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Pteridophytes
Meaning of Pteridophytes:
Pteridophytes (pteron — feather, phyton — plants) are the non-flowering vascular plants. Hence they
may be defined as ‘vascular cryptogams’. They are represented by about 400 genera and about 10,500
species including both the living and fossil plants.
Characteristics of Pteridophytes:
 Pteridophytes are non-flowering (seedless) vascular plats.
 Pteridophytes generally occur in cool, damp, shady places. Some are xerophytic (Selaginella,
Equisetum). Azolla, Salvinia, Marsilea are aquatic.
 Pteridophytes range from small herbaceous annual (Azolla, Salvinia) to large perennials trees (Cyathea,
Alsophila). Mostly, pteridophytes are herbaceous in nature.
 There is a regular heteromorphic alternation of generation where both the sporophytic and
gametophytic generations is small or inconspicuous and are nutritionally independent.
 Sporophyte is the predominant plant body, differentiated into root, stem and leaves.
 The stem is generally branched either dichotomous or monopodial.
 The primary roots are ephemeral and are soon replaced by adventitious roots.
 Leaves can be microphyllous (Selaginella) or macrophyllous (most ferns).
 The leaves may be simple, small and sessile (e.g., Lycopodium, Selaginella); scale like (e.g., Equisetum)
or compound, large and petiolate as in ferns (e.g., Pteris, Marattia).
 Leaves bearing sporangia are called sporophylls. Meiospores are formed inside sporangia by sporic
meiosis
 Pteridophytes are polysporangiate, either homosporous or heterosporous, i.e. with two types of spores,
microspores and megaspores e.g., Selaginella, Salvinia, Marsilea.
 Pteridophytes are free sporangiate where isospores or micro- and megaspores are released through the
dehiscence of sporangia.
 Presence of multicellular sex organs i.e., antheridia and archegonia.
 Water is essential for fertilisation where bi or multi-flagellated sperms swim over a thin film of water
and are attracted chemotactically towards the archegonium.
 The zygote undergoes repeated mitotic divisions to form embryo. The first division of the zygote
determines the polarity of the sporophyte.
 A well-developed vascular system, comprising of xylem and phloem, is present. Cambium is generally
absent, thus secondary growth does not take place in majority of the pteridophytes. The nature of stele
varies in different groups.
Two types of leaves are found in pteridophytes:
(a) Microphylls or Microphyllous Leaves:
 Small & simple leaves.
 Single veined at the middle region.
 No leaf gap formation in the stem
 No tissue differentiation in the leaves.
 Leaf is isobilateral.
 Lycopodium, Selaginella, Isoetes.
(b) Megaphylls or Megaphyllous Leaves:
 Leaves are large &compound (pinnated).
 Venation is complex
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 Mesophyll is palisade and spongy.\

 Leaf is dorsiventreal.
 Prominent leaf gap in the stem.
 Pteris, Marattia, Marsilea.

Stelar System in Pteridophytes:

The central cylinder or core of vascular tissue, consisting of xylem, phloem, pericycle, and,
sometimes, medullary rays and pith, is designated as stele.
Types of Steles in Pteridophytes:
There are two principal types of stelar organisation in pteridophytes — Protostele and Siphonostele.
I. Protostele:
It is characterised by the absence of central column of pith. In its simplest form it is merely
composed of a central strand of primary xylem surrounded by a cylinder of phloem. From the
phylogenetic as well as ontogenic standpoint the most primitive type of stele is the protostele.
All other types of steles have been derived from it in the course of evolution. Protosteles are found in
many living primitive vascular plants e.g., Psilotum, Tmesipteris, Selaginella, Lycopodium.
In advanced vascular cryptogams (ferns), protostele is generally associated with the sporeling stage.
Different types of the protostele encountered in the pteridophytes are:
(a) Haplostele:
It is the simplest and most primitive type of protostele consisting of a central solid and smooth core
of xylem surrounded by a ring of phloem (Fig. 7.133A) e.g., Rhynia, Horneophyton, Cooksonia (fossil
pteridophytes); Tmesipteris, Selaginella (living pteridophytes).
(b) Actinostele:
A protostele in which the contour of the core of xylem is lobed or star- shaped in transectional view
and is known as actinostele (Fig. 7.133B). The phloem, instead of forming a continuous ring, is
situated in the form of small groups in between the radiating arms of the xylem e.g., Psilotum,
Lycopodium serratum (living pteridophytes); Asteroxylon, Sphenophyllum (fossil pteridophytes).
(c) Plectostele:
In this type of protostele, the xylem cylinder breaks into several separate plates (Fig. 7.133C). The
plates are more or less parallel and each xylem plate is surrounded by phloem, e.g., aerial shoots and
cone axis of Lycopodium clavatum and L. volubile.
(d) Mixed Protostele:
In this type of protostele, the solid xylem core is broken into small irregular groups of tracheids that
are embedded in the phloem (Fig. 7.133D). e.g., stem of Lycopodium cernuum.
II. Siphonostele:
It is considered as an advancement in anatomical development over protostele. In siphonostele, the
vascular tissues are arranged in the form of a cylinder, with a distinct pith in the centre. The
siphonostele and its variations are found frequently in the sphenopsids and ferns.
On the basis of position of phloem, siphonostele may be divided into two types:
(i) Ectophloic Siphonostele:
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In this type of siphonostele, the phloem occurs as a single ring only on the external side of the xylem
core (Fig. 7.133E). The pith is central in position and the phloem is externally surrounded by the
pericycle and endodermis. e.g., Equisetum, Osmunda, Schizaea.
(ii) Amphiphloic Siphonostele:
In this case, the vascular cylinder consists of xylem surrounded on both sides (external and internal)
by phloem (Fig. 7.133F). This type of stele characteristically has two endodermal layers (outer
endodermis and inner endodermis) e.g., Marsilea, Adiantum, Dryopteris.
Depending upon the presence or absence of leaf trace and branch trace, Jeffery (1910)
divided siphonostele into two groups:
(i) Cladosiphonic:
It is characterised by the absence of leaf traces e.g., lycopsids.
(ii) Phyllosiphonic:
It is characterised by the presence of both leaf and branch traces, e.g., members of Filicales.

