Course Name-Biology and Diversity of Algae,

Bryophyta and Pteridophyta

(Paper Code: BOT 502)

Unit –11 : General Characters, Classification, Distribution

and Economic Importance of Bryophytes

Dr. Pooja Juyal

Department of Botany
Uttarakhand Open University,
Haldwani
Email id: [email protected]
CONTENT
Introduction
General Characteristics
Distribution
Habitat
Life Cycle
Reproduction
Economic Importance
INTRODUCTION
Bryophytes (nonvascular plants) are the only embryophytes (plants that
produce an embryo) whose life history includes a dominant gametophyte
(haploid) stage. They are an ancient and diverse group of non-vascular
plants. Bryophyta (Gr. Bryon = mass; phyton = plant), a division of
kingdom Plantae comprises of Mosses, Hornworts and Liverworts. They are
groups of green plants which occupy a position between the thallophytes
(Algae) and the vascular cryptogams (Pteridophytes). Bryophytes produce
embryos but lack seeds and vascular tissues. They are the most simple and
primitive group of Embryophyta. Bryophytes grow in two habitats i.e. water
and land so known as amphibians of plant kingdom. They are said to be the
first land plants or non-vascular land plants. They cannot reproduce without
sufficient moisture because without water sex organs neither matures nor
dehisces. Presence of swimming antherozoids is an evidence of their aquatic
ancestory. They comprise three main taxonomic groups: mosses
(Bryophyta), liverworts (Marchantiophyta or Hepatophyta) and hornworts
(Anthocerotophyta) which have evolved quite separately. Most bryophytes
have erect or creeping stems and tiny leaves, but hornworts and some
liverworts have only a flat thallus and no leaves. There are possibly 25,000
species of mosses and liverworts all over the world.
GENERAL CHARACTERISTICS
1. Small group of primitive land dwellers having small leafy or
thalloid green plant body.
2. Like thallophytes plant body is gametophytic, independent,
dominant, autotrophic, either thalloid (i.e., thallus like, not
differentiated into root, stem and leaves) or foliose, containing a
rootless leafy shoot.
3. Plant body is very small and ranges from a few mm to many cm.
Zoopsis is the smallest bryophyte (5 mm.) while the tallest bryophyte
is Dawsonia (50-70 cms.).
4. Leaves and stems found in vascular plants are absent, these
‘leaf’ and ‘stem’ like structures as ‘axis’ and ‘phylloid’ respectively.
5. Roots are absent. Functions of the roots are performed by rhizoids.
Cells are also capable to absorb moisture directly from the ground or
atmosphere. Therefore, Bryophytes can also survive on the moist
soils.
6. Rhizoids may be unicellular, un-branched (e.g., Riccia,
Marchantia, Anthoceros) or multicellular and branched (e.g.,
Sphagnum, Funaria).
7. In members of order Marchantiales (e.g., Riccia, Marchantia)
scales are present. These are violet coloured, multicellular and single
cell thick. They protect the growing point and help to retain the
moisture.
8. Vascular tissue (xylem and phloem) is completely absent. Water
and food material is transferred from cell to cell. However, in some
Bryophytes (e.g., mosses) a few cells in groups of 2-3 are present for
conduction of water and food (photo assimilate). These cells are
known as hydroid (collectively hydrom) and leptoids respectively.
Cuticle and stomata are absent.
9. Sexual reproduction is invariably highly oogamous. The sex organs
are jacketed and multicellular while in algae they are non-jacketed
and unicellular.
10. Female sex organ is archegonium appears for the first time in
bryophytes.
11. Sperms are biflagellate and both the flagella are of whiplash type.
12. Fertilization takes place in the presence of water or moisture.
13. Fertilized egg remains in the venter of the archegonia. It neither
becomes independent from parent gametophyte nor passes into resting
period. In both these respect the bryophyte differs from algae.
14. Zygote undergoes repeated divisions to form an undifferentiated
multicellular structure called the embryo.
15. First division of the zygote is transverse and the apex of the embryo
develops from the outer cell. Such embryogenesis is called exoscopic.
16. The venter wall enlarges to produce a protective multicellular
envelop called calyptra.
17. The embryo by further division and differentiation produce a
relatively small spore producing structure which is not independent. It is
called sporogonium (sporophyte).
18. Sporophyte is rootless and consists of foot, seta and capsule. In some
seta is absent (Corsinia) and rarely the both foot and seta (Riccia).
19. Sporophyte remains attached with gametophyte throughout its life
and also depends on it partially or wholly for nutrition.
20. Sporophyte produces spores which are wind disseminated, non-motile
and cutinized, also called meiospores.
21. Morphologically all the meiospores in a given species are alike, thus
known as homosporous.
22. Each spore under suitable conditions germinates to give rise
gametophyte plant directly or indirectly as lateral bud from protonema.
23. Heterologous type of alternation of generation in the life cycle of
bryophytes while in algae it is of homologous type.
Fig. External Features of Bryophytes (A) Riccia rosette (B) Riccia Thallus (C) Marchantia
(D) Porella (E) Anthoceros (F) Funaria (G) Polytrichum
DISTRIBUTION
Bryophytes are distributed throughout the world, from polar and alpine
regions to the tropics. Water must, at some point, be present in the habitat
in order for the sperm to swim to the egg. Bryophytes do not live in
extremely arid sites or in seawater, although some are found in
perennially damp environments within arid regions and a few are found
on seashores above the intertidal zone. A few bryophytes are aquatic.
Bryophytes are most abundant in climates that are constantly humid and
equable. The greatest diversity is at tropical and subtropical latitudes.
Bryophytes (especially the moss Sphagnum) dominate the vegetation of
peatland in extensive areas of the cooler parts of the Northern
Hemisphere.
The geographic distribution patterns of bryophytes are similar to those of
the terrestrial vascular plants, except that there are many genera and
families and a few species of bryophytes that are almost cosmopolitan.
Indeed, a few species show extremely wide distribution.
Some botanists explain these broad distribution patterns on the theory
that the bryophytes represent an extremely ancient group of plants,
while others suggest that the readily dispersible small gemmae and
spores enhance wide distribution. The distribution of some
bryophytes, however, is extremely restricted, yet they possess the
same apparent dispersibility and ecological plasticity as do
widespread bryophytes. Others show broad interrupted patterns that
are represented also in vascular plants.
Bryophytes are represented by 960 genera and 25,000 species. They
are cosmopolitan in distribution and are found growing both in the
temperate and tropical regions of the world at an altitude of 4000-
8000 feet. In India, Bryophytes are quite abundant in both Nilgiri hills
and Himalayas; Kullu, Manali, Shimla, Darjeeling, Dalhousie and
Uttarakhand are some of the hilly regions which also have a luxuriant
growth of Bryophytes.
Eastern Himalayas have the richest in bryophytic flora. A few species of
Riccia, Marchantia and Funaria occur in the plains of U.P., M.P.
Rajasthan, Gujarat and South India.
In hills they grow during the summer or rainy season. Winter is the rest
period. In the plains the rest period is summer, whereas active growth
takes place during the winter and the rainy season. Some Bryophytes
have also been recorded from different geological eras e.g., Muscites
yallourensis (Coenozoic era), Intia vermicularies, Marchantia spp.
(Palaeozoic era) etc.

