PREFACE

Thank God we pray that the presence of GOD Almighty has bestowed grace
and blessing me so that we can complete the task of this paper without a hitch.
And do not forget we would like to thank Mr / Ms lecture who have guided us, so
that we can complete the task of this paper. We realize in the preparation of this
paper is far from perfect , for that we

expect criticism and constructive

suggestions for the perfection of this paper.

One thing we need to know that we are studying science is dynamic. Both
the natural sciences and social sciences. In this paper we will discuss about the
Taxonomy of Lower Plant , especially in discuss about byriophyta ant its
taxonomy .
We hope that the topic of this paper can provide many benefits for us.
There are explanation about the general charateristic , the morphology and life
cycle.And dont forget also pray to god to give something back to those who
support the preparation of this paper.

Sincerely

Authors

TABLE OF CONTENT

PREFACE ..............................................................................................................1
TABLE OF CONTENTS.......................................................................................2
CHAPTER 1 : INTRODUCTION .......................................................................3
CHAPTER 2 : DISCUSSION ..............................................................................4
2.1

DIVISION BRYOPHYTA ...........................................................5

2.1.1. Class Musci.........................................................................10
2.1.1.1. Subclass Andreaeidae.........................................10

2.2

DIVISION ANTHOROCEROTOPSIDA .................................14

2.2.1. Class Leiosporocerotopsida...............................................15
2.2.2. Class Athocerotopsida.......................................................15

2.3

DIVISION HEPATICOPHYTA ................................................19

2.3.1. Class Marchantiopsida......................................................19
2.3.1.1 Sub class Marchantiidae......................................23
2.3.1.2 Sub class Sphaeocarpidae....................................26
2.3.2 Class Jungermanniopsida...................................................27
2.3.2.1 Subclass Metzgeriidae..........................................28
2.3.2.2 Subclass Jungermanniidae..................................31

CHAPTER 3 : CONCLUSSION ........................................................................34

REFERENCES ....................................................................................................35

CHAPTER I
INTRODUCTION
Bryophyta is the Plant Division that contains mosses. Plants in this
Division have crude stems and leaves, but no roots. Instead of roots, they have
"rhizoids." Rhizoids help anchor the plant to a surface, but they do not absorb
nutrients like roots on other plants do. Instead of using flowers to make seeds,
mosses release spores from their leaves. Spores can travel by water and make new
mosses in new locations. Water is very important to mosses, so they can grow and
spread; however, mosses can survive even when they dry out. When they become
wet again, they revive and continue growing. (S.M. Reddy : 1996 )
After flowering plants and ferns, mosses are the most diverse group of
plants, with more than 10,000 species in 700 genera. This makes mosses almost
twice as diverse as mammals. Mosses don't receive as much attention from us as
flowering plants, ferns, or conifers because most mosses are small and
inconspicuous.
They have no vascular tissue or wood to lend them structural support, nor
do they have large leaves or showy cones or flowers. This does not mean that
mosses are not important; in fact, mosses play important roles in reducing erosion
along streams, water and nutrient cycling in tropical forests, and insulating the
arctic permafrost. (www.ucmp)

CHAPTER II
DISCUSSION
TAXONOMY OF BRYOPHYTA

2.1. DIVISION BRYOPHYTA (MOSSES)

Although the bryophyte is used as a collective term for all of these

-Bryophyta(mosses),

Hepatophyta

(hornworts). Characteristics

(liverworts),

of Mosses

and

, Mosses

Anthoceratophyta

are mostly-terrestrial

bryophytes.Mosses are found in a range of habitats, although moist and shady

habitats are more common. Mosses are often epiphytes. The dominant phase of
the moss life cycle is the gametophyte (haploid). The plant is called a thallus, they
may be erect or prostrate (axis along the ground). Mosses have radial symmetry,
in that a cut down the long axis of an individual gives two similar halves. The
gametophyte has a stem like axis with spirally arranged leaves, which are known
as phyllids . Mosses attach to their substrate with multicellular rhizoids .Moss
leaves are variable in shape.Leaves usually consist of a single cell layer and are
traversed by a midrib that is always more than one cell in thickness. It lacks xylem
and phloem.The plant body may have conducting tissue. The xylem-like
conducting tissue is called hydroid. The phloem-like conducting tissue is called
leptoid. All mosses have a sporic (diplohaplontic) life cycle that is oogamous.
Table 2.1.1. Distinguishing Characters of Division Bryophyta , Division
( Hepatophytaand Division Anthocerotophyta)

Character
Protonema
Gametophyte
form
Leaf
arrangement
Leaf form
Special
organelles
Water

Bryophyta
Filamentous,

Marchantiophyta

Anthocerotophyta

Globose, forming one Globose, forming one

forming many buds bud

bud

Leafy shoot or thallus;

Leafy shoot

thallus simple or with Simple thallus

air chambers

Leaves in spirals

Leaves in three rows Not Applicable

Leaves undivided, Leaves divided into

Not Applicable.

midvein present.

2+ lobes, no midvein

None

Oil bodies

Present in both

Present only in a few Absent.

