THE FAMILIES
AND GENERA
OF VASCULAR PLANTS
Edited by K. Kubitzki
The Families
and Genera
of Vascular Plants
Edited by K. Kubitzki
I
Pteridophytes
and Gymnosperms
Volume Editors:
K. U. Kramer and P. S. Green
Assisted by E. G6tz (Illustrations)
With 216 Figures
Springer-Verlag
Berlin Heidelberg New York
London Paris Tokyo
Hong Kong Barcelona
Professor Dr. KLAUS KUBITZKI
Institut fUr Allgemeine Botanik und Botanischer Garten
hnhorststral3e 18
2000 Hamburg 52
FRG
o
Professor Dr. K. U. KRAMER
Universitat Zurich
Zollikerstral3e 107
8008 Zurich, Switzerland
P. S. GREEN
Royal Botanic Gardens Kew
Richmond, Surrey TW93AB
United Kingdom
Dr. ERICH GOTZ (Illustrations)
Institut fliT Botanik
Universitat Hohenheim
Garbenstral3e 30
7000 Stuttgart 70
FRG
ISBN 3-540-51794-4 Springer-Verlag Berlin Heidelberg New York
ISBN 0-387-51794-4 Springer-Verlag New York Berlin Heidelberg
Llbr.1rY of Congress Cataloging in Publication Data. Pteridophytes and gymnospenns / K U. Kramer and P. S. Green (eds.) ; with
contributions by J. Camus ... let at]. p. em (The Families and genera of vascular plants; vol. I) Includes bibliographical references.
ISBN 0-387-51794-4 (U.S.) 1. Pteridophyta-Classification. 2. Gymnosperms-Classification. I. Kramer, Karl Ulrich (1928- .
II. Green, P,S. III. Camus, J. (Josephine) IV. Series.
QK523.P78
1990 585.012-dc20
89-26217
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Preface
The study of organismic diversity has witnessed an unprecedented upswing in the last two decades. The empirical basis of botanical systematics has been broadened far beyond the realm of
macromorphological traits on which systematists have based their decisions for more than
200 years. Electron microscopy and phytochemistry have become standard tools, rendering evidence from micromorphology and micromolecules accessible to systematic research. The analysis of truly biological phenomena, like the interaction between various groups of organisms, or
the strategy of reproduction, has been added to the already existing fields of floral and dispersal
biology. The study of macromolecules permits genealogical reconstructions with the greatest precision. Considerable progress has also been ·made in the methodology for reconstructing phylogeny. All this is in consonance with the strong intellectual stimulus that the study of phylogeny
seems to exert on taxonomists. It is small wonder then that a large proportion of the taxonomists' activities is absorbed by the attempt to reconstruct phylogeny at all hierarchical levels.
While botanists from Linnaeus to Bentham & Hooker and Engler & Prantl have synthesized
the botanical knowledge of their time, such a synthesis has not been attempted more recently in
spite of, or possibly rather because of, the rising flood and rapid diversification of botanical
knowledge. Yet such a synthesis, by many considered as the foremost goal of systematic.biology,
would serve the needs of all those who are working in the fields of pure and applied biology, and
are interested in and depend on taxonomic information.
Because of this situation, the idea to produce a modem, comprehensive treatise of vascular
plant taxonomy at the generic level was proposed during a symposium held at the University of
Hamburg in 1976. Several colleagues, among them V. H. Heywood, F. Stafleu, and the late
R. Dahlgren, took part in shaping the outline of the work. At later stages, D. M. Bates,
K. U. Kramer, P. S. Green, P. Goldblatt, and S. Shetler offered invaluable assistence in various
matters. Rolf Dahlgren (1932-1987), Professor of Botany at the University in Copenhagen, had
an especially important share in developing the ideas leading to the present work. He also devoted much of his time during his last years to the preparation of the volume that is to deal with the
monocotyledons. Without his enthusiasm and ardour, work on this book would never have started. With deep gratitude the present volume is dedicated to his memory.
It has l)een our intention to bring together in this work information from all disciplines important to modem taxonomy. At the same time, a complete inventory of genera is attempted, including
means for their identification. Practical usefulness is considered to be of prime importance; therefore the keys are designed as tools for identification rather than for expressing phylogenetic
relationships. For the same reason useful generic revisions, where they exist, are cited. Only a limited
number of genera of all vascular plants are treated in the present volume, and therefore rather complete generic descriptions were permissible. It must be noted, however, that in the following volumes the descriptions can only be diagnostic, due to the sheer number of angiosperm genera.
In striking departure from accepted custom it has been decided to arrange the families alphabetically within each major plant group. The rationale for doing so is our awareness of the weak
basis of phylogenetic reconstruction in vascular plants (and especially angiosperms). In spite of
the great progress that is being made in this field, the systematic position of many families is still
unknown or uncertain. The delimitation of major subgroupings of the angiosperms is still unsatisfactory, and even their basic separation in dicotyledons and monocotyledons has been challenged! In establishing phylogenetic schemes the taxonomist very often is faced with problematic choices and has to opt for one of several equally tenable alternatives, or even for an
ill-founded solution. The sequence of families adopted in most systems of classification too often veils our ignorance, and it seems scientifically more honest to us to point explicitly to competing hypotheses or to gaps in our knowledge. Thus we try to avoid an undesired authoritative
VI
Preface
influence that might adversely influence the further development of plant taxonomy. An alphabetic arrangement of families, however, does not mean that consideration of relationships of
families and orders is ignored. On the contrary, the different views on the classification and phylogeny of the major groups are evaluated in special sections, and each family treatment includes a
paragraph discussing its affinities with other families.
Unfortunately, it is an inherent weakness of multiauthor works that a single missing contribution can delay the completion of the whole. Thus, it took much longer than originally anticipated
to complete this volume. I therefore have to apologize for the delay in publication of those contributions that had been submitted by the original target date. For many groups no suitable collaborator could be found. In these cases the treatments were prepared by the editors, with apologies for what may seem presumption; however, necessity for completeness dictated such a
procedure.
Generous support for the preparation of this and the subsequent volume was received from
the Volkswagenstiftung through the Joachim Jungius-Gesellschaft der Wissenschaften in Hamburg, for which I am particularly gratefuL Special thanks go to the volume editors for their meticulous work and to the authors, who generously provided their expert knowledge of the taxa
dealt with. I am also deeply indebted to Dr. E. Glitz for taking care of the illustrations and especially for providing numerous fresh illustrations of conifers, and to all those who very generously
assisted with the photographic illustrations.
Finally, I wish to extend my indebtedness to the Springer-Verlag for the very careful editing
and fine production of this volume and to Dr. K. F. Springer for the personal interest he took in
this work.
Hamburg, Summer 1990
K.KuBITZKI
Contents
Introduction to Pteridophytes and Gymnosperms
A Chemosystematic Overview of Pteridophytes
and Gymnosperms. O. R. GOTTLIEB,
M. A. C. KAPLAN, D. H. T. ZaCHER, and
K. KUBITZKI . . . . . . . . . . . . . . . .
2
Introduction to the Treatment of Pteridophytes
K. U. KRAMER and R. M. TRYON . . . . . . .
12
Conservation of Pteridophytes. A. C. JERMY .
14
General References for Pteridophytes . . . .
14
Glossary for Pteridophytes. K. U. KRAMER .
16
Keys to the Families of Fern Allies and Ferns
K. U. KRAMER . . . . . . . . . . . . . . . . . .
17
Psilotatae
Psilotaceae. K. U. KRAMER . . . . . . . . . . . ..
22
Lycopodiatae
Isoetaceae. A. C. J ERMY . . . .
26
PTERIDOPHYTES
Lycopodiaceae. B. 0LLGAARD
31
Selaginellaceae. A. C. JERMY
39
Equisetatae
Equisetaceae. R. L. HAUKE .
46
Filicatae
Notes on the Higher Level Classification of the
Recent Ferns. K. U. KRAMER . . . . . . . .
49
Aspleniaceae. K. U. KRAMER and R. VIANE
52
Azollaceae. J. J. SCHNELLER. . . . . . . . .
57
Blechnaceae. K. U. KRAMER, T. C. CHAMBERS, and
E. HENNIPMAN . . . . . . . . . . .
60
Cheiropleuriaceae. K. U. KRAMER
68
Cyatheaceae. K. U. KRAMER
69
Davalliaceae. K. U. KRAMER
74
Dennstaedtiaceae. K. U. KRAMER
81
Dicksoniaceae. K. U. KRAMER
94
Dipteridaceae. K. U. KRAMER
99
Dryopteridaceae. K. U. KRAMER, R. E. HOLITUM,
R. C. MORAN, and A. R. SMITH . . . . . . . . . . . 101
VIII
Contents
Gleicheniaceae. K. U. KRAMER.
145
Grammitidaceae. B. S. PARRIS .
153
Hymenophyllaceae. K.IwATSUKI
157
Hymenophyllopsidaceae. K. U. KRAMER and
D. B. LELLINGER . . . . . . . . . . .
163
Lomariopsidaceae. K. U. KRAMER
164
Lophosoriaceae. K. U. KRAMER
170
Loxomataceae. K. U. KRAMER
172
Marattiaceae. J. M. CAMUS .
174
Marsileaceae. K. U. KRAMER
.180
Matoniaceae. K. U. KRAMER
183
Metaxyaceae. K. U. KRAMER
186
Monachosoraceae. K. U. KRAMER
187
Nephrolepidaceae. K. U. KRAMER
188
Oleandraceae. K. U. KRAMER. . .
190
Ophioglossaceae. W. H. WAGNER, JR.
193
Osmundaceae. K. U. KRAMER
.
197
Plagiogyriaceae. K. U. KRAMER.
201
Polypodiaceae. E. HENNIPMAN, P. VELDHOEN,
K. U. KRAMER, and M. G. PRICE . . . . . ..
.. 203
Pteridaceae. R. M. TRYON, A. F. TRYON, and
K. U. KRAMER . . . . . . . . .
230
Salviniaceae. J. J. SCHNELLER .
256
Schizaeaceae. K. U. KRAMER .
258
Thelypteridaceae. A R. SMITH
263
Vittariaceae. K. U. KRAMER. .
272
GYMNOSPERMS
General References for Gymnosperms
Coniferophytina
Taxonomic Concepts in Conifers and Ginkgoids
C. N. PAGE . . . . . . . . . . . . . . . . . . . ..
(Conifers and Ginkgoids)
. . . . . . 280
282
Key to Families ofConiferophytina. C. N. PAGE. 283
Ginkgoatae
Ginkgoaceae. C. N. PAGE . . . . . . . . . .
284
Pinatae
General Traits of Conifers. C. N. PAGE
290
..
Interrelationships Between Families of Conifers
C. N. PAGE . . . . . . . . . . . . . . . . . . . . . . 292
Economic Importance and Conifer Conservation
C. N. PAGE . . . . . . . . . . . . . . . . . . . . . . 293
IX
Contents
Araucariaceae. C. N. PAGE
Cycadophytioa/Cycadatae
with the Single Living Order
Cycadales (Cycads)
294
Cephalotaxaceae. C. N. PAGE.
299
Cupressaceae. C. N. PAGE ..
302
Phylloc\adaceae. C. N. PAGE
317
Pinaceae. C. N. PAGE ....
319
Podocarpaceae. C. N. PAGE.
332
Sciadopityaceae. C. N. PAGE
346
Taxaceae. C. N. PAGE ..
348
Taxodiaceae. C. N. PAGE
353
General Traits of the Cycadales
L. A S. JOHNSON and K. L. WILSON
363
Conservation of Cycads. S. OLDFIELD . . . . . . . 368
. Boweniaceae. L. A S. JOHNSON and K. L. WILSON 369
Cycadaceae. L. A. S.JOHNSON and K. L. WILSON . 370
Stangeriaceae. L. A S. JOHNSON and K. L. WILSON 370
Gnetatae
with the Single Living Order Gnetales
Zamiaceae. L. A S. JOHNSON and K. L. WILSON.
371
General Traits of the Gnetales. K. KUBITZKI
378
Ephedraceae. K. KUBITZKI
379
Gnetaceae. K. KUBITZKI
.
Welwitschiaceae. K. KUBITZKI
383
387
Sources of I1Iustrations . .
393
Index of Scientific Names.
397
List of Contributors
CAMUS, JOSEPHINE M.
Department of Botany, British Museum (Natural
History), Cromwell Road, London, SW7 5BD, United
Kingdom
CHAMBERS, C.
National Herbarium of New South Wales, Royal
Botanic Gardens, Mrs. Macquarie's Road, Sydney,
N. S. W. 2000, Australia
GOTTLIEB, OTTO
R.:
R. L.
HAUKE,
Universidade de Sao Paulo, Instituto de Quimica,
C. P. 20.780, 01498 Sao Paulo SP, Brazil
Department of Botany, University of Rhode Island,
Kingston, Rhode Island 02881, USA
HENNIPMAN,
E.
Vakgroep Systematische Plantkunde, Rijksuniversiteit
Utrecht, Postbus 80.102,3508 TC Utrecht, The
Netherlands
HOLTTUM, R.
E.
Royal Botanic Gardens Kew, Richmond, Surrey
TW93AB, United Kingdom
IWATSUKI, K.
JERMY,
A.c.
JOHNSON,
L. A. S.
Botanic Gardens, Faculty of Science, University of
Tokyo, 3-7-1 Hakusan, Bunkyo-ku, Tokyo 112, Japan
Department of Botany, British Museum (Natural
History), Cromwell Road, London SW7 5BD, United
Kingdom
National Herbarium of New South Wales, Royal
Botanic Gardens, Mrs. Macquarie's Road, Sydney,
N. S. W. 2000, Australia
KAPLAN, MARIA AUXILIADORA C.
Universidade Federal de Rio de Janeiro, Nuc!eo de
Pesquisas de Produtos Naturais, Bloco H, Ilha da
Cidade Universitarja, 20.000 Rio de Janeiro RJ, Brazil
KATO, M.
Botanic Gardens, Faculty of Science, University of
Tokyo, 3-7-1 Hakusan, Bunkyo-ku, Tokyo 112, Japan
KRAMER, KARL ULRICH
Institut fiir Systematische Botanik der Universitat,
Zollikerstral3e 107, 8008 Ziirich, Switzerland
KUBITZKI, KLAUS
Institut fiir Allgemeine Botanik und Herbarium,
Ohnhorststral3e 18, 2000 Hamburg 52, FRG
LELLINGER,
MORAN,
D. B.
R.c.
OLDFIELD, SARA
0LLGAARD,
B.
U. S. National Herbarium, Smithsonian Institution,
Washington, DC 20560, USA
Missouri Botanical Garden, St. Louis, Missouri
63166-0299, USA
22 Mandene Gardens, Great Gransden, Sandy, Bedsh.
SG193AP, United Kingdom
Botanical Institute, University of Aarhus,
Nordlandsvej 68, 8240 Risskov, Denmark
XII
List of Contributors
PAGE, CHRISTOPHER N.
Royal Botanic Garden, Edinburgh EH3 5LR, United
Kingdom
PARRIS, BARBARA S.
Royal Botanic Gardens, Kew, Richmond, Surrey
TW93AB, United Kingdom
PRICE, MICHAEL
G.
University of Michigan Herbarium, North University
Building, Ann Arbor, Michigan 48109, USA
SCHNELLER, J. J.
Institut fUr Systematische Botanik der Universitat,
ZollikerstraBe 107, 8008 ZUrich, Switzerland
SMITH, A. R.
University Herbarium, Department of Botany,
University of California, Berkeley, California 94720,
USA
TRYON, ALICE F.
Harvard University Herbaria, 22 Divinity Ave.,
Cambridge, Massachusetts 02138, USA
TRYON, ROLLA M.
Harvard University Herbaria, 22 Divinity Ave.,
Cambridge, Massachusetts 02138, USA
VELDHOEN, P.
Vakgroep Systematische Plantkunde, Rijksuniversiteit
Utrecht, Postbus 80.102,3508 TC Utrecht, The
Netherlands
VIANE, R.
Instituut voor Plantenmorfologie, -oecologie en
-systematiek, Rijksuniversiteit Gent,
K. L. Ledeganckstr. 35, 9000 Gent, Belgium
WAGNER, W.
H., JR.
University of Michigan Herbarium, North University
Building, Ann Arbor, Michigan 48109, USA
WILSON, KAREN L.
ZoCHER, DOROTHE
National Herbarium of New South Wales, Royal
Botanic Gardens, Mrs. Macquarie's Road, Sydney,
N. S. W. 2000, Australia
H. T.
Universidade Federal de Rio de Janeiro, Nuc1eo de
Pesquisas de Produtos Naturais, Bloco H, Ilha da
Cidade Universitaria, 20.000 Rio de Janeiro RJ, Brazil
List of Families Treated in this Volume
Araucariaceae 294
Aspleniaceae 52
Azollaceae 57
Blechnaceae 60
Boweniaceae 369
Cephalotaxaceae 299
Cheiropleuriacaceae 68
Cupressaceae 302
Cyatheaceae 69
Cycadaceae 370
Davalliaceae 74
Dennstaedtiaceae 81
Dicksoniaceae 94
Dipteridaceae 99
Dryopteridaceae 101
Ephedraceae 379
Equisetaceae 46
Ginkgoaceae 284
Gleicheniaceae 145
Gnetaceae 383
Grammitidaceae 153
Hymenophyllaceae 157
Hymenophyllopsidaceae 163
Isoetaceae 26
Lomariopsidaceae 164
Lycopodiaceae 31
Marattiaceae 174
Marsileaceae 180
Matoniaceae 183
Metaxyaceae 186
Monachosoraceae 187
N ephrolepidaceae 188
Oleandraceae 190
Ophioglossaceae 193
Osmundaceae 197
Phyllocladaceae 317
Plagiogyriaceae 201
Pinaceae 319
Podocarpaceae 332
Polypodiaceae 203
Psilotaceae 22
Pteridaceae 230
Salviniaceae 256
Schizaeaceae 258
Sciadopityaceae 346
Selaginellaceae 39
Stangeriaceae 370
Taxaceae 348
Taxodiaceae 353
Thelypteridaceae 263
Vittariaceae 272
Introduction to Pteridophytes
and Gymnosperms
There is a broad consensus about the delimitation of
vascular plants as against thallophytes, and their major
subdivisions are well-established. Living pteridophytes
are conveniently subdivided into (true) ferns and fern
allies, the latter formed by lycopods, horsetails and
psilotates. Likewise, among gymnosperms the separation of a microphyllous coniferophytalean line (encompassing conifers and ginkgoids) from a megaphyllous cycadophytalean line has hardly been challenged
since the time of Coulter and Chamberlain (1910),
with the only exception that the Gnetales were originally not included in the cycadophytines, while at present they are usually considered as part of them. However, if we look back in time it appears that the fossil
record confronts us with forms that blur this clear-cut
picture, resulting in controversial interpretations of the
interrelationships and the phylogeny of major groups
of both pteridophytes and gymnosperms.
Although this book deals principally with living
tracheophytes and takes fossils into account only
where they contribute towards our understanding of
extant plants, some reference to fossils is germane.
As to the classes of the pteridophytes as defined by
neobotanists, various intermediate forms are known
from the Devonian and Lower Carboniferous. Thus
the characters of lycopods and horsetails are shared by
forms such as Eleutherophyllum. those of horsetails
and ferns by Ibyka. and of ferns and lycopods by
Enigmophyton. although the evaluation of these and
similar fossils is a matter of continuing palaeobotanical inquiry. The Psilotaceae, often considered to belong to a primitive class of fern allies, have more recently been suggested as being closer to true ferns,
although chemical evidence (see next section) does not
favour this. Morphological evidence casts doubt on
the status of Ophioglossaceae and Marattiaceae as
true ferns.
As to gymnosperms, comparable situations exist.
Firstly, the concept of a profound cleft between, and
consequently a very distant phylogenetic origin of, cycadophytinous and coniferophytinous forms has been
disturbed by the discovery of fossils combining charucters of both lineages, such as Phylladoderma. Eoginkgoites. and others. Secondly, the discovery of progymnosperms has brought to light a probable ancestor
lilf both these major gymnospermous groups. This is
most remarkable in view of the numerous characters
common to coniferophytes and cycadophytes, such as
the occurrence of lignified guard cells, the presence of
transfusion tissues in the leaves, the presence of biflavonoids (see next chapter), the near absence of polyploidy in favour of the presence of centromeric fission
or fusion of chromosomes, and so on. Such sharing of
characters would be rather enigmatic -if both groups
really had a distant origin.
It is still a matter of debate among palaeobotanists
whether or not primarily "platyspermic" seeds are of
utmost importance for tracing gymnosperm phylogeny, whether platyspermic and "radiospermic" seeds
originated independently from each other, and whether such characters are relevant to the position of forms
such as Ephedra. Suffice it to say that the clear-cut subdivisions in pteridophytes and gymnosperms as they
exist today are certainly the result pf the great age of
these groups. which has led to the deepening of the gap
between them, notwithstanding the - geologically
speaking - more recent and active diversification of
some of their components, such as the "younger" fern
families.
Thus the following major classification of pteridophytes and gymnosperms, forming the basis of the
present work, is largely a compromise between convenience and present knowledge. The specific problems
inherent in this classification are discussed in the introductory sections for pteridophytes and gymnosperms and the treatments of individual families.
Pteridophyta
Psilotatae with one living family
Lycopodiatae with three living families, attributed
each to one order
Equisetatae with one monogeneric family
Filicatae 33 families recognized in this book,
some of which are difficult to delimit; definition of orders controversial
Gymnospennae
Coniferophytina
Giukgoatae with a single relic species
Pinatae, or conifers,
encompassing nine families attributable to two
orders, Taxales and Pinales
Cycadophytina
Cycadatae with one living order encompassing
four families
Gnetatae with three isolated genera, each attributable to an order and family of its own
A Chemosystematic Overview of
Pteridophytes and Gymnosperms
O. R. GOTTLIEB, M. A. C. KAPLAN, D. H. T. ZocHER,
and K. KUBITZKI
Introduction
On the f01l0wing pages we have organized, in a systematic context, the mass of micromolecular information available for pteridophytes and gymnosperms,
with the aim of investigating, from the chemosystematic point of view, the relationships among them. A complete literature survey for chemical constituents of
these plants has become a task of gigantic magnitude.
This became apparent from our effort in surveying a1l
relevant entries in "Chemical Abstracts" relating to
pteridophytes. Because of limitations in time, we were
forced to extract the data for gymnosperms mostly
from the standard reviews listed under "General References" (p.14/15) and to rely heavily on our own,
necessarily limited judgement, to produce as balanced
a list of compounds as possible. Each of the structures
was then classified according to its biosynthetic origin,
as flavonoid, terpenoid and polyketide. Next, subclasses were recognized, each represented by a co 1Iection of compound types. Where appropriate, for relatively small c01lections, actual compounds, not compound types, were collated. The definition of compound types was based on substituents (e. g. 6,8-dimethylated flavonoids: Table 1, addendum 4) or on
skeletons (e. g. triterpenoids: Table 2, addendum 1).
The next step was the codification of compound types
or compounds. This is a very relevant operation since
codes express the degree of biosynthetic specialization
and at the same time a1l0w for the recognition of
chemical similarities among different taxa. For instance, a1l triterpenoids codified as H belong to the
hopanoid group, i. e. have either the hopane skeleton
itself (H1) or are derived from this by rearrangement
(H2-5), rearrangement and loss (H6) Or loss (H7 -9) of
Table 1 a. Selected flavonoid data of Pteridophyta
Flavones
Psil A~cg
Sela
Lyco ALL Tcg
Isoe ALL T 6,Sor
Equi AA Leg
Biflavonoids
Thel
Blec
Pter
CAALL
6,Sorcg -
Dick
.Q!:YQ A L cg
Loma
Neph
Olea
Azol
Salv
F1avanones
F1avanonols 3-Deoxy·
ProAnthocyanidins
KQSor
N
ArT
AA S.3' 6.0.4'
. AA ~ AA S.3' 6.0.4' 6.3'
K
Mara Acg
Osmu
Ophi
Glei
Schi
Plag
Loxo
Hyrne Acg
Loph
Cyat A L cg
Dava
Mars A L cg
Aspl
Denn Acg
Mona
Poly A cg
F1avonols
AAS.3'
K
QQ
KQ
KQ
AL
K
KQ
KQ
Ln Le 6.6'
K
KQSor
KQ
KKQ
KQ
KQM
KQ
N
NE
N6,Scm
N6,Scm
GQK~QQM
Pfkf§!
PNEE
6,Scm
C~D
C
N
M
Sor6,Scm
CD
P-
AL
AL
AL
C
CD
CD
CD
D
CC
CeD
ce
C£D
C~D
f 6,Scm
CD
CD
PC~
CC
Analysis of the Distribution of the F1avonoids
Table1b. Selected flavonoid data of Gymnospermae
F1avones
Cyca Aeg
Stan Acg
Zami ALcg
AA
Bowe
Gink LT
Arau
Taxa
Ceph
Podo
Scia
Phyl
Taxo
A
ALTcg
F1avonols
CQAcg6,8cm
Ephe A Lcg 8cm
Welw Lj"cg
A~ L Lcg
Flavanones
F1avanonols 3-DeoxyProAnthocyanidins
At! An 8.3' 6.0.4'
AA AA AA 8.3' 8.8
AAAAAA8.3'
AA AA AA AL 8.3'
AA AA AA 8.3' 6.0.4'
6.88.8-6Y
AAAA8.3'
AA AA AA 8.3' 6cm
AA AA AA 8.3' 6.0.4' 8.8
AA8.YAL
ALT
Cupr A LT6,80r
Pina
Biflavonoids
present
AA AA AA An An 8.3'
6.0.4'3.3'
AA AA AA 8.3' 6.0.4'
8.86.3'Ak
ck cc
CD
CD
C
KQQM
CD
CD
Q
Q
KQMeg
CD
CD
CD
Ar~T!
N
PCQD
PCQD
KQQM
CD
KQM
GKQMM
6,8cm
KQ80r
PPNE80r
6cm
ArT
CD
PPArT
AiD 6,8 cm
CD
PCD
P
One!
C
Addenda to Table 1
1. Glossary of symbols (initials of trivial names of compounds belonging to the types indicated in the column headings) for
5,7-dihydroxyflavonoids (additional hydroxylation indicated in first column):
5,7-0H
4'
3',4'
3',4',5'
Flavones
F1avonols
Flavanones
Flavanonol.
3-Deoxy-
chrysin
apigenin
luteolin
galangin
kaempferol
quercetin
myricetin
pinocembrin
naringenin
pinobanksin
aromadendrin
taxifolin
ampelopsin
apigenidin
luteolinidin
tricetin
eriodictyol
tri-OH-pinoc
Len.....
Anthocyanidins
pelargonidin
cyanidin
delphinidin
pelargonidin
cyanidin
delphinidin
2. Underlined symbols refer to partially O-methylated derivatives.
3. These compounds may appear also in O-glycosylated form in most taxa; the presence of C-glycosyl derivatives is indicated
by the suffix -eg.
4. The numbers 6 and 8 refer to position On ring-A of the flavonoids; the suffixes -or and -em indicating respectively hydroxylation (or methoxylation) and C-methylation at one or both of these positions.
5. The coupling mode of dimers is indicated by the bridged positions of the two monomers.
6. By billavonoids capital letters designate flavone parts, small letters designate naringenin (n), eriodictyol (e), kaempferol (k)
and catechin (c).
7. The presence of chalcones is not mentioned explicitly, but indicated by the structure of the corresponding flavanones.
methyl (and isopropyl) groups. Tabulation of these
codes for each family simplified the comparison of
chemical composition. Families with similar compositions were placed as close as possible in the construction of Tables 1, 2, and 3.
In this chemosystematic overview we have followed
the taxonomic framework adopted in this book. From
our analysis it became apparent that several of the
larger families, such as the Pteridaceae, include chemically heterogeneous genera and thus would require
separate consideration. At the moment, however, the
discussion of possible taxonomic rearrangements must
remain outside the scope of the present overview.
Analysis of the Distribution of the Flavonoids (Table 1)
In the fern allies the Psilotaceae possess a very simple
flavonoid composition totally based on the flavone
apigenin either in monomeric or in dimeric form. The
latter mode is further developed by partial O-methylation in the Selaginellaceae. The Isoetaceae and Lycopodiaceae develop only the monomeric flavone theme.
All major oxygenation patterns appear, and partial 0methylation, as well as 6,S-oxygenation, are introduced. Flavone accumulation also occurs in the Equisetaceae. Here also flavonols, which dominate the
flavonoid chemistry of all the remaining fern groups,
appear for the first time.
A Chernosystematic Overview of Pteridophytes and Gymnosperms
4
Table2a. Selected terpenoid data of Pteridophyta
Triterpenoids
Psil
Sela
Lyco
Isoe
Equi
Mara
Osrnu
Ophi
Glei
Schi
P1ag
Loxo
Hyrne
Loph
Cyat
Oava
Mars
Aspl
Denn
Mona
Poly
Thel
Blec
Pter
Dick
Oryo
Lorna
Neph
Olea
Azol
Salv
Steroids
Ecdysons
Oiterpenoids
Sesquiterpenoids
Gi
Sl
Sl,2 C3,4
OnSe
Pl
Al
Sl,6 Cl Ch
Sl Cl
S2,5
Sl
Hl
Hl
H2
Hl
Hl,2,4 Lu On Se
Hl-4,7
Hl,201
Hl-4 Cl 01
Hl,2Cl
Hl-4,7,9 Cl-7 Lu On Se
01-401-3 Vl,2 Bl-4, Po Co
Hl
Hl,2 Cl
Hl-6,8 Cl 04 W2 Sq
Hl
Hl-3 Cl,2,4,5 03 Vl Po Sq
Hl-3
Hl,2Cl
HI-5 01 Wl
Sl
SI
Sl,2 Ch
SI
SI
Sl
Sl,4,5
Sl
Sl Cl
Sl,3 Cl,2 Ch
Pl
Pl,3,11
P3
Pl,3
Al
Gi
L2
Gi
Pl,3,7,11
Kl Pl
Pl-3,5,7 Al,2
P6
Kl At Pl Ll
PI-4
Pl-3
A2 OrRy
Pl-3,11,13 E3
Pl,7
P3,11-14
Pl,11,14,15
Pl,3,7,11 Ml,2 El
Pl
P2
Sl
Sl
Sl Cl Ch
Table 2 b. Selected terpenoid data of Gyrnnospermae
Triterpenoids
Steroids
Ecdysons
Cyca
Stan
Zarni
Bowe
Gink
Arau
Taxa
Ceph
Podo
Scia
Phyl
Taxo
Cupr
C2 HI
Sl
Pl,3,13
H2
Sl
Pl-l0 Ml,2 El,2
Pina
Se 05 Cl,2
Ephe
Welw
Gnet
SI
Sl
SI
Sl
SI
Diterpenoids
Sesquiterpenoids
Gk
Ll,2 P4 Kl-4 Al At Ct
L2 Al Ta Ve Ce
Ha
L2,3 P4,5 Kl-3 Al-3,6-8 Ca La
L2 K3 Ve
K3
L2 P3-5 Kl,3 Al Ve
Ll,2 P3-5 K3,5 AI-4,7
L2 P3,4 Al,5 Ps Ce
Bi Bs Eu
Ca
Bs Ca
A2 Be Co Eu Lo Mu Bs
Ce
Ca Co Mu Bs
AaCeGu Lo
Bs Eu Hi
Analysis of the Distribution of the Flavonoids
Addenda to Table 2
1. Glossary of symbols for triterpenoids
Bl
B2
B3
B4
Cl
C2
C3
C4
CS
C6
C7
Co
DI
baccharanes
norbaccharanes
lemmaphyllanes
shionanes
cyclolaudanes
cycloartanes
norcycloartanes
dimethylcycloartanes
cyclomargene
norcyclolaudanes
bisnorcycloartanes
colypanoxide
dammaranes
2. Glossary of symbols for steroids
Ch cholesterol
Cl campesterol
C2 dehydrocampesterol
C3 ergosterol
3. Glossary of symbols for ecdysons
PI polypodin A
= ecdysterone
= crustecdysone
P2 polypodin B
1'3 ponasterone A
(warabisterone)
1'4 ponasterone B
PS ponasterone C
D2 euphanes
D3 tirrucalanes
D4 citrostanes
D5 lanostanes
HI hopanes
H2 femanes
H3 neohopanes
H4 filicanes
H5 adiananes
H6
H7
H8
H9
norhopane
norfernane
isoadiantane
trisnorhopane
C4
SI
S2
S3
brassicasterol
sitosterol
stigmasterol
dehydrositosteroJ
P7
P8
P9
PIO
PH
P12
P13
1'6 ponasterone D
4. Glossary of symbols for diterpenoids
1\ 1 abietanes
1\2-AS rearr. abo
1\6 trisnorabietanes
1\7 secoabietanes
I\b rearr. secoab.
KI entkauranes
K2 kauranes
K3 phyllocladanes
K4 hibaanes
KS enthibaanes
L1
L2
L3
P1
P2
P3
pterosterone A
podecdysone A
podecdysone B
podecdysone C
ecdysone
2-deoxy-3-epiecdysone
taxisterone
= shidasterone
=osladin
Lu lupanes
01
02
03
04
oleananes
multifloranes
taraxarane
friedelanes
On onoceranes
Po polypodatetraenes
Se serratanes
Sq squalene
UI ursanes
U2 taraxastanes
WI wallichimane
W2 norwallichimane
S4 spinasterol
SS fucosterol
S6 isofucosterol
PI4
PIS
Ml
M2
El
E2
E3
cheilanthone A
cheilanthone B
maldsterone A
maldsterone B
makisterone D
makisterone C
lemmasterone
entlabdanes
labdanes
secolabdanes
Ce cembranes
Ct entclerodanes
Gi gibberellins
entpimaranes
La
Ps
Ta
Ve
Gk
Ha
lauranane
pseudolaritanes
taxanes
verticillanes
ginkgolides
haningtonolide
P2
P3
P4
PS
P6
P7
Ry
norpterosines
isopterosines
norpterosin-C
ptaquiloside
bisnorpterosins
hypacrone
ryomenin
rearr. entpim.
pimaranes
P4 isopimaranes
P5 rearr. isopim.
At entatisanes
Ca camphorane
5. Glossary of symbols for sesquiterpenoids
1\ I abscisic acid
Co copaanes
A2 ionanes
Dr drimane
Eu eudesrnanes
Aa alaskanes
Gu guaianes
lie bergamotanes
Hi himachalanes
IIi bilobalides
Lo longifolanes
lis bisabolanes
('a cadinanes
Mu murolanes
PI pterosines
('c cedranes
6
A Chemosystematic Overview of Pteridophytes and Gymnospenns
In ferns, the flavone character is weakly expressed
in the two ancient and isolated families Marattiaceae
(only apigenin C-g1ycosides) and Osmundaceae (apigenin dimers), both already containing flavonols and
the latter 3-deoxyanthocyanidins. The Ophioglossaceae are aberrant chemically, as demonstrated by their
exclusive terpenoidal flavonoids and dihydrostilbene
as well as partially O-methylated flavonoids, rare features in the pteridophytes. This is most interesting
since it has been argued that the Ophioglossaceae may
perhaps belong to a stock radically different from the
pteridophytes.
Among leptosporangiate ferns, some homogeneity
of flavonol composition is observed for the four families Gleicheniaceae, Schizaeaceae, Plagiogyriaceae and
Loxomataceae. In the Hymenophyllaceae O-glycosylation is added to the common C-glycosylation of apigenin and the Cyatheaceae contain exclusive flavoneflavanone dimers. With regard to flavonoid chemistry,
the Marsileaceae (flavanones, but yet neither 6,8- nor
3',4',5'-hydroxylation) and Davalliaceae (yet no flavanones, but 8-hydroxylation) must be considered transitional to the following group of families. These fern
families, though not closely related but obviously representing a more modem stratum, contain flavanones
and/or chalcones (see Table 1, addendum 7) and
ring-B trihydroxylated compounds represented by leucodelphinidin in the Aspleniaceae, Dennstaedtiaceae,
Thelypteridaceae, Pteridaceae and Dryopteridaceae,
and by myricetin in the latter two families as well as in
the Polypodiaceae and Blechnaceae. The flavonoid
chemistry is particularly similar in the Pteridaceae and
Dryopteridaceae, which both possess not only some
6,8-C-methylated compounds, a feature which they
share with the Thelypteridaceae and Blechnaceae, but
also some compounds exempt of ring-B hydroxylation, in which they agree with the Dennstaedtiaceae.
The Lomariopsidaceae with ring-A 6,8-C-methylated
compounds, as well as the Nephrolepidaceae and
OIeandraceae with ring-B trihydroxylated compounds,
probably also belong to this group, although for these
families basic flavonoid data are yet missing. Insufficient data also exist for the placement of the Salviniaceae itnd Azollaceae, since the presence of leucopelargonidin in the latter and in Lomariopsidaceae is only a
vague clue to affinity. No data exist in our file for
Dicksoniaceae, Lophosoriaceae, Monachosoraceae
and other families.
Apigenin and dimeric forms of this flavone dominate the flavonoid chemistry of the Cycadales (with
the possible exception of the Stangeriaceae), Ginkgo
and the conifers (with exception of the Pinaceae). In
the Gnetales dimers are represented by bicatechols
(Ephedra) and bistilbenes (Welwitschia and Gnetum).
Among the many possible coupling modes between
the two apigenin units, the typical 8.3'- and the
6.0.4' -modes of the pteridophytes are by far the most
common ones. Furthermore, again as in primitive pteridophytes, the common flavonols, kaempferol and
quercetin, are absent or rare in the Cycadales and perhaps even in some families of the conifers (Cephalotaxaceae and Araucariaceae), but appear commonly in
others (Taxaceae, Podocarpaceae, Cupressaceae, Tax0diaceae and Pinaceae). The aberrant and possibly advanced position of the Pinaceae is apparent from the
wide range of ring-B substitution of its flavonoids,
which varies from the absence of oxy-groups to the
presence of three oxy-groups, and from the frequency
of 6- and 8-C-methylation. Biflavonoids are represented by a singular flavone-flavonol dimer. Flavanones and chalcones are also characteristic of the Pinaceae. The presence of dihydroflavonols, and of the
more highly specialized isoflavones, are considered to
point to an affinity of Podocarpaceae with the Cupressaceae.
In summary, flavonoid evolution, both in the pteridophytes and gymnosperms, is considered to have involved the following cumulative introductions, referring to structural type, oxygenation pattern and substitution:
Structural type:
1. Flavones
2. Flavonols
3. Flavanones and chalcones
4.3-Deoxyanthocyanidins
5. Anthocyanidins
6. Leucoanthocyanidins
Oxygenation pattern:
1. Ring B hydroxylation at 4'--+3',4'--+3',4',5'
2. Ring A hydroxylation at 5,7--+5,8(6),7--+5,6,7,8
3. Ring B exempt of hydroxylation
Substitution:
1. C-Glycosylation
2. O-Glycosylation
3. C-Methylation at 6, at 8, or at 6,8
Analysis of Terpenoid Distribution (Table 2)
In the pteridophytes, the distribution of terpenoids not
only corroborates the systematic value of the flavonoid
data, but indicates affinities for some families for
which flavonoid data are inconclusive or missing.
The most comprehensive terpenoidal markers in the
pteridophytes, at least at the level of family, are the triterpenoids. With the exception of the anocerane derivatives in the Lycopodiaceae they have not yet been located in the lycopods, horsetails, Psilotaceae and
eusporangiate ferns. In contrast, nearly all remaining
fern families contain hopanoids representing one or
Analysis of Terpenoid Distribution
7
more skeletal types. Of most frequent occurrence are pounds. Only very few abietanes occur in significantly
the hopanes, femanes, neohopanes, filicanes and adia- primitive pteridophytes but more of them occur in the
nanes. Nevertheless, as it was demonstrated above for angiosperms. Among some seemingly more specialflavonoids, a distinction between ancient and modem ized classes of diterpenes are the verticillanes (Taxorem families can also be based on triterpenoids since diaceae, Taxaceae) and the cembranes (Taxaceae, Pionly families of the latter group are additionally char- naceae). The taxanes (Taxaceae) and the ginkgolides
acterized by cycloartanes, including several related (Ginkgoaceae) possess highly unusual carbon skeletons.
types such as norcycloartanes, cyclolaudanes, etc.
A very different situation is observed for mono- and
The steroids of the pteridophytes are rather common representatives and as widely distributed in this sesquiterpenes which abound in types and number in
group as in the rest of the plant kingdom. Even the ec- . most conifers. Only the Taxaceae (Taxus, but not Tordysons, although preferentially accumulated in mod- reya) and Ginkgoaceae are exceptional in this respect,
ern fern families, appear somewhat sporadically else· both characterized by highly oxidized diterpenoids, rewhere. Here it is not distribution, but increasing spectively taxanes and ginkgolides. The oxidative
diversity and oxygenation of the molecules, which par- power of their biosynthetic machinery seems hardly
allels evolution.
compatible with the presence of easily oxidizable, simMany diterpenoids have been isolated from the ple monoterpenes. Indeed, chiefly from Ginkgo small
pteridophytes. However, from the chemosystematic amounts of highly aromatized mono- and sesquiterpepoint of view, only kauranes and primaranes are use- noid derivatives have also been isolated.
I'ul so far. Both confirm the affinity between Denn·
staedtiaceae and Pteridaceae and between the latter
Analysis of the Distribution
and the Dryopteridaceae.
The sesquiterpenoidal pterosinoids, possessing very of Additional Compound Classes (Table 3)
unusual structures, have so far been found exclusively
in pteridophytes, and are thus excellent clues for rela· Among the few systematically relevant classes of fatty
tionships. A simple derivative occurs in the Equiseta- compounds are the (O-hydroxyacids, accumulated
ceae, confirming the flavonoidal evidence that chemi- preferentially by the more primitive families of the
cally this family approaches the level of organization pteridophytes such as the Psilotaceae, Selaginellaceae,
oi' modem fern families. Diversification of pterosi- Lycopodiaceae and Schizaeaceae. (0- Hydroxyacids
noids, very similar in Pteridaceae and Dicksoniaceae, are most abundant in many families of the gymnoattains its peak in the Dennstaedtiaceae, while a still sperms, in which their interesterification leads to the
more derived representative occurs in the Monacho- typical estolides.
Among the polyketides the largest class is represllraceae. If pterosinoids are really missing in the Davalliaceae and Marsileaceae, these families, which had sented by the abundantly C-methylated acylphlorogluappeared to be borderline cases upon analysis of their cinols of the Dryopteridaceae.
Lignoids, C6Cr monomers and dimers are very rare
Ilavonoids, resemble the ancient stratum of fern families. In contrast, the pterosinoids of the Monachosora- in the pteridophytes but abound in the gymnosperms,
ceae and Dicksoniaceae point to the modem stratum though in lesser structural diversity than is found in
and it is there that the cycloartanes of the Nephro- the angiosperms. The very special C6C5~-lignoids
Icpidaceae confirm the position of the last-mentioned (conioids) are known from the Araucariaceae, Cupressaceae and Thxodiaceae.
lillnily.
Stilbenoids, ~C2"monomers and oligomers are abPrecisely as in primitive pteridophytes, but in conI!"Ust to advanced ones, triterpenoids are very rare in sent from the pteridophytes. The monomers appear in
primitive conifers. Simple hopanes have been located the Pinaceae, while dimers and trimers occur additionin the Podocarpaceae and Taxaceae and only the Pina- ally in Gnetum and Welwitschia, where they seem to be
an important clue to affinity.
ceae contain slightly more diversified representatives.
Pteridophytes and gymnosperms are relatively poor
The steroids of the gymnosperms are totally unex"eptional and ecdysons, most of which have already in nitrogen-containing secondary metabolites. Neverhcen recognized in ferns, are accumulated in the Podo· theless such compounds are worthy of mention in the
present context, in view of their systematic distribu\·a ..paceae and Taxaceae.
Diterpenoids are also of widespread occurrence in tion. This is the case of the lysine-derived "LycopoIhe gymnosperms. However, the distribution of lab- dium alkaloids" (Lycopodiaceae), the inclusively ornidanes, primaranes and kauranes is so uniform that thine-derived palustrines (Equisetaceae), the methylIhey are of little help in respect of chemosystematic azoxymethanol g1ycosides and a-amino-f:l-methylami\'lIdcavour. This is also true for abietanes, the most nopropionic acid (Cycadales), the aminohydroxycindlllracteristic single class of gymnospermous com- namates of taxanes (Taxaceae), the tyrosine-derived
A Chemosystematic Overview of Pteridophytes and Gymnospenns
8
Table3a. Further selected chemical data for Pteridophyta
Unsaturated acids
Hydroxyacids
Psil
Sela
Lyco
Lsoe
Equi
Mara
Osmu
Ophi
Glei
Schi
Plag
Loxo
Hyrne
Loph
Cyat
Dava
Mars
Aspl
Denn
Major
Minor
16:160H
16:160H
16:(OH),
14:140H
Polyketides
Lignans
pyPa
16:118:1-3
16:10,I60H
16:9,160H
Alkaloids
and other
N·cmpds
py Ly
16:118:1-320:4,5
30:3OCO,H 28:280H26:260H
16:160H
Stilbenes
L2
Ll,3
16:118:1-320:4
Xl,2
16:0H
16:118:1-320:4
16:318:1-320:4
Xl
Xl
Xl,2
Mona
Poly
Thel
Blec
Pter
Dick
Dryo
16:318:1-320:4,5 Al
18:1,2
X2
16:1,318:1-3
20:4,524:4
Al,2 em Ll-3 Xl,2
F1 Be
py
Lorna
Neph
Olea
Azol
Salv
homoerythrines and cephalotaxins (Cephalotaxaceae),
the benzyltetrahydroisoquinolines (Onetaceae), taxi·
phyllin, ephedrines, ephedradines and ephedroxane
(Ephedraceae) and the possibly acetate· derived pinidine (Pinaceae),
Conclusion
The analysis of the distribution of chemical characters
shows remarkable agreement between primitive pteridophytes (fern allies plus ancient fern families) and
gymnosperms, This applies to the widespread occurrence of biflavonoids in the Psilotaceae and Selaginellaceae, on the one hand, and in the Cycadales and co·
nifers, on the other. In all these groups biflavonoids
are based on apigenin and show identical coupling
patterns. Also, the rareness of flavonols, flavanones
and flavanonols in the primitive families of the pterid·
ophytes and their at least partial absence from the
gymnosperms are remarkable. Indeed, both the pterid-
ophytes and the gymnosperms could be envisaged as
sharing the same (psilophytalean?) ancestor, character·
ized by the presence of fatty w-hydroxyacids and biflavones (two 8.3'- and 6.0.4'-coupled apigenin units),
as well as by the relative scarcity of flavonols and triterpenoids. The widespread occurrence of oxidative
coupling reactions, leading not only to biflavonoids
but also to dimerous catechols and dimerous and trimerous stilbenes, must be an ancient character in the
evolution of tracheophytes.
Further chemical evolution of the two major groups
was divergent. The pterosinoids and flavonoids of the
Equisetaceae are supposed to have evolved parallel to
those of true ferns. In advanced families of the pteridophytes triterpenoids are well diversified, as shown
chiefly by the presence of many compounds of the hopane and cycloartane types. Among their diterpenes
kauranes predominate. These always belong to the
ent·series and hence do not necessarily show any relationship to the kauranes of the gymnosperms, which
mostly belong to the normal configurational series. An
Conclusion
9
Table3b. Further selected chemical data for Gymnospermae
Hydroxyacids
Major
Cyca
Unsaturated acids Polyketides
Lignans
Stilbenes
and dimers
Minor
16:10,160H 16:9,160H
18:9,10,180H
18:2,320:2,3
AzAm
Stan
Zami
Az
Az
Az
Py
Bowe
Gink
16:10,160H 16:9,160H 16:160H 16: 1-3 18:1-4
20:2-4
Arau
Taxa
Ceph
Podo
Scia
Phyl
Taxo
Cupr
Pina
16:10,160H 16:9,160H 16:0H
Ephe
Welw
Gnet
Alkaloids
and other
n·cmpds
Ll-3 Fl,2 Cl
Ll-3
18:220:3
Di Ta
P2,3
16:9,160H
16:160H
18:9,10,180H
20:3
20:320:4
Ll-3
18:1,220:3,4
20:3,4
18:1-320:3,4
L1,3 Cl,2
P2
Ll-3 Fl,2 Cl,3
Ll-3, Fl,2 Be Pn.!'!!RPcf£ Pi
20:3,4
16:10,160H 16:160H 10:100H
16:10,160H
El-3
RRR
Ac
Cy
RRR~HR
Pl
Addenda to Table 3
1. Glossary of symbols for ",·hydroxyacids; number of carbon atoms of the normal fatty acid chain: position of hydroxyls.
2. Glossary of symbols for unsaturated acids; number of carbon atoms of the normal fatty acid chain: number of double bonds
in one or more derivatives; Cy cyclopropane fatty acids (sterculic acid, malvalic acid)
3. Glossary of symbols for polyketides
Ll osmundalactone
L2 angiospteroside
L3 other simple lactones
4. Glossary of symbols for lignans
Ll dibenzylbutanediols
L2 dibenzyl·y-Iactones/ols
L3 aryltetralins/naphthalenes
Al
A2
Ac
Fl
F2
Be
acylphloroglucinols
methylenebisphloroglucinols
acetophenones
Xl
X2
mangiferin/isomangiferin
other xanthones
arylbenzofurans
diarylfurofurans
arylbenzylfurans
C1
C2
C3
conioids
modified conioids
P3
Pa
Pi
cephalotaxines
palustrines
a-pipecolines
pyridines
taxiphyllin
diarylbutadiene
S. Glossary of symbols for 3,5-dihydroxystilbenes
Pn pinosylvin
Pc
piceatannol (3',4'-OH)
H
hydroxyresveratrol (2',4'-OH)
R resveratrol (4'-OH)
Underlined symbols refer to partially O-methylated derivatives
(,. Glossary of symbols for alkaloids and other N-containing compounds
Am a-amino-p-methylaminopropionic acid
E2 ephedradines
Az azoxymethanol g1ycosides
E3 oxazolidone/dines
Oi p-dimethylamino-a-hydroxycinnamates
Ly lycopodium alkaloids
of diterpenoid taxanes
P1
benzyltetrahydroisoquinolines
El ephedrines
P2 homoerythrines
IInalogous situation prevails for pimaranes. Another
class of compounds characteristic of advanced pteridophytes is constituted by the methylenebisphloroglucinols. With respect to flavonoids, in the advanced pteridophytes flavonols abound and chalcones and flavanones are common. Representatives of the latter com"ound types, absent from primitive pteridophytes, ap-
Py
Ta
pear in the gymnosperms only in the Pinaceae, notably an advanced family. Thus it can be said that the
primitive chemical traits shared by pteridophytes and
gymnosperms may point to their common descent.
The similarities acquired during further evolution appear to be due to the action of similar selective pressures exerted upon them.
10
A Chemosystematic Overview of Pteridophytes and Gymnospenns
Rather strong chemical similarities also exist between advanced pteridophytes and angiosperms. With
regard to flavonoids these include the presence of
flavonols, flavanones, ring-A 6,8-C-methylation and
ring-B trihydroxylation. For terpenoids, triterpenes
(hopanes, cycloartanes), diterpenes (chiefly kauranes
of both stereochemical series, pimaranes, labdanes)
and sesquiterpenes can be adduced. Additionally,
highly C-methylated phloroglucinols (in Dryopteridaceae, Rosaceae, Myristicaceae) can be mentioned.
Since no relationship between ferns and angiosperms
is acceptable, the appearance of these substances in
both plant groups is another example of parallelism.
The notable chemical agreement between the Gnetatae
and some angiosperms, commented upon earlier by us
(Gottlieb and Kubitzki 1984), is also pertinent here.
The widespread occurrence of parallelism may be explained by postulating the action of common selective
pressures, which are related to the plants' defence system.
References
(See also General References to Pteridophytes)
Berti, G., Bottari, F. 1968. Constituents of ferns, In: Reinhold, L., Liwschitz, Y. (Eds) Progress in Phytochemistry.
Vol. 1: 589-685. London: Interscience Publ.
Boralle, N., Braquet, P., Gottlieb, O. R. 1988. Ginkgo biloba: a
review of its chemical composition, In: Braque!, P. (Ed)
Ginkgolides - chemistry, biology, pharmacology and
chemical perspectives. Vol. 1, 9-25. Barcelona: J. R. Prous,
Science Publ.
Cooper-Driver, G. A., Haufler, C. 1983. The changing role of
chemistry in fern classification. Fern Gaz. 12: 283-294.
Gottlieb, O. R., Kubitzki, K. 1984. Chemosystematics of the
Gnetatae and the chemical evolution of seed plants. Planta
Medica 50: 380-385.
Hegnauer, R. 1962. Chemotaxonomie der Pflanzen. Vol. 1,
220-317. Basel: Birkhauser.
Hegnauer, R. 1985. Chemotaxonomie der Pflanzen. Vol. 7,
398-554. Basel: Birkhauser.
Soeder, R. W. 1985. Fern constituents: including occurrence,
chemotaxonomy and physiological activity. Bot. Rev. 51:
442-536.
PTERIDOPHYTES
Pteridophytes: Ferns and Ferns Allies
Plants with a regular alternation between a gametophytic (sexual) and sporophytic (asexual) phase. Sporophytes with leaves, axes and roots (the latter missing
in Psilotaceae), and with well-developed conducting
tissues. The leaves scale-like, with a single vascular
stral)d, or markedly compound and described as
"fronds". Stems protostelic, solenostelic, or dictyostelic, sometimes polystelic; some with limited secondary
thickening. Sporangia thick- or thin-walled, homosporous or heterosporous, borne terminally on an axis or
on leaves. Spores trilete (triradiate) or monolete. Gametophytes autotrophic or mycotrophic. Male gametes
(antherozoids) flagellated. Female gametes (egg-cells)
borne singly in flask-shaped archegonia.
Four classes
Psilotatae
Lycopodiatae
Equisetatae
Filicatae
Introduction to the Treatment
of Pteridophytes
K. U. KRAMER and R. M. TRYON
The following general remarks on the iaxonomy of the
pteridophytes at higher levels focus on the systematics
of the ferns in a narrower sense, especially on what
used to be called the leptosporangiates. The relationships between the classes of the traditional pteridophytes are still very difficult to assess, if indeed they
are at all related in the sense of being derived from a
common source of cormophytes. These relationships
can only be discussed within a palaeobotanical framework (see p. 1), rather than in a work like the present,
which concentrates on living plants.
Fern taxonomy predominantly based on the sorus,
as initiated by Linnaeus (1753) and continued through
the works of W. J. Hooker and his followers, especially
J.G.Baker (e.g., 1865-68, 1874), caused much confusion and retarded the development of fern classification (see, e. g., R. M. Tryon 1952; Holttum 1982). This
viewpoint overwhelmed the pioneering efforts of John
Smith, C. B. Presl, and Fee, in particular, who brought
many characters into the classification of genera. The
authority of the Hookerian school was finally overcome by works such as those of Christ (1897), Diels
(1899-1900), and Christensen (1905-06 and after).
However, the sorus and the structures associated with
it continued to be important in classifying the "higher
leptosporangiate ferns" into genera, tribes, and subfamilies. FInally, the accumulated data from morphological-anatomical studies of representatives of numerous genera, stimulated partly by research on fossils
and admirably integrated by Bower (1923-28), led to
the realization that the complex of characters centering on the sorus is just one group of many characters
that can, and should, be used in fern classification. especially on the levels of genus up to family.
In principle then, the time was ripe for a return to
the best ideas of J. Smith, Presl, and Fee that especially
emphasized the utilization of a multiplicity of characters. The breakthrough was to a great measure due to
R. C. Ching's work (especially 1940), which forms the
basis of a classification that at last emancipated itself
from the dominance of soral characters. Copeland's
"Genera Filicum" (1947) relied heavily on Ching's
work, although he was unjustly critical of it in respect
to rather numerous details on which he disagreed with
Ching.
Major classifications such as those of Christensen
(1938) and Copeland (1947) were unfortunately pub-
Iished before much was known about chromosome
numbers, gametophytes, the surface structure of
spores, or phytochemistry. Cytological techniques and
the resulting records of chromosome numbers in an
ever-increasing array of fern genera from many parts
of the world (Manton 1950 and after) led to a flood of
new data as well as a reassessment of systems of fern
classification. Some fern families like Copeland's Pteridaceae and Aspidiaceae were shown to be cytologically quite heterogeneous and artificiaL Other extensive
work that has added data important for the assessment
of generic relations and classifications include the
study of spores by Lugardon (1972, 1974, etc.) and by
A. Tryon (in Tryon and Tryon 1982), as wen as critical
work on chemical constituents summarized by Swain
and Cooper-Driver (1973, 1980). The enormous
amount of evidence gathered in these new fields has
influenced fundamental ideas on the affinities among
extant pteridophytes.
A major problem with some relatively recent classifications, as of Copeland's (1947), is that while reliance was placed on characters from the morphological-anatomical body of data, the Hookerian influence
stin persisted. Copeland retained many genera on the
basis of differences in sora! shape, presence or absence
of an indusium, sterile-fertile dimorphism, acrostichoid distribution of sporangia, etc. These genera are
now considered as artificial segregates by many workers and the characters as unreliable because the plasticity of such characters in most fern families has become fully realized. The tendency to use easily
observed characters as dimorphism, free vs. anastomosing veins, leaf dissection patterns, etc. for delimiting genera continues into the present, while criteria
based on a body of correlated characters, many of
which may not be so readily observed, are not funy
applied.
A period has begun in which some genera, and to a
lesser extent families that have long been accepted, are
now regarded as artificial, either because they can be
demonstrated to be heterogeneous (genera by definition), or because they were artificially segregated. The
tendency to cancel such genera is still not sufficiently
strong. The opinion is sometimes expressed, but we
believe superficially founded, that it is conservative
(i. e., outmoded) to combine genera. Profound examination of more extensive material often yields greater
emphasis on the points of difference; and points of
similarity tend to be either ignored or else depreciated.
This is particularly unfortunate at the generic level
where classification has such a strong influence on nomenclature.
Hybridization often reflects similarities, and it may
occur without respect for "generic" boundaries. In
such cases, e. g., in Aspleniaceae, it is evident that
characters used to separate genera do not rest on pro-
Introduction to the Treatment of Pteridophytes
found genomic differences and therefore should not
be given particular systematic weight.
One of the major tasks undertaken in the preparation of the present work was the critical evaluation of
every taxon to determine whether it was both sufficiently distinct and clearly an evolutionary group to
warrant recognition and generic separation. This evaluation was based on as many characters as possible as
well as on experience gained from a study of a broad
spectrum of the pteridophytes throughout the world.
Characters shown to be unstable and unusable for the
classification of ranks higher than species in one
group should be evaluated in all cases where they have
been used to distinguish genera. In retrospect, it is difficult to understand why, e. g., the presence or absence
of an indusium, shown to be a character of little relevance in Dryopteridaceae, Thelypteridaceae, and
Cyatheaceae, should still be employed for the segregation of satellite genera from Asplenium, when other
lines of evidence show them to be so intimately related.
The result of this viewpoint is evident in the genera
presented in this classification, in which many genera
recognized by various authors since 1947 have disappeared, and, as we believe, on good ground. This viewpoint may be considered as conservatism. However,
the term "conservatism" is more correctly applied in
reference to the retention of genera of previous classifications without critical assessment of both their distinguishing features and their similarities. It is a basic
tenet of taxomony that the burden of proof rests on
the person who claims to have found differences, and
not on one who merges closely related taxa with each
other. The statement that "they are different" without
supporting evidence and without consideration of similarities is not only objectionable but unscientific.
The opportunity to express degrees of similarity by
recognizing subgenera and sections was badly neglected by Copeland (see, e. g., Morton 1968) and othcrs. Infrageneric ranks should be employed whenever
appropriate and they are used in many cases in the following treatment.
The system of pteridophyte families and genera to
be found below reflects the effect of such consolidation. It should, of course, in no way be regarded as "final", for reassessment must be made as new data become available. In some cases the data indicated the
need for separation, as in the segregation of CalochlaelIa from Culcita. In this example we have been fortunate to receive permission to utilize data prior to their
publication. In other cases, the suspected artificiality
13
of a genus is indicated although we are unable to present a better classification. This applies to Dennstaedtia and perhaps to Blechnum. Doubts are frequently
presented, whether voiced by others or from our own
work, with hope that this will stimulate further studies
in the areas where it is most needed.
We consider that a heavy responsibility rests on
those who place a comprehensive classification before
the scientific world: it will be quoted, requoted (and
likely misquoted) time and again, possibly without a
sufficiently critical attitude, as has happened so often
with other classifications.
We have attempted to utilize many data as the basis
of the present classification. It must be emphasized
that certain genera are still very incompletely known,
especially as to their internal structure. Certain characters either have not, or have only incompletely, been
observed and described for many taxa although we regard them as important for the characterization of certain genera. Among these, which have been provided
as far as possible, are the details of the epidermal appendages of the lamina and its catadromic or anadromic architecture. In other cases both literature and
available materials are inadequate, and accordingly
the taxonomic evaluation of characters must be regarded as provisional. These comments apply to such
taxa as, e. g., Oenotrichia, Psammiosorus, and Thysanosoria. Accordingly, the present treatment does not always represent as critical an assessment as we would
like.
The greatest body of original information is contained in the spore descriptions based on observations
of scanning electron micrographs. Our indebtedness to
A. F. Tryon for contributing this very extensive new information is accordingly great. The treatment of the
Polypodiacea,e also contains newly acquired information on the scales, "paraphyses", venation, etc., provided by E. Hennipman and collaborators at Utrecht.
Certain taxonomists may be critical because we
have consciously refrained from employing cladistic
methods in developing our system of classification.
We regard cladistics as merely one of several possible
methods for arriving at an acceptable classification. In
the pteridophytes we regard evidence for what has
been aptly called "reticulate evolution" and at the generic level might be cautiously termed "reticulate affinities" to be so strong that cladistics is by no means
the best method to use in unravelling the relationships.
For references, see under "General References to Pteridophytes" (p.14/15)
Conservation of Pteridophytes
General References for Pteridophytes
A.C.JERMY
Pteridophytes have evolved to fill almost every ecological niche but the greatest species diversity is clearly
found in the tropical rainforest. The rapid disappearance of this biome throughout the world with many of
their pteridophyte species yet undiscovered, let alone
described, is of great concern. It is fortunate that the
richer montane forests are the last to disappear. Sustainable logging unfortunately takes the mature trees
with large canopies frequently covered with rich epiphytic pteridophytes. It is important to survey as soon
as possible those tropical areas under threat, and to
assess how many and which general families are actually protected in national parks and nature reserves.
In the temperate areas, especially in the developed
northern hemisphere, agricultural, industrial and urban development brings about land-use changes which
destroy fern habitats, especially wetlands and forested
areas. On the other hand, new habitats are created and
pteridophyte diversity is for the most part retained albeit in a mosaic of man-made habitats.
Activity to conserve pteridophytes throughout the
world is being co-ordinated by David Given and Clive
Jenny of the International Association of Pteridologists in conjunction with the Species Survival Commission of I.V.C.N., and a Red Data Book for Pteridophyta is being prepared. Data are being held by the
World Conservation Monitoring Centre at Cambridge,
England, which already has infonnation on 1650
threatened species of pteridophytes, of which 20 may
already be extinct, 67 endangered, 91 vulnerable, 354
rare, and 1318 'candidate species', i. e. likely to be in
one of these categories when more information becomes available.
Taxonony
Baker, J. G. 1865-68. Synopsis Filicum. London: Robert
Hardwicke.
Ching, R C. 1940. On natural classification of the family
"Polypodiaceae". Sunyatsenia 5: 201-268.
Christ, H. 1897. Die Famkrauter der Erde. Jena: Gustav
Fischer.
Christensen, C. 1905-6. Index Filicum. Copenhagen: H. Hagerup.
Christensen, C. 1938. Chapter XX: Filicinae, in: Verdoorn,
F. Manual of Pteridology. The Hague: M. Nijhoff.
Copeland, E. B. 1947. Genera Filicum. Waltham, Mass.:
Chronica Botanica.
Engler, A., Ptant!, K. (Eds.) 1898-1902. Die natOrlichen
Pflanzenfarnilien 1,4. Leipzig: Engelmann, pp.1-80S.
Holttum, R E. 1947. A revised classification of leptosporangiate ferns. 1. Linn. Soc. Lond. Bot 53: 123-158.
Holttum, R E. 1955. A revised flora of Malaya. II. Ferns of
Malaya. Singapore: Government Printer.
Holttum, R E. 1982. Flora Malesiana II. I, 5: 7-20. The
Hague: M. Nijhoff.
Hooker, W.J. 1844-64. Species Filicum I-V. London: William Pamplin.
Kato, M. 1983. Classification of major groups of pteridophytes. J. Fac. Sci. Univ. Tokyo III, 13: 263-283.
Morton, C. V. 1968. The genera, subgenera, and sections of
the Hymenophyllaceae. Contrib. U.S. Nat Herb. 38:
153-214.
Swain, T., Cooper-Driver, G. 1973. Biochemical systematics
in the Filicopsida. Bot. J. Linn. Soc. Supp!. 67: 111-134.
Swain, T., Cooper-Driver, G. 1980: Role offlavonoids and related compounds in fern systematics. Bull. Torr. Bot C!.
107: 116-127.
Tryon, R. M. 1952. A sketch of the history of fern classificarion. Ann. Missouri Bot Gard. 39: 255-262.
Tryon, R M., Tryon, A. F. 1982. Ferns and allied plants with
special reference to tropical America. Berlin/Heidelberg!
New York: Springer.
Verdoorn F. (Ed.). 1938. Manual of pteridology. The Hague:
M.Nijhoff.
Morpbology, Phylogeny
Atkinson, L. R 1973. The gametophyte and family relationships. Bot. J. Linn. Soc. 67, Supp!. 1: 73-90.
Bierhorst, D. W. 1971. Morphology of vascular plants. New
York: Macmillan.
Bower, F. O. 1923, 1926, 1928. The ferns, 3 vols. Cambridge:
Univ. Press.
Cottbem, W. van, 1970. Comparative morphological study of
the stomata in the Filicopsida. Bull. Jardin Bot Nat Belg.
40: 81-239.
Eames, A.J. 1936. Morphology of vascular plants. Lower
groups. New York London: MacGraw-Hill.
General References for Pteridophytes
Erdtman, G. 1957. Pollen and spore morphology/plant taxonomy. Gymnospermae, Pteridophyta, Bryophyta (Illustrations). Stockholm: Almqvist and Wiksell.
Erdtman, G. 1965. Pollen and spore morphology/plant taxonomy. Gymnospermae, Bryophyta (Text). Stockholm:
Almqvist and Wiksell.
Erdtman, G. 1971. Pollen and spore morphology/plant taxonomy. Pteridophyta (Text and additional illustrations).
Stockholm: Almqvist and Wiksell.
Goebel, K. von 1930. Organographie der Pflanzen. 3. Aufl.
Zweiter Teil. Bryophyten und Pteridophyten. Jena: Gustav
Fischer.
Lugardon, B. 1972. La structure fine de l'exospore et de la
perispore des micinoes isosporees 1. Pollen Spores 14:
227-261.
Lugardon, B. 1974. La structure fine de l'exospore et de la
porispore des filicinees isosporees 2. Pollen Spores 16:
161-226.
Nayar, B. K., Kaur, S. 1971. Gametophytes of homosporous
ferns. Bot. Rev. 37: 295-396.
Smith, G. M. 1955. Cryptogamic botany. II. Bryophytes and
pteridophytes. New York: MacGraw-HilI.
Sporne, K. R. 1975. The morphology of pteridophytes. 4th
edn. London: Hutchinson.
Palaeobotany
Meyen, S. V. 1987. Fundamentals of palaeobotany. London
New York: Chapman & Hall.
Niklas, K-J. (Ed.) 1981. Paleobotany, paleoecology and evolution, 2 vols. New York: Praeger
Stein, W. E., Wigh!, D. C., Beck, C. E. 1984. Possible alternatives for the origin of Sphenopsida. Syst. Bot. 9: 102-118.
Stewart, W. N. 1983. Palaeobotany and the evolution of
plants. Cambridge: Univ. Press.
Thomas, B. A. & R. A. Spicer. 1986. The evolution and palaeobiology of land plants. London, Sydney: Croom &
Helm, Portland, Or.: Dioscorides Press.
15
Anatomy
Ogura, Y. 1972. Comparative anatomy of vegetative organs of
the pteridophytes. Handbuch der Pflanzenanatomie, 2nd
edn, Spec. Part 8, 3. Berlin: Borntrager.
Karyology
Jermy, A. C., Walker, T. G. 1985. Cytotaxonomical studies of
the ferns of Trinidad. Bull. Br. Mus. (Nat. Hist.) Bot. Ser.
13 (2): 133-276.
LOve, A., LOve, D., Pichi Sermolli, R. G. 1977. Cytotaxonomical atlas of the Pteridophyla. Vaduz: J. Cramer.
Lovis, J. D. 1977. Evolutionary patterns and processes in
ferns. Adv. Bot. Res. 4: 230-415.
Manton, I. 1950. Problems of cytology and evolution in the
pteridophytes. Cambridge: Univ. Press.
Walker, T. G. 1973. Evidence from cytology in the classification offerns. Bot. J. Linn. Soc. 67, Suppl. 1: 91-110.
Chemistry and Chemosystematies
Berti, F., Bottari, G. 1968. Constituents of ferns. In: L. Reinhold, Liwschitz, Y. (Eds.) Progress in phytochemistry,
vol. 1. London: Interscience, pp. 589-685.
Boralle, N., Braque!, P., Gottlieb, O. R. 1988. Ginkgo hi/oha: a
review of its chemical composition. In: Ginkgolides chemistry, biology, pharmacology and chemical pe~pec­
lives (P. Braquet, Ed.) vol. 1. Barcelona: J. R. Prous, SCience
Publ., pp.9-25.
Cooper-Driver, G. A., Haufler, C. 1983. The changing role of
chemistry in fern classification. Fern Gaz. 12: 283-294.
Gottlieb, o. R., Kubitzki, K- 1984. Chemosystematics of the
Gnetatae and the chemical evolution of seed plants. Planta
Med. 50: 380-385.
Hegnauer, R. 1962. Chemotaxonomie der Pflanzen, vol.l.
Basel: BirkhAuser, pp.220-317.
Hegnauer, R. 1985. Chemotaxonomie der Pflanzen, vol. 7.
Basel: Birkhauser, pp. 398-554.
Soeder, R. W. 1985. Fern constituents: including occurrence,
chemotaxonomy and physiological activity. Bot. Rev. 51:
442-536.
Swain, T., Cooper-Driver, G. A. 1973. Biochemical systematics in the FIlicopsida. Bot. J. Linn. Soc. 67, Suppl. 1:
111-134.
Glossary for Pteridophytes
K. V.KRAMER
A
Terms applying to pteridophytes that may not be generally
known, or applied in the following in a special way, or in a
few cases newly coined.
Allantodioid (of the indusium): an elongate indusium initially
completely wrapped around the sorus, which is of the same
shape, and strongly convex, attached at the anterior margin,
at maturity patent or reflexed and then approximately asplenioid, or sometimes breaking (see Fig. 68 0).
Anadromous (of the dissection pattern of the lamina or of the
venation): with the basal segment or vein(let) issuing from the
anterior side, the one on the posterior side issuing from a distinctly more distal point (as in Fig. 1 0).
Bristles: approximately cylindric epidermal appendages,
usually of the stem, thicker than most scales and not flattened
(although they may appear so in preserved material), at least
basally multiseriate and broader than hairs/trichomes.
Catadromous (of the dissection pattern of the lamina or of the
venation): with the basal segment or veinlet borne at the posterior side (as in Fig. 1 E); or sometimes the basal veinlet issuing from a vein of an order below the one that bears the adjacent, distal veinlet (as in Fig. 59 8).
Clathrate (of a scale): the cell structure having darkened partitions between adjacent cells only (Fig. 18H, I).
Ctenitoid hairs: reddish (except when young), pluricellular
hairs, especially on the axes and veins of leaves, in certain
Oryopteridoid genera like Ctenitis and Tectaria and their relatives (Fig. 600).
Dipladesmic: a special system of veins underlying the sporangia, below the regular venation (as in Fig. 112 B).
Fig.1A-E. Types of fern venation and dissection. A Gonioptemid venation. B Anaxetoid venation. C Sagenioid venation. D Anadromous dissection and venation. E CatadromOllS dissection and venation. For further explanation, see
glossary (A-C from Sadebeck 1899; D, E from Appert 1973)
Goniopteroid (of the venation): as in Fig.1A.
Intestiniform hairs/scales: flattened, largely or entirely uniseriate epidermal appendages of the lamina with strong pattition walls, upon drying contracted at the junction of adjacent
cells and appearing intestiniform, with several contractions.
Isodromous (of the dissection pattern of the lamina or of the
venation): with the basal segments or veins issuing at the same
point on opposite sides.
Lepicystis scales:usuaIly small, pale scales orthe lamina and/
or its axes, darkened at the peltate attachment, often roundish
to broadly lanceolate.
Pseudoanadromous (of the dissection pattern of the lamina):
apparently anadromous, but only so by suppression of one or
a rew basal segment(s) of the posterior side, the loss of divisions shown by the architecture of adjacent pinnae or by that
of related species.
Sagenioid (of the venation): as in Fig.1C.
Spicular cells: elongate, indurated idioblasts of the epidermis,
especially on the dorsal side, that contain silica spicules, typical of Vittariaceae; see Fig. 139 F.
Stem: the leaf-bearing organ of a pteridophyte, here used in
such a way to include the organ usually called "rhizome".
Trophopod: a thickened basal portion of a petiole functioning
as a storage organ that usually persists after the petiole above
decays.
Keys to the Families
of Fern Allies and Ferns
K. U.
KRAMER
1. Key to Families and Subkeys
1. Stem articulate, bearing whorled, partly fused, single·
veined leaves; sporangia borne several together on peltate
sporophylls aggregated in strobili; spores bearing hapters
Equisetaceae
- Stem non-articulate; leaves several~veined, Of, if singleveined, not verticillate nor partly fused; sporophylls not
aggregated in strohili, Of, if so, each one bearing only one
sporangium; spores lacking hapters
2
2. Sporangia enclosed in sporocarps, or else borne several to
many together on the lamina
6
- Sporangia borne singly in the axil or on (in) the base of a
sporophyll
3
3. Sporangia 2-or 3-locular, the wall several-layered; sporangium-bearing leaves bifid
Psilotaceae
- Sporangia unilocular (sometimes septate in /soetes), their
wall uni- or bistratose; sporangium-bearing leaves entire 4
4. Spores of one kind
Lycopodiaceae
.- Spores of two kinds
5
5. Axis of plant weak, elongate; sporangia superficial; ligule
of leaf very fugacious
Selaginellaceae
- Axis of plant thick, mostly very short and with crowded
leaves, rarely more elongate; sporangia partly sunken in
leaf base; ligule evident, persistent
Isoetaceae
6. Sporangia enclosed in sporocarps
7
Sporangia not enclosed in sporocarps
9
7. Rooting plants of seasonally dry to very moist habitats;
sporocarp wall much indurated
Marsileaceae
Free-floating aquatic plants with simple or transversely
bilobed leaves; sporocarps soft-walled
8
X. Leaves alternate, with a dorsal and a ventral lobe; true
roots present, simple; one single megasporangium per
sporocarp
Azollaceae
Leaves verticillate, each node with two simple, floating
leaves bearing large papillae, and one submerged, much
dissected, root-like leaf; several megasporangia per sporocarp
Salviniaceae
(), Lamina, apart from the axes and veins, of only one or a
rew cell layers; epidermis and stomata lacking, or the latter very few (in a few cases also the mesophyll lacking
and the lamina reduced to the veins and their sheaths):
"ft1my ferns"
Key A
Lamina of several to many cell layers; epidermis differentiated, with stomata (usually only on the lower surface of
the lamina)
10
10. Sori dorsal on the lamina, orbicular to elongate, exindusiate, each of relatively few, large, simultaneously matur-
ing, sometimes fused sporangia
Key B
- Sori not dorsal and exindusiate, or, if so, then each of nu~
merous smaller, not simultaneously maturing sporangia
11
11. Sporangia borne on special, modified segments of the
lamina, not covering the entire abaxial surface equally;
indusium generally none
Key C
- Sporangia covering equally the entire abaxial surface of
the lamina or not, not confined to special, modified segments of the lamina
12
12. Sporangia "acrostichoid", covering the entire fertile lami·
na, borne not only on the veins but also on the tissue
be~
tween them
Key D
- Sporangia either confined to definite sori or at least to the
veins and not borne on the tissue between them (beware
of sori that are confluent when mature)
13
13. Sporangia borne in definite groups on the vein-ends of a
reflexed lobe of the lamina, or also spreading between the
vein-ends, these entering the reflexed part
Adiantum (p. 249)
- Sporangia not borne on reflexed marginal lobes entered
14
by veins
14. Sporangia following the veins in groups of indefinite
length, or borne on an intramarginal vein commissure; in-
dusium wanting or simulated by flanges of leaf tissue projecting over the sporangia, or by the reflexed and ± modified leaf margin
15
- Sporangia in orbicular to elongate sori of definite length,
not bome on an intramarginal commissure and simulta-,.
neously without a true indusium
16
15. Veins free, or only joined by a submarginal commissure
Key E
which then bears the sporangia
- Veins variously reticulate, at least forming areoles flanking the costa; a submarginal commissure bearing the
sporangia present or not
Key F
16. Indusium elongate, laterally attached along a Jine to the
soriferous vein, or proximaUy to the submarginal sporan-
gia-bearing commissure
Key G
- Indusium short, attached approximately at a point, or
cup-shaped, inferior, or wanting (if the sori are short and
submarginal with the indusium opening towards the margin, or if the indusium is "double", i.e. besides the true
indusium the sori covered by the reflexed, ± modified
margin or a lobe of the margin, this heading should also
be followed)
17
17. Indusium absent
20
- Indusium present
18
18. Indusium inferior, surrounding the base of the receptacle
Key H
for at least a part of its circumference
- Indusium superior, or, if inferior, attached only at a point
at the posterior side of the receptacle, or cup-shaped, or
small and/or fugacious
19
19. Stem bearing hairs, scales wanting; sori (sub)marginal
Keyf
- Stem bearing flattened scales, hairs may be additionally
present
Key J
20. Stem bearing hairs
Key K
- Stem bearing flattened scales, hairs may be additionally
present
Key L
18
Keys to the Families of Fern Allies and Ferns
2.Subkeys
Key A: "Filmy" ferns, the lamina of one or a few cell layers,
tender, without epidermis and without or with very
few stomata; rarely the laminal parts reduced to the
veins
1. Indusium none
2
- Indusium present
3
2. Sporangia distinct, a few together forming a roundish to
Leptopteris (p. 200)
linear sorns (Fig. 103 E)
- Sporangia fused into linear synangia
Danaea (p. 178)
3. Indusium lateral, elongate, covering a linear group of spoAsplenium (p. 56)
rangia (Fig. 18 A)
- Indusium not lateral and elongate
4
4. Stem scaly; receptacle short
5
- Stem naked or hairy; receptacle cylindric or filiform
Hymenophyllaceae
5. Indusium posteriorly attached, hood-like
Cystopteris (p. 138)
- Indusium cup-shaped, attached at the base and sides
HymenophyfJopsis (p. 163)
Key B: Sporangia few, large, simultaneously maturing, assembled in dorsal, exindusiate sori or fused
1. Sporangia fused
Marattlaceae
- Sporangia distinct
2
2. Lamina pedately branched, free-veined; or forked, but the
ultimate divisions not pectinately arranged
Maloniaceae
- Lamina at least once forked (very rarely not), the ultimate
divisions roundish to linear, pectinately arranged; veins
free
Gleicheniaceae
- Lamina pinnately divided or simple
3
3. Sporangia thick-walled; annulus none; leaves without
sclerenchyma
Marattlaceae
- Sporangia thin-walled, with annulus; axes sclerotic
Cyatheaceae
Key C: Sporangia borne on special, modified segments or
lobes of segments, not assembled in sori (but not the
entire fertile lamina covered with sporangia and ±
modified)
1. Sporangia immersed into the tissue of the fertile spike;
lamina simple, lobed, or forked, with reticulate veins
(Fig.102E, F, G)
Ophioglossum (p. 196)
2
- Sporangia not immersed
2. Sporangial wall thick, multi-layered; no annulus differentiated; fertile segment inserted at the base of the lamina
Ophioglossaceae
- Sporangial wall one-layered; annulus differentiated
(though sometimes only as a small, latero-subapical group
of cells)
3
3. Stem bearing hairs; fertile segments variously distributed 4
7
- Stem bearing flattened scales; fertile segments apical
4. Sporangia borne singly on lamina lobes of otherwise modified or non-modified segments, each covered by an indusium-like flange; rachis twining, leaves scandent
Lygodium (p. 260)
- Sporangia borne in groups (trusses) or many together, the
segments bearing them always modified; rachis not twining
5
5. Lamina pinnately compound
6
- Sterile part of lamina simple or dichotomously branched
Schizaea (p. 262)
6. Annulus complete, subapical, transverse (Fig. 132 f); modified fertile segments basal
Anemia (p. 260)
- Annulus of a small latero-subapical group of cells
(Fig.l03C); modified fertile segments basal, medial or apiOsmnnda (p. 199)
cal
7. Medum-sized to large terrestrial ferns with at least once
pinnate lamina
8
- Small to medium-sized, essentially epiphytic ferns with
simple to pinnatifid or forked leaves
9
8. Leaves simply pinnate; veins forming a fine network
Acrostichum (p. 252)
- Leaves at least twice pinnate; veins free
Ua.ea (p. 247)
9. Spores (with few exceptions) monolete; leaves often scaly;
plants not hairy, or at least lacking acicular hairs
Polypodiaceae
- Spores (with few exceptions) trilete; leaves usually with
persistent acicular hairs, especially on the petiole, never
scaly
Grammitidaceae
Key D: Sporangia with "acrostichoid" distribution, not assembled in distinct sori, nor on modified segments
but evenly covering the (modified or non-modified)
fertile lamina (beware of ferns where the spomngia
are assembled in sori or at least confined to the veins
but appear acrostichoid at full maturity!)
1. Stem hairy or naked
2
- Stem bearing flattened scales
4
2. Annulus consisting of a small, subapico-Iateral group of
indurated cells (Fig. 103 C)
Osmundaceae
- Annulus subapical, transverse (Fig. 132 f) Anemia (p. 260)
- Annulus oblique or vertical
3
3. Lamina simply pinnate, free-veined
Plagiogyriaceae
- Lamina simple or bilobed, reticulate-veined
Cheiropleuria (p. 69)
4. Veins free, also in sterile leaves
5
10
- Veins anastomosing at least in the sterile leaves
5. Fertile leaves "skeletonized", i. e. the laminal parts consisting (nearly) only of the veins
6
- Fertile leaves not "skeletonized", laminal parts present
beside the veins, though sometimes reduced as compared
to sterile leaves
7
6. Stem long-scandent or creeping; veins not joined by an
Polybotrya (p. 119)
intramarginal commissure
- Stein not long-scandent (or long-creeping); veins joined
by an intramarginal commissure
Olfersia (p. 118)
7. Stem scandent
8
- Stem not scandent
9
8. Costa flanked by very narrow, inconspicuous areoles,
veins otherwise free (Fig. 25 B)
StenochJaena (p. 67)
- Costa not flanked by areoles, veins entirely free
Lomariopsidaceae
9. Spomngia confined to a paracostal commissure or to
flanges flanking the costa, only seemingly acrostichoid;
lamina rarely simple
Blechnaceae
- Sporangia truly acrostichoid; lamina not rarely simple
Lomarlopsidaceae
10. Spores trilete
11
- Spores monolete
13
11. Lamina simply pinnate; veins forming a fine network of
meshes with narrow areoles
Acrostichnm (p. 252)
- Lamina simple
12
Keys to the Families of Fern Allies and Ferns
12. Sporangia scattered
Anetium (p. 275)
- Sporangia close
Loxogramme (p. 229)
13. Anastomoses of veins confined to arches along the costa
14
- Veins more amply anastomosing
15
14. Stem creeping, stele strongly dorsiventral
Lomariopsidaceae
- Stem erect, or, if creeping, the stele not dorsiventral
Blechnaceae
15. Stem long-scandent; pinnae articulate Lomariopsidaceae
- Stem not long-scandent; pinnae not articulate
16
16. Petiole articulate at base; perispore usually little prominent; stem scales often peltate
Polypodiaceae
- Petiole not articulate at base, or, if so, the perispore prominent; scales of stem basally attached or occasionally
17
pseudopeltate
17. Stele of stem strongly dorsiventral; leaves borne only on
its dorsal side
Lomariopsidaceae
18
- Stele of stem weakly or not dorsiventral
18. Veins reticulate with free included veinlets; lamina bearing articulate hairs
Dryopteridaceae
- Veins free, or reticulate without free included veinlets;
lamina without articulate hairs but often scaly
Elaphog/osmm (p. 169)
Key E: Sporangia following the veins ("gymnogrammoid") or
on a submarginal commissure; no indusium; veins
free or joined by a submarginal commissure
1. Stem naked when mature
2
- Stem hairy or scaly
3
2. Small epiphytes, often hairy on the leaves, these non-dimorphic
Grammitidaceae
- Medium-sized to large terrestrial ferns; leaves glabrous,
with strong sterile-fertile dimorphism (Fig. 105 A)
Plagiogyriaceae
3.Stem hairy (hairs may be pluriseriate at base: "bristles",
but not flattened)
4
- Stem bearing flattened scales
5
4. Annulus transverse, subapical, complete (Fig. 132 F)
Anemia (p. 260)
. Annulus vertical, interrupted
Pleridaceae
Annulus latero-subapical, consisting of a small group of
cells (Fig 103 C)
Osmundaceae
5. Small to medium-sized epiphytes with simple, lobed, or
forked lamina
6
Terrestrial, mostly large ferns with most often pinnatifid
to more compound leaves
8
6. Lamina simple to pinnatifid, usually bearing acicular
hairs at least on the petiole; spores mostly trilete, chlorophyllous
Grammitidaceae
Lamina simple, or forked, hairless; spores achlorophyllous
7
7. Spicular cells present in leaves (Fig. 139 F)
Vittariaceae
Spicular cells none
Loxogramme (p. 229)
N. Sori dorsal
9
Sori submarginal
Pteridaceae
9. Spores trilete
Pteridaceae
Spores monolete
10
III. Petiole articulate to the stem (phyllopodium), with several
vascular bundles; lamina often bearing peltate scales
Polypodiaceae
Petiole without basal articulation, with one or two vascu11
lar bundles; peltate scales none
19
11. Petiole at base with two strap-shaped xylem strands
Thelypteridaceae
- Petiole at base with two C-shaped xylem strands back to
back,or with one x- or butterfly-shaped one (Fig. 18F)
Asplenium (p. 56)
Key F: Sporangia following the veins ("gymnogrammoid") or
on a submarginal commissure; indusium wanting, or
consisting of the reflexed, ± modified leaf margin only; veins variously reticulate but not only joined by a
submarginal commissure
1. Epiphytic or epilithic ferns with an articulation at the base
of the petiole (phyllopodium); spores usually monolete
Polypodiaceae
- Terrestrial or epiphytic ferns without an articulation at the
petiole base
2
2. Epiphytic (or epilithic) ferns with simple lamina
3
- Terrestrial (occasionally epiIithic) ferns with mostly compound lamina
5
3. Acicular hairs present and mostly persistent on the petiole
and sometimes on the lamina; spores chlorophyllous, trilete
Grammitidaceae
- Acicular hairs wanting; spores monolete or trilete, achlorophyllous
4
4. Spicular cells presentin the leaf (Fig. 139 F) Vittariaceae
- Spicular cells none
Loxogramme(p.229)
5. Reticulation of veins consisting only of areoles flanking
the costa
6
- Veins more amply reticulate
7
6. Sporangia borne on a commissure flanking the costa
Blechnaceae
- Sporangia borne on a submarginal commissure
Pteridaceae
7. Venation reticulate with free included veinlets pointing to
1ectaria (p. 119)
all sides
- Venation without free included veinlets, or with only excurrent veinlets
8
8. Sporangia confined to a submarginal commissure; free included veinlets none
9
- Sporangial lines dorsal
10
9. Stem bearing flattened scales
Pteridaceae
- Stem bearing hairs which may be basally pluriseriate but
not flattened
Dennslaedliaceae
10. Sporangia in a (sometimes interrupted) line parallel to the
Taenitis (p. 240)
costa on either side of it
- Sporangia not so arranged
11
11. Spores monolete
12
- Spores trilete
Pleridaceae
12. Dissection and venation pattern catadromous; petiole
with only two vascular bundles
Thelypteridaceae.
- Dissection pattern anadromous at base; petiole with at
least three vascular bundles
Dryopleridaceae
Key G: Sporangia assembled in definite, elongate sori, these
with elongate, unilaterally attached indusium •
1. Sporangia placed on and indusium attached to the side of
meshes flanking the costa or to a paracostal commissure;
indusium opening towards the costa
Blechnaceae
- Sporangia placed on and indusium attached to a submarginal commissure; indusium a "true" indusium, i. e. not solely consisting of the reflexed, ± modified leaf margin but
opening towards the margin
2
20
Keys to the Families of Fern Allies and Ferns
- Sporangia placed on and indusium laterally arrached to
free or reticulate veins, the indusium opens laterally on the
laminal surface (rarely pronouncedly towards the costa) 5
2. Veins anastomosing, the meshes with free included veinlets; lamina simple
Tecf11.ria ("Dictyoxiphium'J(p.119)
- Veins free, or if anastomosing, without free included veinlets
3
3. Petiole at base articulate to the stem
Da.alIiaceae
4
- Petiole non-articulate at the base
4. Spores monolete, with prominent perispore; petiole with
two vascular bundles (except in very small species); stem
scaly
5
_. Spores trilete, or, if monolete, without prominent perispore; petiole with one vascular bundle; stem scaly or hairy
Dennstaedtiaceae
5. Vascular bundles at petiole base C-shaped, upward merging into one X-shaped or butterfly-shaped one (Fig. 18F);
stem scales clathrate or with occluded cell lumina; or petiole not rarely dark-sclerotic
Aspleniaceae
- Vascular bundles at petiole base strap-shaped, upward (or
in the rachis) fusing into one gutter-shaped one; scales not
or hardly clathrate; petiole rarely dark-sclerotic
Dryopteridaceae-Athyrioideae
Key H: Sori approximately orbicular; indusium present, inferior, surrounding at least a part of the receptacle at its
base, i. e. not only attached at the posterior side; but
indusium not cup-shaped
1. Veins free, or forming costal arches; free included veinlets
2
none
- Veins copiously reticulate, with free included veinlets
3
2. Spores trilete; habit usually that of a tree-fern Cyatheaceae
- Spores monolete; stem may be erect but hardly tree-like
Dryopteridaceae
3. Sori dorsal; lamina simple or lobed (Fig. 58 A) Caraibic reHypoderris (p. 123)
gion
- Sori terminal; veins projecting beyond the margin; lamina
pinnate (Fig. 59 A, B); New Caledonia
Tecf11.ria ("Oonidium'J(p. 119)
Key I: Sori approximately orbicular, indusiate with non-inferior indusium; stem not scaly but bearing hairs (which
may be pluriseriate at the base but not flattened; "bristles")
1. Sporangia borne on a columnar receptacle (Fig. 87 B, C)
Loxomataceae
- Sporangia borne on a flat or convex but not columnar receptacle
2
2. Bow of annulus slightly oblique, continuous, not interrupted by the stalk; stem often trunk-like
Dicksoniaceae
- Bow of annulus vertical, interrupted by the stalk and also
around the stomium; stem not trunk-like Dennstaedtiaceae
Key J: Sori round(ish), indusiate, with non-inferior indusium;
stem bearing flattened scales
1. Fertile leaves with strongly contracted and modified lamina, its edges recurved over and enclosing the indusiate
sori (Fig. 72A, B, 73A, C)
Dryopteridaceae-Athyrioideae-Onocleeae
- Fertile leaves not so formed
2
2. Indusium consisting only of reflexed, ± modified lobes
of the lamina margin ("outer" or "false" indusium), opening inward; no "inner" Of "true" indusium present
3
- A "true" indusium present; a "false" one may be present
besides
4
3. Sporangia few, with transverse, subapical annulus
Mohria(p.261)
- Sporangia one to many per sorus, with vertical, interrupted annulus
Pteridaceae
4. Petiole articulate at its base or higher
5
- Petiole non-articulate but pinna bases articulate
Nephrolepis (p. 190)
- Neither petiole nor pinna bases articulate
6
5. Petiole articulate at its base; sori (sub)terrninal on veins,
(sub)marginal on the lamina; indusium pouch- or shellshaped; lamina usually dissected; pinna bases non-articulate
Davalliaceae
- Petiole articulate usually above its base; sori dorsal (submarginal only in very narrow-leaved species); indusium
reniform to hippocrepiform-suborbicular; lamina simple
(Fig. 100 C)
Oleandra (p. 191)
- Petiole articulate above its base (Fig.l00A); sori terminal; indusium reniform-hippocrepiform; lamina pinnate,
Arthropteris (p. 192)
pinnae articulate at base
6. Sori terminal, submarginal, or projecting beyond the margin; indusium cup-shaped or attached by a narrow base
and free towards the margin of the leaf(let)
7
- Sori dorsal or subterminal, but then without this type of
indusium
9
7. Spores monolete, with prominent perispore
8
- Spores trilete
Dennstaedtiaceae
8. Veins free; lamina (venation) anadromous throughout
Asplenium (p. 56)
- Veins anastomosing; lamina catadromous except for the
lowennost divisions
Dryopteridaceae
9. Petiole base with at least three (mostly more) vascular
bundles
Dryopteridaceae
- Petiole base with two strap-shaped vascular bundles
10
10. Lamina and venation catadromous throughout; stem
. Thelypteridaceae
scales ± hairy
- Lamina usually at least partly anadromous; stem scales
not bearing hairs
Dryopteridaceae-Athyrioldeae
Key K: Sori round(ish), exindusiate; stem bearing (sometimes
basally pluriseriate but not flattened) hairs, these
sometimes minute and the stem then seemingly
naked; flattened scales none
1. Veins amply reticulate, with free included veinlets pointing
to all sides; stem solenostelic; spores monolete
Dipteris(p.l0l)
- Veins free; stem and spores various
2
2. Stem long-creeping, solenostelic; spores monolete
Hypokpis (p. 87)
- Stem usually short-creeping or erect and dictyostelic;
spores trilete
3
3. Annulus vertical, incomplete, interrupted by the stalk and
around the stomium
4
5
- Annulus slightly oblique or transversal, complete
4. Stem bearing minute, few-celled, very inconspicuous hairs
only, seemingly nude (the lamina similarly hairy)
Monachasorum (p. 187)
(Fig. 98 C)
Pteridaceae
- Stem distinctly hairy with pluricellular hairs
Keys to the Families of Fern Allies and Ferns
5. Annulus transversal, subapical (Fig: 132 f) Anemia (p. 260)
- Annulus slightly oblique, almost vertical
6
6. Leaves simply pinnate, pinnae serrate (Fig. 97 A)
Metaxya(p.127)
- Leaves much more dissected, segments crenate-pinnatifid
(Fig. 86 B)
Loplwsoria (p. 171)
Key L: Sori approximately orbicular, exindusiate; stem bearing flattened scales
21
7. Annulus not quite vertical, bypassing the stalk, habit
usually tree fem-like
Cyatheaceae
- Annulus verticual, interrupted by and not bypassing the
stalk; habit scarcely or not tree-fero-like
8
8. Spores trilete
9
- Spores monolete
12
9. Terrestrial or epilithic feros; sporangial stalk 2- or 3-seriate
Pteridaceae
- Epiphytic (or epilithic) ferns; sporangial stalk at least
partly uniserate
10
10. Acicular hairs mostly present at least on the petiole;
1. Epiphytic or epilithic ferns with an articulation at the
petiole base; spores with few exceptions monolete
2
- Terrestrial, epiphytic, epilithic, or scandent ferns without
articulation at the petiole base; spores various
4
2. Pinnae articulate to the rachis; veins free or nearly so
OIeandraceae
- Pinnae not articulate to the rachis, Of, if so, the veins amply reticulate
3
3. Leaves finely dissected; sori (sub)marginal; leaf axes
adaxially sulcate, the groove convex in the middle
Gymnogrammitis (p. SO)
- These characters not combined
Polypodiaceae
4. Stem scandent; veins free, pinnae articulate at the base 5
- These characters not combined
6
5. Fertile leaves much contracted
Jhysanosoria(p.166)
- Fertile leaves not contracted
OIeandraceae
6. At least some stem scales peltate at base
Polypodiaceae
- All stem scales basifixed
7
spores chlorophyllous
Grammitidaeeae
- Acicular hairs wanting; spores achlorophyllous
11
11. Veins free; lamina of at least the larger leaves cleft or pinnate
Vittariaceae
- Veins reticulate; lamina simple
Loxogramme (p. 229)
12. Epiphytes; lamina simple or pectinate-pinnate
13
- Terrestrial ferns; lamina usually pinnate + pinnatifid or
more dissected
14
13. Acicular hairs mostly present at least on the petiole; stem
scales mostly also bearing acicular hairs Grammitidaceae
Polypodiaceae
- Acicular hairs wanting
14. Petiole base with at least three, usually more, vascular
bundles
Dryopteridaceae
15
- Petiole base with two vascular bundles
15. Stem scales hairy andlor glandular; lamina entirely
catadromous
Thelypteridaceae
- Stem scales not hairy (but may bear other marginal appendages), non-glandular; lamina usually at least in part
Dryopteridacea~Athyrioideae
anadromous
Psilotatae
Sporophytes rootless, with dichotomous rhizomes and
aerial branches; leaves, scale-like or leaf-like_ Protostelic. Sporangia thick-walIed, homosporous. Gametophytes axial. Antherowids multiflagellate.
A single orderl and family:
Psilotaceae
K. U.KRAMER
Psilotaceae Kanitz, Nov/my. Att: 43 (1887); White (1977).
Tmesipteridaceae Bertrand, Koidzumi (1939).
Small to medium-sized, often epiphytic plants with
subterranean, dichotomously to laterally branched
stems with endophytic mycorrhiza, without roots, and
aerial, simple or dichotomously branched stems; stele
protostelic, or solenostelic at the transition of subterranean and aerial stem. Aerial stems green, erect or
pendulous, glabrous, slender, angular to sulcate or flattened, the basal part often with rudimentary leaves, the
upper part leafy; leaves spirally arranged or distichous, flaccid or firm, narrow, simple or once forked,
one-veined or veinless. Sporangia sessile on or above
the base of forked sporophylls, thick-walled (eusporangiate), 2- or 3-celled and -lobed, each cell dehiscing
longitudinally with a slit. Spores numerous, small,
achlorophyllous, pale, bean-shaped, monolete. Gametophyte subterranean, heterotrophic.
MORPHOLOGY AND ANATOMY. The external and internal morphology and anatomy have been extensivly
studied (though mostly in Ps. nudum only) because of
the enigmatic and still controversial systematic position of the family. The poorly defined leaves have
been interpreted as reduced lateral axes of limited
growth; or the entire aerial stems have been 'regarded
as homologous to certain kinds of so-called non-ap-
In isolated taxa, which occupy a high taxonomic rank, we
do not attempt to give diagnoses for all subordinate categories, thus avoiding the farce, as Burtt has said, of trying to decide which characters of a single genus should diagnose a
1
family, order, class, etc.
pendicular fern leaves as found in, e. g., Stromatopteris
(Bierhorst 1968 and later). This was much contested,
especially by Kaplan (in White 1977) and is not generally accepted at present. The sporangia are generally
called "synangia", which seems plausible; but, as
pointed out by Eames (1936), the evidence for secondary fusion is slight, and the term "synangium", with its
morphophyletic implications, is avoided here.
Much of the interpretation of the morphology also
depends on whether the Psilotaceae are regarded as
primitive or as derived and simplified (or as a combination of both; e. g. Jonker 1973). The complete lack
of fossil remains makes the decision highly uncertain.
The strongly developed mycotrophy is very suggestive
of reduction, as found in many mycotrophic angiosperms. For a full account of the morphology and
anatomy, see especially Bierhorst (1954 b, 1956, 1971)
and various authors in White (1977). For the spores
see Lugardon (1973, 1979).
GAMETOPHYTE. See again Bierhorst (1954 a, 1971). The
prothallium in both genera is subterranean, non-green,
mycotrophic, ± cylindric and often branched, radially
organized, and consists only of parenchymatic, mycorrhizal parenchyma or has a very weakly developed, often discontinuous, central stele; it sometimes bears
gemmae. Antheridia and archegonia are both borne
diffusely and in large number all over the surface. The
antheridia are rather like those of primitive ferns, with
a one-layered waIl of many ceIls and an opercular cell;
the inner mass may be up to 512-ceIled; the numerous
spermatozoids produced are multi ciliate. The archegonia are sunken, with a short, thick neck most of which
is shed towards maturity (Bierhorst 1954 a).
KARYOWGY AND HYBRIDIZATION. The chromosome
number is known for both genera, the diploid number
being based on n = 52, polyploids (up to octoploid level) being more frequent. No hybrids have so far been
reported.
EcOLOGY AND DISTRIBUTiON. Both genera may grow
terrestriaIly but are more often epilithic or epiphytic.
Tmesipteris often occurs on tree-fern trunks. Both genera mostly grow at lower elevations. See further under
the genera.
AFFINITY. Ideas about this have changed much over
the years. Pritzel (in Engler and Prantl 1902) upheld
23
Psilotatae • Psilotaceae
B,C
i\
E,F
"
... Fig. 2 A-G. Isoetaceae, Lycopodiaceae, spores. A Isoeles cu·
bana, megaspore (x 150). B Isoeles macrospora, megaspore
( x 100). C Isoeles andicola, microspore ( x 1000). D Lycopodium deulerodensum (x 1000). E Lycopodium c1avatum
( x 1000). F Lycopodium obscurum ( x 1000). G Huperzia lini·
folia (x 1000). Phot. A. F. Tryon
Fig. 3 A-C. Selaginellaceae, spores. A Selaginella apoda, microspore (x 1000). B Selaginella galeottii, megaspore with
microspore attached (x 100). C Selaginella selaginoides, microspore (c. x 100). Phot. A. F. Tryon
Ii
T
B
C
Psilotatae . Psilotaceae
24
the idea of affinity to Lycopodiales, taken up again by,
e. g. Jonker (1973). Following a suggestion by Lam
(1948), Ps. were compared with Psilophyta (perhaps
prompted to some extent by the similarity in name)
which is now generally rejected. Bierhorst (1968, 1973,
1977) emphasized similarity to ferns, especially Stromatopteris, which had not been previously considered.
Fig. 4 A-F. Psilotaceae. A, B Psi/olum comp/anatum. A Part
of branch system (x 0.8); B Dehisced sporangium (x 12).
C, D Tmesipteris tnmcata. C Part of branch system (x 1.1).
D Dehisced sporangium (x 12). E, F Tmesipteris sp. E Gametophyte (g) with young sporophyte (sp) (enlarged). F Apex
of gametophyte with archegonia (aT) and antheridia (an) (enlarged) (A-D from Brownlie 1977; E, F from Eames 1936)
A
This idea was much opposed (see above) also on phytochemical grounds (Cooper-Driver 1977; Wallace et al.
1983). But the idea of a certain degree of affinity to
ferns gained some ground (e.g. Tryon and Tryon
1982), also because of similarity in spore structure (Lugardon 1973, 1979). The Ps. diverge, however, too far
from ferns to be included in this class and merit recognition as a class in its own right (Wagner in White
1977).
KEy
TO THE GENERA.
1. Leaves scale- or awl-like, without vein, or with one at the
extreme base; sporangium 3-celled, on the base ofthe bifid
sporophyll
1. Psilotum
- Leaves tongue-shaped to lanceolate, one-veined; sporan-
gium 2-celled, inserted on the sporophyll above its base, at
2. Tmesipteris
the bifurcation
Psi/otum· Tmesipteris
1. Psilotum Sw.
25
Fig. 4
Psi/olUm Sw.• Schrad. J. Bot. 1800': 8, 109 (1802); David
(1965); Reed (1966).
Subterranean stems much branched, with gemmae.
Aerial stems slender, well-developed, to c. 1 m long
but mostly shorter, dichotomously branched, with awlshaped to squamiform leaves, these without veins or
with a vein in the extreme basal part. Sporangia
(sub)sessile, globose, later 3-lobed, 3-celled. Spores ellipsoidal, coarsely rugose.
Several species described, but probably only two
really distinct. Pantropic to warm-temperate except in
dry areas; extending to the SE United States, extreme
SW Europe, central Japan, and South Korea; occurring on many remote, especially Pacific, islands. The
most widespread is Ps. nudum (L.) Pal. Beauv.
2. Tmesipteris Swartz
Figs.4C-F, 5 D
Tmesipteris Swartz, Schrad. J. Bot. 1800': 131 (1802); Reed
(1966); Chinnock (1975,1976); Braithwaite (1986).
Terrestrial or more often epiphytic (often on tree-fern
trunks) plants with little branched underground stems
without gemmae. Aerial stems often short but may
reach c. !h m, often simple, erect to drooping or pendulous, very leafy except at base; in some species indefinite, in others terminated by a leaf; leaves subsessile,
narrowly ovate to lanceolate, or sigmoid, obtuse to
acute, often mucronate. Sporophylls with deeply bifid
lamina, aggregated either in the apical or in the basal
part of the stem, the lobes similar in shape to sterile
leaves. Sporangia sessile at the bifurcation, 2-celled,
the cells rounded or apiculate, equal or unequal.
Spores ellipsoidal, shallowly rugose.
C. 10 species, some as yet imperfectly known, in SE
Asia-Australasia, concentrated in Australia and
New Zealand, extending to New Caledonia, New Guinea, the Philippines, east to the Marquesas and Tahiti.
Selected Bibliography
lIierhorst, D. W. 1954 a. The gametangia and embryo of Psilolum nudum. Amer. J. Bot. 41: 274-281.
lIierhorst, D. W. 1954 b. The subterranean sporophytic axes
of Psilotum nudum. Amer. J. Bot. 41: 732-739.
Bierhorst, D. W. 1956. Observations on the axial appendages
in the Psilotaceae. Phytomorphology 6: 176-184.
Bierhorst, D. W. 1968. On the Stromatopteridaceae
(fam. nov.) and on the Psilotaceae. Phytomorphology 18:
232-268.
Bicrhorst, D. W. 1969. On Stromatopteris and its ill-defined
organs. Amer. J. Bot. 56: 160-174.
lIicrhorst, D. W. 1973. Non-appendicular fronds in the Fili____, _
cales. Bot. J. Linn. Soc. 67 Suppl. 1: 45-57. _
IIraithwaite, A F. 1986. Tmesipleris in Vanuatu (New
Hebrides). Fern Gaz.13: 87-96.
Fig.5. Psilotaceae. Tmesipleris bi/lardieri epiphytic on stem of
Dicksonia; Victoria, Australia. Phot. K.. U. Kramer
Chinnock, R.J. 1975. The New Zealand species of Tmesipleris
(Psilotaceae). New Zeal. J. Bot. 13: 743-768.
Chinnock, R J. 1976. The identification, typification and origin of Tmesipteris lannensis (Psilotaceae). Taxon 25:
115-121.
Cooper-Driver, G. 1977. Chemical evidence for separating
the Psilotaceae from the Filicales. Science 198: 1260-1261.
David, A 1965. Contribution Ii l'etude des Psilotales. Lyon.
Eames, AJ. 1936. Morphology of vascular plants. Lower
groups. New York, London: McGraw-Hill.
Jonker, F. P. 1973. The taxonomic position of the Psilotales in
the light of our knowledge of Devonian plant life. Palaeobotanist 20: 33-38.
Lam, H. J. 1948. Classification and the new morphology. Acta
Biotheor.8: 107-154.
Lugardon, B. 1973. Sur les parois sporales de PsilolUm triquetrum Sw. et leur structure fine. C. R. Acad. Sci. Paris Ser. D
276: 1277-1280.
Lugardon, B. 1979. Sur la formation du sporoderme chez
Psilolum triquetrum Sw. (Psilotaceae). Grana 8: 145-165.
Reed, C. F. 1966. Index Psilotales. Bol. Soc. Brot 11.40:
71-96.
Wallace, 1. W., Pozner, R. S., Gomez, L. D. ·1983. A phytochemical approach to the Gleicheniaceae. Amer. 1. Bot. 70:
207-211.
White, R. A (Ed.). 1977. Taxonomic and morphological relationships of the Psilotaceae: introduction to the symposium. Brittonia 29: 1-68.
Lycopodiatae
Sporophytes with roots, stems and spirally arranged
leaves (microphylls). Protostelic, sometimes polystelic.
Sporangium thick-walled, homosporous Or heterosporous, either borne on a microsporophyll or associated
with one. Some with a water-absorbing ligule. Antherozoids biflagellate or multi flagellate.
Three isolated families, each fonning an order of itself. The ligulate families Isoetaceae and Selaginellaceae more closely related.
lsoetaceae
A. C.JERMY
Isoetaceae Dumortier, Anal. Fam. PI.: 69 (1829) las Isoetineae); Reed (1953); Fuchs (1962).
longitudinal rows above the air canals; ligule narrowly
triangular to ovate-deltate, arising from a sac-like depression on the adaxial face of the leaf base, above the
sporangium when this is present; blade sometimes vestigial and leaf replaced by phyllopodia fonned by persistent scale-like base (Fig. 6). Fertile leaves produced
either continuously throughout the year, or seasonally,
bearing sporangia which are at least initially embedded in the adaxial face of the leaf base, below the
ligule; sporangia ovoid or globose, up to 20 mm long,
surface often appearing foveolate, epithelium cells
thin-walled, with occasional tannin-containing or
thick-walled cells, but without any mechanism for dehiscence, partially septate or with columnar structures
(trabeculae) extending transversely from wall to wall,
sometimes partially or entirely covered by a thin membrane (velum) which may be developed apically fonning the labium or pseudoligule; spores of two kinds,
usually in separate sporangia, but occasionally both in
the same one; megaspores 250-800!.1Ill in equatorial
diam., often varying considerably within a species and
within a single sporangium, trilete, with pronounced
laesurae and equatorial ridge, outer sporodenn containing much silica, fibrillose in texture but variously
patterned: echinate, rugulate, cristate, reticulate, tuberculate, pustulate, laevigate, saccate, clavate, verrucate,
baculate Or retate; microspores 20-40 ].Lm long, monolete, variously patterned: echinate, verrucate, tuberculate, laevigate, scabrate or papillate.
An ancient and widespread family comprising a single genus of some 130 species in which there is considerable endemism due to ecological isolation.
Terrestrial or aquatic plants, either evergreen, Or in
seasonally wet habitats, losing their leaves in the dry
periods. Stem covered by an irregular cortex layer
which sloughs off periodically as new layers are
fonned from a secondary meristem below, often producing bulbils at the edges of epidennallesions, usually erect, elongate or short and conn-like, bilobed or trilobed in transection, or more rarely short-creeping and
rhizome-like; apices naked, usually sunken. Roots
arising from a basipetal growing point beneath the
"conn", or from one Or more median grooves running
longitudinally up the stem from that growing point.
Leaves spirally arranged, 2-100 cm long, up to 10 mm
wide, occasionally filifonn, either terete, often winged
below, with a subulate or tapered apex, few to many, ANATOMY AND MORPHOLOGY. Isoetes lacuslris L. was a
arising in an apical tuft or open rosette, or more Or less subject of much general study by nineteenth century
flat, linear, with a spathulate or rounded, apiculate morphologists. Weber (1922) was one of the first to reapex, fonning an imbricate rosette at the stem apex; view the anatomy of the genus in relation to the systebase frequently spathulate and much broader than the matics, especially of South American species. Rauh
blade, often colourless with broad membranous mar- and Falk (1959) published a very comprehensive acgins Or with blackish-brown sclerotic patches that .count of the morphology and anatomy of 1. andicola
harden and remain as usually tricuspate scales which agg. A detailed account of the developmental anatomy
protect the stem growing point during drought peri- of three North American species was presented by
ods; occasionally viviparous on leaf margin; vascular Paolillo (1963). Other important studies in which the
strands running centrally throughout the length of the usefulness of morphological characters are discussed
leaf, surrounded by four longitudinal air canals which include those of species of the southeastern USA by
are separated from each other by parenchyma and di- Boom (1982), of South America by, Fuchs (1982), of
vided by transverse septa; stomata, when present, in Mrica by Hall (1971), Jenny and Schelpe (1986) and
Lycopodiatae . Isoetaceae
Wanntorp (1970), of India by Panigrahi (1981), and of
Australasia by Croft (1980), Johnson (1984) and Marsden (1976 b). The evolution and morphological diversity of the genus is discussed by Hickey (1986 b).
The roots possess a single vascular bundle, are dichotomously branched, and arise from the stele in a
characteristic way that has much in common with Stigmarian ancestors.
All leaves have the potential to become fertile, there
being no morphological differences between the
blades of sterile leaves and sporophylls; usually those
leaves first formed after a rest period are sterile. In
some semi-aquatic and terrestrial species (e. g. I. histrix
Dur., I. eshbaughii Hickey) the blade of such leaves is
reduced to a vestigial subula at the apex of a broad,
often horny, base (phyllopodium). These horny bases,
seen also on fertile leaves, persist on the corm for
several years, and in some forms of I. histrix, frequent
in seasonal pools in northwest Africa, protect the dormant but succulent corm against foraging animals.
Species of these habitats are also characterized by having elongate cOllenchymatous cells, longitudinally
aligned, just beneath the leaf epidermis, a feature
which helps in supporting the leaf when emerged
(Hickey 1986b). The presence or absence of stomata,
and their size and shape, are not consistent within a
species but are not directly related to habitat. Stomata
are usually absent from the leaves of aquatic species,
or, if present, are non-functional (Maserati and Thomas 1982). They may also be absent from those of
some terrestrial species [e. g. I. andicola (Amstutz)
L. D. Gomez]. When present they are situated in longitudinal bands over the air canals. The function of the
ligule, which frequently withers at an early age, is unclear. Mucous is secreted from the ligular pit, at least
in the young leaf (Michaux 1973) and it has been suggested that such secretion may increase nutrient flow
to the young leaf meristem. Ligule shape has been
used by Hickey (1986) to distinguish between taxa in
the I. andina complex but generally a more useful taxonomic character is the pseudoligule (labium) (Deroux and Pitot 1965) which is associated with the
sporangial wall or velum tissue. It varies in shape and
size, frequently persisting after the ligule has withered.
Studies by Hall (1971) and Jermy (unpub!.) on African
srecies suggest this can be a diagnostic character and
worthy of further investigation. The extent to which
Ihe velum itself covers the sporangium varies with the
age of the plant and of the individual sporophyll (G6mez 1980; Jermy, unpub!.) and should be used as a
laxonomic character only with caution. Pigmentation
or the sporangial wall has been shown by Hickey
( t 986 b) to correlate with the presence of leaf scales
or phyllopodia, and he suggests that this is a primiIive character which has been lost in a number of lineages.
27
SPORE MORPHOLOGY. There is considerable variation
in sporoderm morphology of both megaspores and
microspores (Marsden 1976 b; Tryon and Tryon 1982;
Hickey 1986 c). That of the megaspore was used by
Pfeiffer (1922) to delimit major sections within the genus and has been the principal character used in identifying species in many floras and regional accounts
(e. g. Berthet and Lecocq 1977). As in Selaginella, crystalline silica constitutes a major part of the megaspore
wall (Robert et a!. 1973), and the outer layers are frequently damaged in herbarium specimens. In addition
to sporoderm sculpturing, the form of the equatorial
and proximal (triradiate) ridges can be a diagnostic
feature. There is strong evidence that many sporoderm
patterns are convergent (Hickey 1986 c) and hybridization studies reported by Taylor et a!. (1985) suggest
that different sections of the genus as defined by sporoderm morphology are not as isolated as previously
believed.
GAMETOPHYTE AND SPORE GERMINATION. Gametophytes are endosporic. Rhizoids are formed in both
terrestrial and aquatic species but are absent in species
known to be decaploid. Other distinct differences,
such as the number of archegonia, are seen between
species (Taylor et a!. 1985). Kott and Britton (1980)
compared the germination requirements of five North
American species and found that megaspores which
had been subjected to low temperatures (as over a
hard winter in nature) delayed their germination sufficiently to coincide with the germination of microspores produced in the spring.
PHYTOCHEMISTRY. Few studies have been done on the
species of lsoetes. Voirin and Jay (1978) looked at the
flavones of four unrelated European species and
found that the compound groupings were species-specific. Studies made on North American species, however, suggest that there is more variation within a species than between species (Kott and Britton 1982 b).
KARYOLOGY AND HYBRIDIZATION. Chromosome numbers based on x = 10 or x = 11 have been recorded for
material from all continents. The first number has only
been recorded for l. histrix from England (Manton
1950) and Spain and Morocco (Gibby 1989). Chromosome counts based on x = 11 have been mostly at the
diploid level with records of some tetraploids in closely related species. One hexaploid (I. sampathkumarini
Rao) is recorded from India (Abraham and Ninan
1958). Hybrids have been shown to exist by Taylor et
a!. (1985) between the North American diploid species
l. echinospora Durieu and l. engelmannii A Br.
(= I. x eatonii Dodge) and between the tetraploid species I. riparia Engelm. and I. tuckermanii A A. Eaton
( = I. x harveyi A A Eaton). The authors suggest that
I. riparia is the result of doubling of chromosomes in
28
Lycopodiatae . Isoetaceae
I. x eatonii. A hybrid has also been shown to exist between I. engelmannii and I. tuckermanii. Triploidy coupled with agamospermy has been reported for an Indian species, I. coromandelina L. (Abraham and Ninan
1958). Plants with additional supernumerary chromosomes are recorded for several Indian species (Pant
and Srivastava 1965; Goswami 1974) and need further
investigation.
The circumboreal aggregate of I.lacustris has a decaploid number of 2n = 110 both in Europe and in
North America (as I. macrospora Durieu; Kott and
Britton 1980). Hickey (1984) recorded 2n =126-132 in
I. novo-granadensis H. P. Fuchs from Ecuador, a presumed dodecaploid which needs further investigation.
I. andicola (subgen. Stylites) has been reported by
Rauh and Falk (1959) to have c. 48-52 chromosomes
but Hickey (1984) found it to have 2n = 44, a number
that Gibby (1990) found in its near-relative, I.lech/eri
Mett.
The great range of megaspore sizes, found in many
species and frequently reported, is indicative of cytological unbalance and subsequent sterility (Abraham
and Ninan 1958; Taylor et al. 1985). Sporogenesis, especially in sporangia containing both mega- and microspores (Goswami 1974), and its role in Isoetes evolution need further investigation.
SUBDIVISION. Isoetaceae were regarded as having a
single extant genus until Stylites was discovered in the
Peruvian Andes in 1954 by Raub (Raub and Falk
1959) and independently, in the same locality, by Amstutz in 1956 (Amstutz 1957). Stylites andicola Amstutz
( = I. andicola) has been shown by several workers
(Gomez 1980; Kubitzki and Borchert 1964) to exhibit
the end of a range of variation shown by other South
American Isoetes species (e. g. I.lechleri Mett.) and
therefore is recognized at a subgeneric level only.
DISTRIBUTION, ECOLOGY AND PHYSIOLOGY. Isoetes is
cosmopolitan and widespread, usually as plants of
damper habitats such as springs and seepage channels,
seasonal pools, flowing water and permanent lakes.
Unfortunately many populations have been described
in isolation as endemic species, thus Obscuring wider
relationships and distribution patterns. There are certainly similarities between taxa in Africa, India and
Australia which point to a Gondwanaland origin of
the genus (Jermy, unpubl.; Marsden 1976a). Species
are found in the cold-temperate zones in both hemipheres as submerged aquatics, but in warm-temperate
and subtropical areas which experience a seasonal dry
period (e. g. much of Africa but particularly the south,
the Mediterranean region, the Indian subcontinent
and Australia). Isoetes is often terrestrial, persisting
throughout the dry period as a dormant "corm" beneath the soil surface. I. durieui Bory and I. histrix
Bory, two species characteristic of grassy hollows and
maritime cliff tops in the more oceanic climate of SW
Europe, still interestingly retain this feature of dying
down below ground level in the summer, although
there is no severe drought and the surrounding grasses
and forbs remain green.
Some species (e. g. I. echinospora Durieu, I. herzogii
Weber) can form a close turf on lake bottoms where
competition is low due to scarcity of available nutrients. Detachment of the sporophyll, either by disintegration of the stem or by wave action in the shallower water, allows dispersal by birds which forage
amongst the drift-line debris. Karrfalt and Hunter
(1980) observed in l. andicola that detached mucilagecovered sporophylls were squeezed out of the tussock
by the growth of new leaves and were blown Or
washed away with the sporangium still attached. In
such species, spores are released only on distintegration of the sporangium wall, which was observed in
cultured African material to be the last tissue to decay
(Jermy, unpubl.). Vegetative spread by gemmae
formed at the base of the leaf is prevalent in the northtemperate l. /acustris, allowing it to colonize a lake system quickly. In one form of l. andicola [var. gemmifera
(Rauh & Falk) L. D. Gomez] reproduction is commonly by means of apical dichotomy and subsequent fragmentation (Rauh and Falk 1959). Similar foliar gemmae are reported in the South American I. eshbaughii
by Hickey (1986 b). Rury (1978) described adventitious
buds arising de novo from cauline parenchyma, a useful feature by which the plant could rejuvenate after
aestivation in seasonally dry pans. Spore dispersal of
terrestrial species such as I. histrix is more problematic,
and soil erosion and the foraging of herbivores must
play an important part. Frost et al. (1982) estimated a
population of 69,000 plants in some 500 m2 of highly
grazed rocky turf in West Cornwall, England, suggesting the hooves of grazing animals may help to spread
the spores across short distances.
Along with other aquatic plants, submerged Isoetes
has been reported (Keeley 1982) to use a carbon assimilation pathway of the type normally associated with
desert plants (Crassulacean Acid Metabolism), although some terrestrial species lack this ability. Keeley
et al. (1984) found that the thick cuticle of I. andicola
was essentially impervious to CO2 and H20 vapour
and showed, by isotope labelling, that CO 2 uptake was
entirely through the roots, which were well entrenched
in highly decomposed and therefore CO,-rich peat.
These two metabolic features may well give Isoetes an
ecological advantage over other associated plants.
PALAEONTOLOGY. Isoetes-like plants, attributable to
the family, have been recorded in the Triassic and later
deposits (e. g. as Isoetites); many of the latter are described only from spores. The family shows affinities
with Chaloneriaceae of the Upper Carboniferous
Isoetes
(which apparently also possessed endosporous gametophytes) and with Nathorstiana and other P1euromeiaceae of the Mesozoic, both of which are placed in the
Isoetales by Pigg and Rothwell (1983), suggesting that
direct lsoetaceae ancestors are thus contemporaneous
with Lepidodendrales and not necessarily derived
from them.
A single genus:
lsoetes L.
Figs. 6, 7
[soetes L, Sp. PI. 2: 1100 (1753); Pfeiffer (1922); Reed (1953);
Croft (1980); Panigrahi (1981); Boom (1982); Johnson (1984);
Jermy and Schelpe (1986).
Stylites E. Amstutz, Ann. Missouri Bot.· Gard.44: 121 (1957);
Rauh and Falk (1959 a, b).
Characters of the family.
29
The following subgenera are recognized here following Gomez (1980) but a final definition of the subgenus Stylites must await a review of the South American species (R.J. Hickey, in prep.).
Subgenus Isoetes. Stem usually corm-like, rarely
branched, exhibiting bipolar growth with usually dichotomously branched roots arising in the basipetal
notch, but one species (I. tegetiformans Rury) has a
slender, short-creeping, rhizome-like stem. Leaves terete to more or less flattened, often flaccid and slender
in submerged aquatic species, with or without stomata,
cuticle usually thin Or absent. Sporangia at least partially immersed in the base of the sporophyll.
Subgenus Stylites (Amstutz) L. D. Gomez, Brenesia 18: 4 (1980). Stem more massive, elongate, erect,
often dichotomously branched, elongate, without a basipetal apical notch but with unbranched roots arising
from a single lateral groove running the length of the
stem. Leaves linear, flat, rigid, without stomata, cuticle
'. thick. Sporangia more or less superficial, often several
centimetres above the base of the sporophyll.
Fig. 6 A-C. Isoetaceae. A [soetes tegetiformans. habit (x 5).
B, C [soetes andicola. B Megasporophyll. C Trophophyll
(L~ligule; Sp~sporangium) (A Rury 1978; B, C Rauh and
Falk 1959)
30
Lycopodiatae . Isoetaceae
Fig.7. Isoetaceae. lsoeles andina; Colombia. Phot. J. Aenly
Selected Bibliography
Abraham, A., Ninan, C. A. 1958. Cytology of lsoeles. Curro
Sci. 27: 60-61.
Amstutz, E. 1957. Styliles, a new genus of Isoetaceae. Ann.
Missouri Bot. Gard, 44: 121-123.
Berthe~ P, Lecocq, M. 1977. Morphologie sporate des especes fran~aises du genre ISoeles L Pollen Spores 19:
329-359.
Boom; B. M. 1982. Synopsis of lsoeles in the southeastern
United States. Castanea 47: 38-59.
Croft, J. R. 1980. A taxonomic revision of lsoeles L (Isoetaceae) in Papuasia. Blumea 26: 177-190.
Depoux, M., Pitot A. 1965. Etude anatomique de la region ligulaire de Ia feuille fertile chez lswles melanolheca Alston
et lsoetes pitolii Alston. Bull. Inst. Fr. Afr. Noire 27 Ser. A
(2): 514-550.
Frost, L c., Hughes M. G. B., Nichols C., Lawman J. M.
1982. Total population estimate of the land quillwort
(lsoetes histrix) at the Lizard district and recommendations for its conservation. Univ. Bristol Lizard Proj. Rep. 4:
1-23.
Fuchs, H. P. 1962. Nomenldatur, Taxonomie und Systematik
der Gattung lsoetes Linnaeus in geschichtlicher Betrachtung. Beih. Nova Hedw.3: 1-103.
Fuchs, H. P. 1982. Zur heutigen Kenntnis von Vorkommen
und Verbreitung der siidamerikanischen lsoeles-Arten.
Proc. Koninld. Ned. Akad. Wetensch. C 85: 205-260.
Gibby, M. 1990. New chromosome numbers in lsoetes. Fern
Gaz. (in press).
G6mez-P., L D. 1980. Vegetative reproduction in a Central
American Isoeles(Isoetaceae): its morphological, systematic and taxonomical significance. Brenesia 18: 1-14.
Goswami, H. K. 1974. Chromosome studies in natural populations of lsoetes pantii, with heterosporous sporangia. Cytologia 40: 543-551.
Goswami, H. K., Arya B. S. 1968. Heterosporous sporangia in
lsoetes. Brit. Fern Gaz. 10: 39-40.
Hall, J. B. 1971. Observations on lsoeles in Ghana. Bot. J.
Linn. Soc. 64: 117-139.
Hickey, R. J. 1984. Chromosome numbers of neotropical lsoeles. Amer. Fern J. 74: 9-13.
Hickey, R.J. 1986a. On the identity of lsoetes triquelra
A. Braun. Taxon 35: 243-246.
Hickey, R. J. 1986 b. The early evolutionary and morphological diversity of lsoetes, with descriptions of two new neotropical species. Syst. Bot. 11: 304-321.
Hickey, R.J. 1986 C. lsoetes megaspore surface morphology:
nomenclature, variation and systematic importance. Amer.
Fern J. 76: 1-16.
Jermy, A. C., Schelpe E. A. C. L. E. 1986. Isoetaceae. In:
Schelpe E. A. C. L E. (Ed.) The Pteridophyta of southern
Africa. Cape Town: Botanical Research Institute, Dept. of
Agriculture and Water Supply, Rep. of South Africa.
Johnson, E. R. L 1984. Taxonomic revision of lsoetes L in
western Australia. J. R. Soc. W. Ausl. 66: 28-43.
Karrfalt, E. E., Hunter D. M. 1980. Notes on the natural history of Stylites gemmifera. Amer. Fern J. 70: 69-72.
Keeley, J. E. 1982. Distribution of diurnal acid metabolism in
the genus lsoeles. Amer. J. Bot 69: 254-257.
Lycopodiatae • Lycopodiaceae
Keeley, J. E., Osmond C. B., Raven J. A. 1984.. Stylites, a vascular land plant without stomata absorbs carbon dioxide
via its roots. Nature (London) 310 (5979): 694-695.
Kott, L S., Britton D. M. 1980. Chromosome numbers for Isoetes in northeastern North America. Can. J. Bot. 58:
980-984.
