Malayan Forest Records Mo. 48

i

Series I: Ferns and Lycophytes, Vo

Flora
Edited by ftJi
B.S. Parris
R. Kiew
R.C.K. Chung
L.G. Saw
E. Soepadmo

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Fern and Lycophyte Families in Peninsular Malaysia
(families in Volume 1 in bold)

1. Lycopodiaceac

21. Dennsiaedliaceae

2. Selugindluceac

22. Parkeriaceae

3. Ophioglossaceae

23. Adiantaeeae

4. Psilotaceae

24. Cryptogrammaceae

5. Equisetaccae

25. Sinopteridaceae

6. Marattiaceae

26. Pteridaceae

7. Osmundaccae

27. Aspleniaceae

8. Hymenophyllaceae

28. Thelypteridaceae

9. Gleicheniaceae

29. Woodsiaccac


10. Diptcridaccac

30. Blechnaceae

11. Matoniaccae

3 1. Hypodeniatiaceae

12. Lygodiaceae

32. Dryopteridaccae

13. Schizaeaceae

33. Lomariopsidaceae

14. Marsileaceae

34. Nephrolepidaceae

15. Salviniaceae

35. Tcctariaccac

16. Plagiogyriaceae

36. Oleandraceae

17. Cihotiaccae


37. Davalliaceae

18. Cyatlicaceae

38. I.oxogrammaccae

19. Lindsaeaceae

39. Polypodiaceae

2(1. Saccolomataceae

40. Graminitidaceae


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FLORA
OF
PENINSULAR MALAYSIA
Series I: Ferns and Lycophytes

With compliments from
the Flora of Peninsular Malaysia Project
Forest Research Institute Malaysia
52109 Kepong
Selangor
Malaysia


Produced with the financial support o f
M INISTRY OF SCIENCE, TEC H N O LO G Y AND INNOVATION
GOV ERNM ENT O F MALAYSIA

Advisors to Flora o f Peninsular Malaysia: M.J.E. Coode (Royal Botanic Gardens,
Kew), J. Dransfield (Royal Botanic Gardens, Kew), D.J. Mabberley (Royal Botanic
Gardens, Kew and National Herbarium of the Netherlands, University of Leiden),
D.H. Nicolson (Smithsonian Institute), B.S. Parris (Fern Research Foundation, New
Zealand), J.F, Veldkamp (National Herbarium o f the Netherlands, University of
Leiden), W.J.J.O. de Wilde (National Herbarium of the Netherlands, University of
Leiden) and K M. Wong (University o f Malaya).



M alayan Forest Records N o. 48

FLORA
OF
PENINSULAR MALAYSIA
Series I: Ferns and Lycophytes

Volume 1

Edited by
B.S. Parris, R. Kiew, R.C.K. Chung,
L.G. Saw & E. Soepadmo

Forest Research Institute Malaysia
52109 Kepong, Selangor Darul Ehsan, Malaysia
2010

Ministry o f Natural Resources and Environment, Malaysia


€ Forest Research Institute M alaysia 2010
Date o f Publication: 5th January 2010
All enquiries should be forwarded to:
D irec tor-G eneral
Forest Research Institute M alaysia
52109 Kepong, Selangor Darul Ehsan
M alaysia
Tel:

603-6279 7000
Fax:
603-6 2 731314
Hom epage: m .gov.m y
Perpustakaan N egara M alaysia

Cataloguing-in-Publication Data

Flora o f Peninsular Malaysia: Series 1: Ferns and Lycophytes / edited by:
B.S. Parris, R. Kiew, R.C'.K. Chung, L.G. Saw & E. Soepadm o
(M alayan Forest Records No. 48)
ISBN 978-967-5221-24-8 (v. 1)
I. Ferns—M alaysia. 2. Lycopodiaceae—M alaysia. I. Parris, B.S.
II. Series.
587.3095951
Series Editor: S.S. Fee
M anaging Editor: Y.F. I Io

MS ISO 9001:2008 Certified
C over design: George Wong
Layout: Aslina Baharum
Front and back covers: Ctenopterella hlechnoides (Grev.) Parris (Photo: M. Y. Chew)

Printed in M alaysia by Straits Digital Sdn. Bhd., Subang Jaya, Selangor Darul Ehsan


CONTENTS
Page

FOREWORD

ACKNOWLEDGEMENTS

vii
ix

INTRODUCTION
Conspectus o f Orders, Families and Genera o f Ferns and Lycophytes of
Peninsular Malaysia (B .S. Parris )
Key to Families (B.S. Parris) History o f Botanical Collecting and Observations on Ferns and Lycophytes
in Peninsular Malaysia (B.S. Parris & R. K iew )
Species Assessment and Conservation in Peninsular Malaysia (L.S.L. Chua)

3
15
25
37

FAMILIES
2.
4.
5.
7.
11.
13.
17.
38.
40.

Selaginellaceae (K.M. Wong)
Psilotaccae (R. Kiew)

Equisetaceae (R. K iew )
Osmundaccac (R. Jam an)
Matoniaeeae (R. Jam an & Y. Umi K alsom )
Sehizaeaceae (R. Jam an & Y. Umi K alsom )
Cibotiaceae (A. T. N or-Ezzawanis)
Loxogrammaceae (R. Jam an)
Grammitidaceae (B.S. Parris)

Glossary (B.S. Parris)

49
87
91
95
101
107
119
123
131
207

ABBREVIATIONS FOR LOCALITIES

213

PLATES

215

INDEX TO SCIENTIFIC NAMES


235

INDEX TO VERNACULAR OR COMMON NAMES

249


FOREWORD

It is more than fifty years since the last fern flora for Peninsular Malaysia was published
(Holttum, 1954, revised 1968, Flora of Malaya 2 Ferns) and much has changed in the fern
world, especially with the advent of molecular techniques. This volume, the first in Series I
of the new Flora of Peninsular Malaysia, is a welcome addition to botanical publications not
only for Malaysia but also for the region.
The Flora of Peninsular Malaysia Project was initiated to document biodiversity by
providing reliable and accurate accounts o f plant families. Series I deals w ith the ferns and
lycophytes and Series II includes the gymnosperms and flow'ering plants. Being locally
based, the Flora includes precise ecological information, and correct scientific naming based
on examination of type specimens. The Flora has also incorporated four new' features not
included in many Floras, namely, distribution maps (unless the species is widespread and
common), conservation status for the species in Peninsular Malaysia, specimen identification
lists online (), and colour photographs. By compiling botanical
information for specific groups within one volume, the Flora aims to provide baseline
information that is essential for the management and conservation of plant biodiversity that
is Peninsular Malaysia’s natural heritage.
The Flora of Peninsular Malaysia Project, besides publishing the Flora, has w ider aims
by including strong elements of botanical exploration, particularly of poorly known or
unknown areas, assessment of conservation status, and most important, the training of the
next generation of plant taxonomists. To speed up the production of the Flora, collaboration