HABITAT: Bryophytes grow densely in moist and shady places and

form thick carpets or mats on damp soils, rocks, bark of trees especially
during rainy season. Small in size, but they can be very conspicuous
growing as extensive mats in woodland, as cushions on walls, rocks and
tree trunks, and as pioneer colonists of disturbed habitats.
Majority of the species are terrestrial but a few species grow in fresh
water (aquatic) e.g., Riccia fluitans, Ricciocarpos natans, Riella etc.
Bryophytes are not found in sea but some mosses are found growing in
the crevices of rocks and are being regularly bathed by sea water e.g.,
Grimmia maritima. Some Bryophytes also grow in diverse habitats e.g.,
Sphagnum-grows in bogs, Dendroceros-epiphytic, Radula protensa -
epiphyllous, Polytrichum juniperinum-xerophytic, Tortula muralis-on
old walls. Tortula desertorum in deserts, Porella platyphylla-on dry
rocks, Buxbaumia aphylla (moss), Cryptothallus mirabilis (liverwort) are
saprophytic.

LIFE CYCLE: The life cycle of bryophytes shows two distinct

phases namely a haploid gametophytic phase and a diploid sporophytic
phase alternating with each other. The adult plant body represents
the gametophyte. A short-lived sporophyte occurs as a parasite on the
gametophyte.
Gametophyte: A stage in the life cycle of bryophyte that undergoes alternation
of generations. It is a haploid multicellular organism that develops from
a haploid spore that has one set of chromosomes. The gametophyte is the sexual
phase in the life cycle of bryophytes. It develops sex organs that
produce gametes, haploid sex cells that participate in fertilization to form a
diploid zygote in which each cell has two sets of chromosomes. Cell division of
the zygote results in a new diploid multicellular organism, the second stage in
the life cycle known as the sporophyte, the function of which is to produce
haploid spores by meiosis.

In bryophytes (mosses, liverworts, and hornworts), the gametophyte is the most

visible stage of the life cycle. The bryophyte gametophyte is longer lived,
nutritionally independent, and the sporophytes are typically attached to the
gametophytes and dependent on them. It is the adult plant body in bryophytes.
It is either thalloid or in the form of a leafy shoot with stem-like and leaf-like
structures. Roots are absent and instead thread-like rhizoids are present.
Vascular tissues xylem and phloem are absent.
Water and food are directly transported from a cell to cell. Vegetative
reproduction may sometimes occur by fragmentation. However, sexual
reproduction is common and is of oogamous type. The mature gametophyte
bears male reproductive organs called antheridia and female reproductive
organs called archegonia. The antheridia have a club-shaped body and a stalk.
They produce flagellated male gametes called antherozoids or sperms. The
archegonia are flask-shaped with a well-defined venter and neck. The venter
encloses a venter canal cell and an egg cell while the neck encloses a variable
number of neck canal cells.

