Professional Documents
Culture Documents
Fern Grower's Manual - Revised and Expanded Edition PDF
Fern Grower's Manual - Revised and Expanded Edition PDF
TIMBER PRESS
Portland, Oregon
The in-text Adiantum line drawings are by Lynn Vander Velde; unless otherwise
credited in the captions, all other line drawings are by Barbara Joe Hoshizaki.
Unless otherwise credited in the captions, the in-text black-and-white photographs are
by Arthur Takayama.
Unless otherwise credited in the captions, the color plates are by Barbara Joe Hoshizaki.
Copyright © 2001 by Barbara Joe Hoshizaki and Robbin C. Moran. All rights reserved.
Published in 2001 by
Timber Press, Inc.
The Haseltine Building
133 S.W. Second Avenue, Suite 450
Portland, Oregon 97204, U.S.A.
Preface 7
1. About Ferns 11
2. The Structure of Ferns 13
3. Obtaining Ferns 22
4. Cultural Needs 25
5. Soils and Fertilizers 37
6. Through the Year with Ferns 48
7. Planting 51
8. Propagation 63
9. Landscaping 87
10. Growing Special Ferns 99
11. Troubles with Growing Ferns 118
12. How Ferns Get Their Names 140
13. Ferns and Fern Allies in Cultivation 151
Color plates, including USDA Plant Hardiness Zone Map, follow page 240
5
This page intentionally left blank
Preface
Ferns continue to hold the interest of gardeners as we enter a new millennium. After the Victorian fern
craze, interest in ferns focused mainly on the native species and some of those common in the trade. After
World War II, interest in exotic ferns gradually took hold, and this fostered the formation of new fern
societies and the offering of a greater selection in the trade.
It has been more than 25 years since the first edition of Fern Grower’s Manual was published (Hoshizaki
1975). At the time, the author thought it would interest the horticultural public for only a few years before
ferns would fall out of vogue. She was wrong. A strong interest in ferns continues today as evidenced by the
great increase in fern societies all over the world for both amateurs and professional botanists. Also, the
number of fern species in cultivation has increased tremendously. The contributions of fern botanists in
writing floras and monographs, clarifying species, and supporting amateur interest are important factors in
this growth. The horticultural area has also made great contributions, particularly in pest control, tissue
culture, plug culture, new products, and determining the growing needs of certain ferns.
All this interest in ferns has created a need for a revised edition of Fern Grower’s Manual—one that
updates the information on pest control, propagation, new materials and products, cultural requirements,
and many other topics. But unlike other recent fern books, this revised edition emphasizes identifying cul-
tivated ferns by way of the text and illustrations for the 124 genera and more than 700 species treated (each
species is illustrated). Because diagnostic features are often unclear in photographs, this edition relies heav-
ily on silhouettes and line drawings. With additional help from the book’s glossary and a good hand lens,
the reader should be able to identify most of the cultivated species.
This revised edition treats all the commonly cultivated (and some rarely cultivated) fern species, vari-
eties, and many cultivars in the United States. Besides identification, this new edition will help readers learn
more about a particular fern by including its geographical distribution and author citations after the species
names. This information enables readers to have a better idea of which floras, texts, or journals to consult
for further information about a fern. An expanded bibliography also points to other literature sources.
This book is mainly written for people seriously interested in growing ferns, knowing their names and
what makes them similar or different, and appreciating their diversity. It is not a coffee-table book, nor a
chatty type of garden book meant for light reading. Beginning fern amateurs may find more information
than they need, but they will also find information useful at their level. Although this book primarily is a
reference, it is also for browsing and gleaning bits of information not readily found elsewhere.
The core information in this book will be particularly helpful to plant people who want to grow or iden-
tify different ferns and fern allies. Gardeners, nursery people, commercial growers, plant inspectors, botan-
ical garden staff, horticulturists, botanists, biology teachers, and others will find the information they need
in this one volume. The alternative would be to have a library of horticultural books and floras from the
7
8 P
world over and/or access to a fern botanist with a large herbarium. Botanists needing to grow ferns for their
research will find specific information on growing genera of ferns as well as general cultural information.
Students in horticulture, botany, and biology will find that this book gives an overview of fern infor-
mation that is basic and up-to-date, but they also should note that much remains to be discovered. In addi-
tion to examining the growth response of ferns to various environmental conditions, there is also much to
clarify on the relationships of genera and species. It is hoped that this book will stimulate students to study
horticultural and botanical problems of ferns or use them in their research. Although this type of problem
solving has been traditionally left to the scientist, observant, thoughtful, and innovative growers have made
and will continue to make valuable contributions to studying ferns (pteridology). Gardeners without sci-
entific training have introduced new species to horticulture, produced new cultivars and hybrids, and
found the right combination of conditions to grow difficult species. Much is unknown about temperature
tolerances of ferns, and this too is information that may be observed by growers and added to the store of
knowledge about ferns. Commercial growers have worked out detailed cultural information on growing
Boston ferns and leather ferns. Cultural details for other important trade groups have yet to be fully
reported.
Keeping an accurate list of the species in cultivation has become increasingly difficult, if not impossi-
ble. Many name changes since the first edition of this book have greatly complicated the task. Moreover,
new ferns are constantly being introduced to horticulture. There are about 10 fern societies in the world
whose members often bring new ferns into cultivation, and the number of species that can still be intro-
duced is sizable. An annual list recently offered by a fern society listed 775 packets of spores for exchange.
With so many possible species to list, it was thought better to err on the side of including too many instead
of too few. Several new and promising introductions came to our attention too late to be fully treated.
Several species listed as cultivated could not be verified, and others may have been overlooked. Since few
monographs have been written about problem ferns, identification was sometimes difficult and often com-
plicated by not knowing the country of origin.
Acknowledgments
Writing this book has been a prolonged and immense undertaking, and many people have helped and
encouraged us to reach the finish line. We especially wish to thank those people who read portions of the
manuscript for accuracy and gave helpful suggestions. They are Dr. Wade L. Berry, a plant physiologist and
soil scientist of the University of California, Los Angeles, who edited the cultural information; Dr. A. James
Downer, Farm Advisor, of the University of California Extension, Ventura, who edited the section on
insect pests and diseases; Dr. Ole Becker, of the University of California, Riverside, who edited the section
on nematodes; and botanist Sharon Ishikawa, of the United States Department of Agricultural, Animal and
Plant Health Inspection Service, who edited the section on importing plants.
We are indebted to the kindness of Dr. Alan R. Smith, of the University of California, Berkeley, and
Dr. John T. Mickel, of The New York Botanical Garden, for their assistance and time while we worked at
their herbaria. We also thank Dr. Art Gibson, of the University of California herbarium, for use of their
facilities. For identifying difficult ferns, we wish to thank the following people: Dr. Christopher R. Fraser-
Jenkins for Dryopteris; Dr. Richard Hauke for Equisetum; Dr. Blanca León for Campyloneurum; Dr. John
T. Mickel for Elaphoglossum and hardy ferns; Dr. Alan R. Smith for Thelypteris and other genera; Dr. Rolla
M. Tryon for cheilanthoids and suggestions on genera designations; and Dr. Iván Valdespino for Selaginella.
We were fortunate to have the help of other pteridologists from around the world. For Asian ferns we
were helped by many Japanese botanists. We are particularly grateful for identifications from Dr. Toshiyuki
Nakaike, Prof. Keisuke Yasuda, and Prof. Norio Sahashi. Also of assistance were Prof. Masahiro Kato,
P 9
Prof. Reiko Yoroi, Prof. Ryoko Imaichi, Dr. Haruki Hirabayashi, Dr. Mitsuyasu Hasebe, and Dr. Noriaki
Murakami. The late Dr. Warren H. Wagner Jr., Dr. Dan Palmer, and Dr. Barbara Parris were valuable
sources of information on Pacific Island ferns. Dr. David B. Lellinger supplied information about Costa
Rican ferns, Dr. Lynn Raulerston about Guam and Palau ferns, Martin Rickard about English ferns,
Michael Price about Philippine and Malaysian ferns, and Dr. Peter Bostock and Chris Goudey about Aus-
tralian ferns. Yolanda Orta and Martin Grantham shared their expertise about fern propagation. For keep-
ing us informed of ferns in their area and for generous assistance in other ways, we are indebted to Mari-
lyn Johnson of Florida, Judith Jones of Washington, John and Carol Mickel of New York, and Nancy
Swell of Virginia.
We also thank the following for obtaining specimens and answering specific questions: the late Virginia
Ault, Ed and Phyllis Bates, Betsy Feuerstein, Robin Halley, the late Gerda Isenberg, Hildegard and Bruce
Jackson, Al and Marilyn Johnson, Judith Jones, Miriam Leefe, Robert Manthorne, Dr. Bruce McAlpin,
Duane Petersen, Milton Piedra, Ruth Radcliffe, Martin Rickard, the late Dorothy Skula, Nancy Swell, Roy
Vail, Helen Woodley, and Reggie Whitehead. In addition, we learned much from interacting with many
members of the South Florida Fern Society, the Tampa Fern Society, the New York Chapter of the Amer-
ican Fern Society, the Los Angeles International Fern Society, the San Diego Fern Society, the Hardy Fern
Society, and the Southwestern Fern Society.
Many commercial growers were generous with their help and information. We thank Ades and Gish
Nursery, Barford’s Hardy Ferns, California Ferns, Casa de Flora, Clark’s Nursery, Charles Alford Plants,
Daisy Plant Farm, Eisenbraun Ferns, Fancy Fronds, Fender’s Nursery, Fern Farm, Foliage Gardens, Frank
and Dorothy Skula, Freshwater Fauna and Flora, Glasshouse Works, H. Marugame, Half-moon Bay Nurs-
ery, Henry’s Plant Farm, Huntley Greens, Jerry Horne, Lyndon Horticultural Enterprises, Marilyn John-
son, Monrovia Nursery, MSK Rare Plant Nursery, Plants Unlimited, Santa Rosa Tropicals, Siskiyou Rare
Plant Nursery, Swell’s Nursery, Talnadge Fern Gardens, Tropical Gardens, Varga’s Nursery, Weather-
wood Nursery, Wedel’s Nursery, Weeks Nursery, Yerba Buena Nursery, and Yolanda’s Ferns.
For helping with computers, locating citations, assisting with the illustrations, and just making things
work smoother, we would like to acknowledge the help of Joan DeFato, Carol Brooks-Hoshizaki, Jon and
Madeleine Hoshizaki, Janet Keyes, Harry Le Vine Jr., Alvin Lee, Sylvia Louie, Vickie Milane, Mamie
Moy, Damon Woods, and Leo Yamanaka. We wish to acknowledge Arthur Takayama for taking and
preparing most of the black-and-white photographs and Lynn Vander Velde for the Adiantum line draw-
ings. When the senior author desperately needed editing assistance, Ed and Phyllis Bates, Ann Herrington,
Marilyn Johnson, and Susan MacQueen came to her rescue. Dr. Kenneth Wilson has been a source of
unflagging help, ranging from providing botanical expertise to trouble-shooting computer problems. The
senior author’s husband, Dr. Takashi Hoshizaki, a plant physiologist, suggested doing this revision more
than 15 years ago. He helped with the editing and artwork, but most importantly gave cheerful support and
encouragement over the years.
The authors apologize for any oversights in our acknowledgments and wish to assure all that their con-
tributions are nevertheless deeply appreciated.
The senior author also expresses her gratitude and deepest thanks to the junior author, Dr. Robbin
C. Moran, for consenting to join in producing this book. Not only is he one of today’s leading fern scien-
tists, he has also demonstrated his interest and ability in writing for general fern audiences. He has added
valuable information to the text, clarified complicated topics, and improved the book in many ways. With-
out his broad knowledge of ferns, his patience, and his energy to check on innumerable details, make the
required changes, do the editing, and work with Timber Press, this book would have languished much
longer in manuscript form.
We wish to especially acknowledge and thank our editor, Josh Leventhal. He made our work so much
easier and the book so much better with his careful editing and good judgment.
10 P
Although we relied heavily on the counsel of many people and are greatly indebted to their contribu-
tions, any discrepancies or errors in this publication are entirely our own. The information and advice con-
tained in this work have been brought together with care, but because of the large number of variables we
cannot guarantee complete accuracy or results. The reader acts on his or her own responsibility.
Robbin C. Moran
The New York Botanical Garden
1 About Ferns
Ferns bring to mind pleasant, cool glens and shaded forests. We recall such things when we use ferns in our
gardens and homes. The ferny look is well known and loved, and some of us are content to sit back and relax
as we enjoy the soft green array of patterns and textures in ferns. But there is more to appreciate about ferns
than their mere appearance.
Ferns are usually recognized by their finely divided leaves, a type of leaf so characteristic that it is called
a “ferny leaf.” There are, however, many plants with ferny leaves that aren’t ferns, and many ferns that don’t
have typical ferny leaves. What, then, makes a fern a fern?
Ferns are spore-bearing plants; they lack flowers, fruits, and seeds. The so-called asparagus fern (Aspara-
gus setaceus) is not a fern despite its finely divided appearance because it bears flowers and seeds (it is actu-
ally a member of the lily family; its fruits are the orange berries often seen on the plant). Spores and seeds
greatly differ. Spores are simple one-celled microscopic structures, whereas seeds are complex, many-celled
structures usually visible to the unaided eye. Although plants such as algae, liverworts, and mosses produce
spores, they differ from ferns by lacking the large, thin, true leaves. Ferns further differ from these spore-
bearers by their development of specialized tissue (xylem and phloem) to conduct food and water. These
tissues also strengthen the stem and enable ferns to grow taller than other spore-bearing plants.
Unlike seed plants, ferns depend on water to complete their typical life cycle. They grow in places
where, when the time comes to reproduce, enough water is available for the sperm to swim to the egg.
Seed plants, such as pines and flowering plants, produce cones or flowers that use wind or insects to com-
plete their life cycle. They do not need water in the external environment for fertilization, and therefore they
can grow in drier conditions and dominate more of the landscape.
Nevertheless, some ferns have the surprising ability to thrive in extreme climates. Desert ferns, of which
there are few, often grow in the shade of rocks and boulders, using every bit of available water. Their roots
grow deep in the soil between the cool rocks, and their fronds are often covered with woolly hairs or scales
to protect them from water loss. When water is insufficient for new growth, many desert ferns curl their
leaves or shed their leaflets and suspend growth until the next rain.
Like desert ferns, alpine ferns are also adapted to climatic extremes. They tend to be small and have
hard-textured fronds that can endure the cold, dry winds. They grow only during the short summers as
weather permits.
Most ferns of temperate regions grow in the ground or on rocks, and only a few species grow on trees.
Approximately 200 species of ferns are native to the temperate areas of the United States. Temperate to sub-
tropical areas noted for their abundance of ferns include parts of the Himalayas, Australia, New Zealand,
and Japan. The tropics, however, harbor the largest number of fern species, particularly at middle elevations
in the mountains. Here the mountain fog encourages an abundance of ferns.
11
12 C 1
New ferns are still being discovered in well-known parts of the world, and in the lesser known parts
many more remain to be discovered. There are about 12,000 named species, and many of these have never
been introduced into horticulture. Temperate places like Japan and the cool, mountain slopes of the trop-
ics have many ferns suitable as new introductions for temperate gardens in the United States.
Compared to other plants, ferns have few economic uses. Nevertheless several are important. Ferns and
fern allies dominated the extensive coal-forming swamps of the Carboniferous period 280 to 345 million
years ago. Coal, therefore, is often composed of the compressed remains of ancient ferns and fern allies. If
you are reading this book by electric light and your local power plant is fueled by coal, chances are that the
energy used to generate the electricity for your bulb was captured millions of years ago by a fern or fern ally!
An important economic fern is Azolla, the mosquito fern. It is a rich source of nitrogen, a nutrient usu-
ally limiting to plant growth. Azolla is often used as an organic fertilizer in the rice paddies of southeastern
Asia. Considering how many people rice feeds in that part of the world, Azolla must rank among the world’s
most important economic ferns. It is also used to a lesser extent to supplement the diet of chickens, pigs,
and fish.
Perhaps the best-known use of ferns is the edible fiddleheads of species such as the ostrich fern of the
eastern United States and the bracken fern of Asia. (An early Oregon agricultural bulletin gave recipes on
how to cook bracken, including one on making “creamed bracken.” Subsequent research, however, has
shown that bracken contains carcinogens and should not be eaten.) The foliage of the water fern (Ceratop-
teris) and Diplazium esculentum are eaten in the tropics.
Various ferns with fibrous stems have been used as a substrate to cultivate epiphytic plants. The fibrous
stems of Osmunda and tree ferns have been shredded to make a planting medium for orchids. More recently,
tree-fern trunks have been cut and shaped to make containers for hanging-basket ferns. Because of conser-
vation concerns, however, the export of tree ferns and tree-fern products is restricted. In areas where they
abound, tree ferns have also been used to line roads and to construct small buildings because their trunks
are resistant to rot.
Other uses of ferns include thatching roofs and making tea and dyes. Some fern rhizomes and rachises,
such as those of the Gleicheniaceae and Lygodium, are flexible and tough enough to be used in basketry.
Folklore ascribes many dubious uses to ferns, such as restoring hair, making oneself invisible, and provid-
ing lucky amulets. The use of the male fern to treat worm infections, however, is an example of one such
tale being true. Some ferns, such as the bracken fern, gleichenias, and the hay-scented ferns, are considered
weeds. Ferns have also been used to investigate basic problems in plant biology, such as organ development,
hormones, genetics, evolution, and responses to light. But of all their uses, ferns are by far most important
for their ornamental value.
2 The Structure of Ferns
Like sunflowers, oaks, pines, and other familiar plants in the landscape, the ferns have stems, roots, and
leaves. Although these parts are familiar to most of us, their character and function are sometimes unusual.
Much of this chapter might seem complicated, but you will find that a basic understanding of fern struc-
ture is extremely useful in making use of the rest of this book. Figure 2.1 illustrates the parts of a fern.
Stems
The stems of ferns can become tree-like and conspicuous, or they may be horizontal and go unnoticed.
Inconspicuous stems might be short, partly buried in the soil, or densely covered with leaf bases, hairs, or
scales. If the stem is erect and stout, it may be called a rootstock (Figure 2.2a). If it lies horizontally on the
ground and is anchored to the soil by roots, it is called a rhizome (Figure 2.2b–e). Some rhizomes are long,
freely branched, and climbing or scrambling over soil, rocks, or trees. They can bear fronds close together
or far apart. Some rhizomes may grow into the soil and send out new fronds in unexpected places. Others
branch frequently and form dense clumps. These clumps usually produce fronds in whorls, in irregular clus-
ters, or arranged in two ranks. Erect stems generally produce fronds in a whorl or vase-like cluster. Semi-
erect stems may have less perfect whorls. The slender, long stems on Boston ferns (Nephrolepis exaltata) and
other species are called stolons (Figure 2.2f). These spring from the main stem or rootstock and produce
new plantlets at their tips or from side buds. The stems of tree ferns are so large and wide that they are rightly
called trunks (Figure 2.2g). Some grow up to 18 m (60 ft.) tall. Where abundant, tree ferns have been used
to construct small buildings, and the starch in their stems has been used for food. Some ferns have thick,
hollow stems that house ants. The ants protect the fern and provide some nutrients to the fern in exchange
for shelter.
Stems serve the plant in several ways. First, the stem tip produces new stems, leaves, and roots. If this
tip is damaged, it might kill the fern, particularly if the stem does not branch laterally. Given its importance,
the stem tip is often protected by a covering of hairs or scales (Figure 2.2j, k). Second, the stem supports the
leaves. Leaves can be supported in large clusters at great heights, as in tree ferns, but most ferns have their
leaves supported on short- to long-creeping stems. Whatever the pattern of support, the leaves are always
held up to the light and air. Third, stems conduct water and nutrients between the roots and leaves. For this
purpose they are equipped with strands or bundles of special tissue known as xylem and phloem or, col-
lectively, as vascular or conducting tissue. The vascular tissue can have different patterns depending on the
species or taxonomic group to which the fern belongs (Figure 2.2h, i). Fourth, stems store food and water,
13
costa
costule
segment
rachis costule
segment
Pinnule Detail
rachis
blade
pinna
frond
pinnule
segment
stipe
fiddlehead
scales
rhizome
roots
a d
g
i
h
j
k
Figure 2.2. Kinds of stems: a. erect stem; b. rhizome, short-creeping (Asplenium bulbiferum); c. rhizome, wide-creeping
and branching (Gymnocarpium dryopteris); d. rhizome, wide-creeping and branching from stipe base (Hypolepis
tenuifolia); e. rhizome creeping, branching and fleshy (Phlebodium); f. stolons (Nephrolepis exaltata); g. trunk of a tree
fern; h. vascular or conducting tissue within rhizome (Hypolepis); i. vascular or conducting tissue within rhizome
(Polypodium); j. hairs covering the rhizome of h; k. scales covering the rhizome of i.
16 C 2
particularly if they are thick and fleshy (Figure 2.2e). Some stems, such as stolons and branching rhizomes,
reproduce plants or increase the size of the colony.
Roots
The roots of adult ferns grow from the stem. There is never a main root system consisting of a taproot and
lateral branch-roots. Fern roots are mostly fine, fibrous, densely branched, thick masses; rarely are they
thick and fleshy. They grow close to the soil surface and are easily injured when the soil is tilled or disturbed.
Young, actively growing roots have whitish or yellowish tips. Older portions of the roots are dark brown or
black. The presence of a large number of young roots is a good sign of active growth.
Creeping or reclining rhizomes produce new roots as they grow forward. These roots are usually located
on the underside of the rhizome just behind the tip. Ferns with erect stems produce new roots from the stem
base. Erect-stemmed ferns may have their bases covered with old leaf bases, making it hard for the emerg-
ing roots to establish themselves into the soil (Figure 2.2a). If, however, the old fronds decay and a fine layer
of forest litter accumulates around the stem, emerging roots will find a rooting medium. In cultivation it
may be necessary to remove the old leaf bases of erect stems and replant the fern deeper into the soil to give
the emerging roots a better chance to establish themselves and prolong the life of the fern.
Old tree ferns often have masses of roots on their stems. These roots are called aerial roots and form a
fibrous outer layer around the stem. They absorb water and add strength and rigidity to the stem, which is
necessary to support the heavy crown of fronds. The trunks of some tree ferns—for example, the Hawai-
ian tree fern Cibotium glaucum—readily send out new roots when planted.
The roots of some ferns house fungi in a symbiotic relationship. The fungi (mycorrhizae) help gather
mineral nutrients for the fern in exchange for food. The fungi reside in the cells of the root or the root hairs
that often become swollen and mucilaginous at their tips. The mycorrhizae that have been studied on ferns
are reported to be cup fungi (Ascomycetes) and Glomales (Moteetee et al. 1996). Mycorrhizae of agricultural
crops studied have been found to enhance growth when soil nutrients are low and not when sufficient or
high. Whether these findings apply to ferns is not known.
The primary function of roots is to absorb water and minerals as well as hold the plant in place. Some
ferns have roots with other functions, such as producing new plants or storing food and water.
Leaves
The leaves of ferns are called fronds and are produced at the stem tip. They develop by uncoiling and while
doing so are called fiddleheads or monkey tails (see Figure 2.1). Technically, they are called croziers, from
the term for a shepherd’s crook. Croziers are tender and vulnerable to drying and damage. They are often
protected by a covering of hairs or scales. A few ferns have emerging fronds that are hook-shaped or straight
rather than coiled.
When a crozier uncoils, it produces a frond that typically has two parts: the stipe (petiole) and blade (see
Figure 2.1). The stipe is the stalk, and the thin leafy part it supports is the blade. The stipe commonly
bears hairs or scales that are often important in identification of a fern. In some ferns the stipes may be dark
and highly polished, as in the maidenhair ferns (Adiantum). The stipes may also be long, short, or entirely
absent (sessile). If the stipe naturally breaks or separates from the stem along a predetermined line or joint,
it is said to be jointed or articulate. The tissue found in or near the middle of the stipe is called the vascu-
lar or conducting tissue (stipe bundles), which can be seen if you cut through the stipe with a razor blade.
Many fern genera can be identified by the patterns of this tissue.
T S F 17
Blades come in many shapes and may be undivided, little divided, or much divided into smaller parts.
Names are given to all these shapes, states of division, and their parts. The undivided state is called simple
(Figure 2.3). If the blade is divided to the central vein or midrib, the frond is said to be compound (Figure
2.4). If each resulting leaflet is again divided to its midrib, the frond is twice compound. Tree ferns are often
two- or four-times compound. If the blade is cut so that the leaflets form along the midrib, the frond is said
to be pinnately compound or pinnate. If the blade is cut so the leaflets join at the same point on the stipe,
the frond is said to be palmately compound or palmate. If the lateral leaflets on a palmate frond are cleft into
two or more segments the frond is called pedate. Most ferns are pinnately compound.
a b c d
Figure 2.3. Simple fronds: a. simple and entire; b. bifurcate; c. pinnatifid, with pedate base; d. pinnatifid.
Whether the blade is pinnate, palmate, or pedately compound, the leaflet of the first division is called
a pinna (pinnae is the plural). If the pinna is divided, its divisions are called pinnules or secondary pinnae,
and the frond is two-pinnate (bipinnate). If the secondary pinnae are divided, their divisions are called ter-
tiary pinnae, and the frond is three-pinnate (tripinnate). A general name for these parts, whether primary,
secondary, or tertiary in rank, is leaflet. If a frond is pinnately compound, its midrib appears like a contin-
uation of the stipe and is then called a rachis. The margins of the blade or its leaflets may be variously mod-
ified (Figure 2.5).
Venation patterns differ greatly among ferns (Figure 2.6). Veins may be unforked (simple) or forked.
If forked, they can be branched distinctively. Their ends may be free or joined with others to form meshes,
in which case they are said to be netted (or areolate or anastomosed). Sometimes free veins are enclosed in
a mesh, and the ferns are said to have included veinlets. Some ferns have veins that are deeply immersed in
the blade and hardly visible.
The great variety in frond shape, size, and venation does not affect the ability of the frond to fulfill its
main function—to make food for the plant with the help of sunlight.
Fruiting Bodies
The lower surface of the frond may bear rusty patches or black dots or lines known as sori (sorus is the sin-
gular). These are clusters of spore cases or sporangia. The sori may be round, oblong, linear, or of some other
r
p p
p p
r r r
sp
d
a c
sp
sp p
r p
g
f
tp
sp
e
Figure 2.4. Compound fronds (r = rachis; p = pinna; sp = secondary pinnule; tp = tertiary pinnule): a. pinnatifid above
to pinnate below; b. pinnate; c. pinnate-pinnatifid; d. bipinnate; e. tripinnate; f. pedate, the pinnae pinnate; g. palmate
or radiate, the pinnae pinnate.
T S F 19
d e
a c
b g
f
Figure 2.5. Leaf margins: a. entire; b. undulate; c. crenate; d. serrate; e. lobed; f. pinnatifid; g. incised.
shape (Figure 2.7). Some ferns have loosely arranged sori, not definite clusters. The sporangia may be scat-
tered along the main veins, or they may completely cover the lower surface of the frond. The sori of some
ferns are protected by an indusium, a bit of tissue that, depending on the species, is formed in various ways
from various parts of the blade (Figure 2.8). In some ferns it consists only of the margin of the leaf rolled
over the sori (Figure 2.8c). Such an enrolled leaf margin is called a false indusium. The shape and location
of the sorus and the kind of indusium (if present) are important in identifying ferns.
The spore cases or sporangia are easily seen with a ten-power (10×) magnifying glass, but to see their
finer details requires a microscope. They will be seen as stalked capsules or cases containing spores. If the
sporangia have already shed their spores, they will appear frayed or cracked (Figure 2.9). If the sporangia still
contain spores, the case will be round and intact. In this condition you may be able to see a slightly raised
dark ring of cells around the case. This ring is called the annulus and acts like a catapult to open the case and
hurl out the spores. The annulus may vary in position on the spore case, and in some genera it is only a patch
of slightly thickened cells instead of a well-defined ring (Figure 2.10). In some species all the sporangia in
a sorus open at about the same time, but in most ferns the sporangia open at different times over an
extended period.
b c
a d
Figure 2.6. Veins: a. simple; b. forked with free vein endings; c. netted or areolate; d. netted with included veinlets.
a b
c
d
Figure 2.7. Sori: a. along the veins (Coniogramme); b. round (Macrothelypteris); c. linear and marginal (Pteridium); d.
linear and medial (Athyrium).
b
a c
d
Figure 2.8. Indusia: a. peltate or umbrella-shaped (Polystichum); b. scale-like (Davallia); c. false indusium, formed from
an enrolled leaf margin (Adiantum); d. cup-shaped (Dennstaedtia).
a b c d
Figure 2.10. Annulus positions: a. lateral (Todea); b. apical (Lygodium); c. oblique (Gleichenia); d. vertical
(Polypodium).
A single frond may produce hundreds of thousands of spores. To the unaided eye, spores appear as a
fine dust. Under the microscope, they can be seen to have two basic shapes, depending on the species (Fig-
ure 2.11). Some have bean-shaped spores (monolete), whereas others have more or less pyramidal or slightly
three-angled globose spores (trilete). Both shapes usually have patterns or embellishments on their sur-
faces, and these markings often characterize certain families or genera. The spores of most ferns are brown
or black, but yellow spores are characteristic of the polypods, and green spores are found in some groups.
Green spores are short-lived, usually lasting only one or two weeks. In contrast, nongreen spores usually
remain viable for several years, but with age they germinate more slowly and less dependably.
a b c d e
The life cycle of ferns is complex but of great importance to growers. It is discussed further in Chapter
8 in connection with propagation techniques.
3 Obtaining Ferns
Purchasing Ferns
Before you buy a fern, check its climatic needs to determine whether you can provide it with the proper tem-
perature, shade, and humidity. Also determine whether it will produce the desired landscape effect, espe-
cially given its size when fully grown. Some ferns take up more space than you intended.
Once at the nursery, purchase the fern in a 10 cm (4 in.) to 3.8 liter (1 gal.) size container, unless you
are in a hurry to have a larger plant. Leave the smaller sizes for a time when you have gained more experi-
ence in growing them. Plants larger than the 3.8 liter (1 gal.) size are more expensive, and some may have
passed the stage of rapid growth.
Examine the growing tip. Select a fern that has a firm, large, growing tip. Avoid any that seem discol-
ored, shriveled, or undersized. If the fern is actively growing, see that the tip is forming healthy new fronds.
Avoid ferns with shriveled or deformed growth. If you do not want to repot the fern in the near future, pick
one that has not already overgrown its pot. The growing tip should not be pushing against the edge of the
pot. If you are obtaining a fern that forms clumps of rhizomes, select the one that has the largest number
of healthy rhizome tips.
Nurseries use different soil mixes. Be aware that some mixes require more watering than others. Also,
the smaller the pot and the larger the fern, the more water it will require. If the fern is accustomed to grow-
ing in high humidity at the nursery, it may wilt when you get it home. If you give it just enough humidity
to keep it from wilting, new fronds should adapt to the drier conditions without difficulty. Otherwise, see
the section in Chapter 4 entitled “Air Circulation and Humidity.”
22
O F 23
Collecting Ferns
Certain parts of the world are particularly rich in ferns, and it is tempting for you or your friends to collect
some exotic ferns during travels to these areas. But given the concern about the environment and rare or
endangered species, collecting wild plants is discouraged. Collecting wild plants is forbidden in nearly all
countries unless special collecting permits are obtained. In addition, export permits are also required, and
customs or agricultural inspectors may want to see those permits when you exit a country or re-enter your
country. See Appendix III for more information on permits and restrictions.
If collecting ferns from private property, seek out the owner’s permission. Before collecting wild plants,
be sure to have the proper permits and check that the plants are not on the rare, threatened, or endangered
list. Plants so listed are protected by law (see Appendix III for details). Some plants, such as certain species
of Azolla and Salvinia, are deemed potential pests and are barred from entry into the United States and
within the country cannot be transported without a permit. Even with nonthreatened species, resist the
temptation to collect indiscriminately, wastefully, and without regard for the environment. Thoughtless
collectors have given all collectors a bad reputation. Take only the number of plants that you can grow.
If you live in an area where native ferns abound, you will have access to species suited for the climate
but not necessarily for garden culture. Temperate, deciduous ferns are usually collected in the late fall or
early spring, and other ferns just before the growing season, usually in early or mid-spring. Although ferns
may be collected at any time, their chances of survival are best when they are transplanted just before their
active growing period so that they have a full season to grow and establish themselves. Recently matured
ferns (as opposed to older or long-mature plants) have the best chance of re-establishing themselves. Also,
plants with thick rhizomes generally have a better chance of survival than those with thinner ones. If the fern
is branched or clumped, one piece of the branch or clump is enough to start a new plant. When collecting
a rhizome, take as many of the roots as possible and try not to disturb the soil attached to them. Firm the
soil around the exposed roots of the remaining plants. While transporting the plant, keep the soil and
foliage cool and moist, and replant the fern as soon as possible.
By visiting local nurseries, gardens, and members of plant societies, you might be able to obtain inter-
esting species and establish valuable contacts as well. Above all, keep in mind that spores usually do not
require permits for import and are easier to collect, pack, and mail. Also, spore-grown plants tend to adapt
more readily to cultivation.
Packing Ferns
Water the ferns a day before packing so that excess water has time to drain away. Ferns to be transported
with soil on their roots need only be wrapped in newspaper. If the ferns are in pots, they may be tapped out
of the pots to reduce the weight. When bringing plants into the United States, all soil must be removed from
the plant. Dip the roots into a bucket of water so the soil will fall away, but try to leave intact as many of
the roots as possible. A number of dips and a gentle agitation in the water should get the soil off. Remove
24 C 3
all dead and broken fronds. Double-check to see that all insects are removed. The tip of the stem or rhizome
is the most important part of the fern and must be well protected. If fronds must be removed for packing,
do so, but keep the roots and particularly the rhizome or crown intact. Protect the rhizome and roots from
being crushed by surrounding them with packing material. If packing material other than newspaper is used
and ferns are to be sent into the United States, make certain that the material is on the approved list issued
by the Plant Import Permit Station. If you are going to remote areas, bring your own packaging and wrap-
ping supplies. Stringed tags are useful labels, as are gummed labels that will stick on plastic. Write the
name and place of collection on the labels in pencil or permanent ink.
Some plants can be wrapped for protection in newspaper rolled into a cylinder or a cone and placed in
a plastic bag to retain moisture. Make certain that the plants are not too wet, especially if they are put into
plastic bags without any newspaper to cushion the plant and absorb extra water. Plants that are too wet will
rot readily in plastic bags, especially in warm climates. Air may be blown into plastic bags and sealed to serve
as a cushion. Plants sent to the United States should be in sealed bags. If pests are found in any one open
bag, then all the other open bags in the shipment might be considered infested and would be quarantined
by agricultural inspectors.
Empty plastic water bottles can be made into excellent containers for transporting plants. Cut the bot-
tle in half around the middle. One piece will fit snugly inside the other piece with a slight squeeze when you
wish to close the container.
Cardboard cartons lined with wax paper or plastic can be used to ship ferns. In some areas, woven bas-
kets are easier to obtain and can be used, unless they are made of willow or other wood likely to carry bor-
ing insects. Such baskets are barred from entry into the United States. Secure the individually wrapped ferns
so they will not slip or slide in the box.
Commercial growers have reported that potted, established plants may show stress after 7 to 10 days if
left in the dark, even when other conditions were favorable.
25
26 C 4
The basic needs of plants are not hard to supply, but growing success depends on attending to these
needs with care and exactitude. The remainder of this chapter is devoted to a discussion of these require-
ments, with the exception of mineral needs, which are discussed in Chapter 5.
Temperature
Ferns vary greatly in their temperature requirements. Most grow best at daytime temperatures of 18 to 27°C
(65–80°F). Tropical ferns generally prefer ranges from 21 to 27°C (70–80°F) during the day, and some may
even tolerate similar nighttime temperatures. Most ferns grow best when the nighttime temperature is
about 5°C (10°F) cooler than the daytime temperature. Some ferns need seasonal periods of cooler tem-
peratures to grow well. You cannot predict a fern’s temperature tolerance simply by the looks of the plant.
Knowledge of its native habitat and the temperature tolerances of its relatives will help, but there are some-
times surprising exceptions. When the hardiness of a species is unknown, it is best to use larger individu-
als because they generally withstand temperature extremes better than small plants. Since the minimum
temperature tolerance is not known for many ferns, gardeners can make useful contributions to horticul-
ture by reporting such data from their garden to appropriate fern societies and journals.
Floras, which are books for identifying the plants of a particular region, sometimes provide informa-
tion on determining the temperature tolerance of ferns in a given area. Some relatively recent fern floras
helpful to gardeners have been written for Australia (ABRS/CSIRO 1998; Andrews 1990; Jones and Cleme-
sha 1981), New Zealand (Brownsey and Smith-Dodsworth 1989), South Africa (Burrows 1990; Jacobsen
1983), and Japan (Iwatsuki et al. 1995; Kurata and Nakaike 1964–1997, fully illustrated with maps).
North America and Canada have a flora (Flora of North America Editorial Committee 1993) and there are
many smaller ones. In tropical America, fern floras have been written for the West Indies (Proctor 1977,
1985, 1989), Chihuahua, Mexico (Knobloch and Correll 1962), Oaxaca, Mexico (Mickel and Beitel
1988), Chiapas, Mexico (Smith 1981), Central America (Moran and Riba 1995), Costa Rica and Panama
(Lellinger 1989), the Venezuelan Guayana (Smith 1995), Peru (Tryon and Stolze 1989–1994), and Chile
(Marticorena and Rodríguez 1995). Except for Taiwan (Huang 1994) and Hong Kong (Edie 1978), China
and Korea have not been adequately covered (at least not in English). The ferns of the western Himalayas
have been treated (Khullar 1994) as well as those of Thailand (Tagawa and Iwatsuki 1979–1989), and
western Malaysia (Piggott 1988). Many of the Pacific Islands have floras, but noticeably lacking are floras
for eastern Malaysia, Indonesia, and New Guinea. The British Isles have many recent floras, but those for
Continental Europe in English are not updated.
Ferns may be classified as hardy, semi-hardy, semi-tender, or tender. This classification refers to the
coolest temperature range that a fern can tolerate. It is somewhat arbitrary, since many ferns fall on the bor-
derline between categories or can endure short periods of lower temperatures. The classification, however,
ensures that the temperature preferences of ferns can be more easily known. See the section “Outdoor Ferns”
in Chapter 9 and the individual entries in Chapter 13 for temperature requirements of particular species.
H F
Hardy ferns can be grown outdoors in temperate or colder areas of the United States. They tolerate win-
ter temperatures well below freezing. Also belonging to this group are alpine species that can endure very
cold temperatures for a long time.
Hardy garden species are mostly native to northern and central Europe, North America, and northern
Asia. Many ferns native to the eastern United States are suitable for cultivation. Some adapt to gardens in
warmer climates, but others do not. The cultivation of hardy ferns is treated in several references, includ-
C N 27
ing those by Dyce (1991), Foster (1993), Grounds (1974), Kaye (1968), Mickel (1994), Rush (1984a), and
Rickard (2000).
S- F
Semi-hardy ferns usually tolerate nighttime temperatures above 4°C (40°F) during the cool season.
They survive periods of freezing temperatures that are short and not too severe. These ferns come from
many parts of the world, mostly where warm-temperate climates prevail, such as Japan, Korea, and China.
Semi-hardy ferns grow outdoors in the southeastern United States and warmer northern areas. They are
good choices for inland valleys along warmer coastal regions.
S- F
Semi-tender ferns grow well where nighttime temperatures are mostly above 10°C (50°F) during the
cool season and where freezing temperatures are rare. Frost may burn the foliage or kill the plant, although
some species can survive near-freezing temperatures for a few hours. The semi-tender ferns are native to sub-
tropical and warmer areas with typical daytime temperatures of about 18°C (65°F). Many come from
Latin America, Australia, and New Zealand. Coastal southern California is particularly well suited for
semi-tender ferns, whereas those that tolerate cooler temperatures can be grown inland or farther north
along the coast.
T F
Tender ferns (also known as warm or “stove” species) usually grow poorly when temperatures drop to
16°C (60°F) or below for successive nights. These ferns are mostly native to the lowland tropics where
year-round warmth and humidity prevail. Southern Florida and the Hawaiian lowlands are particularly
favorable places to grow tender ferns outdoors. Here, nighttime temperatures are closer to 21°C (70°F) or
above for most of the year, and daytime temperatures are near 24°C (75°F) or above. In other areas of the
United States, most lowland tropical species must be grown under glass, at least during the cool months,
especially the very tender species that do not like successive nights of temperatures below 21°C (70°F).
Some semi-tender and tender ferns prefer cooler temperatures at night and do not do well in areas
with warm nighttime temperatures. These ferns are usually native to the cooler upland areas of the tropics
and include most of the tree ferns. Conversely, some lowland tropical ferns may not grow in places where
the nighttime temperatures are consistently much lower than the daytime temperatures.
L-T D
Low temperatures (but above freezing) can damage plants. In such instances the plant usually turns yel-
low, produces little growth, and wastes away. But before you dig up a fern to move it to a warmer place, be
sure it is not merely taking its normal rest period. Many ferns take a rest period, especially during the cooler
winter months, when they normally turn yellow and produce little growth. They vigorously renew growth
usually in spring, whereas those that have suffered from low temperatures slowly recover and often have
stunted, undersized rootstocks or rhizomes.
Semi-tender and tender ferns growing in marginally favorable temperatures need to be watered lightly
during the cooler months. At this time growth is slow, and overwatering can often result in root-rot. All
ferns, tender or hardy, are more vulnerable to low temperatures if they are young, weak, recently trans-
planted, or have much new growth.
Various steps can be taken to prevent or reduce damage when temperatures drop, but success will ulti-
mately depend on the fern’s sensitivity to cold and the severity and duration of the low temperatures.
Harden the plants for several weeks or more by giving them less water and exposing them to low but not
injurious temperatures prior to winter weather. Some plants are easier to harden than others. Avoid heavy
pruning or fertilizing late in the season to discourage vulnerable, tender new growth. Keep withered fronds
on the plant to provide some protection through the winter. Transplant ferns early in the growing season
so they will be well established by fall. Protect weak, young, or susceptible plants by moving them to
warmer places such as against buildings, under eaves or trees, or on slopes where the cold air drains away.
Temporary coverings of burlap, straw, leaves, or newspaper will provide some protection during cold spells.
Various foams are available that are nontoxic and work by insulating the plant. Commercial sprays that pro-
tect foliage from frost are also available, but their effectiveness on ferns is unknown.
Once frost or freezing is imminent, increase heat and reduce heat loss from the ground, plant, and sur-
roundings. On clear, still nights the coldest temperatures are likely to come an hour or so before sunrise, and
that is when protective measures may be needed. Shortening the time a plant is exposed to lethal low tem-
peratures from hours to minutes can prevent injury or death. When adding heat with various heaters, be
sure to guard against dehydration, especially from electric heaters in small enclosures. Air movement by
means of fans may bring in warmer air if a low inversion layer is present (that is, a low-lying layer of warm
air above the cold). Inversion layers can be expected when the night is calm and clear but the preceding day
was warm and sunny. This causes the warm air of the day to rise during the night and the heavier colder air
to move downward and settle around the plants.
Fog can be used to protect crops during cold spells with temperatures as low as −2°C (28°F). Farmers
place fog-producing nozzles about 1 m (3 ft.) apart and 2 to 3 m (6.5–10 ft.) above the ground over the aisle
to prevent any dripping water from damaging the plants. The dense fog is confined by a roof of shade
cloth and plastic curtains. The cold temperature is absorbed by the fog, leaving less to damage the plants.
True fog differs from mist in that fog droplets are so small that they do not settle out of the air, whereas mist
droplets are larger and settle out. These larger droplets are less efficient than true fog. Fogging or misting
should be started as soon as temperatures drop below 2°C (35°F), and then maintained continuously or at
regular intervals. It should be continued until the sun has warmed the air above 0°C (32°F). Stopping the
treatment too early may cause the lethal temperature of the plant to be reached.
The formation of frost or ice gives off heat. The layer of frost or ice covering or encasing the plant also
confers some insulation advantage, but the weight of the ice might damage or discolor the plant. Furrow irri-
gation is another means of increasing the temperature, when the water in the furrow freezes, but plants can
become overwatered by this method, and the rise in temperature may be insufficient to protect the plant.
Plants that have been super-cooled (that is, cooled so quickly that damaging ice crystals have not
formed on the plant tissue) should not be moved or disturbed because the movement will cause the sudden
C N 29
formation of ice crystals that, as they grow, pierce and kill the cells of the plant. Super-cooling is apt to occur
when plant surfaces are dry. Allow super-cooled plants to warm slowly.
If plants are frozen, allow them to thaw gradually. This can be done by placing newspaper over the plant
to insulate and shade it from the sun, the warmth of which would otherwise melt the ice crystals within the
plant too rapidly.
Ferns damaged by frost or freezing generally wilt (unless hard tissue is present) and turn black. Less
severely damaged fronds have spots or areas that appear scorched or burned, especially on the more exposed
parts. Damage from desiccation can also occur. For tree ferns and species with woody tissue, particularly
subtropical species, a common practice is to keep the damaged parts on the plant until there are signs of new
growth. Herbaceous ferns that collapse and have completely black fronds may recover if the rhizome is still
firm. Damaged stems or rhizomes often produce distorted, weak growth and will recover slowly, if at all.
Any cold-damaged plants should be watered sparingly because the loss of foliage reduces their water needs.
Assessing the extent of damage may take as long as one growing season on tree ferns. For more details, see
Reasoner (1982).
H-T D
In arid areas, high-temperature damage is difficult to distinguish from damage caused by dry air. Even
with sufficient humidity, high temperatures can scorch or wilt fronds. If the wilted parts do not recover, they
turn brown and brittle. On thicker fronds the scorched areas are usually near the margin or exposed places.
During heat waves, shade the ferns and keep the air humid and the soil moist. In greenhouses, open the
vents and keep the mist nozzle operating. Water sprayed on the roof of the greenhouse will reduce the
temperature appreciably. Never keep plants in a closed car or its trunk on hot days.
C R
Certain temperate and alpine ferns need cold temperatures during the winter to grow well. Without a
cold period these ferns will not renew growth in the spring, or if new growth is produced, it is weak and ulti-
mately wastes away. Growers have established the specific cold requirements of several ferns (see the section
“Forcing Ferns” in Chapter 10). Not all temperate ferns, however, have chilling requirements, and species
that have populations in the southern parts of their ranges may not require cold. Growers in warm climates
should be aware of the difficulties of growing species that require a cold period. Even tropical ferns taken from
higher elevations can prove difficult to grow in hot, lowland tropical climates because of this requirement.
Experiments on Michigan ferns (Hill 1976) showed that different species require different cold peri-
ods, if at all, to renew growth in the spring. Osmunda claytoniana, for example, needs 8 to 12 weeks at 4°C
(40°F) in the dark to achieve 100% bud-break or renewed growth. If the plants are removed from the cold
and dark and placed in a warm, sunny greenhouse (at 20 to 30°C [68–86°F] with 24 hours of light), those
that received the longest cold treatment emerge the fastest (plants with 12 weeks of cold treatment emerged
within 10 days once in the greenhouse). In contrast, bud-break in Adiantum pedatum and Thelypteris palus-
tris was less influenced by the length of the cold treatment, and in Cystopteris fragilis, Onoclea sensibilis,
Deparia acrostichoides, and Diplazium pycnocarpon, no cold treatment was required for bud-break. The lat-
ter species began growth without cold treatment soon after they were brought into the greenhouse. These
experiments explain why it is difficult to grow Osmunda claytoniana in warm climates whereas it is easy to
grow Diplazium pycnocarpon.
30 C 4
Light
H M L
Most ferns grow best in shade or filtered light. The optimal amount of light for most ferns is that pro-
vided by the sun on a dim, overcast day. This measures between 200 and 600 foot-candles. Foot-candles
(f.c.) is a unit for measuring light intensity; another more recent system uses micromole (µmol) units. (See
Appendix I, which describes a method for measuring light intensity with a photographic light meter.)
Adult ferns prefer more light than younger ones, and most spores germinate and grow best in low light.
Contrary to what many people believe, dense shade is unsuitable for most ferns. In dense shade, fronds grow
spindly, are less frequently produced, and tend to yellow and die early.
Ferns growing in low but adequate light are usually large and luxuriant. Under high light, the fronds
become firmer, thicker, more soriferous, and more tolerant of environmental changes. Ferns suffering from
too much light are smaller, less luxuriant, yellowish green, and may have brown margins.
In coastal areas with typically overcast weather, ferns can be planted in full sun. Inland areas may have
brighter days, and the ferns will require more shade. When planting ferns, do not overlook the added light
reflected from walks and buildings. This light can become intense at certain times of the year as the sun’s
angle changes with the seasons.
S F
Trees, laths, painted glass, fiberglass, and plastic cloth all may be used to shade ferns. Plastic cloths made
of polypropylene (sold as Prop-a-Lite, which superseded Saran or Lumite) are much used by the nursery
trade. Different grades give from 30 to 90% shade. In sunny areas such as southern California, 73% shade
cloth (giving 73% shade) has been used with good results. If ferns must be grown in direct sun, they should
be shielded from the sun between noon and 3 .., the brightest and hottest hours of the day.
L Q
Sunlight is composed of a variety of colored lights. These colors are seen separately in a rainbow or
through a glass prism. The red and blue parts of this color spectrum are absorbed by the plant’s pigments
and are used in photosynthesis. Most of the remaining colors in the spectrum are not used by plants and are
passed through the leaf or are reflected. Green light in particular is reflected, which explains why leaves
appear green. Light intensity (discussed earlier under “How Much Light”) should not be confused with light
quality. Light intensity refers to how strong the emitted light is, whereas light quality refers to the kinds of
colored light present. Gardeners who grow plants in sunlight need not be concerned about whether their
plants get enough red and blue light, as they can do little to modify the sunlight’s quality or spectrum. How-
ever, gardeners who grow plants under artificial light to increase the light intensity may also want to know
whether the lamps they are using gives the proper light quality or light spectrum.
A L
Ferns grown indoors, especially in more northerly areas, often suffer from too little light. Supplemen-
tal or full artificial light must be given. Although fluorescent (cool-white) lamps are the usual choice, they
are deficient in red light. Fluorescent lights can be used alone, though they are often used in combination
with incandescent lights (ordinary light bulbs) to supplement the red light. Incandescent lights may also be
used alone, but they produce more heat that can injure plants and are very low in blue light. Gro-Lux
C N 31
lights give a slightly better light quality than cool-white fluorescent but are twice as expensive. High-inten-
sity lights have a good light spectrum and are more energy efficient. They are increasingly easier to install
and more economical. Because they have higher light intensity than other lamps, they can be hung farther
away from the plants. Of the two types of high-intensity lights, the metal halide lamps provide a better spec-
trum for plants and a more pleasing light to the human eye. High-pressure sodium lamps produce a yel-
lowish cast that some people find objectionable. Old lamps may need to be replaced as their intensity
diminishes. Light intensity can be increased by 20 to 50% by walls and benches with reflecting surfaces,
such as by lining them with aluminum foil.
Table 4.1 shows light intensities from two 4-foot fluorescent tubes of 40 watts placed at various heights
above the plants. Light intensity is given at 15 cm (6 in.), 30 cm (12 in.), 45 cm (18 in.), and 60 cm (24 in.)
below the light source. High-intensity lamps will give approximately twice the foot-candle readings shown
in Table 4.1. Use of a reflector will also increase the intensity of light. Most ferns grow best at 200 to 600
foot-candles. Light intensity is measured at the top of the plant. (Also see Appendix I.)
Table 4.1. Light intensity, in foot-candles (f.c.), below two 4-foot, 40-watt cool-white fluorescent tubes at various
distances above the plant material. The light intensity was measured on a horizontal surface with a light meter (Weston
illumination meter model 756).
P R
Little has been reported about the photoperiodic responses of ferns—that is, how they respond to
changes in day length. Such changes can affect growth and the ability to form spores (Patterson and Free-
man 1963). Some ferns seem to become deciduous, resume growth, or produce dimorphic fronds in
response to different day lengths.
32 C 4
Watering
H M W
Most cultivated ferns prefer moist—not soggy—soil. Continuously saturated soil should be avoided as
much as dry soil. A few cultivated ferns might prefer soil that is drier or moister than the average, but these
are exceptions. Ferns should receive less water during cool weather, especially when they are not actively
growing. Chapter 13 gives the water requirements for specific ferns.
H O
There is no precise answer to the question of how often to water ferns. Average conditions require
watering every two or three days, less during cool weather and more during warm. Plants in small pots dry
faster and need to be watered more frequently than those in larger pots. Plants in clay pots need to be
watered more frequently than those in plastic or glazed pots. Because water drains faster through sandy or
coarse soil mixes, plants growing in such soils will need to be watered more frequently than those in silt- or
clay-soil mixes. It is wisest to water a plant when it needs it rather than on a regular time schedule.
H W
Once you have determined that water is needed, thoroughly water the plant with a gentle sprinkle or
flow. Avoid forceful streams because large droplets will compact the soil or wash the soil away from the
roots. Stop when you see water coming through the pot’s drainage hole. If the plant is in the ground, be sure
that the water has reached the soil around the roots. Overhead watering, which wets the foliage, is not rec-
ommended for some ferns, particularly xerophytic species or those with finely divided leaves. These ferns
should be watered at soil level. In areas with high humidity and poor air circulation, you should water
plants in the early part of the day so that the foliage does not remain wet for long. This helps discourage cer-
tain diseases. Scant sprinklings of water will increase the humidity but are seldom sufficient for medium-
sized or larger ferns. Such sprinklings can also cause an accumulation of salts that are injurious to plants.
chances of recovery. Thicker rhizomes contain more stored food, and this fosters new growth to replace the
damaged tissue.
leaching. Salt-encrusted pots should be cleaned or discarded (see the section in Chapter 7 on “Planting in
Pots”). Remember, light sprinklings of water will add salt to the soil, not leach it out.
A M
Ferns need protection from drafts and hot, dry, battering winds. Although some ferns tolerate wind,
their fronds usually look ragged. A small amount of air circulation and ventilation is desirable in greenhouses
or similar enclosures. Air circulation discourages fungi and distributes humidity and temperature evenly.
Small fans, partially open windows, and vents circulate air adequately in the average home greenhouse.
Some outside air flowing into the greenhouse also can be beneficial, as it replenishes the carbon dioxide nec-
essary for plant growth. Incoming air is detrimental, however, when it is extremely dry, hot, cold, smoggy,
or apt to bring in insect pests.
L-H D
Low humidity is one of the most common causes of poor indoor growth. During the winter, the rela-
tive humidity in a typical, centrally heated living room is 40 to 50%—much lower than what most ferns
prefer. Increasing the air temperature worsens the problem because it lowers the relative humidity. Low
humidity can damage ferns in several ways. Emerging fronds tend to shrivel and older fronds yellow more
quickly. Typically, the fronds wilt or the margins become scorched. Barely adequate humidity can result in
a
b
d f
e
Figure 4.1. Increasing the humidity: a. mister or humidifier; b. grouping plants; c. placing pots just above the water
level on pebbles in a tray or pot; d. and e. enclosing plants in plastic frames or glass; f. double potting.
36 C 4
thicker, smaller, harder-textured fronds that may be heavier spore-bearers. An indirect problem of insuffi-
cient humidity is increased infestations of red spider mite.
H-H D
Excessive humidity, especially accompanied by condensation, is harmful for most ferns and should be
controlled, particularly during the night when the temperature may drop quickly. Control can be accom-
plished by watering or misting early in the day or by opening vents to increase air circulation. Excessive
humidity encourages fungal diseases and algae and results in the plants growing soft, weak, and succulent,
making them vulnerable to sudden climatic changes. Under high humidity and warm temperatures, some
ferns with firm, smooth fronds (such as polypodiums) develop brownish-purplish areas, particularly where
the fronds arch or bend. Finely divided fronds, as in some cultivars of Nephrolepis exaltata, might yellow and
shed their leaflets if the humidity is too high.
Soils
There are probably as many soil mixes as there are growers. Experienced growers prepare their own mixes
to best suit their particular style of gardening. If the grower waters frequently, a coarser mix will be used; if
less frequently, a finer mix. If the grower fertilizes regularly, a soil mix without fertilizer will be used. If fer-
tilizers are applied irregularly, the grower will add fertilizer to the mix or use a mix that contains rich gar-
den soil. Whatever mix is used, however, ferns need a growing medium that will hold moisture, provide aer-
ation, support the roots, and provide nutrients or retain added ones.
37
38 C 5
Silty soils hold more water and nutrients than sandy soils, but they might need improved aeration for
good fern growth. Adding organic matter, sand, or perlite will improve the aeration.
Partial or light clay soils need additional coarse and resilient material to improve aeration and to keep
the clay from packing and becoming hard. Add coarse-textured organic matter until it makes up about half
or more of the soil. Perlite or gypsum (calcium sulfate) will also be helpful additives to reduce packing of
the clay. Be cautious about adding sand to certain clays, as binding reactions may occur.
If the soil is primarily dense clay, replace it with potting mix. The replacement is only needed in pock-
ets where the ferns are to be planted. Clay soils are difficult to dilute. Even if an equal amount of organic
matter is added to heavy clay soils, the soil will be suitable for ferns for only about three years. As the added
organic matter gradually decays, clays predominate more and more, and the result is poor aeration and
drainage. These conditions slow plant growth and promote rot, particularly during cool weather. Top
applications of mulch are usually too thin to improve a clay soil, and thicker mulch layers may bury the
ferns. In either case, little organic matter will reach the roots. If the soil has reverted to heavy clay, dig out
the ferns in spring, add organic matter, perlite, or gypsum, and replant. Tree ferns and large ferns are dif-
ficult to replant, so be certain that the soil is suitable before planting in the first place.
S F 39
Clay soils hold more nutrients and water than sandy soils. The main problems with clay soils are their
harsh texture, tendency to expand when wet and shrink when dry, and lack of aeration. Although tropical
ferns grow on clay, they are usually located on slopes with good drainage and in a climate where the clay sel-
dom dries and shrinks.
Coarse granitic soils or decomposed granite formed from the breakdown of granite are unsuitable for
growing ferns. They contain little or no organic matter and the main particles are coarse, resembling crum-
bly gravel. Although granitic soils contain some nutrients, water drains through them so quickly (due to the
coarse particle size) that the roots have little time to absorb the water. The fine roots of ferns will dry rap-
idly in such soil. For most ferns, growing on granitic soil requires incorporating large amounts of water-
holding, humus-forming material such as peat moss and compost.
40 C 5
M
Mulches are loose materials placed over the soil, such as peat moss, leaf mold, and wood or bark prod-
ucts. Mulches serve several functions. They reduce evaporation from the soil, prevent mud from splattering
on the foliage, discourage weeds and some pests, protect exposed roots, and impart a neater appearance to
the garden. On foot paths, medium-sized bark or wood chips reduce soil compaction. Ferns benefit from
mulching, and spring is a good time to do so. The major precaution is to avoid burying the crown of the fern.
S A
Soil additives are incorporated into the soil to improve its quality or suitability for plants. New addi-
tives are constantly being developed. Some are made from recycled products such as rubber or Styrofoam,
and others are old materials that have had their character changed, such as baked clay. Some are products
that are given a new use, such as lichen (reindeer moss). The following are common soil additives used for
growing ferns.
Bark products. Bark can be shredded, ground, or chipped. Bark products decay more slowly than wood by-
products or peat and are used when a well-drained, long-lasting organic material is needed. Composted bark
(partly decayed) is often used in commercial planting mixes. Nursery growers favor using bark products
from fir trees.
Charcoal. Charcoal is added to mixes that might become waterlogged. It absorbs toxic materials produced
when the soil is poorly aerated. There is, however, a limit to what charcoal can do for a soil, and it is best
to avoid waterlogged soils from the start.
Fertilizers. Fertilizers contain minerals needed by the plant. Fish emulsion, blood meal, hooves, and horns
are examples of organic fertilizers often used in soil mixes for ferns. Planting mixes with added organic fer-
tilizers should be used within a week after being prepared. Inorganic fertilizers that can be added to plant-
ing mixes include potassium nitrate, potassium sulfate, superphosphate, dolomite lime, and calcium car-
bonate. Mixes containing inorganic fertilizers can usually be stored for a long time.
Humus. Humus is the dark, resilient, water-holding material that results from the partial decay of plants.
A “humusy” soil is one that contains much partially decayed plant material. Any plant material can be
made into humus. Compost, of course, is very high in humus.
Inorganic matter. Inorganic matter lacks substances derived directly from plants or animals. Inorganic
matter commonly used in planting mixes for ferns includes perlite, vermiculite, sand, and silt.
Leaf mold. Next to peat moss, leaf mold (or partly decayed leaves) is the most popular substance used in soil
mixes for ferns. Quality oak-leaf mold is increasingly difficult to obtain. In California it is illegal to remove
leaf mold from many areas, and if taken from lowland areas it may be infected with Armillaria or oak root-
rot fungus.
S F 41
Lime. See the discussion under “Acidic and Basic Soils” later in this chapter.
Loam. Loam is a soil that has a mixture of different grades of sand, silt, and clay in such proportions that
the characteristics of no one component predominate. An average loam soil contains about 1 part sand, ⅔
part silt, and 1 part clay, although the proportions can vary considerably depending on the grade (size) of
the sand, silt, and clay. Loam is an ideal general garden soil with sufficient drainage, aeration, water-hold-
ing capacity, workability, and nutrient-holding capacity.
Organic matter. Organic matter is anything derived from plants or animals or their remains. It adds nutri-
ents and humus to the soil. Organic matter commonly used in planting mixes for ferns includes peat moss,
leaf mold, sawdust, wood shavings, ground or shredded bark, compost, and manure. Manure and compost
are not favored for commercial planting mixes because of their variable quality and possible contamination
with plant diseases.
Oyster shell. See the discussion under “Acidic and Basic Soils ” later in this chapter.
Peat moss. Peat moss is partly or wholly decomposed moss that provides resilience and aeration to the soil
as well as water-holding capabilities. It may be composed of a mixture of moss species or of individual spe-
cies such as sphagnum moss or hypnum moss. Sedge or black peat does not contain moss and is not rec-
ommended for use with ferns because it might contain excessive salts. Milled peat moss is the common form
used in planting mixes.
Peat moss can be difficult to wet. Try moistening it a day or two before use. It will absorb warm water
faster than cold water, and adding a wetting agent to the water also speeds absorption. Milled peat moss may
need to be mixed and squeezed to ensure an even distribution of moisture. Certain brands of milled peat
moss come in a coarser grade suitable in soil mixes for growing larger ferns and epiphytes. Fine-textured,
milled peat moss breaks down more rapidly but is suitable in mixes used for growing spores or small plants.
Uncut moss consists of intact strands or clumps that have been harvested and dried. It is coarse and
stringy and is often used to line hanging baskets or to plant epiphytes. The best kind comes from various spe-
cies of sphagnum moss. Other uncut moss, usually taken from tree trunks, is a mixture of various moss spe-
cies and has a harsher texture and holds water less evenly. This kind is sometimes sold as green moss and
should not be confused with the live sphagnum moss used to grow insectivorous plants. Uncut moss holds
moisture yet provides excellent aeration if not kept too wet. When used for epiphytes mounted on boards
or tree trunks, pads of uncut moss are placed around the roots and rhizomes. When used in pots, the moss
should not be packed too tightly because it might hold too much water. The purity of commercially avail-
able uncut moss has deteriorated over the years, and nowadays it often contains twigs, grass, and leaves or
is very fragmented. Uncut sphagnum moss from New Zealand is expensive but of excellent quality. Span-
ish moss (Tillandsia usneoides) should not be confused with the uncut sphagnum moss of horticultural use.
Uncut moss that is soaked and drained before use is easier to work. Use gloves when handling wet
uncut moss. Cuts in the skin occasionally become infected by a fungus that lives in the moss. Coconut fiber
and tree-fern trunk fibers are sometimes substituted for uncut moss in lining baskets, but they are stiffer and
retain less water than peat moss.
Perlite. The drainage and aeration of soil can be improved by adding perlite, which is also known by the
brand name Sponge Rok. Perlite is a sterile inorganic material derived by heating and expanding siliceous
rock. It is light, porous, loose, and firm. The particles come in various sizes.
Rockwool (Gordan). Rockwool is a sterile substance made from coal, carbonate, and basalt heated and
blown into fibers. It is 97% air, chemically inert, and does not decompose. Rockwool is used in soilless
mixes. Compressed forms come in blocks, mats, or tubes for special horticultural or floral use. Do not
reuse rockwool that is designed for floral use because it might contain formaldehyde.
42 C 5
Sand. Sand refers to rock particles in a particular size range. The fine grade, commercially designated as
United States Screen Mesh size #30 (approximately 0.5 mm in diameter), is recommended for potting
mixes, although finer sizes to #270 (0.05 mm in diameter) may be used. Coarser grades of size #12, #16,
or #20 (1.6 mm, 1.2 mm, and 0.85 mm in diameter, respectively) are added to soil to increase aeration and
drainage. Size #8 (2.5 mm in diameter) and lower number sizes are designated as gravel. Quartz sand and
washed builder’s sand are suitable for gardening. Do not use sand from ocean beaches or other saline areas
unless it is washed to remove excess salts that might otherwise burn roots.
Sawdust and other wood by-products. Adding wood by-products such as sawdust, shavings, or chips
increases the humus content and improves the soil structure. Wood by-products, however, contain few min-
eral nutrients.
Fresh or partly decayed sawdust (or other wood waste) may be used. Most fresh sawdust and wood
waste contain no appreciable amounts of toxic substances, and those toxins that might be present will
decompose within a few weeks after incorporation into the soil. Reports of toxicity are associated with
finely ground sawdust that rots under conditions of low oxygen, producing toxins in the process. Sawdust
is converted to humus by microorganisms that need air and mineral nutrients to grow. Therefore it is best
to aerate the compost and provide additional minerals for the microorganisms. Add 5.4 kg (12 lbs.) of 10-
6-4 fertilizer for every 45.4 kg (100 lbs.) of sawdust. Other fertilizers rich in nitrogen and phosphorous may
be used. Nitrogen is the most important mineral, and calculations are based on supplying 0.5 kg (1.1 lbs.)
of nitrogen per 45.4 kg (100 lbs.) of sawdust. Apply the minerals at intervals throughout the composting
period. Nitrified sawdust has nitrogen added, sometimes in forms or amounts that can burn fern roots.
Soil. Typically, soil is decomposed, finely divided rock material with a small amount of organic matter.
Sand, silt, and clay (listed from largest to smallest size) are the most important rock particles in garden soil.
Sand provides excellent drainage, aeration, and workability, but retains little water or nutrients. Clay, the
finest textured particle, has poor drainage, poor aeration, and difficult workability, but retains water and
adsorbs and retains nutrients. Silt, being intermediate in size to sand and clay, has essentially intermediate
properties but lacks clay’s chemical ability to attract and retain nutrients.
Vermiculite. Vermiculite is a sterile inorganic material produced by heating bits of mica and causing them
to expand. It comes in various particle sizes and contains some potassium and calcium. Vermiculite adsorbs
minerals and makes them available to plant roots. It also retains water but tends to collapse in a relatively
short time. Wet vermiculite should not be pressed or compacted because its pore structure will be ruined.
Limestone-loving ferns or ferns of basic soil (pH 7–8). Ferns native to basic soils or limestone need more
calcium than other ferns. Chapter 9 includes a list of ferns for such soil. To grow these ferns it might be nec-
essary to add calcium, usually dolomite (CaCO3 · MgCO3), ground limestone (CaCO3), or oyster shells.
Burned or quick lime (CaO) and hydrated or slaked lime (Ca[OH]2) should be used sparingly because small
amounts can greatly increase the pH and harm the plants. Where limestone is available, ferns can be grown
in soil pockets between these rocks, or crushed limestone may be added to the soil. Some growers have had
success using pieces of concrete instead of limestone.
Testing for calcium in the soil is easy. Obtain a teaspoon of a strong acid, such as sulfuric acid or muri-
atic acid (hydrochloric acid or hydrogen chloride). Sulfuric acid may be obtained from a car battery. Muri-
atic acid may be obtained from some toilet bowl cleaners. Place 5 ml (1 tsp.) of the acid on a cup or less of
soil and watch for bubbling. Bubbling indicates the presence of calcium in the form of lime (calcium car-
bonate). If no bubbling occurs, you may not have enough calcium in the soil, or you may have one of the
few soils that has calcium but does not respond to this test, in which case you might have to have the soil
tested commercially.
Acid-loving ferns (pH 4–7). Some acid-loving ferns grow in wet, marshy areas where decaying organic mat-
ter produces carbonic acid. Other acid-loving ferns may grow in well-drained soil that is acidic because its
calcium and basic salts have been leached away. A peaty or highly organic soil will provide acidic conditions
for ferns. Also, soils can be acidified by adding aluminum sulfate, ammonium sulfate, ammonium nitrate,
or finely ground sulfur. The latter is the most economical and efficient additive. Chapter 9 includes a list
of ferns for acid soils.
S S
In the strict sense, soil sterilization means killing all microorganisms, but in general horticulture it
usually refers to destroying disease organisms, insects, and weeds. It will here be referred to as soil “treat-
ment” to distinguish it from true sterilization.
Except for commercial growers, planting mixes for ferns are usually not given soil treatment unless dis-
eases are present or the mix is used for growing fern spores or young ferns. The use of compost, manure,
soils, or leaf mold in soil mixes increases the need for treatment. Soil mixes can be treated using chemicals
or heat (complete soil sterilization is usually done in an autoclave or oven). For information regarding soil
treatments, the commercial grower will find the publication by Matkin and Chandler (1957) particularly
helpful.
Chemical treatments. Chemicals such as methyl bromide and formaldehyde can be used to treat soil but
not, of course, around living plants. However, such chemicals are restricted or no longer available or have
been discontinued for health and environmental reasons. Home growers might find it easier, although not
as effective, to use fungicide drenches such as pentachloronitrobenzene or PCNB (Terraclor), thiram
(Arasan), captan, or quaternary ammonium compounds such as Consan or Physan 20. (See Chapter 11 and
Appendix IV.) These products can be used around live plants.
Steam treatment. Most commercial growers use steam to treat the soil. The steam is applied through var-
ious types of equipment until all parts of the soil have been heated to 71°C (160°F) for 30 minutes. For a
better margin of safety, heat at 82°C (180°F) for 30 minutes. High heat, however, might produce toxic con-
ditions. If this happens, let the soil stand for a few weeks or leach it thoroughly with water.
For small amounts of soil, a homemade steam sterilizer can be devised for treatment. Get a large cook-
ing pot or roasting pan and fill with about 1 cm (0.5 in.) or more of water. Place a rack in the pan to raise
the soil (or to raise pots containing the soil) above the water level. Cover and heat this over the stove or in
44 C 5
the oven at about 120°C (250°F), or high enough to produce steam. Be sure that the steam penetrates all
parts of the soil for at least 30 minutes. This might require placing the oven controls at about 150°C
(300°F) for half an hour if there is much mass to be penetrated. Very moist soil wrapped in aluminum foil
can also be heated in this manner.
Since most plastic pots or boxes cannot be treated with high heat, they have to be disinfected separately
by soaking for 30 minutes in a 10% bleach solution, after which they are rinsed with previously boiled water
and then filled with the treated soil.
Microwave treatment. Treatment in a microwave oven is suitable for small amounts of soil. Add generously
moistened soil to a 2 liter (2 quart) casserole dish, cover, and set the oven on high for 5 to 15 minutes, or
until the steam thoroughly penetrates all the soil. Keep it covered while cooling.
Boiling water treatment. Many growers find that gently pouring boiling water over pots filled with small
amounts of soil and provided with good drainage is a satisfactory treatment, especially if the soil will be used
for growing spores. For a given amount of soil, pour at least twice as much boiling water gently through it.
By placing a clean paper towel over the moist soil before pouring, you can prevent much of the lighter mate-
rial in the mix from floating up to the surface.
Direct boiling treatment. Although it kills soil pathogens, boiling soil in a cooking pot is not recom-
mended. This treatment ruins the soil’s structure, reducing it to a soggy mush that takes a long time to drain
and, upon drying, lacks aeration.
Steam sterilization. Complete sterilization kills all organisms including their spores. Except for scientific
work, such as growing ferns in agar, full sterilization is unnecessary. In steam sterilization, the material to
be sterilized is placed in an autoclave at a pressure of 1 kilogram per square centimeter (15 lbs. per sq. in.)
for 15 minutes or longer (depending on the amount) at a temperature of 121 to 123°C (250–254°F).
Small amounts of soil, pots, dishes, or other items can be sterilized in a home pressure-cooker at 1 kilogram
per square centimeter (15 lbs. per sq. in.) for 20 to 30 minutes.
Dry heat sterilization. In dry heat sterilization, soil is placed in an oven at 160°C (320°F) for two hours or
longer, depending on the amount of soil. This method has the disadvantage of making the soil difficult to
wet; however, it is useful for sterilizing paper envelopes used to store sterilized spores. To sterilize envelopes,
put them in a heavier box before heating them to the oven temperature given above.
Fertilizers
Ferns growing in rich soils are well supplied with mineral nutrients and do not need fertilizers or other sup-
plements. Although most gardeners do not have such fertile soils, they want the best-looking plants possi-
ble. Ferns that receive ample fertilizers or mineral nutrients grow faster and bigger and more luxuriant.
Ferns, however, are extremely sensitive to overfertilizing, and the novice grower, being aware of this fact,
is often hesitant to fertilize. It is therefore important to learn how to select the proper fertilizers and apply
them correctly.
C F
Beginners are advised to select fertilizers that have a reputation for “low burn.” These contain a low per-
centage of highly soluble salts (that is, salts that dissolve quickly in water). Liquid fertilizers with a low-burn
reputation include fish emulsion and those containing urea. Dry fertilizers with low-burn properties include
fish meal, cottonseed meal, castor-bean oil, and activated sewage sludge.
Plants require several elements, but nitrogen is the most important. Trace elements (those needed by
plants in minute amounts) are usually abundant enough in the soil or in fertilizers as impurities. An excep-
tion is calcium. Non-limestone-derived soils in areas of high rainfall often must have calcium added, usu-
ally in the form of dolomite or ground limestone. The amount of calcium in a fertilizer is usually indicated
by percent on the label.
F A
Always follow the manufacturer’s directions for application of a fertilizer. Never give more than the rec-
ommended dosage. If you are uncertain about the fern’s sensitivity to the fertilizer, there is no harm in
reducing the concentration. Some growers regularly use only half the recommended dosage. It is more
beneficial to give smaller amounts over several applications than to give the full dosage all at once. Although
this involves more work, the plant makes better use of the fertilizer.
Fertilizers that must be diluted should be diluted accurately. Dissolve as much of the fertilizer as pos-
sible before application. For watering large numbers of plants, a range of mechanical devices are available.
Great time-savers are those that mix the fertilizer with the water as it flows through the hose. Watering
devices that operate only with a strong stream of water are not desirable, as the force of the water may dam-
age the fronds and wash the soil from the roots.
Powders or granular forms must be sprinkled evenly and thinly over the moist soil surface. Do not mix
these fertilizers into the soil by tilling. Fern roots are fine and fibrous and grow close to the surface. Tilling
will injure the roots. Water thoroughly immediately after applying the fertilizer powder or granules. Any
fertilizer accidentally spilled on the foliage should be washed off immediately. If too much fertilizer has been
accidentally applied to one spot, particularly if near the crown of the fern, remove it or spread it out thinly
to reduce the possibility of burning the fern and encouraging undesirable mold growth. Although organic
fertilizers require microorganisms such as molds to decompose them and release their nutrients, a high
concentration of mold may damage the fern.
F A
The frequency of fertilizer application depends on the manufacturer’s recommendations. On the aver-
age, one application every three weeks is needed for liquid fertilizers and longer intervals for dry fertilizers.
Fertilize infrequently if you do not want the fern to grow too large. Ferns that do not enter any noticeable
rest period but continue to grow actively throughout the year benefit from fertilizers applied at any time.
If you want to discourage new growth prior to winter or for fear of frost damage, do not fertilize late in the
season. Also, do not fertilize ferns during their periods of rest or dormancy because there is little intake of
nutrients at that time.
Where there is heavy watering or frequent rainfall on porous soils, the water will leach the fertilizer.
Ferns growing in such areas should be fertilized more often. On the other hand, soils that do not drain read-
ily retain fertilizer and can be fertilized less frequently. For commercial growers, the recommended nitro-
gen level for maintaining the best fern foliage color is about 150 ppm, which amounts to about 2.5 ml (½
tsp.) of a fertilizer in 3.8 liters (1 gal.) of water if the fertilizer contains a 20% nitrogen content. This should
be applied as needed.
make sure that the soil is well drained. Then water heavily to leach out any remaining fertilizer. Later water-
ing should be sparing so that the soil becomes well aerated. Increase the humidity and remove badly burned
and wilted, dead fronds. Recovery of the plant usually depends on how badly the roots and rhizomes have
been damaged. Young ferns or those with thin, nonfleshy rhizomes are more difficult to save than older ferns
or those with fleshy rhizomes.
