new name for
Nephrodium polytrichum BAK. J. Bot. 29 (1891)
107, non SCHRAD. 1824; C. CHR. Gard. Bull. Str.
Settl 4 (1929) 397.—Dryopteris polytricha
v.A.v.R. Handb. (1908) 187. — Mesoneuron tricho-
podum (C. CHR.) CHING, Acta Phytotax. Sinica 8
(1963) 326. — Type: G. F. HOSE s.n. Lingga Mts,
Sarawak (K).
Dryopteris paleata COPEL. Phillip. J. Sci. 9
(1914) Bot. 228; C. CHR. Gard. Bull. Str. Settl. 4
(1929) 383, 387.— Thelypteris paleata (COPEL.)
[ser. II, vol. 1°
HOLTTUM, Rev. FI. Mal. 2 (1955) 249, f. 141. —
Mesoneuron paleatum (COPEL.) CHING, Acta
Phytotax. Sinica 8 (1963) 326.— Type: BROOKS
136, Lebong Tandai, Benkoelen, Sumatra (MICH).
Nephrodium crinipes sensu RIDL. J. Mal. Br. R.
As. Soc. 4 (1926) 74. — Fig. 6c-e.
Caudex thick, short-creeping; stipe 60cm long,
1.2 cm diameter at base, densely scaly throughout,
scales dark brown, rigid, commonly 10mm long,
1-1.5mm wide at base, some of them narrowed
above base, many small scales also present.
Lamina to 100 cm long; pinnae 20-25 pairs; basal
pinnae narrowed a little near their bases, those of
type 30x4.3cm (sterile). Suprabasal pinnae
commonly to 20cm long, 2.5-3.5 cm wide (sterile
wider than fertile), caudate-acuminate, lobed to
3-4 mm from costa, lobes falcate, basal basiscopic
lobe much longer than acroscopic and curved;
costules 4.5-Smm_ apart (fertile) 6-7.5mm
(sterile); veins to 23 on basiscopic side of largest
sterile lobe, to 18 on fertile lobes (to 15 on acros-
copic side), basal basiscopic vein arising from
costa far from its costule, ends of basal veins
closely parallel to each other on either side of a
narrow sinus-membrane; lower surface of rachis
densely scaly in basal part, throughout with close
short hairs, costae similar, veins on sterile pinnae
bearing slender hairs | mm long (sometimes shor-
ter on fertile pinnae) and very short capitate hairs,
sometimes also red glands. Sori inframedial; in-
dusia small but conspicuous, thin, with many hairs
0.3 mm long.
Distr. Malesia: Borneo, Malaya, Sumatra.
Ecol. Near streams in forest, at 0-1000 m.
Notes. Specimens from Malaya are all smaller
than those from Borneo and Sumatra but are
otherwise not different. Some specimens from G.
Mulu, Sarawak, have fewer, smaller, stipe-scales
and many glands on the lower surface of veins.
6. Mesophlebion oligodictyon (BAK.) HOLTTUM,
Blumea 22 (1975) 227.— Acrostichum oligodic-
tyon BAK. J. Linn. Soc. Bot. 24 (1887) 261.—
Leptochilus oligodictyus (BAK.) C. CHR. Ind. Fil.
(1905) 387; v.A.v.R. Handb. (1908) 735.—
Dryopteris oligodictya (BAK.) C. CHR. Mitt. Inst.
Allgem. Bot. Hamb. 7 (1928) 148. — Cyclosorus
oligodictyus (BAK.) HOLTTUM, Blumea 11 (1962)
530.— Thelypteris oligodictya (BAK.) REED,
Phytologia 17 (1968) 298.— Type: C. Hose 110,
Sarawak, Niah (K).
Syngramma angusta COPEL. Philip. J. Sci. 3
(1909) Bot. 348; v.A.v.R. Handb. Suppl. (1917)
330.— Type: BROOKS s.n. July 1908, Bidi,
Sarawak (MICH; BM, K). — Fig. 6a—b.
Caudex short-creeping, diameter 2.5-3 mm,
scales 2mm long; stipes closely seriate, of sterile
fronds to 7cm long, of fertile 14-18 cm. Lamina
simple, dimorphous, gradually attenuated to both
ends, entire or with slightly sinuous edges which
are strongly cartilaginous. Sterile lamina 20-28 cm
1981]
THELYPTERIDACEAE (Holttum)
381
long, 1.5-1.9cm wide; veins thick and prominent
on lower surface, usually 3 pairs to each group,
basal basiscopic vein arising from midrib of frond
below the rest (lowest acroscopic one also some-
times from midrib), basal veins of adjacent groups
anastomosing irregularly near margin; lower sur-
face quite glabrous apart from small narrow
scales at the base of each group of veins. Fertile
lamina 20-24cm long, 5-6mm wide; sporangia
borne all along the veins, without indusia.
Distr. Malesia: Borneo (Sarawak, Brunei,
Kalimantan), at least 6 collections.
Ecol. On rocks in river bed, low altitudes.
Notes. The evidence that this species belongs
to the genus Mesophlebion is provided by the
gland-bearing hairs on sporangium stalks, also by
the basal veins of each group arising from the
midrib in the same way that basal veins of pinnate
species arise from the costae below the costules.
7. Mesophlebion teuscheri (v.A.v.R.) HOLT-
TUM, Blumea 22 (1975) 228.— Dryopteris teus-
cheri v.A.v.R. Bull. Dép. Agr. Ind. Néerl. 18
(1908) 6; Handb. (1908) 183; C. CHR. Gard. Bull.
Str. Settl. 4 (1929) 383, p.p.— Mesoneuron teus-
cheri (v.A.v.R.) CHING, Acta Phytotax. Sinica 8
(1963) 326.— Thelypteris teuscheri (v.A.v.R.)
REED, Phytologia 17 (1968) 319.— Type: TEUus-
CHER, W. Kalimantan (BO).
Caudex suberect; stipe 5-10cm long, densely
short-hairy, basal scales narrow. Lamina 15-
20cm long; pinnae 20 pairs; 2-3 pairs lower pin-
nae gradually a little narrowed towards their
bases, basal acroscopic lobe of lowest pinnae free,
entire, obovate. Largest pinnae 2.2 cm long, 8 mm
wide above auricled base; apex obtuse; edges
lobed 1/3-1/2 towards costae; costules to 2.5mm
apart; veins to 3 pairs, basal veins not quite meet-
ing at sinus; lower surfaces — short-hairy
throughout, with some large red glands. Sori
medial; indusia densely hairy; sporangia with a
sessile gland on the stalk.
Distr. Malesia: W. Borneo; known from type
collection only.
Note. CHRISTENSEN’s description of 1929
was based on the original description and on small
specimens of M. beccarianum from Borneo and
M. arenicola from Sumatra; he had not seen
TEUSCHER’S specimen.
8. Mesophlebion arenicola HOLTTUM, Blumea 22
(1975) 228.— Type: W. MEIJER 4478, Pajakum-
buh-Taram, Harau Canyon, Sumatra (L).
Caudex short, suberect; stipe 5-15cm long,
minutely hairy with scattered longer hairs, basal
scales very small, dark, with short stiff hairs.
Lamina to 25cm long; pinnae to 15 pairs; basal
pinnae slightly reduced with stalks to 1 mm long.
Largest pinnae 3.5 1.3cm (sterile) 3.00.8 cm
(fertile); base truncate, not auricled; apex a short
broad tip; edges lobed to 1 mm from costa or more
deeply, lobes oblique; costules to 2.5mm apart;
veins 4 pairs, basal ones in lobes near base of
pinnae both to sides of a short sinus-membrane, in
distal lobes both to edge above base of sinus;
lower surface of rachis and costae covered with
short hairs and scattered long ones, whole lower
surface of pinnae closely covered with erect short
hairs, among them many sessile orange glands.
Sori medial, filling lower surface of fertile pinnae;
indusia densely hairy; sporangia with short stalks,
hairs on stalks commonly of 3 cells with terminal
gland.
Distr. Malesia:
tions).
Ecol. On sandstone rocks with dripping water,
at 500 m.
Note. The type is a small specimen with pinnae
to 1.3 x 0.5 cm.
Central Sumatra (3 collec-
9. Mesophlebion endertii (C. CHR.) HOLTTUM,
Blumea 22 (1975) 228.— Dryopteris endertii C.
CHR. Dansk Bot. Ark. 9, 3 (1937) 60, pl. V, f.
7-10.— Type: ENDERT 4433, W. Kutai, Kemul,
Kalimantan (BO). — Fig. 6h.
Stipe 80cm long, glabrous above base; lamina
more than 100cm long; pinnae 12-14 pairs, lower
ones 9cm apart. Basal pinnae 43cm long with
stalks 4cm long and 15 pairs of free pinnules c.
1.5cm apart, then some separately adnate lobes,
distal part of pinnae deeply pinnatifid; basal pin-
nules very unequal (basiscopic 2 cm, acroscopic
4cm long); largest pinnules 5 x 1 cm, in basal half
lobed halfway to costule; veins to 25 pairs, lower
ones pinnate in the pinnule-lobes, thick and prom-
inent, distal ones simple; short hairs present on
lower surface of costae; upper pinnae normal for
genus Mesophlebion. Sori small; indusia very
small with a few short hairs.
Distr. Malesia: Borneo (E. Kalimantan: W.
Kutai) known from type only.
10. Mesophiebion motleyanum (HOOK.) HOLT-
TUM in Nayar & Kaur, Comp. to Bedd. (1974)
209; Blumea 22 (1975) 229.— Nephrodium mot-
leyanum HOOK. Syn. Fil. (1867) 266. — Dryopteris
motleyana (HOOK.) C. CHR. Ind. Fil. (1905) 278;
Gard. Bull. Str. Settl. 4 (1929) 385. — Dryopteris
crassifolia (BL.) O. KTZE var. motleyana
(HOOK.) v.A.v.R. Handb. (1908) 182. — Thelyp-
teris crassifolia (BL.) CHING var. motleyana
CHING, Bull. Fan Mem. Inst. Biol. Bot. 6 (1936)
286. — Thelypteris motleyana (HOOK.) HOLT-
TUM, Rev. FI. Mal. 2 (1955) 247, f. 140. — Type:
MOTLEY, Labuan (K).
Dryopteris vinosicarpa v.A.v.R. Bull. Jard.
Bot. Btzg III, 5 (1922) 198. — Thelypteris vinosi-
carpa (v.A.v.R.) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 255.—M. vinosicarpum
(v.A.v.R.) HOLTTUM, Blumea 22 (1975) 229. —
Type: cult. Hort. Bog. leg. BROOKS, origin Lebong
Tandai, Sumatra (BO; L).
382
FLORA MALESIANA
[ser. II, vol. 1°
Dryopteris motleyana var. dulitensis C. CHR.
Gard. Bull. Str. Settl. 4 (1929) 386.— Type:
MJOBERG, Mt Dulit, Sarawak (BM).
Nephrodium brachyodus sensu BAK. Syn. Fil.
(1867) 295, p.p. quoad pl. Malacc. tantum; BEDD.
Ferns Brit. India Suppl. (1876) 19, pl. 379; Handb.
(1883) 281, p.p. excl. typ.; RIDL. J. Mal. Br. R.
Asiat. Soc. (1926) 74, p.p.— Dryopteris brachy-
odus sensu v.A.v.R. Handb. (1908) 220, 819, p.p.
(all descriptions based partly on specimens from
Malaya).
Caudex short-creeping; stipe of sterile fronds
15-35cm long, of fertile fronds to 50cm,
glabrescent; basal scales few, 2-3mm_ long.
Lamina red when young, 20-50 cm or more long;
pinnae 6-8(-10) pairs, almost opposite; basal
pinnae slightly narrowed near base which is al-
most symmetric, with stalks 2-10 mm long; upper
pinnae sessile and often more asymmetric at base.
Largest sterile pinnae to 20X4cm, acuminate,
lobed about half-way to costa, lobes falcate; cos-
tules 6-10mm apart; veins to 10 on acroscopic
side, to 14 on basiscopic, basal veins from ad-
jacent groups almost parallel for some distance
below the sinus, basiscopic veins arising from the
costa far from their costules and sometimes anas-
tomosing with the veins next above them; lower
surface of rachis and costae short-hairy, many
small scales also present on costae, sparse short
hairs and rudimentary scales on costules. Fertile
fronds smaller than sterile; largest pinnae 12.5 x
2.5 cm, usually lobed a little more than half-way to
costa; orange-red glands often present on lower
surface of costules, minute scales consisting of
2-3 cells, the terminal one glandular, present on
young fronds on and between veins. Sori medial,
lower ones divergent, often somewhat elongate
along veins; indusia very small, often hidden by
sporangia, short-hairy.
Distr. Peninsular Thailand, in
Malaya, Sumatra, Borneo.
Ecol. In forest, at 0-1200 m.
Notes. The type of Dryopteris vinosicarpa
v.A.v.R. was a small plant but fertile. In 1975 I
associated with it some small plants from mid-
mountain forest in Malaya, restricting the name
M. motleyanum to larger lowland plants which
were thinner in texture. But plants from Sarawak,
now in cultivation at Kew, bridge the gap between
the two forms. Some specimens may represent
hybrids between this species and M. crassifolium,
but in general the two species appear to be dis-
tinct.
Malesia:
11. Mesophlebion
nov.
Frondes dimorphae; pinnae usque 18-jugatae,
inferiores stipitulatae, steriles usque 14X2.5 cm,
fertiles usque 11.5 x 1.7 cm, omnes dimidio costam
versus lobatae, lobis falcatis; venae infimae 2-
3mm infra sinum conniventes; indusia minuta,
falcatilobum HOLTTUM, sp.
glabra, sporangiis fere obtecta.— Type: JERMY
13694, Sarawak, G. Mulu (BM).
KEY TO THE VARIEDIES
1. Sterile pinna-lobes with obtuse to rounded
apices Jee oe a. var. falcatilobum
1. Sterile pinna-lobes with apiculate apices
b. var. apiculatum
a. var. faleatilobum
Caudex short-creeping; stipe 45cm _ long,
glabrous, scales short, not persistent, glabrous,
sometimes with red glands on margin near apex.
Lamina 45-50 cm long (sterile and fertile about
equal); pinnae c. 18 pairs, several lower pairs with
stalks, stalks of lowest 5-6mm, somewhat nar-
rowed towards base; aerophores swollen on living
fronds, collapsing on drying. Sterile pinnae to
14x 2.5cm; apex evenly attenuate, apical 2.5cm
entire; edges lobed half-way to costa or a little
less deeply, lobes falcate with obtuse to rounded
forward-pointing apices; costules 7mm _ apart;
veins to 11 on basiscopic side, 8 on acroscopic,
basal basiscopic vein arising from costa far from
its costule, basal veins from adjacent lobes con-
verging to touch the sinus-membrane 2-3mm
below the sinus; lower surface of rachis and cos-
tae bearing a variable number of short hairs, many
narrow scales on costae and a few glands; hairs on
upper surface of rachis and costae pale, thin,
0.3 mm long, rather sparse. Sori a little suprame-
dial, lower ones divergent; indusia very small,
glabrous; spores winged. Chromosomes: 2n=72
(T. G. WALKER, plant from type collection cult.
Kew).
Distr. Malesia: Borneo (Sarawak: Mt Mulu):
besides type, several other plants in cultivation
which are smaller than the wild plant but other-
wise similar.
b. var. apiculatum HOLTTUM, var. nov., a typo
speciei differt: pinnis sterilibus usque 9X 1.6cm,
1/3 costam versus lobatis, lobis valde
apiculatis. —Type: JERMyY 13611, G. Mulu,
Sarawak, in forest at 100-180 m (BM).
Pinnae c. 14 pairs, sterile to 9 X 1.6 cm, fertile to
6.5 1.0cm, lobed less than half-way to costa;
lobes of sterile pinnae very oblique with acute
tips; lobes of fertile pinnae crenulate; sori all
inframedial; lower surfaces of pinnae quite
glabrous.
Distr. Malesia: Borneo (Sarawak: Mt Mulu),
only known from the type.
12. Mesophlebion dulitense HOLTTUM, Blumea 22
(1975) 229. — Type: C. HOSE s.n. 1894, Mt Dulit,
Sarawak. (K; P).
Caudex not seen; stipe more than 75cm long,
glabrescent with dull reddish flush, basal scales
1981]
15mm or more long, thin, narrow. Lamina in-
complete, probably more than 75 cm long, texture
very firm. Largest pinnae probably 30-40 cm long,
6.5cm wide, with stalks 3-Scm long; base very
asymmetric with one basal lobe almost free and
separated from the rest; edges lobed to 3 mm from
costa; lobes slightly falcate, slightly narrowed
distally; costules 10mm apart, almost at right
angles to costa; veins to 25 pairs, rather thick and
prominent on both sides, basal basiscopic vein
arising near base of its costule in basal part of
pinna, remote from costule in distal part; lower
surface of rachis and costae sparsely short-hairy,
when young with many scales to 2mm long. Sori
inframedial distally on lobes, basal ones divergent;
indusia very small, caducous, bearing 1-2 short
hairs.
Distr. Malesia: Borneo. Known from type
locality, and G. Mulu, and Mt Kinabalu (2 collec-
tions).
Note. The Kinabalu plants, from 1500m, are
smaller than the type (pinnae to 20 5cm, with
stalk to 3.5 cm long) but agree in scales, hairs and
indusia.
13. Mesophlebion auriculiferum (v.A.v.R.)
HOLTTUM, Blumea 22 (1975) 231.— Dryopteris
auriculifera v.A.v.R. Bull. Jard. Bot. Btzg III, 5
(1922) 197.— Thelypteris auriculifera (v.A.v.R.)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941)
250. — Type: BUNNEMEIJER 6905, Lingga Is., Mt
Tanda (BO; L).
Caudex short; stipe 50-60 cm long, when young
densely hairy, basal scales rigid, to 7X 1mm.
Lamina 50cm long; pinnae 17-20 pairs, all stal-
ked, mostly not opposite; basal pinnae with stalks
to 7mm long, basal acroscopic lobe short and
often free, basal basiscopic lobe attached further
from rachis than acroscopic; texture firm, brittle
when dry. Largest pinnae 15 x 2 cm; base cuneate
with reduced lowest lobes, apex acuminate with
cauda to 2cm long: edges lobed to 2mm from
costa, lobes falcate; costules 44.5mm apart;
veins 10-11 pairs, basal acroscopic vein passing to
side of short sinus-membrane, basal basiscopic
vein to edge just above base of sinus; lower sur-
face of costae bearing many spreading pale hairs
to 1mm long, similar hairs more sparse on cos-
tules and veins, no scales seen; surface between
veins bearing many short slender erect hairs and
sessile glands. Sori inframedial; indusia thin, pale,
with many short capitate and acicular hairs, also a
few glands like those on surface between veins; a
small glandular cell at end of short hair on stalk of
sporangium; spores not seen.
Distr. Malesia: N.E. Sumatra (Lingga Is.).
Known from type collection only, in forest at
600 m.
Note. The pubescence of the lower surface of
this plant most nearly resembles that of M. teus-
cheri and M. arenicola; the latter are much
THELYPTERIDACEAE (Holttum)
383
smaller plants, growing apparently in a different
kind of habitat.
14. Mesophlebion beccarianum (CESATI) HOLT-
TUM, Blumea 22 (1975) 230.— Nephrodium bec-
carianum CESATI, Atti Acad. Napoli 7, n. 8 (1876)
23.— Dryopteris beccariana (CESATI) C. CHR.
Ind. Fil. (1905) 254; v.A.v.R. Handb. (1908) 185;
C. CHR. Gard. Bull. Str. Settl. 4 (1929) 384. —
Thelypteris beccariana (CESATI) REED, Phy-
tologia 17 (1968) 263.—Type: BECCARI s.n.,
Sarawak (FI).
Dryopteris pallescens BRAUSE, Bot. Jahrb. 56
(1920) 88.— Thelypteris pallescens (BRAUSE)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941)
253. — Mesophlebion pallescens (BRAUSE)
HOLTTUM, Blumea 22 (1975) 230.—Type:
LEDERMANN 9146, N.E. New Guinea, Sepik
Distr. (B; BM). — Fig. 6f-g.
Caudex short- to long-creeping, to 8mm
diameter; stipe to 50 cm or more long, glabrescent,
basal scales firm, to 7mm long, 1mm or more
wide at base with filiform apex. Lamina to 50cm
long, firm, usually drying greenish; pinnae 20 pairs
or more, often almost opposite, basal pinnae with
stalks 5-10mm long, basal pair of lobes often
much reduced, basal acroscopic lobe sometimes
almost free. Largest pinnae 12-18cm long, 1.6—
2.5 cm wide (sterile often wider than fertile); base
broadly cuneate to subtruncate; apex acuminate
with cauda 2-2.5cm long; edges lobed to 1-
1.5mm from costa, lobes oblique and slightly fal-
cate, tips rounded or broadly pointed; costules
3.5-4.5mm apart; veins 10-12(-15) pairs, basal
basiscopic vein arising from costa near base of its
costule, often ending just above base of sinus;
lower surface of rachis hairless or nearly so, of
costae often with sparse hairs 0.2 mm long, costal
scales 1-2 mm long, very narrow with a few mar-
ginal hairs, early caducous. Sori inframedial; in-
dusia large, firm, purplish, glabrous; spores
minutely papillose.
Distr. Malesia: Southern Malaya, Sarawak,
West and East New Guinea.
Ecol. In forest on sloping ground or ridges at
200-1200 m.
Note. The type of Dryopteris pallescens has
rather numerous short hairs on costae; in other
respects it differs little from the type of M. bec-
carianum. A specimen from Western New Guinea
is almost glabrous. Some specimens from Malaya
are intermediate between this and M. chlamy-
dophorum, having hairs on the lower surface of
costae and on indusia; Sarawak specimens seen
are very uniform, and the species seems locally
abundant there.
15. Mesophlebion rufescens HOLTTUM, Blumea 22
(1975) 230. — Type: BRASS 27957, Sudest Island
(K; BO, L, LAE).
Caudex ‘stout, horizontal’’ (BRASS); stipe 50—-
384
FLORA MALESIANA
[ser. II, vol. 1°
80cm long, lightly flushed with red, basal scales
rigid, dark, 4X 1mm, distal part of stipe short-
hairy. Lamina 42 cm long, texture firm, red-brown
when dry; pinnae 12 pairs, mostly opposite; basal
pinnae with stalks 10 mm long, basal pair of lobes
much reduced; fourth pair of pinnae truncate to
full width at base. Largest pinnae 12 x 2.8cm;
apex with cauda Icm long; edges lobed to 2.5-
3mm from costa, lobes slightly falcate with
obtuse to rounded tips; costules 5mm apart (fer-
tile), 6mm (sterile); veins 12-14 pairs, pale and
prominent both sides, basal basiscopic vein arising
from costa near base of its costule, usually ending
just above base of sinus; lower surface of rachis
and costae bearing copious slender pale hairs
0.3 mm long and many narrow scales, on costules
and veins sparse hairs, smaller narrow scales and
large red glands, short erect hairs on surface be-
tween veins. Sori medial, lower ones divergent;
indusia very small, glabrous, early caducous;
glands on stalk of sporangia very large.
Distr. Malesia: Philippines (Biliran: SULIT
PNH 20240; Luzon: Sierra Madre, JACOBS 7625)
and New Guinea (Sudest I. and Geelvink Bay,
SCHONIAN s.n. 1929 in B).
Ecol. In forest at 300 m.
16. Mesophlebion chlamydophorum (C. CHR.)
HOLTTUM, Blumea 22 (1975) 321.— Dryopteris
chlamydophora ROSENST. Med. Rijksherb. n. 31
(1917) 5, nom. nud.; C. CHR. Gard. Bull. Str. Settl.
4 (1929) 384.— Thelypteris chlamydophora (C.
CHR.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 6
(1936) 287; HOLTTUM, Rev. Fl. Malaya 2 (1955)
246, f. 139.— Mesoneuron chlamydophorum (C.
CHR.) CHING, Acta Phytotax. Sinica 8 (1963)
325. — Lectotype (HOLTTUM 1975): KORTHALS
s.n. Borneo (L, sheet n. 908, 342-57).
Lastrea nephrodioides BEDD. Ferns Brit. Ind.
(1866) t. 199, non MOorRE 1858; BEDD. Handb.
(1883) 238. — Type: PARISH, Burma (K, fragment
only).
Caudex short to rather long-creeping, diameter
commonly Smm; stipe to 60cm or more long,
basal scales 3-4 mm long. Fronds of mature plant
usually all fertile, with lamina to 80cm long, dry-
ing brown-olivaceous; pinnae 15-20 pairs, lower
pinnae with stalks 2-3mm or sometimes longer,
1-3 pairs of basal lobes reduced. Largest pinnae
commonly 15x2cm, to 202.5cm; apex acu-
minate; edges lobed to 2-3 mm from costa, lobes
falcate; costules 4-5.5 mm apart; veins commonly
10-12 pairs, basal pair of veins passing to either
side of sinus-membrane, basal basiscopic vein
arising nearer to its costule than in M. crassi-
folium (variable); lower surface of rachis, costae
and costules bearing very short hairs and narrow
scales, rudimentary scales (some consisting of 2-3
cells with glandular end-cell) on all parts of lower
surface, shrivelling and caducous with age. Sori
not very close to costules, basal ones divergent;
indusia fairly large, firm, bearing a variable num-
ber of very short acicular hairs; spores papillose.
Distr. S. Burma and Peninsular Thailand, in
Malesia: Malaya, Sumatra, Borneo, Celebes,
? New Guinea.
Ecol. In lowland forest, especially freshwater
swamp-forest.
Note. ROSENSTOCK published no description
and cited “‘Aspidium crassifollum METT. (non
BL.)”, but METTENIUS also published no des-
cription. The type selected is a specimen named
by ROSENSTOCK.
17. Mesophlebion crassifolium (BL.) HOLTTUM,
Blumea 22 (1975) 232.— Aspidium crassifolium
BL. En. Pl. Jav. (1828) 158; RAciB. FI. Btzg I
(1898) 169.— Lastrea crassifolia (BL.) MOORE,
Ind. Fil. (1858) 89; COPEL. Fern FI. Philip. (1960)
325. — Dryopteris crassifolia (BL.) O. KTZE, Rev.
Gen. Pl. 2 (1891) 812; v.A.v.R. Handb. (1908)
182, excl. var. motleyana; C. CHR. Gard. Bull. Str.
Settl. 4 (1929) 385; ibid. 7 (1934) 242, incl. var.
purpureo-lilacina; BACKER & POSTH. Varenfl.
Java (1939) 39.— Thelypteris crassifolia (BL.)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 6 (1936)
285; HOLTTUM, Rev. FI. Malaya 2 (1955) 246. —
Mesoneuron crassifolium (BL.) CHING, Acta
Phytotax. Sinica 8 (1963) 325.— Type: BLUME,
Java (L, sheet n. 908, 342-64).
Aspidium latum KUNZE ex METT. Farngatt. IV
(1858) 95. — Type: CUMING 266, Luzon (B; BM,G,
K).
Dryopteris divergens ROSENST. in Fedde Repert.
13 (1914) 218. — Thelypteris divergens (ROSENST.)
REED, Phytologia 17 (1968) 273.—Type: J.
WINKLER 36a, 1910, Sumatra (not seen).
Dryopteris subdimorpha COPEL. Univ. Cal. Publ.
Bot. 18 (1942) 220.—Lastrea subdimorpha
(COPEL.) COPEL. Gen. Fil. (1947) 140; Philip. J.
Sci. 78 (1951) 432, pl. 20.— Thelypteris sub-
dimorpha (COPEL.) REED, Phytologia 17 (1968)
317. — Type: BRASS 13666, Idenburg River, New
Guinea (MICH). — Fig. 1k.
Caudex creeping, diameter to 8mm or more;
stipe to 80cm or more long on fertile fronds, on
sterile shorter, basal scales rigid, to 10mm or
more long, with cordate base to 1.5mm wide.
Lamina to 70cm long, + dimorphous, texture
very firm; pinnae to 20 pairs, subopposite, rather
widely spaced; basal pinnae with stalks 3-15 mm
long, basal 1-3 pairs of lobes reduced. Largest
fertile pinnae 15<X3cm (sterile to 25 x 4.5 cm),
apex acuminate and sometimes caudate; edges
lobed to 3-4(—5)mm from costa, lobes falcate;
costules 4-6 mm apart (to 8mm in largest sterile
pinnae), at a wide angle to costa; veins 12-15
pairs, pale and prominent both sides, basal veins
from adjacent costules curved upwards so that
their distal parts are close together on either side
of the sinus-membrane, basal basiscopic vein
usually arising far from its costule; lower surface
1981]
THELYPTERIDACEAE (Holttum)
385
of rachis and costae copiously short-hairy, bearing
also narrow brown scales and sometimes glands
on costules. Sori inframedial, lower ones some-
what divergent; indusia usually rather large but
thin, shrivelling when dried, bearing short hairs,
less commonly quite small.
Distr. Malesia, except Java and Lesser Sunda
Islands.
Ecol. In forest on hill slopes at 700-1800 m.
Notes. This species is variable as regards
development of large sterile fronds and as
regards length of stalks of lower pinnae. Some
specimens appear to be intermediate between
typical M. crassifolium and M. motleyanum or
M. beccarianum; one from Malaya has spores
with perispore intermediate between the winged
type normal in this species and the papillose
spores of M. beccarianum. Most specimens from
Malaya and Borneo have rather large but thin
indusia, but a few have quite small indusia; spe-
cimens from the Philippines (Luzon, Negros,
Leyte) have all quite small indusia.
10. CYCLOSORUS
LINK, Hort. Reg. Bot. Berol. 2 (1833) 128; HOLTTUM, SEN & MITTRA,
Blumea 18 (1970) 200, 212; HoLtTrum, Blumea 19 (1971) 27; Allertonia 1
(1977) 181; of most other authors p.p. min. — Fig. lr, 7a—c.
Caudex long-creeping; scales proportionally broad, their hairs almost all
marginal; lowest pinnae not reduced; all pinnae lobed; basal veins always
anastomosing with a long excurrent vein to the sinus; lower surfaces
Fig. 7. Cyclosorus interruptus (WILLD.) H. ITo. a. Basal pinna, x 1.5; b. part of pinna showing scales,
hairs, glands and position of sori, X 8; c. sorus, showing glands among sporangia, X 32. — Ampelopteris
prolifera (RETZ.) COPEL. d. One primary pinna, x 1; e. two pinna-lobes showing venation, scales and
position of sori, x 3; f. first frond of a young plant proliferating from axil of primary pinna, x § (a-c T.
Loss 19, d-~f MATTHEW s.n.)
386 FLORA MALESIANA (ser. II, vol. 1°
variously hairy, broad flat scales always present on costae, rather large red
spherical glands on costules and veins; sori indusiate; no glands nor hairs
on body of sporangia, long hairs with terminal red glands on sporangium-
stalks; spores closely and irregularly spinulose.
Type species: Cyclosorus gongylodes (SCHKUHR) LINK.
Distr. Pantropic; probably 3 species.
Ecol. Always in freshwater swamps or edges of lakes, in full sun or light shade.
Cytol. All Indian and African records diploid, with n = 36; in the W. Indies and S. America some
plants tetraploid.
Taxon. In his monograph of Dryopteris (1913) CHRISTENSEN adopted the name Cyclosorus with
subgeneric status for a group of tropical American species, most of which are not closely related to C.
gongylodes (of which the type specimen came from Guiana). CHING extended the use of the name,
with generic rank, to cover species of the Old World some of which are related to the atypical
American species of CHRISTENSEN. COPELAND (Gen. Fil. 1947) followed CHING, distinguishing
Cyclosorus from Lastrea (= Thelypteris sensu CHING) solely by anastomosing/free venation, but this
does not result in a natural division. In my judgement, Cyclosorus (as here restricted) is most nearly
related to Ampelopteris (see HOLTTUM, SEN & MITTRA 1970) and Thelypteris s. str. (all have the
same aquatic habitat and a wide distribution), probably also to Mesophlebion, not closely to any other
group. As here restricted, Cyclosorus is a complex which can only be resolved by extensive cytotax-
onomic study of plants from the whole of its range. The fronds of W. Indian tetraploids are large, firm
and almost hairless on the lower surface; South African plants, originally named Polypodium tottum
THUNB., have a similar aspect. Seventeen binomials, with different types, are based on specimens
belonging to this complex.
1. Cyclosorus interruptus (WILLD.) H. ITO, Bot.
Mag. Tokyo 51 (1937) 714, nomen tantum;
HoLttuM, J. S. Afr. Bot. 40 (1974) 152. — Pteris
interrupta WILLD. Phytographia (1794) 13, t.10 f.
1; HoL_trumM, Amer. Fern J. 63 (1973) 81.—
Thelypteris interrupta (WILLD.) K. IwatTs. J.
Jap. Bot. 38 (1963) 314, nomen tantum; FOSBERG
& SACHET, Smiths. Contr. Bot. 8 (1972) 8.—
Type: KLEIN, S. India (B; Herb. Willd. 19, 770).
Aspidium obtusatum Sw. in Schrad. J. Bot.
1800, 2 (1801) 33; Syn. Fil. (1806) 248; WILLD.
Spec. Pl. 5 (1810) 241. — Type: THUNBERG, Java
(BM, fragment).
Nephrodium propinquum R. Br. Prodr. FI. N.
Holl. (1810) 148; BEDpb. Ferns S. India (1863) t.
89.— Type: BANKS, Queensland (BM; not seen).
Aspidium venulosum BL. En. Pl. Jav. (1828)
151. — Type: Java (not seen).
Hypopeltis marginifera BORY in Bél. Voy. Ind.
Or. Bot. 2 (1833) 69. — Type: BELANGER, Java (not
seen).
Hypopeltis propinquoides Bory, ibid. l.c.—
Type: BELANGER, Java (not seen).
Polypodium unitum LINN. Syst. Nat. ed. 10, 2
(1759) 1326 quoad Burm. Zeyl. tantum. — Aspi-
dium unitum sensu SCHKUHR, Krypt. Gew.
(1804) 34, t. 33b; sensu Sw. Syn. Fil. (1806) 47;
sensu BL. En. Pl. Jav. (1828) 150; sensu METT.
Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 230, var. hir-
sutum tantum.— Nephrodium unitum sensu R.
BR. Prodr. Fl. N. Holl. (1810) 148; sensu BEDD.
Handb. (1883) 268; sensu RAciIB. FI. Btzg I
(1898) 182.
Dryopteris gongylodes sensu v.A.v.R. Handb.
(1908) 212; sensu BACKER & POSTH. Varenfl.
Java (1939) 57.—C. gongylodes sensu CHING,
Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 186;
sensu HOLTTUM, Rev. FI. Malaya 2 (1955) 261,
f. 148; sensu COPEL. Fern FI. Philip. (1960) 360.
(All doubtfully based on Aspidium gongylodes
SCHKUHR, Krypt. Gew. (1809) 193, t. 33c).—
Fig. Ir, 7a—c.
Stipe to c. 45cm long, lightly flushed with red
above dark base, glabrescent. Lamina commonly
30-50 cm long; pinnae 20-25 pairs; several pairs
lower pinnae slightly narrowed towards an
asymmetric base. Largest pinnae commonly 10-
15cm long, 1.0-1.8cm wide, in shaded places
sometimes larger, texture firm; apex short-acu-
minate; edges lobed less than half-way to costa,
lobes oblique, broadly rounded with distinct car-
tilaginous edges and a small apiculus; costules
commonly 3.5-4.5mm apart, at less than 60° to
costa; veins 8-10 pairs or more, very oblique
except basal pair, 1-1; pairs anastomosing and
usually 1 pair touching sides of the short sinus-
membrane; lower surface of rachis, costae, cos-
tules and veins bearing a variable number of
short erect pale acicular and sometimes capitate
hairs, a few hairs also sometimes on surface be-
tween veins, a variable number of large red
glands scattered on costules and veins, broad thin
flat fringed scales on costae; upper surface of
rachis covered with short hairs, few short hairs
on costae and edges. Sori supramedial except on
distal veins, absent from basal pair of veins; in-
dusia thin, bearing short acicular hairs and some-
times capitate hairs.
Distr. Tropics and subtropics.
Notes. It seems probable that Aspidium
1981]
THELYPTERIDACEAE (Holttum)
387
gongylodes SCHKUHR was a tetraploid; his spe-
cimens at B and his illustration agree with known
tetraploids. But some other tropical American
specimens are not clearly distinct from the
plants of Asia and Africa known to be diploid.
The morphological distinctions of the tetraploid
need to be established, and the type of Pteris
polypodioides Porr. (1804), an earlier name,
needs to be examined; also P. tottum THUNB.
(1800) from South Africa must be taken into
consideration. In any case, the name gongylodes
is incorrect for the Malesian plants.
CHING, who identified it with the species here
named Amphineuron terminans (HOOK.) HOLT-
TUM, and his identification was copied by many
subsequent authors. In 1801 SWARTZ published
the name Aspidium obtusatum with Pteris inter-
rupta WILLD. as a synonym; this identification
was accepted by WILLDENOw in 1810.
The nature and distribution of hairs on the
lower surfaces of pinnae of Malesian plants
varies greatly. Specimens with many short capi-
tate hairs have been collected in Sabah and
Sarawak, also in Mainland Asia from Peninsular
WILLDENOW’s type was misconstrued by Thailand to Assam and also Hainan.
11. AMPELOPTERIS
KUNZE, Bot. Zeit. 6 (1848) 114; COPEL. Gen. Fil. (1947) 143; CHING, Acta
Phytotax. Sinica 8 (1963) 330; HOLTTUM, SEN & MITTRA, Blumea 18
(1970) 196, 214; HoLTTUM, Blumea 19 (1971) 25.— Meniscium sect. Am-
pelopteris K. IWATS. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 39.—
Thelypteris subg. Meniscium sect. Ampelopteris REED, Phytologia 17
(1968) 255. — Fig. 7d-f.
Caudex creeping; fronds of indefinite apical growth, bearing many buds
at the bases of primary pinnae, the buds developing into new plants;
forked or branched unicellular acicular hairs present on rachis; sori exin-
dusiate; stalks of sporangia bearing hairs each terminating in a large
globular gland; spores similar to those of Cyclosorus.
Type species:
COPEL.
Distr. Monotypic. Old World tropics.
Ampelopteris elegans
KUNZE=A. prolifera (RETZ.)
Cytol. Chromosome number 36 (India & Ceylon), several records (all diploid).
Notes. This genus resembles Cyclosorus in sporangia, spores and in ovate scales on lower surface
of costae; the two genera also agree in creeping rhizome and aquatic habitat. Ampelopteris is the only
Old World member of the family which has branched acicular hairs; in this character it resembles
Goniopteris, a genus of many species in the New World.
1. Ampelopteris prolifera (RETZ.) COPEL. Gen.
Fil. (1947) 144; HoL_LtTTuM, Rev. FI. Malaya 2
(1955) 299; COPEL. Fern FI. Philip. (1960) 377. —
Hemionitis prolifera RETZ. Obs. Bot. 6 (1791)
38. — Meniscium proliferum (RETZ.) Sw. Syn. Fil.
(1806) 19, 207; HooK. 2nd Cent. Ferns (1861)
t. 15, f. 1-3, tantum.—Goniopteris prolifera
(RETZ.) PRESL, Tent. Pterid. (1836) 183; BEDD.
Ferns S. India (1864) t. 172; Handb. (1883) 296, f.
153.— Polypodium luxurians KUNZE, Linnaea
23 (1850) 28, nom. nov. — Phegopteris luxurians
(KUNZE) METT. Farngatt. IV (1858) 25. — Phe-
gopteris prolifera (RETZ.) KUHN in v. Decken
Reise (1879) 44; v.A.v.R. Handb. (1908) 504. —
Dryopteris prolifera (RETZ.) C. CHR. Ind. Fil.
(1905) 286; BACKER & POSTH. Varenfl. Java
(1939) 55. — Cyclosorus proliferus (RETZ.) TARD.
& C. CHR. Notul. Syst. 14 (1952) 346. — Thelyp-
teris prolifera (RETZ.) REED, Phytologia 17
(1968) 306. — Type: KOENIG, S. India (GOET).
A. elegans KUNZE, Bot. Zeit. 6 (1848) 114. —
Type: ZOLLINGER 2360, Java (isotypes at BM,
G, P, W).
Goniopteris meniscioides FEE, Gen. Fil (1852)
253.— Type: CUMING 168 (P;FI-W, G, K,
LE). — Fig. 7d-f.
Primary pinnae to 20 X 2 cm, often with a short
stalk; base truncate to subcordate; apex rather
evenly attenuate; edges lobed to a depth of
2mm; costules 3-4 mm apart; veins 10-12 pairs,
5-6 pairs anastomosing to form a Zig-zag excur-
rent vein; lower surfaces hairless, costae bearing
ovate ciliate scales. Sori exindusiate, somewhat
elongate, on distal parts of veins. Fronds on
plants formed by proliferation from buds much
smaller, usually with pinna-like terminal lamina;
pinnae sometimes fertile from a size of 3.5
0.8 cm.
388 FLORA MALESIANA [ser. II, vol. 1°
f NS Sl See *
Bt pie enor nie ae Rae pHi septa ae
SO a leer a
oe
eu
Fig. 8. Chingia ferox (BL.) HOLTTUM. a. Rachis with terete scales and base of a pinna bearing
coalescent sori, x 4; b. two old sori, x 12.— C. sakayensis (ZEILLER) HOLTTUM. c. Two pinna-lobes,
x2; d. part of a pinna showing glands and position of sori, x4; e. one sorus, x 10.—C. pricei
HOLTTuom. f. Two pinna-lobes, showing venation and position of sori, <4; g. one sorus, X 16.—C.
christii (COPEL.) HOLTTUM. h. Two pinna-lobes, x4.—C. clavipilosa HOLTTUM var. javanica
HOLTTUM. i. Rachis with flat scales and base of a pinna, x 4 (a—b MOUSSET s.n., c—e DAY s.n., f-g PRICE
512, h ELMER 10187, i CLEMENS 30514).
1981] THELYPTERIDACEAE (Holttum) 389
Distr. West Africa to N.E. Australia (to 30°S) Notes No. distinct varieties have been
and New Caledonia; widely in tropical mainland observed. The unnumbered figure of a complete
Asia; throughout Malesia but not often collected. small plant on HOOKER’s plate (1861) represents
Ecol. On banks of rivers and ditches, some- Pronephrium hosei (BAK.) HOLTTUM.
times forming thickets.
12. CHINGIA
HOLTTuM, Blumea 19 (1971) 31; Kalikasan 3 (1974) 13-28. — Fig. 8.
Caudex massive, erect, sometimes forming a trunk 30-100 cm tall. Fronds
usually rather rigid in texture, commonly 150-250 cm long including stipe,
axes usually drying red-brown; basal pinnae not reduced; aerophores not
enlarged; at least the base of the stipe, in many cases the whole of it and
part or all of the rachis persistently scaly, scales at base of stipe narrow,
often rigid, setiferous, in some species grading into quite terete spine-like
scales above base of stipe. Pinnae crenate to deeply lobed, in most species
to 30cm or more long; veins numerous, all except the basal pair very
oblique, basal 1-3 pairs anastomosing (except in C. pricei), the next 1-3 or
more pairs passing to sides of a long sinus-membrane; some acicular hairs
often present on lower surfaces, sometimes only on young plants or those
growing in exposed places; capitate glandular hairs or small sessile glands
abundant in some species on lower or both surfaces of pinnae. Sori near
costules except the lowest 1-2 pairs which may be divergent; indusia small
or lacking; young sporangia in some species bearing small sessile glands or
capitate hairs, in one species rarely setae; spores dark, covered with minute
papillae or wings.
Type species: Chingia ferox (BL.) HOLTTUM.
Distr. Malesia and the Pacific (to Tahiti); in Mainland Asia only to Peninsular Thailand; in all 20
species.
Ecol. Usually on hill slopes in moderately exposed places, at 300-2000 m, sometimes locally abundant
(used for thatching in Mindoro).
Cytol. Chromosome number 36 (C. atrospinosa, C. urens, both diploid).
Notes. This very natural group appears to be closely related to Plesioneuron but not to any other
genus of the present arrangement. CHING and COPELAND placed the species known to them in
Cyclosorus without distinguishing them as a natural group. The nature and distribution of scales are
distinctive in the various species, and some early collections do not show these characters adequately.
New information has recently been added by the collections of M. G. PRICE in the Philippines where
seven species occur. Chemical investigation might yield interesting results; two species are recorded as
malodorous, and one (C. urens) as causing a mild burning sensation in contact with human skin.
Plants of Glaphyropteridopsis CHING (type species G. erubescens (HOOK.) CHING, distributed from
N.E. India to S. China) are similar to Chingia in frond-form, but the veins are always free, aerophores
swollen, and the spores and scales are very different; the two are probably not very nearly related.
KEY TO THE SPECIES
1. Abaxial surface of rachis bearing slender terete spine-like scales.
2. \Pinnaé crenate itolaidepthiofidimm 4 fos). Son We Pa Ae EN), Clee eee racntidens
2. Pinnae much more deeply lobed.
3. Pinnae 8-12 pairs, 6cm or more apart cfu... SORA le ey eC eatrospinosa
3. Pinnae to 20 or more pairs, closer together.
4. Sori all elongate; veins not prominent on lower surface . . . . . . 3. C. muricata
4. At most basal sori elongate; veins rather thick and prominent beneath.
390
5. Pinnae not over 3 cm wide, lobed 1/4-2/5; basal sori only divergent but not elongate
lobed 2/5-3/5; basal 2-
5. Pinnae to 3.5cm or more wide,
elongate
FLORA MALESIANA
(ser. Il, vwoleate
4. C. ferox
3 pairs of sori divergent and lowest
.5. C. imponens
. Abaxial surface of rachis i in “most species lacking scales; scales, if present in this position, distinctly
flat.
6. Basal veins not meeting but passing to sides of the sinus-membrane which is decurrent below their
junction with it
. 6. C. pricei
6. Basal veins anastomosing o or at least meeting below base of sinus- Semiemibranc which i is not decurrent.
7. Stipe scaly throughout; at least lower part of rachis scaly on abaxial side.
8. Pinnae to 3cm wide, thin, drying light greenish; small indusia present
8. Pinae not over 2.5 cm wide, firm, drying brown-olivaceous; no indusia
7. C. sakayensis
8. C. clavipilosa
9. Capitate hairs abundant on lower surface of costae and costules; capitate hairs and sessile glands
between veins
9. Glands mainly sessile, lacking between v veins EL ye:
7. Stipe scaly near base only; rachis not scaly on abaxial surface.
10. Rachis bearing slender terete scales on adaxial surface
. 8a. C. clavipilosa var. clavipilosa
8b. C. clavipilosa var. javanica
. 9. C. bewaniensis
10. Rachis lacking scales on adaxial surface, or such scales flat.
11. Pinnae crenate to a depth of 1.5—
12. Lower surfaces glabrous.
3 mm; no indusia.
12. Lower surfaces covered with slender erect acicular hairs
. 10. C. lorzingii
13. Pinnae thin; veins not prominent beneath; sori close to costules except lowest
11. C. sambasensis
13. Pinnae firm; veins prominent beneath; several pairs of lower sori gradually divergent
12. C. christii
Pinnae lobed 1/3 towards costa or more deeply; indusia present in most cases.
ais Lower surface bearing sessile glands
14. Lower surface bearing acicular and/or capitate hairs.
13. C. paucipaleata
15. Lower surface densely acicular-hairy throughout, tai capitate hairs; no capitate hairs on
sporangia
14. C. horridipes
15. Lower surface bearing short capitate hairs at least | on “ costules and veins; capitate hairs
(rarely setae) present on sporangia.
16. Pinnae thick and rigid; veins prominent on lower surface, grooved on upper surface
15. C. perrigida
16. Pinnae thin; veins not prominent on lower surface nor grooved on upper surface.
17. Basal pair of veins anastomosing to form a slender excurrent vein which is joined by the
second acroscopic vein before entering base of sinus-membrane
16. C. supraspinigera
17. Basal pair of veins meeting, or not quite meeting, at base of sinus-membrane.
18. Pinnae lobed less than half-way to costa; sori indusiate
18. Pinnae lobed more than half-way to costa; sori exindusiate
1. Chingia acutidens HOLTTUM, Kalikasan 3
(1974) 17.— Type: ALSTON 15708, Mt Manim-
porok, Soputan Mts, Minahassa, Celebes (BM).
Stipe more than 85cm long, rather copiously
black-muricate throughout (bases of former
scales), abaxial surface of rachis similarly but less
prominently muricate. Lamina 170 cm long, apex
pinna-like; pinnae nearly 40 pairs; basal pinnae
narrowed to base on basiscopic side, acroscopic
base truncate and slightly auricled. Suprabasal
pinnae to 42 xX 2.4cm; base broadly cuneate; apex
acuminate with slender cauda to 6cm long; edges
crenate-serrate to a depth of 1mm; costules 4.5—
5mm apart, at more than 60°; veins to 10 pairs, 2
pairs anastomosing, 3 or more pairs passing to
sides of sinus-membrane; lower surface of pinnae
glabrous apart from sparse hairs on sinus-mem-
brane and edge; upper surface with short hairs on
costae only. Sori rather large, distinctly free from
costules, at least basal ones elongate; no indusia;
no glands on sporangia.
17. C. urens
. 18. C. tenerior
Distr. Malesia:
HUMUS 2566).
N. Celebes (type and PostT-
2. Chingia atrospinosa (C. CHR.) HOLTTUM, Kal-
ikasan 3 (1974) 19.— Dryopteris atrospinosa C.
CHR. Bot. Jahrb. 66 (1933) 43.— Thelypteris
atrospinosa (C. CHR.) REED, Phytologia 17 (1968)
261.—Type: KJELLBERG 3652, Rante Lemo,
Celebes (S-PA; BO).
Stipe to 75cm long, stramineous; basal scales
10mm long, hardly 1mm _ wide, dark brown,
glossy, with short spreading hairs; above base
throughout bearing many short black spines (bases
of former scales); lower surface of rachis bearing
also scattered spines or terete dark scales. Lamina
to 75 cm long, apex almost pinna-like; pinnae 8-12
pairs, widely spaced, rigid; basal pinnae narrowed
gradually to their bases, with stalks 2-3 mm long.
Suprabasal pinnae 25-30cm long, 2.5-3.0cm
wide; base broadly cuneate; apex acuminate with
sharp-toothed cauda 3cm or more long; edges
1981]
THELYPTERIDACEAE (Holttum)
391
lobed 1/3 towards costa, lobes falcate, narrowed
to acute tip; costules 5mm apart; veins 10-11
pairs, slightly prominent both sides, 2 pairs anas-
tomosing, 2 pairs to sides of sinus-membrane;
lower surface of costae and costules bearing stiff
spreading slender hairs 0.5-1 mm long, scattered
hairs on and between veins; upper surface hairy
on costa and edges only. Sori near costules,
exindusiate, spreading a little along veins; no
glands on sporangia.
Distr. Malesia: Celebes (N to SW); Sabah.
Ecol. At 1000-2000 m, in open places, in some
cases near streams.
3. Chingia muricata (BRAUSE) HOLTTUM, Kal-
ikasan 3 (1974) 20.— Dryopteris muricata
BRAUSE, Bot. Jahrb. 56 (1920) 106, excl. var.
marginata et var. obscura.—Cyclosorus im-
ponens sensu COPEL. Philip. J. Sci. 78 (1951) 457,
p.p.— Thelypteris muricata (BRAUSE) REED,
Phytologia 17 (1968) 295. — Type: LEDERMANN
12720, Sepik Distr., New Guinea (B).
Stipe 45cm long, densely covered with spines
1-2 mm long and scales; scales to 24 x 1 mm, hair-
pointed, bearing short setae, grading in distal part
of stipe to terete hairy spines 8 mm long. Lamina
c. 100cm long; pinnae closely spaced; basal pin-
nae somewhat narrowed to their bases and short-
stalked; texture coriaceous. Largest pinnae of
type 35x3.3cm; base subtruncate, apex acu-
minate, not caudate; edges lobed 1/3-2/5 towards
costa, lobes close, falcate; costules to 5.5mm
apart; veins 10-12 pairs, grooved on upper sur-
face, hardly evident on lower when dry, 1 pairs
anastomosing, 2 pairs to sides of sinus-membrane;
lower surface of rachis with slender terete bristles
to 8 mm long, of pinnae glabrous apart from a few
hairs on sinus-membrane and edges. Sori all
somewhat elongate (elliptic in outline), distal ones
near costule, lower ones a little divergent, exin-
dusiate; no glands on sporangia.
Distr. Malesia: New Guinea, from Idenburg
River to Morobe District.
Ecol. Type from wet rocky place; others from
secondary forest, at 850-1500 m.
4. Chingia ferox (BL.) HOLTTUM, Blumea 19
(1971) 31; Kalikasan 3 (1974) 21.— Aspidium
ferox BL. En. Pl. Jav. (1828) 153. — Nephrodium
ferox (BL.) Moore, Ind. Fil. (1858) 91; Hook.
Spec. Fil. 4 (1862) 77, excl. Wallich, Kamaon;
RACIB. FI. Btzg I (1898) 192. — Phegopteris ferox
(BL.) METT. Ann. Mus. Bot. Lugd.-Bat. 1 (1864)
224. — Dryopteris ferox (BL.) O. KTZE, Rev. Gen.
Pl. 2 (1891) 812; v.A.v.R. Handb. (1908) 221, 819,
excl. var. calvescens; BACKER & POSTH. Varenfl.
Java (1939) 62.— Cyclosorus ferox (BL.) CHING,
Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 167;
COPEL. Fern FI. Philip. (1960) 347. — Thelypteris
ferox (BL.) TAGAWA & K. IwaTs. Acta Phy-
totax. Geobot. 23 (1968) 54.—Type: BLUME,
West Java (L, sheet n. 908, 331-305).
Polypodium scabrum ROXxB. Calc. J. Nat. Hist.
4 (1844) 491.— Type: WALLICH Cat. 2225, Am-
boina (isotype K; ill. as Nephrodium ferox by
BEDD. Ferns Br. India t. 129).
Goniopteris barbata FEE, Gen. Fil. (1852) 252. —
Phegopteris barbata (FEE) METT. Farngatt. IV
(1858) 24. — Polypodium barbatum (FEE) HOOK.
Spec. Fil. 5 (1864) 10.—Type: CUMING 172,
Luzon (orig.?; isotypes BM, K, L, SING). — Fig.
8a-b.
Caudex to 20cm or more tall; stipe commonly
100 cm long, largest basal scales 25 x 1.5 mm, thin,
glossy, setiferous, medium brown; scales on upper
part of stipe and on rachis terete or nearly so,
hairless, dark, 5-10 mm long, often broken leaving
warts or short spines. Lamina 100-200 cm long,
texture varying with altitude and exposure; basal
pinnae slightly narrowed at their bases which are
slightly auricled on acroscopic side. Largest pin-
nae 18—40(-50)cm long, 1.6-3.0cm wide, nar-
rowly acuminate; edges lobed 1/4-2/5 towards
costa; lobes with acute falcate tips; costules 3.5—
4.5mm apart; veins to 16 pairs, not very thick,
variably prominent on lower surface, grooved or
flat on upper, | pair anastomosing, 3-4 pairs pass-
ing to sides of sinus-membrane; lower surface of
costae, costules, sinus-membranes and edges
bearing + abundant erect slender acicular hairs,
sometimes also capitate hairs (which may be
abundant on sterile pinnae) on costules and veins.
Sori exindusiate, close to costules except basal
basiscopic one, not elongate; sporangia of Java
specimens lacking glands, small red glands present
on some specimens from Moluccas and Philip-
pines.
Distr. Malesia: Java, Lesser Sunda Islands,
Borneo, Celebes, Philippines, Moluccas, New
Guinea.
Ecol. Usually in secondary growth, on hill-
slopes; 500-2000 m.
Notes. The specimen n. 2225 from Herb. Wal-
lich among HOOKER’s specimens at Kew is
labelled Kamaon in HOOKER’s hand, though
ROXBURGH gave Amboina as its origin; another
was figured by BEDDOME; both have small red
glands on the sporangia like those from the
Moluccas. No locality is indicated in Wallich’s
Catalogue. Philippine specimens seen are more
shallowly lobed than those from Java.
5. Chingia imponens (CES.) HOLTTUM, Kalikasan
3 (1974) 21. — Polypodium imponens CEs. Rendic.
Acad. Napoli 16 (1877) 27, 29.— Dryopteris im-
ponens (CES.) C. CHR. Ind. Fil. (1905) 271; Dansk
Bot. Ark. 9, 3 (1937) 50, pl. V, f. 5. — Phegopteris
imponens (CES.) v.A.v.R. Handb. (1908) 506. —
Cyclosorus imponens (CES.) COPEL. Gen. Fil.
(1947) 142; Philip. J. Sci. 78 (1951) 437. — Thelyp-
teris imponens (CES.) REED, Phytologia 17 (1968)
284.— Type: BECCARI, Arfak Mts, W. New
392
FLORA MALESIANA
(ser. II, vol. 1°
Guinea (FI, Herb. Becc. 12668).
Dryopteris armata ROSENST. Hedwigia 56 (1915)
351. — Phegopteris armata (ROSENST.) v.A.v.R.
Handb. Suppl. (1917) 318. — Type: BAMLER I11,
Sattelberg, N.E. New Guinea (B; UC).
Dryopteris muricata var. marginata BRAUSE,
Bot. Jahrb. 56 (1920) 107. — Type: LEDERMANN
11937, Sepik Distr., New Guinea (B).
D. muricata var. obscura BRAUSE, l.c. 108. —
Type: LEDERMANN 12022, same locality.
Stipe 100 cm or more long; basal scales 15-20 x
1.5mm, medium dull brown, rest of stipe and
rachis bearing many slender terete dark scales to
10mm long, often broken and reduced to short
spines which bear a few stiff hairs. Lamina
150cm or more long; pinnae close, many pairs;
lowest pinnae gradually narrowed in basal 8-
10cm, with stalks 2mm long. Largest pinnae
commonly 45 x 3.5-4.0 cm; base subtruncate; apex
gradually attenuate; edges lobed 2/5-3/5 towards
costa, lobes falcate distally; costules 6-6.5 mm
apart; veins 12-15 pairs, slender, dark beneath and
slightly prominent, grooved on upper surface
when dry, | pair anastomosing, 2-3 pairs passing
to sinus-membrane; lower surface of costae, cos-
tules, veins and sinus-membranes bearing sparse
short pale acicular hairs; upper surface of costae
bearing slender pale to brownish hairs 0.5 mm or
more long, few hairs on costules. Sori near cos-
tules except basal 2 pairs which are divergent and
slightly elongate; no indusia; no glands on
sporangia.
Distr. Malesia: Eastern New Guinea, at al-
titudes to 1800 m, many collections.
Note. This species is very near C. ferox but has
less rigid and more deeply-lobed pinnae and stiff
hairs on the stumps of broken terete scales on the
rachis.
6. Chingia pricei HOLTTUM, Kalikasan 3 (1974)
22.— Type: M. G. PRICE 512, Mt Santo Tomas,
Benguet Prov. Luzon, exposed place near summit
(PNH; K). — Fig. 8f-g.
Caudex 10cm diameter (collector); stipe 45-
60cm long, basal half covered with glossy light
brown scales to 101mm, thin but firm,
thickened at base, bearing sparse stiff hairs; distal
part of stipe, and rachis, bearing only smaller
scales (which are not terete) on edges of groove of
adaxial surface, abaxial surface smooth. Lamina
to 60cm long; pinnae more than 20 pairs; lowest
pinnae hardly narrowed near base, basal acros-
copic lobe elongate; texture firm. Largest fertile
pinnae 13 cm long, 2.1 cm wide above base (sterile
to 18 x 2.3 cm); base truncate, usually dilated both
sides to a total width of 3cm; apex acuminate;
edges lobed half-way to costa or a little more
deeply, lobes slightly falcate; costules 44.5 mm
apart; veins 10 pairs, basal pair very oblique, not
joining but passing to sides of the sinus-membrane
which is decurrent between them, next 13-2 pairs
also passing to sides of the membrane; lower
surface of rachis, costae, costules, veins and be-
tween veins bearing copious short capitate hairs,
acicular hairs present on sinus-membranes and
edges; upper surface of costae covered with many
thick brownish hairs less than 0.5 mm long, sparse
acicular hairs on costules, scattered short capitate
hairs on and between veins. Distal sori close to
costules, lower ones a little divergent; indusia
fairly large, thin, shrivelling, with capitate and
acicular hairs; capitate hairs on sporangia.
Distr. Malesia: Philippines (Luzon), known
from type only.
7. Chingia sakayensis (ZEILLER) HOLTTUM,
comb. nov.— Nephrodium sakayense ZEILLER,
Bull. Soc. Bot. France 32 (1885) 75; BEDD.
Handb. Suppl. (1892) 78. — Thelypteris sakayensis
(ZEILLER) REED, Phytologia 17 (1968) 311.—
Sphaerostephanos sakayensis (ZEILLER) HOLT-
TUM in Nayar & Kaur, Comp. to Bedd. (1974)
209. — Type: DE MORGAN s.n. 6 Aug. 1884, near
Gunong Riam, Perak, 750 m (P; K).
C. pseudoferox HOLTTUM, Kalikasan 3 (1974)
24. — Type: MATTHEW s.n. 6 Feb. 1908, G. Hijau,
Perak (K).
Cyclosorus ferox sensu. HOLTTUM, Rev. FI.
Malaya 2 (1955) 265, f. 151, quoad pl. Malay.
tantum. — Fig. 8c-e.
Stipe to 200 cm long, densely scaly throughout;
scales to 20x 1.5mm, rigid, flat with thickened
persistent bases, dark brown, bearing short stiff
marginal and superficial setae; abaxial surface of
basal part of rachis bearing very narrow and
shorter similar scales. Lamina to 200cm long;
pinnae many pairs, texture thinner than in C.
ferox, drying light greenish; basal pinnae gradually
much narrowed towards their bases which may be
only 5mm wide. Largest pinnae to 40 x 3 cm (type
of C. pseudoferox) but in young plants sometimes
fertile at 19 x 1.6 cm (type of N. sakayense); apex
rather evenly narrow-attenuate; edges lobed about
half-way to costa, lobes distally falcate and nar-
rowed to an obtuse tip; costules 5 mm apart; veins
10-15 pairs, slender and slightly prominent on
lower surface, 1-2 pairs anastomosing, next 2-3
pairs to sides of sinus-membrane; lower surface of
costae, costules, veins and surface between veins
bearing + abundant small sessile yellowish glands,
short acicular hairs also sometimes present; upper
surface hairy on costae and costules only, no
glands. Sori close to costules except basal ones
which are often a little elongate; indusia very
small, bearing a few small glands; sporangia with
small glands.
Distr. Peninsular Thailand, in
Malaya, Sumatra, Sarawak, W. Java.
Ecol. In forest, often near streams, at 150-
1200 m.
Note. The type of N. sakayense at Paris con-
sists of the apical part of a frond; at Kew are 2
Malesia:
1981]
THELYPTERIDACEAE (Holttum)
393
detached pinnae. The aspect of these specimens is
so different from that of fully grown plants that
formerly I failed to recognize their identity, which
is confirmed by recent gatherings of other small
fertile plants. The single specimen from W. Java is
from G. Pulosari at 600 m (ABDELBERT 492).
8. Chingia clavipilosa HOLTTUM, Kalikasan 3
(1974) 23.—Type: HOLTTUM 44, Mt Kinabalu
2300 m, Sabah (K).
a. var. clavipilosa
Stipe densely scaly throughout, scales all flat,
basal ones 15 x | mm; scales on abaxial surface of
rachis more sparse but present throughout, very
narrow but distinctly flat, not terete, at their bases.
Basal pinnae narrowed towards base which is
1.5cm wide. Largest pinnae 28X2.5cm, very
firm, lobed 1/3 towards costa; costules 4.5mm
apart; veins to 12 pairs, slender and slightly prom-
inent on lower surface, not on upper, 1-13 pairs
anastomosing, 2-3 pairs passing to sides of sinus-
membrane; lower surface of costae (at least distally)
and costules bearing + abundant slender spreading
hairs to 1.5 mm long; on costae, costules and veins
abundant small capitate hairs, on surface between
veins short capitate hairs and very small sessile
glands; upper surface of costae bearing thick
brownish hairs 0.7mm long, rest glabrous. Sori
close to costules except on basal veins; no in-
dusia; sporangia bearing capitate hairs.
Distr. Malesia: N. Borneo (Kinabalu) and
Philippines (Luzon, Mountain Province).
Note. The description is prepared from
Kinabalu specimens; those from N. Luzon are
much smaller (pinnae to 10 1cm) with more
copious acicular hairs on lower surface.
b. var. javanica HOLTTUM, var. nov., ab var.
clavipilosa differt: costis costulis venisque subtus
glandulis sessilibus vel subsessilibus praeditis,
glandulis inter venas nullis; sporangiis immaturis
glandulis pallidis minutis praeditis.— Type: 1221
Herb. Bog., Tjibeureum, G. Gedeh, Java (BO). —
Fig. 8i.
Distr. Malesia: West Java, many specimens,
1800-2000 m; Sarawak, G. Mulu.
Note. These specimens were formerly included
in C. ferox but differ in the distinctly flat scales on
the abaxial surface of the rachis and in the small
glands on voung sporangia.
9. Chingia bewaniensis HOLTTUM, sp. nov.
Stipes basin versus tantum paleatus, paleis
omnibus applanatis; pagina adaxialis rhachidis
paleis teretibus praedita, pagina abaxialis minute
verrucosa; cetera C. feroci similis.— Type:
JERMY 8170, Bewani Mts, N.E. New Guinea,
300 m (BM).
Stipe 70cm long, densely scaly at base only,
above base minutely verrucose; scales 15mm
long, 0.5—1.0 mm wide, thin, medium brown, bear-
ing sparse setae. Lamina firm, drying light
olivaceous, 120cm long; several pairs lower pin-
nae gradually narrowed towards their bases which
are 1.1cm wide with auricle to 9mm long on
acroscopic side, stalk 2mm. Largest pinnae 45 x
2.5cm; apex narrowly acuminate; edges lobed 1/3
towards costa, lobes falcate subacute; costules
5mm apart; veins to 12 pairs on acroscopic side,
14 on basiscopic, | pair anastomosing, 2-3 pairs
passing to sides of sinus-membrane; lower surface
of rachis pale, glabrous, with minute warts as
upper part of stipe, of costae glabrous; upper
surface of rachis, within the groove, bearing
many slender terete scales, costae copiously
antrorse-hairy, rest glabrous. Sori near costules,
basal ones only slightly divergent, exindusiate;
sporangia bearing small red glands.
Distr. Malesia: Papua New Guinea;
known from the type.
only
10. Chingia lorzingii HOLTTUM, sp. nov.
Stipes basin versus tantum paleatus; pagina
abaxialis rhachidis pilis acicularibus patentibus
0.7-0.8 mm longis dense vestita; pinnae tenues,
maximae 20 X 1.8 cm, vix 1/4 costam versus loba-
tae, subtus omnino pilis acicularibus erectis ves-
titae; sori exindusiati; sporangia glandulis non
praedita. — Type: LORZING 16188, Karoplateau,
Dolok-baros, Sumatra, 1750-1950 m (BO).
Stipe 80cm long; basal scales few, flat; upper
part of stipe bearing short pale acicular hairs and
very few scales; abaxial surface of rachis, espe-
cially distally, densely covered with pale erect
acicular hairs 0.7-0.8mm long. Lamina 80cm
long; pinnae c. 30 pairs, texture thin but firm;
basal pinnae narrowed very gradually towards
their bases. Largest pinnae 20X1.8cm; base
broadly cuneate; apex narrowly acuminate; edges
lobed to a depth of 2mm (hardly 1/4 towards
costa); costules 4-5 mm apart, at less than 60°;
veins 8-9 pairs, slender, hardly prominent either
side, 1-1 pairs anastomosing, 3 pairs to sides of
sinus-membrane; all parts of lower surface bear-
ing erect slender acicular hairs c. 0.7 mm long,
most densely on costae, very short capitate hairs
also present on and between veins; upper surface
of costae bearing thicker hairs, shorter ones also
on costules and veins. Sori all close to costules,
not elongate, exindusiate; no glands on sporangia.
Distr. Malesia: N.E. Sumatra; known from the
type and SURBECK 814, from Sibuatan.
11. Chingia sambasensis HOLTTUM, Kalikasan 3
(1974) 24.— Dryopteris penangiana var. cal-
vescens sensu COPEL. Philip. J. Sci. 5 (1910) Bot.
283 quoad pl. Brooks. tantum.—Type: C. J.
BROOKS s5.n. Sept. 1908, Sambas, W. Kalimantan,
900 m (BM; K, L, MICH).
Stipe of type lacking (see also below); rachis
red-brown, quite smooth on abaxial surface. Lar-
5
394 FLORA MALESIANA [ser. II, vol. 1
gest pinnae 43x3.3cm, thin, drying light scales, rest glossy, castaneous, sparsely ver-
olivaceous; apex narrowly attenuate; edges rucose; scales to 20mm long, little more than
crenate to a depth of 2-2.5mm; costules 5mm
apart; veins slender, slightly prominent on upper
surface, not on lower, 10-11 pairs, 1-12 pairs
anastomosing, 4 pairs to sides of sinus-membrane;
lower surface of pinnae quite glabrous; upper
surface glabrous except for a few hairs near base
of costae. Sori close to costules except basal
basiscopic one, all slightly elongate, exindusiate;
no glands on sporangia.
Distr. Malesia: Borneo. Besides the type, an
unlocalized Bornean specimen of KORTHALS.
Note. The Korthals specimens (3 sheets at B, 2
at L) include a stipe of 125cm, rather sparsely
scaly near base, scales flat, narrow, long-acu-
minate, on rest of stipe scattered small black
warts and a few narrow dark flat scales. One sheet
bears the apex of a frond which is not fully
expanded; it shows some short capitate hairs on
both surfaces and on sporangia. The largest pinna
is 24 x 2.1 cm, in shape and venation like the type.
12. Chingia christii (COPEL.) HoLTTUM, Kali-
kasan 3 (1974) 25.— Dryopteris ferox var. cal-
vescens CHRIST, Philip. J. Sci. 2 (1907) Bot.
193. — Cyclosorus christii COPEL. Fern FI. Philip.
(1960) 362 excl. ESCRITOR BS 20707. — Thelyp-
teris zamboangana REED, Phytologia 17 (1968)
325, nom. nov. (not T. christii (C. CHR.)
REED). — Type: COPELAND 1721, San Ramon,
Mindanao, 800 m (MICH; B, NSW). — Fig. 8h.
Stipe to 200cm long (ELMER), castaneous,
scaly at base only; scales 15 x 0.5 mm, thick, dull
brown; upper part of stipe, and rachis, smooth on
abaxial surface. Lamina 100cm or more long;
basal pinnae narrowed at asymmetric base, with
stalks 2mm long. Largest pinnae of type 25-35 cm
long, 2.0-2.7cm wide (but see note below), very
firm, drying red-brown; apex long-attenuate with
subentire cauda 3 cm long; edges crenately lobed
to a depth of 2-3 mm, lobes falcate, acute; cos-
tules 4.5-5.5 mm apart; veins to 10 pairs, prom-
inent on lower surface, 1 pairs anastomosing, 3
pairs to sides of sinus-membrane; lower surface of
type quite glabrous, of ELMER 10182 bearing a
few small capitate hairs and sessile glands on
costules and veins; upper surface hairy on costa
only. Lower sori gradually divergent from costule,
several pairs somewhat elongate; no indusia;
sporangia bearing small glands.
Distr. Malesia: Philippines (Mindanao, Negros,
Mindoro).
Note. With ELMER 10182 (from Negros) in UC
is a note by the collector that the frond was 4m
long, with largest pinnae 52 x 3.3 cm.
13. Chingia paucipaleata HOLTTUM, Kalikasan 3
(1974) 26.—Type: M. G. PricE 781, Tignoan,
Infanta, Quezon Prov., Luzon (PNH; K).
Stipe to 150cm long, basal 15cm covered with
0.5mm wide, thin, contorted when dry, bearing
few setae. Lamina to 100cm long; pinnae to at
least 30 pairs, subopposite, 3.5cm apart; several
lower pairs slightly narrowed at their bases, ses-
sile. Largest pinnae 24 x 1.5 cm; base subtruncate;
apex narrowly attenuate; edges lobed 2/5 towards
costa (to a depth of 3mm), lobes falcate at tips;
costules 3.5mm apart; veins to 9 pairs, slender,
distinct but hardly prominent on lower surface,
1-15 pairs anastomosing, | pair to sides of sinus-
membrane; lower surface of costae and costules
glabrescent (some acicular hairs distally on young
fronds), on veins and surface between them rather
abundant small sessile glands; on upper surface of
rachis sparse warts representing bases of scales,
hairs on costae pale, 0.8mm long. Sori all near
costules, not elongate; indusia small, with a few
small glands or short hairs.
Distr. Malesia: Philippines (Luzon).
Note. ESCRITOR BS 20707, from Tayabas
Province, Luzon, (in Herb. Kew.) agrees with the
above description but has pinnae to 32 x 2 cm and
smaller indusia.
14. Chingia horridipes (v.A.v.R.) HOLTTUM, Kal-
ikasan 3 (1974) 26.— Dryopteris horridipes
v.A.v.R. Bull. Jard. Bot. Btzg II, 28 (1918) 23. —
Thelypteris horridipes (v.A.v.R.) REED, Phy-
tologia 17 (1968) 283.— Type: BUNNEMEIJER
9684, Mt Talamau, Sumatra (BO).
Scales at base of stipe to at least 10x 1.5mm,
short-hairy throughout; above base of stipe scat-
tered small warts (bases of former scales), up-
permost part of stipe, and rachis, with sparse
warts on adaxial surface only. Lamina 100cm or
more long, texture subcoriaceous; lowest pinnae
narrowed towards their bases. Largest pinnae
35 x 2.2 cm; base truncate; apex acuminate; edges
lobed almost half-way to costa; costules to 5mm
apart; veins to 15 pairs, grooved on upper surface,
1-1) pairs anastomosing, 15-2 pairs passing to
sides of sinus-membrane; lower surface of costae
and costules densely covered throughout with
acicular hairs to nearly | mm long, fewer hairs on
and between veins; upper surface hairy on costae
and costules only. Sori near costules; indusia
minute, short-hairy; sporangia lacking glands.
Distr. Malesia: Central West Sumatra (Mts
Talamau and Kerintji), at 2200-2750 m.
15. Chingia perrigida (v.A.v.R.) HOLTTUM, Kal-
ikasan 3 (1974) 27.—Phegopteris perrigida
v.A.v.R. Bull. Jard. Bot. Btzg II, 16 (1914) 27;
Handb. Suppl. (1917) 317. — Type: MATTHEW S33,
Mt Merapi, Sumatra (BO).
Dryopteris ferox var. calvescens sensu v.A.v.R.
Bull. Jard. Bot. Btzg III, 2 (1920) 150, quoad
Koorders 43567 tantum.
Stipe to 100cm long, densely scaly near base
1981]
only; basal scales to at least 15 x 1.5 mm, stiff, flat
at base, apex very slender; no scales on abaxial
surface of rachis. Lamina to 120 cm long; pinnae
to at least 28 pairs; basal pinnae narrowed in basal
3cm, basal lobes somewhat elongate. Largest
pinnae 30X2.3cm; base truncate or broadly
cuneate; apex evenly attenuate; edges lobed 1/3
towards costa or a little more deeply; costules
4.5-5 mm apart; veins 8-12 pairs, grooved on up-
per surface and prominent on lower, basal pair
meeting at base of sinus-membrane, next 2 pairs
to sides of the membrane; lower surface bearing
short capitate hairs on all parts and a varied
number of acicular hairs (sometimes none) on
costules and veins. Sori near costules, basal ones
not divergent; indusia very small, bearing short
capitate hairs; many short capitate hairs on
sporangia.
Distr. Malesia: Central Sumatra, central
Malaya, East Java, Lesser Sunda Is. (Flores).
Ecol. In open places at 1500-2000 m.
Notes. Found in Malaya for the first time at
Genting Highlands in 1977, beside a recently-made
road at 1500 m. Specimens from the Tengger and
Ijang mountains in East Java have fewer capitate
hairs on the lower surface; they were wrongly
named Dryopteris ferox var. calvescens by
v.A.v.R. and one has the MS name D. ferox var.
mitis ROSENST. KOORDERS 37503 (Tengger Mts)
has a seta on a few sporangia, capitate hairs on
others.
16. Chingia supraspinigera (ROSENST.) HOLT-
TUM, Kalikasan 3 (1974) 27.— Dryopteris
supraspinigera ROSENST. Hedwigia 56 (1915)
353.— Cyclosorus supraspinigerus (ROSENST.)
COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78 (1951)
447.— Thelypteris supraspinigera (ROSENST.)
REED, Phytologia 17 (1968) 318. — Type: BAMLER
91, Sattelberg, N.E. New Guinea (S-PA; B, UC).
Stipe more than 40 cm long, scaly at base only;
adaxial side of stipe above base, and of rachis,
bearing scattered small scales (or their wart-like
bases), abaxial side smooth. Lamina 75 cm or more
long, rather thin; basal pinnae slightly narrowed
close to their bases. Largest pinnae 17 x 1.8 cm;
base truncate; apex short-acuminate; edges lobed c.
1/3 towards costa; costules 4mm apart; veins 8
pairs, very slender and hardly prominent either side,
basal pair anastomosing to form a slender excurrent
vein which is joined by the second acroscopic vein
before entering the base of the sinus-membrane,
next 2 pairs to sides of the membrane; lower surface
of rachis and pinnae covered throughout with short
thick hairs with variously thickened and sometimes
wax-encrusted tips; upper surface of costae also
bearing short papillae with a few acicular hairs. Sori
all near costules; indusia small, glandular; sporangia
bearing small glands.
Distr. Malesia: Papua New Guinea (Morobe
Distr., 2 coll.).
THELYPTERIDACEAE (Holttum)
395
Note.The second collection, from 300 m at foot
of the Oomsis Range, is smaller than the type,
with fertile pinnae 12.5 x 1.4m, rather less deeply
lobed.
17. Chingia urens HOLTTUM, Kalikasan 3 (1974)
28.— Type: M. G. PRICE 2367, Mt Makiling,
Batangas Prov. Luzon (PNH; K).
Caudex to 40cm tall; stipe 75cm long, basal
12cm densely scaly, rest with sparse warts; basal
scales to 15 x 1.5mm, rather thin, contorted when
dry, dull brown, not hairy; rachis lacking warts on
abaxial side which is densely covered with short
capitate hairs. Lamina to 75cm long, firm but
rather thin; pinnae 25 pairs; basal pinnae gradually
narrowed in basal 7cm to a quite narrow base
which is auricled, next 4 pairs of pinnae progres-
sively wider at base. Largest pinnae 29 x 2.5cm;
base broadly cuneate; apex narrowly attenuate;
edges lobed almost half-way to costa, lobes fal-
cate distally, acute, fringed with acicular hairs;
costules 4-6 mm apart; veins 10-13 pairs, slender,
not prominent either side, basal pair meeting, or
not quite meeting, just below base of sinus-mem-
brane, 2-3 pairs passing to sides of the membrane;
lower surface bearing + abundant short capitate
hairs throughout, a few acicular hairs on distal
parts of costae only; upper surface of costae
densely covered with thick acicular hairs to | mm
long, sparse shorter acicular hairs on costules and
veins, a few short acicular and capitate hairs
between veins. Sori close to costules except basal
ones, not elongate; indusia small but always dis-
tinct, thin, with short capitate and sometimes 1-2
acicular hairs; sporangia with capitate hairs.
Distr. Malesia: Philippines (Luzon),
known from type.
Ecol. On steep slope inside crater at 960 m.
Note. PRICE reports: ‘““malodorous; skin con-
tact with living fronds produces a mild burning
sensation”.
only
18. Chingia tenerior HOLTTUM, sp. nov.
Pinnae usque 30X*1.9cm, usque 2/3 costam
versus lobatae, textura tenues; venae graciles, non
prominentes, infimae vel ad basin membranae
sinus junctae vel ibi conniventes non junctae, vena
acroscopica secunda vel ad latus membranae vel
ad marginem terminata; sori exindusiati; sporan-
gia pilis capitatis praedita.— Type: RAMOS BS
33066, near Baguio, Ilocos Norte Prov. Luzon (P;
US).
Stipe scaly at base only; scales thin, to 10mm
long, hardly 1mm wide at base. Pinnae to 30
1.9cm, thin; apex gradually attenuate; edges
lobed more than half-way (to 2/3) towards costa;
costules 4mm apart; veins to 15 pairs, very fine
and not prominent either side, basal pair either
meeting just below the sinus-membrane or touch-
ing the sides of the base of the membrane without
meeting (base of membrane not decurrent be-
396 FLORA MALESIANA [ser. II, vol. 1°
Asan
ae ES
Fig. 9. Plesioneuron marattioides (ALSTON) HOLTTUM. a. Base of stipe, X 3, with scale and spine, x 2;
b. lower pinna, x3; c. one pinna-lobe, x 1.— P. tuberculatum (CESATI) HOLTTUM. d. Lower pinna,
x2; e. one pinna-lobe, x3; f. sori (one with indusium removed), x 16.— P. savaiense (BAK.) HOLT-
TUM. g. Basal pinna, x 1; h. one pinna-lobe, <3; i. sorus, X 16 (a-c MILLAR & HOLTTUM 15996, d
FLOYD 5595, e-f BRASS 27112, g-i PRICE 2605).
1981]
THELYPTERIDACEAE (Holttum)
397
tween them), second acroscopic vein ending at
side of membrane or at margin just above base of
sinus; lower surface throughout bearing scattered
slender acicular hairs c. 0.4mm long, short capi-
tate hairs also present on veins and surface be-
tween them; upper surface of costae bearing
long, very few hairs on costules, no others. Sori
near costules, exindusiate, not elongate; short
capitate hairs on sporangia.
Distr. Malesia: Philippines: (Luzon), only
known from the type which was distributed as
Dryopteris extensa.
rather sparse acicular hairs more than 0.5mm
13. PLESIONEURON
HOLTTUM, Blumea 22 (1975) 232; Allertonia 1 (1977) 186; L. R. ATKINSON,
Phytomorphology 25 (1975) 45 (gametophyte).— Mesophlebion subg.
Plesioneuron HoLttTuM, Blumea 19 (1971) 30. — Fig. 1j, 9.
Caudex erect or suberect, rarely prostrate. Stipe always scaly at base,
sometimes throughout, bases of scales usually thickened, in some species
spine-like and persistent; lowest pinnae not or little reduced, variously
narrowed at their bases, sometimes with a free basal lobe (or more than
one); pinnae of firm texture, deeply lobed, usually with somewhat swollen
or elongate basal aerophores, lower surface often verrucose when dry;
veins all free, tips of lowest from adjacent costules usually touching sides
of a sinus-membrane which may be decurrent between them as a hairy
ridge almost to the costa, in the most deeply lobed pinnae both basal veins
passing to the margin above base of sinus; basal basiscopic vein arising
from costa near its costule, or from base of the costule; unicellular acicular
hairs on lower surface of costae and costules various, usually short and
stiff, thick brown hairs also sometimes present, rarely short capitate hairs,
reduced scales few on most species; short acicular or capitate hairs some-
times present between veins on upper surface. Sori usually medial or
inframedial, in a few cases supramedial; indusium, when present, usually
firm and dark; sporangia often bearing small red or yellow glands, or setae,
near annulus; hairs on sporangium-stalk sometimes acicular, never bearing
red glands; spores dark, spinulose, in most observed cases (winged in three
species), in about 15 species not yet observed.
Type species: Plesioneuron tuberculatum (CESATI) HOLTTUM.
Distr. Malesia & Pacific: Borneo (2 spp.); Moluccas (Tidore, Batjan, Buru, Amboina, 2 spp.); Mindanao
(1 sp.); New Guinea & Bismarck Archipelago (32 spp.); islands of the Pacific, to Tahiti (12 spp.).
Ecol. In lowland and mid-mountain forest (and in a few cases to 2700 m) often by streams, sometimes
pendulous from rock-crevices.
Cytol. Chromosome number 36 (P. fulgens only).
Taxon. The name Plesioneuron refers to the basal basiscopic veins in each pinna-lobe which always
arise close to their costules, never far from them as in Mesophlebion. Plants of many species are similar
in general aspect, and to some extent in venation, to Mesophlebion, to which genus they were assigned
as a subgenus in 1971, but they differ from Mesophlebion in sporangia and spores, also, in a majority of
species, in scales, and (in the only case investigated) in gametophytes. They appear to be related to
Chingia and not very closely to any other genus. COPELAND included them in Lastrea, and published a
key to the species of that genus in New Guinea (Philip. J. Sci. vol. 78, 1951) but he did not characterize
them clearly nor distinguish them as a group. There are undoubtedly many species in New Guinea, but
specimens of some are few or imperfect, so that more information about them is needed, and the
present account probably needs some modification. Some type specimens may represent immature
plants (an example is P. ophiura) and more local collecting is needed to establish this; in other cases
small plants probably do represent the mature form of a species. Recent collections certainly include
plants of previously undescribed species, and probably more remain to be discovered.
398 FLORA MALESIANA [ser. II, vol. 1°
Several species show some degree of difference between lobes on acroscopic and basiscopic
sides of a pinna, but there is no sharp distinction between these and those with no clear difference.
However, in a few cases the difference is very marked and can be used in the key to the identification
of species.
The sinus-membrane in Plesioneuron is in all cases distinctly decurrent from the base of the sinus as a
more or less hairy ridge on the lower surface, sometimes almost reaching the costa. This condition is
closely similar to that of Dryopteris subg. Steiropteris C. CHR. (Monogr. Dryopt. I, 1913, 161), a group
of species in the American tropics. These species differ from Plesioneuron as follows: rhizome
wide-creeping; septate hairs on lower surface of rachis and costae; neither glands nor setae on
sporangia; spores with translucent wing and a few cross-wings (this type of spore occurs in a few
species of Plesioneuron). Two aberrant species included with doubt by CHRISTENSEN in subg.
Steiropteris are more similar to Mesophlebion and are mentioned under that genus.
I believe that characters of scales are often of diagnostic significance, but in many specimens they are
not well represented; more information about them may help to clarify specific distinctions.
KEY TO) THE SPECIES
1. Stipe and rachis bearing + copious dark spines (bases of former scales); sori in most cases
exindusiate.
2. Pinnae not over 4.5 cm wide; costules not over 5 mm apart.
3. Pinnae to 4.5 cm long; pinna-lobes concave beneath; stipe-scales to4mm long . . 1. P. pullei
3. Pinnae larger; pinna-lobes not concave beneath; stipe-scales 8 mm or more long.
4. Sori exindusiate.
5. Sporangia bearing several long setae reoteh lat bee ie wr te, oe Bln ee gem oe beam CHISEL
5. Sporangia lacking setae.
6. Pinnae to 9 x 1.2 cm; basal veins both passing to margin above base of sinus 3. P. medusella
6. Pinnae of well-grown plants much larger; basal veins both ending beside sinus-membrane.
7. Stipe-scales terete at base and apex, less than 0.5mm wide in middle; pinnae to 4.5cm
wide fay . . . 4, P. woodlarkense
7. Stipe-scales flat except at base, largest at ‘least Imm wide: pinnae rarely more than 3cm
wide chlo Gaubitta ha sole Macs cca: ccs ca bi ce ee GY aot a ce oe Ee On opteroideum
4. Sori indusiate ae 2 . . . . . 6. P. varievestitum
Me Largest pinnae 7-12 cm wide with costules 10-12) mm n apart.
6a. All pinna-lobes or pinnules auricled at base on basiscopic side; veins forked or pinnate in the
auricles) =.) 5, Psu Li Seip pe, edu Sueen y Gr, Se add ost os teen peas) ye notabile
6a. Pinna-lobes not thus auricled.
7a. Pinnae lobed almost to costae; lobes not separately adnate . . . . . 8. P. septempedale
7a. Pinna-lobes in basal half of pinnae separately adnate to costa . . . . . .49. P. marattioides
1. Stipe (except base) and rachis smooth; scales, if present, not spine-like; sori in almost all species
indusiate.
8. Pinnae 1-5 pairs; apical lamina pinna- -like; costules 5-7 mm apart . . . . . . 10. P. fulgens
8. Pinnae commonly to at least 10 pairs, or if fewer costules 3-4 mm apart.
9. No indusia; no setae on sporangia.
10. Aerophores elongate; sori near costules .......... =. +. +. . M1. P. belense
10. Aerophores not elongate; sorimedial . . . Sere go 2 oe IP Swanriense
a cae present, or if absent all sporangia setiferous.
Sporangia all setiferous; no glands present.
i Sori supramedial; sporangia with 1-3 short setae Cae » 2 OS DS SPs royenit
12. Sori medial or inframedial; sporangia bearing several long setae.
13. Veins 10-12 pairs; largest pinnae not over 12x 2.5cm, almost sessile . . 14. P. savaiense
13. Veins 14-16 pairs; largest pinnae 21 X 3.3 cm, with stalks 34mm long . . . .15. P. altum
11. Sporangia usually with glands; setae, if present, short and not on all sporangia, sometimes
alternating with glands.
14. Sori supramedial.
15. Pinnae to 20 x 3 cm; veins 15-18 pairs.
16. Indusia overlapping margin; basal acroscopic lobe of lower pinnae elongate with forked
WEIS 6 « . . . .16. P. cystodioides
16. Indusia not overlapping margin; basal acroscopic lobe of lower pinnae not enlarged
17. P. subterminale
15. Pinnae smaller; veins not over 12 pairs.
17. Both basal lobes on basal pinnae free and short-stalked . . . . . .18. P. bipinnatum
1981]
THELYPTERIDACEAE (Holttum)
399
17. Basal lobes of basal pinnae not free
14. Sori medial or inframedial.
19. P. quadriquetrum
18. Pinna-lobes on both sides of costa conspicuously falcate.
19. Upper surface glabrous between veins; pinnae 8 pairs
20. P. falcatipinnulum
19. Upper surface bearing hairs between veins; pinnae 20 pairs.
20. Hairs on lower surface of rachis and costae 1mm long, on upper surface between veins
appressed
21. P. kostermansii
20. Hairs on lower surface of achic! and costie 0. 3 mm longs on | upper surface between veins
short-erect
22. P. angiensis
18. Pinna-lobes not conspicuously falcate on ‘both sides of costa.
21. Basal 10-12 pairs of pinna-lobes separately adnate to costa
23. P. sandsii
21. At most 1 free pinna-lobe separately adnate.
22. Pinnae to 25 x S5cm, all except distal ones stalked; lower pinnae with stalks 3-6 mm; indusia
small, caducous
. 24. P. tuberculatum
22. Pinnae smaller; most pinnae almost sessile, basal ones ; with stalks at most 2-3 mm long;
indusia firm, persistent.
23. Veins 20 pairs or more.
24. Basiscopic lobes of all pinnae strongly falcate and shorter than acroscopic lobes; basal
scales to 20 mm long, very narrow
24. Basiscopic lobes,
shorter, usually wider.
. 25. P. attenuatum
if falcate, not or little shorter tani acroscopic; basal scales much
25. Basal acroscopic lobe of basal pinnae free, wider than the rest, with some forked
veins
25. Basal acroscopic lobe of basal pinnae ‘otherwise.
26. Lower surface quite glabrous.
27. Pinnae c. 11 x 2cm; basal acroscopic lobe of basal pinnae reduced. 27.
27. Pinnae c. 22 x 3 cm; basal acroscopic lobe of basal pinnae elongate
. 26. P. platylobum
P. crassum
. 28. P. stenura
26. Lower surface of costules at least bearing acicular hairs.
28. Pinnae lobed to 0.5 mm from costa
28. Pinnae lobed to 1-2 mm from costa
23. Veins 10-15 pairs.
. 29. P. ophiura
30. P. costulisorum
29. Basal pinnae with stalks 2 mm or more long.
30. Pinnae lobed to 1.5—2 mm from costa
. 31. P. doctersii
30. Pinnae lobed to less than 1 mm from costa.
31. Basal scales thick, narrow; sori a little inframedial
31. Basal scales c. 5 x 1.5mm; sori near costules
29. Basal pinnae sessile or nearly so.
32. P. rigidilobum
. 29. P. ophiura
32. Pinnae to 6cm long; lobes on basiscopic side strongly falcate, on acroscopic side almost
straight
. 33. P. wantotense
32. Pinnae longer; fobess on Sbasiscapic| side not greatly different from those on acroscopic
side.
33. Aerophores small, dark; sori medial or a little inframedial.
34. Some sporangia bearing setae
34. Sporangia lacking setae, glands usually present :
33. Aerophores pale, slender, 1 mm long; sori near costules
1. Plesioneuron pullei HOLTTUM, Blumea 22
(1975) 236.—Type: PULLE 905, W. New Guinea,
Mt Hellwig, 2600 m (L; BM).
Caudex short, massive, erect or suberect. Stipe
of fertile fronds 25-30 cm long (shorter on sterile),
red-brown, glossy, bearing many black spine-like
scales 4mm long, scale-bases persistent. Lamina
to 30cm long, texture coriacous; pinnae 30 pairs
or more (but see below); basal 2-3 pairs of pinnae
variably somewhat reduced, lowest on a large
frond 2.2 cm long, basal acroscopic lobe free and
elongate. Largest pinnae 4.5 x 0.9—1.0.cm, sessile;
aerophores not enlarged; base truncate; apex
obtuse; edges lobed to 1 mm from costa or more
deeply; lobes strongly concave on lower surface,
. 34. P. kundipense
35. P. dryas
36. P. croftii
their edges much reflexed, entire, tips rounded;
costules 2.5mm apart; veins 4-5 pairs, thick and
prominent beneath, invisible on upper surface,
basal acroscopic vein passing to side of sinus-
membrane, basal basiscopic vein arising from
costa; lower surface of rachis bearing scattered
glossy spine-like scales 1.5mm long and stiff
spreading hairs 0.7 mm long, costae and costules
hairy as rachis, slender erect hairs present on
surface between veins, copious hairs on and near
margins of lobes; upper surface of costa deeply
grooved with short hairs, capitate hairs present on
rest of surface (often abraded on old specimens).
Sori medial, exindusiate; sporangia lacking glands
or setae.
400
Distr. Malesia: New Guinea; known from the
type and PULLEN S117 (see below).
Note. PULLEN 5117, from Western Highlands
in Papua New Guinea, Kubor Range, at 3450 m,
differs as follows: pinnae 18-20 pairs on a frond
30cm long; costules 34mm apart; veins 6-7
pairs; a very small hairy indusium present. The
plant was on a steep bank near a rock-face in
moss-forest, the fronds pendulous.
2. Plesioneuron fuchsii HOLTTUM, Blumea 22
(1975) 236. —Type: H. P. FUCHS 21477, Sabah,
Mt Kinabalu, North Face, Goking’s valley, 2715 m
(eK US).
Caudex not seen; stipe 60cm long, densely
covered throughout with erect black spines,
mostly broken, all except basal ones terete with
sparse setae; basal scales rigid, dark brown, 10-
15mm long, 0.5mm wide, with sparse setae.
Lamina to at least 75 cm long; pinnae 35 pairs or
more, rigid when dry; basal pinnae a little nar-
rowed at their bases. Largest pinnae 21 X 1.8 cm;
base truncate; apex caudate-acuminate; edges
lobed to 2.5-3 mm from costa; lobes slightly fal-
cate, entire, ciliate; costules 44.5 mm apart; veins
10-11 pairs, prominent beneath, slightly impressed
above, basal veins both touching sides of sinus-
membrane; lower surface of rachis bearing black
spines as stipe, also some intact scales which are
4mm long distally to 7-8mm at base of stipe,
similar scales 4mm long on costae; costae, cos-
tules, veins and surface between veins bearing
rather sparse slender erect acicular hairs and a
few capitate hairs: upper surface bearing terete
scales on edges of the rachis-groove, on costae
copious brown acicular hairs with a few on cos-
tules also. Sori about medial, small, exindusiate;
sporangia bearing long setae.
Distr. Malesia: Sabah (Mt Kinabalu), 2 collec-
tions (the second is CLEMENS 33719).
Ecol. At 2300-2700m, overhanging river in
moss-forest.
3. Plesioneuron medusella HOLTTUM, sp. nov.
Caudex erectus, tenuis; stipes usque 25 cm lon-
gus, cum rachi omnino paleis angustis usque 8 mm
longis vestitus; pinnae usque 9 X 1.2 cm, profunde
lobatae; venae 6~7-jugatae, infimae ambae ad
marginem terminatae; costae subtus pilis usque
1 mm longis vestitae; sori parvi, prope costulas,
exindusati; sporangia non setifera.— Type: A. C.
JERMY 14133, Sarawak, Gunong Mulu National
Park, G. Api, 1500 m (BM).
Caudex erect, 10mm diameter; stipe 12-25cm
long, densely scaly at base, more sparsely up-
wards and on rachis, largest scales c. 8mm long,
terete or flattened at the thickened base, above
base to 0.4mm wide, apex filiform; also on stipe
slender acicular hairs and many short blunt or
capitate hairs. Lamina to 35 cm long; pinnae c. 30
pairs, slightly overlapping; basal pinnae slightly
FLORA MALESIANA
[ser. II, vol. 1°
reduced and narrowed towards their bases, with
elongate crenate basal acroscopic lobe. Largest
pinnae 6.5-9cm long, 1.2cm wide above base;
base truncate with both basal lobes a little longer
than the next pair; apex evenly attenuate to tip;
sides lobed to 1mm from costa or more deeply,
lobes not or slightly falcate; costules 2.5-3 mm
apart, at more than 60° to costa; veins 6-7 pairs,
prominent on lower surface, not on upper, basal
veins both passing to margin above base of sinus;
lower surface of rachis bearing scales 34mm
long, dark and very narrow, also sparse acicular
hairs less than 1mm long and more abundant
short blunt or + capitate hairs, hairs on costae
similar but sometimes a little shorter, sparse aci-
cular and more abundant short capitate hairs
present on surface between veins; upper surface
of rachis and costae with more abundant acicular
hairs, few on costules and veins, between veins
many short capitate hairs and a few acicular ones
(often abraded from old fronds). Sori small, near
costules except basal ones, exindusiate; sporangia
sometimes bearing a shrivelled capitate hair;
spores black, spinulose.
Distr. Malesia: Borneo (Sarawak: Mt Mulu,
1500 m), only known from the type.
Ecol. “In Pandanus zone over limestone pin-
nacles; in crevices of limestone getting consider-
able light”’.
Note. The specific epithet refers to the dense
spreading slender scales which cover the uncoiling
young fronds in a medusa-like manner.
4. Plesioneuron woodlarkense (COPEL.) HOLT-
TUM, Blumea 22 (1975) 236. — Cyathea woodlar-
kensis COPEL. Philip. J. Sci. 9 (1914) Bot. 1;
HOLTTUM, FI. Males. II, 1, pt 2 (1963) 158. — Type:
C. KING 384, Woodlark Island, d’Entrecasteaux Is.
(MICH; NSW).
Caudex erect, 22cm tall (FLOYD); stipe to
100 cm long, bearing very short acicular hairs and
short black spines throughout, scales (fully per-
sistent at base of stipe) c. 12 mm long, dull brown,
terete at base, flat but less than 0.5mm wide
above base, terete distally, bearing short acicular
hairs; rachis also bearing scattered short black
spines on lower surface, on upper surface some
complete bristle-like scales. Lamina to 150cm
long, texture firm-herbaceous; pinnae many pairs;
basal pinnae with stalks 3 mm long, gradually nar-
rowed in basal 4-5 cm, basal lobes 5—7 mm long.
Largest pinnae 42x 4.5cm; base truncate; apex
caudate-acuminate; edges lobed to 1-2 mm from
costa, lobes slightly falcate on both sides of the
costa; costules 5 mm apart, almost at right angles
to costa; veins to more than 30 pairs, slender,
basal ones touching sides of sinus-membrane;
lower surface of rachis and costae bearing rather
sparse pale hairs 1mm long, shorter hairs on
costules and veins, and a few on surface between
veins; upper surface of costae bearing dense red-
1981]
THELYPTERIDACEAE (Holttum)
401
dish hairs 1 mm long, shorter hairs scattered on
veins and surface between them. Sori near cos-
tules, exindusiate; sporangia not setiferous; spores
dark.
Distr. Malesia: Papua New Guinea (Woodlark
Island, New Britain, New Ireland) at 700-900 m.
Notes. In 1914 COPELAND wrongly cited the
number 284 for the type; he again cited the type
wrongly, as 383, in Philip. J. Sci. 77 (1947) 124. Of
other specimens, CROFT 322 from southern New
Ireland, exactly matches the type; FLOYD 6529,
from New Britain, agrees in details of indument
and sori but has pinnae to 33 x3 cm. It should be
noted that the number 384 occurs also on some
specimens which do not belong here; at L, a
specimen of Amphineuron ceramicum, at P a
specimen of A. immersum.
5. Plesioneuron dryopteroideum (BRAUSE)
HOLTTUM, Blumea 22 (1975) 237. —Alsophila
dryopteroidea BRAUSE, Bot. Jahrb. 56 (1920) 70.
—Cyathea atrispora DOMIN, Acta Bot. Bohem. 9
(1930) 95, nom. nov.—Dryopteris atrispora
(DOMIN) C. CHR. Brittonia 2 (1937) 296.—
Thelypteris dryopteroidea (BRAUSE) REED, Phy-
tologia 17 (1968) 273.—Type: LEDERMANN 11897,
N.E. New Guinea, Sepik Distr. 2070 m (B).
KEY TO THE VARIETIES
1. Stipe-scales 10mm long; acicular hairs on
lower surface of costae to 0.5 mm long.
2. Acicular hairs present on lower surface of
rachis and costae a. var. dryopteroideum
2. Hairs on lower surface of rachis and costae
not acicular . . . . . _ b. var. buruense
1. Stipe-scales to 18mm long; acicular hairs on
lower surface of costae 1 mm long
c. var. pilosum
a. var. dryopteroideum
Caudex erect, to at least 30cm tall; stipe 60—-
120 cm long, basal part densely covered with dark
brown rigid glossy scales c. 10mm long, at least
1 mm wide with attenuate apex, distal part of stipe
and both surfaces of rachis bearing less abundant
similar but shorter scales or their spine-like bases.
Lamina to 100 cm or more long, texture firm; lower
pinnae narrowed towards their bases, with stalks
2mm long. Largest pinnae commonly 20 x 2cm,
largest on type 35 x 2.7 cm; apex acuminate with
cauda to 3cm long; edges lobed to c. 2mm from
costa; lobes falcate, entire or slightly crenate,
ciliate; costules 3.5-4.5 mm apart, at more than 60°
to costa; veins commonly 12-15 pairs, basal veins
both passing to sides of sinus-membrane; lower
surface of rachis and costae bearing narrow dark
scales and short acicular hairs, a few short acicular
hairs present on costules with a variable number of
small sessile or subsessile glands which in some
specimens are much swollen and yellow (apparently
infected by a fungus), between veins rather sparse
short erect hairs; on upper surface acicular hairs
confined to rachis and costae, or with a few short
ones on costules. Sori near costules, exindusiate;
sporangia bearing neither setae nor glands; spores
dark, spinulose.
Distr. Malesia: Eastern New Guinea, at 2000-
2700 m.
b. var. buruense HOLTTUM, var. nov.
A typo speciei differt: rachi costisque subtus pilis
brevibus capitatis solum vestitis. — Type: STRESE-
MANN 387, N.W. Buru, G. Fogha, 1900m (L).
Known only from the type collection which
consists of part of a frond with pinnae to 20 x 2 cm.
Distr. Malesia: Moluccas (Buru I.).
c. var. pilosum HOLTTUM, var. nov.
A typo speciei differt: paleis stipitis usque 18 mm
longis; costis costulisque subtus pilis 1 mm longis
copiose vestitis. — Type: J. R. CRorr LAE 68379,
New Ireland, Hans Meyer Range, 1700 m “‘landslip
community” (L).
Known only from the type collection; stipe 50 cm
long, lamina 150 cm long; largest pinnae 25 x 2.2 cra;
veins 18 pairs.
Distr. Malesia: E. New Guinea (New Ireland).
6. Plesioneuron varievestitum (C. CHR.) HOLT-
TUM, comb. nov.— Dryopteris atrispora var.
varievestita C. CHR. Brittonia 2 (1937) 296. —
Lastrea varievestita (C. CHR.) COPEL. Gen. Fil.
(1947) 140; Philip. J. Sci. 78 (1951) 429. — Thelyp-
teris varievestita (C. CHR.) REED, Phytologia 17
(1968) 323. Type: BRASS 4996, E. New Guinea,
Mt Tafa, 2400 m (BM; BO, BRI).
KEY DO THE VARIETIES
1. Rachis and costae bearing acicular hairs on lower
surface BE ot a. var. varievestitum
1. Rachis and costae bearing only short, blunt or
capitate, hairs on lower surface
b. var. obtusipilum
a. var. varievestitum
Caudex to 100cm tall; stipe 47-70cm long,
stramineous above base, base densely covered
with glossy castaneous scales to 10 x 2mm, scat-
tered smaller scales or their broken spine-like
bases present on distal part of stipe and on rachis
and costae. Lamina 40-75 cm long, texture rigid;
pinnae to more than 30 pairs; basal pinnae some-
what narrowed towards their bases, with stalks
1mm long. Largest pinnae of type 15 x 1.5 cm (of
other specimens ranging from 7.51.2 to 21x
2.3m); apex acuminate, with or without a short
cauda; edges lobed to less than 1 mm from costa;
lobes slightly falcate, entire; costules 3-4.5mm
apart; veins of type 9-10 pairs (on another speci-
402
FLORA MALESIANA
(ser. II, vol. 1°
men to 15 pairs), basal acroscopic vein ending
beside sinus-membrane or near it, basiscopic vein
at margin above base of sinus; lower surface of
rachis bearing acicular hairs to 0.7mm long, of
costae and costules shorter acicular hairs and
small colourless or yellow sessile or subsessile
glands, between veins a variable number of short
erect acicular hairs and a few glands; upper sur-
face of rachis and costae bearing acicular hairs,
between veins a variable number of small sessile
glands. Sori near costules; indusia small, thin,
pale, with a few glands or very short acicular
hairs; sporangia sometimes bearing short capitate
hairs or a short seta (not seen on type); spores
dark, spinulose.
Distr. Malesia: Eastern New Guinea, at 1800-
2800 m.
Notes. CHRISTENSEN and COPELAND failed
to see the indusia, which are partly obscured by
the developing sporangia. As in P_ dryop-
teroideum, tha small glands on pinnae are
sometimes swollen and yellow, perhaps through
infection by a fungus.
b. var. obtusipilum HOLTTUM, var. nov.
A typo speciei differt: rachi costisque subtus pilis
0.1 mm longis vel capitatis vel obtusis praedita;
pagina inferiore pinnarum inter venas pilis aci-
cularibus brevibus pilis brevioribus capitatis in-
termixtis vestita.— Type: NAKAIKE 80, N.E.
New Guinea, Morobe Distr., Wau Subdistr., 1300-
1600 m (K).
Scales on stipe much fewer than on var. varie-
vestitum, and thinner; lower surface of rachis and
costae lacking acicular hairs but bearing many
blunt or capitate hairs c. 0.1mm long; between
veins on lower surface of pinnae very copious
erect acicular hairs and shorter capitate hairs (not
sessile glands); indusia lacking acicular hairs;
sporangia apparently lacking capitate hairs or
setae.
Distr. Malesia: Papua New Guinea; known
from type and STREIMANN & KAIRO NGF 44469,
also from Wau Subdistrict, at 1500m, in aban-
doned garden site (K).
Note. The short blunt hairs on these plants
resemble those on Chingia supraspinigera. They
appear to develop a waxing covering of their tips
when old.
7. Plesioneuron notabile (BRAUSE) HOLTTUM,
Blumea 22 (1975) 237.—Dryopteris notabilis
BRAUSE, Bot. Jahrb. 56 (1920) 91.— Lastrea
notabilis (BRAUSE) COPEL. Gen. Fil. (1947) 139;
Philip. J. Sci.- 78 (1951) 429.— Thelypteris
notabilis (BRAUSE) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 253.— Type: LEDERMANN
11663, N.E. New Guinea, Sepik Distr.,
Schraderberg, 2070 m (B).
Caudex not known. Stipe and frond together
3-4m long; stipe seen 75cm long, densely
covered with erect black hair-pointed spines to
2cm long, bearing short acicular hairs. Lamina
very firm; basal pinnae somewhat reduced, with
basal acroscopic lobe 7X 1.4cm and _ pinnatifid
(teste BRAUSE); suprabasal pinnae opposite, 4—
10cm apart. Largest pinnae 38 x 11.5 cm, lobes in
basal half or more separately adnate to costa and
distinctly auricled on their basiscopic bases, the
basal vein forked or pinnate in the auricle, lobes
of distal part of pinna connected by a very narrow
wing along the costa; costules to 1.7cm apart
(11mm on smaller pinnae); veins to more than 30
pairs, slender and prominent both sides; lower
surface glabrous except for hairs on margins of
lobes and sometimes a few on costules; upper
surface glabrous except for hairs on costae and
costules. Sori large, near costules, exindusiate;
sporangia not setiferous; spores very dark,
minutely verrucose.
Distr. Malesia: Papua New Guinea. Known
only from type and another collection from same
locality (LEDERMANN 11991).
8. Plesioneuron septempedale (ALSTON) HOLT-
TUM, Blumea 22 (1975) 238. — Dryopteris septem-
pedalis ALSTON, J. Bot. 78 (1940) 227; Nova Guinea
n.s. 4 (1940) 111, t, 8, f. 11. — Type: L. E. CHEES-
MAN 1383, W. New Guinea, Japen I. (BM; LAE).
Caudex not known; frond with stipe 215 cm
long (collector); base of stipe bearing many dark
spine-like bases of scales which bear short rigid
hairs; pinnae almost opposite, widely spaced,
lower ones slightly reduced, texture rigid. Largest
pinnae 40x12cm; base somewhat narrowed,
basal lobes + deflexed; apex short-acuminate;
edges lobed to within | mm from costa; lobes at
right angles to costa, acuminate, entire; costules to
12mm apart; veins to more than 50 pairs, im-
mersed and only just visible on surface; lower
surface quite glabrous; upper surface of costae
and costules bearing sparse short hairs. Sori near
costules, basal ones divergent, exindusiate; nei-
ther setae nor glands on sporangia; spores dark.
Distr. Malesia: New Guinea (Japen I.), only
known from the type.
9. Plesioneuron marattioides (ALSTON) HOLT-
TUM, Blumea 22 (1975) 238. — Dryopteris marat-
tioides ALSTON, J. Bot. 78 (1940) 227; Nova
Guinea n.s. 4 (1940) 110, pl. 7, f. 7, 8. — Type:
CLEMENS 4809, N.E. New Guinea, Morobe
Distr., Sambanga (BM). — Fig. 9a-c.
Caudex to 30cm tall (HOOGLAND & PULLEN);
stipe 100cm or more long, pale, minutely hairy,
densely covered near base with erect dark spines,
less densely upwards and on rachis, spines to
8mm long, terete to their tips and bearing short
hairs throughout; scales at base of stipe to at least
20mm long, medium brown, glossy, 2mm wide
near base, apex filiform, bearing short stiff hairs
especially on margins. Lamina to 200 cm or more
long, thick; pinnae 25 pairs or more, all sub-
opposite, several pairs of lower pinnae gradually
1981]
THELYPTERIDACEAE (Holttum)
reduced, lowest 20cm long. Largest pinnae 40cm
long, 8-l11cm wide, lobes separately adnate to
costa in basal half of pinna, largest lobes 5.8 x
0.9cm, acuminate, those near base deflexed and
somewhat reduced; distal half of pinna lobed al-
most to costa; costules to 10mm apart near base
of pinna; veins 35-40 pairs or more, thick and
slightly prominent, basal acroscopic vein in larger
lobes sometimes forked; lower surface quite
glabrous apart from short hairs on margins of
lobes; upper surface of costa densely covered
with thick dark brown hairs, a few shorter hairs
also on costules. Sori near costules except basal
ones, exindusiate; no hairs nor glands on sporan-
gia; spores dark with many small wings of irregular
shape.
Distr. Malesia: N.E. New Guinea, at 1800-
2300 m, several collections from widely separated
localities, in Nothofagus forest.
Note. Young plants were grown from spores at
Kew but died after attaining a height of 30cm.
They had rather abundant short orange capitate
hairs on the lower surface of costules and veins,
also a few between veins; such hairs are probably
abraded from fronds of mature plants.
10. Plesioneuron fulgens (BRAUSE) HOLTTUM,
Blumea 22 (1975) 238.—Dryopteris fulgens
BRAUSE, Bot. Jahrb. 56 (1920) 89. — Thelypteris
fulgens (BRAUSE) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 251.— Type: LEDERMANN
11004, N.E. New Guinea, Sepik Distr. 1300 m (B).
Dryopteris hunsteiniana BRAUSE, Bot. Jahrb.
56 (1920) 79. — Thelypteris hunsteiniana
(BRAUSE) REED, Phytologia 17 (1968) 284. —
Type: LEDERMANN 11058, loc. as D. fulgens (B).
Mesoneuron wantotense sensu. HOLTTUM,
Blumea 13 (1965) 134. — Fig. 1j.
Caudex creeping, 4-5 mm diameter; stipe 25-
60 cm long, smooth except base, basal scales 11 x
1.5mm, stiff, hairy. Lamina 30-45 cm long, very
firm; pinnae 4-5 pairs, lowest pinnae a little nar-
rowed towards their bases; apex of lamina pinna-
like. Largest pinnae 16-20cm long, 3.5-4.2 cm
wide; apex acuminate with subentire cauda 2-3 cm
long; edges lobed to 2-3mm from costa;
lobes equal and slightly falcate on the two sides of
costae; costules 7mm apart, at a wide angle to
costa; veins 16-20 pairs, basal acroscopic vein
touching side of sinus-membrane which is decur-
rent as a ridge on the lower surface almost to the
costa, basal basiscopic vein arising from the costa;
lower surface sparsely hairy on sinus-membranes
and margins of lobes, sometimes a few hairs also
on costules; upper surface with coarse dark brown
hairs on costa. Sori near but not touching cos-
tules; indusia firm, dark, with a few short hairs;
slender acicular hairs present on receptacle with
sporangia; sporangia with either a seta or a gland;
spores dark with rather well-spaced short spines.
Distr. Malesia: Eastern New Guinea.
Ecol. At 1300-2000m, on steep banks of
streams, sometimes among rocks, in forest.
11. Plesioneuron belense (COPEL.) HOLTTUM,
Blumea 22 (1975) 239.— Dryopteris belensis
COPEL. Univ. Cal. Publ. Bot. 18 (1942) 220. —
Lastrea belensis COPEL. Gen. Fil. (1947) 138;
Philip. J. Sci. 78 (1951) 433, pl. 21. — Thelypteris
belensis (COPEL.) REED, Phytologia 17 (1968)
263. —- Type: BRASS 11509, W. New Guinea, Bele
River (MICH; L).
Caudex stout, suberect; stipe 40cm long, dark-
hairy in groove only, bearing small (not spine-like)
scales almost throughout, basal scales 4 x 1.5 mm,
not thick. Lamina 45 cm long; pinnae 12-15 pairs,
not opposite; texture very firm. Largest pinnae
15 x 3.cm; base broadly cuneate to rounded, with
one pair of lobes somewhat reduced; apex cau-
date-acuminate; edges lobed to 2mm from costa,
lobes on acroscopic side falcate near their tips
only, those on basiscopic side distinctly falcate
from their bases but not shorter; costules 5mm
apart; veins to 18 pairs, basal pair passing to sides
of the sinus-membrane which is decurrent be-
tween them; lower surface of rachis and costae
bearing scattered stiff brown hairs and also nar-
row scales with marginal setae, costules similar
but scales very small; upper surface of costae
brown-hairy near their bases only, rest of pinna
glabrous. Sori near costules, exindusiate; some
acicular hairs present on receptacle with the
sporangia; sporangia not setiferous, no glands
seen on them; spores dark, large, closely and
minutely papillose.
Distr. Malesia: West New Guinea (Bele
R.). Known only from the type and another spe-
cimen (BRASS 11327) from the same place.
Ecol. On limestone cliff in forest at 2200 m.
Note. COPELAND stated that small deciduous
setulose indusia were present; my observation of
the type is that the only hairs are on the recep-
tacle, as in several other species of this genus.
12. Plesioneuron wariense (COPEL.) HOLTTUM,
Blumea 22 (1975) 239.— Dryopteris wariensis
CoPEL. Philip. J. Sci. 6 (1911) Bot. 73. — Lastrea
wariensis COPEL. Gen. Fil. (1947) 146. — Thelyp-
teris wariensis (COPEL.) REED, Phytologia 17
(1968) 324. — Type: C. KING 101, p.p., N.E. New
Guinea, Waria River (MICH; NSW).
Caudex not seen, nor base of stipe. Lamina
60cm long; pinnae 8 pairs; basal pinnae with
stalks 4mm long, basal 4 pairs of lobes gradually
reduced, successive pinnae with shorter stalks and
fewer reduced basal lobes; apex of frond pinna-
like. Largest pinnae 18-30cm long, to 3.5cm
wide; aerophores not swollen; apex acuminate;
edges lobed to 0.5mm from costa, lobes slightly
falcate, about equal on the two sides of the costa;
costules to 5mm apart; veins 17-20 pairs, con-
colorous and very prominent on lower surface,
less so on upper, both basal veins passing to
margin above base of sinus, sinus-membrane very
404
FLORA MALESIANA
[ser. II, vol. 1°
short but distinct; lower surface of rachis distally
bearing many thick light brown hairs, rest of
rachis and costae with sparse short hairs, no
scales seen; upper surface of costae covered with
very short hairs and scattered hairs 1 mm long,
rest glabrous. Sori medial, rather large, exin-
dusiate; no hairs seen on receptacle; sporangia
bearing yellow glands, often 2; spores not seen.
Distr. Malesia: Papua New Guinea. Known
from two collections by C. KING (second is
n. 430).
Notes. A specimen of KING 101 at Bogor is P.
tuberculatum. The latter differs from P. wariense
in having sori near costules and small caducous
indusia. The sori of the type of P. wariense are
not quite mature.
13. Plesioneuron royenii HOLTTUM, Blumea 22
(1975) 240.— Type: VAN ROYEN 5469, W. New
Guinea, Waigeu Island (L).
Caudex short-creeping; stipe to 40cm long,
basal scales 61mm, setose. Lamina 22-27 cm
long; pinnae 6 pairs; basal pinnae with stalks
1.5-2 mm long, basal 2-3 pairs of lobes gradually
reduced, basal acroscopic lobe free, entire, 5mm
long; apex of frond almost pinna-like but widened
at its base with some transition to upper pinnae.
Largest pinnae 9-10cm long, sterile to 2.8cm
wide, fertile to 2.3cm; base subtruncate; apex
acuminate with cauda 10 mm long; edges lobed to
1.5mm from costa; lobes on both sides of costa
alike, oblique and slightly falcate; costules 3.5 mm
apart (fertile) to 4.5mm (sterile); veins 11-12
pairs, both basal veins passing to margin above
base of sinus; lower surface of rachis and costae
bearing abundant very short erect hairs, distally
on costae also a few longer hairs, few minute hairs
on costules, rest of surface glabrous and slightly
pustular; upper surface short-hairy on costae
only. Sori supramedial, exindusiate, without hairs
on receptacle; sporangia with 1-3 short setae:
spores not seen.
Distr. Malesia: West New Guinea (Waigeu I.),
only known from the type.
Ecol. In forest at 3 m altitude.
14. Plesioneuron savaiense (BAK.) HOLTTUM,
Blumea 22 (1975) 240.— Nephrodium savaiense
BAK. Ann. Bot. 5 (1891) 318.— Dryopteris
savaiensis (BAK.) C. CHR. Ind. Fil. (1905) 291;
Bishop Mus. Bull. 177 (1943) 82.— Thelypteris
Savaiensis (BAK.) REED, Phytologia 17 (1968)
312. — Type: POWELL 183, Samoa (K).
Dryopteris quadriaurita CHRIST, Philip. J. Sci.
2 (1907) Bot. 209.—Lastrea quadriaurita
(CHRIST) COPEL. Gen. Fil. (1947) 139: Fern
Fl. Philip. (1960) 326.— Thelypteris quadriaurita
(CHRIST) REED, Phytologia 17 (1968) 307.—
Type: COPELAND 1714, Mindanao, San Ramon
850 m (US; B, P).
Dryopteris ensipinna BRAUSE, Bot. Jahrb. 56
(1920) 84.— Thelypteris ensipinna (BRAUSE)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941)
251.—Type: LEDERMANN 12773, N.E. New
Guinea, Sepik Distr. 1400 m (B). — Fig. 9g-i.
Caudex short, creeping or suberect; stipe 20-
50cm long, + short-hairy at least distally, basal
scales to 12 1mm, firm, densely covered with
short hairs. Lamina 30-40 cm long; pinnae 12-15
pairs; basal pinnae sessile, basal 1-2 pairs of lobes
+ reduced (more on basiscopic than on acroscopic
side), basal acroscopic lobe free or nearly so,
sometimes with a toothed margin. Largest pinnae
10-12 x 2.0-2.5cm; apex acuminate, sometimes
with a short cauda; edges lobed to 1mm from
costa or more deeply, lobes on acroscopic side of
costa somewhat oblique, straight, those on basis-
copic side slightly falcate; costules 3.5-5 mm
apart; veins 10-12(—15) pairs, concolorous and
slightly prominent, basal pair both ending above
base of sinus; lower surface of rachis, costae and
costules + densely covered with short stiff hairs
of varying length to 0.5mm, some hairs also on
veins and on surface between them; upper surface
usually hairy on rachis and costae only. Sori
medial; indusia variable but always small, with
stiff marginal hairs, absent on some specimens
from New Guinea and the Moluccas; sporangia
bearing several rather long setae; spores dark,
spinulose.
Distr. Samoa, New Hebrides, and Malesia:
New Guinea, Philippines (Mindanao, Negros),
Moluccas (Batjan, Amboina).
Ecol. In New Guinea at 850-1500 m, in primary
or secondary forest.
Notes. One collection by NAKAIKE from
1500m in New Guinea has the largest pinnae
4.5 x 1.0cm and very narrow scales 10mm long. A
Philippine specimen has much less hairy scales
than any from New Guinea.
15. Plesioneuron altum (BRAUSE) HOLTTUM,
Blumea 22 (1975) 241. — Dryopteris alta BRAUSE,
Bot. Jahrb. 56 (1920) 86.—Lastrea alta
(BRAUSE) COPEL. Philip. J. Sci. 78 (1951) 429. —
Thelypteris alta (BRAUSE) REED, Phytologia 17
(1968) 259.— Type: LEDERMANN 11497, N.E.
New Guinea, Sepik Distr. 1300 m (B).
Caudex short-creeping, apical scales to 10x
1.5mm, rigid, bearing short hairs; stipe 70cm
long, glabrescent, scaly near base only. Lamina
70 cm long; pinnae 14 pairs; several pairs of lower
pinnae with stalks 3-4 mm long, only distal ones
sessile; basal acroscopic lobe of basal pinnae little
reduced, basal basiscopic lobe 12 mm long, apex
of frond not pinna-like. Largest pinnae 21x
3.3 cm; aerophores dark and slightly swollen; apex
caudate-acuminate, cauda to 2.5cm; edges lobed
to less than | mm from costa; lobes on acroscopic
side of costa almost straight, those on basiscopic
side rather strongly falcate but little shorter than
acroscopic lobes; costules to 6.5 mm apart; veins
1981]
THELYPTERIDACEAE (Holttum)
405
to 16 pairs, prominent both sides, basal ones not
meeting the distinct short-hairy sinus-membrane;
lower surface sparsely hairy on costae and cos-
tules, short hairs present on margins of lobes;
upper surface of rachis and costae densely hairy,
of costules sparsely, scattered short hairs present
on surface between veins. Sori near costules;
indusia very small, hairy; sporangia setiferous,
with acicular hairs on their stalks; spores dark,
finely spinulose.
Distr. Malesia: Papua New Guinea;
known from the type.
Note. BRAUSE stated that he saw no indusia,
but I observed them on the type. In frond-form
this is near P. tuberculatum, but the latter differs
in less deeply lobed pinnae, more distinct (though
caducous) indusia, and glandular sporangia.
only
16. Plesioneuron cystodioides HOLTTUM, sp. nov.
Pinnae usque 122.5 cm, subsessiles, fere ad
costam lobatae, lobis acroscopicis et basiscopicis
aequaliter leviter falcatis, lobis infimis acroscopi-
cis variabile elongatis; venae usque 15-jugatae;
rachis costaeque subtus pilis 0.1mm_longis
praedita; sori submarginales, indusia magna,
marginem superantia; sporangia nec setifera nec
glandulosa. — Type: T. NAKAIKE 50, N.E. New
Guinea, near Wau (K; TNS).
Caudex not known; stipe 45cm long, glabrous
except distally; basal scales to 9X 1mm, very
thick at base, thinner distally and narrowly
attenuate, sparsely hairy. Lamina to 62cm long;
pinnae 12-18 pairs, basal ones only subopposite;
basal pinnae sessile or nearly so, basal basiscopic
lobe a little reduced, basal acroscopic lobe elongate,
lobed, with pinnate veins in the lobes, to 2.5 x
0.5 cm; basal acroscopic lobe of suprabasal pinnae
sometimes + elongate. Largest pinnae 12 x 2.5 cm;
apex caudate-acuminate, cauda 2cm long, 1mm
wide; edges lobed almost to costa; lobes about equal
on the two sides of the costa, separated by rather
wide sinuses, slightly falcate (basiscopic lobes little
more so than acroscopic), edges cartilaginous and
slightly sinuous; costules 4-5 mm apart; veins to 15
pairs, basal ones passing to margin above base of
sinus, concolorous and prominent on both surfaces;
lower surface of rachis bearing erect hairs 0.1 mm
long with a few thicker brown ones, costal and
costular hairs a little longer, sparse hairs present on
veins and a few between veins; upper surface
of rachis and costae covered with erect hairs
0.2mm long, minute hairs also on costules. Sori
almost at the ends of the veins; indusia large, firm,
glabrous, overlapping margins of pinna-lobes;
no glands seen on sporangia; spores minutely
spinulose.
Distr. Malesia: Papua New Guinea, only
known from the type.
Note. The form of the pinna-lobes, with their
marginal sori, is very similar to that of pinna-lobes
of Cystodium sorbifolium.
17. Plesioneuron subterminale HOLTTUM, Blumea
22 (1975) 241.— Type: BRASS 32049, N.E. New
Guinea, Eastern Highlands, 1400 m (K).
Caudex suberect; scales to 10x 1 mm, thick at
base, narrowed and thinner distally, bearing short
hairs; stipe to 70cm long, glabrescent except in
groove. Lamina to 80cm long, very firm; pinnae
to c. 25 pairs; basal pinnae of type with stalks
2-3 mm long, of other specimens subsessile, basal
1-2 pairs of lobes somewhat reduced; aerophores
slightly elongate. Largest pinnae 20x 3.cm:; apex
acuminate, sometimes caudate; edges lobed to
1-1.5mm from costae, lobes on acroscopic side
slightly falcate distally, those on basiscopic side
falcate from their bases; costules to 5mm apart;
veins to 20 on basiscopic side of costules, to 18 on
acroscopic side, prominent both sides, basal
acroscopic veins usually ending near sinus-mem-
brane; lower surface of rachis densely covered
with erect hairs 0.3mm or more long, a variable
number being rather thick and brown, costal hairs
as rachis but antrorse distally, costular hairs more
sparse, sparse and variable on and between veins;
upper surface hairy about as lower, in some cases
more and longer brown hairs present on rachis.
Sori supramedial but not marginal; indusia firm,
with many short hairs; acicular hairs rare on
receptacle; sporangia sometimes with a small
gland; spores dark, minutely spinulose.
Distr. Malesia: Papua New Guinea (Eastern
Highlands and near Wau), in forest at 1200
2200 m.
18. Plesioneuron bipinnatum (COPEL.) HOLT-
TUM, Blumea 22 (1975) 242. — Dryopteris bipin-
nata COPEL. Philip. J. Sci. 9 (1911) Bot. 2.—
Lastrea bipinnata COPEL. Philip. J. Sci. 78 (1951)
422.— Thelypteris bipinnata (COPEL.) REED,
Phytologia 17 (1968) 264.— Type: C. KING 407,
E. New Guinea, Loane (MICH).
Caudex not known; stipe 32 cm long, closely and
minutely hairy; basal scales 5mm long, narrow,
hairy. Lamina 28cm long, texture rather thin;
pinnae 10-11 pairs. Basal pinnae largest, with
stalk 1mm long; basal pair of lobes free, short-
stalked, 34mm long, next pair of lobes almost
free and longer, rest of pinna lobed almost to
costa; apex acuminate with sinuous cauda 1.5cm
long; lobes slightly falcate, about equal on the two
sides of the costa; costules to 3 mm apart; veins to
11 pairs, concolorous and prominent both sides,
basal veins both passing to margin above base of
sinus except near apex of pinnae; lower surface of
rachis and costae bearing very short and scattered
longer erect hairs, hairs on costules similar, few
on veins; hairs on upper surface of costae as
lower but antrorse. Sori supramedial, impressed
(prominent on the upper surface); indusia small
with many short hairs; no glands or setae seen on
sporangia.
Distr. Malesia: Papua New Guinea. Known
406
only from the type and KING 221, from the
extreme east of New Guinea, opposite Samarai
island.
Note. These lowland coastal plants may
represent the same species as the larger ones from
mountains described here as P. subterminale.
19. Plesioneuron quadriquetrum (v.A.v.R.)
HOLTTUM, Blumea 22 (1975) 242. — Dryopteris
quadriquetra v.A.v.R. Nova Guinea 14 (1924) 16. —
Thelypteris quadriquetra (v.A.v.R.) CHING, Bull.
Fan Mem. Inst. Biol. Bot. 10 (1941) 254. — Type:
LAM 542, W. New Guinea, Mamberamo River, 10 m
(L).
Caudex suberect; stipe 20-40 cm long, glabrous
except for hairs in the groove, basal scales 6mm
long, narrow, hairy. Fronds subdimorphous,
sterile to 25cm long with 7-8 pairs of pinnae,
fertile with 10 pairs; basal pinnae with 2-3 pairs of
basal lobes reduced; apex of frond not pinna-like.
Sterile pinnae to 10 x 2.0cm; apex acuminate with
cauda 1.5cm long; edges lobed to 1mm from
costa; lobes slightly oblique, hardly falcate, about
equal on the two sides of the costa; costules 4mm
apart; veins 10-12 pairs, basal veins usually both
passing to margin above base of sinus; lower
surface of costa bearing rather sparse erect hairs
nearly | mm long and more abundant shorter ones,
short hairs also on costules; upper surface of
costae hairy as lower. Fertile pinnae to 8 x 1.2 cm;
costules 3-3.5 mm apart; lobes well separated; sori
supramedial; indusia firm, persistent, with short
brown hairs; sporangia sometimes bearing a small
colourless gland; spores with a translucent wing
and cross-wings.
Distr. Malesia: West New Guinea; only known
from the type.
Ecol. LAm’s label records the type as growing
epiphytically; confirmation of this unusual habitat
is desirable.
Note. The spores resemble those of P. arch-
boldiae (COPEL.) HOLTTUM from Fiji and P.
phanerophlebium (BAK.) HOLTTUM from the
Solomon Islands.
20. Plesioneuron falcatipinnulum (COPEL.)
HOLTTUM, Blumea 22 (1975) 244.— Dryopteris
falcatipinnula COPEL. Philip. J. Sci. 6 (1911) Bot.
74.— Lastrea falcatipinnula (COPEL.) COPEL.
Gen. Fil. (1947) 138; Philip. J. Sci. 78 (1951) 433. —
Thelypteris falcatipinnula (COPEL.) REED, Phy-
tologia 17 (1968) 276. — Type: C. KING 114, Eastern
New Guinea, in lowlands (MICH; BO).
Caudex not known; stipe 30cm long, glabrous;
basal scales 4mm long, very narrow, rather thin.
Lamina 30cm long, very firm; pinnae 8 pairs,
opposite, well-spaced, apical pair much reduced
and unequal. Largest pinnae 12x 1.5cm; aero-
phores swollen but not elongate; basal 1-2 pairs of
lobes reduced (except on distal pinnae); apex
acuminate with a cauda 10-15 mm long; edges
FLORA MALESIANA
{ser. II, vol. 1°
lobed to less than 0.5mm from costa; lobes on
both sides of costa oblique and much falcate;
costules 3.5mm apart; veins to 12 pairs, slender,
concolorous, prominent, both basal veins passing
to margin above base of sinus; lower surface of
rachis and costae bearing copious short stiff pale
hairs, similar hairs more sparse on costules; upper
surface of pinnae hairy only on costae. Sori in-
framedial; indusia firm, short-hairy; sporangia
bearing orange glands; spores not seen.
Distr. Malesia: Papua New Guinea,
known from the type.
only
21. Plesioneuron kostermansii HOLTTUM, sp. nov.
Stipes 45-50cm longus, omnino pilis erectis
usque | mm longis vestitus; lamina c. 50 cm longa;
pinnae 25-jugatae, usque 12 x 1.8 cm, lobis utrinque
falcatis ; venae 10-jugatae; rachis costaeque subtus
pilis erectis variis usque 1 mm longis dense vestitae,
venae subtus paginaque inter venas pilis multis
tenuibus erectis praeditae; pagina superior pin-
narum pilis adpressis 0.3mm longis vestita; sori
mediales, indusia parva, setosa; sporangia non
setifera. — Type: KOSTERMANS & SOEGENG142,
W. New Guinea, above Jayapura (L; BO).
Caudex short, erect, massive; stipe 45-50cm
long, covered with slender spreading hairs to
1 mm long, basal scales c. 7 x 0.5 mm, firm, glossy,
not thick. Lamina c. 50 cm long, rigid; pinnae c. 25
pairs; basal pair of lobes of basal pinnae reduced
and free. Largest pinnae 12X1.8cm, sessile;
aerophores not elongate; apex acuminate; edges
lobed to 1.5mm from costa, lobes equally falcate
on the two sides of the costa; costules 3 mm apart;
veins 10 pairs, basal pair both passing to margin
above base of sinus; lower surface of rachis and
costae densely covered with erect pale hairs of
varying length, to 1mm long, veins and surface
between them bearing erect slender hairs 0.5 mm
long; upper surface of costa bearing pale hairs
1mm long, whole surface of pinna covered with
appressed hairs 0.3 mm long. Sori medial or a little
inframedial; indusia rather small, firm, dark, with
stiff hairs 0.2mm long; sporangia not setose, no
glands seen on them.
Distr. Malesia: West New Guinea, only known
from the type.
Ecol. At 300m, “on slope of limestone hill,
very wet”.
22. Plesioneuron angiensis HOLTTUM, sp. nov.
Stipes 45 cm longus; lamina 55 cm longa; pin-
nae 24-jugatae, usque 12 X2.1 cm, sessiles; venae
12-13-jugatae; rachis costaeque subtus pilis
0.3 mm longis erectis vestitae; pagina superior in-
ter venas pilis brevibus erectis praedita; sori in-
framediales; indusia margine pilis brevibus
fimbriata; sporangia nec setis nec glandulis
praedita.—Type: KANEHIRA & HATUSIMA
13741, W. New Guinea, Arfak Mts, 1900 m (BO).
Caudex ‘60cm high, 1.5 cm diameter” (collec-
1981]
THELYPTERIDACEAE (Holttum)
407
tors’ label); stipe 45 cm long, glabrous apart from
brown hairs in the groove, basal scales not seen.
Lamina 55 cm long, thin but firm; pinnae 24 pairs;
lowest pinnae not well preserved; apex of frond
pinna-like but larger than pinnae. Largest pinnae
12x 2.1 cm, sessile; aerophores dark and slightly
swollen; base of pinna truncate; apex short-acu-
minate; edges lobed to | mm from costa or more
deeply; lobes falcate, about equally so on each
side of costa, basal lobes not or little reduced;
costules 44.5 mm apart, almost at right angles to
costa; veins 12-13 pairs, slender and prominent both
sides, basal acroscopic vein sometimes touching
side of the short sinus-membrane; lower surface
of costae densely covered with erect brown hairs
0.3 mm long with scattered longer ones, costal and
costular hairs similar but less dense and mostly
pale, no other hairs; upper surface of costae hairy
as lower, rest of upper surface bearing scattered
very short suberect hairs. Sori a little inframedial;
a few acicular hairs present on receptacle; indusia
rather small, dark, firm, fringed with stiff hairs
0.2 mm long; sporangia not setiferous, no glands
seen on them; spores not seen.
Distr. Malesia: West New Guinea (Arfak
Mts), only known from type.
Ecol. “Terrestrial in the poorly drained forest”
near the “male” lake, 1900 m alt.
23. Plesioneuron sandsii HOLTTUM, sp. nov.
Stipes 45cm longus, basi paleis 51mm
praeditus; lamina 50 cm longa; pinnae 25-jugatae,
inferiores 2-jugatae redactae; pinnae maximae
13.5 x 2.0 cm, lobis inferioribus 10—-12-jugatis sin-
gulatim ad costam adnatis; rachis costaeque sub-
tus pilis brunneis erectis vestitae, costae etiam
paleis angustis praeditae; sori inframediales; in-
dusia firma, glabra; sporangia glandulis minutis
praedita.— Type: M. J. S. SANDS 2380 A, New
Ireland, 2150 m (K).
Caudex short-creeping with stipes tufted at its
apex; stipe 45cm long, covered throughout with
stiff erect hairs 0.2-0.3 mm long, basal scales c.
5x 1mm, thick at base only, short-hairy, scales
above base of stipe thin and translucent. Lamina
50cm long, very firm; pinnae 25 pairs, basal 2-3
pairs irregularly reduced, lowest on one frond
3x 1.2cm, on another 7-8cm long, basal acros-
copic lobes of lower pinnae 7 mm long, basiscopic
3mm; apex of frond not pinna-like. Largest pin-
nae 13.5X2.0cm; apex acuminate, sometimes
with cauda 2 cm long; edges lobed almost to costa
in distal part of pinna, basal 10-12 pairs of lobes
separately adnate to costa; lobes on acroscopic
side almost all straight, to 12 mm long, on basis-
copic side strongly falcate, to 10mm long; cos-
tules to 5 mm apart; veins to 10 pairs, concolorous
and prominent on upper surface, pale and less
prominent on lower; lower surface of rachis den-
sely covered with brown erect hairs 0.3 mm long,
costae with shorter and less dense hairs, very
small brown scales also present on costae and
costules, rest of surface glabrous; upper surface
of costae covered with short brown hairs, rest
glabrous. Sori inframedial; indusia dark, firm,
glabrous; sporangia bearing very small elongate
glands (much shrivelled); spores black, closely
and minutely spinulose.
Distr. Malesia: Papua New Guinea
Ireland), only known from the type.
(New
24. Plesioneuron tuberculatum (CESATI) HOLT-
TUM, Blumea 22 (1975) 246.— Nephrodium
tuberculatum CESATI, Rendic. R. Acad. Napoli
16, fasc. 2 (1877) 26, 29. — Dryopteris tuberculata
(CESATI) C. CHR. Ind. Fil. (1905) 299: Dansk Bot.
Ark. 9, 3 (1937) 48, pl. V, f. 6.— Thelypteris
tuberculata (CESATI) CHING, Bull. Fan Mem.
Inst. Biol. Bot. 10 (1941) 255.— Type: BECCARI,
W. New Guinea, Arfak Mts (FI; K).
Dryopteris schlechteri BRAUSE, Bot. Jahrb. 49
(1912) 16, f. 1E, incl. var. djammuense. — Type:
SCHLECHTER 16188, N.E. New Guinea, near
Ketel, 200 m (B; K, L).
Dryopteris schizophylla v.A.v.R. Nova Guinea 14
(1924) 19. — Type: LAM 772, W. New Guinea (BO;
L). — Fig. 9d-f.
Caudex massive, erect; stipe to at least 75cm
long, glabrescent, basal scales to at least 7mm
long, narrow, thick. Lamina to 100cm long;
pinnae 20 pairs, all stalked except distal ones;
basal pinnae with stalks 3-5mm_ long, basal
basiscopic lobe much reduced, acroscopic lobe
not. Largest pinnae 25X5cm; aerophores +
elongate; apex caudate-acuminate, cauda 2.5-5 cm
long; edges lobed to 1-1.5mm from costa; lobes
on acroscopic side almost straight, on basiscopic
side somewhat falcate and sometimes shorter;
costules 5-7 mm apart, a small swelling at the base
of each, at least near bases of pinnae; veins to 23
pairs, slender, concolorous, prominent both sides,
basal acroscopic vein usually passing to side of
sinus-membrane; lower surface of rachis
glabrous, of costae and costules bearing sparse
brown hairs (often persistent only on costules and
appressed), surface between veins often strongly
pustular; upper surface bearing short dark hairs
on groove of rachis and on costae only. Sori near
costules; indusia small, glabrous, soon caducous;
sporangia bearing small red glands on body and a
short acicular hair on stalk; spores dark, minutely
spinulose.
Distr. Malesia: New Guinea and Moluccas
(Batjan: ALSTON 16977).
Ecol. In forest, low country to 1200m in New
Guinea, at 2000 m in Batjan.
Note. The Batjan specimen differs in more
deeply lobed pinnae, basal acroscopic lobe of
basal pinnae free, sori not so near costules, with
many acicular hairs on the receptacle.
25. Plesioneuron attenuatum (BRACK.) HOLT-
408
FLORA MALESIANA
(ser. Il, volate
TUM, Blumea 22 (1975) 245; Allertonia 1 (1977)
191, f. 4, k-m, q.v. for synonymy in the Pacific. —
Lastrea attenuata BRACK. in Wilkes, U.S. Expl.
Exp. 16 (1854) 193, t. 26, f. 2.— Aspidium
brackenridgei METT. Ann. Sci. Nat. Bot. IV, 15
(1861) 75, nom. nov. (not A. attenuatum Sw.).—
Thelypteris brackenridgei (METT.) REED, Phy-
tologia 17 (1968) 265.— Type: U.S. Expl. Exped.
Tahiti (US).
Caudex erect or suberect; stipe to 80cm long,
glabrous, basal scales to 20 mm long, very narrow,
rather thick and not glossy, with few hairs.
Lamina to 120cm long, very firm; pinnae 24 pairs
or more, lower ones with stalks 2-3 mm long, all
with basiscopic lobes strongly falcate and shorter
than the almost straight acroscopic ones: basal
acroscopic lobe on basal pinnae + reduced and
separately adnate to costa; aerophores elongate,
slender. Largest pinnae 25 x3.2cm; apex gradu-
ally attenuate; edges lobed to 1 mm from costa or
more deeply; acroscopic lobes to 2.2cm long,
narrowed to an acute tip, lobes on basiscopic side
to 1.6cm long; costules 4-5 mm apart; veins on
acroscopic lobes to 22 pairs, slender and prom-
inent both sides, basal acroscopic vein touching
side of sinus-membrane which is pellucid and
decurrent almost to the costa; lower surface of
rachis sparsely short-hairy, glabrescent, of costae
+ densely hairy at base, sparsely distally, with
hairs 0.3-0.5mm long, narrow scales sometimes
also present; upper surface hairy in groove of
rachis and on costae, hairs pale, 0.5 mm long. Sori
near costules; acicular hairs present with sporan-
gia on receptacle; indusia firm, persistent,
glabrous or variably short-hairy; sporangia bearing
2-3 small red glands; spores dark, minutely ver-
rucose.
Distr. Pacific, from Tahiti westwards to East
Malesia: New Guinea (New Ireland, New Bri-
tain).
Ecol. In New Ireland and New Britain, in
forest at 1100-1300 m, at lower altitudes in the
Solomon Islands.
26. Plesioneuron platylobum HOLTTUM, Blumea
22 (1975) 246.— Type: JERMY 3610, N.E. New
Guinea, Morobe Distr. (BM).
P. ctenolobum HOLTTUM, ibid. 247.— Type:
JERMY 3604, same locality (BM).
Caudex creeping, | cm diameter, bearing fronds
to 2cm apart; stipe 40-50 cm long, basal scales to
6x 1.5mm, thick, hairy. Lamina 60-80 cm long,
very firm; pinnae 20-30 pairs; basal pinnae with
stalks 1mm or less, their basal acroscopic lobes
free, 2cm long, 7-15 mm wide, with edges vari-
ably crenate or lobed and veins forked in the
lobes, basal basiscopic lobes + reduced, some-
times free but not wider than the next lobes.
Largest pinnae 18-21 x3.5-4.0cm; basal lobes
slightly reduced or not; apex acuminate and +
caudate; edges lobed to 1 mm from costa or more
deeply; lobes on acroscopic side almost straight,
on basiscopic side falcate but not shorter than
acroscopic lobes; costules 4-6mm apart; veins
18-22 pairs, basal acroscopic vein sometimes
touching sinus-membrane; lower surface of rachis
and costae variably short-hairy, few other hairs
present; upper surface hairy on rachis and costae,
some short hairs present on and between veins
distally on lobes. Sori medial; indusia firm, large,
with few to many short hairs; no acicular hairs on
receptacle; sporangia bearing glands or setae;
spores dark, minutely spinulose.
Distr. Malesia: Papua New Guinea,
known from the types.
Ecol. In Castanopsis forest at 1050 m.
only
27. Plesioneuron crassum (COPEL.) HOLTTUM,
Blumea 22 (1975) 243.— Dryopteris crassa
COPEL. Univ. Cal. Publ. Bot. 18 (1942) 220. —
Lastrea crassa COPEL. Gen. Fil. (1947) 138;
Philip. J. Sci. 78 (1951) 435, pl. 23. — Thelypteris
crassa (COPEL.) REED, Phytologia 17 (1968)
269. — Type: BRASS 10878, W. New Guinea, Lake
Habbema, 2650 m (MICH; L).
Caudex creeping, |1cm diameter; stipe 60cm
long, glabrous above base, basal scales to 5x
1.5mm, thin except at base. Lamina to 28cm
long, texture rigid; pinnae 9 pairs, 2 basal pairs
opposite, | pair basal lobes somewhat reduced on
basal pinnae only. Largest pinnae 11x2cm:;
aerophores elongate to 1mm; apex acuminate
with short cauda; edges lobed to 0.5mm from
costa; lobes on acroscopic side hardly falcate, on
basiscopic side distinctly but not strongly so,
margins (reflexed on drying) with a wide car-
tilaginous band; costules 4-4.5mm apart; veins
thick, 12 pairs, both basal veins passing to margin
above base of sinus; lower surface wholly
glabrous except for a few dark hairs on margins of
lobes; upper surface of rachis and costae bearing
stiff dark hairs. Sori near costules; indusia small,
dark, thick, glabrous; sporangia not setiferous, no
glands seen on them; spores of type not seen.
Distr. Malesia: West New Guinea. Known with
certainty from 2 collections from the type locality
(the second is BRASS 10934, from 2800 m); a spe-
cimen collected by C. B. KLOSS on the Carstensz
Mts at 2135 m may also represent this species.
28. Plesioneuron stenura HOLTTUM, sp. nov.
P. crasso affinis, differt: pinnis 18-jugatis, usque
23x 3.cm metientibus, lobis omnibus acutis, lobis
basiscopicis valde falcatis, lobo infimo acrosco-
pico pinnarum mediarum usque 2.5 cm elongato,
lobis
inferioribus utroque_ latere pinnarum
superiorum redactis; sporangiis glanduliferis,
receptaculo sori pilis acicularibus multis
praedito.— Type: L. E. CHEESMAN 152, S.E.
New Guinea, Mafulu, 1220 m (K).
Caudex not known; stipe glabrous, incomplete,
probably 40cm long, basal scales not seen.