The antherozoids liberated from antheridia, swim in a film of water and reach
the archegonia. They are attracted into the archegonia to bring about
fertilization. The zygote develops into the sporophyte.
Fig. Moss Life Cycle
Sporophyte: Zygote represents the first cell of the sporophytic phase. It
divides and develops into a sporophytic plant body called sporogonium. It is
neither independent of the parent gametophyte nor passes into the resting
phase. In both respects differs from the zygote of green algae. Further
development of zygote into embryo occurs within venter of the
archegonium. Zygote undergoes segmentation without a resting period into
multicellular, undifferentiated structure called embryo. Embryo by further
segmentation and differentiation finally develops into full-fledged
sporophyte called sporogonium. The wall of venter forms a protective
covering to the sporogonium, called calyptra. Sporogonium in bryophytes is
leafless and rootless. Most often, the sporogonium is differentiated into a
foot, a seta and a capsule. It remains attached throughout its life to the
gametophytic host with the help of foot. It absorbs nutrients directly from
the gametophyte. In some bryophytes the foot is much reduced and its
function is performed by haustorial collar which develops from the junction
of reduced foot and seta.
Seta conducts the absorbed food to the capsule. The terminal capsule is
considered equivalent to fern sporangium is mainly concerned in the
production of spores which are nonmotile and wind disseminated.
The sporogonium produces haploid spores (meiospores) which get
released from the capsule. Spores are highly specialized cells
differentiated from the diploid spore mother cell by meiosis. They are
thus haploid cells. The spores represent the first cells of gametophytic
generation. They germinate under suitable condition to produce new
gametophytes either directly or through a juvenile stage called
protonema.
Another remarkable feature in the lifecycle of bryophyte in which they
differ from thallophytes is the complete absence of asexual spores called
the mitospores in the life cycle. Asexual reproduction takes place only by
the vegetative method of fragmentation and gemmae.
REPRODUCTION IN BRYOPHYTES
Bryophytes reproduce only by vegetative and sexual means. Asexual reproduction is
absent in these.

A. Vegetative Reproduction in Bryophytes:

Bryophytes possess a characteristic feature and that is their tendency towards extensive
vegetative reproduction. The vegetative reproduction takes place in favourable season for
vegetative growth. Majority of the Bryophytes propagate vegetatively and it is brought about
in many ways.
1. By Death and Decay of the Older Portion of Thallus or by Fragmentation:
In Bryophytes the growing point is situated at the tip of the thallus. The basal, posterior or
older portion of the thallus starts rotting or disintegrating due to ageing or drought. When
this process of disintegration or decay reaches up to the place of dichotomy, the lobes of the
thallus get separated. These detached lobes or fragments develop into independent plants by
apical growth. This is the most common method of vegetative reproduction in Riccia,
Marchantia, Anthoceros and some mosses like Sphagnum
2. By Persistent Apices:
Due to prolonged dry or summer or towards the end of growing season the whole thallus in
some Bryophytes (e.g., Riccia, Anthoceros, Cyathodium) dries and get destroyed except the
growing point. Later, it grows deep into the soil and becomes thick. Under favourable
conditions it develops into a new thallus.
3. By Tubers:
Tubers are formed in those species which are exposed to desiccations (drying effect of the
air). Towards the end of the growing season, the subterranean branches get swollen at their
tips to from the underground tubers. On the periphery of a tuber are two to three layers of
water proof corky, hyaline cells develop.
These layers surround the inner cells which contain starch, oil globules and albuminous
layers. During the unfavorable conditions the thallus dies out but the dormant tubers remain
unaffected. On the return of the favourable conditions each tuber germinates to form a new
plant e.g., Riccia, Anthoceros, Fossombronia etc. Thus, tubers also serve as organ of
perennation.
4. By Gemmae:
Gemmae are green, multicellular reproductive bodies of various shapes. These are produced
in gemma cups, on the surface of the leaves, on stem apex or even inside the cells. They get
detached from the parent plant and after falling on a suitable substratum gemmae give rise to
a new individual directly (e.g., Marchantia) or indirectly (e.g., Mosses).
5. By Adventitious Branches:
The adventitious branches develop from the ventral surface the thallus e.g., Riccia fluitans,
Anthoceros. On being detached from the parent plant these branches develop into new thalli.
In Marchantia, Dumortiera these branches develop from archegoniophore while in Pellia
these branches arise from the dorsal surface or margins of the thallus.
6. By Regeneration:
The liverworts possess an amazing power of regeneration. Part of the plant or any living cell
of the thallus (e.g., rhizoid, scales).are capable of regenerating the entire plant e.g., Riccia,
Marchantia etc.
7. By Innovation:
In Sphagnum one of the branches in the apical cluster instead of forming drooping branches
or divergent branches, develop more vigorously than the others and continues the growth
upwards. This long upright branch has all the characteristics of main axis. It is called
innovation. Due to progressive death and decay of the parent plant these innovation become
separated from the parent plant and establish themselves as parent plants.
8. By Primary Protonema:
Primary protonema is the filament like stage produced by the developing spores of the
mosses. It produces the leafy gametophores. It breaks into short filament of cells by the
death of cells at intervals. Each detached fragment grows into a new protonema which bears
a crown of leafy gametophores e.g., Funaria.
9. By Secondary Protonema:
The protonema formed by other methods than from the germination of spores is called
secondary protonema. It may develop from any living cells of the leafy gametophore i.e.,
from leaf, stem, rhizome, injured portion of the leafy gametophore, antheridium, paraphysis
or archegonium. From this arise the leafy gametophores or lateral buds in the same manner
as in primary protonema e.g., Funaria, Sphagnum.
10. By Bulbils:
These are small resting buds develop on rhizoids. Bulbils are devoid of chlorophyll but full
of starch. On germination bulbils produce a protonema which bears leafy gametophores
11. By Apospory:
The production of diploid gametophyte from the unspecialized sporophyte without meiosis
is known as apospory e.g., Anthoceros. In Funaria green protonemal filaments may arise
from the unspecialized cells of the various parts of sporogonium. These protonemal
filaments bear lateral buds which develop into leafy gametophores.
12. By Rhizoidal Tips:
The apical part of the young rhizoids divide and re-divide to form a gemma like mass of
cells e.g., Riccia glauca. These cells contain chloroplast and are capable to develop into new
thallus.