Successful fertilization works on the principle that a little soluble fertilizer salt is good but too much is
bad. Damage from overfertilization is caused by the release of too much soluble salt. This causes water to flow
out of the roots and into the soil, thus dehydrating, wilting, and eventually killing the plant. If only some
of the roots are injured, there may be a partial wilting and/or the characteristic leaf scorch or burned appear-
ance. The partial loss of roots or foliage sets the plant back, and it may be stunted for some time. Fertilizer
salts on the leaf can also dehydrate and burn the tissue. (See “Water Quality and Salt Injury,” in Chapter 4.)
Year-round Needs
Where ferns grow throughout the year, it is necessary to water, fertilize, and groom year-round. Elsewhere,
for best growth, fertilize more frequently during the growing seasons and less so or not at all during the cool
months. Watch for slugs and snails as well as other pests and for sudden and damaging changes in the
weather.
Remove old or badly damaged fronds from vigorously growing plants. Cut them off at the base of the
stipe. Removing these fronds maintains a neat appearance and reduces hiding places for pests. If the plant
is growing poorly and has only a few fronds, avoid removing any fronds except those that are totally brown
or yellow. Sometimes only the tip or margin of an otherwise lovely frond is disfigured. Instead of remov-
ing the whole frond, the disfigurement may be trimmed away with scissors and the remaining part cut
into a shape that is close to the natural one.
There are advantages and disadvantages to the grooming practice of cutting off all the tattered, leggy,
misshapen fronds along with the undamaged fronds. The advantages are
1. In dense frond clusters it is easier to cut off all the fronds than to single out only the
damaged ones.
2. In a few weeks, fresh new compact growth will appear.
3. The frond shapes will be symmetrical due to greater space for development.
The disadvantages are
1. Certain species are harmed by having all their fronds removed at the same time.
2. Encouraging vulnerable new growth just before a frost or cool weather may bring about the
demise of non-deciduous ferns.
3. The bare look, even if temporary, is unattractive to some gardeners.
If the advantages outweigh the disadvantages, then be sure that the plant is a vigorous grower that ordi-
narily produces many fronds during the growing season (as are the common species of Adiantum, Pteris, and
Microlepia). Also, make sure that the fronds are removed when plenty of growing season lies ahead. In areas
with short growing seasons, cutting back all the old fronds is best done in early spring before the fiddleheads
have emerged. In areas with longer growing seasons, this may be done later. In areas with year-round grow-
ing conditions, the time to cut back fronds is not critical for most non-deciduous ferns.
Ferns benefit from an occasional fine spray to remove dust, dirt, and lurking insects. Fronds with hairy
surfaces tend to catch debris that does not wash off easily. Usually the debris can be loosened with a soft
brush.
48
T Y F 49
Leaf spots left by salts in the water are harmless if not too numerous. If conspicuous and unattractive,
the spots can be removed by misting them gently with a liberal amount of water to wash off the salts. About
half a tablespoon of mild household detergent added to a quart of water (about 15 grams per liter) helps dis-
solve the salt faster. This treatment must be followed with a thorough water rinse. When applying the
detergent solution and rinse, tilt the pot to prevent the detergent from dripping on the soil. Spots can be
avoided if rain or de-ionized water is used instead of tap water. Remember that liquid fertilizers applied on
the leaves will also produce water spots. Commercial leaf polishes can be used on thick, leathery ferns to
remove spots and to shine the surface, but the shine does not necessarily help the plant.
Winter Care
If you live in a temperate area, your outdoor ferns will be dormant during the winter and require little care
other than checking that their protective coverings or shelters are not disrupted. Outdoor ferns in sub-
tropical climates grow slowly during the winter, and there is little to be done besides watering, removing
withered fronds, and fertilizing once or twice a month (or not at all). Winter is the low point in an outdoor
subtropical fern garden, but if the fronds are taken care of in late summer and fall, they will grace the
garden throughout the winter. Watch for early spring growth and remove old fronds before the emerging
fiddleheads become hopelessly entangled in them. Indoor ferns, greenhouse ferns, and those in tropical gar-
dens require their usual care, although species with a rest period should be watered and fertilized less.
Spring Care
In temperate and subtropical areas, ferns grow vigorously in the spring as new fiddleheads emerge and
uncoil. In temperate areas, the winter shelters can be removed in spring. In all areas, dead and old fronds
should be removed before the new fiddleheads emerge. This minimizes injury to the new fiddleheads and
gives them room to develop. Aphids commonly appear on ferns in early spring, so watch for them as well
as other pests. If the winter rains have washed some of the soil away from the roots, replace this soil or cover
with mulch, particularly the area around the tip of the rhizome where new roots emerge. Resume fertiliz-
ing every two or three weeks if maximum growth is desired. All ferns must be protected from early heat
waves, particularly while new fronds are uncoiling since they are soft and easily dehydrated. Spring is a good
time to divide, transplant, or repot ferns, and this should be done before the fiddleheads uncoil.
Some ferns will shed all or most of their fronds just before putting forth new ones. The squirrel’s-foot
fern (Davallia mariesii var. stenolepis) drops its fronds in late winter or early spring; the Canary Island daval-
lia (D. canariensis) drops its fronds in spring or early summer; and the knight’s polypody (Goniophlebium
subauriculatum ‘Knightiae’) drops its fronds in late spring. These ferns look forlorn with their withered
fronds, but new fronds will soon clothe the basket again. Remove any lingering old fronds to make way for
the new.
Summer Care
Most ferns produce new fronds well into the summer. Their older fronds that turn yellow or brown should
be removed. Fertilization should be continued for maximum growth. Watering and keeping the air humid
will be the major problem in some areas, especially for hanging baskets and pots. Watch for thrips and scale
insects that usually appear at this time. Snails, slugs, and other pests will also be active. In many parts of the
50 C 6
country that have a long growing season, it is still possible to divide, replant, and repot in summer. Trim
the more robust ferns when they threaten to shade the smaller and slower growing species. Be ready to mist
your ferns when the weather gets too hot. Avoid overwatering the soil when increasing the humidity; sprin-
kle the walkways instead of saturating the soil. Fern spores are most frequently formed at this time and in
the fall.
Fall Care
From late summer into fall, fern growth wanes and less water is required. Fertilizers should be reduced or
withheld. Deciduous ferns will yellow and brown, and new growth stops until spring. Leave the old fronds
on the plant to protect them through the winter in areas where frost and freezes occur. In very cold climates,
some ferns need mulches and even screens or frames built over them. It is important to mark or remember
where deciduous species are planted so that they will not be accidentally dug up or injured during winter
garden activities.
In subtropical gardens with a long growing period, the ferns will still need to be groomed and watered.
Where there is danger of frost, do not remove green fronds, even if they are tattered, because removing them
will encourage new growth that could be damaged by frost. Resist the urge to trim until the danger from
frost has passed. Also reduce fertilizers to discourage new growth before winter. As the weather cools, ferns
need less fertilizer and water. Most of the fronds that remain on the plant in late summer and fall will stay
on the plant throughout the winter; however, if temperatures are mild, some ferns will continue to produce
a few smaller fronds in the months ahead. If the temperature drops very much, the fronds will yellow.
7 Planting
Planting Time
Spring is generally the best time to plant, replant, transplant, or propagate ferns. It allows for a long grow-
ing period so that the fern can establish itself before cool weather. Ferns that form only one set of fronds per
year—for instance, some osmundas—are best moved in early spring before the new fronds have appeared,
or in fall after the old ones have died. If they must be moved in leafy condition, preserve as much of the
foliage as possible to ensure good growth the following year. Ferns that produce new fronds throughout a
long growing season can be safely moved while in leaf. Avoid moving ferns with soft, newly formed fronds
because they wilt and damage easily. If a fern must be moved at this time, avoid disturbing the roots and
keep the root ball intact (as is done for all transplants). After replanting, water well and keep the plant
shaded and the air humid.
51
52 C 7
kets where they can root over a broader surface. Nevertheless, they are usually planted to best effect in the
ground where they have ample space to grow and are free to climb up trees or moss-covered posts.
Ferns with closely spaced fronds, erect or semi-erect stems, or short-creeping rhizomes are best for
planting in pots where they can display their foliage patterns. Ferns native to dry or seasonal climates are best
planted in pots or isolated places in the garden where their particular watering needs may be more easily
met. Trough gardens are especially suitable for such purposes (see Chapter 10). Bottle gardens or terrari-
ums are used to grow ferns and selaginellas requiring high humidity.
Planting in Beds
S S
Before planting in the ground, check the site for proper light, wind exposure, and soil conditions.
Anticipate the full size of the fern you are planting. Will it fit your landscape needs? It is easier to consider
these factors now rather than later. If possible, start with hardened plants—that is, ones acclimated to
exposed conditions. Greenhouse plants moved outdoors can wilt, especially if they have soft succulent
growth. These plants should be gradually exposed to drier air over a week or more.
Figure 7.1. Removing plants from pots: a. from a small pot; b. from a large pot.
Positioning and planting the fern. Normally, note the soil level of the fern you are moving and replant it
at the same level (Figure 7.2). If an erect or semi-erect fern has been tipping over, remove the old stipe bases
and bury the rootstock or rhizome deeper in the soil, leaving its crown or growing tip uncovered, not
buried (Figure 7.3). Measure the height of the soil ball. Add or remove soil from the hole (or pot) to bring
the crown to the proper level when set in place. For pots, the level of the soil should be about 1 cm (0.5 in.)
or more below the top edge to provide a catch basin for water (Figure 7.4). All added soil should be gently
firmed in place.
If the soil attached to the roots is poor, gently shake most of it loose before replanting. Remove dead,
old, and inactive roots. Cut old broken fronds and stipe bases at the rhizome. If the fern is badly root-
bound, gently loosen some of the surface roots and spread them out.
For creeping ferns, position the growing tip of the rhizome to face the direction in which you wish it
to grow. If rhizomes creeping over the soil surface refuse to stay in place after planting, anchor them with
pieces of bent wire or small rocks. When planting in pots, place the growing tip as far from the edge as pos-
sible (Figure 7.5). This might place the foliage at the edge of the pot instead of the center, but it will pro-
vide maximum room for future growth.
Once the plant is positioned, fill in the spaces around the roots. Gently press the soil into place (but do
not compact it) as it is added until you reach the desired level. Do not cover the crown or growing point if
it was growing exposed before. Water the plant well with a gentle sprinkle. A strong stream of water will flat-
ten the plants, wash soil from the roots, and compact the soil, destroying its aeration. Some growers rec-
ommend commercial vitamin B1 preparations for stimulating root growth of transplanted ferns, but this
treatment generally has no effect on intact plants. The leaves make enough of this vitamin, and it can be
crown
old and new
soil level remove old
stipe bases crown
spread roots
Figure 7.2. Transplanting an established plant: into new soil
soil ball
top, original planting; bottom, new planting.
side view
main growing tip
side view
stored in the stem or passed to the roots. A temporary shade placed over recently transplanted plants will
help them recover and may reduce frond loss. Slug and snail control may also be in order. Within the first
few days after planting, some of the softer fronds might wilt, and if they do not recover should be removed.
New fronds will soon grow to replace them.
S C
Remove dead and damaged fronds. Rake leaves away from the crowns, particularly if slugs and snails
are a problem. Keep alert about trimming fast-growing plants that shade the slower growing ferns. Promptly
replace soil that has been washed away by rain or watering. Apply mulches if desired (see Chapter 5). Some
beds may need to be reworked every few years. Slower growth and packed soil indicate that replanting is
needed.
Planting in Pots
S P
A well-selected pot will provide the best growing conditions possible and display the fern to its best
advantage. Some growers routinely repot in early spring to provide fresh soil and larger pots. If ferns are
growing well and not root-bound, they do not need to be repotted. A fern collection containing pots of
many shapes and sizes and variable soil mixes will need more attention during watering.
Pot size. In general, the diameter of the pot should be about one-third the height of the fern as measured
from the soil level. Repotting will generally require the next size up. For best appearance and growth, avoid
planting ferns in pots that are too big. Many ferns, especially the maidenhairs (Adiantum), grow poorly
when over-potted because the soil holds too much moisture in proportion to the active roots. The roots can-
not use up enough of the surrounding moisture to keep the soil well aerated.
Clay versus plastic pots. If proper adjustments are made, ferns do equally well in plastic or clay pots.
Remember that plastic or glazed pots retain more moisture than porous clay pots or wooden tubs. Thus,
water the plastic or glazed pots less or use a coarser, quicker-draining soil mix than you would with clay pots.
If the soil in a plastic pot has dried and pulled away from the sides, wet the soil thoroughly so that it
expands and the water will not trickle between the soil and the pot without penetrating to the roots. Unlike
clay pots, plastic ones are not as likely to support algae or become heavily encrusted with salt. Plastic pots also
have the advantage of weighing less, and holes for extra drainage are easily made with a hot ice pick or awl.
Pots without drainage holes. Beginners should use pots with drainage holes and avoid those without. If pots
without holes must be used, however, there are two ways to provide drainage. The first is to place one to two
inches of coarse material at the bottom of the pot before planting. Water the plant with distilled water just
enough to moisten (not wet) the soil—do not overwater. The second method is to find a clay or plastic pot
that has drainage holes and will fit inside the undrained pot. Plant the fern in the pot with the drainage hole
and place it within the undrained one. The fern can be removed as needed for watering, permitted to drain,
and then slipped back into the pot lacking the hole. Some gravel or drainage material may be placed at
the bottom of the hole-less pot to prevent the fern from standing in water that may have accumulated at the
bottom.
Pot shape. Most ferns do well in standard pot shapes, particularly the stout pots known in the trade as fern
or bulb pots. If drainage, aeration, and room for soil are adequate, the shape of the pot should not affect the
P 57
growth of the plant. Also consider the aesthetic aspects of the pot. It should highlight the plant, not detract
from it. Bright colors and bold designs draw attention to the pot, not the plant.
When purchasing pots, look inside them to see that the drainage hole is large enough and is either
flush with the bottom or depressed to allow the water to drain freely. Pots with flat, broad bottoms tend to
drain slowly. Add extra gravel or other drainage material if more or larger holes cannot be made. Some nov-
elty containers have such a small capacity that frequent watering may be necessary to keep the plant from
wilting, and roots may fill the space too soon.
S C
Potted plants need to be watched for drying or excessive water.
Poor drainage. Never let a pot stand in water unless it contains a fern that grows naturally in swamps or
aquatic habitats. If the saucer below a pot accumulates water long after watering, it should be emptied. Pots
can be raised on pebbles above the water level to avoid becoming waterlogged. If your plant wilts without
standing in water, it probably needs repotting.
Poor drainage in small- to medium-sized pots can be detected by tapping the soil ball out of the pot a
day or two after watering. Examine the soil to see whether it is waterlogged. Replace the soil ball, and
examine it again the next day. If the soil is still soggy and the plant is not growing well, water it less fre-
quently or improve the drainage. Poor drainage can also be detected by the presence of soft decaying roots
and a strong odor of decaying vegetation (see “Watering,” in Chapter 4). In flower beds or large contain-
ers it may be necessary to check the drainage by digging a narrow hole into the soil to determine whether
it is waterlogged.
Root-bound ferns. A fern that is too big for its pot and becomes root-bound (filling the pot with roots) will
grow slowly, produce small fronds, and have leaves that rapidly turn yellow. Such plants need frequent
watering. If the rhizome branches or clumps, the center may be inactive or dead while the younger parts may
be growing over each other at the periphery of the pot. Such a plant can be kept growing for a time with
careful watering and fertilizing, but its vigor will gradually decline.
Root-bound ferns should be replanted or divided and replanted. If you want to increase the size of the
plant, cut off its dead parts and replant it into a next larger size pot. If you want to keep the fern in the same
size pot, it can be divided if it has many growing tips, and thin it to size by removing the smaller and weaker
rhizome branches and other dead parts. Keep several large rhizomes with their growing tips intact, and
58 C 7
replant in the same size pot. Root-bound tree ferns and ferns with unbranched rhizomes cannot be divided
and usually must be transplanted into larger pots. If there are many old and inactive roots, they can be
trimmed so that the root ball can be replanted in the same pot.
Salt crust on pots. A whitish salt crust on pots is unsightly and injurious to ferns. Unfortunately, it is hard
to dissolve. If possible, change the pot or soak and wash it. For soaking, add Physan (Consan) to the water
because it facilitates scrubbing. For badly encrusted pots, chip or scrape off as much of the crust as possi-
ble, then soak the pot and flush it with water. Vinegar dissolves some of the salt crust, but avoid getting vine-
gar on the soil or plant. If the pot is so heavily encrusted that it cannot be cleaned, then it should be dis-
carded. Remember that salt build-up on pots can be reduced by watering thoroughly (rather than
sprinkling), or by using rainwater, or by using plastic pots or cans instead of clay pots (see Chapter 4).
Planting in Baskets
Ferns that grow on trees are called epiphytes. They are not parasites but merely use the tree for support.
Because soil is absent from tree trunks and branches, epiphytic ferns secure nutrients and moisture by
sending their roots into the bark or the accumulated debris of leaves, mosses, and lichens. Epiphytic ferns
are adapted to rapid drainage and drier air than terrestrial ferns. Most ferns used in hanging containers are
epiphytes, although some terrestrial ferns also grow well in baskets. Ferns with drooping or cascading
fronds look particularly attractive in hanging baskets.
fronds will obscure the design of a basket. Plastic baskets are inexpensive and usually have large enough
meshes to allow thick rhizomes to emerge. Plastic baskets need less moss than wire baskets to line their inner
surfaces, and their meshes are just small enough that the moss does not fall out easily.
wire basket
soil ball
potting mix
moss lining
1. Line the basket with about 5 cm (2 in.) of moist, coarse or uncut moss (green moss or uncut
sphagnum moss), coarsely shredded tree-fern fiber (hapu), or coconut fiber. Shredded
redwood bark also works but is not commonly sold.
2. Add the potting mix to the basket and place the soil ball about 5 to 8 cm (2–3 in.) below
the rim of the basket (Figure 7.6).
If the fern is an epiphyte, plant it in the cavity with uncut or shredded sphagnum moss or
a very friable potting mix high in organic matter, such as coarse peat, or composted bark.
For terrestrial species, use ordinary potting mix in the cavity (see “Basic Planting
Procedures,” earlier in the chapter).
3. Press the soil firmly in place around the soil ball.
4. Cover the potting mix and the exposed part of the soil ball with about 2.5 cm (1 in.) of
lining material. This will prevent subsequent watering from dislodging the loose soil mix.
Extra pieces of rhizomes may be poked through the sides of the basket into the moss.
Rhizomes that will not stay in place can be anchored with bent wires.
5. Water the newly planted basket with a gentle but thorough spray or soak it for a few
minutes in a tub. Check for large leaks, which should be plugged up with additional moss.
Remove the old basket.
Trim off dead rhizomes and old fronds.
b c
Figure 7.8. Keeping basket ferns moist: a. plant a potted fern in a basket lined with moss; b. place a shallow clay saucer
over the moss lining before planting; c. place a piece of plastic sheet, with drainage holes punched into the plastic, over
the moss lining before planting.
62 C 7
S C
If properly planted, a basket fern will last a few years. If it is an epiphyte, the fern must have good
drainage. Terrestrial ferns planted in baskets have a tendency to dry too quickly, and their fronds are sus-
ceptible to dryness, particularly if hung indoors. These plants should be substituted with more adaptable
plants or rotated with fresh ones. If you keep them in pots, they can be lifted out of the moss-lined baskets
and easily replaced (Figure 7.8). In arid climates, hanging baskets tend to dry quickly and can be difficult
to wet again without soaking. To slow the drying, place a small sheet of plastic over the moss lining at the
time of planting. A shallow clay saucer can also be used, but holes for better drainage are punched more eas-
ily into the plastic. A flowerpot may be sunk at the top of the basket, to one side of the fern, to act as a catch
basin for water. The rim of the basket can be built up to act as a catch basin as well. Keep alert for weak or
rusting wires and replace them immediately. As moss or lining material decays or falls away, replace it by
stuffing fresh lining material into the cavities.
8 Propagation
Ferns can be propagated two basic ways: from pieces of the parent plant or from spores. The first involves
vegetative reproduction and gives rise to offspring genetically identical to the parent. The second method
involves sexual reproduction and gives rise to offspring genetically different from the parent and among
themselves. Before learning to propagate ferns by spores, you need to understand the fern life cycle.
63
64 C 8
e
b c d
a i
k j
p
l
n
o
m
Figure 8.1. Fern life cycle: a. familiar fern plant or sporophyte; b. pinna bearing sori; c. sorus; d. indusium; e.
sporangia; f. spore; g. spore germinating; h. young prothallus or gametophyte; i. underside of mature prothallus or
gametophyte; j. rhizoids; k. antheridium; l. sperm; m. archegonium; n. egg; o. embryo; p. young sporophyte.
Figure 8.3. Proliferous buds: a. root bud and a young plant established from Asplenium auritum roots; b. leaf bud on
the underside of a frond tip of Woodwardia radicans.
66 C 8
artificial light). Pteris cretica produced sporophytes in two and a half months after sowing without artificial
light. In a sowing of spores, the sporophytes of apogamous prothalli tend to appear at the same time,
whereas the sporophytes of sexual prothalli appear at different times. Ferns that develop apogamously also
tend to produce leaves before the roots, whereas ferns going through the sexual life cycle tend to produce
roots before the leaves. The leaves appear just below the notch of the prothallus. Ferns that reproduce
entirely by apogamy (those in which no sexual cycle is known) include such familiar cultivated species as
Cyrtomium falcatum, C. fortunei, Dryopteris cycadina, and Pteris cretica. Some ferns reproduce by apogamous
and sexual means.
A single prothallus can produce more prothalli. New ones may form along the margins of the old ones
(Figure 8.6a). If conditions are favorable and if fertilization does not occur, growth continues for years. If
c
first leaf
prothallus b
first leaf
first leaf
stem tip second leaf
stem tip
root
prothallus root
prothallus
Figure 8.5. Various stages of a fern developing apogamously: a. the first leaf emerging from the prothallus; b. stem tip
and root formation; c. the first leaf and roots expanding, as the second leaf emerges. The new fern is formed directly
from the surface of the prothallus, not from within an archegonium as in sexually reproducing ferns.
P 67
fertilization does occur, the prothallus dies as the sporophyte develops. In filmy ferns, shoestring ferns,
grammitids, and some polypodiums, new prothalli are produced from spindle-shaped or rod-shaped struc-
tures (gemmae) borne on short stalks along the margin of the prothalli (Figure 8.6b).
Figure 8.6. Prothallus proliferation: a. an old prothallus forming eight new prothalli and one on the tip of a tapered
process, as shown on Platycerium quadridichotomum; b. gemmae borne on stalks growing from the edge of the
prothallus in Trichomanes, each gemma grows into a new prothallus.
Vegetative Propagation
Plant parts that reproduce new plants are the rhizome, root, and frond. The branching rhizome is
requently used in vegetative propagation. Ferns without branching rhizomes or buds generally cannot be
vegetatively propagated and must be grown from spore (this is true for most tree ferns). Success in vegeta-
tive propagation is greatest when moist soil and sufficient humidity are maintained with as little watering
as possible.
D
To divide a rhizome, look for the growing tips where new fronds arise. Before making the division, you
can stimulate new side buds to form on a thick rhizome by partly cutting through it and waiting for the side
buds to develop. Before you remove a tip or a clump of tips for dividing, be certain that enough growing
tips are left on the parent plant to keep it alive. Look for a joint or naturally weak spot to make the cut from
the parent plant (Figure 8.7a). The larger the rhizome or clump taken, the better the chances of a success-
ful division. Use a clean, sharp knife to make the cut. With a trowel, dig up the division. Avoid injuring the
growing tips, and keep as much of the soil around the roots of the division as possible. Cut off old or bro-
ken fronds and remove parts of the larger fronds to reduce water loss, particularly if the air is dry. Cut the
bladeless and withered stipes as close to the rhizome as possible and remove any old or broken roots.
Before planting the division, dust its cut ends with a fungicide such as Captan. Select a rooting medium
that is well-drained, such as a mixture of one part peat moss and one part perlite. Epiphytes often root well
68 C 8
b c
Figure 8.7. Dividing and planting rhizome divisions: a. dividing the rhizome (roots not shown) (Polypodium); b.
planting the rhizome at half its thickness (Davallia); c. planting the rhizome diagonally (Davallia).
in moist uncut sphagnum moss. Avoid using soil, manure, compost, or other substances that harbor a lot
of bacteria and fungi. Provide the box, flat, or pot with good drainage.
Replant the division at its former soil level. If its roots were previously exposed, set the division slightly
deeper so that the roots are covered. Firm the medium in place and water well. Keep the plant lightly
shaded for a few days.
Pieces of creeping rhizomes without roots should be secured to the rooting medium to keep them from
drying out. Plant these at half their thickness into the medium (Figure 8.7b); bent wire can be used to
hold them in place. If the rhizome pieces are a few inches long, thrust their cut ends diagonally into the soil
to about one-third their length, leaving the growing tip exposed (Figure 8.7c).
After planting, the divisions still need to be cared for. Keep them in a humid, shaded, warm place until
rooted enough to be transplanted. Guard against fungus gnats and other pests that feed on exposed tissue
or new roots. Above all, avoid overwatering that might foster mold or decay. If this develops, increase ven-
P 69
tilation, stop overhead watering, and apply a fungicide drench or Physan. Apply inorganic fertilizer two to
three weeks after dividing.
Plants can be easily propagated by air-layering if they have rhizomes that are long and tend to grow away
from basket or soil. Wrap damp uncut sphagnum moss around the rhizome of the mother plant. Tie the
moss into place with string or strips of plastic. The moss may also be wrapped with a small sheet of plastic
before tying. When roots have developed in the moss, the rhizome can be separated from the mother plant
and planted.
B
Some ferns produce new plantlets from buds on their fronds or, more rarely, roots. Bud-producing
fronds can be anchored to the soil while still attached to the mother plant, or they may be detached and
planted (Figure 8.8). The benefit of leaving the bud on the mother plant is that it will have an adequate sup-
ply of water and food while taking root. The bud can be anchored to the soil with a bit of soil or clean sand.
If detached from the mother plant, bud-bearing fronds may be planted whole or in pieces. These are
anchored to the soil surface with a planting mix or clean sand, watered, and then covered to provide adequate
humidity. Once the buds have rooted, cut the mother frond to separate the plantlets. Some species root more
readily if the bud is left attached to the mother plant (this is particularly true for Adiantum caudatum).
A detached bud should be planted at half its thickness in a rooting medium. Water gently but thor-
oughly and then cover with a piece of glass or plastic. Keep the air humid, but avoid excess humidity and
overwatering.
Detached buds may also be grown conveniently in clear plastic shoeboxes or refrigerator boxes, prefer-
ably with holes for drainage. Line the bottom of the box with an inch of perlite. Place the planting medium
(usually half sand and half peat) over the perlite, then plant the buds. The lid might need to be opened to
control excessively high humidity.
Plants that can be propagated from detached buds include Asplenium bulbiferum, A. daucifolium, Poly-
stichum setiferum Divisilobum Group, Tectaria gemmifera, Woodwardia orientalis, and W. radicans. For
these species, apply a weak solution of inorganic fertilizer every two to three weeks, and gradually remove
the covering over the planting as the buds grow. Transplant the larger individuals to avoid overcrowding.
If mold develops, increase the air circulation and use a fungicide drench.
Root buds develop on ferns such as Ophioglossum, some Platycerium, Diplazium esculentum, and Asple-
nium auritum. They are usually unnoticed until the new plantlets emerge from the soil. When these plants
are several inches tall they can be separated from the mother plant and transplanted.
Figure 8.8. Planting buds: a. layering or rooting a bud still attached to the mother plant (Adiantum caudatum); b.
rooting a bud attached to a section of a frond, top and side view (Polystichum setiferum, a proliferous variant); c.
rooting detached buds in a pot (Woodwardia orientalis); d. rooting detached buds in a plastic container (Tectaria
gemmifera).
P 71
months. For the lady fern, a portion of the rhizome tissue should be left on the stipe. To prevent mold or
rot, treat the stipe pieces with Physan 20 or a similar fungicide. For more details see Dyce (1985) and
Rickard (1986).
T C
Tissue culture is the process of growing entire plants from bits of tissue by planting them in a growth
medium of nutrients and hormones. Aseptic conditions must be maintained throughout the procedure,
which involves transferring the tissue from one formulation to another at certain stages. Most Boston ferns
and many other species are commercially propagated by tissue culture. The advantage of growing ferns by
tissue culture is that large numbers of genetically uniform, disease-free plants can be produced in a small
area. Ferns for which there is limited vegetative propagating material, or those that do not produce viable
spores, are especially suitable for tissue culture.
Some ferns propagated by tissue culture are more attractive than those grown from spores. They tend
to be fuller and more compact due to greater branching of the rhizome. Shorter production time is not
always achieved through tissue culture—it depends on the species and growing conditions. It may take as
long to grow maidenhair (Adiantum) from tissue culture as it does from spores; however, staghorn ferns
(Platycerium) usually take less time. Ferns that have been divided and replanted in tissue cultures for three
or more generations tend to produce many mutations, most of which are misshapen and unattractive. The
more troublesome problems in tissue culture include obtaining uncontaminated tissue (explants) and get-
ting young plants acclimated to greenhouse conditions.
Tissue culture involves three basic stages. In the first or establishment phase, the explants are disinfec-
ted and planted in a growth-inducing medium. Fern explants are usually taken from leaves or young stems
(or stolons in Boston ferns). In the second stage, called the multiplication phase, the explants are planted
on a medium that induces shoot proliferations. Growth at this stage can be subcultured to increase the num-
ber of plantlets. In the final stage, or pre-transplant stage, the clumps of shoots are divided and transferred
to a medium that stimulates further development, particularly of the roots. Growth at this stage can also be
subcultured to increase the number of plantlets. After the plantlets have developed roots, stems, and leaves,
they are transplanted into a soil mix and placed in a greenhouse. The tubes or bottles containing the vari-
ous stages of growth are usually kept at 27°C (81°F) and given 16 hours of artificial light at 300 foot-can-
dles, although some growers use lower temperatures (to 16°C or 60°F) and lower light (100 foot-candles).
The main ingredients in the various culture media are water, inorganic nutrients, sugar, vitamins, and
plant hormones called auxins (which stimulate root growth) and cytokinins (which stimulate shoot growth).
Ferns grow best on a solid rather than liquid medium, which is why agar or agar substitutes are also added.
The recipes for growth media vary from stage to stage and from one species to another. Most ferns grow well
on only two formulations, with the same formulation used for the first and second stages. Murashige’s for-
mulations for in vitro propagation are satisfactory for a wide variety of plants and can be bought premixed.
For the beginner at tissue culture, it is easiest to buy a tissue-culture kit from a scientific or biological sup-
72 C 8
ply house. Occasionally, plant societies, colleges, and universities offer workshops or classes on tissue cul-
ture, and much literature is available for all levels of interest.
Spore Propagation
Although not difficult, growing ferns from spores takes time and patience, but the fascination and
pleasure derived from this activity is a reward in itself. The space and equipment needed are minimal.
Most ferns grown commercially from spore take about 6 to 10 months until they are large enough to be
planted into 5-cm (2-in.) pots. Other ferns take longer, such as filmy ferns, which take a few years to
develop sporelings. For raising fern allies from spores, see “Fern Allies” in Chapter 10. To grow gameto-
phytes for experimental purposes, see Dyer (1979).
C S
To collect spores you need to find the sporangia (spore cases). These are usually grouped in clusters,
called sori, on the underside of the frond. The sori are mostly 2 to 5 mm (about 0.1–0.2 in.) wide and may
be round, oblong, or linear. Less frequently, the sporangia are scattered over the entire lower surface of the
frond or borne on modified portions of the leaf.
When collecting spores, it is important to recognize whether the sporangia are ripe or have already
shed their spores. Immature sporangia usually appear whitish green. As they mature, they turn light brown
and finally medium or dark shiny brown, or in some species blackish (Figure 8.9). (In some ferns, such as
Polypodium, the mature sporangia are yellowish due to the color of the spores within.) The sporangia appear
dull brown and frayed after the spores are shed. Another clue to the ripeness of spores is the appearance of
the indusium. Green or whitish indusia indicate unripe spores; light brown, yellow, or nearly black indu-
sia indicate mature spores. Shed or shriveled indusia usually indicate that the spores have been shed.
Ripeness of the sporangia may be more accurately determined with a 10× hand lens (or jeweler’s loop).
Plump sporangia without a cracked wall contain spores, whereas sporangia frayed or with cracks or slits have
shed their spores. Shriveled sporangia will not produce viable spores. Pick fronds when most of the spo-
rangia are still plump and shiny.
To gather spores for sowing, place the picked frond in an envelope or over a clean piece of smooth paper
with the sporangia side down. Cover the frond to prevent the shed spores from blowing away or other
spores from contaminating your collection. Usually the spores will be shed on the paper within minutes,
but wait a day or two if you want to collect the maximum number of spores. They will resemble fine dust.
Some fronds produce large quantities of spores whereas others do not. The spores are often brownish but
may also be green, yellow, or black, depending on the species. Also shed are whole or fragmented sporan-
gia, bits of hair, scales, or other tissue. Tree ferns of the Cyathea group tend to shed whole sporangia that
later open to shed the spores. Examine the paper with a hand lens to be certain that spores are present. Store
the spores in an envelope or folded paper packet. For reference, write the name of the species, the source,
and the date on the outside of the packet.
Contamination by other fern spores can be a problem during collection. To reduce the risk of con-
tamination, pick only clean fronds, or if these are not available, brush or rinse away dirt and debris. Col-
lecting the spores on a cool or dim day reduces spore loss and contamination from other spores.
Another way to collect spores is to place leaflets bearing ripe sporangia in a plastic bag to keep them from
drying in transit. Then follow the directions given later in this chapter under “Sterilization of spores within
sporangia.”
P 73
Spore envelopes. The best way to collect and store spores is in folded paper packets (Figure 8.10). These
are easy to make and are leak-proof. The problem with using ordinary envelopes is that the spores tend to
leak out, and if the leaks are covered with cellophane tape, many spores can be lost because they stick to the
tape. Plastic bags seem like an obvious alternative, but they have the disadvantage of the spores sticking to
the bag and being hard to remove. Fertile leaf material dries more quickly in paper packets, averting mold
problems.
2 3
4 5 6
Figure 8.10. Making a spore envelope: 1. Fold thin paper in half lengthwise (along the dotted line). 2. Double fold the
upper open edges. 3. The folded and sealed upper edge. 4. Turn the envelope over (keep folded edge on top) and fold
diagonally along the dotted lines. 5. Tuck the folded right corner under the band as shown. 6. To seal the envelope,
tuck the left corner under the band. Label with name, date, and source. Leaflets or spores are best enclosed at step 1
or 5.
C S
Removal of nonspore material (sporangia, scales, hairs) reduces contamination by algae, bacteria, fungi,
and mosses. The larger nonspore material can be picked out with forceps or brushed away with a fine
brush. If pure or nearly pure spores are desired, separate the nonspore material by slightly tilting the paper
74 C 8
upon which spores were collected and gently tapping it. The spores are left behind as the fluffier nonspore
material tumbles ahead and can be brushed away. The spores can then be stored until use.
Small bottles with a very fine (about 0.05 mm) screened mesh in place of the top can be used to sepa-
rate spore from nonspore material and to evenly sow the spores. The finest tea screen available, lens paper,
and even cheesecloth have been used as satisfactory screening material. Check the material with a hand lens
to be sure that the separation is adequate.
S S
Most hobbyists and commercial growers do not sterilize (disinfect) spores before sowing. Scientific
work, however, requires it. Most methods use bleach diluted with water and a wetting agent such as Tween
20, Aerosol OT, Alconox, or a mild household detergent, which will prevent the spores from sticking
together. The following methods of disinfecting spores are representative of the many kinds used. For
more details, see Ford and Fay (1990).
Sterilization of spores freed from the sporangia. Clean the spores to remove the larger nonspore material.
Place the spores in a 5 to 10% bleach solution and add a fraction of a drop of wetting agent (1 part Tween
20 to 2000 parts bleach solution). Soak the spores for one minute. Collect the spores on filter paper. Rinse
two to four times with sterile water. Sow from the water solution or dry and store until needed. Some spe-
cies are sensitive to bleach and may require a weaker solution and less soaking time. For such species, reduce
the concentration of bleach solution to 2% and soak for about 5 to 10 seconds. Researchers report that soak-
ing green spores in 2% bleach solution for two to four minutes gives satisfactory results.
Sterilization of spores within sporangia. Select leaflets or fronds with ripe but unopened sporangia. Soak
in a 5 to 10% (rarely 20%) bleach solution to which a wetting agent has been added as above. Remove all
air bubbles with a small brush, and keep the leaflets immersed 5 to 10 seconds. Do not rinse. Place in a clean
envelope. (Envelopes can be disinfected by placing them in a box and baking in an oven at 160°C [320°F]
for about two hours. Never heat the spores.) The envelopes containing the disinfected leaflets can be placed
between layers of clean paper to absorb the moisture. With regular changing of the absorbing paper, the
spores should be shed and ready for use or storage in about three days.
V S
The fresher the spores, the greater and faster the germination. Depending on the species, fern spores can
remain viable from 2 days to 130 years. Storing nongreen spores under cool conditions, including refrig-
eration, prolongs their viability. Variation exists in the percent germination from spores taken at different
times from the same plant and from different plants of the same species. Spores collected from vigorously
growing plants usually have the highest germination rate. The spores of some species require dormancy
before germination. The dormancy can range from two weeks to one year. For more information, see Rush
(1984a, b).
Green spores, such as those in Osmunda, Todea, Equisetum, and Grammitis, generally have the short-
est viability, averaging about two months (the shortest recorded is two days, and the longest about one year).
Because of their short viability, green spores should be sown soon after collecting. For Osmunda (and prob-
ably other genera with green spores), viability can be prolonged by refrigeration at 4°C (40°F). Osmunda
spores stored in a freezer at −20°C (−4°F) for 2 to 3 years retained their viability (Rush 1984b).
Nongreen spores remain viable longer than green ones, generally from 1 to 48 years, with the extremes
being a few weeks (Cyathea) to 130 years (Marsilea). See Lloyd and Klekowski (1970) for a discussion of via-
bility and germination times in green versus nongreen spores.
P 75
S M
There are many ways to grow ferns from spores, but only the most common methods are discussed here
(Figure 8.11). Before sowing it is helpful to sterilize or treat the soil. This discourages algae, mosses, bacte-
ria, fungi, and other pests that can crowd or attack the spores and prothalli (see “Sterilizing Soils” in Chap-
ter 5). Because fern spores are easily airborne and may contaminate other sowings, avoid successive sowings
of different species on the same day. Several methods have been developed to control the amount of spores
sown, such as mixing the spores with inert, dry material, spraying them through an atomizer, or delivering
aliquot amounts through a syringe (Basile 1973). For sowing small amounts, a cotton swab dipped in the
spore and tapped over the container gives better control than merely tapping the spores out from an enve-
lope. For more details, see Basile (1973).
Growing a few plants. The following methods are suitable for a hobbyist who wants to raise only a few
plants. Milled sphagnum moss is good for beginners, but any of the soil mixes listed under the commercial
glass
soil level, sow here
clear or
a translucent
lid
nutrient solution
glass level, sow here
d
soil level, sow here
clay pot
lid
agar level,
saucer sow here
e
b
Figure 8.11. Common methods of planting spores: a. in a flat; b. in a pot; c. in a plastic container; d. on a nutrient
solution; e. on an agar plate; f. on an inverted pot.
76 C 8
method can be used as a planting medium. For small sowings, adequate space is any small, clear container
that is wide enough to allow fingers or forceps to easily remove the prothalli or sporelings and is deep
enough to hold 5 cm (2 in.) of soil and provide 2.5 to 5 cm (1–2 in.) of overhead space. The container will
need a cover, and transparent lids are best, but plastic wrap or glass can also be used. If the soil is disinfec-
ted by pouring boiling water through it, drain holes will be necessary in the container. Allow enough over-
head space for growth; avoid filling pots too close to the top. If overhead space for the sporelings becomes
inadequate, an inverted clear-plastic cup placed over the pot will provide more room and retain humidity.
Containers without drain holes may be layered on the bottom with 1.5 to 2.5 cm (0.5–1 in.) of perlite
(Sponge Rok) to drain excess water from the soil above. Because additional watering after sowing is infre-
quently needed, many growers use cups or containers without drain holes or omit the layer of perlite. The
container should have about a 5 cm (2 in.) layer of planting medium.
If microwaveable planting containers are used, they can be disinfected along with the soil mix in a
microwave oven. Before microwaving, generously moisten the soil, gently firm it in place, loosely cover with
a lid or wax paper, and cook for about five minutes on high. Upon removal, the soil mix should be steamed
through and moist. Remoisten with boiled water if necessary. Keep these disinfected containers covered
until sowing time. With plastic containers that distort in heat, the soil will need to be disinfected previously
in heat-resistant containers and then spooned into the plastic container when cool. The containers can be
previously disinfected with a 10% solution of bleach or Physan 20 and rinsed with boiled water.
A simpler procedure, but one with greater risk of contamination, is to lightly pasteurize a soil mix in a
clean, undrained plastic cup. To do this, add just enough boiling water to moisten the soil for planting (plas-
tic cups generally do not distort with only this amount of boiling water). A novel use of the pressed-peat pots
(Jiffy-pot pellets, with mesh removed if present) is to place a pellet in a small (180–240 ml or 6–8 oz.) plas-
tic tumbler and moisten with about 120 ml (4 oz.) of boiling water. When the peat has expanded, loosen
it, gently firm it down, cover the tumbler, and let cool. Sow the spores thinly to avoid spindly, tangled pro-
thalli, and minimize crowding. A cotton swab dipped in the spores and gently tapped over the medium
helps to control the amount of spores sown. Although dense sowings can be thinned, the thinning process
tends to disrupt and damage growth. Therefore thin sowings are best. After sowing, the containers are cov-
ered with clear lids or glass or plastic wrap held in place with a rubber band.
Commercial method. The spore-planting media vary from grower to grower. Most growers use a mixture
of one to two parts peat moss and one part fine sand. Some growers add loamy soil, fine perlite, or other
additives to this mix. Commercial planting mixes formulated for growing seeds can also be used for spores.
Finely milled sphagnum moss used alone provides good results but might not be suitable for all ferns.
Whatever soil medium is used, it should be able to pass through a 3 mm (0.1 in.) mesh. Fine texture is
important because it ensures good contact with the prothalli and facilitates transplanting. Some growers use
a fine-textured mix on top of a coarser mix to promote drainage.
Glass or plastic containers may be used as previously discussed. New wood flats and larger clay pots with
drainage holes are traditionally used because they can be sterilized or disinfected easily.
Commercial growers have devised several ways of sowing spores as evenly as possible over a broad sur-
face. Salt shakers and spray bottles (atomizers) with spores suspended in water have been used. Depending
on the species and the transplanting procedures, some growers sow spores densely so that the growth soon
after germination is a solid green mat, which will need early separation. By sowing a small amount of spores
into each plug (a depression in a special plastic growing tray; see “Growing in Plugs” later in this chapter),
growers save labor in transplanting. After sowing, the containers are covered with clean plate glass. Plate
glass is better than plastic wrap, which will sag when stretched over a wide container and cause condensa-
tion to collect and drip on one spot. The resulting wet spots promote the growth of fungi, bacteria, and
algae.
P 77
Nutrient solution method. Many different types of nutrient solutions are used commercially and are of
interest to amateurs. Some growers use distilled or tap water or dilute solutions of balanced inorganic fer-
tilizers (at one-quarter strength), though the more common nutrient solutions are not diluted (Table 8.1).
Other solutions include Beyerincks (Brown 1920) and Knudson C without the sugar and agar (Knudson
1946). The measures given in Table 8.1 are given in the metric and English systems to avoid the need for
special measuring equipment. However, since fractions of ounces or tablespoons are difficult to measure,
the formulation makes 95 liters (25 gal.) of solution—the surplus can be used as a liquid fertilizer. If smaller
amounts are desired, the measurements given in grams make one liter or approximately one quart of solu-
tion. The ingredients are inexpensive and can be obtained from local chemical supply houses, firms selling
fertilizers, and drugstores. Prepare the solution by adding the salts to the water in the order given. Impuri-
ties in the chemicals and from the water (tap water) will probably provide the trace elements needed. If you
want to add trace elements, however, use the formulation in Table 8.1. If the solution is to be stored for a
long time, add the iron or trace elements just before use.
These solutions will have a pH of about 6 if distilled water is used. A pH of 6 is optimum for germi-
nating most fern spores. In general, the pH range for germinating spores is 5–7. A more acidic solution can
be obtained by omitting calcium compounds. Some ferns of limestone habitats have spores that can ger-
minate at pH 7–9.
The nutrient solution can be sterilized by placing it in a pressure cooker for 15 minutes at 1 kilogram
per square centimeter (15 lbs. per sq. in.) or by boiling it for a few minutes. The solution is poured into pre-
viously sterilized dishes or plastic containers (which may be sterilized by boiling or washing with a 10%
bleach solution and rinsed). The depth of the solution may vary, but 6 mm (0.2 in.) is enough for germi-
nation. Cover the solution and allow it to cool. Then sow the spores and replace the cover. To keep the
spores floating on the solution, avoid moving the dish after sowing. Tilt the lid slightly to prevent con-
densation from dripping on the spores.
Nutrient agar solution. This method provides a solid nutrient medium for growing spores. It has no advan-
tage over other methods, and ferns are reported to grow slower by this method. It is used mainly for scien-
tific research or when the spore material is extremely valuable and sterile techniques are necessary to ensure
survival. Agar, the solidifying substance, can be obtained from scientific supply companies. Less expensive
substitutes are now available.
The medium is made by adding 14 g (approximately ½ oz.) of agar to 1 liter (approximately 1 quart)
of any suitable nutrient solution. (Seven drops of a general-purpose, water-soluble fertilizer to one liter of
water makes a convenient nutrient solution.) Dissolve the agar by heating it in the nutrient solution, stir-
ring occasionally. Then pour the dissolved solution 6 to 9 mm (0.2–0.4 in.) deep into shallow dishes, cover
loosely, and sterilize immediately in a pressure cooker for 15 minutes at 1 kilogram per square centimeter
(15 lbs. per sq. in.) pressure. After the dishes have been removed from the pressure cooker and the agar has
cooled and solidified, they are ready to be planted, preferably with disinfected spores. Before and after
planting, keep the dishes covered to avoid airborne contamination and moisture loss. Ordinarily the agar
will remain moist enough to support growth into the mature prothalli stage. Vaseline petroleum jelly can
be used to seal the edges to prevent drying. If the agar should dry or crack, transfer the growth to a fresh
medium as soon as possible. A nutrient solution originally used to grow orchid seeds and containing sugar
(sucrose) has been reported as good for growing fern spores. It works best at pH 5 (Knudson 1946).
Inverted pot method. The inverted pot method is ideal for exhibiting the growth of prothalli and young
ferns. It is inefficient for growing ferns commercially because scraping prothalli off the pot is tedious and
awkward.
Fill a small, clean, porous clay flower pot with uncut sphagnum or peat moss, then invert it into a
saucer or shallow dish. Sterilize the unit with steam or by pouring boiling water over it, or place the whole
Table 8.1. Nutrient Solutions
B’ S
Ammonium nitrate NH4NO3 Technical 0.5 2 4
Potassium phosphate KH2PO4 Technical 0.2 ¾ 1½
Magnesium sulfate MgSO4 · 7H2O Technical 0.2 ¾ 2
Calcium chloride CaCl2 Technical 0.1 ½ 1
Ferric chloride FeCl3 Technical trace trace trace
K’ S
Potassium phosphate KH2PO4 Technical 0.2 ¾ 1½
Potassium nitrate KNO3 Fertilizer 0.2 ¾ 1½
Calcium nitrate Ca(NO3)2 · 4H2O Fertilizer 0.8 3 7
Magnesium sulfate MgSO4 · 7H2O Technical 0.2 ¾ 2
Ferric phosphate FePO4 Technical trace trace trace
H’ S #2 (after Hoagland and Aron 1950, except iron chelate replaces ferrous tartrate)
Ammonium phosphate NH4H2PO4 Technical 0.12 ½ 2
Potassium nitrate KNO3 Fertilizer 0.62 2½ 5
Calcium nitrate Ca(NO3 )2 · 4H2O Fertilizer 0.62 2½ 6
Magnesium sulfate MgSO4 · 7H2O Technical 0.38 1½ 4
Chelates of iron as Commercial trace or 2 ppm to the nutrient solution
Versene or Sequestrene
Trace elements (after Hoagland and Aron 1950). A stock solution is made by adding the following to 1 liter (1 qt.) of
water. Use 1 ml (12 drops or 0.034 oz) of the stock solution to 1 liter (1 qt.) of any of the nutrient solutions given
above. If tap water is used, the addition of trace elements is generally not necessary.
Boric acid H3BO3 2.86 g
Manganese chloride MnCl2 · 4H2O 1.81 g
Zinc sulfate ZnSO4 · 7H2O 0.22 g
Copper sulfate CuSO4 · 5H2O 0.08 g
Molybdic acid H2MoO4 · H2O 0.02 g
P 79
unit in boiling water. When drained and cooled, sow it with spores and cover with a larger glass or jar.
Replace water in the saucer as needed. There is less likelihood of overwatering by this method, and if the
spores are not too densely sown, nicely formed heart-shaped prothalli will develop, free from bits of plant-
ing medium. A dilute solution of fertilizer applied in the saucer will hasten growth. Young ferns may be
loosened and lifted off the clay pot with forceps or a thin, flat blade and planted in potting mix.
Miscellaneous methods. Other media and methods are used to sow fern spores, such as plaster of Paris
blocks, broken clay pots, moistened bricks, tree-fern-trunk fibers, soil, peat moss, and uncut sphagnum
moss. Milled sphagnum is particularly good. The soil and container should be sterilized or treated with
steam or boiling water as a precautionary measure.
A-S C
After sowing, the planting should be placed in the proper light and temperature. The first sign of
growth is the appearance of a very thin green mat about 14 days after sowing. In the early stages of germi-
nation, the emerging green cells are too small to be seen without magnification. Germination usually starts
in 4 to 14 days but may take as long as a year. Ferns grown for the trade usually form mature prothalli in
several months. Others take more than a year to mature.
Light. Place the sown spores in filtered sunlight of low to medium intensity (150 to 500 foot-candles). If
artificial (cool-white fluorescent) light is used, it may be left on continuously or for 8 to 16 hours per day
as desired. Research indicates different cellular responses with blue, red, and far-red light, but these
responses are unimportant to the general growing of spores. Several weeks in total darkness are required for
spore germination in Botrychium dissectum and Ophioglossum, and they may need mycorrhizal fungi to
develop further unless grown in a nutrient medium.
Temperature. The optimal temperature for spores to germinate and grow is 20 to 28°C (68–86°F). Most
growers keep the temperature near 25°C (77°F) and lower it as the plants mature. Uniform temperatures
reduce problems associated with water condensation.
Watering. Watering is unnecessary after sowing if the medium was thoroughly moistened and the plant-
ing covered. The edges of the containers may be sealed with Vaseline petroleum jelly to retain moisture. If
water must be added, set the pots in a saucer of water. Flats and plastic boxes may be watered with a fine mist
or spray. Distilled or cooled boiled water is best during these early stages. Tap water can be an unwanted
source of algae.
Fertilizers. Two to three weeks after the green mat of germinated spores appears, weak solutions of fertil-
izer (about half the recommended strength) can be applied. Commercial growers differ on the concentra-
tion and timing of these applications. Concentrations vary from 100 to 200 ppm, given at every watering
or every other watering. Although researchers have applied sugar (2.5% sucrose) under sterile conditions
to hasten growth, this is not recommended for ordinary purposes because it encourages mold.
Transplanting. Where only a few plants are desired and the prothalli are healthy, transplanting may be
delayed until the sporelings have developed about three leaves. They are then picked out and transplanted,
usually to 6.5 cm (2.5 in.) pots containing a fine-textured potting mix.
In commercial operations, transplanting starts when the green mat of prothalli have formed, about two
months after sowing. Some growers transplant at an early stage when the mat is thick enough to lift off.
Small pieces of the mat 6 to 12 mm (0.2–0.5 in.) across are lifted off with forceps (or the tip of a spoon if
a liquid medium is used). Waiting longer with a liquid medium can result in tangled growth and make it
difficult to get the prothalli to drop off the spoon during transplanting.
80 C 8
The clumps of prothalli are spaced about 12 mm (0.5 in.) apart and firmed onto a moist, finely screened
potting mix of half peat and sand, or whatever mix is desired. Be certain the clumps are in good contact with
the planting medium, otherwise they will dry out. Keep a spray bottle of distilled or boiled cool water
handy, and use it while transplanting. Although not always done, the planting mix should be sterilized or
disinfected. Some growers treat the soil at 88°C (190°F) for three hours before use, and all subsequent
watering is done with cooled boiled water (until the covers are removed from over the planting). Some
growers use fungicide drenches on the transplants at this stage. After they are in place, the transplants
should be watered with a fine spray, preferably of cooled boiled water. Cover the planting with glass or
plastic.
The second transplant is done when the prothalli have enlarged to about 4 mm (0.2 in.) in diameter for
most species. For commercial ferns this is usually about three to four months after sowing. Clumps of pro-
thalli about 10 mm (0.4 in.) or more wide are firmed onto the soil mix and watered as on the first transplant.
Some commercial growers start the second transplant when sporelings appear, at which point a clump of
prothalli along with one or more sporelings is then transplanted, sometimes directly into cell packs or plug
trays, which are plastic trays with small depressions in which the plants are grown and eventually mar-
keted. Keeping the prothalli in small clumps seems to result in better growth than separating them too
finely.
Growers who prefer to sell larger plants usually do a third transplant. This is done when the ferns are
about 1.5 to 2.5 cm (0.5–1 in.) tall or have three well-developed leaves, which in commercial ferns is about
five to seven months after sowing. Sporelings may be planted in fresh flats of soil or 6.5 cm (2.5 in.) pots,
with the usual attention to firm them into place and protect them from drying.
Potted transplants are initially protected in frames under glass or plastic coverings. Once established,
they must be hardened. Hardening is achieved by lifting the glass or plastic covering more and more each
day to allow the foliage to adapt to drier air and higher light. Eventually a lath covering is put in place. The
hardening process may take three to six weeks. Ferns hardened in 6.5 cm (2.5 in.) pots still need protection
and should not be planted directly in the ground unless sheltered conditions can be provided.
During the third transplant, some growers separate the sporelings into uniform size classes before
replanting them in flats or pots. If replanted into flats, the fourth transplant (about nine months after sow-
ing) moves the sporelings into 6.5 cm (2.5 in.) pots, and they are hardened off and marketed in about a
month or two.
Early transplants are almost always made as clumps of prothalli, with or without attached sporelings.
No attempt is made to separate the clumps into individual prothalli because clumps produce more vigor-
ous sporelings than isolated prothalli. This is probably due to greater opportunities for out-crossing with
nearby prothalli. With apogamous ferns, clumps rather than individual prothalli are transplanted for ease
of handling. Between and during transplants it is essential to maintain high humidity and soil moisture
because the transplants are small and will dry quickly. Always use a fine spray of water. If sporelings are slow
to appear, spray the prothalli with enough water to leave a thin film on them for a few hours. This can be
repeated in a few days. Discoloration may indicate a lack of fertilizer or overwatering. Disease may develop
in spite of all precautions (see “Special Care”).
G P
Home and commercial growers sometimes cultivate ferns entirely in plugs. Plugs are plastic trays with
depressions provided with drain holes. The depressions, or “plugs,” come in different sizes and numbers per
tray. Growing ferns in plugs instead of pots saves labor, planting media, and bench space. It also allows for
holding plants in cooling chambers for a limited time until needed. Cooling chambers hold back growth
to fit marketing schedules. Plug trays designed for the home propagator come with a lower tray to catch
P 81
draining water and a clear plastic cover to maintain high humidity. The plug trays with small-diameter
depressions (25 mm [1 in.] or less) are suited to growing young ferns and even spores.
The commercial planting mix for plugs differs from most soilless planting mixes by its finer texture and
by containing a wetting agent. For the home gardener, a finer textured soilless mix suitable for planting seeds
will do.
Because the soil capacity in the plug is small, problems can develop rapidly. Greater care must be taken
with the amount and timing of watering. The even distribution and penetration of water are important.
Avoid allowing water to remain on the foliage by watering at a time of day that allows the foliage to dry.
Good air circulation is also important. Crowded wet plants and poor air circulation promote disease (espe-
cially Botrytis in cooler temperatures). Drying of the soil stresses plants and makes them more vulnerable to
diseases as Rhizoctonia and Pythium.
Etiolation, or weak spindly growth resulting from low light, is another problem. It can be prevented by
placing the trays under a strong light (but not so strong that it causes the foliage to turn yellow). Transplants
that are already etiolated make poor material for plug (or small pot) plantings. They often remain leggy and
do not achieve the sturdy growth required of commercial plants. Firmer tissue can also be promoted by ade-
quate ventilation and judicious use of fertilizers. Some growers prefer to use fertilizers sparingly. They also
use nitrate fertilizers, instead of ammonium ones, which they claim produce stockier plants.
Like other plants grown under protected conditions, those grown in plugs will need to be gradually
hardened off. Care must be taken in deciding whether to increase or decrease exposure during the process.
The ideal result among commercial growers is to have “toned plugs,” or plants that are compact, sturdy, of
good color, and hardened off enough to take the rigors of transport to the market.
S C
Mold may develop on the soil, prothalli, and young sporophytes in spite of all precautions. If this hap-
pens, take the following steps immediately. Stop overhead watering. Water with cooled boiled water. Check
that water is not dripping from condensation overhead. Remove the mold and at least 12 mm (0.5 in.) of
the plant tissue beyond the infected area. Finally, drench the area with fungicide and repeat as directed on
the product label (see “Fungi and Bacteria” in Chapter 11).
Water molds (Pythium and Phytophthora) usually cause the prothalli to turn dark and watery and sub-
sequently collapse. Gray mold (Botrytis) appears as gray tufts on the prothalli and foliage. It flourishes
under cool-moist conditions and can therefore be combated by increasing the temperature and decreasing
the humidity. A cyanobacterium (Oscillatoria) also forms a blackish to grayish mold-like growth, and the
threads form dense mats and inhibit growth (see “Algae, Mosses, and Liverworts” in Chapter 11). A pink,
nonpathogenic fungus (Pyronema) that is very resistant to steam treatment may grow over the prothalli and
crowd them out. Rhizoctonia appears as well-spaced threads. If persistent mold or alga infection occurs
even after materials and equipment have been sterilized and the proper cultural conditions are employed,
the sporing room may need to be washed down or drenched with a disinfectant such as formaldehyde,
bleach, or quaternary ammonium compounds (Physan 20, Consan). In sporing rooms and growing rooms,
it is important to reduce air movement from the outside because it might carry in fungus spores.
Insects are another problem (see Chapter 11). Fungus gnat larvae can attack prothalli. They can be
killed by sprays, but it is best to keep the adult flies out of the cultures by tightly sealing the containers with
plastic wrap, Vaseline petroleum jelly, or cotton batting placed between the pot and glass covering. Cultures
grown in terrariums are said to be undamaged by fungus gnats if an insectivorous plant, such as sundew
(Drosera), is also placed in the terrarium. Foliar nematodes can also damage prothalli.
The failure of prothalli to produce young ferns (sporophytes) long after sowing (6 to 18 months) can
be due to several causes. The most common cause is insufficient water for the sperm to swim to the egg. If
82 C 8
sporelings develop only near the edges of the container (where water condensation is greatest), this is a
good indication that more water is needed. If misting does not produce sporophytes, fertilization can usu-
ally be achieved by flooding the prothalli with a thin film of water for a few hours and then letting it drain
away. Other reasons for failure are more difficult to determine. Sometimes the sowing is so dense that the
crowded prothalli produce only antheridia, not archegonia, in which case sporelings cannot be produced
because there are no eggs to be fertilized. In other cases, ferns need a basic medium to grow healthy prothalli
(as in some Pellaea species). Some ferns just take a long time to form sporelings (two years is not uncom-
mon). Slow-growing cultures often become contaminated by algae and mosses (see Chapter 11).
Hybridizing Ferns
Ferns are famous (or infamous!) for hybridizing. They hybridize in nature, and many fern hybrids have been
artificially produced in the laboratory. Some genera that hybridize frequently are Asplenium, Dryopteris, and
Thelypteris.
Fern hybrids can be recognized because they are intermediate between their parents. If the parents
greatly differ, the hybrids tend to be irregular and vary widely in size, shape, or number of discrete struc-
tures (pinnae, pinnules, sori, and so forth) between the extremes set by the two parents. Hybrid interme-
diacy can usually be seen in frond shape, vein patterns, and sorus shape. Contrary to expectation, hybrids
made at different times from the same parents are not always identical.
Fern hybrids are not always superior to their parents, but some do display hybrid vigor. For example,
Adiantum ×tracyi has physiological qualities superior to both its parents (A. jordanii and A. pedatum). In cul-
tivation, the hybrid grows more robustly than either parent and, in California, it remains green most of the
year. An unfortunate feature of this hybrid (and many others like it) is that it is sterile and can only be prop-
agated by dividing its rhizome. Only rarely are fern hybrids fertile—that is, capable of producing viable
spores.
The simplest method for hybridizing ferns is sowing spores of the two parents together and hoping that
cross-fertilization will occur (Figure 8.12a). The resulting sporelings have to be grown to a fairly large size
to see whether they have the expected intermediate characteristics of the hybrid. This takes a lot of time and
material.
Another method involves sowing the spores of the parents separately and then transplanting the pro-
thalli close to each other in the hope that a hybrid will develop. With this method you can be certain that
spores of both parents have germinated and produced prothalli. For details about this method, see Rasbach
et al. (1994).
Hybrids can also be produced by planting different parts of two mature prothalli close together (Fig-
ure 8.12b). The notched end of the prothallus bears most of the archegonia that contain the eggs; the
remaining part of the prothallus (the tapered end) bears most of the antheridia that contain the sperm. By
cutting one-third of a prothallus at the notched end of one parent and one-third at the tapered end of the
second parent, and by planting these ends together, it is possible to obtain a hybrid. Any sporophytes
resulting from this method are not necessarily hybrids, however. It is possible that the sporeling resulted
from self-fertilization that occurred before separation, or from a selfing after separation (sometimes
antheridia are still present near the notched end of the prothallus), or from new proliferations of the pro-
thallus. Nevertheless, the chances of producing a hybrid by this method are much better than planting
complete prothalli of two species next to each other (Lovis 1968).
Hybrids can also be produced by concentrating the sperm from one parent in water and then adding pro-
thalli of the other parent (Figure 8.12c). The spores from the species intended to be the male parent are
densely sown to encourage antheridia to form. Prothalli used as the female parent are sown thinly or spaced
apart by transplanting to produce more archegonia. The prothallus cultures are sparingly and carefully
watered by soaking the soil from below, so as to prevent a film of water from developing between the soil and
prothalli—this film of water may cause premature dispersal of the sperm and unwanted self-fertilization. Care
must be taken to prevent condensation on the planting cover from falling on the prothalli. Because antheridia
develop before archegonia, the spores of the intended female parent should be sown four to six weeks earlier
so that both parents will be at the proper stage of development simultaneously. Because it is difficult to
know when the parents are at the optimum stage of development for hybridization, it is useful to have sev-
eral cultures of both parents sown on different dates. Prothalli with antheridia suitable for hybridization
usually develop three to four months after sowing, whereas archegonia will develop one to two months after
a
Cut the tapered end off the Cut the notched end off the
intended male parent. intended female parent.
Figure 8.12. Various methods of hybridizing ferns: a. planting spores or young prothalli of intended parents together;
b. exchanging notched and tapered ends of intended parents; c. growing parents separately, mixing them, and then
growing potentially fertilized prothalli separately.
c
the antheridia are ready. Because unwanted self-fertilization may occur, the archegonia must be examined for
signs of sporophyte development before use. This is best done under a dissecting microscope.
The hybridization technique consists of placing about 1.5 square centimeters (¼ sq. in.) of the small
young prothalli bearing the antheridia in a dish (watch glass) with a small amount of warm water, about 25
to 35°C (77–95°F), and teasing the antheridia apart. The dish is then placed 30 cm (12 in.) beneath a 40-
watt light bulb or in sunlight. At 20-minute intervals, the water is checked under a microscope for active
sperm. Once active sperm have been found, 8 to 12 prothalli of the female parent, which have been cleaned
of debris and examined for the absence of sporophytes, are immediately placed, archegonia-side down, on
the water containing the sperm. The dish is covered with glass to prevent drying and returned to the light.
It is left undisturbed for a half-hour to overnight. At the end of this time the female prothalli are removed,
washed several times in fresh water to remove any of the small male prothalli, and planted in fresh soil.
Hybrid sporophytes usually appear four to six weeks after fertilization. Any sporophytes that appear
before this time generally are the result of self-fertilization. A success rate of about 80% has been reported
with this technique in producing hybrids between closely related and compatible strains. Fern sperm is
known to be attracted to malic acid (0.01%) and citric acid, and this may be of use in hybridization work.
Generally, only closely related ferns hybridize. Other abnormal plants are sometimes confused with
hybrids. Abnormal fronds are caused by injury, mutation, or unusual gene recombinations. If a suspected
hybrid can be duplicated by artificial hybridization in the laboratory, then it probably is a true hybrid.
Indirect proof can be found by comparing a suspected hybrid with its parents and looking for intermedi-
ate features. In all cases, keep records of the parents and the methods used in hybridizing because this infor-
mation is scientifically important.
9 Landscaping
When landscaping with ferns, as with any plants, be sure that they receive enough warmth, sunlight,
humidity, protection from wind, and proper soil conditions. Keep in mind their size when fully grown and
any special problems that may arise in their upkeep. Attention to these considerations early on avoids dis-
appointment later.
The planting design you have in mind should be visualized in relationship to its surroundings. Will the
fern harmonize with its surrounding environment? Will it eventually be too shaded or crowded by sur-
rounding trees and shrubs? Are there too many elements in the design that create a sense of “busyness” or
confusion? Sometimes plants that differ greatly provide a refreshing sense of contrast; however, they can be
jarring or disturbing if carelessly selected. The placement of the plants in relationship to the surroundings
should provide a restful sense of balance. If unbalanced, the design can be disturbing to the eye’s sense of
equilibrium. These considerations are basic landscape design principles which when correctly applied pro-
vide the most pleasing aesthetic arrangement to your garden.
C I
Plants used to create center points of interest are called accent plants. The most effective accent plants
have either strong, bold, well-defined patterns to their foliage or, at the other extreme, are soft, fine, and deli-
cate. Accent ferns should not be too small or be crowded among other plants. The general outline of the
plant should have an interesting pattern rather than be just an uninteresting blob of pretty foliage.
87
88 C 9
Figure 9.6. A hanging basket fern offers foliage above eye level.
G C
Where conditions are shady or semi-shady, ferns do well as a ground cover. In such uses, ferns also help
prevent erosion.
O F
The 10 ferns listed here for each section of the United States were selected because they are easy to grow,
attractive, and readily obtainable. Native ferns are favored on these lists. Most parts of the United States
require hardy to very hardy ferns. For additional outdoor species and their temperature tolerances, see
Chapter 13.
Northeastern United States. Hardy to very hardy ferns are the only ones suitable for the northeastern
United States, where prolonged periods of cold are common. This area includes mostly Zones 3–6. Sum-
mers are hot and humid in some areas and often difficult for ferns that prefer cooler summers.
Adiantum pedatum Matteuccia struthiopteris
Deparia acrostichoides Onoclea sensibilis
Dryopteris erythrosora Osmunda claytoniana
Dryopteris goldiana Osmunda regalis
Dryopteris marginalis Polystichum acrostichoides
Southeastern United States. Depending on the locality, ferns for the southeastern United States may be
hardy to semi-tender. This area includes mostly Zones 6–8. For the cooler areas of the Southeast, see the
listing for “Northeastern United States,” and for milder areas see “Southern Florida and Hawaii” and
“Coastal Central and Southern California.” Ferns adapted to hot, humid summers thrive in this area.
Southern Florida and Hawaii. This area includes mostly Zones 10 and 11. Tropical species are favored
here, but a wide range of ferns are adapted to the climate of southern Florida and Hawaii, except those ferns
that like cool summers or need cold temperatures to break dormancy. The occasional cold spells may
require moving the more tender species to protected areas.
Adiantum trapeziforme Microsorum punctatum
Aglaomorpha coronans Nephrolepis falcata ‘Furcans’
Asplenium nidus and relatives Platycerium bifurcatum and cultivars
Davallia, many species Selaginella, many tropical species
Microsorum grossum Sphaeropteris cooperi
Central United States. This area includes mostly Zones 3–6. In the northern part of this area, only very
hardy ferns are suitable; see “Northeastern United States.” For the southern area, see the listings for south-
eastern and southwestern states.
Northwestern United States. This area includes mostly Zones 5–8, with a few pockets of Zone 9. Some
semi-hardy ferns will grow in coastal areas, but the hardy species listed here will be more reliable on coastal
and inland areas. Coastal areas have cooler summers that may not favor species adapted to hot summers.
92 C 9
S-T F
The following ferns will grow in areas that are shadier than usual, but they still must have a certain
amount of light. Provide more light in climates where the weather is frequently overcast. Most of these ferns
will grow in brighter light, but they may be less luxuriant.
Adiantum aleuticum Dryopteris goldiana
Adiantum capillus-veneris Dryopteris marginalis
Adiantum pedatum Gymnocarpium dryopteris
Adiantum raddianum Nephrolepis cordifolia
Arachniodes standishii Osmunda cinnamomea
Asplenium bulbiferum Phegopteris connectilis
Athyrium filix-femina Phyllitis scolopendrium
Blechnum spicant Polystichum acrostichoides
Dennstaedtia punctilobula Polystichum munitum
Dryopteris dilatata Polystichum setiferum
Dryopteris filix-mas
S-T F
Ferns seldom look their best when growing in direct sun (if they grow at all in such intense light). The
following ferns appear reasonably attractive if grown in places with direct morning or late-afternoon sun.
Some tolerate full sun if the skies are often overcast. Elsewhere they need filtered light during the hottest part
L 93
of the day. Those species marked with an asterisk (*) will tolerate full sun only if soil moisture and humid-
ity are adequate.
Aglaomorpha coronans Osmunda, all species*
Athyrium filix-femina Pellaea, most species
Blechnum appendiculatum Phlebodium pseudoaureum
Cheilanthes, most species Pityrogramma, most species
Cibotium glaucum* Platycerium veitchii
Cyrtomium falcatum Polystichum polyblepharum
Dennstaedtia punctilobula* Pteridium aquilinum*
Doodia media Pteris cretica
Dryopteris erythrosora Pteris tremula
Dryopteris ludoviciana Rumohra adiantiformis
Lygodium japonicum Sphaeropteris cooperi
Microlepia platyphylla Thelypteris noveboracensis
Microlepia strigosa Thelypteris puberula
Nephrolepis cordifolia Todea barbara
Onoclea sensibilis*
A F
Aquatic ferns are those that have their foliage submerged (or partly so) or floating on the surface. Such
ferns vary considerably in form compared to bog and wet-soil ferns. Some aquatic ferns can be purchased
from aquarium supply stores. For some species, water that is high in salts may stunt growth or may burn
or blotch the leaves. Changing the old water completely rather than adding more helps reduce salts and
increase aeration. Using a high proportion of sand and adding charcoal to the rooting medium will help
keep the water free of noxious chemicals and odors.
Acrostichum aureum Marsilea, all species
Azolla, all species Microsorum pteropus
Bolbitis heudelotii Salvinia, all species
Ceratopteris, all species
is used seasonally, deciduous species are more frequently cultivated, although some hardy evergreens might
be welcomed. Where seasonal changes are eagerly anticipated, the deciduous species are symbols of spring
as their crowns of beautiful fresh green fiddleheads emerge after a long winter.
One reason it is important to know whether a fern is deciduous is so that you do not think it is dying
when it goes into its rest period. Growing hardy, deciduous ferns in warm climates requires selecting adapt-
able species (sometimes a matter of trial and error), not giving them too much water in the winter (they will
rot), and remembering not to accidentally dig them up when working in the garden. Young plants of some
deciduous species can retain their greenness during their first one or two years of life.
A fern that is deciduous will have all its leaves decline at about the same time. In contrast, an evergreen
fern has leaves that do not decline all at the same time. Contrary to popular belief, evergreen plants do not
have leaves that stay green forever—the older leaves are eventually shed. Whether plants are deciduous or
evergreen, their declining leaves may be self-shedding (articulate) or they may wither in place (marces-
cent). These characteristics affect their landscape use and maintenance.
Deciduous ferns with self-shedding fronds have a special layer (the abscission layer) near the base of the
stipe that allows a clean break at the joint with the stem. Self-shedding fronds require mostly seasonal rak-
ing for maintenance.
Deciduous ferns with fronds that wither in place are more troublesome to groom because their fronds
will need to be cut from the plant to maintain neatness. Because withered, persistent fronds protect the
crown from winter cold, they are usually removed in spring before the new fronds have emerged. For neat-
ness, year-round grooming may be needed to remove any off-season, withered leaves.
Fronds of evergreen ferns generally live a few months to a few years, depending on the species and cul-
tural conditions. Spent fronds of evergreen ferns may be self-shedding (as in Polypodium) or marcescent (as
in Polystichum). Expect a light litter or light grooming problems year-round with most evergreen ferns. For
more details on grooming, see Chapter 6.
The distinction between being evergreen and deciduous is not always clear. Some ferns lose all their
leaves in cold climates but do so only partly in warmer climates. Reclined, marcescent fronds can still con-
tain functional conducting tissue and may stay green and photosynthetic for weeks. Other ferns gradually
reduce growth, and in the absence of new growth, the existing fronds become increasingly tattered but
stay erect. In certain climates the period of dormancy in some ferns is so short that the shedding of the old
fronds is simultaneous with the flush of new fronds, which makes it seem as if the plant is evergreen. Some
species have a rhythm of leaf loss that does not coincide with Northern Hemisphere winters. Many tropi-
cal species grow slowly during the cool months and resume active growth as soon as warm weather returns.
Given these intermediate conditions, discrepancies abound in the literature on the subject. To be clear,
deciduousness or evergreenness should be stated relative to a given climate.
Deciduous ferns. The following ferns are commonly available and obligately deciduous. Their fronds are
deciduous even in warm climates, although their dormancy period may be shorter. Those listed here are
marcescent.
Athyrium filix-femina Gymnocarpium dryopteris
Athyrium niponicum Matteuccia struthiopteris
Athyrium otophorum Onoclea sensibilis
Deparia acrostichoides Osmunda regalis
Diplazium pycnocarpon Phegopteris connectilis
Evergreen ferns. The following common ferns are evergreen even in cold-temperate climates, although their
new growth may be delayed until spring. Those listed here are marcescent.
Blechnum penna-marina Cyrtomium falcatum
Blechnum spicant Dryopteris erythrosora
L 97
A F
Accent ferns have distinctive lines, shapes, textures, or patterns. They can be used as center points of
interest in landscaping.
Aglaomorpha coronans Matteuccia struthiopteris
Asplenium bulbiferum Microsorum punctatum ‘Grandiceps’
Asplenium nidus Nephrolepis exaltata and cultivars
Blechnum brasiliense Osmunda, all species
Blechnum gibbum Phlebodium aureum and cultivars
Cyrtomium falcatum Platycerium, all species
Didymochlaena truncatula Tree ferns, all species
B F
Ferns in hanging baskets or containers are commonly used in today’s house and garden decor. Both epi-
phytes and terrestrial ferns can be used in hanging displays. Species with long fronds that droop over the
98 C 9
edges of the container are particularly attractive. General directions on the selection, planting, and care of
basket ferns are given in Chapter 7, and details on growing specific groups of ferns are provided in Chap-
ter 10. Some of the more popular basket ferns are listed here.
Adiantum capillus-veneris Phlebodium aureum
Adiantum raddianum Phlebodium pseudoaureum
Aglaomorpha, all species Platycerium, all species
Campyloneurum, most species Polypodium, most species
Davallia, all species Pyrrosia lingua
Goniophlebium, all species Rumohra adiantiformis
Nephrolepis, all species
House Ferns
Ferns add grace to a room, and if the room is not too dry and has adequate light, many species can serve as
permanent decorations. Light and humidity often can be improved, but if not, it is best to use several ferns
in rotation, replacing a plant with a fresh one when it shows poor growth. Plants removed from display will
recover if given sufficient light, humidity, and care. Since the kitchen and bathroom are often the most
humid rooms in the house, they are good places to grow ferns if the light is adequate.
Generally, robust ferns with leathery fronds, such as the house holly fern (Cyrtomium falcatum) and
leather fern (Rumohra adiantiformis), grow well indoors. Most davallias are also good as house ferns, and
some are attractive for their finely cut foliage (Davallia fejeensis and D. mariesii var. stenolepis). The most fre-
quently used indoor ferns are the Boston ferns (Nephrolepis exaltata and its cultivars), which come in a
variety of textures.
If possible, start with nearly mature ferns that are acclimated to drier conditions. These “hardened”
plants will grow better in the drier air of a house than those recently removed from a humid greenhouse.