Single plastids with

pyrenoids.

conducting

gametophytes and

cells

sporophytes

Rhizoids
Gametangial
position

Brown,
multicellular

simple thalloid forms

Hyaline, one-celled.
Apical clusters (leafy

Apical clusters

forms) or on upper
surface of thallus

Stomates

Present on

sporophyte capsule. generations

Photosynthetic,
emergent from

Seta

gametophyte early
in development
Complex with

Capsule

operculum, theca
and neck; of fixed
size

Sterile cells in
capsule

Absent in both

Columella.

Capsule

At operculum and

dehiscence

peristome teeth

Hyaline, one-celled
Sunken in thallus,
scattered
Present in both
sporophyte and
gametophyte.

Hyaline, elongating
just prior to spore

Absent.

release
Undifferentiated,
spherical or elongate;
of fixed size

Undifferentiated, hornshaped; growing

continuously from a
basal meristem.

Spirally thickened

Columella and

elaters

pseudoelaters.

Into 4 valves

Into 2 valves.

a. Gametophyte Characteristics
Moss life cycle begins when haploid spores are released from a sporophyte
capsule and begin to germinate. In the majority of mosses, germination is
exosporic, i.e., the spore wall is ruptured by the expanding spore protoplast after
its release from the capsule and prior to any cell division. However, in some
mosses, e.g. Andreaea, Drummondia, and Leucodon, germination is precocious
and endosporic, meaning that cell divisions occur prior to spore release and spore
wall rupture, respectively. There are variations in patterns of germination of moss

( K. Nehira 1983). In most mosses, a highly branched filamentous, uniseriate

protonema are formed.
Cell specialization occurs within the protonema as a result two types of filaments
are formed:

a horizontal system of reddish brown,anchoring filaments (rhizoids),

called the caulonema

upright, green filaments, the chloronema.

Each protonema can spread over several centimeters, forming a fuzzy green
film over its substrate. Usually this protonemal stage is short-lived, but in a few
taxa, e.g., Buxbaumia it persists as the vegetative phase of the plant.
Formation of bud apical cells: As the protonema grows, target cells
usually on the caulonema generate bud initials that will ultimately divide by
sequential oblique divisions to form bud apical cells. This initiates the growth of
the leafy gametophore or shoot stage of the moss.
b. Shoot Morphology and Habit

The leafy shoot continues to grow by mitotic division of its obovoidal to

fusiform apical cell and surrounding meristem.

Divisions occurring in the apical cell form spirally arranged derivatives,

each of which will give rise to a single leaf and a portion of the stem.

The angle of divergence between successive derivatives is responsible for

the spatial arrangement of the leaves or phyllotaxy of the shoot.

Mature leaves of few mosses are clearly ranked; e.g., the leaves of
Fissidens and Bryoxiphium are in two rows, a 1/2 phyllotaxy

Fontinalis and Tetraphis have leaves aligned in three rows, a 1/3

phyllotaxy.

In most mosses, however, the leaves are spirally distributed, with 2/5 and
3/8 phyllotaxies being most common.

The peristomate or true mosses (Superclass V) on the basis of position of

the perichaetia and subsequent sporophytes have traditionally been divided
into two broad morphological groups:1. Acrocarps :-Acrocarps are characterized by erect or ascending
shoot systems that are either unbranched or only sparingly
branched. Branching is typically sympodial with the branches
morphologically comparable to the determinant main shoot from
which they arise. Perichaetia are differentiated at the tip of the
main or primary shoot and terminate its growth, so further plant
growth occurs only if a branch is produced below the perichaetium;
such branches are called subfloral innovations
2.

Pleurocarps are generally characterized by creeping shoot systems,

with extensive lateral branching.In such systems, the indeterminant
main stem may be morphologically distinct from the secondary and
tertiary level branches that arise from it (C. La Farge 1996).
Perichaetia in pleurocarps are produced at the tips of very short,
basally

swollen

lateral

branches

that

are

very

short,

morphologically distinct from the vegetative branches.

Cladocarpic mosses produce perichaetia at the tips of unspecialized lateral

branches that display the same heteroblastic leaf series as the vegetative
branches. Such branches are themselves capable of branching, and these
mosses are neither acrocarpic nor pleurocarpic.

Pleurocarps form a natural, monophyletic lineage of true mosses (B.

Goffinet and W. R. Buck 2004), but cladocarpy has evolved in several
different lineages.

The main stems of Sphagnum (Superclass II) display a furcate or

dichotomous branch architecture (H. A. Crum 1984).

c. Rhizoids
Mosses are anchored to their substrates by filamentous, often
branched, reddish brown rhizoids, except Takakia and Sphagnum. The

rhizoids (As in caulonemata) are multicellular with oblique cross walls;

their walls are smooth or roughened with papillae.

Most rhizoids are slender and only sparingly branched (micronematal

type) arise from any of the epidermal cells of the stem.

But others are larger in diameter and extensively branched (macronematal

type) and is associated only with branch primordia.