B. Sexual Reproduction in Bryophytes:

1. Sexual reproduction is highly oogamous.
2. Male and female sex organs are known as antheridia (Sing, antheridium) and
archegonia (Sing, archegonium), respectively.
3. Sex organs are jacketed and multilayered.
4. Antheridium is stalked, pear shaped or oblong and has an outer one cell thick jacket
which encloses a mass of fertile cells called androcytes. Each androcyte metamorphoses
into biflagellate antherozoid.
5. Archegonium is stalked, flask shaped structure. It has a basal swollen portion called venter and
an elongated neck. The neck is filled with many neck canal cells whereas venter has a large egg
cell and a small venter canal cell,
6. Antherozoids are attracted towards the neck of the archegonium chemotactically by certain
substances (like sugars, malic acid, proteins, inorganic salts of potassium etc.) present in the
mucilaginous substance formed by the degeneration of neck canal cells and venter canal cell.
7. Water is essential for fertilization.
8. The fertilized egg or zygote is the beginning of the sporophytic phase. It is retained within the
venter of the archegonium.

Sporophyte:
1. Without resting period, the zygote undergoes repeated divisions to form a multicellular
structure called the embryo.
2. The first division of the zygote is always transverse and the outer cell develops into embryo.
Such an embryogeny is called exoscopic.
3. Embryo develops into a sporophyte or sporogonium.
4. The sporophyte is usually differentiated into foot, seta and capsule. In certain cases it is
represented only by capsule (e.g., Riccia) or by foot and capsule (e.g., Corsinia).
5. Sporophyte is attached to parent gametophytic plant body throughout its life. It partially or
completely depends on it for nutrition.
6. Foot is basal, bulbous structure. It is embedded in the tissue of parent gametophyte. Its main
function is to absorb the food material from the parent gametophyte.
7. Seta is present between the foot and capsule. It elongates and pushes the capsule through
protective layers. It also conducts the food to the capsule absorbed by foot.
8. Capsule is the terminal part of the sporogonium and its function is to produce spores
9. All Bryophytes are homosporous i.e., all spores are similar in shape, size and structure
10. Capsule produces sporogenous tissue which develops entirely into spore mother cells it
e.g., Riccia) or differentiated into spore mother cells and elater mother cells (e.g.,
Marchantia, Anthoceros).
11. Spore mother cells divide diagonally to produce asexually four haploid spores which are
arranged in tetrahedral tetrads.
12. Elater mother cells develop into elaters (e.g., Marchantia) or pseudo elaters (e.g.,
Anthoceros which are hygroscopic in nature. Elaters are present in liverworts and absent in
mosses.
13. Venter wall enlarges with the developing sporogonium and forms a protective
multicellular layer called calyptra (gametophytic tissue enclosing the sporophyte).

Young Gametophyte:
1. The meiospore (spore formed after meiosis) is the first cell of the gametophytic phase.
2. Each spore is unicellular, haploid and germinates into young gametophytic plant (e.g.,
Riccia or Marchantia) or first germinates into a filamentous protonema on which buds are
produced to give rise to a young gametophytic plant, (e.g., Funaria).
ALTERNATION OF GENERATION: The life-cycle of a bryophyte shows
regular alternation of gametophytic and sporophytic generations. This process of
alternation of generations was demonstrated for the first time in 1851 by Hofmeister.
Thereafter in 1894 Strassburger could actually show the periodic doubling and halving
of the number of chromosomes during the life-cycle.

The haploid phase (n) is the gametophyte or sexual generation. It bears the sexual
reproductive organs which produce gametes, i.e., antherozoids and eggs. With the result
of gametic union a zygote is formed which develops into a sporophyte. This is the
diploid phase (2n). The sporophyte produces spores which always germinate to form
gametophytes.
During the formation of spores, the spore mother cells divide meiotically and haploid
spores are produced. The production of the spores is the beginning of the gametophytic
or haploid phase. The spores germinate and produce gametophytic or haploid phase.
The spores germinate and produce gametophytes which bear sex organs.
Ultimately the gametic union takes place and zygote is resulted. It is diploid (2n). This
is the beginning of the sporophytic or diploid phase. This way, the sporophyte
generation intervenes between fertilization (syngamy) and meiosis (reduction division);
and gametophyte generation intervenes between meiosis and fertilization.
Fig. Bryophytes life cycle (Graphic Representation)
ECOLOGICAL AND ECONOMIC IMPORTANCE
Bryophytes are of great ecological importance due to following reasons:
(a) Pioneer of the land plants. Bryophytes are pioneer of the land plants because they
are the first plants to grow and colonize the barren rocks and lands.
(b) Bryophytes prevent soil erosion. They usually grow densely and hence act as soil
binders. Mosses grow in dense strands forming mat or carpet like structure.
They prevent soil erosion by:
(i) Bearing the impact of falling rain drops
(ii) Holding much of the falling water and reducing the amount of run-off water.
(c) Formation of soil- Mosses and lichens are slow but efficient soil formers. The acid
secreted by the lichens and progressive death and decay of mosses help in the formation of
soil.
(d) Bog succession- Peat mosses change the banks of lakes or shallow bodies of water into
solid soil which supports vegetation e.g., Sphagnum
(e) Rock builders- Some mosses in association with some green algae (e.g., Chara) grow in
water of streams and lakes which contain large amount of calcium bicarbonate. These
mosses bring about decomposition of bi-carbonic ions by abstracting free carbon dioxide.
The insoluble calcium carbonate precipitates and on exposure hardens, forming calcareous
(lime) rock like deposits.
Economic importance
1-Formation of Peat:
Peat is a brown or dark colour substance formed by the gradual compression and
carbonization of the partially decomposed pieces of dead vegetative matter in the bogs.
Sphagnum is an aquatic moss. While growing in water it secretes certain acids in the water
body.
This acid makes conditions unfavorable for the growth of decomposing organisms like
bacteria and fungi. Absence of oxygen and decomposing microorganisms slows down the
decaying process of dead material and a large amount of dead material is added year by year.
It is called peat (that is why Sphagnum is called peat moss).
2-As Packing Material:
Dried mosses and Bryophytes have great ability to hold water. Due to this ability the
Bryophytes are used as packing material for shipment of cut flowers, vegetables, perishable
fruits, bulbs, tubers etc.