with both national and international herbaria is strongly encouraged.
Recent advances in molecular taxonomy have radically changed our view of the
relationships of ferns and fern allies. In fact, the group o f fern allies no longer exists,
lsoetaceae. Lycopodiaccae and Selaginellaceae are now placed among the lycophytes and
the rest of the fern allies (Psilotaceae and Hquisetaceae) are placed among the ferns. This first
volume includes an introductory chapter that lays out these changes for the 40 families, 137
genera and about 650 species of ferns and lycophytes that occur in Peninsular Malaysia.
This fern volume covers about a sixth o f the fern and lycophytes flora with accounts for
9 families, 21 genera and 100 species. Three families have been revised by Forest Research
Institute Malaysia staff, five by taxonomists in three local universities and with international
collaboration. The editors are particularly grateful to Dr B.S. Parris for her guidance as chief
editor for Series 1 (Ferns and Lycophytes), the publication of this volume owes much to her
editorial input and her contribution of the Grammitidaceae.
I congratulate the contributors and editors on ihe timely production of this landmark
volume and look forward to future volumes.

Datn’ DrAbd. Latif Mohmod
Director-General
Forest Research Institute Malaysia

vii


ACKNOWLEDGEMENTS

The successful initiation of the Flora of Peninsular Malaysia is a result of the Malaysian
Government's foresighted policy on biodiversity conservation. We are greatly indebted to
the Ministry of Science, Technology and Innovation (MOSTI) through the National Council
for Scientific Research and Development (MPKSN) for providing generous financial
support to enable the employment and training of young taxonomists, field collection and

herbarium visits by both local and overseas collaborators, and the publication of the Flora
and associated publications. The project was funded under a special science fund (Project No.
01-04-01-0000 Khas 2) entitled “Safeguarding the Forest I )iversity of Peninsular Malaysia”.
It has two components: A Flora o f Peninsular Malaysia and Conservation Monitoring of
Rare and Threatened Plants in Peninsular Malaysia. Special thanks are due to the past and
present Director Generals of the Forest Research Institute Malaysia, Datuk Dr Ilj. Abdul
Razak Mohd. Ali and Dato' Dr Abd. Latif Mohmod, respectively, for their strong institutional
commitment and to the members of the Advisory Bourd for their support.
The Flora has greatly benefited from the support of the Directors and Curators of
herbaria at BKF, BM, BO, CAL, E, K, KLU, L. SAN, SAR, SING and UKMB for assisting
in herbarium visits, the loan o f herbarium specimens, access to types and library material and
databases.
We are also greatly indebted to botanists from local universities. Prof K.M. Wong (KLU),
Razali Jaman (UKMB) and Dr Umi Kalsoni Yusof (UPM), for their contribution of family
revisions in this volume.
To Zainal Mustafa (KLU) we are indebted for his elegant botanical illustrations and to
Razali Jaman for illustrating his account, to Aslina Baharum for her competent, error-free
page-setting of this volume and compiling the indices and to Flamidah Mamat for preparing
the regional maps.
Dr L.S.L. Chua has been the guiding force in conservation assessments of species
and special thanks are due to A.T. Nor-Ezzawunis (Mutoriiaceue, Osmundaeeae and
Schizaeaceae), C.L. Lim and T.L. Yao (Grammitidaccac), A.R. Rafidah and A.R. UmmulNazrah (Selaginellaceae) for preparing materials for the maps and conservation assessments.
C.L. Lim and T.L. Yao assembled the database of specimens examined and T.L. Yao
managed the photo library. D. Filer, Oxford University, and BRAHMS (Botanical Research
And Herbarium Management System) greatly facilitated the databasing and preparation of
maps.
The Flora could not function without the technical support of the KEP botanists,
herbarium and field staff to whom we owe a great debt of gratitude and finally to the project
staff without whose assistance the project would not be able to achieve its goals.


B.S. Parris
R. Kiew
R.C.K. Chung
L.G. Saw
F.. Soepadmo

ix


FLORA OF PENINSULAR MALAYSIA

Conspectus of Orders, Families and Genera of
Ferns and Lycophytes of Peninsular Malaysia
B.S. Parris
Fern Research Foundation,
Bay of Islands, New Zealand

Conspectus of Orders, Families and Genera
In the four decades since llolttum’s publication of an account of ihe ferns of Peninsular
Malaysia and Singapore (Holttum 1968a) there have been many changes in fem classification
at family, genus and species levels. In particular, the recent phytogenies utilising both
morphological and molecular data, summarised by Smith el at. (2006), have revolutionised
our ideas of relationships amongst the lower vascular plants.
Pryer et al. (2001) demonstrated that ferns and fern allies, or pteridophyles, are a
paraphyletic group, showing that lycophytes (the “fern allies" Isoetaceae. Lycopodiaceae
and Selaginellaceae) are a sister clade to the one containing both ferns and seed plants, and
that the “ fem allies” Equisctaccae and Psilotaceae belong w-ith ferns, rather than with the
lycophytes.
The “ fern allies” are included in the current treatment, unlike Holttum (1968a), w'ho
omitted them, us was common in floras of the time, for example Copeland (1960). Piggott

(1988), like Holttum, included only ferns.
The circumscription of orders, families and genera is based on that o f Smith etal. (2006).
The differences between it and the arrangement adopted here arc discussed later. Numbering
of families in the Flora follows Tabic 1.
Table I. Orders, families and genera in Peninsular Malaysia
CLASS 1. LYCOPSIDA

CLASS 2. PSILOTOPSIDA

ORDER 1. LYCOPODIALES

ORDER 3. OPHIOGLOSSALES

FAMILY 1. LYCOPODIACEAE
1. Huperzia Bcmh.
2. Lycopodiclla Holub
3. Lycopodium L.