They function primarily as anchoring structures . Rhizoids are not major sites
of water and nutrient uptake, but can enhance capillary movement of water along
the outer surface of the stem (M. C. F.Proctor 1984).
d. Stem Anatomy
In many mosses, the stem is anatomically complex, consisting of a
differentiated epidermal layer, a cortex, and a central strand of thin-walled,
hydrolyzed water conducting cells, called hydroids.
e. Leaves
Considerable variation in the arrangement and structure of moss
leaves provides some of the most morphologically useful characters for
species identification.<ul

Leaves typically arise from all sides of the stem, most commonly
exhibiting a spiral phyllotaxy, but distichous and tristichous arrangements
can also be found.

Isophyllous:-The mature leaves of a given shoot are usually all similar in

size and shape.

Anisophyllous:- but there are taxa that are anisophyllous, with either
dorsal or ventral leaves decidedly smaller than the lateral leaves.

Except for a few taxa like Fissidens, leaves are attached to the stem along
broad transverse lines.

2.1.1. Class Musci

General Characteristic
The gametophyte consists of prostrate, thalloid, branched protonema and
erect leafy gametophore.The gametophytic plant body consists of the stem,
spirally arranged leaves and the sex organs (antheridia and archegonia) at its
apical portion. The rhizoids are multicellular, branched and obliquely septate. The
sex organs (antheridia and archegonia) develop from the superficial cells of the
gametophore. The sporophyte is differentiated into foot, seta and capsule. The
capsular wall remains interrupted by stomata at several places.The archesporium
or sporogenous mass develops from outer layer of endothecium which in addition
forms columella.The elaters are not present in the sporogonium.
The class Bryopsida (Musci) has been divided into seven sub-classes,
namely : 1. Andreaeidae, 2. Sphagnidae, 3. Tetraphidae, 4. Polytrichidae, 5.
Buxbaumiidae, 6. Bryidae, and 7. Archidiidae.
2.1.1.1. Subclass Andreaeidae
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:

The gametophores are brittle, and can easily be broken. There is

practically no tissue differentiation in plant body. The leaves are generally large,
erect and convolute. The archesporium and colonmella develop from the
endothecium.
a. Order Sphagnidae
The subclass-sphagnidae

includes

the mosses which are often very

importantconstituent

of

peat

bog

vegetation. Hence the name of peat or

bog mosses. The grow of extensive
masses on boggy and peaty soils and
Sphagnidae species

also as submerged aquatics in peaty

pools. The subclass spaghnidae is characterised by the following distinctive

10

feature : The simple, flat, plate like thallose protonema is fixed to the substratum
by numerous rhizoids.
The rhizoids are multicellular. The septa are between the cells and oblique
,The upright, leafy branch originates from a single protonema cell , Usually a
single gametophore develops from one protonema , The leaf of adult bog moss
has a unique structure. It consists two kinds of cells, the narrow, living, green
assimilatory cells and the large, colorless, dead capillary cells , The leaf have ni
midrib , The antheridia occur singly and are axillary in position ,They develop in
special side branches , The archegonia is terminal in position , The young capsule
is invested by the calyptra , The peristome is absent.
b. Order Tetraphidae
The Subclass Tetraphidae is a member of the Class Polytrichopsida. Here is
the complete "parentage" of Tetraphidae: The characteristic of tetraphidae ,
Sporophytes with elongate seta ,Sporangium opening by an operculum exposing
four multicellular peristome teeth that respond to moisture change to release
spores. The Tetraphidaceae is a family of mosses. It includes only the two genera
Tetraphis and Tetrodontium, each with two species. At least 13 species and
subspecies belong to the Family Tetraphidaceae.
c. Order Polytrichidae
The Polytrichaceae is a common
family of mosses. Members of this
family tend to be larger than other
mosses with a thickened central stem
and a rhizome. The leaves have a midrib
that bears lamellae on the upper surface.
Pogonatum brachyphyllum

Species in this group are dioicous.

Another characteristic that identifies them is that they have from 32 to 64

peristome teeth in their sporangium. At least 627 species and subspecies belong to
the Family Polytrichaceae. The Characteristic features , The gametophyte is
perennial and tall. The leaves are narrow and possess longitudinal lamellae on the
upper surface of the midrib , The capsule is terminal. The single annular series of
11

cells gives rise to a peristome in the inner zone of the amphithecium , 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.
d. Order Buxbaumiidae
The species within Buxbaumiidae
are typically found in Northern temperate
to subtropical climates of the world, and
are
Buxbaumiidae Species Buxbaumia piperi

restricted

to

acidic

or

neutral

substrates. Substrates include rotten wood,

rock, and soil. Species are uncommonly

found in the tropics and southern hemisphere.

There is a lot of morphological diversity within Buxbaumiidae.

An

evident pattern is seen between the size of the gametophytic and sporophytic
generations. Typically, the sporophytic generation can be seen macroscopically
while the gametophytic generation is the smallest in comparison to other
bryophytes. The leaves of the gametophyte are acostate and spirally arranged
around the short stem. The reduced male and female shoots are borne from the
same protonemal shoots. One antheridium is surrounded by a single unicellular
flap of tissue. Female shoots are composed of 1-5 archegonia that are surrounded
by 6-10 non-chlorophyllose leaves.

The uniseriate and branched protonemal

phase gives rise to these shoots.

