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Pteridophytes (Ferns)

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Pteridophytes (Ferns)
George Yatskievych, Missouri Botanical Garden, St Louis, MO, USA
Pteridophytes (vascular cryptogams or ferns and fern allies) comprise about 12 000 species
of primitive vascular plants; they do not produce flowers or seeds and reproduce instead via
spores. They occur in most terrestrial habitats and also in some aquatic communities. Some
species are very beneficial to humans, but the group also contains important species of
weeds.
Introduction
Pteridophytes, also known as ‘vascular cryptogams’ and
‘ferns and fern allies’, comprise about 12 000 species of
vascular plants that do not produce flowers or seeds,
reproducing instead via the production of spores. Pter-
idophytes occur in most terrestrial habitats on earth and
are also present in some aquatic communities. They are an
important part of the ground vegetation in many forest
communities and, with about one-third of the species
growing on the trunks and branches of trees, they are also
an important component of many epiphytic plant com-
munities. Some species are very beneficial to humans, but
the group also contains some of the most important weed
species in the world.
Life Cycle
Pteridophytes are characterized by a life cycle that usually
involves an alternation of two free-living generations –
sporophyte and gametophyte – with the sporophyte the
larger phase of the life cycle. Nonvascular plants like
mosses and liverworts also have an alternation of genera-
tions, but in these organisms the gametophyte generation is
generally the dominant phase. In seed plants, the
gametophyte is no longer free-living but remains enclosed
in tissues on the sporophyte and there is a progressive
reduction in the size through various gymnosperm groups
such that in flowering plants (angiosperms) the gameto-
phyte generation is reduced to just a few cells in the
germinating pollen grains and the ovules.
The conspicuous phase of the pteridophyte cycle is the
sporophyte, which is how most people observe the plants in
nature. These are usually perennial. Sporangia are
produced on the leaves of sporophytes (sometimes in
specialized cone-like strobili). In true ferns, these are
commonly on the leaf undersurface and are often clustered
into discrete units called sori. Within each sporangium,
specialized cells undergo a series of mitotic (structural)
divisions followed by meiosis (sexual division) that results
in production of spores with half as many chromosomes as
in the original sporophyte. The more advanced ferns
ENCYCLOPEDIA OF LIFE SCIENCES © 2002, John Wiley & Sons, Ltd. www.els.net
Introductory article
Article Contents
. Introduction
. Life Cycle
. Reproductive Variations
. Cytology
. Morphology and Anatomy
. Systematics and Classification
. Economic Importance
usually have 64 spores per sporangium, but more primitive
ferns and fern allies may have hundreds or even thousands.
At maturity, the sporangium dries and ruptures, dispersing
the spores into the air.
When a spore lands on a suitable substrate, it
germinates, the cells dividing and forming first a filament
and eventually usually a heart-shaped gametophyte (some-
times other shapes in some groups) that is the same species
as the sporophyte but appears very different. Gameto-
phytes are often moss-like in appearance and are quite
small, usually less than 1 cm wide at maturity, but are often
fairly easily located in nature near adjacent sporophytes.
Although in a few genera gametophytes can be long-lived,
in most ferns their lifespan is usually much less than a year.
They are the sexual phase of the life cycle in that they
produce multicellular sex organs at maturity on the side
away from the light. The more or less spherical antheridia
(male gametangia) are produced among the rhizoids
towards the base of the plant and at maturity they pop
open to release motile flagellated spermatozoids. Arche-
gonia (female gametangia) are usually produced at the
opposite end near the notch region, and are flask-shaped
structures containing a single egg cell. A film or droplet of
free-standing water is necessary in order for the sperma-
tozoids to swim to an archegonium of the same or a
different gametophyte. The neck cells of the archegonium
spread at maturity and the spermatozoid swims down the
archegonial canal to fuse with the egg, effecting syngamy
(fertilization) and forming a zygote with twice the number
of chromosomes as the gametophyte. This zygote grows
and develops into a new sporophyte, completing the cycle,
while the maternal gametophyte withers away.
In a typical pteridophyte, each gametophyte is poten-
tially bisexual, producing both antheridia and archegonia.