1981]
THELYPTERIDACEAE (Holttum)
409
Lamina c. 70cm long; pinnae 18 pairs, very firm;
apex of frond widened at base and grading to
upper pinnae; basal pinnae almost sessile with
basal acroscopic lobe elongate and basiscopic lobe
a little reduced, both lobes reduced on upper
pinnae; aerophores slender, | mm long. Suprabasal
pinnae to 23cm long, 3cm wide above the base,
basal acroscopic lobe elongate (maximum 2.5 cm)
and parallel to rachis, basal basiscopic lobe
strongly falcate, 1.2 cm long; apex acuminate with
narrow cauda 4-5 cm long; edges lobed to 1.5 mm
from costa; acroscopic lobes straight, to 1.8cm
long, basiscopic lobes all strongly falcate and
shorter; costules 4.5-5 mm apart; veins to 23 pairs
on acroscopic lobes, slender, concolorous and
slightly prominent both sides, basal acroscopic
vein ending beside or near the sinus-membrane;
lower surface quite glabrous; upper surface of
rachis almost glabrous, of costae copiously hairy
on edges of groove, hairs 0.2 mm long. Sori close
to costules; abundant acicular hairs present on
receptacle with sporangia; indusia firm, dark,
glabrous; sporangia bearing several red-orange
glands.
Distr. Malesia: Papua New Guinea. Known
only from the type collection and a small sterile
frond from the same locality.
Note. This species is also close to P. attenua-
tum but has sessile basal pinnae and quite
glabrous lower surface; scales are needed for a
good comparison, those of P. attenuatum being
very distinctive.
29. Plesioneuron ophiura (COPEL.) HOLTTUM,
Blumea 22 (1975) 248.—Dryopteris ophiura
COPEL. Univ. Cal. Publ. Bot. 18 (1942) 220.—
Lastrea ophiura COPEL. Gen. Fil. (1947) 139;
Philip. J. Sci. 78 (1951) 434, pl. 22. — Thelypteris
ophiura (COPEL.) REED, Phytologia 17 (1968)
298.— Type: BRASS 12471, W. New Guinea,
Idenburg River, 2050m, on open _ rock-slide
(MICH; BO, L).
Description of the type: Caudex erect, apical
scales 5 1.5mm, acute, rigid, copiously short-
hairy; stipe 16cm long, minutely hairy. Lamina
20cm long, very firm; pinnae 7-9 pairs; basal
pinnae with stalks 2 mm long, basal lobes 1-2 mm
long; basal lobes of all other pinnae reduced; apex
of frond not pinna-like. Largest pinna 8 x 1.4cm;
aerophores slightly swollen; apex acuminate with
a cauda 1.5-2.0cm long; edges lobed to less than
1mm from costa; lobes slightly falcate on both
sides of the costa; costules 3.5-4 mm apart; veins
to 10 pairs, immersed (slightly prominent on the
lower surface), basal ones both passing to margin
above base of sinus; lower surface of rachis bear-
ing copious short pale stiff hairs, similar hairs less
copious on costae, also narrow scales, sparse hairs
on costules and veins; upper surface glabrous
apart from costa. Sori near costules; indusia thick,
dark, glabrous; sporangia not setiferous, no glands
seen on them; spores dark, minutely papillose.
Distr. Malesia: West New Guinea, at 2050 m.
Notes. BRASS 12465, from the same locality as
the type but “in young forest at foot of rock-slide,
a clump of fern with fronds to 180cm long”
(specimen at L) was named Lastrea costu-
lisora by COPELAND, and so accepted by me in
1975, but agrees with the type of P. ophiura in
deeply lobed pinnae, and probably represents the
fully developed form of the same species; its
description follows. Stipe 100cm long; basal
scales 10* 1.5mm. Lamina 100cm long; pinnae
22 pairs, lower ones 7 cm apart; basal pinnae with
stalks 3mm long and 3-4 pairs of basal lobes
gradually reduced and separately adnate to costa;
all upper pinnae with much-reduced basal lobes;
aerophores black, firm, more than 0.5mm long.
Largest pinnae 24 x 2.5-3.0cm, at base lobed al-
most to costa, distally to 1 mm from costa, apex
with cauda 3 cm long; lobes on acroscopic side of
pinna straight, on basiscopic side somewhat fal-
cate; costules 6mm apart; veins to 20 pairs;
sporangia sometimes with a small shrivelled gland,
rarely a short seta. If this is included in P. cos-
tulisorum, I think that the type of P. ophiura
should also be so included.
30. Plesioneuron costulisorum (COPEL.) HOLT-
TUM, Blumea 22 (1975) 246.— Dryopteris
basisora COPEL. Philip. J. Sci. 6 (1911) Bot. 73,
non CHRIST 1909. — Lastrea costulisora
CoPpEL. Gen. Fil. (1947) 138, nom. nov.—
Thelypteris costulisora (COPEL.) REED, Phy-
tologia 17 (1968) 269.— Type: C. KING 304, E.
New Guinea (MICH, BO, BRI, NSW).
Caudex lacking on type; stipe incomplete,
glabrescent. Lamina 60cm or more long; pinnae
15-20 pairs; basal pinnae with stalks to 2 mm long,
basal 2 pairs of lobes gradually reduced; aero-
phores not elongate. Largest pinnae 22 3cm;
apex acuminate; edges lobed to I-1.5mm from
costa; lobes on acroscopic side of costa straight,
on basiscopic side somewhat falcate; costules
5mm apart; veins commonly 18-20 pairs, basal
acroscopic vein ending beside or near sinus-
membrane; lower surface of rachis and costae
bearing coarse erect brown hairs, hairs on costules
similar but shorter, very short hairs present on
and between veins, a few narrow fringed scales
present on costae; upper surface of costae den-
sely brown-hairy throughout, rest of surface
glabrous. Sori near costules; indusia dark, firm,
with a few short hairs; acicular hairs present on
receptacle of sorus with the sporangia; sporangia
bearing an orange gland; spores of type not seen.
Distr. Malesia: Eastern New Guinea, in forest,
at altitudes to 1000 m.
Note. Other specimens which agree in frond-
form and,sori have differences in pubescence.
PULLEN 8417 from 250m in Milne Bay District
differs in sparse shorter hairs on the lower surface
410
of rachis and costae and lacks hairs between
veins. CROFT & LELEAN 68533, from a low al-
titude near the coast 93km S.E. of Lae, has
slender pale hairs 1 mm long on the lower surface
of costules and many short hairs between veins.
See also note above on P. ophiura.
31. Plesioneuron doctersii HOLTTUM, Blumea 22
(1975) 246.— Type: DOCTERS VAN LEEUWEN
10357, W. New Guinea, Rouffaer River, 300m
(BO; L).
Caudex short-creeping, 1.5cm diameter; stipe
25-40cm long, glabrescent, basal scales rather
thin, apparently c. 31mm. Lamina 40-50 cm
long, thick and rigid when dry; pinnae 15-18 pairs;
basal pinnae with stalks 3 mm long, only | pair of
basal lobes reduced; successive pinnae with shor-
ter stalks, only distal ones sessile; apex of frond
not pinna-like. Largest pinnae 9-11 cm long, 2.2-
2.6 cm wide; apex short-acuminate; edges lobed to
1.5—2.5 mm from costa; lobes separated by narrow
sinuses, basiscopic lobes more falcate and shorter
than acroscopic lobes; costules 4.5—S mm apart;
veins to 15 pairs, little prominent on either side,
basal pair both passing to sides of the sinus-
membrane; lower surface of rachis bearing rather
dense stiff erect hairs 0.1-0.2mm long, sparse
longer hairs present on costae, costules, sinus-
membranes and margins; upper surface of costae
covered with thick antrorse hairs 0.2-0.3 mm long,
minute hairs on costules. Sori near costules; in-
dusia dark, thick, with a few hairs; sporangia
sometimes bearing a small gland or a seta; spores
not seen.
Distr. Malesia: W. New Guinea, type locality
and Japen Island (L. E. CHEESMAN 1430).
Note. Specimens from Japen Island have some
setiferous sporangia, those from type locality only
glands.
32. Plesioneuron rigidilobum HOLTTUM, Blumea 22
(1975) 248. — Type: T.G. WALKER7741, N.E. New
Guinea, Edie Creek above Wau, on gold-mining
rubble (BM).
Caudex suberect; stipe to 75cm long, basal
scales 5-6mm long, narrow, thick. Lamina to
60 cm long, thick and rigid; pinnae 18 pairs; basal
pinnae with stalks 2mm long and free basal
acroscopic lobes 8mm long, basal basiscopic
lobes shorter, several pairs of successive pinnae
also stalked; basal lobes of middle and upper
pinnae not reduced; apex of frond almost pinna-
like. Largest pinnae of type 14x2.5cm; apex
acuminate, not or little caudate; edges lobed to
1mm from costa or more deeply; lobes on acros-
copic side straight and nearly at right angles to
costa, on basiscopic side slightly falcate and a
little shorter; costules 4.5-5 mm apart; veins to 15
pairs, basal veins both passing to margin above
base of sinus; lower surface of rachis and costae
bearing short pale hairs and thicker brown ones to
FLORA MALESIANA
(ser. II, vol.
0.5 mm long, hairs on costules fewer, not brown,
somewhat antrorse, a few very short hairs present
on veins and on surface between them; upper
surface of rachis and costae bearing many short
hairs. Sori medial or somewhat inframedial, not
touching costules; indusia large, firm, short-hairy;
sporangia bearing small glands and rarely a seta;
spores dark, minutely spinulose.
Distr. Malesia: Eastern New Guinea at 2000-
2500 m.
Note. It is possible that this is connected by
intermediates with P. ophiura and P. costu-
lisorum.
33. Plesioneuron wantotense (COPEL.) HOLTTUM,
Blumea 22 (1975) 245.— Dryopteris wantotensis
COPEL. Univ. Cal. Publ. Bot. 18 (1942) 220. —
Lastrea wantotensis COPEL. Gen. Fil. (1947) 140;
Philip. J. Sci. 78 (1951) 436, pl. 24. — Thelypteris
wantotensis (COPEL.) REED, Phytologia 17 (1968)
324. Type: CLEMENS 11013 bis, N.E. New
Guinea, Morobe Distr. 1200 m (MICH).
Caudex short, erect; stipe to 28cm _ long,
glabrous above base, basal scales small, acu-
minate, hairy. Lamina to at least 20cm long;
pinnae 10 pairs; basal pinnae sessile with 2 pairs
of basal basiscopic lobes reduced, basal one very
small; apex of frond pinna-like but broader than
pinnae. Largest complete pinna 5.5 x 1.2 cm (one
incomplete pinna 1.7cm wide); apex acuminate
with cauda 10mm long; edges lobed to 1.5mm
from costa; lobes on acroscopic side almost
straight, on basiscopic side shorter and falcate;
costules 3-4mm apart; veins 10-12 pairs, basal
acroscopic one ending beside sinus-membrane;
lower surface of rachis, costae, costules, sinus-
membrane and margins of lobes bearing short stiff
erect hairs; upper surface of rachis and costae
short-hairy, rest glabrous. Sori near costules; in-
dusia thick, dark, glabrous; sporangia bearing 2-3
red glands; spores not seen.
Distr. Malesia: Papua New Guinea,
known from the type.
Note. Apart from size, the type differs from P.
attenuatum (which is similar in frond-form) in its
much shorter basal stipe-scales.
only
34. Plesioneuron kundipense HOLTTUM, Blumea
22 (1975) 247.—Type: MILLAR & HOLTTUM
NGF 18593, N.E. New Guinea, Western High-
lands, 2150 m (LAE; BRI, K).
Caudex short, thick, creeping; stipe to 26cm
long, minutely hairy; basal scales thick at base,
thin distally, to 5x 1.5mm. Lamina of type 29cm
long; pinnae 14 pairs; basal pinnae with stalks not
over | mm long, aerophores slightly elongate, |
pair basal lobes slightly reduced; apex of frond
not pinna-like. Largest pinnae 8.5 x 2.0cm; apex
acuminate with short cauda; edges lobed to 1 mm
from costa; lobes slightly falcate on both sides of
costa; costules 3.5-4 mm apart; veins to 12 pairs,
1981]
THELYPTERIDACEAE (Holttum)
41]
prominent both sides, basal acroscopic one ending
near sinus-membrane; lower surface of rachis and
costae rather densely covered with stiff erect hairs
0.3-0.4 mm long, fewer hairs on costules, a few on
veins, rarely between veins; upper surface of
rachis and costae hairy as lower surface, scattered
shorter hairs present on surface between veins.
Sori medial or a little inframedial; indusia firm,
dark, with a variable number of very short hairs;
some sporangia bearing a short seta, glands not
seen; short acicular hairs present on receptacle or
on stalks of sporangia; spores not seen.
Distr. Malesia: Papua New Guinea (Western
Highlands, 2150 m). Known only from the type.
35. Plesioneuron dryas HOLTTUM, Blumea 22
(1975) 247. — Type: BRASS 23447, Eastern New
Guinea, Milne Bay Distr., Mt Dayman, in oak
forest 800 m (BM; L, LAE).
Caudex short, horizontal (teste BRASS); stipe to
60 cm long, glabrescent, basal scales c. 10 x 1 mm,
rigid, covered with short hairs, scales above base
thinner. Lamina 50cm long; pinnae 12 pairs,
several lower pairs opposite; basal pinnae with
stalks to 1mm long, basal acroscopic lobe free
and only slightly reduced, 3-4 pairs of basiscopic
lobes gradually reduced; basal lobes not reduced
on upper pinnae; apex of frond widened at base
with transition to pinnae. Largest pinnae 12 x 2cm
on type (14 x 2.5 cm on BRASS 28451); aerophores
slightly elongate; apex short-acuminate with
entire cauda 10-15 mm long; edges lobed to I-
1.5 mm from costa; lobes slightly falcate, those on
basiscopic side somewhat more oblique than those
on acroscopic side; costules 3.5-4.5mm_ apart;
veins 10-12 pairs, slender and prominent both
sides, basal acroscopic veins usually passing to
margin just above base of sinus; lower surface of
rachis and costae rather densely covered with
short erect hairs, sometimes with scattered longer
brown ones, sparse hairs on costules and veins
and between veins; upper surface of costae den-
sely short-hairy, sparse hairs on costules. Sori a
little inframedial; indusia dark, firm, with a few
short hairs; sporangia bearing red glands, some-
times several; spores not seen.
Distr. Malesia: Eastern New Guinea (2 collec-
tions) and Rossel Island.
36. Plesioneuron croftii HOLTTUM, sp. nov.
Stipes usque 36cm longus, basi paleis brevibus
vestitus; lamina 23cm longa; pinnae 5-jugatae,
suboppositae; pinnae infimae sessiles, aerophoris
tenuibus pallidis 1mm _ longis praeditae, lobis
infimis vix redactis; pinnae maximae 12.5 x
2.5cm, lobis vix falcatis utroque latere fere
aequalibus; venae 12—14-jugatae; sori prope cos-
tulas siti; indusia pilis 0.3mm longis vestita;
sporangia glandulis pallidis praedita.—Type: J.
R. CROFT 413, E. New Guinea, Natter Bay, 93 km
S.E. of Lae, 50m alt. (K; NSW).
Caudex short-creeping; stipe to 36cm long,
pale, sparsely short-hairy, basal scales not well
preserved, short and relatively broad at base,
thinner above base. Lamina 23 cm long; pinnae 5
pairs, subopposite; aerophores pale, slender,
hairy, to 1mm long; apex of frond pinna-like,
11cm long; basal lobes of basal pinnae hardly
reduced, acroscopic lobe not elongate; basal
basiscopic lobes of distal pinnae reduced. Largest
pinnae 12.5X2.5cm (on another frond 10.5 x
1.5cm); apex acuminate with cauda to 15 mm
long; edges lobed to 1-1.5mm from costa, lobes
almost equal on the two sides of the costa,
somewhat oblique, slightly falcate; costules 3-
3.5 mm apart; veins 12-14 pairs, slightly prominent
both sides, basal ones usually both passing to
margin above base of sinus; lower surface of
rachis bearing stiff erect hairs 0.2-0.5 mm long,
similar hairs on costae to 0.8mm, fewer hairs on
costules, slender erect hairs on surface between
veins; upper surface of rachis and costae hairy as
lower surface but hairs thicker, no other hairs.
Sori near costules; no acicular hairs on recep-
tacle; indusia medium brown, firm, not very thick,
with many hairs 0.3 mm long; sporangia not seti-
ferous, sometimes bearing small inconspicuous
pale glands; spores dark, minutely spinulose.
Distr. Malesia: Papua New Guinea; only
known from the type.
Note. The type was growing near larger plants
which are mentioned above under P. costuliso-
rum. It seems probable that P. croftii produces
only small plants; more information is needed.
Apart from size, it differs from the larger plants in
the presence of slender pale aerophores of a kind
I have not seen on other plants of this genus.
14. CYCLOGRAMMA
TAGAWA, Acta Phytotax. Geobot. 7 (1938) 52; CHING, Acta Phytotax.
Sinica 8 (1963) 316; HOLTTUM, Blumea 19 (1971) 28. — Glaphyropteris sect.
Cyclogramma H. ITo in Nakai & Honda, Nov. Fl. Jap. n. 4 (1939)
148. — Thelypteris subg. Cyclogramma K. Iwats. Mem. Coll. Sci. Univ.
Kyoto B, 31 (1964) 26. — Fig. 10a—c.
Caudex short to long-creeping; scales on stipe-base bearing acicular
hairs, sometimes hooked; fronds drying dark-olivaceous, bearing short
412 FLORA MALESIANA [ser. II, vol. 1
Fig. 10. Cyclogramma auriculata (J. SM.) CHING. a. Middle pinna, with an aerophore, x 1; b. venation,
sinus-membrane and sori, x3; c. sorus showing hooked hairs, x 9.— Pseudocyclosorus tylodes
(KUNZE) CHING. d. Basal normal pinna and reduced pinnae, x 1; e. two pinna-lobes showing venation
and sori, X 4 (a-c JACOBS 7299, d—e PRICE 387).
hooked hairs on lower surface of all axes and usually also on sporangia;
lower pinnae reduced or not; aerophores at bases of pinnae swollen or
elongate; pinnae deeply lobed; veins spreading at a wide angle to costule,
basal ones from adjacent costules running to edge close to, or just above,
sinus-membrane, never united; sori exindusiate; no glandular hairs on stalks
of sporangia; spores pale with translucent anastomosing wings.
Type species: Cyclogramma simulans (CHING) TAGAWA.
Distr. About 8 spp., Northern India to S. China, Taiwan; in Malesia: Philippines (Luzon).
Cytol. Base chromosome number probably 36: C. auriculata tetraploid; C. omeiensis octoploid
(n=c. 136; KURITA, J. Jap. Bot. 41, 1966, 176).
Note. A very distinct genus which has no obvious near allies. The species C. auriculata is one of
several ferns which are mainly distributed in Mainland Asia but have established themselves in
Northern Luzon.
1. Cyclogramma auriculata (J. SM.) CHING, Acta
Phytotax. Sinica 8 (1963) 317; HOLTTUM, Kalik-
asan 5 (1976) 110.— Polypodium subvillosum
Moore, Ind. Fil. (1861) 308, nom nud. — Poly-
podium auriculatum WALL. ex HOOK. Spec. Fil.
4 (1862) 237, non LINN., nec RADDI, nec PRESL;
BEDD. Ferns Br. India (1866) t. 203; CLARKE,
Trans. Linn. Soc. Bot. 1 (1880) 543. — Phegopteris
auriculata J. SM. Hist. Fil. (1875) 233, nom. nov.;
BEDD. Handb. (1883) 290, f. 149, excl. syn. Poly-
podium appendiculatum BEDD.— Dryopteris
auriculata (J. SM.) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 2 (1931) 196. — Dryopteris himalayensis
C. CHR. Ind. Fil. Suppl. III (1934) 88, nom. nov.
superfl. — Thelypteris subvillosa CHING, Bull.
Fan Mem. Inst. Biol. Bot. 6 (1936) 279.—C.
himalayensis TAGAWA, Acta Phytotax. Geobot. 7
(1938) 55.— Thelypteris auriculata (J. SM.) K.
1981]
THELYPTERIDACEAE (Holttum)
413
IWwATs. Acta Phytotax. Geobot. 19 (1961) 11;
Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 42. —
Type: WALLICH 314, Nepal (K).
Thelypteris simulans CHING, Bull. Fan Mem.
Inst. Biol. Bot. 6 (1936) 280.— Type: FAURIE s.n.
May 1914, Taiwan, Mt Arisan (S-PA).
Dryopteris squamaestipes sensu HAYATA, Icon.
Fl. Formosa 4 (1914) 179, f. 117, not Polypodium
appendiculatum BEDD. var. squamaestipes
CLARKE. — Fig. 10a-c.
Caudex erect, to c. 10cm tall. Stipe 5-10cm
long, densely covered with short pale hooked
hairs, at base broad brown scales with some
hooked hairs on them. Lamina to 100cm long;
pinnae more than 30 pairs, lower 10 pairs gradu-
ally reduced, a subabrupt transition between the
upper of these and largest pinnae, lowest pinnae
long. Largest pinnae seen on Philippine plants
10x 2cm (on Indian and Chinese plants to 20
2.5cm), lobed to 1-1.5 mm from costa, base trun-
cate, apex acuminate; lobes oblique, oblong with
rounded tips, entire; costules S-7 mm apart; veins
to 11 pairs; hairs on lower surface of rachis 1 mm
long, hooked, shorter hooked hairs on costae and
costules, marginal hairs not hooked; hairs on up-
per surface of costae 1mm long, not hooked;
short suberect hairs between veins, a few of them
hooked. Sori near costules; sporangia usually with
1 hooked hair.
Distr. N. India to S. China & Taiwan, in
Malesia: Philippines (N. Luzon; Mt Pulog, M.
JACOBS 7299 in L,K).
Ecol. Moist gully in mossy forest at 2600-
2700 m altitude.
3-4 mm long; aerophores at bases of pinnae 2mm
15. PPEUDOCYCLOSORUS
CHING, Acta Phytotax. Sinica 8 (1963) 322, excl. P. ciliatus (BENTH.) CHING
and P. caudipinna CHING; HoLttum, J. S. Afr. Bot. 40 (1974) 137.—
Pneumatopteris sensu HOLtTTUM, Blumea 19 (1971) 42, p.p. — Fig. 10d-e.
Caudex erect or short-creeping; scales broad, thin, bearing when young
marginal mucilage-secreting hairs; stipe scaly near base as caudex, scales
appressed. Fronds bearing many deeply-lobed pinnae, at base an abrupt
transition to much smaller ones; aerophores always swollen, in the sole
Malesian species elongate; veins free, basal acroscopic vein to base or side
of sinus-membrane, basal basiscopic vein to edge above base of sinus; short
capitate hairs often present on lower surfaces or on indusia, spherical
sessile glands lacking. Sori indusiate; no glands nor setae on sporangia; on
sporangium-stalk a hair of 2-3 cells, terminal cell largest but not spherical;
spores bearing many small subequal wings.
Type species: Pseudocyclosorus tylodes (KUNZE) CHING (by error as
xylodes).
Distr. About 11 spp. Tropical Africa, Madagascar, Mascarene Islands, S. India & Ceylon, N. India to
Japan; in Malesia: Philippines (Luzon).
Cytol. Chromosome numbers 35 and 36 both reported. P. tylodes, diploid (36, Ceylon and N. India); P.
esquirolii and P. repens, both diploid (35, N. & S. India); P. ochthodes, diploid (35 and 36 both reported from
India). There are erroneous records in LOVE, LOVE & PICHI SERMOLLI, Cytotaxonomical Atlas of
Pteridophyta, p. 221.
Notes. This genus differs from Pneumatopteris in having veins always free, lamina never pustular when
dry, sporangia lacking glands or setae on body. Because of the position of basal veins in relation to the
sinus-membrane CHING (1963) thought Dryopteris patens (Sw.)O. KTZE and D. normalis (SoD.) C. CHR. of
Tropical America should probably be included in Pseudocyclosorus, but in my judgement these species, and
some others allied to them in Africa, are better regarded as constituting a section of Christella (see
HoOLtTTuM, J. S. Afr. Bot. 40, 1974, 144). CLARKE (1880) united all species of Pseudocyclosorus in N. India
under the name Nephrodium prolixum, based on Aspidium prolixum WILLD., but WILLDENOW’s type is a
species of Christella from New Caledonia (HOLTTUM, Amer. Fern J. 63, 1973, 82).
Kew Bull. 34 (1979) 503.— Aspidium tylodes
KUNZE, Linnaea 24 (1851) 244, 283 (xylodes, err.
typ., p. 281). — Lastrea tylodes (KUNZE) MOORE,
1. Pseudocyclosorus tylodes (KUNZE) CHING,
Acta Phytotax. Sinica 8 (1963) 324: HOLTTUM, Brit.
Fern Gaz. 11 (1974) 55; HoLTTUM & GRIMES,
414
Ind. Fil. (1858) 107; BEDD. Ferns S. India (1863) t.
107; COPEL. Fern FI. Philip. (1960) 330. — Neph-
rodium ochthodes var. tylodes BEDD. Handb.
(1883) 240.—Nephrodium tylodes (KUNZE)
Hope, J. Bombay Nat. Hist. Soc. 14 (1903) 724. —
Dryopteris tylodes (KUNZE) CHRIST, Notul.
Syst. 1 (1909) 41; C. CHR. Contr. U. S. Nat. Herb.
26 (1931) 274.— Thelypteris tylodes (KUNZE)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 6 (1936)
296; HENNIPMAN, Blumea 16 (1968) 99, f. le. —
Syntypes cited: SCHMID-KOCH 11, 63, 89, 115,
Nilgiri Hills, S. India (none seen).
Dryopteris crassinervia C. CHR. Hedwigia 74
(1934) 229.—Type: G. STEIN 1179, Timor,
Ramelau (B; BM).
Nephrodium prolixum sensu BAK. Syn. Fil.
(1867) 268, var. tylodes tantum.— Dryopteris
luerssenii (non HARR.) CHRIST, Philip. J. Sci. 2
(1907) Bot. 208. — Fig. 10d-e.
Caudex erect. Stipe short, glabrous, flushed red,
covered at base with broad scales. Reduced pinnae
many pairs, represented by aerophores only.
Lamina 30-80 cm long, subcoriaceous, rigid when
dry; basal pinnae narrowed towards their bases;
aerophores elongate. Largest pinnae 12-20 x 1.2-
2.0cm; base subtruncate with basal acroscopic
lobe elongate; apex caudate-acuminate; edges
FLORA MALESIANA
[ser. II, vol. 1°
falcate acute tips; costules 3.5-4.5 mm apart; veins
10-12 pairs, pale and prominent on both surfaces;
lower surface entirely glabrous or some acicular
hairs present on distal parts of costae and on
sinus-membranes; upper surface hairy only on
costa. Sori inframedial; indusia large, firm,
glabrous; spores dark, finely spinulose.
Distr. India (except N.W.) and Ceylon, Burma
to S.W. China and Hong Kong; in Malesia: Phil-
ippines (N. Luzon) and Lesser Sunda Is. (Timor).
For a similar distribution, see VAN STEENIS, J.
Linn. Soc. Bot. 79 (1979) 127.
Ecol. In Luzon at 1700-2500m, usually on
banks of streams.
Notes. It is evident from KUNZE’s preliminary
list of species (1.c. 1851, p. 244) that he intended
the specific epithet to be tylodes (from the Greek
tylos, meaning a hump or knob, referring to the
aerophores); the name is also so written in the
description of Aspidium ochthodes where the two
species are compared. METTENIUS, HOOKER and
BEDDOME all adopted this spelling. It seems to
me certain that the spelling xylodes on p. 281 was
an error; it has no meaning. The paper was pub-
lished shortly after the death of KUNZE, who
perhaps did not correct a proof. I regard the use
of xylodes by later authors as an error, to be
lobed to 1.5-2 mm from costa, lobes entire with corrected.
16. PREUMATOPTERIS
NAKAI, Bot. Mag. Tokyo 47 (1933) 179; HOLTTUM, Blumea 19 (1973) 42,
excl. Pseudocyclosorus CHING; Allertonia 1 (1977) 226. — Thelypteris subg.
Pneumatopteris K. IWATS. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 33,
excl. sect. Macrocyclosorus. — Cyclosorus sensu CHING, Bull. Fan Mem.
Inst. Biol. Bot. 8 (1938) 230, p.p.; sensu HOLTTUM, Rev. FI. Mal. 2 (1955)
255, p.p.; Lastrea et Cyclosorus sensu COPEL. Gen. Fil. (1947) 135, 140,
p.p. — Fig. 11.
Caudex usually short, erect or decumbent, long-creeping in 2 spp.; stipe
never conspicuously hairy, basal scales in almost all cases broad, thin,
bearing few marginal acicular hairs, in some species also septate hairs
bearing mucilage-cells; fronds usually large with many pinnae, and in
almost all species a varied number of much-reduced basal pinnae, the
transition to these gradual or abrupt; lamina of reduced pinnae in most
species distinct, in the type and a few others greatly reduced; aerophores at
bases of reduced and lower normal pinnae + swollen, in a few cases much
elongate and then small aerophores also present at bases of costules; pinnae
always lobed, edges of lobes distinctly cartilaginous and in some species
toothed at the ends of veins; veins free in a few species which have deeply
lobed pinnae, in most species the basal veins at least anastomosing (in P.
costata small plants with fertile fronds may have free veins, later and larger
fronds having anastomosis); lamina between veins almost always + pustular
when dried (P. costata is an exception); lower surface of rachis, costae and
1981] THELYPTERIDACEAE (Holttum) 415
other parts never densely hairy, short capitate hairs present in some
species, minute colourless glands in a few, conspicuous sessile spherical
glands never present; sori usually covered with rather thin indusia;
sporangia often bearing short club-shaped glandular hairs (rarely setae) near
annulus and on the stalk a hair of 3-4 cells, the terminal cell enlarged but
usually colourless; spores in most cases light brown, bearing many very
small + quadrate wings, in a few species a + continuous longitudinal wing
with cross-wings.
Type species: Pneumatopteris callosa (BL.) NAKAI.
Distr. Tropics of the Old World, in the Pacific southwards to New Zealand; in all c. 80 spp., the majority
Malesian.
Ecol. Almost all species in forest, especially near streams, some in rocky stream-beds; a few species
confined to limestone.
Cytol. Base chromosome number 36; 7 Malesian spp., including P. costata, diploid (T. G. WALKER,
unpublished); P. truncata tetraploid in Ceylon and diploid in N. India and Sarawak; P. pennigera (G.
ForRST.) HOLTTUM tetraploid in New Zealand.
Taxon. NAKAI confined the genus to one species, stating ‘‘there is no other fern with which it can offer
any specific comparison’, but there are three species closely allied to the type in Malesia and four more in
the Pacific. The distinctive character of this group is the extreme reduction of basal rudimentary pinnae
which consist of little more than swollen aerophores which project through the layer of mucilage which
covers very young fronds (GOEBEL, Ann. Jard. Bot. Btzg 36, 1926, 84-96, with figures). TROLL described
the mucilage-bearing scales of another species (probably P. kerintjiensis) which he mis-named Dryopteris
sumatrana v.A.v.R. (Flora 128, 1933, 329-337); this species has shorter aerophores than P. callosa and
quite large reduced basal pinnae. The number, shape and size of such pinnae provide important characters
for distinguishing between species, and in some cases there is still not enough information about them.
Pneumatopteris is closely related to Sphaerostephanos but is distinct in the following characters:
presence of colourless short capitate hairs in place of spherical sessile glands; limited development of
acicular hairs apart from adaxial surface of rachis and costae; pustular appearance of lower surface
between veins of dried fronds; broad thin stipe-scales. There are exceptions to some of these, the most
notable in Malesia being P. costata and a few allies which do not have a pustular surface when dried. The
species with free veins are nearly all in New Guinea. They do not look like a primitive group and may be
derived from ancestors with anastomosing veins. P. truncata looks more like a prototype for the genus.
KEY TOMES PE CES
1. Veins anastomosing.
2. Stipe and rachis bearing many dark spines | mm long irregularly arranged.
3. Indusia lacking.
4. Pinnae to 25 x 1.6-2.7 cm with cauda 3 cm long; spores with many small wings . . 1.P. glabra
4. Pinnae to 12 x 2cm with cauda 1.5 cm; spores with wing and cross-wings . . . 2.P.sibelana
= Indusia present.
. Pinnae short-acuminate (Sumatra) . . . 22 eae ee 2 35Pidicranogramma
: Pinnae narrowly long-acuminate (Philippines) each “A 2A hh Ly ThA ehh elabra
2. Stipe and rachis lacking such spines.
6. Elongate aerophores present at bases of pinnae.
7. Reduced pinnae consisting of a prominent aerophore with a minute rim . . . . 4.P.callosa
7. Reduced pinnae with a distinct lamina.
8. Pinnae crenate; veins 6-7 pairs . . . . . . . . . . 5, P. subappendiculata
8. Pinnae lobed 1/3 towards costa; veins 10-16 pairs jt fons oO) eg 2 (63 Asuperba
6. Aerophores not elongate (in many cases slightly swollen).
9. Indusia lacking.
10. Pinnae commonly 12-15 x 1.5-2.0cm, thin; lower surface of rachis with or without minute
acicular hairs; sporangia with capitate hairs - . . . . 7.P.costata
10. Pinnae to 71cm, rigid when dry; lower surface of rachis densely covered with erect hairs
0.3 mm long; sporangia mostly bearing setae . . .... . .. =. . . 8.P.latisquamata
9. Indusia present.
416 FLORA MALESIANA (ser. II, vol
uae UM tye ea Seng
OC
f oy Sey ay = bt Ah
ARERAAReseINTSS
aR
\ rN
Fig. 11. Pneumatopteris callosa (BL.) NAKAI. a. Reduced basal pinnae, X 1; b. base of normal pinna,
x2; c. part of a fertile pinna showing pustular lamina, aerophore at base of costule and sori, x 8. — P.
truncata (POIR.) HOLTTuM. d. Basal normal pinna and a reduced pinna, x3; e. part of pinna, x 2; f.
showing veins joined along margin in lobes, x 3.— P. laevis (METT.) HOLTTUM. g. Reduced basal
pinnae, x 1.5; h. upper pinna, <P keysseriana (ROSENST.) HOLTTUM. i. Basal pinna, x}; j. one
pinna-lobe, x 3 (a~c W. L. CHEW et al. 1373, d—f MATTHEW s.n., g—h MATTHEW s.n., i-j cult. Kew).
1981] THELYPTERIDACEAE (Holttum) 417
11. Veins in fertile pinnae 2-5 pairs; pinnae in most cases not over 12 cm long.
12. Lower pinnae gradually reduced; stipe 5-15 cm long.
13. Basal pinnae auricled on acroscopic base.
14. Basal large pinnae narrowly cuneate at base on basiscopic side . . . 9. P. egenolfioides
14. Basal pinnae truncate at base on basiscopic side.
15. Brown hairs on rachis both sides; pinnae to8cmlong .. . . . .10.P. patentipinna
15. Lower surface of rachis glabrous; pinnae 3-3.5 cm long.
16. Pinnae lobed 2/5S—1/2 towards costa SoA Ac tissy Desay 11328 cheesmanise
16. Pinnae crenate Sia SW: Oe Ess sande! 442: Palithophila
13. Basal pinnae not auricled on ‘acroscopic ‘base.
17. Basal pinnae sages cuneate on Bo Se side; pinnae to 15cm long including cauda
45cm ae . . . .13.P. angusticaudata
17. Basal pinnae not SO; pinnae, if 1s cm long, with shorter cauda . . . .14.P. microloncha
12. Abrupt transition to reduced pinnae; stipe 20 to 50cm long ics te) a oe SR: brooks
11. Veins in fertile pinnae 7-10 pairs or more; pinnae usually much longer.
18. Basal pinnae not or variably a little reduced.
19. Pinnae 4cm or more wide, lobed less than 1/2 . . . ... . . . . . 16.P.inclusa
19: Pinnae to,2.5 cmiwide, lobed more than 1/2. 2)... 92. =) ea > @ sp’ -| 17.P_longipes
18. At least 1 pair basal pinnae conspicuously reduced.
20. Middle and upper pinnae very oblique, their bases asymmetric; apices of pinnae long and
very narrow.
21. Largest reduced pinnae and lower normal ones distinctly auricled at their bases 18. P. laevis
21. Lower pinnae narrowed towards their bases and not auricled.
22. Pinnae lobed 1/2 way to costa; c. 12 pairs of reduced pinnae, largest 3 cm long
19. P. obliqua
22. Pinnae lobed less deeply; reduced pinnae c. 6 pairs, largest 5 mm long
13. P. angusticaudata
20. Middle pinnae not very oblique, their bases symmetrical.
23. One pair basal pinnae reduced.
24. Reduced pinnae 2 cm long; normal pinnae lobed more than 1/2 . . . 20. P.basicurtata
24. Reduced pinnae | mm long; normal pinnae lobed less than 1/2 . 21. P. microauriculata
23. At least 2 pairs of abruptly reduced pinnae or several pairs gradually reduced.
25. Reduced pinnae and lower normal pinnae with nee serrate or lobed acroscopic auricles.
26. Reduced pinnae 24 pairs, subequal . . Ce fk. e222 PS eeallosa
26. Reduced pinnae to 8 pairs, decreasing downwards" : . . . . 23. P. auctipinna
25. Reduced pinnae and lower normal pinnae not thus auricled.
27. Hairs on upper surface of rachis thick, brown; some similar hairs usually present on lower
surface of rachis and costae.
28. Pinnae lobed less than 1/2; 13 pairs of veins anastomosing : . .24. P. jermyi
28. Pinnae lobed 1/2-2/3; basal veins meeting at base of sinus- -membrane_ . 25. P. papuana
27. Hairs on rachis and costae not thick and brown.
29. Lower pinnae gradually decrescent, at least on fertile fronds.
30. Indusia very small; lower surface bearing minute glands between veins’. 7. P. costata
30. Indusia conspicuous; lower surface lacking such glands . . . . 26. P. rodigasiana
29. Transition from normal to reduced pinnae abrupt.
31. Pinnae lobed 2/3 or more deeply.
32. Pinnae lobed to 1.5mm from costa, not pustular between veins when dried
27. P. japenensis
32. Pinnae lobed less deeply, pustular between veins.
33. Pinna-lobes dentate at ends of veins cova rahe 4.9 43. ae Atobaica
33. Pinna-lobes entire.
34. Pinnae lobed to 2.5 mm from costa; hairs present on all parts of lower surface
29. P. incisa
34. Pinnae lobed to 4mm from costa; lower surfaces quite glabrous 30. P. psilophylla
31. Pinnae lobed at most 3/5 towards costa.
35. Lowest pinnae considerably narrowed towards their bases, gradually or rather ab-
ruptly.
36. Pinnae of well-grown plants lobed more than 1/2.
37. Stipe-scales narrow, spreading ...... . . . . . .31.P.pergamacea
37. Stipe-scales thin, appressed.
38. Pinnae commonly 2.5—3 cm wide; sporangia lacking glands . . 32. P.sogerensis
418 FLORA MALESIANA (ser. II, vol. 1°
38. Pinnae not over 2cm wide; sporangia bearing glands . . . 33. P.micropaleata
36. Pinnae of well-grown plants lobed less than 1/2.
39. Pinnae commonly 20 x 2.5 cm, usually with some hairs on lower surface of costae,
often also between veins. . . . . 34. P. truncata
39. Pinnae not more than 2 cm wide; lower ‘surface quite glabrous.
40. Pinna-lobes almost at right angles to costa; sporangia bearing glands 35. P. lawakii
40. Pinna-lobes oblique; no glands on sporangia . . . . . . . .36.P. michaelis
35. Lowest pinnae not or little narrowed towards their bases.
41. Upper reduced pinnae longer than wide, with dilated bases and lobed distally; lower
surface slightly pustular a ee . . . . 37. P.nitidula
. Upper reduced pinnae as long as wide, not dilated at base nor lobed above base,
strongly pustular between veins . .. . . . . . . . . .38.P. kerintjiensis
1. Veins free.
42. Largest pinnae less than 2 cm long, entire or slightly crenate; many pairs of lower pinnae gradually
decrescent. . . . . . . 39. P. nephrolepioides
42. Largest pinnae in most cases ; larger: lower pinnae otherwise.
43. Both surfaces of pinnae bearing short erect hairs throughout.
44. Lower 1-3 pairs of pinnae progressively smaller, lowest ame Icmlong . 40. P. ligulata
44. Lower 5-6 pairs of pinnae less than 2 mm long : . . . . . 41. P. finisterrae
43. No hairs between veins on upper surface, few or none on lower.
45. Four or more pairs of lower pinnae gradually reduced; pinnae lobed c. half-way to costa
7. P. costata
45. Not this combination of characters.
46. Pinnae 2-3 cm long.
47. Basal pinnae shorter than the next pair ‘ vi el os se 6 oS ea2 ae aversteeshn
47. Six pairs of lower pinnae + gradually reduced.
48. Basal 3-4 pairs of pinna-lobes separately adnate to costa . . . . . 43.P.sumbawensis
48. Pinnae lobed half-way to costa. . . . .. 2. © . . . ©... “S33P) petropinla
46. Pinnae much longer.
49. Much-reduced basal pinnae 0-3 pairs.
50. Sori supramedial; pinnae lobed to 1-2 mm from costa; basal veins from adjacent costules often
meeting, or nearly so, at the sinus.
51. Pinnae to 23 x 4 cm; basal acroscopic lobe of middle pinnae notalmostfree 44. P. keysseriana
51. Pinnae c. 8 X 1.5 cm; basal acroscopic lobe of middle pinnae almost free . 45. P.caudata
50. Sori medial; pinnae lobed to less than 1 mm from costa; basal veins from adjacent costules
ending far apart at the margin.
52. Basal basiscopic lobes of middle pinnae equal to acroscopic lobes . . . 46. P. deficiens
52. Basal basiscopic lobes of middle pinnae shorter than acroscopic.
53. Basal acroscopic lobes of largest reduced pinnae and lowest normal pinnae elongate.
54. Much-reduced basal pinnae 2 pairs; hairs present on lower surface of rachis and costae
47. P. mingendensis
54. Much-reduced pinnae lacking; lower surfaces quite glabrous . . . . 48. P. eburnea
53. Basal’acroscopic lobes of such pinnae reduced! => ~ > = =” = .. : 2). Wes495Reexcisa
49. Much-reduced basal pinnae at least 6 pairs.
55. Pinnae to 15cm or more long.
56. Reduced pinnae 6 pairs, lowest | cm long.
57. Pinnae lobed to 1 mm from costa.
58. Sori inframedial; dark brown hairs present on upper surface of costa . . 50. P. regis
58. Sori supramedial; no dark brown hairs . ... . . . . . . . 44, P. keysseriana
57. Pinnae lobed to 3.5-4mm from costa . . Je Be eS IPSiboridensis
56. Reduced pinnae many pairs, lower ones smaller ow SSS S30 pe 2525 Rsimbricata
55. Pinnae not or little more than 10 cm long.
59. Pinnae lobed half-way to costa ee Ge ate oo M.cost Be a. al SESS SPspetrophila
59. Pinnae lobed almost to costa BW) esr», 412) | hs he rine 2 re SA aI KerI
1. Pneumatopteris glabra (COPEL.) HOLTTUM, Cyclosorus viridis COPEL. Philip. J. Sci. 81
Blumea 21 (1973) 301.—Cyclosorus glaber (1952) 35, pl. 24; Fern Fl. Philip. (1960) 364. —
CopPEL. Philip. J. Sci. 81 (1952) 34; Fern Fl. Thelypteris viridis (COPEL.) REED, Phytologia 17
Philip. (1960) 363. — Thelypteris sevillana REED, (1968) 324. — Type: RAMOS & EDANO BS 37490,
Phytologia 17 (1968) 313.—Type: RAMOS BS- Mt Masigit, Luzon (MICH; US, BO).
42976, Sevilla River, Bohol (UC; BM). Caudex short, creeping. Stipe 5cm long, basal
1981]
THELYPTERIDACEAE (Holttum)
419
scales small, thin, soon disappearing; base of stipe
to first large pinna 25-100cm; stipe and rachis
bearing many dark spines, erect or curved, | mm
long. Reduced pinnae many pairs, c. 3cm apart,
each consisting of an aerophore 1-2 mm long,
upper ones with a lamina 2-3mm long almost
encircling the aerophore; transition to normal
pinnae abrupt or subabrupt; lamina excluding
reduced pinnae to 120cm long (sometimes fertile
at a length of 30cm), basal pinnae narrowed near
their bases. Largest pinnae to 25cm long, 1.6—
2.2cm wide; base broadly cuneate to full width
(asymmetric On upper pinnae); apex caudate-
acuminate (cauda to 3cm long, finely toothed
throughout); edges lobed 1/3 towards costa or a
little more deeply, lobes entire with acute falcate
tips; costules 4-6 mm apart with a small aerophore
at the base of each; veins 8-10 pairs, basal pair
anastomosing, next 2 pairs passing to sides of a
long sinus-membrane; lower surface of rachis
glabrous, of costae and costules bearing a few
small capitate hairs, surface between veins rather
strongly pustular; upper surface of rachis bearing
+ abundant thick brown hairs 0.5-1 mm _ long,
similar hairs near bases of costae, rest of upper
surface glabrous. Sori medial on distal veins,
variably supramedial on lower ones; small firm
indusia sometimes present; sporangia bearing
rather large colourless club-shaped glands.
Distr. Malesia: Philippines (Luzon, Samar,
Bohol, Mindanao).
Ecol. Usually on rocky stream-banks, to 400-
1000 m, also found in ridge forest.
Note. COPELAND distinguished Cyclosorus
viridis by the green (not dark) colour of dried
fronds, narrower pinnae and the presence of small
indusia. All these characters appear to be variable.
Very young fronds of plants in cultivation at Kew
develop a coat of mucilage, as in P. callosa. Small
indusia, not as large as a mature sporangium, are
found in some sori of these plants. RAMOS &
EDANO, BS 49586, from Mindanao (collected in
1927) was distributed as Dryopteris todayensis
CHRIST?.
2. Pneumatopteris sibelana HOLTTUM, Blumea 21
(1973) 301.—Type: ALSTON 16942, Batjan, N.
slope of Mt Sibela, 1500 m (BM).
Stipe less than 5 cm long; base of stipe to first
large pinna 45 cm; stipe and rachis bearing slender
spines 1mm long. Reduced pinnae 10-12 pairs,
each consisting of a semicircular lamina 2-3 mm
wide and an aerophore | mm or more long; lamina
exclusive of reduced pinnae 65-70 cm long; pinnae
c. 28 pairs, basal ones slightly narrowed at their
bases. Largest pinnae 12cm long, 1.8-2.0cm
wide; base subtruncate; apex acuminate with a
finely-toothed cauda 10-15 mm long; edges lobed
c. 1/3 towards costa, lobes entire, falcate,
subacute; costules 5-5.5 mm apart; veins 10 pairs,
basal 1-12 pairs anastomosing, next 2) pairs to
sides of a long sinus-membrane; on lower surface
of rachis short curved brown hairs, sparse minute
hairs on costae, rest of surface glabrous; upper
surface glabrous apart from rachis and costae.
Sori medial, lower ones not or little divergent,
exindusiate; no glands seen on sporangia; spores
light brown with a + continuous translucent wing
and cross-wings.
Distr. Malesia: Moluccas (Batjan), only known
from the type.
3. Pneumatopteris dicranogramma _ (v.A.v.R.)
HOLTTUM, Blumea 21 (1973) 301.— Dryopteris
dicranogramma v.A.v.R. Bull. Jard. Bot. Btzg III,
5 (1922) 202.— Thelypteris dicranogramma
(v.A.v.R.) REED, Phytologia 17 (1968) 272.—
Type: BUNNEMEIJER 10454, Sumatra, G. Kerinci
(BO;-L):
Stipe very short; base of stipe to first large
pinna 45-55cm; stipe and lower part of rachis
bearing short dark spines. Reduced pinnae c. 2.5-
3cm apart, each consisting of a narrow lamina
almost encircling the base of an aerophore.
Lamina excluding reduced pinnae 50-70 cm long,
coriaceous; basal pinnae, several pairs, narrowed
at their bases. Largest pinnae 17.5 x 1.8 cm; base
truncate; apex short-acuminate, serrate to tip;
edges lobed 1/3-1/2 towards costa, lobes falcate,
acute; costules c. 4mm apart, each with a small
aerophore at its base; veins 10-12 pairs, thick,
basal pair anastomosing, then several pairs ending
beside a long sinus-membrane; lower surface of
rachis bearing short stiff brown hairs, a few
similar hairs sometimes on bases of costae, rest
glabrous; upper surface of rachis covered with
brown hairs, of costae glabrous. Sori in an in-
verted V (basal ones divergent); indusia soon
broken, probably hairy when young.
Distr. Malesia: Sumatra (G. Kerinci, 2 collec-
tions at 2100-2400 m; G. Dempo).
Note. The specimen from Malaya named
Cyclosorus dicranogramma in HOLTTUM, Rev.
Fl. Mal. 2 (1955) 267 is P. callosa.
4. Pneumatopteris callosa (BL.) NAKAI, Bot. Mag.
Tokyo 47 (1933) 179; HoL_LTruM, Blumea 21
(1973) 302. — Aspidium callosum BL. En. Pl. Jav.
(1828) 152.— Nephrodium callosum (BL.) KEYS.
Pol. Cyath. Herb. Bung. (1873) 48; RAciB. FI.
Btzg 1 (1898) 192.— Dryopteris callosa (BL.) C.
CHR. Ind. Fil. (1905) 256; v.A.v.R. Handb. (1908)
220; BACKER & POSsTH. Varenfl. Java (1939)
62.— Cyclosorus callosus (BL.) CHING, Bull.
Fan Mem. Inst. Biol. Bot. 8 (1938) 205.—
Thelypteris callosa (BL.) K. IWATS. Mem. Coll.
Sci. Univ. Kyoto B, 31 (1964) 34.— Type:
BLUME, Java (L, n. 908, 339-365).
Goniopteris lobbiana FEE, Gen. Fil. (1852)
251.— Type: Loss, Java (orig.?; isotypes BM,
K).
Aspidium multijugum CHRIST, Ann. Jard. Bot.
420
FLORA MALESIANA
[ser. II, vol. 1°
Btzg 15 (1898) 135. — Lectotype (HOLTTUM 1973):
SARASIN 1323, Celebes (BAS). — Fig. 1la-c.
Caudex stout, erect; apex and fronds in their
early stages covered with mucilage. Stipe very
short; base of stipe to first large pinna 60-100 cm.
Reduced pinnae 2-4 cm apart, each consisting of a
prominent aerophore with a narrow rim at its
base; lamina excluding reduced pinnae 100-
200 cm long, texture very firm; 2-3 pairs of lower
pinnae narrowed near their bases. Largest pinnae
commonly 15-25cm long, 2.0-2.5em wide; base
truncate to broadly cuneate; apex rather evenly
attenuate; edges lobed 1/4-1/3 towards costa,
lobes entire, falcate, subacute, with conspicuous
pale cartilaginous edge; costules 3.5—4.5(-5)mm
apart, at a wide angle to costa, a small aerophore
at the base of each; veins c. 12 pairs, slender and
prominent both sides, basal 1-15 pairs anastomos-
ing, 3-4 pairs passing to sides of the prominent
sinus-membrane; lower surface of rachis and cos-
tae glabrous or with some pale acicular hairs,
short hairs present on sinus-membrane and edges,
lamina between veins finely and copiously pus-
tular; upper surface of rachis (at least distally)
bearing long curved brown hairs, a few such hairs
near bases of costae, rest of surface glabrous. Sori
in an inverted V; indusia firm, glabrous; sporangia
bearing club-shaped glands, stalks of sporangia
bearing hairs with an end-cell which (on young
sporangia) is swollen and orange; spores with
many small wings.
Distr. Malesia: West Malesia, Celebes, Lesser
Sunda Is. (Flores) and Moluccas (Ceram).
Ecol. Near streams in forest, at 600-1750 m.
Notes. BLUME distinguished var. B. with more
rigid fronds and sori confluent; I have not seen the
type, and consider the distinctions not good.
v.A.v.R. described var. sumatrana (Handb. 220)
from a small plant with gradually reduced lower
pinnae; it might represent another species (type
not found). CHRIST gave a brief description of the
SARASIN specimen from Celebes cited under
Aspidium multijugum, referring to Nephrodium
multijugum BAK. Syn. Fil. (1867) 291 and ascrib-
ing the name to WALLICH, but WALLICH in-
cluded specimens of three different species under
his n. 348 and it is not clear to which of them
BAKER’s description applies, though certainly not
to P. callosa. I have not seen the WARBURG
specimens also cited by CHRIST.
5. Pneumatopteris subappendiculata (COPEL.)
HOLTTuM, Blumea 21 (1973) 303.— Dryopteris
subappendiculata COPEL. Univ. Cal. Publ. Bot.
18 (1942) 220.—Cyclosorus subappendiculatus
(COPEL.) COPEL. Gen. Fil. (1947) 143; Philip. J.
Sci. 78 (1951) 449, pl. 30.— Thelypteris subap-
pendiculata (COPEL.) REED, Phytologia 17 (1968)
317.— Type: BRASS 12247, New Guinea, Iden-
burg River (MICH; L).
Stipe 10cm long, covered with short brown
hairs and when young thin scales; base of stipe to
first large pinna 35 cm. Reduced pinnae to 7 pairs,
lowest 5mm long, uppermost 2 cm long, broadly
triangular with auricled acroscopic base. Lamina
excluding reduced pinnae to 90 cm long; pinnae c.
16 pairs, opposite, with conical aerophores at their
bases; basal pinnae auricled on acroscopic side or
+ dilated both sides at base; texture firm. Largest
pinnae 18X3cm (sterile), fertile to 2.5cm wide;
base truncate to subcordate; apex short-acu-
minate; edges crenately lobed, sterile to a depth of
2mm, fertile to 3mm; costules 5-6mm apart;
veins 6-7 pairs, slender, prominent both sides,
basal 15-2 pairs anastomosing, next 1-2 pairs to
sides of sinus-membrane which is not prominent
on lower surface; lower surface of rachis bearing
rather sparse thick brown hairs, costae, costules
and sinus-membrane sparse slender short hairs,
surface between veins slightly pustular; upper
surface of rachis covered with brown hairs, those
on costae paler and more slender. Sori medial
(distal ones inframedial); indusia thin, pale, with a
few short hairs; sporangia lacking glands.
Distr. Malesia: Eastern New Guinea, at 1400-
2200 m.
Note. It is not clear whether this is sharply
distinct from P. superba.
6. Pneumatopteris superba (BRAUSE) HOLTTUM,
Blumea 21 (1973) 303.— Dryopteris superba
BRAUSE, Bot. Jahrb. 56 (1920) 105. — Cyclosorus
superbus (BRAUSE) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 249; COPEL. Philip. J. Sci. 78
(1951) 447.— Thelypteris superba (BRAUSE)
REED, Phytologia 17 (1968) 318.— Type:
LEDERMANN 11733, N.E. New Guinea, Sepik
Distr., Schraderberg 2070 m (B).
Dryopteris deltiptera COPEL. Univ. Cal. Publ.
Bot. 18 (1942) 142.—Cyclosorus deltipterus
CoPEL. Gen. Fil. (1947) 142; Philip. J. Sci. 78
(1951) 449, pl. 29.—Thelypteris deltiptera
(COPEL.) REED, Phytologia 17 (1968) 271.—
Type: BRASS 11260, New Guinea, near Lake
Habbema 2200 m (GH).
Differs from P. subappendiculata: caudex
short, erect; stipes of largest fronds bearing a few
spines less than 0.5 mm long; fronds larger; pinnae
to 23cm long, + dilated at base which in largest
fronds is 3-4cm_ wide, asymmetric (wider
throughout on basiscopic side of costa than on
acroscopic side), edges lobed up to 1/3 towards
costa, more deeply on basiscopic than on acros-
copic side; lobes with broad undulate cartilaginous
edges; costules 6-7 mm apart; veins 10-16 pairs;
basal sori not much divergent.
Distr. Malesia: Eastern New Guinea, many
collections; 2 doubtful from W. New Guinea.
Ecol. In forest at 1700-2700 m.
7. Pneumatopteris costata (BRACK.) HOLTTUM,
Blumea 21 (1973) 305; Allertonia 1 (1977) 229, f.
1981]
THELYPTERIDACEAE (Holttum)
421
9b. — Goniopteris costata BRACK. in Wilkes,
U.S. Expl. Exp. (1854) 28.— Dryopteris costata
(BRACK.) MAXON, Univ. Cal. Publ. Bot. 12 (1924)
26. — Thelypteris costata (BRACK.) REED, Phy-
tologia 17 (1968) 269. — Type: U.S. Expl. Exped.,
Fiji (US).
Lastrea cavitensis COPEL. Philip. J. Sci. 81
(1952) 26; Fern Fl. Philip. (1960) 328.— Thelyp-
teris cavitensis (COPEL.) REED, Phytologia 17
(1968) 267. — Type: MANGUBAT BS 1302, Luzon,
Cavite (MICH).
Dryopteris pennigera (ForST. f.) C. CHR.
subsp., C. CHR. Ind. Fil. (1905) 289. — Dryopteris
luzonica CHRIST, Philip. J. Sci. 2 (1907) Bot. 196,
quoad Mangubat 1302 tantum.
Caudex short, erect; stipe 5-10cm long, basal
scales broad, thin, imbricating. Lamina to 80cm
long; pinnae to more than 25 pairs, basal 4-8 pairs
gradually decrescent, lowest 3-12mm_ long;
reduced pinnae not auricled nor narrowed at their
bases; texture thin but firm. Largest pinnae to
14x 1.5 cm (type of L. cavitensis 8 x 0.8 cm); base
broadly cuneate to subtruncate; apex narrowly
acuminate; edges lobed c. 3 way to costa; costules
to 4mm apart (2.5mm in type of L. cavitensis);
veins in well-grown plants 8-9 pairs, basal veins
anastomosing, next acroscopic vein or pair to
short sinus-membrane (in type of L. cavitensis 4-6
pairs of veins, basal ones meeting without joining
at sinus-membrane); lower surface of rachis and
costae bearing many minute acicular hairs, such
hairs sparse on costules and veins, on and be-
tween veins minute sessile or subsessile glands
which collapse on drying; upper surface of rachis
and costae covered with pale hairs more than
0.5mm long, between veins minute glands as
lower surface. Sori inframedial, exindusiate or
sometimes with very small indusia; sporangia
bearing small capitate hairs; spores pale, with a +
continuous thin translucent wing and cross-wings.
Chromosomes: n= 36 (T. G. WALKER).
Distr. Queensland, Solomon Islands, New
Hebrides, New Caledonia, eastwards to Tahiti;
var. hispida HOLTTUM in Cook Islands, Pitcairn,
Easter Island (see HOLTTUM in Allertonia 1977),
in Malesia: N.E. New Guinea, Philippines (S.
Luzon), and Lesser Sunda Is. (Bali, Sumbawa).
Ecol. In Malesia at 250-1200m, on sloping
ground near streams, apparently not in deep
shade.
Note. This species has been much misinter-
preted; the first author after BRACKENRIDGE to
characterize it clearly was MAXON (1924) but he
did not see the characteristic minute glands. The
fern so named by HOOKER (Spec. Fil. 5, 1864, 7)
is Chingia longissima (BRACK.) HOLTTUM.
DIELS and CHRISTENSEN confused P. costata
with P. pennigera (FORST. f.) HOLTTUM of New
Zealand and S.E. Australia; some specimens from
Samoa were named Dryopteris transversaria and
D. christopherseni by CHRISTENSEN.
8. Pneumatopteris latisquamata HOLTTUM, Blu-
mea 21 (1973) 305.— Type: MILLAR & HOLT-
TUM NGF 15777, N.E. New Guinea, Morobe
Distr., Edie Creek, 2000 m (K; BO, LAE).
Caudex short; apex covered with imbricating
firm, apparently hairless scales; stipe 4cm long,
minutely hairy, scales early caducous. Lamina
35cm long; pinnae c. 24 pairs, lower 8 pairs
gradually shorter, lowest 2cm long, all with
symmetric bases; texture firm, rigid when dried.
Largest pinnae 3.5 x 0.8 cm; base broadly cuneate;
apex short-acuminate; edges lobed c. 2 way to
costa; lobes slightly falcate, their edges reflexed
when dried; costules 2mm apart; veins 4 pairs,
lowest anastomosing, next pair to margin; lower
surface of rachis densely covered with erect aci-
cular hairs 0.3-0.4mm long, more sparse and
shorter similar hairs on costae, costules and veins
and on surface between veins, minute sessile or
subsessile glands on all parts; upper surface of
rachis covered with pale hairs more than 0.5 mm
long, shorter hairs on costae, rest of pinna bearing
many suberect acicular hairs 0.2mm long and
many short capitate hairs or small glands. Sori
probably medial, confluent at maturity and cover-
ing most of lower surface, exindusiate; sporangia
bearing either a short seta or a short capitate hair;
spores with extended sinuous wings which are
joined together irregularly.
Distr. Malesia: Papua New Guinea, only
known from the type.
Note. Allied to P. costata but very distinct.
9. Pneumatopteris egenolfioides HOLTTUM, Blu-
mea 21 (1973) 306.— Type: BRASS 11497, New
Guinea, near Lake Habbema, 2200 m, on limes-
tone cliffs (L; BM, BO, MICH).
Caudex slender, procumbent, 8cm long with a
tuft of stipes at apex; stipe 10-17 cm long, pale,
glabrous, base covered with dark ovate-acute
glabrous scales 1.2mm long. Lamina to 45cm
long; pinnae to 28 pairs, firm to rigid, 3-4 lower
pairs gradually smaller and more widely spaced,
lowest 5-7 mm long. Largest pinnae 2.4-3.2cm
long, 0.6-0.8cm wide, base strongly auricled on
acroscopic side, narrowly cuneate on basiscopic;
apex narrowed, subtruncate with a small mucro;
edges lobed c. 1/3 towards costa; lobes oblique,
near bases of larger pinnae obliquely truncate
with teeth at vein-ends; costules 3mm apart, at
little more than 45° to costa; veins mostly 2 pairs
(3 pairs in basal auricle), | pair anastomosing with
a short excurrent vein to the sinus-membrane, free
in distal part of pinna, pale and prominent on lower
surface; lower surface hairless, with residual very
small scales on costae, costules and veins, surface
between veins pustular; upper surface hairy on
rachis and bases of costae only. Sori only on basal
acroscopic vein of each group, near costule; in-
dusium firm, glabrous; a short capitate hair
present on some sporangia.
422
FLORA MALESIANA
[ser. Il, volar
Distr. Malesia: W. New Guinea (Habbema).
Ecol. Limestone cliffs at 2200 m.
10. Pneumatopteris patentipinna HOLTTUM, Biu-
mea 21 (1973) 307. — Type: T. G. WALKER 8477,
N.E. New Guinea, trail from Moro to Sewi (BM).
Caudex short-creeping; stipe 5—-15cm_ long,
pale, glabrescent or with a few brown hairs, basal
scales thin, caducous. Lamina to 60cm long (of
type 35cm); 5-10 pairs of lower pinnae gradually
smaller, lowest less than | cm long; reduced pinnae
either auricled on acroscopic base or dilated both
sides, middle ones broadly triangular, opposite or
not; all pinnae at right angles to rachis; texture
firm. Largest pinnae of type 4.0cm long, 1.1 cm
wide above auricled base, of largest plants seen
8.5 x 1.8 cm; base truncate; apex short-acuminate,
slightly upcurved; edges of pinnae of largest
fronds lobed 2 way to costa, of smaller fronds less
deeply, lobes truncate with acute falcate apex, +
toothed at ends of veins; costules 3-4 mm apart;
veins 4-6 pairs, pale and prominent, | pair anas-
tomosing, next vein to sinus-membrane or edge;
lower surface of rachis and costae bearing a vari-
able number of brown hairs, some short pale hairs
also on costae, rather sparse large pustules on
surface between veins; upper surface of rachis
covered with thick brown hairs to 1mm long,
hairs on costae shorter, few, pale distally. Sori
medial, indusia large, thin, bearing a variable
number of short acicular hairs; no glandular hairs
on sporangia. Chromosomes: n=36 (T. G.
WALKER).
Distr. Malesia: Eastern New Guinea, at 1650-
2300 m.
Note. The type appeared to be very distinct,
but later collections indicate a possible gradation
to the condition of P. jermyi.
11. Pneumatopteris | cheesmaniae HOLTTUM,
Blumea 21 (1973) 307. — Type: L. E. CHEESMAN
1269, Japen Island, Aian Range, 300m, in forest
on old coral limestone (BM).
Stipe at least 14cm long, pale, glabrous; lamina
26cm long, pinnae c. 14 pairs, basal 3 pairs of
pinnae gradually reduced, lowest 3-5 mm long.
Largest pinnae 3.3cm long, 1.1cm wide above
auricled base, basiscopic base truncate; apex
short-pointed; edges lobed 2/5 or a little more
deeply; lobes + quadrate, apices subtruncate with
slight projections at vein-ends, edge not con-
spicuously cartilaginous; costules 3—3.5 mm apart,
almost at right angles to costa; veins 3 pairs
except in basal auricle which has 4-5 pairs, slen-
der, concolorous, basal ones meeting to form a
very short excurrent vein below the sinus or in
smaller pinnae meeting at the sinus-membrane;
lower surface quite hairless, surface between
veins slightly pustular; upper surface not seen.
Sori inframedial, lowest not divergent, exindusiate;
sporangia often with a pear-shaped small gland.
Distr. Malesia: New Guinea (Japen I.), only
known from type.
Ecol. In forest on old coral limestone, 300m
altitude.
12. Pneumatopteris lithophila HOLTTUM, sp. nov.
Lamina 30 cm longa; pinnae 16—18-jugatae, in-
feriores 6-7-jugatae sensim decrescentes; pinnae
maximae 3.01.0 cm, marginibus crenatis, api-
cibus rotundatis, pagina inferiore fere glabra;
rachis supra pilis brunneis vestita, pinnae supra
inter venas pilis erectis acicularibus minutis
praeditae; sori mediales, indusia glabra. — Type:
M. G. PRICE & B. F. HERNAEZ 779, Western
Samar, on limestone cliff in forest, 700 m (K).
Caudex short; stipe 3-6cm long, pale, hairy in
groove, basal scales very small, broad, thin.
Lamina 30cm long; pinnae 16-18 pairs, 6-7 lower
pairs gradually decrescent, lowest 3 x 3 mm; tex-
ture thin. Largest sterile pinnae 3.0 1.1cm, fer-
tile 2.5 x 1.0 cm; base truncate, a little auricled on
acroscopic side; apex rounded; edges crenate to a
depth of 1-1.5mm; costules 2.5mm apart; veins
3-4 pairs at base of largest pinnae, basal pair
anastomosing to form an excurrent vein to the
short sinus-membrane, next acroscopic vein to the
membrane or the margin; lower surface of rachis
glabrous, very sparse minute acicular hairs
present on costae and costules, sometimes also
between veins, a few short capitate hairs on cos-
tules, veins and on surface between veins
which is slightly pustular; upper surface of rachis
covered with slender brown hairs 1 mm long, hairs
on costae pale and shorter, short erect acicular
hairs rather sparse on surface between veins. Sori
medial; indusia small, glabrous; neither glands nor
setae On sporangia; spores pale, with many small
wings.
Distr. Malesia: Philippines (W. Samar), 2 coll.
from type locality.
Ecol. On limestone cliff in forest at 700m
altitude.
13. Pneumatopteris angusticaudata HOLTTUM,
Blumea 21 (1973) 308.—Type: BRASS 24984,
New Guinea, Goodenough Island, 800m, on a
rock-face beside a stream (BM: LAE).
Fronds pendent (collector); stipe short; base of
stipe to first large pinna 50-55 cm; reduced pinnae
6 pairs, 3-5 mm long, then 2 pairs of intermediate
length. Lamina excluding reduced pinnae 65cm
long; pinnae 25 pairs; several pairs of lower pin-
nae narrowed towards their bases, especially on
basiscopic side; texture firm. Largest pinnae 15 x
1.2cm; base broadly cuneate to full width on
acroscopic side, narrowly cuneate on basiscopic;
apex acuminate with subentire cauda 4-5 cm long
and | mm wide; edges lobed to a depth of 2mm;
lobes very oblique, falcate, subacute, entire; cos-
tules 4mm apart; veins 4-5 pairs, | pair anas-
tomosing, next acroscopic vein to side of sinus-
1981]
THELYPTERIDACEAE (Holttum)
423
membrane; lower surface quite glabrous; upper
surface of rachis bearing a few short hairs, similar
hairs on bases of costae, rest glabrous. Sori
medial; indusia small, thin, with many short hairs;
no glands nor setae on sporangia.
Distr. Malesia: Papua New Guinea
(Goodenough I.), known from the type only.
Ecol. Rock-face beside a stream at 800m al-
titude.
14. Pneumatopteris microloncha (CHRIST) HOLT-
TUM, Blumea 21 (1973) 308. — Dryopteris micro-
loncha CHRIST, Philip. J. Sci. 2 (1907) Bot. 202. —
Cyclosorus microlonchus (CHRIST) COPEL. Gen.
Fil. (1947) 143; Fern Fl. Philip. (1960) 366. —
Thelypteris microloncha (CHRIST) REED, Phy-
tologia 7 (1968) 293.— Lectotype (HOLTTUM
1973): MANGUBAT BS 1304, Cavite, Luzon (P).
Dryopteris caudiculata v.A.v.R. Handb. (1908)
820 (based on Nephrodium caudiculatum J. SM.
nom. nud.).
Caudex erect; stipe 4-10cm long, pale,
glabrous, scales broad, thin, not persistent.
Lamina 25-45(-55) cm long; pinnae 15-25 pairs,
texture thin; 3-6 pairs of lower pinnae gradually
smaller, lowest 3-5 mm long. Largest pinnae 3-
12cm long, 0.6—-1.5 cm wide; base subtruncate, a
little dilated both sides or only on acroscopic side;
apex acuminate, + caudate on longest pinnae;
edges lobed 5 way to costa; lobes obliquely
quadrate, + dentate at ends of veins; costules
2-4mm apart; veins 3-4(-6) pairs, slender and
prominent, | pair anastomosing, next pair to sinus-
membrane on larger pinnae; lower surface bearing
short slender erect hairs throughout; upper sur-
face of rachis bearing slender pale hairs 0.5mm
long, similar hairs sparse on costae, no others.
Sori medial; indusia short-hairy; sporangia with
rather large pale or yellowish club-shaped hairs.
Distr. Malesia: Philippines (Luzon, Negros).
Ecol. At low altitudes on rocky stream-banks.
Note. CHRIST cited several specimens (the
lectotype is the best in his herbarium), among
them CUMING 317, which is one of several
numbers cited by JOHN SMITH under “Neph-
rodium caudiculatum PRESL” (a non-existent
name), the others representing Sphaerostephanos
productus. v.A.v.R., adopting SMITH’s name,
copied CHRIST’s description. Plants are fertile
from a small size.
15. Pneumatopteris brooksii (COPEL.) HOLTTUM,
Blumea 21 (1973) 308.—Dryopteris brooksii
CoPEL. Philip. J. Sci. 3 (1908) Bot. 345; v.A.v.R.
Handb. Suppl. (1917) 185.— Thelypteris brooksii
(COPEL.) REED, Phytologia 17 (1968) 265.—
Type: C. J. BRooKs s.n. April 1908, Bidi,
Sarawak (MICH; BM).
Caudex erect or suberect; stipe 20-50 cm long,
pale, glabrescent, basal scales appressed; base of
stipe to first large pinna 40-70 cm; reduced pinnae
3-4 pairs, 45mm _ long. Lamina excluding
reduced pinnae 75cm long; pinnae more than 40
pairs, texture firm; several pairs of lower pinnae
gradually narrowed towards their bases, not auri-
cled. Largest pinnae 10-12.5 x 1.0-1.3cm; base
abruptly broad-cuneate; apex evenly and narrowly
attenuate; edges lobed c. 1/3 towards costa, lobes
slightly oblique with rounded apices; costules 3.5-
4mm apart; veins 4-5 pairs (to 7 pairs in largest
sterile pinnae), slender, | pair anastomosing with a
long excurrent vein, next veins to edge; lower
surface quite glabrous, hardly pustular; a few
short pale hairs in groove of upper surface of
rachis, few or none on costae. Sori medial; in-
dusia small, firm, glabrous; sporangia lacking
glandular hairs.
Distr. Malesia:
collections.
Ecol. On limestone.
Borneo (Sarawak), several
16. Pneumatopteris inclusa (COPEL.) HOLTTUM,
Blumea 21 (1973) 306.— Dryopteris inclusa
CoPEL. Univ. Cal. Publ. Bot. 14 (1929) 373, pl.
57. — Thelypteris inclusa (COPEL.) REED, Phy-
tologia 17 (1968) 284.— Type: BARTLETT 8576,
Sumatra, Karo Plateau (UC).
Dryopteris berastagiensis C. CHR. Dansk Bot.
Ark. 9, 3 (1937) 59.— Type: RIDLEY s.n. 1921,
Sumatra, Berastagi (K).
Caudex creeping, 10mm diameter; stipe not
seen. Lamina to at least 100 cm long; pinnae to 20
pairs, thin but firm; basal pinnae not or slightly
reduced. Largest pinnae 28 x 5cm; base of lower
pinnae cuneate, of upper ones truncate; apex ab-
ruptly short-acuminate; edges lobed 1/4 towards
costa or less deeply; lobes obliquely truncate with
slight teeth at ends of veins; costules 6.5-7.5 mm
apart, at a little more than 60° to costa; veins
10-12 pairs, slender, 23 pairs anastomosing, 22-3
pairs passing to sides of sinus-membrane; lower
surface glabrous in type, with sparse slender hairs
on costae and costules of another specimen; upper
surface with short hairs on rachis and costae only.
Sori medial; indusia thin with a few short hairs;
hairs on stalks of sporangia with swollen orange
end-cell; spores with many small wings.
Distr. Malesia: Central and north Sumatra,
several collections.
Note. There is some variation among the spe-
cimens included here. One collected by C. G.
MATTHEW at Padang Panjang has thinner pinnae
than the type, with slender hairs on lower surface
of costae and costules; it also has the basal pair of
pinnae distinctly reduced (12 and 17 cm long) with
much-narrowed bases. Other collections show
different parts of fronds and are difficult to com-
pare.
17. Pneumatopteris longipes (BL.) HOLTTUM,
Blumea 21 (1973) 306.— Aspidium longipes BL.
En. Pl. Jav. (1828) 155; MIQUEL, Ann. Mus. Bot.
424
FLORA MALESIANA
[ser. II, vol. 1°
Lugd.-Bat. 4 (1869) 158. — Nephrodium longipes
(BL.) Moore, Ind. Fil. (1868) 95; RAcIB. FI. Btzg
1 (1898) 182.—Dryopteris longipes (BL.) O.
KTZE, Rev. Gen. Pl. 2 (1891) 813; v.A.v.R.
Handb. (1908) 211; BACKER & POSTH. Varenfl.
Java (1939) 58.— Thelypteris longipes (BL.)
REED, Phytologia 17 (1968) 289. — Type: BLUME,
Java, Boerangrang (L, n. 908,335—1062).
Caudex long-creeping, to 10 mm diameter; stipe
c. 100cm long, pale, glabrous, basal scales c.
10x4mm. Lamina c. 100cm long; pinnae to 35
pairs; basal pair of pinnae not opposite, not auri-
cled, variably unequal and reduced, in some cases
2.5cm long, in others much longer, + narrowed
near their bases. Largest pinnae 17-22 x 2.2-
3.0cm; base truncate; apex narrowly acuminate;
edges lobed c. 2/3 towards costa, lobes with
rounded tips, at most slightly dentate at vein-ends;
costules 4 mm apart; veins to 11 pairs, lowest only
anastomosing, next pair to margin or to the short
sinus-membrane; sparse erect hairs on lower sur-
face of costules and distally on costae, on sinus-
membranes and margin, surface between veins
finely and not copiously pustular; upper surface
hairy on rachis and costae only. Sori medial;
indusia small, hairy; sporangia sometimes with a
small capitate hair.
Distr. Malesia: Java, at 1400-1800 m.
18. Pneumatopteris laevis (METT.) HOLTTUM,
Blumea 21 (1973) 308.— Aspidium laeve METT.
Farngatt. IV (1858) 104.— Dryopteris laevis
(METT.) C. CHR. Ind. Fil. (1905) 273; v.A.v.R.
Handb. (1908) 220.— Thelypteris laevis (METT.)
REED, Phytologia 17 (1968) 286.— Neotype
(HOLTTUM 1973): JAGOR, Samar (B, ex Herb.
Mett.).
Dryopteris luzonica CHRIST, Philip. J. Sci. 2
(1907) Bot. 196, excl. var. puberula; v.A.v.R.
Handb. (1908) 821, excl. var. puberula. — Lec-
totype (HOLTTUM 1973): LOHER s.n. Jan. 1906,
Mt Makiling, Luzon (P).
Cyclosorus nitidulus sensu COPEL. Fern FI.
Philip. (1960) 364, p.p. — Cyclosorus microlonchus
sensu COPEL. ibid. 366, p.p. — Fig. 11g-h.
Caudex quite prostrate with closely seriate
fronds; stipe 12-18cm long, pale, glabrous; base
of stipe to first large pinna c. 30cm; reduced
pinnae 3-5 pairs, transition to large ones gradual
or subabrupt, lowest 5-10 mm long, all strongly
auricled on acroscopic base. Lamina in all 50-
80cm long (sometimes fertile at a smaller size);
pinnae to 25 pairs. Largest pinnae to 18cm long,
1.4-2.0 cm wide; middle and upper pinnae oblique,
with very asymmetric base, auricled on acroscopic
side, narrower and rounded on basiscopic; apex
caudate-acuminate (cauda 24cm long); edges
lobed c. 1/3 towards costa; lobes oblique, sub-
truncate with teeth at vein-ends; costules 4-
4.5mm apart; veins 5-8 pairs, slender, basal pair
anastomosing, next acroscopic vein to sinus-
membrane; lower surface quite glabrous, slightly
pustular between veins; upper surface bearing
short pale hairs on rachis and costae (in a few
cases hairs on rachis red-brown). Sori medial,
lower ones not divergent; indusia glabrous or with
a few short acicular or capitate hairs on edges;
sporangia usually lacking glands. Chromosomes:
n= 36 (T. G. WALKER, plant cult. Kew).
Distr. Malesia: Philippines (Luzon, Negros,
Samar, Leyte, Mindanao).
Ecol. On rocky banks of streams, with pen-
dulous fronds; low altitudes.
Notes. Mettenius cited as type “Cuming 83 bis
ex parte’; this is not represented in his herbarium
at Berlin. The neotype is one named by him and
agreeing with his description. Copeland’s treat-
ment is very confused. M. G. Price has sent to
Kew several dried specimens and also living
plants which have supplied information for the
above description. One of his specimens has red-
brown hairs on the rachis and a few capitate hairs
on its sporangia.
19. Pneumatopteris obliqua HOLTTUM, Blumea 21
(1973) 309. — Type: P. & F. SARASIN 127, Mas-
arang, N. Celebes (BAS).
Caudex short-creeping; stipe c. 40 cm long, pale,
glabrous; base of stipe to first large pinna 90 cm or
more; reduced pinnae to 12 pairs, all c. 33mm
or the uppermost a little larger, transition to large
pinnae abrupt. Lamina excluding reduced pinnae
to 65cm long; pinnae 25 pairs, all except lower
ones very oblique; several pairs of lower pinnae
gradually narrowed towards their bases, not auri-
cled. Middle pinnae to 16x 1.6-1.9cm; base very
asymmetric, basal basiscopic lobe much reduced,
basal acroscopic lobe not longer than next; apex
acuminate with almost entire cauda 34cm long;
edges lobed c. 3/5 towards costa, lobes with fal-
cate acute tips and slight teeth at vein-ends; cos-
tules to 5mm apart, at c. 50° to costa; veins
slender, 7-8 pairs, basal pair meeting at base of
sinus-membrane, next acroscopic vein passing to
side of membrane, rest to margin; lower surface
of rachis rather densely covered with hairs 0.1-
0.2 mm long, similar hairs more sparse on costae,
surface between veins with rather sparse large
pustules; upper surface of rachis and costae bear-
ing pale hairs to 0.5 mm long. Sori medial; indusia
thin, glabrous or with a few short hairs; sporangia
sometimes with small capitate hairs.
Distr. Malesia: North Celebes, in forest at
1100-1200 m (5 collections).
20. Pneumatopteris basicurtata HOLTTUM, Blu-
mea 21 (1973) 309. — Type: ROBINSON & KLOss
148, G. Kerinci, Sumatra, 2250 m (BM).
Stipe 42cm long, pale, minutely hairy; lamina
50 cm long, comprising 15 pairs normal pinnae and
one basal pair 2cm long, 1.2cm wide at base
which is strongly auricled on acroscopic side;
1981]
THELYPTERIDACEAE (Holttum)
425
lowest normal pinnae more widely-spaced,
deflexed, much narrowed at their bases on basis-
copic side; texture thin but firm. Largest pinnae of
type 12x2.4cm, of BUNNEMEIJER 10296 19x
2.7 cm; base truncate, slightly dilated both sides;
apex short-acuminate; edges lobed a little more
than > way to costa; lobes falcate with + acute
tips, not toothed at vein-ends; costules 5—5.5 mm
apart, at a wide angle; veins 9-10 pairs, basal pair
anastomosing, next pair to sinus-membrane; lower
surface quite glabrous except for a few slender
hairs on costules and veins near tips of lobes, also
on sinus-membrane and margin. Sori medial; in-
dusia large, rather firm, with a few hairs; sporan-
gia sometimes with capitate hairs.
Distr. Malesia: Sumatra: G. Kerinci (2 collec-
tions) and near Lake Toba.
Note. It is possible that these specimens are
not distinct from P. longipes of Java; information
about the caudex and constancy of basal pinnae is
needed.
21. Pneumatopteris microauriculata HOLTTUM,
Blumea 21 (1973) 311. — Type: CLEMENS 27137,
Mt Kinabalu, Sabah, 850 m (BM).
Caudex short-creeping with closely seriate
fronds; stipe 50-70cm long, pale except near
base, glabrous, basal scales c. 2.52.0mm.
Lamina 35-40cm long, consisting of c. 18 pairs
normal pinnae and | pair 1 mm long 6-8 cm below
basal normal pair; basal pinnae narrowed gradu-
ally towards their bases which are 8 mm wide, not
auricled; texture firm. Largest pinnae 12 x 1.6cm;
base somewhat asymmetric, truncate and slightly
auricled on acroscopic side, narrow and rounded
on basiscopic; apex upcurved, acuminate, serrate
to tip; edges lobed c. 2/5 towards costa; lobes
falcate, broadly pointed, not toothed; costules 3.5-
4mm apart, at a wide angle to costa; veins 8 pairs,
concolorous, prominent both sides, | pair anas-
tomosing, 13-2 pairs passing to sides of sinus-
membrane; lower surface of rachis glabrous, cos-
tae bearing very short pale hairs distally, a few
hairs on sinus-membrane, surface between veins
pustular; upper surface hairy on rachis and costae
only. Sori a little inframedial, lower ones not
divergent; indusia large, thin, glabrous, caducous;
sporangia lacking glands.
Distr. Malesia: Borneo (Sabah: Mt Kinabalu),
3 collections.
22. Pneumatopteris ecallosa (HOLTTUM) HOLT-
TUM, Blumea 21 (1973) 310.— Cyclosorus ecal-
losus HOLTTUM, Gard. Bull. Sing. 11 (1947) 269;
Rev. Fl. Mal. 2 (1955) 272, f. 156. — Thelypteris
ecallosa (HOLTTUM) REED, Phytologia 17 (1968)
274. — Type: Ho_tTTuM 31294, Cameron High-
lands, Malaya (SING; BO, K).
Caudex short-creeping; stipe to 100cm long,
pale except near base, glabrous, scales thin, ad-
herent. Lamina to 100 cm long, consisting of c. 35
pairs normal pinnae and at the base 2-3(-4) pairs
of subequal opposite pinnae c. 2.5 x 2.5 cm, deeply
lobed, strongly auricled on the acroscopic side;
several pairs of lower normal pinnae narrowed
towards their bases with a lobed acroscopic auri-
cle; texture thin. Largest pinnae commonly 20 x
2.2 cm, sterile pinnae to 28 x 3. cm; base truncate;
apex narrowly attenuate; edges lobed c. > way to
costa (sterile pinnae often more deeply); lobes not
falcate, truncate with slight teeth at vein-ends;
costules to 5mm apart, at a wide angle to costa;
veins 7-12 pairs, prominent both sides, lowest pair
anastomosing, next pair to sinus-membrane; lower
surface quite glabrous, minutely pustular between
veins; upper surface of rachis and costae bearing
short pale hairs. Sori inframedial, lower ones not
divergent; indusia thin, glabrous; sporangia bear-
ing many short club-shaped glands.
Distr. Malesia: Malaya (middle and north of
Main Range).
Ecol. Near streams in forest; now abundant in
secondary growth near streams where forest has
been felled at Cameron Highlands, 1500 m.
23. Pneumatopteris auctipinna HOLTTUM, Blumea
21 (1973) 310; Reinwardtia 8 (1974) 499. — Type: P.
& F. SARASIN 119, N. Celebes, Tomohon (BAS).
Aspidium truncatum var. celebicum? MIQUEL,
Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 160. — Type:
DE VRIESE, Menado, N. Celebes (L).
Aspidium truncatum sensu CHRIST, Ann. Jard.
Bot. Btzg 15 (1898) 133, as regards specimens
cited from Celebes.
Caudex short, erect (ALSTON); stipe 20cm
long, pale, minutely hairy; base of stipe to first
large pinna 75cm; reduced pinnae c. 8 pairs, all
with a broad asymmetric base which is enlarged to
form a serrate auricle, uppermost 2.5-4.0 x 2-
3cm, lowest 1.5x1.5cm. Lamina _ excluding
reduced pinnae 80cm or more long; pinnae c. 25
pairs, thin but firm, 2-3 lower pairs with + auri-
cled base as reduced pinnae. Largest pinnae 20 x
2.5cm; base truncate (except sub-basal ones);
apex caudate-acuminate (cauda 2.5 cm, sinuous or
crenate); edges lobed 1/2-1/3 towards costa; lobes
subtruncate to rounded with slight teeth at vein-
ends; costules 44.5 mm apart, at a wide angle to
costa; veins 8-10 pairs, slender, prominent both
sides, often pale reddish on lower surface, basal
1-1; pairs anastomosing, next 2-23 pairs to the
sinus-membrane; lower surface of rachis and cos-
tae rather densely covered with short erect pale
hairs, sparse hairs on costules, veins and on sur-
face between veins which is finely pustular; hairs
on upper surface of costae pale, 0.5 mm long, rest
of surface glabrous. Sori a little inframedial, basal
ones not divergent; indusia small, short-hairy;
sporangia bearing capitate hairs.
Distr. Malesia: North & Central Celebes, East
Java, Lesser Sunda Is. (Flores) and Moluccas
(Buru & Amboina), 900-2000 m.
426
FLORA MALESIANA
[ser. II, vol. 17
Note. SARASIN 124 bears a note that young
fronds are rose-coloured.
24. Pneumatopteris jermyi HOLTTUM, Blumea 21
(1973) 310.—Type: JERMyY 3739, N.E. New
Guinea, Butemu, Moro Trail (BM).
Caudex short, suberect; stipe 23 cm long, pale,
hairs brown, sparse above base; base of stipe to
first large pinna 50 cm; reduced pinnae c. 6 pairs,
not opposite, uppermost 1.5-2.0cm long, 0.8 cm
wide at base which is truncate and slightly auri-
cled on acroscopic side, lowest 2-3 mm long, +
orbicular. Lamina excluding reduced pinnae
60 cm long; pinnae 22 pairs, texture thin but firm;
lowest pinnae slightly narrowed towards their
bases, + auricled on acroscopic side. Largest
pinnae of type 13x 1.8cm (of another specimen
14x 2.5cm); base truncate; apex acuminate with
narrow subentire cauda 2-3.5cm long; edges
lobed > way to costa or a little less deeply, lobes
subtruncate with slight projections at vein-ends;
costules 4mm apart, at a wide angle to costa;
veins to 9 pairs, concolorous, basal pair meeting at
an obtuse angle to produce a slender excurrent
vein which may be joined by another vein before
entering base of sinus-membrane, next pair of
veins to sides of the membrane; lower surface of
rachis bearing stiff curved brown hairs more than
0.5 mm long, similar hairs near bases costae grad-
ing to pale hairs distally, rest of surface almost
glabrous; upper surface of rachis and costae hairy
as lower. Sori inframedial, lower ones not diver-
gent; indusia_ short-hairy; sporangia lacking
glands.
Distr. Malesia: Eastern New Guinea,
2000 m.
1150-
25. Pneumatopteris papuana HOLTTUM, Blumea
21 (1973) 311. — Type: BRASS 22778, Mt Dayman,
E. New Guinea, 2000 m (LAE; BM, L).
Differs from P. jermyi as follows: lamina
excluding reduced pinnae to 150 cm long; largest
pinnae 18 x 3.5cm, lobed 1/2-2/3 towards costa;
costules 5-6 mm apart; basal veins meeting at an
acute angle at base of sinus-membrane, sometimes
without true anastomosis; lower surface of costae
and rachis glabrous or with few brown hairs;
indusia glabrous.
Distr. Malesia: Eastern New Guinea, 1750-
2000 m.
Note. A specimen from near Bulolo has pinnae
to 24x 3.5cm, lobed rather more than 2/3, with
copious brown hairs on lower surface of rachis
and costae. Field study of this and P. jermyi is
needed to confirm distinctions between them, or
possibly to indicate that they should be united.
26. Pneumatopteris rodigasiana (TIT. MOORE)
HOLTTUM, Blumea 21 (1973) 319; Allertonia 1
(1977) 233.—Nephrodium rodigasianum T.
Moore in Linden, Ill. Hort. 29 (1882) 27, pl.
442.— Thelypteris rodigasiana (T. MOORE)
REED, Phytologia 17 (1968) 310.— Dryopteris
transversaria (BRACK.) C. CHR. var. rodigasiana
C. CHR. Bishop Mus. Bull. 177 (1943) 88. — Type:
cult. LINDEN, ex Herb. T. Moore (K).
Cyclosorus rotumaensis ST. JOHN, Occ. Pap.
Bish. Mus. 21 (1954) 180.—Type: St. JOHN
19139, Rotuma I. (BISH; K).
Caudex erect; stipe 9-l18cm long, pale,
minutely hairy. Lamina to 100cm or more long;
pinnae thin, lower 6-9 pairs gradually decrescent,
lowest less than I1cm long, none of them either
narrowed or auricled at their bases. Largest pin-
nae 18x2cm; base truncate; apex narrowly
acuminate; edges lobed > way to costa; lobes
slightly falcate with rounded tips, projections at
vein-ends slight; costules 4-5.5mm apart; veins
slender, hardly prominent, concolorous, 6-9 pairs,
1 pair anastomosing, next acroscopic vein to
sinus-membrane or to margin; lower surface of
rachis and costae bearing very short erect pale
hairs, a few such hairs on costules and veins and
some on the surface between veins; upper surface
of rachis bearing pale hairs 0.5 mm long, shorter
hairs on costae, no others. Sori a little in-
framedial; indusia thin, glabrous or with a few
hairs in the middle; sporangia bearing capitate
hairs. Chromosomes: n= 36 (T. G. WALKER).
Distr. Polynesia (Samoa, Niue I.), Melanesia
(Solomons, New Hebrides, Rotuma I.), and E.
Malesia: Papua New Guinea (Admiralty Is., New
Ireland).
27. Pneumatopteris japenensis HOLTTUM, sp. nov.
Pinnae basales redactae 12-jugatae, superiores
1.5 cm longae, auriculatae; lamina 100 cm longa;
pinnae maximae 24x2cm, profunde lobatae,
tenues; venae infimae anastomosantes, vena
communi excurrenti perbrevi; pagina inferior inter
venas glandulas minutas ferens; indusia parva,
glabra. — Type: AET & IDJAN 426, Japen Island,
W. New Guinea (SING; K, L).
Caudex short, erect or suberect; stipe c. 12cm
long, pale, glabrescent, when young covered with
scales 15 x 3 mm; base of stipe to first large pinnae
65cm; reduced pinnae 12 pairs, upper ones auri-
cled, uppermost 1.5 cm long with auricle 4 x 1mm
on acroscopic base, narrowed to basiscopic base,
lower ones less than 5mm long, an intermediate
pinna c. 8cm long sometimes present. Lamina
excluding reduced pinnae more than 100 cm long;
pinnae more than 30 pairs, thin, drying pale
olivaceous; lower pinnae somewhat narrowed at
their bases which are slightly auricled. Largest
pinnae 24x 2.0cm; base truncate; apex with nar-
row entire cauda 2.5x0.3cm; edges lobed to
1.5mm from costa; lobes at right angles to costa,
separated by broad sinuses, their tips rounded
with slight teeth at vein-ends; costules 4-5 mm
apart; veins 12-13 pairs, basal pair anastomosing
to form a very short vein excurrent to the sinus;
1981]
THELYPTERIDACEAE (Holttum)
427
lower surface of rachis, costae and costules bear-
ing sparse pale hairs to | mm long, between veins
sparse short erect acicular hairs and small colour-
less glands which collapse when dried, surface not
pustular; upper surface of rachis bearing coarse
pale hairs 1mm long, hairs on costae shorter,
minute acicular hairs and glands between veins.
Sori a little inframedial, lower ones not divergent;
indusia small, glabrous; sporangia apparently lack-
ing glands; spores with a fairly broad erose wing
and some anastomosing cross-wings.
Distr. Malesia: New Guinea (Japen I.), only
known from type.
28. Pneumatopteris tobaica HOLTTUM, Blumea 21
(1973) 316. Type: SURBECK 14, N. Sumatra, S
side of Lake Toba, 1900 m (L; BO).
Stipe not seen; base of stipe to first large pinna
more than 50cm; reduced pinnae 56cm apart, c.
7 x7mm, not auricled; basal normal pinnae much
narrowed towards their bases, not auricled, base
of fifth pinna less than | cm wide. Largest pinnae
24x 2.8cm; apex acuminate, serrate to tip; edges
lobed to 4mm from costa; lobes oblong, sub-
truncate with irregular short teeth at vein-ends;
costules 5-5.5 mm apart; veins 11-12 pairs, basal
pair anastomosing, rest all to margin; lower sur-
face of rachis bearing sparse hairs, of pinnae
glabrous apart from a few hairs on sinus-mem-
branes and margin, surface slightly pustular; upper
surface of costa bearing sparse hairs, rest
glabrous. Sori inframedial: indusia firm, with a
few short hairs; sporangia bearing capitate hairs.
Distr. Malesia: N. Sumatra. Known only from
type collection.
Note. The Bogor isotype includes a small plant
with erect caudex, but it is not certain that this
represents the same species as the large frond.
The reduced pinnae are only shown by the Leiden
specimen; apart from them, the fronds are very
like those of P. longipes. More collections are
needed.
29. Pneumatopteris incisa HOLTTUM Blumea 21
(1973) 317. — Type: ALSTON 16623, Ternate, G.
Gamalama (BM).
Stipe 15cm long; base of stipe to first large
pinna 50cm; reduced pinnae 7 pairs, subopposite,
uppermost 1.5x1.0cm, triangular, base sym-
metrically truncate, edges crenate, apex blunt,
lowest 5-6 mm long and wide. Lamina excluding
reduced pinnae 70 cm long; pinnae 30 pairs, texture
firm; lower pinnae sub-opposite, gradually nar-
rowed towards their bases, base of lowest 10 mm
wide, not auricled; middle pinnae with basal 1-2
pairs of lobes somewhat reduced, basal basiscopic
lobe usually longer than acroscopic; upper pinnae
with cuneate bases. Largest pinnae 21 x2.7 cm;
apex acuminate with subentire cauda 2—2.5cm
long; edges lobed to 2.5mm from costa; lobes
slightly falcate, their tips rounded, entire: costules
5mm apart, at a wide angle to costa; veins to 12
pairs, slender, pale and prominent both sides,
basal pair anastomosing, next pair usually both to
margin; lower surface of rachis sparsely hairy, of
costae and costules with short slender pale erect
hairs, somewhat longer hairs on veins and spar-
sely on surface between veins, longer ones on
sinus-membrane and margin, surface between
veins slightly pustular; upper surface of costae
with hairs 1 mm long, rest of pinna glabrous. Sori
inframedial, lower ones not divergent; indusia
thin, with slender hairs; sporangia bearing capitate
hairs.
Distr. Malesia:
hera, PLEYTE 397).
Ecol. By stream in forest, at 600 m.
Moluccas (Ternate: Halma-
30. Pneumatopteris psilophylla HOLTTUM, sp.
nov.
Pinnae redactae 3-jugatae, usque 7x5 mm;
pinnae normales steriles usque 17X2.8cm, 2/
costam versus lobatae, subtus perfecte glabrae;
venae 12-jugatae, infimae vel anastomosantes vel
ad basin membranae sinus conniventes, par
sequens latera membranae tegentes ; pinnae fertiles
ignotae. — Type: JERMY 13741, Sarawak, Gunong
Mulu (BM).
Caudex erect (collector); stipe 10cm long, pale,
basal scales thin, to 8mm long, 1-2 mm wide at
base, small residual ones adherent; base of stipe
to first normal pinna 25-30 cm; reduced pinnae 3
pairs, uppermost 7x5mm, deltoid with almost
symmetrical base, lowest very small; aerophores
not elongate on dried specimens but functional on
very young fronds which are covered with
mucilage (collector). Lamina (sterile) 65 cm long:
pinnae c. 25 pairs, basal pinnae slightly reduced
and narrowed towards their bases in basal 5cm,
base 7-8mm wide, pinnae above base suc-
cessively less narrowed, 6 pairs or more with
basal 1-2 pairs of lobes reduced. Largest pinnae
17 x 2.8 cm; apex acuminate with entire cauda 1.5—
2.0cm long: edges lobed 2/3 towards costa or a
little more deeply, lobes slightly falcate, slightly
narrowed distally, edges entire, cartilaginous
margins well-marked; costules 5mm apart: veins
12 pairs, slender, slightly prominent both sides,
basal pair anastomosing or meeting at the base of
the sinus-membrane, second pair passing to the
sides of the membrane: lower surface quite
glabrous: upper surface with sparse short pale
hairs on rachis and costae. Fertile fronds not
known.
Distr. Malesia: Borneo (Sarawak: Mt Mulu),
only known from type.
Ecol. In humus in limestone crevices at 150m.
Note. This is related to P. truncata but its
pinnae are much more deeply lobed; it is probably
confined to limestone.
31. Pneumatopteris pergamacea HOLTTUM, Blu-
428
FLORA MALESIANA
[ser. II, vol. 1°
mea 21 (1973) 315.—Type: JERMyY 7877, New
Ireland, cult. Hort. Bot. Kew, 123/70, n. 1035
(K).
Caudex short-creeping; stipe 10cm long, bear-
ing many rather persistent + spreading scales 6-
10 x 1 mm with acicular hairs 0.3 mm long on their
dorsal surface; base of stipe to first normal pinna
60cm; reduced pinnae to 15 pairs, uppermost
3.0 1.7 cm, subtriangular, base broadly cuneate
and almost symmetric, edges lobed, tip acute,
lowest c. 6X 4mm with base auricled both sides.
Lamina excluding reduced pinnae 75cm long,
rather thin; pinnae 28 pairs; lowest 3.5 cm wide in
the middle, narrowed towards the base which is
1.8cm wide, not auricled. Largest pinnae c. 27 x
3cm; base broadly cuneate to full width; apex
rather abruptly caudate-acuminate, cauda suben-
tire, to 3cm long; edges lobed 3/5 towards costa;
lobes slightly falcate, tips bluntly pointed, margins
entire or nearly so; costules 5-5.5 mm apart, at c.
60° to costa; veins 11-12 pairs, concolorous,
hardly prominent, basal pair anastomosing to form
a short excurrent vein to base of sinus-membrane,
next I-13 pairs passing to sides of the membrane;
lower surface of rachis and costae sparsely and
minutely hairy (hairs more abundant distally in
both cases), a few short hairs on sinus-membranes
and margins of lobes, rest glabrous, old fronds
distinctly pustular between veins; upper surface
of rachis bearing pale hairs more than 0.5 mm long
and shorter ones, hairs on costae similar, rest of
surface of pinnae of large fronds glabrous but
acicular hairs present between veins of the small
fronds of young plants, also on reduced basal pin-
nae of mature plants. Sori a little inframedial on
distal veins, further from costules on lower veins
especially near apex of pinna; indusia very small,
hidden by mature sporangia, bearing a few short
hairs; sporangia lacking glands; spores medium
brown, bearing many minute wings.
Distr. Malesia: Papua New Guinea (New
Ireland), only known from the type.
Ecol. On mossy limestone boulder in forest at
630 m.
32. Pneumatopteris sogerensis (GEPP) HOLTTUM,
Blumea 21 (1973) 315.— Dryopteris sogerensis
GEPP, J. Bot. 61 (1923) Suppl. 61. — Cyclosorus
sogerensis (GEPP) COPEL. Gen. Fil. (1947) 146;
Philip. J. Sci. 78 (1951) 450.— Thelypteris
sogerensis (GEPP) REED, Phytologia 17 (1968)
314.— Type: FORBES 446, Sogere, S.E. New
Guinea (BM).
P. laticuneata HOLTTUM, Blumea 21 (1973)
312. — Type: JERMY 8044, N.E. New Guinea (BM).
Caudex erect; stipe 15-30cm long, pale,
glabrous, bearing residual appressed scales; base
of stipe to first normal pinna 70cm or more;
reduced pinnae c. 6 pairs, gradually decrescent
downwards, uppermost c. 4cm long, 1.5cm wide
at base which is slightly auricled both sides,
crenate or lobed above base, lowest 10 x 5mm or
smaller. Lamina to 150 cm long, rather thin; pin-
nae to more than 40 pairs; lower pinnae narrowed
near their bases which are about 3 maximum width.
Largest pinnae commonly 27 x 2.5-3 cm, on lar-
gest fronds 36x 4.5cm (fronds often fertile at a
much smaller size); base subtruncate; apex acu-
minate but not narrowly caudate; edges lobed 3
way to costa (less deeply in small plants, up to 2/3
in largest); lobes slightly falcate with slight pro-
jections at vein-ends; costules 4-5.5mm apart,
pale and prominent on lower surface; veins to 14
pairs, slender, concolorous, slightly prominent, 1
pair anastomosing and | pair to sides of sinus-
membrane; lower surface entirely glabrous; upper
surface rather sparsely hairy on rachis and costae,
hairs pale, to 1mm long. Sori inframedial, basal
ones not divergent; indusia usually glabrous,
rarely with some acicular hairs; sporangia lacking
glands; spores pale with many small wings.
Chromosomes: n= 36 (T. G. WALKER).
Distr. Australia (Queensland and north of New
South Wales), Melanesia (Solomon Is.) and
Malesia: Papua New Guinea (Bismarck Arch.)
and Moluccas.
Ecol. In lowland forest,
streams; a common species.
Note. P. laticuneata was based on a small
plant with shallowly lobed pinnae.
especially near
33. Pneumatopteris micropaleata HOLTTUM,
Blumea 21 (1973) 319. — Type: HOLTTUM 58, Mt
Kinabalu, Sabah, on wet rocks near waterfall,
1800 m (K; SING).
Caudex erect; stipe 10cm long, pale, bearing
short pale acicular hairs and appressed scales;
base of stipe to first large pinna 50cm; reduced
pinnae 4-5 pairs, uppermost 1cm long, 0.9mm
wide at subtruncate base, almost semicircular with
projections at vein-ends, lowest 5mm_ long.
Lamina to 100cm long, texture firm; pinnae c. 30
pairs; 1-2 pairs basal pinnae gradually narrowed in
basal 2cm. Largest pinnae 17X1.8cm; base
truncate; apex acuminate with entire cauda to
2 cm long; edges lobed a little more than 3 way to
costa (basal pinnae lobed 3/5); lobes slightly
oblique, not falcate, their apices broadly rounded,
projections at vein-ends slight; costules 3-3.5 mm
apart on a fertile frond; veins to 11 pairs, slender,
concolorous, | pair anastomosing or meeting at
base of sinus-membrane, next pair to sides of
membrane; lower surface of rachis bearing rather
sparse slender pale hairs 0.2-0.4 mm long, hairs on
costae very short, near base only, on costules
slender appressed pale hairs 0.5mm long, small
scales (the smallest linear) at first abundant on
costae and costules; upper surface of rachis
covered with slender pale hairs c. 1mm long,
similar shorter hairs on costae and sparse on
costules and veins. Sori a little inframedial, basal
ones not divergent; indusia glabrous; sporangia
1981]
bearing small capitate hairs; spores not seen
(sporangia immature).
Distr. Malesia: Sabah (Mt Kinabalu), known
from type only.
Ecol. On wet rocks near waterfall at 1800 m.
34. Pneumatopteris truncata (POIR.) HOLTTUM,
Blumea 21 (1973) 314.— Polypodium truncatum
Porr. Encycl. Meth. 5 (1804) 534.— Dryopteris
truncata (Porr.) C. CHR. Ind. Fil. (1905) 299, excl.
Polystichum truncatum GAUD.; v.A.v.R. Handb.
(1908) 227, quoad pl. Males.; BACKER & POSTH.
Varenfl. Java (1939) 54.—Cyclosorus truncatus
(Porr.) FARW. Amer. Midl. Nat. 12 (1931) 259;
CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938)
216; HOLTTUM, Rev. FI. Mal. 2 (1955) 266, f. 152;
COPEL. Fern FI. Philip. (1960) 367. — Thelypteris
truncata (PorIR.) K. WATS. Mem. Coll. Sci. Univ.
Kyoto B, 31 (1964) 33.—Type: “Brézil”’, no
collector cited (P).
Aspidium abortivum BL. En. Pl. Jav. (1828)
154, incl. var. B (var. C?). — Dryopteris abortiva
(Be) eO. Krze. Rev: (Gen. "Pls 2) (1891) 812;
v.A.v.R. Handb. (1908) 217. — Type: BLUME, W.
Java (L, n. 908, 337-855).
Aspidium abruptum BL. En. Pl. Jav. (1828)
154.— Type: KUHL & VAN HASSELT, W. Java
(L, n. 908, 337-817).
Aspidium eusorum THw. Enum. PI. Zeyl. (1864)
391.— Nephrodium eusorum (THW.) BEDD.
Ferns Br. India (1866) pl. 130.—Type:
THWAITES CP 3064, Ceylon (K).
Dryopteris batacorum var. winkleri ROSENST.
Fedde Rep. 13 (1914) 217.— Type: J. WINKLER
158a, Sumatra, in terra Batacorum (S-PA).
Cyclosorus lepidopodus C. CHR. & TARD.
Notul. Syst. 7 (1938) 73; Fl. Gén. I.-C. 7, 2 (1941)
392, f. 45.— Thelypteris lepidopoda (C. CHR. &
TARD.) REED, Phytologia 17 (1968) 287. — Type:
EBERHARDT 5252, Tonkin, Bac Kan (BM).
P. christelloides HOLTTUM, Blumea 21 (1973)
311.—Type: CLEMENS 27451, Sabah, Mt
Kinabalu, 1000 m (K; BM).
Nephrodium truncatum sensu RACIB. FI. Btzg 1
(1898) 190, non (Gaud.) PRESL. — Fig. 11d-f.
Caudex erect; stipe 10-30cm long, pale,
glabrous, bearing residual thin appressed scales;
base of stipe to first normal pinna 60-85 cm;
reduced pinnae 6-10 pairs, not opposite, 3-6cm
apart, uppermost c. 1 xX 1 cm, not auricled, 1 or 2
intermediate pinnae sometimes present. Lamina
to 120 cm long, thin but firm; pinnae to 30 pairs or
more; several pairs of lower pinnae much nar-
rowed near their bases which are less than 1 cm
wide and unlobed for 1-2cm, not auricled. Lar-
gest pinnae commonly 20 x 2.5 cm (to 32 x 3.5 cm);
base broadly cuneate to truncate; apex acuminate,
in broader pinnae rather abruptly; edges lobed c.
1/3 towards costa, less deeply in narrower pinnae;
lobes almost at right angles to costa, truncate,
always with some teeth at ends of veins; costules
THELYPTERIDACEAE (Holttum)
429
4-5 mm apart; veins to 12 pairs, pale and prom-
inent, 15-2 pairs anastomosing, 2 pairs to sides of
the sinus-membrane where they usually unite to
form a distinct vein on each side of the distal part
of the membrane, often continuing close to the
margin of a pinna-lobe to join higher veins there;
lower surface of rachis and costae bearing a vari-
able number of short slender erect pale hairs (in
Sarawak sometimes quite glabrous), sparse hairs
also on costules and veins and sometimes a few
between veins where the surface is always pus-
tular; upper surface of rachis bearing longer pale
hairs, short hairs sparse on costae, minute on
costules, sometimes a few between veins near
apices of pinna-lobes. Sori a little inframedial,
lower ones not divergent; indusia thin, glabrous
(hairy on two specimens from Mt Kinabalu):
sporangia bearing elongate small glands (lacking in
some Sarawak specimens); spores with many
small wings.
Distr. Ceylon & S. India; N.E. India to S.
China; Western Malesia, Lesser Sunda Islands
and Philippines.
Ecol. Near streams in forest, at 100-1200 m.
Notes. The epithet truncatum, as used by 19th
century authors, was based on Polystichum trun-
catum GAUD. CHRISTENSEN cited Polypodium
truncatum PorrR. as basionym in Ind. Fil. (1905),
wrongly placing GAUDICHAUD’s name as a
synonym. CHING was the first to see and des-
cribe POIRET’s type, and accepted its origin as
Brazil, but there is no species at all like it in S.
America, and the specimen closely matches those
from Ceylon and Malaya. A plant examined by
MANTON in Ceylon was tetraploid. Smaller plants
in N. India are diploid and differ in pinna-lobes
and larger glands on sporangia (see HOLTTUM
1973); plants from G. Mulu, Sarawak, lacking
glands on sporangia, are diploid (T. G. WALKER).
The type of Aspidium abortivum BL. was a
lowland plant and agrees well with specimens
from Malaya. The type of A. abruptum BL. was a
mountain plant, differing in entire pinna-lobes, but
among other specimens from Java there is much
variation in the shape of pinna-lobes and it does
not seem possible to divide the specimens on this
character. Existing specimens do not show any
possible distinctions in characters of the reduced
basal pinnae, or of lower normal pinnae. As with
several other species of this family from Java,
good new specimens are needed.
The type of P. christelloides at Kew lacks the
basal pinnae, but these are present on the BM
isotype and show that my description of 1973 was
inaccurate. The type differs chiefly from typical P.
truncata in the unusual abundance of short hairs
between the veins of both surfaces (but not on all
pinnae), also on indusia. Another plant from Mt
Kinabalu, cultivated at Kew, is identical. The
pubescence is very like that of Christella.
Allied species have been described from China,
430
and need to be critically compared with Malesian
specimens.
35. Pneumatopteris lawakii HOLTTUM, nom.
nov. — Polystichum truncatum GAUD. in Freyc.
Voy. Bot. (1827) 332.— Aspidium truncatum
GAUD. ibid. t. 10. — Type: GAUDICHAUD, Rawak
( = Lawak) I., W. New Guinea (P; B, FI).
P. glaberrima sensu HoL_truM, Blumea 21
(1973) 318, p.p.
Agreeing with P. truncata (POIR.) HOLTTUM in
the peculiar venation (which is well figured by
GAUDICHAUD), differing as follows: fronds much
smaller, and thinner; largest pinnae 15 x 1.8 cm;
veins 6-7 pairs; lower surface quite glabrous;
upper reduced pinnae 2.00.9cm, lowest
0.5 cm.
Distr. Malesia: West New Guinea.
Notes. Known from the type and MCKEE
NGF 1940 from Biak Island, from which the
details of the reduced pinnae are described (not
figured by GAUDICHAUD). The fronds of the type
of P. glaberrima (RICHARD) are of similar size
but the pinna-lobes are different. More specimens
are needed from West New Guinea.
36. Pneumatopteris michaelis HOLTTUM, Blumea
21 (1973) 313.—Type: M.-G. PRICE 317, Mt
Makiling, Luzon, 1050 m (K).
Caudex erect; stipe of type 17cm long,
minutely hairy; base of stipe to first normal pinna
50cm; reduced pinnae 4-5 pairs, uppermost
1.2 cm long, 0.8 cm wide at truncate base which is
symmetric or slightly wider on acroscopic side,
margins crenate. Lamina excluding reduced pin-
nae 60cm long; pinnae 20 pairs or more, rather
close together, texture firm; basal pinnae nar-
rowed towards their bases which are 8mm wide,
narrowed part very shallowly lobed. Largest pin-
nae 22cm long, 1.5-1.9cm wide; base broadly
cuneate; apex evenly attenuate, apical 2-2.5cm
subentire; edges lobed c. 1/3 towards costa; lobes
oblique, strongly truncate with broad teeth at
vein-ends and an acute forward-pointing tip; cos-
tules 4-5 mm apart; veins 7-8 pairs, slender, con-
colorous, slightly prominent, 1; pairs anastomos-
ing, 1 pair to sides of sinus-membrane; lower
surface quite glabrous including sinus-membranes
and margin, surface between veins slightly pus-
tular; upper surface of rachis and costae rather
sparsely hairy, hairs pale, 0.5mm long. Sori in-
framedial on distal veins, supramedial on basal
ones, those on basal veins from adjacent costules
sometimes confluent; indusia thin, glabrous;
sporangia lacking glands; spores pale with a
moderate number of distinctly flat small wings.
Distr. Malesia: Philippines (Luzon: Mt Makil-
ing), 2 collections.
Note. Plants have been cultivated at Kew, and
have remained very distinct from a plant of P.
truncata from Mt Kinabalu both in size of plant
FLORA MALESIANA
[ser. II, vol. 1°
and shape of pinnae, and also in requiring a cooler
temperature for good growth.
37. Pneumatopteris nitidula (PRESL) HOLTTUM,
Blumea 21 (1973) 318.— Nephrodium nitidulum
PRESL, Epim. Bot. (1851) 46; HoLTTUM, Novit.
Bot. Univ. Carol. Prag. 1968 (1969) 40. — Cyclo-
sorus nitidulus (PRESL) COPEL. Fern FI. Philip.
(1960) 364, p.p. excl. syn. omn.— Thelypteris
nitidula (PRESL) REED, Phytologia 17 (1968)
296. — Type: CUMING s.n., Philippines (PRC).
Caudex short, erect; stipe c. 10cm long, pale,
bearing thin appressed scales; base of stipe to first
large pinna 30-40 cm; reduced pinnae to 7 pairs,
basal ones 7 X 7mm, almost orbicular, uppermost
2.5-3.0 cm long, 1.5cm or more wide, base trun-
cate, auricled or dilated both sides, distal part
lobed. Lamina excluding reduced pinnae to
100cm long; pinnae to 30 pairs or more, thin,
spreading at a wide angle to rachis; basal pinnae
not or little narrowed at their bases. Largest pin-
nae of type 24 x 2.2 cm (largest seen 34 cm long);
base truncate; apex almost evenly attenuate, nar-
row subentire distal part 2cm or more long; edges
lobed less than 3 way to costa on small fronds,
more than > on type; lobes slightly oblique, their
apices almost rounded on largest pinnae, sub-
truncate on smaller ones, projections at vein-ends
slight or lacking, cartilaginous margins narrow;
costules 4.5-5.5 mm apart; veins 8-10 pairs, slen-
der, concolorous, basal pair anastomosing, next
acroscopic vein to side of short sinus-membrane;
lower surface of rachis and costae bearing minute
erect hairs, similar hairs also + abundant on other
parts, surface between veins often not pustular;
hairs on upper surface of rachis and costae
sparse, pale, short. Sori inframedial, basal ones
not divergent; indusia bearing few to many aci-
cular hairs; sporangia with many rather large club-
shaped glands; spores light brown with many very
small wings.
Distr. Malesia: Philippines (Luzon to Min-
danao).
Ecol. A common species at low altitudes, often
near streams, sometimes in rather open places.
38. Pneumatopteris kerintjiensis HOLTTUM, Blu-
mea 21 (1973) 312.—Type: ALSTON 14149,
Sumatra, Sungei Kering, Kerintji (BM).
Dryopteris sumatrana sensu TROLL, Flora 128
(1933) 329-337.
Stipe 5 cm long; base of stipe to first large pinna
75 cm: reduced pinnae at least 10 pairs, uppermost
to 3.5x3.5cm, almost symmetrically triangular
with shallowly lobed edges and blunt tip, lowest
2x2cm. Lamina excluding reduced pinnae
120 cm long; pinnae 45-50 pairs; basal pinnae not
or little narrowed towards their bases which are
almost symmetrically dilated; texture firm. Lar-
gest pinnae 25 X2.5cm; base truncate; apex acu-
minate, not caudate; edges lobed 1/3-2/5 towards
1981]
THELYPTERIDACEAE (Holttum)
43]
costa; lobes not falcate, entire with rounded tips;
costules 4.5mm apart, at a wide angle to costa;
veins to 12 pairs, hardly prominent either side, 2
pairs anastomosing, next 15 pairs to sides of sinus-
membrane; minute hairs present on lower surface
of costae, costules and veins, surface between
veins strongly pustular; upper surface of costae
bearing short hairs, rest of pinnae glabrous. Sori a
little inframedial, lower ones not divergent; in-
dusia thin, glabrous; sporangia bearing capitate
hairs; spores light brown with many small wings.
Distr. Malesia: North-central Sumatra.
Ecol. In forest at 1150-1400 m; the type found
in a tea estate.
Note. TROLL’s photograph and his data about
mucilage hairs on scales clearly indicate the
present species. He refers to BEDDOME’s des-
cription of the reduced basal pinnae of Neph-
rodium molle var. major, but that description
refers to Christella papilio, q.v.
39. Pneumatopteris nephrolepioides (C. CHR.)
HOLTTuM, Blumea 21 (1973) 320. — Dryopteris
nephrolepioides C. CHR. Brittonia 2 (1937) 268, f.
1, c,d.— Thelypteris nephrolepioides (C. CHR.)
REED, Phytologia 17 (1968) 295. — Type: BRASS
5354, Mafulu, Central div. Papua, on limestone
1700 m (BM; NY).
Caudex short; stipe 3—4cm long, slender, gla-
brous, scales ovate, |!mm long. Lamina to
40cm long; pinnae c. 40 pairs; lower pinnae
gradually reduced and more widely spaced, lowest
2mm long; texture firm, drying light green. Lar-
gest pinnae 1.8xX0.6cm; base truncate to sub-
cordate, slightly auricled on acroscopic and roun-
ded on basiscopic side; apex rounded; edges
slightly sinuous to almost crenate; small hyda-
thodes each with a white scale near ends of veins,
just within the margin; veins 7-8 pairs in each
pinna, oblique, simple except the forked basal
acroscopic one, free, slightly prominent on upper
surface; lower surface of rachis bearing short
acicular hairs, sparse minute capitate hairs present
on costa and veins; upper surface of rachis
copiously hairy, very short erect acicular hairs
also present on whole upper surface of pinnae.
Sori medial on veins, small, exindusiate; sporan-
gia bearing very small capitate hairs (sometimes
3); spores light brown with many small wings.
Distr. Malesia: Papua New Guinea. Only
known from type.
Ecol. On limestone, at 1700 m.
40. Pneumatopteris ligulata (PRESL) HOLTTUM,
Blumea 21 (1973) 320. — Lastrea ligulata PRESL,
Epim. Bot. (1851) 35; CopeL. Fern FI. Philip.
(1960) 327.—Dryopteris ligulata (PRESL) O.
MiPZES wReveasGens Plje2 (189) sis aveAnvaR:
Handb. (1908) 184. — Thelypteris ligulata (PRESL)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941)
252. — Type: CUMING 74, Luzon (PRC; E, K).
Lastrea philippina PRESL, Epim. Bot. (1851)
36. — Type: CUMING 343, Zebu (PRC).
Nephrodium luerssenii HARR. J. Linn. Soc. Bot.
16 (1877) 29. — Dryopteris luerssenii (HARR.) C.
Cur. Ind. Fil. (1905) 276; v.A.v.R. Handb. (1908)
190. — Type: STEERE s.n. Bulukai Isl. (K).
Dryopteris immersa var. ligulata CHRIST,
Philip. J. Sci. 2 (1907) Bot. 208. — Type: CUMING
343, Zebu (P).
Dryopteris foxii CHRIST, Philip. J. Sci. 2 (1907)
Bot. 208; v.A.v.R. Handb. (1908) 814. — Lastrea
foxii (CHRIST) COPEL. Fern FI. Philip. (1960)
328. — Thelypteris foxii (CHRIST) REED, Phy-
tologia 17 (1968) 277.— Lectotype (HOLTTUM
1973): COPELAND 940, Mindanao (US).
Caudex short-creeping; stipe pale or + flushed
reddish, to 40cm long, scales near base to 5 xX
1mm bearing many short acicular hairs on outer
surface. Lamina very variable, in largest plants
70 cm or more long with c. 20 pairs of well-spaced
and oblique pinnae, basal pinnae of large fronds
commonly about half length of largest, sometimes
a smaller, more widely-spaced pair also; smaller
plants have fertile fronds 25cm long, three pairs
lower pinnae gradually reduced and more widely
spaced, lowest 1cm long. Pinnae of large fronds
to 23 x 3 cm; base unequally broadly cuneate with
acroscopic lobe longer than basiscopic; apex
caudate-acuminate; edges lobed to 0.5mm from
costa or more deeply; lobes 2.5-3mm_ wide,
oblique, sometimes slightly toothed at ends of
veins, widened a little above the base, apices
broadly pointed; sinuses between lobes broad with
rounded base and very small sinus-membranes;
costules 5mm apart, at 45° to costa; veins to 12
pairs, slender, pale, prominent both sides; lower
surface of rachis glabrous, of costae, costules,
veins and surface between veins bearing a vari-
able number of slender erect hairs, a few longer
hairs sometimes present on costules and veins,
surface between veins finely pustular; upper sur-
face of costae with pale hairs 0.5 mm long, rest of
surface bearing very short erect pale hairs. Sori
supramedial; indusia thin with short capitate hairs
and sometimes acicular hairs; sporangia bearing
capitate hairs. Pinnae of small fertile fronds c.
10x 1.5cm; lobes 2mm wide; costules 4mm
apart; veins to 9 pairs; sori sometimes almost
medial; indusia usually lacking acicular hairs.
Distr. Malesia: Philippines (Luzon to Min-
danao) and N. Moluccas (Talaud Is.).
Ecol. The smaller plants mostly on rocky
stream-banks (once on the bank of a rice-paddy),
the larger onces in forest, at low and medium
altitudes.
Notes. As reported by CHRIST (1907)
COPELAND at first distinguished D. foxii by its
small fronds, much-reduced basal pinnae and al-
most medial sori, but in all these characters there
is no sharp distinction (and the largest fronds in
herbaria usually lack the base, so that smaller
432
FLORA MALESIANA
[ser. II, vole?
basal pinnae might have been present). I suggest
that the distinction is one of habitat (as with
stream-bank forms of Pronephrium menisciicar-
pon). Middle pinnae of large fronds are similar in
aspect to those of Amphineuron immersum (BL.)
HoLtTtuM, but differ from the latter in very
oblique widely-spaced lobes, in venation and in
glandular hairs.
41. Pneumatopteris finisterrae (BRAUSE) HOLT-
TUM, comb. nov.—Dryopteris __ finisterrae
BRAUSE, Bot. Jahrb. 49 (1912) 20.— Lastrea
finisterrae (BRAUSE) COPEL. Gen. Fil. (1947) 138;
Philip. J. Sci. 78 (1951) 426. — Thelypteris finis-
terrae (BRAUSE) REED, Phytologia 17 (1968)
277. — Type: SCHLECHTER 18134, N.E. New
Guinea, Finisterre Mts (B; P).
Caudex erect; stipe densely short-hairy, basal
scales broad, appressed; base of stipe to first
normal pinna 35cm or more; reduced pinnae 5-6
pairs, all less than 2mm long. Lamina excluding
reduced pinnae to 40 cm long; pinnae to 18 pairs,
thin, basal pinnae not distinctive. Largest pinnae
7.5 x 0.9 cm; base unequally broadly cuneate; apex
acuminate; edges lobed to 0.5mm from costa;
lobes at 45° to costa, their tips rounded, entire or
at most slightly crenate; costules 3mm _ apart;
veins 4-6 pairs, oblique, hardly prominent, basal
ones both ending at margin above base of sinus;
lower surface bearing short erect pale acicular
hairs on all parts, also short capitate hairs or small
sessile glands; upper surface of rachis and costae
bearing short antrorse hairs, whole surface of
pinna bearing short erect acicular hairs. Sori
medial; indusia small, thin, long-hairy; sporangia
bearing short slender setae; spores with a rather
broad entire translucent wing and a few anas-
tomosing cross-wings.
Distr. Malesia: N.E. New Guinea (3 collec-
tions).
Note. This appears to be allied to P. costata. It
is aberrant in the slender setae on sporangia, and
needs further study.
42. Pneumatopteris versteeghii HOLTTUM, Blu-
mea 21 (1973) 321.—Type: VERSTEEGH BW
10259, W. New Guinea, Genifa Mts (L).
Caudex short-creeping, 2mm diameter, stipes
closely seriate on it; stipe 8 cm long, slender, pale,
scales very small and appressed. Lamina 15-
18cm long; pinnae to 20 pairs; basal pinnae
somewhat reduced, at least half as long as the
next pair; texture firm but translucent, light
olivaceous when dried. Largest pinnae 2 x 0.6 cm;
base very asymmetric, on basiscopic side very
narrowly cuneate, on acroscopic side truncate
with a dentate auricle 5-6 mm long; apex obtuse;
in middle lobed half-way to costa; lobes some-
times dentate at vein-ends; veins in basal auricle 4
pairs, in middle lobes 2 pairs, basal acroscopic
vein passing to base of sinus, basiscopic vein to
margin; lower surfaces quite glabrous; upper sur-
face with short hairs on rachis and a few on bases
of costae. Sori on basal acroscopic vein of each
group; indusia short-hairy; sporangia with capitate
hairs.
Distr. Malesia:
known from the type.
Ecol. In forest on steep rocky soil at 1100 m.
West New Guinea. Only
43. Pneumatopteris sumbawensis (C. CHR.)
HOLTTuM, Blumea 21 (1973) 323. — Dryopteris
sumbawensis C. CHR. in Rensch, Hedwigia 74
(1934) 231, t. vii, f. 1.—Type: RENSCH 578,
Sumbawa, Batu Dulang (B; BO).
Caudex unknown; stipe 2-3 cm long. Lamina
30cm long; pinnae 35-40 pairs; 6 pairs lower
pinnae gradually or subabruptly reduced, lowest
3 mm long; lower large pinnae a little narrowed to
their bases on basiscopic side; texture thin, drying
pale olivaceous. Largest pinnae 3.0 x 1.0cm, sub-
pinnate with basal acroscopic lobe quite free and
2-3 pairs of lobes separately adnate to costa, rest
deeply lobed, apex abruptly narrowed to a blunt
tip; costules to 3 mm apart; veins free, 3-4 pairs,
dark but not prominent beneath, basal acroscopic
one sometimes forked; lower surface of rachis
and costae bearing minute capitate hairs and very
short acicular ones, short capitate hairs present all
over lower surface of pinnae as in P. costata;
hairs on upper surface of rachis 0.3mm long,
shorter on costae, rest of pinna with sparse very
short acicular and capitate hairs. Sori medial to
supramedial, exindusiate; neither glands nor setae
on sporangia.
Distr. Malesia: Lesser Sunda Is. (Sumbawa),
only known from type.
Note. This is related to P. costata which also
occurs on Sumbawa, but has very different
pinnae.
44. Pneumatopteris keysseriana (ROSENST.)
HoL_ttuM, Blumea 21 (1973) 320.— Dryopteris
keysseriana ROSENST. Fedde Rep. 10 (Feb. 1912)
333; v.A.v.R. Handb. Suppl. (1917) 157. — Lastrea
keysseriana (ROSENST.) COPEL. Gen. Fil. (1947)
139; Philip. J. Sci. 78 (1951) 430.— Thelypteris
keysseriana (ROSENST.) REED, Phytologia 17
(1968) 286.— Type: KEYSSER 253, N.E. New
Guinea, Sattelberg (not seen).
Dryopteris schultzei BRAUSE, Bot. Jahrb. 49
(August 1912) 19; v.A.v.R. Handb. Suppl. (1917)
156. — Type: L. SCHULTZE 253, Sepik District
(B). — Fig. 11i-j.
Caudex short-creeping with tufted stipes; stipe
to 70cm long, pale except base, scales thin, ap-
pressed; reduced basal pinnae commonly 2-3
pairs, much smaller than normal ones, in one
specimen apparently 8 pairs. Lamina to 100cm
long; normal pinnae 20 pairs or more; basal pinnae
variably narrowed towards their bases, narrower
on basiscopic side where the basal lobe is obsolete
1981]
THELYPTERIDACEAE (Holttum)
433
or very small. Largest pinnae c. 25X4cm
(ROSENSTOCK reports 6cm wide); base un-
equally broadly cuneate, basal basiscopic lobe
shorter than acroscopic; apex acuminate with
cauda 34cm long; edges lobed to 1-2 mm from
costa with wide sinuses between lobes; lobes
slightly tapering, tips rounded to bluntly pointed,
edges near tips slightly crenate at vein-ends; cos-
tules 5-6 mm apart, at a wide angle to costa; veins
to 16 pairs, pale and prominent both sides, basal
pair ending at margin near together, at least in
distal part of pinna; lower surface of rachis bear-
ing very short hairs, similar hairs variably present
on lower surface of pinnae, surface between veins
slightly pustular; upper surface of costae bearing
short pale hairs, rest of pinna glabrous. Sori
supramedial; indusia small with short hairs;
sporangia sometimes with capitate hairs; spores
with many small wings. Chromosomes: n= 36
(T. G. WALKER).
Distr. Malesia: Eastern New Guinea.
Ecol. By streams in lowland forest and to
1000 m, common.
Note. Specimens from the type locality, named
by ROSENSTOCK, have been seen. The type of
Dryopteris schultzei differs chiefly in a greater
abundance of hairs on the lower surface, some of
them between veins.
45. Pneumatopteris caudata (HOLTTUM) HOLT-
TUM, Blumea 21 (1973) 321.— Pseudocyclosorus
caudatus HOLTTUM, Blumea 13 (1965) 133.—
Type: cult. Hort. Bot. Kew. 545/63, n. 12, origin
near Lae, N.E. New Guinea (K).
Caudex erect; stipe pale, basal scales small,
ovate; base of stipe to first normal pinna 15 cm or
more; reduced pinnae 2 pairs, 2-3mm_ long.
Lamina excluding reduced pinnae 30cm long;
pinnae 15 pairs; lower pinnae narrowed both sides
towards their bases. Largest pinnae 8 x 1.5 cm;
base strongly asymmetric with minute basal
basiscopic lobe, basal acroscopic lobe almost free
and not reduced; apex abruptly narrowed to a
cauda 2.5cm long, 2mm wide; edges lobed to
1mm from costa; lobes at 45° to costa, entire or
slightly toothed at vein-ends, apices rounded;
costules 3.5 mm apart; veins 9 pairs, oblique, pale
and prominent on lower surface, basal acroscopic
vein ending at sinus, basiscopic at margin above
base of sinus; lower surface of rachis, costae and
costules bearing minute acicular hairs, short capi-
tate hairs on costules and veins, surface between
veins slightly pustular; upper surface hairy on
tachis and costae only. Sori supramedial; indusia
short-hairy; no glands seen on sporangia; spores
pale with many small wings. Chromosomes: n=
36 (S. K. Roy 1965).
Distr. Malesia: Papua New Guinea.
known from the type.
Note. This is certainly nearly allied to P. keys-
seriana and from a neighbouring locality, but
Only
plants of the two cultivated at Kew remained very
distinct.
46. Pneumatopteris deficiens HOLTTUM, Blumea
21 (1973) 321. —Type: POSTHUMUS 3183, Flores
(BO; K).
Caudex short, erect; stipe to 40cm long, pale,
glabrous, covered with thin small scales when
young. Lamina c. 35 cm long; free pinnae 8 pairs,
texture firm; basal pair of pinnae + reduced, in
most cases only 5mm long; basal large pinnae
narrowed evenly towards their bases. Largest
pinnae 12 x2.2cm; base unequally cuneate; apex
acuminate, dentate to tip; edges lobed to 1mm
from costa; lobes oblique, acute, edges with slight
teeth at ends of veins; costules 5-6mm apart:
veins to 8 pairs,‘pale and prominent on both sides,
basal acroscopic vein ending beside the short
sinus-membrane, basiscopic vein to margin above
base of sinus; lower surface of costae sometimes
with minute acicular hairs, short hairs present on
margins of lobes, surface between veins slightly
pustular; upper surface hairy on costae, rest of
pinna glabrous. Sori a little inframedial; indusia
with a few very short hairs; no glands seen on
sporangia.
Distr. Malesia:
several collections.
Ecol. At 1300m, on moist banks of earth in
wet shady ravines.
Lesser Sunda Is. (Flores),
47. Pneumatopteris mingendensis (GILLI) HOLT-
TUM, comb. nov. — Lastrea mingendensis GILLI,
Ann. Nat. Hist. Mus. Wien 81 (1978) 24. — Type:
GILLI 265, N.E. New Guinea, Chimbu District (W).
Stipe 15cm long, pale, glabrous except in
groove; base of stipe to first normal pinna 30cm;
reduced pinnae 2 pairs, widely spaced, upper ones
2 cm long with basal auricle 7 mm long, lower ones
7mm long with auricle 4mm. Lamina excluding
reduced pinnae 60cm long; basal normal pinnae
somewhat reduced. Largest pinnae 13X3cm,
lobed to less than 1mm from costa, lobes on
basiscopic side shorter and more oblique than on
acroscopic, basal acroscopic lobe 2.0cm long,
basal basiscopic lobe much shorter and 10mm
from base of pinna; apex narrowly caudate, cauda
2-2.5 cm long; costules 5-6 mm apart; veins 10-12
pairs, pale and prominent both sides, basal veins
ending far apart at the margin; lower surface of
rachis and costae bearing many acicular hairs
hardly 0.1 mm long, sparse similar hairs on cos-
tules, surface between veins not distinctly pus-
tular; upper surface of rachis and costae bearing
hairs a little longer than those on lower surface,
no other hairs present. Sori medial; indusia small,
glabrous; capitate hairs present on some sporan-
gia; spores pale with many wings of irregular
shape.
Distr. Malesia: Papua New Guinea. Only
known from the type.
434
Ecol. At 2100 m in Pandanus forest.
Note. GILLI reported that indusia are absent,
but though small they are certainly present.
48. Pneumatopteris eburnea HOLTTUM, sp. nov.
Lamina usque 35cm longa; pinnae usque 18-
jugatae, rigidae, maximae 6.5 1.6cm, infimae
leviter redactae, inferiores plurijugatae basin ver-
sus angustatae, omnes profunde lobatae; venae
liberae, subtus crassae, prominentes, pallidae;
pagina inferior perfecte glabra; indusia parva,
glabra. — Type: JERMyY 14132, Sarawak, Gunong
Mulu, 1500-1600 m (BM).
Caudex short-creeping; stipe 20-40cm long,
pale above base, glabrous, basal scales broad,
thin, not persistent; reduced pinnae lacking.
Lamina to 35cm long; pinnae to 18 pairs, rigid
and brittlhe when dry; basal pinnae slightly
reduced, much narrowed towards their bases on
basiscopic side, less so on acroscopic side which
is slightly auricled; several successive pairs of
pinnae less narrowed at their bases. Largest pin-
nae 6.5 x 1.6cm, sessile, aerophores not elongate;
apex short-acuminate; edges lobed to 1.5 mm from
costa, lobes hardly falcate, their tips rounded,
edges thickened; costules to 3 mm apart, almost at
right angles to costa; veins to 10 pairs, all very
oblique, thick, pale and prominent on lower sur-
face, concolorous on upper, basal veins both end-
ing just above base of sinus, sinus-membrane not
evident; lower surfaces quite glabrous, often with
residual narrow scales; upper surface of rachis
with sparse hairs 0.3mm long, many hairs 0.4—
0.5 mm long on edges of costal groove, no other
hairs. Sori medial; indusia small, thin but firm,
glabrous; sporangia lacking glands, a hair of 3
cells sometimes present on stalks of sporangia;
spores pale with many small wings.
Distr. Malesia: Borneo (Sarawak: Mt Mulu),
known only from type.
Ecol. In gullies on limestone at 1500-1600 m.
49. Pneumatopteris excisa (HOLTTUM) HOLT-
TUM, Blumea 21 (1973) 321. — Pseudocyclosorus
excisus HOLTTUM, Blumea 13 (1965) 133.—
Type: MILLAR & HOLTTUM NGF 18623, N.E.
New Guinea, Eastern Highlands (K; LAE, L).
Caudex erect; base of stipe to first normal pinna
25cm; reduced pinnae 2-3 pairs, very small,
remote from basal normal pinna. Lamina to 40 cm
long; pinnae to 25 pairs; texture firm. Largest
pinnae of type 10x 1.5cm, of another collection
19x 3cm, lobed to less than 0.5mm from costa,
lobes with rounded tips and distinct teeth at vein-
ends, several lobes reduced near base of pinna
especially on basiscopic side, basal acroscopic
lobe not free; apex of pinna caudate-acuminate,
cauda 2.5cm long, crenate; costules 3 mm apart;
veins of type 8 pairs, pale and prominent both
sides, basal veins usually both ending at margin
above base of sinus; lower surface of rachis and
FLORA MALESIANA
[ser. II, vol. 1°
costae bearing very short acicular hairs, scattered
similar hairs on costules and a few short capitate
ones, surface between veins slightly pustular,
glabrous; upper surface short-hairy on costae, a
few hairs present on costules and veins. Sori
medial, filling lower surface at maturity; indusia
minute, glabrous; no glands seen on sporangia;
spores with many very small wings.
Distr. Malesia: N.E. New Guinea, several col-
lections at 1300-2000 m.
50. Pneumatopteris regis (COPEL.) HOLTTUM,
Blumea 21 (1973) 322. — Dryopteris regis COPEL.
Univ. Cal. Publ. Bot. 18 (1942) 220.— Lastrea
regis COPEL. Philip. J. Sci. 78 (1951) 425, pl.
19.— Thelypteris regis (COPEL.) REED, Phy-
tologia 17 (1968) 308.—Type: C. KING 486,
Papua, mountains behind Wedan (MICH).
Stipe not seen. Lamina 120 cm long including 6
pairs of deeply lobed basal pinnae 1-1.5 cm long
and wide; texture firm. Largest pinnae 15x
2.5cm; base truncate, a little dilated, wider on
acroscopic side; apex acuminate; edges lobed to
Imm from costa; lobes 134mm, entire, ends
rounded; costules 5 mm apart, at more than 60° to
costa; veins 8 pairs, oblique, basal veins both
ending at the margin above base of sinus; lower
surface glabrous; upper surface of pinnae
glabrous apart from many dark brown hairs on
costae, hairs on rachis pale. Sori small, in-
framedial; indusia apparently absent (all sori are
old, and indusia might have fallen); no glands seen
on the few remaining sporangia; spores not seen.
Distr. Malesia: Papua New Guinea. Only
known from the type.
51. Pneumatopteris boridensis HOLTTUM, sp. nov.
Pinnae basales redactae 6-jugatae, alternae,
suprema 3X 1.3cm, infima 0.9 cm longa; pinnae
normales 17x2.3.cm, fertiles 2/3 costam versus
lobatae; venae 9-10-jugatae, infimae ambae
membranam sinus tangentes vel in pinnis sterilibus
infra sinum junctae; indusia magna, tenuia,
hirsuta. —Type: FOREMAN & VINAS LAE
60262, S.E. New Guinea, Port Moresby Subdistr.,
near Boridi village (K).
Caudex short-creeping; stipe 22 cm long, pale,
glabrous; base of stipe to first normal pinna 55 cm;
reduced pinnae 6 pairs, alternate, lowest 9mm
long and distinctly auricled, uppermost 3 x 1.3.cm
with slight basal auricle. Lamina excluding
reduced pinnae 70cm long, texture thin; basal
pinnae neither narrowed towards their bases nor
auricled. Largest fertile pinnae 17 x 2.3 cm (sterile
2.5cm wide at base); base very broadly cuneate;
apex caudate-acuminate, cauda to 3cm _ long,
crenate; edges lobed to 3.5-4mm from costa in
fertile pinnae, less deeply in sterile; lobes slightly
toothed at vein-ends especially where fertile; cos-
tules 5 mm apart; veins 9-10 pairs, lowest pair on
fertile pinnae just touching base of the short
1981]
THELYPTERIDACEAE (Holttum)
435
sinus-membrane, on fertile pinnae sometimes uni-
ting below the membrane; lower surface glabrous;
upper surface of rachis bearing pale hairs more
than 0.5mm long, shorter ones on costae, no
others. Sori inframedial, basal ones not divergent;
indusia large, thin, shrivelling when old, bearing in
the middle many slender hairs 0.5mm long; no
glands seen on sporangia.
Distr. Malesia: Papua New Guinea,
known from the type.
Ecol. By stream at 1190 m.
only
52. Pneumatopteris imbricata HOLTTUM, Blumea
21 (1973) 322. Type: BROOKS 17775, Amboina,
Hila, 200 m, on rock by river (BM; BO, L).
Caudex “short and stumpy” (Womersley &
Whitmore); stipe Scm long, basal scales thin,
2mm wide at base; base of stipe to first normal
pinna 45-60 cm; reduced pinnae many pairs, | cm
long, deeply lobed, both basal lobes large and
lobed. Lamina to 110cm long, thin, translucent;
pinnae 35 pairs, all opposite; basal normal pinnae
with enlarged dentate basal lobes, the acroscopic
lobe (when dried in a press) overlapping the lower
surface of the rachis, basiscopic lobe overlapping
the upper surface. Largest pinnae 24x 3.0 cm;
base truncate, + auricled on the acroscopic side;
apex narrowly acuminate, serrate to the tip; edges
lobed to | mm from costa, sinuses between lobes
wide, rounded at their bases; lobes oblong,
oblique, slightly falcate, their tips broadly roun-
ded, entire, little over 3mm wide when dried;
costules 4.5-5.5 mm apart; veins to 12 pairs, basal
ones both to margin above base of the sinus; lower
surfaces glabrous apart from a few short hairs
distally on costae and on sinus-membranes and
margin; upper surface densely brown-hairy on
rachis and costae, rest glabrous. Sori small,
somewhat inframedial, somewhat impressed on
upper surface, exindusiate; sporangia of Guadal-
canal specimen bearing short yellow capitate
hairs; spores with a rather narrow median wing
and some cross-wings.
Distr. Solomon Is. (Guadalcanal) and E.
Malesia: Moluccas (Halmahera & Amboina).
Note. The sori of the type are young and the
sporangia do not show glands; The Halmahera
specimen (PLEYTE 370) has old sori with
shrivelled glands on sporangia. The Guadalcanal
specimen (WOMERSLEY & WHITMORE BSIP
1038) is a little smaller than the type but otherwise
closely similar.
53. Pneumatopteris petrophila (COPEL.) HOLT-
TUM, Blumea 21 (1973) 322.— Dryopteris petro-
phila CopeEL. Univ. Cal. Publ. Bot. 18 (1942)
220. — Lastrea petrophila COPEL. Philip. J. Sci. 78
(1951) 424, pl. 18. — Pseudocyclosorus petrophilus
(COPEL.) HOLTTUM, Blumea 13 (1965) 133. —
Thelypteris petrophila (COPEL.) REED, Phy-
tologia 17 (1968) 303.— Type: BRASS 11326, W.
New Guinea, Bele River (MICH; BM, L).
Caudex short, erect; stipe 46cm long, pale,
minutely hairy, basal scales small, appressed.
Lamina 40cm long, texture very firm; pinnae c.
20 pairs, almost opposite, lower 8-10 pairs
reduced, rather abruptly and then gradually,
lowest 33mm, intermediate pinnae lobed and
distinctly auricled on the acroscopic base. Largest
pinnae of type 4cm long, 8-9mm wide above
base (other specimens to 9 x 1 cm); base subtrun-
cate and + auricled on acroscopic side, narrower
and rounded on basiscopic; apex short-pointed
(acuminate on larger specimens); edges lobed 1/2
towards costa; lobes falcate, narrowed a little to
blunt tips, entire; costules 3mm apart, oblique;
veins 3-5 pairs, slender and prominent, basal
acroscopic vein ending beside sinus-membrane,
basiscopic one at margin above base of sinus;
lower surface of rachis copiously short-hairy,
minute acicular and capitate hairs on costae, cos-
tules and veins, surface between veins pustular
and bearing minute capitate hairs; upper surface
of costae short-hairy (hairs brown on larger spe-
cimens), sparse short hairs present between veins.
Sori medial or inframedial, exindusiate; sporangia
bearing small colourless capitate hairs; spores pale
with many small wings. Chromosomes: n= 36
(T. G. WALKER).
Distr. Malesia: New Guinea. Besides type,
several collections in N.E. New Guinea.
Ecol. On wet rocks at c. 2000 m.
Note. This is related to P. latisquamata; I
found both near Edie Creek.
54. Pneumatopteris walkeri HOLTTUM, Blumea 21
(1973) 323.—Type: T. G. WALKER 9980, N.E.
New Guinea, Eastern Highlands, Waisa (BM).
Caudex short, erect or suberect; stipe 20cm
long, pale, minutely hairy, basal scales thin, ap-
pressed; base of stipe to first normal pinna 45—-
50 cm; reduced pinnae 6-8 pairs, 2-3 mm long, with
1-2 pairs transitional to normal pinnae. Lamina
55cm long, excluding reduced pinnae; pinnae 20
pairs; texture thin, drying pale olivaceous. Largest
pinnae 11x 1.9cm; base asymmetric, truncate on
acroscopic side, narrowly cuneate on basiscopic
side with reduced basal lobe; apex caudate-acu-
minate, cauda 3cm long, serrate; edges lobed to
0.5mm from costa; lobes on acroscopic side of
pinna longer than on basiscopic and almost at
right angles to costa, lobes on basiscopic side at
45°, all lobes + dentate near apices; costules 4mm
apart; veins in acroscopic lobes to 9 pairs, in
basiscopic lobes sometimes fewer, prominent both
sides, pale beneath, basal veins both running to
margin above base of sinus; lower surface of
rachis bearing copious minute acicular hairs, rest
glabrous, pustular between veins; upper surface
with short hairs on rachis and costae only. Sori
medial; indusia thin, glabrous; no glands seen on
sporangia.
[ser. II, vol. 1°
436 FLORA MALESIANA
Distr. Malesia: Papua New Guinea. Only
known from the type.
Doubtful
Aspidium glaberrimum RICHARD, Sert.
Astrolab. (1834) xviii.— Type: D’URVILLE, Prt
Dorei, N.W. New Guinea (P).
There is a photograph at BM of the type, but I
did not find the specimen at Paris. I saw another
specimen so named, collected by D’URVILLE in
New Ireland in 1827. Both specimens differ from
the type of Polystichum truncatum GAUD. in
having entire pinna-lobes. They are very like small
plants of Pneumatopteris sogerensis but the spe-
cimen from New Ireland has capitate hairs on the
sporangia. A specimen from the Markham Valley,
N.E. New Guinea (WAKEFIELD 1505, BM) is
about the same size as the type and has rather
large club-shaped glands on the sporangia, but has
also rather numerous short hairs on all parts of the
lower surface of pinnae.