Kott, L S., Britton D. M. 1982 a. Comparison of chromatographic spot patterns of some North American Isoetes species. Amer. Pern J. 72: 15-18.
Kott, L S., Britton D. M. 1982 b. A comparative study of
spore germination of some Isoetes species of northeastern
North America. Can. J. Bot. 60: 1679-1687.
Kubitzki, K., Borchert R. 1964. Morphologische Studien an
Isoetes triquetra A. Br. und Bemerkungen fiber das Verhaltnis der Gattung Sty/iles E. Amstutz zur Gattung Isoeles L
Ber. DIsch. Bot. Ges. 77: 227-234.
Marsden, C. R. 1976 a. A new subspecies of Isoetes coromonde/ina from northern Australia. Contrib. Herb.
Aust.24: 1-10.
Marsden, C. R. 1976 b. Morphological variation and taxonomy of Isoetes muelleri A. Br. J. Adelaide Bot. Gard.l (19):
37-54.
Maserati, D. L, Thomas B. A. 1982. The stomata of Isoetes.
Phyta, Stud. Living Possil Plants, Pant Commun. Vol. 1982:
155-162.
Michaux, N. 1973. La ligule chez l'Isoiiles setacea Lam. C. R.
Hebd. seances Acad. Sci. Ser. D 277: 41-44.
Ninan, C. A. 1958. Studies on the cytology and phylogeny of
the pteridophytes. V. Observations on the Isoetaceae. J. Indian Bot. Soc. 37: 93-102.
Panigrahi, G. 1981. Systematics of the genus Isoetes L (Isoetaceae) in India. BioI. Mem.6 (2): 129-138.
Pant, D. D., Srivastava, G. K. 1965. Cytology and reproduction of some Indian species of Isoetes. Cytologia 30:
239-251.
Paolillo, D. J. 1963. The developmental anatomy of Isoetes.
III. BioI. Monogr. 31: 1-130.
pfeiffer, N. E. 1922. Monograph of the Isoetaceae. Ann. Missouri Bot. Gard. 9: 79-232.
Pigg, K. 8., Rothwell G. W. 1983. Megagametophyte development in the Chaloneriaceae ram. nov., permineralised Paleozoic Isoetales (Lycopsida). Bot. Gaz. 144: 295-302.
Rauh, W., Falk H. 1959. Stylites E. Amstutz, eine neue Isoetacee aus den Hochanden Perus 1. Sitzungber. Heidelberger
Akad. Wiss. Math. Naturwiss. Kl. 1959: 3-160.
Reed, C. P. 1953. Index Isoetales. Bol. Soc. Brot 11.27: 5- 72.
Robert, D., Roland-Heydacker F., Denizot J., Laroche J.,
Pougeroux P., Davignon L 1973. Etude de la paroi siliceuse chez la megaspore d'Isoetes setaceo Delile. C. R.
Acad. Sci. Paris ser. D 276: 2521-2524.
Rury, P. M. 1978. A new and unique mat-forming Merlin'sgrass (Isoeles) from Georgia. Amer. Pern J. 68: 99-108.
Sharma, U. 1958. Contributions to our knowledge of Isoeles
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'n,ylor, W. c., Luebke N. T., Smith M. 8. 1985. Speciation and
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Edinburgh 86 B: 259-263.
Voirin, B., Jay M. 1978. Etude chimosytematique des Lycopodiales, Isootales, Selaginellales et Psilotales. Biochem. Syst.
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Wanntorp, H.-E. 1970. The genus Isoetes in South West Africa. Svensk Bot. Tidskr.64: 141-157.
Weber, U. 1922. Zur Anatomie und Systematik der Gattung
Isoetes L Hedwigia 63: 219-262.
31
Lycopodiaceae
B.0LLGAARD
Lycopodiaceae Mirbel in Lamarck & Mirbel, Hist. Nat. Veg.
4: 293 (1802); 011gaard (1989).
Terrestrial or epiphytic, erect to pendulous herbs or
climbers. Stems dichotomously branched, rarely with
lateral branching, protostelic, with the xylem arranged
radially or in parallel bands, or forming an almost
closed cylinder (Phylloglossum). Leaves simple, with
one simple vein, arranged in low alternating spirals or
irregular whorls, or decussate, homophyllous Or heterophyllous, isophyllous Or anisophyllous. Sporophylls
like the foliage leaves or modified, sometimes specialized and aggregated into distinct strobili. Sporangia
solitary, in the leafaxils or on the upper side of the
sporophyll base, homosporous, unilocular, reniform to
subglobular, short-stalked, dehiscing by a transverse
slit, dividing the sporangium into two valves. Spores
without chlorophyll, subglobose to tetrahedral, with a
trilete scar. Gametophytes monoecious, tuberous, subterranean and holomycotrophic or surface-living,
green and hemimycotrophic.
MORPHOLOGY AND ANATOMY. Branching patterns
were studied by 0llgaard (1979a), who found three basic patterns corresponding to three genera (as SUbgenera of Lycopodium s.lat.). The morphogenesis of phyllotaxis seems irregularly organized. Stevenson (1976)
interpreted the phyllotaxis of Huperzia lucidula
(Michx.) Trev. as consisting of low, alternating spirals
and found a definite relation between the number of
orthostichies and the number of protoxylem lobes in
the cauline stele. In many species this relation is different, and transitions from spiral to irregularly whorled
phyllotaxis exist. Distinct types of anisophylly evolved
independently in several sections of Lycopodium and
Lycopodiel/a. The cortex is thick and usually not sclerified in Huperzia. usually thin and sclerified in Lycopodium and Lycopodiel/a. The development of cortical
roots (Stokey 1907) is unique to Huperzia. The xylem
of the root steles is crescent-shaped in cross-section in
Huperzia. Phylloglossum. and Lycopodiella sect. Lycopodiella and sect. Carolinianae. In the other groups the
stele of the main root is similar to the stem stele. In'Lycopodium the xylem of the main stem is arranged in
rather fixed, parallel bands, gradually changing to a
radial arrangement in the branchlets. In Lycopodiella
and Huperzia the stele is radial in structure, usually
with a variable, ± anastomosing course of the xylem
lobes. In Lycopodiella sect. Campylostachys the xylem
is highly dissected. In Phylloglossum the stele is highly
reduced and variable; it was interpreted as a mesarch,
medullated protostele with rudimentary phloem; see
32
Lycopodiatae . Lycopodiaceae
Hackney (1950). Bruce (1976a) found the distribution
of mucilage canals in leaves, sporophylls and cortex
strongly taxonomically correlated.
The type of sporophyll differentiation is taxonomically significant. In Huperzia the sporangiate leaves
are paleate, i. e. basally attached without basiscopic
laminar appendages, and continue to assimilate after
dehiscence. In the other genera the sporophylls are
ephemeral and subpeltate or peltate; only Lycopodium
sect. Pseudodlphasium and sect. Pseudolycopodium
have paleate but ephemeral sporophylls; see 011gaard
(1987). The mature sporangia are axillary in Huperzia,
Lycopodiella sect. Lycopodiella, and Lycopodium sect.
Pseudodiphasium. In the remaining groups they are
borne on the sporophyll base. The sporangia are deeply sunken in the contiguous strobilus cortex and coalescent sporophyll bases in Lycopodiella sect. Campylostachys and sect. Lateristachys. The relative size of
the two sporangium valves, and the shape and thickenings of the sporangium epidermis cells, are strongly
taxonomically correlated (0llgaard 1975; Fig. 12).
SPORES. The external spore morphology was studied
by Wilce (1972) who found five distinct spore types,
strongly correlated with taxonomy. The spores of Phylloglossum are very similar to those found in Huperzia
(Breckon and Falk 1974).
GAMETOPHVTES. Although known in rather few species
[detailed studies by Bruchmann (1898) and Holloway
(1916, 1919, 1920)], gametophyte morphology seems
taxonomically significant (Boivin 1950; Bruce 1976b;
Fig. 8). Experimental cultivation led Freeberg and
Wetmore (1957) to the conclusion that the distinctive
morphology of the gametophytes in the wild is determined by the fungus symbiont. However, gametophytes grown without the fungus by Wbittier (1977,
1981) maintained the morphology of wild ones, and
Bruce (1976b) showed that Freeberg and Wetmore's
conclusion was due to experimental error. Gametophytes are rarely observed, most often in disturbed
habitats with scanty pioneer vegetation. In Huperzia
and Lycopodium the spores have highly resistant spore
walls and may germinate only after a long time, developing into slowly maturing, mycotrophic, subterranean gametophytes. In Lycopodiella the spores may
germinate without delay, developing into surface-living, green gametophytes. Outbreeding seems as likely
in the subterranean as in the surface-living gametophytes (Wagner et al. 1985; 011gaard 1985).
KARVOLOGV AND HVBRIDIZATION. Chromosome
studies of the Lycopodiaceae have been hampered by
high numbers and by the peculiar behaviour of meiotic chromosomes. In many species studied there are only approximate counts, and many others need confirmation. In Huperzia two distinct base numbers were
indicated by Chiarugi (1960) and Love et al. (1977) for
groups recoguized as Huperzia (x= 11) and PhlegmariuTUS (x= 17). Later counts by Beitel and Wagner
(1982) and Takamiya and Kurita (1983) do not support
this subdivision. In Lycopodiella distinct numbers are
found in sect. Lycopodiella(n=78) and sect. Carolinianae (n=34, 35, c. 69), whilst the series n=104, 108,
110, 136, c. 165 in sect. Campylostachys eludes the simple deduction of a base number. In Lycopodium there
are distinct counts for sect. Complanata (n=23), sect.
Diphasium (n=c. 90,34-36), sect. Magellanica (n=31),
see 0llgaard (1987). The identical number of n= 34 in
Lycopodium sect. Lycopodium, sect. Annotina, sect. Obscura and sect. Lycopodiastrum may reflect the antiquity of this number, rather than a close affinity of the
groups.
Several species hybrids are known in Lycopodium
sect. Complanata, Lycopodiella sect. Lycopodiella, and
in Huperzia (Beitel 1979; 011gaard 1985; Wagner et
al. 1985). In several cases hybrid meioses in Lycopodium sect. Complanata are reported to be normal (Wagner 1980; Hersey and Britton 1981) and cannot be
used to verify hybridization, while Huperzia hybrids
were reported to have irregular meioses.
VEGETATIVE REPRODUCTION. The bulbils of Huperzia
lucidula were studied by Stevenson (1976) who also reviewed earlier literature on bulbils in the group of
H. selago. Bulbils have been interpreted as homologous with leaves, topographically homologous with
leaves, or arrested, extremely unequal stem dichotomies, the latter interpretation being the most convincing (Fig. 9 B).
Adventitious buds often regenerate ageing or damaged plants of Huperzia from the stem base (see
Chowdhury 1937). Holloway (1917) surveyed the
means of vegetative propagation in the New Zealand
species of Lycopodiaceae.
ECOLOGV AND DISTRIBUTION. Wbile Phylloglossum has
a limited Australian distribution, the three other genera are almost cosmopolitan, being absent only from
arid areas. The greatest species concentration is in humid, tropical, montane forests and in humid, tropical,
alpine vegetation. A few species are very widely distributed, e. g. Lycopodiella cernua (L.) Pichi-Sermolli
(tropics and sUbtropics of all continents), Lycopodium
clavatum L. and Lycopodiella caroliniana (L.) PichiSermolli (both absent from Australia). Many species
establish themselves in disturbed sites and persist
mainly in habitats with low competition (Bruchmann
1898; 0llgaard 1979 b).
AFFINITV AND SUBDIVISION. The Lycopodiaceae are
without close affinity to other groups. The relationship
to other Lycopsida is very remote. Fossils of Lycopodium sensu lato and Lycopodites are generally poorly
Huperzill • Phylloglossum
preserved and cannot be referred with certainty to extant groups of the family.
The subdivision of the Lycopodiaceae has been a
matter of considerable disagreement. The spore study
by Wilce (1972) seems to have settled the delimitation
of taxa in the family, although the opinions of taxonomic rank of the taxa differ (see Holub 1964, 1975,
1983; Bruce 1976a, b, c; Pichi Sermolli 1977). The separation of Huperziaceae (Rothmaler 1944, 1962, as
Urostachyaceae) was followed by some authors and
reflects the isolated position of Huperzia. The four
genera are very distinct, and also the sections within
Lycopodiella and Lycopodium seem to represent ancient, independent evolutionary lines. An index and a
systematic list ofthe species have been given by Herter
(1949, 1949/50) and by 0llgaard (1989). Nessel's
"monograph" (1939) is of limited use. 0llgaard (1987)
revised the classification of the family.
KEY TO THE GENERA.
1. Tiny plants with a subterranean tuber. a small cluster of
leaves, and a compact strobilus on a leafless peduncle
2. Phylloglossum
- Small to large plants with leafy stems; strobili, if present,
sessile or borne on leafy peduncles
2
2. Stems isotomously branched throughout, without elongate,
indeterminate main stems; roots usually forming one basal
tuft; sporophylls and vegetative leaves alike. or the sporophylls, if smaller, paleate. persistent
1. Hupenu.
- Stems anisotomously branched almost throughout, the
branches differentiated into elongate. indeterminate main
stems and determinate branchlet systems; roots emerging
with intervals on the lower side of the main stems; sporophylls strongly modified, peltate. subpeltate. or rarely paleate, ephemeral
3
3. Strobili pendent and sessile; or erect and terminating simple (rarely forked) branches which arise dorsally on the
creeping stem; or erect and seemingly lateral on overtopping vegetative shoots
4. Lyeopodb!Ua
- Strobili erect, sessile or pedunculate, borne on branchlet
systems which arise laterally on the main stem; or pendent
3. Lycopodium
and pedunculate.
I. Huperzia Bernhardi
33
mucilage cavities, persistent; sporangia axillary, reniform, isovalvate, with a slender stalk, the epidermal
ceIl walls sinuate, thickened and lignified except on
the outer surface; spores foveolate-fossulate; gametophytes subterranean, holomycotrophic, cylindrical
with radial or bilateral symmetry, with pluriceIlular
uniseriate hairs among the gametangia (Fig. 8).
Herter (1949/1950 and later additions) recognized
more than 400 species; Tryon and Tryon (1982) estimated 200 species; perhaps c. 300 species are distinguishable; species distinction is problematic in many
groups due to the plasticity of the characters. A nearly
cosmopolitan genus, in tropical, temperate, arctic and
alpine environments. Species diversity is highest in
tropical evergreen montane forests and in the Andean
paramos and subparamos of South America. The genus includes tiny moss-like species, massive, rigid erect
ones and long, lax, pendent epiphytes, and an almost
continuous gradation between such forms (Figs. 9, 10).
Classifications up till now (e. g. Pritzel 1901; Herter
1949/50) based on the mode of leaf transformation
and the growth habit are artificial. Recently the genus
Phlegmariurus Holub (1964) was recognized as distinct
from Huperzia, but without a study of all species.
Chromosome counts were used to argue for their distinctness, but new evidence (Takamiya and Kurita
1983) indicates their similarity.
The only reasonably distinct infra generic entity is
the group of bulbiferous species including Huperzia
selago (L.) Schrank & Mart:! and H. serrata (Thunb.)
Trev. The spores of this group seem to be distinct, with
deep proximal pitting, concave proximal faces, and
truncate corners vs smooth or shaIlowly pitted, usuaIly
flat proximal faces and evenly rounded to angular
corners in the remaining species. The latter. although
their growth habit and morphology are highly diverse,
exhibit almost continuous variation. A sound classification of these is not evident. A synopsis of the terrestrial American species has been given by Rolleri
(1981).
Figs. 8-10, 12 A
lIuperzia Bemhardi, Schrad. J. Bot. 1800 (2): 126 (1801).
I"ycopodium subgen. Selago Baker (1~87).
I.ycopodium subgen. Urostachya Pritzel (1901).
Urostachys Herter (1922).
1'h1egmariurus Holub (1964).
Sporophytes epiphytic or terrestrial, pendent, erect, or
a~cending, isotomously branched throughout, the
hranches all similar, or in some terrestial species slightly differentiated into prostrate vs erect aerial branches.
Roots arising from the stem stele, descending through
the cortex to the stem base, here emerging as one basal
tuft; homophyllous Or heterophyllous; sporophylls
and vegetative leaves alike, or the sporophylls gradually or abruptly smaller than the foliage leaves, without
2. Phylloglossum Kunze
Fig.9D
Phyllog/ossum Kunze. Bot. Z. 1: 721-23 (1843).
Tiny terrestrial plants, up to c. 5 cm talI, consisting of a
subterranean tuber producing a short stem with a cluster of filiform leaves, one to three adventitious roots, a
compact strobilus borne on a leafless peduncle, and a
leafless geotropic branch which develops into a new
marsupial tuber with the apex deeply immersed and
pointed back towards the parent plant. Sporophylls
peltate, without a basal mucilage cavity. Sporangia axillary to basal on the sporophylls, reniform, isovalvate,
with sinuate, thickened and lignified side walls in the
34
Lycopodiatae . Lycopodiaceae
;~,
~>
Fig. 8 A- D. Lycopodiaceae. A Huperzia lucidula, gameto·
phytes with attached sporelings (x 4.2). B Lycopodium digita·
tum, gametophytes, one with attached sporeling (x 8.5).
C Lycopodium clavatum, gametophyte with attached base and
roots of sporeling (x 5). D Lycopodiella cemua, gameto·
phytes, with attached sporeling (ltift) and showing position of
archegonial necks (right) (c. x 16) (Original 0llgaard)
epidermal cells. Spores foveolate·fossulate, with un·
ornamented, plane proximal faces. n=246II+101
(Blackwood 1953). Gametophytes surface-living,
green.
One species, Ph. drnmmondii Kunze (Fig. 9 D), in
Australia, Tasmania, and New Zealand. A highly reduced and specialized species, adapted to seasonally
wet growing conditions. Only the subterranean tubers
survive the dry season. The affinities of the genus are
not clear. Bower'S interpretation (1885) of Phylloglossum as a permanently embryonic (neotenic) form, derived from Lycopodiella cernua, is unlikely, considering
the highly specialized marsupial tubers; see review by
Hackney (1950). The spore type (Breckon and Falk
1974) and sporagium wall cells (011gaard 1975) are
similar to those of Huperzia, whereas the gametophyte
and protocorm resemble Lycopodiella.
Huperzia· Phylloglossum
Fig. 9 A-D. Lycopodiaceae. A Huperzia crassa, habit (x 0.7)
and details of sporophyl\s with large basal air cavities
( x 4.4). B Huperzia selago, part of stem bearing sporangia
and two bulbiferous branchlets, one bulbil detached (x 8).
C Huperzia phylid!olia, habit (x 0.7) and details (x 4) showing basal expanded leaves and fertile division. D Phylloglossum drummondii, habit, with new marsupial tuber (c. x 3)
(OriginaI0I\gaard)
36
Lycopodiatae • Lycopodiaceae
Fig. 10. Lycopodiaceae. Huperzia brevi/olia; Ecuador. Photo
B.011gaard
3. Lycopodium L.
Fig.11 B
Lycopodium L, Sp. PI.: 1100 (1753); Wilee (1965); Hickey
(1977).
Lepidotis Mirbel (1802).
Lycopodium subgen. Lepidotis Baker (1887).
Lycopodium subgen. Rhopalostachya Pritzel (1901).
Diphasium C. Presl ex Rothm. (1944).
Diphasiastrum Holub (1975).
Lycopodiastrum Holub (1983).
Pseudodiphasium Holub (1983).
Pseudolycopodium Holub (1983).
Sporophytes terrestrial, anisotomously branched, with
elongate, indeterminate, subterranean, creeping or
climbing main stems, and determinate lateral, ascending to erect or spreading branchlet systems arising at
inclined angles. Stele of main stems and major branchlets with xylem in parallel bands; in minor branchlets,
peduncles and strobili usually with radially arranged
xylem. Roots emerging directly along the underside of
main stems, with parallel-banded stele in main roots.
Leaves uniform throughout, or branchlets anisophyllous. Strobili erect or pendent, simple or forked, either
sessile and terminating the stronger branchlets or
borne on simple or forked peduncles. Sporophylls
ephemeral, subpeltate with a thin basal decurrent
wing, with or without a basal mucilage cavity, or peltate or paleate. Sporangia on the sporophyll base
(rarely axillary), reniform, with a short, thick stalk, isovalvate to slightly anisovalvate, their epidermal cells
with thin, lignified, sinuate anticlinal walls without
partial thickenings. Spores reticulate, baculate, or scabrate. Gametophytes carrot-shaped or obconic to convoluted disc-shaped, subterranean, holomycotrophic,
without pluricellular uniseriate hairs among the gametangia.
A genus of perhaps 40 species; Ching (1981-82)
added several new Chinese species. Most species are
in north- and south-temperate regions and on tropical
mountains. Growth habits are diverse, including low
matted forms, species with tree-like growth, and high
climbers. There are nine very natural sections (011gaard 1987), several of which were treated as genera
by Holub (1975, 1983). The sections are: Sect. Lycopodium; type species: L. clavatum L. (Fig. 11 B). - Sect.
Annotina (Rouy) Holub; type species: L. annotinum L.
- Sect. Complanata Victorin (Syn.: Diphasiastrum Holub, 1975); type species: L. complanatum L. - Sect. Diphasium (Rothm.) B. 011gaard (Syn.: Diphasium
Lycopodium. Lycopodiella
C. Presl ex Rothm., 1944); type species: L.jussiaei PoiI·et. - Sect. Lycopodiastrum (Holub) B.0IIgaard (Syn.
I"ycopodiastrum Holub 1983); type species: L. casuarilIoides Spring. - Sect. Magellallica B. 0IIgaard; type
species: L. magellanicum (Pal. Beauv.) Sw. - Sect. Ob.feura (Rothm.) Holub; type species: L. obscurum L. Sect. Pseudodiphasium (Holub) B.0IIgaard (Syn.:
Pseudodiphasium Holub, 1983); type species: L. voluhile Forst. - Sect. Pseudolycopodium (Holub) B. 0l1gaard (Syn.: Pseudolycopodium Holub 1983); type species: L. deuterodensum Herter.
These sections are based on characters of growth
habit; leaf differentiation (isophyIIy or type of anisophyIIy); the presence or absence of peduncles; peltate,
subpeltate, or paleate sporophyIIs; structure of sporangium epidermis cell walls; spore ornamentation; shape
of gametophyte; and chromosome number.
4. Lycopodiella Holub
Figs. 11 A, 12B
I,vcopodie//a Holub, Preslia 36: 22 (1964).
I,vcopodium subgen. Cemuistachys Herter sensu Tryon and
Tryon (1982); Lepidotis auct. non Mirbel (1802).
I..vcopodium subgen. Lepidotis auct. non Baker (1887).
I'ulhinhaea Vasco and Franco (1967).
I~'eudolycopodiella
Holub (1983).
I.uleristachys Holub (1983).
37
Fig..tt A, B. Lycopodiaceae., A Lycopodiella alopecuroides.
habit (x 0.7), enlarged vegetative leaves and sporophyll (left).
B LycopodIum c1avatum ssp. contiguum. habit (x 0.7), details
of spOrophylls (x 8) (Original 0l1gaard)
Plants terrestrial, anisotomously branched, with horizontal, creeping, or arching-looping indeterminate
shoots rooting at short to long intervals along the underside, and dorsally arising, determinate, erect, simple or profusely tree-like branched aerial shoots; or
with irregularly, truly laterally branched, subterranean
stems (sect. Lateristachys). Stem steles radial, sometimes with highly dissected xylem. Leaves isophyllous
to anisophyllous. Strobili pendent and sessile, or erect
and terminating simple (rarely forked), dorsally arising
branches. Sporophylls subpeltate, with a thin basal decurrent wing, or with coalescent basal membranes
which almost enclose the sporangia, with a mucilage
cavity in the base and sometimes along the vein. Sporangia reniform to sub globose, anisovalvate or isovalvate, on the sporophyll base, their epidermal cells with
thin, straight, non-lignified walls except nodular or
semiannular, lignified thickenings. Spores rugate.
Gametophytes tuberous and lobed, surface-living,
hemisaprophytic.
38
Lycopodiatae . Lycopodiaceae
B
A
Fig. 12 A, B. Lycopodiaceae. Sporangial wall (x 5(0). A Huperzia funiformis. B Lycopodie/la a/opecuroides. Phot.
B.0llgaard
A genus of perhaps 40 species in almost all moisttemperate and tropical regions. South America has the
highest species diversity. Growth habit is diverse and
in part reflects infrageneric taxonomy. There are four
very natural sections (0llgaard 1987) which were treated as genera by Vasconcellos and Franco (1967) and
Holub (1964, 1983). The sections are: Sect. Lycopodiella (syn.: Lycopodiella Holub, 1964; Lycopodium sect.
Inundata Pritzel, 1901); type species: L. inundata (L.)
Holub. - Sect. Campylostachys (K. Muller) B. 0llgaard
(Syn.: Palhinhaea Vasco & Franco 1967); type species:
L. cemua (L.) Pichi-Sermolli. - Sect. Carolinianae
(Bruce) B. 011gaard (Syn.: Lycopodium sect. Carolinianae Bruce; Pseudolycopodiella Holub, 1983); type species: L. caroliniana (L.) Pichi SermolJi. - Sect. Lateristachys (Holub) B.0llgaard (syn.: Lateristachys
Holub, 1983); type species: L.lateralis (R. Br.) B.0llgaard.
Selected Bibliography
Baker, J. G. 1887. Handbook of the fern-allies. London:
George Bell & Sons.
Beitel, J. M. 1979. Clubmosses (Lycopodium) in North America. Bull. Amer. Fern Soc. 6 (5): 1-8.
Beitel, J. M., Wagner, F. S. 1982. The chromosomes of Lycopodium lucidulum. Amer. Fern J. 72: 33-35.
Blackwood, M. 1953. Chromosomes of Phylloglossum drummondii, Kunze. Nature (London) 172: 591-592.
Boivin, B. 1950. The problem of generic segregates in the
form-genus Lycopodium. Amer. Fern J. 40: 32-41.
Bower, F. O. 1885. On the development and morphology of
Phylloglossum drummond;;. Philos. Trans. R Soc. London
Ser B 176(II): 665-678.
Breckon, G. J., Falk, R. H. 1974. External morphology and
taxonomic affinities of Phylloglossum drummondii Kunze
(Lycopodiaceae). Amer. J. Bot. 61: 481-485.
Bruce, J.G. 1976a. DevelOpment and distnbution of mucilage canals in Lycopodium. Amer. J. Bot. 63: 481-491.
Bruce, J. G. 1976b. Gametophytes and subgeneric concepts in
Lycopodium. Amer. J. Bot. 63: 919-924.
Bruce, J.G.1976c. Comparative studies in the biology of Lycopodium carolinianum. Amer. Fern J.66: 125-137.
Bruchmann, H. 1898. Oller die Prothallien und Keimpl1anzen
mehrerer europiiischer Lycopodien. Gotha.
Chiarugi, A. 1960. Tavole chromosomiche delle Pteridophyta.
Caryologia 13: 27 -150.
Ching, R C. 1981/82. The taxonomy of Chinese Lycopodiaceae (sens. lat.). I-IV. Acta Bot Yunnan. 3: 1-9, 291-305;
4: 119-128, 213-226.
Chowdhury, N. P. 1937. Notes on some Indian species of Lycopodium, etc. Trans. Nat!. Inst. Sci. India 1: 187-226.
Freeberg, J. A., Wetmore, R. H. 1957. Gametophytes of Lycopodium as grown in vitro. Phytomorphology 7: 204-217.
Hackney, F. M. V. 1950. A review of and contnbution to the
knowledge of Phylloglossum drummondii Kunze. Proc. Linn.
Soc. New S. Wales 75: 133-152.
Hersey, R E., Britton, D. M. 1981. A cytological study of
three species and a hybrid taxon of Lycopodium (section
Complanata) in Ontario. Can. J. Genet. Cytology 23:
497-504.
Herter, G. 1949. Index Lycopodiorum. Est. Bot. Reg. Uruguaya 20: I-IV, 1-120.
Herter, G. 1949150. Systema Lycopodiorum. Rev. Sudamer.
Bot. 8: 67-86, 93-116.
Hickey, R J. 1977. The Lycopodium obscurum complex in
North America. Amer. Fern J. 67: 45-48.
Holloway, J. E. 1909. A comparative study of the anatomy of
six New Zealand species of Lycopodium. Trans. New Zeal.
Inst. 42: 356 0 370.
Holloway, J. E. 1916. Studies in the New Zealand species of
Lycopodium l. Trans. New Zeal. Inst. 48: 253-303.
Selected Bibliography
Holloway, J. E. 1917. Studies in the New Zealand species of
Lycopodium II. Trans. New Zeal. Inst. 49: 80-93.
Holloway, J. E. 1919. Studies in the New Zealand species of
Lycopodium III. Trans. New Zeal. Inst. 51: 161-171.
Holloway, J. E. 1920. Studies in the New Zealand species of
Lycopodium IV. Trans. New Zeal. Inst. 52: 193-239.
Holub, J. 1964. Lycopodiella, novY rod fildu Lycopodiales.
Preslia 36: 16-22.
Holub, J. 1975. Diphasiastrum, a new genus in Lycopodiaceae. Preslia 47: 97-110.
Holub, J. 1983. Validation of generic names in Lycopodiaceae: with a description of a new genus PseudolycopodieUa.
Folia Geobot. Phytotax. 18: 439-442.
Uive, A., Love, D., Pichi Sermolli R. E. G. 1977. Cytotaxonomical atlas of the Preridophyta. Vaduz: Cramer.
Nessel, H. 1939. Die Biirlappgewiichse. Jena: Gustav Fischer.
011gaard, B. 1975. Studies in Lycopodiaceae I. Observations
on the structure of the sporangium wall. Amer. Fern J. 65:
19-27.
011gaard, B. 1979a. Studies in Lycopodiaceae II. The branching patterns and infrageneric groups of Lycopodium sensu
lato. Amer. Fern J. 69: 49-61.
011gaard, B. 1979b. Lycopodium in Ecuador - habits and habitats. In: Larsen, K., Holm-Nielsen, L. B. (Eds) Tropical
botany. London: Academic Press.
011gaard, B. 1983. Lycopodiaceae. In: Stolze, R. G. Ferns and
fern-allies of Guatemala. Fieldiana Bot N. S. 12: 20-44.
011gaard, B. 1985. Observations on the ecology of hybridisation in the clubmosses (Lycopodiaceae). Proc. R. Soc.
Edinburgh 86B: 245-251.
011gaard, B. 1987. A revised classification of the Lycopodiaceae s. lat. Opera Bot. 92: 153-178.
011gaard, B. 1989. Index of the Lycopodiaceae. Biolog. Skr.
34, Kongel. Danske Vidensk. Selsk., Copenhagen: 1-135.
I>ichi Sermolli, R. E. G. 1977. Tentamen pteridophytorum in
taxonomicum ordinem redigendi. Webbia 31: 313-512.
"ritzel, E. 1901. Lycopodiaceae. In: A. Engler und K. Prantl
(Eds) Die natOrlichen PIIanzenfamilien I 4, Leipzig:
W. Engelmann, pp. 563-606.
Rolleri, C. H. 1981. Sinopsis de las especies de Lycopodium L.
(Lycopodiaceae, Pteridophyta) de la secci6n Crassistachys
Herter. Rev. Mus. La Plata N.S. Bot. 13: 61-114.
Rothmaler, W. 1944. Pteridophyten-Studien I. Feddes Rep.
54: 55-82.
Rothmaler, W. 1962. Dber einige Diphasium-Arten (Lycopodiaceae). Feddes Rep. 66: 234-236.
Stevenson, D. w. 1976. Observations on phyllotaxis, stelar
morphology, the shoot apex and gemmae of Lycopodium
lucidulum Michx. (Lycopodiaceae). Bot. J. Linn. Soc. 72:
81-100.
Stokey, A. G. 1907. The roots of Lycopodium pithyoides. Bot.
Gaz. 44: 57-63.
'1IIkamiya, M., Kurita, S. 1983. Cytotaxonomic studies on
Japanese species of the genus Lycopodium sensu lato. Acta
Phytotax. Geobot. 34: 66-79.
'1lIkamiya, M., Tanaka, R. 1983. Chromosomes of Lycopodi11m casuarinoides, a fern ally. Chromosome Inf. Servo 35:
27-28.
Vusconcellos, I. C., Franco, J. A. 1967. Breves notas sobre Licopodiaceas. Rol. Soc. Brot. II. 41: 23-25.
Wagner, F. S. 1980. Chromosome behavior in three interspecific hybrids of Diphasiastrum (Lycopodiaceae). Bot. Soc.
Amer. Misc. Ser. 158: 121.
Wagner, W. iI., Jr., Wagner, F. S., Beitel, J. M. 1985. Evidence
for interspecific hybridisation in pteridophytes with subter-
39
ranean mycoparasitic gametophytes. Proc. R. Soc. Edinburgh 86B: 273-281.
Walker, T. G. 1966. A cytotaxonomic survey of the pteri·
dophytes of Jamaica. Trans. R. Soc. Edinburgh 66: 169237.
Whittier, D. P. 1977. Gametophytes of Lycopodium obscurum
as grown in axenic culture. Can. J. Bot. 55: 563-567.
Whittier, D. P. 1981. Gametophytes of Lycopodium digitatum, etc., as grown in axenic culture. Bot. Gaz. 142: 519524.
Witce, J. H. 1965. Section Complanara of the genus Lycopodium. Beih. Nova Hedwigia 19: i-ix, 1-233.
Witce, 1. H. 1972. Lycopod spores I. General spore patterns
and the generic segregates of Lycopodium. Amer. Fern J.
62: 65-79.
SelagineUaceae
A.C.JERMY
Selaginellaceae Willk. in Willk. and Lange, Prodr. F1. Hisp. 1:
14 (1861); Reed, Mem. Sc. Brot. 18: 1-287 (1966); Pichi
Serm, Webbia 35: 235 (1982).
Terrestrial, or very occasionally epiphytic, usually
perennial plants, of varied habit, with an erect or
prostrate main stem and subsidiary branch systems of
varying arrangements, the basal portions not differentiated into a distinct rhizome although some erect species spread by means of creeping basal branches
known as sobols, which in turn give rise to further
erect branches; growing points not protected by scales.
Roots (sometimes termed rhizophores) dichotomously
branChed, varying in thickness, arising from the axils
of branches, either throughout the length of the main
stem, or basally (acting as support in erect species), or
occasionally apparently adventitiously. Main stems either far-creeping and often much-branched, of indefinite growth which may be only seasonally halted or
terminated; or short-creeping and then becoming
erect, often with a distinct unbranched ('caulescent')
region below, and with variously arranged frond-like
complanate branch systems (pseudofronds) of finite
growth above, further growth arising only from basal
branches; sometimes exhibiting a scrambling or climbing habit. Leaves spirally arranged, ligulate, those on
basal creeping portions often distantly arranged, either
all similar or of two kinds, at least on the secondary
and ultimate branches where they are arranged in four
ranks. Strobili terminal on primary or ultimate
branches, compact, Or occasionally more spread out
along secondary branches, apex occasionally reverting
to become a vegetative shoot; or, more rarely, in a lateral position on a primary or secondary branch. Spor-
40
Lycopodiatae • Selaginellaceae
ophylls leaf-like, spirally arranged or in ranks of four,
uniform or, more rarely, dimorphic, subtending sporangia which are on the adaxial surface just above the
ligule; sporangia stalked, of two kinds, variously disposed throughout the strobilus, megasporangia assuming a shape dictated by the developing megaspores,
microsporangia globose or slightly wider than long,
thin-walled but with areas of thickened cells which aid
dehiscence; megaspores trilete, usually four (>42
have been observed) in each sporangium, pale buff or
white, with distinct muri Or ridges around the equator
and along the triradiate scar, sporoderm variously patterned, rugose-reticulate, rugose, papillate, tuberculate,
granulate, rarely plain on proximal face, rugose-reticulate patterns predominate generally there is less ornamentation on the proximal face than on the distal face,
200-600 J1IIl in equatorial diam.; microspores trilete,
> 100 in each sporangium, ranging in colour from
yellow, through orange-brown, to red, sporoderm variously patterned from finely echinulate through papillate to verrucate, sometimes strongly rugose, 20-60 J1IIl
in diam.
A family of a single genus, with some 750 species
mainly in the tropical zones of the world, with a few
species reaching the arctic-alpine zones in both hemispheres.
FLORISTIC STUDIES. The following works by
A. G. H. Alston, who made an extensive study of the
genus during his tenure as pteridologist at the British
Museum (Natural History), 1930-1958, give keys to
Selaginella of selected areas: Africa, central (1957),
West (1959), South (1939a), Madagascar (1932);
America, Carribean (1952), North (1955), South-tem. perate (1939b), South-tropical (with Jermy and Rankin
1981); Asia, China (1934a), India (1945), Indo-China
(1951), Malesia (1934b, 1935a, b, 1937, 1940). Other
useful regional accounts include Jermy (1970: E. Africa), Jermy in Parris et aI. (1984: Borneo), Proctor
(1985: Jamaica), Smith (1981: Mexico), Stolze (1983:
Guatemala), Tagawa and Iwatsuki (1979: Thailand),
Wong (1983: Malaysia).
ANATOMY AND MORPHOLOGY. Stems are basically dichotomously branched, with various, often diagnostic,
branching patterns (Jermy 1986; Wong 1983). Vascular
anatomy ranges from a relatively simple protostele (always present in young sporelings) to two steles (which
in prostrate species are flattened in the plane parallel
to the substrate), to more compound plectosteles of
several plate meristeles (which vary from being flat in
cross-section to V-shaped or a closed circle), to T- and
X-shaped 'actinosteles' (Mickel and Hellwig 1969).
Each stele is surrounded by a conspicuous endodermis. Hieronymus (1901) used the vascular complexity
of the stem to delimit taxonomic sections, but Bierhorst (1971) pointed out that variations can Occur in a
single species depending on the position and width of
stems. In many species adventitious shoots arise from
the primary branch axes. The xylem in some species of
subgenera Selaginella and Tetragonostachys is unique
in having vessels of an advanced type which occurs
elsewhere only in flowering plants. One group of predominantly tropical American species in subgen. Stachygynandrum (Series Articulatae Spring) have weaker
areas in the cortex, usually just below the branch,
which appear nodular although the vascular structure
is unaltered. Very few species actually break naturally
at this point and the function is unclear. Somers (pers.
comm.) has studied the anatomy and morphology of
this group and believes it to be a distinct and highly
advanced section of the genus.
Roots (rhizophores) arise in, or proximal to, the
axils of a stem dichotomy, or occasionally adventitiously on the stem surface. They are dichotomously
branched, generally very smooth and cutinized externally, and in many species are stout and support the
aerial stems; they are also capable of developing, urider certain conditions, into a typical leafy stem and
their morphogenetic origins are a matter of debate.
Root steles are monarch to tetrarch, with the vascular
tissue bilaterally distributed as in Isoetales, Lepidodendrales and most Lycopodiales (Bierhorst 1971).
The leaves of Selaginella are traditionally recognized as microphylls (as in other Lycopsids) and most
species have leaves with a single unbranched vein. Occasional reports indicate a more complex situation (in
S. adunca A. Br. ex Hieron., Mukherjee and Sen 1981;
S. schaffneri Hieron., Wagner et aL 1982). The latter
discuss phylogenetic rdationships. In those genera
with dimorphic leaves, the lower leaves on the primary
stem, especially in erect species, may appear to be uniform and only gradually change shape towards the dimorphic condition on the upper part of the stem. Leaf
shape is diagnostic and is best viewed on the lower
part of a secondary branch. At a branch dichotomy
one lateral leaf appears in the axil of the dichotomy
and is termed 'axillary leaf. It is slightly different in
shape and also diagnostic. Epidermal cells with sinuous (or more rarely straight) anticlinal walls are more
or less consistently isodiametric on the adaxial surface
of lateral and on the abaxial surface of median leaves,
and elongate on the opposing surfaces, respectively;
the cells lying over the midrib may not conform. Callose deposits, which later silicify, may be observed on
young leaves under fluorescent microscopy (Bienfait
and Waterkeyn 1974). Such sculpturing, observed
clearly under the SEM, is consistent and can be useful
in the determination of species (Dahlen, pers. comm.;
Jermy and Quansah, unpublished). Stomata are anomocytic and, for the most part, situated on the abaxial
surface only, either scattered or confined to the midrib
region and margin Or leaf apex (Quansah, pers.
Lycopodiatae • Selaginellaceae
comm.; Gulyas 1961). Ligules can be aciculate-lingulate to obclavate, ranging from 0.10 to 0.45 mm long,
and are consistent within species. Homer et aI. (1975)
have studied the ontogeny and fine structure of the ligule but dismiss any former ideas as to function and
concluded that this is still an enigma. They suggested
that the ligule may be a vestigial organ which has lost
its ability to function.
STROBILUS STRUCTURE AND SPOROPHYLL ARRANGE~
MENT. In a review of 30 species of Selaginella (in subgenera Selaginella and Stachygynandrnm) Homer and
Amott (1963) found three major patterns in the arrangement of sporangia in each strobilus: (1) Strobili
having a basal megasporangiate zone with an upper
zone of microsporangia; (2) strobili having two rows
of megasporangia (occasionally with a few microsporangia) and two rows of microsporangia; (3) strobili
which are wholly megasporangiate. They found a reasonable correlation with the infrageneric classification
proposed by Baker (1883) but more study is required
to ascertain the usefulness of sporophyll arrangement
in the taxon'omy and phylogeny of the genus. The initiation of heterospory occurs sometime between sporangium inception and meiosis (Homer and Beltz
t970), and its significance in phylogeny also needs further investigation.
The morphology of sporophylls reflects that of the
vegetative leaves, basal megasporophylls sometimes
heing slightly larger than the microsporophylls higher
lip the strobilus. In subgen. Heterostachys the larger
sporophylls (which may not always be megasporan·
giate) are folded, forming a lamina flap Or pteryx
(Quansah and Thomas 1985) which partially enfolds
(und protects) the sporangium.
Somers (1982) described an arrangement of thickened cells in the sporangium wall of species in the SeI'ies Articulatae which he claims is unique to that group
and was not seen in other species examined.
SPORE MORPHOLOGY. Spore-wall formation of SelagilIella and its phylogenetic significance within the Pteridophyta has been the subject of research by Pettitt
(1966), Kempf (1970) and A. F. Tryon and Lugardon
( t978). The outer walls of megaspores, composed of
sporopollenin and much silica, have characteristic patlerns which may have taxonomic significance; reticulute and rugose patterns predominate. Similarly, the
sporodenn patterns of microspores, which may be
quite different from those found on the megaspores of
Ihe same species, may show species relationships. Significant regional or systematic surveys have been published (A. F. Tryon 1949; Knox 1950; Hellwig 1969;
Minaki 1984), but more are needed.
(;AMETOPHYTES AND EMBRYOLOGY. There is no special
lI1echanism for spore dispersal, spores of both kinds
41
frequently remaining in the dehisced sporangia until
the surrounding plant tissue decays. In open habitats,
strong winds may help to disperse spores, but ultimately most of the spores will fall around the parent
plant. In mat-forming species [e. g., S. kraussiana
A. Braun, S. apoda (L.) Spring] spores of both kinds
may lie dormant for long periods until the parent plant
dies or is physically removed (e. g., by the foraging of
an animal), when gametophytes of both sexes will germinate and produce sporelings (Webster 1967).
Gametophytes are endosporic and the prothallus is
well developed when the spores are shed. The archegonia and antheridia are similar in structure and mode
of development to those of Isoetes. Antherozoids are
biflagellate and similar to those of Lycopodium. The
embryology is known for only a few species and shows
variation in the development of the suspensor cell and
foot; a wider survey is needed to establish the taxononiic significance of this.
PHYTOCHEMISTRY AND PHARMACEUTICAL USES. No ex·
tensive comparative studies have been carried out on
the Selaginellaceae. Yasuo et aI. (1984) studied the oligosaccharides in nine eastern Asiatic species and
found significant differences. Species are collected locally in India and China and sold in the markets for
medicinal purposes.
CYTOLOGY AND SPOROGENESIS. The first reliable chromosome counts were made by Manton (1950) on three
European species of Selaginella, showing 2n= 18. In
an extensive survey of 76 species, Jermy et aI. (1967)
showed, that four base numbers exist (x= 7, 8, 9 and
10 in subgen. Stachygynandrnm and x=9 in subgen.
Selaginella and Heterostachys), with triploids and
tetraploids being found in several species with x= 9.
There was no correlation between chromosome base
number and subgeneric classification, but with pattern
of coning: those with x=9 producing cones on iso·
lated branches sporadically, those with x= 10 coning
simultaneouslY on all branch endings of the primary
branch system or pseudofrond. The size of the chromosomes shows considerable interspecific variation,
althoug all complements are very small. Subsequently
n= 12 has been reported (Panigrahi). Hybrids have
been reported in subgen. 1I!tragonostachys (R. M. Tryon 1955), determined by intermediate morphology and
spore abortion, but the cytology was not checked.
Pettitt (1971), in investigating developmental mechanisms in heterosporous plants, showed that in the megasporangium of S. sulcata (Desv.) Spring (and presumably most other species) all but one of the
megasporocytes either degenerate, or persist without
completing the meiotic cycle, presumably as diploid
cells. This situation could lead to diploid megaspores
and thus agamospermy, a condition indicated by records of megasporangia with single or two spores (Ly-
42
Lycopodiatae . SeJaginellaceae
on 1902; R. M. Tryon 1955). Agamospermy was also
shown in the triploid S. tenerrima A. Br. ex Kuhn, a
true therophyte which survives the dry seaSOn only as
spores (Kornas and Jankun 1983).
SUBDIVISION. The family is presently regarded as monotypic although Rothmaler (1944) resurrected the
earlier generic names Lycopodioides Boehmer and
Didic/is Beauv. for some European species, without
any significant taxonomic revision. Kunkel (1963)
published Stachygynandrum myosurus (Sw.) Kunkel
[= Selaginella myosurus (Sw.) Alston) but again without discussion.
At the subgeneric level, Baker (1883) divided the genus into four subgenera and this was followed by Walton and Alston (1938). Hieronymus (1901), On the
other hand, followed the arrangement of Spring (1850)
and raised his Sections, Homoeophyllae and Heteraphyllae, to sub generic rank, thus amalgamating Baker's
three subgenera with dimorphic leaves into a single
subgenus, Heterophyllum, a situation followed by Tryon and Tryon (1982). Somers (1982 and pers. comm)
suggested that the Series Articulatae (sub gen. Stachygynandrum) are sufficiently distinct to warrant consideration as a separate subgenus, but made no firm proposal. The group is certainly distinct and mOre investigation is needed. The present treatment propose five
subgenera based On morphology and distribution.
DISTRIBUTION, ECOLOGY AND PHYSIOLOGY. Subgen.
Selaginella: one species, S. selaginoides (L.) link, is a
base-rich mire plant with a circum boreal range extending south to the Canaries, but absent from Africa, and
in North America, south to latitude 42 0 N. The second
species (S. dej/exa Brackenridge) is endemic to bogs in
the Hawaiian archipelago.
Subgen. Tetragonostachys is composed of plants
characteristic of seasonally dry areas, ranging from
southern North America across the tropics in South
America, Africa and the Indian Subcontinent to northern China and Japan, a few species extending to more
mesophytic heathy woodlands in north-temperate
areas. Those living in dry areas have thick cutinized
leaves with fine hair points which protect the growing
shoot apex.
Subgen. Ericetorum: S. uliginosa (Labill.) Spring is
confined to proteaceous heathlands throughout the
Australian continent and two further species of similar
habitats are annuals, one IS. gracillima (Kunze) Spring)
in SE Australia, the other [S.pygmaea (Kaulf.) Alston)
in southern Africa.
Subgen. Stachygynandrum and subgen. Heterostachys: widespread in all continents; predominantly terrestrial plants of lowland to mid-montane primary rainforest but preferring more open glades and river
banks, and therefore a frequent component of secondary forest in these areas. Occasionally found as
epiphytes on mossy tree trunks, and some with a
climbing habit [e. g., S. willdenowii (Desv.) Baker,
S. myosurus (Sw.) Alston) can soon become smothering
weeds in gaps and clearings (Wong 1983). The latter
species commonly displays a metallic blue tinge On its
leaf surface which may act as an interference filter, increasing the transmission of light at the red end of the
spectrum, those wavelengths most used for photosynthesis at the forest floor level. In addition, the leaves of
this species possess epidermal cells with regular COnvex surfaces which may focus light on the singie, large
chloroplast resting at its distal end (Lee 1977). A few
species reaching cooler temperate regions, e. g., S. denticulata (L.) link of the Mediterranean area and S.lepidophylla (Hook. & Oreville) Spring of Mexico, are
adapted for seasonal drought by having the ability to
inroll both leaves and sterns to prevent excessive water
loss. The leaves ot the latter and similar species, sold
under the name of "Resurrection plants", can withstand repeated drying/wetting without showing lesions in the cell membranes (Leopold et al. 1981). An
alternative strategy is seen in S. tenerrima A Br. ex
Kuhn, a typical therophyte from south-central Africa.
This species will live only for the few weeks of the wet
season and then reproduce by agamospermous megaspores, very few, if any, micros pores being formed
(Kornas and Jankun 1983).
PALAEONTOLOGY. ligulate, heterosporous and bisporangiate, herbaceous plants which probably gave rise to
Selaginellaceae appeared amongst the arborescent Lycophytes of the Lower Carboniferous. A significant
fossil seen throughout the Carboniferous is Paurodendron, now regarded as Selaginella jraipontii (Leclerque) Schlanker & Leisman. It shows some Sigillarian characteristics but may also be compared in general morphology to the extant S. selaginoides. The majority of fossil forms of Selaginella-like plants (Selaginellidites, Selaginellites) have been described from megaspores from the Cretaceous and Tertiary sediments
in the course of studies on pollen/spore assemblages.
A single genus:
Selaginella Pal. Beauv.
Figs. 3, 13, 14
Selaginella Pal. Beanv., Mag. Encycl. Paris 9 (5): 478 (1804),
nom. conserv.
Description as for the family. The following subgenera
are recognized here:
Subgenus Selaginella (2 species). Stems erect, new
primary shoots arising from the base upon maturation
of the single terminal strobilus, rooting from a basal
hypocotular node; leaves and sporophylls are spirally
arranged, uniform and herbaceous.
Subgenus Ericetornm Jermy, Fern Gaz. 13: 117
(1986) (3 species). Stems erect, either unbranched or
more compound, arising from a creeping solenostelic
stem; leaves uniform, decussately arranged at least below, more or less herbaceous; sporophylls tetrastichous.
Subgenus Tetragonostachys Jermy, Fern Gaz. 13:
118 (1986) (c. 50 species). Prostrate or low suffruticose
plants, often mat-forming, stems much-branched, rooting throughout; leaves spirally arranged, uniform or
with a tendency towards dimorphism on prostrate
branches (possibly as a result of one-sided illumination), usually coriaceous, Iinear-lanceolate or sometimes needle-like with long hair points; sporophylls
tetrastichous.
Subgenus Stachygynandrnm (PaL Beauv.) Baker, J.
Bot. London 21: 3 (1883) (c. 300 species). Primary
stems either creeping, branches semi-prostrate, rooting
at the axils of branch dichotomies, or erect, rooting
usually at the base of main stem only, suffruticose or
with elaborate compound branching systems; primary
and secondary branches dichotomously branched, frequently in such a way as to form a regular pseudopinnate arrangement in one plane, pseudofronds with a
characteristic outline, ultimately producing strobili simultaneously at the branch apex, or in an irregular
pattern in which only the upper branches produce
strobili, the lower vegetative branches becoming fertile
on the maturation of the terminal one above; leaves
dimorphic, at least on the secondary branches, in four
distinct rows, those of the two upper (dorsal or median) rows being distinctly smaller than those of the lower (ventral or lateral) rows; leaves on erect primaiy
stems often scarcely dimorphic; sporophylls uniform
and tetrastichous.
Subgenus Heterostachys Baker, J. Bot. London 21: 4
(1883) (c. 60 species). Stems creeping and muchbranched, or secondary branches erect and suffructicose, rooting at branch axils; leaves as in sub gen. Stal'hygynandrnm; strobili complanate, sporophylls dimorphic, tetrastichous, those on the ventral side
smaller than those on the dorsal side of the shoot.
1"lg.13A-E. Selaginellaceae. A Selaginella hordeiformis
lIaker, habit (x 1), leaves dimorphic on base of stem. B,
(' Selaginellafirmuloides Warb. B Habit (x 1), leaves on stem
hase monomorphic. C Strobilus with monomorphic sporoI,hylls ( x 6). D, E Selaginella /curzii A. Br. ex Warb. D Part of
,Imbilus with dimorphic sporophylls (x 10). E Part of sterile
portion of stem with median and lateral leaves (x 10) (A-C
rrom Brownlie 1969; D, E from Panigrahi and Dixit 1968)
C
44
Lycopodiatae . Selaginellaceae
Fig. 14 A-C. Selaginellaceae. A, B SelagineUa apoda (L)
Morren. A Habit t x 1.5). B Detail of sterile portion of plant
(x 12).C Selaginella myosurus (Sw.) Alston, strobilus and vegetative branch bearing it(x 6) (A, B from Billington 1952; C
from Alston 1959)
Selected Bibliography
Alston, A. H. G. 1932. Selaginellaceae, in C. Christensen, The
pteridophyta of Madagascar. Dansk Bot. Ark. 7: 193-200.
Alston, A. H. G. 1934a. An enumeration of the Chinese species of Selaginella. Bull. Fan Inst. Biology (Bol) 5:
2611-294.
Alston, A. H. G. 1934 b. The genus Selaginella in the Malay
Peninsula. Gdns' Bull. 8: 41-62.
Alston, A. H. G. 1935a. The Philippine species of Se/aginella.
Philipp. J. Sci. 58: 359-383.
Alston, A. H. G. 1935b. The Selaginellae of the Malay Islands I. Java and the Lesser Sunda Islands. Bull. Jard. Bot.
Buitenzorg III, 13: 432-442.
Alston, A. H. G. 1937. The Selaginellae of the Malay Islands II. II. Sumatra. Bull. Jard. Bot. Buitenzorg III, 14:
175-186.
Alston, A. H. G. 1939a. Notes on Selaginella: IX. The South
African species. J. Bot. London 77: 221-224.
Alston, A. H. G. 1939b. The Se/aginellae of Argentina, Uruguay and Paraguay. Physis (Buenos Aires) 15: 251-257.
Alston, A. H. G. 1940. The Selaginellae of the Malay Islands.
III. Celebes and the Moluccas. Bull. Jard. Bot. Buitenzorg III, 16: 343 - 350.
Alston, A. H. G. 1945. An enumeration of the Indian species
of Selaginella. Proc. Nat!. Inst. Sci. India 11: 211-235.
Alston, A. H. G. 1951. Selaginellacees. In: Aore generale de
l'Indo-Chine, vol. 7. (H. Lecomte, Ed.) Paris: Masson &
Cie. pp.555-594.
Alston, A. H. G. 1952. A revision of the West Indian species
of Selaginella. Bull. Brit. Mus. (Nat. Hist.) Bot. 1: 25-47.
Alston, A. H. G. 1955. The heterophyllous Selaginellae of
continental North America. Bull. Brit. Mus. (Nat. Hist.)
Bot. 11: 219-274.
Alston, A. H. G. 1957. Selaginellaceae. In: Alston, A. H. G.,
Tardieu-Blot M. (Eds) Les pteridophytes de l'Afrique
intertropicale fran""ise. Mem. Inst. Fr. Afr. Noire 50:
26-44.
Alston, A. H. G. 1959. The ferns and fern allies of West Tropical Africa, being a supplement to the second edition of the
flora of West Tropical Africa (R. W. Keay, Ed). London:
Crown Agents.
Alston, A. H. G., Jerrny, A. c., Rankin, J. M. 1981. The genus
Selaginella in tropical South America. Bull. Brit. Mus.
(Nat. Hist.) Bot. 9 (4): 233-330.
Baker, J. G. 1883. A synopsis of the genus SelagineUa. pt. 1.
J.Bot. London 21: 1-5.
Baker, J. G. 1887. Handbook of the fern-allies. London: Bell.
Bienfait, A., Waterkeyn, L 1974. Contribution" l'etude systematique des Selaginella: specificite des formations callosiques foliaires observees en fluorescence. Bull. Jard. Bot.
Nat. Belg. 44: 295-302.
Billington, C. 1952. Ferns of Michigan. Bull. Cranbrook Inst.
Sci., 32:114.
Gulyas, S. 1961. Untersuchungen an der Blattepidermis der
Selaginellen. Acta BioI. Univ. Szeged, N. S. 7: 15-24.
Hellwig, R. L 1969. Spores of the heterophyllous Selaginellae
of Mexico and Central America. Arm. Missouri Bot. Gard.
56: 444-464.
Hieronymus, G. 1901. Selaginellaceae. In: Engier, A., Prant!,
K. (Eds). Die natiirlichen Pflanzenfamilien I, 4. Leipzig:
Engelmann, pp.621-715.
Horner, H. T., Arnott, H. J. 1963. Sporangial arrangement in
North American species of Selaginella. Bot. Gaz. 124:
371-383.
Horner, H. T., Beltz, C. K. 1970. Cellular differentiation of heterospory in Selaginella. Protoplasma 71: 335-341.
Horner, H. T., Beltz, C. K., Jaegels, R., Boudreau, R. E. 1975.
Ligule development and fine structure in two heterophyllous species of Selaginella. Can. J. Bot. 53: 127-143.
Jermy, A. C. 1970. Selaginellaceae. In: Schelpe, E. A. C. L E.
(Ed.) Aora Zambesiaca, Pteridophyta. London: Crown
Agents, pp.22-30.
Jerrny, A. C. 1986. Two new Selaginella species from Gunung
Mulu National Park, Sarawak. Kew Bull. 41: 547-559.
Jerrny, A. c., Jones, K., Colden, C. 1967. Cytomorphological
variation in Selaginella. Bot. J. Linn. Soc. 60: 147-158.
Kempf, E. U. 1970. Elektronenmikroskopie der Sporoderrnis
von Megasporen der Gattung Selaginella (Pteridophyta).
Rev. Palaeobot. Palynologie 10: 99-116.
Knox, E. M. 1950. The spores of Lycopodium. Phylloglossum.
Se/aginella and Isoetes and their value in the study of mi-
Selected Bibliography
crofossils of Palaeozoic age. Trans. Bot. Soc. Edinburgh
35: 209-357.
Kornas, J., Jankun, A. 1983. Annual habit and apomixis as
drought adaptations in Selaginella tenerrima. Bothalia 14:
647-651.
Kunkel, G. 1963. Die Formvariabilitat einiger westafrikanischer Farne. Nova Hedw. 6: 199-217.
Lee, D. W. 1977. On iridescent plants. Gard. Bull. Singapore
30: 21-29.
Leopold, A. c., Musgrave, M. E., Williams, K. M. 1981. Solute leakage resulting from leaf desiccation. Plant Physiol.
68: 1222-1225.
Loyal, D. S. 1976. Chromosome counts in north-western Himalayan species of Selaginella·I. Proc. 63rd Indian Sci.
Congr. Pt. 3: 127-128.
Lyon, F. M. 1902. Two megasporangia in Selaginelia. Bot
Gaz. 36: 308.
Mickel, J. T., Hellwig, R. L 1969. Actino-plectostely, a complex new stelar pattern in Selaginella. Amer. Fern J. 59:
123-134.
Minaki, M. 1984. Microspore morphology and taxonomy of
Selaginelia (Selaginellaceae). Pollen Spores 26 (3-4):
421-480.
'
MukheIjee, R. N., Sen, U. 1981. A forked vein and foliar
fibres in Selaginelia. Fern Gaz.12: 175-177,
Parris, B. S., Jermy, A. C., Camus, J. M., Paul, A, M, 1984. The
Pteridophyta of Gunung Mulu National Park, Sarawak.
In: Studies on the flora of Gunung Mulu National Park,
Sarawak (A. C, Jermy, Ed.). Kuching: Forestry Department, pp.145-233.
Pettitt, J. M. 1966. Exine structure in some fossil and recent
spores and pollen as revealed by light and electron microscopy. Bull. Brit Mus. (Nat. Hist.) Geol. 13: 223-257.
Pettitt, J. M. 1971. Developmental mechanisms in heterospory. I. Megasporocyte degeneration in Selaginelia. Bot J.
Linn. Soc. 64: (3): 237-246.
Proctor, G. R. 1985. Ferns of Jamaica. A guide to the Pteridophytes. London: British Museum (Natural History),
45
Quansab, N., Thomas, B. A. 1985. 'Sporophyll-pteryx' in African and American Selaginella. Fern Gaz. 13(1): 49-52.
Reed, C F. 1%6. Index Selaginellarum. Mem. Soc. Brot. 18:
1-287.
Rothmaler, W. 1944. Pteridophyten-Studien I. Fedd. Rep. 54:
55-82.
Smith, A. R. 1981. Flora of Chiapas II. Pteridophytes, San
Francisco: Cal. Acad. Sci.
Somers, P. 1982. A unique type of microsporangium in Selaginella, Series Articulatae. Amer. Fern J. 72: 88-92.
Spring, A. 1850. Monographie de la famille Lycopodiacees:
2. Selaginella. Mem. Acad. R. Sci. Lett Belg. 24: 52-264,
Stolze, R. G. 1983. Ferns and fern-allies of Guatemala III.
Field. Bot N. S. 12: 1-91.
Tagawa, M., Iwatsuki, K. 1979. Flora of Thailand 3 (1): Pteridophytes: 1-128.
Tryon, A. F. 1949, Spores or'the genus Selaginella in North
America, north of Mexico. Ann, Missouri Bot Gard. 36:
413-431.
Tryon, A. F., Lugardon, B. 1978. Wall structure and mineral
content in Selaginellaspores. Pollen Spores 20: 315-340.
Tryon, R. M, 1955. Selaginella rupestris and its allies. Ann.
Missouri Bot Gard. 42: 1-99.
Wagner, W. H., Beitel, J. M., Wagner, F. S. 1982. Complex
venation patterns in the leaves of Selaginella: megaphylllike leaves of Lycophytes. Science 218: 793-794.
Walton, J" Alston, A. H. G. 1938, Lycopodiinae. In: Verdoorn, F. (Ed.) Manual of pteridology. The Hague: Nijhoff, Chap. 17.
Webster, T. R. 1967. Induction of Selaginella sporelings under
greenhouse and field conditions. Amer. Fern J. 57:
161-166.
Wong, K. M. 1983. Critical observations on peninsular
Malaysian Selaginella, Gard, Bull. Singapore 35: 107 -135.
Yasuo, S., Konishi, T., Kiyosawa, S. 1984, Studies on oligosaccharides of plants in the Selaginellaceae in Japan.
Shoyakugaku Zasshi 38: 207-211.
Equisetatae
Sporophytes with roots, stems and whorled leaves.
Protostelic-derived, medullated, some with secondary
thickening. Sporangia thin-walled, homosporous to
heterosporous. Antherozoids multiflagellate.
A single surviving family.
Equisetaceae
R.L.HAUKE
Equisetaceae L. C. Richard ex DeCandolle, FI. Fr. 2: 580.
(1805).
Rhizomatous perennials with stems characterized by a
jointed appearance, with leaves small, whorled, and
fused into a nodal sheath. A series of ridges and valleys traverse the internodes, and continue up into the
nodal sheaths, alternating in successive internodes.
Stems in cross-section usually with a prominent central canal and small vallecular (under the valleys) and
carinal (under the ridges) canals. Reproduction by
spores borne in elongate, longitudinally dehiscing
sporangia on sporophylls in cones terminal on the vegetative stem, or occasionally on branches, or in some
species on a specialized cone-bearing stem. Spores
spherical, green, of one kind, with elaters (hapters);
gametophytes green, terrestrial. Chromosome number
n = 108. Monogeneric.
ANATOMY AND MORPHOLOGY. Recent accounts by
Bierhorst (1971) and Foster and Gifford (1989) summarize the morphology and anatomy of Equisetum.
The rhizome system is extensive, and of great importance in the persistence and spread of this genus. It
bears roots, tubers (in some species), and aerial
branches. These initiate as nodal buds, at the base of
the internode above and enclosed by the nodal sheath.
Each bud has a root apex and a shoot apex, but often
only one will develop further. AIl are determinate
structures, the tuber developing only one or two internodes which become gorged with starch.
The aerial shoots have sharply defined nodes and
internodes because the leaves are whorled and fused
into a nodal sheath. The internode has ridges and
grooves running its length, and the ridges become the
midribs of the leaves (sheath segments) and continue
up into their unfused tip portions (sheath teeth).
Leaves, and hence the internode ridges, alternate at
adjacent nodes. Stomata are found along the grooves,
either scattered or in bands on either side, or in single
rows on each side.
A transverse section of the internode shows a pattern of collenchyma (often mistakenly called sclerenchyma) and chlorenchyma under the epidermis, vallecular canals in the cortex under the grooves, and
carinal canals under the ridges. These are protoxylem
lacunae. The vascular system is a ring of bundles, each
with a small amount of phloem flanked by xylem, and
with the proto xylem lacuna internal to the phloem. Associated with the vascular system is an endodermis,
which appears as an individual endodermis around
each bundle, or a single outer endodermis enclosing
the ring of bundles, or a double common endodermis,
one outside and the other inside the ring of bundles.
The pith region is occupied by a large central canal.
The collenchyma may be massed under the grooves,
under the ridges, or both. The epidermis is covered
with a silica layer which develops characteristic micromorphology (Hauke 1979).
Branches develop regularly in some species, irregularly in others, and only as an injury response in others. These branches repeat the morphology of the
main stem. At the base of the internode is an intercalary meristem which develops from the lower part of
the leaf primordium. Buds initiate exogenously from
the uppermost cell of the sheath primordium, between
leaf segments. These are enclosed by the nodal sheath
and erupt through it, alternating with the leaves. With
its often dormant nodal buds and intercalary meristems, this genus can be very plastic in gross morphology, and as a result a large number of meaningless taxonomic names have been published (Hauke 1963,
1966, 1979).
Cones terminate the stems. In most species these are
the vegetative stems, but in a few, the cone-bearing
stems lack chlorophyll, are unbranched and succulent,
with enlarged sheaths. These may be ephemeral, or
they may persist and become chlorophyllous and
branched after the spores are shed. The cone consists
of whorls of sporophylls, each a peltate structure with
about six elongate sporangia pendant from its adaxial
surface. Sporangia dehisce longitudinally to release
Equisetum
Fig. 15 A-G. Equisetaceae. A-D Equisetum ramosissimum.
A Habit (x 0.6). B Node with brancb (x 5.25). C Strobilus
( x 4). D Schematic transection through stem (x 8.25).
~;-G Equisetum arvense. E Habit of sterile plant (x 0.5).
F Schematic transection through stem (x 12). G Apex of fertile stem with strobilus (x 1) (A-F from M~dalski 1954)
Ihe spherical, alete, chlorophyllous spores. These have
perispore which becomes four strap-like hygroscopic
daters (hapters) and an unmarked spore wall; SEM
pictures reveal a granulate surface with a scattered
Nl,hericai deposit. The spores have a limited viability
(ahout 1 week) and require light for germination.
Gametophytes are epigeal, may grow to about 1 cm
ill diameter, and have a parenchymatous basal portion
with unicellular rhizoids ventrally and plates of chlowphyllous tissue dorsally. Although the spores are
homosporous, the gametophytes are unisexual in most
cases (Duckett 1970, 1972, 1979; Hauke 1969, 1977,
II
47
1980). The archegonial gametophytes, if unfertilized,
commonly change over to antheridial, and during a
short period may be functionally bisexual and self-fertile.
ECOLOGY AND DISTRIBUTION. Equisetum is primarily a
plant of open, sunny sand banks along river and lake
margins, in marshes, and in other wet places. One
grows in open, standing water and several inhabit
shaded, wet woods. Because of the widespread filling
of low, wet places to make roads and railroad beds,
some species have become common along roadsides
and railroad embankments. This genus is found
around the world from the southern portions of South
America and Africa to above the Arctic Circle. The
greatest concentration of species are found between
40° and 60° north latitude. Of the major land masses,
only Australia, NewZealand, and Antarctica have no
representatives.
48
Equisetatae . Equisetaceae
AFFINITY. This family with its one genus stands alone
among extant vascular plants. The many unique features of its anatomy and morphology justify its being
segregated as a distinct class, subdivision, or even, by
some authors, a division of the plant kingdom.
KARYOLOGY AND HYBRIDIZATION. The chromosome
number of all species that have been counted (12 of
15) is n = 108. There is some indication of slight differences in the form of the chromosomes of the two subgenera. Numerous hybrids have been described be·
tween species within each subgenus, but none are
known between the two subgenera. Evidence for dip·
loidy, notwithstanding the high chromosome number,
was presented by Soltis (1986).
Equisetum L.
Fig. 15
Equisetum L., Sp. Pl.: 1061 (1753).
Hippochaete Milde (1865).
Presfa Dulac (1867).
Characters of the family.
KEy TO SUBGENERA.
1. Stomata superficial, usually scattered or in bands 2 or more
stomata wide; COnes non-apiculate; aerial stems mostly annual, regularly branched, with an outer common endodermis; antheridia with more than two cover cells
Subgenus Equise/llm
- Stomata sunken, usually in single lines; cones mostly apiculate; aerial stems usually perennial, often unbranched,
with a double COmmon endodermis; antheridia with only
2 cover cells
Subgenus Hippoc/uImI
Although a number of workers have elevated these to
genera, the only morphological character which sepa-
rates all the species of one from all the species of the
other is the superficial vs. sunken stomatal apparatus.
Subgenus Equisetum: Eight species, predominately
northern in distribution, with only one species south of
the 20° north latitude.
Subgenus Hippochaete (Milde) Baker: Seven species,
widespread in both southern and northern hemispheres.
Selected Bibliography
Duckett, J. G. 1970. Sexual behaviour of the genus Equisetum. subgenus Equisetum. Bot. J. Unn. Soc. 63: 327-352.
Duckett, J. G. 1972. Sexual behaviour of the genus Equisetum, subgenus Hippochaete. Bot. J. Linn. Soc. 65: 87-108.
Duckett, J. G. 1979. Comparative morphology of the gameto
phytes of Equisetumsubgenus Hippochaete, etc. Bot. J. Unn.
Soc. 79: 179-203.
Foster, A. S., Gifford, E. M. 1989. Comparative morphology
of vascular plants, 3rd edn., Chap. 10: The Sphenophyta.
San Francisco: W. H. Freeman & Co.
Hauke, R. L. 1963. A taxonomic monograph of Eljuisetum
subgenus Hippochaete. Beih. Nova Hedw. 8: 1-123.
Hauke, R. L. 1966. A systematic study of Equisetum arvense.
Nova Hedw. 13: 81-109.
Hauke, R. L. 1969. Gametophyte development in Latin
American horsetails. Bull. Torrey Bot. Club 96: 568-577.
Hauke, R. L. 1977. Experimental studies on growth and sexu·
al determination in Equisetum gametophytes. Amer. Fern
J.67: 18-31.
Hauke, R. L. 1979. A taxonomic monograph of Equisetum
subgenus Eljuisetum. Nova Hedw.30 (1978): 385-455.
Hauke, R. L. 1980. Gametophytes of Equisetum diffusum.
Amer. Fern J. 70: 39-44.
Soltis, D. E. 1986. Genetic evidence for diploidy in Equisetum. Amer. J. Bot. 73: 908-913.
Filicatae
Notes on the Higher Level Classification
of the Recent Ferns
K. U.KRAMER
Sporophytes with roots, stems and spirally arranged
leaves (megaphylls). Protostelic, solenostelic or dictyostelic, sometimes polycyclic. Some with limited secondary thickening. Sporangia thick- or thin-walled,
homosporous or heterosporous, borne on leaves.
Gametophytes axial or flat, mycotrophic or autotrophic. Antherozoids multiflagellate.
33 families.
A firm basis for the alignment of the leptosporangiate
fern families, which by themselves are becoming increasingly understood and well-defined, is still lacking.
In the treatment of the families given below, ideas
about their affinites are presented; but as the family
sequence is alphabetical, in accordance with the editorial rules for ''The Families and Genera of Vascular
Plants", rather than being "natural", it is very difficult
to glean an overview of supposed family relationships
from the present work. I therefore give a resume of
what I think is the present state of knowledge (or, in
many cases, ignorance) about family relationships in
the leptosporangiates (Fig. 16). As the views expressed
below are essentially my own, I have chosen the first
person singular for stating them, in contrast to the
style adopted elsewhere in the work.
Opinions diverge widely as to the relationships (if
any exist) between the two eusporangiate families; see
the remarks given with the family treatments of the
Ophioglossaceae and Marattiaceae. The leptosporangiates only will therefore be dealt with more at length.
When the artificiality of the Polypodiaceae sensu
Christ (1897) and Diels (1899/1900) was recognized see especially Ching (1940) - various new schemes of
classification were proposed for the now greatly increased number of leptosporangiate fern families. In
these schemes, attempts were often made at relating
the segregates of the former, inclusive Polypodiaceae
to each other in various ways, but also, and especially,
to various families of the more primitive leptosporangiates from which they were now often supposed to
have been derived independently. Important attempts
at such a classification were made by Holttum (1949),
Nayar (1970), Mickel (1974), Crabbe et al. (1975), Lovis (1977), Pichi Sermolli (1977), and Tryon and Tryon
(1982); and, on a more regional scale, i. e., not comprising all genera and families of the world, by Ching
(1978).
Of these, the system by Crabbe et al. is not a true
system of classification, as only a linear sequence is
given which of course can only very imperfectly reflect
ideas about affinity; it was devised as a sequence of
families and genera for the fern herbarium. Pichi Sermolli's system (and also Nayar's) is based on a philosophy of emphasizing differences rather than similarities, which is alien to the principles of classification
and the spirit underlying the present work. He tended
50
Notes on the Higher Level Classification of the Recent Ferns
Aspleniaceae
r--------------,
: Blechnaceae
: lomarlopsidaceae
: Davalliaceae
I I Nephrolepidaceae
I: Oleandraceae
:
:
:
I
:
I L_~:~~~~~a~___ J
Thelyptarldaceae
r---- - - - - - - - - - - - - - - - - - - ,
I
Dennstaedtiaceae
:
I
Cyatheaceae
I
Metaxyaceae Lophosonaceae :
Dicksonlaceae
I
Hymenophyllopsldace.e
Monachosoraceae
It.. _____________________ .JI
Hymenophyllaceae
~ataceae
II
Gleichenlaceae
Dipte(od.ce.e Cheiropleuriace.el
. Matoniaceae
--- -- - ----
I
L
Plagiogyriace.e
Osmundaceae
affinities very probable
affinities possible, not unlikely
affinities possible but very uncertain
Fig.16. Graphic representation of supposed affinities between fern families. Primitive families at the hollom. derived
families above. In the right column isolated families or groups
of families. Ophioglossaceae and Marattiaceae do not fit into
this scheme
to retain taxa whose distinctness had long been strongly challenged, like keeping Loxoscaphe apart from Asplenium. Sambirania from Lindsaea. etc. An extreme
example is the recognition of an order Negripteridales,
based on the monogeneric family Negripteridaceae;
other authors like Tryon and Tryon (1982) doubt or deny (R. Tryon, present work) even generic distinctness
of Negripteris from Cheilanthes. Thus, it becomes difficult to find a basis for comparison of Pichi Sermolli's
system with that of most other authors, including the
one adopted here. Lovis' system has very considerable
merits but tends to overemphasize a single character
complex, the chromosome numbers. Recent findings
of divergent numbers in otherwise homogeneous genera have shown that this character, too, must be handled with caution, although it would seem to be of
rather fundamental significance. For example, most
but not all species of Asplenium are based on x= 36, of
Pteris on x= 29, etc., but such (few) divergent numbers
as there are do not make these genera artificial as
some authors (in these cases, not Lovis!) would have
it.
The task of finding relationships among the primitive leptosporangiates is particularly difficult. Several
of the families have a long geological record and are
evidently the last relict representatives of ancient lineages; it may seem downright methodologically incorrect to try and relate them to other extant families
when they have become so strongly isolated by a long
evolutionary history and drastic extinction among
their members and relatives. Families now occupying
such isolated positions are Loxomataceae, Matoniaceae, Dipteridaceae, Cheiropleuriaceae, and Gleicheniaceae. Except for the first, they are sometimes
placed side by side in a primitive alliance, e. g., by Lovis. Except for the obvious relationship between the
third and the fourth, I can see little similarity between
these families, which in my opinion should not be
squeezed into a group of arguable naturalness. The
same holds for the Hymenophyllaceae. I cannot see
any evidence for placing them near the basis of an alliance "Hymenophyllidae" that further comprises the
majority of the "indusiate leptosporangiates", as was
done by Pichi Sermolli; and the placement of the Hymenophyllopsidaceae near them seems altogether fanciful. Uke certain other predominantly epiphytic
groups, the Hymenophy11aceae are morpho-ecologica\1y so highly specialized that their relationships cannot at present be determined, and the structure of their
receptacle is unique.
Further isolated families are Osmundaceae and Plagiogyriaceae, the former of considerable antiquity. I
agree with Holttum (1949) that they may be distantly
related, but no other group seems to have its natural
place near them.
Another difficult case is represented by the Schizaeaceae, also an old family. They are often connected
with the (or some) "Gymnogrammoid" ferns (Pteridaceae); the resemblance of Mohriato Cheilanthesis certainly suggestive but convergence cannot be ruled out.
At the present state of our knowledge the affinity
seems hypothetical at best. A defendable hypothesis but no more than that - seems the derivation of the
Marsileaceae from Schizaeaceous stock. These heterosporous ferns are, however, again so strongly specialized that a more definite statement seems hazardous
until new evidence turns up. The Salviniales, of
course, have nothing to do with the Marsileales, and
their ancestry remains quite obscure.
Holttum and Sen (1961) have re-established the relationship between Dicksoniaceae and Cyatheaceae,
although most other workers would not unite these
two families into one, as they proposed. Lophosoriaceae and Metaxyaceae are clearly also related, and
these four families constitute a natural alliance that
might be treated as an order. Dennstaedtiaceae are in
my qpinion related to Dicksoniaceae, through genera
like Calochlaena. and although they are more derived,
placement in the same order, or at most a neighbouring one (in view of divergent spore characters and
Notes on the Higher Level Classification of the Recent Ferns
chromosome numbers), would be justified. A circumscription of the Dennstaedtiaceae as proposed by
Holttum (1947, 1949) is no longer tenable. There does
not seem to be any other family among the higher leptosporangiate ferns that shows clear affinities with the
Dennstaedtiaceae, with the possible exception of the
Monachosoraceae whose taxonomic disposition is,
however, disputed and obscure.
As pointed out above, the Pteridaceae may be
distantly related to the Schizaeaceae, but convincing
proof is as yet lacking. More probably related to the
former are the Vittariaceae, a genus like Rheopteris (if
this is indeed correctly placed in the Vittariaceae) providing a kind of link with the Pteridaceae. The epiphytic habitat has evidently simplified the Vittariaceae
so strongly that they are likely to be very different in
appearance from any group that may be ancestral.
Holttum has argued repeatedly (1969, 1981) that the
Thelypteridaceae are an offshoot from a Cyatheoid
ancestor. In my opinion anatomical and karyological
evidence does not much support such a derivation; in
spore characters the two diverge strongly, too. I am
still not convinced that there may not be a link between Thelypteridaceae and Athyrioid ferns, but the
admittedly not very close similarities may be deceptive.
A fairly closely knit group of hlgher leptosporangiate fern families is formed, I think, by the Dryopteridaceae (sensu lato, including the Athyrioid ferns with
Woodsia and the Onocleoid ferns), Oleandraceae,
Nephrolepidaceae, Davalliaceae, Lomariopsidaceae,
and Blechnaceae. These families are sufficiently close
to make it a matter of taste where to draw the exact
limits between some of them: Rumohra to be placed in
Davalliaceae or Dryopteridaceae? Nephrolepis placed
near Oleandra or not? These two included in Davalliaceae or not? Lomariopsidaceae separated from Dryopteridaceae or included as a subfamily? The Blechnaceae stand farther apart, in terms of morphology as
well as of karyology. Anatomically, however, they are
not very far from the Dryopteridaceae and I think that
inclusion in the present group (order, if desired) can be
confidently advocated.
Aspleniaceae are in my opinion a rather isolated
family (genus), incertae sedis among the hlgher leptosporangiates. I cannot accept (nor even understand)
that Mickel (1974) placed the Athyrioids nearer to the
Asplenioids than to the Dryopteridoids, and I cannot
see how a family Aspleniaceae as delimited by Crabbe
et aI. can be at all defined. The petiolar anatomy, the
structure of the scales, the chromosome number, to
name only some more obvious characters, seem to preclude any assumption of close affinity to the Dryopteridoid alliance; only spore characters provide some
51
positive evidence. Pichl Sermolli believed the Aspleniaceae to be closer to the Thelypteridaceae, but again I
fail to understand on what grounds. Asplenium seems
to me to be a fairly recent genus, fully in the process of
evolution, with a high degree of interfertility between
morphologically divergent members. Moreover, at present we do not have any idea whlch part of Asplenium
may be basal ("primitive") which causes difficulty in
relating the family to another one. In accepting the
isolated position of Asplenium I agree more with Pichi
Sermolli than with other authors.
I
The last major group to be discussed are the PolypodiaceaeiGrammitidaceae. Whereas an affinity between the two was long denied or at least doubted, recent work has provided arguments for regarding them
as quite close; inclusion in one ,family, as advocated by
Tryon and Tryon, may even be considered. But where
are t.he roots of thls alliance? The Polypodiaceae are
usually derived from a primitive, exindusiate leptosporangiate group of families (see above). I cannot
find sufficient support for such a derivation, and I
think epiphytism has so strongly affected the structure
of these ferns as to obscure their affinities insofar as
they can be traced by currently available methods. Jarrett (1980) has argued convincingly that the Polypodiaceae and Grammitidaceae cannot be regarded as derivatives of a "primitive exindusiate stock" if indeed
there is (or was) such a thing.
Three genera remain that in my opinion cannot at
all be satisfactorily accomodated within the framework of leptosporangiate families. Hymenophyllopsis is
definitely not related to Hymenophyllaceae but may
not be too far away from Dennstaedtiaceae; further
evidence, particularly from karyology, is urgently
needed. The genus is distinct enough for placement in
a family by itself. Monachosorum is less distinct, but
the evidence as to affinity from its various characters is
so equivocal that it is placed in a family of its own,
too, although this constitutes an admission of ignorance rather than of knowledge. Finally, Pleurosoriopsis is evidently a small, reduced, and much derived genus, but derived from what? This is evidently still quite
enigmatic; but the genus lacks distinctive characters
that would warrant the erection of a special family for
it (which has been done). It is appended below to Pteridaceae, not because of any suspicion of affinity but
only because in the general key it will run to that family.
In recognizing a fairly limited number of families in
the leptosporangiate ferns, 31 as compared to Pichi
Serrnolli's 51, I have tried, with all other authors who
have contributed to the present volume, to act in the
spirit of the principles outlined above (Kramer and
R. Tryon, this VoL).
Filicatae . Aspleniaceae
52
References (apart from those included in "General
References to Pteridophytes" [p.14/15]).
Ching, R. C. 1978. The Chinese fern families and genera:
systematic arrangement and historical origin. Acta Phytotax. Sin. 16: 1-37.
Crabbe, J. A., Jermy, A. c., Mickel, J. T. 1975. A new generic
sequence for the pteridophyte herbarium. Fern Gaz. 11
(2/3): 141-162.
Diels, L. 1899/1900. Filicales. In: Engler, A., Prantl, K.. (Eds)
Die natiirlichen Pflanzenfamilien 14. Leipzig. W. Engelmann.
Holttum, R. E. 1947. A revised classification of leptosporangiate ferns. J. Unn. Soc. Bot. 53: 123-158.
Holttum, R. E. 1949. The classification of ferns. BioI. Rev. 24:
267-296.
Holttum, R. E. 1%9. Studies in the family Thelypteridaceae,
etc. Blumea 17: 5-32.
Holttum, R. E. 1981. Thelypteridaceae. Flora Maleso. 11.1,5:
331-599.
Holttum, R. E., Sen, U. 1961. Morphology and classification
of the tree ferns. Phytomorphology 11: 406-420.
Jarrett, F. M. 1980. Studies in the classification of the leptosporangiate ferns: I. The affinities of the Polypodiaceae
sensu stricto and the Grammitidaceae. Kew Bull. 34:
825-833.
Mickel, J. T. 1974. Phyletic lines in the modern ferns. Ann.
Missouri Bot. Gard. 61: 474-482.
Nayar, B. K. 1970. A phylogenetic classification of the homosporous ferns. Taxon 19: 229-236.
Pichi Sermolli, R. E. G. 1977. Tentamen Pteridophytorum
genera in taxonomicum ordinem redigendi. Webbia 31 :
313-512.
.
Aspleniaceae
K. U. KRAMER and R. VIANE
Aspleniaceae Frank in Leunis, Syn. Pflanzenkd., ed. 2, 3:
1465 (1877).
Terrestrial, epilithic, or epiphytic ferns of small to medium, rarely to large size; stem short and (sub)erect or
sometimes long and creeping, dictyostelic, with welldeveloped cortical sclerenchyma, bearing non-peltate,
clathrate, entire, marginally fimbriate, or glandular,
rarely ciliate, dorsally glabrous scales, these sometimes
iridescent. Leaves clustered to remote; petiole non-articulate, almost wanting to well-developed, usually
smooth when mature, usually with paleae at the base,
often hairy and glandular; often dark and sclerotic
and then glabrous, adaxially usually flattened to sulcate, typically with two C-shaped to elliptic vascular
bundles back to back joining above the base to near
the apex of the petiole to form one X-shaped bundle
(Fig. 18 F). Lamina very variable, simple or lobed,
more often once or twice pinnate, occasionally more
strongly dissected, to quadripinnate (+ pinnatifid),
anadromous, often truncate at base, thinly membranous to coriaceous, rarely somewhat dimorphic; surface rarely completely glabrous, often with minute uniseriate (glandular) hairs and slightly scaly; rachis like
the petiole but with smaller scales, often also darksclerotic; proliferous buds often present on the rachis
(on pinnae and sometimes on the petiole), usually
borne singly. Pinnae (if any) usually shortly petiolulate
to sessile, non-articulate, simple to variously dissected;
upper pinnae, pinnules, etc. usually gradually reduced
and confluent; a conform terminal pinna only present
in some simply pinnate species. Ultimate free divisions
of very variable size and shape, usually sessile with
rounded to cuneate base, rarely decurrent, never surcurrent, sometimes pseudodimidiate or dimidiate,
usually costate; edge rarely entire, often crenate-serrate, to pinnatifid, the lobes sometimes bifid. Rachises
of secondary order (and costae) adaxially grooved, the
groove then like that of the primary pinnae, or often
convex in the middle, continuous with that on rachises
of different order or rarely not; costae adaxially flat to
convex or sulcate. Ultimate lobes not rarely narrow
and uninerval. Axes scantily scaly, inconspicuously
hairy or glandular. Veins anadromous, rarely isodromous (sometimes basiscopic veins absent), free and
pinnate or forked, rarely sparingly to fully reticulate or
loop-connected at their ends; free included veinlets always absent. Vein-ends thickened behind the margin;
margin occasionally sclerotic. Sori single, dorsal on the
veins, often only on an anterior vein branch but sometimes also on the posterior branch of an adjacent veinlet or alternately on anterior and posterior branches
and then approximate in pairs. Indusium attached to
the side of the vein, usually long and narrow and with
free, tapering ends; occasionally the sori very short
and submarginal and the indusium then at its lateral
ends often fused with the lamina to form a pouch-like
structure ("Loxoscaphe") (Fig. 19B); when the sori face
each other in pairs, the indusia open against each other and the sori are then seemingly confluent at full
maturity ("Phyllitis", "Diplora''); rarely the sori following the vein network and here and there back-to-back
or branched ("Camptosorus',), or exindusiate and
linear ("Ceterach ". "Pleurosorus''). Sporangia longstalked, most of the stalk uniseriate, eglandular; annulus vertical, interrupted by the stalk; stomium well-developed, of several elongate cells, passing into the
bow, typical of the family. Spores monolete, usually elIipsoid; perispore prominent, consisting of a thin basal
layer (inner perispore) pressed against the exospore,
this inner perispore bearing rod-like (sometimes forking) structures (middle perispore) sustaining the outer
perisporal layer. Middle and outer variously developed, the outer layer often (partly) perforated and/or
thrown into variously shaped ridges or into spines.
53
Filicatae . Asplcniaceae
B
D
F
RARE OCCURRENCE. Stem creeping,
with many hair-like scales and few paleae, in some
very small species like. A. delicatulum C. Pres!. Stem
with buds in A. protensum Schrader; petiole base modilied into a starch-storage tissue: trophopod (Wagner
lind Johnson 1983), in A. platyneuron (L.) Oakes,
A. adiantum-nigrum L. and relatives. Multiseptate
hairs on the leaves in A. pumilum Swartz, A. protensum
CI·IARACTERS OF
Fig. 17 A-F. Aspleniaceae, Blechnaceae, and Cyatheaceae,
spores (an x 1000). A Asplenium falcinellum. B Asplenium
(.. Loxoscaphe·' theciferum. C Woodwardia fimbriata. D Grathea caracasana. E Cyathea cooperi. F Cyathea (Cnemidaria)
horrida. Pho!. A. F. Tryon
54
Filicatae . Aspleniaceae
B
Fig.18A-I. Aspleniaceae. A,B Asplenium tripteropus. A Pinna with son (x 4). B Part of rachis ( x 20). C Asplenium septentrionale. transection of fertile segment (x 25). D,E Asplenium {..Diellia erecta·~. D Fertile pinna (x 3). E Matgin with
SOn (x 8). F Asplenium sp., vascular bundles of petiole, at
base (above) and higher (below) (schematic). G Asplenium nidus, sporangium (x 140). H Asplenium septentrionale, scale
of stem (x 10). I Asplenium car<iiophyllum, scale of stem
(x 20) (A-C, H from Rora Tsinlingensis 1974; D,E Perez
Arbehiez 1928; F from Ogura 1972; G from Haider 1954;
I from Kurata and Nakaike 1981).
Schrader, etc. Amply reticulate veins in, e. g., A. eeleraeh L., A. rhizophyllum L., "Camptosorus sibiricus
Rupr.", A. purdieanum Hooker, A. ghiesbreghlii Fournier, "Diellia laeiniata (Hillebr.) Diels", and their relatives; the character occurs here and there in the genus.
Some slight dimorphism of sterile and fertile leaves in
a few species, e. g., A. platyneuron (L.) Oakes and
A. dimorphum Kunze. Abaxially densely scaly leaves
in A. aethiopicum (Burm.) Bech~rer, A. ceteraeh L. and
relatives. Indusia of neighbouring sori facing each other with their free edges (and often confluent sori) in
A. seolopendrium L. and other species referred to
"Phyllitis", "Antigramma", "Diplora", etc. Partly
"double" sori, back-to-back on one vein ("diplazioid")
in a few species, e. g., A. melanopus Sodiro.
Indusia facing the margin with their free edges and
on veins near and parallel to the margin in "Diellia"
(Fig. 18 D,E). J-shaped indusia in A. onopteris L., etc.
Buds on the prolonged, apically lamina-less rachis in
A. mannii Hooker, A. bipinnatifidum Bak., A.fogax
Christ, and in some neotropical taxa (A. alatum
Humb. and Bonpl. ex Willd., A. radieans L., etc). Linear ultimate divisions in A. septentrionale (L.) Hoffm.,
A. novae-caledoniae Hooker, etc. Rabellate, ecostate
pinnae in some (African) taxa: A.laurentii Bommer ex
Christ, A.jaundeense Hieron., A. megalura Hieron. ex
Brause. Non-dehiscent sporangia in A. lepidum
C. Presl (Brownsey 1977 c). Absence of the outer perisporiallayer in A. seelosii Leybold.
MORPHOLOGY AND ANATOMY. The family as far as investigated is quite homogeneous (Fig. 17). Certain variations are to be found in the rachis structure: concave,
sulcate, or the groove with raised middle; adaxially
grooved vs. non-grooved or raised costae; reticulate or
semi-reticulate vs. free veins; etc. Leaves without lamina that root at the apex and produce new plantlets,
formerly misinterpreted as stolons, occur in A. bipinnatifidum Baker and A. mannii Hooker (Fig. 19 A). As
in Lindsaea, hardly any morphological or anatomical
adaptations to epiphytism can be observed in epiphytic species, except in the so-called nest ferns (Section
Thamnopleris Presl) (Holttum 1974) and a few other,
unrelated species like A. serrafurn L. and A. a/rieanum
Desv. where they are weakly developed.
The stomata are principally polocytic, with staurocytic and paracytic ones occasionally present besides.
The most extensive studies of the stem anatomy are
by Mitsuta et al. (1980) and by Chandra and Nayar
(1975). The rhizome is dictyostelic, radial in species
with erect, dorsiventral in most of the (rather few) species with creepinK stem, e. g., Sect. Hymenasplenium
(Iwatsuki and Kato, 1975); see also Tardieu-Blot
(1932).
The leaf trace and petiolar bundle are single in a
few small, obviously reduced species.
Asplenium
Laminal parts of about two cell layers with few or
without any stomata have been reported for a variety
of A. unilaterale Lam. (Iwatsuki 1975).
Little comprehensive work is available; for a review, see Atkinson (1973). The prothallia
ure of the common cordate or elongate-cordate type;
in some species they are naked, in others they bear papillae and/or uni- to multicellular, usually chlorophyllous hairs. The few that are known in "satellite genera" are not divergent (Wagner 1951; Atkinson 1973).
GAMETOPHYTE.
Nothing definite is known about the affinity
of the Aspleniaceae. Various group have been thought
of as potential relatives, like Thelypteridaceae, Blechnuceae, and Athyriaceae, but the similarities are not
numerous and may be due to convergence. The former
confusion of Asplenium with Diplazium (and Athyr111m) is well known; for a good summary of differenliating characters, see Smith (1976). Any idea about aflinity between Diplazium and Asplenium has now been
dropped. However, perispore structures closely similar
(0 some encountered in Asplenium are found in ThelYflteris (sect. Stegnogramma) and in Polystichum.
AFFINITY.
The more than 700 species of Asplenium are distributed allover the world,
some species even extending into Arctic and arid
IITeas. The majority are in the (sub)humid tropics and
south-temperate regions. Many occur in tropical-mon(line vegetations. Here they grow on the forest floor,
on banks, and in ravines, often on rocks, less often as
high epiphytes; a few species are virtually scan dent as
(hey ascend trees with their rooting-proliferating rachis
(lIJOGRAPHY AND EcoWGY.
Fig. 19A-F. Aspleniaceae. A,B Asplenium r..Loxoscaphe")
mannii. A Entire plant, with "stolons" ( x 0.6). B Fertile segment (x 5). C Asplenium rutifolium. fertile segment (x 5).
D, E Asplenium laurentii. D Habit (x 0.3). E Pinna (x 1).
F Asplenium (,. Ceterach'~ cordatum. leaf (x 0.6) (A-C, F from
Schelpe 1970; D,E from Tardieu-Blot 1964).
tips (e. g., A. sandersonii Hooker). A few species are
obligate tree-fern epiphytes (A. hypomelas Kuhn). The
concentration of species with proliferous buds in Africa is peculiar. The "nest-fern" epiphytes have been
mentioned; they are humus collectors. A very few species are known to be rheophytes (e.g., A. subaquatile
Cesati).
Asplenium is a cosmopolitan genus, and one of the
very few that is represented about equally in all tropical areas [c. 30% in the neotropics, c.22% in Africa,
c.33% in Asia, c. 10% in the Pacific (incl. Australia,
etc.), c.5% in Europe s.l.]. Few species are (sub)cosmopolitan (e. g., A. adiantum-nigrum L., A. trichomanes L.). Islands often harbour endemic species, like
Madagascar (c. 40 species), La Reunion, Hawaii, Juan
Fernilndez, Tristan da Cunha, the Azores, and New
Zealand. Rather few (sub)tropical species occur on
more than one continent, e. g., A. monanthes L., A. normale D. Don. Some species have a very restricted distribution (palaeo- or neo-endemics), e. g., A. jahandiezii (Litard.) Rouy, A. bourgaei Milde, A. balearicum
Shivas, etc. Different centres of (secondary) diversification seem to exist, e. g., North America (Appalachians), the mountains of Central America, the Andes,
the Himalaya, etc.
56
Filicatae • Aspleniaceae
CYTOLOGY AND HYBRIDIZATION. Few families are cytologically as homogeneous as this. An x number of 36,
with diploid to 16-ploid levels based on it (Lovis 1973),
has been recorded in all but a handful of species, and
in nearly all of the (formerly recognized) "satellite genera"; this, together with the lack of sterility barriers, is
very important evidence in favour of not recognizing
these segregates as genera; (see, e.g., Bir et al. 1985).
The exceptions are A. ("Boniniella") cardiophyllum
(Hance) Baker: 2n=76; and A. unilaterale Lam.,
n =40 (Manton and Sledge 1954); a new number of
2n = 112, resulting form hybridization of the latter
with an n =36 species (36 11 +401; Bir 1960,1963), has
been recorded in a few populations. The literature on
hybridization within Asplenium, often involving "satellite genera", with both naturally occurring and artificially produced hybrids, is enormous; see e. g., Meyer
(1952), Wagner (1954), Lovis (1973), Brownsey
(1977b), Reichstein (1981), Werth et al. (1985). Half of
all known temperate taxa are hybridogenous allopolyploids behaving like species. Intraspecific hybrids are
also known. Apogamous taxa (hybridogenous or not)
have not been reported in great number; see, e. g.,
Manton (1950), Manton and Sledge (1954), Mehra and
Bir (1960), Braithwaite (1964), Murakami and Iwatsuki
(1983). Hybridization and subsequent polyploidy often
involve morphologically quite diverse species, resulting in hybrids or species with peculiar intermediate
and/or fluctuating morphology; see, e. g., Wagner
(1954), Meyer (1965), Walter et al. (1982); also Fig. 19.
A single genus:
Characters as for family.
C. 720 species, subcosmopolitan; with growing knowledge from biosystematic sources the number of species
is likely to increase as species complexes are unravelled. SEM work on spores is also yielding important
new data as the genus is very diverse in this respect
(see, e. g., Viane 1977).
As to the recognition of natural infrageneric groups,
whether subgenera or sections, Asplenium is still extremely poorly known. Some of the "satellite genera"
are good candidates for natural sect ion s. Other
natural infrageneric groups are sections Thamnopteris
and Hymenasplenium, the group of A. aethiopicum
(Burm.) Becherer, the A. myriophyllum-abyssinicum
group, the A. laetum-erectum group, the A. trichomanes-monanthes group, etc. Other species groups recognized are often based on superficial resemblance,
and may be convenient but are artificial, like "Darea",
"Loxoscaphe", etc. The species groups recognized by
Mettenius (1859) and Diels (1899) are almost wholly
unnatural. The great majority of modem authors have
refrained from proposing a natural subdivision of the
genus, and we may still be far removed from such a
goal. The great majority of the satellite genera are
based on single, easily observed characters of little taxonomic weight. Their source in Asplenium (i. e., their
nearest non-deviating relatives) can often be pinpointed.
Selected Bibliography
A.pleniumL
Fig. 17-19
Asplenium L. Spec. Plant.2: 1078 (1753); Maxon (1913);
Looser (1944); Sehnem (1963): Sledge (1965); Morton and
Lellinger (1966); Brownsey (1977 a); Nakail<e (1986).
Phyllitis Hill (1756).
Scolopendrium Adans. (1763).
Ceterach Willd. (1804), nom. cons.
Camptosorus Unk (1833).
Antigramma C. Presl (1836).
Neot/opteris J. Smith (1841).
Pleurosorus Fee (1852).
Diellia Brackenridge (1854); Wagner (1951).
Loxoscaphe Moore (1853).
Schajftleria Fee (1857).
Diplora Baker (1873).
Asplenidictyum (Hooker) J. Smith (1875).
Holodictyum Maxon (1908).
Biropteris Kiimmerle (1922).
Boniniella Hayata (1927).
Hymenaspknium Hayata (1927).
Ceterachopsis (1. Smith) Ching (1940).
X Asp/enophyllitis Alston (1940).
X Asplenosorus Wherry 19(37).
X Asplenoceterach D. E. Meyer (1957).
Sinephropteris Mickel (1976).
x Ceterophyllitis Pic. Ser. 1979.
x Phyllitopsis Reichstein (1981).
Bir, S. S. 1960. Cytological observations on the East Himalayan members of Asplenium Unn. Curro Sci. 29: 445-447.
Bir, S. S. 1963. Evolution in the Indian members of the genus
Asplenium Linn. Mem. Indian Bot. Soc. 4: 41-50.
Bir, S. S., Fraser-Jenkins, C. R., Lovis, J. D. 1985. Asplenium
punjabense sp. nov. and its significance for the status of
Ceterach and Ceterachopsis. Fern Gaz. 13: 53-63.
Braithwaite, A. F. 1964. A new type of apogamy in ferns.
New Phytologist 63: 293-305.
Brownsey, P.J. 1977a. A taxonomic revision of the New Zealand species of Asplenium. New ZeaL J. Bot. 15: 39-86.
Brownsey, P. J. 1977b. Asplenium hybrids in the New Zealand
flora. New ZeaL J. Bot. 15: 601-637.
Brownsey, P. J. 1977 Co An example of sporangial indehiscence in the Filicopsida. Evolution 31: 294-301.
Chandra, S., Nayar, B. K 1975. Vascular organization in the
rhizome of spleenworts. J. Indian Bot. Soc. 54: 187-199.
Faden, R. B. 1973. Some notes on the gemmiferous species of
Asplenium in tropical East Mrica. Am. Fern J. 63: 85-90.
Holttum, R. E. 1974. Asplenium Unn., sect. Thamnopteris
PresL Gard. BulL Singapore 27: 143-154.
Iwatsuki, K 1975. Taxonomic studies of pteridophyta X.
13. Asplenium sect. Hymenasplenium. Acta Phytotax. Geobot. 27: 39-55.
Iwatsuki, K, Kato, M. 1975. Stelar structure of Asplenium
unilaterale and allied species. Kalikasan 4: 165-174.
Looser, G. 1944. Sinopsis de los ''Asplenium'' (Filicales) de
Chile. Ulloa 10: 233-264.
Filieatae . Azollaceae
Lovis, J. D. 1973. A biosystematic approach to phylogenetic
problems and its application to the Aspleniaceae. Bot. J.
Linn. Soc. 67 Suppl. 1: 211-228.
Manton, I., Sledge, W. A. 1954. Observations on the cytology
and taxonomy of the pteridophyte flora of Ceylon. Philos.
Trans. R. Soc. London Ser. B 238: 127-185.
Maxon, W. R. 1913. Asplenium trichomanes and its American
allies. Contrib. U. S. Nat. Herb. 17(2): 134-153.
Mehra, P. N., Bir, S. S. 1960. Cytological observations on Asplenium cheilosorum Kunze. Cytologia 25: 17-27.
Mettenius, G. 1859. Dber einige Famgattungen. VI. Asplenium. Abh. Senckenberg Naturf. Ges.3: 110-254.
Meyer, D. E. 1952. Untersuchungen liber Bastardierung in
der Gattung Asplenium. Bibl. Bot. (Stuttgart) 123: 1-34.
Meyer, D. E. 1965. Zum morphologischen Prinzip der Irregu·
larita! von Artbastarden und Bastardarten, vom Blickpunkt
der Systematik. Willdenowia 4: 63-73.
Mitsuta, S., Kato, M., Iwatsuki, K. 1980. Stelar structure of
Aspleniaceae. Bot. Mag. (Tokyo) 93: 275-289.
Morton, C. V., Lellinger, D. B. 1966. The Polypodiaceae subfamily Asplenioideae in Venezuela. Mem. New York Bot.
Gard. 15: 1-49.
Murakami, N., Iwatsuki, K. 1983. Observation on the variation of Asplenium unilaterale in Japan with special reference to apogamy. J. Jpn. Bot. 58: 257-262.
Nakaike, T. 1986. An enumeration of the ferns of Nepal II.
Asplenium L. Bull. Nat. Sci. Mus. Ser. B 12: 37-54.
]>jchi Sermolli, R. E. G. 1979. [Ceterophyllitis}. Webbia 34:
208.
Reichstein, T. 1981. Hybrids in European Aspleniaceae (Pteridophyta). Bot. Helv. 91: 89-139.
Sehnem, A. 1963. 0 genero Asplenium nos Estados Santa Catarina e Rio Grande do SuI. Sellowia 15: 9-37.
Sledge, W. A. 1965. The Ceylon species of Asplenium. Bull.
Brit. Mus. (Nat. Hist.) Bot. 3(6): 235-277.
Smith, A. R. 1976. Diplazium delitescens and the neotropical
species of Asplenium sect. Hymenaspienium. Amer. Fern
J. 66: 116-120.
Tardieu-Blot, M. 1932. Les Aspleniees du Tonkin. Toulouse.
Viane, R. 1977. Spore morphology and stomatal characters in
some Kenyan Asplenium-species. Ber. Dtsch. Bot. Ges. 90:
219-239.
Waguer, W. H. Jr. 1951. The fern genus Diellia. Its structure,
affinities and taxonomy. Univ. Cal. Publ. Bot. 26(1): 1-212.
Waguer, W. H. Jr. 1953. An Asplenium prototype of the genus
Diellia. Bull. Torrey Bot. Club. 80: 76-94.
Waguer, W. H. Jr., 1954. Reticulate evolution in the Appalachian Aspleniums. Evolution 8: 103-118.
Wagner, W. H. Jr., Johnson, D. M. 1983. Trophopod, a commonly overlooked storage structure of potential systematic
value in ferns. Taxon 32: 268-269.
Walter, K. S., Waguer, W. H., Wagner, F. S 1982. Ecological,
biosystematic and nomenclatural notes on Scott's spleenwort, x Asplenosorus ebenoides. Amer. Fern J. 72: 65-75.
Werth, Ch. R., Guttmann, Sh. I., Eshbaugh, W. H. 1985. Electrophoretic evidence of reticulate evolution in the Appalachian Asplenium complex. Syst. Bot. 10: 184-192.
57
Azollaceae
J. J. SCHNELLER
Azollaceae Wettst., Handb. Syst. Bot. 2: 77 (1903).
Free floating aquatic plants with copiously branched,
delicate, horizontal, protostelic stems. Roots simple,
on the ventral side of the axis, chlorophyllous, solitary
or in clusters, extending a short distance downward in
the water, deciduous. Leaves sessile, alternate, in two
rows in the dorsal side of the stem; each leaf divided
into two lobes, a (usually somewhat smaller) upper, aerial, and a lower floating lobe. The upper, aerial lobe
photosynthetic, with epidermis and mesophyll, bearing
on the lower surface a large cavity with mucilage and
usually living colonies of the blue-green alga Anabaena; ihe lower lobe floating with the dorsal side only
immersed, unistratose except at the base, mostly without chlorophyll. Sporocarps borne on the first leaf of a
lateral branch, the lower lobe of this fertile leaf reduced to two, sometimes up to four sporocarps, the
upper lobe forming an involucre covering the sporocarps. Sporocarps containing either micro- or megasporangia; megasporocarps smaller. Sporangia without dehiscence mechanism. Microsporocarps containing numerous microsporangia; microsporangia forming 64 [according to Svenson (1944) 32 or 64) trilete
spores; clusters of spores embedded in 3-10 massulae
(massula: hardened tapetal mucilage containing
groups of spores); surface of the massulae mostly with
anchor-shaped outgrowths, called glochidia, except in
A. nilotica. Megasporocarp containing only one megasporangium; Megasporangium with 8 spore-mother
cells; after meiosis only one spore developing, the others aborting. Mature megaspore on the proximal pole
with a columella, hair-like filaments, and with apical
massulae (called floats). Megaspores plain, perforate,
with sparse excrescences, or granulate, with dense excrescences, or densely filamentose, with fine or coarse
perforations, or compact filaments forming a reticulum over coarse perforations, or rugulate, with mostly
obscure perforations; microspores rugulose, germinating within the massula; microprothalJium reduced,
with one antheridium. MegaprothalJium developing
within megaspore, the growing cell cushion then forcing the megaspore open at the laesura, forming a (in
transversal section) round to triradiate prothalJium
with several archegonia on the upper surface. C. 6 species, in the New World from Argentina to Alaska, in
the Old World in Africa, E. Asia and Australia;
introduced into Europe.
ANATOMY AND MORPHOLOGY. The most important
contributions are by Bonnet (1957), Demalsy (1958),
Filicatae . Azollaceae
58
Eames (1936), Konar and Kapoor (1972, 1974), Stras·
burger (1873), and Warmbrodt and Evert (1978).
Warmbrodt and Evert (1978) described the detailed
structure of the leaf. The lower lobe is achlorophyllous
except for several layers of photosynthetic cells sur·
rounding the single vascular bundle. The aerial lobe
consists of upper and lower epidermis with stomata,
mesophyll tissue differentiated into spongy and palisade parenchyma, and a single vascular bundle. The
upper epidermis is covered by numerous water-repellent trichomes (Pieterse et al. 1977). The palisade parenchyma consists of a single layer of cells, the spongy
parenchyma of one or more layers. Within the spongy
mesophyll a single large cavity is differentiated, which
is connected to the outside through a small pore in the
lower surface of the lobe. This cavity contains a muci-
D
Fig.20A-E. Azollaceae. Azolla microphyUa. A Habit (x 16).
B Leaf with dorsal and ventral lobe (x 45). C Massula with
glochidia (x 120). D Glochidion with septa (x 240). E Me·
gaspore in lateral view (x4O)( A,B Eames 1936; C-E Sven·
son 1944)
laginous substance, colonies of the blue-green alga
Anabaena azollae, and numerous multicellular hairs.
The Anabaena-Azolla symbiosis is well studied (Duckett et al. 1975; Peters et al. 1978; Holst and Yopp
1979).
Single roots, or in A. nilotica groups of roots, develop at the base of the branches; the root is simple,
3-4 cm long, chlorophyllous, with or without root
hairs. The sporocarps originate on the lower leaf lobe
normally in pairs (sometimes in fours); two of a pair
may be of the same or the opposite sex. According to
Duncan (1940), the sporocarp initial forms a columella
on whose base in the indusium initials soon differentiate. The indusium develops into a two-layered sporocarp wall. The mature sporocarp is then surrounded
by a hood-like structure called involucrum, formed by
the upper lobe of the fertile leaf (Fig. 20). The apical
cell of the columella always becomes the apical cell of
the developing megasporangium, so never more than
one megasporangium is formed. While meiosis is taking place in the megasporangium, the columella elongates and some of its epidermal cells become microsporangia initials. If one megaspore in the megasporangium persists, the further development of the microsporangia is suppressed and a megasporocarp develops. If all megaspores disintegrate, the megasporangium collapses, and numerous microsporangia develop, and a microsporocarp is formed. The sporangia of
both types develop in the leptosporangiate manner but
no annulus is found. Micro- and megasporogenesis
were studied in detail by Duncan (1940) and Bonnet
(1957). The massula, and certain characters of the
megaspore, are highly distinctive structures peculiar to
Azolla. The occurrence and the shape of glochidia
(Svenson 1944), the number of floats, and the morphology (sporoderm architecture) of the megaspore
(Fowler and Stennet-Wilson 1978) are important characters for separating taxa.
GAMETOPHYTE. According to Bonnet (1957), the male
prothallium of A. fiJiculoides contains one rhizoid cell,
one protonemal cell, and the cells forming a single antheridium. The microprothalli develop within the massulae. The glochidia become erect when the massulae
are free. According to Fowler (1975), the microspores
of A. caroliniana, A.filiculoides, and fossil Azolla species occupy one of the pseudocellular cavities within
the massula. Each microspore-containing cavity is
closely associated with a funnel-shaped cavity, the
neck of which extends to the periphery of the massula
and opens by a pore to the exterior. The spermatozoids probably escape from the massula··through this
pore. The first stages of the megaprothallus remain
within the megaspore. The mature megaprothallium
breaks out of the megaspore at the laesura. If the first
archegonium is not fertilized, the cushion-like tissue
59
Azalia
develops further archegonia. The external aspect of
the megaspore does not change until fertilization has
taken place. According to Campbell (1893), Konar and
Kapoor (1972), and Lucas and Duckett (1980), both
micro- and megasporocarps remain floating until the
indusial wall disintegrates. The mega- and microsporangia then complete their development at the bottom
of the water bodies on which the parent plants were
growing. Rao (1935) stated in contrast that the megasporocarps float on the surface soon after the archegonia are formed in the megaprothallium. Considering
the possible function of the very distinctive structures
like massulae with glochidia, megaspores with fibers,
columella and "floats", and special grooves in the
spore wall, one is tempted to assume that the massula
(containing micros pores and microprothalli) and the
megaspores (with the female gametophyte) float on
the surface of the water and do not lie on the bottom.
Further investigation on the biology of the haploid
generation is necessary.
BLASTOGENY. According to Campbell (1893) and Konar and Kapoor (1974), the young embryo differentiates
into root, foot, first leaf, and stem. The first leaf is simple and without a cavity and differs in shape and anatomy from the following leaves, which possess a cavity
and develop gradually into two-lobed leaves.
ECOLOGY AND DISTRIBUTION. Species of Azolla are
free-floating and occur on the surface of ponds or in
sloughs of rivers, in warm-temperate or tropical climates. They propagate very successfully by vegetative
means. The genus is of worldwide distribution. It was
introduced to some areas, for example Europe, by
man. Two main distribution centres can be distinguished: one in East Asia and Africa, the other in
North and South America. The Old World species
were originally lacking in Europe and W. Asia.
RELATIONSHIPS. There are similarities to Salvinia, but
since these are not very close, it does not seem indicated to unite the two genera into one family but to distinguish a family Azollaceae, and to include Salviniaceae and Azollaceae in one order, the Salviniales. The
phylogenetic derivation of Azolla remains unknown.
Fossil remains were found in the Cretaceous (Hall
1969; Sweet and Hills 1976). Some affinity to the Hymenophyllaceae has been considered (Eames 1936;
Takhtajan 1953), but most of the anatomical and morphological features are very different from those of
that family.
KARYOLOGY AND HYBRIDIZATION. Only the chromonumbers of A. pinnata, n ~22 (Loyal 1958),
]n ~44 (Loyal et al. 1982) and A. caroliniana, 2n ~48
(Thchermak-Woess and Dolezal-Janisch 1959) are
known. So far no hybrids have been found.
A single genus:
~ome
Azolla Lamarck
Fig. 20
Azalia Lamarck, Encycl. Meth. Bot. 1: 343 (1783); Reed
(1953, 1965).
Characters as for family.
Selected Bibliography
Bonnet, A. L M. 1957. Contribution Ii retude des Hydropteridees. III. Recherches sur Azallajiliculaides Lamk. Rev. Cytol. BioI. Veg. 18: 1-88.
Campbell, D. H. 1893. On the development of Azalia jiliculaides Lam. Ann. Bot. (London) 7: 155-187.
Demalsy, P. 1958. Nouvelles recherches sur Ie sporophyte
d'Azalla. Cellule 59: 235-268.
Duckett, J. G., Toth, R., Soni, S. L 1975. An ultrastructural
study of the Azalia-Anabaena azallae relationship. New
Phytol. 75: 111-118.
Duncan, R. F. 1940. The cytology of sporangium development in Azalia jiliculaides. Bull. Torrey Bot. Club 67:
391-412.
Eames, A. J 1936. Morphology of vascular plants, lower
groups. London: McGraw-Hill.
Follieri, M. 1977. Classification and phylogeny of living and
fossil water ferns of the genus ''Azolla''. Webbia 31:
97-104.
Fowler, K. 1975. An escape mechanism for spermatozoids in
Azalia massulae. Amer. Fern J. 65: 7-10.
Fowler, K., Stennet·Willson, J. 1978. Sporoderm architecture
in modem Azolla. Fern Gaz. 11: 405-412.
Hall, J. W. 1969. Studies on fossil Azalia: primitive types of
megaspores and massulae from the Cretaceous. Amer. J.
Bot. 56: 1173-1180.
Holst, R. W., Yopp, J. H. 1979. Studies of the Azalia-Anabaena symbiosis using Azalia mexicana. 1. Growth in nature
and laboratory. Amer. Fern J. 69: 17-25.
Konar, R. N., Kapoor, R. K. 1972. Anatomical studies on
Azallapinnata. Phytomorphology 22: 211-223.
Konar, R. N., Kapoor, R. K. 1974. Embryology of Azalia pinnata. Phytomorphology 24: 228-261.
Loyal, D. S. 1958. Cytology of two species of Salviniaceae.
Curro Sci. 27: 357-358.
Loyal, D. S., Gollen, A. K., Ratra, R. 1982. Morphological
and cytotaxonomic observations on Azalia pinnata. Fern
Gaz. 12: 230-232.
Lucas, R. C., Duckett, J. G. 1980. A cytological study of the
male and female sporocarps of the heterosporous fern
Azallajiliculoides Lam. New Phytol. 85: 409-418.
Peters, G. A., Toia, R. F., Raveed, D., Levine, N. J. 1978. The
Azalia - Anabaena relationship. VI. Morphological aspects
of the association. New Phytol. 80: 583-593.
Pieterse, A. H., de Lange, L., van Vliet, J. P. 1977. A com·
parative study of Azalia in the Netherlands. Acta Bot.
Neerl. 26: 433-449.
Rao, H. S. 1935. The structure and life history of Azalia pinnata R. Brown with remarks on the fossil history of the Hydropterideae. Proc. Indian Acad. Sci. 2B: 175-200.
Reed, C. F. 1954. Index Marsileata et Salviniata. Bol. Soc.
Brot. II, 28: 5-61.
Reed, C. F. 1965. Index Marsileata et Salviniata. Suppl. Bol.
Soc. Brot. II, 39: 259-302.
Strasburger, E. 1873. Ober Azolla. Jena: Gustav FIScher.
60
Filicatae . Blechnaceae
Svenson, H. K. 1944. The New World species of Azalia. Arner. Fern J. 34: 69-84.
Sweet, A. R., Hills, L. V. 1976. Early tertiary species of Azalia
subg. Azalia sect. Kremalaspora from western and arctic
Canada. Can. J. Bot. 54: 334-351.
Takhtajan, A. L. 1953. Phylogenetic principles of the system
of higher plants. Bot.·Rev.19: 1-45.
Tschermak-Woess, E., Dolezal-Janisch, R. 1959. Ober die
karyologische Anatomie einiger Pteridophyten sowie auffallende Unterschiede im Kernvolumen bei Cyrtomium falcalum. bsterr. Bot. Z. 106: 315-324.
Warmbrodt, R. D., Evert, R. F. 1978. Comparative leaf structure of six species of heterosporous ferns. Bot. Gaz.139:
393-429.
Blechnaceae
K. V. KRAMER with the collaboration of
T. C. CHAMBERS and E. HENNIPMAN
Blechnaceae (c. Presl) Copeland, Gen. Fi!.: 155 (1947).
Blechneae C. Presl, Epimel. Bot.: 103 (1851) ["Blechnaceae"].
Stenochlaenaceae Ching, Acta Phytotax. Sin. 16 (4): 18
(1978).
Terrestrial or epilithic, occasionally scandent, rarely
epiphytic ferns of small to large size. Stem creeping,
erect, or scandent, radially dictyostelic, scaly. Petioles
aggregated to remote, usually, well-developed, adaxialIy grooved, with several vascular bundles usually arranged in a V-pattern. Lamina mostly pinnatifid to
simply pinnate with entire to pinnatifid pinnae, less
frequently simple or lobed, rarely bipinnate. Sterilefertile leaf dimorphism very widespread and developed to various degrees but by no means universal. Veins
free or anastomosing, in the latter case (with one exception) without free included veinlets; dissection pattern and/or venation catadromous, at least in the
upper part of the lamina. Fertile pinnae/segments
with a least one row of costal areoles, their outer arch
mostly forming the receptacle of the sori or of one
long, continuous sorus, or a special soral vascularization present. Rachis adaxially sulcate, the groove not
continuous with those on the pinna costae. Sori short
to long and narrow, with few exceptions having an indusium that is attached along the receptacle and free
towards the costa or costule (this character rarely
found in other ferns); occasionally the indusium wanting, the sporangia spreading along the veins or acrostichoid in arrangement, no sterile appendages borne
among the sporangia. Sporangia with strong, triseriate
stalk; annulus many-celled, the indurated part extending to the stalk, the several-celled stomium weakly differentiated from the many-celled, non-indurated part
of the bow. Spores monolete.
ANATOMY AND MORPHOLOGY. Few representatives
have so far been studied comprehensively, particularly
as to the anatomy of the vegetative organs, the receptacle having received the most attention; e. g., de la Sota
and Gouvea Labouriau (1961). Stenochlaena is better
known, and its taxonomic position in the present family has partly been established on anatomical grounds
(e. g., Mehra and Chopra 1951). The stem is radially
dictyostelic, with well-developed, sometimes sclerotic
pith. The vascular bundles of the petiole are arranged
in a V-pattern, as seen in transection, with the two
adaxial bundles larger and the xylem hippocampiform
(Fig. 25 F), the number of other bundles, of which
there are at least two, dependent upon the size of the
leaf (see, e. g., Tardieu-Blot 1932). The stomata are polocytic with anomocytic ones sometimes also present.
In this character Stenochlaena also agrees with the other genera, but its vascular anatomy is more complicated, probably in connection with its growth habit; see
under the genus. Young, unfolding leaves are very often tinged with red, a character possibly of taxonomic
significance (Tryon and Tryon 1982). Sporangia
spreading onto the indusium have been found in
Blechnum and Salpichlaena, a very rare feature.
GAMETOPHYTE. See especially Stokey and Atkinson
(1952a, b), Stone (1962), Atkinson (1973), HolbrookWalker and Lloyd (1973), and Nayar and Kaur (1971).
The prothallia are cordate, or elongate when mature,
with a distinct, often firm midrib; they often bear simple or few-celled, partly glandular, chlorophyllous
hairs. The gametangia are of the common, advanced
leptosporangiate type, but antheridia with elongate
end cells are known in some species. Gametophytic
characters of Stenochlaena again corroborate its placement in the present family.
KARYOLOGY AND HYBRIDIZATION. Blechnaceae are
one of the cytologically most complex fern families.
Probable or certain base numbers found so far are 32
(Doodia), 28-37 but most commonly 28 and 33 (Blechnum), 33 (Brainea, Sadleria), 34 and 35 (Woadwardia),
and 74 (from 377) (Stenochlaena). Tetraploids are frequent in Blechnum (Walker 1966, 1973). Salpichlaena,
with n = 40, is kept separate from Blechnum partly because of the divergent base number. Some authors tentatively regard x = 33 as the basic number in the family
(Walker 1973).
A hybrid was reported in Daodia (Parris 1972). The
variability of many species in the Blechnum occidentale
complex in largely due to hybridization (see especially
Walker 1973, Fig. 7; Jermy and Walker 1985, Fig. 43).
SUBDIVISION. As Stenochlaena stands rather apart, subfamilial status seems indicated. The other genera are
certainly closely related, and the lines drawn between
them are not always satisfactory, possibly being too
Woodwardia· Pteridoblechnum
greatly based on characters such as soral shape, sporangial distribution, and venation. When Blechnum,
the central genus, has been revised, its borderlines redrawn, and its natural subdivisions determined, the
picture is likely to change.
EcOLOGY AND DISTRIBUTION. The family is distributed
nearly throughout the world. Blechnum is by far largest
genus, with most representatives in the southern hemisphere extending to the mountains of the tropics and
relatively few species in the northern hemisphere.
Woodwardia is a counterpart of Blechnum, nearly confined to the northern hemisphere, where its distribution pattern is somewhat disjunct and possibly relict.
Most of the other genera are Asiatic-Australasiatic-Pacific. Blechnaceae occur in a wide range of habitats,
mostly terrestrial, but are rarely found in dry environments. They play an important part in the vegetation
of some oceanic archipelagos like Tristan da Cunha,
Juan Fernandez, and Hawaii.
AFFINITY. The relationships of the family are not clear.
Ideas about affinity with Aspleniaceae, because of superficial resemblance in soral shape, have been refuted
by anatomy, karyology, etc. As it is not known whether
free or broken sori, free or anastomosing veins, etc.
represent a primitive or a derived condition, ties with
other families of derived leptosporangiate ferns are
difficult to establish. Relationships with Dryopteridaceae have been suggested but are not very close at
best.
KEY TO THE GENERA.
61
- Sori short, discontinuous, or, if long, the fertile parts
strongly contracted; veins copiously anastomosing, or, if
not, the pinnae undivided
6
6. Lamina deeply pinnatifid to once pinnate, mostly tapering
atthe base; veins fonning 1-3 series ofareoles 4. Doodia
- Lamina pinnate+pinnatifid (or almost bipinnate at base)
or, if simply pinnate or simple, with very amply anastomosing veins; lamina base various
7
7. Lamina anadromous at base, catadromous above; crossveins ± arching; Quter areoles often with short, free included veinlets
3. Steenisiobkdmum
- Lamina catadromous throughout (or occasionally pseudoanadromous or isodromous in places); veins straight, the
meshes angular to about hexagonal; free included veinlets
none
8
8. Lamina strongly dimorphic; fertile leaves with greatly re-
duced laminal parts; petiole winged to near the base
(Queensland)
2. Pteridoblechnum
- Lamina hardly or not dimorphic, or, if strongly so, the petiole unwinged or slightly winged above (northern hemisphere)
1. Woodwardia
Subfamily Blechuoideae
Stem with a simple dictyostele and bearing non-peltate
scales; vascular bundles in the petiole forming a simple U in cross-section; sporangia only borne on or
near veins.
1. Woodwardia J. E. Smith
Fig. 21 A
Woodwardia J. E. Smith, Mem. Acad. Turin 5: 411 (1793).
Anchisfea C. Presl (1851).
Lorinseria C. Presl (1851).
Chieniopteris Ching (1964).
I. Sporangia borne on, and (almost) confined to, a longitudi-
nal vein on either side of the costalcostule and parallel to
it, rarely spreading onto the indusium, or borne on vein
arches flanking the costa/costules; indusium at least initially present
3
Sporangia not confined to such a vein or arch (except in
incompletely fertile leaves of Brainea); indusium none 2
2. Pinnae articulate (Fig. 25 C; sporaniga with acrostichoid arrangement; stem scandent
9. StEnochlJJena
Pinnae non-articulate; sporangia (in fully fertile leaves) following the veins, eventually often seemingly acrosticboid
(Fig. 25 G); stem erect, rather trunk-like
8. Brainea
.1. Veins free, except for the paracostal commissure of fertile
leaves bearing the sorus; or a submarginal commissure
rarely present; sterile parts without the paracostal commissure
4
Veins anastomosing beside the paracostal commissure or
vein arch bearing the sori; or this commissure present in
sterile as well as in fertile leaves
5
4. Leaves simple, pinnatifid, or simply pinnate, or, if bipinnate (rare), the rachis erect; veins of sterile parts quite free
5. BIechnum
Leaves (at least) bipinnate; rachis twining; veins always
joined by a submarginal commissure
6. Salpich/aena
~. Sori mostly long, continuous and parallel to the costules;
leaves pinnate+pinnatified or bipinnate; no anastomoses
rresent beside the arches flanking the costules; fertile parts
hardly or not contracted
7. Sadleria
Terrestrial or epilithic ferns. Stem short-creeping to ascending, or slender and long-creeping, bearing nonclathrate scales. Petiole well-developed, usually slramineous with dark base, often ± scaly, adaxially sulcate. Lamina pinnatifid, pinnate + pinnatifid, or occasionally simple, not reduced at base, with (subconform) terminal pinna or pinnatifid apex, monomorphic or less often dimorphic, strongly catadromous (or
occasionally pseudoanadromous, or isodromous in
places); proliferous buds sometimes present in pinna
rodls or elsewhere. Rachis adaxially grooved, the
groove not continuous with those of the costae. Texture herbaceous to subcoriaceous. Margin minutely to
coarsely serrate-dentate, with strongly ascending teeth.
Veins anastomosing, free included veinlets none; one
to many series of areoles present between costa and
margin. Free vein ends mostly short, terminating just
behind the margin. Sori on paracostal vein arches (giving the impression of an interrupted commissure),
flanking the costae and usually also the costules, or in
more dissected species only the costules, sometimes
embossed. Laminal parts of strongly dimorphic species much reduced in fertile leaves. Spores ellipsoidal,
Filicatae . Blechnoideae
62
Fig. 21 A-C. Blechnaceae. A Woodwardia oriemalis. part of a
segment with venation and sori (x 7). B Doodia caudata.
basal part of pinna with sori (x 4). C Doodia aspera. basal
part of pinna (x 4) (A from Ching 1935; B, C from DUncan
and Isaac 1986)
surface bearing irregular, more or less connected folds,
sometimes echinate or with rod-like elements, the
folds often slightly raised.
