Introduction

The Pteridophytes refer to the ferns and their allies.

Pteridophytes are non-seed bearing plants having a vascular system
consisting of xylem and phloem.
The vascular system connects true leaves, roots, and stems.
Different approaches
A single division Pteridophyta,
At class level: Psilotopsida, Lycopsida, Sphenopsida and Filicopsida (or
Pteropsida)
The current approach divided into four major divisions:
Psilotophyta, Lycophyta, Sphenophyta, and Pterophyta
Biologists consider these divisions as lower vascular plants,
primitive vascular plants, seedless vascular plants, or
Pteridophytes
Conversely, gymnosperms (e.g., a pine tree) and angiosperms
(flowering plants) are considered higher vascular plants, seed-
forming vascular plants or spermatophytes.
Vascular plants are also collectively called tracheophytes.
This separation of "higher" and "lower" vascular plants was invented
by early botanists, who believed that all seed-forming plants must be
related, and that plants lacking seeds are similarly related.
According to later investigations ferns represent a separate group of
seedless plants
Reason : the presence of Megaphylls ( large leaves with several to
many veins).
The remaining seedless vascular plants = microphylls, which are
small leaves with one vein and are only somewhat related to ferns,
hence the name "fern allies."
All ferns and fern allies possess sporophylls (sporangia bearing
leaves).
Sporophylls are leaf-like structures of the sporophyte generation
that bear sporangia.
They may be large and have several to many veins (Megaphylls of
ferns) or they may be smaller and have one vein (microphylls of
fern allies).
Thus the "lower vascular plants" include a diverse group of ferns
and fern allies
Evolutionary history
Silurian plants
Land plants needed to develop ways to obtain water and nutrients
Epidermis-reduce water loss,
Stomata to breathe
Devonian
Different groups has developed important characteristics
Five classes of fern allies had arisen by this time the Psilotopsida,
Trimerophytopsida, Zosterophyllopsida, Lycopsida, and
Sphenopsida.
Diagram showing the evolutionary history and relationships of
vascular plants. Adapted from
http://www.hcs.ohio-state.edu/hcs300/svp1.htm
From these early plants the ferns and fern allies as we know them tod
arose.
Below shows how the evolutionary lines developed, with some groups
lacking any modern descendants omitted. The vascular plants are land
dwelling plants one division, the Tracheophyta suggesting a close
evolutionary relationship among the major groups of vascular plants.
The evolutionary history of the vascular plants remains controversial
due to the assumption that the major groups of vascular plants have
evolved along independent lines.
The classification system used in Campbell reflects the latter view and
divides the living vascular plants into nine divisions (divisions:
Psilotophyta, Lycophyta, Spenophyta, Pterophyta, Cycadophyta,
Ginkgophyta, Conferophyta, Gentophyta and Anthophyta).
Vascular plants or tracheophytes, first appear in the fossil record abou
410 million years before present.
All lines of evidence suggest that the tracheophytes developed
directly from some complex, multicellular Chlorophyta.
Unfortunately, an extremely large gap exists between the algae and
vascular plants, and there are no living plants which bridge the gap.
Unfortunately, an extremely large gap exists between the algae and
vascular plants, and there are no living plants which bridge the gap.
Bryophytes are somewhat intermediate but are closer to algae than
even to the simplest vascular plants, and it appears that the
intermediate plants have become extinct.
In addition, vascular plants are identified by independent and
dominant sporophytes and greatly reduced gametophytes. This is in
direct contrast to the bryophytes. The sporophytes of most vascular
plants are differentiated into roots, stems, and leaves. Also, vascular
plants are typically land plants and, like bryophytes, have evolved
terrestrial adaptations.
The transition from water to land was thus gradual over
millions of years.
Problem for early land plants was availability of water.
A successful land plant must possess an efficient
mechanism for absorbing materials from the soil and
transporting them throughout the plant. Once obtained from
the environment the water must be conserved. This problem
led to the evolution of vascular tissues by vascular plants.
Thus, the development of vascular tissue allowed plants to
successfully colonize habitats that until then were not available to
them. Xylem is the tissue responsible for water conduction in
vascular plants, but also provides support due to its lignified cell
walls. Phloem is the tissue responsible in conduction of
photosynthetic products.

.
In conclusion, Pteridophytes are therefore the earliest
vascular plants.
All members exhibit an alternation of generations life cycle,
as do the bryophytes, but here, the sporophyte is the
dominant generation and the gametophyte is reduced and
either free-living or contained within the spore.
Pteridophytes exhibit a range of habits and life forms,
and exist in most habitat types except the marine
environment. The sporophytes range in size from a few
millimeters to tens of meters tall and in mass from a
few milligrams to many tens of kilograms
Pteridophytes habit reflects the environment and substrate in
which they choose to grow and it is convenient to divide to
recognize three major classes: those that grow on the ground,
those that grow on trees and those that grow in or on the
water. However, it should be noted that a habit/habitat
preference may span these classes.
In particular, species that live low on trees might be classed as
epiphytes, sub epiphytes or terrestrial, and plants that grow
on the margins of water bodies might be considered aquatic.
Branching patterns in vascular plants
Monopodial branching: of branching patterns, in which there
is one main shoot with lateral branches emerging from it.(e.g.
Tid, Zigba, etc.) Consider the Christmas tree image!
Dichotomous branching: a type of branching in which the tip
divides into two more-or-less equal apices; stem like a fork.
Quite common in the lower vascular plants such as Psilotum,
Selaginella, and Lycopodium.
Sympodial: when growth continues through a series of
axillary branches that seem to form a single stem. It is with a
single short trunk and irregular branching above the trunk .
An example of sympodial branching is a pine tree.
Major families of Pteridophytes are predominantly terrestrial,
predominantly epiphytic or predominantly aquatic,
demonstrating a significant correlation of habit defined in this
manner with other morphological characters.
Note that the structure of the stem or rhizome has a
particular impact on the habit type, compact or creeping
stems determining the clustering or spacing of the fronds.
Terrestrial Pteridophytes have erect or creeping stems and
the leaves are held more or less upright, either vertical or
spreading and arching. Erect stems are generally unbranched,
radial with a more or less terminal rosette of fronds, and may
be stout and woody.
Epiphytic Pteridophytes have their stems attached to or rooted on
trunks or branches of trees. Epiphytes can have compact or short to
long-creeping stems. The stems can start on the tree or on the
ground, but in any case are attached to the tree and are
Sub epiphytes live low on the base of tree trunks, generally among
bryophytes and decaying detritus. They may also occur on mossy
ground and mossy rocks.
Aquatic Pteridophytes can have compact or shortly creeping stems
and can be free-floating on the surface of the water.
The stem is the main lengthwise growing axis of the fern or fern ally
and bears the roots or root-like organs for attachment to the
substrate and acquisition of nutrients and water, and produces
leaves or fronds in a more or less regular manner. Once leaves or
fronds are produced, they generally do not increase in diameter at
that point for the most part, they are not photosynthetic but in
some epiphytic climbing species there is some chlorophyll
Pteridophytes roots are not produced as the downward proliferation
of the stem but are mostly produced laterally, sometimes scattered
along the length of the stem, and are thus adventitious.