Because the eggs and spermatozoids of an individual all
grew from a single spore and are thus genetically identical,
potentially these plants can become self-fertilized in a way
that renders the resultant sporophyte entirely homozygous
(having only one kind of allele for each gene locus) for its
entire makeup. Various mechanisms exist to promote
cross-fertilization: the gametangia often mature at differ-
ent times; the genome may have deleterious alleles that are
1
 Pteridophytes (Ferns)
fatal to homozygous individuals (genetic load); and there
may exist mating factors that prevent successful self-
fertilization. Some ferns also produce pheromones known
as antheridiogens, in which the first spore to germinate at a
site becomes a female gametophyte and exudes a substance
causing later-germinating spores to develop into male
gametophytes.
Reproductive Variations
Unusual gametophytes
In some ferns and fern allies, including some clubmosses
(Lycopodiaceae), whisk ferns (Psilotaceae), and grape
ferns (Ophioglossaceae), the gametophytes are not surface-
dwelling and green. Instead, they are subterranean and
nonphotosynthetic, often appearing as pale brown or
yellow fuzzy cylinders or pads of tissue. These gameto-
phytes are mycotrophic; that is, they receive their nutrients
from soil-borne fungi that establish connections with their
rhizoids. However, although such gametophytes are
usually slow-growing, they usually produce normal
gametangia and otherwise complete their life cycles in the
typical fashion.
Heterospory
Other ferns and fern allies, including spike mosses
(Selaginellaceae), quillworts (Isoetaceae), and aquatic
ferns (Azollaceae, Marsileaceae, Salviniaceae), depart
from the typical life cycle in producing two different types
of sporangia. One of these produces numerous microscopic
microspores that germinate to produce male gameto-
phytes. The other sporangial type produces many fewer
and much larger megaspores (usually visible to the naked
eye), which grow into female gametophytes. In both spore
types the gametophytes are reduced in structure and
develop mostly within the ruptured spore wall.
Vegetative reproduction
Many pteridophytes supplement their sexual cycles with
various forms of vegetative reproduction. This may be as
simple as the fragmentation of a creeping rhizome into
smaller pieces that become established as separate plants.
Horsetails (Equisetaceae) growing along rivers and
streams are frequently spread over long distances in this
fashion by flooding. Other species develop specialized
structures to effect vegetative propagation. Some ferns
produce stolons, which are specialized long, spreading
stems that root at their tips and form new plants. Others
produce buds or bulbils on their leaves that can germinate
to form new plantlets. Still others produce roots where
their fronds come into contact with soil. A few species
2
produce specialized underground structures, such as tubers
and similar offsets. Some species have sporophytes or
gametophytes that produce gemmae, which are specialized
relatively undeveloped fragments of plants that break off
and are dispersed, eventually germinating to form new
plants. In a few species of filmy ferns (Hymenophyllaceae),
shoestring ferns (Vittariaceae), and other families, the
ability to produce sporophytes has been lost and the plants
exist only as colonies of gametophytes spreading through
the production of tiny air-dispersed gemmae.
Apogamy
Apogamy is a widespread and important mechanism of
reproduction in pteridophytes more than in any other
group of plants. Apogamous ferns, which frequently occur
in environments with seasonal extremes of heat, cold and/
or drought, avoid the necessity for sex. In the sporangia of
such plants, a mechanism during the series of cell divisions
results in the production of spores with the same genetic
constitution as the sporophyte plant (meiosis does not
result in a reduction in chromosomal ploidy). These
‘diplospores’ grow into gametophytes that produce new
sporophytes directly from meristematic tissues near the
notch region. The environmental advantages of apogamy
include the faster development of the gametophyte and the
release from the requirement of standing water for
fertilization to take place. Interestingly, many apogamous
ferns continue to produce antheridia with functional
spermatozoids, which can be released and fertilize eggs
on nearby gametophytes of related sexual species. Once
formed, such hybrids are always apogamous and thus able
to reproduce themselves.
Cytology
Pteridophytes characteristically have high chromosome
numbers. The highest chromosome number recorded for
any organism is in an adder’s tongue fern, Ophioglossum
reticulatum, with about 1260 pairs of chromosomes. The
base chromosome number (x) in various genera is quite
variable: for example, Asplenium (x 5 36), Botrychium
(x 5 45), Osmunda (x 5 22), Pellaea (x 5 29), Polystichum
(x 5 41); and Pteridium (x 5 52). Exceptions occur in a few
ferns, including most aquatic genera (Salvinia, x 5 9).