The sporophytic generation is unique to Buxbaumiidae. The sporangium
typically has an oblique-horizontal and asymmetric surface, with a cucullate or
mitrate calyptra. The neck and seta are short, and there is an endostome and
exostome. Typically, the articulated endostome is arranged into a pleated coneshape, while the exostome is irregular, rudimentary, or absent. The peristome aids
in identifying species. For instance, in Buxbaumia the endostome is colorless,
truncate, and cone-shaped with 32 pleats. The exostome is composed of
alternating grooves and pleats and can form up to 4 rows of concentric teeth. The
uniqueness of Buxbaumiidae is truly a spectacular sight to see.
e. Order Bryidae
12

The Subclass Bryidae is characterized by sporophytic features. A primary

distinguishing trait is the arrangement of the peristome. Species have a double
peristome with alternating teeth and segments, and the endostome is ciliate.
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 ,The leaves of the gametophores are more than one cell in thickness
and possess midtrib on them.The protonema are filamentous. The sporophyte
bears a well differentiated, elongated seta which pushes out the capsule from the
gametophore ,
The sporogenous tissue is derived
from the endothecium , 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 ,
In between spore sac and columella, the
Bryidae species

partitioned air spaces are present , The

mature capsule possesses the complex structure made of many tissues , The
capsule opens at its apex by an operculum; the spore dispersal is regulated by a
teeth like apparatus, the persistome. A few members of Subclass Bryidae include:
Aulacomnium androgynum , Bartramia pomiformis , Bryum argenteum , Buckiella
undulata , Claopodium crispifolium , Fontinalis antipyretica
f. Order Archidiidae
A subclass of the plant class Bryopsida; consists of a single genus, Archidium,
unique in having spores scattered in a single layer of the endothecium and having
no quadrant stage in the early ontogeny of the capsule.
This sub-class have sporophyte with no seta; sporangia containing a
restricted number of large spores (sometimes 4), lacking columella, opening by
decomposition of the jacket.

General Life Cycle of Bryophyta

13

2.2 DIVISION ANTHOCEROTOPHYTA

Characteristic of this division are nonvascular plants , The common name
refers to the elongated horn-like
structure , which is the sporophyte ,
the flattened, green plant body of a
hornwort is the gametophyte plant ,
growth only in places that are damp or

. Anthocerotophya species

humid. gametophytes are small, usually only one to two centimeters across, with
no significant height , the gametophyte is made up of a thallus, a simple plant
body resembling a mound of ruffled,
Morphology of this division are dark-green leaflets , the long slender
sporophyte generations grow upward from the thallus and are typically 0.5 to 12
centimeters in height. Reproduuksinya sexually ie by way of the formation of
gametes.
Morphology the thallus structure of hornworts is simple, they have an
epidermis and air chambers. In general, the gametophyte has a rather greasy
appearance. typically only one chloroplast per cell in the thallus.
Anthocerotophyta divided into two class:

2.2.1. Class Leiosporocerotopsida

14

General characteristic:
Sporophytes very large with characteristic grooves on either side of the
developing capsule, sporogenous region massive, spores scattered among
pseudoelaters, without distinctive columelle large stomata.
a.

Order Leiosporocerotales
General characteristic , Gametophytes are thalloid and solid (no internal

openings) with no mucilage clefts, sporophytes are very large with characteristic
grooves on either side of the developing capsule, the sporogenous region of the
sporophyte is massive with isobilateral spores (not tetrads) scattered among the
elaters (pseudoelaters), which are relatively large
and may be made of more than one cell, the
columella is not distinct, the capsule has large
stomata.

Morphology

of

Leiosporocerotales

Nostoc parallel the main axis of the thallus

Family Leiosporocerotaceae General characteristic usually small spores
that are monolete and unornamented, additionally, there are unique strands of
Nostoc (cyanobacteria) that grow inside the plant parallel with its direction of
growth.
2.2.2. Class Athocerotopsida
General characteristic Sheet-shaped
gametophyte, sporophytes tubular extends
upward , like horns, in the " horn "produced
spores, anatomy of the talus homogeneous
structure , each cell consist the chloroplasts
with a large pirenoid, sporogonium consists
of legs and capsules alone , Spores germinate not form Prototype , asexual
propagation is equal to liverworts.Morphology of athocerotopsida
Has seta in which the capsule is a cylinder with a length of several
centimeters, On the outer capsule contained epidermal cells ( capsule wall ) , and
generally there are stomata . Sporophyte not stemmed and has the shape of a horn.
15

Athocerotopsida divided into two orders:

a. order anthocerotales
General characteristic:
The gametophytes are typically thalloid and
spongy (cavernous) with mucilage canals
and lamellae on the upper part, stomates
occur on the gametophyte thallus and on the
sporophyte capsule, the antheridia, which open apically, occur in large groups and
have thick walls with four tiers, spores are dark (brown to black), which is a
source of fixed nitrogen.

Highly ornamented and darkly pigmented spores; schizogenous mucilage

canals in the dorsal half of the thallus, the occurrence of large dorsal lamellae on
the thallus surface, and the development of tiered antheridia that occur in large
groups and open apically.