The leaves of typical and true ferns are usually called fronds and
those of fern allies referred to as leaves.
The bewildering varieties of leaf form or arrangement provide many
of the most useful characters of Pteridophyte taxonomy. Fern fronds
are Megaphylls and except in reduced species, are large and complex,
have many veins and are often lobed or variously divided, commonly
with a distinct stalk or petiole. The microphyllous leaves of
Psilotaceae and Equisetaceae are reduced to minute brown teeth and
the photosynthetic function has been taken over by the green
phyllodinous stems. Thus, from an evolutionary perspective:
Microphylls are single veined leaves associated with leaf gaps and
evolved as superficial outgrowth of the stem.
Megaphylls have a complex venation pattern, and evolved from a
branch system.
Life Cycle
oThe life cycle of pteridophytes involves two distinct and separate
phases, the gametophyte (also called prothallus) and sporophyte . The
sporophyte, is the most conspicuous phase, and is so-named because
this is the stage that produces spores; it is the sporophyte that most of
us refer to when we describe or name pteridophytes. Under favorable
conditions, the spores shed from the sporophyte and develop into the
gametophyte, which is small, inconspicuous and short-lived.
oThe purpose of the gametophyte is to produce the male and female
sex cells (gametes), the female of which, when fertilized, develops into
a new sporophyte to continue the cycle. This is a regular process and
the alternation of generations, which involves an alternate doubling
(2n) and halving (n) of chromosome numbers at each phase .
In the bryophytes, as you are well aware of, the gametophyte is
the dominant and most conspicuous stage and the sporophyte is
retained as a relatively small dependent appendage on the mature
gametophyte.
pteridophytes exhibit two types of life cycles: homosporic life
cycle (like all members of bryophytes) producing all equal sized
spores or heterosporic life cycle producing two different sized
spores, called megaspores and microspores
Gametophyte
The gametophyte is the sexual or haploid stage of the
pteridophyte life-cycle and contains a single set of chromosomes
(n). It develops from the spore (n) produced on the sporophyte.
This spore germinates and develops by mitosis into a body called
the prothallus (n).
At maturity, regions of the prothallus develop into small separate
sexual organs (n). Antheridia produce the male sex cells (sperm)
and archegonia produce the female sex cells (egg).
The female sex cells are usually sedentary whereas the male are
usually more or less coiled and motile by means of two or more
cilia and swim or are carried by water to the archegonia where
fertilization takes place.
Water is still essential for the process of fertilization. After
fertilization a new sporophyte develops from the fertilized cell and
takes over as the gametophyte withers and dies.
This is the general pattern of behavior but there are fundamental
differences in a number of genera with dimorphic spores and a
number of genera that abandon the haploid phase (apogamy).
In most cases, the prothallus is green and photosynthetic,
developing on the surface of the ground or on moist rocks or bark.
They lack vascular tissue; mostly they are thin and more or less
heart-shaped, less often ribbon-like or filamentous.
The central region is somewhat thickened and bears the
antheridia, archegonia and rhizoids (filamentous root-like
structures).
In the genera of the Ophioglossaceae, Lycopodiaceae, Psilotaceae
and some Schizaeaceae the prothallus is thick, non-photosynthetic,
grows and live in the dark, underground or in the detritus in the
forks of trees, and is saprophytic, obtaining its nutrients from
decaying vegetable matter with the assistance of an endophytic
fungus.
Sporophyte
The sporophyte is the asexual or diploid stage of the pteridophyte
life-cycle. It is mostly large and conspicuous, always green and
photosynthetic, long-lived and produces the spores in special
fruiting bodies; the spores are shed to produce the next
gametophyte generation in the cycle.
 the sporophyte generation is the foundation of all pteridophyte
classification and taxonomy thus the sporophyte alone is the
fundamental unit of a modern pteridophyte specimen.
It is convenient to consider the plant to be composed of three major
parts: the stem which bears the roots, the leaves and the fertile
parts with sporangia.
 Leaves that bear sporangia, which are referred to as fertile as
opposed to sterile for those without sporangia. In the fern allies
sporangia bearing leaves are the sporophylls.
Fertile leaves or fronds may or may not be different to the sterile.
The sporangia of the fern allies are generally borne on the adaxial
(acroscopic) or upper side of the sporophylls, at the base near the
axil of the leaf. In true ferns they are generally produced in various
ways on the abaxial or lower side of the leaf, remote from the base.
Sporangia
The sporangia are the bodies in which the spores are produced.
They are generally small and superficial but may be large and
immersed in the sporophyll (e.g. Isoetes). The mostly minute
spores are shed when the sporangium dehisces or splits although
sometimes the spores are released by decay of the sporangial wall
(e.g. Isoetes). In most cases the sporangia burst under dry
conditions and dispersal is by wind.
The ferns are divided into two main groups on the basis of their
basic sporangia type :
Those with thick walled (several cell layers thick) sporangia that
develop from several cells are known as eusporangiate and are
considered to be more primitive. The bulk of the ferns have thin-
walled sporangia developed from a single cell and are termed
leptosporangiate; these are the advanced ferns.
In the Psilotaceae, perhaps the most primitive group of
pteridophytes, the sporangia are large and united into twos or
threes (synangia) each splitting when matures

Synangia

Psilotum nudum sporophyte bearing synangia. Image adopted from

http://www.botany.hawaii.edu//.
The sporangia of the remainder of the true ferns are all remarkably
similar. They are thin-walled and dehiscence is caused by a region of
cells with thick inner walls and thin outer walls, usually in a ring,
called the annulus.
The microscopic structure of the sporangia yields many useful
taxonomic characters.
Spores
There are two basic types of spores in the pteridophytes: monolete
(or bilateral) and trilete (or tetrahedral). Spores are always produced
in tetrads (groups of four) and the spore shape results from the two
possible cleavage patterns of the initial cell. If the second cleavages
are in the same plane the result is a bilateral spore with a monolete
spore; each spore as it develops has two neighbors, the two contact
faces meet in a single ridge of scar and the free spore is more or less
bean-shaped.
If the second cleavages are in planes perpendicular to each other,
the result is a tetrahedral spore with a trilete scar; each spore as it
develops has three neighbors, the three contract faces meet at a
three-armed ridge or scar, the four spores are positioned at the four
corners of a tetrahedron and the resulting spore is more or less
globular or tetrahedral with a rounded base.

Spore production and conditions (homospory and heterospory)

Homospory
Plants that produce bisexual gametophytes have their
gametophytes germinate from homspores or isospores (iso or
homo=same) that are about all the same size. This is a homosporic
condition.
Homospory is, thus, a condition in which plants produce only one
type and size of spores, (microspores). All bryophytes and
psilophytes and the great majority of lycophytes, sphenophytes and
pterophytes are homosporous.
A typical homosporous life cycle. Note the production of a single type of
bisexual gametophyte that will eventually produce the antheridia (sperm
bearing structures) and archegonia (egg bearing structures). Image taken
from (http://www.sinauer.com/)
Heterospory
Plants that produce separate male and female gametophytes have
those gametophytes germinate from (or within in the case of the more
advanced plants) spores of different sizes (heterospores;
hetero=different)= heterosporic condition

Typical heterosporous life cycle. Note the production of separate,unisexual male

and female gametophytes. Image adapted from (http://www.sinauer.com/
Heterospory is, thus, a condition in which plants produce two
different types and sizes of spores = microspores and megaspores.
As a reproductive strategy, heterospory involves the following steps:
=(1) The mature diploid plant (sporophyte) produces "male" and
"female" spores. Typically, the plant produces numerous small "male"
spores (microspores) and fewer, but larger "female" spores
(megaspores).
=(2) The megaspores are usually retained on the sporophyte, where a
haploid female gametophyte develops inside the spore.
=(3) The microspores may or may not be retained on the sporophyte.
They develop into sperm-producing, haploid gametophytes.
=(4) The sperm are released into the environment, and dispersed
between plants.
=(5) The sperm fertilize female gametophytes grown from
megaspores, producing a diploid zygote (fertilized cell).
=(6) The zygote develops into an embryo and finally a new diploid
sporophyte.
Heterospory evolved a number of times among different ancient
plants. Lycopsids, ferns, sphenopsids and progymnosperms all
developed heterospory. Many of these plants also evolved into large
trees. The obvious advantage for heterosporous trees is that their great
height would enhance the dispersal of windborne microspores.

Heterosporous pteridophytes
Certain groups of pteridophytes (e.g. Selaginellaceae, Isoetaceae,
Marsiliaceae, Azollaceae, Salviniaceae) produce haploid spores of
different sizes (big=megaspores; small=microspores). As described
above, the microspores develop into small microgametophytes and the
megaspores develop into larger megagametophytes. Apart from the
separation of the gametophytes at the sexual stage, the alternation of
generations is essentially the same.

Agamospermous pteridophytes
In some species of ferns and their allies, the sexual process is omitted
and the spores produced on the sporophyte are diploid rather than the
usual haploid. The resulting prothallus is thus diploid and a few
sporophytes are produced vegetatively without the usual union
of gametes.
In conclusion, the life cycle of a vascular plant is conceptually
the same as for bryophytes. The major difference is that the
sporophyte generation is the dominant and persistent phase of
the life cycle, while the gametophyte is reduced and short-lived.
The gametophytes are structurally simple, ranging from the free
living macroscopic thalli of the ferns to the microscopic
gametophytes of the seed plants.
The sporophyte phase consists of a complex plant body
composed of roots, stems and leaves. These plant organs possess
highly differentiated vascular tissues called xylem and phloem.