Avoid selecting plants that appear too soft and green.
Frequency of watering depends on the relative humidity of the room, the size of the fern, the type of
soil, and the kind of pot. Most indoor ferns require a thorough watering two to three times a week, more
if the air is dry and less if cool. Water the pot until the water runs out of the drain hole in order to wash out
the salts that accumulate in the soil. Do not permit pots to sit continually in saucers of water. If you are
watering pots by setting them in saucers, remove the pots as soon as the soil has become moistened through-
out. If the pots lack drain holes, use distilled water or rainwater to avoid salt accumulation, but be careful
not to overwater. Be sure that a layer of coarse drainage material such as gravel, pot shards, or perlite is at
the bottom of pots lacking drain holes. Charcoal added to the soil absorbs noxious chemicals produced by
microorganisms that thrive in the wetness. It is better, however, to avoid overwatering than to rely on the
charcoal’s absorptive power. Always remember that the soil should feel moist, not soggy.
Every two weeks or so, apply a fine spray of water to the foliage to wash off dust and insects. Watch
for scale insects, mealybugs, and aphids. These pests can spread quickly because they have few natural ene-
mies indoors. Furthermore, they are not washed off the plant by rain or overhead watering, as often hap-
pens outdoors.
The main challenge in growing ferns indoors is to provide enough humidity and light. Otherwise the
culture of indoor ferns is similar to that of ferns used elsewhere. The following are some favorite house ferns.
99
100 C 10
Trough Gardens
Trough gardens can be thought of as large, deep-dish gardens. Stone troughs used by farmers to water ani-
mals became much prized as planters for miniature plants. Trough and similar broad containers are ideal
for displaying small ferns, confining species with long-creeping rhizomes, and serving as garden ornaments.
Handsome troughs can be made from easily obtainable materials. Hypertufa troughs are made from one
part cement, one part coarse sand, and two parts shredded peat moss. These ingredients are mixed with
water to the consistency of creamed cottage cheese and poured into a mold to form 5-cm (2 in.) thick
walls. The mold can be fashioned from two cardboard cartons and reinforced with wire mesh. After two
days, the mold is peeled away and the cast allowed to cure for seven days. Any plant toxins in the cement
are neutralized by filling the trough with water and adding about 2.5 g (½ tsp.) of dissolved potassium per-
manganate. After two to three hours, the solution is discarded and the trough is rinsed out. Drainage holes
are drilled into the trough. Troughs are allowed to cure for another two or three days. See MacPhail (1990)
for details on reinforcing thin (less than 5 cm [2 in.] thick) troughs with hardware cloth, creating textural
refinements, using other materials, and readying the trough for final planting. Rock garden magazines
often list other ingredients and directions for building planting troughs. When preparing the soil for a
trough, remember that containers with broad bottoms tend to drain slowly, so use a well-drained mix or
provide ample-sized drain holes.
Plastic containers designed as terrariums are utilitarian. They are lightweight, less likely to break com-
pared to glass, easily opened, have a ventilation hole, and are shaped to accommodate plants. Some prob-
lems, however, are that plastic containers scratch easily and might be unsuitable in certain decors.
plants, the size of the opening, the amount of soil moisture, and the extent of damage to the roots during
planting. If too much condensation appears on the sides of a covered container, uncover it temporarily
or enlarge the opening in the plastic wrap to allow evaporation. For the first few days after planting, keep
the terrarium in a well-shaded place. In covered containers, a decrease in condensation is usually a signal
that watering may be needed. The effects of using commercial products, usually containing silicon or glyc-
erine, for reducing condensation is mixed. In any case, avoid getting these products on the plant or in
the soil.
Maidenhair Ferns
Maidenhair ferns (Adiantum) have a fine billowy appearance and shiny black stalks that, because they
resemble hair, give the group its name. They are found in many parts of the world but are most abundant
in the American tropics. Most of the cultivated species are finely divided, and many have ruffles, fringes,
crests, and other types of fancy foliage.
The common species are easy to cultivate. The three most important factors are moist soil, good drain-
age, and humidity. Let them go to the dry side of moist before rewatering, but not to the point of wilting.
Adiantum is best grown outdoors or in greenhouses instead of indoors, unless a humid place is available such
as the kitchen or bathroom. Shriveling of new fronds is usually a sign of low humidity, especially if the soil
is moist. The other common cause of shriveling is poor root development, possibly the result of over-
watering, lack of nutrients, or poor drainage. Maidenhairs are particularly sensitive to being planted in
G S F 103
pots that are too big for them, as the roots are poorly aerated in oversized pots. Most maidenhairs slow their
growth by the fall and take a rest period until spring. Water them lightly during this inactive period because
too much watering causes the rhizomes to rot.
Dividing, transplanting, or repotting is best done in spring before growth is renewed. A suitable soil is
one consisting of one part sand and one to two parts peat moss or leaf mold. A handful or two of coarser
material (ground bark or perlite) may be added to a 15 to 20 cm (6–8 in.) pot for better drainage. The delta
maidenhair (Adiantum raddianum), Venus’s hair (A. capillus-veneris), and fan maidenhair (A. tenerum)
thrive in calcium-rich soils. The use of balanced fertilizers usually satisfies this requirement, but lime, lime-
stone, or oyster shell may also be added. Fish emulsion or other suitable fertilizers are applied every three
weeks to ferns planted in sand and peat moss alone.
To maintain attractive plants, remove old or discolored fronds. Sometimes, especially with species that
form dense clumps (such as Adiantum capillus-veneris and A. raddianum), the fronds in the center will die.
To avoid the need for complete replanting, remove the dead part and refill the space with fresh soil. In frost-
free climates, all the spent and unsightly fronds may be removed between fall and spring, but in areas with
frosts, wait until spring to remove the dead fronds. In either case, damage to new growth can be avoided by
trimming off all the old fronds in spring just before the new growth uncoils. Although the plant looks bare
after this procedure, the alternative—having to remove the old fronds carefully without injuring the new
ones—is far more tedious.
Maidenhairs are plagued by several pests that must be controlled. Slugs and snails are a constant prob-
lem in some areas, and sowbugs and pillbugs can also cause damage. Aphids usually appear in spring and
can be difficult to eliminate. These pests can be controlled by applying the proper insecticide, but be
extremely careful in doing so because maidenhairs are easily burned by most insecticides. If it is unknown
whether an insecticide will burn the plant, apply it only on a test plant. (Also see Chapter 11 for more
information on controlling pests.)
Some maidenhairs are best grown under particular horticultural conditions. The American or five-
finger maidenhair (Adiantum pedatum) is easy to grow in cold-temperate climates but not in warm ones.
The Venus’s hair (A. capillus-veneris) and the rough maidenhair (A. hispidulum) are good for warm, tem-
perate, and subtropical climates. The delta maidenhair (A. raddianum) is more tender than the Venus’s hair
but possesses the advantage of having many attractive cultivars. The delta maidenhair and Venus’s hair are
sometimes slow to establish, but once in place they grow rapidly, particularly among cement sidewalks
and foundations where presumably more calcium is present. In tropical areas and greenhouses, you can eas-
ily grow A. tenerum and the coarser-leaflet forms such as the silver-dollar fern (A. peruvianum), diamond
maidenhair (A. trapeziforme), two-edged maidenhair (A. anceps), and large-leaved maidenhair (A. macro-
phyllum). The rough maidenhair tolerates lower humidity than most and is suitable as a houseplant.
Several species of maidenhair are especially attractive but difficult to grow. The kidney-shaped fronds
of Adiantum reniforme are handsome and eye-catching, but the plant is tender and hard to cultivate. The
trailing maidenhair (A. caudatum) is interesting for its long fronds that produce a new plantlet at their
whip-like tips. It is easy to grow in well-drained soil but must be cultivated in a warm greenhouse. Adiantum
×tracyi, a semi-hardy, evergreen species, is rarely cultivated because it reproduces only from divisions, not
spores. Large elegant fronds are produced by A. formosum, but because of its wide-creeping habit this spe-
cies is not grown in pots, and few people have greenhouses with beds large enough to accommodate it. It
will grow outdoors in warmer subtropical gardens.
An old favorite in cultivation is the Farley maidenhair (Adiantum tenerum ‘Farleyense’). It has grace-
fully arching fronds and many ruffled segments. It develops most luxuriantly when grown in warm green-
houses with daytime temperatures near 21°C (70°F). Another maidenhair that is smaller but just as lovely
is green petticoats (A. capillus-veneris ‘Imbricatum’). It is a difficult fern for beginners but needs less heat
than ‘Farleyense’. Adiantum raddianum ‘Pacific Maid’ has broadly overlapping segments and the light
104 C 10
fluffy look of the preceding two selections, but it is a more erect, stiffer plant. It is good for beginners and
readily available in the trade.
With a greenhouse or similar means, many of the subtropical and tropical species can be grown with-
out difficulty. Keep in mind their particular needs for humidity and consistently moist but well-aerated soil.
For more details on cultivating Adiantum, see Chapter 13 and also Hoshizaki (1970a).
Staghorn Ferns
Staghorn ferns (Platycerium) are prized for their striking appearance and usefulness as decoration on walls
or tree trunks. In subtropical climates the common ones are grown outdoors year-round, and once estab-
lished, they require minimum care. Because they require only moderate humidity, staghorn ferns are suit-
able indoors, if provided with plenty of filtered light. Plants established and properly mounted need only
be hung in place and watered and fertilized occasionally.
If plants are watered with a hose, water them thoroughly. Small plants may be soaked in water for 10
to 15 minutes. Do not be tempted to water or sprinkle staghorns every time you water other ferns.
Staghorns need less water. More water will be required in a hot spell, less if the weather is cool and cloudy.
Staghorns are susceptible to overwatering by beginner growers because the outer surfaces of the base
fronds may feel dry even if the spongy inner layers are completely saturated with water. To test the mois-
ture, press your fingers firmly against the brown (not green) base fronds. If they are too wet, water will ooze
out. If they feel moist but no water oozes out, the moisture level is just right and the plant should not be
watered. Do not press against the freshly developed green base fronds because this can cause damage. Some
growers wait until the moss at the bottom of the planting is dry and crumbly before watering. Others wait
until the fertile fronds start to become limp. The weight of the plant also indicates relative wetness and dry-
ness. You should learn to determine whether a plant needs water by lifting it and judging its weight.
Inadequate watering usually results in slow or no growth. A sudden and severe lack of water causes
young fronds to wilt and the older ones to become limp and take on a grayish cast. The inner layers of base
fronds will be dry. Sometimes drought-like symptoms are caused by sowbugs, pillbugs, slugs, and snails eat-
ing the emerging roots. These pests are best seen at night.
When deciding where to hang a staghorn fern, be sure that it will not receive a constant drip of water
from overhanging baskets or from the eaves or roof. The symptoms of overwatering include no or poor
growth, absence of new base fronds, soggy base fronds, algae developing on the moss, and (at worst) pur-
plish to blackish decay spots on the base fronds.
Beginners should start with the easily grown species, such as Platycerium bifurcatum, P. hillii, and P.
veitchii. Slightly more difficult to grow and needing warmer conditions are P. willinckii and P. alcicorne.
Staghorn ferns can be propagated from the buds or young plantlets that form on their roots and some-
times their rhizomes. These plantlets are called “pups” by some gardeners. When the pups are big enough,
they can be separated from the parent plant. There is a greater chance of success if the pups are not removed
until they have several brown base fronds. Spring is a good time to remove them, or just before fresh shield
fronds grow out. Use a garden fork or knife to pry or cut beneath and around the oldest shield frond of the
pup (Figure 10.1). If possible, take a little of the base frond from the mother plant. Slip your fingers into
the cut and lift the pup from the mother plant. Plant the pup with the growing tip on the upper side.
The pups are usually mounted on either a tree or a board (Figures 10.2, 10.3). How you mount the pup
on a board depends on the total thickness of the base fronds. If more than 2.5 cm (1 in.) thick, the pup can
be mounted directly on a board and secured with wire (not copper) or stout strips of plastic. If the base
fronds are less than 2.5 cm (1 in.) thick, then place a 2.5 to 5 cm (1–2 in.) pad of sphagnum moss between
the base fronds and the board. If you are mounting the pup on a tree, do not wrap wire around the trunk,
G S F 105
which might girdle the tree and kill it. Instead, use plastic strips, nylon fishing line, discarded hosiery, or
string. If wire must be used, hammer small nails into the trunk and tie the wire to the nails.
It is also possible to grow a large staghorn in such a way that will not require remounting soon. Fill a
broken clay pot, wire basket, or shallow box with a loose, coarse humusy soil, covering or lining it with
sphagnum moss to keep the mixture from falling out (Figures 10.4, 10.5, 10.6). The moss can be kept in
place by covering it with chicken wire in which a hole has been cut for the plantlet. The surface of the plant,
particularly the buds, should be level with or slightly above the surface of the sphagnum moss. Plants so
mounted will have the advantage of more rooting medium, but the disadvantage of being susceptible to
106 C 10
overwatering. Recently mounted pups should be kept moist but well drained. When foliage no longer
appears wilted, the plants can be placed in more exposed places. The wires or strips of plastic will soon be
covered with new base fronds. Remounting established plants is the same as for mounting pups, except that
old base fronds might need to be trimmed away. To grow large symmetrical staghorns, keep one plant and
remove all the pups that appear.
Fertilize the plants about every three weeks, although less so during cooler months. Use liquid fertiliz-
ers because solid fertilizers, such as granules, powders, or pearl types, do not dissolve readily and, if caught
between the base fronds, can burn the tissue or promote infection by mold.
Once fungi or bacteria infect a leaf they are difficult to control. Several control methods are available,
but they are not always effective. Some growers report limited success by removing the decaying spot before
it spreads. They cut at least 1.5 cm (0.5 in.) beyond the decayed area. Others soak the plant in fungicides
(see Chapter 11). Some growers reduce the watering drastically in hope of killing the infection before
killing the plant. Prevention is better than an uncertain cure. Do not overwater, and use only thoroughly
dissolved fertilizer.
Aside from requiring a bright airy place and sufficient water, the common species of staghorn ferns pose
no special problems and will reward you with their striking and unusual foliage. If you have experience
growing the common species, the less common ones are also worth trying, although they can be more dif-
ficult. Most of these other species are large plants that require more space. See the references listed under
Platycerium in Chapter 13 for more details.
G S F 107
Davallia
The squirrel’s-foot fern, Davallia mariesii var. stenolepis (D. trichomanoides of trade), is the most commonly
cultivated Davallia. Its finely divided triangular fronds attract attention, as do its long, chaffy rhizomes that
can creep over an entire basket. Davallias are adaptable to a wide variety of situations. Although usually
grown in hanging baskets or pots, they are also suitable as ground covers and even over rocks if given
enough soil to secure a footing. They can also flourish indoors in sufficient light. Because they are epi-
phytes with firm-textured leaves, davallias can usually withstand the slightly drier air indoors better than
most other ferns. They should not be kept too moist. If fertilized a few times during the growing season,
Davallia plants will multiply readily from their creeping rhizomes.
Most davallias grow seasonally, shedding their fronds once a year between fall and spring and then
entering a period of slow growth. Sometime later a flush of new growth fully refoliates the plant. The exact
time of foliage shedding, and the length of time before new ones appear, varies with the species and climate.
Fronds persist longer on luxuriantly growing plants. In warmer climates such as southern California and
Florida, the deciduous habit of most species is hardly noticeable because new fronds appear before the old
ones have withered. The fronds sometimes turn a beautiful yellow before they wither and drop. If old
fronds have not fallen by the time new ones appear, they should be removed to provide room for the new
ones. The self-shedding of spent fronds is a decided advantage in dealing with high-hanging baskets.
Baskets that become covered with layers of rhizomes should be replanted. In mild climates, the best time
to do this is the fall or spring. Tear the clumps apart, saving those clumps with the most growing tips, and
then discard the dead or old parts of the reusable clumps. Davallia baskets should be lined with uncut
sphagnum moss and filled with a humusy, loose planting mix. Keep the basket moist but well drained, and
set it in a humid place until the plants are established. Further details on planting in baskets are provided
in Chapter 7.
The tender species Davallia fejeensis and D. solida are frequently seen in greenhouses and conservato-
ries. In warm-temperate climates the following species will grow and withstand light frost and short peri-
ods of freezing temperatures: D. canariensis, D. mariesii var. mariesii and var. stenolepis, D. pyxidata, and D.
tyermannii.
tral stem. These can be removed or wound around the plant and tucked inside the pot to keep a tidy look.
Repotting Nephrolepis is best done before active growth, usually in the spring. The taller, older plants
may need to be set deeper into the soil than they were before, but be sure not to bury their stem tips. Cut
away most of the old leaf bases from the stem. Avoid planting them in oversized pots; they tolerate being
slightly root bound. Use an epiphytic soil mix or a mixture of half sand and half peat moss (see Chapter 5).
Most species of Nephrolepis can be propagated by sowing spores. All can be propagated by divisions or
by layering the stolons, which produce new plants when they touch the soil. Planting in benches or shal-
low, broad containers gives the stolons room to spread in all directions and take root. The resulting plants
can be separated and potted when they have two or more fronds. The Boston ferns usually do not produce
spores and must be propagated by rooting the stolons or by meristem culture. Provide bright airy places for
their culture. Commercial greenhouses use as much as 3000 to 3500 foot-candles of light or 73% shade
cloth (see Chapter 8 for details on propagation).
The numerous cultivars of Nephrolepis exaltata offer a wide variety of sizes, shapes, and textures. Some
cultivars produce fronds as long as 1.5 m (5 ft.), whereas others produce fronds that are extremely broad or
narrow. These and other fancy-foliage forms, especially the finely divided ones, often grow slowly.
Boston fern cultivars with finely divided or congested fronds must be watered carefully. Their foliage
cannot support water applied directly from above, which tends to mat and break the fronds. If allowed to
remain on the leaves, the water causes the leaflets to have a dark, water-soaked appearance and can result in
loss, disfigurement, or yellowing of the foliage. If water mats the foliage, gently shake it off or blot it up.
Cultivars with densely clustered leaves need to be groomed regularly. Remove the dead or declining
fronds and unwanted stolons. This will allow more air and light to reach the younger growth below and give
the plant a more attractive appearance.
Besides Boston fern, the tuber sword fern (Nephrolepis cordifolia) is easy to grow. It is probably the most
cold tolerant of all sword ferns, being able to withstand short periods near 0°C (32°F). In subtropical and
tropical climates it is commonly planted as an outdoor border plant—a purpose to which it is well suited
because of its usually stiff, erect fronds. It can even be clipped like a hedge. The tuber sword fern has sev-
eral cultivars, the most popular being ‘Tesselata’ (‘Plumosa’), which has divided leaflets. ‘Tesselata’ grows
more slowly than the species form. Even slower is ‘Duffii’, which has rounded leaflets and is best grown
under glass (except in warm, humid climates where it can be grown outdoors).
The scurfy sword fern (Nephrolepis hirsutula) is rank and generally unattractive except in warm, humid
places. It grows poorly where nighttime temperatures dip below 16°C (60°F) and has no advantage over the
other species except that it grows easily from spores.
Nephrolepis species or cultivars with long, drooping fronds are difficult to find commercially because
growers dislike the difficulties involved with growing and shipping them. They must be displayed in bas-
kets or elevated containers to be attractive. Some long-frond types sold by specialty nurseries include
Nephrolepis falcata, N. pendula, and N. exaltata and its cultivars ‘Gretnae’ and ‘Rooseveltii’.
The rabbit’s-foot ferns or golden polypodies (Phlebodium) are among the most popular polypodiums
grown in the United States. Phlebodium aureum and P. pseudoaureum include several cultivars, some of
which have attractive bluish gray foliage. Once established, the golden polypodies grow rapidly if given
plenty of light. They should be watered when the soil is nearly dry. Some selections tolerate cold better than
others. Variants with broader, thinner lobes and sori in two or more rows on each side of the segment need
warmer temperatures than those with the narrower, firm lobes and sori in one to nearly two rows on each
side of the segment. The bluish green P. aureum ‘Mandaianum’ has thin, ruffled lobes. If unblemished
fronds are desired, grow it indoors or in sheltered places.
Knight’s polypody (Goniophlebium subauriculatum ‘Knightiae’) is another favorite basket fern. The
long, pinnate fronds are fringed and arch down to form a beautiful hanging plant. Established plants tol-
erate irregular watering. Outdoors in warm areas, the old fronds are shed in spring but are soon replaced
with new ones. The cut fronds last a long time in flower arrangements.
Another polypody fern, Microsorum pustulatum, is not widely cultivated but should be. In England it
is a hardy fern, and it should be tried in cooler areas of the United States. Microsorum pustulatum is medium-
sized with deeply divided blades resembling those of the East Indian polypody (M. scolopendria).
Microsorum punctatum ‘Grandiceps’ makes a handsome houseplant. Its thick, glossy leaves, ruffled
and irregularly forked or notched, withstand the drier air of the indoors admirably well. Because it grows
slowly, it seldom needs repotting.
A coarse, large, durable member of the polypody group is Aglaomorpha coronans. Although it grows
slowly, the species tolerates periods of dryness and aridity once established. Remove the old fronds as they
start to decline, because otherwise the individual leaflets will drop and scatter about.
Most of the small polypodium relatives that have simple, entire fronds are tender species and must be
grown in humid greenhouses or terrariums. Because they are small, they are not particularly showy and
therefore seldom seen in collections. One of these small species, Microgramma squamulosa, is attractive
due to its dark veins.
The temperate species of Polypodium vary in their cultural needs. The more common ones are planted
in baskets or among rocks. The licorice fern (Polypodium glycyrrhiza) often volunteers in baskets lined with
moss collected from western Canada and the western United States. A choice species is the leathery polypody
(P. scouleri), which grows along the coast in the Northwest. Its fronds are small- to medium-sized, deeply
lobed, thick, and glossy. It grows slowly and is suitable for pots and baskets in cool humid climates. The res-
urrection fern (P. polypodioides), a common epiphyte in the southeastern United States, is a small fern that
curls its leaves during drought and unrolls them when wet weather returns; it is difficult to establish.
Tree Ferns
Whether planted individually or in groups, tree ferns are sure to be noticed. They can provide a back-
ground for other plants, and their trunks can be used to display small epiphytic orchids, tillandsias, and
ferns.
Most ferns that bear fronds at the top of a tall trunk are called tree ferns by gardeners, although such
growth forms are found in unrelated ferns such as Sadleria, Blechnum, and Ctenitis. Most of the ferns in the
Dicksonia and Cyathea families (Dicksoniaceae and Cyatheaceae) are tree-like and are the ones typically
referred to as tree ferns by botanists. They range from 1 to 15 m (3–50 ft.) tall. In many species, the girth
of the trunk is increased by producing a thick layer of hard, dark, tangled roots—the root mantle. This layer
forms a substrate on which epiphytic plants grow.
Ferns of the Cyathea and Dicksonia families, as well as some tree-fern-like members of Blechnum, are
mostly native to cool, moist mountainous regions of the tropics (a few species occur in warm-temperate
110 C 10
areas). There are about 700 species in these two families, but probably fewer than 50 have been tried in cul-
tivation in Europe and the United States.
Temperature tolerances of the tree ferns are the first consideration in selecting them for cultivation.
Some species tolerate occasional frost and mild freezes, whereas others are intolerant of prolonged periods
at temperatures of 15°C (59°F). Nighttime temperatures are also an important consideration because many
species prefer cooler nighttime than daytime temperatures. Those tree ferns native to lowland tropics where
nighttime and daytime temperatures are relatively constant do well outdoors in tropical gardens. The many
species native to Australia and New Zealand grow well outdoors in subtropical areas, and some (such as
Dicksonia antarctica) grow in mild-temperate areas. In cold-temperate areas, tree ferns must be moved
indoors during the winter.
Tree ferns should be planted in well-drained soil. Add peat and sand (or perlite) to heavy soils, or
replace parts of it. Windy sites must be avoided. Although some species can grow in direct sun, they are
more luxuriant in shade. Water from overhead leaves or hanging baskets should not be allowed to drip into
the crowns of tree ferns. Provide plenty of overhead space for development. Tree-fern diseases continue to
be reported on wild and cultivated species. Rhizoctonia and tip blight (see Chapter 11) infect tree ferns, and
there may be other diseases as well.
Although some tree ferns can be shortened by severing their trunks and replanting them, the risk
involved in doing this is high. Some species root from trunks more readily than others. The Australian tree
fern of the trade, Sphaeropteris cooperi, does not root from severed trunks, whereas Cibotium glaucum, C.
menziesii, and Dicksonia antarctica root fairly well.
Cutting the trunk is best done before the active growing period and before new fronds emerge (usually
in the spring). After cutting, dust the cut ends with a fungicide to protect against rot, then plant in a pot
with a well-drained soil mix, and water sparingly to discourage rotting. Avoid watering the crowns, which
are fleshy and prone to rot. The number of remaining fronds can be reduced by half to prevent too much
water being lost from the trunk. If the trunks are too fleshy and not in danger of drying, it might be best to
wait before planting until a protective “skin” develops over the cut tissue. To encourage this, put the trunk
in a cool, humid place and keep it from drying. After the skin develops, plant the trunks about one-fourth
to one-third their length into moist soil, making sure to keep the trunks shaded and humid. When new
fronds emerge, the plant can be watered more. A few months before cutting the trunk, wrap a layer of
moist uncut sphagnum moss above the cutting point. This encourages roots to form, which then grow into
the moss. The moss layer should be thick enough that it will not dry quickly. Ample humidity and warmth
hasten root formation.
The Australian-native Cooper’s tree fern (Sphaeropteris cooperi) is the most frequently grown tree fern
in the United States. It grows in sun or shade in coastal, central, and southern California, in southern
Florida, and in conservatories in cooler areas. It tolerates frost and short periods of freezing temperatures,
but its foliage might die back. On older plants the fronds tend to spread more horizontally than on younger
plants and may be 6 m (20 ft.) across. The trunks are relatively slender, and cultivated plants seldom form
extensive aerial roots enveloping the stem. Because these roots are usually absent, the trunks are weak and
can snap off in strong winds. Therefore Cooper’s tree fern is best grown under tall trees where their crowns
have room to spread but are protected from wind. The lower fronds will droop or drop several times a
year. This foliage drooping is usually accompanied or preceded by a heavy discharge of spores. For the
sake of neatness, drooping fronds close to the trunk can be removed, but if left on the plant they eventu-
ally dry and fall off on their own. Under optimum conditions this fern may grow 30 cm (1 ft.) per year when
young, and the cultivar S. cooperi ‘Brentwood’ grows even more rapidly. The fronds and trunk of Cooper’s
tree fern are covered with many small scales that can irritate the skin and especially the eyes. Washing the
skin removes the scales and itching. Goggles should be worn to protect the eyes when working with this
fern.
G S F 111
The Tasmanian dicksonia (Dicksonia antarctica) is the second most frequently cultivated tree fern. It
is semi-hardy to semi-tender and has been reported to endure −7°C (20°F), or a bit lower for short periods,
as well as snow on its fronds. It prefers an average winter temperature of about 9°C (48°F). Its fronds are
shorter, narrower, stiffer, harsher, and more numerous than those of Sphaeropteris cooperi. The crown of
foliage reaches about 4 m (13 ft.) across and has a tufted appearance on the stout trunk. It grows more slowly
than Cooper’s tree fern—about one-third as fast. The Tasmanian dicksonia prefers climates influenced by
the ocean or cool and humid conditions. Dicksonia fibrosa and D. squarrosa are seldom cultivated because
they are more difficult to grow.
The Hawaiian tree fern (Cibotium glaucum) is seen now and then in outdoor plantings in southern Cali-
fornia. Bare-root trunks are imported from the Hawaiian Islands, and these can be rooted by planting
them about one-third their length in a well-drained soil mix. Spore-grown plants form more symmetrical
crowns but are infrequently found in the trade. The fronds of the Hawaiian tree fern tend to ascend and arch
more gracefully than those of Cooper’s tree fern. Their leaf bases and trunks are covered with silky, yellowish
tan hairs, unlike the chaffy scales of Cooper’s tree fern.
Occasionally, trunks of Cibotium menziesii and Sadleria cyatheoides are found in shipments of C. glau-
cum trunks due to their similar appearance. Cibotium menziesii is called the “man fern” by the Hawaiians,
possibly because its trunk is covered with stiff black hairs. It requires the same cultural conditions as the
Hawaiian tree fern. Sadleria cyatheoides tends to be more difficult to grow than the other Hawaiian species
and is sensitive to having its roots disturbed. Bare-root trunks of this species may be distinguished from oth-
ers by the presence of scales instead of hairs. Sadleria and Hawaiian cibotiums tend to grow more slowly
than Sphaeropteris cooperi.
The Mexican tree fern (Cibotium schiedei) was once widely sold as a houseplant, but it grows too big
for the average house. It hardly forms a trunk but produces offshoots freely around its base. Its many light
green fronds droop gracefully to give the plant a soft effect.
The silver tree fern (Alsophila tricolor) and the black tree fern (Sphaeropteris medullaris) need more pro-
tection from wind and sun than other tree-fern species to look their best.
Cultivated species that resemble tree ferns include Blechnum gibbum and B. brasiliense. These ferns are
medium-sized and form erect trunk-like stems. They are semi-tender to tender. The plant sometimes called
the American tree fern (Ctenitis sloanei) is native to southern Florida and might form a short, erect stem,
giving it a tree-fern–like appearance. Diplazium esculentum could be considered a small tree fern because it
produces an upright stem, but it is only 2 to 4 cm (0.75–1.5 in.) in diameter. This Asian species has become
naturalized in Louisiana and Florida. It is cultivated in Florida and California.
Xerophytic Ferns
Xerophytes are plants adapted to dry climates. Many xerophytic ferns grow in the arid southwestern United
States. In more humid regions of the country, they are found in exposed places such as on tree branches,
rocks, or cliffs.
Many fern genera have xerophytic species. Genera that are mostly xerophytes include Actiniopteris,
Astrolepis, Cheilanthes, Doryopteris, Notholaena, Pellaea, and Pityrogramma. Hairs, scales, or powder cover
the foliage of many xerophytic ferns to help retain moisture. These coverings can be white, silver, or yellow
and make the fern especially attractive. Ferns that are adapted to dry climates are often small and therefore
suitable for rock or trough gardens. Some xerophytic species are challenging to grow and may require sev-
eral attempts before the proper conditions are found.
In cultivation, xerophytic ferns grow best in bright but indirect sunlight, except along cloudy coastal
areas where they may be planted in full sun. They prefer a well-drained soil kept on the drier side of moist.
112 C 10
Most grow well in an evenly moist soil or one that does not rapidly fluctuate between wet and dry. In gen-
eral, the fronds should be kept free of water droplets resulting from condensation and overhead watering,
although fog does not seem to bother some xerophytic ferns.
Soil mixes range from simple to complex, but all must be well drained. A simple mix consists of one part
peat moss or leaf mold and one to two parts gravelly sand. A more complex mix consists of one to two parts
decomposed granite about 6 mm (0.2 in.) in diameter, two parts compost (leaf mold or fir bark), one part
perlite (#2 size), and one part sand (#20 size). Adding horticultural charcoal to the mix is optional, as is plac-
ing a top dressing of half-inch fine gravel on the soil surface after planting.
Xerophytes should be planted with their crowns slightly above the soil. If using pots, make sure that the
soil level is close to the rim in order to reduce the amount of water that could be caught in the pot during
watering. A process known as double potting maintains uniform soil moisture over a longer time. The
fern is planted in a porous clay pot, which in turn is planted in a larger clay pot, usually 5 to 7.5 cm (2–3
in.) wider than the first. The same soil mix is used in both pots. Xerophytes can be successfully grown in ter-
rariums if the soil moisture is carefully monitored and the humidity not excessive. Soil water evaporates
slowly in a terrarium, thus maintaining a more constant moisture level. In such a protected environment
fronds may develop more fully than in nature.
Outdoors, xerophytes are often planted in trough gardens (see “Trough Gardens” earlier in the chap-
ter), among rocks, or on well-drained sites (also see “Rock Garden and Wall Ferns” in Chapter 9). Xero-
phytic ferns are extremely sensitive to overwatering and can die if overwatered only a few times, and so they
should be planted away from plants that require more water. Whether in the ground or in pots, the plants
should be watered early in the morning so that any water settling on the fronds will evaporate during the day.
Most xerophytic ferns go dormant during the summer in their native habitats. In cultivation, however,
dormancy might not occur. In addition, many xerophytic ferns grow more slowly or go dormant as cool
weather approaches. Dormant or slow-growing plants need less water.
Filmy Ferns
Filmy ferns have been grown successfully in botanical gardens and private collections and were popularly
grown in Wardian cases in England during the Victorian era. These mostly small ferns are distinctive by
their membranous leaves usually one cell thick between the veins. Because of their thinness, the plants
need high humidity and shade. They also require good drainage. In nature most grow as epiphytes or ter-
restrially, and the group reaches its greatest development in tropical montane forests that are covered by
clouds most of the day and have an abundance of mosses on tree trunks and branches. The filmy-fern fam-
ily (Hymenophyllaceae) also includes the kidney fern (Trichomanes reniforme), an eye-catching species cul-
tivated in New Zealand and Australia but rarely in the United States. The elegant Prince-of-Wales plume
(Leptopteris superba) is not a filmy fern even though it has membranous leaves; however, it does require the
same cultural conditions as filmy ferns. It is rarely grown in the United States but is grown in Australia, New
Zealand, and England.
Filmy ferns from warm-temperate areas grow well in temperatures from 4 to 21°C (40–70°F), although
they also tolerate short periods of slightly lower or higher temperatures. Climates where temperatures are
consistently higher than 27°C (80°F) for days are unsuitable. Most tropical species come from cooler mon-
tane forests and grow best in temperatures ranging from 16 to 24°C (60–75°F). In favorable climates filmy
ferns can be grown in terrariums or bottles provided with uncut sphagnum moss or a well-drained potting
soil. In warm climates, air conditioners are needed to keep the temperature low.
Special chambers or houses, often located in a shady corner of a greenhouse, can be built to hold filmy
ferns. An intermittent misting system should be installed and, depending on the climate, possibly some
G S F 113
means of cooling. The chamber’s top and one side should be made out of glass or translucent plastic. If the
chamber is situated in an area with abundant light, only the top needs to be glass. Line the walls with
uncut sphagnum moss held in place by nylon fishing line or chicken wire (do not use copper wire). Plant
the ferns in pots or on pieces of tree-fern trunk and hang them on the moss, or plant them directly into the
wall of moss. Humidity should be kept near 100% by misting several times a day or by automatic humid-
ifiers. Chamber walls can also be made out of hollow concrete blocks filled with sphagnum moss or suit-
able excelsiors. The filled blocks are placed with the open end up, and a pipe set on the top of the block
drips water into the moss to keep the walls moist. Ferns are hung in pots against the wall. Some means of
draining the water away from the enclosure should be provided. The quality of water can also pose prob-
lems. If the salt content is high, salt-free water will have to be used. Farrar (1968) achieved good growth
of filmy ferns in chambers at 100% relative humidity with temperatures between 18 and 24°C (65–75°F)
and indirect natural light at or below 300 foot-candles. The ferns were given a fine mist for 15 minutes
every hour.
Fern Allies
Fern allies are the whisk ferns (Psilotum), ground pines (Lycopodium), spike mosses (Selaginella), and horse-
tails (Equisetum). Like ferns, these plants reproduce by dispersing spores, but none has leaves like ferns.
Instead, the leaves of the fern allies are small, inconspicuous, and often scale-like with only a single vein.
Water clovers (Marsilea), mosquito ferns (Azolla), and water spangles (Salvinia) are ferns but might be
mistaken for fern allies because of their unfernlike appearance.
used in Florida. Warmth, humidity, and constantly moist soil with good drainage are required for most epi-
phytic species.
Tropical epiphytic lycopodiums are usually propagated by divisions or layering because they are diffi-
cult to start from cuttings. Species such as Lycopodium phlegmaria can be propagated by layering or by
anchoring the stem tips in vermiculite or sand and waiting for rootlets to develop. Rooting should take place
in six to eight months, after which the plantlets can be separated from the mother plant.
Besides layering, cuttings are another method of propagating, although it tends to be less successful. The
usual procedure is to take 8 cm (3 in.) long pieces from the tips of stems bearing sterile leaves (that is, those not
bearing sporangia). The leaves are removed from the lower third, and the cutting is rooted in clean potting soil
mixed with four to five parts sand. Be careful not to overwater the potting soil, otherwise rot can easily result.
Provide the cuttings with only enough moisture and humidity to prevent rotting. The temperate Lycopodium
selago has been successfully propagated from cuttings rooted in uncut sphagnum moss and watered with dis-
tilled water. Whether propagating by layering or cutting, some bottom heat will hasten the process.
Growing lycopodiums from spores is rarely done, but the process would be essentially the same as for
selaginellas, discussed in the following section.
probability that both kinds of spores will be present. The methods for hybridizing selaginellas are somewhat
involved due to the presence of male and female spores. For details, see Webster (1979).
F F
Some hardy deciduous ferns that are to be exhibited out of season may be forced into early growth for
the show. Others, such as Woodsia glabella and Polystichum braunii, apparently do not take kindly to forc-
ing. Some will develop deformed fronds, and others will collapse soon after producing fronds.
Thurston (1939) successfully forced many ferns from New England and mid-Atlantic states for a flower
show in March. The ferns were planted in pots or flats in fall or earlier. They were placed in outdoor frames
and covered with lath. As the weather cooled, the ferns were gradually covered with leaves. The ferns were
permitted to freeze in November but under the protection of layers of leaves, branches, lath, glass sash, and
straw so as to be easy to dig out at the end of December. Plants were then removed from the frames and
placed in a dark shed to thaw for four days at temperatures between 4 and 10°C (40–50°F). Afterward they
were moved to a greenhouse and given light and exposed to temperatures of 13°C (55°F) during the day and
4 to 10°C (40–50°F) during the night. After five days, the temperature was raised to 21°C (70°F) during
the day and 10°C (50°F) at night. The water, heat, light, humidity, and ventilation for each species was judi-
ciously adjusted in the greenhouse. The water given was at 21°C (70°F) and was not permitted to touch the
foliage of most of the ferns; the exceptions were Camptosorus and others that like humidity, which were
G S F 117
misted to create relative humidity of 80%. Hardening off of the ferns began in the second week of Febru-
ary, and temperatures dropped to 13 to 16°C (55–60°F) during the day and to 7 to 10°C (45–50°F) at
night. The ferns were ready for the show by the first week of March. Thurston observed that forced ferns
produced good growth for the show, but the continued new growth was not as vigorous as in nonforced
ferns.
Most semi-hardy and more tender ferns can be kept in show condition by keeping them in warm
greenhouses and protecting the fronds from damage. Little is known about forcing tender species that are
deciduous.
11 Troubles with Growing Ferns
118
T G F 119
Root damage caused by overwatering or If roots have deteriorated, trim dead parts
noxious materials. The soil may smell bad. away, replant the rhizome into fresh soil,
keep moist but not wet, and hope for the
best.
S: F , ,
.
Common Cause Possible Solution
Too much humidity. Reduce humidity.
Too much nitrogen fertilizer. Reduce nitrogen fertilizer.
B C
Biological control uses beneficial organisms to kill pests. Familiar control organisms include the pray-
ing mantis, lady bugs, lacewings, ant lions, spiders, lizards, toads, and birds. Less well known are bacteria,
fungi, nematodes, parasitic mites, and wasps. All these controls are harmless to humans and pets. New
control organisms for specific pests are constantly being offered, and inquires should be made to county
agricultural extension services for restrictions and suppliers.
To work effectively, control organisms must be released before infestation becomes severe and well after
application of any long-lasting insecticides. Results take time, and close monitoring might be required to
determine effectiveness. Several releases may be necessary. Control rather than annihilation is the objective.
If control organisms are microscopic (such as bacteria), they can be formulated as a dust or liquid solu-
tion. Insects or animals are released as eggs, juveniles, or adults, depending on the species. Severe infesta-
tions are best treated first with a quickly biodegradable pesticide before the release of the controls. Because
healthy plants resist pests better than unhealthy ones, it is essential that you maintain proper fertilization
and optimum cultural conditions. You can also encourage naturally occurring controls such as spiders,
lizards, and toads.
To maintain beneficial organisms after they have controlled the pest might require advanced plan-
ning, unless you intend to make new releases. Ask the supplier how to provide for the control organisms
until the next pest attack or season. It might require alternative food sources and suitable habitats and pro-
tection from certain pesticides.
0.4 and 2.0%, or about 5 to 10 ml per 480 ml (1–2 tsp. per 2 cups) of water. The more detergent, the more
effective the spray and the more risk of burning the fronds. One test using a 3% solution found no dam-
age to maidenhair ferns (Adiantum). Studies indicate that soap used with acephate (Orthene), pyrethroids,
and chlorohydrocarbon pesticides increases the effectiveness of the spray. Commercial insecticidal soaps are
also available, but commonly used brands are not recommended for most ferns.
Alcohol also kills insects. It must be applied laboriously with a cotton swab by touching it to the insect.
Generally concentrations of 35 to 40% are used.
P-D I
Although plant-derived insecticides are rapidly biodegradable and generally safer to use than synthetic
chemicals, they are not selective and could harm beneficial insects and fish. Sprays and dusts are available,
and protective gear should be used to prevent inhalation. Several new plant-derived insecticides, such as
neem (Margosan), seem promising. Old standbys include rotenone, best if ingested by the pest, and
pyrethrin (pyrethrum), which is best for adult insects. These pesticides kill ants, aphids, caterpillars, cut-
worms, leaf hoppers, mites, thrips, whiteflies, earwigs, mealybugs, millipedes, and sowbugs. They are inef-
fective against adult scale insects.
Nicotine compounds are also plant-derived insecticides. Although they degrade quickly, they are toxic
to mammals and therefore unavailable to home growers. Commercial growers use them against pests such
as aphids, young scale, mealybugs, thrips, and fungus-gnat larvae.
Several synthetic plant-derived insecticides (as pyrethroids modified from pyrethrin) are on the mar-
ket. They have the advantages of being more effective on specific pests or longer lasting. Some are men-
tioned under the pest that they best control.
P
Pheromones are substances given off by plants or animals that cause specific behavioral responses in
other individuals of the same or different species. Some pheromones are sex attractants used to lure insects
into traps. Most pheromones available in the trade are used to control moths, flies, and beetles whose
young feed on plants and may only incidentally feed on ferns. Pheromones are nontoxic to humans, pets,
and the environment.
ultraviolet light, and such lights are used in “bug zappers” with electrically charged grids that kill adult fly-
ing insects on contact. Ultraviolet light can be useful to control moths that in their caterpillar or grub stage
eat plants.
Mulches may help reduce surface-crawling pests, and a layer of ashes often discourages slugs and snails,
but use sparingly because too many ashes could harm the soil. A commercially available diatomaceous
earth dust is reportedly effective against soft-bodied insects. The microscopic needles of silica in the dust
puncture the insect’s guts after ingestion. It should not be applied when beneficial insects are active. Report-
edly, this talc-like powder can be handled safely without gloves.
Spraying with water dislodges some pests and gives a certain measure of control. Adding a few drops of
liquid detergent to the water helps loosen pests attached to stems or leaves. Picking off insects by hand or
dabbing them with an alcohol- or oil-soaked cotton swab is labor intensive. The best time to look for slugs,
snails, and cutworms is at night, especially when the ground is moist. Providing good sanitation will reduce
hiding and breeding places of pests.
5. Test the products on a few plants before applying them to other plants. Damage usually
appears after one day to two weeks and is evident as wilting, yellowing, spotting, marginal
or surface burning, or abnormal growth.
6. Proper dosage is given on the label, but only a few fern species have been tested at the
dosages listed by the manufacturer. The recommended dosage will often damage
susceptible ferns. Reducing the dosage, however, violates the label law, even if a lower
dosage is effective and does not damage the fern.
7. Sprays and dusts are most effective when thinly and thoroughly applied to all surfaces of
the foliage. Fine mist sprays are the best. Water at the soil level after spraying or dusting so
that the residue is not washed off the foliage.
8. Aerosols or smoke generators should be used only in greenhouses or tightly sealed plastic
houses.
9. Spray, dust, or apply granules when the leaves are dry to avoid or minimize damage.
10. Spray early in the day so that plants can dry rapidly.
11. Spray when the air temperature will stay below 29°C (85°F) for at least two hours after
spraying.
12. Move houseplants outdoors for treatment if possible.
13. The healthier the plant, the less likely the possibility of spray damage. Plants most easily
damaged are those that are root bound, in need of water, or very young. Any damage that
does occur might only affect the existing foliage, and any new growth may be unharmed if
the roots and rhizomes have not been damaged.
14. Assume that all pesticides and fungicides are toxic. To reduce the possibility of damage to
your (and others’) health, proper procedures, protective clothes, and equipment must be
used, and re-entry periods observed.
15. For the commercial grower, the suitability of a pesticide or fungicide for large-scale use
should be checked with local county agents, state experiment stations, extension specialists,
or the manufacturer’s technical representatives. These specialists can determine the
suitability of a specific pesticide for local soils, temperatures, moisture conditions, cultural
practices, and dosage rates.
Systemic pesticides are those taken into the plant tissue and are effective in controlling the pest or dis-
ease from within the plant. A drench is a solution applied by sprinkling it over the soil.
A
Aphids are small, soft-bodied insects that are red, green, yellow, orange, or black. The fern aphid is black
with whitish legs. Aphids weaken plants by sucking juices and causing distortion and stunting of foliage.
They also produce a secretion called honeydew that is sipped by ants. Because the ants encourage and
protect the aphids, they must also be controlled (see above). Some aphids are easy to kill but difficult to
control because reinfestation often occurs from nearby areas. The fern aphid is common on new growth,
particularly in the spring (Figure 11.1).
P C. For nonchemical control, see the various measures discussed under
“Alternatives to Chemical Controls,” earlier in the chapter.
Acephate (Orthene) is a systemic, broad-spectrum insecticide. Fern growers prefer the 75% soluble
powder used at the manufacturer’s rate of 5 ml (1 tsp.) to 3.8 liters (1 gal.) of water. Because damage has
been reported on Blechnum gibbum, Phlebodium aureum, Polystichum, and Pteris ensiformis, you should do
a sample test before using acephate powder on your ferns. Emulsions are more easily available and should
also be tested before wider use. The manufacturer recommends an acephate emulsion of 9.4% used at 30
ml per 3.8 liters of water (2 tbsp. per gal.) for aphids and 45 ml per 3.8 liters of water (3 tbsp. per gal.) for
mealybugs and scale insects on ornamentals in general. This dosage, however, could burn some ferns, so test
first. Orthene aerosol might also burn some ferns.
Carbaryl (Sevin) is a broad-spectrum insecticide that can be used around edible plants. Carbaryl is
registered for aphid use, but only for the apple aphid and rose aphid. For the fern aphid, other insecticides
126 C 11
are more effective, though carbaryl is also useful for other pests that attack ferns. It is available as a dust,
emulsion, or wettable powder. Emulsions burn sensitive ferns.
Diazinon is another broad-spectrum insecticide. Diazinon emulsions of 25% are recommended by
the manufacturer to be used at 10 ml to 3.8 liters of water (2 tsp. per gal.) for foliage plants, but not for cer-
tain ferns. Dosages reduced one-quarter to one-half the recommended concentration gave adequate con-
trol with little damage, but such dosages violate label laws. Wettable diazinon powder of 50% at 15 ml per
3.8 liters of water (1 tbsp. per gal.) is recommended for foliage plants; however, the manufacturer also
reports that maidenhairs and other ferns are burned at this concentration. In all cases, sample test on a few
plants before wider use.
Dimethoate (Cygon 2E), a systemic, broad-spectrum insecticide, is effective for three weeks after appli-
cation. When used as a drench to minimize foliage burn, Cygon 2E (23.4%) is used at 5 ml to 3.8 liters of
water (1 tsp. per gal.) for foliage plants in general. As a spray for foliage plants, the recommended dosage is
7.5 ml per 3.8 liters of water (1½ tsp. per gal.). Treatments of 2.5 to 5 ml per 3.8 liters of water (½–1 tsp.
per gal.) have been successful in tests on ferns. This product has restrictions and is unavailable in certain
states.
Disulfoton (Di-syston) granules applied to the soil provide systemic protection against aphids and
other insects. It is often included in rose-care products and is effective against scale insects, red spider-
mite, and thrips.
Fluvalinate (Mavrik Aquaflow) is a synthetic pyrethrin-like chemical. The water-based formulation
makes it safe for plants. Recommended dosage ranges from 1.3 to 3.3 ml per 3.8 liters of water (¼–⅔ tsp.
per gal.) for foliage plants.
Imidacloprid (Marathon) is a broad-spectrum systemic insecticide. It comes in wettable powder or
granular form. The active ingredient does not readily move in the soil, therefore drenches or granules must
be applied to the root zone to be effective. It reportedly remains active in the soil for 8 to 12 weeks after one
application. Some ferns reportedly do not respond to granular forms even if generously applied. It is not
available to home growers.
Kinoprene (Enstar 5E) is safe for Boston fern (Nephrolepis exaltata ‘Bostoniensis’) and its cultivar
‘Fluffy Ruffles’ and for Pteris cretica ‘Parkeri’ when used at the manufacturer’s recommended rate. It severely
damages maidenhair fern (Adiantum), however, and its effect on other ferns is unknown.
Malathion is a broad-spectrum insecticide available as a wettable powder but more commonly as a
dust or emulsion. The latter is most commonly used, and the manufacturer’s recommended dosage for 50%
malathion concentration on foliage plants in general is 10 ml per 3.8 liters of water (2 tsp. per gal.). This
dosage severely burns or injures a variety of ferns, although commercial fern growers encountered little or
no damage when the dosage was reduced to 2.5 ml per 3.8 liters of water (½ tsp. per 1 gal.), though this
reduction violates the label laws.
Nicotine sulfate is no longer available in the United States, although a nicotine alkaloid aerosol is avail-
able to commercial growers. It is highly toxic.
Oxydemetron-methyl (Metasystox-R) is a systemic insecticide. The 25% emulsion is used at 7.5 to 10
ml per 3.8 liters of water (1½–2 tsp. per gal.) for foliage plants. It is reportedly safe for maidenhair and other
ferns when applied to the soil.
Oxamyl (Vydate) controls aphids and a variety of insects, nematodes, and mites; however, it is available
only to commercial growers.
Pyrethrin kills aphids and comes in many formulations. Synthetic pyrethrins (pyrethroids) are also
available.
Resmethrin kills aphids and other pests and comes as a spray or aerosol. The aerosol form is favored by
some commercial growers.
Soap sprays are discussed under “Alternatives to Chemical Controls.”
T G F 127
E
Earwigs are reddish brown insects about 13 mm (0.5 in.) in length, with prominent pinchers at the end
of their tails. They ordinarily eat decaying vegetation, but they can damage ferns by nibbling at tender
new growth. During the day earwigs hide under flower pots and plant debris. They forage at night.
P C. Keep a clean culture. Trap earwigs during the night in a roll of damp
newspaper, and destroy it in the morning.
Propoxur (Baygon) bait is the insecticide of choice. Baits or dusts of carbaryl (Sevin), chlorpyrifos
(Dursban), and diazinon also control earwigs. Scatter the bait where earwigs congregate but not directly on
the fern foliage.
F G
Fungus gnats resemble small mosquitoes less than 3 mm (0.1 in.) long. With a hand lens the feelers
(antennae) are seen to be longer than the head, and the veins of the wing form a Y-shaped pattern. These
features are absent on shore flies, a similar gnat. Fungus gnats frequent damp places and soils with high
organic matter. When disturbed, the adults often run rapidly on the plant or soil before taking flight.
Both shore flies and fungus gnat adults can spread disease, but the fungus gnat causes the most dam-
age when in its larval stage. In greenhouses and enclosed places they eat roots, root hairs, stems, and pro-
thalli. Shore-fly larvae are maggot-like and eat detritus, whereas fungus gnats eat living plant tissue. Fun-
gus-gnat larvae are small, whitish, translucent, legless worms with black heads. They can be found beneath
the soil level or in the plant tissue being eaten. Affected plants grow slowly, lack vigor, may wilt, show yel-
low leaves, and have rotted roots. Prothalli appear wilted or have small but noticeable holes where the lar-
vae have fed. The greatest damage is seen during plant growth stages in which there are few roots and the
larvae have fed on them.
P C. Keep a clean culture. Remove weeds and plant debris, and take steps
to avoid excessive moisture in or on the soil and surrounding area. To control algae, which often flourish
in unclean cultures and attract fungus gnats, use clean or new flats and pots (see “Algae, Mosses, and Liv-
erworts”). On spore pots and flats covered with glass, seal any space between the glass cover and pot or flat,
preferably with plastic foam strips instead of cotton. If feasible, reduce the organic content in the soil mix.
Ammonium nitrate used as a drench at three-quarters the normal fertilizer concentration works for larval
control, and acts as a fertilizer as well.
If drenches or sprays are used, remove surrounding debris before application. Avoid irrigation for one
day after application. Drench, dust, or spray the plant, the soil, under benches, and any places where the
gnat or its larvae might be. Yellow sticky cards are useful for trapping adult gnats and for monitoring the
effectiveness of other control measures.
Biological controls include Bacillus thuringiensis and other organisms (see “Alternatives to Chemical
Controls”). Bacillus thuringiensis H-14 (Gnatrol) kills larvae. The drench works within 24 hours after
ingestion. Ferns are unharmed by the recommended dosages of 5 to 10 ml per 3.8 liters of water (1–2 tsp.
per gal.) for light infestations, or 20 to 40 ml (4–8 tsp.) for heavy infestations. A nematode that kills the lar-
val stages of the fungus gnat is sold as Exhibit.
Chemical treatments of choice include diazinon (also used in a capsule form called Knox-out), kino-
prene (Enstar 5E), oxamyl (Vydate 10% granules for the larvae and Vydate 2L spray for the larvae and
adults), and resmethrin (a synthetic pyrethrin). One percent resmethrin aerosol (PT1200 Resmethrin) did
not damage Boston ferns, and growers reported no damage to the commonly grown species of Pteris and
Pellaea. Resmethrin spray (SBP-1382-2EC) is recommended for foliage plants at 5 ml per 3.8 liters of
water (1 tsp. per gal.); its toxicity to ferns is unknown. Less effective, but also registered as a control for fun-
128 C 11
gus gnats, are dimethoate (Cygon 2E), carbaryl (Sevin), and malathion. See “Aphids” for details and
precautions.
Although unregistered against fungus gnat larvae, fluvalinate (Mavrik Aquaflow) seems a promising
control. Others products that are reportedly effective on fungus gnats include Fenoxycarb (an insect growth
regulator), Citation, Orthene, and Dycarb.
M
Several species of mealybugs attack fern fronds and (less frequently) roots. These small insects are char-
acterized by a white mealy look (Figure 11.2). They weaken a plant by feeding on its sap. Ants are attracted
to the mealybugs’ honeydew secretion.
P C. Adult mealybugs are more difficult to control than the immature
stages (crawlers). Therefore you should inspect plants frequently for infestation and employ control meas-
ures promptly. Control the ants (as discussed earlier); also see “Alternatives to Chemical Controls.” For root
mealybugs, use the spray as a drench.
Acephate (Orthene), diazinon, and malathion are the insecticides of choice. Also used are dimethoate
(Cygon 2E), imidacloprid (Marathon), kinoprene (Enstar 5E), oxydemetron-methyl (Metasystox-R), and
oxamyl (Vydate). If you have a choice, use systemics. See “Aphids” for more details.
T G F 129
M
Millipedes generally feed on decaying plant debris but also eat healthy fern foliage. They appear worm-
like but have many legs. The garden species is reddish brown, about an inch long, and emits a sweet, pun-
gent odor when crushed (reminiscent of toasted almonds). Millipedes may be confused with predatory
centipedes, which differ by having only one instead of two pairs of legs per body segment, and by generally
moving faster. Millipedes are most active at night. During the day look for them under flowerpots and
debris.
P C. Keep a clean culture. Remove debris.
Propoxur (Baygon) bait is the pesticide of choice. Carbaryl (Sevin) dust or a spray from 50% wettable
powder, at 30 ml per 3.8 liters (2 tbsp. per gal.), will also control millipedes. Spray into cracks, under con-
tainers, and in other hiding places, thoroughly wetting these surfaces. Avoid spraying on fern foliage.
Pesticides such as acephate (Orthene), chlorpyrifos (Dursban), diazinon, pyrethrin, and rotenone also
kill millipedes. Be sure to follow the directions on the label, and avoid getting the insecticide on foliage.
N N
Leaf nematode blight is caused by minute worms (Aphelenchoides fragariae). The nematodes form red-
dish brown or black spots that are typically sharply marked on the leaf. They typically feed on the leaf tis-
sue between the veins, seldom across the veins. Therefore the spots have an angular shape. On bird’s-nest
fern (Asplenium nidus) the discoloration is limited to tissue between the parallel veins. In other species and
on young ferns, the affected area may be more irregular due to the pattern of the veins. Nematodes can be
seen by placing a bit of infected tissue in a drop of water, then lightly mincing and examining it under about
20× magnification. With good light, you can see the characteristic thrashing movement of these nearly
transparent, slender worms (about 0.5 mm long). Leaf nematodes thrive under cool moist conditions.
They travel in a film of water or in drops from splashed water. Besides weakening and discoloring the
plant, nematodes spread disease. Root-knot nematodes have not been reported to attack ferns.
130 C 11
It is unlikely that nematodes spread through fern spores, unless fragments of infected leaf tissue are sown
with the spores. Rapid drying of the leaf tissue prevents nematodes from entering their rest stage. Other-
wise, they can remain alive for a year or more.
Certain ferns are more susceptible to leaf nematode damage than others. These include Asplenium
(A. bulbiferum and A. nidus in particular), Blechnum, Pteris, Tectaria, and Woodwardia (W. fimbriata and
W. radicans in particular). Also susceptible are Adiantum, Cyrtomium falcatum, Dicksonia antarctica, Diplaz-
ium dilatatum, Diplazium proliferum, Dryopteris filix-mas, Dryopteris hondoensis, Nephrolepis cordifolia,
Polypodium, Polystichum munitum, Rumohra adiantiformis, Sphaeropteris cooperi, and Thelypteris.
P C. Prevention is easier than control. Start with disease-free stock and use
only clean flats or pots. Beware of soil, materials, or tools that might harbor nematodes. Disinfect con-
taminated materials or tools, pasteurize suspected soil with steam at 82°C (180°F) for about 30 minutes,
or treat with a soil fumigant. Because nematodes thrive in debris, discolored leaves should be discarded to
where they will not reinfect other plants. Remove infected leaves and burn them. Avoid overhead watering
and other conditions that wet the leaves. Remove weeds that might harbor nematodes. For the fern hob-
byist with only a few infected plants, the best course is to dispose of them properly and enact preventative
measures to protect against further spread. Bird’s-nest fern (Asplenium nidus) has been successfully freed of
nematodes by immersing the plant in hot water at 46°C (115°F) for 10 to 15 minutes.
If the infection is from the ground, the plants should be grown in pots on raised benches. Do not allow
the infected soil to be splashed on benches, pots, or foliage. If planting the ferns in the ground, allow the
soil to remain fallow and weed-free for a season or so. This will decrease but not eliminate the nematode
population. Drying the soil also reduces the nematode population. Live mature marigolds (Tagetes patula)
will release from their roots a chemical (ozone) that kills nematodes. This control practice is only effective
if many marigolds are planted.
The chemicals used to control nematodes are generally unavailable to home growers because of their
toxicity to humans and pets and their ability to remain in the soil or plants for a long time if improperly
applied. Nematicides used on ferns include ethoprop (Mocap), oxamyl (Vydate), and phenamiphos
(Nemacur). All of these have restrictions on their use. With the constant changes in labeling and laws, it is
best to contact local agricultural agencies for current information. For more details, see Krusberg (1992) and
anonymous (1986).
S I
Adult scale insects live in a stationary, wax-like shell under which they suck plant juices. Extra juices are
secreted as honeydew, which attracts ants and serves as the medium for the growth of a blackish mold.
Several species of scale insects feed on ferns. The most common is the brown soft-scale. It has black or
brown spots on its back. Young scale insects (crawlers) continually emerge from the mother scale for about
two months and grow to full size in two more months (Figure 11.3).
P C. Adult scale insects are difficult to control, and therefore frequent
inspection and enacting controls at an early stage are important. Although biological controls such as kino-
T G F 131
prene (Enstar 5E) are available, natural predators can be encouraged by spraying the plants with water to
keep off dust. Soaps may be effective against crawlers. Adults can be controlled by dabbing with alcohol-
or oil-soaked swabs. Chemicals of choice to control the adults are diazinon, dimethoate (Cygon 2E), imi-
dacloprid (Marathon), malathion, acephate (Orthene, more effective against the crawlers). Other pesticides
that kill scale insects include oxydemetron-methyl (Metasystox-R), oxamyl (Vydate), and disulfoton (Di-
syston) granules. See “Aphids” for more details.
moss are other mulches that snails dislike to traverse. Slugs do not seem to object to sand. Snails are stopped
by vertical copper bands at least 4 cm (1.5 in.) wide, though slugs are less inhibited by this. Insecticidal soaps
reportedly kill slugs and snails.
Decollate snails are predators of brown garden snails but will eat tender foliage, including that of some
ferns. Slugs are rarely attacked by decollate snails. Toads, opossums, and other wildlife feed on snails and
slugs.
For chemical controls, various baits, granules, and sprays can be applied on less exposed areas of flats,
fences, and walls where snails and slugs travel and feed. The pests tend to return to the same hiding and feed-
ing areas.
Methaldehyde attracts and stuns slugs and snails, and death is dependent on subsequent dehydration.
Slugs and snails can recover and may live to develop resistant strains. Keep pets, particularly dogs, away from
methaldehyde baits, which are poisonous. Baits in pellet form can become unsightly due to mold; other
forms do not mold or the mold is less conspicuous. Methaldehyde is less effective at temperatures below
20°C (68°F) and loses potency in sunlight and high humidity. A special formulation (which can be obtained
as Deadline, Corry’s, and other brand names) resists sunlight and humidity damage and lasts longer. Sprays
have the advantage of penetrating the soil where slugs may be hiding, but earthworms will also be harmed.
Avoid spraying fern foliage.
Methiocarb (Mesurol) kills snails or slugs instead of stunning them. It also kills earthworms. The gran-
ular form should be applied when the foliage is dry. Increased humidity increases its toxicity; where humid-
ity fluctuates widely, using methiocarb in combination with methaldehyde will give better control. For use
in commercial greenhouses, methiocarb comes in aerosol form. It also kills aphids, scale insects, spider
mites, moths, and whiteflies.
Mexacarbate (Zectran) is infrequently sold. Because direct sprays will burn maidenhair and possibly
other ferns, it should be applied only to the soil and walkways. The dehydrating effect of aluminum sulfate
applied as a drench of 56.7 grams per 3.8 liters of water (2 oz. per gal.) is reported to kill snails and slugs,
but its effect on ferns is unknown. It might acidify the soil. For further details, see Parrella et al. (1985).
T
Thrips are small, slender insects that cause noticeable white or bleached areas on ferns by sucking the
plant juices. Bleached areas may become especially conspicuous during the summer months. Tiny, scattered
black or brown dots (mold-infected honeydew) left by the thrips are visible with a hand lens on the under-
side of the damaged area of the frond (Figure 11.4). By the time the bleached areas appear, none or only a
few lingering thrips will be seen. Greenhouse thrips, the species that attacks ferns, have dark brown to
black adults, about 1 to 1.5 mm long. This species is common and found worldwide. It may attack ferns
indoors or outdoors. The flower thrip does not form black dots on the foliage and has not been reported to
attack ferns.
T G F 133
P C. Keep the surrounding grass mowed and remove weeds and badly dam-
aged fronds. Ferns susceptible to thrips (such as Polystichum and some Dryopteris) should be placed in
cooler, more humid sites of the garden. If in a greenhouse, use 0.178 mm (0.007 in.) brass mesh screen
across openings. The chemical controls of choice are acephate (Orthene), disulfoton (Di-syston), malathion,
and oxydemetron-methyl (Metasystox-R). Others include chlorpyrifos (Dursban), dimethoate (Cygon
2E), fluvalinate (Mavrik Aquaflow), oxamyl (Vydate), and imidacloprid (Marathon). See “Aphids” for
more details; also see “Alternatives to Chemical Controls.”
W
Whiteflies are sucking insects about 3 mm (0.1 in.) long that fly upward when the foliage is disturbed.
Examination of the leaf’s underside will reveal many flattish, oval larvae and pupae adhering to the surface.
Whiteflies are uncommon on ferns, although in some areas newly introduced species are attacking some
ferns and are difficult to control.
P C. See “Alternatives to Chemical Controls” for biological controls, soaps,
and insect growth-regulators.
Chemical controls for greenhouse whiteflies are acephate (Orthene), diazinon, dimethoate (Cygon
2E), fluvalinate (Mavrik Aquaflow), malathion, oxydemetron-methyl (Metasystox-R), and resmethrin,
among other registered controls. See “Aphids” for more details. Insecticides used for greenhouse whiteflies
may not be effective against other whitefly species.
P C. The chemical control of choice is carbaryl (Sevin), but also used are
acephate (Orthene), diazinon, and chlorpyrifos (Dursban). See “Aphids” for more details. Also see “Alter-
natives to Chemical Controls.”
interiorscapes), for certain types of plants, and for certain users. As with pesticides, use of these chemicals
other than as described on the label violates the law.
In the following section the common names of fungicides and some of their trade names are given.
Trade names are used for convenience and do not imply endorsement. See Appendix IV for cross-references
between trade names and the active ingredient.
Armillaria R-R
Armillaria root-rot has been reported to kill Dicksonia fibrosa in southern California. It is widespread
and attacks woody tissue, but some woody plants are more susceptible than others. The infected plants
decline and eventually die. Prying into a dead stem will reveal the whitish growth of the mold. Eventually
the characteristic honey-colored mushrooms may emerge, usually around the base of the stem.
P C. Avoid planting Dicksonia fibrosa or woody plants in areas susceptible
to Armillaria unless the species are resistant. The soil can be fumigated before planting, but recontamina-
tion is likely to occur.
B
Blights are caused by fungi or bacteria that attack young growing tissue. Affected young plants will
wither and die, and the fronds of older plants will dry at the tips and margins (especially in Boston ferns).
Blights can appear as water-soaked spots on bird’s-nest ferns (Asplenium nidus). Also see the section “Tip
Blight” later in the chapter.
P C. Remove and destroy blighted parts and disinfect tools. Avoid getting
the foliage wet, reduce humidity, and space the plants farther apart to facilitate drying and improve air cir-
culation. If the specific disease organism is unknown, use a broad-spectrum spray (such as Banrot). Chem-
icals with the active ingredients propiconazole (Banner), chlorothalonil (Daconil 2787), iprodione (Chipco
26019), or mancozeb (Dithane M-45) might also be effective. Also see “Rhizoctonia” and “Water Molds.”
D-o
Damping-off diseases attack young plants near the soil surface and cause them to fall over, wither, and
die. Prothalli often reveal the symptoms of damping-off. Several different pathogens cause the disease.
P C. Sterilize the soil if possible. Provide good ventilation, keep tops of
young plants dry, and reduce the moisture level to a minimum. Broad-spectrum fungicides must be used
unless the pathogen is known. Commercial growers favor Banrot or a combination drench of iprodione
(Chipco 26019) and metalaxyl (Subdue). See also “Blights,” “Rhizoctonia,” and “Water Molds.”
maneb (Dithane M-22), thiophanate-methyl (Zyban), and zineb (Dithane Z-78). All are registered for
use on foliage plants, but often the only ferns that have been properly tested are the Boston fern and leather
fern. Spray test before widely applying any fungicide to your ferns.
L S
Spots and blotches on leaves are caused by factors such as poor air quality (see Chapter 4), physical dam-
age, excessive moisture or shade, fungi, bacteria, and nematodes. Healthy ferns in well-ventilated places sel-
dom become infected. Blemishes usually appear on older fronds, but new fronds can also become infected.
P C. With minor infections, the disfigured fronds should be removed to
prevent spreading. Cut the foliage back to healthy tissue, destroy the infected parts, and disinfect all tools.
Reducing humidity, spacing the plants farther apart, increasing air circulation, and avoiding overhead
watering or splashing will help prevent further infection. If the spots are few, smearing them with a cotton
swab dipped in a strong solution of baking soda, a 10% bleach solution (avoid healthy tissue), or a quater-
nary ammonium compound (Physan 20 or Green Shield) may help stop the spots from enlarging. This
treatment is particularly effective for the larger spots appearing on staghorn ferns. Experiments indicate that
insecticidal soaps can increase or decrease leaf spots, depending on the disease and host. A 0.5% baking-
powder solution has successfully been used to treat leaf spots but is not registered for such use. It is report-
edly more effective when combined with a horticultural oil, such as Volck Oil Spray, a petroleum oil.
Chemical sprays are a last resort. A broad-spectrum spray (see under “Blights”) must be used unless the par-
ticular pathogen is known.
Some specific leaf-spot pathogens that have been identified on ferns include Cercospora, Cladosporium,
Colletotrichum, Cylindrocladium, Fusarium, Myrothecium, Peyronellaea, Phoma, Pseudomonas, and Xan-
thamomas campestris. The latter two are bacteria that produce rot with a wet appearance (especially on base
fronds of Platycerium). They can be controlled by Physan 20 applied with a cotton swab. Leaf blister (Taph-
rina), which forms a blister on both sides of the leaf, attacks mostly native ferns.
Rhizoctonia
Rhizoctonia is a fungus that usually attacks plants near the soil surface. The stems or stipes rot at the
point of infection, and the weakened stalk cannot support the frond. The infection can also spread below
the soil. Thicker roots turn yellow or brown and have cankers. Prothalli and young plants wither and damp
off. Fronds may wilt, grow slowly, be deformed, or stop emerging altogether. This condition is called “hard
crown” by some growers. Rhizoctonia often causes black-brown lesions on stems, leaf stalks, and roots.
The base fronds (shields) of staghorn ferns may show wrinkly brown lesions.
The decaying parts of infected plants have brownish threads that can be seen with a hand lens. These
threads are coarse enough to cling to soil particles (water molds have fine threads that do not cling to soil par-
ticles). Under the microscope, the characteristic T-shaped cells where the filaments branch are diagnostic.
Rhizoctonia also attacks tree ferns. The new fronds do not uncoil but seem to dry in place. Mature
fronds will wilt, and the crown hardens and turns brown. Eventually the plant dies.
Overwatered plants or ones growing in poorly drained soils are most susceptible to infection. Infected
plants continue to grow but usually languish and succumb to minor stresses. High humidity and temper-
atures of 21 to 27°C (70–80°F) favor the disease.
P C. Discard infected plants. Make sure contaminated soil or equipment
does not touch clean soil or equipment. Sterilize the soil before replanting contaminated areas. The disease
is soil-borne and can easily spread by splashing or flowing water. Several chemicals can be applied as a
drench, mainly as a preventative. They will inhibit the disease but not destroy it. The chemical of choice is
T G F 137
pentachloronitrobenzene (PCNB, Terraclor), but it burns certain ferns. Also effective are chlorothalonil
(Daconil 2787), Cleary’s 2226, iprodione (Chipco 26019), captan, etridiazole (Truban), and the broad-
spectrum Banrot. In all cases it is a good idea to test spray first.
R
Many species of fungi and bacteria can cause decay or rots. Rots caused by bacteria appear wetter and
slimier than those caused by fungus.
P C. Cut away rotting spots. Disinfect tools and treat as for blights (which
see). Bacterial rots may respond to such bactericides as streptomycin (Agri-strep), copper sulfate pentahy-
drate (Phyton 27), and others. Some bactericides are expensive, and ferns may be sensitive to them. Small
spots of rot can be dabbed with a cotton swab, as mentioned under “Leaf Spots.” As a preventative meas-
ure, dust cut surfaces of the plant with fine sulfur or dip in a solution of general disinfectants (Physan 20
or Green Shield).
R
Rust diseases have long been known to affect wild ferns, but they are less frequently encountered on cul-
tivated ones. The disease produces orange-brown pustules or white spots on the underside of the leaf. Dis-
coloration can appear on the upper surface of the leaf opposite the pustules. Rust spores may be airborne.
Moist, cool nights and warm days favor rust. The rust Milesia is common in England on hart’s-tongue fern
(Phyllitis scolopendrium).
P C. Remove infected leaves to minimize airborne rust spores that can rein-
fect other leaves. Winter cleanup, particularly the removal of old fronds, will reduce infection in the spring.
Propiconazole (Banner), folpet (Phaltan), chlorothalonil (Daconil 2787), mancozeb (Dithane M-45),
maneb (Dithane M-22), and zineb (Diathane Z-78) are chemical sprays that kill rusts. Their effect on
ferns is not known.
S M
Sooty mold is a fungus (usually Fumago) that appears as black smutty patches on leaves. It also grows
on the honeydew drops secreted by aphids and mealybugs. Although unsightly, sooty molds do not harm
the plant. They do, however, intercept light to the fronds.
P C. Control scale insects, aphids, mealybugs, and ants. Promote air
circulation.
T B
The development of irregular, deformed fronds and a subsequent slowing or cessation of growth may
appear among young and well-established species of tree ferns (such as Dicksonia antarctica and Sphaeropteris
cooperi). Laboratory tests on diseased Dicksonia plants have tentatively identified the pathogen as the fun-
gus Phyllosticta. This fungus is sometimes called tip blight because it causes the leaf tissue to appear sun-
burned along the margins, but the brown coloration can spread throughout the leaflet. Leaflets become dis-
torted, undersized, appear moth-eaten, or are absent in places. Emerging fiddleheads can be irregularly
twisted, and leaf-stalk cross sections sometimes reveal a central area of dead, darkened tissue. This disease
has appeared in southern California, and similar symptoms have been reported in the Hawaiian Islands.
P C. Generous applications of fertilizer may retard the disease symptoms for
a time, but they eventually return. Broad-spectrum fungicides are reportedly effective against tip blight,
138 C 11
including chlorothalonil (Daconil 2787) for home growers, and iprodione (Chipco 26019), mancozeb
(Dithane M-45), and zineb (Dithane Z-78) for commercial growers. Overhead watering of tree ferns
should be avoided.
A
Of all the algae found in spore pans, the green algae and cyanobacteria (blue-green algae) cause the most
trouble. Oscillatoria (a cyanobacteria) forms a blackish slime when growing under bright light, and a gray-
ish mold-like mat under high humidity and dimmer light. It also emits a distinct odor. Studies indicate that
some toxins produced by cyanobacteria reduce respiration in mammals and seed plants, which might
account for the inhibition of sporeling growth when these algae are present. Nevertheless, some of the
T G F 139
diminished growth probably results from the cyanobacteria robbing the plant of light, crowding it, and
attracting fungus gnats. Ferns that are inherently slow growing or those growing in poor conditions in
spore pans are most susceptible to algae. By providing optimum conditions for rapid fern growth, some of
the algae problems may be averted.
P C. Practice sanitation in planting and growing areas. Sterilize the soil
before sowing spores. Use only distilled water because tap water might be contaminated with algae, and
avoid any other possible contamination sources, such as unclean watering cans or pots. If algae appear, pick
out the growth. In small plantings a light dab with a cotton swab will usually pick up the algae mat with-
out disturbing the ferns. Dry soil additives such as pumice and sawdust placed on top of soilless mixes can
reduce algae growth.
Algae can be controlled with quaternary ammonium compounds (Physan 20 or Green Shield). Sprays
of these algaecides were harmless to small plants of Cyrtomium falcatum, Dicksonia antarctica, Pteris cretica,
Sphaeropteris cooperi, and Sphaeropteris medullaris when used at a rate of 400 ppm (1.67 ml per 3.8 liters of
water, or ⅓ tsp. per gal.). With either chemical, reapplications will probably be necessary.
A pale blue solution (0.1 ppm) of copper sulfate (bluestone) used as a drench will also inhibit algae
growth. Do not allow any of the solution to get on the ferns. Various algaecides have been found to hurt
young ferns, whereas others (such as Algamycin) do not. The synthetic antibiotic chloramphenicol (Parke-
Davis or Rochelle) has been used to control cyanobacteria as a drench (at 20–40 ppm) but is not registered
for this use. Chlorothalonil (Daconil 2787) and mancozeb (Dithane M-45) are other fungicides that have
been reported as effective against some algae.
To prevent the spread of algae and to destroy the habitat of fungus gnats and other pests, the algae on
benches, walkways, and walls should be controlled. Many commercial preparations are available for this
purpose, but some are toxic to plants and must be used with care. Drenches or sprays of hydrated lime (at 681
g to 3.8 liters of water, or 1.5 lbs. per gal.) or copper sulfate (454 g to 3.8 liters, or 1 lb. per gal.) are effective.