FAMILY 3. OPIIIOGLOSSACEAE
1. Botrychium Sw.
2. Helminthostachys Kaulf.
3. Ophioglossum L.
Reference: Wiefifering (1964)

ORDER 2. SELAGINELLALES

ORDER 4. PSILOTALES

Family 2. SELAGINELLACEAE
1. Selaginella P.Beauv.

Reference: Wong (1983), 2010

FAMILY 4. PSILOTACEAE
1. Psilotum Sw.
Reference: Kiew (2010)

3


FERNS & LYCOPIIYTES VOL. 1 (2010)

1. Dicranoptcris Bemh.
2. Diplopterygium (Diels) Nakai
3. Gleichenia Sm.
4. Sticherus C.Presl

CLASS 3. EQUISETOPSIDA
ORDER 5. EQUISETALES
FAMILY 5. EQU1SETACEAE

FAMILY 10. DIPTER1DACEAE
1. Cheiropleuria C.Presl
Reference: Laferriere (1998a)
2. Dipteris Reinw.

Reference: Laferri£re ( 1998b), Kiew (2010)
1. Equisetum L.

CLASS 4. MARATTIOPSIDA
FAMILY 11. MATOMACEAE

Reference: Kato (1998),
Jaman &Umi Kalsom (2010)
1. Matonia R.Br.

ORDER 6. MARATTIALES
FAMILY 6. YIARATTIACEAE
1. Angiopteris I lolfm.
2. Chrlstensenia Maxon
Reference: Rolleri (1993)
3. Ptisana Murdock
Reference: Maideen et al. (2002)

ORDER 10. SCHIZAEALES
FAMILY 12. LYGODIACEAE
Reference: Holttum (1959b)
1, Lygodium Sw.
FAMILY 13. SCHIZAEACEAE
Reference: Holttum (1959c),
Jaman &Umi Kalsom (2010)
1. AcUnostachys Wall.
Hook.
2. Schizaea Sm.

CLASS 5. POLYPODIOPSIDA
ORDER 7. OSMUNDALES
FAMILY 7. OSMUNDACEAE
Reference: Bidin (1984), Jaman (2010)
1. Osmunda L.

ORDER 11. SALVINIALES

FAMILY 14. MARSILEACEAE
1. Marsilea L.

ORDER 8. HYMENOPHYLLALES

Reference: Johnson (1988)
FAMILY 8. HYMENOPHYLLACEAE
References: Ebihara et al. (2006), Ivvatsuki
(1985)
1. Abrodietyum C.Prcsl
2. C'allistopteris Copel.
3. Ceplialonianes C.Presl
4. Crepidoinanes C.Presl
5. Didvnioglossum Dcsv.
6. Hymenophyllum Sm.
7. Vandenboschia Copel.

FAMILY 15. SALVINIACEAE
1. Azolla Lam.
Reference: Saunders (1998)
2. Salvinia Seg.

ORDER 12. CYATHEALES
FAMILY 16. PLAGIOGYRIACEAE
Reference: Zhang & Nooteboom (1998)
1. Plagiogyria (Kun/.e) Mett.

ORDER 9. GLEICHENIALES
FAMILY 17. CIBOTIACEAE
Reference: Holttum (1963a),

Nor-Ezzawani (2010)
1. Cibolium Kaulf.

FAMILY 9. GLEICIIENIACEAE
Reference: llolttum (1959a)

4


ORDERS, FAMILIES, GENERA (PARRIS)

FAMILY 18. CYATHEACEAE
References: 1lolttum ( 1963b). I lolttum &
Molesworth Allen (1967)
l. Cyalhea Sm.

2. Syngramma J.Sm.
Reference: I lolttum (1975)
3. Taenitis Willd. ex Schkuhr
References: Holttum (1968b, 1975)

ORDER 13. POLYPODIALES

FAMILY 27. ASPLENIACEAE
1. Asplenium L.
Reference: Holttum (1974)

FAMILY 19. LINDSAEACEAE
Reference: Kramer (1971)
1. Lindsaca Dryand. ex Sm.

2. Splieiiomeris Maxon
3. Tapeinidium (C.PresI) C.C'hr.

FAMILY 21. DENNSTAEDT1ACEAE
1. Dennslaedtia Bemh.
2. Histiopteris (J.Agardli) J.Sm.
3. Hypolepis Bemh.
Reference: Brownsey (1987)
4. Microlcpia C.PresI
5. Monachosorum Kunze
6. Pacsia A.St.-Hil.
7. Pteridium Gled. ex Scop.

FAMILY 28. THELYPTERIDACEAE
Reference: I lolttum (1981)
1. Ampelopteris Kunze
2. Amphineuron Holttum
3. Chingia Holttum
4. Christella H.L6v.
5. Coryphopteris Holttum
6. Cyclosorus Link
7. Macrothclypteris (H.lto) Ching
8. Mesophlebion Holttum
9. Metathelypteris (H.lto) C'hing
10. Parathelypteris (H.lto) Ching
11. Pneuinatopteris Nakai
12. Pronephrium C.PresI
13. Pseudophegopteris Ching
14. Sphaerostephanos J.Sm.
15. Trigouospora Holttum


FAMILY 22. PARKERIACEAE
1. Acrostichum L.
2. Ceratnpteris Rrongn.

FAMILY 29. WOODSIACEAE
1. Acystopteris Nakai
2. Athyrium Roth

FAMILY 23. ADIANTACEAE
1. Adiantum L.
2. Antrophyum Kaulf.
3. Haplopteris C.PresI
4. Monogramma Comm, ex Schkuhr

4. Deparia Hook. & Grev.
Reference: Kato (1984)
5. Diplaziopsis C.Chr.
6. Diplazium Sw.

FAMILY 20. SACCOLOMATACEAE
1. Saccoloma Kaulf.

3. Callipteris Rory

FAMILY 30. BLECHNACEAE
1. Blechnum L.
Reference: Chambers & Farrant (2001)
2. Brainea J.Sm.
3. Stcnochlacna J.Sm.