Thirteen species, very widespread in the northern
hemisphere, but with large gaps in the area, south to
Costa Rica and western Indonesia, lacking in tropical
Africa, the Pacific, and the Antilles. Greatest species
concentration in eastern Asia.
2. Pteridobkchnum Hennipman
Fig. 22
Pteridoblechnum Hennipman, Blumea 13: 397 (1966); Hen-
nipman (1976).
Medium-sized, terrestrial ferns; stem long-creeping,
unbranched, cylindrical, with three vascular strands;
scales narrowly triangular, acuminate, non-clathrate,
brown, basally attached, sometimes auriculate, irregularly denticulate. Leaves up to a few cm apart, spirally
inserted; petiole stout, well-developed, with 5 vascular
bundles, brownish, adaxially sulcate in the upper half,
Fig. 22. B1echnaceae. Pteridoblechnum neglectum, habit with
one sterile and one fertile leaf (x 0.16) (Jones and Clemesha
1976)
with a lobed wing near its base. Lamina dimorphic, catadromous, deeply pinnatifid, sterile lamina ovate,
± abruptly decurrent onto the petiole, with small lobes
between the segments, segments 20-50, linear-lanceolate, coarsely serrate, glabrous, smooth. Venation sagenioid-reticulate without free included veinlets. Fertile
lamina similar but with linear segments, each segment
consisting of a costa flanked by two linear, indusiate
sori and a vestigial laminal part; indusia opening towards the costa; the wing connecting the segments
shortly continuing into their bases. Spores ellipsoidal,
bearing prominent, coarse tubercles, especially on
their distal face.
A single species, Pt. neglectum (F. M. Bailey) Hennipman, terrestrial in forests, confined to Queensland,
Australia.
Steenisioblechnum . Doodia· Blechnum
3. Steenisioblechnum Hennipman
Steenisiohlechnum Hennipman, Blumea 30: 17 (1984).
Pteridoblechnum Hennipmann (1976), p. p., excl. type.
Medium-sized, terrestrial ferns; stem short-creeping,
ascending, terete, unbranched; scales narrowly triangular, acuminate, brown with paler edge, entire, auriculate. Petioles aggregated, spirally arranged, well-developed, stramineous, terete, the upper half adaxially
with a median and two lateral grooves. Lamina pinnate, anadromous at base, catadromous above, dimorphic, glabrous except for short, stiff, fuscous, multicellular hairs occurring especially adaxially at pinna
insertions. Sterile lamina widest somewhat below the
middle, with up to 10 lateral pinnae and a conform
terminal one which may be trilobed; pinnae widest
somewhat below the middle, the lowest shortly petiolulate, the upper ones sessile, base rounded, apex narrowed, margin serrate. Veins anastomosing in an intricate network, about sagenioid near the margin, forming distinct areoles along costae and major lateral
veins some of which contain free included veinlets
pointing to all sides. Fertile leaves long-petiolate with
linear pinnae; indusium hyaline, marginal. Spores ellipsoidal, bearing distant, coarse echinae or cristae.
A single species, S. acuminatum (C. T. White and
Goy) Hennipman, terrestrial in forests; like the preceding confined to Queensl;tnd.
4. Doodia R. Brown
Fig.21B, C
Doodia R. Brown, Prodr. A. Nov. Holl.: 151 (1810); Goy
(1939); Blasdell (1956); Parris (1972).
Small to medium-sized terrestrial ferns; stem short,
erect or ascending, sometimes stoloniferous, bearing
dark scales. Leaves close, with well-developed petiole,
the latter usually dark, adaxially grooved in the upper
part, often ± scaly, sometimes pubescent and/or verrucose. Lamina very deeply pinnatifid to simply pinnate,
with few exceptions reduced at base, with pinnatifid or
caudiform apex; rachis like the petiole, largely pale, its
adaxial groove not continuous with those of the costae. Pinnae sessile or adnate, ovate to linear, herbaceous to coriaceous, mostly glabrous, with percurrent,
adaxially sulcate costa; margin sharply serrate-dentate.
Venation catadromous or isodromous; veins forked,
forming one or a few series of areoles by means of
urching cross-veinlets; vein ends at the margin beIween the teeth. Fertile pinnae sometimes slightly to
moderatley contracted. Sori short, on the cross-veinlets, sometimes somewhat embossed, in one series next
10 the costa or also in a second series closer to the
margin, the sori of the two series alternating or partly
superimposed; indusium membranous, often dark.
Spores ellipsoidal, low'rugose or papillate, sometimes
nearly plain.
63
About 12 species, but the taxonomy imperfectly
known, several species being quite similar; in Australia, New Zealand, the Pacific Islands east to Hawaii
and Easter Island; a few rare species occurring locally
in Ceylon, Java, the Lesser Sunda Islands, and New
Guinea. Some species are very locally distributed.
This genus seems to be closely related to Woodwardia.
S. Blechnum L
Fig. 23
Blechnum L, Spec. Plant. 2: 1077 (1753) (for revisionallitera-
ture see below).
Struthiopteris Scopoli (1760), non Willdenow (1809,
~ Matteuccia).
Lomaria Willdenow (1809).
Blechnidium Moore (1859).
Diploblechnum Hayata (1928).
Spicantopsis Nakai (1933).
Terrestrial or epilithic, rarely scan dent, small to large
ferns with creeping or erect, sometimes trunk-like or
scandent stem bearing scales that are often dark, long,
and narrow. Petiole usually well-developed, sometimes
winged to base. Lamina typically pinnatifid or simply
pinnate with undivided, long and narrow pinnae, occasionally simple or lobed to pinnatifid, rarely pinnate + pinnatifid to bipinnate. Rachis like the petiole,
often ± scaly, less often hairy or glandular. Pinnae catadromous, or the lower isodromous or weakly anadromous, often adnate and decurrent (or also surcurrent), sometimes with small, interspersed lobes, usually
non-petiolulate, often firm and glabrous; margin often
crenate to sharply dentate. Costa percurrent, adaxially
grooved. Veins free, usually forked, with clavate ends
terminating just behind the margin, often with hydathodes, ending in or between the teeth (if any). Fertile
pinnae similar to the sterile, varying to strongly contracted; veins forming a straight, continuous vascular
commissure flanking the costa and parallel to and often very close to it, the sporangia being borne there but
often strongly spreading at full maturity and covering
the fertile pinnae in a seemingly almost acrostichoid
manner. The vascular commissure may be on a level
different from that to the regular veins, then not being
a true commissure; or it may consist of diffuse conducting tissue (see Holttum 1955, Fig. 259, and de la
Sota and Gouvea Labouriau 1961); in species with
much reduced fertile laminal parts the commissure
may seem to be marginal. Indusium long and often
narrow, initially covering the sporangia, eventually
sometimes fugacious or hidden by the sporangia, in
strongly dimorphic species often seemingly marginal.
Sporangia inserted on the commissure but sometimes
spreading onto the indusium. Spores ellpisoidal, surface very variable: slightly papillate to nearly smooth;
rugose or with wing-like folds; coarsely echinate to
64
Filicatae . Blechnoideae
tingsh. from New Zealand which also diverges by having a slender, high-climbing stem; broken sori and sori
at an angle to the costa in forms of the African
B. punctulatum Sw.; articulate pinnae in B. serrulatum
L. C. Rich. of tropical America and Australasia.
A subcosmopolitan genus of an estimated
150-200 species, taxonomically very incompletely
known; modern revisions exist only for a few species
groups, more for some limited areas (e. g., Looser
1947, 1958; Copeland 1950; Schelpe 1952; Brownlie
1969; A. R. Smith 1985, etc.). The genus is poorly and
only locally represented in north-temperate regions,
very richly in south-temperate regions, in tropicalmontane to subalpine areas, and on oceanic islands;
e. g., New Caledonia has 16 species, New Zealand 15,
Venzuela 24, Chile 11, etc. There are few species in
continental Africa. Many species grow in forests and
thickets, often by streams, other in open places.
A natural subdivision of the genus has not been given. Sterile-fertile dimorphism has been much used in
the past, also for splitting off parts of the genus ("Lomaria", "Struthiopteris"), but this does not bring together natural alliances. Natural entities, as species
groups, have been described for the Neotropics by
Tryon and Tryon (1982).
It is not advisable to exclude species that diverge
only in certain single aberrant characters, as indicated
above.
Strongly dimorphic species have in the past been
confused with Plagiogyria. The latter genus is easily
recognized by the complete absence of scales and by
its adaxially non-sulcate costae.
6. Salpichlaena Hooker
Salpichlaena Hooker in Hooker and Bauer, Gen. RI.: pI. 93
(1842).
Blechnum auctt., p. p. min.
Fig. 23 A, B. Blechnaceae. A, B Blechnum penna-marina.
A Plant with sterile and fertile leaves (x 0.6). B Detail of fer·
tile leaf (x 3) (Looser 1958)
reticulate, the reticulate elements sometimes thin; rarely with spheroidal deposit.
CHARACTERS OF RARE OCCURRENCE. Pinnate+pinnatifid, or even bipinnatifid leaves in the East Asiatic
B.fraseri (Cunn.) Luerssen and a "variety" of the
Madagascan B. microbasis (Baker) C. Chr. Venation as
in Doodia, also in sterile leaves, but with long sori of
Blechnum type, in a few species like the Asiatic
B. ("Blechnidium") melanopus Hooker and the South
American B. heringeri Brade; di- or trimorphic sterile
leaves in a few species like B.filiforme (A. Cunn.) Et-
Very large terrestrial ferns with long-creeping stem
bearing small, rigid, dark (or pale-edged) scales.
Leaves remote; petiole long, stout, stramineous to tan,
adaxially sulcate. Lamina to several m long, with twining, stramineous, smooth or lightly pubescent, adaxially sulcate rachis bearing numerous subopposite, large
primary pinnae, these long-stalked and with long rachidule, the adaxial groove of the latter evanescing towards the stalk. Pinnae pinnate or rarely bipinnate at
base, with several, usually sub opposite, relatively longstalked pinnules, the basal rarely trifoliolate. Pinnules
chartaceous-subcoriaceous, narrowly oblong to linear,
entire, acuminate-cuspidate, with rounded base, the
upper sessile but scarcely reduced and not confluent, a
conform terminal pinnule present. Costa percurrent,
adaxially sulcate; veins close, numerous, evident, simple or basally forked, straight almost to their tips,
joined there by an intramarginal commissure just be-
Salpichlaena· Sadleria
hind the pale·sclerotic margin. A few tan scales some·
times present on axes and costae beneath. Usually the
lower pinnules anadromous, the upper catadromous in
venation. Fertile pinnules weakly to strongly contracted, sometimes almost lamina-less, narrowly oblong to
linear; sori linear, continuous, paracostal, on a continuous vein runnig parallel to the costa and flanking it
on either side, the sporangia confined to this vein or
sometimes spreading to the indusium. Indusium very
long and narrow, continuous, initially completely
covering the sporangia, the edges of two opposite ones
meeting over the costa; no sterile organs among the
sporangia. Spores monolete, ellipsoidal, faintly papillate-rugose.
A single species, S. volubilis (Kaulf.) Hooker, widespread in tropical America, from Nicaragua and the
Lesser Antilles to Bolivia and SE Brazil. Scandent in
trees and shrubs, in forests and thickets, at lower to
higher elevation.
Forms with varying degrees of contraction of fertile
pinnules do not merit taxonomic recognition.
65
Fig. 24. Blechnaceae. Sadleria pallida. abaxial side of fertile
lamina; cult. Berkeley, California. Phot. K. U. Kramer
7. Sadleria Kaulfuss
Fig. 24
Sadleria Kaulfuss, En. Fil.: 161 (1824).
Medium-sized to large terrestrial ferns, often growing
on lava. Stem erect, often trunk-like with age, bearing
entire, non-clathrate, brown scales. Leaves borne in a
rosette; petiole stramineous or less often dark, mostly
stout, adaxially sulcate, bearing very long, narrow,
partly filiform, light brown scales, especially at base.
Lamina pinnate+pinnatifid or bipinnate; rachis like
the petiole, stramineous, often ± scaly, adaxially with
at least one groove which ist not continuous with the
grooves on rachidules and/or costae. Texture of lamina firm; upper pinnae, pinnules, etc. gradually reduced and confluent; basal pinnae somewhat shortened. Pinnae sessile, catadromous (or isodromous);
basal segments sometimes springing from rachis rather
than from rachidule. Segments numerous, ovate to
tongue-shaped, mostly adnate, connected or free, subentire to crenate, the upper confluent; surfaces glabrous or scaly beneath. Costae evident; veins forked,
free but joined by costal arches whether sterile or fer-
66
Filicatae . Blechnoideae • Stenochlaenoideae
C
Fig. 25A-G. Blechnaceae. A-E Stenochlaena palustris.
A Parts of sterile and fertile lamina (x 0.25). B Tip of sterile
pinna with venation (x 1). C Insertion of sterile pinna on rachis (x 2.5). D Thmsection of fertile pinna (x 5). E Peltate
scale of stem (x 20). F, G Brainea insignis. F Transection of
petiole ( x 12.5). G Base and apex of pinna showing arrangement of sporangia and venation (x 1.3) (A-D from Holttum
1955; E,F from Tardieu-Blot 1932; G from Flora Taiwan
1975)
tile. Sori as in Blechnum but mostly shorter, slightly
embossed. Spores distinctive for the species (lloyd
1976), ellipsoidal, bearing small, irregular sheaths,
these usually more or less eroded and exposing the
lower reticulate formation.
Probably four species; the only fern genus restricted
to Hawaii. Some species are pioneers on lava (see,
e.g., Holbrook-Walker and lloyd 1973). Very closely
related to Blechnum and perhaps only infragenerically
distinct.
8. BraineaJ. Smith
Fig. 25
Brainea J. Smith, Catal. Kew Ferns: 5 (1856).
Rather large terrestrial ferns with erect, ±trunk-Iike
stem bearing an apical rosette of leaves; apex bearing
long, narrow, entire, centrally dark-banded scales.
Petiole well-developed, stramineous, adaxially unisulcate. Lamina medium-sized to large, firm, simply pinnate (very rarely bipinnate at base), with numerous
sessile, linear pinnae, the lower somewhat reduced, the
upper gradually reduced and confluent; rachis like the
petiole, scaly or glabrescent, its adaxial groove not
continuous with those of the costae. Costa evident,
percurrent, scaly or glabrescent beneath; veins forked,
parallel, free except for the basal parts which are connected by a series of short paracostal arches in both
sterile and fertile leaves. Venation catadromous. Margin crenate, ± revolute. Sparingly fertile pinnae with
naked sporangia borne on the vein arches; fully fertile
pinnae so~ewhat contracted, bearing the sporangia
along most of the length of the veins, almost entirely
covered by them at full maturity. Spores ellipsoidal, irregularly low-rugose to nearly plain.
A single species, B. insignis (Hooker) J. Smith, growing in open places and thickets, distributed from
Brainea • Stenochlaena
NE India, S. China, and Taiwan to Sumatra and
Malaya. Closely related to Blechnum, differing in venation, sporangial disposition, and lack of an indusium.
Subfamily Stenochlaenoideae
Stem long-scan dent, dictyosteIic, with a few larger,
central and many smaller, peripheral vascular bundles,
bearing peltate scales. Vascular bundles of the petiole
with many vascular bundles in two circles; sporangia
with acrostichoid disposition.
Monogeneric.
9. Stenoch1aena J. Smith
Fig. 25A-E
Stenochlaena J. Smith, Hooker's J. Bot. 3: 401 (1841); Holttum (1932, 1971).
Stem very long-creeping or scandent, the outer bundles of the dictyostele roughly arranged in 2 or 3 concentric circles; surface of stem green, sparsely beset
with small, peltate, round to narrow scales, eventually
glabrescent except for the apex. Leaves remote; petioles non-articulate, glabrous or slightly hairy, well-developed, adaxially narrowly sulcate and with some additional ridges and grooves, with a continuous lateral
aerating line that extends onto the stem. Lamina
strongly dimorphic. Sterile lamina simply pinnate with
an odd, conform terminal pinna; rachis like the
petiole; pinnae usually alternate, (sub)sessile,
± distinctly articulate at base except for the terminal
pinna, oblong-Ian ceo late with unequal base, not lobed,
margin sharply toothed, lower surface sometimes
short-hairy; costa percurrent, adaxially shallowly
grooved, the groove evanescent at base. Veins close,
parallel, simple or forked, ending at the sclerotic margin in (and between) the teeth, free except for a series
of long, very narrow, inconspicuous costal areoles;
venation anadromous at base of lamina, in the upper
pinnae ± distinctly catadromous; anterior base of pinnae with a callose gland, similar glands often present
on rudimentary, deciduous pinnae below the normal
ones. Fertile lamina once (or in one species twice) pinnate, with very narrow, articulate pinnae, these beneath covered by the sporangia except for a narrow
sterile marginal band; venation similar to that of the
sterile pinnae, the commissure more evident. Sterile
organs among the sporangia wanting. Spores ellipsoidal, bearing distant tubercles, mostly on distal face, or
these sometimes fused in parallel rows.
Six species (Holttum 1971) in tropical and warmtemperate parts of the Old World, two in Africa (to
South Africa) and Madagascar, the others from India
to NE Australia, east to Tonga and Samoa. Formerly
credited with a pantropic distribution and many more
species, due to confusion with the genera Teratophyl-
67
lum and Lomariopsis, here placed in Lomariopsidaceae. Holttum (1932) was the first to elucidate this
confusion.
Somewhat divergent from the other genera of Blechnaceae, mainly in anatomical characters; but venation,
stomata, spores, and gametophyte agree best with this
family. If the base number would be shown to be really x ~ 37, this would constitute another confirmation.
The ontogeny of the venation was studied by Leist
(1975).
Selected Bibliography
Blasdell, R. F. 1956. A comparative study of Hawaiian species of the fern genus Doodia. Bull. Torrey Bot. Club 83:
62-71.
Brownlie, G. 1969. Hore de la Nouvelle-Caledonie et Dependances.3. Pteridophytes. Paris: Museum National
d'Histoire Naturelle.
Ching, R C. 1978. The Chinese fern families and genera, etc.
Acta Phytotax. Sin. 16: 1-37.
Copeland, E. B. 1950. Aspleniaceae and Blechnaceae of New
Guinea. Philipp. J. Sci. 78: 207-229.
Goy, D.A. 1939. The rasp fern (Doodia) of Australia.
Queensl. Natural. 11: 23-38.
Hennipman, E. 1966. Pteridoblechnum, a new genus of Blechnoid ferns from Australia. Blumea 13: 397 -403.
Hennipman, E. 1976. A second species of Pteridoblechnum
(Blechnaceae) formerly referred to Leptochilus acuminatus
C. T. White and Goy. Proc. R. Soc. Queensl. 87: 95-98.
Holbrook-Walker, S. G., Lloyd, R. M. 1973. Reproductive biology and gametophyte morphology of the Hawaiian fern
genus Sadleria (Blechnaceae), etc. Bot. J. Linn. Soc. 67:
157-174.
Holttum, R. E. 1932. On Stenochlaena, Lomariopsis and
Teratophyllum in the Malay region. Gard. Bull. S. S. 5:
245-312.
Holttum, R. E. 1971. The genus Stenochlaena J. Smith with
description of a new species. Amer. Fern J. 61: 119-123.
Leist, N. 1975. Die Entwicklungsgeschichte der Blattoervatur
beim Farn Stenochlaena tenuifolia (Blechnaceae). Plant
Syst. Eyol. 124: 213-228.
Lloyd, R M. 1976. Spore morphology of the Hawaiian genus
Sodleria(Blechnaceae). Amer. Fern J. 66: 1-7.
Looser, G. 1947. Los Blechnum (Filices) de Chile. Rev. Univ.
(Santiago) 32: 7 -1 06.
Looser, G. 1958. Clave de los Blechnum (Filicales) de Chile.
Rev. Uniy. (Santiago) 43: 123-128.
Mehra, P. N., Chopra, N. 1951. Anatomy of Stenochlaena
palustris (Bunn.) Bedd. Ann. Bot. II., 15: 37-45.
Murillo, M. T. 1968. Blechnum subgenero Blechnum en sur
American, con especial referencia a las especies de Colombia. Nova Hedwigia 16: 329- 366.
Parris, B. S. 1972. The genus Doodia R. Br. (Blechnaceae: Filicales) in New Zealand. New Zeal. J. Bot. 10: 585-604.
Schelpe, E. A. C. L. E. 1952. A revision of the African species
of Blechnum. J. Linn, Soc. 53: 487-510.
Smith, A. R. 1985. Pteridophytes of Venezuela, an annotated
list. Berkeley, Cal.
De la Sota, E. R, GOUyea Labouriau, L 1961. Correlaciones
entre esporangios y nervadura comisural en Blechnum 0ccidentale L. (Filices, Blechnaceae). An. Acad. Bras.
Ci~nc, 33: 225-239.
68
Filicatae . Cheiropleuriaceae
Stokey, A. G., Atkinson, L. R. 1952a. The gametophyte of
Stenochlaena palustris (Bunn.) Bedd. Phytomorphology 2:
1-9.
Stokey, A. G., Atkinson, L. R. 1952b. The gametophyte of
Blechnwn spicant (L.) Wither and B. buchtienii Rosenst.
Phytomorphology 2: 9-15.
Stone, I. G. 1962. The ontogeny of the antheridium in some
leptosporangiate ferns with particular reference to the funnel-shaped wall. Austr. J. Bot. 10: 76-92.
Tardieu-Blot, M. 1932. Les Aspleniees du Tonkin. Toulouse.
Walker, T. G. 1%6. A cytotaxonomic study of the pteridophytes of Jamaica. Trans. R. Soc. Edinburgh 66(9):
169-237.
Walket; T. G. 1973. Additional cytotaxonomic notes on the
pteridophytes of Jamaica. Trans. R. Soc. Edinburgh 69(5):
109-135.
Cheiropleuriaceae
K. V_KRAMER
Cheiropleuriaceae Nakai, Bot. Mag. (Tokyo) 42: 210 (1928).
Terrestrial or epilithic ferns of medium size with creeping, protostelic, rarely branching stem covered with
long, uniseriate, articulate hairs_ Petioles more or less
close, hairy at the base only, non-articulate, severalranked, terete, upward gradually flattened and marginate, with two vascular strands at base which divide
upward Lamina naked, strongly dimorphic; sterile
lamina firmly herbaceous, more or less deeply bilobed
or not rarely simple, rarely 4-lobed, hypostomatic, the
main veins dichotomously forking with anadromous
branching, the smaller veins areolate with smaller areoles included in the larger ones and numerous free included ultimate veinlets, all veins more or less prominent; stomata paracytic. Fertile lamina simple, long,
and narrow, with three longitudinal main veins, abaxially almost wholly covered by the sporangia, these
served by special branches of the veins running to near
the surface. Sporangia of mixed maturation, intermingled with hairs with swollen terminal cell; stalk
short, massive, quadriseriate, constricted below the
head; head with almost vertical yet uninterrupted annulus largely consisting of the bow, with well-differentiated stomium_ Spores, achlorophyllous, c_ 128 per
sporangium, trilete, tetrahedral, with prominent ridge
near the aperture; surface smooth to rugulose_
ANATOMY. The occurrence of protostely in a stem of
fair dimension is a rare phenomenon in recent leptosporangiate ferns; so is the purely dichotomic branching pattern of the petiolar vascular supply whose
branches lie almost in one plane; both characters
point to the primitive nature of the genus- The acrosti-
Fig_26A-D_ Cheiropleuriaceae. A-D Cheiropleuria hiscuspis. A Habit (x 0.4). B Detail of venation (x 3.5). C Trichome from stem (x 14). D Sterile appendage from among
the sporangia (x 18) (Hu and Ching 1930)
choid disposition of the sporangia and the special vein
branches serving them are, on the other hand, derived
characters. See further Nozu (1955).
GAMETOPHYTE_ The prothallium, described by Stokey
and Atkinson (1954) and by Nayar and Kaur (1971), is
very similar to that of Dipteris, including the structure
of the gametangia, which, however, are confined to the
ventral side_
AFFINITY. Except for Diptem, Cheiropleuria has no obvious relatives. The differences in stem, anatomy, sori,
spores, etc_ seem to warrant separation at the family
level. As discussed under Diptem, the Polypodiaceae
. (s_ str.) do not form a natural place for these two genera, and it is surprising that they were retained there
by, e. g., Copeland (1947) whose families are not usually inclusive. Cheiropleurla was associated by TardieuBlot and Christensen (1941) with Platycerium, which is
quite untenable in the light of modern insight. For a
further discussion see Jarrett (1980) where alliance
Cheiropieuria
with very primitive leptosporangiate families like Matoniaceae and Gleicheniaceae is discussed.
KARYOLOGY. Apogamy was reported by Stokey and
Atkinson (1954). A chromosome number of n =33 was
reported in passing by de la Sota (1973), but the source
of this information was not disclosed.
A single genus:
Cheiropleuria C. Presl
Fig. 26
Cheiropieuria C. Presl, Epimel. Bot. 189 (1849).
Characters as for family.
A single species, Ch. biscuspis (Blume) C. Presl, ranging from Indo-China and Honshu to eastern Malesia,
growing on the ground or on rocks in shaded places,
in the tropics at middle and higher elevation. The simple-leaved form has been distinguished as "var. integrifolia"but is not worthy of taxonomic recognition.
Selected Bibliography
De la Sota, E. R. 1973. On the classification and phylogeny of
the Polypodiaceae. Bot. J. Unn. Soc. 67 Suppl.1: 229-244.
Jarrett, F. M. 1980. Studies in the classification of the Lepto·
sporangiate ferns: I. The affinities of the Polypodiaeeae
sensu stricto and the Grammitidaceae. Kew Bull. 34:
825-833.
Nozu, Y. 1955: On Cheiropieuria biscuspis var. anguslifoiia.
etc. Bot. Mag (Tokyo) 68: 86-93.
Stokey, A. G., Atkinson, L. R. 1954. The gametophyte of
Cheiropieuria bicuspis (BI.) Presl. Phytomorphology 4:
192-201.
Tardieu-B1ot, M., Christensen, C. 1941. Cryptogames vaseulaires. In: Lecomte, H. (Ed.) Aore Gen. Indo·Chine 7(9):
1-596. Paris.
Cyatheaceae
K.U.KRAMER
Cyatheaceae Kaulfuss, D. Wesen d. Farrenkr.: 119 (1821).
Large to very large, less often medium-sized, terrestrial
ferns with short to usually long, erect, slender to robust, apically scaly stem; stele radially symmetric,
forming a complicated dictyostele with much sclerenchyma on both sides of the bundles, often with accessory medullary and cortical bundles; the surface often
hearing a cover of densely matted adventitious roots,
rarely spiny, the leaf scars prominent, often the upper
ratt with petiole base remnants, rarely branched.
69
Leaves forming a rosette; petioles usually close, stout,
short to long, bearing scales often inserted on wart- to
spine-like epidermal outgrowths, mostly hairy in addition, the adaxial side flattened to sulcate and, with few
exceptions, like the other leaf axes bearing antrorse,
septate hairs, laterally with 1 (-3) discontinuous lines
of pale pneumathodes; petiole bases sometimes with
so-called aphlebiae (see below). Lamina pinnate to
(more often) bipinnate (+ pinnatifid), herbaceous to
coriaceous, usually large, often oblong or deltoid,
truncate at the base or the lower pinnae somewhat to
Oess often) strongly and gradually reduced; leaf architecture entirely catadromous. Vascularization of the
petiole essentially of an abaxial and an adaxial arch of
many small bundles, but often with additional series
(Fig. 27 A). Rachis adaxially sulcate, the groove usually
raised in the middle; additional flanking grooves often
present in larger species, with a ± persistent indument
of .scales and/or hairs, often also warts or spines;
adaxial indument as in the petiole. Leaf axes stramineous to tawny, less often dark and polished. Upper pinnae, pinnules, etc., with few exceptions gradually reduced and confluent. Pinnae stalked or sessile, the
petiolule often with a dark articulation-like discontinuity at its base or occasionally truly articulate. Pinna rachis adaxially sulcate, the groove raised in the middle,
mostly strigose, the groove not continuous with that on
the primary rachis. Lamina at least initially scaly and
(with few exceptions) hairy, these appendages persistent or not; scales on abaxial side of costae, costules,
etc. not rarely bullate and/or pale. Ultimate free
divisions ± symmetrical, very often Iiguliform and to
some degree falcate, sometimes roundish, entire to crenate or pinnatifid, mostly adnate but sometimes narrowed at the base, sessile, costate. Lower surface
sometimes glaucous. Veins issuing from the costae/
costules in a pinnate pattern, catadromous or rarely
isodromous, the posterior basal one usually issuing
from the costule, often from its base; veins simple or
usually (subpinnately) forked, free or, especially in the
Cnemidaria group, the basal posterior branches anastomosing in pairs and forming costal arches, their free
excurrent branches connivent to the sinus (Fig. 28 A).
Costae and costules adaxially raised, mostly strigose.
Veins not quite reaching the margin. Sori dorsal on the
veins, most often at a fork, round; receptacle strongly
to hardly raised, containing a vein branch. Indusium
attached round the receptacle base, apically with a
small to large opening, in the former case irregularly
rupturin at maturity (Fig. 28D); or more unilateral, attached at the costular side only, small to large and
scale-like (Fig. 28C); or wanting altogether (Fig. 28E);
in a few species of subgen. Sphaeropteris replaced by a
group of scales placed round the receptacle base.
Sporangia numerous, rather small, with a short, 4-seriate stalk, mostly intermingled with pluricellular hairs
70
Filicatae . Cyatheaceae
G
Fig. 27 A-H. Cyatbeaceae. A Cyathea lepifera. vascular bundles in transections of petiole and rachis, at successively higher levels (schematic). B-H Details of petiolar scales of Cyathea species (all except F c. x 30). B Cyathea mollicula.
C Cyathea marginalis. D Cyathea myosuroides. E Cyathea
condnna. F Cyathea manniana (x 90). G Cyathea salvinii.
H Cyathea nockii (A from Ogura 1972; B-H from R. M. Tryon 1970)
with or without glandular end cell and occasionally
broadened and scale-like at base. Annulus almost vertical, complete, bypassing the stalk; stomium well-developed, consisting of several narrow cells. Spores trilete, globose, mostly 64, sometimes only 16 per sporangium.
CHARACfERS OF RARE OCCURRENCE. Adaxially nonstrigose axes up to costules in the Cnemidaria group
and a few other species. Vein arches also along the
bases of the costules in a few species of the same
group. Lamina simple in a single species, C. sinuata
Hooker and Grev., from Ceylon. Contracted fertile divisions, sometimes also with revolute lobes, in sect
Gymnosphaera. Scandent stems in C. ("'Thysanobotrya") biformis (Rosenst.) Cope\., which has almost
lamina-less fertile segments and whose sterile leaves
are also dimorphic, with abortive lamina apex. A prostrate stem in C. colensoi (Hook. f.) Domin (New Zea-
land), a short-creeping, rhizome-like stem in the
E. Asiatic C. hancockii Cope\. Constantly and regularly branched stems in C. /Uyamae Ohba from the
Bonin Is. Anadromous pinnae occur very occasionally
in one species or another. Stem branches serving as
struts and for vegetative propagation in a few African
species (Halli: 1966; Tryon and Tryon 1982).
ANATOMY AND MORPHOLOGY. Much work has been
done on the family, but only a few species have been
thoroughly investigated. The most important studies
are by Ogora (1927, 1972), Holttum and Sen (1961),
Lucansky (1974), Lucansky and White (1974), Sen
(1964), A. F. Tryon and Feldman (1975), Holttum
(1957) (scales), Gastony and Tryon (1976) (spores),
Gastony 1973 (id.).
The stomata are anomocytic, polocytic, and acyclicparacytic (van Cotthem 1970). Tindale (1956) drew attention to the pneumathodes on the leaf axes and their
taxonomic relevancy. Holttum (1957) first demonstrated the presence of two types of scales, viz. the
setiferous type with entire margin and sharply offset
marginal hairs (Fig. 27 D, E), and the flabelloid type,
without such hairs but with irregularly fringed margins
and peltate base (Fig. 27 F, G, H). These scales proved
to be much more important for the taxonomy of the
family than the indusia on which former subdivisions
were based. This was first elaborated by Holttum and
Sen (1961) and by R. M.lIyon (1970), who empha-
Filicatae . Cyatheaceae
71
A
c
sized the difference between scales with and scales
without a terminal seta, structurally marginate vs. nonmarginate scales; and the so-called squaminate scales
of the Nephelea group (see also Gastony 1973), where
some spines fonn part of a scale, transitions between
the organs being present (but see Holttum and Edwards 1983). It is still a matter of contention whether
the indusium is homologous with the inner indusium
of the Dicksoniaceae or not (A. F. Tryon and Feldman
vs. Holttum). The presence of characteristic, so-called
cubical cells with crystals on the border of sclerenchyrna and vascular bundles was descnbed by Holttum
and Sen (1961) and adduced as evidence for the close
relationship between scaly (Cyatheaceae) and hairy
(Dicksoniaceae, Metaxyaceae, Lophosoriaceae) tree
ferns; there are further important points of taxonomic
similarity.
The peculiar elaminate, capillary, highly branched
outgrowths near the petiole base found in a number of
species have been called aphlebiae and have attracted
much attention (e. g., Goebel 1900; Tardieu-Blot 1941;
Conant 1983). There are sometimes transitions to reduced but otherwise normal pinnae (e. g., in C. setosa;
see Gastony 1973).
For detailed descriptions of the spores, see especially Gastony (1974, 1979), Gastony and Tryon (1976),
lind Liew and Wang (1976). The spore structure of the
infrageneric groups as recognized by Holttum (in
Holttum and Edwards 1983) may be summarized as
l"allows: Subgen. Sphaeropteris: surface plain with
coarse, echinate elements or low ridges and granulate-
Fig.28A-G. Cyatheaceae. A Cyathea (Cnemidaria) grandifolia, detail of pinna with venation and sori (x 1). B Cyalhea
roraimensis, part of median pinna with venation and hemitelioid indusia (x 3). C Cyathea heterochlamydea. hemitelioid
indusium (x 10). D Cyathea aenulata, cyatheoid indusium
( x 10). D Cyathea crenulata, cyatheoid indusium (x 10).
E Cyathea glabra, indusium none (x 10). F,G Cyathea capensis, sporangium, viewed from two different sides (A from
Stolze 1974; B from Holttum and Edwards 1983; C-E from
Holttum and Sen 1961; F,G from Holttum 1963)
verrucate with rodlets, rarely foraminate; subgen. Cyathea: sect. Nephelea, surface bearing low, long or
short ridges to nearly cristate, not echinate; sect. Cyathea, surface bearing rodlets over a plain or verrucate exospore, rarely foraminate or lacking rodlets;
sect. Alsophila, surface bearing branched and usually
echinate ridges, or rodlets and a granulate deposit
over a plain or verrucate surface.
The peculiar surface structure of the group treated
as the genus Cnemidaria by R. Tryon (1970, etc.) and
Stolze (1974), together with the lack of trichomes on
the adaxial $ide of the lamina and the characteristic
venation, seem to indicate that it merits recognition as
an infrageneric group, a status which Holttum (in
Holttum and Edwards 1983) did not accord it. The
spores are pitted, usually with three large, equidistant
lobes in the equatorial area.
KARYOLOGY AND HYBRIDIZATION. Cytologically the
family is very homogeneous, with universally n = 69,
72
Filicatae • Cyatheaceae
Fig. 29. Cyatheaceae. Cyathea excelsa, crowns forming part
of canopy of montane forest; La Reunion, Mascarenes. Phot.
K.U.Kramer
no polyploids and no naturally occurring cases of
apogamy being known.
A considerable number of hybrids have been described, some involving different sections (Conant
1975,1983; R. M. Tryon 1976; Holttum 1984), providing additional evidence against recognizing any segregates of Cyathea (see below). Hybridization is supposed to provide a mechanism for speciation, even in
the absence of allopolyploidy (Conant and CooperDriver 1980).
No hybrids involving parent species from different
subgenera have so far been found.
GAMETOPHYTE. This was most fully described by Stokey (1930). Mature prothallia are heart-shaped or occasionally more elongate, with a thick midrib. Peculiar
multicellular, usually multiseriate, green hairs occur on
both surfaces of older prothallia and offset the family
from the Dicksoniaceae where they are lacking. Antheridia are of a primitive, five-celled type with one or
two stalk cells. Archegonia are also primitive, with
long, rather straight necks. See also Mukherjee and
Sen (1986) for further data and for points of similarity
with the prothallia of Dicksoniaceae.
EcOLOGY AND DISTRIBUTION, The species are concentrated in the tropics where they are most numerous in
montane to (locally) alpine vegetation (Fig.29). Many
species occur in the undergrowth of moist forests, often in ravines; others prefer more open habitats, even
swamps, and some grow preferentially in cleared areas
(especially species of subgen. Sphaeropteris in Malesia), where they often form an important constitutent
of secondary vegetation. Such species are often gregarious, notwithstanding the apparent absence of vegetative propagation (for cases where such propagation
has been established, see Halle 1966 and Brade 1971).
A number of species extend far into south-temperate areas, e. g., in New Zealand and southern South
America, but only into very few parts of the north-temperate zone such as India, South China, and southern
Japan. Even remote tropical islands usually harbour
one or a few, often endemic, species. Continental Africa has only 14 species (Holttum 1981).
Some species are widely distributed, but local endemics are quite numerous, especially on islands and
on tropical mountains (Andes, New Guinea). The
Cyathea
73
problem of endemism in tree ferns and its possible origin was discussed by R. Tryon and Gastony (1975).
Few autecological studies on tree ferns have been
undertaken, and little is known about the age of individual plants and the longevity of their leaves. Notable
exceptions are to be found in Seiler (1981), Tanner
(1983), Holttum and Edwards (1983), and Ortega
(1984).
the subdivisions are constant in indusial characters,
other not at all, thus underscoring the unnaturalness of
the three "old" genera Alsophila, Hemitelia. and Cyathea, as defined by indusial characters alone.
A single genus:
AFFINITV. Formerly the Cyatheaceae were usually associated with the Dicksoniaceae, but this idea was
abandoned at Bower's (1926) suggestion. The association was re-established by Holttum and Sen (1961)
who even merged the two families, an idea that has
not won general acceptance. Metaxya and Lophosoria
have been excluded but are usually regarded as related
to both. The affinity of Cyatheaceae and Dicksoniaceae is now generally recognized.
Little ist known about affinity to other fern families.
Points of resemblance to Thelypteridaceae are thought
to reflect affinity (see especially Holttum 1971), but
this is again not generally accepted.
(1930); Tindale (1956); Holttum (1954, 1963, 1964,
1965 a, b, 1974, 1981, 1982); R. M. Tryon (1976); Brownsey
(1979); Holttum and Edwards (1983).
Sphaeropteris Bernhardi (1802); R. M. Tryon (1971); Win·
disch (1977, 1978).
Alsophiia R. Brown (1810); Riba (1967, 1%9); Conant (1983).
Hemite/ia R. Brown (1810).
Trichipteris ~ Trichopteris C. Presl (1822); Barrington (1978).
Gymnosphaera Blume (1828).
Cnemidaria C. Presl (1836); Stolze (1974).
Schizocaena Hooker (1838).
Nephe/ea R. M. Tryon (1970, 1973).
SUBDIVISION. The classification into three genera,
based on the structure of the indusium, or its absence,
was challenged by Domin (1929, 1930, with a predecessor in Copeland 1909), but only gradually abandoned. This author's proposal to recognize only a sino'
gle genus gained increasing acceptance, although
Copeland (1947) still recognized a few minor segregates. Holttum (1955 and later) has only one genus. A
system recognizing five genera was recently proposed
by Lellinger (1988).
In 1970 R. M. Tryon proposed a new system with
six, partly new, partly revived and redefined genera of
scaly tree ferns, based in part on the subdivisions of
Cyathea sens. lat. as adopted by Holttum (1963). This
system was much followed, particularly in America,
but rejected by, among others, Brownlie (1969, 1977),
Holttum (1977), Kramer (1978), Holttum and Edwards
(1983), etc., and again defended by Tryon (1977). Holttum's (and others') rejection of Tryon's segregates is
followed here, for reasons to be found in the cited papers. The presence of inter"generic" hybrids and the
c-onstancy of the chromosome number bear out this
treatment (see, e. g., Jermy and Walker 1985). Even a
seemingly clear-cut group like Cnemidaria (Stolze
(974), at first also accepted by Holttum (1963), can be
delimited only with difficulty, as the limits come out
differently whether one lets macromorphological characters or spore characters prevail (cf. Tryon 1970 with
Stolze 1974).
The subdivision as given by Holttum in Holttum
lind Edwards (1983), with two subgenera, Cyathea and
Sphaeropteris, mainly based on characters of the petiolar scales, each subgenus with two sections and a number of subsections, seems the best founded. Some of
Cyathea J. E. Smith
Fig. 27-29
Cyathea J. E. Smith, Mem. Acad. Thrin 5: 416 (1793); Domin
Characters of the family. About 600-650 species, distributed in the warmer parts of the world, lacking in
dry areas and largely in the north-temperate zone.
Selected Bibliography
Barrington, D. S. 1978. A revision of the genus Trichipteris.
Contrib. Gray Herb. 208: 3-93.
Brade, A C. 1971. Cyathea sampaioana Brade et Ros. somente uma "forma" de Cyathea sternbergii Poh!. Bradea
1(10): 73-76.
Brownlie, G. 1969. Flore de la Nouvelle-Caledonie et Dependances. 3. Pteridophytes. Paris: Museum National
d'Histoire Naturelle.
Brownlie, G. 1977. The Pteridophyte flora of Fiji. Vaduz:
Cramer.
Brownsey, P.J. 1979. Cyathea cunninghamii in New Zealand
(with key to the New Zealand species). New Zeal. J.
BOI.17: 97-107.
Conant, D. S. 1975. Hybrids in American Cyatheaceae. Rhodora 77: 441-455.
Conant, D. S. 1983. A revision of the genus Alsophiia (Cyatheaceae) in the Americas. J. Arnold Arbor. 64: 333-382.
Conant, D. S., Cooper-Driver, G. 1980. Autogamous a110homoploids in A/sophila and Nephe/ea (Cyatheaceae): a
new hypothesis for speciation in homoploid homosporous
ferns. Amer. J. Bot. 67: 1269-1288.
Copeland, E. B. 1909. New species of Cyathea. Philipp. J.
Sci. 3: 353-357.
Domin, K. 1929. Pteridophyta. Praha (in Czech). Nakladem
Ceske Akademie VOrl a Umeni.
Domin, K. 1930. The species of the genus Cyathea J. E. Sm.
Acta Bot. Bohem. 9: 85-174.
Gastony, G.J. 1973. A revision of the fern genus Nephe/ea.
Contrib. Gray Herb. 203: 81-148.
Gastony, G. J. 1974. Spore morphology in the Cyatheaceae. I.
The perine and sporangial capacity: general considerations. Amer. J. Bot. 61: 672-680.
Gastony, G.J. 1979. Id. III. The genus Trichipteris. Amer. J.
Bot. 66: 1238-1260.
74
Filicatae . Davalliaceae
Gastony, G.J., Tryon, R. M. 1976. Spore morphology in the
Cyatheaceae. II. The genera Lophosoria. Metaxya. Sphaeropteris. Alsophi/a. and Nephe/ea. Amer. J. Bot. 63:
738-758.
Goebel, K. 1900. Organographie der Pflanzen. II. Jena:
Gustav Fischer.
Halle, F. 1966. Etude de la ramification du tronc chez
quelques fougeres arborescentes. Adansonia N. S. 6:
405-424.
Holttum, R. E. 1957. The scales of Cyatheaceae. Kew Bull.
1957: 41-45.
Holttum, R. E. 1963. Cyatheaceae. Fl. Males. II. 1': 65-176.
Holttum, R. E. 1964. The tree ferns of the genus Cyathea in
Australasia and the Pacific. B1umea 12: 241-274.
Holttum, R. E. 1965 a. Tree ferns of the genus Cyathea in
Java. Reinwardtia 7: 5-8.
Holttum, R. E. 1965b. Tree ferns of the genus Cyathea Sm. in
Asia (excluding Malaysia). Kew Bull. 19: 463-487.
Holttum, R. E. 1971. Studies in the family Thelypteridaceae.
III. A new system of genera in the Old World. Blumea 19:
17-52.
Holttum, R. E. 1974. The tree ferns of the genus Cyathea in
Borneo. Gard. Bull. Singapore 27: 167-182.
Holttum, R. E. 1977. An exchange of views on the Cyatheaceae. Fl. Males. Bull. 30: 2835-2839.
Holttum. R. E. 1981. The tree ferns of Africa. Kew Bull. 36
463-482.
Holttum, R. E. 1984. A remarkable Cyathea hybrid. Amer.
Fern J. 74: 19-21.
Holttum, R. E., Edwards, P.J. 1983. The tree ferns of Mount
Roraima and neighbouring areas of the Guayana Highlands with comments on the family Cyatheaceae. Kew
Bull. 38: 155-188.
Holttum, R. E., Molesworth-Allen, 8. 1967. The tree ferns of
Malaya. Gard. Bull. Singapore 22: 41-51.
Holttum, R. E., Sen, U. 1961. Morphology and classification
of the tree ferns. Phytomorphology 11: 406-420.
Jermy, A. C., Walker, T. G. 1985. Cytotaxonomic studies of
the ferns of Trinidad. Bull. Brit. Mus. (Nat. Hist.) Bot.
Ser. 13(2): 133-276.
Kramer. K. U. 1978. The pteridophytes of Suriname. Uitg.
Natuurwetensch. Studiekr. Surin. Ned. Ant. 93. Utrecht.
Lellinger, D.8.. 1988. The disposition of 1richopteris (Cyatheaceae). Amer. Fern J. 77 (1987): 90-94.
Liew. E S., Wang, S. Ch. 1976. Scanning electron microscopical studies in the spores of pteridophytes. VIII. The tree
fern family (Cyatheaceae) and its allied species found in
Taiwan. Taiwania 21: 251-267.
Lucansky, T. W. 1974. Comparative studies of the nodal and
vascular anatomy in the neotropical Cyatheaceae. II. Squamate genera. Amer. J. Bot. 61: 472-480.
Lucansky, T. W., White, R. A. 1964. Id. III. Nodal and petiole
patterns; summary and conclusion. Amer. J. Bot. 61 :
818-828.
Mukherjee, A. K., Sen, T. 1986. Gametophytes of some tree
ferns and their impact on phylogenetic relationships. Indi-
an Fern J. 3: 70-81.
Ogura, Y. 1927. Comparative anatomy of Japanese Cyatheaceae. J. Fac. Sci. Univ. Tokyo (Bot.) 1: 141-350.
Ohba. H. 1982. A branched tree fern. J. Jpn. Bot. 57:
321-326.
Ortega, E 1984. Notas sobre la autecologia de Sphaeropteris
senilis(KJ.) Tryon, etc. Pittiera 12: 31-53.
Riba, R. 1%7. Revision monogr"fica del complejo A/sophi/a
swartziana Martius (Cyatheaceae). An. lnst. BioI. Univ.
Nac. Auton. Mex. 38 Ser. Bot. 1: 61-100.
Riba, R. 1969. The Alsophi/a swartziana complex (Cyatheaceae). Rhodora 71: 7-17.
Seiler, R.1.. 1981. Leaf turnover rate and natural history of
the Central American tree fern Alsophi/a salvinii. Amer.
Fern J. 71: 75-81.
Sen, U. 1964. Importance of anatomy in the phylogeny of
tree ferns and their allies. Bull. Soc. Bot. Bengal. 18: 26-34.
Stokey, A. G. 1930. Prothallia of the Cyatheaceae. Bot.
Gaz. 90: 1-45.
Stolze, R. G. 1974. A taxonomic revision of the genus Cnemidaria (Cyatheaceae). Fieldiana Bot. 37: 1-98.
Tanner, E. V.J. 1983. Leaf demography and growth of the tree
fern Cyathea pubeseens Mett. ex Kuhn in Jamaica. Bot. J.
Unn. Soc. 87: 213-227.
Tardieu-Blot, M.1.. 1941. Sur les aphlebia des Cyatheacees
malgaches. Bull. Soc. Bot. Fr. 88: 522-531.
Tindale, M. D. 1956. The Cyatheaceae of Australia. Contrib.
New S. Wales Nat. Herb. 2(4): 327-361.
Tryon, A. E, Feldmann, I..J. 1975. Tree fern indusia: studies
of diversity and development. Can. J. Bot. 53: 2260-2273.
Tryon, R. M. 1970. The classification of the Cyatheaceae.
Contrib. Gray Herb. 200: 1-53.
Tryon, R. M. 1971. The American tree ferns allied to Sphaeropteris horrida. Rhodora 73: 1-19.
Tryon, R. M. 1976. A revision of the genus Cyathea. Contrib.
Gray Herb. 206: 19-98.
Tryon, R. M.1977. An exchange of views on the Cyatheaceae.
Fl. Males. Bull. 30: 2839-2842.
Tryon. R. M., Gastony, G. J. 1975. The biogeography of endemism in the Cyatheaceae. Fern Gaz. 11 (2/3): 73-79.
Windisch, P. G. 1977. Synopsis of the genus Sphaeropteris
(Cyatheaceae) with a revision of the neotropical exindusiate species. Bot. Jahrb. Syst. 98: 176-198.
Windisch, P. G. 1978. Sphaeropteris (Cyatbeaceae). The syste·
matics of the group of Sphaeropteris hirsuta. Mem. New
York Bot. Gard. 29: 1-22.
Davalliaceae
K.U.KRAMER
Davalliaceae Mett. ex Frank in Leunis, Syn. Pllanzenkd.
ed. 2,3: 1747 (1877).
Gymnogrammitidaceae Ching (1966).
Epiphytic, less often epilithic or terrestrial ferns with
short- to long-creeping, fleshy stem; cortex little sclerotic; stele a peculiar type of strongly dorsiventral dictyostele, only some perforations associated with leaf
traces. Stem densely and permanently clothed in scales
with often cordate or peltate base, the scales often
toothed, sometimes clathrate; surface and/or margin
of scales bearing hairs, these glandular or not; rarely
hairs on stem beside scales. Roots mostly ventral.
Leaves inserted in two alternate rows on the dorsal
side of the stem, on short phyllopodia, with a functional articulation at the base of the petiole. Petiole ±
Filicatae . Davalliaceae
75
F
sclerotic, with two lateral, almost continuous pneumatophores, with two large adaxial and a varying number
of smaller abaxial bundles (in some species only one
or none), all together forming a U or a semi-circle;
petiole usually long, stramineous to brown, adaxially
nattened or more often with a groove, this continuous
with the grooves on the axes of higher order, the middle of the groove usually raised, ridge-like. Lamina
mostly simply pinnate to several times pinnate + pinnatifid, rarely simple, occasionally weakly to strongly
dimorphic; axes adaxially grooved, the grooves with
ruised centre; edges of laminal parts continuous with
Fig.30A-F. Spores
Dryopteridaceae (all
staedtia cieutaria. C
var. arachnoideum.
Pho!. A. F. Tryon
of Davalliaceae, Dennstaedtiaceae, and
x 1(00). A Davallia tenuiseeta. B DennHypo/epis erassa. D Pteridium aquilinum
E Ctenitis vellea. F Dryopteris patu/a.
the ridges (wings) bordering the axis groove; costae
and costules adaxially convex (Fig. 31 G). Lamina often firm in texture, usually triangular, less often narrowed at the base, when mature mostly without macroscopical epidermal appendages; dissection pattern
anadromous or occasionally catadromous or isodro-
76
Filicatae· Davalliaceae
Fig. 31A-K. Davalliaeeae. A-D Sori with receptacles and
vascularization (x 26.5). A,B Araiostegia pu/Chra. CoD Davallia solida. E Araiostegia yunnanensis. relation between sori
and venation (x 2). F Gymnogrammttts dareiformis. the same
( x 2). G Davallia dentieulala. adaxial side of part of rachis
with pinna base (x 3). H-K Davallodes hirsutum. H Hair of
lamina ( x 50). I Transection of petiole ( x 10). J Transection
of stem (x 10). K Stem scale (x 12) (A-D from Sen et al.
1972; E-F from Kato 1985; G from Holttum 1955;
H - K from Perez Arbelaez 1928)
mous. Ultimate free divisions dentate to pinnatifid,
less often entire. Veins pinnately branched, free, ending behind the margin, rarely reaching it; costae adaxially not grooved; "false veins" occasionally present
between the true veins. Stomata polocytic. Sori terminal on the anterior fork of a vein, or situated in the
fork (but one of the branches may be reduced), near
the margin, nearly always uninerval, isodiametric or
longer than wide; indusium attached at the base, often
also at the sides or part of them, rarely reniform, with
a short point of attachment, or absent; outer edge free,
equaling the margin or not; segments not rarely protracted into small projections flanking the sori. Soral
trichomes present or not. Receptacle not elevated;
sporangia long-stalked, the stalk proximally 1- or
2-seriate, distally 3-seriate. Annulus vertical, the indurated part not reaching the stalk nor the stomium.
Spores monolete.
MORPHOWGY AND ANATOMY. This was most fully described for the type species of three important genera
by Sen et aI. (1972). Other important contributions are
by Perez Arbelaez (1928), Nayar and Bajpai (1977),
and Kato and Mitsuta (1980). The cauline stele, in taxa
where it is of complex structure, has been called a perforated dictyostele (but see Kato 1985). The ring of
vascular bundles, as seen in transection, consists of a
large dorsal and a large ventral bundle connected by a
few to many secondary meristeles that form a network
with many areoles and large gaps between them, some
of the dorsal areoles giving off leaf traces (Fig. 311).
The minor meristeles are regarded as homologous to
leaf traces by Kato and Mitsuta (1980).
The fine structure of the articulation between phyllopodium and petiole was described by Phillips and
White (1967).
The false veins occurring in some species of Davallia are homologous with the sclerotic lamina margin,
PiHeatae . Davalliacea.
as interpreted by Iwatsuki and Kato (1976); they are
supposed to be indicative of phylogenetic simplification of the lamina through fusion of divisions; representatives regarded as primitive indeed have in general
a complex lamina structure.
The different types of innervation of the sorus, and
the attachment of the stem scales, and the taxonomic
implications of these characters were described and
discussed by Kato (1975, 1985). The vein branch bearing the receptacle is aborted in some species, the sorus
then appears to be lateral (Davallodes) or at the bottom
of a vein fork (Araiostegia spp., Davallia spp.).
The stem scales often have marginal appendages
consisting of projections of two adjacent cells.
CHARACTERS OF RARE OCCURRENCE. Stem scales with
broad basal attachment, i. e., neither cordate nor peltate, in Leucostegia. where they may also bear hairs.
Amphistomatic leaves reported for a species of Araiostegia. The sori of Gymnogrammitis are exindusiate; in
this genus the large ventral bundle of the stem is also'
lacking. Strongly dimorphic leaves are found in some
species of Davallia. A subcontinuous, multinerval sorus following the margin is present in one species of
Davallia ("Parasorus'').
GAMETOPHYTE. Gametophytes of Davalliaceae were
briefly described by Sen et al. (1972), Atkinson (1973),
and Nayar and Bajpai (1977). They are not very distinctiYe among higher leptosporangiate fern prothallia,
except for the presence of very short, occasionally
branched hairs on the margin and body, and for the
antheridia being sometimes short-stalked.
KARYOLOGY AND HYBRIDIZATION. Not all genera are
as yet karyologically known. The known numbers are
consistently based on n ~40, except for Leucostegia
which has n~41. Polyploidy has occasionally been
found. No hybrids seem to have been found; an apogamous triploid form of "Humata" repens was reported by Manton and Sledge (1954).
ECOLOGY AND DISTRIBUTION. Davalliaceae are essentially epiphytes of tropical and warm-temperate regions. Some often occur on rocks, and Leucostegia,
which may be the most primitive genus, is mostly or
always terrestrial. Some species, especially those with
thin, finely cut leaves, occur in moist, montane forests.
A number of species occurring in strongly seasonal climates shed their leaves in the dry (cool) season.
Dayalliaceae (exclusive of Oleandraceae and NephI'Olepidaceae) are one of the very few larger fern families entirely restricted to the Old World, where they occur in the extreme SW of Europe, in Macaronesia,
tropical and South Africa and its islands, throughout
South and East Asia north to Korea and Hokkaido,
cast to Tahiti and south to SE Australia and the exheme north of New Zealand; they are absent from
Hawaii.
77
AFFINITY. Ideas about affinity with other groups of
ferns with cup-shaped, submarginal indusia, still cited
by Holttum (1955), are no longer tenable in the light of
modem taxonomy. Sen et al. (1972) argued in favour
of affinity to Dryopteridaceae, which is now accepted
by many and supported by karyology. The genus Rumohra has been shifted from one of the two families to
the other; it is here placed in Dryopteridaceae. Genera
like Oleandra and Nephro/epis, here excluded from
Davalliaceae but certainly not far removed, show a
mixture of certain Davalliaceous and Dryopteridaceous features. See also under Oleandraceae.
Similarities to Polypodiaceae (stem histology, peltate scales, petiole articulation, spores), believed at one
time to be indicative of affinity, are now thought to be
due to convergence.
SUBDIVISION AND TAXONOMY. No formal subdivision
of the family has been proposed. The four following
groups, which might be treated as tribes, can easily be
discerned.
LEUCOSTEGIA GROUP: Lamina anadromous, macroscopically glabl'Ous when mature, broadest at base; no
lacunae without leaf traces in stem stele; hairs often
present beside scales on the stem; terrestrial; sori indusiate: Leucostegia.
DAVALLIA GROUP: Lamina anadromous, macroscopically usually glabrous, mostly broadest at base; many
lacunae without leaf traces in the stem stele; stem
bearing only scales; epiphytic or epilithic; sori indusiate: Araiostegia, Davallia.
DAVALLODES GROUP: Lamina catadromous, hairy,
narrowed to base; stem stele as in the last; stem scaly;
epiphytic; sori indusiate: Davallode•.
GYMNOGRAMMITIS GROUP: Lamina anadromous,
broadest at base; stem stele as in the last but the large
ventral bundle absent; stem scaly; epiphytic; sori exin·
dusiate: Gymnogrammitis.
In the past several species were removed from Davallia, mainly on the basis of soral/indusial structure:
Parasorus with subcontinuous marginal ("coeno")sorus; Humata with laterally free indusia and leaves of
simpler structure, etc. Various authors have commented on the artificiality of such genera, and they are here
sunk in Davallia. The spore morphology bears out this
classification.
Kato (1985), though supporting the general taxonomy as adopted here, attempted to subdivide Davallia
anew and recognized five genera. Parasorus remains a
"single character genus"; the other segregates can be
defined by combinations of characters only, such characters being virtually all found in Davallia s. str. These
genera are therefore not recognized here, either. Even
the distinctness of Araiostegia on the generic level is
open to some doubt.
78
Filicatae . Davalliaceae
KEy TO THE GENERA.
1. Lamina always compound, catadromous, somewhat to
strongly narrowed at the base; at least the axes of mature
leaves hairy
2. DavaUodes
- lamina anadromous, rarely isodromous or exceptionally
catadromous, or simple, not narrowed to the base except
when simple, with few exceptions glabrous when mature 2
2. Sori exindusiate; spores c1avate-baculate
5. Gymnogrammitis
- Sori indusiate; spores smooth to rugose or verruculose,
never clavate-baculate
3
3. Stem scales attached along a basal line; hairs often present
on stem beside scales. Stele of stem with only two large
meristeles that are connected by short, simple, lateral ones.
Sori terminal
1. Leucostegia
- Stem scales attached at a point, cordate or peltate; hairs on
stem lacking. Stele of stem with many branched lateral
meristeles connecting the large dorsal and ventral ones. So-
ri at forking or "bending" point of veins (but often appearing terminal)
4
4. Lamina much dissected, thin in texture, glabrous or bear-
ing short. pluricellular hairs; scales inserted on a small
stalk, cordate; indusium nearly always free at the sides
3. Araiostegia
- Lamina much dissected to simple, firm in texture, macroscopically glabrous; scales peltate or cordate, not stalked;
indusium laterally free or attached
4. DavaJ/iJz
Medium-sized epiphytic ferns; stem creeping, fleshy,
with a stele typical of the family; roots confined to
short, detenninate side branches associated with a
phyllopodium. Scales peltate, with broad base, the apical part abruptly contracted, deciduous, with indurated cell walls; apex glandular. Petiole well-developed,
the base with three vascular bundles, these upward
fusing into one U-shaped one; adaxial side grooved,
± persistently hairy, the hairs rather short, severalcelled, simple or branched, glandular or not. Lamina
pinnate + pinnatifid or bipinnate + pinnatilobate, thinly herbaceous, somewhat to strongly narrowed to the
base, catadromous; upper pinnae and segments reduced, confluent. Rachis adaxially convex or with two
dorso-Iateral grooves, persistently hairy like the other
leaf axes; veins and often lamina tissue with hairs like
those of the petiole. Pinnae oblong, not deltoid; segments (pinnules) much decurrent, dentate-lobed. Sori
typical of the family; indusium renifonn to pouchshaped, attached at the base or also laterally. Spores
ellipsoidal, low-verrucate.
About 10 species, epiphytic in moist places, from
Sumatra to the Philippines and New Guinea.
3. Araiostegia Copeland
Leucostegia C. Presl, Tent. Pteridogr.: 94 (1836); Copeland
(1927); Holttum (1955).
Fig.31H-K
Daval/odes Copeland, Philipp. J. Sci. 3: 33 (1908); Copeland
(1927); Holttum (1972).
ParadavaJ/odes Ching (1966), p. p., excl. type.
land (1931); Tagawa and Iwatsuki (1970); Sen, Sen and
Holttum (1972); Kato (1975).
Paradaval/odes Ching (1966), p. p., quoad typo
Medium-sized, usually terrestrial ferns. Stem ± longcreeping, thick, fleshy; stele as described in the key;
tannin cells present; roots borne on all sides; scales
non-peltate, sometimes setiferous; hairs often present
besides scales. Leaves (sub)tripinnate to quadripinnate + pinnatifid, glabrous when mature, anadromous, herbaceous-chartaceous; petiole long, stramineous, adaxially grooved like the other leaf axes.
Upper pinnae, pinnules, etc. gradually reduced and
confluent. Ultimate free divisions rhombic-Ianceolate,
dentate to pinnatifid, edge thickened at the base and
decurrent as a wing onto the edge of the rachis. Veins
immersed, evident, simple or forked, ending well behind the margin. Sori often impressed; indusium renifonn-suborbicular, shortly basally attached, or suborbicular and attached also by the basal half of its sides,
delicate, reaching the margin or not; spores ellipsoidal, coarsely verrucate, also a thin outer layer present
that fonns a flaky cover. Two species, from S. China,
the central Himalaya, and S. India to E. Polynesia.
2. Davallodes Copeland
Fig. 31A, B, E
Araiostegia Copeland, Philipp. J. Sci. 34: 240 (1927); Cope-
1_ Leuctmegia C. Presl
Medium-sized, epiphytic or epilithic ferns with creeping, fleshy stem bearing thin, basally cordate, hairy,
non-acuminate scales; roots borne only ventrally; stele
typical of the family. Petiole well-developed, adaxially
grooved, with three vascular bundles that higher up
fuse into one; at least the secondary rachises with a
raised portion in the middle of the groove. Lamina
tripinnate + pinnatifid to quadripinnate + pinnatifid,
anadromous, rarely catadromous or isodromous, not
narrowed at the base, thin, in one species arnphistomatic; surface glabrous or pUberulent with short, simple, or branched, sometimes glandular hairs, persistently scaly on the axes; pinnae short-stalked or sessile. Veins evident, forked, not reaching the margin.
Ultimate divisions mostly small, narrow, acute; sori as
typical of the family, terminal on the anterior branch
of a vein, the other branch serving a lobe flanking the
soruS. Indusium delicate, suborbicular or reniform, attached at the base (in one species also laterally), not
reaching the margin. Spores ellipsoidal, coarsely verrucate, with a thin surface covering of papillate or compressed rodlets.
Eleven species (Tagawa and Iwatsuki 1970), growing in moist, wann-temperate to tropical or montane
forests; from the Himalaya and S. China to southern
India, Taiwan, and Malesia. Seven more species were
Leucostegia' Davallodes· Araiostegia • Davallia
transferred by Kato (1975) from Davallia on account
of the scale attachment; the borderline between the
two genera (if any) needs to be critically re-assessed.
4. Dava/liaJ.
E. Smith Figs. 30, 31C,D,G, 32A,B,E,F
Davallia J. E. Smith, Mem. Acad. Thrin 5: 414 (1793); Copeland (1940); Morton (1957); Joe-Hoshizaki (1981); Kato
(1985).
Burnata Cav. (1802); Copeland (1940).
Pachypleuria (c. Presl) C. Presl (1849); Kato (1985).
Scyphufllrla Fee (1852).
Parasorus v. A. v. R. (1922).
Trogostolon Copeland (1927).
Small to rather large, epiphytic or epilithic ferns with
creeping, fleshy stem with a stele typical of the family,
bearing peltate scales with marginal teeth formed by
projections of two adjacent cells, glandular or not,
bearing superficial hairs or not, not rarely differentiated into a roundish-oblong basal and a narrowed,
sometimes bristle-like apical portion which may be deciduous. Petioles usually well-developed, in two ranks
79
Fig. 32A-F. Davalliaceae. A,B Davallia trichomanoides.
A Stem with leaf (x 0.3). B Leaf segment with sori (x 2).
C,D Gymnogrammitis dareiformis. C Part of rachis with pinna (x 1). D Scale of stem (x 25). E,F Davallia pentaphylla.
E Stem with leaf (x 0.7). F Part of pinna with sori (x 2.5)
(A,B,E,F from Backer and Posthumus 1939; C,D from
Tardieu-Blot and Christensen 1939)
on the dorsal side of the stem the ventral side of which
bears roots, with two larger dorsal and three smaller
ventral bundles, these uniting upward; adaxial face
sulcate, the groove usually raised in the middle; small
species sometimes with subsessile leaves; petiole occasionally persistently scaly. Lamina large, deltoid-oblong, or small, pentagonal or lanceolate, occasionally
dimorphic, anadromous or rarely isodromous, tripinnate + pinnatifid or bipinnate + bipinnatifid to simple,
mostly firm in texture, macroscopically glabrous when
mature but bearing minute, few-celled, glandular hairs
when young which may persist in places. Edge of segments decurrent as wings onto axes. Fertile lamina of
80
Filicatae . Davalliaceae
dimorphic species with reduced leaf tissue andlor
more dissected. Veins mostly evident, forked, ending
behind, or in a few species at the margin; in a few species sclerotic lines ("false veins") interspersed between
true veins. Ultimate divisions linear to lanceolate or
ovate, toothed to pinnatifid; in some small species the
lowest pinnae basitonically produced. Sori near the
margin, the vein branch bearing them sometimes very
short, the next higher vein forked, giving off a branch
to a tooth which is often hom-like and flanks the sorus. Indusium attached at the base or also at the sides
or part of them, thus from shell-shaped to pouchshaped, in the latter case often longer than broad,
± firm, reaching the margin or not; in a single species
a continuous sorus on many to all vein ends following
the margin of the entire fertile lamina, the indusium
then resembling that of Lindsaea. No sterile appendages among the sporangia. Spores ellipsoidal, coarsely verrucate, tuberculate, or rugose, a thin outer layer
or spherical deposit sometimes present.
As defined here, in a broad circumscription, about
90 species in tropical and oceanic-temperate areas of
Asia, from the Himalaya and northern Japan to Australia and Tahiti; only two species in Africa and Madagascar; one in NW Africa, Macaronesia, and SW Europe.
S. Gymnogrammitis Griffith
Fig. 31 F,32C,D
Gymnogrammitis Griffith, Icon. Plant. Asiat. II: pI. 129
(1849).
Araiostegia sensu Copeland (1927, 1931, 1947). p. p. min.,
excl. type.
Epiphytic ferns with creeping, pruinose stem containing tannin cells; stele as typical of the family but lacking the large ventral meristele. Scales cordate, lanceolate, dentate-ciliate. Petiole well-developed, with two
large and three small bundles. Lamina tripinnate +
pinnatifid, anadromous, deltoid, thin, the ultimate segments Iinear-Ianceolate, acute; larger axes grooved in
the middle. Sori on short or completely reduced vein
branches, thus sometimes seemingly dorsal, flanked by
one or two extended branches, roundish, exindusiate,
without sterile appendages. Spores ellipsoidal; surface
low-papillate to rugose, bearing scattered rodlets and
spherical deposit.
A single species, G. dareifonnis (Hooker) Ching in
Tardieu-B1ot and C. Chr., from the eastern Himalaya
and S. China to Indo-China; epiphytic in montane
forests at middle and higher elevation.
A derived element in the family, probably closest to
Araiostegia. with which it has been confused. In spite
of the unique spore structure and the naked sori the
genus fits the family quite well.
Selected Bibliography
Ching, R. C. 1966. Gymnogrammitidaceae Ching, a new fem
family. Acta Phytotax. Sin. 9: 11-16.
Copeland, E. B. 1927. Daval/odes and related genera. Philipp.
J. Sci. 34: 239-256.
Copeland, E. B. 1931. Miscellaneous oriental pteridophytes.
Univ. Cal. Publ. Bot. 12: 383-407.
Copeland, E. B. 1940. Oleandrid ferns (Davalliaceae) of New
Guinea. Philipp. J. Sci. 73: 345-356.
Holttum, R. E. 1972. The genus Daval/odes. Kew Bull. 27:
245-249.
Iwatsuki, K., Kato, M. 1976. Evolution of fern leaves through
fusion. Phytomorphology 26(3): 234-239.
Hoshizaki, B. Joe 1981. The fern genus Davallia in cultivation; Davallia relatives in cultivation. Baileya 21: 1-42;
43-50.
Kato, M. 1975. Taxonomic notes on the cordate scales in Davallia. Acta Phytotax. Geobot. 26: 152-159.
Kato, M. 1985. A systematic study of the genera of the fern
family Davalliaceae. J. Fac. Sci. Univ. Tokyo III. 13:
553-573.
Kato, M., Mitsuta, S. 1980. Stelar organization in davallioid
ferns. Phytumorphology 29: 362-369.
Manton, I., Sledge, W. A 1954. Observations on the cytology
and taxonomy of the pteridophyte 110ra of Ceylon. Philos.
Trans. R. Soc. London, Ser. B 238: 127-185.
Morton, C. V. 1957. Observations on cultivated ferns. IV. The
species of Davallia. Amer. Fern J. 47: 143-148.
,Nayar, B. K., Bajpai, N. 1977. Morphology in relation to phylogeny of the Davallioid-Oleandroid group of ferns. Phytomorphology 26 ("1976") 333-354.
Perez Arbelaez, E. 1928. Die natfirliche Gruppe der Davalliaceen (Sm.) Kfs. Bot. Abh. (Jena) 14: 1-96.
Phillips, D. A, White, R. A 1967. Frond articulation in species of Polypodiaceae and Davalliaceae. Amer. Fern J. 57:
78-88.
Sen, T., Sen, U. 1971. Morphology and anatomy of the fern
genus Gymnogrammitis. Ann. Bot. (London) II, 35:
229-235.
Sen, T., Sen, U., Holttum, R. E. 1972 Morphology and anatomy of the genera Davallia, Araiostegia and Daval/odes,
with a discussion on their affinities. Kew Bull. 27: 217-243.
Tagawa, M., Iwatsuki, K. 1970. New or interesting ferns from
Thailand 6. 39. Araiostegia. Acta Phytotax. Geobot. 24:
178-181.
Filicatae . Dennstaedtiaceae
Dennstaedtiaceae
K. V.KRAMER
Dennstaedtiaceae Pichi Sermolli, Webbia 24: 704 (1970).
Hypolepidaceae Pichi Sermolli, Webbia 24: 705 (1970).
Lindsaeaceae Pichi Sennolli, Webbia 24: 707 (1970).
Pteridiaceae Ching, Acta Phytotax. Sin. 13: 96 (1975).
Pteridaceae sensu Copeland (1947), p. p.
Terrestrial or epilithic, occasionally epiphytic ferns;
stem usually long- or short-creeping, with a solenostele
or a protostele with internal phloem, rarely short and
dictyostelic, bearing articulate hairs, or rarely peltate,
non-clathrate, entire scales, or both, with transitions.
Petioles close or more often remote, non-articulate,
usually with one or two, less often several vascular
bundles. Leaf axes usually adaxially sulcate, with continuous grooves; mature leaves often glabrous at maturity, or bearing (mostly articulate) hairs, not scaly;
hairs often persistent on leaf axes. Lamina often bi- or
tripinnate to decompound, less often simply pinnate,
rarely simple, mostly distinctly anadromous. Veins
free, ending with ± thickened ends behind the margin,
less often anastomosing, free included veinlets none.
Stomata anomocytic and/or polocytic. Sori essentially
terminal on a vein but often on a submarginal commissure joining two to very many vein ends; sterile appendages among sporangia none or if present not or
little differentiated from epidermal appendages. Indusium various, shell- to pouch-shaped, or linear in taxa
with elongate sori borne on a commissure; or wanting
and replaced by the reflexed, ±modified leaf margin;
sometimes such a "false" indusium present beside a
"true" one. Sporangial stalk triseriate, capsule usually
with well-differentiated stomium; spores monolete or
trilete, variously sculptured.
ANATOMY AND MORPHOLOGY. See under the subfamilies.
KARYOLOGY AND HYBRIDIZATION. See under the genera.
GAMETOPHYTE. This is known in comparatively few
representatives; the variability may be larger than
thought at present. The prothallia are hairless, cordate,
and the plate stage is reached soon after germination.
See Nayar and Kaur (1971) and Atkinson (1973). The
sex organs are of the advanced leptosporangiate type.
EmLOGY AND DISTRIBUTION. The family occurs
worldwide, but much of this is due to the enormous
area of one genus, Pteridium. Of 16 genera, only 6 are
confined to one or two continents, the majority is
again worldwide, though largely confined to warmer
regions. Several genera extend, however, into warmtemperate or even cool-temperate areas. Only a few
genera, like Oenotrichia, Leptoiepia, and Ormoioma,
81
are more narrowly distributed. The family is certainly
geologically old.
Many species are confined to the ground of moist,
undisturbed forests, like representatives of Paesia, Biotiella, Lindsaea, and Tapeinidium. Others are colonizers of open places, are endowed with long-creeping,
much-branching stems, and thus become troublesome
weeds in clearings and disturbed habitats, especially
species of Hypoiepis, Histiopteris, and Pteridium (see,
e.g., Gruber 1981). A few members of the family are
sprawling climbers ("Spreizklimmer") whose leaves
climb by getting entangled in the surrounding vegetation: some species of Odontosoria, Dennstaedtia, Hypoiepis, Histiopteris, etc. Spiny leaf axes occur in some,
especially scandent, species of Hypoiepis, Dennstaedtia, and Odontosoria, a comparatively rare character.
Epiphytes are few; see under Lindsaea and Denn-
staedtia.
. AFFINITY AND SUBDIVISION. Concepts about the affinity of the genera and genus groups united here in one
family have greatly varied, and continue to do so.
Those with short sori and an approximately pouchshaped indusium have mostly been associated with the
Davaliia group (see especially perez Arbelaez 1928, for
discussion and review), those with elongate sori and a
"false" indusium with Pteris, or even included in it
(see Page 1976, for a more general review). Copeland's
(1947) very broadly circumscribed Pteridaceae go back
to Bower's ideas but are recognized by hardly any contemporary worker. Alston (1956) was the first to recognize the Dennstaedtioideae in their present circumscription, to which little was added by Mickel (1973).
Association of the Lindsaeoideae with Dennstaedtiaceae was first suggested by Holttum (1947); genera regarded as primitive in both groups are fairly close, derived taxa diverging much more. Tryon and Tryon's
(1982) treatment of the subordinate groups as tribes
tends to underrate their differences.
KEy TO THE GENERA.
1. Sori with an outer, "false" indusium fonned by the reflexed and ± modified leaf margin, or with an outer,
"false", and an inner, "true" indusium (Fig. 34A, B); rarely indusium none
2
- Sori only with an inner, "true" indusium, this never wanting; leaf margin opposite the indusium not reflexed and
not or hardly modified, though sometimes laterally fused
with the indusium
8
2. Veins regularly and copiously anastomosing, or at least
forming regular costal! costular areoles; spores monolete
3
- Veins free or with occasional anastomoses; spores various
4
3. Stem slender, long-creeping; mature leaves without macroscopic hairs
10. Histiopteris
- Stem massive, ascending-erect; mature leaves thinly to
densely hairy at least on the veins beneath, rarely glabrous
9. BlotJella
82
Dennsta~dtiaceae • Saccolomatoideae . Dennstaedtioideae
4. Sori short, on single veins, protected by a short, about semi-orbicular, marginal, reflexed flap. or this vestigial and
the sori naked; spores monolete
- Spores monolete; receptacular appendages pluricellular,
evident; sori on one or two, rarely 3 vein ends; Old World
8. Hypolepis
13. Tapeinidium
- Sori long, on several to many veins, the outer indusium
16. Ultimate free divisions lanceolat.. equal-sided or auricled
at the anterior base; veins free except as joined by the receptacle; spores monolete; receptacular appendages pluricellular, evident; Malesia
14_ Xyropteris
(reflexed margin) occupying most or all of the margin of a
5
fertile segment
5_ Spores trilete; petiole with two or more vascular bundles 6
- Spores monolete; petiole with a single vascular bundle 7
6. Petiole with two vascular bundles; margin oflamina only
reflexed over the sori; rachises without dark, callose spots
- Ultimate free divisions not lanceolate and equal-sided, or,
if so, the spores trilete; veins free or reticulate; widespread
15. Lirulsaea
11. Lonchitis
- Petiole with several vascular bundles; margin of lamina
always reflexed and ± modified, even in the absence of
sori ("sterile indusium"); dark, callose spots present on
rachis at insertion oflarger pinnae
6. Preridium
7. ··Inner" indusium present beside the "outer", often con-
spicuous; pirmae alternate
7. Paesia
- Only an "outer" indusium present; at least the larger pin10. Histiopteris
nae and pirmules (sub)opposite
8. Stem ascending-erect, dictyostelic, scaly; leaf axes glabrous; sori on single veins
1. Saccoloma
- Stem creeping or scandent, protostelic or solenostelic,
hairy or scaly; sori on several veins, or, if on single veins,
the leaf axes often hairy
9
9_ Stem solenostelie, bearing (often reddish) articulate hairs,
which are usually also present on leaf axes; sori on single
veins
10
- Stem protostelic with internal phloem (very rarely dictyostelic), bearing (occasionally very narrow) scales; leaf
axes nearly always macroscopically glabrous; sori on sin-
gle or more often on two to several vein ends
13
10_ Indusium falling short of the margin, the laminallobe or
portion of the lamina opposite it scarcely or not modified
12
- Indusium equaling the margin or nearly so, the adaxial
lobe or portion of the lamina opposite it modified or not
11
11. Stem short-creeping, with approximate leaves; indusium
attached at the base only, reflexed at maturity
3. Oenotrichia
- Stem usually long-creeping, bearing remote, mostly large
leaves; indusium attached at the sides as well as at the
base, not reflexed at maturity, forming a cup-like structure
with the part of the lamina opposite it
2. Dennstaedtia
12. Indusium attached at the base and sides, only the outer
edge free
5. Micro/epia
- Indusium attached only at the base, laterally free
4. Leptolepia
13. Sori on 1-8 vein ends; indusium laterally largely to entirely adnate to the lamina; ultimate divisions never dimidiate; veins free except as joined by the receptacular commissure (if any)
14
- Sori on many vein ends, or, if on 8 or less, its sides free, Of
the pinnules dimidiate, or the veins anastomosing. Of
combinations of these characters present
16
14. Ultimate (nearly) free divisions of a linear or cuneate-di-
varicate type, with the soruslsori on the apical margin;
receptacular appendages usually small, few-celled, or
wanting
12. Odontosoria
- Ultimate (nearly) free divisions not of a linear or cuneatedivaricate type; sori on the lateral margin; receptacular
appendages various
15
15_ Spores trilete; receptacular appendages none or fugacious; sori strictly uninerval; neotropical
16. Ormoloma
Subfamily Saccolomatoideae
Stem short-creeping to erect, dictyostelic, bearing only
scales; macroscopical hairs wanting, lamina axes glabrous. Sori tenninating single veins.
Monotypic:
1. SaccoloflUl Kaulfuss
Fig.33A
Saccoloma Kaulfuss, Berl. lahrb_ Pharm. 1820: 51.
Orthiopterls Copeland (1929).
IthycauJon Copeland (1929).
Stem short-creeping to erect or somewhat trunk-like,
terrestrial, dictyostelic, scaly, the scales sometimes peltate (Nair 1979)_ Petioles arising on all sides, well-developed, with a single vascular bundle and two lateral
pneumatic lines. Lamina glabrous, usually decompound with pinnatifid apex, in one species simply pinnate and with an odd tenninal pinna; dissection anadromous (or isodromous in S. elegans); rachis adaxially sulcate, the lateral ridges bordering the groove
continuous with ridges bordering the costae of the ultimate divisions; grooves continuous with one another
or separated by shallow sills at junctions_ Veins free,
simple or forked, not reaching the margin. Sori terminal on single veins, with baso-Iaterally attached, cupor pouch-shaped indusium that sometimes almost
equals the margin. Lamina opposite the indusium not
modified to slightly modified_ Spores trilete, tetrahedral-globose, with prominent, ± parallel, branched
ridges, these often partly obscured by a granulate deposit
Chromosome counts of 2n = 376 and 2n = c. 63 for
two different neotropical species are puzzling.
C. 12 species, pantropical but lacking in continental
tropical Africa; most numerous in eastern Malesia and
the western Pacific, only three species in the New
World and one in Madagascar_
The leaf architecture of the type species, the South
American S_ elegans Kaulf., with a simply pinnate
lamina with confonn tenninal pinna, diverges from
that of the other species that are decompound and
have been split off as Orthiopteris and Ithycaulon.
R_ M. Tryon (1962) has shown the close agreement in
other characters. Sterile leaves of the decompound
species are very much like those of Dennstaedtia and
Microlepia but the absence of articulate hairs is distinctive_
Saccoloma • Dennstaedtia
83
~
"
'"
":OC:-
'-
A
C
Subfamily Dennstaedtioideae
Stem (mostly long-)creeping, solenostelic (occasionally
polycyclic), bearing articulate, often reddish hairs
(Fig. 35 C). Petioles borne only dorsally on the stem,
usually with one gutter-shaped, or else with two adjacent, upward merging vascular bundles (see Keating
1968). Upper pinnae, pinnules, etc. confluent; veins
free, or less often anastomosing without included veinlets. Distinction of certain genera in sterile state may
be difficult. Sori on one to very many vein ends.
Epipetiolar buds or stem branches have been described for Dennstaedtia, Hypo/epis, Paesia, and Histiopteris (Troop and Mickel 1968) and may be more
widespread. Hagemann (1976) stressed the unusual
orientation of the leaves on the stem in some genera,
where the adaxial face of the petiole is in a line with
the longitudinal axis of the stem; he concluded that
such leaves are essentially non-appendicular.
The petiole bears on each side a lateral band of
aerating tissue that continues onto the stem and may
be elevated. The stomatal pattern passes from predominantly polocytic into entirely anomocytic; its taxonomic relevancy seems slight.
For chromosome numbers see under the genera.
2. Dennstaedtia Bernhardi
Fig.33B-E
Dennstaedtia Bernhardi, Schrad. J. Bot. 18002: 124 (1801);
Conard (1908); R. M. Tryon (1960).
Costaricia Christ (1909).
Coptodipteris !"Coptidipteris'1Nakai and Momose (1937).
Fuz!filix Nakai and Momose (1937).
Paradennstaedtia Tagawa (1952).
Emodiopteris Ching (1978).
Fig. 33A-E. Dennstaedtiaceae. A Saccoloma elegans. pinna
margin with sori ( X 4). B, C Dennstaedtia bipinnata. B Transection of stem (x 5). C Transection of petiole (x 5). D Dennstaedtia globulifera, fertile pinnnie (x 3.5). E Dennstaedtia
dissecta, fertile segments (x 7) (A from Bower 1926;
B,C from Perez Arbelaez 1928; D,E from Stolze 1981)
Terrestrial ferns with (usually long-)creeping, solenostelic, sometimes polycyclic stem bearing yellow or
reddish, articulate hairs; petioles usually well-spaced,
adaxially sulcate, well-developed, in a few species the
basal part with spine-like processes. Lamina usually
de compound, in a few species simply pinnate, anadromous or the primary divisions weakly catadromous,
sometimes scandent; primary rachis occasionally
spiny. Rachises of various order adaxially sulcate,
bearing articulate hairs or glabrescent. Grooves on rachises of higher order continuous with those of lower,
or incompletely so, or fading on the lateral ridges below the junctions. Edges of ultimate leafy divisions decurrent as wings onto their rachises. Segments usually
obtuse and lobed to pinnatifid, often asymmetric, the
posterior side less deeply or not lobed. Veins free, suppinnate in the ultimate divisions, not reaching the margin, often with thickened ends. Stomata predominantly polocytic. Sori terminal on single veins, in a few
species on marginal lobes, often as broad as long or
broader. Indusium with convex base, free only on the
distal side, equaling the leaf margin, the part of the
lamina opposite it ± modified, the sorus thus cupshaped, often pointing downward from the plane of
84
Dennstaedtiaceae • Dennstaedtioideae
the lamina. Spores trilete, tetrahedral-globose to globose, tuberculate, ridged, or reticulate [with coarse,
discrete tubercles in "Coptodipteris" wilfordii (Moore)
Nakai and Momose].
A confusing array of chromosome numbers has
been found in this genus: n=30, 33 (347),34,46,47,
60, 64, 65, 94, and 2n numbers based on these. Walker
(1973) attempted to connect these numbers by invoking aneuploid changes and allopolyploid additions,
and arrived at base numbers of 15, 16, or 17. Lovis
(1977) claimed that a single base number, e. g., 16, with
subsequent aneuploid and (allo)polyploid changes,
could account for the various numbers known. The ge- .
nus may even turn out to be artificial. No hybrids have
been reported. Sterile leaves are often confusingly similar to those of Microlepia. Saccoloma. and even Hypolepis.
CHARACTERS OF RARE OCCURRENCE. Articulate primary pinnae in a few species; simply pinnate leaves in
a small number of species; small, delicate, crowded
leaves in D. wilfordii (Moore) Koidz. ex Tagawa
("Coptodipteris"); a scandent to epiphytic stem in "Costarida" werclcleana Christ (to be included in Dennstaedtia); an erect stem with approximate leaves in
D. ("Emodiopteris") appendiculata (Hooker) J. Sm.
An estimated 45 species, but the taxonomy is quite
inadequately known; many species are much alike,
and many herbarium specimens are grossly inadequate, consisting only of fragments of the lamina. The
concentration of species is very uneven; most occur in
tropical to warm-temperate parts of the world, but the
genus is weakly represented or lacking in large areas
like the Amazon Basin, the Malay Peninsula, and continental Africa. Regional revisions are available for only small portions of the area (e. g., Tryon and Tryon
1982; Copeland 1950, 1958; etc.). A few species occur
in north-temperate regions, e. g., D. punctilobula
(Michx.). Moore in eastern North America, D. hirsuta
(Sw.) Mett. in NE China and northern Japan. A modern subdivision of the genus is yet to be given.
3. Oenotrichia Copeland
Fig. 341, J
Oenotrichia Copeland, Univ. Cal. Publ. Bot. 16: 82 (1929).
Very similar to Dennstaedtia and Microlepia. and apparently closely related to both; differing chiefly by
subreniforrn, only basally attached indusium, and by
the presence of receptacular appendages. Spores
trilete, tetrahedral-globose, bearing prominent,
± anastomosing ridges that may be partly obscured
by granulate deposit.
Tho species in New Caledonia, Oe. maxima
(Fourn.) Copel. and Oe. macgillivrayi (Fourn.) Brownlie; other species ascribed to the genus are misplaced.
4. Leptolepia Met!.
Leptoiepia Mett. [ex Kuhn, Chaeopt.: 348 (1882)1 ex Diels in
Engler und Pranti, Nat. Pfl. Fam. L 4: 212 (1899).
Stem terrestrial, long-creeping, solenostelic, bearing
reddish articulate hairs. petioles remote, long, adaxialIy sulcate, with a single vascular bundle. Lamina tripinnate + pinnatifid, anadromous, glabrous, herbaceous; segments acute. Sori terminal on single veins,
small, roundish, close to but distinctly behind the margin; indusium ovate-subreniform, attached at the truncate or slightly concave base only, laterally free or
slightly adnate to its vein; outer edge lacerate. Spores
trilete, tetrahedral-globose, bearing coarse, compact
verrucae.
A single species, L. novae-zelandiae (Col.) Mett. ex
Diels, in New Zealand, also reported from New Guinea and Queensland, perhaps incorrectly so. Closely
related to Dennstaedtia and Microlepia; see the key.
The chromosome number of n = 47 also separates it.
S. Microlepia C. Presl
Fig. 35
Microiepia C. Presl, Tent. Pteridogr.: 124 (1836).
Medium-sized to large ferns with terrestrial, mostly
long-creeping, solenostelic stem clothed with articulate
hairs; petioles mostly remote, lacking basal buds,
adaxially grooved. Lamina bipinnate to decompound,
rarely simply pinnate, anadromous; rachises adaxially
grooved, the grooves continuous, or more often fading
on the lateral ridges of grooves of lower order, below
the junction. Axes with a short tomentum of reddish,
articulate hairs; longer, pale, also articulate hairs often
present besides. Stomata predominantly polocytic.
Pinna-bases non-articulate; leaf axes spineless. Upper
pinnae, pinnules, etc. gradually reduced and confluent. Ultimate segments much as in Dennstaedtia, also
in venation, their edges posteriorly thickened and continuous with the lateral ridges on their axes. Costae
slightly raised, not grooved. Sori terminal on single
veins, usually not very near the margin, but in any case
the indusium laterally not fused to the lamina lobe opposite it to form a cup; indusium pouch-shaped, attached at the convex base and the sides, sometimes
hairy. Spores trilete, tetrahedral-globose, usually with
prominent angles, echinate; rodlets often connected by
strands, then forming a reticulum.
Karyologically the genus is not quite as diverse as
Dennstaedtia, most counts being based on x =43 or
being close to multiples of this number. Reports of
2n = 84 and 160 also exist but are perhaps erroneous.
Hybrids have not been described.
CHARACTERS OF RARE OCCURRENCE. A simply pinnate lamina with subentire, auriculate pinnae in
M. hookeriana (Wall. ex Hooker) C. Presl; M. tenera
Oenotrichia • Lepto/epia . Micro/epia • Pleridium
85
B
'iI!
.
I
~
.
A
!
G
Fig. 34A-J. Dennstaedtiaceae. A Pleridium aquilinum. part
of segment from abaxial side showing "true" and "false" indusium, the latter reflexed at right (x 10). B Paesia acciivis,
two fertile segments from abaxial side showing sori with outer and inner indusium (x 2). C Hypo/epis repens, stem with
aerating band (ae) and lateral branches on petiole bases (s)
( x 0.3). D- F Hypo/epis rugosula. D Base of pinna (x 0.8).
E Fertile segment ( x 3). F From left to right hairs from lamina, from rachidule, and (2) from rachis (x 16). G,H Hypolepis punctata. G Fertile segment ( x 3). H Hairs, from lift to
right from lamina, rachidule, and rachis (x 16). ( Oenotrichia
maxima, part of fertile pinnule (x 4). J Oenotrichia macgillivrayi, part of fertile pinnule (x4) (A from Hyde et aI.1969;
B from Mettenius 1858; C from Hagemann 1976; D-H from
Duncan and Isaac 1986; (,J from Brownlie 1969)
Christ, described as having only basally an attached
indusium, is perhaps misplaced.
An essentially Asiatic-Pacific genus, with only one
species, M. speluncae (L.) Moore, in Africa and the
Neotropics; in the latter area it may not be native
(Tryon and Tryon 1982), but as it is very widespread
and euryoecious in the Old World, it may have crossed
the Atlantic naturally; it is also the only species reaching Australia. Otherwise the genus occurs in tropical to
warm-temperate Asia, where there are an estimated
45 species; but critical revisional work on the genus is
much needed. The genus seems to be more natural
than Dennstaedtia. Most species occur on the floor of
rorests, in thickets, etc. and some have leaves several
m long that are supported by the surrounding vegetation.
6. Pteridium Scopoli
Fig.34A
Pleridium Scopoli, A. Carniol. 169 (1760); R. M. Tryon
(1941); Perring and Gardiner (1976).
Eupteris Newman (1845).
Pleris auctt., p. p. min.
Medium-sized to large terrestrial ferns with very longcreeping and much branched and vegetatively spreading stem with two concentric, much corrugated solenosteles; differentiation of long, branching, and
short, leaf-bearing shoots; indument of multicellular
hairs. Petioles remote, long and stout, pale, with several vascular bundles arranged in a horseshoe pattern
in cross-section. All axes adaxially grooved, the
grooves± hairy; grooves on axes of higher order continuous with those of lower but often very shallow at
Dennstaedtiaceae . Dennstaedtioideae
F
Fig. 35A-F. Dennstaedtiaceae. A-E Micro/epia hookeriana.
A Part of lamina (x 1). B Part of fertile pinna, abaxial side
( x 3). C Transection of stem (x 10). D Transection of petiole
( x 10). E Trichome of lamina ( x 20). F Microlepia platyphylla, margin of fertile segment from abaxial side (x 5) (A-E
from Ching et al. 1959; F from Perez Arbelaez 1928)
the junctions. Lamina in large plants developing slowly and stepwise; dissection anadromous or catadro·
mous, but usually constant in a single lamina, at least
bipinnate+pinnatifid, up to 4-pinnate; primary pinnae usually subopposite, often strongly spreading, at
least the larger with a dark callose spot on the rachis
near their base which in developing leaves functions as
a nectary. Upper pinnae, pinnules, etc. gradually reduced and confluent. Penultimate divisions pinnately
compound or lobed, often with a long, entire apical
portion; segments oblong, obtuse, adnate, often surcurrent and! or decurrent, often interspersed with
much smaller, broad and short lobes. Leaf tissue
usually firm, variously hairy to g1abrescent. Costae
and costules adaxially sulcate, abaxially flattened and
usually also broadlyandsnallowly -grooved.' veins
close, forked, free; margin entire, always revolute and
± modified even in the absence of a sorus ("sterile indusium"). Sori occupying all vein ends, borne on a
commissure, in addition to the outer indusium a membranous, occasionally rudimentary or obsolete inner
indusium present (Fig. 34A). Receptacular append·
ages none. Spores trilete, tetrahedral-globose, irregularly granulate.
According to judgment a single, variable, subcosmopolitan species, Pt. aquilinurn (L.) Kuhn in v. d.
Decken, with two subspecies and a number of
± geographically vicariant varieties; or a small number of distinct species. As the various forms intergrade
where they occur together, treatment as a single species is preferred here, as advocated by R M. Tryon
(1941), Page (1976), and Tryon and Tryon (1982).
The great majority of chromosome counts yielded
n = 52; 2n = 52 and 2n = 208 have been found on single occasions.
7. Paesia St. Hilaire
Fig.34B
Paesia Sl. Hilaire, Voy. Distr. Diam. 1: 381 (1833).
Medium-sized to large, terrestrial or epilithic ferns
with long-creeping, solenostelic stem bearing reddish-
Paesia • Hypo/epis
brown pluricellular trichomes. Leaves remote; petiole
well-developed, with a single vascular bundle, like the
other leaf axes brown to blackish, adaxially with a
groove that may be very shallow, hairy like the stem or
glabrescent; bases of trichomes often like minute
warts, the leaf axes then scabrous. Lamina anadromous (except in one species), bipinnate or more often
tripinnate (+pinnatipartite); rachis slightly zigzag. Primary pinnae short-stalked, alternate or the lower often
in subopposite pairs. Upper pinnae, pinnules, etc.
gradually reduced and confluent. Texture usually firm.
Leaf axes adaxially grooved, the groove continuous
between axes of different order. Ultimate segments
about ovate-Ian ceo late, often unequal-sided. Veins
free, often obscure, ending behind the margin; costae
adaxially grooved or not. Surface of lamina glabrous
or pubescent, glandular or not. Sori on a marginal
commissure occupying a few to all vein ends of a segment; outer indusium, i. e., the modified and reflexed
leaf margin, firm with thinner edge, oblong to linear;
inner indusium evident, chartaceous, often lacerate.
No appendages among the sporangia. Spores monolete, ellipsoidal, coarsely and irregularly rugose. There
are counts of n - 26 and n -104, the latter evidently
octoploid; but few samples have been counted.
About a dozen species, only two in tropical America, the others in eastern Asia and the western Pacific,
from Taiwan and Sumatra to New Caledonia and
New Zealand. The New Zealand species P. scaberula
(A. Rich.) Kuhn may become a troublesome weed in
disturbed habitats, like Pteridium. In many ways, Paesia is like Pteridium, including the chromosome number, but the spores differ widely.
8. Hypolepis Bernhardi
87
Fig. 36. Dennstaedtiaceae. Hypo/epis sparsisora, intennittent
development of lamina; La Reunion, Mascarenes. Phot.
K. V.Kramer
Fig. 34C-H, 36
Hypo/epis Bernhardi, Schrad. J. Bot. 1(2): 34 (1806);
R. M. Tryon (1964); Brownsey and Chinnock (1984); Biswas (1985); Brownsey (1987).
Medium-sized to large, terrestrial or epilithic ferns, the
stem long-creeping, often much branched and forming
large clones, solenostelic, bearing pluricellular trichomes. Petioles remote, well-developed, with a single
vascular bundle that in cross-section is U-shaped,
sometimes corrugated; adaxially grooved, the central
groove sometimes flanked by shallower ones, often
pubescent, not rarely also verruculose or even spiny.
Lamina often large, then intermittently developing
rrom one pinna pair to the next and often straggling,
at least bipinnate + pinnatifid, up to 4-pinnate + pinnatifid, anadromous or more often catadromous, depending on the species, usually not thick in texture;
surface usually pubescent with pluricellular, sometimes glandular trichomes; rachis like the petiole, often pubescent, the grooves of secondary rachises not
joining that of the primary, or strongly decurrent and
shallow where joining it far below the point of insertion. Larger pinnae/pinnules often subopposite and
spreading at an open angle. Upper pinnae, pinnules,
etc. gradually reduced and confluent. Ultimate divisions mostly small, often crenate to incised, adnate,
pubescent or not. Costae adaxially grooved to the base
or nearly so, the edges of the groove sometimes winglike; costules not grooved. Veins pinnate in the segments, free, ending behind the margin. Sori terminal
(less often dorsal) on the anterior branch of a vein,
uninerval, roundish, the margin opposite them usually
reflexed as an oblong, ± modified lobe, the sorus thus
located at a sinus (Fig. 34E); in other cases the sori
are unprotected and then not placed near a sinus
(Fig. 340); transitions occur. Appendages among the
sporangia none. Spores monolete, ellipsoidal, reticulate or usually coarsely echinate, often with connecting
strands that form a raised reticulum.
As in Dennstaedtia, there is a confusing array of
chromosome numbers, with n -29, 39, 51-53, c.92,
Dennstaedtiaceae . Dennstaedtioideae
88
B
Fig. 37 A-E. Dennstaedtiaceae. A,B Bloliella currorii. A Fertile pinna (x 0.5). B Detail, with venation and sori (x 1.5).
C Bloliella gracilis. detail of pinnule with sori and venation
(x 3). D, E Hisliopleris incisa. D Part of rachis .nd part of
pinna (x 0.5). E Fertile segment with venation and sori
(x 1.5 (A-C from Tardieu-Blot 1964; D,E from Schelpe
1970)
98, etc.; 2n=c.92, 104, c.150, 208; see Brownsey
(1983) and Brownsey and Chinnock (1984, 1987). Occasional intermediate specimens are probably hybrids;
substantiated reports of hybridization are notably by
Brownsey and Chinnock (1984).
The number of species is estimated at about 40, but
the genus is not well known. It is distributed in tropical and south-temperate regions of the world, with local extensions into north-temperate areas (E. Asia,
Florida); the species are often of local occurrence but
are not rarely abundant in open places and secondary
vegetation, where they may become somewhat weedy,
e. g., H. sparsisora (Schrader) Kuhn in Africa.
9. Biotie/la R. M. Tryon
Fig.37A-C
Blotielfa R. M. Tryon, Contrib. Gray Herb. 191: 96 (1962).
£ondli/is auett., p. p. maio
Pleris .uett., p. p. min.
Medium-sized to very large terrestrial ferns with ascending to erect, up to a few m tall, usually massive
stem with dictyostele, clothed with pluricellular trichomes. Leaves close; petioles well-developed, pale to
purplish, adaxially deeply grooved, in large plants the
groove flanked by two additional ones, bearing pluricellular trichomes, especially in the groove; vascular
bundle one, much corrugated in larger leaves. Lamina
once pinnate or more often bipinnate or bipinnate +
pinnatifid, catadromous or anadromous, truncate at
base. Rachis ridges not interrupted at the pinna insertions; axes of primary pinnae adaxially not grooved;
at least the axes and larger veins pubescent with articulate trichomes, very rarely glabrous. Divisions of
lower order often sessile or short-stalked and subopposite; upper pinnae, pinnules, etc. gradually reduced
and confluent, or sometimes abruptly so, but a conform terminal division never present. Ultimate free divisions roughly lanceolate, entire to pinnatilobate with
rounded lobes; margin entire or sinuate-pinnatifid, not
serrate. Lobes distinctly costuIate; venation areolate,
without free included veinlets; costal and costular areoles elongate; rarely only costal areoles present; venation isodromous or usually distinctly catadromous.
Sori short, confined to the sinuses, or longer and occupying most of the margin and only shortly interrupted,
the outer indusium formed by the reflexed, modified
leaf margin; inner indusium none. Receptacular trichomes abundant, several-celled, often taxonomically
diagnostic. Spores monolete, ellipsoidal, coarsely echinate, rarely with connecting strands between the appendages.
The few chromosome counts extant yielded numbers based on n = 38. About a dozen species in Africa
and adjacent islands, greatly in need of critical study;
a single species, B. lindeniana (Hooker) R. M. Tryon,
in the Neotropics. One of the few fern genera strongly
centred in Africa. Usually growing on forest floor, in
moist to rather dry places, from low to middle elevation.
Bloliella • Histiopteris . Lonchitis
10. Histiopteris(Agardh) J. Smith
Fig. 37D-E
Histiopteris (Agardh) J. Smith, Hist. Fil.: 294 (1875). Basion.
Pteris sect. Histiopteris Agardh, Rec. Gen. Pteridogr.: 76
(1839).
Lepidocaulon Copeland (1942).
Medium-sized to rather large ferns; stem long-creeping, solenostelic, terrestrial, occasionally scandent or
epiphytic, bearing bristles (pluriseriate at base) but not
true scales. Leaves remote; petiole well-developed,
dark-sclerotic, adaxially faintly flattened-sulcate, with
a single vascular bundle. Lamina bipinnate or more
dissected, catadromous (not always distinctly so), truncate at base, firmly herbaceous, often glaucous beneath; primary and often also larger secondary divisions (sub)opposite, developing stepwise from one
pair to the next. Leaf axes reddish-brown, mostly
smooth; segments often spreading at a very open
angle. Pinnae sessile to long-stalked; basal secondary
divisions often reduced, somewhat stipule-Iike, touching the rachis; upper segments reduced and confluent.
Ultimate divisions lanceolate to tongue-shaped, large,
entire to pinnatifid, the greater part adnate, often also
decurrent; costae adaxially not grooved. Sterile margin
entire. Veins reticulate without free included veinlets,
or forming only costal areoles, less often quite free;
venation pattern isodromous or weakly catadromous,
rarely anadromous. Sori occupying all vein ends of a
fertile segment, the commissural receptacle continuous; leaf margin reflexed, strongly modified, forming
the outer indusium; inner indusium none. Soral trichomes numerous, filiform, several-celled. Spores
monolete, ellipsoidal, coarsely tuberculate, the tubercles often partly fused.
The few chromosome counts amount to n = 48 and
n =96; the base number is not known with certainty.
One pantropic to south-temperate species, H. incisa
(Thunb.) J. Smith; in addition about half a dozen more
local Malesian species that are taxonomically poorly
known. Species of Histiopteris grow in thickets, by the
forest edge, in open, disturbed ground; H. incisa may
locally be a troublesome weed. Holttum (1966)
showed that LepidocauIon is congeneric.
11. LOIIdIitis L.
Ftg.38D-E
Lonchitis L. Spec. Plant. 2: 1078 (1753).
Anisosorus Trev. ex Maxon (1926), nom. supertl.
Pteris auctt., p. p. min.
Rather large terrestrial ferns with rather stout and
fleshy, solenostelic stem bearing pale,
pi uri cellular trichomes. Petioles rather close, well-developed, pale to dark brown, with two vascular bundles, adaxially grooved, in large plants additional lateral grooves present. Lamina often large, succulent
when fresh, thin when dry, pinnate + pinnatifid to tri-
~hort-creeping,
89
pinnate + pinnatifid, dissection pattern fluctuating between anadromy and catadromy; base truncate. Pinnae sub opposite, subsessile to stalked; upper pimiae
and segments gradually reduced and confluent. Segments mostly adnate, often decurrent, broad, margin
entire. Leaf axes bearing persistent pluricellular trichomes, adaxially sulcate, the ridges bordering the
groove not interrupted at pinna insertions; pinna rachises with a median ridge flanked by two laterally
winged grooves. Veins free or casually and irregularly
joining, without free included veinlets; venation pattern catadromous. At least the major veins pubescent.
Sori on a submarginal commissure joining several
veins, confined to the sinuses or often more extensive,
often concave; outer indusium formed by the reflexed,
much modified margin; inner indusium none. Receptacular trichomes numerous, filiform, several-celled.
Spores trilete, globose, the aperture prominently projecting; surface granulate.
The few chromosome counts available yielded
n = 50 and 100, respectively.
Two species, L. hirsuta L., widespread in tropical
America, and L. occidentalis Baker in tropical Africa
and Madagascar; in forests, often in locally moist
places, from low to middle elevation. For the nomenclature see R. M. Tryon (1962) and Lellinger (1977).
The genus was formerly construed to include what
is here called Blotiella; in view of their widely diverging spore morphology and chromosome numbers their
closeness is open to doubt.
Subfamily Liodsaeoideae
Stem short- to long-creeping, terrestrial or epiphytic,
protostelic with internal phloem strand, rarely soleoostelic, bearing basally attached, non-clathrate, entire
scales which may be so narrow as to simulate trichomes. Petioles aggregate to remote, with a single
vascular bundle, adaxially grooved, the groove continuous with those on axes of higher order. Mature leaves
nearly always macroscopically glabrous; lamina very
variable, simply pinnate to decompound, rarely simple; upper pinnae and segments reduced and confluent or not, in the latter case with a free terminal leaflet; architecture anadromous. Veins free or anastomosing without free included veinlets. Sori terminal, most
often on a submarginal commissure uniting two to
many vein ends, or occasionally uninerval. Indusium
always present, attached at the base or also at the
sides, opening towards the margin; receptacular trichomes mostly present. Spores trilete, less often monolete.
Species with decompound lamina in Lindsaea, Tapeinidium, and Odontosoria are thought to be primitive, those with simpler leaf architecture derived. The
epiphytic habit of some species of Lindsaea is not accompanied by any evident morphological-anatomical
90
Dennstaedtiaceae . Lindsaeoideae
Fig.38A-E. Dennstaedtiaceae. A,B Odontosoria j/exuosa.
A Part of stem with petiole and portion of lamina with immature apex (x 0.4). B Fertile segment ( x 2.6). C Odontosoria
chinensis, fertile pinnnie (x 4). D,E Lonchitis occidentalis.
D Secondary pinnnie (x 1). E Lobes of pinnnie (x 2) (A,B
from Proctor 1977; C from Kramer 1971; D,E from Christensen 1932)
adaptations, but such species have a distinctly dorsiventral stele in the stem, with the internal phloem
strand displaced towards the dorsal side. Anastomosing veins occur only in Lindsaea but there they have
evidently arisen in several distinct sections and cannot
be used all by themselves for a subdivision of the genus.
A true solenostele occurs in a few species of Tapeinidium and Odontosoria, where it may be regarded as a
primitive character. The "Lindsaea type" protostele
with internal phloem but lacking a medulla and an inner endodermis may be regarded as a derived simplification of the solenostelic condition. For a review of
the anatomy and morphology of the subfamily see
Perez Arbelaez (1928) and Kramer (1957, 1971).
There is a confusing array of chromosome counts
for Lindsaea and Odontosoria (Sphenomerisj, with
probable or possible base numbers of 34, 38, 39, 44,
47,50, and 51, polyploids being fairly frequent. See ai-
91
Odontosoria· Tapeinidium
so Walker (1973). Probable hybrids in Lindsaea were
recorded by Kramer (1957,1963) and Kramer and Tindale (1976).
The subfamily is of pantropical distribution.
12. Odontosoria Fee
Fig.38C
Odontosoria Fee, Mem. Foug. 5: 325 (1852); Maxon (1913).
Stenoloma Fee (1852), p. p. min. and of other authors, p. p.
Sphenomeris Maxon (1913); Kramer (1971).
Twenty-two species, pantropic, extending north to
Korea and to Florida; many species of narrow distribution, e. g., in New Caledonia and around the Caribbean; absent from Australia and New Zealand. Growing on forest floor or more often in thickets, even in
open places, from low to rather high elevation. One
species, O. chinensis (L.) J. Smith, may become somewhat weedy.
13. Tapeinidium (C. Presl) C. Christensen
Medium-sized to large, rarely small, terrestrial ferns
with creeping stem having a Lindsaeoid protostele or
rarely a solenostele; scales of stem sometimes very
narrow, the uniseriate ones resembling trichomes. Petioles well-developed, close or somewhat remote. Lamina sometimes scandent, decompound, at least bipinnate +pinnatifid, often more dissected; axes spiny
and/or flexuous in a few species. Axes of mature
leaves minutely hairy in one species. Ultimate divisions very often narrowly oblong-cuneate or linear,
rarely lanceolate or dimidiate. Texture membranous to
coriaceous. Sori terminal on a single vein or more often on a commissure uniting several vein ends; indusium attached at the base and the greater part of the
sides. Veins free, simple or once to several times
forked in the ultimate divisions, ending behind the
margin. Receptacular trichomes mostly small, fewcelled, large and many-celled in one species. Spores
trilete, globose, with projecting aperture, or monolete,
ellipsoidal; surface often plain, sometimes finely granulate.
Fig. 39
Thpeinidium (c. Pres!) C. Christensen, Ind. AI.: 631 (1906);
Kramer (1968,1971).
Microlepia § Tapeinidium C. Presl, Epimel. Bot.: 96 (1849).
Terrestrial ferns with rather short-creeping stem, this
with a solenostele or in small species with a Lindsaeoid protostele, bearing narrow, long-acuminate scales.
Petioles well-developed, rather close; lamina mediumsized, simply pinnate to tripinnate + pinnatifid; upper
pinnae, pinnules, etc. gradually reduced and confluent
or a conform terminal pinna present. Ultimate divisions oblong, lanceolate, or linear-spathulate, often
acuminate, frequently lobed. Texture herbaceous to
coriaceous. Secondary rachises and costae adaxially
usually only shallowlY to obsoletely grooved. Less dissected species with a percurrent costa; veins simple,
forked, or pinnate in the ultimate divisions, free, not
reaching the margin. Sterile margin often lobulate-serrate. Sori terminal on one vein, or on a commissure
joining two, rarely three vein ends, submarginal; indusium firm, attached at the base and at least part of the
Frg. 39 A,
B. Dennstaedtiaceae. Tapeinidium novoguineense.
A Part of lamina (x 0.4). B Fertile pinnule (x 2) (Kramer
1971)
92
Dennstaedtiaceae . lindsaeoideae
A
D
B
Fig.40A-F. Dennstaedtiaceae. A Lindsaea repens. fertile
pinnule (x 1.5). B Lindsaea flSsa. part of fertile lamina
( x 2.5). C Lindsaea bifida, apex of lamina and middle pinna
( xl). D Lindsaea pendula, apex of lamina ( xl). E Lindsaea
reniformis, lamina (x 0.4). F Lindsaea semilunata, lamina
(x 0.5) (A,B from Kramer 1971; C-F from Kramer 1957)
sides. Receptacular trichomes evident, filiform, multicellular. Spores monolete, ellipsoidal or subglobose,
with prominent, projecting aperture; surface plain or
sparsely granulate.
Seventeen species in Asia and the western Pacific,
from S. India and the Ryukyus to Samoa; wanting in
Australia, New Zealand, and New Caledonia. The
greatest species concentration is in New Guinea In
forests and thickets, from lower to middle, occasionally to higher elevation.
14. Xyropteris Kramer
Xyropleris Kramer, Acta Bot. Neerl. 6: 599 (1957).
In many respects like a large species of Tapeinidium.
Lamina simply pinnate; pinnae large, lanceolate, anteriorly auriculate; terminal division free, trilobed. Sori
of fully fertile pinnae borne on a commissure joining
all vein ends. Spores as in Thpeinidium.
A single species, X. stortii (v. A. v. R.) Kramer, in
Borneo and Sumatra, on rocks by creeks and waterfalls, rare. This might be treated as a subgenus of Tapeinidium.
15. Lindsaea Dryander in J. E. Smith
Fig. 40
Lindsaea Dryander in J. E. Smith, Mem. Acad. Thrin 5: 401
(1793); Kramer (1957, 1963, 1971, 1972a, b); Kramer and
Tindale (1976); Dixit and Ghosh (1983).
Schizoloma Gaudichaud (1824), p. p.
[soloma 1. Smith (1841).
Humbloliella Tardieu-Blot (1956).
Sambirania Tardieu-Blot (1956).
Schizolegnia Alston (1956), nom. supeif/.
Small to medium-sized, terrestrial or epiphytic ferns
with short- to long-creeping stem, its stele a radial to
dorsiventral Lindsaeoid protostele, bearing non-clathrate (or very weakly clathrate), (sub)entire, ovate to
narrowly lanceolate, rarely trichome-like scales. Petioles very short to well-developed, aggregate to remote, glabrous when mature, stramineous to dark.
Lamia subsessile to long-petiolate, simply pinnate to
decompound or in a few species simple; ultimate divisions often dimidiate and entire or lobed-cleft, less often cuneate to linear and cleft or lanceolate and entire;
in one section the pinnae articulate at base. Costa absent, or present in a few species with undivided, equalsided pinnae, adaxially hardly sulcate; veins subpinnately issuing from the vein bordering the posterior
margin in dimidiate species, otherwise simple or once
to several times forked, ending behind the margin, free
or reticulate without free included veinlets. Texture
membranous to coriaceous, surface glabrous or very
rarely the axes minutely hairy. Sori terminal on single
veins, or on a commissure joining all vein ends of a
segment, with all possible transitions; indusium accordingly short to elongate, if short attached at the
base only, opening towards the margin. Receptacular
trichomes small, few-celled, often hardly evident.
Xyropteris • Lindsaea . Orm%ma
Spores usually trilete, tetrahedral, or less often monolete, ellipsoidal, the aperture often prominent; surface
irregularly granulate or with curved rodlets.
About 150 species, pantropic, but very few in continental Africa; extending into temperate areas in
South Brazil, Australia, New Zealand, and Japan. Terrestrial or epiphytic, mostly in seasonally moist or
everwet, often submontane forests; a few species occur
preferably in open habitats; one species, L. ensifolia
Swartz, may be slightly weedy. Occurring from sea level to c. 2000 m.
Two subgenera: Lindsaea;terrestrial, with (sub)radial stele, pantropic; and Odontoloma, epiphytic, with
long-creeping stem and dorsi ventral stele, from the
Mascarenes and Ceylon to Hawaii. Twenty-one sections distinguished by Kramer (1957-1976), some not
well defined. Certain species with divergent appearance have been placed in segregate genera, but these
are never based on characters of any fundamental importance.
16. Ormoloma Maxon
Orm%ma Maxon, Proc. BioI. Soc. Washington 46: 143
(1933); Kramer (1957); Tryon and Tryon (1982).
Sacc%ma auett., p. p.
Medium-sized, terrestrial ferns with long-creeping
stem, stele a Lindsaeoid protostele; scales ovate-Ianceolate. Petioles remote, dark, well-developed. Lamina
simply pinnate, with conform terminal pinna, glabrous, herbaceous; pinnae lanceolate, almost symmetric, costate, with free, forked veins ending behind the
margin. Sori terminal on single veins; indusium semiorbicular to semi-ovate, attached only at the base, submarginal. Soral trichomes none. Spores trilete, tetrahedral-globose with prominent angles and sometimes
prominent aperture; surface bearing curved, often
± coalescent rodlets.
Two species described in Costa Rica, the Lesser Antilles, Hispaniola, and the Guianas; but perhaps
monotypic; O. imrayanum (Hooker) Maxon. Terrestrial in forests, at lower and middle elevation. Close to
Lintisaea, but with a combination of characters nowhere present in that genus, and without an obvious
source there.
Selected Bibliography
Alston, A. H. G. 1956. The subdivision of the Polypodiaceae.
Taxon 5: 23-25.
Biswas, A. 1985. The genus Hypo/epis Bernb. in India. J.
Eeon. Taxon. Bot. 7: 111-124.
Brownsey, P. J. 1983. Polyploidy and aneuploidy in Hypo/epis, and the evolution of the Dennstaedtiales. Amer. Fern
J. 73: 97-108.
Brownsey, P. J. 1987. A review of the fern genus Hypo/epis
93
(Dennstaedtiaceae) in the Malesian and Pacific regions.
Blumea 32: 227-276.
Brownsey, P. J., Chinnock, R J. 1984. A taxonomic revision
of the New Zealand species of Hypo/epis. New Zeal. 1.
Bot. 22: 43-80.
Brownsey, P.J., Chinnock, RJ. 1987. A taxonomic revision
of the Australian species of Hypo/epis. J. Adelaide Bot.
Gard. 10: 1-30.
Conard, H. S. 1908. The structure and life history of the hayscented fern. Washington, D. C.: Carnegie Institution.
Copeland, E. B. 1950. Pteridaceae of New Guinea. Philipp. J.
Sci. 78: 5-41.
Copeland, E. B. 1958. Fern flora of the Philippines I. Manila.
Dixit, R. D., Ghosh, B. 1983. The genus Lindsaea Dryand. ex
Smith in India. Proc. Indian Acad. Sci. (Plant ScL) 92:
233-258.
Gruber, T. M. 1981. The branching pattern of Hypo/epis repens. Amer. Fern J. 71: 41-47.
Hagemann, W. 1976. Sind Fame Konnophyten? Eine Alternati"e zur Telomtheorie. Plant Syst. Evol. 124: 251-277.
Holttum, R E. 1947. A revised classification of I.eptosporangiate ferns. Bot J. Linn. Soc. 53: 123-158.
Holttum, R E. 1966. The geuera Lepidocau/on Copel. and
Histiopteris J. Sm. Kew Bull. 20: 458-460.
Keating, R. C. 1968. Trends of specialization in the stipe anatomy of Dennstaedtia and related genera. Amer. Fern J. 58:
126-140.
Kramer, K. U. 1957. A revision of the genus lindsaea in the
New World with notes on allied genera. Acta Bot. Neerl. 6:
97-290.
Kramer, K. U. 1963. The fern genus Lindsaea in Santa Catarina and Rio Grande do Sui, Brazil. Sellowia 15: 115-121.
Kramer, K. U. 1968. The Lindsaeoid ferns of the Old World
II. A revision of Tapeinidium. Blumea 15: 545-556.
Kramer, K. U. 1971. Lindsaea Group. Flora Males. II. 1(3):
177-254.
Kramer, K. U. 1972a. The Lindsaeoid ferns of the Old World
IX. Africa and its islands. Bull. Jard. Bot. Nat. Belg.42:
305-345.
Kramer, K. U. 1972b: Id. VI. Continental Asia, Japan and
Taiwan. Gard. Bull. Singapore 26: 1-48.
Kramer, K. U., Tindale, M. D. 1976. The Lindsaeoid ferns of
the Old World VII. Australia and New Zealand. Telopea 1:
91-128.
I.ellinger, D. B. 1977. The identity of Lonchitis au rita and the
generic names Anisosorus and Lonchitis. Taxon 26:
578-580.
Maxon, W. R 1913. The genus Odontosoria. Contrib. U. S.
Nat. Herb. 17: 157-168.
Mickel, J. T. 1973. The classification and phylogenetic position of the Dennstaedtiaceae. Bot. J. Linn. Soc. 67 Supp!.l:
135-144.
Nair, G. B. 1979. Peltate scales in Saccoloma. Fern Gaz.
12(1): 53-55.
Page, C. N. 1976. The taxonomy and phytogeography of
bracken - a review. Bot. J. Linn. Soc. 73: 1-34.
Perez Arbelaez, H. 1928. Die natiirliche Gruppe der Davalliaceen (Sm.) Klfs. Bot. Abh. (Jena) 14: 1-96.
Perring, F. H., Gardiner, B. G. (Eds.) 1976. The biology of
bracken. Bot. J. Linn. Soc. 73: 1-302.
Troop, J. E, Mickel, J. T. 1968. Petiolar shoots in the Denn·
staedtioid and related ferns. Amer. Fern J. 58: 64-69.
Tryon, R M. 1941. A revision of the genus Pteridium. Contrib. Gray Herb. 134: 1-31, 37-67.
94
Filicatae . Dicksoniaceae
Tryon, R. M. 1960. A review of the genus Dennslaedlia in
America. Contrib. Gray Herb. 187: 23-52.
Tryon, R. M. 1962. Taxonomic fern notes III. Contrib. Gray
Herb. 191: 91-107.
Tryon, R. M. 1964. The ferns of Peru [Hypo/epis]. Contrib.
Gray Herb. 194: 34-39.
Walker, T. G. 1973. Additional cytotaxonomic notes on the
pteridophytes of Jamaica. Trans. R. Soc. Edinburgh 69(5):
109-135.
Dicksoniaceae
K. U.KRAMER
Dicksoniaceae Bower, Orig. Land PI.: 591 (1908).
Thyrsopteridaceae C. Presl, Gerossb. i. Stip. d. Farm: 38
(1847).
Culcitaceae Pichi Sermolli, Webbia 24: 702 (1970).
Cystodiaceae Croft, Kew Bull. 41: 797 (1986).
Cyatheaceae auctt. div., p. p., e. g., Holttum, Fl. Males. II. I':
65-176 (1963).
Large to very large terrestrial ferns with massive,
creeping to erect and then mostly ±trunk-like stem often bearing adventitious roots and/or persistent petiole bases; stele a solenostele or a dictyostele; indu·
ment of very long, simple multicellular, yellowish to
dark brown trichomes. Leaves close; petiole non·articulate, often long, strong, terete, adaxially flattened or
centrally grooved, in large species with additional ventro-lateral grooves, with one to three, often corrugated
vascular bundles or occasionally with more numerous,
smaller, strap-shaped bundles arranged in an horseshoe-like pattern (Fig. 41 D) (but see under Cystodium). Lateral pneumathodes present, forming a continuous or interrupted line. Hairs of axes often persistent
at junctions or on the adaxial face. Lamina at least bipinnate, usually more dissected, firm, monomorphic to
dimorphic, with few exceptions catadromous, bearing
multicellular trichomes at least adaxially on the axes.
Stomata diacytic, paracytic and/or anomocytic. Ultimate divisions usually of an approximately lanceolate
type, often crenate to pinnatifid; upper divisions gradually reduced and confluent. Veins forked or pinnate,
free, ending near the margin. Fertile leaf divisions not
rarely with weakly to strongly reduced laminal parts
("skeletonized" in extreme cases). Sori single on vein
ends, submarginal, protected by an abaxial ("inner")
indusium and a hardly to strongly modified lobe of
the leaf segment ("outer indusium"), these together
forming a cup-like to box-like structure. Multicellular,
filiform trichomes usually present among the sporangia. Receptacle often transversely enlarged, occasion·
ally columnar and free from the lamina. Sporaniga
maturing in gradate sequence, short- to long-stalked,
the stalk up to 6-seriate; capsule with longitudinal,
vertical, or slightly oblique annulus that bypasses the
stalk, most of it thickened as the bow. Spores trilete.
ANATOMY AND MORPHOWGY. Comprehensive studies,
even of single genera, are few and often not recent.
See Bower (1926), Ogura (1930), Holttum and Sen
(1961), and for individual taxa and special aspects,
Chandra (1970), de Rezende-Pinto (1943), and Croft
(1986).
In the non-arborescent species the stem is creeping
or ascending, rarely erect, and solenostelic; in arborescent species it is erect and usually dictyostelic; transitions exist. The meristeles of the upright-stemmed species (Dicksonia) are highly corrugated and accompa·
nied by layers of very hard, sclerotic tissue on both
sides that follow the corrugations and have a reinforc·
ing function. The pith is large and (except in Thyrsopteris) without medullary bundles or sclerenchyma.
The leaf traces are simple and horseshoe-shaped in
transection at their extreme bases but divide a little
higher into several strands, at least in some species;
sclerotic tissue may be present. So·called cubical cells,
described by Holttum and Sen (1961), accompany the
sclerotic cell masses; they are absent in Cibotium but
connect the other genera with Cyatheaceae, Metaxyaceae, and Lophosoriaceae. Thyrsopteris diverges in its
purely anomocytic stomata.
GAMETOPHYTE. This does not show any remarkable
features, except for its primitive antheridia with several wall cells. See Mukherjee and Sen (1986), also for
resemblances to the gametophyte of Cyatheaceae.
KARYOWGY. The chromosome numbers, as far as
known, show a wide range, enhancing the relict nature
of this ancient family. Counts have been published as
follows: Thyrsopteris n=c. 76-78; Culcita 71=68; Calochlaena 71=55-58; Dicksonia 71=65 (271=130); and
Cibotium 71 = 68.
SUBDIVISION. The subdivision as given by Holttum and
Sen (1961) and Holttum (1963) seems natural, except
for placing Dicksonia and Cystodium in a tribe with
the Cyatheoideae, the latter being put here in a different family. We may thus distinguish the subfamilies
Dicksonioideae, with Dicksonia, Thyrsopteridoideae,
with Thyrsopteris, Culcita, and Calochlaena, and Cibotioideae, with Cibotium. Cystodium is usually placed
near Dicksonia, but Croft (1986) showed that it is quite
distinct in its vascular anatomy and in its spore sculpture. His solution to put it in a family by itself is not
accepted here, but placement in another subfamily is
probably justified. The other subfamilies are also characterized by anatomical characters, and in addition
features of the rachis structure and of the "outer indusium". Thyrsopteris is distinct ~nough to be placed in a
tribe by itself, but if it is put in a distinct family (Pichi
Thyrsopleris
Sermolli) or even in a distinct order (Kunkel 1965), its
divergent characters are overemphasized. The subdivision proposed by de Rezende-Pinto (1943) is based on
superficial characters.
ECOLOGY AND DISTRIBUTION. Most species occur in
the undeIgrowth of not too dense forests, often in
thickets or more open vegetation, and on mountain
sides; some species ascend to over 3000 m. The scattered distribution in tropical (often montane) and
north- and south-temperate areas of the world underscores the relic nature of this family, which has a long
fossil record. See, e. g., the map of Cibotium in van
Steenis and van Balgooy (1966).
AFFINITY. Like other primitive leptosporangiate families, the Dicksoniaceae occupy a somewhat isolated
position. They were formerly associated with, or included in, the Cyatheaceae, because of similar habit
and sporangial structure. Bower (1908, 1926) removed
them from that family, mainly because of their marginal sori, and placed them near the Dennstaedtia alliance; Copeland (1947) included these two in his Pteridaceae, now regarded as artificial. Holttum and Sen
(1961) and Holttum (1963) reverted to the first-named
classification, sinking Dicksoniaceae again in Cyath-
95
Fig. 41 A-E. Dicksoniaceae. A-C Cibolium schiedei. A Pinna rachis with pinnules (x 1). B Fertile segment from abaxial
side (x 4.5). C Sori (x 10). D Cibotium barometz. transection
of petiole and rachis with vascular bundles (schematic).
E Cibolium menziesii, sporangia from three different angles
(x 50) (A-C from Hooker 1846; D from Ogura 1972;
E from Bower 1926).
eaceae. Although the character of the marginal vs. dorsal sorus is at present not so heavily weighted, few authors followed Holttum's merger. As pointed out by
Tryon and Tryon (1982), Metaxyaceae and Lophosoriaceae, though having dorsal sori, agree with Dicksoniaceae in having only trichomes and lacking scales,
rather than with Cyatheaceae in which they were formerly included.