Family anthocerotaceae
General characteristic

16

The sporophyte is long,silindrical structure which aries from the dorsal

surface of the thallus.at the base,it is surrounded by a tubular sheath called
involucres, the capsule wall is several layered thick, the stomata are present on the
epidermis, the cental columella is endothecial in origin
.Morphology of anthocerotaceae

The dorsal surface may be velvety do to the presence of flat, The ventral
surface of the thalllus bears numerous smooth walled rhizoids, have sporophytes
in the middle of the capsule color green.
b. Order notothyladales
General characteristic of notothyladales
Highly variable and diverse, these taxa form a clade (or two clades) without
distinct structural synapomorphies their spores are light and do not have tiered
antheridia.

Family notothyladaceae
General characteristic
17

Each proximal spore surface with a small central hollow, irregularly

arranged, rectangular to quadrate epidermal cells of the capsule, ebsence of a
special dehiscence line in the capsule.

The presence of a single chloroplast with a pyrenoid per cell and the
number of antheridia per chamber.

Family Dendrocerotaceae
General characteristic
The gametophyte is yellowish-green and usually less than one-half cm

wide, the thallus branches in a bifurcating pattern. In the subgenus Apoceros, there
are cavities in the central strand of the thallus, the edges of the thallus are only a
single layer of cells thick and have an undulating margin. It is common to find
symbiotic colonies of blue-green bacteria (usually Nostoc) growing among the
cells. Under a microscope, the epidermal cells have trigones, the sporophyte is
erect when mature, growing up to 5 cm tall,like other hornworts, its surface has
stomata,the interior of the sporophyte differentiates into a central column and a
surrounding mass of spores and elater cells, with a distinct spiral,the spores are
both green and relatively large with an ornamented surface.

18

Morphology : growing on the bark of a tree, grows on humid ground,

rocky outcrops, and on the sides of trees. Its name literally means "tree horn". its
surface has stomata.
2.3 DIVISION HEPATICOPHYTA
2.3.1. Class Marchantiopsida
The species in the class Marchantiopsida are characterized as being
complex thalloid liverworts. They have a global distribution and are commonly
found growing on moist earth or mud. A few species can also be observed
growing in water.
The gametophore of the species in this class is always thallose. The
thallus, which is generally several cell thick, typically exhibits dichotomous
branching.

The differentiated tissues of the thallus is why species in this class are
termed the complex thalloid liverworts, and in turn, it distinguishes them from
the simple thalloid liverworts seen in the class Jungermanniopsida.
The internal differentiation of the thallus can be made apparent due to the
presence of several distinct tissues. The upper surface (dorsal) of the thallus is
composed of an epidermis that is unistratose. The cells forming the epidermis
contain no or very little chlorophyll. Furthermore, the epidermis of most genera in
19

this class has pores, which can either be simple or complex, that serve as an
opening to the photosynthetic tissue in the air chambers beneath. The outline and
distribution of the air chambers can be seen when looking at the reticulated pattern
on the dorsal side of the thallus.

Unlike stomata seen in higher plants, simple pores of these thalloid

liverworts cannot open and close. However, some species have complex pores, in
which the pores are surrounded by cells, giving the pore a barrel-like appearance.
The cells at the bottom of the barrel-like pore can remain turgid when in a moist
environment, thereby keeping the pore open. When dry, these cells can collapse,
and thus can greatly narrow the opening of the pore.

As previously mentioned, these air chambers house the photosynthetic

tissue. The chloroplasts are typically located in the cells that make up the walls of
these chambers or in uniseriate filaments found growing on the bottom of the air
chamber.Below the chlorophyllous layer is where one can see a layer of storage
cells, predominantly parenchymous cells. Some cells within this layer will contain
a single large brownish complex oil body, while other cells may be filled with
mucilage. Not only is the parenchymous tissue used for storage, but it can also be
used to house fungi in order to benefit from a symbiotic relationship.
A ventral epidermis is present on the underside of the thallus, from which
can emerge colorless rhizoids and scales.
20

The rhizoids present can either pegged or smooth. Pegged rhizoids are
thought to be involved in water transport, whereas the smooth rhizoids aid in
fixing the thallus to the substrate and are often considered a gateway for
endophytic fungi. The ventral unistratose scales are typically arranged in one or
two rows and are thought to function in water retention and conduction.
The gametophyte of the species in this class can either be monoicous or
dioicous. The antheridia can be found in chambers that opens to the upper surface
of the thallus.

In some genera, the antheridia are present on a specialized structure that

extends from the gametophyte called the antheridiophore, as seen in this
picture.The antheridia are located at the top of the disk and the sperm is dispersed
via rain drops.
The archegonia can be found embedded in the thallus, as in Ricciocarpos,
or elevated on umbrella-like called archegoniophores, as in Marchantia. The
location of the archegonia differs from that of the antheridia, such that they are
typically found on the margin of the archegoniophore. Furthermore, as the

21

archegonia mature, the upper portion of the archegoniophore grows more than the
lower portion and as a result, the archegonia are transferred to the lower surface

(The archegonia in the following picture are labeled).