Some fern allies also have low chromosomal base numbers
(Selaginella, x 5 mostly 7–10).
Several theories have been advanced as to why ferns have
so many chromosomes. Among the most intriguing is that
of palaeopolyploidy. Botanists have long known that
polyploidy (the development of extra sets of chromosomes
over the basic diploid level) is very widespread and
common in pteridophytes. Both autopolyploidy and
allopolyploidy have been documented in numerous genera.

Allopolyploidy involves hybridization between species,
resulting in a sterile hybrid that regains its fertility by
doubling its chromosome number during spore production
in some sporangia. Autopolyploidy involves the same
doubling of chromosome number during spore production
but without a hybridization event. With polyploidy so
pervasive among pteridophytes, geneticists have long been
puzzled that most fern species are apparently functionally
diploid, having only two alleles for each gene locus. In the
last two decades, ‘gene-silencing’ – the selective shutting off
of duplicate copies of genes found in polyploid species –
has been documented in several fern genera. This has given
rise to the hypothesis that, over time, ferns undergo regular
rounds of polyploidy (which increases their chromosome
numbers) followed by gradual diploidization of the
genomes (selective silencing of the extra gene copies to
return the species to a functionally diploid level). Evidence
for this mechanism is circumstantial, but it seems likely to
function in at least those cases that have been better studied
thus far.
Morphology and Anatomy
Sporophytes
Stems
Most ferns have specialized stems called rhizomes that are
positioned at the level of the substrate or somewhat buried.
Rhizomes vary greatly in size, thickness and orientation.
Most commonly, they are horizontal and creeping, but
many species have short upright rhizomes. In some groups,
notably the tree ferns, specialized stems are trunk-like and
may be 20 m or more tall. These modified stems produce
only adventitious roots and are usually covered with dense
scales or hairs, at least towards the growing tip.
Other types of stems occur in some primitive ferns and in
most fern allies. Grape ferns (Ophioglossaceae) usually
have somewhat tuberous stems. Horsetails (Equisetaceae)
have both rhizomes and fluted or ridged aerial stems.
Quillworts (Isoetaceae) have very short stout stems with
the nodes very close together (corms). Most clubmosses
(Lycopodiaceae) have relatively unspecialized stems
Rhizomes are structurally simpler than those of most
seed plants in that they do not produce secondary growth
(wood). Even the tree ferns have only primary growth, and
a thick mantle of interwoven roots is produced to help with
structural support. In most ferns, the vascular system of the
stem is in the form of a hollow cylinder interrupted (with
gaps) where traces branch off to the leaves. In cross-
section, most fern rhizomes thus appear as an irregular ring
of vascular bundles. In some primitive ferns and fern allies,
the vascular system is a solid uninterrupted cylinder.
Pteridophytes (Ferns)
Leaves
Pteridophytes exhibit an amazing variety of leaf morphol-
ogies. In most fern allies and a few primitive ferns, the
leaves are reduced and scale-like, needle-like or grass-like,
with at most a single vein. In most true ferns, however, the
leaves are the dominant organ of the sporophyte and can be
extremely complex in their pattern of division. In a few
genera (especially in the Gleicheniaceae), the leaves are
indeterminate in growth; that is, they continue to elongate
at the tip, often reaching several metres in length and
clambering over surrounding vegetation. The petiole
(stipe) of fern leaves may be circular, angled, or U-shaped
in cross-section, and is sometimes hairy or scaly. There are
one to several vascular strands, and the number and
position of these in the petiole are often diagnostic for
individual families or genera.
In most ferns, the development of the leaf follows a
pattern known as ‘circinate vernation’. This produces a
characteristic fiddlehead or crozier as the leaf uncurls. In a
few genera, this pattern has become modified so that the
unfurling leaf produces a hook-like structure. The fern
allies and the grape ferns (Ophioglossaceae) do not exhibit
circinate vernation but expand by unfolding or in an
indefinite pattern.
The leaf blade (lamina) varies from entire to highly
divided, with pinnate, pedate and palmate patterns of
division in various species, but most commonly is one or
more times pinnately compound. The continuation of the
petiole as the central axis of the leaf blade is known as the
rachis, to which the pinnae (primary divisions or leaflets)
are attached. The pinnae may themselves be entire or one
or more times compound. The ultimate divisions of the leaf
are called pinnules, which may be entire or lobed. Venation
of the leaves can be quite complex, with several orders of
successively finer midveins (costae) and lateral veins. The
venation of the pinnules may be unbranched or branched
with free to variously anastomosing veinlets.