KEy TO THE GENERA.
1. Lamina anadromously dissected; all axes adaxially
grooved, the grooves open to one another
2
- Lamina catadromously dissected; or, if the larger divisions
of lower order are seemingly anadromous, the axes are
adaxially raised
3
2. Lamina strongly dimorphic, the fertile segments skeletonized; receptacle columnar, partly free; no trichomes among
1. ThyrsopU!rls
the sporangia
96
Filicatae . Dicksoniaceae
- Lamina not dimorphic, fertile segments not contracted; receptacle not columnar and free; trichomes present among
2. CaJocbJDena
the sporangia
3. Rachises of highest order adaxially grooved
4
5
- Rachises of highest order adaxially raised
4. Lamina bipinnate, with oblong pinnae
5. Cystodium
- Lamina more dissected; pinnae and pinnules deltoid, ta3. Cul.ita
pering from base to apex
5. Outer indusium sharply offset from the laminal part bearing it; stem creeping-ascending or erect, mostly solenostelie; fertile segments hardly modified; lamina not narrowed
at base
4. Qbotium
- Outer indusium not sharply offset from the laminal part
bearing it; stem erect, dictyostelic; fertile segments weakly
to strongly modified, with reduced laminal parts; lamina
6. Dicksonw
narrowed at base
FJg.42A-G. Dicksoniaceae. A,B Thyrsopteris eiegans.
A Part of fertile pinna ( x 5). B Vertical section through sorus
( x 110). C,O Calochlaena dubia. C Fertile pinnule (x 1).
o Fertile segment from abaxial side ( x 5). E Culcita macrocarpa, vertical section through sorus (x 92). F, G Cystodium
sorbifolium. F Part offertile pinna (x 3). G Edge of lamina
with sori from abaxial side (x 10) (A from Diels 1899;
B,E from Holttum and Sen 1961; C,O,F,G from Hooker
1846)
1. Thyrsopteris Kunze
Fig. 42 A, B
Thyrsopleris Kunze, Unnaea 9: 507 (1834).
Terrestrial ferns with ascending to erect, solenostelic
stem bearing runners and stiff, long, multicellular trichomes. Leaves long-petiolate; petiole adaxially
grooved, with one, in transection horseshoe-shaped,
corrugated vascular bundle, petiole base long-hairy.
Lamina 3- to 5-pinnate, firm, glabrous when mature,
anadromous, the axes adaxially grooved up to the costae of the segments, all grooves open to one another;
ultimate divisions narrowly cuneate. Veins free, not
reaching the margin. Stomata anomocytic. Fertile portions mostly borne in the basal part of the lamina,
much dissected and with almost absent leaf tissue
("skeletonized"), up to 7-pinnate; sori terminal on
stalk-like veins, the outer and inner indusia fused to
form an asymmetric, cup-shaped structure, a short,
free, columnar receptacle in its middle; no trichomes
among the sporangia. Sporangia maturing basipetally,
with a very short stalk of several cells; annulus complete, bypassing the stalk, the indurated part not
reaching the stalk nor the several-celled stomium,
Spores globose-tetrahedral with prominent angles;
exospore thick, two-layered; surface formed on the
lower layer of the perispore, mucro-verrucate by irregular granulate perispore deposit,
CaJochlaena • Culcita
97
A single recent species, Th. elegans Kunze, confined
to the Juan Fernandez Islands in the SE Pacific. Fossil
representatives of the genus ('!) are widespread, and its
present distribution is obviously a relic. Th. elegans
grows in moist woods and thickets, generally between
400 and 700 m (Looser 1966).
2. Calochiaena (Maxon) White & Turner
Fig. 42 C, D
Calochlaena (Maxon) R. A. White and M. D. Turner, Amer.
Fern 1. 78: 91 (1988).
Cu/cita subgenus Calochlaena Maxon, 1. Washington Acad.
Sci. 12: 454 (1922); Holttum (1963).
Terrestrial ferns with stout, solenostelic, creeping-ascending stem bearing long, dark, multicellular trichomes. Pneumathodes on petiole narrow, inconspicuous, forming a somewhat interrupted line. Vascular
bundles at petiole base converging, with incurved
edges; higher in the petiole a single U-shaped bundle.
All leaf axes adaxially grooved, the grooves continuous. Lamina chartaceous·subcoriaceous, bipinnate +
bipinnatifid to tripinnate +pinnatifid, anadromous;
axes and veins initially or permanently pale-hairy. Ultimate divisions lanceolate, subacute-acute, dentate to
pinnatifid. Veins pinnate, the branches mostly simple.
Sori terminal on a vein, up to c. 1 mm wide; outer indusium little differentiated, shortly connate with the
inner indusium at base, the two forming a rellexed, bilobed cup; receptacle not transversely elongate. Stomata paracytic and anomocytic. Pluricellular trichomes present among the sporangia. Spores globose;
outer layer of exospore forming contours; surface
bearing irregular, basall y conll uent tubercles with papillate surfaces, a thin granulate layer formed by the
perispore.
Five species, from the Philippines and Java to Australia and Samoa; incorrectly reported from Taiwan
through confusion with Dennstaedtia. Long included
in Culcita, but the differences are obvious and consistent; chromosome numbers and characters of the pro·
thallia also confirm the separation (MukheIjee and
Sen 1986).
3. Culcita C. Presl
Figs. 42 E, 43
Culcita C. Presl, Tent. Pteridogr.: 135 (1836); Maxon (1922)
p.p.; de Rezende-Pinto (1943).
Balanlium auctt., excL type.
Rather large terrestrial ferns with stout, creeping or ascending stem with solenostele, bearing persistent petiole bases and articulate trichomes. Leaves rather
close; petiole with one in transection U-shaped vascular bundle, adaxially grooved like the other leaf axes,
the grooves continuous; long, articulate, yellowishbrownish hairs persistent at the base, sometimes also
elsewhere. Pneumathodes in two lateral series of dis-
Fig.43. Dicksoniaceae. Culcita macrocatpa, sterile and fertile
pinnules from abaxial side; Pico, Azores. Phot. K. U. Kramer
crete knobs. Lamina firm, 4-5-pinnate + pinnatifid, ca·
tadromous, sparingly hairy to glabrous when mature,
with ovate-lanceolate, acute ultimate segments. Veins
free, often forked. Fertile portions conform or with
somewhat reduced laminal parts. Sori large, to 3 mm
wide, terminal on a vein, with trichomes among the
sporangia. Receptacle transversely elongate, attached
to the inner indusium; outer indusium scarcely differentiated from the lamina, basally shortly connate with
the inner indusium and covering its edge when young.
Spores tetrahedral·globose, ± depressed between the
angles or these prolonged; surface shallowly tuberculate to rugose or bearing rodlets, especially on the distal face.
When Calochlaena is excluded, this is a genus of
two closely related species: C. coniifolia (Hooker)
Maxon in tropical America from southern Mexico to
Bolivia and Venezuela, in the Greater Antilles, and in
SE Brazil; and C. macrocarpa C. Presl in the Azores,
Madeira, Tenerife, and the western Iberian Peninsula.
The species grow in open forests, thickets, and more
open vegetation.
98
4. Obotium Kaulfuss
Filicatae . Dicksoniaceae
Fig.41
Cibotium Kaulfuss, Berliner Jahrb. Pharm. 21: 53 (1820);
Maxon (1912); Ogura (1930); Becker (1984).
Very large terrestrial ferns; stem massive, creeping to
erect, occasionally to several m tall, solenostelic or
sometimes dictyostelic, bearing roots and persistent
petiole bases, hairy with many long, yellowish-brown,
pluricellular trichomes. Leaves close, forming a crown
at the stem apex; petiole with three corrugated vascular bundles, one abaxial, two adaxial, with much incurved ends; pneumatophores forming a line on each
side; adaxial face of petiole flattened. Lamina bipinnate + pinnatifid to tripinnate + pinnatifid, firm, not
rarely glaucous beneath, glabrous when mature or
± persistently pale-hairy on axes and veins. Venation
catadromous; dissection pattern essentially also catadromous, but on larger divisions often a few basal
basiscopic pinnules lacking and the dissection then
pseudo-anadromous (Kramer 1987). Leaf axes adaxially raised, hairy in the middle, in dried material the
larger axes often sulcate in the middle of the ridge
which is flanked by two grooves. Stomata paracytic.
Veins free, simple or forked to pinnate; costules adaxially convex, costae convex or grooved in the middle.
Sterile segments entire or crenate; fertile segments not
differentiated. Sori single on a vein end, on the posterior branch if the vein is forked, confined to the basal
or middle portion of a segment or extending to its
apex, isodiametric, or the lower sori broader than long,
much reflexed; outer indusium strongly differentiated
from the lamina, firm like the inner, approximately
tongue-shaped indusium, the two shortly connate at
the base; trichomes mostly present among the sporangia. Spores tetrahedral-globose with prominent angles;
outer layer of exospore sometimes forming a prominent flange and ridges; surface coarsely ridged, especially distally, with prominent equatorial flange; perispore thick, bistratose.
Eleven species, three in Asia, from north-eastern India, southern China, and the Ryukyus to Borneo; two
in Mexico and northern Central America; the remainder, six species (Becker 1984), in the Hawaiian Islands.
5. Cystodium J. Smith
Fig.42F,G
Cystodium J. Smith in Hooker and Bauer, Gen. Fil.: pI. 96
(1841); Holttum (1963); Croft (1986); nom. cons. propos.,
non Cystodium Fee (1837).
Cystodiopteris Rauschert, Taxon 31: 555 (1982).
Stem creeping to erect, dictyostelic, bearing long, multicellular trichomes. Petiole densely long-tawny hairy
at base, with two vascular bundles with short branches
near their ends, these lateral and outward concave in
the basal, dorsoventral in the upper part of the petiole,
a zone with one X-shaped bundle in between. Lamina
bipinnate, catadromous; rachises adaxially grooved up
to those of the highest order, the grooves of axes of
different order not continuous among each other. Pinnules tongue-shaped, sessile, adnate, serrate, with simple or forked veins and adaxially grooved costa. l\ichomes of lamina sparse, long-articulate. Fertile pinnules slightly contracted, shallowly lobed. Sori terminal on single veins in small lobes, rather as in
Dicksonia, with abundant multicellular trichomes
among the sporangia. Spores globose; exospore thick,
of two layers, the outer rugulose; inner layer of perispore forming low, sinuous ridges, surface with granular deposit on these.
A single species, C. sorbifolium (J. E. Smith) J. Smith
in Hooker and Bauer; in Malesia from North Borneo
to the Admiralty Islands and Louisiades; a distinct
subspecies in the Solomon Islands (Croft 1986). In
lowland forest, to 400 m.
6. Dicksonia L'Heritier
Dicksonia L'Heritier, Sert. Angl.: 30 (1789); Holttum (1963).
Balantium Kaulfuss (1824), as to type.
Large terrestrial ferns; stem ascending or erect, rarely
prostrate, trunk-like, to several m tall, dictyostelic; vascular bundles strongly corrugated and accompanied
by much sclerotic tissue; surface bearing many roots,
the apex bearing also yellowish to dark brown, multicellular trichomes, Petiole not very long, usually with
three corrugated vascular bundles with incurved ends,
adaxially not concave, base clothed in many long trichomes; pneumathodes forming a continuous line on
the petiole. Lamina mostly large, firm, bipinnate + pinnatifid to quadripinnate, somewhat narrowed at base,
not glaucous beneath but often persistently hairy; leaf
axes adaxially raised and strigose-hairy; dissection
pattern and venation catadromous. Leaf divisions subsessile, ultimate ones serrate to pinnatifid. Veins free,
simple or the basal forked. Stomata paracytic or diacytic. Fertile segments with faintly to strongly reduced leaf tissue but hardly "skeletonized". Sori terminal on a single vein, usually approximately isodiametric, much deflexed; outer indusium little differentiated
from the lamina; inner indusium about similar or thinner, basally shortly connate with the outer indusium;
receptacle elevated, somewhat transversely extended;
filiform multicellular trichomes with differentiated
terminal cell usually present. Sporangia distinctly
stalked; annulus oblique, indurated around the base,
with lateral stomium. Spores globose, sometimes with
prolonged angles; exospore two-layered, usually pitted, perispore filling the pits and forming an irregular
surface, surface plain or coarsely reticulate, a particulate deposit ± fused into rodlets.
Between 20 and 25 taxonomically not well known
species in tropical and south-temperate areas; in
Filicatae . Dipteridaceae
America from Mexico to Venezuela, Bolivia, and Vruguay, and in the Juan Fernandez Islands; in Australasia from the Philippines and Sumatra to Tasmania,
New Zealand, and Samoa; and in St. Helena. The
greatest concentration of species in New Guinea.
Selected Bibliography
Becker, R. 1984. The identification of Hawaiian tree ferns of
the genus Cibotium. Amer. Fern J. 74: 97-100.
Chandra, S. 1970. Vascular organization of the rhizome of
Cibotium barometz. Amer. Fern J. 60: 68-72.
Croft, J. R 1986. The stipe and rachis vasculature of the
Dicksonioid fern, Cystodium sorbifolium (Cystodiaceae).
Kew Bull. 41: 789-803.
HoltlUm, R E. 1963. Cyatheaceae. FI. Males. II. I, 2: 65-176.
Holttum, R. E., Sen, U. 1961. Morphology and classification
of the tree ferns. Phytomorphology 11(4): 406-420.
Kramer, K. U. 1987. A brief survey of the dromy in fern
leaves, with an expanded terminology. Bot. Helv. 97:
219-228.
Kunkel, O. 1965. Catalogue of the pteridophytes of the Juan
Fernandez Islands (Chile). Nova Hedwigia 9: 245-284.
Looser, O. 1966. Los pterid6fitos 0 helechos de Chile. III.
Cyath~aceae. Rev. Univ. Cat. Chile 50/51: 75-93.
Maxon, W.R. 1912. The American species of Cibotium. Contrib. U. S. Nat. Herb. 16(2): 54-58.
Maxon, W. R. 1922. The genus Cukita. J. Washington Acad.
Sci. 12: 454-460.
MukheIjee, A. K., Sen, T. 1986. Oametophytes of some tree
ferns and their impact on phylogenetic relationships. Indian Fern J. 3: 70-81.
Ogura, Y. 1930. On the structure of Hawaiian tree ferns with
notes on the affinities of the genus Cibotium. Bot. Mag.
(Tokyo) 44: 467-478.
Rezende Pinto, M. e. de 1943. Culcita macrocarpa Pr. Contribuiyiio para 0 seu estudo monografico.· Bol. Soc.
Brot. II. 17: 94-146.
Sen, U. 1964. Importance of anatomy on the phylogeny of
tree ferns and their allies. Bull. Bot. Soc. Bengal 18 : 26-34.
van Steenis, e.O.G.J., van Balgooy, M. M.J. 1966: Pacific
Plant Areas 2. Blumea Suppl. Vol. V: 1-312.
White, R A., Turner, M. 0.1988. Calochlaena, a new genus of
Dicksonioid ferns. Amer. Fern J. 78: 86-95.
99
Dipteridaceae
K. V.KRAMER
Dipteridaceae [Diels (1899) in Engler and Prant!, Nat. Pflanzenfam. I, 4: 202, as tribe of "Aspidinae"j Seward and
Dale, Philos. Trans. R Soc. London Ser.B 194: 487 (1901).
Medium-sized to rather large, terrestrial Or often cpilithic ferns with rather long-creeping stem clothed with
dark, very long-tapering, non-clathrate bristles resembling scales in dried material; stele a radial solenostele. Petioles remote, weakly two-ranked, non-articulate, with one V-shaped vascular bundle or at the base
with several strands arranged in a V (depending on
the species), if one, the "V" bifurcating upward; petiole externally subterete or adaxially concave. Lamina
at least once cleft dividing it into two (sub)equal parts
(in most species again one to several clefts), the primary divisions anadromously divided, acroscopically
more strongly developed, in living plants at the base
twisted through 90 0 , in pressed specimens through
1800 , their abaxial surface then facing upward; lamina
hypostomatic with paracytic stomata, bristly with pale
trichomes when young, glabrescent to glabrous when
mature. Larger veins slightly unequally dichotomous,
the larger branch facing outward; smaller veins forming a dense network with many free included veinlets
(Fig. 44), abaxially prominent. Sori small, round, exindusiate, compital, scattered over the surface, or in narrow-lobed species irregularly few-seriate. Sporangia
intermingled with hair-like soral trichomes with swollen top cell, small, several per sorus, maturing simultaneously or not, with short, 4-seriate stalk; annulus longitudinal, slightly oblique, almost entirely indurated,
the stomium ill-defined from the bow. Spores monolete, ellipsoidal, achlorophyllous, 64 or fewer per sporangium; perispore thin; surface smooth or sometimes
rugulose.
ANATOMY AND MORPHOLOGY. See the family description.
GAMETOPHYTE. Desribed by Stokey (1945) and Nayar
and Kaur (1971) as naked, cordate-thalloid with thick
midrib, slowly growing, eventually ± elongating, bearing gametangia on the ventral and often also on the
dorsal side; antheridium of a primitive, massive type
with rew sperms; archegonium also primitive, particularly in its long, straight neck.
AFFINITY. Dipteris bears a certain resemblance to the
equally isolated genus Cheiropleuria but has otherwise
no close relatives. Points of similarity with Matonia
were discussed by Bower (1926), but the differences
are considerable. Association with Polypodiaceae rests
on superficial resemblances, the antheridium, sporan-
100
Filicatae • Dipteridaceae
Fig. 44A-E. Dipteridaceae. DipteTis conjugata. A Half of
lamina (x 0.3). B Base of lamina from abaxial (left) and
adaxial (right) side, showing basal twisting (x 0.8). C Part of
lamina with venation and sori (x 1.4). D Detail of segment
with venation and sori (x 5). E Sporangium (x 130) (A-D
from Holttum 1955; E from Haider 1954)
gium, stelar anatomy, trichomes, etc. being entirely different. Stokey (1945) stressed affinity with Gleicheniaceae, mainly on the basis of gametophytic characters;
Jarrett (1980) arrived at a similar conclusion. See also
under Cheiropleuriaceae. Holttumiella Copel. was described as a relative of Dipteris and placed in tbe same
family by Nayar and Kaur (1971), although it had already been shown to be a synonym of Taenitis.
The family is of considerable antiquity; various
Mesozoic fossils, from the uppermost Triassic onward,
are referable to it, some probably to the genus Dipteris
itself (see, among otbers, Postbumus 1928a, b). The
group was formerly of almost worldwide distribution,
and Dipteris in its present area is a relict.
KARYOLOGY AND HYBRIDIZATION. Chromosome numbers of n = c. 33, 2n = 66 and 132, have been reported,
emphasizing again the distinctness from tbe great majority of the Polypodiaceae. Apogamy in otherwise
sexually reproducing prothallia was observed by Stokey (1945) and Nayar and Kaur (1971). The spore germination is according to the "Gleichenia type" (Nayar
and Kaur 1968); filament formation slight or absent.
Fig. 45. Dipteridaceae. Dipteris novoguineensis; New Guinea.
Pho!. E. Zogg
Filicatae . Dryopteridaceae
A probable hybrid was described by Nakaike
(1983).
A single, palaeotropical-subtropical genus:
Dipf£ris Reinwardt
Figs. 44, 45
Reinwardt, Syll. PI. 2: 3 (1824); Posthumus
(1928 a, b).
Phyma/odes C. Presl (1836), p. p. min., sed quoad typum.
101
Dryopteridaceae
K. UKRAMER
with contributions by R. E. HOLTTUM, R. C. MORAN,
and A. R. SMITH
Dip/ens
Characters of the Family.
About eight species, from NE India, southern China,
and the southern Ryukyus to NE Queensland and Fiji.
Most species grow on rocks, on soil, in exposed
places, clearings, thickets, etc.; one species, D. lobbiana (Hooker) Moore, is a facultative rheophyte (van
Steenis 1981, with Fig.).
Selected Bibliography
Jarrett, F. M. 1980. Studies in the classification of the leptosporangiate ferns: I. The affinities of the Polypodiaceae
sensu stricto and the Grammitidaceae. Kew Bull. 34:
825-833.
Nakaike, T. 1983. A contribution to the fern flora of New
Guinea (V.). Bull. Nat. Sci. Mus. (Tokyo) B9: 98.
Nayar, B. K., Kaur, S. 1968. Spore germination in homosporous ferns. J. Palynol. 4: 1-14.
Posthumus, O. 1928a. Einige Eigentiimlichkeiten der Blattform bei Dip/elis und bei anderen noch lebenden oder fossilen Pflanzen. Rec. Trav. Bot. Neerl. 25: 241-292.
Posthumus, O. 1928b. Dip/elis novo-guineensis, ein "Iebendes
Fossil". Rec. Trav. Bot. Neerl. 25: 244-249.
van Steenis,C.G.G.J.1981. Rheophytes of the world. Rockville, Md.; Alphen a. d. Rijn. Sijthoff and Noordhoff.
Slokey, A. G. 1945. The gametophyte of Dip/elis coryuga/a.
Bot. Gaz. 106: 402-411.
Dryopteridaceae Ching, Acta Phytotax. Sin. 10: 1 (1%5).
Aspidiaceae Frank in Leunis, Syn. Pflanzenfam. ed. 2, 3: 1469
(1877), nom. iIIeg. (basion. iIIeg.).
Peranemataceae [as Perenemaceaej (C. Presl) Ching, Sunyatsenia 5: 246 (1940), based on: Filicaceae trib. Peranemaceae C. Presl, Tent. Pteridogr.64 (1836).
Woodsiaceae (Diels) Herter, Rev. Sudam. Bot. 9: 14 (1949),
based on: Polypodiaceae trib. Woodsieae Diels in Engler &
Prantl, Nat. Pflanzenfam. I, 4: 159 (1899).
Athyriaceae Alston, Thxon 5: 25 (1956).
Onocleaceae Pichi Sermolli, Webbia 24: 708 (1970).
Hypodematiaceae Ching, Acta Phytolax. Sin. 13: 96 (1975).
Tectanaceae Lellinger Arner. Fern J.77: 102 (1988).
Terrestrial or epilithic, very rarely epiphytic ferns of
very different size. Stem erect or ascending, sometimes
creeping, dictyostelic (mostly radially), very often
bearing old leaf bases which may be incrassate to form
trophopods, scaly; scales basally attached, more often
non-clathrate than clathrate, superficially glabrous, entire or, if dentate, the teeth usually consisting of appendages of two adjacent cells; glands may be present
on or beside the teeth. Petioles close, less often remote,
not articulate at base, often ± persistently scaly, hairs
present besides or not, or exclusively so; vascular bundles two or more, depending on the subfamily. Lamina
often of an oblong, triangular, pentagonal, or ovate
type, simply pinnate to decompound, rarely simple,
the basal pinnae not rarely basitonically produced;
dissection pattern anadromous, catadromous, or often
anadromous at base and catadromous above, less often isodromous. Lamina glabrous, hairy, and/or scaly;
glands may be present in addition. Secondary rachises,
costae, etc. adaxially sulcate or not. Veins free, forkedpinnate, or variously anastomosing, then usually with
free included veinlets; veins mostly not reaching the
margin. Fertile leaves conform or weakly, less often
more strongly to completely difform. Sporangia very
often in approximately orbicular sori, dorsal or less often terminal or compital, or not rarely elongate, the indusium superior, lateral, or rarely inferior, or wanting.
Sporangial stalk 2- or 3-seriate; annulus vertical or
nearly so, the bow strongly interrupted; stomium weB
differentiated from annulus, usually 2-celled (exc.
Onocleeae). Spores without exception monolete, with
prominent perispore. Stomata polocytic; anomocytic
and/or staurocytic ones often present besides. Chromosome number most often 40 or 41.
MORPHOLOGY AND ANATOMY. The dictyostele of the
stem is radial, or weakly dorsiventral in taxa with
creeping stem; the stelar anatomy is an important
102
Filicatae . Dryopteridaceae
A
c
D
Fig.46A-E. Spores of Dryopteridaceae, Hymenophyllaceae,
and Lomariopsidaceae (all x 1000). A Polystichum muricatum. B Athyrium filix-femina. C Elaphoglossum pascoense.
D Lomariopsis hederacea. E 1Iichomanes pinnatum. Pho!.
A F. Tryon
character for delimiting the family from the related
Dava1liaceae and Lomariopsidaceae. Internally the
parenchymatous portions of the stem are often green.
The petiole bases are very often persistent and may be
developed as storage organs (trophopods); they sometimes bear adventitious buds. An erect, trunk-like stem
is found in a few cases, e. g., in species of Diplazium.
Matteuccia. and Polystichum. The vascular pattern of
the petiole is a character of taxonomic importance (see
under the subfamilies).
The dissection pattern of the lamina, whether anadromous or catadromous, or the one at the base, the
other in the upper part of the lamina, is an important
but as yet insufficiently studied character; it seems to
be constant in at least most species of a genus. Another taxonomically diagnostic character is the absence or presence of a groove on the adaxial side of
the leaf axes, its discontinuity or continuity on axes of
different order, etc., as stressed by Holttum (1947 and
after) but too little employed by other authors.
E
A tendency towards reticulate venation is found in
various tribes and even genera (but in some instances
free veins may be derived, e. g., in Tectaria); it must be
regarded as an independently derived feature, occurring in different groups, that is neither indicative of affinity nor sufficient for distinguishing genera, as is true
in many other leptosporangiate families.
Sterile-fertile leaf dimorphism also appears in various groups and, by itself, cannot be used for separating genera. As a whole it is rather weakly expressed in
the present family, except for the Onoc\eeae. A completely "gymnogrammoid" arrangement of sporangia
is rare in the present family, "acrostichoid" sporangia
even rarer (e. g., species of Tectaria and Heterogonium). In the Athyrioideae acrostichoid sporangia are
unknown, and even when the sori are very long the indusium is nearly always conserved (Diplazium, Dictyodroma).
The peculiar reddish, articulate hairs found in Ctenitis and Tectaria and their relatives ("ctenitoid hairs")
are taxonomically important; so are the equally articulate but pale and stiff, scale-like appendages found in
Peranema, Psomiocarpa, Stenolepia, etc. ("intestiniform hairs"). Both these and the ctenitoid hairs may be
morphologically nearer to scales than to true trichomes; the "intestiniform hairs" of Nothoperanema
are up to 3-seriate at their base. In the Athyrioideae
the indument is more diverse.
Filicatae • Dryopteridaceae
GAMETOPHYTE. This is fairly constant in shape, being
almost universally cordate, with simple, derived gametangia; there is more variation in the presence or absence and structure of superficial appendages which
may be hair-like, glandular, even branched (Atkinson
1973).
KARYOLOGY AND HYBRIDIZATION. The great constancy
in chromosome number is one of the features that
hold the family together (see, e. g., Sledge 1973 b).
Base numbers of 40 and 41 are almost equally frequent; some other numbers are found in the Athyrioideae. These two numbers may occur in apparently
quite closely related genera, and the numbers have actually been used for assigning borderline species to
one or the other (e. g., in Athyrium vs. Diplazium; Jermy 1964), but the naturalness of such a procedure is
open to doubt. Auto- and allopolyploidy are of very
frequent occurrence.
As in the Aspleniaceae, the fairly constant chromosome number in the family and in the majority of genera has resulted in the formation of hybrids between
species (groups) formerly placed in different genera;
evidence from hybridization in the present treatment is
often used as an argument for merging genera (see,
e. g., under Tectaria). There are even reports of hybrids
between genera that would scarcely be merged: the
first credible report of a Dryopterisx Polystichum hybrid (Wagner in Barrington 1985 b).
The incidence of intrageneric hybridization in the
family is again large, as in Asplenium; e. g., in Dryopteris (Chandler 1948; W. H. Wagner 1971; Fraser-Jenkins and Reichstein 1984; Gibby 1985; Fraser-Jenkins
1986), Polystichum (Sleep 1966; Daigobo 1974; Vida
and Reichstein 1975; D. Wagner 1979; Barrington
1985 a; Gibby 1985), etc. In the Athyrioideae fewer hybrids have been described, but many probably still
await discovery; see, e. g., Sarvela (1980) for Gymnocarpium. Sledge (1956) and Schneller and Rasbach
(1984) for Athyrium. Butters (1941) for Woodsia. Jermy
and Walker (1985) for Diplazium. See Knobloch's list
(1976) to which many more recent examples may be
added.
Apogamy is also fairly widespread; it is concentrated in, but not limited to, taxa that are hybridogenous
and have an odd 2n chromosome number; see, e. g.,
Gibby (1985) and Fraser-Jenkins (1986) for Dryopteris.
Vida and Reichstein (1975) and D. Wagner (1979) for
Polystichum, Lloyd (1971) for Matteuccia, etc.
ECOLOGY AND DISTRIBUTION. The great majority of
Dryopteridaceae grow on the ground or on rocks,
especially in temperate and montane areas. Low
epiphytes occur occasionally, but only a few species of
Rumohra have been described as true habitual epiphytes. Many species occur in forests and thickets, at
103
higher elevation often in more exposed places. See also under the genera.
The family is of cosmopolitan distribution; many of
the larger, more conspicuous ferns of especially nontropical areas belong to it, beside Dennstaedtiaceae
and Thelypteridaceae. There is a strong concentration
of genera and species in north-temperate regions for
Onocleeae, Woodsieae, Dryopterideae, and certain
genera of Physematieae; Tectarieae and other Physematieae are preponderantly tropical. The by far greatest
species concentrations of most non-tropical genera are
in the hills and mountains of east-central, eastern, and
south-eastern Asia. Comparatively few genera are species-rich in tropical lowlands, e. g., Tectaria, Triplophyllum. and Diplazium. Genera like Woodsia. Cystopteris.
Dryopteris. Polystichum. and Athyrium extend to very
high latitudes.
AFFINITY. Physematieae (the ''Athyrium group" of genera) were formerly confused with and included in the
Asplenioid ferns, whereas the Thelypteridoid ferns
were classed with the "Aspidioid" group (e.g., as late
as 1938 by Christensen). Such ideas have now been
abandoned, at the latest with the advent of karyology,
and need not be refuted here, although they still linger
on in some floras. Relationships to Davalliaceae, with
Oleandraceae to a certain extent as intermediates, are
accepted by most workers; the same is true for the
Lomariopsidaceae which are sometimes included,
which can well be defended. Blechoaceae, too, are
often placed near Dryopteridaceae, on anatomical
grounds, but this is less obvious and less generally accepted. Inclusion of the Dryopteridaceae in a broadly
circumscribed family Aspleniaceae (e. g., Crabbe and
Jermy 1975) seems much less advisable, in view of very
considerable differences in morpho-anatomical and
karyological respects (see under Aspleniaceae). While
admitting a close relationship, some taxonomists exclude the Physematieae; Onocleeae and Woodsia may
then also be raised to family rank. This, however, is a
matter of opinion. Following Sledge (1973 b), these
subfamilies and tribes are here included in one more
comprehensive family. A few genera have been classed
with both the Dryopteridoid and the Athyrioid ferns,
viz. Gymnocarpium (also placed in Thelypteridaceae!),
Adenoderris. and Hypodematium. They certainly occupy somewhat isolated positions within the Dryopteridaceae and their affinity is not known with certainty. In
a key to the genera, Dryopteridoid and Athyrioid genera tend to be mixed, again testifying to their close relationship.
The subdivision adopted here is essentially based on
Holttum's (1947), with some changes:
Subfamily Dryopteridoideae: petiole with at least 3,
mostly more, vascular bundles arranged in a semi-circle; spores achlorophyllous.
104
Tribe Rumohreae:
(genus 1)
Dryopteridaceae . Dryopteridoideae
stele of stem strongly dorsiventral; some scales peltate; costae
adaxially flat; n =41.
Tribe Dryopterideae: stele of stem radial or nearly so;
(genera 2-15)
scales all basally attached; costae adaxially sulcate; ctenitoid
hairs none; n =41.
Tribe Tectarieae:
(genera 16-29)
stele and scales as in the last;
costae adaxially non-sulcate;
ctenitoid hairs often present;
n=40 or 41.
Subfamily Athyrioideae: petiole with two strap-shaped
vascular bundles upward merging into one guttershaped one; n = usually 40 or 41.
Tribe Physematieae: leaves not or weakly dimorphic;
(genera 31-42)
sori not covered by the strongly
revolute margin of the fertile
segments; spores achlorophyllous
Tribe Onocleeae:
(genera 43 -45)
leaves very strongly dimorphic;
fertile leaves with strongly reduced laminal parts, their edges
recurved and enclosing the sori;
spores chlorophyllous
(genus 30, Adenoderris. is "incertae sedis").
CHEMOTAXONOMY. Partly because of the toxicity of at
least some Dryopteris species and their former use for
medicinal purposes, chemotaxonomic studies have
been focused on Dryopteris and related genera. They
have also yielded much important evidence for the derivation of allopolyploids and F, hybrids. For yielding
evidence for classification on a higher level the data
are still too scanty. The distribution of phloroglucinols,
widespread in Dryopteridoid and lacking in Athyrioid
genera, confirms the distinctness of these two groups
(Widen et al. 1976 b). For further data, see, e.g., von
Euw et aI. (1980); Widen et al. (1970, 1975, 1976 b,
1978,1981), Gibby et aI. (1978).
CHARACTERS OF RARE OCCURRENCE (see also under
Dryopteris and Polystiehum). Epiphytism (not quite
consistent) in Rumohra; an articulation in the petiole
above its base in some species of Woodsia. An inferior
indusium in Woodsia. Peranema. and Hypode"is. Articulations at pinna (pinnule) bases in eye/opeltis and
Didymochlaena. A "filmy" lamina in a species of Cys-
topteris.
TAXA LIKELY TO BE CONFUSED WITH DRYOPTERIDACEAE. As stated above, Thelypteridaceae were included
in Dryopteridaceae for a long time. Their soral structure is often strikingly similar to that of the Dryopter-
idoideae, but the vascular anatomy of the petiole is
more like that of the Athyrioideae. The combination of
hairiness (sometimes very scanty) and catadromy of
the lamina in Thelypteridaceae will usually render distinction easy. Certain species of Bolbitis may look
much like Teetaria when sterile; the venation patterns,
sagenioid or anaxetoid, are strikingly similar. The ctenitoid hairs mostly present in Teetaria and the adaxially
grooved costae of at least some Bolbitis species help to
avoid confusion.
Simple-leaved species of Teetaria may be taken for
Polypodiaceae; but stem morphology and anatomy,
scales, and spores are entirely different.
The genus Thysanosoria of the Lomariopsidaceae is
difficult to exclude from Dryopteridaceae except by
anatomical characters and is therefore included in the
following key to the genera.
KEy TO THE GENERA (INCLUDING ATHYRIOIDEAE).
[A separate key to the genera of Athyrioideae is provided on p.131]
1. Lamina of mature plauts simple or shallowly to deeply
lobed, not once fully pinnate
2
- Lamina of mature plants at least at base with one to mauy
free pinnae
8
2. Veins free
3
5
- Veins reticulate
3. Sori orbicular. with (sub)peltate indusium
4
- Sori elongate. with lateral indusium
32. Diplazium
4. Lamina glandular-hairy
30. Adenotkrris
- Lamina essentially glabrous or scaly. not glandular-hairy
52
5. Sori orbicular, with inferior indusium
19. Hypoderm
- Sori not orbicular, or, if so, not with inferior indusium
6
6. Sori linear, indusiate
33. DictyodrtmUJ
- Sori orbicular to elliptic, or, iflinear, exindusiate
7
7. Lamina glabrous
45. Onocletl
- Lamina hairy, at least in the adaxial axis grooves
16. Tecta";"
8. Sporangia assembled in round to elongate sori of definite
size and shape, usually indusiate
13
- Sporangia following the veins, not assembled in sori of
definite size and shape; or not arranged in sori, acrosti-
choid (beware of ferns with definite sori that may appear
confluent with age!): indusium none
9
9. Costae of ultimate divisions adaxially convex or flat;
sporangia variously arranged
10
- Costae of ultimate divisions adaxially sulcate; sporangia
acrostichoid, on segments with much reduced lamina 12
10. Adaxial leaf surface bearing intestiniform hairs between
the veins (as in Fig. 48 0)
20. Psomiocarpa
- Adaxial leaf surface without such hairs
11
11. Lamina pinnate + pinnatifid: basal segments oflowermost
17. Heterogonium
pinnae reduced
- Lamina with different dissection (or simple); or, if pinnate + pinnatifid, basal segments of lowermost pinnae the
largest
16. Tecta";"
12. Lamina simply pinnate, with conform terminal pinna:
veins joined by a submarginal commissure, otherwise
free; stem creeping
14. Olfersia
- Lamina more than once pinnate, or, if not so, without
confonn tenninal pinna; veins free or anastomosing but
Key to the Genera
not joined by an intramarginal commissure; stem mostly
scandent
15. Poly/Jotrya
13. Sterile and fertile leaves strongly difform; sori indusiate
and also enclosed by the revolute, narrow laminal parls of
the "skeletonized" fertile lamina (Onocleeae)
74
14
- Without this combination of characters
14. Indusium inferior
15
- Indusium superior, lateral, or wanting
17
15. Indusium attached only at the posterior (basal) side of the
41
receptacle
- Indusium surrounding the receptacle with its base
16
16. Lamina simply pinnate or pinnate+pinnatifid, more or
less tapering at base; basal pinnae equal-sided; petiole often with an articulation above its base
42. Woodsia
- Lamina at least bipinnate + pinnatifid, broadest at base;
basal pinnae with the posterior side wider than the anterior; petiole without an articulation
3. Peranema
17. Indusium wanting or minute and fugacious; sori approximately orbicular
18
- Indusium present, distinct, or, if absent, the sori elongate
37
18. Basal pinnae (segments) with an articulation at base
40. Gymnocarpium
- All pinnae except the terminal with an arliculation at base
Thysanosoria (l.omariopsidaceae)
- No pinna bases with an articulation
19
19. Veins at least partly anastomosing
20
- Veins quite free
24
20. Anastomoses of veins goniopteroid
23
- Anastomoses of veins sagenioid or anaxetoid; or only
costal/costular areoles present
22
21. Costae adaxially convex
29. CoW!nklla
- Costae adaxially sulcate
23
22. Sinus bottom with a tooth, a free excurrent veinlet termi18. Pleocnemia
nating behind it
- No such tooth with a vein end present
11
23. Lamina and venation preponderantly anadromous or
isodromous; leaf tissue not pellucid-punctate
13. Cye/odium
- Lamina and venation preponderantly catadromous; leaf
tissue pellucid-punctate
12. Stigmatopteris
24. Indusium minute, replaced by a reflexed marginal lobe
42. Woodsia ("arn/anthopsis'~
No such lobes protecting the sori
25
25. Costae adaxially distinctly unisulcate
26
- Costae adaxially obscurely concave. flat, or convex, or, if
apparently grooved. the groove with a ridge along its cen~
~
26. Bases of costae and costules adaxially corniculate with
37. Comopteris
fleshy outgrowths
- No such outgrowths present at costa and costule bases 27
27. Lamina preponderantly anadromous or isodromous
29
- Lamina preponderantly and distinctly catadromous, except often for the basal pair(s) of pinnae
28
28. Lamina pellucid-punctate; veins simple 12. Stigmatopteris
- Lamina not pellucid-punctate; veins usually forked
6. Dryopteris
29. Petiole with two vascular bundles, these merging above
31. Athyrium
- Petiole with at least three vascular bundles. in transection
30
in a semi-circle, the two adaxial larger
30. Stem erect
10. Polysticlwm
- Stem creeping
31
31. Indusium with few exceptions peltate; lamina usually
simply pinnate or pinnate + pinnatifid. rarely more dis-
105
sected, not basitonically produced; axes and costae minutely hairy in the adaxial grooves
13. Cye/odium
- Indusium deeply reniform, not peltate; lamina often more
dissected. the basal pinnae mostly basitonically produced; adaxial grooves of rachises lacking hairs
7. Arachniotks
32. Septate (ctenitoid or not) or acicular hairs present on axes,
sometimes only sparingly so
34
- Axes non-hairy
33
33. Sinus bottom with a projection (tooth) (Fig. 62 B); lamina
not pellucid-punctate
27. heridrys
- Sinus bottom without a tooth; lamina pellucid-punctate
12. Stigmatopteris
34. Stem short, (sub)erect; ctenitoid hairs present
35
- Stem creeping
36
35. Scales on leaf axes at least partly clathrate; unicellular,
cylindric glands present on lamina, young indusia. etc.
22. Ctenitis
- Scales of axes (if any) non-clathrate; such glands absent
17. Heterogonium
36. Lamina with the basal pinnae much basitonically produced
58
- Lamina regularly bipinnate + pinnatifid to tripinnate, not
basitonically produced at base; articulate (but not ctenitoid) hairs present
39. Acystopteris
37. Sori elongate, either exindusiate, or a false indusium
formed by the reflexed margin; veins reticulate
11
- Sori approximately orbicular, or, if elongate, with a "true"
indusium
38
38. Soms and indusium about orbicular
40
- Soms and indusium elongate
39
39. Lamina regularly bipinnate. with conform terminal (primary) pinna; at least the lower pinnules arliculate; all
pinnules dimidiate (Fig. 52 A, B)
11. Didymochlllena
- Without this combination of characters
76
40. Indusium attached at the posterior (basiscopic) side of the
receptacle
41
- Indusium (sub)centrally attached, reniform, horseshoeshaped, or peltate
44
41. Scales of petiole and leaf axes clathrate, the larger ones
inserted on verrucae
4. Stenolepia
- Scales neither clathrate nor inserted on verrucae
42
42. Lamina minutely hairy to glabrous
38. Cystopteris
- Lamina bearing evident hairs or hair-like scales
43
43. Intestiniform, hair-like appendages borne adaxially on the
lamina; pinnae and larger pinnules opposite; flaccid,
·subpersistent scales present at attachment points of pin2. Acrophorus
nae and pinnules (Fig. 48 A. D, G)
- Acicular and/or glandular hairs present on lamina; only
the lowermost pinnae (sub)opposite; no scales present at
pinna and pinnule insertions
41. Hypodematium
44. Indusium peltate, centrally attached
45
- Indusium reniform
54
45. Lamina simply pinnate. the pinnae articulate at base
28. CycIope/tis
- Lamina, if simply pinnate, without articulation at pinna
bases
46
46. Cost.e adaxially flattened or convex
47
- Costae adaxially distinctly grooved (unisulcate)
48
47. Stem long-creeping, with dorsiventral dictyostele; petiole
not dark; some scales of stem with peltate base; lamina
not hairy, anadromous, free-veined
1. Rumohra
- Stem (mostly short-)creeping or erect. with radial dictyostele; all scales basally attached; petiole often dark-scle-
106
Dryopteridaceae • Dryopteridoideae
rotic; lamina anadromous only at base, catadromous
above; axes (and veins) usually bearing ctenitoid hairs
16. Tectaria(see also 30. Adenoderm)
48. Lamina catadromous
49
- Lamina anadromous or isodrornous
51
49. Leaf axes hairless; laminal parts pellucid-punctate
12. Stigmatoptem
- Leafaxes scaly or hairy; laminal parts not pellucid-punctate
50
50. Leaf axes bearing ctenitoid hairs
26. Dryopolystichum
- Leafaxes scaly
10. Polystkhum
51. Lamina hairy; stem long-creeping or -scan dent; fertile
lamina strongly skeletonized
9. Maxonia
- Lamina not hairy (but may bear narrow scales); stem not
scandent, rarely long-creeping; fertile lamina not or weakly reduced
52
52. Laminal parts aristate at edge (Fig. 51 C, D, E); lamina often bipinnate or more dissected; veins free; sori uniseriate
10. Poly3tichum
- Edge of laminal parts entire to serrate, not aristate; sori
bi- to pluriseriate
53
53. Edge of laminal parts serrate; lamina simply pinnate;
veins free or casually anastomosing
10. Polystkhum ("Cyrtomium',
- Laminal parts entire or lobed, or serrate towards the apex
only; veins free; or goniopteroid-anastomosing
13. Cye/odium
mostly strongly basitonically produced; intestiniform
21. Trip/ophyl/um
hairs present or not
- Stem relatively stouter, ascending or erect, with aggregate
64
leaves
64. Scales (at least partly) clathrate; intestiniform hairs absent
22. Ctenin.
- Scales non-clathrate; intestiniform hairs present on adaxial surface (Fig. 61 A-C)
23. Aenigmoptem
65. Pluricellular, basally thickened hairs present on margins
of adaxial side of rachis grooves; adaxial pinna-rachis
grooves starting above their bases, thus not continuous
with the groove on the primary rachis
24. Dryopsis
- No such hairs present, but intestiniform hairs may be
present
66
66. Costae adaxially sulcate
67
- Costae adaxially flattened or convex
56
67. Lamina simply pinnate, with entire to lobed pinnae
68
- Lamina more dissected
70
68. Petiole base with two vascular bundles
31. Athyrium ("Anisocampium'')
- Petiole with more than two vascular bundles, these arranged in a semi-circle in transection
69
69. Stem erect, or, if creeping, the divisions not with sharply
serrate-aristate edge
6. Dryoptem
- Stem wide-creeping; laminal divisions with sharply serrate-aristate margin
7. Arachniodes
54. Lamina scaly or not, glabrous or bearing ctenitoid hairs,
largely or entirely catadromous; petiole with more than
two vascular bundles, not swollen at base
56
- Lamina bearing pale hairs (scales may be present), dissection pattern various; petiole with two vascular bundles,
often swollen or broadened at base
55
55. Lamina largely or entirely anadromous, bearing acicular
hairs; petiole swollen at base; stele of stem dorsiventral
70. Lamina anadromous, usually rigid; stem usually creeping
71
- Lamina catadromous, or the basal pinnae (rarely most
pinnae) anadromous; texture usually herbaceous; stem
erect or ascending
72
71. Indusium very firm, rupturing at maturity; ultimate segments very narrowly elliptic, almost linear, with entire
margin (Fig. 50 C); edge of ultimate leaflets decurrent onto the margin of the rachis groove
8. Litlwstegia
41. Hypodematium
- Indusium not so firm, not rupturing at maturity; ultimate
- Dissection pattern of lamina various; hairs pale, septate,
not acicular; petiole broadened at base or not; stem stele
36. Deparia
radial
56. Pinnae pinnatifid; sinus bottom with a lobe containing a
57
veinlet (Figs. 57 B, 62 B)
- Pinnae not incised, Of, if so, the sinus without a lobe containing a veinlet
58
57. Veins free
27. PteriJirys
- Veins anastomosing with at least costal/ costular arches
18. Pleocnemill
58. Veins copiously reticulate
11
- Veins free or with some casual anastomoses
59
59. Lamina with the basal pinnae basitonically produced;
60
veins free or casually anastomosing
- Lamina without this dissection pattern; veins free
65
60. Posterior edge of ultimate free (or adnate) divisions thickened at base and running into the lateral rachis ridge on
25. Lostreopsi.
its side (Fig. 61 D)
- Posterior edge of ultimate free (or adnate) divisions not
segments not so narrow, very often sharply dentate or
aristate, their posterior side adaxially somewhat thickened
inside the margin, this thickening continuous with a ridge
7. Amchniodes
bordering the rachis groove.
thickened at base and tenninating on the axis on the same
side outside (i. e., below, as seen from the adaxial side) the
lateral rachis ridge
61
61. Axes bearing ctenitoid hairs
62
- Ctenitoid hairs wanting
66
62. Vein of basal posterior segment springing from the same
63
vein as those of the segments just distal from it
- Vein of basal posterior segment springing from the vein of
11
one order below
63. Stem slender, creeping, with spaced leaves; basal pinnae
72. Adaxial side of lamina bearing intestiniform, hair-like
scales
73
_ Lamina without such scales, but narrow, hair-like (not rigid and intestiniform) scales may be present 6. Dryoptem
73. Adaxial pinna-rachis groove continuous with the groove
5. No/hopelY1llema
on the primary rachis
- These grooves not continuous, the pinna-rachis groove
not extending to base
24. Dryopsis
74. Fertile segments (sub)globose; veins of sterile lamina reticulate (Fig. 73)
75
- Fertile segments linear; veins free (Fig. 72 A)
43. Matteuccill
75. Lower pinnae not reduced; indusium distinct, cup45. Onoclea
shaped; fertile leaves indurated, persistent
- Lower pinnae reduced; fertile leaves not indurated, not
persistent; indusium tender, fugacious
44. Onocleopsis
76. Multicellular hairs mostly present on leaf axes ; adaxial
groove of costae (or secondary rachises) not continuous
with that of the primary rachis
36. Deparia
- Multicellular hairs mostly absent; adaxial groove of secondary rachises or costae continuous with that of the primary rachis
77
77. Adaxial rachis groove with narrow bottom (V-shaped in
transection); sori orbicular, U- or J-shaped, or, if linear,
on only one side of a vein (not "double"); veins free
31. Athyrium
Rumohra
107
- Adaxial rachis groove with broad bottom (V-shaped in
transection); sori linear, nearly always at least some on
both sides of a vein ("double"); veins free or anastomos~
n
78. Veins free or occasionally goniopteroid-anastomosing
32. Dip/adum
- Veins at least partly sagenioid-anastomosing
79
79. Veins anastomosing only in the outer (marginal) parts of
the pinnae, free near the costa; lamina simply pinnate
with conform terminal pinna
80
- Veins wholly anastomosing, or preponderantly so near the
costa; lamina usually with pinnatifid apex (always so if
veins anastomosing only distally)
81
80. Vein ends connected by an intramarginal commissure; indusium diplazioid (Fig. 68 E)
35. Hemidktyum
- Vein ends free, or connected in pairs; indusium allanto-
dioid (Fig. 68 D)
34. DiplazWpsis
81. Veins ending near the margin; both parts of a "double"
sorus of about equal length
32. Diplazium
- Veins ending well behind the margin; both parts of a
"double" sorus in many cases of unequal length and thus
only partly "back to back" (Fig. 67 B)
33. Dictyodroma
1. Rumohra Raddi
Fig. 47
Rumohra Raddi, Opusc. Sci. Bol. 3: 290 (1819); Ching (1934)
(p. p. min.); Tardieu-Blot (1958).
Terrestrial, epilithic, or low-epiphytic ferns with longcreeping stem bearing lanceolate, basally attached,
and also some peltate scales, or sometimes eventually
scaleless; stele a dorsiventral dictyostele with several
anastomosing bundles in each leaf gap that run to the
petiole. Leaves borne in two rows dorsal on the stem;
petiole non-articulate, well developed, at least basally
usually persistently fuscous-scaly and minutely warty,
adaxially unisulcate, the groove with raised centre;
vascular bundles several, in a semi-circle, the larger,
adaxial ones hippocampi form. Lamina oblong-deltoid, usually firm, bipinnate+ pinnatifid or more dissected, anadromous; rachis like the petiole, scaly to
glabrescent. Pinnae inserted on the lateral ridges, the
grooves of their axes ± continuous with that of the primary rachis. Divisions of higher order anadromous, or
occasionally some catadromous. Primary pinnae
stalked. Free divisions of highest order lanceolate,
lobed-pinnatifid, their edges running into the lateral
ridges of their axes, costae adaxially flat. Veins free,
pinnate in the lobes; veinlets forked, almost reaching
the margin. Surfaces naked, or often the axes and larger veins bearing some filiform scales. Sori large, round,
dorsal or terminal on the veins, in two rows flanking
the costae and costules, with persistent or deciduous,
peltate, orbicular indusium. Spores ellipsoidal to
somewhat globose, with prominent folds or shorter, inflated tubercles; n=41.
Seven described species retained in the genus (the
remainder placed in Arachniodes); one, R. adiantiformis (Forst. f.) Ching, of circum-austral distribution,
north to Bermuda, Zimbabwe, Madagascar, New Gui-
Fig. 47 A-D. Dryopteridaceae. Rumohra adiantiformis.
A Habit ( x 0.25). B Pinna (x 0.6). C Sorus (x 9). D Petiolar
scale (x 10) (Tardieu-Blot 1958)
nea, and Polynesia; one endemic to Juan Fernandez;
the others in Madagascar. In forests and open places;
some species said to be obligate epiphytes; one Madagascan species apparently an obligate epiphyte of
Pandanus, with naked stem.
Holttum (1947) and others showed Rumohra to be
distinct from Arachniodes, and how to separate them.
Rumohra was variously included in the Davallioid
(Holttum 1947; Tardieu-Blot 1958) or Aspidioid ferns,
108
Dryopteridaceae . Dryopteridoideae
as it has characters in common with both alliances.
Kato's (1974) arguments for inclusion in the latter
group seem strongest, and the chemical evidence presented by Widen et al. (1983) bears out this conclusion. The perispore structure is certainly unlike anything known from Davalliaceae. The affinity of Dryopteridaceae with Davalliaceae is confirmed by the
characters of Rumohra.
2. Acrophorus C. Presl
Fig.48A-G
Acrophorus C. Presl, Tent. Pteridogr.: 93 (1836); Nayar and
Kaur (1963); Wu (1983).
Medium-sized to large terrestrial ferns with short,
erect, dictyostelic stem bearing broad, brown, marginally hairy, entire scales. Leaves close; petiole well-developed, stramineous, with 6- 8 vascular bundles;
adaxial side 1- or 3-sulcate; base± persistently scaly,
scales when shed often leaving a small, wart-like scar.
Lamina deltoid, highly compound, tripinnate+pinnatifid to quadripinnate + pinnatifid, with at least the
larger pinnae (sub)opposite; rachis stramineous to medium brown, adaxially like the petiole, swollen at the
insertions of the larger pinnae whose bases bear a
dense, dark brown indument of small, hair-like scales
(see below); similar, less developed structures on the
secondary axes. Dissection pattern catadromous. Pinnae and pinnules sessile; basal divisions of larger pinnae and pinnules somewhat reduced; divisions of
higher order gradually alternate; basal pinnae broader
on the posterior side; pinnae and pinnules strongly
spreading. Bases of pinna and pinnule rachises abaxially each with a single, large, broad, flaccid, brown,
persistent, appressed scale. Leaf and pinna apices
gradually reduced, with confluent upper divisions.
Small, several-celled hairs abundant on (young) lamina; narrow, appressed, brownish, hair-like, largely uniseriate scales scattered on the adaxial leaf surface, especially on rachidules, costae, costules, and veins,
abundant on the swellings at the pinna insertions. Rachises adaxially grooved, the grooves evanescing at
pinna/pinnule insertion points. Rachidules adaxially
narrowly 2-winged Ultimate divisions tender, asymmetrically oblong-ovate, obtuse, crenate to lobed.
Veins free, forked, with clavate endings behind the
margin. Sori dorsal-subterminal, round; indusium tender, ± semicircular, very shallowly cup-shaped, basally
attached, reflexed at maturity. Sporangial stalk often
bearing few-celled hairs. Spores brown, monolete, elIipsoidal; perispore with long-prominent folds, sometimes perforate; 2n = 82.
Two closely related species (6 recognized by Wu
1983), A. nodosus C. Presl and A. blumei Ching in
C. Chr., in moist mountain forests from NE India and
southern Japan to Fiji.
3. Peranema D. Don
Fig. 48 H-J
Peranema D. Don, Prodr. Fl. Nepal.: 12 (1825) (non Dujardin
1841, Euglenophyta); Nayar and Kaur (1963).
Diacalpe Blume, En. Plant Jav.: 241 (1828); Wu (1983).
Medium-sized to rather large ferns; stem erect, stout,
dictyostelic, densely vested in brown scales with minutely hairy margins. Petioles close, well-developed,
with several vascular bundles, the adaxial side sulcate.
Lamina oblong-triangular to subpentagonal, tripinnate
to quadripinnate, firm, catadromous except for the
basal primary divisions; axes, especially petiole base,
± persistently scaly, often only the wart-like scale
bases persistent; sometimes also bearing minute, several-celled hairs, adaxially grooved, the grooves on axes
of different order not connected. Basal pinnae subopposite; pinnae and pinnules strongly spreading; basal
pinnae usually broader on the posterior side. Segments
oblong, adnate or decurrent, obtuse, crenate to lobed;
upper divisions reduced and gradually confluent.
Veins free, forked to pinnate, with clavate ends behind
the margin; axes of higher order, and adaxial side of
costae, costules, and veins, bearing intestiniform
scales. Sori dorsal, round, sessile or with a slender
stalk up to a few mm long; indusium inferior, at first
globose, enclosing the sporangia, later bursting irregularly from the apex, the opening near the anterior side.
Sporangium stalks often bearing unicellular hairs.
Spores ellipsoidal; perispore with prominent folds;
x=41.
Three or four species (8 according to Wu 1983), terrestrial in mountain forests, from north-eastern India,
southern China, and Taiwan to Malesia. For the congenerity of Diacalpe and Peranema, see, e. g., Sledge
(1973 a).
4. Stenolepia v. A. v. R.
Slenolepia v. A. v. R., BUll. Dep. Agric. Ind. Neerl. 27: 45
(1909).
Rather large terrestrial ferns; stem erect, stout, bearing
narrowly lanceolate, long-acuminate, non-clathrate,
tan scales. Leaves clustered; petiole well-developed,
stramineous to medium brown, densely scaly at base,
the scales thinning upward, narrow, clathrate, grading
into filiform, trichome-like ones; such scales also present on the smaller leaf axes ; larger scales inserted on
dark warts, the axes thus dark-verrucose. Lamina deltoid, chartaceous-subcoriaceous, tripinnate + pinnatifid to quadripinnate at base; lower pinnae subopposite, short-stalked, the upper alternate, (sub)sessile;
upper pinnae, pinnules, etc. gradually reduced and
confluent. Basal primary divisions anadromous, lamina otherwise ± distinctly catadromous. Axes adaxially
sulcate, the grooves on the axes of second and higher
order evanescing at base. Pinnae caudate, pinnules ac-
Acrophorus. Peranema . Steno[epia. Nothoperanema
109
H
J
Fig. 48A-J. Dryopteridaceae. A-G Acrophorus nodosus.
A Habit (x 0.2). B l\vo pinnules (x 2). C Sorus (x 10).
D Part of rachis with bases of two pinnules and scales
(x 0.5). E Scale from pinna base (x 1.5). F Scales from
petiole (x 4). G Multicellular hairs (x 15). H-J Peranema
cyatheoides. H Fertile pinnule (x 1). I Segment with sorus
(x 4). J Single sorus (x 8) (Hora Taiwan 1975)
uminate, segments usually obtuse to subacute, sinuate
to pinnatifid. Leaf tissue naked; costules abaxially
with scattered, intestiniform scales (like the smallest of
the leaf axes); venation abaxially dark. Veins forked,
almost reaching the margin. Sari dorsal on anterior
vein branches, orbicular; receptacle elevated. Indusium approximately orbicular, affixed at the posterior
side of the receptacle, later reflexed and caducous.
Spores ellipsoidal, surface bearing prominent, coarse
folds; n~?
A single species, S. tristis (BI.) v. A. v. R, in central
and eastern Malesia, in mountain forests at middle
and higher elevations.
Closely related to Peranema, perhaps not generically distinct. The dark-verrucose axes and the fugacious
indusia are distinctive, but the two genera are sometimes confused in the herbarium.
5. Nothoperanema (Tagawa) Ching
No/hoperanema (Tagawa) Ching, Acta Phytolax. Sin. 11: 25
(1966).
Dryopteris subgen. Nothoperanema Tag'w" Acta Phytol,x.
Geobot. 7: 199 (1938).
110
Dryopteridaceae • Dryopteridoideae
Medium-sized terrestrial ferns; stem erect or ascending, bearing dark, thick, glabrous, entire scales. Petioles close, well-developed, stramineous to medium
brown, ± persistently scaly. Lamina tripinnate or in
addition lobed to pinnatifid, herbaceous to subcoriaceous, catadromous except for the basal pinnae; lower
pinnae the largest, subopposite, the basal pair basitonically produced. Segments of a Iiguliform-oblong type,
obtuse, subentire to pinnatifid; leaf axes often persistently scaly, adaxially sulcate, the grooves of axes of
higher order evanescing at base. Veins free, simple or
forked, ending well behind the margin. Sori dorsal to
subterminal, round, with reniform indusium or this
rarely lacking. Adaxial surface of lamina bearing reddish-brown, vermiculiform trichomes composed of
subquadratic cells, especially at the bases of costae
and costules. Spores ellipsoidal to somewhat globose,
perispore with prominent folds, these sometimes shorter and coarsely tuberculate; n - 41.
Five described species (Ching 1966) but probably
only two or three; in India, China, Taiwan, and Japan,
one of them, N. squamisetum (Hooker) Ching, ranging
to Africa and Madagascar. Terrestrial in shady, moist,
montane forests.
6. Dryopteris Adans.
Fig. 49
Dryopteris Adans., Fam. Plants 2: 20, 551 (1763), nom. cons.;
Christensen (1911, 1913); Ching (1936-38); Sledge (1973);
Serizawa (1976); Price (1977); Fraser·Jenkins (1986, 1989).
Nephrodium Richard in Marthe ex Michaux, Fl. Bor.-Amer. 2:
266 (1803).
Acrorumohra (H. Ito) H. Ito in Nakai & Honda, Nov. Fl.
Jpn.4: 101 (1939), based on: Rumohra sect. Acrorumohra
H. Ito, J. Jpn. Bot. 11 : 583 (1935).
Aspidium auctt., p. p., excl. type.
Terrestrial ferns with ascending or erect, rarely creeping, usually thick, often branched stem clothed with
old leaf bases and bearing light brown to fuscous, nonclathrate, usually (sub)entire, glabrous scales at apex.
Leaves mostly close and in a rosette, medium-sized to
large, rarely smlill; petiole short to well-developed,
scaly at least at base, adaxially sulcate like the rachis
and other leaf axes. Lamina pinnate + pinnatifid to
quadripinnate, rarely more highly compound, in a few
species once pinnate with undissected pinnae, usually
catadromous, at least in the upper part of the lamina,
occasionally largely or entirely anadromous, or isodromous, triangular, oblong or lanceolate, sometimes pentagonal in outline, then with basitonically produced
basal pinnae, most often herbaceous; pinnae usually
(sub)sessile if lamina less than twice pinnate; upper
pinnae, pinnules, etc. gradually reduced and confluent, or in some simply pinnate species a conform terminal pinna present. Segments not rarely asymmetric
at base or (sub)auriculate anteriorly, very often adnate
and ± decurrent; margin entire or usually toothed with
obtuse to hair-pointed teeth; segment apex usually
without a terminal arista or hair point. Surface glabrous or variously scaly, especially the axes and larger
veins, often glandular but never hairy, but sometimes
the scales very narrow and hair-like; or sometimes bullate scales present. Adaxial grooves of pinna and/or
pinnule axes most often not continuous with those of
the primary rachis. Costa adaxially sulcate; edges of
leafy parts at base not running into the ridges bordering the rachidule groove but ending on or decurrent
onto the lateral face of the rachidule. Fertile leaves or
divisions in a few species somewhat contracted. Sori
sometimes confined to about the apical half of the
lamina. Sori orbicular, usually dorsal (rarely terminal),
with a reniform to cordate, often convex indusium
which may be glandular, or sometimes the indusium
deciduous or wanting. Spores ellipsoidal, with prominent long folds or shorter inflated tubercles and slightly rugulose, echinate, or spinulose; or the folds sometimes compressed into wings or short, cristate elements
with echinate borders; x - 41.
C. 225 species, some very complex due to hybridization, allopolyploidy, etc.; of subcosmopolitan distribution but rare or absent in tropical lowlands; most
often in forests and in open vegetation, sometimes on
rocks, often in montane to alpine zones, most strongly concentrated in the north-temperate to warm-temperate parts of Central, S. E., and E. Asia; comparatively well represented in montane and S. Africa and
Hawaii; only one species in Australia, none in New
Zealand.
Formerly, following Diels's and Christensen's work,
much more broadly circumscribed and including alliances now regarded as distinct genera like Stigmatopteris, Cye/odium, Nothoperanema, and Ctenitis, as well
as (nearly) all Thelypteridaceae.
SUBDIVISION, The first subdivision that was based on a
study of the entire genus is by Fraser-Jenkins (1986),
with four subgenera and 17 sections. It is as follows:
Subgenus Pycnopteris (Moore) Ching: Lamina simply
pinnate, with a conform terminal pinna, of firm texture.
Subgenus Dryopteris: Lamina variously incised, without conform terminal pinna, catadromous, subcoriaceous to herbaceous; scales of lamina non-bullate.
Eleven sections.
Subgenus Erythrovariae (H.lto) Fraser-Jenkins: Lamina as in the last; bullate or bullate-based scales present on the abaxial side of the leaf axes. Three sections.
Subgenus Nephrocystis (H.lto) Fraser-Jenkins: Lamina as in the two last but sometimes anadromous;
bullate scales none; pinnules strongly asymmetric.
Two sections.
111
CHARACTERS OF RARE OCCURRENCE. A slender, widecreeping stem in D. angustifrons (Moore) O. Ktze. An
entirely anadromous lamina in D. patula (Swartz) Underw. and D. subexaltata (Christ) C. Chr. Primary rachis zigzag, with somewhat deflexed pinnae, in section
Nephrocystis ( ~ genus Acrorumohra): D. diffracta
(Baker) C. Chr., D. hasseltii (BI.) C. Chr., etc. Proliferous buds on the rachis in D. manniana (Hooker)
Fig. 49A-E. Dryopteridaceae. A-C Dryopteris atrata.
A Leaf (x 0.4). B Part of pinna from abaxial side (x 2).
C Petiolar scale (x 8). D Dryopteris erythrosora. portion of
rachidule with bullate scales and segment with sori (x 5).
E Dryopteris lacera. transections of petiole and rachis with
vascular bundles (schematic) (A-D from Kurata and Nakaike
1985; E from Ogura 1972).
Dryopteridaceae . Dryopteridoideae
112
C. Chr. Somewhat contracted fertile segments in
D. eristata (L.) A. Gray, D. eoehleata (D. Don) C. Chr.,
etc. Sparingly but regularly present anastomoses of
veins in D. yakusilvieola Kurata (Damaedi and Iwatsuki 1986).
7. Arachniodes Blume
Fig.50A, B
Arachniodes Blume, En. Plant. Jav.: 241 (1828); Tindale
(1961 b);
Ching (1962); Ohwi (1962); Kurata (1962).
Leptorumohra H. Ito in Nakai and Honda (1939); Nakaike
(1970).
Polystichopsis (J. Smith) Holttum (1947), based on: Lastrea
subgen. Polystichopsis J. Smith (1875), nom. subnud.
Byrsopteris Morton (1960).
Phanerophlebiopsis Ching (1965).
Rumohra auctt., p. p., excl. type, e. g., Ching (1934), Copeland
(1947).
Dryopteris auctt., p. p., excl. type, e. g., Tryon and Tryon
(1982).
Medium-sized to large terrestrial ferns; stem longcreeping to short and ascending, or rarely erect, bearing non-clathrate, non-peltate scales with entire margin or teeth foimed by protrusions of two adjacent
cells. Leaves remote to close; petiole well-developed,
at least its base ± persistently scaly, not dark-sclerotic,
adaxially sulcate. Lamina usually bipinnate or bipinnate+pinnatifid, to more highly compound, rarely
simply pinnate, often deltoid or pentagonal, basally
truncate, chartaceous to subcoriaceous, glabrous or
with a few, narrow scales, rarely septate-hairy; dissection anadromous. Primary rachis like the petiole, often
with seattered, dark, small, fibril-like scales, adaxially
sulcate, the grooves continuous with those on axes of
higher order. Fertile lamina rarely somewhat contracted. Pinnae usually short-stalked, the basal ones often
basitonically produced. Upper divisions gradually reduced and confluent, the apex often acuminate or caudate or less often a subconform terminal pinna present. Ridges bordering adaxial grooves or rachidules,
costae, etc. not running into pinnule/segment edges
but continuous with ridges bordering grooves on costae, etc. of higher order (if any). Ultimate divisions sessile, crenate to pinnatifid, often asymmetrically ovatesubrhombic or elliptic, in many species resembling
those of Polystiehum species, the anterior base often
auriculate, the margin very often sharply dentate to
aristate; posterior side just inside the margin adaxially
somewhat thickened, this thickening (not the edge itself) continuous with the rachis ridge (difference from
Lithostegia and Lastreopsist). Veins immersed or ±
elevated, free, pinnate or forked, not quite reaching
the usually narrowly sclerotic margin; veins and axes
of higher order sometimes persistently scaly, the scales
flaccid, or occasionally bullate; rarely these parts long
septate-hairy. Sori orbicular, dorsal or terminal on the
veins, in the latter case often supramedial-submargi-
nal; indusium reniform-hippocrepiform, sometimes
fugacious. Spores ellipsoidal, with prominent folds or
shallow, inflated tubercles, often perforate and echinate, sometimes compressed into crests with echinate
borders; x=41.
A still imperfectly known genus, many species being
quite similar, others variable; the specific boundaries
are apparently also obscured by hybridization (see Kurata 1962). An estimated 50-70 species, very widely
distributed but much concentrated in the hills and
mountains of tropical to warm-temperate Central and
East Asia; a few species in tropical America; one extending to Australia, New Zealand, and the Pacific,
three to Fiji; only one, A.foliosa (c. Chr.) Schelpe, in
continental Africa, one in Madeira. Terrestrial or
epilithic, most often in forests and thickets, commonest at middle and higher elevations.
The distinctness of this genus is still disputed. In appearance it is rather like Po/ystiehum, with which it
was long united, in indusial characters like Dryopteris;
the pronouncedly anadromous architecture linked
with a creeping stem are distinctive. The character of
the apical tooth of the segments, emphasized by Fraser-Jenkins (1986, p.185), is not dependable, as it occurs in some Dryopteris species, too. The continuity of
all adaxial axis gro~oves is an important character but
is not unknown in Polystiehum.
Chemosystematic investigations did not clear up the
problem of generic distinctness; but phloroglucinols,
widespread in Dryopteris and rare in Polystiehum, have
also been found in Araehniodes (Widen et al. 1976 b,
1978, 1981, 1983).
The characters given for separating Phanerophlebiopsis Ching are on a specific level; the similarity of
"Ph." blinii (A. Lev.) Ching to Araehniodes assamiea
(Kuhn) Ohwi was already noted by Ching.
The few septate-hairy American species are often
segregated as Po/ystiehopsis; but, as pointed out by
Proctor (1985), this character is too fickle to be used
for delimiting a genus.
8. Lithostegia Ching
Fig.50C
Lithostegia Ching, Sinensia 4: 2 (1933).
Rather large terrestrial ferns; stem ascending, stout,
clothed in many brown, ovate, acuminate, shortly
fringed scales. Leaves close; petiole well-developed,
stramineous to reddish-brown, densely scaly at base,
scabrous by persistent bases of shed scales. Lamina
narrowly oblong, somewhat reduced at base, apex
acuminate, 4 or 5 times pinnate (+pinnatifid); axes
adaxially sulcate, bearing very narrow, fibril-like, twisted scales. Dissection anadromous; texture chartaceous. Upper divisions gradually reduced and confluent. Leaf divisions close, ± contiguous, ultimate divi-
Arachniodes· Lithostegia. Maxonia
113
B
Fig.50A-C. Dryopteridaceae. A,B Arachniodes maximowiczii. A Leaf (x 0.4). B Fertile pinnule ( x 2.5). C Lithostegia
joenicuiacea, fertile pinnule (x 2.5) (A, B from Kurata and
Nakaike 1979; C from Ching et aI. 1983)
sions very narrowly lanceolate, acute, with a single
vein terminating in a thickened end, not close to the
margin. Costules and veins abaxially bearing brown,
hair-like scales. Sori single in the segments, dorsal or
terminal on a short lateral vein serving a bulge or tooth
on the anterior margin of the segment; indusium very
firm, dark, entire, at first reniform and covering the sorus, later splitting into irregular segments. Spores ellipsoidal, bearing prominent, sometimes long folds; n =?
A single species, L.foeniculacea (Hooker) Ching, in
the eastern Himalaya and Yunnan, in mountain forests at higher elevation. The taxonomic position of the
genus is not quite clear, and it is placed only tentatively near Arachniodes.
9. Maxonia C. Christensen
Maxonia C.Christensen, Smiths. Misc. Coli. 66: 3 (1916);
Walker (1972); Chandra (1975).
Stem terrestrial becoming scandent, elongate, with almost radial dictyostele, the ventral meristele slightly
larger, bearing leaves on all sides, and clothed in golden-brown, basally attached, long and narrow, irregularly toothed scales. Leaves remote; petiole long-decurrent as a ridge on the stem, well-developed, adaxially unisulcate; vascular bundles many, typical of the
subfamily. Petiole and rachis with long, narrow, continuous pneumathodic streaks. Lamina large, triangular, basally truncate, 3- to 4-pinnate, anadromous,
114
Dryopteridaceae • Dryopteridoideae
firm, shortly pubescent and bearing a few narrow
scales, strongly dimorphic. Sterile lamina with trapeziform-Ianceolate, toothed to lobed segments; penultimate axes adaxially unisulcate, the grooves decurrent
into those of next lower order. Veins free, pinnately
branched, with simple branches that almost reach the
margin. Fertile leaves only produced on scandent
stems, like the sterile but with strongly reduced lamina; sori terminal on short veins, very close; indusium
hippocrepiform-cordate or (pseudo?)peltate; spores
ellipsoidal to somewhat spheroidal, bearing prominent
folds with rugulose to echinate elements; n =41.
A single species, M. apiifolia (Swartz) C. Chr., in
central and western South America, Cuba, and Jamaica; in forests and more open vegetation, at lower elevation. Apparently most closely related to Arachniodes.
10. Polysdchum Roth
Figs. 46, 51
Polystichum Roth, Tent. Fl. Germ.3: 31, 69 (1799): Maxon
(1908); Tagawa (1940); Kurata (1964); Looser (1968):
Sledge (1973 a); Nakaike (1982).
Cyrtomium C. Presl (1836); Christensen (1930); Ching (1936):
Morton (1957): Kurata (1963); Shing (1965).
Phanerophlebia C. Presl (1836); Underwood (1899); Maxon
(1912).
Plecosoms Fte (1852).
Papuapteris Christensen (1937).
Cyrtogonellum Ching (1938).
Cyrtomidictyum Ching (1940).
Acropelta Nakai (1953).
Medium-sized to rather large, less often small ferns,
usually terrestrial, less often epilithic; stem ascending
or usually erect, sometimes slightly trunk-like, often
massive, scaly; scales pale to blackish or bicolorous,
basifixed, entire to dentate or ciliate-spiny at the edge,
like those of the lamina axes often of great diagnostic
value. Leaves close, in a rosette; petiole very short to
well-developed, most often±scaly, adaxially unisulcate or with additional dorsolateral grooves, sometimes bearing several kinds of scales, stramineous or
dark. Lamina triangular and truncate, or very often
narrowly oblong to lanceolate and ± tapering at the
base, or linear, very often bipinnate or simply pinnate
(very rarely less dissected), or up to tripinnate+ pinnatifid, herbaceous or more often subcoriaceous to rigidly coriaceous, anadromous, or, when bipinnate, the
primary pinnae often isodromous, very rarely catadromous; rachis like the petiole, very often ± persistently
scaly; axes hardly ever dark-sclerotic. Secondary rachises, if present, adaxially grooved, the groove as a
rule not (quite) continuous with that of the primary
rachis. Upper pinnae, pinnules, etc. gradually reduced
and confluent, or in some simply pinnate species with
a (sub)conform terminal pinna. Rachis sometimes apically prolongate, rooting and proliferous: buds may
also be present elsewhere on the lamina. Ultimate free
divisions very often sessile or basally cuneate, shortstalked, mostly asymmetric at base, narrowly cuneate
on the posterior, broadly rounded or often auriculate
on the anterior side; margin sinuate or crenate to serrate or lobed, the teeth often aristate, even pungent.
Fertile leaves occasionally difform, with contracted divisions, or the lamina hemidimorphic, with the sori
confined to an eventually deciduous apical portion
with somewhat reduced laminal parts. Costa adaxially
grooved at least near its base; in bipinnate leaves this
groove often continuous with that of the secondary rachises. Veins immersed, pinnate-forked, anadromous
or isodromous, not quite reaching the margin, free or
less often anastomosing, then with oblique, elongate
areoles, at least the inner with excurrent included veinlets. Costae and veins glabrous or variously scaly,
some of the laminar scales, especially smaller ones, often very narrow and fibril- or hair-like. Sori dorsal or
also terminal, subcostal or submarginal or more often
about medial, or in several series, circular, with hardly
convex receptacle, in most cases with an orbicular, peltate, entire to laciniate, sometimes caducous, rarely
quite absent, very rarely reniform indusium. No receptacular appendages among the sporangia. Spores ellipsoidal or somewhat globose, with prominent, inflated
tubercles, often perforate and spinulose to cristate or
echinate, or with longer folds that may be compressed
and wing-like; x=41.
A subcosmopolitan genus of an estimated
180-230 species, rare or absent in tropical lowlands,
best represented in the mountains of tropical and
warm-temperate areas, e. g., in east-central to southeast Asia (21 species in Nepal according to Nakaike
1982), the Greater Antilles, and the Andes; few species
in continental Africa; rather well represented in temperate areas, especially south-temperate. Many species
are superficially similar, and in many parts of the area
the species of this genus are taxonomically imperfectly
known. This is enhanced by the fact that some widespread and common species are allotetraploids (see,
e. g., Kurata 1964; Vida and Reichstein 1975;
D. Wagner 1979), sterile Ft-hybrids also being fairly
frequent and obscuring species limits.
Most species grow terrestrially or epilithically in forests and thickets, at higher elevation and higher latitudes also in open vegetation; some prefer rock crevices.
The species with simply pinnate lamina, non-aristate
margin, multiseriate sori, and, in some cases, anastomosing veins, are usually segregated as Cyrtomium, or
the neotropical species as yet another genus, Phanerophlebia. These characters are neither sufficiently correlated nor constant for drawing generic lines. Cyrtomium and Phanerophlebia are moreover probably independent derivatives from Polystichum proper, en-
hancing the artificiality of a genus or genera segregated from the core of species.
Polystichum is rather similar, and apparently closely
related, to Dryopteris. but the first credible hybrid was
only very recently reported by W. H. Wagner (in Barrington 1985b). The two genera are, however, separated with ease, and the perispore structure supports
the traditional borderline drawn between them (Tryon
and Tryon 1982). See also under Arachniodes.
A natural subdivision of the genus has so far only
been given on a local basis, e. g., by Tagawa (1940) for
East Asia.
Fig.51A-E. Dryopteridaceae. A,B Polystichum (Cyrtomium)
macrophyllum. A Leaf ( x 0.5). B Part of pinna with venation
and sori (x 2.5). C Polystichum setiferum. fertile pinnule
(x 5.5). D,E Polystichum presconianum. D Habit (x 0.5).
E Fertile pinna (x 4) (A,B from Kurata and Nakaike 1979;
C from Hyde et aL 1969; D,E Ching et aL 1983)
Dryopteridaceae . Dryopteridoideae
116
D
B
Fig. 52A-F. Dryopteridaceae. A,B Didymochlaena IrnncalUla. A Part of lamina (x 0.3). B Fertile pinnule with sori
(x 2.5). C, D Cye/odium inerme. C Apex oflamina (x 0.5).
D Fertile pinna (x 0.5). E,F eye/odium meniscioides. E Sterile pinna (x 0.5). F Fertile pinna (x 0.5) (A,B from Holttum
1955; C-F from A. R. Smith 1986)
CHARACTERS OF RARE OCCURRENCE. Sori borne on the
adaxial (!) surface of the pinnules in a form of
P. anomalum (Hooker & Am.) J. Smith (Ceylon; see
Sledge 1973 a). Exindusiate sori covered by reflexed
leaf lobes in P. ("Plecosorus',) speciosissimum (Kunze)
Tryon & Tryon (Central America). Exindusiate, eventually confluent sori with slightly reflexed leaf margin
over them in P. ("Sorolepidium'') duthiei (Hope) C. Chr.
(Asia). Bipinnate but essentially tripartite leaves in
p. hancock;; (Hance) Diels and P. tripteron (Kunze)
C. Presl (E. Asia). Neither aristate nor dentate pinnule
margins in P. copelandii Christ (Luzon), P. harrisii
Maxon (Jamaica), and the species included in Cyrtomidictyum by Ching (1957). A tripinnate+ pinnatifid
lamina with linear ultimate segments in P. (''Acropelta'') carvifolium Diels (China). Lamina simply pinnate
with sometimes protracted-rooting apex and sori confined to the apical part in the West Indian P. rhizophyllum (Swartz) C. Presl. A bipinnate, linear lamina with
very short pinnae and globe-like, strongly revolute pinnules nearly hidden by the enormous mass of scales in
the almost Jamesonia-Iike New Guinean P. ("Papuapteris'') lineare (c. Chr.) Copel. Somewhat anastomosing veins in a few species like P. ("Cyrtomidictyum '')
lepidocaulon (Hooker) J. Smith (E. Asia).
11. Didymoch/aena Desvaux
Fig. 52 A, B
Didymochlaena Desvaux, Ber\. Mag. Naturf. Fr. 5: 303 (1811).
Large terrestrial ferns with massive, erect, dictyostelic
stem bearing long, narrow and small, hair-like scales
and ± persistent petiole bases. Leaves in a rosette; petiole well-developed, stramineous, at least the base ±
persistently scaly, adaxially grooved. Lamina bipinnate, anadromous or isodromous; neither upper nor
lower pinnae reduced, the apical pinna conform. Rachis like the petiole, bearing very narrow scales as do
the secondary rachises, adaxially grooved, the grooves
of the secondary rachises evanescing at base; adaxial
side of secondary rachises with spine-like processes on
the lateral ridges, especially near the pinnule bases.
Pinnules of a rhombic, tongue-shaped type, ± dimidiate, subsessile, at least the basal ones articulated at
base; main vein near the posterior margin or diagonal,
weakly developed, not grooved. Lateral veins free,
forked, with clavate ends behind the subentire to crenate margin. Upper pinnules abruptly reduced and
confluent. Sori terminal on a - usually somewhat abbreviated - vein, supramedial, not rarely slightly embossed; indusium long and narrow, often quite elongate but essentially horseshoe-shaped, attached along
a line; receptacle elongate, bearing sporangia on both
sides and around its distal extremity. Spores ellipoidal
to somewhat globose, with prominent, inflated folds or
tubercles, usually echinate; n =41.
A single, pantropic species, D. truncalUla (Swartz)
J. Smith, or possibly a second species in Madagascar;
in (often montane) forests and ravines; south to northern Argentina and South Africa, east to Fiji; not in
Australia.
Dldymochlaena • Stigmatopteris
117
Fig. S3A-C. Dryopteridaceae. Stigmatopteris ichtiosma.
A Fertile lamina (x 0.2). B Part of rachis and pinna bases of
fertile lamina (x 2.5). C Costal scale ( x 8) (Original Moran)
12. Stigmatopteris C. Chr.
Fig. 53
Stigmatopteris C. Chr., Bot Tidsskr.29: 292 (1909; 1913,
1919).
Dryopteris subgen. Stigmatopteris (c. Chr.) C. Chr., Kongel.
Dansk. Vid.-Selsk. Skr. VII.10 (2): 73 (1913).
Medium-sized terrestrial ferns; stem short-creeping to
suberect, the apex bearing fuscous, narrowly triangular, acuminate scales. Petioles rather close to aggregate, often persistently scaly, stramineous to pale
brown, adaxially sulcate, with 4 to many vascular bundles. Lamina pinnate to pinnate + pinnatifid (or rarely
bipinnate + pinnatifid), of an oblong-deltoid shape,
pinnatifid apically, herbaceous to chartaceous, glabrous, pellucid-punctate by the presence of internal
glands; dissection preponderantly catadromous. Pinnae sessile or, at least the upper ones, adnate and de-
current by a wing, apically long-attenuate and serrate.
Rachis and costae adaxially grooved, the grooves ±
continuous, glabrous or bearing minute cylindrical
hairs, these sometimes present also elsewhere; major
axes abaxially bearing scattered thin, flaccid, ovate to
lanceolate, tan to dark brown scales having glandular
marginal hairs. Venation largely catadromous, at least
in the pinna bases. Veins simple, free or somewhat irregularly anastomosing in pairs and producing a free
excurrent veinlet; veins not reaching the margin, ending in a conspicuous clavate tip. Sori dorsal, subbasal
to subapical, orbicular to oblong, exindusiate. Spores
ellipsoidal to somewhat globose, with prominent,
sometimes long folds, sometimes perforate; x=41.
Between 20 and 25 neotropical species, from southern Mexico and the Antilles to Bolivia and SE Brazil;
mostly in rainforest at lower and middle elevation.
118
Dryopteridaeeae . Dryopteridoideae
Two Antillean species bear gemmae in the axils of distal pinnae.
Formerly included in Dryopteris or Cyclodium. or
placed near these genera, but now regarded as less
closely related; alliance within the tribe somewhat uncertain.
13. Cydodium C. Presl
Fig.52C-F
eye/odium c. Presl, Tent. Pteridogr.: 85 (1836); A. R Smith
(1986).
Dryopteris subgen. Stigmatopteris (c. Chr.) C. Chr. "group"
Peltochlaena Fee ex C. Chr., Kongel. Dansk. Vid.-Selsk.
Skr. VI 1.1 0: 74 (1913).
Stigmatopteris auett., p. p.
Terrestrial or hemi-epiphytic ferns of medium size
with short- to rather long-creeping stem bearing fuscous, very narrowly lanceolate, subentire to denticulate, deciduous scales. Leaves close to remote; petiole
well-developed, stramineous, deciduously scaly at
base, adaxially sulcate. Lamina of an oblong to deltoid
shape, chartaceous to coriaceous, not pellucid-punctate, glabrous to the unarmed eye, often simply pinnate but up to bipinnate + pinnatifid, dissection and
venation anadromous; upper pinnae, etc. gradually reduced and confluent, or a conform odd pinna present.
Rachis(es) and costae adaxially sulcate, with continuous grooves, in the grooves very short (0.05-0.3 mm),
blunt hairs present (sometimes also abaxiaIly). Fertile
leaves sometimes difform, then longer petiolate and
with narrower laminal parts. Pinna margin entire or
sinuate to deeply pinnatifid, somewhat scierotic, sometimes auricled at the anterior base; lower pinnae shortpetiolulate. Costa percurrent; veins free, or connivent
below sinus, or (sometimes except the basal) anastomosing in pairs to form one or a few series of elongate
areoles, an excurrent vein then present, this free or
joining the next outer vein; veins ending at or near the
margin, their ends not enlarged. Sori dorsal, in one or
mostly a few irregular series between costa and margin, or flanking the costules; indusium peltate, or in
two species round-reniform, often deciduous, glabrous
or glandular-hairy. Spores ellipsoidal, with prominent,
perforate, wing-like folds, more or less echinate;
x-41.
Ten species, from Panama and Trinidad to Bolivia,
NE Argentina, and southern Brazil; greatest concentration in the Guianas. Terrestrial and on rocks, or hemi-epiphytic, in lowland and lower montane forests, to
1500 m, sometimes in swamps; one species a rheophyte.
14. Olfersia Raddi
Fig. 54
Olfersia Raddi, Opusc. Sci. 3: 283 (1819); Moran (1987 a).
Polybotrya auctt., p. p. min., excl. type.
Fig. 54A-C. Dryopteridaceae. O/fersia cervina. A Sterile leaf
(x 0.27). B Fertile leaf (id.). C Sterile lateral pinna (x 0.6)
(Original Moran)
Medium-sized, usually terrestrial, occasionally scandent ferns; stem densely scaly, bearing linear, goldenbrown scales. Leaves close; petiole well-developed,
stramineous, glabrescent, adaxially sulcate. Lamina
strongly dimorphic (with intermediate forms occasionally present); sterile lamina simply pinnate with conform terminal pinna, anadromous, glabrous, subcoriaceous; pinnae ovate-Ianceolate to lanceolate, alternate,
4-12 to a side, short-petiolulate, entire, with cartilaginous margin, acute-acuminate, unequally narrowed at
base, the basiscopic side excavate; veins forking at or
near the base, ciose, parallel, free but joined by a submarginal commissure (this sometimes difficult to observe in dried material). Fertile leaves bipinnate (rarely
unipinnate), produced on terrestrial and on scan dent
stems, more erect than the sterile leaves and with longer petiole, soon withering after the shedding of the
spores; pinnae linear, occasionally entire, but usually
with pectinately arranged pinnules which are up to
1 x 113 em; sori soon confluent and evenly covering
Cyclodium • O(fersia. Polybotrya. Teetaria
the segments beneath, without indusia; sporangial
stalks bearing gland-tipped hairs. Spores about as in
the next genus; n ~ 41.
A single species, O. cervina (L.) Kunze, neotropical,
from southern Mexico and the West Indies south to
Bolivia, Paraguay, and SE Brazil; in forests, terrestrial,
epilithic, or on tree trunks, at lower and middle elevation.
119
species and endemics is in the Andes from Colombia
to Bolivia; a secondary centre is in SE Brazil with 5
endemic species. Most species inhabit primary forests
at lower and middle elevation. Two of the three species of section Sorbifolia are terrestrial.
16. Tectaria Cavanilles
Figs. 56, 59
Teetaria Cavanilles, Anal. His!. Nat. 1: 115 (1799); Tardieu-
Fig. 55
Blot (1951); Morton (1966); Iwatsuki (1968); Sledge
(1972); Holttum (1981, 1988).
Poiybotrya Willdenow, Spec. Plant. 5: 99 (1810); Moran
Aspidium Swartz (1802), nom. superfl., and of most later au-
15. Polybotrya Willdenow
(1987b, c).
Medium-sized to large, scandent (rarely terrestrial)
ferns; stem long-scandent, comparatively stout, ± angular-ridged, attached by numerous ventral roots, at
least the apical portion densely scaly, with 4-12 meristeles in transection, these arranged in a circle and
each surrounded by a dark sclerenchymatous sheath;
scales large, very long and narrow, yellowish-brown to
fuscous, their structure often of diagnostic value.
Leaves remote, strongly dimorphic; petiole stramineous, adaxially sulcate, well-developed, glabrescent or
the base with persistent scales. Sterile lamina 2- to
4-pinnate (subgroup Polybotrya) or simply pinnate
(subgroups Soromanes and Sorbifolia), chartaceous to
coriaceous, anadromous or catadromous, depending
on the species; axes especiaUy abaxiaIly loosely to
densely hairy, the hairs in some species extending to
the larger veins, narrow scales sometimes present on
the axes in addition; all axes incl. costae adaxially sulcate, the grooves continuous, shaUower at junctions or
truncate; no articulations at insertions of leaf divisions. Ultimate divisions very variable in size and
shape, depending on the species and the place in the
lamina; edge entire or more often serrate-lobulate to
deeply pinnatifid; upper divisions graduaUy reduced
and confluent, no conform terminal divisions present.
Veins subpinnately forked to pinnate, the pattern
anadromous or more often catadromous; veins free or
(in section Soromanes) anastomosing without free included veinlets, the excurrent veins again joined, forming very elongate areoles; vein ends not close to the
margin, hardly enlarged. Fertile leaves strongly difform, "skeletonized" with suppression of most of the
laminal parts, in the scandent species only produced
on climbing stems, not terrestrially; divisions very narrow, short to long, usually entire; sporangia in discrete, round, often eventually confluent sori without
indusia or acrostichoid, evenly covering the fertile divisions. Sporangial stalks often bearing a glandular
hair. Spores ellipsoidal, bearing prominent folds with
dense echinate elements, or echinate; x~41.
Confined to the neotropics, with 35 species, occurring from Mexico and the West Indies south to Paraguay and southernmost Brazil. The greatest number of
thors, at least in part.
Amphiblestra C. Presl (1836).
Stenosemia C. Presl (1836).
Fadyenia Hooker in Hooker and Bauer (1840).
Dietyoxiphium Hooker (1840).
Camptodium Fee (1852).
Cionidium Moore (1857).
Luers'senia Kuhn (1882).
Hemigramma Christ (1907).
Tectaridium Copeland (1926).
Querc!/ilix Copeland (1928).
PleurodelTis Maxon (1934).
Ctenitopsis Ching (1938), p. p. maL
? Pseudoteetaria Tardieu-Blot (1955).
Small'to rather large, terrestrial or epilithic ferns with
rather long-creeping to short, ascending or erect stem
bearing long and narrow, acuminate, (often dark)
brown, non-clathrate scales. Petioles remote to clustered, weIl-developed, stramineous to dark and then
often polished, often scaly at least near the base, with
several vascular bundles in a semi-circle, adaxially
grooved, in large species often with additional
grooves, occasionaUy winged, sometimes hairy, especially in the adaxial groove, the hairs short, reddish,
articulate, or also with thinner, paler hairs. Lamina
small to more often medium-sized to large, pinnatifid
to bipinnate + pinnatifid, less often lobed or entire and
simple, when simply pinnate usually with a basiscopic
lobe on the basal pinnae, herbaceous to subcoriaceous; basal pinnae segments very often basitonicaUy
produced; rachis like the petiole, rarely dark, usuaUy
clothed with reddish, articulate hairs in the adaxial
groove; upper pinnae often reduced and confluent into a pinnatifid leaf apex, or this trilobed, less often
conform. Pinnae sessile or the lower short- to longstalked; or the pinnae, especially the upper ones, adnate and decurrent (or also surcurrent). Secondary rachises (if any) adaxially flat to convex, bearing ctenitoid hairs, or, if shortly grooved at base, the groove not
joining that of the primary rachis. Architecture catadromous, only the (usually single) basal pair anadromous; in upper pinnae/pinnules the basal posterior
primary vein may spring from the rachis/rachidule
rather than from the costalcostule. Scales on axes of
higher order few or none. Costae occasionally adaxialIy bearing proliferous buds; rarely the leaf apex root-
120
Dryopteridaceae • Dryopteridoideae
Fig. 55 A-H. Dryopteridaceae. A Polybotrya speciosa, habit
with sterile and fertile leaf ( x 0.25). B Polybotrya osmundacea. sterile pinnule (x 0.6). C Polybotrya speciosa, sterile pinnules (x 0.6). D Polybotrya pubens, sterile pinnule (x 0.6).
E Polybotrya jractiserialis, sterile pinnule (x 0.6). F Polybot-
rya serratifolia, sterile pinnule (x 0.6). G Polybotrya osmundacea, fertile segments from abaxial side (x 3). H Polybotrya
sp., transection of stem, meristeles surrounded by sclerenchyrna sheath (SH) (AP aerophore; LTieaf trace; RTroot trace)
( x 3) (Original Moran)
Tectaria
121
B
E
F
ing and proJiferous. Veins sparingly (along costae and
costules) to copiously anastomosing, usually with
some to many free, included veinlets which point in all
directions or not; rarely all veins free ("Ctenitopsis'');
venation catadromous, basal posterior veinlets springing from costa, not from costule; costal areoles sometimes subdivided. Veins glabrous to pubescent; costae!costules adaxial\y usually bearing ctenitoid hairs,
at least near their bases. Margin often ciliate. Leaf tissue pubescent or glabrous between the veins. Sinuses
sometimes sclerotic, not protracted into a tooth (as
they are in Pleocnemia), often with a patch of ctenitoid
hairs on the adaxial side. At least some marginal veinlets free. Fertile leaves conform or difform, then usually weakly to strongly contracted. Sori of highly varying
shape and position but most often dorsal or terminal
Fig. 56A-F. Dryopteridaceae. Teetaria (Quercifilix) zeilaniea.
A Habit (x 1). B Part of abaxial surface of sterile lamina
showing indument and venation (x 2.5). C Detail of venation of sterile lamina (x 5). D Young fertile lamina (x 3).
E Hairs of lamina (c. x 5). F Stem scale (x 24) (A,B,E from
Flora Taiwan 1975; C,D,F from Ching 1935)
or not rarely compital, round, elongate, or less often
linear, usually biseriate and!or scattered between adjacent costules, sometimes following the margin, very
rarely on tiny lobes terminating veins on the margin or
beyond ("stalked") ("Cionidium", Fig.59A-C); when
round to shortly elongate usually with a peltate, hippocrepiform, or pseudopeltate indusium, sometimes
exindusiate or the indusium early caducous; very rare-
122
Dryopteridaceae • Dryopteridoideae
Iy the sari linear and following the margin, then with a
conform indusium opening outward as in Lindsaea
("Dictyoxiphium'') or exindusiate and±covered by the
reflexed margin (''Amphiblestra ''); very rarely the sporangia following the veins, not assembled in sari, or
even eventually covering the entire fertile surface
("Quercifilix''); but these extremes usually connected
by intermediates and/or hybrids. Spores subellipsoid,
bearing wing-like folds, these often cristate or echinate; areas between the folds usually reticulate-echinate or spinulose; n =40 or multiples.
A genus of perhaps 150 species, widely distributed
in the tropics of the whole world, extending into
south-temperate and, very locally, also into north-temperate areas (Florida, E. Asia); very often terrestrial in
forests, not rarely cremnophytic on road banks, also
often on rocks and in thickets, rarely in open places.
A subdivision into two sections was proposed by
Holttum (1987): section Tectaria with amply anastomosing veins and branched free included veinlets; and
section Sagenia (c. Prest) Holttum with either free
veins, or veins forming costal areoles without included
veinlets and few included veinlets in the other areoles.
Several segregates have been proposed (see synonymy) on the basis of deviating venation, soral shape, absence of indusium, sporangia not being assembled in
sori, extreme dimorphism, etc., but none of these "genera" is distinct enough for recognition in a system
whose genera are based on a combination of many
characters, including those of indument, anatomy, architecture, etc.
Pseudotectaria is treated as distinct by Holttum
(1987); it comprises a small number of Madagascan
species.
The fern originally described as Gymnopteris hollrungii Kuhn in Schumann and Hollrung, later placed
in Hemigramma, was recently made the type (and only) species of the new genus Chlamydogramme Holttum (1987). It is distinguished from Teetaria chiefly by
the linear sori and should probably be included in Tectaria sens.lat., as defined here.
17. Heterogonium C. Presl
Heterogonium C. Presl, Epimel. Bot.: 142 (1851); Holttum
(1949b,1975).
Tectaria auctt., p. p. min.
Small to medium-sized, terrestrial ferns; stem erect,
bearing rigid, dark, narrow scales. Petioles close, welldeveloped, medium to usually blackish brown, deciduously scaly, bearing ctenitoid hairs, to glabrescent.
Lamina deltoid to oblong, herbaceous to chartaceous,
pinnate + pinnatifid, catadromous except for the anadromous or isodromous basal pinnae; pinnae not numerous, sessile or the basal short-stalked, the upper
adnate-decurrent, the uppermost gradually confluent;
rachis adaxially sulcate, bearing ctenitoid hairs, these
also present on the laminal parts or confined to small
groups of hairs near the sinuses; costae abaxially
sometimes strigose. Pinnae lobed to deeply pinnatifid,
the lower ones often broader at the posterior side but
always broadest above the base, the basal basiscopic
segment(s) reduced. Veins free or anastomosing along
costae and costules, or fully reticulate, without or with
few free included veinlets. Sporangia in round to elongate sari, or following the veins, or acrostichoid in arrangement; sari if short dorsal or terminal, indusiate
or not, the indusium reniform. Fertile leaves often with
somewhat contracted lamina, this strongly contracted
in case of acrostichoid sporangia and with longer petiole. Spores ellipsoidal, bearing prominent connected
folds with echinate borders, echinate also in the areolae; n=40.
About 20 species, from southern China and Burma
to western Melanesia, and one species in Mauritius.
Terrestrial in forests and thickets, or epilithic, at lower
elevation to c. 1200 m. Some species confined to limestone.
Very close to Tectaria and perhaps not generically
distinct, especially as there is some evidence for hybridization.
18. Pieocnemia C. Presl
Fig. 57
Pleocnemia C. Presl, Tent. Pteridogr.: 182 (1836); Holttum
(1951 a, 1974).
Arcypteris Underwood (1903); Holttum (1951 b).
Terrestrial or epilithic ferns with ascending or erect,
sometimes trunk-like stem bearing narrow brown
scales with short, marginal projections from two cells.
Leaves close, often large; petiole well-developed,
adaxially flattened or sulcate, with one U-shaped series of vascular bundles with larger adaxial ones, in
larger species with additional bundles in the centre
and on the adaxial side; base bearing narrow, thin,
fuscous scales. Lamina pinnate + pinnatifid to bipinnate + pinnatifid or subtripinnate, of a deltoid type;
basal pinnae the largest, basitonically enlarged; leaf
pattern catadromous except (always?) in the basal
pair. Primary rachis adaxially grooved, often with additional lateral ridges and grooves; pinnae inserted on
the ridges bordering the central groove, their grooves
not continuous with that of the primary rachis. All
axes with ctenitoid hairs, or glabrescent with age. Ultimate free divisions lobed to pinnatifid; upper ones reduced, confluent. Bottom of sinus between lobes with
a distinct (or very small) tooth served by a vein which
in life points out of the plane of the pinna. Margin of
lobes entire to crenate. Veins anastomosing to form at
least a series of narrow costal areoles; costular areoles
often also present; outer veins free or casually to regularly anastomosing, without or with a few simple, free
Heterogonium . Pieocnemia . Hypoderris. Psomiocarpa
123
Fig. 57 A, B. Dryopteridaceae. Pleocnemia conjugata. A Part
of rachidule with sterile pinnules (x 0.6). B Part of rachidule
with pinnule base (x 4) (Holllum 1955)
included veinlets; veins beneath often with scattered,
few-celled glandular hairs. Sori on conform or slightly
contracted segments, dorsal, compital, or terminal,
orbicular, with a reniform indusium or exindusiate;
sporangial stalk often with a short glandular hair.
Spores ellipsoidal to somewhat spheroidal, with prominent folds which may be wing-like or interrupted, or
sometimes prominently echinate; n=41.
Nineteen species (Holttum 1974), from NE India,
SE China, and Taiwan throughout Malesia to Samoa.
In forests and thickets, some species epilithic.
19. Hypoderris R. Brown
-.
"
B
.
.
}
Fig. 58
Hypoderris R. Brown in IWallich, Icon. PI. As. Rar. 1: 16
(1830) or] Hooker and Bauer, Gen. Fil.: pI. 1(1838).
In most respects like a medium-sized Teetaria. Lamina
non-dimorphic, simple or lobed below, the lobes small
or when large often with a small basiscopic lobule;
veins amply anastomosing, with free included veinlets;
sori small, often near the secondary veins, dorsal or
compital, suborbicular to elliptic; indusium inferior,
forming a thin, irregular, lacerate, ciliate ring. Spores
ellipsoidal or somewhat globose, irregularly echinate,
the elements basally perforate.
A single species, H. brownii J. Smith in Hooker and
Bauer, in moist woods and on rocks, at lower to middle elevation; Caribbean region from Hispaniola
south to Trinidad and northern Venezuela.
Very similar to Teetaria (also n = 40); but the inferior indusium is unique (Fig. 58 B).
Fig. 58 A, B. Dryopteridaceae. Hypoderris brownii. A Habit
( x 0.25). B 80rus (x 15) (Diels 1899)
20. Psomiocarpa C. Presl
Psomiocarpa C. Presl, Epimel. Bot.: 161 (1849); Zamora and
Chandra (1977).
Rather small terrestrial ferns with ascending or erect
stem bearing clathrate, non-glandular, marginally
fringed scales. Leaves close; petiole well-developed,
stramineous, adaxially sulcate, sparingly scaly, chiefly
at base, shaggy with articulate hairs, especially adaxially, minutely dark-warty abaxially, with three vascular bundles. Leaves strongly dimorphic; sterile
leaves±spreading; lamina oblong, pinnate+pinnatitid to bipinnate+pinnatitid, catadromous, the basal
pair often basitonically produced; rachis like the pe-
124
Dryopteridaceae . Dryopteridoideae
Fig.S9A-D. Dryopteridaceae. A-C Tee/aria (.. Cionidium'J
moorei. A Fertile pinnule ( x 0.6). B Part of fertile segment
(x 1.5). C Sorus (x 5). D Tee/aria sinuala, fertile segment
(x 1.5) (Brownlie 1969)
tiole; upper pinnae and pinnules reduced, abruptly
confluent, all ascending, (sub)sessile, the basal posterior segments of higher pinnae springing from the rachis; margin dentate to pinnatifid; veins free, pinnate,
the larger bearing ctenitoid hairs; upper surface bearing scattered intestiniform hairs between the veins;
leaf axes other than primary rachis not grooved, pinnule edges running into narrow wings bordering secondary axes. Fertile leaves erect, more long-petiolate,
bi- to tripinnate; laminal parts very much reduced,
consisting of sessile, roundish segments covered with
sporangia; indusium none. Sporangia bearing 2-celled
glandular hairs on the stalk, mixed with long, onecelled, and shorter, articulate hairs. Spores ellipsoidal;
surface with prominent, compact, dentate wings of
compressed folds; n = 40.
A single species, Ps. apiifolia (1. Smith ex Kze.)
C. Presl, widespread in the Philippines. Regarded variously as related to Tectaria or Ctenitis; the chromosome number (n =40) agrees better with the former,
the morphology more with the latter; or perhaps it is
closer to Heterogonium. The intestiniform hairs are peculiar.
21. Triplophyllum Holttum
Triplophyllum Holttum, Kew Bull. 41 : 239 (1986).
Clenitis auctt., p. p. min.
Terrestrial, medium-sized to rather large ferns; stem
slender, short- to rather long-creeping, bearing narrow,
non-clathrate scales. Petioles close to remote, well-developed, light to dark brown, dull, thinly scaly with
narrow, non-clathrate scales and tomentose with short
ctenitoid hairs to glabrescent, adaxially narrowly sulcate; sometimes sparse longer hairs also present. Lamina of a triangular to pentagonal type, the basal pinnae
usually the largest, stalked, subopposite, and basi tonically produced, sometimes the lamina tripartite (i. e.,
each of the basal pinnae about equivalent to the remainder of the lamina); texture herbaceous to subcoriaceous; primary dissection pattern of the basal pinna
pair anadromous, otherwise the entire lamina including the venation catadromous; upper pinnae, pinnules,
etc. gradually reduced and confluent (except in one
species). Lamina at least pinnate+pinnatilobate (at
base), often more compound, to tri- or quadripinnate + (bi)pinnatifid. Rachis in some species bearing
adventitious buds in its upper part, hairy like the petiole, adaxially more distinctly grooved, the grooves of
axes of higher order (if present) evanescing near the
base; secondary rachises and costae occasionally also
bearing adventitious buds. Ultimate free divisions of
an ovate or lanceolate type, rarely auriculate at their
anterior base, asymmetric, mostly very obtuse, usually
adnate, especially posteriorly usuaUy adnate-decurrent, their edges neither thickened nor running into
adaxial rachis ridges. Abaxial surface sometimes
glandular; ctenitoid hairs mostly present on costae
and costules, especially adaxially; intestiniform hairs
sometimes present adaxially on veins; cylindric glands
none. Veins pinnate in the segments, immersed, not
reaching the margin, free or rarely casually to copiously anastomosing without free included veinlets.
Basal veinlet springing from the costule. Sori orbicular,
dorsal or (sub)terminal, with hippocrepiform-reniform, ± persistent, sometimes hairy or glandular indusium, rarely exindusiate and somewhat elongate
[T. varians (Moore) Holttum]. Spores rather like those
of Tectaria, bearing cristate or echinate, wing-like
folds, the surface between the folds ± densely reticulate-echinulate; n =41.
Fifteen species in West and Central Africa and
Madagascar, five in tropical America; there present in
the West Indies and northern South America; terrestrial in rainforest.
Agreeing in dissection pattern and scales with Tectaria, but differing in venation and chromosome number in which it approaches Ctenitis; but the latter diverges in scales and leaf architecture.
Triplophyllum
125
B
Fig.60A-D. Dryopteridaceae. A,B Ctenitis cirrhosa. A Habit (x 0.5). B Part of fertile pinna (x 1). C Ctenitis distam,
scale from abaxial face of costa (x 30). D Ctenitis hairs
(much enlarged) (A,B from Schelpe 1970; C,D from Christensen 1912)
126
22. Ctenitis C. Chr.
Dryopteridaceae • Dryopteridoideae
Fig. 60
Ctenitis (c. Chr.) C. Chr. in Verdoom, Man. Pteridol.: 543
(1938), based on: Dryopteris subgenus Ctenitis C. Chr., BioI.
Arb. til E. Warming: 77 (1911).
Ata/opteris Maxon (1922).
Ataxipteris Holttum, B1umea 30: 10 (1984).
Terrestrial or epilithic, medium-sized to large ferns;
stem ascending to erect or even trunk-like and a few m
tall, bearing pale to dark brown, pointed, clathrate
scales. Petioles close, well-developed, usually stramineous to medium brown, adaxially sulcate, in large species sometimes with additional grooves and ridges,
bearing reddish, articulate hairs at least in tbe adaxial
groove and pale to dark brown, ± distinctly clathrate
scales at least at base, tbere often in a dense mass.
Lamina at least pinnate + pinnatifid, usually more
highly compound, up to tripinnate + (bi)pinnatifid,
herbaceous to subcoriaceous, catadromous (very rarely
anadromous at very base), usually broadest at base,
tbe lowest pinnae not rarely basi tonically enlarged,
rarely tbe lower pinnae reduced and reflexed; rachis
like the petiole, adaxially grooved only in at least bipinnate species, tbe groove not continuous with tbat of
the primary rachis; pinnule!segment-bearing axes!
costae adaxially convex or flattened, rarely grooved at
base; pinnae stalked or in smaller species often sessile.
Upper pinnae, pinnules, etc. gradually reduced and
confluent; apices often obtuse to subacute. Laminal
parts at posterior base not thickened and decurrent
onto the ridges of their axes, these not being laterally
ridged. Axes up to costae often, especially abaxially,
scaly, tbe scales ± distinctly clathrate, occasionally bullate; long, pale, pointed hairs may be present beside
ctenitoid hairs and! or scales. Veins, sometimes also
surfaces, adaxially bearing ctenitoid hairs; veins free,
simple, forked, or subpinnate, catadromous, tbe basal
vein of a segment rarely springing from tbe costa instead of the costule. (Pen)ultimate divisions often oblique, with unequal base, decurrent and wing-connected; margin entire or crenate to pinnatifid. Surface
bearing few to many unicellular, cylindrical glands.
Fertile segments rarely contracted to strongly reduced
("Atalopteris"). Sori dorsal or subterminal (not truly
terminal) on an anterior vein branch, orbicular; indusium reniform to subpeltate, rarely fugacious or wanting, with small glands at its margin. Spores saccate,
spinulose, or echinate, the spines sometimes fused
into ± parallel ridges; n=41.
A taxonomically difficult genus never studied in its
entirety. Species groups for the American representatives were proposed by Christensen (1919) and modified by Tryon and Tryon (1982); a formal subdivision
is still to be given. Delimitation against neighbouring
genera, especially Tectaria, has been somewhat fluctuating; and some groups were recently taken out and
treated as genera: Aenigmopteris, Dryopsis, Triplophyllum. About 150 species in tropical and soutb-temperate areas (not in Australia), here and tbere also in
northern warm-temperate areas (Asia); often terrestrial
in forests, sometimes On rocks, in thickets, or in more
open situations. When Atalopteris is included, tbe
name Clenitis is in need of conservation.
The group of species centred around Ct. vil/osa (L.)
Copel., mainly in tropical America but extending to
Mrica and the Mascarenes, was recently segregated as
a new genus, Megalastrum, by Holttum (1987).
23. Aenigmopteris Holttum
Fig. 61 A-C
Aenigmopteris Holttum, Blumea 30: 3 (1984).
Rather large terrestrial ferns. Stem ascending, apically
bearing narrow, fuscous, non-clathrate scales. Petioles
close or not, well-developed, dark, sparsely scaly and
short-hairy, adaxially grooved. Lamina herbaceous,
narrowly triangular, bipinnate + pinnatifid to tripinnate, with several pairs of pinnae, the lower subopposite, tbe lowermost basitonically produced; dissection
and venation catadromous, only the basal primary division anadromous. Upper pinnae, pinnules, etc. gradually reduced and confluent, with narrow segments.
Proliferous buds sometimes present at pinna bases.
Axes fuscous, with few, small, tbin, non-clatbrate
scales and bearing ctenitoid hairs; thick, articulate, intestiniform hairs present on tbe adaxial surface between the veins, sometimes also abaxially on the veins.
Axes adaxially not grooved, or the pinna rachis slightly grooved. Penultimate segments narrow, acute to
acuminate, decurrent; ultimate segments lanceolate to
tongue-shaped. Veins pinnate, tbe branches simple or
forked in the segments, ending behind tbe margin;
vein of basal posterior segment arising from costule,
not from axis of next lower order. Sori terminal, suborbicular; indusium firm, dark, reniform to asymmetrically horseshoe-shaped. Spores oblong, the perispore
forming thin, slightly areolate ridges with spinulose
margins; n = ?
Five described species, terrestrial in forest at middle
elevation; in tbe Philippines, Borneo, and New Guinea.
In aspect ratber like Ctenitis, differing, e. g., in the
intestiniform hairs on tbe adaxial side of the lamina
and in tbe non-clathrate scales; and from free-veined
Tectaria species in tbe divergent venation pattern. The
form "Aenigmatopteris" would be linguistically more
correct.
24. Dryopsis Holttum and Edwards
Dryopsis Holttum and Edwards, Kew Bull. 41: 179 (1986).
Ctenitis auctt., p. p. min.
Rather small to medium-sized ferns; stem short, ascending or erect, sometimes massive, densely scaly at
Ctenitis· Aenigmopteris' Dryopsis
apex; scales non-clathrate, entire, ovate, tan, and flaccid to very narrowly triangular, long-acuminate, fuscous, and glossy. Leaves aggregate; petiole well-developed, longer or shorter than the lamina, stramineous
to fuscous, adaxially sulcate, scaly, especially at the
base, the scales tan and flaccid to blackish-brown and
rigid, weakly clathrate in the lower part or not at all so,
intermingled with multicellular, basally thickened
hairs; hair/scale intermediates may be present. Lamina deltoid to oblong, pinnate+deeply pinnatifid to
sUbtripinnate, tertiary divisions, if present, narrowly
wing-connected; basal pinnae not enlarged, sometimes
reduced, but the posterior pinnules often larger than
the anterior; dissection weakly to pronouncedly catadromous, or the basal pinnae anadromous; pinnae
sessile to short-stalked, the lower very nearly opposite;
upper pinnae, pinnules, etc. gradually reduced and
confluent; texture herbaceous to subcoriaceous. Rachis at least abaxially scaly, often densely and conspicuously so, the scales pale and flaccid to dark and rigid; similar, smaller scales usually present on secondary
axes, often also on costae; in certain species some or
all scales basally bullate. Axes adaxially shallowly to
pronouncedly sulcate, the groove without ctenitoid
hairs but at its edge bearing multicellular, basally
broadened and short-celled hairs; groove of pinna axis
starting above the base, therefore not continuous with
that of the primary rachis. (Pen)ultimate divisions often obtuse, ± parallel-sided, entire to pinnatifid. Stiff
127
Fig_ 61 A-G. Dryopteridaceae. A, B Aenigmopteris elegans.
A Pinnule (x 3.5). B Pinnule with sori (x 7.5). C Aenigmoptelis katoi, adaxial side of pinnule with hairs (x 18).
D,E Lastreopsis munita. D Adaxial side of junction of secondary and tertiary rachises, showing hairy grooves and continuous lateral ridges (x 7). E Fertile tertiary segment showing catadromous venation (x 5). F, G Dryopolystichum phaeostigma. F Part of lamina with two pinnae (x 0.5). G Fertile
segment (x 5) (A-C from Holttum 1984; D, E from Tindale
1%1; F, G from Copeland 1947)
hairs like those of the rachis sometimes also present on
the veins; slender, appressed hairs may also be present
on laminal parts. Veins catadromously or anadromously branched, the branches simple or less often
forked, free, ending well behind the margin. Sori orbicular, dorsal or subterminal on the veins, sometimes
borne only in the apical parts of the segments; indusium reniform, the sinus often so narrow as to give the
appearance of a peltate indusium, flaccid, thin, in a
few species small or absent. Spores ellipsoidal to
somewhat spheroidal, usually echinate, or with coarse,
inflated tubercles; x=41.
About 26 species (one probably a hybrid), in the
mountains of east-central to east, south, and SE Asia,
from Nepal to Ceylon, Taiwan, and New Guinea; the
greatest concentration of species in southern China.
Terrestrial in montane forests.
128
Dryopteridaceae • Dryopteridoideae
In aspect often much like Nothoperanema, but easily
distinguished by the adaxial groove of the secondary
rachis that does not extend to its base.
25. Lostreopsis Ching
Fig. 61 D, E
Lastreopsis Ching, Bull. Fan Mem. Inst. BioI. BQt. 8: 157
(1938); Tindale (1957, 1961 b).
Parapoiystichum (Keyserting) Ching (1940).
Medium-sized terrestrial or less often epilithic ferns, or
low epiphytes; stem short· to long-creeping or rarely
erect, dictyostelic, bearing thin, narrow, brown to
blackish, entire to denticulate, non-clathrate scales.
Petioles remote to close, well·developed, with several
vascular bundles in a semi·circle, adaxially sulcate or
with two additional grooves, stramineous to medium
brown with darker base, there, or also elsewhere, ±
persistently scaly. Lamina herbaceous to chartaceous,
pentagonal, much divided, at least bipinnate, up to
5-pinnate + pinnatifid, the basal pinnae basi tonically
enlarged; lower pinnae usually subopposite; dissec·
tion often anadromous at base, catadromous higher
up, the divisions of higher order also catadromous;
less often quite catadromous, rarely quite anadro·
mous; rachis occasionally bearing proliferous buds,
adaxially grooved, the groove continuous onto the
axes of higher order, bearing short ctenitoid hairs or
longer, thinner hairs, occasionally also scaly; occasion·
ally the entire leaf bearing long, slender hairs up to the
petiole; ctenitoid hairs sometimes covering the axes
but densest in the groove, the latter rarely glabrescent.
Glandular hairs may be present in addition. Grooves
of axes at least of higher order raised in the middle
and hairy. Upper divisions gradually of simpler structure and confluent. Ultimate free divisions crenate or
lobed to deeply (bi)pinnatifid, rarely spinulose-dentate, unequal-sided, narrowed at base, their posterior
basal edge thickened and continuous with the lateral
rachis ridge; veins adaxially convex, not grooved.
Lowest segments sometimes inserted on the rachis of
lower order. Veins free, pinnate (+ forked), variously
catadromous or anadromous, ending behind the mar·
gin. Sori dorsal or terminal on a vein or its anterior
branch, round or rarely elliptic; indusium reniform to
peltate or wanting. Sporangia long-stalked, the stalk
mostly gland·bearing; bow of 13-16 indurated cells.
Spores ellipsoidal, usually rugulose-saccate, subcristate, or bearing long folds with echinate surface between them; n=41.
C. 40 species, pantropic and south-temperate, the
greatest concentration in Australia; to New Zealand
and Polynesia; only 6 species in America. Mostly terrestrial in forests, in the tropics often montane; a few
species often grow as epiphytes.
26. Dryopolystichum Copeland
Fig. 61 F, G
Dryopo[ystichum Copeland, Gen. Fil.: 125 (1947).
Rather large terrestrial ferns; stem short, erect, bearing
liguliform, tapering, to linear, fuscous, firm, entire
scales to 2 cm long. Leaves approximate, forming a
crown; petiole about as long as the lamina, adaxially
sulcate, with 6 vascular bundles near the base, bearing
scales similar to those of the stem, especially at the
base, these grading into fibrils and ± lost with age.
Lamina deltoid-lanceolate, pinnate + pinnatisect, or
the basal pinnae fully pinnate at base, somewhat
abruptly reduced to an acuminate apex, catadromous
above the basal pinnae; pinnae short·stalked at the
base of the lamina, or subsessile, patent to ascending,
not contiguous, acuminate, abaxially with scattered fibrils, the costa adaxially grooved and prominently
2-ridged, the groove not continuous with that of the rachis, adaxially with hair· like processes, especially at
the junction with the rachis; upper pinnae sessileadnate, reduced and confluent; ultimate segments basally entire, apically remotely toothed, asymmetrically
triangular-lanceolate, ascending, ± falcate, weakly decurrent; venation catadromous, the veins mostly once
forked. Sori borne on a vein branch or at a fork, about
medial, orbicular, covered by a large, peltate, orbicular
indusium, this minutely glandular-ciliate, usually persistent. Spores ellipsoidal, with few prominent, long
wings which echinulate edges, surface sparsely echinu·
late between them; n = ?
A single species, D.phaeostigma (Cesati) Cope!., terrestrial in forest and by watercourses at lower elevation; in New Guinea, the Bismarck Archipelago, and
the Solomon Is.
27. Pteridrys C. Christensen and Ching
Figs. 62 A-D,63
Pteridrys C. Christensen & Ching, Bull. Fan Mem. Inst. Bi01.5: 125 (1935).
Terrestrial or epilithic, medium·sized to large ferns
with short, ascending to suberect stem bearing narrow,
subentire, basally attached, cordate scales and some
pluricellular hairs. Petioles close, well-developed,
adaxially deeply grooved, the groove with raised centre. Lamina pinnate + pinnatifid, herbaceous; rachis
adaxially grooved, the centre of the groove raised,
minutely hairy to glabrescent; dissection oflamina catadromous; or a few basal pinnae anadromous, the
lamina upward gradually catadromous. Pinnae subsessile to stalked, lanceolate, pinnatifid, with ± caudate
apex, the base or stalk marginate, the margins passing
into the segment edges; costae adaxially flat. Lower
pinnae sometimes basitonically produced; upper pinnae reduced or not, leaf apex subconform to pinnati-
Lastreopsis· Dryopolystichum • Pteridrys· Cyclopeltis
129
Fig. 62A-F. Dryopteridaceae. A-D Pteridrys syrmatica.
A Basal pinna (x 1). B 'l\vo fertile segments (x 4). C Sorus
(x 30).0 Stem scale (0. x 7). E,F Cyclopeltis crenata. E Part
of fertile pinna (x 3). F Pinna base with articulation (x 10)
(A-D from Christensen and Ching 1934; E,F from Holttum
1955)
lid; margin of pinnae serrate, especially towards apex;
surfaces glabrous except sometimes the costae. Veins
free, forked; venation strongly catadromous, the basal
basiscopic veinlet often springing from the costa rather
than from the costule; basal veinlets ± connivent below the sinus. Sinus with a lobe-like tooth served by a
veinlet; posterior veinlet branches running to a tooth
of the margin and almost reaching the edge. Sori on
the shorter, anterior veinlet branch, terminal or subterminal, in an approximately medial series in the
lobes, orbicular, with reniform, sometimes asymmetric
indusium. Spores eJlipsoidal to subspherical, bearing
prominent, compressed, wing-like folds with echinate
borders, these sometimes interrupted, and perforate;
Fig. 63. Dryopteridaceae. Pteridrys cnemidaria, part of lamina from adaxial side showing sinus bottom teeth; Thiwan.
Photo Ch.-M. Kuo
n=41.
Eight species, in forest, terrestrial or epilithic, ranging from Sikkim, Ceylon, and SE China to Taiwan and
New Guinea.
28. eye/ope/tis J. Smith
Fig.62E, F
Cyclopeltis J. Smith, Camp. Curtis Bot. Mag. 72: 36 (1846).
Terrestrial or epilithic ferns with thick, ascending or
erect stem bearing long and narrow, fuscous scales on
short projections, with marginal teeth consisting of
parts of one cell. Petioles close, well-developed, adaxi-
ally grooved and with some additional lateral grooves,
minutely hairy with few-celled hairs and bearing some
dark, very narrow scales. Lamina simply pinnate, with
subconform terminal pinna; rachis like the petiole,
abaxially minutely hairy. Pinnae numerous, long and
narrow, (sub)entire, inserted on lateral ridges of the
adaxial side of the rachis, sessile, articulate at base,
eventually deciduous; posterior base often subauriculate and overlying the rachis; veins isodromous or
130
Dryopteridaceae • Dryopteridoideae • Athyrioideae
weakly catadromous; basal pinnae reduced or not;
upper pinnae abruptly and slightly reduced. Veins
free, subdichotomously pinnate, the first branch acroscopic, ending at various distances from the margin;
costa adaxially flattened, abaxially ± minutely hairy
and slightly scaly, glabrescent. Sori dorsal on the veins
but often near their apices, in one or a few series parallel to the costa, orbicular, with peltate, sometimes deciduous indusium. Spores subglobose, with prominent,
usually echinate folds, some shorter, inflated tubercles
also present; n=41.
A small genus of c. 5 species in SE Asia from southern China and Burma to the western Pacific; not in
Australia; in shaded places, not rarely on limestone. A
single American species, C. semicordata (Swartz)
J. Smith, widespread from Mexico to Peru and northern Brazil. Species delimitation insufficiently known,
and alliance not quite clear.
29. Coveniella Tindale
Coveniella Tindale, Gard. Bull. Singapore 39: 169 (1986).
Medium-sized terrestrial fern; stem long-creeping,
knotted, with radial dictyostele, densely clothed in
dark brown, ovate, denticulate scales with sparsely
glandular margin. Petioles rather close, well-developed, erect, medium brown, adaxially monosulcate, basally sparsely scaly and densely very short-pubescent,
otherwise glabrous. Lamina oblong, simply pinnate,
paucijugate, with conform terminal pinna, chartaceous, catadromous except at the extreme base, with
scattered, minute, unicellular, yellow glands. Rachis
stramineous, adaxially elevated in the middle, the
ridge flanked by two narrow grooves, these interrupted
at pinna insertions. Pinnae subsessile, subopposite,
lanceolate, crenate-serrate or distally lobulate, basally
unequally cuneate; costa adaxially convex; edges
thickened, continuous with the lateral rachis ridges;
surface subglabrous, a few ctenitoid hairs present on
rachis and veins; costae with a few minute, deciduous
scales. Costules distinct, flexuous; veins distinct, very
oblique, simple, most of them anastomosing in opposite pairs which are terminated by a short, excurrent
veinlet; basal pair and outer veinlets free; basal posterior veinlet usually springing from costa (not from costule). Sori orbicular, dorsal or less often compital, irregularly biseriate between costules; indusium none.
Sporangia glandular on the pedicel; spores shortly ellipsoid to subglobose, monolete; perispore rugulosesaccate; 2n = 82.
A single species, C. poecilophlebia (Hooker) Tindale,
terrestrial in rainforest, narrowly distributed in NE
Queensland, Australia.
Incertae seelis:
30. Adenoderris 1. Smith
AdenodelTi. J. Smith, His!. Fi!.: 222 (1875); Maxon (1905);
Kato (1984 b).
Small terrestrial or epilithic ferns with short, erect stem
bearing tan, lanceolate, long-tapering, glandular
scales. Leaves fascicled, short-petiolate to subsessile;
petiole ± scaly and glandular-hairy, with two vascular
bundles; lamina lanceolate, gradually narrowed to
both ends, deeply pinnatifid or pinnate (+pinnatifid),
shortly glandular-hairy, herbaceous or spongy; primary divisions oblong, obtuse to subacute, unequalsided at base, decurrent and adnate or subsessile, sinuate to pinnatifid; rachis adaxially grooved. Costa
deflexed at base, not percurrent; venation anadromous
or isodromous. Veins forked or the basal anterior pinnate, not or almost reaching the margin. Sori dorsal, or
terminal on short anterior branches, medial or supramedial, round; indusium peltate (to subcordate),
glandular-hairy. Spores bearing coarse, short, inflated
folds, somewhat perforate and papillate; X=?
Two species, in shaded situations at lower and middle elevation, both rare and local: A. glandulosa (Presl)
J.Smith in Cuba and Jamaica, A.sororia Maxon in
Guatemala.
Of controversial affinity; placed in Athyrioideae
(Physematieae) by Tryon and Tryon (1982), with which
it agrees in the number of petiolar bundles and the indument; near Tectaria by Proctor (1985) and near Polystichum by Kato (1984 b), which it approaches in the
shape of the indusium and of the xylem strands in the
petiolar bundles.
Subfamily Atbyrioideae
M. KAro; final version by K. V. KRAMER
Synon. Woodsiaceae (Diels) Herter, Rev. Sudam. Bot. 9: 14
(1949).
Athyriaceae Alston, Taxon 5: 25 (1956).
OnocJeaceae Pichi Sermolli, Webbia 24: 708 (1970).
Hypodematiaceae Ching, Acta Phytotax. Sin.13: 96 (1975).