4. Woodwardia Sm.

FAMILY 24. CRYPTOGRAMMACEAE
1. Coniogramme Fee
FAMILY 25. SINOPTERIDACEAE
1. Cheilanthes Sw.
2. Doryopteris J.Sm.
3. Hcmionitis L.
4. Pityrogramma Link

FAMILY 31. HYPODEMATIACEAE
1. Didymochlaena Desv.
2. Hypodematiuni Kunze
Reference: Holttum (1985)

FAMILY 26. PTERIDACEAE
1. Pleris L.

5


FERNS & LYCOPHYTES VOL. i (2010)

3. Leucostegia C. Presl
Reference: Nooteboom (1998a)

FAMILY 36. OLEANDRACEAE
1. Oleandra Cav.

FAMILY 32. DRYOPTERIDACEAE

1. Acrophoms C.Presl
2. Arachniodes Rlume
3. Rolbitis Schott
References: Hennipman (1978), Holttum
(1979)
4. Ctenitis (C.Chr.) C.Chr.
Reference: Holttum ( 1991a)
5. Dryopteris Adans.
6. Elaphoglossum Schott ex J.Sm.
Reference: Holttum (1978a)
7. Lomagramma J.Sm.
Reference: Holttum ( 1978b)
8. Peranema D.Don
9. Polystichum Roth
10. Teratophyllum Mett. ex Kuhn
Reference: Holttum (1978c)

FAMILY 37. DAVALLIACEAE
Refeierice: Nooteboom (1998b)
1. Davallia Sm.

FAMILY 38. LOXOGRAMMACEAE
Reference: Jaman (2010)
1. Loxogramme (Blurne) C.Presl
FAMILY 39. POLYPODIACEAE
Reference: I lovenkamp et al. (1998)
1. Aglaomorpha Schott
2. Belvisia Mirb.
3. Cliristopteris Copel.
4. Drynaria J.Sm.

5. Goniophlebium C.Presl
6. Lecanopteris Reinw.
7. I .emmaphyllum C.Presl
8. Lepisorus Ching
9. Leptochilus Kaulf.
10. Microsoruni Link
11. Platycerium Desv.
12. Pyrrosia Mirb.
13. Selliguca Bory
14. Thylacopteris Kunze ex J.Sm.

FAMILY 33. LOMARIOPSIDACEAE
1. Cyclopeltis J.Sm.
Reference: Holttum ( 19 9 1b)
2. Lomariopsis Fee
Reference: Holttum (1978d)
FAMILY 34. NEPHROLEPIDACEAE
Reference: Hovenkamp & Miyamoto (2005)
1. Nephrolepis Schott

FAMILY 40. GRAMMITIDACEAE
Reference: Parris (1986, 2007, 2010)
1. Acrosorus Copel.
2. Calymmodon C.Presl
3. Chrysogrammitis Parris
4. Ctenopterella Parris
5. Dasygrammitis Parris
6. Oreogrammitis Copel.
7. Prosaptia C.Presl
8. Radiogranimitis Parris

9. Scleroglossuni Alderw.
10. Themeliuni (T.Moore) Parris
11. Tomophyllum Parris
12. Xipliopterella Parris

FAMILY 35. I EC 1AR1AC EAE
Reference: Holttum (1991c)
1. Arthropteris J.Sm.
Reference: Holttum (1966)
2. Hcterogonium C.Presl
3. Pleocnemia C.Presl
4. Ptcridrys C.Chr. & Ching
5. Tectaria Cav.
Reference: Rusea et al. (2004)

6


ORDERS, t AMILIES. GENERA (PARRIS)

Differences between Smith et al. (2006) and the family system adopted for the Flora of
Peninsular Malaysia are as follows:
1.

Ptcridaceae has been divided into five monophyletic families, Parkeriaceae, Adiantaccae,
Cryptogrammaceae, Sinopteridaceae and Pteridaceae, for the convenience of Flora users.
Smith et o/.’s (2006) concept o f Pteridaceae includes 56 Peninsular Malaysian species
and, given that genera and species are nowadays arranged alphabetically for both floras
and herbaria, it seems practical to divide them into five families. The majority of modem
fern floras do not use such a broad concept o f Pteridaceae as do Smith et al. (2006).


2.

Hypodematiaceae is separated from Dryopteridaceae. Smith et al. (2006) noted
that the inclusion of Didymochlaena, Hypodematium and Leucastegia may render
Dryopteridaceae paraphyletic, but tentatively included them in Dryopteridaceae pending
further studies. Schuetlpelz & Pryer (2007) clarify the affinities of the genera and their
relationship to Drypteridaceac.

3.

Nephrolcpidaceae is recognised as a family distinct from Lomariopsidaceae. It contains
one genus. Nephrolepis. which is tentatively included in Lomariopsidaceae by Smith et
al. (2006) pending possible future recognition. It is treated as a monogeneric family by
Kramer in Kramer & Green (1990) on the basis of its very distinct anatomical and spore
characters, and is recognised as such in some recent floras and floristic accounts.

4.

Polypodiaccae has been divided into three families, Loxogrammaceae. Polypodiaceae
and Grummitidaceac. Polypodiaccae, in the sense of Smith et al. (2006), is sister to
Davalliaccae. Within Polypodiaceae. Loxogrammaceae is basal to the rest of the family,
while Grammitidaceae nests within one of the three major eludes (Ranker et al. 2004:
Schneider et al. 2004). Grammitidaceae is morphologically distinct from Polypodiaceae
in a number of characters, including veins free (in most taxa), lack of scales on fronds,
simple unjointed hairs ("setae”) on fronds (in most taxa). sporangial stalks one cell
wide, spores chlorophyllous, globose-tetrahedral (in most taxa) and gametophytes
ribbon-shaped. Smith et al. (2006) recognise "grammitid ferns” us an unranked informal
taxon. For the Flora of Peninsular Malaysia it is more practical to treul this taxon as a
family rather than treat it within Polypodiaceae, for the same reason of convenience that

Pteridaceae is divided into five families. Polypodiaceae (excluding Loxogrammaceae)
has 64 species and is the largest fern family in Peninsular Malaysia; Grammitidaceae,
with 52 species, is the third largest fern family. Combination of the two would be very
unwieldy for users of the Flora and for herbarium curalion. Separation o f the two leads
logically to the separation of Loxogrammaceae from Polypodiaceae. Grammitidaceae
is currently recognised in Old World floras, while Loxogramme is usually treated in
Polypodiaceae.