3- As Bedding Stock:
Because of great ability of holding and absorbing water, in nurseries beds are covered with
thalli of Bryophytes.
4. In Medicines:
Some Bryophytes are used medicinally in various diseases for e.g.,
(a) Pulmonary tuberculosis and affliction of liver—Marchantia spp.
(c) Acute hemorrhage and diseases of eye—Decoction of Sphagnum.
(d) Stone of kidney and gall bladder—Polytrichum commune.
(e) Antiseptic properties and healing of wounds—Sphagnum leaves and extracts of some
Bryophytes for e.g., Conocephalum conicum, Dumortiera, Sphagnum protoricense, S.
strictum show antiseptic properties.
5. In Experimental Botany:
The liverworts and mosses play an important role as research tools in various fields of
Botany such as genetics. For the first time in a liverwort, Sphaerocarpos, the mechanism of
sex determination in plants was discovered.

6. As Food:
Some Bryophytes e.g., mosses are used as food by chicks, birds and Alaskan reindeer etc.
CLASSIFICATION OF BRYOPHYTES
The term Bryophyta was first introduced by Braun (1864); however, he included algae,
fungi, lichens and mosses in this group. Later, algae, fungi and lichens were placed in a
separate division Thallophyta and liverworts, mosses in division Bryophyta.
Eichler (1883) was the first to divide Bryophyta into two groups:
Group I. Hepaticae
Group II. Musci.

Engler (1892) recognised Hepaticae and Musci as two classes and divided each class
into the following three orders:
Division- Bryophyta:
Class I. Hepaticae: Divided into three orders:
Order 1. Marchantiales
Order 2. Jungermanniales
Order 3. Anthocerotales
Class II. Musci: Divided into three orders:
Order 1. Sphagnales
Order 2. Andreaeales
Order 3. Bryales
International code of Botanical Nomenclature (ICBN) suggested in 1956-that the suffix-
opsida should be used for the classes and such usage had already been proposed by
Rothmaler (1951) for the classes of Bryophytes. He changed the class names as:
Class I. Hapaticae as Hepaticopsida.
Class II. Anthocerotae as Anthoceropsida
Class III. Musci as Bryopsida.

Proskauer (1957) suggested that the class name Antheoceropsida should be

Anthocerotopsida. Parihar (1965) and Holmes (1986) followed Proskauer’s
classification and divided Bryophyta into three classes:
Clase I. Hepaticopsida
Class II. Anthocerotopsida
Class III. Bryopsida.
Class I. Hepaticopsida (Liverworts):
General Characters:
1. This class includes about 280 genera and 9500 species.
2. The name of this class is derived from a Latin word Hepatica which means liver.
Hence members of this class are commonly known as liverworts.
3. Plant body is gametophytic and the gametophyte is either thalloid or foliose.
4. Thalloid forms are prostrate, lobed, dorsiventral and dichotomously branched.
5. In foliose forms, ‘leaves’ are entire, lobed or divided and without ‘midrib’. ‘Leaves
arranged in two to three rows on the axis.
6. Rhizoids are unicellular and branched.
7. Photosynthetic cells contain many chloroplasts.
8. Pyrenoids are absent.
9. Sex organs are borne dorsally or apically, superficial or embedded in gametophytic
tissue
10. Members may be monoecious or dioecious.
11. Sporophyte is either simple or represented by capsule only (e.g., Riccia) or may
differentiated into foot, seta and capsule (e.g., Marchantia).
12. Archesporium is endothecial in origin.
13. Sporogenous tissue either forms only spores (e.g., Riccia) or is differentiated into
sterile elater mother cells and fertile spore mother cells.
14. Columella is absent in the capsule.
15. Elaters are unicellular, hygroscopic with spiral thickenings.
16. Capsule wall is one to several layers thick and without stomata.
17. Dehiscence of the capsule is irregular or in definite number of valves.
18. Spores on germination form the gametophytic plant body.
19. Plants show heteromorphic alternation of generation. Schuster (1957) divided the
class Hepaticae into two sub-classes:
Sub-class 1. Jungerinanniae. It includes four orders:
Order 1. Calobryales (e.g., Calobryum)
Order 2. Takakiales (e.g., Takakia)
Order 3. Jungermanniales (e.g., Pellia)
Order 4. Metzgeriales (e.g., Metzgeria)

Sub-class 2. Marchantiae: It includes three orders:

Order 5. Sphaerocarpales (e.g., Sphaerocarpos)
Order 6. Marchantiales (e.g., Marchantia).