However, it is important to keep in mind that the morphology of the male
and female reproductive structures can vary considerably in this class. For
example, the antheridiophore of some aquatic species is highly reduced. Instead of
being elevated, the antheridia are embedded in the thallus, and the sperm are
dispersed by water movement. However, the locomotive and biflagellate sperm is
a distinctive trait that all species in this class share.

Asexual reproduction can be achieved by various ways depending on the

species. In Marchantia, gemmae develop inside a cup-like structure on the surface
of the thallus. Bifurcation, which is the splitting of the main thallus in two, is
another means of asexual reproduction. This process is followed by the decay of
the older posterior regions of the thallus, thus allowing the species to colonize
nearby areas rapidly.

2.3.1.1 Sub class Marchantiidae

22

The plants of this subclass are usually thalloid and rarely leafy. The
thallus, which typically differentiated on the dorsal and ventral surface, will
usually have air chambers and pores present. Scales on the ventral surface of the
gametophyte can generally be observed in several species.
Artheridia are present in chambers in various places depending on the
species. Some perigonial chambers can found scattered on the dorsal surface of
the thallus, while others can be clustered together on the main thallus or on a
specialized raised receptacle. The location of the archegonia can also vary
depending on the species. While some are found on the dorsal surface of the
thallus, others can be raised on a stalked receptacle termed the archegoniophore.
Even when the sporophyte reaches maturity the seta is typically short, and
maybe lacking in some species. The sporangial wall is typically unistratose and
the sporophyte generally dehisces by longitudinal valves or slits.
The Marchantiidae are thalloid plants, usually in the form of
dichotomously branching small rosettes or large flattened ribbons. The thalli
attach themselves to the soil by means of rhizoids. The male and female
gametangia are embedded in the tissue of the thallus or are raised on special
podetia. The plants grow predominantly in soil; a few species grow on rocks on
rocks or in water. Distributed throughout the world, they are especially
widespread in the tropics. The subclass has 16 known families, embracing 35
genera and about 420 species, including the fossil Naiadita from the Mesozoic.
Sub-classes Marchantiidae are divided into three orders there are : Marchantiales,
Monociales and Ricciales
a. Order Machantiidae
23

(Conocephalum conicum - a thallose liverwort)

Marchantiales is an order of thallose liverworts that includes species like
Marchantia polymorpha, a widespread plant often found beside rivers, and
Lunularia cruciata, a common and often troublesome weed in moist, temperate
gardens and greenhouses.
As in other bryophytes, the gametophyte generation is dominant, with the
sporophyte existing as a short-lived part of the life cycle, dependent upon the
gametophyte.
The genus Marchantia is often used to typify the order, although there are
also many s

pecies of Asterella and species of the genus Riccia are more

numerous. The majority of genera are characterized by the presence of (a) special
stalked vertical branches called archegoniophores or carocephala, and (b) sterile
cells celled elaters inside the sporangium.
b. Order Monocleales

(Monoclea fosteri)

24

Monocleales is a monotypic order represented by a few species of its only

genus Monoclea. Plant body is thalloid and remarkably large, reaching up to 20cm
or more in length and 5cm or more in breadth, and that is why these are
commonly called giant-thallose-liveworst. Monocleales possessing several
characters like ; lack of air chambers in the gametophyte, lack of ventral scales,
long seta, elongate elates, elaborate calyptra, and an archegonial pouch.
Monocleales are a thalloid type and is the largest species of livewort in
New Zealand. Its often found in the bush near waterfalls or seeps and has
extremely large lobes. The thallus does not have surface pores or gemma cups and
there is only one species found in New Zealand, Monocleales fosteri.
c. Order Ricciales

Order Ricciales is aquatic habitat and asymmetric plant body. The thallus
segments are 2 to 3 times dichotomous branched. In diameter they are 4 to 6 times
as wide as high. The plant grow vertical, in water and have been found asssociated
with a number of algae and aquatic angiosperms.the rhizoids are simple occuring
at the base of the midrib when plants become prostrate. The plants also have
several minute lanccolate ventral scales present on ventral surface of the midrib.
Anatomy of the thallus is mre or less similar to that of sphaerocarpus.
Reproduction by vegetative and sexual methods. In vegetative propagation take
place either by one celled or by many celled spherical gemmae produced at the
older part of midrib. Sometimes gemmae are also found in between ventral and
lateral scales. In sexual reproduction some species are both monoecious and
diocious. Some species are monoecious (homothalic) both antheridia and
archegonia are produced on separate independent plants.