The similarities and differences between Holtlum ( 1968a). Piggott (1988). various Flora
Malesiana accounts and the system adopted here are discussed below. Table 2 shows the
families used here, and their equivalents in both I lolttum (1968a) and Piggott (1988); because
the Flora Malesiana fern and lycophyte accounts are incomplete they are not included in this
table.

7


FERNS & LYCOPHYTES VOL. I (2010)

Table 2. Flora of Peninsular Malaysia fern and lycophyte families and their treatment here
and in llolttum (1968a) and Piggott (1988)
Here_______________________

llo lt tu m

Piggoo__________________

1. LYCOPODIACEAE

excluded


excluded

2. SELAGINELLAC EAE

excluded

excluded

3. OPHIOGLOSSACEAE

Ophioglossaceae

Ophioglossaceae

4. PS ILOTAC EAE

excluded

excluded
excluded

5. EQUISETACEAE

excluded

ft. MARATTIACEAE

Marattiaceae


Marattiaceae

7. OSMUNDACEAE

Osmundaecac

Osmundaecac

8. HYMENOPHYLEACEAE

Hymenophylluceae

Hymenophyllaceae

9. GLEICHENIACEAE

Gleiehoniaceae

Gleicheniaecac

l«. DIPTERIDACEAE

Polypodiaceae

Polypodiaceae

11. MATONIACEAE

Matoniaceae


Matoniaceae

12. LYGODIACEAE

Schizacaccac

Schizacaccac

13. SCHIZAEACEAE

Sclii/aeaeeae

Schizacaccac

14. MARSII.EACEAE

Marsileaceae

Marsileaceae

15. SALVINIACEAE

Salviniaceae, Azollaceae

Salviniaceae, Azollaceae

16. PLAGIOGYRIACEAE

Plagiogyriaceae


Plagiogyriaceae

17. CIBOTIACEAE

Dickson iaceae

Cyathcaceae

1 8 . f ' Y A T 11E A G F A F .

C’yalheaceae

Cyatheaceae

19. LINDSAEACEAE

Dcnnstaedtiaccac

Lindsaeaceae

20. SACCOLOMATACEAE

Dennstaedtiaceae

Dennstaedtiaceae

21. DENNSTAEDTIACEAE

Dennstaedtiaceae


Dennstaedtiaceae. Hypolepidaccac

22. PARKERIACEAE

Adiantaccac, Dennstaedtiaceae

Parkcriaccae. Ptcridaceae

23. A D1ANTACEAE

Adiantaccae

Adiantaccac. Vittariaccac

24. CRYPTOGRAMMACF.AE

Adiantaceae

Hcmionilidaceae

25. SINOPTF.RIDACEAE

Adiantaceae

Hcmionilidaceae, Sinopteridaceae

26. PTERIDACEAE

Adiantaceae. Dennstaedtiaceae


Hemionitidaccae, Pteridaecac,
Taenilidaccae

27. ASPLENIACEAE

Dennstaedtiaceae

Aspleniaceac

28. THF.I A’PTF.R IDACE A E

Thelypteridaceae

Thelypteridaceae

29. WOODSIACEAE

Dennstaedtiaceae

Athyriaceae

30. BLECHNACEAE

Dennstaedtiaceae

Blcchnaccac

31. HYPODEMAT1ACEAE

Dennstaedtiaceae


Aspidiaceae. Davall iaceae

32. DRYOPTERIDACEAE

Dennstaedtiaceae

Aspidiaceae, I .omnriopsidaccac

33. I OM A RIOPS1 DACE A E

Dennstaedtiaceae

Aspidiaceae. Lomariopsidaccae

34. NEPHROLEPIDACEAE

Dennstaedtiaceae

Nephrolcpidaceae

35. TECTARIACEAE

Den iistaedt iaceae

Aspidiaceae. Nephrolcpidaceae

36. OLEANDRACKAE

Dennstaedtiaceae


Oleandraceae

37. DAVALLIACEAE

Dennstaedtiaceae

Davalliaceae

38. I.OXOG R AM M ACE A E

Polypodiaceae

Polypodiaceae

39. POLY PODIACEAE

Polypodiaceae

Polypodiaceae

40. GRAM Mi l l DACE AE

Cirammitidaceae

Grammitidaceae

8



ORDERS, FAMILIES. GENERA (FARRIS)

As mentioned previously, Equisetaceae, Lycopodlaceae, Psilotaceac and
Selaginellaceae were not included in Holttum (1968a) and Piggott (1988). Equisetaceae has
been revised for Flora Malesiana (Laferriere, 1998b).
The primitive fern families Ophioglossaceae, Marattiaeeae. Osmundaceae,
Hymenophyliaceae, Gleicheniaceae, IMatoniaceac and Plagiogyriaceae remain unchanged
here. Gleichcniaceac (Holttum. 1959a) and Matoniaceae (Kato, 1998) have been revised for
Flora Malesiana.
Diptcridaceae here includes Cheimpleuria and Dip ter is, both treated as primitive generu
o f Polypodiaceae in Holttum ( 1968a) and Piggott (1988). Cheimpleuria has been revised for
Flora Malesiana (Laferriere, 1998a) as the single genus in Cheiropleuriaceae.
Schizaeaceae is here treated in a narrow sense, with Lygodium separated into the
monogeneric family Lygodiaceae. Schizaeaceae, including Lygodiaceae, has been revised
for Flora Malesiana (Holttum, 1959b,c).
The heterosporous water ferns Mars ilea (Marsileaceae). Azolla (Azollaceae) and Salvinia
(Salviniaccac) were previously (e.g. Holttum, 1968a) considered to be very advanced. Here
they are included with more primitive families, Azolla is now included in Salviniaceae.
Azollaceae has been revised for Flora Malesiana (Saunders. 1998).
Cibotiurn, previously included in Dicksoniaceae, is here separated into the monogeneric
family Cibotiaccac. It hus been revised for Flora Malesiana in C’yatheaceae (Holttum.
1963b). Cyathea is treated here as a monogencric family. Cyathcaceae.
Holttum ( 1968a) maintained Dennstacdtiaccac as a large family with 11 subfamilies, each
of which is treated here at family level or divided into families. In several cases, genera are
treated in families not equivalent to 1lolttum’s subfamilies. Piggott (1988) divided 1iolttum’s
broadly based Dennstaedtiaceac into numerous families, some of which are not maintained
in the present system.
HoIttunTs Dennstaedtiaceae subfam. Lindsaeoideae is treated here as family
Lindsaeaceae. It has been revised for Flora Malesiana as "Lnuisaea group” (Kramer.
1971).