All these 6 orders have their characteristic features and all these are described
here. The important features of these orders are:
1. Order- Calobryales (2 genera- Calobryum (8 spp.) and Haplomitrium (1 spp.):
The characteristic features are as follows:
1. They possess erect leafy gametophytes with leaves in three vertical rows.
2. The leaves are dorsiventrally flattened.
3. They have a pale, subterranean, sparingly branched rhizome from which arise erect
leafy branches.
4. Erect branches bearing sex organs have the uppermost leaves close together and in
more than three rows.
5. They are devoid of rhizoids.
6. The antheridia are ovoid, stalked, and borne at the apex of the stem.
7. The jacket of the neck of archegonium has only four vertical rows of cells.
8. The sporophyte bears an elongate capsule whose jacket layer is only one cell in
thickness except at the apex.
9. The number of chromosomes is-n=9.
Since there is single family Calobryaceae the characters are similar to that of the order.
Two genera - Calobryum and Haplomitrium.

Order-Takakiales (1 genus; 2 species):

The characteristic features are as follows:
1. They possess cylindrical, rhizomatic and erect gametophores.
2. They are devoid of rhizoids.
3. They possess copious beaked or non-beaked mucilage hairs on them.
4. They possess bifid-trifid-quadrifid leaves or phyllids.
5. The gametophores are about 1 to 1.5 cm. in height.
6. The leafless branches facing downward known as 'flagella' or 'stolons' may be
present.
7. Asexual reproduction is not known.
8. Only female (archegonial) shoots are known. They bear conspicuous pedestalled
archegonia.
9. The male (antheridial) shoots and the sporophytes are not known.
10. They have lowest chromosome number (i.e., n=4).
11. They are supposed to be most primitive and sometimes known as living fossil.
There is one family Takakiaceae, and one genus Takakia.

3. Order- Jungermanniales (220 genera; 8, 500 species):

The characteristic features of this order are as follows:
1. The gametophyte is differentiated into stem and leaves; the leaves are borne in a
regular spiral succession along the stem.
2. The apical cell is pyramid-like with three cutting faces.
3. The stem generally bears three rows of leaves; two rows are lateral and consist of
leaves of normal size; the third row consists of the under leaves which are generally
smaller than the lateral leaves.
4. The archegonia are always restricted to the apices of the axis and its branches.
5. The sporophytes are always terminal in position.
6. The antheridia are borne singly or in groups in the axis of leaves.
Family- Porellaceae (single genus- Porella):
1. The leaves are arranged in three rows on the stem; ventral leaves are well developed
and usually decurrent at the base.
2. The rhizoids are scarce and arise from the lower side of the stem in tufts generally
near the base of under leaves (ventral leaves).
3. The archegonia are borne in terminal cluster on small lateral branches; the archegonia
remain surrounded by a large inflated perianth.
4. The spherical capsule dehisces by four valves which split only to half way down.

Family- Frullaniaceae (three genera; important genus Frullania):

1. The thallus is pinnately branched and differentiated into stem and leaves.
2. The leaves are arranged in three rows two laterals unequally lobed and a ventral
lobule.
3. The ventral leaves are bifid and trumpet-shaped.
4. The archegonia develop in a group.

4. Order- Metzgeriales (23 genera; 550 species):

The characteristic features of this order are as follows:
1. The gametophyte may be thalloid or differentiated into stem and lateral leaves.
2. In most cases the gametophytes are without internal differentiation of tissues but
certain genera have a central strand of thick-walled cells.
3. The ventral surface of a gametophyte bears smooth-walled rhizoids.
4. The sex-organs are found to be scattered on dorsal surface of thallus.
5. The archegonia arise from the young segments cut off by the apical cell.
6. The mature sporophytes lie some distance back from the growing apex of a
gametophyte.
7. The sex organs (antheridia and archegonia) are produced on any branch of the
gametophyte or only on special branches.

Family-Pelliaceae (Three genera- Pellia, Noteroclada and Calycularia):

1. The thallus is prostrate, dorsiventral and very often lobed by irregular incisions.
2. The rhizoids are simple, non-septate, smooth and thin-walled. The scales are absent.
3. The sex organs (antheridia and archegonia) remain scattered on the dorsal surface of
the thallus.
4. The archegonial cluster always remains surrounded by an involucre which is an
outgrowth of the thallus.
5. The capsule (sporogonium) is globose or oval in shape. It possesses a basal
elaterophore.

Family- Riccardiaceae (two genera):

1. The gametophytes are wholly thallose or have thallose terminal branches.
2. The cells of thallus possess finely segmented oil bodies.
3. The sex organs (antheridia and archegonia) are borne on short lateral branches.
4. A well-developed calyptra is present but there is no involucre.
5. A distal elaterophore is present to which some elaters are attached.
6. The capsule is ovoid to cylindrical and dehisces longitudinally into four parts
extending to the base.