World distribution and habitat use

Pteridophytes are distributed throughout the world, depending
on the particular division.
Psilotophyta are considered tropical plants, Psilotum, for
example, grows wild in Florida wood lands.
The Lycopodium and Selaginella species are examples of Lycophyta
and can be found in various places throughout the Earth.
Lycopodium usually grows beneath other plants, while the more
delicate and more tropical Selaginella usually grow in damp
locations.
Equisetum is widely distributed, finds its home in wet-lands,
marshes, and anywhere where the soil is saturated with water.
Ferns encompass all continents except Antarctica and most islands
(Knee, 2001). Although they favor moist temperate and tropical
regions, ferns have the ability to withstand most climates except for
extreme cold and extreme arid environments. There are no marine
species, but water is required for fertilization and production on new
generations of plants. Environments with low light and high levels of
moisture and humidity are perfect homes for ferns. Pteridophytes
flourish in tropical forests (Knee, 2001).
Ecological roles
The pteridophytes formed approximately 290-360 m.y.a., during
the Carboniferous period. These plants formed in forests, and
they left fossilized fuel in the form of coal. Although coal has
formed over many geological periods, the strata of the
Carboniferous period have by far the largest coal deposits.
Pteridophytes play a role in providing food for wildlife because
they are primary producers .
Impacts on human society
Provide beauty in their natural habitat.
Ferns also have a horticultural value in society.
People are knowledgeable and can recognize ferns because they are
unique and catch our undivided attention.
Ferns are intentionally added to indoor and outdoor gardens for
decoration and enjoyment. Believe it or not, ferns do have an impact on
historic life. The Industrial Revolution was "fueled" by coal, and coal is
becoming more and more important as society uses up nonrenewable
fuels such as gas and oil.
The sphenophytes, or horsetails, got their name "scouring rushes"
because people use them to scrub pots and pans. The rough texture of
their stems makes them ideal for this purpose. See later for the rest
under each particular lesson.
Comparison with other vascular plants
The ferns and their allies share a lot of common morphology with the
other vascular plants thus in many cases the same descriptive
terminology is used.
The most obvious difference between the pteridophytes and the
remainder of the vascular plants (seed plants) is that they do not
produce large floral or reproductive structures that give rise to seeds,
which eventually develop, into the next generation of plants.
Pteridophytes reproduce by microscopic spores not seeds
Release of spore into the gametophyte=primitive mode
In contrast to algae, fungi and bryophytes, the sporophytes of
vascular plants possess a well-developed vascular system that serves to
conduct water, mineral salts, and food.
Taxonomy
Divisions: Psilotophyta (whisk ferns), Lycophyta (club mosses),
Sphenophyta (horsetails or scouring rushes), and Pterophyta (true
ferns).
Divisions of Extinct Seedless Vascular Plants
There are three divisions of extinct seedless vascular plants which
include Rhyniophyta, Zosterophyllophyta and
Trimerophytophyta.The genera Rhynia, Zosterophyllum and
Trimerophyton are members of these divisions. They are the
earliest known, which go back about 425 m.y.a. and most went
extinct by the end of the Devonian about 370 m.y.a.. These three
groups were the dominant vegetation from the mid-Silurian to the
mid-Devonian, 425 to 370 m.y.a.
For the most part they were relatively simple plants 18 in to 36
inches tall. They had the following characteristics:
Naked photosynthetic stems ,Terminal sporangia (some lateral)
No roots or leaves ,They were all homosporous
They had protosteles
Pterophytes, lycophytes and progymnosperms are more
complex groups that were dominant from the Late Devonian
through the Carboniferous, from about 370 to 290 m.y.a. Seed
plants arose starting in the Late Devonian period, about 380
m.y.a., and evolved many new lines by the Permian (290 to 248
m.y.a.). Gymnosperms dominated the land floras throughout the
Mesozoic until about 100 m.y.a. Angiosperms appeared in the
fossil record about 125 m.y.a. It became the dominant group
about 30 to 40 m.y.a. and has remained so until the present.
Division Rhyniophyta
oSeedless; produced spores.
oDichotomous branching.
oTerminal sporangia.
oHomosporous.
oPlant body was not differentiated into roots, stems and leaves.
oUnderground rhizome with rhizoids.
oProtostele consisting of a core of xylem surrounded by
one or two layers of phloem cells.
oSome had conducting cells similar to hydroids rather
tracheids; they are called protracheophytes.
oThere is evidence of isomorphic alternation of
generations.
Examples: Rhynia, Cooksonia, Aglaophyton.

Cooksonia fossil specimen (Left) and reconstruction (Right).

Image taken from the UCMP Berkeley website.
Division Zosterophyllophyta
oGametophyte was likely photosynthetic and structurally complex.
Sporophyte was herbaceous; with dichotomously branching stems
covering with lateral spines.
oZosterophyllum is dichotomously branched, but frequently lateral
branches further differentiated into one axis that grew upward and
another downward.
oAerial stems were covered with cuticle and had stomata only on
the upper branches; leaves lacking.
oRoots lacking but probably had a branching rhizomatous system
with rhizoids. The lower branch may have function as a root.
Zosterophylls were homosporous.
Thick-walled eusporangia were globose or reniform occurring on
short lateral stalks on the sides of the fertile stems; some
Zosterophylls were naked or had small spine-like enations.
A slender solid exarch vascular bundle; xylem matured from the
periphery toward the center: centripetal differentiation.
oAll plants extinct from the Devonian period. Fossils indicate
that the plants grew in shallow aquatic habitats.
oThey are considered to be the ancestral group that gave rise
to the Lycopods; the sporangia of both groups are borne
laterally and are similar in shape, and the xylem in both
groups differentiates centripetally.

Zosterophyllum sp. Image from http://www.botany.hawaii.edu//.

Division Trimerophytophyta
oThey lacked leaves.
oLateral branches forked dichotomously several times.
oThey were homosporous.
oSome branches were vegetative while others bore elongated
sporangia.
oVascular strand was more massive than that of the rhyniophytes
oThe xylem differentiated centrifugally

Trymerophyton sp.
Image from http://www.botany.
hawaii.edu//.
DIVISION PSILOTOPHYTA: The Wisk Ferns
Introduction
single family, the Psilotaceae, the Whisk fern family, with two
genera, Psilotum, the whisk fern and Tmesipteris.
They are 2-3 feet tall, and are native to southeastern United
States, Japan and Australia.
They have simple plant body (with very primitive
morphology) and with virtually no true leaves and roots.
They are herbaceous and dichotomously branched with
underground stems (rhizome- rhizomatous), from which the
above ground parts, stems, branch off.
The branches are terminated by small sporangia fused
together in threes.
Members are homosporous with sporangia being bilobed or
trilobed developing laterally on stem, subtended by a pair of
scales, called enations.
The trilobed sporangium is called synangium (plural-
synangia)
Spores released from the sporangium germinate in the soil to
gametophytes, which are found beneath the surface of the soil
(subterranean) and are easily overlooked since they are
colorless. The gametophytes are saprobic and obtain their
nourishment through association with mycorrhiza fungi.
Interestingly, this is mycorrhizal associations; perhaps they are
necessary in the absence of true roots. After fertilization the
zygote develops a foot and a rhizome. As soon as the rhizome
establishes itself with the aid of the mycorrhizal fungi, upright
stems are produced and the rhizome separates from the foot.
Evolution
Common during the Devonian period
The evolutionary significance of the psilophytes is that they
are the most primitive known group of vascular plants and
some plant biologists consider them to be a transitional group
linking the aquatic algae with more advanced pteridophytes
(club mosses, horsetails, and ferns).
Psilotum nudum morphology. A. Sporophyte plant. B. A bit of stem
with scaly appendages. C. A gemma on rhizome. D. A fertile twig.
Image adopted from http://www.botany.hawaii.edu//.
However, there are others which disagree with this idea saying
that there are no organisms which bridges this gap.
they suggest how club mosses, horsetails, and ferns may have
originated by various leaf modifications and developments.
Certain psilophytes with small leaves developed as emergences
and may have given rise to the Lycophyta line.
Others with whorled branches may have been the forerunners of
the Sphenophyta line.
The fern line may have developed from forms in which the
branch tips were flattened, possibly leading to the evolution of
large leaves.
Psilophytes best represent what some of the first vascular land
plants in the Devonian would have looked like. It has only
dichotomously branching stems, terminal sporangia, and leaf-like
enations. It is thus resembles the Devonian vascular plants such as
Rhynia and Cooksonia.
 Division characteristics
Sporophyte with a dichotomously branching aerial and
subterranean stem system.
True roots and leaves lacking.
Underground stems with rhizoids and with a fungal association,
an endomycorrhizal zygomycete.
Aerial stems lacking leaves but with scale-like or larger leaf-like
structures (enations).
Gametophytes bisexual, subterranean, resembling a small piece
of rhizome.
Gametophyte with a symbiotic fungus. Some have a vascular
tissue.
The sperm is multiflagellated and requires water to swim to the
archegonium.
Homosporous.
.
Sporangia, a 2 or 3-chambered synangium, borne at the apex
of small side-branches (appear to be arranged along the sides of
the major stems of the plant, the sporophyte).
Sporophyte remains attached to the gametophyte in the early
stages by a foot and derives nourishment from the gametophyte
Classification
two extant genera,
Psilotum and Tmesipteris, from tropical and
subtropical regions of the world. Consider
Psilotum f
Its taxonomic status is:
Class: Psilotopsida
Order: Psilotales
Family: Psilotaceae
Genus: Psilotum
oFamily Psilotaceae
o Small to moderate-size terrestrial or epiphytic herbs ,rhizome shortly
creeping, dichotomously branched, protostelic, lacking roots, densely
covered with fine rhizoids.
oStem in Psilotum repeatedly dichotomous upward.
oHomosporous
Distribution
oIt is a bigeneric tropical, subtropical and southern temperate family
with approximately 12 species.
Xylem occupies the center but the phloem occupies the indentations in
the xylem. This generates the star-like pattern, forming actinostele.
Note that cells in the center of this heterotrophic thallus are invaded by
an endophytic fungus. The gametophyte consumes digestible
constituents of the fungal hyphae as a food source, leaving the
indigestible parts in a knot-like lump, the "hyphal plexus", in the center
of the cell. Surplus carbohydrates are stored as starch in amyloplasts
Dichotomously branched Psilotum gametophyte bearing antheridia,
archegonia and rhizoids (Photo by Karen Renzaglia). Image taken from (
http://www.sinauer.com/)
Life cycle
In the reproductive cycle of whisk ferns, a dominant
sporophyte plant (1), bearing trilobed sporangia (2), produces
homosporous spores (3) which germinate into haploid
gametophytes (4). The underground gametophyte forms
mycorrhizal associations with fungi for nutrition and develops
archegonia (5), each with 1 egg, and antheridia (6) with
flagellated sperm.
After fertilization a diploid zygote (7) forms and eventually
develops into a sporophyte plant (8).
Asexual reproduction can take place by means of gemmae.
While gemmae are means of asexual reproduction in non-
vascular plants such as the liverworts, the difference is that in
the non-vascular plants the gemmae develop on the
gametophyte while in Psilotum the gemmae develop on the
sporophyte.
The life cycle of Psilotum. Image from http://www.botany.hawaii.edu
//.
DIVISION LYCOPHYTA: The Club Mosses
Introduction
oAlthough they are a relatively inconspicuous component of the
modern flora, the lycophytes included large trees of the late
Paleozoic. These trees, some of which grew to be more than 35
meters tall, formed forests in the Carboniferous coal swamps of
North America and Europe. The large number and good
preservation of these ancient plants has allowed us to learn much
about their biology, which will eventually be presented in a
separate exhibit.
oThis Division contains three main modern genera:
Lycopodium (lyco = wolf) (poda = foot) also known as ground pine
or wolf’s claw ,which is the largest of the rest
Selaginella
Isoetes
Running pine of Lycopodium complanatum. Image from
http://www.botany.hawaii.edu//.
oMany tropical species in this Division are epiphytes.
oEpiphytes are plants that use another plant as a substrate upon
which to live but are not parasitic.
oMost species have a rhizome from which the roots and branches
arise.
oThese plants possess true vascularized stems, leaves and roots.
oThe leaves are described as microphylls, i.e. enations or reductions
of the stem in which the vascular strand is simple, a trace, or a single
vein (a central vein that does not branch).