Diverging from Dryopteridoideae in the following
characters: Petiole base not rarely swollen ("trophopod") or winged. A dorsiventral stele in the stem present in a few representatives (Hypodematium. some
Athyrium species). Petiole and rachis adaxially most
often grooved, the grooves on axes of higher order
continuous or not. Petiole with two vascular bundles,
the xylem in each omega-shaped in transection, the
two uniting above, sometimes in the 'rachis, into one
which is V-shaped in transection; only in some very
large species of Diplazium some subsidiary abaxial
bundles may be present. Veins more often free than in
Corenie/Ja • Adenoderris
Dryopteridoideae, when anastomosing the pattern goniopteroid, rarely sagenioid, never anaxetoid. Stomata
polocytic, anomocytic ones not rarely present besides.
Sori dorsal, rarely terminal (in Deparia section Deparia), most often elongate, linear, U- or J-shaped (then
crossing the vein), but may also be (sub)orbicular, then
with cup- or hood-shaped or reniform indusium, rarely exindusiate. Sporangia nearly always on long, slender pedicels. Spores chlorophyllous in the tribe Onocleeae.
ANATOMV. For anatomical data see, e. g., Tardieu-Blot
(1932), Bir (1969b), Kato (1972); the dermal appendages were described by Kato (1973).
131
1956 seems soundest; but here Sledge (1973b) is followed who treated the Athyrioid ferns as a subfamily
of Dryopteridaceae, pointing out the close morphological and karyological resemblance. Two very natural
tribes may be distinguished:
Tribe Physematieae (= Athyrieae)
Leaves not or only weakly dimorphic; sori not enclosed by revolute laminal parts of the fertile leaves;
sporangia of average size; spores achlorophyllous.
Genera 31 to 42.
Tribe Onocleeae
GAMETOPHVTE. This is much like that of Dryopteridoideae. Schlumberger's statement (1911) that the prothallium of Woodsia is somewhat like that of Cyatheaceae in having antheridia with divided cap cells has
proved incorrect. The prothallium is hairy on the
wings in Cystopteris and some species of Deparia, Di-
Sterile and fertile leaves strongly difform; fertile lamina much reduced, the laminal parts reflexed and enclosing the sori; sporangia large, with 24-35 indurated
bow. cells; spores chlorophyllous, short-lived (Lloyd
1971). Genera 43 to 45.
plaziopsis, Gymnocarpium, Hypodematium, Onoclea,
and Woodsia. The hairs are papilla-like and glandular;
setiform hairs, like those of the leaves, occur in Hypodematium.
KEv TO THE GENERA OF TRIBE PHVSEMATIEAE,
SUBFAMILY ATHVRIOIDEAE.
KARVOLOGV. The basic chromosome numbers are
somewhat more diverse than they are in the Dryopteridoideae, but 40 and 41 also prevail, the former in
Athyrium, nearly all Deparia species, Cornopteris,
Gymnocarpium, Hypodematium, Matteuccia, and some
species of Diplazium, the latter in Dictyodroma, Diplaziopsis, most species of Diplazium, some of Hypodematium, Deparia, and Cornopteris. 42 occurs only in Cystopteris, 37 in Onoc/ea, 31 in Hemidictyum, and 33, 38,
39 (besides 41) in Woodsia. The numbers are not quite
so constant in many genera as formerly believed, and
it is not advisable to draw generic lines only on the basis of this character. More karyological data may, however, help in delimiting the genera around Athyrium
more clearly, like Deparia, Diplazium, and Cornopteris.
Triploid, apogamous species have been found in Diplazium and Gymnocarpium. Polyploidy and hybridization are frequent, having been found especially in
Athyrium, Deparia, Diplazium, Gymnocarpium, Cystopferis, Cornopteris, and Woodsia. Intrageneric hybridization has been found between species of Athyrium
and Cornopteris,' see Lovis (1977).
ECOLOGY. Most species of Athyrioideae
shady, mesic habitats, whereas many species
dematium. Gymnocarpium. Cystopteris, and
occur on rocks. Some rheophytes have been
Deparia and Diplazium
grow in
of Hypo-
Woodsia
found in
SUBDIVISION. Ching (1978) split the group into four
families, which certainly goes too far; but his segregates are natural alliances. Alston's circumscription of
(these are also keyed out in the general key to the genera of Dryopteridaceae)
1. Sterile-fertile leaf dimorphism pronounced; sori round,
± distinctly indusiate, also covered by the revolute leaf
segments; spores chlorophyllous (Onocleeae)
see bracket 74 in the key on p.106
- Sterile-fertile leaves conform or weakly dimorphic; sori
not covered by the revolute leaf segments, not rarely elongate; spores achlorophyllous (Athyrieae)
2
2. Sori round, with inferior indusium
3
- Sori elongate, or, if round, the indusium not inferior, Or
wanting
4
3. Indusium attached to the posterior side of the receptacle;
38. Cystopteris
petiole never with an articulation
- Indusium surrounding the receptacle at its base; petiole
sometimes with an articulation above its base 42. Woodsia
4. Lamina glabrous, or, if hairy, the hairs not acicular; sari
various
5
- Lamina bearing acicular and (or) glandular hairs; sori
rounded, with reniform indusium
41. Hypo/kmatium
5. Indusium present, distinct
9
- Indusium wanting (or minute)
6
6. Basal pinnae articulate at base
40. Gymnocarpium
- No pinnae articulate at base
7
7. Bases of costae and costules adaxially corniculate with
37. Comopteris
fleshy, horn-like projections
- No such projections present
8
8. Stem slender, creeping; leaves with few to many multiseptate hairs·
39. Acystopteris
- Withoutthis combination of characters
31. Athyrium
9. Grooves of rachis and costae not continuous at junction;
leaf axes bearing multicellular, articulate hairs 36. Deparia
- These characters not combined
10
10. Veins free; rachis groove V-shaped in transection; sori
31. Athyrium
round-reniform, U- or J-shaped, or linear
- Veins free or anastomosing; rachis groove U-shaped in
11
transection; sori linear
Dryopteridaceae . Athyrioideae
132
o
Fig.64A-O. Dryopteridaceae. A Athyrium schimperi. fertile
segments (x 4). B Athyrium scandicinum. fertile segments
( x 4). C, O. Athyrium sheareri. C Rachis with two fertile pinnae (xO.8). 0 Indusium (x20) (A,B from Schelpe 1970;
C,O from Ching 1935)
31. Athyrium Roth
11. Lamina simply pinnate, with conform terminal pinna;
veins free and parallel in their proximal parts. anastomos12
ing in their distal parts
- Veins free, or anastomosing in a different pattern; lamina
various, sometimes simple
13
12. Indusium allantodioid; veins not connected by an intramarginal commissure
34. Diplaziopsis
- Indusium not allantodioid; veins connected by an intramarginal commissure
35. Hemidictyum
- Indusium not aIlantodioid; veins not connected by an intramarginal commissure
32. Diplazium
13. Indusium aIlantodioid, conspicuous
34. Diplaziopsis
- Indusium not aIlantodioid, evident, rarely evanescent 14
14. Veins free, Or goniopteroid- or sagenioid-anastomosing,
ending near the margin; both parts of "double" sari of
about equal length
32. Diplazium
- Veins with sagenioid anastomoses, ending well behind the
margin; both parts of "double" sari in many cases only
partly "back-to-back", a considerable part of such a sorus
simple
33. Dictyodroma
Stem long-creeping or ascending to erect, nearly always with a radial dictyostele; scales brown or blackish, entire. Leaves remote to (usually) aggregate; petiole very short to well-developed, the base often
swollen ("trophopod") and winged, bearing pneumatho des ; adaxial side sulcate, the groove usually Vshaped in transection; scales often present at base, less
often the petiole and rachis densely and persistently
scaly. Lamina linear to broadly triangular, reduced at
base or not, mostly herbaceous; dissection pattern
variable, in the primary divisions often anadromous or
isodromous at base and catadromous towards the leaf
apex, or quite anadromous, rarely entirely catadromous. Groove of rachis continuous with that on the
axes of higher order. Pinnae entire or lobed to pinnatilid, or up to tripinnate ( + pinnatilid); upper pinnae,
pinnules, etc. gradually reduced and confluent. Surface of lamina glabrous or minutely (not rarely glandular-) hairy; axes often minutely hairy in the groove,
rarely densely and persistently scaly. Costae and costules adaxially sulcate. Not rarely spine-like appendages present on the adaxial side at the bases of costae
Fig. 64
Athyrium Roth, Tent. Fl. Germ. 3: 58 (1799).
Anisocampium C. Presl (1849); Sledge (1962).
Pseudocystopteris Ching (1964).
Cystoathyrium Ching (1966).
Kuniwatsukia Pichi Sermolli (1973).
Athyrium • Diplazium
and costules. Pinnae and pinnules not rarely asymmetric or the anterior side auriculate. Fertile divisions occasionally somewhat contracted. Veins most often anadromously branched, free, rarely anastomosing. Sori
dorsal, from inframedial to subterminal, most often
unequally horseshoe-shaped, J-shaped, or linear, less
often equally horseshoe-shaped or reniform to orbicular, indusiate; the indusium of the same shape as the
sorus, sometimes fugacious, rarely lacking. Spores ellipsoidal, with prominent, short, inflated, connected
folds, rarely bearing wing-like projections; x- 40.
About 180 species, of worldwide distribution but
strongly concentrated in the northern hemisphere, especially in eastern and south-eastern Asia and the Himalaya and adjacent mountain chains; comparatively
few species in tropical and southern Africa and South
America, very few in Europe. Terrestrial in thickets
and forests, also in open vegetation and on rocks, especially at higher elevation; in warmer regions virtually confined to higher altitudes. The taxonomy is poorly known, the genus being in great need of critical
revisional work. A natural subdivision is also lacking;
a first attempt in this direction was made by Hsieh
(1986).
CHARACTERS OF RARE OCCURRENCE. Entirely catadromous leaves are found in, e. g., A. cuspidatum (Bedd.)
M. Kato; goniopteroid anastomosing veins in A. cu-
133
Fig. 65. Dryopteridaceae. Diplazium kawakamii. part of fer·
tile lamina with sori; Taiwan. Phot. Ch.-M. Kuo
mingianum (C. Presl) Milde; sori that are exindusiate
from an early stage of development in A. distentifolium
(Tausch) Ryl.; a creeping, dorsiventral stem in A. cumingianum (c. Presl) Milde. The last-named species is
usually placed in the satellite genus Anisocampium, but
this is connected with Athyrium by the intermediate
A. sheareri (Baker) Ching and the genus cannot be well
maintained. Simply pinnate leaves with lanceolate, entire pinnae, veins that reach the margin, and contracted fertile pinnae are found in A. pycnocarpon (Sprengel) Tidestr.
32. Dip/azium Swartz
Figs. 65, 66
Dip/azium Swartz, Schrad. J. Bot. 18002 : 61 (1802).
Callipteris Bory (1804).
Rhachidosorus. Ching (1964).
Trib/emma (J. Smith) Ching, Acta Bot. Sin. 11: 24 (1978).
Medium-sized to large, less often small, terrestrial or
epilithic ferns; stem creeping or more often erect,
sometimes trunk-like (up to 1 m tall), bearing brown to
blackish, entire or dentate scales, the teeth consisting
of the upturned extremities of two adjacent marginal
cells. Petiole usually well-developed, often stout, occa-
Dryopteridaceae • Athyrioideae
F
Fig. 66A-G. Dryopteridaceae. A Diplazium nemorale. pinnule lobe with sori (x 4). B Dip/azium zanzibaricum. pinnule
lobes with sori (x 4). C-G Diplazium asperum. C Rachis and
pinna base from adaxial side (x 1.5). D Transection of rachidule (x 3). E Part of rachidule and bases of pinnules from
adaxial side (x7). F Lobes of fertile pinnule (x3.5). G "Single" sorns showing fringed indusium (x 10) (A,B from
Schelpe 1970; C-G from Holttum 1955)
sionally muricate, rarely persistently scaly, adaxially
sulcate. Lamina simple or pinnatifid, or more often
pinnate to quadripinnate (+ pinnatifid), not rarely
large; terminal pinna conform, or the lamina and pinnae gradually reduced to apex; basal pinnae rarely reduced; dissection pattern as in Athyrium but the divisions usually less cut and less unequal-sided at base;
segments not rarely approximately parallel-sided and
lanceolate to tongue-shaped. Rachis occasionally bearing adventitious buds. Adaxial groove of rachis usually with flat bottom, glabrous; no spine-like appendages at bases of costae and costules which are
adaxially grooved. Ultimate divisions costate, the costa
anadromously pinnate, veins simple or forked, free or
less often goniopteroid or sagenioid anastomosing,
without free included veinlets. Sori dorsal, often medial, linear, on an acroscopic veins branch, nearly always
distinctly indusiate, the indusium linear, attached
along the vein, anteriorly free, or usually on both sides
of the largest basal vein branch of a costal costule,
then the free indusium edges directed to both sides;
rarely the indusium very small or fugacious. Spores
with long, usually prominent, wing-like folds, these often with echinate borders, sometimes cristate or echinate; X= 40 or 41,
An estimated 400 species (but probably fewer), terrestrial in forests and thickets, occasionally epilithic;
very widespread in the warmer parts of the world,
sparingly and only locally extending into temperate
areas. In the tropics occurring from the lowlands to
the mountains. Pew species in continental Africa.
A taxonomically very difficult and quite insufficiently known genus, in great need of monographic
study. Not yet full-grown plants may be fertile and difficult to assign to a species. A natural subdivision of
the genus has not been given.
Closely related to Athyrium. and for some time
united with it, following Copeland. Currently again
Dictyodroma . Diplaziopsis
135
treated as a distinct genus, especially in view of the
divergent chromosome numbers; but n- 41 is not
quite consistent in Diplazium. It seems artificial to assign species to one or the other genus on the basis of
the chromosome number alone (e.g., Jermy 1964).
CHARACTERS OF RARE OCCURRENCE. Simple, freeveined leaves in, e. g., D. plantaginifolium (L.) Urban,
D. subsinuatum (Wall. ex Hooker & Grev.) Tagawa,
and D. subserratum (B!.) Moore; simple leaves with
(distally) sparingly to amply anastomosing veins in
D. praestans (Cope!.) Morton and D. cordifolium B!.;
in D. aberrans Maxon & Morton even the sori are reticulate. Basally pinnate, upward pinnatifid leaves with
broad, very obtuse pinnae and sparingly anastomosing
veins in D. pinnatifidum Kze., some relatives freeveined. Amply goniopteroid anastomosing veins in
D.proliferum (Lam.) Kaulf., D.accedens B!., and their
relatives ("Callipteris").
33. Dictyodroma Ching
Fig. 67
Dictyodroma Ching, Acta Phytotax. Sin. 9: 57 (1964).
Small to medium-sized, terrestrial ferns with rather
short, ascending to erect stem bearing brown, entire
scales. Leaves close; petiole well-developed, fuscous,
bearing fleshy scales at base, adaxially unisulcate.
Lamina narrowly to broadly triangular, simple and
pinnatifid or usually pinnate at the base, the upper
pinnae gradually shortened and very broadly ad nate
and connected; rachis adaxially shallowly sulcate,
bearing small scales and multicellular hairs. Texture
herbaceous or fleshy. Pinnae acuminate, entire or sinuate; costa percurrent, when dry also abaxially shallowly grooved; venation catadromous except in the basal
pinnae. Veins amply sagenioid-reticulate without free
included veinlets, ending behind the margin with often
free, thickened ends. Sori linear, single, or the larger
ones at least partly double, i. e., on both sides of a
vein, but the sori on two different sides of a vein usually only partly overlapping; often some sori weakly reticulate; indusium of the same shape. Spores as in Diplazium; x-41.
Five described species, but probably only two or
three, from the central Himalaya and South China to
Indo-China and Taiwan; terrestrial in forest. Perhaps
better included in Diplazium, like the next.
32. Diplaziopsis C. Chr.
Fig.68C,D
Diplaziopsis C. Chr., Ind. Fil.: 227 (1905), as renaming of AIlantodia Wallich (1830), non R. Brown (1810); Ching
1964a); Bir (1969a, b).
Medium-sized, terrestrial ferns with rather short to
long, stout, erect, fleshy stem bearing brown, entire
scales that are glandular when young. Leaves close,
Fig. 67 A-D. Dryopteridaceae. Dictyodroma formosana.
A Habit (x 0.5). B Part of fertile pinna showing partly "back
to back" sori (x 3). C Scale of stem (c. x 15). D Glandular
trichome from abaxial side of lamina (enlarged) (Fl. Taiwan
1,1975)
medium-sized to rather large; petiole well-developed,
not very dark, adaxially unisulcate, sparsely scaly at
base. Lamina simply pinnate with conform terminal
pinna, oblong, not reduced at base, the upper pinnae
scarcely shortened; surface naked, texture herbaceous.
Pinnae sessile, oblong, acuminate, not numerous, entire; costa percurrent, adaxially shallowly sulcate, the
groove evanescing at base; venation isodromous beneath, catadromous above in lamina. Veins forked at
their extreme bases, the posterior branch about at right
angles with the costa; about the outer half of the venation reticulate, forming hexagonal meshes without included veinlets (sagenioid); outermost veins ending
behind the inargin, thickened, mostly free. Sori linear,
on the anterior vein branch up to about the first mesh,
single or double; indusium very convex, initially allantodioid, enveloping the sporangia, bursting irregularly
or asplenioid at maturity, bearing glandular hairs.
Spores as in Diplazium; x-41.
Three or four species; one, D.javanica (B!.) C. Chr.,
widespread in S. and E. Asia east to Samoa; the others
Dryopteridaceae • Athyrioideae
136
B
Fig.68A-E. Dryopteridaceae. A,B Depariajaponica. A Por·
tion of rachis and two pinna bases, adaxial side (x 2.5).
B Abaxial side of fertile segment (x 5). C,D Diplaziops/s javanica. C Habit (x 0.5). D Part of fertile pinna showing allantodioid indusium (x 3). E Hemidictyum marginatum. part
of fertile pinna (x 1) (A,B from Duncan and Isaac 1986;
C from Flora Thiwan 1, 1975; D from Tardieu-Blot and
Christensen 1939; E from Vareschi 1969)
more localized in Japan and China. Terrestrial in forest, at middle elevation.
Included in Diplazium by some authors, which is
not unnatural; but the combination of erect, fleshy,
soft stem, half-reticulate venation (about as in Hemidictyum), and allantodioid indusia is distinctive.
35. Hemidictyum C. Presl
Fig.68E
Hemidictyum C. Presl, Tent. Pteridogr.: 110 (1836).
Diplaz;um auctt., p. p. min.
Stem stout, erect, bearing brown, entire scales; roots fibrous. Petiole stout, well-developed, scaly at base, pale
to dark brown. Leaves very large, simply pinnate with
subopposite, sessile, plurijugate pinnae, the lower not,
the upper little reduced, and with conform terminal
pinna; outline elongate-oblong, base subtruncate, apex
shortly acuminate, margin entire to sinuate, surface
naked. Costa pale, percurrent, adaxially sulcate; venation catadromous. Veins free and forked in the proximal % or 31., reticulate without free veinlets in the outer
part, their ends fully connected by a continuous intramarginal commissure. Sori very long and narrow,
on the free, basal parts of the veins, on their anterior
branches, single or double; indusium membranous,
not allantodioid. Spores ellipsoidal to subspheroidal,
with prominent, inflated, perforate folds; x=31.
A single species, H. marginatum (L.) C. Presl [when
included in Diplazium the species must be called
D.limbatum (Will d.) Proctor]; terrestrial in forest, especially in creek ravines; widespread in tropical America.
Kato (1975) and others indicated relationship to Diplaziopsis and to certain species of Diplazium; but the
chromosome number militates against this; see also
Jermy and Walker (1985).
36. Deparia Hooker and Greville
Fig. 68 A, B
Deparia Hooker and Greville, Icon. Fil.: pl.154 (1829); Kato
(1977,1984a).
Lunathyrium Koidzumi (1932); Ohba (1965,1966).
Dryoathyrium Ching (1941).
Parathyrium Holttum (1959).
Athyriops/s Ching (1964).
Small to medium-sized or occasionally large ferns with
long-creeping to erect, often fleshy stem bearing entire
scales, these often with multicellular hairs, Leaves remote to close; petiole slender to stout, well-developed,
stramineous to medium brown with darker base, adaxially unisulcate, basally winged and then often bearing
pneumathodes, or unwinged, often hairy and/or scaly.
Lamina triangular to linear, basally truncate, simply
Hemidictyum . Deparia· Carnaptens
pinnate to bipinnate + pinnatifid, weakly catadromous
or anadromous; texture herbaceous or somewhat
fleshy. Rachis adaxially sulcate, the groove not open to
those of the pinna axes or costae. Septate, sometimes
somewhat scale-like hairs mostly present on leaf axes.
Venation anadromous; veins free, forked or pinnate,
not reaching the margin. Sori orbicular and then often
at a vein fork, or V- or J-shaped to linear and then
dorsal, or exceptionally terminal; indusium of the
same shape as the sorus, laterally attached. Spores ellipsoidal to subspheroidal; surface with diffuse echinae projecting from an appressed reticulum; perispore
2-layered, the reticulum formed by the outer; X= 40,
rarely 41.
About 40 species, widespread in the tropical and
warm-temperate parts of the Old World, east to
Hawaii; several in Madagascar, one in Africa; one
species in North America; locally naturalized in Europe and tropical America.
Four quite distinct sections were established by
Kato, with the following characters: Section Deparia:
Lamina pinnate+pinnatifid to bipinnate, sometimes
gemmiferous; septate hairs wanting or very few; sori
linear, or round and terminal, on vein ends projecting
beyond the margin in some forms of D. prolifera
(Kaulf.) Hooker & Oreville (for which the genus was
originally erected), with seemingly almost Dicksonialike indusium; edge of indusium entire. One species in
the Bonin Is., four in Hawaii, all local.
Section Lunathyrium (Koidzumi) M. Kato: Petiole
base winged, with pneumathodes; septate hairs present; lamina pinnate + pinnatifid to bipinnate + pinnatifid, pinnae enlarged at anterior base; sori linear,
J- or V-shaped; indusia rather frrm, ± bullate, entire
or dentate. C. 10 species, mostly in central and eastern
Asia, one in North America.
Section Dryoathyrium (Ching) M. Kato: Leaves
usually large, pinnate + pinnatifid to tripinnate + pinnatifid; petiole base with or without pneumathodes;
septate hairs evident in most species; sori rather small,
numerous, orbicular, horseshoe- or J-shaped, or linear;
indusium lacerate. C 10 species in the Old World, especially in East and South-East Asia and Madagascar;
D. boryana (Willd.) M. Kato widespread.
Section Athyriopsis (Ching) M. Kato: Leaves medium-sized, simply pinnate to bipinnate; petiole base
not winged; septate hairs present; sori linear or
J-shaped, indusium membranous, lacerate. At least
14 species, from central Asia to the Pacific Islands and
Australia; the most widespread is D.petersenii(Kunze)
M. Kato, with many local forms, introduced outside its
native area in various parts of the world. For the relationship between the sections and hybrids see Kato
(1984 a).
137
Fig. 69 A-C. Dryopteridaceae. Comopteris hirii. A Habit
( x 0.75). B Fertile segment (x 7). C Multicellular laminar tricbome (x 120) (Bir 1964)
37_ Conwpteris Nakai
Fig. 69
Comopteris Nakai, Bot. Mag. (Tokyo) 44: 7 (1930); Kato
(1979).
NeoathyriumChing & Z. R. Wang, Acta Phytotax. Sin. 20: 76
(1982).
Medium-sized to rather large terrestrial ferns with
creeping to short and erect stem bearing fleshy roots
and brown, entire scales. Leaves close; petiole welldeveloped, with fleshy base. Lamina elongate-triangular, truncate at base, pinnate + pinnatifid to tripinnate+pinnatifid; lower pinnae subopposite, upper
138
Dryopteridaceae . Athyrioideae
pinnae, pinnules, etc. gradually reduced and confluent; dissection pattern fluctuating. Rachis adaxially
grooved, the groove continuous with those of axes of
pinnae, etc.; axes and veins septate-hairy or glabrescent. Veins anadromously pinnately branched, the
branches simple or forked; costae and costules adaxially with fleshy, hom-like appendages at their bases.
Sori orbicular to linear, or rarely small and reniform;
indusium wanting. Spores ellipsoidal to somewhat
spherical, bearing prominent, short, connected folds;
x=40 or 41.
Nine species, from the Himalaya to eastern and
south-eastern Asia. Terrestrial in forests at middle to
higher elevations. For the non-recognition of Neoathyrium. see Kato (1986).
38. Cystopteris Bernhardi
Cystopteris Bernhardi, Schrad. J. Bot. I': 26 (1806), nom. conserv.; Blasdell (1963); Pearman (1976); Monso (1982).
Small to medium-sized, terrestrial or often epilithic
ferns with creeping to ascending stem bearing membranous, entire scales. Leaves close to remote; petiole
well-developed, slender, stramineous to medium
brown, adaxially sulcate. Lamina thinly herbaceous (in
one species "filmy": C. membranifolia Mickel), glabrous, or the axes deciduously scaly, broadly to very
narrowly triangular, the lower pinnae hardly or not reduced, bipinnate to tripinnate + pinnatifid; basal pinnae sometimes basitonically produced. Dissection pattern anadromous, rarely isodromous; upper divisions
gradually reduced and confluent. Rachis occasionally
bearing bulbils; axes adaxially grooved, the grooves
continuous. Veins pinnately branched, anadromous,
free, with simple or forked branches, their ends scarcely thickened, placed very near the margin in teeth or
between them. Sori dorsal, orbicular, with distinct,
scale-like, eventually often reflexed indusium attached
below the sporangia at the posterior side of the receptacle. Spores ellipsoidal to spheroidal, echinate, usually coarsely so, sometimes perforate; surface sometimes
coarsely low-rugose with papillate elements; X= 42.
Twelve species, on forest floor or in open places, especially on rocks; very widespread in temperate zones,
montane to alpine in tropical areas.
39. Acystopteris Nakai
Acystopteris Nakai, Bot. Mag. (Tokyo) 47: 180 (1933).
Cystopteris subgen. Acystopteris (Nakai) Blasdell (1963).
In many respects like Cystopteris. Medium-sized, terrestrial ferns with long-creeping stem. Leaves remote,
long-petiolate; lamina elongate-triangular, bipinnate + pinnatifid to tripinnate + pinnatifid, isodromous
or, especially above, catadromous; pinnae sessile.
Lamina, especially its axes, bearing non-glandular,
multi septate hairs. Indusium small, not cup-shaped,
fugacious or wanting; spores very densely verrucose,
x=42.
Two species in Asia, from the Himalaya to Japan
and Malesia. The perispore is very different from that
of Cystopteris. In appearance not unlike certain species of Thelypteridaceae.
40. GymlWCarpium Newman
Gymnocatpium Newman. Phytol. 4: 371 (1851); Sarvela
(1978. 1980).
Cu"ania Copeland (1909).
Carpogymnia D & A. LOve (1966), nom. superfl.
Rather small, terrestrial ferns with slender, long-creeping, dark stem; scales membranous, entire, glandularhairy or not. Leaves remote; petiole well·developed,
slender, pale, naked or scaly at base, or bearing short
glands. Lamina deltoid to pentagonal in outline, thinly
herbaceous, borne at an angle to the petiole, pinnatifid
to tripinnate, almost consistently anadromous, naked
or minutely glandular; larger pinnae (sub)opposite.
Rachis and costae adaxially sulcate, the grooves not
continuous. Pinna rachises and larger costae articulate
at base. Veins anadromously branched, the branches
mostly simple, at least the outer almost reaching the
margin. Sori dorsal, orbicular to elliptic, naked (a reniform indusium rarely found in one species). Spores ellipsoidal, bearing slightly to prominently perforate
folds, these sometimes contracted to verrucae; n=40.
Six species, terrestrial in forests and open places
and on rocks and rubble: four north-temperate, one in
Taiwan, one from the Himalaya to New Guinea.
Subgenus Cu"ania (Copel.) Sarvela: lamina pinnatifid; sori distinctly elliptic. Only G.oyamense (Baker)
Ching. Subgenus Gymnocarpium: lamina bipinnate or
more dissected; sori orbicular or weakly elliptic. The
remaining five species.
The genus has been variously placed, e. g., in Thelypteridaceae, or next to Dryopteris. or in the Athyrioid
ferns. The evidence is not unequivocal, but vascular
anatomy, leaf architecture, and karyology point to the
last-named placement.
41. Hypodematium Kunze
Fig. 70
Hypodematium Kunze. Flora 1833': 690; Ching (1935); Iwatsulci (1964).
Terrestrial or mostly epilithic, medium-sized ferns;
stem short-creeping, with dorsiventral dictyostele, the
leaves restricted to the dorsal, the roots to the ventral
side, densely covered with large, rufous-brown, entire,
glabrous scales. Leaves close; petiole swollen at base,
well-developed, stramineous, scaly at base, thinly hairy
with acicular hairs, adaxially like the other axes sul-
Cystopteris • Acystopteris . Gymnocarpium • Hypodematium
139
C
cute, all grooves continuous. Lamina triangular, truncute at the base, firm-herbaceous, bipinnate + pinnatilid to tripinnate + pinnatifid, the basal pinnae basitonil'ully produced; upper divisions gradually reduced and
confluent Dissection largely or entirely anadromous.
I.cafaxes and veins bearing acicular and! or glandular
Fig.70A-E. Dryopteridaceae. Hypodematium crenatum.
A Stem and leaf (x 0.5). B Fertile tertiary pinnule (x 6).
C Scale from petiole base (x 10). D Soms (x 18). E Hairs
from indusium (x 30) (A,D,E from Sche\pe 1970; B,C from
Ching 1935)
140
Dryopteridaceae • Athyrioideae
hairs. Ultimate divisions obtuse, incised. Veins ana·
dromously pinnate, the branches simple or forked,
ending at the margin. Sori dorsal or at a bifurcation,
large, orbicular; indusium convex, reniform, hairy, in
one species as in Cystopteris; spores ellipsoidal, bear·
ing prominent, inflated tubercles with papillate ele·
ments; X= 40 or 41.
An Old World genus of four species, one widespread from West and South Africa to East Asia, the
others East Asiatic only. Growing preferentially on
limestone. A somewhat isolated genus of controversial
position, now generally placed in the Athyrium group.
42. Woodsia R. Brown
Fig. 71
Woodsia R. Brown, Prodr. Fl. Nov. Hall.: 158 (1810);
D. Brown (1964).
Physematium Kaulfuss (1829).
Hymenocystis C. A Meyer (1831); Askerov (1986).
Cheiianthopsis Hieronymus (1920).
Protowoodsia Ching (1945).
Terrestrial or very often epilithic, small to mediumsized ferns with short, creeping or ascending stem
bearing persistent petiole bases and entire scales.
Leaves close; petiole slender, dark at least at base,
mostly shorter than the lamina, in some species with
an oblique articulation above the base. Lamina thin,
herbaceous, to firm, ovate to linear-Ian ceo late, the basal pinnae usually reduced and remote, the upper divisions reduced and confluent; dissection pattern anadromous or often isodromous. Narrow, approximately
hair-like scales and one- or few-celled hairs usually
present on the lamina. Pinnae rarely pseudo-articulate
at base. Veins pinnate, the branches simple or forked,
free. Sori dorsal to subterminal, orbicular; indusium
inferior, subglobose and sac-like or composed of a ring
of hairs, with various intermediates; occasionally reflexed, modified laminar lobes representing a kind of
second, "outer" indusium. Spores ellipsoidal to somewhat spherical, bearing folds, these sometimes compressed into short, wing-like structures, somewhat
echinate, sometimes perforate or inflated and coarsely
rugose; x=33, 38, 39, 41About 25 species; several· hybrids; very widespread
in the north-temperate zone and in the mountains of
tropical America; south-temperate in South-America
and South-Africa; absent from Australia, New Zealand, SE Asia, and the Pacific. Terrestrial, or very often growing in rock crevices.
Formerly the genus was placed in various alliances
on account of its unusual inferior indusium; but when
the morphology of the plant as a whole is considered,
it finds its natural place in the Athyrioideae. Nevertheless, its place there is somewhat isolated, also in its
chromosome number which in only some species with
x=41 agrees with what is found otherwise in many of
the genera. Ma's suggestion (1985), made on karyolog-
FIg.71A-C.
Dryopteridaceae. Woodsia montevidensis.
A habit (x 0.5). B Base of fertile pinna (x 10). C Lamina!
hairs (x 40) (de la Sola 1977)
ical grounds, that W. is related to Gleicheniaceae can
hardly be seriously considered.
43. Matteuccia Todaro
Fig. 72
Matteuccia Todaro, Syn. Plant Vase. Sicil.: 30 (1866); Lloyd
(1971); Kato and Sakahashi (1980).
Struthiopteris Willdenow (1809), non Scopoli (1760) nee Weis
(1770) nee Bernhardi (1802).
Pteretis Rafinesque (1818), nom. superfl.
Medium-sized ferns with erect, trunk-like, stoloniferous stem, the apex bearing membranous, entire scales.
Leaves tufted in a crown; petiole short, winged above
and again tapering towards the base, there dark, other-
Woodsia· Matteuccia . Onocleopsis· Onodea
141
growing season, chlorophyllous, short-lived, ellipsoidal to somewhat spherical, bearing few large folds with
minutely cristate to echinate elements; X= 40.
Terrestrial in forests and by streams; two species,
M. strutniopteris (L.) Tod., widespread in the northern
hemisphere, and M. intermedia C. Chr., a hybrid of the
first species with Onodea orientalis (see there).
The presence of cataphylls, without a lamina, was
described by, among others, Von Aderkas and Green
(1986).
44. Onocleopsis Ballard
Onocleopsis Ballard, Amer. Fern J. 35: 1 (1945; 1948); Uoyd
(1971).
Fig.72A, B. Dryopteridaceae. Matteuccia orientalis. A Part
of fertile lamina (x 1). B Transection of fertile pinnae (x 9)
(Flora Tsinlingensis 1974)
wise lighter, adaxially sulcate like the rachis. Lamina
strongly dimorphic, catadromous (or isodromous),
much reduced towards the base. Sterile lamina oblanceolate, pinnate + pinnatifid, herbaceous, thinly and
deciduously hairy, more persistently so at the pinna
bases. Pinnae numerous, sessile, truncate at the base,
lhe basal segments often somewhat enlarged; upper
divisions gradually reduced and confluent. Costa
adaxially sulcate, the groove not continuous with that
of the rachis. Veins pinnate in the segments, catadromously branched, the branches simple, free, almost
reaching the margin. Fertile lamina indurated and with
greatly reduced laminar parts, simply pinnate; margin
of pinnae reflexed, covering the sori. Sori orbicular,
terminal, with membranous, cup-like, extrorse indusium. Spores retained in the fertile leaves until the next
Medium-sized to large terrestrial ferns; stem stout,
short-creeping or ascending to erect, stoloniferous or
not; apex bearing pale brown scales. Leaves aggregated in a crown, rather short-petiolate; lamina slightly
scaly, short-pubescent, strongly dimorphic. Sterile
leaves c. 0.5-1.8 m long, simply pinnate; pinnae numerous, with truncate-subcordate base, narrowly lanceolate, acuminate, serrate-crenate to lobed; lower
pinnae reduced, upper pinnae abruptly confluent.
Veins reticulate except for the free excurrent branches,
with elongate costal areoles, without free included
veinlets. Fertile leaves shorter, bipinnate + pinnatifid
to tripinnate, not indurated, not persisting till the next
growing period; segments free-veined, thin, with revolute margin coyering the sori, irregularly rupturing at
maturity; sporangia assembled in round, terminal sori,
borne at different levels .on an erect receptacle, initially
covered by a hyaline, fugacious indusium. Spores el,lipsoid, bearing coarse, prominent folds with dense
echinate-rugose elements; x=40.
A single species, O. hintonii Ballard, in Mexico and
Guatemala, in moist mountain canyons. Included in
Matteuccia by Kato and Sakahashi (1980) but diverging in its thin, non-persistent fertile leaves and its reticulate venation, in which character it is clo~er to the
next genus.
45. Onoclea L.
Fig. 73
Onoclea L., Spec. Plant. 2: 1062 (1753); Uoyd (1971); Kato
(1980).
Pentarhizidium Hayata (1928).
Medium-sized terrestrial ferns with creeping stem
bearing membranous, entire scales. Leaves rather
close; petiole well-developed, like the rachis adaxially
flat, non-sulcate. Lamina triangular or oblong, deeply
pinnatifid to (more often) pinnate + pinnatifid, strongly dimorphic; sterile pinnae narrowly elliptic to Iinearlanceolate, the basal weakly to strongly narrowed at
base; veins free or reticulate without freeveinlets, catadromous. Fertile leaves shorter, indurated, persisting
142
Dryopteridaceae . Athyrioideae
linear pinnae; x=40. O. orientalis (Hooker) Hooker in
the Himalaya and East Asia. Formerly included in
Matteuccia, with which it agrees in chromosome number and free venation, and with which it forms a hybrid; in other characters nearer to O. sensibilis.
Selected Bibliography
Fig. 73A-D. Dryopteridaceae. Onoclea sensibilis. A Habit
(x 0.3). B Part of sterile pinna (x 2). C Fertile pinnules
(x 6.5). 0 Vascular bundles of petiole and rachis (schematic)
(A-C from Petrik-Ott 1979; 0 from Ogura 1972)
to the next growing season; lamina once or twice pinnate, the laminal parts much reduced, revolute and
covering the sori. Sori, indusia, and spores as in Mat-
teuccia.
Two species, terrestrial in forests and thickets in
north-temperate regions, one in each of the following
sections:
Section Onoclea: Sterile leaves with anastomosing
veins; fertile leaves bipinnate, with bead-like segments/ pinnules; X= 37. O. sensibilis L., widely distributed in central and eastem Asia and eastern North
America.
Section Pentarhizidium (Hayata) M. Kato: Sterile
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Filicatae • Gleicheniaceae
Gleicheniaceae
K.U.KRAMER
Gleicheniaceae (R. Brown) C. Presl, ReI. Haenk. 1: 70 (1825)
("Gleichenieae").
Dicranopteridaceae Ching, Acta Phytotax. Sin. 3: 94 (1954).
Stromatopteridaceae (Nakai) Bierhorst, Phytomorph. 18 (2):
263 (1968).
Terrestrial ferns of rather small to very large size with
slender, at least partly creeping, protostelic (with one
exception) stem bearing scales andlor hairs. Petiole
non-articulate, terete or adaxially flattened, with a single, in cross-section C-shaped bundle. Lamina at least
once pinnate; segments adnate, ±firm in texture, freeveined. Sori round or nearly so, exindusiate, dorsal,
with rather few to very few sporangia, these maturing
simultaneously, short-stalked with large capsule and
many spores; annulus with oblique, nearly complete
bow and ± defined stomium, not interrupted by the
stalk. Spores achlorophyllous.
Two subfamilies, distinguished as follows:
- Lamina simply pinnate; leaves borne on erect stem
branches; venation anadromous
Stromatopteridoideae
- Lamina at least once forked, above the fork(s) pectinatepinnatifid or -pinnate; leaves borne directly on the hori-
zontal stem; venation catadromous
145
dodichotomously forked with dormant apices lying
between the bifurcations ("leaves indeterminate"), or
sometimes the primary or secondary rachis continued
for one or a few nodes beyond the primary dichotomous branching; apices protected by rudimentary segments andlor dermal appendages, together forming a
dormant bud. Leaf architecture and venation catadromous. Ultimate segments pectinately arranged on the
ultimate rachis branches, sometimes also borne on
some of lower order, roundish or more often linear to
tongue-shaped, numerous, of firm texture, with costa
and subpinnately branched lateral veins or, in Gleichenia, subecostate; veins free, with slightly thickened
ends near the margin. Sori round, exindusiate, usually
only one per (lowermost anterior) vein branch; receptacle small, raised. Scales or branched hairs present
among the sporangia or not. Sporangia few (c. 2-20)
per sorus, with a few- to multiseriate stalk. Stomium
cells several. Spores monolete or trilete. Gametophyte
elongate to subcordate, with long-necked archegonia
on the midrib.
ANATOMV AND MORPHOLOGY. The most important
contributions are by Chrysler (1943, 1944) and Holttum (1957). The former described the peculiar, Cshaped petiolar xylem strand with strongly involute
margins which may touch and then form a tubular
strand. He also showed that the dichotomous branch-
Gleichenioideae
Subfamily Gleichenioideae
Stem horizontal, long-creeping; leaves dorsal, uniserilite, at least once, usually several to many times pseu-
Fig. 74A~C. Spores of Gleicheniaceae, Nephrolepidaceae,
and OphlOglossaceae (all x 1000). A Dicranopteris dichotorna. B Nephrolepis exaltata. C Botrychium lunaria. Pho!.
A. F.Tryon
146
Gleicheniaceae . Gleichenioideae
J
Fig. 75 A-J. Dermal appendages of Gleicheniaceae. A Diplopteryg;um longissimum. B Sticherus hispidus. C Sticherus
truncatus. D Detail of C. E Dicranopteris curranii. F, G Dicranopteris pubigera. H Dicranopleris linearis var. subferruginea. I Dicranopteris linearis var. linearis. J Dicranopteris
linearis var. subspeciosa (all x 75 except C which is x 25)
(Holttum 1957)
ing is in reality pseudodichotomous. Wagner (1957)
described the blastogeny of Dicranopteris linearis
which begins with pinnate leaves, their basal pinnae
becoming more and more enlarged in successive stages
and leaf-apex aborting, i. e., remaining donnant. Simply pinnate early stages of leaves with gradual development of apical donnancy were described for Gleichenia (s. str.) and for Diplopterygium by Holttum
(1957 a).
The venation pattern is catadromous, the basal vein
of a segment, on the posterior side, springing from the
costule base or from the costa. The segments themselves can often (but not always distinctly) be seen to
be catadromously arranged.
Holttum (1975a) also discussed the dermal appendages (Fig.75); he showed the branched hairs of Dicranopteris to be essentially different from truly stellate
hairs of other fern genera (see Fig. 88). The leafy parts
are very densely hypostomatic, the stomata anomocytic and diacytic.
As shown again by Holttum, the sori are dorsal in
all genera. Bierhorst (1971) argued on ontogenetic
grounds that the receptacle must be regarded as tenninal, not superficial, and the leaf must be regarded as a
three-dimensional structure. His contention that the
leaves themselves are non-lateral organs was refuted
by Hagemann and Schulz (1978), who also described
the unique ontogenetic patterns of branching for representatives of three genera.
The sporangium has a short, usually multiseriate
stalk, a large capsule, and an oblique annulus with the
bow interrupted only by the stomium. The occurrence
of both trilete and· monolete spores in the family and
even within single genera is an uncommon feature.
The spore structure of a few species was described in
detail by Lugardon (1974). External features are few;
the perispore is relatively thick and ± smooth (Fig. 74).
Diplopterygium • Sticherus
GAMETOPHYTE. This is long-lived, green, subcordate to
ribbon-shaped, with a thick central strand, as described by Rauwenhoff (1889) and Stokey (1950); the
marginal wings are ruffled. The archegonia are longnecked and curve forward, the antheridia many-celled
and scattered over the lower surface. Two-celled hairs
arising from a wedge-shaped initial are frequent.
Spore germination is by a protonema-like filament
with transverse divisions (Nayar and Kaur 1971).
ECOLOGY AND DISTRIBUTION. Most species occur in
open, often strongly disturbed and!or pioneer habitats, on lateritic soil; the scandent species may cover
bushes and trees at forest edges. Erect species (especially Diplopterygium) may occur in dense, low, thicket-like stands. All Gleicheniaceae are apparently heliophilous. The furcate leaves with often strongly divergent forks anchor themselves as sprawling climbers
("Spreizklimmer") in the surrounding vegetation. The
long-creeping, freely branching stems permit much vegetative spreading, some species, e. g. Dicranopteris linearis, becoming actually weedy. Gleicheniaceae often
grow on mineral-poor substrates. The reason that they
are difficult to cultivate and thus rarely found in botanical gardens may be the fact that they are killed by
excessive mineral supply. A few species occur in bogs
and other open places on higher mountains.
Three of the four genera are pantropic; only Gleichenia s. str. is confined to the Old World. Diplopterygium has only a single neotropical species and is absent from Mrica. The greatest diversity of Diplopterygium and Dicranopteris is in Malesia. Continental
Africa is poor in species, and so is Madagascar. The
American representatives belong mostly to Sticherns.
AFFINITY. This is an old family. dating back to the early Mesozoic and perhaps even to the Carboniferous.
Holttum (1957 a) discussed some remote resemblance
to Schizaeaceae, Matoniaceae, and Cyatheaceae. The
family is rather isolated.
KARYOLOGY AND HYBRIDIZATION. Chromosome
counts are as yet available for rather few species only.
but they bear out the taxonomy as established on the
basis of morphological characters. The counts show
base numbers of 20 and 22 in Gleichenia, 56 in Diplopterygium. and 34 in Sticherus. The two subgenera of
Dicranopteris have different base numbers, 39 in subgenus Dicranopteris and 43 (44?) in subgen. Acropterygium; morphologically these are, however, not very
divergent.
There are rather few reports of hybridization in the
family: for Cuba by Duek (1974) and for Trinidad by
Jermy and Walker (1985).
SUBDIVISION. Neither Holttum (1959) nor Tryon and
Tryon (1982) proposed a formal subdivision. However,
it seems evident that Dicranopteris stands somewhat
147
apart in the subfamily, and the distinction of two
tribes, one for it and one for the other genera, seems
natural.
KEy TO THE GENERA.
1. Veins once forked or simple; sori consisting of 2-5 sporangia; indument of peltate, ciliate scales and fugacious stellate hairs
2
- Veins twice or more times forked; sori of c. 8 or more spor·
angia; scales none, indument of branched hairs
4. Dicranopteris
2. Branches of lamina flanking dormant apices of the highest
2. Sticherus
order pinnatifid
- Branches of lamina flanking dormant apices of the highest
order pinnate + pinnatifid
3
3. Ultimate segments about as long as wide, not opposite
(Fig. 76); costa flexuous, ill-defined; sori single on the seg·
ments
3. Gleichellia
- Ultimate segments opposite, much longer than wide; costa
distinct, straight; each segment with several sori
.
1. Diplopterygium
1. Diplopterygium Nakai
Diplopterygium Nakai, BulL Nat. Sci. Mus. Tokyo 29: 47
(1950).
Hicriopteris auett. non C. Presl, e. g., Ching (1940) Copeland
(1947).
Sticherus subgen. Hicriopteris sensu C. Chr. in Verdoorn,
Man. Pteridol: 530 (1938).
Gleichenia subgen. Diplopterygium (Nakai) Holttum, Reinwardtia 4: 261 (1957); Tryon & Tryon (1982).
Stem creeping, protostelic; leaves petiolate, scandent
or often erect, in mature plants with the rachis apex
dormant at a higher node than the first, often the second; indument as in the next genus but the scales often entire. Lowest leaflets of pinnae often reduced,
pinnate + pinnatifid, stipule-like, flanking the dormant
apex; semipenultimate divisions often broadly oblong,
pinnate; penultimate divisions narrowly oblong, deeply pectinately pinnatifid; ultimate divisions, soral and
sporangial characters as in the next genus; spores trilete, tetrahedral with prolonged angles; exospore with
a sheath-like structure extending from the aperture to
near the flange in the middle layer; surface levigate to
shallowly rugose, scarcely perforate.
About 25 species, from northern India, southern
China and southern Japan to Queensland and Polynesia (incl. Hawaii); not in Africa; a single species,
D. bancroftii (Hooker) A. R. Smith, in tropical America.
2. Sticherus C. Presl
Sticherus C. Presl, Tent. Pteridogr. 51 (1836).
Gleichenia subgenus Menensia Hooker, Spec. FiL 1: 4 (1844),
(not Menensia Willd. 1804, nec Roth 1797) e. g., Holttum
(1959); Tryon & Tryon (1982).
Gleichenia auctt., p. p. min.
Dicranopteris auett., p. p. min., e.g. Maxon (1909).
148
Gleicheniaceae . Gleichenioideae
Fig.76A-C. Gleicheniaceae. Gleichenia elongata. A Habit
(x 0.7). B Part of pinna (x 4). C Fertile segments from abaxial side (x 14) (Schelpe 1970)
Stem creeping, protostelic, bearing peltately attached,
lanceolate scales, glabrescent. Leaves usually scandent; primary rachis in mature plants none, the apex
of the petiole dormant beyond the first pinnae; all further rachis apices dormant beyond their first nodes;
young parts bearing peltate scales and branched hairs,
these deciduous or locally persistent. Pinnae paired
Gleichenia . Dicranopteris
("opposite"), equal or variously unequal, ± divergent.
Penultimate divisions deeply pectinately pinnatifid;
ultimate divisions narrowly oblong to tongue-shaped,
costate, with once-forked lateral veins, such segments
often also present on axes of penultimate or lower order. Dormant apices, often also costae, at least initially, clothed with long and narrow, fimbriate scales and
stellate hairs. Sori dorsal on the acroscopic branch of
each vein, round, with 2-5 large sporangia; sterile appendages among sporangia present or not. Spores
monolete, ellipsoidal; exospore as in Diplopterygium;
surface rugulose, usually perforate.
C. 80 species, pantropical and austral, very few in
Mrica and its islands.
Fig. 76
3. Gleichenia J. E. Smith
Gleichenia. J. E. Smith, Mem. Acad. Turin 5: 419 (1793), nom.
cons.; Holttum (1959).
Calymella C. Presl (1836); Nakai (1950).
Gleicheniastrum C. Presl (1848); Nakai (1950).
Stem creeping, protostelic; leaves of young and stunted plants with pinnate + pinnatifid lamina, without
dormant apices of axes, of well-developed plants with
a dormant apex at the first or second rachis node,
usually also on the first or second nodes of secondary
rachises; lamina often erect and non-scandent. Indument as in the last genus. Penultimate divisions narrowly oblong, deeply pinnatifid to subpinnate; accessory branches none (Fig. 76). Ultimate segments borne
only on the ultimate rachis branches, at least in well-
149
developed plants, shortly truncate.ovate to suborbicular, with a flexuous costa and a few simple or onceforked lateral veins, sometimes much excavate beneath
and almost pouch-like; sori single on the segments,
dorsal on the basal acroscopic veins, sometimes immersed in the leaf-tissue and covered by a flange of
the reflexed margin. Sporangia few per sorus. Spores
trilete, as in Diplopterygium.
About 10 paleotropical and austral species, absent
from the New World; from Mrica to Indo-China,
Malesia, Australia, and New Zealand.
4. DicrtlllOpteris Bemhardi
Figs. 74A, 75D-J, 77,
78A-C,79
Dicranopteris Bemhardi, Schrad. Joum. Bot. 1 (2): 38 (1806);
only in part of many other authors, e. g., Underwood
(1907); Maxon (1909).
Mertensia Willdenow (non Roth 1797), (not Gleichenia subgen. Mertensia Hooker).
Acropterygium (Diels) Nakai (1950).
Gleichenia auct!. div. p. p.
F'l!. 77 A-F. Gleicheniaceae. A, B Dicranopteris pectinata.
A Part of fertile leaf segment from abaxial side (x 4). B Sporangia in ventral and dorsal view (x 120). C-F Dicranopteris
linearis. C Tip of stem with unfolding leaf (x 0.6). D Apex
of developing lamina with crozier, pinna bases, and "stipules" (x 1.2). E Part of fertile segment from abaxial side
(x4). F Branched hair (much enlarged) (A,B from
G. M. Smith 1938; C-F from Holttum 1955)
B
F
150
Gleicheniaceae . Gleichenioideae . Stromatopteridoideae
~.:"
~
. E
Fig.7SA-Eo Gleicheniaceae. A-C Dicranopteris linearis.
A Habit (x 0.5). B Fertile segments from abaxial side (x 3).
C Sorus (xI5). D,E Stromatopteris moniliformis. D Habit,
with stem (x 0.7). E Fertile segment (x 4) (A-C from
Schelpe 1970; D,E from Brownlie 1969)
Stem creeping, protostelic or rarely solenostelic, bearing septate, at least basally branched hairs. Leaves
scandent, or erect in smaller species; lamina several
times, equally or unequally, pseudodichotomously
forked, the primary rachis sometimes continued between the forkings of lower order; often a pair of ac-
cessory, pectinately pinnatifid segments present at the
forks; penultimate branches sometimes so unequally
forked as to suggest an almost subbipinnate dissection
pattern. Dormant apices bearing rudimentary segments and branched hairs. Ultimate branches deeply
pectinately pinnatifid, the basal segments occasionally
enlarged and pinnatilobate, other ultimate segments
linear, rigid, costate, with subpinnately at least twice
forked veins. Sori· usually single on an acroscopic
branch of a secondary vein, with c. 8-15 sporangia not
mixed with sterile appendages; spores trilete and tetrahedral with prolonged angles (subgen. Dicranopteris)
or monolete and ellipsoidal (subgen. Acropterygium);
Dicranopteris - Stromatopteris
surface levigate, foveolate, low-rugulose, or rarely
granulate; otherwise as in Gleichenia.
About 12 species, pantropical and austral, most
strongly represented in Malesia. Two distinct subgenera, Dicranopteris (Fig.78, 79) with protostelic stem
and with accessory segments present at least at some
forkings; all but one species; subgenus Acropterygium
(Diels) Holttum with solenostelic stem and without accessory segments; only D.pectinata (Willd.) Underw.,
tropical America (Fig. 77).
In the past this genus was much more broadly circumscribed, e. g., by Underwood (1907), Maxon
(1909), etc., and contained all species with elongate ultimate segments. After preliminary work by Copeland
(1947) and Nakai (1950), Holttum (1957a, b) showed
that a genus so defined rests only on superficial characters. He proposed a more natural alignment of the
species and genera in the family.
Subfllmily Stromlltopterldoidelle Nakai
Fig. 78D, E
Stromatopteridoideae Nakai, Bull. Nat. Sci. Mus. Tokyo 29:
32 (1950).
Stromatopteridaceae (Nakai) Bierhorst, Phytomorphology 18
(2): 263 (1968).
Stem horizontal, protostelic, with vertical, slender, unequally dichotomous branches bearing the leaves;
young parts clothed in non-peltate scales; long, sim-
151
Flg.79. Gleicheniaceae. Dicranopteris linearis; Darjeeling, India. Phot K. U. Kramer
pie, bristle-like hairs present besides. Leaves several,
rigidly erect; petiole long, dark, subterete, adaxially
flattened-sulcate, with a single C-shaped bundle.
Lamina±erect, simply pinnate, with numerous, partly
overlapping, simple, ovate-rounded, entire or faintly
lobed, very coriaceous pinnae borne on and fully adnate to a dark, sulcate rachis, reduced and confluent at
the lamina apex; margin±revolute. Venation with illdefined, flexuous costa and free branches, anadromous. Sori usually one per pinna, occupying both
branches of the basal acroscopic veinlet, roundish, of
c. 15-20 rather large sporangia intermingled with
small, irregularly shapes scales. Sporangial stalk uniseriate; capsule large, with almost complete, oblique to
nearly transversal annulus bow and ill-defined stomium of smaller cells; annulus not interrupted at the
stalk. Spores monolete, ellipsoidal; exospore with
sheath-like structure, as in Gleichenia; perispore thin,
adherent to the irregular exospore; surface rugulose,
low ridges forming a nearly reticulate surface.
ANATOMV AND MORPHOLOGY. This was most fully described by Bierhorst (1968, 1969, 1971, 1973). He
(over)stressed the difficulty in separating the leaves, as
152
Gleicheniaceae . Stromatopteridoideae
lateral, appendicular organs, from the stem, as central
organ: as a similar condition prevails in the aerial
stems of Psilotaceae, he regarded the latter as true
ferns and Stromatopteris as a kind of connecting link;
but other characters are not at all in agreement with
this idea. The so-called non-appendicular nature of
the leaves of Stromatopteris was challenged by Kaplan
(1977). Stromatopteris is strongly mycorrhizal, like Psilotaceae and (less so) at least some Gleichenioideae
(Boullard 1958), and it seems reasonable to assume
that this has influenced the morphology to a certain
degree, resulting in simplification of vegetative parts.
GAMETOPHYTE. This was again well described and illustrated by Bierhorst (1968, 1971). It is subterranean,
cylindric, non-green, mycotrophic, bears rhizoids and
superficial gametangia, and is rather like the stem of
the sporophyte. The antheridia are large, many-celled,
and produce numerous spermatozoids. The archegonia are long-necked and straight or curved, variously
oriented.
ECOLOGY AND DISTRIBUTION. The sole representative
is confined to New Caledonia, where it is frequent in
serpentinic areas in the southern third of the island, in
open places and macchia-like vegetation (Brownlie
1969); Bierhorst (1968) also reported it from rainforests.
AFFINTY. The similarity to Psilotaceae, emphasized by
Bierhorst and regarded as indicative of affinity, was
very differently interpreted by many other authors,
notably Kaplan (1977; see also Kato 1983). When the
points of similarity are regarded as due to convergence
rather than to true affinity, one again has to turn towards Gleicheniaceae, which Bierhorst did not rule
out, either. The similarity in stelar structure, indument,
sporangial structure, spores, and gametangia can hardly all be fortuitous, and Nakai's (1950) and Holttum's
(1959) classification as a subfamily of Gleicheniaceae
is here reinstated. Phytochemistry also strongly supports this conclusion (Wallace et al. 1983).
A chromosome number of 2 n=c. (probably exactly) 39 was reported by Bierhorst (1968). The same basic number occurs in Dicranopteris subgenus Dicrano-
pteris.
A single genus:
5. Stromatopteris Mettenius
Stromatopteris Mellenius, Ann. Sci. Nat. IV 15: 84 (1861).
Characters of the subfamily.
Sole species S. moniliformis Mett. of New Caledonia.
Selected Bibliography
Bierhorst, D. W. 1968. On the Stromatopteridaceae (fam.
nov.) and on the Psilotaceae. PhytomofPhology 18:
232-268.
Bierhorst, D. W. 1969. On Stromatopteris and its ill-defined
organs. Amer. J. Bot. 56: 160-174.
Bierhorst, D. W. 1971 a. MOfPhology of vascular plants. New
York, London. Chapter 13: Stromatopteridaceae.
Bierhorst, D. 1971 b. Soral and pinnule ontogeny in Gleichenia linearis. Amer. J. Bot. 58: 417-423.
Bierhorst, D. W. 1973. Non-appendicular fronds in Filicales.
Bot. J. Linn. Soc. 67 Suppl. 1: 45-57.
Boullard, R. 1958. La mycotrophie chez les pteridophytes.
Bordeaux.
Brownlie, G. 1969. Flore de la Nouvelle-CaIOdonie et Dependances. 3. Pteridophytes. Paris.
Chrysler, M.A. 1943. The vascular structure of the leaf of
Gleichenia. I. The anatomy of the branching regions. Amer.
J. Bot. 30: 735-743.
Chrysler, M.A. 1944. Id. II. The petiolar bundle. Amer. J. Bot.
31: 483-491.
Duek, J. J. 1974. A newly recognized Gleichenia hybrid from
Cuba. Amer. Fern J. 64: 74-76.
Hagemann, w., Schulz, U. 1978. Wedelanlegung und Rbizomverzweigung bei einigen Gleicheniaceen. Bot. Jahrh.
Syst. 99: 380-399.
Holttum, R. E. 1957a. MOfPhology, growth habit and classification in the family G1eicheniaceae. PhytomofPhology 7:
168-184.
Holttum, R. E. 1957b. Florae Malesianae Praecursores XVI.
On the taxonomic subdivision of the Gleicheniaceae, etc.
Reinwardtia 4: 257-280.
Holttum, R. E. 1959. Gleicheniaceae. Flora Malesiana II. 1:
1-36.
Jermy, A. c., Walker, T. G. 1985. See General Referenes.
Kaplan, D. R. 1977. MOfPhological status of the shoot systems of Psilotaceae. Brittonia 29: 30-53.
Kato, M. 1983. The classification of major groups of pteridophytes. J. Fac. Sci. Univ. Tokyo III. 13: 263-283.
Lugardon, B. 1974. La structure fine de l'exospore et de la
perispore des filicinees isosporees. Pollen & Spores 16:
161-226.
Maxon, W. R. 1909. Gleicheniaceae. North Amer. Fl. 16 (1):
53-63.
Nakai, T. 1950. A new classification of Gleicheniales. Bull.
Nat. Sci. Mus. Tokyo 29: 1-71.
Nayar, B. K., Kaur, S. 1971. See General References.
Rauwenhoff, N. W. P. 1889. De geslachtsgeneratie der Gleicheniaceeen. Natuurk. Verh. Kon. Ned. Akad. Wetensch.
28: 1-54.
Stokey, A. G. 1950. The gametophyte of the Gleicheniaceae.
Bull. Torrey Bot. Club 77: 323-339.
Tryon, R. M., Tryon, A. F. 1982. See General References.
Underwood, L. M. 1907. A preliminary review of the North
American Gleicheniaceae. Bull. Torrey Bot. Club 34:
243-262.
Wagner, W. H. Jr. 1952. Types of foliar dichotomy in living
ferns. Amer. J. Bot. 39: 578-592.
Wagner, W. H. Jr. 1957. Heteroblastic leaf mOfPhology in
juvenile plants of Dicranopteris linearis (Gleicheniaceae).
PhytomofPhology 7: 1-6.
Wallace, J. W., Pozner, R. S., GOmez, L. D. 1983. A phytochemical approach to the Gleichenjaceae. Amer. J. Bot. 70:
207-211.
Filicatae . Grammitidaceae
Grammitidaceae
B.S. PARRIS
Grammitidaceae (c. Presl) Ching, Sunyatsenia 5: 264 (1940)
("Grammitaceae") based on: tribe Grammitideae (as
"Grammitaceae") C. Presl (1836).
Epiphytic, sometimes terrestrial or rupestral ferns,
with creeping to erect, simple or ± branched solenostelic stem usually bearing scales, occasionally glabrous. Petiole usually non-articulated to rhizome, terete, with a single vascular strand. Leaf usually bearing
unicellular or multicellular, occasionally glandular,
hairs which sometimes have unicellular or multicellular outgrowths. Lamina simple to tripinnate; veins free
or occasionally casually anastomosing without free included veinlets, sometimes ending in a thickened apex
or hydathode which may bear a white scale. Sori exindusiate, round or elongate, on the abaxial side of the
lamina or submarginal, sometimes sunken in laminar
tissue on the abaxial surface or at or near the margin,
sometimes protected by the lamina being folded over
them; receptacular appendages sometimes present;
sporangia many, sometimes bearing hairs near the annulus, maturing at different times; annulus longitudinal, bow of c. 7-20 cells; stalk usually of one row of
cells. Spores usually 64, sometimes 16 or 32 per sporangium, chlorophyllous, thin-walled, usually globose
to tetrahedral with trilete laesura, papillate with scattered spherical deposit, sometimes somewhat verrucate, somewhat echinate in Calymmodon.
ANATOMY AND MORPHOWGY. Very few studies have
been made on Grammitidaceae. Hayata (1928) illustrated the protostele of "Micropolypodium" pseudotrichomanoides. Van Cotthem (1970) found anomocytic
and polocytic stomata consistently throughout the
family, he studied all genera recognized below except
Acrosorus.
GAMETOPHYTE. This is slow-growing with a prolonged
filamentous phase giving rise to a spathulate or cordate plate, with a lobed apex, which is usually flat but
occasionally lobed and rumed. The filament tends to
break up and the fragments can then grow as independent gametophytes. The hair types already described
for the leaves may also occur on the gametophyte; for
a given species the hair type or types of the gametophyte also occur on the sporophyte, where additional
hair types may be present besides. The antheridia are
borne on filaments or plates, the archegonia on the
plate behind the apical notch; both antheridia and
archegonia are of the usual higher fern type. Spore
germination is usually directly by a filament, sometimes by a mass or plate which produces filaments; it
153
often takes place within the sporangium. Rhizoid development is delayed until the filament is at least three
cells long (Stokey and Atkinson 1958; Stone 1960).
ECOLOGY AND DISTRIBUTION. Most species occur in
tropical montane primary rainforest, but some grow in
tropical lowland primary rainforests and others are
found in lowland to montane temperate rainforest or
subalpine scrubland and alpine communities; usually
epiphytic, occasionally terrestrial, sometimes epilithic,
occasionally as facultative or obligate rheophytes. Few
species succeed in cultivation, even in botanical gardens, presumably because the habitat requirements for
nutrition, light, humidity, and air movement are precise and not easily duplicated.
One of the four genera is pantropic to (mostly
south-) temperate; the other three occur in Southeast
Asia and the Pacific.
AFFINITY. No fossils attributable to the family are
known except for Grammitis succinea described from
Oligocene amber deposits in the Dominican Republic
(Gomez 1982).
The Grammitidaceae have traditionally been regarded as Polypodiaceae (e. g., Christensen 1938;
Copeland 1947) or derived from them (Ching 1940).
Holttum (1947) pointed out the similarity of their stiff,
unicellular hairs on scales, leaves, and sporangia with
the hairs of the Thelypteridaceae, suggesting that they
may both have originated from Gleicheniaceae and
suggesting that the occurrence of leaves which rest
temporarily from apical growth in two species of
Grammitidaceae and in Gleicheniaceae may be significant. Nayar (1970) derived Grammitidaceae from
Cyatheoideae together with Aspleniaceae and Thelypteridaceae without discussing this decision. More recently, however, evidence has been presented that
Grammitidaceae and Polypodiaceae are indeed closely related (Price 1983; Wagner 1985). Jarrett (1980)
thought that both Grammitidaceae and Polypodiaceae
are related to the indusiate rather than the Gleichenioid ferns.
KARYOWGY AND HYBRIDIZATION. Relatively few chromosome counts are available for the family and all refer to Grammitis sensu lato. Most of these are of diploids, usually n=37 although n=32, 33, and 36 are also reported, but tetraploids, n= 74, are also ·known.
No counts are available for Acrosorus. Calymmodon.
and Scleroglossum.
All hybrids presently known or suspected in the
family are between simple-leaved and pinnate or pinnately lobed species, most of which would be placed
in different genera if Grammitis is treated in a narrow
sense, but this may reflect the ease of detection of such
morphologically distinct crosses rather than the scarcity of hybrids between species with a similar degree of
154
Filicatae • Grarnmitidaceae
leaf dissection. They are aenopteris curtisii (syn. Polypodium decrescens) x Grammitis fasciata (= G. x 10heriana) (Copeland 1952 b, 1%0), C. curtisii x G.loheriana (Copeland 1952 b), C. heterophyOa x G. billardieri (Parris 1977), C. heterophylla x G. mageOanica
subsp. nothofageti (Parris and Barnett, in prep.), C.longiceps x G. sumatrana (Parris 1984), G. subfasciata x
Xiphopteris conjunctisora ( = G. crenulata?) (Parris
1983), Adenophorus oahuensis x A. pinnatifidus
(Bishop 1974), G.jungermannioides x X. taenifolia?
(A. R. Smith, pers. comm.)
SUBDIVISION. Ching (1940) subdivided the family into
the tribe Grammitideae ("Grammitieae''), with sori
polypodioid, oblong or round, superficial or immersed, terminal on dorsal vein ends, containing
Grammitis, Calymmodon, Acrosorus, Prosaptia, and
Holcosorus, and the tribe Cochlidieae with sori coenosoroid, drymoglossoid, superficial, or sunk in grooves
parallel to the midvein, containing Cochlidium, ScleFig. 80 A-D. Grammitidaceae. A Grammitis curtisii, habit
(x 0.75). B Grammitis harrisii, leaf (x 0.75) and details of
lamina with sori (x 1.5). C,D Grammitis (., Cochlidium'~ seminuda. C Habit (x 0.75). D Detail of fertile part of lamina
(x 0.3) (A from Flora Taiwan 1975; B from Christensen
1936; C,D from Proctor 1977)
roglossum, Nematopteris, and Oreogrammitis. Holcosorus is now accepted as a member of the Polypodiaceae, the only species of Oreogrammitis is regarded as
a member of Grammitis (Parris 1983), while one species of Nematopteris is now included in Grammitis
(Copeland 1952 b; Parris 1983) and the other in Scleroglossum. As Cochlidium and Scleroglossum are more
closely related to genera treated as members of the
tribe Grammitideae than to each other, it seems best to
ignore Ching's treatment and to defer subdivision of
the family until the generic limits and relationships are
better known.
KEy TO THE GENERA.
1. Sori immersed in marginal or submarginal grooves, one on
each side of the midvein of the lamina but not contiguous
to it (Fig. 82 E, f)
4. Sc/eroglossum
- Sori superficial (sometimes protected in folds) or if immersed in the lamina, not in grooves one on each side of
the midvein of the lamina
2
2. Sori superficial or sunk in grooves or pouches in the lami1. Grammitis
na, not protected by folds
- Sori protected by folds of the lamina
3
3. Laminar fold protecting the sori opening towards the leaf
apex (Fig. 82 D); laminar texture membranous
2. Catymmodon
- Laminar fold protecting the sori opening towards the lower
surface of the leaf (Fig. 82 B); laminar texture coriaceous
3. Acrosorus
T
D
1. Grammitis Swartz
Figs. 80, 81
Grammitis Swam, Schrad. J. Bot. 1800': 3, 17 (1802); Cope-
land (1952 b); Morton (1967); Parris (1975, 1983, 1986);
Parris and Given (1976); Bishop (1977).
Cochlidium Kaulf. Ber!' Jahrb. Pharm. 21: 36 (1820); Christensen (1929); Bishop (1978).
Xiphopteris Kaulf. Ber!' Jahrb. Pharm. 21: 35 (1820); Copeland (1952 a).
Adenophorus Gaudich., Ann. Sci. Nat. 3: 508 (1824); Bishop
(1974).
Amphoradenium Desv., Prodr.: 335 (1827).
Prosaptia C. Presl. Tent. Pteridogr.: 165 (1836).
Ctenopteris Blume ex Kunze, Bot. Z. 4: 425 (1846); Copeland
(1956).
Austrogramme Fournier, Ann. Sci. Nat. Bol V. 18: 278 (1873).
p.p.
Glyphotaenium (1. Smith) J. Smith, Hist. Fi!.: 187 (1875).
Enterosora Baker. TImehri 5: 218 (1886).
Oreogrammitis Copeland. Philipp. J. Sci. C 12: 64 (1917).
Micropolypodium Hayata, Bot. Mag. Tokyo 42: 341 (1928).
Stem erect to long-creeping, scaly, the scales medium
to dark brown, sometimes iridescent and/or clathrate,
sometimes with hairs on the margin, occasionally
adaxially or abaxially. Petiole elongate or ± winged to
base. Lamina simple to tripinnate, lanceolate, oblanceolate, elliptic or linear in outline, occasionally dichotomously branched, membranous to coriaceous in
texture, occasionally with a dark sclerotic border, glabrous or with hairs of various lengths and types; simple, branched, acicular, glandular, catenate, sometimes
branches of the same hairs being of two different
types; veins free or anastomosing; sori usually on the
abaxial surface of the lamina and unprotected, occasionally sunken in pits in the lamina, occasionally ±
marginal in pouches in the lamina.
About 400 species in wet tropical, subtropical and
south-temperate regions, most numerous in southeast
Asia and tropical America.
The genus has sometimes been divided into eight
separate genera (Grammitis, Glyphotaenium, Cochlidium, Oreogrammitis, Xiphopteris, Ctenopteris, Adenophorus, and Prosaptia, e. g., Copeland 1947), which are
more often recognized in the Old World than in the
New, where the concept of Grammitis provisionally
adopted here is used and the constituent genera treated essentially as subgenera (e. g., Proctor 1985) or sections (e. g., Tryon and Tryon 1982). The treatment of
Grammitis here as a large and diverse genus reflects
our current lack of knowledge concerning relationships within the family. The presence of a number of
hybrids which would be considered intergeneric if
Grammitis were treated in the narrow sense weakens
the case for dividing the genus into genera, subgenera,
or sections along conventional lines. It may be possible, however, to draw up new generic boundaries in
the future using characters other than the classical
ones of leaf dissection to divide the bulk of the spe-
Fig. 81 A-F. Grammitidaceae. A, B Grammitis delitescens.
A Habit (x 2). B Detail offertile lamina (x 4). C,D Grammitis (.,Xiphopteris") serrulata. C Habit (x 1.5). 0 Apex of fertile lamina (x 5). E,F Grammitis (.,Prosaptia") contigua.
E Leaf (x 0.3). F TIp of fertile pinna (x 2.5)(A-D from Proctor 1985; E,F from Holttum 1955)
cies, and the classification of the Grammitidaceae into
four genera proposed here must be regarded as a temporary expedient pending further research. Although
the type species of Prosaptia and its close relatives may
appear distinct enough to warrant generic separation,
there is a gradation to the species group containing the
type species of Ctenopteris and from there to the species groups with links to Xiphopteris and Grammitis s.
str. The glandular receptacular appendages said to
characterize Adenophorus are found in other non-Hawaiian members of the family. The following three
small genera are kept separate because of their distinctive characters (see key) which do not intergrade with
those of Grammitis s.lat.
Filicatae . Grammitidaceae
156
..
3. Acrosorus Copel.
~~
Fig. 82 A, B
Acrosorus Copel., Philipp. J. Sci. 1 (Suppl. 2): 158 (1906).
Stem erect or ascending, bearing medium to dark red
brown scales, these glabrous except for an occasional
apical hair. Petiole ± winged to base. Lamina pinnately lobed to pinnate, linear in outline, coriaceous,
usually with short, stout, simple, or unilaterally
branched hairs < 0.5 mm long on rachis, margin or
around the sori; sori solitary on each pinna or lobe,
protected by ± equal folds of lamina on both sides,
sometimes sunken in laminar tissue.
About seven species, from Southeast Asia to Polynesia.
B
4. Scleroglossum v. A. v. R
Fig.82E,F
Scleroglossum v. A. v. R., Bull. Jard. Bot. Buit. 2, 7: 37 (1912);
Christensen (1929).
Nematopteris v. A. v. R. (1918); Christensen (1929).
~
F
Fig. 82 A-F. Grammitidaceae. A, B Acrosorus triangularis.
A Habit (x 0.37). B Part of lamina with sori (x 2.5). C,
D Calymmodon curtus. C Lamina (x 0.7). D Part of fertile
lamina (x 2). E, F Scleroglossum pusillum E Habit (x 0.5).
F Transection of fertile lamina (x 3) (Holttum 1955)
Stem erect or ascending, bearing medium brown
scales, these glabrous except for an occasional apical
hair. Petiole ± winged to base. Lamina linear to linear-spathulate, simple, coriaceous in texture, usually
with short, stout hairs ( < 0.5 mm long) on margin and
costa; sori in two rows, one each side of the costa,
near the margin to ± midway between margin and
costa, deeply sunken in grooves in the lamina.
About 8 species, from Ceylon through Southeast
Asia to Australia and Melanesia.
Scleroglossum superficially resembles some small
species of Vittaria. but can be distinguished easily by
its medium brown, rather than clathrate, scales, chlorophyllous spores and the absence of branched receptacular appendages characteristic of Vittaria.
Selected Bibliography
2. Ca/ymmodon C. Presl
Fig.82C,D
Calymmodon C. Presl, Tent. Pteridogr.: 203 (1836); Copeland
(1927).
Stem erect to ascending, bearing medium to dark
scales, these glabrous except for an occasional apical
hair. Petiole ± winged to base. Lamina pinnately
lobed to pinnate, linear in outline, membranous in texture, usually with short, slender, medium brown, simple, or unilaterally branched hairs < 0.5 mm long on
rachis and margin. Sori solitary on each pinna or lobe,
protected by a fold formed by the lower side of the
pinna or lobe.
About 30 species, from Ceylon through Southeast
Asia to Australia and Polynesia, with Borneo the centre of diversity.
Bishop, L. E. 1974. Revision of the genus Adenophorus
(Grammitidaceae). Brittonia 26: 217-240.
Bishop, L E. 1977. The American species of Grammitis Sect.
Grammitis. Amer. Fern J. 67: 101-106.
Bishop, L E. 1978. Revision of the genus Cochlidium (Grammitidaceae). Amer. Fern J. 68: 76-94.
Christensen, C. 1929. Taxonomic fern studies I. Cochlidium.
Dansk Bot. Ark. 6: 17-34.
Christensen, C. 1938. Filicinae. In: Verdoorn, F. (Ed) Manual
of pteridology. The Hague: Nijhoff, pp.522-550
Copeland, E. B. 1927. The genus Calymmodon. Philipp. J. Sci.
34: 259-269.
Copeland, E. B. 1952 a. The American species of Xiphopteris.
Amer. Fern J. 42: 41-52, 93-110.
Copeland, E. B. 1952 b. Grammitis. Philipp. J. Sci. 80: 93-276.
Copeland, E. B. 1956. Ctenopteris in America. Philipp. J. Sci.
84: 381-473.
Copeland, E. B. 1%0. Fern flora of the Philippines. Manila:
Bureau of Printing.
Filicatae . Hymenophyllaceae . Hymenophylloideae
Gomez, L D. Grammitis succinea, the first New World fern
found in amber. Amer. Fern J. 72: 49-52.
Hayata, B. 1928. On the systematic importance of the stelar
system in the Filicales III. Bot. Mag. (fokyo) 42: 334-348.
Jarrett, F. M. 1980. Studies in the classification of the leptosporangiate ferns: I. The affinities of the Polypodiaceae
sensu stricto and the Grammitidaceae. Kew Bull. 34:
825-833.
Morton, C. V. 1967. The genus Grammitis in Ecuador. Con·
!rib. U. S. Nat. Herb. 38 (3): 85-123.
Nayar, B. K. 1970. A phylogenetic classification of the homosporous ferns. Thxon 19: 229-236.
Parris, B.S. 1975. A revision of the genus Grammitis Sw. (Filicales: Grammitidaceae) in Australia. Bot. J. Linn. Soc. 70:
21-43.
Parris, B. S. 1977. A naturally occurring hybrid in Grammitidaceae (Filicales): Ctenopteris heterophylla x Grammitis billardieri. New Zeal. J. Bot. 15: 597-599.
Parris, B. S. 1983. A taxonomic revision of the genus Grammitis Swartz (Grammitidaceae: Filicales) in New Guinea.
Blume. 29: 13-222.
Parris, B. S. 1984. Another intergeneric hybrid in Grammitidaceae: Ctenopteris Jongiceps x Grammitis sumatrana. Fern
Gaz. 12 (6): 337-340.
Parris, B. S. 1986. Grammitidaceae of Peninsular Malaysia
and Singapore. Kew Bull. 41: 491-517.
Parris, B. S., Given, D. R. 1976. A taxonomic revision of the
genus Grammitis Sw. (Grammitidaceae: Filicales) in New
Zealand. New Zeal. J. Bot. 14: 85-111.
Price, M. G. 1983. Pecluma, a new tropical American fern genus. Amer. Fern J. 73: 109-116.
Proctor, G. R. 1985. Ferns of Jamaic•. London: British Museum (Natural History).
Stokey, A. G., Atkinson, L R. 1958. The gametophyte of the
Grammitidaceae. Phytomorphology 8: 391-403.
Stone, I. G. 1960. The gametophytes of Grammitis billardieri
and Ctenopteris heterophylla. Aust. J. Bot. 8: 11-37.
Wagner, F. S. 1985. Bilateral spores in New World grammitid
ferns. Amer. Fern J. 75: 6-11.
Hymenophyllaceae
K. IWATSUKI
Hymenophyllaceae Link, Handb. z. Erk. d. Gew. 3: 36
(1833); Copeland (1938); Taton (1946); TIndale (1963);
Sledge (1968); Croxall (1975); Lellinger (1984); Iwatsuki
(1985).
Trichoman.ceae Kunkel, Feddes Rep. 70: 155 (1965).
Epiphytic, saxicolous, or terrestrial ferns of small to
medium size; stem creeping, or less often erect, with a
so-called Hymenophyllacean protostele, bearing only
hairs; petiole non-articulated, terete, commonly with
one leaf trace; lamina simple to pinnately decompound, or flabellate, digitate, dichotomous, or even irregularly divided; segments one-veined, unistratose, or
rarely with two to four cell layers without intercellular
spaces and stomata; sori terminal on the veins, soli-
157
tary, at the apex of the ultimate segments, or marginal
on simple to pinnatifid leaves; involucres cup-shaped
to deeply 2-c1eft nearly to the base; receptacles terminating a vein, short, capitate or clavate, or long and exserted; sporangia short-stalked to (sub)sessile, maturing basipetally; annulus oblique, not interrupted;
dehiscence irregular; spores globose-trilete, tetrahedral, containing chloroplasts, usually short-lived.
Two Subfamilies:
1. Leaves simple to pinnately decompound, if simple and reniform, not large nor long-stalked; lamina unistratose except
for the veins, or at most of 3 cell layers. Hymenophylloideae
- Leaves simple, reniform, large, commonly over 5 em diam.,
long-stipitate, petioles over 7 em long; lamina mostly of 4
cell layers
Cardiomanoideae
Subfamily Hymenopbylloideae
Stem creeping, ascending, or erect. Petioles very short
to elongate, rarely longer than laminas. Lamina simple
to pinnately decompound, or flabellate, forked, digitate, dichotomous, or even irregularly divided, from
less than 2 mm when mature to over 40 cm long; leaf
architecture anadromous or catadromous. Veins free,
rarely somewhat anastomosing. In pinnate laminas the
ultimate segments linear, one-veined, membranous to
herbaceous; ultimate segments or lobes entire or with
teeth that are usually independent from vein ends, or
each tooth with a vein end (in Hymenoglossum, Crepidomanes sect. Gonocormus, and Cephalomanes subgen. Cephalomanes). Sori at the ends of veinlets, at the
apices of ultimate segments, or on the margin oflobes;
involucres cup-shaped with truncate, dilated or bilabiate mouth, or variously bivalvately cleft, up to the
base; receptacles short, capitate to clavate, or elongate
and long-exserted. Sporangia maturing basipetally,
short-stalked. Spores often starting cell division in the
sporangium. Gametophytes filamentous or ribbonshaped.
ANATOMY AND MORPHOLOGY. Fundamental observations on anatomical characters were published by
Boodle (1900); additions were giyen by many botanists
(see, e. g., Ogura 1972). In most species the roots are
weakly deyeloped and simple in internal structure; the
stele is monarch or diarch, with one to seyen protoxylem groups in each strand. There are species that lack
roots; they must haye lost them in the COurse of evolution. The stem is creeping or erect; in Hymenoglossum,
Serpyllopsis, Hymenophyllum, and Sphaerocionium filiform or wiry; in the other genera erect, or, if creeping,
thicker or densely coyered with black hairs. The stele
is a so-called Hymenophyllaceae-type of protostele;
three Iypes were proposed by Ogura (1972). The petiole is lerete, often winged throughout, bul the wings
are often lost with age. Three types of internal slruc-
158
Hymenophyllaceae • Hymenophylloideae
ture of the petioles were described by Nozu (1950) according to the arrangement of the vascular elements;
generally the leaf trace is single and its structure is
simplified to some extent.
The lamina is mostly unistratose, except in the veins.
There are species with 2-4 cell layers, but even then
there are neither intercellular spaces nor stomata.
False veinlets are known in some species; morphologically they are veinlets without tracheids (Wessels Boer
1%2; Iwatsuki 1978). The venation pattern is anadromous in most representatives, catadromous in Trichomanes (s. str.), but in leaves of simple structure this is
hard to observe. In accordance with the venation pattern the sori are epitactic or paratactic in position
(Prantl 1875), i. e., in determinate leaves a proximal
veinlet mayor may not also be terminated by a sorus
(epitactic), or the sorus is borne on the tip of a proximal veinlet, so that a distal vein is free to continue
growth and branch further (paratactic). The sori are
terminal on the veinlets, solitary at the apex of an ultimate segment, or placed along the margin of lobes.
The involucres are cup-shaped with truncate or dilated
mouth, obconic with bilabiate mouth, or bilabiately
cleft nearly to base. Histological and developmental
comparison shows that these variations appear only in
the course of the ontogeny of the involucres (Iwatsuki
1977).
GAMETOPHYTE. The spores of Hymenophyllaceae contain chloroplasts and start to divide within the spore
coat. Accordingly, they are short-lived. The development of the prothallium, however, is slow, often taking
a few years until it is fully mature. It is filamentous or
ribbon-like; the ribbons are,green, unistratose, and attain 2 em in length. The early development was described by Stokey (1940, 1948) and Atkinson (1960).
Yoroi (1972) described the development and morphology of the gametophytes of 14 Japanese species. The
distribution of filamentous and ribbon-like prothallia
is in general accordance with Copeland's system of
generic classification, at least in the species where it is
known. If, however, other important features are taken
into consideration, the variation is greater. The diversification of the gametophytes in this family may be related to ecological adaptations.
ECOLOGY AND DISTRIBUTION. The Hymenophyllaceae
occur predominantly in mossy forest in tropical-montane and south-temperate areas. Most species are epiphytes and grow on moss-covered tree trunks. Many
terrestrial species are also concentrated in mossy forests. The filmy habit is apparently closely related to
the moist air in which these ferns grow. When the air
becomes drier the filmy ferns curl up as if dead but
upon moistening the leaves unfold again, as bryophytes do (e. g., Komas 1977). There are species growing near streams in lowlands, up to middle elevation.
Most species of Cephalomanes subgen. Cephalomanes
seem to be rheophytes. Some species appear to have
xerophytic features (Sphaerocionium subgen. Apteropteris), but they, too, are limited to areas with high rainfall. The evolution of the filmy ferns shows a tendency
towards progressive reduction in size, which may be
considered as adaptive to the extremely moist mossy
zone, again a parallel with bryophytes. The species assortment is richest in the tropics and in south-temperate areas. Among seven genera recognized here, two
are monotypic and confined to Chile and subantarctic
South America, respectively. Sphaerocionium and Trichomanes are centred in the New World but also represented in the Old World; for Cephalomanes the reverse is true and the same holds for Crepidomanes.
Hymenophynum occurs throughout the area of the
family. Although Copeland (1939) suggested an Antarctic origin for the family, it may be safely assumed
that the present distribution of the Hymenophyllaceae
arose in the tropics and that they subsequently dispersed from there (Iwatsuki 1979).
In eastern North America vegetatively reproducing
gametophytes of Trichomanes have been found far to
the north of where sporophytes are known (Farrar
1967,1985; Farrar et al. 1983).
AFFINITY. Fossil records are scarce, as is to be expected from the filmy habit. The extremely reduced morphology is interpreted here as derived; the family may
be supposed to have evolved with the apparition of
dense tropical forests. It is usually placed in the "Marginales," often next to Schizaeaceae. The involucres
are often considered as peculiar structures, but except
for their unistratose tissues they much resemble those
of the Dicksoniaceae (Iwatsuki 1977). The Hymenophyllaceae may safely be placed in the series "Marginales," probably as a small offshoot adapted to a humid, warm climate.
KARYOWGY AND HYBRIDIZATION. Chromosome
counts are available for more than 120 species (Love et
al. 1977), slightly less than one-fifth of the supposed
overall number of species. The commonest number is
2n= 72, distributed among almost all subgroups, as far
as known. The information on chromosome numbers
is still insufficient, and there are divergent numbers
even within a single taxon. Tatuno and Takei (1969) attempted a karyotype analysis of the Japanese species,
but the results were limited, as the morphology of only
some of the chromosomes could be observed. Braithwaite (1975) discussed a case of difference in karyology between groups with the same chromosome number. There are suggestions that some particular forms
may be natural hybrids (e.g., Bierhorst 1974), but the
evidence is nowhere conclusive. There are no experimental observations on the occurrence of self-fertilization vs. cross-fertilization, and the likelihood for the
Hymenoglossum • Serpyllopsis. Hymenophyllum
formation of hybrids cannot be assessed. A few cases
of apogamy are known in Trichomanes s.l., e. g., in
T. pinnatum Hedw. (Bierhorst 1975) and in T. prolifernm Bl. (T. minutum BI.)(BeIl 1960; Yoroi and Iwatsuki
1977).
Phytochemistry. Markham and Wallace (1980) investigated the variation of flavonoids in the family. They
pointed out their diversity, but the taxonomic implications were not quite clear. There was a resemblance in
flavonoid pattern to the Marsileaceae.
SUBDIVISION. Traditionally two genera were recognized: Hymenophyllum. with bivalved involucres, and
Trichomanes. with cup-shaped to tubular or infundibuliform involucres (e. g., Christensen 1906, 1938; Morton 1968). These are, however, heterogeneous, and
their delimitation becomes impossible when all species
are taken into account. Copeland proposed a new generic system with 34 genera (1938, 1947), but this involves too much splitting and is inconvenient for the
non-specialist. Recently, a system with seven genera in
the Hymenophylloideae, besides the monotypic Cardiomanioideae, was proposed (Iwatsuki 1984).
KEy TO THE GENERA OF THE HYMENOPHYLLOIDEAE.
1. Lamina simple. over 5 em long, long-stipitate, the marginal
cells in two rows, forming a dark border (Chile)
1. Hymenoglossum
- Lamina pinnatifid to pinnately decompound, digitate, or
flabellate, rarely simple, then less than 5 em long and not
2
long-stipitate
2. Leaves minute, moss-like, simply pinnate with entire, leaflike pinnae; petioles ill-dermed from the stem (Fig. 83 A, B)
(southernmost South America)
2. SerpyfJopsis
- Leaves otherwise
3
3. Dissection/venation catadromous or lamina simple; sori
epitactic; involucres tubular, truncate at apex, or with dilated lips
6. Trichomanes
159
Stem long-creeping, wiry, irregularly bearing wiry
roots; petioles longer than laminas, terete, wingless,
naked. Lamina simple, large, about 10 cm long, subentire to crenate, rhomboid or broadly lanceolate, acute,
with cuneate base, glabrous; veins (sub)opposite, simple, parallel, reaching the margin; lamina unistratose
with a broad marginal band of two cell layers. Sori
marginal, semi-immersed; involucre bivalvate, the immersed portion cup-shaped; receptacle clavate to capitate, included. Spores spheroidal, papillate, with quite
dense, low prominences.
A single species, H. crnentum (Cav.) C. Presl, in
Chile and Juan Fernandez.
2. SerpyUopsis van den Bosch
Fig. 83 A, B
SerpyUopsis van den Bosch, VersL Meded. KoninkL Akad.
Wetensch. Afd. Natuurkd. 11: 318 (1861); Christensen
(1910); Copeland (1938, 1947); Morton (1968).
Stem long-creeping, wiry, irregularly bearing roots;
petioles proportionately short, terete, sparsely hairy.
Lamina pinnate, indeterminate, lanceolate, subacute to
rounded, the base rounded to broadly cuneate, about
5 em long; pinnae simple, entire, narrowly ovoid, subacute, one-veined, unistratose; hairs on rachis and
veins beneath reddish-brown, thick-walled, multicellular. Sori solitary at the ends of veins, on the distal margin of the pinnae; involucres immersed, cup-shaped,
with narrow lips protruding from the pinna margin;
receptacles long, exserted. Spores spheroidal, densely
echinate with many blunt projections.
A single species, S. caespitosa (Gaud.) C. Chr., in
southernmost South America, the Falkland Islands,
and Juan Fernandez. Some infraspecific taxa have
been described.
- Dissection/venation anadromous: lamina rarely simple
(sometimes in Crepidomanes)
4
3. Hymenophyllum J. E. Smith
4. Leaves bearing stellate hairs or long, pale hairs, or at least
leaves hairy on the basal part of the petioles and on young
leaves with long, pale, often forked hairs 4. Sphaerocionium
- Leaves lacking stellate hairs and long, pale, simple or
forked hairs
5
5. Involucres bivalvate to the base or to the middle, or, if conical, with bilabiate mouth and serrate margin of lobes, or
with thick cell walls and brown hairs; false veinlets wanting
3. HymenophyfJom
- Involucre tubular or conical, with truncate or dilated
mouth, or with two lips occupying about half the length of
the involucres; if the mouth of the involucre is bilabiate,
Hymenophyllum J. E. Smith, Mem. Acad. Turin 5: 418 (1793);
Copeland (1938, 1947); Morton (1968), p. p.
Meringium C. Presl (1843).
Leptocionium C. Presl (1843).
Hemicyatheon (Domin) Copeland (1938).
Buesia (Morton) Copeland (1938).
Amphipterum (CopeL) Copeland (1938).
Myriodon (CopeL) Copeland (1938).
Craspedophyllum (c. Presl) Copeland (1938).
Mecodium (CopeL) Copeland (1938).
Rosenstockia Copeland (1947); Morton (1968).
the segments possess false veinlets
Fig. 83 C-E
6
6. Stem long-creeping or short but slender; plants epiphytic,
saxicolous, or rarely terrestrial
5. Crepidomanes
- Stem erect, or, if creeping, short and thick; terrestrial or
saxicolous plants
7. CephaIomanes
1. Hymenoglossum C. Presl
Hymenoglossum C.Presl, Abh. bohm. Ges. V.3: 35 (1843);
Copeland (1938, 1947); Morton (1968).
Stem long-creeping, wiry, bearing sparse, brownish
hairs; petioles distinct, terete, winged or not. Lamina
pinnately divided, pinnate and pinnatifid to quadripinnate; rachis like the upper part of the petiole, naked
or hairy. Pinnae alternate; in some species wings present on rachis, pinna-rachis, and veins, in others scalelike accessory lobes on rachis and pinna-rachis beneath. Ultimate segments one-veined, entire or dentic-
160
Hymenophyllaceae • Hymenophylloideae
Fig. 83 A-I. Hymenophyllaceae. A, B Serpyllopsis caespitosa.
A Habit (x 1). B Sori (x 10). C-E Hymenophyllum javanicum. C Habit (x 1). D Pinna (x3). E Sori (x 12). F-H Crepidomanes paucinervium. F Habit (x 1). G Pinna (x 4).
H Sorns with the anterior part of the involucre cut away
(x 14). 1 Cephalomanes cumingii, fertile segment with sorus
(x 15) (A, B from Hooker 1846; C-E from DeVol 1970;
F-H from Flora 'Thinlingensis 1974; 1 from Flora Taiwan I,
1975)
ulate, no relation between teeth and veins. Lamina unistratose except for the veins. Sori solitary at the apices
of the ultimate segments; involucres bivalved, cleft
nearly to the base, or the basal portion immersed in
the segments and cup-shaped, with bivalved lips; receptacles capitate, clavate, or filiform, included to
long-exserted. Spores spheroidal, with very thin perispore and low-papillate, rarely echinate surface.
Nearly 250 species, throughout the tropics and
south-temperate areas, a few extending to north-temperate areas in oceanic regions.
Hymenophyllum is here construed in a broader
sense, excluding only Sphaerocionium. The soral structure is in some species similar to that in some Crepidomanes species. The vegetative structure is simple yet
variable, and several subgenera can easily be recognized.
4. Sphoerodonium C. Presl
Sphaerocionium C. Presl, Abh. BOhm. Ges. V. 3: 33 (1843);
Copeland (1933, 1947); Iwatsuki (1982).
HymenophylJum sect. Sphaerocionium Morton (1947, 1968).
Sphaerocionium· Crepidomanes. Trichomanes· Cephalomanes
Apteropteris Copeland (1938).
Microtrichomanes (Mett. ex Prantl) Copeland (1938), p. p.
Stem long-creeping, filiform, bearing rather sparse,
brownish hairs, these simple, forming a downy indument. Petioles distinct, remote, terete, usually hairy,
the hairs like those on the lamina. Lamina pinnately
compound, pinnate and pinnatifid to bipinnate and
tripinnatifid, or in dwarfed species seemingly digitate,
of medium size: rachis like the upper part of the petiole, or very short and winged, usually hairy. Lamina
of one cell layer except for the veins. Ultimate segments one-veined, entire, usually hairy; hairs on leaves
simple, forked or stellate, when simple usually long,
pale brown, forming a downy indument; in some species accessory wings with lamina-like structure present
on axes. Laminar cells regularly arranged; internal
walls thin, straight. Sori solitary on the tips of the ultimate segments; involucres 2-valved, deeply cleft nearly to the base, or basally immersed in the segments
and forming a shallow cup with bilabiate mouth; receptacles filiform or clavate, included or shortly protruding. Spores spheroidal, with low, usually rounded,
papillate projections. About SO species, in both hemispheres but predominantly neotropical.
5. Crepidomanes C. Presl
Fig.S3 F-H
Crepidomanes (C. Presl) C. Presl, Epimel. Bot.: 258 (1851),
based on Trichomanes subgen. Crepidomanes C. Presl, ibid.:
17; Copeland (1938,1947).
Trichomanes L, auct. pp.
Pleuromanes (c. Presl) C. Presl (1849).
Gonocormus van den Bosch (1861).
Vandenb08chia Copeland (1938).
Microtrichomanes (Mett. ex Prantl) Copeland (1938).
Reediella Pichi Sermolli (1970).
Stem long-creeping, slender, bearing more or less
dense, blackish hairs. Petioles distinct, terete, winged
or not. Lamina flabellate or digitate to pinnately divided, then uni- to quadripinnate. Rachis like the
upper part of the petiole, rarely very shortly and
broadly winged like the segments. Lamina generally of
one cell layer except for the veins; margin entire; ultimate segments glabrous or setiferous; in subgenus
Crepidomanes false veins present, these submarginal
and oblique; in subgenus Crepidium the marginal cells
differentiated; in subgenus Pleuromanes lamina unistratose except for the veins and margin of segments,
the internal cell walls usually thin and straight. Sori
solitary on the tips of the ultimate segments; involucres cup-shaped with subtruncate, dilated, or bilabiate
mouth, in subgenus Crepidomanes the lips separated
for one-third of the length of the involucres; receptacles long, exserted. Spores spheroidal, with. thin perispore and low, dense papillae.
Nearly 120 species, mostly palaeotropical, some
neotropical.
161
6. TrkhotnlllU1S L.
TrichomanesL, Spec. Plant. 2: 1097 (1753); Copeland (1938,
1947); Morton (1968), p.p.
Didymogl08sum Desvaux (1827).
Microgonium C. Presl (1843).
Feea Bory (1824).
Lecanolepis Pichi Sermolli (1973).
Stem short, ascending, or long-creeping, in some species rootless (subgen. Didymoglossum), usually bearing
dense, blackish hairs; petioles obsolete to distinct,
terete. Lamina simple, irregularly divided, pinnatifid,
or simply pinnate, of very small to medium size; rachis
obsolete or distinct. Pinnae of pinnate leaves and lobes
of simple leaves entire or lobed, one-veined, manyveined, or veinless, uni- or rarely bistratose (in fertile
leaves of sect. Feea). False veinlets present (subgen.
Didymoglossum), oblique or submarginal. Marginal
hairs (sect. Didymoglossum) and scale-like accessoary
marginal emergences (sect. Lecanium) sometimes present. Venation if pinnate catadromous. Sori epitactic,
often immersed in the margin of the lamina: involucres campanulate or cup-shaped, the mouth dilated or
truncate; receptacles long, exserted. Spores spheroidal; presence ofperispore uncertain; surface shallowly
papillate or echinate.
Nearly SO species, most of them in the neotropics, a
few in the Old World.
7. Cepludomanes C. Presl
Fig.S3 I
Cephalomanes C.PresI, Abh. Biihm. Ges. Y.3: 17 (1843):
Capel. (1933, 1947).
Abrodictyum C. Presl (1843).
Davalliopsis van den Bosch (1863).
Macroglena (C. Presl) Capel. (1938, 1947).
Selenodesmium (Prantl) Capel. (1938, 1947).
CallistopleTis Capel. (1938, 1947).
NesopleTis Capel. (1938, 1947).
Stem short, erect, or creeping, relatively thick, bearing
numerous wiry roots and lustrous, multicellular hairs.
Petioles distinct, terete, usually narrowly winged, often
densely hairy. Lamina simply pinnate to elaborately
pinnately decompound, commonly comparatively
large, often c. 10-15 cm long; rachis like the upper
part of the petiole, usually winged. Venation anadromous. Ultimate segments one-veined, unistratose except for the veins, glabrous, or rarely hairy on the
veins. Lamina cells of a special type and specially arranged in some species, often with thick and coarsely
pitted internal walls, or the expanded laminar parts extremey reduced, of one row of cells on each side of a
vein. Sori usually at the tips of the ultimate segments,
paratactic. Involucres cup-shaped or campanulate, the
mouth truncate or dilated; receptacles usually very
long, exserted. Spores spheroidal, with short spines.
About 60 species, throughout the tropics but concentrated in the Old World.
162
Hymenophyllaceae . Cardiomanoideae
Subfamily Cardiomanoideae
Stem long-creeping, comparatively thick. Petioles remote, elongate, more than three times as long as the
lamina. Lamina simple, reniform, subentire or sinuatecrispate in the soriferous region, relatively large, commonly over 5 cm in diam., chartaceous, of 4 cell layers; venation dichotomous, repeatedly forked; veins
free, with slightly thickened ends just behind the sclerotic margin. Sori terminal on·the veins, sometimes occupying all vein ends but often confined to the central
portion of the lamina; involucres immersed, campanulate with truncate mouth; receptacles long, exserted.
Sporangia as for the family; spores spheroidal, with
thin, granulate perispore, surface coarsely low-papillate.
ANATOMY AND MORPHOLOGV. Roots adventitious on
the stem, wiry, with monarch to triarch stele; branching scarce and irregular. Stem stele of the so-called
Hymenophyllaceous protostelic type. Rhizoids rather
densely borne on the roots, the stem bearing sparse articulated hairs. Petioles terete, winged at least above,
usually naked, erect, well-developed. Lamina distinctive in its reniform shape with deeply cordate base and
usually fringed by sori. Although there are usually
four layers of cells, in which this genus is very distinctive in the family, there are no intercellular spaces.
GAMETOPHVTE AND EMBRVOWGV. The fundamental
observations were made by Holloway (1930, 1944).
The gametophyte is ribbon-like, at the base irregularly
branched, each ribbon attaining a length of 10-12 mm,
growing very slowly and reaching its definitive size only after c. 6.5 years. The lobes of the prothalIium are
green and unistratose except for the thicker base and
cushions. The archegonia have straight necks; the antheridia are comparatively large and contain a large
number of spermatocytes; there is no cap cell and a
large number of wall cells. The first or basal partition
wall of the embryo is more or less transverse to the
longitudinal axis of the archegonium. The epibasal
and hypobasal cells divide next to form a 4-celled embryo. The following divisions do not form a fixed pattern; the first leaf and other organs are differentiated
relatively late.
ECOLOGV AND DISTRIBUTION. The sole species of the
only genus, C. reniforme (Forst.) C. Presl, is endemic to
New Zealand, where it occurs in moist forests at low
elevation, often as an epiphyte not too far from the
ground, but also terrestrial, especially after clearing of
the forest (Holloway 1923).
CYTOWGY. 2n=72 is known as the chromosome number. As it is common in many other genera of the family, it is not indicative of the relationship within the
family.
PHYTOCHEMISTRY. Wallace and Markham (1978) investigated the flavonoids and compared them with those
of other primitive ferns.
A single genus.
8. Cardiomanes C. Presl
Cardiomanes C. Presl, Abh. SOhm. Ges. V. 3: 12 (1843).
Monotypic; see above.
Selected Bibliography
Atkinson, L. R. 1960. A new germination pattern for the Hymenophyllaceae. Phytomorphology 10: 26-36.
Bell, P. R. 1960. The morphology and cytology of sporogenesis of 1Hchomanes proliferum BI. New Phytol. 59: 53-59.
Bierhorst, D. W. 1974. A peculiar new species of Trichomanes
sect. Selenodesmium and several of its probable hybrids.
Amer. Fern J. 64: 18-26.
Bierhorst, D. W. 1975. The apogamous life cycle of 1Hchomanes pinna/um - a confirmation of Klekowski's predictions on homoeologous pairing. Amer. J. Bot. 62: 448-456.
Boodle, L. A. 1900. Anatomy of Hymenophyllaceae, Schizaeaceae and Gleicheniaceae. Ann. Bot. (London) 14:
455-496.
Braithwaite, A. F. 1975. Cytotaxonomic observations on some
Hymenophyllaceae from the New Hebrides, Fiji and New
Caledonia. Bot. J. Linn. Soc. 71: 167-189.
Christensen, C. 1910. On some species of ferns collected by
Dr. Carl Skottsberg in temperate South America. Ark. fOr
Bot. 10 (2): 1-32.
Christensen, C. 1938. Filicinae. In: Verdoorn, F. (Ed) Manual
of pteridology. The Hague: Nijhoff, pp 522-550.
Copeland, E. B. 1933. Trichomanes. Philipp. J. Sci. 51:
119-280.
Copeland, E. B. 1937. Hymenophyllum. Philipp. J. Sci. 64:
1-188.
Copeland, E. B. 1938. Genera Hymenophyllacearum. Philipp.
J. Sci. 67: 1-110.
Copeland, E. B. 1939. Fern evolution in Antarctica. Philipp. J.
Sci. 70: 157-189.
Croxall, J. P. 1975. The Hymenophyllaceae of Queensland.
Aust. J. Bot. 23: 509-547.
Farrar, D. R 1967. Gametophytes of four tropical fern genera
reproducing independently of their sporophytes in the
southern Appalachians. Science 155: 1266-1267.
Farrar, D. R 1985. Independent fern gametophytes in the
wild. In: Dyer, A. F., Page, C. N. (Eds) Biology of Pteridophyta. Proc. R Soc. Edinburgh Ser B 86: 361-369.
Farrar, D. R., Parks, 1. C., McAlpin, B. W. 1983. The fern genera Vitlaria and 1Hchomanes in the northeastern United
States. Rhodora 85: 83-91.
Holloway, J. E. 1923. Studies in the New Zealand Hymenophyllaceae Part I. The distribution of the species in Westland and their growth forms. Trans. N. Zeal. Inst. 54:
577-618.
Holloway, J. E. 1930. The experimental cultivation of the
gametophytes of HymenophyHum pulcherrimum, Col. and
of Trichomanes reniform., Forst. f. Ann. Bot. (London) 44:
269-284.
Holloway, 1. E. 1944. The gametophyte, embryo and developing sporophyte of Cardiomanes reniforme (Forst.) Presl.
Trans. R Soc. N. Zeal. 74: 196-206.
Filicatae . Hymenophyllopsidaceae
Iwatsuki, K. 1977. Studies in the systematics of filmy ferns
III. An observation on the involucres. Bot. Mag. (Tokyo)
90: 259-267.
Iwatsuki, K. 1978. Id. IV. Notes on the species with false
veinlets. Mem. Coli. Sci. Univ. Kyoto B 7: 31-43.
Iwatsuki, K. 1979. Distribution of the filmy ferns in palaeotropics. In: Larsen, K., Holm-Nielsen, L. B. (Eds) Tropical
botany. London: Academic Press.
Iwatsuki, K. 1982. Studies in the systematics of filmy ferns
VI. The genus Sphaerocionium in Asia and Oceania. J. Fac.
Sci. Univ. Tokyo III. 13: 203-215.
Iwatsuki, K. 1984. Id. VII. A scheme of classification based
chiefly on the Asiatic species. Acta Phytotax. Geobot. 35:
165-179.
Iwatsuki, K. 1985. The Hymenophyllaceae of Asia, excluding
Malesia. J. Fac. Sci. Univ. Tokyo III. 13: 501-551.
KomaS, J. 1977. Ufe-forms and seasonal patterns in the pteridophytes in Zambia. Acta Soc. Bot. Pol. 46: 669-690.
Lellinger, D. B. 1984. Hymenophyllaceae, in: B. Maguire &
collab., Botany of the Guayana Highland XII. Mem. New
York Bot. Gard. 38: 9-46.
LOve, A.., LOve, D., Pichi Sermolli, R. E. G. 1977. Cytotaxonomical atlas of the Pteridophyta. Vaduz: Cramer.
Markham, K. R., Wallace, J. W. 1980. C-glycoxylxanthone
and flavonoid variation within the filmy ferns (Hymenophyllaceae). Phytochemistry 19: 415-420.
Morton, C. V. 1947. The American species of Hymenophyllum
section Sphaerocionium. Contrib. U. S. Nat. Herb. 29 (3):
139-201.
Morton, C. V. 1968. The genera, subgenera, and sections of
the Hymenophyllaceae. Contrib. U. S. Nat. Herb. 38 (5):
153-214.
Nozu, Y. 1950. Morphological observation of the Hymenophyllaceae from Japan I-II. Bot. Mag. (Tokyo) 63: 71-76;
132-138.
Prantl, K. 1875. Untersuchungen zur Morphologie der
GefaBkryptogamen I. Leipzig: W. Engelmann.
Sledge, W. A. 1968. The Hymenophyllaceae of Ceylon. Bot. J.
Unn. Soc. 60: 289-308.
Stokey, A. G. 1940. Spore germination and vegetative stages
of the gametophytes of Hymenophyllum and Trichomanes.
Bot Gaz. 101: 759-790.
Stokey, A. G. 1948. Reproductive structures of the gametophytes of Hymenophyllum and Trichomanes. Bot. Gaz. 109:
363-380.
Taton, A. 1946. Revision des Hymimophyllacees du Congo
Beige. Bull. Soc. R. Bot. Belg. 78: 5-42.
Tatuno, S., Takei, M. 1969. Karyological studies in Hymenophyllaceae I. Chromosomes of the genus Hymenophyllum
and Mecodiumin Japan. Bot. Mag. (Tokyo) 82: 121-129.
Tindale, M. D. 1963. Hymenophyllaceae. Contrib. N. S.
Wales Nat. Herb. Flora Ser. 201: 1-49.
Wallace, J. W., Markham, K. R. 1978. Flavonoids of the primitive fems. Stromatopteris. Schizaea. Gleichenia. Hymenophyllum and Cardiomanes. Amer. 1. Bot. 65: 965-969.
Wessels Boer, J. G. 1962. The New World species of Trichomanes sect Didymoglossum and Microgonium. Acta Bot.
Neerl. 11: 277-330.
Yoroi, R. 1972. Studies in spore germination and gametophyte of Japanese Hymenophyllaceae. Sci. Rep. TokYO
Kyoiku Daig. B 15: 81-110.
Yoroi, R., Iwatsuki, K. 1977. An observation on the variation
of Trichomanes minutum and allied species. Acta Phytotax.
Geobot 28: 152-159.
163
Hymenophyllopsidaceae
K. U. KRAMER and D. B. LELLINGER
Hymenophyllopsidaceae [Christensen in Verdoom, Man.
Pteridol.: 532 (1938), sine descr. lat.;] Pichi Serm., Webbia
24: 712 (1970).
Small, delicate, epilithic or terrestrial ferns with shortto long-creeping, solenostelic, non-sclerotic stem
clothed with stramineous to dark brown, non-peltate,
non-clathrate, long and narrow, entire or denticulate
scales. Petioles spirally arranged, close, non-articulate,
sulcate, bisulcate, or terete, scaly at base (or villous in
H. superba Lellinger), with two vascular bundles that
coalesce upward. Lamina hairy when young, pinnate
+ pinnatifid to 3-pinnate + pinnatifid, anadromous,
free-veined, sometimes scaly like the petiole-base (villous in H. superba), of c. 3 or 4 layers of cells, only the
epidermis chlorophyllous; stomata wanting. Ultimate
segments often dichotomous, uninerval. Sori terminal
on a vein (but the veins often bearing branches flanking the sorus), subterminal on their segment; receptacle short. Indusium attached at the base and part of
the sides, short, anatomically like the lamina, forming
a kind of pouch with the laminal lobe opposite it, this
of about the same length, entire, lobed, or deeply lacerate. Sporangia few, intermingled with some hairs;
pedicel short, pluriseriate; capsule large, very asymmetric, with somewhat oblique annulus running past
the insertion and interrupted by the well-differentiated, pluricellular stomium. Spores globose, trilete, granulate, non-chlorophyllous. Gametophyte and karyology undescribed.
A single genus:
HymenophylJopsis Goebel
Hymenophyllopsis Goebel, Flora 124: 3, 21 (1929);
A.C.Smith (1931); Vareschi (1969); Lellinger (1984).
Eight described species, from Mt. Roraima (Guyana!
Brazil/Venezuela) to the mountains of western Venezuela; growing on moist rocks and moist earth, in sheltered places. The first known species is H. dejecta
(Baker) Goebel.
AFFINITY. This isolated genus is without known relatives. The resemblance to Hymenophyllaceae is only
superficial; anatomically the two families are very different, and the similarities must be due to convergence
through ecological parallelism. Goebel compared the
sporangium to that of the Cyatheaceae, which it resembles, but otherwise the two families are again very
dissimilar. Copeland (1947) called the genus "of Pteridaceous ancestry" and suspected affinity to "Orlhiopteris" (Sacc%ma), which seems possible but for the
164
Filicatae . Lomariopsidaceae
moment rests on insufficient evidence. The chromo·
some number and gametophytic characters may shed
more light on the affinity. At present Hymenophyllopsis
must be regarded as a leptosporangiate fern genus incertae sedis.
Selected Bibliography
Lellinger, D. B. 1984. Hymenophyllopsidaceae (Filicales). In:
The botany of the Guayana Highland XII. Mem. New
York Bot. Gard. 38: 2-9.
Smith, A.c. 1931. Pteridophyta. In: Gleason, H. A. (Ed.) The
Tyler-Duida expedition. Bull. Torrey Bot. Club 58:
299-315.
Vareschi, V. 1969. Hymenophyllopsidaceae. In: Lasser, T.
(Ed.) Flora de Venezuela I, 1. Caracas: Instituto BotAnico,
pp.254-258.
Lomariopsidaceae
K.. V. KRAMER
Lomariopsidaceae Alston, Taxon 5: 25 (1956).
Elaphoglossaceae Pichi Sermolli, Webbia 23: 209 (1968).
Bolbitidaceae (Pichi Sermolli) Ching, Acta Phytotax. Sin. 16
(3): 15 (1978).
Terrestrial, epilithic, scandent, or epiphytic, small to
rather large ferns with short- to long-creeping (or
-scan dent), less often erect stem bearing basifixed or
narrowly peltate, clathrate or non-clathrate scales that
are often marginally toothed or ciliate; stele most often strongly dorsiventral, occasionally radial, dictyostelic. Leaves close to remote, continuous or in one genus mostly articulate to phyllopodia; petiole usually
well-developed, sometimes winged, never dark-sclerotic and polished, variously scaly or glabrescent, with
several vascular bundles in a V-arrangement, the
adaxial ones larger and with hooked xylem strands;
adaxial face of petiole usually with a median groove,
sometimes with further grooves. Lamina simple, lobed
to pinnatifid, or simply pinnate, rarely bipinnate, pedate, or dichotomously compound; pinnae or pinnules
(if any) often articulate at base. Hairs, if present, very
small and usually glandular; scales often present on
leaves. Lamina with conform terminal pinna or with
gradually reduced and confluent upper divisions; dissection pattern often anadromous at base, catadromous or isodromous upward. Rachis like the petiole,
not rarely winged. Basal pinnae occasionally basitonically enlarged. Veins free, simple or forked, or variously anastomosing with or without free included veinlets.
Fertile leaves mostly non-conform, usually with contracted laminal parts, often diplodesmic. Sporangia
nearly always evenly covering the fertile parts abaxialIy, short- to rather long-stalked, the stalk mostly 3-seriate; capsule with vertical, interrupted annulus; stomium 2- to few-celled, not continuous with the bow.
Spores monolete, mostly with very prominent, folded
perispore. Sterile appendages sometimes present
among the sporangia.
MORPHOLOGY AND ANATOMY. The peculiar type of
stele is of particular diagnostic value for the family;
the dictyostele is strongly dorsiventra!, with a large,
strap- to gutter-shaped (in transection) ventra! strand
and 1-5 smaller, variously connected, dorsal strands
(e.g., Bell 1950; Nayar and Kaur 1964a, b; Hennipman 1977) (Fig. 84 C). In accordance with the placement of the leaf gaps the leaves are borne on the dorsal side only, in 2 or more (up to 5) rows; only some
species of Elaphoglossum have an erect, radially symmetric stem with the leaves borne all around. The
roots are in the former case inserted on the ventra! side
only.
The petiole has the vascular pattern typical of
Dryopteridaceae-Dryopteridoideae. An articulation
between stem or phyllopodium and petiole base occurs in Teratophyllum and most species of Elaphoglos-
sum.
Leaf dimorphism is nearly always present to some
degree, only in some species of Elaphoglossum fertile
and sterile leaves are nearly identical. Lamina! parts of
fertile leaves are usually shorter and/or narrower than
those of trophophylls, rarely wider ("Microstaphyla".
"Peltaptens"); in extreme cases the sporophylls may be
skeletonized (species of Bolbitis ser. Egenoffianae).
Sporophylls often have a special vein system underlying the sporangia (diplodesmic condition).
An additional heteromorphy is present in Teratophyllum and - much more weakly - in Lomagramma
and some species of Lomariopsis. The divisions of
leaves of juvenile, or at least non-adult, scandent
plants that have not yet reached the canopy with its
higher level of light incidence are ± deeply dissected
and often of thinner texture than the adult leaves
(acrophylls); such much-dissected leaves are called
bathyphylls. Transitions occur.
The range of dissection and venation patterns within the family is very wide, from deeply divided leaves
with one vein per segment to simple or simply pinnate,
free-veined leaves; to variously dissected (less often
single) leaves with sparingly to abundantly anastomosing veins with or without included, free veinlets. In
this respect even some genera (Bolbitis. less so Elaphoglossum) are variable.
The stomata are polocytic, anomocytic ones occurring much more sporadically.
Lomariopsis
Lateral aerating bands (pneumatophores) occur on
the petioles or petiole bases of most (all but the smallest?) representatives; they often extend onto the stem,
below the petiole insertion (e. g., Lloyd 1970).
The spores are monolete, ellipsoidal, usually with
prominent, wing-like folds or with reticulate surface.
GAMETOPHYTE. See Nayar and Kaur (1971) and Atkinson (1973). Most species have cordate prothallia, without, or sometimes with, multicellular marginal hairs.
In Elaphoglossum and some species of Bolbitis the prothallia are long-lived, slow-growing, and strap-shaped
(Fig. 85 J). The gametangia are of the usual advanced
type (Stokey and Atkinson 1957).
KARYOLOGY AND HYBRIDIZATION. Most counts are
based on n=41 or x=41, with tetraploids fairly frequent; triploids are known in, e. g., Bolbitis. Lomariopsis is heterogeneous, with base numbers from 32 to 41,
and multiples (Roy and Manton 1966). Elaphoglossum
subsect. Peltapteris has n=40 as opposed to 41 elsewhere. Thysanosoria has not yet been counted.
Hybrids are fairly numerous in Bolbitis, 13 instances
having been reported by Hennipman (1977); they are
as a rule sterile but may spread vegetatively by proliferous buds. Hybrids in other genera may still await detection, the species often being quite similar.
EcOLOGV AND DISTRIBUTION. The family is of pantropic distribution, with a concentration of genera in
the Old World and of species in the New. Only a few
species of Elaphoglossum extend significantly beyond
the tropics, to south-temperate or, much less, to northtemperate areas.
Ecologically the family is rather diverse; see under
the genera.
AFFINITY. Most authors agree that the family is closely
related to Dryopteridaceae where it is even included
by some as a subfamily or tribe (e. g., Tryon and Tryon
1982). Once the confusion of Lomariopsis and relatives
with Stenoehlaena was cleared up (Holttum 1932), it
was shown that the two could not even be placed in
the same family; anatomical characters and especially
the chromosome numbers bear this out well. Kaur
(1974) postulated derivation from the Teetaria group
of genera, but the dermal appendages are quite different. Holttum (1978) also envisaged a relationship with
Arthropteris and thus with Oleandraceae.
SUBDIVISION. Some authors exclude Elaphoglossum,
e. g., Pichi Sermolli (see above). But Holttum (e. g.,
1978) has emphasized its affinity to Lomariopsis. The
discrete, round sori of Thysanosoria, unique in the
family, may be indicative of relationship to Dryopteridaceae, but in its morphological-anatomical characters
the genus is highly specialized and typical of Lomariopsidaceae. On the basis of scale characters Holttum
(I. c.) believed Bolbitis and Lomagramma to be particu-
165
larly close. Neither a formal nor an informal subdivision has been proposed so far.
KEy TO THE GENERA.
1. Petiole base articulate (sometimes obscurely so); lamina
(with rew exceptions) simple, free-veined (if with reticulate
veins, without included veinlets), never pinnate with articu-
late pinnae; stem never scandent (may be long-creeping in
6. Elaphog/ossum
epiphytes)
- Petiole base non-articulate, decurrent; or, if articulate. the
leaves pinnate and the stem scandent; veins various
2
2. Sporangia in discrete, round sori on the outennost parts of
free veins; stem scandent
2. Thysanosoria
- Sporangia acrostichoid, at least when mature, rarely only
following the veins; stem various
3
3. At least the lateral pinnae articulate at base; or lamina simple and free-veined
4
- Pinnae (if any) non-articulate; lamina divided, or, if simple, with reticulate veins; veins very often amply reticulate
5. Bo/bitis
4. Terminal pinna jointed like the lateral ones; leaves pinnate
or bipinnate; veins various
5
- Tenninal pinna not jointed; leaves once pinnate, or lamina
simple; sterile leaves free-veined
1. Lommiopsis
5. Veins of sterile pinnae free; lamina often anadromous to
apex
3. TeratophyUum
- Veins of sterile pinnae amply reticulate, without secondary
(main) veins and without free included veinlets; lamina
cata-(or iso-)dromous towards apex
4. Lomagramma
1. Lomariopsis Fee
Fig.84D,E
Lomariopsis Fee, 2' Mem. 10: 66 (1845); Holttum (1932, 1939,
1940, 1966, 1978).
Stenochlaena auett., p. p., e. g., Underwood 1906 a, b.
Medium-sized to large ferns with long-scan dent stem
bearing shallowly peltate, lanceolate, non-clathrate, ±
ciliate scales, soon glabrescent, ridged. Leaves not
close; petioles usually well-developed, non-articulate,
decurrent, at least initially scaly, not dark-sclerotic.
Lamina simply pinnate, with few to many lateral and a
conform terminal pinna (in one species simple); rachis
like the petiole, deciduously scaly, sometimes winged,
adaxially shallowly grooved. Pinnae usually sessile,
suborbicular to narrowly lanceolate, herbaceous to
subcoriaceous, glabrous or minutely scaly, inserted on
the edges of the ridges bordering the adaxial rachis
groove, the lateral ones articulated at base, with a percurrent, adaxially flattened to grooved costa, the
groove not continuous with the rachis groove; terminal pinna conform, not articulated, or aborted. Veins
close, parallel, forked, quite free, anadromous in the
basal, catadromous or isodromous in the upper pinnae, (nearly) reaching the entire to sinuous, cartilaginous margin. Fertile leaves with much narrowed pinnae, their veins diplodesmic, sometimes somewhat
reticulate; lower surface entirely covered by sporangia,
these intermingled with small scales. Spores ellipsoidal, bearing prominent, echinate or cristate wings; or
surface bearing broad, reticulate wings.
166
Filicatae . Lomariopsidaceae
Fig. 84A-E. Lomariopsidaceae. A Teratophyllum aculeatum,
parts of stem with bathyphylls (left) and acrophylls (right)
(x 0.3) B Teratophyllum sp., stem ascending tree with bathy·
phylls (x 0.7). C Bolbitis appendiculata, stele of stem from
dotSal side (schematic). D, E Lomariopsis brackenridgei.
D Part of rachis with sterile pinna ( x 0.9). E Part of rachis
with fertile pinna (x 0.9) (A from Backer and Posthumus
1939; B from Holttum 1954; C from Flora Taiwan 1, 1975;
D, E from Brownlie 1977)
C.45 species, pantropic, about equally distributed
over the continents and extending to eastern Polynesia; only one in Australia. In forest shade, preferring
lower and middle elevations; sometimes on very moist
ground. The adult, scandent form is developed when a
creeping stem meets a tree trunk and becomes scandent; fertile leaves are apparently only produced when
more light is available. Bathyphylls none.
Formerly much confused with Stenochlaena (Blech.
naceae); easily distinguished by the lack of a vein
commissure flanking the costa, and by the stem structure.
2. Thysanosoria Gepp in
Gibb~
Thysanosoria Gepp in Gibbs, Dutch NW New Guinea: 19
(1917); Holttum (1978).
In most respects like Lomariopsis. Sterile pinnae herbaceous, with simple or basally forked veins. Fertile
pinnae much narrower than the sterile; sporangia in
small, round, exindusiate sori on the terminal parts of
veins, near the margin; the small lobe containing them
may be somewhat reflexed to the adaxial side.
A little known genus with a single species, Th. pteridiformis(Cesati) C. Chr., only known from two collections, from the coastal regions of NW New Guinea.
Hennipman (1977) regarded it as an atavistic form of
Lomariopsis. Further collections must be awaited.
Thysanosoria· Teratophyllum. Lomagramma· Bolbitis
3. Teratophyllum Mettenius ex Kuhn
Fig. 84 A, B
Terratophyllum Mettenius ex Kuhn, Ann. Ludg. Bat. 4: 296
(1869); Holttum (1932, 1966 a, 1978).
Arthrobotrya J. Smith (1875).
Stem as in Lomariopsis but more slender, often spiny;
scales pel tate, incised, deciduous. Leaves borne in 2,
rarely more, dorsal rows on the stem; petiole base
non-decurrent, swollen and ± functionally articulate.
Sterile leaves differentiated into bathyphylls and acrophylls, with transitions. Bathyphylls borne on juvenile,
or low on adult, scandent plants, with small, closely
set, lobed to pinnate (+ pinnatifid) pinnae that are
jointed to the (often winged) rachis. Acrophylls rather
as in Lamariopsis, but also the terminal pinna articulate; in a few species the leaves bipinnate. Some species with weakly to strongly asymmetric pinnae!pinnules, auriculate anteriorly, cuneate posteriorly, resembling Arthropteris species. Dissection anadromous near
apex or throughout, occasionally isodromous. Costae
(costules) adaxially not grooved. Fertile pinnae much
contracted, diplodesmic, sometimes with anastomosing veins, narrowly lanceolate to linear, sometimes auriculate, rarely again pinnate. Small, stalked scales
among the sporangia. Spores spheroidal to ellipsoidal,
echinate or reticulate, sometimes short folds present.
Nine species, from southern Burma to eastern
Polynesia and Queensland; most species in Malesia.
Ecology as in Lomariopsis.
4, Lomagramma J. Smith
Fig.85E
Lomagramma J. Smith, Hooker's J. Bot. 4: 152 (1841); Holttum (1966 a, 1978).
Medium-sized to large ferns with scandent stem;
scales peltate, (partly) clathrate, deciduous. Leaves remote, or in remote clusters, in 2-5 rows dorsally on the
stem; petioles decurrent as strong ridges on the stem,
well-developed. Leaf axes stramineous, often persistently scaly, the scales clathrate, the smaller ones bullate. Lamina simply pinnate, rarely bipinnate, mostly
with a conform terminal pinna, except in bathyphylls;
pinnae approximately lanceolate, sessile, herbaceous
to chartaceous, articulate at base. Terminal pinna, if
conform, also articulate. Leaves of juvenile plants and
bathyphylls often sinuate to lobed. Costa adaxially
convex, percurrent; no secondary veins; network of
tertiary veins uniform, with a few series of areoles between costa and margin; free included veinlets none;
vein ends free or loop-connected just behind the margin. Costae (and veins) abaxially sometimes bearing
some bullate scales. Margin entire to serrate. Venation!dissection catadromous, occasionally isodromous. Bathyphylls not strongly difform, with smaller,
often crenate pinnae. Sporophylls with much narrower
pinnae, diplodesmic, the veins less reticulate than
167
in sterile leaves; sporangia evenly covering their
lower side, intermingled with stalked scales, these
± filiform. Spores ellipsoidal, bearing broad,
echinate wings, or plain, with scattered, granulate
deposit.
C. 20 species, from NE India and southern China to
eastern Polynesia; not in Australia. A single species,
£. guianensis (Aublet) Ching, rather widespread in the
Neotropics, divergent by non-bullate scales, different
intersporangial appendages, and prominent, folded
perispore; to be excluded according to Holttum
(1978); but much less divergent than the genera
amongst each other and thus probably better left in L.
(Tryon and Tryon 1982), perhaps as a distinct subgenus.
5, Boibitis Schott
Figs. 84 C, 85 E- I
Bolbitis Schott, Gen. Fil.: pI. 13 (1835); Hennipman (1977,
1978).
Egenolfia Schott (1835); Nayar and Kaur (1964 b).
Campium C. Presl (1836); Copeland (1928), p. p.
Edanyoa Copeland (1952).
Leptochilus auctt., p. p., excl. typo
Medium-sized, less often small ferns, usually terrestrial
or epilithic, occasionally high-scandent; stem short- to
less often long-creeping or -scandent; scales basally attached to pseudopeltate, somewhat to strongly clathrate, with some, often gland-tipped, marginal hairs.