There are about 12,000 species of ferns worldwide, but only about 30 are commonly cultivated. Most peo-
ple know these cultivated species by easy-to-remember common names that are sometimes charmingly
appropriate or even whimsical. Common names are useful when dealing with only a few species and when
local usage is uniform, but they can cause confusion if many species and people from distant lands are
involved. One reason for the confusion is that several different common names might apply to the same
plant. Phlebodium aureum, for example, is known variably as the rabbit’s-foot fern, the bear’s-foot fern, the
serpent fern, and golden polypody. Conversely, several different species might be referred to by the same
common name; the name “lace fern” is used to describe plants in five different genera (Cheilanthes gracil-
lima, Microlepia strigosa, Nephrolepis exaltata ‘Smithii’, Odontosoria chinensis, and Paesia scaberula). No
rules are established for designating the correct common name for a species. Any number of names might
exist, and new ones can be coined by anyone. Another problem with common names is that most plants
don’t have them.
Scientific names have the disadvantage of often being long, difficult to pronounce, and hard to remem-
ber. Because they are in Latin, their meaning is usually obscure to most people. But scientific names offer
several advantages over common names. They are understood internationally. A trained botanist in Indo-
nesia, Russia, or Mexico will know exactly what you mean when you utter “Dryopteris.” Using scientific
names reduces confusion and increases accuracy in communication. The other great advantage is that they
indicate a degree of evolutionary relationship. You can tell by the name, for example, that Dryopteris inter-
media and Dryopteris marginalis are more closely related to each other than either is to Polystichum acrosti-
choides. If the common names of these three species were used—glandular wood fern, marginal shield fern,
and Christmas fern—relationships are not apparent. If you deal with many species and want to be precise
when communicating with other plant people, then it is worth the time to learn about scientific names.
Every species has only one correct scientific name. The situation is complicated, however, because
many species have synonyms, or alternative scientific names, that arise as a result of new discoveries or
hypotheses about relationships. For example, in the 1800s the Massachusetts fern was called Dryopteris
simulata, but this name is now considered a synonym of Thelypteris simulata because the plant belongs
with the thelypterioid ferns instead of the dryopteroids. The change resulted from greater knowledge based
on new evidence about the relationships of the plant.
In the encyclopedic portion of this book, commonly encountered synonyms are given after the accepted
scientific name, which is the one currently used in most floras, manuals, and monographs. The synonyms
are given because they continue to be used by some gardeners and trade people. There is no mandate that
new names be immediately adopted. Sometimes they are controversial, and even professional botanists
disagree as to which name is correct. To avoid confusion, plant labels can give both names, placing the syn-
140
H F G T N 141
onym in parentheses. The word “formerly” is sometimes added before the synonym. Adding this extra
name to labels or catalog lists might be an inconvenience to the commercial growers, but gardeners are
becoming more knowledgeable about species names and appreciate greater accuracy in labeling.
Naming Species
The scientific name of a species consists of two words. The first is like the surname of a person (Doe), the
second is like the given name (Jane). The first word denotes the genus or larger group to which the plant
belongs and indicates some degree of relationship. The second word denotes a particular kind of plant in
the group. For instance, Asplenium bulbiferum is the scientific name for the mother fern. The first word,
Asplenium, is the genus to which the mother fern belongs. The second word, bulbiferum, indicates the par-
ticular kind of Asplenium. There is a similar fern called Asplenium daucifolium. It is apparent from this
name that the species is related to the mother fern because both are in Asplenium. It is also clear that the fern
is not the same species as the mother fern because it has a different second word, or specific epithet. These
two species differ not only in size and color but more significantly by the shape of the leaflets and the rhi-
zome scales (Figure 12.1). Having the same specific epithet means nothing. Botrychium virginianum and
Polypodium virginianum have the same specific epithet, but they belong to different genera and families. In
this case the specific epithet indicates that the original specimens from which these species were described
were collected in the State of Virginia.
How does a plant get a scientific name? A plant suspected as new to science is studied by a botanist who
compares it to its relatives. If no match is found, and the plant is not a freak or variant of a known species,
then the botanist names the plant according to the rules set forth in the International Code of Botanical
Nomenclature, or the Code for short.
If the plant differs greatly from known plants, the botanist might decide to give it a genus name all its
own. If, on the other hand, the botanist believes the plant belongs to an established genus, a specific epithet
is added to the genus name to complete the species name of the plant. The botanist can name the plant after
anything; it may be the name of the person who discovered it, the place where it was discovered, or a struc-
tural feature.
The botanist who names the plant is known as the author or authority. The author’s name appears after
the scientific name, although this is sometimes omitted in general use. The author’s name may be given in
full, or sometimes only the initials or a surname abbreviation are used. The author citations are important
to botanists in determining the priority and application of names. The scientific name for the bird’s-nest
fern, for example, is Asplenium nidus Linnaeus (sometimes abbreviated “L.” or “Linn.”) for the famous
Swedish botanist who began the consistent practice of giving species two names. This practice is called the
binomial system, and it permits a systematic arrangement of species into a filing system.
Figure 12.1. The fronds of the closely allied species Asplenium bulbiferum (left) and A. daucifolium (right).
C V
A cultivated variety, or cultivar, is a distinctive plant arising through selection from cultivated or wild
stock or by hybridization in cultivation. Its distinctive attributes are uniform and stable and are retained
when propagated. Whether or not the variant is distinct and worthy of being assigned a name is left to the
individual naming the cultivar to decide. Not every fern variation is named, and many are unworthy of
receiving a name. Some common variations seen in cultivated varieties of ferns include the following:
H F G T N 143
• Changes in the leaf margin, such as fringing, lobing, ruffling, cresting, or forking.
• Changes in size of the parts, such a depauperate, narrower, wider, smaller, or large pinnae or
pinnules.
• Changes in position of the parts, such as overlapping or more distant pinnae.
• Changes in overall size of the plant, such as dwarfism or gigantism.
• Changes in color, such as variegation.
• Miscellaneous changes, such as formation of buds, failure to form spores, maintenance of
juvenile foliage, and even physiological differences such as greater resistance to drought or
cold.
Figure 12.2 shows the variations possible in cultivars of the lady fern.
The name of a plant cultivar is enclosed in single quotation marks and follows the species name. Older
usage might have the abbreviation “cv.” preceding the cultivar name. Valid cultivar names given after 1959
are in non-Latin form and in roman type, not italic. The main words of the name have the first letter cap-
italized, as in Adiantum raddianum ‘Pacific Maid’. The species epithet can be omitted if there is no likeli-
hood of confusion with another cultivar bearing the same name in the genus. Thus Adiantum ‘Pacific
Maid’ is permissible.
Cultivar-groups are used to name an assemblage of two or more similar, named cultivars. (The word
“named” was added to the definition of cultivar-group in the 1995 International Code of Nomenclature for
Cultivated Plants. This precludes using the cultivar-group as a catchall category for variants belonging to the
group but unworthy of recognition. This wording is currently being met with considerable controversy.)
This category may also be used to recognize distinct groups within a species when the species itself is no
longer recognized. For instance, plants known as Davallia fejeensis have been reduced by some botanists to
D. solida. Plants known as D. fejeensis are prized by horticulturists and in this reduction to D. solida would
be left without a name. However, it may continue to be recognized by using the cultivar-group category D.
solida Fejeensis Group. If a particular cultivar in the Fejeensis Group needs to be identified, the group
name is enclosed in parentheses, followed by the cultivar name in single quotation marks, as in Davallia sol-
ida (Fejeensis Group) ‘Plumosa’.
Naming of cultivated plants is governed by rules set forth in the International Code of Nomenclature for
Cultivated Plants, commonly referred to as the Cultivated Plant Code or ICNCP (Trehane 1995). The
Cultivated Plant Code promotes uniformity, accuracy, and stability in naming horticultural plants. Grow-
ers, plant businesses, and producers of cultivated varieties should use these rules to minimize confusion. The
assignment and use of cultivar names is a voluntary matter. It should not be confused with trademark
names or patents, which are legal matters involving the government (discussed later in the chapter).
Unfortunately, despite these rules, the naming of fern cultivars in the trade and among hobbyist grow-
ers often makes it difficult to determine whether a given cultivar is new or a renamed older one. Most cul-
tivars in circulation have not been named according to the rules in the International Code of Nomenclature
for Cultivated Plants, and thus, technically speaking, have invalid names. Yet it is impossible to ignore these
names because they are an important part of the commercial trade and of many gardens. Until a Fern Reg-
istration Authority is established, the problems with the valid naming and identification of many cultivars
will be difficult to solve.
11
5 12
6
8
7 13
Figure 12.2. Variation in cultivars of the lady fern (Athyrium filix-femina): 1. ‘Caput-Medusae’; 2. ‘Foliosum
Grandiceps’; 3. ‘Grandiceps’ (dwarf form); 4. ‘Gemmatum Bolton’ (original clone); 6. ‘Acroladon’ (original clone); 7.
‘Frizelliae’ (original clone); 8. ‘Frizelliae Cristatum’ (from spore); 9 ‘Corymbiferum’; 10. ‘Pritchardii’ (original clone);
11. ‘Angustocruciatum Cristatum’; 12. ‘Fieldii’ (original clone); 13. ‘Victoriae’ (original clone);
14. ‘Victoriae Foliosum’; 15. ‘Flabellipinnulum’; 16. ‘Minutissimum Congestum’; 17. ‘Congestum Minus’; 18.
‘Crispum Coronans’; 19. ‘Veroniae’ (original clone); 20. ‘Plumosum Stansfield’ (original clone); 21. ‘Plumosum
Druery’ (original clone); 22. ‘Plumosum Furcillans’; 23. ‘Plumosum Penny’ (original clone); 24. ‘Plumosum
Multifidum’; 25. ‘Howardii’ (original clone); 26. ‘Clarissima Jones’ (original clone); 27. ‘Cristatum’; 28 ‘Clarissima
Cristatum Garnett’ (original clone). After Kaye (1965).
14
21
20
22
15
23
24 25
17
16
26
27
19
28
18
146 C 12
States there is P. aquilinum var. pubescens, the western bracken; common in the northeastern United States
is P. aquilinum var. latiusculum, the eastern bracken. Besides geographical differences, the western bracken
has pinnules nearly at right angles to the rachis and hairs on the indusia, while the eastern bracken has
obliquely placed pinnules and hairless indusia. A third and more southeastern variety, P. aquilinum var.
pseudocaudatum, the tailed bracken, has long, narrow terminal segments. A fourth variety occurs in south-
ern Florida: P. aquilinum var. caudatum, the lacy bracken, with very stiff, wiry fronds. (Some botanists now
believe var. caudatum to be a separate species and call it P. caudatum.) The abbreviations “subsp.” or “ssp.”
stand for subspecies, a category used to imply an incipient species.
Figure 12.3. Comparison of the foliage of the botanical varieties of Pteridium aquilinum (left to right): var. latiusculum,
var. pubescens, and var. caudatum.
A variation that appears sporadically among plants in nature is placed in a category known as a form
(forma or f.). Additionally, the variation should be found sporadically throughout the range of the species.
An example is Blechnum spicant f. bipinnatum. The typical form of the species (B. spicant f. spicant) has
entire pinnae, but f. bipinnatum has deeply lobed pinnae. The lobed form, which is atypical, was found
growing among typical plants in nature (Figure 12.4).
Sometimes botanical forms can be environmentally induced. Examples are the lobed types of sensitive
fern (Onoclea sensibilis) and Christmas fern (Polystichum acrostichoides). In certain individuals of these spe-
cies, abnormally deeply lobed pinnae are produced on plants that also produce typical pinnae. The deep
lobes are caused by trauma from frost, mowing, fire, or transplanting (Beitel et al. 1981; Wagner 1942).
These variants are often referred to as Onoclea sensibilis f. obtusilobata and Polystichum acrostichoides f.
incisum, but they are unworthy of taxonomic recognition. If they were named, then two different form
names would apply to the same plant—an illogical situation.
H F G T N 147
Figure 12.4. Comparison of the pinnae of the botanical forms of Blechnum spicant: forma spicant (left) and forma
bipinnatum (right).
Early use of the category “varietas” did not distinguish between cultivated varieties, botanical varieties,
and forms; hence, many plants listed as “var.” are nowadays considered cultivars or forms. Actually, the dis-
tinction between cultivated varieties, botanical varieties, and forms is poorly defined and overlapping. A few
plants might end up having both a botanical name and a cultivar name. The naming of subspecies, varieties,
and forms is governed by rules set forth in the International Code of Botanical Nomenclature.
H
A hybrid is the offspring from a cross between two different species. Fern hybrids are often sterile
because the chromosomes from the two parents cannot pair with each other during meiosis—the kind of
cell division that gives rise to spores. The result is spore abortion, and under the microscope the spores
appear irregular, misshapen, and often blackened. Ferns with aborted spores must be propagated by divi-
sion of the rhizome or by buds.
When sterile, hybrids are designated by the hybrid sign “×” placed between the genus name and spe-
cific epithet. Examples are Adiantum ×tracyi (= A. aleuticum × A. jordanii), Dryopteris ×triploidea (= D.
carthusiana × D. intermedia), and Polystichum ×potteri (= P. acrostichoides × P. braunii ). A name such as
“Adiantum ×tracyi” is the binomial; “A. aleuticum × A. jordanii” is the formula name.
Occasionally a sterile hybrid doubles its chromosome number, a process referred to as polyploidy.
Polyploidy automatically confers fertility to the hybrid. The plant can now produce normal (not mis-
shapen) spores that germinate upon sowing. Such fertile hybrids, also known as “nothospecies,” take on an
existence of their own, sometimes spreading far beyond the ranges of the parents. In this book, fertile spe-
cies of hybrid origin are given binomials without using the hybrid sign “×.” Examples of such plants are
Cystopteris tennesseensis, which arose from a cross involving C. bulbifera and C. protrusa, and Dryopteris
clintoniana, which arose from a cross involving D. celsa and D. ludoviciana.
Crosses between different genera may be placed in hybrid genera (Figure 12.6). To form the name of
a hybrid genus, the generic names of the two parents are combined, and the hybrid sign “×” is placed before
the name. For example, a hybrid between a species of Aglaomorpha and a species of Drynaria would be
placed in the hybrid genus ×Aglaonaria.
148 C 12
a × b
Figure 12.5. Hybrid of species in the same genus:
a. parent Adiantum jordanii; b. parent A.
aleuticum; c. hybrid offspring A. ×tracyi.
If naturally occurring hybrids are brought into cultivation, they must keep their hybrid name unless
they were given an additional cultivar name at the time of publication (this rarely, if ever, happens). For
example, Adiantum ×tracyi must be listed as such in horticultural use (Figure 12.5). But had this hybrid
originated in cultivation (it did not), it would be permissible to list it as Adiantum ‘Tracyi’.
a × b
Figure 12.6. Hybrid of species of different genera: a. parent Asplenium
adiantum-nigrum; b. parent Phyllitis scolopendrium; c. hybrid offspring
c ×Asplenophyllitis jacksonii. After Lovis and Vida (1969).
b
a
genus Asplenium (Figure 12.7). What distinguishes this genus are the sori and indusia, not the shape of the
frond. The sori and indusia of the two Asplenium species are long and narrow, and although longer in the
bird’s-nest fern, they are basically the same in both species. The indusium in both species is attached to the
upper fork of a vein branch, and the rhizome scales are latticed or resemble a stained-glass window
(clathrate) when observed through a hand lens. (They also all have two vascular bundles in the stipe.) Thus
the relationships between ferns is not always obvious to the naked eye.
The fern student interested in learning how to recognize ferns in general should start by studying the
genera. Start with a fern that you know, such as the maidenhair ferns of genus Adiantum. Select a fertile
frond, and look up the scientific name of the plant in the index of this book. The discussion in Chapter 13
will describe a few of the more conspicuous features that distinguish this genus from others. Locate on
your fern the features mentioned, then check your observations with the illustrations of the genus. A hand
lens of 8 to 15× magnification will assist you greatly. Once you are satisfied that you can recognize the fea-
tures mentioned, keep them in mind; the next time you see another kind of maidenhair fern, look for these
features again. Soon you will associate maidenhairs with a certain set of features. You will also recognize fea-
tures that do not belong with the maidenhairs. Select other fern genera that you know and repeat the proc-
ess: (1) look up the scientific name; (2) locate the distinguishing features on your plant; (3) check your
observations with the diagrams; and (4) examine other members of the genus for the same features to rein-
force your observations. At this point you may be able to recognize some of the more technical differences
between the species within the genus, but it is most important to concentrate on the features of the genus
as a whole. The features used to distinguish a genus are generally less diversified and less confusing than
those used to distinguish species. Besides, the species will be easier to recognize once you have become
familiar with the genera. Most of the generic features are associated with the position, shape, and type of
sorus and indusium. Also important is the rhizome, which may be erect or creeping, hairy or scaly. The
venation and the shape of the frond may also be important. Chapters 2 and 13 and the glossary discuss or
illustrate most of these distinguishing features.
13 Ferns and Fern Allies
in Cultivation
This chapter describes the ferns and fern allies commonly cultivated in the United States. It omits those spe-
cies found only in special collections and botanical gardens. Some species treated here may no longer be in
cultivation; conversely, it was impractical to include all recent introductions. A few genera not cultivated
in the United States are included because they were deemed worthy of culture, challenging to grow, or im-
portant in teaching botany. Most of the entries have been verified against botanical specimens and/or de-
scriptions; however, specimens or sufficient data were not always available, and many species are poorly
known. The correctness of the names must be considered in view of these problems.
S E
Species within each genus are listed alphabetically by scientific name. Synonyms or misapplied names
and common names, if any, follow. Cultural information is followed by a brief botanical description and
information on the geographical area to which the fern is native.
Where species important in horticulture were reduced by botanists to another species (as is the case in
the genus Davallia), the horticulturally important species name was retained with a note in the entry indi-
151
152 C 13
cating the discrepancy. Keeping such names is the least disruptive stance to take at this time. If necessary,
they may later be given cultivar status.
Only the most important synonyms are listed for a species; many synonyms not used in horticulture are
omitted. You can get an idea of the other genera under which a species might be synonymized by looking
at the generic synonyms after the genus name, or by referring to the Index of Plant Names. If the fern is
widely known under an incorrect name, this name will be noted as being “misapplied to” or as a “trade
name.”
Subspecies, varieties, and cultivars. Within the species entries, plants below the rank of species, including
subspecies, varieties, forms, and cultivars, are also discussed. Due to the tremendous number of cultivars—
there are at least a thousand—including all cultivars is beyond the scope of this book and many are omit-
ted. We have focused on those cultivars that are more common in the trade or among hobbyists and have
a fairly distinct appearance. Cultivars with names that are believed to be in agreement with the International
Code of Nomenclature for Cultivated Plants are also favored. As discussed in the previous chapter, the culti-
var names circulating in the trade are not always reliable because they are often invalid or inconsistently or
erroneously used. Where a name is known to be commonly misapplied to a particular plant, that is indi-
cated in the text.
C I
For each species, information is provided on the temperature, light, soil, and water requirements of the
plants. Ornamental attributes, including height and habit, and landscape uses are also discussed. The cul-
tural information is meant only as a guide. Often little is known about a fern’s requirements for growth, and
what is known can vary greatly with different gardens, microclimates, and genetic races of the plants.
Height. As used in this chapter, fern size refers to the height of the whole plant, not the frond length. Be-
cause some ferns vary greatly, the given heights may have exceptions. The following general terms are used
to describe fern height:
Temperature. It is difficult to give the minimum temperature tolerances of most introduced ferns. Little
data exist on the subject, and what is reported is often insufficiently documented. Knowing the tempera-
tures of the fern’s native habitat helps, but ferns often grow well in colder or hotter places than might be ex-
pected from their natural distribution. The categories of hardiness used in this book are fairly broad and may
overlap, but they offer workable categories for sorting out the hardiness of different ferns. The range of av-
erage annual minimum temperatures in the following section is based on the USDA Plant Hardiness Zone
Map (United States Department of Agriculture, Agricultural Research Service, Miscellaneous Publication
Number 1475; also see the map at the end of the color plates section). These ratings indicate the minimum
temperature range at which a plant may be expected to survive. They do not necessarily represent the op-
timal winter temperature for the plant. Also keep in mind that the timing and duration of the minimum
temperatures can profoundly affect a fern’s survival, as can many other factors. Microclimates and genetic
variations within a species often create exceptions to hardiness ratings. Two ferns might be listed as toler-
ating the same minimum temperatures, yet one might survive only with cool nights whereas the other only
with warm nights.
F F A C 153
Very hardy: Severe and extended winters, such as in alpine, tundra, taiga, and northern
coniferous forest. The range of annual minimum temperature is −46°C and below to −
29°C (−50°F and below to −20°F).
Hardy: Cold-temperate areas; tolerates long periods below freezing. The range of average
annual minimum temperature is −29 to −15°C (−20 to 5°F).
Semi-hardy: Warmer temperate areas; tolerates short periods below freezing. The range of
average annual minimum temperature is −15 to −10°C (5 to 15°F).
Semi-tender: Subtropical areas, with frost or subfreezing temperatures very rare; winter night
temperatures often reaching 4 to 10°C (40 to 50°F). The range of average annual
minimum temperature is −10 to −4°C (15 to 25°F).
Tender: Tropical areas, indoors, or greenhouses not below 16°C (60°F) at night, 18°C (65°F)
or higher during the day. The range of average annual minimum temperature is −4 to 4°C
and above (25 to 40°F and above).
Very tender: Temperatures should be maintained at about 21°C (70°F) or higher most of the
time; some species will tolerate infrequent dips to 18°C (65°F). The range of average
annual minimum temperature is 4°C and above (40°F and above).
With the dearth of data on introduced ferns, we have used broader categories of hardiness than the U.S.
Department of Agriculture’s hardiness zone map. Where USDA zones are known from reliable sources,
they are cited. The rough equivalent between the USDA zones and those used in this book are given in
Table 13.1.
Geographical area Range of average minimum temperature USDA zones Ratings in this text
Light. Because light intensity interacts with humidity and temperature, these factors must be considered
along with the light ratings given in this book. Light intensity is usually measured in units called foot-can-
dles. Estimating foot-candles by eye is difficult. See Appendix I for information on calculating foot-candles
with a camera light meter.
Low light: Dense shade, about 200 to 400 foot-candles.
Medium light: Partial shade and sun or medium shade, about 400 to 600 foot-candles.
High light: Bright shade, about 600 to 1000 foot-candles.
Direct sunlight: On a clear day, direct sunlight measures from 5,000 to 10,000 foot-candles.
Plants tolerating direct sun usually can withstand higher temperatures as well.
Soil. The following terms are used in the plant entries to categorize soil types:
Garden soil: Garden soil, preferably a loam with a lot of humus.
Potting mix: Mostly soilless mixes; must be well-drained if containing soil.
Uncut moss: Coarse, uncut, or unground moss (green moss or sphagnum moss). Used in
hanging baskets or tied to boards to give good drainage.
Drained: Provide with good drainage, which may be increased by planting on boards, hanging
baskets, or mixing material such as bark pieces, coarse perlite, or coarse sand in the
medium.
Acidic: Prefers acidic conditions (pH 4–7).
Basic: Prefers basic or limestone conditions (pH 7–8).
Water. Most ferns should be watered deeply but not too often; avoid light sprinkles. Expect the soil to be
saturated after a thorough watering. Fortunately for most plants, this saturated condition does not last
long as the extra water will drain away, the soil will start to dry, and much of the water will be replaced by
air. The basic requirement is to keep the soil at the moisture level preferred by the plant for most of the time
and not let it dry out. Water less during cool weather and more frequently during hot, dry weather.
Wet: The soil is saturated with water and the excess allowed to drain away (a condition known
as field capacity). Plants that prefer such conditions have to be watered more frequently to
maintain this level of wetness.
Moist-wet: The soil is not saturated with water but feels wet and tends to be sticky when
handled. It is a condition between those described for wet and moist.
Moist: The soil feels moist but is not so wet as to be sticky when handled. Most ferns prefer
this range of moisture.
Moist-dry: The soil feels damp but is not sticky to the hands. It is a condition between those
described for moist and dry.
Dry: The soil feels dry and is light in weight due to the low water content.
Evergreen. As used in this text, evergreen refers mainly to hardy ferns that remain green even in freezing
temperatures. These ferns grow primarily in temperate climates. Tender ferns can be assumed to be ever-
green (or nearly so) unless noted otherwise.
Deciduous. As used in this text, deciduousness indicates fronds that markedly wither and fall from the plant
or wither in place (marcescent) even if temperatures remain constant. Some subtropical or tropical species
(mainly native to seasonal climates) are deciduous at certain times of the year regardless of temperature
changes.
D F
Each species entry includes the main diagnostic characteristics, not a full botanical description. Because
these descriptions are short, the accompanying illustration of the species should be examined. Refer to the
preceding chapters and the glossary for explanations of unfamiliar terms.
Acrostichum 13.1.1, 2 sake of “acrostichoid sori,” a term used to describe sori in
Leather fern which the sporangia are spread across the lower surface of
The genus Acrostichum includes bold-looking ferns with the leaf, making the entire sorus resemble a mat of brown
large, erect fronds up to 4 m (13 ft.) tall. Although the felt. Acrostichoid sori always lack indusia. This type of
plants grow tall in the wild, they tend to be shorter in cul- sorus characterizes Acrostichum and other, unrelated genera
tivation. In tropical climates or large indoor spaces with ad- such as Elaphoglossum and Platycerium.
equate light, they can be grown in ponds or in pots kept The genus occurs throughout the tropics, thriving pri-
wet or set in water. Acrostichum is subject to damage by marily in mangroves and brackish marshes. Its name comes
scale insects and slugs. from the Greek akros, summit, and stichos, row, referring to
All three species of Acrostichum have fronds that are one- the fertile pinnae borne toward the leaf apex in Acrostichum
pinnate and leathery, with veins closely and uniformly net- aureum.
ted. The fertile pinnae are slightly narrower than the sterile
ones. The sporangia cover the entire lower surface of each Acrostichum aureum Linnaeus 13.1.3
pinna, and the sori lack indusia. Acrostichum is the name- Leather fern, coastal leather fern
Tender, thrives in Zone (9)10
A very large fern with erect rhizomes and fronds in clus-
ters. Best grown under high light in garden soil or potting
mix kept constantly wet. Acrostichum aureum can grow
with its stems submerged but is typically found rooting in
mud with the foliage held above water. It grows natively in
brackish water but can be cultivated in fresh water.
b
Figure 13.1.1. Acrostichum danaeifolium: habit.
Figure 13.1.2.
Acrostichum
danaeifolium: fertile Figure 13.1.3. Acrostichum
pinna (left) and sterile aureum: a. habit, bar = 25 cm
pinna (right). (10 in.); b. vein pattern, bar =
5 mm (0.2 in.). After Proctor
(1985).
F F A C 157
The blades of this species have up to 30 pairs of pinnae together than those of A. aureum. If the plants are fertile,
that are glabrous on their lower surfaces. The veins form another distinction can be seen: the fertile pinnae of A.
areoles along the costae, and the areoles are about three danaeifolium typically occur along the entire length of the
times longer than wide. See Acrostichum danaeifolium for a leaf (on the basal, middle, and apical portions) whereas
comparison with that species. Acrostichum aureum occurs those of A. aureum occur only toward the apex (usually the
throughout the tropical regions of the world. apical one to seven pairs). In nature A. danaeifolium grows
along coasts as well as farther inland—thus the common
Acrostichum danaeifolium Langsdorff & Fischer name inland leather fern—whereas A. aureum grows only
13.1.1, 2, 4 near the coast.
Giant fern, inland leather fern
Tender, Zone 9 Actiniopteris
A very large fern with erect rhizomes and fronds in clus- Grown as novelty plants for the attractive fan-shaped
ters. Best grown under high light in garden soil or potting fronds, Actiniopteris is a genus of small, terrestrial ferns with
mix kept constantly wet. short-creeping rhizomes and clustered fronds. Although at-
The fronds of Acrostichum danaeifolium tend to be taller tractive, the plants are difficult to grow. They seem to pre-
and more erect than those of A. aureum. The blades have fer moderately humid conditions, but excessive moisture
40–60 pairs of pinnae. The areoles formed by the veins around their roots should be avoided.
along the costae are less than three times longer than broad. Actiniopteris is characterized by fan-shaped blades re-
This species occurs only in the tropics of the New World. peatedly forked into linear segments. Along the segment
Acrostichum danaeifolium resembles A. aureum but can margins are the sori, which are protected by the enrolled
be distinguished by hairs on the lower surface of the blade. edges of the segments. In the wild the species grow in dry
The hairs are less than 0.5 mm long and are best seen with habitats, and the blades, when dry, bend toward the leaf
a hand lens. The pinnae of this species are also usually closer stalk.
The genus consists of five species and is native to Africa
and the arid parts of tropical Asia. Its name comes from the
Greek aktis, ray, and pteris, fern, alluding to the radiating
segments of the blade. For additional information on this
genus, see Pichi-Sermolli (1962) and the section on “Xero-
phytic Ferns” in Chapter 10 of this book.
Adiantopsis
This genus consists of small- to medium-sized, terrestrial or
rock-inhabiting ferns. Only one species is cultivated, grown
primarily for the attractive radiating arrangement of its pin-
nae; other species have pedate or pinnate fronds. All species
in the genus have erect or short-creeping and decumbent
rhizomes. Adiantopsis is closely related to Cheilanthes but
can be distinguished by the grooved upper surface of the
rachis and the sharp wings on either side of the groove. The Figure 13.3.1. Adiantopsis radiata: fronds, bar = 5 cm
ultimate segments are asymmetrical and eventually fall off, (2 in.).
F F A C 159
Adiantopsis radiata has blades that consist of four to nine Hoshizaki (1970a), and the section on “Maidenhair Ferns”
one-pinnate pinnae radiating from the top of the stipe. The in Chapter 10.
species occurs throughout most of the American tropics.
Adiantum aethiopicum Linnaeus 13.4.2
Adiantum 13.4.1 Semi-tender, Zone 8
Maidenhair fern A small to medium, clump-forming fern with fronds in
Most maidenhairs are small- to medium-sized, terrestrial loose-spreading clusters on medium- to short-creeping rhi-
ferns prized for their fine, billowy foliage and shiny dark zomes. Best grown under medium light in moist garden
stalks. They generally require medium light, humidity, and soil or potting mix. This species is easy to grow.
moist soil but will not tolerate soggy soil. For this reason, Adiantum aethiopicum is characterized by finely divided
keep the plants well drained, and do not overpot. Adiantum blades up to three- or four-pinnate and by ultimate seg-
is mostly tender, but a few species are hardy or semi-hardy. ments that are fan-shaped to broadly obovate, with up to
three deep notches on the distal margin. Typically each seg-
ment has one to three sori. The plants are completely
glabrous, and in fresh material the base of the segment
stalks have an inconspicuous pale green swelling. The in-
dusium is kidney-shaped with a deep, C-shaped sinus. The
species is native to South Africa, Australia, New Zealand,
and New Caledonia. The specific epithet aethiopicum sug-
gests that the species occurs in northern Africa; however, in The bluish green fronds are pedate with one-pinnate
Africa it occurs only near the Cape. branches. The blade is fan-shaped or funnel-shaped in out-
Most of the plants sold as Adiantum aethiopicum in the line. The medial segments are more than three times longer
United States are actually A. raddianum ‘Triumph’, which than they are broad or somewhat long-triangular or reni-
can be distinguished by its lack of a swelling at the base of form, their tips are sharply denticulate-lobed, and the lobes
the segment stalk and by veins that end in minute sinuses are separated by deep sinuses, 0.6–4.0 mm deep. Adiantum
instead of teeth. Adiantum capillus-veneris, a similar spe- aleuticum is indigenous to the western United States and
cies, differs by oblong indusia and the absence of swellings western Canada.
on the segment stalk base. This species is very close to Adiantum pedatum, an east-
ern United States species that differs in having a rounder
Adiantum aleuticum (Ruprecht) C. A. Paris blade outline and shorter medial segments with rounded,
13.4.3 crenulate or crenulate-toothed lobes. The lobes are sepa-
syn. Adiantum pedatum subsp. aleuticum Ruprecht rated by shallow sinuses (mostly 0.1–2.0 mm deep).
Western five-finger fern In the wild this species often grows on serpentine, a
Hardy, Zone 4(5) metamorphic rock that is often green and slippery to the
A small to medium fern with fronds in clusters on a clump- touch. Serpentine has several traits inimical to plant
forming rhizome. Grows in low to medium light in moist growth. It is low in essential nutrients, such as nitrogen,
potting mix. Adiantum aleuticum can be difficult to grow in calcium, potassium, and phosphorous, and high in toxic
arid climates. elements such as nickel and chromium. Its pH may be
either highly acidic or basic. Soils derived from serpentine
are often sterile and support unusual endemic plants.
subsp. subpumilum (W. H. Wagner) Lellinger (forma
imbricatum Hoshizaki; ‘Imbricatum’). Dwarf western five-
finger fern. Smaller than the typical subspecies, with
densely overlapping leaflets.
c
b
Adiantum diaphanum is characterized by smooth stipes, Adiantum flabellulatum Linnaeus 13.4.13
and blades that are pinnate to some having one pair of pin- Tender or slightly hardier
nae at the base. The membranous segments bear black or A small to medium fern with fronds in a loose cluster from
reddish brown hairs on the lower surface. The indusium a clump-forming rhizome. Grows under medium light in
varies from round to nearly kidney-shaped and usually has potting mix kept moist. This species is easy to start from
dark hairs on the surface. An extremely unusual character- spores.
istic is the small barrel-shaped tubers borne by the roots.
The species is native to Asia, Australia, New Zealand, and b
the Pacific Islands. It is a delicate fern that in nature usually
thrives near water.
b b
Figure 13.4.15.
Adiantum fragile:
a. frond, bar = 5 cm
(2 in.); b. leaflets
a shedding, bar = 1 cm
a (0.4 in.).
pinnate with a terminal pinna resembling the lateral ones; Some young plants of Adiantum hispidulum are circu-
fertile pinnules with tips mostly straight and obtuse, not lating as Adiantum ‘Birkenheadii’, a name originally used in
falcate; and round apices. The blade features often overlap early English horticultural literature for a plant thought to
with A. tetraphyllum, which is distinguished by a long- have originated from A. diaphanum.
creeping rhizome and more falcate segments. Adiantum
fructuosum is native to the American tropics. Adiantum jordanii K. Müller 13.4.18
California maidenhair
Adiantum hispidulum Swartz Semi-hardy to semi-tender
13.4.17; 9 A small to medium fern with fronds in a cluster from a
Rosy maidenhair, rough maidenhair clump-forming rhizome. Best grown under high light in
Semi-hardy or hardier, Zone 7 moist-dry garden soil or potting mix. The leaves usually
A small fern with clustered fronds from a clump-forming wither in summer, and the soil should be kept drier at this
rhizome. Grows in low to high light in moist to moist-dry time. This species is difficult to grow.
garden soil or potting mix. Adiantum hispidulum tolerates
drier conditions than other maidenhairs. The plants are
easy to cultivate and frequently seen in the United States
trade. Young fronds are reddish.
Adiantum hispidulum is characterized by hairy stipes, pe-
date blades that are firm and have whitish hairs on the sur-
face, and rounded indusia with slender, reddish brown hairs
on the surface. The species is native to southern India, east-
ern Africa, and the Pacific Islands. It has escaped from cul-
tivation in the United States (Connecticut and Georgia). b
The name Adiantum pubescens Schkuhr applies to a vari-
ant with thinner and laxer hairs.
b a
b
b
b
a
a b
Figure 13.4.25. Adiantum pedatum: frond, bar = 5 cm Figure 13.4.26. Adiantum pentadactylon: a. frond, bar =
(2 in.). 5 cm (2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).
170 C 13
b
Figure 13.4.31. Adiantum pulverulentum: a. frond, bar =
5 cm (2 in.); b. fertile leaflet, bar = 5 mm (0.2 in.).
b
b
Figure 13.4.32. Adiantum pyramidale: a. frond, bar = Figure 13.4.33. Adiantum raddianum: a. frond, bar =
5 cm (2 in.); b. fertile leaflet, bar = 5 mm (0.2 in.). 5 cm (2 in.); b. fertile leaflets, bar = 1 cm (0.4 in.).
Adiantum reniforme Linnaeus 13.4.35 Figure 13.4.36. Adiantum seemannii: a. frond, bar =
Tender to semi-tender 5 cm (2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).
A small fern with fronds clustered from a short-creeping
rhizome. Grows under low to medium light in moist pot-
ting soil. This species is usually difficult to grow.
are usually several sori per pinna. This species resembles A.
Adiantum reniforme has small, kidney-shaped fronds
anceps, but its blade is whitish green (glaucous) beneath.
with entire margins. It is native to Africa (Kenya), Madeira,
Adiantum seemannii is native from Mexico to Panama and
the Canary Islands (Tenerife), and Comoros.
Colombia.
b
a
a
Figure 13.4.38. Adiantum tenerum cultivars: Figure 13.4.39. Adiantum tetraphyllum: a. frond, bar =
a. ‘Farleyense’; b. ‘Fergusonii’. Bar = 5 cm (2 in.). 5 cm (2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).
F F A C 175
b
Adiantum trapeziforme Linnaeus 13.4.41
Diamond maidenhair
Tender
A medium-sized fern with clustered fronds from a short-
creeping rhizome. Grows under low to medium light in
moist potting mix.
Adiantum trapeziforme is characterized by triangular Figure 13.4.42.
blades up to four-pinnate, with a terminal pinna similar to Adiantum venustum:
the lateral ones. The segments are trapeziform and stalked, a. frond, bar = 5 cm
with the stalks 4–7 mm (0.2–0.3 in.) long. The dark stalk (2 in.); b. fertile
a
color ends abruptly at the segment blade. This species is leaflets, bar = 1 cm
native to Central America and the West Indies. (0.4 in.).
176 C 13
b
Figure 13.5.6. Aglaomorpha
drynarioides: fertile frond, bar
= 10 cm (4 in.).
Figure 13.5.5.
Aglaomorpha
coronans: a. frond,
bar = 10 cm
(4 in.); b. part of
fertile pinna, bar
= 2.5 cm (1 in.).
a a
Figure 13.5.8. dle-like hairs. The soral patches resemble those of A. splen-
Aglaomorpha dens, but the latter species differs by having larger fronds,
meyeniana: a. fertile shorter-creeping rhizomes, humus-collecting frond bases,
frond, bar = 10 cm and hairless sporangial capsules. Aglaomorpha pilosa is na-
(4 in.); b. part of tive to the Philippines.
fertile pinna, bar = Some plants circulating in gardens are misidentified as
2.5 cm (1 in.). Aglaomorpha brooksii Copeland.
brown and papery at the base but remain green and folia-
ceous above. The blades are pinnatisect with apices pin-
nately lobed to a small terminal segment (or the terminal
segment is absent). The pinnae are connected by a thin,
cartilaginous margin, and each pinna is slightly narrowed
above the broadly adnate base. The fertile pinnae resemble
the sterile ones and are not narrowed as in most Aglaomor-
pha species. The sori vary from round to sublinear, with
one or two sori united or separated and placed between the
lateral veins.
This plant is a hybrid of horticultural origin and does
not grow in the wild. Its parental species belong to the gen-
era Aglaomorpha and Drynaria; thus the generic name of
the hybrid is a compounding of those two names. In ac-
cordance with the International Code of Nomenclature for
Cultivated Plants, the hybrid is also assigned the cultivar
name ‘Santa Rosa’. Santa Rosa Tropicals, California, was
the nursery that first tissue cultured the plant.
‘Starburst’. The pinna margins are sharply and narrowly
incised; less robust than ‘Santa Rosa’. Figure 13.6.2. Alsophila tricolor: frond.
Anemia phyllitidis has one-pinnate fronds with a termi- These ferns bear fronds up to 7 m (23 ft.) long, although
nal pinna that resembles the lateral ones. The veins are net- the fronds tend to be smaller in cultivation. The roots and
ted. This species is native to tropical America. stems are usually thick and fleshy, often massive; stems
sometimes slender and creeping. The middle of the stipe
Anemia rotundifolia Schrader 13.7.5 often has a swelling or bump. The blades vary from one- to
Semi-tender
A medium-sized fern with erect rhizomes and buds at the
tips of the fronds. Grows best under medium light in moist
garden soil or potting mix.
The fronds of Anemia rotundifolia are arching and one-
pinnate with roundish pinnae, and the rachis is greatly ex-
tended, whip-like, and roots at the tip. This species is native
to Brazil.
Figure 13.8.3. Angiopteris evecta: swollen pinna bases. Angiopteris angustifolia C. Presl 13.8.5
Tender
Very large, with erect, stout, massive stems and whorled
two-pinnate and often have a swelling at the juncture of fronds. Best grown under medium light in well-drained
the rachis and pinnae. The sori consist of a double row of garden soil or potting mix. Angiopteris angustifolia prefers a
free (unfused) sporangia, and an indusium is absent. humid environment; do not permit the soil to dry.
Angiopteris resembles the closely related Marattia, and The pinnules are 12–20 mm (0.5–0.75 in.) wide and
sterile specimens are difficult to assign to either genus. have translucent false veins that run from the margin to
Plants from both genera have fleshy flaps of tissue called nearly the midrib of the pinnule. This species occurs from
stipules on either side of the stipe base. The stipules hug Malaysia to the Philippines.
the swollen base of the fleshy stipe. The genera have similar
cultural requirements.
New plants can be propagated from stipules cut from
the stipe. The stipules should be planted in a moist, loose
medium (gravel, perlite, sphagnum, or a loose, well-drained a
planting mix) at a slant or flat on the medium with either
side up. Some growers notch the thin edges slightly to en-
courage plantlet formation at these points; otherwise,
plants tend to form at the base of the stipule, usually one on
each side. Apply bottom heat at 27°C (80°F) to hasten the
appearance of buds, which may take several months or up
c
to 2 years to appear; large stipules are slower to produce
buds. The buds should be kept moist, but not so wet as to
rot. After the buds produce roots and two or three leaves,
they can be transplanted. Angiopteris can also be propagated
by offshoots (more commonly produced on young plants)
or spores. On all established plants, new roots that emerge
from the stem base should be kept moist with a covering of
mulch. For more information, see Hill (1984).
Figure 13.8.4.
Angiopteris evecta:
stipules around Figure 13.8.5. Angiopteris angustifolia: a. pinna, bar =
bases of leaf stalks, 5 cm (2 in.); b. pinnule, bar = 2 cm (0.8 in.); c. veins
bar = 5 cm (2 in.). with long false vein (left) and sporangia (right), bar =
1 cm (0.4 in.).
186 C 13
Angiopteris evecta (G. Forster) Hoffmann powder on the lower surface of the blades. The life cycle of
13.8.1–4, 6 Anogramma is highly unusual because the gametophytes
Mule’s-foot fern are perennial and the sporophytes are annual. In contrast,
Tender to marginally semi-tender the gametophytes of all other ferns are ephemeral, dying
Very large, with erect, stout, massive stems and whorled after the sporophyte is formed. Thus, the leaves of Ano-
fronds. Best grown under medium light in well-drained gramma plants live less than a year and reappear the fol-
garden soil or potting mix. The plants prefer a humid en- lowing growing season.
vironment; do not permit the soil to dry. Anogramma grows in tropical and warmer temperate
Angiopteris evecta is characterized by pinnules that are areas throughout the world and consists of five species. The
about 25 mm (1 in.) wide and translucent false veins that genus name comes from the Greek ano-, upward, and
extend less than halfway from the margin to the midrib. gramme, line, referring to the elongate sori on the distal seg-
The species occurs from Malaysia to Polynesia. One of the ments.
most spectacular plantings of A. evecta can be seen in the
Foster Botanical Garden in Honolulu, Hawaii. Anogramma chaerophylla (Desvaux) Link
13.9.1–3
Semi-tender
a
A small fern with erect rhizomes. Grows best under me-
dium light in moist potting mix. Volunteer plants appear in
spring, densely and luxuriantly, and usually wither within
a few months. In terrariums it reproduces readily from
spores.
c
fertile fronds are taller and more contracted than the sterile.
This species is native to Japan and China and has become
naturalized in the southeastern United States.
b A variegated form sometimes circulates under the mis-
applied name of Arachniodes aristata ‘Variegatum’.
c
a
b b
related to Pellaea but can usually be distinguished by the Argyrochosma dealbata is distinguished from A. jonesii
presence of a waxy powder (farina) on the lower surfaces of and A. microphylla by the white powder on the lower sur-
the blades (this is absent in Argyrochosma jonesii and A. mi- face of the blades. It is further characterized by dark reddish
crophylla) and other technical characteristics. brown, shiny stipes and triangular blades two- to three-pin-
The blade surfaces of Argyrochosma lack hairs or scales. nate and blue-green on the upper surface. The dark color of
The segments are untoothed and distinctly stalked. The the stalk enters the segment bases and merges with the
sporangia run along the veins, usually covering the outer green of the blades. The ultimate segments are round to
one-third to two-thirds of the segments. The fertile margins ovate-oblong, 3 mm (0.1 in.) or less long, and their margins
are flat or slightly enrolled and have the same color and tex- enroll, covering the dark sporangia that are borne on the
ture as the rest of the blade—that is, the fertile margins are distal third of the secondary veins. The species is native to
not texturally modified like those of some other cheilan- the south-central United States, where it grows on calcare-
thoid ferns. ous cliffs and ledges.
About 20 species occur in the genus, which is entirely
native to the New World. Its name comes from the Greek Argyrochosma jonesii (Maxon) Windham
argyros, silver, and chosma, powder, referring to the whitish 13.11.2
powder found on the lower blade surfaces of most species. syn. Cheilanthes jonesii (Maxon) Munz, Notholaena
jonesii Maxon
Argyrochosma dealbata (Pursh) Windham Jones’s cloak fern
13.11.1 Semi-hardy, Zone 8(9)
syn. Cheilanthes dealbata Pursh, Notholaena dealbata A small fern with compact, erect to ascending, usually un-
(Pursh) Prantl branched rhizomes and clustered fronds. Best grown under
Powdery cloak fern high light in drained, basic(?) garden soil kept moist-dry to
Hardy, Zone 5 dry.
A small fern with compact, erect to ascending, usually un-
branched rhizomes and clustered fronds. Apparently prefers
medium light with basic(?), moist-dry to dry, drained gar-
den soil with sand.
b b
Figure 13.11.1.
Argyrochosma dealbata:
a. frond, bar = 2.5 cm
a (1 in.), after Tryon Figure 13.11.2. Argyrochosma jonesii:
(1956); b. segments a. frond, bar = 2.5 cm (1 in.);
with sori and indusia, b. fertile segment, bar = 1 mm.
bar = 1 mm. After Tryon (1956).
190 C 13
Argyrochosma jonesii has dark brown stipes and shiny, four-pinnate, glaucous (but not white-powdery) blades;
ovate-lanceolate, leathery blades that are two- to three-pin- and rachises grooved on the upper surface. The segment
nate. The dark color of the segment stalks extends into the margins are enrolled, and the sporangia are borne on the
base of the segments and merges with the green of the outer third of the veins. The species is native to the south-
blade; the stalks are not jointed to the segments, nor does western United States and northern Mexico, where it in-
the stalk color stop abruptly as in A. microphylla. The lower habits rocky, limestone hillsides and cliffs.
surface of the blade lacks white powder and has flat or
slightly enrolled margins, with sporangia borne on outer Arthropteris 13.12.1, 2
third to half of the veins. The species is native to the south- Joint fern
western United States and Mexico. Some species of Arthropteris look like a small Thelypteris
with a long-creeping rhizome. Nevertheless, one species,
Argyrochosma microphylla (Mettenius ex Kuhn) Arthropteris tenella, is distinct for its small- to medium-
Windham 13.11.3 sized, one-pinnate fronds. This small fern is handsomely
syn. Cheilanthes parvifolia (R. M. Tryon) Mickel, displayed in baskets or by training it to grow up the trunks
Notholaena parvifolia R. M. Tryon of trees. The latter growth habit can be started by planting
Small-leaved cloak fern the fern in the soil around the tree and training the rhi-
Semi-hardy, Zone 7 zomes up the trunk, holding them in place with string or
A small fern with compact, erect to ascending, usually less visible plastic fishing line. A collar of uncut sphagnum
unbranched rhizomes and clustered fronds. Grows under moss placed around the base of the tree encourages rooting
high light in basic(?), moist-dry, well-drained garden soil and more vigorous growth.
with sand. These terrestrial or epiphytic ferns have long-creeping,
Unlike the previous two species, Argyrochosma micro- slender, scaly rhizomes, and the stipes are jointed. The joint
phylla has its ultimate segments jointed to their stalks. This is slightly raised and sometimes distant from the stipe base,
can be seen by the dark color of the stalk stopping abruptly often appearing as a faint dark line. Blades vary from one-
at the segment base, not passing into the segment and pinnate to one-pinnate-pinnatifid and have sessile pinnae
merging with the green of the blade. The species is further jointed to the rachis. The veins are either simple or forked,
characterized by brown stipes; triangular to ovate, three- to never netted. The round sori are covered by rounded, kid-
ney-shaped indusia (this is absent in some species). The
genus resembles certain species of Thelypteris but can be
distinguished by the tiny, jointed, usually reddish (when
dry) hairs on the upper surfaces of the rachises and costae;
Arthropteris does not bear needle-shaped hairs, as is often
the case in Thelypteris. Another distinction is that
b Arthropteris stipes have three or more vascular bundles,
whereas Thelypteris stipes have only two.
a Figure 13.11.3.
Argyrochosma microphylla:
a. frond, bar = 2.5 cm
(1 in.); b. segment with
sori and indusia, bar =
1 mm. After Tryon
(1956).
This genus is native to the Old World tropics, Japan, A similar species, Arthropteris orientalis (J. F. Gmelin)
Polynesia, and temperate areas of Australia and New Zea- Posthumus, is not yet known in cultivation but has been
land. One species occurs in South America (Juan Fernán- confused with A. monocarpa. It differs by having stipes
dez Islands); the remaining 12 to 15 species are found in the jointed in the upper (distal) half, one to eight sori per lobe,
Old World. The genus name comes from the Greek and the presence of whitish (not dark) hydathodes on the
arthron, joint, and pteris, fern, alluding to the jointed stipes upper surface of the blade. The joint on the stipe appears as
of some species. a faint dark line and is easily overlooked. This species is also
native to tropical Africa. Introduced plants came from
Arthropteris monocarpa (Cordemoy) C. Christensen Zimbabwe.
13.12.3
Semi-tender to semi-hardy Arthropteris tenella (G. Forster) J. Smith ex Hooker f.
A small to medium fern with medium- to long-creeping 13.12.1, 2, 4
rhizomes. Best grown under medium light in moist-dry syn. Polypodium tenellum G. Forster
garden soil or potting mix. The plants can be used as a Joint fern
ground cover or confined in a wide container. They can Semi-hardy
also be used over logs or tree trunks if the substrate is suffi- A small to medium fern with medium- to long-creeping
ciently moist. The fronds are deciduous. rhizomes. Best grown under low to medium light in moist
Arthropteris monocarpa is characterized by stipes jointed to moist-dry garden soil or potting mix or uncut moss. The
in the lower (basal) half, one-pinnate-pinnatifid blades, usu- plants are versatile: they can form handsome displays if
ally one sorus per lobe, and usually dark hydathodes on the trained to grow up tree trunks, or they can be grown in bas-
vein tips of the upper surface of the blade. The indusium is kets or as a ground cover. Any divisions should be made
kidney-shaped. This species is native to tropical Africa. during periods of active growth.
192 C 13
Aspidotis
Lace fern
These small ferns typically grow in dry, rocky areas. Aspi-
dotis species are sometimes included in Cheilanthes, though
they can be distinguished by their elongate, distally toothed
segments with shiny, striated upper surfaces and broad, pa-
pery indusia. See Cheilanthes for a comparison with related
genera. Aspidotis consists of four species, three of which
occur primarily in western North America, and the re-
maining one in Mexico. The generic name comes from the
Greek word aspidotes, shield-bearer, in allusion to the
shield-like false indusia.
a
c
Figure 13.14.7.
Asplenium auritum:
frond, bar = 5 cm (2 in.).
c
scales on the lower surfaces. The indusium is inconspicu-
ous. The species is widespread in tropical America, where it a
typically grows as an epiphyte on tree trunks.
Asplenium australasicum (J. Smith) Hooker Asplenium bulbiferum G. Forster 13.14.9
13.14.8 Mother fern
syn. Asplenium australis Horticulture Semi-hardy to semi-tender
Semi-tender A medium-sized fern with short-creeping to suberect rhi-
A medium to large fern with erect, stout, unbranched rhi- zomes and pale green, arching foliage. Grows under low to
zomes. Grows well under medium light in well-drained, medium light in moist potting mix. This species is recom-
moist to moist-dry potting mix or uncut moss. The plants mended for places with low light. It is easy to grow and can
do particularly well in pots, but be careful not to overpot. be easily propagated from the buds on the fronds; however,
Collectively, the fronds form a funnel or bowl. This species it has the disadvantage of being particularly susceptible to
tends to be more cold hardy and less vulnerable to slugs attack by slugs and snails. Narrowly segmented forms prop-
than Asplenium nidus, which it resembles. agated from spores do not breed true.
The fronds of Asplenium australasicum vary greatly in Asplenium bulbiferum is characterized by highly divided
outline but are generally oblanceolate. They differ from blades (mostly two-pinnate-pinnatifid but sometimes up
those of A. nidus by having keeled undersides of the mid- to nearly four-pinnate) that may bear numerous buds on
ribs, and the sori are usually spread along more than half the the upper surfaces. The blades are oblong-ovate and thick.
length of the vein. The species is native to Australia and the Each incurved lobe of the pinnules contain a single sorus.
Pacific Islands. The species is native to India, Malaysia, Australia, New
Whether fronds are arranged in a narrow funnel or wide Zealand, and the Pacific Islands.
bowl is not consistent in Australian plants or those in cul-
tivation. The most reliable feature for separating Asplenium Asplenium ×crucibuli Horticulture 13.14.10
australasicum from A. antiquum and A. nidus is the keeled syn. ×Asplenosorus crucibuli Horticulture
rather than rounded midribs. Although infrequently Hardy to semi-hardy, Zones 5
grown, A. serratum also has simple leaves and a keeled mid- A small fern with erect rhizomes. Grows best under low to
rib, but it can be distinguished because its veins are free at medium light in moist garden soil or potting mix. These
the tips, not connected. evergreen plants are excellent for rock gardens.
F F A C 197
Figure 13.14.13.
Asplenium
×ebenoides:
fronds, bar =
5 cm (2 in.).
Figure 13.14.25.
Asplenium
prolongatum: habit,
bar = 5 cm (2 in.).
The plant called Asplenium mayii in horticulture is the The fronds of Asplenium thunbergii are narrowly oblong,
same as this species. In 1894 the original young plant of A. up to two-pinnate near the base, with numerous pinnae di-
mayii was presumed to be a horticultural hybrid of A. bap- vided into narrowly oblong, one-veined segments. There
tistii and A. pteroides, a species sometimes confused with A. is one sorus per segment. The rachis is broadened by nar-
surrogatum. row wings on both sides, and buds form at the rachis-costa
junctions. The species is native from southeastern Asia to
Asplenium thunbergii Kunze 13.14.31 Indonesia and Borneo.
syn. Asplenium belangeri (Bory) Kunze
Tender Asplenium trichomanes Linnaeus
A medium-sized fern with suberect, branched, clumped 13.14.32; 13
rhizomes and bud-bearing fronds. Grows under medium Maidenhair spleenwort
light in moist, well-drained garden soil or potting mix. The Very hardy, Zone 2?
plants are readily attacked by slugs and snails. Damage from A small fern with erect rhizomes. Grows under low light in
these pests can be lessened by growing the plants in hang- moist potting mix. Some plants adapt more easily to culti-
ing pots. The species is easily propagated by leaf buds. vation than others. Plants from China have adapted well
to gardens in southern California. Asplenium trichomanes
is an evergreen fern. The fronds often grow in a rosette pat-
tern.
Asplenium trichomanes is characterized by dark rachises
and one-pinnate blades with rounded to oblong pinnae. It
resembles A. resiliens, but the pinnae are less elongate and
the blade tissue is thinner and a brighter green (not dull
blue-green). A cosmopolitan species, A. trichomanes occurs
on every continent but mainly in temperate areas.
An infrequently seen but closely related species is Asple-
nium tripteropus Nakai, from eastern Asia, which differs by
rooting at the frond tip.
In North America Asplenium trichomanes consists of two
chromosomal races: one with two sets of chromosomes
(diploid) and another with four (tetraploid). The two races
differ in habitat, with the diploids tending to occur on
acidic rocks such as sandstones, granites, and basalts, and
the tetraploids occurring on basic rocks such as limestones
and dolomites (Moran 1982).
‘Cristatum’. A crested form.
‘Incisum’. Pinna margins incised.
Figure 13.14.31.
Asplenium thunbergii:
frond, bar = 5 cm (2 in.).
206 C 13
Astrolepis
Star-scale cloak fern
The genus Astrolepis consists of small to medium ferns that c
typically grow in dry, rocky places. They tolerate dryness
and make excellent rock garden plants.
The blades of star-scale cloak ferns are linear to linear-
oblong, one-pinnate to pinnate-pinnatifid, and somewhat
leathery. The segments are shallowly lobed or entire and
covered with star-shaped scales on the upper surface. The
sporangia run along the veins and lack an indusium, the
segment margins being more or less flat, not strongly en-
rolled, and of the same color and texture as the rest of the
blade tissue (not differentiated as in other cheilanthoid
ferns). b
Until recently, the species of Astrolepis were classified
with either Notholaena or Cheilanthes; however, they can
be separated from these genera by the star-shaped scales on a
the upper surfaces of the blades and the two vascular bun-
dles in the stipe (the other genera have only one). See Chei-
lanthes for a comparison to that genus and other cheilan-
thoid ferns.
Astrolepis is native to the Americas and contains about
eight species. The common and generic names refer to the
star-shaped scales on the upper surfaces of the blades: astro Figure 13.15.1. Astrolepis beitelii: a. frond, bar = 2.5 cm
is Greek for “star” and lepis means “scale.” (1 in.); b. upper surface of lobe with scales, bar = 1 mm,
by H. Fukuda; c. star-shaped scale, bar = 1 mm, by H.
Fukuda. Courtesy of John T. Mickel.
F F A C 207
Figure 13.16.8.
Figure 13.16.6. Athyrium filix-femina var. asplenioides: Athyrium filix-
a. frond, bar = 5 cm (2 in.); b. pinnule with elongate sori femina var. filix-
and indusia, bar = 3 mm (0.1 in.), courtesy of Knobloch femina: frond, bar
and Correll (1962); c. rhizome, bar = 2 cm (0.8 in.), = 5 cm (2 in.).
courtesy of Knobloch and Correll (1962).
F F A C 211
Plants of Athyrium niponicum are often misidentified in Athyrium otophorum is characterized by broad, triangu-
the trade as A. iseanum Rosenstock or its synonym A. goe- lar, two-pinnate-pinnatifid, slightly leathery fronds with
ringianum (Kunze) T. Moore. Athyrium iseanum is not long, narrowly triangular pinnae. The stipe, rachis, and
known in cultivation in the United States. costa are reddish tinged. The species is native to Japan,
‘Pictum’. Plate 14. Japanese painted fern. A variegated Korea, and China.
form with reddish and gray-silver colors. The colors de- Athyrium epirachis (H. Christ) Ching, a species from
velop best in high light. Crested, ruffled, and other forms China and Japan with similar red-tinged rachises and
have been reported. The ghost fern, a suspected hybrid be- costae, has been introduced into cultivation. It differs from
tween ‘Pictum’ and Athyrium filix-femina, is becoming A. otophorum by narrower, less foliaceous blades and erect
available commercially and shows great horticultural prom- rhizomes. It is a more vigorous grower and more evergreen
ise. Its medium-sized leaves are a dull, “ghostly” grayish than A. otophorum.
green.
Athyrium vidalii (Franchet & Savatier) Nakai
Athyrium otophorum (Miquel) Koidzumi 13.16.12
13.16.11; 15 Semi-hardy to hardier, Zone 6
Hardy, Zone 4(5) A medium-large fern with ascending to suberect rhizomes
A medium-sized fern with short-creeping, clump-forming and clustered, deciduous fronds. Grows well under me-
rhizomes and deciduous fronds. Grows under medium dium light in moist garden soil or potting mix.
light in moist garden soil or potting mix. Athyrium vidalii is characterized by triangular, two-pin-
nate to two-pinnate-pinnatifid blades, sessile to short-
stalked pinnae, spines present on the costule or base of the
pinnule midribs (upper surface), and elliptic to oblong sori.
The indusium is persistent, conspicuous, lunate, hooked
or horseshoe-shaped, with margins irregularly toothed or
entire. The species is native to China, Japan, Korea, and
Taiwan.
b
b
a
a
nitrogen enters the plant by the action of a cyanobacterium Azolla pinnata is broadly triangular and regularly pin-
(blue-green algae), Anabaena azollae, that lives within the nately branched. The roots are branched and feathery, un-
leaves of Azolla. The cyanobacterium combines, or “fixes,” like the other species in the genus, all of which have un-
atmospheric nitrogen with hydrogen to make ammonia, a branched, nonfeathery roots. It occurs in tropical Asia,
molecule that can be taken up by organisms and used to Africa, Australia, New Zealand, and New Caledonia.
form nitrogen-containing compounds. Besides its use as
fertilizer, Azolla is also fed to livestock and certain fish. For Belvisia
a popular account of the economic importance of Azolla, Members of this genus are small- to medium-sized, epi-
see Moran (1997); more technical accounts can be found in phytic ferns with undivided, narrow fronds bearing a fertile
Moore (1969), Lumpkin and Plucknett (1982), van Hove spike at the tip. They are infrequently cultivated but are
(1989), and Wagner (1997). generally grown in greenhouses or, in tropical areas, out-
doors in pots or baskets. The gracefully arching fronds are
Azolla caroliniana Willdenow 13.17.3a closely spaced and best displayed in an elevated position.
Hardy, Zone (4)5; survives frost and ice The most distinctive characteristics of Belvisia are the
Small-sized. Prefers high light. This floating fern is the best simple leaves with narrowed, tail-like fertile tips. The rhi-
species for growing on mud. The plants turn crimson in zome is short- to long-creeping, unbranched, and covered
the fall, and several weeks later they die and sink to the bot- by black or red-brown, clathrate scales. The veins are netted
tom. with irregular areoles containing forked veinlets. The sori
Azolla caroliniana plants are roughly circular to trian- lack indusia and are spread on the lower surface of the tail-
gular and about the size of a dime. The largest hairs (papil- like tip. Minute, scale-like paraphyses are mixed among the
lae) on the upper surfaces of the leaves are two-celled. The sporangia.
megaspores lack angular bumps or pits and are densely cov- The genus, which consists of eight species, occurs in the
ered with tangled filaments. Compared to A. filiculoides, Old World tropics, northeastern Australia, and Polynesia in
the clumps of A. caroliniana tend to fragment more often tropical and warm-temperate climates. It is named for
and are more compact and rounder in outline. The species A. M. F. J. Palisot de Beauvois (1752–1820), a French trav-
occurs in the Americas, Europe, and Asia. eler and agrostologist (grass specialist) whose Latinized
Azolla mexicana C. Presl is less hardy but may be in cul- name is Belvisius. For a technical treatment of Belvisia, see
tivation. It differs by having megaspores sparingly covered Hovenkamp and Franken (1993).
with tangled filaments.
Belvisia mucronata (Fée) Copeland 13.18.1
Azolla filiculoides Lamarck 13.17.1, 2, 3b syn. Belvisia callifolia (H. Christ) Copeland
syn. Azolla filiculoides var. rubra (R. Brown) Strasburger, Tender
A. rubra R. Brown A medium-sized fern with short-creeping rhizomes. Best
Hardy, Zone (6)7 grown under medium light in well-drained, moist potting
Small-sized. Grows best in high light. This floating species mix or uncut moss.
forms dense mats on the water’s surface. Growth peaks in The blades of Belvisia mucronata vary tremendously, es-
late spring and noticeably declines in winter. The plants pecially in width, ranging mostly from 1 to 5 cm (0.5–2
can survive under thin ice. Most of the cultivated material in.) wide, and they are gradually narrowed below the fertile
in southern California is Azolla filiculoides. spike. The rhizome scales have dentate to ciliate margins,
Clumps of Azolla filiculoides are less likely to fragment and the cells of the main body of the scales are all thick.
than those of A. caroliniana, and they have a more open, The species is native to Sri Lanka, southeastern Asia, Ma-
branched aspect. The largest hairs on the upper leaf lobe laysia, Australia, and the Pacific Islands.
are one-celled. The megaspores are warty with raised an-
gular bumps. This species occurs in the Americas, Europe, Blechnum 13.19.1, 2
northeastern Asia, southern Africa, and the Pacific Islands. syn. Lomaria
Most Blechnum species are terrestrial plants with leaves that
Azolla pinnata R. Brown 13.17.3c; 16
are reddish when young but turn green with age. Nearly all
Ferny azolla
the species have pinnatifid or one-pinnate blades; never are
Tender
they finely cut. The trunk-forming species, such as Blech-
Small-sized. Best grown under high light. This floating fern num brasiliense and B. gibbum, are particularly valued in
has been used for centuries in China and Vietnam as a pots for accent use. The species with creeping rhizomes are
green manure for growing rice. The federal and some state useful as ground covers or bedding plants. Blechnum aus-
governments list Azolla pinnata as an aquatic weed and pro- trale, B. penna-marina, and B. spicant are favored in warm-
hibit its import, possession, and distribution. temperate regions, whereas B. appendiculatum is used in
F F A C 215
Figure 13.19.3.
Blechnum amabile:
habit, showing
dimorphic fertile
frond (center) and
sterile fronds, bar =
5 cm (2 in.).
a b
soil or potting mix. Fronds gradually wither in place upon
light frost. This species is a slow grower.
Blechnum amabile bears dimorphic, one-pinnate fronds.
The sterile blades are lanceolate and narrow, 2–4 cm (0.75–
1.5 in.) wide, and the fertile fronds are barely longer. Ster-
ile pinnae are numerous, linear, and decurrent on the
rachis, with the lower pinnae shorter and triangular. The Figure 13.19.4. Blechnum appendiculatum variants:
fertile pinnae are narrower and more contracted than the a. monomorphic sterile frond of the less red form with
sterile ones. The species is native to China and Japan. glandular hairs; b. monomorphic sterile frond of the
This plant is similar to Blechnum niponicum, which see. redder form with needle-like hairs. Bar = 5 cm (2 in.).
a b
Plants circulating as B. capense in the United States may gins strongly enrolled when old. The plant is native to
not be the New Zealand species. South America.
In the United States and England, plants circulating as
Blechnum cordatum (Desvaux) Hieronymus Blechnum chilense (Kaulfuss) Mettenius, B. tabulare (Thun-
13.19.8 berg) Kuhn, and B. magellanicum (Desvaux) Mettenius all
Semi–hardy to hardy, Zone 7 seem to be B. cordatum. Blechnum wattsii Tindale of Aus-
A medium-large fern with erect fronds that may be spaced tralia is rarely cultivated in the United States. It can be dis-
along a creeping rhizome as well as clustered on an erect tinguished from B. cordatum by the absence of aerophores,
rhizome. Grows under medium light in moist-wet garden more oblong pinnae that are abruptly long acuminate at
soil or potting mix. In the northwestern United States it the tip, and fertile pinnae that are farther apart.
may take full sun if well watered. Rhizomes and stolons
below the ground usually survive freezing weather. The Blechnum falciforme (Liebmann) C. Christensen
plants spread by stolons. 13.19.9
Blechnum cordatum has dimorphic fronds. The sterile Semi-tender
fronds have oblong, dark green blades that are somewhat A large fern with ascending to erect rhizomes and whorled
hard and leathery, with the terminal pinna resembling the fronds. Grows under medium light in moist garden soil or
lateral pinnae; the pinnae are oblanceolate and finely potting mix.
toothed. On the lower surface of the rachis opposite each The fronds of Blechnum falciforme are one-pinnate, di-
pinna stalk is a small, elongated bump, a type of aerophore. morphic, and noticeably scaly. The stipe has a dense mat-
Aerophores may be inconspicuous or absent on dried fronds ting of twisted hair-like scales, and the sterile pinnae have
but are easy to see on fresh fronds. They are usually whitish cartilaginous, serrate margins. The basal pinnae are not
yellow and contrast strongly with the color of the sur- reduced, and the undersurfaces of the costae are scaly. The
rounding tissue. The fertile pinnae are linear, with the mar- species is native to Central and South America.
a b
b
a
Blechnum niponicum has dimorphic fronds, the fertile latum. The species is widespread in tropical America.
fronds taller than the sterile ones. The sterile fronds are one- Most plants identified as Blechnum occidentale in gar-
pinnate and 5–10 cm (2–5 in.) wide. The scales on the base dens in the United States are actually B. appendiculatum.
of the stipe are linear. This species is native to Japan.
Blechnum spicant resembles B. niponicum but differs by Blechnum penna-marina (Poiret) Kuhn
narrower, less oblanceolate sterile fronds and much taller 13.19.14; 18
fertile fronds. The similar B. amabile has a creeping rhi- Hardy, Zone 5
zome, narrower sterile fronds, shorter fertile fronds, and A small fern with fronds spaced along a much-branched,
ovate stipe scales. creeping, stoloniferous rhizome. Grows under medium or
high light in moist but well-drained, acidic garden soil or
Blechnum occidentale Linnaeus 13.19.13 potting mix. The plants spread by stolons and make a use-
Hammock fern ful ground cover in areas with mild winters.
Probably tender Blechnum penna-marina bears one-pinnate, dimorphic
A small to medium fern with erect rhizomes and clustered fronds, the fertile ones much contracted and about twice as
fronds. Grows under medium light in moist garden soil or tall as the sterile. The species is native to South America,
potting mix. The plants spread by stolons. southeastern Australia, New Zealand, and the circum-
Blechnum occidentale is characterized by blades that are antarctic islands.
one-pinnate at the base but pinnatisect above and blade An unnamed dwarf cultivar circulates in the trade, as
apices that are gradually tapered to the tip. The blades are well as the following:
usually narrowly triangular, with the lowest pinnae point- ‘Cristata’. Tip of frond crested.
ing downward and heart-shaped at the base. The rhizomes
usually bear several stolons. The lower surface of the rachis Blechnum polypodioides Raddi 13.19.15
is glabrous, not hairy as in the closely related B. appendicu- syn. Blechnum unilaterale Swartz
Tender
A small to medium fern with erect, stoloniferous rhizomes
and clustered fronds. Grows under medium light in moist
garden soil or potting mix.
Blechnum polypodioides is characterized by pinnatisect
to one-pinnate, lanceolate fronds tapered at both ends. The
sterile and fertile fronds are monomorphic and minutely
hairy on the lower surfaces. The pinnae tend to be numer-
ous, oblong to linear, and essentially entire, and the lower
ones are distant and much reduced. The species is wide-
spread in tropical America.
Figure 13.19.14.
Blechnum penna-
marina: habit,
showing sterile
frond (shorter
outer fronds) and
dimorphic fertile
fronds (taller
a b inner fronds), bar
= 5 cm (2 in.).
a b
Figure 13.19.16.
Blechnum
punctulatum: Figure 13.19.17. Blechnum serrulatum: a. monomorphic
a. dimorphic fertile fertile frond; b. young sterile frond. Bar = 5 cm (2 in.).
frond; b. sterile
a
frond. Bar = 5 cm
(2 in.).
leaves are monomorphic. Sori are borne only on the distal
pinnae and are covered by erose-lacerate indusia. This spe-
cies is native to the United States (Florida), the West Indies,
Blechnum serrulatum L. C. Richard 13.19.17 Central and South America, India, Malaysia, and Australia.
syn. Blechnum indicum of authors, not N. L. Burman In the Old World this species has long been called Blech-
Saw fern, swamp fern num indicum N. L. Burman; however, that name is a syn-
Tender, Zone 9 onym of Asplenium longissimum Blume (Morton 1967).
A large fern with creeping rhizomes and clustered, shiny,
thick fronds. Grows under medium to high light or direct Blechnum spicant (Linnaeus) Smith 13.19.18
sun. This species prefers moist garden soil or potting mix Deer fern
but tolerates wet soils. The fronds tend to grow upright in Hardy to semi-hardy, Zone 7
full sun. During the dry season, the pinnae are shed from A medium-sized fern with erect, sometimes-branched rhi-
the rachis by means of the abscission layer, leaving behind zomes and fronds in a rosette or cluster. Best grown under
erect, naked rachises. The species often forms dense colo- low-medium light in moist, well-drained, acidic potting
nies in the wild. mix. Be sure to provide the plants with humidity and acidic
Blechnum serrulatum is characterized by fronds one-pin- soil, and avoid using hard water. This evergreen species has
nate throughout, with 20–40 pinna pairs and a single api- proven difficult to grow in the southwestern United States,
cal segment. The pinna margins are finely serrate, and care- possibly because of high summer heat and humidity. It
ful inspection will reveal that the pinnae are jointed to the grows readily from spores.
rachis, with the joint visible as a faint dark line or swelling. Blechnum spicant bears fronds that are one-pinnate and
The stipes are often densely clustered. The sterile and fertile dimorphic. The sterile fronds vary from lanceolate to nar-
222 C 13
Figure 13.20.3. Bolbitis portoricensis: sporangia spreading Figure 13.20.4. Bolbitis heteroclita: sterile frond, bar =
on under surface. 5 cm (2 in.).
lar for terrariums. Bolbitis heudelotii in the United States plants. Many of the species reduced to B. heteroclita are rec-
and B. fluviatilis in Europe are sometimes sold in aquar- ognized in horticulture and are here given cultivar status.
ium supply stores as underwater plants. Some species are Difformis Group (Edanyoa difformis Copeland). Figure
easy to grow but are rarely seen in cultivation, such as B. 13.20.5. A diminutive creeping fern often used in terrari-
aliena, which needs warm temperatures year-round. ums and easy to grow. Fronds are usually less than two-pin-
Bolbitis contains 85 species and occurs throughout the nate-pinnatifid and lack the terminal elongated pinna of
tropics of the world. The genus name comes from the the species. Native to the Philippines and long known as
Greek bolbition, diminutive of bolbos, bulb, alluding to the the species Edanyoa difformis. Hennipman believes this
small bulblets borne on the leaves of some species. For a plant is a neotenous form (represents juvenile characteris-
technical monograph of the genus, see Hennipman (1977). tics) of Bolbitis heteroclita. The International Code of No-
Figure 13.20.7.
Bolbitis heteroclita
Simplicifolia
Group: habit
with sterile Figure 13.20.8. Bolbitis heteroclita Tenuissima Group:
fronds, bar = habits, showing variations in sterile fronds, bar = 5 cm
5 cm (2 in.). (2 in.).
F F A C 225
(1977) listed the plant as ‘tenuissima’, a Latin name mean- Bolbitis portoricensis (Sprengel) Hennipman
ing “more slender.” 13.20.1–3, 10
syn. Bolbitis cladorrhizans (Sprengel) Ching
Bolbitis heudelotii (Bory ex Fée) Alston Tender
13.20.9 A medium to large fern with medium-long-creeping rhi-
Tender zomes and elongate blade apices bearing buds at their tips.
A small fern with medium- to long-creeping rhizomes. Can grow submerged in an aquarium or out-of-water in
Grows under medium light on rocks or in soil. This species pots with garden soil or a sand-peat mix in medium light.
can be used as an underwater plant for aquariums, or it can If grown in pots, the soil must be kept wet.
be potted in a peat-sand-soil mix and kept wet. It is rarely The fronds of Bolbitis portoricensis are triangular and
cultivated. one-pinnate, with the lower pinnae longer and lobed on
Bolbitis heudelotii has one-pinnate fronds, usually with the basiscopic side. The veins are netted. The apical por-
ascending pinnae. If the plants are submerged, the sterile tions of well-developed fronds often are elongate (some-
pinna margins are entire, but when held above water, they times whip-like or tailed) and bud-bearing at the tips. The
are irregularly and shallowly serrate. The fertile frond has species is native to tropical America.
narrower pinnae and emerges above the water during non-
flood periods. This species is native to Africa, where it often Bommeria
grows along streambanks prone to flooding. Copper fern
Bommeria is a genus of attractive small ferns that produce
triangular fronds covered with silky white hairs. They are
rarely cultivated and are native to dry, rocky places.
The most distinctive characteristics of this genus are its
small, triangular to pentagonal blades and sori that run
Figure 13.20.9.
Bolbitis heudelotii: Figure 13.20.10. Bolbitis
habit with sterile portoricensis: sterile frond
frond, bar = 5 cm and rhizome, bar = 5 cm
(2 in.). (2 in.).
226 C 13
b
F F A C 227
Camptosorus
syn. Asplenium, in part
Walking fern
The walking ferns are small, lime-loving terrestrials used in
rock gardens or terrariums. The common name derives
from the plant’s ability to form plantlets at the tips of its
long-tapered fronds, making the plant appear to “walk”
over the rock surface. Although it can be grown in warmer
climates outdoors or in terrariums, Camptosorus must have
humidity and drainage. The plants grow best in temperate
climates.
The genus is characterized by netted veins and fronds
that taper to a long, whip-like apex, which roots at the tip. a
b
The rhizome is short and erect. The sori are elongate and
scattered (not regularly arranged as in the closely related
genus Asplenium) and covered by a flap-like indusium. This Figure 13.23.1. Camptosorus rhizophyllus: a. habit, bar =
genus is often classified in Asplenium because it hybridizes 5 cm (2 in.); b. sori and indusia, bar = 5 mm (0.2 in.),
with several species of that genus. courtesy of T. Hoshizaki..