17. SPHAEROSTEPHANOS
J. SM. in Hook. Gen. Fil. (1839) t. 24; CopEL. Univ. Cal. Publ. Bot. 16
(1929) 60; CHING, Sunyatsenia 5 (1940) 240; COPEL. Gen. Fil. (1947) 144;
emend. HOLTTUM, Blumea 19 (1971) 39; Kalikasan 4 (1975) 47; Allertonia 1
(1977) 201. — Thelypteris subg. Sphaerostephanos K. Iwats. Mem. Coll.
Sci. Univ. Kyoto B, 31 (1964) 32.
Mesochlaena R. BR. in Benn. & Br. Pl. Jav. Rar. (1838) 5, nom. illeg.; J.
SM. in Hook. J. Bot. 3 (1840) 18; C. Cur. Ind. Fil. (1905) xxii.
Proferea PRESL, Epim. Bot. (1851) 259; HoLtruM, Novit. Bot. Inst. Bot.
Univ. Carol. Prag. 1968 (1969) 48.
Cyclosorus sensu CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 162,
p.p.; sensu COPEL. Gen. Fil. (1947) 140, p.p.; sensu HOLTTUM, Rev. FI.
Mal: 2 (1955) 255; p-p.
Theylpteris subg. Glaphyropteridopsis sect. Neocyclosorus K. IwaTs.
Mem. Coll. Sci. B, 31 (1964) 30, quoad spec. typ.
Thelypteris subg. Pneumatopteris sect. Macrocyclosorus K. Iwats. lL.c.
34, quoad spec. typ. — Fig. 1a—h, q, 12, 13.
Caudex in most species erect or short-creeping, rarely long-creeping or
scandent; scales usually narrow, thin, bearing superficial acicular hairs;
much-reduced pinnae present at base of fronds, the transition downwards
to these from normal pinnae usually abrupt but in some cases quite gradual
(a few species are included which have no reduced basal pinnae but have
deeply lobed normal pinnae with glands typical of this genus); aerophores
at pinna-bases often swollen, in some species 1-2 mm long (very young
fronds then covered with mucilage); apex of frond rarely pinna-like; pinnae
in most species lobed (in some very deeply) or crenate, in a few subentire;
sinus-membrane almost always distinct, of varying length; veins anas-
tomosing, but in a few species just meeting at the sinus or ending above it;
sessile spherical yellow glands usually present on parts of the surface of
pinnae or on indusia or sporangia, in a minority of species quite lacking;
short capitate hairs lacking; acicular hairs almost always present on both
surfaces of costae and costules, lacking on the lower surface in a few
species, often present also between veins on either or both surfaces, either
erect or appressed, rarely if ever septate; lower surface rarely pustular
when dry; sori usually round, in some species (indusiate or not) more or
less elongate; indusia usually present; sporangia often bearing yellow
1981] THELYPTERIDACEAE (Holttum) 437
glands distally, less often setae, usually with a gland-tipped hair on the
stalk; spores usually bearing many small wings, in some species a con-
tinuous wing with cross-wings.
Type species: Sphaerostephanos asplenioides J. Sm. = S. polycarpus (BL.)
COPEL.
Distr. Throughout Malesia (152 spp.), tropical mainland Asia (12 spp.), Mascarene Islands and E.
Africa (4 spp.), Australia and the Pacific (17 spp., only 1 reaching Tahiti).
Ecol. Almost all are forest plants. Nearly all those adapted to open places have long-creeping
rhizomes and are widely distributed; allied to one of these are a few species in New Guinea and the
Solomons which have a slender scandent caudex, not known in any other genus of the family.
Cytol. Base chromosome number 36; 8 spp. examined, all diploid.
Taxon. JOHN SMITH’s name Sphaerostephanos refers to the yellow glands which fringe the
indusium of the type species. COPELAND and CHRISTENSEN treated this name as feminine, but as the
Greek stephanos is masculine, I regard the compound name also as masculine. SMITH’s original
specimen had broken indusia (carefully illustrated by FRANCIS BAUER) from which he gained an
erroneous idea of their structure. Soon afterwards SMITH received from ROBERT BROWN a better
specimen, and his comments on it were published in 1840 under BROWN’s name Mesochlaena which
SMITH regarded as having priority; he then realized that it differed from Nephrodium (as interpreted by
SCHOTT) only in having an elongate sorus. BROWN’s name Mesochlaena had been published in 1838
with a very slight description and without any indication of its status, so that it cannot be regarded as a
valid generic name, as pointed out by COPELAND in 1929. The first good illustration of the sorus was
published by METTENIUS (Fil. Hort. Lips. t. 18, fig. 13. 1857) with the name Mesochlaena javanica; this
was changed to Nephrodium javanicum by HOOKER who published an excellent illustration of a whole
plant (Fil. Exot. t. 62. 1859), with Aspidium polycarpon BL. as a synonym and reference to SMITH’s
comments of 1840.
When BEDDOME published the Supplement to his Handbook (1892) he also reverted to the genus
Nephrodium for Aspidium polycarpon BL., and described an additional Malayan species N. larutense
which also has elongate sori. In Index Filicum (1905) CHRISTENSEN retained the generic name
Mesochlaena for M. polycarpa but transferred BEDDOME’s N. larutense to Dryopteris. VAN ALDER-
WERELT accepted the generic name Mesochlaena in his Handbook (1908), transferring N. larutense to it
and later adding two more species. In the third Supplement to Index Filicum CHRISTENSEN followed
COPELAND in adopting the name Sphaerostephanos and transferred to it v.A.v.R.’s names.
The species thus brought together in Sphaerostephanos are not a closely allied group, and there are
others (e.g. S. norrisii) which have slightly elongate sori, so that no clear line can be drawn between
species with elongate and those with round sori. For these reasons, in 1955 I transferred the Malayan
species with elongate sori to Cyclosorus, a genus which I accepted in the sense of CHING (1938), though
commenting on the unsatisfactory nature of generic delimitation in the family Thelypteridaceae at that
time. CHING however had misinterpreted the sorus of the type species of Sphaerostephanos and had
wrongly reported the spores as trilete, for which reasons he proposed for it a new family (Sunyatsenia 5,
1940, 240).
When I had made a preliminary study of all known species of Thelypteridaceae in preparation for the
present work, I made a survey of all existing generic names and found that Sphaerostephanos was the
oldest name for a large group of mainly Malesian species most of which do not have elongate sori. For
this reason in 1971 I published a revised concept of the genus, which is here adopted. Species of this
genus were variously placed in IWATSUKI’s classification of 1964, which makes no reference to the
majority of Malesian species; I cite above only infrageneric taxa which he typified by Malesian species.
The main characters of this genus are the presence of much-reduced basal pinnae and of sessile
non-resinous spherical yellow glands of an easily recognized type; these glands are not destroyed in the
process of drying specimens for the herbarium as those of Coryphopteris and Amphineuron often are.
The species can be arranged in a series beginning with those that have glands on both surfaces of pinnae
and on indusia and sporangia, then those lacking glands on the upper surface (there are no species
bearing glands on the upper surface and not on the lower), those with glands only on sori and finally
those with no glands at all. Species without glands agree with Pneumatopteris in having reduced basal
pinnae but have a much greater complement of acicular hairs on all parts (especially on sporangia); they
lack also short capitate hairs and the peculiar pustules on the lower surface of dried fronds which are
characteristic of Pneumatopteris.
I also found that some species had very few reduced pinnae and that there is not a sharp distinction in
this character between Sphaerostephanos and Pronephrium. This matter is discussed under Proneph-
rium; I believe that the great majority of species belong clearly to one genus or the other.
438 FLORA MALESIANA [ser. II, vol. 1°
Fig. 12. Sphaerostephanos polycarpus (BL.) COPEL. a. Reduced pinnae, x 3; b. venation and position of
sori, X 3; c. sorus, X 12. — S. larutensis (BEDD.) C. CHR. d. Venation, sinus-membrane and sori, X 3; e.
sorus, X 12. — S. norrisii (ROSENST.) HOLTTUM. f. Base of lower pinna, x 1.5; g. venation and sori,
x 3; h. sorus, x 12.— S. pilososquamatus (v.A.v.R.) HOLTTUM. i. Base of sterile pinna, X 1.5; j. base
of fertile pinna, x 1.5; k. sorus, x 12. — S. hispidifolius (v.A.v.R.) HOLTTUmM. I. Middle pinna, x 1; m.
venation and sori, x 8.— S. inconspicuus (COPEL.) HOLTTUM. n. Base of lower pinna, X 2; 0. upper
pinna, X 1.5; p. venation and sori, x 4 (a—b YAPP 215, c MATTHEW S.n., d—e ERNST s.n., f KUNSTLER
2360, g-h MATTHEW s.n., i-k cult. Kew, |-m G. F. HOSE 91, n-p P. S. SHIM SAN 81677).
1981] THELYPTERIDACEAE (Holttum) 439
As the sessile spherical yellow glands are the most distinctive feature of the genus, I regard the
development of these on all parts of the frond as a primitive character. Assuming also that the original
species had an erect caudex, S. polycarpus might be regarded as a prototype for the genus. It is one of
the largest species vegetatively. But it is not a forest plant (a reason for its wide distribution) and it has
deeply lobed pinnae, whereas the most widely distributed Malesian forest species with glands on both
surfaces is S. penniger, which has shallowly lobed pinnae with several pairs of anastomosing veins; it is
allied to the arborescent S. arbuscula (WILLD.) HOLTTUM of Southern India and the Mascarene Islands. So
I suggest that the primitive Sphaerostephanos had shallowly-lobed pinnae with anastomosing veins, and
that S. arbuscula is the nearest surviving species. Several New Guinea species have a slender erect
caudex and shallowly-lobed pinnae (e.g. S. arfakianus) but have no glands on the upper surface. At least
it can be said with some certainty that (as in Christella) the few species which have free veins are
probably not primitive but represent a secondary development occurring locally in a few areas (e.g. S.
inconspicuus in Borneo, S. novoguineensis in New Guinea and S. pycnosorus in Samoa); they do not
form a coherent natural group.
A few species have quite gradually reduced basal pinnae (e.g. S. hastatopinnatus and S. arfakianus,
both with an erect caudex) but most have an abrupt (or almost abrupt) transition from normal to reduced
pinnae at the base of the frond. A few species which have many reduced pinnae are somewhat
intermediate, showing first a gradual reduction downwards and then an abrupt transition to much smaller
pinnae (e.g. S. hispiduliformis). I suggest that the condition of gradual reduction is the more primitive
arrangement, as it alone occurs in Cyathea. Apart from Sphaerostephanos and Pneumatopteris, which
are predominantly Malesian in distribution and certainly allied, most other genera have basal pinnae
gradually or not at all reduced.
The species with long-creeping rhizome are S. unitus, a gland-bearing species very widely distributed
with distinct eastern and western varieties, and S. invisus which lacks glands and is mainly distributed in
the Pacific (its distribution is similar to that of Pneumatopteris costata). S. hirtisorus (C. CHR.)
HOLTTUM in N.E. India and W. China appears to be related to S. invisus.
The following key is based on the distribution of glands on the surfaces of pinnae and on sori. I
believe that this represents an evolutionary trend, but within it there are many separate bifurcations
which, in such a complex group, are not easy to discern. For example, there is a group of species which
are related to S. stipellatus and have closely appressed hairs on the lower surface of costae and
costules: this character may be associated with elongate aerophores and lack of glands on the lower
surface of pinnae (as in S. stipellatus itself) or with varying combinations of these and other characters.
Varying trends may lead on to other groups (e.g. elongate aerophores may be associated with erect hairs
on the lower surface of costae) and I cannot see how to follow all the trends or delimit sub-groups. It
seems to me evident that the present condition of the genus is due to rapid and complex recent
evolution, especially in New Guinea, resulting in a difficulty in the delimitation of species, with some
necessary inequality of treatment. The destruction of Malesian forests may put a stop to this process or
alter it in unpredictable ways.
In some species where glands are not abundant there seems to be some variation in their occurrence;
in such cases a species may appear in more than one place in the key.
KEY HOV HE SPECIES
Glands present between veins on upper surface of pinnae.
5 Six or more pairs of lower pinnae gradually reduced . . bak, Uwe Sgn eee & Bel Sxdebilis
2. Transition to reduced basal pinnae (if any) abrupt or subabrupt.
3. Sori elongate.
4. No basalireduced pinnae: .0 |.) =) hres.) oe Dee 2 ein Se ee Ssndestiens
4. Basal reduced pinnae present.
5. Middle pinnae sessile : wi ea hs 25 Ss 8) 2 232Sapolycarpus
5. Middle pinnae with stalks 3- 10n mm ane Sicmal eel | ee ae. 2 3 eS zeminens
3. Sori not elongate.
6. Sori exindusiate.
7. Basal veins quite free suse 3S. ob: 445 meeebatbdcetenetoo es Ble aaa
7. Basal veins anastomosing.
8. Pinnae to 10x 1.5cm; reduced pinnae at least 5 pairs . . . . . . . . 6,8. stenodontus
8. Pinnae to 21 x 2.5cm; reduced pinnae 2-3 pairs . ... . . =... . 7.S. flavoviridis
6. Sori indusiate.
9. Reduced pinnae c. 30 pairs é 5 Qee iG) Sol bl et Pea erees
9. Reduced pinnae rarely more than 12 pairs.
10. Lower surface of rachis and costae quite glabrous, or with very short hairs on costae only.
440 FLORA MALESIANA (ser. II, vol. 1°
FF
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Meda ‘
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feb)
ion
UN WRERAN laden netic
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nig mi 9.4 4S RA APN LAE RUS RW A NS Nat gb NSC ‘Sis Da niddl
7 "
AN \ ) . *
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—
Fig. 13. Sphaerostephanos alatellus (CHRIST) HOLTTUM. a. Base of fertile pinna, x 13; b. base of sterile
pinna, x li; c. fertile pinna-lobes, x 4. — S. archboldii (C. CHR.) HoLTTum. d. Base of frond, x 1; e.
pinna-lobes, x 6. — S. baramensis (C. CHR.) HOLTTUM. f. Base of pinna, x 13; g. lobes of fertile pinna,
x 3.— S. batacorum (ROSENST.) HOLTTUM. h. Base of frond, x3; i. base of pinna, x2; j. lobes at
middle of pinna, x 4; k. costa and costule, with appressed hairs, and sorus, = 12 (a-c PULLEN 8258,
d—e BRASS 30856, f-g HOLTTUM 1, h-k MATTHEW 946).
1981] THELYPTERIDACEAE (Holttum) 44]
11. Fronds dimorphous; reduced pinnae all minute owes eee ees bye esSsalatellus
11. Fronds not dimorphous; reduced pinnae conspicuous . . . . ». . 10.8. daymanianus
10. Hairs present throughout lower surface of rachis and costae.
12. Sporangia copiously setiferous, rarely also with a gland.
13. Pinnae c. 7 pairs, to 13 X2.7 cm eee al Steers i 2te }-.. eee Ge peiioes,
13. Pinnae, if only 7 pairs, much smaller.
14. One pair of veins anastomosing, next pair to sinus-membrane . . 11. S. appendiculatus
14. 132 pairs anastomosing, b pairs to sinus-membrane . . . . 12.S. pseudomegaphyllus
12. Sporangia usually bearing glands, sometimes with 1-2 setae.
15. Reduced pinnae 0-3 pairs.
16. Pinnae not more than 10 pairs, reduced ones usually lacking . . . . 13.S.diversilobus
16. Pinnae at least 20 pairs, reduced ones always present.
17. Pinnae lobed not more than half-way to costa ee | eee 42S idichrotrichus
17. Pinnae lobed fully; . . . Sa eres it? ISAS cyestioiatas
15. Reduced pinnae on well-grown plants at least 5 pairs.
18. Pinnae not more than 4cm wide, their lobes not acute.
19. Pinnae crenate to subentire.
20. Pinnae subentire; sinus-membrane hardly detectable . . . . . .16.S. morotaiensis
20. Pinnae crenate, at least distally; sinus-membrane evident.
21. Pinnae to 3.5 x 1.0cm; reduced pinnae all very small . . . . . . 17.8. kalkmanii
21. Pinnae (at least sterile ones) to 7cm or more long, upper reduced pinnae at least 7 mm
long.
22. Pinnae to 12-15 cm long; sinus-membrane distally widened and translucent.
23. Bases of pinnae cuneate, not auricled nor dilated; costules at 45° to costa
18. S. perglanduliferus
23. Bases of pinnae truncate, lower ones auricled and dilated; costules at c. 60° to costa
19. S. veitchii
22. Pinnae to 8 cm long; sinus-membrane distally not widened nor translucent 20. S. lamii
19. Pinnae lobed at least 3 towards costa.
24. Nerophores, clongatega b-ieclt at okie he ee eek oe 21S: hirsutus
24. Aerophores not elongate.
25. Bases of pinnae, at least lower ones, dilated both sides.
26. Hairs on lower surface of rachis and costae erect, to | mm long on sterile fronds
22. S. acrostichoides
26. Hairs on lower surface of rachis and costae shorter, antrorse or appressed
23. S. melanorachis
25. Bases of pinnae not dilated.
27. Only | pair of veins truly anastomosing, second acroscopic vein sometimes touching
sinus-membrane.
28. Basal pinnae much narrowed at their bases . .. . . . . 24.S.heterocarpus
28. Basal pinnae not or little narrowed at their bases.
29. Reduced pinnae all very small ee dame 2 eel = Sq 2548S sisomorphus
29. Reduced pinnae to 1.5 cm or more long.
30. Hairs on lower surface of rachis and costae less than 0.1mm long; no setae on
sporangia . . . . . . 26.8. dimidiolobatus
30. Hairs on lower surface of rachis ‘and costae 0.2mm or more long; sporangia
setiferous SMA: . . . 30.8. productus
27. At least 13 pairs of veins ‘truly ‘anastomosing uu 1 vein touching sinus-membrane.
31. Pinnae lobed 3 towards costa, lower ones auricled on acroscopic base 27. S. solutus
31. Pinnae lobed not OVET 5, lower ones not auricled.
32. Pinnae lobed fully 3, widest in distal half ae . . . . 28.8. hernaezii
32. Pinnae in most cases lobed less than 3, not widened distally.
33. Fertile pinnae little more than 2 cm wide; 13-25 pairs of veins truly anastomosing.
34. Reduced pinnae 3-5 pairs, all very small . . . . <a *. 9.2929. ‘Samoseleyi
34. Reduced pinnae commonly 8 pairs or more, upper ones at least 1 x 1 cm, rounded
30. S. productus
33. Fertile pinnae commonly 2.5-3.0cm wide; 3-3 pairs of veins truly anastomosing
31. S. penniger
18. Pinnae to 7cm wide, deeply lobed, lobes narrowly acute . . . . . . 31.S.penniger
1. Glands lacking or rare on upper surface of pinnae.
35. Glands present on lower surface of pinnae between veins.
442 FLORA MALESIANA
(ser. II, vol. 1°
36. Pinnae 2-4 cm long, 3-4 mm wide, subentire, several lower pairs gradually reduced
32. S. warburgii
36. Pinnae, if subentire, much wider; transition to reduced basal pinnae + abrupt.
37. Sporangia bearing several setae, sometimes also a gland.
38. Normal pinnae 3-4 pairs, 11 x 4cm or larger; reduced pinnae 12-15 pairs
38. Normal pinnae more numerous or smaller.
39. Pinnae opposite, 6-7 pairs, 10 x 2.7 cm or larger
39. Not this combination of characters.
40. Sori exindusiate.
41. Pinnae c. 12 x 1.5cm, lobed over 3; reduced pinnae 15 pairs
41. Pinnae c. 6X 1.8 cm, lobed less than 3; reduced pinnae 2 pairs
40. Sori indusiate.
42. Reduced pinnae 1I-2 pairs, normal pinnae 2-4 pairs 3.5 cm long
42. Not this combination of characters.
43. Reduced pinnae at least 6 pairs; transition to large pinnae abrupt.
44. Reduced pinnae 25-30 pairs
44. Reduced pinnae rarely more than 12 pairs.
45. Pinnae lobed conspicuously more than 3 way to costa.
46. Pinnae c. 12 pairs, 12.5 x 2.4cm
46. Pinnae 25-30 pairs, proportionately narrower.
47. Pinnae to 21 x 2.5cm
47. Pinnae to 16 x 1.5 cm
45. Pinnae lobed about } or little more deeply.
48. Reduced pinnae all small, to 3 mm long
48. Reduced pinnae all conspicuous :
43. Reduced pinnae not more than 4 pairs, transition ‘not abrupt.
49. Texture rigid; pinna- lobes dentate at vein-ends
49. Texture thin; pinna-lobes entire
37. Sporangia in most cases bearing glands, sometimes ¢ a seta with the glands.
50. Pinnae subentire, very narrowly cuneate at basiscopic base
oe Pinnae otherwise.
Reduced pinnae 0-2 (—3) pairs, small, irregularly spaced.
a Basal veins free ake ,
52. Basal veins anastomosing.
53. 1 or 2 pairs of pinnae 6-8 cm long and | pair very small
53. Normal pinnae several pairs.
54. Upper surface of pinnae bearing appressed hairs between veins.
= Free pinnae 3-4 pairs below long apical lamina
Free pinnae more numerous; apical lamina not elongates
A Sterile pinnae not more than 3.5 x 1.2 cm.
57. Hairs on lower surfaces all short and mostly appressed.
58. Pinnae 12-15 pairs, 3.5 cm long, symmetrically many-lobed
58. Pinnae 5 pairs, c. 16 cm long, with few irregular lobes
57. Hairs on lower surface of rachis and costae erect, to | mm He
56. Sterile pinnae to8x1.8cm . .
54. Upper surface of pinnae ae appressed hairs between veins.
59. Pinnae 7-10 x 1 cm
59. Pinnae not over 5.5cm long, more . than tre cm wide.
60. Pinnae not dimorphous; apical lamina narrow, 10cm long
60. Pinnae dimorphous; apical lamina broadly triangular, shorter
51. Reduced pinnae at least 4 pairs, regularly spaced;
abrupt.
61. Aerophores distinctly elongate.
62. Basal veins quite free, not meeting at sinus :
62. Basal veins meeting at sinus, or anastomosing below it.
63. Pinnae entire or slightly crenate.
33. S. tandikatensis
. 34.8. carrii
. 35. S. exindusiatus
36. S. lucbanii
. 37.8. major
38. S. polyotis
. 39. S. grandescens
40. S. magnus
41. S. santomasii
42. S. trichochlamys
. 11. S. appendiculatus
43. S. adenostegius
14. S. dichrotrichus
. 44.8. aquatilis
. 45. S. hispidifolius
. 46. S. uniauriculatus
47. S. urdanetensis
48. S. batjanensis
49. S. humilis
50. S. lastreoides
50. S. lastreoides
51. S. cataractorum
. 52.S. menadensis
. 53. S. subcordatus
transition to normal pinnae gradual or
. 54. S. novoguineensis
64. Pinnae c. 51cm, distinctly crenate; transition to 15 pairs of reduced pinnae subabrupt
55. S. hispiduliformis
64. Pinnae larger, almost entire; 12 pairs of lower pinnae gradually reduced
56. S. hastatopinnatus
63. Pinnae distinctly lobed.
1981} THELYPTERIDACEAE (Holttum) 443
65. Appressed hairs present between veins on upper surface of pinnae.
66. 12 pairs of lower pinnae gradually reduced and 20 pairs of smaller ones
55. S. hispiduliformis
66. Subabrupt transition to reduced lower pinnae.
67. Pinnae lobed not or little more than > way to costa.
68. Reduced pinnae conspicuous, upper ones 1-3 cm long.
69. Pinnae 12 x 1.6 to 20 x 2. cm, gradually acuminate; lower surface of costae glabrous or
with sparse short hairs oe 4 . . . 57.8. latebrosus
69. Pinnae to 14x 2.5cm, abruptly short- acuminate: low. er ‘surface of costae densely
covered with appressed hairs . . ..... . . . . . 458.8. porphyricola
68. Reduced pinnae all very small.
70. Lower surface of rachis and costae almost glabrous . . . . 59.8. caulescens
ps Lower surface of rachis and costae bearing hairs to | mm long.
. Largest pinnae commonly 15-20cm long: hairs on rachis pale . . 21.S.hirsutus
a Largest pinnae 7x 1.5cm; hairs onrachis brown . . . . . . 60.S.reconditus
67. Pinnae lobed conspicuously more than > way to costa.
72. Hairs on lower surface of costules antrorsely appressed.
73. Sori supramedial; pinnae to 2.5 cm wide.
74. Basal normal pinnae not narrowed at base . . . . 61.8. cyrtocaulos
74. Several pairs of lower pinnae much narrowed at their ‘bases . . 62.8. baramensis
73. Sori inframedial; pinnae to 1.5 cm wide -9 £Seyeoee Alera 63585 batekitessis
72. Hairs on lower surface of costules not antrorse.
75. Reduced pinnae all conspicuous, lowest 5 mm long . . . . . 64.8. subalpinus
75. Reduced pinnae all small, uppermost 5 mm long.
76. Dark hairs on both sides of rachis . . . ..... . =. - . 65.8. lobatus
76. Pallid hairs on both sides of rachis.
77. Lowest pinnae little narrowed at base; reduced pinnae evident . 66. S. ellipticus
77. Lower pinnae much narrowed at base; all reduced pinnae minute 67. S. foxworthyi
65. No appressed hairs on upper surface between veins.
78. Hairs on lower surface of costules copious, closely appressed.
79. Reduced pinnae all very small, basal normal pinnae narrowed at base.
80. Pinnae lobed less than > tow ardsicosta) ney 4ial (92) Seay. 9). = 6S: Ss iedrapurae
80. Pinnae lobed about ¢ Sige . . . 62.8. baramensis
79. Reduced pinnae conspicuous, many 10-15 1 mm 1 long: basal normal pinnae not narrowed
at base ee. aut oe -. (69) Sc hbatacorum
78. Hairs on lower surface of costules short, ‘not appressed.
81. Sori indusiate; sporangia glandular, rarely with a seta.
82. Lowest normal pinnae narrowed at base Stes cguct BO 2A e Se aneestibass
82. Lowest normal pinnae not narrowed atbase . ... . . . . 57.8. latebrosus
81. Sori exindusiate; neither glands nor setae on sporangia.
83. Pinnae thin. lobed to2mmfromcosta ...... . . . . 71.S8.nudisorus
83. Pinnae subcoriaceous, lobed little more than 3 . «eat & 6722S. paripomains
61. Aerophores not or slightly elongate.
84. Basal veins both passing to sides of sinus-membrane or uniting just below it, not or rarely
forming a distinct excurrent vein.
85. Reduced pinnae all with distinct lamina; costules 2.5mm apart . 73.S.novae-britanniae
85. Reduced pinnae without distinct lamina; costules 4 mmapart .. . . 74.S.convergens
84. Basal veins uniting to form an excurrent vein, at least near base of pinna.
86. Sori supramedial; caudex a long-creeping rhizome > ces. ceeds Ss Seis
86. Sori medial or inframedial; caudex not long-creeping.
87. Pinnae to 2.5 cm long; dark hairs on upper surface of rachis . . . 76.8. sessilipinna
87. Not this combination of characters.
88. Lower surface of rachis glabrous or nearly so; hairs on lower surface of base of costae
none or very short.
89. Reduced pinnae all very small.
90. Largest pinnae c.6cmlong .......-.-.- - - - - - 77.S.lobangensis
90. Largest pinnae at least 10cm long. :
91. Fertile pinnae to 16X2cm,lobed atmosts ... . .- . - 78.S.gymnorachis
91. Fertile pinnae to26x3cm,lobed? . . . . . 79.8. pullenii
89. Reduced pinnae: upper ones 1.5-2 cm long. lower gradually smaller.
92. Pinnae lobed toadepthof2mm ........ . . . . -80.S.gregarius
444 FLORA MALESIANA [ser. Il; voli
92. Pinnae lobed at least 3 wayatorcostal 4 = > i s . . « 57Slatebrosus
88. Lower surface of rachis and costae hairy throughout.
93. Pinnae crenate.
94. Pinnae not more than 4.5 x 1.4 cm; veins 4-5 pairs, I-13 pairs anastomosing.
95. Pinnae 5-6 pairs SAS Ree an SO BRS ee y SI Satephrophyllus
95. Pinnae 18 pairs . . . . . 17.8. kalkmanii
94. Sterile pinnae to 8 x 1.7 cm; 23 pairs of veins anastomosine:! sinus-membrane very short
20. S. lamii
93. Pinnae distinctly lobed.
96. Fronds dimorphous; sterile pinnae to 25 x 3.5 cm . . . . $2.8. pilososquamatus
as Fronds not or little dimorphous, pinnae not over 2.5 cm wide.
. Pinnae to 9.5X0.8-0.9cm . . . . . . .83.S. angustifolius
a Pinnae proportionately wider, to at least 1. 5 cm n wide.
98. Lowest normal pinnae much narrowed at base . . . . . 24,8. heterocarpus
98. Lowest normal pinnae not narrowed at base.
99. Hairs | mm long on lower surface of rachis and costae; indusia with long hairs.
100. Pinnae lobed less than 1/2 way tocosta . . . . . . . . .84. S. efogensis
100. Pinnae lobed more than 1/2 way to costa.
101. Reduced pinnae CASmpairsie Ya 20 TE SS 85. Seidichnotnichoides
101. Reduced pinnae many pairs aa . . . 3. S. polycarpus
99. Hairs inmost cases less than 0.5 mm long on nilower surface of rachis and costae; glands
present on indusia.
102. Lowest veins sometimes both touching sides of sinus-membrane; pinnae lobed
2/3-3/4 Hy as . . . 86.8. nakaikei
102. Lowest veins always aReCtomosine: pinnae lobed & ae
103. Pinnae rigid, lobes of larger ones toothed distally; reduced pinnae not over 8
pairs.
104. Reduced pinnae 4-5 pairs; upper surface of pinnae bearing appressed hairs
between veins aah . . . . 87. S. woitapensis
104. Reduced pinnae 8 pairs; upper surface of pinnae lacking appressed hairs
88. S. ekutiensis
103. Pinnae thin, lobes not toothed; reduced pinnae 12-15 pairs.
105. Hairs on lower surface of costae not appressed.
106. Hairs on lower surface of costae all very short . . . . 57.8. latebrosus
106. Some hairs 0.5 mm or more long on lower surface of costae 66. S. ellipticus
105. Lower surface of costae covered with closely appressed hairs
58. S. porphyricola
35. Glands between veins on lower surface of pinnae lacking.
107. Glands present on lower surface of costules and veins.
108. Pinnae 3-4 pairs, to 1.1 x 0.5 cm, basal pinnae somewhat reduced . . . . 89.S. omatianus
108. Pinnae more numerous or larger, or several pairs reduced basal pinnae.
109. All pinnae with narrowly cuneate basiscopic base . . oe. ee ee). 44,8. aquatilis
109. Pinnae symmetrically broadly cuneate or truncate at base.
110. Few to many pairs of lower pinnae gradually reduced.
111. Pinnae subentire to crenate.
112. Pinnae to 3.5 x 1.0cm; pinna-lobes with strongly reflexed edges.
113. Sporangia setiferous; 12-14 pairs of lower pinnae gradually reduced . -. 90.8. alticola
113. Sporangia not setiferous; 4-5 pairs of lower pinnae gradually reduced . 91.8. rigidus
112. Pinnae larger, edges of lobes not strongly reflexed.
114. Sori exindusiate; lower surface of costae and costules glabrous
56. S. hastatopinnatus
114. Sori indusiate; short hairs abundant on lower surface of rachis and costae
2 92. S. arfakianus
111. Pinnae lobed 3-5 towards costa.
115. 1-4 pairs of basal pinnae gradually reduced . . . . . . . . +. 43.8. adenostegius
115. Many pairs of basal reduced pinnae.
116. 7-12 pairs of lower pinnae gradually reduced with subabrupt change to 12 pairs much
omgilie 5 5 « . . . . . . 55.8. hispiduliformis
116. Similar transition but 2 2 6) pairs ae grail ihasal pinnae eee ee = 95>) Ssarchboldit
110. Transition to reduced basal pinnae (if any) abrupt or nearly so.
1981] THELYPTERIDACEAE (Holttum) 445
117. Reduced pinnae 0-3 pairs.
118. Pinna-lobes strongly concave beneath . .... . woah 28% & °91; Sarigidus
118. Pinnae-lobes not concave beneath.
119. Pinnae widened distally pe ee Tass AOR Aree ee LAS diversilobus
119. Pinnae not widened distally.
120. Caudex short-creeping; pinnae lobed c. § towards costa.
121. Fronds not dimorphous; sori supramedial.
122. Pinnae 3.5 X 1.4cm, narrowed evenly toapex . .. . . . . . 94.S. tibangensis
122. Pinnae larger, abruptly short-acuminate ....... . . . . 9§5.S.norrisii
121. Fronds dimorphous; sorimedial . . . . . . . . . . . 82.8. pilososquamatus
120. Caudex slender, erect; pinnae crenate ...... . .. . . .96.S.semimetralis
117. Reduced pinnae at least 4 pairs.
123. Aerophores elongate.
124. Pinnae crenate.
125. Hairs on lower surface of costae minute .... . . . . . 97.8. semicordatus
125. Hairs on lower surface of costae appressed . . .. . . . . . 98.S. pterosporus
124. Pinnae lobed at least 2/5 towards costa.
126. Hairs on lower surface of costules appressed fy ee ee Sie 9SaSspterosporus
126. Hairs on lower surface of costules not appressed.
127. Reduced pinnae 20-30 pairs; normal pinnae lobed less than > way to costa.
128. Reduced pinnae acuminate Lee Re es Pg BAe OE 99 Sssagittifolius
128. Reduced pinnae with rounded tips ayn: . . . 100.8. foliolosus
127. Reduced pinnae not more than 12 pairs; normal pinnae lobed ) way to costa or more
deeply.
129. Pinnae lobed ; to costa.
130. Basal veins anastomosing; upper reduced pinnae 12mm long . . . 101.S. alpinus
130. Basal veins free; upper reduced pinnae 3 mm long . . . . 54,8. novoguineensis
129. Pinnae lobed about 2 to costa.
131. Pinnae c. 6.5 X 1.3 cm, lowest narrowed at base; indusia glabrous 102. S. polisianus
131. Pinnae c. 11 x 2.cm, lowest not narrowed at base; indusia bearing long hairs
103. S. hoalensis
123. Aerophores not elongate.
132. Sorielongate .. see PREP A ee eee > ey Pen O45 Sslarutensis
132. Sori not or little elongate.
133. Reduced pinnae c. 30 pairs, deflexed, overlapping.
134. Reduced pinnae acuminate; normal pinnae 20cmlong . . . . . 99.S. sagittifolius
134. Reduced pinnae with rounded tips; normal pinnae smaller . . . . 38.8. polyotis
133. Reduced pinnae not over 12 pairs, not overlapping.
135. Sori supramedial.
136. Pinnae lobed 3, sporangia setiferous . . oe. . we) 6. 105.8. loherianus
136. Pinnae lobed less than 3; sporangia glandular ORT se 7S sSsunitus
135. Sori not supramedial.
137. Pinnae lobed more than }.
138. Sori indusiate; sporangia with glands.
139. Largest reduced pinna 7 mm long; hairs on lower surface of costae not appressed
106. S. erectus
139. Largest reduced pinna 2 mm long; hairs on lower surface of costae appressed
98. S. pterosporus
138. Sori exindusiate; sporangia setiferous . . . . . . . . . 107.S.stresemannii
137. Pinnae crenate or lobed less than >.
140. Pinnae not over 1 cm wide; indusia very small or lacking . . . 17.8. kalkmanii
140. Pinnae more than 1 cm wide; indusia all distinct.
141. Some hairs on lower surface of rachis and costae | mm or more long.
142. Pinnae commonly more than 2 cm wide, crenate to a depth of | mm.
143. Apex of frond pinna-like, crenate to base . . . . . . 108. S. simplicifolius
143. Apex of frond deeply lobed at base « OR? SOR 2 eS A109SSsspenceni
142. Pinnae to 1.7 cm wide, distinctly lobed . . . . . . . . 84.8. efogensis
141. Hairs on lower surface of rachis and costae all very Short
144. Largest reduced pinnae 2 mm long » 2, oe 2) a ee 2 ee 98aSopterosporus
144. Largest reduced pinnae at least 7 mm long.
446 FLORA MALESIANA [ser. II, vol. 1°
145. Sporangia not setiferous.
146. Pinnae opposite; caudex 40 cm tall . oe. . . . . . . 110.8. mengenianus
146. Pinnae not opposite; caudex suberect SVs Se oe S05Ssoreganins
145. Sporangia setiferous . . soon A eee aye SMS Sconfertus
107. Glands lacking on lower surface of costules and \ veins.
147. Reduced pinnae c. 30 pairs.
148. Reduced pinnae acuminate. STAT a ce hae ee eee ease bene ep 299RSSsacittifolius
148. Reduced pinnae with rounded tips Via) bose toe Sk) .e 8 Veer QS8aSspolyotis
147. Reduced pinnae much fewer.
149. Sporangia bearing glands (sometimes a seta also).
150. Hairs on lower surface of costules appressed.
151. Aerophores elongate.
152. Several pairs of lower pinnae gradually reduced . o. . . . . 112.8. multiauriculatus
152. Transition to reduced pinnae + abrupt.
153. Upper reduced pinnae 12mm, lowest6mmlong ... . . . . . 113.S.stipellatus
153. All reduced pinnae very short.
154: *Pinnae lobed léssithania. = = < Aino) gb istesi ).. eee Soucy AGREES! pterosporus
154. Pinnae lobed more than 3.
155. Indusia large, firm . .watsis- 2. eee 3) eee SS lonitotine
155. Indusia small, thin.
156. Basal veins anastomosing, second acroscopic vein to sinus-membrane
115. S. hendersonii
156. Basal veins often connivent at sinus-membrane, next veins both to margin
116. S. posthumii
151. Aerophores not elongate.
157. Basal acroscopic vein ending beside sinus-membrane, basiscopic vein at margin above base
of sinus.
158. Pinnae 15-20 pairs; reduced pinnae 4-7 pairs . oo. . . . . . . 117.8. inconspicuus
158. Pinnae to 30 pairs; reduced pinnae 2-3 pairs . . . . . .118.S. subulifolius
157. Basal veins both touching sinus-membrane or uniting below it.
159. Gradual transition to reduced pinnae at base of frond.
1605sindustaspresent’ 0 A seen Sea le eee es (eo 2a Saantakianns
160. Indusia absent eS: . oo. . . . . . 112.8. multiauriculatus
159. Abrupt transition to very ‘small reduced pinnae . oe... . . . . 98. S. pterosporus
150. Hairs on lower surface of costules not appressed.
161. Basal veins not connivent below sinus.
162. Pinnae deeply lobed.
163. Pinnae to 3.6 x 1.0cm; veins 3-4 pairs 8S 4 een ee Sea. Deo eS eSSmiyohersii
163. Pinnae to 10 x 1.5 cm; veins 9-10 sa . oe ee ee). 54.8. novoguineensis
162. Pinnae almost entire . . . . . . 120.8. mutabilis
161. Basal veins connivent below sinus or uniting to form an excurrent vein.
164. Pinnae to 4.2 x 1.1 cm; sori elongate with small indusia . . . . . . 4121.S.uaniensis
164. Pinnae in most cases much larger; sori otherwise.
165. Ten or more pairs of basal pinnae gradually reduced.
166. Pinna-margins sinuous or slightly crenate . . . . . . . 56.8. hastatopinnatus
166. Pinnae deeply lobed A ee . oe. ee ws). . 122.8. echinosporus
165. Abrupt transition to reduced pinnae if present.
167. Reduced pinnae 0-4 pairs on mature plants.
168. Pinnae almost entire, 3.5cm or more wide ... . . . . . 123.8. peltochlamys
168. Pinnae distinctly lobed, narrower.
169. Pinnae lobed more than uitiecte tlaniee: “Abid diad. oxv', Wer seegiaaass irayensis
169. Pinnae lobed less than}. . spike alot), Se RES SS jnornisit
167. Reduced pinnae at least 6 pairs on mature plants.
170. Sori supramedial.
171. Caudex a long-creeping rhizome ecb aivnd Je. 1c. cee A SaSamitus
ijt Caudexeshontenectar), pace tees tes) tees Goes Sel eA SAI25SSalithophyllus
170. Sori medial or inframedial.
172. Pinnae crenate : PURtye AG abe) he ay Wenst Be sues LOSS sspenceri
172. Pinnae lobed at begi2
173. Pinnae lobed more than } Rea tye Ges (eee $590ae0.2> 391262 Sshamiferus
1981] THELYPTERIDACEAE (Holttum) 447
173. Pinnae lobed less than> . . . jucret. 10. tere See BSOFSZerecarius
149. Sporangia lacking glands, in most cases bearing setae.
174. Aerophores elongate.
175. Pinnae entire or slightly crenate; veins 18-20 pairs . . . . . . . 127. S. plurivenosus
175. Pinnae distinctly lobed; veins rarely more than 12 pairs.
176. Pinnae lobed less than >
177. Hairs on lower surface of costae and costules appressed.
178. Lower 12-15 pairs of pinnae gradually reduced, lowest 1 cm long
112. S. multiauriculatus
178. Transition to small pinnae at base of frond abrupt . . . . . . 128.S.squamatellus
177. Hairs on lower surface of costae and costules not appressed.
179. Reduced pinnae more than 12 pairs; sporangia not setiferous . . 55.S. hispiduliformis
179. Reduced pinnae to 5 pairs; sporangia setiferous Ay ovata Wie) ee 29S. canescens
176. Pinnae lobed more than 3.
180. Sporangia always with several setae.
181. Basal 2-3 pairs of pinnae somewhat narrowed at their bases.
182. Pinnae to 11 x 2.1 cm, lobed <3; hairs on lower surface of costules antrorse
130. S. suboppositus
182. Pinnae to 17 x 2.8cm, lobed more than 4; hairs on lower surface of costules erect
131. S. makassaricus
181. All pinnae in lower half of frond narrowed at their bases, lowest much narrowed
132. S. sarasinorum
180. Sporangia rarely with a seta.
183. Pinnae lobed 4; or more deeply; reduced pinnae all very small.
184. Lower surface of rachis glabrous vee . . . . 62.8. baramensis
184. Lower surface of rachis densely covered with erect brown hairs . 133. S. muluensis
183. Pinnae lobed at most 3; reduced pinnae 1-1.5 cm long . . . . . 126.8. hamiferus
174. Aerophores not elongate.
185. Sporangia not setiferous.
186. Pinnae not or little more than 5 cm long.
187. Pinnae crenate; sori inframedial.
188. Pinnae 10-20 pairs; 2-3 pairs of lower ones gradually reduced.
189. Caudex slender, erect; pinnae slightly crenate distally . . . . 134.S. telefominicus
189. Caudex very short; pinnae crenate throughout . . . . . . . 135.S.roemerianus
188. Pinnae to 30 pairs; abriet change to reduced ones at base . . . 136.S. hellwigensis
187. Pinnae lobed more than 2; sori supramedial at. hes «ate els 7iSsbenoitianus
186. Pinnae more than 10cm long.
190. No reduced pinnae; normal pinnae distinctly stalked . . . . . 138.8. neotoppingii
190. At least 5 pairs lower pinnae gradually or abruptly reduced; all pinnae sessile.
191. Sori elongate and indusiate.
192. Normal pinnae not or little more than 2 cm wide; upper reduced pinnae 4 mm long
139. S. oosorus
192. Normal pinnae commonly 2.5—3.5 cm wide; reduced pinnae much larger
104. S. larutensis
191. Sori not elongate, or if so exindusiate.
193. Abrupt transition to 6 pairs of very small basal pinnae . .. . . . . 140.S.rudis
193. Basal pinnae not very small.
194. Rachis quite glabrous on lower surface; pinnae almost entire . . 141.S.dimorphus
194. Rachis hairy on lower surface; pinnae crenate or lobed throughout 142. S. potamios
185. Sporangia setiferous.
195. Caudex terrestrial and long-creeping, or scandent.
196. Caudex terrestrial; indusia distinct : = laut, tee ee 1435S sinvisus
196. Caudex scandent; indusia very small or lacking.
197. Several pairs of lower pinnae gradually smaller and auricled on acroscopic base
144. S. austerus
197. Transition to basal reduced pinnae abrupt . . . . . . . . +. . +. 145.S. mundus
195. Caudex various, not long-creeping nor scandent.
198. Lower 1-S pairs of pinnae gradually reduced.
199. Pinnae to2cmlong;noindusia . . 50a. itd an = 6S obtusifohus
199. Pinnae to at least 4cm long; indusia present J 2” 5 eactldewiat Leoeee 435 Syadenostegius
448 FLORA MALESIANA [ser. II, vol. 1°
198. At least 3 pairs of reduced pinnae distinct from normal ones.
200. Pinnae lobed 3 way to costa or more deeply.
201. Reduced pinmae 20 pairs, all very small
201. Reduced pinnae to 12 pairs, all eee
202. Caudex short, erect; pinnae lobed 2
202. Caudex tall, slender, if free-standing; pinnae lobed 3 =
200. Pinnae lobed less than 3 way to costa.
. 147. S. alcasidii
148. S. pilosissimus
. 145. S. mundus
203. Pinnae to 3.5 cm long, slightly crenate
149. S. erwinii
203. Pinnae in most cases larger, deeply crenate or lobed.
204. Stipe 40-50 cm long; reduced pinnae 3 pairs, small
82. S. pilososquamatus
204. Stipe much shorter; reduced pinnae more numerous, conspicuous.
205. Pinnae crenate; no long hairs on lower surface of rachis and costae.
206. Caudex erect; largest fertile pinnae 17x 2.5 cm
206. Caudex short-creeping; largest pinnae 8 x 2cm
. 150. S. atasripii
111. S. confertus
205. Pinnae distinctly lobed; long hairs usually present on lower surface of rachis and
costae.
207. Stipe-scales 1 mm long; pinnae to 4 Icm
207. Stipe-scales 7 mm long; pinnae to 12 x 2cm
1. Sphaerostephanos debilis (METT.) HOLTTUM,
comb. nov.— Phegopteris debilis METT. Ann.
Mus. Bot. Lugd.-Bat. 1 (1864) 123, t. 6 f. 1;
v.A.v.R. Handb. (1908) 501.— Dryopteris debilis
(METT.) C. CHR. Ind. Fil. (1905) 260.—
Cyclosorus debilis (METT.) COPEL. Gen. Fil.
(1947) 142, non CHING 1941. — Thelypteris debilis
(METT.) REED, Phytologia 17 (1968) 270. — Type:
ZIPPELIUS, Amboina (L).
Caudex short, erect; stipe 34cm long. Lamina
30-40 cm long, gradually attenuate both to base
and apex; basal pinnae 5mm long; texture thin.
Largest pinnae 3.0cm long, 0.7cm wide above
base; base truncate, 1.0cm wide, auricled on
acroscopic side, slightly dilated on basiscopic,
distal part gradually narrowed to a rounded apex,
edges slightly crenate; costules 2.5mm _ apart;
veins 2 pairs, lower pair anastomosing; sparse
hairs | mm long on lower surface of rachis, sparse
minute hairs on costae and costules, small spheri-
cal glands throughout; minute suberect hairs and
glands between veins on upper surface. Sori in-
framedial, small, exindusiate; sporangia bearing
1-2 short setae and sometimes a gland distally and
a sessile gland on the stalk.
Distr. Malesia: Moluccas (Amboina, without
collector’s name at B, K, L, also one cult. Hort.
Bog. at L); W. New Guinea.
Ecol. In New Guinea on rocks by river at
100 m.
Note. The New Guinea specimens (KANE-
HIRA & HATUSIMA 12837) have lamina 30 cm
long with 50 pairs of pinnae, largest pinnae 2.0 cm
long, 0.4-0.5cm wide above base, basal pinnae
3 mm long; a few long hairs are present on lower
surface of costae.
2. Sphaerostephanos sudesticus
nov.
Pinnae_ redactae
54cm longus;
DCM Dast
HOLTTUM, sp.
verisimiliter nullae; lamina
pinnae 15-jugatae, maximae 17 Xx
late cuneatae, 3/5 costam versus
. 151. S. albosetosus
152. S. wauensis
lobatae, utrinque pilis minutis erectis glandulisque
praeditae; venae 9-jugatae, basales solum anas-
tomosantes; sori mediales, inferiores praesertim
elongati, indusiis tenuibus breviter hirsutis et
glandulosis tecti; sporangia setifera.— Type:
BRASS 27772, Sudest Island, Louisiade
Archipelago, terrestrial in openings in forest, alt.
2m (L).
Caudex not known; stipe 47cm long, minutely
hairy. Reduced pinnae not seen; if present, minute
and few. Lamina 54cm long; pinnae 15 pairs,
alternate; 3-4 pairs upper pinnae adnate to rachis
with asymmetric bases; lowest pinnae 2.5cm
wide. Rachis minutely hairy on lower surface,
hairs on upper surface pale, less than 0.5 mm long.
Middle pinnae to 17X2.2cm, base _ broadly
cuneate, aerophores not swollen, apex with cauda
2cm long, edges lobed to 3.5-4mm from costa
(less than 2/3), lobes slightly falcate; costules
5mm apart, at 60°; veins to 9 pairs, | pair anas-
tomosing, next acroscopic vein to sinus-mem-
brane; hairs on lower surface of costae 0.2mm
long, slightly antrorse, a little longer distally,
similar shorter hairs more sparse on costules,
veins and between’ veins, glands present
throughout; hairs on upper surface of costae
0.2 mm long, scattered hairs 0.5 mm long on cos-
tules and veins, very short erect hairs and glands
between veins. Sori medial, almost all elongate,
lower ones longest; indusia very thin, rather large,
bearing sparse short hairs and several glands;
sporangia bearing I-2 short setae.
Distr. Malesia: Papua New Guinea (Sudest I.),
only known from the type.
Ecol. Terrestrial in openings in forest at 2 m.
3. Sphaerostephanos polycarpus (BL.) COPEL. Un.
Cal. Publ. Bot. 16 (1929) 60; Fern FI. Philip. (1960)
379. — Aspidium polycarpon BL. Enum. PI. Jav.
(1828) 156.—Didymochlaena polycarpa (BL.)
BAK. Syn. Fil. (1867) 248; RacisB. Fl. Btzg 1
(1898) 197.—Nephrodium polycarpum (BL.)
1981]
KEYS. Pol. Cyath. Herb. Bung. (1873) vii; BEDD.
Handb. Suppl. (1892) 74.— Mesochlaena_ poly-
carpa (BL.) BEDD. Ferns Br. Ind. Suppl. (1876)
13; Handb. (1883) 199; v.A.v.R. Handb. (1908)
232.— Dryopteris polycarpa (BL.) CHRIST,
Philip. J. Sci. 2 (1907) Bot. 202.— Cyclosorus
polycarpus (BL.) HOLTTUM, Rev. FI. Malaya 2
(1955) 283.—Thelypteris polycarpa (BL.)
K. TWAT. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964)
32.— Type: BLUME, Noesa Kambangan, Java
(L).
Aspidium heterodon BL. Enum. PI. Jav. (1828)
157. — Type: from Celebes, no collector cited (L).
S. asplenioides J. SM. in Hook. Gen. Fil. (1839)
9245 KUNZE, Farnkr. 1 (1840) t. 11, 12.—
Mesochlaena asplenioides J. SM. in Hook. J. Bot.
3 (1840) 18.— Stegnogramma asplenioides (J.
SM.) FEE, Gen. Fil. (1852) 204.— Type: origin
unrecorded (BM in Herb. J. Sm.).
Mesochlaena javanica R. BR. ex METT. Fil.
Hort. Lips. (1856) 96, t. 18 f. 13.— Aspidium
jJavanicum MeEtTT. Farngatt. IV (1858) 103.—
Nephrodium javanicum (METT.) Hook. Fil.
Exot. (1859) t. 62. — Type: “Java” (not seen).
Lastrea microchlamys DE VRIESE, Tijdschr.
Wisk- Nat. Wet. Amst. | (1848) 155. — Type: Cult.
Hort. Bot. Lugd.-Bat. (L).
Nephrodium microchlamys BAK. J. Linn. Soc.
Bot. 15 (1876) 107.—Type: MOSELEY, Chal-
lenger Exp. Kei Isl. (K).
Aspidium perakense BEDD. J. Bot. 26 (1888)
4.— Nephrodium perakense (BEDD.) BAK. Ann.
Bot. 5 (1891) 319; BEDD. Handb. Suppl. (1892)
80. — Thelypteris perakensis (BEDD.) REED, Phy-
tologia 17 (1968) 303.—Type: J. DAy, Perak,
Birch’s Hill, on exposed rocks (K; E).
Mesochlaena toppingii COPEL. Philip. J. Sci. 12
(1917) Bot. 57; C. CHR. Gard. Bull. Str. Settl. 7
(1934) 254. — S. toppingii (COPEL.) C. CHR. Ind.
Fil. Suppl. III (1934) 172. — Type: TOPPING 1902,
Mt Kinabalu (Herb. Am. Fern Soc.).
S. unijugus COPEL. Philip. J. Sci. 60 (1936) 109,
t. 15. — Type: BRASS 2692, Solomon Is., San Cris-
tobal (UC; BRI, MICH). — Fig. 12a—.
Caudex erect; stipe c. 10cm long, bearing
many narrow scales 10mm long. Reduced pinnae
20 pairs or more, 2.5-3.5cm apart, lowest 1 cm
long, uppermost 2.5-3cm, spreading, triangular
with broad base dilated almost equally both sides
and acute apex, edges of larger ones incised.
Lamina above reduced pinnae 100-150 cm long;
pinnae many pairs, rather close; basal pinnae not
narrowed at base, aerophores slightly swollen.
Largest pinnae commonly 25x 1.8cm, largest
seen 35*2.5cm; base rather asymmetric, basal
acroscopic lobe often elongate; edges lobed more
than > way to costa; lobes slightly oblique, slightly
falcate; costules commonly to 4mm apart; veins
12-15 pairs, basal pair anastomosing with excur-
rent vein to sinus, I-2 pairs passing to sides of
sinus-membrane (in smaller fronds sometimes
THELYPTERIDACEAE (Holttum)
449
only acroscopic vein); lower surfaces bearing
glands throughout, acicular hairs on rachis, costae
and costules normally short, spreading, in some
specimens to | mm long; acicular hairs on upper
surface of costae to 1mm long, scattered similar
hairs on costules and veins, between veins a vari-
able number of glands and very short erect aci-
cular hairs. Sori medial, elongate, commonly
Imm long; indusia with many yellow glands;
sporangia also bearing glands; spores with a
moderate number of wings of varying size.
Distr. Peninsular Thailand; throughout Malesia
except Philippines north of 10° and Lesser Sunda
Islands east of Lombok; Solomon Islands, Samoa.
Ecol. In open not too dry places, in low country
and to 1500 m.
Notes. Quite small plants may be fertile; in
these sometimes the second veins may both reach
the margin above base of sinus. Specimens with
long hairs on the lower surface do not differ from
the normal condition in other ways, and length of
hairs varies; there is no evident correlation of long
hairs with drier habitats, but this is possible.
Aspidium perakense BEDD. was based on young
sterile plants growing on a rock; they are very
hairy. DAy collected with them fragments of a
fertile frond of an abnormal Christeila; this is the
source of BEDDOME’S description of sori.
Mesochlaena toppingii COPEL. was _ based
on a small hairy specimen. S. unijugus was also
based on a small specimen in which at most the
second acroscopic vein passes to the sinus-mem-
brane; the upper surface was almost or quite
glandless but similar specimens from New Britain
have many glands. Another specimen from the
Solomon Islands, otherwise similar, has a fair num-
ber of glands on the upper surface.
4. Sphaerostephanos eminens (BAK.) HOLTTUM,
comb. nov. — Nephrodium eminens BAK. J. Bot.
18 (1880) 213.— Dryopteris eminens (BAK.) C.
Cur. Ind. Fil. (1905) 263; v.A.v.R. Handb. (1908)
214.— Type: BECCARI 455, Sumatra, G. Sing-
galang 1700 m (FI; K).
Dryopteris pilososquamata_ var. obtusata
v.A.v.R. Bull. Jard. Bot. Btzg II, 16 (1914) 12;
Handb. Suppl. (1917) 182. — Type: MATTHEW 645,
G. Singgalang (BO).
Mesochlaena talamauensis v.A.v.R. Bull. Jard.
Bot. Btzg II, 28 (1918) 34.—S. talamauensis
(v.A.v.R.) C. CHR. Ind. Fil. Suppl. II (1934) 172. —
Type: BUNNEMEIJER 873, Sumatra, Talamau,
1750 m (BO).
Caudex erect or suberect. Stipe c. 50cm long,
glabrous; base of stipe to first large pinna 110 cm,
this part bearing 7 pairs widely-spaced reduced
pinnae, upper ones broadly triangular, 2.2 cm long
and wide, lowest hardly |cm long. Lamina to
90 cm long, pinnae 16 pairs, lowest almost sessile
and slightly narrowed at base, middle ones with
stalks 3-12 mm long; apex of frond almost pinna-
450
FLORA MALESIANA
[ser. II, vol. 1°
like; aerophores not swollen. Largest pinnae 30 x
3cm, base cuneate, apex long-acuminate, lobed
about 2 way to costa, lobes distinctly falcate,
blunt, entire; costules 7 mm apart at 60°; veins to
12 pairs, concolorous and prominent on lower
surface but not above, basal 1-15 pairs anas-
tomosing, 2 pairs to sides of sinus-membrane; pale
curved hairs 0.2-0.5 mm long on lower surface of
rachis and costae, shorter and somewhat antrorse
on costules, glands (no hairs) on veins and surface
between veins; hairs 0.5 mm long on upper surface
of rachis and costae, much shorter on costules,
glands present between veins. Sori medial, lower
ones somewhat divergent, all except distal ones +
elongate; indusia and sporangia bearing glands.
Distr. Malesia: Middle to northern Sumatra,
several collections.
Notes. The length of pinna-stalks varies in
different specimens; those of the type of M.
talamauensis only 3mm, on BECCARIs type
12mm, on a specimen of MATTHEW’s from the
same locality 5mm. YATES 699 from Brastagi is
rather intermediate between this species and S.
penniger, with sessile pinnae and _ setiferous
sporangia.
5. Sphaerostephanos williamsii (COPEL.) HOLT-
TUM, Kalikasan 4 (1975) 53.— Dryopteris willi-
amsii COPEL. Brittonia 1 (1931) 67, pl. 1.—
Thelypteris williamsii CHING, Bull. Fan Mem.
Inst. Biol. Bot. 10 (1941) 255. — Lastrea williamsii
COPEL. Gen. Fil. (1947) 140; Fern FI. Philip.
(1960) 322.— Type: R. S. WILLIAMS 2216, Min-
danao, San Ramon (UC; B, G, NY, US).
Caudex short, erect. Stipe 10cm long, very-
short-hairy, basal scales 2 mm long, ovate; base of
stipe to first normal pinna 20-30mm. Reduced
pinnae 3 pairs, not opposite, irregularly spaced,
lowest 4mm long; transition to normal pinnae
abrupt or with one pair intermediate. Lamina of
type 30cm long (of NY isotype 40cm); pinnae
12-14 pairs, 1-2 lower ones narrowed at base;
aerophores not enlarged. Largest pinnae of type
6.5 xX 1.lcm (of NY isotype 11 x 1.8cm) lobed to
less than 1 mm from costa, apex acuminate; lobes
oblique, slightly falcate; costules 2.5-3.5 mm
apart, at 60°; veins 8-10 pairs, both basal veins
ending above base of sinus; acicular hairs on
lower surface of rachis and costae short, spread-
ing; whole lower surface of pinna covered with
short slender erect hairs and glands; upper surface
minutely hairy and glandular throughout. Sori
medial, exindusiate; sporangia bearing many small
yellow glands, less often 2-3 short setae; spores
with thin translucent wing and cross wings.
Distr. Malesia: Philippines (Mindanao, at
150 m), only known from the type.
6. Sphaerostephanos stenodontus (COPEL.)
HOLTTUM, Kalikasan 4 (1975) 56. — Cyclosorus
stenodontus COPEL. Philip. J. Sci. 81 (1952) 28, t.
20; Fern Fl. Philip. (1960) 344. — Thelypteris
stenodonta REED, Phytologia 17 (1968) 315.—
Type: RAMOS & EDANO BS 30849, Panay (US).
Caudex not known; base of stipe lacking.
Reduced pinnae at least 5 pairs, lowest 3-4mm
long; transition to normal pinnae subabrupt.
Lamina 50cm long; pinnae 30 pairs, lower ones
not narrowed at base. Largest pinnae 10cm long,
15cm wide at dilated base, short-acuminate,
lobed 3/5 towards costa, lobes slightly oblique, not
falcate; costules 2.5—3 mm apart, at more than 60°;
veins 8-9 pairs, | pair anastomosing, next acros-
copic vein to short sinus-membrane; hairs more
than | mm long on lower surface of rachis, fewer
with shorter ones on costae and costules, glands
all over surface; scattered long hairs on upper
surface of costules and veins, short slender ap-
pressed hairs and a few glands between veins.
Sori medial, lacking indusia, basal ones not
divergent; sporangia bearing glands.
Distr. Malesia: Philippines (Panay), only
known from the type.
7. Sphaerostephanos flavoviridis HOLTTUM, sp.
nov.
Pinnae redactae 2-3-jugatae, superiores 1-2.cm
longae; lamina 120cm longa; pinnae inferiores
basi angustatae; pinnae maximae 21 X2.5 cm,
profunde lobatae; venae 12-—14-jugatae, infimae
anastomosantes; costae subtus pilis minutis
praeditae; pagina inter venas utrinque copiose
glandulosa; sori mediales, exindusiati; stipites
sporangiorum pilis glanduliferis praediti. — Type:
JERMY 3694, N.E. New Guinea, Butemu, Morobe
Distr. on steep bank in open place (BM).
Caudex not known. Stipe 30cm long, glabrous.
Reduced pinnae variable, 2-3 pairs on 13cm
length of rachis, lowest very small, uppermost
1-2 cm long. Lamina 120 cm long, somewhat yel-
low-green when dry; basal pinnae much narrowed
at base, next two pairs less so; aerophores not
elongate. Middle pinnae to 21x2.5cm, almost
sessile, base broadly cuneate to subtruncate, apex
with cauda 3cm long, lobed to 2.5-3mm from
costa, lobes hardly falcate, oblong; costules 5 mm
apart, at more than 60°; veins 12-14 pairs, basal
pair anastomosing with short excurrent vein to
sinus, next pair to sides of short sinus-membrane;
hairs on lower surface of rachis and costae
minute, on costules 0.2 mm long, longer on sinus-
membrane and edge, glands throughout lower
surface; long antrorse hairs on upper surface of
costae, short on costules and veins, glands and
minute hairs between veins. Sori medial, basal
ones divergent; no indusia seen; sporangia lack-
ing glands or setae on body, hairs on stalks
with glandular end-cell; spores with many small
wings.
Distr. Malesia: Papua New Guinea. Known
only from type and another specimen from same
locality.
1981]
THELYPTERIDACEAE (Holttum)
451
8. Sphaerostephanos trimetralis HOLTTUM, sp.
nov.
Caudex erectus, brevis; stipes 10cm longus, e
basi stipitis usque pinna normalis infima 150 cm;
pinnae redactae 30-jugatae, late triangulares;
lamina 150cm longa; pinnae maximae 14x
1.3. cm, crenatae; venae 6—7-jugatae; rachis cos-
taeque subtus pilis 0.2mm _ longis adpressis
praeditae; pagina inter venas utrinque glandulosa;
sori mediales, indusia pilis brevibus et glandulis
praedita; sporangia glandulifera. — Type:
HOOGLAND 3391, Papua, Lake Koena (CANB;
Is, IL)
Caudex short, erect. Stipe 10cm long, minutely
hairy, basal scales 7mm long, narrow; base of
stipe to lowest normal pinna 150cm or more.
Reduced pinnae c. 30 pairs, sub-opposite, 4-6 cm
apart, broadly deltoid with symmetric base, upper
ones 2X1.8cm, crenate, apex acute. Lamina
150cm long; pinnae subcoriaceous, drying light
red-brown; aerophores not enlarged. Largest
pinnae 14x 1.3 cm, base truncate, apex acuminate,
edges crenate to depth of 1mm or little more;
costules 3mm apart, at more than 60°; veins 6-7
pairs, 23 pairs anastomosing to form a Zig-zag
excurrent vein, | pair to short sinus-membrane;
lower surface of rachis, costae and costules
covered with pale antrorse hairs 0.2mm long, a
few short hairs and many glands between veins;
hairs on upper surface of costae 0.5mm long,
between veins short erect hairs and glands. Sori
medial; indusia thin, bearing short hairs and
glands; sporangia with glands; spores minutely
papillose.
Distr. Malesia: Papua New Guinea.
known from the type.
Ecol. Near shore of lake, in marshy vegetation
of tall grasses and sedges, alt. 10 m.
Only
9. Sphaerostephanos alatellus (CHRIST) HOLT-
TUM, comb. nov.—Nephrodium alatellum
CHRIST in K. Schum. & Laut. FI. Schutzgeb.
(1900) 112.— Aspidium alatellum (CHRIST)
CHRIST, Bull. Herb. Boiss. II, 1 (1901) 454.—
Dryopteris alatella (CHRIST) C. CHR. Ind. Fil.
(1905) 251; v.A.v.R. Handb. (1908) 210. — Cyclo-
sorus alatellus (CHRIST) COPEL. Gen. Fil. (1947)
142; Philip. J. Sci. 78 (1951) 445, excl. syn. Dry-
opteris stenobasis C. CHR., D. logavensis
ROSENST. & D. pseudostenobasis COPEL. —
Type: LAUTERBACH 2354, N.E. New Guinea,
600 m(B).
Cyclosorus heterocarpus sensu HOLTTUM &
Roy, Blumea 13 (1965) 134. — Fig. 13a—c.
Caudex short, erect. Stipe 15cm long, basal
scales thin, to 7 X 2 mm; base of stipe to first large
pinna 40-60 cm. Reduced pinnae 4-6 pairs, very
small. Lamina dimorphous, to 60 cm long; pinnae
20 pairs or more, texture thin, aerophores not
elongate, basal pinnae somewhat narrowed at
base. Largest sterile pinnae 20 x 2.6cm, fertile to
14x 1.4cm; base unequally broadly cuneate, apex
short-acuminate, edges lobed 2/3-3/4 to costa,
lobes slightly falcate with rounded tips; costules
of sterile fronds 4.5-Smm apart, of fertile 3.5-
4.5 mm; veins 8-9 pairs (sterile) 6-8 pairs (fertile),
basal pair spreading at a wide angle to costule and
anastomosing with a short excurrent vein or
sometimes meeting just below sinus; next veins
both to edge; lower surface quite hairless or with
some very short hairs on costa only, glands
abundant; hairs on upper surface of costae short,
very short and sparse on costules, between veins
many glands. Sori medial; indusia small, glan-
dular; sporangia bearing glands.
Distr. Malesia: Widely in Eastern
Guinea, in forest at 400-1300 m.
Notes. CHRIST described only a fertile frond
and noticed neither glands nor reduced pinnae.
COPELAND confused this species with Am-
phineuron attenuatum (O. KTZE) HOLTTUM in
the Philippines (Fern FI. Philip. 341) and with
Dryopteris logavensis ROSENST. (Amphineuron
pseudostenobasis (COPEL.) HOLTTUM) in New
Guinea. HOLTTUM and Roy misidentified a cul-
tivated plant as Cyclosorus heterocarpus and
reported a chromosome count 2n = 72.
New
10. Sphaerostephanos HOLTTUM,
sp. nov.
Caudex erectus, 30-40cm altus; stipes usque
30cm longus; pinnae redactae c. 6-8-jugatae,
superiores 1.5-2.5 cm longae, integrae, aerophorae
vix 1mm longae; lamina 150 cm longa; pinnae
maximae 25 X2.cm vel majores, usque 1/3 costam
versus lobatae; venae 9-jugatae, 1-—13-jugatae
anastomosantes; pagina utrinque inter venas
glandulosa; sori inframediales, inferiores leviter
divergentes et leviter elongati; indusia glabra,
glandulosa; stipites sporangiorum pilis glanduli-
feris praediti.— Type: BRASS 23557, Papua,
Milne Bay District, Mt Dayman (BM; A, L, LAE).
Caudex erect, 30-40cm tall, 4-5cm diam.
(BRASS). Stipe to 30cm long, base covered with
broad thin scales 10-15 mm long; base of stipe to
first large pinna 70cm or more. Reduced pinnae
6-8 pairs, uppermost 1.5—2.5 cm long, broadly tri-
angular, lowest less than 1mm long. Lamina
150cm long; lower 3-4 pairs normal pinnae
gradually narrowed and more shallowly lobed
towards base; aerophores hardly | mm long. Lar-
gest pinnae of type 25x 2.0cm (of BRASS 23226
36x2.4cm), base subtruncate, apex caudate-
acuminate (cauda 34cm), lobed 1/3-2/5 to costa,
lobes slightly falcate; costules 4.5mm apart (to
6 mm in BRASS 23226) at more than 60°; veins to 9
pairs, 1-1; pairs anastomosing, 13-2 pairs to sinus-
membrane; all lower surfaces glandular
throughout, no acicular hairs; hairs on upper sur-
face of costa 0.5 mm long, very short on costules,
glands on all parts of surface. Sori inframedial,
lower ones slightly divergent and slightly elongate;
daymanianus
452
indusia thin, bearing glands only; sporangia lack-
ing glands on body, hairs on stalks with terminal
gland which is red in young sori.
Distr. Malesia: Eastern New Guinea (Milne
Bay and Morobe Districts), Bougainville.
Ecol. In forest at 700-1500 m.
Notes. PULLEN 7964 (Milne Bay Distr.) has
pinnae to 2.6cm wide, lobed a little more than ,
with veins to 13 pairs.
WOMERSLEY & MILLAR 8373 (Morobe Distr.)
is reported to have had a caudex 150 cm tall; its
sori are young and not elongate.
CRAVEN & SCHODDE 277 from Bougainville
had a caudex 60 cm tall, base of stipe to first large
pinna 95 cm, fronds in all 3.25 m long; my record
is that it has glands on body of sporangia, which
needs checking.
11. Sphaerostephanos appendiculatus (BL.)
HOLTTUM, Kalikasan 4 (1975) 62.—Gym-
nogramme appendiculata BL. Enum. Pl. Jav
(1828) addendum to p. 112; Fl. Jav. Fil. (1828) 92,
t. 39. — Goniopteris appendiculata (BL.) PRESL,
Tent. Pterid. (1836) t. 7, f. 11.— Aspidium
pilosiusculum METT. Ann. Mus. Bot. Lugd.-Bat.
1 (1864) 233, nom. nov.— Nephrodium ap-
pendiculatum (BL.) RAciIB. FI. Btzg 1 (1898)
186.— Dryopteris appendiculata (BL.) C. CHR.
Ind. Fil. (1905) 252: v.A.v.R. Handb. (1908) 213;
BACKER & POSTH. Varenfl. Java (1939) 52, f.
9.— Thelypteris appendiculata (BL.) REED, Phy-
tologia 17 (1968) 260.— Type: BLUME, Java,
Boerangrang (L).
Dryopteris perakensis var. sumatrensis v.A.v.R.
Bull. Jard. Bot. Btzg II, 11 (1913) 13.— Type:
MATTHEW 516, Sumatra, Padang Pandjang (BO;
K).
Dryopteris brunnescens C. CHR. Bot. Jahrb. 66
(1933) 44. — Thelypteris brunnescens REED, Phy-
tologia 17 (1968) 265.— Type: KJELLBERG 3534,
S. W. Celebes, Rante Lemo (S-PA; BO).
Caudex short-creeping. Stipe short, densely
short-hairy; base of stipe to first large pinna 15-
40 cm. Reduced pinnae 6-12(-20) pairs, somewhat
deflexed, lowest 3-S mm long, upper ones | cm
long, broadly triangular with asymmetric base.
Lamina 20-45(-70) cm long, reddish when young;
apical lamina deeply lobed with gradual transition
to upper pinnae, several pairs of which are adnate;
free pinnae 10-20 pairs, oblique with upturned
tips. Largest pinnae commonly to 8.5 x 1.5 cm (to
17 x 2.2 cm); base truncate and distinctly auricled
on acroscopic side (bases of upper ones asym-
metric), apex short-acuminate, lobed + way to
costa (largest ones more than 3); lobes oblique
with falcate obtusely angled tips; costules 3.5 mm
apart in small frond, to 5 mm in large ones; veins
6-7(-9) pairs, one pair anastomosing, next pair to
sinus-membrane; lower surface of costae and
rachis bearing short and long spreading hairs,
sparse on costules and veins, very short hairs and
FLORA MALESIANA
[ser. II, vol. 1°
glands between veins; short hairs on upper surface
of costae, very short erect hairs abundant between
veins, usually some glands also. Sori medial,
lower ones not divergent; indusia small with
glands and short hairs; sporangia with several
setae (to 10), sometimes also a gland.
Distr. Malesia: Sumatra, Java, Lesser Sunda
Is. (Bali, Flores), SW. & N. Celebes, Philippines
(Luzon).
Ecol. On rocks by streams or waterfalls at
400-1200 m.
Notes. D. brunnescens was based on a small
plant with pinnae to 4.5cm long. PRICE reports
red young fronds on plants in Luzon but I have
seen no such report from Java.
12. Sphaerostephanos pseudomegaphyllus
(v.A.v.R.) HOLTTUM, comb. nov. — Dryopteris
megaphylloides v.A.v.R. Bull. Jard. Bot. Btzg II,
20 (1915) 16, non ROSENST. 1913.— Dryopteris
pseudomegaphylla v.A.v.R. Handb. Suppl. (1917)
180. — Type: BURCK, Sumatra, Padang Pandjang
(BO).
Aspidium megaphyllum var. hirsutum METT.
Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 233; MIQUEL,
ibid. 4 (1869) 159. — Type: KORTHALS, Sumatra
(lz):
Differs from S. penniger: fronds smaller; pinnae
7x1 to 16x 2.2cm, lobed about 1/4-1/3 to costa;
veins 8-10 pairs, 13-2 pairs anastomosing, 13 pairs
to sides of sinus-membrane; dense spreading hairs
to 0.5mm long on lower surface of rachis and
costae; indusia densely hairy or with many glands
and fewer hairs; sporangia copiously setiferous.
Distr. Malesia: Sumatra (Padang Highlands).
13. Sphaerostephanos diversilobus (PRESL)
HOLTTUM, Kalikasan 4 (1975) 57. — Nephrodium
diversilobum PRESL, Epim. Bot. (1851) 47;
HOLTTUM, Novit. Bot. Univ. Carol. Prag. 1968
(1969) 39.—Dryopteris diversiloba (PRESL)
CHRIST, Philip. J. Sci. 2 (1907) Bot. 199, nomen
tantum; COPEL. Philip. J. Sci. 40 (1929) 297. —
Phegopteris diversiloba (PRESL) v.A.v.R. Handb.
(1908) 503.—Cyclosorus diversilobus (PRESL)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941)
244; COPEL. Fern Fl. Philip. (1960) 357.—
Thelypteris diversiloba (PRESL) REED, Phytolo-
gia 17 (1968) 273. — Lectotype (HOLTTUM 1969):
CUMING 102, Luzon (PRC; E, FI-W, G, K, LE).
Goniopteris asymmetrica FEE, Gen. Fil. (1852)
253.— Thelypteris asymmetrica (FEE) REED,
Phytologia 17 (1968) 261.—Specimens cited:
CUMING 51, 102; FEE’s specimens of these num-
bers not seen.
Dryopteris acromanes CHRIST, Philip. J. Sci. 2
(1907) Bot. 200.— Phegopteris acromanes
(CHRIST) v.A.v.R. Handb. (1908) 505. — Type not
selected; specimens cited (which include one of
CUMING 51) not seen.
Caudex short, creeping. Stipe of sterile frond to
1981]
20 cm, of fertile to 30cm, sparsely hairy. Reduced
pinnae 2 pairs, very small, seen in isotype at E
only. Lamina 25 cm long; pinnae 7-9 pairs; basal
pinnae gradually narrowed at base, basal basis-
copic lobe missing. Fertile pinnae to 6.5 x 2.0cm,
sterile to 8cm long, 3cm wide near apex; lower
pinnae and sometimes others widened and more
deeply lobed distally, some with irregularly elon-
gate subapical lobes. Middle pinnae with = dilated
base, abruptly short-pointed apex, lobed 2/3 to
costa apart from widened apical part, lobes hardly
falcate, normally entire; costules 4-5 mm apart;
veins 7-8 pairs except in elongate distal lobes. |
pair anastomosing, next acroscopic vein to sinus-
membrane; lower surface of rachis and costae
bearing short erect acicular hairs, also some
longer ones, especially on sterile pinnae; short
erect hairs between veins on both surfaces, glands
few on both surfaces, those on lower surface
usually confined to costules and veins. Sori
medial; indusia thin, densely hairy with a few
glands; sporangia with glands; spores with trans-
lucent wing and cross-wings.
Distr. Malesia: Philippines (Luzon, Panay), in
forest at 300-1000 m.
Notes. Some pinnae are irregularly lobed on
most specimens; this suggests hybridization. One
sterile frond of CUMING 51 at Kew has very long
distal lobes which are crenate with pieocnemioid
venation. The only specimen seen with reduced
basal pinnae is one of CUMING 102 at Edinburgh.
The specimens listed by CHRIST in 1907 under D.
diversiloba represent several other species; he
re-described PRESL’s species as D. acromanes
from CUMING S51 and other specimens now ap-
parently lost.
14. Sphaerostephanos’ dichrotrichus (COPEL.)
HOLTTUM, comb. nov. — Dryopteris dichrotricha
COPEL. Philip. J. Sci. 6 (1911) Bot. 74. — Thelyp-
teris dichrotricha (COPEL.) REED, Phytologia 17
(1968) 272. — Type: C. KING 294, Papua “main-
land opposite Samarai island’ (MICH: BRI,
NSW).
Caudex short-creeping. Stipe 40cm to first
reduced pinna, dull reddish with pale hairs 0.5 mm
or more long. Lamina 65 cm or more long: pinnae
to 20 pairs, basal 1-3 pairs gradually reduced,
lowest 14cm long: aerophores slightly swollen.
Largest pinnae 182.0cm; base truncate, apex
acuminate with entire cauda to 1.5 cm long; edges
lobed +3 way to costa, lobes slightly falcate with
round tips; costules 4.5-Smm apart, at 60° or
more; veins 7 pairs, 1-1: pairs anastomosing, next
+1 pair to sinus-membrane; rather sparse hairs
1mm long and many shorter ones on lower sur-
face of rachis and costae, shorter on costules and
veins, glands and short erect hairs between veins:
minute erect hairs and glands between veins on
upper surface. Sori medial; indusia thin, with
many hairs, also glands; sporangia usually with
THELYPTERIDACEAE (Holttum)
453
1-2 slender setae (rarely a gland) on body, on stalk
a sessile gland; spores with a thin translucent wing
and cross-wings.
Distr. Malesia: extreme east of New Guinea,
Admiralty Islands, at low altitudes.
Notes. Some specimens have more erect hairs
than glands on upper surface. In one (LAE 65333)
there are 2 pairs of small pinnae near base of stipe
and then a gap of 20cm.
15. Sphaerostephanos (COPEL.)
HOLTTUM, comb. nov.— Dryopteris vestigiata
COPEL. Un. Cal. Publ. Bot. 18 (1942) 220.—
Cyclosorus vestigiatus (COPEL.) COPEL. Gen.
Fil. (1947) 143; Philip. J. Sci. 78 (1951) 446, pl.
27.— Thelypteris vestigiata (COPEL.) REED,
Phytologia 17 (1968) 323.— Type: BRAsSs 12317,
W. New Guinea, Idenburg River, 1700 m, in forest
(MICH; BM, L).
Caudex slender, erect, to 60cm tall. Stipe to
50cm long, glandular, hairless. Reduced pinnae 3
pairs, all very small, Scm apart. Lamina more
than 100cm long; pinnae 40 pairs, not dimor-
phous, basal pinnae somewhat narrowed at base,
with stalks to 1 mm long, aerophores not swollen.
Largest pinnae 21 x 2.2 cm, base broadly cuneate,
apex caudate-acuminate (cauda 2.5-5.0 cm suben-
tire) lobed to 2-2.5mm from costa, lobes hardly
falcate with rounded tips; costules 44.5 mm apart
at 60° or more; veins 9-11 pairs, 1 pair anas-
tomosing, next veins both to edge: hairs on lower
surface of rachis and costae 0.1 mm long (longer
distally on costules), surface between veins
copiously glandular; upper surface of costules and
veins with scattered hairs like those on costae,
between veins minute erect hairs and glands. Sori
medial or little supramedial: indusia small, thin,
with glands only; sporangia bearing glands on
body and gland-tipped hairs on stalk; spores with
many small wings.
Distr. Malesia: Several collections from east-
ern New Guinea at c. 1500 m.
Note. Some specimens have quite hairless
lower surface of rachis and base of costae.
vestigiatus
16. Sphaerostephanos morotaiensis HOLTTUM,
sp. nov.
Caudex brevis, repens: stipes Scm longus;
pinnae redactae usque 12-jugatae, patentes,
superiores 1.5-2 cm longae: lamina 50cm longa:
pinnae 16-jugatae, maximae 7.51.5 cm, subin-
tegrae; venae 4-5-jugatae: pagina inter venas
utrinque pilis minutis erectis glandulisque
praedita; sori mediales; indusia brevi-pilosa;
sporangia glandulifera. — Type: MAIN & ADEN
758, Morotai, Totodahu, 30m (K; BO, L, SING).
Caudex short-creeping. Stipe S5cm_ long,
minutely hairy; base of stipe to first normal pinnae
50cm. Reduced pinnae 1.5-2.5cm apart, to 12
pairs; upper ones 1.5-2cm long, spreading,
broadly triangular, entire: lowest 4mm _ long.
454
Lamina 50cm long; pinnae 16 pairs, lower ones
not narrowed at base nor auricled; aerophores not
elongate. Largest pinnae 7.5 x 1.5cm; base trun-
cate, sometimes a little dilated; apex short-acu-
minate; edges subentire; costules 3—3.5 mm apart
on fertile pinnae, 4.5mm on sterile; veins 4—5
pairs, 3-3; pairs anastomosing, sinus-membrane
hardly detectable; whole lower surface (including
rachis) covered with very short erect hairs and
glands; hairs on upper surface of costae 0.3-
0.4 mm long, on costules and veins sparse, surface
between veins with very short erect hairs and
glands. Sori medial; indusia small with many short
hairs; sporangia bearing glands.
Distr. Malesia: Moluccas. Apart from the type
and other specimens from Morotai, I refer here
SA-AANAM 49 from Obi Island (L) which has
larger crenate pinnae (to 18 x 3.0 cm), agreeing in
other respects.
17. Sphaerostephanos kalkmanii HOLTTUM, sp.
nov.
Caudex erectus; pinnae redactae 4-jugatae,
omnes minutae; pinnae normales 18-jugatae,
usque 3.5X1.0cm, crenatae, costae costulaeque
subtus pilis minutis appressis glandulisque
praeditae; indusia parva vel nulla; sporangia nec
glandulis nec setis instructa. —Type: KALKMAN
4070, W. New Guinea, Orion Mts, Tenma River,
1500 m (BM; LAE, L, SING).
Caudex erect, to at least 15cm tall. Stipe 4-
5 cm long, dark, sparsely short-hairy, basal scales
c. 4X 1mm; base of stipe to first large pinna
10-15 cm. Reduced pinnae to 4 pairs, all about
1mm long. Lamina 20cm long; pinnae 18 pairs,
subcoriaceous, lower ones not narrowed at base;
aerophores slightly swollen. Hairs on lower sur-
face of rachis 0.3mm long, brown, on upper sur-
face a little longer. Largest pinnae 2.6-3.5cm
long, 0.7-1.0 cm wide, fertile ones sometimes, not
always, smaller than sterile; base subtruncate to
broadly cuneate; apex rather abruptly short-poin-
ted; edges crenate, a little more deeply in fertile
than in sterile pinnae; costules 2mm apart; veins
all very oblique, 4-5 pairs, 13 pairs anastomosing,
next vein to short sinus-membrane; lower surface
of costae and costules covered with minute ap-
pressed hairs, also glands, a few glands sometimes
present on surface between veins; hairs on upper
surface of costae to 0.5 mm long, on costules and
veins shorter, sparse, a few glands sometimes
present (?) on surface between veins. Sori medial,
sometimes covering whole surface at maturity;
indusia very small with short hairs and glands,
sometimes lacking; sporangia bearing neither
glands nor setae; spores minutely spinulose.
Distr. Malesia: West New Guinea, 1500 m.
Only known from type.
Ecol. “In abandoned garden, rather common”.
Note. Possibly nearest to the lowland species
S. lamii, which has glands between veins on both
FLORA MALESIANA
[ser. II, volaale
surfaces. S. lamii has longer pinnae, upper
reduced pinnae 10mm long, and veins not very
oblique.
18. Sphaerostephanos perglanduliferus (v.A.v.R.)
HOLTTUM, Kalikasan 4 (1975) 59.— Dryopteris
perglandulifera v.A.v.R. Bull. Jard. Bot. Btzg III,
2 (1920) 150.—Thelypteris perglandulifera
(v.A.v.R.) REED, Phytologia 17 (1968) 303.—
Type: BROOKS 281, Sumatra, Lebong Tandai,
Benkoelen (BO).
D. perglandulifera var. firmior v.A.v.R. l.c.—
Type: DOCTERS VAN LEEUWEN-REYNVAAN
3284, Sumatra, Haboko (BO).
Phegopteris pseudoarfakiana HOSOKAWA, Tr.
Nat. Hist. Soc. Formosa 28 (1938) 147. — Menis-
cium pseudoarfakianum (HOSOKAWA) HOSOK-
AWA, ibid. 32 (1942) 286.— Type: HOSOKAWA,
Caroline Is., Palau (TAI).
Dryopteris arbuscula sensu KJELLBERG & C.
CHR. Bot. Jahrb. 66 (1933) 43.
Caudex massive, short, erect; stipe 5-15cm
long, densely short-hairy; basal scales 10 mm long,
narrow; base of stipe to first normal pinna 20-
50cm; reduced pinnae to 10 pairs, lowest 7 x
7mm, upper ones to 1.51.5 cm, base broadly
cuneate, symmetric, apex obtusely pointed.
Lamina of type 75cm long; transition at base to
reduced pinnae subabrupt; lowest normal pinnae
not auricled at base; aerophores not elongate.
Largest pinnae of type 14x1.7cm (on small
plants, fertile, 71cm), base broadly cuneate,
apex short-acuminate, edges shallowly crenate;
costules 3-4 mm apart, at c. 45° to costa; veins 6—7
pairs, 3 pairs anastomosing to form a slightly
Zig-zag excurrent vein, | pair to sides of sinus-
membrane; lower surface of rachis covered with
slender appressed hairs less than 0.5mm _ long,
hairs on costae minute, glands throughout lower
surface; hairs on upper surface of rachis pale,
1mm long, on costae less than 0.5 mm, copious
glands between veins. Sori medial; indusium
bearing glands; sporangia with short setae and
glands, a gland at end of short hair on sporangium-
stalk.
Distr. Malesia: S. Sumatra, Borneo, S.W. Cele-
bes, Philippines (Negros) and Micronesia (Palau).
Ecol. In rocky stream-beds at low to medium
altitudes.
Note. This is closely related to S. penniger,
but appears to be distinct in its erect caudex (seen
on several specimens) and narrow crenate pinnae;
the stream-bed habitat is recorded for several
specimens.
19. Sphaerostephanos veitchii HOLTTUM, AIl-
lertonia 1 (1977) 204.—Type: J. G. VEITCH,
Solomon Islands, San Cristobal (K).
Nephrodium amboinense var. subglandulosum
BAKER, J. Linn. Soc. Bot. 19 (1882) 295. — Type:
R. B. COMINS, Solomon Islands (K).
1981]
THELYPTERIDACEAE (Holttum)
455
Caudex short, creeping or suberect. Stipe of
type lacking, of others 10-20cm, minutely hairy;
base of stipe to first large pinna 35-60 cm.
Reduced pinnae about 7 pairs, lowest 5 mm long,
upper ones 1.5—2.5cm long and wide, broadly
triangular with asymmetric base (wider on acros-
copic side), crenate. Lamina 50-90 cm or more
long; basal pinnae not narrowed at base; aero-
phores not elongate. Largest pinnae of type 13 x
1.7cm, of others to 20 x 2cm; base broadly trun-
cate to full width, sometimes a little dilated but
not auricled; apex acuminate, not caudate; edges
crenate to depth of 1-I.5mm; costules 4mm
apart, at 60° to costa; veins 6 pairs, 23. pairs
anastomosing to form a zig-zag excurrent vein,
15-2 pairs to sides of sinus-membrane; hairs on
lower surface of rachis 0.4 mm long (much shorter
on some plants from New Guinea), pale, curved,
on costae and costules shorter, antrorse, many
glands and short erect hairs between veins; upper
surface of rachis covered with brown hairs more
than 0.5mm long, similar hairs on costae and
scattered on costules and veins, glands on surface
between veins and sometimes short erect hairs.
Sori medial, lower ones divergent, those on basal
veins from adjacent costules sometimes confluent;
indusia small, bearing short hairs and glands;
sporangia bearing glands.
Distr. Solomon Islands, and Malesia: Eastern
New Guinea.
Ecol. In the Solomons by streams in low
country and to 800m; in New Guinea plants in
forest at higher altitudes (to 1400 m) are smaller
but otherwise not distinct.
20. Sphaerostephanos lamii HOLTTUM, sp. nov.
Caudex brevis, repens vel suberectus; stipes
20cm longus; pinnae redactae 3-5-jugatae,
superiores 10mm longae; lamina 30cm longa,
pinnae 12-15-jugatae, dimorphae; pinnae Steriles
usque 8.5X1.8cm, crenatae, pagina utrinque
glandulifera; pinnae fertiles usque 6.0 1.2cm;
sori inframediales; indusia parva, pilis brevibus
glandulisque praedita; sporangia glandulifera. —
Type: H. J. LAM 748, W. New Guinea, Prauwen
bivak, 10m (L; BO).
Caudex short, creeping or suberect; stipe 12-
20 cm long, short-hairy in groove; base of stipe to
first normal pinna 30-40 cm; reduced pinnae 3-5
pairs, not opposite, upper ones to 10mm long,
deflexed with slightly auricled asymmetric base,
lowest 3mm long. Lamina 30cm long; pinnae
8-15 pairs, more or less dimorphous, basal pinnae
not narrowed at base, not auricled; aerophores not
elongate; rachis bearing stiff brown hairs to 1 mm
long on both surfaces. Sterile pinnae to 8.5 x
1.8 cm, base truncate, apex short-acuminate, edges
crenate to a depth of 1mm; costules 3.5-4mm
apart, at 60° to costa; veins 5-6 pairs, 23 pairs
anastomosing, one pair to sides of sinus-mem-
brane; on lower surface hairs at base of costae
very short, distally 0.2-0.3mm (but see note
below), rest of lower surface of pinnae glabrous
with sessile glands; hairs on upper surface of
costae hardly 0.5 mm long, rest of upper surface
glabrous or with some minute hairs, glands
present throughout but fewer than on lower sur-
face, sometimes lacking. Fertile pinnae to 6.0 x
1.2 cm, slightly crenate; veins 3-4 pairs; sori near
costules, basal ones not divergent; indusia small,
with very short hairs and a few glands; sporangia
bearing glands.
Distr. Malesia: W. New Guinea, several col-
lections from Rouffaer and Mamberamo Rivers, at
10-200 m; Admiralties (Los Negros: GRETHER &
WAGNER 4003); a distinct variety in Eastern New
Guinea?
Note. A specimen from 820m in the Southern
Highlands of Eastern New Guinea differs in the
presence of hairs 1 mm long on the lower surface
of costae and scattered long hairs on costules and
veins on the upper surface.
21. Sphaerostephanos hirsutus (KUNZE ex METT.)
HOLTTUuUM, Kalikasan 4 (1975) 63. — Nephrodium
hirsutum PRESL, Epim. Bot. (1851) 48, non D.
DON 1825; HOOK. Sp. Fil. 4 (1862) 70 p.p. & t.
240B. — Aspidium hirsutum KUNZE ex METT.
Farngatt. IV (1858) 107, nom. nov. — Dryopteris
hirsuta (KUNZE ex METT.) O. KTZE, Rev. Gen.
Pl. 2 (1891) 813; v.A.v.R. Handb. (1908) 214. —
Dryopteris adenophora C. CHR. Ind. Fil. (1905)
251, nom. nov. superfl. — Cyclosorus adenophorus
(C. CHR.) CHING, Bull. Fan Mem. Inst. Biol. Bot.
10 (1941) 242; CopEL. Fern FI. Philip. (1960)
343. — Thelypteris adenophora (C. CHR.) REED,
Phytologia 17 (1968) 258.— Type: CUMING 82,
Luzon (PRC; K, P).
KEY TO THE VARIETIES
1. Pinnae commonly to 202.2cm; sporangia
glanduliferous ae . a. var. hirsutus
1. Pinnae to 15*1.4cm; sporangia setiferous
b. var. celebicus
a. var. hirsutus
Caudex short, erect, with many branches at its
base; stipe short; base of stipe to first normal
pinna 30-90 cm, basal scales broad, thin, not per-
sistent; reduced pinnae to 12 pairs or more, each
consisting of an aerophore with a very small
rounded blade, uppermost less than 1 cm long.
Lamina to 120cm long; transition at base to
reduced pinnae abrupt or with an intermediate
pair; lowest normal pinnae somewhat narrowed at
their bases and sometimes slightly auricled;
aerophores to Imm _ long. Largest pinnae
commonly 12-20 x 1.5-2.2 cm (to 30 x 2.5 cm, teste
COPELAND), base truncate and somewhat dilated;
apex caudate-acuminate (cauda 1.5-2.5 cm long),
lobed 2/5-3/5 towards costa; lobes distinctly fal-
456
FLORA MALESIANA
[ser. II, vol. 1°
cate with rounded or obtuse-pointed tips; costules
3-4 mm apart, at 60° or more to costa; veins 7-8
pairs, | pair anastomosing, next pair to sides of
sinus-membrane; lower surface of rachis and cos-
tae with a variable number of long coarse hairs
(sometimes only distally on costae) with very
short hairs and glands, costules similar, abundant
glands between veins; hairs on upper surface of
rachis and costae 1.5mm or more long, scattered
hairs 1mm long on costules and veins, between
veins abundant appressed hairs and a variable
number of glands (often few). Sori medial, lower
ones not divergent; indusia small. with glands and
short hairs; glands on sporangia.
Distr. Malesia: throughout
forest, at 400-1200 m.
Note. Pubescence in var. hirsutus is variable;
it is most clearly distinguished from S. todayensis
(which see), also variable in pubescence, by the
small size of its basal reduced pinnae. A more
nearly allied species is S. trichochlamys.
Philippines, in
b. var. celebicus HOLTTUM, var. nov.
A typo speciei differt: pinnis usque 15 x 1.4cm;
pagina inferiore inter venas pilis tenuibus ad-
pressis vestita; indusiis nullis vel minutis; spo-
rangiis 3-4 setis ornatis.— Type: POSTHUMUS
2743, S.W. Celebes, G. Bonthain, 1500m (BO).
Distr. Malesia: S. W. and Central Celebes &
Lesser Sunda Is. (Flores: JAAG 1647).
22. Sphaerostephanos acrostichoides (DESV.)
HOLTTUM, comb. nov.— Nephrodium acrosti-
choides DESV. Mém. Soc. Linn. Paris 6 (1827)
255. (Not Thelypteris acrostichoides (MICHX)
NEWL. nom. inval.). — Type: Collector unnamed,
Timor (P).
Polystichum riedleanum GAUD. in Freyc. Voy.
Ur. Phys. Bot. (1828) 327. — Dryopteris riedleana
(GAUD.) v.A.v.R. Handb. (1908) 230.— Type:
GAUDICHAUD, Timor (G).
Nephrodium smithianum PRESL, Epim. Bot.
(1851) 46; HOLTTUM, Novit. Bot. Inst. Bot. Univ.
Carol. Prag. 1968 (1969) 41 excl. syn. Aspidium
productum KAULF.—S. smithianus (PRESL)
HOLTTuM, Kalikasan 4 (1975) 65. — Type: CUM-
ING 279, Guimaras (PRC; B, BM, E, G, K, L).
Dryopteris perpilifera v.A.v.R. Bull. Jard. Bot.
Btzg II, 11 (1913) 12; Handb. Suppl. (1917) 183. —
Thelypteris perpilifera (v.A.v.R.) REED, Phytolo-
gia 17 (1968) 303. — Type: LAUTERBACH 68, N.E.
New Guinea, Finschhafen (BO; BRI, L, SING).
Dryopteris angusta COPEL. Philip. J. Sci. 9 (1914)
Bot. 3; v.A.v.R. Handb. Suppl. (1917) 186. —
Thelypteris angusta (COPEL.) REED, Phytologia 17
(1968) 259. — Type: C. KING 408, E. New Guinea
(MICH; K, NSW).
Dryopteris pseudoarbuscula yv.A.v.R. Philip.
J. Sci. 11 (1916) Bot. 106; Handb. Suppl. (1917)
504.— Type: C. B. ROBINSON 1962, Amboina
(BOskKe ea):
Dryopteris riparia COPEL. Univ. Cal. Publ. Bot.
18 (1942) 221.—Cyclosorus riparia (COPEL.)
COPEL. Gen. Fil. (1947) 143; Philip. J. Sct. 78
(1951) 456, pl. 37. — Thelypteris riparia (COPEL.)
REED, Phytologia 17 (1968) 309. — Type: BRASS
13048, New Guinea, Idenburg River, 800m (UC;
BM, BO, L, LAE).
Cyclosorus boholensis COPEL. Philip. J. Sci. 81
(1952) 31, pl. 23; Fern FI. Philip. (1960) 348. —
Thelypteris boholensis (COPEL.) REED, Phy-
tologia 17 (1968) 204.— Type: RAMOS BS 42990,
Bohol (UC; G).
Cyclosorus reederi COPEL. Amer. Fern J. 43
(1953) 12.— Thelypteris reederi (COPEL.) REED,
Phytologia 17 (1968) 308. — Type: REEDER 889,
N.E. New Guinea, Finsch-hafen (US).
Aspidium arbuscula sensu CHRIST, Ann. Jard.
Bot. Btzg 15 (1898) 135.— Cyclosorus hispidula
sensu COPEL. Philip. J. Sci. 78 (1951) 455. — Cyclo-
sorus productus sensu COPEL. Fern Fl. Philip.
(1960) 348, p.p.
Caudex erect, not or little branched at the base.
Base of stipe to lowest reduced pinna 3-10 cm, to
lowest large pinna 10-30cm or more; at base of
lamina a subabrupt transition to reduced pinnae;
number of reduced pinnae varying with size of
frond, larger ones spreading, triangular with a
broad base. Lamina of smaller plants fertile at
20 cm long with 15 pairs pinnae, of large plants to
50cm long. Largest fertile pinnae of small plants
3-5 x 0.7-1.0 cm, of largest plants to 10 x 1.0 cm or
larger; base broadly cuneate and + dilated on both
sides, apex evenly tapered, edges lobed 1/3-1/2
way to costa; costules at less than 60° to costa,
2-2;mm apart in small plants; veins 3-4 pairs in
small plants, 7-8 pairs in large ones, I-12 pairs
anastomosing, next acroscopic vein to sinus
membrane; lower surface of rachis with copious
spreading pale hairs 1 mm long and shorter ones,
long hairs less abundant on costae, lower surface
between veins bearing copious glands and some
erect short hairs; upper surface of rachis with pale
hairs |mm long, shorter hairs on costae, also
scattered on costules and veins, between veins
copious short erect hairs and glands. Sori medial,
on small pinnae covering lower surface; indusia
hairy and glandular; sporangia bearing glands.
Distr. Malesia: East Java, Lesser Sunda Is.
(Sumba, Timor), Central & S.E. Celebes, Philip-
pines (Bohol, Mindanao, Guimaras), Moluccas
(Amboina), and New Guinea incl. Admiralty Is.
Ecol. Usually in rocky stream-beds at 0-500 m
(to 1500 m in Mindanao and New Guinea). In New
Guinea plants of all sizes have been collected, but
elsewhere none as large as the larger ones in New
Guinea. Types of the various basionyms differ
chiefly in size, agreeing well in frond-form, dilated
bases of pinnae, pubescence and the distribution
of glands. If the larger New Guinea plants (which
apparently do not grow in stream-beds) are judged
to represent a distinct species, the name Dry-
1981]
THELYPTERIDACEAE (Holttum)
457
opteris perpilifera v.A.v.R. should be _ its
basionym.
23. Sphaerostephanos melanorachis HOLTTUM,
sp. nov.
Stipes rachisque atrobrunneus; stipes 7cm
longus; pinnae redactae 10-jugatae, superiores
usque 1.7X1.5cm; lamina 55cm longa; pinnae
dimorphae, steriles usque 13X1.9cm, _ fertiles
minores, omnes c. 1/3 costam versus lobatae;
venae 8-jugatae, 2-jugatae anastomosantes ; rachis
costaeque subtus pilis brevibus antrorsis vestita;
pagina inter venas utrinque glandulosa; sori
mediales; indusia pilis paucis praedita.— Type:
A. H. G. ALSTON 16927, Moluccas, Batjan, Mt
Sibela 750 m (BM).
Caudex short, apparently erect. Stipe 7cm,
dark brown, minutely hairy; base of stipe to first
large pinna 35-40cm; rachis also very dark.
Reduced pinnae 10 pairs, upper ones 1.7 x 1.5 cm,
triangular with asymmetric base, crenate, slightly
deflexed, lowest 4-5 mm long. Lamina 55 cm long;
pinnae 25 pairs, dimorphous; aerophores not
elongate. Sterile pinnae to 13 cm long, 1.9 cm wide
above truncate dilated base, apex caudate-acu-
minate (cauda to 1.5cm); edges lobed to a depth
of 3mm (c. 1/3), lobes slightly falcate; costules
4mm apart, to 60°; veins to 8 pairs, 2 pairs anas-
tomosing, forming a zig-zag excurrent vein, | pair
to sinus-membrane; lower surface of rachis bear-
ing rather sparse pale appressed hairs 0.5mm
long, those on costa also antrorse, 0.2 mm long,
sparse short appressed hairs on costules and
veins; glands on surface between veins; hairs on
upper surface of rachis pale, | mm long, on costae
a little shorter, similar hairs scattered on costules
and veins, glands between veins. Fertile pinnae to
9.5cm long, 1.6cm wide at base, 1.2cm above
base; sori medial; indusia rather small, thin, with a
few short hairs; sporangia rarely with a gland.
Distr. Malesia: Moluccas (Batjan, Mt Sibela),
750 m.
24. Sphaerostephanos heterocarpus (BL.) HOLT-
TUM in Nayar & Kaur, Comp. to Bedd. (1974)
209.— Aspidium heterocarpon BL. Enum. PI.
Jav. (1828) 155; MeETT. Ann. Mus. Bot. Lugd. —
Bat. 1 (1864) 233.— Nephrodium heterocarpum
(BL.) MOORE, Ind. Fil. (1858) 93; RACIB. FI. Btzg
1 (1898) 187.— Dryopteris heterocarpa (BL.) O.
KTZE, Rev. Gen. Pl. 2 (1891) 813; v.A.v.R.
Handb. (1908) 228; C. CHR. Gard. Bull. Str. Settl.
4 (1929) 380; BACKER & POSTH. Varenfl. Java
(1939) 54.—Cyclosorus heterocarpus (BL.)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 180;
HOLTTUM, Rev. Fl. Malaya 2 (1955) 269, f. 155;
TARD. FI. Gén. I.-C. 7, pt. 2 (1941) 385; COPEL.
Fern Fl. Philip. (1960) 345, p.p.— Thelypteris
heterocarpa (BL.) MORTON, Amer. Fern J. 49
(1959) 113.—Type: BLUME, Java, ad pedem
Boerangrang (L, n. 908, 332-1158).
Polypodium longifolium ROxB. Calc. J. Nat.
Hist. 4 (1844) 462.— Type from Amboina (BR).
Nephrodium pubescens BRACK. in Wilkes U.S.
Expl. Exp. 16 (1854) 186, non DON 1825. — Type:
U.S. Expl. Exp. Luzon, near Banos (US).
Dryopteris bordenii CHRIST, Philip. J. Sci. 2
(1907) Bot. 204; v.A.v.R. Handb. (1908) 822. —
Cyclosorus bordenii (CHRIST) CHING, Bull. Fan
Mem. Inst. Biol. Bot. 10 (1941) 243; COPEL. Fern
Fl. Philip. (1960) 345, p.p.— Thelypteris bordenii
(CHRIST) REED, Phytologia 17 (1968) 264.— S.
heterocarpus var. bordenii HOLTTUM, Kalikasan
4 (1975) 67.—Lectotype (HOLTTUM 1975):
BORDEN 1237, Luzon, Mt Mariveles (US).
Dryopteris suprastrigosa ROSENST. Fed.
Rep. 10 (1912) 335.—Cyclosorus suprastrigosus
(ROSENST.) COPEL. Gen. Fil. (1947) 143; Philip.
J. Sci. 78 (1951) 446. — Thelypteris suprastrigosa
(ROSENST.) REED, Phytologia 17 (1968) 318. —
Type: Frau BAMLER 37, N.E. New Guinea, Sat-
telberg (not seen; at BM is KEYSSER 37 from
Sattelberg, named by ROSENST.).
Dryopteris mixta ROSENST. Fed. Rep. 12 (1913)
172. — Lastrea mixta (ROSENST.) COPEL. Philip.
J. Sci. 78 (1951) 426.— Thelypteris mixta
(ROSENST.) REED, Phytologia 17 (1968) 293. —
Type: KEYSSER S. 141, N.E. New Guinea, Sat-
telberg (S-PA). — Fig. 1q.
Caudex erect, rarely very tall, usually with
several branches from base. Stipe 5cm or little
more. Reduced pinnae commonly 10 pairs (dis-
tance apart, and number, depending on size of
plant), never less than 6 pairs, uppermost com-
monly 10mm long, narrow, auricled, lowest very
small. Lamina variable in length; pinnae com-
monly 20-30 pairs; several pairs lower pinnae
much narrowed towards their bases, basal one at
most slightly auricled; aerophores slightly swollen.
Largest pinnae 10-20cm long, 1.2-2.5cm wide,
fertile sometimes narrower than sterile and with
shorter hairs, lobed 5/8-3/4 towards costa; cos-
tules 3-3; mm apart; veins 6-12 pairs, basal veins
anastomosing at least in basal half of pinna with
excurrent vein to sinus, second acroscopic vein
sometimes touching side of sinus-membrane; hairs
on lower surface of costae very short, erect, with
scattered longer ones usually not over 0.5mm
long, more sparse similar hairs on costules and
veins, copious yellow glands and a variable num-
ber of short erect hairs between veins; upper
surface of costae bearing antrorse pale hairs to
1 mm long with scattered similar hairs on costules
and veins; surfaces between veins bearing few to
many appressed hairs 0.3-0.4 mm long, sometimes
also glands which are never as abundant as on
lower surface. Hairs on upper surface of rachis
pale to brownish, curved, not over | mm long; on
lower surface erect hairs as on costae. Sori
medial, lower ones not divergent; indusia firm,
usually with a few short hairs and glands;
sporangia with glands near annulus.
458
FLORA MALESIANA
[ser. Il, voluale
Distr. Peninsular Thailand; Hainan, Hong Kong;
throughout Malesia; North Queensland, Solomon
Islands, New Hebrides, Fiji, Samoa.
Ecol. In forest, low country to 1500 m.
Notes. As here interpreted, this is a very
variable species and varieties are not always
sharply distinct; no formal names for them are
here given. The following key attempts to dis-
tinguish the various forms.
KEY TO) THE LOCAL FORMS
1. Lower pinnae lobed at least 2/3 towards costa
and at least 1.5 cm wide.
2. Glands none or rare on upper surface of
pinnae.
3. Fertile pinnae of large plants 2cm wide
(sterile ones sometimes wider); costules
4mm apart, second acroscopic vein usually
touching side of sinus-membrane
typical form
3. Fertile pinnae not over 1.5 cm wide; costules
3mm apart; second acroscopic vein always
to margin (Dryopteris suprastrigosa
ROSENSDID) ee) ee oe New Guinea
2. Glands + abundant on upper surface of
pinnae.
4. Appressed hairs to 0.5mm _ long between
veins on upper surface . Borneo
4. Short suberect hairs between veins on upper
surface Kipel. eee. . Moluccas
1. Lower pinnae lobed not or little more than 1/2
way to costa, rarely more than 1.5 cm wide.
5. Glands and appressed hairs (sometimes dense)
on upper surface . .Borneo (limestone)
5. Glands lacking on upper surface; appressed
hairs variable, never dense.
6. Pinnae lobed distinctly more
second acroscopic vein to margin.
7. Pinnae to 16 x 1.6 cm; costules almost at right
angles to costa; pinnae more than 20 pairs;
sori supramedial . Mindanao
7. Pinnae to 10 x 1.3 cm; costules at 60°; pinnae
than 1/2;
to 15 pairs; sori medial (Nephrodium
pubescens BRACK., Dryopteris bordenii
CHRIST) Eee? (oa ore ee Luzon
6. Pinnae lobed 1/2 or less deeply; second
acroscopic vein usually ending beside sinus-
membrane lowlands of W. Malesia
The typical form occurs in mountain forest in
Western Malesia and to Flores. In Malaya there
seems to be no sharp distinction between the
mountain form and the lowland one which comes
at the end of the key. Plants like the lowland form
of Malaya occur in Thailand, Hainan and Hong
Kong, also in the lowlands of Borneo.
BLUME distinguished varieties B and C in Java.
I have found no authentic specimen of the former;
var. C has rather large almost sterile fronds with sori
on basal veins only and is not a distinct variety.
The Borneo form with many glands and ap-
pressed hairs on the upper surface occurs in
Sarawak and Sabah at low to medium altitudes.
The limestone Bornean form is abundant in the
Gunong Mulu National Park; at higher altitudes
and in more exposed positions fronds are smaller
and very densely covered with hairs on the upper
surface, but there seems to be no sharp distinction
between these and plants with larger fronds less
densely covered with hairs.
In the Philippines neither the typical form nor
the lowland form of Western Malesia has been
found. From Palawan is one collection somewhat
resembling the Borneo form; from Mindanao are
two specimens (RAMOS & PASCASIO BS 34476,
Surigao Province, and EDANO PNH 12581, Davao
Province) which seem distinct and are keyed as
the Mindanao form, but a third (COPELAND 1643,
San Ramon, 500m) is nearer to the New Guinea
form. In Luzon the form represented by Dry-
opteris bordenii (small, with short fronds) appears
common; in herbaria it has been confused with the
large specimens of S. lobatus (COPEL.)
HOLTTUM.
Dryopteris suprastrigosa appears to be the
common form in eastern New Guinea; few col-
lections have been made in the west. Specimens
from North Queensland are similar but less hairy
on the upper surface of pinnae. D. mixta was
described from a small fertile plant with most
veins free but does not appear distinct in other
characters.
The Moluccan form is represented by BROOKS
18112 (Amboina) and KORNASSI 1222 (West
Ceram).
BACKER & POSTHUMUS cite Aspidium
dimorphum KUNZE as a synonym. This was
based on a specimen of JUNGHUHN’s from Mt
Pangerango in W. Java; I have not found it.
METTENIUS (l.c. 1864) described it briefly as a
synonym of A. heterocarpon var. B BL. He stated
that its aerophores were enlarged, for which
reason I doubt whether it belongs here.
25. Sphaerostephanos isomorphus HOLTTUM, sp.
nov.
Stipes ignotus ; pinnae redactae 6-jugatae, omnes
parvae; lamina 90 cm long; pinnae inferiores basi
non angustatae; pinnae maximae 14.5 x 2.0 cm,
profunde lobatae; venae 12-jugatae, infimae anas-
tomosantes; costae subtus pilis erectis minutis
vestitae; pagina inter venas utrinque glandulosa;
indusia parva, glandulifera.-_Type: FLENLEY,
ANU 2062, Papua New Guinea, W. Highlands, Lai
valley (K).
Caudex and stipe not known. Reduced pinnae 6
pairs, 4cm apart, all very small with enlarged
aerophores; one intermediate pair of pinnae also
present. Lamina 90 cm long; pinnae 40 pairs, light
green when dry, firm; lower pinnae not narrowed
at base. Largest pinnae 14.5 x 2.0 cm, sterile and
1981]
fertile isomorphous; base truncate and slightly
dilated; apex caudate-acuminate (cauda 22.5 cm),
edges lobed to 2mm from costa, lobes slightly
falcate; costules 3-3.5 mm apart, at more than 60°;
veins to 12 pairs, basal pair only anastomosing
with excurrent vein to short sinus-membrane, next
veins both to edge; lower surface of rachis, costae
and costules bearing very short erect hairs and
glands, between veins many glands but no hairs;
hairs on upper surface of costae pale, less than
1 mm long, scattered hairs 0.5 mm long on costules
and veins, glands between veins. Sori medial,
lower ones divergent, those on basal veins from
adjacent costules often touching; indusia very
small, glandular; sporangia bearing glands.
Distr. Malesia: Papua New Guinea, only known
from the type.
26. Sphaerostephanos dimidiolobatus HOLTTUM,
sp. nov.
Pinnae redactae usque 12-jugatae, superiores
2.5-3.0 cm longae; aerophori non elongati; pinnae
normales usque 27 X2.3cm, dimidio costam ver-
sus lobatae; venae 8—10-jugatae, infimae solum
anastomosantes; costae costulaeque subtus
glanduliferae, glandulae inter venas paucae;
pagina superior inter venas glandulis pilisque
minutis praedita; indusia parva; sporangia
glandulifera. —Type: J.R. CRorr 324, New
Ireland, 7km WNW of Taron, 700m (K; NSW).
Stipe 22-27 cm long, almost glabrous, slightly
flushed with red, scales thin, c. 10 x 2 mm; base of
stipe to first normal pinna 50-55 cm; reduced pin-
nae 8-12 pairs, uppermost 2.5-3.0cm_ long,
broadly triangular with subtruncate base slightly
auricled both sides, edges crenate, lowest reduced
pinna | cm long. Lamina excluding basal pinnae c.
100cm long; basal pinnae slightly narrowed in
basal third, basal lobes both slightly elongate;
aerophores not elongate. Largest pinnae 20-27 cm
long, to 2.3 cm wide; base truncate, not dilated nor
auricled; apex narrowly acuminate, not caudate;
edges lobed about half-way to costa (on one frond
slightly more than half), lobes hardly falcate; cos-
tules 4.5-S mm apart, at more than 60° to costa;
veins 8-10 pairs, slender, slightly prominent when
dry, basal pair at a broad angle to costule, anas-
tomosing with an excurrent vein to the sinus, next
pair usually both to sides of sinus-membrane;
lower surface of rachis bearing very short hairs
and some to 0.5mm long, costae minutely hairy
near base only, with glands throughout, few short
hairs and many glands on costules, few glands
between veins; upper surface of rachis and costae
bearing hairs 0.5mm long, minute hairs on cos-
tules and veins, between veins a variable number
of glands and very short erect hairs. Sori medial;
indusia small, shrivelled, with a few short hairs
and (?) glands; glands present on body of some
sporangia, more commonly sessile on stalks of
sporangia; spores not seen (sori old).
THELYPTERIDACEAE (Holttum)
459
Distr. Malesia: Papua New Guinea (S. New
Ireland), 2 collections.
Ecol. In forest at 650-700 m.
27. Sphaerostephanos solutus HOLTTUM, sp. nov.
— Aspidium hispidulum DECNE var. solutum
MIQUEL, Ann. Mus. Bot. Lugd.-Bat. 4 (1868)
160. — Aspidium hispidulum var. B METT. ibid. |
(1864) 234. — Type: SPANOGHE (?), Timor (L, n.
908, 333-406).
Base of frond lacking. Lamina of specimen
55cm long with 20 pairs pinnae. Largest pinnae:
19 x 2.1 cm, base truncate with acroscopic lobe of
many pinnae elongate, apex caudate-acuminate,
edges lobed to 2.5mm from costa, lobes falcate;
costules 4-4.5mm apart, at 60°; veins 12 pairs,
1-15 pairs anastomosing, one vein or a pair to
sinus-membrane; hairs on lower surface of costae
and rachis sparse, erect, 0.3mm long, similar but
shorter and more sparse on costules, glands also
present; between veins glands and sparse short
erect hairs; hairs on upper surface of costae
0.3mm long, similar hairs scattered on costules
and veins, very short erect hairs and glands be-
tween veins. Sori medial, lower ones not diver-
gent; indusia thin, with a few glands and short
hairs; sporangia with glands on body.
Distr. Malesia: Lesser Sunda Is. (Timor), only
known from the type.
Note. The type bears the name Aspidium
solutum n. sp. in BLUME’s hand, also Aspidium
hispidulum var. B METT. and is evidently the one
referred to by METTENIUS and MIQUEL, but they
give ZIPPELIUS as collector whereas the speci-
men here cited bears the collectors name
SPANOGHE. A. hispidulum DECNE (P) =Christ-
ella hispidula.
28. Sphaerostephanos hernaezii HOLTTUM, sp.
nov.
Caudex suberectus: stipes 10-15cm longus:
pinnae redactae c. 6-jugatae, superiores 5-6mm
longae; lamina 45cm longa; pinnae 9-jugatae,
inferiores usque 11.5 x 2.5 cm, basin versus sensim
angustatae, dimidio costam versus lobatae; cos-
tulae 7mm inter se distantes; venae 9-jugatae;
pagina utrinque pilis erectis brevibus glandulisque
praedita; sori mediales, indusia pilis brevibus
vestita. —Type: M. G. PRICE & B. E. HERNAEZ
75, Western Samar (K).
Caudex suberect. Stipe 10-15 cm long, glabrous
apart from short hairs in groove; basal scales
8x 1mm, copiously setiferous; base of stipe to
first large pinna 45 cm. Reduced pinnae c. 6 pairs,
uppermost 5-6 mm long, lowest 2mm. Lamina to
45 cm long; pinnae to 9 pairs, widely spaced, basal
ones longest; apical lamina almost pinna-like but
with widened base; aerophores not elongate.
Basal pinnae to 11.5X2.5cm, widest 1/3 from
apex and gradually narrowed to narrowly cuneate
base, in middle lobed half way to costa, apex
460
FLORA MALESIANA
fser. II, vol. 1°
short-acuminate; lobes slightly falcate, tapered
from base to an obtuse tip; costules to 7mm
apart; veins to 9 pairs, 15 pairs anastomosing, one
vein or a pair to short sinus-membrane; hairs on
lower surface of rachis and costae 0.2-0.3 mm
long, antrorsely curved, shorter hairs on costules
and veins, between veins minute erect hairs and
copious glands; hairs on upper surface of rachis
and costae to 0.5 mm, rest as lower surface. Upper
pinnae gradually less narrowed at base but up-
permost somewhat narrowed. Sori medial, basal
ones a little divergent and a little elongate along
veins; indusium with copious short hairs and some
glands; no glands nor hairs on sporangia.
Distr. Malesia: Philippines (Samar), only known
from the type.
Ecol. In primary forest on limestone and
limestone-derived soil.
Note. Young plants have fewer and short
pinnae but almost equally wide, with few lobes
and widely-spaced costules.
29. Sphaerostephanos
nov.
S. pennigero affinis, ab eo differt: fronde
minore; pinnis redactis 3—S-jugatis, omnibus par-
vis; pinnis normalibus 7-12-jugatis, fertilibus
usque 15 X2.1 cm; venis 23-jugatis anastomosan-
tibus; sporangiis nec setiferis nec glanduliferis. —
Type: MOSELEY s.n. Challenger Exp., Aru
Islands (K; BM).
Caudex short-creeping; stipe 20-35cm long,
basal scales 7 x 1 mm; base of stipe to first normal
pinna 35-65 cm; reduced pinnae 3-5 pairs, lowest
very small, uppermost c. 5mm long. Lamina to
50cm long; pinnae 7-12 pairs. Largest fertile pin-
nae 15 x2.1cm, shaped as in S. penniger but with
costules at a very wide angle to costa; veins 8
pairs, 2; pairs anastomosing, | pair to sinus-mem-
brane; pubescence as in S. penniger but some
short erect hairs in addition to glands between
veins on both surfaces. Sori medial, lower ones
divergent, those on basal veins from adjacent
costules sometimes confluent; indusia copiously
glandular; sporangia lacking glands or hairs dis-
tally, glands at ends of hairs on sporangium-stalks
conspicuous.
Distr. Malesia: S.E. Moluccas (Aru Is.).
Notes. In addition to the type are BUWALDA
4984 and 5169, both from Kobroor I., the former
much like the type. Number 5169 (from forest in
steep rocky limestone country) is sterile, with
longer fronds than the type and pinnae to 24x
2.7 cm, with 13-14 pairs of veins but only 23 pairs
anastomosing, 2 or 3 pairs passing to a long sinus-
membrane.
moseleyi HOLTTUM, sp.
30. Sphaerostephanos productus (KAULF.)
HOLTTUM, Kalikasan 4 (1975) 59.— Aspidium
productum KAULF. Enum. Fil. Chamisso (1824)
237; METT. Farngatt. IV (1858) 109. — Dryopteris
producta (KAULF.) C. CHR. Ind. Fil (1905) 286;
v.A.v.R. Handb. (1908) 230.— Cyclosorus
productus (KAULF.) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 248; COPEL. Fern FI. Philip.
(1960) 348, nomen tantum. — Thelypteris producta
(KAULF.) REED, Phytologia 17 (1968) 306.—
Type: CHAMISSO, Manila (LE).
Nephrodium philippinense BAK. Ann. Bot. 5
(1891) 327.— Dryopteris basilaris C. CHR. Ind.
Fil. (1905) 254, nom. nov. superfl.; CHRIST, Philip.
J. Sci. 2 (1907) Bot. 186; v.A.v.R. Handb. (1908)
820. — Dryopteris philippinensis (BAK.) COPEL.
Philip. J. Sci. 56 (1935) 100. — Lectotype (HOLT-
TUM 1975): CUMING 10, Luzon (K).
Dryopteris luzonica var. puberula CHRIST,
Philip. J. Sci. 2 (1907) Bot. 197.—Type: BOL-
STER 175, Luzon (P).
Dryopteris kotoensis HAYATA, Ic. Pl. Formosa 5
(1915) 279, f. 107.— Cyclosorus truncatus var.
kotoensis H. Iro, Bot. Mag. Tokyo 51 (1937)
729. — Thelypteris kotoensis (HAYATA) K. IWATS.
Acta Phytotax. Geobot. 21 (1964) 42; Mem. Coll.
Sci. Univ. Kyoto B, 31 (1965) 189. — S. kotoensis
(HAYATA) HOLTTUM in C. M. Kuo, FI. Taiwan 1
(1975) 436.— Type: KAWAKAMI & SASAKI S.n.
July 1912, Taiwan, Kotosho I. (TI; seen by
IWATSUKI).
Dryopteris pseudohirsuta’ ROSENST. Meded.
Rijksherb. n. 31 (1917) 7.— Thelypteris pseudo-
hirsuta (ROSENST.) REED, Phytologia 17 (1968)
306. — Type: CUMING 82, partim (L).
Cyclosorus serratus COPEL. Philip. J. Sci. 81
(1952) 36; Fern Fl. Philip. (1960) 365.— Type:
COPELAND P. P. E. 19, Luzon, Lamao River
(MICH; K).
Cyclosorus weberi COPEL. Philip. J. Sci. 81
(1952) 36, t. 25; Fern Fl. Philip. (1960) 366. —
Thelypteris weberi (COPEL.) REED, Phytologia 17
(1968) 324.—Type: WEBER 1174, Mindanao
(US; K).
Cyclosorus rigidus COPEL. Philip. J. Sci. 81
(1952) 27; Fern FI. Philip. (1960) 340, not C. rigi-
dus (RIDL.) COPEL. 1951. — Thelypteris ugoensis
REED, Phytologia 17 (1968) 322, nom. nov. —
Type: RAMOS BS 5744, Luzon, Mt Ugo (MICH).
Cyclosorus leucadenius COPEL. Philip. J. Sci.
81 (1952) 27; Fern Fl. Philip. (1960) 339.—
Thelypteris leucadenia (COPEL.) REED, Phy-
tologia 17 (1968) 287.—Type: ELMER 8999,
Luzon, Baguio (MICH; K, E, L, BO).
S. punctatus HOLTTUM, Kalikasan 4 (1975)
60.— Type: ELMER 16642, Luzon, Mt Bulusan
(US; K, L, G, BO, NSW).
Cyclosorus megaphyllus sensu COPEL. Fern FI.
Philip. (1960) 367.
Cyclosorus nitidulus sensu COPEL. Fern FI.
Philip. (1960) 364, p.p.
Caudex short-creeping, branched. Size of
fronds very variable, with pinnae from 101 to
30 2.7cm. Stipe short; basal scales 10-20mm
long, narrow. Reduced pinnae 6-12 pairs, not
1981]
deflexed, broadly ovate to bluntly triangular,
subentire, uppermost commonly I-1.5 cm long and
wide, lowest 0.5cm. Normal pinnae crenate in
smallest fronds, commonly lobed almost 1/3, in
largest fronds 1/2 towards costa; costules always
at about 45° to costa except in largest fronds;
veins 4-11 pairs, in almost all cases 12 pairs anas-
tomosing and | pair to sinus-membrane; hairs on
lower surface of costae near base very short
(rarely with some more than 0.5mm long), to
0.2mm or more long distally, very short and
sparse on costules and veins, many glands and
few hairs between veins; hairs on upper surface
of costae to 0.5 mm long, on costules and veins all
very short, glands between veins abundant. Hairs
on lower surface of rachis variable, 0.2-0.5 mm or
more long, on upper surface to Imm long. Sori
medial, lower ones not or little divergent; indusia
firm, bearing glands and sometimes a few short
hairs; sporangia bearing 1-2 setae or glands on
body and a spherical gland at end of hair on stalk;
spores with many small wings.
Distr. Malesia: Philippines (Luzon to Min-
danao); Taiwan (Orchid Island only).
Ecol. In forest at low and medium altitudes.
Notes. In 1975 I distinguished S. punctatus,
with large pinnae lobed fully half-way to costa and
costules at a rather broad angle, but these do not
appear to be sharply distinct; such specimens
occur also on Orchid Island (S.E. of Taiwan)
along with others indistinguishable from the type
of Nephrodium philippinense BAK. The specimen
originally described by KAULFUSS has pinnae
9x0.9cm; the type of Cyclosorus serratus
CoPEL. 10x0.8cm, the type of Nephrodium
philippinense 18 x 1.4cm; some other types come
between these extremes.
The name Nephrodium basilare PRESL (Epim.
Bot. 258) was published as a substitute for N.
caudiculatum J. SM. non PRESL (Hook. J. Bot.
3:411), but SMITH’s name was a nomen nudum,
and the first valid name based on the specimens
cited by SMITH was N. philippinense BAK. When
transferring this to Dryopteris, CHRISTENSEN
(1905) preferred the epithet basilaris because of
the existence of Phegopteris philippinensis METT.
METTENIUS evidently did not see _ the
CHAMISSO specimen (he cited CUMING LI,
which I cannot trace), but he gave a fairly good
description, mentioning the glands on the lower
surface which no other author noticed. HOOKER
and BAKER made no reference to Aspidium
productum KAULF. CHRISTENSEN, CHING,
COPELAND and REED transferred the species to
other genera without describing it. When describ-
ing Dryopteris producta, VAN ALDERWERELT
referred to METTENIUS, but did not copy MET-
TENIUS’s description; he gave a better descrip-
tion under D. basilaris. COPELAND misconstrued
the species in his Fern Flora of the Philippines,
placing it as a synonym of Nephrodium nitidulum
THELYPTERIDACEAE (Holttum)
461
PRESL, which belongs to the genus Pneumatop-
teris.
31. Sphaerostephanos penniger (HOOK.) HOLT-
TUM in Nayar & Kaur, Comp. to Bedd. (1974)
209.— Aspidium pennigerum sensu BL. Enum.
Pl. Jav. (1828) 153.— Nephrodium pennigerum
Hook. Spec. Fil. 4 (1862) 82, nom. nov.; RACIB.
Fl. Btzg 1 (1898) 190.—Type: BLUME, Java,
Boerangrang (L).
Aspidium megaphyllum METT. Ann. Mus. Bot.
Lugd.-Bat. 1 (1864) 233; MIQUEL, ibid. 4 (1869)
159. — Dryopteris megaphylla (METT.) C. CHR.
Ind. Fil. (1905) 277: v-A.v.R. Handb. (1908) 218:
BACKER & POSTH. Varenfl. Java (1939) 50.—
Cyclosorus megaphyllus (METT.) CHING, Bull.
Fan Mem. Inst. Biol. Bot. 8 (1938) 225; HOoLT-
TUM, Rev. Fl. Mal. 2 (1955) 268, f. 154.—
Thelypteris megaphylla (METT.) K. IWATS. Mem.
Coll. Sci. Univ. Kyoto B, 31 (1964) 34.— Type:
METTENIUS cited REINWARDT and ZIPPELIUS,
Java (L).
Aspidium multilineatum WALL., nom. nud. —
Nephrodium pennigerum var. multilineatum
CLARKE, Trans. Linn. Soc. Bot. | (1880) 532. —
Nephrodium multilineatum BEDD. Handb. Suppl.
(1892) 80 (not Aspidium multilineatum METT.
1858).— Dryopteris multilineata (BEDD.) O.
KiTrZE; ‘Rev; Gen! (PL? (1891) she wAy
Handb. (1908) 219.—Type: WALLICH 353,
Penang (K).
Aspidium abortivum sensu METT. Farngatt. IV
(1858) 110, quoad ZOLLINGER 3525 tantum.
Nephrodium abruptum sensu HOOK. Spec. Fil.
4 (1862) 78, p.p., t. 241B. — Fig. la-h.
KEY TO THE VARIETIES
1. Lower pinnae not over 4cm wide, lobed not
more than 1/3 towards costae.
2. Normal pinnae to 25 pairs, reduced pinnae
10-15 pairs : . a. var. penniger
. Normal pinnae 7 pairs: reduced pinnae 5 pairs
c. var. karoensis
lobed to 3mm
. b. var. excellens
1. Lower pinnae to 7cm wide,
from costa
a. var. penniger
Caudex short-creeping. Stipe 8-10cm long;
scales to 10x 1mm, closely setiferous; base of
stipe to first large plant 80cm. Reduced pinnae
10-15 pairs, upper ones to 32cm, spreading,
triangular with a not quite truncate symmetric
base, blunt apex and subentire edges, lowest on a
large plant 1x 1cm. Lamina to 100 cm long; pin-
nae 25 pairs; basal pinnae not or slightly narrowed
at base; apex not pinna-like; aerophores slightly
swollen. Largest pinnae commonly 20 x 2.8 cm (to
30*x4cm), lobed to a depth of 3-Smm, lobes
oblique, subdeltoid, broadly pointed; costules 5.5-
7mm apart, at 60°; veins 8-12 pairs, 3-33 pairs
anastomosing to form a zig-zag excurrent vein,
462
1-2 pairs to sinus-membrane; hairs on lower sur-
face of rachis and base of costae very short,
distally on costa and costules 0.2-0.3 mm, some-
what antrorse, between veins many glands; hairs
on upper surface of costae 0.5 mm long, minute on
costules, many glands between veins. Sori medial,
lower ones not divergent; indusia glandular with a
few short acicular hairs; sporangia with short
setae and glands on body, hair on stalk often with
a conspicuous terminal gland.
Distr. Peninsular Thailand, in Malesia: Malaya,
Sumatra, Borneo, Java, Lesser Sunda Is. (Bali,
Lombok), S.W. and N. Celebes.
Notes. On p. 78 of Species Filicum vol. 4
(which was printed off in fascicles of 16 pages)
HOOKER cited his specimens of this species un-
der Nephrodium abruptum and with them speci-
mens belonging to six other species; his plate
241B certainly represents a specimen of S. pen-
niger. In the next fascicle, on p. 82, he revised his
opinion and cited all his specimens of the present
species under N. pennigerum, based on Aspidium
pennigerum BL. stating correctly that they agreed
with a specimen so named by BLUME. HOOKER’s
description is a translation of BLUME’s; his name,
in Nephrodium, ranks as a new name.
BLUME’s name was based on a_misinter-
pretation of Polypodium pennigerum FORST.
which represents a New Zealand species very
different from the Java one, and in citing BLUME
HOOKER excluded the FORSTER name; in his
fifth volume HOOKER retained Polypodium pen-
nigerum FORST. for the New Zealand fern, which
is exindusiate.
HOOKER’s confusions on p. 78 misled BED-
DOME, who united an Indian species with the
Malesian S. penniger under the name Nephrodium
abruptum in Ferns of S. India, and then under N.
pennigerum (Handb. p. 277). In the Supplement to
his Handbook BEDDOME separated the two,
adopting WALLICH’s name multilineatum for the
Malesian species; this in turn led to confusions in
the name of another Indian species.
b. var. excellens (BL.) HOLTTUM, Stat. nov. —
Aspidium excellens BL. Enum. Pl. Jav. (1828)
160; C. CHR. Ind. Fil. (1905) 73.— Proferea
excellens (BL.) PRESL, Epim. Bot. (1851) 259;
HOLTTUM, Novit. Bot. Univ. Carol. Prag. 1968
(1969) 48.— Pleocnemia excellens (BL.) v.A.v.R.
Handb. (1908) 171.— Tectaria excellens C. CHR.
Ind. Fil. Suppl. III (1834) 179. — Type: BLUME,
Java (PRC; L, K).
Upper part of frond indistinguishable from the
typical form of S. penniger; lower pinnae greatly
enlarged, lowest 30 7cm, lobed to 3mm from
costa, costules 9mm apart, lobes falcate and
tapered to an acute tip, venation in lobes pleoc-
nemioid, in largest lobes aeroles of more than one
series along costule.
Distr. Malesia: Java, only one collection.
FLORA MALESIANA
[ser. II, vol. 1°
Note. In the Rijksherbarium at Leiden are two
sheets, one bearing the apical part of a frond, the
other a lower part of the same frond, but not the
base. The base of the frond appears to have been
divided between HOOKER (one pair of pinnae)
and PRESL who had the three lowest pairs of
large pinnae and two pairs of reduced pinnae.
c. var. karoensis HOLTTUM, var. nov.
A typo Sspeciei differt: fronde minore; pinnis
redactis S-jugatis; pinnis evolutis 7-jugatis, usque
13 x 2.7 cm, infimis basin versus valde angustatis ;
sporangiis setiferis, non glanduliferis. —Type:
MOLESWORTH ALLEN 2407, Sumatra, Prapat by
Lake Toba (K).
Distr. Malesia: N. Sumatra, only known from
the type.
32. Sphaerostephanos warburgii (KUHN &
CHRIST) HOLTTUM, Allertonia 1 (1977) 202.—
Aspidium warburgii KUHN & CHRIST in War-
burg, Monsunia 1 (1900) 81.— Dryopteris war-
burgi (KUHN & CHRIST)» 9G io 3@ar:
Ind. Fil. (1905) 98; v.A.v.R. Handb. (1908)
180. — Cyclosorus warburgii (KUHN & CHRIST)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941)
250; COPEL. Philip. J. Sci. 78 (1951) 441.—
Thelypteris warburgii (KUHN & CHRIST) B. C.
STONE, Micronesica 2 (1966) 3.— Type: WAR-
BURG, N.E. New Guinea, Sattelberg (B).
Dryopteris calcicola C. CHR. Bot. Jahrb. 66
(1933) 44.—Type: KJELLBERG 2611, S.E.
Celebes, Kosali-Porema (S-PA; BM).
Caudex short, erect. Stipe 1-3 cm long. Lamina
10-15 cm long; pinnae to 18 pairs, several lower
pairs gradually smaller, lowest 3-5 mm long. Lar-
gest pinnae 1.5—1.8cm long, 0.3-0.4 cm wide, dis-
tinctly stalked, base unequally cuneate, edges
entire or rarely crenulate, apex rounded or
obtusely pointed; costules 1.5mm apart, at 45°,
distal ones unbranched, rest forked once or twice,
branches of neighbouring ones anastomosing;
lower surface of costae bearing short hairs, some-
times with scattered long ones, glands throughout
between veins; upper surface glabrous apart
from costae and costules, not glandular. Hairs on
rachis to 1 mm long, both sides. Sori uniseriate on
each side of the costa; indusia with a few short
hairs; sporangia bearing glands, on specimens
from Tenimbar Islands and Celebes also setae.
Distr. Micronesia (Guam, Admiralty Is.), and
Malesia: New Guinea, S.E. Moluccas (Tenimber
Is.), and Central & S.E. Celebes.
Note. Specimens from Guam only differ from
those of New Guinea by their larger size (frond to
30cm long, pinnae to 3cm long), agreeing in
glandular sporangia. The type of D. calcicola,
from limestone, differs in fewer glands on lower
surface of pinnae, densely hairy indusia and
setose sporangia.
Ecol. On rocky stream banks.
1981]
33. Sphaerostephanos tandikatensis (v.A.v.R.)
HOLTTUM, comb. nov.— Dryopteris tandikaten-
sis v.A.v.R. Bull. Jard. Bot. Btzg II, 11 (1913)
11; Handb. Suppl. (1917) 178, 503.— Type:
MATTHEW 635, Sumatra, G. Tandikat (BO).
Caudex short-creeping. Stipe 4-5 cm long, short-
hairy; basal scales 3-4 = 1mm; base of stipe to
first large pinna 35-45cm. Reduced pinnae 12-
15 pairs, broadly ovate, uppermost 10 x 8 mm with
a small auricle on acroscopic base, lowest 3-4 mm
long. Lamina to 45cm long; pinnae 3-4 pairs,
lowest pinna usually a little reduced; apex of
lamina pinna-like, larger than pinnae on fronds of
young plants; aerophores slightly swollen. Pinnae
of type to 11 x 4cm, of another specimen to 17 x
6.5cm, widest at or above middle, base abruptly
broad-cuneate, of lower ones asymmetric; apex
abruptly short-cuspidate; edges entire, or crenate
distally; costules 3.5-4.5 mm apart, at 60° or rather
less; veins 12-14 pairs, almost all anastomosing,
combined excurrent veins almost straight, mostly
continuous; lower surface of costae and costules
densely covered with erect hairs 0.1mm long,
veins and surface less densely, glands present on
veins and surface between them: upper surface
covered with appressed hairs 0.3mm long, no
longer hairs present. Hairs on both sides of rachis
also short. Sori all supramedial, those on veins
from adjacent costules often coalescing; indusia
small, short-hairy; sporangia with many setae.
Distr. Malesia: South and Central Sumatra,
720-1400 m, several collections.
34. Sphaerostephanos carrii HOLTTUM, sp. nov.
Pinnae redactae 4-jugatae, superiores 1.5cm
longae; lamina usque 45cm longa; pinnae 6-7-
jugatae, omnes oppositae, 10 x 2.7 cm vel majores,
vix usque 1/3 costam versus lobatae; lamina inter
venas subtus glandulosa, supra glabra eglan-
dulosa; indusia tenuia, pilis brevibus glandulisque
praedita; sporangia setifera. Type: C. E. CARR
14202, Papua, Boridi, 1500 m (K; BM, L, LAE).
Caudex short-creeping or suberect. Stipe 12-
20cm long; basal scales 7-10 x 1.5-2 mm, thin,
setiferous; base of stipe to first large pinna 40—
50cm. Reduced pinnae 4-5 pairs, lowest 4mm
long, uppermost 1.5cm, deflexed and auricled.
Lamina 304S5cm_ long; pinnae 6-7 pairs,
opposite; lowest pinnae narrowed in basal 1/3 on
basiscopic side, truncate but not auricled on
acroscopic side; apical lamina almost pinna-like
but larger and more deeply lobed at base; texture
thin; aerophores not elongate. Suprabasal pinnae
10-17 x 2.7-3.5cm, base truncate, apex abruptly
short-acuminate, edges lobed rather less than 1/3
to costa, lobes slightly falcate with rounded tips;
costules 5-6 mm apart; veins 10-12 pairs, 23 pairs
anastomosing, I-l> pairs passing to sinus mem-
brane; lower surface of costae and costules bear-
ing short spreading hairs, distally on costa to
0.5mm long, short erect hairs and sparse glands
THELYPTERIDACEAE (Holttum)
463
between veins; hairs of upper surface of costae
mixed in length, longest 1 mm or more, scattered
similar long hairs on costules and veins, surface
between veins glabrous. Hairs on lower surface of
rachis rather sparse, 1-1.5mm long, on upper
surface copious, mostly 0.5 mm, longest I-1.5 mm.
Sori medial, basal ones not divergent; indusia
bearing short hairs and a few glands: sporangia
setiferous.
Distr. Malesia: Eastern New Guinea, 3 col-
lections.
35. Sphaerostephanos exindusiatus HOLTTUM, sp.
nov.
Pinnae redactae 15-jugatae, superiores vix
10mm longae; lamina 60cm longa; aerophora
2mm longa; pinnae maximae 12X1.5cm, 3/5
costam versus lobatae; venae 8-jugatae: costa
subtus pilis erectis 0.5 mm longis praedita, lamina
inter venas glandulosa; pagina superior omnino
pilis appressis vestita; sori exindusiati, sporangia
setifera.— Type: POSTHUMUS 3494, S.W.
Celebes, near Patapang, 1200m, on river bank
(BO).
Caudex short, thick, suberect. Stipe 15-20cm
long, densely scaly throughout when young: scales
to 10 x 2 mm, thin; base of stipe to first large pinna
50cm. Reduced pinnae at least 15 pairs, 1.5-2.cm
apart, lowest 4mm long, uppermost hardly 10 mm.
Lamina 60cm long; pinnae 25 pairs or more:
lower pinnae not or little narrowed at base; aero-
phores 2mm long. Largest pinnae 12 1.5cm,
base truncate and a little dilated both sides, apex
narrowly acuminate, edges lobed about 3/5 to
costa, lobes slightly falcate; costules 3-3.5mm
apart; veins to 8 pairs, 1 pair anastomosing,
next acroscopic vein to side of sinus-membrane;
lower surface of costae and costules bearing
rather sparse erect hairs 0.5mm long, surface
between veins copiously glandular; hairs on upper
surface of rachis and costae 1 mm long, scattered
similar hairs on costules and veins, whole surface
covered with slender appressed hairs. Sori in-
framedial, exindusiate; young sporangia bearing
several short setae.
Distr. Malesia: S.W. Celebes, only known from
the type.
36. Sphaerostephanos lucbanii HOLTTU™, sp. nov.
Pinnae redactae 2-jugatae; lamina 25 cm longa:
pinnae liberae 7-jugatae, tenues, maximae 6X
1.8 cm, 1/3-2/5 costam versus lobatae; venae 5—6-
jugatae: costae subtus pilis usque 1 mm longis
vestitae, pagina inter venas pilis erectis glan-
dulisque praedita; pagina superior pilis appressis
vestita: sori exindusiati; sporangia setifera. —
Type: M. G. Price 2720A, Luzon, Mt Bana-
haw, streamside at 1000 m (K).
Caudex slender, short-creeping. Stipe 5cm
long, densely short-hairy, scales 3 mm long; base
of stipe to first large pinna 11 cm. Reduced pinnae
464
FLORA MALESIANA
2 pairs on largest frond, lower ones 63mm,
upper one deflexed, 106mm, crenate, base
asymmetric. Lamina 25cm long, thin, apex nar-
rowly triangular, 13cm long, deeply lobed and
grading to upper pinnae, venation in its basal lobes
as in Haplodictyum; free pinnae 7 pairs, lowest
distinctly stalked, somewhat narrowed at base on
basiscopic side, aerophores swollen. Largest pin-
nae 6X 1.8cm, base truncate, apex shortly poin-
ted, edges lobed 1/3-2/5 to costa, lobes rounded,
entire; costules 4.5mm apart, at 60°; veins 5-6
pairs, | pair anastomosing with long excurrent
vein, next acroscopic vein to short sinus-mem-
brane; hairs on lower surface of costae rather
sparse, pale, slender, 0.5-1 mm long, on costules
same but few, between veins slender erect hairs
0.2 mm long and glands; hairs on upper surface of
costae 0.5 mm long, rest of upper surface covered
with slender appressed hairs 0.2-0.3mm_ long.
Hairs on both sides of rachis slender, to 1.5mm
long. Sori medial, exindusiate; sporangia seti-
ferous.
Distr. Malesia: Philippines (Luzon), only known
from the type.
Note. Under the same number and near the
same locality Mr PRICE also collected very much
smaller plants with lamina to 7 cm long, pinnae 2
pairs, | cm long, subentire, but fully fertile. These
small plants (separated as n. 2720B) agree in sori
and in pubescence with the larger ones above
described. They agree also closely in all respects
with Dryopteris diminuta COPEL. from Mindanao,
which I have transferred to Pronephrium. Mr
PRICE calls my attention to the plants described
by COPELAND as Dryopteris bakeri in Philip. J.
Sci. 38 (1929) 135; one of them had much larger
pinnae than the rest. It is thus possible that the
whole collection 2720 represents different stages
of plants on one species.
37. Sphaerostephanos major (COPEL.) HOLTTUM,
comb. nov.— Haplodictyum majus COPEL. in
Elmer, Leafl. Philip. Bot. 9 (1920) 3110; Fern FI.
Philip. (1960) 378; HoLtTTuM, Kalikasan 2 (1973)
61.— Type: RAMOS BS 13992, Luzon, Mountain
Prov., Apayao (MICH).
Cyclosorus dimorphus COPEL. Philip. J. Sci. 83
(1954) 99, pl. 5, non (BRAUSE) COPEL. 1960. —
Type: EDANO PNH = 17282, Luzon, Mountain
Prov., Mt Magnas (MICH).
? Dryopteris bakeri sensu COPEL. Philip. J. Sci.
38 (1929) 135.
Caudex short-creeping; fronds dimorphous.
Sterile frond. Stipe to 4cm long; base of stipe to
first large pinna 6-12cm; reduced pinnae 1-2
pairs, very small; apical lamina 12-20cm long,
deeply lobed, 2.5—4 cm wide at base, veins in lobes
mostly forked, the branches uniting to form a
series of costular areoles, base of lamina grading
into 2-4 pairs of pinnae closely placed; pinnae on
largest frond 3.5 x 2.cm, broadly pointed, lobed to
[ser. II, vol. 1°
about 1/3, with long spreading hairs on lower
surface of costae, short erect hairs and glands
between veins, upper surface covered with short
appressed hairs. Fertile frond. Stipe 13cm or
more, base of stipe to first large pinna 20-27 cm;
reduced pinnae as sterile but more widely spaced;
apical lamina to 12 x2.cm, veins mostly not for-
ked; pinnae to 2.5 1.4cm, more widely spaced
and less deeply lobed than sterile; sori on both
apical lamina and pinnae, basal ones sometimes a
little elongate; indusium small with many hairs;
sporangia with 2-3 long setae and sometimes also
a gland.
Distr. Malesia:
northern Luzon).
Philippines (mountains of
38. Sphaerostephanos polyotis (C. CHR.) HOLT-
TUM, comb. nov. — Dryopteris polyotis C. CHR.
Bot. Jahrb. 66 (1933) 46. — Thelypteris polyotis (C.
CHR.) REED, Phytologia 17 (1968) 305.— Type:
KJELLBERG 1718, S.W. Celebes, Todjamboe,
below 1000 m (BO; BM).
Caudex short, suberect. Stipe 5-10cm long;
base of stipe to first normal pinna 35-70cm.
Reduced pinnae 25-30 pairs, |cm apart on type
(to 2.5cm on other specimens), lowest 4-5 mm
long, uppermost 1.5-2.0cm long, deflexed with
basal acroscopic lobe 7 mm long, basiscopic lobe
shorter, edges incised, apex obtuse or rounded.