Petioles close to remote, very short to well-developed,
non-articulate, in 2-6 rows on the dorsal side of the
stem, often shortly decurrent onto it, adaxially sulcate
and ridged, or convex. Lamina triangular, oblong, less
often narrowly oblong or linear, truncate at base or cuneate in simple-leaved forms, glabrous when mature
but for the rachis; in a few species the basal divisions
basitonically developed (as in Tectaria). Dissection
anadromous (rarely isodromous) at base, catadromous
or isodromous above, pinnate to pinnate + pinnatilobate, less often pinnatifid or simple; upper pinnae
gradually reduced and confluent or an odd terminal
pinna present; leaf tip sometimes protracted, rooting
and proliferous. Pinnae adnate or sessile to shortstalked, non-articulate, costate, margin sometimes serrate. Rachis sometimes winged and!or scaly, adaxially
with a central ridge, sometimes with additional lateral
ones; costa ad axially convex, or grooved at base, the
groove then usually continuous with a dorsolateral rachis groove. Texture herbaceous to subcoriaceous. Pinna bases sometimes with a few persistent, small scales.
Margin in Series Egenolfianae with small, weak spines
terminating the most distal acroscopic vein branch.
Veins free, forked to subpinnate, or sparingly to amply
anastomosing, with or without free included veinlets,
these excurrent or pointing to a1I'sides. Marginal veins
free or arcuate-connected. Fertile leaves with moder-
168
Filicatae . Lomariopsidaceae
Fig. 8S A- K. Lomariopsidaceae. A Elaphoglossum buchii.
habit (x 0.7). B Elaphoglossum glaucum. plant with sterile
and (left) fertile leaf (x 0.5). C, D Elaphoglossum ("Microstaphyla'") b!{urcatum. C Stem with sterile leaf (x 0.5). D Segment of sterile leaf (x 4). E Lomagramma sumatrana. detail
of venation of sterile pinna of acrophyll (x 0.25). F Boibitis
("Egeno!fia'") appendiculata, sterile pinna (x 1.5). G Bolbitis
angustipinna. venation of sterile pinna (x 0.8). H Bolbitis
("Egenolfia') appendicu/ata ssp. vivipara. venation of sterile
pinna (x 0.8). I Bolbitis oligarchica. venation of sterile pinna
(x 0.8). J Elaphoglossum cuspidatum. prothallium (x 3).
K Elaphoglossum ("Peltapteris") peltatum. plant with sterile
and (right) fertile leaf (xO.5) (A from Christensen 1936);
B from Stolze 1981; C, D, K from Mickel 1980; E from
Holttum 1955; F from Aora Taiwan 1, 1975; G-! from Hennipman 1977; J from Atkinson 1973)
ately narrowed to almost wanting (skeletonized) laminal parts, usually less dissected, often more longstalked and borne strictly erect, beneath evenly
covered with sporangia (except in rare cases where
these follow the veins only); venation similar to that of
the sterile leaves, non-diplodesmic. No sterile organs
among the sporangia. Spores ellipsoidal or spheroidal,
bearing prominent, wing-like, usually echinate folds,
the wings sometimes reticulate to fenestrate.
44 species (Hennipman 1977) and a few hybridogenous ones, in the warmer parts of the world, extending
far east into the Pacific, and to NE India and southern
Japan; greatest species concentration in SE Asia; well
represented in continental Mrica; absent from dry
areas. Terrestrial or often epilithic, occasionally scandent or epiphytic, mostly in forests, or by streams, at
lower and middle elevations.
Elaphoglossum
Iwatsuki (1959) and Hennipman (loc. cit.) have
shown that the distinction of a genus Egeno{fia, with
free veins, is untenable.
6. Elaphoglossum J. Smith
Fig.85A-D,J,K
Elaphoglossum J. Smith, Hook. J. Bot. 4: 148 (1841),
nom. cons.; Christ (1899); Morton (1948); Alston (1958);
Sledge (1967); Schelpe (1969); Holttum (1978); Mickel and
Atehortua (1980).
Acrostichum L, p.p. min., and of many later authors, e. g., Fee
(1845}.
Peltapteris Link (1841); G6mez (1975).
Rhipidopteris Fee (1845), nom. superfl.
Hymenodium Fee (1845).
Microstaphyla C. Presl (1849).
Small to medium-sized ferns, epiphytic or epilithic,
less often terrestrial; stem creeping and dorsiventral or
erect and radial, bearing non-clathrate, often basally
cordate scales, their edge entire or toothed to (often
glandular-) ciliate. Leaves close to remote, in 2 or
more dorsal rows, or in a rosette in species with erect
stem, usually inserted on (often darker) phyllopodia of
varying length and articulate to them, rarely continuous. Petiole wanting to well-developed and exceeding
the lamina, often winged and!or subpersistently scaly,
not dark-sclerotic above the phyllopodium, ad axially
grooved, often with additional adaxio-lateral grooves,
or convex in small species. Pneumatophores mostly
present, of varying shape, sometimes forming outgrowths (Bell 1955; Lloyd 1970). Lamina simple, ±
dimorphic, the sterile often tongue-shaped to oblong
or lanceolate, also elliptic, spathulate, or linear; in a
few species once to several times deeply dichotomously incised ("Pe/tapteris'') or deeply pinnatifid with simple or forked segments ("Microstaphyla") (see Wagner
1952, for connecting shapes); when mature glabrous or
more often minutely to conspicuously and densely
scaly, the scales not rarely more persistent on costa beneath and margin, or on the entire lower surface, the
upper surface mOre often moderately to hardly scaly
when mature. Scales pale to dark, often of several
kinds together, usually incised to deeply fimbriate or
even resembling stellate hairs, sometimes with bristlelike, enrolled apex, in a few cases reduced to glandlike dots, diagnostically important. Texture of lamina
usually firm to coriaceous, less often soft and herbaceous. Costa percurrent (except in dichotomously
branched leaves which are non-costate), adaxially flattened or sulcate; veins lax or more often close, frequently immersed and not easily observed, simple or
usually forked, ± parallel, free or loop-connected just
behind the margin, very rarely reticulate; margin often
sclerotic; hydathodes not rarely present at vein ends.
Fertile leaves weakly to strongly difform, usually on
longer stalks and with iliortened and!or narrowed
lamina, usually not produced in large numbers or
169
throughout the year, beneath densely and evenly
covered with sporangia, sometimes with interspersed
scales; in some cases hair-like, multicellular appendages on the sporangium stalk or between the sporangia
(Anderson and Crosby 1966; Lloyd 1970). In the species with divided leaves the sporophylls are simple and
roundish-oblong (pedate in E. cardenasit). Sporangia
sometimes initially in circular sori which subsequently
coalesce (Lloyd 1970). Spores ellipsoidal to somewhat
spherical, bearing low folds which are often short, inflated, and perforate, or wing-like and fenestrate or
echinate, the elements sometimes irregular or fused into a reticulum.
CHARACTERS OF RARE OCCURRENCE. Reticulate veins,
without free included veinlets, in E. crinitum (L.)
Christ (Caribbean) and E. crassifolium (Gaud.) Anderson & Crosby (Hawaii). Proliferous laminas in a few
species, e. g., E. wageneri (Kunze) Moore (South
America). A pedately branched lamina in E. cardenasii
Wagner (South America) (see Mickel 1980). Erect,
short fertile leaves combined with pendulous, much
longer sterile leaves in E. herminieri (Bory ex Fee)
Moore (trop. America).
Estimated to have over 400 species; pantropic to
south-temperate, extending to remote Pacific and other islands; one in Australia, none in New Zealand;
very locally also north-temperate (Azores, Japan).
Greatest species concentration in tropical America, the
species there still imperfectly known. Many are epiphytes of montane forests; at higher altitudes increasingly epilithic and terrestrial, e. g., in paramos above
the timberline. Species occurring in seasonally dry
areas often have their leaves folded or curled up when
dry.
The African (Schelpe 1969) and Asiatic (Holttum
1978) representatives are rather well known.
Attempts at subdividing the large genus in a natural
way were first made by Christ (1899), subsequently by
Holttum (1966 b, 1978) for the Asiatic and by Mickel
and AtehortiIa chiefly for the American species (1980).
The former classification is informal, and the latter
tentative, in view of the very imperfect knowledge of
many neotropical species.
Selected Bibliography
Alston, A H. G. 1958. The Brazilian species of Eiaphoglossum. Bol. Soc. Brot. II. 32: 1-32.
Anderson, W. R., Crosby, M. R. 1966. A revision of the Hawaiian species of Elaphoglossum. Brittonia 18: 380-397.
Bell, P. R. 1950. Studies in the genus Elaphoglossum. I. Stelar
structure in relation to habit. Ann. Bot. (London) II 14:
545-555.
Bell, P. R. 1951 a. Studies in the genus Elaphoglossum. II. The
root and bud traces. Ann. Bot. (London) II 15: 333-346.
Bell, P. R. 1951 b. Studies in the genus Eiaphoglossum. III.
Anatomy of the rhizome and frond. Ann. Bot. (London) II
15: 347-357.
170
Filicatae • Lophosoriaceae
Bell, P. R. 1955. Studies in the genus Elaphoglossum. IV. The
morphological series in the genus and their phylo·
genetic interpretation, I. Ann. Bot. (London) II 19: 173199.
Bell, P. R. 1956. Studies in the genus Elaphoglossum IV. The
morphological series in the genus and their phylogenetic
interpretation II. Ann. Bot. (London) II 20: 69-88.
Christ, H. 1899. Monographie des Genus Elaphoglossum.
Denkschr. Schw. Naturf. Ges. 36: 1-159.
Copeland, E. B. 1928. Leptochilus and genera confused with
it. Philipp. J. Sci. 37: 333-416.
G6mez, L. D. 1975. Contribuciones a la pteridologia costarri·
cense. VI. El genera Peltapteris Link en Costa Rica. Brenesia 6: 25-31.
Hennipman, E. 1977. A monograph of the fern genus Bolbitis
(Lomariopsidaceae). Leiden Bot. Ser. 2: 1-331.
Hennipman, E. 1978. Bolbilis. In: R. E. Holttum 1978.
Holttum, R. E. 1932. On Stenochlaena, Lomariopsis and Teralophyllum in the Malay Region. Gard. Bull. S. S. 5:
245-313.
Holttum, R. E. 1939. The genus Lomariopsis in Madagascar
and the Mascarene Islands. Not. Syst. 8: 48-62.
Holtturn., R. E. 1940. New species of Lomariopsis. Kew Bull.
1939: 613-628.
Holtturn., R. E. 1966 a. The genera Lomariopsis, Teratophyllum
and Lomagramma in the islands of the Pacific and Australia. Blumea 14: 215-223.
Holttum, R. E. 1966 b. Florae Malesianae Precursores XLIV.
The fern genus Elaphoglossum in Malesia, with descriptions of new species. Blumea 14: 317-326.
Holttum, R. E. 1978. Lomariopsis group. In: Flora Malesiana
Ser. II, 1 (4): 255-330. SijthofT & NoordhofT, Alphen a. d.
Rijn.
Iwatsuki, K. 1959. Taxonomic studies of pteridophytes IV.
Emendation of Bolbilis, with special reference to the Far
Eastern species. Acta Phytotax. Geobot. 18: 44-59.
Kaur, S. 1974. The family Lomariopsidaceae (Filicopsida)
and its probable ancestors. J. Unn. Soc. Bot. 68: 153162.
Lloyd, R. M. 1970. A survey of some morphological features
of the genus Elaphoglossum in Costa Rica. Amer. Fern J.
60: 73-83.
Mickel, J. T. 1980. Relationships of the dissected Elaphoglossoid ferns. Brittonia 32: 109-117.
Mickel, J. T., Atehortua, G. L. 1980. Subdivision of the genus
Elaphoglossum. Amer. Fern J. 70: 47-68.
Morton, C. V. 1948. Notes on Elaphoglossum - II. The species
of the French West Indies. Amer. Fern J. 38: 202-214.
Nayar, B. K. 1966. Morphology of the fertile leaves of the
Lomariopsidaceae with special reference to the venation.
New Phytol. 65: 221-238.
Nayar, B. K., Kaur, S. 1964 a. Ferns of India - XI. Bolbitis.
Bull. Nat. Bot. Gard. India 88: 1-74.
Nayar, B. K., Kaur, S. 1964 b. Ferns of India - XIII. Egenolfla. Bull. Nat. Bot. Gard. India 100: 1-38.
Roy, S. K., Manton, I. 1966. The cytological characteristics of
the fern subfamily Lomariopsidoideae sensu Holttum. J.
Unnol. Soc. Bot. 59: 343-347.
Schelpe, E. A. C. L. E. 1969. Reviews of tropical African pter·
idophyta I. 4. The Lomariopsidaceae of continental tropical Africa. Contrib. Bolus Herb. 1: 25-43.
Sledge, W. A. 1967. The genus Elaphoglossum in the Indian
Peninsula and Ceylon. Bull. Brit. Mus. Nat. Hist. (Bot.) 4
(2): 81-96.
Stokey, A. G., Atkinson, L. R. 1957. The gametophyte of some
American species of Elaphoglossum and Rhipidopleris. Phytomorphology 7 (3/4): 275-292.
Underwood, L. M. 1906a. The genus Stenochlaena. Bull. Torrey Bot. Club 33: 35-50.
Underwood, L. M. 1906 b. American ferns - VII. The American species of Stenochlaena. Bull. Torrey Bot. Club 33:
591-605.
Wagner, W. H. 1952. Types of foliar dichotomy in ferns.
Amer. J. Bot 39: 578-592.
Lophosoriaceae
K. U.KRAMER
Lophosoriaceae Pichi Sermolli, Webbia 24: 700 (1970).
Rather large to large, terrestrial ferns with massive, ascending stem to a few rn tall, radial in structure, solenostelic (with transitions to dictyostely), bearing
long, pluricellular, tawny to castaneous trichomes;
creeping part bearing reduced leaves, normal leaves
borne on the erect part. Petioles rather close, well-developed, adaxially sulcate, bearing long, pluricellular,
tawny hairs at the base, often bearing adventitious
buds on their abaxial bases which may develop into
runners; vascular bundles three, highly corrugated,
one abaxial, two adaxial, all with involute margins.
Lamina bipinnate + pinnatifid to tripinnate + pinnatifid, elongate·triangular, truncate at base; rachis
adaxially broadly sulcate with centrally convex
groove, bearing shorter, ± deciduous, tawny hairs, especially in the groove. Pinnae numerous, petiolulate,
the grooves on their adaxial side not continuous with
that of the rachis, more persistently short-hairy; bases
of largest petiolules with a rudimentary articulation.
Primary pinnae (pseudo 1)anadromous, secondary
(pseudo?)ana- or catadromous, divisions of higher order catadromous (Kramer 1987). Upper divisions
gradually reduced and confluent; ultimate segments
sessile, lobed to pinnatifid, oblong-Ianceolate, firm, often glaucous beneath and ± pubescent. Rachises of
higher order ridged in the middle, laterally grooved on
each side by the presence of narrow wings formed by
the decurrent pinnules and segments. Veins free, mostly simple. Stomata paracytic. Sori single and dorsal on
the veins, with hardly raised receptacle, round, exindusiate; pluricellular, filiform trichomes present among
the sporangia. Sporangia with a short, c. 6-seriate stalk
and a complete, subvertical annulus bypassing the
stalk, with weakly differentiated stomium. Spores 64,
trilete, globose with large equatorial flanges, the wall
structure unique, formed by a double wall on the
proximal face, the distal face perforate; surface tuberculate, ± covered with a granulate deposit.
171
ANATOMY. See references under Metaxyaceae (Bower;
Holttum and Sen; Lucansky).
GAMETOPHYTE. Described as obcordate-elongate,
mostly glabrous, with about 5-celled antheridia; overall structure somewhat irregular, not unlike that of
Cyatheaceae (Stokey 1930).
KARYOLOGY. The number of n=65 diverges from
those of both Metaxya and Cyathea. The number of
n = 69 in all Cyatheaceae (s. str.) is only doubtfully indicative of affinity.
Fig. 86 A-C. Lophosoriaceae. Lophosoria quadripinnata.
A Base of petiole (x 0.6). B Pinna base and part of rachis
(x 0.6). C Sporangia seen from two sides (x 100) (A, B from
A. R. Smith 1981; C from Bower 1926)
AFFINITY. According to Lucansky (Ioc. cit.), this is a
primitive genus, most closely related to Metaxya but
also showing relationship to Cyatheaceae and Dicksoniaceae. Lophosoria and Metaxya are certainly not
close enough to constitute a family "Protocyatheaceae" (Bower 1926). A single genus:
172
Lophosoria C. Presl
RHeatae . Loxomataceae
Fig. 86
Lophosoria C. Presl, Gemssb. i. Stipes d. Farm: 36 (1847).
Alsophila auett., p. p. m.
CHARACTERS OF THE FAMILY. A single species, L. quadripinnata (Gmelin) C. Chr., widespread in montane
areas of tropical America, from Cuba and southern
Mexico to Bolivia and SE Brazil; also occurring disjunctly in moist, temperate areas of central and southern Chile and in the Juan Fernandez Islands (Looser
1933), there at lower altitude. A high-altitude form
with compact, erect leaves has been described as var.
contracta (Hieron.) R. & A. Tryon.
Selected Bibliography
Kramer, K U. 1987. A brief survey of the dromy in fern
leaves, with an expanded terminology. Bot. Helv. 97:
219-228.
Looser, G. 1933. Sobre las Ciateilceas chilenas y en especial
sobre Lophosoria quadripinnala. Ostenia 1933: 141-151.
Stokey, A. G. 1930. Prothallia of the Cyatheaceae. Bot. Gaz.
90: 1-45.
For further references see under Metaxyaceae.
Loxomataceae
K.U.KRAMER
Loxomataceae C. Presl, Gefiissb. i. Stipes d. Farrn: 31 (1847)
("Loxsomaceae").
Terrestrial, medium-sized to large ferns with longcreeping, branched, solenostelic stem bearing dark,
stiff, multicellular, basally pluriseriate bristles (similar
to Dipteris). Leaves remote, non-articulate; petiole
well-developed, with a single, in transection guttershaped vascular bundle with incurved base, naked except for the base, adaxially convex or flattened-sulcate. Lamina bipinnate + pinnatifid to tripinnate +
lobed, firm, hypostomatic, narrowly to broadly deltoid, catadromous, at least above the base; ultimate
segments deltoid-lanceolate. Stomata cryptopore, paracytic. Veins free, forked-subpinnate, catadromously
branched. Sori terminal on a vein, marginal, protruding beyond the margin; indusium urceolate, laterally
free; receptacle columnar. Sporangia on thick, short,
pluriseriate stalks, ± pyriform; annulus subvertical,
slightly oblique, continuous but only partly indurated;
multicellular trichomes present among the sporangia.
Spores trilete, non-chlorophyllous.
MORPHOLOGY AND ANATOMY. Only Loxoma has been
somewhat closely studied; see Bower (1926), von
Goebel (1930), and Ogura (1972). The essential characters, as outlined above, are shared by both genera.
GAMETOPHYTE. See Stokey and Atkinson (1956). In external shape, which is obcordate to elongate, the prothallium offers few remarkable features except for the
presence of multicellular, usually pluriseriate hairs,
especially on the ventral cushion. The antheridia are
of a primitive, pluricellular type with often divided cap
cell, and sometimes extend to the upper surface; they
are somewhat reminiscent of those of Cyatheaceae.
KARYOLOGY. The numbers for Loxsomopsis: n =46,
and for Loxoma: n = 50 (Brownsey 1975) do not agree
with any other genus whose affinity can be considered
on morphological grounds.
AFFINITY. Very little is known about this; suggestion of
relationship with Hymenophyllaceae were based only
on superficial similarity of the soms and are not substantiated by any other character. In prothaIIiai and
stomatal features there is a certain resemblance to
Cyatheaceae, in stomata also to Dipteridaceae, with
which the type of hairs is in agreement, too; but the
evidence from such characters is slight. Loxomataceae
are undoubtedly an isolated and ancient group. A Jurassic genus, Stachypteris. has been referred to the
family with some confidence.
ECOLOGY AND DISTRIBUTION. The very disjunct distribution of the two genera, and also of the known populations of the first, strengthens the impression of a relic
group. Loxsomopsis occurs in open places of moist
tropical forests; it is found from Costa Rica South to
Bolivia. Loxoma ist confined to lowland forests of a
small area in the north of the North Island of
New Zealand.
KEy TO THE GENERA.
1. Lamina catadromous, sometimes persistently hairy on the
axes and veins abaxially, not glaucous. Pinnae (sub)sessile,
asymmetric, the basal posterior divisions reduced
(Fig. 87 A). Sporangia with transverse dehiscence
(Fig. 87 D). New World
1. Loxsomopsis
- Lamina anadromous at base, otherwise catadromous, glabrous when mature, often glaucous beneath. Pinnae±
equal-sided; lower pinnae stalked. Sporangia with longitudinal dehiscence. New Zealand.
2_ Loxoma
1. Loxsomopsis Christ
Fig. 87A-C
Loxsomopsis Christ, Bull. Herb. Boiss. II, 4: 399 (1904).
Stem long-creeping; leaves remote, up to 5 m long.
Lamina narrowly oblong, at base bipinnate + pinnatifid; catadromous; pinnae (sub)sessiIe, subopposite,
asymmetric, the posterior basal pinnules/segments reduced. Abaxial side of lamina sometimes persistently
hairy on axes and veins, the hairs articulate. Secondary
pinnae and segments obliquely ascending, decurrent;
Loxsomopsis· Loxoma
margin crenate-dentate to entire. Sori directed downwards from plane of lamina. Sporangia with almost
vertical, complete annulus bypassing the stalk, indurated except for a group of laterobasal, smaller, thinner cells; dehiscence transversal. Spores tetrahedralglobose; surface tuberculate or rugose with ± fused
rugae and pits.
Four described species, but more probably only
forms of a single species (Tryon and Tryon 1982): L.
costaricensis Christ. More or less scattered, isolated
popUlations in Costa Rica and in South America from
southern Colombia to Bolivia.
2. Loxoma
Fig. 87D-F
Loxnma R. Brown in A. Cunningham, Hooker's Compo BoL
Mag. 2: 366 (1836) (often misspelled "Loxsoma').
Stem creeping, relatively stout; leaves not close. Petiole well-developed; lamina deltoid, up to C. 60 em
long, coriaceous, often glaucous to whitish beneath,
173
Fig, If] A-F. Loxomataceae. A-C Loxsomopsis COSlarieensis.
A Part of lamina with one complete pinna (x 1). B Two fertile segments (x 3.5). C Longitudinal section of sorus (x 40).
D-F Loxoma cunninghamii. D Pinna (x 1). E Sorus (x 5).
F Sporangium (x 50) (A-C from Christ 1904; D, E from
Diels 1899)
naked, at the base tripinnate + pinnatifid; lower pinnae s ubopposite, stalked; primary dissection anadromous at base, catadromous above. Pinnae about
equal-sided; smaller pinnae, pinnunules and segments
decurrent, upper parts of axes winged by their decurrent bases. Sori much as in the preceding genus. Sporangia with lateral, ± vertical, complete annulus of
which only the distal cells are indurated; dehiscence
longitudinal. Spores tetrahedral-globose; surface rugulose, pitted, with low, coalescent ridges.
A single species, L. cunninghamii R. Brown in
A. Cunningh., confined to northern New Zealand.
174
Filicatae . Marattiaceae
Selected Bibliography
Brownsey, P.J. 1975. A chromosome count in Loxsoma. New
Zeal. J. Bot. 13: 355-360.
Goebel, K. von 1930. Organographie der Pflanzen. 3 Aufl.
2. Bd. Jena: Gustav Fischer.
Stokey, A. G., Atkinson, L R. 1956. The gametophyte of Loxsoma cunningham;; R Br. and Loxsomopsis costaricensis
Christ. Phytomorphology 6: 249-261.
Marattiaceae
I.M.CAMUS
Marattiaceae Bercht & J. S. Presl, Pfirozen. Rostl. 1: 272
(1820).
Terrestrial ferns. Stem mycorrhizal, generally unbranched, radial or dorsiventral, slender and creeping
or ± stout and erect, often apparently massive since
clothed with pairs of large, persistent photosynthetic
stipules which protect the young croziers. Roots fleshy,
often produced aerially. Leaves clustered or ± distant.
Petiole sometimes grooved on the upper side, with one
swollen area (pulvinus) at the base, another at the base
of the laminate portion of the leaf and sometimes one
or more pulvini in the middle region, sparsely to
densely covered in non-clathrate, pel tate (except in
Christensenia) scales which may be borne on podia.
Lamina simple to 4-pinnate (subpedate in Christensenia), catadromous, with pulvini at the junctions of the
different orders of pinnae and pinnules with rachises
and also at the junctions of the different orders of rachises; rachises terete, sometimes winged. Pinnae or
pinnules thick (membranous only in Danaea trichomanoides Spruce ex Moore), sparsely scaiy, rarely with
uniseriate multicellular hairs, costate, margin with an
often barely distinguishable tooth or lobe at each vein
ending. Venation catadromous, veins characteristically
free, generally simple or once-furcate (merging to form
a submarginal vein in some species of Danaea), but
reticulate in Ozristensenia; occasional veins bifurcate.
Sori exindusiate, generally superficial (strongly sunken
in Danaea) placed along lateral veins (or on a veinlet
in Christensenia), composed of a double row or ring of
free sporangia, or sporangia united to form a synangium. Sporangia or synangia sessile on ·the receptacle
but development of the receptacle very variable, generally glabrous but deciduous scales occur on synangia
of Christensenia; uniseriate multicellular branched
hairs may be attached to the receptacle but not in
Christensenia or Danaea; dehiscence by longitudinal
slits (pores in Danaea), maturation ± simultaneous,
spore output several thousand per sporangium or synangial compartment. Spores monomorphic, typically
globose and trilete or ellipsoid and monolete, sometimes intermediate or alete, longest dimension c.
20-40 ,urn; exospore ornamentation granular or warty
to spiny, spines simple, bifurcate to multifurcate in
some species of Angiopteris (Fig. 88); perispore present
but of variable thickness. Germination results in a
plate-like mass which develops into a large, dark green
gametophyte that resembles an anthoceroid liverwort.
MORPHOLOGY AND ANATOMY. Marattiaceae are unique
amongst extant ferns in the following features: endarch protophloem, multicellular as well as unicellular
root hairs; mucilage canals both in the roots and
leaves as well as in the stem; massive starchy stipules
(trophopods); pulvini; ±strongly cyclocytic stomata;
and the tight packing of the sporangia or synangial 10cules within the sorus or synangium.
Other unusual characters are found in members of
this family. Their protoxylem is usually endarch, a
character shared with Ophioglossaceae but not with
Filicales. The stem and stipe vasculature in many species is distinctive and more complex than in other extant ferns; in three dimensions it is like a series of internested and interlinked cones and shows concentric
rings of meristeles in transverse section. Stems range
from monocyclic with a central strand in Christensenia
to 14-cyclic in some Angiopteris plants. Mucilage is a
very characteristic feature of this group, but its function is unknown. The pulvini lack collenchyma in their
internal tissue, with the result that the leaf, pinnae, and
pinnules droop at these places when under water
stress. The massive stipules of large plants have lenticels on both their adaxial and abaxial surfaces. Stipule
form is very variable, but is constant for a plant and
may be of taxonomic use (Fig. 91). Lenticels are borne
all round the petioles of the larger plants, unlike large
members of Filicales in which the lenticels occur in
two discrete rows. The fronds are long-lived except for
fertile ones of Danaea. The degree of pinnation of
leaves is apparently under loose control and is easily
affected, sometimes at least by environmental factors:
a leaf or part of one may show a greater or lesser degree of pinnation than the rest of the leaf or leaves
(this is especially evident in adolescent leaves). Apices
of frond pinnae or pinnules of many plants (excluding
Christensenia) may be imparipinnate or paripinnate,
or so alternately paripinnate as to appear imparipinnate. Stomata of Christensenia are over 80 p.m, possibly the largest known in vascular plants, and are permanently open. Pinnae or pinnules of some species of
Angiopteris show conspicuous fibre bundles (often referred to as "venulae recurrentes") between and parallel with the lateral veins. Microscopic projections of
unknown function but probably composed of cellulose
Filicatae • Marattiaceae
175
D,C
A
D
E,F
occur on the exterior of the mesophyll cell walls.
These are known to occur in other ferns, for example,
Helminthostachys of the Ophioglossaceae. The wall of
the sporangium or synangium that dehisces (ventral
wall) is±one cell thick at maturity; the other walls
(i. e., dorsal and lateral or compartment walls) are 2 or
3 cells thick at maturity. Sporangia of Angiopteris show
the greatest differentiation of cells in the outer layer of
the walls, typically with a ± conspicuous group
( = "hood") of ± isodiametric thickened cells near the
apex above the dehiscence slit. Together with the elongate thickened cells of the lateral wall this has formerly been referred to as an annulus by some authors.
Differentiation of cells in the outer layer of the sporangial or synangial walls may be of taxonomic value
(Fig. 89).
KARYOLOGY. A basic chromosome number x=40 occurs in every genus studied (e. g., Manton and Sledge
1954; Walker in Manton 1959; Walker 1966, 1979,
1981,1985). Two records of x=39 for Marattia (Ninan
1956; Brownlie 1%1) and a count of n =116 for
Danaea nodosa (L.) Smith (Sorsa in Odum and Pigeon
1970) require further investigation. All genera show
polyploidy and all recorded species of Danaea are te-
Fig. 88A-F. Unacetolyzed spores of Marattiaceae. A Angiopteris somai, bar=8.3.urn.
B Angiopteris tonkinensis.
bar- 8.5 /l-m. C Christensenia aesculifolia. bar- 8.5 /l-m.
D Danaea elliptica. bar-8.5/l-m. E Anglopteris smith;;.
bar-8.3 /l-m. F Marattia fraxinea. bar-B.l /l-m. (Original
Camus)
traploids, although the occurrence of a triploid
Danaea hybrid (D. jenmanii Underw. xjamaicensis
Underw.; T. G. Walker 1966) suggests (as may Sorsa's
count) that diploids occur in this genus.
VEGETATIVE REPRODUCTION. Vegetative propagation
occurs through growth of adventitious buds on the stipules, at least in the larger plants, and provides a
means for bringing plants into cultivation. Jenman
(1909) considered that in Marattia alata Swartz stipular reproduction is more common than that by spores.
Plants growing from stipule buds appear to pass
through the same developmental stages of leaf form as
those From gametophytes. Adventitious buds may be
produced near the apex of the rachis in Danaea plants
of lax habit.
GAMETOPHYTE. Gametophyte assumed to be very longlived and remarkably slow-growing in the wild as it is
176
FiJicatae . Marattiaceae
B
A
Fig. 89 A, B. Sori of Marattiaceae. A Marattia brassii,
bar- 200 11m. B Angiopteris evecta. bar= 200 11m (Original
Camus)
in culture; glabrous, mycorrhizal, cordate, up to several em long, sometimes branched; more than one cell
thick except at the margin, midrib not clearly defined.
Rhizoids unicellular and multicellular. Antheridia
borne dorsally and ventrally, large embedded, producing more than 100 spermatozoids. Archegonia borne
ventrally, short, broad, with scarcely projecting necks
and a poorly developed jacket layer. Embryo emerges
through the dorsal surface of the prothallus (Brebner
1896; Stokey 1942; Atkinson and Stokey 1964; Bierhorst 1971).
SUBDIVISION AND RIlLATlONSHIPS. Marattiaceae are
considered here to comprise all extant marattialean
ferns. Many alternative schemes regarding subdivisions and relationships within Marattiales have been
proposed by various authors (see Hill and Camus 1986
for a general review). Preliminary cladistic analysis
based on morphology and anatomy by Hill and Camus showed that only Christensenia and Danaea are
monophyletic and did not resolve the relationships between Danaea and the paraphyletic genera Marattia
and Angiopteris. The sequence of first recorded stratigraphic appearances of these and closely related taxa in
the fossil record is consistent with their view of the
phylogeny of extant taxa.
AFFINITIES. This family has only distant affinity with
other extent ferns and (excluding Ophioglossales) is
generally considered the most primitive surviving
group.
DISTRIBUTION. Marattiaceous ferns occur throughout
the tropics and subtropics. Marattia is found worldwide, Danaea in the New World only. Christensenia
has a disjunct distribution in Asia and Malesia. An-
giopteris grows more widely: from Madagascar in the
west, throughout Asia and Malesia, and east to Tahiti.
ECOLOGY. Plants of wet tropical and subtropical primary and secondary forests on steep clay banks, e. g.,
sides of streams, gullies, and ravines, where the canopy is dense; up to c. 3000 malt.
USES. Whilst members of Marattiaceae are of no economic importance, the stipules of Marattia have been
eaten by Maoris in New Zealand. Those of both Marattia and Angiopteris have been eaten by man elsewhere in times of extreme food shortage, and are frequently eaten by wild pigs. An alcoholic drink has
been brewed from stipules and an aromatic oil obtained from Angiopteris plants (May 1978). Pinnae of
Angiopteris have been used as head-dresses at feasts
and the huge leaves as temporary beds afterwards.
Marattia fraxinea Smith is grown as a garden plant in
New Zealand.
PALAEOBOTANY. The distinctive stem anatomy and
structure of the sori or synangia link members of this
group with several fossil genera, the best known being
Psaronius Cotta (stem) and Scolecopteris Zenker (fertile leat). The first securely identifiable fossils are of
these two genera from Lower to Middle Pennsylvanian age rocks (Upper Carboniferous, Westphalian A).
Qasimia Hill, Wagner & EI-Khayal, which closely resembles taeniopteroid species of Marattia but with
rounded pinnule apices, is known from the Late Permian (Hill et al. 1985), Marattia is first recorded from
the Triassic onwards, and Angiopteris (Hill 1987) with
any certainty from the Jurassic. Other notable fossil
genera are Danaeopsis Heer (Triassic, e. g., P'an 1936),
Millaya Mapes & Schabilion, Eoangiopteris Mamay
and possibly (1) the gigantopterids (Li and Yao 1983).
KEy TO THE GENERA.
1. Fertile leaves bearing sori composed of separate or almost
separate sporangia
1. Angropteris
- Fertile leaves bearing synangia
2
177
Angiopteris. Christensenia
D
M
F
A
2. Synangia circular, veins reticulate
2. Christensenia
- Synangia linear, veins free (rarely fusing to fonn a submarginal vein)
3
3. Each synangium a fused double row of spore-bearing compartments, each compartment dehiscing by a ± circular
pore, leaves dimorphic
3. DanQ£{l
- Each synangium a bivalved structure, valves gaping apart
at maturity, each compartment dehiscing by a longitudinal
slit, leaves monomorphic
4. Marattia
1. AlIgWpteris Hoffmann
~
Q!Q2
E
m
G
Fig.90A-G. Marattiaceae. Maraltia laevis. A Two secondary
pinnae on secondary rachis (x 1). B Two segments with synangia· (x 2). C Dehisced synangium (x 5). D-G Schematic
transections of sporangia and synangia. D Angiopteris.
E Christensenia. F Maraltia. G Danaea (A-C from Vareschi
1969; D-G after Eames 1936, redrawn)
Figs. 88 E, 89 B, 90 D, 91, 93
Angiopleris Hoffmann, Commenta!. Soc. Regiae. So. Got!. 12:
29 (1796), nom. cons.; Ching 1958, Chang 1975, Holttum
1978
Archangiopteris Christ & Giesenhagen (1899)
Macroglossum Copeland (1909)
Protomarattia Hayata (1919)
Protangiopleris Hayata (1928)
Stem creeping and slender or erect, sometimes massive. Leaves clustered, arising radially, 1· to 2'pinnate,
up to 7 m long, bearing deeply fimbriate scales that
usually have long auricles; scales composed of elon·
gate cells. Leaves of juvenile plants typically with one
pulvinus (0 in one species, several in one species) in
middle region of petiole; leaves of adult plants lack
this pulvinus in bipinnate species, pulvinus «()..) 1
(-several) in one-pinnate species. Pinnules sparsely
scaly (one species with multicellular, uniseriate hairs
on lower surface), shortly petiolate, typically strapshaped (lanceolate to falcate in some species), unequal
to cuneate at base, margin serrate or scarcely serrulate,
apex acuminate and more strongly toothed. Most species have conspicuous fibre bundles between and parallel with main lateral veins. Sori short and submarginal or long and medial, composed of a double row of
sporangia borne on a receptacle; receptacle usually
bearing conspicuous uniseriate multicellular branching hairs. Sporangia typically superficial but slightly
sunken in lamina tissue in a few species, occasionally
showing some fusion but of their dorsal walls only and
rarely extensive enough to form a "pseudosynan-
gium"; cells of outer layer of sporangial wall highly
differentiated. Spores typically globose, trilete; exospore ornamentation of warts to simple or multifurcate
short spines.
Some 200 mostly poorly described "species" from
Madagascar, India, Sri Lanka, Indo-China, China,
throughout Malesia, Solomon Islands, New Hebrides,
north to Japan and east to Tahiti and recently naturalized in Jamaica. Extensive fieldwork and breeding
studies are required for worthwhile monographing of
this genus. Vegetative plants of Angiopteris may be difficult to distinguish from those of some species of Marattia.
2. Christensellia Maxon
Figs. 88 C, 90 E, 92,93
Christensenia Maxon, Proc. BioI. Soc. Wash. 18: 239-240
(1905).
Stem creeping, dorsiventral. Leaves arising dorsally,
± remote, up to 1 m long; petiole bearing scales and
uniseriate multicellular hairs; lamina (simple, trifoliate
or) subpedately 5-foliate, pinnae shortly petiolate,
broad, lanceolate to falcate, glabrous above, with
scales and uniseriate multicellular hairs below, base
cuneate to oblique, margin entire to serrate, apex acute
to acuminate; stomata polycyclocytic, c. 80 pm or
more in diameter. Soms a circular synangium with up
to 20 compartments; synangia either scattered or ± in
2 rows parallel with main lateral veins. Spores typically ellipsoid, monolete; exospore spiny.
178
Filicatae . Marattiaceae
One variable species, C. aesculifolia (Blume) Maxon, in lowland forests of Assam, Arunachal Pradesh
(India), SE Yunnan (China), extreme south of Thailand, Peninsular Malaysia, Sumatra, Java, Moluccas,
Borneo, Philippines, New Ireland and the Solomon Islands (not collected in mainland New Guinea). A tetraploid plant (n = 80) was reported by Braithwaite
(1977) from San Cristobal, Solomon Islands, and
a diploid (n=40) recorded from Sarawak by T.G.
Walker (1979). The distribution of the two cytotypes
would be of interest.
3. Danaea J. E. Smith
Fig. 880, 90G, 93
Danaea J. E. Smith, Mem. Acad Sci. Thrin 5: 420, t.9, r.l1
(1793), nom. cons.; Underwood (1902).
Stem creeping or erect. Roots with sclerified pith and
hypodermal cortical wne. Leaves arising radially, clustered to ±remote, up to c. 2 m long, dimorphic; petiole
Fig. 91. Matattiaceae. Angiopteris evecta. young unfurling
crozier issuing from among the massive, paired stipules, white
spots ate lenticels; New Guinea. Pho!. F. Greenaway bar=
1,4 em
with 0 to several pulvini in the middle region, bearing
scales composed of ± isodiametric cells. Lamina 1-pinnate (simple in 1 species), rachis conspicuously winged
in some spp.; pinnae sparsely scaly, shortly petiolate,
strap-like to lanceolate; base cuneate to obtuse,
margin ± entire; apex usually acuminate and serrulate;
pinnae of fertile leaves shorter, narrower and apex less
acuminate than those of pinnae of vegetative leaves.
Stomata with lateral subsidiary cells in ± paracytic position. Sorus an elongate syoangium composed of a
double row of spore-bearing compartments extending
across almost the whole of the pinna half-width; compartments fused laterally and ventrally. Spores typically ellipsoid, monolete; exospore ornamentation of
simple spines or some spines fused laterally. About 30
Danaea· Marattia
179
Fig. 92. Marattiaceae. Christensenia aesculifolia; Sabah,
Borneo. Photo A. C. Jermy
ANGIOPTERIS
spp. described, few of which would survive monographic revision. Limited to the lowland tropics and subtropics of the New World.
Presl's (1845) subdivisions have not been generally
accepted.
:!:
.;
.,;
W
~
Z
'zw"
I-
4. Marattia Swartz
Fig. 88F, 90A-C, F
Marattia Swartz, Prodr.: 8, 128 (1788)
Stem erect. Leaves arising radially, clustered, up to 4 m
long; petiole never with a pulvinus in the middle region, bearing deeply fimbriate scales with long auricles; scales composed of elongate cells. Lamina 1- to
4-pinnate, ultimate rachises usually winged, pinnules
sparsely scaly, sessile to shortly petiolate, strap-like
to ± oblong to lanceolate, base cuneate to unequal,
margin scarcely to conspicuously serrate, apex acuminate to serrulate. Fibre bundles lacking. Soms a median or submarginal elongate bivalved synangium, each
valve attached to the receptacle and composed of
several compartments; receptacle development variable (synangium appearing stalked in M. laevis Smith),
receptacle usually bearing uniseriate multicellular
branched hairs. Spores typically ellipsoid, monolete;
exospore granular to spiny.
'"a:
J:
0
0(
0(
;::
W
I0(
w
0
"a:
a:
..
I-
a:
c;
0(
c
::;
0(
0(
::J
'"'"0
z
0(
::;
z
,.J
s. I.
'"a:
..
W
I-
0
c;
Z
0
0(
0
J:
0
0(
::;
a:
0(
Fig. 93. Representation of relationships between the genera
of Marattiaceae (Original Camus)
180
Filicatae . Marsileaceae
A pantropical genus of some 70 described species,
extending to the southern subtropics, typically growing at higher altitudes and in denser shade than do
species of Angiopteris. In need of monographic revision. The subgeneric divisions of Presl (1845) and Rosenstock (1908) are not generally accepted but may
prove to have some value.
Selected Bibliography
Atkinson, L R, Stokey, A.G. 1964. Comparative morphology
of the gametophyte of homosporous ferns. Phytomorphology 14: 51-70.
Berchtold, F. von, Presl, J. S. 1820. 0 pnrozenosti rostlin,
aneb rostlinlif, etc. 1. Prague: Joz. Krarse.
Braithwaite, A. F. 1977. A chromosome count and range extension for Christensenia (Marattiaceae). Amer. Fern. J. 67:
49-50.
Brebner, G. 1896. On the prothallus and embryo of Danaea
simplicifolia Rudge. Ann. Bot. (London) 10: 109-1'21.
Brownlie, G. 1961. Additional chromosome numbers New Zealand ferns. Trans. R. Soc. New Zeal. Bot. 1: 1-4.
Cambie, R. C. 1961. A New Zealand phytochemical survey 3.
The ferns and fern allies. New Zeal. J. Sci. (Wellington) 4:
707-714.
Camus, J. M. (in press) The limits and affinities of marattialean fern genera in China and the West Pacific. Proc. Int.
Symp. Systematic Pteriodology, 1988, Beijing.
Chang, C.-y. 1975. Morphology of Archangiopteris Christ &
Gies. and its relationship with Angiopteris Hoffm. Acta
Bot. Sin. 15: 235-247.
Ching, R. C. 1958. A revision of the fern genus Archangiopteris Christ & Giesenhagen. Acta Phytotax. Sin. 7: 212-224.
Hill, C. R. 1987. Jurassic Angiopteris (Marattiales) from North
Yorkshire. Rev. Palaeobot. Palynol. 51: 65-93.
Hill, C. R., Camus, J. M. 1986. Evolutionary cladistics of
marattialean ferns. Bull. Brit. Mus. (Nat. Hist.) Bot. 14:
219-300.
Hill, C. R., Wagner, R. H., EI-Khayal, A. A. 1985. Qasimia
gen. nov., an early M arattia-like fern from the Permian of
Saudi Arabia. Scr. Geol. 79: 1-50.
Holttum, R. E. 1978. The morphology and taxonomy of Angiopteris (Marattiaceae) with description of a new species.
Kew Bull. 32: 587-594.
Jenman, G. S. 1909. The ferns and fern allies of the British
West Indies and Guiana. Thnidad: Bull. Misc. Int. Bot.
Dept.
Li X., Yao Z. 1983. Fructifications of gigantopterids from
South China. Palaeontographica B, 185: 11-26.
Manton, I. 1959. Cytological information on the ferns of
West Tropical Africa; In: A. H. G., Alston, The ferns and
fern allies of West Tropical Africa. (Suppl. R W. J. Keay,
Ed.) Flora of West Tropical Africa, 2nd. edn. London,
Crown Agents, pp.75-81.
Manton, I., Sledge. W. A 1954. Observations on the cytology
and taxonomy of the pteridophyte flora of Ceylon. Philos.
Trans. R. Soc. London Ser. B. 238: 127-185.
May, L W. 1978. The economic uses and associated folklore
of ferns and fern allies. Bot. Rev. 44: 491-528.
Ninan, C. A 1956. Studies on the cytology and phylogeny of
the pteridophytes. 1. Observations on the Marattiaceae. 1.
Indian Bot. Soc. 35: 233-239.
P'an, C. H. 1936. Older Mesozoic plants from North SchensL
Palaeontologia Sinica. Ser. A 4, fasc. 2: 1-49. xv pI.
Presl, C. B. 1845. Supplementum Tentaminis Pteridographiae.
Prague, Abh. bohm. Ges. d. Wiss. V. 4.
Rosenstock, E. 1908. Filices novo-guineenses novae. Rep.
Spec. Nov. RegnL Veg. 5: 33-44 & 370-376.
Sorsa, V. 1970. Fern cytology and the radiation field. In:
Odum, H. T., Pigeon, R. F. (Eds) A tropical rainforest. Oak
Ridge, Tenn.: Office of Information Services, U. S. Atomic
Energy Commission, pp. G 39-G 50.
Stokey, A. G. 1942. Gametophytes of Marattia sambucina and
Macroglossum smithii. Bot. Gaz. (Crawfordsville) 103:
559-569.
Underwood, L M. 1902. American ferns V. A review of the
genus Danaea. Contrib. Dep. Bot. Columbia Univ. 199:
669-679.
Walker, T. G. 1966. A cytotaxonomic survey of the pteridophytes of Jamaica. Trans. R. Soc. Edinburgh 66: 169-235.
Walker, T. G. 1979. A further chromosome count for Christensenia (Marattiales). Fern Gaz. 12: 51-52.
Walker, T. G. 1981. A chromosome count for Macroglossum
(Marattiales). Fern Gaz.12: 161-162.
Walker, T. G. 1985. Cytotaxonomic studies of the ferns of
Trinidad. 2. The cytology and taxonomic implications.
Bull. Brit. Mus. (Nat. Hist.) Bot. 13: 149-249.
Wallace, J. W. 1987. The C-glycosylflavones of Archangiopteris henryi. Biochem. Syst. Ecol. 15: 529-530.
Marsileaceae
K. V.KRAMER
Marsileaceae Mirbel, Hist. Nat. Veg. 5: 126 (1802). - Pilulariaceae Wettstein, Handb. d. Syst. Bot. 2: 81 (1903).
Small or even minute ferns of periodically moist to
flooded places. Stem slender, creeping, often branching, with a radial solenostele, with little sclerenchyma
and with air canals, bearing septate hairs, often glabrescent. Leaves circinate in bud, septate-hairy when
young but often glabrescent at maturity, borne in two
dorsal rows, usually remote; petiole non-articulate,
terete, green, with one V-shaped vascular bundle.
Lamina simply pinnate, the segments two or four, so
close as to make the lamina appear palmate, or lacking. Sori borne in globose to ellipsoid, firm, at least
initially septate-hairy, stalked sporocarps with very
firm wall, these inserted on simple or sometimes
branched stalks on the lower part or the base of the
petiole or on the stem near the petiole base, dehiscing
with two valves; each sporocarp with at least two sori
each containing mega- and microsporangia. Annulus
wanting. Spores trilete, of very different size, one in
each megasporangium, many in each microsporangium. Microspores free from each other. Gametophyte
strongly reduced, not becoming free from the spore.
ANA TOMY AND MORPHOLOGY. The stele of the stem is
often reduced in small species. Air canals are usually
Marsilea
present in the cortex of rhizome and petioles. Regnellidium possesses latex ducts in its vegetative parts in
which it is unique in the ferns (Labouriau 1952).
The lamina, where present, is pseudopalmate but in
reality pinnate, with one or two pairs of strongly
crowded pinnae. The veins are forked, ± reticulate,
without free included veinlets. The stomata are borne
on both surfaces and are diaCy1ic and anomOCy1ic. The
pluricellular hairs are non-basally attached and are appressed to the epidermis.
The nature of the sporocarp has been variously interpreted, but it is now generally agreed that it represents a fertile pinna with two reflexed and connate
segments each bearing one or more often a number of
elongate sori following the veins; it has been claimed
that the ventral halves of the valves represent accessory flanges rather than the marginal parts of the segments. Abnormally developed organs intermediate between pinnae and sporocarp valves have been described.
KARYOLOGY AND HYBRIDIZATION. The known chromosome numbers of the family are all based on n = 20
or multiples thereof (up to 2 n =60), except for
Regnellidium where counts of n = 19 (besides 20) have
been made; older, deviating counts seem to be incorrect.
There is one report of a sterile hybrid in Marsilea
(Knobloch 1976). As the species in this, the principal
genus, are mostly so similar, other existing hybrids
may have been overlooked. None are mentioned in local revisions like Gupta's (1962) or Launert's (1984).
Mahlberg and Baldwin (1975) found evidence for
parthenogenesis in Regnellidium and, much less, in
Marsilea. These are the only known instances in ferns.
GAMETOPHYTE. As in all heterosporous pteridophy1es,
the gametophy1es are very much reduced. The very
rapid germination of the spores has been much remarked on, which contrasts with the enormous longevity of the sporocarps. A microspore produces spermatozoids within a day after germination. The microprothallium consists only of a few cells and does not
leave the spore. Two spermatogenous cells form 16
spermatozoids each. The megaprothallium constists of
a small prop of cells protruding from the spore and
bears one very simple archegonium with very short
neck.
ECOLOGY AND DISTRIBUTION. Most species occur in
periodically wet to flooded places, though some, especially Australian, Marsilea species are very drought-resistant. Sporocarps are apparently not produced when
the plants are submerged and the laminal parts are
floating. Upon moistening the sporocarp swells and
bursts by water uptake of the gelatinous interior, and
in Marsilea a worm-like mass of gelatine protrudes
181
outward, carrying the sori with it. Spore germination
and fertilization take place very rapidly. The enormous
longevity of the spores within their sporocarps has
been reported several times; see, e.g., Johnson (1985):
up to 100 years!
For the geographical distribution, see under the
genera.
AFFINITY. An order "Hydropteridales", still upheld by
Copeland (1947), consisting of Salviniaceae, Azollaceae, and Marsileaceae, characterized by heterospory
and the presence of sporocarps, is since long regarded
as artificial. Alliance of the Marsileaceae to Schizaeaceae has also been claimed long ago; it is based on
overall morphology, similarity in indument, etc., but it
is remote enough to call for inclusion in two different
orders.
KEY.TO THE GENERA.
1. Lamina wanting, leaves consisting only of the filiform petiole; sori 2 or 4 per sporocarp, seemingly longitudinally
3_ Pilularia
oriented in it; sporocarp single
- Lamina present; sori several per sporocarp, distinctly
transversely oriented in it; sporocarps single or several together
2
2. Lamina of four leaflets; veins usually ± reticulate; sporo1_ Marsika
carp one-celled
- Lamina of two leaflets; veins free or usually joined at their
2. Regnellidium
ends only; sporocarp two-celled
1. Marsilea L.
Fig. 94
Marsilea L., Spec. Plant. 2: 1099 (1753); Gupta (1962); Launert (1984); Johnson (1986) (often misspelled "Marsilia").
Characters as given for the family; besides: lamina of
two (very rarely, as an abnormality, three) pairs of leaflets borne very closely together, sometimes very small,
not rarely toothed at their distal margin, flabellate to
cuneate. Veins close, forked, sparingly to amply anastomosing and with joined ends. Sporocarps borne on
the petiole or on special stalks inserted at or near its
base, single to many together, very rarely on a several
times branched system of stalks (Ballard 1962), with
one or two tooth-like appendages near the insertion of
the stalk (these taxonomically important), borne at
about right angles to the stalk, subglobose to ellipsoid
or bean-shaped, the upper edge often concave, of two
connate segments, each with a row of several parallel
sori running at about right angles to the suture, each
sorus enclosed by a delicate indusium. Megaspores
spheroidal to slightly ellipsoidal, papillate, nearly
plain; microspores spheroidal, surface slightly rugulose, nearly plain.
A taxonomically not well understood genus of between 50 and 70 species which are often very similar in
appearance and plastic under varying ecological conditions; of almost world-wide distribution but lacking
in cool-temperate areas. The greatest concentration of
182
Filicat.e . Marsile.ce.e
Fig. 94 A-I. Marsileaceae. A-C Marsilea polycarpa. A Habit
of fertile plant (x 1). B Sporocarp (x4). C Lamina with venation (x 2.5). D, E Marsilea vera. D Habit on fertile plant
(x 0.7). E Sporocarp in lateral view (x 4). F Marsilea quadrifolia, longitudinal section of sporocarp (x 12). G-I Marsilea
drummondii. G Sporocarp in lateral view ( x 2). H Dehisced
sporocarp (x 3). I Dehisced sporocarp with protruded mucilaginous ring and attached sori (x 1) (A-C from Tardieu-B1ot
1957; D, E from Schelpe 1970; F from M~dalski 1954;
G-I from Wettstein 1935)
species is in Africa. The genus is absent from most
oceanic islands.
2. RegneUidium Lindman
Regnellidium Lindman, Ark. f. Bot. 3(6): 2 (1904); Johnson
and Chrysler (1938); Chrysler and Johnson (1939).
Diverging from the last by the presence of only two,
rather large, broadly flabellate pinnae; veins very
close, forked, free or joined at their ends; vegetative
parts with latex vessels. Sporocarp single, on a stalk inserted on the basal part of the petiole, spherical to
bean-shaped, two-celled, the stalk joined to it on the
upper suture, the sori at right angles to this; sori sever-
Filicatae . Matoniaceae
al (c. 6) in each cell, one each in a soral canal, with a
delicate indusium; sporangia attached along the septa
separating the soral canals. Mega- and microspores
spheroidal, bearing dense, prominent folds or verrucate elements.
.
A single species, R. diphyllum lindman, in south~rnmost Brazil and adjacent Argentina.
3. Pilularia L
Pilularia L. Spec. Plant. 2: 1100 (1753); Bonnet (1955).
Stem very slender; leaves distant, consisting only of
the filiform petiole which is circinate in bud; vascular
tissue reduced. Sporocarp single on a stalk inserted at
the petiole base, subglobose-ovoid, borne erect or inclined, attached to the stalk at its narrow edge, with
two or four sori seemingly longitudinally oriented but
not truly so (Johnson 1933); a rudimentary tooth present near the sporangium insertion. Spores spheroidal;
megaspores somewhat ornate, rugose; microspores
bearing compact folds on a coarsely papillate surface.
Between three and six species (Reed 1954), northtemperate, in western South America, and in temperate South America, Australia, and New Zealand; in
swamps and temporary pools. P. minuta Our. ex A Br.
is among the smallest of all ferns.
Selected Bibliography
Ballard, F. 1962. Marsilea botryocarpa F. Ballard. Hooker's
Icon. Plant. 3600.
Bonnet, A L. M. 1955. Contribution a I'etude des Hydropteridees I. - Recherches sur Pilularia globulifera L. et P. minuta
Dur. CeUule 57: 131-239.
Chrysler, M. A, Johnson, D. S. 1939. Spore production in
Regnellidium. BuU. Torrey Bot. Club 66: 263-279.
Gupta, K. M. 1962. Marsilea. Bot. Monogr. New Delhi 2.
Johnson, D. M. 1985. New records of longevity for Marsilea
sporocarps. Amer. Fern J. 75: 30-31.
Johnson, D. M. 1986. Systematics of the New World species
of Marsilea(Marsileaceae). Syst. Bot. Monogr. 11: 1-87.
Johnson, D. S. 1933. The curvature, symmetry and homolo·
gies of the sporocarps of Marsilia and Pilularia. Amer. J.
Bot. 25: 141-156.
Johnson, D. S., Chrysler, M. A. 1938. Structure and development of Regnellidium diphyllum. Amer. J. Bot. 25: 141-156.
Knobloch, I. W. 1976. Pteridophyte hybrids. Publ. Mus.
Mich. State Univ. BioI. Ser. 5(4): 273-352.
Labouriau, L. G. 1952. "Regnellidium diphyllum" Lindm., a
lactescent fern. Rev. Bras. BioI. 12: 181-183.
Launer!, E. 1984. A revised key to and new records of African
species of the genus Marsilea. Garcia de Orta Ser. Bot. 6:
119-140.
Mahlberg, P. G., Baldwin, M. 1975. Experimental studies on
megaspore viability, parthenogenesis and sporophyte formation in Marsilea, Pilularia, and Regnellidium. Bot. Gaz.
136: 269-273.
Reed, C. F. 1954. Index Marsileata et Salviniata Bol. Soc.
Brot. II. 28: 5-61.
183
Matoniaceae
K. U.KRAMER
Matoniaceae C. Presl, Gefassb. i. Stipes d. Farrn: 32 (1847).
Terrestrial or epilithic, medium-sized ferns with longcreeping stem densely clothed with pluricellular, uniseriate hairs. Stem containing 2 or 3 concentric steles,
the outer solenostelic. Leaves remote; petiole non-articulate, dark and polished, adaxially ± sulcate, with a
simple, marginally involute vascular bundle. Lamina
trichotomous-pedate or pseudodichotomous, catadromous, glabrous at least when mature, firm in texture;
stomata anomocytic. Ultimate divisions long and narrow, costate, with forked, free or slightly reticulate,
prominent veins. Sori dorsal on the segments, terminal
or compital, round; sporangia few, large, simultaneously maturing, arranged in a circle; indusium umbrella-shaped, centrally attached, eventually deciduous. Sporangial stalk short, massive, with a central and
a number of peripheral cells; capsule subgiobose; annulus oblique, incomplete; stornium none, dehiscence
not predefined. Spores trilete, 48-64 per sporangium..
ANATOMY AND MORPHOLOGY. The stele of Matonia
consists of three concentric solenosteles which are
connected in a complicated way by leaf traces (Bower
1926, Fig. 498, etc.). The thinner stem of Phanerosorus
has an inner protostele and a concentric outer solenostele.
The vascular strand of the petiole is simple, Ushaped, in cross-section with involute ends. The leaf
architecture of Phanerosorus was redescribed by Kato
and Iwatsuki (1985).
GAMETOPHYTE. This was described by Stokey and Atkinson (1952) for Malonia and by Yoroi and Kato
(1987) for Phanerosorus. It is cordate, naked, with ruffled wings bearing antheridia on both sides; gametangia of both sexes ofa primitive type; the antheridia are
large, with many-celled wall and a large spermatozoid
output; the archegonium has a long, inclined neck. The
resemblance to the gametophyte of Dipteris and (less
so) of Gleicheniaceae is evident. Germination is by a
filament. Polyembryony is another primitive feature.
The differences between the two genera are small.
ECOLOGY AND DISTRIBUTION. The recent members of
the family are confined to Malesia; fossil members are
very widespread. The family is of great antiquity, going back to the Lower Mesozoic, Matonia (? peetinata)
to the Cretaceous; the present distribution is undoubtedly a relic area. The species occur very locally
on isolated tropical hills and mountains; at least one
prefers soil poor in minerals. See further under the
genera.
Filicatae . Matoniaceae
184
D
Fig.9SA-G. Matoniaceae. A-D Matoniapectinata. A Lami-
KEy
na (right-hand portion cut away) (x 0.2). B Part of fertile
pinna (x 0.5). C Base of segment with sori (x 2). D Three
different views of sporangiwn (x 50). E-G Phanerosorus sarmentosus. E Habit ( x 0.17). F Segment of leaf with inhibited
bud (X 1). G Detail of fertile segment (x 7) (A-C from Holttum 1955; D from Bower 1926; E-G from Diels 1900)
1. Lamina pedate, unevenly dichotomously branched, with
central shank; sori compital (Fig. 95 A-C).
1. MaloniD
TO THE GENERA.
- Lamina with prolongate, trailing primary rachis bearing
simple or paired primary segments with (usually dormant)
axis apices between them; sori terminal or compital
2. P/wnerosorus
(Fig. 95 E).
1. Matonia R. Brown
In view of the great antiquity it is not surprising that no closely related, recent groups are
known. Points of similarity with Gleicheniaceae and
especially Dipteridaceae are too numerous to be all
ascribed to convergence, but the relationship must be
remote in geological time.
AFFINITY.
Matonia has a number of n = 26 (Manton in Holttum 1955); Phanerosorus proved to have the
same number (Jermy, in litt.).
KARYOLOGY.
Figs. 95 A-D,96
Matonia R. Brown in WaIlich, PI. Rar. As. I: 16 (1829).
Morphology and anatomy as described above. Veins
free or slightly anastomosing in sterile segments, connivent and joining in spots midway between costa and
margin in fertile segments, the sorus borne on the
junction.
Spores trilete, tetrahedral-globose; surface bearing
an irregular, ± coalescent, granulate deposit over more
diffuse rodlets formed by the perispore. M. peetinata
Matonia . Phanerosorus
185
Fig. 96. Matoniaceae. Matonia pectinata; Selangor, Malaysia.
Phot. E. Zogg
R Br. in Wall. in Malaya, M. foxworthyi Copel. in
Borneo, doubtfully distinct.
2. Phallerosoms Copeland
Fig.95E-G
Phanerosarus Copeland, Philipp. J. Sci. 3C: 344 (1905).
Epilithic on limestone. Characters as described above.
Branching of the lamina rather as in Lygodium, but
primary pinnae may be lacking. Veins free; sori terminal, between costa and margin; or occasionally some
anastomoses present and sori compital. Spores trilete,
tetrahedral-globose; surface bearing a granulate perispore deposit or some rodlets over a verrucate surface
formed by the exospore.
Ph. sarmentosus (Baker) Copel. in Sarawak, Ph. major Diels on islands off western New Guinea and in
the southern Moluccas (Kato and Iwatsuki 1985).
Selected Bibliography
Kato, M., lwatsuki, K. 1985. Juvenile leaves and leaf ramification in Phanerosorus major (Matoniaceae). Acta Phytotax. Geobo!. 36: 139-147.
Stokey, A. G., Atkinson, L. R. 1952. The gametophyte and
young sporophyte of Malonia peelinala R. Br. Phytomorphology 2: 138-150.
Walker, T. G., Jenny, A. C. 1982. The ecology and cytology of
Phanerosorus (Matoniaceae). Pern Gaz. 12 (4): 209-214.
Yoroi, R., Kate, M. 1987. Wild gametophytes of Phanerosorus
major(Matoniaoeae). Amer. J. Bot. 74: 354-3459.
Filicatae . Metaxyaceae
186
Metaxyaceae
Stem creeping-ascending, short, stout, dorsiventral,
solenostelic, clothed with long, pluricellular, pointed,
tawny trichomes. Petioles rather close, inserted laterally and dorsally on the stem, long, stout, brown, sublerete, adaxially grooved, with long lateral pneumathodes; vascular bundle one, horseshoe-shaped and
corrugated in transection; petiole base sometimes with
adventitious buds, hairy, glabrescent. Rachis like the
petiole, glabresecent, adaxially grooved, the groove
not or only incompletely continuous with that of the
costae. Lamina rather large, simply pinnate with conform terminal pinna; lateral pinnae alternate, short-
stalked to subsessile, very elongate-oblong, subglabrous when mature, chartaceous-subcoriaceous, crenate-serrate towards the caudate-acuminate apex. Costa percurrent, adaxially sulcate; lower pinna bases
anadromous, upper ones ± distinctly catadromous.
Veins evident, close, free, simple or forked near the
base, occasionally the two branches fusing outwards,
ends clavate at the sclerotic margin. Sori dorsal on the
veins, irregularly arranged, often near the costa but also extending towards the middle, sometimes two or
even three sori borne on the same vein (unique in recent ferns), orbicular to elliptic; receptacle densely
covered with filiform, several-celled, sometimes
branched trichomes. Sporangia numerous, maturing
together, with a short, 4-seriate stalk and large capsule;
annulus nearly vertical, just bypassing the stalk:, with
distinct, several-celled stomium. Spores 64, trilete, globose; exospore of 2 layers; perispore thick:, 3-layered,
the outer layer forming a granulate deposit.
Fig. 97 A-D. Metaxyaceae. Metaxya rostrata. A Upper part
of lamina (x 0.4). B Part of stem with petiole base (x 0.5).
C Part of fertile pinna (x 3). D Sporangia (xl00)(A-C from
Proctor 1977; D from Bower 1926)
MORPHOLOGY AND ANATOMY. This was most fully described by Bower (1926), Holttum and Sen (1961), and
Lucansky (1974, 1982). The stomata are of a paracyticlaterocytic type (van Cotthem 1970; Holttum and Sen
1961). Unexpectedly, juvenile plants have much dis-
K. U.KRAMER
Metaxyaceae Pichi Sennolli, Webbia 24: 701 (1970).
A
Filicatae . Monachosoraceae
sected leaves (Roy and Holttum 1965; Tryon and Tryon 1982).
GAMETOPHYTE. This was first described by Tryon and
Tryon (1982) as cordate, with the sex organs borne
ventrally, and a rather primitive type of antheridium
with several-celled wall.
KARYOLOGY. An exact count is as yet not available,
but a number of n between 94 and 96 underlines the
distinctness from both Cyatheaceae and Lophosoriaceae.
AFFINITY. Metaxya seems closest to Lophosoria but
differs in too many points to be included in the same
family. Differences from Cyatheaceae, even Alsophila,
where it was formerly placed, are numerous and obvious. A single genus:
Metaxya C. B. Presl
Fig. 97
Metaxya C. B. Presl, Tent. Pteridogr. 59 (1836).
Amphidesmium J. Smith (1866).
Alsophiia auctt, p. p. min.
A single species, M. rostrata (H. B. K.) C. Presl, a fern of the
floor of moist forests of lower and middle elevation; tropical
America, from southern Mexico to Bolivia, the Guianas, and
Amazonian Brazil; also in Trinidad and Guadeloupe.
Selected Bibliography
Gastony, G.J., Tryon, R. M. 1976. Spore morphology in the
Cyatheaceae. II. The genera Lophosoria, Metaxya, etc.
Amer. J. Bot. 63: 738-758.
Holttum, R. E., Sen, U. 1961. Morphology and classification
of the tree ferns. Phytomorphology 11 (4): 406-420.
Lucansky, T. W. 1974. Comparative study of the nodal and
vascular anatomy in the neotropical Cyatheaceae - I. Metaxya and Lophosoria. Amer. J. Bot. 61: 464-471.
Lucansky, T. W. 1982. Anatomical studies of the neotropical
Cyatheaceae. II. Metaxya and Lophosoria. Amer. Fern J.
72: 19-25.
Lucansky, T. W., White, R.A.1974. Comparative study of the
nodal and vascular anatomy in the neotropical Cyatheaceae - III. Nodal and petiole patterns: summary and conclusions.
Morton, C. V. 1959. The correct name of the fern usually
called Amphidesmium blechnoides. Amer. Fern J. 49:
151-153.
Roy, S. K, Holttum, R. E. 1965. Cytological and morphological observations on Metaxya rostrata (H. B. K.) Presl.
Amer. Fern J. 55: 158-164.
187
~onachosoraceae
K. V.KRAMER
Monachosoraceae Ching, Acta Phytotax. Sin. 14: 17 (1978).
Dennstaedtiaceae tribe Monachosoreae (Ching) Tryon and
Tryon (1982).
Terrestrial ferns of rather small to large size with shortcreeping or ascending, dictyostelic stem. Petioles close,
non-articulate, well-developed, in certain species some
without lamina, consisting only of a short, very thick,
finger-like trophopod. Stem without scales or larger
trichomes, bearing only minute, few-celled, cylindrical,
glandular hairs, these also occurring on the foliar
parts. Lamina once pinnate to quadripinnate + pinnatifid, herbaceous; primary divisions catadromous
(Monachosorum subdigitatum) or anadromous (M.
jlagellare, M. maximowiczii). Petiole with two strapshaped vascular bundles with hippocampiform xylem,
these merging upward into one gutter-shaped bundle;
adaxial side with a single groove with slightly raised
centre, this continuous with the grooves on the secondary rachises (if present). Leaf axes bearing, beside the
minute glandular hairs, also small, scarce, reddish, articulate, non-glandular hairs and, like the laminal
parts, minute, club-like, mostly 2-celled hairs. Rachis
gemmiferous at the tip, or at the insertion of a pinna,
depending on the species, the gemma consisting of a
small rosette of finger-like trophopods. Apices of lamina, etc. of gradually simpler structure, with confluent
segments. Divisions of higher order, if present, anadromous. Ultimate segments unequal-sided, oblong-Ianceolate, crenate to deeply pinnatifid. Veins free, ending well behind the margin. Sori terminal, roundish,
exindusiate, consisting of rather few simultaneously
maturing sporangia intermingled with cylindrical, fewcelled trichomes with or without glandular end cell.
Sporangial stalk 2- or 3-rowed, bearing some fewcelled trichomes. Annulus interrupted by the stalk;
stomium well-differentiated, of 2 cells. Spores trilete,
tetrahedral-globose, with prominent angles, irregularly
tuberculate, the tubercles beset with delicate echinae.
Gametophyte (only known in M. subdigitatum) ribbon-like, lobed, the antheridia surrounding one or a
few archegonia. All chromosome counts seem to be
based on n = 56.
AFFINITY. The single genus is here placed in a family
by itself mainly because it does not fit very well in any
other family. It is most often associated with Dennstaedtiaceae (Copeland, 1947, between Microiepia and
Oenotrichia in his Pteridaceae; Tryon and Tryon 1982,
as tribe), but this seems a rather arbitrary placement,
also in view of the divergent chromosome number.
Christensen (1938) suggested affinity with Thelypteri-
188
Filicatae . Nephrolepidaceae
daceae, the evidence for which is equally weak. There
is even less support for assigning it a place in or near
Anogramma, of the Pteridaceae, as advocated by Posthumus (in Backer and Posthumus 1939) and taken up
by Nair and Sen (1974). There is some phytochemical
evidence for affinity with Dennstaedtiaceae.
A single genus:
Monachosorum Kunze
Fig. 98
Monachosorum Kunze, Bot. Z. 119 (1848), Tagawa (1937);
O. B. Nair and Sen (1974).
PtilopteTis Hance (1884), quoad typum.
Monachosorella Hayata (1927).
Six species have been described, but probably only
three are truly distinct; they range from the central Himalaya and YUnnan to Honshu, Thiwan, and Malesia.
Tagawa segregated the simply pinnate M. maximowiczii (Baker) Hayata under the genns Ptilopteris,
but the characters employed are on the specific level;
and Copeland (1947) already pointed out correctly
that M. flageYare (Maxim.) Hayata, although it is bipinnate + pinnatifid, is closer to the simply pinnate
M. maximowiczii than to the highly dissected M. subdigitatum (B1.) Kuhn. The latter is a widespread, variable species, local forms of which have been split off
as distinct species, but without sufficient justification.
Selected Bibliography
Backer, C. A., Posthumus, O. 1939. Varenflora voor Java.
Buitenzorg, Java: 's Lands Plantentuin.
Ching, R C. 1959. Flora Reipublicae Popularis Sinicae 2:
Monachosorum. Peking: Chinese Academy of Sciences,
pp. 252-256 (in Chinese).
Nair, O. B.. Sen, U. 1974. Morphology and anatomy of Monachosorum subdigitatum (BI.) Kuhn with a discussion on its
affinities. Ann. Bot. (London) 38: 749-756.
Tagawa, M. 1937. Monachosorum and PtilopteTis. Jpn. J. Bol
9: 107-120.
Nephrolepidaceae
Flg.98A-C. Monachosoraceae. A Monachosorum maximowiczii. pinnae (x 2.5). B Monachosorum flagelfare, pinnules
(x 3). C Monachosorum subdigitatum, hairs from stem,
petiole, and lamina (enlarged) (A, B from Tagawa 1937;
C from Nair and Sen 1974)
K.U.KRAMER
Nephrolepidaceae Pichi Sermolli, Webbia 29: 8 (1974).
Terrestrial, epilithic, or epiphytic ferns; stern erect or
short-creeping, rarely elongate, with a radial dictyostele, bearing peltate scales with few-celled marginal
hairs, projecting teeth, and caducous superficial hairs.
Protostelic runners often present, their stele turning in-
Nephrolepis
189
to a dictyostele as they fonn adventitious buds; in a
few species they also bear tubers. Petioles close, nonarticulate, sometimes very short, with two larger ventral and a few smaller dorsal bundles arranged in a Ushape, adaxial side flattened to sulcate, not dark-sclerotic, without pneumatophores, sometimes persistently
hairy and/or scaly. Lamina long and narrow, simply
pinnate, with numerous, often close pinnae, the apex
maturing slowly, pinnatifid; rachis like the petiole,
adaxially grooved. Pinnae sessile, articulate at base, inserted on the ridges bordering the rachis groove, often
lanceolate, with adaxially grooved, sometimes scaly
and/or hairy, percurrent costa; base of pinnae unequal, the anterior side often auricled. Fertile pinnae
occasionally somewhat diffonn, more incised and/or
narrower than the sterile pinnae. Veins free, simple or
forked, or pinnate in the basal auricle; venation ±
isodromous. Vein ends bearing hydathodes behind the
margin; pinna margin entire, or lobed to pinnatifid to
the middle. Stomata polocytic or monocyclic-staurocytie. Surface of pinnae sometimes persistently hairy.
Basal pinnae not rarely reduced; upper part of lamina
very gradually narrowed to the tip. Sori tenninal on
acroscopic branches of veins (groups), medial to submarginal, usually round or oval-renifonn, sometimes
placed in the lobes near their apices, in one species linear and occupying mimy vein ends. Indusium renifonn to suborbicular-subpeltate, linear in one species;
no sterile appendages among the sporangia. Spores
monolete, ellipsoidal or spheroidal, surface irregularly
low-tuberculate to rugose, sometimes with spherical
deposit.
ANATOMY AND MORPHOLOGY. The long, protostelic
runners are present in many species; such organs are
otherwise rare in ferns. The tubers borne on them in
species like N. occidentalis Kunze, N. cordifolia (L.)
C. Presl, etc., are unique; their function was described
by Heinricher (1907).
Certain cultivars have more than once pinnate
leaves, leaves with forked rachises ("N. duffii'), etc.;
they are, or were, popular as ornamental plants. For
general accounts of the anatomy and morphology, see
Nayar and Bajpai (1976) and Sen and Sen (1973).
GAMETOPHYTE. See Atkinson (1973) and Sen and Sen
(1973). The prothallium is cordate and often bears
several-celled hairs; the antheridia may be borne on
short stalks.
KARYOLOGY AND HYBRIDIZATION. The base number is
n -41, as mostly in Oleandraceae and often in Dryopteridaceae. Tetraploids are known (Walker 1966). Hybrids may have largely gone undetected; one was reported by Nauman (1979).
ECOLOGV AND DISTRIBUTION. Nephrolepis is one of the
few genera where one and the same species may occur
Fig. 99A-C. Nephrolepidaceae. A, B Nephro/epis undulala.
A Stem with tuber and leaf (x 0.7). B Fertile pinna, abaxial
side (x 2). C Nephrolepis acuti/olia. abaxial side of fertile pinna (x 2) (Schelpe 1970)
190
Filicatae . Oleandraceae
terrestrially or epiphytically. Most species grow in
thickets, on rocks, and in similar, somewhat open
places; others are pioneers on lava flows. Some species prefer very acid-humose habitats like peat swamps
or leaf bases of palms with litter accumulations. N. delicatula (Dcne.) Pichi Serm. and N. occidentalis Kunze
are said to be annuals (Pichi Sermolli 1969; Tryon and
Tryon 1982), a very rare phenomenon in ferns apart
from the genus Anogramma; but they reproduce vegetatively by runners.
AFFINITV. Nephrolepis is usually placed in the Davalliaceae, or, when that family is subdivided, into Oleandraceae, where it is closely associated with Arthropteris
which may be very similar in appearance. Anatomical
and spore characters, however, are so divergent that
the genus is better placed in a special family.
Monotypic.
Nephrokpis Schott
Fig. 99
Nephrolepis Schott, Gen. Fil. pI. 3 (1834); Morton (1958);
Copeland (1940); Tryon (1964); Pichi Sennolli (1969).
Lindsayoides Nakai (1943).
About 30 species whose taxonomy is partly very incompletely understood; the genus is greatly in need of
a modem revision. It is distributed throughout the
warmer parts of the world; the greatest species concentration is in SE Asia. Certain species have been introduced into new areas through cultivation.
Selected Bibliography
Copeland, E. B. 1940. Oleandrid ferns (Davalliaceae) of
New Guinea. Philipp. J. Sci. 73: 345-357.
Heinricher, E. 1907. Zur Kenntnis der Farngattung Nephro/epis. Flora 97: 43-74.
Morton, C. V. 1958. Observations on cultivated ferns, V. The
species and fonns of Nephrolepis. Amer. Fern J. 48: 18-27.
Nauman, C. E. 1979. A new Nephrolepis hybrid from Florida.
Amer. Fern J. 69: 65-70.
Nayar, B. K., Bajpai, N. 1976. Morphology in relation to phylogeny of the Davallioid-Oleandroid group of ferns. Phytomorphology 26: 333-354.
Pichi Sennolli, R. E. G. 1969. Taxonomical notes on Nephrolepis cordifolia (L) Presl and related species. Ann. Mus.
Civ. Stor. Nat. Genova 77: 270-277.
Sen, U., Sen, T. 1973. Anatomical relationships between the
Oleandra and Nephrolepis groups. Bot. J. Linn. Soc. 67
Suppl. 1: 155- 172.
Tryon, R. M. 1964. The ferns of Peru. Polypodiaceae (Dennstaedtieae to Oleandreae). Contrib. Gray Herb. 94; NephroIepis, pp.225-234.
Walker, T. G. 1966. A cytotaxonomic survey of the pteridophytes of Jamaica. Trans. R. Soc. Edinburgh 66 (9):
169-237.
o leandraceae
K. UKRAMER
OIeandraceae (J. Smith) Ching ex Pichi Senn., Webbia 20:
745 (1965).
Oleandreae J. Smith (1866).
Terrestrial, epilithic, or often scandent or epiphytic
ferns; stem elongate, with a perforated dictyostele and
bearing peltate scales. Lamina simple, articulate to
phyllopodia, or pinnate, the pinnae articulate to the
rachis, or the petiole also articulate; vasculature of
petiole consisting of two larger, ventral and a number
of smaller, dorsal bundles, in transection arranged in
about a semi-circle. Petioles usually remote, adaxially
flattened or unisulcate, or virtually absent. Lamina
simple or once pinnate; leaf apex various. Sori terminal on the veins, less often dorsal, nearly always
round, exindusiate or with a reniform to suborbicular
(but not peltate) indusium. Spores monolete, with
strong sculpture derived from the perispore.
ANATOMY AND MORPHOLOGY. Oleandra is better
known than Arthropteris, and no information is available on Psammiosorus; see Ogura (1938), Nayar et al.
(1968), Nayar and Bajpai (1976), and Sen and Sen
(1973). The two genera are fairly similar. The phyllopodia agree anatomically with the petiole. Hydathodes
are present at the vein ends. The stomata are polocytic,
in Oleandra also anomocytic. Certain species of
Oleandra have erect stem branches with aggregated
leaves; much-branched plants thus form thickets. Such
a shrubby habit is almost unique in ferns. For the
structure of the roots see Wetter (1951).
GAMETOPHYTE. This is known for a few species only;
see Nayar et aI. (1968), Atkinson (1973), also Stokey
(1960). In Oleandra it is cordate, often hairy, and
sometimes bears short-stalked antheridia; in Arthropteris it is more variable, cordate to strap-shaped, also
hairy.
KARYOLOGY AND HYBRIDIZATION. A base number of
n =41 has been found in the two principal genera, in
Oleandra also n = 40. A tetraploid species of Oleandra
was reported by Manton and Sledge (1954). Hybrids
have apparently not been found so far.
AFFINITY. Oleandraceae have been variously associated with, or included in, Davalliaceae and Dryopteridaceae. Like Nephrolepidaceae, they seem to be related
to both and may be close to the common source of the
two families. Nephrolepis is excluded from the family
because of divergent anatomy and spore sculpture.
Oleandra . Arthropteris . Psammiosorus
KEY TO THE GENERA.
Lamina simple; sari dorsal or subterminal
1.
2.
-
Lamina simply pinnate; sari various
Sori terminal, indusiate or not
Sori dorsal, exindusiate (Madagascar only)
1. Oleandra
2
2. Arthl'Opteris
3. Psammiosoms
1. Oleandra Cavanilles
Fig. 100C-F, 101
Oleandra Cavanilles, Anal. Hist. Nat. 1: 115 (1799); Maxon
(1914); Copeland (1940); Scamman (1961); Pichi Sermolli
(1965); Sen and Sen (1973).
191
Fig.100A-F. Oleandraceae. A, B Arthropteris monocarpa.
A Stem with phyllopodia and leaf (x 0.7). B Fertile pinna,
abaxial side (x2). C-F Oleandra wallichii. C Habit(xO.3).
D Scale of stem (x 14). E Portion of lamina along costa with
sori (x3). F Edge of lamina (x3) (A, B from Schelpe 1970;
C-F from Flora Taiwan 1, 1975)
192
Filicat.e . Ole.ndrace.e
apically triseriate, often bearing short lateral hairs;
stomium of two well-differentiated cells. Spores monolete, ellipsoidal, surface bearing prominent wing-like
folds, these usually echinate, often perforate or reticulate; or with coarse, low-rugose surface.
The c. 40 species are pantropic in distribution, extending only locally into warm-temperate areas, e. g.,
in South Africa and the Himalaya; the genus reaches
eastern Polynesia but is poorly represented in Africa
and its islands. Species of Oleandra occur in forests,
where they are often scandent or even epiphytic;
"shrubby" species often form large stands in more
open places. Some species ascend to considerable altitude on tropical mountains.
A formal subdivision of the genus has not been given. Many species are similar in appearance, and a
modem monograph is required.
2, Arthropteris J. Smith in Hooker f.
Fig.100A, B
Arthropteris J. Smith in Hooker r., Fl. New Zeal. 2: 53 (1854);
Holttum (1966)
Fig. 101. Ole.ndr.ce.e. Oleandra dislenla, "shrubby" habit;
Mt. Kenya. Pho!. K. U. Kramer
Stem terrestrial or epilithic, sometimes with erect,
shrub-like branches, or scandent to epiphytic, its scales
hair-pointed, appressed or squarrose, bearing marginal
and superficial hairs. Leaves articulated to phyllopodia which are short and knob-like to several em long,
the articulation often evident as a swelling; leaves
borne on all sides of the stem or in creeping species
only on the dorsal side, sometimes aggregated into
pseudowhorls; petioles almost wanting to well-developed, hairy andlor scaly, g1abrescent, adaxially sulcate
at least near apex, often winged by the decurrent blade
base. Lamina simple, elongate-ovate to narrowly lanceolate, entire, often firm in texture, in two species
contracted in fertile leaves, abaxially deciduously or
persistently hairy andlor scaly, bearing non-peltate
scales, especially along the percurrent, adaxially sulcate or flattened costa. Veins close, issuing at about
right angles, forked (often near the base), free or with
occasional, irregular anastomoses, with thickened, hydathode-bearing ends behind the margin. Sori single
on acroscopic vein branches, dorsal or rarely subterminal, often forming an irregular line parallel to the costa, round; indusium reniform to hippocrepiform-suborbicular, attached by a broad to narrow sinus;
sporangia mixed with hairs, long-stalked, the stalk bi-,
Stem long-creeping, terrestrial, epilithic, or scandent
(hemi-epiphytic); stele dorsiventral, with few meristeles; scales with toothed margin and deciduous
hairs. Leaves borne on the (Iatero) dorsal side, articulated to short to elongate, two-ranked phyllopodia
(Fig. 100 A). Petiole short to long, adaxially flattened
to sulcate, deciduously scaly, at least initially pubescent with several-celled, partly glandular hairs; 3 vascular bundles. Lamina small to medium-sized, pinnate
or pinnate + pinnatifid, with pinnatifid apex or a
(sub)conform terminal pinna; pinnae (sub)sessile, articulate to the adaxially sulcate rachis, inserted on or
just beside the edge of the groove, in the latter case
two additional, ventro lateral grooves present. Pinnae
elliptic to lanceolate, often basally asymmetric with
the posterior side narrower, entire to pinnatifid,
glabrous or pubescent, with percurrent, adaxially convex costa; venation isodromous or weakly catadromous, the veins simple or forked or the basal pinnate,
ending in hydathodes behind the margin, sometimes
with white pellets overlying the hydathodes. Sori medial or supramedial, terminal on an acroscopic vein
branch, round, with reniform indusium or exindusiate;
sporangia mixed with hairs. Spores monolete, ellipsoidal to spheroidal; surface bearing irregular, winglike or cristate folds with papillate or echinate elements.
Between 12 and 15 species, paleotropical to southtemperate, from tropical and South Africa and tropical
Arabia to the Mascarenes, and from Ceylon, Burma,
and the Ryukyus to Australia, New Zealand, and eastern Polynesia; absent from the American continent,
but one endemic species in the Juan Fernandez Is-
Filicatae . Ophioglossaceae
lands. The greatest concentrations of species are in
Madagascar and in New Guinea. No formal subdivision has been given.
193
Ophioglossaceae
W. H. WAGNER, JR.
3. I'sIlInmiosorus C. Christ.
Psammiosorus C. Christ., Dansk Bot. Ark. 7: 73 (1932).
Much resembling species of Arthropteris with entire
pinnae. Petioles articulate at base. Lamina simply pinnate with conform terminal pinna, all pinnae articulate
at base; pinnae sessile, lanceolate, sinuate; lower pinnae slightly shortened. Veins pinnate, sometimes sparingly anastomosing in pairs, without free included
veinlets. Sari dorsal, medial, exindusiate, without appendages among the sporangia. Spores monolete, ellipsoidal, bearing compressed folds forming an irregular, wing-like, more or less perforate, echinate surface.
A little known genus of one species, Ps. paucivenius
(C. Chr.) C. Chr., confined to NW Madagascar, collected three times.
Selected Bibliography
Copeland, E. B. 1940. Oleandrid ferns (Davalliaceae) of
New Guinea. Philipp. J. Sci. 73 (3): 345-357.
Holttum, R. E. 1966. The genus Arthropteris. J. Sm. in Malesia. Blumea 14: 225-229.
Manton, I., Sledge, W. A. 1954. Observations on the cytology
and taxonomy of the peridophyte flora of Ceylon. Philosoph. Trans. Roy. Soc. London B 238: 127-185.
Maxon, W. R. 1914. The American species of Oleandra. Contrib. U. S. Nat. Herb. 17: 392-398.
Nayar, B. K., Bajpai, N. 1976 (1978?). Morphology in relation
to phylogeny of the Davallioid-Oleandroid group of ferns.
Phytomorphology 26: 333-354.
Nayar, B. K., Bajpai, N., Chandra, S. 1968. Contributions to
the morphology of the fern genus Ofeandra. Bot. J. linn.
Soc. 60: 265-282.
Ogura, Y. 1938. Anatomy and morphology of OIeandra Waflichii (Hk.) Pr., with some notes on the affinities of the genus OIeandra. Jpn. J. Bot. 9: 193-211.
Pichi Sermolli, R. E. G. 1965. Adumbratio florae aethiopicae
11. Oleandraceae. Webbia20: 745-769.
Scamman, E. 1961. The genus OIeandra of Costa Rica. Rhodora 63: 335-340.
Sen, U., Sen, T. 1973. Anatomical relationships between the
OIeandra and Nephro/epis groups. Bot. J. Linn. Soc. 67
Suppl. 1: 155-172.
Stokey, A. G. 1960. Multicellular and branched hairs on the
fern gametophyte. Amer. Fern J. 50: 78-87.
Wetter, C. 1951. Dber die Luftwurzeln von OIeandra. P1anta
39: 471-475.
Ophioglossaceae (R. Br.) Agardh, Aphor. Bot. 8: 113 (1822),
based on: Ophioglosseae (tribe) R. Brown, Prodr. Fl. Nov.
Hall. 163 (1827). For synonyms see below.
Mainly terrestrial herbs, some epiphytic, usually small
and fleshy, laCking sclerenchyma. Stem mostly short,
subterranean, upright, rarely horizontal, the stele with
collateral vascular tissue and resembling a eustele, the
xylem mainly of radially aligned tracheids with circularly bordered pits or in some scalariformly bordered
pits. Roots laCking root hairs, mycorrhizal, some with
well-developed cork layer; the stele 2-S-arch. Leaf
bases dilated and enclosing 2-5 primordia, the oldest
enclosing the next oldest and so on. Primordial leaf
blade usually conduplicate, not circinate. Epidennal
surfaces glabrous or provided with uniseriate hairs.
Mature leaves usually one per stem per growing season, in some taxa more, usually divided into a bladelike sterile photosynthetic portion (trophophore) and a
centrally attached skeletonized spore-bearing portion
(sporophore), the latter commonly aborted. Trophophore absent in several species. Common stalk
glabrous to hairy. Trophophore divided into segments,
deltoid to oblong and free-veined, or simple-spathulate to trullate to linear (rarely palmatifid), and veins
simple-reticulate (without included veinlets) or complex-reticulate (free veinlets and!or areolets within
primary areoles). Segment margins lacerate, dentate,
crenate, or entire. Lamina shiny or glaucescent. Stomata anisocytic. Sporophore single and arising at base of
trophophore stalk, at base of trophophore blade, or on
the trophophore rachis, or rarely plural and arising
from top of petiole and base of blade. Sporangia marginal or terminal, globose, massive with thick wall and
containing thousands of spores, non-annulate, dehiscing along a single lateral or vertical line, either exposed on branches of sporophore or deeply sunken in
two rows on the sides of a simple terminal "spike".
Spores tetrahedral-trilete, mainly coarsely or finely
verrucate. Gametophytes subterranean, mycotrophic,
fleshy.
ANATOMY AND MORPHOWGY. Bierhorst (1971) described the structural features in detail. The sporophore is the most distinctive feature of the family. Always stalked, it arises upon the trophophore axis and
dies after discharging spores. Its homologies are not
fully understood, but it is commonly believed to be the
fusion product of the two basal pinnae (analogous to
the fertile basal pinnae of Anemia, but with fusion).
Lower pinnae of trophophores in Botrychium commonly produce scattered sporangia, and part or all of
194
Filicatae . Ophioglossaceae
Fig.102A-G. Ophioglossaceae. A-C Botrychium austral•.
A Habit, fertile plant (x 0.5). B Part of lamina (x 1.2).
C Part of sporophore (x 3.5). D Ophioglossum palmalum.
fertile leaf (x 0.25). E-G Ophioglossum lusilanicum. E Habit,
fertile plant (x 2). F Lamina with venation (x 3). G Fertile
sporophore (x 4) (Duncan and Isaac 1986)
the sporophore may be transformed into sterile tissue.
For the sterile lamina portion the term trophophore is
recommended rather than trophophyll, because it is
believed that the ophioglossaceous leaf (Phyll) is actually made up of two parts, as judged by ontogenetic
and anatomical evidence. The unique leaf blade of
Ophioglossum appears to be phyllodial and to result
from loss of the morphological blade and retention of
only the axis which becomes expanded laterally together with elaboration of the vein system (Fig. 102).
In some cases, diagnostic structures of critical species
can be obliterated by dwarfism as in Ophioglossum ellipticum Hook. & Grev. (Wagner et al. 1984b). Among
the rare structures are, in Botrychium, the tracheidal
idioblasts of the major leaf axes of B. lunarioides
(Michx.) Sw. (Arnott 1960), the absence of trophophore in B. paradoxum Wagner (Wagner et al. 1985),
the clusters of brood bodies on stems of B. campestre
Wagner, and the production of a sporophore on the
trophophore rachis in B. lanuginosum (Wall.) Hook. &
Grev.; in Helminthostachys the creeping stem and cylindrical sporophore with sterile projections; in Ophioglossum the bulbous stems and gametophytes of O.
crotalophoroides Walter (Mesler 1973), the basally attached sporophore in O. bergianum Schlecht., the loss
of tropophore in O. kawamurae Tagawa and certain
other species (Sahashi 1980), and the palmatifid trophophore and multiple sporophore branches in O. palmalUm L.
GAMETQPHYTE. The prothallia are subterranean,
achlorophyllous, whitish to brownish, fleshy, subspherical, cylindrical, or cushion-shaped (BotrychiumJ
or cylindrical and linear, unbranched or branched
(OphioglossumJ. All have simple hairs on the surfaces.
Their nutrition is mycoparastic, and one may find
Filicatae . Ophioglossaceae
them in the wild by digging where there are young
sporophyte leaves protruding. Only recently have they
been successfully grown in axenic culture (Whittier
1976). Potentiality of gametic exchange between prothallia is evidenced by interspecific hybrids (Wagner et
al. 1985) although it has been proposed that they are
entirely selfed (St. John 1950). The gametangia are
large and sunken. Embryogenesis is diverse (Bierhorst
1971): In Ophioglossum the embryos are exoscopic,
but in Helminthostachys they are endoscopic, and in
Botrychium some species are endoscopic and others
are exoscopic. The prothallia of the Indian representatives were extensively described and figured by Pant et
al. (1984).
ECOLOGY AND DISTRIBUTION. Except for Helminthostachys, the genera are nearly cosmopolitan. The most
widespread species are probably Botrychium lunaria
(L.) Sw. of temperate and polar lands and Ophioglossum nudicaule L. f. of the tropics and warm-temperate
regions. Botrychiums are most frequent at high latitudes and altitudes, ophioglossums at low latitudes
and altitudes. Helminthostachys is restricted to low altitudes in the Australasian region. All Ophioglossaceae
(except the epiphytic ones) are associated with disturbance and especially early to middle succession. Even
those in mature forests are favoured by disturbances
(edges of pathways, old lumber roads, blow-downs,
grazing). In extreme drought the leaf primordia may
remain dormant and not appear until normal moisture
returns. Depending on appropriate periods of rainfall,
some ophioglossums may come up two or more times
per year. The plants are gregarious and grow, often
several species together, in genus communities (Wagner and Wagner 1983). The best habitats for both botrychiums and ophioglossums are open pastures, old
fields, young second-growth woods, and grassy slopes.
Epiphytic ophioglossums tend to grow in rotten soil of
root masses of ferns, bromeliads, and leaf bases of
palms. Many of the species seem to be very rare, although they are possibly overlooked because they resemble young shoots or juvenile leaves of flowering
plants. Many of the species are poorly collected therefore and are represented by few, often inadequate,
samples in herbaria.
AFFINITY. There is little indication of the precise relationships of Ophioglossaceae. Of modern plants normally classified as "ferns," the Ophioglossaceae are
the most isolated. Such characteristics as upright
stems, dilated sheathing leaf bases, conduplicate leaf
primordia, collateral leaf traces, eustele-like vascular
organization, large circular bordered pits, cork cambium, massive eusporangia, thick parenchymatous gametophytes with large sunken gametangia, and endoscopic embryos suggest closer affinities to progymnosperms or cycadophytes than to typical modern ferns.
195
However, there are no convincing fossils to back up
speculations about the nearest relatives. Among living
plants the Ophioglossaceae seem to lie phenetically
roughly between the Cycadales and the Marattiales,
but the presence of the sporophore is unique and not
found elsewhere in present-day fern-like plants.
SUBDIVISION. As treated here, the Ophioglossaceae are
perhaps "underclassified." The family as a whole is
relatively small, and the characters used in their taxonomy are unfamiliar to most taxonomists working with
typical ferns and flowering plants. In keeping with the
hierarchical inflation in other groups of vascular
plants that is current today, a greater amount of splitting may be called for, as illustrated by the following:
Botrychiaceae Nakai
Botrychioideae
Botrychium Sw.
Sceptridium Lyon
BotrypWi Michx.
Japonobotrychium Masamune
Helminthostachyoideae
Helminthostachys Kaulf.
OphiogJossaceae (R. Brown) Agardh
Ophioglossoideae
Ophioglossum L.
Ophioderma (Blume) End!.
Cheiroglossa C. Presl
Some recent authors (e. g., Pichi Sermolli 1977) have
even more extreme taxonomic arrangements, but for
the present it is probably best to retain the conservative classification given below. Kato's novel classification (1987) that divides the family into six equivalent
genera arranged from most primitive to most advanced (viz. Botrypus, Japanobotrychium, Sceptridium,
Helminthostachys, Botrychium, and Ophioglossum) is
based on some character analyses that are questionable or erroneous. In particular this system ignores the
numerous autapomorphies that separate Helminthostachys from Ophioglossum and these two respectively
from Botrychium. Clausen's (1938) monograph of the
family was conservative at the higher levels, but split
extensively at the infraspecific level, basing varieties
and subspecies on trivial characters such as plant
size.
KARYOLOGY AND HYBRIDIZATION. The base numbers
of chromosomes in the Botrychium-Helminthostachys
group are x=44, 45, 46, and 47; x=45 seems to be the
most common number in Botrychium. Tetraploids
(n = 90, 92, 94) are common, but there are only a few
instances of hexaploids and octoploids (n = 135, 180).
The base number for Ophioglossum is evidently x = 30;
present polyploids range from n = 120 to n = c. 660,
the latter the highest known number in all plants.
Twelve examples of sterile hybrids are known involving species of Botrychium where they occur together
(Sahashi 1982; Wagner et al. 1985). The hybrids are in-
196
Filicatae . Ophioglossaceae
termediate between their parental species and display
abortive spores and, where known, irregular meiosis.
KEy TO THE GENERA.
1. Sporophore pinnately or radially branched; sporangia ex-
posed on branchlet tips, not sunken; trophophore mostly
pinnately lobed or divided, veins free
2
- Sporophore simple, unbranched (except in subgenus Cheirog/ossa); sporangia sunken in two rows on the sides of a
single linear spike; trophophore mostly unlobed or divided, veins reticulate
3. Ophioglossum
2. Sporophore pinnately branched, mostly in one plane: stem
upright; sporangia solitary and lacking associated sterile
projections
1. Botrychium
- Sporophore radially branched, cylindrical; stem horizontal; sporangia more or less clustered in clumps of several,
associated with sterile projections
2. He/minthostatiJys
abortive; leaf primordia hairy; n =45, 90, 135. Approximately 25-30 species, both hemispheres, but the
majority of species found in eastern North America
and eastern Asia, especially Japan, where studies by
Sahashi have revealed several new species and hybrids.
Subgenus Botryehium: Leaf sheath closed: plants
mostly inconspicuous and of small stature, deciduous;
blade ternate to oblong, fleshy, absent in B. paradoxum; sporophore always developed (exceptions very
rare); leaf primordia glabrous; n = 45, 90, 135. Approximately 15-25 species, both hemispheres, concentrated in central and western North America, where 12
new species were recently discovered.
2. Helminthostachys Kaulfuss
1_ Botrychium Sw.
Fig.102A-C
Bolrychium Sw.• Schrad. J. 1800: 110 (1801).
BOlrypus Michaux (1803).
Sceptridium Lyon (1905).
Japanobotrychium Masamune (1931).
Stem fleshy, up to 10 em long; leaves pinnate (rarely
simple); pinnae and pinnules linear to flabellate, freeveined; sporophore pinnate with long stalk, attached
at base of trophophore stalk, at or near blade base, or
upon the rachis; sporangia terminal on sporophore
branches. Spores trilete, globose or somewhat trilobate; exospore of three, generally distinct, continuous
layers; perispore of very fine matrix; surface shallowly
to prominently rugose or the ridges ± fused (Seeptridium), irregularly papillate to verrucate (BOlrychium),
reticulate with striate ridges (Japanobotryehium), or
with coarse, distant verrucae (Osmundopteris).
Approximately 45-55 species of temperate and polar regions and tropical mountains, concentrated in
North America and Asia, many very rare and local.
Four subgenera (raised to genera by Sahashi and
others, see above):
Subgenus Osmundopteris (Milde) Clausen: Leaf
sheath open; sporophore arising at base of trophophore blade; blade ternate, of rather thin texture;
leaves glabrescent at maturity; leaf primordia hairy;
n =92, 94. Two or three species. B. virginianum (L.)
Sw. widespread in the northern hemisphere, extends
into South America as the end of a cline [B. cicutarium
(Sav.) Sw.).
Subgenus Japanobotrychium (Masamune) Kato &
Sahashi: Leaf sheath closed, sporophore arising on rachis of trophophore blade; blade ternate, of rather thin
texture; leaves mostly hairy at maturity; leaf primordia
hairy; n = 92, 94. One or two species. Confined to
southern Asia.
Subgenus Sceptridium (Lyon) Clausen: Leaf sheath
closed; plants mostly of medium stature, mainly wintergreen; blade ternate, fleshy; sporophore commonly
He/minthoslatiJys Kaulfuss, Flora 1822: 103.
Stem horizontal; leaves pinnate to subpalmate; pinnae
linear, the lower ones coarsely divided, the segments at
base strongly excavated anteriorly and decurrent posteriorly, free-veined; sporophore cylindrical, the stalk
attached at base oftrophophore blade; sporangia radially arranged in stalked clusters interspersed with
small sterile projections. Spores trilete, globose; surface with coarse, prominent, ± fused baculae with
delicate projections; n = 94.
A single species, H. zeylaniea (L.) Hooker, widespread in Asia from India and Ceylon to South China,
Taiwan, tropical Australia, and the western Pacific, at
lower elevation.
3_ Ophioglossum L.
Fig.102D-G
Ophiog/ossum L., Spec. Plant. 2: 1062 (1753).
Ophiodenna (Blume) End/. (1836).
Rhizoglossum C. Presl (1845).
Cheiroglossa C. Presl (1845).
Stem short, upright; sporophore unbranched and
borne centrally at base of, or upon, the expanded portion of the trophophore or forming branches at the tip
of the sporophore stalk and base of blade; sporangia
sunken in tissue on either side of a single linear spike,
with a more or less developed narrow sterile apex; trophophore simple, midrib less, veins anastomosing
(sporophore absent in several taxa); plants mostly
clone-forming by root proliferations. Spores trilete,
globose, rarely monolete; exospore of 3 layers, perispore thin, fine, fibrillar, easily detached; surface irregularly reticulate with many surface granules that are
echinate in prome.
Approximately 25- 30 species. Cosmopolitan but
most common in the tropics. Three subgenera:
Subgenus Ophioglossum (Fig.102E-G): Plants terrestrial; veins with free included veinlets or areolets
within the areoles; blades (when present) linear, ob-
Filicatae • Osmundaceae
long, ovate or cordate, usually less than 10 cm long;
sporophore centrally located, one per blade. Around
20-25 species, occurring especially in open soil in
lawns and grassy fields and resembling simple leaves
of angiosperms. Sporophore commonly abortive or
absent. The Indo-Pacific species have been revised by
Wieffering (1964).
Subgenus Ophioderma (Blume) Clausen: Plants epiphytic (or terrestrial, then usually reduced); veins with
large areoles that lack included veinlets; blade oblong
to strap-shaped, sometimes reaching 2 m in length and
becoming repeatedly branched; leaf parts pendent
when epiphytic but upright when terrestrial. One to
four variable species. O. pendulum L. is widespread in
the Old World tropics and the other "species" of subg.
Ophioderma may actually be reduced forms. This and
the following taxon should perhaps be treated as genera.
Subgenus Cheiroglossa (C. Presl) Clausen (Fig.
102D): Bizarre epiphytic plants; blades when young
simple but becoming broadly fan-shaped and coarsely
palmatifid when mature; petiole bases and stem apices
.hairy; sporophore branching at juncture of petiole and
blade in a number of pendent spikes (single and central in juvenile fertile plants). One variable species of
tropical and subtropical regions of the New World,
and disjunct in Indo-China, Madagascar, and La
Reunion.
Selected Bibliography
Arnott, H.J. 1960. Tracheidal idioblasts in Botrychium. Trans.
Amer. Microsc. Soc. 78: 97-103.
Clausen, R. T. 1938. A monograph of the Ophioglossaceae.
Mem. Torrey Bot. Club 19: 1-177.
Kato, M. 1987. A phylogenetic classification of the Ophioglossaceae. Gard. Bull. Singapore 40: 1-14.
Mesler, M. R. 1973. Sexual reproduction in Ophioglossum crotalophoroides. Amer. Fern. J. 63: 28-33.
Pant, D. D., D. D. Nautiyal, D. R. Misra. 1984. Gametophytes
of Ophioglossaceae. Pob!. Phyla Monogr. (Allahabad) 1:
1-111,
Pichi Serrnolli, R. E. F. 1977. Tentamen Pteridophytorum genera in taxonomicum ordinem redigendi. Webbia 31 :
313-512.
Sahashi, N. 1980. Comparative morphology of spores of
some species in Ophioglossales. J. Jpn. Bot. 55: 73-80.
Sahashi, N. 1982. Morphological and taxonomic studies on
Botrychiaceae in Japan and adjacent regions. Doct. Thesis,
Toho Univ., Jpn.
St. John, E. P. 1950. The evolution of the Ophioglossaceae of
the eastern United States. Quart. J. A. Acad. Sci. 12:
207-218.
Wagner, W. H, Wagner, F. S. 1983. Genus communities as a
tool in the study of New World Botrychium (Ophioglossaceae). Taxon 32: 51-63.
Wagner, W. H., Wagner, F. S., HauOer, c., Emerson, J. K.
1984a. A new nothospecies of moonwort (Ophioglossaceae: Botrychium). Can. J. Bot. 62: 629-634.
Wagner, W. H., Allen, C. A., Landry, G. P. 1984b. Ophioglos-
197
sum ellipticum Hook. and Grev. in Louisiana and the taxonomy of O. nudicaule L. f. Castanea 49: 99-110.
Wagner, W. H., Wagner, F. S., Beitel, J. M. 1985. Evidence for
interspecific hybridisation in pteridophytes with subterranean mycoparasitic gametophytes. Proc. R. Soc. Edinburgh
86B: 273-281.
Whittier, D. P. 1976. Tracheids, apogamous leaves, and sporophytes in gametophytes of Botrychium dissectum. Bot. Gaz.
137: 237-241.
Wieffering, J. H. 1964. A preliminary revision of the Indo·Pacific 'pecies of Ophioglossum (Ophioglossaceae). Blumea
12: 321-337.
Osmundaceae
K.V.KRAMER
Osmundaceae Bercht. & J. C. Presl, Prirozen. Rast!. 1: 272
(1820).
Terrestrial ferns with mostly unbranched, often massive, erect, sometimes trunk-like stem clothed in roots
and persistent leaf bases, hairy at apex. Stele consisting of a ring of distinct xylem strands, these partly
conduplicate or twice conduplicate; leaf gaps small.
Petioles close, spirally arranged, non-articulate, laterally winged at the base, there bearing mucilaginous hairs
when young, with a single V-shaped vascular bundle;
sclerenchytna strongly developed. Lamina once to
twice pinnate, catadromous, when young bearing
wool-like, uniseriate hairs, these deciduous or, especially on the axes, persistent. Rachis adaxially with a
complicated system of ridges and grooves, either with
a central, medially grooved ridge on the sides of which
the pinnae are inserted, or this ridge flanked by two
further grooves and ridges, the edges of the latter then
running into the pinna edges. Bases of lateral pinnae
nearly always distinctly articulate (but the articulations
functional only in some species; see Thgawa, 1941);
pinnule bases sometimes also articulate, though less
distinctly so. Veins free, subpinnately furcate. Sporangia not assembled in sori, following the veins of not or
only slightly modified fertile segments, or entirely
covering strongly contracted fertile segments. Sporangia with a short, stout stalk and a large capsule with a
more or less developed "annulus" consisting of a
group of thickened cells on one side of the capsule,
sometimes near its base; dehiscence by a longitudinal
slit over the apex; maturation ± simultaneous; spore
output per sporangium large, over 100 to several thousand. Spores subgJobose, trilete, nearly smooth, chlorophyllous, with bipolar germination, the young gametophyte with apical rhizoid and basal meristem.
ANATOMY AND MORPHOLOGY. This was extensively described by Hewitson (1962). The stem anatomy and
Filicatae . Osmundaceae
D
A
Fig.103A-F. Osmundaceae. A, B Osmunda javanica.
A Fertile lamina (x 0.3). B Sterile leaf (xO.1). C Osmunda
lancea. dehisced sporangium (x315). D Todea barbara, apex
of fertile segment with dehisced sporangia (x 4). E LeptopteTis hymenophylloides. fertile pinnule ( x 3). F Todea barbara,
xylem of stem in cross-section (schematic) (A from Backer
and Posthumus 1939; B, C, E, F from Hewitson 1962;
D from Schelpe 1970)
the flanged petiole bases are peculiar features; so is
the disposition of the sc\erenchyma in the leaf bases:
small, scattered strands outside and usually also inside
the sclerenchymatic sheath surrounding the vascular
bundle; these characters pe1TIlit one to assign fossils to
this family with confidence (Fig. 103). They date back
to the Pe1TIlian. The stomata are of an anomocytic
type, but wanting in Leptopteris, which is peculiar in
having a "filmy" lamina structure of a few cell layers,
without mesophyll, epide1TIlis, or stomata.
GAMETOPHYTE. The prothallium is cordate to ribbonshaped, a few to several cm long, with a strong midrib,
naked, very long-lived, sometimes branched, and not
Todea . Osmunda . Leptopteris
rarely polyembryonic. Antheridia and archegonia are
mostly ventrally borne; the fonner are of a primitive
type, with many wall cells and very large spenn output. See Stokey and Atkinson (1956) and DeMaggio
(1961).
ECOLOGY AND DISTRIBUTION. Osmunda. the principal
genus, is of almost cosmopolitan distribution, with the
greatest concentration of species in East and SouthEast Asia; it is absent from cold and arid areas and
from many islands. Leptopteris occurs in Australia,
New Zealand, and from New Guinea to Samoa and
New Caledonia. Todea is restricted to South Mrica,
Australia, New Guinea, and New Zealand. Fossils testify to the greater distribution and particularly to the
greater diversification of the family in the past, showing the present representatives to be relics; see Bobrov
(1967). These are, however, often quite numerous
where they occur, and some species are quite widespread. Many occur in sites with either high edaphic
or high atmospheric moisture (or both); the most extreme in this respect is Leptopteris with its "filmy" habit, which may, however, tolerate partial, temporary
desiccation.
AFFINITY. As is to be expected in view of its 'antiquity,
this family has no near relatives among other living
ferns. Certain points of similarity to Plagiogyriaceae
(type of hairs, winged petiole bases) may be due to a
certain measure of affinity. Some characters of the Osmundaceae are indicative of eusporangiate relationships, e. g., the sporangium which may develop from
more than one epidennal cell, and the very large spore
output.
SUBDIVISION. No recent subdivision of the family has
been proposed. Leptopteris is usually placed very near
to Todea, or even united with it (as late as 1961 by Allan), but Hennipman (1968) has pointed out that'anatomical features indicate a closer relationship of Todea
with Osmunda than with Leptopteris. At any rate, the
three genera are certainly closely related.
KARYOWGY AND HYBRIDIZATION. The basic number
of n = 22 seems to prevail throughout the family.
n=22 and 2n=44 having been found consistently in
the three genera. At present most species have been
cytologically sampled. A series of induced autopolyploids of O. rega/is was produced and described by
Manton (1950, Chap. 3).
The only hybrid known with certainty in Osmunda,
O. c/aytoniana x rega/is, was described by R M.
Tryon (1940) and re-investigated by W. H. Wagner et
al. (1978). A hybrid in Leptopteris was reported by
Brownsey (1981).
KEy TO THE GENERA.
1. Sporangia on hardly or non-contracted fertile segments, at
least initially distinctly confined to the veins
2
199
- Sporangia on strongly contracted fertile segments without
green portions, covering them entirely except over the
veins
2. Osmunda
2. Lamina coriaceous, possessing mesophyll and stomata
t. Totka
- Lamina membranous, "filmy", without stomata or mesophyll
3. Leptopreris
1. Todea Willdenow in Bemhardi
Fig.103D,F
Todea Willdenow in Bernhardi, Schrad. J. Bot. 1800 (2): 126
(1801); Hennipman (1968).
Stem erect, massive, up to c. 2 m tall. Leaves pinnate
+ pinnatifid to bipinnate, rigid, subglabrous when
mature; ultimate pinnules (segments) tongue-shaped,
serrate-crenate, adnate and decurrent. Sporangia following the veins, mostly confined to the basal pinnae
and pinnules, seemingly acrostichoid at full maturity;
fertile pinnules not or hardly contracted. Spores like
those of Osmunda.
Two species, T. barbara (L) Moore in South Mrica,
Australia, and New Zealand, and T. papuana Hennipman in New Guinea.
2.0smundaL
Fig.103A-C
Osmunda L, Spec. Plant. 2: 1063 (1753); Bohrov (1967);
Panigrahi and Dixit (1969).
Plenasium C. Presl (1836).
Osmundastrum C. Presl (1847).
Stem erect, massive, not usually elongate and trunklike; leaves simply pinnate to bipinnate, always notably catadromous; ultimate pinnules (segments)
adnate and decurrent, or articulate. Mature leaves
glabrous or ± woolly on the axes. Fertile leaves or
segments strongly contracted, without expanded, leafy
parts, bearing the sporangia everywhere except over
the veins, the sporangia often seemingly marginal by
crowding. Fertile segments basal, medial, or apical in
the lamina, or the entire lamina fertile, this depending
on the species and being a diagnostic character. Sporangia numerous, ± contiguous at full maturity; annulus cells not close to the stalk. Spores globose; surface coarsely tuberculate with fine echinate elements
fonned by fascicles of perispore strands.
A varying number of species recognized, seven by
Hewitson (1962), up to 30 by Bobrov (1967, incl. P1enasium and Osmundastrum); perhaps c. 10 sufficiently
distinct species, of subcosmopolitan distribution; one
very variable, perhaps aggregate species, O. rega/is L,
very widely distributed, the others in North America
and especially in East and South-East Asia.
The following subdivision into subgenera (raised to
generic rank by Bobrov and others) seems very natural:
Subgenus Osmunda: leaves deciduous, bipinnate,
apically fertile or dimorphic and the fertile leaves en-
200
Filicatae . Osmundaceae
tirely sporangia-bearing; principal species O. regalis
L., O. japonica Thunb., and O. lancea Thunb. (the latter two E. Asia).
Subgenus Osmundastrum (C Presl) C. Presl: Leaves
deciduous, pinnate + pinnatifid, fertile in the middle,
or dimorphic and the fertile leaves entirely sporangiabearing. O. cinnamomea L. and O. claytoniana L.,
chiefly in E. Asia and North America.
Subgenus Plenasium (C Presl) J. Smith: Leaves evergreen, rigid, simply pinnate, pinnae usually toothed;
lamina fertile at the base or in the middle. Principal
species O. javanica BI., O. bromeliifolia (C. Presl)
Copel., and O. banksiifolia (C Presl) Kuhn, all in E.
and SEAsia.
These subgenera, treated as genera by Bobrov
(1967) and others, were further subdivided by him into
section and/or series. His whole system is an example
of splitting by raising the rank of taxa, without adducing further evidence. Some of the species recognized
by him and even placed in different series are treated
by other authors as local forms of O. regalis L. and
other widespread species. Critical re-investigation is
required.
Fig. 104. Osmundaceae. LeptopteTis fraseri, part of fertile
lamina with sori; New South Wales, Australia. Phot.
K. UKramer
3. Leptopteris C Presl
Figs. 103 E, 104
LeptopteTis C. Presl, Tent. Plerid. Suppl.: 70 (1845).
Stem stout, erect, not rarely trunk-like and to c. 1 m
tall; leaves in a terminal rosette, medium-sized to
large; leaf axes often ± persistently hairy. Lamina bipinnate, with dentate to deeply pinnatifid (+ bifid)
pinnules (segments), these adnate, or, if not, articulate;
leaf tissue of a few undifferentiated layers of cells,
without epidermis or stomata. Sporangia rather few,
on the veins of unmodified segments; annulus large,
many-celled, adjacent to the sporangium stalk. Spores
like those of Osmunda.
Six species, divided among four series of Bobrov;
distribution given above.
Selected Bibliography
Allau, H. H. 1961. Flora of New Zealand; vol.l. Wellington,
New Zealand: Government Printer.
Bobrov, A E. 1967. The family Osmundaceae (R Br.) Kaulf.,
its taxonomy and geography. Bot. Zh. 52: 1600-1610 (in
Russian).
Brownsey, P. J. 1981. A biosystematic study of a wild population of LeptopteTis hybrids in New Zealaud. New Zeal. J.
Bot. 19: 343-352.
DeMaggio, A E. 1961. Morphogenetic studies on the fern
Todea barbara (L.) Moore I. Life history. Phytomorphology 11: 46-79.
.
.
Hennipmau, E. 1968. A new Todea from NewGumea WIth
remarks on the generic delimitation in recent Osmundaceae (Filices). Blumea 16: 105-108.
Hewitson, W. 1962. Comparative morphology of the Osmundaceae. Ann. Missouri Bot. Gard. 49: 57-93.
Ninan, C.A 1956. Studies on the cytology and phylogeny of
Preridophytes. III. Observations on Osmunda rega/is L. J.
Indian Bot. Soc. 35: 248-251.
Pauigrahi, G., Dixit, RD. 1969. The family Osmundaceae in
India. J. Indian Bot. Soc. 48: 90-101.
Stokey, A. G., Atkinson, L. R 1956. The gametophyte of the
Osmundaceae. Phytomorphology 6: 19-40.
Tagawa, M. 1941. Osmundaceae of Formosa. J. Jpn. Bot 17:
692-703.
Tryon, R M. 1940. An Osmunda hybrid. Amer. Fern J. 30:
65-68.
Wagner, W. G., Wagner, F. S., Miller, C. N. Jr., Wagner, D. H.
1978. New observations on the royal fern hybrid Osmunda
x ruggii. Rhodora 80: 92-106.
Filicatae • Plagiogyriaceae
Plagiogyriaceae
K. V.KRAMER
P/agiogyriaceae Bower, Ann. Bot. (London) 40: 484 (1926).
Terrestrial, medium-sized ferns with creeping or more
often erect, mostly unbranched, dictyostelic stem without scales or hairs. Leaves close, spirally arranged;
petiole non-articulate, well-developed, expanded at
base, bearing two rows of knob-like aerophores on the
expansions, with a single vascular bundle that is V- or
V-shaped in cross-section, or with 3 bundles merging
above into one, in section quadrangular, trigonous, or
rarely terete. Lamina (and petiole above the base)
when young densely covered with pluricellular hairs
with swollen, glandular, mucilage-secreting end cell,
later glabrescent, pinnatifid or simply pinnate, dimorphic; rachis adaxially sulcate, often with lateral
flanges or wings; aerophores like those on the petiole
base present on the lamina of croziers, sometimes extending all the way from petiole to leaf tip. Sterile pinnae sessile or adnate, rarely short-stalked, long and
narrow, herbaceous to coriaceous, serrate-dentate at
least near the apex; costa convex above. Vpper pinnae
of sterile and fertile leaves reduced and confluent, or a
subconfonn terminal pinna present. Stomata anomocytic. Veins simple or forked, free, with slightly enlarged ends very near the margin, in the teeth, if any.
Fertile leaves stimy erect, their pinnae very narrow,
linear; veins sometimes somewhat anastomosing towards their ends. Sporangia borne on the enlarged distal parts of the veins, seemingly acrostichoid at fuU
maturity; indusium and sterile appendages among the
sporangia none but the young sporangia covered by
the somewhat reflexed leaf margin; stalk long, c.
6-seriate; capsule asymmetric, with complete, slightly
oblique annulus, the greater part thickened, the bow
interrupted by a several-celled stomium near the stalk,
dehiscence transverse. Spores trilete, tetrahedral, depressed between the angles; outer layer of bistratose
perispore forming rodlets; surface bearing coarse tubercles with papillae and rodlets.
ANATOMY AND MORPHOLOGY. The stem is very sclerotic and devoid of all indument which is very rare in
ferns. The secretory hairs were described and figured
by Hennipman (1968). Occasionally stolons spring
from a leaf base. Sclerenchyma in the leaf axes is variously developed. The mesophyll contains secretory
cells (Nayar and Kazmi 1962a). A wax-like indument
on the abaxial surface of the pinnae is present in a few
species, e. g., P. glauca (B\,) Mett. Leaves with mixed
characters of sterile and fertile laminas occur occasionally (DeVol 1972). Fertile dwarf forms of certain
species have been found; they are confusing and have
201
been described as species but do not merit taxonomic
recognition (id.).
GAMETOPHYTE. This is known in only a few species;
see Stokey and Atkinson (1956), Nayar and Kazmi
(1962b), Nayar and Kaur (1971). The prothallium is
large, thick, cordate, long-lived, with strongly developed wings, without appendages, with strong regenerating capacity. Sex organs are found on both surfaces, in
both sexes of a rather primitive type: antheridia with
many-celled wall, archegonia with thick neck and
several neck-canal cells.
KARYOLOGY AND HYBRIDIZATION. Chromosome counts
are few and divergent: n = 66 for P. semicordata
(C. Presl) Christ, P. matsumureana Makino, and P.
tuberculata Cope\., c. 132 for P. glauca (B\,) Mett., c.
75, 100, and 125, respectively, for Japanese and Taiwanese species (Kurita 1963; Tsai 1973; Lovis 1977). A
base number cannot be established with confidence.
Hybrids were described from Japan by Nakaike
(1971).
ECOLOGY AND DISTRIBUTION. Plagiogyria species grow
on the ground or sometimes on rocks, in moist places
in the hills and mountains, often up to considerable altitude. They are often locally common.
The genus is confined to tropical and warm-temperate parts of Asia and America. The greatest concentration of species is in South China. In Asia the genus occurs from eastern Himalaya to South Korea, Hokkaido, and Anhwei Prov., China, east to the Philippines
and New Guinea; it is not known from Australia or
the Pacific. In America it ranges from Mexico to Venezuela, Bolivia, and SE Brazil; also in the Greater Antilles.
AFFINITY. The species of Plagiogyria were long confused with strongly dimorphic species groups of Blechnum ("Lomaria"), a striking but merely superficial resemblance. Bower (1926) first recognized the isolated
position of the genus. Its affinities are not clear. Similarities to Osmundaceae, stressed by Bower (1926) and
van Cotthem (1970), and to Schizaeaceae, pointed out
by Ching (1958), are perhaps more indicative of a
primitive position within the leptosporangiate ferns
than of true affinity.
A single genus:
Plagiogyria ( Kunze) Mettenius
Fig. 105
P/agiogyria (Kunze) Mettenius, Abh. Senckenb. Naturf. Ges.
2: 265 (1858); Copeland (1929); Ching (1958); Nayar and
Kazmi (1962a); Lellinger (1971); De Vol (1972); Dixit and
Das (1981). Basion.: Lomaria sect. P/agiogyria Kunze,
Farrnkr. 2: 61,63 (1850).
CHARACTERS OF THE FAMILY. Ching (1958) gave a subdivision into two sections, one with 3 subsections, and
202
Filicatae . Plagiogyriaceae
Fig.105A-G. Plagiogyriaceae. A-C Plagiogyria koitlzumii.
A Habit of plant with sterile and fertile leaves (x 0.5).
B Transection of fertile pinna (x 10). C Trichomes as found
among sporangia (x 15). D Plagiogyria euphlebia, trichomes
from base of young leaf (x 20). E, F Plagiogyria distinctissi-
rna. Portion of fertile pinna with sporangia ( x 5). F Sporangia seen from two different sides (x 100). G Plagiagyria sp.,
antheridia (left) and archegonia (right) (enlarged) (A-C from
DeVol 1972; D-G from Nayar and Kazmi 1962)
Filicatae . Polypodiaceae
reported 42 Asiatic species which is too many. In the
New World there are only 6 species (Lellinger 1971) or
perhaps even only one (Tryon and Tryon 1982); they
belong to section Carinatae Ching ex Lellinger which
also includes a few Asiatic species. The species concept varies widely according to author. Few species are
truly widespread, many of quite local occurrence.
Selected Bibliography
203
Polypodiaceae
E. HENNIPMAN, P. VELDHOEN, and K. U. KRAMER; with
contributions by M. G. PRICE (Anarthropteris.
Loxogramme)
Polypodiaceae Berchtold & J. C. Presl, I'firozen. RostIin. 1:
272 (1820).
P1atyceriaceae Ching, Sunyatsenia 5: (1940).
Loxogrammaceae Pichi Serm., Webbia 29: 11 (1974).
Drynariaceae Ching, Acta Phytotax. Sin. 16: 1-37 (1978).
Ching, R. C. 1958. The fern genus Piagiogyria on the mainland of Asia. Acta Phytotax. Sin. 7: 105-154.
Usually terrestrial or epiphytic, sometimes epilithic,
Copeland, E. B. 1929. The fern genus Piagiogyria. Philipp. J.
Sci. 38: 377-417.
usually small to medium-sized, sometimes large ferns
DeVol, Ch.E. 1972. The PiagiogyriaofTaiwan. Taiwania 17:
with short- to long-creeping stem bearing basifixed,
277-292.
pseudopeltate or peltate, clathrate, opaque or hyaline
Dixit, R. D., Das, A. 1981. The family Plagiogyriaceae Bower
scales that are marginally entire, toothed, ciliate, or
in India. Proc. Indian Acad. Sci. (Plant. Sci.) 90: 371-387.
lacerate, and provided with at least a terminal glanduHennipman, E. 1968. The mucilage secreting hairs on the
lar cell; stele dorsiventral, dictyostelic. Leaves close to
young fronds of some leptosporangiate ferns. Blumea 16:
97-103.
remote, usually articulate to phyllopodia, mono- or
Kurita, S. 1963. Cytotaxonomical studies on some leptosposometimes dimorphic. Petiole variously developed,
rangiate ferns. J. Coli. Arts Sci. Chiba Univ. Nat. Sci. Ser.
sometimes winged, variously scaly or glabrous, with
4: 43-52.
several vascular bundles in a U-arrangement, the
Lellinger, D. B. 1971. The American species of Piagiogyria .
adaxial ones larger and with hooked xylem strands;
sect. Carinatae.Amer. Fern J. 61: 110-118.
adaxial face of petiole usually terete or with a median
Nakaike, T. 1971. Notes on Plagiogyria from Japan and adjagroove. Lamina simple, lobed to pinnatifid and coscent regions. Bull. Nat. Sci. Mus. (Tokyo) 14: 257-268.
Nayar, B. K., Kazmi, F. 1962a. Ferns of India - IV. Piagiotate, or simply pinnate, rarely bipinnate, pedate, or digyria. Bull. Nat. Bot. Gard. Lucknow 64: 1-37.
chotomously compound; pinnae costate, often articuNayar, B. K., Kazmi, F. 1962b. Morphology of the spores
late at base; terminal segment, if present, conform to
and prothalli of five speices of Piagiogyria. Bull. Bot. Soc.
pinnae or the lamina terminally with gradually reBengal 16: 3-8.
duced and confluent divisions; laminal hairs glanduStokey, A. G., Atkinson, L. R. 1956. The gametophytes of Plalar, usually 2- or 3-celled, sometimes otherwise; scales
giogyria giauca (BI.) Mett. and P. semicordata (Pr.) Christ.
Phytomorphology 6: 239-249.
often present, variously shaped; veins free, simple or
Tsai, J. L. 1973. Chromosome numbers of some Formosan
forked, or variously anastomosing with or without inferns (2). J. Sci. Eng. (Taichung) 10: 261-275.
cluded free veins. Fertile leaves sometimes with conWalker, T. G. 1966. A cytotaxonomic survey of the pteridotracted laminal parts. Sori usually rounded, sometimes
phytes of Jamaica. Trans. R. Soc. Edinburgh 66: 169-237.
elongate, linear, or sporangia acrostichoid, sori often
diplodesmic; sporangia short- to rather long-stalked,
the stalk mostly 3-seriate; capsule with vertical, interrupted annulus; stomium 2-celled, not continuous
with the bow; receptacular trichomes and/or scales of
varying shape often present; sporangial appendages
and sporangiasters sometimes present. Spores usually
64 per sporangium, rarely 8 (Platycerium ridleyi) or 16
(Lecanopteris p. p.), monolete, usually± bean-shaped,
rarely fusiform, very rarely trilete, white, yellow or
brown, with or without a prominent, variously shaped
perispore (Fig. 106).
