Division Pteridophyta

■ The Pteridophyta are the ferns. The

ferns are vascular plants with a
dominant sporophyte plant body, a
predominantly leafy anatomy and
megaphylls.
■ Ferns have been prevalent since the
Middle Devonian and are believed to
include 10,000 species.
Bold proposes the following
phylogeny for ferns:
Class 1: Ophioglossopsida
Order: Ophioglossales (eusporangiate ferns)

Class 2: Marattiopsida (eusporangiate ferns)

Order Marattiales

Class 3: Filicopsida (leptosporangiate ferns)

Order: Filicales
Order: Marsileales
Order: Salviniales
Class Ophioglossopsida
These ferns are eusporangiate. Their
sporangia arise from several cells. These
sporangia are large, sessile or have massive
stalks, and produce a large number of
spores.

Ophioglossum and Botrychium are examples of

the Ophioglossales.
Vegetative Morphology
Both of these ferns are common in temperate
North America. Ophioglossum is commonly
called the adder’s tongue fern. Botrychium is
commonly called the grape fern.

Both ferns produce a single leaf each year from

an underground stem. The leaf of
Ophiglossum is entire. The leaf of
Botrychium is dissected. (Different species
have different degrees of dissection.)
Ophioglossum has
an entire leaf.
The leaf of
Botrychium is
dissected.
Ophioglossum
Ophioglossum,
adder’s tongue
fern
Ophioglossum with mature fertile spikes
Ophioglossum,
whole plants
Two species of
Botrychium

corm

Dehiscent
sporangium
Botrychium
 Botrychium
virginianum,
rattlesnake
fern
Botrychium
dissectum
Botrychium
multifidum
Stems
■ The underground stems of both are
erect, fleshy (a corm) and slow growing.
The development of the stem occurs by
divisions of a single apical cell.
Botrychium is the only extant fern genus
with secondary vascular tissue.
Leaves
■ Leaves in these two genera appear as
primordial near the slow-growing stem
tip several years before they are raised
above the ground, and their
development is extremely slow.
Roots
■ The roots of both genera develop from
single apical cells. Roots contain a well
developed pericycle and endodermis.
The epidermal cells lack root hairs.
Endophytc fungi are associated with the
roots.
Root tip of Botrychium

Apical cell
Spore production
Ophioglossum and Botrychium are the only two
genera of ferns which produce sporangia on
special fertile spikes. These spikes arise at
the junction of the leaf blade and the petiole.
These are interpreted anatomically as a pair
of fertile, lateral leaflets. The fertile spike is
unbranched in Ophioglossum and branched
in Botrychium. Sporangia are large.
Thousands of spores are produced by each
sporangium.
 Ophioglossum

Fertile
spike
Botrychium fertile
spikes
Gametophyte
In both genera, the gametophyte is
fleshy, non-green, subterranean and
contains an endophytic fungus. Their
nutrition is saphrophytic. Gametangia
arise from superficial cells of the
gametophyte, but at maturity are mostly
sunken in the gametophytic tissue.
Embryonic sporophyte

In Ophioglossum, the young sporophyte

begins with the differentiation of a foot,
root and primordial leaf. The stem
apical cells appear later.
Class Marattiopsida
This class contains one order, the
Marattiales. These plants are
exclusively tropical. Two genera are
representative of the seven in this order,
Angiopteris and Marattia.
Angiopteris evecta
Angiopteris evecta
Marattia attenuata
Vegetative Morphology
The fronds are several times pinnately
compound. The young leaves are
circinately vernate, or curled in the bud.
Curled fern buds are also called croziers.
The fronds in some species are several
meters in length.
The stems are fleshy, tuberous and erect.
The stems are covered by the persistent
paired stipules at each leaf base. The stem
develops from a single apical cell.
Fronds of
Angiopteris
smithi
Angiopteris leaf, close up
Spore production
Sporangia in Angiopteris are located on the
lower surfaces of leaflets near the leaf
margins along small veins in rings of 10 to 20.

Each sporangium of Angiopteris dehisces into

two valves or halves. The sporangia of
Marattia are united into a unit called a
synangium. The entire synangium dehisces
along a single suture to produce two valves.
Angiopteris
sporangia
Marattia synangia, along leaflet margins
Gametophyte

The gametophytes are large and

liverwortlike, similar to Pellia.
Gametangia are embedded in the
gametophyte tissue. Sperm are
multiflagellated.
Major differences between the
Ophioglossopsida and Marattiospsida

1) The sporangia in the Ophioglossopsida are

located on fertile spikes. In the Marattiopsida
they are on the lower surface near the
margins of vegetative leaflets.

2) The gametophytes of the Ophioglossopsida

are subterranean, non-green. The
gametophytes of the Marattiopsida are above
ground, liverwortlike.
Major similarities

1) Both are eusporangiate ferns.

2) Both have megaphylls.