Proposed steps in the evolution of the microphyll leaf. Note that microphylls do not
leave a leaf gap in the stem's vascular cylinder. If we wanted to place Psilotum-like
plants on the left, we would have Lycopodium-like plants on the right. Image from (
http://www.sinauer.com/).
oThe spore producing bodies are borne on special branches and the
structure is a strobilus (plural: strobili).
oThus, lycophytes have their sporangia organized into strobili, which
are the spore producing bodies of the sporophyte generation.
oThe sporangia which contain the spores are associated with fertile
leaves known as sporophylls.
oThe leaves of the living members of this division are generally small
and usually arranged in a spiral.
oLeaf gaps are never formed at the junction of stem and leaf in the
Lycophyta, so that a stem of a member of this division has a core of
xylem (protostele)
oThe mature sporophyte has two types of stem: upright and horizonta
stems. The upright stems grow from the horizontal stem . Both types o
stems are sheathed with small green leaves. Small but well-developed
adventitious roots arise from the underside of the horizontal stem . The
stem lacks pith as do roots of most of higher vascular plants. The xylem
is composed of tracheids whereas phloem contains sieve cells.
oSome of these lycopodalean gametophytes may persist for 15
years before releasing their sperm or producing a sporophyte.
oOther lycophytes, such as Selaginella, have gametophytes
which develop entirely within the wall of the spore (endosporic),
never venturing out into the environment. These gametophytes
may begin their sexual phase while still within the parent's
sporangium, producing the new sporophyte generation from
within the tissues of the previous sporophyte generation.
oTheir sporangia are borne on sporophylls. Sporophylls are either
grouped in terminal aggregations to form a strobilus or are
distributed in alternating clusters with non sporangium bearing
leaves.
oBoth Selaginella and Isoetes-heterosporous
Evolution
The lycophytes, also commonly known as Lycopods dominated
the world flora during the Devonian. During the Carboniferous
Period of geological time, this group was represented by some of
the largest and most numerous plants.
During the Carboniferous Period of geological time this group was
represented by some of the largest and most numerous plants. As
described above, members of this group possess narrow pointed
leaves that are vascularized by a single strand of vascular tissue
(microphylls), which cover the stem. When they fall away, they leave
the stem covered by characteristic diamond shaped scars. Modern
representatives, the "club mosses," are a sad remnant of a once
mighty radiation. During the Mississippian and Pennsylvanian, their
tree-sized relatives, including Lepididendron and Sigillaria were the
largest forest plants. The sporangia of lycophytes are born on the
upper surface of specialized leaves which grow in groups, forming
cones or strobili at the ends of stems. Earliest representatives of
lycophyte lineage may be Zosterophyllopsida, (Zosterophyls)
resembling rhyniophytes but with alternating sporangia on stem.
oOn the other hand, the lycophytes are also believed to have
evolved from the psilophytes. Although members of this group
are of no real economic importance, the group does illustrate
some noteworthy evolutionary advances over the psilophytes.
oThe presence of true roots, and leaves, increased development
of the vascular tissue, and the organization of sporophylls into
cones (strobili) are all viewed as advances over the psilophytes.
o It is not clear why the lycophytes have not given rise to more
advanced groups of plants. Some have developed secondary
growth (e.g. Isoetes) and exhibit early stages of the seed habit.
Ecology
oModern lycophytes are low-growing understory plants,
epiphytes, and submerged aquatics -they are not a significant
fraction of the modern vegetation.
oTheir greatest diversity (as with virtually everything else) occurs
in the wet tropics, though there are several alpine and subarctic
species.
oA number of species of Selaginella are known for their survival in
extremely xeric conditions. These species occur in the dry, rocky
regions of the tropics and subtropics where they cling to the sides of
slopes and along the edges of outcrops. One species, Selaginella
lepidophylla, is so tolerant of drying out that it has been dubbed the
"resurrection plant", for it can recover from several months of
complete drying. This is uncommon among vascular plants. Many
tropical lycophytes experience xeric conditions as well. Those plants
which live as epiphytes in the rainforest canopy are subject to intense
light and drying winds.
oLike ferns and other seedless vascular plants, lycophytes produce
gametophyte plants which develop independently of their parent
sporophyte. In the club mosses , the gametophyte is usually quite
small, and may even develop underground like psilotophytes. It relies
on a mycorrhizal fungus to help provide nutrients and water.
oboth homosporous and heterosporous members
oThe homosporous members have exosporic bisexual gametophytes
(e.g. Lycopodium).
oThese gametophytes are free living and either photosynthetic or
subterranean and dependent on a fungus for nutrition.

Life cycle
oThe gametophyte that arises from the microspore is a micro-
gametophyte.
oendosporic
onot photosynthetic but heterotrophic
oforms antheridium (sterile jacket) containing up to 32 sperms
oThus, the microgametophyte can be thought of as a male structure.
oThe microspore wall and sterile jacket rupture in free water to release
the sperm to swim to the egg.
omicrogametophyte lacks rhizoids or sterile thallial cells
oMegaspore give rise to a megagametophyte
oIt too is endosporic and heterotrophic
oIts large volume means that it can contain considerable reserves, usuall
oils.
oThe female gametophyte is less reduced than the male.
Division characteristics
oSmall, usually underground, saprophytic gametophyte; which grows
within the bounds of the spore wall in some.
oSporophyte is herbaceous; extant species generally creeping and
with upright dichotomizing axes.
oHomosporous to heterosporous.
oThick-walled eusporangia usually associated with microphylls
(sporophylls) often united into a strobilus.
oVariable stele; actinostelic to siphonostelic.
oMicrophyllous leaves, which are spirally arranged on the stems.
oRoots present; protostelic.
ofrom terrestrial and epiphytic to aquatic and semi-aquatic;
found from tropics to subarctic regions .
Classification
othe extant lycopods or club mosses have 1000 species.
oThe Lycophyta has two classes: Aglossopsida (leaves without
ligules) and Glossopsida (leaves associated with ligules).
oThese characters also correlate with the occurrence of a ligule
with the sporophyll (the ligulate species are heterosporous).
Order Lycopodiales
oThese plants are extant but have a fossil history which goes back to
the upper Devonian.
oThe microphylls are closely spaced on the stem and sporophylls are
clustered in strobili. Examples: Lycopodium, Huperzia
Order Selaginellales
These plants are extant and have a fossil history from the
Carboniferous. Plants with scale-like leaves; some are anisophyllous.
Example: Selaginella,
Order Isoetales
oThese plants are extant with a fossil history which dates back
to the Cretaceous (although it is probably much older).
oThe stems are reduced but exhibit secondary growth. The
leaves are quill-like. Example: Isoetes
Extant taxa of Lycophytes
Three most important extant orders of division Lycophyta are
order Lycopopsida, Selaginelopsida and Isoetopsida.
The three prominent families and genera of these order, which
this course emphasize on include:

oLycopodiaceae, Lycopodium, or club mosses;