MORPHOLOGY AND ANATOMY. Stem. Almost all members of the family share a perforated dictyostele; a
more complex stele occurs in the drynarioids and the
platycerioids (Ogura 1972; Schmid 1982). The number
of vascular bundles varies within the family; in some
of the species the stem possesses sclerenchyma strands
which are scattered in the ground tissue, and! or sc1e-
204
Filicatae . Polypodiaceae
C
A
Fig.106A-E. Spores of Polypodiaceae and Pteridaceae
(all x 1000). A Polypodium decumanum. B Polypodium lepidotrichum. C Solanopteris brnnei. D Pteris grandifolia. E Doryopteris pedata. Photo A. F. Tryon
renchymatic sheaths around the meristeles, or a subepidermal sclerenchymatic sheath. Leaves nearly always in two dorsal rows. Roots inserted on the ventral
side only. In Lecanopteris the stem is swollen and
ant-inhabited, in Soianopteris the stem forms tubers
which are also ant-inhabited.
Leaves. These are usually articulated to a sometimes
conspicuous phyllopodium. The shape of the leaves
does not show much variation in the family, varying
from entire to variously lobed or pinnate, rarely more
compound. Exceptions include the unique and eyecatching leaf shape and dimorphism in Piatycerium
and the leaf with internal dimorphism in part of the
drynarioids. The meristeles of the petiole are arranged
in a U-shaped configuration.
The character of the "dromy" of the venation, where
it applies at all, is usually difficult to observe. Where it
is discernible, both anadromous and catadromous patterns prove to be present, but the latter seem to be
more common. The bipinnatifid Polypodium lindenianum Kunze is catadromous. The character does not
seem to be of much relevancy in the present family.
The family is otherwise characterized by the presence of exindusiate, usually rounded to linear sori; the
arrangement of the sporangia is sometimes acrostichoid. The range of the venation pattern within the
family is very broad (Fig. 107). Beside free venation, a
great diversity of anastomosing patterns is also present, either without or with included, ex- andlor recurrent, free veins (Hetterscheid and Hennipman 1984).
The stomata are usually (co)polycytic andlor anomocytic, but more specialized stomatal types also occur (Sen and Hennipman 1981).
Indument. The scales of the stem may be basally attached, pseudopeltate, or peltate, and either opaque or
(partly) clathrate. The presence of a terminal glandular
cell is the rule. The leaf indument may include scales
similar to those of the stem or less complex ones, always including (usually 2- or 3-celled, sometimes more
complex) glandular hairs (Figs. 108-110). In the platycerioids, in which the stem is scaly, unique stellate
hairs are present on the leaves only. Receptacular trichomes, when present, show much variation (Baayen
and Hennipman 1987); deviating, non-functional
sporangia (sporangiasters) are present in a few genera,
then usually in a single species.
Spores. The greater part of the species have spores
with a thin, usually ± smooth exospore with a linear
laesura similar to that found in families such as Blechnaceae and Aspleniaceae. Spores of Polypodium are
often coarsely verrucate. A more complex exospore is
Filicatae . Polypodiaceae
205
B
B
Fig.t08A-C. Leaf indument of various Polypodiaceae.
A Pyrrosia dielsii. B Pecluma plumula. C Polypodium (Microgramma) vaccinlifolium (enlarged) (from Hennipman and
Hennipman 1987)
Fig.t07 A-D. Venation pattern of various Polypodiaceae.
A Dictymia brownii ( x 2). B Polypodium (Goniophlebium) Iwrthalsii (x 1.5). C Polypodium (Phlebodium) decumanum (x 1).
D Phymatosorus subgeminalUs ( x 2) (Original Hennipman)
present in, for instance, the lepisorioids, e. g., Belvisia,
Lepisorns proper, Drymotaenium, and Lemmaphyllum.
These spores can be seen to have the exospore
throughout traversed by a multitude of narrow canals
when studied with the transmission electron microscope. In part of the family the exospore is covered by
a (surprisingly) complex and sometimes elaborate
perispore (Fig. 106), e. g., in Pyrrosia (van Uffelen and
Hennipman 1985); in Lecanopteris (Mynnecopteris)
mirabilis a single mature sporangium contains a packet
of 16 or 32 spores each which are connected by twisted, rope-like perisporial threads (A. F. Tryon 1985;
Walker 1985). Symbiosis of polypods with ants is reported by 'fryon and Tryon (1985), Huxley (1980), and
Janzen (1974).
GAMETOPHYTE. See Nayar and Kaur (1971) and Atkinson (1973). The prothallium is usually cordate, broader
than long, with a thin to prominent midrib, with
broad, spreading wings. Unicellular, papillate, secretory hairs (both marginal and superficial) as well as
club-shaped, multicellular, branched, glandular hairs
may be present on the prothallia. In some species,
such as Lepisorns nonnalis and L. lorifonnis, the prothallia are elongate and strap-like, with notched or
cordate apex, narrow wings, and an ill-differentiated
(often interrupted) midrib. The sex organs are of the
advanced leptosporangiate type.
KARYOWGY AND HYBRIDIZATION. See Lovis (1977),
LOve et aI. (1977), Jermy and Walker (1985); a survey
of earlier literature is given in Walker (1973). Most
genera have n = 36 and! or 37. In Selliguea and Belvisia the number may be 33 or 35. Lepisorns possesses a
wide range of aneuploid chromosome numbers; 22,
23,25,26,35,36, 37, and 47. The last probably belongs
to an alloploid hybrid of plants with 22 and 25 chromosomes, respectively. Tetra-, penta-, and hexaploids
occur in some genera.
According to Wagner and Wagner (1975) hybrids
between strongly distinctive elements are rare in the
family and are of special interest as they indicate closer relationships. Some examples are: Polypodium x
206
Filicatae . Polypodiaceae
Fig. 109 A-J. Scales from leaves of various Polypodiaceae.
A Lemmaphyllum drymoglossoides. B Polypodium (Goniaphlebium) verrucosum. C Drynaria rigidula. D Po[ypodium
(P1eopeltis) percussum.
E Polypodium (Microgramma)
vacciniifolium. F Polypodium polypodioides. G Polypodium
(Pleopeltis) wiesbaurii. H Polypodium jallax. I Niphidium americanum. J Dicranoglossum desvauxii (A-D, F-H, Jx 57;
Ex 25; I X 38) (from Baayen and Hennipman 1987)
schneideri hort. is a cross between P. vulgare L. and P.
(Phlebodium) aureum L.; P. leucosporum Klotzsch is
apparently P. (Pleopeltis) lanceolatum L. X P. thyssanolepis Klotzsch; P. (P1eopeltis) lanceolatum L. var. sinuatum Sim is an inferred hybrid of P. polypodioides (L.)
Watt ssp. ecklonii (Kunze) Schelpe and P. lanceolatum
L. [Pleopeltis macrocarpa (Bory ex Willd.) Kaulf.) for
which Anthony and Schelpe (1985) erected the hybrid
genus x Pleopodium. P. x semipinnatifidum (Fee)
Mett. is apparently a cross between P. glaucophyllum
Kze. ex Klotzsch and an as yet unknown species of
Polypodium s. str. Microgramma x moraviana Gomez
is a hybrid between P. (Microgramma) lycopodioides L.
and P. plesiosorum Kunze (Gomez 1975). The existence of these hybrids militates against recognition of
Goniophlebium, Phlebodium, Pleopeltis, and Microgramma as genera distinct from Polypodium s. str.
CLASSIFICATION. The family is here taken in a restricted sense, i. e., excluding Grammitidaceae but in-
cluding Loxogrammaceae, Drynariaceae, and Platyceriaceae. Other authors (e. g. Jarrett 1980) prefer a
larger family in which Grammitidaceae are given subfamily rank.
SUBDIVISION. The subdivision of the family into lower
categories and especially genera is problematical
(Hennipman and Roos 1983). Results of recent work
On Polypodiaceae has clarified the classification of
several genera and especially those of the platycerioids
and drynarioids. However, much work remains to be
done, especially in the alliance of Polypodium sensu
lato. The classification given here is provisional and
partly artificial. It attempts as much as possible to give
taxonomic weight to morphologically distinct groups.
The subdivision adopted here is the following:
Subfamily Platycerioideae: Leaves bearing stellate
hairs. Genera 1 and 2.
SUbfamily Polypodioideae: Leaf indument of scales
and/or hairs, the latter never stellate.
Tribe Drynarieae: Leaves dimorphic; the dimorphism either (sterile) humus-collecting, so-called
nest leaves present, or often the bases of the leaves
differentiated into humus-collecting organs. Stem
scales opaque. Genera 3 and 4.
Roos (1985) mono graphed this group and reduced
the number to two; earlier, Chandra (1979-1982),
Filicatae . Polypodiaceae
207
'fL T-'"
C
having studied most representatives in detail, also
recognized Photinopteris, here included in Ag/aomorpha.
Tribe Selligueeae: No humus-collecting leaves or
leaf bases present; lamina entire or unipinnate. Perispore thick, exospore thin. Stem scales opaque.
Genera 5 and 6.
Tribe Lepisoreae: Leaves as above, usually entire.
Exospore thick, perispore thin. Stem scales clathrate. Genera 7-10.
D ;,\
Fig. 110 A-K. Sterile appendages from receptacles of various
Polypodiaceae. A Polypodium australe. B Polypodium (Mierogramma) vacciniifolium. C Solanopteris brymei. D Drynaria rigidula. E Polypodium (Pleopeltis) percussum. F Pyrrosia
pilose/loides. G Anarthropteris dietyop/eris. H Belvisia novoguineensis. I Polypodium (Goninphlebium) verrueosum. J Lepisorus obseurevenulosus. K Polypodium (Pleopeltis) anguslUm.
(Ex40;all others x 57) (from Baayen and Hennipman 1987)
208
Filicatae . Po]ypodiaceae
3. Stem swollen and with cavities or bearing tubers, ant·inhabited
4
5
- Stem never with cavities or tubers
4. Tubers present; stem bearing opaque scales (neotropical)
24. So/auopteris
- Tubers absent; stem naked Or bearing rounded, marginally hyaline scales (Asia)
12. Lecauopteris
5. Humus-collecting basal leaves or lamina bases nearly always present; nectaries present along the costa (rachis)
6
- No humus-collecting (parts of) leaves, nor nectaries present
7
6. Apical lobe of lamina usually aborted, the uppermost lateral lobe then in an almost tenninal position; abscission
Fig. ttl A-D. Polypodiaceae. A Aglaomorpha parkinsonii,
sporangium. B Niphidium americanum, sporangium. C Polypodium lachnopus, sporangium bearing appendages on pedi·
cel. D "Sporangiaster" of Polypodium virginianum. (All x 57)
(from Baayen and Hennipman 1987)
Tribe Microsoreae: Leaves as above, simple or variously dissected. Exospore thin, perispore various.
Stem scales usually clathrate. Genera 11-18.
Tribe Po\ypodieae: Leaves as above, simple or usually deeply pinnatifid to pinnate, rarely more dissected; exospore thin and smooth or thick and ± verrucate; perispore usually inconspicuous. Stem scales
usually opaque or hyaline, or sometimes ± clathrate.
Genera 19-27.
Tribe Loxograrnmeae: Leaves as above, simple, their
basal articulation usually non-functional, obscure or
not; internal sclerenchyma lacking except in the
roots. Genera 28 and 29.
ECOLOGY AND DISTRIBUTION. The members of this
predominantly tropical-subtropical family occupy a
variety of habitats. With epiphytism prevailing, still
not a few representatives regularly occur terrestrially
or on rocks. Many prefer everwet forests at low to
middle elevation, but some are also found in monsoon
vegetation and drier habitats. Some species of, e. g.,
Drynaria, Niphidium, and Polypodium occur to an altitude of about 4000 m.
The number of genera in the Old World is very
much higher than that occurring in the New World, although the number of species there is very considerable. The great majority of these belong to genera in
the tribe Polypodieae, which is only comparatively
weakly represented in the Old World.
KEy TO THE GENERA.
1. Leaves bearing stellate hairs (Fig. 108A) (nearly entirely
Old World)
2
- Leaves naked, scaly, or bearing non-stellate hairs
3
2. Leaves dimorphic, differentiated into basal ("nest") leaves
and forked foliage (soriferous or not) leaves 1. P/atycerium
- Leaves monomorphic, or, if dimorphic, no "nest leaves"
present; lamina usually entire, rarely at base pedately or
hastately lobed
2. Pyrrosia
tissue present along the costa (rachis) and between the
lobes; distinct humus-collecting leaves present (absent in
one species)
4. Drynaria
- Apical lobe of lamina not aborted; base of leaves mostly
differentiated into a humus-collecting organ; abscission
tissue only present in the sinuses between the lobes
3. Aglaomorpha
7. Sporangia assembled in sori or in elongate soral lines
("coenosori")
8
- Sporangia not assembled in sari but with acrostichoid arrangement
30
8. Sari orbicular, elliptic, or oblong
9
- Sori elongate 10 linear
28
9. Lamina subdichotomously branched, bearing marginally
clathrate, centrally hyaline scales (fig. 109J)
23. Dicrauoglossum
- Lamina simple or pectinately divided (i. e., with broadly
adnate, almost or quite free divisions)
10
10. Lamina pectinately divided
11
- Lamina simple, or, if divided, not pectinately divided 13
11. Stem short-creeping, with tufted leaves, these usually
bearing small, unicellular hairs on the abaxial side; stem
25. Peelllma
scales basally affIxed (New World)
- Stem often long-creeping; leaves without such hairs; stem
scales often peltate or pseudopeltate
12
12. Scales peltate, opaque; veins anastomosing, areoles usually without free included veinlets;
superficial; leaves
son
herbaceous (Borneo)
26. Polypodiopteris
- Petiolar and some stem scales pseudopeltate, clathrate;
veins free; sari pustulatl'-embossed; leaves thin (Asia)
27. Thylacopteris
- Stem scales clathrate or opaque, base various; veins free
or anastomosing, then with a free excurrent veinlet
springing from the arch of the costal areole; sari superficial or embossed; leaves often firmly herbaceous to coriaceous (widespread)
22. Polypodium
13. Stem scales opaque or hyaline-margined (fig. 109 F)
14
- Stem scales clathrate (Fig. 109 B)
18
14. Leaves always fully pinnate; pinnae, at least the lower,
narrowed at base, articulate; petiole adaxially sulcate
5. Arthromeris
- Leaves, if fully pinnate, without an articulation at the base
of the pinnae; or tess or not incised; petiole of pinnate
leaves adaxially non-sulcate
15
15. Scales basally attached; sclerenchyma none; lamina sim~
M
- Scales usually peltate; sclerenchyma present; lamina various
17
16. Receptacular scales dark, linear, clathrate (New Zealand)
28. Anarthropteris
- Receptacular appendages filiform or wanting; Old World
(exc. one species)
29. Loxogramme
Platycerium
17. Spores brown; leaf margin usually notched; sori often
elongate (Asia)
6. SelJiguea
- Spores yellowish; leaf margin rarely notched; sori rarely
22. Polypodium
elongate (widespread)
18. Sori in a single series of at least 5 between two adjacent
secondary veins (Fig. 119 F, 0); or leaves densely clothed
with long-ciliate scales (New World)
21. Niphidium
- Sori differently arranged; no dense indument of long-ciliate scales present
19
19. Sori, or most of them, subtenninal on free, excurrent, included veinlets, mostly 2 or 3 per areole (Fig. 119 B); lami·
na simple, rarely non-pectinately pinnate (America)
19. Campyloneurum
- Sori otherwise arranged
20
20. Young sori covered by clathrate scales
21
- Young sori not so protected
22
21. Leaves monomorphic, usually with blackish, clathrate
scales
10. Lepisorus
- Leaves monomorphic or dimorphic, naked
9. Lemmaphyllum
22. Stem scales bearing a tuft of root-like hairs on their dorsal
23
surface (Old World)
- No such hairs present on the stem scales
24
23. Leaves entire or pedately lobed; several similar main areoles formed by adjacent secondary and connecting veins
16. Neocheiropteris
- Leaves simple or (shallowly to deeply) lobed; one large
main areole formed by the primary vein, adjacent secondary veins, and the second connecting vein
18. Phymatosorus
24. Several similar main areoles, fonned by adjacent secondary and connecting veins
15. MicrosoYUm
- One large main areole formed by the primary vein, adjacent secondary veins, and the second connecting vein 25
25. Sori over 2 mm0, pustulate-embossed; venation underlying sori diplodesmic
18. Phymatosorus
- Sori under 2 mm0, usually superlicial; venation underlying sori not diplodesmic
15. Microsorum
26. Veins free, or, if anastomosing, with free excurrent veinlets
springing from the arch of the costal areoles (widespread)
22. Polypodium
- Veins anastomosing~ free included veinlets none, or costal
areoles with recurrent free included veinlets (Old World)
27
27. Free included veinlets wanting
17. Dictymia
- Costsal areoles with recurrent free veinlets
18. Phymatosorus
28. Sori elongate, linear, flanking the costa Or costules, Or at
29
an angle to the costa (Fig. 114C)
- Sori linear, "coenosorioid," parallel to the margin
36
29. Lamina coriaceous, margin often notched; stem scales
opaque; spores brown (Asia)
6. Selliguea
- Lamina herbaceous to coriaceous, margin not often
notched; stem scales with dark central band; spores yellowish (widespread)
22. Polypodium
- Lamina herbaceous, without notched margin; stem scales
clathrate; spores whitish (Asia)
13. Colysis
30. Sporangia confined to a distinct, ± contracted apical seg7. BeMsia
ment ("spike") (Fig. 115) (Old World)
- Sporangia not borne on a modified apical segment
31
31. Lamina lobed, at least at the base
32
- Lamina entire
33
32. Lamina pinnately lobed or cleft
15. Microsorum
- Lamina trilobed, or its base sagittately lobed
11. Chrisriopteris
209
33. Fertile leaves with virtually absent laminal parts, the sporangia borne on the costa (Fig. 118A) (Asia)
14. Leptochilus
- Sori borne on well-developed laminal parts, or at least not
on the costa
34
34. Secondary veins none; sclerenchyma lacking; scales basi29. Loxogramme
fixed
- Secondary veins present; sclerenchyma present; scales
(pseudo)peltate
35
35. Secondary veins immersed, evanescing not far from the
14. Leptochilus
costa (Fig. 118 A)
- Secondary veins prominent, distinct to near the margin
(Fig. 117)
13. Colysis
36. Lamina subdichotomously forked to deeply pinnatifid,
bearing few to many small scales not resembling those of
the stem (neotropical)
23. Dicranoglossum
- Lamina entire, naked or bearing scales resembling those
of the stem
37
37. Sterile vs. fertile leaf dimorphism present, or leaves with
internal dimorphism (sterile vs. fertile parts) (Old World)
38
- Leaves quite monomorphic
40
38. Dimorphism between sterile and fertile parts present;
sporangia on fertile leaves not confined to a±modified
terminal portion of the lamina (Fig. 116A)
9. LemmaphylJum
- Leaves with internal dimorphism, the sporangia confined
to a ± contracted apical portion
39
39. Sporangia assembled in orbicular sori on the contracted
terminal portion of the lamina
9. Lemmaphylillm (L. accetkns)
- Sporangia on the contracted apical "spike" in linear
"coenosori" or acrostichoid (Fig. 115)
7. Bebisia
40. Lamina narrowly linear; sporangia in two very long, linear, immersed sori
8. Drymotaenium
- Lamina lanceolate; sporangia in two ]inear, continuous or
interrupted, superficial soral lines, these confined to the
apical part of the lamina (Fig. 119D) (neotropical)
20. Neurodium
1. Platycerium Desv.
Fig. 112 A,
B
Platycerium Desv., Mem. Soc. Linn. Paris: 213 (1827); Boyer
(1964); Hoshizaki (1%4, 1970, 1972); Hennipman and
Roos (1982).
Usually epiphytic, rarely epilithic ferns with shortcreeping, stout stem bearing usually basifixed, rarely
peltate, opaque, often costate scales, their margin entire or usually bearing 1-celled or multicellular, unbranched or branched, glandular, sometimes nonglandular projections. Leaves approximate, dimorphic.
Base leaves non-articulate, usually sessile, rarely shortstalked, sterile, in most of the species forming a socalled nest or basket, in others forming an approximately globular mass of persisting old leaves. Lamina
erect or recurved, upper part erect, spreading, or appressed to the substrate, variously shaped and incised,
lower part appressed to the substrate, margin entire,
sinuate, or dentate, densely pubescent, with or without
a fringe; hairs stellate; veins anastomosing in a complex network with re- and excurrent included free
210
Filicatae . Polypodiaceae
B
Fig. 112A-F. Polypodiaceae. A Platycerium holttumii. fertile
lamina from abaxial side (x 0.1). B Platycerium elephantotis.
diplodesmic venation underlying the soral patch, the thicker
veins are the xylem of the receptacular venation (x 4). C,
D Pyrrosia subfoifuraeea. C Habit (x 0.7). D Detail of vena·
tion (x 1.5). E, F Pericytic stomata of Pyrrosia. E Pyrrosia
nummulariifolia. F Pyrrosia lingua var. heteraetis ( x 280) (A
from de Joncheere and Hennipman 1970; B from Bower
1923; C, D from Ching 1934; E, F from Nayar and Chandra
1965)
Pyrrosia • AgJaomorpha
veins. Foliage leaves articulate, short-stalked, usually
fertile. Lamina variously shaped, entire to many times
forked, erect to pendulous, usually thick-pergamentaceous, sometimes leathery; sporangia crowded in
acrostichoid patches, these usually situated terIilinally
on apices of lobes or in sinuses, in two species in the
central part of the leaves; sporangia long-stalked, stalk
(2- or) 3-seriate, annulus with about 10-22 indurated
cells; receptacular hairs stellate, in two species deviating from the lamina indument. Spores yellow (green in
P. wallichii Hooker), ellipsoidal, with short aperture
one-fourth to one-half of the spore length; exospore
thick, 2-layered, to outer plain or slightly tuberculate,
perispore conforming to exospore; surface nearly
plain, shallowly tuberculate; P. ridleyi Christ with only
8 spores per sporangium; n = 37.
Platycerium represents one of the few pantropical
genera of Polypodiaceae. It is most diversified in Mrica and Madagascar (6 species) and Asia (8 species).
One species, (P. andinum Baker) is confined to the
Andes of Peru and Bolivia. The greater part of the species occurs in seasonal dry forest, two species in lowland evergreen forests of Malesia. Ecological data
were given by Boyer (1964) and Tryon and Tryon
(1982). Several species are grown as ornamentals. The
systematic position is near Pyrrosia with which it
forms a monophyletic group.
2. Pyrl'Osia Mirbel
Figs. 108 A, 110 F, 112 C-F
Pyrrosia Mirbel, Hist. Nat. Gen. 4: 70 (1803); Shing (1983);
Hovenkamp (1986); Ravensberg and Hennipman (1986).
CycJophorus Desv. (1811).
NiphoboJus Kaulf. (1824), nom. superfl.
Drymoglossum C. Presl (1836), nom. cons.
Neoniphopsis Nakai (1928).
Saxiglossum Ching (1931).
Epiphytic or epilithic, sometimes terrestrial, small to
medium-sized ferns with short- to long-creeping stem
bearing basifixed, pseudopeltate, or peltate, round to
lanceolate, opaque scales that are marginally entire,
toothed, or ciliate, or sometimes bearing glandular
projections. Leaves approximate to remote, articulate,
mono- or dimorphic. Monomorphic and sterile leaves:
petiole absent or very short to long; lamina entire,
rarely hastately or pedately dissected, pergamentaceous to (thick-)coriaceous, often succulent and then
to 2 mm thick, usually densely set with stellate hairs;
veins anastomosing, areoles more or less rectangular
with either free, simple or forked, or anastomosing
veins, a marginal zone with a row of free, excurrent
veins usually present. Fertile leaves conform to the
sterile or contracted. Sori rounded to slightly elongate,
sometimes confluent, sometimes in rows parallel to the
margin, usually dorsal and terminal on the included
veins, rarely at the junction of tertiary veins; sporangia
211
sessile to long-stalked, annulus usually with (9-)15-22
indurated cells; receptacular hairs stellate. Spores yellow, ellipsoidal, the aperture one-third to two-thirds
the length of the spore; exospore thick, of two layers,
the outer smooth or tuberculate; perispore with coarse
spherical material on or below the surface, or coarsely
folded; surface often tuberculate to verrucate, the verrucae may be prolonged in echinate elements; folds
parallel to longitudinal axis. n =36, 37, 74, 108-111;
2n =72, c. 73, 74, c. 144, c. 149, c. 179,216.
Fifty-one species including those formerly referred
to Drymoglossum and Saxiglossum; monographed by
Hovenkamp (1986) and Ravensberg and Hennipman
(1986). Most of the species are epiphytic in lowland
and lower montane forests, others are terrestrial at low
to high elevations. The spores show a most striking
and significant diversity in perispore (van Uffelen and
Hennipman 1985). Synaptospory in the genus was
studied by van Uffelen (1985).
Chromosome number: n=36, 37, 74, 108-111;
2n =72,±73, 74, ± 144, ± 149,± 179, 216.
3. Aglaomorpha Schott
Figs. 111 A, 113
Aglaomorpha Schott, Gen. Fil.: ad tab. [191 (1936); Roos
(1985) (monogr.).
Photinopteris J. Smith (1841).
Thayeria Copel. (1906).
Merinthosorus Copel. (1911).
Drynariopsis (Copel.) C. Chr. in Verdoom (1938). based on:
Polypodium sect. Drynariopsis (Copel. (1905).
Pseudodrynaria (c. Chr.) C. Chr. in Verdoom (1938). based
on: Aglaomorpha sect. Pseudodrynaria C. Chr. (1934).
Hemistachyum (Copel.) Ching (1940). based on: Aglaomorpha
sect. Hemistachyum Copel. (1911).
Holostachyum (Copel.) Ching (1940). based on: Aglaomorpha
sect. Holostachyum Copel. (1914)
Usually epiphytic, sometimes epilithic, large to very
large ferns with long-creeping, stout, massive stem
bearing usually pseudopeltate, sometimes basifixed,
rarely peltate, commonly linear-triangular, sometimes
triangular-ovate, opaque scales that are usually marginally toothed or ciliate with 1- or 2-celled glandular
projections; stem often with a perforated polycyclic
stele. Leaves approximate or remote, non-articulate,
monomorphic, usually internally dimorphic. Petiole
absent or well-developed, usually winged, scaly. Lamina usually lobed to pinnatifid, rarely pinnate, rigidly
coriaceous, with conform terminal pinna, base of sessile leaves often dilated, shell-shaped; pinnae articulate, ovate to lanceolate, with scattered hairs to pubescent [in A. speciosa (Blume) Roos], scaly on the
primary (and secondary) veins; veins anastomosing,
often forming series of areoles between secondary
veins, usually with simple or forked free veins pointing
to all sides. Fertile pinnae usually linear to filiform,
sometimes conform to sterile pinnae, venation reduced. Sori in one to many rows between the second-
212
Filicatae • Polypodiaceae
Fig. 113. Polypodiaceae. Ag/aomorpha coronans,
habit (x 0.2) (Nayar 1961)
ary veins, or in one row on either side of the primary
vein, rounded, elongate, elliptic or quadrangular,
usually confluent, superficial to slightly embossed;
sporangia with long 3-seriate stalk, annulus with 11-16
indurated cells; sporangial appendages often present
as aciculair hairs; receptacular hairs 2-5-celled, glandular. Spores yellowish or brown, ellipsoidal, often with
obtuse ends, the aperture one-fourth to one-half the
spore length; exospore thick, 2-layered, the outer layer
usually plain; perispore conform to outer layer of exospore; surface low tuberculate to irregularly verrucate, with spheroidal deposit sometimes prolonged into short echinate elements; n -36,37.
The genus is taken in a broad sense, following Roos
(1985), including all genera recognized in the Drynaria
group apart from Drynaria, e, g., by Copeland (1947)
and Pichi Sermolli (1977), thus also including Photinopteris. The later, although possessing several unique
characters, nevertheless is shown to be close to A. pilosa (J. Smith) Cope\. and A.parkinsonii (Baker) Croxall
& Roos. The genus is confined to Asia where it deploys a remarkable diversity in leaf dimorphism; there
are 32 species.
Several species are facultative lithophytes. A. heraclea (Kunze) Cope\. and A. drynarioides (Hooker) Roos
are amongst the largest epiphytic ferns known. The
humus-collecting habit of the leaves is most conspicuously developed in A. (17Jayeria) cornucopiae (Cope\.)
Roos, the epithet being most appropriate.
4. Drynaria (Bory) J. Smith
Figs. 110 D, 114 A
Drynaria (Bory) J. Smith, Hooker's J. Bot. 3: 397 (1841); Roos
(1985), based on: Polypodium sect. Drynaria Bory, Ann.
Sci. Nat. 5: 464 (1825).
Usually epiphytic, sometimes epilithic, rarely terrestrial, medium-sized to large ferns with long-creeping
stem bearing usually peltate, rarely basifixed or pseudopeltate, commonly triangular to ovate, sometimes
linear or rounded, opaque scales that are marginally
toothed or ciliate and mostly provided with 1- or
2-celled, rarely multicellular, glandular projections;
stem sometimes with black slerenchyma strands.
Leaves approximate to remote, non-articulate, usually
dimorphic, rarely monomorphic. Base leaves wanting
in Drynaria parishii (Bedd.) Bedd. Sterile leaves sessile; lamina entire to lobed, ovate to elongate, firmly
coriaceous. Foliage leaves with well-developed, usually winged, scaly or glabrescent petiole, the wing rarely
absent. Lamina pinnatifid, rarely pinnate, apical pinna
usually aborted, rarely a conform terminal pinna present or apex pinnatifid; pinnae lanceolate or linear,
entire, usually coriaceous, sometimes herbaceous;
veins anastomosing, main areoles between adjacent
secondary veins of about similar size, with or without
simple or once-dichotomous free veins pointing to all
sides. Sori in one series on either side of the primary
vein, or in one or two rows on either side of the secondary veins; sporangia long-stalked, annulus with
Drynaria . Arthromeris
B
12-15 indurated cells; sporangial appendages mostly
absent, rarely 2-celled glandular hairs present; receptacular hairs branched or not, 2- to 10-celled, with 1-4
glandular cells, receptacular scales hair-like to complex, opaque, intermediates between hairs and scales
present. Spores yellow or brown, ellipsoidal, often
with blunt ends, the aperture one-half to two-thirds the
spore length; exospore thick, of 2 layers, the outer
plain or tuberculate; perispore thin, conforming to exospore; surface tuberculate or verrucate, sometimes irregular, usually with blunt and constricted echinate
elements, or only a spherical deposit present; n =36,
37.
A genus of about 15 species occurring in Africa and
throughout Asia to N. E. Australia. Morphologically it
is rather uniform, showing more diversity in geographical distribution and ecological preferences of its
species. It is most diversified in continental Asia, especially in China.
Fig. 114A-C. Polypodiaceae. A Drynaria propinqua. habit
with nest leaves and fertile leaf (x 0.25). B Drynaria quercifolia. part of fertile lamina with venation and sori (x 3).
C Selligueafeei. habit (x 0.3) (A, B from Nayar 1961; C from
Backer and Posthumus 1939)
5. Arthromeris (Moore) J. Smith
Arthromeris (Moore) J. Smith, His!. FiI.: 110 (1875). based on:
Pleopeltis sect. Arthromeris Moore, Index FiI. lxxviii (1857).
Terrestrial or epiphytic, medium-sized ferns with longcreeping, stout stem bearing peltate, lanceolate-attenuate, opaque, entire to shallowly toothed scales with lor 2-celled, glandular marginal projections; stem with
sclerenchyma strands. Leaves remote, articulate,
monomorphic. Petiole long, glabrous, adaxially with a
median groove. Lamina pinnate with a conform or
hastate terminal pinna; lateral pinnae articulate at
214
Filicatae . Polypodiaeeae
base, (sub)opposite, lanceolate, acuminate, finn, not
scaly, cartilaginous-margined, entire; costa percurrent;
veins anastomosing, secondary veins conspicuous,
otherwise veins immersed, areoles irregular, with simple or forked free included veins pointing to all sides.
Sori in one or less often two series between the secondary veins, rounded, compital, superficial; sporangia with long 3-seriate stalk, annulus 14- to 16-celled;
receptacular hairs 2- or 3-celled, glandular. Spores
brown, ellipsoidal, the aperture one-third to one-half
the length of the spore; exospore of two thick layers,
the outer verrucate, persispore conforming to exospore; surface (often coarsely) verrucate, with often
constricted, acuminate, or echinate elements; 11 - 36,
37; 211-72, 74, 144.
A genus of a few species, closest to Selliguea; extending from northern India east to Taiwan (and
Borneo ?). Is is unique in the shape of the perispore
which consists of hollow tubercles set with conspicuous spines.
6. Selliguea Bory
Fig. 114 C
Se/liguea Bory, Diet Class. Hist. Nat. VI: 587 (1824), non
Blume (1826) (= Colysis).
Crypsinus C. Presl (1951); Nakaike (1987).
Holcosorus T. Moore (1857).
Grammatopteridium v. A. v. R. (1924), based on: Grammatopteris v. A. v. R. (1922), nom. iIleg., non Renault (1891)
(foss.).
Pyenoloma C. Chr. (1919).
Oleandropsis Copel. (1942).
Crypsinopsis Piehi Serm. (1977).
irregularly placed, rounded, elongate, or confluent, or
sporangia in continuous or interrupted, linear bands
parallel to the secondary veins; pedicel of sporangia
long, 3-seriate; annulus with about 14 indurated cells;
receptacular hairs usually 2- to 6-celled, glandular.
Spores brown, ellipsoidal, often with blunt ends; aperature one-fourth to three-fourths the length of the
spore; exospore thick, 2-layered, the outer thin,
forming the contours; perispore conforming to exospore; surface tuberculate or verrucate, with spherical
deposit and usually densely echinate elements that are
often long and constricted, sometimes capitate; 11 = 33,
35, 36; 211 = 66, 72, 74, 144. Tetraploids have been reported.
Fifty or more species in Asia, from India eastward
to Japan and southward to New Guinea, Australia
(one species), and the Pacific; one species in South
Africa and one in Madagascar (Parris, pers. comm.);
the majority of species in New Guinea where they are
found from the lowlands to high in the mountains.
The spores always have a well-developed perispore
with a distinct, usually verruculate basal layer often
provided with small spines or baculae. As taken in its
present circumscription, the genus includes several
genera formerly segregated on the basis of only one or
a few conspicuous, deviating characters like elongate
sori (Selliguea), setaceous leaves with sunken, elongate
sori (Holcosorus), or linear, acrostichoid fertile leaves
(Pyc1loloma, Grammatopteridium). The genus is in
need of monographic study.
7. Bel.isia Mirbel
Epiphytic, small to medium·sized ferns with usually
long-creeping stem bearing peltate, elongate, lanceolate, attenuate or setaceous, opaque, entire, toothed or
ciliate scales, these mostly with 1- or 2-celled, sometimes multicellular, glandular marginal projections;
stem with conspicuous black sclerenchyma strands.
Leaves remote, articulate, monomorphic or dimorphic.
Petiole of sterile and monomorphic leaves short to
long, glabrous, sometimes scaly at base; lamina simple, ovate, rhombic, lanceolate, pinnatifid, or hastately
divided or (sub)pinnate with broadly adnate pinnae,
herbaceous to thick-coriaceous, sometimes glaucous
beneath, margin cartilaginous, usually sinuate or
notched; veins anastomosing, secondary veins mostly
conspicuous, reaching almost to the margin, interconnected by transverse lateral tertiary veins, other veins
mostly immersed, usually forming a regular series of
areoles with sometimes forked, free included veins
pointing to all sides. Fertile (parts of) lamina commonly narrower than the sterile, occasionally longer, sometimes skeletonized and virtually without laminar tissue. Sori in one series on either side of the costa, in
one or two rows between adjacent secondary veins, or
Figs. 110 H, 115
Bel.isia Mirbel, Hist. Nat. Gen. PI. 4: 65 (1803).
Hymenolepis Kaulf. (1824), nom. iIleg., non Cassini (1817);
Christensen (1929).
Epiphytic, small or medium-sized ferns with short- to
long-creeping, unbranched, ± terete stem clothed in
peltate, ovate to linear-Ianceolate, black, clathrate
scales bearing 1- or 2-celled glandular marginal projections; stem with sclerenchyma strands. Leaves close
or remote, articulate, monomorphic. Petiole short,
sometimes winged, near the stem terete, near the apex
abaxially terete, adaxially shallowly grooved. Lamina
simple, entire, lanceolate-elliptic to linear, rarely scaly,
chartaceous to coriaceous, occasionally glaucous; laminal hairs, if present, glandular, 1- to 3-celled, sometimes branched; veins anastomosing, secondary veins
prominent, the others usually immersed, areoles irregular with forked free veins pointing to all sides. Fertile
leaves apically gradually or abruptly contracted into a
mostly linear segment (spike) bearing the sporangia in
linear sori on either side of the primary vein, sometimes leaving a sterile zone along the costa and/or
Selliguea • Belvisia· Drymotaenium • Lemmaphyllum
215
margin; margin of (immature) fertile segment COmmonly ± revolute; sporangial arrangement acrostichoid; sporangial stalk long, 2· or 3-seriate, annulus
with 12-16 indurated cells; receptacular hairs, when
present, glandular, 5- to 12-celled, receptacular scales
basifixed or peltate, hyaline to clathrate. Spores white
or yellowish, ellipsoidal, the aperture one-fourth to
one-half the length of the spore; surface coarsely rugose, the rugae of irregularly fused tubercles. n = 33;
tetraploids have been reported.
About 10 species, epiphylic (often common) in the
tropics and warm-temperate parts of the Old World
from NE India to South China, NE Australia, and Polynesia, and in Mrica and its islands.
The fertile segment is most strongly offset in widely
distributed species, e. g., in B. mucronata (Fee) Copel.,
B. revoluta (Bl.) Copel., and B. spicata (L. f.) Mirbel. In
other species this is far less pronounced, e. g., in B. dura (Copel.) Copel. (restricted to Rapa) and B. platyrhynchos (Kunze) Copel. (Philippines).
Fig. 115. Poiypodiaceae. Belvisia mucranata. fertile plant;
New Guinea. Pho!. E. Zogg
8. Drymotaenium Makino
Drymotaenium Makino, Bot. Mag. (Tokyo) 15: 120 (1901).
Medium-sized epiphylic ferns with short-creeping
stem bearing peltate, lanceolate, acuminate, dark
brown, clathrate, toothed scales. Leaves approximate,
articulate, monomorphic; petiole very short, glabrous.
Lamina simple, entire, narrowly linear, coriaceous,
naked, broadly costate, with revolute margin; veins
anastomosing, forming irregular areoles with or without free included veinlets. Sori sunken, forming two
continuous lines; sporangia long-stalked, annulus with
14-16 indurated cells; receptacular hairs multicellular,
glandular, receptacular scales peltate, usually clathrate, rarely hyaline, toothed. Spores yellow, ellipsoidal,
the aperture one-half to three-fourths of the spore
length; surface shallowly rugose, the rugae irregular,
of fused tubercles; n = 36.
Monotypic; the single species, D. miyoshianum
(Makino) Makino, in Japan, China, and Taiwan; in
forests at lower to higher altitude.
9. Lemmaphyllum C. Presl
Figs. 109 A, 116 A, B
Lemmaphyllum C. Presl, Epimel. Bot.: 157 (1851); Donk
(1954).
Lepidogrammitis Ching (1940).
Weatherbya Copel. (1947).
Small epiphylic or epilithic ferns with long-creeping,
slender, branched stem bearing peltate, ovate-lanceolate, clathrate, toothed scales provided with 1- or
2-celled glandular marginal projections. Leaves re-
mote, articulate, usually dimorphic, rarely monomorphic. Sterile and monomorphic leaves: petiole absent
or short to long, sometimes winged, glabrous. Lamina
entire, rounded, elliptic, obovate, or lanceolate, naked
or rarely bearing a few scales, chartaceous to subcoriaceous or fleshy; veins anastomosing, areoles regular,
complex, with usually recurrent, sometimes excurrent
free veins, or free veins pointing to all sides. Fertile
leaves orbicular, lanceolate to linear, in L. aceedens
contracted above the sterile lower half. Sori rounded
or linear, the lines parallel to the margin, rarely interrupted; or the sporangia not assembled in sori and
acrostichoid, long-stalked; annulus with 14 or 15 indurated cells; receptacular hairs multicellular, glandular,
receptacular scales peltate or basifixed, mostly clathrate, usually toothed. Spores yellow, ellipsoidal, the
aperture one-half to three-fourths the length of the
spore; surface prominently tuberculate, the tubercles
often irregular, ± fused into rugae, n = 36.
About six species, e. g., L. carnosum (J. Smith)
C. Presl, from NE India eastward to Japan and IndoChina, and L. accedens (Blume) Donk, distributed
throughout Malesia. Common epiphyles in lowland
and montane forests, there often forming dense, rather
large colonies. Fertile leaves of dimorphic species are
held more erect than the more or less appressed sterile
ones. Sterile juvenile plants of L. accedens are common low epiphyles, whereas fertile plants occur higher
on the trees where they catch more light. In habit the
genus is not unlike Polypodium Group Microgramma.
216
Filicatae . Polypodiaceae
Fig. 116A-D. Polypodiaceae. Lemmaphyllum microphyllum.
A Habit (x 1). B Scale of stem (x 50). C Sterile leaf with
venation (x3). D Lecanopteris mirabilis. habit (b~phyllopo­
dium) (xO.17) (A, C from Flora Taiwan 1, 1975; B from
Ching 1934; D from Hagemann 1969)
Lepisorus· Christiopteris' Lecanopteris
10. Lepisorus(J. Smith) Ching
Fig.110J
Lepirorus O. Smith) Ching, Bull. Fan Mem. Inst. BioI. 4: 47
(1933); Kurata (1965); Bir and Trikha (1969, 1974); Mitui
(1971 a, b); Ching et al. (1983).
Paragramma (Blume) Moore (1957), based on: Grammitis
sect. Paragramma Blume (1828).
Pleopeltis auctt., p. p., excl. type.
Terrestrial, epilithic, or epiphytic, small to mediumsized ferns; stem short- to long-creeping, bearing peltate, ovate to lanceolate or linear, clathrate, entire or
toothed scales sometimes having 1- or 2-celled glandular projections. Stem sometimes with a wax-like
covering. Leaves approximate to remote, articulate,
monomorphic; petiole commonly short, usually
naked, rarely scaly. Lamina simple, entire, lanceolate
to linear, (sub)coriaceous, less often herbaceous, sometimes bearing clathrate scales; veins anastomosing,
areoles irregular, with many free included veinlets
pointing to all sides. Sori in one series on both sides of
the costa, sometimes submarginal, rounded, oblong, or
elongate, not rarely somewhat, occasionally strongly
embossed, rarely confluent; sporangia long-stalked,
annulus with c. 14 indurated cells; receptacular hairs,
if any, multicellular, glandular; receptacular scales all
or in part peltate, clathrate, entire. Spores yellow, ellipsoidal, the aperture one-half to three-fourths the
length of the spore; surface rugose to ± reticulate, the
rugae coarse, distant, ± fused; spherical deposit on or
below surface; n =36,72; aneuploid series have been
reported.
Widespread in the tropical and subtropical parts of
the Old World; one species in Hawaii; in Asia largely
confined to the mainland, extending north to the Russian Far East (Ussuriland). Number of species uncertain, in the absence of a monographic treatment;
about 40 according to Ching which is certainly too
many. Merged with Pleopeltis by Copeland (1947) and
Pichi Sermolli (1977), from which it differs in, among
others, leaf indument, receptacular appendages, and
stem scales. Epiphytic, epilithic, or occasionally terrestrial, in forests and open places, from low to higher altitude, to over 3000 m.
11. OIristiopteris Cope\.
Christiopteris Copel., Perkins' Fragm.: 188 (1905) ['Christopteris']; Hennipman and Hetterscheid (1984).
Rather small to medium-sized, usually epiphytic,
sometimes terrestrial ferns; stem long-creeping, bearing peltate, linear-lanceolate, opaque, entire or basally
slightly toothed scales with 2- or 3-celled, sometimes
217
branched, glandular marginal projections. Leaves remote, articulate, dimorphic. Sterile leaves with welldeveloped petiole; lamina firm, when young sometimes bearing peltate scales, trilobed, the lobes entire;
veins anastomosing, areoles regular with many
branched free included veinlets pointing to all sides.
Fertile leaves similar but with strongly contracted laminal parts. Sporangia with acrostichoid disposition;
pedicel long, 3-seriate; annulus with about 14 indurated cells; receptacular hairs 2- to 5-celled, usually
branched, glandular. Spores hyaline, ellipsoidal, the
aperture one-fourth to one-third the length of the
spore; surface shallowly tuberculate, with usually
abundant spherical deposit. Chromosome number as
yet unknown.
Two species, C. sagitta (Christ) Cope\. in the Philippines, C. tricuspis (Hooker) Christ from NE India to
Indo-China and Hainan. Both seem to be of quite local occurrence.
Perhaps the genus is better placed in the tribe Selligueeae. Pichi Sermolli (1977) assigned it an isolated
position in the family as he was incorrectly informed
about its stomata (Sen and Hennipman 1981). A third
species placed in the genus, from New Caledonia, was
transferred to Microsorum by Hennipman and Heterscheid (1984).
12. Lecanopteris Reinwardt
Fig. 116 D
Lecanopteris Reinwardt, Flora 1825: Beil. 3, 48; Jermy and
Walker (1975); Hennipman (1986): Hennipman and Verduyn (1987).
Onychium Reinwardt (1824), nOm. illeg" non Kaulf. (1820).
Myrmecophila (Christ) Nakai (1929), nom. illeg.• non Rolfe
(1917), based on: Polypodium sect. Myrmecophila Christ
(1897).
Myrmecoptem Pichi Serm., Webbia 31; 239 (1977).
Epiphytic, small to medium-sized ferns with creeping,
usually hollow, ant-inhabited, much-branched stem,
the branches closely appressed, usually covering other
branches and their roots, eventually forming a clumplike structure to over 50 cm in diameter, stem sometimes long-creeping, sometimes more sparsely
branched, naked, inconspicuously hairy, or scaly;
scales peltate, rounded, centrally clathrate, marginally
hyaline, entire, with many 1- or 2-celled glandular
marginal projections; naked stems usually sparsely to
densely set with (usually rigid) spines. Leaves remote,
articulate to a usually distinct, short phyllopodium
which is sometimes situated on flask-like swellings of
the stem, monomorphic; petiole absent or short to
long, glabrous; lamina entire, lobed, or deeply pinnatifid, or the fertile part bipinnatifid leaving a narrow
wing on either side of the primary vein, herbaceous to
subcoriaceous, glabrous, entire; marginal lobes usually
218
Filicatae • Polypodiaceae
broadly rounded, sometimes acute; veins anastomosing, forming a complex network with ex- or recurrent
included free veins. Sori rounded, laminal or submarginal on semi-circular projections covered by a deeply
immersed, acroscopically exposed sorus; sporangia
long-stalked, annulus with 12-14 indurated cells; receptacular trichomes 2- to 4-celled, glandular hairs;
rarely (in L. spinosa Jermy and Walker) sporangiasters
present. Spores hyaline or yellowish, ellipsoidal, the
aperture one-fourth to one-third the length of the
spore; exospore of two layers, the outer plain; perispore usually with spherical deposit and sheaths,
usually near the aperture, sometimes forming twisted
cables around the spore; surface slightly papillate,
sometimes with long filaments; n - ?
Lecanopteris s. lat. consists of 13 exclusively Malesian species. The fleshy, swollen, ant-inhabited stem is
a unique feature amongst Asian ferns. The genus can
be subdivided into two distinct parts: Lecanopteris
proper including 9 species with a scaleless stem, and
Myrmecopteris. Lecanopteris proper is most diversified
in Sulawesi (Hennipman 1986). The mature sporangia
of L. (Myrmecopteris) mirabilis (Co Chr.) Copel. consist
of four sets of 16 spores each enveloped by twisted
threads. A. F. Tryon (1985) thought this to be an adaptation to dispersal by ants, but Walker (1985) regarded
the discharged groups of 16 spores held together by
these filaments as a means of maintaining genetic variability by promoting intergametophytic mating of the
prothallial populations subsequently produced. This,
if true, constitutes an excellent example of synaptospory as first named by Kramer (1977).
The systematic position of the genus is as yet not
fully established. There are a number of differences
between the species of Lecanopteris proper and those
referred to Myrmecopteris. As yet it cannot be decided
whether these two groups merit separate generic recognition. Holttum accommodated the species with scaly stems in Phymatosorus ("'Phymatodes"), leaving the
others in Lecanopteris proper.
13. Colysis C. Presl
Fig. 117
Colysis C. Presl, Epimel. Bot.: 146 (1851).
Sel/iguea Blume (1826), nom. illeg., non Bory (1824).
Dendroglossa C. Presl (1851).
Myuropteris C. Chr. (1929).
Paraleptochilus CapeL (1947).
Epilithic, terrestrial, or epiphytic, small to mediumsized ferns with (Iong-)creeping stem clothed with
pseudopeltate or peltate, ovate-lanceolate, dark brown,
clathrate, entire or slightly toothed scales, these usually
with 1- or 2-celled, rarely multicellular, glandular marginal projections. Leaves remote, articulate, monomorphic or dimorphic. Sterile and monomorphic leaves
with usually well-developed and (at least above)
Fig. 117. Polypodiaceae. Co/ysis e/liptica, part of fertile lamina; Taiwan. Photo K. U. Kramer
winged petiole. Lamina simple, entire, palmately
lobed, digitate, pinnatifid, or pinnate with the pinnae
adnate to the rachis, (thinly) herbaceous to subcoriaceous; veins anastomosing, secondary veins prominent, reaching almost to the margin; usually two,
sometimes more rows of areoles between adjacent secondary veins, areoles with ex- or recurrent free veins.
Fertile leaves usually conform to sterile ones or sometimes smaller and (much) contracted. Sori usually between adjacent secondary veins, rounded or elongate
to linear, sporangia rarely acrostichoid; sporangial
stalk long, 3-seriate, annulus with 12-17 indurated
cells. Spores hyaline to light brown, ellipsoidal, the aperture one-fourth to three-fourth of the spore length;
surface shallowly tuberculate, usually plain with
abundant spherical deposit, sometimes with short
echinate elements; n=36, 2n-72, 108, 144,216.
An Asian genus generally reported to have about 30
species, erroneously reported from Africa by Ching
(1933b). It shows a most interesting variation in the arrangement of the sporangia. Although most species
Colysis • Leptochilus. Microsorum • Neocheiropteris
have linear sori on tertiary veins parallel to secondary
veins, others have either a strongly contracted fertile
lamina (e.g., C.poilanei C.Chr. & Tardieu-Bl.), or
round to elongate sori, e. g., C. hemionitidea (Wall. ex
Mett.) C. Presl which rarely has the sporangia acrostichoid. The delimitations between Colysis, Leptochilus,
and Microsorum need further research.
14. Leptochi1us Kaulf.
Fig. 118 A, B
Leptochilus Kaulf., En. Fil.: 147 (1824); Copel., (1928),
p.p.min.
Nistarika Nayar, Madhusoodanan & Molly, Fem Gaz. 13: 33
(1985).
Campium auctt., p. p., excl. type.
Epiphytic, rather small ferns with long-creeping stem
clothed with peltate, ovate, acuminate, clathrate scales
that are marginally entire or bear some long teeth with
1- or 2-celled, glandular marginal projections. Leaves
remote, articulate, dimorphic. Sterile leaves: petiole
absent or very short, lamina simple, entire, lanceolate,
herbaceous; veins anastomosing, areoles regular, decreasing in size towards the margin, with re- and excurrent free veins. Fertile leaves: petiole short to long,
lamina extremely contracted, linear; sporangia marginally inserted, acrostichoid in appearance, longstalked; annulus with 14 indurated cells; receptacular
hairs multicellular, glandular. Spores ellipsoidal, the
aperture one-fourth of the spore length; surface plain
with spherical deposit and short echinate elements;
n=36.
Three species, one in south-western India (L.lanceola/us Fee), one in Ceylon [L. wallii (Baker) C. Chr.],
and one [L. axillaris (Cav.) Kaulf.] from north-eastern
India to New Guinea. The systematic position is as yet
uncertain but seems closest to Microsorum and Colysis.
Nistarika (Nayar et al. 1985) seems to belong here, or
possibly in Colysis, but does not constitute a distinct
genus.
1S. Microsorum Link
Fig. 118 C-G
Microsorum Link, Hort. Berol. 2: 110 (1833); Bosman (1987b)
[often misspelled "Microsorium··'].
Dlblemma J. Smith (1841).
Dendroconche Copel. (1911).
Kaulinia Nayar (1964).
Podosorus Holttum (1966).
Medium-sized to rather large, epiphytic ferns with
short- or long-creeping stem sometimes covered by a
bluish-white ceraceous substance and bearing peltate
or pseudopeltate, rounded or ovate to lanceolate
scales, these entire or toothed and usually provided
with 1- or 2-celled, glandular marginal projections.
Leaves remote to approximate, usually articulate,
monomorphic, rarely dimorphic. Monomorphic and
219
sterile leaves: petiole if present naked; lamina simple,
regularly to irregularly lobed or pinnatifid, rarely pinnate, herbaceous, chartaceous, or coriaceous; veins
anastomosing in simple or complex patterns; first-order areoles regularly rectangular, regularly semi-elliptic, or irregular, with simple or forked free veins pointing to all sides and always terminated by a hydathode.
Fertile leaves partly or completely contracted; sori
usually orbicular, sometimes elliptic, linear and parallel to secondary veins or to the margin, or sunken in
rows parallel to the margin; sporangia sometimes arranged in irregular patches or acrostichoid, rarely assembled in marginal sori, long-stalked; annulus with
14-16 indurated cells; receptacular hairs multicellular,
glandular. Spores hyaline or somewhat yellowish, ellipsoidal, the aperture one-fourth to three-fourths the
length of the spore; exospore of two layers, with canals near the aperture; perispore thin, often with
sheaths; surface plain or ± rugose, usually with abundant spherical deposit, the spheres may be prolonged
and somewhat echinate; n = 36, 37.
A strongly diversified genus of c. 60 species, the
greater part Asian, a few elsewhere in the Palaeotropics. As construed here it includes acrostichoid representatives formerly referred to other genera, e. g., Microsorum (Christiopteris) varians (Mett.) Hennipm. &
Hettersch.
Microsorum
(Diblemma)
samarense
(J. Smith) Bosman has marginal sori as sometimes also
found in Microsorum bamlerianum (Rosenst.) Copel.
Phymatosorus (syn.: Phymatodes) is regarded as distinct; the type and its allies have characteristically entire leaves bearing a multitude of small sori consisting
of but few sporangia Bosman reinstated the name
Microsorum and gave (1987b) preliminary results of
monographic studies.
The spores of M. membranaceum (D. Don) Ching
are divergent in being manifestly tuberculate; their ultrastructure is reminiscent of that generally found in
Grammitidaceae. This must be a case of parallel development as most characters of this species agree well
with those of other microsoroid Polypodiaceae.
16. Neocheiropteris Christ
Neocheiropteris Christ, Bull. Soc. Bot. Fr. 52 Mem.: 1-21
(1905), based on Cheiropteris Christ, Bull. Herb. Boiss. 6:
876 (1898), nom. illeg., non KUIT (1858) (foss.).
Neolepisorus Ching (1940).
1Ticholepidium Ching (1978).
Platygyria Ching & Wu, Acta Bot. Yunnan. 2: 67 (1980).
Small to medium-sized, epiphytic ferns with longcreeping stem bearing usually pseudopeltate, sometimes peltate, normally ovate to lanceolate, sometimes
orbicular, clathrate, entire or toothed scales with 1- or
2-celled, rarely branched, glandular marginal projections; blade of scale opposite the point of attachment
220
Filicatae . Polypodiaceae
Fig. 118A-G. Polypodiaceae. A, B Lepfochilus axillaris.
A Habit with sterile and fertile lamina (x 0.3). B Venation of
sterile lamina (x 1.2). C-E Microsorum brachylepis. C Habit
(x OJ). D Detail of lamina with venation, placement of sori
indiCated by dotted circles (x 203). E Scale of stem (x 10.5).
F, G Microsorum membranaceum. F Habit (xO.14). G Part
of lamina with venation and sori (x 3) (A, B from Holttum
1955; C-E from Kurata and Nakaike 1981; F, G from Bir
and Shukla 1971)
invested with long, brownish, unicellular hairs. Leaves
remote, articulate, monomorphic. Petiole long, scaly.
Lamina usually entire, sometimes pedately lobed, irregularly lobed, or hastate, herbaceous or chartaceous,
scaly; veins anastomosing, areoles regular or irregular,
simple to complex, with simple to often forked, reand excurrent included free veins. Sori in one or two
rows on either side of the primary vein, or in one row
on either side of the secondary veins, rarely only borne
Dictymia • Phymatosorus • Campyloneurum
in the basal part of the lamina, oblong or rounded to
confluent, sometimes about linear or slightly irregular;
sporangia long-stalked, annulus with 16 indurated
cells; receptacular hairs glandular, receptacular scales
peltate, clathrate, intermediates between hairs and
scales present. Spores yellow, ellipsoidal, the aperture
three-fourths the length of the spore; surface plain to
slightly papillate, with spherical deposit; n -36.
About 5 to 10 species, distributed from north-eastern India east to Japan and south to Malesia. The
systematic position has been traditionally with Lepisorus. This, however, is not confirmed by the spore characters of its type species, N. palmatopedata (Baker)
Christ, which has microsoroid spores. In the emended
description of the genus given by Ching (1933 a) the
character of the stem scales showing a tuft of long,
brown, unicellulair hairs dorsally opposite the stalk
has been given much weight; this feature, when fully
expressed, is indeed most striking. However, detailed
studies of the scales of living plants of N. palmatopedata (unpublished) show that such unicellulair hairs are
absent from scales taken from supraterraneously growing stems, being present on scales of subterraneously
growing stems only. Besides, it was found that scales
with hairs on their dorsal side are also found in other
Microsorum species, e.g., in M.pteropus (Blume)
Ching, and also in Phymatosorus commutatus (Blume)
Pichi-Serm. The specific epithet of N. palmatopedata is
very appropriate.
11. Dictymia J. Smith
Fig. 107 A
Dictymia 1. Smith, Bot. Mag. 72 Comp.: 16 (1846); Walker
and Page (1982); Chandra (1983).
Dictyopteris C. Presl (1836), nom. iIIeg.. non Lamoureux
(1809).
Epiphytic or epilithic, small or medium-sized ferns
with long-creeping stem bearing peltate, broad, ovate,
clathrate, lacerate scales with usually glandular marginal projections. Leaves close to remote, articulate,
monomorphic. Petiole short, glabrous. Lamina simple,
linear or lanceolate, (sub)coriaceous, glabrous, broadly
costate; margin entire or sinuate; veins anastomosing,
areoles irregular, rarely with free included veins. Sori
large, rounded to elliptic or narrowly oblong, slightly
to strongly embossed, sporangial stalk 2-seriate with a
single small cell at the base, annulus with 12-21 indurated cells; receptacular trichomes usually absent,
sometimes 2- or 3-celled ones present; glandular hairs
present. Spores yellow, shallowly or coarsely rugose,
ellipsoidal, the aperture one-half the length of the
spore; 2n-70.
An isolated genus of two (three?) species occurring
from eastern Australia to Fiji, including New Caledonia. Its type of spore is similar to that found in Phymatosorus and in the lepisorioid Polypodiaceae.
18. Phymatosorus Pichi Serm.
221
Fig. 107 D
Phymatosorus Pichi Serm., Webbia 28: 457 (1973), based on:
Phymatodes, C. Presl, Tent. Pteridogr.: 195 (1836), nom. illeg.
Medium-sized to rather large, terrestrial, epiphytic, or
epilithic ferns; stem long-creeping, sometimes with
bluish, wax-like indument, clothed with peltate, broad,
clathrate scales bearing 1- or 2-celIed, glandular marginal projections. Leaves remote, articulate, monomorphic or sUbdimorphic. Sterile and monomorphic
leaves with long, stramineous, naked petiole; lamina
entire to very deeply pinnatifid (the shape sometimes
strongly fluctuating within one species or individual
plant), usually glabrous, rarely slightly pubescent, occasionally glaucous; margin entire, scarcely sclerotic;
veins anastomosing; main areoles semi-elliptic, in
broad leaves and leaf segments accompanied by polygonal areoles, included veins much anastomosing,
with simple or forked, re- and excurrent free veins.
Fertile leaves sometimes (slightly) contracted. Sori in
one, two, or three rows on either side of the primary
vein and/or secondary veins, underlain by soral veins
(diplodesmic), rounded to elliptic, more or less embossed; sporangia with long, uni- or biseriate stalk, annulus with about 14-16 indurated celIs; receptacular
hairs multicellular, glandular, cells in part inflated.
Spores yellow, ellipsoidal, the aperture one-half to
three-fourths the length of the spore; surface shallowly
and coarsely rugose; n = 36,37; 2n = 12, 14.
A small genus of about 10 species, in Africa, Madagascar, and the Mascarenes, and throughout the
warmer parts of Asia to Australia and Polynesia; as a
neophyte in tropical America [Ph. scolopendria
(Burm. f.) Pichi Serm.). Often accommodated in Microsorum, but differing in long-creeping or sometimes
pendulous (then up to 6 m long) stems, uniquely
shaped receptacular scales, and spores. A number of
species, including Ph. commutatus (Blume) Pichi
Serm., Ph. nigrescens (Blume) Pichi Serm. and Ph.longissimus (Blume) Pichi Serm., are provisionally included here; they may prove to be closer to Microsorum. The spores resemble those of Dictymia.
19. Campyloneurum C. Presl
Fig. 119 A-C
Campyloneurum C. Presl, Tenl Pteridogr.: 189 (1837).
Hyalotrichia Copel. (1953), nom. iIIeg., non Dennis (1949).
Hyaiotrichopterls W. H. Wagner, Taxon 27: 548 (1978).
Usually epiphytic, sometimes terrestrial or epilithic,
small to medium-sized ferns with a short- to longcreeping, stout to thin stem bearing peltate or pseudopeltate, ovate to linear-lanceolate, brownish, usually
clathrate scales that are entire or sometimes bearing
marginal, multicellular, glandular projections, sometimes a waxy bloom on the stem. Leaves usually ap-
222
Filicatae . Polypodiaceae
proximate, rarely remote, articulate, monomorphic.
Petiole short to long, glabrous. Lamina simple, entire,
rarely pinnate with adnate pinnae, narrowly linear to
broadly lanceolate, chartaceous to (firmly) coriaceous,
usually glabrous, entire, not rarely with revolute margin, often bearing a few basifixed clathrate scales;
veins anastomosing, forming series of areoles between
adjacent secondary veins, costal areoles with one excurrent free vein, other areoles with usually 2, sometimes more, parallel, occasionally branched, excurrent
free veins which may connect with the next cross-vein
and then subdivide the areole. Sori usually subterminal, on (free) included, excurrent veins, usually in 2 (or
3) rows between the secondary veins, rounded, superficial; sporangia long-stalked, smooth, annulus with
12-14 indurated cells; receptacular trichomes absent.
Spores whitish or yellowish, ellipsoidal, the aperture
one-third to three-fourths the length of the spore;
exospore of two layers, the outer verrucate; perispore
of one layer, conform to exospore; surface shallowly
to prominently verrucate with coarse spherical deposit; n-37, 74; 2n-74, 148.
A neotropical genus of c. 20 species occurring from
Florida and Mexico to Argentina, best characterized
by the small, round sori subterminally inserted on excurrent, free included veins. Some species are common
epiphytes, in humid forests especially of the lowlands,
but the genus is most diversified in the Andes. C. (Hyalotrichopteris) anetioides (Christ) R. & A. Tryon has
been recorded from vertical mossy rocks in dark rainforest (Wagner and Farrar 1976).
20. Neurodium Fee
Fig. 119 D, E
Neurodium Fee, Mem. Soc. Mus. Hist. Nat. Strasbourg 4; 201
(1850).
Heteropteris Fee (1843), nom. iIleg.• non H. B. K. (1821).
Paltonium C. Presl (1851), nom. superj/.
Epiphytic or rarely terrestrial, medium-sized ferns with
short-creeping, stout stem bearing peltate, linear-Ianceo late to ovate, clathrate, entire scales. Leaves approximate, articulate, monomorphic. Petiole short,
abaxially terete, adaxially flattened or shallowly sulcate, glabrous. Lamina simple, entire, Ian ceo late, coriaceous, glabrous, costate; veins anastomosing, areoles
irregular, with simple or forked free veins pointing to
all sides apical portion of fertile leaves soriferous,
somewhat contracted. Sporangia in continuous or interrupted, marginal soral lines, long-stalked; annulus
with about 13 indurated cells; receptacular hairs 2-,
rarely 3-celled, glandular. Spores hyaline, ellipsoidal,
the aperture one-fourth to three-fourths the length of
the spore; surface with prominent, dense tubercles and
abundant spherical deposit; n -37.
A single species, N. lanceolalum (L.) Fee, in Central
America and the West Indies, north to the Bahamas
and the Florida Keys; also in French Guiana? Epiphytic in the lowlands, up to 1000 m; of local occurrence.
21. Niphidium J. Smith
Figs. 109 I, 111 B, 119 F-J
Niphidium J. Smith, Hist. Fit.: 99 (1875); Lellinger (1972).
Pessopleris Underwood ex Maxon (1908).
Anaxetum Schott (1834), nom. iIleg., non Gaertner (1791) nec
Schrank (1824).
Epiphytic or epilithic, medium-sized to rather large
ferns with short- to long-creeping stem clothed with
peltate, ovate, lanceolate-acuminate, centrally clathrate
scales, these entire, apically toothed, or ciliate-toothed
and sometimes bearing 1- or 2-celled, glandular marginal projections, a wax-like bloom sometimes present.
Leaves approximate, articulate, monomorphic. Petiole
short to long, scaly at the base. Lamina simple, entire,
elliptic-Ianceolate to oblong or linear, costate, coriaceous, naked or scaly, often glaucous; hydathodes often present on adaxial face; veins anastomosing, secondary veins conspicuous, reaching almost to the
margin, tertiary veins irregular, areoles with forked,
re- or excurrent free veins. Sori orbicular or subelliptic,
large, in rows of 5-12 between two secondary veins,
one in each of the main areoles, apparently compital
but underlain by an irregular ring of veins; sporangia
long-stalked, annulus with 12-15 indurated cells; sporangial appendages of 1- or 2-celled, acicular hairs
situated near the annulus; receptacular trichomes
absent. Spores hyaline, ellipsoidal, sometimes with
obtuse ends, the aperture one-fourth to two-thirds the
length of the spore; surface plain or slightly papillate,
with sometimes dense spherical deposits; n -74,
2n -148.
Ten described species (Lellinger 1972) in the New
World from Cuba and central Mexico south to Argentina and Uruguay. Some species are very common epiphytic, terrestrial, or epilithic ferns in the lowlands as
well as in the mountains. N. americanum (Hooker)
J. Smith is characterized by the dense, ciliate scales on
the lamina beneath, associated with the occurrence of
cyclocytic stomata. This species occurs as a lithophyte
at higher elevations from 2300 to 2900 m (Lellinger
1972). As already pointed out by Tryon and Tryon
(1982), perhaps too many segregates have been split
off from the common and widespread N. crassi/olium
(L.) Lellinger.
22. Polypodium L.
Polypodium L., Spec. Plant. 2: 1082 (1753).
For synonyms, see under the infrageneric groups.
Usually epiphytic, sometimes terrestrial or epilithic,
most often medium-sized, sometimes small or rather
large ferns with long-creeping stem clothed with basifixed, peltate, or pseudopeltate, opaque or (partly)
Neurodium . Niphidium· Polypodium
clathrate scales. Leaves close to remote, articulate,
usually monomorphic. Monomorphic and sterile
leaves: petiole absent, short, or long, naked or scaly.
Lamina simple, pinnatifid, pinnatisect, or i-pinnate,
rarely more highly compound, herbaceous to coriaceous, glabrous, pubescent, or scaly; veins free or variously anastomosing. Fertile leaves conform to the
sterile ones or longer and! or narrower; sori in one or
two, rarely more rows on either side of the primary or
223
Fig. 119 A-J. polypodiaceae. A-C Campyloneun.lm phyllitidis. A Habit (x 0.5). B Detail of lamina with venation and
sari (x 2). C Scale of stem (x 7.5). D, E Neurodium lanceo/alum. D Habit (x 0.4). E Apex of fertile lamina with soral
lines (x3.5). F-J Niphidium crassifolium. F Habit (x 0.3).
G Portion of lamina with venation and sori (x 1.5). H Scale
of stem (x 3.5). I Detail of stem scale, partly clathrate
(x 20). J Part of bow of annulus with appendages (x 50)
(A-C, F, H-J from de la Sota 1977; D, E from Stolze 1981;
G from Wagner 1972)
224
Filicatae . Polypodiaceae
secondary vein(s) usually rounded, rarely elongate or
linear; sporangia long-stalked, annulus with 12-20 indurated cells; receptacular hairs and/or scales usually
present. Spores hyaline to yellow, ellipsoidal, the aperture one-half to three-fourths the length of the spore;
exospore of two layers forming the coarse contours;
perispore thin, conforming to exospore contours; surface usually prominently tuberculate to verrucose, often with spherical deposit, sometimes with prominent
verrucae with echinate tufts, rarely with coarse, winglike folds.
A cosmopolitan genus of 200 species; by far the
greatest concentration is in the American tropics. It is
here provisionally subdivided into four groups, all of
which are sometimes given generic status (e. g. by Tryon and Tryon 1982). Formerly the genus was even
more broadly circumscribed; here a number of distinct
groups are segregated as genera, e. g., Pecluma and Solanopteris. It is anticipated that further studies in Polypodium s.lat. will (again) lead to the recognition as
genera of groups here treated as subordinate to Polypodium. In the absence of such work the present classification must be regarded as rather provisional. Hybrids between "Pleopeltis" and Polypodium (s. str.) and
between the latter and "Microgramma" are known.
22.1 MicrogrammaGroup
Figs. 108 C, 109 E, 110 B
Microgramma C. Presl, Tent. Pteridogr.; 213 (1836); Tryon
and Tryon (1982).
Usually epiphytic, sometimes terrestrial or epilithic,
small ferns with long-creeping stem clothed with
usually peltate, rarely basifixed, opaque or centrally
clathrate, entire or toothed scales usually bearing 1- or
2-celled, unbranched, glandular marginal appendages.
Leaves remote, articulate, monomorphic to dimorphic.
Monomorphic and sterile leaves: petiole short to long,
scaly or glabrous. Lamina simple, herbaceous to coriaceous, glabrous or scaly, margin entire; veins anastomosing in a complex network with usually excurrent
free included veins, or free veins pointing to all sides.
Fertile leaves conform to sterile ones or weakly to
strongly contracted and then with simpler, sometimes
free venation. Sori usually in one, sometimes more series on either side of the costa, usually rounded, sometimes elongate; sporangia long-stalked, annulus with
about 12-20 indurated cells; receptacular scales usually present in various forms. Spores hyaline to yellow,
ellipsoidal to somewhat ovate, the ends obtuse; surface coarsely verrucate to irregularly tuberculate or
rugose, often bearing papillae; n =36 (7), 37, 74;
2n =148.
A group of c. 15 species, mostly confined to the
warmer parts of the New World; a single species in
Africa, Madagascar, and the Mascarenes; often common epiphytes in lowland forests, incidentally occur-
ring in montane vegetations. The group is close to Polypodium proper; a hybrid between one species each of
the two groups was described by Gomez (1977). The
group is easily distinguished from the Pleopeltis group
by the occurrence of hyaline, centrally clathrate scales.
The spores of P. megalophylium Desv. have an outer
surface ornamentation similar to that found in lepisorioid ferns which obviously represents a parallel development.
22.2 Polypodium Group
Figs. 106 A, 107 C, 109 B, F, H, 110 A, I, 111 D
Synammia C. Presl (1836).
Phlebodium (R. Brown) J. Smith (1841), based on Polypodium
sect. Phlebodium R. Brown (1838).
Lepicystis (J. Smith) J. Smith (1842), based on Goniophlebium
sect. Lepicystis J. Smith (1841).
Pseudocolysis Gomez (1977).
Epiphytic, epilithic, or terrestrial, small to large ferns;
stem rather short- to long-creeping, sometimes bearing
a wax-like bloom, clothed with usually peltate or pseudopeltate, rarely basifixed, opaque, clathrate or centrally clathrate, entire to toothed scales, their margin
usually bearing 1- or 2-celled, unbranched, glandular
appendages. Leaves usually remote, articulate at base,
monomorphic to dimorphic. Monomorphic and sterile
leaves with short to long, scaly or glabrous petiole;
lamina simple, pinnatifid, pinnatisect, or 1-pinnate,
rarely up to bipinnate + pinnatifid, most often deeply,
pectinately pinnatifid to pinnate, truncate or narrowed
at base, herbaceous to coriaceous, glabrous, pubescent, or variously scaly; veins free or anastomosing;
venation simple to complex, usually with one row of
areoles with one excurrent free vein, sometimes several
rows of areoles with free, simple or forked veins that
are usually excurrent or sometimes pointing to all
sides. Fertile leaves conform to sterile ones or larger
and/or narrower. Sori usually in one, sometimes more
rows on either side of the primary or secondary veins,
terminal or compital, usually rounded, rarely elongate
or linear; sporangia long-stalked, annulus with about
12-20 indurated cells; receptacular hairs and/or scales
of one form or another usually present. Spores hyaline
to yellow, ellipsoidal, the ends often obtuse; surface
coarsely verrucate or low-verrucose, tuberculate, or
papillate with dense globules, or with prominent
wings; exospore of two layers, the outer verrucate or
tuberculate; perispore thin, conforming to exospore,
or in some species forming prominent folds; n=37,
74; 2n=74, 111, 148.
In its present sense the group is an assemblage of
heterogeneous elements, even when Microgramma,
Goniophlebium. and Pleopeitis are recognized as groups
or genera. It is especially the New World representa-
Microgramma. Polypodium • PIeopeltis • Goniophlebium • Dicranoglossum
tives that need detailed study, as the group is most
strongly represented there. Tryon and Tryon (1982)
were in part successful in drawing distinct lines between 6 subgroups in this group of c. 150 species, even
though these are not all quite natural; e. g., they placed
Polypodium (Phlebodium) aureum L. and P. (PhI.) decumanum Willd. in different groups, which we cannot
continn, and also suggested to include Campyloneurum fendleri (Eaton) J. Smith and C. decurrens (Raddi)
C. Presl in Polypodium.
Certain authors like de la Sota (1965, 1966). Weatherby (1939,1947), and Maxon (1916 a, b) published
revisional studies of more or less distinct elements
in this alliance. Present knowledge suggests that
the group of species with "Lepicystis" scales fonns a
monophyletic group, together with the Pleopeltis group
and Dicranoglossum. The remaining group of species,
including those with goniophlebioid venation, is still
heterogeneous.
22.3 P1eopeltis Group
Figs. 109 D, G, 110 E, K, 120 A-F
Pleopeltis Humb. et Bonpl. ex Willd., Sp. PI. 5: 211 (1810).
Marginariopsis C. Chr. (1929).
Small, epiphytic or epilithic ferns with long-creeping
stem bearing peltate, linear or linear-lanceolate, clathrate, toothed scales with 1- or 2-celled glandular marginal appendages. Leaves approximate to remote, articulate, mono- or dimorphic. Monomorphic and sterile
leaves: petiole short or long, sometimes winged, scaly;
lamina usually simple, entire, sometimes pinnatifid,
(sub)coriaceous, variously densely scaly; lamina or
lobes lanceolate. entire; veins anastomosing in a simple or complex pattern; areoles regular with free veins
pointing to all sides. Fertile parts or leaves not or
somewhat contracted. Sori in one (or two) series on either side of the primary or secondary veins, usually
rounded, sometimes elliptic or elongate; sporangia
long-stalked, annulus with about 14 indurated cells;
receptacular hairs 2-celled or multicellular, sometimes
scale-like, glandular; receptacular scales peltate,
stalked, usually clathrate, rarely hyaline. Spores yellow, ellipsoidal, low-verrucate with sparse globules;
n=35, 37, 74; 2n=74, 148, 185, 210; aneuploid numbers have also been reported.
In its present circumscription, i. e., excluding Lepisorus, the Pleopeltis group is a neotropical group of
about ten species, only one, P.lanceolatum L. [= Pleopeltis macrocarpa (Bory ex Willd.) Kaulf.] extending
to Africa and India It is near the species group of Polypodium s. str. where "Lepicys/is" scales are found; instances of hybridization between the two groups are
known (see Wagner and Wagner 1975; Anthony and
Schelpe 1985, and Mickel and Beitel 1987); Dicrano-
225
glossum is also related. For the inclusion of Marginariopsis see Wagner (1986).
22.4 Goniophlebium Group
Figs. 107 B, 109 B, 110 I
Goniophlebium C. Presl, Tent. Pteridogr.: 185 (1836); Roedl-
Linder (1990).
Schellolepis J. Smith (1866).
Epiphytic, medium-sized to large ferns with longcreeping stem clothed with peltate, clathrate, toothed
scales usually bearing 1- or 2-celled glandular marginal appendages. Leaves remote, articulate, monomorphic: petiole long, glabrous. Lamina pinnatifid to pinnate with a confonn tenninal pinna; pinnae articulate,
lanceolate or linear, herbaceous, glabrous or pubescent, rarely scaly, margin entire or toothed; veins anastomosing, areoles regularly angular, costal areoles
large, with one excurrent free included vein; other
areoles, if present, similar or smaller, sometimes with
an excurrent free vein at each junction. Sori in one
row on either side of the secondary veins, tenninal on
free veins, rounded; sporangia long-stalked, annulus
with about 14 indurated cells; receptacular hairs multicellular, sometimes scale-like, unbranched, glandular,
receptacular scales basifixed or peltate, clathrate, entire or toothed, intennediates between hairs and scales
present. Spores yellow, with obscure aperture; exospore with tuberculate surface; perispore of one layer,
raised in folds; surface with prominent, wing-like folds
running parallel to the long axis; n=37; 2n=74.
A group of five to ten Asian species to which several
others from Asia (and America?) here left in the Polypodium group may have to be added.
23. Dicranoglossum J. Smith
Figs. 109 J, 120 G
Dicranoglossum J. Smith, Bot. Yoy. Herald: 232 (1854).
Eschatogramme Trevisan (1851), nom. nud.; Christensen
(1929).
Small, epiphytic ferns with short-creeping stem bearing peltate, ovate to lanceolate, clathrate and entire
scales. Leaves approximate, subpinnatitid, monomorphic. Petiole short, mostly winged, scaly, articulate.
Lamina subdichotomously forked, with a confonn terminal segment; segments connected at base, lanceolate
to linear, acuminate, entire, firmly herbaceous or subcoriaceous; scales centrally hyaline, marginally clathrate; veins free to anastomosing, up to twice forked,
sometimes with one row of regular areoles without
free included veins, distally from these sometimes with
excurrent free veins. Sori in one row on either side of
the costa, or usually midway between costa and margin, sometimes closer to the margin, usually continuous in the entire segment, linear, sometimes interrupted, occasionally rounded to elliptic, superficial; sporangia long-stalked, annulus with 13-16 indurated
Filicatae . Polypodiaceae
226
.~{.,~~:
........ ~ ..
".~.'.~'
D
B
Fig. 120A-G. Polypodiaceae. A-E Polypodium lanceolalum
(= Pleopeitis macrocarpa). A Habit (x 0.5). B Part of fertile
lamina with venation and sori (x 3.75). C Scales from stem
(x 15). D Sorus with peltate scales (x 7.5). E Stalked peltate
soral scale (x 15). F Polypodium (Pleopeltis) anguslum, habit
(x 0.5). G Dicranoglossum desvauxtt. habit (x 0.8) (A-E from
de la Sota 1977; F from Stolze 1981; G from Proctor 1977)
cells; receptacular trichomes absent. Spores yellow, ellipsoidal, the aperture one-half to three-fourths the
length of the spore; surface shallowly tuberculate, with
sparse spherical deposit; n = 36, 72.
About six species in the Neotropics from Central
America south to Bolivia. The division of the lamina is
distinctive. The genus is closest to Pleopeltis.
24. Solanopteris Copel.
Figs. 106 C, 110 C, 121 A-C
Solanopteris Copel.. Amer. Fern J. 41: 75 (1951) ("Solenopteris'~.
Mlcrogramma subg. Solanopteris (Copel.) Lellinger. Amer.
Fern J. 67: 59 (1977).
Soillnopteris. Pecluma· Po[ypodiopteris
227
G
B
Epiphytic, small ferns with long-creeping stem bearing
hollow pseudotubers inhabited by ants, clothed with
peltate, round, opaque, entire or slightly toothed scales
with 1· or 2-celled, glandular marginal appendages.
Leaves remote, articulate, dimorphic. Sterile leaves:
petiole short, winged; lamina entire to pinnately
lobed, lanceolate, herbaceous, glabrous or pubescent,
sparsely scaly; veins anastomosing, areoles irregular
with simple, sometimes forked, free included veinlets
pointing to all sides. Fertile leaves longer and narrower than the sterile. Sori orbicular, in one row on either
side of the costa, rounded to elongate, sometimes confluent, the receptacle underlain by several joined veinlets; sporangia long-stalked, annulus with about 12-14
indurated cells; receptacular hairs multicellular, scalelike. Spores yellow, ellipsoidal, the aperture one-half
the length of the spore; exospore of two layers, the
outer plain or shallowly tuberculate; perispore of one
layer forming the echinate elements; surface echinate,
Fig.i2iA-H. Polypodiaceae. A-C Solanopteris bifrons.
A Sterile leaf (x 1,5). B Part of fertile, leaf ( x 1.5). C Long
shoot with tubers, tuber pedicels (Kn) and leaf scars (BN)
(schematic). D, E Pecluma filicula. D Habit (x 0.5). E Part
of rachis with segments (x 3.75). F-H Loxogramme salicifolia. F Habit ( x 0.5). G Part of lamina with venation and sori
(x 2.5). H Scale of stem (x 14) (A, B from Wagner 1972; C
from Hagemann 1969; D, E from de la Sota 1977; F-H from
Kurata and Nakaike 1981).
with long spines, these sometimes constricted, or with
granulate deposit; n=37.
A genus of only three (or four) species, all epiphytes, from the lowlands to the upper montane forests of continental tropical America, from Costa Rica
to Peru. The genus has been merged with Micrograrnrna from which it diverges in several characters including the morphology of the spores. For a detailed study
further collections are required.
Filicatae . Polypodiaceae
228
25. Pecluma M. Price
Figs. 108 B, 121 D, E
Pecluma M. Price, Amer. Fern J. 73; 109 (1983).
Polypodium subg. Pectinatum Lellinger, Amer. Fern J. 71: 93
(1981).
"Polypodium pectinalUm Group" of Evans (1969).
Epiphytic or epilithic, usually medium-sized ferns with
often short-creeping stem bearing basifixed, usually
narrowly to linear-triangular, sometimes cordate or
ovate, opaque, mostly entire scales, these dorsally
mostly bearing tufted, unicellular hairs. Leaves approximate, articulate, monomorphic. Petiole usually
short to long, rarely absent, terete, usually pubescent
throughout and scaly at base. Lamina pectinate with
pinnatifid apex, narrowly elliptic, ovate, or deltoid, often reduced at base; lobes (very) narrowly oblong to
linear, usually entire, rarely crenate or crenulate, usually thinly pubescent with many multicellular glandular
hairs and ± hyaline acicular hairs, scaly on the rachis
only, base dilated, adnate; veins usually free, simple or
forked, rarely anastomosing, costal areoles then with a
single free vein. Sori terminal on the proximal acropetalous vein, rounded; sporangia long-stalked, with 13
or 14 indurated cells; usually some of the sporangia
with 2- or 3-celled hairs near the annulus, receptacular
hairs multicellular, glandular, rarely branched. Spores
ellipsoidal to subspherical, the aperture three-fourths
of the length of the spore; surface prominently verrucate, with sparse spherical deposit; n ~37.
About 25 species, in the warmer parts of the New
World from Florida to northern Argentina with a
unique combination of characters: short-creeping
stem, basally attached scales, pectinate leaves, sterile
parts of the lamina with uniquely shaped acicular cells
beside multicellular branched or unbranched glandular hairs. The systematic position is ambiguous. In the
past some species have been assigned to the Grammitidaceae because of acicular hairs which are, however,
essentiaJly different in both groups. The habitual similarity to species like Grammitis (Ctenopteris) taxi/olia
(L.) Proctor is, however, close and striking. Some authors include Pecluma in Po[ypodium.
26. Polypodiopteris Reed
Polypodiopteris Reed, Amer. Fern J. 38: 87 (1948).
Polypodiopsis Copel. (1947), nom. il/eg., non Carr. (1867).
Epiphytic, small to medium-sized ferns with longcreeping stem bearing peltate, Iinear-lanceolate,
opaque, toothed scales. Leaves remote, articulate,
monomorphic. Petiole long, glabrous, stramineous.
Lamina pectinate, gradually narrowed into a conform
terminal segment; segments very narrowly oblong, adnate, usually glabrous, rarely with a few hairs near the
rachis; margin crenulate or notched; base dilated;
veins anastomosing in one. row of areoles without in-
cluded free veins, otherwise free, excurrent. Sori in one
row on either side of the secondary vein, usually situated on the connecting veins flanking the costae, rarely
terminal on short, excurrent veins, rounded, superficial; sporangia long-stalked, annulus with about 14 indurated cells; receptacular trichomes absent. Spores
brown, baculate, otherwise smooth; n ~?
A genus of possibly three species (Copeland), not
only one (Christensen 1937), endemic to Borneo, in
montane forests (1000-1500 m), locally common. Apparently related to Selliguea ("Crypsinus"), as suggested by Christensen.
1:1. Thylacopteris Kunze ex J. Smith
Thylacopfetis Kunze ex J. Smith, Hist. Fil.: 87 (1875).
Epiphytic or terrestrial, medium-sized ferns with longcreeping stem clothed with pseudopeltate, ovate to
lanceolate, clathrate, entire scales sometimes bearing
1- or 2-celled glandular marginal appendages. Leaves
remote, articulate, monomorphic; petiole long, glabrous. Lamina pectinate, oblong, with a conform terminal lobe; lobes oblong, membranous, glabrous or
pubescent; veins free, simple or once forked. Sori in
one row on either side of the costa, terminal on an acroscopic vein, rounded, deeply embossed in T. papillasa; sporangia long-stalked, annulus with 12(-14) indurated cells; receptacular trichomes absent. Spores
yellow, ellipsoidal, the aperture one-half of the length
of the spore; surface plain or with shallow tubercles
and dense spherical deposit; n ~35, 36, 37.
Delicate ferns, the systematic position of which is
still uncertain. T. papillosa (Blume) Kunze ex J. Smith
is widespread in Malesia.
28. Anarthropteris Copel.
Fig. 110 G
Anarfhropteris Copel., Gen. Fil.: 217 (1947).
Stem very short-creeping, unbranched, roots often proIiferous; scales clathrate, brown, concolorous, lanceo late-deltoid, entire, basally attached. Articulation
between phyllopodium and leaf evident but nonfunctional. Entire plant, except roots, lacking sclerenchyma. Lamina simple, monomorphic, narrowly oblanceolate-elliptic, entire, thick-papyraceous, base
evenly attenuate to the petiole-like portion, apex
bluntly acuminate. Costa adaxially prominent,
rounded. Margins not cartilaginous, drying slightly revolute. Laminar surfaces glabrous except for minute
2-celled clavate glandular hairs. Veins regularly anastomosing without free included veinlets, no strong
main lateral veins, no hydathodes. Sori very slightly
sunken, orbicular to oblong, uniseriate on each side
and subparallel to costa, inframedial. Receptacular
scales dense, persistent, dark, clathrate, linear, gland-
Thylacopteris' Anarthropteris' Loxogramme
tipped, longer than sporangia. Sporangial stalk uniseriate at base, annulus with c. 16 indurated cells. Spores
yellow, oblong, monlete, aperture c. one-third the
length, surface minutely verrucose; n =37.
Monotypic. The single species, A.lanceolata
(J. D. Hooker) L. B. Moore, is endemic to New Zealand, epiphytic on trunks, petrophytic, and on steep
embankments in forest, 0-800 m.
29. Loxogramme(Blume) C. Presl
Fig. 121 F-H
Loxogramme (Blume) C. Presl, Tent. Pteridogr.: 214 (1836),
based on Antrophyum sect. Loxogramme Blume (1828).
Stem very short- to long-creeping, occasionally
branching, roots forming a spongy mass; scales clathrate, orange-brown to blackish, concolorous, entire,
elongate, basifixed. Articulation between phyllopodiurn and leaf none, or evident but not functional, or
functional. Entire plant, except for the roots, lacking
internal sclerenchyma. Lamina simple, monomorphic
to dimorphic, linear, narrowly elliptic, oblanceolate,
spathulate to orbicular, entire, thin- to thick-papyraceous, 1-90 em long and 0.3-15 em broad. Petiole-like
leaf base and costa not cartilaginous, prominent above
or below or the latter on both sides. Margins not cartilaginous, drying revolute or involute. Laminar surface
glabrous except for minute 2-celled clavate glandular
hairs. Veins regularly anastomosing with numerous,
few, or without free included veinlets; hydathodes
lacking. Sori mostly elongate and oblique to costa,
rarely orbicular or confluent and the sporangia acrostichoid. Receptacular appendages filiform or wanting.
Sporangial stalk 1- or 2-celled at base, annulus with
12-16 indurated cells. Spores green (at time of dispersal), globose-trilete or oblong-monolete, laesura c. onethird to two-thirds of the length, surface finely verrucose; n=35, 36.
About 33 species, one in Central America, one in
the Pacific from Fiji to Samoa, four in Mrica, the majority in Malesia and in tropical and warm-temperate
continental Asia. From sea level to high elevation,
growing as trunk epiphytes and shade petrophytes.
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229
Bir, S. S., Trikha, C. K. 1974. Taxonomic revision of the polypodiaceous genera of India VI. Lepisorns excavatus group.
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Bosman, M.1987. Preliminary results of monographic studies
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Chandra, S. 1979. Taxonomic use of foliar epidermis and hypodermis in drynarioid ferns. KaJikasan 8: 211-226.
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Copeland, E. B. 1928. Leptochilus and genera confused with
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Donk, M. A 1954. Notes on Malesian ferns - I. On the genus
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Gomez, L D. 1975. Contribuciones a la pteridologia
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Gomez, L D. 1977. Contribuciones a la pteridologia centroamericana XI. Novitates. Brenesia 10/11: 115-119.
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1-10.
Hennipman, E., Roos, M. C. 1982. A monograph of the fern
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Hennipman, E., Roos, M. C. 1983. Phylogenetic systematics
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Hennipman, E., Verduyn, G. P. 1987. A taxonomic revision of
230
Filicatae . Pteridaceae
the genus Lecanopteris (Polypodiaceae) in Sulawesi, indonesia. Blumea 32: 313-319.
Hetterscheid, W. LA., Hennipman, E. 1984. Venation patterns, soral characteristics and shape of the fronds of the
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Joe Hoshizaki, B. 1970. The rhizome scales of Platycerium.
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~
Hovenkamp, P. H. 1986. A monograph of the fem genus Pyrrasia. Leiden Bot. Ser. 9: 1-280.
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Janzen, D. H. 1974. Epiphytic mynnecophytes in Sarawak:
mutualism through the feeding of plants by ants. Biotropica 6: 237-259.
.
Jarrett, F. M. 1980. Studies in the classification of the ieplosporangiate ferns - I. The affinities of the Polypodiaceae
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825-833.
.
Jenny, A. C., Walker, T. G. 1975. Lecanopteris spinosa - a new
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Pteridaceae
R.M.TRYON
with the collaboration of A. F. TRYON and
K. U. KRAMER (Pteridoideae)
Pteridaceae Reichb., Handb. Nat. Pflanz.: 138 (1837), as Pteroideae.
Parkeriaceae Hook., Exot. Fl. 2: t. 147 (1825).
Acrostichaceae Frank, Syn. Pflanzenkd. (Leunis) ed. 2,3:
1458 (1877).
Sinopteridaceae Koidzumi, Acta Phytotax. Geobot. 3: 50
(1934).
Adiantaceae (C. Presl) Ching, Sunyatsenia 5: 229 (1940). nom.
cons. (over Sinopteridaceae).
Negripteridaceae Pichi Serm., Nuovo Giorn. Bot. It. N. S.
53: 160 (1946).
Platyzomataceae Nakai, Bull. Natl. Sci. Mus. (Tokyo) 29: 4
(1950).
Actiniopteridaceae Pichi Serm., Webbia 17: 5 (1962).
Cryptogrammaceae Pichi Senn., Webbia 17: 299 (1963).
Hemionitidaceae Pichi Senn., Webbia 21: 487 (1966).
Cheilanthaceae Nayar, Taxon 19: 233 (1970).
Taenitidaceae (C. Presl) Pichi Serm., Webbia 29: 1 (1974).
Filicatae . Pteridaceae
Terrestrial or rupestral, or (Ceratopteris) aquatic ferns
of mostly small size, exceptionally leaves are to 6 m
long. Stem erect to long-creeping, with a medullated
protostele, a solenostele, or a dictyostele, bearing trichomes or scales or both. Petiole with 1 to 4 vascular
bundles (or to many in Ceratopteris) near the base, terete to adaxially sulcate or two-ridged. Lamina usually
pinnate, or entire, pedate, palmate, radiate, or the petiole furcate apically into two strongly recurved rachises; veins usually free, if anastomosing the areolae
without free included veinlets. Sporangia in exindusiate, short to long soral lines along the veins or in
marginal sori or soral lines, and then usually covered
by a modified marginal indusium, with or without special trichomes among the sporangia. Sporangia usually
long-stalked, the stalk of 2 or 3 rows of cells (rarely to
5 rows), annulus vertical or rarely oblique, interrupted
by the stalk. Spores trilete, rarely a1ete, without chlorophyll.
NOMENCLATURE. Although Pteridaceae Reichb. is a
later name, it has consistently been used for a segregate group including Pteris, and is adopted here.
MORPHOLOGY AND ANATOMY. Most of these characters
are discussed under the subfamilies, but a few matters
are more appropriate here.
The stele types are based on Ogura (1972).
. The catadromous or anadromous arrangement of
the lamina architecture is variable in some species and
within most genera. However, in Afropteris, Onychium,
and Uavea it is strictly anadromous, in Pityrogramma
and Adiantum commonly anadromous, and in Ceratopteris usually catadromous.
Stomatal types have been variously classified, and
the scheme of van Cotthem (1970) is adopted (see also
Kondo 1%2; Fryns-Claessens and van Cotthem 1973;
Probst 1973).
Hydathodes are circular areas of modified vein ends
on the upper surface of leaves. They secrete an aqueous solution, especially in young leaves, when conditions for transpiration are reduced (Sperry 1983). In
most ferns hydathodes are identified on the basis of
their external appearance, and by this criterion they
occur in several genera of the Pteridaceae. Adaxial superficial vein ends, however, vary from circular to
elongate and slender and the application of the term
hydathode to all of these is uncertain. In several
genera apparent hydathodes are both present and absent, as in Doryopteris, Hemionitis, and Pterozonium In
others, elongate superficial vein ends are sometimes
present, as in Eriosorus, Jamesonia, and Pityrogramma,
but these may not be hydathodes. A special study of
these structures is needed to understand their nature
and to provide a clear definition of the term.
SUBDIVISION. The Pteridaceae have been variously
classified from a very large family (Copeland 1947) to
231
a restricted one that includes only Pteris and its nearest
allies (Pichi Sermolli 1977). Copeland included diverse
elements such as Dicksoniaceae and Dennstaedtiaceae
in the family, but treated Ceratopteris in its own family
(Parkeriaceae). Pichi Sermolli recognized 10 families
among the genera of Pteridaceae, a classification that
emphasizes some less distinctive groups. Others recognize only one or a few families.
The relations of the genera within the Pteridaceae
are complex and there are distinctive groups and poorly defined ones. The' chromosome number predominantly of n=29, 30, or multiples, trilete spores, and the
sporangia generally borne in exindusiate soral lines
along the veins, or covered by a modified marginal indusium, relate the genera in a natural group.
AFFINITY. The Vittariaceae, based on n- 60, are clearly
related and have sometimes been placed in the same
family with genera of Pteridaceae (Holttum 1949, and
Crabbe et at. 1975, as part of Adiantaceae). The Pteridaceae have no other obvious close relations among
living ferns, although they are often regarded as having a general affinity with the Schizaeaceae. The Pteridaceae are undoubtedly an ancient family and evidence of their evolutionary relations has been lost
through extinction.
DISTRIBUTION AND EcOLOGY. The Pteridaceae are essentially worldwide in distribution, althOUgh centred
in the tropics. Eleven of the genera are confined to the
New World and nine to the Old World. Most species
grow in rather open, often rocky, habitats, although
some, such as Adiantum and Pteris, are frequently in
forests. There is unusual ecological diversity, with Ceratopteris an aquatic genus, Acrostichum often associated with mangroves, and Jamesonia a characteristic
paramo genus. Genera of the subfamily Cheilanthoideae are often significant elements in the flora of xeric
habitats. Their leaves are often dormant when desiccated and revive with rainfall.
KEY TO THE SUBFAMIUES.
1. Wholly sterile leaves strongly dimorphic, some short and
filiform, others long and l-pinnate with several 100 small,
pouch-like, articulate pinnae; Australia Platyzomatoideae p. 232
- Wholly sterile leaves monomorphic or not markedly dimorphic, if l-pinnate then with few pinnae
2
2. Sporangia borne on a circular to elongate modified and
strongly recurved margin, facing the abaxial side of the
segment, which the veins enter
Adiaotoideae p. 247
- Sporangia borne on the abaxial side of the lamina or segment surface, or at its margin and then covered or not by a
modified marginal indusium, which lacks veins
3
3. Sporangia borne distantly on the veins; leaves dimorphic,
the sterile lamina thin, with evident anastomosing veins;
spores with prominent parallel ridges, without an equatorial flange
Ceratopteridoideae p. 232
- Sporangia approximate, in sori or sorallines on veins or on
a marginal commissure, Or also on the leaf tissue, rarely
232
Pteridaceae • Platyzomatoideae . Ceratopteridoideae
single at the end of a vein; spores with various surface features, if strongly few-ridged, then with an equatorial flange
4
4. Stem with trichomes only
Taenitidoideae p. 234
- Stem with scales, trichomes sometimes also present
5. Trichomes among the sporangia unlike other lamina indument, or, if these rarely absent, then the sporangia on a
marginal commissure and the lamina pinnate, anadromic
at and near the base and catadromous beyond and with
Pteridoideae p. 249
mostly adnate ultimate segments
- Trichomes absent among the sporangia, or if present then
similar to those on the leaf tissue; sporangia variously
borne, if (rarely) on a marginal commissure then the lamina pedate or, if pinnate, then either wholly anadromous or
with stalked ultimate segments
6
6. Spores with an equatorial flange, or, if not, then the petiole
with two or more vascular bundles near the base and the
lamina farinose beneath
Taenitidoideae p. 233
- Spores without an equatorial flange, the petiole with 1 vascular bundle near the base, or, if with 2, then the lamina
not farinose beneath
Cheilanthoideae p. 240
Subfamily PIatyzomatoideae
(Nakai) A Tryon, Amer. J. Bot. 51: 942 (1964). - Platyzomataceae Nakai, Bull. Natl. Sci. Mus. (Tokyo) 29: 4 (1950).
Terrestrial, small plants growing in dense colonies, the
stem growth and production of new leaves at the periphery. Stem short-creeping, with a medullated protostele with inner endodermis, bearing a dense covering of rigid, reddish, multicellular trichomes. Leaves
trimorphic, some sterile ones small, filiform, without
pinnae, others much longer, narrowly linear and 1-pinnate, the fertile similar to the large sterile leaves but
bearing somewhat larger fertile pinnae in zones along
the leaf. Petiole with 1 vascular bundle near the base,
nearly terete or adaxially slightly sulcate towards the
apex. Lamina 1-pinnate, determinate, with up to several 100 very small, nearly sessile, pouch-like, articulate
pinnae, glabrate or glandular with yellow farinose exudate especially when young; veins free. Sporangia
borne along the veins, behind the apex, in short, exindusiate sorallines, short-stalked, with a spherical capsule and broad, more or less oblique, nearly complete
annulus interrupted by the stalk, small ones bearing 32
mostly small spores and large ones bearing 16 large
spores, these commonly on the same pinna. Spores trilete, globose, coarsely reticulate, the large spores with
strong ridges parallel to the equator, the surface with
more or less dense granulate deposit, especially on
small spores that lack an equatorial flange; 2n=76.
A single monotypic genus.
MORPHOLOGY AND ANATOMY. The first studies were by
Thompson (1916, 1917, 1919), and later ones by
A. Tryon (1961, 1964). Platyzoma is a very unusual
fern, especially in its large number of small and
pouch-like pinnae, dimorphic sporangia and spores,
and incipient heterospory. Filiform leaves c. 3-8 em
long are borne by juvenile plants and also occur in
zones on adult plants. The pinnate leaves are up to
c.75 cm long. Capitate glands bearing a yellow farinose exudate occur especially in young leaves. The
stomata are anomocytic (van Cotthem 1970).
GAMETOPHYTE. The unusual incipient heterosporous
condition of Platyzoma was recognized by A. Tryon
(1964). The small spores produce a small, short-filamentous, apically slightly expanded gametophyte
without rhizoids and with antheridia on all surfaces.
The large spores produce larger, spathulate gametophytes with rhizoids and a few archegonia in a marginal position. Heterospory is not completely developed
since there is some variation in sporangia and spore
size, and old archegoniate gametophytes may produce
some antheridia. Additional details were published by
Duckett and Pang (1984).
KARYOLOGY. A. Tryon and Vida (1967) reported
2n= 76. Although this number is known in other genera, it is more meaningful in respect to evolutionary relations to that known in some species of Anemia and
Mohria, with n= 76, in the Schizaeaceae.
DISTRIBUTION AND EcOLOGY. Platyzoma has a scattered distribution in northern and northeastern Australia where it grows in open vegetation, primarily in
deep white granitic sand ("nicotine sand") that overlies heavy clay and is often flooded in the rainy season.
AFFINITY. Platyzoma was first described in the Gleicheniaceae (Brown 1810) and treated as a separate
family within the. Gleicheniales (Nakai 1950). Comparisons were made especially to Jamesonia by Holttum (1957) who indicated its affinity was with the
"Gymnogrammoid" ferns (= Taenitidoideae). The
chromosome number and surface contours of the
spores are similar to some of the Schizaeaceae, while
the granulate surface deposit on the spores as well as
the farinose indument on the leaf indicate relations
with the Pteridaceae (Wollenweber et aI. 1987).
1. Platyzoma R. Brown
Fig. 122
Platyzoma R.Brown, Prodr. A. Nov. Holl.: 160 (1810);
A Tryon (1%1, 1964).
Platyzoma microphyllum R. Br. is the only species. The
characters are discussed under the subfamily.
Subfamily Ceratopteridoideae
(J.Smith) R. Tryon, Amer. Fern J. 76: 184 (1987). - Tribe Ceratopterideae J. Smith, Hist. Fil.: 170 (1875).
233
Platyzoma • CerafOpferis
Palustral or often aquatic, rather small, short-lived
plants. Stem small, erect, with a perforated dictyostele,
bearing a few thin scales. Petiole with 4 to many vascular bundles. Leaves dimorphic (the sterile lobed to
3-pinnate, the fertile 1- to 5-pinnate, more erect and
with narrower segments). Lamina glabrous, catadromous; veins anastomosing without included veinlets.
Sporangia dispersed on the veins, more or less covered
by a strongly modified marginal indusium. Spores tetrahedral-globose, with coarse, nearly parallel ridges,
the surface more or less covered by rodlets and granulate deposit, without an equatorial flange.
A single genus.
MORPHOLOGY AND ANATOMY. Kny (1875) published
the first thorough account of the morphology of Ceratopteris. A recent review by Pal and Pal (1962) includes
the studies made after Kny as well as some additional
observations.
The stem has a protostele in young plants and a dictyostele in mature plants. Ceratopteris pteridoides
(Hook.) Hieron. is a floating species with inflated petioles 3 -4 em thick, with many vascular bundles.
Roots are primarily borne at the base of the petioles.
The leaves have dormant bud initials in the sinuses
that may develop into young plants, especially when
an old leaf decays and plantlets become separated.
The stomata are polocytic (van Cotthem 1970). '
The sporangia are globose, nearly sessile, and are
unusual in the variation of the number of indurated
cells of the annulus as well as the irregularity of the
annulus. In Ceratopteris pteridoides the annulus is absent or nearly so.
GAMETOPHYTE. Pal and Pal (1963) presented a modem
account of the gametophyte. It is generally cordate but
asymmetrical, with a lateral meristem and no apical
cell, glabrous and with a poorly developed midrib; antheridia are borne on the lower surface and the margins, archegonia are mostly on the midrib.
KARYOLOGY. The chromosome numbers of Ceratopteris indicate aneuploidy and polyploidy. Confirmed
counts are n~39, 77, 78, and there are less certain reports of 2n~80 and 154.
AFFINITY. The relation of Ceratopteris to other ferns is
not clear, probably due to the modification of characters as a consequence of the aquatic habit. While the
spores and chromosome numbers suggest an ancient
affinity with Anemia of the Schizaeaceae, the sporangia, borne along the veins, and the strongly modified
marginal indusium ally it to the Pteridaceae.
DISTRIBUTION AND EcoLOGY. Ceratopteris is widespread through the tropics of both hemispheres. Ceratoptens pteridoides is only American, C. cornuta
(Pal. Beauv.) Lepr. is mostly African but extends east
to Australia, and C. thalictroides (L.) Brongn. is pan-
Fig.t22. Pteridaceae.
Phot. A C. Jermy
Platyzoma microphy/lum;
Queensland.
tropical. Plants grow in a variety of aquatic or wet habitats, such as lakes, ponds, rivers, open swamps,
ditches, and rice and taro fields.
2. Ceratopterls Brongniart
Cerafopferis Brongniart,
Bull. Soc. Philom.: 186 (1821); Uoyd
(1974); Tryon and Tryon (1982).
This is the only homosporous fern with a floating aquatic habit. Three species are recognized by Tryon and
Tryon (1982), four by Lloyd (1974); the characters of
Ceratopteris are discussed under the subfamily.
234
Pteridaceae . Taenitidoideae
Subfamily Taenitidoideae
(C. Presl) R. Tryon, Amer. Fern J. 76: 184 (1987). - Tribe
Taenitideae C. Presl, Tent. Pteridogr.: 222 (1836).
Terrestrial or rupestral, mostly small plants. Stem erect
to long-creeping, with a solenostele, with transitions
towards a dictyostele, or with a dictyostele, bearing trichomes, bristles, or rather undifferentiated scales. Petiole with 1-4 vascular bundles near the base, terete to
adaxially sulcate. Lamina diverse, simple, entire, with
f1abellate venation to 6-pinnate, gradually reduced to
the apex or rarely with stalked ultimate segments and
an odd terminal pinna, glabrous to usually, sometimes
densely, glandular, pubescent or farinose (not scaly)
beneath; veins usually free, sometimes anastomosing.
Sporangia borne along the veins in exindusiate soral
lines, or in an inframarginal soral band, or along a
marginal commissure, with trichomes often borne on
or among the sporangia similar to or different from
those on the leaf tissue. Spores trilete, usually with
prominent coarse ridges, and an equatorial flange.
CHARACTERS OF RARE OCCURRENCE. The stele of Anogramma is a medullated protostele with internal phloem (Mehra and Sandhu 1976) and that of Actiniopteris
a dictyostele.
The venation of the pinnatisect lamina of Pityrogramma lehmannii (Hieron.) R. Tryon is unique, with
many veins separately entering the primary segments
from the costa.
MORPHOLOGY AND ANATOMY. The stem usually has a
siphonostele, often grading towards a dictyostele when
the leaves are closely spaced. In most genera the stem
bears trichomes or bristles (trichomes with a thickened, multicellular base), or less often scales. Unusual
leaves in the subfamily occur in some species of Eriosorus with scan dent leaves to over 4 m long, in Jamesonia with leaves of indeterminate growth, and some
species of Pterozonium with simple, entire and xeromorphic leaves. The stomata are polocytic or anomocytic, or in Actiniopteris they are hypocytic, and some,
in Afroptens, are pericytic or desmocytic (van Cotthem
1970). In some genera there is an indument among the
sporangia that is similar to that on the leaf tissue. Specialized trichomes of 3 to many cells occur in others.
These are often curved and sometimes distally enlarged and secretory. They may be borne on the sporangium stalk as well as the fertile vein (Walker 1968;
Hennipman 1975). These have been referred to as paraphyses. They are sometimes useful to distinguish
species although they are not consistent in genera.
GAMETOPHYTE. The gametophyte of the Taenitidoideae is usually initially spathulate with a lateral meristem and later may develop lobes with additional meristems (Stokey 1948; Nayar 1%4; Atkinson 1970;
A. Tryon 1970; Nayar and Kaur 1971; Mehra and
Sandhu 1976). Old gametophytes are asymmetrically
cordate or irregularly lobed with archegonia and antheridia variously distributed. The gametophytes of
Anogramma are exceptional as they produce special
branches bearing archegonia and antheridia, which
give rise to slightly subterranean tubercles. These may
become dormant and produce a new gametophyte at
the beginning of a growing season. Baroutsis (1976) reported tubercles in four species but not in Anogramma
osteniana Dutra.
KARYOLOGY. The known chromosome numbers are
quite uniform, mostly n~ 29 or multiples or less often
n~ 30 or multiples. Taenitis is widely different with
n~44, 108, 110, and 114 reported.
PHYTOCHEMISTRY. A farinaceous exudate from stalked
glands occurs prominently on the abaxial surface of
the lamina of Pityrogramma and Onychium siliculosum,
and especially among the sporangia in Pterozonium
and Cerosora. The flavonoid chemistry has been extensively studied in Pityrogramma, especially by Wollenweber (1978) and Wollenweber and Dietz (1980).
The genus is characterized by the dominance of chalcones and dihydrochalcones. Certain species tend to
have a particular combination of compounds but due
to their variation the present res~lts do not provide significant data for systematics.
AFFINITY. The subfamily Taenitidoideae'is clearly related to others of the Pteridaceae by the predominant
chromosome numbers of n ~ 29 or 30 or multiples. The
few exceptions do not alter relationships which are
evident in other characters.
DISTRIBUTION AND ECOLOGY. Eriosorus, Jamesonia,
Pterozonium, and Nephopteris are confined to tropical
America. Pityrogramma and Anogramma centre in the
American tropics, the former extending eastward to
Madagascar, the latter subcosmopolitan. Actinioptens
is primarily African, and the other genera centre in
southeastern Asia.
The species mostly grow in moist, open, terrestrial
or rupeslral habitats. The rupestral species are nearly
all confined to either sandstone or to limestone. Others
are forest species, especially growing along or near
streams, where a few may be rheophytes. Austrogramme francii (Rosenst.) Hennipm. is sometimes an
epiphyte.
KEy TO THE GENERA.
1. Stem annual, bearing only living leaves, erect, very small,
with a few whitish trichomes or scales
9. Anogramma
- Stem perennial, bearing petiole bases of previous leaves,
usually decumbent or creeping, with many brownish or
dark trichomes, bristles, Or scales
2
2. Stem bearing only trichomes or bristles
3
- Stem with scales
10
Eriosorus • Jamesonia . Pterozon;um
3. Small trichomes different from those on the leaf tissue
among the sporangia or on their stalk, or rarely none and
then the veins anastomosing Or their ends joined at the
margin
4
- Trichomes, if present among the sporangia, similar to
those on the leaf tissue; veins wholly free
7
4. Veins anastomosing from costa to margin, or, if only costal areolae present, then the lamina nearly orbicular
15.1Ilenitis
- Veins joined only at the margin or in 1 or 2 series of marginal areolae. or veins free at the margin and the lamina
elongate
5
5. Veins joined at the margin or in 1 or 2 series of marginal
areolae
14. Syngramma
- Veins free at the margin, all veins free or only costal areolae present
6
6. Stem with rigid, dark bristles; petiole with one vascular
7. Cerosora
bundle
- Stem with short, brown trichomes or bristles; petiole with
two vascular bundles
13. Austrogramme
7. Fertile leaves erect, much longer than the sterile
6. Nephopreris
- Fertile leaves similar to the sterile
8
8. Pinnae entire to shallowly lobed, the lamina indeterminate, or if detenninate the pinnae adnate
4. Jamesonia
- Pinnae pinnatifid to decompound, or, if entire, the lamina
determinate and the pinnae stalked
9
9. Pinnae cleft, lobed, pinnatifid or more complex, if entire
then the largest c.1 ctn broad; stem short-ereeping;
3. EriosortlS
spores trilete
- Larger pinnae cleft to pinnate; stem long-creeping, spores
monolete
(genus incertae sedis: 34. P1eurosoriopsis)
- - Pinnae entire, the largest c.2 em broad, spores trilete
5. Pterozonium
10. Fertile lamina entire, or 1-pinnate with entire pinnae with
flabellate venation
5. Pterozonium
- Fertile lamina dichotomously cleft
12. Actiniopleris
- - Fertile lamina pinnatisect, or, if pinnate, then the pinnae
with a costa
11
11. Sporangia on a marginal commissure, covered by a modified marginal indusium
12
- Sporangia in soral lines along the veins, the margin not
modified
8. Pityrogramma
12. Spicular cells present in the epidermis; basal pinnae with
the basal acroscopic segment c. one-half as long as the basiscopic one
10. Afropteris
- Spicular cells absent in the epidermis; basal pinnae with
the basal aeroscopic and basi scopic segments nearly of
the same length
11. Onydrillm
3. Eriosorus Fee
Fig. 123A-C
£riosorus Fee, 5' Mem. Gen. Fil.: 152 (1852); Scamman
(1962); A. Tryon in R. Tryon (1964); A. Tryon (1970); Tryon
and Tryon (1982).
Gymnogramma auctt., at least in part.
Stem decumbent or creeping, with golden brown to
blackish, usually rigid trichomes or bristles, or very
rarely with light brown, narrow scales. Leaves monomorphic. Petiole with 1 vascular bundle near the base,
terete or subterete, or adaxially sulcate, especially near
the apex. Lamina 1-pinnate to 6-pinnate, anadromous,
usually determinate, usually pubescent or glandular;
235
veins free, ending at or somewhat behind the margin,
the ends slightly to clearly enlarged. Sporangia borne
along the veins in short, sometimes branched, exindusiate sorallines, these sometimes with trichomes similar to those on the leaf tissue. Spores trilete, tetrahedral, with coarse, often fused tubercles, the distal face
with coarse ridges more or less extended at the angles,
with a prominent equatorial flange; n=87, 174.
Eriosorus flexuosus (HBK.) Copel. var. galeanus
A. Tryon, from Guerrero, Mexico, the northernmost
occurrence of the species, has narrow scales on the
stem, undoubtedly derived from the trichomes of var.
flexuosus.
Nine hybrids in Eriosorus, five of them within the
genus and four with Jamesonia, have been recognized
by A. Tryon (1970). Some of the variation in common
species such as Eriosorus flexuosus is also considered
to be derived from hybridization.
A tropical American genus of 25 species, primarily
Andean, from Mexico and the West Indies south to
Bolivia, Brazil and Uruguay; also on Tristan da Cunha
and on Gough Island
4. Jamesonia Hooker & Greville
Fig. 123D-F
Jamesonia Hooker & Greville, Icon. Fil.: pI. 78 (1830); A. Tryon (1962); A. Tryon in R. Tryon (1%4); Tryon and Tryon
(1982).
Psilogramme Kuhn (1882).
Stem decumbent or usually long-creeping, with light
brown to black, usually rigid trichomes or bristles.
Leaves monomorphic. Petiole with 1 vascular bundle
near the base, terete to adaxially slightly sulcate. lamina 1-pinnate, linear, usually indeterminate, often pubescent, sometimes densely so, or glandular, pinnae
small, usually roundish and with strongly recurved
margins; veins free, ending at or near the margin, the
ends not or slightly enlarged. Sporangia borne along
the veins in short, sometimes branched, exindusiate
sorallines, these often with trichomes similar to those
on the leaf tissue. Spores trilete, tetrahedral, with
coarse, often fused tubercles, distal face with three
prominent ridges more or less prolonged at the angles,
with a prominent equatorial flange; n=87.
A tropical American genus of 19 species, almost exclusively Andean but locally also in southern Mexico,
Costa Rica, and in southeastern Brazil (Mt. Jtatiaia).
Jamesonia represents an unusual evolutionary development among ferns. Its species have radiated in
the Andes where they are evidently adapted to the special paramo environment by their indeterminate,
1-pinnate, linear leaves bearing many small, imbricate
pinnae. The genus is closely related to Eriosorus and
evidently derived from the group of Eriosorus cheilanthoides (Sw.) A. F. Tryon.
Pteridaceae . Taenitidoideae
C
Fig. 123A-K. Pteridaceae. A, B Eriosorus hirtus. A Habit
(x 0.3). B Fertile pinnule, abaxial side (x 0.7). C Trichomes
from adaxial side of lamina (x 40). D-F Jamesouia rotundifolia. D Pinna from abaxial side (x 5). E Trichome from
rachis (x 30). F Trichomes from stem (x 30). G Pterozonium
reniforme. habit (x 0.5). H Pityrogramma calomelanos. pinna
( x 0.5). I Pityrogramma ferruginea. part of rachis with two
pinnae (x 0.5). J, K Pityrogramma trifoliata. J Sterile pinna
(x 0.5). K Fertile segment from abaxial side (x 0.5) (A-C
from A. F. Tryon 1970; D-F from A. F. Tryon 1962; G from
R. M. Tryon 1964; H-K from R. M. Tryon 1962)
5. PteroZOIIium Fee
Fig. 123G
Pterozonium Fee,S' Mem. Gen. Fil.: 178 (1852); Lellinger
(1%7); Tryon and Tryon (1982).
Syngrammatopsis Alston (1952).
Stem creeping to erect, with brownish to atropurpureous, lax or rigid bristles, or dark, rigid, narrow scales.
Leaves monomorphic. Petiole with 1 or 2 vascular
bundles near the base, terete to adaxially sulcate, rare-
Iy narrowly alate. Lamina usually entire, or l-pinnate
with an odd tenninal pinna, often xeromorphic, with
or without sparse indument; veins free, ending behind
the margin, the ends not or definitely enlarged. Sporangia borne along the veins in long to short, exindusiate soral lines, or confined to the vein ends, sometimes with glands with farinose exudate among the
sporangia. Spores trilete, tetrahedral-globose, somewhat tuberculate, with more or less dense granulate
deposit, distal face with a strong triradiate ridge extending into the angles, with a prominent equatorial
flange; n=unknown.
An American genus of 13 species, centred on the
Roraima sandstones of Venezuela where all species
occur. Also east to Surinam and southwest to Colombia and northern Peru.
6. Nephopteris Lellinger
Nephopteris Lellinger, Amer. Fern J. 56: 180 (1966); Tryon
and Tryon (1982).
Nephopteris· Cerosora· Pityrogramma. Anogramma
Stem short, with dark brown, rigid trichomes. Leaves
dimorphic (the fertile larger and more complex). Petiole with 1 vascular bundle near the base, terete to
adaxially sulcate. Sterile lamina 1-pinnate, fertile lamina partly 2-pinnate, densely pubescent beneath; veins
free, ending at the margin or nearly so, the ends not
enlarged. Sporangia borne along the veins in often
branched, exindusiate sorallines, these with trichomes
similar to those on the leaf tissue. Spores trilete, tetrahedral-globose tuberculate expecially on the proximal
face, three prominent ridges on the distal face more or
less extended at the angles, with a prominent equatorial flange; n=unknown.
The only species is Nephopteris maxonii Lell., known
from three localities in central Colombia at c. 3000 m.
It is perhaps close to Eriosorus but is recognized as a
separate evolutionary line on the basis of the small (to
10 cm long), dimorphic leaves and reduced lamina architecture.
7. Cerosora (Baker) Domin
Cerosora(Baker) Domin, Acta Bot. Boh. 8: 3 (1929); Holttum
(1959). Based on: Gymnogramma Sect. Cerosara Baker,
J. Linn. Soc. 24: 260 (1887).
Anogramma Sect. Monosorus Domin (1928).
Medium-sized ferns with creeping stem bearing dark,
rigid trichomes or bristles. Leaves monomorphic; petiole well-developed, brown-sclerotic, with a single
vascular bundle, adaxially sulcate. Lamina narrowly
triangular, bipinnate to tripinnate or partly quadripinnate, anadromous, not reduced at base, with ascending, short-petiolulate pinnae, glabrous. Ultimate divisions rhombic-obtriangular, narrowed at base, cleft-laciniate, with narrow, subacute lobes. Veins free, ending
nearly at or some way behind the margin, their ends
not enlarged. Sporangia borne along the veins in short,
sometimes branched, exindusiate soral lines, these
sometimes yellow-farinose from the exudate of fewcelled simple trichomes. Spores trilete, tetrahedral-globose, with compact tubercles denser and somewhat
fused on the distal face, with a prominent flange somewhat above the equator on the proximal face; n=unknown.
A small genus of three species. Cerosora chrysosora
(Baker) Domin in Borneo, C. sumatrana Holttum in
Sumatra, and C. microphylla (Hooker) R Tryon in the
Himalaya. The ultimate segments of this last species
are small and elliptic-acute.
8. Pityrogramma Link
Fig. 123H-K
Pifyrogramma Link, Handb. d. Gew. 3: 19 (1833); R. Tryon
(1962); Tryon and Tryon (1982).
Ceropferis Link (1841), nom. ilIeg.
1Iismeria Fee (1852); Copeland (1947).
Gymnogramma aucll., p.p.
237
Stem short-creeping to erect, with light brown to
atropurpureous, rather narrow and usually thin scales.
Leaves monomorphic or slightly dimorphic (the fertile
more erect). Petiole with 2-4 vascular bundles near the
base, rarely terete, usually adaxially sulcate. Lamina
2-pinnate to 4-pinnate-pinnatifid, or rarely only pinnatisect, anadromous (exc. P. triangularis), usually
abundantly white-, yellow- or orange-farinose beneath,
or sometimes densely pubescent, or glabrous; veins
free, ending at or behind the margin, the ends not or
somewhat enlarged. Sporangia borne along the veins
in usually short, sometimes branched, exindusiate soraI lines, these usually with glands bearing farinose indument similar to those on the leaf tissue. Spores trilete, tetrahedral-globose, rugose to reticulate, the distal
face usually with coarse ridges parallel to the equatorial flange, or spores sometimes lacking a flange and
more or less rugose or granulate; n= 30,58, and multiples.
A genus of about 16 species, native in the American
tropics, where most species occur, and in the southwestern United States and Pacific Coast (1 species),
and in Africa-Madagascar (4 or 5 species). Ten species
occur in the Andes of Colombia, Ecuador, and Peru.
The genus is widely adventive, especially represented
by Pityrogramma calomelanos (L.) Link, through the
paleotropics.
9. Allogramma Link
Anogramma Link, Fil. Sp. Cult.: 137 (1841); Tryon and Tryon
(1982).
Gymnogramma auctt., p. p.
Stem erect, very small, with usually very few thin,
whitish trichomes or scales. Leaves monomorphic. Petiole with 1 or 2 vascular bundles near the base, adaxially sulcate, rarely with few to many trichomes. Lamina 1- to 4-pinnate, anadromous, glabrous or rarely
somewhat pubescent; veins free, ending nearly at or
behind the margin, the ends not enlarged. Sporangia
borne along the veins in short, sometimes branched,
exindusiate soral lines. Spores trilete, tetrahedral-globose, tuberculate-rugose, the distal face with prominent ridges parallel to the equatorial flange, the surface usually with a granulate deposit; 11=29,58.
A genus of about five species, widely distributed in
the American tropics from Mexico to Argentina; also
in the Azores south to the Cape Verde Islands, Africa,
and Madagascar, south-western and southern Europe
to northern India and Ceylon, east and south to Taiwan, Australia, and New Zealand.
Although technically hardly different from Pityrogramma, Anogramma is recognized as a significant
evolutionary line on the basis of the annual stem that
does not bear leaf bases of previous years and the unusual gametophyte that produces a dormant tubercle
238
Pteridaceae . Taenitidoideae
F"1g.124A-H. Pteridaceae. A,B A/ropteris repens. A Habit
(x 0,7). B Fertile segment from abaxial side (x 2). C Syngramma cartilagidens, tip of fertile lamina with sora! lines
( x 2,5). D Syngramma alismifolia, edge of fertile lamina with
venation and sorallines (x 1). E Taenitis blechnoides, portion
of fertile pinna with venation, receptacle, and soral lines
(x 1,5). F Cheilanthes /arinosa, part of fertile pinna from
abaxial side ( x 9). G Cheilanthes rn/opunctata, base of fertile
pinna from abaxial side (x 4). H Adiantopsis radiata, lamina
(xO,25) (A-B from Tardieu·Blot 1964; C-E from Holttum
1955; F original R. M. Tryon; G,H from R. M. Tryon 1964)
from which a new gametophyte develops at the beginning of a growing season.
10. Afropteris Alston
Fig. 124A,B
A/ropteris Alston, Bol. Soc. Brot. 11.30: 5 (1956) napsu ''Aropleris"].
Pteris auctt., p. p. min.
Pel/aea auct!., p. p. min.
Stem creeping, bearing entire, brown, lanceolate
scales. Leaves arising closely to moderately distantly,
nearly monomorphic. Petiole with a single U-shaped
vascular bundle, adaxially sulcate. Lamina 3-pinnatepinnatifid to 4-pinnate anadromous, subpentagonal,
glabrous; upper pinnae, pinnules, etc. gradually reduced and confluent. Basal basiscopic pinnules of basal pinnae enlarged, Veins pinnate in the ultimate,
Ajropteris . Onychium . Actiniopteris . Austrogramme
about Iigulifonn segments, the branches free, ending
behind or nearly at the margin, their ends hardly enlarged. Sterile margin entire or with a few teeth. Sori
occupying part of the margin of the segments, on both
sides or in smaller segments only on the anterior side;
sporangia borne on a submarginal vascular commissure, covered by reflex ed, narrow, entire lobes of the
margin. Soral trichomes none. Spores trilete, tetrahedral, rather triangular in distal view, somewhat irregularly tuberculate or rugose, especially on the distal
face, with more or less dense surface rodlets, with an
equatorial or nearly equatorial flange. Chromosome
number unknown.
Two species, terrestrial in forests at lower elevation,
A. repens (c. Chr.) Alston in tropical West Mrica and
A. barklyae (Baker) Alston in the Seychelles.
The genus has usually been associated with Pteris,
but clearly has a closer affinity with the Taenitidoideae. It is allied to Onychium in the following characters: adaxially sulcate axes, the grooves continuous between axes of different order, anadromous lamina,
sporangia on a vascular commissure lacking trichomes, and spore with a moderately to well-developed equatorial flange. It differs from Onychium especially in the presence of spicular cells in the epidennis
and the enlarged posterior pinnules of the basal pinnae, and in the spores. The consistently anadromous
lamina excludes it readily from Pteris.
239
C. Chr. is the only species that has the fertile segments
yellow-farinose beneath.
The following must be deleted from the revision by
Kilmmerle: the erroneous Peru locality cited under
Onychium vivlparum (Cav.) Kiimm. and the American
Onychium strictum Kunze, which is Anopteris hexagona (L) C.Chr.
12. Actiniopteris Link
Actiniopteris Link, Fil. Spec. Cult.: 79 (1841); Pichi Sennolli
(1962); KornaS et al. (1982).
Stem very short-creeping, with rather narrow, brownish, concolorous or bicolorous scales with a dark-sclerotic centre. Leaves monomorphic to rather dimorphic
(the fertile with longer segments). Petiole with 1 vascular bundle near the base, adaxially and abaxially sulcate, at least towards the apex. Lamina dichotomously
forked or branched into c. 6-30 long, narrow ultimate
segments, mostly glabrous to somewhat scaly, especially at the base, sterile veins free, ending nearly at the
margin, the ends hardly enlarged. Cells of the upper
epidennis with spiral thickenings. Sporangia borne in
a long soral line along a marginal commissure lacking
trichomes and connecting the vein ends, covered by a
continuous, well-modified marginal indusium. Spores
trilete, tetrahedral-globose, tuberculate, especially on
the distal face, with a prominent equatorial flange;
n=58.
11. Onychium Kaulf.
Onychium Kaulf., lahrb. d. Phannac. 1820: 45; KUmmerle
(1930); Ching (1934).
Stem short- to rather long-creeping, with narrow,
brownish, thin to dark-sclerotic, rigid scales. Leaves
nearly monomorphic to dimorphic (the fertile with
fewer and larger ultimate segments). Petiole with 2
(rarely 1) vascular bundles near the base, adaxially sulcate. Lamina 3- to 5-pinnate, anadromous, nearly glabrous or the fertile segments yellow-farinose beneath;
sterile veins free, ending at or near the margin, the
ends somewhat enlarged. Sporangia in a soral line
along a marginal commissure lacking trichomes and
covered by a continuous, well-modified marginal indusium. Spores trilete, tetrahedral-globose, tuberculate, distal face reticulate with prominent coarse ridges
parallel to the equatorial flange; n= 29, 58, 87.