Saccolomataceae contains only Saccoloma. treated by Holttum ( 1968a) as Orthiopteris
in his Dennstaedtiaceae subfam. Dennstaedtiodeae.
Dennstaedtiaceae, as defined here, contains the genera treated in HoIttunTs (1968a)
Dennstuedliuceue subfam. Dennstaedtiodeae, with the exception of Saccoloma (syn.
Orthiopteris), together with Histiopteris and Pteridiwn, placed by Holttum in his
Dennstaedtiaceae subfam. Pteridoideae.
Parkeriaceae here contains Acrostichum, regarded by Holttum (1968a) as a member
of Dennstaedtiaceae subfam. Pteridoideae, and Ceratopteris, treated by Holttum in
Adiantaceae.
Here, Adiantaceae contains Adiantum and the vittarioid terns Antrophyum, Haplopteris
(syn. Vittaria) and Monogramma. HoIttunTs (1968a) Adiantaceae also contained genera
treated here in Cryptogrammaceae (Coniogramme), Pleridaceae (Svngramma, Taenitis) and
Sinopteridaceae (Cheilanthes, Doiyopteris, Hemionitis, Pityrogramma).
Pleridaceae here contains only Pteris, Svngramma and Taenitis. HoIttunTs (1968a)
Dennstaedtiaceae subfam. Pteridoideae included Acrostichum (now Parkeriaceae),
Histiopteris and Pteridium (now Dennstaedtiaceae) and Stenochlaena (now Bleehnaceac),
as well as Pteris.
HoIttunTs (1968a) Dennstaedtiaceae subfam. Asplenioideae is treated here as family
Aspleniaceae. and his Dennstaedtiaceae subfam. Athyrioideae is now family Woodsiaccae.
Thelypteridaceae remains unchanged from Holttum (1968a). It has been revised for

9


ILRNS & LYCOPHYTES VOL. 1 (2010)

Flora Malesiana (Holttum, 1981).
Bleclinuceae liere contains Holttum "s ( 1968a) Dennstaedtiaceae subfam. Blechnoideae,
with the addition o f Stenochlaena, which he placed in Dennstaedtiaceae subfam.
Pteridioideae.

Hypodematiaceae here contains Didymochlaena from Holttum’s (1968a)
Dennstaedtiaceae subfam. Dryopteridoideae, Hypodematium from his subfam. Tectarioideae
and Leucostegia from his subfam. Davallioidcae. Leucostegia has been revised for Flora
Malesiana (Nooteboom 1998a) in Davalliaceae.
Dryopteridaceae as defined here is much larger than Holttum’s Dennstaedtiaceae subfam.
Dryopteridoideae. which contained only Aavphonts, Arachniodes (syn. Polystichopsis),
Didymochlaena, Dryopleris, Peranema (syn. Diacalpe) and Polystichum. Apart from
Didymochlaena, now in Hypodematiaceae, these genera are included here in Dryopteridaceae,
together with Ctenitis from Holttum’s (1968a) subfam. Tectarioideae and Hoibit is,
Elaphoglossum, Lomagramma and Teratophyllum from bis subfam Lomariopsioidcac.
Ctenitis has been revised for Flora Malesiana (Holttum, 1991a) in “Tectaria group". Bolbitis
(Hennipman. 1978), Elaphoglossum (Holttum, 1978a), Lomagramma (Holttum. 1978b)
and Teratophyllum (Holttum. 1978c) have been revised for Flora Malesiana. together with
Lomariopsis (Holttum, I978d). as “Lomariopsis group".
I.omariopsidaceae here is a smaller family than Holttum’s Dennstaedtiaceae subfam.
Lomariopsidoideae, containing only Cyclopeltis, formerly in Dennstaedtiaceae subfam.
Tectarioideae and revised for Flora Malesiana (Holttum, 1991b) in “ Tectaria group", and
Lomariopsis, revised for Flora Malesiana (Holttum, 1978d) in “Lomariopsis group”.
Ncphrolepidaccae here containsNephrolepis, from Holttum’s ( 1968a) Dennstaedtiaceae
subfam. Olcandroideae.
Tectariaceae here differs from Holttum’s ( 1968a) Dennstaedtiaceae subfam. Tectarioideae
by the transference of Ctenitis to Dryopteridaceae, Hypodematium to Hypodematiaceae (see
above), and Cyclopeltis to Lomariopsidaceae (see above), together with the addition of
Arthwpteris from Dennstaedtiaceae subfam. Oleandroidcae. Tectariaceae in part was revised
for Flora Malesiana (Holttum. 1991c) as **Tectaria group”.
Oleandraceae here contains only Oleandra Iogether with Arthropteris and Nephrolepis,
it formed Holttum’s ( 1968a) Dennstaedtiaceae subfam. Olcandroideae. Arthropteris is now
in Tectariaceae and Nephrolepis is in Nephrolepidaceae.
Davalliaceae is similar to Holttum’s (1968a) Dennstaedtiaceae subfam. Davallioideae,
with the transference of Leucostegia to Hypodematiaceae. Davalliaceae has been revised for

Flora Malesiana (Nooteboom, 1998b).
Loxogrammaceae has been separated from Holttum’s Polypodiaceae. Apart from the
removal o f Cheiropleuria andDipteris to Dipteridaceae and Loxogramme to Loxogrammaceae,
his Polypodiaceae has the same circumscription as in this account.
(ii'am m itidaceae is also circumscribed by Holttum ( 1968a) in the same way as here.