25

2.3.1.2 Sub class Sphaeocarpidae

The thalloid plant body is bilaterally symmetrical. The family has two
genera Sphaerocarpus and Geothallus. Habitat prefer very moist and colder place
for their luxuriant growth. The morphological characters of the thallus vary if
habitat is changed. The gametophyte plant body is thalloid. The thallus is
prostrate, dorsiventrally flattened, dichotomously branched and dark green in
colour. Most of the species have notched margin and each notch bears a growing
point.
a. Order Sphaerocarpales
Plants leafy, stems bearing 2 rows of longitudinally inserted lateral leaves
(Sphaerocarpaceae) or small lateral and ventral scale-like leaves and one (rarely
two) large dorsal wing (Riellaceae),leaves and dorsal wing, unistratose, rhizoids
smooth, idioblastic oil cells absent or present (Riella), perigonial chambers
scattered dorsally on the stem or near the margin of the dorsal Wing, each
archegonium and sporophyte enclosed only by a pseudoperianth, involucres
absent, seta very short; capsules cleistocarpous, elaters absent, spores shed singly
or in tetrads.
Plants terrestrial, dioicous, strongly dimorphic, with the male plants
minute, often reddish, and the female plants much larger and green (only slightly
dimorphic in Geothallus), ventral slime papillae present at plant apices; idioblastic
oil cells absent; oil bodies absent, perigonial chambers emergent, pyriform to
bottle-shaped, pseudoperianths cylindrical to flask-shaped, covering the dorsal
surface of the stem, spores shed singly or in tetrads, gemmae absent.
Plants aquatic, submerged to emergent, dioicous, rarely monoicous, when
dioicous with male and female plants of similar size, ventral scales present, in 2
rows, without appendages, idioblastic oil cells present, perigonial chambers
embedded in the dorsal wing, opening along the margin, pseudoperianths
pyriform to flask-shaped.

26

2.3.2 Class Jungermanniopsida (Leafy Liverworts)

Class Jungermanniopsida includes leafy and simple thalloid liverworts.
There is a lack of shared characteristics between these two groups.The leaves of
leafy liverworts may be entire or lobed with various forms of insertion. The
thalloid liverworts may be branched.Oil bodies may or may not be present. When
present, oil bodies are typically found in both gametophytic and sporophytic
regions of the liverwort. There are multiple oil bodies per cell. Most liverworts
are small, usually from 220 millimetres (0.080.8 in) wide with individual plants
less than 10 centimetres (4 in) long, so they are often overlooked. The most
familiar liverworts consist of a prostrate, flattened, ribbon-like or branching
structure called a thallus (plant body); these
liverworts are termed thallose liverworts.
However, most liverworts produce flattened
stems with overlapping scales or leaves in two
A thallose liverwort, Lunularia
cruciata
conspicuously
different from

or more ranks, the middle rank is often

the outer ranks; these are called leafy liverworts or

scale liverworts. (See the gallery below for examples.)

Liverworts can most reliably be distinguished from the apparently similar
mosses by their single-celled rhizoids.Other differences are not universal for all
mosses and all liverworts; but the lack of clearly differentiated stem and leaves in
thallose species, or in leafy species the presence of deeply lobed or segmented
leaves and the presence of leaves arranged in three ranks, all point to the plant
being a liverwort. Unlike any other embryophytes, most liverworts contain unique
membrane-bound oil bodies containing isoprenoids in at least some of their cells,
lipid droplets in the cytoplasm of all other plants being unenclosed. The overall
physical similarity of some mosses and leafy liverworts means that confirmation
of the identification of some groups can be performed with certainty only with the
aid of microscopy or an experienced bryologist.

27

2.3.2.1 Subclass Metzgeriidae

Metzgeriidae is a subclass found within the class of Jungermanniopsida
and the species within are characterized as being simple thalloid liverworts.
This

subclass

differs

from

Jungermanniidae such that most of its

species have thallose body rather than
having leaves. These thalloid liverworts
differ from those found in the class
Aneura pinguis

Marchantiopsida

(complex

thalloid

liverworts) because of the lack of tissue

differentiation within the thallus.
a. Order Haplomitriales
Small (to 1 cm tall), upright
plants of H. Hookeri grow sparsely
Haplomitrium mnioides

among other bryophytes and usually

require a conscious search to be found. They are distinctively bright green, with
irregularly shaped leaves arranged around the stem rather than in rows like other
British liverworts. This dioicous species is often fertile, male plants (see inset
photograph) being particularly striking thanks to the orange male organs (similar
in colour to those of Fossombronia).
It is unique in having a branched, rather thick rhizome, instead of the thin
rhizoids of other liverworts. Unlike many liverworts, the spore capsule is
elongated (see photograph); this is only produced occasionally.The most similar
species is Fossombronia incurva (p. 232), which is also dioicous and grows as
small, scattered plants. However, it is pale green rather than the opaque, bright
green of H. Hookeri, and has relatively wider leaves and purple rhizoids.
Damp, gravelly ground, often where shallow water lies in winter, is the
typical habitat of H. hookeri. Such conditions are met on streamsides, edges of
upland tracks,

28

loch shores and the margins of flushes. Archidium alternifolium, Blasia

pusilla and tall rushes (Juncus) are especially characteristic associates. It is also
sometimes found in dune slacks, and in corries and below late-lying snow patches
in the mountains. Montane plants are often much taller (to 7 cm) than those found
elsewhere
b. Order Brasiales