A genus of perhaps eight species, distributed from
northeastern Mrica and Iran eastward to India, China,
Japan, and New Guinea. All occur in China except
Onychium divaricalum (Poir.) Alston, of Mrica and the
Near East.
In soral characters Onychium is very much like Pteris, but easily distinguished by the wholly anadromous
lamina architecture. Onychium siliculosum (Desv.)
A genus of five species, primarily of Mrica and adjacent islands, but also extending east to Mghanistan,
Nepal, India, and Ceylon. It grows in more or less xeric habitats and Actiniopleris radiala (Sw.) Link is especially well adapted to survive a hot and dry season.
When desiccated the lamina turns downward towards
the petiole.
13. Austrogramme Fournier
Austrogramme Fournier, Ann. Sci. Nat. V. 18: 278 (1873);
Hennipman (1975).
Asp/eniopsis Kuhn (1882).
Stem creeping, with short, brown bristles. Leaves monomorphic or nearly so. Petiole with 2 vascular bundles
near the base, adaxially sulcate. Lamina simple and
entire, 1-pinnate, or 2-pinnate to 2-pinnate-pinnatifid,
isodromous, glabrous; veins free or fonning costal areolae without included veinlets, extending nearly to the
margins, the ends not enlarged. Sporangia borne on
the veins in short to long, sometimes branChed, exindusiate soral lines, these with short, partly curved trichomes usually with enlarged apical cells. Spores trilete, tetrahedral-globose, often more or less depressed
between the aperture anns, with papillae or small tubercles associated with rodlets or with granulate deposit, without an equatorial flange; n=58.
240
Pteridaceae • Cheilanthoideae
A genus of five species in New Caledonia,
New Guinea, the Moluccas, and the New Hebrides.
Austrogramme may be divided into two sections: Sect.
Austrogramme with two species with a simple lamina,
and Sect. Aspleniopsis (Kuhn) Hennipm. with three
species with the lamina l-pinnate to 2-pinnate-pinnatifid.
14. Syngramma J. Smith
Fig. 124C,D
Syngramma J. Smith, London J. Bot. 4: 168 (1846); Holttum
(1975).
Toxopteris Trevisan (1877).
Craspedodictyum Copel. (1911).
Stem creeping, with dark, rigid bristles. Leaves monomorphic or nearly so. Petiole with 1-3 vascular bundles near the base, subterete or adaxially sulcate, narrowly alate. Lamina simple, entire or 3- to rarely
6-foliolate, anadromous (where applicable), glabrous;
veins free to near the margin where they join to form a
marginal strand or 1 or 2 series of areolae without included veinlets. Sporangia borne along the veins in
rarely short, usually long, sometimes branched, exindusiate soral lines, or rarely along the anastomosing
veins, usually with small trichomes with an enlarged,
often dark red apical cell among the sporangia. Spores
trilete, tetrahedral-globose, usually depressed between
the aperture arms, with low tubercles, with papilJae
more or less overlaid by rodlets, without an equatorial
flange; n=58, 116.
Malaya to the Philippine Islands, New Guinea, and
east to Fiji and the Caroline Islands (Palau). About 15
species, seven of them in Borneo.
Three sections may be recognized: Sect. Syngramma
with small areolae near the margin of the simple lamina, Sect. Toxopteris (Trev.) C. Chr. with the vein ends
joined to form a strand along the margin of the simple
lamina, and Sect. Craspedodictyum (Cope!.) R. Tryon,
with a usually 3- to 5-foliolate lamina.
15. Taenitis WiJldenow ex Sprengel
Fig. 124E
Taenitis Willdenow ex Sprengel, Anleit. z. Kenntn. d. Gew.
3: 374 (1804); Holttum (1968,1975).
Schizo/epton Fee (1850).
P1atytaenia Kuhn (1882).
Ho/ttumiella Copeland (1941).
Stem rather short-creeping to long-creeping, with dark,
rigid bristles. Leaves monomorphic or slightly to
strongly dimorphic (the fertile with narrower lamina or
pinnae). Petiole with 1, 2, or 4 vascular bundles near
the base, adaxially sulcate, at least towards the apex.
Lamina simple, entire and nearly orbicular to elongate, or l-pinnate, glabrous, veins anastomosing to
form oblique areolae without included veinlets from
the costa to the margin, or free at the margin. Sporan-
gia borne along the veins in exindusiate soral lines,
following the anastomosing veins, or partly so, sometimes of varying length with some very short, or in a
long inframarginal soral band, or confined to the fork
of free veins, usually with small trichomes among the
sporangia, sometimes these with an enlarged apical
cell or curved. Spores trilete, tetrahedral-globose, with
low, compact tubercles usually with projecting rodlets,
usually with an equatorial flange; n=44, 108, 110, 114.
A complex genus of about 15 species, distributed
from southern India and Ceylon to Hainan, through
Malesia to northern Queensland and Fiji. Nine of the
species occur in Borneo.
The four sections of Holttum (1975) are not recognized because major chracters such as lamina architecture, the flange on the spores, and the relation of the
sporangia to the venation are insufficiently correlated.
Subfamily Cheilauthoideae
(J. Smith) W.-c. Shieh, J. Sci. Eng. 10: 211 (1973). - Tribe
Cheilantheae J. Smith, Hist. Fil.: 277 (1875).
Terrestrial or rupestral, mostly small plants. Stem erect
to long-creeping, with a solenostele, or this grading to
a dictyostele, bearing scales sometimes intergrading to
trichomes. Petiole with 1 or 2 vascular bundles near
the base, terete to adaxially sulcate or two-ridged.
Lamina simple to 1- to 5-pinnate, gradually reduced to
the apex or sometimes with stalked ultimate segments
and an odd terminal segment, glabrous, pubescent,
scaly, glandular, or sometimes farinose beneath; veins
usually free or sometimes anastomosing. Sporangia
borne along the veins in exindusiate soral lines, or
covered by a modified marginal indusium, or on a
marginal commissure, or at the vein ends, or in a narrow to broad band on anastomosing veins and between them, without special trichomes among the
sporangia, or rarely with a few inconspicuous ones.
Spores trilete, often cristate, without an equatorial
flange.
CHARACTERS OF RARE OCCURRENCE. Sporangia are
rarely solitary on a vein end in Cheilanthes and Notholaena, and then are unusually large, nearly sessile,
and with a broad annulus (Christensen and Ching
1933; Pichi Sermolli 1946).
MORPHOLOGV AND ANATOMV. The stem of Cryptogramma stelleri (Gme!.) Prant! is unique among ferns.
The short-creeping living stem is yellow-green, succulent and brittle. It shrivels in its second year so that the
living tissues are always less than 2 years old.
The leaves are diverse. Most unusual are the
scrambling leaves of Pellaea ovata (Desv.) Weath.
more than 1 m long, and the fertile leaves of Llavea
cordifolia Lag. with dimorphic segments, the fertile ap-
Key to the Genera
ical and the sterile basal. The stomata are mostly
anomocytic. In Cheilanthes they are rarely also polocytic, and in Coniogramme they are both anomocytic
and polocytic (van Cotthem 1970).
Croziers are fully circinate as the leaf unfolds in
most ferns but in Bommeria (Haufler 1979), in Cheilanthes (Knobloch 1965), and in Paraceterach, while
the lamina bud is circinate, the petiole is bent downward distally instead of being coiled.
Genera such as Cheilanthes, Paraceterach, Hemionitis, Bommeria, and Trachypteris often have an exceptionally dense lamina indument of trichomes or scales.
GAMETOPHYTE. The gametophyte is known in several
genera (Nayar and Bajpai 1964; Nayar and Kaur
1971; Nayar and Devi 1973). It is epigeal, cordate,
with thin margins and a thickened central cushion
bearing mostly archegonia, with antheridia among the
rhizoids or scattered, sometimes marginal. Old
gametophytes often become irregularly lobed.
The occurrence of indument on gametophytes of
Notholaena and Cheilanthes, similar to that of the
sporophyte, is an unusual feature. Notholaena gametophytes with farinose glands are reported by A. Tryon
(1947) and Giauque (1949) and farinose glands or
trichomes in Cheilanthes by Giauque (1949).
KARYOLOGY. The chromosome numbers of n=29 or
30 or multiples are nearly consistent within the subfamily, as in most members of the family Pteridaceae.
Reports of n=27 in Notholaena and n=56 in Cheilanthes are evidently unusual cases of aneuploid numbers.
AFFINITY. The subfamily Cheilanthoideae is clearly related to other groups of the Pteridaceae by the nearly
uniform chromosome numbers and spore characters.
However, evolutionary lines within the subfamily and
the relations of these to other groups in the Pteridaceae are scarcely known. Alloploidy and extinction
have undoubtedly obscured the evolutionary history.
DISTRIBUTION AND ECOLOGY. The subfamily has an essentially worldwide distribution, with its greatest development in or near the tropics of North and South
America Six of the 12 genera are confined to America,
Cheilanthes, Pellaea and Doryopteris are nearly pantropical, Cryptogramma is amphitropical, and Paraceterach and Coniogramme are confined to the Old
World.
The species grow in a variety of habitats, but especially those of Cheilanthes, Pellaea, and Notholaena
often constitute a significant element of the flora of
xeric and semi-xeric regions.
241
KEy TO THE GENERA.
1. Fertile segments with a modified marginal indusium that
wholly or mostly conceals the sporangia
2
- Margin of the fertile segments not modified, or if so then
not or hardly concealing the sporangia
7
2. Fertile leaves hemi-dimorphic, the apical portion with
narrow fertile segments, the basal with broad sterile segments (Fig. 127 E)
27. LIIlvea
- Fertile leaves with all of the segments fertile, or the sterile
segments similar to the fertile
3
3. Stem not persistent. surviving 1 year and part of the next
26. Cryptogramma
- Stem persistent. indurated
4
4. Lamina pinnate, with an odd terminal segment. most ultimate segments sessile or stalked
19. Pellaea
- Lamina of various architecture, usually pinnate and the
lamina gradually reduced to the apex and the ultimate
segments usually adnate or joined
5
5. Lamina entire, lobed, palmate, or usually pedate and not
farinose beneath, and the petiole terete or rarely adaxially
20. Doryopteris
flattened or very slightly ridged
- Lamina radiate or pinnate, or pedate and then farinose
beneath, or the petiole adaxially sulcate or two-ridged 6
6. Spores cristate, rugose, verrucate, or granulose; lamina
axes terete or adaxially flattened, Of, if the spores are echinate or the axes adaxially two-ridged, then the ultimate
16. OJeiJanthes
- Spores echinate, lamina axes adaxially two-ridged, many
segments adnate or symmetrical
to all ultimate segments subsessile to stalked and asymmetrical
17. Adiantopsis
7. Sporangia borne in a narrow to broad marginal band on
and between anastomosing veins
24. Trachypteris
- Sporangia borne only on veins, on a vascular receptacle,
or on a marginal commissure
8
8. Leaves strongly dimorphic, lamina 2-pinnate or more
complex
26. Cryptogramma
- Fertile and sterile leaves monomorphic Or nearly so, or, if
dimorphic, then simple
9
9. Lamina with large, herbaceous, stalked ultimate segments,
the lamina and pinnae with an odd tenninal segment;
spores nearly smooth
25. Coniogramme
- Lamina simple or with small ultimate segments, or with
large segments and the spores with pronounced surface
elements
10
10. Lamina farinose beneath, or, if indument lacking, then 2to 4-pinnate and with an odd terminal segment
18. Notholaena
- Lamina not farinose beneath, if 2- to 3-pinnate then
pubescent beneath or gradually reduced to the apex
11
11. Lamina pedate, 2- to 3-pinnatifid, sporangia along all of
the veins or at least the apical one-fourth 23. Bommeria
- Lamina simple, pedately lobed, or pinnate, or, if pedate,
12
then the sporangia confined to the vein ends
12. Sporangia in sori or single at or nearly at the vein ends
16. OJeiJantlres
- Sporangia in soral lines along anastomosing veins or
along the sometimes short free veins
13
13. Lamina densely pubescent or scaly beneath, the indument
21. Paracetemdr
concealing the leaf tissue
- Lamina glabrous, slightly scaly, or moderately to sparsely
pubescent beneath, the indument not concealing the leaf
tissue
14
14. Sporangia in sori at or near the vein ends or on the apical
portion of the veins
19. PeUaea
- Sporangia in long soralHnes along the veins
22. Hemionitis
242
Pteridaceae • Cheilanthoideae
16. Cheilanthes Swartz
Fig. 124F,G
Cheilanthes Swartz, Syn. Fil. 5: 126 (1806), nom. conserv.;
R. Tryon (1964); Quirk et al. (1983); Anthony (1984); Tryon
and Tryon (1982).
Allosorus Bernhardi (1805).
Myriopteris Fee (1852).
A/euritopteris Fee (1852), Wu (1981, 1983), Saiki (1984).
Cheilop/ecton Fee (1857).
Cosentinia Todaro (1866), Pichi Senn. (1985).
Neurosoria Kuhn (1869).
Mildella Trevisan (1876); Hall and Lellinger (1%7).
Sinopteris C. Chr. & Ching (1933).
Negripteris Pichi Senn. (1946).
Aspidotis Copel. (1947).
Lepto/epidium Hsing & S.K. Wu, Acta Bol. Yunnan. 1: 115
(1979).
Notholaena auctt., p.p., e.g. R. Tryon (1956).
Stem erect to long-creeping, with light brown, concolorous or bicolorous to wholly dark-sclerotic, linear to
ovate-Ianceolate, usually thin scales. Leaves usually
monomorphic, to slightly or rarely strongly dimorphic
(the fertile erect, more complex than the sterile).
Petiole with 1 or 2 vascular bundles near the base,
terete to adaxially sulcate or two-ridged. Lamina 1- to
5-pinnate, or less often pedate and to at least 3-pinnatifid, anadromous or isodromous, rarely catadromous,
glabrous, pubescent, scaly, glandular, or white-, yellow- or orange-farinose beneath; veins free or rarely
anastomosing without included veinlets, ending at or
near the margin, the ends not to clearly enlarged.
Sporangia usually in sori at the vein ends, or rarely
single or in soral lines along an incomplete to complete marginal commissure, the margin unmodified to
strongly modified as a marginal indusium covering
one or more sori or a soral line. Spores trilete, tetrahedral-globose to globose, both faces usually prominently cristate, sometimes rugose or verrucate, rarely
tuberculate or echinate, without an equatorial flange;
n=29, 30, 56, 58, 60.
A complex genus of about 150 or more species. It is
essentially worldwide in distribution but poorly represented or absent in boreal regions. It is especially welldeveloped in xeric and semi-xeric habitats in Mexico
and the Andes, also in southern Mrica, and to a lesser
extent in the Sino-Himalayan region and Australia.
Numerous, mostly small and sometimes monotypic
genera have been segregated, but these represent only
a portion of the diversity within the genus. Perhaps
30 groups could be recognized in Chei/anthes. Ten
groups and seven morphologically isolated species are
recognized in America by Tryon and Tryon (1982). A
formal infrageneric classification, based on a review of
all characters, has not been developed.
The problems of classification relate to highly distinctive species or small groups and to character convergence. The following species, for example, are not
evidently related to any others. Cheilanthes vellea
(Aiton) F. v. Muell., with spores similar to those of Pityrogramma; C. sinuata (Sw.) Domin, with 2 vascular
bundles in the petiole; and C. eoriaeea Dcne., with the
indurated marginal indusia overlapping or nearly so.
Examples of convergent characters that occur in unrelated species include the following: a pedate lamina in
Cheilanthes cone%r (Langsd. & Fisch.) R & A. Tryon
and C. geraniifolia (Weath.) R. & A. Tryon; a lamina
with farinose indument beneath in Chei/anthes argentea (GmeJ.) Kunze and C. aurantiaea Moore; long,
narrow fertile segments with long, modified indusia in
Chei/anthes smithii (C. Chr.) R Tryon and C. si/iquosa
Maxon; single sporangium in a sorus in Chei/anthes
grevilleoides Christ and C. rigida (Sw.) Domin.
17. Adiantopsis Fee
Fig. 124H
Adiantopsis Fee 5' Mem. Gen. Fil.: 145 (1852); Tryon and
Tryon (1982).
Chei/anthes auctl., p.p., e.g., Copeland (1947).
Stem usually erect to decumbent, to long-creeping,
bearing scales that are usually bicolorous with brown
margins and a dark-sclerotic centre or are wholly
darksclerotic. Leaves monomorphic. Petiole with 1-3
vascular bundles near the base, it and the lamina axes
ad axially two-ridged. Lamina 1- to 4-pinnate, anadromous, or sometimes pedate or radiate, glabrous to
somewhat short-pubescent; veins free, usually ending
well behind the margin, the ends somewhat enlarged.
Sori separate on the vein ends, each (rarely two)
covered by a lunate or roundish marginal indusium.
Spores trilete, tetrahedral-globose, echinate, without
an equatorial flange; n=60.
A tropical American genus of about seven species.
The small Adiantopsis minutula Sehnem has leaves to
c.5 cm long, the petiole and other axes terete, the vein
ends not enlarged, and echinulate spores.
18. Notholaellll R Brown
Fig. 125A-D
Notho/aena R. Brown, Prodr. A. Nov. Holl.: 145 (1810);
R. Tryon (1956); 1l:yon and Tryon (1982).
Argyrochosma (J. Smith) Windham. Amer. Fern J. 77: 38
(1987), based on Notholaena sect. Argyrochosma J. Smith
(1841).
Chei/anthes auell., p.p., e.g., Copeland (1947).
Stem suberect to creeping, with rufous, thin, con colorous scales or the scales bicolorous with a dark-sclerotic centre. Leaves monomorphic. Petiole with 1 vascular bundle near the base, usually terete, rarely adaxialIy flattened, sulcate, or slightly ridged. Lamina 1- to
4-pinnate or rarely pedate and 2-pinnatifid or 2-pinnate-pinnatifid, usually catadromous or isodromous,
usually white- or yellow-farinose beneath and often
with scales and (or) trichomes, rarely glabrous; veins
free, ending behind the margin, the vein ends not to
Cheilanthes • Adiantopsis. Notholaena . Pellaea
243
H
D
E
C
G
~'r
ft
,
~
.<1
'~l
I)
,
".
.'
"'
1({r',
1-
A!I '/
B
F
clearly enlarged. Sori borne on the vein ends, or rarely
single or along most of a vein, exindusiate, the margin
not or only slightly modified. Spores trilete, globose to
tetrahederal-globose, usually cristate or low rugose, to
plain with a granulate deposit, without an equatorial
flange; n=27, 29, 2n=60.
An American genus of about 40 species, distributed
from the central United States south to Argentina and
Chile. Most of the species occur in the southwestern
United States and Mexico.
Two sections may be recognized: Sect. Notholaena
with 22 species, with dark-sclerotic (at least the central
portion), rigid stem scales and blackish, rather granulate spores; Sect. Argyrochosma J.Smith with 17 species, with lax, rufous stem scales and brownish, cristate
spores.
Species 22-58 of the revision by R. Tryon (1956)
form the present genus Notholaena. Species 1-21 of
that treatment and nearly all species of the Old World
have been placed in Cheilanthes; N.marantae is now
included in Paraceterach.
Fig. 125A-I. Pteridaceae. A,B Notholaena aschenbomiana.
A Habit (x 0,5). B Two segments from abaxial side, left bearing scales, right with scales removed to show venation (x 6).
C Notholaenafendler~ habit (x 0,5). D Notholaena lumholtzi~
fertile segment from abaxial side ( x 4,5). E Pel/aea rufa, fertile segment, abaxial side, indusium reflexed at left (x 12).
F, G Doryopteris pedata, F Fertile leaf ( x 0,25), G Edge of
fertile segment with venation and receptacle (x 5,5), H Doryoptens paradoxa, part of sterile segment with venation
( x 5,5). I Doryopteris omithopus, edge of fertile segment with
receptacle and indusium (schematic) (A-D from R. M. Tryon
1956; E from A. F. Tryon 1955; F-I from R, M, Tryon 1942)
19. Pe/Ioea link
Fig. 125E
Pel/aea Link, Fit. Spec. Cult.: 48, 59, (Sept. 1841), nom. conserv.; A. Tryon (1957); Anthony (1984); Tryon and Tryon
(1982).
Platyloma J. Smith (Aug. 1841).
Ormopteris J. Smith (1875).
Stem decumbent to rather long-creeping, with narrow
to ovate-lanceolate, rufous, rather thin and concol-
244
Pteridaceae • Cheilanthoideae
orous scales, or the scales bicolorous with a dark-sclerotic centre. Leaves nearly monomorphic. Petiole with
1 vascular bundle near the base, terete to adaxially sulcate or rarely ridged. Lamina 1- to 4-pinnate, nearlyalways with an odd terminal segment, anadromous, a
few species catadromous or isodromous, ultimate segments sessile or short-stalked, usually glabrous or
somewhat pubescent or with a few scales; veins free,
or in a few species anastomosing without included
veinlets, ending somewhat behind the margin, the vein
ends not or clearly enlarged. Sori borne at the vein
ends, or on their apical portion, or sporangia in a soral
line on an extended marginal commissure, rarely with
short, often farinaceous glands among the sporangia,
partly to wholly covered by a slightly to clearly modified marginal indusium. Spores trilete (or alete in
apogamous plants), tetrahedral-globose or globose,
usually low-rugose, sometimes with granulate deposit,
or cristate, or rarely low-tuberculate to verrucate, without an equatorial flange; n= 29,30, 58.
A genus of about 35 species, mainly centred in the
southwestern United States and Mexico and in southern Africa. It is distributed in North and South America, Africa and eastward to India and New Zealand;
also in northeastern Spain and introduced in the
Azores.
Four sections may be recognized. Section Pellaea:
stem short- to long-creeping, the segments usually not
articulate, the indusium slightly to moderately differentiated, and the spores more or less rugose to cristate;
about 16 species in America and one in Africa. Section
Ormopteris (J. Smith) R. & A. Tryon: stem short- to
long-creeping, the segments not articulate, the indusium strongly differentiated, and the spores rugose,
cristate or tuberculate; six species of South America.
Section Holcochlaena Hooker & Baker: stem short, the
segments articulate, the indusium strongly differentiated, and the spores rugose, rugulose, or verrucate;
about 10 species of Africa to India and Ceylon. Section Platyloma (J. Smith) Hooker & Baker: stem slender, long-creeping, the segments articulate, the indusium rather moderately differentiated, and the
spores echinate; three species, India and Ceylon east
to Tasmania, New Zealand, and New Caledonia.
20. Doryopteris J. Smith
Figs. l06E, 125F-I
Doryopleris J. Smith, J. Bot. (Hooker) 4: 162 (1841); R. Tryon
(1942); Brade (1965); Tryon and Tryon (1982).
Cassebeera Kaulfuss (1824).
Tryonella Pichi Senn. (1974).
Stem decumbent to suberect to rarely long-creeping,
usually with rigid, tawny, narrow, concolorous scales
or the scales bicolorous with a dark-sclerotic centre, or
to ovate-Ianceolate and with lighter margins. Leaves
monomorphic to dimorphic (the fertile with narrower
segments and often a more complex lamina). Petiole
with 1 or 2 vascular bundles near the base, terete to
adaxiaUy sulcate or rarely slightly ridged. Lamina simple and cordate to pedate or palmate and pinnatifid to
rarely 2-pinnatifid, glabrous; veins free or anastomosing without included veinlets, the veins ending near
the margin, the ends slightly to clearly enlarged. Sporangia usually in long soral lines on a marginal commissure, or in an inframarginal band on and between
anastomosing veins, or rarely in roundish sori at the
vein ends, covered by a continuous modified marginal
indusium. Spores trilete, tetrahedral-globose, usually
cristate, or plain to slightly rugose, rarely echinate,
without an equatorial flange; n=30, 60, 116.
A genus of 25 species, distributed from tropical
America to the Old World, eastward to New Guinea.
Most of the species occur in tropical America and are
centred in southeastern Brazil. Brade (1965) treated
25 species in Brazil. 'TWo sections may be recognized.
Section Lytoneuron Kl. with linear stem scales, 2 vascular bundles in the petiole, and free veins; 13 species
in America. Doryopteris triphylla (Lam.) Christ may
have ovate- laneeolate stem scales; it and sometimes
D. paradoxa (Fee) Christ have a single vascular bundle
in the petiole; and D.omithopus (Hooker & Baker)
J. Smith has anastomosing veins. These species are
clearly members of Section Lytoneuron in their other
characters. Section Doryopteris with broader stem
scales, 1 vascular bundle in the petiole and anastomosing veins; 12 species, 7 in America and 5 from Madagascar east to New Guinea.
Doryopteris concolor (Langsd. & Fisch.) Kuhn and
its allies (species 22- 24 of R. Tryon 1942) are now
placed in Cheilanthes.
21. Paraceterach (F. v. Mueller) Copel.
Paraeelerach (F. v. Mueller) Copel., Gen. Fil: 75 (1947);
R Tryon, Amer. Fern J. 76: 86 (1987), based on: Grammilis
section Paraeeleraeh F. v. Mueller (1866).
Gymnopleris auctt., e.g. Underwood (1902); Copeland (1947).
Hemionitis auctt., e.g. Mickel (1974).
Stem short-creeping to very compact, with usually
light brown to somewhat reddish, narrow, concolorous
scales, or the scales bicolorous with a narrow to broad,
dark-sclerotic centre, sometimes grading into trichomes. Leaves nearly monomorphic. Petiole with
1 vascular bundle near the base, terete. Lamina 1- to
2-pinnate and sometimes with an odd terminal segment, isodromous, densely pubescent or scaly, especially beneath; veins free or occasionally partly anastomosing without included veinlets, ending near the
margin, the ends scarcely enlarged. Sporangia borne
along the veins in short to long, sometimes branched,
exindusiate soral lines. Spores trilete, tetrahedralglobose to globose, cristate or sometimes echinate, rarely
nearly plain, without an equatorial flange; R= 29,30.
Doryopteris· Paraceterach . Hemionitis
Seven species, in the Canary Islands and Madeira,
southern Europe, Ethiopia, northern India to China
and Australia. Most species of Paraceterach have a
multicipital stem and croziers that are only partly
circinate; the genus also differs from the American
Hemionitis in the lamina indument.
The two elements in the genus may represent derivatives from different sources in the Old World. Paraceterach muelleri (Hooker) Cope\., P. reynoldsii (F. v.
Muell.) Tindale, P. delavayi (Baker) R. Tryon, ana P.
marantae (L.) R. Tryon, have the lamina 1-pinnate or
1-pinnate-pinnatifid and densely scaly beneath. Paraceterach vestita (Hooker) R. Tryon, P. bipinnata (Christ)
R. Tryon, and P. sargentii (Christ) R. Tryon, have the
lamina 1- or 2-pinnate and densely pubescent beneath.
"Hemionitis" arifolia (Burm.) Moore is similar to Paraceterach in some technical characters, but differs in its
dimorphic leaves and simple, cordate to hastate lamina with sparse indument. It is not readily included in
any recognized genus.
245
Fig. 126A-F. Pteridaceae. A,B Hemionitis palmata. A Fertile leaf with venation ( x 0,5). B Sterile leaf with proliferous
buds (x 0,5). C, D Bommeria hispida. C Habit (x 0,7).
D Fertile segment from abaxial side (x 3). E, F Trachypteris
induta. E Leaf (x 0,5). F Abaxial side of segment (xl,5)
(A,B,E,F from R. M. Tryon 1964; C,D from Knobloch and
Correll 1962)
22. Hemionitis L.
Fig. 126A,B
Hemionitis L., Spec. Plant. 2: 1077 (1753), Gen. Plant. ed.
5: 485 (1754); Tryon and Tryon (1982).
Gymnopteris Bemhardi (1799).
Gymnogramma Desv. (1811), as to type.
Stem erect to short-creeping, with light brown, thin,
concolorous scales or the scales bicolorous with a dark
sclerotic centre, grading into brownish trichomes.
Leaves monomorphic to dimorphic (the fertile erect,
the sterile forming a rosette). Petiole with 1 vascular
bundk near the base, terete to adaxially sulcate. Lami-
246
Pteridaceae • Cheilanthoideae
na entire to deeply 7-lobed and pedate, or 1- to 2-pinnate, or rarely nearly 3-pinnate, and with an odd terminal segment, approximately isodromous, slightly to
moderately pubescent beneath and sometimes with a
few scales; veins free or partly to completely anastomosing without included veinlets, the ends nearly
reaching the margin, not or clearly enlarged. Sporangia borne along the veins in long, usually branched,
exindusiate soral lines. Spores trilete, tetrahedral-globose, cristate, echinate, or verrucate to irregularly tuberculate, without an equatorial flange; n= 30, 60.
A genus of tropical America with seven species.
There are three elements in the genus which may have
been derived from different American sources. Hemionitis subcordata (Davenp.) Mickel has a strawcoloured, sulcate, glabrous petiole and a 1-pinnate
lamina with partly areolate venation. Hemionitis TUfa
(L.) Sw. and H. tomentosa (Lam.) Raddi have a
dark-coloured, terete petiole with strongly dimorphic
pubescence and a 1- to 3-pinnate lamina with free venation. Hemionits levyi Fourn., H.palmata L., H.pinnatifida Baker, and H. elegans Davenp. have a darkcoloured, mostly terete petiole that is glabrous or pubescent with long trichomes intergrading to short ones,
and a simple to subpedately or pedately lobed lamina
with wholly areolate venation.
For the Old World "Hemionitis" arifolia, see under
Paraceterach.
23. Bommeria Fournier
Fig. 126C,D
Bommeria Fournier, Diet. Bot. (Baillon) 1: 448 (1876); HaurIer (1979); Tryon and Tryon (1982).
Stem short- to long-creeping, with lanceolate to ovatelanceolate, thin, light brown scales or the scales bicolorous with a dark-sclerotic centre or nearly wholly
dark-sclerotic, one species also with trichomes. Leaves
monomorphic. Petiole with 1 vascular bundle near the
base, terete or adaxially sulcate. Lamina pedate, 2- to
3-pinnatifid, pubescent and with some scales; veins
free or partly anastomosing without included veinlets,
ending nearly at the margin, the ends not enlarged.
Sporangia borne in exindusiate soral lines along the
veins near the margin. Spores trilete, tetrahedral-globose, slender strands forming a cristate to reticulate
surface, without an equatorial flange; n=30.
An American genus of four species, distributed
from the southwestern United States south to Costa
Rica_ All of the species occur in Mexico.
Stem decumbent to erect, with broad, brownish or
sometimes pink, nearly concolorous scales. Leaves
monomorphic, the lamina pinnatifid and pedate, or dimorphic (the fertile erect, the lamina 1-pinnate, the
sterile leaves in a rosette, nearly sessile, the lamina
obovate, entire). Petiole with 1 vascular bundle near
the base, terete or adaxially sulcate. Lamina densely
scaly beneath, glabrous to sparsely scaly above; veins
anastomosing without included veinlets. Sporangia
borne in an exindusiate soral band on and between the
veins. Spores trilete, tetrahedral-globose, strongly cristate, without an equatorial flange; n=either 29 or 30.
A genus of three species. Trachypteris induta (Maxon) R. & A. Tryon and T. pinnata (Hook. f.) C. Chr. are
South American, the latter also in the GalApagos Islands. The sterile leaves of T. drakeana C. Chr. of
Madagascar have an apical bud by which the plants
may reproduce vegetatively. Scale cells with peculiar
striations have been figured by Ballard (1962).
25. Coniogramme Fee
Coniogramme Fee, 5' Mem. Gen. Fil.: 167 (1852), nom. conserv.; Hieronymus (1916); Dixit and Das (1979).
Stem short- to moderately long-creeping, with brown,
more or less lanceolate, rather rigid scales. Leaves
monomorphic. Petiole with 1 large vascular bundle
near the base, adaxially sulcate. Lamina 1- to 3-pinnate, with large ultimate segments, with an odd terminal segment, variously anadromous, isodromous, or
catadromous, usually glabrous or somewhat pubescent; veins free, or anastomosing near the costa without included veinlets, ending near the margin, the ends
enlarged. Sporangia borne on the veins in usually
long, sometimes branched, exindusiate soral lines, or
following the anastomosing veins; usually sparse,
hardly visible, few-celled, simple trichomes often present among the sporangia. Spores trilete, tetrahedralglobose with prominent angles, more or less depressed
between the aperture arms, low tuberculate or rugulose, with rodlets, without an equatorial flange; n=30,
60.
Twenty or more species, distributed from tropical
Africa east to the Himalaya, China, Java, and Japan,
and in the Pacific to the Hawaiian Islands and Samoa.
Coniogramme americana Maxon of Mexico and
Guatemala is Hemionitis subcordata (Davenp.) Mickel.
26. Cryptogramma R. Brown
Fig. 126E,F
Cryptogramma R. Brown, Bot. Append. Franklin, Narrative
Polar Sea, Addenda: 767 (1823); Fernald (1935); Tryon
and Tryon (1982).
Allosorus auct. non Bernh.
Trachypteris Christ, Denkschr. Schweiz. Naturf. Ges. 36: 150
(1899); Tryon and Tryon (1982).
Saffordia Maxon (1913).
Stem decumbent or short-creeping, with small to rather large, more or less concolorous, brownish, thin
24. Trachypteris Christ
Bommeria· Trachypteris· Coniogramme. Cryptogramma . Ilavea
scales. Leaves somewhat to strongly dimorphic (the
fertile more erect and with entire rather than toothed
segments). Petiole with 1 vascular bundle near the
base, adaxially sulcate. Lamina 2-pinnate to 2-pinnatepinnatisect, anadromous, glabrous, or the fertile sometimes yellow-farinose beneath; veins free, ending at or
near the margin and the ends scarcely enlarged, or behind the margin and the ends enlarged. Sporangia
borne in soral lines along the veins, or in sori at the
vein ends, more or less covered by a modified marginal indusium. Spores trilete, tetrahedral-globose with
prominent angles, verrucate, without an equatorial
flange; n= 30, 60.
A genus of two distinctive species, primarily boreal
and austral. It occurs in the United States and Canada,
northwest to Alaska, in Europe from Spain to the Balkan, Scandinavia, and the Urals, and eastward to the
Himalaya, China, and northeast Asia; also southern
Chile and adjacent Argentina.
The stem of Cryptogramma crispa (L.) R.Br. is short,
indurated, scaly, and persistent while in C. stelleri
(Gmelin) Prantl it is short-creeping, brittle-succulent,
greenish-yellow, nearly naked, and withers after the
first year. Cryptogramma stelleri can be recognized as
Sect. Homopteris (Rupr.) C. Chr.
27. Uavea Lagasca
Fig. 127E
Llavea Lagasca, Gen. Spec. Plant.: 33 (1816); Tryon and Try-
on (1982).
Stem decumbent to nearly erect, with large, dark
brown to atropurpureous, rigid scales. Fertile leaves
hemi-dimorphic (the apical fertile portion with longer
and narrower segments). Petiole with dense, large,
whitish-yellow scales at the base, with 1 vascular bundle near the base, adaxially sulcate. Lamina 2- to
4-pinnate, anadromous, with an odd tenninal segment; sterile segments often glaucous beneath, the fertile often glandular or white-farinose beneath; veins
free, extending to the margin, the ends not enlarged.
Sporangia borne in short soral lines along the veins,
more or less covered by a modified marginal indusium. Spores trilete, tetrahedral-globose, irregularly tuberculate, without an equatorial flange; n=29.
A monotypic genus of Mexico and Guatemala.
Uavea cordifolia Lag. is of uncertain relationship within the subfamily.
Subfamily Adiantoideae
(C. Presl) R. Tryon, Amer. Fern J. 76: 184 (1987).
Adianteae [as Tribe Adiantaceaej C. Presl, Tent. Pteridogr.: 139 (1836).
Terrestrial or rupestral ferns of small to large (over 2 m
high) size. Stem suberect to short- or long-creeping,
247
with a solenostele or this grading to a dictyostele, with
acicular to ovate-lanceolate, brownish or darker,
usually rather thin, dull to iridescent scales. Leaves
monomorphic or nearly so. Petiole with 1 or 2 vascular
bundles near the base, usually adaxially sulcate. Lamina simple and entire to sagittate or 1- to rarely 6-pinnate, very predominantly pronouncedly anadromous,
or the petiole furcate apically into two strongly recurved rachises, commonly with an odd tenninal segment or pinna, usually glabrous, less often glaucous,
pubescent, scaly, glandular, or farinose beneath; veins
usually free, or sparingly to fully anastomosing without included veinlets, ending at or near the margin, the
ends not to somewhat enlarged. Sporangia borne
along the veins in short sorallines, or also between the
veins, on a strongly recurved, orbicular to long, modified indusioid margin, the capsules facing the abaxial
side of the lamina. Spores trilete, tetrahedral-globose
with prominent angles, more or less depressed between the aperture arms, irregularly tuberculate or
rugose below irregular sheaths, without an equatorial
flange.
A single genus.
CHARACTERS OF RARE OCCURRENCE. In Adiantum the
veins of sterile segments rarely end in a definite tooth,
as in Adiantum deflectens Mart. and A. capillus-veneris
L. In most species the veins end in a sinus between the
definite or indefinite teeth.
MORPHOLOGY AND ANATOMY. Nayar (1962) presented
a detailed account of the morphology and anatomy of
Adiantum, based on study of many species.
The stem has a solenostele in species with widely
spaced leaves; it has transitions to a dictyostele in species with closely spaced leaves and a perforated
dictyostele in Adiantum capillus-veneris L. The rachis is
rarely prolonged into a rooting tip. The petiole is
usually dark brown to blackish and highly polished, as
is the rachis and other lamina axes. Some species have
articulate ultimate segments and only the axes remain
in old leaves.
Young leaves of Adiantum are generaly tinged with
red. The upper and lower epidennis are usually
chlorophyllous and cells especially of the upper epidermis have pronounced internal projections (papillae), and in some species the mesophyll is lacking
(Wylie 1948). There are especially thick walled cells in
the epidennis, usually associated with the veins which
have been referred to as false veins or spicular cells.
Nayar (1962) pointed out differences between them
and the spicular cells of the Vittariaceae and did not
regard them as evidence for a relation between the two
groups. The stomata are anomocytic (van Cottbem
1970).
Trichomes of 1 or 2 cells, with a globular apical cell,
have been described from among the sproangia in a
248
Pteridaceae . Adiantoideae . Pteridoideae
Fig. 127 A-E. Pteridaceae. A Adiantum philippeme, fertile
lamina (x 0,5). B Adiantum macroc1adum, base of fertile pinna from abaxial side (x 1). C Adiantum poeppigianum, fertile
pinna from abaxial side (x 0,5). D Adiantum davidi~ pinnule
from abaxial side, soral lobe lifted to show the sporangia
(x 5,8). E Uavea cordifolia, base of petiole and upper part of
fertile lamina (xO,5) (A from Scamman 1960; B,C from
R M. Tryon 1964; D from Ching 1935; E from A. R. Smith
1981)
ion and thin borders. Antheridia are mostly in the basal region among rhizoids, the archegonia more apical
on the thickened portion.
few species, including Adiantum tenerum Sw. (Nayar
AFFINITY.
1962).
Kachroo and Nayar (1953) discussed
the gametophyte of Adiantum in detail, based on a
study of seven species and a review of other reports. It
is cordate, symmetrical, with a thickened central cushGAMETOPHYTE.
KARYOLOGY. There are many chromosome counts for
Adiantum, exhibiting both aneuploid and polyploid
series, for example, n=29, 30, 57, 58, 60, 90,114,116,
122, 150, and c. 180. Walker (1985) discussed SOme of
the variations.
Adiantum is evidently an old and morphologically isolated genus, with general relations to the
Pteridaceae but without clear alliance to any particular
group within it.
DISTRIBUTION AND ECOLOGY. The genus is pantropical
with extensions northward to Newfoundland, Alaska,
Adiantum· Ochropteris . Anopteris
and northeastern Asia, and southward to southern
South America, southern Africa, and New Zealand.
Many species grow in primary or secondary forest,
others on cliffs or on rocky slopes. The species centre
in the mountains of the American tropics. Several
neotropical species are adventive in the paleotropics,
especially in oil palm and tea plantations.
249
- Lamina more than once pinnate, (almost) entire1y anadro-
mous
4
4. Stem (sub)erect, short, dictyostelic; scales clathrate; fertile
segments distinctly contracted (West Indies) 30. Anopteris
- Stem creeping, solenostelic; scales non-clathrate; fertile
segments scarcely contracted (Madagascar, Mascarenes)
29. Ochropteris
29. Ochropteris J. Smith
28. Adiantum L.
Fig. 127A-D
Adiantum L, Spec. Plant. 2: 1094 (1753); Gen. Plant. ed.
5: 485 (1754); Keyserling (1875); Kuhn (1881); Ching
(1957); Scamman (1960); Nayar (1961); R. Tryon (1964);
Sledge (1973); Kramer (1978); Palacios-Rios and Riba
(1985); Tryon and Tryon (1982).
Hewardia J. Smith (1841 a); Ching (1957).
A large and diverse genus of about 150 species; its
characters are reviewed under the subfamily. Tryon
and Tryon (1982) presented a tentative organization of
the species into eight groups.
Subfamily Pteridoideae
Terrestrial or epilithic, rather small to very large ferns
with creeping to erect, solenostelic or dictyostelic (in
large species sometimes polycyclic) stem bearing basifixed, (sub)entire, mostly non-clathrate scales. Petiole
adaxially sulcate, with 1, 2, or 2 large and a number of
smaller, dorsal bundles. Lamina simply pinnate to
several times pinnate, rarely simple; veins free or reticulate without free included veinlets. Stomata predominantly polocytic, anomocytic ones usually present besides. Sori on a submarginal commissure joining 2 to
very many vein ends, often on somewhat contracted
laminal part..; or sori none, the sporangia "acrostichoid". Sporangia intermingled with pluricellular trichomes with usually enlarged, ± capitate-glandular
end cell, these trichomes rarely wanting. Leaves nonhairy, or if hairy the hairs not resembling the trichomes
mixed with the sporangia. Indusium consisting of the
reflex ed, modified leaf margin or wanting. Spores
trilete, sometimes with an equatorial flange; n=29 or
mUltiples in most representatives but n=30 in Acrostichum and 27 in one species of Pleris; the number is
not known for Ochropteris.
KEy TO THE GENERA.
1. Sporangia assembled in short to very long submarginal sori
3
- Sporangia not assembled in sori but borne on the veins as
well as on the leaf tissue; veins reticulate; leaves simply
2
ScariOllS, ± reflexed (circumCaribbean)
32. NeurocaDis
- Leaf margin undifferentiated (widespread) 33. Acrostkhum
pinnate
2. Leaf margin differentiated,
3. Lamina (venation) quite catadromous, or the lamina anadromous at base and becoming catadromous upward; rare-
ly lamina simple (widespread)
31. Ptetis
Ochropteris J. Smith in Hooker, J. Bot. 4: 158 (1841).
Medium-sized to rather large ferns with creeping,
solenostelic, scaly stem; scales castaneous, lanceolate,
non-clathrate. Leaves rather closely distichous; petiole
naked except at base, stramineous with dark base, exceeding the lamina, adaxially sulcate, all leaf axes similarly grooved, with continuous grooves; vascular bundle one, omega-shaped in cross-section. Lamina deltoid, chartaceous to subcoriaceous, glabrous, quite
anadromous, tripinnate + pinnatifid or to 5-pinnate at
base; larger divisions stalked; upper divisions gradually reduced and confluent. Ultimate divisions ± pronouncedly asymmetrically rhombic-cuneate, pinnatifid
to subentire, ascending, with narrow base, the larger
with flexuous costa; veins simple or forked, free, ending near the margin, behind its teeth. Stomata polocytic, some anomocytic. Sori on both lateral margins,
only on the anterior margin, or on (part of) the truncate apex of a segment, depending on the size of the
segment, suborbicular or elliptic, on a commissure
joining two to several vein ends; indusium consisting
of a reflexed, modified marginal lobe, firm, dark at
base, pale towards the margin, entire to sinuate. Soral
trichomes numerous, pluricellular, filiform. Spores tetrahedral, with prominent equatorial flange and rough
surface.
A single species, O. pal/ens (Sw.) J. Smith, in the
Mascarenes and locally in Madagascar; terrestrial in
lowland thickets and at forest edge, in climatically
moist places. Additional species described by Fee and
Tardieu-Blot do not merit recognition.
30. Anopteris (Prantl) Diels
Anopteris (Prantl) Diels in Engler & Prantl, Nat. Pflanzenfam. 1,4: 288 (1899); Morton (1957), based on: Cryptogramma sect. ('1) Anopteris Prantl, Engl. Bot. Jahrb. 3: 414
(1882).
Rather small epilithic ferns; stem short, (sub)erect, dictyostelic, bearing stiff, broad, dark brown, pale-margined, erose, clathrate scales. Leaves close; petiole
about equaling the lamina, with one vascular bundle.
Lamina deltoid, bipinnate to quadripinnate + (bi)pinnatifid or even 5-pinnate, (firmly) herbaceous, anadromous in dissection and venation or not quite consistently so, naked; leaf axes pale, thinly scaly to naked,
adaxially grooved, all grooves continuous. Larger divi-
250
Pteridaceae . Pteridoideae
sions stalked; stalks of smaller divisions ± winged by
decurrent laminal parts. Upper divisions gradually reduced, but a free terminal segment usually present.
Segments lanceolate-cuneate, very variable in size and
shape, about linear in highly dissected forms, contracted in fertile leaves; veins pinnate, the branches simple,
ending behind the margin in clavate tips, or very narrow segments uninerval; sterile margin lobulate-dentate at least in the distal half, such teeth often also present on the distal sterile parts of fertile segments.
Fertile segments somewhat contracted, soriferous on
parts of their lateral margins; sori borne on a submarginal commissure connecting a few·to most vein ends;
indusium thin, pale, the edges of two opposite ones often meeting over the costa. Soral trichomes numerous,
filiform, pluricellular, gland-tipped. Spores tetrahedral-globose, with prominent angles, without equatorial flange, coarsely tuberculate or rugose on distal,
much less so on proximal face.
A single species (Morton 1957, 1962), A. hexagona
(L.) C. Chr., on limestone rocks in the Greater Antilles
and Bermuda. The non-catadromous leaves separate
this readily from Pteris, the erect, dictyostelic stem
from Ochropteris.
31. Pteris L.
Figs. l06D, 128A-D
Pteris L., Spec. Plant. 2: 1073 (1753); Walker (1958, 1962);
Scamman (1961); Shieh (1966); Keating (1968); Schelpe
(1969); Saiki (1983).
Pycnodoria C. Presl (1849).
Schizostege Hillebrand (1888).
Lathyropteris Christ (1907).
Hemipteris Rosenstock (1908).
Idiopteris Walker (1957).
Schizostegopsis Copel. (1958); Stone (1968), nom. invalid.
Copeiandiopteris Stone (1973, 1976).
Terrestrial or epilithic, small to very large ferns; stem
short-creeping to erect, slender to massive, solenostelic
or dictyostelic (sometimes polycyclic), often strongly
sclerotic, bearing basifixed, elongate to very narrow,
entire or less often dentate, mostly brownish to blackish, glabrous, non-clathrate scales, their margins sometimes pale. Leaves rather close to aggregated; petioles
usually well-developed, with one, in cross-section Uor V- or about omega-shaped, less often 2 C-shaped
vascular bundle(s), or in large species sometimes with
two larger adaxial and a semi-circle of smaller abaxial
ones; petiole adaxially grooved, additional ventrolateral grooves sometimes present in large species; colour
pale to dark, not rarely dark-polished; surface often
naked, or ± persistently scaly, less often hairy. Lamina very rarely simple, usually once to several times
(occasionally to 5 x) pinnate, the ultimate divisions
often pectinately lobed to pinnatifid; lamina occasionally palmate or pedate. Basal pinnae very often basitonically produced, not rarely with a single, large, bas-
al posterior segment qonform to the one bearing it
("forked basal pinnae"); at least the basal pinnae
anadromous, the rest of the lamina gradually to
abruptly catadromous (or subisodromous), or in a few
species (subgen. Pteris) entirely catadromous. Leaf
axes not rarely winged, especially distally, below pinna
insertions, adaxially grooved, the grooves with few exceptions continuous on axes of different order. Short
spines sometimes present on the axes, especially abaxially, or only on their bases ("axils"). Larger divisions
sessile or short- (rarely long-) stalked, very often of an
elongate-oblong to lanceolate-Iinear shape, usually
symmetric; apex often abruptly acuminate-caudate
above the pinnatifid part (if any); upper divisions
lobed-confluent, or a (sub)conform terminal division
present. Surface glabrous or less often variously hairy,
rarely scaly when mature; texture thin-herbaceous to
coriaceous. Stomata polocytic and anomocytic.
(Pen)ultimate disivions costate, with percurrent costal
costule, the latter adaxially usually grooved, the ridges
bordering the groove not rarely bearing a short to
long, ascending spine at the insertion of a costule, or
sometimes on the costules (or even the veins). Veins
simple or usually forked, catadromously arranged,
reaching the margin or not, free, or forming costal!
costular arches, or more freely to entirely reticulate;
free included veinlets wanting. "False veinlets" rarely
present between the true veins (see below). Margin entire to sharply dentate or serrate. Fertile divisions subconform to strongly contracted; fertile lamina nearly
always with the same architecture as the sterile. Sori
on a short to long submarginal commissure uniting a
few to many veins, or even all veins of a segment, often absent from its apex and/or the sinus; indusium
always present and evident, elongate to linear or rarely
short and lobe-like, formed by the modified, reflexed
leaf margin, occasionally slightly intramarginal. Soral
trichomes filiform, several to many-celled, usually evident, sometimes few, apparently wanting in a few species. Spores tetrahedral or globose, usually with an, often prominent, equatorial flange; surfaces with low
tubercles ± fused into ridges; or, in subgen. Pteris,
reticulate; spherical deposit present.
MORPHOLOGY AND ANATOMY. The venation pattern is
variable, closely related species not rarely differing in
this respect. It is therefore unsuited as a basis of intrageneric classification. The epidermal venuloid idioblasts ("false veinlets") present in Pt. grevilleana Wall.
ex Ag. and some other species were described and discussed by Wagner (1978), who regarded them as evidence for alliance with Adiantum and Vittariaceae.
The continuous, mostly very long receptacles are ±
typical of the genus but not consistently present, some
species having often [Pt. ("Schizostege'') lydgatei (Hillebr.) Christ) or always [Pt. ("Hemip/eris'') werneri (Ro-
Pteris
senst.) Holttuml short interrupted sori; nor is the sorus
form of typical Pteris confined to this genus but occurs
also in, e.g., Ajropteris, Onyehium, Doryopteris, Teetaria
("Amphiblestra"), etc.
CHARACTERS OF RARE OCCURRENCE. Simple (sterile
and/or fertile) leaves in Pt.simplex Holttum and
Pt. deltodon Baker; a pectinately (sub)pinnate, otherwise undivided lamina in sect. Syrrhaca and in Pt. subindivisa Clarke and Pt. redueta Baker; very asymmetrical, only posteriorly pectinately pinnatifid pinnae in
Pt. semipinnata L. and Pt. dispar Kunze, only anteriorly pinnatifid in Pt.jormosana Baker; abaxially spiny
axes in Pt. hamufosa Christ, etc.; sori partly overlain
by a laminal flange arising opposite the indusium in
Pt. warburgii Christ; a scandent rachis in Pt.fathyropteris C. Chr.; proliferous buds on the rachis in Pt. profifera Hieron.
The pale marks on the lamina of a few species (e.g.,
Pt. argyraea Moore), mostly on the basal parts of the
segments, flanking the costa, make them attractive as
ornamentals but have little taxonomic significance.
251
Fig, 128A-G. Pteridaceae. A, B Pteris gigantea. A Fertile
pinnule from abaxial side (x 1). B Lateral pinna (x 0,5).
C.D Pterisfauriei. C Costa and fertile segments from abaxial
side (x 2,5). D Costa with spines and segment bases from
adaxial side (x 1,4). E-G Pleurosoriopsis makinoi. E Habit
(x 1,8). F Detail of lamina (x 7,3). G Hairs on margin of
lamina (x 180) (A,B from Vareschi 1969; C,D from Ching
1935; E-G from Kurata and Nakaike 1979)
KARYOLOGY AND HYBRIDIZATION. With a single exception, all chromosome numbers found in the genus are
based on X= 29; only Pt. ("Jdiopteris") hookeriana Ag.
has n=27. Many species in subgen. Litobrochia are interfertile, and the great part played by hybridization in
the genus was elegantly demonstrated by Walker
(1958, 1962); hybrids not rarely become stabilized as
species through the occurrence of apogamy. These are
then difficult to recognize as hybrids, and the difficulty met with in trying to arrive at a satisfactory spe-
252
Pteridaceae . Pteridoideae
cies concept and species distinction in the genus, especially in the group of species centred around Pt.
quadriaurita Retz., is at least partly due to this phenomenon. In a few cases even hybrid swarms are
known, of which few instances have been recorded in
pteridophytes. Polyploidy also plays a part (Walker
1962), and races with different ploidy may occupy different areas (Nakato 1981).
Another curious aspect of hybridization in the genus
is the fact that some hybrids instead of showing intermediate characters fluctuate between the characters of
one parent species and the other; e.g., in one pinna
there may be alternating entire and pinnatifid zones, if
the parents differ in this feature (Walker 1958; Wagner
1962). Such plants are of course readily recognized as
hybrids.
SUBDIVISION. Nearly all subdivisions proposed so far
are based on easily observed characters of little taxonomic weight, like dissection pattern, venation, etc. A
natural subdivision covering the entire genus has so
far not been proposed. A hopeful start, but involving
only E. Asiatic species and still partly artificial, was
made by Shieh (1966), who as a whole employed more
natural characters including scales.
Many species are so large that most (if not all) extant herbarium specimens are incomplete, lacking
petiole bases, stems with their scales, etc. This stands
in the way of correct identification and even of a satisfactory classification.
TWo subgenera can be distinguished as follows:
1. Lamina entirely catadromous, often basally ± reduced;
grooves on costae not continuous with those of the primary
rachis, or costae not grooved; pinnae always simple, entire,
and free-veined; spores tetrahedral, trilete, rarely monolete,
ellipsoidal, distal face coarsely reticulate, rugose elements
forming an incomplete equatorial flange closer to the aperture than to the equator
subgenus Pteris
- Lamina anadromous at least in the basal pinnae, truncate
at the base, or very rarely the lamina simple; grooves on
axes of various order continuous with one another; pri-
mary pinnae very often "forked" (at least the basal ones),
often pinnate or pinnatifid; veins free or reticulate; spores
trilete, tetrahedral or spheroidal, usually tuberculate, more
prominently so on the distal face; flange closer to the equator than to the aperture, or wanting, both sides uniformly
reticulate or tuberculate
subgenus Litobrochia
The group of three species from the Philippines with
adaxially non-grooved costae and gradually confluent
upper pinnae, segregated as section Syrrhacra M.G.
Price by Price (1975), may also merit subgeneric rank;
this requires further study.
The great majority of species belong to subgenus
Litobrochia. A natural subdivision of this into sections,
etc. is still to be given.
An estimated 250 species, partly depending on what
species are recognized in the complex of species relat-
ed to Pt. quadriaurita Retz. The genus is worldwide in
tropical, warm-temperate, and south-temperate areas
but lacking in very dry regions and poorly represented
in temporarily dry regions; of local occurrence in
north-temperate areas, but well represented in East
Asia. The species are rather evenly distributed over the
continents, even continental tropical Africa having a
fair number. Some species are local endemics of small
islands like St. Helena and La Reunion.
Pteris species grow on the ground in ± shaded
places, from sea level to high in the mountains, less often in open places, on calcareous and other rocks, etc.
A number have a certain importance as garden ornamentals and some have in this way become established
far outside their natural ranges, like Pt. tripartita Sw.,
Pt. vittata L., and Pt. multifida Poiret.
32. Neuroca/lis Fee
Fig. 129
Neurocallis Fee, 2' Mem.: 19, 89 (1845).
Large terrestrial ferns with stout, ascending, dictyoste1ic, fuscous-scaly stem; leaves close; petiole well-developed, dark and scaly at base, paler and glabrescent
above, adaxially (and laterally?) sulcate, with a single,
in transection omega-shaped vascular bundle. Lamina
simply pinnate, anadromous at base, catadromous
above, with sessile (often subopposite) lateral pinnae
and a conform terminal one, dimorphic; sterile pinnae
lanceolate, acuminate, (sub)glabrous, entire, with percurrent costa; secondary veins none; tertiary venation
reticulate, with roughly hexagonal meshes decreasing
towards the margin; free included veinlets none. Stomata polocytic. Fertile leaves taller, with linear pinnae;
sporangia not assembled in sori, covering the pinnae
beneath except for a narrow zone along the costa, the
pinna margin narrowly reflexed and modified; soral
trichomes numerous, mostly capitate. Spores tetrahedral-globose, without equatorial flange, shallowly
rugulose, especially distally.
A single species, N. praestantissima Fee, in Costa
Rica, northern Venezuela, and the Lesser Antilles; rare
in most parts of its area. Terrestrial in moist forest at
lower and middle elevation.
The genus is closely related to Pteris, differing mainly in the disposition of the sporangia and spore sculpture.
33. Acrostichum L.
Acrostichum L., Spec. Plant. 2: 1067 (1753); Schumann (1915);
Troll (1934); Adams and Tomlinson (1979); only p.p. min.,
or not at all, of most authors between 1753 and c. 1900.
Terrestrial ferns of moist to swampy places, of medium
to large size. Stem short, creeping to erect, stout,
dictyostelic, with a lesser system of medullary bundles
Neurocallis . Acrostichum . P1eurosoriopsis
(diplostelic), scaly; scales often large, up to a few em
long, multistratose, thin at the entire edge; roots fleshy.
Leaves approximate; petiole well-developed but shorter than the lamina, glabrescent, stramineous to dark
brown, adaxially sulcate, additional ventro lateral
grooves may be present; sometimes spine-like, reduced pinnae present on petiole; petiole base with two
larger, adaxial, in section C-shaped bundles and a ring
of smaller abaxial bundles, the pattern becoming gradually more complicated upward. Lamina simply pinnate (simple in juvenile plants), glabrous when mature,
with numerous close to remote, often ascending pinnae, at least the larger short-stalked; rachis adaxially
grooved; lower pinnae not reduced (strongly reduced
and spine-like in A. aureum), upper pinnae somewhat
reduced, a free conform terminal pinna present Pinnae tongue-shaped to elongate-oblong, chartaceous to
thick-coriaceous, entire, obtuse to acuminate, with
percurrent costa, this adaxially sulcate, the groove evanescing at the pinna base and thus not quite continuous with that on the rachis. Basal pinnae anadromous
(i.e., the leaf tissue extending farther down on the anterior side), the upper gradually becoming catadromous. Veins immersed but usually evident; secondary
veins none; tertiary veins fOrming a dense reticulum of
ascending, narrow, ± hexagonal areoles, only the costal areoles elongate and about parallel to the costa;
free included veinlets none. Stomata polocytic and
anomocytic. Fertile pinnae borne in the apical part of
the lamina or on most or all of the fertile lamina,
somewhat (not strongly) contracted, otherwise similar
to the sterile, abaxially everywhere, except on costa
and very near the margin, evenly covered by the sporangia; margin not modified nor reflexed. Sporangia
mixed with numerous trichomes, these with swollen,
entire to lobed apex. Spores tetrahedral-globose, without equatorial flange, coarsely diffuse-papillate or tuberculate, with rods or sparse strands, these denser distally; n=30.
Three species in brackish to saline swamps and
mangrove vegetation, often also occurring farther inland in fresh-water swamps. A. aureum L. is pantropic
to warm-temperate, A. speciosum Willd. paleotropic
from SE Asia to tropical Australia, and A. danaeifolium Langsd. & Fisch. neotropical. Possible hybrids
were reported by Garcia de L6pez (1978).
Morphologically the genus is very close to Pteris,
but the distribution of the sporangia, the soral trichomes, and the chromosome number exclude it readily. Formerly it was much more broadly circumscribed,
including (nearly) all ferns with "acrostichoid" sporangia, and was one of the least natural fem genera of the
18th and 19th centuries.
253
Fig. 129. Pteridaceae.Neurocallis praestantissima, venation
(Original A. F. and R M. Tryon)
Genus incertae sedis:
34. P1eurosoriopsis Fomin
Fig. 128 E-G
Pleurosoriopsis Fomin, Bull. Jard. Bot. KiefT 11: 8 (1930);
Kurita and Ikebe (1977).
Small epilithic ferns; stem long-creeping, dictyostelic
with only two meristeles, bearing short, 2-celled hairs
and longer acicular, ferrugineous hairs often inserted
in pairs or small groups on an epidermal outgrowth.
Leaves remote; petiole well-developed, non-articulate,
hairy, with a single, terete vascular bundle. Lamina
herbaceous, narrowly oblong, pinnate + pinnatifid, or
bipinnate at base, anadromous, bearing short, articulate hairs like the stem, especially on the rachis and the
margin. Pinnae oblong, obtuse, with few entire or
lobed segments; veins free, forked, with clavate ends
far behind the margin. Stomata polocy1ic. Sporangia
following the veins in their lower and middle parts; indusium none. Spores monolete, with little surface
sculpture; 2n= 144.
A single species, P. makinoi (Maxim.) Fornin, in the
mountains of eastern Asia, from the Ussuri Province
to Korea and Japan.
The genus is of disputed and uncertain affinity;
placed in Pteridaceae by Copeland (1947), Tagawa
(1959) and Ohwi (1965); in Aspleniaceae by LOve et al.
(1977); and in Grammitidaceae by Tryon and Tryon
(1982). None of these placements seems convincing
254
Pteridaceae . Pteridoideae
(the last-named perhaps the most); in every one of
these families Pieurosoriopsis would form a discordant
element. It is certainly significant that Kurita and
Ikebe (1977) who studied the genus rather closely
failed to arrive at a conclusion about its taxonomic
position and suggested to place it in a family by itself; this was subsequently formally described by
Ching: Pleurosoriopsidaceae Ching, Acta Phytotax.
Sin. 16: 17 (1978). The genus is, however, too little distinctive to justify the erection of such a family, and
"incertae sedis" expresses better that a satisfactory
conclusion about its affinity is yet to be reached. It is
here appended to Pteridaceae only because it will key
out to this family in the general key, without any implication that its natural place is there.
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Salviniaceae
J. J. SCHNELLER
Salviniaceae Dumortier, Anal. Fam. PI. 67 (1829).
Free-noating aquatic plants with horizontal protostelic
stems. Roots lacking. Leaves in grottpS of three along
the length of the stem, each group consisting of two
dorsal noating, green, simple, entire leaves and one
ventral, strongly branched, submerged leaf. Floating
leaves nearly sessile or short-stalked, usually ovate to
cordate, with costa and anastomosing lateral veins
without free included veinlets. Upper surface with special water-repellent hairs. Submerged leaf usually with
a well-defined petiole. Shoots, submerged leaves, and
lower parts of noating leaves bearing uniseriate, multicellular, apiculate hairs. Spore-producing organs borne
on the submerged leaves, consisting of sori surrounded
by a globose indusium ("sporocarp"). One or a few
basal sporocarps containing up to about 40 megasporangia, the other (more numerous) sporocarps containing numerous microsporangia. Sporangia without dehiscence mechanism. Microsporangia containing either 64 or 32 trilete spores, these all together enclosed
in a hardened, alveolar mass: the massula. Megasporangium originally containing 32 spores, only one of
which develops into a mature trilete spore. Spores
spheroidal; megaspores shallowly rugose to nearly
plain and perforate; microspores shallowly rugose to
rugulose. Microprothallium reduced, only a few cells
long, with two reduced antheridia. Megaprothallium
free-floating, about 0.5-1 mm long, consisting of more
or less heart-shaped tissue of a few cell layers, connected with the megasporangium, normally with only
a few archegonia.
ANATOMY AND MORPHOLOGY. Important contributions
have been made by Pringsheim (1863), Zawidski
(1912), and Croxdale (1978, 1979, 1981). As already
mentioned by Pringsheim (1863), the leaves OCCUr in
phyllotactic units of six, within each unit the first three
leaves are initiated in one direction, the next three in
the other. Croxdale (1978) showed that the noating
leaf nearer to the submerged leaf is produced prior to
the initiation of the other noating leaf and therefore
she preferred to speak of groups rather than of whorls.
There is always a resting bud (or a grown-out lateral
branch) between the floating leaf further from the submerged leaf and the submerged leaf. Therefore one
finds a bud (or the lateral branch) in one group
("whorl") on the right and in the next on the left side.
The development of the leaves, especially the noating
leaves, is unique amongst the ferns. Again Croxdale
(1979) showed that the upper side of the noating leaf
(which is folded in bud) is anatomically the abaxial
surface. The hairs on the upper side (abaxial) furnish
important differential characters. In one group of species they are borne on papillae (normally four hairs together), in another they are borne directly on the surface of the leaf; these hairs may be arranged irregularly or in rows. The hairs on the papillae are free or
connected at the tip ("egg-beater type"). It is not exactly known if the highly branched submerged leaf may
have some root-like functions or if its role is merely
that of a stabilizing organ. The fertile parts of the submerged leaves consist of a chain- or cyme-like organ
on which many pedunculate or sessile sporocarps are
borne. The initials of the mega- and microsporangia
appear early at the apex of the young sporocarp. They
develop on a stout columnar receptacle on the base of
the sporocarp. In the megasporangia there are up to
about 40 sporangia arranged in small groups on
branched stalks or solitary on short stalks. In the microsporocarp the sporangia are numerous and borne
on slender, branched stalks. The sporangia develop in
the same mode as in other leptosporangiate ferns, except that no annulus is formed. Mega- and microspores (the latter assembled in the massulae) remain in
the sporangium. Sporangia containing spores are distributed as entities and become free when the sporocarps decay. The wall surrounding the sori is interpreted as a two-layered indusium (Zawidski 1912). It
forms the approximately globose sporocarp.
GAMETOPHYTE. Important contributions to the gametophyte development and structure were published by
Pringsheim (1863), Belajeff (1898), and Yasui (1911).
All published data are based on observations made on
S. natans. The micros pores germinate inside the spo-
Salvinia
257
o
rangium and the small prothalli penetrate the sporangium wall (in predetermined regions?). The microprothallia remain fixed to the sporangium. Belajeff (1898)
and Yasui (1911) could clearly show that the much reduced prothallium forms two reduced antheridia. This
seems to be true also for S. sprucei (Schneller unpubl.).
Pringsheim (1863) and others showed that on older
megaprothallia two wing-like appendages are formed
which are thought to be stabilizing organs. Such wings
could not be found in the two tropical species S. sprucei and S. bi/oba (Schneller unpubl.). According to
Schneller (1976), the megaprothallium floats on the
water surface with the archegOnia directed downwards
and not upwards as stated in all textbooks (based on
Pringsheim's publication).
BLASTOGENY. The sporophyte generation initially
forms a more or less long-stalked, free-floating leaflike structure with stomata on the upper surface.
Below this "first leaf' the shoot apex develops which
then forms regular floating leaves and submerged
leaves. In the tropical S. biloba this "first leaf' differentiates two lobe-like appendages and then the first
regular leaves appear (Schneller unpubl.). The "first
leaf' and the appendages contain small vascular bundles. In S. sprucei a small protuberance on the upper
side of the "first leaf' may be observed, but no clearly
distinguishable appendages. In all species observed so
far the first two floating leaves bearing water-repellent
hairs on the upper side are not inserted in an exactly
opposite position and no submerged leaf. is formed.
The following group of leaves then consists of two
floating leaves and one submerged leaf.
Fig. 130 A- o. Salviniaceae. A, B Salvinia auricula/a. A Habit (x 0,6). B Upper leaf surface with "egg-bea~er" hairs
( x 20). C Salvinia na/ans, habit (x 0,8). 0 Salvmla has/ala,
upper leaf surface (x 20) (A,B,O from Schelpe 1970; C from
Wettstein 1933)
ECOLOGY AND DISTRIBUTION. Most species occur in
nutrient-rich water in tropical and subtropical regions.
They show strong vegetative propagation and a normally very high growth rate. Some species have become tropical weeds, especially on man-made reservoirs and in irrigation systems. One, S. molesta Mitchell, has become pantropic (through human agency).
There are two main distribution centres of the genus,
one in Africa, the other in South and Central America.
Two Old World species are extratropical in the northern hemisphere. Two other species occur in the palaearctic region.
AFFINITY. According to some authors there are affinities to Azolla, but as so many characters are quite different this affinity does not seem to be very close. Inclusion in one order, Salviniales, seems indicated. The
phylogenetic derivation of Salvinia remains unknown.
Fossil remains were found in the Cretaceous (Hall
1974). Some authors suppose a derivation from the
Hymenophyllaceae (see Takhtajan 1953) because of
the cup-shaped indusia. However, the other morphological and anatomical features are very different ~rom
the Hymenophyllaceae; there is, in fact, not a slllgie
further point of resemblance.
KARYOLOGY AND HYBRIDIZATION. The chromosome
numbers of about one-fourth of the species are known.
258
Filicatae . Schizaeaceae
The base number is 9. Polyploids up to heptaploids are
known (Schneller 1981). Some hybrids occur mainly in
South America; S. molesta. the notorious weed, is hybridogenous, too.
Schizaeaceae
A single genus:
Schizaeaceae Kaulf., D. Wesen d. Farrnkr.: 119 (1827).
Anemiaceae Link (1833).
Lygodiaceae C. Presl (1845).
Mohriaceae (c. Presl) Reed (1948).
Sal.inia Seguier
Fig. 130
Salvinia Seguier, PI. Veron. 3: 52 (1754); Reed (1954); de la
Sota (1962,1963,1964, 1976 a, b).
Characters as for family.
Ten species, widespread in the Old World except in SE
Asia-Australasia; and in the New World north to the
southern United States.
Selected Bibliography
Belajeff, W. 1898. Vber die mfinnlichen Prothallien der Wasserfame. Bot. Z. 56: 141-194.
Bonnet, A. L. M. 1955. Contribution it I'etude des Hydropteridees: Recherches sur Salvinia auricuiata Aubl. Ann. Sci.
Nat. Bot. 16: 529-599.
Croxdale, J. G. 1978. Salvinia leaves. II. Origin and early differentiation of floating and submerged leaves. Can. J. Bot.
56: 1982-1991.
Croxdale, J. G. 1979. Salvinia leaves. II. Morphogenesis of
the floating leaves. Can. J. Bot. 57: 1951-1959.
Croxdale, J. G. 1981. Salvinia leaves III. Morphogenesis of
the submerged leaf. Can. J. Bot. 59: 2065-2072.
Hall, J. W. 1974. Cretaceous Salviniaceae. Ann. Missouri Bot.
Gard. 61: 354-367.
Herzog, R. 1934. Anatomische und experimentell-morphologische Untersuchungen ober die Gattung Salvinia. Planta
22: 490-514.
Pringsheim, N. 1863. Zur Morphologie von Salvinia natans.
lahrb. Wiss. Bot 3: 464-472.
Reed, C. F. 1954. Index Marsileata et Salviniata. Bol. Soc.
Brat. II, 28: 5-61.
Reed, C. F. 1965. Index Marsileata et Salviniata. Suppl. Bol.
Soc. Brat. II. 39: 259-302.
Schneller, 1.1. 1976. The position of the megaprathallus of
Salvinia natans. Fern Gaz. 11: 217-219.
Schneller, J. J. 1981. Chromosome number and spores of Salvinia auriculata Aublet s. str. Aquat. Bot. 10: 81-84.
de la Sota, E. R. 1962. Contribucion al conocimento de las
"Salviniaceae" neotropicales I-III. Darwiniana 12:
465-520.
de la Sola, E. R. 1963. Contribucion al conocimento de las
"Salviniaceae" neotropicales IV. Darwiniana 12: 612-623.
de la Sota, E. R. 1964. Contribucion al conocimento de las
"Salviniaceae" neotropicales V. Darwiniana 12: 529-536.
de la Sota, E. R. 1976a. Sinopsis de las especies argentinas
del genera Salvinia Adanson (Salviniaceae, Pteridophyta).
Bol. Soc. Argent. Bot. 17: 47-50.
de la Sota, E. R. 1976b. Indice sistematico y bibliogrMico de
los taxa viventes del genera "Salvinia" Adans. (Salviniaceae, Pteridophyta). Obr. Centeno Mus. La Plata 3: 229-235.
Takhtajan, A. 1953. Phylogenetic principles of the system of
higher plants. Bot. Rev. 19: 1-45.
Yasui, K. 1911. On the life-history of Salvinia natans. Ann.
Bot. (London) 25: 469-483.
Zawidski, S. 1912. Beitrfige zur Entwicklungsgeschichte von
Salvinia narans. Bot. Centralbl. 28: 17-65.
K. U.KRAMER
Terrestrial or epilithic, small to large ferns with creeping or erect stem bearing multicellular hairs, rarely basifixed, non-clathrate scales. Them protostelic, solenostelic, or dictyostelic, radial or dorsiventral. Leaves
distichous or polystichous to spirally arranged; petiole
non-articulate, with a single solid, ± distinctly Cshaped, or 3-lobed xylem strand; axes adaxially convex, flattened, marginate, or with a single groove.
Lamina rarely simple, more often dichotomously or
pinnately compound, at least at the base usually distinctly anadromous, mostly firm, hairy or g1abrescent,
rarely scaly. Veins forked or pinnately branched, free
or less often reticulate without free included veinlets.
Sporangia usually assembled on strongly modified leaf
segments or entire leaves, rarely grouped in distinct sori, Or in one genus single on modified leaf lobes, the
head large, sessile or with a short, thick stalk, subglobose, flask-shaped, or asymmetrically flask-shaped,
with transverse, subapical, continuous annulus and
distinct stomium, dehiscing longitudinally (Fig. 131).
Indusium none, or when sporangia borne singly, each
with a special, small, indusium-like lobe inserted on
the sporangium-bearing vein and opening distally.
Spores trilete or less often monolete, with strongly
sculptured surface.
ANATOMY AND MORPHOLOGY. Anatomically the family
is quite diversified. The stem is protostelic in Lygodium. medullated-protostelic in Schizaea, solenostelic or
dictyostelic in Anemia, and dictyostelic in Mohria. The
leaf trace is unlobed in Lygodium; in the other genera
the xylem is C-shaped or trilobed. In some species the
leaves are amphistomatic or epistomatic, but most are
hypostomatic: the patterns are variable according to
genus.
Thorspecken and Hagemann (1983) argued on ontogenetic evidence that Anemia possesses sori, but the
applicability of the term is debatable.
KARYOLOGY AND HYBRIDIZATION. An acceptable base
number for the family has not been proposed; the genera are diverse in themselves and between one another.
Numbers of n= 38, 76, and 114, and 2n= C. 76, 114,
and 152 have been reported for Anemia by Mickel
(1962), who also found considerable numbers of hybrids and polyploids, up to 14 n (Mickel 1982). In Lygodium numbers of n=29, 30, 58, and 60 have been
Lygodium • Anemia. Mohria
259
B
A
c
D
found, with 2 n~56, 112, 116, and 120. In Schizaea
numbers of n~c. 72,77,94,96,103, and approximate
numbers up to c. 540 have been found (Jermy and
Walker 1985). MoMa is constant with n-76.
The high ploidy numbers make interpretation very
difficult. The large number of hybrids in Anemia has
made unravelling of the species quite difficult (Mickel
1982, etc.). Jermy and Walker (1985) described a hybrid in Lygodium.
GAMETOPHYTE. This is again very diverse. Some species of Schizaea have filamentous, partly green prothallia with special rhizoid-bearing cells, antheridia on
short branches, and archegonia on the filaments or on
archegoniophores; others (subgenus Actinostachys)
have subterranean, fleshy, tuber-like, non-green,
mycorrhizal prothallia. Cordate, thalloid prothallia occur in the other genera, hairy in Anemia and Mohria,
glabrous in Lygodium. The antheridia are rather large
and of a primitive, pluricellular type.
Fig. 131 A-D. Spores of Schizaeaceae and Theiypteridaceae
(all x 1000). A Anemia phyllitidis. B Mohria eaffrorum.
C Schizaea peetinata. D Thelypteris eheilanthoides. Pho!.
A. F. Tryon
SUBDIVISION. The family has been variously divided,
most excessively by Reed (1948) and others, who place
each genus in a distinct family; but although the genera are diverse and specialized in different ways, they
share certain important characters, sometimes in pairs
or 3's of genera; thus, according to the weight attached
to the characters, the family can be subdivided in various ways. Lygodium stands more apart in its asymmetric sporangia each covered by a kind of indusium,
in which it is unique, and in the twining rachis; biochemically it is also divergent (Lal and Bhambie 1981)
and is best placed by itself in a subfamily.
ECOLOGY AND DISTRIBUTION. The family is essentially
tropical and southern warm-temperate, but exceeds
260
Filicatae . Schizaeaceae
the Tropic of Cancer considerably in North America
and East Asia. Except for Lygodium, the genera often
inhabit mineral·poor and/or periodically dry or
swampy, open habitats. Lygodium is more shade-loving, preferring thickets and forest edges.
The family is of considerable geological age,
but perhaps not all fossil genera placed there in the
past really belong to it. Anemia is known to have formerly occurred in presently north-temperate areas.
Some taxonomists regard the Schizaeaceae as the
family from which the so-called Gymnogrammoid
ferns (Pteridaceae subfam. Cheilanthoideae) have
sprung. Mohria is not rarely confused with Cheilanthes
in the herbarium; but this may be due to ecologically
induced convergence. The spores are not entirely unlike those of certain "Gymnogrammoid" genera; on
the other hand, karyological data militate against closer affinity. Marsileaceae are often regarded as being
derived from Schizaeaceae, and here the case is much
stronger; but the relationship is remote at best.
AFFINITY.
KEy TO THE GENERA.
1. Stem and lamina scaly; sporangium-bearing parts not differentiated from the lamina
3. Mohria
- Stem hairy, lamina hairy or glabrous; sporangia nearly always borne on strongly modified laminas or parts of laminas
2
2. Rachis twining; primary divisions of lamina consisting of
(mostly short) axes with dormant apex and a pair of lateral
secondary pinnae; sporangia solitary, each protected by an
indusium-like flange
1. Lygodium
- Rachis not twining; lamina simple or, if divided, not with
donnant apices on the primary divisions; sporangia neither
solitary nor indusiate
3
3. Apices of segments or of the simple lamina bearing pinnately or pseudodigitately arranged sporangium-bearing,
strongly modified segments
4. Schizaea
- Sporangia borne on (nearly always strongly modified) basal segments or on entirely fertile, "skeletonized" leaves
2. Anemia
1. Lygodium Sw.
Fig. 132A,B
Lygodium Sw., Schrad. J. Bot. 1800': 106 (1802), nom. conserv.; Duek (1978).
Medium-sized or large terrestrial ferns with creeping,
rotostelic stem bearing stiff, articulate hairs. Petioles
2-ranked, with a single, non-lobed vascular trace; rachis twining, adaxially flattened or convex and narrowly winged on the sides, like its branches. Primary
rachis branches alternate, short to very short, their
apex dormant, hairy, each bearing a pair of branches,
these foliaceous or further (often catadromously) pinnately or palmately branched. Lamina hairy or glabrescent, often rather firm in texture. Ultimate divisions Iiguliform-Ianceolate or deltate, sometimes
palmatilobate or pinnatilobate, sometimes articulate at
base, their stalks adaxially grooved and laterally
winged up to the laminal part; costa convex above,
giving off forked or pinnately branched veins, the basal pair often opposite; veins free or less often reticulate without free included veinlets; margin of segments
entire or serrate. Stomata diacytic or occasionally
anomocytic. Fertile segments with hardly to strongly
reduced laminal parts but not "skeletonized", bearing
sporangia on most or all of their veins; these in their
apical parts serving as sporangiophores, each lateral
vein bearing a sporangium and an antrorse indusiumlike flange covering it abaxially. Sporangia shortstalked, very asymmmetrically ovoid, the annulus just
below the outward-pointing apex. Spores trilete, tetrahedral-globose, rarely monolete; surface bearing spheroid tubercles, or coarsely verrucate; distal face and
equatorial area coarsely reticulate.
C.4O species, pantropic and south-temperate, one in
the eastern USA; the greatest concentration of species
in Malesia and Central America. A natural subdivision
of the genus has not been given. Reticulate veins have
probably developed more than once and cannot be
used for infrageneric classification.
2. Anemia Sw.
Fig.131A
Anemia Sw., Syn. Fil.: 6; 155 (1806), nom. conserv.; Mickel
(1962, 1981, 1982); de la Sota and Mickel (1968); Duek
(1980) [often misspelled ':4neimia'J
Trochopteris Gardner (1842).
Coptophyllum Gardner (1842).
Hemianemia (Prantl) Reed (1947).
Terrestrial or epilithic, small to medium-sized ferns
with creeping to suberect, solenostelic or dictyostelic
stem bearing articulate, often yellow or reddish hairs.
Petioles distichous or polystichous, with one C-shaped
xylem trace, adaxially usually sulcate. Lamina once
pinnate or bipinnate + pinnatifid or rarely only pinnatilobate (very rarely simple), usually hairy with glandular and/or acicular hairs, often anadromous at base
and catadromous above. Veins forked or subpinnately
branched, free or less often reticulate without free included veinlets. Leaf apex pinnatifid or less often with
an odd terminal pinna. Costae not or only shallowly
grooved adaxially. Stomata peri-, desmo- and polocytic. Sporangia on two skeletonized basal segments
which are borne below or nearly at the level of the first
sterile pinnae and often held erect, rarely on scarely
modified basal pinnae ("Trochopteris'~; in a few species the entire sporangia-bearing lamina skeletonized
and fertile. Sporangia in two rows flanking the veins,
sometimes under very narrow wings of leaf tissue, sessile, slightly asymmetrically ovoid or pear-shaped;
annulus subapical, uniseriate, with stomium. Spores
trilete, globose to tetrahedral-globose, with prominent
angles (except in subgen. Anemiorrhiza); surface with
coarse, often echinulate or baculate ridges, or reticulate in species of sub gen. Anemiorrhiza.
Somewhat over 100 species, the great majority in the
Lygodium • Anemia· Mohria· Sehizaea
warmer and drier parts of America from Texas to
Argentina, a few in Africa (north to Nigeria and
Ethiopia) and in Madagascar, one in peninsular India.
Three subgenera recognized by Mickel (1962 and
later), largely based on characters of spores, stomata,
and stem morphology. The subdivision of the genus by
Reed (1948) has not met with approval. A species with
entire, tongue-shaped lamina was described by Mickel
(1984).
3. Mohria Sw.
Fig. 131 B
Mohria Sw., Syn. Fil.: 6, 159 (1806).
Terrestrial or epilithic, rather small ferns with shortcreeping dictyostelic stem bearing basally attached,
261
F'1g.132A-F. Schizaeaceae. A Lygodium mierophyl/um, fertile pinna (x 0,7). B Lygodium kerstenii, fertile pinna (x 0,7).
C-F Sehizaea peetinata. C Whole plant (x 0,5). D Fertile
apex of lamina (x 1,3). E Fertile segment from abaxial side
(x 6,5). F Sporangium (x 3,3) (Scbelpe 1970)
non-clathrate scales. Petioles close, with a single vascular trace. Lamina in appearance reminiscent of Cheilanthes, bearing scales and multicellular hairs, pinnate+pinnatifid to bipinnate+ pinnatifid, narrowly
oblong, anadromous basally, catadromous apically; all
axes adaxially grooved. Ultimate segments subflabellate, lobed-dentate, with subpinnate, free venation.
Stomata polocytic. Fertile segments not or hardly contracted; sporangia in small groups near the vein ends,
262
Filicatae . Schizaeaceae
initially ± covered by tbe reflexed, hardly modified
pinnule margin, lacking a true indusium, sessile, subglobose, with subapical annulus of ratber few cells.
Spores trilete, tetrahedral-globose; surface with coarse,
compact, tuberculate ridges with keel-like connections
at the angles.
Three species; one, M. caffrorum (L.) Desv., widespread in southern and eastern Africa, Madagascar,
and tbe Mascarenes; another, M.iepigera (Baker)
Baker, in central and SE tropical Africa and in Madagascar; the third, M. hirsuta Roux, in the Drakensberg, South Africa (Roux 1984).
Species of Mohria prefer rather dry, open places or
half-shade, grassland, rocky sites or open forests, at
lower to middle elevation.
4. Schizaea J. E. Smitb
Figs. 131 C, 132C-F, 133
Schizaea J. E. Smith, Mem. Acad. Thrin 5: 419 (1793), nom.
conserv.
Lophidium L. C. Rich. (1792).
Actinostachys Wallich (1829).
Microschizaea Reed (1947).
Small to medium-sized terrestrial ferns; stem creeping
or erect, with medullated protostele, bearing articulate
hairs. Petiole with a single, often trilobed xylem trace,
often winged above. Lamina simple or dichotomously
branched, sometimes consisting only of two wings of
tissue along the midrib, in other cases expanded, flabellate, lobed to deeply dichotomously incised; stomata hypocytic, in a few cases leaves amphistomatic.
Sporangia borne on terminal sporangiophores; modified segments with hardly developed laminal parts
which are pinnately or pseudodigitately (with very
short rachis) arranged. Sporangia inserted on the sides
of an abaxial ridge and often originally ± covered by
tbe reflexed edge of the laminal part, exposed at maturity, sessile, almost symmetric, ovoid or ellipsoid, witb
subapical, uniseriate annulus. Spores monolete, ellipsoidal to subglobose; surface various, rugose, papillate, foveolate, striate, ridged, or reticulate.
C.30 species in tropical and soutb-temperate areas,
in Africa only in the Soutb; one outlying species from
the n. e. USA to Newfoundland. Most species on substrates poor in minerals, at least one regularly on decaying wood.
The species with well-developed lamina have been
separated as Lophidium, but that is artificial. Species
with pseudodigitate sporangiophores are more often
segregated as Actinostachys, which also diverges by
having tuberous rather tban filamentous protballia,
and by otber, minor characters (see especially Bierhorst 1968). Treatment as a subgenus is preferred by
others, as is done here.
Selected Bibliography
Fig_ 133. Schizaeaceae. Schizaea digitata; Taiwan. Photo
Ch.-M.Kuo
Bierhorst, D. W. 1966. The fleshy cylindrical subterranean
gametophyte of Schizaea me/anesica. Amer. J. Bot. 53:
123-133.
Bierhorst, D. W. 1968. Observations on Schizaea and Actinostachys spp., including A. oligostachys sp. nov. Amer. J. Bot.
55: 87-108.
Bierhorst, D. W. 1971. Morphology and anatomy of new species of Schizaea and Actinostachys. Amer. J. Bot. 58:
634-648.
Britton, E. G. Taylor, A 1901. Life history of Schizaea pusilla.
Bull. Torrey Bot. Club 28: 1-19 [Contrib. New York Bot.
Gard.l1].
Brownlie, G. 1965. Chromosome numbers in some Pacific
pteridophyta. Pac. Sci. 19: 493-497.
Duele, J. J. 1978. A taxonomic revision of Lygodium (Filicinae) in America. Feddes Rep. 89: 411-423.
Duek, J.J. 1980. A taxonomical monograph of Anemiasubgenus Anemiorrhiza (Filicinae). Feddes Rep. 91: 69-87.
Holttum, R. E. 1959. Schizaeaceae. Flora Malesiana Ser. II. 1 :
37-61.
Lal, S. D., Bhambie, S 1981. Amino acid differentiation in
Schizaeaceae. Isr. J. Bot. 30: 11-12.
Lellinger, D. B. 1969. The botany of the Guayana Highland Part VIII. Schizaeaceae (Filicales). Mem. New York Bot.
Gard. 18 (2): 2-11.
Filicatae • Thelypteridaceae
Lovis, J. D., Roy, S. K. 1964. A chromosome count in Mohria.
Nature (London) 201: 1348.
Mickel, J. T. 1962. A monographic study of the fern genus
Anemia subgenus Coptophyllum. Iowa State J. Sci. 36:
349-482.
.
Mickel, J. T. 1967. The phylogenetic position of Anemia colimensis. Amer. J. Bot. 54: 432-437.
Mickel, J. T. 1981. Revision of Anemia subgenus Anemiorrhiza (Schizaeaceae). Brittonia 33: 413-429.
Mickel, J. T. 1982. The genus Anemia (Schizaeaceae) in Mexico. Brittoni. 34: 388-413.
Mickel, J. T. 1984. New tropical American ferns. Amer. Fern
J. 74: 111-119.
Prantl, K.. 1881. Untersuchungen zur Morphologie der
Gefal3kryptogamen II. Die Schizaeaceae, etc. Leipzig:
W. Engelmann.
Reed, C. F. 1948. The phylogeny and ontogeny orthe Pteropsida. I. Schizaeales. Bo!. Soc. Brot. 11.21: 71-197.
Richter, A. 1914. Phylogenetisch-taxonomische und physiologisch-anatomische Studien iiber Schizaea. Math. Naturwiss. Ber. Ungarn 30: 213-297.
Roux, J. P. 1984. Mohria hirsuta. a new fern species from the
Drakensberg. J. S. Mr. Bot. 50: 435-441.
Selling, O. 1944. Studies in the recent and fossil species of
Schizaea, with particular reference to their spore characters. Medded. Gliteborg Bot. TliIdg.16: 1-112.
Selling, O. 1946. Two new species of Schizaea and their affinities. Svensk Bot. Tidskr. 41: 431-450.
de la Sota, E. R., Mickel, J. T. 1968. Sinopsis de las especies
argentinas del genero ''Anemia'' Swartz (Schizaeaceae).
Rev. Mus. La Plata N. S. Bot 11: 133-152.
Thorspecken, A., Hagemann, W. 1983. Besitzt Anemia Sori?
Untersuchungen iiber die Entwicklung des Sporophylls
von Anemia phyllitidis (Schizaeaceae). Plant Syst. Evo!.
143: 133-150.
Thelypteridaceae
AR.SMITH
Thelypteridaceae Pichi-Senn., Webbia 24: 709 (1970).
Dryopteris auctt., p. p., e. g., Christensen (1907, 1913, 1920).
Terrestrial (rarely epiphytic) ferns with branched or
unbranched stems, these long- to short-creeping, suberect, or erect and trunk-like, bearing scales at apex,
these entire, usually pubescent along margin and
sometimes surface, basally attached. Vascular system a
radially symmetrical dictyostele. Petioles non-articulate, scaly at base, with two lateral pneumatic lines
along the length, in cross-section nearly always with
two hippocampus-shaped vascular strands at base,
these fusing into a V-shaped strand distally. Lamina
simple to pinnate to most often pinnate + pinnatifid,
catadromous, infrequently twice-pinnate Or more divided (but never with basitonically enlarged pinnae),
monomorphic or less often dimorphic. Pinnae usually
263
numerous, gradually reduced distally into a confluent,
lobed, apical segment, sometimes with a conform apical division. Indument of lamina and axes almost always of hairs, these variously unicellular or multicellular, acicular, hooked, furcate, or stellate (Iwatsuki
1962), sometimes also with sessile or stalked glands,
occasionally also with scales. Rachis adaxially
grooved, this groove not continuous with adaxial
grooves of costae. Aerophores often present at bases
of pinnae abaxially, these swollen or elongate. Veins
simple or less often forked, reaching the margin or not,
± cdistinctly catadromously pinnately arranged in ultimate segments; free, connivent below the sinus, or
united to form one to many regular areoles between
costa and pinna margin, then with (usually only) excurrent free veinlets. Sporangia nearly always in discrete sori, these orbicular to occasionally elongate,
dorsal (occasionally submarginal) on the veins, with a
short to long stalk three cells in cross-section, with a
well-developed vertical annulus, dehiscing by a horizontal slit, maturation mixed. Indusia present or absent, if present then reniform or infrequently athyrioid,
often very reduced and obscured in mature sori.
Spores 64 (infrequently 32) per sporangium, nearly always bilateral with a monolete scar (tetrahedral with
trilete scar in Trigonospora), non-chlorophyllous; perispore variously ornamented (reticulate, winged, spinulose); gametophytes surficial, cordate, often with marginal glands or hairs; x=27, 29-36.
ANATOMY AND MORPHOLOGY. There is no comprehensive survey of this large family. Holttum et al. (1970)
compared the types of Theiypteris, Cyclosorus, and
Arnpeiopteris, along with a Goniopteris, and Holttum
(1982 and references therein) provided much taxonomically useful information in numerous revisions.
Salient features of the family include the petiole with
two hippocampus-shaped vascular strands uniting distally and acicular hairs especially along costae and rachis above. Some groups have aerophores - swellings
or projections bearing numerous stomata - at the pinna bases. Such structures may protrude through a layer of mucilage on young leaves (Hennipman 1968;
Holttum 1973b; Smith 1980), and are presumably adaptations for air exchange. Various species produce
vegetative buds in the axils of pinnae. The stomata are
polocytic, with one or two subsidiary cells (van Cotthem 1970). Sporangia often bear distinctive glands or
hairs arising from the stalk or capsule; such structures
'have been used extensively in the delimitation of
genera (Holttum 1982, and references therein).
GAMETOPHYTE. The prothallium is cordate or elongatecordate, often with broad wings, symmetrical, chlorophyllous, relatively thin, fast-growing, lacking a strong
midrib, and often with hairs or stipitate glands (similar
to those found on the sporophyte) on the margin and
264
Filicatae • Thelypteridaceae
surfaces. Rhizoids are colourless or pale tan and
abundant on the lower half of the cushion. Antheridia
and archegonia are of the type common for higher
leptosporangiate ferns, with three-celled antheridia dehiscing by a cap cell and with long, slender-necked
archegonia directed away from the meristematic
notch. It is questionable whether differences in gametophyte morphology can be used to characterize any
of the genera (subgenera) of the family. See Atkinson
(1973,1975) for literature review.
SPORES. The spores are bilateral, with a monolete scar,
except in Trigonospora (Holttum 1971) and generally
range from 25-60 pm long x 16-35 pm wide. A variously ornamented perispore which may be winged,
rugose, echinate, papillate, or reticulate, surrounds the
exospore. The most important modern surveys include
those by Wood (1973) for the family, Smith (1980) for
Steiropteris. Grimes (1980) for Pseudocyclosorus. and
Tryon and Tryon (1982) for New World species. In
general, closely related species have similar spores. In
a few cases, spore morphology appears to characterize
particular groups, e. g., Trigonospora and Amauropelta.
Other groups, such as Stegnogramma (echinate spores)
and Steiropteris (winged spores) also seem relatively
homogeneous, but not unique in their spores. In most
groups, spore morphology is variable or insufficiently
studied. Within a related group, ploidy is often correlated with spore size (Smith 1971).
ECCOLOGY AND DISTRIBUTION. Thelypteridaceae are
one of the largest families of ferns, comprising nearly
1000 species mostly in tropical and subtropical regions; fewer than 2% are temperate. Both number and
diversity of species are greatest in Malesia, where
440 species in 22 groups occur (Holttum 1982), but significant numbers - perhaps 300 species in six groups are native to the Neotropics. Mrica and nearby islands
are relatively depauperate, with 55 species (Holttum
1974a). Related groups (genera of Holttum 1971) are
largely confined to either the Old or the New World;
only two (Cye/osorus subg. Cye/osoriopsis and subg.
Cyclosorus) are pantropic. Only two species are considered naturally pantropic. Most members of the family
occur at middle and lower elevations in or at edges of
rain forests; a few species extend to 4500 m in the
tropics. Still other groups are primarily temperate in
distribution (e. g., Phegopteris. Theiypteris subg. Parathelypteris). Most species are found in wet, primary,
undisturbed forests or in secondary forests. Others are
most common in shaded areas under cultivation, especially coffee and cacao plantations. Still others are
facultative or apparently obligate rheophytes [e. g.,
Theiypteris aspidioides (Willd.) R. Tryon, Th. sancta
(L.) Ching.] Some species [e. g., Cye/osorus dentatus
(Forssk.) Ching] are common and invasive "weeds" of
road banks, ditches, and greenhouses, and have great-
ly expanded their range under man's influence. Most
species are terrestrial, but some are epipetric; many
species of Cyclosorus subg. Goniopteris have an affinity
for calcareous rocks. A few species inhabit swamps,
e. g., Thelypteris palustris Schott, Th. serrata (Cav.) Alston = Cye/osorus palustris (Raddi) ined. The earliest
fossil that may belong to this family is the
Jurassic Aspidistes (Holttum 1971), but its affinities,
even if thelypteroid, are very uncertain because of its
2-3-pinnate leaves and trilete spores. Both conditions
are unusual in extant species and occur in different
genera.
AFFINITY. This family apparently has no close living
relatives, and its affinities are not known with confidence. Until recently, species were included in Dryopteris. First Ching (1940) and later Holttum (1971,
1973 a) have effectively refuted that relationship. Following an intensive study of the family, particularly
Old World elements, Holttum (1971, 1973 a, 1982) concluded that Thelypteridaceae are most closely related
to, and probably had a common origin with. Cyathea
s.1. Shared characters include a radially symmetric
stem, erect in Cyathea and (according to Holttum)
primitively erect in Thelypteridaceae; presence of similar acicular hairs, especially on costae above; presence of aeration tissue at the base of pinnae and along
petiole and rachis; similar leaf form (catadromous and
with progressively reduced lower pinnae) and segment
venation (symmetrically pinnate in each lobe). The
presence of trilete spores in the thelypterid Trigonospora (all other groups have monolete spores) was also
cited by Holttum as a reflection of cyatheaceous affinity. However, it seems more likely that this condition is
derived in Thelypteridaceae. Chromosome number
(27 -36 in Thelypteridaceae, 69 in Cyathea s.I.), indusial and sporangial characters, and vascular anatomy
of the stipe and rachis also separate the two families
and make a relationship between them tenuous. Aside
from the reniform indusia, bilateral perisporiate
spores, and similar sporangia, there is little to tie Thelypteridaceae to Dryopteris and other Aspidiaceae sensu Copeland.
KARYOLOGY AND HYBRIDIZATION. Base numbers include x=27 (Thelypteris subg. Parathelypteris. pt.); 29
(subg. Amauropelta); 30 (Phegopteris); 31 (Thelypteris
subg. Parathelypteris. Macrotheiypteris. Pseudophegopteris);32 and 33 (Coryphopteris. Thelypterissubg. Parathelypteris); 34 (subg. Lastrea); 35 (subg. Theiypteris.
subg. Metathelypteris. Cyclosorus subg. Pseudocyclosorus); and 36 (Cye/osorus. all subgenera). Most genera
(or subgenera) appear to have only a single base number, but several base numbers have been reported in
subg. Metathelypteris (x=31, 35, 36), subg. Parathelypteris (x=27, 31, 32), and Cye/osorus subg. Pseudocye/osorus (x= 35, 36). Both diploids and tetraploids are
Thelypteris . Phegopterls
known in many subgenera; a few higher ploids are also known. Natural hybrids occur in Cyclosorus subg.
Pelazoneuron (Smith 1971). Hybrids have been synthesized between several species in subg. Cyclosoriopsis
(Panigrahi and Manton 1958). Apogamy is known only in Phegopteris connectilis (Manton 1950) and possibly a species of subg. Goniopteris (Walker 1966).
CLASSIFICATION. Holttum (1971, 1982) grouped the
Old World species into 25 genera. Utilizing similar criteria, at least four additional American genera would
be recognized. Although many of Holttum's genera
seem natural (i. e., monophyletic), a combination of
characters must be used to circumscribe them. Some
of the characters concern minute glands and hairs and
require 30 x magnification or greater for observation.
Others require that complete specimens be at hand (including lower part of blade and stem). Even then,
identification to genus may be difficult, as generic
lines are not always sharp. Hybridization between genera is suspected (by Holttum himself) and transitional
species and species groups are known (such as between Sphaerostephanos and Pronephrium sect. Dimorphopteris in the Old World; between Goniopteris and
Meniscium in the New World). Unfortunately, some of
the best characters for circumscribing Holttum's
genera involve chromosome number and charactes of
spores best seen under the scanning electron microscope. Because of these problems a conservative classification is adopted herein. Three more or less discrete major lines of evolution can be discerned: (1)
Thelypteris; (2) Phegopteris, Pseudophegopteris, and
Macrothelypteris; and (3) Cyclosorus.
.KEy TO THE
GENERA.
1. Costae grooved adaxiaIly; veins free or anastomosing,
usually not forking
2
- Costae not grooved adaxially; veins always free, frequently
forked in the ultimate segments
3
2. Veins free. meeting the margin above the sinus; spores with
a fine to coarse reticulum
1. T/relypteris
- Veins meeting the margin at the sinus, connivent below the
sinus, or one or more pairs uniting below the sinus to fonn
areoies, rarely the veins ending above the sinus; spores
5. Cyclosorus
winged, rugose, or spinulose
3. Pinnae, at least in the distal half of the lamina, connected
by wings along the rachis, these wings often forming semicircular lobes between the pinnae, their veins arising from
the rachis; indusia absent or minute
2. Phegopterls
- Pinnae not adnate, the rachis not winged; indusia present4
m_
4. Sari exindusiate, often slightly elliptic; hairs unicellular;
basal segments of pinnae frequently enlarged
3_ Pseudophegopteris
- Sori indusiate (except in M. ornata), orbicular; hairs various; basal segments of pinnae not enlarged
5
5. Lamina bipinnate or more divided; hairs on abaxial side of
costae septate, often over 1 mm long
4. Macrothelypteris
- Lamina usually pinnate+pinnatifid (except in T.hattorii);
hairs unicellular, less than 1 mm long
1. The/ypleris subgenus
Metathelypteris
1. TheWteris Schmidel
265
Figs. 131 D, 134A,B, 138
Thelypteris Schmidel, Icon. PI. (ed. Keller): 3rd page, t. xi text
(Oct 1763), nom. cons.
Lastrea Bory (1824), nom. supeif!..
Amauropelta Kunze (1843).
Parathelypteris (H. Ito) Ching (1963).
Metathelypteris (H. Ito) Ching (1963).
Oreopteris Holub (1969), Holttum (1981).
Coryphopte/;s Holttum (1971, 1976a).
Stem creeping to erect. Lamina pinnate+pinnatifid, rarely pinnate or bipinnate [e. g., Th. pteroidea
(Klotzsch) R. Tryon, Th. hattorii (H. Ito) Tagawa], apex
never pinna-like; lower pinnae reduced or not; veins
always free, usually simple and reaching the margin
(except Metathelypteris) above the sinus. Sori indusiate
(except some Amauropelta species); sporangial capsule
glabrous, rarely with setae (a few species of Amauropelta) or glands (subg. Thelypteris). Spores ellipsoidal,
reticulate, with sometimes prominent elements, these
sometimes slender, somewhat contracted; or the surface echinate; or bearing low folds that have echinate
margins and are perforate; x=27, 29, 31, 32, 33, 34, 35.
Here treated as comprising five subgenera.
Subgenus Thelypteris: Stem long-creeping; lowermost pinnae not or only slightly reduced; aerophores
lacking; veins forked; broad, flat, tan scales on costae
below; sori indusiate; sporangial caspules with shortstipitate glands; X= 35; two species, Th. palustris
Schott in north-temperate areas to the Himalaya and
Th. confluens (Thunb.) Morton in Argentina, Africa,
India, Malesia, New Zealand; in wet or marshy
ground.
Subgenus Amauropelta (Kunze) A. R. Smith: Stem
creeping to erect, lowermost pinnae in most species reduced; veins usually simple; aero ph ores often swollen
or elongate at pinna bases; sori indusiate or exindusiate; x=29; c. 200 species, primarily in tropical
America, but nine in Africa, the Mascarenes, and Sri
Lanka, and three on Pacific islands, including Hawaii;
0-4500 m, mostly in or at edges of forests. Smith
(1974) divided the neotropical species into nine sections.
Subgenus Parathelypteris (H. Ito) R. & A. Tryon:
Stem slender, long-creeping; lowermost pinnae reduced or not; veins simple; aerophores absent; sori indusiate; x=27, 31, 32; c. 15 species, mostly East Asia
to Malesia and Sri Lanka, three in temperate North
America [Th. nevadensis (Baker) Clute ex Morton,
Th. noveboracensis, (L.) Nieuwl., Th. simulata (Oavenp.) Nieuwl.].
Subgenus Lastrea (Hook.) Alston (syn. Oreopteris):
Stem suberect; lowermost pinnae gradually reduced;
veins forked or simple; aerophores swollen (Th. elwesii) or lacking; lamina below with sessile glands; sori
indusiate; x=34; three species, Th.limbosperma (All.)
H. P. Fuchs in temperate Europe and eastern North
266
Filicatae . Thelypteridaceae
A
D
H
Fig. 134A-H. Thelypteridaceae. A,B Thelypteris palustris.
A Sterile segments (x 7). B Mature stem scale (x 40).
C-F Phegopteris decursive-pinnata. C Three scales from
rachis and costa (x25). D Two scales from stem (x 10).
E Detail of fertile lamina (x 2). F Scale from petiole (x 10).
G, H Cyc/osorus (Christella) dentatus. G Pinna bases with
rachis from adaxial side (x 3). H Enlarged portion of G
(x 12) (A,B from Holttum et al. 1970; C,D from Iwatsuki
1962; E,F from Flora Taiwan 1, 1975; G,H from Holttum
1977b)
America, Th. quelpaertensis (Christ) Ching in NE Asia
and NW North America, and Th. elwesii (Baker)
Ching in Sikkim (Holttum 1981).
Subgenus Metathelypteris (H. Ito): Stem short-creeping or erect; lowermost pinnae not or little reduced;
aerophores lacking; costae not grooved adaxially;
veins often forked, ending short of margin; sori indusiate, indusia often greenish; X= 35 (31 and 36 also reported); c. 12 species, East Asia south to Malesia,
Solomon Is., Sri Lanka, Africa, Fernando Poo, Sao
Tome.
Subgenus Coryphopteris (Holttum): Stem shortcreeping [Th. hirsutipes (Clarke) Ching] to usually
erect; lowennost pinnae not reduced; aerophores
swollen; veins simple, reaching the margin; lamina below often with sessile glands; indusiate (large); x=32,
33; 47 species, NE India and East Asia, especially
Malesia, to Samoa and the Marquesas; plants of peaty
soil on mountain crests (Holttum 1976a).
These subgenera are all treated as genera by Holttum (1971, 1982). Together they represent a well-defined genus, with subgenera more closely related to
each other than to any other genus of the family. Holttum allied Metathelypteris to Macrothelypteris and
Pseudophegopteris on the basis of the species having
forked veins ending in a clavate tip behind the margin
and having non-grooved costae. Its relationship is
probably closer to Thelypteris because of the large, persistent indusia, pinnate + pinnatifid blades, lack of
sporangial hairs and glands, and chromosome number; it may be an evolutionarily transitional group.
Holttum also allied Thelypteris s. str. to Cyc/osorus s.
str. because of their similar long-creeping stem and
costal scales, but venation, chromosome number, and
spore ornamentation suggest the relationship proposed here.
2. Phegopteris (c. Presl) Fee
Fig. 134C-F
Phegopteris (c. Presl) Fee, Gen. Fil.: 242 (1852), emend.
Ching (1963); Holllum (1969).
Stem long-creeping or suberect. Lamina bipinnatifid
or tripinnatifid in lower part, with deeply lobed adnate
pinnae connected by a wing along the rachis, wing of-
267
Pseudophegopteris· Macrothelypteris· Cye/osorus
ten fonning a lobe between pinnae and served by a
vein arising from rachis; veins free, simple or forked,
the distal ones of a segment reaching the margin or
nearly so; rachis and costae below with spreading lanceolate scales. Sori exindusiate or minutely indusiate;
sporangial capsule often bearing acicular hairs or
stipitate glands; spores ellipsoidal, near plain, tuberculate, or perforate with raised ridges; x~ 30.
Three species, P. hexagonoptera (Michx.) Fee in
eastern North America; P. connectilis (Michx.) Watt,
circum boreal and north-temperate; and P. decursivepinnata (van Hall) Fee, in East Asia.
3. Pseudophegopteris Ching
Pseudophegopteris Ching, Acta Phytotax. Sin. 8: 313 (1963);
Holttum (1969).
Stem creeping or erect. Lamina bipinnate + pinnatifid
to pinnate+pinnatifid; petiole and rachis often castaneous; pinnae opposite to subopposite, sessile or
partially adnate to rachis, lowennost pinnae usually
somewhat reduced, basal pinnules often enlarged;
aerophores lacking; veins free, often forked, tips adaxially enlarged, not reaching the margin; scales usually
lacking on costae below; hairs unicellular. Sori exindusiate, often oblong; sporangial capsule often bearing hairs or stipitate glands. Spores ellipsoidal, with
low, reticulate surface features; x~31.
About 20 species, tropical and subtropical Asia,
Malesia, Samoa, Hawaii, the Mascarenes, Madagascar, Africa, Fernando Poo, Silo Tome, and St. Helena,
mostly at middle elevations, 1000-2800 m.
4. Macrothelypteris (H. Ito) Ching
tions, 0-1000 m [M. po[ypodioides (Hooker) Holttum to
2150 m], often in somewhat weedy habitats.
5. Cyclosorus Link
Figs. 134G,H, 135, 136
Cye/osorus Link, Hort. Reg. Bot. Berol. 2: 128 (1833).
Meniscium Schreber (1791); Stegnogramma Blume (1828);
Goniopteris C. Presl (1836); Sphaerostephanos J. Smith (1839),
HolttuIIi (1979); Leptogramma J.Smith (1841); Ampelopteris
Kunze (1848); Pronephrium C. Presl (1849), Holttum (1972);
Haplodictyum C. Presl (1849), Holttum (1973c); Abacopteris
Fee (1852), Iwatsuki (1959); Dictyocline Moore (1855); Glaphyropteris Fee (1873); Christel/a Uveille (1915), Holttum
(1976b); Pneumatopteris Nakai (1933), Holttum (1973 b); c)!"
clogramma Tagawa (1938); Menisorus Alston (1956); Dimorphopteris Tagawa & Iwats. (1961); Glaphyropteridopsis Ching
(1963); Pseudocyclosorus Ching (1%3), Holttum and Grimes
(1979); Trigonospora Holttum (1971); Mesophlebion Holttum
(1971); Chingia Holttum (1971); Nannothelypteris Holttum
(1971, 1973c); Amphineuron Holttum (1971, 1977a); Steiropteris (C. Chr.) Pichi Sermolli (1973); Plesioneuron (Holttum)
Holttum (1975).
B
Fig. 137
Macrothelypteris (H. Ito) Ching, Acta Phytotax. Sin. 8: 308
(1963), Holttum (1969); based on: Thelypteris sect. Macrothelypteris H. Ito in Nakai and Honda (1939).
Stem short-creeping, thick. Lamina bipinnate + pinnatifid; petiole and rachis stramineous to tan; pinnae opposite to mostly alternate distally, lowermost not reduced, largest petiolulate; basal pinnules usually reduced; veins free, often forked, tips not reaching the
margin; scales sometimes present on costae below,
these from a thickened base; costae and costules below usually with septate hairs mostly over 1 mm long.
Sori indusiate [except M. ornata (Wall. ex Bedd.)
Ching], indusia often small, obscured in mature sori;
sporangial capsules bearing short-stipitate glands.
Spores ellipsoidal, usually reticulate or with perforate
folds; x~ 31.
About 10 species in tropical and SUbtropical Asia,
Malesia, Queensland, islands of the Pacific, Africa.
One species, M. torresiana (Gaud.) Ching, is widely
naturalized in the New World. Mostly at low eleva-
Flg.135A-C. Thelypteridaceae. A,B Cyclosorus (Stegnogramma) aspidioides. A Sterile pinna (x 0,7). B Portion of
fertile pinna showing extent of sori (x 3). C Cyclosorus
(Stegnogramma) subcalcaratus, portion of fertile pinna (x 6)
(Holttum 1982)
Filicatae . Thelypteridaceae
L.',e.,.:;
L . '-f.
.
~
i""
F:'
L;:
....
.
B
".
.,
."
11
E
-'.
.
C.
F
A
Fig. 136A-O. Thelypteridaceae. A,B Cyclosonls (SpaerostephanOs) pen niger. A Leaf (x 0,25). B Reduced basal pinna
with aerophore (x 2). C Cyclosonls (Spaerostephanos) pyenaSOnlS, fertile segments (x 4). D Cyclosonls (Spaerostephanos)
deeadens, part of segment with soms and indument (x 16).
E, F Cyclosonls (Leptogramma) totloides. E Detail of fertile,
pinna (x 10). F Stem scale (x 100). G Cyclosonls normalis,
scale from petiole base (x 20). H Cyclosonls serra, scale from
costa (x 20). I,J Cyelosonls (Steiropteris) po/yphlebius. I Part
of rachis with pinna bases (x 0,4). J Fertile segment
(x 1,25). K,L Cyclosonls (Meniscium) salzmann;;. K Base of
lamina ( x 0,3). L Part of sterile pinna (x 2). M Cyclosonls
(Dietyocline) griffith;;, part of pinna showing venation (x 1,5).
N,O Cye/osoros (Goniopteris) tetragonus. N Scale of stem
(x 10). 0 Detail of margin (x 25) (A,B from Holttum 1982;
C,D from Holttum 1977b; E,F,M from Flora Taiwan 1,
1975; G,H from A.R.Smith 1971; I-L from A.R.Smith
1983; N,O from Iwatsuki 1%2)
Filicatae • Thelypteridaceae
Stem creeping to erect. Lamina simple to usually pinnate or pinnate + pinnatifid, apex sometimes pinnalike; lower pinnae reduced or not; veins free to usually
connivent at the sinus or anastomosing below it, often
forming a regular network of areoles; sori indusiate or
exindusiate; sporangial capsule glabrous or often with
sessile glands or setae, stalks also often with glands or
hairs. Spores monolete and ellipsoidal or rarely trilete
and somewhat spheroidal; surface often reticulate or
with short, low ridges, or prominently winged, the borders sometimes ciliate, or short-cristate to echinate;
X= 36 (perhaps also 35 in Pseudocyclosorus).
Here treated as comprising 20 subgenera, none of
which have nomenclatural combinations available. As
here circumscribed, Cyclosorus is antedated by both
Meniscium and Stegnogramma, both aberrant elements
but connected to more typical elements of the genus
by intermediates. Nomenclatural combinations for
most species already exist in Cyclosorus, which if
maintained with this circumscription must be conserved.
Natural hybrids have been found between, e. g.,
subgen. Cyclosoriopsis and subgen. Pneumatopteris
(Quansah and Edwards 1986).
Subgenus Cyclosorus: Stem long-creeping; lowermost pinnae not reduced; basal veins anastomosing
with excurrent vein to sinus; broad scales on costae
below; sori indusiate; sporangial stalks with stalked
globose reddish glands; 2-4 species, pantropical and
subtropical in freshwater swamps, often in sun (Holttum 1974a).
Subgenus Ampelopteris (Kunze): Stem creeping;
lamina indeterminate, bearing buds (and plantlets) at
bases of pinnae, lowermost pinnae not reduced; veins
anastomosing (5 or 6 pairs) to form areoles and zigzag
ex current vein; sori exindusiate; sporangial stalks with
stalked globose glands; monotypic; West Africa, tropical Asia to NE Australia, river banks and ditches.
Closely related to subg. Cyclosorus.
Subgenus Mesophlebion (Holttum): Stem shortcreeping; lowermost pinnae not reduced; veins free,
meeting the margin at or near the sinus, basal basiscopic vein of each group arising from the costa; sori
indusiate [except c. oligodictyus (Baker) Holttum]:
sporangial stalks bearing stalked globose orange-red
glands; 17 species, Malesia, in forests, 0-1500 m. Related to subg. Cyclosorus, as indicated by the sporangial glands (Holttum 1975).
Subgenus Steiropteris (c. Chr.): Stem usually creeping; lowermost pinnae not reduced [except. Th. deltoidea (Swartz) Proctor]; aerophores often elongate at
pinna bases; veins free to connivent below sinus; a
cartilaginous pseudovein often leading from sinus towards costa; sori indusiate (most species) or exindusiate; 21 species, Neotropics, in forests, 0-1500 (2100)
m (Smith 1980; Fig. 136). Closely related to
269
B
Fig. 137 A-C. Thelypteridaceae. Macrothelypteris torresiana.
A Part of rachis and pinna bases, adaxial side (x 2). B Part
of fertile segment (x 24). C Transections of petiole (x 2)
(Holttum 1977)
Mesophlebion as indicated by similar aspect and venation [cf., e. g., Th. glandulosa (Desv.) Proctor and
Th. insignis(Mett.) Ching (Steiropteris) with Th. motleyana (Christ) Holttum and Th. crassifolia (Blume)
Ching (Mesophlebion)] and also the stalked paraphyses
of Th. valdepilosa (Baker) Reed.
Subgenus Goniopteris (C. Presl): Stem creeping to
suberect; lowermost pinnae usually not reduced; proIiferous buds often present at pinna bases; aerophores
usually lacking; veins free to anastomosing in one to
many series; axes and often lamina with sessile or
stalked furcate or stellate hairs (absent in 3 species);
sori indusiate or exindusiate; c. 60-75 species, Neotropics, in or at edges of forest (Christensen 1913).
Most closely related to subg. Steiropteris through such
species as 1h.fraseri (Mett. ex Kuhn) A. R Smith and
Th.lugubriformis (Rosenst.) R. Tryon (the only species
with swollen aerophores).
Subgenus Meniscium (Schreber): Stem creeping;
lamina simple or usually I-pinnate with conform apical segment, pinnae entire to serrate, lowermost not re-
270
Filicatae • Thelypteridaceae
duced; a bud often present in the axil of a basal pinna; veins regularly uniting, with cross-veins usually
bearing a single elongate sorus; hairs acicular or lacking on lamina; sori exindusiate; c. 20 species, New
World tropics and subtropics, in forests (occasionally
swamps), especially along streams, 0-1500 m (Maxon
and Morton 1938). Closely related to and derived from
subg. Goniopteris.
Subgenus MeniwIUS (Alston): Stem erect; lamina 1pinnate, pinnae serrate, lowermost not reduced; a bud
present at the base of a distal pinna; veins anastomosing; sori exindusiate; monotypic; tropical Africa,
along streams, at lower elevation.
Subgenus Abacopteris (Fee): Stem creeping to suberect; pinnae crenate to entire, veins usually regularly
uniting; lower surface of lamina often pustular when
dried and with sessile yellowish glands; sori exindusiate or indusiate; sporangial capsules often setose or
glandular; comprising 4 sections, from India throughout Malesia to Queensland and Fiji: sect. Abacopteris
(syn. Pronephrium, Haplodictyum), 15 species; sect. Dimorphopteris (syn. Nannothelypteris), 36 species; sect.
Menisciopsis (Holttum), 10 species; sect. Grypothrix
(Holttum), 11 species (Holttum 1972, 1982).
Subgenus Sphaerostephanos (1. Smith): Stem creeping to erect; lowermost pinnae reduced; aerophores
often swollen; veins usually anastomosing; sessile yellowish glands on lamina below; sori usually indusiate;
sporangia often bearing yellow glands or setae on capsule; c. 176 species, tropical Africa and Asia, Pacific
islands, in forests (Holttum 1982).
Subgenus Stegnogramma (Blume); syn. Leptogramma, Dictyocline: Lowermost pinnae not or little reduced; distal pinnae adnate to rachis; aerophores
usually lacking [except Th. aspidioides (Willd.) R Tryon]; veins free to anastomosing; sori exindusiate,
spreading along veins; sporangia setiferous; 15 species, Africa, Europe, Asia, Malesia, 1 species in the
New World, Alabama to Honduras (Iwatsuki 1963).
Two southern Brazilian species often included here
form a separate, unnamed subgenus most closely allied to Goniopteris.
Subgenus Glaphyropteridopsis (Ching): Lowermost
pinnae not reduced, aerophores swollen; veins free;
sori exindusiate; 4 species, NE India, China.
Subgenus Pelazoneuron (Holttum): Lowermost pinnae not reduced or 1-6 pairs reduced; aerophores
lacking; veins connivent just below the sinus or meeting the margin at or just above sinus; sori indusiate;
23 species, tropical and sUbtropical Africa and America, mostly low to middle elevations (Smith 1971 ; Holttum 1974a).
Subgenus Cyclosoriopsis (Iwatsuki); syn. Christella:
Lowermost pinnae usually reduced; aerophores lacking; veins ususally anastomosing in one or more series
below sinus; sori indusiate; sporangia with stalked
pyriform glands on stalk; c. 60 species, most of them
in India and SE Asia; two endemic to Hawaii; several
in Africa and adjacent islands; only two species [Th.
hispidula (Decne.) Reed, Th, conspersa (Schrader)
A. R Smith] native to Neotropics (Holttum 1976b;
Smith 1971).
Subgenus Amphineuron (Holttum): Lowermost pinnae not reduced or in a few species 1 or 2 (3) pairs reduced; aerophores not swollen; veins free to anastomosing below sinus; sori indusiate or exindusiate;
elongate glandular hairs on sporangial stalks; c.
12 species, tropical East Africa, Asia, Malesia,
Queensland to Tahiti (Holttum 1977 a, 1982). One species widely naturalized in Neotropics [Th. opulenta
(Kaulf.) Fosberg].
Subgenus Cyc/ogramma (Tagawa): Lowermost pinnae reduced or not; elongate aerophores usually present at pinna bases; hooked hairs on axes below and
often on sporangia; veins free; sori exindusiate; c.
8 species, northern India to South China, Taiwan,
Philippines. Closely related to Stegnogramma [cf., e. g.,
Th. ("S. 'j pozoi (Lag.) Morton and Th. ("e. 'j omeiensis
(Ching) Reed which are very similar in blade form and
venation; the latter has echinate spores similar to those
of Stegnogramma].
Subgenus Pneumatopteris (Nakai): Lowermost pinnae reduced; aerophores at pinna bases ± swollen
and sometimes elongate; veins usually anastomosing;
lamina ± pustular when dried; sori usually indusiate;
c. 80 species, tropical Africa, Asia, Malesia, and Pacific islands to Hawaii and New Zealand, in forests,
mostly at low to middle elevations (Holttum 1973b,
1982).
Subgenus PseudocyciosolUS (Ching): Lowermost
pinnae reduced; aerophores at pinna bases ± swollen; veins free, lowermost meeting the margin at or
just above the sinus; lamina not pustular; sori indusiate; 12 species, tropical Africa, India, and SouthEast Asia, in Malesia only in the Philippines (Holttum
and Grimes 1979).
Subgenus Trigonospora (Holttum): Stem erect; lowermost pinnae not or little reduced; aerophores lacking; veins free, lowermost meeting the margin at or
just above sinus (anastomosing in Th. khamptorum
Holttum); sori indusiate; spores trilete, minutely papillose; c. 9 species, Sri Lanka (7 species) and India to
South-East Asia (Holttum 1982; Sledge 1981).
Subgenus Chingia (Holttum): Stem erect, sometimes
trunk-like; lowermost pinnae not reduced; aerophores
not enlarged; stipe bases (at least) with persistent, rigid, setiferous scales; veins usually anastomosing; sori
usually near costules; sori exindusiate or in a few species indusiate; 20 species, Malesia and Pacific islands
to Tahiti, in mainland Asia only west to peninsular
Thailand (Holttum 1974b, 1982).
Subgenus Plesioneuron (Holttum): Stem short-creep-
Selected Bibliography
271
defining characters of the group; at least four species
of Cyclosoriopsis lack the characteristic pyriform
glands on sporangial stalks; and a few species of Goniopteris lack stellate or furcate hairs. There is considerable variation in other characters used by Holttum to
define genera, e. g., stem habit (long-creeping to erect
in Cyclosoriopsis and Sphaerostephanos), aerophores
[absent in Cyclosoriopsis except in Ch. distans (Hooker)
Holttum] (Fig. 138), spores, pustular lamina (in most
but not all Pneumatopteris. also in some Goniopferis
and Pronephrium). Reversals of character states seem
likely to have occurred, e. g., free venation is derived in
Cyclosoriopsis and Sphaerostephanos (Holttum 1982),
while being primitive for the family as a whole. Hybridization is suspected among certain subgenera, e. g., between Pneumatopteris afra and Cyclosoriopsis (Holttum
1974a), Cyclosoriopis and Trigonospora (Sledge 1981),
and Sphaerostephanos and Pronephrium (Holttum
1982, p.495). Convergent, parallel, and reticulate evolution have played such an important part in the development of Cyclosorus s.1. that the interests of taxonomists seem best served by the recognition of a single
genus, no more variable than many other fern
genera (as, e.g., Asplenium, Adiantum, Blechnum).
Selected Bibliography
Fig. 138. Thelypteridaceae. Thelypteris thomsoni~ crozier with
pale aerophores on primary and secondary rachises; Mexico.
Pho!. D. Breedlove
ing to erect; lowermost pinnae not or little reduced;
aerophores usually somewhat swollen or elongate;
lower surface often verrucose when dry; veins free;
basal basiscopic vein arising from costa; sori indusiate
or exindusiate; c. 45 species, Malesia and Pacific islands to Tahiti, mostly in montane forests, often by
streams (Holttum 1975).
Holttum (1982 and revisions cited therein) recognized all of the Old World subgenera of Cyclosorus
(except Pelazoneuron) at generic rank. These groups
(as subgenera or genera) are generally definable only
by a combination of variable characters, if at all. Traditional characters used for generic circumscription
cannot be applied. For example, both indusial presence and absence occur in 9 of 20 subgenera; 10 of
20 subgenera have both free-veined and anastomosing-veined species; and presence and absence of reduced basal pinnae are conditions found in 8 of
20 subgenera. Holttum relied most heavily on characters of sporangial glands and hairs. These, too, are
variable within a group, e. g., about 15 species of Sphaerostephanos lack sessile glands - one of the principal
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Atkinson, L. R. 1975. The gametophyte of five Old World
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Ching, R. C. 1963. A reclassification of the family Thelypteridaceae from the mainland of Asia. Acta Phytotax. Sin. 8:
289-335.
Christensen, C. 1907. Revision of the American species of
Dryopteris of the group of D.opposita. Kongel. Danske
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Christensen, C. 1913. A monograph of the genus Dryopteris J.
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Kongel. Danske Vid. Selsk. Skr., Naturvid. Afd. VII, 10:
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Christensen, C. 1920. A monograph of the genus Dryopteris
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Grimes, J. W. 1980. Spore morphology in Thelypteridaceae. I:
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Hennipman, E. 1968. The mucilage secreting hairs on the
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Holttum, R. E. 1960. Vegetative characters distinguishing the
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sori. Gard. Bull. Singapore 17: 361-367.
Holttum, R. E. 1969. Studies in the family Thelypteridaceae.
The genera Phegopteris, Pseudophegopteris. and MacrotheIyptens. Blumea 17: 5-32.
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272
Filicatae . Vittariaceae
Holttum, R. E. 1972. Id. IV. The genus Pronephrium Presl.
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Holttum, R. E. 1973 a. The family Thelypteridaceae in the Old
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Holttum, R. E. 1973c. Id. VI. Haplodictyum and Nannothelypteris. Kalikasan 2: 58-68.
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VII: The genus Chingia. Kalikasan 3: 13 - 28.
Holttum, R. E. 1975. Studies in the family Thelypteridaceae
VIII. The genera Mesophlebion and Plesioneuron. Blumea
22: 223-250.
Holttum, R. E. 1976a. Studies in the family Thelypteridaceae
X. The genus Coryphopteris. Blumea 23: 18-47.
Holttum, R. E. 1976b. Studies in the family Thelypteridaceae
XI. The genus Christella Leveille, sect. Christella. Kew
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Holttum, R. E. 1977a. Id. XII. The genus Amphineuron Holttum. Blumea 23: 205-218.
Holttum, R. E. 1977b. The family Thelypteridaoeae in the Pacific and Australasia. Allertonia 1: 169-234.
Holttum, R. E. 1979. Sphaerostephanos in Asia, excluding
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Holttum, R. E. 1981. The genus Oreopteris (Thelypteridaceae). Kew Bull. 36: 223-226.
Holttum, R. E. 1982. Thelypteridaceae. Flora Malesiana II, 1
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Holttum, R. E., Grimes, J. W. 1979. The genus Pseudocye/osorus Ching (Thelypteridaceae). Kew Bull. 34: 499-516.
Holttum, R. E., Sen, U., Mittra, D. 1970. Studies in the family
Thelypteridaceae II. A comparative study of the type-species of Thelypteris Schmidel, Cye/osorns link, and Ampelopteris Kunze. Blumea 18: 195-215.
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Iwatsuki, K. 1965. Taxonomy of the thelypteroid ferns, with
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