Fern leaves may be glabrous or variously covered with
hairs and/or scales. In some species, the leaves are
glandular and sticky. Other species secrete a powdery
farina, usually on the leaf undersurface, which may be
white or bright yellow or orange. The leaves also vary
greatly in thickness and texture. The thinnest leaves occur
in the filmy ferns (Hymenophyllaceae), in which the leaves
are often only two cell layers thick. The production of
thick, leathery leaves or leaves with dense vestiture of hairs,
scales, glands or farina is generally explained as adaptation
to droughty habitats and/or areas of high sunlight.
Sporangia
In fern allies and a few primitive ferns, relatively large
sporangia are produced either in complex cone-like strobili
at the stem or branch tips or in the axils at the bases of
leaves. Some fern species produce dimorphic leaves, with
vegetative (trophophyll) and fertile (sporophyll) leaves
3
 Pteridophytes (Ferns)
having different morphologies. In other ferns, the leaf is
divided into specialized fertile and vegetative regions.
However, in most ferns, the sporangia are produced on the
undersurface of normal leaves.
The positional patterns and other details of sporangia on
the leaf undersurface are often diagnostic for particular
families or genera and are a principal tool in fern
classification. At one extreme, the sporangia may entirely
cover the leaf undersurface (acrostichoid). In contrast, in
some primitive ferns, the sporangia are sparsely scattered
along some veins. In most ferns, however, the sporangia
are grouped into discrete lines or clusters known as sori.
Sori may be circular to linear, positioned along the margin
or towards the midvein (costa), surficial or in a groove or
channel, etc. In some cases, the developing sori are
protected by a recurved leaf margin (false indusium), a
covering of deciduous scales, or a more permanent small
flap of tissue, the indusium. Indusia vary greatly in shape,
size, texture and persistence, ranging from umbrella-
shaped to globose to linear. In the water ferns (Azollaceae,
Marsileaceae, Salviniaceae), the sporangia become en-
closed in hardened capsular structures called sporocarps
that are formed either from modified leaflets or from
modified indusia.
The sporangia themselves are usually positioned on a
somewhat thickened vein ending or along a portion of a
vein. In most cases, the sporangium consists of a stalk, of
varying length and cell number, and a multicellular
capsule. In most ferns, the capsule is differentiated into
thin-walled cells and an annulus, a ring or region of cells
with only some of the walls thickened. The annulus
functions in spore release.
Spores
Spores are the main structures by which ferns are dispersed
to form new populations. As such, in most ferns, they are
relatively impervious, long-lived and metabolically inac-
tive. Although the majority of spores produced fall within a
few metres of the parental sporophyte, the spores of some
ferns have been recovered from air currents in the
stratosphere during high-elevation atmospheric sampling
studies and ferns are among the most successful colonists
of highly isolated oceanic islands. Although the life of most
spores is measured in terms of months or a few years, in
some cases fern spores have been induced to germinate
after more than a hundred years of storage. In a few groups
scattered throughout the ferns and fern allies, the spores
are relatively thin-walled, green and photosynthetically
active, and relatively short-lived, reflecting an adaptation
to rapid establishment of new plants following dispersal.
Developmentally, spores are the direct products of
meiosis, which begins with a single spore mother cell
undergoing two separate rounds of division, and yields a
tetrad of products that breaks apart into four individual
4
spores. The spatial relationship between the plane of
division of the two rounds of meiotic division affects the
shape and markings of the resulting spores. Two main
types are recognized. Trilete (tetrahedral) spores vary from
nearly spherical to somewhat three-angled and have a
three-branched scar where each spore was attached to the
others in the tetrad. Monolete (bilateral) spores are
ellipsoid to bean-shaped and have a linear attachment
scar along one side. The attachment scar is usually where
the spore ruptures during germination. As it matures, each
spore develops two or three outer protective layers, the
relatively thin endospore, the thick perispore, and in some
cases an outermost exospore. The exospore and also
portions of the perispore actually develop from materials
produced by the inner sporangium wall and deposited over
the spore, rather than by the spore itself.