Family- Fossombroniaceae (4 genera; Fossombronia, Simodon, Petalophyllum and

Sewardiella):
1. Thallus is distinctly foliose.
2. The thallus is dorsiventral and prostrate.
3. The stem is branched. Growth of the main axis and of its branches is by means of an
apical cell with two cutting faces.
4. The leaf is thin, one cell-thick except the basal portion which is 2 or 3 cells in
thickness.
5. Antheridia develop in acropetal succession singly or in small groups.
6. Archegonia occur in small groups.
7. Young sporophyte remains surrounded by a calyptra and which is ensheathed by a
cuplike involucre.
8. The mature sporophyte is surrounded and protected by a bell-shaped sheathing
perianth.
9. Important genus-Fossombronia.
5. Order- Sphaerocarpales (3 genera)-two families:
1. Family- Sphaerocarpaceae - Sphaerocarpos (seven species) and Geothallus (single
species).
2. Family- Riellaceae- Riella (17 species).
The characteristic features of the order are as follows:
1. Vegetative structure of gametophyte is similar to that of order-Metzgeriales, but in
which development and structure of sex organs, as well as the structure of sporophyte
are similar to those of order-Marchantiales, and because of this the genera are placed in
separate order Sphaerocarpales.
2. The main diagnostic feature by which the order is recognized is the presence of
globose or a flask-like envelope or involucre around each of the sex organs (i.e.,
antheridia and archegonia).

6. Order-Marchantiales (32 genera; 400 species):

The characteristic features are as follows:
1. The ribbon-like, dichotomously branched and dorsiventral thalli grow prostrate upon
suitable substrata.
2. Excluding Dumortiera, Monoselenium and Monoclea, the rest of the genera possess
internally differentiated air chambers on the dorsal side of the thallus; such chamber
opens outside by an air pore of a particular design.
3. The ventral portion of the thallus consists of parenchyma which acts as storage tissue;
oil and mucilage cells may be present in this region.
4. The scales and rhizoids are present on the ventral side of the thallus; the rhizoids are
of two types (smooth-walled and tuberculate).
5. The antheridia and archegonia may be found directly on the dorsal surface of the
thallus or they may be present on the special branches known as antheridiophores and
archegoniophores respectively.
6. In most of cases the capsules of the sporophytes possess single layered jacket.
7. The capsule may be simple as in Riccia or it may be differentiated into foot, seta and
capsule as in Marchantia.
8. The elaters may or may not be present. According to Campbell (1940), there are five
families in this order. The characteristic features of two families are given here:

I. Family- Ricciaceae (3 genera; 140 species):

1. The gametophyte consists of a rosette-like dichotomously branched thallus.
2. In the thallus, the dorsal portion consists of chlorophyllous strips which may or may
not have air canals among them; the ventral portion of the thallus is parenchymatous
and acts as storage tissue.
3. The sex organs (antheridia and archegonia) are found in the longitudinal groove on
the dorsal side from the growing apex to backward in basipetal succession.
4. The sporogonium consists of a simple capsule which is not differentiated into foot,
seta and capsule.
5. Elaters not present.
6. The archesporium produces only the spores.
The important genera are- Ricciocarpus and Riccia.

II. Family- Marchantiaceae (23 genera; 250 species):

1. The thallus is dorsiventral; it has distinct assimilatory and storage regions.
2. The assimilatory region remains divided into several chambers and each chamber
contains branched assimilatory filaments.
3. The pores of the thallus may be simple or barrel-shaped.
4. The archegonia are borne upon special erect, stalked, vertical branches, the archego-
niophores.
5. The antheridiophores may or may not present; however, in Marchantia, the antheridia
are borne upon these erect, stalked antheridiophores.
6. The typical sterile elaters are found in the sporogonium mixed with the spores. The
important genera are - Conocephalum, Cryptometrium, Lunularia, Marchantia, etc.

Class II. Anthocerotopsida (Hornworts):

General Characters:
1. This class is represented by about 6 genera and 300 species.
2. Plant body is flat, dorsiventral, thalloid, gametophytic and variously lobed.
3. Smooth walled rhizoids are present.
4. Tuberculated rhizoids and scales are absent.
5. Internally the thallus is not differentiated into zones.
6. All cells are alike.
7. Air chambers or air pores are absent.
8. Each cell has a single chloroplast and each chloroplast contains a single pyrenoid.
9. Mucilage cavities open on the ventral surface by slime pores.
10. Sex organs are embedded in the thallus.
11. Antheridia develop either singly or in groups in closed cavities called antheridial
chambers.
12. The sporophyte is differentiated into foot, an intermediate zone or meristematic
zone and capsule.
13. Due to the presence of the meristematic zone, the sporophyte shows indeterminate
growth i.e., it continues to grow indefinitely.
14. Archesporium is amphithecial in origin.
15. Sporogenous tissue forms the fertile spores and sterile elaters. Elaters do not have
spiral thickenings and are known as pseudo elaters.
16. Capsule wall is four to six layered thick and epidermis has the stomata.
17. Capsule matures from apex to base and usually dehisce by two valves. The class
Anthocerotopsida has only a single order Anthocerotales.

Muller (1940), Proskauer and Reimers (1954) divided the order Anthocerotales in two
families:
Family 1. Anthocerotaceae (e.g., Anthoceros)
Family 2. Notothylaceae (e.g., Notothylas).
Family-Anthocerotaceae (4 or 5 genera; important genus Anthoceros):
1. The capsule is linear and vertical.
2. The stomata are present on the wall of capsule.
3. The archesporium develops from amphithecium.
4. The elaters are four-celled, smooth or thick-walled; thickening band may or may not
be present.

Family-Notothylaceae (single genus- Notothylas):

1. The capsule is cylindrical and horizontal.
2. The stomata are not found on the wall of capsule.
3. Archesporium arises from endothecium and amphithecium.
4. Elaters are short and stumpy; they have irregular thickening bands.