Lamina of type 75cm long; normal pinnae 25
pairs, basal ones a little narrowed towards their
bases, with enlarged basal lobes; aerophores not
elongate. Lower surface of rachis densely short-
hairy, hairs to 0.5mm long; upper surface with
antrorse pale hairs. Largest pinnae of type 7 x
10cm (of WALKER 12187 14x2.0cm); base
truncate; apex acuminate; edges lobed 1/2-2/3
towards costa; costules 3—3.5 mm apart, at 60° to
costa; veins of type to 5 pairs, of WALKER 12187
10-12 pairs, in both cases | pair anastomosing and
next acroscopic vein ending beside sinus-mem-
brane; lower surface of costae densely covered
with erect hairs of mixed length to 0.5 mm, more
sparse hairs and sometimes glands on costules
(not seen on type), surface between veins of
WALKER 12187 bearing short erect hairs and a
few glands; upper surface + closely covered with
short appressed hairs. Sori medial; indusia bearing
short hairs and sometimes a few glands; sporangia
bearing glands and sometimes setae.
Distr. Malesia: S.W. Central Celebes (Lati-
modjong Mts).
Notes. Additional collections are T.G. WALKER
12186-12188, 12270, 12273, 12282 (BM). Most
of these are larger than the type, but 12188
has normal pinnae of the same size as the type
with fewer reduced pinnae. WALKER’S specimens
also have some glands on the lower surface of
costules and between veins, but setae more
abundant than glands on sporangia. This species
and S. foliolosus are related to S. hispiduliformis
1981]
THELYPTERIDACEAE (Holttum)
465
of New Guinea, but the transition from normal to
reduced pinnae at the base of fronds of the latter
is not abrupt.
39. Sphaerostephanos grandescens HOLTTUM, sp.
nov.
Pinnae redactae c. 6-jugatae, omnes parvae;
lamina 45cm longa; pinnae 12-jugatae, 2/3 vel
ultra costam versus lobatae; costulae 5 mm inter
se distantes; pagina subtus inter venas sparsim
glandulifera, supra omnino pilis appressis vestita;
indusia pilis brevibus paucis praedita, sporangia
setifera.— Type: MERRILL 6094, Mindoro, Mt
Halcon (MICH).
Stipe 7-14.cm long, glabrescent; base of stipe to
first large pinna 45 cm or more. Reduced pinnae 6
pairs, 3rd from base 3—4mm long, upper ones
apparently not much larger. Lamina 45 cm long;
pinnae 12-13 pairs, basal pair widest above
middle, narrowed to base which is 1.2cm wide,
not auricled; aerophores elongate, less than | mm
long. Largest pinnae 12.5 x 2.4 cm, base broadly
cuneate, apex acuminate, edges lobed to 3.5mm
from costa (more than 2/3), lobes oblique, slightly
falcate, a little narrowed distally; costules 5mm
apart, at less than 60°; veins to 9 pairs, 1 pair
anastomosing, next acroscopic vein usually to side
of sinus-membrane; hairs on lower surface of
costae less than 0.5 mm long, antrorse distally, on
costules same but sparse, sparse glands on cos-
tules and on surface between veins; hairs on upper
surface of costae 0.6mm, scattered similar hairs
on costules and veins, whole surface covered with
slender hairs 0.3-0.4mm long. Hairs on lower
surface of rachis pale, curved, spreading, 0.6mm
long, on upper surface similar but appressed. Sori
medial, lower ones not divergent; indusia rather
firm, with a few short hairs only; sporangia seti-
ferous.
Distr.
Halcon).
Note. This specimen was named Dryopteris
bordenii (here placed as a variety of S. hetero-
carpus) by CHRIST, but has much wider pinnae
and setiferous sporangia and elongate aerophores;
it is near S. magnus, but smaller.
Malesia: Philippines (Mindoro: Mt
40. Sphaerostephanos magnus (COPEL.) HOLT-
TUM, comb. nov. — Cyclosorus magnus COPEL.
Philip. J. Sci. 81 (1952) 30; Fern FI. Philip. (1960)
346. — Thelypteris magnus (COPEL.) REED, Phy-
tologia 17 (1968) 291.—Type: MERRILL 6952,
Negros (MICH; B, P).
Stipe 10cm, glabrescent; scales broadly ovate,
thin, 2.5mm wide; base of stipe to first normal
pinna to 85cm. Reduced pinnae many pairs, each
consisting of an aerophore more than | mm long
and a minute green blade. Lamina to 100 cm long;
pinnae c. 30 pairs; basal pinnae apparently not
narrowed at their bases. Largest pinnae 21x
2.5cm, base subtruncate, apex acuminate, edges
lobed 3/5 to costa, lobes falcate, somewhat nar-
rowed distally; costules 5mm apart, at more than
60°; veins to 9 pairs, basal pair anastomosing, next
pair to sides of sinus-membrane; hairs on lower
surface of rachis sparse, to | mm long, on costae
and costules very short throughout with much
longer ones also distally; many glands throughout
lower surface; upper surface covered with slender
appressed hairs, with scattered longer ones on
costules and veins. Sori small, medial, lower ones
not divergent; indusia bearing glands; sporangia
setiferous, sometimes also a gland present.
Distr. Malesia: Philippines (Negros). Known
from two collections, the other being ELMER 9845
which consists in part of the present species, in
part of Pneumatopteris nitidulus (PRESL)
HOLTTUM.
41. Sphaerostephanos santomasii HOLTTUM,
Kalikasan 4 (1975) 62. — Type: M. G. PRICE 1034,
Luzon, Benguet Prov., Mt Santo Tomas, 2000 m
(K; PNH).
Caudex short, erect. Stipe 5-10 cm long. basal
scales narrow, 8mm long; base of stipe to first
large pinna 40-75cm. Reduced pinnae 34cm
apart, upper ones 3mm long, lowest very small.
Lamina to 55cm long; pinnae to 30 pairs; several
pairs lower pinnae narrowed towards their bases,
a basal short pair sometimes present; aerophores
to 1mm long. Largest pinnae to 16x1.5cm
(rarely to 2cm wide), base truncate, apex acu-
minate, edges lobed 3/5—2/3 to costa, lobes slightly
falcate; costules 34mm apart, at 60° or more;
veins 7-10 pairs, basal pair anastomosing, next
pair to edge or acroscopic one to sinus-membrane;
lower surface of costae of sterile pinnae bearing
pale spreading hairs 1mm long and more
numerous shorter antrorse hairs, longer hairs on
fertile pinnae less than | mm, similar hairs more
sparse on costules, surface between veins bearing
glands and a variable number of slender appressed
hairs; hairs on upper surface of costae 1 mm long,
scattered similar hairs on costules and veins,
whole surface covered with slender appressed
hairs 0.3-0.5 mm long. Sori medial, lower ones not
divergent; indusia small, bearing a few short hairs;
sporangia copiously setiferous.
Distr. Malesia: Philippines (Northern Luzon:
Mt Santo Tomas, Mt Nangaoto and Mt Data:
ALCASID PNH 1750), at 2000 m.
42. Sphaerostephanos trichochlamys HOLTTUM,
sp. nov.
Caudex brevis, erectus; stipes 8cm longus;
pinnae redactae usque 9-jugatae, superiores 3 mm
longae; lamina 45cm longa; pinnae maximae
13 X 1.7 cm, dimidio costam versus lobatae; rachis
costaeque subtus pilis multis usque 1.5 mm longis
vestita, pagina inter venas pilis erectis glandu-
lisque praedita; indusia pilosa; sporangia
setifera. — Type: HOLTTUM 4, Mt Kinabalu (K).
466
FLORA MALESIANA
[ser. II, volar
Caudex erect, short. Stipe 8cm long, dark red-
dish; base of stipe to first large pinna 50cm.
Reduced pinnae to 9 pairs, uppermost 3 mm long,
rest very small, with aerophores to 1 mm long.
Lamina to 45cm long; pinnae to 20 pairs, red
when young, lower ones narrowed towards their
bases. Hairs on rachis, both surfaces, to more
than 1 mm long, with shorter ones also on lower
surface. Largest pinnae 13X1.7cm (sterile to
2cm wide), base (except lower ones) truncate,
apex short-acuminate, edges lobed 2 way to costa;
costules 3-4 mm apart, falcate distally; veins 7-8
pairs, thick and prominent below in sterile fronds,
basal pair anastomosing, next acroscopic vein to
sinus-membrane; lower surface of costae densely
covered with spreading hairs 1-1.5mm long and
much shorter ones, similar but more sparse hairs
on costules and veins; surface between veins
bearing short erect hairs and glands; upper surface
of costae densely antrorse-hairy, sparse long hairs
on costules and veins, surface between veins with
a variable number of fine appressed hairs. Sori
medial, lower ones not divergent; indusia firm,
bearing many hairs 0.3mm _ long; sporangia
copiously setiferous.
Distr. Malesia: Borneo (Sabah: Mt Kinabalu,
many specimens; Sarawak: G. Mulu).
Ecol. In forest at c. 1500 m, not near streams;
young fronds red, not mucilaginous.
Notes. This is closely allied to S. hirsutus but
is smaller and much more densely hairy, with
copiously setose sporangia. A specimen from
2275 m on G. Mulu has few sori which have small
glabrous indusia.
43. Sphaerostephanos adenostegius (COPEL.)
HOLTTUM, comb. nov. — Dryopteris adenostegia
COPEL. Univ. Cal. Publ. Bot. 18 (1942) 220. —
Cyclosorus adenostegius (COPEL.) COPEL. Gen.
Fil. (1947) 142; Philip. J. Sci. 78 (1951) 442, pl.
6B.— Thelypteris adenostegia (COPEL.) REED,
Phytologia 17 (1968) 258.— Type: BRASS 10282,
near Lake Habbema, New Guinea, 2800m, in
forest (GH).
Pronephrium nothofageti HoLTTUM, Blumea
20 (1972) 118. — Type: T. G. WALKER 8788, New
Guinea, Finisterre Range, 2400 m, in Nothofagus
forest (BM).
Rhizome short-creeping, 3-4 mm diameter. Stipe
commonly to 30cm long, minutely hairy, basal
scales 5X 1mm. Lamina 20-30cm long; pinnae
8-10 pairs, lower ones with stalks 1-2 mm long,
lowest 1-4 pairs + reduced, lowest sometimes
1.3 cm long; texture firm; aerophores not elongate;
frond-apex deltoid, deeply lobed. Largest pinnae
5-8 cm long, 1.3-1.6cm wide, base subtruncate
and sometimes slightly auricled, edges lobed about
1/3 to costa, lobes slightly dentate at ends of
veins; costules 4-4)mm apart; veins to 5 pairs,
one pair anastomosing, next acroscopic vein to
side of sinus-membrane; hairs on lower surface of
rachis pale, + curved, to 0.5 mm long, on costae
0.2-0.3 mm long, slightly antrorse, sparse glands
usually present on costules and on surface be-
tween veins; hairs on upper surface of rachis as
lower, on costae short and sparse, scattered longer
hairs on costules and veins, slender appressed
hairs to 0.5mm long more or less abundant be-
tween veins. Sori medial; indusia small, with a
few short hairs and glands; sporangia bearing 1-3
slender setae; spores with a rather broad trans-
lucent wing and cross-wings.
Distr. Malesia: Eastern New Guinea, several
localities in forest at 2300-2900 m.
44. Sphaerostephanos aquatilis (COPEL.) HOLT-
TUM, comb. nov. — Dryopteris aquatilis COPEL.
Philip. J. Sci. 6 (1911) Bot. 75.— Thelypteris
aquatilis (COPEL.) REED, Phytologia 17 (1968)
260.— Type: C. KING 182, Papua (MICH, BO,
NSW, P).
Dryopteris caudiculata ROSENST.
Repert. 9 (1911) 426, non v.A.v.R.
Type: as above (BO).
Caudex short, erect. Stipe to 18cm long, pale
with dark base, short-hairy. Lamina to 45cm
long, apical section narrowly triangular and deeply
lobed; pinnae 20 pairs or more, well spaced, lower
3-4 pairs gradually smaller, lowest 0.8-1.3 cm
long; in smaller fronds only | pair pinnae some-
what reduced. Largest pinnae of type collection
8.5 x 1.0cm, of BRASS 6725 14 x 0.8 cm; base very
narrowly cuneate on basiscopic side, at c. 45° on
acroscopic; apex acuminate; edges very slightly
crenate; costules little over 2mm apart, at 45°;
veins of type 3 pairs, 2 pairs anastomosing; sinus-
membrane hardly evident; hairs on lower surface
of costae copious, antrorse, on costules sparse,
rest of lower surface glabrous, glands present on
costae and costules only of type (also between
veins on BRASS 6725); upper surface bearing hairs
on costae only, sparse distally. Sori medial to
supramedial; indusia rather large, glabrous or
sometimes bearing glands or a few short hairs;
sporangia bearing glands.
Distr. Malesia: S.E. Papua New Guinea; 2
collections (BRASS 6725, Fly River).
Ecol. At low altitude by streams in flood zone.
Note. Possibly not distinct from S. mutabilis.
Fedde
1908. —
45. Sphaerostephanos _hispidifolius (v.A.v.R.)
HOLTTUM, comb. nov.— Dryopteris hispidifolia
v.A.v.R. Bull. Jard. Bot. Btzg II, 20 (1915) 15;
Handb. Suppl. (1917) 186.— Thelypteris hispidi-
folia (v.A.v.R.) REED, Phytologia 17 (1968) 283. —
Lectotype (selected here): JAHERI 1124, Borneo
(BO).
Nephrodium hispidulum (non (DECNE) BAK.)
CHRIST, Ann. Jard. Bot. Btzg 20 (1906) 107. — Fig.
12]-m.
Caudex short-creeping, with a dense mass of
roots. Stipe 6-10 cm long, slender, reddish, short-
1981]
THELYPTERIDACEAE (Holttum)
467
hairy; basal scales 4mm long, narrow; base of
Stipe to first large pinna 10-17 cm. Reduced pinnae
usually 1-2 pairs, 1-2mm long, rarely lacking.
Lamina 20-30cm long, apex not pinna-like; pin-
nae 10-15 pairs, lower ones with stalks 1-2 mm
long; texture firm, aerophores not enlarged. Lar-
gest fertile pinnae 3.5-6cm long, 0.6-0.8 cm wide
(sterile to 7.5 x 0.9cm), base narrowly cuneate on
basiscopic side, more broadly on acroscopic, apex
acuminate, edges lobed 2 way to costa (or more
deeply in largest pinnae), lobes oblique; costules
3mm apart, at 45° to costa; veins 3-4 pairs, basal
acroscopic vein ending exactly at base of sinus,
basal basiscopic vein to edge very near base of
sinus; rather sparse stiff hairs to 1mm long on
lower surface of rachis, base of costae and mar-
gins of lobes, costa distally glabrous, some glands
present on lower surface between veins; upper
surface of pinnae glabrous apart from slender hairs
0.5 mm long on costae. Sori medial; indusia firm,
glabrous; sporangia bearing glands.
Distr. Malesia: Borneo; collections
several localities.
Ecol. On rocks and earth of stream banks, in
the flood zone.
Note. v.A.v.R. in the Supplement to his
Handbook describes the basal veins “‘uniting at or
a little below the sinus”, but in the specimens
examined by me they usually end separately,
though near together, their tips sometimes touch-
ing near base of pinnae.
from
46. Sphaerostephanos_ uniauriculatus (COPEL.)
HOLTTUM, comb. nov.—Dryopteris — uni-
auriculata COPEL. Philip. J. Sci. 9 (1914) Bot.
3.— Thelypteris uniauriculata (COPEL.) REED,
Phytologia 17 (1968) 322. — Type: C. KING 406,
Papua (MICH; BM, P).
Caudex short, suberect or creeping. Stipe 20-
27cm long, minutely hairy. Reduced pinnae one
on each side of rachis, not opposite, c. 5 mm long.
Lamina consisting of a large terminal leaflet and 2
pinnae (not opposite), sometimes a second pair
also; aerophores slightly swollen. Apical lamina c.
18 cm long, fertile 3.4-3.7 cm wide, sterile 4.2 cm;
base truncate or cordate, edges crenate; costules
5 mm apart along midrib; veins 8 pairs, almost all
anastomosing; texture firm. Sterile pinnae to 8 x
3.2cm, widest 1/3 from apex, base broadly roun-
ded to truncate, apex short-pointed, edges irre-
gularly sinuous; costules 4-4.5 mm apart; veins 6
pairs, 3 distal veins reaching the margin, rest
anastomosing; sinus-membrane hardly developed;
hairs on lower surface of costae, costules and
veins very short, somewhat antrorse, a few short
hairs and some glands on surface between veins;
upper surface glabrous apart from hairs on costae
and costules. Fertile pinnae to 6.5 X2.5cm; sori
exindusiate, distal ones medial, lower ones diver-
gent, those on basal veins of adjacent costules
sometimes confluent; sporangia bearing glands.
Distr. Malesia: Papua, known only from type
and KING 383 (P).
Note. KING 383 includes a young plant which
has a cordate-based apical lamina and one pair of
pinnae 6 x 8mm, also an older plant with 2 pairs
of pinnae and a single reduced pinna. COPELAND
stated that there is a group of hairs in place of an
indusium, but I did not see this.
47. Sphaerostephanos urdanetensis (COPEL.)
HOLTTUM, comb. nov. — Dryopteris urdanetensis
COPEL. in Elmer Leafl. Philip. Bot. 5 (1913)
1682. — Cyclosorus urdanetensis (COPEL.)
COPEL. Gen. Fil. (1947) 143; Fern Fl. Philip.
(1960) 355.— Thelypteris urdanetensis (COPEL.)
REED, Phytologia 17 (1968) 322. —Type: ELMER
13762, Mindanao, Mt Urdaneta (MICH).
Dryopteris matutumensis COPEL. Philip. J. Sci.
40 (1929) 299, pl. 3.— Cyclosorus matutumensis
(COPEL.) COPEL. Gen. Fil. (1947) 143; Fern FI.
Philip. (1960) 355.— Thelypteris matutumensis
(COPEL.) REED, Phytologia 17 (1968) 291.—
Type: COPELAND s.n. May 1917, Mindanao, Mt
Matutum, 2000 m (MICH; UC).
Caudex short, creeping; base of stipe to first
normal pinna 3-5 cm (sterile), 10-18 cm (fertile);
reduced pinnae 2-3 pairs, to 4-5 mm long. Lamina
excluding reduced pinnae to 20 cm long, consisting
of an apical lamina 12-15 cm long, to 3cm wide,
deeply lobed and grading to pinnae at its base,
with 3-5 pairs of free pinnae; basal pinnae nar-
rowed towards their bases on basiscopic side,
widest 1/3 from apex. Largest pinnae 1.5 x 0.8 cm
(type of D. matutumensis); apex broadly pointed
to rounded; edges crenate; costules 2mm apart;
veins 2-3 pairs, one pair anastomosing near base
of pinna, free elsewhere; lower surface of rachis
bearing thick curved pale brown hairs 0.5mm
long, hairs on costae shorter, yellow glands
present throughout; upper surface of rachis as
lower, of pinnae covered throughout with slender
appressed hairs. Sori on apical lamina medial or
supramedial, on pinnae near costae, one on each
basal vein; indusia small, dark, firm, glabrous or
with a few glands or short hairs; sporangia bearing
glands.
Distr. Malesia: Philippines (Mindanao).
Note. The type of D. matutumensis has larger
fronds than that of D. urdanetensis and has a few
hairs on indusia, but agrees in other characters.
48. Sphaerostephanos batjanensis (ROSENST.)
HOLTTUM, comb. nov. — Dryopteris batjanensis
ROSENST. Meded. Rijksherb. n. 31 (1917) 5.—
Thelypteris batjanensis (ROSENST.) REED, Phy-
tologia 17 (1968) 263.— Type: DE VRIESE &
TEYSMANN 589, Moluccas, Batjan (L).
Aspidium canescens forma gymnogrammoides
CHRIST, Ann. Jard. Bot. Btzg 15 (1898) J31.—
Type: SARASIN 137, N. Celebes, G. Masarang
(BAS).
468
Caudex short, erect. Base of stipe to first large
pinna of sterile fronds 5-12 cm, of fertile to 30 cm.
Reduced pinnae 1-2 pairs, very small, irregularly
spaced. Lamina to 30 cm long; pinnae 12-15 pairs,
basal 2-3 pairs deflexed and slightly reduced,
lowest narrowed towards base on basiscopic side.
Sterile pinnae to 3.5cm long, 1.2cm wide above
base which ts a little dilated and sometimes auri-
cled on acroscopic side; apex abruptly short-
pointed; edges lobed to a depth of 1.5mm; cos-
tules 3 mm apart; veins to 6 pairs, pale and prom-
inent on lower surface, 13 pairs anastomosing,
next pair to sinus-membrane; lower surface of
rachis, costae and costules covered with ap-
pressed pale hairs 0.3-0.4 mm long, copious glands
on surface between veins; upper surface of pinnae
covered throughout with slender appressed hairs
0.3 mm long, no glands; hairs on rachis more than
0.5mm long. Fertile pinnae a little smaller than
sterile; sori almost covering lower surface; in-
dusia firm, rather large, hairy; sporangia bearing
glands.
Distr. Malesia: N. Celebes & Moluccas (Bat-
jan, Ceram).
Note. The specimen of DE VRIESE & TEYS-
MANN 589 at Kew is labelled Ceram, not Batjan: it
agrees with the Leiden specimen.
49. Sphaerostephanos humilis HOLTTUM, sp. nov.
Pinnae 5-jugatae, usque 1.6 = 1.1 cm, inferiores
non decrescentes, subtus ubique glanduliferae,
supra pilis adpressis vestitae; sori mediales;: in-
dusia magna, glandulis pilisque brevibus praedita:
sporangia copiose’ glandulifera. —Type: W.
MEER 9826, Central Celebes, Mt Nokilalaki,
1500-2000 m (L).
Caudex short-creeping; stipe of sterile fronds
4-5 cm, of fertile 6-13 cm, short-hairy throughout,
basal scales thin, c. 3x 1.5mm. Sterile frond to c.
14cm long, consisting of an apical lamina 8.5 cm
long, 1.2cm wide, lobed to a depth of 2-2.5 mm,
and 5 pairs of pinnae, basal pinnae variably
somewhat reduced, no very small reduced pinnae
present; largest pinnae 1.6 1.1cm, subsessile,
widening from the base to widest part near apex,
irregularly shallowly lobed (most deeply towards
apex) with 3-4 lobes on each side: costules 3-
5mm apart; veins to 4 pairs, basal pair anas-
tomosing, next pair to margin; lower surface of
rachis bearing rather sparse pale hairs 0.4mm
long, costae covered sparsely with very short ap-
pressed hairs, glands copious throughout lower
surface which is not pustular; upper surface of
rachis hairy as lower, whole upper surface of
pinnae covered with appressed hairs 0.3 mm long.
Fertile fronds with apical lamina a little narrower
than sterile, edges of lobes dentate at ends of
veins; pinnae to 1.4 0.9cm:; sori medial; indusia
rather large, firm, with a few glands and usually a
few short hairs; sporangia copiously glandular;
spores spinulose.
FLORA MALESIANA
[ser. II, vol. 1°
Distr. Malesia: Central Celebes. Only known
from the type.
50. Sphaerostephanos lastreoides (PRESL) HOLT-
TUM, Kalikasan 4 (1975) 54.— Pronephrium las-
treoides PRESL, Epim. Bot. (1851) 259: Holttum,
Novit. Bot. Univ. Carol. Prag. 1968 (1969) 49.
Type: “Java (JUNGHUHN, comm. Vriese 1850)”
(PRC).
Cyclosorus duplosetosus COPEL. Philip. J. Sci.
81 (1952) 32; Fern Fl. Philip. (1960) 354. —
Thelypteris duplosetosus (COPEL.) REED, Phy-
tologia 17 (1968) 274.—Type: MERRILL 9530,
Palawan, Mt Capoas (MICH).
Caudex short, creeping. Stipe to 5cm (sterile),
to 15cm (fertile) covered with short pale hairs;
base of stipe to first large pinna 10-20cm.
Reduced pinnae 1-2 pairs, small, irregularly
spaced. Lamina to 15cm long; free pinnae c. 10
pairs, lower ones with short stalks and narrowed
at base on basiscopic side; apical lamina short,
acuminate; aerophores not elongate. Supramedial
pinnae to 2.3x0.9cm, base truncate, apex ab-
ruptly obtuse-pointed, lobed to about 3 way to
costa; costules 3 mm apart; veins 3-4 pairs, basal
pair anastomosing with excurrent vein to sinus,
next pair to edge; sparse pale spreading hairs
1 mm long and many much shorter ones on lower
surface of costae and costules, glands on surface
between veins; scattered long hairs on upper sur-
face of costae, costules and veins, whole surface
covered with slender appressed hairs 0.2-0.3 mm
long. Rachis also with spreading hairs 1 mm long
and shorter ones on both surfaces. Sori medial;
indusia with short hairs; sporangia bearing glands.
Distr. Malesia: Philippines (Palawan, Luzon,
Negros, Mindanao).
Notes. Part of CUMING 251 from Luzon, in
several herbaria, exactly matches the type of this
species; nothing like it is otherwise known from
Java. Probably the type was a CUMING specimen
misplaced by PRESL. M. G. PRICE 2734, from
Mindanao, Agusan del Sur Prov., is like the type
in all essentials but a little larger; its fertile pinnae
are the same size as sterile pinnae on the Kew
specimen of CUMING 251, which has smaller fer-
tile ones.
M. G. PRICE 690, from Mt Halcon, Mindoro,
differs from the above description only in larger
size: lamina to 35 cm long, pinnae 20 pairs, largest
7x1.7cm. It may represent a distinct local
variety.
A. LOHER s.n. March 1915, from Rizal Pro-
vince, Luzon, differs in having shallowly crenate
pinnae, and slender appressed hairs on lower sur-
face of costules.
51. Sphaerostephanos cataractorum (WAGN. &
GRETH.) HOLTTUM, comb. nov. — Cyclosorus
cataractorum WAGN. & GRETH. Un. Cal. Publ.
Bot. 23 (1948) 50, pl. 16. — Thelypteris catarac-
1981]
THELYPTERIDACEAE (Holttum)
469
torum (WAGN. & GRETH.) REED, Phytologia 17
(1968) 266. — Type: GRETHER & WAGNER 3971,
Admiralty Islands, Manus I. (UC; NSW).
Caudex short, erect. Stipe 10-20 cm long, pallid;
basal scales narrow, 5mm long. Lamina to 30cm
long; pinnae to 16 pairs; basal pinnae reduced,
sometimes one or both to a very narrow leaflet
less than |1cm long; apex of frond sometimes
almost pinna-like; texture thin; aerophores not
elongate. Largest pinnae to 10x 0.9cm, widest at
middle, base very narrowly cuneate, apex acu-
minate, edges lobed } way to costa, lobes very
oblique, slightly falcate, acute; costules 44.5 mm
apart, at 45°; veins 5-6 pairs, basal pair anas-
tomosing with excurrent vein to sinus, next
acroscopic vein to side of short sinus-membrane;
lower surface of pinnae quite glabrous with
resinous pale glands between veins; upper surface
short-hairy on costae, otherwise glabrous. Rachis
glabrous on lower surface or with very short
hairs; hairs on upper surface thick, brown, curved,
0.3-0.4mm long. Sori medial; indusium firm,
glabrous; sporangia bearing glands.
Distr. Admiralty Islands, New _ Britain,
Bougainville, in Malesia: Eastern New Guinea.
Ecol. By streams in flood-zone at altitudes to
250 m.
Note. This is very similar to S. hispidifolius of
Borneo, but has veins always anastomosing and
almost glabrous pinnae. The resinous glands on
lower surface are usually flattened when dry, not
spherical as normally in this genus.
52. Sphaerostephanos menadensis HOLTTUM, sp.
nov.
Pinnae redactae 2-jugatae, minutae; lamina
20 cm longa, pars apicalis angusta 10cm longa;
pinnae 6-jugatae, infimae basin versus valde
angustatae, mediales 5.5 x 1.6 cm, dimidio costam
versus lobatae; costae subtus pilis minutis ves-
titae; pagina inter venas subtus glandulifera,
supra eglandulosa; indusia glabra, sporangia
setifera.— Type: KOORDERS' 17133, Celebes,
Menado, 1800-2000 m (L).
Base of stipe to first large pinna 25 cm. Reduced
pinnae 2 pairs, very small and widely spaced.
Terminal lamina 10cm long, narrowly deltoid;
free pinnae 6 pairs, lowest deflexed and gradually
much narrowed towards their bases, widest above
middle; aerophores not elongate. Middle pinnae to
5.5 x 1.6 cm, base truncate, apex abruptly caudate,
edges lobed 3 way to costa, lobes falcate; costules
3.5mm apart, at more than 60°; veins 6 pairs
(fertile) 7-8 pairs (sterile), 1 pair anastomosing,
next pair of the acroscopic one only to sinus-
membrane; long coarse hairs on lower surface of
rachis, very short ones on costae and costules,
surfaces between veins bearing glands only; hairs
on upper surface of costules and veins all very
short, neither hairs nor glands between veins. Sori
medial; indusia glabrous; sporangia bearing glands.
Distr. Malesia:
from the type.
N.E. Celebes; only known
53. Sphaerostephanos subcordatus HOLTTUM, sp.
nov.
Pinnae redactae unijugatae, parvae; frondes
dimorphae; pinnae usque 8-jugatae, steriles 3
1.2cm, fertiles 1.7X0.9cm, usque 1/3 costam
versus lobatae; costae subtus brevi-pilosae,
pagina inter venas glabra glandulis paucis
praedita; pagina superior inter venas glabra,
eglandulosa. indusia parva, glabra; sporangia
glandulosa — Type: M. G. PRICE & B. F. HER-
NAEZ 61, Western Samar (K).
Caudex short, creeping, 3mm diameter when
dry. Stipe of sterile fronds 5-7 cm, of fertile 15-
18 cm to first large pinna below which are 2 much-
reduced pinnae, not opposite. Lamina 15 cm long,
apex broadly deltoid and deeply lobed, free pinnae
to 8 pairs, basal ones slightly reduced, defiexed,
with stalks 1mm long and asymmetric base;
aerophores not elongate. Largest sterile pinnae
3 x 1.2cm, in some cases 1.5 cm wide near apex;
base truncate to subcordate; apex abruptly blunt-
pointed; edges lobed to a depth of 2mm, more
deeply near apex if it is widened, lobes rounded;
costules 4mm apart, at less than 60°; veins 4-5
pairs, basal pair anastomosing, next acroscopic
vein to sinus-membrane; hairs on lower surface of
costae 0.2mm _ long, antrorsely curved, more
sparse and shorter on costules, surface between
veins glabrous with a few glands; hairs on upper
surface of costae sparse, 0.4mm long, similar or
longer ones scattered on costules and veins; sur-
face between veins glabrous. Hairs on lower sur-
face of rachis 0.3-0.4 mm long, thick and curved,
on upper surface rather sparse, | mm long. Sori
medial, lower ones sometimes a little elongate;
indusia small, glabrous; sporangia with glands.
Distr. Malesia: Philippines (W. Samar), only
known from the type.
Ecol. On limestone-derived soils, in forest.
54. Sphaerostephanos novoguineensis (BRAUSE)
HOLTTUM, comb. nov.—Dryopteris novo-
guineensis BRAUSE, Bot. Jahrb. 49 (1912) 21;
v.A.v.R. Handb. Suppl. (1917) 159.— Lastrea
novoguineensis (BRAUSE) COPEL. Gen. Fil.
(1947) 139; Philip. J. Sci. 78 (1951) 426. — Thelyp-
teris novoguineensis (BRAUSE) REED, Phytologia
17 (1968) 297. — Type: SCHLECHTER 17719, N.E.
New Guinea, Kani Mts, 1000 m (B; P).
Dryopteris glaucescens BRAUSE, Bot. Jahrb.
56 (1920) 85; CoPEL. Philip. J. Sci. 78 (1951)
426. — Thelypteris glaucescens (BRAUSE) CHING,
Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 251. —
Type: LEDERMANN 13034, N.E. New Guinea,
Sepik Dist. 1400 m (B).
Caudex short, creeping. Stipe 10-15 cm long;
basal scales thin, 10 x 2.5 mm; base of stipe to first
large pinna 20-45 cm. Reduced pinnae 3-8 pairs,
470
FLORA MALESIANA
[ser. II, vol. 1°
not more than 3mm long. Lamina to 80cm long;
pinnae to 25 pairs, lower ones not narrowed at
base; aerophores to more than I mm long. Pinnae
somewhat dimorphous, sterile to 10 x 1.7 cm, fer-
tile to 7.5 x 1.2 cm; base truncate, basal acroscopic
lobe to 4mm longer than next; apex short-acu-
minate; edges lobed to less than | mm from costa,
lobes falcate; costules 3 mm apart (sterile), 2.5 mm
(fertile); veins 10 pairs, basal acroscopic vein
passing to side of sinus-membrane, basiscopic
vein to edge above it; lower surface of costae
bearing many short hairs and less abundant long
ones to Imm (long hairs sometimes lacking),
similar hairs on costules, glands sometimes few
present on surface between veins; upper surface
with hairs | mm long scattered on costae, costules
and veins, surface between veins + closely
covered with fine appressed hairs. Sori medial to
supramedial; indusia large, firm, with short hairs
and sometimes glands; sporangia bearing glands;
spores finely spinulose.
Distr. Malesia: Eastern New Guinea, several
collections, in forest at 850-1400 m.
Note. The type of D. glaucescens differs from
that of D. novoguineensis in the absence of long
hairs on lower surface of rachis and costae and
the presence of glands on indusia.
55. Sphaerostephanos hispiduliformis (C. CHR.)
HOLTTUM, comb. nov.— Dryopteris hispiduli-
formis C. CHR. Ind. Fil. Suppl. III (1934) 88, new
name for Dryopteris hispidula BRAUSE, Bot. Jahrb.
56 (1920) 102, non (Sw.) O. KTZE 1891. — Thelyp-
teris hispiduliformis (C. CHR.) REED, Phytologia 17
(1968) 283.—Type: LEDERMANN 11758, N.E.
New Guinea, Sepik Distr. 2070 m, in forest (B).
KEY TO THE VARIETIES
1. Caudex to 150. cm tall; lower surface of pinnae
lacking glands between veins
a. var. hispiduliformis
1. Caudex short; lower surface of pinnae between
veins copiously glandular.
. Fronds to 90cm long; pinnae to 14 1.5 cm;
hairs on lower surface of costae 0.1 mm long
b. var. vinkii
2. Fronds to 40 cm long; pinnae to 5 x | cm; hairs
on lower surface of costae 1 mm long
c. var. brassii
nN
a. var. hispiduliformis
Caudex erect, to 150cm tall; stipe 10-15cm
long, basal scales broad, thin; base of stipe to
pinnae of maximum size 60cm or more, whole
frond to 150cm long; reduced pinnae 1.5-2.0cm
apart, broadly triangular, c. 12 pairs gradually
increasing upwards from very small basal ones to
lcm, then 7-12 pairs grading to pinnae of maxi-
mum size; aerophores more than | mm_ long.
Largest pinnae 11cm long, to 1.7cm wide at
dilated base, rather evenly tapered to apex, lobed
c. 2/5 towards costa; costules to 3.5mm apart, at
more than 60° to costa; veins 7-9 pairs, 12 pairs
anastomosing, 1-15 pairs ending beside sinus-
membrane; lower surface of costae with spreading
pale hairs 1mm long and shorter ones, hairs on
costules more sparse, a few short hairs between
veins, glands confined to costules and veins,
sometimes lacking; hairs on upper surface of cos-
tae more than | mm long, scattered similar hairs
on costules and veins, fine appressed hairs all over
surface. Sori medial; indusia bearing hairs and
glands, sporangia with neither; spores covered
with very small wings.
Distr. Malesia: Eastern
mountain forest at c. 2000 m.
New Guinea, in
b. var. vinkii HOLTTUM, var. nov.
A typo speciei differt: caudice breve; pinnis 1/3
costam versus lobatis; costis subtus pilis minutis
erectis vestitis; pagina inferiore inter venas
copiose glandulifera.— Type of variety: VINK
17605, N.E. New Guinea, W. Sepik Distr., in low
secondary growth on limestone (L).
Distr. Malesia: Papua New Guinea (Sepik).
c. var. brassii HOLTTUM, var. nov. — Dryopteris
strigosissima COPEL. Univ. Cal. Publ. Bot. 18
(1942) 221. — Cyclosorus strigosissimus (COPEL.)
COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78
(1951) 456, pl. 36.— Thelypteris strigosissima
(COPEL.) REED, Phytologia 17 (1968) 316.—
Type: BRASS 11436, New Guinea, Bele River,
2200 m, on limestone cliff in shade (MICH).
Differs from var. vinkii: fronds much smaller;
reduced pinnae contiguous, lower ones wider than
long; largest pinnae 5 x | cm, crenate to a depth of
1mm; long hairs present on lower surface of
costae.
Distr. Malesia: Papua New Guinea.
Notes. The greater number of glands on the
lower surface may be due to the limestone habitat
of var. vinkii and var. brassii (plants of S.
heterocarpus on limestone are densely glandular),
also their short caudex. But there is no evidence
that LEDERMANN’s type did not grow on limes-
tone. When more information is available it may
be better to regard var. vinkii and var. brassii as
representing a distinct species, for which
COPELAND’s epithet strigosissimus would be
available.
56. Sphaerostephanos hastatopinnatus (BRAUSE)
HOLTTUM, comb. nov. — Dryopteris hastatopin-
nata BRAUSE, Bot. Jahrb. 56 (1920) 112.—
Thelypteris hastatopinnata (BRAUSE) REED,
Phytologia 17 (1968) 281. — Type: LEDERMANN
8237, N.E. New Guinea, Sepik Distr., 200m on
rocks in stream-bed in forest (B; K).
Plants varying much in size according to habi-
tat. Caudex erect, short. Stipe 3-4cm on type, to
1981]
THELYPTERIDACEAE (Holttum)
471
15cm on large plant. Lamina of type 45 cm long,
of large plant 130cm; pinnae 28 pairs on type, 35
pairs on large plant; 10-20 pairs lower pinnae in all
cases gradually smaller, lowest 5S-10mm_ long;
apex of frond almost pinna-like; aerophores dis-
tinctly swollen, to almost 1mm long in large
fronds. Reduced pinnae broadly and symmettric-
ally triangular, spreading, subentire. Largest pin-
nae of type 10X1.3cm, of a large plant 20x
3.0cm; base truncate, dilated and more or less
auricled both sides (upper ones auricled on
acroscopic side only), apex short-acuminate,
edges sinuous or slightly crenate; costules of small
plants 2.5-3 mm apart, of a large one to 4.5 mm, at
more than 60°; veins 3-7 pairs, almost all anas-
tomosing; sinus-membrane hardly developed;
lower surface of pinnae quite glabrous or with a
few hairs on costae, glands sometimes present on
and between veins; hairs on upper surface of
costae slender, brown, to 0.5mm _ long, none
elsewhere, a few glands rarely present. Lower
surface of rachis glabrous or with sparse ap-
pressed hairs, upper surface bearing copious red-
brown hairs | mm long. Sori medial, lower ones +
elongate and sometimes confluent; no indusia;
sporangia bearing glands.
Distr. Malesia: Central Celebes, Moluccas
(Ceram) and widely distributed in New Guinea.
Ecol. On river banks in forest at low altitudes,
up to 650m. The type was probably stunted
owing to its rocky habitat and perhaps exposed
position.
57. Sphaerostephanos_ latebrosus (KUNZE ex
METT.) HOLTTUM in Nayar & Kaur, Comp. to
Bedd. (1974) 209. — Aspidium latebrosum KUNZE
ex METT. Farngatt. IV (1858) 104; BAK. Syn. Fil.
(1867) 294; MIQUEL, Ann. Mus. Bot. Lugd.-Bat. 4
(1869) 159.— Dryopteris latebrosa (METT.) C.
Cure. Ind. Fil. (1905) 274; v.A.v.R. Handb. (1908)
221. — Thelypteris latebrosa (METT.) REED, Phy-
tologia 17 (1968) 287. — Type: ZOLLINGER 354 pt,
Java (L, n. 908, 333-517 only).
Nephrodium glaucostipes BEDD. Handb. Suppl.
(1892) 80.— Dryopteris glaucostipes (BEDD.) C.
Cure. Ind. Fil. (1905) 268; Gard. Bull. Str. Settl. 4
(1929) 389; v.A.v.R. Handb. (1908) 226. — Cyclo-
sorus heterocarpus var. glaucostipes (BEDD.)
HOLTTuM, Rev. Fl. Mal. 2 (1955) 271. — Thelyp-
teris heterocarpus var. glaucostipes (BEDD.)
REED, Phytologia 17 (1968) 282. — Type: KING’s
Collector (KUNSTLER) 2046, Perak, Larut (K;
CAL, SING).
Dryopteris todayensis CHRIST, Philip. J. Sci. 2
(1907) Bot. 193; v.A.v.R. Handb. Suppl. (1917)
184. — Cyclosorus todayensis (CHRIST) CHING,
Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 249;
COPEL. Fern FI. Philip. (1960) 344. — Thelypteris
todayensis (CHRIST) REED, Phytologia 17 (1968)
319. — Sphaerostephanos todayensis (CHRIST)
HOLTTUM, Kalikasan 4 (1975) 58.—Type:
COPELAND 1463, Mindanao, Mt Apo 1220 m (P;
WiSiB):
Dryopteris angustipes COPEL. Philip. J. Sci. 7
(1912) Bot. 60; v.A.v.R. Handb. Suppl. (1917)
184.— Thelypteris angustipes (COPEL.) REED,
Phytologia 17 (1968) 260.— Type: BROOKS 110,
Sarawak (MICH; BM).
Cyclosorus sagittifolioides COPEL. Philip. J.
Sci. 81 (1952) 29, 30, pl. 21; Fern FI. Philip. (1960)
344. — Thelypteris sagittifolioides (COPEL.)
REED, Phytologia 17 (1968) 311. —Type: EDANO
BS 24832, Samar, Catubig River (US).
Cyclosorus halconensis COPEL. Philip. J. Sci.
81 (1952) 29; Fern Fl. Philip. (1960) 345.—
Thelypteris halconensis (COPEL.) REED, Phy-
tologia 17 (1968) 305. — Type: EDANO PNH 3849,
Mindoro, Mt Halcon (MICH; L).
Cyclosorus polypterus COPEL. Philip. J. Sci. 84
(1955) 161.—Thelypteris polyptera (COPEL.)
REED, Phytologia 17 (1968) 305. — Type: EDANO
7852 (PNH 21373), Negros, Canlaon Volcano
650 m (MICH).
Caudex erect, not or little branched at base.
Stipe 6-12cm long; base of stipe to first large
pinna 45-70 cm. Reduced pinnae 10-15 pairs, 2.5—
4 cm apart, lowest commonly 5 mm long, gradually
increasing upwards to 2.5-3.cm, larger ones nar-
rowly triangular, somewhat deflexed, with acros-
copic auricle, edges incised, apex broad-pointed;
transition to large pinnae subabrupt; aerophores
elongate, in some cases | mm or rarely more (type
of C. sagittifolioides). Lamina above reduced
pinnae to 60cm or more long; basal large pinnae
not narrowed at base. Rachis lower surface
commonly quite glabrous, in some specimens with
sparse long hairs or some very short ones; hairs
on upper surface to | mm long, usually appressed
(spreading in some Philippine specimens). Largest
pinnae 12 X 1.6 to 20 x 2.0 cm; base truncate, usu-
ally a little dilated both sides; apex acuminate;
edges lobed about half-way to costa or a little
more in large pinnae; costules 3.5-4.5 mm apart;
veins 7-8 pairs, basal pair anastomosing, one or
both of next pair to sinus-membrane; lower sur-
face of costae usually glabrous at base with very
short antrorse hairs distally, in some cases very
short hairs and a few somewhat longer
throughout; costules as costae but with shorter
hairs (if any); rest of lower surface usually quite
glabrous with many glands; upper surface with
long hairs on costae, similar hairs scattered on
costules and veins, rest of surface bearing a vari-
able number of fine appressed short hairs. Sori
medial, lower ones not divergent; indusia glabrous
or with a few short hairs, usually also glands;
sporangia bearing glands, rarely a seta; spores
with many very small wings.
Distr. Malesia: Western Malesia, Philippines.
Ecol. In forest, low altitudes to 800 m.
Notes. In Malaya, Sumatra and Java all lower
surfaces are almost glabrous, but in nearly all
472
cases there are short hairs on distal parts of
costae; aerophores are little elongate. In Sabah
(Mt Kinabalu) and the Philippines some hairs are
nearly always present and aerophores are more
developed. In the type of Cyclosorus sagit-
tifolioides some aerophores are 2mm long, and
short erect hairs are present on lower surface
between veins.
Some Philippine specimens seem rather inter-
mediate between this species and S. hirsutus; they
may be hybrids. Hybrids with S. heterocarpus in
Sabah are also possible. The solitary erect caudex
of S. latebrosus, large upper reduced pinnae and
broad base of lowest normal pinnae seem dis-
tinctive.
58. Sphaerostephanos porphyricola) (COPEL.)
HOLTTUM, Kalikasan 4 (1975) 59.— Dryopteris
porphyricola COPEL. Philip. J. Sci. 7 (1912) Bot.
60; v.A.v.R. Handb. Suppl. (1917) 186. — Thelyp-
teris porphyricola (COPEL.) CHING, Bull. Fan
Mem. Inst. Biol. Bot. 6 (1936) 287, nomen
tantum. — Cyclosorus porphyricola (COPEL.)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941)
248; HOLTTUM, Rev. FI. Malaya 2 (1955) 271. —
Type: BROOKS 112, Sarawak, Bau (MICH; BM).
Dryopteris angustipes sensu. C. CHR. Gard.
Bull. Str. Settl. 4 (1929) 389.
Caudex erect, short. Stipe 10cm long; base of
stipe to first large pinna 35 cm. Reduced pinnae to
12 pairs, 1.5cm apart, uppermost I-1.5 cm long,
deflexed, narrowly triangular with auricle on
acroscopic base, edges incised, lowest ones 3-
5mm long. Lamina 55 cm long; pinnae to 20 pairs
or more, lowest ones narrowed a little at base on
basiscopic side only; aerophores elongate, to less
than 1mm long; young fronds covered with
mucilage. Lower surface of rachis almost
glabrous or with very short hairs, also many
glands; upper surface copiously hairy. Largest
pinnae 14x 2.5 cm (sterile to 15 x 3 cm), base sub-
truncate, apex abruptly short-acuminate; edges
lobed 2/5 to costa, lobes distinctly falcate with
forward-pointing tips; costules 3-4.5mm_ apart;
veins 7-9 pairs, | pair anastomosing, next pair to
sinus-membrane; lower surface of costae, costules
and veins covered with fine short closely ap-
pressed hairs, also glands; surface between veins
bearing copious glands and a variable number of
short fine appressed hairs; hairs on upper surface
of costae antrorse, pale, similar hairs sometimes
scattered on costules and veins, surface between
veins typically covered closely with slender ap-
pressed hairs 0.5mm long, fewer such hairs on
specimens from Malaya. Sori medial, lower ones
not divergent but sometimes a little elongate; in-
dusia bearing short hairs and glands; sporangia
with glands.
Distr. Malesia: Malaya and Borneo (Sarawak;
Sulu Arch.: Tawi-Tawi).
Ecol. In forest at low altitudes.
FLORA MALESIANA
[ser. II, vol. 1°
Note. This is closely allied to S. latebrosus;
more field observations are needed.
59. Sphaerostephanos caulescens HOLTTUM, sp.
nov. — Dryopteris porphyricola (non COPEL.) C.
Cur. Gard. Bull. Str. Settl. 7 (1934) 244.
Caudex erectus; stipes Scm longus; pinnae
redactae usque 8-jugatae, omnes minutae; aero-
phora |mm longa; pinnae majores 13.5 Xx 1.5-
2.0cm, dimidio costam versus lobatae; rachis
subtus fere glabra; pagina inter venas subtus
glandulifera, supra pilis appressis vestita; indusia
fere glabra; sporangia glandulifera. — Type: G. P.
LEWIS 284, Sarawak, G. Mulu (K).
Caudex erect, to at least 20cm tall. Stipe 5 cm
long, base covered with large thin scales; base of
stipe to first large pinna 40 cm. Reduced pinnae to
8 pairs, when young covered with mucilage, all
very small, with aerophores |mm long. Lower
surface of rachis often bearing narrow scales,
otherwise almost glabrous, upper surface with
hairs Imm long closely appressed, with some
longer and spreading. Lamina to 75 cm long; pin-
nae to more than 30 pairs, not opposite; basal
pinnae (at least sterile ones) narrowed at their
bases. Largest fertile pinnae 13.5 X 2.0 cm (sterile
sometimes larger, to 20*3cm); base truncate;
apex acuminate (sometimes with cauda 1.5 cm
long); edges lobed about 3 way to costa; costules
c. 3.5mm apart, at 60°; veins c. 7 pairs, 1 pair
anastomosing, next pair to sides of sinus-mem-
brane; lower surface of costae almost glabrous at
base, with appressed hairs to 0.2 mm long distally,
costules similar, surfaces between veins bearing
copious glands; hairs on upper surface of costae
to 1mm long, similar hairs scattered on costules
and veins, short appressed hairs on surface be-
tween veins. Sori medial, lower ones not diver-
gent; indusia glabrous or with few very short
hairs; sporangia bearing glands.
Distr. Malesia: Borneo (N. Sarawak; Sabah),
N. Celebes, and Moluccas (Batjan).
Ecol. In forest near streams at 500-1500 m.
Note. This is near S. magnus, but pinnae of S.
caulescens (except largest sterile omnes) are
generally smaller and less deeply lobed, with
different pubescence and glandular sporangia.
60. Sphaerostephanos reconditus HOLTTUM, sp.
nov.
Pinnae redactae c. 6-jugatae, parvae; pinnae
normales usque 7.2 X 1.5 cm, dimidio costam ver-
sus lobatae; venae 5-6-jugatae, infimae solum
anastomosantes; costulae subtus pilis adpressis
vestitae; pagina inferior inter venas pilis adpressis
glandulisque praedita; sori mediales; indusia
dense pilosa; sporangia copiose glandulifera. —
Type: B. S. PARRIS 6872, Sarawak, Gunong Mulu
(CGE).
Caudex short, erect or suberect; stipe Scm
long; base of stipe to first normal pinna 30cm;
1981]
THELYPTERIDACEAE (Holttum)
473
reduced pinnae all very small, 3-4cm apart.
Lamina excluding reduced pinnae 30cm long;
pinnae c. 12 pairs; basal pinnae much narrowed in
basal third, above base 2cm wide; aerophores
when dried barely 0.5mm long, on living plant
“vellow’’. Lower surface of rachis densely hairy,
hairs thick, curved, brown, | mm long with shorter
pale ones (brown hairs more numerous on basal
part); upper surface with thicker brown hairs and
shorter pale ones. Suprabasal pinnae to 7.2
1.5cm; base truncate and slightly auricled; apex
short-acuminate; edges lobed not quite half-way
to costa; costules 3.5-4.5 mm apart, at c. 60°; veins
5-6 pairs, hardly prominent, | pair anastomosing,
next acroscopic vein or pair passing to sinus-
membrane; lower surface of costae bearing pale
antrorse hairs which are appressed distally,
0.5mm long, also glands, costules covered with
slender appressed hairs, between veins some ap-
pressed hairs and rather sparse glands; upper sur-
face of costae bearing pale hairs, no long hairs on
costules or veins, between veins sparse appressed
hairs. Sori medial or a little inframedial, lower
ones not divergent; indusia densely hairy, some-
times with a gland; sporangia bearing many
glands.
Distr. Malesia: Borneo (Sarawak: Mt Mulu),
only known from type.
Ecol. In forest clearing in Hidden Valley at
450 m; very young fronds covered with mucilage.
61. Sphaerostephanos cyrtocaulos (v.A.v.R.)
HOLTTUM, comb. nov. — Dryopteris cyrtocaulos
v.A.v.R. Bull. Jard. Bot. Btzg III, 5 (1922) 201. —
Thelypteris cyrtocaulos (v.A.v.R.) REED, Phy-
tologia 17 (1968) 270. — Lectotype (here selected):
BUNNEMEIJJER 9922, Sumatra, G. Kerinci, 2100 m
(BO; L).
Caudex erect. Stipe 10-15cm long; base of
stipe to first large pinna 40-75 cm. Reduced pinnae
8 pairs or more, 2-3 mm long; aerophores to 2mm
long. Lamina to 75 cm long; pinnae more than 20
pairs, well spaced; lowest pinnae not narrowed at
base, nor auricled. Hairs on both sides of rachis
short. Largest pinnae of lectotype 12x2.5cm
(possibly larger on sterile fronds); base truncate;
apex acuminate with cauda to 1.5 cm; edges lobed
to 2.5-3 mm from costa (about 3/4), lobes slightly
falcate; costules 44.5 mm apart at more than 60°;
veins 8-11 pairs, basal pair anastomosing, next
pair both to edge above base of sinus; hairs on
lower surface of costae and costules 0.2 mm long,
appressed, a few also on veins, glands present on
and between veins; hairs on upper surface of
costae 1 mm long, antrorse, similar hairs scattered
on costules and veins, pale appressed hairs over
whole surface. Sori rather supramedial; indusia
firm, rather large, with short hairs; sporangia
bearing glands; spores densely short-spinulose.
Distr. Malesia: Sumatra (on and near G.
Kerinci), 1800-2300 m, in forest.
Note. VAN ALDERWERELT cited three col-
lections by BUNNEMEIJER without indicating a
type; the largest one is here selected. One other
collection is known from a neighbouring locality.
62. Sphaerostephanos baramensis (C. CHR.)
HOLTTUM, comb. nov. — Dryopteris baramensis
C. CHR. Gard. Bull. Str. Settl. 7 (1934) 246. —
Thelypteris baramensis (C. CHR.) CHING, Bull.
Fan Mem. Inst. Biol. Bot. 6 (1936) 287. — Type:
MJOBERG 5 partim, Sarawak, Baram Valley, 900—
1200 m (BM). — Fig. 13f-g.
Caudex short, erect. Stipe 5-10 cm long, basal
scales thin, to 2mm wide at base; base of stipe to
first large pinna 40-60 cm. Reduced pinnae to 10
pairs, 4-6 cm apart, c. 2mm long, with aerophores
c. 1mm long. Lamina to at least 70cm long;
pinnae to 20 pairs, lower ones often opposite,
upper ones usually not; 3-4 pairs lower pinnae
much narrowed towards their bases; middle pin-
nae often distinctly stalked; aerophores 1-2 mm
long. Lower surface of rachis glabrous; hairs on
upper surface less than | mm long, brown, ap-
pressed. Largest pinnae commonly 15 x2.5cm,
largest on type 24 4cm; apex of wider pinnae
rather abruptly caudate-acuminate; edges lobed to
3mm from costa; lobes falcate; costules com-
monly 4mm apart, on large fronds to 7 mm; veins
10-12 pairs, prominent on lower surface, tips of
basal veins touching sides of short sinus-mem-
brane or sometimes uniting just below the sinus to
form a short excurrent vein; lower surface of
costae and costules covered with antrorsely cur-
ved hairs 0.2-0.3mm long, those on costules
closely appressed; surface between veins bearing
a variable number of small glands and short ap-
pressed hairs; upper surface of costae bearing
pale antrorse hairs less than 1 mm long, hairs on
costules much shorter, surface between veins of
type covered with short appressed hairs, of other
specimens glabrous or with appressed hairs near
costa only. Sori a little supramedial; indusia rather
large, firm, glabrous or with short hairs; sporangia
sometimes with a small gland, not setiferous.
Distr. Malesia: Borneo (Northern Sarawak, E.
Kalimantan and Sabah), in forest at 900-1500 m.
63. Sphaerostephanos batulantensis HOLTTUM, sp.
nov.
Caudex brevis, repens; stipes 10cm longus;
pinnae redactae usque 10-jugatae, superiores 4—
5mm longae; pinnae maximae 111.5 cm, 2/3
costam versus lobatae; costulae subtus pilis bre-
vibus appressis vestitae, pagina inter venas glan-
dulifera; pagina superior omnino pilis appressis
vestita; sporangia glandulifera.— Type: Kos-
TERMANS 18850, W. Sumbawa, Mt Batulante,
900 m (L; BO, K).
Caudex short, prostrate, c. 5 mm diameter. Stipe
10 cm long; base of stipe to first large pinna 40 cm
or more. Reduced pinnae to 10 pairs, upper ones
474
4-5mm long. Lamina 45cm long; lowest pinnae
narrowed to a base 5 mm wide; aerophores hardly
1mm long. Lower surface of rachis densely
covered with pale erect hairs 0.3mm long, hairs
on upper surface to |mm long, pale. Largest
pinnae 11 x 1.5 cm; base subtruncate, not auricled;
apex acuminate; edges lobed 2/3 to costa or a little
more deeply, lobes falcate; costules 3-4 mm apart,
at more than 60°; veins 8-9 pairs, basal pair anas-
tomosing, next acroscopic vein sometimes to
sinus-membrane; lower surface of costae hairy as
rachis, hairs on distal part longer and antrorse;
lower surface of costules covered with short ap-
pressed hairs; surface between veins bearing
glands and a few short erect hairs; upper surface
covered entirely with fine appressed hairs, no long
hairs on costules or veins. Sori a little inframedial,
lower ones not divergent; indusia thin with many
hairs 0.3 mm long; sporangia bearing glands.
Distr. Malesia: Lesser Sunda Is. (W. Sum-
bawa: Mt Batulante, 900 m); only known from the
type.
64. Sphaerostephanos subalpinus (v.A.v.R.)
HOLTTUM, comb. nov.— Dryopteris subalpina
v.A.v.R. Bull. Jard. Bot. Btzg III, 5 (1922) 200. —
Thelypteris subalpina REED, Phytologia 17 (1968)
317. Type: BEGUIN 1496, Ternate, N. Formad-
jahi, 1200 m (BO; L).
Polypodium acutum ROXB. Calc. J. Nat. Hist. 4
(1844) 492, non BURM. f.; MORTON, Contr. U.S.
Nat. Herb. 38 (1974) 335.—Type: ‘““Amboina”
(probably C. SMITH, Ceram).
Aspidium hispidulum DECNE var. ternatense
Miq. Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 160. —
Type: TEYSMANN, Ternate (L).
Caudex short, creeping. Stipe 10cm long; base
of stipe to first large pinna 30-35cm. Reduced
pinnae 1.5-2.0cm apart, c. 10 pairs, upper ones
1.5cm long, deflexed, narrowly triangular, deeply
lobed, with basiscopic auricle to 10mm _ long,
lowest 5mm long. Lamina 60-70 cm long; pinnae
to 30 pairs, basal ones not narrowed at base;
aerophores c. | mm long. Lower surface of rachis
covered with very short hairs and sparse longer
ones to | mm long; hairs on upper surface pale or
brownish, to 1mm long. Largest pinnae to 15 x
2.0cm, base subtruncate and a little dilated both
sides; apex caudate-acuminate; edges lobed to
2mm from costa, lobes falcate, narrowed above
their bases; costules to 4mm apart; veins 8-10
pairs, basal pair anastomosing near base of pinna,
near apex both touching sides of short sinus-
membrane; hairs on lower surface of costae near
base 0.2-0.3 mm long, suberect, distally to 0.7 mm,
hairs on costules. similar, glands present
throughout lower surface; hairs on upper surface
of costa mostly less than 1 mm long, with scat-
tered longer ones, similar longer hairs scattered on
costules and veins; a few appressed hairs some-
times present between veins. Sori a little
FLORA MALESIANA
[ser. II, vol. 1°
supramedial, basal ones not divergent; indusia
fairly large, with a few short hairs and sometimes
glands; sporangia with 1-2 glands.
Distr. Malesia: Moluccas (Ternate,
collections; Halmahera; Ceram?).
several
65. Sphaerostephanos lobatus (COPEL.) HOLT-
TUM, Kalikasan 4 (1975) 64. — Cyclosorus lobatus
(COPEL.) COPEL. Philip. J. Sci. 81 (1952) 33; Fern
Fl. Philip. (1960) 356.— Thelypteris lobata
(COPEL.) REED, Phytologia 17 (1968) 289.—
Type: COPELAND 1966, Luzon, Baguio (MICH).
Dryopteris canescens var. lobata CHRIST,
Philip. J. Sci. 2 (1907) Bot. 198, quoad
COPELAND 1866 (1966).
Cyclosorus bordenii sensu COPEL. Fern FI.
Philip. (1960) 345, p.p.
Caudex short, creeping. Stipe 5-8 cm long; base
of stipe to first large pinna 40 cm or more (shorter
on sterile fronds). Reduced pinnae 4-8 pairs, all
very small with thick aerophores Imm _ long.
Lamina commonly to 35cm long; pinnae to c. 16
pairs; basal pinnae much narrowed towards their
bases. Rachis bearing thick curved dark red hairs
to 1 mm long on both surfaces. Largest suprabasal
pinnae acuminate, to 15 x 1.7 cm (basal pinnae to
2cm wide) on larger plants, on type abruptly
short-pointed, c. 8 x 1.7 cm, lobed 2/3-3/4 to costa;
costules to 4mm apart; veins 7-9 pairs, basal pair
anastomosing, next pair both to edge; hairs on
lower surface of costae at base 0.1-0.2 mm, dis-
tally to almost 1 mm long, not appressed; hairs on
costules sparse; copious glands on surface be-
tween veins; hairs on upper surface of costae less
than | mm long, scattered similar hairs on costules
and veins at least distally on pinna-lobes, surface
between veins bearing a variable number of fine
appressed hairs. Sori inframedial; indusia small,
glabrous or with a few glands; sporangia bearing
glands.
Distr. Malesia: Philippines (Luzon, many col-
lections), in forest at 700-1000 m.
Note. On Mt Makiling a large form of this
species, above described, is common. On moun-
tains further north are plants (including the type)
with shorter abruptly pointed pinnae but closely
similar in pubescence and in reduced pinnae to the
plants on Mt Makiling. COPELAND placed most
of the Mt Makiling plants in Cyclosorus bordenii
(here included in Sphaerostephanos heterocarpus).
He failed to notice the reduced basal pinnae of the
type of C. lobatus. The plants here named S.
sessilipinna and S. urdanetensis are very similar
to the type of S. lobatus except in size of pinnae.
66. Sphaerostephanos ellipticus (ROSENST.)
HOLTTUM, Kalikasan 4 (1975) 66.— Dryopteris
elliptica ROSENST. Meded. Rijksherb. Leiden n.
31 (1917) 6.—Cyclosorus ellipticus (ROSENST.)
COPEL. Gen. Fil. (1947) 142; Fern Fl. Philip.
(1960) 351, nomen tantum. — Thelypteris elliptica
1981]
THELYPTERIDACEAE (Holttum)
475
(ROSENST.) REED, Phytologia 17 (1968) 274.—
Type: ELMER 13976 partim, Mindanao, Mt
Urdaneta (L, n. 914, 130-95; FI, G, K, U).
KEY, LOPTHE VARIETIES
1. Basal veins free on distal part of pinnae; lower
surface of rachis hairy . a. var. ellipticus
1. Basal veins not free in distal part of pinnae; lower
surface of rachis almost glabrous
b. var. glabrior
a. var. ellipticus
Caudex short, probably erect. Stipe 4-7cm
long, covered with very short hairs; base of stipe
to first large pinna 35cm. Reduced pinnae to 6
pairs, 34cm apart, uppermost 3-5 mm _ long,
lowest 2mm or less. Lamina 45 cm long; pinnae
to 18 pairs; basal pinnae slightly narrowed at base;
aerophores hardly 1mm long. Lower surface of
rachis covered with very short hairs and rather
sparse ones to imm long; upper surface with
uniform hairs 1mm. Largest pinnae 8.5 X 1.6cm,
base truncate, apex short-acuminate, edges lobed
2/3 towards costa; costules 3.5 mm apart, at more
than 60°; veins 8-9 pairs, basal pair near base of
pinnae anastomosing with short excurrent vein to
sinus, near apex of pinnae just meeting at the
sinus-membrane; lower surface of costae hairy as
rachis but longest hairs less than 1 mm, hairs on
costules sparse, between veins very short erect
hairs and glands; hairs on upper surface of costae
1mm long, similar or shorter hairs scattered on
c>stules and veins, short appressed hairs copious
to rather sparse between veins. Sori inframedial;
indusium firm with short stiff hairs; sporangia
sometimes with glands.
Distr. Malesia: Philippines (Mindanao), only
known from type.
Note. ELMER’s n. 13976 included specimens
of another species S. norrisii; COPELAND’s des-
cription of 1960 applies to the latter.
b. var. glabrior HOLTTUM, var. nov.
A speciei typica differt: pinnis 3/5 costam ver-
sus lobatis; venis semper anastomosantibus;
rachidi subtus fere glabra; costis subtus pilis 0.1-
0.2mm longis praeditis. —Type: M. G. PRICE
2558, Negros Oriental, Amlan, near river, 600—
800 m (K).
Distr. Malesia: Philippines (Negros Oriental),
only known from the type.
67. Sphaerostephanos foxworthyi HOLTTUM, sp.
nov.
S. elliptico (Rosenst.) Holttum affinis, differt:
pinnis redactis omnibus minutis; pinnis evolutis
inferioribus basin versus valde angustatis. —
Type: FOXwWworRTHY BS 2441, Luzon, Tayabas
Prov. Mt Banajao (K).
Caudex short, creeping. Stipe
12cm _ long,
glabrescent; scales broad and thin; base of stipe to
first large pinna 36cm. Reduced pinnae 5 pairs,
each consisting of an aerophore without detect-
able blade. Lamina 50 cm long; pinnae c. 20 pairs;
lowest 2 pairs much narrowed towards their
bases; aerophores thick, to 1mm long. Rachis
bearing pale, erect hairs 0.2mm long on lower
surface, not dense. Largest pinnae 13x 1.8cm;
base subtruncate; apex acuminate, sometimes
with cauda I-2cm long; edges lobed 2/3-3/4 to
costa; costules 4-4.5mm apart, at 60°; veins 9
pairs, | pair anastomosing, next pair usually both
to margin; lower surface of costae near base
bearing erect hairs 0.1-0.2 mm long, distally hairs
to 0.4 mm, antrorsely curved, not appressed, hairs
on costules few, between veins many glands, not
hairs; upper surface of costae bearing antrorse
hairs less than I mm long, rest of surface covered
with fine appressed hairs 0.2-0.3mm long. Sori
medial, lower ones divergent; indusia thin with
glands and a few short hairs; sporangia bearing
glands.
Distr. Malesia: Philippines (Luzon), only known
from the type.
68. Sphaerostephanos indrapurae HOLTTUM, sp.
nov.
Pinnae redactae usque 10-jugatae, superiores
3mm _ longae, aerophoris elongatis praeditae;
pinnae sSteriles usque 13X2.3cm, fertiles 9x
1.7 cm, c. 2/5 costam versus lobatae; rachis subtus
glabra, costae costulaeque pilis appressis vestitae;
pagina inter venas glandulifera; pagina superior
inter venas glabra; indusia glabra, sporangia
glandulifera. — Type: C. G. MATTHEW 655,
Sumatra, G. Kerinci 1500 m (BO; K).
Stipe 10cm long; base of stipe to first large
pinna 50cm. Reduced pinnae 10 pairs, blade of
largest 3mm long, with elongate aerophore.
Lamina 50-60 cm long; pinnae c. 20 pairs, several
lower pairs much narrowed towards their bases,
only upper ones truncate to full width at base.
Rachis glabrous on lower surface; hairs on upper
surface slender, less than 1mm long. Largest
pinnae 13X2.3cm (sterile), 9x 1.7cm (fertile),
lobed c. 2/5 towards costa, lobes with blunt for-
ward-pointing tips; apex abruptly — short-
acuminate; costules 5mm (sterile) 4mm (fertile)
apart; veins 6-8 pairs, basal pair anastomosing,
one or both of next pair ending at sinus-mem-
brane; lower surface of costae and costules
covered with fine closely appressed hairs 0.2-
0.4mm long, surface between veins bearing
glands; hairs on upper surface of costae 0.5mm
long, sparse very short ones on costules, no
others. Sori medial, lower ones a little divergent;
indusia glabrous; sporangia bearing glands.
Distr. Malesia: Central Sumatra (Mt Kerinci),
1500 m, only known from the type.
(ROSENST.)
69. Sphaerostephanos batacorum
476
FLORA MALESIANA
[ser. II, vol. 1°
HOLTTUM, comb. nov.— Dryopteris batacorum
ROSENST. Fedde Rep. 13 (1914) 217, excl. var.
winkleri; v.A.v.R. Handb. Suppl. (1917) 185.—
Thelypteris batacorum (ROSENST.) REED, Phy-
tologia 17 (1968) 263.— Type: J. WINKLER 158,
Sumatra, Batak Lands (S-PA).
Dryopteris stipellata var. obtusata v.A.v.R.
Bull. Jard. Bot. Btzg II, 2 (1920) 151. — Syntypes:
BROOKS 277, Sumatra, Lebong Tandai; 249,
Tambang Sawah (BO; BM). — Fig. 13h-k.
Caudex short; erect (?). Stipe 5-10cm long;
base of stipe to first large pinna 30-50cm.
Reduced pinnae to 20 pairs, 1.5-2cm apart,
deflexed, largest 1.5-2cm long, 8mm wide, with
acroscopic auricle 8mm long, edges shallowly
lobed, apex broadly obtuse. Lamina to 60cm
long; pinnae to 30 pairs, basal ones sometimes
narrowed a little at base on basiscopic side;
aerophores thick, to 2mm long. Rachis beneath
bearing short pale appressed or + spreading hairs
0.1-0.2 mm long, hairs on upper surface 1mm
long. Largest pinnae 9X 1.6cm; base truncate;
apex abruptly short-pointed; edges lobed 1/3-2/5
to costa; costules 3—3.5 mm apart; veins to 8 pairs,
basal pair anastomosing, one or both of next pair
to sinus-membrane; hairs on lower surface of
costa and costules pale, closely appressed, 0.2-
0.3 mm long; surface between veins bearing glands
which in some specimens are sparse (most abun-
dant on reduced pinnae); hairs on upper surface
of costae less than 1 mm long, sometimes no long
hairs on costules, rest of upper surface glabrous.
Sori medial; indusia rather large, glabrous, some-
times with a few glands; sporangia sometimes
with a gland.
Distr. Malesia: Sumatra, several collections,
at 850-1000 m.
Note. The type is a small specimen; the above
description is prepared partly from larger speci-
mens collected on G. Singgalang and G. Kerinci
which agree in shape and pubescence of frond and
of reduced pinnae.
70. Sphaerostephanos angustibasis HOLTTUM, sp.
nov.
Caudex erectus, gracilis; pinnae redactae 6-
jugatae, superiores 5mm _ longae, auriculatae;
aerophora elongata; pinnae usque 9.5 X2.1 cm,
inferiores basi angustatae, 2/3 costam versus
lobatae; costae subtus patenti-pilosae; pagina in-
ter venas subtus glandulosa; indusia sporangiaque
glandulifera. — Type: ALSTON 16724, Tidore, G.
Kiematuba (BM).
Caudex slender, to 20 cm tall. Stipe 15 cm long,
short-hairy; base of stipe to first large pinna
60cm. Reduced pinnae c. 6 pairs, subopposite,
upper ones 5mm long, deflexed, narrow, with
basal acroscopic auricle 4 mm long. Lamina 38 cm
long; pinnae to 15 pairs, almost all opposite, lower
3-4 pairs narrowed towards their bases, basal pair
5-7 mm wide at base; aerophores 1mm _ long.
Lower surface of rachis bearing curved brown
hairs and very short ones, brown hairs on upper
surface more uniform. Largest pinnae 9 x 2.1 cm,
base truncate, apex short-acuminate, edges lobed
up to 2/3 towards costa, lobes slightly falcate;
costules 5mm apart, at more than 60°; veins 7-8
pairs, all at a wide angle to costule, pale and
prominent both sides, basal pair anastomosing,
next pair both to edge; most hairs on lower sur-
face of costae minute, with some longer, erect, on
costules all short, glands present throughout lower
surface; upper surface of costae covered with
brown hairs, similar hairs scattered on costules
and veins, surface between veins glabrous. Sori
medial; indusia small, with glands and a few hairs;
sporangia with many glands, sometimes also with
a seta; spores with many small wings.
Distr. Malesia: Moluccas (Tidore), only known
from the type.
71. Sphaerostephanos nudisorus HOLTTUM, sp.
nov.
Pinnae redactae 12-jugatae, superiores 3mm
longae; aerophora 2mm longa; pinnae normales
usque 15X1.8cm, profunde lobatae; rachis sub-
tus glabra, costae subtus pilis minutis erectis
praeditae; pagina inter venas subtus glandulifera,
supra glabra; sori mediales, exindusiati; sporan-
gia nec glandulis nec setis praedita. — Type: T. G.
WALKER 12269, Central Celebes (BM).
Caudex not known. Stipe 5cm long, glabrous;
base of stipe to first large pinna 50cm. Reduced
pinnae c. 12 pairs, upper ones 3mm long, with
aerophores 2 mm long. Lamina 95 cm long; pinnae
c. 40 pairs, lowest not narrowed at base; one pair
of intermediate length between normal and
reduced pinnae. Lower surface of rachis bearing
glands and small scales only, upper surface
covered with hairs 1mm long. Largest pinnae
15x 1.8cm; base truncate with basal acroscopic
lobe 2 mm longer than next; apex acuminate with
cauda 1.5-2cm long; edges lobed to 2mm from
costa, lobes oblong, hardly falcate, with rounded
ends; costules 4mm apart, at more than 60°; veins
8 pairs, basal pair anastomosing near base of
pinna, near apex just meeting at the sinus; hairs
on lower surface of costae very short, erect,
glands abundant all over lower surface; hairs on
upper surface of costae 1 mm long, similar hairs
scattered on costules and veins, rest of upper
surface glabrous. Sori medial, exindusiate; neither
glands nor setae on sporangia; spores minutely
spinulose.
Distr. Malesia: Central Celebes, only known
from the type.
72. Sphaerostephanos paripinnatus (COPEL.)
HOLTTUM, comb. nov.— Dryopteris paripinnata
COPEL. Univ. Cal. Publ. Bot. 18 (1942) 220.—
Cyclosorus paripinnatus (COPEL.) COPEL. Gen.
Fil. (1947) 143; Philip. J. Sci. 78 (1951) 448, pl.
1981]
THELYPTERIDACEAE (Holttum)
477
28.— Thelypteris paripinnata (COPEL.) REED,
Phytologia 17 (1968) 302. — Type: BRASS 12435,
New Guinea, Idenburg River (MICH; BO, L).
Caudex probably erect. Stipe 10cm _ long,
glabrous, basal scales narrow; base of stipe to first
large pinna 60cm. Reduced pinnae to 10 pairs,
each consisting of an aerophore 2 mm long and a
minute blade; transition to normal pinnae abrupt.
Lamina to 75cm long; apical section long and
deeply lobed with gradual transition to pinnae;
pinnae 20 pairs, their ends much upcurved, sub-
coriaceous. Largest pinnae 21X2.5cm; base
truncate to subcordate, apical 3-S cm gradually
attenuate and crenate, middle part lobed more
than >» way to costa, lobes somewhat narrowed,
not falcate; costules to 6 mm apart, almost at right
angles to costa; veins 11-12 pairs, at a very wide
angle to costule, basal pair anastomosing, next
pair both to sides of short sinus-membrane; lower
surface of rachis, costae and costules with
spreading hairs 0.4mm long, shorter erect hairs
and glands between veins; hairs on upper surface
of costa 1mm long, similar hairs scattered on
costules and veins, no hairs nor glands between
veins. Sori medial, basal ones divergent; no in-
dusia; sporangia lacking hairs and glands on body,
hairs on stalk with yellow gland, often 2; spores
not seen.
Distr. Malesia: W. New Guinea. Only known
from the type.
Ecol. At 1700 m, “a large clump-fern common
in rain-forest gullies”’.
73. Sphaerostephanos novae-britanniae HOLT-
TUM, Sp. nov.
Caudex erectus; pinnae redactae 7-jugatae,
superiores 7mm longae; pinnae normales opposi-
tae, usque 8 X 1.2 cm, 2/3 costam versus lobatae;
venae infimae raro anastomosantes; costae subtus
pilis erectis praeditae; pagina inter venas subtus
glandulifera, supra pilis brevibus suberectis
praedita; indusia brevi-pilosa, sporangia
glandulifera. — Type: STEVENS & LELEAN LAE
58644, New Britain, Subdistr. Pomio, 890m (K).
Caudex erect, to 15cm tall. Stipe 4cm long,
scales at base thin, 8x 1mm, closely setiferous;
base of stipe to first large pinna 27cm. Reduced
pinnae 3cm apart, c. 7 pairs, uppermost 7mm
long, 5mm wide at slightly asymmetric base,
edges lobed, apex acute. Lamina 40cm long;
pinnae 22 pairs, almost all opposite, basal ones
with 3 basal lobes gradually smaller, base 6mm
wide, not auricled; aerophores not elongate. Hairs
on lower surface of rachis 0.3-0.4 mm long, erect,
on upper surface more than 0.5 mm long. Largest
pinnae 8X1.2cm; base broadly cuneate, apex
acuminate with cauda 10 x 2mm; edges lobed 2/3
to costa; costules 2.5 mm apart, at 60°; veins to 7
pairs, basal veins either meeting just below the
short sinus-membrane or touching its sides, rarely
joining to form an excurrent vein below the sinus;
hairs on lower surface of costae as rachis, few on
costules, surface between veins densely glandular
with some short erect hairs; hairs on upper sur-
face of costae pale, more than 0.5mm _ long,
similar hairs scattered on costules and veins, sur-
face between veins bearing suberect hairs 0.2 mm
long and sometimes a few glands. Sori in-
framedial, basal ones not divergent; indusia large,
firm, short-hairy; sporangia sometimes bearing
glands.
Distr. Malesia: New Guinea (New
only known from the type.
Britain),
74. Sphaerostephanos convergens HOLTTUM, sp.
nov.
Pinnae redactae 4—S—jugatae, omnes parvae:
aerophora non elongata; pinnae usque 6.5 X
1.3cm, 3/5 costam versus lobatae; venae infimae
conniventes, non anastomosantes; costae subtus
pilis erectis usque 1mm longis praeditae: pagina
inter venas utrinque pilis brevibus erectis praedita,
subtus etiam glandulifera; indusia pilosa;
sporangia glandulifera. — Type: M. G. PRICE &
B. F. HERNAEZ 164, Western Samar, 500m (K).
Caudex short, suberect, bearing a close tuft of
fronds. Stipe 4cm long; basal scales 7 x 0.8mm,
thin; base of stipe to first large pinna 20cm.
Reduced pinnae 4-S pairs, uppermost 2 mm long;
aerophores swollen, hardly elongate. Lamina
28cm long; pinnae 10-12 pairs, lowest 1-2 pairs
somewhat narrowed at their bases. Hairs on lower
surface of rachis erect, pale, 0.3mm long, with
some more than | mm long also; hairs on upper
surface more than 1mm long. Largest pinnae
6.5 x 1.3. cm; base truncate; apex short-acuminate;
edges lobed c. 3/5 to costa, lobes slightly falcate,
separated by sinuses nearly | mm wide; costules
4mm apart, at 60°; veins to 6 pairs, basal pair both
touching sides of short sinus-membrane or meet-
ing just below it without fusing, next pair to edge;
hairs on lower surface of costae and costules
erect, as on rachis but shorter, surface between
veins bearing many short erect hairs and glands;
hairs on upper surface of costae 1mm long,
similar hairs scattered on costules and veins, sur-
face between veins bearing suberect hairs 0.2 mm
long. Sori inframedial, basal ones not divergent;
indusia bearing hairs 0.4 mm long; sporangia bear-
ing glands.
Distr. Malesia: Philippines (W. Samar), only
known from the type.
Ecol. In forest on limestone soils.
75. Sphaerostephanos unitus (L.) HOLTTUM, J. S.
Afr. Bot. 40 (1974) 165; Kalikasan 4 (1975) 63. —
Polypodium unitum L. Syst. Nat. ed. 10. 2 (1759)
1326, excl. syn.— Aspidium unitum (L.) Sw. in
Schrad. J. Bot. 1800, 2 (1801) 32, nomen tantum;
WILLD. Sp. Pl. ed. 4, 5 (1810) 241. — Dryopteris
unita (L.) O. KTZE, Rev. Gen. Pl. 2 (1891) 811;
BACKER & PosTH. Varenfl. Java (1939) 51.—
478
FLORA MALESIANA
[ser. II, vol. 1°
Cyclosorus unitus (L). CHING, Bull. Fan Mem.
Inst. Biol. Bot. 8 (1938) 192; HOLTTUM, Rev. FI.
Mal. 2 (1955) 260, f. 147; COPEL. Fern FI. Philip.
(1960) 360.— Thelypteris unita (L.) MORTON,
Amer. Fern J. 49 (1959) 113.—Type: without
locality (LINN).
Tectaria serrata CAV. Descr. (1802) 251; C.
CHR. Dansk Bot. Ark. 9, n. 3 (1937) 16, t. 1, f.
10-11.— Nephrodium serratum (CAV.) PRESL,
Rel. Haenk. (1825) 34.— Type: NEE, Marianas
(MA).
Nephrodium insculptum DeEsv. Mém. Soc.
Linn. Paris 6 (1827) 254.—Type: Réunion, no
collector named (P).
Aspidium cucullatum BL. En. Pl. Jav. (1828)
151.— Nephrodium cucullatum (BL.) BAK. Syn.
Fil. (1867) 290; BEDD. Handb. (1883) 270; RACIB.
Fl. Btzg I (1898) 184. — Dryopteris cucullata (BL.)
CHRIST, Philip. J. Sci. 2 (1907) Bot. 194; v.A.v.R.
Handb. (1908) 213.— Type: BLUME, Java (L, n.
908, 337-5).
Nephrodium haenkeanum PRESL, Epim. Bot.
(1851) 46; HoLtTruM, Novit. Bot. Univ. Carol.
Prag. 1968 (1969) 17.— Thelypteris haenkeana
(PRESL) REED, Phytologia 17 (1968) 281. — Type:
HAENKE, Marianas (PRC).
KEY TO THE VARIETIES
1. Glands present on some part of lower surface
of pinnae.
2. Rather large glossy brownish glands present
on and between veins on lower surface
a. var. unitus
2. Smaller dull yellow glands present on costules
and veins only .b. var. mucronatus
1. Glands lacking on lower surface; cells between
veins all papilliform . ¢. var. papilliferus
a. var. unitus
Rhizome long-creeping, commonly to 5mm
diameter; stipe 10-20 cm long, base of stipe to first
normal pinna 30-60cm; reduced pinnae 6-12
pairs, lower ones very small, upper ones triangular
with symmetric base, to 10mm long. Lamina 30-
60cm long; pinnae closely placed, lowest not or
little narrowed at base, aerophores not elongate.
Hairs on both sides of rachis less than | mm long.
Largest pinnae 10-15 x 0.8-1.5cm; base broadly
cuneate; apex rather evenly attenuate; edges
lobed 1/3 towards costa; lobes rounded with a
slight point at falcate end of costule; costules
3-4mm apart; veins 8-10 pairs, 13 pairs anas-
tomosing, next 2-4 pairs passing to sides of sinus-
membrane which forms a ridge on the lower sur-
face; hairs on lower surface of costae c. 0.5 mm
long, slender, always antrorse but not closely ap-
pressed, similar hairs on costules; rather large
brownish glossy glands present on costules and
veins, and between veins; upper surface of pinnae
hairy on costae only. Sori supramedial; indusia
firm, glabrous or with a few short hairs, some-
times also glands; sporangia bearing glands near
annulus, also copious glands at the ends of hairs
on sporangium-stalks.
Distr. E. Africa; Mascarene Islands; Ceylon &
S. India; N.E. India, Burma & Thailand (few
specimens); Malesia (Malaya, Sumatra, Java,
S.W. Celebes, Lesser Sunda Is., few specimens
from Luzon and Mindanao, | from W. New
Guinea); Guam, N. Queensland.
Ecol. At low altitudes, in open places, in moist
(not swampy) ground.
Notes. The identity of this species is fixed by
the type. LINNAEUS’s description is not adequate,
and his references to plants from Asia all indicate
the species here named Cyclosorus interruptus
(WILLD.) CHING. ROBERT BROWN and others
interpreted Polypodium unitum according to the
references, and BEDDOME followed this usage. It
is now customary to accept the type as fixing the
application of a name in such cases; it is to be
noted that the specimen in the Willdenow Her-
barium agrees with the Linnaean type.
The next name, Tectaria serrata CAV., was
based on a specimen collected by NEE in Guam.
Specimens of the same species were also collected
there by HAENKE, and PRESL at first (1825)
identified them with T. serrata, but later (1851)
thought this was wrong and published the new
name Nephrodium haenkeanum for them. There is
no doubt that the two collections were identical.
b. var. mucronatus (CHRIST) HOLTTUM, comb.
nov.— Dryopteris cucullata var. mucronata
CHRIST, Philip. J. Sci. 2 (1907) Bot. 195; v.A.v.R.
Handb. (1908) 819. — Type: CUMING 182, Luzon
(CHRIST’s specimen not seen; isotypes FI-W, K,
IE):
Aspidium multilineatum METT. Farngatt. IV
(1858) 108. — Syntypes: CUMING 182, 278, Luzon
(duplicates of 278 at FI-W, G, K, LE, PRC).
Nephrodium haenkeanum sensu BAK. Syn. Fil.
(1867) 290.
Pinnae 15-20 1.0-2.0cm; veins 10-14 pairs;
glands present only on costules and veins of lower
surface, small, dull, yellow; indusia always hairy.
Distr. Polynesia (Samoa, Fiji, Tonga), Micro-
nesia (Palau), and Malesia: New Guinea, Moluc-
cas, Lesser Sunda Is. (Sumba, Wetar, Timor),
Celebes, Philippines and Borneo.
Notes. The characteristic glands on the lower
surface of costules and veins were described by
METTENIUS but ignored by all others. MET-
TENIUS however made a confusion by adopting
the name multilineatum from WALLICH and
citing WALLICH 353 from Penang, which
represents the species here named S. penniger,
very different from the CUMING specimens
which were clearly the basis of METTENIUS’s
description. BEDDOME later adopted the WAL-
LICH epithet for S. penniger.
1981]
BAKER wrongly referred CUMING 182 to
Nephrodium haenkeanum. FOURNIER also adop-
ted this name in his work on the ferns of New
Caledonia, but his citation of specimens shows
that he confused it with other species. The New
Caledonia specimens which he so named are S.
invisus.
The varietal name mucronatus was first pub-
lished by J. SMITH in 1843, as a nomen nudum
under which he cited specimens of three species;
it was not validated until CHRIST’s description of
1907. Both var. unitus and var. mucronatus occur
in the Lesser Sunda Islands; they may occupy
different habitats; records are inadequate.
c. var. papilliferus HOLTTUM, var. nov.
A typo speciei differt: costulis venisque subtus
eglandulosis; pagina subtus inter venas omnino
minute papillifera.—Type: HOOGLAND 9084,
N.E. New Guinea, Huon Peninsula, 1320m (K).
Distr. Malesia: New Guinea, many specimens,
at 1300-2000 m.
Note. All cells on the lower surface of pinnae
have small papilliform colourless outgrowths.
Stomata presumably occur, but they are not
detectable. The condition is probably an adap-
tation to extreme exposure.
76. Sphaerostephanos sessilipinna (COPEL.)
HOLTTUM, Kalikasan 4 (1975) 53.— Dryopteris
sessilipinna COPEL. Philip. J. Sci. 6 (1911) Bot.
145.— Cyclosorus sessilipinna (COPEL.) CHING,
Bull. Fan Mem. Inst. Biol. Bot. 10 (1941) 249;
COPEL. Fern FI. Philip. (1960) 353. — Thelypteris
sessilipinna REED, Phytologia 17 (1968) 313. —
Type: MERRILL 6934, Negros, Mt Canlaon
(MICH).
Dryopteris canlaonensis COPEL. Philip. J. Sci.
40 (1929) 300.— Thelypteris canlaonensis
(COPEL.) REED, Phytologia 17 (1968) 266. —
Type: MERRILL 6934, Negros (MICH).
Dryopteris austrophilippina COPEL. Philip. J.
Sci. 40 (1929) 300. — Cyclosorus austrophilippinus
(COPEL.) COPEL. Gen. Fil. (1947) 142; Fern FI.
Philip. (1960) 354. — Thelypteris austrophilippina
(COPEL.) REED, Phytologia 17 (1968) 262.—
Type: COPELAND 1705, Mindanao, San Ramon
(MICH).
Caudex short, creeping. Stipe 5cm long; base
of stipe to first large pinna 15-20cm. Reduced
pinnae 4-6 pairs, all less than 2 mm long. Lamina
to 20cm long; apical section 10-12cm long,
deeply lobed, tapering very gradually to tip, at
base with gradual transition to pinnae; free pinnae
10-12 pairs, very close; basal pinnae deflexed and
narrowed towards their bases; aerophores not
elongate. Rachis with thick curved brown hairs
both sides. Middle pinnae 2.3x0.8cm, lobed
about 1/3 to costa, apex obtuse; costules 2mm
apart; veins 2-4 pairs, basal pair anastomosing in
basal half of pinna, free near its tip; hairs on lower
THELYPTERIDACEAE (Holttum)
479
surface of costae pale, antrorsely curved; glands
present on surface between veins; upper surface
of type bearing rather sparse appressed hairs be-
tween veins, of other specimens many. Sori
medial or inframedial; indusia glabrous or with a
few short hairs; sporangia bearing glands.
Distr. Malesia: Philippines (Negros,
danao).
Notes. The type of D. austrophilippina has a
less tapering apical lamina than the other type,
and paler hairs on lower surface of rachis. In 1975
I included D. urdanetensis COPEL. also as a
synonym of the present species, but it lacks very
small basal reduced pinnae. Apart from size, there
is little to distinguish this from S. lobatus.
Min-
77. Sphaerostephanos lobangensis (C. CHR.)
HOLTTUM, comb. nov. — Dryopteris lobangensis
C. CHR. Gard. Bull. Str. Settl. 7 (1934) 245.—
Thelypteris lobangensis (C. CHR.) REED, Phy-
tologia 17 (1968) 289.— Type: CLEMENS 10728,
Mt Kinabalu, Pakka Cave to Lobang (MICH;
BO).
Caudex short, suberect. Stipe 12cm long; base
of stipe to first large pinna 50cm. Reduced pinnae
10 pairs, upper ones 3 mm long. Lamina to 40 cm
long; pinnae to 20 pairs; basal pinnae a little
reduced and deflexed, narrowed towards their
bases; aerophores not elongate. Lower surface of
rachis glabrous. Largest pinnae 6X 1.4cm; base
truncate; apex short-acuminate; edges lobed 1/3 to
costa; costules 3 mm apart, at 60°; veins to 6 pairs,
1 pair anastomosing, next acroscopic vein to side
of sinus-membrane; lower surface of pinnae quite
glabrous, with yellow glands throughout; hairs on
upper surface of costae short, on costules shorter,
few on veins, no others. Sori inframedial, basal
ones not divergent; indusia small, glandular;
sporangia bearing glands.
Distr. Malesia: Sabah (Mt Kinabalu), in forest
at 1500-2000 m; two collections, the second being
an unnumbered CLEMENS specimen from Marei
Parei spur “on rocks in stream” (BO).
Notes. The above description is prepared from
the Bogor isotype; the type at MICH is somewhat
smaller. CHRISTENSEN described the rhizome as
creeping with widely-spaced stipes, but this is not
true of the Bogor specimen.
78. Sphaerostephanos gymnorachis HOLTTUM, sp.
nov.
Caudex erectus; pinnae redactae usque 8-10-
jugatae, superiores 3 mm longae; pinnae normales
fertiles usque 16X1.9cm, 1/2-5/8 costam versus
lobatae; rachis subtus glabra, costae costulaeque
pilis brevibus antrorsis vestitae, pagina inter venas
glandulosa; indusia glabra; sporangia glanduli-
fera.—Type: HoLttTumM 34, Mt Kinabalu,
Pinosok area, Nov. 1972 (K).
Caudex erect, at least to 10cm tall. Stipe 10cm
long, dull reddish; base of stipe to first large pinna
480
FLORA MALESIANA
[ser. II, vol. 1°
of largest fronds 50cm. Reduced pinnae 4-5 cm
apart, 8-10 pairs, uppermost 3mm long, lowest
with no evident blade. Lamina to 70cm long;
pinnae more than 20 pairs, in most cases all
opposite; basal pinnae gradually narrowed
towards base in basal 4cm, next 2-3 pairs less
narrowed; aerophores not elongate. Lower sur-
face of rachis glabrous; hairs on upper surface
0.5mm long, brown. Largest fertile pinnae 16x
1.9cm (sterile to 2.3cm wide); base broadly
cuneate; apex acuminate with cauda 1-1.5cm;
edges lobed a little more than > way to costa
(about 3 way on upper pinnae or those of small
fronds); lobes falcate, distinctly tapered to a blunt
tip; costules 3.5-4mm apart (to 5mm in sterile
fronds); veins to 12 pairs, basal pair anastomosing
with long excurrent vein to sinus, second pair both
to sides of sinus-membrane; lower surface of cos-
tae near base covered with hairs 0.1-0.2 mm long,
near apex 0.3-0.4mm, antrorse but not closely
appressed, on costules similar, sparser and shorter
on veins, many glands on surface between veins
(hairs on lower surface of sterile pinnae somewhat
longer, distal ones on costae 0.5mm); hairs on
upper surface of costae brown, 0.7 mm long, on
costules 0.2 mm, pale, rarely with a solitary longer
hair; surface between veins glabrous or with a few
pale appressed hairs. Sori medial, basal ones
somewhat divergent; indusia large, glabrous;
sporangia bearing glands.
Distr. Malesia: Sabah (Mt Kinabalu), in forest
at 1500 m.
Notes. In addition to the type collection,
CLEMENS 28233 and 28382 (from Tenompok,
1500 m) have pinnae not opposite and upper sur-
face between veins covered with many appressed
hairs; CLEMENS 29050 (Silau Basin,.2100 m) is a
smaller plant (pinnae to 9x 1.5cm) with almost
sterile fronds, differing in scattered pale hairs
1mm long on lower surface of rachis. S. gym-
norachis may be a fully developed form of S.
lobangensis.
79. Sphaerostephanos pullenii HOLTTUM, sp. nov.
Pinnae redactae 6-jugatae vel ultra, omnes
parvae; pinnae normales 263.0cm, 3/4-4/5
costam versus lobatae; rachis costaeque subtus
glabra vel pilis minutis paucis praedita; pagina
subtus inter venas glandulosa, supra pilis erectis
brevibus praedita; indusia parva, glandulifera,
sporangia nec glandulis nec setis instructa. —
Type: R. PULLEN 672, N.E. New Guinea, East-
ern Highlands (BM; L, LAE).
Caudex erect. Stipe 30cm long, glandular; base
of stipe to first long pinna 85cm. Reduced pinnae
6 cm apart, at least 6 pairs, all very small. Lamina
85cm or more long; pinnae to 30 pairs; basal
pinnae narrowed towards their bases, basal lobes
6-10mm long; aerophores not elongate. Lower
surface of rachis glabrous or with sparse minute
erect hairs; upper surface with pale hairs less than
Imm long. Largest pinnae 26 x 2.0 cm; base trun-
cate; apex caudate-acuminate (cauda to 3cm);
edges lobed to 3-3.5 mm from costa, lobes slightly
falcate; costules 5-7 mm apart, at more than 60°;
veins to 12 pairs, basal pair at a wide angle to
costule meeting to form a long excurrent vein, one
or both of next pair passing to sinus-membrane;
lower surface of costae near base with sparse
minute hairs, somewhat longer hairs distally, cos-
tules similar; copious glands between veins; hairs
on upper surface of costae 0.5mm long, shorter
on costules with a few long ones near apices of
lobes, surface between veins bearing very short
suberect hairs. Sori medial, lower ones divergent,
lowest somewhat elongate; indusia small, thin,
with marginal glands; sporangia bearing neither
glands nor setae.
Distr. Malesia: Papua New Guinea.
Ecol. In Nothofagus forest at 2000 m.
80. Sphaerostephanos gregarius (COPEL.) HOLT-
TUM, comb. nov. — Cyclosorus gregarius COPEL.
J. Arn. Arb. 24 (1943) 440; Philip. J. Sci. 78 (1951)
451, pl. 31. — Thelypteris gregaria (COPEL.) REED,
Phytologia 17 (1968) 280.— Type: BRASS 6759,
Papua, Fly River (MICH; BM, BO, L).
Caudex short; erect or suberect. Stipe 25-30 cm
long, dull reddish; base of stipe to first large pinna
60cm. Reduced pinnae 7 pairs, lowest 3-4mm
long, uppermost 2.2 cm long, broadly triangular,
subentire, base slightly asymmetric. Lamina
60 cm long; apex on type pinna-like, on isotype at
L not pinna-like; pinnae 16 pairs, lower ones 4cm
apart, subopposite, not narrowed towards their
bases, upper ones close and alternate; aerophores
not elongate. Rachis glabrous on lower surface or
with minute appressed hairs; hairs on upper sur-
face not over 1mm long. Largest pinnae 15 x
1.5cm; base subequally broadly cuneate and a
little dilated both sides; apex acuminate; edges
lobed to a depth of 2-2.5mm (on BM and BO
isotypes little more than | mm); costules 3.5-4 mm
apart, at 60° or more; veins 6-7 pairs, 2-3 pairs
anastomosing, | pair to short sinus-membrane;
lower surface of costa glabrous or with minute
appressed hairs, no other hairs on lower surface,
sparse glands present on costules and on surface
between veins; hairs on upper surface of costae
1mm long, rest of upper surface glabrous apart
from short hairs on margin. Sori medial or in-
framedial, lowest not divergent; indusia very
small, thin, bearing a few very short hairs;
sporangia bearing glands.
Distr. Malesia: Eastern New Guinea, at al-
titudes to 1000 m, in forest.
81. Sphaerostephanos_ tephrophyllus (COPEL.)
HoOLtTTuM, Kalikasan 4 (1975) 58.— Dryopteris
tephrophylla COPEL. Philip. J. Sci. 40 (1929)
296. — Cyclosorus tephrophyllus (COPEL.)
CopEL. Gen. Fil. (1947) 143; Fern FI. Philip.
1981]
(1960) 350.— Thelypteris tephrophylla (COPEL.)
REED, Phytologia 17 (1968) 319.—Type:
COPELAND s.n. 18 April 1905, Mindanao, Zam-
boanga, San Ramon (MICH).
Caudex suberect, short. Stipe 3-4 cm long; base
of stipe to first large pinna 12-15cm. Reduced
pinnae 3-5 pairs, lowest 3-5 mm long, uppermost
to 1.5x0.8cm, entire, deflexed, auricled on
acroscopic base; transition to normal pinnae
not abrupt. Lamina to 25cm long; apical sec-
tion to 12cm, deeply lobed and grading to upper
pinnae; free pinnae 5-6 pairs, lower ones slightly
reduced and sometimes a little auricled on
acroscopic base; aerophores not elongate.
Hairs on lower surface of rachis short, with
sparse longer ones, on upper surface similar but
thicker, the long ones more than 1mm. Largest
pinnae 4.5 x 1.4cm (sterile), 3.5 x 1.2 cm (fertile):
base truncate; apex short-pointed; edges crenate
to a depth of less than 1 mm; costules 3-3.5mm
apart; veins to 4 pairs, 1 pair anastomosing, next
pair to sides of sinus-membrane; hairs on lower
surface of costae and costules short, with a few
longer ones, between veins short hairs and glands:
short hairs on upper surface of costae, between
veins very short suberect hairs. Sori medial; in-
dusia large, with a few hairs; sporangia sometimes
with a gland.
Distr. Malesia: Philippines (Mindanao), 3 col-
lections.
Note. Allied to S. spenceri, differing in little but
much smaller size.
82. Sphaerostephanos pilososquamatus (v.A.v.R.)
HOLTTUM, comb. nov. — Dryopteris pilososqua-
mata v.A.v.R. Bull. Dép. Agr. Ind. Néerl. 21
(1908) 4; Handb. (1908) 222: Handb. Suppl. (1917)
181, excl. var. obtusata.— Thelypteris piloso-
squamata (v.A.v.R.) REED, Phytologia 17 (1968)
304.— Type: cult. Hort. Bog., origin W. New
Guinea (BO; BM, L).
Dryopteris paraphysata COPEL. Philip. J. Sci. 6
(1911) Bot. 74. — Thelypteris paraphysata REED,
Phytologia 17 (1968) 301.— Type: C. KING 306,
Papua (MICH; NSW, P).
Dryopteris megaphylloides ROSENST. Fedde
Repert. 12 (1913) 174.— Thelypteris megaphyl-
loides (ROSENST.) REED, Phytologia 17 (1968)
292. Type: KEYSSER 120, N.E. New Guinea,
Sattelberg (orig.?; UC). — Fig. 12i-k.
Caudex short, creeping. Stipe on a well-grown
plant 30-40cm long; base of stipe to first large
pinna 45-SScm (sterile) 60-70cm (fertile).
Reduced pinnae 2-3(-4) pairs, largest 5 mm long;
an intermediate pair sometimes present. Lamina
dimorphous; pinnae to 15 pairs, sometimes with
stalks 1-2mm long; basal pinnae narrowed
towards their bases; apical lamina almost pinna-
like; aerophores not elongate. Hairs on rachis of
sterile fronds to 1mm long both sides, rather
sparse on lower surface. Largest sterile pinnae
THELYPTERIDACEAE (Holttum)
481
20-27 x 3.5-4cm; base broadly cuneate; apex
acuminate with cauda to 2 cm; edges lobed 1/4~-2/5
to costa, lobes falcate, narrowed to tips in more
deeply lobed pinnae; costules 5—5.5 mm apart, at
60° or rather less; veins 12-15 pairs, 2+33 pairs
anastomosing, 2-3 pairs to sides of sinus-mem-
brane; hairs on lower surface of costae spreading,
0.5—1 mm long, not dense, on costules and veins
similar but fewer, glands usually present on cos-
tules and veins and between veins, not abundant:
scattered hairs on upper surface of costules and
veins like those on costae, appressed hairs be-
tween veins sometimes present. Fertile pinnae
commonly to 18x2cm, less deeply lobed than
sterile and with closer costules, hairs on all parts
shorter and more sparse; veins 7-9 pairs: sori
medial, lower ones not or little divergent; indusia
bearing glands and hairs; sporangia usually with a
seta, sometimes also a gland, near annulus, and a
gland at end of hair on stalk.
Distr. Malesia: New Guinea,
Archipelago, in forest, 0-1400 m.
Note. One specimen assigned to this species
appears to be quite eglandular. Another (NAK-
AIKE 73) is small, lacks reduced pinnae and has
no glands on sporangia.
Bismarck
83. Sphaerostephanos angustifolius (PRESL)
HOLTTwuM, Kalikasan 4 (1975) 66. — Nephrodium
angustifolium PRESL, Epim. Bot. (1851) 48:
Hook. Spec. Fil. 4 (1862) 69.— Cyclosorus
angustifolius (PRESL) COPEL. Fern Fl. Philip.
(1960) 349. — Thelypteris setulosa REED, Phytolo-
gia 17 (1968) 313, nom. nov. — Type: CUMING 268,
Luzon (PRC; B, BM, E, FI-W, G, K, LE, US).
Dryopteris hispidula sensu C. CHR. Ind. Fil.
(1905) 271, p.p.; v.A.v.R. Handb. (1908) 228.
Caudex short, erect. Stipe 46cm long, dark
reddish, densely short-hairy; base of stipe to first
large pinna 15cm or more. Reduced pinnae to 5
pairs, highest 3mm long. Lamina to 35cm long:
pinnae c. 17 pairs; lower 2-3 pairs somewhat
narrowed towards their bases with basal acros-
copic lobe a little elongate: aerophores not elon-
gate. Lower surface of rachis covered with short
erect hairs and scattered longer ones (some of
them dark red), upper surface covered with pale
hairs 0.5mm long. Largest pinnae 9.5 x 0.8cm;
base truncate and not dilated: apex a subentire
cauda 1.5-2.0 cm long; edges lobed 3 way to costa,
lobes oblique; costules 3mm apart, at 60°; veins
4-5 pairs, 1 pair anastomosing, next pair to mar-
gin; hairs on lower surface of costae not dense,
very short, with scattered hairs to 0.8 mm long, on
costules similar but more sparse, minute erect
hairs present between veins, glands throughout
lower surfaces; hairs on upper surface of costae
0.5mm long, similar hairs scattered on costules
and veins, suberect hairs 0.1-0.2 mm long between
veins. Sori medial; indusia short-hairy; sporangia
bearing glands.
482
FLORA MALESIANA
(ser. II, vol. 1°
Distr. Malesia: Philippines (Luzon); a second
collection is M. G. PRICE 2553, Zambales Prov.,
from exposed place by stream (small plants).
Note. S. angustifolius is very near S. hetero-
carpus; it is peculiar in its very narrow pinnae.
84. Sphaerostephanos efogensis HOLTTUM, sp.
nov.
S. conferto (Brause) affinis, differt: membranis
sinuum brevibus; costis subtus pilis usque 1mm
longis vestitis; costulis subtus glanduliferis; soris
supramedialibus, sporangiis glanduliferis. — Type:
J. R. Crorr LAE 61873, Papua, Central Distr.,
Port Moresby Subdistr. Efogi (K).
Caudex short, decumbent (collector). Stipe 12-
20cm long, basal scales short and thick, ap-
parently 1.5 1mm; base of stipe to first large
pinna 40-60 cm. Reduced pinnae 12 pairs, lowest
2-4 mm long, middle ones 10 x 10mm, uppermost
1.7cm long, 1.3cm wide at truncate base, trian-
gular, crenate. Lamina 43cm long; apex broadly
deltoid and deeply lobed; pinnae 20 pairs, basal
ones not narrowed at base; aerophores not elon-
gate. Hairs on lower surface of rachis not dense,
mostly 0.1-0.3mm long with some more than
Imm, on upper surface long hairs more
numerous, to 1.5mm long. Largest pinnae 8.5 x
1.7 cm: base subtruncate, almost symmetric; apex
short-acuminate; edges lobed to a depth of 2mm,
lobes oblique, subtriangular; costules to 3.5mm
apart; veins 7-8 pairs, all except basal ones very
oblique, 2-25 pairs anastomosing to form a zig-zag
excurrent vein, | pair to short sinus-membrane;
hairs on lower surface of costae as on rachis, on
costules shorter, glands present on costules and
veins, between veins sparse short erect hairs and
sometimes a few glands; hairs on upper surface of
costae mostly 0.3mm long with scattered long
ones which occur also on costules and veins,
surface between veins bearing appressed hairs
0.2-0.3 mm long. Sori a little supramedial; indusia
thin, with a few hairs 0.5mm long; sporangia
bearing glands.
Distr. Malesia: Papua New Guinea.
Ecol. At 1500 m in riverine hill forest.
85. Sphaerostephanos dichrotrichoides (v.A.v.R.)
HOLTTUM, Kalikasan 4 (1975) 65.— Dryopteris
dichrotricha COPEL. Philip. J. Sci. 7 (1912) Bot.
54, non COPEL. 1911.—Dryopteris di-
chrotrichoides v.A.v.R. Handb. Suppl. (1917)
Corr. 48, nom. nov. — D. weberi COPEL. Philip. J.
Sci. 38 (1929) 135, nom. nov. superfl. — Lastrea
dichrotrichoides (v.A.v.R.) COPEL. Fern FI.
Philip. (1960) 329.— Thelypteris dichrotrichoides
(v.A.v.R.) REED, Phytologia 17 (1968) 272.—
Type: WEBER 1173, Mindanao, Mt Hilong-Hilong
(MICH; E, G, K, NSW, P, US).
Caudex short, creeping. Stipe 15cm long, pale,
short-hairy; base of stipe to first large pinna
30cm. Reduced pinnae to 5 pairs, upper ones
4mm long, auricled, lowest very small. Lamina
35cm long; pinnae 18 pairs, basal pinnae a little
narrowed towards their bases, strongly auricled on
acroscopic base; aerophores swollen. Hairs on
both surfaces of rachis 1 mm long, pale. Largest
pinnae 7-12cm long, 1.5cm wide at broadly
cuneate base from which they are gradually
attenuate to apex; edges lobed to 2mm from
costa, lobes oblique, slightly falcate; costules to
3mm apart; veins 8-9 pairs, basal pair anas-
tomosing near base of pinna, passing to sides of a
short sinus-membrane towards apex of pinna;
coarse rather sparse hairs 1mm long on lower
surface of costae and costules, shorter hairs on
veins and surface between them, glands present
on costules and veins, few between veins; hairs on
upper surface of costae 1mm long, similar hairs
scattered on costules and veins, whole upper sur-
face covered with fine appressed hairs nearly
0.5mm long. Sori medial, basal ones not diver-
gent; indusia with copious hairs 0.5mm _ long;
sporangia often with 2-3 glands.
Distr. Malesia: Philippines
Panay”).
(Mindanao:
86. Sphaerostephanos nakaikei HOLTTUM, sp.
nov.
Pinnae redactae S-jugatae, superiores 10mm
longae, valde auriculatae; pinnae normales usque
14 x 2.3 cm, 2/3-3/4 costam versus lobatae; venae
infimae anastomosantes vel ad basin sinus con-
niventes; costae subtus pilis erectis brevibus ves-
titae; pagina inter venas subtus glandulifera,
supra pilis appressis vestita; indusia glandulifera,
sporangia non glandulifera. — Type: T. NAKAIKE
358, E. New Guinea, Mt Wilhelm 2500-3500 m (K;
TNS).
Caudex ‘“‘massive or ascending” (NAKAIKE).
Stipe 20cm long, short-hairy, basal scales broad
and thin; base of stipe to first large pinna 45 cm.
Reduced pinnae 5 pairs, lowest 2mm long, up-
permost 10 mm with basal acroscopic auricle 5mm
long; 1-2 pairs pinnae of intermediate length
sometimes present. Lamina 60-70 cm long; pinnae
25 pairs; basal large pinnae not narrowed at base,
slightly auricled; aerophores slightly elongate.
Hairs on lower surface of rachis dense, brownish,
erect, 0.3-0.5 mm long, on upper surface a little
longer. Largest pinnae 13-14 cm long, 1.8-2.3cm
wide; base truncate; apex acuminate; edges lobed
2/3-3/4 to costa, lobes falcate, separated by dis-
tinct sinuses; costules 5mm apart; veins 8-10
pairs, basal pair anastomosing with a short excur-
rent vein to sinus, less commonly both touching
sides of short sinus-membrane; lower surface of
costae densely covered with erect hairs 0.2mm
long, hairs on costules sparse, glands present on
costules and veins and on surface between veins;
hairs on upper surface of costae 0.7mm long,
similar hairs scattered on costules and veins, sur-
face between veins rather sparsely covered with
1981]
THELYPTERIDACEAE (Holttum)
483
appressed hairs. Sori medial; indusia bearing
glands; sporangia with neither setae nor glands.
Distr. Malesia: Papua New Guinea (Mt Wil-
helm and Mt Giluwe: PARRIS & CROXALL 5900,
at 2950 m).
87. Sphaerostephanos woitapensis HOLTTUM, sp.
nov.
S. nakaikeo affinis, differt: pinnis minoribus,
brevi-stipitatis, inferioribus sensim decrescenti-
bus; lobis pinnarum distaliter dentatis; venis
infimis semper anastomosantibus. — Type:
NAKAIKE 576, Papua, Central Distr., Woitape
(TNS; K).
Caudex thick, short-creeping or ascending.
Stipe 25cm long, copiously short-hairy. Lamina
to 75cm long: pinnae to more than 20 pairs,
distinctly stalked: 5-6 lower pairs gradually
decrescent and more widely spaced, lowest 8 mm
long; aerophores slightly swollen. Hairs on lower
surface of rachis suberect, 0.2-0.4mm long, on
upper surface 0.7mm long. Largest pinnae 9.5 x
2.0cm; base truncate and in some cases slightly
dilated both sides; apex short-acuminate: edges
lobed c. 3/5 to costa, lobes oblique. slightly fal-
cate, with distinct teeth at ends of some distal
veins; costules 44.5mm apart at 60°; veins 6~7
pairs, pale and prominent both sides, basal pair
always anastomosing, second pair both to margin:
hairs on lower surface of costae near base 0.1 mm,
distally 0.3 mm long, on costules and veins sparse
and very short, glands present on surface between
veins; upper surface of costae hairy as rachis,
scattered similar hairs on costules and veins, ap-
pressed hairs 0.2mm long between veins. Sori
medial; indusia with short hairs and many glands:
sporangia lacking glands and setae.
Distr. Malesia: Eastern New Guinea at 2500—
2800 m.
Note. Other collections have smaller pinnae
than the type, on smallest fronds 5.0 1.2cm,
rather rigid; these do not show teeth at ends of
distal veins.
88. Sphaerostephanos ekutiensis HOLTTUM, sp.
nov.
Pinnae redactae 8-jugatae. usque 10mm lon-
gae; pinnae normales usque 7.0*1.8cm, c. 3/5
costam versus lobatae: venae 7-jugatae, infimae
solum anastomosantes: costae subtus pilis paucis
praedita, pagina inferior pinnarum cetera glabra,
glandulifera, glandulis inter venas paucis: sori in-
framediales; indusia tenuia, glandulis multis
omata: sporangia nec _ setis nec glandulis
praedita.— Type: B. S. PARRIS & J. P. CROXALL
6022, N.E. New Guinea, Morobe Distr., Ekuti
Range (CGE; K).
Caudex not known; stipe 7 cm long, glabrous on
abaxial surface, scales at base 7 x 1.5 mm; base of
stipe to first normal pinna 30 cm; reduced pinnae 8
pairs, uppermost 104mm with an auricle 8mm
long, lowest 5mm long, deflexed, narrowly trian-
gular with acroscopic auricle 3 mm long; two pairs
of pinnae of intermediate length present between
reduced and normal pinnae. Lamina excluding
reduced pinnae 37cm long; pinnae 18 pairs, all
opposite, texture firm; basal pinnae not narrowed
at their bases. Both surfaces of rachis bearing
thick curved pale brown hairs | mm long. Largest
pinnae 7.0 1.8cm, sessile; aerophores slightly
swollen; base subtruncate, not auricled; apex ab-
ruptly short-caudate; edges lobed 3/5 towards
costa, lobes slightly falcate with rounded tips;
costules 4mm apart. at more than 60° to costa:
veins to 7 pairs, basal pair anastomosing, next pair
to margin; lower surface of costae bearing rather
sparse thick pale hairs 0.3mm long and glands,
rather large glands and no hairs on costules and
veins, rather sparse glands between veins: upper
surface of costae covered with pale hairs less than
1 mm long, shorter hairs scattered on costules and
veins, no other hairs. Sori inframedial; indusia
thin, with many glands: sporangia bearing neither
setae nor glands.
Distr. Malesia:
known from the type.
Ecol. In Nothofagus forest at 2250 m.
Papua New Guinea: only
89. Sphaerostephanos omatianus HOLTTUM, sp.
nov.
Stipes 2.5 cm longus; lamina 7-8 cm longa, pars
apicalis profunde lobata 5.5cm longa, pinnae
liberae 3-4-jugatae. crenatae. infimae leviter
redactae; venae liberae: costae costulaeque subtus
pilis longis glandulisque praeditae: sori exindusi-
ati, sporangia nuda.— Type: WOMERSLEY &
SIMMONDS 5076A, Papua. Gulf Div., Omati
(BRI).
Caudex apparently short-creeping. Stipe 2.5cm
long, covered with pale spreading hairs 1 mm long.
Lamina 7-8 cm long. apical 5.5 cm deeply lobed,
grading to pinnae; free pinnae 34 pairs, lowest
only reduced, 6-7 mm long, deflexed and a little
narrowed on basiscopic side. Largest pinnae 1.1 x
0.45cm, almost sessile; base broadly cuneate:
apex obtusely pointed: edges crenate near base,
entire distally; costules near base of pinnae hardly
2mm apart. twice forked, distal ones forked or
simple, all veins free: lower surface of rachis,
costae and costules bearing hairs 1 mm long and
some shorter ones, slender erect hairs on surface
between veins, glands present on costae and cos-
tules: slender hairs 1 mm long on upper surface of
costae and costules. appressed hairs on surface
generally. Sori in an uneven row on each side of
costa, on acroscopic branches of forked costules;
no indusia: neither glands nor setae on sporangia:
spores minutely papillose.
Distr. Malesia: Papua, known only from type
collection.
Ecol. At an altitude of 30m, in shallow soil
over pinnacle limestone, in dense rain-forest.
484
FLORA MALESIANA
[ser. II, vol. 1°
90. Sphaerostephanos alticola HOLTTUM, sp. nov.
Caudex brevis, crassus; stipes 12cm longus;
lamina 45-55 cm longa; pinnae 40-jugatae, 12-14-
jugatae inferiores sensim decrescentes, maximae
3.3 cm longae, 7-8 mm latae, leviter lobatae, rigidae,
lobis cucullatis; costae costulaeque subtus
patenti-pilosae paulo glandulosae; indusia parva,
glandulis pilisque praedita; sporangia setifera. —
Type: HOOGLAND 9762, N.E. New Guinea,
Huon Peninsula, Salawaket Range (LAE; L).
Caudex “thick, + horizontal”. Stipe 12 cm long,
base covered with glossy scales 10x 1.3-3 mm.
Lamina 45-55 cm long; pinnae to 40 pairs, rigidly
coriaceous, lower 12-14 pairs gradually smaller
and more widely spaced, lowest 5mm long, 6mm
wide at truncate base, 3-lobed, upper ones trian-
gular, crenate; aerophores not elongate. Rachis
covered with dense pale spreading hairs 0.7 mm
long. Largest pinnae 3.3cm long, 7-8mm wide
above base, base truncate and a little dilated; apex
shortly obtuse; edges lobed to a depth of | mm or
a little more deeply; lobes rounded, their edges
much deflexed so that the lower surface is con-
cave; costules little more than 2mm apart; veins
3-5 pairs, pale, prominent both sides, | pair anas-
tomosing, next pair to short sinus-membrane;
lower surface of costae covered with erect thick
pale hairs to | mm long, similar hairs sparse on
costules; sparse glands present on costules and
veins; upper surface of costae densely hairy,
similar hairs more sparse on costules and veins,
some erect hairs between veins. Sori medial; in-
dusia small, bearing glands and short hairs;
sporangia setiferous; spores with many small
wings.
Distr. Malesia: Papua New Guinea (Salawaket
Range); 2 collections.
Ecol. “In low open forest on landslide, lime-
stone’. 2500-3200 m.
91. Sphaerostephanos rigidus (RIDL.) HOLTTUM,
comb. nov.—Goniopteris rigida RtIDL. Trans.
Linn. Soc. Bot. 9 (1916) 258. — Phegopteris wol-
lastonii v.A.v.R. Handb. Suppl. (1917) 515, nom.
nov., not P. rigida (HOOK. & GREV.) METT. —
Thelypteris rigida (RIDL.) REED, Phytologia 17
(1968) 309.— Type: C. B. KLOss s.n. 18 Feb.
1913, W. New Guinea, Mt Carstensz, Camp VIc,
1680 m (BM; K).
Caudex lacking from type and stipe incomplete.
Lamina to 40cm long; pinnae to 30 pairs; basal
3-4 pairs pinnae gradually decrescent, lowest 2.2 x
0.6 cm, below them 3 pairs reduced pinnae 3-4 mm
long; aerophores c. 0.5mm long. Rachis bearing
throughout stiff spreading brown hairs to 1.2mm
long. Largest pinnae 6.0 1.3cm; base truncate
and slightly dilated both sides; apex narrowed to a
short obtuse or rounded tip; edges lobed 1/4-1/3
to costa, lobes rounded with edges strongly
reflexed (lower surface thus concave); costules
2.5mm apart, pale and prominent on lower sur-
face, grooved above; veins 4-5 pairs, prominent
beneath, one pair anastomosing, next acroscopic
vein to sinus-membrane; stiff brown hairs to
0.7mm long on lower surface of costae, more
sparse on costules, veins and edge, glands present
on costae and costules; upper surface of pinnae
glabrous apart from sparse short hairs on costae.
Sori medial; indusia very small with a few stiff
hairs (sometimes absent?); sporangia with neither
glands nor hairs on body, yellow glands some-
times on their stalks.
Distr. Malesia: Western New Guinea; 2 col-
lections at 1700 m; on limestone?
Note. The second collection, EYMA 4986 from
Wissel Lake region, is small, with lamina of
fronds 15cm long, pinnae to 2 cm long, 3-4 pairs
lower ones gradually reduced, lowest 8 mm long;
the caudex is short-creeping. It seems probable
that the type was found on limestone.
92. Sphaerostephanos arfakianus (BAK.) HOLT-
TUM, comb. nov.— Polypodium arfakianum
BAK. in Beccari, Malesia 3 (1880) 45.— Dry-
opteris arfakianus (BAK.) C. CHR. Ind. Fil. (1905)
253; Dansk Bot. Ark. 9, n. 3 (1937) 50. — Phegop-
teris arfakiana (BAK.) v.A.v.R. Handb. (1908)
502.— Cyclosorus arfakianus (BAK.) COPEL.
Gen. Fil. (1947) 142; Philip. J. Sci. 78 (1951)
451. — Thelypteris arfakiana (BAK.) REED, Phy-
tologia 17 (1968) 260. — Type: BECCARI s.n. 1872,
W. New Guinea, Mt Arfak at Putat (FI; K).
Dryopteris sepikensis BRAUSE, Bot. Jahrb. 56
(1920) 101; COPEL. Philip. J. Sci. 78 (1951) 443. —
Type: LEDERMANN 12053, N.E. New Guinea,
Schraderberg (B).
Dryopteris arborea v.A.v.R. Bull. Jard. Bot.
Btzg II, 28 (1918) 24, non BRAUSE 1914. — Dry-
opteris rosenburghii C. CHR. Ind. Fil. Suppl. III
(1934) 96. — Type: RUTTEN 161, Ceram (BO).
Caudex slender, erect, to at least 100cm tall.
Stipe 10-20 cm long, dark, short-hairy. Lamina to
80cm long, + dimorphous, thin; pinnae to 40
pairs, lowest 6-10 pairs gradually smaller, lower
ones broadly triangular with slightly asymmetric
base, entire; aerophores not elongate. Hairs on
both sides of rachis less than 0.5 mm_ long,
brownish, + appressed. Largest sterile pinnae 12-
17x 2.2-4cm, fertile commonly to 10x 1.5cm,
sometimes larger; base broadly cuneate to trun-
cate, sometimes a little dilated both sides; apex
acuminate; edges shallowly crenate or sinuous;
costules to 4.5mm apart on large pinnae; veins
5-7 pairs, slender, concolorous, slightly prominent
both sides, almost all anastomosing to form zig-
zag excurrent veins between costules; hairs on
lower surface of costae 0.1-0.2 mm long, slender,
appressed, similar hairs on costules and veins,
also sometimes a few glands; hairs on upper sur-
face of costae 0.3-0.4 mm long, on costules shor-
ter and sparse, usually a few slender appressed
hairs on surface between veins at least near mar-
1981]
gin. Sori inframedial; indusia small, thin, usually
with a few hairs; sporangia bearing glands.
Distr. Malesia: Moluccas (Ceram;? Amboina,
BROOKS 18102) and widely in New Guinea.
Ecol. In forest at altitudes to 2000 m.
Note. The type of D. sepikensis and some
others from an altitude of about 2000m are
smaller than BECCARI’s type and other lowland
specimens, with fewer reduced pinnae and dis-
tinctly lobed sterile ones, but agree in details of
pubescence and sori.
93. Sphaerostephanos archboldii (C. CHR.)
HOLTTUM, comb. nov.— Dryopteris archboldii
C. CHR. Brittonia 2 (1937) 297.— Thelypteris
archboldii (C. CHR.) REED, Phytologia 17 (1968)
260. — Type: BRASS 4873, Papua, Mt Tafa, valley
forest, 2400 m (BM; BRI).
Dryopteris protecta COPEL. Univ. Cal. Publ.
Bot. 18 (1942) 221.—Cyclosorus protectus
(COPEL.) COPEL. Gen. Fil. (1947) 143; Philip. J.
Sci. 78 (1951) 453, pl. 33. — Thelypteris protecta
(COPEL.) REED, Phytologia 17 (1968) 306. —
Type: BRASS 10933, New Guinea, 9km N.E. of
Lake Habbema, 2800m (MICH; BM, L). — Fig.
13d-e.
Caudex slender, erect, to 100cm or more tall.
Stipe 5-15cm long, minutely hairy, when young
covered with rather firm brown scales 7 X 1mm.
Lamina to 65cm long, texture very firm; lower
8-15 pairs pinnae gradually smaller, lowest 3-6
pairs of these more widely spaced with asym-
metric base, lowest c. 1 mm long; aerophores not
elongate. Hairs on lower surface of rachis erect,
to 0.5mm long, on upper surface a little longer,
also spreading. Largest pinnae of type to 6x
10cm (type of D. protecta 9x1.5cm); base
truncate; apex evenly attenuate; edges lobed
about 1/3 to costa; costules to 3mm apart; veins
4-6 pairs, prominent on lower surface, 1-1} pairs
anastomosing, next acroscopic vein or pair to
short sinus-membrane; hairs on lower surface of
costae 0.1-0.3 mm long, antrorse distally, sparse
glands and hairs on costules and veins (glands best
seen on Sterile fronds); hairs on upper surface of
costae short, sparse on costules, veins and be-
tween veins. Sori near costules; indusia bearing
hairs and glands; sporangia sometimes with a seta.
Distr. Malesia: Eastern New Guinea at 1800—
2800 m, many collections.
94. Sphaerostephanos_ tibangensis (C. CHR.)
HOLTTUM, comb. nov.— Dryopteris tibangensis
CG Gur. Dansk Bot Ark 9) 13° (1937)66.—
Pronephrium tibangense (C. CHR.) HOLTTUM,
Blumea 20 (1972) 118.—Type: MJOBERG s.n.
Oct.-Dec. 1925, E. Kalimantan, Mt Tibang, 1400-
1700 m (BM; S-PA).
Caudex short, suberect; stipe to 20cm long,
short-hairy. Lamina to 20cm long; no reduced
basal pinnae; normal pinnae 8-10 pairs, lower
THELYPTERIDACEAE (Holttum)
485
ones with stalks to 2mm long; basal pinnae
slightly shorter than next pair, with 3 pairs of
basal basiscopic lobes slightly reduced. Largest
pinnae 3.5 1.4cm; base truncate and slightly
auricled on the acroscopic side, narrowed rather
evenly from base to acute apex; edges lobed 2/5
towards costa or rather more deeply; lobes sub-
falcate, acute; costules 3mm apart, at more than
60° to costa; veins 6 pairs, basal pair anastomos-
ing, second pair to sides of sinus-membrane, rest
to margin; lower surface of costae and costules
bearing pale spreading hairs 0.2-0.6mm _ long,
short erect hairs present between veins, many
glands on costules and veins; upper surface of
costae copiously short-hairy, rest of surface bear-
ing short suberect hairs. Sori supramedial; indusia
small, densely short-hairy; sporangia not seti-
ferous.
Distr. Malesia:
1400-1700 m.
Note. This is certainly very near S. norrisii.
The type consists of one fertile and one sterile
frond, the latter not quite fully expanded, with
immature sori. The fertile frond is smaller than
any specimen undoubtedly referable to S. norrisii,
and the pinnae are of a different shape; mature
sori might show further differences.
East Borneo (Mt Tibang),
95. Sphaerostephanos norrisii (ROSENST.)
HOLTTUM, comb. nov.— Dryopteris — norrisii
ROSENST. Med. Rijksherb. n. 31 (1917) 8.—
Type: W. NORRIS, Malaya (L; K).
Nephrodium pennigerum var. malayense BEDD.
Handb. Suppl. (1892) 74, excl. Parish,
Tenasserim. — Dryopteris indica var. malayensis
(BEDD.) v.A.v.R. Handb. (1908) 224. — Lectotype
(selected here): KUNSTLER 2360, Perak (K).
Dryopteris subfalcinella v.A.v.R. Bull. Jard.
Bot. Btzg III, 2 (1920) 151. — Thelypteris subfal-
cinella (v.A.v.R.) REED, Phytologia 17 (1968)
317. — Lectotype (selected here): LORZING 5338,
Sumatra, Bandar-baroe (BO; L).
Dryopteris elmerorum COPEL. Philip. J. Sci. 40
(1929) 295, pl. 2.—Cyclosorus elmerorum
(COPEL.) COPEL. Gen. Fil. (1947) 142; Fern FI.
Philip. (1960) 351.— Thelypteris elmerorum
(COPEL.) REED, Phytologia 17 (1968) 274. —S.
elmerorum (COPEL.) HOLTTUM, Kalikasan 4
(1975) 53.—Type: COPELAND s.n. Nov. 1911,
Mindanao, San Ramon (MICH).
Nephrodium indicum sensu RIDL. J. Mal. Br. R.
As. Soc. 4 (1926) 73.
Dryopteris toppingii sensu C. CHR. Gard. Bull.
Str. Settl 4 (1929) 391.—Cyclosorus toppingii
sensu HOLTTUM, Rev. FI. Mal. 2 (1955) 280, f. 161.
Cyclosorus ellipticus sensu COPEL. Fern FI.
Philip. (1960) 351. — Fig. 12f-h.
Caudex short-creeping. Stipe 30-90cm long,
dull reddish, minutely hairy; reduced pinnae 0-3
pairs, present only on largest fronds, irregularly
spaced and very small. Lamina to 100cm long
486
(type), on type of D. elmerorum 30cm long; pin-
nae 10-18 pairs, several pairs distinctly stalked;
basal pinnae often somewhat reduced if small
reduced pinnae absent, with stalks 1-3 mm long,
narrowed towards their bases, not auricled; aero-
phores not elongate. Hairs on lower surface of
rachis dense, short, stiff, erect; on upper surface
copious, appressed, less than 0.5 mm long. Largest
pinnae of type 24x3.4cm, of type of D.
elmerorum 6.5 X 1.8cm; base subtruncate on large
fronds, subcordate on small ones; apex rather
abruptly short-acuminate; edges lobed c. 2/5
towards costa (less deeply on small fronds), lobes
falcate with broadly pointed or rounded forward-
pointing tips; costules 5-6 mm apart (3.5-4 mm in
type of D. elmerorum); veins to 12 pairs, 1-13
pairs anastomosing, 2-23 pairs passing to sides of
sinus-membrane; hairs on lower surface of costae
and costules dense, short, erect, some short hairs
also between veins, glands present on costules at
least near apex of pinna-lobes; hairs on upper
surface of costae and costules short, some short
appressed or suberect hairs on surface between
veins. Sori supramedial, somewhat elongate,
especially the lower ones; indusia large, densely
short-hairy, sometimes with a _ few glands;
sporangia sometimes with a gland or a short seta.
Distr. Malesia: Malaya, Sumatra, Borneo,
Philippines (Mindanao).
Ecol. In forest at 1000-1500 m; apparently not
common anywhere.
Note. Though the type of D. elmerorum is
small, other specimens from Mindanao have pin-
nae to 15x3cm. No small fertile fronds have
been found on plants in Sumatra and Malaya.
Some specimens have very few glands on the
lower surface of costules. Specimens from Suma-
tra and Malaya have appressed hairs between the
veins on the upper surface, specimens from Bor-
neo and Mindanao have shorter suberect hairs.
96. Sphaerostephanos semimetralis HOLTTUM sp.
nov.
S. arfakiano affinis, differt: frondibus minori-
bus; pinnis redactis 2-3-jugatis, minutis; pinnis
fertilibus usque 5X1.2cm; costis costulisque
subtus pilis sparsis suberectis instructis. — Type:
PULLE 518, W. New Guinea, Perameles Mts,
1100 m, on limestone (L).
Caudex erect, slender, 50cm tall. Stipe 15cm
long, basal scales c. 8 x 1 mm; base of stipe to first
large pinna 25-30cm. Reduced pinnae 2-3 pairs,
all very small, with above them an apparently
much larger intermediate pair (incomplete).
Lamina 35 cm long; pinnae 10-12 pairs, very firm,
dimorphous; basal pinnae not narrowed at base;
aerophores less than 0.5 mm long. Hairs on lower
surface of rachis thick, brown, 0.5mm long, on
upper surface similar, to 1 mm long. Largest sterile
pinnae 9X2.2cm; base truncate with a winged
stalk 1mm long; apex abruptly short-acuminate;
FLORA MALESIANA
[ser. II, vol. 1°
edges crenate to a depth of 1-2mm; costules
3.5-4mm apart, at 60°; veins 6-8 pairs, very
oblique except basal ones, prominent both sides, 2
pairs anastomosing to form a zig-zag excurrent
vein, | pair to short sinus-membrane; lower sur-
face of costae bearing sparse erect hairs 0.1 mm
long, similar hairs very sparse on costules and
veins with a few glands, sparse erect hairs be-
tween veins; hairs on upper surface of costae
pale, 0.5 mm long, rest of surface glabrous. Fertile
pinnae to 5 x 1.2. cm; veins 5 pairs; pubescence as
sterile; sori near costules; indusia small with a few
short hairs and glands; sporangia sometimes with
a gland.
Distr. Malesia: New Guinea. A specimen from
N.E. New Guinea, Western Highlands, at Kompiai
in the Jimmi valley at 1900m (MANNER &
STREET 454), probably not from limestone, is
similar to the type.
Note. This differs from the mountain form of
S. arfakianus in the very small reduced pinnae,
and in pubescence of lower surface; but better
specimens are needed to characterize it clearly.
97. Sphaerostephanos semicordatus HOLTTUM, sp.
nov.
Pinnae redactae 8-jugatae, omnes minutae;
lamina 35 cm longa; pinnae 10-12-jugatae, omnes
oppositae, subintegrae; costulae subtus glabrae,
glanduliferae; sori exindusiati; sporangia Setifera;
sporae alatae. —Type: PULLEN 1476, N.E. New
Guinea, Sepik Distr., Prince Alexander Range,
900 m (CANB).
Caudex short, creeping. Stipe 8cm long, red-
dish, glabrescent, basal scales 3 x 1.5 mm; base of
stipe to first large pinna 60 cm. Reduced pinnae 8
pairs or more, 5-6cm apart, all very small.
Lamina 35cm long; pinnae 10-12 pairs, all
opposite, basal pinnae not narrowed at their
bases; aerophores slender, to 1 mm long. Hairs on
rachis both sides 0.5mm long, on upper surface
brown. Largest sterile pinnae 14x2.5cm, fertile
to 10x 2.0cm; basiscopic base cordate and over-
lapping rachis, acroscopic truncate or slightly
cordate; apex acuminate; edges irregularly sinu-
ous to slightly crenate; costules 4.5 mm (sterile)
3.5mm (fertile) apart, at more than 60°; veins 6-7
pairs, 32 pairs anastomosing, one vein to short
sinus-membrane; lower surface of costae minutely
hairy, rest of lower surface glabrous, glands
present on costules; on upper surface base of
costae covered with brown hairs as rachis, above
base spare short hairs, rest of upper surface
glabrous. Sori on distal veins medial, on lower
ones supramedial, those on veins from adjacent
costules often coalescent; no indusia; sporangia
with 4-6 setae; spores with a + continuous trans-
lucent wing and a few irregular other wings.
Distr. Malesia: Papua New Guinea (Sepik).
(v.A.v.R.)
98. Sphaerostephanos _ pterosporus
1981]
THELYPTERIDACEAE (Holttum)
487
HOLTTUM, comb. nov. — Dryopteris pterospora
v.A.v.R. Bull. Jard. Bot. Btzg III, 2 (1920) 148. —
Thelypteris pterospora (v.A.v.R.) REED, Phy-
tologia 17 (1968) 307.—Type: BRooKs 447,
Sumatra, Benkoelen, Tambang Sawah (BO; BM).
Dryopteris bungoensis C. CHR. Dansk Bot. Ark.
9, n. 3 (1937) 59. — Type: BROOKS 10, April 1909,
Sarawak, Bungo Range (BM).
KEYerO THE VARTIEERIES
1. Pinnae lobed at least 1/3 towards costa.
2. Pinnae lobed less than 1/2 towards costa; basal
pair of veins often anastomosing
a. var. pterosporus
2. Pinnae lobed 3/5—2/3 towards costa; basal veins
rarely anastomosing . b. var. altilobus
1. Pinnae crenate . ¢. var. crenatus
a. var. pterosporus
Caudex short, erect. Stipe 3-10 cm long, basal
scales 8 X 1 mm; base of stipe to first normal pinna
20-30 cm (sterile fronds) 30-45 cm (fertile fronds).
Reduced pinnae 6-8 pairs, all less than 2 mm long.
Lamina c. 40cm long; pinnae to 18 pairs, basal
pinnae narrowed to a width of 6-8 mm or less at
base; aerophores less than | mm long. Hairs on
lower surface of rachis of sterile fronds thick,
brown, curved, to | mm long, on fertile paler and
shorter; hairs on upper surfaces as on lower sur-
face of sterile. Largest sterile pinnae to 12 x2cm;
base truncate; apex rather abruptly caudate-acu-
minate; edges lobed less than 1/2 towards costa,
lobes falcate with forward-pointing tips; costules
4.5-5 mm apart, at 60° to costa; veins 5-7 pairs, all
very oblique, basal pair uniting to form an excur-
rent vein to sinus or touching just below sinus,
next acroscopic vein to side of sinus-membrane or
to margin; hairs on lower surface of costae 0.3 mm
long, pale, closely appressed, on costules and
veins similar, sometimes also a few glands, on
surface between veins some fine appressed hairs
may be present but not glands; hairs on upper
surface of costae 0.5 mm long with some to | mm,
on costules and veins short and sparse, sometimes
a few appressed hairs between veins. Fertile pin-
nae to 7X 1.4cm, lobed a little more deeply than
sterile; costules 3.5mm apart; basal veins more
often connivent at the sinus without joining. Sori
medial; indusia firm with short hairs; sporangia
bearing glands; spores with many small separate
wings.
Distr. Malesia: Southern Malaya (several
localities), S. Sumatra, W. Sarawak, in forest at
9-700 m.
b. var. altilobus HOLTTUM, var. nov.
A typo speciei differt: pinnis maximis sterilibus
usque 14X2.6cm, fertilibus 10 1.8 cm, 3/5-2/3
costam versus lobatis; venis infimis plerisque ad
sinum conniventibus, raro anastomosantibus. —
Type: MOLESWORTH ALLEN 2736, Perak,
Maxwell’s Hill, 730m, in forest, “deep green
fronds, brown rachis, quite distinct” (K). Also
from Taiping Hills: KUNSTLER 6345, at 750-
900 m, “‘pinnae dark green”.
Distr. Malesia: Malaya.
c. var. crenatus HOLTTUM, var. nov.
A typo speciei differt: pinnis crenatis; venis
1 : =
l5-jugatis anastomosantibus, vena sequente
acroscopica ad membranam sinus terminata;
pinnis fertilibus usque 101.5 cm.—Type:
JERMY 13870, Sarawak, G. Mulu National Park,
by Tapin River (BM). Also M. HoTTa 13809,
Brunei, along Sungei Lacquan, 50-300 m (L).
Distr. Malesia: Borneo (Sarawak; Brunei).
99. Sphaerostephanos sagittifolius (BL.) HOLT-
TUM, comb. nov.— Aspidium sagittifolium BL.
Enum. Pl. Jav. (1828) 153; MIQUEL, Ann. Mus.
Bot. Lugd.-Bat. 4 (1869) 150.— Nephrodium
sagittifolium (BL.) Moore, Ind. Fil. (1858) 103;
RACIB. FI. Btzg 1 (1898) 191. — Dryopteris sagit-
tifolia (BL.) O. KTZE, Rev. Gen. Pl. 2 (1891) 813;
v.A.v.R. Handb. (1908) 222; BACKER & POSTH.
Varenfl. Java (1939) 52.— Thelypteris sagittifolia
(BL.) REED, Phytologia 17 (1968) 311.— Type:
BLUME, Java (L, n. 908, 336-24).
Caudex short, massive, decumbent or suberect:
young fronds covered with slime. Stipe 15cm
long; scales (fide RACIBORSKI) 4 X 3 mm; base of
stipe to first large pinna 50-80 cm. Reduced pinnae
more than 30 pairs, 1.5-2 cm apart, deflexed, up-
per ones 2.5-3.0cm long with basal auricle to
1.5cm, apex acuminate, edges crenate: lowest
lcm long. Lamina 100cm or more long; several
pairs lower pinnae with elongate basal acroscopic
auricle in which veins are forked; aerophores
thick but hardly 1 mm long. Rachis covered with
thick spreading hairs more than | mm long, both
sides. Largest pinnae to 20 2.3cm; base trun-
cate, basal acroscopic lobe somewhat elongate:
apex caudate-acuminate; edges lobed 2/5 to costa,
lobes with falcate tips; costules 5-5.5 mm apart at
more than 60°; veins to 10 pairs, pale and prom-
inent on lower surface, 1:2 pairs anastomosing,
2-23 pairs to sides of sinus-membrane; hairs on
lower surface of costae stiff, spreading, of varying
length to 1 mm, on costules similar but sparse, a
few glands on costules and veins; hairs on upper
surface of costae 1mm long, scattered similar
hairs on costules and veins, no others. Sori
medial, basal ones divergent; indusia small with
stiff hairs 0.3mm long; sporangia bearing glands
or setae; spores with many small wings.
Distr. Malesia: Sumatra, Java, and Lesser
Sunda Is. (Lombok).
Ecol. In forest at 1000-1500 m.
Note. v.A.v.R. (Handb. Suppl. 182) reported
specimens from New Guinea; I did not find these
in Herb. Bog.
488
FLORA MALESIANA
[ser. II, vol. P?
100. Sphaerostephanos foliolosus HOLTTUM, sp.
nov.— Aspidium hirsutum and A. cucullatum
sensu CHRIST, Ann Jard. Bot. Btzg 15 (1898) 134,
quoad Sarasin 745 tantum.
S. polyoti affinis, differt: pinnis c. 2/5 costam
versus lobatis; venis inferioribus 23-3-jugatis vel
anastomosantibus vel ad membranam sinus con-
niventibus; aerophoris usque 1mm longis; pinnis
redactis superioribus minoribus.— Type: P. & F.
SARASIN 745, N. Celebes, G. Matinang 350m
(BAS).
Stipe to 15cm long; scales broad, thin; base of
stipe to first large pinna 60cm. Reduced pinnae c.
30 pairs, 1-1.5 cm apart, upper ones to 12x 6mm,
lowest 4-8mm long, shape as in S. polyotis,
covered with mucilage when young. Lamina 65-
70cm long; pinnae 30 pairs, lower ones not nar-
rowed at base; aerophores slender, to 1 mm long.
Lower surface of rachis densely short-hairy
(sometimes with longer hairs?); brown hairs to
1mm long on upper surface. Largest pinnae 15 x
1.8cm, base subtruncate, symmetric (more
cuneate and less symmetric on upper pinnae);
apex gradually attenuate, not caudate; edges lobed
c. 2/5 to costa, lobes falcate with obtusely pointed
tips; costules 3mm apart, at 60° or more; veins
8-10 pairs, 13 pairs anastomosing, 1+-2 pairs pass-
ing to sides of sinus-membrane; hairs on lower
surface of costae short, dense, mixed with long
ones on sterile fronds, sparse hairs with glands on
costules and veins, short erect hairs between
veins; upper surface of costae bearing pale hairs
1 mm long, rest of surface + closely covered with
slender appressed hairs. Sori medial; indusia
small, pilose; sporangia usually with 1-2 glands,
sometimes a seta.
Distr. Malesia: N. Celebes, Moluccas (Talaud
Islands: LAM 2786, 2788, on river bank at 80-
350 m).
Note. LAM’s sterile specimens differ from the
type (which is fertile) in the presence of hairs
1 mm long on lower surface of rachis and costae.
101. Sphaerostephanos
nov.
Pinnae redactae 6—7-jugatae, superiores 12mm
longae; lamina 45cm longa; pinnae maximae
9.5 1.8cm, 3/4 costam versus lobatae; costae,
costulae venaeque subtus glandulosae, pagina in-
ter venas pilosa; pagina superior pilis appressis
vestita; indusia glandulis et pilis brevibus
praedita; sporangia nec glandulis nec setis
praedita.— Type: A. C. JERMy 5248, N.E. New
Guinea, Western Highlands, Keglsugl (BM).
Caudex short, creeping. Stipe 15—20cm long,
minutely hairy, basal scales broad, thin; base of
stipe to first large pinna 45-50 cm. Reduced pinnae
6-7 pairs, 34cm apart, lowest 5-7 mm long, up-
permost 12mm long, somewhat deflexed, nar-
rowly triangular, with auricle 6-8mm long on
acroscopic base. Lamina 45 cm long; pinnae to 24
alpinus HOLTTUM, sp.
pairs; basal pinnae not or little narrowed at their
bases; aerophores thick, hardly 1 mm long. Hairs
on lower surface of rachis dense, short, spread-
ing, on upper surface | mm long. Largest pinnae
9.5cm long, 1.8cm wide above slightly dilated
base; apex acuminate with cauda 1.5cm; edges
lobed almost or quite 3/4 to costa, lobes slightly
falcate; costules 3.5-4mm apart, their bases al-
most at right angles to costa; veins 9 pairs, at a
wide angle, pale and prominent both sides, basal
pair anastomosing, next pair usually both to mar-
gin; lower surface of costae densely covered with
short spreading hairs, hairs less dense, with
glands, on costules, surface between veins with
short erect hairs; hairs | mm long on upper sur-
face of costae, similar hairs scattered on costules
and veins, whole upper surface covered with ap-
pressed hairs. Sori medial; indusia bearing glands
and short hairs; sporangia with neither glands nor
setae; spores with many small wings.
Distr. Malesia: Papua New Guinea.
Ecol. Type locality at 2500-2750m, and Mt
Wilhelm at 3300-3500 m (NAKAIKE 274).
102. Sphaerostephanos _ polisianus HOLTTUM,
Kalikasan 4 (1975) 65.— Type: M. G. PRICE 431,
Luzon, Mountain Prov. Mt Polis (CAHUP; K).
Caudex creeping, 5-6mm diameter, bearing a
close series of fronds. Stipe 4-5 cm long, minutely
hairy, basal scales short and firm; base of stipe to
first large pinna 25-30 cm. Reduced pinnae 34cm
apart, all very small. Lamina to 28 cm long; pin-
nae 14 pairs, lowest narrowed at their bases;
aerophores 1mm long. Lower surface of rachis
covered with very short erect reddish hairs, upper
surface with thick curved dark red hairs 0.5 mm
long. Largest pinnae 6.5 x 1.3cm; base subtrun-
cate; apex short-acuminate; edges lobed > way to
costa, lobes oblique with forward-pointing obtuse
tips; costules 3-3.5mm apart at 60°; veins 6-7
pairs, slender and prominent on lower surface,
basal pair anastomosing, next acroscopic vein to
side of short sinus-membrane; lower surface of
costae covered with hairs 0.1 mm long or a little
more, antrorse at least distally, few similar hairs
on costules, glands on costules and veins; hairs on
upper surface of costae 0.3 mm long, rest of upper
surface glabrous. Sori medial; indusia small,
glabrous; sporangia sometimes with a gland.
Distr. Malesia: Philippines (Luzon);
known from the type.
Note. This should perhaps be regarded as a
form of S. lobatus.
only
103 Sphaerostephanos hoalensis HOLTTUM, sp.
nov. — Dryopteris ceramica ROSENST. MS, non
v.A.v.R.