F F A C 229
Campyloneurum
Strap fern
The more frequently cultivated strap ferns, such as Campy-
loneurum angustifolium and C. phyllitidis, are handsomely
displayed in hanging containers, where their dense cluster
of narrow fronds form a spreading or gracefully arching
pattern. The frond width can vary greatly within a species.
Nearly all species of Campyloneurum have simple, en-
tire, strap-shaped leaves. Their blades are glabrous except
along the rachis, where there may be scattered scales. The
veins are always netted, although this may be difficult to
see in species with thick blades. In many species the main a b
lateral veins are parallel and connected by curved cross-
veins to form areoles, and the areoles enclose veinlets that
point toward the margin. The sori are round, lack indusia,
and in most species are borne in one to four rows between Figure 13.24.1. Campyloneurum angustifolium: a. habits
the main veins. The rhizome varies from short- to long- of two plants, bar = 5 cm (2 in.); b. sori and vein
creeping and may be clothed by clathrate or nonclathrate pattern, bar = 1 cm (0.4 in.).
scales.
The genus name comes from the Greek kampylos,
arched, and neuron, nerve, referring to the arched veinlets
230 C 13
caudate apices, and white dots on the upper blade surfaces. to go from spores to mature, spore-bearing plants, though
Veins are less obscure on the lower surfaces. The areoles are three or four months is more typical. Because of this quick
irregularly shaped, with generally five to eight between the cycle, the plants are favorites of laboratory geneticists who
costa and margin, and the sori are in two rows between the study differences in heritable traits from one generation to
main lateral veins. The species occurs from southern Mex- the next.
ico to Costa Rica. The plants live about one year and are usually perpetu-
ated by the numerous buds that form on the blade surface,
Ceratopteris 13.25.1 typically in the margins of the sinuses. The buds detach
Water fern, water sprite and float away from the parent plant. Mature plants can be
Ceratopteris is commonly used as an aquarium plant. These propagated by division of the rhizome, which may be al-
ferns either float on water, grow fully or partially sub- lowed to float or, if grown submerged, held in place by
merged, or root in mud. The fertile leaves are erect and are coarse gravel. If plants are to be rooted in mud, garden soil
produced above water. In Asia the foliage of Ceratopteris with little organic matter will do. Some growers recom-
thalictroides is used in salads and is said to have a peppery mend a mixture of about half peat and half sand mixed with
taste. 10% top soil. Plants that are rooted in mud need their
Ceratopteris is characterized by short, erect, inconspicu- fronds kept moist. Temperatures must be maintained close
ous rhizomes bearing a few thin scales. The sterile and fer- to 27°C (80°F) for good growth. The plants and buds de-
tile fronds are dimorphic—the fertile ones are more finely cline when temperatures are below 20°C (68°F), and if lost,
cut with narrower segments—but both have triangular, new plants must be started from spores.
glabrous blades. The veins are netted, not free. The large, The genus, which consists of four species, is native to
marginal sporangia are protected by a continuous enrolled the tropics and subtropics worldwide. The name comes
indusium. from the Greek keras, horn, and pteris, fern, alluding to the
This genus has the distinction of having the fastest life antler-like fertile leaves. The species are sometimes difficult
cycle of any fern. The plants can take as little as one month to identify because the fronds vary considerably in form.
Only Ceratopteris thalictroides and C. pteridoides are com-
monly cultivated. For more details on cultivation, see Curt-
wright (1995) and Lloyd (1974).
Figure 13.26.3.
Ceterach aureum:
fronds, bar =
5 cm (2 in.).
that may be glabrous, glandular, hairy, scaly, or covered Cheilanthes: Upper surface of the blade with-
with a colored powder on the lower surface. Some species out stellate scales, not striate; lower surface
have numerous bead-like segments that are circular or of the blade not powdery; margin of the fer-
nearly so and often cupped, with the open end of the cup tile segments slightly enrolled to cover the
facing downward. sporangia; ultimate segments entire, stalked
Cheilanthes contains about 180 species and is wide- or sessile. (Cheilanthes alabamensis, C. argen-
spread in the temperate and tropical regions of the world. tea, C. arizonica, C. bonariensis, C. buchtienii,
The genus name comes from the Greek cheilos, lip, and an- C. covillei, C. eatonii, C. farinosa, C. feei, C.
thos, flower, alluding to the position of the sporangia be- fendleri, C. gracillima, C. hirta, C. lanosa, C.
neath the lip-like false indusium. lendigera, C. lindheimeri, C. newberryi, C. tomentosa, C. vil-
Some species of Notholaena and Pellaea are difficult to losa, C. wrightii)
distinguish from Cheilanthes. In view of this problem,
Notholaena has been redefined to include only those species Mildella: Upper surface
with a dense white or yellow waxy powder on the lower sur- of the blade without stel-
faces of the blades and with sporangia at the tips of the veins late scales, not striate;
in a more or less continuous marginal band covered by an lower surface of the blade
enrolled but texturally unmodified frond margin. This def- not powdery; margin of
inition, accepted here, places all the cultivated species for- the fertile segments en-
merly listed as Notholaena into Cheilanthes. Many Cheilan- rolled to cover the spo-
thes species go in and out of cultivation rather frequently, rangia; ultimate segments
and therefore all could not be treated here. The reader is entire, sessile, adnate; in-
referred to floras, particularly of drier regions, for more dusium linear, attached
information. below the margin of the segment and thus leaving a nar-
Pteridologists now distinguish several genera formerly row band of blade tissue between the edge and the indu-
included in Cheilanthes. To help with identification, these sium. (Mildella intramarginalis)
genera are described here.
Notholaena: Upper surface
Argyrochosma: Upper surface of the blade of the blade without stel-
without stellate scales, not striate; lower sur- late scales, not striate;
face of the blade usually has a yellowish or lower surface of the blade
whitish powder; margin of the fertile seg- often with a yellowish or
ments enrolled to cover the sporangia or not; whitish powder; margin of
ultimate segments entire and stalked. (Argy- the fertile segments en-
rochosma dealbata, A. jonesii, A. microphylla) rolled to cover the sporan-
gia; ultimate segments en-
Aspidotis: Upper tire, sessile to subsessile.
surface of the (Notholaena standleyi)
blade without stel-
late scales, striate;
lower surface of the blade not Cheilanthes alabamensis (Buckley) Kunze
powdery; margin of the fertile 13.27.4
segments en- Alabama lip fern
rolled to cover Hardy, Zone 7
the sporangia, A small to medium fern with short-creeping to compact
white-papery; ul- rhizomes and clustered fronds. Grows best under medium-
timate segments entire and short-stalked. high light in moist-dry to dry, basic garden soil and sand.
(Aspidotis californica, A. carlotta-halliae, A. Cheilanthes alabamensis has black stipes, rachises, and
densa) costae, with the color entering the pinnules. The hairs on
the upper rachis surface are dense, short, and twisted; those
Astrolepis: Upper surface of the blade with beneath are long and sparse or absent. The blade is lanceo-
stellate scales, not striate; lower surface of late to linear-oblong, two-pinnate to two-pinnate-pinnati-
the blade not powdery; margin of the fertile fid, with segments elliptic to elongate-triangular, not bead-
segments flat, not enrolled to cover the spo- like. The blade surface varies from hairy to nearly glabrous.
rangia; pinnae stalked to subsessile. (As- The species is native to the United States and Mexico,
trolepis beitelii, A. sinuata) where it typically grows on limestone.
F F A C 237
Figure 13.27.5.
b Cheilanthes argentea:
a. frond, bar = 2.5 cm
(1 in.); b. lower surface of
fertile pinna, bar = 2 cm
a (0.8 in.).
b
Figure 13.27.12.
Cheilanthes feei: a. frond,
bar = 2.5 cm (1 in.);
b. upper surface of
Figure 13.27.13.
pinnule, bar = 5 mm
Cheilanthes fendleri: a.
(0.2 in.), after Knobloch
habit, bar = 2.5 cm (1 in.);
and Correll (1962);
b. lower surface of fertile
c. lower surface of pinnule,
pinnule, bar = 2 mm, after
bar = 5 mm (0.2 in.), after
Knobloch and Correll
a Knobloch and Correll
(1962).
(1962).
c
b
Figure 13.27.18.
Cheilanthes
lindheimeri: a.
habit, bar =
2.5 cm (1 in.);
b. upper surface
of pinnule, bar =
2 mm, after
Knobloch and
Correll (1962);
c. lower surface
Figure 13.27.17. of pinnule and
Cheilanthes lendigera: a enlargement of
a. frond, bar = 2.5 cm scale, bar =
(1 in.); b. lower surface 2 mm, after
of fertile pinnule, Knobloch and
a Correll (1962).
showing pouch-like
indusia, bar = 2.5 mm c
(0.1 in.).
b
Figure 13.27.22.
Cheilanthes wrightii:
a. habit, bar =
2.5 cm (1 in.);
b. lower surface of
fertile pinnule, bar =
2.5 mm (0.1 in.).
After Knobloch and
Correll (1962).
a b
a b
a c
has gracefully arching fronds, but older plants are often too
big for most homes. Plantlets sometimes emerge as off-
shoots at the base of the crown. These may be separated
and rooted, but are usually hard to establish.
The stipe of Cibotium schiedei is clothed with yellow-
brown silky hairs, and the blades are large, 2–3 m (6.5–10
ft.) long, and light green on the upper surface but glaucous
on the lower. The segments lack ears, and the sori often pro-
trude beyond the margin. The species is native to Mexico.
A similar species, Lophosoria quadripinnata (J. Gmelin)
C. Christensen, is becoming more frequently cultivated. It
might be identified as a Cibotium because it also has woolly
hairs on the trunk and stipe base and large, finely divided
leaves that are glaucous beneath. It can be distinguished
from Cibotium schiedei by having sori borne on the lower
surface of the leaf, rather than at the margin, and by the
lack of indusia.
terminates in an apical segment that resembles the lateral Coniogramme japonica (Thunberg) Diels
pinnae, and the veins tips are conspicuously enlarged on 13.29.1, 2, 4
the upper surface of the leaf. The sori run along the veins Bamboo fern
between the midrib and margin of the segments. Indusia Semi-hardy, Zone (7)8
are absent. The cultivated species of Coniogramme mimic a A medium-large fern with deciduous fronds and medium-
large Pteris cretica but differ by having longer-creeping rhi- to long-creeping rhizomes. Grows well under low light in
zomes, deciduous fronds, and sori that run between the moist garden soil or potting mix.
midrib and margin of the segments, not along the margin Coniogramme japonica resembles the preceding one but
as in Pteris. has netted veins. It is native to China, Taiwan, Korea, and
The genus consists of about 20 species native to the Old Japan.
World tropics, Japan, and in the Pacific to Hawaii. Its name ‘Variegata’. Segments with yellow along the midribs.
comes from the Greek konios, dusty, and gramme, line, re-
ferring to the lines of sori along the veins.
Some botanists consider Cryptogramma acrostichoides to Cryptogramma crispa resembles C. acrostichoides but has
be a variety of the European species, C. crispa, but they dif- thinner, deciduous leaves, and sterile blades up to four-pin-
fer consistently in blade cutting and chromosome number nate. It is native to Europe.
(2n = 60 in C. acrostichoides and 2n = 120 in C. crispa).
They also differ in habitat. Cryptogramma acrostichoides Cryptogramma sitchensis (Ruprecht) T. Moore
grows on noncalcareous rocks, often in slightly drier habi- 13.30.6
tats than is usual for other species in the genus. In contrast, Alaska parsley fern
C. crispa grows on calcareous rocks in more humid places. Very hardy, Zone 2
A small fern with suberect to erect rhizomes and tufted
Cryptogramma crispa (Linnaeus) R. Brown evergreen fronds. Grows well under medium light in gar-
13.30.5; 23 den soil with good drainage. Probably does best if the soil is
European parsley fern moist in spring and drier at other times.
Very hardy, Zone 2 Cryptogramma sitchensis greatly resembles C. acrostichoi-
A small fern with erect to suberect rhizomes. Grows best des and is believed to have arisen by hybridization between
under medium light in well-drained garden soil. The plants that species and possibly C. raddeana Formin of eastern
seem to do best if the soil is kept moist in the spring but Asia. It differs from C. acrostichoides by its obovate seg-
moist-dry other times of the year. ments with two to six deep lobes and at least some three- to
four-pinnate sterile fronds. It is native to northwestern
Canada and Alaska, where it typically grows among rocks
in acid soils.
Cryptogramma stelleri (S. G. Gmelin) Prantl Ctenitis contains 150 species and is native to tropical re-
13.30.7 gions of the world. The genus name comes from the Greek
Slender rock brake, cliff brake, fragile rock brake kteis, comb, referring to the comb-like lobes of the pinnae
Very hardy, Zone 2 in Ctenitis submarginalis (Langsdorff & Fischer) Ching, the
A small fern with deciduous fronds distributed along a slen- type species of the genus.
der (1–1.5 mm wide), creeping rhizome with few branches.
Thrives under medium light in well-drained, moist, basic Ctenitis sloanei (Poeppig ex Sprengel) C. V. Morton
garden soil. The fronds die back by late summer and need 13.31.1–4
less water at that time. syn. Ctenitis ampla auct. non (Humboldt & Bonpland ex
Cryptogramma stelleri bears stipes that are dark at the Willdenow) Ching
base but greenish above. The blades are up to two-pinnate, American tree fern, Florida tree fern
herbaceous to membranous, and thin. The segments of Tender, Zone 10(11)
sterile leaves are ovate-lanceolate to fan-shaped and lobed A large tree fern with erect stems that can form trunks up to
halfway to the midrib. The species is native to northern 1.3 m (4 ft.) tall, although it infrequently does so in culti-
North America, Europe, and Asia, where it typically grows vation. Grows under medium light in moist, drained pot-
on calcareous rocks, often forming dense colonies in crev- ting mix. This species is sensitive to transplanting and is
ices or ledges. difficult to establish.
The stipe scales of Ctenitis sloanei are dense, linear or
thread-like, orange (rarely brown), and toothed. The veins
reach all the way to the margin, and the indusium falls away
early. The species is native to the United States (Florida)
and American tropics.
Ctenitis
These medium- to large-sized, terrestrial ferns are infre-
quently cultivated, except in Florida. The Florida tree fern
(Ctenitis sloanei ) can form a trunk up to 1.3 m (4 ft.) tall.
Ctenitis can usually be distinguished by the following
combination of characteristics: scaly rhizomes; three or
more vascular bundles in the stipe; minute, jointed, often
reddish hairs on the upper surfaces of the rachises and
costae; free veins (not netted); and round sori. Usually the
rachis is grooved on its upper surface and the costae are not.
In those rare instances where both are grooved, the grooves
are interrupted at the junctures, not continuous. This is in
contrast to dryopteroid ferns such as Cyrtomium, Dryop- Figure 13.31.2. Ctenitis: rachis, groove not opening into
teris, and Polystichum, in which the grooves are prominent the costae, costae not grooved (left), costae grooved
on both the rachises and costae and run into one another. (right).
252 C 13
Cyathea
syn. Hemitelia
Tree fern
Cyathea is a genus of elegant tree ferns that typically bear a
crown of large fronds on top of a tall, erect trunk. They can
reach 15–20 m (50–65 ft.) tall in the wild, and some
mountain slopes support dense stands of these stately trees.
In temperate areas this genus is seen mainly in conservato-
ries or large greenhouses. Tree ferns can be grown as house-
plants, but most species seem to do best outdoors in sub-
tropical or tropical gardens, where they can grow up to 5 m
(16 ft.) in height. Most thrive under filtered light, but they
will grow in full sun in coastal areas where heat and light are
not too intense. Growth is best in rich, organic, moist soil.
(See Chapter 10 for more information.)
Tree ferns have several uses. In some cultures they are
used for food, fodder, or building material. The trunks are
highly valued in backwoods construction because they are
strong and resist decay. The thick layer of tough aerial roots
(the root mantle) found around the trunk of some species
is used as a growing medium for epiphytes, especially or-
chids. The scales on the trunk and stipes can shed readily,
and care should be taken to avoid getting the scales into
your eyes and on your skin. Although not toxic, the scales
can cause severe irritation.
Figure 13.31.3. Ctenitis sloanei: sori and indusia. Most species of Cyathea have upright, scaly trunks and
a layer of tough aerial roots, at least toward the base. The
fronds are borne in a whorl at the top of the stem, forming
a palm-like growth habit, and they usually have finely di-
vided, two- to three-pinnate blades. The stipe scales lack
a dark bristles at the apex, in contrast to those of the similar
genus Alsophila, and the marginal cells differ in orientation
and often in color from the central cells (when viewed
under 20× magnification). The round sori are borne on the
undersides of the blades, and the indusia, when present,
vary from scale-like to cup-like or globose.
To distinguish cyatheaceous ferns from other tree-fern
b groups, look at the indument or covering on the stem apex
or stipe bases. Cyathea has a covering of scales, whereas
dicksoniaceous tree ferns such as Cibotium and Dicksonia
have hairs. Sadleria has scales on the trunk and stipe, but its
fronds are narrow and the sori are covered by a conspicuous
oblong indusium.
Cyathea is native to the Old and New World tropics,
occurring mainly in cloud forests. It consists of about 115
species. The genus name comes from the Greek kyathos, a
wine cup, referring to the cup-like shape of the indusium in
certain species.
c Cyathea is defined here in the narrow sense, excluding
Alsophila, Cnemidaria, and Sphaeropteris. Although pteri-
Figure 13.31.4. Ctenitis sloanei: a. frond from mature dologists agree that these genera represent natural groups,
plant, bar = 5 cm (2 in.); b. frond from immature plant, they disagree over whether the groups should be recognized
bar = 5 cm (2 in.); c. veins reaching the margin, bar = as separate genera. Some pteridologists classify the species
5 mm (0.2 in.). of all four groups into one large, inclusive genus, Cyathea.
F F A C 253
For a reference to tree-fern taxonomy, see Kramer and Cyathea arborea (Linnaeus) Smith 13.32.1, 2
Green (1990). The following guide to the cultivated genera West Indian tree fern
of Cyatheaceae is provided to facilitate identification: Tender to very tender
A large tree fern. Grows best under medium light in moist
potting mix. This species grows in southern Florida and
seems to require warm nights.
The fronds of Cyathea arborea eventually fall away from
the trunk. The stipe scales are mostly whitish, and the base
of the segment midrib on the underside have one or two
white, puffy scales. The saucer-shaped indusium forms a
cup beneath the sporangia. The species is native to the West
Indies, Venezuela, and Colombia.
Cyclopeltis
These medium-sized terrestrial ferns are grown in south-
ern Florida and are sometimes used in public parks in the
West Indies. The rhizome is thick, densely scaly, and as-
cending or erect. The blades are one-pinnate with many
Alsophila: Stipe scales tipped with a dark bristle; marginal sessile pinnae and a terminal one resembling the laterals.
cells of scales are different from the central cells in orienta-
tion, size, and often color; stipes spiny, with spines black
and shiny. (Alsophila australis, A. tricolor)
b
a
b
c
Figure 13.33.1.
Figure 13.32.2. Cyathea arborea: a. pinnule, bar = 1 cm Cyclopeltis
(0.4 in.); b. stipe scale, bar = 1 cm (0.4 in.); c. detail of semicordata: a
segment with bullate scale at base, bar = 1 mm, a. habit, bar =
illustration by P. J. Edwards, used with permission; 10 cm (4 in.);
d. enlargement of bullate scale, bar = 0.1 mm, b. sori with peltate
illustration by P. J. Edwards, used with permission. indusia, bar = 5
mm (0.2 in.).
The veins are free, and the sori are round and covered by
peltate indusia. Cyrtomium 13.34.1, 2
The peltate indusium immediately separates Cyclopeltis syn. Phanerophlebia
from Dryopteris and Thelypteris, which it superficially re- This genus of terrestrial or rock-inhabiting ferns is well rep-
sembles. It differs from Polystichum by having entire, not resented by the house holly fern (Cyrtomium falcatum),
spiny, margins and pinnae that lack an inner, upward- widely grown indoors in pots for its leathery, glossy green,
pointing ear. durable foliage. The most common cultivar in nurseries is
The genus, which consists of five species, is native to the Rockford fern (C. falcatum ‘Rockfordianum’), which
tropical America and southeastern Asia. The genus name has irregularly fringed pinna margins. This plant is easy to
comes from the Greek kyklos, circle, and pelte, shield. grow, but its soil should be kept on the moist-dry side. It
has become naturalized in parts of the United States with
Cyclopeltis semicordata (Swartz) J. Smith mild winters. In the wild Cyrtomium grows in the soil or on
13.33.1 rocks, not as epiphytes.
Tender Cyrtomium can usually be distinguished by erect, scaly
A medium-sized fern with erect rhizomes. Best grown rhizomes; one-pinnate, evergreen fronds; netted veins; and
under medium light in moist garden soil. The plants are round sori covered by an umbrella-shaped indusium. The
occasionally grown in Florida. lateral pinnae tend to be nearly equal in size, and the ter-
Cyclopeltis semicordata has one-pinnate fronds with ses- minal pinna usually resembles the lateral ones, except that
sile pinnae that have a semicircular basal lobe overlapping the base can be lobed.
the rachis. The upper surfaces of the rachises and costae Some pteridologists classify the New World species of
bear short hairs, as is typical for ferns related to Ctenitis and Cyrtomium (which are rarely cultivated) in Phanerophlebia
Tectaria. It is widespread in the American tropics where it because those species have casually to sparingly netted veins
typically grows on wet forest floors. like Phanerophlebia and in contrast to the Old World spe-
cies of Cyrtomium. At the other extreme, Cyrtomium and
F F A C 255
b
b
c
d
var. clivicola (Makino) Tagawa. Upper surfaces of the tured blades about 30 cm (12 in.) long and 9 cm (3.5 in.)
blades matte yellow-green; pinnae typically in about 10 wide. The pinnae are distant from one another, nearly op-
(15) pairs, usually to 4 cm (1.5 in.) long, mostly eared at the posite or alternate, and crenate-serrate. The apex is unusual
base; indusium margins somewhat erose. Native to China in being short and deeply pinnatifid. The indusium mar-
and Japan. gins are erose. The species is native to China.
var. fortunei. Upper surfaces of the blades deep green,
shiny; pinnae in 15–25 pairs, less than 2–3 cm (0.75–1.2 Cyrtomium macrophyllum (Makino) Tagawa
in.) wide, generally truncate and nearly earless; indusium 13.34.8, 9
margins entire. The indusia are grayish white throughout Big-leaf holly fern
(concolorous). Native to Japan, South Korea, southeastern Semi-hardy, Zone 6(7)
China, Indochina, and Thailand. A medium-sized fern with erect rhizomes. Grows under
var. intermedium Tagawa. Pinnae about 15 (8–19) pairs, medium light in moist-dry garden soil or potting mix. The
bases widely wedge-shaped, weakly eared or not, gradually evergreen fronds are a lighter green than those of Cyrto-
narrowed toward an attenuate apex. The indusia are black- mium falcatum.
ish in the center. Native to Japan. Some of the trade mate- Cyrtomium macrophyllum has 2–8 (12) pinna pairs, with
rial in the United States named as this variety probably rep- the lateral pinnae more than 4 cm (1.5 in.) wide and entire
resents var. clivicola. or slightly toothed toward the tip. The pinna bases are
rounded to broadly wedge-shaped, without ears, and the
Cyrtomium lonchitoides (H. Christ) H. Christ terminal pinna is larger than or nearly the same size as the
13.34.7 lateral pinnae. The indusia are nearly entire. The species is
Semi-hardy (?) native to Japan, China, and the Himalayas.
A medium-sized fern with erect rhizomes. Best grown var. tukusicola (Tagawa) Tagawa (Cyrtomium tukusicola
under medium light in moist garden soil or potting mix. Tagawa). Figure 13.34.9. Separated from var. macrophyl-
Cyrtomium lonchitoides is characterized by stipes only lum by wedge-shaped pinnae and indusia that are black-
5–10 cm (2–4 in.) long and elongate, lanceolate, thin-tex- brown in the center but pale at the margins. Native to
Japan (Honshu and Kyushu). Evergreen and hardy in Salt
Lake City, Utah.
Figure 13.34.9.
Cyrtomium
macrophyllum var.
tukusicola: frond,
bar = 5 cm (2 in.).
Figure 13.35.1. Cystopteris fragilis: habit.
The genus name comes from the Greek kystos, bladder, and
pteris, fern, and refers to the inflated indusium covering the
sorus.
Figure 13.35.6.
Cystopteris fragilis:
a. habit, bar = 5 cm
(2 in.); b. sorus and
indusia, bar = 1 mm,
after Knobloch and
Correll (1962).
Figure 13.35.5.
Cystopteris dickieana:
habit, bar = 5 cm (2 in.).
a
Figure 13.35.9.
Cystopteris reevesiana:
frond, bar = 5 cm
(2 in.).
Figure 13.35.10.
Cystopteris tennesseensis:
habit, bar = 5 cm (2 in.).
rachis and (sometimes) costae, but the bulblets are smaller, Davallia 13.36.1–3
scalier, and less fleshy. Minute gland-tipped hairs occur on syn. Araiostegia, Humata, Pachypleuria, Scyphularia,
the rachises and costae (magnification needed), a trait im- Trogostolon
parted from C. bulbifera. The veins end in notches or teeth. Davallias are among the most beautiful and treasured cul-
A characteristic of the Tennessee bladder fern not shared by tivated ferns. Their broad, lacy, often shiny fronds are usu-
other bladder ferns is that nearly all the leaves, even the ally displayed in hanging baskets in conservatories or green-
smallest, bear sori. This species is native to the central houses, where the entire basket may be hidden by the
and eastern United States, where it grows on limestone. It foliage. The most sought-after davallias are those with
is the common bladder fern in the Ozarks of Missouri and finely divided fronds, such as certain selections of the Fiji
Arkansas. davallia (Davallia fejeensis) that bear numerous segments
about the width of a pencil lead. The warm-temperate spe-
Cystopteris tenuis (Michaux) Desvaux cies are grown outdoors in mild climates of the United
13.35.11 States, although most of the nursery stock is used indoors.
syn. Cystopteris fragilis var. mackayi G. Lawson Besides hanging baskets, davallias are planted over rocks
Mackay’s brittle fern or as ground covers so that their creeping rhizomes can be
Very hardy, Zone 4 seen—the rhizomes are often described as “furry feet.” One
A small fern with short-creeping rhizomes. Grows best species with such rhizomes is the bear’s-foot fern (Davallia
under medium light in moist soil or potting mix. The leaves tyermannii). It is a choice fern of small to medium size with
are deciduous and absent during the winter months. durable, triangular, moderately fine-divided fronds. Al-
Cystopteris tenuis resembles C. fragilis and can be diffi- though slower growing than many other common species,
cult to distinguish. It typically has pinnules that are wedge- D. tyermannii less frequently needs repotting. Resembling
shaped at the base, whereas those of C. fragilis are broadest the bear’s-foot fern, the squirrel’s-foot fern, D. mariesii var.
at the base and triangular. The pinnae and pinnules also stenolepis (D. trichomanoides of trade) is the most common
tend to be more ascending than those of C. fragilis, although species in the trade. It has light brown, furry rhizomes and
F F A C 263
ing and scaly, the scales being mostly ovate to lanceolate Araiostegia was delineated primarily by its thin blades,
and peltate; if basally attached, the scales are reniform or finely divided fronds, scaly rachises and costae, and deli-
cordate with overlapping bases. The stipes are jointed to cate, roundish or kidney-shaped indusia attached only at
the rhizomes and fall off cleanly, leaving a circular scar. The the base. The species in this group are rarely cultivated but
blades are anadromous and vary from simple to finely dis- include Davallia hymenophylloides and D. pulchra.
sected (up to four-pinnate) and from lanceolate to triangu- In his monograph on the genus Davallia, Nooteboom
lar in shape. The rachis is raised and grooved on the upper (1994) changed the names of several well-known cultivated
surface, with the grooves running into those of the costae. species. These changes leave many plants with distinct and
The veins are free, not netted. Some species have false veins consistent characteristics (at least in horticulture) essentially
that run parallel to the main veins but are not connected to without names. Because this will bring confusion to gar-
them. Unlike true veins, the false veins do not conduct deners, many of the species synonymized by Nooteboom
water or mineral nutrients; presumably they lend mechani- are still recognized in this book. Eventually, most of these
cal support to the blade tissue. The sori are close to the mar- reduced species should be given cultivar status under the
gin and covered by an indusium that may be round, scale- cultivar-group category discussed in Chapter 12.
like, shell-shaped, pouch-shaped, or more often cup- or This genus is named for Edmond Davall (1763–1798),
tubular-shaped. a Swiss plant collector. It contains about 35 species native
The two recent studies on the delineation of davalliod to the Old World tropics, mainly in Asia and the eastern
genera differ considerably. Kato (1985) recognized several Pacific.
genera whereas Nooteboom (1994) united most of them.
In this work we have followed Nooteboom, but for those Davallia canariensis (Linnaeus) Smith 13.36.4
wishing to use the genera recognized by Kato, we list their syn. Davallia portugal Horticulture
differences below. Canary Island davallia, hare’s-foot fern
Davallia, in the traditional sense, has indusia that are Semi-hardy to semi-tender
attached all the way up the sides, forming a pocket with a
A medium-sized fern with medium- to long-creeping rhi-
single opening toward the leaf margin. The cultivated spe-
zomes. Grows well under medium light or full sun in
cies that belong here are Davallia canariensis, D. denticu-
drained, moist-dry garden soil, potting mix, or uncut moss.
lata, D. divaricata, D. embolstegia, D. epiphylla, D. fejeensis,
The plants are easily cultivated but are moderately slow
D. griffithiana (though with the indusia attached only
growing. The new growth is seasonal.
slightly up the sides), D. lorrainii, D. mariesii, D. pyxidata,
Davallia canariensis is characterized by thick rhizomes
D. solida, and D. tasmanii.
up to 18 mm (0.75 in.) wide that are white-waxy beneath
Humata was separated from Davallia by its leathery
the scales. The blades are broadly triangular, about 40 cm
fronds and indusia attached only at the base, not along the
(16 in.) long and wide, and have a fleshy lip present at the
sides. The cultivated species that belong here include Da-
junction of the costa and rachis on the upper ridge. The
vallia falcinella, D. heterophylla, D. parvula, D. repens, and
indusia are cup-shaped. This species is native to Portugal,
D. tyermannii.
Spain, the Canary Islands, and Morocco.
Kato (1985) recognized several segregate genera from
Humata, such as Trogostolon and Pachypleuria. Under this
system, the genus Trogostolon includes Davallia falcinella. Davallia denticulata (N. L. Burman) Mettenius ex
The characters of Trogostolon are dark brown, peltate, Kuhn 13.36.5
toothed rhizome scales with needle-like apices; fronds syn. Davallia elegans Hedwig
finely dissected with one vein in the ultimate segments; sori Toothed davallia
at a vein fork; and round indusia attached by the base and Tender
halfway up the sides. If Kato’s findings are followed, Pachy- A medium-large fern with medium- to long-creeping rhi-
pleuria includes D. parvula and D. repens. The characters zomes. Grows well under medium light in moist-dry pot-
that define Pachypleuria are peltate, ciliate rhizome scales ting mix or uncut moss with good drainage. The plants are
with wart-like outgrowths on the lower surface; fronds easy to grow and tend to volunteer in suitable climates.
mostly small, leathery, linear to deltate, simple to bipin- The rhizome scales of Davallia denticulata are toothed
natifid, rarely finely dissected; sori at a vein fork or bend; and narrowed abruptly from a broad base. The blade seg-
and firm indusia attached at the base or slightly up the ments are oblong, often with a faint line (false vein) paral-
sides. lel to and between the true veins. The indusia are small and
Scyphularia was segregated from Davallia mainly by the cup-shaped. This species is native to the Old World tropics.
needle-like and bristly scale tips of its rhizome. Cultivated A crested form with somewhat narrower fronds is pres-
species include Davallia pentaphylla and D. triphylla, al- ently cultivated.
though the latter’s rhizome scales are not strongly needle-
like and bristly.
F F A C 265
c c
d
d
a
b
b a
Davallia divaricata Blume 13.36.6 scales of Davallia embolstegia are mostly narrowly triangular
Tender to lanceolate and long-acuminate. Indusia are cup-shaped
A medium-large fern with medium- to long-creeping rhi- with an elongated free lip. This species resembles D. epi-
zomes. Grows well under low to high light in moist-dry phylla but is larger and has ungrooved or only weakly
potting mix or uncut moss with good drainage. The young grooved rachises and nonpeltate rhizome scales. Davallia
fronds are red. embolstegia is native to Borneo and the Philippines.
Davallia divaricata has rhizome scales that are thin,
large, and uniformly reddish brown. The rachises and cos- Davallia epiphylla Sprengel 13.36.8
tae are reddish; the indusia, cup-shaped. This species is na- syn. Davallia denticulata var. elata (G. Forster)
tive to tropical Asia and Malaysia. Mettenius ex Kuhn
Tender
Davallia embolstegia Copeland 13.36.7 A medium-sized fern with medium- to long-creeping rhi-
Tender zomes. Grows well under medium light in drained, moist-
A medium-large fern with medium- to long-creeping rhi- dry potting mix or uncut moss.
zomes. Grows well under medium light in well-drained, Davallia epiphylla has peltate rhizome scales, and the
moist-dry potting mix or uncut moss. This species needs larger stipe scales are broadly heart-shaped at the base and
heat and humidity to grow well. It is rarely cultivated. abruptly narrowed above to a thread-like tip. The indusia
The rhizome scales attach at a notch in the cordate base are cup-shaped with an elongated free lip. This species is
(not peltate), and the segments lack false veins. The stipe native to Malaysia, New Guinea, and the Pacific Islands.
266 C 13
b c
c
b
Figure 13.36.6. Davallia divaricata: a. frond, bar = 5 cm
(2 in.); b. rhizome scales, bar = 5 mm (0.2 in.); c. fertile a
segments, bar = 5 mm (0.2 in.).
Figure 13.36.8. Davallia epiphylla: a. frond, bar = 5 cm
(2 in.); b. rhizome scales, bar = 5 mm (0.2 in.); c. fertile
e segments, bar = 5 mm (0.2 in.); d. indusia, bar = 1 mm;
c e. stipe scales, bar = 2 mm.
c d
a
d
c
b
a
Figure 13.36.11. Davallia fejeensis variations: all lower pinnae (except h, a middle pinna), all unnamed cultivars, except d.
‘Dwarf Ripple’, e. ‘False Plumosa’, i. ‘Plumosa’, and j. ‘Major’. Bar = 5 cm (2 in.).
Figure 13.36.12. Davallia griffithiana: a. habit, with a variant frond on left, bar = 5 cm (2 in.); b. rhizome scales, bar =
5 mm (0.2 in.); c. fertile segments, bar = 5 mm (0.2 in.); d. indusium, bar = 1 mm.
270 C 13
e d
c
b
a
a
Davallia mariesii has triangular fronds up to three-pin- Figure 13.36.17. Davallia mariesii var. stenolepis: a.
nate at the base, with stipe scales triangular to ovate or ob- habit, bar = 5 cm (2 in.); b. rhizome scales, bar = 5 mm
long, mostly narrowing abruptly to a long, twisted apex. (0.2 in.); c. rhizome scale cilia, bar = 2 mm; d. fertile
The whitish tan to brown rhizome scales are lightly pressed segments, bar = 5 mm (0.2 in.); e. stipe scales, bar =
to the rhizome, and the margins bear short cilia and teeth. 2 mm.
The indusia are short and tubular. This species is native to
Korea, Japan, Taiwan, and China.
Nooteboom (1994) considered Davallia mariesii to be
the same as D. trichomanoides var. trichomanoides. The foot fern. Figures 13.36.1–3, 17. Fronds to about 20 cm (8
plants called D. mariesii in horticulture, however, are dis- in.) long, nearly evergreen or with a short dormant period
tinct and important enough in the trade to require main- between shedding of the old fronds and emergence of the
taining the name. new ones. The pinnae and pinnules do not noticeably over-
var. mariesii. Japanese ball fern. Figure 13.36.16. Fronds lap. Variety stenolepis resembles var. mariesii but is only
to 15 cm (6 in.) long, triangular, deciduous, the pinnae slightly deciduous, larger, and has less crowded, more at-
closely placed, and the pinnules sometimes overlapping. tenuate pinnae. Native to the Ryuku Islands, South Korea,
This variety intergrades with var. stenolepis. Native to Korea China, and Taiwan. The variety is easily cultivated but less
and Japan. In Japan the live rhizomes are gathered when cold hardy than var. mariesii. It is the most widely grown
the plant is in its leafless period and tied into balls or other Davallia in the United States and is usually sold, incor-
figures and subsequently hung out to grow as a basket fern. rectly, as D. trichomanoides. Most Japanese botanists con-
var. stenolepis (Hayata) Hoshizaki (Davallia stenolepis sider var. stenolepis merely a semi-evergreen, southern vari-
Hayata, D. trichomanoides of trade, not Blume). Squirrel’s- ant of D. mariesii.
272 C 13
b b
d
c
e
b
a
b
a
The rhizomes of Davallia pyxidata tend to grow stiffly Davallia repens (Linnaeus f.) Kuhn 13.36.22
upward and are covered by easily detached, nearly black syn. Humata repens (Linnaeus f.) Diels, H. vestita
scales with tapered apices and long-hairy margins. The (Blume) T. Moore, Pachypleuria repens (Linnaeus f.)
stipes contain four (sometimes three or five) vascular bun- M. Kato
dles. The secondary pinnules are mostly pointed, with the Semi-tender to tender
apex ending in a dominant tooth. The indusia are short A small fern with medium-creeping rhizomes. Grows best
and tubular. This species is native to Australia. under warm, humid conditions, preferring medium light
Nooteboom (1994) classified Davallia pyxidata as a va- and moist-dry, drained potting mix or uncut moss.
riety of D. solida; however, D. solida var. solida differs most The rhizomes of Davallia repens are white-waxy beneath
conspicuously by having rhizomes that generally do not the scales. The blades are triangular and deeply pinnately
grow stiffly upward. Other differences are the lighter-col- lobed to two-pinnate at the base, with the lower lobes
ored rhizome scales, which are difficult to detach whole; broader and lobed on the lower margin. The indusium is
stipes with six (three to nine) vascular bundles; more leath- semi-circular and attached only at the base. This species is
ery and glossy blades; more obtuse secondary pinnules with native from northern India to Japan, and the Mascarene
apices not ending in a dominant tooth; more obscure veins; Islands to Australia.
and longer indusia. Furthermore, D. pyxidata is hardier Nooteboom (1994) pointed out the tremendous varia-
than the tender D. solida. Given these differences, D. pyxi- tion in the group and reduced Davallia vestita and other
data merits recognition as a distinct species. species to D. repens.
274 C 13
c d
e
b
a
a c
a
d
b
a
a
Figure 13.38.2. Dennstaedtia davallioides:
segments bearing sori.
278 C 13
Dennstaedtia cicutaria can be distinguished from the Dennstaedtia punctilobula (Michaux) T. Moore
other cultivated Dennstaedtia by the numerous needle- 13.38.7
shaped hairs on the lower surfaces of the blades. The rhi- Hay-scented fern
zomes are 12 mm (0.5 in.) thick or more, and branch-buds Very hardy, Zone 3
arise near the stipe bases. The triangular blades have dull A medium-sized fern with long-creeping rhizomes and de-
green surfaces and are densely hairy within the rachis ciduous fronds. Prefers low to high light in moist, acidic
grooves. The species is native to the American tropics. garden soil. This species spreads well in gardens, sometimes
becoming a weed, but it is easy to control by pulling up the
Dennstaedtia davallioides (R. Brown) T. Moore shallow rhizomes. New fronds are produced throughout
13.38.1, 2, 6 the growing season. When crushed, the fronds emit an odor
Lacy ground fern of freshly cut hay. The substance that produces the odor is
Semi-hardy to semi-tender contained within the enlarged, glandular cell of the hair
A medium-large fern with long-creeping rhizomes. Grows covering the blades.
well under medium light in moist or moist-dry garden soil. The blades of Dennstaedtia punctilobula are narrowly
The rhizomes of Dennstaedtia davallioides grow to about triangular to lanceolate and two-pinnate-pinnatifid, with
6 mm (0.2 in.) thick, and the blades are triangular, me- gland-tipped hairs on the lower surface. Epipetiolar buds
dium green, with hairs more or less evenly distributed over frequently occur at the stipe bases. The species is native to
the rachis. The species is native to Australia. eastern North America.
Figure 13.38.7.
Dennstaedtia punctilobula:
frond, bar = 5 cm (2 in.).
Figure 13.40.3.
Dicksonia antarctica:
a. pinna, bar = 5 cm
(2 in.); b. primary
pinnule, bar = 1 cm
(0.4 in.).
Figure 13.40.4.
Dicksonia fibrosa:
a. pinna, bar = 5 cm
(2 in.); b. primary
pinnule, bar = 1 cm b
(0.4 in.).
Dicksonia sellowiana Hooker 13.40.5 Figure 13.40.5. Dicksonia sellowiana: a. pinna, bar =
syn. Dicksonia gigantea H. Karsten 5 cm (2 in.); b. primary pinnule, bar = 1 cm (0.4 in.);
Semi-tender to slightly hardier c. sori and indusia, bar = 1 mm.
A large tree fern, with trunks to 6 m (20 ft.) tall in its native
habitat. Grows best under medium light in moist garden
soil or potting mix. This species is particularly attractive for
its dark, glossy fronds, which are generally broader and and sculpted into animals such as ducks, swans, squirrels,
glossier than those of Dicksonia antarctica. In the coastal and goats. These sculptures are used to adorn lawns and
areas of central and southern California this species can be patios, much like cement statuary is used in the United
grown year-round outdoors without difficulty. States. The Mexicans call this type of sculpture maquique.
The trunk of Dicksonia sellowiana tends to be thickest at Segments of the trunk are also carved into pots for growing
the base because of the wide, dense layer of black rootlets. plants or sawed into squares and used as a substrate for
The fronds are widest above the middle and taper toward growing epiphytic orchids. In nature, many ferns and other
the base to a short stipe. The trunk apex and stipes are cov- epiphytes grow on the root mantles, some species being al-
ered with long hairs. The blades are two- to three-pinnate- most entirely restricted to it.
pinnatifid, up to 3 m (10 ft.) long, and slightly leathery. The similar Dicksonia berteriana (Colla) C. Christensen,
The pinnae are sessile, and the ultimate fertile pinnules are an endemic to the Juan Fernández Islands, is cultivated in
narrowly oblong, each bearing six or more sori. This species a few special collections. It differs from D. sellowiana by
is widespread in the American tropics, where it generally the fertile segments lacking a leafy apex or having only a
grows at elevations above 2000 m (6500 ft.). small one.
In Latin America (particularly Mexico and Central
America) the trunks of Dicksonia sellowiana are harvested
284 C 13
Dicksonia squarrosa (Forster f.) Swartz (many irregular areoles that mostly lack included free vein-
13.40.6 lets). Lepisorus further differs by the presence of peltate
Slender tree fern, rough tree fern, New Zealand paraphyses, at least on young sori. See Polypodium for a fur-
dicksonia ther discussion of related genera.
Semi-tender; tolerates light frost Only two species are found in Dictymia, occurring from
A large tree fern, 5–6 m (16–20 ft.) tall. Grows best under eastern Australia to Fiji. The genus name comes from the
medium light in moist to moist-wet garden soil or potting Greek diktyon, net, alluding to the netted veins.
mix. This species prefers cool, humid climates. It tends to
form offshoots from stolons and can be propagated by re- Dictymia brownii (Wikström) Copeland
moving the offshoots. 13.41.1
The trunk of Dicksonia squarrosa is slender and black, Semi-tender (?)
and the stipes are rough with stiff hairs when young. The A small-medium fern with creeping rhizomes. Grows un-
rachises and costae vary from red-brown to blackish, and der medium light in moist potting mix.
the undersides of the blades are glaucous green. This species Dictymia brownii has stiff, leathery, and erect fronds.
is smaller than D. antarctica. It is endemic to New Zealand. The sori are large, round or oblong, and raised. The species
is native to eastern Australia.
a a
b
b
a
Figure 13.43.5.
Diplazium
dilatatum: frond,
bar = 5 cm (2 in.).
Figure 13.43.6.
Diplazium esculentum:
frond, bar = 5 cm (2 in.).
Diplazium esculentum (Retzius) Swartz
13.43.6
syn. Callipteris esculenta (Retzius) J. Small
Vegetable fern
Semi-tender, Zone 9 pinnatifid above. The species is native to the southern
A medium-sized fern, to 60 cm (2 ft.) tall, with erect rhi- United States (Louisiana) and Mexico to northern South
zomes that can become trunk-like on older plants. Grows America.
well under medium to high light in moist-wet garden soil
or potting mix. Colonies of this species resemble a grove of Diplazium proliferum (Lamarck) Thouars
short-stemmed, coarse-leaved tree ferns. The fiddleheads 13.43.8
are edible. New plants frequently arise from root buds. syn. Callipteris prolifera (Lamarck) Bory
Diplazium esculentum is characterized by broad, two- Tender
pinnate fronds and veins that unite along the costae. The A large fern with erect rhizomes that form offshoots. Grows
scales of the rhizomes and stipe bases have black margins well under medium light in moist-wet garden soil or pot-
and forked teeth (use hand lens). It is native to Africa, ting mix; it does best with warm nights. The plants are large
southeastern Asia, and Polynesia; it has been introduced in and coarse but interesting for the large buds on the midrib.
Florida and Louisiana. Stems of older plants may become trunk-like.
Diplazium proliferum has one-pinnate fronds with
Diplazium lonchophyllum Kunze 13.43.7 coarsely serrate-lobed pinnae and netted veins. The scales of
Semi-tender, Zone 9 the rhizomes and stipe bases have black margins and forked
A medium-sized fern with suberect rhizomes that often teeth (use hand lens). This species is native to Africa, Mada-
produce offshoots. Grows well in moist garden soil or pot- gascar, and from Malaysia to Polynesia.
ting mix; not adapted to climates with cool nights.
The blades of Diplazium lonchophyllum are triangular-
lanceolate and two-pinnate at the base but one-pinnate-
288 C 13
Figure
13.43.8. Diplazium subsinuatum (Hooker & Greville) Tagawa
Diplazium 13.43.10
proliferum: syn. Athyrium dubium (D. Don) Ohwi, Diplazium
a. frond, bar lanceum (Thunberg) C. Presl
= 5 cm Semi-hardy, Zone 7(8)
(2 in.); A small fern with branched, medium- to long-creeping rhi-
b. leaf buds, zomes. Grows well under medium light in moist garden
bar = 5 cm soil or potting mix. The plants are useful for edging or as
(2 in.). ground cover.
F F A C 289
a b
Figure 13.45.5.
Doryopteris ludens:
habit, bar = 5 cm
(2 in.).
b c
Figure 13.46.7.
Drynaria rigidula:
a. foliaceous frond
(center) and
humus-collecting
frond (right), b
foliaceous pinnae
of ‘Whitei’ (lower
left), bar = 10 cm
(4 in.); b. pinna
base and sori, bar =
2.5 cm (1 in.).
a
a
China, Malaysia, Indonesia, the Philippines, Australia, Figure 13.46.8. Drynaria sparsisora: a. foliaceous frond
New Guinea, and southeastern Pacific Islands. (left) and humus-collecting frond (right), bar = 10 cm
‘Whitei’. Plate 25. Pinnae often broader, lacerate to ir- (4 in.); b. rhizome scales, bar = 3 mm (0.1 in.); c. sori,
regularly lobed. bar = 2.5 cm (1 in.).
d
a
e
b
b
Figure 13.47.4. Dryopteris affinis: a. frond, bar = 5 cm
(2 in.), after Hoshizaki and Wilson (1999); b. stipe
scales, bar = 1 cm (0.4 in.), after Hoshizaki and Wilson
Figure 13.47.3. Dryopteris aemula: a. frond, bar = 5 cm (1999); c. pinnules from medial pinnae, bar = 1 cm
(2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After (0.4 in.); d. subsp. borreri, bar = 5 cm (2 in.); e. subsp.
Hoshizaki and Wilson (1999). cambriensis, bar = 5 cm (2 in.).
F F A C 299
Figure 13.47.6.
Dryopteris
case the names may correctly apply to D. filix-mas. In the
amurensis:
United States the hybrid between D. affinis and D. filix-
a. frond, bar =
mas, known as D. complexa (misnamed D. filix-mas ‘Un-
5 cm (2 in.);
dulata-Robusta’) was widely sold.
b. stipe scales,
subsp. borreri (Newman) Fraser-Jenkins. Figure
bar = 1 cm
13.47.4d. Fronds lanceolate to oblong, bases truncate, sur-
(0.4 in.). After
faces not glossy; deciduous in early winter in temperate
Hoshizaki and
climates.
Wilson (1999).
subsp. cambriensis Fraser-Jenkins. Figure 13.47.4e.
Fronds oblanceolate to linear-elliptic, bases tapering, upper
surfaces slightly glossy; deciduous in temperate climates.
‘Congesta’. Figure 13.47.5b. Fronds dwarf, the pinnae
and pinnules congested but neatly overlapping. a
‘Congesta Cristata’. Figure 13.47.5d. Fronds dwarf,
congested, crested.
‘Crispa’. Crisped male fern. In Victorian times this b
name applied to plants with dwarf and broad fronds, to 20
cm (8 in.) long and 14 cm (5.5 in.) wide, with ruffled and
congested foliage. Plants in the current trade are normal-
sized with twisted segments. They are perhaps correctly
called ‘Paleaceo Crispa’.
300 C 13
b
Figure 13.47.10. Dryopteris
×boottii: frond, bar = 5 cm
(2 in.). After Hoshizaki
and Wilson (1999).
Dryopteris carthusiana (Villars) H. P. Fuchs Dryopteris caucasica (A. Braun) Fraser-Jenkins & Corely
13.47.12 13.47.13
syn. Dryopteris spinulosa (O. F. Müller) Watt Hardy, Zone 5
Spinulose wood fern, toothed wood fern, narrow buckler A medium-sized fern, to 80 cm (32 in.) tall, with deciduous
fern fronds and erect rhizomes that form offshoots. Grows well
Very hardy, Zone 2 under medium light in moist garden soil or potting mix.
A medium-sized fern, to 40 cm (16 in.), with ascending to Dryopteris caucasica resembles D. filix-mas (one of its
erect rhizomes. Grows well under medium light in moist, parents) but differs by paler blades, pinnae widest at the
acidic garden soil or potting mix. The plants are deciduous middle (not the base), acute segments that are doubly
and unsuited to subtropical climates. toothed at the apex with acute teeth, and white, lacerate in-
The blades of Dryopteris carthusiana are two-pinnate- dusia. It is native to the Middle East.
pinnatifid to three-pinnate and narrowly triangular-lance-
olate. The lowermost, downward-pointing pinnule is less
than two times as long as the basal upward-pointing pin- a
nule above it. Dryopteris carthusiana is distinguished from
the similar D. dilatata by its medium to light green fronds,
flat margins, and stipe scales of one color. It also resembles
D. intermedia but lacks the minute glandular hairs charac-
teristic of that species. The species is native to North Amer- Figure 13.47.13.
ica, Europe, and Asia. Dryopteris caucasica:
A foliose form of Dryopteris carthusiana is often sold in a. frond, bar = 5 cm
the trade and incorrectly identified as D. stewartii. (2 in.); b. stipe
scales, bar = 1 cm
(0.4 in.). After
Hoshizaki and
Wilson (1999).
Figure 13.47.14.
Dryopteris celsa:
a. frond, bar = c
5 cm (2 in.);
b. stipe scales, bar
= 1 cm (0.4 in.).
After Hoshizaki
and Wilson
(1999).
b
a
b
Figure 13.47.15. Dryopteris championii: a. frond, bar =
5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.);
c. pinnule from medial pinna, bar = 5 mm (0.2 in.);
Dryopteris championii (Bentham) C. Christensen ex d. bullate scales from costa, bar = 1 mm. After Hoshizaki
Ching 13.47.15 and Wilson (1999).
Champion’s wood fern
Hardy, Zone 5
A medium-sized fern, to 90 cm (36 in.), with erect rhi-
zomes and evergreen fronds. Grows well under medium to Dryopteris clintoniana is a hybrid of D. cristata and D.
high light in moist-dry garden soil or potting mix. In the goldiana. It resembles D. cristata, but its basal pinnae are
northeastern United States the fertile leaves are not pro- longer than they are wide and not twisted into a horizontal
duced until the late fall. This species is particularly attrac- plane. The species is native to northeastern North America.
tive in winter because its evergreen fronds remain upright,
not reclining as in many species with evergreen fronds. Dryopteris complexa Fraser-Jenkins 13.47.17
Dryopteris championii has narrowly triangular fronds Robust male fern
and pinnules that are eared on both sides at the base. The Hardy, Zone 4(5)
stipe, rachis, and costae bear a dense covering of shiny red- A medium-large fern, to 130 cm (52 in.), with erect rhi-
brown scales, and those on the costae become strongly in- zomes that form offshoots and deciduous to semi-evergreen
flated. The species is native to eastern Asia. fronds (in warmer climates). Grows well under medium
light in moist to moist-dry garden soil or potting mix. The
Dryopteris clintoniana (D. C. Eaton) Dowell plants are easy to cultivate and become rapid, vigorous
13.47.16 growers with full foliage. The fronds are deciduous, but less
Clinton’s wood fern so in warmer climates.
Very hardy, Zone 3 Dryopteris complexa is a fertile hybrid of D. affinis and D.
A medium-sized fern, to 95 cm (38 in.), with short-creep- filix-mas. The stipe scales are narrowly triangular and mem-
ing rhizomes and deciduous fronds. Grows well under me- branous. The middle and distal pinnae often overlap
dium light in acidic, moist-wet garden soil or potting mix. slightly, and the pinna midrib has a blackened spot on the
304 C 13
c
c
a
b
a
natifid blades, longer and narrower segments with margins Dryopteris cycadina (Franchet & Savatier)
entire or weakly crenate-serrate, and rounded and entire C. Christensen 13.47.20
apices. The costae lack a black blotch at the base. The spe- Shaggy wood fern, black wood fern
cies is native to northeastern Asia. Hardy, Zone 6
A medium-sized fern, to 1 m (3 ft.), with erect rhizomes
Dryopteris cristata (Linnaeus) A. Gray and semi-evergreen fronds. Grows well under medium
13.47.19 light in moist-dry garden soil or potting mix.
Crested wood fern The common names of Dryopteris cycadina come from
Very hardy, Zone 3 the dense, dark scales that cover the stipes and rachises. The
A medium-sized fern, to 50 cm (20 in.) tall, with deciduous blades are one-pinnate and leathery with about 30 pairs of
fronds and short-creeping to erect rhizomes that form off- pinnae, these mostly coarsely serrate to lobed one-third of
shoots. Grows well under medium light in moist-wet gar- the way to the costa. The basal pinnae point downward.
den soil or potting mix. The plants are easy to cultivate. The species is native to eastern Asia.
The fronds of Dryopteris cristata are one- to two-pin- The cultivated plants of Dryopteris cycadina are often
nate, narrowly lanceolate, and often have pinnae oriented misidentified as D. atrata (Wallich) Ching or D. hirtipes
more or less horizontally—the “venetian blind” effect. The Kuntze.
basal pinnae are triangular, and fertile fronds are taller and
more erect than sterile ones. The fertile pinnae occur mostly Dryopteris cystolepidota (Miquel) C. Christensen
toward the apex of the blade and are slightly contracted. 13.47.21
The species is native to the northern and southeastern syn. Dryopteris nipponensis Koidzumi
United States. It commonly hybridizes in nature with D. Hardy, Zone (5)6
intermedia to produce D. ×boottii. A medium-sized fern, to 60 cm (2 ft.), with branched,
short-creeping rhizomes and evergreen, glossy fronds.
Grows well under medium light in moist garden soil or pot-
ting mix. The plants are easy to grow.
The blades of Dryopteris cystolepidota are broadly trian-
gular, up to nearly three-pinnate at base, and thin-leath-
ery. The segments have small spines turned up from the
surface. The species is native to Japan and Korea.
b
a
b
306 C 13
a
b
b
Dryopteris dilatata (Hoffmann) A. Gray
13.47.23, 24
Broad wood fern, broad buckler fern
Hardy, Zone 4(5)
A medium to large fern, to 1 m (3 ft.) or more, with decid-
c uous fronds and ascending to erect rhizomes that form off-
shoots. Grows well under low to medium light in acidic,
e
e
c
d
cultivate. New growth has an attractive reddish coppery Dryopteris expansa has tan stipe scales with a dark central
color, turning dark, shiny green at maturity. New fronds stripe. The blades lack glands and are broadly triangular
are produced throughout the growing season, and in win- and three-pinnate-pinnatifid at the base, with the down-
ter the fronds remain upright, not reclining as in many ward-pointing pinnule next to the rachis two to three times
evergreen ferns. longer than the opposite upward-pointing pinnule. The
Dryopteris erythrosora is highly variable but can be char- pinnule margins are flat, not enrolled. The species is native
acterized as follows: the stipe scales are blackish and the to North America, Europe, and eastern Asia. It is a parent
blades are two-pinnate, mostly oblong-ovate, broad, and of D. campyloptera and is difficult to distinguish from the
arching. The lowest basiscopic pinnules next to the rachis offspring.
are short and have marginal teeth spinulose-tipped and in-
curved. The costal scales are strongly inflated and dark to- Dryopteris filix-mas (Linnaeus) Schott
ward the tip. Set close to the midrib, the indusia are red or, 13.47.27, 28
rarely, greenish white (forma viridosora (Nakai ex H. Ito) Common male fern
H. Ito). The species is native to eastern Asia. Hardy, Zone 4(5)
The variability of this species has caused considerable A large fern, to 130 cm (52 in.), with deciduous fronds and
confusion in the trade. Most of the current plants circulat- short-creeping to erect rhizomes that form offshoots.
ing as Dryopteris bissetiana and D. purpurella are actually Grows well under low to medium light in moist garden soil
D. erythrosora. or potting mix. The usually deciduous fronds become semi-
forma prolifica (Maximowicz ex Franchet & Savatier) evergreen if the plants are grown in warmer climates. This
H. Ito. Blades shorter, 37 cm (15 in.), more triangular, usu- species is easy to grow and has many garden variations.
ally with buds; pinnules narrow-linear, pointed. Dryopteris filix-mas is of hybrid origin (but fertile) from
a naturally occurring cross between D. caucasica and D. ore-
Dryopteris expansa (C. Presl) Fraser-Jenkins & Jermy ades. The fronds are one-pinnate-pinnatisect (nearly two-
13.47.26 pinnate) with segments slightly tapered and toothed along
syn. Dryopteris austriaca (Jacquin) Woynar
Arching wood fern, northern spreading wood fern
Very hardy, Zone 3(4)
A medium-sized fern, to 60 cm (2 ft.), with ascending or
erect rhizomes and deciduous fronds. Grows well under
low to medium light in acidic, moist garden soil or potting
mix. The plants do not do well in subtropical climates.
d
b
a b
a
a c
d
h i
f g
Figure 13.47.28. Dryopteris filix-mas cultivars: a. ‘Linearis Polydactyla’, an extreme form; b. ‘Linearis Polydactyla’; c.
‘Cristata’; d. ‘Grandiceps’; e. ‘Linearis Congesta’; f. ‘Barnesii’; g. ‘Crispa Cristata’; h. a dwarf, crisped, crested cultivar; i.
‘Ramo Cristata’. Bar = 5 cm (2 in.).
310 C 13
the sides and apex. The teeth are acute and typically point
toward the segment apex. The upper surface is puckered
opposite the sori, and the young indusia have a flat, thin
rim all around the edge. The species is native to North
America and Europe. c
‘Barnesii’. Figure 13.47.28f. Barne’s male fern. Fronds
upright, narrow; pinnae short, to 5 cm (2 in.) long, wide,
blunt; pinnules narrowed at base.
‘Crispa Cristata’. Figure 13.47.28g. Frond apices and
pinnae crested, pinnules ruffled.
‘Cristata’. Figure 13.47.28c. Frond apices and pinnae
crested; many different types.
‘Grandiceps’. Figure 13.47.28d. Frond apices heavily
tasseled, pinna tips less so.
‘Linearis Congesta’. Figure 13.47.28e. Fronds to 37 cm
(15 in.), pinnae crowded, pinnules narrow. b
‘Linearis Polydactyla’. Slender crested male fern. Figure
13.47.28a, b. Blades broad and elliptic, the blade and pinna
apices crested, the segments finger-like, the segments of the a
pinnae skeletonized, mostly linear to nearly filiform. May
grow better in acidic soil.
‘Ramo Cristata’. Figure 13.47.28i. Fronds to 60 cm (24
in.), branched and crested.
‘Undulata-Robusta’. See Dryopteris complexa.
Figure 13.47.29. Dryopteris formosana: a. frond, bar =
5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.);
Dryopteris formosana (H. Christ) C. Christensen c. pinnule from medial pinna, bar = 5 mm (0.2 in.).
13.47.29 After Hoshizaki and Wilson (1999).
Hardy, Zone (6)7
A medium-sized fern, to 45 cm (18 in.), with short-creep-
ing to erect rhizomes and evergreen fronds. Grows well
under medium light in moist garden soil or potting mix.
Dryopteris formosana has broad, pentagonal fronds up
to three-pinnate. The lowest basiscopic pinnule next to the
rachis is quite long, and the segment teeth are short-aristate
and tend to turn upward from the plane of the blade. The
scales on the costa undersides are weakly inflated, appearing
puffy (not flat). The species is native to Japan, Taiwan, and
the Philippines.
Figure 13.47.31.
Dryopteris fuscipes:
a. frond, bar =
5 cm (2 in.); b. Figure 13.47.32. Dryopteris goldiana: a. frond, bar =
stipe scales, bar = 5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After
1 cm (0.4 in.). Hoshizaki and Wilson (1999).
After Hoshizaki
and Wilson
(1999).
lie close to the midrib. The species is native to eastern
North America.
a c Figure 13.47.34.
Dryopteris
intermedia:
a. frond, bar =
5 cm (2 in.);
b. stipe scales, bar
= 1 cm (0.4 in.).
b After Hoshizaki
and Wilson
(1999).
a b
This plant is often confused with Dryopteris pycnopter- as wide at the base as in the middle, and the pinnae are pin-
oides (H. Christ) C. Christensen of Japan, a larger plant natisect with many oblong, shiny segments. Segments have
with sori scattered over the pinnae. rounded apices bearing sharp teeth. The species is native
from eastern India to China and Taiwan.
Dryopteris lacera (Thunberg) O. Kuntze
13.47.36
Hardy, Zone 5
Figure 13.47.37.
A medium-sized fern, to about 60 cm (2 ft.), with ascend- Dryopteris
ing to erect rhizomes and deciduous fronds. Grows well lepidopoda: a.
under medium light in moist garden soil or potting mix. frond, bar = 5 cm
This species is easy to cultivate. The normally deciduous (2 in.); b. stipe
fronds tend to be slightly evergreen in warmer climates. scales, bar = 1 cm
The broad lanceolate blades of Dryopteris lacera are one- (0.4 in.). After
pinnate-pinnatifid to two-pinnate and fertile only toward Hoshizaki and
the tip. The segments or lobes are entire or nearly so, unlike Wilson (1999).
many Dryopteris species that are lobed. At maturity the
fronds are glabrous, without scales on the rachises or costae.
The fertile pinnae are more contracted than the sterile and
wither early. The species is native to eastern Asia.
Figure 13.47.36.
Dryopteris lacera: a.
frond, bar = 5 cm (2 in.); a
b. stipe scales, bar = 1 cm
(0.4 in.). After Hoshizaki
and Wilson (1999).
a
a
b
c
Figure 13.47.38. Dryopteris ludoviciana: a. frond, bar =
5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After
Hoshizaki and Wilson (1999). Figure 13.47.39. Dryopteris marginalis: a. frond, bar =
5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.);
c. medial pinna, bar = 1 cm (0.4 in.). After Hoshizaki
and Wilson (1999).
Dryopteris marginalis (Linnaeus) A. Gray
13.47.39
Eastern wood fern, marginal shield fern
Very hardy, Zone 2(3)
A medium-sized fern, to 60 cm (2 ft.), with erect rhizomes
and evergreen fronds. Grows well under medium light in
moist garden soil or potting mix. The plants are easy to
grow and have bluish green blades. They do not do well in
subtropical climates.
Dryopteris marginalis can be distinguished from many Figure 13.47.40. Dryopteris
other Dryopteris by having sori that are close to the margin. mindschelkensis: a. frond, bar =
The blades are one-pinnate-pinnatifid to two-pinnate and 5 cm (2 in.); b. stipe scales,
slightly leathery. The species is native to northeastern bar = 1 cm (0.4 in.). After
North America, where it grows on or among rocks. Hoshizaki and Wilson (1999).
a a d
b
c
a
Figure 13.47.46.
Dryopteris
sacrosancta: a.
frond, bar = 5 cm
Figure 13.47.45. Dryopteris remota: a. frond, bar = 5 cm
(2 in.); b. stipe
(2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After
scales, bar = 1 cm
Hoshizaki and Wilson (1999).
(0.4 in.). After
a Hoshizaki and
Wilson (1999).
Dryopteris sacrosancta has large stipe scales with pale bor-
ders, and the scales on the undersides of the costae are
weakly inflated (puffy) and dark tipped. The blades are tri-
angular, three-pinnate, slightly glossy and leathery, and flat
margined. The lowermost downward-pointing pinnule
next to the rachis is elongate. The species is native to Japan,
China, and Korea.
c Figure 13.47.49.
Dryopteris stewartii:
a a. frond, bar = 5 cm
(2 in.); b. stipe
scales, bar = 1 cm
(0.4 in.); c. basal
pinna, bar = 4 cm
(1.5 in.). After
Hoshizaki and
Wilson (1999).
b
c
ting mix. This species grows best in mildly cool, humid cli-
mates. The fronds are arranged in a handsome shuttlecock
pattern.
Dryopteris wallichiana is characterized by one-pinnate-
pinnatifid blades with 20–35 pairs of sessile pinnae. The
segments of the pinnae are squared or oblong. The rachis is
conspicuously covered by black, linear or thread-like scales.
The species is native to subtropical and tropical mountains
of the world.
Elaphoglossum crinitum (Linnaeus) H. Christ and heart-shaped. The blades of Elaphoglossum decoratum
13.48.1, 4 are linear-lanceolate to oblong, dark, thick, leathery,
syn. Hymenodium crinitum (Linnaeus) Fée abruptly short-acuminate at the apex, and roundish to
Elephant-ear fern acute at the base. The surfaces of the blade are covered by
Tender minute, star-shaped scales; these are much less conspicu-
A small-medium fern with short-creeping rhizomes. Grows ous than the marginal scales. The species is native to trop-
best with year-round heat under low to medium light in ical America.
well-drained, moist-wet potting mix or uncut moss.
Elaphoglossum crinitum is a striking species, character- Elaphoglossum herminieri (Bory ex Fée) T. Moore
ized by large, oblong blades with scattered, erect, black 13.48.6
scales on both surfaces. Unlike most Elaphoglossum species, Tender
the veins are netted (without included veinlets). The species A medium to large fern with short-creeping rhizomes and
is native to the American tropics. often long, pendulous fronds. Grows under low to medium
light in drained, moist potting mix or uncut moss. The
Elaphoglossum decoratum (Kunze) T. Moore fronds are attractive for their bluish green color and pen-
13.48.5 dulous habit.
Tender Elaphoglossum herminieri is characterized by pendent,
A small-medium fern with short-creeping rhizomes. Grows nearly sessile fronds up to 1 m (3 ft.) or more long. Linear,
under low to medium light in well-drained, moist potting metallic blue-green, leathery blades are dotted with minute,
mix or uncut moss. This species is difficult to grow. Showy star-shaped, glandular scales on both surfaces. The fertile
scales along the stipes and margins of the blades make for fronds, which are rarely produced, are rounded or kidney-
attractive foliage.
The scales on the stipes and blade margins are about 3.5
mm (about 0.1 in.) long, overlapping, yellowish brown,
Figure 13.48.5.
Elaphoglossum
decoratum: a. habit,
bar = 5 cm (2 in.);
b. marginal scale,
bar = 1 mm.
b a
shaped and much smaller than the sterile fronds. The spe- The creeping rhizomes of Elaphoglossum maxonii are 8–
cies is native to tropical America. 15 mm (0.3–0.6 in.) wide, with phyllopodia hidden among
the rhizome scales, the scales being narrowly triangular,
Elaphoglossum luridum (Fée) H. Christ golden or light brown, acuminate, and irregularly ciliate.
13.48.7 The fronds are erect, leathery, and have ovate to narrowly
syn. Elaphoglossum schomburgkii (Fée) T. Moore elliptic blades, decurrent at the base. Minute and irregular
Tender star-shaped scales often with blackened centers are scattered
A medium-sized fern with short-creeping to nearly erect on both blade surfaces. The fertile fronds are smaller than
rhizomes. Grows well under low to medium light in well- the sterile ones. The species is native to the Greater Antilles.
drained, moist potting mix or uncut moss.
Elaphoglossum luridum is easy to recognize by its ob-
lanceolate, medium-sized blades and dark, shaggy scales on
the stipes and midribs. The lower surfaces of the blades
have scattered small scales, and the rachises have dark star-
shaped scales with twisted rays. The species is native to low- Figure 13.48.8. Elaphoglossum
land forests of the American tropics. maxonii: habit, bar = 5 cm (2 in.).
a
b
collected in Mexico. Plants from elsewhere should be re-
ferred to the E. latifolium complex, which also has not been
fully studied. The rarely grown E. maxonii, also of the E.
latifolium complex, differs from E. sartorii by its thicker
rhizome and phyllopodia hidden by scales.
Elaphoglossum vestitum (Schlechtendal & Chamisso) Figure 13.48.12. Elaphoglossum vestitum: a. habit, with
Schott ex T. Moore 13.48.12 fertile frond on left, sterile fronds on right, bar = 5 cm
Semi-tender to tender (2 in.); b. blade scales, bar = 1 mm.
A small-medium fern with short-creeping rhizomes. Grows
well under low to medium light in drained, moist potting
mix or uncut moss.
Elaphoglossum vestitum has distinct phyllopodia and rhi- Equisetum 13.49.1
zome scales that are red-brown, shiny, linear-lanceolate, Horsetail, scouring rush
toothed, and 4–6 mm (0.2 in.) long. The stipes are slender Equisetum is usually planted around ponds, bogs, and
and covered with narrowly triangular, red-brown, ciliate marshes, or it can be grown in pots, tubs, and small dish-
scales. The narrowly elliptic, gray-green blades are densely gardens. Garden soil or potting mix are suitable if the plants
covered on the upper surface with whitish scales bearing are grown in pots and the soil is kept moist and drained. If
filiform rays. Similarly shaped hairs on the undersurface are the pots are set in standing water, a sand and peat-moss mix
reddish brown. The species is native to southern Mexico. should be used to discourage the soil from “souring.” Vig-
This species has been misidentified in horticulture as orously growing species such as Equisetum hyemale become
Elaphoglossum muelleri (Fournier) C. Christensen, which weedy in gardens and are difficult to eradicate because of
lacks phyllopodia and is not cultivated. their deep rhizomes. Equisetum is often used in Japanese
flower arrangements.
The plants can be propagated by dividing the rhizomes
or by burying the stem segments shallowly in the soil.
When buried in soil, new roots are eventually produced at
326 C 13
the joints of the stem segments, and new stems soon follow.
This method of producing new growth is how sterile hy-
brids of Equisetum, which cannot reproduce by spores be-
cause they abort, can become locally abundant or spread to
distant sites where their parents do not grow. b
The genus is easy to recognize by its green, usually hol-
low, jointed stems and spore-bearing cones at the stem tips;
the cones, however, are not always present. The sheath
around the stem joints represents a whorl of greatly reduced
F F A C 327
The sterile stems of Equisetum arvense have dense whorls Equisetum ×ferrissii Clute 13.49.4
of branches from the joints, and sheaths of the main, erect Ferriss’s scouring rush
stems tend to flare slightly outward and bear 8–12 teeth, Very hardy, Zone 3
some adhering in pairs. The fertile stems appear in the A medium-sized plant with creeping rhizomes that send up
spring before the sterile ones. They are smaller, nongreen, irregularly spaced or clumped stems. Grows under medium
and short-lived, withering soon after shedding their spores. to high light in moist-wet to dry garden soil. When grown
This species is widely distributed in temperate North in pots, this hybrid produces fewer stems compared to its
America, Europe, and Asia. parents.
Equisetum ×ferrissii is of hybrid origin between E. hye-
Equisetum diffusum D. Don 13.49.3 male and E. laevigatum, and its characteristics are interme-
Semi-tender and perhaps hardier, Zone (8)9 diate between those two species. For example, E. hyemale
A medium-sized plant with creeping, branching rhizomes. has evergreen stems, and E. laevigatum has deciduous ones;
Grows in medium to high light in moist to wet garden soil. the hybrid’s stems are intermediate, being evergreen toward
The plant’s closely placed stems produce an attractive pot the base but deciduous in the upper part. The stem sheaths
plant. of the hybrid are also intermediate: those on the lower stem
The aerial stems of Equisetum diffusum have branches are ashy-white with black borders above and below (as in E.
in regular or irregular whorls. The stems have four ridges, hyemale), but those on the upper part of the stem are green
with each ridge grooved and each side of the groove typi- with a black border only at the top (as in E. laevigatum).
cally lined with a row of closely set, minute, blocky saw The spores of the hybrid are whitish and aborted, not green
teeth (magnification needed). Ovoid-cylindrical cones may and full as in the parental species. Equisetum ×ferrissii is na-
be produced at the tips of the stem late in the growing tive to and widespread in North America.
season (July–November). Tattered late-season stems are
quickly replaced by new growth. This species is endemic
to the Himalayas.
Figure 13.49.4. Equisetum
×ferrissii: a. habit of small
plant, bar = 5 cm (2 in.);
a b. stem with cone, bar =
1 cm (0.4 in.); c. lower stem
sheaths, bar = 1 cm (0.4 in.).
a
b
Figure 13.49.3. Equisetum diffusum: a. habit of stems
from a young plant, bar = 5 cm (2 in.); b. outline of
stem cross-section, enlarged to show stem with grooves
in each of the four ridges.
b c
328 C 13
Equisetum hyemale Linnaeus 13.49.1, 5 Equisetum laevigatum A. Braun 13.49.6
Scouring rush, rough horsetail, winter scouring rush Smooth horsetail
Very hardy, Zone 2 Very hardy, Zone 3
A medium to large plant with short- to long-creeping rhi- A medium-sized plant with creeping rhizomes that send up
zomes. Grows well under medium to high light in moist irregularly spaced or clumped stems. Grows well under me-
to wet garden soil. It is easy to grow but becomes weedy in dium to high light in moist-wet garden soil.
gardens and is difficult to eradicate because of its deep rhi- Equisetum laevigatum is characterized by smooth, usu-
zomes. When grown in full sun the plants are more likely to ally unbranched (although sometimes sparsely and irregu-
produce cones. larly branched, especially if injured), deciduous stems and
Equisetum hyemale is characterized by rough, evergreen green (not ash-colored) sheaths with a black rim. The teeth
stems, and sheaths with a wide, ash-gray band bordered by at the rim typically fall off early and are therefore absent.
a lower black band and an upper black rim. The stems are The cones are blunt at the apex. The species is native to
unbranched, but injury to the apex will result in lateral North America, often occurring in open, sunny habitats.
branches developing at some of the lower joints. The teeth
are 14 or more at a joint, but most of them fall off at matu-
rity. The cones have a minute point at the tip. This species
is native to North America, Europe, and Asia. It typically
forms large colonies along shaded riverbanks, streams, and
in railroad ballast.
The thick-stemmed selection from the Washington area
reverted to typical stem diameters when pot cultivated in
southern California, whereas the slender-stemmed variants
from Alaska maintained their distinctness.
a
b
b d
b c b
a
Figure 13.49.7. Equisetum scirpoides: a. habit, bar = 5 cm
(2 in.); b. stem with cone, bar = 1 mm; c. sheath, bar =
1 mm. Figure 13.49.8. Equiseum telmateia: a. habit of sterile
stem, bar = 5 cm (2 in.); b. dimorphic fertile stem with
cone, bar = 5 cm (2 in.).
b c
Figure 13.50.1. Gleichenia
dicarpa: habit, bar = 5 cm
(2 in.).
on its fronds, and for this reason is widely used in Japan for
New Year’s Day decoration. In parts of southeastern Asia,
baskets are woven from the tough, flexible rachises of several
species. Sticherus flabellatus (R. Brown) H. St. John, S. lo-
batus N. A. Wakefield, and Dicranopteris linearis (N. L. Bur-
man) Underwood are harvested in Australia and exported to
Japan and elsewhere for cut-foliage use.