3) The leaf is the dominant organ of the

sporophyte.
Class Filicopsida
Orders: Filicales
Marsileales
Salviniales
Filicopsida
■ The vast majority of the representatives
of this class belong to the order
Filicales. These are homosporous,
leptosporangiate ferns. Most of the
ferns found in the temperate regions of
North America are in a single family of
the Filicales, the Polypodiaceae.
■ Leptosporangiate sporangia are small,
arise from a single initial cell and
produce a definite number of spores,
usually fewer than in a eusporangium.
Order Filicales, Family
Polypodiaceae
■ As in the eusporangiate ferns, the
leaves of the leptosporangiate ferns are
the dominant organs of the sporophyte.
A fern leaf is called a frond. A frond
leaf is circled in the bud to form
circinate vernation or a crozier.
Fronds are large compound leaves.
Complex, finely divided fern leaves are
considered to be the primitive condition.
Croziers in the
Polypodiaceae
Fronds
The primary leaflets are called pinnae.
The ultimate divisions are called
pinnules. The continuation of the
petioles among the leaflets is the
rachis. The petiole of each leaf is
connected to the stem by one or more
traces. Venation in the leaflets can be
dichotomous, pinnate or reticulate.
Examples of fern fronds:
Adiantum pedatum, maiden-hair fern
Asplenium viride
Athyrium distentifolium filix-femina
Dryopteris felix
Onoclea
sensibilis
 Onoclea
cinnamomea
Platycerium,
staghorn
ferns
Polypodium
pellucidum
Polystichum
Christmas tree
fern
Pteridium
aquilinum
Stem
The stem of the ferns in the Filicales can be a
horizontal rhizome or a short, slow growing
erect structure. In most cases the stem is
underground. The stem is covered with many
adventitious roots which emerge between the
leaf bases. Growth of the stem occurs by the
activity of a single apical cell and its
derivatives. Most ferns have only tracheids in
their xylem. However, some species
(Pteridium) show the beginnings of vessels.
The phloem contains sieve cells, but no sieve
tubes.
Vegetative
morphology
Roots
The roots in all ferns are adventitious
except for the embryonic radicle. The
adventitious roots develop between the
leaf bases. Secondary growth is absent
in both stems and roots of
leptosporangiate ferns.
Polypodium life cycle
Gametophyte,
macroscopic, green, free
living, 1N

gametes

Sporophyte, 2N

meiosis

Spores (1N) in
sporangium
Spore production
In most leptosporangiate ferns the initial
leaves are vegetative, subsequently
produced fronds will be both vegetative
and fertile. Groups of sporangia are
arranged in sori on the bottom side of
the leaf. Some ferns produce sporangia
on special fronds such as Onoclea
sensibilis and Polystichum
acrostichoides.
 Sori

Sporangium
with annulus
Fertile and sterile fronds
in Onoclea
Two fern sori, notice that these sori are associated
with the smaller veins of the vascular tissue.
The indusium is an important
feature for the phylogeny of ferns.
■ The sori are often covered by a layer of
tissue called the indusium. A true
indusium is a single layer of cells and of
epidermal origin. If sporangia mature
simultaneously, the development is said
to be simple. If the sori contain
sporangia in different stages of
development, development is said to be
mixed.
 Nephrolepis sori

indusium

Sorus
containing
many
sporangia
Polystichum frond with sori
covered with indusia
Sori with a marginal indusium in Pteridium,
bracken fern
Leaf and sorus cross-section
from Pteridium
A sorus containing many sporangia
Onoclea with a fertile frond
Dryopteris sori
 Osmunda fertile
pinnae on a fertile
frond
Asplenium nida sori
Athyrium sori
 Osmunda frond
containing both fertile
and sterile pinnae
Sori without indusia
“Naked sori”
 Adiantum (maiden hair fern) sori
covered with a marginal indusium
Sporangium structure
A vertical row of cells called the annulus
forms an incomplete ring around the
sporangium. The radial and inner
tangential walls of the annulus cells are
thickened. At one end of the annulus a
group of thin walled cells form the lip
cells or stomium. The annulus and the
lip cells function in spore dispersal.
Fern sporangium structure
annulus