Mature spores vary greatly in size and surface sculptur-
ing. Spores of some Marattiaceae are only about 15 mm in
diameter, whereas the megaspores of some Selaginella
species may approach 1 mm. The surface sculpturing is
often diagnostic for various species, genera and/or
families, ranging from smooth to wrinkled, spiny and/or
with wing-like ridges.
Gametophytes
Upon germination of spores, cell divisions produce first a
filamentous structure. In most ferns, this subsequently
continues to divide in two or more planes and eventually
differentiates into the mature gametophyte. The typical
fern gametophyte is a flat, heart-shaped structure, with two
lobes and an intervening notch at one end and the other end
narrowed or rounded. It can vary in size from a few
millimetres to about 1 cm in size. A number of variations
exist but are not widespread, including filamentous, strap-
shaped and irregularly lobed gametophytes. In pterido-
phytes with subterranean mycorrhizal gametophytes, these
can mature to various shapes, but most are either tubular
or cushion-shaped.
Gametophytes are moss-like in that they lack vascular
tissue and roots. Slender hair-like structures called rhizoids
function to absorb water and nutrients and act to anchor
the gametophyte to the substrate. The gametangia (sex
organs) generally are formed on the side of the gameto-
phyte away from the light (except in subterranean
gametophytes). The antheridia are positioned among the
rhizoids and are more or less spherical structures consisting
of a jacket of cells enclosing the spermatozoids. The
archegonia are usually positioned near the notch on a
slightly thickened pad of tissue. They are flask-shaped and
somewhat sunken into the tissue. The neck of the
archegonium consists of four columns of cells that separate
at maturity, opening a canal and exposing the egg cell in the
base for fertilization by the spermatozoid.

Systematics and Classification
Classification of pteridophytes remains somewhat con-
troversial. The terms ‘pteridophytes’, ‘ferns and fern allies’
and ‘vascular cryptogams’ continue to be used informally
by botanists who wish to avoid becoming enmeshed in the
technical details of competing systems of fern classifica-
tion. Although many of the groups of ferns and fern allies
are distinctive and have been recognized since antiquity,
the relationships among these groups and the taxonomic
level at which they should be recognized still has not been
fully resolved. In recent years, a consensus has begun to
emerge, and molecular phylogenetic studies involving
mostly the comparison of various gene sequences have
helped to refine theories of pteridophyte evolution and
taxonomy.
In general, pteridophytes have a long fossil record and
the main lineages trace their origins to the first vascular
land plants. Pteridophytes were dominant plants in the
swamps of the Carboniferous Period more than 300 million
years ago, which gave rise to the world’s major coal
deposits. In some groups, such as the horsetails (Equise-
taceae), the relatively few modern species are the remnants
of formerly much more diverse lineages. On the other hand,
some modern fern species have existed for long times, as
shown by the fossils indistinguishable from the modern
sensitive fern (Onoclea sensibilis) dating back to the
Palaeocene Epoch more than 60 million years ago.
In recent years, a fundamental shift in our under-
standing of primitive pteridophyte classification has
occurred. Traditionally, three or four main groups had
been recognized. These included the clubmosses and
related groups (lycophytes), the horsetails (sphenophytes,
sometimes called arthrophytes), the true ferns (filicaleans),
and sometimes the whisk ferns (psilophytes). The psilo-
phytes, an unusual group with structurally relatively
simple plants, were considered the most primitive group
of extant vascular plants by some botanists and true ferns
with reduced simplified structure by others. Recent
anatomical and molecular studies have shown that the
latter interpretation is probably correct – the Psilotaceae
are primitive ferns whose stems, leaves and sporangia have
become simplified over time. These same studies have
yielded an even more fundamental conclusion. The
lycophytes are apparently the most primitive group of
extant vascular plants. The lineage leading to the seed
plants (gymnosperms and angiosperms) has its origins
within the pteridophyte lineage before the divergence of
both the true ferns and the horsetails. Thus, the most recent
hypothesis of pteridophyte evolution advocates the ex-
istence of two fundamental groups, the lycophytes and the
remaining ferns and fern allies.
There are a number of relatively primitive fern families,
many of which are represented by relatively few modern
species but some of which have extremely long fossil
records. The greatest diversity of modern species exists
Pteridophytes (Ferns)
among the most advanced fern groups, and the number of
families to be accepted and the relationships among these
families are the topic of intensive systematic research at
present.