Pinnae redactae c. 13-jugatae, superiores 15 x
8mm, infimae 10mm longae; aerophora elon-
gata; pinnae maximae 11 X2cm, dimidio costam
versus lobatae; costae subtus pilis erectis usque
1981]
THELYPTERIDACEAE (Holttum)
489
Imm longis vestitae, costulae pilis brevioribus
glandulisque praeditae; pagina superior pilis ap-
pressis vestita; indusia_ pilifera; sporangia
glandulifera. — Type: STRESEMANN 110, Ceram,
Central Mts, G. Hoale, 1000 m (L).
Stipe lacking. Reduced pinnae 2.5 cm apart, c.
13 pairs, one near the base 10 mm long, uppermost
15Sx8&mm, deflexed, slightly auricled on acros-
copic base, edges crenate, apex obtuse to roun-
ded. Lamina c. 60cm long; lowest pinnae not
narrowed at base; aerophores elongate. Lower
surface of rachis covered with erect hairs 0.2-
0.3 mm long and some longer ones, hairs on upper
surface | mm long. Largest pinnae 11 x 2. cm; base
truncate; apex abruptly short-acuminate; edges
lobed c. 3 way to costa, lobes oblique, slightly
falcate; costules 3mm apart, at more than 60°;
veins 9 pairs, 1> pairs anastomosing, next acros-
copic vein to sinus-membrane; lower surface of
costae hairy as rachis, costules and veins less
densely covered with shorter hairs and some
glands, surface between veins bearing short erect
hairs; hairs on upper surface of costae 1 mm long,
shorter ones on costules, appressed hairs all over
surface. Sori small, medial to supramedial, lower
ones a little divergent; indusia with many rather
long hairs; sporangia bearing | or 2 glands.
Distr. Malesia: Moluccas (Central Ceram, 2
collections, second one n. 57).
Ecol. In the mountains, 1000 m.
104. Sphaerostephanos larutensis (BEDD.) C. CHR.
Ind. Fil. Suppl. III (1934) 172.— Nephrodium
larutense BEDD. Handb. Suppl. (1892) 73.—
Mesochlaena larutensis (BEDD.) v.A.v.R. Handb.
(1908) 232, incl. var. borneensis. — Cyclosorus
larutensis (BEDD.) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 245; HOLTTUM, Rev. FI. Mal.
(1955) 284, f. 165.—Thelypteris larutensis
(BEDD.) REED, Phytologia 17 (1968) 286. — Lec-
totype (here selected): KUNSTLER 2398, Perak,
Larut (K).
Mesochlaena sumatrensis v.A.v.R. Bull. Jard.
Bot. Btzg III, 2 (1920) 160.—S. sumatrensis
(v.A.v.R.) C. CHR. Ind. Fil. Suppl. III (1934) 172. —
Type: BROOKS 391, Sumatra, Benkoelen, Lebong
Tandai (BO; BM). — Fig. 12d-e.
Caudex short, creeping. Stipe 20-30cm long,
above base covered with spreading hairs to 1 mm
long; base of stipe to first large pinna 40-70 cm.
Reduced pinnae 6-12 pairs, lower ones less than
lcm long, upper ones c. 1.5 x 1.2 cm, spreading,
triangular, edges crenate, apex acute; an inter-
mediate pair sometimes present. Lamina 50-
70cm long; apex pinna-like; pinnae c. 12 pairs,
well spaced; lower pinnae narrowed towards their
bases; middle pinnae with truncate bases, upper
ones broadly cuneate; aerophores not elongate.
Rachis covered throughout with slender light
brown hairs !-1.5mm long. Largest pinnae 20-
30 cm long, 2.5—-3.8 cm wide, acuminate with cauda
1-3 cm long; edges lobed 1/4~1/3 to costa, lobes
falcate with forward-pointing obtuse tips; costules
4.5-6 mm apart; veins to 11 pairs, 13-2 pairs anas-
tomosing, next 2 pairs to sides of long sinus-
membrane; lower surface of costae covered with
erect hairs 0.5-1mm long, shorter hairs on cos-
tules, veins and between veins, some glands with
the hairs on costules and veins; upper surface of
costae covered with fine pale antrorse hairs, on
costules and veins much shorter hairs with scat-
tered long ones, surface between veins glabrous or
with suberect hairs. Sori medial, about twice as
long as wide; indusia large, thin, bearing slender
hairs and glands; sporangia bearing neither glands
nor setae on body, hair on stalk of several cells,
terminal one acicular and thick-walled; spores
with many small wings.
Distr. Malesia: Malaya, Sumatra, Borneo.
Ecol. In forest at 50-1000 m.
Note. v.A.v.R. distinguished var. borneensis
which has many glands and no hairs on indusia.
His M. sumatrensis was based on a rather small
specimen. Glands are apparently sometimes lack-
ing on lower surface of costules.
105. Sphaerostephanos loherianus (CHRIST)
HOLTTUM, Kalikasan 4 (1975) 61.— Aspidium
loherianum CHRIST, Bull. Herb. Boiss. 6 (1893)
191.— Dryopteris loheriana (CHRIST) C. CHR.
Ind) Fil 1@905), 275: CurismePhilipa ws Scie 2
(1907) Bot. 207; v.A.v.R. Handb. (1908) 221,
820. — Cyclosorus loherianus (CHRIST) COPEL.
Gen. Fil. (1947) 142; Fern FI. Philip. (1960) 342. —
Thelypteris loheriana (CHRIST) REED, Phytologia
17 (1968) 289. — Type: LOHER 900, Luzon, Mon-
talban, Oct. 1890 (P; K).
Caudex creeping, sometimes with well-spaced
fronds. Stipe 5-30cm long, reddish, glabrescent;
base of stipe to first large pinna 40-50 cm.
Reduced pinnae 5-15 pairs, defiexed, strongly
auricled, uppermost 6-10mm long, lowest 2-
5mm. Lamina to 60cm or more long; pinnae well
spaced, several lower pairs narrowed towards
their bases; aerophores slightly swollen. Lower
surface of rachis densely covered with short erect
hairs, those on upper surface to 1mm _ long.
Suprabasal pinnae of type 11 x 1.8 cm, of largest
specimen 303.2cm; basal lobes somewhat
reduced; apex acuminate, sometimes with cauda
to 2cm; edges lobed to 2mm from costa, lobes
slightly falcate, narrowed gradually from base;
costules 44.5 mm apart, almost at right angles to
costa; veins of type to 14 pairs, of largest speci-
men 24 pairs, basal veins near base of pinna often
anastomosing to form a short excurrent vein,
those near apex of pinna connivent at the sinus
without uniting; lower surface of costae densely
covered with erect hairs 0.3-0.5 mm long, on some
specimens to | mm, on costules and veins shorter
hairs and some glands, short erect hairs between
veins; hairs on upper surface of costae to 1mm
490
long, scattered similar hairs on costules and veins,
between veins a variable number of short hairs.
Sori all close to margin; indusia small, covered
with short hairs and sometimes a few glands;
sporangia setiferous, rarely with a gland.
Distr. Malesia: Philippines (N. Luzon).
Ecol. In mountain forest, 1500-2000 m; smaller
specimens apparently from streamsides in open
places.
106. Sphaerostephanos erectus (COPEL.) HOLT-
TUM, Kalikasan 4 (1975) 64. — Cyclosorus erectus
COPEL. Philip. J. Sci. 81 (1952) 30, t. 22; Fern FI.
Philip. (1960) 346. — Thelypteris erecta (COPEL.)
REED, Phytologia 17 (1968) 275. — Type: RAMOS
BS 41550, Leyte, Cabalian (US; B).
Caudex slender, erect, to 100cm tall. Stipe
10cm long, reddish, minutely hairy; base of stipe
to first large pinna 45cm. Reduced pinnae to at
least 6 pairs, uppermost 5mm long. Lamina to
50cm long; pinnae 15 pairs, lower ones narrowed
on basiscopic side near base; apex of frond nar-
rowly triangular and deeply lobed with abrupt
transition to pinnae; aerophores not elongate.
Hairs on rachis, both sides, to | mm long, thick,
brown. Suprabasal pinnae to 14x2.8cm; base
truncate; apex rather abruptly caudate-acuminate;
edges lobed 3/5-2/3 to costa, lobes falcate; cos-
tules 4-5.5mm apart, at more than 60°; veins
10-12 pairs, prominent on lower surface, basal
pair anastomosing, second pair touching sides of
sinus-membrane; hairs on lower surface of costae
spreading, of varied length to 1mm, on costules
shorter with glands; hairs on upper surface of
costae 1mm long, on costules short, sometimes
with a few glands, rest of upper surface glabrous.
Sori medial; indusium firm, large, short-hairy;
sporangia bearing glands.
Distr. Malesia: Philippines (Leyte, 2 collec-
tions; Biliran).
Ecol. In mossy forest at 1000 m (M. G. PRICE
3278).
Note. M. G. PRICE informs me that SULIT
PNH 20306 from Biliran, which he identifies with
this species, has a few glands on upper surface of
costules; I have seen none in this position on the
specimens from Leyte.
107. Sphaerostephanos stresemannii HOLTTUM,
sp. nov.
Basis frondis imperfecta; pinnae infimae 12 cm
longae, medio 3.2 cm latae, basin versus angus-
tatae; pinnae suprabasales usque 10X2.7cm,
oppositae, 3/4 costam versus lobatae; costae sub-
tus pilis erectis usque 0.5 mm longis vestitae, cos-
tulae venaeque pilis sparsis glandulisque praedi-
tae; pagina superior inter venas pilis brevibus
suberectis instructa; sori exindusiati; sporangia
setifera.— Type: STRESEMANN 416, Moluccas,
Buru, G. Fogha 1400 m (L).
Caudex and stipe lacking. Reduced pinnae at
FLORA MALESIANA
[ser. II, vol. 1°
least 2 pairs, 3-4mm long; above them are an
intermediate unequal pair 1.0 and 2.0cm long.
Lamina 42cm long; apex narrowly deltoid with
gradual transition to upper pinnae; pinnae 15
pairs, almost opposite; basal pinnae largest, 12 x
3.2cm, widest in middle and narrowed to base
both sides, basiscopic lobes longer than acros-
copic; aerophores swollen, not elongate. Hairs on
lower surface of rachis dense, short, erect, with
scattered longer ones to 0.7mm, hairs on upper,
surface longer and appressed. Suprabasal pinnae
to 10x3.0cm; base truncate; apex short-acu-
minate; edges lobed fully 3/4 to costa, lobes
hardly falcate, slightly tapered; costules 4.5mm
apart at more than 60°; veins 10 pairs, slender,
pale both sides, | pair anastomosing, next acros-
copic veins to side of short sinus-membrane; hairs
on lower surface of costae as on rachis, on cos-
tules and veins more sparse and shorter spreading
hairs with glands; short erect hairs on surface
between veins; hairs on upper surface of costae
more than 0.5 mm long, similar hairs scattered on
costules and veins, between veins short suberect
hairs. Sori medial; indusia lacking; sporangia
copiously setose; spores with many small wings.
Distr. Malesia: Moluccas (Buru), 1400 m.
108. Sphaerostephanos simplicifolius (J. SM. ex
HoOoK.) HOLTTUM, Kalikasan 4 (1975) 61.—
Aspidium simplicifolium J. SM, ex HOoK. Ic. Pl.
10 (1854) t. 919. — Polypodium simplicifolium (J.
SM.) HOOK. Sp. Fil. 5 (1863) 2, excl. SEEMANN
736 (Fiji). — Dryopteris canescens var. C. CHR.
Ind. Fil. (1905) 256. — Dryopteris simplicifolia (J.
SM.) CHRIST, Philip. J. Sci. 2 (1907) Bot. 206. —
Phegopteris simplicifolia (J. SM.) v.A.v.R.
Handb. (1908) 500. — Cyclosorus simplicifolius (J.
SM.) COPEL. Gen. Fil. (1947) 143; Fern FI. Philip.
(1960) 371.— Thelypteris simplicifolia (J. SM.)
REED, Phytologia 17 (1968) 314. — Type: CUM-
ING 315, Samar (K; E, G, L).
Caudex short-creeping; stipe to 20cm long,
pale; basal scales 7 x | mm; base of stipe to first
normal pinna 40-50cm; reduced pinnae to 6-7
pairs, lowest 3mm long, uppermost 10 mm, trian-
gular with asymmetric base. Lamina to 50cm
long; apex pinna-like, pinnae to 8 pairs, lowest
somewhat narrowed at base; aerophores not
elongate. Hairs on both sides of rachis pale,
longest more than | mm rather sparse, with shor-
ter ones. Largest pinnae to 164cm; base
broadly cuneate, not auricled; apex abruptly
short-acuminate; edges crenate to subentire; cos-
tules 4-4.5 mm apart, at 60°; veins to 10 pairs, pale
and prominent on lower surface, almost all anas-
tomosing to produce a zig-zag excurrent vein;
copious stiff short erect hairs on lower surface of
costae with scattered long ones, on costules and
veins fewer short hairs and also glands; hairs on
upper surface of costae and costules short with
scattered long ones. Sori near apices of veins,
1981]
basal ones on veins from adjacent costules some-
times confluent; indusia with many short hairs;
sporangia with many glands; spores with many
broad translucent wings which anastomose.
Distr. Malesia: Philippines (Samar,
Biliran).
Note. The original description was based on a
partially fertile small plant with apical lamina
much larger than the pinnae. The above descrip-
tion is based on fully grown plants from other
collections. Both the type and larger specimens
have glands on sporangia, not setae as stated by
COPELAND. In Sp. Fil. vol. S HOOKER misidentified
a specimen of the exindusiate Pronephrium bec-
carianum (from Fiji) with the present species, for
which reason he transferred the latter to Poly-
podium.
Leyte,
109. Sphaerostephanos spenceri (CHRIST) HOLT-
TUM, Kalikasan 4 (1975) 58. — Dryopteris spenceri
CHRIST, Philip. J. Sci. 2 (1907) Bot. 206; v.A.v.R.
Handb. Suppl. (1917) 188.— Phegopteris spen-
ceri (CHRIST) v.A.v.R. Handb. (1908) 508. —
Cyclosorus spenceri (CHRIST) COPEL. Gen. Fil.
(1947) 143; Fern Fl. Philip. (1960) 350. — Thelyp-
teris spenceri (CHRIST) REED, Phytologia 17
(1968) 314. Type: COPELAND 1464, Mindanao,
Davao Distr., Todaya (P; B, US).
Caudex short, suberect or creeping. Stipe S-
10cm long, densely short-hairy; base of stipe to
first large pinna 35 cm. Reduced pinnae to at least
10 pairs, lowest S-10 mm long, uppermost to 2cm
long, 1.5 cm wide above auricled acroscopic base,
triangular, crenate. Lamina 45 cm long; apex nar-
rowly triangular, deeply lobed at base; pinnae to
at least 10 pairs; basal pinnae not narrowed at
base; aerophores not elongate. Hairs on both sides
of rachis dense, c. 0.3mm long, with some less
abundant | mm or more long. Largest pinnae 8.5 x
2.0cm; base broadly rounded to subcordate on
basiscopic side, truncate to broadly cuneate on
acroscopic; apex short-acuminate; edges crenate
to a depth of 1mm or a little more; costules
4-5 mm apart, at 60°; veins 7-8 pairs, prominent
on lower surface, 2-23 pairs anastomosing to form
a Zig-zag excurrent vein, 2-2) pairs passing to
sides of sinus-membrane; lower surface of costae
hairy as rachis, hairs on costules shorter with
some glands, short erect hairs on surface between
veins; hairs on upper surface of costae 0.3-0.4 mm
long, on costules shorter. Sori medial, distinctly
elongate; indusia with short hairs; sporangia bear-
ing glands.
Distr. Malesia: Philippines (E. & S.E. Min-
danao).
Note. CHRIST failed to see the indusia and
thus at first misled v.A.v.R. The short stipe and
deeply lobed apical lamina appear to be distinctive
and are shown by M. G. PRICE 2790 from Agusan
del Sur Province, which is a rather small sterile
plant differing from the type in scattered long
THELYPTERIDACEAE (Holttum)
491
hairs on the upper surface of costules and veins
and in abundant glands on lower surface of cos-
tules and veins.
110. Sphaerostephanos mengenianus HOLTTUM,
sp. nov.
Caudex 40cm longus, erectus; pinnae omnes
oppositae, basi late cuneatae; pinnae redactae 8-
jugatae, maximae 22cm; pinnae normales
13.5X2.0cm, crenatae; pili paginae inferioris
omnes breves; costulae subtus glanduliferae; in-
dusia_ pilifera; sporangia glandulifera? (non
setifera). —Type: STEVENS & LELEAN LAE
58680, E. New Britain, in Castanopsis forest,
900 m (K; L).
Caudex erect, 50cm tall (collectors). Stipe
15cm long, minutely hairy, dull reddish; base of
stipe to first large pinna 60cm. Reduced pinnae
opposite, S-6cm apart, lowest 7mm long, up-
permost 22cm, almost symmetrically rhombic
with basal angle more than 90°, apical angle less
than 90°, subentire; one pair intermediate pinnae
4cm long also present. Lamina 55cm long; apex
not pinna-like; pinnae 20 pairs, all opposite; basal
pinnae not narrowed at base; aerophores not
elongate. Hairs on lower surface of rachis 0.2 mm
long, slender, appressed, on upper surface thicker,
antrorsely curved, 0.5mm long. Largest pinnae
13.5 2.0cm:; base broadly cuneate and slightly
dilated both sides; apex acuminate, not caudate;
edges crenate to a depth of 1.5 mm or little more;
costules 4.5-5 mm apart, at less than 60°; veins 7-8
pairs, 2-23 pairs anastomosing to form a zig-zag
excurrent vein, 1>2 pairs passing to rather long
sinus-membrane; lower surface of costae covered
with slender appressed hairs 0.2 mm long, similar
sparse hairs with glands on costules, surface be-
tween veins glabrous, eglandular; hairs on upper
surface of costae 0.5 mm long, rest of upper sur-
face glabrous. Sori medial; indusia rather small,
thin, with short hairs; sporangia probably with
glands, not setiferous.
Distr. Malesia: Papua New Guinea (New Bri-
tain), only the type.
Ecol. In Castanopsis forest, 900 m.
111. Sphaerostephanos confertus (BRAUSE)
HoOLTTUM, Webbia 30 (1976) 194. — Dryopteris
conferta BRAUSE, Bot. Jahrb. 49 (1912) 22, f. IF;
v.A.v.R. Handb. Suppl. (1917) 175.— Cyclo-
sorus confertus (BRAUSE) COPEL. Gen. Fil. (1947)
142; HoLtTuM & Roy, Blumea 13 (1965) 134. —
Thelypteris conferta (BRAUSE) REED, Phyto-
logia 17 (1968) 268.—Type: SCHLECHTER
17864, N.E. New Guinea, Rani Mts 700m (B; K,
EYP Ue):
Dryopteris terrestris COPEL. Univ. Cal. Publ.
Bot. 18 (1942) 221.—Cyclosorus terrestris
(COPEL.) COPEL. Gen. Fil. (1947) 143; Philip. J.
Sci. 78 (1951) 454, pl. 34. — Thelypteris terrestris
(COPEL.) REED, Phytologia 17 (1968) 319.—
492
Type: BRASS 13660, New Guinea, Idenburg
River, in forest at 700 m (MICH; BO, L).
Caudex short, creeping, branched, apices of
branches with tufts of fronds. Stipe 5-15 cm long,
minutely hairy; basal scales narrow, firm; base of
stipe to first large pinna 15-30 cm. Reduced pinnae
5-6 pairs, uppermost 6-12mm long and wide,
broadly triangular, lowest 3-5 mm long. Lamina
30-40 cm long; apex deltoid, deeply lobed; pinnae
to 15 pairs, basal ones slightly narrowed at base
which in some cases is slightly auricled; aero-
phores not elongate. Lower surface of rachis
covered with short erect pale hairs; upper surface
with long curved brown hairs and shorter ones in
the groove. Largest pinnae 5-10cm long, 1.5-
2.0 cm wide, in some cases widest 1/3 from apex;
base truncate, in some cases a little dilated both
sides or only on acroscopic side; apex abruptly
short-acuminate, with or without a short cauda;
edges crenate throughout (near apex only in small
pinnae) to a depth of 1-2 mm; costules commonly
4mm apart, at 60°; veins 5-6 pairs, 2 pairs anas-
tomosing to form a zig-zag excurrent vein, 1-2
pairs to rather long sinus-membrane; lower sur-
face of costae covered with antrorse (not closely
appressed) hairs 0.1-0.3 mm long, hairs on cos-
tules and veins sparse, sometimes with a few
glands, short erect hairs on surface between veins;
hairs on upper surface of costae 0.3-0.4 mm long,
a few minute hairs on costules, between veins a
variable number of short appressed hairs. Sori
medial; indusia with many short stiff hairs;
sporangia bearing several setae, sometimes also a
gland; spores minutely papillose.
Distr. Malesia: New Guinea and Waigeo; at
low altitudes in forest.
Note. Many collections from eastern New
Guinea, none from west except Waigeo.
112. Sphaerostephanos multiauriculatus (COPEL.)
HOLTTUM, comb. nov.— Dryopteris multiauri-
culata COPEL. Univ. Cal. Publ. Bot. 18 (1942)
221.—Cyclosorus multiauriculatus (COPEL.)
COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78
(1951) 453, pl. 32.— Thelypteris multiauriculata
(COPEL.) REED, Phytologia 17 (1968) 294. —
Type: BRASS 12850, New Guinea, Idenburg
River, 1150 m, in gullies in forest (MICH; BM, L).
Caudex erect, to 40cm tall, 1.5cm diameter.
Stipe 5-10 cm long, reddish, minutely hairy, with
many narrow scales 10mm long. Lamina 150 cm
long, somewhat dimorphous; pinnae to 45 pairs,
lower 12-15 pairs gradually reduced, lowest 1 cm
long, suprabasal ones spreading, broadly trian-
gular with symmetric base; aerophores elongate
barely to 1 mm. Lower surface of rachis of fertile
frond densely covered with short appressed hairs,
hairs on upper surface similar but with some thick
brown hairs 1.5mm long; similar long thick hairs
sometimes present on lower surface of sterile
fronds. Largest sterile pinnae to 15 x 1.8cm; base
FLORA MALESIANA
(ser. II, vol. 1°
subtruncate; apex short-acuminate, not caudate;
edges crenate to a depth of 1mm; costules 3 mm
apart, at 60°; veins to 7 pairs, slender and prom-
inent both sides, 3-32 pairs anastomosing to form
a zig-zag excurrent vein, | pair to very short
sinus-membrane; lower surface of costae, costules
and veins covered with appressed hairs 0.1-
0.3mm long, a few similar hairs on surface be-
tween veins, glands lacking; upper surface of
costae covered with hairs less than 0.5 mm long,
rest of surface glabrous. Fertile pinnae to 9x
1.0cm; sori medial, often spreading a little along
veins; no indusia; sporangia sometimes bearing a
few glands, not setae.
Distr. Malesia: Eastern New Guinea at 750-
1220 m.
113. Sphaerostephanos stipellatus (BL.) HOLT-
TUM, comb. nov.— Aspidium stipellatum BL.
Enum. Pl. Jav. (1828) 152; MIQUEL, Ann. Mus.
Bot. Lugd.-Bat. 4 (1869) 160.— Nephrodium
stipellatum (BL.) Moore, Ind. Fil. (1858) 105;
RACcIB. FI. Btzg 1 (1898) 188. — Dryopteris stipel-
lata (BL.) O. KTZE, Rev. Gen. Pl. 2 (1891) 813;
v.A.v.R. Handb. (1908) 227; BACKER & POSTH.
Varenfl. Java (1939) 49.— Thelypteris _ stipel-
lata (BL.) K. Iwats. Acta Phytotax. Geobot. 21
(1965) 168.—Type: BLUME, Java (L, n. 908,
337-123).
Caudex thick, short-creeping, apex and very
young fronds covered with slime. Stipe 10cm
long, glabrescent; scales very thin; base of stipe to
first large pinna 65cm. Reduced pinnae 2-3cm
apart, to 15 pairs or more, lowest 6mm long,
uppermost 12 mm long, deflexed, narrowly trian-
gular and strongly auricled. Lamina to 70 cm long;
pinnae to 30 pairs, lower ones not narrowed at
their bases; aerophores thick, to 2 mm long. Hairs
on lower surface of rachis of fertile fronds
0.3mm long, closely appressed, on sterile ones
sparse longer hairs also; on upper surface pale,
spreading, | mm long. Largest pinnae commonly
12x 1.8cm; base subtruncate; apex acuminate,
apical 1.5cm entire; edges lobed c. 1/3 to costa
(always less than 3), lobes of fertile fronds dis-
tinctly falcate; costules 3.5-4.5mm apart, at a
wide angle; veins 6-8 pairs, | pair anastomosing,
next acroscopic vein to side of short sinus-mem-
brane; hairs on lower surface of costae and cos-
tules short, closely appressed, with a few longer
ones distally on costae, few or none on veins and
surface between them, no glands; hairs on upper
surface of costae 1mm long, scattered similar
hairs on costules and veins. Sori medial; indusia
firm, with a few glands; sporangia bearing glands;
spores finely spinulose.
Distr. Malesia: Java, Sumatra.
Ecol. In forest at 1000-1700 m.
114.
HOLTTUM,
Sphaerostephanos _plurifolius (v.A.v.R.)
comb. nov.— Dryopteris plurifolia
1981]
THELYPTERIDACEAE (Holttum)
493
v.A.v.R. Bull. Jard. Bot. Btzg III, 5 (1922) 201. —
Thelypteris plurifolia (v.A.v.R.) REED, Phy-
tologia 17 (1968) 305.—Type: LORZING 6393,
Sumatra, Deli, Bandar-baroe, 800 m (BO).
Caudex erect, 20-30cm tall, apex and young
fronds covered with mucilage; stipe 10-l1Scm
long; base of stipe to first normal pinna 40-50 cm;
reduced pinnae c. 4cm apart, to 8 pairs, lowest
very small, upper ones 6mm long. Lamina 70cm
or more long; pinnae to 25 pairs, 3 or more pairs
of lower ones distinctly narrowed towards their
bases, basal ones much narrowed; aerophores to
2mm long. Hairs on lower surface of rachis very
short, erect, on upper surface 0.5 mm long, or to
1 mm on sterile fronds. Largest suprabasal pinnae
commonly 15 x2 cm (sterile to 25 x 3.2 cm): base
truncate; apex acuminate; edges lobed 2/3-3/4
towards costa, lobes falcate; costules 3.5-S mm
apart, at more than 60° to costa; veins 10-13 pairs,
basal pair anastomosing, next pair both passing
to margin or the acroscopic vein to sinus-mem-
brane; lower surface of costae, costules and veins
covered with short pale appressed hairs, without
glands, area between veins glabrous; hairs on upper
surface of costae short, with some longer ones,
similar long hairs scattered on costules and veins.
Sori medial; indusia large, firm, with a few hairs;
sporangia usually bearing glands.
Distr. Malesia: Sumatra.
Note. The type has distinctly dimorphous
fronds, but a specimen from Gunong Kerinci
(ROBINSON & KLOSS s.n. at BM) has fertile pinnae
to 3.2 cm wide.
115. Sphaerostephanos hendersonii HOLTTUM, sp.
nov.
Pinnae redactae c. 8-jugatae, usque 6mm lon-
gae; pinnae normales usque 14 Xx 2.3 cm, 3/5 cos-
tam versus lobatae, basi aerophoris longis praedi-
tae, subtus eglandulosae; venae 10-jugatae,
infimae solum anastomosantes; rachis, costae
costulaeque subtus pilis adpressis vestitae; sori
mediales; indusia parva, tenuia, sparsim pilosa;
sporangia glandulifera. —Type: HENDERSGN
23352, Malaya, Cameron Highlands, 1500m
(K; SING).
Caudex short, erect or suberect; stipe 10-15 cm
long; base of stipe to first normal pinna 40-50 cm;
reduced pinnae c. 4cm apart, c. 8 pairs, uppermost
6mm long, rest very small. Lamina excluding
reduced pinnae 70cm long; pinnae more than 20
pairs; basal pinnae narrowed towards their bases
which are 1.2 cm wide; aerophores to almost 2mm
long. Largest pinnae (fertile) 14x 2.3cm; base
truncate; apex abruptly short-acuminate with nar-
row entire cauda 1l-2cm long; edges lobed
to 4mm from costa; costules 4-5 mm apart, at
more than 60°; veins to 10 pairs, basal pair anas-
tomosing, next acroscopic vein passing to the
sinus-membrane, sometimes a basiscopic vein
also; lower surface of rachis bearing slender ap-
pressed pale hairs 0.2mm _ long (sometimes
sparse), costae and costules bearing many ap-
pressed hairs 0.3-0.4mm long, on and between
veins neither hairs nor glands; upper surface of
rachis bearing pale curved hairs 0.5 mm long, hairs
on costae to !mm or more, similar hairs rare on
costules and veins, surface between veins bearing
a variable number of appressed hairs. Sori medial,
basal ones not divergent; indusia small, thin, with
a few hairs; sporangia bearing glands.
Distr. Malesia: Malaya (Cameron Highlands,
3 collections); N. Sumatra at 1700 m.
Note. The Sumatran specimens (SURBECK
747, 1201) are somewhat smaller than those from
Malaya, with pinnae lobed a little less deeply, and
have a seta on some sporangia.
116. Sphaerostephanos posthumii HOLTTUM, sp.
nov.
Pinnae redactae c. 10-jugatae, usque 4 mm lon-
gae; pinnae normales usque 12.5 1.5 cm, basi
aerophoris 1mm longis praeditae, profunde loba-
tae; venae infimae prope sinum conniventes, vel
liberae vel anastomesantes; pagina inferior pin-
narum eglandulosa, pagina superior pilis adpressis
vestita; indusia parva, brevi-pilosa; sporangia
glandulifera. — Type: POSTHUMUS 3157, Flores,
near Sita, 600-700 m (BO).
Caudex not known; stipe 24cm long, basal
scales broad, thin, with rather long superficial
hairs; base of stipe to first normal pinna 80cm;
reduced pinnae c. 5cm apart, uppermost 4mm
long. Lamina 60cm long, excluding reduced pin-
nae; pinnae 20 pairs; lowest large pinnae much
narrowed towards their bases, several successive
pairs gradually less narrowed; aerophores 1mm
long. Rachis on both surfaces covered with pale
appressed hairs c. 0.6mm long. Largest pinnae
(fertile) 12.5x1.5cm, sessile; apex gradually
attenuate, not caudate; edges lobed to 1.5-2mm
from costa, lobes slightly falcate; costules 3-
35mm apart, at 60°; veins 10-12 pairs, basal pair
either both touching sides of a short sinus-mem-
brane or meeting just below it to form a short
excurrent vein; lower surface of costae and cos-
tules closely covered with pale appressed hairs,
between veins some shorter appressed hairs,
glands lacking; upper surface throughout covered
with pale appressed hairs 0.3-0.4 mm long. Sori a
little inframedial, basal ones not much divergent;
indusia rather small, thin, with many hairs 0.2 mm
long; sporangia often with a gland.
Distr. Malesia: Lesser Sunda Is. (Flores), only
known from type.
117. Sphaerostephanos inconspicuus (COPEL.)
HOLTTUM, comb. nov. — Dryopteris inconspicua
CoPEL. Philip. J. Sci. 12 (1917) Bot. 55; C. CHR.
Gard. Bull. Str. Settl. 7 (1934) 242. — Thelypteris
inconspicua (COPEL.) CHING, Bull. Fan Mem.
Inst. Biol. Bot. 10 (1941) 252. — Type: TOPPING
494
1543, Sabah, Mt Kinabalu, Kiau (isotype NY). —
Fig. 12n-p.
Caudex short, creeping. Stipe 4-8cm long,
minutely hairy; base of stipe to first large pinna
15-30cm. Reduced pinnae 4-7 pairs, all very
small, not opposite. Lamina 25-40 cm long; pin-
nae to 25 pairs, basal pinnae narrowed at base;
aerophores not elongate. Hairs on lower surface
of rachis variable, sometimes very few on fertile
fronds, on sterile often many, thick, curved,
0.5 mm or more long; hairs on upper surface 0.8-
1.0mm long. Largest pinnae 3.5x0.8 to 6.0
1.3cm; base subtruncate; basal acroscopic lobe
sometimes elongate; apex acuminate, sometimes
caudate; edges lobed to 1-1.5mm from costa,
lobes oblique; costules 3-3.5 mm apart, usually at
less than 60°; veins 4-5 pairs, slender, basal
acroscopic vein touching short sinus-membrane,
basal basiscopic vein to edge above base of sinus;
hairs on lower surface of costae 0.2-0.3 mm long,
pale, appressed, fewer on costules, sometimes
with rudimentary scales which have a glandular
tip, sessile spherical glands usually lacking; hairs
on upper surface of costae less than 0.5 mm long,
no others. Sori medial; indusia rather large with a
few short hairs or glabrous; sporangia bearing a
gland or a seta; spores not seen.
Distr. Malesia: Borneo; several localities, in
forest at 1200-1800 m.
118. Sphaerostephanos subulifolius (v.A.v.R.)
HOLTTUM, comb. nov.— Dryopteris subulifolia
v.A.v.R. Bull. Jard. Bot. Btzg II, 28 (1918)
22.— Thelypteris subulifolia CHING, Bull. Fan
Mem. Inst. Biol. Bot. 10 (1941) 254.— Type:
BROOKS 334 partim, Sumatra, Benkoelen, Lebong
Simpang (BO; BM).
Closely allied to S. inconspicuus, differing as
follows: reduced pinnae 2-3 pairs, all very small;
normal pinnae to 30 or more pairs; pinnae ab-
ruptly short-acuminate, lobed to 1 mm from costa;
hairs on lower surface of costae 0.5 mm long, not
closely appressed.
Distr. Malesia: Southern Sumatra, 3 collec-
tions.
Notes. The differences between this and S.
inconspicuus need checking from further collec-
tions from Sumatra. BROOKS 334 at BM includes
specimens of Coryphopteris hirsutipes (CLARKE)
HOLTTUM.
119. Sphaerostephanos mjobergii HOLTTUM, sp.
nov.
S. inconspicuo (Copel.) Holttum affinis, differt:
pinnis evolutis 27-jugatis, usque 3.6*1.0cm
metientibus, ad alam 0.5 mm latam lobatis; venis
infimis omnibus ad marginem supra basin sinuum
terminatis; costis subtus pilis patentibus
vestitis. — Type: E. MJOBERG s.n. Sarawak, Mt
Poi, 1370-1520 m (BM).
Caudex and stipe missing. Reduced pinnae all
FLORA MALESIANA
[ser. II, vol. 1°
very small, 2-3 pairs on specimen. Lamina 30cm
long; pinnae 27 pairs; basal pinnae apparently
narrowed on basiscopic side only; aerophores not
elongate. Hairs on lower surface of rachis 0.2-
0.3 mm long, copious. Largest pinnae 3.6 X 1.0 cm;
base symmetric, subtruncate; apex rather abruptly
acute; edges lobed to 0.5mm from costa, lobes
hardly falcate; costules to 2.5 mm apart; veins 3-4
pairs, basal ones both passing to margin above
base of sinus: lower surface of costae covered
with somewhat antrorse hairs 0.2-0.3mm long,
longer hairs scattered on costules; hairs on upper
surface of costae short, with longer ones scattered
on distal parts of costae and on costules. Sori
inframedial; indusia sparsely hairy; sporangia
bearing glands; spores with a narrow erose wing,
surface otherwise bearing scattered small wings or
papillae.
Distr. Malesia: Borneo (Sarawak), only
known from the type.
120. Sphaerostephanos mutabilis (BRAUSE)
HOLTTUM, comb. nov.— Dryopteris mutabilis
BRAUSE, Bot. Jahrb. 56 (1920) 97. — Cyclosorus
mutabilis (BRAUSE) COPEL. Gen. Fil. (1947) 143;
Philip. J. Sci. 78 (1951) 443. — Thelypteris mutabilis
(BRAUSE) REED, Phytologia 17 (1968) 295.—
Type: LEDERMANN 9745, N.E. New Guinea,
Sepik Distr., on rocks, 200-400 m (B; UC).
Caudex short, erect. Stipe 5-8 cm long, reddish,
glabrous; scales c. 2% 1mm. Lamina to 20cm
long; apex pinna-like; pinnae 10 pairs, lower 3-4
pairs gradually smaller, lowest S-10mm_ long;
aerophores not elongate. Largest pinnae com-
monly 60.3cm (to 8.5x0.5cm, fide BRAUSE);
base and apex narrowly attenuate, edges entire;
costules 1.5mm apart, very oblique, forked or
simple, without anastomosis; lower surface quite
glabrous; upper surface hairy in groove of rachis
only. Sori in one row on each side of costa, often
on both branches of a costule; indusia large, firm,
with a few glands; sporangia sometimes bearing
glands; spores minutely spinulose.
Distr. Malesia: New Guinea. Known only
from type and BRASS 13047 from Idenburg
River.
Ecol. In forest, “massed in semi-shade on
steep flood-swept bank of river’ (BRASS), 200-
850 m.
121. Sphaerostephanos uaniensis HOLTTUM, Sp. nov.
Stipes 5cm longus; pinnae redactae 3-4-juga-
tae: lamina 33cm longa, pinnae normales 12-
jugatae, superiores plures non liberae; pinnae
liberae subintegrae, basi auriculatae; costae sub-
tus pilis erectis 0.1-0.2 mm longis vestita, pagina
inferior cetera pilis brevioribus praedita, eglan-
dulosa; sori elongati; indusia parva (vel nulla?);
sporangia interdum glandula una_praedita. —
Type: T. G. WALKER 10132, cult. Kew, origin
New Britain, Uani River (BM).
1981] THELYPTERIDACEAE (Holttum) 495
Stipe 5cm long; lamina in all 33cm long, con- from the type; ‘““common” at the type locality.
sisting of 3 pairs of reduced pinnae (basal one
7x5mm with stalk 1.5mm, uppermost 1.2 123. Sphaerostephanos peltochlamys (C. CHR.)
0.6cm with auricled base), 1 intermediate pair
(unequal) 2.0 and 2.7 cm long, and 12 pairs normal
pinnae of which several upper ones are adnate to
the rachis. Largest pinnae 4.2cm long, 1.4cm
wide at the auricled base, 1.1 cm wide above base;
apex rather abruptly obtuse; edges irregularly
sinuous; costules 2.5—3 mm apart, at c. 45°; veins 3
pairs, 2 pairs anastomosing, slender, concolorous;
lower surface of rachis bearing sparse stiff curved
pale hairs 0.2-0.3mm long, costal hairs 0.1-
0.2 mm, erect, very short hairs on costules and on
surface between veins, no glands; upper surface
hairy only on costae and rachis. Sori all con-
siderably elongate, those on basal veins of ad-
jacent costules almost meeting, distal ones shorter
and near costules; indusia very small with 2 or 3
short hairs, apparently sometimes _ lacking;
sporangia not setiferous, sometimes with a gland.
Distr. Malesia: New Guinea (New Britain),
only known from type.
122. Sphaerostephanos_ echinosporus (v.A.v.R.)
HOLTTUM, comb. nov. — Dryopteris echinospora
v.A.v.R. Bull. Jard. Bot. Btzg III, 2 (1920)
149. — Thelypteris echinospora (v.A.v.R.)
REED, Phytologia 17 (1968) 274.—Type:
BROOKS 451, Sumatra, Seblat River, Benkoelen
(BO; BM).
Caudex not known; stipe 5cm long; base of
stipe to largest pinna 80cm; reduced pinnae c. 15
pairs, uppermost (v.A.v.R.) 61cm, a middle
one seen 4.5 cm long, 7 mm wide in the middle and
gradually narrowed towards auricled base, lobed
half-way to costa, one near base 1.8cm long,
2mm wide apart from a narrow basal auricle.
Lamina excluding reduced pinnae 100cm long;
basal pinnae narrowed towards their bases which
have a narrow auricle 6mm long, upper pinnae
truncate to full width at base; aerophores to 2mm
long. Largest pinnae 16x 2.1cm; apex narrowly
caudate and entire; edges lobed to 2mm from
costa; lobes slightly falcate, separated by wide
sinuses; costules 3.5-4mm apart; veins to 11
pairs, basal pair, at a wide angle to costule, anas-
tomosing to form a short excurrent vein, next pair
both to margin above base of sinus; lower surface
of costae densely covered with hairs 0.2-0.3 mm
long, hairs on costules more sparse and somewhat
antrorse, sparse minute erect hairs present be-
tween veins, no glands; upper surface of costae
covered with antrorse hairs 0.6mm long, rest of
surface covered with slender appressed hairs
0.3mm long. Sori medial, covering lower surface
of pinna-lobes at maturity; indusia large, firm,
with sparse short hairs; a small proportion of
sporangia bearing a single seta or (less com-
monly?) a gland.
Distr. Malesia: South Sumatra. Known only
HOLTTUM, comb. nov.— Dryopteris peltoch-
lamys C. CHR. Dansk Bot. Ark. 9, 3 (1937) 65;
BACKER & POSTH. Varenfl. Java (1939) 338, excl.
syn.— Abacopteris peltochlamys (C. CHR.)
HOLTTUM, Rev. Fl. Mal. 2 (1955) 295, f. 171. —
Thelypteris peltochlamys (C. CHR.) REED, Phy-
tologia 17 (1968) 303.— Type: C. W. FRANCK
s.n. E. Java, Tanggore (BM).
Nephrodium urophyllum sensu RAciB. FI. Btzg
1 (1898) 184.
Caudex a short or long-creeping rhizome. Stipe
20-30cm long, glabrous, stramineous; scales
broad, thin; base of stipe to first large pinna
30-60cm (longest on fertile fronds). Reduced
pinnae 2-4 pairs (lacking on young plants), less
than 5mm long. Lamina to 60cm long; apex
pinna-like or with one basal lobe; pinnae 6-10
pairs, lower ones sometimes a little reduced, al-
ways with a narrowed base; aerophores not
swollen. Lower surface of rachis glabrous; upper
surface with short hairs in groove. Largest pinnae
of type 21 x 3.5cm; sterile pinnae to 4.5 cm wide,
fertile ones on small plants 14 x 2.8 cm; suprabasal
pinnae widest at or above the middle, slightly and
gradually narrowed to an abruptly contracted
broadly cuneate base; apex abruptly short cau-
date-acuminate; edges entire to crenate; costules
4mm apart, at 60°; veins 7-10 pairs, almost all
anastomosing to form a zig-zag excurrent vein:
sinus-membrane only distinct where margin is
crenate; lower surface quite glabrous; between
veins + verrucose when dry; upper surface of
costae with slender hairs 0.3 mm long. Sori medial
or somewhat inframedial; indusia large, thin,
glabrous, rarely with hairs or a few glands;
sporangia often bearing glands; spores with a
continuous translucent wing and several other
separate small wings.
Distr. Burma, in Malesia: Malaya (one collec-
tion in Kelantan), Sumatra and Java (many locali-
ties), in lowland forest.
Note. Apart from the reduced pinnae, this is
very similar to Pronephrium lineatum (BL.)
PRESL. It seems uniform over a wide area, and
may be a tetraploid of hybrid origin.
124. Sphaerostephanos irayensis (COPEL.) HOLT-
TUM, comb. nov. — Cyclosorus irayensis COPEL.
Philip. J. Sci. 81 (1952) 28; Fern FI. Philip. (1960)
340.— Thelypteris irayensis (COPEL.) REED,
Phytologia 17 (1968) 285. — Plesioneuron irayense
(COPEL.) HOLTTUM, Blumea 22 (1975) 239. —
Type: RAMOS BS 80344, Batanes Islands, Mt
Iraya (MICH; SING).
Sphaerostephanos fenixii HOLTTUM, Kalikasan
4 (1975) 55.— Type: FENIX BS 3780, Batanes
Islands (P; US).
Caudex short-creeping; stipe 35-70cm long,
496
glabrescent, reddish. Lamina 35-40 cm long; pin-
nae to 15 pairs, of firm texture; 1 pair reduced
basal pinnae 1.5cm long sometimes present;
lowest pinnae variably narrowed towards base
with stalk 1 mm long, to 2.5 cm wide at the middle;
aerophores not elongate. Lower surface of rachis
densely short-hairy. Suprabasal fertile pinnae to
12 x 2.2 cm (sterile to 2.8cm wide); base broadly
cuneate to full width; apex acuminate; edges
lobed more than 1/2 way to costa, in basal sterile
pinnae to more than 2/3; lobes falcate, those of
basal pinnae attenuate; costules 5 mm apart, at 60°
or more; veins 10-12 pairs, pale and prominent on
both surfaces, basal pair always connivent below
sinus, either anastomosing or touching sides of
sinus-membrane without fusing; lower surface of
costae hairy as rachis but less densely, of costules
and veins sparsely, hairs longer on sterile than on
fertile pinnae; upper surface densely short-hairy
on costae, sparsely on costules. Sori medial, im-
pressed on upper surface when dry, lower ones
distinctly elongate; indusia small, thin, with short
hairs; sporangia bearing 2-3 spherical yellow
glands; spores with a narrow erose translucent
wing and a few cross-wings.
Distr. Malesia: Philippines (Batanes Islands, 3
collections).
Note. This very interesting and isolated spe-
cies needs further study. The type of C. irayensis
resembles species of Plesioneuron in aspect and
texture, but its glands (on sporangia), and venation
and sinus-characters are unlike that genus.
125. Sphaerostephanos lithophyllus = (COPEL.)
HOLTTUM, comb. nov.— Dryopteris lithophylla
CoPEL. Philip. J. Sci. 12 (1917) Bot. 57; C. CHR.
Gard. Bull. Str. Settl. 7 (1934) 206, 245. — Thelyp-
teris lithophylla (COPEL.) REED, Phytologia 17
(1968) 289.—Type: TOPPING 1850, Sabah, Mt
Kinabalu (AFS; US).
Caudex short, suberect; apex covered with
slime. Stipe 1-2cm long, dark; base of stipe to
first large pinna 20-35cm. Reduced pinnae near
base 1 cm apart, distally 2-3 cm, in all to at least
10 pairs, all with rigid aerophore, lower ones
without a perceptible blade, upper ones 2 mm long
and wide. Lamina to 30cm long, red when young;
pinnae 12-18 pairs, fleshy when living, thick and
rigid when dried, 2-3 basal pairs slightly narrowed
at base; aerophores elongate. Rachis flushed with
red; both surfaces bearing a varied number of
short pale to dark red erect hairs; upper surface
deeply grooved. Largest pinnae 5-6.5 x 1.1cm;
base truncate; apex short-acuminate; edges lobed
about 1/3 to costa, lobes rounded, their margins
revolute on drying; costules to 3mm apart; each
with a small swelling at the base; veins to 8 pairs,
prominent on the upper surface and not below,
basal pair anastomosing, 2 pairs to sinus-mem-
brane which is very prominent on the lower sur-
face; lower surface of costae almost glabrous or
FLORA MALESIANA
(ser. II, vol. 1°
with pale short spreading hairs, some short hairs
present on surface between veins; sparse hairs
present on edges of groove of costa on upper
surface, no others. Sori supramedial; indusia with
a few short hairs and glands; sporangia bearing
glands, often several; spores copiously minutely
spinulose.
Distr. Malesia: Sabah (Mt Kinabalu).
Ecol. In wet ground in fully exposed places,
often among rocks, at 1500-2000 m.
126. Sphaerostephanos hamiferus (v.A.v.R.)
HOLTTUM, comb. nov.— Dryopteris hamifera
v.A.v.R. Bull. Jard. Bot. Btzg II, 16 (1914) 12;
Handb. Suppl. (1917) 181.— Thelypteris hamifera
(v.A.v.R.) REED, Phytologia 17 (1968) 281. — Type:
MATTHEW 5.n. Sumatra, G. Singgalang (BO).
Caudex erect. Stipe 12cm long; basal scales
thin, narrow; base of stipe to first large pinna
50cm. Reduced pinnae at least 10 pairs, lowest
5 mm long, upper ones 1.0-1.5 cm long, spreading,
narrowly triangular with auricle 5mm long on
acroscopic side, short on basiscopic; on type 1
pair intermediate pinnae also present. Lamina
70 cm long; pinnae 30 pairs; lowest pinnae slightly
narrowed and slightly auricled at base; aerophores
1-2 mm long (at least on reduced pinnae). Hairs on
lower surface of rachis dense, erect, to 0.5mm
long on fertile frond, mixed with many hairs 1 mm
long on sterile; hairs on upper surface 0.5-1 mm
long, less difference between fertile and sterile.
Largest fertile pinnae of type 10x 1.5cm; base
truncate; apex acuminate, not caudate; edges
lobed more than 3 way to costa (on a larger
specimen to 2/3), lobes slightly falcate; costules
3.5-4 mm apart, at more than 60°; veins 7-8(-10)
pairs, basal pair anastomosing, next pair both to
margin or acroscopic one to the short sinus-
membrane; lower surface of costae hairy as rachis
but. hairs a little shorter, hairs on costules sparse,
on fertile fronds antrorse but not closely ap-
pressed, glands absent or rare; hairs on upper
surface of costae to 1 mm long, scattered similar
hairs on costules and veins, surface between veins
+ closely covered with slender appressed hairs.
Sori medial; indusia large, firm, with short hairs;
sporangia sometimes with a gland, not setiferous.
Distr. Malesia: Central Sumatra, at 1500 m or
less.
Note. There is variability in size of fertile
pinnae from 6.4 0.9cm to 12 x 2.0cm, the latter
lobed fully 2/3.
127. Sphaerostephanos HOLTTUM,
sp. nov.
Caudex brevis, repens; pinnae redactae 20-30-
jugatae, aerophoris 3 mm longis constitutis; pin-
nae 10-jugatae, fertiles usque 305.5 cm; venae
18-20-jugatae; pagina inferior pinnarum glabra;
sori exindusiati; sporangia nec setis nec glandulis
praedita.—Type: JERMY 4038, N.E. New
plurivenosus
1981]
THELYPTERIDACEAE (Holttum)
497
Guinea, Madang Distr., gorge below Moro (BM).
Caudex massive, prostrate. Stipe c. 5cm long;
base of stipe to first true pinna 30-50 cm. Reduced
pinnae consisting of aerophores 3mm long with
no evident blade, 10-15 mm apart, 20-30 pairs.
Lamina 100cm or more long; apex pinna-like;
pinnae c. 10 pairs, opposite, basal ones not or little
reduced, lower ones 10cm apart. Rachis reddish,
glossy, quite glabrous apart from small hairs in
groove of upper surface. Largest fertile pinna
30 x 3.5 cm (sterile to 35 X 6.5 cm) with stalk 2mm
long and stiff aerophores | mm; base rounded to
broadly cuneate; apex acuminate with slender
cauda to 3cm long; edges parallel for most of
their length, entire or slightly crenate; costules
445mm apart, at almost 90°, falcate distally;
veins 18-20 pairs, close, at a wide angle to cos-
tules (more oblique in sterile pinnae), excurrent
veins mostly free, 2-3 veins from each costule
passing to margin; both surfaces quite glabrous,
eglandular. Sori supramedial, those on veins from
adjacent costules touching or fusing, exindusiate,
lower ones elongate; sporangia bearing neither
glands nor setae; spores with a continuous wing of
irregular width and some _ smaller’ wings.
Chromosomes: n= 36 (T. G. WALKER).
Distr. Malesia: Papua New Guinea; Madang
and Morobe Districts at 1300-1700 m; additional
specimens are CLEMENS 4841, Sattelberg;
HOOGLAND 9094, Mt Rawlinson.
Ecol. HOOGLAND’s specimen was from a
shaded vertical limestone cliff near a creek.
128. Sphaerostephanos squamatellus HOLTTUM,
sp. nov.
Pinnae redactae 8-jugatae, superiores 3mm
longae; pinnae normales 2/5 costam versus loba-
tae; aerophora elongata; costae costulaeque sub-
tus pilis appressis vestitae, eglandulosae; pagina
superior pinnarum pilis appressis vestita; indusia
pilosa; sporangia setifera. —Type: HOLTTUM 56,
Sabah, Mt Kinabalu, near waterfall in forest,
1900 m (K).
Caudex not preserved. Stipe 12cm _ long,
minutely hairy with many small scales; base of
stipe to first large pinna 35 cm. Reduced pinnae 8
pairs, uppermost 3mm long, with elongate aero-
phores. Lamina 40cm long; pinnae 16 pairs, all
almost opposite; basal pair of pinnae somewhat
shortened and a little narrowed at base which is
0.8cm wide. Rachis stramineous; lower surface
covered at base with very short erect hairs and
small appressed scales, distally with longer hairs
and few scales; upper surface bearing brown hairs
1mm long. Largest pinnae 10.5 1.7cm; base
broadly cuneate; apex rather abruptly short-acu-
minate; edges lobed about 2/5 to costa, lobes
slightly falcate; costules 44.5 mm apart, at 60° or
more; veins 7 pairs, basal pair anastomosing, 1-13
pairs passing to sinus-membrane which is not
prominent on the lower surface; lower surface of
costae covered with slender appressed hairs
0.3mm long, costules and veins the same with
minute uniseriate scales, appressed hairs 0.3 mm
long scattered on surface between veins; hairs on
upper surface of costae 0.7 mm long, rest of upper
surface bearing slender appressed hairs 0.3 mm
long. Sori medial; indusia firm with a few short
hairs; sporangia all with 2-4 short setae.
Distr. Malesia: Sabah (Mt Kinabalu) (also
CLEMENS 32548A at 2450 m, BO, L).
129. Sphaerostephanos canescens (BL.) HOLTTUM,
comb. nov. — Polypodium canescens BL. Enum.
Pl. Jav. (1828) 133.— Gymnogramme canescens
(BL.) BL. FI. Jav. Fil. (1829) 93, t. 40. — Phegop-
teris canescens (BL.) METT. Ann. Mus. Bot. Lugd.-
Bat. 1 (1864) 223 (non MEeETT. 1858); v.A.v.R.
Handb. (1908) 507, excl. var. omn.— Dryopteris
canescens (BL.) C. CHR. Ind. Fil. (1905) 250,
quoad typus tantum; BACKER & POSTH. Varenfl.
Java (1939) 56.— Haplodictyum canescens (BL.)
CHING, Sunyatsenia 5 (1940) 251.— Type:
REINWARDT, Moluccas, Tidore (L, n. 908,
300-133).
Caudex short, creeping. Stipe 9-12cm long,
densely short-hairy in groove; base of stipe to first
large pinna 19-24cm. Reduced pinnae to 5S pairs,
2-3mm long, 3cm apart. Lamina 27cm long
consisting of a deeply lobed apical section 18 cm
long, 2.7 cm wide at base, with gradual transition
to pinnae, and 5 pairs free pinnae, lowest longest,
all widest near their apices; aerophores elongate.
Rachis bearing hairs 1mm long both sides, less
abundant on lower surface with many shorter
hairs. Largest pinnae 4—4.5 cm long, 1.6cm wide
at 1/3 from apex; base truncate; apex very ab-
ruptly short-pointed; edges lobed 1/3 to costa;
costules to 3 mm apart at more than 60°; veins 7-8
pairs, | pair anastomosing and 2-2; pairs passing
to long sinus-membrane which is prominent on the
lower surface; hairs on lower surface of costae
pale, spreading, 0.4-0.7mm long, on costules
similar, erect veins present on surface between
veins; hairs on upper surface of costae more than
0.5mm long, similar hairs scattered on costules
and veins, whole surface covered with fine pale
appressed hairs. Sori medial, exindusiate;
sporangia bearing several setae; spores with many
small wings.
Distr. Malesia: West Java (Mt Gedeh), N.
Celebes, Moluccas (Tidore, Halmahera).
Notes. BLUME (1829) stated that the name
was given by REINWARDT to the specimen col-
lected by REINWARDT in Tidore, which may have
been the model for BLUME’s plate; BLUME stated
that he had also specimens from G. Gedeh (n. 908,
337-391) and these are certainly like REIN-
WARDT'’S.
This species was construed in a very broad
sense by CHRIST to include specimens which are
here referred to several other species, and in part
498
FLORA MALESIANA
(ser. II, vol. 1°
he was copied by v.A.v.R. The veins on some of
the lower lobes of the apical lamina are sometimes
forked as in Haplodictyum heterophyllum
PRESL, for which reason CHING transferred this
species to Haplodictyum.
130. Sphaerostephanos suboppositus HOLTTUM,
sp. nov.— Aspidium procurrens sensu CHRIST,
Ann. Jard. Bot. Btzg 15 (1898) 130, quoad
SARASIN 128.— Aspidium heterocarpus sensu
CHRIST, l.c. 133, quoad SARASIN 1326.
Pinnae redactae 9-jugatae, superiores 5mm
longae; pinnae normales usque 11 cm longae, ad
basin auriculatam 2.5 cm latae, suboppositae, 3/5—
2/3 costam versus lobatae, infimae basi leviter
angustatae; costae costulaeque subtus pilis
antrorsis non appressis vestitae, eglandulosae;
pagina superior pilis appressis vestita; sporangia
setifera. — Type: ALSTON 15744, Celebes, Mina-
hassa, G. Lokon (BM).
Caudex short, suberect. Stipe 8cm_ long,
glabrescent; scales thin, 5 X 1.5mm; base of stipe
to first large pinna 30-45 cm. Reduced pinnae 9
pairs, subopposite, lowest very small, uppermost
5mm long. Lamina 50-70 cm long; pinnae 18-21
pairs, subopposite in basal half of frond, basal 2-3
pairs somewhat narrowed at base; aerophores
slender, to 1mm long. Hairs on lower surface of
rachis dense, short, pale; on upper surface pale,
less than 0.5mm long. Middle pinnae to 11cm
long, to 2.5cm wide at auricled base (which is
sometimes a little dilated also on basiscopic side),
2.1cm wide above base; apex acuminate; edges
lobed 3/5—2/3 to costa, lobes slightly falcate with
obtuse tips; costules 4.5mm apart, at more than
60°; veins slender, 9-11 pairs, basal pair anas-
tomosing, next acroscopic vein or pair to sinus-
membrane; lower surface of costae with dense
somewhat antrorse short hairs, on sterile fronds
scattered long ones also present, hairs on costules
and veins short, antrorse, not appressed, short
erect hairs on surface between veins; upper sur-
face of costae covered with pale hairs more than
0.5mm long, similar hairs scattered on costules
and veins, surface generally covered with pale
appressed hairs. Sori medial, lower ones a little
divergent; indusia small, thin, with a few hairs;
sporangia bearing copious short setae; spores
short-spinulose.
Distr. Malesia: N. Celebes at 1100-1400 m.
131. Sphaerostephanos makassaricus HOLTTUM,
sp. nov.
Pinnae redactae c. 10-jugatae, usque 2 mm lon-
gae; pinnae normales usque 17 X 2.8 cm, profunde
lobatae; aerophori elongati; venae 12-jugatae,
infimae tantum anastomosantes; costae costu-
laeque subtus pilis 1mm longis brevioribus in-
termixtis vestitae, eglandulosae; indusia parva,
tenuia, brevi-pilosa; sporangia setifera.— Type:
POSTHUMUS 2727, S.W. Celebes, G. Bonthain,
S.W. slope, 1500 m, above Karoenglowe (BO).
Caudex short, prostrate; stipe 8cm long; base
of stipe to first normal pinna 50cm or more;
reduced pinnae c. 10 pairs, uppermost 2 mm long,
lower ones very small. Lamina excluding reduced
pinnae more than 50 cm long; 3 pairs lower pinnae
somewhat narrowed at their bases which are not
auricled; upper pinnae with almost symmetric
bases. Lower surface of rachis bearing copious
erect hairs 0.2mm long and scattered pale hairs
1mm long; upper surface with slender pale hairs
0.7mm long. Largest pinnae 17 X2.8cm, almost
sessile; aerophores more than Imm long; base
broadly cuneate; apex gradually attenuate with
cauda to 2cm long; edges lobed to 3mm from
costa, lobes falcate and somewhat tapered to a
blunt apex; costules 4.5-5.5mm apart, at 60° to
costa; veins to 12 pairs, | pair anastomosing,
next pair both ending at margin; lower surface
of costae bearing copious erect hairs 0.2-0.4 mm
long with many to 1 mm or more, on both sterile
and fertile pinnae, hairs on costules similar but
fewer, few hairs on veins, no hairs and no glands
between veins; upper surface of costae with pale
hairs 1mm long, shorter hairs scattered on cos-
tules and veins, appressed hairs 0.3—-0.4 mm long
on surface between veins. Sori medial,
lower ones not divergent; indusia small, thin, with
many short hairs; sporangia bearing 3-4 slender
setae.
Distr. Malesia: S.W. Celebes; a second spe-
cimen is POSTHUMUS 3476, from Patapang.
132. Sphaerostephanos sarasinorum HOLTTUM,
sp. nov. — ASpidium longipes sensu CHRIST, Ann.
Jard. Bot. Btzg 15 (1898) 133.
Caudex erectus; stipes 7cm longus; pinnae
redactae c. 9-jugatae, omnes parvae; lamina
100 cm longa; pinnae inferiores plurimae basin
versus valde angustatae; pinnae omnes profunde
lobatae, apicem versus abrupte caudatae; rachis
costaeque subtus pilis brevibus vestitae; pagina
inter venas utrinque glabra; sporangia setifera. —
Type: JERMyY 7458, Central Celebes, Masim-
bolong River, 1700 m (BM).
Caudex short, erect, Stipe 7cm long; basal
scales 10 x 1.5 cm, acuminate; base of stipe to first
large pinna 50cm or more. Reduced pinnae c. 9
pairs, all very small with swollen aerophores.
Rachis short-hairy on both surfaces. Lamina to
100 cm long; pinnae in basal half of frond all much
narrowed towards their bases which in lower ones
are c. 5mm wide; upper pinnae with basal lobes
only reduced. Largest pinnae of type 16 x 3 cm (of
SARASIN 996 21%X3.5cm); apex abruptly nar-
rowed to a subentire cauda 3 cm long; edges lobed
fully 3/4 towards costa, lobes falcate; costules
4.5-6.5mm apart; veins 12-16 pairs, basal ones
anastomosing with a short excurrent vein to sinus,
next acroscopic vein sometimes to sinus-mem-
brane; hairs on lower surface of costae short,
1981]
THELYPTERIDACEAE (Holttum)
499
distal ones and those on costules appressed, sur-
face between veins glabrous, eglandular; hairs on
upper surface of costae | mm long, similar hairs
scattered on costules and veins, surface between
veins glabrous or sometimes a few hairs present
near tips of pinna-lobes. Sori medial, lower ones
slightly divergent; indusia sparsely short-hairy;
sporangia setiferous.
Distr. Malesia: Central Celebes;
specimens are SARASIN 993, 996.
Ecol. In partial shade on very steep hillside.
Note. The SARASIN specimens lack base of
fronds, for which reason JERMY’s more perfect
specimen is chosen as type. Very young fronds
are covered with mucilage (JERMY).
additional
133. Sphaerostephanos muluensis HOLTTUM, sp.
nov.
Pinnae redactae 8-jugatae, omnes parvae;
aerophori leviter elongati; pinnae normales usque
15 x 2.5 cm, profunde lobatae; venae 11-12-juga-
tae, infimae non anastomosantes; pagina inferior
pinnarum eglandulosa; costulae et pagina inter
venas subtus pilis adpressis vestita; sori
supramediales; indusia pilosa; sporangia inter-
dum _setifera.—Type: B. S. PARRIS 6729,
Sarawak, Gunong Mulu, 1680 m (CGE).
Caudex short-creeping; stipe 15cm long, basal
scales 10mm long, narrow; base of stipe to first
large pinna 50 cm; reduced pinnae 8 pairs, all very
small. Lamina 70 cm long; pinnae 22 pairs, middle
ones 3-4cm apart, basal pair much narrowed
towards their bases (several successive ones
gradually less narrowed); aerophores (when dried)
less than 1mm long. Lower surface of rachis
rather densely covered with erect stiff brown hairs
0.3-0.4 mm long; upper surface bearing similar but
longer hairs, longest distally. Largest pinnae 15 x
2.5 cm, sometimes with a stalk to 1 mm long; apex
rather abruptly acuminate with subentire cauda
1.5-2cm long; edges lobed to 1.5-2mm from
costa; lobes almost at right angles to costa,
slightly narrowed distally, their tips obtuse; cos-
tules 5mm apart; veins 11-12 pairs, slender,
slightly prominent both sides, basal ones usually
both touching sides of the sinus-membrane, less
often both ending just above base of sinus; lower
surface of costae covered with antrorse but not
closely appressed pale hairs 0.6 mm long, hairs on
costules a little shorter, closely appressed, slender
appressed hairs 0.2-0.3mm long on surface be-
tween veins; upper surface bearing hairs about the
same length as on lower surface but thicker, no long
hairs on costules and veins, surface between veins
bearing appressed hairs 0.3 mm long. Sori distinctly
supramedial; indusia rather thin, covered with
hairs 0.3 mm long; some sporangia bearing a short
seta.
Distr. Malesia: Borneo (Sarawak), 3 collec-
tions.
Ecol. In forest at 1680-1800 m.
134. Sphaerostephanos telefominicus HOLTTUM,
sp. nov.
S. roemeriano (Rosenst.) Holttum affinis, ab ea
differt: caudice tenue, erecto vel scandente; lamina
longiore, pinnis 20-jugatis constata; pinnis dis-
taliter solum crenatis; pagina supra et subtus pilis
adpressis vestita. — Type: W. R. BARKER & J. R.
CROFT LAE 67649, N.E. New Guinea, W. Sepik
Distr., Telefomin Subdistr. (K; LAE).
Caudex slender, erect, sometimes creeping up
small shrubs (collectors); stipe 5—-12cm long,
copiously scaly, scales 5 mm long, less than | mm
wide, firm, hairy. Lamina to 25cm long; pinnae
more than 20 pairs, isomorphous; basal 2-3 pairs
slightly and gradually reduced, smallest 1.2 cm
long, subsessile, base unequally cuneate, not
auricled; apical lamina not pinna-like. Largest
pinnae 3.00.8 cm, with stalk 0.5 mm long; base
broadly cuneate to subtruncate, sometimes
slightly dilated both sides; apex short-acuminate;
edges subentire or slightly crenate distally; cos-
tules 2mm apart or a little more; veins to 4 pairs,
2 pairs anastomosing, next acroscopic vein pass-
ing to short sinus-membrane; lower surface of
rachis and costae bearing copious erect hairs
0.2 mm long (on sterile pinnae 0.3 mm), costules
and veins sparsely hairy, surface between veins of
sterile pinnae bearing many slender appressed
hairs; upper surface of rachis bearing hairs
0.5mm long with scattered ones |_mm, shorter
hairs present on costae and a few on costules and
veins, between veins slender appressed hairs
0.3 mm long. Sori medial; indusia thin, fairly large,
with short hairs; sporangia of type immature, not
setiferous, no glands seen on them.
Distr. Malesia: Papua New Guinea (Tele-
fomin), only known from the type.
Ecol. In disturbed montane forest near river, at
1500 m.
135. Sphaerostephanos roemerianus (ROSENST.)
HOLTTUM, comb. nov. — Dryopteris roemeriana
ROSENST. Nova Guinea 8 (1912) 723; v.A.v.R.
Handb. Suppl. (1917) 179. — Thelypteris roemeri-
ana (ROSENST.) REED, Phytologia 17 (1968)
310. — Type: VON ROEMER 1023, W. New Guinea,
1350 m (BO).
Caudex short, erect. Stipe 3cm long, covered
with short pale hairs; scales narrow. Lamina to
12cm long; apex pinna-like; pinnae 10-12 pairs,
lower 2-3 pairs gradually smaller, lowest 3-7 mm
long. Lower surface of rachis covered with pale
spreading hairs 0.3mm long. Largest pinnae to
3.0 x 0.8 cm; base unequally cuneate, sometimes a
little auricled on acroscopic side; apex slightly
tapered to a rounded tip; edges crenate; costules
2.5mm apart, at 45°; veins to 3 pairs, basal pair
anastomosing; lower surface of costae, costules
and veins bearing rather sparse short hairs; hairs
present on upper surface of costae, scattered on
costules and veins. Sori inframedial; indusia large,
500
FLORA MALESIANA
[ser. II, vole
glabrous; sporangia not setose, no glands seen.
Distr. Malesia: W. New Guinea, only known
from the type.
136. Sphaerostephanos hellwigensis HOLTTUM, sp.
nov.
Caudex erectus, usque 30cm altus; pinnae
redactae 4—S-jugatae, usque 8-9 mm longae; pin-
nae normales usque 4.5 x 0.9 cm, crenatae, eglan-
dulosae, costae costulaeque subtus pilis erectis
brunneis 0.5-0.6 mm longis vestitae; sori mediales,
indusia parva, pilis brevibus praedita; sporangia
non setifera, ut videtur non glandulosa. — Type:
PULLE 884, W. New Guinea, Mt Hellwig, 2600 m
(BO).
Caudex erect, to 30cm tall (collector); stipe
5-10 cm long, scales rather firm, 5 x 1 mm; base of
stipe to first normal pinna 20cm; reduced pinnae
4-5 pairs, lowest 1-2 mm long, uppermost 8-9 mm
long with an acroscopic auricle 4mm long, basis-
copic base narrower, edges almost entire. Lamina
excluding reduced pinnae 40 cm long; pinnae to c.
30 pairs, texture rigid; basal normal pinnae not
narrowed at their bases. Largest pinnae 4.5 x
0.9cm; base truncate and slightly dilated; aero-
phores c. 0.5mm long; apex short-acute; edges
crenate to a depth of 1 mm or a little more deeply;
costules 2.5 mm apart; veins to 4 pairs, prominent
both sides, 1 pair anastomosing, next acroscopic
vein or pair to the sinus-membrane; lower surface
of rachis bearing thick erect brown hairs 1mm
long, of costae similar hairs 0.5-0.6mm_ long,
fewer on costules, no hairs between veins, no
glands; upper surface of rachis covered with
antrorse brown hairs, brownish antrorse hairs less
than 0.5 mm long on costae, a few short hairs on
costules distally. Sori medial; indusia small, thin,
with a few short hairs; sporangia not setiferous,
no glands seen on them.
Distr. Malesia: W. New Guinea (Mt Hellwig),
2600 m.
137. Sphaerostephanos benoitianus (GAUD.)
HOLTTUM, comb. nov. — Polystichum benoiti-
anum GAUD. in Freyc. Voy. Uran. Phys. Bot.
(1827) 331. — Aspidium benoitianum GAUD. ibid.
t. 11.— Dryopteris benoitiana (GAUD.) v.A.v.R.
Handb. (1908) 225. —Type: GAUDICHAUD, New
Guinea, Rawak Island (G; BM, FI-W).
Caudex short, creeping. Stipe 7-9cm long,
short-hairy; basal scales dark, firm, 4mm long.
Lamina 27cm long; apical 10-12cm narrowly
triangular and deeply lobed, grading to upper pin-
nae; free pinnae 10-12 pairs; basal 1-2 pairs
somewhat reduced, narrowed at base on basis-
copic side, not on acroscopic, in middle wider on
basiscopic than on acroscopic side of costa.
Lower surface of rachis bearing short erect hairs
and scattered pale ones 1 mm long, hairs on its
upper surface thick, brown, spreading. Largest
pinnae 5.5 x 1.4cm; base subtruncate, symmetric;
apex short-acuminate; edges lobed a little more
than 3 way to costa (to 2/3 on fertile fronds), lobes
slightly falcate; costules 3mm apart; veins 6-8
pairs, basal pair anastomosing, next acroscopic
vein to sinus-membrane or to margin; lower sur-
face of costae and costules hairy as rachis; hairs
on upper surface of costae sparse, short and long,
rest of surface usually glabrous. Sori submarginal;
indusium small, bearing short hairs; sporangia
with neither glands nor setae; spores dark,
minutely papillose.
Distr. Malesia: W. New Guinea (Rawak I.),
only known from the type.
Note. This appears to be an isolated species,
possibly related to S. invisus.
138. Sphaerostephanos neotoppingii HOLTTUM,
nom. nov. — Dryopteris toppingii COPEL. Philip.
J. Sci. 12 (1917) Bot. 56; C. CHR. Gard. Bull. Str.
Settl 7 (1934) 246.— Thelypteris toppingii
(COPEL.) K. IwaTs. Acta Phytotax. Geobot. 21
(1965) 168, not Sphaerostephanos _ toppingii
(COPEL.) C. CHR. 1934. — Type: TOPPING 1766,
Sabah, Mt Kinabalu, Lobang (AFS, not seen).
Caudex short, creeping. Stipe 50-90 cm long,
dull reddish, minutely hairy; scales narrow, to
10mm long. Reduced pinnae lacking. Lamina 50-
80cm long; pinnae 10-15 pairs, lowest not or little
reduced; several pairs lower pinnae much nar-
rowed towards their bases and stalked 34mm,
distal ones broadly cuneate to subtruncate, almost
sessile; aerophores not elongate. Rachis densely
short-hairy (0.2-0.3 mm) on both surfaces. Largest
pinnae commonly 15-20 x 2.0-2.5cm (maximum
24x 3.5cm); base cuneate; apex acuminate with
subentire cauda 1.5—2.5 cm; edges lobed 1/3-2/5 to
costa, lobes distinctly falcate with narrow blunt
forward-pointing tips; costules 5-6.5 mm apart at
more than 60°; veins to 12 pairs on acroscopic side
of costule, to 14 on basiscopic side, prominent on
lower surface, 13-23 pairs anastomosing, 1-2 pairs
to sides of sinus-membrane; hairs on lower sur-
face of costae antrorse but not closely appressed,
0.2-0.3 mm long with some 0.5mm, similar and
more sparse hairs on costules and veins, on sur-
face between veins a variable number of short
suberect hairs, glands lacking; hairs on upper sur-
face of costae 0.3mm long, shorter on costules,
between veins (especially on young fronds) some
short appressed hairs. Sori supramedial, those on
lowest veins from adjacent costules sometimes
touching; indusia firm, glabrous, sometimes a little
elongate and asymmetric at base; sporangia lack-
ing glands and setae, hair on sporangia-stalk end-
ing in a red gland.
Distr. Malesia: Borneo (Sabah and Kaliman-
tan), in forest at 1200-1500 m.
139. Sphaerostephanos oosorus (BAK.) HOLTTUM,
comb. nov.— Nephrodium oosorum BAK. Kew
Bull. (1896) 41.— Dryopteris oosora (BAK.) C.
1981]
THELYPTERIDACEAE (Holttum)
501
Cur. Ind. Fil. (1905) 280; Gard. Bull. Str. Settl. 7
(1934) 245. — Type: G. F. HOSE 334, Sabah, Pulo
Gaya (K).
Caudex short, creeping. Stipe 25-40cm long,
minutely hairy; base of stipe to first large pinna
40-55 cm. Reduced pinnae 2-4(—6) pairs, upper-
most 4mm long. Lamina 60 cm long; pinnae to 20
pairs, sessile; 2-3 pairs lower pinnae narrowed at
their bases; aerophores not elongate. Lower sur-
face of rachis covered with short erect pale hairs
0.2-0.4mm long, in some specimens mixed with
long ones to 1mm; upper surface with uniform
pale hairs 0.5 mm long. Largest pinnae commonly
12-15 x 1.8-2.0 cm (largest seen 21 x 2.4 cm); apex
acuminate with cauda to 1.5cm; edges lobed a
little less than 2 way to costa, lobes oblique, nar-
rowed to obtuse tips; costules 44.5 mm apart, at
60°; veins to 8 pairs, basal 12 pairs anastomosing,
1-1 pairs to sinus-membrane; lower surface of
costae covered with erect hairs varying from 0.2
to 0.5 mm, hairs on costules and veins similar but
shorter, surface between veins bearing many short
erect hairs; hairs on upper surface of costae
0.5mm long, shorter on costules, abundant short
suberect hairs between veins. Sori medial, elon-
gate, 1.5 mm long; indusia large, pale, firm, bearing
many short hairs and rarely a gland; neither glands
nor setae on sporangia; spores minutely papillose.
Distr. Malesia: Borneo (Sabah & P. Balam-
bangan).
Ecol. In forest at low altitudes.
140. Sphaerostephanos rudis (RIDL.) HOLTTUM,
comb. nov.—Goniopteris rudis RiDL. Trans.
Linn. Soc. Bot. 9 (1916) 259.— Phegopteris
ridleyana v.A.v.R. Handb. Suppl. (1917) 515, nom.
nov.— Dryopteris rufopilosa BRAUSE, Bot.
Jahrb. 56 (1920) 106, nom. nov. (not D. rudis
(KUNZE) C. CHR. nor D. ridleyana BRAUSE). —
Type: KLoss s.n. W. New Guinea, Mt Carstensz,
Camp I-III, 150-750 m (BM; K).
KEY TO THE VARIETIES
1. Lower surface of rachis and costae bearing
thick reddish hairs : . a. var. rudis
1. Lower surface of rachis and costae glabrous
b. var. micans
a. var. rudis
Caudex short-creeping; stipe 10cm long, dark
reddish, dark-hairy in groove, scales 7 x 1mm,
thin; base of stipe to first normal pinna 35cm;
reduced pinnae 5-6 pairs, all very small, with short
aerophores. Lamina 50cm long; apex pinna-like;
pinnae 4 pairs, opposite, all widest at or above
middle and gradually narrowed to an abruptly
cuneate base. Lower surface of rachis covered
with thick spreading dark red hairs 1 mm long.
Largest pinnae 20x4.5cm; apex abruptly acu-
minate with cauda 2-2.5cm long; edges entire;
costules 5mm apart; veins 8 pairs, most of them
united to form a zig-zag excurrent vein; lower
surface of costae bearing thick erect reddish hairs
0.5 mm long which are more sparse distally, hairs
on costules short and sparse, none on surface
between veins; upper surface glabrous apart from
a few hairs in groove of costa. Sori medial, exin-
dusiate; sporangia bearing neither glands nor
setae.
Distr. Malesia: W. New Guinea and Japen
Island, at low altitudes.
b. var. micans (BRAUSE) HOLTTUM, Stat. nov. —
Dryopteris micans BRAUSE, Bot. Jahrb. 56
(1920) 98.—Cyclosorus micans (BRAUSE)
COPEL. Gen. Fil. (1947) 143; Philip. J. Sci. 78
(1951) 458.— Thelypteris micans (BRAUSE)
REED, Phytologia 17 (1968) 293.—Type:
LEDERMANN 12468, N.E. New Guinea, Sepik
Distr. 1400-1500 m (B).
Differs from var. rudis in quite glabrous lower
surface of rachis and costae; all specimens also
are larger, with up to 10 pairs of reduced pinnae
and 6 pairs of normal ones which are not opposite,
largest 27 x 5.5 cm.
Distr. Malesia: New Guinea, from Idenburg
River eastwards, at 850-1750.
141. Sphaerostephanos dimorphus (BRAUSE)
HOLTTUM, comb. nov.— Dryopteris dimorpha
BRAUSE, Bot. Jahrb. 56 (1920) 100. — Cyclosorus
dimorphus (BRAUSE) COPEL. Philip. J. Sci. 78
(1951) 442.— Thelypteris dimorpha (BRAUSE)
REED, Phytologia 17 (1968) 273.— Type:
LEDERMANN 12622, N.E. New Guinea, Sepik
Distr., 1400-1500 m (B).
Dryopteris morobensis COPEL. Univ. Cal. Publ.
Bot. 18 (1942) 221.—Cyclosorus morobensis
(COPEL.) COPEL. Gen. Fil. (1947) 143; Philip. J.
Sci. 78 (1951) 459, pl. 38. — Thelypteris moroben-
sis (COPEL.) REED, Phytologia 17 (1968) 294. —
Type: CLEMENS s.n. N.E. New Guinea, Morobe
Distr., Finugan, 1400 m (MICH).
Caudex erect, 100-150cm tall. Stipe 10-20cm
long, dull red, glabrous, with many scales to 15 x
2mm. Lamina to 140cm long; apex pinna-like;
pinnae to 40 pairs, dimorphous, lower 10-12 pairs
gradually reduced, lower ones narrowly triangular
with + hastate base, lowest 7-15 mm long; aero-
phores swollen, not or little elongate. Rachis
glabrous apart from dark red hairs in groove of
upper surface, most abundant distally. Largest
sterile pinnae commonly 20 2.0cm (on type to
36 x 1.8cm); base asymmetric, rounded on basis-
copic side, cuneate on acroscopic; apex narrowly
acuminate; edges slightly undulate; costules 4—
5mm apart, at a wide angle to costa; veins 4-6
pairs, all very oblique, pale and prominent on
lower surface, 2-23 pairs anastomosing and 1 pair
to sides of sinus-membrane; both surfaces of
pinnae quite glabrous. Fertile pinnae 7-14 cm long,
502
0.7-1.2 cm wide; edges entire as sterile, or crenate
(rather strongly crenate in type of D. morobensis);
veins 4 pairs; sori confluent at maturity; indusia
glabrous or with a few short hairs; sporangia
lacking glands and setae; spores minutely papil-
lose.
Distr. Malesia: Eastern New Guinea, in forest
at 1200-2000 m, many collections.
142. Sphaerostephanos potamios HOLTTUM, sp.
nov.
Caudex erectus, c. 20cm altus; frondes dimor-
phae, omnes pinnis redactis 6-8-jugatis praeditae;
pinnae Steriles 15*1.6cm, fertiles usque 8.0
0.8 cm, non ultra 1/4 costam versus lobatae, subtus
eglandulosae; rachis pilis brunneis vestita; sori
mediales; indusia parva, glabra vel pauci-pilosa;
sporangia nec setis nec glandulis praedita. — Type:
J. CROFT & Y. LELEAN NGF 34913, S.E. New
Guinea, Port Moresby Subdistr. near Lake Myola
(I):
Caudex erect, 20cm tall; stipe 8 cm long, basal
scales 7-8 X 1 mm, covered with many short hairs:
base of stipe to first large pinna of sterile frond
25cm, of fertile 35-42 cm; reduced pinnae of fer-
tile frond 6-8 pairs, not opposite, uppermost
1.5cm long with basal acroscopic auricle 8mm
long, apex acute, lowest reduced pinna 3-4 mm
long. Lamina 63 cm long (sterile); pinnae 30 pairs,
basal pinnae with slightly dilated truncate bases;
texture firm; apex of frond not pinna-like; aero-
phores slightly swollen. Sterile pinnae to 15x
1.6 cm; apex narrowly acuminate; edges lobed to a
depth of 1-2.5mm, lobes with falcate tips and
slight projections at ends of veins; costules 3.5-
4.5 mm apart; veins 6-7 on basiscopic side, 5-6 on
acroscopic, pale and prominent on lower surface,
reddish on upper, basal 1-15 pairs anastomosing,
13 pairs passing to the rather long sinus-mem-
brane; lower surface of rachis bearing rather
sparse thick brown hairs 0.7 mm long, hairs on
costae sparse, pale, erect, 0.3-0.4mm long, hairs
on costules shorter, + antrorse, not appressed,
surface between veins hairless, slightly pustular;
upper surface of rachis with brown hairs to 1 mm
long, of costae with pale hairs 0.50.7 mm, no
other hairs. Fertile pinnae to 8.0x0.8cm (ap-
parently not quite fully expanded); veins 3-4
pairs; hairs on both surfaces shorter and more
sparse than on sterile pinnae; sori medial; indusia
small, thin, glabrous or with a few short hairs;
sporangia not setiferous, no glands seen on them.
Distr. Malesia: S.E. New Guinea; only known
from the type.
Ecol. In forest close to stream-bank, at 1900 m.
143. Sphaerostephanos invisus (FORST. f.) HOLT-
TUM, Webbia 30 (1976) 195; Allertonia 1 (1977)
211, f. 7B. — Polypodium invisum ForstT. f. FI.
Ins. Austr. Prodr. (1786) 81.— Nephrodium in-
visum (ForRST. f.) CARR. in Seem. FI. Vit. (1873)
FLORA MALESIANA
[ser. II, vol. 1°
362. — Dryopteris invisa (FORST. f.)O. KTZE, Rev.
Gen. Pl. 2 (1891) 813; v.A.v.R. Handb. (1908) 212;
COPEL. Bishop Mus. Bull. 59 (1929) 45; 93 (1932)
38. — Cyclosorus invisus (FORST. f.) COPEL. Gen.
Fil. (1947) 142; BROWNLIE in Aubrév. FI. Nouv.
Cal. 3 (1969) 215, excl. syn. Nephrodium haen-
keanum PRESL.— Thelypteris forsteri MORTON,
Contr. U.S. Nat. Herb. 38 (1967) 60, nom. nov. (not
T. invisa (DESV.) PROCTOR). — Type: FORSTER,
locality unrecorded (BM; K, LE).
Aspidium dissectum sensu METT. Ann. Mus.
Bot. Lugd.-Bat. 1 (1864) 232, excl. synon.
Caudex a long-creeping rhizome to 7mm
diameter. Stipe 25-45cm or more long. Lamina
commonly 40-75 cm long (to 120 cm); basal 2 pairs
of pinnae gradually smaller and irregularly spaced,
lowest sometimes 2cm long, usually longer;
aerophores not elongate. Largest pinnae com-
monly 15 x 1.3cm (to 24x 1.8cm), narrowly acu-
minate, lobed 1/3-2/5 to costa, lobes falcate, dis-
tinctly pointed, oblique; costules commonly 3-
3.5mm apart; veins 7-8 pairs, close, very oblique
except basal pair, 1-1) pairs anastomosing, 1-2
pairs to sinus-membrane; short spreading hairs on
all part of lower surface, a little antrorse on cos-
tules, erect (sometimes lacking) between veins;
upper surface almost glabrous apart from costae.
Sori supramedial; indusia thin, hairy; sporangia
bearing a variable number of setae; spores dark,
with irregular + continuous brownish wing and a
few cross-wings.
Distr. Polynesia (many islands in the Pacific
eastward to Tahiti and Hawaii), New Caledonia,
Queensland; in Malesia: New Guinea (lowlands of
north and east) and Moluccas (Kei & Tenimber
Is., Amboina).
Ecol. In open or lightly shaded places, some-
times in wet ground, at low altitudes.
144. Sphaerostephanos austerus (BRAUSE)
HOLTTUM, comb. nov.—Dryopteris austera
BRAUSE, Bot. Jahrb. 56 (1920) 108. — Cyclosorus
austerus (BRAUSE) COPEL. Philip. J. Sci. 78
(1951) 446.— Thelypteris austera (BRAUSE)
REED, Phytologia 17 (1968) 262.—Type:
LEDERMANN 11750, N.E. New Guinea, Sepik
Distr., 2070 m (B).
Caudex slender, scandent. Stipe 11-18 cm long.
Lamina 65-100cm long; 5-6 pairs lower pinnae
gradually reduced, all with an acroscopic auricle,
lowest 1.3cm long, subtriangular with rounded
apex; aerophores not elongate. Hairs on lower
surface of rachis dense, short; on upper surface a
little longer. Largest pinnae 9-13cm long, 2.2-
3.0cm wide; base subtruncate; apex abruptly
short-acuminate; edges lobed 3 way to costa or a
little more deeply, lobes falcate, blunt-pointed;
costules 5—S.5mm apart, at 60°; veins 7 pairs,
slender, basal pair anastomosing, next pair usually
both to margin; lower surface of costae bearing
dense short pale somewhat antrorse hairs of uni-
1981]
THELYPTERIDACEAE (Holttum)
503
form length, similar hairs sparse on costules,
scattered on surface between veins; hairs on up-
per surface of costae short, scattered longer ones
present on costules and veins, surface between
veins bearing scattered short appressed hairs. Sori
slightly inframedial, exindusiate; sporangia with a
few setae; spores minutely spinulose.
Distr. Malesia: Papua New Guinea (Sepik),
only known from type.
145. Sphaerostephanos mundus (ROSENST.)
HOLTTUM, comb. nov.— Dryopteris munda
ROSENST. Meded. Rijksherb. n. 31 (1917) 5.—
Thelypteris munda (ROSENST.) CHING, Bull.
Fan Mem. Inst. Biol. Bot. 10 (1941) 252. — Type:
ATASRIP s.n. 1903, W. New Guinea (L).
Dryopteris oblonga BRAUSE, Bot. Jahrb. 56
(1920) 109.—Cyclosorus oblongus (BRAUSE)
CoPEL. Philip. J. Sci. 78 (1951) 446. — Thelypteris
oblonga (BRAUSE) REED, Phytologia 17 (1968)
297.— Type: LEDERMANN 10100, N.E. New
Guinea, Sepik Distr., 1000 m (B).
Dryopteris farinosa BRAUSE, Bot. Jahrb. 56
(1920) 111. — Cyclosorus farinosus (BRAUSE) Co-
PEL. Philip. J. Sci. 78 (1951) 446. — Thelypteris far-
inosa (BRAUSE) REED, Phytologia 17 (1968) 276.
— Type: LEDERMANN 12103, Sepik Distr., 2070 m
(B).
Caudex a scandent rhizome 7-10 mm diameter,
or sometimes free, slender, erect. Stipe 5-10cm
long, dark; base of stipe to first large pinna 25-
35 cm. Reduced pinnae 5-7 pairs, smallest 3-4 mm
long, uppermost 10 mm, spreading, triangular with
base auricled both sides (more on acroscopic
side). Lamina 60-100 cm long; pinnae c. 25 pairs,
not dimorphous; lower pinnae a little narrowed to
symmetric base; aerophores not elongate. Lower
surface of rachis sparsely short-hairy; upper sur-
face with hairs to 1mm long. Largest pinnae
commonly 9-12 cm long, 2.0-2.5 cm wide (on type
of D. oblonga 18 x 2.6cm); base broadly cuneate;
apex rather abruptly acuminate with cauda 1.5—
3cm long; edges lobed 2/3-3/4 to costa, lobes
falcate with blunt forward-pointing tips; costules
4-5 mm apart (to 6.5mm on type of D. oblonga);
veins 7-9 pairs, basal pair anastomosing to form a
short excurrent vein to sinus, next pair both to
margin; lower surface of costae bearing rather
sparse short erect hairs (dense with some longer
ones on type of D. oblonga), sparse short hairs on
costules, veins and surface between veins; hairs on
upper surface of costae less than 1 mm long,
sparse short hairs on costules and veins, a variable
number of fine appressed hairs between veins (in
some cases very sparse). Sori inframedial, lowest
not or little divergent; indusia absent or very small
with a few hairs; sporangia setiferous; spores
closely papillose.
Distr. Malesia: New Guinea, from West to East.
Ecol. Usually climbing on trees in forest, at
1000-2000 m.
Note. BRAUSE distinguished D. farinosa by a
grey-yellow deposit on the lower surface; this
consists of a fungus. In the type of D. munda and
others from Western New Guinea, basal veins
sometimes meet at the sinus without anas-
tomosing.
146. Sphaerostephanos_ obtusifolius (ROSENST.)
HOLTTUM, comb. nov. — Dryopteris obtusifolia
ROSENST. Fedde Repert. 10 (1912) 336. — Phe-
gopteris obtusifolia (ROSENST.) v.A.v.R. Handb.
Suppl. (1917) 315.—Cyclosorus obtusifolius
(ROSENST.) COPEL. Gen. Fil. (1947) 143; Philip.
J. Sci. 78 (1951) 442.— Thelypteris obtusifolia
(ROSENST.) REED, Phytologia 17 (1968) 297. —
Type: Frau BAMLER 43, N.E. New Guinea, Sat-
telberg (not seen).
Caudex short, creeping. Stipe 2-8cm long.
Lamina 8 cm long; apex not pinna-like; pinnae c.
8 pairs, | or 2 basal pairs reduced, lowest 4-6 x 3-
5 mm; aerophores slightly swollen. Hairs on lower
surface of rachis minute, on upper surface 0.4 mm
long, brown. Largest pinnae 1.5—2.0 cm long, 0.8-
10cm wide; base subtruncate, slightly asym-
metric; apex broadly rounded, edges sinuous with
very slight development of sinus-membranes;
costules 2.5mm apart; veins 2-3 pairs, 15 pairs
anastomosing; all hairs on lower surfaces minute;
on upper surface short hairs on costae only or a
few between veins near margin. Sori exindusiate,
those on basal veins from adjacent costules
sometimes fusing; sporangia setiferous; spores
pale, with a moderate number of wings of varying
extent.
Distr. Malesia: New Guinea
several collections; Manus Island).
Notes. At BM is a specimen labelled KEyYS-
SER 43, from the type locality, named D. obtusi-
folia by ROSENSTOCK;; this is probably an error
for BAMLER 43 (ROSENSTOCK was describing
specimens from both collectors). Also from
Sattelberg is Rosenst. Fil. Novoguin. exsicc. 115,
coll. BAMLER (B, BM, W). M. SANDS 2682 (K)
from limestone on Manus Island is closely similar
to the Sattelberg specimens but less hairy on both
surfaces of pinnae.
(Sattelberg,
147. Sphaerostephanos alcasidii HOLTTUM, sp.
nov.
Pinnae redactae plurimae, 1.5 cm inter se dis-
tantes, aerophoris tumidis tantum constitutae;
lamina 80cm longa; pinnae 35-jugatae, maximae
15 1.5cm, 3/5 costam versus lobatae, eglan-
dulosae; pagina utroque latere inter venas pilis
adpressis vestita; indusia parva brevi-pilosa;
sporangia setifera.— Type: ALCASID PNH 1747,
Luzon, Mountain Prov., Mt Data (MICH).
Caudex not known. Stipe 10cm long; base of
stipe to first large pinna 50cm. Reduced pinnae
many pairs, 1.5 cm apart, all except the very small
upper ones consisting of an aerophore without
504
perceptible blade. Lamina 80cm long; pinnae 35
pairs, lowest narrowed to 8mm wide at the base.
Lower surface of rachis near base covered with
appressed hairs 0.5 mm long, distally with longer
spreading hairs; hairs on upper surface similar.
Largest pinnae 15x 1.5cm; base truncate; apex
acuminate with entire cauda 1-2cm long; edges
lobed a little more than > way to costa, lobes
falcate; costules 3.5mm apart at more than 60°;
veins 8-9 pairs, basal pair anastomosing, next
acroscopic vein to sinus-membrane; hairs on lower
surface of costae near base appressed, 0.5 mm
long, stally and on costules longer and spread-
ing, appressed hairs on lower surface between
veins; upper surface of costae bearing hairs 1 mm
long, similar hairs scattered on costules and veins,
copious appressed hairs 0.3mm long on sur-
face between veins. Sori medial; indusia small
with short hairs; sporangia sometimes with 1-2
setae.
Distr. Malesia: Philippines (Luzon: Mt Data),
only known from the type.
148. Sphaerostephanos pilosissimus HOLTTUM, sp.
nov.
Pinnae redactae 12-jugatae, infimae 0.7. cm
longae, superiores 2.5cm, anguste triangulares,
crenatae; pinnae evolutae usque 16.5 X2.3 cm,
dimidio costam versus lobatae; costae costulaeque
subtus pilis erectis vestitae, eglandulosae; indusia
pilifera; sporangia setifera.— Type: HOLTTUM
20222, Sumatra, G. Kerinci, 1800-2100 m (SING;
BO).
Caudex erect, short. Stipe 15 cm long, glabrous;
base of stipe to first large pinna 40cm. Reduced
pinnae at least 12 pairs, deflexed, basal one 7 mm
long, uppermost 2.5 cm long, narrowly triangular
with acute apex, crenate edges and slight basal
acroscopic auricle. Lamina 65 cm long; pinnae 25
pairs; basal pinnae narrowed and less deeply lobed
to base which is 1.3cm wide, basal basiscopic
lobes of several successively higher pinnae shor-
tened; aerophores swollen, not elongate. Hairs on
lower surface of rachis dense, erect, 0.3 mm long,
on upper surface more than 0.5mm long, pale.
Largest pinnae 16.5 X 2.3 cm; base truncate; apex
acuminate with subentire cauda to 2.5cm long;
edges lobed about 3 way to costa, lobes slightly
oblique with rounded tips; costules 4.5 mm apart;
veins 10-14 pairs, basal 2 pairs anastomosing, |
pair to sinus-membrane; lower surface of costae
and costules densely covered with erect hairs of
varied length to 0.5 mm, less dense hairs on veins
and surface more than 0.5 mm long; hairs on upper
surface of costae 0.7 mm long, costules, veins and
surface between them covered with slender ap-
pressed hairs 0.3mm long. Sori a little in-
framedial, lower ones not divergent; indusia
fringed with short hairs and a few on surface;
sporangia bearing 2-3 short setae.
Distr. Malesia: West Central Sumatra (G.
FLORA MALESIANA
[ser. II, vol. 1°
Kerinci), 2 collections (ALSTON
1800-2100 m.
14185, BM),
149. Sphaerostephanos — erwinii
nov.
Frondes parvae, lamina apicali 7-8 cm longa,
pinnis evolutis 6-jugatis pinnisque redactis 4-
jugatis constitutae; pinnae redactae usque 8mm
longae; pinnae evolutae usque 3.21.3 cm,
subintegrae; indusia parva, pilosa; sporangia
setifera. — Type: STRESEMANN 43, Ceram, Cen-
tralgeb., G. Hoale, 1000 m (L).
Caudex short, suberect. Stipe 7cm long, slen-
der, short-hairy; base of stipe to first large pinna
17cm. Reduced pinnae 4 pairs, ovate, uppermost
8 mm, lowest 3-4 mm long. Apical lamina 7-8 cm
long, 2.5 cm wide at base, deeply lobed and grad-
ing to pinnae; free pinnae 6 pairs. Lower surface
of rachis bearing thick brown hairs and shorter
pale ones. Largest pinnae 3.2 x 1.3cm, widest a
little above middle; base truncate, sometimes with
a slight acroscopic auricle; apex abruptly nar-
rowed to a rounded or obtuse tip; edges shallowly
crenate; costules 3.5 mm apart, at little more than
45°: veins 3 pairs, 1 pairs anastomosing; hairs on
lower surface of costae short, erect, with some
longer, sparse hairs on costules and veins, short
erect hairs on surface between veins; scattered
long hairs on upper surface of costules and veins
like those on costae, appressed hairs on sur-
face between veins. Sori medial; indusia small
with hairs more than 0.5 mm long; sporangia seti-
ferous.
Distr. Malesia: Moluccas (Ceram) and West
New Guinea.
Note. The plant from W. New Guinea (Kos-
TERMANS 2209a, Angi-gita Lake, at 2000 m, BO)
is smaller than the type, with pinnae to
2.2x0.8cm, and only 2 pair of reduced basal
pinnae.
HOLTTUM, | sp.
150. Sphaerostephanos atasripii (ROSENST.)
HOLTTUM, Webbia 30 (1976) 194.— Dryopteris
atasripii ROSENST. Meded. Rijksherb. n. 31
(1917) 6; C. CHR. Brittonia 2 (1937) 298.—
Thelypteris atasripii (ROSENST.) REED, Phy-
tologia 17 (1968) 261.— Type: ATASRIP 211, W.
New Guinea (L).
Cyclosorus diminuens HOLTTUM, Blumea 13
(1965) 135.—Type: BRASS 32106, N.E. New
Guinea, Eastern Highlands, Arau, 1400 m (K).
Caudex massive, erect. Stipe 13cm long; base
of stipe to first large pinna 75 cm. Reduced pinnae
15 pairs, spreading, broadly triangular, lowest 6—
7mm long and 10mm wide at base, uppermost
4-5 x 2.5-3.0cm. Lamina to more than 100cm
long; apex not pinna-like; lower pinnae not nar-
rowed at base; aerophores slightly swollen. Lower
surface of rachis densely covered with erect hairs
0.1-0.2 mm long; upper surface covered with ap-
pressed pale hairs 0.5mm long with thicker
1981]
spreading brown hairs | mm long on each side of
the groove. Largest fertile pinnae 17*2.5cm,
sterile to 22 x 3.1 cm; base truncate and somewhat
dilated both sides; apex short-acuminate; edges
lobed to a depth of 2-3 mm; costules 44.5 mm
(fertile), to 6mm (sterile) apart; veins 9-11 pairs,
233 pairs anastomosing, 2-3 pairs to sides of
sinus-membrane; lower surface of costae densely
covered with short hairs as rachis, hairs on cos-
tules and veins more sparse, on sterile pinnae
longer than those on costae, copious short erect
hairs on surface between veins; hairs on upper
surface of costae 0.7 mm long, a few similar hairs
on costules and veins, surface between veins +
densely covered with fine appressed hairs. Sori
medial, lower ones not divergent; indusia covered
with hairs 0.3mm _ long; sporangia setiferous,
sometimes with an acicular hair on the stalk;
spores closely and minutely _ spinulose.
Chromosomes: n= 36 (T. G. WALKER).
Distr. Malesia: throughout New Guinea, in
forest at 700-1400 m.
Notes. This species is very similar to S. con-
fertus in all characters except the massive erect
caudex and much larger fronds which never bear
glands. In cultivation the two remain quite distinct
in size and habit.
151. Sphaerostephanos albosetosus (COPEL.)
HOLTTUM, comb. nov.— Dryopteris albosetosa
CopEL. Univ. Cal. Publ. Bot. 18 (1942) 221.—
Cyclosorus albosetosus (COPEL.) COPEL. Gen.
Fil. (1947) 142; Philip. J. Sci. 78 (1951) 455, pl.
35.— Thelypteris albosetosa (COPEL.) REED,
Phytologia 17 (1968) 259.— Type: BRAss 11435,
W. New Guinea, 18km north of Lake Habbema,
on dry face of a limestone wall, in shade (MICH:
BM, BO, L).
Caudex short, creeping, 4-Smm_ diameter,
bearing close but seriate stipes; stipe 6cm long,
basal scales ovate, 1 mm long; base of stipe to first
normal pinna 12-24cm; reduced pinnae 6 pairs,
lowest 4mm long, others gradually longer; tran-
sition to normal pinnae subabrupt, an intermediate
pair 1.5cm long, 6mm wide above auricled base,
apex broadly obtuse. Lamina 21 cm long; pinnae
10 pairs, lower ones not auricled at their bases;
aerophores not elongate. Lower surface of rachis
bearing short pale erect hairs with some to 1mm
long; hairs 1 mm long also on upper surface. Lar-
gest pinnae 4.0 1.0cm; base truncate on acros-
copic side, rounded on basiscopic; apex abruptly
narrowed to obtuse tip: edges lobed to a depth of
1.5-2 mm, lobes strongly concave beneath (spe-
cimen at BO); costules 3.5mm apart; veins 4-6
pairs, thick and prominent on lower surface, basal
pair anastomosing, second pair passing to sinus-
membrane or margin; lower surface of costae
bearing short erect hairs mixed with longer ones
to 1mm, more sparse similar hairs on costules,
THELYPTERIDACEAE (Holttum)
505
short slender erect hairs on and between veins;
upper surface of costae bearing hairs less than
1 mm long, scattered similar hairs on costules and
veins, surface between veins covered with fine
appressed hairs. Sori medial, basal ones much
divergent; indusia small with copious long hairs;
sporangia setiferous.
Distr. Malesia: W. New Guinea (near Lake
Habbema), only known from the type.
Ecol. On dry face of limestone wall, in shade.
152. Sphaerostephanos wauensis HOLTTUM, sp.
nov.
Pinnae redactae usque 9-jugatae, infimae 5 x
4mm, superiores 1.4 * 1.0 cm, triangulares; pinnae
evolutae usque 12*2.0cm, 2/5 costam versus
lobatae; rachis utrinque pilis patentibus brunneis
1mm vel ultra longis pilisque brevioribus vestita;
costae costulae venaeque subtus pilis pallidis
1mm longis praeditae, eglandulosae; indusia
magna, tenuia, pilis brevibus instructa; sporangia
setifera.— Type: T. NAKAIKE 96, N.E. New
Guinea, Morobe Distr., Wau, Mt Missim 1300-
1600 m (K; TNS).
Caudex massive, suberect. Stipe 17cm long;
basal scales narrow, 7mm or more long; base of
stipe to first large pinna 5S cm. Reduced pinnae to
9 pairs, alternate; basal ones 5 x 4mm, uppermost
1.5x1.0cm, triangular with symmetric base,
crenate; one pair intermediate pinnae present.
Lamina 60 cm long: pinnae 21 pairs; lower pinnae
slightly narrowed towards base which may be
slightly auricled both sides; aerophores not elon-
gate. Rachis bearing on both sides spreading thick
brown hairs more than 1mm long (sometimes
lacking on lower surface), on lower surface also
short erect pale hairs, on upper surface short
brown hairs. Largest pinnae 12 x2.0cm, sessile;
base broadly cuneate to subtruncate, not dilated;
apex short-acuminate; edges lobed rather less than
> way to costa; lobes oblique and slightly falcate;
costules 4-5mm apart, at 60° or a little less;
veins to 8 pairs, 1> pairs anastomosing and 1 pair
to sinus-membrane; lower surface of costae bear-
ing many spreading pale hairs more than 1mm
long with a varied range of shorter ones, hairs on
costules and veins sparse, to 1mm long, short
erect hairs between veins; upper surface of costae
bearing slender pale hairs 1 mm long, similar hairs
scattered on costules and veins, short appressed
hairs on surface between veins not abundant. Sori
inframedial, lower ones not divergent; indusia
rather large, thin, bearing hairs 0.3mm _ long;
sporangia setiferous.
Distr. Malesia: Papua New Guinea (Morobe
District), at 1300-1800 m, several collections.
Note. A smaller plant has fertile pinnae 6.8 x
1.3 cm which are slightly stalked (WOMERSLEY &
MILLAR 8309, from 1800m), otherwise agreeing
with the type.
[ser. II, vol. 1°
FLORA MALESIANA
506
“CN
hae Nast ‘ as Mi
Gy kam ry ay —
Bho 7 yay ay ala
Spin ppp als
b. venation and position of sori in apical
2
35
lamina, x 3.— P. heterophyllum (PR.) HOLTTUM. c. Fertile and sterile fronds, x
of apical lamina, x 3.— P. micropinnatum HOLTTUM. e. Frond, x
Fig. 14. Pronephrium womersleyi HOLTTUM. a. Frond, xX
3; d. venation of part
2
3
2
3
3; f. part of apical lamina, x 4. — P.
h. venation and position of sori, x 2; i. one sorus, X< 16 (a
2
33
NAKAIKE 69, b BRASS 31670, c-d PRICE & HERNAEZ 162, e-f cult. Kew, g-i HENDERSON 21863).
asperum (PR.) HOLTTUM. g. One pinna, xX
1981] THELYPTERIDACEAE (Holttum) 507
18. PRONEPHRIUM
PRESL, Epim. Bot. (1851) 258, excl. P. lastreoides PRESL; HOLTTUM, Novit.
Bot. Inst. Bot. Univ. Carol. Prag. 1968 (1969) 48; Blumea 19 (1971) 34;
Blumea 20 (1972) 105—126.— Type species: Pronephrium lineatum (BL.)
PRESL, l.c. (designated by HOLTTUM, I.c. 1969).
Haplodictyum PRESL, l.c. 50; CHING, Sunyatsenia 5 (1940) 251, p.p.;
COPEL. Fern Fl. Philip. (1960) 377; HoLTTUM, Blumea 19 (1971) 37;
Kalikasan 2 (1973) 59, excl. H. majus. — Thelypteris subg. Haplodictyum K.
IWATS. Mem. Coll. Sci. Univ. Kyoto B, 31 (1964) 32.—Type species:
Haplodictyum heterophyllum PRESL, l.c.
Abacopteris FEE, Gen. Fil. (1852) 309; CHING, Bull. Fan Mem. Inst. Biol.
Bot. 8 (1938) 230; Acta Phytotax. Sinica 8 (1963) 331; HOLTTUM, Rev. FI.
Malaya 2 (1955) 285, excl. A. peltochlamys (C. CHR.) HOLTTUM. — Dry-
opteris sect. Abacopteris C. CHR. Gard. Bull. Str. Settl. 7 (1934) 247. —
Thelypteris subg. Abacopteris K. IWATS. Mem. Coll. Sci. Univ. Kyoto B,
31 (1964) 34.— Type species: Abacopteris philippinarum FEE, l.c. (desig-
nated by HOLTTUM, I.c. 1969) = Pronephrium asperum (PRESL) HOLTTUM.
Dimorphopteris TAGAWA & K. IWATS. Acta Phytotax. Geobot. 19 (1961)
8.— Type species: Dimorphopteris moniliformis TAGAWA & K. IWATS.
l.c. — Fig. In-o, 14-16.
Caudex creeping or suberect (never truly erect); fronds simple or simply
pinnate, basal pinnae not reduced but often narrowed at base on basiscopic
side, less often on acroscopic; apical lamina pinna-like or widened towards
its base and then merging with the upper pinnae; pinnae crenate to entire;
veins several pairs (except in a few species with small pinnae), almost all
anastomosing, the excurrent veinlets arising from their union in some
species free, more often joining to form zig-zag composite veins alternating
with the costules; sinus-membranes short or (in species with entire pinnae)
lacking; lower surface of old fronds usually pustular when dried; acicular
hairs usually present on lower surfaces, hooked in sect. Grypothrix; sessile
spherical yellow glands present on lower surface, indusia and/or sporangia
in some species; sori in some species exindusiate and then often spreading
along the veins; sporangia often bearing short setae, less often spherical
glands or both glands and setae; spores of most species with a + continuous
longitudinal wing and cross-wings, three species with many small separate
wings or papillae.
Distr. India & Ceylon; from S. China southwards throughout Malesia; North Queensland; Solomon
Islands, New Hebrides, Fiji; c. 68 species, of which 12 not Malesian.
Ecol. Most species are terrestrial in forest or on stream-banks in shade (in several cases on rocks
only); two are limestone ferns in more exposed positions (P. simillimum, P. scopulorum).
Cytol. Chromosome number 36; 10 species investigated, of which P. pentaphyllum, P. peltatum var.
peninsulare, P. womersleyi and P. triphyllum are tetraploid.
Taxon. CHRISTENSEN (Ind. Fil. 1905) accepted Abacopteris FEE (which he included in Dryopteris) as
published in 1843, but that publication consisted of the generic name only with no description. CHING
gave generic status to Abacopteris, and cited Aspidium lineatum BL. as type, but FEE did not mention
that species when the generic name was validly published by him in 1852; he then listed Pronephrium
PRESL as a genus not adopted, as he had seen none of the species (p. 358).
508 FLORA MALESIANA [ser. II, vol. 1°
cS
= ig NE TST ee
Tt Gi & :
Fig. 15. Pronephrium amboinense (WILLD.) HOLTTUM. a. Lower pinnae, <3. — P. celebicum (BAK.)
HOLTTUM. b. Sterile pinna, x 1.5; c. fertile pinna, x 1.5; d. two indusia, x 100. — P. hewittii (COPEL.)
HOLTTuo. e. Middle sterile pinna, x 1.5; f. basal sterile pinna, x 1.5; g. fertile pinna, x 1.5; h. venation
of sterile pinna, x 4; i. sorus, x 16.— P. rhombeum (CHRIST) HOLTTUM. j. Fertile frond, x3; k. ster-
ile frond, <3; i. sorus, X 16.— P. granulosum (PR.) HOLTTUM. m. Fertile frond, <3; n. sorus, X24 (a
from photo of type, b DRANSFIELD 3776, c-d CURTIS 431, e-i MOLESWORTH ALLEN 3048, j—n cult.
Kew).
1981] THELYPTERIDACEAE (Holttum) 509
Pronephrium PRESL (1851) comprised four species, which I discussed in 1969, citing P. lineatum
(BL.) PRESL as type and accepting the statement by METTENIUS and CHRISTENSEN that the second
species P. affine (BL.) PRESL was not distinct from the first. Subsequently (HOLTTUM 1971) I examined
the types of both of BLUME’s species and found them to be quite distinct from each other. The third of
PRESL’S species is here named P. rhombeum; the fourth (P. lastreoides PRESL) is here transferred to
Sphaerostephanos.
PRESL’s most distinctive character is dimorphism of fertile and sterile fronds; this is doubtfully true
of P. lineatum (the fertile frond of the type is not fully expanded) but is true of species 2 and 3. FEE
distinguished Abacopteris mainly by subentire pinnae with meniscioid venation but the excurrent
veinlets not free, and indusiate sori. CHING (1938) extended it to admit species in mainland Asia with
exindusiate sori and typical meniscioid venation; he also cited absence of sinus-membranes as a generic
character, but short membranes are often present (the line of distinction is not a sharp one).
CHRISTENSEN (1934) had already added species from Malesia as Dryopteris sect. Abacopteris. In 1971
and 1972, under the earlier name Pronephrium, 1 accepted the generic concepts of CHING and
CHRISTENSEN, adding the character absence of reduced basal pinnae, and divided the whole into three
sections: Pronephrium, Dimorphopteris and Grypothrix. But some of the species then placed in sect.
Pronephrium have resemblances to species in sect. Dimorphopteris and some to species in sect.
Grypothrix. I here attempt to adjust this situation by re-arranging the species in two subgenera
Pronephrium and Menisciopsis. Haplodictyum PRESL (which I treated as a separate genus in 1971) is
included in sect. Pronephrium. But though sections Dimorphopteris and Grypothrix, as re-arranged,
appear to be natural groups, the other two sections are probably not; problems of relationships are
discussed under the sections.
KEY TO THE SUBGENERA AND SECTIONS
. Veins in pinnae less than 10 pairs, or if more than 10, sori indusiate; yellow spherical glands often
ene at least on sporangia sigh: . . . . 1. SUBG. PRONEPHRIUM
2. Apical lamina pinna-like or much larger than pinnae: cainnce not auricled on acroscopic base; fronds
of young plants simple, in P. beccarianum all fronds simple . . . . . .41. Sect. Pronephrium
. Apical lamina not ae or if so pinnae auricled on acroscopic base; fronds of young plants
rarely Simpleia ee . . . . 2. Sect. Dimorphopteris
1. Veins in pinnae 10 pairs or more, or if Tease excurrent veinlets all free; sori exindusiate; spherical
glands never present a Fe 7) ee 22 SUBGS MENTISCIOPSIS
3. No hooked hairs on any part of plant; sori rarely ‘coalescent . . . . . . 3, Sect. Menisciopsis
3. Hooked hairs present on sporangia or on some other part of plant; sori often elongate or coalescent
4. Sect. Grypothrix
1. Subgenus Pronephrium
1. Section Pronephrium
Haplodictyum PRESL, Epim. Bot. (1851) 50.
The following species, included here in 1972, are now transferred to other sections: the non-Malesian
P. lakhimpurense and allied species, also the Malesian P. nitidum, P. repandum and P. acanthocarpum,
to sect. Menisciopsis; P. aquatiloides and P. menisciicarpon to sect. Dimorphopteris.
The species here included belong to three groups:
1. Species which appear to be related to the type, P. lineatum. Of these, P. euryphyllum has glands on
the lower surface and appressed hairs on the upper surface as in P. glandulosum of sect. Dimor-
phopteris. P. asperum and P. gymnopteridifrons lack glands but have the same frond-form as the type.
None of these have the reddish tinge shown by species of subg. Menisciopsis. But they show
resemblances to Sphaerostephanos peltochlamys, S. simplicifolius and S. spenceri, which differ only in
the presence of several pairs of much-reduced basal pinnae.
2. Species which have an elongate apical lamina and a few much smaller pinnae, and sori which lack
indusia. These are specialized forms the relationship of which it is difficult to suggest. Three of them
have glands on their sporangia as in P. lineatum. The most aberrant species, lacking glands, is P.
micropinnatum; it may belong elsewhere.
3. Species included in Haplodictyum in my arrangement of the genera in 1971, but excluding H.
majus COPEL. which is now placed in Sphaerostephanos because it has much-reduced basal pinnae in
addition to normal ones. These species have glands on the lower surface of pinnae and indusiate sori.
510 FLORA MALESIANA [ser. II, vol. 1°
I NG 2
Fig. 16. Pronephrium rubicundum (v.A.v.R.) HOLTTUM. a. Complete frond, <3; b. venation of sterile
frond, x 3; c. venation of fertile frond and position of sori, x 4; d. costa and costule showing hooked
hairs, x 12.— P. ramosii (CHRIST) HOLTTUM. e. Upper part of frond, x3; f. bud at base of a pinna,
x 8.— Sporangia, x35, and spores, 235: g—h. P. nitidum HOLTTUM; i-j. P. repandum (FEE)
HOLTTuM; k-l. P. triphyllum (Sw.) HOLTTUM (a-d MATTHEW s.n., e-f ELMER 17851, g-I after
HOLTTuUM, Blumea 19: 24, fig. 8-9 and SEM photos).
1981] THELYPTERIDACEAE (Holttum) ei
Two of them have the peculiar venation in their terminal lamina which caused PRESL to establish the
genus Haplodictyum: the veins in the lower part of the apical lamina are forked, their branches
anastomosing to form a series of areoles along the costules, also sometimes additional areoles below the
sinuses. This venation resembles that of Pleocnemia, and on that account CHING placed Haplodictyum
in his family Aspidiaceae, though it is unquestionably thelypteroid. But the third species (P. bakeri) does
not have the veins thus forked, though in every other character it resembles the other two. The
Haplodictyum type of venation occurs to a small extent at the base of the apical lamina in several
species of other genera of Thelypteridaceae where the transition from apical lamina to pinnae is not
sharply defined. The peculiar venation is like that of pinnae, of which the upper ones are more or less
adherent to the base of the apical lamina. There is a North American fossil named Goniopteris
claiborniana BERRY (Bull. Torr. Bot. Cl. 44, 1917, 331, t. 22) which shows the Haplodictyum venation.
KEY TO LAE SPECIES
1. Pinnae of fronds of adult plants about equal in size to apical lamina.
2. Glands present on lower surface of pinnae
2. Glands lacking on lower surface of pinnae.
3. Pinnae 2-3 pairs; glands present on indusia or sporangia or both
.1. P. euryphyllum
. 2. P. lineatum
3. Pinnae to at least 7 pairs; no glands on indusia and sporangia.
4. Pinnae to 30 x 5 cm; spores with a median wing and cross-wings
4. Pinnae to 15 x 2.5 cm; spores with many separate small wings
F . 3. P. asperum
. 4. P. gymnopteridifrons
1. Apical lamina much larger than pinnae, or fronds simple.
5. Sori exindusiate; glands usually lacking on lower surface between veins.
6. No free pinnae; sometimes one small pair broadly adnate
6. Free pinnae present on adult plants.
7. Sporangia bearing glands near annulus.
.5. P. beccarianum
8. Apical lamina to 20 x 3.5 cm; midrib of apical lamina and rachis glabrous on lower surface
6. P. womersleyi
8. Apical lamina to 6X 1.7cm; hairs present on lower surface of midrib of apical lamina and
rachis
7. P. melanophlebium
7. Sporangia bearing setae, “not glands, near annulus.
9. Terminal lamina lobed 1/4—1/3 towards midrib; pinnae to at least 1 cm long.
10. Pinnae 4-8 cm long
SF Pinnae not over 2 cm long.
. No glands on lower surface
Glands present on lower surface
9: eee lamina crenate; pinnae 2-S mm long. 3
. Sori indusiate; glands always present on lower surface between veins.
8. P. pentaphyllum
. 9. P. brauseanum
. 10. P. diminutum
11. P. micropinnatum
5 Veins in basal part of apical lamina forked, their branches anastomosing.
13. Apical lamina widest near its base; pinnae 1 pair
12. P. bulusanicum
13. Apical lamina not widest near its base; pinnae 3 pairs, decreasing downwards
12. Veins in apical lamina not forked
1. Pronephrium euryphyllum (ROSENST.) HOLT-
TUM, Blumea 20 (1972) 112.— Dryopteris eury-
phylla ROSENST. Meded. Rijksherb. n. 31 (1917)
7.— Thelypteris euryphylla (ROSENST.) REED,
Phytologia 17 (1968) 276.— Type: KORTHALS,
Sumatra (L; B).
Dryopteris urophylla var. teysmannii v.A.v.R.
Bull. Jard. Bot. Btzg II, 28 (1918) 24.— Type:
TEYSMANN, Sumatra, Loeboe Alang (BO?; not
found).
Sterile fronds unknown. Fertile fronds consis-
ting of terminal lamina to 21 x 5.5 cm and at least 3
pairs of pinnae. Pinnae to 18 X 6cm, base broadly
cuneate, sides parallel for most of their length,
crenate to depth of 1-1.5 mm, apex abruptly nar-
rowed to a cusp 1-2 cm long; costules 44.5 mm
apart, at more than 60° to costa, slightly falcate;
13. P. heterophyllum
14. P. bakeri
veins c. 15 pairs, almost all anastomosing, excur-
rent veinlets not free; lower surface of rachis,
costae and costules covered densely with short
erect hairs, similar shorter hairs with many glands
on surface between veins; upper surface densely
covered throughout with short appressed hairs.
Sori supramedial, spreading a little along veins;
indusia hairy, persistent but shrivelling; sporangia
bearing several short setae; spores pale, with con-
tinuous wing and cross-wings.
Distr. Malesia: Sumatra; the two types cited.
Notes. The KORTHALS collection at L con-
sists of 5 sheets (2 also at B); on one of the
smaller fronds the basal pinnae are somewhat
reduced and narrowed at their base, but the lowest
(of three pairs) on the largest specimen are not, so
that there may have been more pairs of pinnae on
il
the latter. VAN ALDERWERELT’Ss short descrip-
tion of TEYSMANN’s specimen is adequate for
identification; the number of pairs of pinnae is not
specified.
2. Pronephrium lineatum (BL.) PRESL, Epim. Bot.
(1851) 259; Ho_truM, Blumea 19 (1971) 34; 20
(1972) 112.— Aspidium lineatum BL. Enum. PI.
Jav. (1828) 144; METT. Farngatt. IV (1858) 264, p.p.
— Meniscium lineatum (BL.) KUNZE, Bot. Zeit.
6 (1848) 259.—Nephrodium lineatum (BL.)
PRESL, Epim. Bot. (1851) 48; HooK. Spec. Fil. 4
(1862) 74, p.p. — Cyclosorus lineatus (BL.) TARD.
& C. CHR. Notul. Syst. 7 (1938) 74, nomen
tantum. — Abacopteris lineata (BL.) HOLTTUM,
Rev. Fl. Malaya 2 (1955) 292, nomen tantum. —
Thelypteris lineata (BL.) K. IwaTs. Mem. Coll.
Sci. Univ. Kyoto B, 31 (1964) 34.— Type:
BLUME, Java (L. n. 909, 27-60; isotypes P, PRC).
Alsophila fragilis ZOLL. & MOoORITZI, Nat.
Geneesk. Arch. Ned. Ind. 1 (1844) 400. — Menis-
cium sp. Morirzi, Syst. Verz. (1846) 116.—
Meniscium fragile (ZOLL. & MOR.) KUNZE, Bot.
Zeit. 6 (1848) 259.—Type: ZOLLINGER 1019,
Java, Tjikoya river (L; FI).
Dryopteris verruculosa v.A.v.R. Bull. Jard. Bot.
Btzg II, 11 (1915) 12; Handb. Suppl. (1917) 177. —
Thelypteris verruculosa (v.A.v.R.) REED, Phy-
tologia 17 (1968) 323.—Type: BACKER 3954,
Java, Pasuruan (BO).
Dryopteris menisciicarpa (BL.) POSTH. in
Backer & Posth. Varenfl. Java (1939) 61, p.p.
Caudex short-creeping or suberect. Stipe of
sterile frond 20cm, of fertile 40cm long, pale,
glabrescent except for hairs in groove. Sterile
frond. Apical lamina to 15 x 4 cm; pinnae 2-3 pairs,
to 9.5 3.0cm, widest 1/3 from apex, gradually
narrowed to abruptly broad-cuneate base, apex
rather abruptly short-cuspidate, edges crenate
distally, sinuous towards base; costules 3mm
apart; veins 7-8 pairs, almost all anastomosing,
excurrent veinlets not free; lower surface with
rather sparse erect hairs on costae, much shorter
on costules, surface between veins glabrous and
finely pustular, no glands present; upper surface
hairy on costae only. Fertile pinnae of type not
fully expanded, largest possibly 3.0 1.5cm, of
ZOLLINGER 1019 9X2.5cm, of BACKER 3954
7 x 2.2 cm; sori inframedial; indusia small, bearing
a few short hairs and glands; sporangia with
glands, or both glands and setae; spores with
translucent wing and cross-wings.
Distr. Malesia: Central & East Java (3 collec-
tions) and Sabah (Bettotan, KLOss SFN 19083);
Sulu. Archipelago, Tawi-Tawi (ALCASID &
CELESTINO PNH 7507).
Notes. METTENIUS (1858) included Aspidium -
affine BL. as a synonym, and his description is
composite; HOOKER (1862) copied him; later
authors mostly described A. affine under the name
lineatum, not noticing that BLUME’s descriptions
FLORA MALESIANA
{ser. II, vol2 1?
of the two species indicate fronds of quite
different form.
BLUME gave Noesa Kambangan as type local-
ity, but the only specimen named in his hand is
unlocalized; it agrees well with his description.
The published description of Alsophila fragilis
ZOLL. & Mor. is doubtfully valid, but the spe-
cimens of ZOLLINGER 1019 certainly agree with
BLUME’s. The sporangia of the type have both
glands and setae; those of ZOLLINGER 1019
glands only.
3. Pronephrium asperum (PRESL) HOLTTUM,
Blumea 20 (1972) 112.— Polypodium asperum
PRESL, Rel. Haenk. (1825) 24, t. 3, f. 4, non
LINN.; HOLTTUM, Novit. Bot. Inst. Bot. Univ.
Carol. Prag. 1968 (1969) 18.— Goniopteris aspera
PRESL, Tent. Pterid. (1836) 183, nom. nov. —
Aspidium asperum (PRESL) MEtTT. Ann. Mus.
Bot. Lugd.-Bat. 1 (1864) 225, excl. syn. Neph-
rodium latifollum PRESL.— Dryopteris presliana
CHING in C. CHR. Ind. Fil. Suppl. 3 (1934) 95,
nom. nov. superfl.— Abacopteris presliana
(CHING) CHING, Bull. Fan Mem. Inst. Biol. Bot. 8
(1938) 248, p.p.— Abacopteris aspera CHING,
Acta Phytotax. Sinica 8 (1963) 332. — Thelypteris
aspera REED, Phytologia 17 (1968) 261. — Type:
HAENKE, Luzon (PRC).
Abacopteris philippinarum FEE, Gen. Fil.
(1852) 310, t. 18C f. 1. — Cyclosorus philippinarum
(FEE) COPEL., Fern Fl. Philip. (1960) 372.—
Nephrodium latifolium PRESL, Epim. Bot. (1851)
45, p.p.— Type: CUMING 16, Luzon (isotypes
BM, G, K, LE).
Meniscium kennedyi F.v.M. Fragm. 4 (1864)
165.— Goniopteris kennedyi (F.v.M.) BAILEY,
Handb. Queensl. Ferns (1874) 41, f. 31.— Type:
DALLACHY & KENNEDY, Rockingham Bay,
Queensland (MEL; K).
Dryopteris urophylla (WALL. ex HOOK.) C.
CHR. var. novoguineensis ROSENST. in Fedde
Rep. 10 (1912) 336; v.A.v.R. Handb. Suppl. (1917)
177.— Type: G. Bamler 80, Sattelberg, New
Guinea (orig. not seen; probable isotypes BM, L,
W, as Rosenst. fil. novoguin. exsic. 174).
Abacopteris multilineata (WALL. ex HOOK.)
CHING var. malayensis HOLTTUM, Rev. FI.
Malaya 2 (1955) 297, f. 173, nom. illeg. — Thelyp-
teris multilineata (WALL. ex HOOK.) CHING var.
malayensis (HOLTTUM) REED, Phytologia 17
(1968) 294 (type not cited). — Lectotype: HOLT-
TUM SFN_ 15303, Trengganu, Kuala Telumong
(SING; K).
Aspidium repandum sensu BL. Enum. PI. Jav.
(1828) 144, excl. syn.
Nephrodium glandulosum sensu J. SM. in Hook.
J. Bot. 3 (1841) 411 (misidentification of CUMING
16); DIELS in E. & P. Nat. Pfl. Fam. I Abt. 4 fig.
92G “nach Fée”’ (inaccurate copy of FEE, Gen.
Fil. t. 18C, f. 1).— Aspidium glandulosum sensu
KUNZE, Bot. Zeit. 6 (1848) 260, quoad Cuming 16
1981]
THELYPTERIDACEAE (Holttum)
S13
tantum; METT. Farngatt. IV (1858) 111, quoad
syn. Abacopteris philippinarum FEE tantum.
Polypodium urophyllum sensu. BENTH. FI.
Austral. 7 (1878) 766, excl. BEDD. Ferns Brit. Ind.
t. 3.—Nephrodium urophyllum sensu. BEDD.
Handb. (1883) 274, p.p.; sensu RAcIB. FI. Btzg 1
(1898) 184. — Dryopteris urophylla sensu v.A.v.R.
Handb. (1908) 216, p.p.; sensu BACKER &
PosTH. Varenfl. Java (1939) 64, p.p. — Fig. 14g-i.
Caudex short-creeping. Stipe to 70cm or more
long, slightly flushed red, glabrescent, base at first
covered with narrow thin scales. Lamina to 60 cm
or more long; pinnae 6-8 pairs, lower ones +
narrowed at base, lowest sometimes reduced, tex-
ture thin; apical lamina more strongly crenate than
pinnae. Largest pinnae to 37 x 6cm, sessile, base
abruptly broad-cuneate, apex short-acuminate,
edges subentire to distinctly crenate, parallel for
most of their length; costules 5—5.5 mm apart, at
60° to costa; veins 12-15 pairs, at c. 45°, nearly all
anastomosing, excurrent veinlets usually not free;
sinus-membrane short but distinct; lower surface
of costae, costules and veins bearing short hairs,
surface between veins glabrous or rarely with
short erect hairs, finely verrucose when dry; upper
surface hairy only on costa. Sori medial, lowest
ones supramedial and + coalescent; indusia small,
with or without short hairs; sporangia of type
lacking setae, a hair of 2 cells on the stalk; spores
dark with a continuous wing and cross-wings.
Distr. Polynesia (Fiji), Solomon Is. (Bougain-
ville), North Queensland and throughout Malesia.
Ecol. In lowland forest.
Notes. Until the work of CHING (1938) this
species and others were much confused with
Polypodium urophyllum WALL. (= Pronephrium
repandum (FEE) HOLTTUM), the present species
also with Nephrodium latifollum PRESL (=
Pronephrium menisciicarpon) and Aspidium
glandulosum BL. (Pronephrium glandulosum);
under the last-named METTENIUS also cited spe-
cimens of Sphaerostephanos peltochlamys (C.
CHR.) HOLTTUM (ZOLLINGER 2608, 2920). Spe-
cimens from New Guinea and Queensland often
have 1-2 short setae on sporangia.
4. Pronephrium gymnopteridifrons (HAYATA)
HOLTTuUM, Blumea 20 (1972) 112; C. M. Kuo, FI.
Taiwan 1 (1975) 429, pl. 150; EprE, Ferns Hong
Kong (1978) 151, f. 73.— Dryopteris gymnop-
teridifrons HAYATA, Ic. Pl. Formos. 8 (1919) 148,
fig. 75, 76.—Abacopteris gymnopteridifrons
(HAYATA) CHING, Bull. Fan Mem. Inst. Biol.
Bot. 8 (1938) 251.—Type: T. SOMA s.n. 1912,
Taiwan (isotypes TAI).
Polypodium urophyllum var. uniseriale HOOK.
Spec. Fil. 5 (1863) 10, new name for Polypodium
granulosum sensu BENTH. Fl. Hongkong (1861)
459. — Lectotype (HOLTTUM Ke 1972):
URQUHART, Hong Kong (K).
Cyclosorus pustulosus COPEL. Philip. J. Sci. 81
(1952) 37; Fern Fl. Philip. (1960) 374. — Thelyp-
teris pustulosa (COPEL.) REED, Phytologia 17
(1968) 307. — Type: Copeland 218, Lamao River,
Luzon (MICH).
Dryopteris glandulosa sensu CHR. Philip. J. Sci.
2 (1907) Bot. 205, p.p.
Abacopteris presliana sensu CHING, Bull. Fan
Mem. Inst. Biol. Bot. 8 (1938) 249, p.p.
Differs from P. asperum as follows. Caudex (at
least in some cases) long-creeping; frond smaller,
pinnae commonly to 15 x 2.5 cm (exceptionally to
25x 4cm); sori more often coalescent (in some
cases many pairs); sporangia often setiferous;
spores with many small wings.
Distr. Southern China and Malesia: Philip-
pines (Luzon).
5. Pronephrium beccarianum (CESATI) HOLT-
TUM, Blumea 20 (1972) 107; BROWNLIE, Pterid.
Fiji (1977) 256, pl. 26, f. 2.— Meniscium bec-
carianum CESATI, Rendic. Acad. Napoli 16 (1877)
27, 30.— Dryopteris cesatiana C. CHR. Ind. Fil.
(1905) 257, nom. nov. (not D. beccariana (CESATI)
C. CHR.); COPEL. Bishop Mus. Bull. 59 (1929)
48. — Phegopteris beccariana (CESATI) v.A.v.R.
Handb. (1908) 509.—Cyclosorus beccarianus
(CESATI) COPEL. Gen. Fil. (1947) 142; Philip. J.
Sci. 78 (1951) 460.— Thelypteris cesatiana (C.
CHR.) REED, Phytologia 17 (1968) 267 (not T.
beccariana (CESATI) REED).—Type: Beccari,
Andai, W. New Guinea (FI, Herb. Becc. 12727).
Goniopteris simplicifolia (J. SM. ex HOOK.)
CARR. var. vitiensis CARR. in Seem. FI. Vitiensis
(1893) 366. — Type: SEEMANN 736, Fiji (K).
Dryopteris oblanceolata COPEL. Philip. J. Sci. 9
(1914) Bot. 3.—Phegopteris oblanceolata
(COPEL.) v.A.v.R. Handb. Suppl. (1917) 320.—
Type: C. KING 394, Taupota, Papua (MICH;
SING).
Phegopteris rutteniana v.A.v.R. Bull. Jard. Bot.
Btzg II, 28 (1918) 36.— Type: KORNAssI 725,
Ceram (BO).
Caudex short-creeping. Stipe 2-10cm_ long,
usually longest in fertile fronds, near base
minutely hairy, above base hairy in groove.
Lamina simple, to 35cm long, sterile to 5cm
wide, fertile usually narrower, widest 1/3 from
short-acuminate apex, very gradually narrowed to
base, edges entire or sinuous, base sometimes
dilated or with a pair of small broadly adnate
separate leaflets; main veins 3.5-4mm apart, at
broad angle to midrib, distally upcurved; veins
9-10 pairs, almost all anastomosing, excurrent
veinlets mostly not free; lower surface between
veins pustular, sparse short hairs present on main
veins and veins, sometimes also rudimentary
scales with red end-cell; upper surface short-hairy
on midrib, sparsely on main veins. Sori exin-
dusiate, spreading along veins, those on connivent
veins often joining, sori rarely extending a little
along excurrent veinlets; sporangia bearing small
514
FLORA MALESIANA
{ser. II, vol. 1°
glands and sometimes a short seta; spores pale
with rather broad minutely erose translucent wing
and a few cross-wings.
Distr. Polynesia (Fiji), Solomon Is., in
Malesia: New Guinea and Moluccas (Ceram).
Ecol. In forest, usually near streams, to 1000 m
or a little higher.
Notes. In Fiji the basal part of the lamina is
more often strongly sinuous and there may be 3
pairs of broadly adnate basal leaflets; glands on
sporangia are larger than in New Guinea.
6. Pronephrium womersleyi HOLTTUM, Blumea
20 (1972) 108.— Cyclosorus beccarianus sensu
HOLTTUM & Roy, Blumea 13 (1965) 134.—
Type: FLOYD & WOMERSLEY NGF 6308, N.E.
New Guinea, Eastern Highlands, Goroka Subdistr.
2200 m (LAE; BM). — Fig. 14a-b.
Differs from P. beccarianum as follows. Adult
plants always with several (to 10) pairs of free +
orbicular pinnae 7-10mm long, 7-8mm_ wide;
veins in pinnae forked, the branches joining to
form a series of areoles; apical lamina smallest on
fronds with most pinnae; lower surface of midrib
and veins glabrous or nearly so; sori (on apical
lamina only, or rarely on a pinna) on lower 1-3
pairs of veins coalescent, more distal sori on basal
part of each vein; sporangia bearing small glands,
not setae.
Distr. Solomon Is. (Guadalcanal), in Malesia:
Papua New Guinea, 1400-2400 m.
Note. A plant cultivated at Kew was found to
be tetraploid (HOLTTUM & Roy l.c.).
7. Pronephrium melanophlebium (COPEL.)
HOLTTuM, Blumea 20 (1972) 108.— Dryopteris
melanophlebia COPEL. Philip. J. Sci. 6 (1911) Bot.
147. — Phegopteris —melanophlebia (COPEL.)
v.A.v.R. Handb. Suppl. (1917) 319. — Cyclosorus
melanophlebius (COPEL.) COPEL. Fern FI. Philip.
(1960) 359. — Thelypteris melanophlebia (COPEL.)
REED, Phytologia 17 (1968) 292. — Type: MER-
RILL 6959, Canlaon Volcano, Negros (isotypes B,
BO, E, K, L, NSW).
Dryopteris canescens’ var. subsimplicifolia
CHRIST, Philip. J. Sci. 2 (1907) Bot. 199. — Type:
WHITFORD 784, Luzon (not seen).
In habit similar to P. womersleyi but much
smaller. Stipe 2-3cm (sterile), to 8cm (fertile),
covered with hairs more than 0.5 mm long. Apical
lamina of sterile fronds to 5 x 1.8 cm, of fertile to
6X 1.4cm; pinnae 3 pairs, sterile 6 x 5 mm, fertile
5x4mm and more widely spaced than sterile,
oblong-ovate with midrib and 4 pairs of forked
veins, the branches anastomosing; lower surface
of costa with copious spreading pale hairs to 1 mm
long, sparse shorter hairs on veins; on upper sur-
face a few short appressed hairs between veins,
especially near margins; sori from base of veins,
+ elongate, exindusiate, those on connivent veins
sometimes meeting; sporangia bearing glands.
Distr. Malesia: Philippines (Negros, Mindanao
(ELMER 10784, Mt Apo), (?) Luzon).
Note. ELMER’s specimen from Mindanao was
distributed under the name Dryopteris canescens
var. subsimplicifolia; it agrees with CHRIST’s
brief description of that variety, but ELMER
10278, also so named, is Pronephrium granulosum.
8. Pronephrium pentaphyllum (ROSENST.)
HoLtTtruM, Blumea 20 (1972) 108.— Dryopteris
pentaphylla ROSENST. in Fedde Rep. 12 (1913)
529. — Phegopteris —_ pentaphylla (ROSENST.)
v.A.v.R. Handb. Suppl. (1917) 317. — Thelypteris
pentaphylla (ROSENST.) REED, Phytologia 17
(1968) 303.— Type: KEYSSER 186, N.E. New
Guinea, Sattelberg 1400 m (S-PA; B, BM).
Caudex long-creeping, 2-3 mm diameter when
dry, with fronds 1-2cm apart. Stipe of sterile
frond c. 10cm long, of fertile to 20cm, bearing
pale spreading hairs more than 0.5mm _ long;
fronds consisting of apical lamina and 1-2 pairs of
pinnae. Apical lamina 10-15 x 1.5-3.0cm, lobed
1/3 towards midrib, narrowed slightly to + trun-
cate base, apex acuminate; main veins c. 5mm
apart, veins to 10 pairs, 2 pairs anastomosing, 13-2
pairs passing to long sinus-membrane; lower sur-
face of midrib and veins bearing pale spreading
hairs | mm long, rather sparse slender erect hairs
between veins, no glands (but see note below);
upper surface glabrous except for midrib. Pinnae
opposite, 4-8 cm long, 1-2.5 cm wide (sterile wider
than fertile), widest above middle, lobed 1/4-1/3
towards costa; veins 4-6 pairs; pubescence as
apical lamina. Sori inframedial, spreading a little
along the veins, basal ones only sometimes
confluent; sporangia bearing 2-6 setae, on stalk a
hair of 2 cells; spores pale, opaque, with rather
broad translucent wing and _ cross-wings.
Chromosomes: n= 72 (T. G. WALKER).
Distr. Malesia: N.E. New Guinea (also New
Ireland).
Ecol. In forests, 1200-2000 m.
Notes. One specimen from Mt Oga, Western
Highlands (B. S. CROXALL 4362) differs from all
others seen in having scattered yellow glands in-
stead of hairs between veins on lower surface of
sterile fronds, and a yellow spherical gland at tip
of hair on stalk of sporangium; also in the
presence of fairly abundant short suberect hairs
between veins of upper surface.
9. Pronephrium brauseanum HOLTTUM, Blumea
20 (1972) 107. — Dryopteris canescens var. novo-
guineensis BRAUSE, Bot. Jahrb. 49 (1912) 22;
CoPEL. Philip. J. Sci. 78 (1951) 460.— Type:
SCHLECHTER 18844, N.E. New Guinea, Bismarck
Mts 1100 m (B; K, L, BISH).
Caudex short-creeping; fronds dimorphous.
Sterile fronds. Stipe 1.5-3 cm long; frond consis-
ting of terminal lamina and 1-2 pairs of pinnae
close below it; terminal lamina 5-6 x 1.7-2.1 cm,
1981]
THELYPTERIDACEAE (Holttum)
515
lobed 1/4 towards midrib, main veins 4—4.5 mm
apart, veins 5-6 pairs, 15 pairs anastomosing, 1
pair to sinus-membrane, strongly prominent on
lower surface and bearing stiff hairs to 1 mm long,
a few glands and short erect hairs present on
surface between veins; upper surface bearing
copious appressed hairs 0.3-0.4 mm long between
veins; pinnae 1.0-2.0cm long, 0.5-1.1cm wide,
sessile, edges crenate, pubescence as apical
lamina. Fertile frond. Stipe to 12cm long; pinnae
more widely spaced than sterile, not opposite, to
1.2 x 0.6cm with stalks 1 mm long, subentire with
rounded apex, lowest pinna longest; sori (mainly
on the terminal lamina) a little elongate, exin-
dusiate; sporangia copiously setose.
Distr. Malesia: Eastern New Guinea at 1100-
1700 m, in forest (5 collections).
10. Pronephrium diminutum (COPEL.) HOLT-
TUM, Blumea 20 (1972) 115.— Dryopteris
diminuta COPEL. Philip. J. Sci. 40 (1929) 298. —
Cyclosorus diminutus (COPEL.) COPEL. Gen. Fil.
(1947) 142; Fern FI. Philip. (1960) 359. — Type:
COPELAND s.n. Nov. 1911, Mindanao, San
Ramon 1200 m (MICH).
Caudex creeping, 1mm diameter; scales 2mm
long. Stipe of sterile fronds 1-3 cm long, of fertile
to 6cm, covered with short pale erect hairs; frond
consisting of apical lamina and 1-2 pairs of pinnae
(not opposite). Sterile fronds. Apical lamina 2-
4cm long, 1.5 cm wide, lobed 1/4 towards midrib;
main veins 4-4.5 mm apart, veins 3-4 pairs, 1-2
pairs anastomosing, prominent on lower surface;
lower surface of midrib bearing stiff pale hairs to
1mm long, shorter hairs on veins with a few
glands, slender erect hairs and a few glands on
surface between veins; upper surface covered
with slender appressed hairs; pinnae broadly
ovate, and slightly lobed, to 8 mm long, on stalks
more than 1mm long. Fertile fronds somewhat
smaller than sterile, with more widely-spaced
pinnae; sori exindusiate, those on lower veins
elongate and coalescing, on distal veins round;
sporangia bearing several short setae distally and
sometimes a yellow gland on the stalk.
Distr. Malesia: Philippines (Luzon, Mindanao).
Note. COPELAND made 2 collections at San
Ramon, one earlier than the type (1907). The only
other collection is by M. G. PRICE (n. 2720B)
from Mt Banahaw, Luzon, at 1000m “on rocky
slope 10m above stream”’; it only differs from the
type in slightly larger fronds with fertile pinnae to
1.1 x 0.8 cm.
11. Pronephrium micropinnatum HOLTTUM,
Blumea 20 (1972) 108. — Type: DARBYSHIRE &
HOOGLAND 8014, N.E. New Guinea, Sepik Distr.
(CANB; BM, L, LAE). — Fig. 14g-f.
Caudex short, suberect. Stipe to 6cm long,
minutely hairy on abaxial side, hairs in adaxial
groove to 1mm long. Fronds uniform, consisting
of terminal lamina and 2-3 pairs of pinnae 1-
1.5cm apart. Terminal lamina to 18 x 1.4m, nar-
rowed both to apex and base, edges crenate; main
veins 4-5 mm apart, each with 4-5 pairs of veins
of which 1) pairs anastomose, | pair passing to
sides of short sinus-membrane; lower surface of
midrib covered with pale erect hairs almost 1 mm
long, similar hairs sparse on main veins, short
erect hairs between veins; upper surface glabrous
apart from midrib; sori all medial, not elongate,
exindusiate; sporangia bearing several setae
nearly as long as the body; spores with many
small wings. Pinnae of type c. 1mm long, 2mm
wide; of plants cultivated at Kew to 53mm;
veins free; sori present on largest pinnae.
Distr. Malesia: N.E. New Guinea (Sepik and
Madang Districts, 3 collections).
Ecol. On shaded river bank (type) and sand-
stone walls of river gorge, at 30-200 m.
12. Pronephrium bulusanicum (HOLTTUM)
HOLTTUM, comb. nov. — Haplodictyum bulus-
anicum HOLTTUM, Kalikasan 2 (1973) 61.—
Type: ELMER 16585, Mt Bulusan, Luzon (K; BO,
PGE):
Stipe of sterile frond 5-6 cm, of fertile 9-14 cm
long. Lamina to 12cm long, 2.2-3.0cm wide,
short-acuminate, at the middle lobed 2/5 towards
midrib, more deeply lobed at the base which is not
narrowed, the basal lobes almost or quite free;
veins in lobes near apex of frond 7-8 pairs, not
forked, 13-2 pairs anastomosing, next pair to sides
of sinus-membrane; veins in lower lobes forked,
forming areoles on each side of costule and ad-
ditional irregular ones on each side of sinus-
membrane and below it; veins and costules prom-
inent on both surfaces; on lower surface of midrib
stiff pale hairs 1 mm long and shorter ones abun-
dant, shorter hairs present on costules and veins,
yellow glands and fine erect hairs between veins;
upper surface covered with short appressed hairs,
longer ones also on midrib and costules. Pinnae
0.9-1.5 x 0.5-0.8 cm, crenate, narrowed towards
base on basiscopic side. Sori inframedial; indusia
rather large, bearing a variable number of short
hairs and a few glands; sporangia glandular;
spores with a wing and cross-wings.
Distr. Malesia: Philippines (Luzon, Sorsogon
Province, 2 collections from type locality, | from
Mt Pulog).
13. Pronephrium heterophyllum (PRESL) HOLT-
TUM, comb. nov. — Haplodictyum heterophyllum
PRESL, Epim. Bot. (1851) 51; FEE, Gen. Fil.
(1852) 309, t. 18C, f. 2; COPEL. Fern Fl Philip.
(1960) 378; HoLtTruM, Kalikasan 2 (1973) 62.—
Aspidium heterophyllum (PRESL) HOOK. Ic. PI.
10 (1854) t. 920; COPEL. Polypod. Philip. (1905)
39. — Pleocnemia heterophylla (PRESL) v.A.v.R.
Handb. (1908) 171.— Type: CUMING 322, Samar
(PREABRES KEE):
516
FLORA MALESIANA
[ser. II, vol. 1°
Aspidium blumei KUNZE ex METT. Fil. Hort.
Lips. (1856) 94, t. 22, f. 5. — Aspidium blumei var.
subpinnata METT. Farngatt. IV (1858) 98, excl.
syn. Polypodium canescens BL. et CUMING
251. — Type: CUMING 322 (formerly at LZ, now
lost).
Aspidium canescens (BL.) CHRIST var.;
CHRIST, Farnkr. der Erde (1897) 244.— Dry-
opteris canescens (BL.) C. CHR. var.; C. CHR.
Ind. Fil. (1905) 256. — Fig. 14e-d.
Caudex short-creeping; fronds dimorphous.