10


ORDERS, FAMILIES. GENERA (PARRIS)

References
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Malay. 9, 1:28-31.
Brownsey, P.J. 1987. A review o f the fern genus Hypolepis (Dennstaedtiaceae) in the
Malesian and Pacific regions. Blumea 32: 227 276.
Chambers, T.C. & Farrant. P.A. 2001. Revision of Blechnum (Blechnaceae) in Malesia.
Blumea 46: 283-350.
Copeland, E.B. 1960. Fern Flora o f the Philippines. Bureau of Printing, Manila.
Ebihara, A., Dubuisson, J.-Y., Iwatsuki. K., Hennequin, S. & Ito. M. 2006. A taxonomic
revision of Uymenophyllaceae. Blumea 5 1: 221 -280.
Hennipman, E. 1978. Bolhitis, in Lomariopsis group. FI. Malesiana 2, 1:314 330.
I lolttum. R.E. 1959a. Gleieheniaceae. FI. Malesiana 2, 1: 1-36.
Holttum, R.E. 1959b. Lygodium, in Sehizaeaccac. FI. Malesiana 2, I: 44-61.
1lolttum, R.E. 1959c. Schizaea, in Schizaeaceae. FI. Malesiana 2. I: 37-44.
Holttum, R.E. 1963a. Cibofium, in Cyatheaceae. FI. Malesiana 2, 1: 164-166.
Holttum. R.E. 1963b. Cyathea, in Cyatheaceae. FI. Malesiana 2. 1: 65-158.
Holttum, R.E. 1966. The genus Arthrapteris J.Sm. in Malesia. Blumea 14: 225 229.
Holttum. R.E. 1968a. Revised Flora o f Malaya. Vol. 2, Ferns. 2nd ed. Government Printing
Office. Singapore.

Holttum, R.E. 1968b. A re-definition of the fern-genus Taenitis Willd. Blumea 16: 87-95.
Holttum. R.E. 1974. Asplenium L.. sect. Thamnopteris Presl. Card. Bull. Sing. 21:
143-154.
Holttum, R.E. 1975. A comparative account of the fern-genera Syngramma J.Sm. and
Taenitis Willd., with discussion of their relationships to each other and to other genera.
Kew Bull. 30: 327-343.
Holttum, R.E. 1978a. Elaphoglossum, in Lomariopsis group. FI. Malesiana 2, 1: 289-314.
Holttum. R.E. 1978b. Lomagramma, in Lomariopsis group. FI. Malesiana 2, 1: 276-289.
Holttum, R.F.. 1978c. Teratophyllum. in Lomariopsis group. FI. Malesiana 2, 1: 265-276.


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Holttum, R.E. I978d. Lomariopsis, in LomariopsLs group. FI. Malesiana 2. 1: 258 265.
Holttum, R.E. 1979. A new look at the fern-genus Bolbitis in Malaya. Malay. Nat. J. 33:
79 82.
Holttum, R.E. 1981. Thelypteridaceae. FI. Malesinna 2, 1: 331-599.
Holttum, R.E. 1985. Two new species of Tectaria from limestone in Peninsular Malaysia,
with comments on some other species. Gant Hull. Sing. 38: 145 148.
Holttum, R.E. 1991a. Ctenitis, in Tectaria group. FI. Malesiana 2, 2: 21-36.
Holttum. R.E. 1991b. Cyclupeltis, in Tectaria group. FI. Malesiana 2, 2: 116-119.
Holttum. R.E. 1991c. Tectaria group in part. FI. Malesiana 2. 2: 1-132.
Holttum. R.E. & Molesworth Allen, B. 1967. The tree-ferns of Malaya. Gard. Bull.
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Hovenkatnp, P. & Miyamoto, F. 2005. A conspectus of the native and naturalized species of
Nephmlepis in the world. Blumea 50: 279-322.
Hovenkamp, H.P., Bosnian. M.T.M., Hennipman. E.. Nooteboom, H.P., Roedl-I mder. G.
& Roos, M.C. 1998. Polypodiaceae. FI. Malesiana 2, 3: 1-234.
Iwatsuki, K. 1985. The Hymenophyllaeeae of Asia, excluding Malesia. J. Fac. Sci.,
Univ. Tokyo 3, 13: 501-551.

Johnson, D M 1988. Marsilea scalaripes. a new member of Marsilea section Clemvs from
the Asian tropics. Amer. Fern J. 78: 68-71.
Jaman. R. 2010. Loxogrammaceae. FI. Fen. Malay. I, 1: 123-130.
Jaman, R. 2010. Osmundaceae. FI. Pen. Malay. I. 1: 95 99.
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Jaman, R. & IJnii Kalsom, Y. 2010. Schizaeaceae. FI. Pen. Malay. I, 1: 107-117.
Kato, M. 1984. A taxonomic study of the Athyrioid fern genus Deparia. with main
reference to the Pacific species. J. Fac. Sci., Univ. Tokyo 3, 13: 375—429.
Kato, M. 1998. Matoniaceae. FI. Malesiana 2. 3: 289-294.
Kiew, R. 2010. Equisetaccac. FI. Fen. Malay. I, 1: 91-94.
Kiew. R. 2010. Psilotaceae. FI. Fen. Malay. I. 1: 87-90.

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ORDERS, FAMILIES. GENERA (PARRIS)

Kramer, K.U. 1971. Lindsaea group. FI. Malesiana 2,1:177-254.
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Laferriere, J.E. 1998a. Cheiropleuriaceae. FI. Malesiana 2,3: 285 28ft.
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Zhang, X.C. & Nooteboom. H.P. 1998. Plagiogyriaceae. FI Malesiana 2, 3: 295-316.

14


FLORA OF PENINSULAR MALAYSIA

Key to Families
B.S. Parris
Fern Research Foundation,
Ray of Islands, New Zealand
1.

Leaves with single unbranched vein, sometimes bilid when subtending synangia, or vein
absent but plants not floating: sporangia or synangia solitary in leaf axils or aggregated
into cone-like structures.......................................................................................................2
Fronds with branched veins, or fronds reduced to vascular tissue except lor apex with
lobes bearing sporangia, or veins absent and plants floating; sporangia in clusters of
various shapes or covering under surface of frond or in sporocarps..................................5

2.

Stems and branches grooved and jointed, leaves in whorls at each node, fused at base to
form a sheath; sporangia aggregated into cone-like structures................5. Equisetaccae
Stems and branches (if present) not grooved, not jointed, leaves spirally or oppositely
arranged, sporangia or synangia solitary in leaf axils..................................................... 3


3.

Leaves widely spaced, up to 2 mm long; sporangia fused to form 2-3-lobed synangia.
................................. ,...................................................................................... 4. Psilotaceae
Leaves closely spaced, usually more than 2 mm long, sporangia separate, not fused to
form synangia........................................................................................................................ 4

4.