Goniophlebium
syn. Polypodium, in part; Schellolepis
Most gardeners know the genus Goniophlebium as Poly-
podium. It differs, however, by having jointed pinnae and
goniophleboid venation—that is, several rows of regularly
shaped, angular areoles with one free, included veinlet
pointing toward the pinna margin. Some authors disregard
the jointed pinnae and instead emphasize the presence of
scattered, black, fibrous strands in the rhizome, clathrate
rhizome scales, and (usually) hairy and scaly paraphyses in
the sori. In this book, the species with nonjointed pinnae
Figure 13.50.2. Sticherus are left in Polypodium (P. amoenum and P. formosanum),
flabellatus: habit, bar = except for the hybrid Goniophlebium ×ekstrandii. The
5 cm (2 in.). plants with long fronds, such as the knight’s polypody (Go-
niophlebium subauriculatum ‘Knightiae’), are handsomely
displayed in hanging baskets, where their fronds may reach
2.6 m (8.5 ft.) long.
The rhizome of Goniophlebium is creeping and often
glaucous, and when viewed in cross section, many scat-
tered, black fibers appear among the circle of vascular bun-
dles. The rhizome scales are clathrate, and the blades vary
from pinnatifid to one-pinnate. The pinnae are jointed,
rarely scaly, with entire or toothed margins. The veins are
netted, and the areoles have a regularly angular shape, those
rooted. Because the United States Department of Agricul-
along the midribs large and containing a veinlet that points
ture requires that soil be removed from imported plants,
outward (toward the pinna margin). The sori are round-
any imported Gleicheniaceae stand little chance of success.
ish, in one row on either side of the costa, and terminal on
It is generally thought that gleichenias need a mycorrhizal
the veinlet contained within the areole. Usually hairs or
fungus in their roots to grow. The successes with spore-
scales are mixed among the sporangia. Like all polypodia-
grown plants, however, raise questions about this need.
ceous ferns, this genus lacks indusia.
Members of this family can usually be recognized by
Goniophlebium, which contains about 23 species, is na-
their forked pinnae with a dormant bud in the fork’s angle.
tive to Asia, Australia, and some Pacific Islands. The genus
In nature the plants are terrestrial, not epiphytic, although
name comes from the Greek gonia, angled, and phleps, a
their leaves may rest on other plants for support. The rhi-
vein, referring to the angular mesh of veins.
zomes are always long-creeping, and the fronds are scram-
bling or trailing, one to many times forked, with the stipe
containing a single, C-shaped vascular bundle. The pinnae Goniophlebium ×ekstrandii Hoshizaki
are one to several times forked and have free veins. The sori 13.51.1
are always round, located on the undersurface of the leaf, Semi-tender
and lack an indusium. A medium-large fern with long-creeping rhizomes. Grows
In Alabama and Florida the native Dicranopteris flexuosa under medium light in moist, drained potting mix.
(Schrader) Underwood has not become well established, Goniophlebium ×ekstrandii has one-pinnate fronds with
and colonies are known to have died out. The Hawaiian pinnae broadly attached to the rachis. The foliage is inter-
Islands have several native species of gleichenoids, some of mediate in shape between that of its parents, Polypodium
which form extensive stands. formosanum and Goniophlebium subauriculatum. This ster-
Plants of this family have only a few uses. A Japanese spe- ile hybrid is known only in horticulture, and its cultivar
cies of Diplopterygium has yearly growth that is well marked name is ‘Nola’.
332 C 13
a a
Goniophlebium persicifolium resembles G. percussum but The fronds of Goniophlebium subauriculatum are up to
differs by lacking needle-shaped hairs on the blades. In ad- 2 m (6.5 ft.) long and one-pinnate with a pinnatifid apex.
dition, the sori of this species are only slightly sunken, There are many pinna pairs and these are jointed, mostly
wider than 1.5 mm, and lack the ring of dark scales. Gonio- cordate at the base, and sessile or nearly so. The species is
phlebium persicifolium is native from India to China, south- native to China, southeastern Asia, Indonesia, Australia,
eastern Asia to Malaysia, Indonesia, and the Pacific Islands. and the Philippines.
‘Knightiae’. (Polypodium knightiae Horticulture).
Knight’s polypody. Figure 13.51.4c; Plate 28. Pinnae ir-
Goniophlebium subauriculatum (Blume) C. Presl
regularly and deeply incised. The plants tolerate more cold
13.51.4
(semi-tender) and more irregular watering than the species.
syn. Polypodium subauriculatum Blume
Jointed polypody
Tender Grammitidaceae 13.52.1
A large fern with short- to medium-creeping rhizomes. Grammitid family
Grows well under medium light in moist, drained potting Although not cultivated in the United States, the family
mix. The fronds are long and pendulous, and the old fronds Grammitidaceae is included here because of its large size
are shed in early to mid-spring as new ones emerge. and horticultural potential. Most plants in the family are
small, tropical epiphytic ferns. Their attractive frond shapes
and delicate aspect are reminiscent of tiny polypodiums;
other species have fronds that resemble narrow strips
Figure 13.51.4. Goniophlebium trimmed by pinking shears. Among the genera in this fam-
subauriculatum: a. frond, bar = ily are Acrosorus, Adenophorus, Calymmodon, Cochlidium,
5 cm (2 in.); b. pinna, bar = Ctenopteris, Grammitis, Lellingeria, Melpomene, Micropoly-
1 cm (0.4 in.), after Hoshizaki podium, Prosaptia, Scleroglossum, Terpsichore, Themelium,
(1982); c. ‘Knightiae’ frond, and Zygophlebia. They are important in the tropics, espe-
bar = 5 cm (2 in.). cially in montane and cloud forests. Some species occur in
subtropical or warm-temperate forests where humid con-
ditions prevail. The Grammitidaceae contain about 450
species.
Although a few species were grown in Victorian times
and some have been recommended for cultivation, no one
seems to have cultivated the plants long-term. Usually they
languish in terrariums and eventually die. Their cultural
requirements are unknown. The proper combination of air
movement, humidity, and light seem to be the main factors
affecting their growth, but nutrients and mycorrhizal fungi
might also be important.
The senior author has established Cochlidium serrulatum
(Swartz) L. E. Bishop in a 3.8 liter (1 gal.) glass bottle (may-
onnaise jar type) covered with a piece of glass. Medium in-
direct light is supplied through a large north-facing win-
dow. The fern grows in moist uncut sphagnum moss, which
was lightly firmed into a 60-degree angle against one side of
the bottle. Room temperature ranges from 16 to 24°C
(60–75°F). Ordinary tap water is used but seldom required
c in the covered container. The plant appeared as a volunteer
in a small clump of moss (collected in Costa Rica) that was
planted at the top of the sphagnum moss slope. The plant is
now over a year old, thriving and producing spores.
a The plants resemble small polypodiums because of their
pinnatifid or one-pinnate blades and round sori that lack
b indusia. They differ by having green, tetrahedral spores
(not yellow, bean-shaped ones) and, usually, the rhizomes
are erect or short-creeping, not long-creeping, as in many
polypodiums.
334 C 13
b c
d
a
f
e g
two vascular bundles, broadly triangular to pentagonal Gymnocarpium dryopteris (Linnaeus) Newman
blades, and round (rarely elliptic), non-indusiate sori. The 13.53.4; 29
blades vary from pinnatifid to three-pinnate-pinnatifid and syn. Polypodium dryopteris Linnaeus
may have minute glands, but hairs are lacking. There is usu- Common oak fern
ally a slight swelling where the pinna joins the rachis. The Very hardy, Zone (3)4
veins are free, not netted. A small fern, 15–25 cm (6–10 in.) tall, with slender, long-
The genus consists of five species and is native to North creeping rhizomes and deciduous fronds. Thrives under
America, Europe, and Asia. Its name comes from the Greek low to medium light in moist garden soil or potting mix. Be
gymnos, naked, and karpos, fruit, referring to the sori careful not to allow the soil to dry.
(“fruit”) that lack an indusium. Gymnocarpium dryopteris has blades that are triangular-
pentagonal, two-pinnate-pinnatifid, 3–14 cm (1.2–5.5 in.)
Gymnocarpium disjunctum (Ruprecht) Ching long, and glabrous. The second pinna pair from the base is
13.53.1–3 sessile, and the basal pinnules on both sides are essentially
Western oak fern equal in length. The sori are round. The species is native to
Very hardy, Zone 4 northern North America, north-central Europe, and north-
A medium-sized fern with slender, long-creeping rhizomes ern Asia to China and Japan.
and distant, deciduous fronds. Prefers low to medium light ‘Plumosum’. Pinnules thin, delicate, dense, the margins
in moist garden soil or potting mix. This species is unsuit- turned under.
able for areas with warm winters.
The blades of Gymnocarpium disjunctum are broadly tri- Gymnocarpium oyamense (Baker) Ching
angular-pentagonal, three-pinnate-pinnatifid, 8–24 cm 13.53.5
(3–9 in.) long, and glabrous. The second pinna pair from syn. Currania oyamensis (Baker) Copeland
the base is sessile, and its basiscopic basal pinnule is longer Hardy
than the acroscopic one. The sori are round. The species is A small-medium fern with slender, long-creeping rhizomes
native to northeastern Asia and the northwestern coast of and distant, deciduous fronds. Grows well under medium
North America. light in moist garden soil or potting mix.
Figure 13.53.3. Gymnocarpium disjunctum: a. frond, bar = 5 cm (2 in.); b. frond base, showing uneven basal pinnules in
second pinna pair, bar = 2 cm (0.8 in.).
336 C 13
Figure 13.53.5. Gymnocarpium oyamense: a. frond, bar = Figure 13.53.6. Gymnocarpium robertianum: a. frond,
5 cm (2 in.); b. sori, bar = 2 cm (0.8 in.). bar = 5 cm (2 in.); b. sori, bar = 1 cm (0.4 in.).
F F A C 337
times with a few scales, and they vary from simple and en-
tire to deeply palmately or pinnately lobed to one-pinnate
(noncultivated species can be up to three-pinnate). The
veins are either free or netted, and if the latter, the areoles
lack included veinlets. The sporangia run along the veins;
an indusium is absent. Some species of Hemionitis might be
confused with Bommeria, but the sori in that genus run
along the margin, not between the margin and midrib as in
Hemionitis.
The genus consists of seven species native to the Amer-
ican tropics. Hemionitis arifolia, which questionably be-
longs here, occurs in Asia. The genus name comes from the
Greek hemionos, mule, and is of uncertain derivation.
Histiopteris
Bat fern, bat-winged fern
Histiopteris is a medium- to large-sized terrestrial fern in-
frequently cultivated. Because of its long-creeping rhi-
zomes, it is best confined to large pots or tubs. The com-
mon name alludes to the wing-like appearance of the sessile
pinnules. Histiopteris incisa is widely distributed in the trop-
Figure 13.54.4. Hemionitis palmata: fronds, bar = 5 cm ics and subtropics and seems to vary in size, degree of blue-
(2 in.). gray greenness, and adaptability to cold.
Histiopteris is easily recognized by long-creeping rhi-
zomes; opposite, sessile (or nearly so) pinnae and sessile
pinnules; and enrolled leaf margins that serve as indusia.
Hemionitis pinnatifida Baker 13.54.5 The fronds are distant on the rhizome, and their tips grow
Tender continuously, never completely unfurling. The stipe con-
A small fern with short-creeping to suberect rhizomes and tains one vascular bundle, and the blades are two-pinnate or
bud-bearing fronds. Grows well under medium light in more divided, often glaucous beneath. The basal pinnules
moist potting mix. of each pinna are opposite, often reduced and touching the
Hemionitis pinnatifida resembles H. palmata and is of rachis. The veins vary from free to netted. The sori run
hybrid origin between it and another unknown species in along the blade margin.
the genus. Hemionitis pinnatifida is distinguished by deeply The genus is south-temperate to pantropical (mostly
pinnatifid terminal lobes and several buds that usually de- Malaysia) and contains five species. The genus name comes
velop in smaller marginal notches. It is native to Central from the Greek histion, sail, and pteris, fern, alluding to
America. shape of the basal pinnae.
F F A C 339
Figure 13.56.2.
a Hymenophyllum: sorus
and indusium.
Figure 13.55.1. Histiopteris
incisa: a. frond, bar = 5 cm
(2 in.); b. pinnule with sori
and indusia, bar = 1 cm
(0.4 in.).
340 C 13
c
a
Figure
13.59.5.
Lastreopsis
effusa: frond
and bud, bar =
5 cm (2 in.).
Lecanopteris b
syn. Myrmecophila, Myrmecopteris
Ant fern
The genus Lecanopteris has enlarged and hollow rhizomes
that provide homes for ants. The ants, in turn, protect the
plant by attacking anything that disturbs it. These small- to
medium-sized epiphytes are grown as novelties where tropi-
cal conditions can be provided.
The rhizomes of Lecanopteris are bluish green when
young and become blackish with age. The rhizome scales a
are absent or dense, round, peltate, and marginally mem-
branous. The stipes are jointed to a small raised projection
(phyllopodium), and the sterile blades vary from entire to
d
lobed to deeply pinnatifid. The veins are netted, not free.
The sori are round or oblong, usually sunken, and like all
polypodiaceous ferns, lack indusia. Figure 13.60.1. Lecanopteris carnosa: a. habit, bar = 5 cm
The species with scaly rhizomes, such as Lecanopteris (2 in.); b. fertile portion of frond, upper surface, bar =
sinuosa, have been placed in Phymatosorus (Phymatodes), 1 cm (0.4 in.); c. sori on projection bent to upper frond
but some pteridologists put them in Myrmecopteris (Myrme- surface, bar = 2 mm; d. swollen rhizome, bar = 1 cm
cophila). More study is needed to resolve these differences. (0.4 in.).
For details, see Johns (1995). See Polypodium for a com-
parison of Lecanopteris to the other polypodiaceous genera.
346 C 13
a b
a
Figure 13.61.1. Lemmaphyllum microphyllum: habit.
Figure 13.61.2. Lemmaphyllum microphyllum: sori on Figure 13.61.4. Lemmaphyllum subrostatum: a. habit,
narrowed fertile fronds. bar = 5 cm (2 in.); b. fertile frond, bar = 5 cm (2 in.);
c. rhizome scale, bar = 1 mm.
348 C 13
Lepisorus
syn. Polypodium in part, Pleopeltis in part
The species of Lepisorus grow in soil, on rocks, or on tree
trunks and branches. These are small- to medium-sized, a b
creeping polypodiaceous ferns. The widespread Lepisorus
thunbergianus is often seen on trunks and branches in the
Hawaiian Islands, and many cultivars of this species are
commonly grown in pots or baskets in Japan. The culti-
Figure 13.62.1. Lepisorus longifolius: a. habit, bar = 5 cm
vated species can be difficult to establish, but with patience
(2 in.); b. fertile portion of frond, bar = 1 cm (0.4 in.);
the second or third attempt may succeed. Other species of
c. peltate scale of sori, bar = 0.1 mm.
Lepisorus are extremely difficult to grow.
The rhizomes vary from short- to long-creeping and
contain scattered, blackish fiber strands (seen in cross sec-
tion). Peltate, clathrate, entire or toothed scales cover the peltate scales. This species is native from India to Malaysia
rhizomes, sometimes with one or two glandular projections and the Philippines.
on the margins. The stipes are short, and the blades are sim-
ple, entire, and lanceolate to linear, sometimes bearing Lepisorus thunbergianus (Kaulfuss) Ching
clathrate scales. The fertile fronds are nearly the same size 13.62.2
and shape as the sterile ones. The veins are netted with syn. Polypodium lineare Thunberg
many irregular areoles, within which are free veins. The sori Hardy to very hardy
vary from round to elongate and are protected when young
A small fern with medium-creeping rhizomes. Grows
by partly or fully peltate, clathrate, entire scales. Like all
under medium to high light in well-drained, moist-dry
polypodiaceous ferns, Lepisorus lacks indusia. The species of
potting mix or uncut moss. This species is difficult to estab-
Lepisorus were first classified in Polypodium and later in
lish but is a popular pot plant in Japan, with many named
Pleopeltis; see Polypodium for a comparison of this genus to
cultivars.
other polypodiaceous genera.
The rhizome scales of Lepisorus thunbergianus are dark
Lepisorus is native mainly to the Old World tropics and
brown, ovate-cordate at base, and long-attenuate. The
subtropics. The number of species is unknown, but is prob-
fronds are simple, leathery, tapered at both ends, linear-
ably less than 40. The genus name comes from the Greek
elliptic, 10–30 cm (4–12 in.) long, and 5–15 mm (0.2–0.6
lepis, scale, and soros, mound, referring to the scaly sorus.
in.) wide. Because of the leathery blades, the veins are hard
to see. The sori are round or oval and disposed in one row
Lepisorus longifolius (Blume) Holttum on either side of midrib. The species is native to Japan,
13.62.1 South Korea, China, Taiwan, Indochina, the Philippines,
syn. Paragramma longifolia (Blume) T. Moore and the Hawaiian Islands.
Probably tender The species is extremely variable, not only in blade size
A medium-sized fern with short-creeping rhizomes. Grows and shape, but also in habitat. The plants grow on rocks,
well under medium light in moist, drained uncut moss or steep banks, tree trunks, stone walls, and the roofs of
potting mix. This species is reportedly the easiest Lepisorus houses.
to grow in Florida.
Lepisorus longifolius has linear-lanceolate fronds. Oval
to oblong, marginal sori are covered and surrounded by
F F A C 349
b
Leptochilus Kaulfman
syn. Colysis C. Presl
These small- to medium-sized ferns have variously shaped
leaves and are attractive in pots, baskets, or (in warmer cli-
mates) the ground. Leptochilus, better known to gardeners
as Colysis, resembles Microsorum but differs by having elon-
gate sori located in an even row between two veins. They
are epiphytic or epilithic.
The genus has creeping rhizomes covered by peltate or
false-peltate, clathrate or subclathrate scales. The fronds are
simple or pinnatifid, with the fertile fronds contracted or
not. The veins are visible, netted, and form areoles con- a
taining branched included veinlets pointing in various di-
rections. The sori are mostly elongate or linear (rarely acros-
tichoid) and usually appear in one line or a fairly even row
that is parallel to the secondary veins. The indusium is
absent.
Leptochilus consists of about 10 species and is native to
Asia, Malaysia, Australia, and the Solomon Islands. The
genus name comes from the Greek lepto, meaning slender,
and -chilus, meaning lipped, alluding to the blade tips of
some species, which end in a slender, curled tip. For a tech-
nical comparison of Leptochilus to related genera, see
Nooteboom (1997); also see Polypodium for a comparison Figure 13.63.1. Leptochilus amplus: a. habit, bar = 5 cm.
of this genus to other polypodiaceous genera. (2 in.); b. sori and vein pattern, bar = 1 cm (0.4 in.).
350 C 13
Leptopteris
The genus Leptopteris resembles large filmy ferns with strik-
ing delicate, plume-like fronds. The filmy texture and
translucent appearance are due to the blades being only a
few cell layers thick. These ferns are native to moist, humid
habitats, where they can form woody trunks to 1 m (3 ft.)
a tall. Leptopteris should be grown in protected sites with
b above-freezing temperatures and ample humidity. If these
conditions are unavailable, the plants should be grown in-
doors in large, shaded terrariums kept cool and humid year-
round. They may be planted in high-quality uncut sphag-
num moss.
Figure 13.63.2. Leptochilus ellipticus: a. habit, bar = 5 cm Although rarely cultivated in the United States, Lep-
(2 in.); b. sori and vein pattern, bar = 1 cm (0.4 in.). topteris is so striking that it merits further attempts. The
F F A C 351
genus is now on the endangered list in New Zealand, which is easier to grow than Leptopteris superba and tolerates
prohibits the export of mature plants. They can be grown slightly drier conditions.
from spores, but the spores are green and presumably have Leptopteris hymenophylloides has ovate to triangular blades
short viability. with flat segments (that is, in the same plane of the blade).
The rhizomes of Leptopteris are erect and trunk-forming. The species is endemic to New Zealand.
Medium- to large-sized fronds are arranged in a whorl and
have finely hairy rachises and costae. The blades may be up Leptopteris superba (Colenso) C. Presl 13.64.2
to three-pinnate, are very thin, and have ultimate divisions Crape fern, Prince-of-Wales plume
that are often bilobed. The sporangia are scattered on the Semi-tender to semi-hardy
lower surfaces of the blades. Indusia are lacking. A medium-sized fern with erect rhizomes and fronds that
Leptopteris, which contains six species, is native to Aus- collectively form a basket. Grows under low light and in
tralia, New Zealand, and New Guinea to Samoa and New moist-wet, humusy potting mix or uncut sphagnum moss.
Caledonia. The genus name comes from the Greek leptos, This species grows slowly and is difficult to cultivate, re-
thin, and pteris, fern, referring to the extremely thin fronds. quiring cool temperatures and constant high humidity. In
nature it grows near waterfalls.
Leptopteris hymenophylloides (A. Richard) C. Presl In contrast to the preceding species, Leptopteris superba
13.64.1 has blades that taper gradually toward the base, ending in
Semi-tender to semi-hardy small pinnae. Another difference is that the segments turn
A medium-sized fern with erect rhizomes and fronds that up, out of the plane of the blade. The species is endemic to
collectively form a basket. Prefers low light and moist-wet, New Zealand.
humusy potting mix or uncut sphagnum moss. The plants
require high humidity and cool temperatures. This species Leucostegia
Although commonly used as a pot plant in Malaysia, Leu-
costegia is rarely cultivated in the United States. The genus
is related to Davallia, which it resembles by its much-di-
vided, broad fronds.
The plants are usually terrestrial and medium in size.
Their rhizomes are thick, fleshy, more or less long-creeping,
and root-bearing on all sides. They are covered by non- This fern resembles Leucostegia immersa C. Presl, which
peltate scales, and hairs are often present as well. Grooves is not cultivated in the United States but is reported to be
appear on the upper surfaces of the stipes. The blades are cultivated in Australia. That species differs by indusia at-
nearly three-pinnate to four-pinnate-pinnatifid, glabrous tached only by a narrow base.
when mature, anadromous, and herbaceous or papery
(chartaceous). The ultimate segments are rhombic-lanceo- Lindsaea 13.66.1
late. The sori are often impressed and borne at the vein tips. These attractive ferns are often seen in the tropics and
They are covered by delicate, kidney-shaped or rounded sometimes in temperate areas, but they are difficult to grow
indusia—if the indusium is reniform, then it is attached at and rarely cultivated in the United States. They are prima-
a point at the base; if rounded, then attached halfway up rily seen in botanical gardens and special fern collections. In
the sides. England, four species of Lindsaea are currently cultivated.
Leucostegia, which consists of two species, occurs in Some species establish themselves if provided with a well-
southern China, the Himalayas, and southern India to drained planting mix and anchored in place on uncut moss
Polynesia. The genus name comes from the Greek leucos, or a turf-like growing medium. The plants should be kept
white, and stege, cover, referring to the whitish indusia. evenly moist in terrariums. The genus is mentioned here
because of its horticultural potential and importance in the
Leucostegia pallida (Mettenius) Copeland tropics.
13.65.1 Some of the species resemble maidenhair ferns (Adian-
Tender tum), but there are several differences. Lindsaea almost al-
A medium-sized fern with creeping rhizomes and stipes 2– ways has light-colored (tan, green, or yellowish) stipes,
3 cm (0.75–1.2 in.) apart. Grows well under medium to rachises, and costae, and these are grooved on their upper
high light in moist garden soil or potting mix. The plants surfaces, often with a sharp wing on either side of the
can be grown in baskets or pots. The fronds tend to spread groove. Furthermore, the indusium is formed on the lower
and should be given plenty of room.
Leucostegia pallida has triangular blades, up to four-pin-
nate, with obovate or rhombic ultimate segments. The sori
appear as raised bumps on the upper surface of the blade. b
Roundish indusia are attached at the base and halfway up
the sides. The species is native to the New Hebrides, Bor-
neo, and Malaya.
d
Figure 13.65.1.
a Leucostegia pallida:
a. habit, bar = 5 cm Figure 13.66.1. Lindsaea: a. frond (Lindsaea ensifolia);
(2 in.); b. segment b. habit (L. repanda); c. frond (L. stricta), bar for a,
with sori and indusia, b, and c = 5 cm (2 in.); d. sorus and indusium, bar =
bar = 2 mm. 2 mm.
F F A C 353
Llavea
Llavea consists of one species, endemic to Mexico and Gua-
temala, that is rarely cultivated. Llavea cordifolia has an at-
tractive airy appearance and bears many gray-green, oval
segments. The plants are easy to grow from spores. They are Figure 13.67.1. Llavea cordifolia: habit.
sensitive to overwatering and susceptible to leaf-spot dis-
eases when grown in extremely humid environments.
The genus has several distinctive characteristics, the the other, Lonchitis occidentalis Baker, to Africa and Mada-
most obvious being the strong differentiation in the shape gascar. The genus name comes from the Greek lonche,
and position of its sterile and fertile segments. The sterile lance, and -itis, a suffix indicating a close connection. It
segments are oval or nearly so and borne throughout the comes from an ancient name that was used by the Greek
basal two-thirds of the blade, whereas the fertile ones are physician and botanist Dioscorides.
linear and borne only in the apical third. The sori are pro-
tected by the enrolled margin of the segments. The rhizome
scales are mostly black but grade into bright yellow ones at
the bases of the leaf stalks. The genus is named for Pablo de
la Llave (1773–1833), a traveler in Mexico. b
Lonchitis
syn. Anisosorus
These terrestrial ferns are rarely cultivated. Lonchitis has c
rather coarsely cut fronds with a soft, hairy appearance. The
rhizome is creeping, thick, and fleshy and bears hairs and
scales. The stipe contains two vascular bundles. The fronds
are slightly succulent, two- to three-pinnate-pinnatifid, a
broad, and hairy. The veins are free or occasionally netted.
The sori are marginal, being borne at the sides of the seg-
ments or lobes. The indusium is formed by the enrolled
leaf margin.
This genus is sometimes confused with Blotiella (not
cultivated), which has hard, erect rhizomes and fully netted
veins. Lonchitis is distinguished from the similar Pteris by
hairy (not scaly) rhizomes. Figure 13.67.2. Llavea cordifolia: a. frond, bar = 5 cm
The genus consists of two species that hardly differ. (2 in.); b. distal portion of a fertile frond, bar = 5 cm
One, Lonchitis hirsuta, is native to tropical America, and (2 in.); c. stipe base, bar = 5 cm (2 in.).
354 C 13
a b
c
c
Figure 13.69.12. Lycopodium phlegmaria: a. branches Figure 13.69.13. Lycopodium phlegmarioides: a. branches
with sterile and fertile leaves in spikes, bar = 5 cm with sterile and fertile leaves in spikes, bar = 5 cm
(2 in.); b. sterile leaf bar = 3 mm (0.1 in.); c. fertile leaf, (2 in.); b. sterile leaf, bar = 3 mm (0.1 in.); c. fertile leaf,
bar = 3 mm (0.1 in.). bar = 3 mm (0.1 in.).
360 C 13
rangia are shorter than the fertile leaves. This species is na-
tive from Australia and Malaysia to New Caledonia and
the Pacific Islands.
Figure 13.69.15.
Lycopodium selago Linnaeus 13.69.14
b Lycopodium
syn. Huperzia selago (Linnaeus) C. Martius & Shrank
squarrosum:
Mountain club moss, fir club moss, alpine club moss
a. branches with
Very hardy, Zone 2
fertile and sterile
A small plant with evergreen, clump-forming stems. Grows leaves in spikes,
under medium light in moist-wet sandy soil or drained pot- bar = 5 cm
ting mix. This species is difficult to grow. (2 in.); b. sterile
a
The stems tend to be yellow-green (at least in cultiva- leaf, bar = 3 mm
tion), forked only once or twice, and produce buds near the (0.1 in.); c. fertile
tips. The sterile leaves are mostly narrowly triangular, leaf, bar = 3 mm
pressed close to the stem, and have papillate or entire mar- c
(0.1 in.).
gins. The fertile leaves resemble the sterile. Lycopodium se-
lago is similar to L. lucidulum but differs by having stomata
on both surfaces of the leaves, whereas in L. lucidulum these
are only on the lower surface. A hand lens is needed to see
the stomata, which appear as tiny white dots. Lycopodium se-
lago is native to northern North America (chiefly Canada).
Figure 13.70.7.
Lygodium
palmatum: leaflets,
sterile (above) and
fertile (below), bar
= 5 cm (2 in.).
Figure 13.73.7.
Marsilea mutica:
a. habit, bar =
2.5 cm (1 in.);
Figure 13.73.5. Marsilea macropoda: a. habit, bar = b. sporocarp, bar =
2.5 cm (1 in.); b. sporocarps, bar = 5 mm (0.2 in.). 5 mm (0.2 in.).
b
Marsilea minuta Linnaeus 13.73.1, 6 a
Tender to semi-tender
A very small fern with medium-creeping to erect rhizomes.
Grows well under high light in aquatic conditions or moist
to wet sand-peat mix or garden soil. Marsilea quadrifolia Linnaeus 13.73.8
Marsilea minuta is one of the smallest cultivated species, Hardy, Zone 5
growing to 7 cm (3 in.) tall. Some roots are present on the A small fern with medium-creeping to erect rhizomes.
internodes. The distal margin of the leaflet tends to be cre- Grows well under high light in aquatic conditions or moist-
nate. The sporocarp stalk branches near the base, bearing wet garden soil or sand-peat mix. The plants are used in
two or three (rarely four) sporocarps that are small, 2.6–4.1 China for treating infections.
mm long, with one conspicuous tooth. Marsilea minuta is Unlike many species of Marsilea, Marsilea quadrifolia
a common, widespread, and often weedy species in Africa has leaves glabrous or nearly so. Also, the stipes of the non-
and India. It occurs in a few scattered locations in the New immersed fronds tend to recline on the soil instead of being
World tropics, but these populations probably represent stiffly erect as in many other species. The branched sporo-
escapes from cultivation. carp stalks of M. quadrifolia are found elsewhere only in
M. macropoda, a species that differs by its hairy leaves.
Roots are present on nodes and internodes. The species is
native to the northeastern United States, southeastern Eu-
a rope, and Asia.
Figure 13.73.6.
b Marsilea minuta:
Marsilea schelpeana Launert 13.73.9
a. habit, bar =
Notched water fern
2.5 cm (1 in.);
Semi-tender to (perhaps) hardier
b. sporocarp, bar =
5 mm (0.2 in.). A small fern with medium-creeping to erect rhizomes.
Grows well under high light in aquatic conditions or moist-
wet garden soil or sand-peat mix.
F F A C 369
Figure 13.73.10.
Marsilea vestita: a.
habit, bar = 2.5 cm
(1 in.); b. sporocarps,
bar = 5 mm (0.2 in.).
b
Matteuccia 13.74.1, 2
syn. Pteretis, Struthiopteris
Ostrich fern
The ostrich fern native to the United States and Canada,
a Matteuccia struthiopteris, is a medium- to large-sized plant
Figure 13.73.9. Marsilea that produces an attractive vase-shaped cluster of fresh
schelpeana: a. habit, bar = green fronds in spring. It forms dense stands in damp places
2.5 cm (1 in.); b. sporocarp, as it sends out new plants from stolons. The common name
bar = 5 mm (0.2 in.). refers to the feathery, plume-like fronds that resemble os-
trich feathers.
370 C 13
Figure 13.74.2. Matteuccia struthiopteris var. Figure 13.74.3. Matteuccia orientalis: fertile frond (left)
pensylvanica: fertile pinnae. and sterile frond (right), bar = 10 cm (4 in.).
F F A C 371
Usually one or two new plants are produced each year from
the stolons.
The sterile leaves of Matteuccia struthiopteris are green
and oblanceolate, shaped like an ostrich feather, and grad-
ually taper toward the base, ending in highly reduced pin-
nae. The fertile leaves are brown, smaller than the sterile
ones, and have strongly ascending pinnae with enrolled
margins that protect the sporangia within. The species is
native to Europe, Asia, and North America.
var. pensylvanica (Willdenow) C. V. Morton. American
ostrich fern. Figures 13.74.1, 2, 4. Stipe scales about the
same color throughout (not darker in the center); pinna
lobes rounded. This is the common and widely planted va-
riety in North America.
var. struthiopteris (Figure 13.74.5). European ostrich
fern. Stipe scales bicolorous, darker in the center; pinna
lobes more truncate. This variety has been recently intro-
duced into cultivation in the United States.
Megalastrum
syn. Ctenitis section Subincisae (C. Christensen) Tindale
These medium- to large-sized ferns have decumbent to
erect rhizomes. The genus Megalastrum was previously clas-
sified in Ctenitis, but its basal basiscopic veins on the distal
pinnules spring from the costa, not the costule, and the lobe
supplied by this vein is widely adnate to the costa. Another
Figure 13.74.4. Matteuccia struthiopteris var. difference, best seen on the upper surface of the blade, is
pensylvanica: sterile frond (left) and fertile frond (right), that the veins end behind the margin in slightly enlarged,
bar = 10 cm (4 in.). club-shaped tips—that is, they do not run all the way to
372 C 13
a
c
a b
a b
b
Figure 13.76.7. Microgramma reptans: a. habit of a small
plant, bar = 5 cm (2 in.); b. sterile fronds (three on the
Figure 13.76.5. Microgramma percussa: a. habit, bar = left) and fertile frond (right), bar = 5 cm (2 in.), after
5 cm (2 in.); b. fertile portion of frond, bar = 1 cm Hoshizaki (1982); c. scales on frond, bar = 1 cm
(0.4 in.); c. blade scales, bar = 1 mm. (0.4 in.), after Hoshizaki (1982).
F F A C 375
Figure 13.77.5. Microlepia strigosa: a. frond, bar = 5 cm (2 in.); b. lower pinna, bar = 2 cm (0.8 in.); c. lower pinnae,
showing variations, bar = 2 cm (0.8 in.); d. forma macfaddeniae frond, bar = 5 cm (2 in.); e. forma macfaddeniae frond
detail, bar = 1 cm (0.4 in.).
F F A C 379
a a
b b
Figure 13.77.6. Microlepia substrigosa from European Figure 13.77.7. Microlepia substrigosa from Hawaiian
growers: a. frond, bar = 5 cm (2 in.); b. lower pinna, bar gardens: a. frond, bar = 5 cm (2 in.); b. lower pinna, bar
= 2 cm (0.8 in.). = 2 cm (0.8 in.).
have long, arching fronds, and these are attractive as hang- Most species of Microsorum have included veinlets
ing plants. pointing in all directions, not just toward the blade margin
Besides the cobra plant, the East Indian polypody (Mi- as in many other polypodiaceous ferns. The large sunken
crosorum scolopendria) is commonly cultivated. In tropical sori of Phymatosorus (Phymatodes) intergrades into Micro-
climates this species and its close relatives are used as sorum, and because other reliable distinguishing characters
ground covers, basket plants, or over rock walls. Although have not been found, Nooteboom (1997) reduced Phyma-
the branching rhizome is capable of growing to 6 m (20 ft.) tosorus to Microsorum. For more details, see Bosman (1991)
in length, cultivated plants are seldom allowed to grow this and Nooteboom (1997). Microsorum species were formerly
long. As with many ferns, the fronds may differ in shape listed under Polypodium and may be more familiar to gar-
on the same plant, ranging from entire to pinnatifid. deners under that name. See Hoshizaki (1982) for details
The genus consists of medium- to large-sized epiphytes on cultivated species. The entry for Polypodium in this
with long- or short-creeping rhizomes bearing peltate or chapter includes a comparison of Microsorum with other
false-peltate, clathrate or subclathrate scales. The blades are polypodiaceous genera.
either entire, lobed, or pinnatifid, or (in Microsorum lu- Microsorum consists of about 50 species native to tropi-
cidum) pinnate. The veins are netted (this is hard to see in cal and subtropical regions of Africa, southeastern Asia,
species with thick blades) and have within the areoles northern Australia, and the Pacific Islands. It has become
branched veinlets that may point in any direction. The sori naturalized in several areas in the New World. The genus
are separate, small to larger, round or less often elongate, name comes from the Greek mikros, small, and soros,
and irregularly scattered or in partly irregular rows, and mound, referring to the small, scattered sori, although some
they lack an indusium. species have larger sori.
380 C 13
a
b
overhead watering, which can cause leaf spots. The veins Microsorum pteropus (Blume) Copeland
form a handsome pattern. 13.78.11
The fronds of Microsorum musifolium are oblanceolate, syn. Colysis pteropus (Blume) Bosman
to 1 m (3 ft.) long, 10 cm (4 in.) wide, and sessile; the fronds Tender
are truncate to obtuse at the base and often narrowed A small fern with short-creeping rhizomes. Grows well
above. The venation is regular and conspicuous, with 6–10 under medium light in wet, coarse garden soil or potting
primary connective veins between the pairs of secondary mix with sand; can also be planted in drained pots or pots
veins. The species is native to southeastern Asia and the set in water. The plants can be grown submerged and are
Malayan Archipelago. often used in aquariums. They can remain submerged for
We have not followed Nooteboom (1997), who con- years without ill effects; however, sori are formed only on
cluded that this species was the same as Microsorum punc- emergent fronds, not those under water. The fronds are
tatum. Microsorum musifolium differs by its conspicuous vulnerable to leaf spots, which usually can be controlled by
veins and firm but thin translucent blades. In contrast, live fungicide sprays.
plants of M. punctatum have inconspicuous veins and leath- The fronds of Microsorum pteropus are simple to
ery blades. trilobed, with the center lobe the largest. Submerged fronds
are small and entire. The buds often form on the lower sur-
Microsorum papuanum (Baker) Parris face where sori would normally form. The species is native
13.78.10 from India to southern China, the Malay Archipelago, New
Tender, Zone 9 Guinea, and Japan. In nature it typically grows along stream
A medium-sized fern with creeping rhizomes bearing fronds banks prone to flooding, or in the spray of waterfalls.
close or more distant from each other. Grows under high to
medium light in well-drained, moist to moist-dry potting
mix.
Microsorum papuanum differs from the closely related
Microsorum scolopendria by its leathery and often crisped
blade that is simple and entire or has one to three lobes,
with the central lobe the longest. The sori are round to oval,
often irregularly so, and deeply sunken into the frond (em-
bossed on the upper surface). In cultivated plants, the sori
reach 13 mm (0.5 in.) in length and to 8 mm (0.3 in.) in Figure 13.78.11. Microsorum
width. It is native to Malaysia, New Guinea, and the Pacific pteropus: habit, bar = 5 cm
Islands. (2 in.), after Hoshizaki (1982).
a b
with some moisture-retaining pockets in which to root. green, leathery, and flat. The veins are hard to see. The sori
The fronds are evergreen and fragrant when crushed. vary from round or oblong and are usually arranged in two
The fronds of Microsorum scandens are entire or pinnat- irregular rows on each side of the midrib. The species is na-
ifid, herbaceous, narrowly elliptic, to about 60 cm (24 in.) tive to the tropics of the Old World.
long, 16 cm (6.5 in.) wide. The frond base is long-decur- As construed by Nooteboom (1997), Microsorum scolo-
rent on the stipe. The lobes of the blade are narrow, 4–10 pendria consists of a single variable species; however, other
mm (0.2–0.4 in.) wide, and separated by broad, U-shaped botanists believe it consists of several. Plants of M. scolopen-
sinuses. The sori are round or elongate, borne in one row dria with more than four pairs of lobes are recognized by
between costa and margin, and slightly raised on the upper some authors as M. grossum, whereas those with four or
surface of the frond. The species is native to Australia and fewer are called M. scolopendria (and these plants are by no
New Zealand. means uniform). Most plants cultivated in the United States
are M. grossum.
Microsorum scolopendria (N. L. Burman) Copeland
13.78.17 Microsorum steerei (Harrington) Ching
syn. Phymatosorus scolopendria (N. L. Burman) Pichi- 13.78.18
Sermolli, Polypodium scolopendria N. L. Burman Tender
East Indian polypody A small-medium fern with short-creeping rhizomes and
Tender thick and leathery fronds up to 40 cm (16 in.) long, though
A medium-large fern with long-creeping rhizomes. Grows usually shorter in cultivation. Prefers low to medium light
well under high light in drained, moist to moist-dry gar- and a well-drained, moist potting mix. Plants grow some-
den soil, potting mix, or uncut moss. what slowly and seem to do best at temperatures of 21°C
The long-creeping rhizome bears triangular or ovate- (70°F) or more year-round. This species is striking because
triangular scales that are spiny or ciliate on the margins. of its blue-green iridescent fronds.
The blade of Microsorum scolopendria is deeply lobed into The fronds of Microsorum steerei are like those of a small,
four or fewer pairs of segments that are somewhat yellowish fleshy M. punctatum but are more pointed and distinctly
a c
Figure 13.78.17.
Microsorum
scolopendria: a. habit,
bar = 5 cm (2 in.); b
b. sori, bar = 1 cm
(0.4 in.); c. rhizome
scale, bar = 1 mm.
388 C 13
Figure 13.78.19.
Microsorum superficiale:
a. habit, bar = 5 cm
(2 in.); b. sori, bar =
1 cm (0.4 in.).
culated among growers as “Mt. Elgan fern,” a name of un- long-pentagonal blades are one-pinnate-pinnatifid to two-
certain origin. pinnate, with linear, entire or serrate, adnate segments. The
Microsorum viellardii has rounded rhizome scales up to surfaces of the blades are smooth except for some fine hairs
6 mm (0.2 in.) wide. The blade is deeply lobed, and the on the upper surface. The veins are prominent and free.
lobes are narrowed toward the base. The sori are submar- The indusium is linear, distinctly submarginal, and up to
ginal and in one row between the midrib and margin. This 0.8 mm wide. The species occurs from Mexico to Central
species is endemic to New Caledonia. America and the Galápagos Islands.
a
c
b
are lax and hairy in general aspect. Plants volunteer readily
from spore, accounting for most of the plants in cultiva-
tion. Requires winter protection in southern California.
Nephrolepis biserrata can be separated from many others
in Nephrolepis by the hairy lower surfaces of the pinnae,
rachises, and costae. The stipe bases bear loose, spreading,
narrow, light brown, concolorous scales, and the pinnae
are typically 7 cm (3 in.) long or more with doubly crenate
margins (large crenations alternating with small ones). The
c
indusia are submarginal and round-reniform. The species is
native to tropical America, Africa, and southeastern Asia.
Trade plants by this name are actually Nephrolepis
falcata.
e
d
Figure 13.80.6. Nephrolepis cordifolia, fronds of cultivars: a. ‘Duffii’, pinnae bar = 1 cm (0.4 in.); b. ‘Kimberly Queen’;
c. ‘Lemon Buttons’, pinnae bar = 1 cm (0.4 in.); d. ‘Plumosa’; e. ‘Pom-pom’. All fronds, bar = 10 cm (4 in.).
America, Africa, Asia, Australia, Japan, and New Zealand ‘Lemon Buttons’. Figure 13.80.6c. Fronds to about 25
(where it reportedly lacks tubers). cm (10 in.) long, to 2.5 cm (1 in.) wide; pinnae reduced,
‘Duffii’. Figure 13.80.6a. Fronds less than 30 cm (12 oblong to roundish, not symmetrical at the base, not dou-
in.), erect, forked; pinnae small, to about 1 cm (0.5 in.) bled as in ‘Duffii’.
long, rounded or fan-shaped, often in pairs. This plant does ‘Petticoats’. Fronds to about 70 cm (28 in.) long, ends
not produced sori, and although considered a cultivar of forked several times in different planes; pinnae mostly ob-
Nephrolepis cordifolia, its parentage is doubtful, in part be- long-rectangular, sometimes shallowly and bluntly forked
cause it does not produce tubers and is more tender than N. at their tips. Forms with long fronds have been called ‘Den-
cordifolia. nis Petticoats’; those with short fronds, ‘Pom-pom’ (Fig-
‘Ecuadorian Fern’. Pinnae ruffled, undulate to irregu- ure 13.80.6e); and a shorter, more erect form, ‘Can-can’.
larly lobed, finely toothed. A recent introduction from cul- The weight of the terminal tassel, especially on long fronds,
tivation in Ecuador. causes the blades to bend and become pendent.
‘Kimberly Queen’. Figure 13.80.6b. Fronds to 1 m (3 ‘Tesselata’. Fronds to about 75 cm (30 in.) long; pin-
ft.) or more long and over 12.5 cm (5 in.) wide, sometimes nae one-pinnate in the distal half, except for lower pinnae;
slightly narrowly obovate; more arching and softer in ap- pinnules oblong-ovate. Variations in this cultivar include
pearance than the species. This cultivar grows vigorously. It pinna and pinnule length, the degree to which the pinnae
is often misidentified in the trade as Nephrolepis obliterata are bent backward, and the degree of emargination.
(R. Brown) Carruthers, an Australian fern with narrow, Whether these variations are due to cultural conditions or
distantly spaced pinnae, not known in cultivation. are stable is not certain. Related (if not the same) plants are
F F A C 393
circulating as ‘Plumosa’ (Figure 13.80.6d) and ‘Mildred round to round–kidney-shaped with a U-shaped sinus,
Murray’. mostly opening toward the margin. This species is native to
Florida, the West Indies, and the Pacific Islands.
Nephrolepis exaltata (Linnaeus) Schott Young plants of Nephrolepis exaltata are difficult to dis-
13.80.2, 7, 8; 5 tinguish from those of N. cordifolia (which are often mis-
Common sword fern, Boston fern takenly sold as N. exaltata). In N. exaltata the scales on the
Tender to semi-tender upper surfaces of the rachises are uniform in color or, if
A medium-large fern with short, erect, stoloniferous rhi- darker in the center, then also with irregular darker spots on
zomes. Grows best under medium to high light in moist- the rays of the scales. On young plants of N. cordifolia, these
dry potting mix. The wild form is stiff, upright, and has a scales are usually whitish with a dark center sharply marked.
rank appearance. Although the wild form is rarely culti- The fronds are also narrower than those of N. exaltata. Ster-
vated, the cultivars derived from it are among the most ile, adult plants of N. cordifolia are distinguished by their
loved and widely grown ferns. Natural hybrids between erect, stiff, narrower fronds, crowded pinnae, and tuber-
Nephrolepis exaltata and other species may be cultivated, producing runners. For commercial purposes N. cordifolia
especially a hybrid with N. multiflora in Hawaii. More in- is grown mainly from spores, whereas the sterile Boston
formation on cultivating Boston ferns is provided in Chap- ferns are cloned mainly by tissue culture.
ter 10. Nephrolepis exaltata has given rise to numerous culti-
The stipe bases of Nephrolepis exaltata have spreading, vars, and these generally have the same cultural require-
completely brown (concolorous) scales. The pinnae are ments as the wild form. So commonly grown are these ferns
usually less than 5 cm (2 in.) long, with the surfaces essen- that commercial growers have established the precise cul-
tially glabrous or with a few hair-like scales on the upper tural conditions for optimum growth: 2000 foot-candles
surface. The rachises are sparsely to densely covered with of light, temperatures between 18 and 35°C (65–95°F),
hair-like scales on the upper surfaces, and the indusia are fertilizers with a 3-1-2 composition, micronutrients, and
dolomite to adjust the pH to 5.5. Use a mix with good
drainage, and water when the soil surface is slightly dry. To
prevent breakage and decay of fronds, water should be
applied at the soil level, not on the foliage. Poor foliage re-
sults when temperatures are below 18°C (65°F) for more
than a month. Repotting just before the active growth in
spring is recommended.
Those ferns with more finely cut or congested fronds
might grow more slowly and need more care. You can ex-
pect cultivars with dense, compact foliage to show aborted
pinnae where light does not penetrate into the center of the
cluster. This does not harm the plant as long as the outer fo-
liage is healthy. All declining fronds or pinnae, however,
should be removed to promote air circulation and light
a penetration to underlying foliage, discouraging decay. The
cultivars with long fronds are seldom found in the trade be-
cause these large ferns are not well accommodated to
smaller homes, and nursery growers find it difficult to trans-
port long fronds without damage.
b English growers report that buds form on the foliage of
their cultivars, but in the United States this has been doc-
umented only for ‘Trevillian’. The buds are reported to
produce plants that revert to the ordinary Boston fern,
‘Bostoniensis’.
c The names of Boston fern cultivars are greatly confused.
Both old and new names have been loosely used and are
poorly documented, except for the work of Sessions (1978).
The delineation of cultivars is further complicated by their
tendency to intergrade. Sometimes a given cultivar will pro-
Figure 13.80.7. Nephrolepis exaltata: a. frond, bar = 10 duce fronds with different characteristics on the same plant.
cm (4 in.); b. middle part of frond, bar = 2.5 cm (1 in.); Cultivar variations include size and shape of divisions,
c. pinna, bar = 1 cm (0.4 in.). number of times pinnate, margin patterns, and degree of
g
b c
a
h i
d
e f
m n
Figure 13.80.8. Nephrolepis exaltata cultivars: a. ‘Bostoniensis’, middle part of frond; b. ‘Dallas’; c. ‘Elsevier’ (frond
enlarged at right); d. ‘Fluffy Ruffles’; e. ‘Gretnae’, middle part of frond; f. ‘Massii’; g. ‘Mini-ruffles’ (frond enlarged at
right); h. ‘Norwoodii’, showing pinna; i. ‘Rooseveltii’, upper half of frond; j. ‘Sassy’; k. ‘Smithii’; l. ‘Splendida’; m.
‘Verona’, showing pinna; n. ‘Wanamaka’. Bar = 10 cm (4 in.).
F F A C 395
ruffling, twisting, forking, or overlapping. Some of these times forked and heavily crested. The pinna apex is acute or
variants tolerate cold better, tend to revert less, and retain to three times forked. A large spectacular fern when well
their pinnae longer. grown.
Only a small number of the hundreds of cultivars of ‘Verona’. Figure 13.80.8m. Fronds three-pinnate, to
Nephrolepis exaltata are mentioned here. They are selected about 40 cm (16 in.) long and half as wide, spreading to
to show the great diversity within the group. pendent, the segments ovate. All divisions are distant from
‘Bostoniensis’ (var. bostoniensis Desvaux). Boston fern. each other, hence the blade appears open and imparts an
Figure 13.80.8a. Fronds one-pinnate, up to about 120 cm airy appearance to the plant.
(4 ft.) long, narrow, arching; pinnae slightly wavy; sori (if ‘Wanamaka’. Figure 13.80.8n. Fronds one-pinnate,
present) with aborted sporangia. ‘Bostoniensis’ is the parent dwarf, less than 30 cm (12 in.) long, narrow, stiff, upright,
of the many Nephrolepis exaltata cultivars. It is more com- leathery, dark green; pinnae closely overlapping, curled and
pact and graceful than the wild plant, with the pinnae twisted, crenulate.
broader and not as flat as those of the wild form. It was first
noticed in a Boston nursery in 1821, hence the name Bos- Nephrolepis falcata (Cavanilles) C. Christensen
ton fern. 13.80.9
‘Dallas’. Dallas fern. Figure 13.80.8b. A compact plant Broad sword fern, macho fern
with short fronds bearing oblong pinnae that are notice- Tender
ably serrate with round apices.
A large fern with erect, stoloniferous rhizomes and erect or
‘Elsevier’. Figure 13.80.8c. Fronds two-pinnate, minia-
pendulous, long fronds. Grows well under medium to high
ture, less than 30 cm (12 in.) long. Resembles ‘Mini-ruffles’
light in moist-dry potting mix or uncut moss.
but with narrowly elliptic, slightly twisted fronds.
The older stipe bases on Nephrolepis falcata have scales
‘Fluffy Ruffles’. Figure 13.80.8d. Fronds two-pinnate,
that are spreading to appressed, lanceolate or triangular,
dwarf, less than 30 cm (12 in.) long, elliptic, stiff, upright;
and nearly black with a pale margin. The pinnae are up to
stipes thick; pinnae and pinnules overlapping, with entire
about 10 cm (4 in.) long and often curved toward the apex
to crenate margins. Tends to stay greener in winter.
(falcate). The fertile pinnae are slightly narrowed compared
‘Gretnae’. Figure 13.80.8e. Like ‘Bostoniensis’ but with
to the sterile ones. A few scattered scales are found on both
stouter pinnae forked at the apex into blunt lobes, the frond
surfaces of the blades, but these tend to fall off with age.
apex often several times forked.
The upper surface of the pinna midrib is glabrous or with a
‘Golden Boston’ (‘Aurea’). Like ‘Bostoniensis’ but with
few scales. The sori are near the margin, and the indusia
yellow-green fronds.
are round with a narrow sinus. The species is native to Ma-
‘Gracillima’ (‘Irish Lace’). Resembles ‘Smithii’, but the
laysia and extends northward into Burma and Indochina.
fronds are three-pinnate and more ovate, and the pinnae
Nephrolepis falcata is often misidentified as N. biserrata
arise uniformly, one behind the other.
and N. ensifolia C. Presl.
‘Massii’. Figure 13.80.8f. Short frond form with pinnae
‘Furcans’. Fishtail sword fern. Figure 13.80.9b. Fronds
slightly twisted at apex.
shorter and less pendent than typical for the species; pinna
‘Mini-ruffles’. Figure 13.80.8g. Fronds miniature, to
tips once- or twice-forked. In the tropics this plant is com-
about 10 cm (4 in.) long, three-pinnate, broadly triangular,
monly grown in pots in full sun, although it produces bet-
and erect, with overlapping pinnae.
ter color in the shade.
‘Norwoodii’. Figure 13.80.8h. Fronds three-pinnate,
‘Ram’s Horn’. Figure 13.80.9c. Pinnae twisted and
less than 40 cm (16 in.) long, ovate, spreading; pinnae and
curled backward.
pinnules more or less in regular layers, overlapping and
congested, pinnules divided into long narrow segments.
The dense foliage on the blade has a fluffy but layered ap- Nephrolepis hirsutula (G. Forster) C. Presl
pearance with delicate details. The two-pinnate form also 13.80.10
circulating as ‘Norwoodii’ is closer to ‘Fluffy Ruffles’. Rough sword fern, scurfy sword fern
‘Rooseveltii’. Figure 13.80.8i. Fronds to 1 m (3 ft.) long, Tender
pinnae wavy, usually eared on both sides of the base. A medium-large fern with short, erect, stoloniferous rhi-
‘Sassy’. Figure 13.80.8j. Fronds short; pinnae moder- zomes. Grows well under medium to high light in garden
ately twisted, ruffled, and irregularly finely dentate-serrate. soil or potting mix kept moist-dry.
‘Smithii’. Lace fern. Figure 13.80.8k. Fronds to four- The older stipe bases of Nephrolepis hirsutula bear ap-
pinnate, dwarf, about 30 cm (12 in.) long, widely ovate, pressed, convex scales with nearly black centers and pale
arching to pendent, lacy; segments ovate, small, 2 mm or margins. The pinnae are about 9 cm (3.5 in.) or more long,
less long, spaced apart. closely placed, and usually sharply eared. They bear fila-
‘Splendida’. Figure 13.80.8l. Fronds one- to two-pin- mentous scales on the lower surface. The upper surface of
nate, pendent, to 1 m (3 ft.) long, with the distal half many the pinna midrib has long hairy scales or is glabrous. The
396 C 13
a
b c
c
a
about twice as long as broad and have rounded apices and
mostly entire margins. The upper part of stipe and rachis
(except at the point of pinna attachment) essentially lacks
hairs or scales. The species is native to tropical America.
Niphidium
syn. Pessopteris, Polypodium in part
These epiphytic, epilithic, or terrestrial plants have me-
dium- to large-sized, stiff, undivided, upright fronds. They
are reminiscent of the bird’s-nest fern (Asplenium nidus),
but instead of the latter’s neat rosette of light green fronds,
Niphidium fronds are dark green and irregularly clustered. b
The two genera can be distinguished by the sori, which are
linear and indusiate in Asplenium but round and non-
indusiate in Niphidium. Niphidium also resembles the trop-
ical American genus Campyloneurum, but that genus has
two or more rows of sori between the main lateral veins, c
whereas Niphidium only has one. d
The rhizomes of Niphidium vary from short- to long-
creeping. The fronds are simple, entire, elliptic-lanceolate
to oblong, and leathery, often with white dots on the upper
surfaces. The main lateral veins are prominent and oblique Figure 13.81.1. Niphidium: a. N. crassifolium frond, bar
to the rachis. In the cultivated species the sori are large, = 5 cm (2 in.); b. N. crassifolium sori, bar = 2.5 cm
about 3 mm (0.1 in.) wide, usually round, and produced in (1 in.), after Hoshizaki (1982); c. N. crassifolium
a single row between the lateral veins. Each sorus is served rhizome scale, bar = 1 mm, after Lellinger (1972); d. N.
by a small vein ring. As in all polypodiaceous ferns, indusia albopunctatissimum rhizome scale, bar = 1 mm, after
are lacking. Lellinger (1972).
F F A C 399
Notholaena
Cloak fern
A mostly rock-inhabiting fern of dry climates, Notholaena
is difficult to grow and is found only in special collections.
These small ferns have short-creeping, compact rhizomes.
The genus resembles certain species of Cheilanthes and Pel-
laea but differs by the dense covering of white, cream, or Figure 13.82.1. Notholaena standleyi: fronds, the
yellow waxy powder (farina) on the undersides of the underside (left) with dense powder, and the upperside
fronds and by the marginal, hardly enrolled, narrow indu- (right), bar = 2.5 cm (1 in.).
sia. The stipes are frequently longer than the blades, and
the fronds may be linear-lanceolate to pentagonal and pin-
nate-pinnatifid to four-pinnate. Many species formerly
classified in Notholaena have been transferred to Cheilanthes Odontosoria
(which see for a comparison). syn. Sphenomeris
Notholaena consists of about 25 species native to North
The cultivated species of Odontosoria are finely cut terres-
America, Central and South America, and the West Indies.
trial plants of rocky areas. They are used in tropical gardens
The genus name comes from the Greek nothos, false, and
and are rarely seen elsewhere. Like many ferns that grow
chlaena, cloak, referring to the blade margins, which are
between rocks, these plants are difficult to remove without
not reflexed as in the similar genus Cheilanthes. For addi-
damaging the roots. Transplants with damaged roots often
tional information on growing cloak ferns, see the section
do not reroot readily, hence the genus is best propagated by
on “Xerophytic Ferns” in Chapter 10.
spores.
The short-creeping to suberect rhizomes are covered
Notholaena standleyi Maxon 13.82.1 with narrow or hair-like dark scales. The fronds are set close
Hardy together and have two- to four-pinnate blades. The ulti-
A small fern with compact rhizomes and fronds in a cluster. mate segments are linear or wedge-shaped and dichoto-
Requires high light in moist-dry, well-drained garden soil mously lobed, with a single vein or the vein once or twice
preferably mixed with coarse sand or gravel. forked. Marginal sori are borne at the vein tips or, more
Notholaena standleyi has pentagonal blades densely cov- rarely, are supplied by two or three veins. The rounded to
ered with a whitish powder on the lower surface, and the oval indusium is attached by the base and more or less by
hardly enrolled indusium is narrow. It might be confused the sides. It opens toward the margin of the segments.
with the more frequently grown Cheilanthes argentea, The genus, which consists of 22 species, is native to the
which is less divided, less densely powdery, and has a more tropics and subtropics of America, Africa, and Japan. Its
enrolled, broader indusium. Notholaena standleyi is native name comes from the Greek odous, tooth, and soros, mound,
to the southwestern United States and Mexico. and refers to the sori borne at the extreme tips of small,
marginal teeth.
400 C 13
c
a
Olfersia
Rarely cultivated, Olfersia grows in soil in the wild or
scrambles over fallen logs or rocks. It occasionally climbs
tree trunks, in which instances the rhizomes become long-
creeping. Younger plants tend to maintain a short rhizome
for some time and therefore can be confined to a wide pot.
Olfersia is medium-sized and can be distinguished from
all other ferns by its creeping, densely scaly rhizomes and
b strongly dimorphic sterile and fertile fronds. The sterile
fronds are one-pinnate with a terminal segment resembling
the lateral pinnae. The veins are long-parallel and free for
most of their length, joining just before the margin to form
a submarginal connecting strand. The fertile fronds are bi-
402 C 13
pinnate with linear pinnae and sori that merge. The indu- Onoclea can be identified by its netted veins, triangular
sium is absent. and deeply pinnatifid or (more often) one-pinnate-pinnat-
This genus consists of one species widespread in the ifid sterile blades, and strongly dimorphic sterile and fer-
American tropics. Its name honors Ignaz Franz Werner von tile fronds. Other characteristics are medium-wide-creep-
Olfers (1793–1871), a professor who collected in Brazil. ing rhizomes and fertile pinnae that are strongly ascending
See Moran (1986) for a technical treatment of the species. and contracted into rounded, bead-like lobes enclosing the
sori.
Olfersia cervina (Linnaeus) Kunze 13.85.1 The genus, which consists of only one species, is native
syn. Polybotrya cervina (Linnaeus) Kaulfuss to North America and Asia. The genus name comes from
Tender the Greek onos, vessel, and kleiein, to close, referring to the
A medium-sized fern with medium- to long-creeping rhi- pinnules of the fertile leaf, which roll up into bead-like seg-
zomes. Grows well under medium light in moist garden ments to enclose the sori.
soil. Olfersia cervina is easy to grow from spores. See the de-
scription of the genus for distribution and additional char- Onoclea sensibilis Linnaeus 13.86.1–3
acteristics. Very hardy, Zone 2
A medium-sized fern with medium- to long-creeping rhi-
zomes and deciduous fronds. Grows well under medium
to high light in moist-wet garden soil. This species is easy to
grow.
Onoclea sensibilis is characterized by pinnatisect fronds
with lobed pinnae narrowed toward the base, and veins
netted throughout. The margins of the blade are finely
serrulate. The fertile fronds are brown, erect, and rigid
with strongly ascending pinnae. The species is native to
eastern North America and eastern Asia, often growing in
wet habitats.
Onoclea sensibilis has been around for a long time. Fos-
sils identical to modern plants have been found in Paleo-
cene rocks in Canada—rocks that date back more than 54
million years (Rothwell and Stockey 1991).
Onychium
Claw fern
Onychium is a genus of terrestrial, clump-forming ferns that
can be planted in the ground or in pots. Their finely di-
vided fronds have a delicate, lacy appearance.
The rhizomes are short- to long-creeping and scaly. The
Figure 13.85.1. Olfersia cervina: sterile frond (left) and fronds are dimorphic or monomorphic. The sterile blades
fertile frond (right), bar = 5 cm (2 in.).
Onoclea
Sensitive fern, bead fern
The bead-like appearance of the fertile fronds accounts for
this genus’s common name of bead fern. Some say that the
name sensitive fern originates from the fronds’ sensitivity to
frost (they wither after the first subfreezing temperatures).
These medium- to large-sized natives of temperate areas
have coarsely cut but attractive foliage. The plants spread by
creeping, branching rhizomes. Onoclea is useful in wet,
temperate areas but will grow well in warmer climates even
though it is deciduous. The fertile fronds are often used in
dried flower arrangements. Figure 13.86.1. Onoclea sensibilis: habit.
F F A C 403
a c
a
b
Figure 13.89.5.
Osmunda claytoniana:
frond, middle pinnae
fertile, bar = 5 cm
(2 in.).
ing prior to withering and appear bunched together. The var. regalis. Figure 13.89.7. Fronds are completely de-
plants seem adapted to neutral or only slightly acidic soil. ciduous. The fronds are more leathery than those of the
The cultivated material of this fern was collected in Brazil. other varieties and have thicker stipes and rachises. The
Some botanists do not consider this variety distinct from pinnae and pinnules are sessile (or nearly so) and closer to-
var. spectabilis. gether than in var. spectabilis. The hairs on the rachis are
ample and darker than those of var. spectabilis. The fertile
pinnae spread at nearly right angles to the rachis and are
borne in the same plane as the sterile pinnae. This robust
grower is native to Europe and Asia. The following cultivars
are derived from it: ‘Cristata’, pinnules or segments crested
at the tips; ‘Purpurescens’, with purplish stipes, rachises,
and costae.
var. spectabilis (Willdenow) A. Gray. Fronds are com-
pletely deciduous. The pinnae and pinnules are stalked and
farther apart than in var. regalis. Also, the hairs on the
rachises are fewer and lighter colored than in var. regalis.
The fertile pinnae are slightly falcate and ascending, spread-
ing in the same plane as the sterile pinnae. This variety is
native to North America and might not be distinct from
var. brasiliensis.
Paesia
Plants of the genus Paesia typically have finely divided
fronds and zigzag rachises that allow them to scramble over
other plants. Paesia scaberula, however, the only cultivated
Figure 13.89.7. Osmunda
species, has only a slightly a zigzag rachis and less elongate
regalis: fertile frond, bar =
fronds.
5 cm (2 in.).
Paesia has slender, 3–5 mm (0.1–0.2 in.) long, creep-
ing, hairy rhizomes. Other characteristics of the genus are
two- to four-pinnate blades, free veins, and rachises and
costae that are strongly ridged and grooved on the upper
surfaces. The sori of Paesia have two indusia, an outer one
Figure 13.89.8. Osmunda formed from the modified, enrolled margin of the blade,
regalis var. brasiliensis: fertile and an inner one that is thin, colorless, and hidden by the
frond, bar = 5 cm (2 in.). outer one and, typically, by the sporangia. The sori are
borne on an outer, marginal vein supplied by two or more
lateral veins. In contrast, Hypolepis and Odontosoria (also
in the family Dennstaedtiaceae) have only one indusium,
and the sori are supplied by a single vein. Paesia is closely re-
lated to Pteridium but differs by having only one vascular
bundle in the stipe, whereas Pteridium has several arranged
in a horseshoe pattern.
Paesia consists of 12 species distributed in the Ameri-
can tropics, eastern Asia, and the western Pacific. It is
named for Duke Fernando Diaz Paes Leme, who visited
Brazil in 1660 on government service from Portugal.
a
c
faces, and the pinnae at the base of the blades are reduced
abruptly to small, downward-pointing lobes. The veins are
once-forked. The species is widespread in the American
tropics.
Pellaea andromedifolia is characterized by bicolorous rhi- narrower than the sterile ones. This species occurs from
zome scales, tan stipes, up to three-pinnate blades, and oval, Canada to Guatemala.
noncordate segments. The fiddleheads are densely scaly. In the wild this species grows on calcareous rocks such as
This species is native to the United States (California and dolomite and limestone. It resembles Pellaea glabella, with
Oregon) and Mexico (Baja California). which it often grows, but can be easily distinguished by the
hairy stipe and stalked pinnae and pinnules.
Pellaea atropurpurea (Linnaeus) Link 13.92.4
Purple cliff brake Pellaea brachyptera (T. Moore) Baker 13.92.5
Very hardy, Zone 4 Sierra cliff brake
A small to medium fern with stout, ascending rhizomes and Hardy, Zone 7
evergreen, clustered fronds. Prefers medium to high light A small fern with stout, ascending rhizomes and clustered
and moist-dry, basic garden soil with coarse sand. This spe- evergreen fronds. Grows under medium light in drained,
cies can tolerate annual minimum temperatures of −28 to moist-dry garden soil with coarse sand. The plants are dif-
−35°C (−23 to −31°F). ficult to cultivate, but they have been recorded to endure
Pellaea atropurpurea has rusty-tan and matted rhizome temperatures as low as −14°C (7°F).
scales. The stipe and rachis vary from purplish to black and Pellaea brachyptera has bicolorous rhizome scales, and
have a scruf consisting of minute, twisted hairs. The blade the stipes are dark brown and shiny. The blades are linear-
is bluish gray and two-pinnate, with the sterile segments oblong, two- to three-pinnate, and bluish green. The basal
oval to ovate-lanceolate. The fertile segments are longer and pinnae are one-pinnate, with the largest ones divided into
3–11 segments. The linear, tufted segments are crowded
on a short costa. The segment margins are greenish and
strongly enrolled, concealing nearly the entire lower sur-
Figure 13.92.3. Pellaea face of the pinnule. This species is native to the western
andromedifolia: frond, United States (California, Washington, and Oregon).
bar = 5 cm (2 in.).
Pellaea bridgesii Hooker 13.92.6
Bridges’ cliff brake
Hardy, Zone (5)6
A small fern with stout, ascending rhizomes and clustered,
deciduous fronds. Grows under high light in moist-dry,
drained garden soil with coarse sand. This species is difficult
to grow.
Some of the rhizome scales of Pellaea bridgesii are weakly Pellaea cordifolia is one of the few species in the genus
bicolorous. The stipes and rachises are dark brown and that has tan stipes and rachises. The blades are ovate-trian-
shiny. The linear blades are one-pinnate throughout, with gular and up to three-pinnate, with rachises and costae
the pinnae round to ovate, entire, and leathery. The fertile straight, not zigzag. The segments vary from round-cor-
pinnae are often folded lengthwise along the midvein. Spo- date to triangular-cordate, and the segment stalks are glab-
rangia are borne in a marginal band, but the margins re- rous. Hairs occasionally occur on the enrolled margin of
main flat, not enrolled as in many other species; P. bridge- indusium. This species is native to the southern United
sii therefore lacks a false indusium. The species is native to States (Texas) and Mexico.
the western United States.
Pellaea falcata (R. Brown) Fée 13.92.1, 8
Pellaea cordifolia (Sessé & Mociño) A. R. Smith syn. Platyloma falcata (R. Brown) J. Smith
13.92.7 Australian cliff brake
syn. Pellaea cordata (Cavanilles) J. Smith, P. sagittata var. Semi-hardy, Zone (7)8
cordata (Cavanilles) A. F. Tryon A small-medium fern with dark green fronds in loose clus-
Semi-hardy, Zone 8 ters from short- to medium-creeping rhizomes. Grows well
A medium-sized fern with short-creeping rhizomes and lax, under medium to high light in drained, moist-dry garden
scrambling, bluish gray-green fronds. Grows well under or potting mix. This species can withstand temperatures
high light in drained, moist-dry garden soil with coarse down to −7°C (19°F) for one week.
sand. The longer fronds tend to become tangled and will The stipes and rachises of Pellaea falcata bear spreading
need support if grown upright. scales, and the blades are one-pinnate. The pinnae, which
b a
Figure 13.92.7. Pellaea cordifolia: a. frond, older, bar = Figure 13.92.8. Pellaea falcata fronds: a. typical form;
5 cm (2 in.); b. frond, younger; bar = 5 cm (2 in.). After b. var. nana; c. ‘Star Glow’, older (left) and younger
A. F. Tryon (1957). (right). Bar = 5 cm (2 in.).
F F A C 415
currently consider these genera distantly related within the Pentagramma pallida (Weatherby) Yatskievych, Wind-
Pteridaceae and best placed in different subgroups of the ham & E. Wollenweber differs by having dull and pow-
family. dery stipes and blades that are white-powdery on both sur-
Pentagramma contains two species and is native to the faces, appearing grayish when fresh. It is endemic to the
southwestern United States and northwestern Mexico. The foothills of the Sierra Nevada in California and may be cul-
genus name comes from the Greek penta, five, and gramme, tivated by local hobbyists. Its cultural requirements are the
line, alluding to the pentagonal outline of the blade. same as for P. triangularis.
year from short runners. The green of the fronds lasts until
the first hard frost.
The rhizomes of Phegopteris decursive-pinnata are erect
or suberect, and the blades are narrowly elliptic and tapered
toward both ends, rather than broadly triangular as in the
other two species. The pinnae are hairy on both surfaces,
and their bases are connected to the rachis by a broadly
lobed wing, but the lower pinnae are often free. The rachis
is also hairy and bears ciliate scales. The species is native to b
eastern Asia.
Phlebodium
syn. Polypodium in part
Rabbit’s-foot fern, golden polypody
Phlebodium aureum is well known in the trade as a pot or
basket plant, though it can also be grown in soil with good
drainage. Its fronds and those of P. pseudoaureum vary con-
siderably, from gray-green to blue-green shades, from stiff
and erect to arching or pendant, from medium to large in
b size, from entire to fringed lobes, from sparsely to heavily
crested tips, and so on. These characteristics are affected by
the amount of nutrients and exposure to light. Phlebodium
a aureum is less cold hardy and needs more protection than P.
pseudoaureum, which is also a robust grower. For more de-
tails on Phlebodium cultivars, see Hoshizaki (1982).
Phlebodium is closely related to Polypodium, and some
Figure 13.94.2. Phegopteris decursive-pinnata: a. habit, pteridologists prefer to classify it in that genus. Like Poly-
bar = 5 cm (2 in.); b. winged rachis, bar = 1 cm (0.4 in.). podium, it has creeping rhizomes with two ranks of fronds
420 C 13
Figure 13.95.4.
Phlebodium
×schneideri: frond,
bar = 10 cm (4 in.).
Phyllitis
Hart’s-tongue fern
The hart’s-tongue ferns are small, terrestrial, evergreen
plants with tongue-shaped fronds. The fiddleheads of Phyl-
litis were used as the model for the scrolls at the ends of vio-
lins. There is wide variation in spore-grown plants, and
many garden forms are seen in cultivation. Phyllitis may
also be propagated from basal pieces of the stipe, a process
described in Chapter 8. The genus is susceptible to root-
rot, so avoid overwatering and provide good drainage. For
details about cultivars, see Druery (1912), Kaye (1968),
and Dyce (1972).
The distinguishing characteristics of the genus are its
simple, entire blades and paired sori in which each sorus in
the pair opens toward its partner. Additional traits include
the heart-shaped blade bases, leathery texture, once- or
twice-forked veins, and linear indusiate sori. Phyllitis is
closely related to Asplenium and hybridizes with it. Some
pteridologists prefer to classify Phyllitis under Asplenium;
it is only a matter of opinion whether to do so. Phyllitis is
used here because of its long use by horticulturists.
The genus contains five species and is native to North
America, Europe, and Asia. The name comes from the
Greek phyllos, meaning leaf, which was an ancient Greek
name for the fern.
Pilularia
Figure 13.96.3. Phyllitis scolopendrium: sori Pillwort
and indusia. The pillworts are small, inconspicuous, sedge-like or grass-
like plants. They can be distinguished from grasses and
sedges by the coiled tips of their young leaves. Pilularia is
Phyllitis scolopendrium has tongue-shaped fronds with related to the water clover (Marsilea) but lacks the distinc-
cordate bases. The stipes are short and scaly. The species is tive clover-like leaves. Of little ornamental value, this genus
native to North America and Europe. is best used as part of a small, aquatic dish-garden or in bog
Only the European variety (var. scolopendrium) is culti- or marsh plantings.
vated. The smaller, American variety (var. americana Fer- The rhizome is slender and creeping, bearing a filamen-
nald) is rare and difficult to grow. It tends to bear sori in the tous stipe at each node. The fertile part, which is not al-
distal half of the frond but is otherwise similar to the Euro- ways present, is a stalked, bean-like structure (sporocarp)
pean variety. borne at the base of the stipe. The sporocarp encloses the
The hart’s-tongue fern is extremely variable. In the late sporangia, which contain either male or female spores.
1800s, one British fern grower listed 445 varieties (Lowe Pilularia is cosmopolitan and consists of five species.
The genus name comes from the Greek pilula, little ball,
and refers to the globose sporocarps.
e Figure 13.97.1.
Pilularia americana:
habit, bar = 1 cm
(0.4 in.).
Figure 13.98.5.
Pityrogramma
calomelanos: Figure 13.98.6. Pityrogramma
a
underside of frond, ebenea: a. frond, bar = 5 cm
bar = 5 cm (2 in.). (2 in.); b. underside of
pinnule, bar = 2.5 cm (1 in.).
P. quadridichotomum
stipe
Base frond
cross-
P. wallichii of Malayan-
section
Asiatic species
P. andinum
except
P. ridleyi
P. grande
P. superbum
P. stemaria
P. elephantotis
P. wandae
P. madagas-
cariense
P. ellisii
P. holttumii
P. alcicorne
P. coronarium
Javan-Australian species
P. veitchii
P. ridleyi
a d
Figure 13.99.5. Platycerium andinum: a. habit, bar = Figure 13.99.6. Platycerium bifurcatum: a. habits, bar =
10 cm (4 in.); b. sporangial area placement; c. rhizome 10 cm (4 in.); b. sporangial area placement; c. rhizome
scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar = scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar =
1 cm (0.4 in.). 1 cm (0.4 in.); e. stipe bases, bar = 2 cm (0.8 in.).
The base fronds of Platycerium bifurcatum typically have ations of the members of this complex. Some of these trade
the distal part slightly to moderately extended and shal- plants may be recognized as belonging to one of the four
lowly to moderately lobed. The foliage fronds are erect to species in the complex, whereas others are more difficult to
arching or pendent, typically with a short stipe that has the identify because of intergrading or inconstant characters.
vascular bundles arranged in a circle. The blades are several The particular frond form of a plant may not be the same in
times forked into narrow segments, and the sporangial later growth or under different cultural conditions. Fronds
patches are borne on the ultimate forks. Emerging fronds can vary by season and age.
soon turn to a horizontal position, instead of remaining Platycerium willinckii var. venosum Hughes is closer to P.
vertical for some distance as in the similar P. willinckii. The bifurcatum than to P. willinckii because of its strongly
species is native to eastern Australia. stalked (not sessile) foliage fronds. It differs from typical P.
The preceding description excludes Platycerium hillii, bifurcatum by having widely spreading base fronds, usually
P. veitchii, and P. willinckii, which are sometimes consid- longer foliage fronds, and strongly raised major veins on the
ered varieties or subspecies of P. bifurcatum but are here upper surface (but depressed on the lower) especially near
treated as separate species. Collectively, these species and the blade base. It was collected in northeastern Australia.