Class III. Bryopsida (Mosses):

General Characters:
1. It is the largest class in Bryophyta and includes about 700 genera and 14,000 species.
2. The main plant body is gametophytic and can be differentiated into two stages-
juvenile stage and leafy stage or gametophore.
3. Juvenile stage is represented by green, filamentous branched structures called
protonema. It develops from the germination of the spore.
4. Gametophores are erect leafy branches which develop on the protonema.
5. Gametophores can be branched or un-branched and can be differentiated into three
parts-rhizoids, ‘stem’ and ‘leaves’.
6. Branches arise below the ‘leaves’.
7. ‘Leaves’ are with midrib, un-lobed and arranged spirally in three to eight rows on the
axis or
8. Rhizoids are multicellular, filamentous, branched with oblique septa.
9. The axis is differentiated into central conducting strand enclosed by cortex.
10. The sex organs (antheridia and archegonia) develop from the superficial cells of the
gametophore.
11. The sporophyte is green in early stages and can be differentiated into foot, seta and
capsule.
12. The seta is usually elongated and rigid.
13. The capsular wall remains interrupted by stomata at several places.
14. Columella is usually present and endothecial in origin.
15. Archesporium (spore forming tissue) is differentiated only in spores.
16. Elaters are absent.
17. Dehiscence of capsule takes place by separation of lid or operculum.
18. Peristome helps in the dispersal of spores.
19. Spores on germination produce the protonema.
The class Bryopsida (Musci) has been divided into three sub-classes
(1) Sphagnobrya (Sphagnidae); (2) Andreaeobrya (Andreaeidae) and (3) Eubrya
(Bryidae).

I. Sub-class: Sphagnobrya:
The sub-class has a single order, the Sphagnales and a single family, the Sphagnaceae.
(Single genus Sphagnum with 326 species). The characteristic features are as follows:
1. They are called 'bog mosses' or 'peat mosses'.
2. The protonema is broad and thallose; it produces one gametophore; the leaves or
gametophores lack mid-rib and usually composed of two types of cells-(i) the narrow
living green cells and (ii) large hyaline dead cells.
3. The branches arise in lateral clusters in the axis of the leaves borne on the stem.
4. The antheridia are borne in the axis of leaves on the antheridial branch.
5. The archegonia are terminal and formed acrogenously.
6. The sporogenous tissue of a sporophyte develops from the amphithecium.
7. The sporogonium remains elevated above the gametophyte due to elongation of a
stalk of gametophytic tissue, the pseudopodium.

II. Sub-class. Andreaeobrya:

This sub-class has a single order, the Andreaeales, and a singly family, the
Andreaceceae. The important genus is Andreaea. The characteristic features are as
follows:
2. There is practically no tissue differentiation in plant body.
3. The leaves are generally large, erect and convolute.
4. The archesporium and colonmella develop from the endothecium.

III. Sub-class Eubrya (650 genera; 14,000 species):

This sub-class has been further divided into three cohorts and fifteen orders. The true
mosses are included in this sub-class. The characteristic features are as follows:
1. The leaves of the gametophores are more than one cell in thickness and possess
midrib on them.
2. The protonema is filamentous.
3. The sporophyte bears a well differentiated, elongated seta which pushes out the
capsule from the gametophore.
4. The sporogenous tissue is derived from the endothecium.
5. The archesporium does not overarch the columella; the columella continues upto the
apex of the capsule; both columella and archesporium have been derived from the
endothecium.
6. In between spore sac and columella, the partitioned air spaces are present.
7. The mature capsule possesses the complex structure made of many tissues.
8. The capsule opens at its apex by an operculum; the spore dispersal is regulated by
teeth like apparatus, the peristome.
Order-Funariales (26 genera; 356 species):
Characteristic features:
1. The plants are terrestrial; they are small in size and may be annual or biennial.
2. The leaves possess distinct mid-ribs and arranged in rosettes at the apex of the
gametophyte.
3. The capsule is wide and provided with an unbeaked operculum.
4. The peristome of the capsule is double and consists of inner and outer peristome
called endostome and exostome respectively.
5. There are five families in this order, of which Funariaceae is most important.

Family-Funariaceae (9 genera; 200 species):

1. The leaves are one cell in thickness except at the mid-rib region.
2. The small mosses form the velvety appearance on the surface of the sustratum.
3. The calyptra is soon detached from the opercula of the capsules; the calyptra is
provided with long beaks.
4. The capsules are pyriform and situated on the long, elongated setae.

Order-Polytrichales:
Characteristic features:
1. The gametophyte is perennial and tall.
2. The leaves are narrow and possess longitudinal lamellae on the upper surface of the
midrib.
3. The capsule is terminal.
4. The single annular series of cells gives rise to a peristome in the inner zone of the
amphithecium.
5. There are 32 to 64 pyramidal teeth in peristome; the tips of the peristome teeth
remain joined above to a thin membrance, the epiphragm covers the mouth of the
capsule.
6. There is a single family, the Polytrichaceae in this order; the important genera of this
family are — Polytrichum and Pogonatum.

Apogamy and Apospory in Bryophytes

Bryophytes are endowed with a remarkable regeneration capacity. Parts of the plant or
any living cell of the thallus are capable of regenerating the entire plant. The
sporophytic cells regenerate to form a protonema on which appear gametophytes. This
regeneration of diploid gametophyte from a sporophyte without the formation of spores
is called apospory.
Conversely a gametophyte may form a mass of cells which may regenerate a
sporophyte. This regeneration of a diploid sporophyte from a gametophyte, without the
formation of gametes is called apogamy. Aposory and apogamy are rarely found in life
cycle of Bryophytes.