oSelaginellaceae, Selaginella, or Spike mosses; and
oIsoetaceae, Isoetes, or Quillworts
oThus, in this course, the Lycophyta are well represented by the
three genera: Lycopodium, Selaginella and Isoetes, representing
Lycopodaceae, Selaginellaceae and Isoetaceae, respectively
Class Lycopopsida
Terrestrial or epiphytic, small to large size, stems erect,
pendulous, creeping or scandent, rooted at the base or stolon-like
and rooted at intervals,
 simple or few- to many-times dichotomously branched, or
monopodial, protostelic, the xylem stellate, or mixed with phloem
(haplostelic), or plate-like (plectostelic)
 Leaves small, numerous, sessile, simple with a single central vein,
iso- or less often dimorphic, spirally arranged or decussate
(vegetatively reproducing bulbils may replace the apical leaves in
some species.
Sporophylls contracted or similar to the leaves and aggregated
into fertile regions along the stem, or modified and arranged in
compact strobili, sporangia solitary, borne in the axes of
sporophylls, reniform, bivalvate; spores isomorphic, trilete, smooth
to variously sculptured
A cosmopolitan family of 2 genera, considered to be 5 or more by
some authors, containing over 450 species
The array of descriptive stele types given below is
overwhelming but do not panic, you do not need to memorize
them all. You should become familiar with the protostele, a
solid interior core of xylem surrounded by a cylinder of
phloem, siphonostele, a central pith (parenchyma)
surrounded by a cylinder of vascular tissue, and eustele,
separate vascular bundles in the cortex with phloem to the
outside of the xylem. Eustele is characteristic of dicot
angiosperms. You will also encounter dictyosteles, which are
complex siphonosteles in which the vascular cylinder is broken
up by many leaf gaps. This gives the stem cross section the
appearance of concentric, broken rings. Similarly, actinosteles
are protosteles in which the central vascular strand is lobed;
giving is a star-shaped shape in cross section (e.g. Psilotum
stem). There are many other elaborations on these basic
steles that will crop up occasionally to be on the lookout.
Type of stele: 1. Protostele, 2. Actinostele, 3. Plectostele, 4. Siphonostele, 5.
Eustele, 6. Atactostele, 7. Solenostele, 8. Dictyostle, 9. Polystele. Note that the basic
stele types in vascular plants include protostele, siphonostele, and eustele. Image
taken from http://www.botany.hawaii.edu//.
Family Lycopodiaceae
The Club Moss Family
Consists of about 400 species, mostly tropical.
Terrestrial or epiphytic, moss-like herbs that produce roots directly on
stem and leaves, no ligule (flap at base of each leaf).
Sporophyte with true leaves, stems, and roots.
Dichotomous branching rhizome from which aerial branches and roots
arise.
Stems and roots are protostelic or siphonostelic. Leaf gaps absent.
Leaves are microphyllous and spirally arranged, sometimes opposite or
whorled.
Sporangia borne on upper side of sporophylls which are aggregated
into strobili (strobilus, cluster of overlapping non-photosynthetic
sporophylls) that may be terminal or axillary.
In Huperzia and Phlegmariurus, the sporophylls are similar to ordinary
microphylls and are interspersed among the sterile microphylls.

 One sporangium per sporophyll, near the base and on the adaxial side
Homosporous.
Gametophyte bisexual, either green or subterranean, non-
photosynthetic, mycorrhizal structures, depending on the genus.
Gametangia, antheridia and archegonia, may require 6 to 15 years to
mature.
Self-fertilization is rare.
Biflagellated sperm requires water to reach the archegonium.
Example is Lycopodium which is commonly known as club moss,
ground pine, running cedar.
Genus Lycopodium
Lycopodium is frequently called ground pines or club mosses.
This is because sporangia are carried in structures called strobili
which are "club-shaped" and resemble pine cones.
Its species can be found throughout the world - in a wide range of
habitats but usually growing beneath other plants.
They possess true roots, stems, and leaves.
Their upright stems develop from rhizomes.
The stems are clothed in microphylls.
The leaves may be whorled or in a tight spiral manner.
Adventitious roots arise from the rhizome. Most asexual
reproduction by club mosses occurs via rhizomes.
The sporangia bearing fertile leaves, the sporophylls, may be
spread all over the plant or they may be clustered in a cone-like
strobilus . And they all are homosporous.
Lycopodium is ev­ergreen, as are some ferns, most
gymnosperms, and some angiosperms.
 a foliage plant.
Sporophyte
 Lycopodium may reproduce asexually or through an alteration of
generations involving distinctive gametophytic and sporophytic
generations.
Spores are borne in sporangia in or near the leaf axil, sheathing the
stem. Spores and sporangia of a given species are all alike.
Leaves bearing sporangia are the sporophylls. The sporophylls are
grouped together closely at the ends of stems, forming a cone or
strobilus
A = antheridia
B = archegonia

The spores of Lycopodium germinate to produce a tiny, short-lived, peg-

shaped gametophyte. This usually forms an association with a fungus and
some species of Lycopodium have lost their photosynthetic capacity and
rely on the fungus
to provide carbohydrate! The gametophyte bears antheridia and archegonia
and motile sperm cells swim to the eggs to fertilize these. The young
sporophyte, as for ferns, develops initially as a parasite on the
gametophyte.
Gametophyte

The spores germinate and develop into gametophytes.

Gametophytes in some species grow above ground and
are green; in other species, gametophytes are subterranean
and lack chlorophyll.
The gametophyte is always associated with fungus. Both
male and female gametangia are found on the same
gametophyte.
Gametangia are borne on the upper portion of the
gametophyte.
Fertilization occurs when sufficient free water is present to
allow sperms to swim to mature archegonium.
Life cycle of Lycopodium. Image from http://www.botany.hawaii.ed
Class Selaginellopsida
The vegetative body of Selaginellopsida (e.g. Selaginella) is not
particularly different from that of the other seedless vascular plants in
the division Lycophyta.
The leaves are microphylls with limited cutinized epidermis, mesophyll
only a few cells thick and entirely spongy, and a single haplostelic
vascular bundle.
Typically the stem is dichotomously branched (primitive) and the
spirally-arranged leaves are flattened dorso-ventrally into two morphs.
The stem is held horizontally in most species and is suspended a few
centimeters over the soil on rhizophores.
These branch dichotomously but are positively gravitropic. Upon
contact with soil (thigmomorphogenesis), they dichotomize into a
fibrous root system with root hairs.
At the tips of the branches are found strobili .
In various species the strobili may be held horizontally or may be
vertically aligned. The microphylls in the strobilus are sporophylls. Each
sporophyll has a sporangium in its axil .
The sporangium consists of a stalk and a sterile jacket of cells.
Inside the sterile jacket is one or more sporocytes which ultimately
divide by meiosis to produce spores.
So far this description could be valid for any member of Lycophyta.
Family Selaginellaceae
The Spike Moss Family
about 700 species in this family, most of them are tropical.
are terrestrial, rarely epiphytic, small to moderate size,
stems erect and rooted near the base, and creeping and rooted at
intervals by slender rhizophores that branch dichotomously on
reaching the ground, monopodial with alternate or more or less
dichotomous branching, mostly in one plane.
Selaginella is the only genus in the family, making the family a
monogeneric family.
Characteristics
 Plants small, prostrate, herbaceous, annual or perennial,
sometimes remaining green over winter.
Stems leafy, branching dichotomously, regularly or irregularly
forked or branched
Roots adventitious (produced at the end of small branches called
rhizomorphs); moss-like, rhizomorph producing herbs
dichotomously branched, ligulate .
Stems and roots protostelic (sometimes with many protosteles or
meristeles), siphonostelic, or actino-plectostelic; protostele held in
place by trabeculae.
Rhizophores (modified leafless shoots producing roots) present or
absent, geotropic, borne on stems at branch forks, throughout, or
confined to base of stems.
Leaves microphylls, on one plant dimorphic or monomorphic,
small, with adaxial ligule near base, single-veined, rarely veins
forked.
Strobili sometimes ill-defined, terminal, cylindrical, quadrangular,
or flattened.
Sporophylls (fertile leaves) monomorphic or adjacently different,
microsoporophylls and megasporophylls.
Sporophylls and microphylls with ligule.
Sporangia short-stalked, solitary in axil of sporophylls, opening by dista
slits.
Spores of two types (plants heterosporous), megaspores (1-2-4), large
microspores numerous (hundreds), very small.
Gametophytes develop inside the spore wall: endosporic developmen
are unisexual.
Microgametophyte lacks chlorophyll.
At maturity, it consists of a single prothallial cell and an antheridium;
produces many of biflagellated sperms.
Microspore wall ruptures to liberate the sperms.
Megagametophyte multicellular; protrudes through a rupture in the
spore wall.
Archegonia develop in the exposed area.
Sperms require water to swim to the archegonia.
The suspensor develops and pushes the developing embryo deep into th
female gametophyte.
Genus Selaginella
Life cycle
The sporophyte of Selaginella is the larger of the two genera of living
club mosses.
They are sometimes called spikemosses. Like other members of
lycophytes, club mosses have true roots, stems, and leaves.
They tend to branch more freely than Lycopodium.
have ligules near the base of the leaf.
the strobili of Selaginella reveals that the sporangia are different in
size and color.
four very large yellow spores
These four are the result of meiosis from a single sporocyte. The
spores are called megaspores because of their large size.
This makes the sporangium a megasporangium and the sporophyll a
megasporophyll
The sporangia near the apex of a vertically-held strobilus or
along the upper side of a horizontally-held strobilus are darker
in color (typically orange) and are oval (not lumpy).
These contain many very tiny orange spores, the products of
meiosis of several sporocytes.
These nearly-microscopic spores are called microspores .
This makes their sporangium a microsporangium and the
sporophyll a microsporophyll .
It is not known mechanistically how the strobili achieve their
dimorphic construction with respect to light and gravity vectors.
Selaginella has both microspores and megaspores =the plant is
called heterosporous.
The other fern-allies we have studied were homosporous
This significant advance in evolution is important; sexual
dimorphism is being expressed now much earlier in the life cycle, in
the sporophyte. If the gametophyte is to be reduced and engulfed,
then sexual expression must be shifted to the sporophyte.
the Selaginella sperm must swim through the water-film up the
rhizoids and into the cracked megaspore wall to reach the
megagametophyte inside.
The sperm swim down the open archegonium neck to arrive at the
egg.
The syngamy of egg and sperm results in a zygote.
This develops into an embryo and ultimately an adult sporophyte.
The embryo will demonstrate root and shoot apices. Near its
middle, a few embryonic cells will form a suspensor. These will
elongate to push the embryo deeper into the surrounding
megagametophyte tissue within the megaspore wall.
At this point we have an embryo surrounded by storage tissue
(megagametophyte in this case) and the whole contained in a coating
(megaspore wall). This is almost a seed; but lacks a multicellular seed
coat.
Nevertheless this formation is suggestive of seeds which we will
study in your next higher course, the Seed Plants.
The life history of Selaginella is interesting as it has three
characteristics that are transitional between seedless vascular
plants and the seed plants (gymnosperms and angiosperms):
heterosporous.
endosporic gametophytes .
seed-like embryonic phase.