P. bifurcatum are referred to as the “P. bifurcatum com- Native var. venosum reportedly intergrades into typical P.
plex.” They all have a circular arrangement of vascular bun- bifurcatum plants (Roy Vail, personal communication).
dles in the stipes and lack dark tissue encircling the bundles. ‘Netherlands’. Fronds arching, grayish, the base fronds
(This trait can be seen by making a clean cut across the stipe swept upward, lobed on the upper margins. Common in
with a razor blade or sharp knife.) Most of the numerous, the trade.
loosely given Platycerium names in the trade represent vari-
F F A C 431
Platycerium coronarium (König ex Müller) Desvaux Platycerium ×elemaria Hoshizaki & M. G. Price
13.99.7 13.99.8
syn. Platycerium biforme (Swartz) Blume Tender
Disk staghorn A medium-large fern with a short rhizome and bud-form-
Tender ing roots. Grows well under medium to high-medium light
A large fern with a short rhizome that may branch and be- in drained, moist to moist-dry, coarse potting mix or uncut
come longer; new plants form when the tip of the branch moss.
reaches the surface. Grows well under high light in drained, Platycerium ×elemaria is a sterile hybrid of P. elephanto-
moist-dry, coarse potting mix or uncut moss. tis and P. stemaria. Its base fronds resemble those of the
The base fronds of Platycerium coronarium have the dis- parents, and the foliage fronds resemble those of P. elephan-
tal part extended, deeply lobed, and forward-arching. The totis but have the apical margins shallowly and irregularly
foliage fronds are long, pendent, and several times forked. forked or lobed. Hydathodes are absent. This plant origi-
The sporangial patch is typically a kidney-shaped lobe. This nated as a hybrid in horticulture and does not occur in the
species is native to southeastern Asia, Malaysia, Indonesia, wild. The cultivar name of this hybrid is ‘Sanchez’.
and the Philippines.
c
Figure 13.99.8. Platycerium ×elemaria: a. habit, bar =
10 cm (4 in.); b. sporangial area placement; c. rhizome
scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar =
1 cm (0.4 in.).
a
c
a c
d
Figure 13.99.12. Platycerium hillii: a. habit, bar = 10 cm
(4 in.); b. sporangial area placement; c. rhizome scales,
bar = 5 mm (0.2 in.); d. stipe cross section, bar = 1 cm
(0.4 in.).
a c
d
b
c
Platycerium ridleyi H. Christ 13.99.17
Tender
A medium-sized fern with short rhizomes. Grows under
high light in drained, moist-dry to dry, coarse potting mix
or uncut moss. This species is difficult to grow and is at-
tacked by a variety of insect pests.
Platycerium ridleyi has round to kidney-shaped base
fronds with unlobed or coarsely dentate margins. The fo-
liage fronds are erect, and the sporangial patch is borne on
an obovate lobe. This species is native to western Malaya
Figure 13.99.15. Platycerium ×mentelosii: a. habit, bar = and Indonesia.
10 cm (4 in.); b. sporangial area placement; c. stipe cross
section, bar = 1 cm (0.4 in.).
F F A C 435
b a
a d
d
Figure 13.99.18. Platycerium stemaria: a. habit, bar = Figure 13.99.19. Platycerium superbum: a. habit, bar =
10 cm (4 in.); b. sporangial area placement and frond 10 cm (4 in.); b. sporangial area placement; c. rhizome
variations; c. rhizome scales, bar = 5 mm (0.2 in.); scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar =
d. stipe cross section, bar = 1 cm (0.4 in.). 1 cm (0.4)in.
436 C 13
a
b c
d
c d
Figure 13.99.21. Platycerium wallichii: a. habit, bar = Figure 13.99.22. Platycerium wandae: a. habit, bar =
10 cm (4 in.); b. sporangial area placement; c. rhizome 10 cm (4 in.); b. sporangial area placement; c. rhizome
scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar = scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar =
1 cm (0.4 in.). 1 cm (0.4 in.).
F F A C 437
e
d
Campyloneurum: Rhizomes short- veins pointing toward the margins. Native to Africa and
creeping; rhizome scales clathrate; Asia. (Lepisorus longifolius, L. thunbergianus)
fronds medium-sized, simple, en-
tire, strap-shaped or linear-lance- Leptochilus (Colysis): Rhizomes
olate, monomorphic; veins net- short- to long-creeping; rhizome
ted, lateral veins parallel and scales peltate, clathrate; fronds
connected by curved cross-veins medium-sized; stipes often
to form the areoles, the included winged; blades simple to one-
veinlet unbranched and pointing pinnate, monomorphic or di-
toward the margin; sori round, morphic; veins netted, included
borne below the free veinlet tip. veinlet pointing in various direc-
Native to tropical and subtropical tions; similar to Microsorum ex-
America. (Campyloneurum angustifolium, C. brevifolium, cept sori are mostly elongate to
C. costatum, C. phyllitidis, C. tenuipes, C. xalapense) linear and oblique to the axis.
Native to Asia. (Leptochilus amplus, L. ellipticus, L. macro-
Dictymia: Rhizomes short-creep- phyllus var. wrightii )
ing; rhizome scales peltate, broadly
ovate, clathrate, usually with glan- Microgramma: Rhizomes long-creeping, often
dular marginal projections; fronds slightly flattened; rhizome scales elongate, not
medium-sized, simple, entire, lin- clathrate, attached well above their base; fronds
ear to lanceolate, subleathery, less than 39 cm (15 in.) long; blades simple, en-
smooth; veins netted, included tire, lanceolate to oblong, slightly dimorphic;
veinlets absent or rare; sori round veins netted with unbranched included veinlets
to narrowly oblong, slightly to pointing outward toward the margin; sori round.
strongly embossed on the blade. Native to Australia and Native to the American tropics and subtropics.
the islands of the southwestern Pacific. (Dictymia brownii ) (Microgramma heterophylla, M. lycopodioides, M.
megalophylla, M. nitida, M. percussa, M. pilosel-
Goniophlebium: Rhizomes medium- loides, M. reptans, M. squamulosa, M. vacciniifolia)
to long-creeping; rhizome scales pel-
tate, clathrate, toothed, often with Microsorum (Phymatosorus):
one- or two-celled, glandular marginal Rhizomes short- to long-
appendages; fronds medium- to large- creeping; fronds medium- to
sized; blades pinnatifid to pinnate large-sized, simple, margins
with a terminal pinna resembling the entire, lobed, or pinnatifid;
lateral ones; pinnae jointed; veins net- veins netted, included vein-
ted, areoles angular and in several lets simple or branched, vari-
rows, costal areoles large, with one included veinlet point- ously directed, always ending
ing outward. Native to Asia. (Goniophlebium ×ekstrandii, in a hydathode; sori scattered
G. percussum, G. persicifolium, G. subauriculatum) dots or larger, round to oval, immersed in the blade tissue
or not. Native to Africa, Asia, Australia, and islands of the
Lecanopteris: Rhizomes short-creep- southwestern Pacific. (Microsorum commutatum, M. for-
ing and swollen, with internal tunei, M. grossum, M. insigne, M. linguiforme, M. longissi-
chambers inhabited by ants; rhi- mum, M. lucidum, M. membranifolium, M. musifolium,
zome scales peltate, clathrate cen- M. pteropus, M. punctatum,
trally but not at the margin; fronds M. pustulatum, M. samarense,
small-sized; blades entire to deeply M. scandens, M. scolopendria,
pinnatifid; veins netted, the areoles M. steerei, M. superficiale, M.
with included free veinlets. Native viellardii)
to Malaysia. (Lecanopteris car-
nosa, L. crustacea, L. sinuosa) Niphidium: Rhizomes creep-
ing to ascending; fronds clus-
Lepisorus: Resembles Pleopeltis, tered, medium- to large-
except the rhizome scales are en- sized, simple, entire, upper
tirely clathrate and the outer row surfaces often have many
of areoles do not produce free white dots; sori round, large
440 C 13
(3–5 mm in diameter), in a single row between the main paraphyses (if present) not peltate. Cosmopolitan. (Polypo-
lateral veins, not sunken; sporangial capsules with minute dium amoenum, P. appalachianum, P. attenuatum, P. bom-
hairs. Native to the American tropics and subtropics. bycinum, P. brasiliense, P. calirhiza, P. cambricum, P. dis-
(Niphidium crassifolium) simile, P. formosanum, P. glycyrrhiza, P. guttatum, P.
hesperium, P. hirsutissimum, P. interjectum, P. ×leucospo-
Pecluma: Rhizomes very short-creep- rum, P. levigatum, P. loriceum, P. maritimum, P. menisci-
ing; rhizome scales attached along ifolium, P. pellucidum, P. plebeium, P. polypodioides, P.
their base; fronds small- to medium- ptilorhizon, P. pyrrholepis, P. rhodopleuron, P. sanctae-rosae,
sized; blades deeply pinnatisect, comb- P. scouleri, P. thyssanolepis, P. triseriale, P. virginianum, P.
shaped, the many divisions narrow vulgare)
and close together, with small one-
celled hairs on lower surfaces; sori Selliguea (Crypsinus): Rhizomes
round. Native to the American tropics long-creeping, conspicuous
and subtropics. (Pecluma dispersa, P. pectinata, P. plumula) black fiber strands within; rhi-
zome scales peltate; fronds me-
Phlebodium: Rhi- dium-sized, simple, entire, lan-
zomes medium- to ceolate to oblong-lanceolate,
long-creeping; rhi- some species (formerly in Crypsi-
zome scales usually nus) hastate, pinnatifid to pin-
orangish when freshly nate with cartilaginous and often
mature, not clathrate; notched margins; fertile fronds
fronds medium- to taller, more contracted; veins
large-sized, pinnati- netted, lateral veins conspicuous; sori brown, round but
lobed or pinnatifid; usually elongate or linear, borne between adjacent lateral
areoles next to the midrib mostly narrow and without in- veins in one or two rows. Native to southern Africa, Asia,
cluded veinlets; other areoles with included veinlets or not, Australia, and the islands of the Pacific. (Selliguea feei)
if present the veinlets are united or, if free, usually pointing
outward toward the margin; sori supplied by two veinlets. Solanopteris: Rhizomes long-creep-
Native to the American tropics and subtropics. (Phlebodium ing, with ant-inhabited tubers,
aureum, P. decumanum, P. pseudoaureum, P. ×schneideri) without black fiber stands within;
rhizome scales peltate, round;
Pleopeltis: Rhizomes long-creeping; rhi- fronds small-sized, simple to pin-
zome scales clathrate centrally but not at natifid; fertile fronds usually nar-
the margins; fronds small- to medium- rower and longer than the sterile;
sized, simple, entire (rarely dichoto- veins netted with included veinlets;
mously forked), surfaces with peltate sori round or linear. Native to the
scales; veins netted with included American tropics. (Solanopteris
branched(?) veinlets pointing in various bifrons, S. brunei)
directions, the marginal row of areoles
with free veins pointing toward Even with the removal of these
the margins; sori usually round, segregate genera, Polypodium con-
covered with peltate paraphyses tains many species. To facilitate identification of these spe-
when young. Native to the Amer- cies, Polypodium is divided into three groups.
ican tropics and subtropics, and
Africa. (No cultivated species; The Vulgare Group (Polypodium vulgare complex). This
plants brought into cultivation group is characterized by free veins (rarely with a few netted
have not survived.) ones) and pinnatifid (or pinnatilobed) blades with scaleless
surfaces. The sori are always in one row between the mid-
Polypodium: Rhizomes short- to rib and margin of the blade. The vulgare group contains
long-creeping; fronds small to about 25 species, most of which are native to the boreal and
large; blade pinnatifid or pinnate; temperate regions of North America, Europe, and Asia. A
pinnae not jointed; veins free or few occur in Mexico, and one species (Polypodium pellu-
netted, with or without a free in- cidum) is native to Hawaii.
cluded veinlet; sori round or ob- The most commonly cultivated species in the group are
long, at tip or junction of vein, of European origin. Other species may be cultivated but
F F A C 441
are usually difficult to establish. Pteridologists have named Polypodium amoenum Wallich ex Mettenius
and described many species and hybrids that may be diffi- 13.101.3
cult for nonspecialists to identify, especially if the plant’s syn. Goniophlebium amoenum (Wallich ex Mettenius) J.
original habitat is unknown. Even if this is known, fertile Smith ex Hooker & Greville
fronds and a microscope are usually needed for accurate Semi-tender to hardier
identification. A small to medium fern with long-creeping rhizomes.
The cultivated species of the vulgare group treated here Grows well under low to high light in moist-dry, well-
include Polypodium appalachianum, P. calirhiza, P. cambri- drained garden soil or potting mix. The plants are subject
cum, P. glycyrrhiza, P. hesperium, P. interjectum, P. pelluci- to slug and snail attack and require a lot of humidity.
dum, P. scouleri, P. virginianum, and P. vulgare. They are The rhizomes of Polypodium amoenum contain scattered
illustrated together in Figure 13.101.34. dark fibers, and the fronds are generally 14–50 cm (5.5–20
The Loriceum Group (Polygoniophlebium). This group is in.) long and 8–27 cm (3–11 in.) wide, with pinnatifid,
characterized by pinnatifid to pinnate, rarely simple blades dull green blades. The segments typically have pentagonal
that have scaleless surfaces (a few scales might be present areoles next to the rachis. The sori are in one row between
on the rachis or costae). The veins form at least one row of the pinna midrib and margin. The species is native to the
regular areoles along the pinna midrib. Each areole con- Himalayas, southeastern Asia, China, and Taiwan.
tains a veinlet that points toward the blade margin. The Some pteridologists classify this species in Goniophle-
pinnae are continuous with the rachis, not jointed to it by bium, but it lacks jointed pinnae (a defining characteristic
an abscission layer. The sori appear in one to several rows of that genus) and is therefore retained in Polypodium until
between the costae and margins. The group contains about further study.
40 species, most of which are native to the New World.
Two widely grown species in the loriceum group are Polypodium appalachianum Haufler & Windham
Polypodium triseriale and P. menisciifolium. The former is 13.101.4, 34d
grown as a basket plant in gardens in Florida. Polypodium Appalachian polypody
menisciifolium, which is cultivated in baskets in southern Very hardy, Zone 3
California, grows vigorously and tolerates irregular water- A small fern with short- to medium-creeping rhizomes.
ing. Another species in the group, P. formosanum, is often Grows under medium light in drained, moist potting mix.
found in the trade and is prized for its attractive pale gray- This species adapts poorly to gardens in southern California.
ish green fronds and rhizomes. A cultivar with repeatedly
forked and congested rhizomes, ‘Cristatum’, adds further
interest to this species.
The cultivated species treated here are Polypodium amoe- b
num, P. attenuatum, P. brasiliense, P. dissimile, P. formo-
sanum, P. levigatum, P. loriceum, P. maritimum, P. meniscii-
folium, P. ptilorhizon, P. rhodopleuron, and P. triseriale.
The Scaly-Polypody Group. This group is characterized
by pinnatifid (or pinnatilobed) blades that are scaly, espe-
cially on the lower surface. Often the scales are dense and
conspicuous. The veins vary from free to netted, but they d
are usually hard to see because of the thick blade tissue or
because they are hidden by the scales. The sori are in one c
row between the costa and margin. All the species in the
scaly-polypody group, of which there are about 40, are na-
tive to tropical America. a
This group is closely related to Pleopeltis, and probably
all the species should be transferred to that genus, as has al-
ready been done with some species. The cultivated species
treated here are Polypodium bombycinum, P. guttatum, P.
hirsutissimum, P. ×leucosporum, P. plebeium, P. polypodi-
oides, P. pyrrholepis, P. sanctae-rosae, and P. thyssanolepis. Figure 13.101.3. Polypodium amoenum: a. habit, bar =
5 cm (2 in.); b. vein details along the rachis, bar = 1 cm
(0.4 in.), after Hoshizaki (1982); c. rhizome cross
section, showing vascular bundles and scattered fibers,
bar = 1 cm (0.4 in.); d. rhizome scale, bar = 1 mm.
442 C 13
a
c
b
a b
a
Figure 13.101.8. Polypodium
calirhiza: a. habit, bar = 5 cm
(2 in.); b. base of segment with
one areole, bar = 5 mm (0.2 in.).
tips), and the sori vary from round to oval and lack para-
physes. The species is native to Asia and the western coast
of North America.
a a
Figure 13.101.16.
Polypodium interjectum:
a. frond variations, bar
= 5 cm (2 in.);
b. sporangium, bar =
0.5 mm, after Shivas
(1962).
a b
a
by Wagner and Wagner (1975). The spores are misshapen, Polypodium loriceum Linnaeus 13.101.19
irregular, and aborted. Sterile hybrids such as this one are Tender
one reason pteridologists believe that the scaly-polypody A small-medium fern with medium-creeping rhizomes.
group is more closely related to Pleopeltis than to other Grows well under medium light in drained, moist potting
groups of Polypodium. mix.
Polypodium loriceum resembles P. maritimum, but its
Polypodium levigatum Cavanilles 13.101.18 rhizome scales are less distinctly clathrate, its blades are
syn. Polypodium glaucophyllum Kunze ex Klotzsch glabrous, and only one row of sori is found between the
Tender pinna midrib and margin. This species is widespread in the
A small fern with creeping, gray-green rhizomes bearing a American tropics.
few minute, scattered scales. Grows in medium light in As presently defined, Polypodium loriceum is extremely
moist to moist-dry, well-drained potting soil or uncut moss. variable and includes several geographic entities that prob-
The simple and entire fronds of Polypodium levigatum ably represent different species. This matter, however,
are unusual. They are gray-green, elliptic-lanceolate, and needs further research.
smooth except for a few ovate-acuminate scales on the
lower surface of the rachis. The veins are netted. It is native
from Central America to South America.
a b
a
c
den soil or potting mix kept well drained. The plants are
easy to cultivate in hanging baskets or in the ground.
Polypodium pyrrholepis has tufted, bright rusty-red rhi-
zome scales. The fronds are set close together, with blades
narrowly linear-oblong, pinnatisect, and densely scaly on
both surfaces. A small, slightly swollen gland is on the in-
side, upper edge of the segments. This species is native to
southern Mexico.
b
b
a
Figure 13.101.29. Polypodium
scouleri: a. frond, bar = 5 cm
(2 in.); b. rhizome scales, bar
= 3 mm (0.1 in.).
d c
b
a
b
Figure 13.101.32. a
Polypodium virginianum:
a. habit, bar = 5 cm
(2 in.); b. paraphysis,
top view, bar = 0.1 mm.
b
b
c
b
Figure 13.101.33. Polypodium
Figure 13.101.31. Polypodium triseriale: a. habit, bar = vulgare: a. habit, bar = 5 cm
5 cm (2 in.); b. pinna with sori, bar = 1 cm (0.4 in.); (2 in.); b. sporangium, bar =
c. rhizome scale, bar = 1 mm. After Hoshizaki (1982). 0.5 mm, after Shivas (1962).
a
b d e
c f
l
m
k
i
Figure 13.101.34. Frond silhouettes of species in the Polypodium vulgare complex. European species: a. P. cambricum;
b. P. vulgare; c. P. interjectum. Eastern North American species: d. P. appalachianum; e. P. sibericum; f. P. virginianum.
Western North American species: g. P. californicum; h. P. glycyrrhiza; i. P. calirhiza; j. P. amorphum; k. P. hesperium;
l. P. scouleri. Hawaiian species: m. P. pellucidum. Bars = 5 cm (2 in.). Plants c, f, i, and k of hybrid origin; see text entries
for details.
454 C 13
cept in very hot, humid places. The northwestern species Fronds two-pinnate-pinnatifid or up to three-pinnate, at
can be grown in the southwest if provided with high hu- least at the base of the lower pinnae: Polystichum proliferum,
midity and acid soils. Because only a few species produce P. richardii, P. silvaticum, P. tagawanum, P. tsus-simense,
multiple crowns that can be divided, most polystichums P. vestitum
are propagated from spores. Many species also form buds
along their rachises or at the tips of the fronds. These buds Polystichum acrostichoides (Michaux) Schott
are easily rooted by pegging the leaf to the soil. Thrip dam- 13.102.4
age is often conspicuous on fronds of Polystichum, appear- Christmas fern
ing as a grayish silvering on the blade. Very hardy, Zone 3
Polystichum has erect, ascending (rarely short-creeping) A medium-sized fern with ascending, clump-forming rhi-
rhizomes. The fronds are mostly one- to two-pinnate, less zomes and evergreen leaves. Grows well under low to me-
often three-pinnate, pinnatifid. They are elongate, often dium light in moist garden soil or potting mix. Although
narrowed at the base, with the apex gradually reduced. The easily cultivated in the eastern United States, the plants are
pinnae often have an upward-pointing “ear” at their base— difficult to grow on the West Coast. In winter, the fronds
a characteristic helpful in distinguishing the genus from recline to the ground, forming a rosette around the rhizome
Dryopteris, which often lacks the ear. Some one-pinnate apex.
species have terminal pinnae that resemble the lateral ones. Polystichum acrostichoides is characterized by one-pin-
The marginal teeth generally bear a short bristle or spine. nate blades with the distal, fertile pinnules narrower than
The veins are anadromic and free, not netted. Round sori the sterile ones. The pinnae have an upward-pointing “ear”
are formed in one row (or infrequently two) between the at the base, and the margins are spinulose. When the sori
midrib and margin. Indusia may be present or absent; when mature they tend to become confluent, resembling acrosti-
present, they are peltate. choid sori. The species is native to eastern North America.
Some authors believe that Cyrtomium (including Pha- A sterile hybrid, Polystichum ×potteri Barrington (= P.
nerophlebia) is so closely related to Polystichum that it acrostichoides × P. braunii ), is sometimes cultivated. The
should be combined with it. Cyrtomium, however, differs hybrid is native to Pennsylvania, New England, Nova Sco-
by having netted veins and sori in two or more series. tia, New Brunswick, and Quebec.
The genus Polystichum contains 180 species and is cos- Various cultivar names are listed in the trade, such as
mopolitan. For details on the many varieties, see Dyce ‘Crispum’, ‘Cristatum’, and ‘Incisum’.
(1963), Kaye (1968), and Mickel (1994). The genus name
comes from the Greek poly, many, and stichos, row, refer-
ring to the sori of the type species (Polystichum aculeatum),
which are arranged in many regular rows on the pinnae.
Because Polystichum contains so many species, the fol-
lowing subgroups are given to aid identification. Once an
unknown species is limited to one of these subgroups, it
can be compared to the descriptions and illustrations of Figure 13.102.4. Polystichum
other species in the group. The groups are based on typical acrostichoides: frond, bar =
fronds (not cultivars) from mature plants. 5 cm (2 in.).
Fronds one-pinnate, the pinnae entire, serrate, or crenate,
not pinnatifid: Polystichum acrostichoides, P. craspedosorum,
P. imbricans, P. lepidocaulon, P. lonchitis, P. munitum, P.
nepalense, P. stenophyllum, P. stimulans
c
Figure 13.102.5. Polystichum
aculeatum: a. frond, bar =
5 cm (2 in.); b. pinna, bar =
1 cm (0.4 in.). Figure 13.102.6. Polystichum
andersonii: a. frond, bar =
5 cm (2 in.); b. pinna, bar =
1 cm (0.4 in.); c. bud on
rachis, bar = 2 mm.
a
b
Polystichum braunii (Spenner) Fée
13.102.7; 6
Braun’s holly fern, prickly shield fern
Very hardy, Zone 3(4)
A small to medium fern with clump-forming rhizomes and
dark green, shiny, evergreen fronds. Grows well under me-
dium light in moist potting mix. The plants do best if
F F A C 457
b
placed in a cool site. The fiddleheads are particularly at-
tractive because they are densely covered by silvery scales,
which turn light brown with age.
Polystichum braunii has short stipes, usually about one-
quarter the length of the frond. The stipes are densely scaly,
and the blades are two-pinnate and elliptic-lanceolate, grad-
ually narrowed toward the base. The pinnules are short-
stalked and bristly. This species is native to North America,
Europe, and Asia.
Figure 13.102.11.
Polystichum imbricans:
a. frond, bar = 5 cm
(2 in.); b. pinnae, bar
= 1 cm (0.4 in.), after
D. Wagner (1979).
b
b
Figure 13.102.18.
Polystichum munitum: a
a. frond, bar = 5 cm
(2 in.); b. pinna, bar =
1 cm (0.4 in.).
b
b
a c
Figure 13.102.21.
Polystichum
polyblepharum:
a. frond, bar =
5 cm (2 in.);
b. pinna, bar =
1 cm (0.4 in.);
c. pinnule, bar =
5 mm (0.2 in.).
a
Polystichum polyblepharum (J. Roemer ex Kunze) Polystichum proliferum (R. Brown) C. Presl
C. Presl 13.102.1, 2, 21 13.102.2, 3, 22
Bristle fern, Japanese sword fern Mother shield fern
Hardy, Zone 5(6) Hardy, Zone 5
A medium-sized fern with erect rhizomes and dark green, A medium-sized fern with erect rhizomes and dark green
glossy, evergreen fronds. Prefers high-medium light and fronds that are evergreen in warmer climates. Grows well
moist or moist-dry garden soil or potting mix. This species under medium light in moist garden soil or potting mix.
is easy to grow. This species is easy to grow and can be propagated from
Polystichum polyblepharum has densely scaly stipes and the bulbils on the fronds.
rachises. The blades are elongate-elliptic to oblong and two- The blades of Polystichum proliferum are lanceolate to
pinnate. The pinnules are eared, with the acroscopic ear broader at the base and two- to three-pinnate, with a bud
prominent and spine-tipped and the basiscopic ear incon- forming on the apical part. The pinnules are toothed. The
spicuous with its spine often bent over the upper pinnule species is native to Australia and New Zealand.
surface. This species is native to Japan, southern Korea, and This species resembles Polystichum vestitum but is iden-
eastern China. tifiable by the bulbils produced near the frond apices and by
The plants in the trade are widely misidentified as Poly- the glossy stipe scales with dark centers and pale borders.
stichum setosum Horticulture. The name Polystichum proliferum might be confused with
Polystichum retrosopaleaceum (Kodama) Tagawa some- P. setiferum cultivars that bear buds on their rachis and have
what resembles P. polyblepharum but differs by lower stipe circulated incorrectly under the same name.
scales broadly ovate with a long point (versus lanceolate-
triangular), the pinnae apices more acuminate-pinnatifid
(versus acute and subconform), and the foliage clear green
(versus dark green). It is rarely cultivated in the United
F F A C 463
a b c
d
e f
Figure 13.102.27. Polystichum setiferum cultivars: a. ‘Congestum’; b. Divisilobum Group; c. Imbricatum Group; d.
Multilobum Group; e. Plumoso-Divisilobum Group; f. ‘Trilobum’. Bar = 5 cm (2 in.).
Multilobum Group. Figure 13.102.27d. Resembles Di- and congested; fronds bud-bearing. A very distinct, com-
visilobum Group but with pinnules stouter, nearly rectan- pact plant. Sometimes circulates as ‘Divisilobum Grandi-
gular, and not overlapping; some variants are bud-bearing. ceps’ or ‘Divisilobum Cristatum’ in the United States trade.
Plumoso-Divisilobum Group (‘Plumoso-Multilobum’).
Divided soft shield fern. Figure 13.102.27e; Plate 46.
Fronds 60–100 cm (2–3 ft.) long, horizontally spreading, Polystichum setigerum (C. Presl) C. Presl
to three-pinnate; pinnae overlapping; basiscopic pinnules 13.102.28
linear-lanceolate, longer than the acroscopic pinnules. This syn. Polystichum braunii subsp. alaskense (Maxon) Calder
beautiful fern has a feathery, neatly three-dimensional as- & R. L. Taylor
pect due to the spreading angle of the overlapping pinnae. Alaskan holly fern
‘Trilobum’ (‘Divisilobum Grandiceps’). Figure Hardy, Zone
13.102.27f. Pinnules have the same shape as those of Di- A medium-sized fern with ascending to erect rhizomes and
visilobum Group, but the distal half of the blade is a mass evergreen fronds. Grows under medium light in moist pot-
of branches. Blades to about 30 cm (12 in.) long, rachises ting mix. The plants do best in cool, moist climates.
branched, branches often irregular and branched one or The stipes of Polystichum setigerum are about half the
more times making the far (distal) part of the blade broad length of the fronds, and the blades are lanceolate and one-
466 C 13
b
Figure 13.102.30. Polystichum stenophyllum: a. frond, bar
= 5 cm (2 in.); b. pinnule, bar = 1 cm (0.4 in.); c. stipe
base scale, bar = 3 mm (0.1 in.). After Hu and Ching
(1930).
Figure 13.102.34.
Polystichum tsus-simense
var. tsus-simense:
a. frond, bar = 5 cm a
(2 in.); b. pinna, bar =
1 cm (0.4 in.).
Figure 13.102.35.
Polystichum tsus-
Polystichum tripteron is distinctive by its greatly elon-
simense var.
gated basal pinnae, which impart an inverted T-shape to
mayebarae: a.
the blades. The blades are two-pinnate. The long apex is
frond, bar = 5 cm
one-pinnate above the lateral pinnae. The pinnules are dull
(2 in.); b. pinna,
green (not shiny) and coarsely toothed, without bristly
bar = 1 cm
teeth. This species is native to eastern Siberia, eastern
(0.4 in.). a
China, Korea, and Japan.
Pteridium
Bracken
This well-known native of open fields and slopes is a me-
dium- to large-sized terrestrial fern that spreads from a slen-
der, wide-creeping, underground rhizome. Its triangular
fronds are spaced far apart and are several times divided.
Pteridium is rarely used as an ornamental because it spreads
invasively in the garden. It is a terrible weed in many parts
of the world.
The fiddleheads of bracken are harvested in Japan for
Figure 13.103.1. Psilotum nudum: habit. food, and in the United States the packages are sold in some
F F A C 471
d
e
Figure 13.105.2.
Pteris vittata:
sorus and
indusium.
a b
c
a
d
b
h
e
f g
Figure 13.105.9. Pteris cretica cultivars: a. ‘Childsii’; b. ‘Mayii’; c. ‘Ouvrardii’; d. ‘Parkeri’; e. ‘Rivertoniana’; f. ‘Roweri’; g.
‘Wilsonii’; h. ‘Wimsettii’. Bar = 5 cm (2 in.).
mate divisions narrowly linear. This species is native to near the midrib but netted toward the margin. This spe-
India, China, Malaysia to Australia, and Polynesia. cies is native to the American tropics.
‘Evergemiensis’. Variegation more cream-white than in
‘Victoriae’, and the white extends between the lateral veins. Pteris longifolia Linnaeus 13.105.13
‘Victoriae’. Victorian brake. Figure 13.105.11b. Cream- Ladder brake
white band down the center of pinnae and pinnules, with Tender
only some white extending along the lateral veins. A medium fern with short-creeping rhizomes and clustered
fronds. Grows well under medium to high light in moist
Pteris grandifolia Linnaeus 13.105.12 garden soil or potting mix.
Tender Pteris longifolia has oblanceolate, one-pinnate blades.
A large fern with short-creeping rhizomes and clustered The stipes and rachises of unweathered plants are amply
fronds. Grows well under high-medium light in moist gar- covered with scales and hairs. The pinnae are numerous,
den soil or potting mix. Too large for most gardens, this simple, and jointed to the rachis. The veins are free. The in-
huge plant is grown only in south Florida. dusium is erose-fimbriate. This species is native to the
Pteris grandifolia has one-pinnate blades throughout, United States (Florida), Mexico, Guatemala, and the West
and the pinnae are simple and entire. The veins are free Indies.
b
a b
a
Figure 13.105.10. Pteris Figure 13.105.12. Pteris grandifolia: a. frond, bar = 5 cm
dentata: a. frond, bar = (2 in.); b. pinnae, bar = 5 cm (2 in.).
5 cm (2 in.); b. pinna tip,
bar = 1 cm (0.4 in.).
b
a
Most of the plants sold in the United States under this winged only between two upper pinna pairs or less.) This
name are actually Pteris vittata. The two species are often species is native to China and Japan.
difficult to distinguish. The best characteristic to separate A few crested and compact-growing cultivars are some-
them is the jointed pinna bases of P. longifolia. These bases times seen.
have a dark abscission layer where the pinna attaches to the
rachis, and the juncture is usually thickened. (Pteris vittata Pteris nipponica W. C. Shieh 13.105.15
never has thickened rachises at the juncture with the pin- syn. Pteris cretica var. albolineata Hooker
nae.) These characteristics are often hidden by scales, which Semi-hardy, Zone 8
must be scraped away to identify the plant. A small to medium fern with short- to medium-creeping
rhizomes and fronds forming somewhat open clumps.
Pteris multifida Poiret 13.105.14 Grows well under medium light in moist to moist-dry gar-
syn. Pteris serrulata Linnaeus f. den soil or potting mix. This species is more commonly
Spider brake known by its synonym: Pteris cretica var. albolineata (or
Hardy, Zone 6 ‘Albo-lineata’).
A small fern with clump-forming rhizomes and fronds in Pteris nipponica is distinguished by a pale band of tissue
dense, irregular clusters. Grows well under medium light in on both sides of the pinna midrib. It differs from P. cretica
moist to moist-dry garden soil or potting mix. The plants by having a more open-spreading habit, by the absence of a
tend to be weedy. distinct wing on the distal part of the rachis, and by other
Pteris multifida resembles P. cretica but differs by nar- microscopic spore details. This species is native to Japan,
rower pinnae and the rachis fully winged between each of South Korea, and Taiwan.
the three upper pinna pairs. (Pteris cretica is unwinged or
Pteris quadriaurita Retzius 13.105.16
Semi-tender
A medium-large fern with clustered fronds and infrequently
branched, short-creeping rhizomes. Grows best under me-
dium light in moist-dry garden soil or potting mix.
The blades of Pteris quadriaurita are herbaceous and
one-pinnate-pinnatifid except for the basal pinnae that are
forked at the base. The costa (but not the costules) bears
a
b
b
small appressed scales. The veins are free, not netted, along
the midrib. This species is native to tropics worldwide.
Pteris quadriaurita comprises a complex of variable
plants not fully studied. Plants examined in the trade la-
beled as P. fauriei Hieronymus are actually P. quadriaurita.
True P. fauriei, native to eastern Asia, has leathery fronds
and pinnae that are widest in the middle with caudate
apices.
The blades of Pteris tremula are triangular, membran- Pteris vittata Linnaeus 13.105.2, 20
ous, and two- to four-pinnate-pinnatifid. The ultimate seg- Chinese brake, rusty brake
ments are narrowly oblong to more linear and serrate with Semi-hardy, Zone (7)8
blunt teeth. The fertile segments are narrower than the ster- A medium-large fern with short-creeping rhizomes and
ile ones. The veins are free, not netted. This species is native clustered fronds. Grows well under high-medium light to
to Australia, Tasmania, New Zealand, and Fiji. full sun in moist-dry garden soil or potting mix. The plants
are easy to start from spore, but they tend to become weedy.
Pteris tripartita Swartz 13.105.19 They grow on old masonry and bricks, as can be seen in
Trisect brake the French Quarter in New Orleans.
Tender, Zone 9 Pteris vittata is characterized by one-pinnate blades wid-
A large fern, to 2 m (6.5 ft.) tall, with clustered fronds and est above the middle, with narrow pinnae and a terminal
clump-forming rhizomes. Grows well under medium light pinna that resembles the lateral ones. The stipes and
in continually moist-wet garden soil or potting mix. The rachises are sparsely scaly-hairy, and the pinnae are not
plants grow rapidly from spores but become too big for jointed to the rachis. The veins are free, and the indusia
most gardens. have entire margins. This species greatly resembles Pteris
Pteris tripartita has broadly triangular blades divided longifolia. It is native to eastern Asia but has become widely
into three parts: the central part is ovate-triangular and one- naturalized throughout the tropics and subtropics of the
pinnate-pinnatifid, and the two lateral parts (the basal pin- world.
nae) are as long as the central part, and each forks twice on
the lower side. The veins are netted. This species is native to
the Old World but has become naturalized in tropical
America.
b
Pyrrosia 13.106.1–3
syn. Cyclophorus, Drymoglossum, Niphobolus
Felt fern
a
Most Pyrrosia species are small-medium-sized and have
simple, entire blades (a few are deeply lobed) with star-
shaped hairs. Be careful not to overwater pyrrosias. The
Figure 13.105.19. Pteris tripartita: a. small frond, bar = epiphytic species in particular need a coarser soil mix and
5 cm (2 in.); b. large frond, bar = 15 cm (0.5 ft). good drainage. The plants can be easily propagated by di-
vision of the rhizomes.
F F A C 481
a b
Figure 13.106.4.
Figure 13.106.5. Pyrrosia confluens: a. var. confluens
Pyrrosia angustata:
habit, bar = 5 cm (2 in.); b. var. confluens sori, bar =
a. habit, bar = 5 cm
1 cm (0.4 in.); c. var. dielsii habit, bar = 5 cm (2 in.).
(2 in.); b. sori, bar =
1 cm (0.4 in.).
Figure 13.106.7.
Pyrrosia eleagnifolia:
a. habit, bar = 5 cm
(2 in.); b. sori, bar =
b 1 cm (0.4 in.).
b
Pyrrosia fallax (Alderwerelt) M. G. Price
13.106.8
Tender (?)
A small fern with medium-creeping rhizomes. Grows well
a under medium light in moist-dry potting mix or uncut
moss with good drainage.
Figure 13.106.6. Pyrrosia costata: a. habit and larger Pyrrosia fallax has slender rhizomes covered by round-
frond, bar = 5 cm (2 in.); b. stipe cross section, bar = ish, dark scales with pale erose-ciliate margins. The phyl-
1 mm. lopodia are about 1–1.5 cm (about 0.5 in.) apart, and lat-
eral buds occurs on the rhizome about midway between
phyllopodia. The fronds are dimorphic. The sterile ones
are obovate or rounder, 1–4.5 cm (0.5–2 in.) long, and
Pyrrosia eleagnifolia (Bory) Hovenkamp nearly glabrous above but with pale hairs below. The fertile
13.106.7 fronds are narrowly linear, about 4–10 cm (1.5–4 in.) long.
syn. Pyrrosia rupestris of authors, not (R. Brown) Ching, The sori are arranged in two or three irregular rows or are
P. serpens of authors, not (N. L. Forster) Ching
Semi-tender to hardier
A small fern with medium-creeping rhizomes. Grows
under medium light in moist-dry potting mix or uncut
moss with good drainage. The plants are difficult to estab- Figure 13.106.8. Pyrrosia fallax: a. habit,
lish but otherwise are tough and adaptable to drier condi- bar = 5 cm (2 in.); b. fertile frond with
tions. linear sori, bar = 1 cm (0.4 in.).
Pyrrosia eleagnifolia has entire to dentate rhizome scales,
and the fronds are dimorphic, thick-leathery, and densely
covered with tan hairs on the lower surfaces. Hydathodes
are absent. The sterile fronds are almost rounded, rarely to
3 cm (1.2 in.) long, and the fertile ones are longer and nar-
rower. The distinctly sunken sori are round, slightly spaced,
and appear in two or three rows. Native to New Zealand. a
This species was recently introduced from New Zealand
and has shown promise in warm-temperate areas. Botanists
have confused it with Pyrrosia rupestris (R. Brown) Ching of
Australia and P. serpens (N. L. Forster) Ching of the South b
Pacific. Pyrrosia rupestris differs in having hydathodes. Pyr-
rosia serpens differs in having two rows of sori with the sec-
ond row weakly developed. Pyrrosia serpens is doubtfully
cultivated in the United States.
484 C 13
long, and the fertile ones are longer, narrower, and more
erect. The sori are mostly 1 mm wide, three or four on each
c
side of midrib, scattered, and sometimes fusing. This spe-
cies is native to India, southeastern Asia to Indonesia, and
the Philippines. The forms with longer fronds were for-
d merly called P. obovata.
Figure 13.106.13. Pyrrosia longifolia: a. fronds, bar = Pyrrosia piloselloides (Linnaeus) M. G. Price
13.106.15
5 cm (2 in.); b. sori, bar = 1 cm (0.4 in.), after Hoshizaki
syn. Drymoglossum piloselloides (Linnaeus) C. Presl
(1981); c. rhizome and stipe bases, bar = 1 cm (0.4 in.),
Tender
after Hoshizaki (1981); d. rhizome scales, bar = 1 mm.
A small fern with medium-creeping rhizomes. Grows well
under medium light in moist-dry potting mix or uncut
moss with good drainage. The plants are easy to grow and
Pyrrosia nummulariifolia (Swartz) Ching suitable for terrariums.
13.106.14 The rhizome scales of Pyrrosia piloselloides are dark with
syn. Pyrrosia obovata (Blume) Ching pale margins and bear long hairs. The fronds are dimor-
Tender phic, thick and fleshy, glossy green, and have deciduous
A small fern with medium-creeping rhizomes. Grows well hairs. The sterile fronds are roundish-oblong, sessile or
under medium light in moist-dry potting mix or uncut nearly so, and to 6 cm (2.5 in.) long by 2 cm (0.75 in.)
moss with good drainage. The plants are easy to grow and wide. The fertile fronds are short stalked, linear-oblong,
make an ideal terrarium plant. and to 12 cm (4.75 in.) long by 1 cm (0.5 in.) wide. The
Pyrrosia nummulariifolia has fleshy, dimorphic fronds. sori occur in a marginal band around the apex. This species
The sterile fronds are roundish, mostly to 2 cm (0.75 in.) is native from India to Malaysia and China (Hainan).
F F A C 487
Figure
13.106.16.
a Pyrrosia
polydactyla:
frond, bar =
5 cm (2 in.).
c
b
b
a
Figure 13.106.19. Pyrrosia
rupestris: a. habit, bar = 5 cm
(2 in.); b. fertile frond with
sori, bar = 1 cm (0.4 in.).
After Hoshizaki (1981).
c
with conspicuous hydathodes in a row near the apex. The
fertile fronds are longer and narrower than the sterile ones,
mostly to 8–20 cm (3–8 in.) long. The sori are relatively
Figure 13.106.18. Pyrrosia princeps: a. habit, showing
large (1–2.5 mm wide) and slightly sunken, appearing in
frond with fertile area, bar = 5 cm (2 in.); b. trough-like
one to four rows on each side of the midrib. This species is
rhizome scale with marginal teeth, bar = 0.5 mm; c. stipe
native to Australia.
cross section, with the stippled area showing the dark,
thick-walled cells (sclerenchyma) that strengthen the
stipe, bar = 1 mm. Pyrrosia samarensis (C. Presl) Ching
13.106.20
Tender
A small-medium fern with medium- to longer-creeping
and scattered. The round sori are 0.5 mm wide and densely rhizomes. Grows well under medium-high light in moist-
packed between the main veins in the apical part of blade. dry potting mix or uncut moss with good drainage. The
This species is native to Sulawesi, the Moluccas, and New plants are easy to grow.
Guinea. Pyrrosia samarensis has dimorphic sterile and fertile
Pyrrosia princeps can be confused with P. splendens, a fronds covered with loose white hairs that eventually fall
species that differs by having entire rhizome scales and off. The sterile fronds are narrowly lanceolate and tapered
blade hairs with a distinct, long, spine-like ray. at both ends; the fertile ones are narrowed apically and cov-
ered beneath with roundish sori running together. This
Pyrrosia rupestris (R. Brown) Ching species is native to the Philippines.
13.106.19 Pyrrosia angustata also has loose, deciduous hairs but dif-
Rock felt fern fers by a more erect habit, thicker and grooved stipes, acute
Semi-tender to hardier (?) blade apices, and discrete, sunken sori.
A small fern with medium- to long-creeping rhizomes.
Grows well under medium light in moist-dry potting mix Pyrrosia sheareri (Baker) Ching 13.106.21
or uncut moss with good drainage. Semi-tender or hardier (?)
Pyrrosia rupestris has entire rhizome scales without mar- A medium-sized fern with short-creeping rhizomes. Grows
ginal hairs. The fronds are dimorphic, with the sterile ones well under medium-high light in moist-dry potting mix or
to about 7 cm (3 in.) long, typically angular-rhomboid, uncut moss with good drainage. The plants grow slowly.
F F A C 489
Pyrrosia stigmosa (Swartz) Ching 13.106.23 creeping and bear fronds 1–3 cm (0.5–1.2 in.) apart. At
Tender the top of the stipe are two leaflets that have veins free or
A medium-sized fern with short-medium-creeping rhi- netted at their ends.
zomes. Grows well under medium-high light in moist-dry The genus has only one species, native to southern Bra-
potting mix or uncut moss with good drainage. zil and adjacent Argentina. The genus name honors André
The rhizomes of Pyrrosia stigmosa are about 4.5 mm (0.2 Frederick Regnell (1807–1884), a Swedish botanist who
in.) thick, with the stipes spaced 1–2 cm (0.5–0.75 in.) collected and studied plants in Brazil.
apart. The fronds are 11–48 cm (4.5–19 in.) long by 1.5–
5.2 cm (0.6–2 in.) wide. They are wedge-shaped toward Regnellidium diphyllum Lindman 13.107.1
the base and acuminate to rounded at the apex. The hyda- Very tender
thodes are distinct. The sori are about 0.5 mm wide and A small fern with medium-creeping rhizomes. Grows well
scattered. This species is native to southeastern Asia and under high light in moist-wet soil (a mixture of sand and
Indonesia. peat) or fully submerged. The plants are typically grown in
Pyrrosia stigmosa is similar to P. costata, but the latter pots set in water or in an aquarium with plants partly sub-
species has a stouter rhizome and little or no stipe. merged. Regnellidium diphyllum is easy to grow but apt to
die if the temperature drops below 21°C (70°F). Further
Regnellidium characteristics are described under the genus.
Horticulturists interested in aquatic plants might consider
growing Regnellidium. The genus is related to Marsilea, the Rumohra
clover fern, but differs by having two leaflets instead of four. Leather fern
The leaves are produced too far apart on the rhizome to The genus Rumohra consists of medium-sized, terrestrial or
make an attractive pot plant; the plants are mainly used as epiphytic ferns with broad, dark green, coarsely divided,
a novelty in aquariums. leathery fronds. The durable fronds are used extensively by
Regnellidium roots in mud, although sometimes it is florists in cut flower arrangements. The plants grow in beds
submerged and the fronds are floating. The rhizomes are or pots or baskets and require only moderate watering and
humidity. Over the years, commercial growers, mainly lo-
cated in Florida, have made many selections from their
stock, and today’s leather fern looks quite different from
wild plants, though the wild plants vary considerably as well.
The rhizomes of Rumohra are medium-creeping,
branched, and covered by peltately attached scales. The
cies is native to the southeastern United States, West In- Web site at http://nas.er.usgs.gov/ferns for more informa-
dies, and Central and South America. tion. Moran (1992c) discusses the plant’s weediness and
At different times of the year, cultivated material may the biological control methods that have been used to com-
have many atypical hairs. These hairs might be lacking, rep- bat it.
resented by a bump, or stoutly cone-shaped. The free
branch-hairs borne at the tip of the cone-like hair may be Salvinia natans (Linnaeus) Allioni 13.110.5
poorly developed and number less than four. When sporo- Semi-tender to hardier
carps are not present, Salvinia minima might be confused A small aquatic fern with short-creeping rhizomes. Grows
with S. natans except for the roundish leaves of the former. well under direct sunlight.
Salvinia natans has oblong-truncate to ovate fronds. The
Salvinia molesta D. S. Mitchell 13.110.1, 4 hairs on the upper surfaces of the blades have four free
Semi-tender to tender branches, not united. The sori (sporocarps) are spherical,
A small aquatic fern with short-creeping rhizomes. Grows about six in a tight cluster. The species is native to Europe
well under direct sunlight. and Asia.
Salvinia molesta has rounded fronds, and the mature
ones are often folded lengthwise along the midrib, espe-
cially in crowded conditions. As in S. auriculata, stiff hairs a
b
are present on the upper surfaces of the blades. These hairs
consist of a stalk tipped by four hairs that unite at their c
apices to form an egg-beater- or cage-like structure. The
fertile part of the frond is chain-like (unbranched) and to-
ward the base has short-stalked sori (sporocarps) that are
ovate with a slight beak at the tip. The sori gradually be-
come sessile toward the apex of the fertile part. The spores
are mostly aborted. Salvinia auriculata can be distinguished
from this species by its branched, corymb-like fertile part. Figure 13.110.5. Salvinia natans: a. habit, bar = 1 cm
Salvinia molesta is native to southern South America (0.4 in.); b. hair from upper surface, bar = 1 mm;
(southern Brazil, Uruguay, and Paraguay) but has become c. sporocarps, bar = 1 cm (0.4 in.).
naturalized in Africa, India, Sri Lanka, Indonesia, Australia,
and New Zealand.
This species is a terrible weed in the Old World tropics
and is becoming a serious pest in the southern United Schaffneria
States from Texas to Florida. Although killed by hard frosts
This small, charming fern bears clusters of rounded, dia-
and prolonged subfreezing temperatures, it should not be
mond-shaped fronds. It grows on mossy rocks and ledges
cultivated anywhere in the United States because of its po-
(usually limestone) in tropical areas and is well suited for
tential to escape. In the United States it has been declared
terrarium culture.
a Federal Noxious Weed, which makes it illegal to import,
The rhizomes are erect and bear small, clathrate scales.
own, cultivate, transport, or sell the species. In some states,
The clustered fronds are 2.5–15 cm (1–6 in.) tall, and the
laws also prohibit its possession. Check the World Wide
blades are entire, rhombic-obovate to fan-shaped, glabrous,
and lacking a midrib. The veins spread and divide in a fan-
shaped pattern, forming areoles without included veins.
b c The sori are short to linear, with indusia shaped like the
sori and attached on one side to a vein. Some pteridologists
a classify this genus in Asplenium.
Schaffneria consists of only one species native to Central
America and Cuba. The genus is named for Wilhelm H. J.
Schaffner (1830–1882), a German pharmacist who col-
lected plants in Mexico.
age but kept constantly moist. The plants are easy to grow
from spores.
See the characteristics of the genus for a description.
lateral leaf
median
leaf
axillary stem
leaf
upper surface lower surface
a a b
b b
c
c
a
Figure 13.112.10. Selaginella erythropus: a. branches, bar Figure 13.112.11. Selaginella harrisii: a. branches, bar =
= 5 cm (2 in.); b. leaf arrangement, bar = 2 mm; c. 5 cm (2 in.); b. leaf arrangement, bar = 2 mm; c. leaves,
leaves, lateral (below) and median (above), bar = 1 mm. lateral (below) and median (above), bar = 1 mm.
b
c
Selaginella martensii Spring 13.112.16 Figure 13.112.16. Selaginella martensii: a. branches, bar
Semi-tender to hardier = 5 cm (2 in.); b. leaf arrangement, bar = 2 mm; c.
leaves, lateral (below) and median (above), bar = 1 mm.
A medium-sized plant with erect to suberect, frond-like
growth arising from short-creeping horizontal stems.
Grows well under medium light in moist to moist-wet gar-
den soil or potting mix. This species likes humidity. eral leaves are oblong, to 5 mm (0.2 in.) long. The apex is
Selaginella martensii has leaves that are dark green, rounded, the base unequal, acroscopic side rounded, basis-
smooth, and shiny above but lighter green below. The lat- copic side with a small ear, margins ciliate to denticulate at
the base to more entire toward the apex. The median leaves
are ovate, keeled, and usually long-aristate, the aristae to
b about half the blade length; the base has two uneven ears,
a and the leaf margins are ciliate, becoming more denticu-
late toward the apex, often with a white border. This species
is native to Central America.
‘Albolineata’. Snowy club moss. Leaves variegated with
scattered creamy-white to white streaks.
c
a
a c
b
b
c
a
c
a
b b
c
a
ones turn buff. The lateral leaves are ovate, acute, and cili-
ate to denticulate. The medial leaves are ovate, acuminate at
the apex, ciliate to denticulate, with or without an indis-
tinct white border. The axillary leaves lack ears. This species
is native to Mexico.
Selaginella oaxacana is sometimes misidentified in the
trade as S. pulcherrima.
The lateral leaves of Selaginella serpens are ovate, obtuse, Selaginella uncinata (Desvaux ex Poiret) Spring
cordate at the base, and ciliate (less so toward the apex). 13.112.25; 49
The median leaves are ovate-acuminate, ciliate, and some- Blue spike moss, peacock plant, rainbow moss
times larger or broadly eared on the outer basiscopic side. Hardy, Zone 6
This species is native to the West Indies. A small to medium plant with mat-like growth from long-
creeping or trailing, branched stems. Grows well under me-
Selaginella umbrosa Lemaire ex Hieronymus dium light in moist garden soil or potting mix. The plants
13.112.3, 24 reportedly overwinter well in coastal New York (Zone 6).
Red spike moss This species is attractive for its blue-green, iridescent fo-
Tender liage. This color is best developed when the plants are
A medium-sized plant with frond-like growth arising from grown in low light.
long-creeping, branched horizontal stems. Grows well Selaginella uncinata has ovate or ovate-oblong lateral
under medium light in moist-wet garden soil or potting leaves with transparent and entire margins. Some are
mix. It likes humidity. The species is attractive because of weakly stalked, and the leaf base is truncate to weakly cor-
its red stems. date, with the acroscopic side often overlapping the stem.
The main, erect stems of Selaginella umbrosa are red, The median leaves are overlapping, ovate-lanceolate, the
and the blade-like part is irregularly and densely flabellate, margins transparent and entire, the apex abruptly attenu-
flat, and bright green. The lateral leaves of the main ate, and the base often oblique and weakly cordate. This
branches are ovate-lanceolate, unequally sided, and strongly species is native to southern China and has become natu-
ciliate at the base. The median leaves are ovate-acuminate ralized in the southeastern United States.
and slightly keeled. This species is native to the American
tropics and has become naturalized in many parts of the Selaginella viticulosa Klotzsch 13.112.26
tropics throughout the world. Tender
A small-medium plant with frond-like growth arising from
creeping horizontal stems. Grows well under medium light
a in moist potting soil and in humidity. This species has
softer and laxer foliage than other selaginellas.
c
b b
The lateral leaves on the distal half of the main stem are Selaginella willdenovii (Desvaux ex Poiret) Baker
oblong, two times longer than wide, spreading (not imbri- 13.112.28
cate), and faintly streaked. The acroscopic margin is ciliate Peacock fern, vine spike moss
toward the base to denticulate toward the apex, and the Tender, Zone 9
basiscopic base is truncate. Selaginella viticulosa is native to A medium to large plant with sprawling to erect, frond-like
Costa Rica, Panama, Colombia, and Venezuela. growth. Grows well under medium-high light in moist gar-
den soil. Give the plants a lot of space to grow. The scram-
Selaginella vogelii Spring 13.112.27 bling stems may reach 6 m (20 ft.) long and typically bear
Tender small leaves with a strikingly beautiful sky-blue iridescence
A small plant with frond-like growth arising from short- on their upper surface. The iridescence, however, develops
creeping horizontal stems. Grows well under medium light only on plants grown in shade; those plants grown in full or
in moist garden soil or potting mix. This species likes partial sun are not iridescent. Iridescence in pteridophytes
humidity. tends to occur in species that grow on deep-shaded forest
The main, erect stems of Selaginella vogelii are about as floors, which suggests that this trait is an adaptation to low
long as the blade-like part. The latter is broadly triangular light. See Moran (1995a) for a popular account of irides-
and bears branches with many short, stiff hairs on the lower cence in ferns.
surface. The lateral leaves are narrowly triangular, con- All the leaves are entire on Selaginella willdenovii. The
stricted near the base then decurrent, and entire (not cili- lateral leaves are whitish eared on the acroscopic side, and
ate). The median leaves are long-acuminate, unequal at the the axillary ones have two elongate-rounded ears. This spe-
base, and entire. This species is native to tropical Africa. cies is native to tropical Asia and Indonesia and has occa-
sionally become naturalized elsewhere in the tropics.
F F A C 507
Solanopteris
Potato fern Figure 13.114.1. Solanopteris
Comprising small epiphytes, the genus Solanopteris is bifrons: habit, bar = 5 cm (2 in.).
grown as a novelty for its globose tubers (modified stems),
which in nature are inhabited by ants. Growers report that
the plants often do well for a while then suddenly die, al-
though plants at the Marie Selby Botanical Garden in
Florida have been flourishing for a long time. Solanopteris
grows high in trees in its native, wet-forest habitat.
This genus and Lecanopteris are the only ferns that have
stems inhabited by ants. The ants fiercely attack anything
that disturbs the plant. In Solanopteris, they eventually fill
the tuber with organic debris and frass, at which point the
ants seek another tuber in which to live. The plant’s roots
then grow inside the tuber and extract water and mineral
nutrients from the organic matter left behind by the ants.
Thus, the plant receives two benefits from the ants’ activi-
ties: protection and nutrients. The plants seem to grow bet-
ter when inhabited by ants; however, the tubers will form
even if ants are absent. For more about the natural history
of this fern, see Moran (1992b).
In addition to the tubers, Solanopteris is characterized
by long-creeping rhizomes covered by minute, circular
scales. The sterile and fertile fronds are dimorphic, with the
sterile fronds entire to pinnately lobed, and the fertile ones
longer and narrower. The sori vary from round to elongate
and, like in all polypodiaceous ferns, they lack indusia.
Some pteridologists consider Solanopteris close enough
to Microgramma to be placed in that genus. It is kept dis-
tinct here because the name has been used in many floras
and is well established in horticulture.
The genus is native to the American tropics and con-
sists of four species. The name is derived from the potato
genus, Solanum, and the Greek pteris, fern, in reference to
the potato-like tubers.
Figure 13.115.5.
Sphaeropteris
cooperi: tip of
stipe scale,
not bristle tipped,
bar = 1 mm.
comes from the Greek sphaera, sphere, and pteris, fern, al-
luding to the globose sori.
c b
a c b
a
a b
b
b
Figure 13.117.5.
Figure 13.117.4. Tectaria decurrens: a. frond, bar = 8 cm a Tectaria gemmifera:
(3 in.); b. sori and indusia, bar = 5 mm (0.2 in.). a. frond, bar = 8 cm
(3 in.); b. bud, bar =
2 cm (0.8 in.).
the sterile. The stipes are elongate and the blades greatly
contracted, with the sporangia acrostichoid, spreading
b across the lower surface and lacking indusia. This species is
native to Sri Lanka, India, southern China, Vietnam, and
c Taiwan.
Figure 13.117.9.
Tectaria vivipara:
a. frond, bar = 8 cm
(3 in.); b. bud at base
a
of pinna, bar = 5 cm
(2 in.); c. indusium,
bar = 5 mm (0.2 in.).
apices, elongate pinnae, and veins of each segment united narrowed frond apices and irregularly crimped hairs on the
with the vein of an adjacent segment and forming an ex- lower pinna surfaces.
current vein running to the sinus. Native to eastern Asia.
Thelypteris dentata (Forsskål) E. St. John
Thelypteris augescens (Link) Munz & I. M. Johnston 13.118.5
13.118.4 syn. Christella dentata (Forsskål) Brownsey & Jermy,
Abrupt-tipped maiden fern Cyclosorus dentatus (Forsskål) Ching
Tender, Zone 10 Downy thelypteris
A large fern, up to 120 cm (4 ft.) tall, with medium-creep- Semi-tender, Zone (7)8
ing rhizomes. Grows well under high light in moist, basic A medium-sized fern with short-creeping to suberect rhi-
garden soil or potting mix. zomes. Grows well under medium light in moist garden
The abruptly narrowed frond tip and long, narrow pin- soil or potting mix. The plants are easy to grow but do
nae are distinct features of Thelypteris augescens. The apical poorly in areas with cool nights. They often become weedy
portions of the leathery blades are about five times longer in greenhouses.
than they are broad, and the lateral pinnae are less than 1.5 Thelypteris dentata can be identified by the combination
cm (0.6 in.) wide. On the undersurfaces of the pinnae the of one-pinnate-pinnatifid blades, basal pinnae that are usu-
costae, costules, and veins are sparsely covered with hairs ally reduced and bent forward and downward, and basal
and scales. This species is native to Florida, Cuba, the Ba- veins that unite and form an excurrent vein that runs to the
hamas, and Central America. base of the sinus. Other helpful characteristics are the often
Much of the cultivated material by this name is actually
Thelypteris puberula, which differs by having less abruptly
b
Figure 13.118.5.
Thelypteris dentata:
a. habit, bar = 5 cm
(2 in.); b. segments,
bar = 1 mm.
a
b
Figure 13.118.4. Thelypteris augescens: a. frond, bar =
5 cm (2 in.); b. segments, bar = 1 mm.
520 C 13
Figure 13.118.8.
Thelypteris limbosperma:
a. habit, bar = 5 cm
(2 in.); b. segments, bar
= 1 mm.
Figure 13.118.9. Thelypteris
nevadensis: a. habit, bar =
5 cm (2 in.); b. segments
dotted with glands, bar =
1 mm.
a
a
moist garden soil or potting mix. The plants are easy to
b grow and may become invasive, tending to spread irregu-
larly throughout the garden if not contained. They are not
adapted to subtropical climates.
Thelypteris noveboracensis is characterized by one-pin-
nate-pinnatifid blades gradually tapered toward the base,
with the lowest pinnae appearing as mere stubs or nubbins.
The rhizomes are long-creeping, and the blades either lack apex and widest (or nearly so) at the base, and fertile pinnae
or are sparsely beset with sessile glands. The costae are con- narrower and more contracted than the sterile ones. The
spicuously hairy on the lower surface. The veins are un- veins are free, once-forked, and meet the margin above the
branched and meet the margin above the sinus. This species sinus. The lower surfaces of the costae bear scattered scales.
is native to eastern North America. The species is native to North America, Mexico, the West
Indies, and Europe.
Thelypteris ovata R. St. John 13.118.11 A crested variant (forma pufferae A. A. Eaton) is re-
Semi-hardy, Zone 8 ported in cultivation.
A medium-sized fern with medium-creeping rhizomes.
Grows well under medium light in moist to moist-dry gar- b
den soil or potting mix. The plants are easy to grow.
Thelypteris ovata is characterized by blade apices less
than five times long as wide, more or less attenuate and dif-
ferent from the lateral pinnae, and sessile pinnae with the
basal segment slightly longer than the other segments and
often toothed. Unlike in similar species, the costules and
veins on the upper surface are glabrous and the sori are sub-
marginal. The species is native to the southeastern United
States, Central America, and the West Indies.
b c
Figure 13.118.11.
Thelypteris ovata: a.
habit, bar = 5 cm Thelypteris parasitica (Linnaeus) Fosberg
a 13.118.13
(2 in.); b. segments,
with elongate basal syn. Christella parasitica (Linnaeus) H. Léveillé,
segments, bar = Cyclosorus parasiticus (Linnaeus) Farwell
1 cm (0.4 in.). Semi-tender to slightly hardier
A medium-sized fern with medium-creeping rhizomes.
Grows well under medium light in moist garden soil or pot-
ting mix. The plants volunteer readily from spores.
In Thelypteris parasitica, the basal veins on a segment
unite with those of adjacent segments to form an excurrent
Thelypteris palustris Schott 13.118.12 vein that runs to the base of the sinus. Both surfaces of the
syn. Thelypteris thelypteroides (Michaux) Holub blade–particularly the veins on the lower one–have hairs
Marsh fern and stalked, orange glands. The indusia are long-ciliate.
Very hardy, Zone (2)3 The species is native to tropical Asia and the islands of the
A medium-sized fern with long-creeping rhizomes and de- Pacific.
ciduous fronds. Grows well under medium-high light in
moist-wet garden soil or potting mix.
Thelypteris palustris can be identified by long-creeping
rhizomes, one-pinnate-pinnatifid blades tapered toward the
F F A C 523
b
Figure 13.118.13.
Thelypteris
parasitica: a.
habit, bar =
5 cm (2 in.); b.
segments with
minute glands on
segment midrib,
bar = 1 mm.
a
c
Figure 13.118.14.
Thelypteris patens: a. habit,
bar = 5 cm (2 in.); b. pinna
base, bar = 1 cm (0.4 in.);
b c. stipe scales, bar = 1 cm
(0.4 in.).
two to five lateral ones. The pinnae are elliptic with suben-
tire to coarsely serrate-crenate margins. Minute, forked, or
star-shaped hairs are present only on the rachis near the
junctions with the costae. The sori lack indusia and form a
double row between the main lateral veins. This species is
native to, and widespread in, the American tropics.
Figure 13.118.17.
Thelypteris puberula (Baker) C. V. Morton Thelypteris reptans:
13.118.1, 2, 16 fronds, bar = 5 cm
Semi-tender, Zone (8)9 (2 in.).
A medium-sized fern with medium- to long-creeping rhi-
zomes. Grows well under high-medium light in moist-dry
garden soil or potting mix.
The blades of Thelypteris puberula are broadest at or near
the base, gradually or somewhat abruptly tapered toward
the apex, and glabrous on the upper surface except for
crimped hairs on the costae. The basal veins of each seg-
ment end at or above the sinus. The indusia are densely
hairy. This species is native to the southwestern United
States, Mexico, and Central America.
a
Figure 13.118.18. Thelypteris resinifera: a. habit, bar =
5 cm (2 in.); b. segments dotted with glands, bar =
1 mm.
Todea
a
A handsome accent or pot plant, Todea forms a large, erect
funnel of bright green foliage. In their native habitat these
terrestrial ferns can become large plants with stout trunks to
1 m (3 ft.) tall and fronds to 1.2 m (4 ft.) long. The trunks
b
Figure 13.118.21.
Thelypteris serrata:
a. frond, bar = 5 cm b
(2 in.); b. margin and
veins of pinna (young
plant), bar = 1 mm.
F F A C 527
Trachypteris
This genus of small, terrestrial or rock-inhabiting ferns has
short-creeping rhizomes, densely scaly leaves (at least on
the undersurface), and netted veins. The sori lack an indu-
sium. The genus name comes from the Greek word trachys,
rough, and pteris, fern, referring to the rough texture im-
parted by the scales to the fronds.
The fronds of Trichomanes auriculatum are sessile or Figure 13.121.3. Trichomanes holopterum: a. frond, bar =
have stipes to 4 mm (0.2 in.) long. The blades are linear-ob- 5 cm (2 in.); b. indusium and sorus with extended
long, 10–40 cm (4–16 in.) long, and one-pinnate, with the receptacle, bar = 1 mm.
sterile pinnae lobed and crenate; the fertile pinnae are
deeply and sharply incised. This species is native from India
to southeastern Asia, Japan, Malaysia to Indonesia and
Micronesia.
Figure 13.122.4.
Vittaria lineata:
a. habit, bar = 5 cm
(2 in.); b. sori, bar =
3 mm (0.1 in.).
a
b
Figure 13.123.8.
Figure 13.123.7. Woodsia Woodsia oregana:
obtusa: a. frond, bar = 2.5 cm a. frond, bar =
(1 in.); b. sorus and indusium, 2.5 cm (1 in.);
bar = 1 mm. b. sorus and
indusium, bar =
1 mm.
a
a
b
b
Figure 13.124.5.
a Woodwardia fimbriata:
a. frond, bar = 10 cm
(4 in.); b. lowest pinna,
Figure 13.124.4. bar = 5 cm (2 in.).
Woodwardia areolata:
a. habit with fertile
frond (right), sterile
frond (left), bar =
10 cm (4 in.); b. b
fertile pinnae, bar =
1 cm (0.4 in.).
b
Figure 13.124.6. Woodwardia orientalis: a. frond, bar =
10 cm (4 in.); b. lowest pinna, bar = 5 cm (2 in.); c. leaf
buds, bar = 2 cm (0.8 in.). Figure 13.124.7. Woodwardia radicans: a. frond (left)
and leaf bud (right), bar = 10 cm (4 in.); b. lowest
pinna, bar = 5 cm (2 in.).
Woodwardia ×semicordata Mickel & Beitel
13.124.8
Semi-tender, Zone 9 more pairs of decurrent pinnae. Woodwardia martinezii
The general culture of this fern is similar to that for Wood- Maxon ex Weatherby, native to Mexico, is unknown in
wardia spinulosa except that it might be more cold toler- cultivation. It differs from W. ×semicordata by long-creep-
ant. According to Mickel and Beitel (1988), W. ×semicor- ing rhizomes and sori along the costules and entire length of
data resulted from a cross between W. martinezii and W. the costae and rachis. Woodwardia ×semicordata is native
spinulosa, but further research is needed to determine to Mexico.
whether it is actually of hybrid origin. At least some spores
appear to be normal and well formed, and the plants might Woodwardia spinulosa M. Martens & Galeotti
be apogamous. 13.124.9
In cultivation, Woodwardia ×semicordata has been con- Mexican chain fern
fused with both of its putative parents. It resembles W. spin- Semi-tender, Zone (9)10
ulosa by having sori distributed along the costules (the mid- A large fern with short-creeping rhizomes and evergreen
ribs of the pinnules) but differs by having sori also fronds. Grows well under high light in moist garden soil.
distributed in the distal half or less of the costae (the mid- The plants are easy to grow.
ribs of the pinnae). Beneath the pinnatifid apex, only one Woodwardia spinulosa can be distinguished from W.
pair of free pinnae with decurrent bases is present (all other fimbriata by the triangular blades with fimbriate scales on
pinnae are nondecurrent), whereas W. spinulosa has two or the lower surfaces. The lower pinnae have basal segments
F F A C 539
a
b
b
Woodwardia virginica (Linnaeus) Smith
13.124.11
syn. Anchistea virginica (Linnaeus) C. Presl
Figure 13.124.9. Woodwardia spinulosa: a. small frond, Virginia chain fern
bar = 10 cm (4 in.); b. lowest pinnae with basal pinnules Very hardy, Zone 3
overlaying the rachis, bar = 5 cm (2 in.); c. sori along A large, deciduous fern with long-creeping rhizomes up to
costules only, bar = 2.5 cm (1 in.). 1 cm (0.5 in.) thick. Grows in medium to high light in wet,
acidic garden soil. This species is considered invasive in
some areas.
540 C 13
Figure 13.124.11.
Woodwardia virginica:
a. habit, bar = 10 cm (4 in.);
b. pinnule with sori and
a indusia, bar = 4 mm.
The human eye often has difficulty judging light intensity. For this reason, special light meters can be pur-
chased to measure light intensity in foot-candle units. Photographic light meters can also be used, and the
readings can be converted to foot-candle units. The formula is as follows:
foot-candles = 20f 2 / TS
where f is the aperture in f-stop, T the shutter speed in seconds, and S the film speed in ASA units.
To measure light intensity using a photographic light meter, place a large sheet of white paper on the
surface to be measured. Set an appropriate ASA film speed on the meter and read the shutter speed required
for proper exposure at a given f-stop. Read the meter at about 30 cm (12 in.) from the paper, and use the
preceding formula to calculate the equivalent foot-candle measurement. For example, if the ASA film speed
setting is set at 100 and the proper exposure is ½ second at an f-stop of 16, the foot-candle value is 102:
20 × (16)2 / ½ × 100 = 102 foot-candles
This formula provides a workable approximation. Results will vary according to the accuracy of the light
meter and the cone of light measured by the meter.
For scientific work, precise light intensities need to be measured. A quantum meter accurately measures
the blue and red part of the spectrum (the photosynthetically active radiation, or PAR). Plants use the blue
and red light but not yellow and green light. With quantum meters, the intensity is measured in units
known as micromoles, instead of foot-candles. A reading of 200 micromoles (µmol m−2 sec−1) is equivalent
to about 1000 foot-candles. Quantum meters are specifically designed for plants, but the average fern
grower will find photometers sufficiently accurate for monitoring the light needs of ferns.
Other light-measuring units that may be encountered, and their approximate equivalency to foot-can-
dles, are as follows:
1 lumen/cm2 = 1000 foot-candles
1 lux = 1⁄ 10 foot-candle
1 micromole = 5 foot-candles
541
Fern Societies
Updated information and addresses for various fern societies in the United States and abroad can be
obtained from the American Fern Society. Contact the secretary of the American Fern Society:
Dr. W. Carl Taylor
Botany Department
Milwaukee Public Museum
800 W. Wells Street
Milwaukee, Wisconsin 53233-1478
The information is also available from the American Fern Society’s World Wide Web page:
http://www.amerfernsoc.org
Membership information for the American Fern Society can be obtained from the membership secretary:
Dr. David B. Lellinger
Botany Department, MRC-166
Smithsonian Institution
Washington, D.C. 20560
A highly informative Web site about ferns is maintained by the San Diego Fern Society. It has links to nearly
all other fern Web sites and is an excellent way to search for current information about the fern society near-
est you.
http://www.inetworld.net/sdfern/society.htm
Another helpful Web site for growers is that maintained by the British Pteridological Society.
http://www.nhm.ac.uk/hosted_sites/bps/
An e-mail exchange for fern information is available and called Fernet. Inquire about the service at
[email protected]
542
Importing Ferns
The following information will be helpful for importing plants to the United States and knowing what to
expect as your plants pass through customs. When plants are carried through as personal luggage and they
comply with all regulations, a United States Department of Agriculture (USDA) Plant Import Permit is not
required for one-time or infrequent importers (though see the regulations and restrictions applicable to
ferns). Frequent importers need a plant-import permit. This permit also allows you to send plants to the
United States by mail or separate carrier to an inspection station. If the plants pass the inspection, they are
forwarded to you.
You must pay attention to the regulations and directions that come with the import permit. These rules
are not given here, except in a general way applicable to ferns. Because regulations and procedures are sub-
ject to change, always check with the agencies involved for the latest information.
543
544 A III
or export requirements. Also check with the CITES office for updated information and the address of the
CITES office abroad. Once at your destination, the local customs or plant protection or regulating agency
should be able to inform you of their regulations.
mit separately from non-CITES plants. The plant inspector will routinely send any CITES plants to the
inspection station before release. Before relinquishing the plant, be certain that the CITES permit and the
plant are securely together, preferably in their own bag. Also make sure that the customs invoice (Post-entry
papers or Manifest) indicates that the plant passed customs. In addition to the usual USDA plant import
requirements, tree-fern trunks may not be more than 45 cm (18 in.) long measured from the soil level to
the tip of the stem (apical bud). The latest CITES regulations indicate that all tree ferns on the CITES list,
with or without trunks, are subject to the CITES regulations (formerly, those without trunks were exempt).
Avoid traveling to a second country with a CITES plant even if you have an export permit issued from
the first country you visited. You will need to get a second export permit (re-export) from the second coun-
try, and this can take several days. It may be better to change your itinerary or mail the ferns along with their
CITES export permit from the original country directly to the United States. CITES regulations specify
that CITES export permits must come from the country of origin, and that may be interpreted to mean the
last country visited. That last country may refuse to issue an export permit.
The preceding procedures and restrictions are meant to protect the United States from introduced
pests and diseases, and to prevent the extinction of threatened or endangered species. The restrictions may
be inconvenient and complicated, but they are imposed for the well being of agriculture, the environment,
and ultimately all of us.
Names of Pest and Disease
Control Substances
The active ingredients of common pest and disease control products are given in the following charts. The
trade names appear in upper case. Some products are available to commercial applicators only. Formula-
tions vary, and no attempt has been made to account for these variations. The Federal and State Environ-
mental Protection Agencies (EPA) regulate the label content and use of pesticides, fungicides, bactericides,
disinfectants, and other substances. Their directions are the law. EPA regulations may change, and new
products continue to be produced, so check with county agricultural services or other reputable sources for
the latest information. Always test a product on one plant before wider application. The reader acts on his
or her own responsibility. No endorsement is intended in the use of trade names, nor is criticism implied of simi-
lar products not mentioned.
Pesticides
Alphabetical listing Corresponding names Applications
547
548 A IV
553
554 A V
Nephrolepidiaceae
Nephrolepis
Horsetails
Isoetaceae
Selaginellaceae Lycopods
Lycopodiaceae
An evolutionary tree (cladogram) showing the current ideas about the relationships of major groups of vascular plants.
The lycopod line of evolution has been distinct from other plants since the early Devonian, about 395 million years
ago. It is based on evidence from morphology, anatomy, and DNA sequences of living and fossil plants.
F C F G T T 557
Grammitidaceae
Polypodiaceae
Polypodiaceae (Loxogramme)
Davalliaceae
Nephrolepidaceae
Lomariopsidaceae
Dryopteridaceae
Blechnaceae
Paleocene or more recent Thelypteridaceae
Aspleniaceae
Eocene
Pteridaceae
Pteridaceae (Adiantum)
Paleocene Vittariaceae
Dennstaedtiaceae
Dennstaedtiaceae (Lonchitis)
Lindsaeaceae
Triassic
Dicksoniaceae
Early Cretaceous Metaxyaceae
Cyatheaceae
Plagiogyriaceae
Late Cretaceous
Salviniaceae (Azolla)
Late Cretaceous Salviniaceae (Salvinia)
Middle Jurassic Marsileaceae
Permian Schizaeaceae
Gleicheniaceae
Permian Hymenophyllaceae
Osmundaceae
Ophioglossaceae
Marattiaceae
Late Carboniferous
Cycadaceae
An evolutionary tree (cladogram) showing the current ideas about the relationships of major families of ferns. The tree
is based on a combined analysis of data from morphology and DNA and is modified from Pryer et al. (1995). The
geologic periods given for some of the branches of the tree are times of earliest origin based on fossil evidence
(Collinson 1996).