Sterile fronds. Stipe 1-3 cm long, with spreading
pale hairs 1mm long; fronds consisting of ter-
minal lamina and 1-3 pairs of pinnae. Terminal
lamina to 17 X2.8cm, apex rather abruptly poin-
ted, base somewhat narrowed, middle part lobed
not quite half-way to midrib, main veins 5-6 mm
apart, veins in lobes 6-7 pairs, forked and anas-
tomosing as in lower part of lamina of P. bulus-
anicum; base of lamina more deeply lobed, with
transition to 2-3 pairs of pinnae close below it:
hairs and glands as in P. bulusanicum. Fertile
fronds. Stipe to 7cm long; terminal lamina 13 x
1.6cm, acuminate, lobed less deeply than sterile,
veins closer and fewer than in sterile; pinnae 2
pairs, rather irregularly spaced, lowest 3-4mm
long; sori in terminal lamina medial or inframedial
on the veins, not elongate; indusia with short
hairs; sporangia usually with 1 gland; spores with
translucent wing and cross-wings.
Distr. Malesia: Philippines (Samar, Southern
Luzon, Mindanao).
Ecol. On rocks in stream-beds at 200-350 m
(M. G. PRICE).
14. Pronephrium bakeri (HARR.) HOLTTUM,
comb. nov. — Nephrodium bakeri HARR. J. Linn.
Soc. Bot. 16 (1877) 29; BAk. in Hook. Ic. Pl. 17
CoPEL. Philip. J. Sci. 2 (1907) Bot. 405; v.A.v.R.
Handb. (1908) 208.— Haplodictyum _ bakeri
(HARR.) CHING, Sunyatsenia 5 (1940) 251;
HOLTTUM, Kalikasan 2 (1973) 63.— Cyclosorus
bakeri (HARR.) COPEL. Gen. Fil. (1947) 142; Fern
Fl. Philip. (1960) 358.— Thelypteris bakeri
(HARR.) REED, Phytologia 17 (1968) 262. — Type:
STEERE, Panay (MICH, K).
Aspidium canescens (BL.) CHRIST var. sensu
CHRIST, Farnkr. der Erde (1897) 244.— Dry-
opteris canescens (BL.) C. CHR. var. sensu C.
Cur. Ind. Fil. (1905) 256.
Caudex short-creeping. Stipe of sterile frond
5 cm, of fertile to 11 cm long; fronds consisting of
terminal lamina and 1 pair (rarely 2 pairs) of
pinnae; terminal lamina to 13 x2.1cm, apex ab-
ruptly short-pointed, middle part lobed 1/3
towards midrib, somewhat narrowed towards base
which is sometimes dilated; main veins to 5mm
apart, veins to 7 pairs, not forked except some-
times in basal lobes of largest fronds; lower sur-
face with pubescence and glands as in P. bulus-
anicum; pinnae opposite or not, to 107mm,
apex rounded, veins anastomosing. Sori medial on
veins of terminal lamina, sometimes also on pin-
nae, not elongate; indusia hairy; sporangia some-
times with a gland; spores with a wing and cross-
wings.
Distr. Malesia: Philippines (Luzon; Panay;
Negros Oriental; Mindanao, cf. COPELAND, not
seen).
Ecol. In forest at 500-800 m, near river (M. G.
PRICE).
Notes. CLEMENS 16530, from Isabella Prov.,
Luzon, has dimorphic fronds; sterile fronds have
an apical lamina to 2.5cm wide, fertile to 1.4 cm.
Some fronds also have a pair of very small pinnae
some distance below the normal ones.
(1886) t. 1664.— Dryopteris bakeri (HARR.)
2. Section Dimorphopteris
(TAGAWA & K. Iwats.) HoLtTTuM, Blumea 19 (1971) 36; Blumea 20 (1972)
113-121. — Dimorphopteris TAGAWA & K. Iwarts. Acta Phytotax. Geobot. 19
(1961) 8. — Thelypteris subg. Dimorphopteris K. wats. Mem. Coll. Sci. Univ.
Kyoto B, 31 (1964) 35.
Type species: Pronephrium moniliforme (TAGAWA & K. IwaTs.) HOLTTUM.
Plants of moderate size, always with a relatively short deltoid apical lamina
on fronds of adult plants, young plants rarely (as in P. menisciicarpon) having
simple fronds 10 cm or more long; sterile and fertile fronds often dimorphous,
the latter with longer stipes and smaller pinnae than the former; pinnae in most
cases distinctly auricled on the acroscopic base.
Distr. 45 species, all Malesian (P. beccarianum extending to Fiji) except P. articulatum (HOULST. &
Moore) HOLTTvuM in Ceylon, India to S. China, and P. palauense (HOSOKAWA) HOLTTUM in Palau
island.
Notes. I suggest that a probable prototype for this section is shown by the Indian species P.
1981] THELYPTERIDACEAE (Holttum) 547
articulatum (see HOLTTUM 1972: 116); the Malesian P. glandulosum differs from it in smaller size and
greater abundance of glands. The other species are mostly still smaller, with a pronounced tendency to
dimorphism, the most extremely dimorphic being the type species P. moniliforme.
This section agrees with Sphaerostephanos in the presence of spherical yellow glands, on either the
lower surface of pinnae or on indusia or sporangia, in a majority of species; it differs from Sphaeroste-
phanos in the absence of much-reduced basal pinnae and (in most species) in the pustular nature of the
lower surface of dried fronds. But there appears to be an evolutionary trend within Sphaerostephanos
towards reduction in the number of such small basal pinnae, and there are species in which some fronds
may have 1-2 pairs of reduced pinnae and others none, on the same plant. I see no sharp distinction
between such species and others which have no reduced basal pinnae. Thus on this character there
appears to be no clear separation of this section from Sphaerostephanos. The section does however give
the impression of being natural if one excludes from it species with deeply lobed non-pustular normal
pinnae (e.g. Nephrodium diversilobum PRESL); such species are here placed in Sphaerostephanos (in
some cases they have obvious relatives in that genus). Even when this is done, the present section may
still include species of two different ancestral origins: those derived from ancestors with, and without,
reduced basal pinnae. The whole problem of recognizing affinities is complex; the present arrangement is
one of convenience pending further investigation of an experimental nature.
The present arrangement includes some species placed in sect. Pronephrium in 1972; P. buwaldae
HOLTTUM, included here in 1972, is now transferred to Christella.
KEY 1O THE SPECIES
1. Sori exindusiate.
2. Pinnae not more than 3 cm long.
35 Upper surface short-appressed-hairy; som elongate << . . 2. = . . . =. . 15. PB: millarae
3. Upper surface glabrous; sori not elongate «Lee 8 4 OE ee PE OLGA: peramelense
2. Pinnae longer on mature plants.
4 RRS. pMMTAO MATING. 5 0 216 5 obo oie go ow 6 6 6 6 oo no JU einen
4. Fertile pinnae to at least 1 cm wide.
5. Surfaces between veins glabrous.
G, SoOminGR SoG 5 66 456 o coon is 6 6 4616 4 5p o o o olen IE Amini:
6. Sporangia not setiferous.
Veeeinnae.2—sicm wide: 22h". 75 “2 Slo. 30 bie PU eee ee AO ee fiemulom
7. Pinnae not over | cm wide eer Beet ce. 1 Set ile Cees Ge) Ree OE faquatiloides
5. Surfaces both short-hairy between veins.
SambinnactwGwlSipairseus’ “koe cake ie eee | ee 28S fee ee ee ye ee ee simillimum
Seekinnae 2=3 pairs: Wes Ae GI Are i ees eis B22 PS oiluwense
1. Sori indusiate.
9. Fronds simple, 10-20 cm long with cordate base away ee ees] Ae 7- 4 Pmenisciicarpon
9. Fronds pinnate or, if simple, smaller and sterile.
10. Lower surface between and/or on veins bearing sessile spherical glands.
11. Veins free aie aren). (oe ee), Saar Pee ae 23 SP akielibercii
11. Veins anastomosing.
12. Free pinnae not over 7 pairs, not over 2.0 cm wide.
13. Hairs on lower surface of costules and on upper surface between veins not appressed
24. P. amphitrichum
13. Hairs on lower surface of costules and on upper surface ate
14. Indusium bearing glands is ate 3 a aL, . . . . 25. P. clemensiae
14. Indusium bearing hairs . . . Bry... Sigh nn 2a) +: 25b. P. clemensiae var. degenerum
12. Free pinnae more numerous or wider.
15. Lower pinnae with stalks 1-2mmlong . ..... . . . . . . 26. P. glandulosum
15. Lower pinnae sessile.
16. Hairs on lower surface of costae and costules erect . . ea. See 7 Bdebile
16. Hairs on lower surface of costae and costules slender, appressed.
17. Sterile and fertile pinnae both 8-9cmlong . . . . . 25b. P. clemensiae var. degenerum
17. Sterile and fertile pinnae dimorphous, smaller.
18. Sterile pinnae c. 4.5 x 1.4 cm; rachis almost glabrous on abaxial surface 28. P. minahassae
18. Sterile pinnae c. 2.0 x 0.8 cm; rachis bearing copious erect hairs on abaxial surface
29. P. solsonicum
10. Lower surface between and on veins lacking sessile spherical glands.
19. Glands present on sporangia, in some cases setae also.
518
20. Fertile pinnae 3
FLORA MALESIANA
—4mm wide; sterile to 10 mm wide
[ser. II, vol. 1°
30. P. celebicum
20. Fertile pinnae 10mm or more wide; little dimorphism.
21. Pinnae 6-10 pairs; sporangia bearing small glands only.
22. Pinnae 2-3 cm wide
ses Pinnae not over 1 cm wide
. Pinnae 3-4 pairs; sporangia often with both ‘glands and setae
19. P. firmulum
20. P. aquatiloides
31. P. granulosum
a ae lacking glands; setae present in most cases.
23. Fertile pinnae less than 1.5 cm long; sori in | row on each side of costa
. 32. P. samarense
23. Fertile pinnae to at least 2cm long; sori in several rows.
24. Pinnae and/or aa on both young and adult plants + dilated and irregularly lobed
distally
24. Pinnae not thus dilated nor irregularly lobed.
33. P. X xiphioides
25. Basal pinnae narrowly cuneate at base both sides, not auricled; fertile pinnae not or little
narrower than sterile.
.18. P. amboinense
25. Basal pinnae otherwise, usually auricled; fertile pinnae in ) most cases narrower than sterile.
26. Indusia not very small, bearing many short hairs.
27. Upper surface of pinnae bearing short suberect hairs between veins.
28. Pinnae 7 pairs; indusia thin with some superficial hairs
28. Pinnae 15 pairs; indusia firm with stiff marginal hairs
27. Upper surface glabrous between veins.
29. Sporangia setiferous.
30. Sterile pinnae barely 1 cm wide
. 34. P. trachyphyllum
. 35. P. thysanoides
. 36. P. hosei
30. Sterile pinnae commonly 2 cm or more wide:
31. Sterile pinnae entire, to 13 x 4cm
31. Sterile pinnae crenate, to 11 x 2.5cm
29. Sporangia lacking setae.
32. Indusia firm, almost circular, with hairs in the middle
37. P. menisciicarpon
.31. P. granulosum
. 38. P. hewittii
32. Indusia thin, shrivelled when dry, hairy throughout.
33. Sterile pinnae crenate, at least distally
33. Sterile pinnae entire
26. Indusia small, with dark red cell- walls; “marginal hairs few | or none.
34. Fertile pinnae 1.5 cm or more wide
34. Fertile pinnae not more than | cm wide.
39. P. affine
37. P. menisciicarpon
40. P. borneense
35. Sterile pinnae not or little more than 1 cm wide, fertile 3-6 mm wide.
36. Pinnae thick; lower surface glabrous with thick pale prominent veins
41. P. exsculptum
36. Pinnae thin; hairs present on lower surface of costae; veins concolorous, not prominent.
37. Pinnae to 15 pairs, sterile ones 4-6 cm long, narrowly acuminate
37. Pinnae not over 10 pairs, sterile ones shorter, abruptly pointed
35. Sterile pinnae 1.5 cm or more wide, fertile 5-10 mm wide
15. Pronephrium millarae HOLTTUM, Blumea 20
(1972) 115.—Type: WOMERSLEY & MILLAR
NGF 8500, N.E. New Guinea, Morobe Distr.,
Wau-Salamaua Road 1600 m (LAE).
Caudex short-creeping; fronds dimorphous.
Sterile fronds. Stipe 7-8 cm long, covered densely
with hairs 0.1-0.2 mm long. Lamina 14cm long;
pinnae 7-8 pairs, all with stalks 1-1.5 mm long;
basal pinnae slightly reduced and deflexed, basis-
copic base rounded, acroscopic truncate and
slightly auricled. Largest pinnae 2.5-3.5 x 1cm;
base subtruncate, apex rounded to bluntly poin-
ted, edges slightly crenate near base, more dis-
tinctly so distally; costules 2.5-3.5mm apart, at
little more than 45° to costa; veins slender and
slightly prominent beneath, 3-4 pairs, 12 pairs
anastomosing, sinus-membrane not distinct; lower
surface of rachis and costa bearing stiff erect
hairs 0.2-0.4 mm long, hairs on costules and veins
shorter and somewhat antrorse, on surface be-
. 42. P. merrillii
. 43. P. rhombeum
44. P. peltatum
tween veins many appressed hairs 0.1—0.2 mm
long, no glands; upper surface throughout bearing
stiff suberect hairs 0.2-0.3 mm long. Fertile fronds.
Pinnae 2.0-2.3cm long, 0.8cm wide, lower ones
rather widely spaced, shape and pubescence as
sterile; sori exindusiate, occupying whole of basal
veins and sometimes the base of an excurrent
vein, on middle part of distal veins; sporangia
sometimes with a gland, not setiferous.
Distr. Malesia: Eastern New Guinea,
Goodenough Island.
Ecol. On Goodenough Island
moist gully” (BRASS 24636).
“common in a
16. Pronephrium peramelense HOLTTUM, Blumea
20 (1972) 115. — Type: PULLE 415, Western New
Guinea, Perameles bivouac, 1000 m (BM; L).
Caudex horizontal, 2-2.5 mm diameter, bearing
stipes 5mm or more apart; fronds not dimor-
phous. Stipe 10-20cm long, dark, short-hairy;
1981]
THELYPTERIDACEAE (Holttum)
oye)
basal scales 3-4 mm long, narrow, rather firm with
few hairs. Lamina to iScm long; apex broadly
triangular, pinnae to 8 pairs, basal ones a little
deflexed with somewhat asymmetric and some-
times slightly auricled base. Middle pinnae to 3.0 x
1.3cm; base truncate, apex rather abruptly nar-
rowed to a rounded tip, edges irregularly slightly
crenate to subentire; costules to 4 mm apart; veins 2
pairs, 15 pairs anastomosing; sinus-membrane not
evident; lower surface of rachis covered with dense
spreading dark rigid curved hairs 0.3 mm long, rest
glabrous; upper surface of costa hairy near base
only, rest of surface glabrous. Sori inframedial, on
all veins, not elongate, exindusiate; sporangia
without glands or setae.
Distr. Malesia: Western New Guinea; only
known from the type.
17. Pronephrium moniliforme (TAGAWA & K.
IWATS.) HOLTTUM, Blumea 20 (1972) 115.—
Dimorphopteris moniliformis TAGAWA & K.
IwaTs. Acta Phytotax. Geobot. 19 (1961) 8, f.
14-16. — Thelypteris moniliformis (TAGAWA &
K. Iwats.) K. Iwats. Mem. Coll. Sci. Univ.
Kyoto B, 31 (1964) 36.— Type: HARADA s.n. 30
June 1944, Halmahera (KYO).
Caudex short-creeping, 3-3.5mm_ diameter;
fronds dimorphous. Sterile fronds. Stipe 7-10 cm
long, pale, minutely hairy; lamina 15-20 cm long;
pinnae c. 15 pairs; basal pinnae deflexed, not
reduced; middle pinnae 41cm, base truncate
with acroscopic auricle, apex acute, edges ser-
rulate; costules 2.5mm apart; veins 4-5 pairs, 2
pairs anastomosing; lower surface minutely hairy.
Fertile frond. Stipe 20-24cm long; lamina 20-
24 cm long; pinnae more than 20 pairs, 4cm long,
2mm _ wide, entirely covered beneath with
sporangia (which spread to the surface between
veins); veins forming a single row of narrow
areoles with a short free vein in each crenature of
margin; sporangia bearing glands; spores with
longitudinal wing and cross-wings.
Distr. Malesia: Moluccas (Halmahera; Batjan:
ALSTON 16908, BM).
18. Pronephrium amboinense (WILLD.) HOLT-
TUM, Blumea 20 (1972) 120.— Aspidium am-
boinense WILLD. Sp. Pl. ed. 4, 5 (1810) 228. —
Thelypteris amboinensis (WILLD.) REED, Phy-
tologia 17 (1968) 259. — Type: VENTENAT, Am-
boina (B, Herb. Willd. 19751).
Aspidium canescens forma nephrodiiformis
CHRIST, Ann. Jard. Bot. Btzg 15 (1898) 131,
quoad SARASIN 975, Celebes, Palopo (BAS).
Dryopteris subconformis C. CHR. Bot. Jahrb. 66
(1933) 47.—Syntypes: KJELLBERG 1999,
Celebes, Lamasie, Palopo (BO); KJELLBERG
1360, S.E. Celebes, Labibia (BO).
Pronephrium palopense HOLTTUM, Blumea 20
(1972) 116.—Type: KJELLBERG 1999, Celebes
(BO). — Fig. 15a.
Caudex short, suberect; fronds slightly dimor-
phous. Stipe 20-40cm long (longest on fertile
fronds), minutely hairy; lamina c. 20cm long,
apex acuminate, deeply lobed with gradual tran-
sition to pinnae; free pinnae 10-15 pairs, basal pair
not reduced, narrowed to base both sides. Middle
pinnae c. 5.0X1.3cm (sterile), to 6.0 1.1cm
(fertile); base truncate and slightly auricled on
acroscopic side, apex short-acuminate (more ab-
ruptly pointed on pinnae of smaller fronds), edges
crenate to a depth of 1-2 mm; costules to 3mm
apart; veins 4-5 pairs, I-13 pairs anastomosing, >]
pairs to sides of short sinus-membrane; lower
surface of rachis, costae and costules bearing
rather sparse minute hairs, surface between veins
slightly pustular; hairs on upper surface of costae
less than 0.5mm, sparse and shorter hairs on
costules, a few minute hairs sometimes between
veins near margin. Sori medial; indusia small,
short-hairy (in some cases absent?); sporangia
with several setae; spores pale with translucent
wing and a few cross-wings.
Distr. Malesia: S.E. & Central
Moluccas (Amboina).
Ecol. At low altitudes, in forest, on stream-
bank (KJELLBERG 1360).
Notes. Four collections known. The type from
Amboina is a small frond (pinnae to 3.0 x 0.8 cm)
of which the sori are in poor condition; a few
sporangia are certainly setiferous but no indusia
are present. SARASIN 975 from Palopo agrees
closely with the type in shape of frond and pinnae
(to 4.5 1.2cm) and certainly has small indusia.
One of KJELLBERG’s specimens on_ which
CHRISTENSEN based Dryopteris subconformis (n.
1999), also from Palopo, is somewhat larger than
SARASIN’S and apparently lacks indusia; KJELL-
BERG’S second specimen (n. 1360) from S.E.
Celebes has indusia. In 1972 I confusedly cited
KJELLBERG 1999 both as type of D. subcon-
formis and of Pronephrium palopensis; I had in-
tended to cite 1360 as type of the former.
BLUME misinterpreted Aspidium amboinense
WILLD. and described under that name small
specimens of Christella subpubescens which had
many club-shaped glands on them. This error was
copied by later authors and all subsequent cita-
tions of WILLDENOW’s species are erroneous.
Celebes,
19. Pronephrium firmulum (BAK.) HOLTTUM,
Blumea 20 (1972) 116.— Polypodium firmulum
BAK. Kew Bull. (1893) 211. — Dryopteris firmula
(BAK.) C. Cue. Ind. Fil. (1905) 266; Gard. Bull.
Str. Settl. 7 (1934) 249.— Phegopteris firmula
(BAK.) v.A.v.R. Handb. (1908) 501.— Type:
C. HOSE 295, Sarawak, Mt Dulit (K).
Caudex long-creeping, 3-5 mm diameter, bear-
ing fronds 1-1.5cm apart; fronds not or little
dimorphous. Stipe to 45 cm long, glabrous; lamina
to 35cm long, apical section subentire with
widened base, small on largest fronds, pinnae to
520
FLORA MALESIANA
[ser. II, vol. 1°
12 pairs, of firm texture; basal pinnae narrowed
both sides towards the base, acroscopic base
sometimes with a slight auricle. Middle pinnae
from Mt Dulit to 11 x 2.5 cm, from Mt Kinabalu to
15 x 3.3cm, base subtruncate and distinctly auri-
cled (more on fertile than sterile pinnae), apex
acuminate, edges irregularly slightly sinuous; cos-
tules 3.5—4.5 mm apart, at less than 60°; veins 5S—7
pairs, prominent on both surfaces, almost all anas-
tomosing with continuous zig-zag excurrent veins;
lower surface of rachis covered with thick curved
hairs barely 0.5mm long, costae of Dulit speci-
mens bearing sparse short antrorse (not ap-
pressed) hairs, many such hairs on Kinabalu spe-
cimens, rest glabrous, surface between veins pus-
tular; upper surface of rachis as lower, of costae
bearing rather sparse hairs less than 0.5 mm long,
no others. Sort somewhat inframedial; indusia
none or very small, glabrous; sporangia many,
with rather small red or yellow glands, no setae;
spores not seen.
Distr. Malesia: Borneo (Sarawak; Sabah).
Ecol. On rocks by streams in forest at 250+
1000 m.
Note. Kinabalu specimens have broader and
somewhat thinner pinnae than those from Dulit;
indusia have only been seen on some sori of the
former.
20. Pronephrium aquatiloides (COPEL.) HOLT-
TUM, Blumea 20 (1972) 108.— Dryopteris aqua-
tiloides COPEL. Philip. J. Sci. 7 (1912) Bot. 59;
v.A.v.R. Handb. Suppl. (1917) 177. — Thelypteris
aquatiloides (COPEL.) REED, Phytologia 17 (1968)
260. — Type: BROOKS 9, Sarawak, Bongo Range
(MICH; BM, L).
Cyclosorus jacobsii HOLTTUM, Blumea 11
(1962) 530.— Thelypteris jacobsii (HOLTTUM)
REED, Phytologia 17 (1968) 285. — Type: JACOBS
5086, Sarawak, Mt Penrissen (L; K).
Caudex long-creeping, 2-3 mm diameter; fronds
c. 5mm apart. Stipe 15-30cm long, minutely
hairy; lamina to 27cm long, apex pinna-like, pin-
nae 6-8 pairs; basal pinnae narrowed towards base
both sides, with stalks to 4mm long. Middle pin-
nae 9-11x1.0cm; base auricled on acroscopic
side; apex acuminate with rounded tip 2 mm wide;
edges almost entire; costules 3mm apart; veins
3-4 pairs, anastomosing to form a zig-zag excur-
rent vein, slightly prominent; hairs on lower sur-
face of rachis thick, brown, curved, 0.5 mm long,
on costae same length, slender, antrorsely ap-
pressed, no others hairs; surface between veins
pustular; upper surface of rachis hairy as lower,
costal hairs shorter, no others hairs. Sori medial,
basal ones sometimes coalescent; indusia very
small, glabrous or glandular; sporangia bearing red
glands; spores not seen.
Distr. Malesia: Borneo (W. Sarawak).
Ecol. On rocks by streams in forest at 300-
1000 m.
21. Pronephrium simillimum (C. CHR.) HOLT-
TUM, Blumea 20 (1972) 116.— Dryopteris simil-
lima C. CHR. Ind. Fil. (1905) 292, new name for
Nephrodium simulans BAK. J. Bot. 26 (1888) 325,
non BAK. 1874; v.A.v.R. Handb. (1908) 223. —
Thelypteris simillima (C. CHR.) K. IwaTs. Acta
Phytotax. Geobot. 21 (1965) 169.— Type: G. F.
HOSE 231, Sarawak, limestone hills (K).
Caudex short-creeping to suberect; fronds
hardly dimorphous. Stipe 12-25(—45)cm_ long,
minutely hairy, basal scales dark, narrow, rigid, to
8mm long with stiff pale hairs 0.2mm _ long;
lamina very firm, 25-40 cm long, apex deltoid and
deeply lobed, pinnae 12-16 pairs; basal pinnae
slightly reduced, narrowed to base on basiscopic
side. Largest pinnae of type 4.7 x 1.6cm (width
above base), largest seen 6.0 x 1.7 cm; base trun-
cate and distinctly auricled on acroscopic side;
apex abruptly narrowed, rounded or broadly
pointed; edges lobed to depth of 1.5-2mm (on
young plants almost entire), lobes oblique, roun-
ded, more or less distinctly dentate at ends of
veins; costules to 4mm apart, at c. 50° to costa;
veins 5-6 pairs, slightly prominent both sides, 13-2
pairs anastomosing, | pair to sides of sinus-mem-
brane; lower surface of rachis covered with stiffly
erect pale hairs 0.2-0.4mm long, similar hairs
0.1-0.2 mm long on costae, costules, veins and on
surface between veins; upper surface similarly
hairy, hairs between veins more dense than on
lower surface. Sori small, medial, exindusiate;
sporangia with 6-8 slender setae; spores with
rather broad finely erose translucent wing and
cross-wings.
Distr. Malesia: Borneo; S.W. Celebes (BROOKS
16853, Maros).
Ecol. On limestone, 0-1700 m.
Notes. One specimen from a shaded place at
entrance to a cave has thinner pinnae more deeply
lobed than above described, to 2.2 cm wide, cos-
tules 5mm apart, pinna-lobes distinctly dentate at
all vein-ends, veins often forked, only 1 pair
anastomosing. A specimen from 1700m on G.
Mulu is small, with 5 pairs of free pinnae, largest
3.0 x 1.3. cm, hairs on lower surface of rachis and
costae more than 0.5 mm long.
22. Pronephrium giluwense HOLTTUM, sp. nov.
Pinnae paucijugatae, inferiores leviter redactae
auriculatae stipitataeque, pinnae majores 5 cm
longae, 1.5 cm latae, vix dimorphae; pagina inter
venas utraque pilis brevibus vestita; sori exin-
dusiati, sporangia setis 3-6 praedita.— Type:
WOODHAMS 138, Papua, Mt Giluwe, 2400 m, cult.
Hort. Kew. (K).
Caudex short-creeping; stipe to 14cm long,
short-hairy; lamina thin, to 16 cm long consisting of
apical section 7x2.4cm lobed as pinnae and 3
pairs of pinnae; basal pinnae 2.5 x 1.4cm, rather
strongly auricled, with stalks 1.5 mm long. Largest
pinnae 5.0 x 1.5 cm with stalks 1 mm; base broadly
1981]
THELYPTERIDACEAE (Holttum)
324
cuneate and slightly auricled on acroscopic side,
apex abruptly short-acuminate; edges lobed 1/3
towards costa; costules 4mm apart at 45° to costa;
veins slender, slightly prominent, 5-6 pairs, | pair
anastomosing, 1-1} pairs to sides of sinus-mem-
brane; lower surface of rachis and costae bearing
spreading hairs of varied length to 0.8 mm, shorter
slender hairs on costules, veins and surface be-
tween them; upper surface covered with suberect
hairs 0.2-0.3mm long, longer hairs present on
costae and scattered on costules and veins. Sori
medial, lowest supramedial, exindusiate; sporan-
gia bearing 3-6 rather long setae; spores with a
longitudinal wing and cross-wings.
Distr. Malesia: Papua New Guinea. The only
other collection is CLEMENS 12110 D, from a
branch of the Buso River, Morobe District, at
1500-1800 m (MICH); this is a less mature plant
with longer apical lamina than the type; it differs
also in having a few glands on the lower surface
of pinnae.
Note. The type plant perhaps did not attain its
full size before dying. Possibly this species should
be transferred to Sphaerostephanos but it does
not match any known species in that genus.
23. Pronephrium kjellbergii HoLTrumM, Blumea
20 (1972) 117.—KJELLBERG 2638, Central
Celebes, Mt Porema 1400 m (BO).
Caudex short-creeping; fronds subdimorphous.
Stipe of sterile frond 3-4 cm long, of fertile 10cm,
short-hairy, basal scales 4x 1mm bearing short
hairs; lamina 8cm long, thin; pinnae 7 pairs,
lower sterile ones sometimes a little reduced, fer-
tile not; apical lamina deltoid and deeply lobed.
Largest pinnae 1.5<0.6cm (sterile) 1.0 x 0.6cm
(fertile); base truncate and a little dilated on
acroscopic side; apex blunt; edges crenate; cos-
tules 2.5mm apart; veins 2 pairs (sometimes 3
pairs in basal acroscopic lobe), all free; lower
surface of rachis, costae and costules bearing
sparse hairs 0.5mm long, many glands on and
between veins; upper surface throughout bearing
short antrorse hairs with longer ones scattered on
costules and veins. Sori near bases of veins; indusia
bearing a few hairs and glands; sporangia some-
times with a gland.
Distr. Malesia: Central Celebes, 1400 m. Only
known from the type.
Note. This specimen was named Dryopteris
urdanetensis COPEL. by CHRISTENSEN (Bot.
Jahrb. 66, 1933, 48) but differs from that species in
having a much shorter apical lamina and no much-
reduced basal pinnae.
24. Pronephrium amphitrichum HOLTTUM, sp.
nov.
P. clemensiae (Copel.) Holttum affinis, differt:
pinnis multo minoribus; pilis costarum costu-
larumque subtus patentibus, pilis inter venas
paginae superioris non appressis; soris venarum
inferiorum elongatis. —Type: EDANO BS 75790,
Luzon, Camarines Sur, Mt Potianay (MICH;
NY).
Caudex suberect; fronds subdimorphous. Stipe
3 cm long (sterile) 6-7 cm (fertile), short-hairy, basal
scales 3 mm long, narrow; lamina 5 cm long; pinnae
4-5 pairs; basal pinnae slightly reduced, much
narrowed towards base on basiscopic side. Largest
pinnae to 1.8 x 0.9 cm (sterile) 1.5 x 0.8 cm (fertile);
base truncate and slightly auricled on acroscopic
side, narrowed and rounded on basiscopic; apex
abruptly blunt-pointed; edges crenate to depth of
1mm; costules to 3mm apart at little over 45° to
costa; veins 3 pairs, | pair anastomosing; lower
surface of rachis bearing thick pale spreading hairs
0.5 mm long, hairs on costae and costules similar but
shorter, abundant slender erect hairs and a few
glands on surface between veins; hairs on upper
surface of rachis thick, curved, to | mm long, on
costae 0.2-0.3 mm, on costules and veins scattered
longer hairs, between veins copious suberect hairs
0.2 mm long. Sori inframedial to medial, lower ones
elongate; indusium small with short stiff hairs;
sporangia sometimes with a gland.
Distr. Malesia: Philippines
known from the type.
(Luzon). Only
25. Pronephrium clemensiae (COPEL.) HOLTTUM,
Blumea 20 (1972) 118.— Dryopteris clemensiae
CoPEL. Philip. J. Sci. 46 (1931) 213. — Cyclosorus
clemensiae (COPEL.) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 10 (1941) 243; COPEL. Fern FI. Philip.
(1960) 369.— Thelypteris clemensiae (COPEL.)
REED, Phytologia 17 (1968) 268.— Type:
CLEMENS 16490, Luzon, Isabella Prov., Mt Moises
(UC; US).
a. var. clemensiae
Caudex short-creeping; fronds subdimorphous.
Stipe 8-10cm long (sterile), 16cm (fertile), hairy
as rachis distally, basal scales to 3 x 1 mm, firm;
lamina 13cm long, texture firm, apex broadly
triangular and deeply lobed, pinnae 5 pairs, basal
pinnae not reduced, narrowed towards base of
basiscopic side. Largest pinnae 4.3 1.8cm
(sterile) 3.2x1.5cm (fertile); base subequally
broadly cuneate; apex abruptly narrowed to acute
tip; edges lobed to depth of 2mm; costules 3.5—
4mm apart, at 45° to costa; veins 4-5 pairs, prom-
inent both sides, 1-1; pairs anastomosing; lower
surface of rachis covered with coarse erect hairs,
hairs on costae and costules shorter and antror-
sely appressed, glands abundant on lower surface
generally; hairs on upper surface of costae, cos-
tules and veins antrorse as lower surface, between
veins rather sparse appressed hairs. Sori medial;
indusia rigid, dark, with yellow glands and some-
times a few hairs; sporangia often with a yellow
gland.
Distr. Malesia:
known from the type.
Philippines (Luzon); only
522
b. var. degenerum (CHRIST) HOLTTUM, comb.
nov.— Dryopteris canescens var. degenera
CHRIST, Philip. J. Sci. 2 (1907) Bot. 199. — Phe-
gopteris canescens var. degenera (CHRIST) v.A.v.R.
Handb. (1908) 507. — Cyclosorus degener (CHRIST)
CopeEL. Fern FI. Philip. (1960) 356. — Thelypteris
degenera (CHRIST) REED, Phytologia 17 (1968)
271. — Lectotype (HOLTTUM 1972): LOHER s.n.
March 1906, Luzon, Montalban (P).
Differs from var. clemensiae: indusia densely
setose; sporangia bearing glands or setae. Spores
have a complete translucent wing.
Distr. Malesia: Philippines, Luzon (C. B.
ROBINSON BS 9455, Tayabas Prov. Infanta;
RAMOS BS 1791, Rizal Prov.; M. G. PRICE 2824,
Zambales Prov., Mts above Palauig at 1000 m).
Notes. PRICE’s specimen is much larger than
the others (pinnae to 8.5 x 2.0cm) and has rather
sparse glands on lower surface but agrees in other
respects. I designate the LOHER collection from
Montalban as lectotype because I did not find the
other syntype, also collected by LOHER, at Paris.
26. Pronephrium glandulosum (BL.) HOLTTUM,
Blumea 20 (1972) 118.— Aspidium glandulosum
BL. En. Pl. Jav. (1828) 144; Metr. Farngatt. IV
(1858) 111, quoad descr. tantum.— Nephrodium
glandulosum (BL.) J. SM. in Hook. J. Bot. 3 (1841)
411, nomen tantum; PRESL, Epim. Bot. (1851) 45,
excl. pl. Zoll.; HOOK. Spec. Fil. 4 (1862) 76, excl.
syn. omn. praeter BI.; RActiB. Fl. Btzg 1 (1898)
185.— Abacopteris glandulosa (BL.) FEE, Gen.
Fil. (1852) 310. — Dryopteris glandulosa (BL.) O.
KTZE, Rev. Gen. Pl. 2 (1891) 812; v.A.v.R.
Handb. (1908) 215.— Dryopteris malayensis C.
CHR. Dansk Vid. Selsk. Skr. VII, 10 (1913) 171,
nom. nov. illegit.; BACKER & POSTH. Varenfl.
Java (1939) 59.—Cyclosorus glandulosus (BL.)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938)
227; HOLTTUM, Rev. FI. Malaya 2 (1955) 278, f.
160. — Thelypteris malayensis (C. CHR.) REED,
Phytologia 17 (1968) 291.—Type: BLUME s.n.,
W. Java (L, n. 908, 337-89).
Dryopteris iridescens v.A.v.R. Bull. Jard. Bot.
Btzg II, 11 (1913) 11; Handb. Suppl. (1917) 176. —
Thelypteris iridescens (v.A.v.R.) REED, Phy-
tologia 17 (1968) 285.—Type: MATTHEW S517,
Sumatra, Padang Panjang (BO).
Dryopteris excrescens COPEL. Univ. Cal. Publ.
Bot. 14 (1929) 374. — Type: RAHMAT SI TOROES
142, Sumatra (UC; L).
Dryopteris bartlettii COPEL. ibid. l.c.—
Thelypteris bartlettii (COPEL.) REED, Phytologia
17 (1968) 263. — Type: BARTLETT 6692, Sumatra,
Asahan (UC).
Nephrodium lineatum sensu. BEDD. Handb.
Suppl. (1892) 71, excl. syn. Aspidium affine BL.
Caudex short-creeping; fronds dimorphous.
Sterile fronds. Stipe 20-30cm long, minutely
hairy; lamina 30-40cm long; pinnae 12 pairs;
basal pinnae narrowed at base, more on basiscopic
FLORA MALESIANA
[ser. II, vol. 1°
than acroscopic side, with stalks 2mm long. Lar-
gest pinnae commonly to 12 x 2.5 cm; base trun-
cate, auricled on acroscopic side, apex abruptly
short-acuminate; edges obliquely lobed to a depth
of 1-2 mm or sometimes more deeply; costules to
5 mm apart; veins to 8 pairs, 2-4 pairs anastomos-
ing (according to depth of lobing), 15-2 pairs to
sides of sinus-membrane; lower surface of rachis
covered with thick curved hairs more than 0.5 mm
long, similar but shorter hairs on costae, very
short and sparse on costules and veins, surface
between veins pustular, bearing many glands and
sometimes very short erect hairs; upper surface
covered throughout more or less closely with fine
appressed hairs 0.2-0.3mm long. Fertile fronds.
Stipe to 50cm long; pinnae more widely spaced
than sterile, commonly to 7 x 1.5 cm, edges shal-
lowly crenate; sori medial, lower ones at least
somewhat elongate along veins; indusia thin bear-
ing many very short hairs; sporangia with glands
on body and a hair with glandular tip on stalk;
spores with a complete wing and cross-wings.
Distr. Malesia: Java, Sumatra, Malaya.
Ecol. Lowland, up to 800m, in forest near
streams.
Notes. The lobing of pinnae is variable. One
Java specimen at Kew (coll. LOBB) has both ster-
ile and fertile pinnae almost entire. A specimen
collected by KURZ on G. Salak (PR) has fertile
pinnae 7 x 1.7 cm, lobed fully half-way to costa.
The name Aspidium glandulosum BL. was
variously misinterpreted by J. SMITH, KUNZE
and HOOKER, who cited specimens of other spe-
cies; these citations were copied by METTENIUS
and PRESL who both gave good descriptions.
Apart from the presence of copious glands on
lower surface and appressed hairs on the upper,
this species is very similar to A. affine BL., with
which BEDDOME confused it under the name
Nephrodium lineatum.
27. Pronephrium debile (BAK.) HOLTTUM, Blu-
mea 20 (1972) 118. — Nephrodium debile BAK. J.
Bot. 18 (1880) 212. — Dryopteris pseudoreptans C.
Cure. Ind. Fil. (1905) 286, nom. nov. (not D. debilis
(METT.) C. CHR.); v.A.v.R. Handb. (1908) 223. —
Thelypteris pseudoreptans (C. CHR.) REED, Phy-
tologia 17 (1968) 306.— Type: BECCARI 433,
Sumatra, G. Singgalang 1700 m (K).
Nephrodium pilosiusculum RAciB. FI. Btzg 1
(1898) 189. — Dryopteris pilosiuscula (RACIB.) C.
Cur. Ind. Fil. (1905) 284; v.A.v.R. Handb. (1908)
215; BACKER & POSTH. Varenfl. Java (1939) 65,
excl. syn. Aspidium pilosiusculum METT. —
Thelypteris pilosiuscula (RACIB.) REED, Phy-
tologia 17 (1968) 304. — Type: RACIBORSKI, Java,
G. Salak, Tjiapoes (BO; K, L, P).
Caudex short-creeping; fronds slightly dimor-
phous. Stipe 15-20 cm long, glabrous except base
which is covered with minute hairs and firm seti-
ferous scales 2x 1mm; lamina 15-25cm long,
1981]
THELYPTERIDACEAE (Holttum)
323
firm in texture; apical lamina narrowly acuminate;
pinnae 10-15 pairs; basal pinnae slightly narrowed
at base on basiscopic side, sessile. Largest pinnae
4.5x1.5cm (sterile) 3.0 1.0cm (fertile); base
truncate and slightly auricled; apex abruptly short-
acuminate with rounded tip; edges crenate or in
the largest sterile pinnae lobed to a depth of
2-3 mm; costules 3-4 mm apart; veins 3-4 pairs, |
pair anastomosing, next pair to sides of sinus-
membrane; lower surface of rachis covered with
thick spreading brown hairs 0.7 mm or more long
and shorter pale ones, hairs on costae and costules
dense, pale, 0.4mm long, somewhat antrorse,
hairs on veins shorter with some yellow glands,
surface between veins bearing many short erect
hairs and a few glands (especially distal on the
lobes); upper surface of rachis bearing erect thick
brown hairs Imm long, pinnae covered closely
with appressed hairs 0.5mm long. Sori medial;
indusia covered with short hairs, 1-2 glands also
sometimes present; sporangia bearing glands.
Distr. Malesia: Sumatra, Java, Flores.
Ecol. On wet rocks by streams at 1400-1800 m.
Note. BECCARI’s specimen from G. Singgalang
has more deeply lobed pinnae than RACIBORSKI’s
from Tjiapoes but differs little in other respects.
RACIBORSKI cited a specimen of ZIPPELIUS (to
whom he attributed the name) as well as his own;
I have not found the former and take RACIBOR-
SKI’s own, which he described, to be the type.
Confusion has been caused because METTENIUS
independently published the name Aspidium
pilosiusculum Zipp. as a new name for Gym-
nogramme appendiculata BL. (here transferred to
Sphaerostephanos). Both species were collected
by RACIBORSKI at Tjiapoes (Mt Salak, W. Java).
28. Pronephrium minahassae HOLTTUM, Blumea
20 (1972) 119.—Type: ALSTON 16410, N.
Celebes, G. Tetawiran (BM).
Caudex short, suberect; fronds dimorphous.
Sterile fronds. Stipe to 15 cm long, minutely hairy,
basal scales small. Lamina to 25 cm long, texture
thin; apical lamina pinna-like, to 10cm long; pin-
nae to 10 pairs, basal pair more remote and nar-
rowed to base on basiscopic side, widest in
middle. Largest pinnae 4.5cm long, 1.5cm wide
above base; base truncate, slightly auricled espe-
cially on lower pinnae; apex abruptly short-poin-
ted, edges lobed to a depth of 1.5-2 mm; costules
3.5-4mm apart; veins 6-7 pairs, 1) pairs anas-
tomosing, | pair to sides of sinus-membrane;
lower surface of rachis glabrous or sparsely hairy,
of costae, costules and veins covered with slender
appressed hairs, glands abundant throughout; upper
surface covered with slender appressed hairs.
Fertile fronds. Pinnae 2.8 cm long, 0.7-0.9 cm wide
above base, lobed as sterile and similarly hairy and
glandular; sori at maturity covering whole surface;
indusia hairy; sporangia bearing glands; spores with
a translucent wing and cross-wings.
Distr. Malesia: N. Celebes; only known from
the type.
29. Pronephrium solsonicum HOLTTUM, Kalik-
asan 3 (1974) 197.—Type: M. G. PRICE 2903,
Luzon, Ilocos Norte, Solsona, 1100 m, in forest
(K; PNH).
Caudex short-creeping; fronds dimorphous.
Sterile fronds. Stipe 5-6 cm long, short-hairy near
base, basal scales thin, 3mm long. Lamina to
15 cm long; apex 8 cm long, deeply lobed; pinnae 9
pairs, basal pair slightly reduced and narrowed to
base on basiscopic side. Largest pinnae 2.1
0.8 cm; base truncate and slightly auricled; apex
obtuse; edges lobed to a depth of 1 mm; costules
2.5mm apart; veins slender, 3 pairs, basal pair
anastomosing, next pair to edge; lower surface of
rachis bearing coarse spreading pale hairs | mm
long, hairs on costae and costules much shorter,
slender, appressed, many glands on surface be-
tween veins; upper surface of rachis as lower,
hairs on costae 0.5mm long, similar hairs scat-
tered on costules and veins, no others. Fertile
fronds. Stipe 12-20cm long; pinnae to 1.4
0.7 cm, edges subentire or slightly crenate; sori
near costules, indusia large, glabrous; sporangia
bearing glands.
Distr. Malesia:
known from the type.
Philippines (Luzon); only
30. Pronephrium celebicum (BAK.) HOLTTUM,
Blumea 20 (1972) 119.— Acrostichum celebicum
BAK. Kew Bull. (1901) 741.— Leptochilus cele-
bicus (BAK.) C. CHR. Ind. Fil. (1905) 384;
v.A.v.R. Handb. (1908) 741. — Dryopteris celebica
(BAK.) COPEL. Philip. J. Sci. 37 (1928) 410.—
Thelypteris celebica (BAK.) REED, Phytologia 17
(1968) 267. — Lectotype (HOLTTUM 1972): DE LA
SAVINIERRE 61, N. Celebes (K; P).
Aspidium canescens forma _ acrostichoides
CHRIST, Ann. Jard. Bot. Btzg 15 (1898) 132.—
Dryopteris acrostichoides (CHRIST) v.A.v.R.
Handb. Suppl. (1917) Corr. 49, non O. KTZE 1891
excl. var. rhombea & var. lanceola. — Lectotype
(HOLTTUM 1972): KOORDERS 17153, N. Celebes
(BOs):
(2?) Meniscium hosei var. sumbensis v.A.v.R.
Bull. Dép. Agr. Ind. Néerl. 21 (1908) 7. — Phegop-
teris hosei var. sumbensis v.A.v.R. Handb. (1908)
510.— Type: TEYSMANN 10693, Sumba (BO?;
not seen). — Fig. 15b-—d.
Caudex short-creeping; fronds dimorphous.
Sterile fronds. Stipe 6-15 cm long, pale, minutely
hairy or glabrescent, basal scales thin, narrow,
3mm long. Lamina thin, 15-18cm long; apex
acuminate and deeply lobed; pinnae 8-10 pairs,
basal pinnae narrowed to base on basiscopic side,
sessile or with a very short stalk. Largest pinnae
4.2 1.2cm (width above base); base truncate,
strongly auricled on acroscopic side; apex short-
acuminate; edges serrate-crenate to a depth of
524
FLORA MALESIANA
1-1.5mm; costules 2.5mm apart; veins slender
and little prominent, 4-5 pairs, 2 pairs anastomos-
ing, sinus-membrane hardly evident; lower surface
of rachis bearing hairs 0.3 mm long, pale, sometimes
sparse, on costae and costules hairs much shorter
and always sparse, surfaces between veins pus-
tular; hairs on upper surface of rachis thick, to
0.5 mm long, on costae 0.2 mm, rather sparse, no
others. Fertile fronds. Stipe 20-30 cm long; lamina
to 25cm; pinnae widely spaced, 2.5-4.0 cm long,
2-4 mm wide above auricled base, edges crenate;
veins 3 pairs, basal pair only anastomosing; lower
surface covered with sori (also upper surface near
apex?); indusia very small with a few hairs;
sporangia of type bearing both glands and short
setae, of Curtis specimen (also cited by Baker)
glands only; spores light brown, with a translucent
wing and cross-wings.
Distr. Malesia: Celebes, Moluccas (Ceram,
Tenimber Is.), West New Guinea (Vogelkop), and
? Lesser Sunda Is. (Sumba).
Notes. SARASIN 1022, from Central Celebes,
cited by CHRIST with those of KOORDERS, has
sterile pinnae to 6 x 1.5 cm and sporangia strongly
setiferous. v.A.v.R.’s description of D. acrosti-
choides was based on KOORDERS’S specimens at
Bogor named by CHRIST; these were not cited by
CHRIST in 1898; I have selected KOORDERS
17153 as the type of v.A.v.R.’s name.
31. Pronephrium granulosum (PRESL) HOLTTUM,
Blumea 20 (1972) 119. — Polypodium granulosum
PRESL, Rel. Haenk. (1825) 24, pl. 4, f. 2; HOLT-
TUM, Novit. Bot. Inst. Bot. Univ. Carol. Prag.
1968 (1969) 19. — Dryopteris granulosa (PRESL) C.
Cure. Ind. Fil. (1905) 269; CHRIST, Philip. J. Sci. 2
(1907) Bot. 217. — Phegopteris granulosa (PRESL)
v.A.v.R. Handb. (1908) 503, p.p.— Cyclosorus
granulosus (PRESL) COPEL. Fern FI. Philip. (1960)
373, p.p. — Thelypteris granulosa (PRESL) REED,
Phytologia 17 (1968) 280.—Type: HAENKE,
Luzon (PRC; specimen at W is marked HAENKE
98).
Dryopteris chamaeotaria CHRIST, Philip. J. Sci.
2 (1907) Bot. 203.— Phegopteris chamaeotaria
(CHRIST) v.A.v.R. Handb. (1908) 505.— Thelyp-
teris chamaeotaria (CHRIST) REED, Phytologia 17
(1968) 267.—Lectotype (COPELAND 1960):
WHITFORD 1369, Luzon, Mt Mariveles (not found
at MICH, P, US); Neotype (HOLTTUM 1972):
ELMER 6970, same locality (K).
Dryopteris maquilingensis COPEL. Philip. J.
Sci. 56 (1935) 103, pl. 8. — Type: COPELAND s.n.
Nov. 1932, Luzon, Mt Makiling (formerly in
PNH); Neotype (HOLTTUM 1972): ELMER 18169,
same locality (K; BISH, G, L).
Cyclosorus subdimorphus COPEL. Philip. J. Sci.
81 (1952) 38; Fern Fl. Philip. (1960) 374.—
Thelypteris palawanensis REED, Phytologia 17
(1968) 300, nom. nov.—Type: EDANO PNH
13925, Palawan (MICH). — Fig. 15m-n.
[ser. II, vol. 1°
Caudex short to long-creeping, 3 mm diameter;
fronds close or 1l-2cm apart, subdimorphous.
Stipe 15-20cm long (sterile) 30-45 cm (fertile);
lamina of mature plants 15-24cm long, thin but
firm, consisting of apical section 8-15cm long,
1.5-3.0cm wide, and 2-4 pairs of pinnae; lowest
pinnae stalked 1-2mm, widest at about the
middle, base asymmetric and + auricled. Middle
pinnae to 11 x2.5cm (sterile; fertile smaller, to
9x2.2cm); base truncate and slightly auricled;
apex acuminate (narrowly on longest pinnae);
edges crenate to depth of 1 mm (entire on pinnae
of young plants); costules to 4mm part; veins
5-6(—8) pairs, slender and slightly prominent, 2-5
pairs anastomosing (according to width of pinnae);
lower surface of rachis bearing a variable number
of coarse pale hairs to 0.8mm long; hairs on
costae sparse and much shorter, surface between
veins pustular and sometimes with very short erect
hairs; upper surface of rachis covered with thick
pale hairs 1mm long, costal hairs rather sparse,
0.3mm long, no other hairs. Sori inframedial;
indusia small, soon caducous, dark red with short
hairs on surface and margin; sporangia bearing
glands and also short setae; spores with trans-
lucent wing and cross-wings.
Distr. Malesia: Philippines (Luzon, Palawan,
Balabac, Panay, Mindanao).
Ecol. In forest at low altitudes, to 500 m.
Notes. One frond in HAENKE’s collection at
Prague has two pinnae widened and irregularly
deeply lobed distally; PRESL named this var.
lobata. This feature may indicate hybridity; it
needs experimental investigation.
Young plants have much smaller pinnae than
above described; the smallest fertile pinna seen
measures 4.5 1.2cm. A specimen from Balabac
(J. B. STEERE) has no glands on sporangia, and
pinnae crenate to a depth of 2 mm; otherwise it is
very near the type. A specimen from Mindanao
has very narrow pinnae, the largest fertile one
5.0 x 0.8 cm.
32. Pronephrium samarense (COPEL.) HOLTTUM,
Blumea 20 (1972) 119.—Cyclosorus samarensis
CoPEL. Philip. J. Sci. 81 (1952) 35. — Thelypteris
samarensis (COPEL.) REED, Phytologia 17 (1968)
311.— Type: GACHALIAN PNH _ 15240, Samar
(MICH; 15220 at SING).
Caudex short-creeping, 2mm diameter; fronds
dimorphous. Sterile fronds. Stipe 3-5 cm long,
basal scales to 2mm long, pale stiff hairs 0.7 mm
long in the groove. Lamina 5-8cm long; apex
narrowly triangular, 2.5cm long, lobed at base
only; pinnae to 5 pairs, stalked 1mm, lowest a
little reduced and narrowed to base on basiscopic
side. Largest pinnae 2-3 cm long, 0.8-1.0 cm wide;
base subtruncate, slightly auricled on acroscopic
side, rounded on basiscopic; apex obtusely poin-
ted to rounded; edges entire; costules 2 mm apart;
veins mostly 3 on basiscopic side of costule, 2 on
1981]
THELYPTERIDACEAE (Holttum)
D2
acroscopic, anastomosing to form one series of
areoles with irregular additional ones; hairs on
lower surface of rachis 0.3 mm long, on costae and
costules shorter and sparse, no other hairs, sur-
face between veins pustular; upper surface of
rachis and costae similarly hairy, a few minute
hairs present near margin. Fertile fronds. Stipe
8-18 cm long; lower pinnae widely spaced, largest
pinnae to 1.2x0.5cm, shaped and stalked as
sterile; costules simple or forked; where forked
the branches anastomosing near margin; sori in
one row on each side of costa with additional ones
present in basal acroscopic lobe; indusia small,
dark red, bearing short hairs; sporangia with 1 or 2
short setae.
Distr. Malesia: Philippines (Samar, 3 collec-
tions).
Ecol. Streambanks at c. 100 m.
33. Pronephrium x xiphioides (CHRIST) HOLT-
TUM, Blumea 20 (1972) 119.— Dryopteris
xiphioides CHRIST, Philip. J. Sci. 2 (1907) Bot.
201. — Phegopteris xiphioides (CHRIST) v.A.v.R.
Handb. (1908) 501.—Cyclosorus xiphioides
(CHRIST) COPEL. Gen. Fil. (1947) 143; Fern FI.
Philip. (1960) 349.— Thelypteris xiphioides
(CHRIST) REED, Phytologia 17 (1968) 324. —
Type: COPELAND s.n. April 1905, Mindanao, San
Ramon (MICH).
Cyclosorus edanyoi COPEL. Philip. J. Sci. 81
(1952) 37; Fern Fl. Philip. (1960) 370. — Thelyp-
teris edanyoi (COPEL.) REED, Phytologia 17
(1968) 274. — Type: Edano BS 46065, Panay (UC).
Dryopteris diversiloba sensu CHRIST, Philip. J.
Sci. 2 (1907) Bot. 199, excl. var. acrostichoides.
Dryopteris rhombea sensu COPEL. Philip. J.
Sci. 56 (1935) 102, pl. 6, p.p. — Cyclosorus rhom-
beus sensu COPEL. Fern FI. Philip. (1960) 357,
p.p.
Plants with a frond-form intermediate between
P. granulosum and P. rhombeum, with pinnae of
irregular shape; young plants with lamina 4-5 cm
long often with 2-3 pairs of pinnae; adult plants
mostly with 4-6 pairs of pinnae (rarely 8 pairs); in
almost all cases some or all pinnae, often also the
apical lamina of the frond, irregularly dilated and
irregularly + deeply lobed distally (see
COPELAND 1935, line drawings on pl. 6); at least
the lower pinnae + stalked as in P. granulosum;
indusia small with a few short hairs; sporangia
always setiferous.
Distr. Malesia: Philippines, widely.
Notes. These plants look like a hybrid swarm,
which needs experimental cytotaxonomic in-
vestigation. The type collection of P. granulosum
includes a frond with one pinna dilated and deeply
lobed at the apex, indicating perhaps a slight
degree of hybridity. Both parent species have very
similar indusia; the sporangia of P. granulosum
have glands on sporangia, but the hybrids do not. I
have regarded this as an important distinction.
The type of D. xiphioides is the only specimen
seen in which no pinnae are dilated distally.
34. Pronephrium trachyphyllum HOLTTUM, sp.
nov.
Stipes frondis fertilis 23cm longus; lamina
14 cm longa pinnis 7-jugatis laminaque 5 cm longa
constituta; pinnae usque 3.00.8 cm, crenatae,
eglandulosae, supra inter venas pilis minutis
suberectis multis vestitae; sori inframediales, non
elongati; indusia parva tenuia atrorubra, pilis
multis 0.2-0.3 mm longis vestita; sporangia setis
pluribus praedita.— Type: A. B. COLINA 772,
Mindanao, Surigao (K ex CAHP).
Caudex short-creeping, bearing small sterile and
much larger fertile fronds. Sterile fronds. Stipe
1.5cm long; lamina 4cm long, apex deltoid and
lobed, pinnae 3-4 pairs, to 1.3 x 0.5 cm, subentire.
Fertile fronds. Stipe 10-23cm long, minutely
hairy; basal scales thin, narrow, to 3mm long.
Lamina 8-14cm long; apex 3—4cm long, rather
deeply lobed throughout with some _ subapical
lobes irregularly elongate; pinnae to 7 pairs; basal
pinnae of largest frond not reduced but more
widely spaced, not opposite, 3.0cm long, 0.8 cm
wide above base, somewhat dilated distally with a
few lobes irregularly slightly elongate. Suprabasal
pinnae 2.7cm long, 0.8cm wide above truncate
and auricled base; apex abruptly obtuse; edges
lobed to a depth of 1mm; costules 2.5 mm apart;
veins to 4 pairs, 1: pairs anastomosing; lower
surface of rachis covered with pale hairs 0.2-
0.3 mm long, costae bearing pale spreading hairs
of mixed length to 0.6mm, sparse short hairs on
costules and veins and between veins; hairs on
upper surface of rachis 0.5 mm, on costae 0.2 mm,
very short suberect hairs present on surface be-
tween veins. Sori inframedial, not elongate; in-
dusia small, thin, dark red, with many stiff pale
hairs 0.2-0.3mm; sporangia with several short
setae; spores pale with translucent wing and
cross-wings.
Distr. Malesia: Philippines (Mindanao), only
known from type.
Notes. It is possible that larger sterile fronds
might be produced on other plants. The fertile
fronds differ from those of P. amphitrichum (in
which the upper surface is hairy in a similar way)
in having larger pinnae, sori not elongate and
setose sporangia. The smaller fertile fronds of the
type have shallowly crenate pinnae, but agree with
the largest in having irregular long lobes on the
apical lamina. Such irregular lobes are sometimes
a sign of hybridity, but there are abundant good
spores.
35. Pronephrium thysanoides HOLTTUM, sp. nov.
P. simillimo (C. Chr.) Holttum affinis, ab ea
differt: frondibus minoribus; pinnis non auri-
culatis; rachide costisque subtus glabris; soris
indusiatis, indusiis parvis, rigidis, setis brevibus
526
FLORA MALESIANA
(ser. Il, voleate
ciliatis. — Type: JERMY 14049, Sarawak, Gunong
Mulu, Ulu Sungei Air Jernih, north wall (BM).
Caudex short-creeping; stipe 15-20cm long,
glabrous, basal scales narrow, firm, c. 3mm long
with few hairs; fronds of type all fertile. Lamina
to 24cm long; pinnae 15 pairs, lower ones only
subopposite; apical lamina 4 cm long, deeply lobed
at base and grading to upper pinnae; basal pinnae
slightly narrowed towards their bases, more on
basiscopic than on acroscopic side, not auricled.
Largest pinnae 3.5 xX 1.1cm; base truncate; apex
abruptly acute, + upcurved; edges crenate to a
depth of less than 1 mm; costules 3 mm apart, at c.
60° to costa; veins 4 pairs, slender, | or 13 pairs
anastomosing, next vein passing to the short
sinus-membrane; lower surface of rachis and cos-
tae glabrous, of costules, veins and surface be-
tween veins bearing rather sparse very short hairs
which may be abraded from old fronds; upper
surface of rachis and costae bearing rather sparse
pale hairs 0.3mm long, rather sparse very short
hairs between veins. Sori medial, small; indusia
small but firm, with many short stiff marginal
hairs; sporangia with several short setae; spores
with many small wings.
Distr. Malesia: Borneo (Sarawak: Mt Mulu),
only known from the type.
Ecol. “In clay in crevice of open rock-face’’,
on limestone at 700 m.
36. Pronephrium hosei (BAK.) HOLTTUM, Blumea
20 (1972) 120.— Meniscium hosei BAK. J. Linn.
Soc. Bot. 22 (1886) 142.—Dryopteris hosei
(BAK)! Gy 1GHrs Inds) Fill (1905) 92712. COREL.
Philip. J. Sci. 10 (1915) Bot. 146. — Phegopteris
hosei (BAK.) v.A.v.R. Bull. Dép. Agr. Ind. Néerl.
21 (1908) 7; Handb. (1908) 510.— Cyclosorus
hosei (BAK.) COPEL. Gen. Fil. (1947) 142; Fern
Fl. Philip. (1960) 352. — Thelypteris hosei (BAK.)
K. IwatTs. Acta Phytotax. Geobot. 21 (1965)
170. — Type: G. F. HOSE 160, Sarawak (K).
Meniscium proliferum sensu. HOOK. 2 Cent.
Ferns (1861) t. 15, quoad plantam totam et fig. 2
tantum.
Caudex short-creeping; fronds dimorphous.
Sterile fronds. Stipe 5-15 cm long, minutely hairy
at base, scales c. 2X 1mm, caducous. Lamina
12-15 cm long, firm; apex to 7 cm long, narrowly
triangular, lobed near base only; pinnae 6-8 pairs;
lower pinnae not or little reduced, in most cases
narrowed slightly near base on basiscopic side,
edges sometimes crenate. Middle pinnae to Scm
long, 0.9cm wide above base; base truncate, al-
ways auricled on acroscopic side, often slightly
also on basiscopic; distal half of pinna evenly
attenuate to narrow rounded apex; edges entire or
+ crenate distally; costules 2mm apart; veins 4-5
on basiscopic side of costules, 3-4 on acroscopic,
to 7 pairs in basal auricle, 2 pairs anastomosing;
sinus-membrane not evident; lower surface of
tachis bearing curved hairs 0.5 mm long, hairs on
costae 0.1-0.2 mm, no other hairs except on mar-
gin, surface between veins very pustular; hairs of
upper surface of rachis to 1mm long, on costae
copious, 0.3mm long, some longer hairs present
near edge. Fertile fronds. Stipe to 25cm long;
lamina 10-12 cm long; apex 2.5 cm long; pinnae to
9 pairs, to 3.0cm long, 0.4 cm wide above auricled
base; pubescence as sterile but hairs on rachis
shorter; sori near costules, spreading somewhat
long veins; indusia persistent, bearing many hairs
0.1mm long; sporangia with 1-2 short setae;
spores pale, with longitudinal wing and cross-
wings. Chromosomes: n= 36 (T. G. WALKER).
Distr. Malesia: Borneo; Philippines (Min-
danao: Zamboanga Prov., SANTOS 4114).
Ecol. On rocky stream-banks in forest, at low
altitudes.
37. Pronephrium menisciicarpon (BL.) HOLTTUM,
Blumea 20 (1972) 111. — Aspidium menisciicarpon
BL. En. Pl. Jav. (1828) 142. — Dryopteris menisci-
icarpa (BL.) PosTH. Bull. Jard. Bot. Btzg III, 13
(1933) 93; BACKER & POSTH. Varenfl. Java (1939)
61, excl. syn. Dryopteris verruculosa v.A.v.R. —
Abacopteris menisciicarpa (BL.) HOLTTUM, Rev.
Fl. Malaya 2 (1955) 290, f. 168.— Thelypteris
menisciicarpa (BL.) K. IwATs. Acta Phytotax.
Geobot. 21 (1965) 171.— Type: BLUME, W. Java
(L, n. 908, 333-724).
Nephrodium latifolium PRESL, Epim. Bot.
(1851) 45, excl. CUMING 16; HOLTTUM, Novit.
Bot. Inst. Bot. Univ. Carol. Prag. 1968 (1969)
40. — Cyclosorus latifolius (PRESL) COPEL. Fern
Fl. Philip. (1960) 370.— Thelypteris latifolia
(PRESL) REED, Phytologia 17 (1968) 287. — Lec-
totype (HOLTTUM 1969): CUMING 298, Leyte
(PRC).
Abacopteris truncata FEE, Gen. Fil. (1852)
310. — Type: CUMING 298 (not seen).
Dryopteris holophylla C. CHR. Ind. Fil. (1905)
271, nom nov. for Polypodium holophyllum BAK.
J. Bot. 26 (1888) 325, non BAK. 1879. — Phegop-
teris holophylla (C. CHR.) v.A.v.R. Handb. (1908)
500. — Type: C. HOSE 242, Sarawak, Niah (K; E).
Dryopteris cordifolia v.A.v.R. Bull. Jard. Bot.
Btzg II, 11 (1913) 19, pl. 5; Handb. Suppl. (1917)
320. — Type: AMDJAH 322 p.p. N. Borneo (BO;
Kee):
Dryopteris mirabilis COPEL. Philip. J. Sci. 6
(1911) Bot. 137, pl. 19; v.A.v.R. Handb. Suppl.
(1917) 174; C. CHR. Gard. Bull. Str. Settl. 4 (1929)
391; 7 (1934) 249. — Type: BROOKS |6a, Sarawak,
Bidi (MICH; BM).
Dryopteris _korthalsii ROSENST. Meded.
Rijksherb. n. 31 (1917) 5.—Type: KORTHALS,
Sumatra (L; K).
Dryopteris verruculosa var. sumatrana v.A.v.R.
Bull. Jard. Bot. Btzg III, 2 (1920) 151.— Type:
Brooks 157/S, Sumatra, Lebong Tandai (BO;
BM).
Dryopteris urophylla var. peraspera v.A.v.R.
1981]
THELYPTERIDACEAE (Holttum)
“yal
Bull. Jard. Bot. Btzg III, 5 (1922) 303.— Type:
LAM 666, W. New Guinea (BO; L).
Nephrodium glandulosum sensu BEDD. Handb.
Suppl. (1892) 70, p.p.
Caudex short-creeping; fronds tufted, dimor-
phous. Sterile fronds. Stipe to 20cm_ long,
minutely hairy; basal scales thin, broad, soon
shrivelling. Lamina 30-40 cm long, consisting of
4-6(-8) pairs of pinnae and an apical lamina which
is longest on fronds with few pinnae; young plants
in Borneo (D. holophylla) have simple fronds to
25x8cm with cordate base. Pinnae to 13.5 x
4.5cm, usually widest at or above the middle;
base truncate with a slight acroscopic auricle
(especially on lower ones); apex abruptly short-
pointed; edges + crenate distally; costules to
4mm apart; veins to 10 pairs, anastomosing to
form a zig-zag excurrent vein; both surfaces of
rachis covered with coarse pale hairs, those on
costae much shorter, sparse on costules, surface
between veins strongly pustular. Fertile fronds.
Stipe to 45cm long; pinnae more widely spaced
than sterile ones, commonly 7 x 1.5 to 10 x 2.5 cm
(on some plants much smaller), shape as sterile
but edges almost entire; sori medial to suprame-
dial, slightly elongate; indusia with many short
hairs; sporangia bearing 0-2 setae; spores with
longitudinal wing and cross-wings.
Distr. Malesia: Sumatra, Malaya, Borneo, W.
Java; Philippines (Samar, Leyte); W. New Guinea,
Papua, New Britain.
Ecol. In forest near streams at low altitudes.
Notes. In Malaya (Pahang and Perak) plants
grow on earthen banks of streams; in such places
they are swept away by floods before they have
attained their full size, but they commonly
produce fertile fronds with pinnae to c. 3.0
0.6cm. Young plants with very large simple
fronds like the type of Dryopteris holophylla have
not been found in Malaya, but the largest adult
specimens from Malaya are not different from
those from Borneo. In the Philippines the largest
simple fronds seen are 18 x 6cm (sterile) and 13 x
4cm (fertile). The type of Dryopteris korthalsii
has up to 6 pairs of rather small fertile pinnae
(4x 1.3 cm); there are similar fronds from Borneo
with pinnae 10 x 2 cm.
Under the name Aspidium menisciicarpon BL.
METTENIUS described a specimen of a species of
Tectaria from the Philippines; his description was
copied by HOOKER who transferred the name to
Polypodium. BEDDOME’s description of Neph-
rodium glandulosum (Handb. Suppl. 70) was
based mainly on specimens cited which belong to
the present species, but in part on the true Aspi-
dium glandulosum BL. CHRISTENSEN (Ind. Fil.
1905) confused the present species with A. linea-
tum BL. and A. affine BL.
38. Pronephrium hewittii (COPEL.) HOLTTUM,
comb. nov. — Dryopteris hewittii COPEL. Philip.
J. Sci. 3 (1909) Bot. 344; v.A.v.R. Handb. Suppl.
(1917) 189. — Thelypteris hewittii (COPEL.) REED,
Phytologia 17 (1968) 282.—Type: Brooks &
HEWITT s.n. Feb. 1908, Sarawak, Bongo Range
(MICH).
Dryopteris compacta COPEL. Philip. J. Sci. 6
(1911) Bot. 137, pl. 18; v.A.v.R. Handb. Suppl.
(1917) 176.— Thelypteris compacta (COPEL.)
REED, Phytologia 17 (1968) 268.— Type:
BROOKS 4, Sarawak, Bongo Range (MICH;
BM). — Fig. 15e-i.
Caudex short-creeping; fronds subdimorphous.
Sterile fronds. Stipe 10-15cm long, glabrous;
basal scales thin, to 31mm. Lamina to 25cm
long; apex narrowly acuminate, to 5 x | cm, lobed
at base; pinnae to 20 pairs, close, spreading at
right angles to rachis; lower pinnae deflexed, nar-
rowed to base on both sides, with a small acros-
copic auricle. Middle pinnae 4-6cm long, 0.9-
1.1 cm wide above the truncate and strongly auri-
cled base; apex acuminate; edges crenate
throughout, distally to a depth of 1mm; costules
2.5-3 mm apart, at 45° to costa; veins to 3 pairs,
15-2 pairs anastomosing, sinus-membrane distinct
but very short; lower surface of rachis glabrous,
of costae bearing scattered hairs 0.5mm long, a
few shorter hairs on costules and veins, surface
between veins slightly pustular; upper surface of
rachis bearing thick pale hairs | mm long, costae
covered with hairs 0.2-0.3 mm long, a few longer
ones near ends of veins. Fertile frond. Stipe to
25 cm long; pinnae to 4cm long, 0.5—0.9 mm wide,
auricled as sterile but usually less deeply crenate;
sori near costules; indusia firm, dark, almost cir-
cular with hairs in the middle; sporangia not seti-
ferous.
Distr. Malesia: Borneo (Western Sarawak, 4
collections).
Ecol. “On small rock well above stream”
(MOLESWORTH ALLEN 3048, foot of Gunong
Santubong).
Notes. The type of D. hewittii is a small plant
with dimorphic fronds; that of D. compacta much
larger with fertile pinnae nearly as wide as sterile;
two later collections have narrower fertile pinnae.
In 1972 I united this species with P. merrillii but
the latter is quite distinct in venation and sori.
39. Pronephrium affine (BL.) PRESL, Epim. Bot.
(1851) 259; HoLTTUM, Blumea 20 (1972) 121, excl.
syn. praeter BL. et Gymnogramme macrotis
KUNZE. — Aspidium affine BL. En. PI. Jav. (1828)
148.— Type: BLUME, W. Java (L, n. 908, 333-
740).
Gymnogramme macrotis KUNZE, Bot. Zeit. 6
(1848) 114.—Dryopteris oxyotis ROSENST.
Meded. Rijksherb. n. 31 (1917) 5, nom. nov. (not D.
macrotis (HOOK.) O. KTZE).—Type: ZOL-
LINGER 324z, Java (L).
Aspidium lineatum sensu METT. Farngatt. IV
(1858) 110, quoad syn. A. affine BL. et G. macrotis
528
FLORA MALESIANA
(ser. II, vol. 1°
KUNZE tantum.— Nephrodium lineatum sensu
HooK. Spec. Fil. 4 (1862) 75, p.p.; RACIB. FI.
Btzg 1 (1898) 186.— Dryopteris lineata sensu C.
Cur. Ind. Fil. (1905) 63, 275, p.p.; sensu v.A.v.R.
Handb. (1908) 209; sensu BACKER & POSTH.
Varenfl. Java (1939) 60.
Caudex short-creeping or suberect; fronds
dimorphous. Sterile fronds. Stipe 15-20cm long,
glabrous except in groove, basal scales very thin,
to 4x2mm. Lamina to 25cm long, thin; apex
small and deeply lobed; pinnae to 12 pairs; basal
pinnae somewhat narrowed towards base both
sides with acroscopic auricle. Middle pinnae of
type 4x 1.7 cm (width above base), of a Sumatran
specimen 7X2.0cm; base truncate with acros-
copic auricle 2-4 mm long; apex of type abruptly
short-pointed, of Sumatran specimen short-acu-
minate; edges crenate throughout in smaller
plants, distally on larger ones; costules 3-3.5 mm
apart, at less than 60°; veins 5-6 pairs, 3-33 pairs
anastomosing, sinus-membrane very short; lower
surface of rachis bearing coarse hairs to 0.7 mm
long, hairs on costae 0.1mm, sometimes with a
few longer ones, similar hairs distally on costules
and veins, surface between veins pustular; hairs
on upper surface of rachis to 0.5mm long, very
short on costae and scattered distally on veins,
none between veins. Fertile fronds. Stipe to 35 cm
long; pinnae of type 3.0x0.7cm, of others to
4.0 x 0.9 cm, base strongly auricled (sometimes on
both sides), edges almost entire; veins to 4 pairs;
sori medial, slightly elongate; indusia thin, bearing
copious acicular hairs 0.1-0.2mm or sometimes
short capitate hairs; sporangia lacking setae;
spores pale with a rather broad continuous wing
and cross-wings.
Distr. Peninsular Thailand, in Malesia: Central
& S. Sumatra, West Java.
Ecol. By streams in forest, low altitudes.
Notes. The description by METTENIUS of A.
lineatum was evidently based on specimens of
several distinct species, doubtfully including the
type of A. affine; it was copied by HOOKER who
varied the list of synonyms, and the status of A.
affine as a synonym of A. lineatum was accepted
by CHRISTENSEN, who cannot have read
BLUME’s descriptions. But the descriptions of
RACIBORSKI, VAN ALDERWERELT and BACKER
& POSTHUMUS clearly apply to the present spe-
cies only.
BLUME’s type matches specimens from a low
altitude in Sumatra which, like BLUME’s, were
found on limestone rocks by streams. A specimen
from Pattani in Thailand is similar.
40. Pronephrium borneense (HOOK.) HOLTTUM,
comb. nov.— Polypodium borneense HOOK.
Spec. Fil. 5 (1863) 11. — Dryopteris labuanensis C.
Cure. Ind. Fil. (1905) 273, nom. nov. (not D. bor-
neensis (HOOK.) O. KTZE).— Phegopteris bor-
neensis (HOOK.) v.A.v.R. Handb. (1908) 500. —
Thelypteris borneensis (HOOK.) REED, Phytolo-
gia 17 (1968) 264. — Type: T. Loss, Labuan (K).
Caudex short-creeping; fronds not or little
dimorphous. Stipe to 12cm long, pale, minutely
hairy; basal scales thin, c. 2 1mm. Lamina thin,
to 18cm long; apex 6-9 cm long, triangular, lobed
near base which is 3-4 cm wide; pinnae 4—S pairs,
not opposite; basal pinnae narrowed towards base
on basiscopic side, slightly so to auricled acros-
copic base. Middle pinnae 3.5 x 1.5 to 5.0 x 2.2 cm;
base truncate with auricle 2-3 mm long; apex ab-
ruptly short-pointed; edges shallowly crenate
throughout; costules 2.5-3mm apart; veins 6
pairs, slender, at a broad angle to costules, almost
all anastomosing to form a zig-zag excurrent vein
(rarely with gaps in it), sinus-membrane not
evident; lower surface of rachis and costae spar-
sely short-hairy, rest glabrous and slightly pus-
tular; upper surface of rachis with copious hairs
0.2-0.7 mm long, costal hairs 0.3 mm, a few short
hairs also present near margin. Sori small, not
elongate, mostly medial; indusia very small
(sometimes lacking?), red when young, variable in
shape, with a few mostly marginal hairs 0.1 mm
long; sporangia with 4-5 short setae; spores with a
narrow wing and a few cross-wings.
Distr. Malesia: Borneo (Sarawak, Sabah).
Ecol. ‘Hill forest, on clay bank, 420m”
(JERMY 13072, G. Mulu); ‘on wet rocks, 300 m”
(BROOKS l6c, Bongo Range).
Notes. The type is a single fertile frond. Living
plants of JERMY 13072, cultivated at Kew,
resemble the type closely and have sterile and
fertile fronds of similar size; the above description
is based on the type and cultivated plants. Dried
specimens of JERMY 13078, also from G. Mulu,
are larger, with stipes 30cm long and dimorphous
fronds to 30 cm long with 6 pairs of pinnae; sterile
pinnae to 8.5 2.8cm (veins 7 pairs), fertile to
4.5 x 1.6cm.
41. Pronephrium exsculptum (BAK.) HOLTTUM,
Blumea 20 (1972) 117. — Acrostichum exsculptum
BAK. J. Bot. 26 (1888) 326.— Leptochilus
exsculptus (BAK.) C. CHR. Ind. Fil. (1905) 9, 385;
v.A.v.R. Handb. (1908) 740.— Dryopteris
exsculpta (BAK.) COPEL. Philip. J. Sci. 37 (1928)
410; C. CHR. Gard. Bull. Str. Settl. 7 (1934) 248,
p.p.— Thelypteris exsculpta (BAK.) K. IWATS.
Acta Phytotax. Geobot. 21 (1965) 170. — Type: C.
HOSE 244, Sarawak, Niah (K).
Meniscium stenophyllum BAK. J. Bot. 29 (1891)
108. — Phegopteris stenophylla (BAK.) v.A.v.R.
Handb. (1908) 510.— Thelypteris stenophylla
(BAK.) REED, Phytologia 17 (1968) 315. — Type:
G. F. HOSE 20, Sarawak (K).
Caudex short-creeping; fronds dimorphous.
Sterile fronds: Stipe 10-17cm long, glabrous
except distally; basal scales to 51mm, dark,
firm. Lamina 25cm long, very firm; apex small,
triangular, deeply lobed; pinnae 20-25 pairs; basal
1981]
THELYPTERIDACEAE (Holttum)
529
pinnae slightly narrowed both sides towards auri-
cled base. Middle pinnae 3.5 X 1.0cm; base trun-
cate, slightly auricled; apex short-acuminate (pin-
nae of type of M. stenophyllum to 2.0 x 0.8 cm,
apex shortly obtuse); edges crenate to a depth of
0.5mm; costules 2—2.5mm apart; veins 3 pairs,
thick, pale and prominent on lower surface, 2
pairs anastomosing, sinus-membrane not evident;
lower surface of rachis bearing thick curved hairs
less than 0.5mm long, a few minute hairs on
costae, rest of surface glabrous, not or hardly
pustular; upper surface of rachis as lower, sparse
short hairs on costa. Fertile fronds: Stipe 25 cm or
more long; pinnae to 2.0cm long (type of M.
stenophyllum 1.0 cm), 0.3—-0.4 cm wide above base
which is strongly auricled (sometimes both sides);
veins 2 pairs; sori near costules, at maturity filling
lower surface; indusia small with a few very short
marginal hairs; sporangia not setiferous; spores
not seen.
Distr. Malesia: Borneo (Sarawak).
Ecol. Probably on limestone.
Note. CHRISTENSEN (l.c. 1934) referred here
specimens with thinner pustular pinnae and seti-
ferous sporangia; these are now placed in P.
rhombeum and P. peltatum.
42. Pronephrium merrillii (CHRIST) HOLTTUM,
Blumea 20 (1972) 117.—Dryopteris merrillii
CHRIST, Philip. J. Sci. 2 (1907) Bot. 201. — Phe-
gopteris merrillii (CHRIST) v.A.v.R. Handb. Suppl.
(1917) 505.—Cyclosorus merrillii (CHRIST)
CopEL. Gen. Fil. (1947) 143; Fern Fl. Philip.
(1960) 352.— Thelypteris merrillii (CHRIST)
REED, Phytologia 17 (1968) 292.— Type: Fox-
WORTHY 742, Palawan (P).
Differs from 38. P. hewittii as follows: fertile
pinnae to 2.7 x 0.4 cm, entire or nearly so; indusia
very small, asymmetric, with a few very short
marginal hairs.
Distr. Malesia: Philippines (Palawan, 2 collec-
tions; second is ELMER 13031).
43. Pronephrium rhombeum (CHRIST) HOLTTUM,
Blumea 20 (1972) 120.— Dryopteris diversiloba
var. acrostichoides subvar. rhombea CHRIST,
Philip. J. Sci. 2 (1907) Bot. 200.— Dryopteris
acrostichoides var. rhombea (CHRIST) v.A.v.R.
Handb. Suppl. (1917) Corr. 49.— Dryopteris
rhombea (CHRIST) COPEL. Philip. J. Sci. 56
(1935) 102, pl. 6 quoad icon. photogr. tantum. —
Cyclosorus rhombeus (CHRIST) CHING, Bull.
Fan Mem. Inst. Biol. Bot. 8 (1938) 192; COPEL.
Fern Fl. Philip. (1960) 357, p.p.— Thelypteris
rhombea (CHRIST) REED, Phytologia 17 (1968)
309. — Lectotype (COPELAND 1935): CUMING
149, Luzon (orig. PNH destroyed; isotype K).
Dryopteris diversiloba var. acrostichoides sub-
var. lanceola CHRIST, lI.c.— Dryopteris acrosti-
choides var. lanceola (CHRIST) v.A.v.R., l.c.—
Dryopteris lanceola (CHRIST) COPEL. Philip. J.
Sci. 56 (1935) 102, pl. 7. — Cyclosorus lanceolus
(CHRIST) COPEL. Gen. Fil. (1947) 142; Fern FI.
Philip. (1960) 353.— Thelypteris —_lanceola
(CHRIST) REED, Phytologia 17 (1968) 286.—
Lectotype (COPELAND 1935): COPELAND 250,
Luzon, Mt Mariveles (orig. PNH destroyed; iso-
types K, MICH, US). — Fig. 15j-1.
Caudex short-creeping; fronds dimorphous.
Sterile fronds. Stipe 5-8cm long with pale hairs
1 mm long throughout. Lamina to 15 cm long, thin
but firm; apex narrowly triangular, longer than
pinnae; pinnae 8(-10) pairs; basal pinnae not or
little narrowed towards base. Middle pinnae 2.0-
3.5 cm long, 0.7—1.0 cm wide above auricled base;
apex abruptly acute or obtuse, sometimes a little
dilated; edges crenate, more deeply towards apex;
costules 2mm apart, at little more than 45° to
costa; veins mostly 3 pairs, concolorous and
slightly prominent, 2 pairs anastomosing, sinus-
membrane not evident; lower surface of rachis
bearing pale hairs 0.7 mm long, a few similar hairs
with shorter ones present on costae, sparse short
hairs on costules and veins, surface between veins
pustular; upper surface of rachis hairy as lower,
hairs on costae 0.2-0.3 mm long, a few short hairs
near margin. Fertile fronds. Stipe to 25cm long;
lamina 10-20cm long, in the latter case pinnae
more widely spaced; pinnae 2.0-3.0 cm long, 0.4—
0.6cm wide, base auricled, edges almost entire;
sori near costules, not elongate; indusia small,
red, with marginal hairs 0.2-0.3 mm long; sporan-
gia mostly with 2-3 short setae (in some cases
none); spores with translucent wing and cross-
wings.
Distr. Malesia: Philippines (Luzon to Min-
danao), N. Borneo, Celebes.
Notes. The Kew isotypes of the lectotypes
cited do not differ in the ways indicated in
COPELAND’s keys and descriptions of 1960, and
among many specimens seen I cannot see two
distinct forms. The Kew specimen of COPELAND
250 has pinnae more dilated distally than that of
CUMING 149, though COPELAND indicates the
contrary. The pinnae with irregular large distal
lobes figured (as line drawings) with the pho-
tograph of the type of D. rhombea by COPELAND
(1935) are representative of a group of forms,
almost always with 4-5 pairs of pinnae, which
appear to be hybrids and are here treated as
P. x xiphioides.
44. Pronephrium peltatum (v.A.v.R.) HOLTTUM,
comb. nov.— Dryopteris peltata v.A.v.R. Bull.
Jard. Bot. Btzg II, 16 (1914) 12; Handb. Suppl.
(1917) 190. — Thelypteris peltata (v.A.v.R.) REED,
Phytologia 17 (1968) 303. — Type: MATTHEW 632,
Sumatra, Gunong Tandikat (BO).
Dryopteris zippelii ROSENST. Meded.
Rijksherb. n. 31 (1917) 6.—Type: ZIPPEL, Java
(L)
Dryopteris lineata var. subacrostichoides
530
v.A.v.R. Bull. Jard. Bot. Btzg II, 28 (1918) 24. —
Type: CRAMER 37, S. Sumatra, Kota Agoeng
(BO).
BEY TO THE VARTEGIES
1. Indusia glabrous; sporangia not setiferous.
2. Sterile pinnae thin, margin deeply serrate dis-
tally nee Bess . a. var. peltatum
. Sterile pinnae rigid when dry, entire
b. var. tenompokense
1. Indusia with a few short hairs; sporangia seti-
ferous or not.
3. Sporangia not setiferous
3. Sporangia setiferous.
4. Sporangia with several setae; basal pinnae
not reduced d. var. persetiferum
4. Sporangia with 0-2 setae; reduced basal
pinnae present on most specimens
e. var. aberrans
in)
c. var. peninsulare
a. var. peltatum
Stipe of sterile frond 14cm long, of fertile
20cm. Lamina to 21cm long; pinnae c. 10 pairs.
Sterile pinnae to 6cm long, base rounded on
basiscopic side and auricled on acroscopic, 1.5 cm
wide above base, apex short-acuminate, edges
crenate throughout or in distal part only; costules
2.5mm apart; veins 3-4 pairs, almost all anas-
tomosing; lower surface glabrous, pustular when
dry; upper surface hairy on costae only. Fertile
pinnae to 4X 1 cm; sori medial; indusia firm, dark,
lacking acicular hairs; sporangia not setiferous.
Distr. Malesia: Central & S. Sumatra, W.
Java, at 900-1500 m.
Note. Specimens from Tjibodas in W. Java
have sterile pinnae to 8x 1.6cm. One specimen
from Sumatra has lower sterile pinnae auricled on
both sides at the base.
b. var. tenompokense (C. CHR.) HOLTTUM, Stat.
nov. — Dryopteris tenompokensis C. CHR. Gard.
Bull. Str. Settl. 7 (1934) 248. — Type: HoLTTUM
25388, Sabah, Mt Kinabalu, Tenompok (BM; K,
SING).
Differs from var. peltatum as follows: texture
of lamina firm; edges of pinnae almost entire;
FLORA MALESIANA
[ser. II, vol. 1°
lower surface of costae glabrous or sparsely
minutely hairy.
Distr. Malesia: Sabah (Mt Kinabalu).
Ecol. In forest, on rocks by streams or near
streams, at 1500 m.
c. var. peninsulare HOLTTUM, var. nov.—
Abacopteris lineata sensu HOLTTUM, Rev. FI.
Mal. 2 (1955) 292, excl. syn., f. 169. — Type: A. G.
PIGGOTT 1668, Malaya, Negri Sembilan, G.
Telapak Burok, 900 m (K).
A var. tenompokense differt indusiis semper
breviter pilosis.
Distr. Malesia: Malaya, on Main Range and
Taiping Hills.
Ecol. In forest, not (or not always) by streams,
900-1500 m.
Note. A plant of this variety from Taiping
Hills, cultivated at Kew, was found by MANTON
to be tetraploid (HOLTTUM I.c. p. 625).
d. var. persetiferum HOLTTUM, var. nov. —
Dryopteris exsculpta sensu C. CHR. Gard. Bull.
Str. Settl. 7 (1934) 248, p.p.— Type: HOLTTUM
s.n. 1931, Mt Kinabalu, crest of Penibukan Ridge,
1200 m (SING; BM).
A var. tenompokense_ differt pinnis
crenatis, sporangiis setis pluribus praeditis.
Distr. Malesia: Borneo. Besides type, known
from G. Mulu, Sarawak, in stream-bed at 1760 m.
leviter
e. var. aberrans HOLTTUM, var. nov.
A var. tenompokense differt: frondibus pinnis
inferioribus redactis usque 12-jugatis, aequaliter
bilobatis (lobis usque 1.2cm longis) praeditis;
pinnis normalibus sterilibus usque 122.5 cm,
distaliter crenatis, fertilibus usque 7X 1.7cm;
sporangiis 0-2 setis praeditis. —Type: CLEMENS
33702 & 32020’, Sabah. Mt Kinabalu, Colombon
Basin, 1400-1600 m (K; G, MICH).
Distr. Malesia: Sabah.
Note. These specimens have the frond-form of
Sphaerostephanos, but the reduced basal pinnae
are not like those of that genus; apart from size,
the normal pinnae agree with the present species.
Spores of a specimen at MICH are normal.
2. Subgenus Menisciopsis HOLTTUM, subg. nov.
Pinnae subintegrae, plurivenosae, venis plerisque more generis Meniscii
Schreb. anastomosantibus; sori
exindusiati,
saepe elongati, interdum
coalescentes; sporangia vel nudis vel pilis rectis vel hamatis praedita.
Type species: Pronephrium lakhimpurense (RosENST.) HOLTTUM, Blumea 20
(1972) 110.
Taxon. The type species, P. lakhimpurense, is distributed from N.E. India to southern China and
southwards to northern Thailand and Vietnam. It has large entire pinnae with many pairs of anastomos-
ing veins, the excurrent veinlets all free; the sori of large plants spread all along the veins, the sporangia
1981] THELYPTERIDACEAE (Holttum) 531
lacking setae. In these characters it is very close to the species of Meniscium of tropical America, and
differs from them most conspicuously in the red colour of dried fronds. Another species of the same
distribution in Asia, P. penangianum (HOOK.) HOLTTUM, has narrower pinnae which are strongly
crenate-serrate and distal excurrent veinlets not free. In Malesia, P. rubrinerve, P. rubidum and P.
scopulorum are clearly related to the two species of mainland Asia, but their sori are not more than
slightly elongate; only in P. rubidum are excurrent veinlets all free.
Agreeing with the species mentioned above in the reddish colour of dried fronds and also in elongate
sori are species of another Malesian group which are distinguished by having hooked hairs on
frond-axes and usually also on sporangia; formerly these species were all included in Meniscium.
Several of them have buds at the bases of pinnae. They are here included in section Grypothrix.
There remain three species which appear to be somewhat intermediate between the two groups above
mentioned. Of these, P. acanthocarpum has buds at the bases of its pinnae as in sect. Grypothrix but
sori not elongate and straight setae on its sporangia. The other two, P. nitidum and P. repandum, have
similar sori and sporangia but no buds on the frond; they are very similar to each other in general aspect
but differ in sporangia and spores. They are possibly a connecting link between subgenera Pronephrium
and Menisciopsis. They are here placed in section Menisciopsis.
The earlier-described species of this group were much confused by HOOKER, and it was not until the
work of CHING in 1938 that their taxonomy and nomenclature were clarified. CHING however did not
notice the hooked hairs which are the distinctive feature of sect. Grypothrix.
3. Section Menisciopsis
KEY TO THE SPECIES
1. Body of sporangium lacking setae.
2. Pinnae 4-6 cm wide: edges distinctly serrate throughout
2. Pinnae 2.5—3.5 cm wide; edges entire or nearly so.
3. Excurrent veinlets mostly free; sori close to costules
= Excurrent veinlets mostly not free; sori not close to costules
. Body of sporangium bearing setae.
m Pinnae one pair, a bud at the base of each
4. Pinnae several pairs on adult plant; no buds present.
5. Caudex long-creeping: veins very oblique; several acicular hairs on stalk of sporangium
49. P. nitidum
5. Caudex short-creeping; veins at a broad angle to costules; a hair of 2-3 cells, not acicular on stalk
of sporangium 50. P. repandum
. 45. P. rubrinerve
.46. P. rubidum
.47. P. scopulorum
48. P. acanthocarpum
45. Pronephrium rubrinerve (METT.) HOLTTUM, _ pinna-like; basal pinnae distinctly stalked, nar-
Blumea 20 (1972) 110; Allertonia 1 (1977) 213;
BROWNLIE, Pterid. Fiji (1977) 258. — Phegopteris
rubrinervis METT. in Kuhn, Linnaea 36 (1869) 116;
KUHN, Verh. Zool. Bot. Ges. Wien 19 (1869) 576;
v.A.v.R. Handb. Suppl. (1917) 316. — Goniopteris
rubrinervis (METT.) CARR. in Seem. FI. Vit.
(1873) 366.— Polypodium rubrinerve (METT.)
BAK. Syn. Fil. ed. 2 (1874) 315.— Cyclosorus
rubrinervis (METT.) COPEL. Gen. Fil. (1947) 143;
J. Arn. Arb. 30 (1949) 438.— Thelypteris
rubrinervis (METT.) K. IWATS. Mem. Coll. Sci.
Univ. Kyoto B, 31 (1965) 195.— Lectotype
(HOLTTUM 1972): TURNER, New Ireland (B).
Caudex massive, short-creeping to suberect;
fronds closely tufted; scales small, thin, broad-
ovate, glabrous or nearly so. Stipe to 145 cm long,
glossy, glabrous apart from a few very short aci-
cular hairs near base (often lost), drying light
reddish; lamina to 170cm long, thin but firm,
drying brown-olivaceous with red costae and in
some cases costules; pinnae to 24 pairs, upper
ones on large fronds gradually decrescent; apex
rowly cuneate at base, the lamina on basiscopic
side ending 1.5-2.0 cm from rachis. Largest pinnae
to 436cm; base broadly cuneate: apical half
gradually narrowed to short-acuminate tip: edges
crenate to a depth of 1mm, cartilaginous margin
thick, pale to reddish; costules 4-5 mm apart, at
60° to costa, falcate distally; veins 15-20 pairs, at
45° to costules, almost straight, fine and slightly
prominent on lower surface, almost all anas-
tomosing, excurrent veinlets, except basal ones,
rarely free: lower surface of rachis and pinnae
quite glabrous (young fronds may have thin red-
dish scales on rachis and costae), surface between
veins slightly pustular; upper surface of both
rachis and costae glabrous. Sori medial or
supramedial, basal ones sometimes meeting and
slightly elongate; no indusia; sporangia lacking
setae, a hyaline hair of 2 cells on the sporangium-
stalk; spores red-brown, with a rather broad
translucent wing and cross-wings.
Distr. Polynesia (Fiji), Melanesia (New Heb-
rides) and Malesia: New Guinea (New Ireland).
532
Ecol. On river banks in forest at low altitude
(to 900 m in Fiji).
Notes. In 1977 I reported the caudex of this
species to attain a height of 30cm, on the evi-
dence of a herbarium label; but BRAITHWAITE
reports it to be prostrate, and this is also true of a
plant he sent from the New Hebrides, cultivated
at Kew.
46. Pronephrium rubidum (HOOK.) HOLTTUM,
Blumea 20 (1972) 111.— Polypodium rubidum
HooK. Spec. Fil. 5 (1863) 12, excl. DE VRIESE
41.— Dryopteris rubida (HOOK.) C. CHR. Ind.
Fil. (1905) 289; CHRIST, Philip. J. Sci. 2 (1907)
Bot. 207, excl. WHITFORD 272. — Phegopteris
rubida (HOOK.) v.A.v.R. Handb. (1908) 502. —
Cyclosorus rubidus (HOOK.) COPEL. Gen. Fil.
(1947) 143; Fern Fl. Philip. (1960) 375. — Thelyp-
teris rubida (HOOK.) K. IwWATS. Mem. Coll. Univ.
Kyoto B, 31 (1965) 195.— Type: CUMING 415,
Euzoni(K; E; FIEW,G, LE; P).:
Caudex short, suberect. Stipe dark reddish,
glabrous except for minute hairs near base, to
100cm long; basal scales not seen, reported by
CHRIST to be 2cm long. Lamina to 120cm long;
apex pinna-like; pinnae to 20 pairs, subopposite,
drying dark reddish; basal pair not or little
reduced, with stalks 1-2 mm long. Largest pinnae
of type 21 x 2.7cm, largest seen 28 x 3.8 cm; base
cuneate (angle c. 45° each side); apex caudate-
acuminate (cauda 2.5-3.0cm long); edges irre-
gularly slightly sinuous to slightly crenate, car-
tilaginous margin rather broad and red; costules
3-5 mm apart, at 60°; veins 9-13 pairs, at 45° to
costules, slightly curved, dark and not prominent
on lower surface, excurrent veinlets all free; lower
surface of rachis and pinnae quite glabrous; upper
surface of rachis with a few short hairs in the
groove, hairs on costae sparse, slender. Sori very
close to costules except basal basiscopic ones on
large pinnae, sporangia few; no indusia; no setae
on body of sporangia; hair on sporangium-stalk
3-celled, distal cell not much enlarged.
Distr. Malesia: Philippines (Luzon; Palawan,
4 collections).
Ecol. “Streamside, in open pine
1700 m” (PRICE 2831; no other record).
Notes. The largest specimen is FOxWORTHY
684 from Palawan (MICH). HOOKER described
var. B, but he cited no specimen and I have not
found one so marked. DE VRIESE 41, wrongly
cited by HOOKER, is P. ramosii, as also WHIT-
FORD 272 wrongly cited by CHRIST.
forest,
47. Pronephrium scopulorum HOLTTUM, sp. nov.
A P. rubido differt: pinnis omnibus basi anguste
inaequaliter cuneatis, pinnis basalibus non stipi-
tatis; venulis excurrentibus plerisque non liberis;
soris costulas non _ contingentibus. — Type:
BUWALDA 5168, Aru Is., P. Kabroor, on lime-
stone (K; BO).
FLORA MALESIANA
[ser. II, vol. 1°
Caudex not seen. Stipe probably to 50 cm long,
dark reddish, glabrous; basal scales not seen.
Lamina c. 50 cm long; apex pinna-like; pinnae to 8
pairs, firm, drying dull reddish, subopposite; bases
of all pinnae narrowly unequally cuneate, lowest
pinnae with narrowest base. Middle pinnae 18 x
2.8-3.5 cm, widest 1/3 from base; acroscopic base
narrowed to attachment to rachis, lamina on
basiscopic side ending 3-4 mm from rachis; apex
narrowly caudate; edges entire or slightly sinuous
distally; costules 3-4mm apart at 60° to costa;
veins to 10 pairs, at c. 45° to costules, almost
straight, almost all anastomosing to form zig-zag
excurrent veins between costules, basal basiscopic
vein arising from costa; both surfaces quite
glabrous (upper surface of rachis and costae
probably have some hairs when pinnae are
young). Sori near costules but not touching them,
basal ones sometimes supramedial; no indusia;
sporangia lacking setae.
Distr. Malesia: S.E. Moluccas (Aru Is.: P.
Kabroor) and West New Guinea (Waigeu L:
CHEESMAN 1240-1243, BM).
Ecol. On steep limestone which carries little
other vegetation.
48. Pronephrium acanthocarpum (COPEL.)
HOLTTUM, Blumea 20 (1972) 107.— Dryopteris
acanthocarpa COPEL. Philip. J. Sci. 6 (1911) Bot.
136, pl. 17. — Phegopteris acanthocarpa (COPEL.)
v.A.v.R. Handb. Suppl. (1917) 315. — Thelypteris
acanthocarpa (COPEL.) REED, Phytologia 17
(1968) 257.—Type: BRooKs 54, Sarawak, Mt
Penrissen (MICH; BM).
Caudex short-creeping; bases of stipes less than
lcm apart. Stipe to 50cm long, dark at base with
small dark scales and sparse minute hairs, rest
glabrous, dull reddish-brown. Lamina consisting
of one pair of opposite pinnae with a bud at the
base of each and an apical section larger than
pinnae, the whole dull reddish when dry. Apical
lamina to 23 x 10 cm, widest near broadly rounded
and symmetric base, apex short-acuminate, edges
irregularly slightly sinuous; main veins 6mm
apart, at wide angle to midrib, falcate distally;
veins more than 20 pairs, not prominent, at a wide
angle to main veins, almost all anastomosing with
occasional irregularities, excurrent veinlets free;
lower surface of midrib copiously minutely hairy
(less than 0.1mm), similar hairs sparse on main
veins, surface between veins not pustular; sori
medial or inframedial, those on basal veins some-
times supramedial, in many cases slightly elon-
gate; sporangia bearing several short straight
setae; spores not seen. Pinnae to 17 X7cm; base
broadly cuneate and very asymmetric, the basis-
copic side 4.5 cm wide at 1/3 length of pinna, acro-
scopic side 3cm; veins to 18 pairs on basiscopic
side; pubescence and sori as terminal lamina.
Distr. Malesia: Borneo (Western Sarawak,
two localities; the second is Mt Merinjak).
1981]
THELYPTERIDACEAE (Holttum)
533
Ecol. On rocks near streams at 900 m.
49. Pronephrium nitidum HOLTTUM, Blumea 20
(1972) 109.— Dryopteris urophylla (METT.) C.
Cur. var. nitida HOLTTUM, Gard. Bull. Str. Settl.
7 (1934) 249, 251 (fig.).— Thelypteris urophylla
var. nitida (HOLTTUM) K. IWATs. Acta Phytotax.
Geobot. 22 (1966) 94.— Type: HoLTTUM SFN
25592, Sabah, Mt Kinabalu, Menetendok (SING;
BO, K).
Cyclosorus urophyllus sensu COPEL. Fern FI.
Philip. (1960) 373, excl. loc. Malacca. — Fig. 16g—h.
Caudex long-creeping, 5-8mm diameter. Stipe
commonly 70-100 cm long; base dark with dark
rigid scales c. 5X 1mm bearing many short stiff
hairs, above base brownish to slightly reddish,
glabrous. Lamina 50cm long, firm; apex pinna-
like, often with almost symmetrical base; pinnae
6-7 pairs, subopposite, lowest pinnae usually with
a Stalk to 4mm long, narrowly and asymmetrically
cuneate at base, upper pinnae sessile with broader
and more symmetric bases (very variable). Middle
pinnae commonly 20 x 4-5 cm, narrowly elliptic, +
abruptly narrowed to cuneate base and to caudate
apex (cauda commonly 2 cm long); edges crenate-
serrate with a thick pale cartilaginous margin;
costules 3.5-4 mm apart, at a broad angle, falcate
distally; veins 18-20 pairs, slender and slightly
prominent both sides, very oblique and slightly
S-curved, excurrent veinlets almost all free; lower
surface of rachis and costae covered with pale
thin hairs 0.2-0.3 mm long (antrorse but not ap-
pressed on costae), similar and sometimes also
longer hairs on costules, surface between veins
glabrous or with some very short erect hairs, at
most slightly pustular; upper surface of rachis and
costae glabrous or with sparse short hairs, surface
between veins smooth and glossy. Sori supra-
medial, those on connivent veins sometimes +
confluent; receptacle bearing many short acicular
hairs with the sporangia; sporangia short-stalked
with several acicular hairs on stalks, also 8-12
short acicular hairs distally on body; spores
closely and minutely papillose.
Distr. Malesia: Banka, Borneo, N. & Central
Celebes, Philippines (Sulu Archipelago, Min-
danao).
Ecol. At 0-1200 m, usually in secondary forest,
often on edges of forest or rather open places on
stream-banks.
Note. COPELAND (1960) reports from
Palawan, but I have seen no specimen thence.
Both sporangia and spores are markedly different
from those of P. repandum.
50. Pronephrium repandum (FEE) HOLTTUM,
Blumea 20 (1972) 109.—Goniopteris repanda
FEE, Gen. Fil. (1852) 251.—Type: GAvuDI-
CHAUD, Penang (FI-W, isotype).
Polypodium cuspidatum ROxB. Calc. J. Nat.
Hist. 4 (1844) 491 (not Pronephrium cuspidatum
(BL.) HOLTTUM).— Type:
Penang (BR; K).
Goniopteris dalhousiana FEE, 8e Mém. (1857)
92.— Type: Lady DALHOUSIE, “Indes Orien-
tales” (Herb. Graham, not seen; probable isotype,
from Penang, K).
Phegopteris urophylla MetTT. Farngatt. IV
(1858) 26, excl. var. aspera (PRESL).— Poly-
podium urophyllum (METT.) HOOK. Spec. Fil. 5
(1863) 9, excl. syn. et var.— Nephrodium urophyl-
lum (METT.) KEYSERL. Pol. et Cyath. Herb.
Bung. (1873) 49; BEpp. Handb. (1883) 274, p.p. —
Dryopteris urophylla (METT.) C. CHR. Ind. Fil.
(1905) 299, p.p.; v.A.v.R. Handb. (1908) 216, p.p.;
BACKER & PosTH. Varenfl. Java (1939) 64,
p.p.— Abacopteris urophylla (METT.) CHING,
Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 251;
HOLT TUM, Rev. Fl. Malaya 2 (1955) 296, f.
172. — Cyclosorus urophyllus (METT.) TARD.
Notul. Syst. 7 (1938) 77; TARD. & C. CHR. FI.
Gén. I.-C. 7, pt. 2 (1941) 391, p.p. — Thelypteris
urophylla (METT.) K. IWATS. S.E. Asian Studies
3 (1965) 81; Acta Phytotax. Geobot. 22 (1966)
94.— Type: WALLICH 299, Penang (B?; isotype
K).
Polypodium pinwillii BAK. Ann. Bot. 5 (1891)
460.— Nephrodium uropyllum_ var. _ pinwillii
BEDD. Handb. Suppl. (1892) 72.—Type: PIN-
WILL, Malacca (K). — Fig. In, 16i-j.
Caudex short-creeping, fronds tufted at its
apex. Stipe to 80cm long, slightly flushed with
red, densely short-hairy with narrow very hairy
scales at base, glabrescent above base. Lamina to
60cm long, firm, drying green but rachis, costae
and costules + flushed with red; apex pinna-like
with + asymmetric cuneate base; pinnae 3-6 pairs;
lowest pinnae usually slightly smaller than next,
always more narrowed at base than the rest,
which are very variable in width in relation to
length. Middle pinnae commonly c. 22 *4.0cm,
largest 30x 7cm, widest at, or more commonly
above, the middle, gradually narrowed to a rather
abruptly cuneate base, apex abruptly caudate-
acuminate, cauda 3-4cm long; edges + deeply
crenate-serrate with very narrow _ pale
cartilaginous margin; costules 4-5 mm apart, at
45-60°, slightly curved; veins 15-18 pairs, slender,
slightly prominent both sides, slightly curved;
at 45° to costule, excurrent veinlets often but not
always free (if free, extending almost to the
junction of the next pair of connivent veins); lower
surface of rachis and costae + densely covered with
erect pale hairs 0.2-0.3 mm long, similar hairs more
sparse on costules, surface between veins finely
pustular, bearing a variable number of short erect
hairs some of which may be hooked; upper surface
of rachis and costae hairy as lower surface, rest
glabrous, surface between veins minutely pustular.
Sori medial or supramedial, not elongate, exin-
dusiate; sporangia bearing 10 or more short setae
distally, stalk longer than in P. nitidum witha hair of
WALLICH- 299,
534 FLORA MALESIANA [ser. II, vol. 1°
one cell which is in some cases almost spherical; Moulmein southwards to Malesia: Malaya,
spores with a continuous wing and cross- Sumatra, Riouw, Lingga & Anambas Is., and West
wings. Java.
Distr. South Thailand & Vietnam, from Ecol. In lowland forest, not in exposed places.
4. Section Grypothrix
HOLTTUM, Blumea 19 (1971) 36.
Type species: Pronephrium cuspidatum (BL.) HOLTTUM.
Taxon. See discussion under subg. Menisciopsis. The earlier-known species of this section were
placed in the genus Meniscium SCHREB. by SWARTZ, BLUME, PRESL, HOOKER, BEDDOME and
RACIBORSKI, and in Phegopteris sect. Meniscium by METTENIUS and VAN ALDERWERELT. CHRIS-
TENSEN (Monogr. Dryopteris 1, 1913, 248) was the first author to regard them as belonging to a group
distinct from the type of Meniscium. CHING (1938) clarified the taxonomy and nomenclature of the
species of Mainland Asia but did not note the hooked hairs by which these species are distinguished
from those here placed in sect. Menisciopsis.
P. triphyllum is distributed throughout the range of the genus Pronephrium; the others are more local
in distribution. There are plants which appear to be hybrids of which one parent is P. triphyllum. In
Malesia P. x parishii is so designated; there are others, more irregular in frond-form, in Sri Lanka and
the Ryukyu Islands. FEE described a species Abacopteris elegans (Gen. Fil. 1852, 310), based on a
specimen from Cochinchina which I have not seen; he stated that it had hooked hairs on sporangia,
indusiate sori, and excurrent veinlets not free, a combination of characters which I have never seen, and
I doubt the accuracy of his observations.
KEY LO THESSRECIES
uuCauder long- creeping with well-spaced fronds.
. Pinnae one pair, opposite, always asymmetric : wh
Di Pinnae 2-6 pairs, distal ones small and adnate, rest with variable asymmetry
Caudex short-creeping, fronds close together.
a Pinnae all narrowly cuneate at base, fertile ones ee lcm wide; hooked hairs lacking on
lamina and sporangia 3 . . . . 53. P. salicifolium
3. Pinnae not all narrowly cuneate at base, fertile ones always much 1 more than | cm wide; hooked
hairs present on some parts of lamina and usually on sporangia.
4. Pinnae 1-3 pairs; no buds at their bases.
. Pinnae always opposite; fertile pinnae with undulate margins; hooked hairs present on basal
scales and on sporangia 3 54. P. rubicundum
5. Pinnae not always opposite; fertile pinnae sharply serrate- crenate distally; no hooked hairs on
scales nor on sporangia : 55. P. sulawesiense
4. Pinnae of well-grown plants more than 3 pairs; a 1 bud always present at base of uppermost pinna.
6. Bud confined to base of uppermost pinna; pinnae widest in basal half; hydathodes on upper
surface conspicuous : : . 56. P. cuspidatum
6. Buds present at bases of several pinnae: pinnae “usually widest above middle; hydathodes not
conspicuous. . Sat ®.. ene tatty ee ee . 2) J See ramos
7. Hooked hairs present on sporangia 57a. P. ramosii var. ramosii
7. Hooked hairs lacking on sporangia . 57b. P. ramosii var. minahassae
. 51. P. triphyllum
52. P. Xx parishii
51. Pronephrium triphyllum (Sw.) HoLtTTum,
Blumea 20 (1972) 122; Allertonia 1 (1977) 214;
BROWNLIE, Pterid. Fiji (1977) 257. — Meniscium
triphyllum Sw. in Schrad. J. Bot. 1800, 2 (1801)
triphylla (Sw.) CHING, Bull. Fan Mem. Inst. Biol.
Bot. 8 (1938) 241; Ho_truM, Rev. FI. Malaya 2
(1955) 287, f. 166.— Cyclosorus triphyllus (Sw.)
TARD. Notul. Syst. 7 (1938) 78; TARD. & C. CHR.
16; HooK. & GREV. Ic. Fil. (1820) t. 120; BEDD.
Ferns S. India (1863) t. 56; Handb. (1883) 397:
Racis. Fl. Btzg 1 (1898) 66.— Phegopteris
triphylla (Sw.) Metr. Fil. Lechl. (1859) 21;
v.A.v.R. Handb. (1908) 509. — Dryopteris triphylla
(Sw.) C. CHr. Ind. Fil. (1905) 298; BACKER &
PosTH. Varenfl. Java (1939) 59.— Abacopteris
Fl. Gén. I.-C. 7, pt. 2 (1941) 386; COPEL. Fern FI.
Philip. (1960) 371. — Thelypteris triphylla (Sw.) K.
IwaATs. Mem. Coll. Sci. Univ. Kyoto B, 31 (1965)
190. — Type: China (S-PA).
Meniscium cumingii FEE, Gen. Fil.
222.— Type: CUMING s.n., Philippines
seen). — Fig. 16k-I.
(1852)
(not
1981]
THELYPTERIDACEAE (Holttum)
535
Caudex long-creeping, 2-3mm_ diameter,
covered with short pale hooked hairs; scales c.
3mm long, narrow, bearing many short straight
hairs. Stipe of sterile fronds 7-20cm long, of
fertile 20-50cm, + tinged with dull red, rather
sparsely hairy, hairs as caudex. Lamina consisting
of an apical section and one pair of opposite
pinnae attached 0.5-2.0 cm below it. Sterile apical
lamina 10-18cm long, 2.5-4.0cm wide; base al-
most symmetrical, broadly cuneate to rounded,
rarely somewhat cordate; apex short-acuminate;
edges entire or slightly sinuous; main veins 4—
4.5 mm apart along midrib; veins 10-12 pairs, pale,
slender, slightly prominent on lower surface, at a
broad angle to main veins, excurrent veinlets free
or meeting the junction of the next pair of con-
nivent veins; lower surface of midrib and main
veins bearing pale hooked hairs 0.3-0.4 mm long,
rest of surface glabrous, or with a few short erect
hooked hairs, slightly pustular between veins;
upper surface of midrib bearing copious short
hooked hairs, hairs on main veins sparse. Fertile
apical lamina 1.0-2.5cm wide; veins to 7 pairs,
almost at right angles to main veins; excurrent
veinlets almost always short and free; sori
extending all along each vein so that sporangia are
distributed in a crescent-shaped row along each
pair of connivent veins; sporangia bearing hooked
hairs distally and sometimes a hair of 2 cells on
the stalk; spores bearing many separate but not
crowded small wings. Sterile pinnae 5-10 cm long,
1.5-3.0 cm wide, with stalks 1-3 mm long, shaped
as apical lamina but asymmetric, the lamina wider
on basiscopic side of costa than on acroscopic;
veins to c. 8 pairs on basiscopic side. Fertile
pinnae 1.0-2.5 cm wide, sori as apical lamina.
Distr. Tropical and subtropical mainland S.E.
Asia, Taiwan, Ryukyu Islands; throughout
Malesia; N. Queensland, Fiji (only 1 record from
Fiji).
Ecol. Low altitudes in light shade, sometimes
abundant under fruit trees or palms in villages;
few collections from Philippines and New Guinea.
52. Pronephrium x parishii (BEDD.) HOLTTUM,
Blumea 20 (1972) 123.— Meniscium parishii
BEDD. Ferns Brit. India (1866) t. 184. — Menis-
cium triphyllum var. parishii BEDD. Handb. (1883)
399. — Abacopteris triphylla var. parishii (BEDD.)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938)
241; HOLTTUM, Rev. FI. Malaya 2 (1955) 287. —
Thelypteris triphylla var. parishii (BEDD.) K.
IwaTs. Mem. Coll. Sci. Univ. Kyoto B, 31 (1965)
191. — Thelypteris parishii (BEDD.) PANIGRAHI,
Phytologia 31 (1975) 372.— Type: PARISH 135,
Burma, Moulmein (K).
Caudex long-creeping, to 4mm diameter, as P.
triphyllum in hairs and scales (hairs on scales of
some Indian specimens not hooked). Lamina
variable; apical section about as in P. triphyllum
but often with 1 or 2 narrow lobes at the base, or
1-2 small narrow adnate free pinnae just below it,
no buds at the bases of upper pinnae; pinnae to 6
pairs, usually opposite but sometimes not, usually
increasing in size from apex to base of frond,
upper ones always + adnate to rachis at basis-
copic base; largest 15 x 2-3cm, sometimes with
lamina symmetrically divided by the costa, some-
times the two halves unequal in width as in P.
triphyllum; pubescence as in P. triphyllum but
more numerous erect hooked hairs often present
on lower surface between veins. Sori as in P.
triphyllum; sporangia sometimes with longer’
hooked hairs; spores not seen.
Distr. Assam, Burma, Thailand, Vietnam; in
Malesia: Malaya (Perak and Pahang).
Ecol. In Malaya along stream banks in rather
open forest, at low altitudes.
Notes. The specimens referred here are vari-
able, and in their small adnate upper pinnae agree
with Meniscium thwaitesii HOOK. of Ceylon, but
the pinnae of the latter are more irregular and
some of its hairs are not hooked. Variability
among the present specimens suggests hybridity,
one parent certainly being P. triphyllum; it is
possible that plants in different areas have
different genetic origins.
53. Pronephrium salicifolium (WALL. ex HOOK.)
HOLTTUM, Blumea 20 (1972) 123.— Meniscium
salicifollum WALL. ex Hook. Ic. Pl. 10 (1854) t.
990; BEDD. Ferns Brit. India (1866) t. 207; Handb.
(1883) 399.— Phegopteris salicifolia (WALL. ex
Hook.) METT. Fil. Lechl. 2 (1859) 22; v.A.v.R.
Handb. (1908) 511.—Abacopteris salicifolia
(WALL. ex HooK.) HOLTTUM, Rev. FI. Malaya 2
(1955) 288, f. 167.— Thelypteris — salicifolia
(WALL. ex HOOK.) REED, Phytologia 17 (1968)
311.— Type: WALLICH 63, Penang (K).
Caudex short-creeping, 7-8 mm diameter, bear-
ing sparse hooked hairs. Stipe 20-60 cm long; near
base a few hooked hairs and rigid dark scales to
10 x 1 mm bearing pale hooked hairs, rest of stipe
glabrous, glossy, + castaneous. Lamina 30-80 cm
long, very firm, somewhat dimorphous, slightly
flushed with red when dry; apex pinna-like, not or
little larger than distal pinnae; pinnae to 18 pairs,
a bud sometimes present at base of uppermost
pinna; basal pinnae asymmetric at base, the blade
decurrent to the attachment to rachis on acros-
copic side, ending 5mm or more from rachis on
basiscopic side. Middle pinnae commonly 10—
15cm long, fertile ones 0.6-1.2cm wide, sterile
sometimes wider, largest sterile pinnae 20
2.5cm, fertile rarely to 2cm wide; base always
very narrowly cuneate; apex narrowly acuminate
but not caudate; edges quite entire; costules 3-
4mm apart, at 45° to costa; veins 3-10 pairs
(according to width of pinna), almost all anas-
tomosing, not prominent on either side, excurrent
veins all free except distal ones; both surfaces
quite glabrous. Sori supramedial, elongate along
536
FLORA MALESIANA
[ser. II, vol. 1°
veins, those on connivent veins usually meeting;
no hairs on body of sporangia and none seen on
stalk; spores with a + continuous wing and cross-
wings.
Distr. Malesia: Malaya, Sumatra, Borneo.
Ecol. On rocks in and beside streams, in
forest, in flood zone, at 0-500m; plants in the
more exposed places on rocks in a stream-bed
have smallish fronds with narrow ‘pinnae. The
very firm narrow pinnae are probably an adap-
tation to dry conditions when the stream is low
and also to survival in the periodic rush of flood-
water; in form they closely resemble the pinnae of
Dipteris lobbiana (HOOK.) MOORE, which grows
in similar habitats.
54. Pronephrium rubicundum (v.A.v.R.) HOLT-
TUM, Blumea 20 (1972) 123. — Phegopteris rubi-
cunda v.A.v.R. Bull. Jard. Bot. Btzg III, 2 (1920)
162. — Abacopteris rubicunda (v.A.v.R.) HOLT-
TUM, Rev. Fl. Malaya 2 (1955) 292, f. 170.—
Thelypteris rubicunda (v.A.v.R.) K. IWATS.
Mem. Coll. Sci. Univ. Kyoto B, 31 (1965) 195. —
Type: BROOKS 232/S, Sumatra, Lebong Tandai
(BO; BM). — Fig. lo, 16a-—d.
Caudex short-creeping, to 8 mm diameter, bear-
ing hooked hairs. Young plants bearing simple
fronds up to 30x 12cm, sometimes with fertile
ones 20 x 5 cm; later fronds on these plants having
lateral pinnae, the apical lamina progressively
smaller. Stipe 20-30 cm long on sterile fronds, to
70cm on largest fertile ones, base bearing thin
narrow scales to c. 8X1mm with a variable
number of hooked hairs on them, above base
+ flushed with dull red and bearing sparse pale
short hooked hairs. Lamina 20-35 cm long, firm,
drying dull reddish, consisting of an apical
segment and 1-3 pairs of opposite pinnae;
no buds at bases of pinnae; all pinnae asym-
metric, broader on basiscopic side of costa
than on acroscopic; basal pinnae short-stalked.
Apical lamina of fronds with 3 pairs of pinnae
15-20 x 5-8 cm (proportion of length to width very
variable), base broadly rounded to broadly
cuneate, apex short-acuminate, edges entire to
rather strongly undulate. Largest pinnae usually at
a wide angle to rachis, 10-24 x3-7cm, widest
about the middle or sometimes above the middle,
base cuneate to rounded, apex abruptly caudate
(cauda commonly to 2cm long), edges entire
(sterile) or undulate (most fertile ones); costules
3.5mm apart (fertile) or 4.5mm (sterile); veins
12-15 pairs, slightly prominent on lower surface,
where Sterile at 45° to costules, where fertile
closer and almost at right angles to costules,
excurrent veinlets all free; lower surface of rachis
and costae sparsely to rather copiously hairy,
most abundantly on young plants, hairs all
hooked, similar hairs sometimes present on sur-
face between veins; upper surface of rachis and
costae bearing + abundant pale hooked hairs less
than 0.5mm long. Sori medial, + elongate, those
on connivent veins often uniting; sporangia usu-
ally with 2 hooked hairs distally, hairs on stalk not
seen; spores with rather narrow wing and a few
cross-wings.
Distr. Peninsular Thailand, in Malesia: Suma-
tra, Malaya.
Ecol. In primary forest at 0-800m, not on
stream-banks.
55. Pronephrium = sulawesiense (K. IWATS.)
HOLTTUM, sp. nov.—P. rubicundum _ ssp.
sulawesiense K. IWATS. Acta Phytotax. Geobot.
28 (1977) 162, f. 2.— Type: K. SOMA et al. 218,
249, Central Celebes, inland from Polo, Kelawi
Distr. 1000-1700 m (TUS, not seen).
Differs from P. rubicundum as follows: scales
on rhizome 4X 2.5mm, on stipe 5 x 1.8 mm, lack-
ing hooked hairs; pinnae not always opposite, to
24 x 4.5 cm including apical cauda 4 cm long, more
oblique to rachis, distal ones longest, apparently
almost symmetric in shape, widest 1/3 from apex,
edges of fertile pinnae strongly crenate-serrate
distally, of sterile ones less strongly; sori medial,
those on connivent veins not meeting; sporangia
lacking hooked hairs.
Distr. Malesia: Central Celebes, only known
from the type.
Notes. This is geographically remote from P.
rubicundum and differs in scales, proportionately
narrower more oblique more symmetrical pinnae
with different edges. The figure shows a fertile
frond with distinctly alternate pinnae. In frond-
form this is nearer to P. ramosii of the Philippines
but differs in other characters.
56. Pronephrium cuspidatum (BL.) HOLTTUM,
Blumea 20 (1972) 123, excl. syn. praeter Bl. —
Meniscium cuspidatum BL. En. Pl. Jav. (1828)
114; Fl. Jav. Fil. (1828) 102, t. 45; RActB. Fl. Btzg
1 (1898) 65. — Phegopteris cuspidata (BL.) METT.
Farngatt. [TV (1858) 25; v.A.v.R. Handb. (1908)
511. — Dryopteris cuspidata (BL.) CHRIST, Philip.
J. Sci. 2 (1907) Bot. 205, p.p.; BACKER & POSTH.
Varenfl. Java (1939) 63, f. 10. — Abacopteris cus-
pidata (BL.) CHING, Bull. Fan Mem. Inst. Biol.
Bot. 8 (1938) 245.— Cyclosorus cuspidatus (BL.)
CopEL. Gen. Fil. (1947) 142; Fern Fl. Philip.
(1960) 375, nomen tantum.— Thelypteris cuspi-
data (BL.) K. IwaTs. Mem. Coll. Sci. Univ.
Kyoto B, 31 (1965) 192.—Type: BLUME, Java
(L).
Dryopteris amaiensis ROSENST. Meded.
Rijksherb. n. 31 (1917) 6. — Type: HALLIER 3157,
Java (L).
Caudex short-creeping, 7-l10mm_ diameter,
bearing hooked hairs. Stipe to 50 cm or more long,
dark at base with sparse hooked hairs and firm
dark brown scales c. 101mm bearing a few
hooked hairs, above base light-castaneous,
glabrous. Lamina commonly 30-5S0cm long (to
1981]
THELYPTERIDACEAE (Holttum)
537
120 cm, fide BACKER & POSTHUMUS), firm, dry-
ing dull reddish; apical segment of similar size to
upper pinnae, base cuneate and slightly asym-
metric; pinnae commonly to 6 pairs (10 pairs on
type), lowest usually opposite, the rest often not
so, a bud always at base of highest pinna only of
old fronds; basal pinnae with stalks to 5 mm long,
slightly asymmetric on either side of costa,
otherwise like the rest. Pinnae commonly to 16 x
3.5cm, sometimes dimorphous, smallest fertile
ones 9X 2cm, largest sterile 20 x 6cm, symmetric
on either side of costa, usually widest 1/4-1/3
from base, base narrowly to broadly cuneate,
slightly asymmetric with the acroscopic edge
decurrent to junction with rachis, the basiscopic
edge less decurrent; apex caudate-acuminate to
abruptly cuspidate (narrow tip 1.5—2.5cm long);
edges entire or nearly so; costules 3.5—-4 mm apart,
at 45° to costa, grooved above when dry and
slightly prominent beneath; veins 10-12 pairs, not
prominent either side, dark on lower surface, at a
narrower angle to costules in sterile than in fertile
fronds, excurrent veinlets mostly free and ending
in a conspicuous hydathode on upper surface
(with white incrustation on old fronds); lower
surface glabrous throughout or (especially in
Sumatra) with a few hooked hairs on costae and
costules; upper surface quite glabrous, even in
groove of rachis. Sori near costules, somewhat
elongate, those: on connivent veins near costa
uniting, distal ones sometimes not; sporangia
lacking hooked hairs on Java specimens, such
hairs present on specimens from Sumatra.
Distr. Malesia: Sumatra, Borneo, Java and
Lesser Sunda Is. (Flores).
Ecol. On rocks or steep valley-sides in forest
at altitudes to 1200 m.
Note. There may be some introgressive hybri-
dization between this species and P. rubicundum
in Sumatra.
57. Pronephrium ramosii (CHRIST) HOLTTUM,
comb. nov. — Dryopteris ramosii CHRIST, Philip.
J. Sci. 2 (1907) Bot. 203.— Lectotype (selected
here): RAMOS BS 1792, Luzon, Rizal Prov. (P; FI,
K).
Cyclosorus cuspidatus sensu COPEL. Fern FI.
Philip. (1960) 375, excl. syn. Meniscium cuspida-
tum BL. — Fig. 16e-f.
a. var. ramosii.
Differs from P. cuspidatum as follows: scales
at base of stipe to 4 x 1 mm, lacking hooked hairs;
pinnae always alternate, including basal ones, in
almost all cases widest above the middle and
narrowly cuneate at base (exceptions are speci-
mens from Mindoro and Sibuyan), commonly c.
11 x2.5cm, buds present at the bases of several
pinnae (to 4 on each side of rachis); veins 6-8
pairs, no conspicuous hydathodes present at
apices of excurrent veinlets; hooked hairs usually
more abundant on lower surface of costae and
costules, often present between veins on lower
surface, present also in groove of upper surface of
costae and in groove of rachis at base of a costa;
sori medial; sporangia always bearing hooked
hairs.
Distr. Malesia: Philippines (Luzon to Min-
danao), Moluccas (Halmahera), N.E. New Guinea.
Ecol. “Common at 600-800 m along streams in
damp gulches” (COPELAND).
b. var. minahassae HOLTTUM, var. nov.
A var. ramosii differt: pinnis usque 23x4cm
(steriles), 18 X 3.5 cm (fertiles); pilis hamatis sub-
tus inter venas carentibus ; sporangiis pilis hamatis
non praeditis. —Type: ALSTON 16551, N.
Celebes, G. Manembo-nembo (BM).
Distr. Malesia: N. Celebes.
Note. Meniscium liukiuense CHRIST, which I
listed in the synonymy of P. cuspidatum in 1972,
should also be included in P. ramosii, perhaps as a
distinct variety; its excurrent veins are mostly not
free. In the Solomon Islands is another possibly
distinct variety which has no hooked hairs on
sporangia and lower surface; they occur only at
the base of costae on the upper surface.
19. NANNOTHELYPTERIS
HOLTTUM, Blumea 19 (1971) 38; Kalikasan 2 (1973) 64. — Fig. 17.
Caudex short-creeping; fronds somewhat dimorphous, sterile ones with
shorter stipes and larger pinnae than fertile; pinnae 16-35 pairs, not more
than 2.5 cm long, close together except in some fertile fronds, lower pinnae
more widely spaced but hardly reduced; veins free or the basal ones
anastomosing; lower surface of pinnae between veins + pustular when dry,
not glandular, in some cases with minute acicular hairs; upper surface
between veins glabrous or with minute erect acicular hairs; sori indusiate or
exindusiate; indusia bearing hairs or glands; sporangia bearing spherical
538
FLORA MALESIANA
[ser. II, vol. 1°
yellow glands, as in Sphaerostephanos; spores with a +
some Cross-wings.
complete wing and
Type species: Nannothelypteris aoristisora (HARR.) HOLTTUM.
Distr. Malesia: Philippines (Luzon, Panay, Samar, Mindanao).
Ecol. In forest, in some cases on rocks.
Cytol. n= 36 (M. G. PRICE, N. camarinensis).
Taxon. A natural group of species closely related to Pronephrium sect. Dimorphopteris, with which it
might be united. No transference is here made to Pronephrium because of the uncertain status of the
latter.
COPELAND’s treatment of these species in his Fern Flora of the Philippines is confused because he
did not examine details. Some earlier confusion was caused by the inclusion by CUMING of some
specimens of Sphaerostephanos lastreoides in his n. 251,
nothelypteris philippina.
together with the type collection of Nan-
KEY *tO THE SPECIES
1. One pair of veins regularly anastomosing, at least near bases of larger pinnae; short hairs present between
veins on upper surface.
2. Sori exindusiate; pinnae less than 2 cm long
2. Sori indusiate; pinnae to 3 cm long
1. N. aoristisora
. 2. N. camarinensis
Veins all free, or with casual slight anastomosis; upper surface between veins glabrous, or nearly so.
S Pinnae to 1.3cm long, almost entire; costules mostly forked,
sometimes anastomosing
the branches of adjacent ones
3. N. nervosa
3. Pinnae (at least sterile ones) to 2cm or more e long, distinctly lobed at least o on acroscopic side; veins
all free.
4. Sterile pinnae to 2.5 x 1.0cm, lobed equally both sides half-way to costa or more deeply, not
conspicuously auricled; indusia setose
4. N. philippina
4. Sterile pinnae usually not over 2 cm long, 5-6 mm n wide above the auricled base, lobed more deeply
on acroscopic than on basiscopic side; indusia glandular
1. Nannothelypteris aoristisora (HARR.) HOLTTUM,
Blumea 19 (1971) 38; Kalikasan 2 (1973) 65.—
Polypodium aoristisorum HARR. J. Linn. Soc.
Bot. 16 (1897) 30.— Phegopteris aoristisora
(HARR.) v.A.v.R. Handb. (1908) 501.— Cyclo-
sorus aoristisorus (HARR.) COPEL. Gen. Fil.
(1947) 142; Fern Fl. Philip. (1960) 355, p.p.—
Thelypteris aoristisora (HARR.) REED, Phytolo-
gia 17 (1968) 260.—Type: STEERE, Panay
(MICH; K).
Stipe 3-12 cm long (longest in fertile fronds),
short-hairy; lamina to 20cm long; pinnae to 25
pairs. Sterile pinnae to 1.7 X0.6cm; base slightly
to distinctly auricled on acroscopic side, rounded
on basiscopic; apex obtusely pointed to rounded;
edges crenate to lobed, more deeply on acroscopic
than on basiscopic side; costules 2.5mm apart;
veins 2-3 pairs, basal pair anastomosing except
near apex of pinna; lower surface of rachis den-
sely short-hairy, hairs more sparse on costules and
veins; upper surface of rachis with thicker and
more sparse hairs, surface of pinnae between
veins bearing scattered very short erect hairs.
Fertile pinnae 1.1 x 0.35 cm; sori at maturity filling
lower surface, exindusiate.
Distr. Malesia: Philippines (Panay; Luzon: Mt
Makiling).
2. Nannothelypteris camarinensis HOLTTUM, Kal-
5. N. inaequilobata
ikasan 5 (1976) 119.— Type: M. G. Price 3101,
Luzon, Camarines Norte, Basud, Cone Mt (PNH;
K). — Fig. 17e-f.
Sterile fronds of young plants: stipe 3-6cm
long; lamina 15cm long with 10-16 pairs of pin-
nae; pinnae to 2.0cm long, 0.4cm wide above a
rounded basiscopic auricle, edges subentire, apex
rounded. Larger fronds, partially fertile: stipe S—
10cm long; pinnaé to 2.5 x 0.7 cm, basal auricles
smaller; apex more narrowly obtuse; edges stron-
gly crenate; costules 3mm apart at 45° to costa;
veins to 4 pairs, basal pair anastomosing; lower
surface of rachis bearing copious pale hairs
0.3mm long, shorter hairs on costae with scat-
tered thick ones 0.5mm long, minute hairs on
surface between veins; upper surface of rachis
with more sparse and thicker hairs, whole surface
of pinna bearing abundant short erect hairs. Fully
fertile fronds: stipe slender, to 22 cm long; pinnae
to 1.5x0.5cm; sori medial, slightly elongate on
the veins, with short-hairy indusia and glandular
sporangia.
Distr. Malesia:
known from the type.
Ecol. On forested rocky slope at 250-350 m.
Philippines (Luzon), only
3. Nannothelypteris nervosa (FEE) HOLTTUM,
Kalikasan 2 (1973) 66.— Phegopteris nervosa
FEE, Gen. Fil. (1852) 244; 6e Mém. (1853) 13, t. 2,
1981]
THELYPTERIDACEAE (Holttum)
539
NAA
sin
SS
\y
AM
TARR
AN
VN
ae
oe EID)
Fig. 17. Nannothelypteris nervosa (FEE) HOLT-
TUM. a. Frond, <3; b. pinna, x 3.—N. philippina
(PR.) HOLTTUM. c. Pinna, x 2.—N. inaequilo-
bata HoxLttum. d. Sterile pinna, x2.—N.
camarinensis HOLTTUM. e. Pinna, x2; f. sorus,
x 24 (a-b PRICE & HERNAEZ 71, c isotype K, d
holotype, e-f holotype).
f. 4.— Aspidium exiguum KUNZE ex METT.
Farngatt. IV (1858) 76, var. a. — Lastrea nervosa
(FEE) COPEL. Philip. J. Sci. 81 (1952) 32: Fern FI.
Philip. (1960) 323, nomen tantum. — Type: Cum-
ING S.n., Philippines (FEE’s specimen not seen:
specimens at B, BM, G, K agree with FEe’s figure
and are perhaps isotypes).
Polystichum —auriculatum var. — nervosum
CHRIST, Bull. Herb. Boiss. 6 (1898) 192: v.A.v.R.
Handb. (1908) 165.
Cyclosorus aoristisorus sensu COPEL. Fern. FI.
Philip. (1960) 355, p.p. — Fig. 17a—b.
Fronds almost uniform, but stipe of sterile ones
7-10 cm long, fertile 15 cm. Lamina to 24 cm long;
pinnae 30-40 pairs. Largest pinnae 1.3cm long,
3-3.5mm _ wide above the dilated base which is
44.5 mm wide (base more auricled on acroscopic
side in fertile than in sterile pinnae); apex obtuse;
edges entire or slightly crenate distally; costules
1.5mm apart, simply forked or with one branch
forked again, in basal acroscopic lobe pinnate with
2-3 pairs of veins; veins free or with casual anas-
tomosis; lower surface of rachis densely short-
hairy, hairs sparse on costae and costules; upper
surface of rachis with more sparse and shorter
hairs than lower, sparse minute hairs on costae.
Sori near margins of pinnae, on acroscopic branch
of each costule; indusia small, hairy.
Distr. Malesia: Philippines
Samar), at low altitudes.
Notes. The name Lastrea exigua J. SM., on
which Aspidium exiguum KUNZE ex METT. was
based, was published (without description) with
citation of CUMING 251 and 272, types of N.
philippina and N. inaequilobata; Mettenius cited
P. nervosa FEE under his var. a. If transferred to
Thelypteris, N. nervosa will need another specific
epithet, as T. nervosa (KLOTZSCH) TRYON
represents another species.
(Mindanao,
4. Nannothelypteris philippina (PRESL) HoLt-
TUM, Kalikasan 2 (1973) 66.— Physematium
philippinum PRESL, Epim. Bot. (1851) 192:
HOLTTUM, Novit. Bot. Univ. Carol. Prag. 1968
(1969) 44, excl. syn. Phegopteris nervosa FEE. —
Thelypteris philippina (PRESL) CHING, Bull. Fan
Mem. Inst. Biol. Bot. 10 (1941) 283. — Type:
CUMING 251, Luzon, S. Ilocos (PRC; G, K, P,
US).
Dryopteris confusa CopEL. Philip. J. Sci. 8
(1911) Bot. 146. — Lastrea confusa COPEL. Fern
Fl. Philip. (1960) 324.— Thelypteris confusa
(COPEL.) REED, Phytologia 17 (1968) 268.—
Type: CUMING 251 partim (M, destroyed); neo-
type (HOLTTUM 1973): RAMOs BS 8271 (MICH;
US).
Aspidium exiguum KUNZE ex METT. Farngatt.
IV (1858) 76, var. b, p.p.— Dryopteris exigua
sensu v.A.v.R. Handb. (1908) 190. — Fig. 17c.
Fertile and sterile fronds almost alike. Stipe c.
20 cm long, pale, copiously short-hairy. Lamina to
540
30cm long; pinnae to 20 pairs, lower ones more
widely spaced and slightly reduced. Largest pin-
nae 2.5 1.0cm, gradually narrowed from trun-
cate base to abruptly obtuse or rounded apex,
lobed equally on both sides 1/3-1/2 towards costa;
lobes oblique, basal acroscopic one often slightly
enlarged; costules 2.5—-3 mm apart; veins 3-4 pairs,
basal acroscopic one ending at base of sinus,
basiscopic one passing to margin above base of
sinus; lower surface of rachis bearing coarse pale
hairs 0.3-0.5 mm long, hairs on costae very short
with a few longer ones, sparse on costules; upper
surface of rachis as lower, hairs on costae minute.
Sori inframedial; indusia small, setiferous:
sporangia bearing glands.
Distr. Malesia: Philippines (Luzon).
Note. In some herbaria specimens of CUMING
251 and 272 are mounted without distinction on
the same sheet. The number 251 here refers to the
specimens at Prague and elsewhere as indicated,
but at BM the specimen of the present species
bears the number 272.
5. Nannothelypteris inaequilobata HOLTTUM,
Kalikasan 2 (1973) 67.—N. nervosa sensu
HOLTTuUM, Blumea 19 (1971) 38, p.p.— Type:
FLORA MALESIANA
[ser. II, vol. 1°
CUMING 272, Luzon (K; B, E, G, L, P).
Aspidium exiguum KUNZE ex METT. Farngatt.
IV (1858) 76, var. b, p.p. — Fig. 17d.
Fronds distinctly dimorphous; pubescence as N.
philippina. Sterile fronds: stipe 5S-9cm_ long;
lamina to 20cm long, pinnae 25 pairs or more,
lower 3-4 pairs not reduced but more widely
spaced, lowest with free basal acroscopic lobe;
largest pinnae 1.5-2.0cm long, 0.5—0.6cm wide
above base, base rather narrowly rounded, on
basiscopic side, auricled on acroscopic, basiscopic
margin slightly crenate distally, acroscopic margin
deeply crenate; costules 2 mm apart, very oblique,
veins 2 pairs (3-4 pairs on basal auricle), basal
acroscopic vein ending at base of sinus. Fertile
fronds: stipe to 15 cm long; lamina 20-30 cm long,
pinnae to 30 pairs, lower ones more widely spaced
than in sterile fronds; largest pinnae 1.0-1.3cm
long, to 0.4mm wide above base, on acroscopic
side lobed 1/2 way to costa, costules less than
2mm apart; sori inframedial, indusia bearing
glands, not hairs.
Distr. Malesia: Philippines (Luzon: Mt Makil-
ing and Nueva Vizcaya Province).
Ecol. On Mt Makiling at 300m, near mud-
springs in open place in forest.
20. STEGNOGRAMMA
BLUME, En. Pl. Jav. (1828) 172; CHING, Sinensia 7 (1936) 90; Acta Phy-
totax. Sinica 8 (1963) 329; COPEL. Gen. Fil. (1947) 144; K. IwaTs. Acta
Phytotax. Geobot. 19 (1963) 112; Mem. Coll. Sci. Univ. Kyoto B, 31 (1964)
18-21; Amer. Fern J. 54 (1964) 141; HoLTrumM, Blumea 19 (1971) 38. —
Leptogramma J. Sm. in Hook. J. Bot. 4 (1841) 51; Hist. Fil. (1875) 231;
CHING, Sinensia 7 (1936) 96. — Fig. 18.
Caudex short-creeping or erect; stipes densely hairy, hairs unicellular
(pluricellular in sect. Haplogramma, not Malesian); scales bearing uni-
cellular acicular hairs on edges and surface; fronds simply pinnate; apical
lamina always deltoid and deeply lobed with basal basiscopic veins of some
of the middle or lower lobes springing directly from the rachis; pinnae
subentire to deeply lobed, upper ones always + adnate to rachis, in some
species only the lowest fully free; lowest pinnae not or little reduced;
spherical glands lacking; hairs between veins on upper surface few, rather
long, not appressed; veins free or anastomosing; sori exindusiate, extending
along the veins (where veins anastomose, sori also along the intermediate
excurrent veins between costules); sporangia setiferous; spores with many
small wings.
Type species: Stegnogramma aspidioides BL.
Distr. Pantropic (excluding New Guinea and the Pacific) and subtropic, c. 18 spp.; in Malesia on
mountains in Sumatra, Java, Borneo, Celebes, Luzon, Mindanao.
Cytol. Chromosome number 36 in sect. Leptogramma (diploid in Mexico, Ceylon and Japan;
tetraploid in Madeira): in other sections unknown.
Taxon. IWATSUKI’s arrangement of 1963 is here adopted. In it he recognized four sections, two of
1981] THELYPTERIDACEAE (Holttum) 541
Fig. 18. Stegnogramma aspidioides BL. a. Pinna, <1; b. part of fertile pinna showing extent of sori,
x 3. — S. subcalcarata (v.A.v.R.) HOLTTUM. c. Pinna, x 1; d. venation and sori, * 6; e. part of sorus,
x 24 (a—b T. LOBB s.n., c-e isotype K).
which (Haplogramma and Dictyocline) do not occur in Malesia. The New World species (see
IWATSUKI 1964) are not yet well studied. In Malesia, the widely-distributed species S. pozoi is
represented by a variety in Java; the other plants of sect. Leptogramma, from four widely-separated
small areas, are here regarded as representing separate species, but more material is needed to
characterize them clearly; only S. dissitifolia has been well collected.
Stegnogramma is related to those species of Sphaerostephanos which have no spherical glands, but is
distinct in the combination of characters mentioned in the generic description. In the 19th century,
species of this genus were usually placed in Gymnogramme.
The peculiar Hawatian species Pneumatopteris sandwicensis (BRACK.) HOLTTUM shares with
Stegnogramma the characters of venation of the apical lamina and few thick hairs between veins on the
upper surface of pinnae, and has somewhat elongate sori, but differs in sporangia and spores.
KEY TO THE SPECIES
1. Veins anastomosing (sect. Stegnogramma) .......... .. . . . 41.8. aspidioides
1. Veins free (sect. Leptogramma).
2. Pinnae to 7.5 1.6cm or yeti lower surface of costae bearing many hairs, short ones more
abundant than longer . . . .2. 8. pozoi var. petiolata
2. Pinnae, if over 4cm long, not more than if cm . wide: lower surface of costae bearing rather sparse
hairs 0.5 mm or more long and a few short ones.
3. Pinnae not over 1.5 cm long, only basal ones lobed; sporangia sparsely setose 3. S. gymnocarpa
3. Pinnae longer, several pairs lobed; sporangia all freely setose.
4. Pinnae to 5.0 x 0.8 cm with narrowly triangular apex 1|O0mmlong .. . . . 4.8. subcalcarata
4. Pinnae proportionately wider with short entire apex.
5. Fronds to 30 cm long; basal pinnae lobed more than half-way towards costa .5. S. dissitifolia
5. Fronds c. 16cm long; basal pinnae less deeply lobed 6. S. celebica
1. Stegnogramma aspidioides BL. En. Pl. Jav. KAULF. 1824 nec DESv. 1827 nec BL. 1828; Ic.
(1828) 173.— Gymnogramme stegnogramma BL. PI. 10 (1854) t. 950; Spec. Fil. 5 (1864) 150, excel. pl.
Fl. Jav. Fil. (1829) 98, t. 44.—Gymnogramme Khasya & Ceylon; RACIB. FI. Btzg | (1898) 70. —
aspidioides HooK. Gen. Fil. (1841) t. 120B non Phegopteris stegnogramma METT. Fil. Hort. Lips.
542
(1856) 84; v.A.v.R. Handb. (1908) 508. — Dry-
opteris stegnogramma (BL.) C. CHR. Ind. Fil.
(1905) 294, p.p.; Gard. Bull. S. S. 7 (1934) 250;
BACKER & POSTH. Varenfl. Java (1939) 63.—
Thelypteris stegnogramma (BL.) REED, Phy-
tologia 17 (1968) 466. — Type: BLUME, Java.
Phegopteris stegnogramma var. meniscioides
v.A.v.R. Bull. Jard. Bot. Btzg II, 16 (1914) 27;
Handb. Suppl. (1917) 319.—Type: MATTHEW
584a, G. Singgalang, Sumatra (BO). — Fig. 18a—b.
Stipe to 40cm long, dark, bearing long pale
hairs mixed with short ones; basal scales c. 6
Imm; hairs on rachis to 2mm long. Lamina
commonly 55cm long; pinnae 8-10 pairs, adnate
to rachis except basal 2-3 pairs; lowest pinnae
slightly reduced, narrowed to base on basiscopic
side and slightly auricled on acroscopic, veins in
auricle forked and anastomosing; texture thin,
dark when dried. Largest pinnae commonly 10 x
3cm (to 133.5cm); aerophores swollen; base
truncate to subcordate; apex short-acuminate;
edges crenate; costules 5mm apart, at a wide
angle to costa; veins 8 pairs, 3 basal pairs anas-
tomosing with zig-zag excurrent veins, next 2-3
pairs to sides of a long sinus-membrane; lower
surface of costa bearing copious long spreading
hairs mixed with shorter ones, costules and veins
with shorter spreading hairs; rather sparse short
erect hairs on surface between veins; copious
antrorse hairs on upper surface of costae, shorter
spreading hairs on costules and veins, sparse erect
hairs between veins. Sori spread all along lower
veins and also on the excurrent intermediate veins
arising from them, usually on basal part only of
distal veins.
Distr. Malesia: West and Central Java; Suma-
tra (G. Kemiri, G. Singgalang, G. Kerinci); Sabah
(Mt Kinabalu). Wrongly recorded from Khasya
and Ceylon by HOOKER and BEDDOME.
2. Stegnogramma pozoi (LAGASCA) K. IWATsS.
Acta Phytotax. Geobot. 19 (1963) 124, var.
petiolata (CHING) HOLTTUM, comb. nov. — Lep-
togramma petiolata CHING, Acta Phytotax.
Sinica 8 (1963) 319.—Type: G. WALL (“WAL-
LICH’’), Ceylon (PE).
Gymnogramme aspidioides BL. En. Pl. Jav.
(1828) 112, non KAULF. 1824 nec DESv. 1827. —
Gymnogramme totta sensu BL. Fl. Jav. Fil.