General habit of Selaginella kraussiana. Image taken from

from (http://www.sinauer.com/)
 Microsporangia

Megasporangia
Three features distinguish Selaginella from Lycopodium:

o Heterospory: Selaginella produces microspores

and megaspores; Lycopodium is homosporous,
producing only microspores.
o Endospory: Selaginella microspores and
megaspores develop their gametophytes within the
actual walls of the spores; in Lycopodium the
gametophyte grows out of the spore.
o Ligule: Each leaf in Selaginella (fertile and sterile)
bears in its axil a tiny tongue-like structure called a
ligule; these are not present in Lycopodium.
Lycopodium sp. and Selaginella sp. strobili comparison. Note the
dissimilarly-sized microspores and megaspores and the ligule on each
sporophyll of Selaginella sp. but absent from Lycopodium. Note also
that the sporangia of Lycopodium is horse-shoe shaped. Image from
http://www.botany.hawaii.edu//.
Life cycle of Selaginella. Image from
http://www.botany.hawaii.edu//.
the sporophyte (1), some branches bear at their tips aggregations
of sporangium-bearing leaves, called sporophylls. They form loose,
cone like structures called strobili (2).
 Each strobilus usually contains two kinds of sporophylls,
microsporophylls bearing microsporangia and megasporophylls
bearing megasporangia.
The outer wall of the sporangia is a sterile jacket of cells that
surrounds and protects the enclosed sporogenous tissue.
 In the microsporangia, the diploid sporocyte cells undergo meiosis
to form numerous spores which in turn form haploid microspores.
In the megasporangium, the entire contents of the
megasporangium form one large megasporocyte that, by meiosis,
produces four haploid megaspores.

Upon rupture of mature microsporangia, the small, lightweight
microspores (3) are readily dispersed by air currents but the larger,
heavier megaspores often remain in place in the strobili.
Germination of a microspore produces a male gametophyte
(microgametophyte) that develops totally within the confines of
the microspore wall (endosporic). At maturity, the entire male
gametophyte consists of a one-cell-layer-thick antheridium (4) and
fertile tissue that forms biflagellated sperm. Sperm are released by
rupture of the microspore wall (6).
Megaspores (3) germinate to begin development of a female
gametophyte (megagametophyte) (5) within the confines of the
megaspore wall (endosporic). Absorptive hairlike structures,
rhizoids which absorb water and dissolved minerals, develop on
the female gametophyte. Archegonia also develop on the exposed
upper surface of the female gametophyte. Each archegonium
contains a single egg, and at maturity, the neck and ventral canal
cells disintegrate to form an open
passageway to the egg (7). Sperm, attracted by substances
released from the matured archegonium, swim through a
film of water and enter the open passageway. One sperm
eventually fertilizes the egg to form a zygote (8).
Development of the embryonic sporophyte occurs within
the archegonium in the female gametophyte. During its
early development as an embryo, the sporophyte is
parasitic upon the female gametophyte. Breakdown of
the parent sporophytic strobili releases the germinated
megaspores (female gametophyte), and the young
sporophytes soon develop a root and shoot system to
become independent plants (9).
 Class Isoetopsida
Submerged or emergent aquatic, sometimes terrestrial in moist
places, herbs with a 2-5 lobed, corm-like rootstock, bearing a
rosette of leaves, and dichotomous roots from between the
lobes. Leaves linear and grass-like, with 4 longitudinal air-canals,
and a single central vascular strand, expanded basally, glabrous, a
small triangular ligule on the adaxial surface above the
sporangium. Sporangia very large, borne adaxially in depressions
at the base of the sporophylls, containing either large, trilete
megaspores, or minute, monolete microspores, sometimes
partially or entirely covered by a membranous vellum, extending
down from the apex of the sporangium.
A class of a single family, which consists of two genera (considered a
single genus by many authors) of more or less cosmopolitan
distribution, with over 125 species
Family Isoetaceae
Quillwort Family
A family of one genus, Isoetes (about 150 spp.), found world-wide,
especially in temperate areas.
Characteristics

Plants tufted grass-like, perennial, evergreen aquatics or emergent

to ephemeral terrestrials.
Underground stem brown, corm-like, lobed (Fig. 3.28).
Roots arising along central groove separating each rootstock lobe,
simple or dichotomously branched, containing eccentric vascular
strand and surrounding lacuna.
Leaves linear, simple, spirally or distichously arranged, dilated
toward base, tapering to apex, containing 4 transversely septate
longitudinal lacunae, a central collateral vascular strand, and
frequently several peripheral fibrous bundles.
Each leaf is a potential sporophyll .
Ligule inserted above sporangium on each sporophyll .
Heterosporous; megasporophylls and microsporophylls usually borne
in alternating cycles; hardened scales and phyllopodia occasionally
surround leaves.
Sporangia solitary, adaxial, embedded in basal cavity of leaf, velum
(thin flap extending downward over sporangium) partly to completely
covering adaxial surface of sporangium .
Megasporangium with several to hundreds of megaspores.
Microsporangium with thousands of microspores.
Megagametophytes white, endosporic, exposed when megaspore
opens along proximal ridges; archegonia one to several, indicated by
quartets of brownish neck cells.
Microgametophytes nine-celled, endosporic, antheridium releasing
four multitailed spermatozoids.
Isoetes, quillwort, is an example.
 leaf

corm

Rhizoids
Isoetes coromandeliana
Only two genera are known, Isoetes and Stylites. They resemble
clumps of grass-like leaves.
Stylites was only discovered in 1957 in the Andes. It is a swamp-dwelling
plant with no stomata, taking in CO2 through its roots!
Isoetes is more widespread.
Image from http://www.botany.hawaii.edu
Genus Isoetes.
oThe genus Isoetes represents one of the most unusual in the division
Lycophyta.
othis genus is small inconspicuous plants may be found in mud, lakes
and ponds.
oThey are heterosporous and monoecious .