spores

stomium
Fern sporangia, note the uneven thickness of the
walls of the cells in the annulus.
Sporangium dehiscence
Water evaporates from the sporangium
through the thin outer walls of the
annulus cells. This causes the annulus
to shorten and rupture the sporangium
transversely at the lip cells. Continued
water evaporation causes the ruptured
sporangium to snap back or catapult,
dispersing the spores.
Fern spores resemble the pollen
of flowering plants.
The walls of the spores of ferns, like the
spores of other land plants, are
differentiated into two layers, the exine
on the outside, and the intine on the
inside. The exine is variously
ornamented. The pattern of the
ornamentation can be used to identify
the type of fern which produced it. Note
the similarity to the structure of pollen.
Fern gametophytes
The spores of most ferns require light for
germination. The spore initial germinates to
produce a green, filamentous structure similar
to the protonema of mosses. The filament
will develop rhizoids. The terminal cell of the
filament will begin to differentiate segments in
two directions, thus initiating the production of
the heart-shaped fern gametophyte.
Early stages of fern spore
germination
Fern gametophyte
Fern
gametophytes
are green, free-
living, and
macroscopic.
Antheridia and
archegonia are
usually located
near the
“notch.”
Prothallia
■ The gametophytes of ferns are often
called prothallia. The antheridia and
archegonia develop from single
superficial cells of the gametophyte.
The antheridia are protuberant; only the
necks of the archegonia project from the
surface.
Fern antheridia
 Fern
archegonia
Antheridiogen
Young gametophytes secrete a substance
chemically related to the giberillins of higher
plants called antheridiogen. The
antheridiogen stimulates nearby
gametophytes to develop antheridia.
Gametophytes become progressively less
sensitive to antheridiogen and then start to
produce archegonia. Thus, gametophytes
are often protandrous. The archegonium
secretes a chemical stimulant which attracts
sperm.
Embryo development
The young fern
sporophyte
contains a
stem, leaf,
foot and root.
Order Marsileales
■ One family, the Marsileaceae is
contained in the order Marsileales. This
order contains three genera: Marsilea
and Pilularia and Regnellidium. These
are water ferns which are
heterosporous. Marsilea and Pilularia
are widespread in the United States.
Regnellidium is found only in Brazil.
Vegetative morphology
Growth of the stem, root and leaves is
from apical cells.
Air chambers are present in the leaves
and stem and represent an adaptation
to an aquatic habitat.
Roots arise adventitiously from nodes.
Marsilea and Regnellidium have vessels
in their xylem, Pilularia lacks vessels.
Spore production
Sporangia of Marsilea are borne in
sporocarps. Sporocarps are modified fertile
pinna. At maturity the sporocarps become
dark in color and hard. Each sporocarp
contains two elongate sori. The sori are
covered with a delicate indusium.

Development of the sporangia is

leptosporangiate. Heterospory is evident in
the sporocarp.
■ Within each receptacle both
microsporangia and megasporangia
develop. Both micro- and mega-
sporoangia lack an annulus. Spores
are not dispersed until the sporocarp
wall decomposes.
Gametophytes: similar to
Selaginella and Isoetes
Development of the male gametophyte begins
with a 2 cell stage containing a small
prothallial cell and a large antheridial initial.
The entire antheridium develops from the
antheridial cell. The male gametophyte is
composed of an antheridium surrounded by a
sterile jacket of cells. It protrudes slightly
from the microspore wall, but as in
Selaginella and Isoetes, the male
gametophyte is small and only approximately
the size of the spore itself.
The female gametophyte is also much
reduced. Cell division produces a small
amount of vegetative tissue with a
single apical archegonium.
Embryo
The embryo contains a foot, leaf, stem
and root.
The development of the gametophytes
and embryonic sporophyte is very rapid.
Order Salviniales
Two genera are present in the Salviniales,
Salvinia and Azolla.
Vegetative plant
Salvinia is rootless. The leaves are in
whorls of three, two of which are floating
and the third submerged. The floating
leaves are covered with hairs and waxy
papillae.
Azolla
■ Azolla is also an aquatic, floating plant.
It is highly branched with dense
alternate bilobed leaves. Each leaf is
composed of a chlorophyllous dorsal
lobe and a submerged non-green
ventral lobe. Azolla grown in strong
sunlight often becomes orange-red.
The colored pigment is an anthocyanin.
Azolla has true roots.
Spore production
Both Salvinia and Azolla are heterosporous
and produce their spores in sporocarps. The
sporocarps represent single sori enclosed by
a modified indusium. The sporocarps of
Salvinia are located at the tips of the short
branches of the submerged, branching
rootlike leaves. The sporocarps of Azolla are
formed in groups of two or four in the axil of
the dorsal lobe of a basal leaf of a baranch.
The vegetative plants are monoecious; a
sporocarp contains only micro- or mega-
sporangia.
Gametophytes
The gametophytes have little vegetative
tissue and are small, as in the other
heterosporous vascular cryptogams.
Fossil ferns
Fossil Ophioglossopsida are unknown. The
Marattiopsida are known from the
Pennsylvanian. One family (Osmundaceae)
of the Filicopsida is represented beginning
with the Permian, but most are Mesozoic or
Cenozoic. The Polypodiaceae do not appear
until the Jurassic and thus are as recently
evolved as the Angiosperms.
Primitive fern characteristics
1) elaborate, compound leaves
22) erect, epiterranean stems
33) massive sporangia with no annulus,
producing indefinitely large numbers of
spores
44) sporangia marginal on leaves and
at the ends of leaves
Advanced fern characteristics

11) simple leaves

22) prostrate, rhizomes for stems
33) small sporangia with annuli, fewer spores
44) sporangia borne on the abaxial surface of
leaves instead of marginally