Table 1 summarizes current hypotheses concerning
pteridophyte classification, from most primitive to most
advanced. An estimate of the number of extant genera and
species, in parentheses, follows each family name, and also
the common names of selected well-known examples.
Economic Importance
Relatively few species of pteridophytes are economically
important. Perhaps the best-known current use is horti-
cultural, as garden plants, house plants and specimen
plants in conservatories and greenhouses. One species,
Ruhmora adiantiformis is often called florist’s fern; its finely
divided but thick and leathery leaves resist wilting and are
used in cut flower arrangements. Another horticultural
practice has been the use of chunks of the dense rot-
resistant root mantles covering the stems of tree ferns
(known as orchid bark) as a substrate for growing orchids
and other plants that are epiphytic in nature. However, this
has caused the decline and endangerment of numerous tree
fern species, with the result that commercial trade in tree
fern products is now strictly regulated by international law
and trade treaties.
A number of ferns have been used in handicrafts.
Petioles of some members of the climbing fern family,
Schizaeaceae, as well as other groups, are used in some
tropical countries for colour designs in basketry and
bracelets. Pteridium (bracken) leaves have been used to
make a green dye. The rhizomes of the tree fern Cibotium,
which are covered with dense, long, golden hairs, have been
fashioned since antiquity into animal statue curios some-
times known as ‘vegetable lamb of Tartary’.
One group of pteridophytes with an extensive history of
use is the clubmosses (Lycopodiaceae). The microscopic
spores of these fern allies contain nonvolatile oils that
made them useful as dry industrial lubricants. They have
also been used to keep latex items such as surgical gloves
and condoms from sticking together, but this practice has
been mostly discontinued since it was discovered that the
spores caused skin irritation and allergic reactions in some
people. Other uses of the spores have been in flash powder
for photography and in fingerprint powder used in forensic
investigation.
Various ferns are also eaten as food, with the young
foliage usually steamed as a vegetable or dried and used as
an additive in stews and sauces. Several species are eaten,
including Diplazium esculentum (which is cultivated for this
purpose in parts of Asia), but the commercially most
important species in the western hemisphere is Matteuccia
struthiopteris, the ostrich fern, whose fiddleheads are a
5
 Pteridophytes (Ferns)

Table 1 Summary of the classification of extant pteridophyte families

Lycophytes (Division Lycopodiophyta) (4 extinct orders 1 3 extant orders (each with 1 family))
Lycopodiaceae (4/380), clubmosses
Isoetaceae (1/130), quillworts
Selaginellaceae (1/800), spikemosses
Ferns (Division Pteridophyta)
Eusporangiate ferns (3 extant orders, each with 1 family)
Psilotaceae (2/12), whisk ferns
Ophioglossaceae (3–8/80), grape ferns
Marattiaceae (4/100), giant ferns
Sphenophytes (3 extinct orders 1 Equisetales)
Equisetaceae (1/15), horsetails
Leptosporangiate ferns (1 extinct order 1 about 10 extant orders)
Primitive isolated groups (6 orders (each with 1 modern family except that Cheiropleuriaceae is in Dipteridales))
Osmundaceae (3/20), cinnamon and royal ferns
Hymenophyllaceae (2 or 3/650), filmy ferns
Stromatopteridaceae (1/1)
Gleicheniaceae (4/150), scrambling ferns
Cheiropleuriaceae (1/1)
Dipteridaceae (1/8)
Schizaeaceae (5/200), climbing ferns, curly grass fern
Heterosporous aquatic groups (Order Marsileales, also called Hydropteridales)
Marsileaceae (3/70), water clovers
Azollaceae (1/6), mosquito ferns
Salviniaceae (1/11), water spangles
Tree ferns (Order Cyatheales)
Loxomataceae (2/2)
Plagiogyriaceae (1/11)
Matoniaceae (2/4)
Metaxyaceae (1/2)
Dicksoniaceae (5/28–33), tree ferns
Lophosoriaceae (1/1)
Hymenophyllopsidaceae (1/8)
Cyatheaceae (14–20/620–675), tree ferns
Advanced groups (Order Polypodiales)
Lindsaeaceae (5/200)
Dennstaedtiaceae (14/350), cup ferns, hay-scented fern, bracken
Pteridaceae (35–45/1150), cliff brakes, lip ferns, goldback ferns, maidenhair ferns, shoestring ferns
Aspleniaceae (1/800), spleenworts
Thelypteridaceae (1–35/900)
Blechnaceae (8/250), chain ferns
Dryopteridaceae (including Tectariaceae) (30/950), wood ferns, shield ferns, halberd ferns, ostrich fern, sensitive fern
Woodsiaceae (including Athyriaceae) (10–13/500), fragile ferns, lady ferns, oak ferns
Lomariopsidaceae (6/550), paddle ferns, tongue ferns
Davalliaceae (14/120), rabbit’s foot ferns, Boston fern
Polypodiaceae (including Grammitidaceae) (35–45/1050), polypodies, staghorn ferns

common sight in markets of the northeastern United States
in late spring. Formerly, Pteridium aquilinum (bracken)
was quite important in some cuisines, particularly in parts
of eastern Asia. However, medical studies have linked this
species to stomach cancer and its use has declined.