DIVISION SPHENOPHYTA: The Scouring Rushes or Horsetails

Introduction
oThe main genus is Equisetum (equis = horse; setum = bristle).
oMonogeneric
oThe stem epidermal cells contain silica,
oused as pot scrubbers or scouring pads= the common name of
“scouring rush”.
ohorsetails (branched) and the scouring rushes (unbranched).
oMost species of Equisetum have an underground rhizome with
vertical stems.
oArial stems (are upright) are jointed and side branches projecting
from the aerial stem resemble a horse’s tail.
oSome species have whorls of small, brownish or grayish (dead)
leaves at the joints instead of branches, thus sort-of resembling
rushes or bamboo .
oThe leaves are slender and are reduced megaphylls (originally
contained more than one strand of vascular tissue).
oSpores are produced in a strobilus at the end of the stem
the gametophyte stage produced from these spores is
inconspicuous.
oGeographic distribution currently is still widespread with the
smaller forms represented (3-4ft) throughout the tropics and
redwood forests, and unevenly through more temperate areas.
These plants may be widespread in moist or damp places. Forms
standing 15 ft tall may still be found.
Evolutionary history
oThey arose some 400 m.y.a. and once reached 18m+ in height (now
herbaceous).
oThey were widely represented in the Carboniferous forests.
oEquisetum is possibly the oldest living plant genus (300 million year
old fossils essentially identical).
oThey are considered to be a separate line derived from the
psilophytes which did not give rise to other plant groups.
oLike lycophytes, these plants protect their sporangia in strobili that
develop at the end of stems.
oThey are distinguished by the unique morphology of their stems.
oThese consist of cylindrical sections separated by nodes and whorls of
slender leaves radiating from the nodes.
oModern sphenopsids are known for rendering themselves
unpalatable by incorporating silica phytoliths into their tissues.
Sphenophyllum sp. Image from http://www.botany.hawaii.edu//.
Economic uses
of little economic significance-, although together with the
lycophytes, they contributed their vegetative parts to the formation
of coal during the Carboniferous period.
Equisetum was used to clean pots and pans, "sand" wooden floors,
and scour plow shears,
thus accounting for its common name of "scouring rush."
Division characteristics
Structure
•Small, usually photosynthetic; heterothallic gametophyte.
•Herbaceous; extant species generally upright ribbed axes
sporophyte.
•Sporangia in strobili of sporangiophores.
•Stems are the primary photosynthetic organs have monopodial
growth habit.
•Mostly homosporous (a few Calamitales are heterosporous).
•Spores have elaters.
•Thick-walled eusporangia usually associated with sporophores
(homology not clear; they may be compound strobili).
•Unusual branching pattern at the nodes.
•Leaves are whorled, reduced. They may be either megaphylls
or microphylls.
•Roots present; and are protostelic.
•Ecology variable, from terrestrial and epiphytic to aquatic and
semi-aquatic; found from tropics to temperate regions.
Classification
othe extant horsetails have 25 species.
oThe Sphenophyta has one class: Equisetopsida or
Sphenospsida and four orders, which consists of extant and
extinct members.
Order Equisetales
a single extant genus, Equisetum.
Fossil history of this group goes back to the late Paleozoic.
reduction in branching and loss of secondary growth. Example:
Equisetum
Family Equisetaceae
Characteristics
•The sporophyte is differentiated into an underground rhizome that
bears adventitious roots and an upright, photosynthetic stem with
whorls of microphylls.
•Tough perennial herbs with jointed, ridged aerial stems with distinct
nodes.
•Stems rough, accumulating silica and metals, and complex
anatomically.
•The aerial stems contain a large central pith region, which in mature
plants is hollow.
•Surrounding the pith cavity are discrete bundles of vascular tissue; this
arrangement of conducting tissue is known as a eustele.
oThe bundles contain both xylem and phloem, and are marked by the
presence of large canals known as carinal canals (under the ridges),
which also function in water conduction.
oExternal to the vascular bundles is another set of canals, the
vallecular canals or cortical canals (under the valleys).
oThese canals line up with the depressions between the ribs on the
surface of the stem. Most fossil sphenophytes had very similar stem
morphology.
oLeaves are small, whorled, non-photosynthetic, fused together to
various degrees and adpressed to the stem.
oSporangia borne on peltate sphorangiophores aggregated into a
strobilus.
oStrobili consist of tightly packed appendages =sphorangiophores.
oHomosporous; spores with elaters.
oGametophytes green, epigeal, bisexual or unisexual, male
gametophytes smaller than female; with rhizoids.
oArchegonia develop before antheridia to increase the probability of
cross-fertilization.
oSperms are multiflagellated and require water to swim to the egg.
Several eggs on a single gametophyte may be fertilized.
Distribution
monogenic, Equisetum (approximately 24 species),
nearly worldwide distribution in damp habitats such as riverbanks,
lakeshores, and marshes.

Genus Equisetum

oEquisetum belongs to the family Equisetaceae

ocharacterized by erect, homosporous, herbs with hollow, jointed,
fluted silicaceous stems, leaves reduced to whorled scales at the
nodes, sporangia borne in terminal strobili , and spores containing
chlorophyll.
ovegetative structure identifies the plant better than its reproductive
structure.
oThe above-ground stems are photosynthetic and contain whorls of
small leaves or branches.
oEquisetum is distinguished from other fern allies by its jointed and
ribbed stem
oIf you examine the living Equisetum plants, you will feel the ribbed
stem of an Equisetum plant.
oIts rough texture results from siliceous deposits in its epidermal cells.
oStrobili of Equisetum, which bears sporangia form atop um­brella-like
structures called sporangiophores , which are modified branches.
oThis is a fertile shoot tip of Equisetum.
oThe sporangiophores each protect a mass of spore producing
sporangia.
oElaters in sporangia of Equisetum help disperse spores.
oThe spores have wings which expand out when dry and aid in
dispersal.
Equisetum.T.S of young stem. Image taken from from (http://www.sinauer.com/)
Equisetum. C.S. of stem (Left at top) plant vegetative morphology (Right). Image
from http://www.botany.hawaii.edu//.
oThe wings or elaters are sensitive to humidity- when they hit a moist
area they coil up again
oAnother unique feature is the presence of Photosynthetic spores.
oThe spores of Equisetum are dark green when alive and are capable
of photosynthesis.
oThis presumably confers some survival advantage on the germinating
prothallus.
oIn addition to photosynthetic pigments, Equisetum spores are
provided with special coiled structures called elaters that extend and
contract with changes in humidity, aiding dispersal.
DIVISION PTEROPHYTA: The Ferns
Introduction
oNext to the flowering plants, ferns, the pterophytes (ptera,
ptero = wing, feather; phyta=plant), are the most diverse
group of living land plants.
oAbout 97% of living Pteridophytes are ferns.
oRecent estimates place their diversity at about 12,000
species in 300 genera in the world.
oMost of these are found in the tropics.
oThe most important is order Filicales, which include all the
homosporous leptosporangiate ferns (Unlike the fern allies
which are a relic group, the ferns are highly successful and are
virtually found in any habitat flowering plants are found.
oThey inhabit almost all kinds of environments, and possess
characteristics of the more advanced seed plants as well as the
less advanced bryophytes.
oThey are widely distributed in both tropics and temperate
zones and are almost exclusively perennial plants.
oThey typically have a horizontal (often underground) stem or
rhizome, swollen with food reserves, from which the leaves and
roots arise.
oThere are also climbing ferns, aquatic ferns, tree ferns, filmy ferns
with leaves just one cell thick and epiphytic ferns with a more
conventional herbaceous habit.
oTheir diversity, which is strik­ing, is in general ranging from
majestic tree ferns to bizarre staghorn ferns. For example, tree
ferns reach heights up to 16 m. Along with other plants, these
ferns once formed forests that were transformed into coal
deposits.
oToday, humans use ferns as decorations, fossil fuels, and in rice
cultivation.
othe sporophyte is the dominant and independent generation to
the extent that this is the generation recognized as a fern.
oIt is with well-developed vascular tissue.
oThe gametophyte (prothallus) is short-lived, a few mm in size and
virtually hidden from view.
oFerns and their allies were the dominant form of vascular plants
until the Mesozoic Era (the age of the Dinosaurs) when seed
bearing plants came into prominence.
o There is fossil evidence of ferns in the Devonian Era (345-395
m.y.a.) and they evolved from the first vascular plants that had
evolved in the Silurian Era (395-435 m.y.a.).
Division characteristics
oSporophyte is differentiated into true roots, stem (rhizome) and
leaves (megaphylls-fronds).
oThe stele varies from a protostele to a complex dictyostele.
oThe xylem is composed of tracheids.
oFronds develop in curled position at the apex of the rhizome
(stem) and uncoil as they mature, a condition called circinate
vernation.
oLeaves usually differentiated into stipe (petiole) and blade with a
central rachis or vein.
oLeaf gaps are conspicuous.
oMost ferns are homosporous; a few aquatic genera are
heterosporous.
oSporangia normally develop on the abaxial (dorsal, lower side) or on
the margin of the leaf.
oGametophytes are inconspicuous and arise directly from spores.
oGametophytes are bisexual in homosporous ferns and unisexual in
heterosporous ferns.
oSperms are flagellated and coiled.
oMost of the ferns that we deal with in this course are
leptosporangiate ferns, which belong to Order Filicales.
Structure
oFerns are more complicated in structure than most people would
suspect.
oTheir structures, though similar in some ways to those of flowering
plants are different enough to warrant a distinctive terminology.
oThe most conspicuous part of the fern sporophyte is the leaf.
oThe typical fern leaf, often termed a frond, is an elaborate structure
composed of numerous leaflets
oThe frond arises from the stem which is a rhizome growing on or just
beneath the soil surface.
oYoung fronds are tightly coiled and are commonly referred to as
"Fiddleheads".
oThe rhizome also bears numerous roots, termed adventitious roots
since they arise from stem tissue.
oThis distinguishes them from the primary root (derived from the
embryo) and secondary or lateral roots (derived from a pre-existing
root).
oAll tissues of the shoot (leaf and stem) are derived from a single
meristematic cell located at the apex of the rhizome.
oSimilarly all tissues of the root are derived from a single meristematic
cell located at the apex of the root.
oThe fronds, the rhizomes and the roots make up the sporophyte
generation.
 Frond

Fiddle head

Rhizome

Features of a mature fern, leaves, rhizome and young leaf (fiddlehead).