Glossary
acroscopic: the side facing the apex. The acroscopic margin of a pinna is that which faces the apex of the leaf.
The acroscopic margin of a pinnule is that which faces the apex of the pinna. Compare to basiscopic.
acrostichoid: with the sporangia spread throughout the surface, not in discrete lines or dots.
acuminate: tapering to a long point with the margins pinched or concave just before the tip.
acute: short-tapering to a point, the margins not pinched or concave just before the tip.
adnate: grown to or united with an unlike part.
aerophore: the aerating tissue of the leaf, usually in the form of a light-colored line running lengthwise along
the leaf stalk and sometimes down into the stem. In genera such as Blechnum and Thelypteris, aerophores
may occur as short white or yellowish pegs at the bases of the pinnae where they join the rachis. Stomata
are abundant in aerophores, and thus these structures allow air to diffuse into the leaf. Aerophores are
characteristic of fern leaves, being absent from leaves of other plants.
anadromic (anadromous): with the first basal branch or vein (as on a pinna) arising from the side toward
the frond tip. Compare to catadromic.
anastomosed: said of veins that unite to form areoles; net-veined.
annulus: a complete or partial ring or cluster of thick-walled cells on the spore case functioning to open the
spore case.
antheridium: the male sex organ containing the sperm. Borne on the underside of the prothallus.
apogamy: a form of asexual reproduction in which new sporophytes are produced directly from the pro-
thallus tissue instead of from the fertilized egg cell (zygote).
apospory: a form of asexual reproduction in which prothalli are produced directly from young sporophyte
tissue instead of from spores.
appressed: pressed close to the surface, not spreading.
archegonium: the female sex organ containing the egg.
areolate: having netted veins.
areole: the area enclosed by netted veins.
aristate: having stiff, bristle-like appendages.
articulate: having nodes or joints that naturally separate at maturity; jointed.
attenuate: gradually tapering to a very narrow, slender point.
axes: a collective term referring to the petiole, rachis, costae, costules, and midrib of a leaf.
basiscopic: the side facing the base. The basiscopic margin of a pinna is that which faces the base of the leaf.
The basiscopic margin of a pinnule is that which faces the base of the pinna. Compare to acroscopic.
bicolorous: having two colors, usually referring to scales in which the central part is darker than the mar-
gins. Compare to concolorous.
558
anadromous veins
VEINS
FROND
catadromous veins
distal portion
of blade
PINNAE
medial
portion of
blade
anadromous pinna
acroscopic side
of pinna
basiscopic side
proximal of pinna catadromous
portion of pinna
blade
PINNULE
acroscopic side
of pinnule
Location of plant parts. Courtesy of Carl Taylor, Paul Nelson, and the Milwaukee Public Museum.
560 G
Leaf shapes (left to right): linear, lanceolate, ovate, obovate, cuneate, triangular, pentagonal.
binomial: the species name, which is the two words consisting of the genus name and the specific epithet.
bipinnate: twice pinnate (two-pinnate). Both primary and secondary divisions of the frond are pinnate.
bipinnatifid: twice pinnatifid. The divisions of a pinnatifid frond are again pinnatifid.
blade: the thin, broad part of a leaf or frond. Also called the lamina.
bud: in ferns the term usually applies to a proliferous bud or lump of tissue that grows into a new fern plant.
bulbil or bulblet: a small bulb-like body borne upon a stem or leaf and serving to vegetatively reproduce the
plant.
bullate: puffy or inflated, humped. Usually refers to the base of a scale, especially in tree ferns.
catadromic (catadromous): with the first basal branch or vein (as on a pinna) arising from the side toward
the frond base. Compare to anadromic.
caudate: having a slender, tail-like appendage.
ciliate: having short hairs (cilia) along the margin.
clathrate: refers to scales composed of cells that have dark side cell walls and broad, clear central portions
(the “body” of the cell); thus the scale resembles leaded glass. A hand lens or microscope is needed to
see this characteristic.
compound: said of a leaf having two or more leaflets.
concolorous: having one uniform color throughout, usually referring to scales. Compare to bicolorous.
cone: a tight cluster of highly modified, spore-bearing leaves borne at the branch tips.
confluent: running together.
cordate: heart-shaped.
costa (plural, costae): the midrib of a simple frond, or of the pinna (secondary rachis) or pinnules of a
compound frond.
costule: the midrib of a pinnule.
crenate: having low, rounded teeth.
crested: having forked tips, usually many. Usually refers to the frond, pinnae, or segments.
crown: the tip of the stem where the leaves arise. Usually this term is applied to thick, upright stems.
crozier: a young coiled fern frond; the fiddlehead.
cultivar: a cultivated variety of plant, originating in cultivation or selected from among wild plants, gener-
ally chosen for a desirable trait(s).
decurrent: having the base of a blade or blade division extending down the petiole, rachis, or costa as a
winged expansion or ridge. A decurrent base is winged downward, at least for a short distance.
deltate: triangular.
dentate: toothed, usually with broad teeth directed outward.
denticulate: minutely dentate.
dichotomous: forked regularly into pairs.
G 561
dimorphic: having two forms. In ferns the term usually refers to differences in size or shape of the sterile and
fertile leaves (or segments). Compare to monomorphic.
distal: the part farthest away from the main body or leaf axis. Compare to proximal.
dorsal: relating to the back or lower side of a leaf.
emarginate: having a shallow notch in the margin.
entire: having an unmodified margin (not divided, lobed, or toothed).
epilithic: growing on rocks; also epipetric, saxicolous, or rupestral.
epiphyte (epiphytic): a plant that grows upon another plant. An epiphyte uses its host plant only for sup-
port; it is not parasitic.
erose: appearing as if gnawed. Usually refers to an irregular margin.
excurrent vein: a vein running toward the margin, not the midrib.
falcate: scythe-shaped, curved and flat, tapering gradually.
false indusium: an indusium formed by the enrolled margin of the leaf. Characteristic of many genera in
the Pteridaceae, such as Adiantum, Cheilanthes, and Pellaea.
false peltate: appearing peltate but with a narrow, inconspicuous sinus running to the centrally attached
stalk; also referred to as pseudopeltate. See peltate.
family: a group of related genera. Plant family names end with the suffix -aceae.
farina: a meal-like powder, usually white or yellow and found on the lower surfaces of fronds. Character-
istic of such fern genera as Argyrochosma, Pentagramma, and Pityrogramma.
farinose: covered with a meal-like powder (farina).
fern allies: vascular plants that reproduce by spores and have a life cycle similar to that of true ferns, but dif-
fer by how they bear their sporangia and by having small leaves and usually simple, unbranched veins.
Most of the fern allies are not closely related to the true ferns. The living families of fern allies are the
Equisetaceae, Isoëtaceae, Lycopodiaceae, Psilotaceae, and Selaginellaceae.
fertile: in ferns usually referring to leaves that bear sori.
fiddlehead: a fern leaf, young or in bud, that is coiled in a spiral pattern. Also see crozier.
fimbriate: having a fringed margin, the fringe often consisting of numerous hairs or narrow segments.
flora: a list of all the species growing in a region, or a collective term for all the species growing in a region.
It also refers to books that identify the plants within a certain geographical area.
foot-candle: a unit of measuring light intensity. One foot-candle is equal to the amount of light cast by a
standard candle one foot away from the flame.
friable: describing soil that is moist and loose.
frond: the leaf of a fern, typically consisting of the petiole and blade.
fused: grown to or united with a similar part.
gametophyte: a small, usually flat plant bearing the sex organs (archegonia and antheridia) that in turn pro-
duce the gametes. Each cell in the body of the gametophyte has one set of chromosomes (1n). Game-
tophytes grow from spores.
gene: a unit on a chromosome that determines the inheritance of a particular trait.
genus: a group of related species.
glabrous: lacking hairs or scales, smooth.
glaucous: having a whitish, waxy bloom.
hardened plants: plants that have adjusted physiologically to harsher growing conditions.
hastate: arrowhead-shaped, but with the basal lobes pointing outward at wide angles.
heterosporous: having two kinds of spores, male and female, the males being smaller than the females and
produced in separate sporangia.
homosporous: having one kind of spore.
hybrid: an offspring of two different species.
562 G
hydathode: an enlarged vein tip on the upper surface of a blade. It often secretes water and minerals. In some
ferns, such as Nephrolepis, the minerals may accumulate as a white deposit over the hydathode.
imbricate: overlapping.
immersed: sunken into the surrounding tissue.
incised: cut sharply and irregularly, more or less deeply.
included veinlet: a vein within an areole or mesh formed by other veins.
indusium: the structure covering the sorus. Also see false indusium.
internode: the portion of a stem between two successive nodes.
jointed: able to separate naturally at a certain point, leaving a scar; articulate.
laciniate: slashed or cut into narrow-pointed lobes.
lanceolate: lance-shaped.
leaflet: one of the divisions in a compound leaf.
linear: long and narrow, usually 10 times longer than wide, the sides parallel or nearly so.
lunate: shaped like a crescent or half-moon.
marcescent: withering (senescing) without falling off; in contrast to jointed or articulate, which indicate
withering and falling off at a joint.
marginal: at or relating to the margin or edge of a leaf.
meiosis: the type of cell division that gives rise to spores. During meiosis, the cell replicates its chromosomes
once and divides twice; the result is four cells with only half the chromosome number of the original
cell.
mesh: refers to netted veins.
monomorphic: having only one form. In ferns the term usually refers to sterile and fertile leaves that are not
differentiated. Compare to dimorphic.
mucronate: tipped with a short, abrupt point or mucro.
mutation: a sudden heritable change appearing in animals or plants due to changes in the genes or chro-
mosomes.
node: the point on a stem where a leaf emerges.
nonclathrate: refers to scales in which the cells lack dark, easily visible cell walls outlining the body of the
cell. Compare to clathrate.
oblanceolate: lanceolate with the broadest part near the tip.
obovate: ovate or egg-shaped with the broader end apical.
obtuse: blunt or rounded at the tip.
ovate: egg-shaped in outline.
palmate: having main veins, segments, or lobes radiating from a common point.
paraphysis (plural, paraphyses): a sterile hair that is mixed among the sporangia.
pedate: palmate with the side lobes cleft into two or more segments.
peltate: having the stalk attached centrally rather than at the edge (like the handle on an open umbrella).
See false peltate.
petiole: a leaf stalk; also known as a stipe.
phyllopodium (plural, phyllopodia): a stump-like extension from the rhizome to which the fronds are
attached, usually by a distinct abscission layer.
pinna (plural, pinnae): the primary division of a pinnately divided frond; a leaflet.
pinnate: having a feather-like arrangement, with a single mid-vein from which leaflets arise.
pinnate-pinnatifid: referring to a blade that is once-pinnate and with the pinnae deeply lobed or cut, but
not to their midrib.
pinnatifid: cut half to three-fourths to the rachis.
pinnatisect: cut almost all the way to the rachis.
G 563
stellate: star-like; having arms or hairs that radiate from a central point. Usually refers to hairs or scales.
sterile: refers to leaves that do not produce sori and to hybrids in which spores are aborted.
stipe: the leaf stalk or petiole.
stipule: a basal appendage of a stipe or petiole, usually two. In ferns the term is applied to the Marattiaceae
and sometimes to the flared leaf bases in Osmunda.
stolon: a long, slender stem capable of producing a new plant at its tip or along its length.
strigose: having curved, sharp, forward-pointing hairs.
synonym: an alternative scientific name, but not the one currently accepted by taxonomists.
terrestrial: growing in the ground, not on trees.
tomentum: a dense covering of woolly hair.
ultimate segment: the final division of a frond.
undulate: having a wavy surface or margin.
vascular: pertaining to specialized tissue (xylem and phloem) that conducts water, mineral nutrients, and
sugars.
vascular bundle: a bundle or strand of vascular tissue.
veins: the strands of vascular tissue in a leaf. “Free veins” are those unconnected to other veins; “netted
veins” are those so connected.
wing: a thin expansion or flat extension of an organ or structure.
xerophyte: a plant adapted to dry habitats.
zygote: a fertilized egg cell. The first cell in the development of a sporophyte.
Literature Cited
ABRS/CSIRO Australia. 1998. Flora of Australia. Vol. 48, Ferns, Gymnosperms and Allied Groups.
Melbourne.
Andrews, S. B. 1990. Ferns of Queensland: A handbook to the ferns and fern allies. Queensland Depart-
ment of Primary Industries Information series Z189008. Queensland Herbarium, Brisbane.
Anonymous. 1986. Guidelines for the control of plant diseases and nematodes. Agricultural Handbook No.
656. United States Department of Agriculture, Washington, D.C.
Basile, D. A. 1973. A simple method of initiating axenic cultures of pteridophytes from spores. American
Fern Journal 63: 147–151.
Beddome, R. H. 1864. Ferns of Southern India, Being Descriptions and Plates of the Ferns of the Madras
Presidency. Madras.
Beitel, J. M. 1979. The clubmosses Lycopodium sitchense and L. sabinaefolium in the upper Great Lakes area.
Michigan Botanist 18: 3–13.
Beitel, J. M., W. H. Wagner Jr., and K. S. Walter. 1981. Unusual frond development in sensitive fern Ono-
clea sensibilis L. American Midland Naturalist 105: 396–400.
Best, R. 1980? Growing Ferns. Bay Books Australian Gardening Library, Bay Books, Sydney.
Boodley, J. W. 1972. Soilless mixes. Horticulture 50: 38–39.
Bosman, M. T. M. 1991. A monograph of the fern genus Microsorum (Polypodiaceae). Leiden Botanical
Series 14: i–ix, 1–161.
Brown, E. D. W. 1920. The value of nutrient solutions as culture media for fern prothallia. Torreya 20:
76–83.
Brownsey, P. J., and J. C. Smith-Dodsworth. 1989. New Zealand Ferns and Allied Plants. David Bateman,
Aukland.
Burrows, J. E. 1990. Southern African Ferns and Fern Allies. Frandsen Publishers, Sandton, Republic of
South Africa.
Ching, R. C., and Hsen Hsu Hu. 1930. Icones Filicum Sinicarum. Bishen Singh Mahendra Pal, Singh,
India.
Cody, W. J., and D. M. Britton. 1989. Ferns and Fern Allies of Canada. Canadian Government Publish-
ing Center, Ottawa.
Collinson, M. E. 1996. “What use are fossil ferns?”—20 years on: With a review of the fossil history of
extant pteridophyte families and genera. Pages 349–394. In: Pteridology in Perspective, edited by J. M.
Camus, M. Gibby, and R. J. Johns. Royal Botanic Gardens, Kew.
Crane, E. H. 1997. A revised circumscription of the genera of the fern family Vittariaceae. Systematic
Botany 32: 509–517.
565
566 L C
Curtwright, D. 1995. Ceratopteris thalactroides and its relatives. Journal of the Los Angeles International
Fern Society, Inc. 22: 28–33.
Dittmer, H. J., E. F. Castetter, and O. M. Clark. 1954. The ferns and fern allies of New Mexico. Univer-
sity of New Mexico Publications in Biology 6: 1–139.
Druery, C. T. 1912. British Ferns and Their Varieties. Routledge, London.
Dyce, J. W. 1963. Variation in Polystichum in the British Isles. British Fern Gazette 9: 97–109.
———. 1972. British fern varieties: The Scolopendriums. Newsletter No. 10 (November). British Pteri-
dological Society, London.
———. 1985. New fern varieties—wild and cultivated. Pteridologist 1: 78–80.
———. 1987. Classification of fern variations in Britain. Pteridologist 1: 154–155.
———. 1991. The variation and propagation of British ferns. Special Publication No. 3. British Pterido-
logical Society, London.
Dyer, A. F. 1979. The culture of fern gametophytes for experimental investigation. In: The Experimental
Biology of Ferns, edited by A. F. Dyer. Academic Press, London.
Edie, H. H. 1978. Ferns of Hong Kong. Hong Kong University Press, Hong Kong.
Evans, A. M. 1969. Interspecific relationships in the Polypodium pectinatum–plumula complex. Annals of
the Missouri Botanical Garden 55: 193–293.
Farrar, D. R. 1968. A culture chamber for tropical rain forest plants. American Fern Journal 58: 97–102.
Flora of North America Editorial Committee. 1993. Flora of North America North of Mexico. Vol. 2,
Pteridophytes and Gymnosperms. Oxford University Press, Oxford.
Ford, M. V., and M. F. Fay. 1990. Growth of ferns from spores in axenic culture. Methods in Molecular
Biology 6: 171–180.
Foster, F. G. 1984. Ferns to Know and Grow. 3rd ed. Timber Press, Portland, Oregon.
Franks, W. 1969. Platycerium-Fern Facts. Published privately, Los Angeles, California.
Fraser-Jenkins, C. R. 1986. A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae). Bul-
letin of the British Museum (Natural History), Botany Series 14: 183–218.
Goudey, C. J. 1985. Maidenhair Ferns in Cultivation. Lothian Publishing Co., Melbourne, Australia.
Grounds, R. 1974. Ferns. Pelham Books, London.
Hennipman, E. 1977. A monograph of the fern genus Bolbitis (Lomariopsidaceae). Leiden Botanical Series
2: i–xiii, 1–331.
Hennipman, E., and M. C. Roos. 1982. A monograph of the fern genus Platycerium (Polypodiaceae). Ver-
handelingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afdeeling Natuurkunde,
Tweede Reeks 80: 1–126.
Hickey, R. J. 1977. The Lycopodium obscurum complex in North America. American Fern Journal 67:
45–49.
Hickman, J. C., ed. 1993. The Jepson Manual: Higher Plants of California. University of California Press,
Berkeley.
Hill, R. H. 1976. Cold requirements of several ferns in southeastern Michigan. American Fern Journal 66:
83–88.
Hill, R. S. 1984. Propagation of Marattiaceae. Australian Fern Journal 74: 11–15.
Hoagland, D. R., and D. I. Arnon. 1950. The water-culture method for growing plants without soil. Cali-
fornia Agricultural Experiment Station, Circular 347. College of Agriculture, University of California,
Berkeley.
Hoshizaki, B. J. 1970a. The genus Adiantum in cultivation (Polypodiaceae). Baileya 17: 97–196.
———. 1970b. Rhizome scales of Platycerium. American Fern Journal 60: 144–160.
———. 1972. Morphology phylogeny of Platycerium species. Biotropica 4: 93–117.
———. 1975. The Fern Growers Manual. Alfred A. Knopf, New York.
L C 567
———. 1975. A staghorn fern (Platycerium) hybrid. American Fern Journal 65: 99–101.
———. 1981. The fern genus Davallia in cultivation (Davalliaceae). Baileya 21: 1–50.
———. 1982. The genus Polypodium in cultivation (Polypodiaceae). Baileya 22: 1–98.
———. 1991. An “intergeneric” hybrid: Aglaomorpha × Drynaria. American Fern Journal 81: 37–43.
Hoshizaki, B. J., and M. G. Price. 1990. Platycerium update. American Fern Journal 80: 58–69.
Hoshizaki, B. J., and K. A. Wilson. 1999. The cultivated species of the fern genus Dryopteris in the United
States. American Fern Journal 89: 1–100.
Hovenkamp, P. H. 1986. A monograph of the fern genus Pyrrosia. Leiden Botanical Series 9: i–xiii, 1–
310.
Hovenkamp, P. H., and N. A. P. Franken. 1993. An account of the fern genus Belvisia Mirbel (Polypodi-
aceae). Blumea 37: 511–527.
Hu, I. H., and R. C. Ching. 1930. Icones Filicum Sinicarum. Fascicle 1.
Huang, Tsen-Chieng, ed. 1994. Flora of Taiwan. 2nd ed. Vol. 1, Pteridophyta and Gymnospermae. Edi-
torial Committee of the Flora of Taiwan, Ta’an Taipei, Taiwan.
Iwatsuki, K., T. Yamazaki, D. Boufford, and H. Ohba. 1995. Pteridophyta and Gymnospermae. Vol. 1 of
Flora of Japan. Kodansha, Tokyo, Japan.
Jacobsen, W. B. G. 1983. The Ferns and Fern Allies of Southern Africa. Butterworth Publishers, Durban,
South Africa.
Joe, B. 1958. Pteris species cultivated in California. Lasca Leaves 8: 26–29.
———. 1964. A review of the species of Platycerium (Polypodiaceae). Baileya 12: 69–126.
Johns, R. J. 1995. Lecanopteris lomarioides, Polypodiaceae. Curtis’s Botanical Magazine 12: 89–95.
Johnson, D. M. 1986. Systematics of the New World species of Marsilea (Marsileaceae). Systematic Botany
Monographs 11: 1–87.
Jones, D. L. 1987. Encyclopaedia of Ferns. Timber Press, Portland, Oregon.
Jones, D. L., and S. C. Clemesha. 1981. Australian Ferns and Fern Allies. A. H. and A. W. Reed, Sydney,
Australia.
Kato, M. 1984. A taxonomic study of the athyrioid fern genus Deparia with main reference to the Pacific
species. Journal of the Faculty of Science, University of Tokyo, section 3, 13: 375–429.
———. 1985. A systematic study of the genera of the fern family Davalliaceae. Journal of the Faculty of
Science, University of Tokyo, section 3, 13: 553–573.
Kaye, R. 1965. Variations in Athyrium in the British Isles. British Fern Gazette 9: 197–204.
———. 1968. Hardy Ferns. Faber and Faber, London.
Khullar, S. P. 1994. An Illustrated Fern Flora of West Himalaya. International Book Distributor, Dehra
Dun, India.
Knobloch, I. W., and D. S. Correll. 1962. Ferns and Fern Allies of Chihuahua, Mexico. Texas Research
Foundation, Renner, Texas.
Knudson, L. 1946. A new nutrient solution for the germination of orchid seed. American Orchid Society
Bulletin 15: 214–217.
Kramer, K. U., and P. S. Green, eds. 1990. Pteridophytes and Gymnosperms. Vol. 1 of The Families and
Genera of Vascular Plants, edited by K. Kubitzki. Springer-Verlag, Berlin.
Krusberg, L. R. 1992. Best nematode control. Flower and Garden 36 (2): 8.
Kurata, S. 1963. On Japanese ferns belonging to the genus Cyrtomium. Science Report of the Yokosuka
City Museum 8: 23–47.
Kurata, S., and T. Nakaike. 1964–1997. Illustrations of Pteridophytes of Japan. Vols. 1–8. University of
Tokyo Press, Tokyo.
Langsdorff, G. H., and F. Fischer. 1810. Plantes recueillies pendant le voyage des Russes autour du monde.
Published privately by the authors, Tübingen.
568 L C
Launert, E. 1968. A monographic survey of the genus Marsilea Linnaeus. I: The species of Africa and
Madagascar. Senckenbergiana Biologica 49: 273–315.
Lee, H. 1887. The vegetable lamb of Tartary, a curious fable of the cotton plant. Sampson, Low, Marston,
Searle, and Rivington, London.
Lellinger, D. B. 1972. A revision of the fern genus Niphidium. American Fern Journal 62: 101–120.
———. 1985. A Field Manual of the Ferns and Fern Allies of the United States and Canada. Smithson-
ian Institution Press, Washington, D.C.
———. 1988. Some new species of Campyloneurum and a provisional key to the genus. American Fern
Journal 78: 14–35.
———. 1989. The ferns and fern allies of Costa Rica, Panama, and the Chocó. Part 1: Psilotaceae through
Dicksoniaceae. Pteridologia 2A: 5–364.
———. 1994. Useful fern books in print: 1993–1994. Fiddlehead Forum 21: 19–21.
Little, E. L., Jr., and F. H. Wadsworth. 1964. Common trees of Puerto Rico and the Virgin Islands. Agri-
cultural Handbook No. 249. United States Department of Agriculture, Forest Service, Washington,
D.C.
Lloyd, R. M. 1974. Systematics of the genus Ceratopteris Brongn. (Parkeriaceae). II: Taxonomy. Brittonia
26: 139–160.
Lloyd, R. M., and E. J. Klekowski Jr. 1970. Spore germination and viability in Pteridophyta: Evolution-
ary significance of chlorophyllous spores. Biotropica 2: 129–137.
Lovis, J. D. 1968. Fern hybridists and fern hybridising. II: Fern hybridising at the University of Leeds. Brit-
ish Fern Gazette 10: 13–20.
Lovis, J. D., and G. Vida. 1969. The resynthesis and cytogenetic investigation of ×Asplenophyllitis microdon
and ×A. jacksonii. British Fern Gazette 10: 53–67.
Lowe, E. J. 1890. British Ferns and Where Found. Swan Sonnenschein, London.
Lumpkin, T. A., and D. L. Plucknett. 1980. Azolla: Botany, physiology, and use as a green manure. Eco-
nomic Botany 34: 111–153.
MacPhail, J. 1990. Trough gardening. Pacific Horticulture (spring): 31–39.
Marticorena, C., and R. Rodríguez. 1995. Pteridophyta–Gymnospermae. Vol. 1 of Flora de Chile. Uni-
versidad de Concepción, Concepción, Chile.
Matkin, O. A., and P. A. Chandler. 1957. The U.C.-type soil mixes. In: U.C. System for Producing
Healthy Container-Grown Plants, edited by K. F. Baker. Extension Service Manual No. 23, Califor-
nia Agricultural Experiment Station.
Maxon, W. R. 1912. Studies of tropical American ferns. Contributions to the United States National
Herbarium 16: 54–58.
Mickel, J. T. 1979. How to Know the Ferns and Fern Allies. W. C. Brown, Dubuque, Iowa.
———. 1992. Pteridophytes. In: Gymnosperms and Pteridophytes, Vol. 17 of Flora Novo-Galiciana,
edited by R. McVaugh. University of Michigan, Ann Arbor.
———. 1994. Ferns for American Gardens. MacMillan Publishing Company, New York.
Mickel, J. T., and J. M. Beitel. 1988. Pteridophyte flora of Oaxaca, Mexico. Memoirs of the New York
Botanical Garden 46: 1–568.
Moore, A. W. 1969. Azolla: Biology and agronomic significance. The Botanical Review 35: 17–35.
Moran, R. C. 1982. The Asplenium trichomanes complex in the United States and adjacent Canada. Amer-
ican Fern Journal 72: 5–11.
———. 1986. The neotropical fern genus Olfersia. American Fern Journal 76: 161–178.
———. 1987. Monograph of the neotropical fern genus Polybotrya (Dryopteridaceae). Illinois Natural
History Survey Bulletin 34 (1): i–v, 1–138.
———. 1992a. The vegetable lamb of Tartary—A pteridological tale. Fiddlehead Forum 19: 2, 7, 8.
L C 569
Rush, R. 1983. A hardy hybrid Adiantum. British Pteridological Society Bulletin 2: 261–262.
———. 1984a. A guide to hardy ferns. Special Publication No. 1. British Pteridological Society, London.
———. 1984b. Raising pteridophytes from spores: The special cases. Pteridologist 1: 3–5.
Scamman, E. 1960. The maidenhair ferns (Adiantum) of Costa Rica. Contributions of the Gray Herbar-
iumof Harvard University 187: 3–22.
Sessions, A. 1978. Key to the cultivars of Nephrolepis exaltata ‘Bostoniensis’. Privately published.
Shivas, M. G. 1962. The Polypodium vulgare complex. British Fern Gazette 9: 65–70.
Smith, A. R. 1981. Pteridophytes. In:Flora of Chiapas, Part 2, edited by D. E. Breedlove. California Acad-
emy of Sciences, San Francisco.
———. 1995. Pteridophytes. In:Flora of the Venezuelan Guayana, Vol. 2, edited by J. Steyermark, P. E.
Berry, and B. K. Holst. Timber Press, Portland, Oregon.
Snyder, L. H., Jr., and J. G. Bruce. 1986. Field Guide to the Ferns and Other Pteridophytes of Georgia.
University of Georgia Press, Athens.
de la Sota, E. R. 1963. Contribución al conocimiento de las “Salviniaceae” neotropicales. IV. Darwiniana
12: 612–623.
———. 1966. Revisión de las especies Americanas del grupo “Polypodium squamatum” L. “Polypodi-
aceae” (s. str.). Revista del Museo de La Plata, Sección Botânica 10: 69–186, t. I–VII.
Sowerby, J. E., and C. Johnson. 1855. The Ferns of Great Britain. Self-published, London.
Tagawa, M., and K. Iwatsuki. 1979–1989. Flora of Thailand. Vol. 3, Pteridophytes. Chutima Press,
Bangkok.
Taylor, W. C. 1984. Arkansas Ferns and Fern Allies. Milwaukee Public Museum, Milwaukee, Wisconsin.
Thurston, S. H. 1939. Forcing a collection of native ferns of New England and the middle Atlantic states
for exhibition. American Fern Journal 29: 85–94.
Tindale, M. 1965. A monograph of the genus Lastreopsis Ching. Contributions to the New South Wales
National Herbarium 3: 249–339, t. 1–23.
Trehane, P., ed. 1995. International Code of Nomenclature for Cultivated Plants. Regnum Vegetabile
133: 1–175.
Tryon, A. F. 1957. A revision of the fern genus Pellaea section Pellaea. Annals of the Missouri Botanical
Garden 44: 125–193.
Tryon, R. M. 1956. A revision of the American species of Notholaena. Contributions of the Gray Herbar-
iumof Harvard University 179: 1–106.
———. 1962. Taxonomic fern notes. II: Pityrogramma and Anogramma. Contributions of the Gray
Herbariumof Harvard University189: 5–76.
Tryon, R. M., and R. G. Stolze. 1989. Pteridophyta of Peru. Part I: 1. Ophioglossaceae–12. Cyatheaceae.
Fieldiana, Botany, new series 20: 1–145.
———. 1989. Pteridophyta of Peru. Part II: 13. Pteridaceae–15. Dennstaedtiaceae. Fieldiana, Botany, new
series 22: 1–128.
———. 1991. Pteridophyta of Peru. Part IV: 17. Dryopteridaceae. Fieldiana, Botany, new series 27: 1–
176.
———. 1993. Pteridophyta of Peru. Part V: 18. Aspleniaceae–21. Polypodiaceae. Fieldiana, Botany, new
series 27: 1–176.
———. 1994. Pteridophyta of Peru. Part VI: 22. Marsileaceae–28. Isoëtaceae. Fieldiana, Botany, new
series 32: 1–190.
Tryon, R. M., and A. F. Tryon. 1982. Ferns and Allied Plants, with Special Reference to Tropical Amer-
ica. Springer-Verlag, Berlin.
Tutin, T. G., V. H. Heywood, N. A. Burges, D. H. Valentine, S. M. Walters, D. A. Webb. 1964. Flora
Europaea. Vol. 1, Lycopodiaceae to Platanaceae. Cambridge University Press, Cambridge.
L C 571
Vail, R. 1984. Platycerium Hobbyist’s Handbook. Desert Biological Publications, Mena, Arkansas.
van Hove, C. 1989. Azolla and its multiple uses, with emphasis on Africa. Food and Agriculture Organi-
zation of the United Nations, Rome.
Vareschi, V. 1968. Helechos. In:Flora de Venezuela, Vol. 1, parts 1 and 2. Edición Especial del Instituto
Botânico, Caracas.
Wagner, D. H. 1979. Systematics of Polystichum in western North America north of Mexico. Pteridologia
1: 1–64.
Wagner, G. M. 1997. Azolla: A review of its biology and utilization. Botanical Review 63: 1–26.
Wagner, W. H., Jr. 1942. Bipinnate Christmas fern. American Fern Journal 32: 27–29.
Wagner, W. H., Jr., and F. S. Wagner. 1975. A hybrid polypody from the New World tropics. Fern
Gazette 11: 125–135.
Walker, T. G. 1970. Species of Pteris commonly in cultivation. British Fern Gazette 10: 143–151.
Weatherby, C. A. 1939. The group of Polypodium polypodioides. Contributions of the Gray Herbariumof
Harvard University 124: 22–35.
Webster, T. R. 1979. An artificial crossing technique for Selaginella. American Fern Journal 69: 9–13.
Wherry, E. T. 1972. The Southern Fern Guide. Doubleday, Garden City, New York.
Windham, M. D. 1987. Argyrochosma, a new genus of cheilanthoid ferns. American Fern Journal 77:
37–41.
Yatskievych, G. 1996. A revision of the fern genus Phanerophlebia (Dryopteridaceae). Annals of the Mis-
souri Botanical Garden 83: 168–199.
This page intentionally left blank
Subject Index
573
574 S I
Plants listed in boldface have entries in Chapter 13, “Ferns and Fern Allies in Cultivation.” Cultivars and
common names are cross-referenced to species or genus. Cultivars are listed at the end of the genus.
abrupt-tipped maiden fern, see Thelypteris augescens, 519 Adiantum jordanii, 148, 166
Acrosorus, 333 Adiantum latifolium, 166
Acrostichum, 156 Adiantum lucidum, 167
Acrostichum aureum, 93, 156 Adiantum lunulatum, 167
Acrostichum danaeifolium, 93, 157 Adiantum macrophyllum, 95, 103, 167
Actiniopteris, 111, 157 Adiantum ×mairisii, see Adiantum capillus-veneris
Actiniopteris radiata, 158 ‘Mairisii’, 162
Actiniopteris semiflabellata, 157 Adiantum monochlamys, 168, color plate 10
Acystopteris japonica, 258 Adiantum patens, 168
adder’s-tongue fern, see Ophioglossum, 404 Adiantum pedatum, 29, 91, 92, 97, 103, 168
Adenophorus, 333 Adiantum pedatum subsp. aleuticum, see Adiantum
Adiantopsis, 158 aleuticum, 160
Adiantopsis radiata, 158 Adiantum pentadactylon, 169
Adiantum, 102–104, 159 Adiantum peruvianum, 103, 170
Adiantum aethiopicum, 159 Adiantum petiolatum, 170
Adiantum aleuticum, 92, 148, 160 Adiantum philippense, see Adiantum lunulatum, 167
Adiantum aleuticum subsp. subpumilum, 98, 160 Adiantum poiretti, 170
Adiantum anceps, 103, 160 Adiantum polyphyllum, 171
Adiantum banksianum, see Adiantum capillus-veneris Adiantum pubescens, 166
‘Mairisii’, 161 Adiantum pulverulentum, 171
Adiantum bellum, 160 Adiantum pyramidale, 171
Adiantum capillus-veneris, 82, 92, 98, 101, 103, 161 Adiantum raddianum, 82, 92, 94, 95, 97, 98, 103, 172
Adiantum caudatum, 69, 70, 103, 162 Adiantum reniforme, 173
Adiantum concinnum, 162 Adiantum seemannii, 173
Adiantum cristatum, see Adiantum pyramidale, 171 Adiantum tenerum, 94, 103, 173
Adiantum cultratum, 163 Adiantum tetraphyllum, 174
Adiantum cuneatum, see Adiantum raddianum, 172 Adiantum ×tracyi, 83, 103, 148, 175
Adiantum diaphanum, 163 Adiantum trapeziforme, 91, 103, 175
Adiantum excisum, 164 Adiantum venustum, 98, 175
Adiantum flabellulatum, 164 Adiantum villosum, 176
Adiantum formosum, 103, 164 Adiantum cultivars
Adiantum fragile, 165 ‘Banksianum’, see Adiantum capillus-veneris ‘Mairisii’,
Adiantum fructuosum, 165 162
Adiantum hispidulum, 82, 95, 97, 98, 100, 101, 103, ‘Birkenheadii’, see Adiantum diaphanum, 162
166, color plate 9 ‘Croweanum’, see Adiantum raddianum, 172
581
582 P N I
Christella parasitica, see Thelypteris parasitica, 522 Coniogramme ‘Variegata’, see Coniogramme japonica, 248
Christmas fern, see Polystichum acrostichoides, 455 Cooper’s tree fern, see Sphaeropteris cooperi, 91, 92, 511,
Cibotium, 110, 111, 244 color plates 2, 3
Cibotium barometz, 245 copper fern, see Bommeria, 225
Cibotium chamissoi, 245 cotton fern, see Cheilanthes newberryi, 242
Cibotium glaucum, 16, 93, 95, 111, 245, color plate 3 couplet fern, see Dennstaedtia bipinnata, 278
Cibotium menziesii, 111, 246 Coville’s lip fern, see Cheilanthes covillei, 238
Cibotium schiedei, 94, 100, 111, 246 crape fern, see Leptopteris superba, 351
cinnamon fern, see Osmunda cinnamomea, 407, color creeping star-hair fern, see Thelypteris reptans, 524
plate 8 creeping wood fern, see Thelypteris reptans, 524
Cionidium, see Tectaria, 515 Crepidomanes, see Trichomanes, 529
claw fern, see Onychium, 402 Crepidomanes auriculatum, see Trichomanes auriculatum,
cliff brake, see Aspidotis densa, 193; Cryptogramma 528
stelleri, 251; Pellaea, 412 crested lady fern, see Athyrium filix-femina ‘Cristatum’,
cliff fern, see Woodsia, 531 212
climbing fern, see Lygodium, 360 crested tongue fern, see Pyrrosia lingua ‘Cristata’, 485
climbing vine fern, see Microgramma heterophylla, 372 crested wood fern, see Dryopteris cristata, 305
Clinton’s wood fern, see Dryopteris clintoniana, 303 Cretan brake, see Pteris cretica, 474
cloak fern, see Notholaena, 399; see also Argyrochosma, crisped male fern, see Dryopteris affinis ‘Crispa’, 299
188; Astrolepis, 206 crowe maidenhair, see Adiantum raddianum
club moss, see Lycopodium, 354 ‘Croweanum’, 172
coastal leather fern, see Acrostichum aureum, 156 Crypsinus, see Selliguea, 507
coastal wood fern, see Dryopteris arguta, 300 Cryptogramma, 248
coast polypody, see Polypodium scouleri, 451 Cryptogramma acrostichoides, 249
Cochlidium, 333 Cryptogramma cascadensis, 249
Cochlidium serrulatum, 333 Cryptogramma crispa, 98, 250, color plate 23
coffee fern, see Pellaea andromedifolia, 412 Cryptogramma raddeana, 250
coin-spot tree fern, see Sphaeropteris cooperi, 91, 92, 511, Cryptogramma sitchensis, 250
color plates 2, 3 Cryptogramma stelleri, 251
Colysis, see Leptochilus, 349 Ctenitis, 251
Colysis ampla, see Leptochilus amplus, 349 Ctenitis ampla, see Ctenitis sloanei, 251
Colysis ellipitica, see Leptochilus ellipticus, 349 Ctenitis decomposita, see Lastreopsis microsora, 344
Colysis insignis, see Microsorum insigne, 381 Ctenitis pentangularis, see Lastreopsis microsora subsp.
Colysis pothifolia, see Leptochilus ellipticus, 350 pentangularis, 344
Colysis pteropus, see Microsorum pteropus, 384 Ctenitis sloanei, 251
Colysis sayeri, see Leptochilus amplus, 349 Ctenitis subincisa, see Megalastrum subincisum, 372
Colysis wrightii, see Leptochilus macrophyllus var. wrightii, Ctenitis sect. Subincisae, see Megalastrum, 371
350 Ctenitis submarginalis, 251
comb fern, see Pecluma, 410 Ctenopteris, 333
common golden-scaled male fern, see Dryopteris affinis, cup fern, see Dennstaedtia, 277
298 Currania, see Gymnocarpium, 334
common horsetail, see Equisetum arvense, 326 Currania oyamensis, see Gymnocarpium oyamense, 335
common male fern, see Dryopteris filix-mas, 308 Cyathea, 71, 252
common oak fern, see Gymnocarpium dryopteris, 335, Cyathea aramaganensis, see Sphaeropteris armaganensis,
color plate 29 510
common polypody, see Polypodium vulgare, 452 Cyathea arborea, 253
common staghorn, see Platycerium bifurcatum, 429, Cyathea australis, see Alsophila australis, 182
color plates 1, 5, 42 Cyathea brownii, see Sphaeropteris excelsa, 511
common sword fern, see Nephrolepis exaltata, 393; Cyathea cooperi, see Sphaeropteris cooperi, 91, 92, 511,
Polystichum munitum, 460 color plates 2, 3
common tassel fern, see Lycopodium phlegmaria, 358 Cyathea dealbata, see Alsophila tricolor, 182, color plate
Coniogramme, 247 11
Coniogramme fraxinea, see Coniogramme intermedia, 248 Cyathea medullaris, see Sphaeropteris medullaris, 512
Coniogramme intermedia, 248 Cyathea princeps, see Sphaeropteris horrida, 512
Coniogramme japonica, 248 Cyclopeltis, 253
P N I 587
Elaphoglossum vestitum, 325 fragile bladder fern, see Cystopteris fragilis, 260
elephant-ear fern, see Elaphoglossum crinitum, 322 fragile rock brake, see Cryptogramma stelleri, 251
elkhorn fern, see Platycerium, 427 fragrant fern, see Microsorum scandens, 386
Equisetum, 74, 93, 115, 326 Fritz Luth maidenhair, see Adiantum raddianum ‘Fritz
Equisetum arvense, 326 Luth’, 172
Equisetum diffusum, 327
Equisetum ×ferrissii, 327 ghost fern, see Athyrium niponicum ‘Pictum’, 212
Equisetum hyemale, 328 giant chain fern, see Woodwardia fimbriata, 537
Equisetum laevigatum, 328 giant fern, see Acrostichum danaeifolium, 157
Equisetum scirpoides, 329 giant horsetail, see Equisetum telmateia, 329
Equisetum telmateia, 329 giant maidenhair, see Adiantum polyphyllum, 171
Equisetum variegatum, 329 giant staghorn, see Platycerium superbum, 435, color
erect sword fern, see Nephrolepis cordifolia, 391, color plate 43
plate 35 giant sword fern, see Nephrolepis biserrata, 390
European chain fern, see Woodwardia radicans, 537, giant wood fern, see Dryopteris goldiana, 311
color plate 50 glade fern, see Deparia acrostichoides, 280; Diplazium
European lady fern, see Athyrium filix-femina var. filix- pycnocarpon, 288
femina, 212 glandular wood fern, see Dryopteris intermedia, 311;
European ostrich fern, see Matteuccia struthiopteris var. Thelypteris resinifera, 524
struthiopteris, 371 Gleichenella, 330
European parsley fern, see Cryptogramma crispa, 250, Gleichenia, 330
color plate 23 Gleichenia dicarpa, 330
evergreen wood fern, see Dryopteris intermedia, 311 Gleichenia microphylla, 330
Gleichenia rupestris, 330
fairy moss, see Azolla, 213 Gleicheniaceae, 330
fancy fern, see Dryopteris intermedia, 311 glossy spleenwort, see Asplenium oblongifolium, 200
fan maidenhair, see Adiantum tenerum, 173 goldback fern, see Pentagramma, 417; Pityrogramma,
Farley maidenhair, see Adiantum tenerum ‘Farleyense’, 424
173 golden polypody, see Phlebodium, 419; Phlebodium
felt fern, see Pyrrosia, 480 aureum, 420
Fendler’s lip fern, see Cheilanthes fendleri, 240, color Goldie’s fern, see Dryopteris goldiana, 311
plate 22 Goniophlebium, 98, 331, 439
ferny azolla, see Azolla pinnata, 214, color plate 16 Goniophlebium amoenum, see Polypodium amoenum, 441
Ferriss’s scouring rush, see Equisetum ×ferrissii, 327 Goniophlebium ×ekstrandii, 331
fibrous dicksonia, see Dicksonia fibrosa, 282 Goniophlebium formosanum, see Polypodium formosanum,
field horsetail, see Equisetum arvense, 326 444, color plate 44
Fiji davallia, see Davallia fejeensis, 267 Goniophlebium percussum, 332
filmy fern, see Hymenophyllum, 339; Trichomanes, 528 Goniophlebium persicifolium, 332
filmy maidenhair, see Adiantum diaphanum, 163 Goniophlebium subauriculatum, 94, 333, color plate 28
fir club moss, see Lycopodium selago, 360 Goniophlebium verrucosum, see Goniophlebium percussum,
fish-bone fern, see Nephrolepis, 389 332
fishtail sword fern, see Nephrolepis falcata ‘Furcans’, 395 Goniophlebium cultivars
five-finger fern, see Adiantum pedatum, 168 ‘Knightiae’, see Goniophlebium subauriculatum, 333,
flat-branch club moss, see Lycopodium complanatum, 357 color plate 28
flat-branched tree club moss, see Lycopodium obscurum, ‘Nola’, see Goniophlebium ×ekstrandii, 331
358 Goniopteris, see Thelypteris, 517
flexuous cliff brake, see Pellaea ovata, 415 Goniopteris poiteana, see Thelypteris poiteana, 523
Florida tree fern, see Ctenitis sloanei, 251 Goniopteris reptans, see Thelypteris reptans, 524
flowering fern, see Anemia, 182; Osmunda regalis, 408, Grammitidaceae, 333
color plate 6 Grammitis, 74, 333
fluffy ruffles, see Nephrolepis exaltata ‘Fluffy Ruffles’, 395 grape fern, see Botrychium, 226
forest brake, see Pteris altissima, 473 grass fern, see Vittaria lineata, 530
forked spleenwort, see Asplenium septentrionale, 204 green cliff brake, see Pellaea viridis, 416
forking fern family, see Gleicheniaceae, 330 green petticoats, see Adiantum capillus-veneris
four-leaved maidenhair, see Adiantum tetraphyllum, 174 ‘Imbricatum’, 162
P N I 591
green spleenwort, see Asplenium trichomanes-ramosum, Humata falcinella, see Davallia falcinella, 266
206 Humata griffithiana, see Davallia griffithiana, 267
green staghorn, see Platycerium hillii, 432 Humata heterophylla, see Davallia heterophylla, 268
grid-scale maiden fern, see Thelypteris patens, 523 Humata parvula, see Davallia parvula, 272
ground cedar, see Lycopodium complanatum, 357 Humata repens, see Davallia repens, 273
ground fern, see Hypolepis, 341 Humata tyermannii, see Davallia tyermannii, 276
ground pine, see Lycopodium, 113, 354; Lycopodium Humata vestita, see Davallia repens, 273
obscurum, 358 Huperzia carinata, see Lycopodium carinatum, 356
Gymnocarpium, 334 Huperzia filiformis, see Lycopodium filiforme, 357
Gymnocarpium disjunctum, 335 Huperzia lucidula, see Lycopodium lucidulum, 358
Gymnocarpium dryopteris, 92, 96, 97, 98, 335, color Huperzia nummulariifolium, see Lycopodium
plate 29 nummulariifolium, 358
Gymnocarpium oyamense, 335 Huperzia phlegmaria, see Lycopodium phlegmaria, 358
Gymnocarpium robertianum, 94, 335 Huperzia phlegmarioides, see Lycopodium phlegmarioides,
Gymnocarpium ‘Plumosum’, see Gymnocarpium 359
dryopteris, 335, color plate 29 Huperzia polytrichoides, see Lycopodium filiforme, 357
Gymnopteris, see Hemionitis, 336 Huperzia selago, see Lycopodium selago, 360
Huperzia squarrosa, see Lycopodium squarrosum, 360
hacksaw fern, see Doodia, 289 Hymenodium, see Elaphoglossum, 320
hairy lip fern, see Cheilanthes lanosa, 241 Hymenodium crinitum, see Elaphoglossum crinitum, 322
halberd fern, see Tectaria heracleifolia, 515 Hymenophyllum, 339
hammock fern, see Blechnum occidentale, 220 Hymenophyllum demissum, 340
hand fern, see Doryopteris palmata, 292; Ophioglossum, Hymenophyllum polyanthos, 340
404 Hypoderris, 340
Haplopteris, 530 Hypoderris brownii, 341
Haplopteris elongata, see Vittaria elongata, 530 Hypolepis, 94, 97, 341
hapu, see Cibotium glaucum, 245 Hypolepis millefolium, 342
hard fern, see Paesia scaberula, 409 Hypolepis punctata, 342
hard male fern, see Dryopteris affinis, 298 Hypolepis repens, 342
hard shield fern, see Polystichum aculeatum, 456 Hypolepis tenuifolia, 82, 342
hare’s-foot fern, see Davallia canariensis, 264;
Phlebodium aureum, 420 imbricate sword fern, see Polystichum imbricans, 458
Hartford fern, see Lygodium palmatum, 363 Indian’s dream, see Aspidotis densa, 193
hart’s-tongue fern, see Phyllitis scolopendrium, 422, color Indian staghorn, see Platycerium wallichii, 436
plate 39 inland leather fern, see Acrostichum danaeifolium, 157
Hawaiian tree fern, see Cibotium glaucum, 245 intermediate polypody, see Polypodium interjectum, 446
hay-scented fern, see Dennstaedtia punctilobula, 279 interrupted fern, see Osmunda claytoniana, 407, color
hay-scented wood fern, see Dryopteris aemula, 298 plate 37
Hemigramma, see Tectaria, 513
Hemigramma latifolia, see Tectaria hilocarpa, 516 Japanese ball fern, see Davallia mariesii var. mariesii, 271
Hemionitis, 101, 336 Japanese bird’s-nest fern, see Asplenium antiquum, 195
Hemionitis arifolia, 337, color plate 30 Japanese claw fern, see Onychium japonicum, 402
Hemionitis palmata, 338 Japanese climbing fern, see Lygodium japonicum, 362,
Hemionitis pinnatifida, 338 color plate 31
Hemitelia, see Cyathea, 252 Japanese felt fern, see Pyrrosia lingua, 484
Himalayan holly fern, see Polystichum lentum, 459 Japanese osmunda, see Osmunda japonica, 408
Himalayan maidenhair, see Adiantum venustum, 175 Japanese painted fern, see Athyrium niponicum ‘Pictum’,
Histiopteris, 338 212, color plate 14
Histiopteris incisa, 339 Japanese sword fern, see Polystichum polyblepharum, 462
holly fern, see Cyrtomium, 254; Polystichum, 454 Japanese wood fern, see Dryopteris sieboldii, 317
horseshoe fern, see Marattia salicina, 365 Java staghorn, see Platycerium willinckii, 437
horsetail, see Equisetum, 325 jointed polypody, see Goniophlebium subauriculatum,
house holly fern, see Cyrtomium falcatum, 255, color 333, color plate 28
plate 24 joint fern, see Arthropteris, 190
Humata, see Davallia, 262 Jones’s cloak fern, see Argyrochosma jonesii, 189
592 P N I
net-veined chain fern, see Woodwardia areolata, 537 Ophioglossum palmatum, 404
Newberry’s lip fern, see Cheilanthes newberryi, 242 Ophioglossum pendulum, 405, color plate 36
New York fern, see Thelypteris noveboracensis, 521 Ophioglossum petiolatum, 405
New Zealand cliff brake, see Pellaea rotundifolia, 416 Ophioglossum vulgatum, 406
New Zealand dicksonia, see Dicksonia squarrosa, 284 Oregon spike moss, see Selaginella oregana, 502
Niphidium, 398, 439 Oregon woodsia, see Woodsia oregana, 534
Niphidium albopunctatissimum, 399 Oreopteris, see Thelypteris, 517
Niphidium crassifolium, 398 Oreopteris limbosperma, see Thelypteris limbosperma, 69,
Niphobolus, see Pyrrosia, 480 520
nodding club moss, see Lycopodium cernuum, 356 Oriental chain fern, see Woodwardia orientalis, 537
Norfolk Island tree fern, see Sphaeropteris excelsa, 511 Oriental ostrich fern, see Matteuccia orientalis, 370
North American walking fern, see Camptosorus Oriental water fern, see Ceratopteris thalictroides, 233,
rhizophyllus, 228 color plate 19
northern holly fern, see Polystichum lonchitis, 460 Osmunda, 12, 74, 82, 93, 97, 406
northern lady fern, see Athyrium filix-femina var. Osmunda cinnamomea, 92, 407, color plate 8
angustum, 211 Osmunda claytoniana, 29, 91, 94, 407, color plate 37
northern spreading wood fern, see Dryopteris expansa, Osmunda japonica, 408
308 Osmunda regalis, 91, 93, 96, 408, color plate 6
northern woodsia, see Woodsia alpina, 532 Osmunda regalis var. brasiliensis, 408
northwestern lady fern, see Athyrium filix-femina var. Osmunda regalis var. regalis, 409
cyclosorum, 212 Osmunda regalis var. spectabilis, 409
notched water fern, see Marsilea schelpeana, 368 Osmunda ×ruggii, 408
Notholaena, 111, 236, 399 Osmunda vachellii, 407
Notholaena aurea, see Cheilanthes bonariensis, 238 Osmunda cultivars
Notholaena buchtienii, see Cheilanthes buchtienii, 238 ‘Cristata’, see Osmunda regalis var. regalis, 409
Notholaena dealbata, see Argyrochosma dealbata, 189 ‘Purpurescens’, see Osmunda regalis var. regalis, 409
Notholaena jonesii, see Argyrochosma jonesii, 189 ostrich fern, see Matteuccia, 369
Notholaena newberryi, see Cheilanthes newberryi, 242
Notholaena parvifolia, see Argyrochosma microphylla, 190 Pachypleuria, see Davallia, 262
Notholaena sinuata, see Astrolepis sinuata, 206 Pachypleuria parvula, see Davallia parvula, 272
Notholaena standleyi, 399 Pachypleuria repens, see Davallia repens, 273
Notholaena vestita, see Cheilanthes lanosa, 241 pacific maid, see Adiantum raddianum ‘Pacific Maid’,
173
oak fern, see Gymnocarpium, 334 paddle fern, see Elaphoglossum, 320
oak-leaf fern, see Drynaria, 293 Paesia, 409
Oaxaca spike moss, see Selaginella oaxacana, 502 Paesia scaberula, 409
Obake pyrrosia, see Pyrrosia lingua ‘Cristata’, 485 Palhinhaea cernua, see Lycopodium cernuum, 356
Odontosoria, 399 Paragramma longifolia, see Lepisorus longifolius, 348
Odontosoria chinensis, 400 Parahemionitis, see Hemionitis, 336
Oleandra, 400 Parahemionitis arifolia, see Hemionitis arifolia, 337, color
Oleandra articulata, 400 plate 30
Oleandra neriiformis, 401 Parathelypteris, see Thelypteris, 517
Oleandra nodosa, see Oleandra articulata, 400 Parathelypteris nevadensis, see Thelypteris nevadensis, 520
Oleandra pilosa, 401 Parathelypteris noveboracensis, see Thelypteris
Oleandra pistillaris, 401 noveboracensis, 521
Olfersia, 401 Parathelypteris simulata, see Thelypteris simulata, 526
Olfersia cervina, 402 parsley fern, see Cheilanthes hirta, 241; Cryptogramma,
Onoclea, 402 248
Onoclea sensibilis, 29, 91, 92, 93, 94, 96, 97, 146, 402 peacock fern, see Selaginella willdenovii, 506
Onoclea struthiopteris, see Matteuccia struthiopteris, 371, peacock plant, see Selaginella uncinata, 505, color plate
color plate 33 49
Onychium, 402 peacock pyrrosia, see Pyrrosia lingua ‘Nankin-shishi’, 485
Onychium densum, see Aspidotis densa, 193 Pecluma, 410, 439
Onychium japonicum, 403 Pecluma dispersa, 410
Ophioglossum, 69, 404 Pecluma pectinata, 410
596 P N I
Polypodium dryopteris, see Gymnocarpium dryopteris, 335, Polypodium persicifolium, see Goniophlebium
color plate 29 persicifolium, 332
Polypodium ×ekstrandii, see Goniophlebium ×ekstrandii, Polypodium phyllitidis, see Campyloneurum phyllitidis,
331 230
Polypodium formosanum, 444, color plate 44 Polypodium phymatodes, see Microsorum scolopendria, 380
Polypodium fortunei, see Microsorum fortunei, 380 Polypodium piloselloides, see Microgramma piloselloides,
Polypodium fraxinifolium, 448 374
Polypodium glaucophyllum, see Polypodium levigatum, Polypodium plebeium, 448
447 Polypodium polycarpon, see Microsorum punctatum, 384,
Polypodium glaucopruniatum, see Phlebodium color plate 34
pseudoaureum, 420, color plates 5, 38 Polypodium polypodioides, 449
Polypodium glaucum, see Phlebodium pseudoaureum, 420, Polypodium polypodioides var. aciculare, 449
color plates 5, 38 Polypodium polypodioides var. burchellii, 449
Polypodium glycyrrhiza, 444 Polypodium polypodioides var. ecklonii, 449
Polypodium grossum, see Microsorum grossum, 380 Polypodium polypodioides var. michauxianum, 449
Polypodium guttatum, 445 Polypodium polypodioides var. minus, 449
Polypodium hancockii, see Microsorum insigne, 381 Polypodium pseudoaureum, see Phlebodium pseudoaureum,
Polypodium hesperium, 445 420, color plates 5, 38
Polypodium heterophyllum, see Microgramma heterophylla, Polypodium pteropus, see Microsorum pteropus, 384
372 Polypodium ptilorhizon, 449
Polypodium hirsutissimum, 446 Polypodium punctatum, see Microsorum punctatum, 384,
Polypodium integrifolium, see Microsorum punctatum, color plate 34
384, color plate 34 Polypodium pustulatum, see Microsorum pustulatum, 385;
Polypodium interjectum, 446 Microsorum scandens, 386
Polypodium irioides, see Microsoum punctatum, 384 Polypodium pyrrholepis, 98, 449, color plate 45
Polypodium knightiae, see Goniophlebium subauriculatum Polypodium rhodopleuron, 450
‘Knightiae’, 333, color plate 28 Polypodium sanctae-rosae, 450
Polypodium kuhnii, see Polypodium attenuatum, 442 Polypodium scandens G. Forster, see Microsorum scandens,
Polypodium latum, see Campyloneurum brevifolium, 230 386
Polypodium ×leucosporum, 446 Polypodium scandens Labill, see Microsorum pustulatum,
Polypodium levigatum, 447 385
Polypodium lineare, see Lepisorus thunbergianus, 348 Polypodium scolopendria, see Microsorum grossum, 380;
Polypodium linguiforme, see Microsorum linguiforme, 382 Microsorum scolopendria, 387
Polypodium longissimum, see Microsorum longissimum, Polypodium scouleri, 451
382 Polypodium sibericum, 452
Polypodium loriceum, 447 Polypodium sinuosum, see Lecanopteris sinuosa, 346
Polypodium lucidum, see Microsorum lucidum, 382 Polypodium squamulosum, see Microgramma squamulosa,
Polypodium lycopodioides, see Microgramma lycopodioides, 375
373 Polypodium subauriculatum, see Goniophlebium
Polypodium mandaianum, see Phlebodium aureum subauriculatum, 333, color plate 28
‘Mandaianum’, 420 Polypodium tenellum, see Arthropteris tenella, 191
Polypodium maritimum, 447 Polypodium tenuilore, see Microsorum samarense, 385
Polypodium megalophyllum, see Microgramma Polypodium thunbergianum, see Lepisorus thunbergianus,
megalophylla, 373 348
Polypodium menisciifolium, 447 Polypodium thyssanolepis, 451
Polypodium musifolium, see Microsorum musifolium, 383 Polypodium triseriale, 452
Polypodium myriocarpum, see Microsorum longissimum, Polypodium vacciniifolium, see Microgramma
382 vacciniifolia, 375
Polypodium nigrescens, see Microsorum membranifolium, Polypodium verrucosum, see Goniophlebium percussum,
382 332
Polypodium normale, see Microsorum fortunei, 380 Polypodium virginianum, 452
Polypodium palmeri, see Microgramma nitida, 373 Polypodium vitiense, see Microsorum commutatum, 380
Polypodium pectinatum, see Pecluma pectinata, 410 Polypodium vulgare, 92, 452
Polypodium pellucidum, 448 Polypodium vulgare var. occidentale, see Polypodium
Polypodium percussum, see Microgramma percussa, 374 glycyrrhiza, 444
P N I 599
princess pine, see Lycopodium, 113, 354; Lycopodium ‘Ouvrardii’, see Pteris cretica, 475
dendroideum, 357; Lycopodium obscurum, 358 ‘Parkeri’, see Pteris cretica, 475
Prosaptia, 333 ‘Rivertoniana’, see Pteris cretica, 475
Pseudodrynaria, see Aglaomorpha, 176 ‘Roweri’, see Pteris cretica, 475
Psilotum, 113, 470 ‘Victoriae’, see Pteris ensiformis, 95, 101, 476
Psilotum complanatum, 470 ‘Wilsonii’, see Pteris cretica, 101, 475
Psilotum ×intermedium, 470 ‘Wimsettii’, see Pteris cretica, 475
Psilotum nudum, 470 purple cliff brake, see Pellaea atropurpurea, 412
Pteretis, see Matteuccia, 369 Pyrrosia, 480
Pteretis nodulosa, see Matteuccia struthiopteris, 371, color Pyrrosia adnascens, see Pyrrosia lanceolata, 484
plate 33 Pyrrosia angustata, 481
Pteridium, 93, 470 Pyrrosia beddomeana, see Pyrrosia costata, 482
Pteridium aquilinum, 94, 471 Pyrrosia confluens, 482
Pteridium aquilinum var. caudatum, 146, 471 Pyrrosia confluens var. confluens, 482
Pteridium aquilinum var. latiusculum, 146, 471 Pyrrosia confluens var. dielsii, 482
Pteridium aquilinum var. pseudocaudatum, 471 Pyrrosia costata, 482
Pteridium aquilinum var. pubescens, 471 Pyrrosia davidii, see Pyrrosia porosa, 487
Pteris, 472 Pyrrosia dielsii, see Pyrrosia confluens var. dielsii, 482
Pteris altissima, 473 Pyrrosia eleagnifolia, 483
Pteris argyraea, 95, 473, color plate 47 Pyrrosia fallax, 483
Pteris aspericaulis, 95, 474 Pyrrosia hastata, 484
Pteris aspericaulis var. tricolor, 474 Pyrrosia heteractis, see Pyrrosia lingua var. heteractis,
Pteris biaurita, 474 485
Pteris catoptera, 474 Pyrrosia lanceolata, 484
Pteris cretica, 66, 82, 92, 94, 98, 100, 474 Pyrrosia linearifolia, 484
Pteris cretica var. albolineata, see Pteris nipponica, 478 Pyrrosia lingua, 95, 484
Pteris dentata, 475 Pyrrosia lingua var. heteractis, 485
Pteris dispar, see Pteris semipinnata var. dispar, 479 Pyrrosia lingua var. lingua, 485
Pteris ensiformis, 95, 475 Pyrrosia longifolia, 486
Pteris fauriei, 479 Pyrrosia nummulariifolia, 486
Pteris flabellata, see Pteris dentata, 475 Pyrrosia obovata, see Pyrrosia nummulariifolia, 486
Pteris grandifolia, 476 Pyrrosia pekinensis, see Pyrrosia porosa, 487
Pteris longifolia, 476 Pyrrosia piloselloides, 486
Pteris multifida, 82, 97, 98, 101, 478 Pyrrosia polydactyla, 487, color plate 48
Pteris nipponica, 95, 478 Pyrrosia porosa, 487
Pteris quadriaurita, 478 Pyrrosia princeps, 487
Pteris quadriaurita var. argentea, see Pteris argyraea, 473, Pyrrosia rupestris, 488
color plate 47 Pyrrosia samarensis, 488
Pteris quadriaurita var. argyraea, see Pteris argyraea, 473, Pyrrosia serpens, 483
color plate 47 Pyrrosia sheareri, 488
Pteris semipinnata, 479 Pyrrosia splendens, 489
Pteris semipinnata var. dispar, 479 Pyrrosia stigmosa, 490
Pteris serrulata, see Pteris multifida, 478 Pyrrosia tricuspis, see Pyrrosia hastata, 484
Pteris tremula, 82, 93, 94, 100, 479 Pyrrosia varia, see Pyrrosia lanceolata, 484
Pteris tricolor, see Pteris aspericaulis var. tricolor, 474 Pyrrosia cultivars
Pteris tripartita, 480 ‘Cristata’, see Pyrrosia lingua, 485
Pteris vittata, 82, 94, 95, 480 ‘Monstrifera’, see Pyrrosia lingua, 485
Pteris cultivars ‘Nankin-shishi’, see Pyrrosia lingua, 485
‘Albo-lineata’, see Pteris cretica, 474; Pteris nipponica, ‘Nokogiri-ba’, see Pyrrosia lingua, 485
478
‘Childsii’, see Pteris cretica, 475 queen staghorn, see Platycerium wandae, 436
‘Cristata Mayii’, see Pteris cretica, 475 Quercifilix, see Tectaria, 513
‘Distinction’, see Pteris cretica, 475 Quercifilix zeylanica, see Tectaria zeylanica, 517
‘Evergemiensis’, see Pteris ensiformis, 95, 476
‘Mayii’, see Pteris cretica, 475 rabbit’s-foot fern, see Davallia 262; Phlebodium, 419
P N I 601
rainbow moss, see Selaginella uncinata, 505, color plate scented fern, see Paesia scaberula, 409
49 Schaffneria, 494
rasp fern, see Doodia, 289 Schaffneria nigripes, 494
rattlesnake fern, see Botrychium virginianum, 227 Schellolepis, see Goniophlebium, 331
recurved broad-buckler fern, see Dryopteris dilatata Schneider’s rabbit-foot fern, see Phlebodium ×schneideri,
‘Recurved Form’, 307 421
red spike moss, see Selaginella umbrosa, 505 Scleroglossum, 333
Regnellidium, 490 Scott’s spleenwort, see Asplenium ×ebenoides, 198
Regnellidium diphyllum, 490 scouring rush, see Equisetum, Equisetum hyemale, 328
resurrection fern, see Polypodium polypodioides, 449 scurfy sword fern, see Nephrolepis hirsutula, 395
resurrection plant, see Selaginella lepidophylla, 501 Scyphularia, see Davallia, 262
Rhipidopteris peltata, see Elaphoglossum peltatum, 324 Scyphularia pentaphylla, see Davallia pentaphylla, 272
Richard’s holly fern, see Polystichum richardii, 463 Scyphularia pycnocarpa, see Davallia pentaphylla, 272
rigid buckler fern, see Dryopteris mindshelkensis, 314 Scyphularia simplicifolia, see Davallia triphylla, 275
robust male fern, see Dryopteris complexa, 303 Scyphularia sinusora, see Davallia pentaphylla, 272
rock brake, see Cryptogramma, 248 Scyphularia triphylla, see Davallia triphylla, 275
rock felt fern, see Pyrrosia rupestris, 488 scythian lamb, see Cibotium barometz, 245
Rockford holly fern, see Cyrtomium falcatum sea spleenwort, see Asplenium marinum, 199
‘Rockfordianum’, 256 Seemann’s maidenhair, see Adiantum seemannii, 173
rock polypody, see Polypodium virginianum, 452 Selaginella, 91, 95, 97, 114, 495
rock sword fern, see Polystichum scopulinum, 464 Selaginella apoda, 497
rock tassel fern, see Lycopodium squarrosum, 360 Selaginella braunii, 497
rosy maidenhair, see Adiantum hispidulum, 166, color Selaginella caulescens, see Selaginella involvens, 499
plate 9 Selaginella denticulata, 500
rough horsetail, see Equisetum hyemale, 328 Selaginella diffusa, 497
rough maidenhair, see Adiantum hispidulum, 166 Selaginella douglassii, 498
rough sword fern, see Nephrolepis hirsutula, 395 Selaginella emmeliana, see Selaginella pallescens, 503
rough tree fern, see Alsophila australis, 182; Dicksonia Selaginella erythropus, 498
squarrosa, 284 Selaginella harrisii, 499
royal fern, see Osmunda regalis, 408, color plate 6 Selaginella heterodonta, 499
Rumohra, 490 Selaginella involvens, 499
Rumohra adiantiformis, 92, 93, 94, 98, 100, 491 Selaginella kraussiana, 95, 500
Rumohra ‘Davis’, see Rumohra adiantiformis, 491 Selaginella lepidophylla, 501
running pine, see Lycopodium clavatum, 356; Lycopodium Selaginella martensii, 501
complanatum, 357 Selaginella moellendorffii, 501
rusty brake, see Pteris vittata, 480 Selaginella oaxacana, 502
rusty woodsia, see Woodsia ilvensis, 532 Selaginella oregana, 502
rusty-back fern, see Ceterach, 233 Selaginella pallescens, 503
Selaginella plana, 503
Sadleria, 491 Selaginella pulcherrima, 503
Sadleria cyatheoides, 111, 492 Selaginella serpens, 504
Sadleria pallida, 492 Selaginella umbrosa, 505
sago tree fern, see Sphaeropteris medullaris, 512 Selaginella uncinata, 505, color plate 49
Salvinia, 93, 493 Selaginella viticulosa, 505
Salvinia auriculata, 493 Selaginella vogelii, 506
Salvinia minima, 493 Selaginella willdenovii, 506
Salvinia molesta, 494 Selaginella cultivars
Salvinia natans, 494 ‘Albolineata’, see Selaginella martensii, 501
Salvinia rotundifolia, see Salvinia auriculata, 493 ‘Aurea’, see Selaginella kraussiana, 501
saw fern, see Blechnum serrulatum, 221 ‘Brownii’, see Selaginella kraussiana, 501
scale fern, see Ceterach officinarum, 233, color plate 20 ‘Gold’, see Selaginella kraussiana, 501
scaly buckler fern, see Dryopteris remota, 316 ‘Gold-tips’, see Selaginella kraussiana, 501
scaly male fern, see Dryopteris affinis, 298 Selliguea, 439, 507
scaly polypody, see Polypodium thyssanolepis, 451 Selliguea feei, 507
scaly wood fern, see Dryopteris polylepis, 315 sensitive fern, see Onoclea, 402
602 P N I
serpent fern, see Phlebodium aureum, 420 spear-leaved fern, see Doryopteris palmata, 292
serpent’s-tongue fern, see Ophioglossum, 404 Sphaerocionium, see Hymenophyllum, 339
serrate lattice-vein fern, see Thelypteris serrata, 525 Sphaeropteris, 253, 509
shaggy wood fern, see Dryopteris cycadina, 305 Sphaeropteris armaganensis, 510
Shasta holly fern, see Polystichum lemmonii, 459 Sphaeropteris cooperi, 91, 92, 93, 94, 95, 110, 111, 511,
shield fern, see Dryopteris, 296; Polystichum, 454; color plates 2, 3
Thelypteris kunthii, 520 Sphaeropteris excelsa, 95, 511
shining club moss, see Lycopodium lucidulum, 358 Sphaeropteris horrida, 512
shiny shield fern, see Lastreopsis acuminata, 343 Sphaeropteris medullaris, 512
shoestring fern, see Vittaria lineata, 530 Sphaeropteris cultivars
sickle spleenwort, see Asplenium polyodon, 202 ‘Brentwood’, see Sphaeropteris cooperi, 511
Siebold’s wood fern, see Dryopteris sieboldii, 317 ‘Robusta’, see Sphaeropteris cooperi, 511
sierra cliff brake, see Pellaea brachyptera, 413 Sphenomeris, see Odontosoria, 400
sierra water fern, see Thelypteris nevadensis, 520 Sphenomeris chinensis, see Odontosoria chinensis, 400
silverback fern, see Pentagramma, 417; Pityrogramma, Sphenomeris chusana, see Odontosoria chinensis, 400
424 spider brake, see Pteris multifida, 478
silver brake, see Pteris argyraea, 473, color plate 47 spike moss, see Selaginella, 495
silver-dollar fern, see Adiantum peruvianum, 170 spinulose wood fern, see Dryopteris carthusiana, 302
silver fern, see Pityrogramma calomelanos, 425 spleenwort, see Asplenium, 193
silver glade fern, see Deparia acrostichoides, 280 spreading selaginella, see Selaginella kraussiana, 500
silver tree fern, see Alsophila tricolor, 182, color plate 11 squirrel’s-foot fern, see Davallia mariesii var. stenolepis,
silvery felt fern, see Pyrrosia confluens var. dielsii, 482 271, color plate 5; Davallia trichomanoides, 275
silvery spleenwort, see Deparia acrostichoides, 280 staghorn club moss, see Lycopodium clavatum, 356
slender cloak fern, see Cheilanthes bonariensis, 238 staghorn fern, see Platycerium, 427
slender crested male fern, see Dryopteris filix-mas star-scale cloak fern, see Astrolepis, 206
‘Linearis Polydactyla’, 310 Stegnogramma, see Thelypteris, 517
slender lip fern, see Cheilanthes feei, 239 Stenochlaena, 512
slender rock brake, see Cryptogramma stelleri, 251 Stenochlaena palustris, 512
slender tree fern, see Dicksonia squarrosa, 284 Stenochlaena tenuifolia, 512
small-leaved climbing fern, see Lygodium microphyllum, Sticherus, 330
362 Sticherus flabellatus, 330
small-leaved cloak fern, see Argyrochosma microphylla, Sticherus lobatus, 331
190 strap fern, see Campyloneurum, 229
smooth cliff brake, see Pellaea glabella, 415 strawberry fern, see Hemionitis palmata, 338
smooth horsetail, see Equisetum laevigatum, 328 striped brake, see Pteris argyraea, 473, color plate 47
smooth rock spleenwort, see Asplenium fontanum, 199 Struthiopteris, see Matteuccia, 369
snowy club moss, see Selaginella martensii ‘Albolineata’, Struthiopteris pensylvanica, see Matteuccia struthiopteris,
501 371, color plate 33
soft shield fern, see Polystichum setiferum, 464, color swamp fern, see Blechnum serrulatum, 221; Woodwardia
plate 46 areolata, 537
Solanopteris, 440, 508 sweat plant, see Selaginella pallescens, 503
Solanopteris bifrons, 508 sword fern, see Nephrolepis, 389; Polystichum, 454
Solanopteris brunei, 508
southern beech fern, see Phegopteris hexagonoptera, 419 Tagawa’s holly fern, see Polystichum tagawanum, 467
southern fragile fern, see Cystopteris protrusa, 260 tailed bracken, see Pteridium aquilinum var.
southern grape fern, see Botrychium biternatum, 227 pseudocaudatum, 471
southern lady fern, see Athyrium filix-femina var. Tasmanian dicksonia, see Dicksonia antarctica, 282,
asplenioides, 212 color plate 3
southern maidenhair, see Adiantum capillus-veneris, 161 Tasman’s davallia, see Davallia tasmanii, 274
southern polypody, see Polypodium cambricum, 443 tassel fern, see Lycopodium, 354
southern shield fern, see Thelypteris kunthii, 520 Tectaria, 513
southern wood fern, see Dryopteris ludoviciana, 313; Tectaria beccariana, 514
Thelypteris kunthii, 520 Tectaria cicutaria, 515
southwestern brittle fern, see Cystopteris reevesiana, 261 Tectaria decurrens, 514
sparse-lobed grape fern, see Botrychium biternatum, 227 Tectaria gemmifera, 69, 70, 515
P N I 603
walking fern, see Camptosorus, 228 woodrustic fern, see Polystichum dudleyi, 458
Wallich’s wood fern, see Dryopteris wallichiana, 320, Woodsia, 531
color plate 27 Woodsia alpina, 532
wall-rue, see Asplenium ruta-muraria, 203 Woodsia glabella, 532
water clover, see Marsilea, 365 Woodsia ilvensis, 532
water fern, see Ceratopteris 232; see also Marsilea Woodsia manchuriensis, 533
schelpeana, 368; Thelypteris nevadensis, 520 Woodsia obtusa, 92, 98, 533
water spangle, see Salvinia, 493 Woodsia oregana, 98, 534
water sprite, see Ceratopteris, 232 Woodsia polystichoides, 534
water tassel fern, see Lycopodium squarrosum, 360 Woodsia scopulina, 535
wax-dot maiden fern, see Thelypteris resinifera, 524 Woodsia subcordata, 535
western bracken, see Pteridium aquilinum var. pubescens, Woodwardia, 535
471 Woodwardia areolata, 539
western cliff fern, see Woodsia oregana, 534 Woodwardia chamissoi, see Woodwardia fimbriata, 537
western five-finger fern, see Adiantum aleuticum, 160 Woodwardia fimbriata, 537
western holly fern, see Polystichum scopulinum, 464 Woodwardia martinezii, 538
western oak fern, see Gymnocarpium disjunctum, 335 Woodwardia orientalis, 69, 70, 537
western polypody, see Polypodium hesperium, 445 Woodwardia radicans, 65, 69, 95, 537, color plate 50
western sword fern, see Polystichum munitum, 460 Woodwardia ×semicordata, 538
West Indian tree fern, see Cyathea arborea, 253 Woodwardia spinulosa, 538
whisk fern, see Psilotum, 470 Woodwardia unigemmata, 539
white-rabbit’s-foot fern, see Davallia tyermannii, 276 Woodwardia virginica, 92, 95, 539
Whiteside’s crisped broad-buckler fern, see Dryopteris woolly lip fern, see Cheilanthes tomentosa, 243
dilatata ‘Crispa Whiteside’, 307 woolly tree fern, see Dicksonia fibrosa, 282
widespread maiden fern, see Thelypteris kunthii, 520 Wright’s lip fern, see Cheilanthes wrightii, 243
winged beech fern, see Phegopteris decursive-pinnata, 418
winter grape fern, see Botrychium biternatum, 227 yellow-golden-scaled male fern, see Dryopteris affinis, 298
winter scouring rush, see Equisetum hyemale, 328
wood fern, see Dryopteris, 296; see also Thelypteris, 517 Zygophlebia, 333
woodland bladder fern, see Cystopteris protrusa, 260