Leaves with sporangia in their axils sometimes modified and aggregated into cylindrical
strobili at ends of branches; spores of one size.................................... I. Lycopodiaceae
Leaves with sporangia in their axils aggregated into bilateral or tetragonous strobili at
ends of branches; spores of two sizes...................................................2. Selaginellaeeae

5.

Plants free-floating in water, some fronds or lamina lobes submerged and root-like; sori
submerged and spores o f two sizes....................................................... 15. Salviniaceae
Plants not free-floating with some fronds or lamina lobes submerged and root-like; if freefloating, sori on fronds above water and spores of one size............................................ 6

6.

Plants growing in water, at least seasonally, free-floating or rooted in mud. excluding
temporarily submerged rheophytes......................................................................................7
Plants not seasonally growing in water...............................................................................8

7.

Leaflets four; spores of two sizes, borne in sporocarps at stipe base...................................

....................................................................................................................14. Marsileaceae
Leaflets more than four; spores of one size, borne on under surface of leaflets.................
.......................................................................................... 22. Parkeriaccac (Ceratopteris)

15


FERNS & LYCOPHYTES VOL, I (2010)

8.

Sporangia embedded in or seated on a stalked spike attached at or near the base o f the
lamina...................................................................................................3. Ophioglossaeeae
Sporungiu borne on the under surface or the margin of the lamina, if on spike-like
outgrowths then these unstalked and at the apex or edges of lamina............................. 9

9.

Rhizome lleshy, stipes with stipules at their base, sporangia aggregated into sori or fused
to form synangia......................................................................................6. M arattiaceae
Not the above combination of characters...................................................................... 10

10.

Rhizome woody, stipes with a pair of stipules at their base, laminae simply pinnate,
veins free, sterile pinnae entire, sporangia completely covering under surface of fertile
pinnae which are at the base or in the middle of the lamina..................7. Osmundaceae
Not the above combination of characters......................................................................11

11.


Lamina thin, ± translucent, usually only one cell thick, sori at ends of veins, sporangia on
a± elongated receptacle sunk in a tube or protected by two flaps, marginal.......................
.........................................................................................................8. Hymenophyllaceae
Not the above combination of characters......................................................................12

12.

Fronds with pairs of branches, usually large, with frond apex dormant while each pair
o f branches develops, branches bipinnate or repeatedly forked; sori in small groups on
lower surface of lamina, w ithout indusium....................................... 9. Gleicheniaceae
Not the above combination o f characters......................................................................13

13.

Terrestrial or lilhophytic plants, rhizomes covered in hairs, veins copiously
anastomosing........................................................................................ 10. Diptcridaccac
Not the above combination of characters......................................................................14

14.

Terrestrial or lilhophytic plants, rhizomes covered in hairs, veins forked, sometimes
anastomosing, sori covered by umbrella-shaped indusia, fronds forked at apex, the
branches branching again many times in a series of unequal forkings very close
together.................................................................................................... 11. Matoniaceae
Not the above combination of characters..................................................................... 15

15.

Fronds (not rhizomes) climbing by twining, sporangia on narrow marginal lobes...........

............................................................................................ ...................... 12. Lygodiaccac
Not the above combination of characters...................................................................... 16

16.

Fronds erect, simple or dichotomously branched; sporangia in two or four rows on
narrow lobes at apex o f frond or of its branches................................. 13. Sclii/aeaceae
Not the above combination of characters...................................................................... 17

17.

Bases of stipes broad, stipes covered with mucilage when young and with aerophores;
scales absent; fronds simply pinnate, fertile fronds with sporangia covering their under
surface................................................................................................ 16. Plagiogyriaceae
Not the above combination of characters...................................................................... 18

16


KEY TO FAMILIES (PARRIS)

18.

Rhizome massive, upright, usually forming a conspicuous trunk, apex of trunk and
at least base of stipe densely covered with scales; fronds pinnate to bipinnate or
tripinnate in part, with sori on the under surface, sometimes with indusia.......................
..................................................................................................18. Cyatheaceac (in part)
Not the above combination of characters......................................................................19

19.


Rhizome stout, with long shining brown hairs; fronds tripinnatifid, with sori on the
margin, each protected by two convex indusia.................................... 17. Cibotiaceae
Not the above combination of characters....................................................................20

20.

Epiphytes with broad sterile nest fronds and pendulous dichotomously branched fertile
fronds........................................................................ 39. Polypodiaceae (P/atyceriunt)
Not the above combination of characters....................................................................2 1

21.

Fronds simple and entire or almost entire...................................................................22
Fronds lobed, pinnate or more deeply divided........................................................... 43

22,

Fertile fronds more or less completely covered with sporangia on lower surface or a
narrow apical part covered with sporangia..................................................................23
Fertile fronds, or a narrow apical pan. not more or less completely covered with
sporangia...................
26

23.

Sporangia confined to narrow apical part of frond...... 39. Pnlvpndiaccae (Belvisia)
Sporangia usually over whole under surface o f frond................................................. 24

24.


Veins free or joining only at frond m argin... 32. Dryopteridaccae (Llap/wglossum)
Veins anastomosing.........................................................................................................25

25.

Fronds not jointed to rhizome, free veins in arcoles pointing towards edge of frond.....
........................................................................................ 32. Dryopteridaceae (Bolbitis)
Fronds jointed to rhizome, free veins in arcoles pointing in all directions.....................
...................................... ......................... .............................. 39. Polypodiaceae (in part)

26.

Fronds not more than 1.5 mm wide, sori 3-6 mm long..... 23. Adiantaceae (in part)
Fronds wider or sori longer. ............................................................................
27

27.

Sori sunken in grooves on or near the frond margin..................................................28
Sori on frond surface in a line parallel to margin and often close to it. or along veins,
or in lines oblique to midrib of frond, or more or less round.................................... 29

28.

Rhizome scales dark brown to blackish, clathrate (latticed), conspicuous paraphyses
present in sori..................................................23. Adiantaceae (Haplopteris, in part)
Rhizome scales pale to medium red-brown, not clathrate (latticed), without conspicuous
paraphyses in sori............................................40. Gram mitidaccae (Scleroglossunt)


29.

Sort on frond surface in a line parallel to margin and often close to it.....................30
Sori along veins, or in lines oblique to midrib of frond, or more or less round.......32

17