In wild world known 1 family 2 genera, 5 species . Brasiales is a family of

liverworts in the order Brasiales. Species are large and leafy, and were previously
classified among the Metzgeriales.
c. Order Fossombroniales
is an oligotypic genus the relationships
of which remain ambiguous. , Plants relatively
large, 0.2-1.2 cm wide, 1-5 cm long, nearly
flat or with strongly undulate or crispate
margins, pale to yellowish or deep green often
Calycularia crispula

with secondary goldish, fuscous, reddish to

purplish red or red-brown pigmentation. Branching terminal, less often ventral,

simple or pseudodichotomously , Male usually smaller than female plants with
dorsal scales mostly in several rows along midrib or (rarely) scattered over the
entire apical portion of the thallus
d. Order Metzgeriales
Metzgeriales is an order of
liverworts. The group is sometimes
29

Riccardia multifida

called the simple thalloid liverworts: "thalloid" because the members lack
structures resembling stems or leaves, and "simple" because their tissues are thin
and relatively undifferentiated. All species in the order have a small gametophyte
stage and a smaller, relatively short-lived, spore-bearing stage. Although these
plants are almost entirely restricted to regions with high humidity or readily
available moisture, the group as a whole is widely distributed, and occurs on
every continent except . Members of the Metzgeriales typically are small and thin
enough to be translucent, with most of the tissues only a single cell layer in
thickness. Because these plants are thin and relatively undifferentiated, with little
evidence of distinct tissues, the Metzgeriales are sometimes called the "simple
thalloid liverworts".
Members of the Metzgeriales also differ from the related Jungermanniales
in the location of their archegonia (female reproductive structures). Whereas
archegonia in the Jungermanniales develop directly from the apical cell at the tip
of a fertile branch, archegonia in the Metzgeriales develop from a cell that is
behind the apical cell. As a result, the female reproductive organs, and the
sporophytes that develop within them, are always located on the dorsal surface of
the plant.
2.3.2.2 Subclass Jungermanniidae
a. Order Jungermanniales
Leaves flattened, in 2 or 3 rows, usually broadened to attachment, often
lobed; shoots reclining, erect, or pendent; rhizoids smooth-walled .

Scappania sp

30

b. Order Porellales
Is a small foliose, rheophytic liverwort. The plants grow in dense, palegreenish mats with short, creeping, stoloniform primary stems and ascending to
erect secondary stems up to 3 cm long, which arise in bundles from the creeping
primary stems. The leaves are laxly inserted, not imbricated, obovate with a very
short line of insertion; the lobules are reduced to a small flat. Underleaves are
lacking. The plants are autoicous, with long male spikes (6-20 pairs of bracts) and
gynoecia arranged in cymose clusters of up to 10 obpyriform, 5-keeled perianths.
Sporophytes are often present.Vegetative propagation by multicellular disciform
gemmae arising from leaf-surfaces
c. Ordo Pleuroziales :
Pleurozia is the only genus of liverworts
in the family Pleuroziaceae, which is classified
within the order Jungermanniales. The genus
includes eleven species, and as a whole is both
physically distinctive and widely distributed. The
lower leaf lobes of Pleurozia species are fused,
Riccardia multifida

forming a closed water sac covered by a movable

lid similar in structure to those of the angiosperm genus Utricularia. These sacs
were assumed to play a role in water storage, but a 2005 study on Pleurozia
purpurea found that the sacs attract and trap ciliates, much in the same way as
Utricularia. Observations of plants in situ also revealed a large number of trapped
prey within the sacs, suggesting that the species in this genus obtain some benefit
from a carnivorous habit. After Colura, this was the second report of zoophagy
among the liverworts .
The gametophytes are typically small and the thalli are generally no more
than one cell layer in thickness. Here are a few examples from different orders
within this subclass: The Metzgeriidae include about 30 morphologically diverse
genera that include both leafy and thalloid growth forms. They clearly do not form
a monophyletic group, instead forming a paraphyletic grade comprised of two
31

distinct lineages. A characteristic feature shared with the complex thalloid

liverworts is the development of wings and leaves from a single initial cell that
forms in the newly produced 3-cell apical derivative. Simple thalloid liverworts
are distinguished from the leafy liverworts in that archegonia are produced along
the middle of either the main, lateral, or ventral shoots.
General Life Cycle of Hepaticophyta

32

CHAPTER III
CONCLUTION
Plant bryophyte have crude stems and leaves, but no roots. Instead of
roots, they have "rhizoids." Rhizoids help anchor the plant to a surface, but they
do not absorb nutrients like roots on other plants do. Instead of using flowers to
make seeds, mosses release spores from their leaves. Spores can travel by water
and make new mosses in new locations. Water is very important to mosses, so
they can grow and spread; however, mosses can survive even when they dry out.
When they become wet again, they revive and continue growingLiverworts can
most reliably be distinguished from the apparently similar mosses by their singlecelled rhizoids. Other differences are not universal for all mosses and all
liverworts; but the lack of clearly differentiated stem and leaves in thallose
species, or in leafy species the presence of deeply lobed or segmented leaves and
the presence of leaves arranged in three ranks, all point to the plant being a
liverwort. Unlike any other embryophytes, most liverworts contain unique
membrane-bound oil bodies containing isoprenoids in at least some of their cells,
lipid droplets in the cytoplasm of all other plants being unenclosed. The overall
physical similarity of some mosses and leafy liverworts means that confirmation
of the identification of some groups can be performed with certainty only with the
aid of microscopy or an experienced bryologist.

33

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