(1829) 90, t. 38.— Grammitis blumeana PRESL,
Tent. Pterid. (1836) 209, nom. nov. — Phegopteris
totta sensu METT. Farngatt. IV (1858) 18, quoad
pl. javan. tantum:; v.A.v.R. Handb. (1908) 497,
p-p.; Handb. Suppl. (1917) 515, excl. var.
subcalcarata. — Leptogramma totta sensu BEDD.
Handb. (1883) 377, quoad pl. zeyl. tantum. —
Dryopteris africana (DESV.) C. CHR. Ind. Fil.
(1905) 251, p.p.; BACKER & PosTH. Varenfl. Java
(1939) 36.— Type: BLUME, Waterfall Tjikundal,
G. Gedeh, Java (L).
Stipe 15-30cm long; basal scales c. 5X 1mm.
FLORA MALESIANA
[ser. Il, volawe
Lamina 25-40cm long; pinnae 12-15 pairs of
which 4-5 pairs are free, the rest + adnate to the
rachis; basal 1-2 pairs in smaller fronds somewhat
reduced, at least the lowest with stalks | mm long
and reduced basal basiscopic lobe. Largest pinnae
seen 7 1.6cm; base subequally truncate; apex
entire, triangular, 8 x 3.5mm; edges lobed almost
half-way to costa, lobes oblique, hardly falcate,
entire; costules 4.5-S mm apart; veins 6 pairs (or
7 veins on basiscopic side of costule), basal veins
from adjacent costules usually both touching sides
of sinus-membrane which may be decurrent as a
ridge between them; hairs on lower surface of
costae abundant, spreading, of varied length,
longest 0.6-0.7 mm, costules and veins with shor-
ter hairs, copious erect short hairs between veins;
hairs on upper surface of costae mostly not over
0.5mm long with some to 1mm, sparse shorter
hairs on costules and veins, a few thick hairs
0.5mm long between veins. Sori on basal veins
from costule along more than half length of vein,
on distal veins shorter, medial.
Distr. Ceylon; in Malesia: Java.
Ecol. In Java on higher mountains, on steep
earth banks in forest (BACKER & POSTHUMUS);
in Ceylon at 1800 m.
Notes. The plants of Java and Ceylon here
included are closely related to S. pozoi, the type
of which was collected in northern Spain. Similar
plants found in Madeira are tetraploid (MANTON).
The latter differ from var. petiolata (diploid) in
having all pinnae except the lowest adnate to the
rachis, hairs on lower surfaces, confined to costae
and costules, uniformly | mm long, and basal sori
not extending to the bases of veins. Plants named
S. pozoi occur throughout Africa; they are vari-
able and no chromosome counts have been made.
BLUME’s type has pinnae to 4.0 x 1.3 cm, and all
pinnae above the basal ones are adnate; BACKER
& POSTHUMUS report plants with fronds to 50 x
25 cm but I have not seen any so large.
Prof. CHING (in lit.) informs me that the type of
Leptogramma petiolata bears the collector's name
WALL, which he interpreted as WALLICH, but it
was surely G. WALL, who collected ferns in Cey-
lon, not WALLICH.
3. Stegnogramma gymnocarpa (COPEL.) K.
IwatTs. Acta Phytotax. Geobot. 19 (1963) 122. —
Dryopteris gymnocarpa COPEL. in Elmer, Leafi.
Philip. Bot. 3 (1910) 807.— Phegopteris gym-
nocarpa (COPEL.) v.A.v.R. Handb. Suppl. (1917)
313.— Lastrea gymnocarpa COPEL. Gen. Fil.
(1947) 139; Fern Fl. Philip. (1960) 325, quoad typ.
tantum. — Leptogramma gymnocarpa (COPEL.)
CHING, Acta Phytotax. Sinica 8 (1963) 318.—
Thelypteris gymnocarpa (COPEL.) MORTON,
Amer. Fern J. 56 (1966) 179.—Type: ELMER
11508, Mt Apo, Falls of Cati Creek, 1750 m, Min-
danao (MICH; BO, K, L etc.).
Stipe 2-3cm long; hairs 1mm long rather
1981]
THELYPTERIDACEAE (Holttum)
543
sparse, many much shorter; scales little over 1 mm
long. Lamina 9-15 cm long (COPEL.), distal half
or more pinnatifid, free or adnate pinnae 3-4 pairs,
lowest with stalks 1mm long; basal pinnae 1.0 x
0.7 cm, distinctly lobed; other pinnae, and lobes of
terminal lamina, entire. Veins in lobes of basal
pinnae 2-3 pairs, in lobes of terminal lamina sim-
ple or forked; lower surface of rachis bearing
many hairs 1mm long and few short ones, on
costae and costules more sparse similar hairs, no
others; on upper surface. of rachis many short
hairs and fewer | mm long, hairs on costae, cos-
tules and between veins sparse, 0.5 mm long. Sori
on basal acroscopic veins on pinnae, sometimes
on both branches of a forked vein on apical
lamina; a majority of sporangia lacking setae.
Distr. Malesia: Philippines (Mindanao), only
known from type.
Note. COPELAND included the Luzon plants
here named S. dissitifolia.
4. Stegnogramma subcalcarata (v.A.v.R.) HOLT-
TUM, comb. nov. — Phegopteris totta var. sub-
calcarata v.A.v.R. Handb. Suppl. (1917) 515.—
Type: BECCARI 430, G. Singgalang, Sumatra,
1700 m (BO; FI, K, MEL). — Fig. 18c-e.
Stipe 10-12cm long; hairs 1mm long sparse,
with shorter ones; scales small, narrow. Lamina
to 18cm long; free pinnae 4-5 pairs, above these
4-5 pairs gradually more broadly adnate to rachis;
basal pinnae on some fronds reduced, in all cases
narrowed towards base on both sides (more so on
basiscopic), basal acroscopic lobe 2-3.5 mm long,
almost free, stalks 1.5-2 mm long. Largest pinnae
3.5-5cm long, 6.5-8mm wide; base broadly
cuneate; apex entire, triangular, 7-12 mm _ long,
2.5mm wide at base; edges lobed half-way to
costa, lobes subtriangular; costules 3 mm apart, at
less than 60° to costa; veins 4-5 pairs, acroscopic
basal vein passing to side of short sinus-mem-
brane, basal basiscopic vein to edge above base of
sinus; hairs on lower surface of costa sparse,
0.7 mm long with a few shorter ones, very few on
costules and veins, none between veins; on upper
surface hairs few apart from costa. Sori on basal
acroscopic vein almost from its base, medial on
other veins.
Distr. Malesia: Sumatra. Apart from type,
known from one poor specimen from 2000m on
G. Kerinci (BUNNEMEIJER 10480).
5. Stegnogramma dissitifolia HOLTTUM, - sp.
nov. — Lastrea gymnocarpa sensu COPEL. Fern
Fl. Philip. (1960) 325, p.p.
Stipes usque 18 cm longus; lamina usque 30 cm
longa; pinnae liberae usque 12-jugatae, inferiores
6-jugatae stipitatae, maximae 3.5 cm longae, basi
dilatata usque 1.2cm lata, supra basin 0.9 cm
latae, dimidio costam lobatae; pinnae infimae
valde dissitae. —Type: M. G. PRICE 1668, Baguio
City, Luzon (K).
Stipes 10-18cm long, bearing sparse hairs I-
1.5mm long and shorter ones; basal scales to
5x1lmm. Lamina to 30cm long; pinnae c. 12
pairs below deeply lobed apical lamina, c. 6 pairs
of lower pinnae distinctly stalked, stalks to
1.5mm; basal pinnae 2-3cm from next pair (in
another collection 5 cm), wider on basiscopic side of
costa than on acroscopic and lobed more than
half-way to costa, much narrowed to base on
basiscopic side. Largest pinnae 3.5cm_ long,
commonly to 0.9cm wide above dilated base
which is 1.0-1.2cm wide; base subtruncate to
broadly cuneate; apex abruptly short-pointed;
edges lobed about half-way to costa; costules
3-3.5(-4) mm apart; veins 3—4 pairs; on lower sur-
face of rachis sparse hairs 1mm long and many
much shorter, on costae sparse hairs 0.5 mm long
and a few short ones, sparse slender erect hairs
between veins; hairs on upper surface of rachis
more than 1 mm long with many short ones, costal
hairs 0.3-0.5mm, on costules fewer, scattered
thick hairs 0.3-0.5 mm long between veins. Sori
on basal half of basal veins, in middle of distal
veins; sporangia all with several setae.
Distr. Malesia: Philippines; mountains of
middle-north of Luzon at 1400-2700 m (7 collec-
tions).
Notes. Young plants of these Luzon coliec-
tions differ from the small plants of comparable
size from Mindanao which constitute the type of
S. gymnocarpa; the former have more pairs of
free pinnae, which are all lobed, and sporangia all
have more than one seta.
Leptogramma amabilis TAGAWA, based on
small plants from Okinawa (Ryukyu Is.) cannot be
effectively compared until better known. Plants
described as S. pozoi on Orchid Island (near the
south of Taiwan) have pinnae 5 cm long.
6. Stegnogramma celebica (CHING) HOLTTUM,
comb. nov.—Leptogramma celebica CHING,
Sinensia 7 (1936) 99, t. 5.—S. gymnocarpa
(COPEL.) K. IWATS. ssp. celebica K. IWATs.
Acta Phytotax. Geobot. 19 (1963) 123. — Thelyp-
teris bunnemeijeri REED, Phytologia 17 (1968)
265.— Type: BUNNEMEJJER 12073, G. Bonthain,
Celebes (NY; BO, L).
Stipe to 7cm long; hairs hardly 0.5 mm long.
Lamina to 16cm long; free pinnae 5 pairs, 5 pairs
adnate to rachis; lowest pinnae slightly reduced,
smallest seen 1.3cm long, with stalks 1 mm long,
narrowed to base on basiscopic side. Largest
pinnae 3 cm long, 7-8 mm wide; base broadly and
rather unequally cuneate; edges lobed, almost to
apex, less than half-way to costa, lobes small,
subtriangular; costules 2.5 mm apart, at little more
than 45°; veins 3 pairs, basal acroscopic veins
passing to side of sinus-membrane, basiscopic
vein to edge above base of sinus (in sterile fronds,
veins sometimes uniting just below sinus); hairs
on lower surface of costae rather sparse, on fertile
544
pinnae hardly 0.5mm long, on sterile pinnae
longer, few hairs on costules, none between veins
on fertile pinnae (a few on sterile ones); a few
hairs at least 0.5 mm long between veins on upper
surface. Sori along basal half of basal veins,
FLORA MALESIANA
[ser. II, vol. 1°
sometimes also on second veins.
Distr. Malesia: S.W. Celebes. Known from
two collections from about 2000m on G. Bon-
thain; the second is EVERETT s.n. Oct. 1895 (K,
SING).
21. AMPHINEURON
HOLTTUM, Blumea 19 (1971) 45; Blumea 23 (1977) 205. — Fig. 19.
Caudex erect, or short- or long-creeping; scales narrow, setiferous; stipe
minutely hairy, scaly at the base only except in A. distinctum; lamina often
very large, pinnate, pinnae in most species deeply lobed; basal pinnae much
narrowed at their bases; in a few species 1-2 (rarely 3) pairs of irregularly-
placed and -shaped small pinnae sometimes present below the normal ones;
aerophores at the bases of pinnae usually narrowly elliptic and (when dry)
discoloured, not swollen; veins pinnate in the pinna-lobes, simple, basal
veins either free and passing to the margin separately, or connivent at the
sinus-membrane, or anastomosing to form an excurrent vein, these con-
ditions sometimes not constant in a single frond; sinus-membrane usually
ending in a prominent tooth; short acicular hairs always present on some
part of the lower surface, also glandular hairs of varying size and shape,
spherical to pyriform or club-shaped, the larger ones sometimes collapsing
in drying to form resinous spots which may be faint and hardly detectable;
sori in most species medial or supramedial; indusia usually present, bearing
glands and/or hairs, apparently lacking in A. distinctum; glandular cells
often present on the stalks of sporangia, directly attached to the stalk or at
the end of a short hair, capitate hairs seen on the body of sporangia only in
A. subattenuatum; spores usually dark, irregularly rugose or with irregular
thick + branched ridges.
Type species: Amphineuron opulentum (KAULF.) HOLTTUM.
Distr. S.E. Asia; Malesia; Queensland; in the Pacific to Tahiti; East Africa; in all c. 12 species.
Cytol. Base chromosome number 36; A. opulentum (Singapore) and A. terminans (Ceylon) both
tetraploid; no experimental work reported.
Taxon. This genus resembles Christella in its spores and in its mainly elongate glandular hairs; the
latter are more varied than in Christella. The name Amphineuron is intended to indicate the fact that in
several species the anastomosis of veins is inconstant.
The two most widely-distributed species, A. opulentum and A. terminans, are both variable, and both
have a complex taxonomic and nomenclatural history. Hybrids between them probably exist, at least in
Thailand. As indicated in the key, the species may be divided into two groups. Rather large glands, of
varied form, occur on most species of the second group and are undoubtedly distinctive, but they are
often not well preserved on herbarium specimens and are sometimes not detectable, so that such
specimens are difficult to name with certainty, and I have found it impossible to be sure how many
species can be recognized. The present arrangement is tentative. Specimens dried without heat retain
their glands perfectly, as I have discovered when dealing with plants in cultivation at Kew. It may be
that alcohol, sometimes used to preserve specimens temporarily, also has an effect on glands in this
genus.
KEY TO THE SPECIES
1. Indusia rather large, persistent; lamina thin, veins slender, slightly prominent on both sides.
2. Caudex long-creeping; pinnae lobed half-way to costa or less deeply; apex of frond + pinna-like
1. A. terminans
1981]
THELYPTERIDACEAE (Holttum)
545
2. Caudex short-creeping or erect;
pinna-like.
pinnae lobed more than half-way to costa; apex of frond not
3. Pinnae lobed to | mm from costa or more deeply; basal veins free or uniting to form very short
excurrent veins.
4. Basal veins not meeting; sori in slight depressions
2. A. immersum
4. Basal veins meeting just below the sinus, sometimes forming. a short excurrent vein; sori not
immersed
3. A. subattenuatum
3. Pinnae lobed less deeply: ‘basal veins ; anastomosing to form an excurrent vein on most parts of a
frond
4. A. opulentum
. Indusia small, almost hidden by sporangia at maturity of s sorus, or “apparently lacking; lamina firm,
veins not prominent on either surface.
. Hairs more than 0.5 mm long present on both sides of costa; stipe and abaxial surface of rachis
bearing scales throughout
5. All hairs on costa very short; scales confined to base of stipe.
5. A. distinctum
6. Lower surface between veins (sometimes in part) bearing very short erect acicular hairs; short
capitate hairs also in this position on the upper surface
6. Both surfaces lacking hairs between veins.
6. A. attenuatum
7. Pinnae lobed to c. 2mm from costa, basal veins often anastomosing.
8. Many glandular hairs on lower surface between veins, some spherical, some + elongate; similar
hairs also on indusia
7. A. ceramicum
8. Glandular hairs less abundant, always spherical tending to collapse and form resinous patches or
to disappear on drying
8. A. pseudostenobasis
7. Pinnae lobed more deeply; basal veins not PAS eoroSite!
9. Pinnae thin, to 3cm wide; glands on lower surface of pinnae and on indusia elongate
9. A. paraphysophorum
9. Pinnae firm, to 1.5 cm wide; glands on lower surface glabular to pyriform, on indusia spherical,
resinous
1. Amphineuron terminans (HOOK.) HOLTTUM,
Amer. Fern J. 62 (1973) 82; Blumea 23 (1977)
207. — Nephrodium terminans HOOK. Spec. Fil. 4
(1862) 73, excl. syn. N. conioneuron FEE & Las-
trea malaccensis PRESL. — Thelypteris terminans
(HOOK.) TAGAWA & K. IwatTs. Acta Phytotax.
Geobot. 26 (1975) 169.—Type: WALLICH 386,
Burma, Kamoun (= Kumon) (K).
Nephrodium oreopteris FEE, Gen. Fil. (1852)
306, non (EHRH.) DESv. 1827. — Type: CUMING
48, Luzon (holo?; isotypes G, K, LE, W).
Thelypteris wagneri FosB. & SACHET, Smiths.
Contr. Bot. 8 (1972) 6, excl. syn. Polypodium
pteroides RETZ.— Type: RACIBORSKI, Java
(US).
Nephrodium pteroides sensu BEDD. Handb.
(1883) 269; RActB. FI. Btzg 1 (1898) 183. — Dry-
opteris pteroides sensu v.A.v.R. Handb. (1908)
209.
Dryopteris interrupta sensu BACKER & POSTH.
Varenfl. Java (1939) 56. — Cyclosorus interruptus
sensu HOLTTUM, Rev. FI. Malaya 2 (1955) 262, f.
149; CopEL. Fern Fl. Philip. (1960) 361. — Fig.
19a.
Caudex long-creeping, c. 5mm diameter (dry);
stipe commonly to 50cm long, + flushed dull
reddish, glabrescent abaxially, basal scales c.
8mm long. Lamina to 50cm long; pinnae to 25
pairs, basal pair somewhat reduced, rarely very
small, always distinctly narrowed at their bases;
apex of frond usually pinna-like but variable.
Largest pinnae commonly 17-20 x 1.7-2.0cm, if
longer not more than 2cm wide; base of middle
. 10. A. kiauense
pinnae broadly cuneate to truncate; apex acu-
minate; edges lobed 1/3 towards costa or less
deeply, lobes as wide as long (or wider) with
broad asymmetric apex and forward-pointing tip;
costules 4-5 mm apart, usually at less than 60° to
costa; veins 6-9 pairs, basal pairs spreading at a
broad angle to their costules and uniting to form a
rather long excurrent vein to the sinus, next veins
very oblique, 1 or 2 ending beside the sinus-
membrane; lower surface of rachis, costae, cos-
tules and veins bearing short acicular hairs, longer
hairs usually lacking, subsessile almost spherical
rather pale glandular hairs abundant on distal
veins, usually few and smaller on lower veins,
very short acicular hairs often present between
veins; upper surface of costae bearing antrorse
pale acicular hairs, similar hairs scattered on cos-
tules and veins. Sori close to margins of lobes, not
on lower veins; indusia large, thin, often with
some short acicular hairs and a few small glan-
dular hairs which are not marginal.
Distr. Ceylon & S. India; Burma to Hainan and
Macao; throughout Malesia; Queensland (to
18° S); one specimen from Central Africa and one
from Fernando Poo.
Ecol. In Malesia only abundant in areas with a
distinct dry season, in rather open but not too dry
places, spreading by long rhizomes.
Notes. The complex nomenclatural and tax-
onomic history of this species is set forth and
discussed in HOLTTUM 1977. It is probable that in
Thailand this species has become hybridized with
A. opulentum but I have not seen intermediates
546 FLORA MALESIANA [ser. II, vol. 1°
f Nn /
Ne % Se
PS y
i A fy \
Se ee ane = 3
t n
Fe SO TTY ES SEA poe
Femnhsesunscteunere eaubneeh
fubina tet asl ene)
JCD
Fig. 19. Amphineuron terminans (HOOK.) HOLTTUM. a. Venation and sori, xX 4.— A. opulentum
(KAULF.) HOLTTUM. b. Two pinna-lobes, showing differences in course of basal veins, x 3; c. sorus,
x 18.— A. subattenuatum (ROSENST.) HOLTTUM. d. Venation and sori, X 3.— A. immersum (BL.)
HOLTTUM. e. Venation and sori, x 3.— A. pseudostenobasis (COPEL.) HOLTTUM. f. Base of basal
pinna, X 1; g. two pinna-lobes, = 4; h. upper surface of costa with capitate hairs, x 16 (a FORMAN 145,
b-c SEEMANN S.n., d WOMERSLEY & HOLTTUM 17692, e MOUSSET 39, f-h HOOGLAND & CRAVEN
10122).
1981]
THELYPTERIDACEAE (Holttum)
547
from Malesia. FOSBERG and SACHET used the
pinna-like apex of fronds as the main distinguishing
character, ignoring the evidence of venation, hairs
and glands which appear to me more significant.
This is the only species of Amphineuron in which
anastomosis of veins is invariable.
2. Amphineuron immersum (BL.) HOLTTUM in
Nayar & Kaur, Comp. to Bedd. (1974) 203; Blu-
mea 23 (1977) 211.— Aspidium immersum BL.
En. Pl. Jav. (1828) 156; RActrB. FI. Btzg 1 (1898)
169. — Lastrea immersa (BL.) MOorRE, Ind. Fil.
(1857) Ixxxix; BEDD. Ferns Br. India (1867) t. 252;
Handb. (1883) 234; COPEL. Fern FI. Philip. (1960)
327. — Dryopteris immersa (BL.) O. KTZE, Rev.
Gen. Pl. 2 (1891) 813; v.A.v.R. Handb. (1908) 188;
BACKER & POSTH. Varenfl. Java (1939) 39.—
Thelypteris immersa (BL.) CHING, Bull. Fan
Mem. Inst. Biol. Bot. 6 (1936) 306; HOLTTUM,
Rev. Fl. Malaya 2 (1955) 243. — Parathelypteris
immersa (BL.) CHING, Acta Phytotax. Sinica 9
(1963) 303. — Type: BLUME, Java (L, n. 908, 335-
404).
Lastrea caudiculata PRESL, Epim. Bot. (1851)
36; HoLTTUM, Novit. Bot. Univ. Carol. Prag.
1968 (1969) 35. —Type: CUMING s.n. Philippines
(PRC).
Lastrea verrucosa PRESL, Epim. Bot. (1851) 36;
CoPEL. Fern FI. Philip. (1960) 327. — Thelypteris
verrucosa (PRESL) CHING, Bull. Fan Mem. Inst.
Biol. Bot. 6 (1936) 308.— Type: CUMING 72,
Luzon (PRC).
?Dryopteris diversifolia v.A.v.R. Bull. Dép. Agr.
Ind. Néerl. 18 (1908) 7; Handb. (1908) 189. —
Type: RAAP 27, Sumatra, Batu Isl. (BO).
Dryopteris besukiensis v.A.v.R. Bull. Jard. Bot.
Btzg II, 1 (1911) 7; Handb. Suppl. (1917) 156. —
Lectotype (HOLTTUM 1977): KOORDERS 15436,
Java, Besoeki (BO; L).
Thelypteris subimmersa CHING, Bull. Fan
Mem. Inst. Biol. Bot. 6 (1936) 306. — Parathelyp-
teris subimmersa CHING, Acta Phytotax. Sinica 8
(1963) 303. — Type: S. K. LAu 1395, Hainan (PE,
not seen). — Fig. 19e.
Caudex short, erect; stipe to 70cm or more
long, green when living; basal scales thin, to
15 mm long. Lamina to at least 120 cm long (often
fertile at a much smaller size); pinnae close, tex-
ture thin, drying pale-olivaceous; 1-2 pairs of
irregularly spaced small pinnae sometimes present
at base of frond. Largest pinnae of type 25 x 3. cm,
largest seen-45 x 5cm, lobed to 1 mm from costa
or more deeply; lobes, except distally, almost at
right angles to costa, separated by wide sinuses;
costules 3.5-6mm apart; veins 14-20 pairs, basal
acroscopic vein ending beside the short sinus-
membrane, basiscopic one passing to margin
above base of sinus; lower surface of costae and
costules of type lacking acicular hairs except near
apex of pinna, of other specimens bearing a vari-
able number of hairs up to 1 mm long, costules
and veins usually bearing small pale yellow glands
(abundant on the type), between veins sometimes
short erect acicular and capitate hairs; upper sur-
face with antrorse hairs on costae and scattered
hairs on costules and veins. Sori supramedial, in
depressions in the lamina (forming convexities on
the upper surface); indusia thin with a variable
fringe of short yellow capitate hairs.
Distr. Assam; Hainan; southern Thailand;
throughout Malesia; Queensland (to 16°S); New
Hebrides, New Caledonia.
Ecol. At low altitudes, in sheltered places but
not in deep shade, common on limestone in
Malaya, found also on stream banks.
Notes. All the living plants observed by me
have an erect caudex, but BACKER & POST-
HUMUS state ‘‘wortelstok kruipend”. ERYL
SMITH 2432, from Timor, is a small plant with
definitely creeping caudex; it might be a hybrid
with A. opulentum. The types of Dryopteris
diversifolia v.A.v.R. and D. besukiensis v.A.v.R.
are small plants, perhaps stunted owing to
exposure. There is much variation in_ the
development of acicular hairs and of small yel-
lowish glandular hairs on the lower surface of
pinnae. The types of both Lastrea caudiculata
PRESL and L. verrucosa PRESL have acicular
hairs, that of Thelypteris subimmersa CHING has
none, but the published description gives no other
character which would distinguish it from the type
of Aspidium immersum BL. I know of no other
record of this species in China.
3. Amphineuron subattenuatum (ROSENST.)
HOLTTuM, Blumea 23 (1977) 412.— Dryopteris
subattenuata ROSENST. in Fedde, Rep. 10 (1912)
332.— Thelypteris subattenuata (ROSENST.)
REED, Phytologia 17 (1968) 317.— Type: BAm-
LER 37, N.E. New Guinea, Logaueng, 300 m (S-
PA?; BM). — Fig. 19d.
Caudex short, erect; stipe to at least 50 cm long,
pale, basal scales c. 10 1.5mm. Lamina 150cm
or more long; pinnae well-spaced, the lowest with
very narrow bases bearing auricles 5 mm long on
both sides; 1-3 pairs of much-reduced pinnae,
similar auricled, present below normal ones. Lar-
gest pinnae 35cm long, 2.5-3.5 cm wide, lobed to
1-1.5mm from costa, lobes separated by wide
sinuses and almost at right angles to the costa, not
falcate, tips broadly rounded; costules to 6mm
apart; veins to 20 pairs, basal veins spreading at a
wide angle, their tips usually touching the sides of
a short sinus-membrane or sometimes uniting to
form a very short excurrent vein; lower surface
bearing a variable number of very small colourless
spherical glands, sometimes also very short aci-
cular hairs; upper surface of costae bearing pale
acicular hairs 0.6 mm long, few hairs on costules,
between veins a variable number of short acicular
and capitate hairs. Sori a little supramedial, not
impressed; indusia bearing very small glandular
548
hairs; sporangia sometimes with a small capitate
hair; spores pale, with highly prominent thick
ridges.
Distr. Malesia: Eastern New Guinea (6 collec-
tions).
Ecol. In somewhat exposed places near
streams in forest.
4. Amphineuron opulentum (KAULF.) HOLTTUM,
Blumea 19 (1971) 45; Blumea 23 (1977) 212. —
Aspidium opulentum KAULF. Enum. Fil.
Chamisso (1824) 238.— Thelypteris opulenta
(KAULF.) FOSBERG, Smiths. Contr. Bot. 8 (1972)
3, excl. syn. Aspidium terminans WALL. — Type:
CHAMISSO, Guam (LE).
Nephrodium impressum DESv. Mém. Soc.
Linn. Paris 6 (1827) 259.— Dryopteris impressa
(DESvV.) POSTH. Verh. K. Akad. Wet. Amst. 36, 5
(1937) 14; BACKER & POSTH. Varenfl. Java (1939)
57.— Thelypteris impressa (DESV.) REED, Phy-
tologia 17 (1968) 284. — Type: collector not cited,
Timor (P).
Aspidium extensum BL. En. Pl. Jav. (1828)
156. — Nephrodium extensum (BL.) MOORE, Ind.
Fil. (1858) 91; BEDpb. Handb. (1883) 269. — Dry-
opteris extensa (BL.) O. KTZE. Rev. Gen. Pl. 2
(1891) 812; v.A.v.R. Handb. (1908) 210. — Cyclo-
sorus extensus (BL.) CHING, Bull. Fan Mem.
Inst. Biol. Bot. 8 (1938) 182; HoLTTuM, Rev. FI.
Malaya 2 (1955) 264, f. 150; COPEL. Fern FI.
Philip. (1960) 368.— Thelypteris extensa (BL.)
Morton, Amer. Fern J. 49 (1959) 113. — Type:
no collector cited, Pulu Pinang (L).
Lastrea malaccensis PRESL, Epim. Bot. (1851)
35. — Type: CUMING 391, Malacca (PRC; K).
Nephrodium conioneuron FEE, Gen. Fil. (1852)
308. — Aspidium conioneuron (FEE) METT. Farn-
gatt. IV (1858) 102.— Type: GARDNER, Ceylon
(Isotype W).
Aspidium nephrodioides Hook. Spec. Fil. 4
(1862) 162, t. 235, non KLOTZSCH 1847. — Aspi-
dium hookeri BAK. Syn. Fil. (1867) 257, nom. nov.
non KLOTZSCH 1847 — Dryopteris orbicularis C.
Cure. Ind. Fil. (1905) 281, nom. nov. — Thelypteris
orbicularis (C. CHR.) REED, Phytologia 17 (1968)
299.— Type: SEEMANN, Indian Archipelago
(K). — Fig. 19b-c.
Caudex short-creeping; stipe to 70cm long,
rufescent, basal scales 10mm long, hardly 1mm
wide. Lamina to 80 cm long; pinnae to 25 pairs or
more; a pair of much-reduced basal pinnae some-
times present; apex of frond narrowly acuminate,
deeply lobed in its basal part and grading into the
upper pinnae. Largest pinnae commonly to 25 x
2.5cm, largest seen 403.5cm, lobed 3/5-3/4
towards costa; lobes slightly oblique, slightly fal-
cate; costules commonly 4mm apart, on large
sterile fronds to 6 mm; veins 8-10(-15) pairs, basal
pair both touching sinus-membrane or meeting
below it at a varying angle to produce an excur-
rent vein; lower surface of rachis, costae and
FLORA MALESIANA
(ser. II, vol. 1°
costules bearing very short acicular hairs with
scattered longer ones (to 0.5mm) on costules,
veins, sinus-membranes and margin, small pale
yellowish glands variously distributed along cos-
tules and veins, surface between veins usually
bearing some short erect acicular hairs and small
colourless to yellowish capitate hairs; upper sur-
face of costae covered with pale acicular hairs,
similar hairs scattered on costules and veins. Sori
confined to lobes of pinnae, supramedial, in slight
depressions; indusia thin, shrivelled when old,
bearing marginal yellow glandular hairs and
sometimes a few acicular hairs which are not
marginal; an elongate gland sometimes present on
stalks of sporangia.
Distr. East Africa; Seychelles; S. India and
Ceylon; Burma, Thailand; Malesia; N. Queensland;
islands of the Pacific to Tahiti; naturalized at
various places in tropical America.
Ecol. Few records; apparently adapted to
semi-exposed situations among rocks, especially
in areas with a dry season; in S.E. New Guinea
(dry season area) found in secondary swamp-
forest.
Notes. A fuller synonymy, and commentary on
it, is given in HOLTTUM 1977. METTENIUS pub-
lished the first good description (as Aspidium
conioneuron) with a note on the diversity of
venation. The species seems not to be common in
most parts of Malesia (doubtfully native in Java,
few specimens from Borneo and the Philippines).
Plants have long been cultivated in Singapore
(origin unrecorded) and occur spontaneously on
the edges of drains and elsewhere near the
Botanic Garden; these plants are tetraploid. The
center of distribution of the species is uncertain;
possibly southern India.
5. Amphineuron distinctum (COPEL.) HOLTTUM,
Blumea 23 (1977) 215.—Dryopteris distincta
CoPEL. Univ. Cal. Publ. Bot. 18 (1942) 220.—
Cyclosorus distinctus (COPEL.) COPEL. Gen. Fil.
(1947) 142; Philip. J. Sci. 78 (1951) 444, pl. 26. —
Thelypteris distincta (COPEL.) REED, Phytologia
17 (1968) 273. — Type: BAMLER W.11, N.E. New
Guinea, Wareo, 150m (UC).
Dryopteris longissima _ var.
ROSENST. Hedwigia 56 (1915)
BAMLER 132, same locality (B).
Caudex unknown; stipe incomplete, dark at
basé, distally reddish, minutely hairy, bearing thin
narrow scales throughout, basal ones 12 mm long;
abaxial surface of rachis also bearing similar
scales or their small wart-like bases. Size of
lamina not known; basal pinnae narrowed in their
basal 4cm, base 4mm wide. Largest pinnae 25 x
3cm, rather short-acuminate, lobed to 1.5-2 mm
from costa, lobes slightly oblique and slightly fal-
cate; costules 44.5 mm apart; veins to 18 pairs,
basal pair anastomosing to produce an excurrent
vein to the sinus or passing to sides of sinus-
novoguineensis
351.— Type:
1981]
THELYPTERIDACEAE (Holttum)
549
membrane; lower surface of rachis sparsely short-
hairy, of costae bearing copious erect hairs more
than 0.5mm long, fewer such hairs on costules
and veins, between veins copious slender erect
acicular hairs and small + yellowish capitate hairs;
upper surface of costae bearing thick acicular
hairs and small capitate ones, shorter hairs of both
kinds present between veins. Sori inframedial,
exindusiate; hairs on stalks of sporangia consis-
ting of 3 cells, distal ones club-shaped, orange;
spores dark with irregular thick ridges.
Distr. Malesia: Papua New Guinea; only
known from the type.
6. Amphineuron attenuatum (O. KTZE) HOLT-
TUM, Blumea 23 (1977) 215. — Aspidium attenua-
tum KUNZE ex METT. Farngatt. IV (1858) 96, non
Sw. 1801.— Nephrodium attenuatum BAK. Syn.
Fil. (1867) 263, non T. MOORE 1858. — Dryopteris
attenuata O. KTZE, Rev. Gen. PI. 2 (1891) 812,
nom. nov.; v.A.v.R. Handb. (1908) 184.— Dry-
opteris stenobasis C. CHR. Ind. Fil. (1905) 294,
nom. nov. superfl.— Thelypteris stenobasis (C.
CHR.) CHING, Bull. Fan Mem. Inst. Biol. Bot. 10
(1941) 254. — Mesoneuron attenuatum (O. KTZE)
CHING, Acta Phytotax. Sinica 8 (1963) 326.—
Thelypteris attenuata (O. KTZE) MORTON,
Contr. U.S. Nat. Herb. 38 (1967) 35.— Type:
CUMING 327, Samar (B; BRI, E, G, K, L, LE,
SING, US).
Dryopteris superficialis v.A.v.R. Bull. Jard. Bot.
Btzg II, 20 (1915) 12; Handb. Suppl. (1917) 155. —
Type: SA-ANAM 125, Obi I. (BO; L).
Dryopteris erubescens sensu CHRIST, Philip. J.
Sci. 2 (1907) Bot. 210. — Lastrea erubescens sensu
CoPEL. Fern FI. Philip. (1960) 329.
Cyclosorus alatellus sensu COPEL. Fern FI.
Philip. (1960) 341, excl. syn.
Caudex short, erect or suberect; stipe to at least
60 cm long, glabrous, rufescent, basal scales thin,
to c. 10x 1mm. Lamina to 80cm or more long;
lower pinnae much narrowed towards their bases,
the narrowed part several cm long, distinctly
lobed throughout; no reduced pinnae seen. Lar-
gest pinnae 30 x 2 cm, lobed to 1.5 mm from costa;
apex gradually attenuate to a cauda 3-5 cm long;
lobes slightly oblique and slightly falcate; cos-
tules 4-4.5 mm apart; veins to 18 pairs, basal pair
with upcurved tips passing to sides of sinus-
membrane or rarely joining just below the mem-
brane; lower surface of costae and costules bear-
ing scattered minute capitate hairs, between veins
slender short acicular hairs variably present on
different parts of the same frond, also short capi-
tate hairs and almost spherical red resinous
glands; upper surface of costa bearing very short
capitate hairs, also acicular hairs distally, some
capitate hairs present between veins. Sori some-
what inframedial, lower ones divergent; indusia
small with copious marginal spherical red resinous
glands; spores dark, rugose.
Distr. Malesia: Philippines (Luzon, Samar,
Mindanao), Moluccas (Obi Island), N. Celebes?
Ecol. ‘Edge of forest, petrophytic and ter-
restrial’’ on limestone and limestone-derived soils
(M. G. PRICE, on Samar), but probably not
confined to limestone.
Note. The hairs on the lower surface of pinnae
are inconstant. I am not sure of a clear distinction
between this species and A. ceramicum. A spe-
cimen collected by H. H. BARTLETT (n. 7693)
from Asahan, Sumatra has hairs on the lower
surface as in the type of A. attenuatum, but glands
are not well preserved. ENDERT 4165 from E.
Kalimantan is similar.
7. Amphineuron ceramicum (v.A.v.R.) HOLTTUM,
Blumea 23 (1977) 217.— Phegopteris ceramica
v.A.v.R. Bull. Dép. Agr. Ind. Néerl. 18 (1908) 15;
Handb. (1908) 506.— Type: TREUB s.n. Ceram
(BO).
Polypodium erubescens sensu HOOK. Spec. Fil.
4 (1862) 236, quoad pl. Molucc. tantum. — Poly-
podium erubescens var. amboinense BAK. Syn.
Fil. (1867) 306.— Type: collector not cited, Am-
boina, ex Herb. Webb (K; FI, P).
Dryopteris logavensis ROSENST. in Fedde Rep.
10 (1912) 232.—Lectotype (HOLTTUM 1977):
BAMLER L34 (S-PA; BM).
Phegopteris mamberamensis v.A.v.R. Bull.
Jard. Bot. Btzg II, 24 (1917) 3; Handb. Suppl.
(1917) 516.—Type: THOMSON 645, W. New
Guinea, Mamberamo River (BO; L).
Dryopteris moluccana C. CHR. Dansk Bot. Ark.
9 (1937) 64.—Type: FORBES 3273, Amboina
(BM; B).
Differing uncertainly from A. attenuatum as
follows: basal pinnae gradually contracted at their
bases to a narrow wing along the costa; no short
acicular hairs on lower surface of pinnae between
veins, no capitate hairs in that position on the
upper surface; glands between veins on the lower
surface varying from spherical to pyriform; glands
on indusia spherical, resinous.
Distr. Malesia: Moluccas (Amboina, Halma-
hera, Ceram), New Guinea.
Note. The type of Phegopteris ceramica
v.A.v.R. does not show glands on the lower sur-
face; it has pinnae to 22x 1.6cm. HOOKER’s
Amboina specimen which he wrongly included in
Polypodium erubescens is larger, with pinnae to
30x 3cm, the glands on the lower surface well
preserved, those near sinuses distinctly elongate.
A specimen of DE VRIESE from Ceram at Kew is
similar but sterile.
8. Amphineuron pseudostenobasis (COPEL.)
HOLTTUM, comb. nov.— Dryopteris pseudo-
stenobasis COPEL. J. Arn. Arb. 10 (1929) 176. —
Thelypteris pseudostenobasis (COPEL.) REED,
Phytologia 17 (1968) 306.— Type: BRAss 1000,
S.E. New Guinea, Vailala River (A; BRI, UC).
550
FLORA MALESIANA
[ser. II, vol. 1°
Cyclosorus alatellus sensu COPEL. Philip. J.
Sci. 78 (1951) 445, p.p. — Fig. 19f-h.
Differing from A. attenuatum as follows: no
acicular hairs on lower surface between veins, no
capitate hairs in this position on upper surface;
glands between veins on lower surface fewer,
apparently spherical, tending to collapse and form
irregular thin patches of resinous substance or to
disappear on drying of specimens; indusia very
small, none seen bearing glands (which are pos-
sibly present on living fronds).
Distr. Solomon Islands and Malesia: S.E. New
Guinea.
Ecol. In riverine forest “in large masses”
(BRASS).
Notes. The type has pinnae to 26x 1.8cm,
veins 11 pairs; pinnae on BRAITHWAITE’Ss spe-
cimens from the Solomon Islands are up to 40 x
4cm (n. 4505) with veins to 25 pairs. Two of
BRAITHWAITE’S specimens (4505 from Kolom-
bangara and 4014 from Guadalcanal) certainly
have a creeping caudex; his n. 4188 from San
Cristobal has the note “rhizome short, erect’’, but
otherwise is little different from the others.
Spreading resinous glands are only observable on
n. 4505.
9. Amphineuron paraphysophorum = (v.A.v.R.)
HOLTTUM, Blumea 23 (1977) 217, excl. syn. Dry-
opteris kiauensis C. CHR. — Dryopteris paraphy-
sophora v.A.v.R. Bull. Jard. Bot. Btzg III, 2 (1920)
143.—Type: C. J. BrRooKxs 250S, Sumatra,
Lebong Tandai (BO; BM).
Caudex short, erect or suberect; stipe to 80cm
long, basal part (up to 20cm in large fronds)
copiously scaly, scales thin, narrow, 7-8 mm long.
Lamina to 100 cm long (but plants of smaller size
may be fertile), texture rather thin; pinnae to 28
pairs; lower pinnae not reduced but 8-10 pairs
gradually narrowed towards their bases, lobes on
basal 2cm of lowest pinnae 1-2 mm long, then
gradually increasing to a maximum at 7cm from
base. Largest pinnae 25-35cm long, 2.0-3.5 cm
wide (widest on sterile fronds), apex acuminate
but not long-caudate; edges lobed to 1mm from
costa or more deeply; costules 3-4mm_ apart;
lobes hardly falcate, at more than 60° to costa:
veins to 20 pairs, basal acroscopic vein spreading
at a wide angle and abruptly upcurved near its tip
which touches the sinus-membrane or ends just
above it, basal basiscopic vein more often ending
above base of sinus; lower surface of rachis and
costae bearing very short capitate hairs only, on
costules, veins and between veins many pyriform
to almost cylindrical yellow glands; upper surface
of rachis bearing capitate hairs only, basal parts of
costae the same, acicular hairs up to 0.3 mm long
present on distal parts, very short ones also on
costules. Sori medial, lower ones divergent; in-
dusia small, lacerate, with elongate yellow glands
on the margin; sporangia sometimes with similar
glands on their stalks; spores with few large thick
protuberances of varied shape.
Distr. Malesia: S. Sumatra
(Sarawak).
Ecol. In Sarawak, in open places in forest at
90 m alt.
Notes. Part of the above description is based
on a plant from Gunong Mulu in Sarawak cul-
tivated at Kew. This shows the abundance of
glands of a distinctive shape on the lower surface
of pinnae and on indusia. The glands on a dried
specimen from the same locality are so shrivelled
as to be hardly distinguishable.
and Borneo
10. Amphineuron kiauense (C. CHR.) HOLTTUM,
comb. nov. — Dryopteris kiauensis C. CHR. Dansk
Bot. Ark. 9, 3 (1937) 64. — Type: ENDERT 4632
(wrongly cited as 4433), E. Kalimantan, Kutai,
Kiau, 700 m (BO; L, SING).
Description of type: stipe to 100cm_ long;
lamina to 100cm long, texture firm; pinnae (fer-
tile) to 18 x 1.5 cm, lobed to I-1.5 mm from costa;
basal 2cm of basal pinnae consisting of a narrow
wing above which is a gradual transition to full
width of the pinna; veins 8-10 pairs, basal pair
touching sides of sinus-membrane without anas-
tomosis; short capitate hairs present on lower
surface of costae, no glands detectable between
veins; sori inframedial; indusia small, lacerate.
Specimens from Sabah (KAKAWA & HOTTA
1291, distributed as Thelypteris erubescens) are
very similar but have many spherical to pyriform
glands on the lower surface, especially near the
sinuses, and spherical resinous reddish glands on
indusia. It seems probable that these represent the
same species as the type of A. kiauense; there is a
similar specimen (CLEMENS 29765) from Mt
Kinabalu, 1200 m.
Distr. Malesia: Borneo.
22. CHRISTELLA
LEVEILLE, FI. de Kouy-tchéou (1915) 472, emend. Ho_trum, Taxon 20
(1971) 533, Blumea 19 (1971) 43, Kew Bull. 31 (1976) 293. — Nephrodium
SCHOTT, Gen. Fil. (1834) t. 10 et sp. N. molle tantum, non RICHARD
1801. — Thelypteris subg. Cyclosoriopsis K. IWATS. Mem. Coll. Sci. Univ.
Kyoto B, 31 (1964) 28, p.p. — Cyclosorus sensu auctt. plur. p.p. — Fig. 1i, p,
20.
1981] THELYPTERIDACEAE (Holttum) 551
Caudex erect, suberect or creeping, in some species slender and wide-
creeping; scales almost always narrow with many superficial hairs. Lamina
in almost all species with I—5 (rarely to 10) pairs of lower pinnae gradually
decrescent, the lowest usually not less than 2cm long and in almost all
cases auricled on the acroscopic base, aerophores at the bases of pinnae not
swollen; largest pinnae shallowly to deeply lobed, bearing erect acicular
hairs (in some species minute) on all parts of the lower surface, also in
almost all species between veins on the upper surface (these hairs never
appressed), small capitate hairs sometimes also present; thick orange-red
glandular hairs (not erect) present in some species on costules and veins on
lower surface, sessile spherical glands lacking; veins free in a few species,
in most species at least the basal veins from adjacent costules anastomos-
ing, in some cases several pairs; sori indusiate (except in C. nana and C.
buwaldae); sporangia lacking hairs or glands distally (except in some
specimens of C. subpubescens) but bearing unicellular elongate glandular
hairs on their stalks (except in spp. 1-4); spores dark, variously tuberculate
or ridged, lacking thin wings.
Type species: Christella parasitica (L.) LEv.
Distr. About 50 spp.; throughout the warmer parts of the Old World; one species (C. hispidula) also
in the neotropics; in addition c. 15S—20 spp. in Africa and the neotropics the status of which still needs to be
established.
Ecol. In Malesia, almost all are ferns of open places; species confined to forest occur in the region
from N.E. India and Burma to S. China.
Cytol. Base chromosome number 36. C. dentata, C. parasitica and C. subpubescens are tetraploid, C.
hispidula diploid. Experimental hybridization of these species was undertaken at Leeds (see HOLTTUM
1976, p. 295); C. dentata and C. parasitica were shown to be allotetraploids with C. hispidula as one
parent of each. C. arida has been shown to be diploid in northern India but has not been experimentally
hybridized with tetraploids. It is evident that natural hybrids between some of the commoner species
have also developed, but it is difficult to assign a parentage to them.
Taxon. LEVEILLE characterized his genus as follows: pinnae lobed, with simple veins pinnately
arranged in the lobes, sori with reniform indusia, seated on the veins in a row on each side of the
costules. No type species was indicated. The characters apply to most species of Thelypteridaceae, and
LEVEILLE’s list included representatives of four genera as arranged in the present treatment, also two
which do not belong to the family. Three of his species were transferred to other genera by CHING;
several others do not strictly conform to LEVEILLE’s own generic definition. Three of his species are
closely allied and belong to a group recognized by me in my preliminary studies of the family, for which
I was seeking an appropriate generic name; I therefore chose a species from this group as type. In 1964
IWATSUKI had cited an allied species, C. dentata, as type of Thelypteris subg. Cyclosoriopsis, but his
definition of the subgenus would include many species which seem to me not nearly allied.
In the earlier literature the species of this genus were not clearly distinguished, so that there is much
confusion in the use of names. I have not attempted to assign meanings to all names cited; e.g. Aspidium
patens, A. nymphale and A. parasiticum in BLUME’s ‘Enumeratio’ of 1828 are not clearly distinguished
and I have not found the particular specimens to which he gave those names, which were copied from
earlier works by others, who had described them very briefly.
The single most distinctive character is the presence of an elongate unicellular gland on the stalks of
sporangia (shown in SCHOTT’s figure of 1834 but mentioned by no-one else); similar glands are also
present on the lower surface of pinnae in some species. This character is associated with others less
precisely definable, among them the rather thick protuberances or ridges of the perispore, a character
shared by Amphineuron. But the elongate gland on the sporangium-stalk is lacking in a group of four
species in New Guinea, for which I propose a new section as follows:
Leptochristella HOLTTUM, sect. nov. — Plantae parvae, calcicolae; pinnae 2-4.5 cm longae,
tenues, subtus omnino piliferae; venae liberae vel inferiores anastomosantes; sori indusiati
vel exindusiati; pedicelli sporangiorum glandulis destituti, interdum pilis acicularibus
praediti.
552 FLORA MALESIANA [ser. Il, volauty
SR PAAR OS BIR fA SD
SE AGEITES AAO EAA
Bees
Ny
QA
LAW Rs atten SESE MC RL AAV eee Ge MS
SCD
Fig. 20. Christella dentata (FORSSK.) BROWNSEY & JERMY. a. Two pinna-lobes, x 4.— C. hispidula
(DECNE) HOLTTum. b. Two pinna-lobes, x 4: c. upper surface of costa and base of costule, showing
acicular and capitate hairs, x 18.—C, papilio (HOPE) HoLTTUmM. d. A reduced basal pinna, X 1; e.
venation and sori, showing minute hairs on costa, x 6. — C. parasitica (L.) LEv. f. Pinna-lobe, showing
hairs and glands, x 6.—C. subpubescens (BL.) HOLTTUM. g. Venation and sori, x 4.—C. harveyi
(METT.) HOLTTUM. h. Two pinna-lobes, X 4; i. tip of a pinna-lobe showing sori, glands and hairs, x 16
(a PIGGOTT 1479, b-c YAPP 201, d~-e MOLESWORTH ALLEN 4949, f HOLTTUM s.n., g WALLICH 354,
h-i BRYCE 5).
1981] THELYPTERIDACEAE (Holttum) 553
Type species: Christella perpubescens (ALSTON) HOLTTUM.
The plants of this group of species, small in stature and consequently simplified in structure, are
difficult to characterize and need more study. They have the frond-form of Christella with lower pinnae
gradually and slightly reduced (shown most notably by C. perpubescens), abundant slender acicular
hairs (fewest in C. nana) and spores of the Christella type. | see no other genus to which they could be
considered allied.
In Africa also are species with free veins and lacking (so far as I have observed) glandular hairs on
the stalks of sporangia. For them I proposed the sectional name Pelazoneuron (J. S. Afr. Bot. 40, 1974,
144) but they appear to be related to a group of tropical American species in which glandular hairs of the
Christella type do occur on the stalks of sporangia of at least two species (A. R. Smith, Univ. Cal. Publ.
Bot. 29: 15, 66, 79) and apart from their free veins I see no clear distinction between these American
species and C. hispidula. A few species in the Western Pacific (including C. harveyi in the present work)
also have free veins; I suggest that they are derived from Malesian species with anastomosing veins.
KEY TO THE SPECIES
1. Limestone plants; pinnae not over 4.5cm long; no elongate glands on stalks of sporangia (sect.
Leptochristella).
2. Indusia distinctly present.
3. Pinnae to 27 pairs, S—7 lower ones gradually reduced .... . . . . I. C. perpubescens
3. Pinnae c. 6 pairs, lowest only slightly reduced . ....... 2... =. =. 2 C.minima
2. Indusia lacking or not clearly distinguishable.
4. Pinnae lobed almost to costae; veins all free nie: POOR eE Mee S33 MC Anana
we Pinnae lobed less deeply; veins near bases of pinnae anastomosing . . . . «4. C. buwaldae
. Plants not confined to limestone; pinnae commonly 10cm or more long; elongate glands present on
stalks of sporangia (sect. Christella).
5. Veins all free.
6. Red ellipsoid glands present on lower surface of pinnae sep Re De ee en OS Caiharveyi
6. Such glands lacking et AT a, PESTA Be aie i ee OE CS neckelil
5. At least the basal veins anastomosing.
7. At least 3 pairs of lower pinnae gradually reduced.
8. Sinus-membrane prominent on lower surface, 3 Sea of veins ending beside it; reduced pinnae
hardly-auricled’ 54s): JU eA: arida
8. Sinus-membrane not prominent, at most ‘3 pairs of veins s ending beside it: reduced pinnae clearly
auricled.
9. Caudex strictly erect.
10. Reduced pinnae c. 8 pairs; hairs on lower surface of costae c.0.1mmlong ._ 8. C. papilio
10. Reduced pinnae 3—4 pairs; some hairs on lower surface of costae 0.3 mm or more long.
11. Pinnae lobed less than half-way to costa; hairs on lower surface of costae all less than 0.5 mm
long : . . . 9, C. adenopelta
. Pinnae lobed n more than half- -way to costa: hairs « on lower surface of costae mostly more than
0.5Smmilong .. . Sate & 2 Fe seo e0h Cahispidula
9. Caudex short-creeping, or ‘at most suberect distally.
12. Pinnae lobed at least half-way to costa; only 1 pair of veins truly anastomosing; some hairs
0.3 mm long present on lower surface of costae st04 . . . . 11. C. dentata
12. Pinnae lobed 1/4-1/3 towards costa; at least 13 pairs of veins ; truly anastomosing; all hairs on
lower surface of costae c.0.1mmlong . . fo ES RI ee 123 | subpubescens
7. At most 3 pairs of lower pinnae gradually reduced.
13. Hairs on lower surface of costae all very short.
14. Pinnae lobed c. 1/4; minute capitate hairs present in addition to acicular hairs on lower surface
of pinnae : . . . . . . . 12. C. subpubescens
14. Pinnae lobed c. 2/5: no capitate hairs on lower surface Ate . . . . 13. C. subdentata
13. Hairs on lower surface of costae always conspicuous, to | mm long.
15. Caudex long-creeping.
16. Pinnae lobed more than 1/2; apices of pinna-lobes not apiculate.
17. Basal 1-15 pairs of veins anastomosing, 1 pair passing to sinus-membrane 14. C. timorensis
17. Basal 1 pair of veins ea Sea at most the next acroscopic vein meeting the sinus-
membrane : Loy Ree ree S='Caparasitica
16. Pinnae lobed less than 1/2; apices of pinna- lobes apiculate 2002. 216") acuminata
15. Caudex short-creeping or erect.
554
FLORA MALESIANA
[ser. II, vol. 1°
18. Pinnae lobed much more than 1/3; some hairs more than 0.5 mm long present on lower surface.
19. Caudex erect; 2 or more pairs of lower pinnae reduced
19. Caudex not erect; basal pinnae not reduced
10. C. hispidula
15. C. parasitica
18. Pinnae lobed little more than 1/2; hairs on lower surface all less than 0. 5 mm long
1. Christella perpubescens (ALSTON) HOLTTUM,
Kew Bull. 31 (1976) 304.— Dryopteris per-
pubescens ALSTON, J. Bot. 78 (1940) 227; Nova
Guinea n.s. 4 (1940) 111, pl. 8, f. 9, 10. — Thelyp-
teris perpubescens (ALSTON) REED, Phytologia
17 (1968) 303. — Type: CLEMENS 7902C, N.E. New
Guinea, Morobe Distr., Kalasa (B; BM).
Caudex short-creeping; stipe 3-5 cm long, den-
sely short-hairy throughout, basal scales 5mm
long, narrow, soft, covered with minute hairs.
Lamina 22cm long; texture thin; pinnae to 27
pairs, c. 7 pairs of lower ones gradually
decrescent, basal pinnae 3-5 mm long; apex of
frond also gradually attenuate. Largest pinnae of
type 2.8x0.9cm (sterile), 2.2x0.8cm (fertile),
sessile, short-acuminate, lobed to 0.5mm from
costa, basal pair of lobes sometimes separately
adnate to the costa, the acroscopic lobe not elon-
gate but sometimes with a sinuous margin; cos-
tules 2.5-3 mm apart; veins to 4 pairs, free, basal
acroscopic one sometimes touching side of sinus-
membrane, basiscopic one to margin above base
of sinus; lower surface of rachis and costae den-
sely covered with short hairs, scattered hairs more
than 0.5 mm long also present, costules, veins and
surface between veins bearing many short erect
acicular hairs, also pale capitate hairs; upper sur-
face throughout bearing slender erect acicular
hairs, also some capitate hairs. Sori medial or a
little supramedial; indusia firm, hairy; no hairs
seen on sporangium-stalks; spores dark, closely
and irregularly tuberculate.
Distr. Solomon Islands; Malesia: New Guinea.
Ecol. On limestone cliffs near sea-level (Wai-
geo Island and Solomons); in eastern New Guinea
reported to occur in rock crevices (type and
BRASS 23774) at 240-500 m.
Note. BRASS 23774, from Milne Bay District,
lacks capitate hairs, and also BRAITHWAITE 4841
from the Solomon Islands; the latter also differs in
longer stipes and less reduced basal pinnae. L. E.
CHEESMAN 1227 from Waigeo Island has pinnae
to 4.5 cm long, and a few capitate hairs.
2. Christella minima HOLTTUM, Kew Bull. 31
(1976) 304. — Type: JERMY 7875, New Ireland, on
limestone (BM).
Caudex short, apparently erect; stipe to 4.5 cm
long, slender, short-hairy, basal scales c. 3mm
long. Lamina to 9cm long, texture very thin;
pinnae 6 pairs; basal pinnae a little reduced, with
enlarged basal acroscopic lobes; apex of frond
gradually attenuate. Largest pinnae 1.8 x0.9cm,
lobed a little more than halfway to costa; apex
obtuse, lobes rounded, entire; costules 3mm
11. C. dentata
apart; veins free, to 4 pairs in the largest lobes,
basal acroscopic vein ending at base of sinus,
basiscopic one above base of sinus; lower surface
covered throughout with slender short hairs, some
longer ones present on rachis and costae; upper
surface covered with short slender hairs, some
longer ones also present on costules and veins.
Sori supramedial, on basal acroscopic veins, a few
on the second vein; indusia bearing many slender
hairs; spores very dark with irregular blister-like
protuberances.
Distr. Malesia: New Guinea (New Ireland),
only known from the type.
Note. This may be an immature state of C.
perpubescens.
3. Christella nana HOLTTUM, Kew Bull. 31 (1976)
304. — Type: MCKEE 1938, W. New Guinea, Biak
Island (L).
Caudex short-creeping; stipe 4-Scm_ long,
glabrous, basal scales 1.5mm_ long, narrow.
Lamina dimorphous, c. 7cm long, thin; pinnae 6
pairs, wider on basiscopic than on acroscopic side
of costa, basal pair a little reduced. Sterile pinnae
to 3.0 1.0cm, with stalks 1mm long, lobed al-
most to the costa near base; lobes oblique, entire,
with rounded tips, basal acroscopic lobe a little
enlarged; apex of pinnae obtuse; costules to 3 mm
apart; veins to 4 pairs, free, both basal ones
passing to margin above base of sinus; lower
surface of rachis covered with stiff erect hairs of
varying length, costae and costules bearing very
short somewhat antrorse hairs, rest of surface
glabrous; upper surface glabrous apart from the
costae. Fertile pinnae to 2.0 0.6cm; sinuses be-
tween lobes wider than in sterile pinnae; veins in
largest lobes 3 pairs, costules in upper pinnae once
forked; sori confined to the pinna-lobes, on upper
pinnae | or 2 in each lobe; no indusia; spores as in
S. minima.
Distr. Malesia:
known from type.
Ecol. Beneath overhanging coral rock just
above beach.
New Guinea (Biak I.), only
4. Christella buwaldae (HOLTTUM) HOLTTUM,
comb. nov. — Pronephrium buwaldae HOLTTUM,
Blumea 20 (1972) 115.— Type: BUWALDA 4979,
Aru Islands, P. Kobro6r (L; K).
Caudex short-creeping; stipe to 8cm _ long,
slender, covered with short pale erect hairs, basal
scales to 5mm long, narrow, with superficial short
hairs. Lamina of type 16cm long consisting of an
apical section 13cm long, widening downwards
and deeply lobed, the lobes veined as pinnae, with
1981]
THELYPTERIDACEAE (Holttum)
555
2-3 pairs of free pinnae; texture very thin, trans-
lucent; basal pinnae somewhat reduced. Largest
pinnae 2.8 1.0cm, sessile or with very short
stalks; base symmetrically broadly cuneate; apex
abruptly obtuse; edges lobed 1/3 towards costa,
lobes subtriangular; costules 3mm apart; veins
3-4 pairs, basal pair anastomosing except near
apex of pinna; lower surface of rachis, costae and
costules covered with erect hairs of varying
length, longest almost 1 mm long, surface between
veins bearing very slender erect hairs; upper sur-
face similarly hairy. Sori small, on pinnae in-
framedial, on lobes of apical lamina supramedial;
no indusia but acicular hairs present on the recep-
tacle (or a very small hairy indusium?); sporangia
lacking glands or setae; spores light brown.
Distr. Malesia: Moluccas (Buru & Aru Is.).
Ecol. On P. Kobro6r “in forest” (the island is
mainly limestone); on Buru “limestone, 1200 m”
(TOXOPEUS s.n. 4 Sept. 1921, BO).
Note. The Buru specimen is larger than the
type, with an apical lamina 8cm long and c. 6
pairs of free pinnae, largest pinnae 3.2 x 1.3.cm,
lobed more deeply than those of the type; in other
respects it agrees. An acicular hair was observed
on the stalk of a sporangium of the Buru speci-
men, and the pubescence of the frond is very like
that of C. perpubescens, not like that of any spe-
cies of Pronephrium, from which genus the spe-
cies is here transferred. But the spores of the type
need a careful re-examination; sporangia of the
Kew isotype are immature.
5. Christella harveyi (METT.) HOLTTUM, Kew
Bull. 31 (1976) 306; Allertonia 1 (1977) 219, f. 8
A-D. — Aspidium harveyi METT. in Kuhn, Lin-
naea 36 (1869) 115.— Dryopteris harveyi (METT.)
O. KTZE, Rev. Gen. Pl. 2 (1891) 812; C. CHR.
Bishop Mus. Bull. 177 (1943) 83.— Thelypteris
harveyi (METT.) PROCTOR ex K. IWATS. Amer.
Fern J. 53 (1963) 133. — Type: HARVEY, Fiji (B;
K).
Dryopteris euaensis COPEL. Univ. Cal. Publ.
Bot. 12 (1931) 391.— Type: PARKS, Tonga (UC;
BM, K).
Thelypteris novae-hiberniae HOLTTUM, Dansk
Bot. Ark. 25, 2 (1967) 50.— Type: KOIE 1848,
New Ireland (C).
Nephrodium patens sensu HOOK. Spec. Fil. 4
(1862) 95, p.p.— Dryopteris patens sensu v.A.v.R.
Handb. (1908) 189, p.p. — Fig. 20h-i.
Caudex long-creeping, 5-7 mm diameter when
dry; stipe 20-30cm long, hairy in groove, basal
scales to 8X 1.5mm. Lamina 50-70 cm long, tex-
ture thin; pinnae 20-25 pairs; lower pinnae
variously reduced, lowest sometimes only 3cm
long; reduced pinnae more widely spaced, some-
times with basal acroscopic lobe enlarged and
lobulate. Largest pinnae commonly 15-20 x 1.5-
2.2cm, rarely to 3cm wide; base not auricled;
apex caudate-acuminate; edges lobed to 1-1.5 mm
from costa; lobes oblique, slightly falcate, entire,
with rounded apices; costules 4-4.5mm_ apart;
veins 10-12 pairs, basal acroscopic one passing to
side of the short sinus-membrane, basiscopic one
to margin above base of sinus; lower surface of
rachis, costae and costules bearing copious short
hairs and a variable number of longer ones, some
thick orange glandular hairs present on costules
and veins, between veins a variable number of
slender short erect acicular hairs and glandular
hairs; upper surface of costae densely hairy, scat-
tered long hairs present on costules and veins,
between veins a variable number of short suberect
hairs and sometimes glandular hairs. Sori near
margin; indusia firm, glabrous or with a few hairs;
an orange glandular hair on the stalks of some
sporangia.
Distr. Solomon Islands, New Hebrides, Fiji,
Samoa, Wallis Isiand, Tonga, in Malesia: New
Guinea (Admiralty Islands, Bismarck
Archipelago).
Ecol. In open places in forest at 0-1000 m.
6. Christella peekelii (v.A.v.R.) HOLTTUM, Kew
Bull. 31 (1976) 306. — Dryopteris peekelii v.A.v.R.
Bull. Dép. Agr. Ind. Néerl. 18 (1908) 7; Handb.
(1908) 188.— Type: PEEKEL 44, “New Guinea”
(BO).
Caudex probably long-creeping; stipe probably
to 20cm long, copiously short-hairy, basal scales
not seen. Lamine 3040cm long; pinnae 18-22
pairs; basal pinnae slightly reduced, their basal
pair of lobes almost free, the acroscopic one
enlarged and deeply dentate. Largest pinnae 10
1.6cm (sterile), 9*1.3cm (fertile), acuminate,
lobed to less than 1mm from costa, lobes
entire and slightly falcate; costules of sterile
pinnae 4mm apart, of fertile pinnae 3 mm; veins
8-10 pairs, arranged as in C. harveyi; lower sur-
face bearing very short hairs throughout, no
glandular hairs present; upper surface bearing
scattered long hairs on costules and veins and
short suberect hairs between veins. Sori near
margins of lobes; indusia short-hairy; orange
glands present on stalks of sporangia.
Distr. Malesia: Papua New Guinea (New Bri-
tain).
Note. PEEKEL collected in the Bismarck
Archipelago; the type was probably found in New
Ireland. It is closely related to C. prolixa
(WILLD.) HoL_trumM (Aspidium obliquatum
METT.) of New Caledonia, which also lacks
glandular hairs on the lower surface of pinnae.
7. Christella arida (D. DON) HOLTTUM in Nayar
& Kaur, Comp. to Bedd. (1974) 206; Kew Bull. 31
(1976) 320; Allertonia 1 (1977) 172, 225, f. 9A. —
Aspidium aridum D. Don, Prodr. Fl. Nepal
(1825) 4. — Nephrodium aridum (D. Don) J. SM.
in Hook. J. Bot. 4 (1841) 188; BEDpD. Handb.
(1883) 272.—Dryopteris arida (D. DON) O.
556
FLORA MALESIANA
[ser. II, vol. 1°
KTZE, Rev. Gen. Pl. 2 (1891) 812; v.A.v.R.
Handb. (1908) 212; BACKER & POSTH. Varenfl.
Java (1939) 50.—Cyclosorus aridus (D. DON)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938)
194; HOLTTUM, Rev. FI. Malaya 2 (1955) 259, f.
146; COPEL. Fern FI. Philip. (1960) 362.—
Thelypteris arida (D. DON) MORTON, Amer.
Fern J. 49 (1959) 113. — Type: WALLICH, Nepal
(not found at BM; possibly at BR, see MORTON
1974, infra).
Aspidium obscurum BL. En. Pl. Jav. (1828)
150. — Nephrodium obscurum (BL.) T. MOORE,
Ind, Fil. (1858) 98; Racrs. FI. Btzg 1 (1898)
125. — Type: BLUME, Java (L).
Polypodium acuminatum ROXxB. Calc. J. Nat.
Hist. 4 (1844) 490, non HoutTtT. 1786; MORTON,
Contr. U.S. Nat. Herb. 38 (1974) 335.— Type:
ROXBURGH, “Ind. Or.” (BR; G).
Dryopteris arida var. ebeneorachis COPEL. in
Elmer, Leafl. Philip. Bot. 2 (1908) 390. — Type:
ELMER 1044, Negros (MICH; BO, G). — Fig. 1i.
Caudex long-creeping, 4-5 mm diameter; stipe
15-30cm long, glabrous except in groove, basal
scales 5 mm long, narrow. Lamina to 150 cm long;
pinnae to 30 pairs or more; lower 3-5 pairs
gradually or subabruptly decrescent and more
widely spaced, not or little auricled, lowest com-
monly 5S-10mm long; apex of frond almost pin-
na-like; texture very firm. Largest pinnae to 16x
1.8cm (width above the somewhat dilated base),
acuminate, lobed c. 1/4 towards costa. lobes with
a short stiff point; costules 3-4 mm apart; veins to
10 pairs, prominent beneath, 15 pairs, at a broad
angle to costule, anastomosing, next 3 pairs very
oblique and passing to the sinus-membrane which
is prominent on the lower surface; lower surface
of costae bearing rather sparse stiff erect hairs
0.2mm long and a few very narrow scales, cos-
tules and veins with scattered short acicular hairs
and thick yellow glandular hairs, short erect hairs
sometimes present between veins; upper surface
of costae covered with short antrorse hairs,
costules, veins and surface between veins almost
or quite glabrous. Sori medial, lower ones diver-
gent; indusia glabrous or with some glandular and
short acicular hairs.
Distr. N.W. India to southern China, Thailand
and Vietnam; throughout Malesia: N. Queens-
land, New Hebrides, New Caledonia, Fiji, Samoa.
Ecol. In open places in low country, often
among tall grasses, less commonly at higher al-
titudes (once in Malaya at 1800 m). Plants growing
in tall grass have a greater number of reduced
basal pinnae than those in more exposed places.
Note. Reports on the distribution of this spe-
cies are often erroneous because it has been con-
fused with Sphaerostephanos invisus and S. unitus
which have a similar habit (see Holttum in Al-
lertonia, 1977). LoyAL found a plant in North
India to be diploid; some specimens are possibly
hybrids with C. subpubescens.
8. Christella papilio (HOPE) HOLTTUM in Nayar
& Kaur, Comp. to Bedd. (1974) 208; Kew Bull. 31
(1976) 321.—Nephrodium papilio Hope, J.
Bombay Nat. Hist. Soc. 12 (1899) 625, t. 12.—
Cyclosorus papilio (HOPE) CHING, Bull. Fan
Mem. Inst. Biol. Bot. 8 (1938) 214; MOLESWORTH
ALLEN, Gard. Bull. Sing. 22 (1967) 180, 185. —
Thelypteris papilio (HOPE) K. IwaTs. Mem. Coll.
Sci. Univ. Kyoto B, 31 (1965) 175.— Lectotype
(HOLTTUM 1976): LEVIGNE s.n. 1880, N.E. India,
below Darjeeling 1200 m (K).
Nephrodium molle var. major BEDD. Handb.
Suppl. (1892) 76, p.p. — Fig. 20d-e.
Caudex massive, erect; stipe 10-30cm long,
minutely hairy, basal scales thin, c. 7X 1mm.
Lamina to 150cm or more long; c. 10 pairs of
lower pinnae gradually decrescent, broadly trian-
gular, strongly auricled on acroscopic base,
broadly truncate on basiscopic, margins above
base shallowly lobed, apex acuminate; lowest
pinnae I|-3cm long. Largest pinnae to 17 x2cm;
base subtruncate; apex acuminate with short
cauda; edges lobed 1/4-1/3 towards costa (most
deeply in fertile pinnae), lobes slightly falcate and
rounded at their tips; costules 4-4.5mm apart;
veins 7-9 pairs, 1) pairs anastomosing, 1-2 pairs
ending beside the sinus-membrane; lower surface
of all parts bearing very short erect hairs (less
than 0.1 mm long on rachis and bases of costae, a
little longer on distal parts of pinnae), some thick
orange glandular hairs sometimes present on veins
in pinna-lobes; upper surface of costae covered
with hairs 0.3-0.5 mm long, scattered shorter hairs
on costules, minute erect hairs on surface between
veins. Sori medial; indusia rather large, thin, with
short hairs as lamina.
Distr. Southern India & Ceylon; N.E. India,
Thailand; Malesia: to northern Malaya.
Ecol. In Malaya, in forest on sloping ground at
600-900 m.
Notes. The Indian specimens lack orange
glandular hairs on the lower surface, but such
glands are present on all specimens from Thailand
and Malaya. MANTON found a Ceylon plant to be
tetraploid, LOYAL found plants in N. India to be
diploid.
9. Christella adenopelta HOLTTUM, Kew Bull. 31
(1976) 322. — Type: W. A. SLEDGE 1698, Samoa
(K).
Near C. papilio, with the same erect caudex and
similar pubescence, but only 3-4 pairs of lower
pinnae gradually decrescent, the lowest 3.5 cm
long; largest pinna 12x 1.8cm; hairs between
veins on the lower surface a little longer than in C.
papilio and many short capitate hairs also present
with them, some scattered hairs up to 0.5 mm long
present distally on costae and costules; indusia
large, bearing glandular hairs like those on the
lower surface of veins in addition to many very
short acicular hairs.
1981]
THELYPTERIDACEAE (Holttum)
55 i]
Distr. Samoa and Malesia: S.E. New Guinea
(Brown River, near Port Moresby).
Ecol. In secondary swamp forest, low altitude.
10. Christella hispidula (DECNE) HOLTTUM, Kew
Bull. 31 (1976) 312.—Aspidium hispidulum
DECNE, Nouv. Ann. Mus. Hist. Nat. Paris 3
(1834) 346.— Dryopteris hispidula (DECNE) O.
KTZE, Rev. Gen. Pl. 2 (1891) 813; C. Cur. Ind.
Fil. (1905) 271, excl. syn. Nephrodium angusti-
follum PRESL & N. smithianum PRESL. —
Thelypteris hispidula (DECNE) REED, Phytologia
17 (1968) 283.— Type: GUICHENOT, Timor (P).
Dryopteris —_ contigua ROSENST. Meded.
Rijksherb. n. 31 (1917) 8; C. CHR. Gard. Bull. Str.
Settl 7 (1934) 244.—Cyclosorus contiguus
(ROSENST.) CHING, Bull. Fan Mem. Inst. Biol.
Bot. 10 (1941) 243; HoLTTum, Rev. FI. Mal. 2
(1955) 282, f. 163.— Thelypteris contigua
(ROSENST.) REED, Phytologia 17 (1968) 269. —
Type: TEUSCHER, Borneo (L).
Dryopteris hirtopilosa ROSENST. Meded.
Rijksherb. n. 31 (1917) 7; M. G. PRICE, Kalikasan
2 (1973) 112. — Thelypteris hirtopilosa
(ROSENST.) REED, Phytologia 17 (1968) 283. —
Type: MERRILL 7671, Luzon (L; K).
Dryopteris repandula v.A.v.R. Nova Guinea 14
(1924) 20.—Cyclosorus repandulus (v.A.v.R.)
CHING, Bull. Fan Mem. Inst. Biol. Bot. 10 (1941)
248; PANIGRAHI & MANTON, J. Linn. Soc. Bot.
55 (1958) 729-743.— Thelypteris repandula
(v.A.v.R.) REED, Phytologia 17 (1968) 308. —
Type: H. J. LAM 1058, W. New Guinea, Mam-
beramo River (BO; L).
Dryopteris parasitica var. falcatula CHRIST,
Philip. J. Sci. 2 (1907) Bot. 197.— Cyclosorus
falcatulus (CHRIST) COPEL. Fern FI. Philip.
(1960) 339.— Thelypteris falcatula (CHRIST)
REED, Phytologia 17 (1968) 276.—Type:
COPELAND 1677, Mindanao (MICH). — Fig.
20b-c.
Caudex erect; stipe 15-25cm long bearing
slender pale hairs, basal scales 7-10mm long.
Lamina 30-50 cm long, texture thin; pinnae 20-30
pairs, 2-4(—6) lower pairs gradually decrescent
with dentate acroscopic auricles, lowest com-
monly 3 cm long. Largest pinnae 8-12 (—15) x 1.2-
1.5(—2.0) cm; base truncate with basal acroscopic
lobe a little elongate; apex short-acuminate; edges
lobed 3/4 towards costa, lobes slightly oblique;
costules 3-4mm apart; veins 7-9 pairs, lowest
pair, at a wide angle to the costules, uniting to
form a short excurrent vein to the sinus, next pair
to margin; lower surface of rachis bearing rather
sparse slender pale hairs 1-1.5 mm long, shorter
hairs on costae, costules and veins, short slender
hairs on surface between veins, some short capi-
tate hairs usually present on costules and veins,
no thick glandular hairs; upper surface of rachis
and costae hairy as lower surface, scattered long
hairs present on costules and veins, short capitate
hairs sometimes present between veins. Sori
medial or a little supramedial; indusia thin, rather
small, bearing hairs of varying length.
Distr. Tropical America and wetter parts of
tropical Africa; Ceylon & S. India; Khasya Hills
southwards to Malaya; throughout Malesia:
Caroline Islands.
Ecol. Less common than C. parasitica and in
more sheltered places, low altitudes to 1500 m.
Notes. Plants from Florida, Ascension Island,
Ghana, Nigeria, Ceylon and Sarawak have proved
to be diploid. For cytotaxonomic studies involving
this species, see PANIGRAHI & MANTON 1955
(under Cyclosorus repandulus).
Only synonyms based on Malesian specimens
are cited above; see HOLTTUM 1976 for a fuller
synonymy. The latest information on plants in
tropical America is by A. R. SMITH, Univ. Cal.
Publ. Bot. 59 (1971) 64, under the name Thelyp-
teris quadrangularis (FEE) SCHELPE. Christella
siamensis (TAGAWA & K. IWATS.) HOLTTUM, in
Thailand and further north, differs only in less
deeply lobed pinnae, and a distinction from T.
hispidula seems doubtful; study of plants in cul-
tivation is desirable.
11. Christella dentata (FORSSK.) BROWNSEY &
JERMY, Brit. Fern Gaz. 10 (1973) 338; HOLTTUM,
Kew Bull. 31 (1976) 314. — Polypodium dentatum
Forssk. Fl. Aegypt. Arab. (1773) 185.— Dry-
opteris dentata (FORSSK.) C. CHR. Vid. Selsk.
Skr. VIII, 6 (1920) 24; BACKER & POSTH. Varenfi.
Java (1939) 58, excl. syn. Aspidium parasiticum &
A. patens.— Thelypteris dentata (FORSSK.) E.
St. JOHN, Amer. Fern J. 26 (1936) 44. — Cyclo-
sorus dentatus (FORSSK.) CHING, Bull. Fan
Mem. Inst. Biol. Bot. 8 (1938) 206; COPEL. Fern
Fl. Philip. (1960) 337, p.p.— Type: FORSSKAL,
Arabia (C).
Polypodium nymphale G. Forst. FI. Ins. Austr.
Prodr. (1786) 81.—Aspidium nymphale (G.
FORST.) SCHKUHR, Kr. Gew. 1 (1806) 36, t. 34;
BLUME, En. Pl. Jav. (1828) 157.— Thelypteris
nymphalis (G. ForsT.) REED, Phytologia 17
(1968) 297.—C. nymphalis (G. FORST.) PICHI
SERMOLLI, Webbia 31 (1977) 252.— Type: G.
FORSTER, New Zealand (BM).
Polypodium molle JAca. Collect. Bot. 3 (1789)
188, non SCHREB. 1771. — Aspidium molle Sw. in
Schrad. J. Bot. 1800, 2 (1801) 34, nom. nov. —
Nephrodium molle (Sw.) R. BR. Prodr. Fl. N.
Holl. (1810) 149; Racip. FI. Btzg 1 (1898) 188,
p.p.?— Dryopteris mollis (Sw.) HTlERON. Hed-
wigia 46 (1907) 348; v.A.v.R. Handb. Suppl. (1917)
183, p.p.? — Type: Cult. Vienna (W).
Dryopteris mindanaensis CHRIST, Philip. J. Sci.
2 (1907) Bot. 194.—Cyclosorus mindanaensis
(CHRIST) COPEL. Gen. Fil. (1947) 143; Fern FI.
Philip. (1960) 363.— Thelypteris mindanaensis
(CHRIST) REED, Phytologia 17 (1968) 293.—
Type: COPELAND 607, Mindanao (MICH).
558
FLORA MALESIANA
[ser. II, vol. 1°
Dryopteris submollis v.A.v.R. Bull. Jard. Bot.
Btzg III, 2 (1920) 152.—Type: LORZING 6040,
Sumatra, Karo Plateau (BO; L).
Cyclosorus subpubescens sensu. HOLTTUM,
Rey. FI. Malaya 2 (1955) 273, f. 157. — Fig. 1p, 20a.
Caudex short-creeping; stipe very variable, to
50cm long, short-hairy, basal scales c. 8 mm long.
Lamina to 90cm long; pinnae 15-25 pairs; lower
2-4 pairs of pinnae gradually decrescent, lowest
commonly 4-5 cm long, sometimes less, all stron-
gly auricled on the acroscopic base, the auricles
lobed with forked veins in the lobes. Largest
pinnae commonly 8-10 x 1.5-1.8 cm (largest seen
23x2.2cm, type of Dryopteris mindanaensis);
apex acuminate; edges lobed 1/2-2/3 towards
costa, lobes slightly oblique with rounded tips;
costules commonly 4mm apart; veins 8-9 pairs,
basal ones anastomosing with excurrent vein to
the sinus, next acroscopic vein (basiscopic some-
times also) passing to side of the short sinus-
membrane; lower surface of rachis bearing slen-
der pale hairs 0.2-0.4 mm long, hairs on costae and
costules mostly 0.2mm long with some longer
ones, rarely to 0.5mm long, distally on pinnae
short hairs present on surface between veins;
hairs on upper surface of costae to 0.5mm or
more long, scattered similar hairs on costules and
veins, very short hairs between veins. Sori medial
apart from lowest ones which sometimes touch
those on veins from neighbouring costules; in-
dusia thin, copiously short-hairy.
Distr. Throughout tropics and subtropics of
the Old World, since 1930 adventive in various
places in the Americas.
Ecol. In open or lightly shaded places, at 0-
1500 m.
Notes. For a fuller synonymy, see HOLTTUM
1976. This is a very variable species which has
greatly multiplied with the clearing of forest in the
past 100 years. Plants examined from various
sources have all been tetraploid; some experi-
mental hybridization has been effected with C.
hispidula (q.v.) and C. parasitica. It is probable
that natural hybrids also occur; these are difficult
to discriminate. Earlier authors did not distinguish
between this species and its near allies, so that
statements in literature are unreliable. Much more
local study is needed.
12. Christella subpubescens (BL.) HoOLtTTuM,
Webbia 30 (1976) 193; Kew Bull. 31 (1976) 323. —
Aspidium subpubescens BL. En. Pl. Jav. (1828)
149. — Dryopteris subpubescens (BL.) C. CHR.
Gard. Bull. Str. Settl. 4 (1929) 390; BACKER &
PosTH. Varenfl. Java (1939) 65.— Thelypteris
subpubescens (BL.) K. IwWATS. Mem. Coll. Sci.
Univ. Kyoto B, 31 (1965) 173, excl. syn. Aspidium
jaculosum CHRIST. — Type: BLUME, Java, river
bank at Tanjong Krukot, Batavia (L, n. 910, 327-
113, large sheet collection).
Aspidium subpubescens BL. var. B. BL. En. PI.
Jav. (1828) 149.—Type: BLUME, Java, Noesa
Kambangan (L, n. 922, 220-249, large sheet collec-
tion).
Aspidium subpubescens BL. var. C BL. L.c.—
Type: BLUME, W. Java, Kolleket (L).
Aspidium molle var. latipinna BENTH. FI.
Hongkong. (1861) 455.— Nephrodium latipinna
Hook. Syn. Fil. (1867) 292.— Dryopteris
latipinna (BENTH.) O. KTZE, Rev. Gen. Pl. 2
(1891) 813; v.A.v.R. Handb. (1908) 217. — Cyclo-
sorus latipinna (BENTH.) TARD. Notul. Syst. 7
(1938) 73; HOLTTUM, Rev. FI. Malaya 2 (1955)
276, f. 159. — Thelypteris latipinna (BENTH.) K.
IWATS. Acta Phytotax. Geobot. 21 (1865) 166;
MorRTON, Contr. U.S. Nat. Herb. 38 (1974) 361. —
Lectotype (MORTON 1974): HANCE 135, Hong
Kong (K).
Dryopteris sumatrana v.A.v.R. Handb. (1908)
227. — Nephrodium molle var. major BEDD.
Handb. Suppl. (1892) 76, quoad pl. Sumatr.
tantum. — Dryopteris subpubescens var. major C.
CHR. Gard. Bull. Str. Settl. 4 (1929) 390.—
Cyclosorus sumatranus (v.A.v.R.) CHING, Bull.
Fan Mem. Inst. Biol. Bot. 10 (1941) 249; HoLr-
TUM, Rev. Fl. Malaya 2 (1955) 275, f. 158.—
Thelypteris sumatrana (v.A.v.R.) TAGAWA & K.
IwatTs. Acta Phytotax. Geobot. 22 (1967) 101. —
Lectotype (HOLTTUM 1955): C. MILLER s5.n.
1778, Sumatra (BM).
Dryopteris — pseudoamboinensis ROSENST.
Meded. Rijksherb. n. 31 (1917) 7.— Thelypteris
pseudoamboinensis (ROSENST.) PANIGRAHI,
Phytologia 30 (1975) 410, pl. III]. — Lectotype
(PANIGRAHI 1975): KORTHALS 270, Sumatra (L).
Dryopteris acuminata ROSENST. Meded.
Rijksherb. n. 31 (1917) 7, non Watts 1916. —
Thelypteris blumei PANIGRAHI, Phytologia 30
(1975) 409, nom. nov.; op. cit. 31 (1975) 369. —
Type: ZOLLINGER 735, Java, Ufer des Tjidurians
(L, n. 908, 333-35).
Aspidium amboinense sensu BL. En. Pl. Jav.
(1828) 148; KUNZE, Bot. Zeit. 6 (1848) 261; METT.
Farngatt. IV (1858) 105. — Nephrodium molle var.
amboinense BEDD. Handb. (1883) 278, p.p. — Fig.
20g.
Caudex short-creeping or suberect; stipe short,
minutely hairy. Lamina varying much in size;
several pairs of lower pinnae _ gradually
decrescent, their acroscopic auricles rather small,
entire or slightly crenate; largest pinnae lobed less
than 1/2 towards costa (in small plants 1/4); in
small plants at least 13 pairs of veins anastomosing
and the next pair ending beside the sinus-mem-
brane, in large plants 2-23 pairs of veins anas-
tomosing to form a zig-zag excurrent vein and +1
pair ending beside the sinus-membrane; lower
surface of rachis, costae and costules bearing
minute hairs (0.1 mm long or less), in some spe-
cimens also orange glandular hairs on costules and
veins, surfaces between veins minutely hairy,
usually with some very short capitate hairs; upper
1981]
surface of costae bearing hairs c. 0.3mm long,
scattered longer hairs sometimes present on cos-
tules and veins, minute suberect hairs usually
present between veins. Sori medial; indusia rather
large, thin, glabrous or minutely hairy; in a few
cases, capitate hairs present on sporangia.
Distr. N.E. India to S.W. China; Burma,
Thailand, Vietnam; throughout Malesia; N.
Queensland, New Hebrides, Fiji.
Ecol. On stream-banks in moderate shade in
forest, also in lightly shaded places elsewhere, in
lowlands.
Notes. It would be possible to make a key
distinguishing the types of the basionyms above
cited, but I find it impossible to fit all other spe-
cimens into such a key. More field study, and
experimental study of cultivated plants, are needed.
The types of Aspidium subpubescens BL. var.
C and Nephrodium iatipinna HOoOK., also the
specimens of BLUME and ZOLLINGER on which
the misinterpretations of the name Aspidium am-
boinense WILLD. were based, were small fertile
plants. I have seen such plants on the banks of
small streams in the forest in Pahang, where the
streams are subject to periodic flooding with
swiftly-flowing water after a sudden storm. In
such places, plants are washed away before they
attain their full size; a plant from the Tahan river
in Malaya, grown at Kew, is much larger than any
on the river banks, and is very like BLUME’s var.
B, though less rigid in texture. Plants found grow-
ing spontaneously in and near the Botanic Garden
in Singapore (also others growing similarly at
Bogor) are very like the type of A. subpubescens
BL. and differ from the fully-grown plants of
stream-bank origin in having somewhat broader
pinnae. I have no evidence that the Singapore and
Bogor plants are fertile from an early age, and
think that they are probably genetically different
from stream-bank plants, but I cannot point to
clear distinctions.
Thick orange glandular hairs are present on the
lower surface of pinnae of most of the stream-
bank plants from Malaya, and of the Java plants
named amboinense by BLUME, KUNZE, MET-
TENIUS and HOOKER. Such glands are absent
from the type of Nephrodium latipinna HooK.
and other specimens from Hong Kong, also from
the type of A. subpubescens BL. and similar spe-
cimens from Singapore and Bogor. But other
specimens very similar to typical A. subpubescens
have a few such glandular hairs, while on the
other hand some stream-bank plants from north-
ern Malaya are quite glandless like the type of N.
latipinna from Hong Kong.
Some stream-bank plants from Malaya, also
from Luzon, have small capitate hairs on some of
their sporangia; I have not observed this character
in any other specimens of Christella. The capitate
hairs appear to be just like those on the lower
surface of pinnae between veins.
THELYPTERIDACEAE (Holttum)
559
13. Christella subdentata HOLTTUM, Kew Bull. 31
(1976) 335. — Type: VILLAFLORES 57, Mindoro,
Lubang Island (MICH).
Caudex short, erect or suberect; stipe 25cm
long, minutely hairy, basal scales c. 10 1mm.
Lamina 50cm long; pinnae 15 pairs; lowest 1-2
pairs of pinnae somewhat reduced (basal pinnae
5cm long), not auricled on the acroscopic side.
Largest pinnae 9.5 * 1.5cm; base truncate; apex
acuminate with cauda 2cm long; edges lobed 2/5
towards costa; costules 3.5mm apart; veins to 8
pairs, lowest pair anastomosing, second acros-
copic veins passing to side of sinus-membrane;
lower surface of rachis, costae and costules bear-
ing hairs 0.1mm long, similar hairs present be-
tween veins; upper surface of rachis covered with
hairs 0.5 mm long, shortet hairs present on costae,
sparse minute hairs between veins. Sori medial:
indusia short-hairy.
Distr. Malesia: Philippines (Mindoro), only
known from the type.
Note. This may be a hybrid.
14. Christella timorensis HOLTTUM, Kew Bull. 31
(1976) 316. — Type: BLOEMBERGEN 3364, Timor,
Mt Timan (L; BO, K).
Caudex long-creeping, 5mm diameter, bearing
fronds 2-3 cm apart; stipe short-hairy, basal scales
c. 6X 1.5mm, rigid. Lamina of type incomplete,
probably 70cm or more long with pinnae more
than 20 pairs; 4-5 pairs of lower pinnae
decrescent, with enlarged entire basal acroscopic
lobes (to 2.0cm long), lowest pinna seen 7.5cm
long. Largest pinnae 15cm long, 2.2cm wide
above base; apex narrowly acuminate but not
caudate; edges lobed 3/5—2/3 towards costa; lobes
oblique, a little falcate, broadly pointed; costules
4.5-5 mm apart; veins to 12 pairs, basal 1-15 pairs
anastomosing, next acroscopic vein, or next pair,
passing to side of sinus-membrane; lower surface
of rachis bearing rather sparse hairs 0.7-1.0mm
long, costal hairs erect, to 0.7 mm, shorter hairs on
costules and veins, sometimes also a few thick
glandular hairs, slender erect hairs present be-
tween veins; hairs on upper surface of costae to
0.5mm long, similar hairs scattered on costules
and veins, no hairs between veins. Sori suprame-
dial; indusia with short stiff hairs.
Distr. Malesia: Lesser Sunda Is. (Timor), only
known from type.
Ecol. In grassland along stream at 1300 m.
15. Christella parasitica (L.) Lev. FI. Kouy-
tchéou (1915) 475: HoLTTUM, Kew Bull. 31 (1976)
309. — Polypodium parasiticum L. Sp. Pl. (1753)
1090. — Dryopteris parasitica (L.) O. KTZE, Rev.
Gen. Pl. 2 (1891) 811; @ Cur Ark: Bots 9)! 11
(1910) 26, f. 4.—Cyclosorus parasiticus (L.)
FARW. Amer. Midl. Nat. 12 (1929) 259; HoLt-
TUM, Rev. Fl. Malaya 2 (1955) 281, f. 162.—
Thelypteris parasitica (L.) K. Iwats. Mem. Coll.
560
Sci. Univ. Kyoto B, 31 (1965) 172.—Type:
OSBECK, Canton (S-PA in Herb. Swartz).
Aspidium procurrens METT. Ann. Mus. Bot.
Lugd.-Bat. 1 (1864) 231.— Dryopteris procurrens
(METT.) O. KTZE, Rev. Gen. Pl. 2 (1891) 813;
v.A.v.R. Handb. (1908) 211, 819, p.p.— Cyclo-
sorus procurrens (METT.) COPEL. Fern FI. Philip.
(1960) 340, nomen tantum. — Thelypteris procurrens
(METT.) REED, Phytologia 17 (1968) 306. — Type:
ZIPPELIUS, Java (L, n. 908, 335-152).
Nephrodium didymosorum PARISH ex BEDD.
Ferns Br. India (1866) t. 200.—N. molle var.
didymosorum BEDD. Handb. (1883) 279. — Dry-
opteris didymosora (BEDD.) C. CHR. Ind. Fil.
(1905) 262; v.A.v.R. Handb. (1908) 225.— Type:
PARISH, Burma, Moulmein (not found in Herb.
Bedd. at K).
Nephrodium tectum BEDD. Handb. Suppl.
(1892) 79, excl. KING 8757.— Lectotype (HOLT-
TUM 1976): WALLICH 394, Singapore (K-W).
Dryopteris albociliata COPEL. J. Arn. Arb. 10
(1929) 177.— Thelypteris albociliata (COPEL.)
REED, Phytologia 17 (1968) 259. — Type: BRASS
566, E. New Guinea, Basiatibu (GH; UC).
Cyclosorus benguetensis COPEL. Philip. J. Sci.
81 (1952) 28; Fern Fl. Philip. (1960) 341.—
Thelypteris benguetensis (COPEL.) REED, Phy-
tologia 17 (1968) 263.—Type: MERRILL 7679,
Luzon, Benguet Subprov. (MICH) — Fig. 20f.
Caudex short- to long-creeping; stipe to 40cm
long, copiously hairy. Lamina to 40 cm long, tex-
ture thin; pinnae c. 20 pairs, closely placed; basal
pinnae (as dried for herbaria) deflexed, not widely
spaced, not or little reduced, auricled at the
acroscopic base, auricle curved towards the
rachis, sometimes dentate. Largest pinnae com-
monly 162cm, short-acuminate, lobed 2/3-3/4
towards costa, lobes slightly oblique; costules c.
4mm apart; veins 8-10 pairs, basal pair anas-
tomosing, next pair passing to margin above base
of sinus or sometimes the second acroscopic vein
touching the sinus-membrane; lower surface
typically covered with slender erect hairs to 1 mm
long, thick orange glandular hairs usually present
on veins in the lobes; hairs on upper surface of
costae thicker than on lower, similar hairs scat-
tered on costules and veins, very short hairs be-
tween veins. Sori medial, lower ones divergent;
indusia hairy.
Distr. Wetter parts of tropics and subtropics
of Asia; throughout Malesia; Queensland; in the
Pacific to Tahiti and Hawaii; E. Africa (Uganda,
Kenya); St Helena (introduced ?).
Ecol. In open places, mainly at low altitudes
but also to 1500 m; apparently nowhere abundant.
Notes. The type was discovered by CHRIS-
TENSEN (1910); previously the name was used
confusedly. In Kwangtung (source of the type)
almost all plants have a rather short-creeping
caudex and many glandular hairs on veins, but
some have a longer rhizomatous caudex and few
FLORA MALESIANA
[ser. II, vol. 1°
or no glands. The type of Aspidium procurrens
METT. lacks glands. BEDDOME at first used the
name procurrens for a mixture of species which
had in common a creeping rhizome; v.A.v.R. also
used the name confusedly. Under Cyclosorus
procurrens COPELAND described a fern with
narrow pinnae lobed half-way to the costa; I have
not seen specimens. Some specimens have the
frond-form of this species but short hairs; some
agree in hairs and glands but have some reduced
basal pinnae (type of Cyclosorus benguetensis);
some such specimens are probably hybrids.
MANTON found that C. parasitica in Ceylon is
tetraploid and her co-workers produced hybrids
with the diploid C. hispidula and with other
tetraploids. In breeding, the absence of glandular
hairs is recessive to their presence.
16. Christella acuminata (HoutTtT.) Lev. FI.
Kouy-tchéou (1915) 476; HOLTTUM, Kew Bull. 31
(1976) 333.— Polypodium acuminatum HourTt.
Nat. Hist. 14 (1783) 181, pl. 99, f. 2. — Thelypteris
acuminata (HOUTT.) MORTON, Amer. Fern J. 48
(1958) 139; K. Iwatrs. Mem. Coll. Sci. Univ.
Kyoto B, 31 (1965) 186.— Type: THUNBERG,
Japan (UPS).
Polypodium unitum THUNB. FI. Jap. (1784) 336,
non LINN.— Polypodium sophoroides THUNB.
Tr. Linn. Soc. 2 (1794) 341, nom. nov. — Neph-
rodium sophoroides (THUNB.) DESV. Mém. Soc.
Linn. Paris 6 (1827) 256; BAK. Syn. Fil. (1867)
289. — Type: as for P. acuminatum HOUTT.
Caudex long-creeping, slender; stipe 25cm or
more long, sparsely hairy. Lamina to 60cm long,
texture firm; lower pinnae more widely spaced
than the rest but not or little reduced. Largest
pinnae to 9X 1.5cm (fertile), 15 x 2cm (sterile),
lobed a little less than half-way to costa; lobes
acute, basal acroscopic lobe enlarged, + curved
towards the rachis, with some forked veins; cos-
tules 3.5—4.5 mm apart; veins commonly 7 pairs (to
10), pale and prominent on both surfaces, basal
pair anastomosing, next pair ending beside sinus-
membrane; lower surface of costae and costules
bearing a variable number of short erect hairs and
a few longer ones, sparse short hairs present on
and between veins, glandular hairs lacking; upper
surface of costae bearing hairs to 0.5mm long,
similar hairs often scattered on costules and veins,
a variable number of very short hairs between
veins. Sori medial or a little supramedial; indusia
large, copiously short-hairy.
Distr. Southern China and southern Japan, in
Malesia: Philippines (Babuyan Is. and N. Luzon).
Notes. For a fuller synonymy, see CHING
1938 and IWATSUKI 1965. The specific epithet
sophoroides is used in most of the older lit-
erature. In general aspect this species is similar to
C. arida but lacks reduced basal pinnae and
glandular hairs; the larger pinnae of C. acuminata
are also rather strongly auricled.
ADDENDA, CORRIGENDA ET EMENDANDA
As has been done in Series I, Flowering Plants, it seems useful to complete the volume with worthwhile
additions and corrections.
Page numbers are provided with either a or b denoting the left and right columns respectively.
13b
14a
20a
20b
22a
22b
34a
Gleicheniaceae
Gleichenia volubilis var. peninsularis.
Replace by the following:
7a. Gleichenia gigantea WALL. ex HOOK.
Spec. Fil. 2 (1844) 5, t. 3A; BEDD. Ferns
Brit. India (1865) t. 30.— Diplopterygium
giganteum (HOOK.) NAKAI, Bull. Nat. Sci.
Mus. Tokyo n. 29 (1950) 50.— Type:
WALLICH 157, Nepal (K).
G. glauca auct. non (THUNB.) HOOK.;
BEDD. Handb. (1883) 2, quoad pl. Ind.
Differs from G. volubilis JUNGH. of Java
and Sumatra as follows: lower surface of
lamina distinctly glaucous; stipular leaflets
branched and more deeply lobed; scales on
lower surface of rachis and costae rusty
brown like the felt of stellate hairs.
Distr. N.E. India at c. 1500 m; Vietnam;
N. Thailand; in Malesia still only known
from the collection cited from the Malay
Peninsula.
Note. Apart from the copious scales
and hairs on rachis and costae this differs
little from G. longissima BL. Further field
study on the higher mountains of Malaya is
desirable.
Gleichenia clemensiae
TUM.
Add the following: Stipular leaflets lacking
(thus resembling G. bullata, p. 13).
Gleichenia reflexipinnula C. CHR.
Add the following: BRASS 30382 from Mt
Wilhelm, 2900 m, has ultimate branches to
42 by 4.5cm with costules 5 mm apart and
veins distinctly prominent.
Gleichenia truncata (WILLD.) SPR. var.
truncata.
Add the following synonym: G. bifurcata
BL. En. Pl. Jav. (1828) 250.
Gleichenia milnei BAKER.
Add the following synonym: Sticherus
kajewskii St JOHN, Occ. Pap. Bishop Mus.
17 (1942) 81.
Add to Distr.: Mindanao (OLSEN 986).
(COPEL.) HOLT-
Insert the following after Gleichenia
erecta:
25a. Gleichenia hooglandii HOLTTUM,
Blumea 14 (1967) 327. — Type:
HOOGLAND & SCHODDE 7692, W. High-
lands of Papua New Guinea (CANB).
Intermediate between G. erecta and G.
bolanica (p. 24). Differs from G. erecta as
follows: largest lamina-segments on the
ultimate branches 8-15 mm long, costules
3-4 mm apart.
Dicranopteris linearis (BURM. f.) UNDERW.
34a
35a
37
39
40
44b
var. ferruginea (BL.) HOLTTUM.
Correction for synonymy: The com-
bination D. ferruginea was first published
by HosoKAWaA, Trans. Nat. Hist. Soc.
Formosa 25 (1935) 435. Additional
synonym: D. tomentosa COPEL. Univ.
Cal. Publ. Bot. 18 (1942) 217.—Type:
CLEMENS 41228, N.E. New Guinea (UC).
Dicranopteris linearis var. subferruginea
(HIERON.) NAKAI.
Add the following synonym: Gleichenia
caudata COPEL. Bishop Mus. Bull. 59
(1929) 9.— Type: GILLESPIE 4389, Fiji
(UC).
Dicranopteris linearis var. subpectinata
(CHRIST) HOLTTUM.
Add to Distr.: Palawan.
Additional varieties of Dicranopteris
linearis which need study:
var. bidentata v.A.v.R. Bull. Jard. Bot.
Btzg III, 5 (1922) 204, Karimon I.
var. crassifrons v.A.v.R. Ibid., Ternate.
var. stipulosa CHRIST, Bull. Herb. Boiss.
II, 6 (1906) 1010, Luzon.
Schizaeaceae
Schizaeaceae: Fossils.
Add the following: JENNINGS & EGGERT,
Amer. J. Bot. 59 (1972) 66 state that the
anatomy of Senftenbergia is like that of
Ankyropteris and conclude that Senften-
bergia is not related to Schizaeaceae. But
the sporangia and spores of Senftenbergia
are very similar to those of some living
Schizaeaceae; the latter are certainly much
reduced and specialized vegetatively. The
subject needs a much more comprehensive
study.
Cytology of Lygodium.
Add the following: L. longifolium from
Singapore: n=58, tetraploid; L. salici-
folium from Perak: n= 28?, diploid. (In-
formation from I. MANTON, in litt.).
Schizaea SM.
Add the generic synonym: Ripidium
BERNH. in Schrader, J. Bot. 1800, pt 2
(1801) 127.
Schizaea malaccana BAK. var. robustior
C. CHR.
Add the synonym: S. robusta BAK. Syn.
Fil. (1868) 429.—Type: HILLEBRAND,
Hawaii (K).
Lygodium SWARTZ.
Delete the generic
BERNH.
synonym Ripidium
(561)
562
FLORA MALESIANA
53b
59a
61b
63
64
TA
76
Lygodium flexuosum (L.) Sw.
Add the synonym: Ugena polymorpha
CAV. Teones61(1801)) 75; t. S95, te 1:
Lygodium longifolium (WILLD.) Sw.
Add to Distr.: Palawan.
Lygodium versteegii CHRIST.
Add to Distr.: Philippines:
Luzon.
Mindanao,
Isoetaceae
Tsoetes LINNE.
See revision of the genus in New Guinea
by J. R. CROFT, Blumea 26 (1980) 177-190,
with key, description of new species I.
hopei and I. stevensii, and under I. neo-
guineensis BAKER a new variety rheo-
phila; also SEM photographs of spores of
all species and much new information on
vegetative morphology and distribution.
Isoetes sp. has been found in Central West
Sumatra, in addition to Mindanao and New
Guinea the third island where the genus is
hitherto discovered in Malesia. Cf. FI.
Males. Bull. 30 (1977) 2767 and J. R.
FLENLEY & R. J. MORLEY, J. Biogeogr. 5
(1978) 57-58. It is assumed to be an un-
described species, differing from the Philip-
pine one in the megaspores. The exact local-
ity is on the westside of Mt Kerintji, midway
the lakes Sati and Landak Panjang, at c.
2080 m altitude, in small, shallow, muddy
depressions in swampy forest, 1°42’S and
101°11'E.
Cyatheaceae
Conspectus of the family Cyatheaceae: new
comments:
I would now raise the subfamilies to the
rank of family, but see no reason for other
changes. PICHI SERMOLLI (Webbia 31,
1977, 333-334, 423-427) includes all in the
order Dicksoniales, with suborders and
families thus:
Thyrsopteridineae:
(Thyrsopteris ).
Culcitineae: Culcitaceae (Culcita).
Dicksoniineae: Dicksoniaceae (including
Cibotium and Cystodium), Lophosoriaceae
(Lophosoria).
Cyatheineae: Cyatheaceae (Sphaeropteris,
Alsophila, Nephelea, Trichopteris, Cyathea,
Cnemidaria).
Metaxyineae: Metaxyaceae (Metaxya).
Subdivision of the genus Cyathea: recent
proposals:
R. M. TRYON (Contr. Gray Herb. 200, 1970,
1-53) has recognized Sphaeropteris
BERNH. as a distinct genus, in which he
includes the tropical American species
Thyrsopteridaceae
80
82b
[ser. II, vol. 1°
mentioned on p. 124 of the present work,
and also several others which disagree in
scales and sori from the specification on p.
65; in my opinion the latter should be
excluded. TRYON limits the genus Cyathea
to some tropical American species, dis-
tinguishing them from all Malesian species
here included in Cyathea subg. Cyathea
solely on the lack of a seta at the apex of
stipe-scales. He has transferred all
Malesian species of Cyathea subg. Cyathea
sensu HOLTTUM 1963 to the genus AI-
sophila (type species A. australis R. BR.).
I agree that subg. Sphaeropteris as
defined on p. 76 is clearly distinct from all
other members of the family, but I cannot
agree that the remaining species, including
those of tropical America, are divisible into
natural groups on such clearly defined
characters. CONANT has’ shown _ that
hybrids exist between species of different
genera as recognized by TRYON, and in
one case such hybrids have good spores
(Rhodora 77, 1975, 441-455).
All species so far examined, of several
different genera as recognized by TRYON
(including Sphaeropteris) have the
chromosome number 69. This is a very
strong indication that Cyathea in the broad
sense adopted in Flora Malesiana is a phy-
letic unity, and I adhere to my recognition
of it as a single genus. As Sphaeropteris is
the only sub-group separable on well-
defined characters, I still place all the rest
in a subgenus Cyathea, the further sub-
division of which seems to me still un-
certain. The species C. australis (R. BR.)
DOMIN (type of Alsophila) has spores
which differ considerably from those of the
majority of Malesian species transferred to
Alsophila by TRYON (see GASTONY,
Amer. J. Bot. 63, 1976, 754, f. 110 and
compare with figures of Malesian species
on p. 751).
Taxonomy of Malesian species of Cyathea:
Since 1963 many new collections have been
made, especially in New Guinea. I have not
been able to examine all of these. The
following data are based on specimens
which have come to my attention; probably
more new species remain to be recognized,
and new information about many species
remains unrecorded.
In Key to the species, after lead 77, add:
77a. Pinnules to 22 mm long, bearing free
tertiary leaflets.
69a. C. nothofagorum
77a. Pinnules to at least 50 mm long, lack-
ing free tertiary leaflets.
Cyathea hunsteiniana BRAUSE.
Add to description: Scales on pinna-rachis
1981]
Addenda, corrigenda et emendanda
563
10la
103b
118b
121b
130b
13S5a
138
140b
140b
elongate, thin, dull brown with a few mar-
ginal setae (WOMERSLEY & VANDEN-
BERG 37293, Western Highlands, 1500 m).
Cyathea oosora HOLTTUM.
Add to Distr.: Pulau Tioman (KLU 19781).
The Celebes specimens here included
should probably rank as a distinct species;
more material is needed.
Insert additional species:
69a. Cyathea nothofagorum HOLTTUM,
Blumea 14 (1966) 327.— Type: PULLEN
5358, N.E. New Guinea, W. Highlands,
Kubur Range, in Nothofagus forest at
2775 m (K).
Aspect of C. microphylloides and C.
perpelvigera (p. 82), having small fronds
and pinnate pinnules, differing in (large)
hemitelioid indusia and lack of hairs on
lower surface of pinna-rachis and costae,
also in scales not bullate.
Cyathea biformis (ROSENST.) COPEL.
Add the following: PULLEN 7295 from the
Upper Fly River at 100m has 1-2 pairs on
small pinnae (the pinnules of which do not
have an incised lamina) near the base of
stipes, and unusually large pinnules; sterile
pinnules to 9 by 2cm, fertile to 10 by
12cm:
Cyathea lurida (BL.) COPEL.
Add the following: A sterile specimen from
a low altitude on limestone near Kuala
Lumpur (J. BoEY 4, KLU) closely resem-
bles this species but differs in pale brown
stipe and rachis. Typical C. lurida occurs
only on high mountain ridges.
Cyathea sangirensis (CHRIST) COPEL.
Reduce to a synonym of the following:
Cyathea felina (ROXB.) MORTON, Contr.
U.S. Nat. Herb. 38 (1974) 344. — Polypodi-
um felinum ROXB. Calc. J. Nat. Hist. 4 (1844)
496.— Lectotype: from Amboina (G);
others at BR and in Herb. J.E. Smith.
Cyathea angiensis (GEPP) DOMIN.
Add the following note: Stipes always spiny,
length of spines 2-7 mm.
Key to the species of subsection Fourniera:
alter as follows:
7. Scales absent from lower surface of veins.
8. Spines on stipe scattered, 1-3 mm long;
soral scales many, covering sorus to
maturity 150. C. tripinnata
8. Spines on stipe copious, to 5 mm long;
soral scales not covering sorus at
maturity .150a. C. jacobsii
Cyathea tripinnata COPEL.
Delete synonym C. arachnoidea (non
HOOK.) BACKER & POSTH.
After Cyathea tripinnata, insert the fol-
lowing:
150a. Cyathea jacobsii HOLTTUM, Rein-
wardtia 8 (1974) 499. — Type: M. JACOBS
141
166a
167
214a
8113, S. Sumatra at 1200-1300 m (L).
This species, occurring in South Sumatra
and West Java, differs from C. tripinnata
in the characters shown in the above
modified key; it also has a different dis-
tribution. Specimens named C. arach-
noidea in W. Java by BACKER & POSsT-
HUMUS (see above) belong to C. jacobsii.
In Key to the species of subsection
Schizocaena, lead 13: alter the word ‘pin-
nules’ at end of line 2 from bottom to
‘segments’.
Cyathea rosenstockii BRAUSE.
Add the following: NAKAIKE 717, Central
District, Papua, Woitape, 1500 m, has flat
pale scales to 4mm long and dark glossy
thick hairs on lower surface of pinna-
rachis; costal scales narrow, pale with dark
bullate bases, smaller distally; on costules
very few small scales.
Cibotium barometz (L.) J. SM.
Add to Distr.: N.E. New Guinea (B. S.
PARRIS in Fern Gaz. 11, 1979, 428).
Culcita PRESL.
Correction in Cytol.: C. macrocarpa n=
66; C. villosa 2n = 232 (tetraploid with base
number 58); from unpublished observations
by G. VIDA on plants in cultivation at Kew.
Lindsaea-group (KRAMER)
Sphenomeris
TAGAWA.
Mapped in Pac. Pl. Areas 3 (1975) 344-345.
Sphenomeris chinensis (L.) MAXON.
Mapped in Pac. PI. Areas 3 (1975) 346-347.
Tapeinidium denhamii (HOOKER) C. CHR.
Mapped in Pac. PI. Areas 3 (1975) 348-349.
Lindsaea bouillodii CHRIST.
For this species MORTON (Contr. U.S.
Nat. Herb. 38, 1974, 385) made the new
combination Lindsaea interrupta (ROXB.)
MorRTON, based on Vittaria interrupta
ROXBURGH, referring L. cambodgensis
CHRIST to its synonymy (following my
earlier misinterpretation of this name:
KRAMER, Blumea 15, 1967, 563). In view
of the gross incompleteness of ROx-
BURGH’s original description, and of the
complete lack of indication of provenance
on the label of the specimen regarded as
type by MorTON, I think it unwise to
adopt ROXBURGH’s (and MORTON’s)
names which I prefer to regard as names of
uncertain application. The same holds in
my opinion for the interpretation of Vit-
taria lunulata ROXBURGH, despite MorR-
TON’s assertions to the contrary (I.c. 386).
Lindsaea tetragona KRAMER.
This East Malesian—Pacific species, map-
ped in Pac. Pl. Areas 3 (1975) 340, was
biflora (KAULFUSS)
564
FLORA MALESIANA
(ser. II, vol. 1°
recently collected as far west as the Nico-
bar Is. (Great Nicobar), and may thus be
expected in western Malesia, too.
Lindsaea polyctena KRAMER.
Mapped in Pac. Pl. Areas 3 (1975) 340-341.
Lindsaea tenuifolia BLUME.
Mapped in Pac. PI. Areas 3 (1975) 340-341.
Lindsaea adiantoides J. SMITH.
Mapped in Pac. PI. Areas 3 (1975) 342-343.
Lindsaea gueriniana (GAUD.) DESVAUX.
Mapped in Pac. Pl. Areas 3 (1975) 332-333.
For the record from Fiji see BROWNLIE,
Pterid. Fl. Fiji (1977) 134.
Lindsaea repens (BORY) THWAITES, with
varieties.
Mapped in Pac. Pl. Areas 3 (1975) 336-337.
Lindsaea carvifolia KRAMER.
This was placed in sect. Odontoloma of
subg. Odontoloma. Its occurrence in
Celebes was reported with a ?. A good
series of specimens from that island
(HENNIPMAN .S. 5264, L) confirms its
presence; furthermore, it shows that sim-
ply pinnate as well as bipinnate leaves
occur side by side, and that the species
should be transferred to sect. Lindsaenium,
where it must be placed between L.
rosenstockii BRAUSE and L. versteegii
(CHRIST) v.A.v.R. In appearance and
width of the pinnule segments it is more
245a
245b
like the latter but differs in less divergent
and usually spathulately broadened seg-
ments. It is undoubtedly distinct from
both; the simply pinnate leaves are indis-
tinguishable from specimens previously
examined. The distinction between sect.
Odontoloma and sect. Lindsaenium may
require reconsideration.
Lindsaea rigida J. SMITH.
Mapped in Pac. PI. Areas 3 (1975) 334-335;
for the record from Fiji see BROWNLIE,
Pterid. Fl. Fiji (1977) 139.
Lindsaea sarawakensis KRAMER.
This was described from a single, incom-
plete collection and was only provisionally
placed next to L. rigida. Two additional
collections from Sarawak, NIELSEN 815
(AAU) and JERMyY 14334 (BM, Z), both
from Mt Mulu, confirm the taxonomic
position assigned to it and provide ad-
ditional data. The rhizome anatomy and
morphology agree with that of Lindsaea
subg. Odontoloma sect. Lindsaenium, and
the difference from L. rigida, as given in
the key (p. 203), proves to be constant.
Further additional data: petiole to c. 35 cm
long; rhizome 1-1.5mm_ diam., long-
creeping, castaneous; scales very much
like those of L. rigida. Fertile pinnae al-
ways with a short apical sorus only.
81
113a
265
266
269b
ADDENDA, CORRIGENDA ET EMENDANDA 2
At couplet 101 in key delete °94, C. in-
cisoserrata’, and insert afterwards:
10la. Apex of stipe-scales setiform; bullate
scales on costules dull brown, smooth
94. C. incisoserrata
10la. Apex of stipe-scales not setiform;
bullate scales on costules almost
white, clathrate. 94a. C. decurrens
After 94. C. incisoserrata, insert:
94a. Cyathea decurrens (HOOK.) COPEL.
Univ. Cal. Publ. Bot. 14 (1929) 356; HoLt-
TUM, Blumea 12 (1964) 248.—Alsophila
decurrens HOOK. Spec. Fil. 1 (1844) 51.—
Type: NIGHTINGALE s.n., Pacific (K).
Stipe-scales broad, castaneous with broad
paler margins, apex broadly rounded or +
attenuate, not setiform; pinnules-lobes
oblique, almost all free but decurrent, all
deeply lobulate; costules covered beneath
with almost white clathrate bullate scales;
indusia very small, hidden by sporangia.
Distr. Solomon Is., New Hebrides, Fiji,
Samoa, Cook I.; in Malesia: New Ireland.
Note. Collected in New Ireland at 2300 m
(CROFT 301); on other islands found at
much lower altitudes. CROFT’s specimen
differs from others in its firmer bullate scales
and in lacking the thick white hairs which are
usually + abundant on the lower surface of
costae in this species; it may represent a
distinct variety. C. decurrens and allied
Pacific species have stipe-scales like those of
the Neotropic species to which TRYON
limits the genus Cyathea; no other known
Paleotropic species have this character.
Teratophyllum METT. ex KUHN.
Add to Distr.: Ceylon.
Teratophyllum, section Teratophyllum.
Add to Distr.: Ceylon.
Teratophyllum aculeatum (BL.) METT. ex
KUHN.
Add to Distr.: Ceylon; rare, first collected by
THWAITES, recently recollected in Sin-
haraja Forest, KOSTERMANS 27853, Oct.
1979 (L).
325a- Bolbitis, series Heteroclitae.
326b
361b
393b
452a
494b
502b
HENNIPMAN has reduced some taxa
which I regard as species, notably B. sim-
plicifolia (HOLTTUM) CHING, B. malac-
censis (C. CHR.) CHING and B. nitens
HOLTTUM. See my paper on Malayan
Bolbitis, Mal. Nat. J. 33 (1980) 79-82 for an
alternative opinion.
Coryphopteris gymnopoda (BAK.) HOLT-
TUM var. bintangensis HOLTTUM.
Add to the description: The type has many
such hairs but recent collections by A. G.
PIGGOTT on Gunong Ulu Kali (Genting
Highlands) at 1830 m show much variation.
Specimens from Borneo are all quite
glabrous between veins.
Line 7 from bottom: Dryopteris penangiana
(HOOK.) C. CHR. was based on Polypodium
penangianum HOOK. The type of this was a
WALLICH specimen which bore the local-
ity Penang, but the species is otherwise only
known from northern India, so I am sure it
was wrongly localized. I did not know any-
thing about this species when I wrote my
book on Malayan ferns.
Dryopteris perakensis was based on Aspi-
dium perakense BEDD.,a species I could not
account for and omitted from my book on
Malayan ferns. I could not find a type in Kew
herbarium, but some years later it was found,
among other BEDDOME specimens, in the
basement. It is based on an immature plant of
Sphaerostephanos polycarpus mixed with a
small fragment of a fertile pinna from ano-
ther plant, an aberrant Christella. VAN
ALDERWERELT VAN ROSENBURGH of
course misinterpreted it.
Sphaerostephanos_ mutabilis
HOLTTuo. Add:
Note. This is near S. aquatilis (no. 44),
differing in narrower pinnae, veins free, and
lower surface quite glabrous; the two should
perhaps be united.
Sphaerostephanos
HOLTTUM.
Add to Ecol.: less commonly to 1000 m.
(BRAUSE)
invisus (FORST. f.)
(565)
F 2 ae o 7 | C?.3 ea te
ary
INDEX TO SCIENTIFIC PLANT NAMES
compiled by
M. J. VAN STEENIS-KRUSEMAN
Epithets or taxa above generic rank have been entered under their name.
Infrageneric epithets have been entered immediately under the generic name to which they belong,
preceded by the indication of their rank (subgenera, sections, etc.). If possible sections are arranged
under the subgenera to which they belong.
Epithets of wrongly identified species precede the correct ones.
Infraspecific epithets have been entered under the specific name to which they belong, preceded by
the indication of their rank (subspecies, varieties, formas, etc.).
Epithets of new names and new combinations have been printed in bold type, synonyms in italics.
An asterisk behind a page number denotes the presence of an illustration of the concerned taxon;
‘map’ printed behind a page number denotes that a map of the concerned taxon is present on that page.
Page numbers in bold type denote main treatment.
Some minor printing errors in plant names have been corrected.
Of synonyms with a double authority, the latter has not always been cited in full. The full authority
can easily be derived from the text.
Scientific plant names mentioned in the Dedication have not been indexed.
(Acrostichum)
crispatulum Clarke
Acrosorus il, XVii
Acrosticheae 255
Abacopteris Fée xxi, 333, 336,
337, 507, 509
aspera Ching 512 Acrostichopteris 37 var. contaminans Clarke
cuspidata (BI.) Ching 536 Acrostichum sensu Fée 289 321
elegans Fée 534 Acrostichum L. ili, vi, vii, xx, cumingii sensu Baker 304,
glandulosa (BI. ) Fée 522
gymnopteridifrons (Hayata)
Ching 513
lineata (Bl.) Holtt. 512, 530
menisciicarpa (BI.) Holtt.
526
multilineata (Wall. ex Hook.)
Ching
var. malayensis Holtt. 512
peltochlamys (C.Chr.) Holtt.
495, 507
philippinarum Fée 507, 512,
513
presliana sensu Ching 513
presliana (Ching) Ching 512
rubicunda (v.A.v.R.) Holtt.
536
salicifolia (Wall. ex Hook.)
Holtt. 535
triphylla (Sw.) Ching 534
var. parishii (Bedd.) Ching
535
truncata Fée 526
urophylla (Mett.) Ching 533
Aconiopteris Pres] 289, 290, 291
Acrophorus ti, xix
adiantoides (Bl.) Moore 249
cuneifolius (Presl) Moore 180
hemipterus (Bory) Moore 237
hymenophylloides (BI.)
Moore 237
repens (Bory) Moore 237
PSS, OI; AOS, QO, Pik
291, 314, 318, 335
sect. Chrysodium (Fée)
Hook. 277
sect. Elaphoglossum Hook.
289
sect. Lomariopsis Hook. 259,
265
sect. Polybotrya Hook. 275
sect. Stenochlaena 265
aculeatum (BI.) Racib. 269
aemulum sensu BI. 308
angulatum BI. 297, 313
apodum sensu Fée 301
appendiculatum Willd. 323
var. costulatum Hook. 325
var. hamiltonianum Baker
322
articulatum Hook. 276
beccarianum Baker 301
blumeanum Fée 311
blumeanum (Fée) Hook. 279,
281
borneense Burck 314
callifolium Bl. 307
celebicum Baker 523
commutatum Mett. ex Kuhn
308, 309
conforme sensu BI. 297, 308
var. Javanicum Mett. 314
conforme Sw. 290, 313
contaminans Wall. 321
307
cumingii Fée 307, 314
decurrens sensu BI. 303, 304
decurrens sensu Hook. 304
decurrens Desv. 292, 303, 314
var. cumingii v.A.v.R. 304
var. major Mett. ex Kuhn
304
var. ornatum Fée 304, 307
dichotomum L. 41
digitatum L. 41
diversifolium Bl. 326
exsculptum Baker 528
flagelliferum Wall. ex Hook.
325
gorgoneum sensu BI. 314
gorgoneum sensu Christ 298
gorgoneum Kaulf. 292, 313
gracile Racib. 269
hamiltonianum Wall. 322
heteroclitum Presl 325
hirtum Sw. 314
junghuhnianum Kunze 307
latifolium Sw. 314
laurifolium sensu Christ 262
laurifolium sensu Fée 297
laurifolium sensu Thw. 308
laurifolium Thouars 314
lessonii Mett. 301
melanostictum Bl. 301
micradenium Fée 314
miniatum Christ 311
(567)
568
(Acrostichum)
modestum Baker 325
norrisii sensu Cesati 301
norrisii Hook. 302
obtusifolium sensu BI. 303
obtusifolium Willd. 304
oligodictyon Baker 380
ophioglossoides Goldm. 298
pellucido-marginatum Christ
298
proliferum Bl. 325
quoyanum Gaud. 327
repandum BI. 329
scandens Hook. 269
sinense Baker 325
smithii Racib. 262
sorbifolium L. 259, 261, 263
speciosum Presl 262
spectabile (Kunze)
262
squamosum sensu Sw. 314
stelligerum Wall. ex Baker
311
subdiaphanum Hook. &
Grev. 291
variabile Hook.
var. rasamalae Racib. 326
viscosum sensu BI. 311
viscosum 292
zollingeri Kunze 326
Actinostachys Wali. ex Hook.
40
boninensis Nakai 42
digitata Wall. ex Reed 41
inopinata Reed 42
wagneri Reed 44
Racib.
Adiantum ii, xvii
chinense (L.) Burm. 182
chinense L. 182
chusanum L. 182
cultratum Willd. 222
ensifolium (Sw.) Poir. 211
falcatum Blanco 180
heterophyllum (Dryand.)
Poir., non Colenso 210
lanceolatum (Labill.) Poir.
211
lobatum (Poir.) Poir. ex
Steud. 219
orbiculatum Lamk 206
tenuifolium Lamk 182
Adiantum group ii, ix, x, xi, xii,
XIll, XIV, XVil
Aglaomorpha ii, xvi
Agyratae 71
Alsophila sensu C.Chr. 115
Alsophila R.Br.—See index p.
169-171 and p. 562. Sub-
sequent references are:
australis R.Br. 562
decurrens Hook. 565
FLORA MALESIANA
(Alsophila)
dryopteroidea Brause 401
fragilis Zoll. & Mor. 512
polypodioides Hook. 348, 349
Amauropelta 334, 338, 341
Ampelopteris Kunze ili, xxi,
340, 341, 342, 375, 378,
386, 387
elegans Kunze 387
prolifera (Retz.) Copel. 385*,
387
Amphicosmia Gardn. 73
alternans Moore 145
javanica (Pr.) Moore 111
manilensis (Pr.) Moore 110
Amphineuron Holtt. 340, 341,
342, 343, 437, 544, 551
attenuatum (O.Ktze) Holtt.
451, 545, 549, 550
ceramicum (v.A.v.R.) Holtt.
401, 545, 549
distinctum (Copel.) Holtt.
544, 545, 548
immersum (BI.) Holtt. 370,
401, 432, 545, 546*, 547
kiauense (C.Chr.) Holtt. 545,
550
opulentum (Kaulf.) Holtt.
544, 545, 546, 547*, 548
paraphysophorum (v.A.v.R.)
Holtt. 545, 550
pseudostenobasis (Copel.)
Holtt. 451, 545, 546*, 549
subattenuatum (Rosenst.)
Holtt. 544, 545, 546*, 547
terminans (Hook.) Holtt. 387,
544, 545, 546*
Anapausia zollingeri Presl 326
Anemia 37, 38, 39
Anemiaceae 39
Angiopteris il, vi, xv, 70
Anisocampium ii, xviii
Ankyropteris 561
Anogramma Il, xVil
Antrophyum iil, xxi
Araiostegia ii, Xvill
Arcypteris ili, xx
Arthrobotrya J.Sm. ii, xx, 267,
275, 276
articulata J.Sm. 276
Arthromeris ti, xvi
Arthropteris tii, xx, 257, 258
obliterata (R.Br.) J.Sm. 253
tenella (Forst.) J.Sm. 258
Aspidiaceae 255, 511
tribe Aspidieae 255
Aspidiales 339
Aspidistes Harris 65, 67
thomasii Harris 333
Aspidium Sw. 335, 336, 338
abortivum sensu Mett. 461
[ser. II, vol. 1°
(Aspidium)
abortivum BI. 429
var. B (var. C?) 429
abruptum Bl. 429
adiantoides Bl. 249
affine Bl. 512, 522, 527, 528
alatellum (Christ) Christ 451
amboinense sensu BI. 558,
559
amboinense Willd. 519, 559
arbuscula sensu Christ 456
aridum D.Don 555
asperum (Presl) Mett. 512
attenuatum Kunze ex Mett.
non Sw. 549
barometz Willd. 165
benoitianum Gaud. 500
blumei Kunze ex Mett. 516
var. subpinnata Mett. 516
brackenridgei Mett. 408
calcaratum BI. 374
callosum Bl. 419
var. B 420
canescens (BI.) Christ 516
f. acrostichoides Christ 523
f. gymnogrammoides
Christ 467
f. nephrodiiformis Christ
519
var. sensu Christ 516
ciliatum Benth. 375
conioneuron (Fée) Mett. 548
crassifolium Bl. 384
crassifolium Mett. 384
cucullatum sensu Christ 488
cucullatum BI. 478
dayi Bedd. 352
decursive-pinnatum (van
Hall) Kunze 354
dimorphum Kunze 458
dissectum sensu Mett. 502
echinatum sensu Christ 379
echinatum Mett. 379
eusorum Thw. 429
excellens (Bl.) Holtt. 462
exiguum Kunze ex Mett. 539
var. b 538, 540
extensum BI. 548
fasciculatum Fourn. 367
ferox Bl. 391
flaccidum BI. 351
glaberrimum Richard 436
glanduligerum Kunze 351,
373
glandulosum sensu. Kunze
512
glandulosum BI. 513, 522, 527
gracilescens BI. 351
grammitoides Christ 373
hallieri Christ 379
harveyi Mett. 555
1981]
(Aspidium)
heterocarpon BI. 457
var. B Blume 458
heterocarpus sensu
498
heterodon BI. 449
heterophyllum (Presl) Hook.
515
hirsutum sensu Christ 488
hirsutum Kunze ex Mett. 455
hispidulum Decne 459, 557
var. B Mett. 459
var. solutum Mig. 459
var. ternatense Miq. 474
hookeri Baker 548
immersum BI. 370, 547
jaculosum Christ 558
javanicum Mett. 449
koordersii Christ 375
laeve Mett. 424
latebrosum Kunze ex Mett.
471
latum Kunze ex Mett. 384
lineatum sensu Mett. 527,
528
lineatum BI. 507, 512, 527
loherianum Christ 489
longipes sensu Christ 498
longipes Bl. 423
megaphyllum Mett. 461
var. hirsutum Mett. 452
menisciicarpon Bl. 526, 527
molle Sw. 557
var. latipinna Benth. 558
multijugum Christ 419, 420
multilineatum Mett. 461, 478
multilineatum Wall. 461, 462
nephrodioides Hook. 548
novo-pommeranicum Brause
ex Rech. 327
nymphale (G.Forst.) Schkuhr
Spills Ey /
obliquatum Mett. 555
obscurum BI. 556
obtusatum Sw. 386, 387
ochthodes 414
opulentum Kaulf. 548
paludosum BI. 347
parasiticum 551, 557
patens sensu BI. 551, 557
pennigerum sensu BI.
462
perakense Bedd. 449, 565
pilosiusculum Mett. 452, 522
pilosiusculum Zipp. 523
polycarpon Bl. 448
procurrens sensu Christ 498
procurrens Mett. 560
productum Kaulf. 456, 460,
461
prolixum Willd. 413
Christ
461,
Index to scientific plant names
(Aspidium)
reinwardtianum Kunze ex
Mett. 374
repandum sensu BI. 512
sagittifolium Bl. 487
setigerum sensu Racib. 348
simplicifolium J.Sm. ex
Hook. 490
simulatum Davenp. 371
solutum BI. 459
sorbifolium Willd. 289
squamigerum Fée 377
stipellatum Bl. 492
subpubescens Bl. 558, 559
var. C BI. 558, 559
terminans Wall. 548
thelypteris
var. squamigerum
Schlechtend. 377
triste Bl. 158
truncatum sensu Christ 425
truncatum Gaud. 430
var. celebicum Miq. 425
tylodes Kunze 413, 414
uliginosum Kunze 348
unitum sensu Mett. 386
unitum sensu Schkuhr 386
unitum (L.) Sw. 477
venulosum BI. 386
vile sensu Racib. 349
warburgii Kuhn & Christ 462
Aspleniaceae 339
Asplenieae
subtribe Blechninae 255
Asplenium ii, xviii, 266
adiantum-nigrum L. 61
bakeri C.Chr. 235
epiphyticum Copel. 266, 270
erectum Willd. 354
Asplenium group ii, ix, Xi, Xiii,
XVii
Astralopteris 315
Athyrium ii, v, xviii, 69, 335,
337, 355
benguetense Christ 351
horizontale Rosenst. 366
hyalostegium Copel. 373
ledermannii Hieron. 368
Athyrium group ii, Xi, Xiil, xiv,
XVIil
Azolla iii, xxi
Balantium Kaulf. 71, 158, 166,
167
auricomum 167
blumei Kunze 159
chrysotrichum Hassk. 159
copelandii Christ 168
culcita 167
glaucescens Link 165
javanicum Copel. 168
569
(Balantium)
pilosum Copel. 168, 169
Belvisia ii, xv
Blechnum ii, vii, xviii
filiforme (A.Cunn.) Ettingsh.
263
punctulatum Sw. 61
Blechnum group ii, x, Xi, Xi,
XViil
Bolbitis Schott ti, xix, 255, 256,
257, 258, 278, 289, 314
ser. Bolbitianae Hennipm.
315, 316, 318, 321
ser. Egenolfianae Hennipm.
3155318, '322;329
ser. Heteroclitae Hennipm.
316, 318, 325, 329, 565
ser. Quoyanae Hennipm. 316,
318, 327
angustipinna (Hayata) Ito
S172 3195320*3321
appendiculata (Willd.) Iwats.
258, 318, 319, 322 map
ssp. appendiculata 317*,
318, 320%, 322; 323, 329
ssp. vivipara (Hook.)
Hennipm. 322 map, 323
var. neglecta Hennipm.
320*, 322 map, 323
var. vivipara 322 map
x arguta (Fée) Ching 329
cladorrhizans (Spr.) Ching
258
coloradica R.W.Brown 315
contaminans Ching 321
copelandii Iwats. 323
cuspidata Ching 325, 326
diversifolia (Bl.) Schott 326,
329
edanyoi Copel. 326
enormis C.Chr. 327
flagellifera Schott 325
x foxii Copel. 329
foxworthyi Ching 326
heteroclita (Presl) Ching 258,
315, 316*, 318, 319, 324%,
325, 329, 330
heteroclita x rhizophylla
Hennipm. 329
hookeriana Iwats. 258, 322
hydrophylla Ching 326
inconstans Ching 326
interlineata (Copel.) Ching
319, 320*, 330
intermedia Iwats. 323
linnaeana C.Chr. 325
malaccensis (C.Chr.) Ching
326, 327, 565
membranacea C.Chr. 326
modesta Ching 325
naumannii Ching 327
570
(Bolbitis)
nigra Ching 326
nitens Holtt. 326, 327, 565
novoguineensis Hennipm.
319, 320*, 324*, 328
ovata (Copel.) Ching 330
parva Ching 327
portoricensis (Spr.) Hennipm.
258
pseudoscalpturata Ching 326
quoyana auct. 329
quoyana (Gaud.) Ching 319,
321, 324*, 327, 328 map,
330
repanda (BI.) Schott 258,
S85 3202) 321327, 329:
330
rhizophylla (Kaulf.) Hen-
nipm. 319, 320*, 322, 323
map, 329, 330
rivularis (Brack.) Ching 319,
327, 328 map
scalpturata (Fée) Ching 319,
320*, 321, 330
serratifolia (Kaulf.) Ching
278
serrulata Iwats. 323
simplicifolia (Holtt.) Ching
326, 565
sinensis (Baker) Iwats. 257,
SIS, SiR SIs spAlies sys}
map, 325, 329
var. costulata Tagawa &
Iwats. 325
x singaporensis Holtt. 320%,
329
sinuata (Presl) Hennipm. 315,
318, 319% 320%, 326; 327
map, 329, 330
x sinuosa (Fée) Copel. 321,
329
nm. foxii (Copel.) Hen-
nipm. 320*, 330
nm. sinuosa Hennipm. 330
stenophylla Ching 325
subcrenata (Hook. & Grev.)
Ching 325, 330
var. prolifera 330
subsimplex Ching 326
taylorii 327
tenuissima Copel. 326
virens (Hook. & Grev.)
Schott 316*, 3118, 321)
322 map
var. compacta Hennipm.
S19 5205509 2p S22 emap
var. deltigera 322 map
var. virens 322 map
vivipara C.Chr. 327
Botrychium ii, xv
Brainea ii, xviii
FLORA MALESIANA
Callipteris ii, xviii
Calymella Presl 6
borneensis Ching 6
circinnata Ching 7, 9
dicarpa Presl 9
microphylla Presl 7
squamosissima Nakai 9
vulcanica Presl 9
Calymmodon il, xvii
Campium Presl 255, 315, 318
angustipinnum Copel. 321
argutum Copel. 329
cuspidatum Copel. 325
diversifolium Copel. 326
enorme Copel. 327
foxworthyi Copel. 326
heteroclitum (Presl) Copel.
325
var. eurybasis Copel. 326
hydrophyllum Copel. 326
interlineatum Copel. 330
kajewskii Copel. 328
linnaeanum Copel. 325
membranaceum Copel. 326
modestum Copel. 325
nigrum Copel. 326
ovatum Copel. 330
parvum Copel. 327
pseudoscalpturatum Copel.
326
quoyanum Copel. 327
repandum Pres] 329
rivulare Copel. 328
scalpturatum Copel. 321
sinense C.Chr. 325
subcrenatum sensu Back. &
Posth. 330
subsimplex Copel. 326
tenuissimum Copel. 326
validum Copel. 327
virens Pres] 321
viviparum Kjellb. 327
Cathetogyratae 71
Ceratopteris 1, Xvii
Cerosora ll, XVil
Cheilanthes 11, xvii
fuscata Bl. 61
gigantea Cesati 349
hirta Sw. 61
multifida Sw. 61
setigera Bl. 349
stenophylla Kunze 349
Cheilolepton Fée 277, 278
blumeanum Fée 279
Cheiropleuria ii, Xv
Chingia Holtt. 334, 340, 342,
378, 389, 397
acutidens Holtt. 389, 390
atrospinosa (C.Chr.) Holtt.
389, 390
bewaniensis Holtt. 390, 393
[ser. Il, vol@ay
(Chingia)
christii (Copel.) Holtt. 388%,
390, 394
clavipilosa Holtt. 390, 393
var. Clavipilosa 390, 393
var. javanica Holtt. 388%,
390, 393
ferox (Bl.) Holtt. 388%, 389,
390, 391, 392, 393
horridipes (v.A.v.R.) Holtt.
390, 394
imponens (Ces.) Holtt. 390,
391
longissima (Brack.) Holtt. 421
lorzingii Holtt. 390, 393
muricata (Brause) Holtt. 389,
391
paucipaleata Holtt. 390, 394
perrigida (v.A.v.R.) Holtt.
390, 394
pricei Holtt. 388*, 389, 390,
392
pseudoferox Holtt. 392
sakayensis (Zeiller) Holtt.
334, 388%, 390, 392
sambasensis Holtt. 390, 393
supraspinigera (Rosenst.)
Holtt. 390, 395, 402
tenerior Holtt. 390, 395
urens Holtt. 389, 390, 395
Chnoophora Kaulf. 73
glauca BI. 135
lanuginosa Jungh. 132
lurida Bl. 121
tomentosa BI. 132
Chorizopteris Moore 277
Christella Lév. 333, 334, 335,
338, 340, 341, 342, 343,
413, 429, 439, 449, 517,
550, 559
sect. Leptochristella Holtt.
551
sect. Pelazoneuron Holtt. 553
acuminata (Houtt.) Lév. 553,
560
adenopelta Holtt. 553, 556
arida (D.Don) Holtt. 332*,
551, 553, 555, 560
buwaldae (Holtt.) Holtt. 551,
553, 554
crinipes (Hook.) Holtt. 378
dentata (Forssk.) Brownsey
& Jermy 332*, 551, 552*,
553, 554, 557
harveyi (Mett.) Holtt. 552*,
55555
hispidula (Decne) Holtt. 459,
SSilk 55275 S53 oS 4s Sue
558, 560
minima Holtt. 553, 554
nana Holtt. 551, 553, 554
1981]
(Christella)
nymphalis (G.Forst.) Pichi
Sermolli 557
papilio (Hope) Holtt. 431,
552515535556
parasitica (L.) Lév. 551, 552*,
S)3}, Sse Spi, Sey Seb
560
peekelii (v.A.v.R.) Holtt. 553,
555
perpubescens (Alston) Holtt.
S524 559,554,555
prolixa (Willd.) Holtt. 555
siamensis (Tagawa & Iwats.)
Holtt. 557
subdentata Holtt. 553, 559
subpubescens (BI.) Holtt.
3B4E Sil, Sil, Seve, Sis
556, 558
timorensis Holtt. 553, 559
Christella group 340
Christensenia ii, Xv
Christiopteris ii, xvi
Chrysodium 255, 291
cuspidatum Kuhn 325
heteroclitum (Presl) Kuhn
325
var. subcrenatum Kuhn
325
zollingeri Kuhn 326
Cibotium Kaulf. ti, xix, 65, 67,
69; 70, 71, 72, 159; 164,
169, 562
arachnoideum (C.Chr.) Holtt.
164, 166
assamicum Hook. 165
barometz (L.) J.Sm._ 164,
165*, 563
var. cumingii v.A.v.R. 166
var. sumatranum Christ
165
chamissoi Kaulf. 164
crassinerva Rosenst. 166
cumingii Kunze 164, 166
var. arachnoideum C.Chr.
166
djambianum Hassk. 165
glaucescens Kunze 165
glaucum sensu Bedd. 165
glaucum sensu J.Sm. 166
regale Linden 165*
Cnemidaria 65, 69, 70, 72, 562
horrida 69, 70
Colysis ti, xvi
cf. pedunculata (Hook. &
Grev.) Ching 330
Condyloneura Christ 291
Coniogramme ii, vi, xvii
Coniopteris 65, 67
hymenophylloides (Brongn.)
Seward 65
(Coniopteris)
murrayana Brongn. 65
Cornopteris ii, Xviii
Coryphopteris Holtt. 331, 333,
334, 338, 340, 341, 342,
343, 354, 370, 437
andersonii Holtt. 357, 358,
364
andreae Holtt. 358, 364
arthrotricha Holtt. 355, 357,
363
athyriocarpa (Copel.) Holtt.
358, 365
athyrioides Holtt. 357, 363
atjehensis Holtt. 357, 362
badia (v.A.v.R.) Holtt. 355,
356*, 359, 369
borealis Holtt. 358, 364
brevipilosa Holtt. 358, 368
coriacea (Brause) Holtt. 358,
368
diaphana (Brause) Holtt. 357,
362
dura (Copel.) Holtt. 359, 368
engleriana (Brause) Holtt.
358, 366
fasciculata (Fourn.) Holtt.
354, 358, 367
gymnopoda (Baker) Holtt.
361
var. bintangensis Holtt.
357, 361, 565
var. gymnopoda 357, 361
var. humilis Holtt. 357, 361
habbemensis (Copel.) Holtt.
355, 356*, 359, 362, 366
hirsutipes (Clarke) Holtt. 355,
356*, 357, 359, 364, 494
horizontalis (Rosenst.) Holtt.
358, 366
hubrechtensis Holtt. 358, 367
inopinata Holtt. 354, 359, 370
iwatsukii Holtt. 357, 362
klossii (Ridl.) Holtt. 356%,
357, 361
lauterbachii (Brause) Holtt.
358, 367
ledermannii (Hieron.) Holtt.
358, 368
meiobasis Holtt. 358, 364
microlepigera Holtt. 358, 366
multisora (C.Chr.) Holtt. 355,
356*, 357, 359, 363
obtusata (v.A.v.R.) Holtt.
357, 362
oligolepia (v.A.v.R.) Holtt.
358, 368
pectiniformis (C.Chr.) Holtt.
355, 357, 363, 364
var. hirsuta Holtt. 357, 363
var. minor Holtt. 357, 364
Index to scientific plant names 57]
(Coryphopteris pectiniformis)
var. pectiniformis 357, 363
platyptera (Copel.) Holtt.
359, 369
plumosa (C.Chr.) Holtt. 357,
360
propria (v.A.v.R.) Holtt. 358,
367
pubirachis (Baker) Holtt. 365
var. major Holtt. 358, 365
var. philippinensis Holtt.
358, 365
var. pubirachis 358, 365
var. sulawesica Holtt. 358,
365
squamipes_ (Copel.)
354, 358, 364, 365
stereophylla (v.A.v.R.) Holtt.
355, 358, 363, 366
subnigra (Brause) Holtt. 359,
369
tahanensis Holtt. 355, 357,
363
tanggamensis Holtt. 358, 365
unidentata (Bedd.) Holtt. 355,
356*, 357, 359
viscosa sensu Holtt. 363
viscosa (Baker) Holtt. 332*,
355, 356*, 360, 365
var. borneensis Holtt. 358,
361
var. poiensis Holtt. 357,
361
var. viscosa 357, 360
Coryphopteris group 339
Crypsinus ii, xvi
Cteisium Michx 39, 45
Ctenitismi} veo 1672 256; 2575
258, 336, 338
habbemensis (Copel.) Copel.
362
Ctenopteris ii, xvii
Culcita Presl ii, xix, 65, 67, 69,
70, 72, 159, 166, 562, 563
subg. Calochlaena 65, 166,
167
subg. Culcita 65, 166, 167
coniifolia (Hook.) Maxon 167
copelandii Maxon 168
dubia (R.Br.) Maxon 70, 167,
168*
formosae (Christ) Maxon 167
javanica (BI.) Maxon 167,
168*
macrocarpa Presl 70, 167, 563
pilosa Christ 168
straminea (Labill.)
167, 169
villosa C.Chr. 167, 169, 563
Culcitaceae 562
Culcitineae 562
Holtt.
Maxon
572 FLORA MALESIANA [ser. Il, volivee
(Cyclosorus)
ecallosus Holtt. 425
edanyoi Copel. 525
Currania ii, Xix Cyclosorus sensu Copel. 414,
Cyathea Smith—See index p. 436
171-175. Subsequent Cyclosorus sensu Holtt. 414,
references are: Ii, iv, Vi, 436 ellipticus sensu Copel. 485
VIGEX VS 75 3935598539) Cyclosorus Link iii, vi, xxi, 333, matutumensis (Copel.)
439, 562, and: 335, 337, 338, 339, 340, 474
subg. Cyathea sensu Holtt. 341, 342, 375, 378, 385, elmerorum (Copel.) Copel.
562 387, 389, 437 485
subg. Sphaeropteris (Bernh.) adenophorus (C.Chr.) Ching erectus Copel. 490
Holtt. 455 extensus (BI.) Ching 548
sect. Schizocaena (J.Sm.)
Holtt. 338
subsect. Fourniera (Bommer)
Holtt. 563
subsect. Schizocaena 563
angiensis (Gepp) Domin 563
arachnoidea sensu Back. &
Posth. 563
atrispora Domin 401
biformis (Rosenst.) Copel.
563
decurrens (Hook.) Copel. 565
felina (Roxb.) Morton 563
hunsteiniana Brause 562
incisoserrata Copel. 565
jacobsii Holtt. 563
lurida (BI.) Copel. 563
microphylloides Rosenst. 563
nothofagorum Holtt. 562, 563
oosora Holtt. 563
perpelvigera v.A.v.R. 563
rosenstockii Brause 563
sangirensis (Christ) Copel.
563
tripinnata Copel. 563
woodlarkensis Copel. 400
Cyatheaceae ii, xi, xii, xv, 3, 65-
169, 291, 562
subfam. Cibotioideae 72
subfam. Cyatheoideae 71
tribe Cyatheae 71, 72
tribe Dicksonieae 71, 72
tribe Lophosorieae 72
subfam. Metaxyoideae 72
subfam. Thyrsopteridoideae
72
tribe Culciteae 72
tribe Thyrsopterideae 72
Cyatheineae 562
Cyclogramma Tagawa 334, 340,
341, 343, 411
auriculata (J.Sm.) Ching 412*
himalayensis Tagawa 412
omeiensis 412
simulans (Ching) Tagawa
412
Cyclopeltis tii, xx
Cyclosorus sensu auctt. plur.
550
Cyclosorus sensu Ching 414,
436
adenostegius (Copel.) Copel.
466
alatellus sensu Copel. 549,
550
alatellus (Christ) Copel. 451
albosetosus (Copel.) Copel.
505
angustifolius (Presl) Copel.
481
aoristisorus sensu Copel. 539
aoristisorus (Harr.) Copel.
538
arfakianus (Baker) Copel.
484
aridus (D.Don) Ching 556
austerus (Brause) Copel. 502
austrophilippinus (Copel.)
Copel. 479
bakeri (Harr.) Copel. 516
beccarianus sensu Holtt. &
Roy 514
beccarianus (Cesati) Copel.