 Sori

Dryopteris sp. A. Habit of sporophyte. B. Fertile frond. Note that all

above ground part is a frond. Image taken from from (
http://www.sinauer.com/)
Pteridium sp. Fern habit with rhizome and frond
Roots
Roots are formed from rhizomes , all developing from apical cells,
commonly near leaf bases, or sometimes from the stipe are
adventitious .
The roots usually do not divide once they grow from the rhizome.
Tree fern roots grow down from the crown and help thicken and
strengthen the trunk. As in other plants, the roots anchor the plant to
the ground and absorb water and minerals.
Rhizome
The stems of Filicales are mainly true rhizomes, i.e., stems that run
horizontally at or just beneath the surface of the ground.
Some have upright stems like tiny tree trunks (Cyatheceae and
Decksoncaceae) and are unable to form colonies except for occasional
upright branching.
Rarely, Filicales may have both upright stems and horizontal rhizomes.
In many cases, the rhizome can be inconspicuous or even entirely
underground. Rhizomes of tree ferns on the other hand may be 60 cm
in diameter and up to 12 m tall.
In some epiphytic ferns (ferns that grow in trees) and in
terrestrial creeping ferns the rhizome roams widely and is quite
visible .
The rhizomatous habit of many ferns results in extensive
vegetative reproduction. Thus, it should be noted that although
fern stems are often called rhizomes, strictly speaking this term
refers to those with a long-creeping or climbing habit.
If the stem is short and compact and radially symmetric, erect,
prostrate or ascending, it is called a caudex or rootstock (or
simply stock).
In some cases the caudex is erect and tree-like with an apical
radial tuft of leaves (Cyathea, Dicksonia, Leptopteris, some
Blechnum, Thelypteridaceae) in which case it is often called a
trunk.
Creeping rhizomes can be either long-creeping with widely
spaced fronds or short-creeping with fronds relatively close
together.
The rhizomes can be radial or radially symmetric or
dorsiventral with leaves or fronds produced on the dorsal
or upper side and roots produced on ventral or lower
side. That is, fronds arise from the rhizome. The fronds
that arise from this "stem" arise from the upper side or
occur in one or more rows laterally on each side.
The rhizome contains the conducting tissues (xylem
and phloem) and the strengthening tissues
(sclerenchyma fibers.
The conducting tissue, known as the vascular bundle,
carries the water, minerals, and nutrients throughout the
plant .
The stele in juvenile sporophytes is usually a protostele.
As the stem increases in size a siphonostele may develop.
Anatomically, the stems of pteridophytes are simpler
than in most seed plants because they are made up only
of primary tissues.
Rhizome branching
Overall, the rhizome or caudex may be unbranched
(most compact rhizomes are unbranched) or branched in
various ways.
Creeping rhizomes may be unbranched or branched
irregularly. Sometimes the branching is regularly
dichotomous (or isotomous), sometimes with alternate
branches reduced to a great or lesser extent
(anisotomous). Sometimes the fork or branching is
associated with the production of a leaf or frond in the
axil.
The rhizomes or caudices of some species that do not
usually branch (e.g. tree ferns) may sometimes be
induced to branch following physical damage. The
branching patterns of the stem of some groups of
pteridophytes are quite distinctive and may be used
taxonomically.
Rhizome protection
The rhizomes of ferns well protected from the extremes of
temperature, desiccation, predation and physical
abrasion( epidermal hairs, bristles or scales, or a
combination of any of these. secreting a mucilaginous
slime that engulfs the growing tip and the developing
leaves (e.g. Plagiogyria). Sometimes the surface of the
rhizome is glaucous with a bluish grey waxy sheen and
sometimes with a dense white waxy covering; this often
useful diagnostic character may be lost with collecting and
preservation techniques involving excessive heat or
solvents such as ethanol.
Bulbils
Bulbils are vegetative buds for reproduction.
Fern fronds
Fronds develop in a distinctive manner
young fronds developing by uncurling along the main axis from
the base towards the tip; this type of vernation (leaf
development) is the circinnate vernation
distinctive characteristic of ferns; the uncurling frond is a
fiddlehead or crosier .
The crozier is produced by a process of growth and unfolding.
Leaf development that is initiated at the tip is called acropetal,
and this development is characteristic of most modern ferns.
Leaves of seed plants, in contrast, tend to develop by overall
growth and expansion, i.e., by intercalary differentiation and
enlargement.
The frond is a fern leaf attached to the main axis of growth; the
point of attachment may be below or above ground .
two main parts, the stipe (leaf stalk or petiole) and the
blade (the leafy expanded portion of the frond).
The blade form and shape varies depending on the species.
Besides, fronds vary greatly in size, from tree ferns with 12
foot fronds to the mosquito ferns with fronds only 1/16 of
an inch long.
The branching of a fern frond is nearly always in the one
plane and the horizontal twisting of the secondary
branches of scandent species.
Fern fronds are either simple or compound and composed
of several to many leaflets which may in turn by composed
of several to many leaflets, and so on until the ultimate
division is a simple leaflet or lobe.
A simple lamina is continuous on either side of the midrib,
but may be shallowly or deeply dissected or lobed or
unlobed and entire.
A frond axis is the central supporting tissue about which the
lamina and/or subsequent axes are arranged. The main axis
consists of the stipe (if present) and its continuation into the
lamina of the frond; in simple fronds this is the midrib or costa,
in compound fronds, the rachis.
The rachis is the extension of the petiole into the blade
The blade may be undivided (simple) or divided multiple
times . Most commonly, the leaflets or secondary rachises are
borne alternately.
This is the pinnate condition and the leaflets are known as
pinnae (pinna-singular); the pinnae themselves may be
similarly divided and the frond is described as being bipinnate
and the pinna of each leaflet is known as a pinnule.
tripinnate or even quadripinnate fronds; in each case the
ultimate simple division is referred to as a pinnule
 If the lamina is only partially incised towards the axis, the
segment is described as being pinnatifid (the term
pinnatisect describes very deep incisions, but this distinction
is not often drawn). In a similar manner to pinnate division,
it is possible to have bipinnatifid or tripinnatifid fronds. It is
possible to have combinations of these forms of dissection
and such structures as pinnate-bipinnatifid or bipinnate-
pinnatifid etc. are common. Finely divided fronds are often
described with the general term of decompound. For
example, some species possess pinnatifid pinnae . For
example, a pinnate-pinnatifid fern has pinnae that are
pinnate, attached to the rachis as in , but each pinna is itself
pinnatifid. Another type of division is one where the green
leafy tissue is not completely separated from the rachis but
rather it spreads along the rachis, instead this degree of
division is called "pinnatifid" .
Midrib/costa

pinnae
stipe
Rachis
 Stipe
Leaf blade

Palmate leaf blade. Image taken from from (http://www.sinauer.com/)

Palmate blades have pinnae that are directly attached to the

end of the petiole and extend outward from the center of the
frond, extending like fingers from a person's palm

Stipe
Stipe bases generally bear the same type of hairs or
scales that cover the stem and are similarly very important
taxonomically
.
Venation
The vascular traces of the stipe and axes eventually end up in
the venation of the lamina (leaf blade) and the manner and
pattern in which this is achieved is particularly important in fern
classification. Veins are the ultimate vascular strands (and their
supporting structure) running through the leaf, and are for the
most part clearly visible on either surface of the lamina, being
raised or impressed with cells of different color, shape and
orientation of the rest of the lamina.
Veins with no branches or unions are called simple.
Frond dimorphism
In addition to differences that may exist between sterile and
fertile leaves juvenile leaves are sometimes different to the adult
leaves.
In conclusion, two features of ferns especially stand out:
 Large, typically pinnate,
fronds bearing
sporangia
Fern showing sori on
underside of leaf.

Circinate vernation with spiral like

a shepherd's crook or bishop's
crozier or a fiddle head.
The leaves also bear sporangia grouped together
in roundish sori (see the above photograph) and
covered by protective kidney-shaped indusium.
Such leaves are regarded as fertile fronds.
Reproductive structures
Underneath a fern blade, you may find small spore
casings containing many individual spores. The
spore casings are the sporangia, and clusters of
sporangia are the sori
Sori typically form distinct patterns that are helpful
for identifying different species. Sori may be
shaped like circular dots as in, or be in short or
long linear lines. Sori may follow the margins,
midveins, or venation patterns of the pinnae or
pinnules. Sometimes, sporangia completely cover
the undersides of the pinnae
Osmunda regalis. A. Frond. B. Fertile pinnae. C. Sporangium. D. Fertile pinna of Leptopteris. Image
taken from from (http://www.sinauer.com/)
Gametophyte
The ferns immature stage is called a prothallus.
Terrestrial ferns are homosporous (undergo
homospory). The fern gametophyte is a macroscopic
thallus that is free living and independent of the
sporophyte phase. The gametophyte thallus is
composed of parenchyma cells containing chloroplasts,
ventral rhizoids, archegonia and antheridia
The young sporophyte, derived from the zygote, consists
of a primary leaf, a primary root, a rhizome growing point
(shoot apical meristem) and a foot. The sporophyte
remains attached to the gametophyte thallus until the
young sporophyte becomes functionally independent.
Soon after the gametophyte thallus degenerates. e.
The gametophytes may be filamentous, ribbon like
cylindrical or cordate
The gametophytes of almost all Filicales are bisexual