Perhaps the most economically valuable species of
pteridophyte is Azolla, a genus of tiny floating aquatic
ferns. For centuries farmers in parts of eastern Asia
jealously guarded strains of this plant, which they used to
inoculate rice paddies in the spring for markedly increased
yields. During the Vietnam War era, this practice came to

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the attention of western scientists. They ‘discovered’ that
hollow chambers in Azolla leaves contain symbiotic
cyanobacteria (Anabaena azollae) that are able to convert
atmospheric nitrogen into the nitrate form that serves as a
major plant nutrient. Thus, the fast-growing plants of
Azolla acted as a living source of fertilizers. During the past
few decades, millions of dollars have been spent to locate
superior strains of this fern and to make the process more
efficient, in an attempt to increase rice production in
developing countries.
A few ferns have had negative economic impacts because
of their weediness. Two of the best examples include
Salvinia and Pteridium. Salvinia molesta (Kariba weed,
giant salvinia) is a floating aquatic fern that is weedy
throughout the warmer parts of the world. In some
situations it can form a mat several inches thick on the
surface, that prevents light and oxygen penetrating into the
water. In places such as New Guinea, this fern has at times
threatened to destroy local fishing economies, and it is
being carefully eradicated where found outside of its
natural range in southern Brazil. Pteridium aquilinum
(bracken) is a coarse fern with an immense creeping
rhizome capable of reaching lengths of 400 m. The plant
quickly invades open habitats, competing vigorously with
other plants. Because the plants are toxic to livestock,
bracken has ruined the pasturage on large acreages of land,
especially in parts of Europe.
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Pteridophytes (Ferns)
Further Reading
Camus JM, Gibby M and Johns RJ (eds) (1996) Pteridology in
Perspective. Kew, UK: Royal Botanic Gardens.
Galston AW (1975) The water fern–rice connection. Natural History
84(12): 10–11.
Hoshizaki BJ and Moran RC (2001) Fern Grower’s Manual, revised edn.
Portland, OR: Timber Press.
Kramer KU and Green PS (eds) (1990) Pteridophytes and Gymnosperms;
vol. 1 in Kubitzki K (ed.) The Families and Genera of Vascular Plants.
Berlin: Springer-Verlag.
May LW (1979) The economic uses and associated folklore of ferns and
fern allies. Botanical Review (Lancaster) 44: 491–528.
Perring FH and Gardiner BG (eds) (1976) The biology of bracken.
Botanical Journal of the Linnaean Society 73: 1–302.
Pryer KM, Schneider H, Smith AR et al. (2001) Horsetails and ferns are a
monophyletic group and the closest living relatives to seed plants.
Nature 409: 618–622.
Sheffield E, Wolf PG and Haufler CH (1989) How big is a bracken plant?
Weed Research 29: 455–460.
Tryon RM and Tryon AF (1982) Fern and Allied Plants, with Special
Reference to Tropical America. New York: Springer-Verlag.
Wolf PG (ed.) (1995) Use of molecular data in evolutionary studies of
pteridophytes. American Fern Journal 85: 101–428.
Wolf PG, Sipes SD, White MR et al. (1999) Phylogenetic relationships of
the enigmatic fern families Hymenophyllopsidaceae and Lophosor-
iaceae: evidence from rbcL nucleotide sequences. Plant Systematics
and Evolution 219: 263–270.
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