513
benguetensis Copel. 560
boholensis Copel. 456
bordenii sensu Copel. 474
bordenii (Christ) Ching 457
callosus (Bl.) Ching 419
cataractorum Wagn. &
Greth. 468
christii Copel. 394
clemensiae (Copel.) Ching
521
confertus (Brause) Copel.
49]
contiguus (Rosenst.) Ching
557
cuspidatus sensu Copel. 537
cuspidatus (BI.) Copel. 536
debilis (Mett.) Copel. 448
degener Copel. 522
deltipterus Copel. 420
dentatus (Forssk.) Ching 557
dicranogramma 419
diminuens Holtt. 504
diminuta (Copel.) Copel. 515
dimorphus (Brause) Copel.
464, S01
distinctus (Copel.) Copel. 548
diversilobus Ching 452
duplosetosus Copel. 468
falcatulus (Christ) Copel. 557
farinosus (Brause) Copel.
503
ferox sensu Holtt. 392
ferox (Bl.) Ching 391
glaber Copel. 418
glandulosus (Bl.) Ching 522
gongylodes sensu Ching 386
gongylodes (Schkuhr) Link
335, 386
granulosus (Presl) Copel. 524
gregarius Copel. 480
halconensis Copel. 471
heterocarpus sensu Holtt. &
Roy 451
heterocarpus (Bl.) Ching 457
var. glaucostipes (Bedd.)
Holtt. 471
hispidula sensu Copel. 456
hosei (Baker) Copel. 526
imponens sensu Copel. 391
imponens (Ces.) Copel. 391
interruptus sensu Holtt. 545
interruptus (Willd.) H.Ito
332*, 385*, 386, 478
invisus (Forst.f.) Copel 502
irayensis Copel. 495
jacobsii Holtt. 520
lanceolus (Christ) Copel. 529
larutensis (Bedd.) Ching 489
latifolius (Presl) Copel. 526
latipinna (Benth.) Tard. 558
lepidopodus C.Chr. & Tard.
429
leucadenius Copel. 460
lineatus (Bl.) Tard. & C.Chr.
512
lobatus (Copel.) Copel. 474
loherianus (Christ) Copel.
489
magnus Copel. 465
matutumensis (Copel.)
Copel. 467
megaphyllus sensu Copel.
460
megaphyllus (Mett.) Ching
461
melanophlebius (Copel.)
Copel. 514
merrillii (Christ) Copel. 529
micans (Brause) Copel. 501
1981] Index to scientific plant names 573
(Cyclosorus) (Cyclosorus) (Davallia)
microlonchus sensu Copel.
424
microlonchus (Christ) Copel.
423
mindanaensis (Christ) Copel.
557
morobensis (Copel.) Copel.
501
multiauriculatus (Copel.)
Copel. 492
mutabilis (Brause) Copel. 494
nitidulus sensu Copel. 424,
460
nitidulus (Presl) Copel. 430
oblongus (Brause) Copel. 503
obtusifolius (Rosenst.)
Copel. 503
oligodictyus (Baker) Holtt.
380
papilio 556
parasiticus (L.) Farw. 559
paripinnatus (Copel.) Copel.
476
philippinarum (Fée) Copel.
512
polycarpus (Bl.) Holtt. 449
polypterus Copel. 471
porphyricola (Copel.) Ching
472
procurrens (Mett.) Copel. 560
productus sensu Copel. 456
productus (Kaulf.) Ching 460
proliferus (Retz.) Tard. 387
protectus (Copel.) Copel. 485
pustulosus Copel. 513
reederi Copel. 456
repandulus (v.A.v.R.) Ching
557
rhombeus sensu Copel. 525
rhombeus (Christ) Ching 529
rigidus (Ridl.) Copel. 460
riparis (Copel.) Copel. 456
rotumaensis St.John 426
rubidus (Hook.) Copel. 532
rubrinervis (Mett.) Copel. 531
sagittifolioides Copel. 471,
472
samarensis Copel. 524
serratus Copel. 460, 461
sessilipinna (Copel.) Ching
479
simplicifolius (J.Sm.) Copel.
490
sogerensis (Gepp) Copel. 428
spenceri (Christ) Copel. 491
stenodontus Copel. 450
strigosissimus (Copel.)
Copel. 470
subappendiculatus (Copel.)
Copel. 420
subdimorphus Copel. 524
subpubescens sensu Holtt.
558
sumatranus (v.A.v.R.) Ching
558
superbus (Brause) Ching 420
supraspinigerus (Rosenst.)
Copel. 395
suprastrigosus (Rosenst.)
Copel. 457
tephrophyllus (Copel.) Copel.
480
terrestris (Copel.) Copel. 491
todayensis (Christ) Ching 471
toppingii sensu Copel. 485
triphyllus (Sw.) Tard. 534
truncatus (Poir.) Farw. 429
var. kotoensis H.Ito 460
unitus (L.) Ching 478
urdanetensis (Copel.) Copel.
467
urophyllus sensu Copel. 533
urophyllus (Mett.) Tard. 533
vestigiatus (Copel.) Copel.
453
viridis Copel. 418, 419
warburgii (Kuhn & Christ)
Ching 462
weberi Copel. 460
xiphioides (Christ) Copel. 525
Cyrtogonium acuminatum
Brack. 325
laciniatum J.Sm. 327
rivulare Brack. 328
Cystodium J.Sm ii, xix, 65, 69,
72, 162, 562
sorbifolium (Sm.) J.Sm. 67,
163*, 405
Cystopteris il, xvili, 335
Davallia J.Sm. ii, iv, xviii, 167,
179, 185, 198, 256, 258
subg. Odontoloma Hook.
235
achillaeifolia Wall. 266
achilleifolia Wall. ex Hook.
269
adiantifolia Hook. 249
amboynensis Hook. 187
biflora Kaulf. 182
biserrata Bl. 194
blumeana Hook. 214, 216
boryana Presl 237, 241
chinensis (L.) J.Sm. 182
chusana (L.) Willd. 182
decipiens Cesati 180
delicatula Christ 237, 239
denhami Hook. 186
didyma Hedwig 183
firmula Baker 193
flagellifera Hook. & Grev.
193
gracilis Bl. 190
hemiptera Bory 237
hosei Baker 191
intramarginalis Cesati 197
kunzeana Hook. 219
lanceolata Baker 235
lobata (Poir.) Desv. 219
longipinnula Cesati 197
luzonica Hook. 191
microcarpa J.Sm 183
oligophlebia Baker 191
philippinensis Harr. 191
pinnata Cav., non Mett. ex
Kuhn 193
pulchella (J.Sm.) Hook. 249
repens (Bory) Desv. 237
retusa Cav. 180
serrata Roxb. ex Griff. 193
spec. 253
stenoloba Baker 187
tenuifolia (Lamk) Swartz 182
var. chinensis (L.) Moore
182
var. lata Hook. 182
triquetra Baker 216
veitchii Baker 184
Davallia group Ul, 1x, Xi, Xil, xiv,
XViil
Davallioideae 256
Davallodes ii, xviii, 258
Dendroconche ii, xvi
Dendroglossa il, xvi
linneana Fée 325
Dennstaedtia Ii, vil, xviii, 38, 68,
69, 71, 159, 167, 178, 256,
339
ampla (Baker) Bedd. 161
cuneata (Hook.) Moore 161
erythrorachis (Christ) Diels
162
flaccida (Forst.) Bernh. 162
glabrata (Ces.) C.Chr. 162
gomphophylla (Baker) C.Chr.
162
javanica Christ 168
moluccana (BI.) Moore 162
multifida v.A.v.R. 168
paraphysata v.A.v.R. 168
remota (Christ) Diels 162
thombifolia (Baker) C.Chr.
162
scabra (Wall.) Moore 161, 162
scandens (BI.) Moore 162
smithii (Hook.) Moore 162
straminea J.Sm. 169
Dennstaedtia group ii, x, Xi, xiii,
XiV
Dennstaedtiaceae 178
574
FLORA MALESIANA
[ser. II, vol. 1?
(Dennstaedtiaceae)
subfam. Lomariopsidoideae
Holtt. 255, 256
Dennstaedtiales 339
Diacalpe ti, xix
aspidioides BI. 158
Diblemma ii, xvi
Dichorexia Presl 73, 76
latebrosa Presl 115
Dicksonia L’Hérit. ii, vil, xix,
65, 67, 68, 69, 70, 71, 72,
74, 108, 158, 167
sect. Eudicksonia Hook. &
Baker 158, 166
subg. Balantium Hook. 158,
166
ampla Baker 161
antarctica Labill. 69, 160*
arborescens L’Hérit. 158,
159, 167
archboldii Copel. 159, 160
assamicum Griff. 165
baranetz Link 165
blumei sensu C.Chr. 159
blumei (Kunze) Moore 69,
159, 160*
chrysotricha (Hassk.) Moore
159
copelandii Christ 168
culcita L’Hérit. 167
cuneata Hook. 161
deltoidea Hook. 161
erythrorachis Christ 162
flaccida (Forst.) Sw. 162
glabrata Cesati 162
gomphophylla Baker 162
grandis Rosenst. 159, 161
hieronymi Brause 159, 161
japonica Sw. 162
javanica BI. 168
kingii Bedd. 162
lanigera Holtt, 159, 161
ledermannii Brause 161
linearis Cav. 162, 190
mollis Holtt. 159, 160*
moluccana BI. 162
moluccana Roxb., non BI.
163
neglecta Fée 162
nephrolepioides Christ 162
papuana F.v.M. 163
remota Christ 162
repens Bory 235, 237, 241
rhombifolia Baker 162
scabra Wall. ex Hook. 162
scandens Bl. 162
schlechteri Brause 161
sciurus C.Chr. 159, 160, 161
smithii Hook. 162
sorbifolia Sm. 163
straminea Labill. 169
(Dicksonia )
strigosa Thunb. 162
torreyana Brack. 169
zippeliana Kunze 162
Dicksonia group Il, x, Xili, xix
Dicksoniaceae 65, 71, 562
Dicksoniales 339, 562
Dicksoniineae 562
Dicranopteris sensu Und. 11,
18
Dicranopteris Bernh. ii, viii, xv,
le 5 3B IO 4 SUS Silks
S59
subg. Acropterygium 29
subg. Dicranopteris 29
clemensiae Holtt. 31*, 32
crassifolia Nakai 34
curranii Copel. 3, 28*, 29*,
B0N3IEI357
dolosa Copel. 26
ferruginea Copel. 34
ferruginea Hosok. 561
lessonii Nakai 31
linearis (Burm. f.) Und. 2, 3,
DIE 2963 0S 332533
var. alternans (Mett.)
Holtt. 29*, 31*, 33, 35, 36
var. altissima Holtt. 33, 36
var. bidentata v.A.v.R. 561
var. crassifrons v.A.v.R.
561
var. demota Holtt. 33, 36
var. ferruginea (BI.) Holtt.
33, 34, 561
var. inaequalis (Rosenst.)
Holtt. 33, 36
var. latiloba Holtt. 33, 34
var. linearis 29*, 31*, 33,
34, 35*, 36
var. montana Holtt. 29*,
33, 36
var. rigida (BI.) Holtt. 33,
34
var. stipulosa Christ 561
var. subferruginea
(Hieron.) Nakai 31*, 33,
34, 561
var. subpectinata (Christ)
FOltt 297 33555556;
561
var. subspeciosa
BS Bik, BB), BG
var. tetraphylla (Rosenst.)
Nakai 29*, 33, 36
opposita Nakai 32
pectinata (Willd.) Und. 1, 3
pubigera (BI.) Nakai 29%, 31*,
32*, 34
speciosa (Presl) Holtt. 2, 3,
PE). Bil, 2944, 3S)
tomentosa Copel. 561
Holtt.
(Dicranopteris)
warburgii sensu Nakai 35
Dictyocline ili, xxi
Dictyoglossum J.Sm 289, 291
Didymochlaena li, xix, 335
polycarpa (Bl.) Baker 448
Dimorphopteria Tagawa &
Iwats. 507, 516
moniliformis Tagawa &
Iwats. 507, 519
Diplaziopsis ti, xviii, 157
Diplazium 11, xviti, 157
Diplopterygium Nakai 11
blotianum Nakai 17
conversum Nakai 17
elmeri Nakai 14
giganteum (Hook.) Nakai 561
laevissimum Nakai 11
longissimum Nakai 14
norrisii Nakai 15
novoguineense Nakai 15
volubile Nakai 13
Diplora il, xviii
Dipteris ti, xv, 3
lobbiana (Hook.) Moore v,
536
novoguineensis Posth. 7
Disphenia Pres 73, 76
orientalis Kunze 86
Doodia ii, xviii
Doryopteris ii, xvil
Drymoglossum ii, xvi
Drynaria ti, Xvi
Dryoathyrium 11, xviii
Dryopolystichum iti, xx
Dryopterideae 255
Dryopteridoideae C.Chr. 257
Dryopteris Adans. ii, v, xix, 65,
67, 69, 158, 167, 256, 257,
266; 335; 53365e5Si-eSSSs
339, 507
sect. Abacopteris C.Chr. 507,
509
subg. Cyclosorus C.Chr. 386
subg. Steiropteris C.Chr. 378,
398
abortiva (Bl.) O.Ktze 429
acanthocarpa Copel. 532
acromanes Christ 452, 453
acrostichoides (Christ)
v.A.v.R. 523, 524
var. lanceola (Christ)
v.A.v.R. 523, 529
var. rhombea _ (Christ)
v.A.v.R. 523, 529
acuminata Rosenst. 558
adenophora C.Chr. 455
adenostegia Copel. 466
africana (Desv.) C.Chr. 542
alatella (Christ) C.Chr. 451
albociliata Copel. 560
1981]
(Dryopteris)
albosetosa Copel. 505
alta Brause 404
amaiensis Rosenst. 536
angustata Copel. 456
angustipes sensu C.Chr. 472
angustipes Copel. 471
appendiculata (BI.) C.Chr.
452
aquatilis Copel. 466
aquatiloides Copel. 520
arborea v.A.v.R. 484
arbuscula sensu Kjellb. &
C.Chr. 454
archboldii C.Chr. 485
arfakianus (Baker) C.Chr.
484
arida (D.Don) O.Ktze 555
var. ebeneorachis Copel.
556
arisanensis Rosenst. 351
armata Rosenst. 392
atasripii Rosenst. 504
athyriocarpa Copel. 365
atrispora (Domin) C.Chr.
158, 401
var. varievestita C.Chr. 401
atrospinosa C.Chr. 390
attenuata O.Ktze 549
aureoviridis Rosenst. 353
auriculata (J.Sm.) Ching 412
auriculifera v.A.v.R. 383
aurita (Hook.) C.Chr. 345
austera Brause 502
austrophilippina Copel. 479
backeri v.A.v.R. 349
badia v.A.v.R. 369
bakeri (Harr.) Copel. 464, 516
baramensis C.Chr. 473
bartlettii Copel. 522
basilaris C.Chr. 460
basisora Copel. 409
batacorum Rosenst. 476
var. winkleri Rosenst. 429
batjanensis Rosenst. 467
beccariana (Cesati) C.Chr.
383, 513
beddomei (Baker) O.Ktze 371
var. nadiwonensis v.A.v.R.
371
belensis Copel. 403
benoitiana (Gaud.) v.A.v.R.
500
berastagiensis C.Chr. 423
besukiensis v.A.v.R. 547
bipinnata Copel. 405
bordenii Christ 457, 458, 465
borneensis (Hook.) O.Ktze
528
brachyodus sensu v.A.v.R.
382
Index to scientific plant names
(Dryopteris)
brassii C.Chr. 371
brooksii Copel. 423
brunnea (Wall.) C.Chr. 347
brunneo-villosa C.Chr. 349
brunnescens C.Chr. 452
bungoensis C.Chr. 487
calcarata (Bl.) O.Ktze 374
calcicola C.Chr. 462
callosa (Bl.) C.Chr. 419
var. sumatrana v.A.v.R.
420
calva Copel. 351
canescens (Bl.) C.Chr. 497
var. sensu C.Chr. 490, 516
var. degenera Christ 522
var. lobata Christ 474
var. novoguineensis Brause
514
var. subsimplicifolia Christ
514
canlaonensis Copel. 479
caudiculata v.A.v.R. 423
caudiculata Rosenst. 466
celebica (Baker) Copel. 523
ceramica Rosenst. non
v.A.v.R. 488
cesatiana C.Chr. 513
chamaeotaria Christ 524
chlamydophora Rosenst. 384
christopherseni Christ 421
ciliata (Benth.) C.Chr. ex Wu
375
clemensiae Copel. 521
compacta Copel. 527
conferta Brause 491
confusa Copel. 539
conterminoides C.Chr. 367
contigua Rosenst. 557
cordifolia v.A.v.R. 526
coriacea Brause 368
var. elata Brause 368
costata (Brack.) Maxon 421
crassa Copel. 408
crassifolia (Bl.) O.Ktze 377,
384
var. motleyana (Hook.)
v.A.v.R. 381, 384
var. purpureo-lilacina
C.Chr. 384
crassinervia C.Chr. 414
creaghii (Baker) C.Chr. 353
cucullata (Bl.) Christ 478
var. mucronata Christ 478
cuspidata (Bl.) Christ 536
cyrtocaulos v.A.v.R. 473
dayi (Bedd.) C.Chr. 353
debilis (Mett.) C.Chr. 448,
522
decursive-pinnata (van Hall)
O.Ktze 354
575
(Dryopteris)
deltiptera Copel. 420
dentata (Forssk.) C.Chr. 557
diaphana Brause 362
dichrotricha Copel. 453, 482
dichrotrichoides v.A.v.R. 482
dicranogramma v.A.v.R. 419
didymosora (Bedd.) C.Chr.
560
diminuta Copel. 464, 515
dimorpha Brause 501
distans sensu v.A.v.R. 347
distincta Copel. 353, 548
divergens Rosenst. 384
diversifolia v.A.v.R. 547
diversiloba (Presl) Christ 452,
453
var. acrostichoides (J.Sm.)
Christ 525
subvar. lanceola Christ
529
subvar. rhombea Christ
529
diversivenosa v.A.v.R. 353
dura Copel. 368
echinata (Mett.) O.Ktze 379
echinospora v.A.v.R. 495
elliptica Rosenst. 474
elmerorum Copel. 485, 486
eminens (Baker) C.Chr. 449
endertii C.Chr. 381
engleriana Brause 366
var. hirta C.Chr. 367
ensipinna Brause 404
erubescens sensu Christ 549
euaensis Copel. 555
euryphylla Rosenst. 511
var. teysmannii v.A.v.R.
511
excrescens Copel. 522
exigua sensu v.A.v.R. 539
exsculpta sensu C.Chr. 530
exsculpta (Baker) Copel. 528
extensa (Bl.) O.Ktze 397, 548
falcatipinnula Copel. 406
farinosa Brause 503
ferox (Bl.) O.Ktze 391
var. calvescens sensu
v.A.v.R. 394, 395
var. calvescens Christ
391, 394, 395
var. mitis Rosenst. 395
finisterrae Brause 432
firmula (Baker) C.Chr. 519
flaccida (Bl.) O.Ktze 352
flavo-virens Rosenst. 353
foxii Christ 431
fulgens Brause 403
glanduligera (Kunze) Christ
373
glandulosa sensu Christ 513
576
FLORA MALESIANA
[ser. II, vol Rit?
(Dryopteris)
glandulosa (Bl.) O.Ktze 522
glaucescens Brause 469, 470
glaucostipes (Bedd.) C.Chr.
471
gongylodes sensu v.A.v.R.
386
gracilescens (Bl.) O.Ktze 351
var. calva (Copel.) C.Chr.
351
var. chinensis Christ 359
gracilis Copel. 371
granulosa (Presl) C.Chr. 524
gymnocarpa Copel. 542
gymnopoda (Baker) C.Chr.
361
gymnopteridifrons Hayata
513
habbemensis Copel. 362
hallieri (Christ) C.Chr. 379
hamifera v.A.v.R. 496
harveyi (Mett.) O.Ktze 555
hastatopinnata Brause 470
heterocarpa (BI.) O.Ktze 457
hewittii Copel. 527
himalayensis C.Chr. 412
hirsuta (Kunze ex Mett.)
O.Ktze 455
hirtisquamata Hayata 352
hirtopilosa Rosenst. 557
hispidifolia v.A.v.R. 466
hispidula sensu Brause 470
hispidula sensu C.Chr. 481
hispidula (Decne) O.Ktze 557
hispiduliformis C.Chr. 470
holophylla C.Chr. 526, 527
horizontalis (Rosenst.)
v.A.v.R. 366
horridipes v.A.v.R. 394
hosei (Baker) C.Chr. 526
hunsteiniana Brause 403
immersa
var. ligulata Christ 431
imponens (Ces.) C.Chr. 391
impressa (Desv.) Posth. 548
inclusa Copel. 423
‘inconspicua Copel. 493
indica
var. malayensis (Bedd.)
v.A.v.R. 485
indochinensis Christ 359, 360
interrupta sensu. Back. &
Posth. 545
invisa (Forst. f.) O.Ktze 502
iridescens v.A.v.R. 522
japonica
var. stricta sensu Rosenst.
364
keysseriana Rosenst. 432
kiauensis C.Chr. 550
kinabaluensis Copel. 361
(Dryopteris)
klossii (Ridl.) v.A.v.R. 361
koordersii (Christ) C.Chr. 375
korthalsii Rosenst. 526, 527
kotoensis Hayata 460
labuanensis C.Chr. 528
laevis (Mett.) C.Chr. 424
lanceola (Christ) Copel. 529
larutense 437
latebrosa (Mett.) C.Chr. 471
latipinna (Benth.) O.Ktze 558
lauterbachii Brause 367
leucolepis (Presl) Maxon 349
ligulata (Presl) O.Ktze 431
linearis Copel. 369
lineata sensu C.Chr. 528
var. subacrostichoides
v.A.v.R. 529
lithophylla Copel. 496
lobangensis C.Chr. 479
logavensis Rosenst. 451, 549
loheriana (Christ) C.Chr. 489
longipes (BI.) O.Ktze 424
longissima
var. novoguineensis
Rosenst. 548
luerssenii sensu Christ 414
luerssenii (Harr.) C.Chr. 431
luzonica Christ 421, 424
var. puberula Christ 424,
460
macgregorit (Baker) C.Chr.
367
macrotis (Hook.) O.Ktze 527
malayensis C.Chr. 522
maquilingensis Copel. 524
marattioides Alston 402
marthae v.A.v.R. 374
mataanae Brause 365
matutumensis Copel. 467
media v.A.v.R. 352
megalocarpa v.A.v.R. 359,
360
megaphylla (Mett.) C.Chr.
461
megaphylloides v.A.v.R. 452
megaphylloides Rosenst. 481
melanophlebia Copel. 514
menisciicarpa (BI.) Posth.
$12, 526
merrillii Christ 529
micans Brause 501
microcarpa v.A.v.R. 371
microloncha Christ 423
mindanaensis Christ 557, 558
mirabilis Copel. 526
mixta Rosenst. 457, 458
mollis (Sw.) Hieron. 557
moluccana C.Chr. 549
monodonta C.Chr. 359
morobensis Copel. 501
(Dryopteris)
motleyana (Hook.) C.Chr.
381
var. dulitensis C.Chr. 382
moussetii Rosenst. 346
multiauriculata Copel. 492
multilineata (Bedd.) O.Ktze
461
multiseta (Baker) C.Chr.
349
multisora C.Chr. 359
muricata Brause 39]
var. marginata Brause 391,
392
var. obscura Brause 391,
392
mutabilis Brause 494
nephrolepioides C.Chr. 431
normalis (Sod.) C.Chr. 413
norrisii Rosenst. 485
notabilis Brause 402
novoguineensis Brause 469,
470
oblanceolata Copel. 513
oblonga Brause 503
obtusata v.A.v.R. 362
obtusifolia Rosenst. 503
oligodictya (Baker) C.Chr.
380
oligolepia v.A.v.R. 368
oosora (Baker ) C.Chr. 500
ophiura Copel. 409
orbicularis C.Chr. 548
oxyotis Rosenst. 527
paleata Copel. 380
pallescens Brause 383
paraphysata Copel. 481
paraphysophora v.A.v.R. 550
parasitica (L.) O.Ktze 559
var. falcatula Christ 557
paripinnata Copel. 476
patens sensu v.A.v.R. 555
patens (Sw.) O.Ktze 413
pectiniformis C.Chr. 363
peekelii v.A.v.R. 555
peltochlamys C.Chr. 495
penangiana (Hook.) C.Chr.
565
var. calvescens sensu
Copel. 393
pennigera (Forst. f.) C.Chr.
421
perakensis 565
var. sumatranus v.A.v.R.
452
perglandulifera v.A.v.R. 454
var. firmior v.A.v.R. 454
perpilifera v.A.v.R. 456, 457
perpubescens Alston 554
persquamifera v.A.v.R. 379
petrophila Copel. 435
1981]
(Dryopteris)
philippinensis (Baker) Copel.
460
pilosiuscula (Racib.) C.Chr.
522
pilososquamata v.A.v.R. 481
var. obtusata v.A.v.R. 449,
481
pinnata Copel. 375
platyptera Copel. 369
plumosa C.Chr. 360
plurifolia v.A.v.R. 492
polycarpa (BI.) Christ 449
polyotis C.Chr. 464
polytricha v.A.v.R. 380
porphyricola’ sensu C.Chr.
472
presliana Ching 512
procurrens (Mett.) O.Ktze
560
producta (Kaulf.) C.Chr. 460,
461
prolifera (Retz.) C.Chr. 387
propria v.A.v.R. 367
protecta Copel. 485
pseudoamboinensis Rosenst.
558
pseudoarbuscula v.A.v.R.
456
pseudohirsuta Rosenst. 460
pseudomegaphylla v.A.v.R.
452
pseudoreptans C.Chr. 522
pseudostenobasis Copel. 451,
549
pteroides sensu v.A.v.R. 545
pterospora v.A.v.R. 487
pubirachis (Baker) C.Chr.
365
quadriaurita Christ 404
quadriquetra v.A.v.R. 406
ramosii Christ 537
regis Copel. 434
repandula v.A.v.R. 557
repentula Christ 373
rhombea sensu Copel. 525
rhombea (Christ) Copel. 529
ridleyana Brause 501
ridleyi (Bedd.) C.Chr. 361
riedleana (Gaud.) v.A.v.R.
456
rigidifolia v.A.v.R. 369
riparia Copel. 456
robinsonii (Ridl.) C.Chr. 361
roemeriana Rosenst. 499
rosenburghii C.Chr. 484
rubidum (Hook.) C.Chr. 532
rudis (Kunze) C.Chr. 501
rufopilosa Brause 501
sagittifolia (B!.) O.Ktze 487
savaiensis (Baker) C.Chr. 404
(Dryopteris)
schizophylla v.A.v.R. 407
schlechteri Brause 407
var. djammuense Brause
407
schultzei Brause 432, 433
sepikensis Brause 484, 485
septempedalis Alston 402
sessilipinna Copel. 479
setigera sensu C.Chr. 348
setigera (Bl.) O.Ktze 349
var. pallida v.A.v.R. 348,
349
simillimum C.Chr. 520
simplicifolia (J.Sm.) Christ
490
singalanensis (Baker) C.Chr.
352
sogerensis Gepp 428
spenceri Christ 491
squamaestipes sensu Hayata
413
squamipes Copel. 364
stegnogramma (BI.) C.Chr.
542
stenobasis C.Chr. 451, 549
stereophylla v.A.v.R. 366
stipellata (Bl.) O.Ktze 492
var. obtusata v.A.v.R. 476
strigosissima Copel. 470
subalpina v.A.v.R. 474
subappendiculata Copel. 420
subattenuata Rosenst. 547
subconformis C.Chr. 519
subdimorpha Copel. 384
subfalcinella v.A.v.R. 485
sublaxa Hayata 351
submollis v.A.v.R. 558
subnigra Brause 369
subpubescens_ (BI.)
558
var. major C.Chr. 558
subulifolia v.A.v.R. 494
subviscosa v.A.v.R. 361, 364
subviscosa Rosenst. 364
sumatrana sensu Troll 430
sumatrana v.A.v.R. 415, 558
sumbawensis C.Chr. 432
superba Brause 420
superficialis v.A.v.R. 549
supraspinigera Rosenst. 395
suprastrigosa Rosenst. 457,
458
supravillosa C.Chr. 362
tandikatensis v.A.v.R. 463
tenompokensis C.Chr. 530
tephrophylla Copel. 480
terrestris Copel. 491
teuscheri v.A.v.R. 381
thelypteris
var. squamigera C.Chr. 377
@Chr:
Index to scientific plant names BI)
(Dryopteris)
tibangensis C.Chr. 485
todayensis Christ 419, 471
toppingii sensu C.Chr. 485
toppingii Copel. 500
transversaria (Brack.) C.Chr.
421
var. rodigasiana
426
trichodes Rosenst. 348
trichopoda C.Chr. 380
triphylla (Sw.) C.Chr. 534
truncata (Poir.) C.Chr. 429
tuberculata (Ces.) C.Chr. 407
tylodes (Kunze) Christ 414
uliginosa sensu. Back. &
Posth. 349
uliginosa (Kunze) C.Chr. 348
uniauriculata Copel. 467
unita (L.) O.Ktze 477
uraiensis Rosenst. 352
urdanetensis Copel. 467, 479,
521
urophylla sensu v.A.v.R. 513
urophylla (Mett.) C.Chr. 532
var. nitida Holtt. 532
var. novoguineensis
Rosenst. 512
var. peraspera v.A.v.R. 526
verruculosa v.A.v.R. 512, 526
var. sumatrana v.A.v.R.
526
vestigiata Copel. 453
villosipes Gepp 369
vinosicarpa v.A.v.R. 381, 382
viscosa (Baker) O.Ktze 360
var. kamborangana C.Chr.
361
wantotensis Copel. 410
warburgii (Kuhn & Christ)
C.Chr. 462
wariensis Copel. 403
weberi Copel. 482
williamsii Holtt. 450
xiphioides Christ 525
zippelii Rosenst. 529
Dryopteris group ii, Xi, xii, xix
@CChr
Eatoniopsis Bommer 73
Edanyoa Copel. 315
difformis Copel. 318, 326, 329
Egenolfia Schott ii, xix, 255,
256, 258, 315, 318, 322
appendiculata J.Sm. 322, 329
bipinnatifida J.Sm. 325
fluviatilis Copel. 323
hamiltoniana Fée 322
intermedia Fée 323
rhizophylla Fée 323
serrulata Fée 323
sinensis Maxon 325
578
FLORA MALESIANA
[ser. II, vol. 1°
(Egenolfia)
vivipara C.Chr. 322
Elaphoglossaceae_ Pichi Ser-
molli 255
Elaphoglossum J.Sm. ii, iv, xix,
255250; 25>, 25852289,
314
alstonii Tard.-Blot 297
amblyphyllum Bell 290, 294,
303, 304*
angulatum sensu Copel. 307
angulatum (BI.) Moore 292,
296*, 297, 314
angustatum (Schrad.) Hieron.
313
angustifrons Holtt. 293, 303
annamense C.Chr. & Tard.-
Blot 293, 299%, 303
apiculatum Holtt. 293, 299
apoense Holtt. 293, 302, 305*
arachnoideum Holtt. 294, 303
archboldii Copel. 293, 299, 303
austrosinicum Matt. & Chr.
310
basilanicum Copel. 301
beccarianum C.Chr. 301
blumeanum (Fée) J.Sm. 292,
294, 311, 312*, 313, 314
var. blumeanum 311
var. philippense Christ MS
311
bolanicum Rosenst. 292, 295*
borneense (Burck) C.Chr. 314
brassii C.Chr. 309
brevifolium Holtt. 294, 307
brunneum Copel. 292, 293,
295
calanasanicum Holtt. 293, 301
callifolium sensu Holtt. 308
callifolium (BI.) Moore 292,
294, 306*, 307, 308, 314,
S17
commutatum sensu v.A.v.R.
307
commutatum (Mett. ex
Kuhn) v.A.v.R. 294,
296*, 308, 314
conforme sensu Bl. 308
conforme sensu Copel. 307
conforme (Sw.) J.Sm. 290,
291, 292, 309, 313
copelandii Christ 311
cumingii
var. papuanum C.Chr. 304
decurrens sensu v.A.v.R. 303
decurrens sensu C.Chr. 303
decurrens Desv. 314
var. crassum C.Chr. 299
dolichocaulon v.A.v.R. 297
elmeri Copel. 304, 307
(Elaphoglossum)
fuscum Copel. 297
gayanum (Fée) Moore 290
gorgoneum (Kaulf.) Brack.
313
habbemense Copel. 292, 295
hellwigianum Rosenst. 292,
295
heterolepium v.A.v.R. 303,
S11F 312+, 313"
heterostipes Holtt. 293, 294,
301
hirtum (Sw.) C.Chr. 314
idenburgensis Holtt. 293, 298
indrapurae Holtt. 294, 310
junghuhnianum Moore 307
latemarginatum Holtt. 293,
298
laticuneatum Copel. 297
latifolium sensu Bedd. 297
latifolium (Sw.) J.Sm. 314
laurifolium sensu_ v.A.v.R.
307
laurifolium sensu Bedd. 297
laurifolium (Thouars) Moore
308, 314
lepidopodum C.Chr. ex
Ogata 304
luzonicum Copel. 294, 304,
305*, 314
macgregorii Copel. 307
malayense Holtt. 294, 308,
310
melanochlamys Holtt. 293,
302
melanostictum sensu Holtt.
302
melanostictum (BI.) Moore
256, 290, 293, 301
merrillii Christ 298
microphyllum v.A.v.R. 314
minahassae v.A.v.R. 297
miniatum (Christ) Christ 294,
311, 312*
muscosum 292
negrosensis Holtt. 294, 307
nesioticum Holtt. 293, 302
nigripes Holtt. 293, 302
norrisii (Hook.) Bedd. 293,
300*, 301
novoguineense Rosenst. 294,
309
obtusifolium Bell 303
obtusifolium Brack. 303
obtusifolium J.Sm. 304, 314
ogatai C.Chr. 297
ophioglossoides (Goldm.)
Holtt. 293, 298, 301
pallescens Holtt. 293, 303
pellucido-marginatum (Christ)
C.Chr. 293, 298, 314
(Elaphoglossum)
pellucidum Gaud. 314
peninsulare Holtt. 301
permutatum v.A.v.R. 308
var. mutatum v.A.v.R. 307,
308
petiolatum sensu v.A.v.R.
311
petiolatum (Sw.) Urban 311
planicosta Holtt. 294, 307
pullenii Holtt. 293, 294, 297
pumilum Lam & Verhey 292,
295
rapaense Copel. 292
recommutatum Holtt. 294,
308, 313
reineckei Hieron. & Laut. 307
repens Copel. 293, 298
resiniferum Holtt. 290, 294,
313
robinsonii Holtt. 294, 310
salicifolium (Willd. ex Kaulf.)
Alston 311
samoense Brack. 292
sclerophyllum v.A.v.R. 293,
297
societarum Copel. 292
sordidum Christ 294, 309*
spathulatum (Bory) Moore
292
spongophyllum Bell ex Holtt.
294, 310
stenolepis Bell ex Holtt. 294,
303
sumatranum Holtt. 294, 310,
311
thamnopteris Holtt. 293, 298
vepriferum Holtt. 293, 302
yunnanense sensu Holtt. 311
yunnanense (Baker) C.Chr.
311
Equisetaceae ii
Equisetales 1
Equisetum ii
Filicales 11, vii
Fourniera Bommer 73, 138
Gisopteris Bernh. 39, 44
Glaphyropteridopsis Ching 389
erubescens (Hook.) Ching
389
Glaphyropteris
sect. Cyclogramma H.Ito 411
Gleichenia Smith ti, v, xv, 1, 2,
3, 4, 6, 29, 70, 71, 169
subg. Diplopterygium Holtt.
1, 35:45. 7, 10% 1a Sao»
20, 33
subg. Eugleichenia Diels 6
subg. Gleichenia 1, 2, 3, 4, 6
1981]
(Gleichenia)
subg. Mertensia Hook. 1, 2,
3, 5, 18, 19, 20
sect. Dicranopteris v.A.v.R.
11, 18, 27
sect. Diplopterygium Diels
11
sect. Heteropterygium Diels
27
sect. Holopterygium Diels
18
sect. I Hook. 11
sect. IJ Hook. 18
sect. III Hook. 27
sect. IV Hook. 27
alstonii Holtt. 5, 23
amboinensis v.A.v.R. 27
amoena v.A.v.R. 20, 26
angustiloba Holtt. 5, 14
arachnoidea (Hassk.)
v.A.v.R. 13
arachnoidea Cunn. 13
arachnoides Mett. 13
articulata J.Sm. 230
barbula C.Chr. 26
bifurcata Bl. 561
blotiana C.Chr. 5, 17
bolanica Rosenst. 6, 24, 561
borneensis C.Chr. 6
bracteata Bl. ex Hook. &
Baker 21
brassii C.Chr. 6, 24
brevipinnula Holtt. 5, 17
bullata Moore 5, 13, 561
candida Rosenst. 26
caudata Copel. 561
circinnata Sw. 7, 11
var. borneensis Baker 6
clemensiae (Copel.) Holtt. 5,
14, 561
var. clemensiae 14
var. membranacea Holtt. 14
conversa v.A.v.R. 5, 10*, 17
crassifolia Copel. 34
cunninghamii 3
deflexa Holtt. 5, 15
dicarpa R.Br. 4, 7*, 9, 10, 11
var. volcanica Christ 9
dichotoma Hook. 33
var. alternans Mett. 36
var. ferruginea Racib. 34
var. malayana v.A.v.R. 31
var. normalis Mett. 33
var. pubigera Mett. 32
var. rigida Mett. 34
var. tenera Mett. 31
dolosa C.Chr. 26
var. virescens Hieron. ex
Brause 27
elmeri Copel. 5, 14
erecta C.Chr. 5, 6, 22, 561
Index to scientific plant names
(Gleichenia)
excelsa J.Sm. ex Hook. 14
ferruginea Bl. 34
flabellata R.Br. 6, 25
gigantea Wall. ex Hook. 3,
561
glauca auct. 561
glauca v.A.v.R. 14
glauca (Thunb.) Hook. 33
var. arachnoides C.Chr. 13
hallieri Christ 26
hantonensis Wanklyn 2
hermannii R.Br. 33
var. pubigera BI. 32
var. rigida Bl. 34
var. tenera BI. 31
var. venosa BI. 31
hirta BI. 6, 18*, 19*, 25
var. amboinensis (v.A.v.R.)
Holtt. 27
var. amoena (v.A.v.R.)
Holtt. 26
var. candida (Rosenst.)
Holtt. 26
var. hirta 26, 27
var. lanuginosa (v.A.v.R.)
Holtt. 27
var. ornamentalis
(Rosenst.) Holtt. 27
var. paleacea (Baker)
@iGhis22226
var. virescens (Hieron.)
Holtt. 27
hispida Mett. ex Kuhn 6, 18*,
19*, 22, 24, 349
hooglandii Holtt. 561
kajewski Copel. 22
koordersii Christ 24
laevigata Hook. 20
var. bracteata v.A.v.R. 21
laevissima Christ 4, 11
linearis (Burm. f.) Clarke 33
var. alternans (non Mett.)
Holtt. 35, 36
var. altissima Holtt. 36
var. ferruginea v.A.v.R. 34
var. inaequalis Rosenst. 36
var. montana Holtt. 36
var. normalis v.A.v.R. 33
var. pubigera v.A.v.R. 32
var. rigida v.A.v.R. 34
var. subferruginea Hieron.
ex Brause 34
var. tetraphylla Rosenst.
36
loheri Christ 6, 25
var. loheri 25
var. major Holtt. 5, 25
longissima BI. 5, 10*, 12*, 13,
17, 561
matthewii Holtt. 5, 17
3/9
(Gleichenia)
microphylla R.Br. 2, 3, 4, 7*,
8*) 9; 14
var. semivestita v.A.v.R. 7
milnei Baker 5, 22, 561
monticola Ridl. 24
norrisii Mett. 5, 15, 16*, 17
var. floccigera C.Chr. 15, 17
novoguineensis Brause 15
oceanica Kuhn 5, 22
opposita v.A.v.R. 32
ornamentalis Rosenst. 27
var. lanuginosa v.A.v.R. 27
paleacea (Copel.) Holtt. 4, 13
papuana Holtt. 14
var. membranacea Holtt.
14
parallela Ridl. 32
peltophora Copel. 2, 4, 6
var. peltophora 6
var. schizolepis C.Chr. ex
Holtt. 7
peninsularis Copel. 26
polypodioides (Thunb.) Smith
3
pseudoscandens v.A.v.R. 5,
23
pteridifolia Ridl. 35
pulchra (Copel.) Holtt. 5, 23
reflexipinnula C.Chr. 5, 20,
561
sordida Copel. 5, 15
squamosissima Copel. 9
subpectinata Christ 35
subulata v.A.v.R. 24
sumatrana Holtt. 15
truncata (Willd.) Spr. 5, 18%,
19*, 20, 22
var. bracteata (BI. ex
Hook. & Baker) Holtt.
20521522
var. celebica Holtt. 20, 22
var. involuta Holtt. 20, 22
var. plumaeformis (Presl)
Holtt. 20, 21
var. truncata 20, 21, 561
venosa (Copel.) Holtt. 5, 20
vestita BI. 6, 19*, 23*
var. paleacea Baker 26
volubilis sensu v.A.v.R. 13
volubilis Jungh. 4, 13, 14, 561
var. peninsularis Holtt. 13,
561
var. volubilis 13
vulcanica BI. 2, 4, 7*, 8*, 9,
10
warburgii Christ 34, 35
weatherbyi Fosb. 32
Gleicheniaceae iii, v, vii, X, Xi,
Xiii, Xiv, XV, 1-36, 38, 71,
339, 561
580
(Gleicheniaceae)
ser. Marginales 71
ser. Superficiales 71
Gleicheniastrum Presl 6
lowei Nakai 7
microphyllum Pres] 7
var. semivestitum Nakai 7
Gleicheniopsis |
Gleichenites gracilis Zigno 1,
20
Goniophlebium ii, vi, Xvi
Goniopteris 336, 338, 341, 387
appendiculata (Bl.) Presl 452
aspera Presl 512
asymmetrica Fée 452
barbata Fée 391
claiborniana Berry 334, 511
costata Brack. 421
dalhousiana Fée 533
kennedyi (F.v.M.) Bail. 512
lobbiana Fée 419
meniscioides Fée 387
prolifera (Retz.) Presl 387
repanda Fée 533
rigida Ridl. 484
rubrinervis (Mett.) Carr. 531
rudis Ridl. 501
simplicifolia (J.Sm. ex
Hook.) Carr.
var. vitiensis Carr. 513
Grammatopteridium ii, xvi
Grammitidaceae i, ii, ix, Xiii,
XIV, XVil
Grammitis ti, vi, xvii, 291, 336
blumeana Presl 542
Gymnocarpium ii, xix, 335
Gymnogramme 335, 336, 541
appendiculata Bl. 452, 523
aspidioides Bl. 542
aspidioides Hook. 541
aurita Hook. 345
borneensis Hook. 314
canescens (Bl.) BI. 497
macrotis Kunze 527
pteridiformis Cesati 263
rhizophylla Kaulf. 323
stegnogramma BI. 541
subtrifoliata Hook. 261
totta sensu Bl. 542
Gymnopteris Christ 276, 277,
278, 318
flagellifera Bedd. 325
inconstans Copel. 325
linnaeana Christ 325
naumannii Diels 327
repanda Christ 329
var. quoyana (Gaud.) Diels
327
subrepanda J.Sm. 326
subsimplex Fée 326
Gymnosphaera BI. 73, 76, 115
FLORA MALESIANA
(Gymnosphaera)
atropurpurea (Copel.) Copel.
118
biformis (Rosenst.) Copel.
118
bipinnatifida Copel. 151
burbidgei (Baker) Copel. 151
dinagatensis Copel. 151
glabra sensu Copel. 151
glabra Bl. 120
gracillima Copel. 123
hewittii Copel. 118
holttumii Copel. 146
hornei Copel. 121
kingii (Copel.) Copel. 121
melanoclada Copel. 121
melanorachis Copel. 121
mollis (Copel.) Copel. 151
obliqua (Copel.) Copel. 146
papuana (Ridl.) Copel. 155
pulchra Copel. 152
ramispina (Hook.) Copel. 117
recommutata Copel. 118
sarawakensis (C.Chr.) Copel.
152
schlechteri (Brause) Copel.
123
squamulata sensu J.Sm. ex
Hook. 118
squamulata B1. 152
subbipinnata Copel. 146
trichophora Copel. 151
vexans (Ces.) Copel. 120
Haplodictyum Presl xxi, 334,
335, 336, 507, 509, 511
bakeri (Harr.) Ching 516
bulusanicum Holtt. 515
canescens (BI.) Ching 497
heterophyllum Presl 498, 507,
515
majus Copel. 464, 507, 509
Helicogyratae 71
Helminthostachys ii, xv
Hemigramma iii, xx
latifolia auct. non Copel. 330
Hemionitis ii, xvii
prolifera Retz. 387
Hemipteris iii, xx
Hemitelia R.Br. 69, 71, 73, 77
alsophiliformis v.A.v.R. 111
alternans Hook. 145
arfakensis (Gepp) v.A.v.R.
barisanica v.A.v.R. 89
bicolor (Copel.) v.A.v.R. 108
capensis sensu Hook. 111
capensis (L. f.) R.Br. 61
caudata (J.Sm.) Mett. 110
caudiculata Rosenst. 106,
108
[ser. II, vol. 1°
(Hemitelia)
caudipinnula v.A.v.R. 89
confluens v.A.v.R. 102
crenulata Mett. 111
fallax v.A.v.R. 111
var. major v.A.v.R. 111
glaucophylla v.A.v.R. 111
hemichlamydea (Copel.)
v.A.v.R. 110
heterochlamydea v.A.v.R.
108
horrida 73
horridipes v.A.v.R. 114
javanica Pres] 110
junghuhniana (Kunze) Mett.
111
var. dissoluta Racib. 111
latebrosa (Wall.) Mett. 111,
115
var. paraphysata v.A.v.R.
115
latipinnula v.A.v.R. 105
ledermannii Brause 140
leptolepia v.A.v.R. 115
manilensis Pres] 110
merapiensis v.A.v.R. 111
montana v.A.v.R. 109
multiflora (Sm.) R.Br. 77
paraphysophora v.A.v.R. 114
perpunctulata v.A.v.R. 114
rudimentaris v.A.v.R. 115
salticola v.A.v.R. 114
singalanensis v.A.v.R. 102
subconfluens v.A.v.R. 102
sumatrana v.A.v.R. 114
tonglonensis v.A.v.R. 108
truncata sensu Christ 139
warthon (Copel.) v.A.v.R.
108
Heterogonium ili, xx, Xxi
Heteroneuron argutum Fée 329
cuspidata Presl 325
heteroclitum Fée 325
naumannii Kuhn 327
scalpturatum Fée 321
sinuosum Fée 329
Hicriopteris sensu Ching 11
Hicriopteris Presl 3, 27, 33
astrotricha Copel. 15
blotiana Ching 17
bullata Ching 13
clemensiae Copel. 14
glauca 3
laevissima Ching 11
longissima Ching 14
norrisii Ching 15
novoguineensis Copel. 15
paleacea Copel. 13
speciosa Presl 32
volubilis Ching 13
Histiopteris ti, xix
1981]
Index to scientific plant names 581
Holcosorus ii, Xvi
Holostachyum ti, xvi
Hugona Cav. ex Roem. 451
Humata ii, Xvili
Hydroglossum Willd. 39, 45
auriculatum Willd. 54
circinnatum (Burm. f.) Willd.
59
flexuosum (L.) Willd. 53
japonicum (Thunb.) Willd. 51
longifolium Willd. 59
pedatum (Burm. f.) Willd. 59
pinnatifidum Willd. 39, 47, 53
Hydropteridales viii
Hydropterideae vili
Hymenophyllaceae ii, iv, vii,
viii, iX, X, XIll, XV
Hymenophyllum ti, xv
ramosissimum Ham. ex
D.Don 183
Hypodematium Kunze ili, xx,
331, 333
Hypolepis ii, xix, 38
Hypopeltis marginifera Bory
386
propinquoides Bory 386
Isoetaceae 11, 63-64, 562
Isoetales 63
Isoetes L. ti, 63, 562
habbemensis Alston 62*, 64
hopei Croft 562
hystrix Durieu 64
lacustris L. 64
neoguineensis Baker 64
var. rheophila Croft 562
philippinensis Merr. & Perry
64
sp. 562
stevensii Croft 562
Isoetites 64
Isoloma J.Sm. ii, xix, 178, 198,
229, 230
angustum (Copel.) Tard.-Blot
232
auriculatum (v.A.v.R.)
Tagawa 231
dicksonioides (Christ) Tard.-
Blot 253
divergens (Hook. & Grev.)
J.Sm. 233
fuligineum (Copel.) Copel.
230
querinianum (Gaud.) Fée 230
induratum (Baker) Tard.-Blot
230
jamesonioides (Baker)
Tagawa 232
lanuginosum J.Sm. 253
lindsayae (Christ) Tard.-Blot
253
(Isoloma)
ovatum (J.Sm.) Pres] 231
pellaeiforme (Christ) Tard.-
Blot 232
schizolomae (v.A.v.R.)
Tagawa 230
walkerae (Hook.) Pres] 229
Kaulinia Nayar 291
Klukia 37
Kylikipteris 65
Lastrea sensu Copel. 414
Eastrea Bory? 335," 336;- 337,
338, 341, 386, 397
alta (Brause) Copel. 404
attenuata Brack. 408
beddomei (Baker) Bedd. 371
belensis Copel. 403
bipinnata Copel. 405
brassii (C.Chr.) Copel. 373
calcarata
var. ciliata Bedd. 375
calva (Copel.) Copel. 351
caudiculata Pres] 547
cavitensis Copel. 421
ciliata Hook. 375
confusa Copel. 539
conterminoides (C.Chr.)
Copel. 367
coriacea (Brause) Copel. 368
costulisora Copel. 409
crassa Copel. 408
crassifolia (Bl.) Moore 384
dayi (Bedd.) Bedd. 353
decurrens J.Sm. 354
dichrotrichoides (v.A.v.R.)
Copel. 482
dryopteroidea Copel. 158
dura (Copel.) Copel. 368
erubescens sensu Copel. 549
eugracilis Copel. 371
exigua J.Sm. 539
fairbankii Bedd. 377
falcatipinnula (Copel.)
Copel. 406
finisterrae (Brause) Copel.
432
flaccida (Bl.) Moore 352
flavo-virens (Rosenst.)
Copel. 353
foxii (Christ) Copel. 431
gracilescens Bedd. 359, 371
var. glanduligera Bedd. 373
gracilescens (Bl.) Moore 351
grammitoides (Christ) Copel.
373
gymnocarpa Copel. 542, 543
hirsutipes (Clarke) Bedd. 359,
360
immersa (Bl.) Moore 547
(Lastrea)
keysseriana (Rosenst.) Copel.
432
klossii Holtt. 361
leucolepis Pres] 349
ligulata Pres] 431
macgregorii (Baker) Ridl. 367
malaccensis Presl 545, 548
microchlamys de Vriese 449
mingendensis Gilli 433
mixta (Rosenst.) Copel. 457
nephrodioides Bedd. 384
nervosa (Fée) Copel. 539
notabilis (Brause) Copel. 402
novoguineensis (Brause)
Copel. 469
ophiura Copel. 409
petrophila Copel. 435
philippina Presl 431
platyptera (Copel.) Copel.
369
pyrrhorhachis sensu Copel.
347
quadriaurita (Christ) Copel.
404
regis Copel. 434
ridleyi Bedd. 361
robinsonii Ridl. 361
singalanensis (Baker) Bedd.
352
squamipes (Copel.) Copel.
364
subdimorpha (Copel.) Copel.
384
subnigra (Brause) Copel. 369
tenericaulis (Hook.) Moore
348
thelypteris
var. squamigera Bedd. 377
tylodes (Kunze) Moore 413
unidentata Bedd. 359
uraiensis (Rosenst.) Tagawa
352
varievestita (C.Chr.) Copel.
401
verrucosa Presl 547
viscosa (Baker) Bedd. 360
viscosa J.Sm. 360
wantotensis Copel. 410
wariensis Copel. 403
williamsii Copel. 450
Lastreopsis ili, xxi
Lecanopteris ii, xvi
Lemmaphyllum ii, xvi
Lepidocaulon ili, xx
Lepisorus Ii, xvi
Leptochilus Kaulf. ti, xvi, 255,
277, 318
sect. Bolbitis Christ 255
sect. Lomagramma_ Christ
255
582
(Leptochilus)
angustipinnus Hayata 321
celebicus (Baker) C.Chr. 523
cuneatus R.Bonap. 285
cuspidatus (Presl)
3255327)
var. crenatus Rosenst. 321
var. marginalis Rosenst.
327
var. quoyanus C.Chr. ex
v.A.v.R. 327
decurrens 330
diversifolius C.Chr. 326
exsculptus (Baker) C.Chr.
528
heteroclitus C.Chr. 325
var. eurybasis Christ 326
var. foxworthyi Christ 326
var. inconstans Christ 326
var. linnaeanus Christ 325
hydrophyllus Copel. 326
inconstans Christ 326
linnaeanus Fée 325
lomarioides Bl. 279
var. pteroides v.A.v.R. 283
malaccensis C.Chr. 326
modestus C.Chr. 325
naumannii C.Chr. 327
novoguineensis Brause 281
oligodictyus (Baker) C.Chr.
380
ovatus Copel. 330
perakensis (Bedd.) C.Chr.
281
reimersii Rensch 321
rivularis C.Chr. 328
scalpturatus C.Chr. 321
simplicifolius Holtt. 326
stolonifer Christ 330
sumatranus v.A.v.R. 326
x trifidus v.A.v.R. 319, 320*,
330
virens C.Chr. 321
zollingeri Fée 326
Leptogramma J.Sm. 337, 540
amabilis Tagawa 543
aurita (Hook.) Bedd. 345
celebica Ching 543
decursive-pinnata (van Hall)
J.Sm. 354
gymnocarpa (Copel.) Ching
$42
petiolata Ching 542
totta sensu Bedd. 542
Leptopteris ii, xv
Leucostegia ii, xviii
Lindsaea Dryander ii, v, vii,
xix, 177, 178, 179, 198
group A 200
group B 200, 201
group C 200, 201, 203
©@Chr-
FLORA MALESIANA
(Lindsaea)
group D 200, 202
group E 200, 202
group F 200, 202
group G 200, 203
sect. Paralindsaea 178
sect. Sambirania Tard.-Blot
229
subg. Lindsaea 204
sect. Davalliastrum 204
sect. Isoloma (J.Sm.)
Kramer 177, 199, 202,
230
sect. Lindsaea 178, 199,
202, 225
sect. Osmolindsaea
Kramens tla liss 199;
200, 227, 229
sect. Psammolindsaea
Kramer 199, 200, 229
sect. Schizoloma (Gaud.)
Kramer 177, 178, 199,
200, 204, 230
sect. Stenolindsaea Kramer
199, 202, 233
sect. Synaphlebium (J.Sm.)
Diels 199, 201, 204, 213,
216
sect. Temnolindsaea
Kramer 199, 201, 213,
Dilfer 233
sect. Tropidolindsaea
Kramen 17759199" 200,
229
subg. Odontoloma (Hook.)
Kramer 203, 204, 235
sect. Lindsaenium (Fée)
Kramer 199, 244, 564
sect. Odontoloma 199, 235,
244, 564
sect. Penna-arborea
Kramer 199, 203, 249
sect. Pseudolancea Kramer
199, 244
acutifolia Desv. 253
adiantifolia (Hook.) Copel.
249
adiantoides (Bl.) Kuhn 249
adiantoides J.Sm. 200, 215*,
229, 564
alpestris v.A.v.R. 249, 251
ambigens Cesati 218
amboynensis (Hook.) Mett.
ex Kuhn 187, 253
angusta Copel. 240
apiculata Kunze 228
apoensis Copel. 203, 240,
242*
arcuata Kunze 178, 223
azurea Christ 201, 223
var. mambae v.A.v.R. 223
[ser. Il, volar
(Lindsaea)
baker (C:Chr.)§@iehraw202.
235
var. bakeri 209*, 235
var. pycnophylla Kramer
235
bantamensis Bl. 237, 239
billardieri (Gaud.) Carruth.
ex Seemann 211
binervata C.Chr. 249
bipinnata Roxb. 253
blanda Mett. ex Kuhn 251
blumeana (Hook.) Kuhn 216
bonii Christ 206
borneensis Hook. ex Baker
202, 225, 254
boryana (Presl) Brause 239
bouillodii Christ 200, 204,
209*, 254, 563
brachypoda 178
brevipes Copel. 234
bullata v.A.v.R. 228
calomelanos Kunze 228
cambodgensis auct. 204
cambodgensis Christ 563
canaliculatipes v.A.v.R. 214
capillacea Christ 203, 238%,
243
carvifolia Kramer 203, 241,
242*, 564
ceramica v.A.v.R. 218
chienii Ching 178, 207, 208
chinensis (L.) Mett. ex Kuhn
182, 253
colobodes Kunze 234
commixta Tagawa 207
‘concinna’ 178
concinna J.Sm. 233
copelandi C.Chr. 206
cordata (Gaud.) Desv. 253
crassipes Rosenst. 228
crenulata Fée 234
crispa Baker 201, 217, 224
cultrata auct. 227, 228
cultrata (Willd.) Swartz 201,
202, 217, 222 map
var. concinna (J.Sm.)
Domin 233
var. lobbiana (Hook.)
Becc. 234
var. pallens Hook. 228
var. parvula Holtt. 228
var. plumula (Ridl.) Holtt.
228
var. securifolia (Presl)
Baker 233
var. varia Copel. 228
cultripinna Copel. 237, 239
cuneifolia Pres] 180, 253
cyathicola Copel. 249, 251
davallioides auct. 218
1981]
(Lindsaea)
davallioides Bl. 219
f. parallelogrammoides
v.A.v.R. 220
decomposita auct. 218
decomposita Willd. 178, 222
f. longipinnula” v.A.v.R.
218
f. minor v.A.v.R. 218
var. davallioides (Bl.)
Domin 219
decrescens Copel. 253
delicatula Christ 237
denhami (Hook.) Mett. ex
Kuhn 186, 253
diplosora v.A.v.R. 245
var. acrosora C.Chr. 245
divergens Hook. & Grev.
202, 230, 233
doryphora Kramer 178, 202,
226*, 227 map, 244, 253
ensifolia Swartz 177, 201,
204, 211
ssp. agatii (Brackenr.)
Kramer 211
ssp. coriacea (v.A.v.R.)
Kramer 178, 211, 212
ssp. ensifolia 178, 204, 211,
73 VAI)
var. heterophylla (Dryand.)
Benth. 210
erecta Mirbel 211
fissa Copel. 203, 238*, 243,
254
flabellulata Dryand. 206
var. gigantea Hook. 208
var. polymorpha (Hook. &
Grev.) Hook. 206
foersteri Rosenst. 237, 239
furcata Copel. 218
gigantea (Hook.) C.Chr. 208
glandulifera v.A.v.R. 203,
236, 241*, 252 map
gomphophylla Baker 200,
205*, 207
gracilis B1. 233
var. major Mett. ex Kuhn
234
gracillima Copel. 234
grandifolia J.Sm. 253
griffithiana Hook. 211
gueriniana (Gaud.) Desv. 202,
230, 231*, 564
guianensis (Aubl.) Dryand.
ssp. lanceastrum Kramer
224
havicei Copel. 240
hemiacroscopica Kramer 200,
204, 205*
heterophylla Dryand. 201,
205*, 210 map, 212
Index to scientific plant names
(Lindsaea)
heterophylla Prentice 210
hewittii Copel. 202, 224
hosei C.Chr. 253
humilis Kuhn 229
hymenophylloides auct. 243
hymenophylloides Bl. 237,
239
impressa Christ 224
indurata Baker 230
integra Holtt. 178, 201, 221*,
223
intermedia Hook. 222
var. minor Hook. 222
interrupta (Roxb.) Morton
563
jamesonioides Baker 202, 232
japonica (Baker) Diels 229
javanensis BI. 200, 208
kinabaluensis Holtt. 224
kingii Copel. 201, 213, 215*
kjellbergii C.Chr. 213
klotzschiana Moritz. 213
lancea auct. 244
lancea auct. spec. Asiat. 227
lancea (L.) Bedd. 225
lanceolata Labill. 211
lanuginosa Wall. ex Hook.
253
ledermannii Brause 235
linearis 178
lobata Poir. 202, 219
var. incisa Mett. ex Kuhn
219
lobbiana Hook. 234
loheriana Christ 228
longa Copel. 237, 239, 253,
254
longifolia Copel. 202, 219,
Dies
longipes C.Chr. & Tard.-Blot
206
longissima Christ 245
lowei hort. 253
lucida BI. 202, 231*, 233
ssp. brevipes (Copel.)
Kramer 234
ssp. lucida 234
lunulata v.A.v.R. 236
macraeana (Hook. & Arn.)
Copel. 178, 239, 253
malayensis Holtt. 201, 217
marginata Brause 249
var. falcata Brause 251
var. marginata 251
media R.Br. 201, 208, 209*
membranacea Kunze 211
merrillii Copel. 203, 241
ssp. merrillii 238*, 241
microstegia Copel. 203, 204,
238*, 246, 247*, 248
583
(Lindsaea)
modesta Kramer 202, 221*,
225
monocarpa Rosenst. 203, 245
monosora sensu Rosenst. 245
monosora Copel. 245
montana Copel. 206, 207
multisora v.A.v.R. 201, 213,
215*
napaea v.A.v.R. 201, 202, 217
natunae Baker 201, 214, 215*
nitens Bl. 222
‘nitida’ 178
nitida sensu Holtt. 222, 223
nitida Copel. 222
oblanceolata v.A.v.R. 178,
203, 236, 244
oblongifolia Reinw. ex Hook.
23H]
obscura Brause 202, 224
obtusa J.Sm. 178, 202, 218,
DIO; 22224
odorata Roxb. 178, 200, 227,
228
oligoptera Kunze 211
orbiculata (Lamk) Mett. ex
Kuhn 177, 200, 201, 204,
206, 207, 254
var. commixta (Tagawa)
Kramer 206, 207, 208,
254
var. gigantea (Hook.) Mett.
ex Kuhn 208
var. integra Mett. ex Kuhn
254
var. odontosorioides
Copel. 204, 206
var. orbiculata 205*, 206,
207
var. polymorpha (Hook. &
Grev.) v.A.v.R. 206
var. sumatrana_ Rosenst.
204
ovata J.Sm. 202, 231
papuana Copel. 201, 223
parallelogramma v.A.v.R.
178, 202, 220 map, 221*
parasitica (Roxb.) Hieron.
203, 227, 244, 253, 254
parishii Baker 253, 266, 269
‘pectinata’ 178
pectinata auct. 236
pectinata Bl. 237, 239
f. dimorpha Rosenst. 239
var. brevipinnula Rosenst.
246
pectinata (Bory) Thw. 237
pellaeiformis Christ 202, 231
pentaphylla Hook. 211
philippinensis Kramer 202,
232
584 FLORA MALESIANA [ser. II, vol. 1°
(Lindsaea) (Lindsaea) (Lindsaea)
pinnata (Cav.) Mett. ex roemeriana’ Rosenst. 203, vieillardii 178
Kuhn 193, 253 209*, 252 viridis 178
var. bipinnata Mett. ex
Kuhn 191, 253
plumula Ridl. 228
polyctena Kramer 201, 214,
215*, 216 map, 564
polymorpha Wall. ex Hook.
& Grev. 206
portoricensis 178
prolongata 178
propria v.A.v.R. 234
pteroides Desv. 211
pulchella (J.Sm.) Mett. ex
Kuhn 203, 249, 250*, 251
map
var. blanda (Mett. ex
Kuhn) Kramer 178, 249,
250*, 251 map
var. falcata (Brause)
Kramer 249, 250, 251
map
var. lomatosora Kramer
249, 250*, 251 map
var. pulchella 249, 250*,
251 map
ramosii Copel. 201, 219
recurvata (Hook.) Hook. 222
regularis Rosenst. 204, 246,
247*
repens (Bory) Thw. 178, 203,
239; 237,238". 241, 253,
564
f. angusta (Copel.) Kramer
240
f. truncatiloba Rosenst.
237, 239
var. delicatula
Kramer 237, 239
var. hemiptera
v.A.v.R. 237, 239
var. intermedia Christ 237,
239
var. laciniata Mett. ex
Kuhn 236
var. pectinata (Bl.) Mett.
ex Kuhn 238*, 239,
240
var. pseudohemiptera
v.A.v.R. 239, 240
var. sessilis (Copel.)
Kramer 178, 237, 238%,
239, 254
var. submarginalis Kramer
238*, 239, 240
retusa (Cav.) Merr. 180, 253
rhombifoliolata v.A.v.R. 251
rigida J.Sm. 203, 204, 244,
245, 246, 254, 564
f. acutata v.A.v.R. 245
(Christ)
(Bory)
rosenstockii Brause 203, 204,
238*, 248, 564
roxasii Copel. 237
sagincola Wagner & Grether
237), 239
sarasinorum Christ 222
sarawakensis Kramer 203,
245, 564
scandens Hook. 244
var. terrestris Holtt. 178,
227
schizoloma Ettingsh. 211
schlechteri Brause 235
schultzei Brause 218, 219
securifolia Presl ex Gold-
mann 233
seemannii J.Sm. 229
sepikensis Brause 245
sessilis Copel. 237, 239
sinuato-crenata v.A.v.R. 218,
219
striata Bl. 254
subalpina auct. 217
subalpina v.A.v.R. 202, 218,
DOA
sublobata Kunze 211
subsemilunularis
236
subtripinnata Copel. 208
tenera auct. 204
tenera Dryand. 207, 254
var. commixta (Tagawa)
Iwats. 207
var. gigantea
Holtt. 208
tenuifolia auct. 214
tenuifolia Bl. 201, 216 map,
564
tetragona Kramer 201, 214,
563
var. brassiana Kramer 214
var. tetragona 214
trapeziformis auct. 227
trapeziformis Dryand. 198,
225
trapezoidea Copel. 222
trichophylla Copel. 235
tricrenata Baker 254
trilobata Baker 253
trimarginata C.Chr. 229
tripartita Bl. 218, 219
triplosora v.A.v.R. 245
triquetra (Baker) Christ 216
tropidorachis v.A.v.R. 229
variabilis Hook. & Walker
Arnott 210
versteegii (Christ) v.A.v.R.
203, 204, 238*, 248, 564
v.A.v.R.
(Hook.)
vittata Zoll. & Mor. 253
vrieseana Rosenst. 231
walkerae Hook. 200, 229, 230
map
werneri Rosenst. 203, 238%,
252 map
wollastonii v.A.v.R. 252
Lindsaea group ii, ix, X, Xi, XIV,
xix, 177-254, 563
Lindsaenium Fée 198, 244
rigidum (J.Sm.) Fée 245
Lindsaya (sphalm.) 198
Lindsaynium Fée 198, 244
Lomagramma sensu Copel. 275
Lomagramma J.Sm. ii, xx, 255,
256, 257, 2585325952765
278
angustipinna Copel. 278, 279,
281
articulata Copel. 276
bipinnata Copel. 276
borneensis (erron.) 285
brassii Holtt. 277, 279, 287,
289
brooksii Copel. 279, 285
copelandii Holtt. 279, 287,
289
cordata Copel. 283, 285
cordipinna Holtt. 277, 278
guianensis (Aubl.) Ching 257,
289
leucolepis Holtt. 279, 285
lomarioides (Bl.) J.Sm. 278,
279, 289
matthewii (Ching) Holtt. 289
melanolepis v.A.v.R. 278,
279, 282250 2862875
288*, 289
merrillii Holtt. 278, 279, 281
novoguineensis (Brause)
C.Chr:. 2783279 5aest
282*
pedicellata Copel. 283
perakensis Bedd. 278, 279,
280*, 281, 283*
polyphylla Brack. 255, 277
praestantissimum (Bory)
Gries. 289
pteroides J.Sm. 277, 278, 279,
281, 283*, 289
var. negrosensis Copel. 287
var. subcoriacea Copel.
283
sinuata C.Chr. 277, 278, 279,
282*, 284*, 285, 289
f. papuana C.Chr. 279, 285,
287
sorbifolia (Willd.) Ching 289
1981]
(Lomagramma)
sp. 285
subcoriacea Copel. 283
sumatrana v.A.v.R. 277, 278,
279, 283*, 285
Index to scientific plant names
Lycopodium ti
Lycopsida ii
Lygodiaceae 39
Lygodictyon J.Sm. 45
Lygodium Swartz ii, v, xv, 37,
wilkesiana (Brack.) Copel.
289
Lomaria 266
aculeata Bl. 266, 267, 269
gracilis Bl. 269
ludens Fée 275
polymorpha Zoll. & Mor. 269
spectabilis Kunze 262
variabilis sensu BI. 262
Lomariopsidaceae Alston 255,
256
Lomariopsis Fée ii, xx, 255,
256) 25715 2585 259; 261,
263, 265, 266, 267, 276,
PINT, Pass)
sect. Eulomariopsis Mett. ex
Kuhn 259
sect. Stenochlaena Mett. 267
brackenridgei Carr. 261
cochinchinensis Fée 259, 262
hiigelii Presl 263
intermedia (Copel.) Holtt.
260*, 261, 317*
kingii (Copel.) Holtt. 261,
262226355177
leptocarpa sensu Holtt. 262
leptocarpa Fée 267, 270
lineata (Presl) Holtt. 259, 261,
262, 270
ludens Fée 267
oleandrifolia (Brack.) Mett.
261
palustris (Hook.) Mett. 259
papyracea Copel. 262
raciborskii (C.Chr.) Holtt.
262
setchellii (Maxon) Holtt. 261
smithii Fée 262
sorbifolia Christ 257, 259,
261, 266
spectabilis (Kunze) Mett.
260*, 262
spinescens Fée 267
subtrifoliata (Copel.) Holtt.
261, 287
variabilis (Willd.) Fée 256, 259
Lomariopsis group ii, vi, ix, x,
Xxli, x1x, 255-330, 564
Lophidium Rich. 39, 40
Lophosoria 562
Lophosoriaceae 562
Loxogramme Il, xvi
Loxoscaphe li, xvii
Luerssenia ili, xxi
Lycopodiaceae ii
Lycopodiales ti
38, 39, 40, 44, 561
auriculatum (Willd.) Alston
45, 46, 54, 55*, 57
basilanicum Christ 59
borneense v.A.v.R. 45, 46,
AW, SRY Sif
f. samarindae v.A.v.R. 55
chaerophylloides Desv. 51
circinnatum (Burm. f.) Sw.
39, 46, 47*, 58, 59
var. cristatum v.A.v.R. 59
var. monstruosum v.A.v.R.
59
var. semihastatum Fosb.
54
var. trifurcatum Christ 54
cochinchinense Desv. 51
derivatum v.A.v.R. 59
dichotomum sensu Bedd. 59
dichotomum (Cav.) Sw. 59
digitatum Pres] 59
dimorphum Copel. 45, 46,
52*, 54
dissectum Desv. 51
flexuosum sensu Gaud. 54
flexuosum sensu Holtt. 51
flexuosum sensu Prantl 54
flexuosum (L.) Sw. 46, 51,
5255 535562
var. setulosum Tard. &
@Chi2s3
japonicum (Thunb.) Sw. 37,
39, 46, 50*, 51
f. elongata v.A.v.R. 51
f. microstachya (Desv.)
Marda cae: Chri
kingii Copel. 51
longifolium (Willd.) Sw. 46,
A]z S659; SOLE SOL
matthewii Copel. 61
mearnsii Copel. 51
merrillii Copel. 46, 60*, 61
microphyllum Link 51
microphyllum (Cav.) R.Br. 37,
39, 46, 47*, 48*, 49*, 61
microstachyum Desv. 51
moszkowskii Brause 61
novoguineense Rosenst. 54
pedatum (Burm. f.) Sw. 59
pinnatifidum sensu Prantl 47
pinnatifidum sensu Racib. 51
pinnatifidum Sw. 53
polystachyum Wall. ex
Moore 37, 45, 46, 47*,
50*
pubescens Kaulf. 51
585
(Lygodium)
reticulatum Schkuhr 46, 61
salicifolium Pres] 37, 46, 51,
S37 0
scandens (L.) Sw. 39, 47
var. intermedium Ces. 47
var. microphyllum (Cav.)
Luerss. 47
semibipinnatum R.Br. 53
semihastatum sensu v.A.v.R.
55
semihastatum Desv. 54
serrulatum Bl. 53
tenue Bl. 51
teysmannii v.A.v.R. 59
trifurcatum sensu v.A.v.R.
54
trifurcatum Baker 45, 46, 54
versteegii Christ 46, 61, 562
Macroglossum ii, xv
Macrothelypteris sensu Pichi
Sermolli 343
Macrothelypteris (H.Ito) Ching
324, 333, 339, 340, 345,
347, 374
leucolepis (Presl) Ching 349
multiseta (Baker) Ching 348,
349
oligophlebia (Baker) Ching
347
ornata 348
paludosa (Bl.) Love & Love
347
polypodioides (Hook.) Holtt.
344*, 345, 348, 349
setigera (BI.) Ching 344*, 348,
349
torresiana (Gaud.) Ching 335,
344*, 345, 348, 349
uraiensis (Rosenst.) Love &
Love 352
viridifrons 348
Marattia ll, xv
Marattiaceae Ii, X, Xll, XV
Marattiales ii, vil
Marsilea iii, Xx1
Marsileaceae ili, vill, ix, Xii, XXi
Matonia ii, iv, xv, 71
Matoniaceae il, X, Xilil, xv, 38
Meniscium Schreb. 335, 337,
338, 531, 534
sect. Ampelopteris Iwats. 387
beccarianum Cesati 513
cumingii Fée 534
cuspidatum Bl. 536, 537
fragile (Z. & M.) Kunze 512
hosei Baker 526
var. sumbensis v.A.v.R.
523
kennedyi F.v.M. 512
586
FLORA MALESIANA
[ser. II, vol. 1°
(Meniscium)
lineatum (BI.) Ktze 512
liukiuense Christ 537
parishii Bedd. 535
proliferum sensu Hook. 526
proliferum (Retz.) Sw. 387
pseudoarfakianum (Hosok.)
Hosok. 454
salicifolium Wall. ex Hook.
535
sp. Moritzi 512
stenophyllum Baker 528, 529
thwaitesii Hook. 535
triphyllum Sw. 534
var. parishii Bedd. 535
Menisorus Alston 340, 374
Meringium xv
Merinthosorus il, Xvi
Mertensia Roth 3
Mertensia Willd. 3, 18, 27
arachnoides Hassk. 13
crassifolia Pres] 34
laevigata Willd. 20
lessonii A.Rich. 31
plumdeformis Presl 21
pteridifolia Presl 33
spissa (non Fée)
var. pubigera Nakai 32
truncata Willd. 3, 20, 21
Mesochlaena R.Br. 436, 437
asplenioides J.Sm. 449
javanica R.Br. ex Mett. 437,
449
larutensis
489
var. borneensis
489
polycarpa
449
sumatrensis v.A.v.R. 489
talamauensis v.A.v.R. 449
toppingii Copel. 449
Mesoneuron Ching 377, 378
(Bedd.) v.A.v.R.
v.A.v.R.
(BI.) Bedd. 437,
attenuatum (O.Ktze) Ching
377, 549
chlamydophorum (C.Chr.)
Ching 384
crassifolium (Bl.) Ching 384
echinatum (Mett.) Ching 379
hallieri (Christ) Ching. 379
paleatum (Copel.) Ching 380
persquamiferum (v.A.v.R.)
Ching 379
teuscheri (v.A.v.R.) Ching
381
trichopodum (C.Chr.) Ching
380
wantotense sensu Holtt. 403
Mesophlebion Holtt. 335, 340,
BAIENS 425s 528 S771.) 386,
397, 398
(Mesophlebion)
subg. Plesioneuron Holtt.
377, 378, 397
arenicola Holtt. 378, 379, 381,
383
auriculiferum (v.A.v.R.)
Holtt. 379, 383
beccarianum (Ces.) Holtt.
376*, 378, 379, 381, 383,
385
caroli Holtt. 378, 380
chlamydophorum (C.Chr.)
Holtt. 341, 378, 379, 383,
384
crassifolium (BI.) Holtt. 332*,
334, 378, 379, 382, 384
dulitense Holtt. 379, 382
echinatum (Mett.) Holtt. 378,
379
endertii (C.Chr.) Holtt. 376*,
377, 379, 381
falcatilobum Holtt. 378, 379,
382
var. apiculatum Holtt. 382
var. falcatilobum 382
hallieri (Christ) Holtt. 378,
379
motleyanum (Hook.) Holtt.
377, 378, 379, 381, 385
oligodictyon (Baker) Holtt.
376*, 377, 378, 380
pallescens (Brause)
383
persquamiferum
Holtt. 378, 379
rufescens Holtt. 379, 383
teuscheri (v.A.v.R.) Holtt.
378, 379, 381, 383
trichopodum (C.Chr.) Holtt.
376*, 378, 379, 380
vinosicarpum (v.A.v.R.)
Holtt. 381
Mesosorus Hassk. 11, 18, 27
arachnoides Hassk. 13
Metathelypteris (H.Ito) Ching
333, 334, 339, 342, 350
dayi (Bedd.) Holtt. 332%,
3505351352
decipiens (Clarke) Ching 351
flaccida (BI.) Ching 350*, 351
gracilescens (BI.) Ching 350*,
351, 352
singalanensis (Baker) Ching
350*, 351, 352
var. singalanensis 352
var. surbeckii Holtt. 351,
352
uraiensis (Rosenst.)
350*, 351, 352
Metaxya 65, 72, 291, 562
Metaxyaceae 562
Holtt.
(v.A.v.R.)
Ching
Metaxyineae 562
Mezonevron Desf., non Unger
378
Microlepia ii, iv, xix, 169, 185,
256
sect. Tapeinidium Presl 184
biflora (Kaulf.) Mett. 182
biserrata (Bl.) Pres] 194
chinensis (L.) Mett. 182
denhami (Hook.) Moore 186
gracilis (Bl.) J.Sm. 190
philippinensis (Harr.) Copel.
191
pinnata (Cav.) J.Sm. 193
strigosa (Thunb.) Pres] 162
tenuifolia (Lamk) Mett. 182
tenuis Brack. 186
Microschizaea Reed 40
fistulosa Reed 44
hallieri Reed 44
malaccana Reed 44
Microsorium li, Xvi
pteropus (BI.) Ching 291
punctatum (L.) Copel. 291
Microstaphyla Pres] 255
Mohria 37, 39
caffrorum (L.) Desv. 61
Monachosorum ii, xviii
Monogramma iii, XX1
Myrmecophila ti, xvi
Nannothelypteris Holtt. 340,
343, 537
camarinensis Holtt. 538, 539*
inaequilobata Holtt. 538,
539*, 540
nervosa sensu Holtt. 540
nervosa (Fée) Holtt. 538,
539*
aoristisora (Harr.) Holtt. 538
philippina (Presl) Holtt. 538,
539*
Nathorstiana 63, 64
Nematopteris tl, xvii
Neocheiropteris ti, Xvi
Nephelea 562
Nephrodium Schott 258, 335,
336; 337; 3383935459437.
550
abruptum sensu Hook. 461
abruptum Bedd. 462
acrostichoides Desv. 456
alatellum Christ 451
amboinense
var. subglandulosum
Baker 454
angustifolium Presl 481, 557
appendiculatum (Bl.) Racib.
452
aridum (D.Don) J.Sm. 555
arutense Bedd. 489
1981]
(Nephrodium)
attenuatum Baker 549
bakeri Harr. 516
basilare Pres] 461
beccarianum Cesati 383
beddomei Baker 371
brachyodes sensu Baker 382
calcaratum (BI.) Hook. 360,
374
callosum (BI.) Keys. 419
caudiculatum J.Sm. 423, 461
conioneuron Fée 545, 548
creaghii Baker 353
crinipes sensu Ridl. 380
cucullatum (BI.) Baker 478
cuspidatum Pres|l 325
debile Baker 522
decursive-pinnatum (van
Hall) Hook. 354
didymosorum Parish ex
Bedd. 560
diversilobum Pres] 452, 517
eminens Baker 449
eusorum (Thw.) Bedd. 429
extensum (BI.) Moore 548
falcilobum Hook. 374
fasciculatum (Fourn.) Baker
367
ferox (Bl.) Moore 391
flaccidum (BI.) Hook. 352
glandulosum sensu Bedd. 527
glandulosum sensu J.Sm. 512
glandulosum (BI.) J.Sm. 522
glaucostipes Bedd. 471
gracilescens (Bl.) Hook. 351
var. hirsutipes Clarke 359
griseum Baker 375
gymnopodum Baker 361
haenkeanum sensu Baker
478, 479
haenkeanum Presl 478, 502
heterocarpum_ (BI.) Moore
457
hirsutum Presl 455
hispidulum sensu Christ 466
impressum Desv. 548
indicum sensu Ridl. 485
insculptum Desv. 478
invisum (Forst. f.) Carr. 502
javanicum (Mett.) Hook. 437,
449
larutense Bedd. 437
latifolium Pres] 512, 513, 526
latipinna Hook. 558, 559
lineatum sensu Bedd. 522
lineatum sensu Hook. 528
lineatum (BI.) Pres] 512
longipes (Bl.) Moore 424
luerssenii Harr. 431
macgregorii Baker 367
microchlamys Baker 449
(Nephrodium)
molle (Sw.) R.Br. 557
var. amboinense Bedd. 558
var. didymosorum Bedd.
560
var. major Bedd. 431, 556,
558
motleyanum Hook. 381
multijugum Baker 420
multilineatum Bedd. 461
multisetum Baker 349
nitidulum Presl 430, 461
obscurum (BI.) T.Moore 556
ochthodes
var. tylodes Bedd. 414
oosorum Baker 500
oreopteris Fée 545
papilio Hope 556
patens sensu Hook. 555
pennigerum Hook. 461, 462
var. malayense Bedd. 485
var. multilineatum Clarke
461
perakense (Bedd.) Baker 449
philippinense Baker 460, 461
pilosiusculum Racib. 522
polycarpum (BI.) Keys. 448
polytrichum Baker 380
prolixum sensu Baker 414
var. tylodes 414
prolixum Clarke 413
propinquum R.Br. 386
pteroides sensu Bedd. 545
pubescens Brack. 457
pubirachis Baker 365
rodigasianum T.Moore 426
sagittifolium (Bl.) Moore 487
sakayense Zeiller 392, 393
savaiense Baker 404
serratum (Cav.) Pres] 478
setigerum sensu. Hook. &
Baker 348
simulans Baker 367, 520
singalanense Baker 352
smithianum Pres] 456, 557
sophoroides (Thunb.) Desv.
560
sorbifolium
289
squamulosum Hook. f. 377
stipellatum (Bl.) Moore 492
tectum Bedd. 560
(Willd.) Pres!
tenericaula (Hook.) Hook.
348
terminans Hook. 545
thelypteris
var. squamulosum Hook.
377
truncatum sensu Racib. 429
tuberculatum Cesati 407
tylodes (Kunze) Hope 414
Index to scientific plant names 587
(Nephrodium)
unitum sensu R.Br. et auct.
386
urophyllum sensu Racib. 495
urophyllum (Mett.) Keys. 533
var. pinwillii Bedd. 533
viscosum Baker 360
vulcanicum Baker 351
Nephrolepis iii, xx, 257, 258,
263, 335
acutifolia (Desv.) Christ 253
dicksonioides Christ 162, 253
schizolomae v.A.v.R. 230
Nephrolepis group Ill, ix, Xi, Xil,
ROK
Neurocallis Fée 277, 278
sect. Cheilolepton Moore 277
praestantissimum (Bory) Fée
289
Notholaena ii, xvii, 255
pteridiformis (Ces.)
263
Baker
Odontoloma J.Sm. 198, 235
adiantoides (Bl.) Pres] 249
blumeanum (Hook.) Mett.
216
boryana (erron.) 237
boryanum (Presl) J.Sm. 237
hymenophylloides (BI.) J.Sm.
237
pulchellum J.Sm. 249
repens sensu J.Sm. 241
repens (Bory) Pres] 237
tenuifolium (BI.) J.Sm. 216
triquetra (erron.) 216
triquetrum (Baker) J.Sm. 216
Odontopteris Bernh. 39, 44
Odontosoria Fée 177, 178, 179
biflora (Kaulf.) C.Chr. 182
chinensis (L.) J.Sm. 182
var. divaricata Christ 183
var. tenuifolia (Lamk)
Matsum. 183
chusana (L.) Masam. 182
var. tenuifolia (Lamk)
Masam. 183
decipiens (Ces.) Christ 180
lindsayae v.A.v.R. 180
retusa (Cav.) J.Sm. 180
tenera Ridl. 252
tenuifolia (Lamk) J.Sm. 182
tsoongii Ching 182
versteegii Christ 248
Oleandra ii, iv, xx, 258, 335
Oleandropsis ti, xvi
Olfersia Pres] 289, 291
aculeata Bl. 269
angulata Pres] 297
blumeana Pres] 303, 311
callifolia Pres] 307
588
(Olfersia)
corcovadensis 291
gracilis Bl. 269
lineata Pres] 262
Olfersia Raddi 291
Oligocarpia 1
Oligolepideae Fée 291
Onychium ii, xvii
Ophioglossaceae ii, ix, xX, Xi,
XV
Ophioglossales ii, vil, viii
Ophioglossum ii, xv
circinnatum Burm. f. 59
filiforme Roxb. 47
flexuosum L. 39, 53
furcatum Roxb. 59
japonicum Thunb. 51
pedatum Burm. f. 59
scandens L. 39, 53
Oreogrammites ii, xvii
Ormoloma 177
Orthiopteris ti, xviii
kingii (Bedd.) Holtt. 162
Osmunda ii, xv, 71
dichotoma Spr. 41
Osmundaceae ii, vii, X, Xili, Xv
Paesia ii, xix, 38
Palmifilix alba Rumph. 131
Paragramma ii, xv
Paraleptochilus ovatus Copel.
330
Parapolystichum Ching 331
Parasorus ii, xviii
Parathelypteris (H.Ito) Ching
340, 342, 343, 370
FLORA MALESIANA
(Parathelypteris)
pectiniformis (C.Chr.) Ching
363
simozawae Ching 360
simulata (Davenp.) Nieuwl.
370
subimmersa Ching 547
viscosa (Baker) Ching 360
Pecopteris 1
Pellaea ii, xvii
pteroides Prantl 61
Pelletiera 37
Peltapteris Link 255, 259
Peranema Ii, xix
Phanerosorus li, xv
Phegopteris sensu Tagawa 343
Phegopteris Fée 333, 336, 337,
338, 339, 342, 353
sect. Lastrella H.Ito 343
sect. Meniscium 534
acanthocarpa (Copel.)
v.A.v.R. 532
acromanes (Christ) v.A.v.R.
452
aoristisora (Harr.) v.A.v.R.
538
arfakiana (Baker) v.A.v.R.
484
armata (Rosenst.) v.A.v.R.
392
auriculata J.Sm. 412
aurita (Hook.) J.Sm. 345
barbata (Fée) Mett. 391
beccariana (Ces.) v.A.v.R.
$13
borneensis (Hook.) v.A.v.R.
[ser. II, vol. 1°
(Phegopteris)
holophylla (C.Chr.) v.A.v.R.
526
hosei (Baker) v.A.v.R. 526
var. sumbensis v.A.v.R.
523
imponens (Ces.) v.A.v.R. 391
luxurians (Kunze) Mett. 387
mamberamensis v.A.v.R. 549
melanophlebia (Copel.)
v.A.v.R. 514
merrillii (Christ) v.A.v.R. 529
moussetii (Rosenst.) v.A.v.R.
346
nervosa Fée 538, 539
oblanceolata (Copel.)
v.A.v.R. 513
obtusifolia (Rosenst.)
v.A.v.R. 503
oppositipinna v.A.v.R. 346
pentaphylla (Rosenst.)
v.A.v.R. 514
perrigida v.A.v.R. 394
philippinensis Mett. 461
polypodioides Fée 353
prolifera (Retz.) Kuhn 387
pseudoarfakiana Hosok. 454
ridleyana v.A.v.R. 501
rigida (Hook. & Grev.) Mett.
484
rubicunda v.A.v.R. 536
rubida (Hook.) v.A.v.R. 532
rubrinervis Mett. 531
rutteniana v.A.v.R. 513
salicifolia (Wall. ex Hook.)
Mett. 535
sect. Melanostipes Ching 354
angulariloba (Ching) Ching
360
angustifrons (Miq.) Ching 373
beddomei (Baker) Ching 370,
S7i 32%
var. beddomei 371, 373
var. brassii (C.Chr.) Holtt.
371
var. eugracilis
Holtt. 371
cystopteroides (Eaton) Ching
370, 373
glanduligera (Kunze) Ching
370, 373
grammitoides (Christ) Holtt.
373
hirsutipes (Clarke) Ching 359
immersa (BI.) Ching 547
indochinensis (Christ) Ching
359
nipponica (Fr. & Sav.) Ching
371
noveboracensis (L.) Ching
370, 371
(Copel.)
528
canescens (BI.) Mett. 497
var. degenera (Christ)
v.A.v.R. 522
ceramica v.A.v.R. 549
chamaeotaria (Christ)
v.A.v.R. 524
cheilanthoides (Baker)
v.A.v.R. 349
connectilis 354
cuspidata (Bl.) Mett. 536
debilis Mett. 448
decursive-pinnata (van Hall)
Fée 343, 354
diversiloba (Presl) v.A.v.R.
452
engleriana (Brause) v.A.v.R.
366
ferox (Bl.) Mett. 391
firmula (Baker) v.A.v.R. 519
granulosa (Presl) v.A.v.R.
524
gymnocarpa (Copel.)
v.A.v.R. 542
hexagonoptera 354
simplicifolia (J.Sm.) v.A.v.R.
490
smithii v.A.v.R. 371
spenceri (Christ) v.A.v.R. 491
stegnogramma Mett. 541
var. meniscioides v.A.v.R.
542
stenophylla (Bak.) v.A.v.R.
528
totta sensu Mett. 542
var. subcalcarata v.A.v.R.
$42, 543
triphylla (Sw.) Mett. 534
urophylla Mett. 533
wollastonii v.A.v.R. 484
xiphioides (Christ) v.A.v.R.
525
Photinopteris ii, xvi
Physematium Pres| 336
philippinum Pres! 539
Pityrogramma Ii, vi, XVii
Plagiogyria iii, vii, xx, 340
Plagiogyria group iii, x, xiii, Xx
Platycerium il, xvi
Platyzoma 1, 2
1981]
Pleocnemia ili, xx, 257, 334,
336, S11
cumingiana Presl 158
excellens (BI.) v.A.v.R. 462
heterophylla (Presl) v.A.v.R.
515
Plesioneuron Holtt. 335, 340,
342, 378, 389, 397
altum (Brause) Holtt. 398,
404
angiensis Holtt. 399, 406
archboldii (Copel.) Holtt. 406
attenuatum (Brack.) Holtt.
399, 407, 409
costulisorum (Copel.) Holtt.
399, 409, 410, 411
crassum (Copel.) Holtt. 399,
408
croftii Holtt. 399, 411
ctenolobum Holtt. 408
cystodioides Holtt. 398, 405
belense (Copel.) Holtt. 398,
403
bipinnatum (Copel.) Holtt.
398, 405
doctersii Holtt. 399, 410
dryas Holtt. 399, 411
dryopteroideum (Brause)
Holtt. 398, 401, 402
var. buruense Holtt. 401
var. dryopteroideum 401
var. pilosum Holtt. 401
falcatipinnulum (Copel.)
Holtt. 399, 406
fuchsii Holtt. 398, 400
fulgens (Brause) Holtt. 332*,
397, 398, 403
irayense (Copel.) Holtt. 495
kostermansii Holtt. 399, 406
kundipense Holtt. 399, 410
marattioides (Alston) Holtt.
396*, 398, 402
medusella Holtt. 398, 400
notabile (Brause) Holtt. 398,
402
ophiura (Copel.) Holtt. 397,
399, 409, 410
phanerophlebium (Baker)
Holtt. 406
platylobum Holtt. 399, 408
pullei Holtt. 398, 399
quadriquetrum (v.A.v.R.)
Holtt. 398, 406
rigidilobum Holtt. 399, 410
royenii Holtt. 398, 404
sandsii Holtt. 399, 407
savaiense
396*, 398, 404
septempedale (Alston) Holtt.
398, 402
stenura Holtt. 399, 408
(Baker) Holtt.
Index to scientific plant names
(Plesioneuron)
stenura Holtt. 399, 408
subterminale Holtt. 398, 405,
406
tuberculatum (Ces.) Holtt.
396*, 397, 399, 404, 405,
407
varievestitum (C.Chr.) Holtt.
398, 401
var. obtusipilum Holtt. 401,
402
var. varievestitum 401
wantotense (Copel.) Holtt.
399, 410
wariense (Copel.) Holtt. 398,
403
woodlarkense (Copel.) Holtt.
398, 400
Pleuromeia 64
Pneumatopteris sensu Holtt.
413
Pneumatopteris Nakai 333, 334,
338, 340, 343, 413, 414,
437, 439, 461
angusticauda Holtt. 417, 422
auctipinna Holtt. 417, 425
basicurtata Holtt. 417, 424
boridensis Holtt. 418, 434
brooksii (Copel.) Holtt. 417,
423
callosa (BI.) Nakai 415, 416*,
419, 420
caudata (Holtt.) Holtt. 418,
433
cheesmaniae Holtt. 417, 422
christelloides Holtt. 429
costata (Brack.) Holtt. 414,
415, 417, 418, 420, 421,
432, 439
var. hispida Holtt. 421
deficiens Holtt. 418, 433
dicranogramma (v.A.v.R.)
Holtt. 415, 419
eburnea Holtt. 418, 434
ecallosa (Holtt.) Holtt. 417,
425
egenolfioides Holtt. 417, 421
excisa (Holtt.) Holtt. 418, 434
finisterrae (Brause) Holtt.
418, 432
glaberrima sensu Holtt. 430
glabra (Copel.) Holtt. 415,
418
imbricata Holtt. 418, 435
incisa Holtt. 417, 427
inclusa (Copel.) Holtt. 417,
423
japenensis Holtt. 417, 426
jermyi Holtt. 417, 422, 426
kerintjiensis Holtt. 415, 418,
430
589
(Pneumatopteris)
keysseriana (Rosenst.) Holtt.
416*, 418, 432, 433
laevis (Mett.) Holtt. 416*,
417, 424
laticuneata Holtt. 428
latisquamata Holtt. 415, 421,
435
lawakii Holtt. 418, 430
ligulata (Presl) Holtt. 418, 431
lithophila Holtt. 417, 422
longipes (BI.) Holtt. 417, 423,
425, 427
michaelis Holtt. 418, 430
microauriculata Holtt. 417,
425
microloncha (Christ) Holtt.
417, 423
micropaleata Holtt. 418, 428
mingendensis (Gilli) Holtt. 418,
433
nephrolepioides (C.Chr.)
Holtt. 418, 431
nitidula (Presl) Holtt. 418,
430, 465
obliqua Holtt. 417, 424
papuana Holtt. 417, 426
patentipinna Holtt. 417, 422
pennigera (G.Forst.) Holtt.
415, 421
pergamacea Holtt. 417, 427
petrophila (Copel.) Holtt.
418, 435
psilophylla Holtt. 417, 427
regis (Copel.) Holtt. 418, 434
rodigasiana (T.Moore) Holtt.
417, 426
sandwicensis (Brack.) Holtt.
541
sibelana Holtt. 415, 419
sogerensis (Gepp) Holtt. 417,
428, 436
subappendiculata (Copel.)
Holtt. 415, 420
sumbawensis (C.Chr.) Holtt.
418, 432
superba (Brause) Holtt. 415,
420
tobaica Holtt. 417, 427
truncata (Poir.) Holtt. 415,
416*, 418, 427, 429, 430
versteeghii Holtt. 418, 432
walkeri Holtt. 418, 435
Poecilopteris contaminans
Moore 321
stenophylla Kurz ex T. & B.
325
subrepanda Presl 326
Polybotrya sensu Fée 275
Polybotrya Humb. & Bonpl.
276; 277-291 318
590
FLORA MALESIANA
[ser. II, vol. 1°
(Polybotrya)
sect. Arthrobotrya v.A.v.R.
2555215)
sect. Egenolfia Diels 255, 315
sect. Lomagramma Kuhn
277, 278
sect. Teratophyllum Christ
275, 276
appendiculata (Willd.) J.Sm.
322
var. costulata Bedd. 325
var. hamiltoniana auct. 323
var. hamiltoniana Bedd.
323
var. marginata C.Chr. 322
var. rhizophylla auct. 323
arfakensis Gepp 118, 119
articulata J.Sm. ex Fée 275,
276
duplicato-serrata Hayata 323
exaltata Brack. 323
hamiltoniana (Presl) Fée 322
lomarioides (BI.) Kuhn 279
marginata Bl. 322, 323
neglecta Fée 323
pteroides (J.Sm.) Kuhn 283
rhizophylla Pres] 323
serrulata (J.Sm.) Fée 323
vivipara Ham. ex Hook. 322,
323
Polylepideae Presl 291
Polypodiaceae 1, ix, x, Xii, Xiil,
xv, 71, 256, 257, 291
tribe Aspidieae 336
tribe Leptogrammeae 336
tribe Meniscieae 336
tribe Phegopteridiae 336
tribe Polypodieae 336
Polypodiopsis ti, xvi
Polypodium 256, 258, 304, 335,
336, 527
sect. Phegopteris Pres] 336,
353
acuminatum Houtt. 560
acuminatum Roxb. 556
acutum Roxb. 474
alternans Wall. 145
aoristisorum Harr. 538
appendiculatum Bedd. 412
var. squamaestipes Clarke
413
arboreum Lour. 157
arfakianum Baker 484
asperum Presl, non L. 512
auriculatum Wall. ex Hook.
412
auritum (Hook.) Lowe 345
barbatum (Fée) Hook. 391
barometz L. 165
borneense Hook. 528
canescens Bl. 497
(Polypodium)
cheilanthoides Baker 349
contaminans Wall. 135
cuspidatum Roxb. 533
decursive-pinnatum van Hall
354
dentatum Forssk. 557
dichotomum Thunb. 3, 33, 34
distans sensu Racib. 347
distans Don
var. minor Clarke 346
erubescens sensu Hook. 549
var. amboinense Baker 549
felinum Roxb. 563
firmulum Baker 519
fragile Baker 348
granulosum sensu Benth. 513
granulosum Presl 524
var. lobata Pres] 524
guianense Aubl. 278, 289
holophyllum Baker 526
imponens Ces. 391
intermedia (J.Sm.) Fée 323
invisum Forst. f. 502
latebrosum Wall. 115
lineare Burm. 33, 34
longifolium Roxb. 457
lunulatum Forst. 131
luxurians Kunze 387
molle Jacq. 557
nymphale G.Forst. 557
paludosum BI. 347
parasiticum L. 559
penangianum Hook. 565
pennigerum Forst. 462
phegopteris L. 353
pinwillii Baker 533
pteroides Retz. 545
rectangulare Zoll. 346
rubidum Hook. 532
rubrinerve (Mett.) Baker 531
scabrum Roxb. 391
scolopendria Burm. f. 303
simplicifolium (J.Sm.) Hook.
490
sinuatum Presl 326
sophoroides Thunb. 560
subvillosum Moore 412
tenericaule Hook. 348
tottum Thunb. 386, 387
trichodes Houlst. & Moore
348
truncatum Poir. 429
unitum L. 386, 477, 478
unitum Thunb. 560
urophyllum sensu Benth. 513
urophyllum sensu Racib. 513
urophyllum (Mett.) Hook.
533
var. uniseriale Hook. 513
urophyllum Wall. 513
Polypodium group xi
Polystichopsis ii, xix
Polystichum Roth i, xix, 335
aculeatum 335
auriculatum
var. nervosum Christ 539
benoitianum Gaud. 500
cuspidatum Pres] 325
riedleanum Gaud. 456
torresianum Gaud. 348
truncatum Gaud. 429, 430,
436
Proferea Presl 336, 436
excellens (Bl.) Pres] 462
Pronephrium Pres] 333, 335,
336, 338, 340, 341, 343,
437, 464, 507, 555
subg. Menisciopsis Holtt.
509, 530
sect. Grypothrix Holtt. 334,
340, 341, 507, 509, 531,
534
sect. Menisciopsis 509, 531,
534
subg. Pronephrium 509
sect. Dimorphopteris
(Tagawa & Iwats.) Holtt.
509, 516, 538
sect. Pronephrium 509, 517
acanthocarpum (Copel.)
Holtt. 509, 531, 532
affine (BI.) Pres! 509, 518, 527
amboinense (Willd.) Holtt.
508*, 517, 519
amphitrichum Holtt. 517,
§21, 525
aquatiloides (Copel.) Holtt.
509, 517, 520
articulatum (Houlst. &
Moore) Holtt. 516, 517
asperum (Presl) Holtt. 506*,
507, 509, 511, 512, 513
bakeri (Harr.) Holtt. 511, 516
beccarianum (Ces.) Holtt.
491, 511, 513, 514, 516
borneense (Hook.) Holtt.
518, 528
brauseanum Holtt. 511, 514
bulusanicum (Holtt.) Holtt.
511, 515
buwaldae Holtt. 517, 554
celebicum (Baker) Holtt.
508*, 518, 523
clemensiae (Copel.) Holtt.
517, 521
var. clemensiae 521, 522
var. degenerum (Christ)
Holtt. 517, 522
cuspidatum (BI.) Holtt. 533,
534, 536, 537
debile (Baker) Holtt. 517, 522
1981] Index to scientific plant names 591
(Pronephrium) (Pronephrium) Pseudophegopteris Ching 333,
diminutum (Copel.) Holtt. pentaphyllum (Rosenst.) 339, 340, 342, 343, 345
$11, 515 Holtt. 507, 511, 514 aurita (Hook.) Ching 344*,
euryphyllum (Rosenst.) peramelense Holtt. 517, 518 345
Holtt. 509, 511 ramosii (Christ) Holtt. 510*, cyclocarpa Holtt. 345, 346
exsculptum (Baker) Holtt. 532, 534, 536, 537 kinabaluensis Holtt. 345, 346,
518, 528 var. minahassae Holtt. 534, 347
firmulum (Baker) Holtt. 517, 537 oppositipinna (v.A.v.R.)
518, 519 var. ramosii 534, 537 Ching 346
giluwense Holtt. 517, 520 repandum (Fée) Holtt. 332*, paludosa (BI.) Ching 344%,
glandulosum (BI.) Holtt. 509, 5095, S10*; 5135531, 533 345, 347
S17 522 rhombeum (Christ) Holtt. pyrrhorhachis (Kunze) Ching
granulosum (Presl) Holtt. 508*, 509, 518, 525, 529 345, 347
508*, 514, 518, 524, 525 rubicundum (v.A.v.R.) Holtt. rectangularis (Zoll.) Holtt.
gymnopteridifrons (Hayata) 33275105534. 536, 537 344*, 345, 346
Holtt. 509, 511, 513
heterophyllum (Presl) Holtt.
506*, 511, 515
hewittii (Copel.) Holtt. 508%,
518, 527, 529
hosei (Baker) Holtt. 389, 518,
526
kjellbergii Holtt. 517, 521
lakhimpurense (Rosenst.)
Holtt. 509, 530
lastreoides Pres] 468, 507,
509
lineatum (BI.) Pres] 495, 507,
509, 511, 512
melanophlebium (Copel.)
Holtt. 511, 514
menisclicarpon (BI.) Holtt.
432, 509, 513, 516, 517,
518, 526
merrillii (Christ) Holtt. 518,
527, 529
micropinnatum Holtt. 506%,
509, 511, 515
millarae Holtt. 517, 518
minahassae Holtt. 517, 523
moniliforme (Tagawa &
K.I wats.) Holtt. 516, 517,
519
nitidum Holtt. 509, 510*, 531,
533
nothofageti Holtt. 466
palauense (Hosok.) Holtt. 516
palopense Holtt. 519
x parishii (Bedd.) Holtt. 534,
535
peltatum (v.A.v.R.) Holtt.
518, 529
var. aberrans Holtt. 530
var. peltatum 530
var. peninsulare Holtt. 507,
530
var. persetiferum Holtt. 530
var. tenompokense (C.Chr.)
Holtt. 530
penangianum (Hook.) Holtt.
531
ssp. sulawiense Iwats. 536
tubidum (Hook.) Holtt. 531
532
rubrinerve (Mett.) Holtt. 531
salicifolium (Wall. ex Hook.)
Holtt. 534, 535
samarense (Copel.)
518, 524
scopulorum Holtt. 507, 531,
532
simillimum (C.Chr.)
507, 517, 520, 525
solsonicum Holtt. 517, 523
sulawiense (Iwats.) Holtt. 534,
536
thysanoides Holtt. 518, 525
tibangense (C.Chr.) Holtt.
485
trachyphyllum Holtt. 518, 525
triphyllum (Sw.) Holtt. 335,
507, 510*, 534, 535
womersleyi Holtt. 506*, 507,
511, 514
x xiphioides (Christ) Holtt.
518, 525, 529
Prosaptia il, XVii
Protolindsaya Copel. 184
brooksii Copel. 191
Pseudocyclosorus Ching 335,
340, 341, 343, 373, 374,
377, 413, 414
caudatus Holtt. 433
caudipinna Ching 413
ciliatus (Benth.) Ching 375,
413
esquirolii 413
excisus Holtt. 434
falcilobus Ching 374
ochthodes 413
Holtt.
Holtt.
petrophilus (Copel.) Holtt.
435
repens 413
tylodes (Kunze) Ching 412*,
413
xylodes 413
Pseudogyratae 71
sumatrana Holtt. 345, 346
tenggerensis Holtt. 345, 346
Psilotaceae ii
Psilotales ii
Psilotopsida ti
Psilotum ti
Psomiocarpa Ili, xx
Pteridaceae 71
Pterideae 255
Pteridium 11, vii, xix
Pteridrys ili, xx, 257
Pteris ili, iv, v, yli, xx, 253, 335
angulata Pres] 211
confluens Thunb. 377
interrupta Willd. 386, 387
polypodioides Poir. 387
stricta Poir. 211
tottum Thunb. 387
Pteris group ili, ix, xX, Xi, Xil,
xiii, xx, 178
Pteropsida ii, ix, xil
Pycnoloma ii, xvi
Pyrrosia il, vi, XV1
Quercifilix iii, xx
Ramondia Mirbel 39, 44
flexuosa (L.) Mirbel 53
Ripidium Bernh. 39, 44, 561
dichotomum Bernh. 41
Rhipidopteris Fée ex Schott
255
Saccoloma adiantoides (BI.)
Pres] 249
boryanum (Presl) Pres! 237
cuneifolium Presl 180
hemipterum (Bory) Pres] 237
pinnatum (Cav.) Pres] 193
sorbifolia Christ 163
sp. 197
Salvinia iii, xxi
Salviniaceae iii, viii, ix, xii, xxi
Schizaea Sm. ii, vi, viii, xv, 37,
38, 39, 40, 45, 561
asperula Wakef. 39
592
(Schizaea)
australis Gaud. 44
biroi Richter 41
chilensis Phil. 44
copelandica Richter 41
dichotoma (L.) Sm. v, 38%,
39, 40, 41*, 43
digitata (L.) Sw. 37, 40, 41,
42*, 44
fistulosa Labill. 37, 40, 43%,
44
var. malaccana Prantl 44
forsteri Spr. 41
hallieri Richter 44
inopinata Selling 37, 40, 42*
malaccana Baker 37, 40, 43*,
44
var. malaccana 44
var. robustior C.Chr. 44,
561
papuana Brause 44
paucijuga Holtt. 44
ponapensis Hosok. 42
propinqua A.Cunn. 44
robusta Baker 561
spirophylla Troll 37, 40, 42
wagneri Selling 37, 40, 43*, 44
Schizaeaceae ii, vii, X, Xill, Xv,
37-61, 561
Schizaeales 39
Schizaeopsis 37
Schizocaena J.Sm. 73, 76, 141
alternans J.Sm. 145
arthropoda (Copel.) Copel.
143
brunonis J.Sm. ex Hook. 141,
143
capitata Copel. 142
gaudichaudii Fée 143
kinabaluensis (Copel.) Copel.
143
moluccana (R.Br.) Copel.
143
Schizolegnia ensifolia (Swartz)
Alston 211
heterophylla (Dryand.) Al-
ston 210
javanensis (Bl.) Alston 208
orbiculata (Lamk) Alston 206
stortii (v.A.v.R.) Alston 253
walkerae (Hook.) Alston 229
Schizolepton ii, xvii
cordatum (Gaud.) Fée 253
Schizoloma Gaud. 198, 204,
229, 230
agatii Brack. 211
angustum Copel. 232
auriculatum (v.A.v.R.)
v.A.v.R. 231
billardieri Gaud. 204, 211
cordatum Gaud. 253
FLORA MALESIANA
(Schizoloma)
coriaceum v.A.v.R. 212
davallioides (Bl.) Moore 219
divergens (Hook. & Grev.)
Kuhn 233
var. auriculata v.A.v.R.
231
ensifolium (Swartz) J.Sm.
211
f. grifithianum Domin 211
f. praelongum Domin 211
f. pteroides Domin 211
f. subsimplex Domin 211
f. typicum Domin 211
var. attenuatum Domin 211
var. borneense Domin 211
var. clarkeanum Domin
211
var. heterophyllum
and.) Domin 210
var. intercedens Domin 211
var. lanceolata (Labill.)
R.Bonaparte 211
var. longipinnum
211
var. medium (R.Br.) Domin
208
ferulaceum (Moore) Kuhn
253
fuligineum Copel. 230
griffithianum (Hook.) Fée 211
guerinianum Gaud. 230
heterophyllum (Dryand.)
J.Sm. 210
var. speluncae Copel. 211
hosei (C.Chr.) Copel. 253
(Dry-
Domin
induratum (Baker) C.Chr.
230
jamesonioides (Baker) Copel.
japy
javae Fée 211
jJavanense (BI.) Holtt. 208
lanceolatum (Labill.) Presl
211
lobatum (Poir.) Bedd. 219
medium (R.Br.) Kuhn 208
nitens (Bl.) Bedd. 222
orbiculatum (Lamk) Kuhn
206
ovata (erron.) 231
ovatum (J.Sm.) Copel. 231
pellaeiforme (Christ) C.Chr.
232
pentaphyllum (Hook.) Fée
211
recurvatum (Hook.) Moore
222
retusum (Cav.) Kuhn 180,
253
schizolomae (v.A.v.R.)
v.A.v.R. 230
[ser. Il, vole
(Schizoloma)
stortii v.A.v.R. 197, 253
tenerum auct. 204
walkerae (Hook.) Kuhn 229
Schizostege iii, xx
Scleroglossum ii, xvii
Scyphularia ii, xviii
Selaginella ii
Selaginellaceae ii
Selliguea Bory ii, xvi, 291
Senftenbergia 37, 38, 45, 561
Sitolobium stramineum Brack.
169
Sphaeropteris Bernh. 124, 562
Sphaerostephanos J.Sm. iii, xxi,
334, 337, 338, 340, 341,
343, 415, 436, 509, 517,
521,523,530) 58855411
acrostichoides (Desv.) Holtt.
441, 456
adenostegius (Copel.) Holtt.
442, 444, 447, 466
alatellus (Christ) Holtt. 440*,
441, 451
albosetosus
448, 505
alcasidii Holtt. 448, 503
alpinus Holtt. 445, 488
alticola Holtt. 444, 484
angustibasis Holtt. 443, 476
(Copel.) Holtt.
angustifolius (Presl) Holtt.
444, 481
appendiculatus (BI.) Holtt.
441, 442, 452
aquatilis (Copel.) Holtt. 442,
444, 466
arbuscula (Willd.) Holtt. 439,
44]
archboldii (C.Chr.) Holtt.
440*, 444, 485
arfakianus (Baker) Holtt. 439,
444, 446, 484, 486
asplenioides J.Sm. 437, 449
atasripii (Rosenst.) Holtt.
448, 504
austerus (Brause) Holtt. 447,
502
baramensis (C.Chr.) Holtt.
440*, 443, 447, 473
batacorum (Rosenst.) Holtt.
440*, 443, 475
batjanensis (Rosenst.) Holtt.
442, 467
batulantensis Holtt. 433, 473
benoitianus (Gaud.) Holtt.
447, 500
canescens (BI.) Holtt. 447, 497
carrii Holtt. 442, 463
cataractorum (Wagn. &
Greth.) Holtt. 442, 468
caulescens Holtt. 443, 472
1981] Index to scientific plant names 593
(Sphaerostephanos) (Sphaerostephanos) (Sphaerostephanos)
confertus (Brause) Holtt. 446,
448, 482, 491, 50S
convergens Holtt. 443, 477
cyrtocaulos (v.A.v.R.) Holtt.
443, 473
daymanianus Holtt. 441, 451
debilis (Mett.) Holtt. 439, 448
dichrotrichoides (v.A.v.R.)
Holtt. 444, 482
dichrotrichus (Copel.) Holtt.
441, 442, 453
dimidiolobatus Holtt. 441, 459
dimorphus (Brause) Holtt.
447, 501
diversilobus (Presl) Holtt.
441, 445, 452
echinosporus (v.A.v.R.) Holtt.
446, 495
efogensis Holtt. 444, 445, 482
ekutiensis Holtt. 444, 483
ellipticus (Rosenst.) Holtt.
443, 444, 474, 475
var. ellipticus 475
var. glabrior Holtt. 475
eminens (Baker) Holtt. 439,
449
erectus (Copel.) Holtt. 445,
490
erwinii Holtt. 448, 504
exindusiatus Holtt. 442, 463
fenixii Holtt. 495
flavoviridis Holtt. 439, 450
foliolosus Holtt. 445, 464, 488
foxworthyi Holtt. 443, 475
grandescens Holtt. 442, 465
gregarius (Copel.) Holtt. 443,
445, 447, 480
gymnorachis Holtt. 443, 479
hamiferus (v.A.v.R.) Holtt.
446, 447, 496
hastatopinnatus (Brause)
Holtt. 439, 442, 444, 446,
470
hellwigensis Holtt. 447, 500
hendersonii Holtt. 446, 493
hernaezii Holtt. 441, 459
heterocarpus (BI.) Holtt.
332*, 441, 444, 457, 465,
470, 472, 474, 482
var. bordenii Holtt. 457,
465
hirsutus (Kunze ex Mett.)
Holtt. 441, 443, 455, 466,
472
var. celebicus Holtt. 455,
456
var. hirsutus 455
hirtisorus (C.Chr.) Holtt. 439
hispidifolius (v.A.v.R.) Holtt.
438*, 442, 466, 469
hispiduliformis (C.Chr.) Holtt.
439, 442, 443, 447, 464,
470
var. brassii Holtt. 470
var. hispiduliformis 470
var. vinkii Holtt. 470
hoalensis Holtt. 445, 488
humilis Holtt. 442, 468
inconspicuus (Copel.) Holtt.
438*, 439, 446, 493, 494
indrapurae Holtt. 443, 475
invisus (Forst. f.) Holtt. 339,
439, 447, 479, 500, 502,
556, 565
irayensis (Copel.) Holtt. 446,
495
isomorphus Holtt. 441, 458
kalkmanii Holtt. 441, 444,
445, 454
kotoensis (Hayata) Holtt. 460
lamii Holtt. 441, 444, 454,
455
larutensis (Bedd.)
438*, 445, 447, 489
lastreoides (Presl) Holtt. 442,
468, 538
latebrosus (Kunze ex Mett.)
Holtt. 443, 444, 471, 472
lithophyllus (Copel.) Holtt.
446, 496
lobangensis (C.Chr.) Holtt.
443, 479, 480
lobatus (Copel.) Holtt. 443,
458, 474, 479, 488
loherianus (Christ) Holtt. 445,
489
lucbanii Holtt. 442, 463
magnus (Copel.) Holtt. 442,
465, 472
major (Copel.) Holtt. 442, 464
makassaricus Holtt. 447, 498
melanorachis Holtt. 441, 457
menadensis Holtt. 442, 469
mengenianus Holtt. 446, 491
mjobergii Holtt. 446, 494
morotaiensis Holtt. 441, 453
moseleyi Holtt. 441, 460
multiauriculatus (Copel.)
Holtt. 446, 447, 492
muluensis Holtt. 447, 499
mundus (Rosenst.) Holtt. 447,
448, 503
mutabilis (Brause) Holtt. 446,
466, 494, 565
nakaikei Holtt. 444, 482, 483
neotoppingii Holtt. 447, 500
norrisii (Rosenst.) Holtt. 437,
438*, 445, 446, 475, 485
novae-britanniae Holtt. 443,
477
©@Chr:
novoguineensis (Brause) Holtt.
439, 442, 445, 446, 469
nudisorus Holtt. 443, 476
obtusifolius (Rosenst.) Holtt.
447, 503
omatianus Holtt. 444, 483
oosorus (Baker) Holtt. 447,
500
paripinnatus (Copel.) Holtt.
443, 476
peltochlamys (C.Chr.) Holtt.
446, 495, 509, 513
penniger (Hook.) Holtt. 332*,
439, 441, 450, 452, 454,
460, 461, 478
var. excellens (BI.) Holtt.
461, 462
var. karoensis Holtt. 461,
462
var. penniger 461
perglanduliferus
Holtt. 441, 454
pilosissimus Holtt. 448, 504
pilososquamatus _(v.A.v.R.)
Holtt. 438*, 444, 448, 481
plurifolius (v.A.v.R.) Holtt.
446, 492
plurivenosus Holtt, 447, 496
polisianus Holtt. 445, 488
polycarpus (BI.) Copel. 437,
438*, 439, 444, 448, 565
polyotis (C.Chr.) Holtt. 442,
445, 446, 464, 488
porphyricola (Copel.) Holtt.
443, 444, 472
posthumii Holtt. 446, 493
potamios Holtt. 447, 502
productus (Kaulf.) Holtt. 423,
441, 460
pseudomegaphyllus (v.A.v.R.)
Holtt. 441, 452
pterosporus (v.A.v.R.) Holtt.
445, 446, 486
var. altibus Holtt. 487
var. crenatus Holtt. 487
var. pterosporus 487
pullenii Holtt. 443, 480
punctatus Holtt. 460, 461
pycnosorus 439
reconditus Holtt. 443, 472
rigidus (Ridl.) Holtt. 444, 445,
484
roemerianus (Rosenst.) Holtt.
447, 499
rudis (Ridl.) Holtt. 447, 501
var. micans (Brause) Holtt.
501
var. rudis 501
sagittifolius (Bl.) Holtt. 445,
446, 487
(v.A.v.R.)
594 FLORA MALESIANA [ser. II, volo a
(Sphaerostephanos) (Sphaerostephanos) (Stegnogramma)
sakayensis (Zeiller) Holtt. vestigiatus (Copel.) Holtt. 441, subcalcarata (v.A.v.R.) Holtt.
392 453 541*, 543
santomasii Holtt. 442, 465
sarasinorum Holtt. 447, 498
semicordatus Holtt. 445, 486
semimetralis Holtt. 445, 486
sessilipinna (Copel.) Holltt.
443, 474, 479
simplicifolius (J.Sm. ex
Hook.) Holtt. 445, 490,
509
smithianus (Presl) Holtt. 456
solutus Holtt. 441, 459
spenceri (Christ) Holtt. 445,
446, 481, 491, 509
squamatellus Holtt. 447, 497
stenodontus (Copel.) Holtt.
439, 450
stipellatus (Bl.) Holtt. 439,
446, 492
stresemannii Holtt. 445, 490
subalpinus (v.A.v.R.) Holtt.
443, 474
subcordatus Holtt. 442, 469
suboppositus Holtt. 447, 498
subulifolius (v.A.v.R.) Holtt.
446, 494
sudesticus Holtt. 439, 448
sumatrensis (v.A.v.R.)
C.Chr. 489
talamauensis (v.A.v.R.)
C.Chr. 449, 450
tandikatensis (v.A.v.R.) Holtt.
442, 463
telefominicus Holtt. 447, 499
tephrophyllus (Copel.) Holtt.
444, 480
tibangensis (C.Chr.) Holtt.
445, 485
todayensis (Christ) Holtt.
456, 471
toppingii (Copel.) C.Chr. 449,
500
trichochlamys Holtt. 442, 456,
465
trimetralis Holtt. 439, 451
uaniensis Holtt. 446, 494
uniauriculatus (Copel.) Holtt.
442, 467
unijugus Copel. 449
unitus (L.) Holtt. 339, 439,
443, 445, 446, 477, 556
var. mucronatus (Christ)
Holtt. 478
var. papilliferus Holtt. 478,
479
var. unitus 478
urdanetensis (Copel.) Holtt.
442, 467, 474
veitchii Holtt. 454
warburgii (Kuhn & Christ)
Holtt. 441, 462
wauensis Holtt. 448, 505
williamsii (Copel.) Holtt. 439,
450
woitapensis Holtt. 444, 483
Sphaerostephanos group 340,
341
Sphenomeris Maxon ii, xix,
177, 178, 179
alutacea 179
biflora (Kaulf.) Tagawa 177,
178, 179, 180, 182, 184,
563
chinensis (L.) Maxon 178,
563
var. chinensis 181*, 182,
184
var. divaricata (Christ)
Kramer 181*, 183, 184
var. rheophila Kramer
181*, 183
var. tenuifolia (Lamk)
C.Chr. 183
chusana (L.) Copel. 182
var. divaricata (Christ)
Tard.-Blot 183
var. littoralis (Tagawa)
H.Ito 182
var. tenuifolia (Lamk)
Holtt. 183
clavata (L.) Maxon 178, 179
retusa (Cav.) Maxon 178, 179,
180, 181*, 184, 253
veitchii (Baker) C.Chr. 180
181*, 184
Sphenopsida ii
Sphenopteris 1
Stegnogramma BI. ili, xxi, 257,
334, 335, 337, 340, 341,
342, 540
sect. Dictyocline Iwats. 541
sect. Haplogramma _ Iwats.
540
sect. Leptogramma_ Iwats.
540
aspidioides Bl]. 540, 541*
asplenioides (J.Sm.) Fée 449
celebica (Ching) Holtt. 541,
543
dissitifolia Holtt. 541, 543
gymnocarpa (Copel.) Iwats.
541, 542, 543
ssp. celebica Iwats. 543
pozoi (Lagasca) Iwats. 542,
543
var. petiolata (Ching) Holtt.
541, 542
Stenochlaena J.Sm. ili, iv, xx,
255, 261, 265, 266, 267
sect. Eustenochlaena Diels
259, 265
sect. Lomariopsis 259, 261,
267, 276
sect. Stenochlaena Und. 267
sect. Teratophyllum Und.
261, 263, 265, 267
abrupta v.A.v.R. 262
aculeata sensu v.A.v.R. 269
aculeata (Bl.) Kunze 269
var. crassior v.A.v.R. 269,
270
arthropteroides Christ 270
cochinchinensis (Fée) Und.
262
dubia v.A.v.R. 118
gracilis Kunze 269
var. williamsii v.A.v.R. 270
hiigelii Und. 263, 276
intermedia Copel. 261
kingii Copel. 261
leptocarpa (non Fée) Und.
262
palustris (Burm.
266, 269
raciborskii C.Chr. 262
rotundifoliata R.Bonap. 270
scandens J.Sm. 266, 269
smithii (Racib.) v.A.v.R. 262
smithii Und. 262
sorbifolia sensu Bedd. 261,
262, 275
sorbifolia (L.) J.Sm. 253
ssp.) C:Chr. 270
var. spectabilis
262
subtrifoliata Copel. 261
williamsii Und. 270
Stenolepia li, xix
tristis (BI.) v.A.v.R. 158
Stenoloma Fée 179
biflorum (Kaulf.) Ching 182
blumeanum (Hook.) Fée 216
chinense (L.) Bedd. 182
chusanum (L.) Ching
var. littorala (Tagawa) Ito
182
clavatum (L.) Fée 179
dumosum Fée 179
littorala Tagawa 182
retusum (Cav.) Fée 180
tenuifolium (Lamk) Fée 182
veitchii (Baker) C.Chr. 184
Stenoneura Christ 291
Stenosemia ili, xx
dimorpha Copel. 327
f.) Bedd.
v.A.v.R.
1981]
Index to scientific plant names
595
Sticherus Pres] 3, 18
sect. Eu-Sticherus C.Chr. 18
sect. Hicriopteris C.Chr. 11
amoenus Nakai 26
barbulus Nakai 26
bolanicus Copel. 24
brassii Nakai 24
caudatus Copel. 24
erectus Copel. 22
flabellatus H.St.John 25
habbemensis Copel. 22
hallieri Nakai 26
hirsutus (erron.) 26
hirtus Ching 26
var. candidus Nakai 26
var. virescens Nakai 27
hispidus Copel. 24
kajewskii Copel. 22
kajewskii St.John 561
laevigatus Pres] 20
lamianus Copel. 27
loheri Copel. 25
milnei Ching 22
monticola Nakai 24
myriapoda Nakai 20, 21
oceanicus St.John 22
peninsularis Ching 26
perpaleaceus Copel. 25
pinnatus Copel. 24
plumaeformis Nakai 22
pseudoscandens Copel. 23
pulcher Copel. 23
reflexipinnula Copel. 20
truncatus Nakai 20
venosus Copel. 20
vestitus Ching 23
Stromatopteris 1, 2
Stylites 63
gemmifera W.Rauh 64
Synaphlebium J.Sm. 179, 198
davallioides (Bl.) J.Sm, 219
nitens (Bl.) J.Sm. 222
recurvatum Hook. 217, 222
Syngramma ii, xvii, 291
angusta Copel. 380
borneensis (Hook.) J.Sm. 314
Taenitis ii, xvii
blechnoides (Willd.) Swartz
253
cordata (Gaud.) Holtt. 253
trilobata Holtt. 253
Tapeinidium (Presl) C.Chr. ii,
xix, 177, 178, 179, 184,
197
acuminatum Kramer 185,
194, 196*
amboynense (Hook.) C.Chr.
186, 187*, 253
amplum Copel. 187
(Tapeinidium)
atratum Kramer 185, 187*,
189
bartlettii Copel. 197, 198
biserratum sensu. v.A.v.R.
191
biserratum (BI.) v.A.v.R. 185,
194
brooksii (Copel.) C.Chr. 191
buntifolium Kramer 185, 186
calomelanos Kramer 185, 190
denhamii (Hook.) C.Chr. 185,
186, 187*, 253, 563
firmulum (Baker) C.Chr. 193
gracile (BI.) v.A.v.R. 185, 190
map
longipinnulum (Ces.) C.Chr.
185, 194, 195*, 197
luzonicum (Hook.) Kramer
185, 186, 191, 194, 253
var. leptophyllum Kramer
191
var. luzonicum 191, 192%,
193
var. thelypteridoides
Kramer 191, 193
marginale Copel. 197
melanesicum 194
moluccanum sensu Wagn. &
Teratophyllum Mett. ex Kuhn
LU PRX See ee ONT
258, 259, 261, 263, 266,
275, 565
sect. Polyseriatae Holtt. 257,
259, 265, 275
sect. Teratophyllum Holtt.
265, 276, 565
aculeatum (BI.) Mett. ex
Kuhn 264*, 266, 267,
268*, 269, 275, 565
var. aculeatum 269
var. inermis Mett. ex Kuhn
269, 275
var. montanum Holtt. 267,
269
arthropteroides (Christ)
Holtt. 267, 270, 271*
articulatum (J.Sm. ex Fée)
Mett. ex Kuhn 255, 259,
DOE 27475255276
brightiae (F.v.M.) Holtt. 263,
276
clemensiae Holtt. 267, 270,
VIB 275
gracile (BI.) Holtt. 267, 268%,
269
koordersii Holtt. 267, 271*
leptocarpum (Fée) Holtt. 267,
270
ludens (Fée) Holtt. 266, 267,
269) 2727) 275
luzonicum Holtt. 267, 269,
270, 275
rotundifoliatum (R.Bonap.)
Holtt. 266, 267, 269, 270,
D125
wilkesianum (Brack.) Holtt.
275, 276, 289
Greth. 186
moluccanum (BI.) C.Chr.
187, 197
novoguineense Kramer 186,
188, 192*
obtusatum v.A.v.R. 186,
187*, 189
oligophlebium (Baker) C.Chr.
185, 191
philippinense (Harr.) C.Chr.
191
pinnatum (Cav.) C.Chr. 162,
185, 193, 194 map, 253
prionoides Kramer 185, 194,
Thayeria li, xvi
Thelypteridaceae 331-560
Thelypteris Adans. 335
Thelypteris sensu Ching 386
Thelypteris Schmidel i, v, vi,
1954
stenocarpum v.A.v.R. 185,
188
stenolobum (Baker) Wagner
187
sumatranum v.A.v.R. 191
tenue sensu Kramer 186
tenue (Brack.) Copel. 197
tenuius Copel. 186
Megan (Cay, ii, VW, xox Wail
258, 266, 334, 335, 338,
339, 527
excellens C.Chr. 462
serrata Cav. 478
Tectaria group ill, x, Xi, Xi, Xili,
Gh, 0. ZAI
Tempskya 37
Teratophyllum sensu Holtt. 265
KX1,655)1 6681583395559.
336, 337, 338, 339, 340,
341, 342, 375, 378, 386,
539
sect. Macrothelypteris H.Ito
347
sect. Parathelypteris H.Ito
370
subg. Abacopteris Iwats. 507
subg. Cyclogramma_ Iwats.
411
subg. Cyclosoriopsis Iwats.
550, 551
subg. Dimorphopteris Iwats.
516
subg. Glaphyropteridopsis
sect. Mesoneuron Iwats. 377
596
(Thelypteris)
sect. Neocyclosorus Iwats.
436
subg. Glaphyropteris Christ
333
subg. Haplodictyum Iwats.
507
subg. Meniscium
sect. Ampelopteris Reed
387
subg. Phegopteris
sect. Lastrella Iwats. 343,
353
sect. Phegopteris Iwats.
353
subg. Pneumatopteris Iwats.
414
sect. Macrocyclosorus
Iwats. 414, 436
subg. Sphaerostephanos
Iwats. 436
subg. Thelypteris
sect. Metathelypteris
Iwats. 347, 348, 350
sect. Thelypteris Iwats. 370
acanthocarpa (Copel.) Reed
532
acrostichoides (Michx) Newl.
456
acuminata (Houtt.) Morton
560
adenophora (C.Chr.) Reed
455
adenostegia (Copel.) Reed
466
albociliata (Copel.) Reed 560
albosetosa (Copel.) Reed 505
alta (Brause) Reed 404
amboinensis (Willd.) Reed
519
angulariloba Ching 359, 360
angustata (Copel.) Reed 456
angustipes (Copel.) Reed 471
aoristisorus (Harr.) Reed 538
appendiculata (Bl.) Reed 452
aquatilis (Copel.) Reed 466
aquatiloides (Copel.) Reed
520
archboldii (C.Chr.) Reed 485
arfakiana (Baker) Reed 484
arida (D.Don) Morton 556
aspera Reed 512
asymmetrica (Fée) Reed 452
atasripii (Rosenst.) Reed 504
atrospinosa (C.Chr.) Reed
390
attenuata (O.Ktze) Morton
549
aureoviridis (Rosenst.) Reed
353
auriculata (J.Sm.) Iwats. 412
FLORA MALESIANA
(Thelypteris)
auriculifera (v.A.v.R.) Ching
383
aurita (Hook.) Ching 345
austera (Brause) Reed 502
austrophilippina (Copel.)
Reed 479
badia (v.A.v.R.) Ching 369
bakeri (Harr.) Reed 516
baramensis (C.Chr.) Ching
473
bartlettii (Copel.) Reed 522
batacorum (Rosenst.) Reed
476
batjanensis (Rosenst.) Reed
467
beccariana (Ces.) Reed 383,
513
beddomei (Baker) Ching 371
belensis (Copel.) Reed 403
benguetensis (Copel.) Reed
560
bipinnata (Copel.) Reed 405
blumei Panigrahi 558
boholensis (Copel.) Reed 456
bordenii (Christ) Reed 457
borneensis (Hook.) Reed 528
brackenridgei_ (Mett.) Reed
408
brassii (C.Chr.) Reed 373
brooksii (Copel.) Reed 423
brunnea sensu Ching 347
brunnescens Reed 452
bunnemeijeri Reed 543
calcarata (BI.) Ching 374
callosa (Bl.) Iwats. 419
canlaonensis (Copel.) Reed
479
cataractorum (Wagn. &
Greth.) Reed 469
cavitensis (Copel.) Reed 421
celebica (Baker) Reed 523
cesariana (C.Chr.) Reed 513
chamaeotaria (Christ) Reed
524
chlamydophora (C.Chr.)
Ching 384
christii (C.Chr.) Reed 394
ciliata (Benth.) Ching 375
clemensiae (Copel.) Reed 521
compacta (Copel.) Reed 527
conferta (Brause) Reed 491
confluens (Thunb.) Morton
BSI
confusa (Copel.) Reed 539
conterminoides (C.Chr.)
Reed 367
contigua (Rosenst.) Reed 557
coriacea (Brause) Ching 368
costata (Brack.) Reed 421
costulisora (Copel.) Reed 409
[ser. II, vol.
(Thelypteris)
crassa (Copel.) Reed 408
crassifolia (Bl.) Ching 377,
384
var. motleyana Ching 381
cuspidata (BI.) Iwats. 536
cyrtocaulos (v.A.v.R.) Reed
473
dayi (Bedd.) Nayar & Kaur
353
debilis (Mett.) Reed 448
decursive-pinnata (van Hall)
Ching 343, 354
degenera (Christ) Reed 522
deltiptera (Copel.) Reed 420
dentata (Forssk.) E.St.John
557
diaphana (Brause) Ching 362
dichrotricha (Copel.) Reed
453
dichrotrichoides (v.A.v.R.)
Reed 482
dicranogramma (v.A.v.R.)
Reed 419
dimorpha (Brause) Reed 501
distincta (Copel.) Reed 548
divergens (Rosenst.) Reed
384
diversiloba (Presl) Reed 452
diversivenosa (v.A.v.R.) Ching
353
dryopteroidea (Brause) Reed
401
duplosetosus (Copel.) Reed
468
dura (Copel.) Reed 368
ecallosa (Holtt.) Reed 425
echinata (Mett.) Reed 379
echinospora (v.A.v.R.) Reed
495
edanyoi (Copel.) Reed 525
elliptica (Rosenst.) Reed 474
elmerorum (Copel.) Reed 485
engleriana (Brause) Reed 366
ensipinna (Brause) Ching 404
erectus (Copel.) Reed 490
erubescens 550
eugracilis (Copel.) Reed 371
euryphylla (Rosenst.) Reed
S11
exsculpta (Baker) Iwats. 528
extensa (BI.) Morton 548
falcatipinnula (Copel.) Reed
406
falcatula (Christ) Reed 557
farinosa (Brause) Reed 503
fasciculata (Fourn.) Ching
367
ferox (Bl.) Tagawa & Iwats.
391
finisterrae (Brause) Reed 432
1981] Index to scientific plant names 597
(Thelypteris) (Thelypteris) (Thelypteris)
flaccida (Bl.) Ching 352
flavo-virens (Rosenst.) Reed
353
forsteri Morton 502
foxii (Christ) Reed 431
fulgens (Brause) Ching 403
glanduligera (Kunze) Ching
370, 373
glaucescens (Brause) Ching
469
gracilescens (Bl.) Ching 351
grammitoides (Christ) Ching
373
granulosa (Presl) Reed 524
gregaria (Copel.) Reed 480
grisea (Baker) Ching 375
gymnocarpa (Copel.) Morton
542
haenkeana (Presl) Reed 478
halconensis (Copel.) Reed
471
hallieri (Christ) Reed 379
hamifera (v.A.v.R.) Reed 496
harveyi (Mett.) Proctor ex
Iwats. 555
hastatopinnata (Brause)
Reed 470
herbacea Holtt. 360
heterocarpa (BI.) Morton 457
var. glaucostipes (Bedd.)
Reed 471
hewittii (Copel.) Reed 527
hirsutipes (Clarke) Ching 359,
370
hirtopilosa (Rosenst.) Reed
557
hispidifolia (v.A.v.R.) Reed
466
hispidula (Decne) Reed 557
hispiduliformis (C.Chr.) Reed
470
horridipes (v.A.v.R.) Reed
394
hosei (Baker) Iwats. 526
hunsteiniana (Brause) Reed
403
immersa (BI.) Ching 158, 547
imponens (Ces.) Reed 391
impressa (Desv.) Reed 548
inclusa (Copel.) Reed 423
inconspicua (Copel.) Ching
493
indochinensis (Christ) Ching
359
interrupta (Willd.) Iwats. 386
irayensis (Copel.) Reed 495
iridescens (v.A.v.R.) Reed
522
jacobsii (Holtt.) Reed 520
japonica 370
keysseriana (Rosenst.) Reed
432
khamptorum Holtt. 374
klossii (Ridl.) Ching 361
kotoensis (Hayata) Iwats. 460
laevis (Mett.) Reed 424
lanceola (Christ) Reed 529
larutensis (Bedd.) Reed 489
latebrosa (Mett.) Reed 471
latifolia (Presl) Reed 526
latipinna (Benth.) Iwats. 558
lauterbachii (Brause) Reed
367
lepidopoda (C.Chr. & Tard.)
Reed 429
leucadenia (Copel.) Reed 460
leucolepis (Presl) Ching 349
ligulata (Presl) Ching 431
lineata (BI.) Iwats. 512
lithophylla (Copel.) Reed 496
lobangensis (C.Chr.) Reed
479
lobata (Copel.) Reed 474
loheriana (Christ) Reed 489
longipes (BI.) Reed 424
magnus (Copel.) Reed 465
malayensis (C.Chr.) Reed 522
matutumensis (Copel.) Reed
467
megalocarpa (v.A.v.R.)
Ching 359
megaphylla (Mett.) Iwats.
461
megaphylloides
Reed 481
melanophlebia (Copel.) Reed
$14
menisciicarpa_ (BI.) Iwats.
526
merrillii (Christ) Reed 529
micans (Brause) Reed 501
microcarpa (v.A.v.R.) Ching
371
microloncha (Christ) Reed
423
mindanaensis (Christ) Reed
557
mixta (Rosenst.) Reed 457
moniliformis (Tagawa &
Iwats.) Iwats. 519
morobensis (Copel.) Reed
501
motleyana (Hook.) Holtt. 381
multiauriculata (Copel.)
Reed 492
multilineata (Wall. ex Hook.)
Ching
var. malayensis 512
multiseta (Baker) Ching 349
multisora (C.Chr.) Reed 359
(Rosenst.)
munda (Rosenst.) Ching 503
muricata (Brause) Reed 391
mutabilis (Brause) Reed 494
nephrolepioides (C.Chr.) Reed
43]
nervosa (Klotzsch) Tryon 539
nitidula (Presl) Reed 430
notabilis (Brause) Ching 402
novae-hiberniae Holtt. 555
novoguineensis (Brause)
Reed 469
nymphalis (G.Forst.) Reed
Spy//
oblonga (Brause) Reed 503
obtusata (v.A.v.R.) Ching
362
obtusifolia (Rosenst.) Reed
503
oligodictua (Baker) Reed 380
oligolepia (v.A.v.R.) Ching
368
ophiura (Copel.) Reed 409
oppoOsitipinna (v.A.v.R.)
Ching 346
opulenta (Kaulf.) Fosb. 548
orbicularis (C.Chr.) Reed 548
palawanensis Reed 524
paleata (Copel.) Holtt. 380
pallescens (Brause) Ching
383
palustris Schott 375
var. squamigera Weath.
377
papilio (Hope) Iwats. 556
paraphysata Reed 481
parasitica (L.) Iwats. 559
paripinnata (Copel.) Reed
477
parishii (Bedd.) Panigrahi
535
pectiniformis (C.Chr.) Ching
355, 363
peltata (v.A.v.R.) Reed 529
peltochlamys (C.Chr.) Reed
495
pentaphylla (Rosenst.) Reed
514
perakensis (Bedd.) Reed 449
perglandulifera (v.A.v.R.)
Reed 454
perpilifera (v.A.v.R.) Reed
456
perpubescens (Alston) Reed
554
petrophila (Copel.) Reed 435
philippina (Presl) Ching 539
pilosiuscula (Racib.) Reed
S22
pilososquamata
Reed 481
(v.A.v.R.)
598
(Thelypteris)
pinnata (Copel.) Ching 375
platyptera (Copel.) Reed 369
plumosa (C.Chr.) Reed 360
plurifolia (v.A.v.R.) Reed 493
polycarpa (BI.) Iwats. 449
polyotis (C.Chr.) Reed 464
polyptera (Copel.) Reed 471
porphyricola (Copel.) Ching
472
procurrens (Mett.) Reed 560
producta (Kaulf.) Reed 460
prolifera (Retz.) Reed 387
protecta (Copel.) Reed 485
pseudoamboinensis
(Rosenst.) Panigrahi 558
pseudohirsuta (Rosenst.)
Reed 460
pseudoreptans (C.Chr.) Reed
522
pseudostenobasis (Copel.)
Reed 549
pterospora (v.A.v.R.) Reed
487
pubirachis Reed 365
pustulosa (Copel.) Reed 513
quadrangularis (Fée) Schelpe
557
quadriaurita
404
quadriquetra (v.A.v.R.)
Ching 406
rectangularis (Zoll.) Nayar &
Kaur 346
reederi (Copel.) Reed 456
regis (Copel.) Reed 434
repandula_ (v.A.v.R.) Reed
Si,
rhombea (Christ) Reed 529
rigida (Ridl.) Reed 484
rigidifolia (v.A.v.R.) Ching
369
riparia (Copel.) Reed 456
robinsonii (Ridl.) Ching 361
rodigasiana (T.Moore) Reed
(Christ) Reed
426
roemeriana (Rosenst.) Reed
499
rubicunda (v.A.v.R.) Iwats.
536
rubida (Hook.) Iwats. 532
rubrinervis (Mett.) Iwats. 531
sagittifolia (Bl.) Reed 487
Sagittifolioides (Copel.) Reed
471
sakayensis (Zeiller) Reed 392
salacifolia (Wall. ex Hook.)
Reed 535
samarensis
524
savaiensis (Baker) Reed 404
(Copel.) Reed
FLORA MALESIANA
(Thelypteris)
sessilipinna Reed 479
setigera Ching 349
setulosa Reed 481
sevillana Reed 418
simillima (C.Chr.) Iwats. 520
simozawae Tagawa 360, 370
simplicifolia (J.Sm.) Reed
490
simulans Ching 413
singalanensis sensu Holtt.
353
singalanensis (Baker) Ching
352
sogerensis (Gepp) Reed 428
spenceri (Christ) Reed 491
squamigera Ching 377
squamipes (Copel.) Reed 364
squamulosa (erron.) 377
stegnogramma (BI.) Reed 542
stenobasis (C.Chr.) Ching
549
stenodonta Reed 450
stenophylia (Baker) Reed 528
stereophylla (v.A.v.R.) Ching
366
stipellata (Bl.) Iwats. 492
strigosissima (Copel.) Reed
470
subalpina Reed 474
subappendiculata
Reed 420
subattenuata (Rosenst.) Reed
547
subdimorpha (Copel.) Reed
384
subfalcinella (v.A.v.R.) Reed
485
subglandulifera Ching 363
subimmersa Ching 547
subnigra (Brause) Ching 369
subpubescens (BI.) Iwats. 558
subulifolia Ching 494
subvillosa Ching 412
sumatrana (v.A.v.R.)
Tagawa & Iwats. 558
superba (Brause) Reed 420
supraspinigera (Rosenst.)
Reed 395
suprastrigosa (Rosenst.) Reed
457, 458
tephrophylla (Copel.) Reed
481
terminans (Hook.) Tagawa &
Iwats. 545
terrestris (Copel.) Reed 491
teuscheri (v.A.v.R.) Reed 381
todayensis (Christ) Reed 471
toppingii (Copel.) Iwats. 500
torresiana (Gaud.) Alston 348
triphylla (Sw.) Iwats. 534
(Copel.)
[ser. Il, ‘volar
(Thelypteris)
var. parishii (Bedd.) Iwats.
535
truncata (Poir.) Iwats. 429
tuberculata (Ces.) Ching 407
tylodes (Kunze) Ching 414
ugoensis Reed 460
uliginosa (Kunze) Ching 348
uniauriculata (Copel.) Reed
467
unidentata (Bedd.) Holtt. 359
unita (L.) Morton 478
uraiensis (Rosenst.) Ching
352
urdanetensis (Copel.) Reed
467
urophylla (Mett.) Iwats. 533
var. nitida (Holtt.) Iwats.
533
varievestita
401
verrucosa (Presl) Ching 547
verruculosa (v.A.v.R.) Reed
512
vestigiata (Copel.) Reed 453
vinosicarpa (v.A.v.R.) Ching
381
viridis (Copel.) Reed 418
viscosa sensu Holtt. 361
viscosa (Baker) Ching 355,
360
vulcanica (Baker) Reed 351
wagneri Fosb. & Sachet 545
wantotensis (Copel.) Reed
410
warburgii (Kuhn & Christ)
B.C.Stone 462
wariensis (Copel.) Reed 403
weberi (Copel.) Reed 460
williamsii Ching 450
xiphioides (Christ) Reed 525
zamboangana Reed 394
Thelypteris group ili, ix, xi, xii,
Xili, XiV, XX1, 257, 340
group 2 Ching 370
group 4 Ching 343
group 10 Ching 437
Thylacopteris tl, xvi
Thyrsopteridaceae 562
Thyrsopteridineae 562
Thyrsopteris 65, 69, 70, 72, 167,
562
Thysanobotrya v.A.v.R. 73, 115
arfakensis (Gepp) v.A.v.R.
118
Thysanosoria Gepp iii, xx, 255,
256, 258, 263
dimorphophylla Gepp 263
pteridiformis (Cesati) C.Chr.
263*
Tmesipteridaceae ti
(C.Chr.) Reed
1981]
Tmesipteris ii
Todea barbara Moore 61
Toppingia Deg. 343
Trichomanes ii, vii, Xv
sect. Crepidomanes vi
chinense L. 182
malayanum Roxb. ex Griff.
183
Trichopteris 562
falcata Llanos 146
Trigonospora Holtt. 331, 333,
338, 339, 340, 342, 373
calcarata (BI.) Holtt. 372*,
374
ciliata (Benth.) Holtt. 372*,
374, 375, 377
koordersii (Christ) Holtt. 374,
375
zeylanica Ching 374
Trigonospora group 340
Trogostolon ii, xviii
Index to scientific plant names
Ugena Cav. 39, 44
dichotoma Cav. 59
microphylla Cav. 47
polymorpha Cav. 562
semihastata Cav. 54
Vaginularia ili, XXi
Vallifilix Thouars 39, 45
Vittaria ili, vi, vill, Xxi
divergens (Hook. & Grev.)
Roxb. ex Griff. 233
interrupta Roxb. ex Griff.
254, 563
lunulata Roxb. ex Griff. 254,
563
parasitica Roxb. ex Griff.
244
Vittaria group Ill, ix, Xili, Xxi
Wagneriopteris Love & Love
370, 371
aoe
(Wagneriopteris)
angulariloba (Ching) Love &
Love 360
ogasawarensis 370
Wibelia auct. 184
Wibelia amboynensis
Kuhn 187
bipinnata Fée 191
denhami (Hook.) Kuhn 186
gracilis (BI.) Christ 190
javae Fée 193
oligophlebia (Baker) Christ 191
pinnata (Cav.) Bernh. ex Fée
193
Woodwardia il, Xvili
(Hook.)
Xiphopteris ii, xvii
Xyropteris Kramer 177, 178,
179, 197
stortil (v.A.v.R.) Kramer 197,
253
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