On the basis of non-overlapping or overlapping gaps, the following three types of

siphonostele are categorised:
(i) Solenostele:
It is a type of siphonostele where the leaf gaps are successive so that there is only one break in the
vascular cylinder at any one given point (non-overlapping). The solenostele may be ectophloic (Fig.
7.133G) or amphiphoic (Fig. 7.133H).
(ii) Dictyostele:
A siphonostele with overlapping leaf gaps is known as dictyostele (Fig. 7.1331). In this case, the leaf
gaps are large and overlap each other (many occur at a point), and the vascular cylinder thus appears
to be highly dissected. It is usually found in ferns where rhizome is very small and crowded with
leaves e.g., Dryopteris, Pteris, Ophioglossum. A dictyostele has many scattered vascular strands.
Each strand is known as ‘meristele’. All the meristeles are surrounded by a common endodermis.
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(iii) Polycyclic Stele:

This is a complex type of siphonostele, where vascular tissue is present in the form of two or more
concentric cylinders (Fig. 7.133J). e.g., two rings of vascular tissue in Pteridium aquilinum, three in
Montania peotinata and four in Pteris podophylla.
Polycyclic stele may be of two types: i.e., Polycyclic solenosteles and polycyclic dictyostele. A
polycyclic stele having an outer cylinder of solenostelic type is called polycyclic solenostele, while the
outer cylinder of dictyostelic type is called Polycyclic dictyostele.

Reproduction in Pteridophyta
The basic life cycle pattern of pteridophytes shows a regular heteromorphic alternation of generations
between a gametophyte (sexual) phase and a sporophyte (asexual) phase.
The main plant body is sporophytic which forms a dominant phase in the life cycle. The gametophytic
generation bears male and female sex organs, antheridia and archegonia.
The antherozoids (male gamates) produced in large numbers are motile, while the eggs (female gametes)
are non-motile and are borne singly in archegonia. Fusion between an egg and an antherozoid results in the
formation of a diploid (2n) zygote. The zygote develops directly by mitotic divisions into sporophyte. The
sporophyte plant develops sporangia which produce haploid spores through reductional division (i.e.,
meiosis). The life cycle is then completed when these spores germinate and grow into haploid gametophytes
(n).
Sporophyte
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The sporophytic plant reproduces by means of spores produced in the sporangia (singular: sporangium).
Sporangium:
 The position of sporangia may vary in different groups;
 They may be borne on the stems i.e., cauline (e.g., Psilotum; Rhynia) or on the ventral (adaxial) surface
of the leaves i.e., foliar (e.g., Lycopodium, Selaginella) or in the axil of the leaves (e.g., Ophioglossum).
 The sporangia containing leaves are called sporophylls.
 The sporophylls may be scattered (e.g., Lycopodium selago), uniformly distributed (e.g., Pteris,
Adiantum and other ferns) or grouped in definite areas to form strobili (Selaginella, Equisetum).
 In some aquatic pteridophytes the sporangia are present within a specialised structure, called
sporocarps (e.g., Azolla, Salvinia, Marsilea).

On the basis of mode of development, the sporangia are of two types, the eusporangiate and the
Leptosporangiate.

In some forms (e.g., ferns) the sporangia are aggregated in clusters termed sori (singular sorus).
On the basis of maturity of the sporangia, the sori are of three types:
(a) Simple Sorus:
A sorus in which all the sporangia originate, grow and mature at the same time (e.g., Botrychium,
Ophioglossum). The forms showing such condition are grouped together as simplices (Fig. 7.2A).
(b) Gradate Sorus:
Here sporangia develop over a period of time, where the central part of the sorus has mature sporangia and
the peripheral part has younger sporangia (e.g., Hymenophyllum, Marsilea, Cythea). The forms showing
this condition is called Gradatae (Fig. 7.2B).

(c) Mixed Sorus:

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Here the mature and immature sporangia of different ages are arranged in an irregular fashion (e.g.,
Pteridium, Pteris, Adiantum), and the condition is termed as Mixtae (Fig. 7.2C).
Spores:
 Meiotic (reduction) divisions of spore mother cells produce numerous haploid spores inside the
sporangium. If all the spores produced are of equal sizes and shapes, then the plant is called
homosporous (e.g., Lycopodium, Equisetum, Dryopteris), and if they are of two different sizes and
shapes the plant is called heterosporous (Selaginella, Isoetes, Marsilea).
 In the heterosporous type, the two different types of spores are produced in separate sporangia. The
smaller spores are called microspores or male spores and are produced in microsporangia. The
microspores are produced in large numbers.
 The larger spores which are produced in smaller numbers are termed megaspores and are developed in
megasporangia. Microspores, after germination, produce male gametophyte, while megaspores produce
female gametophyte.
 Sporophylls with megasporangia are called megasporophylls, while sporophylls with microsporangia are
called microsporophylls.
 In homosporous member, spores germinate to produce monoecious i.e., homothalic gametophyte
bearing both male and female sex organs. In this case, the sex determination takes place at the time of
formation of antheridium and archegonium
 In heterosporous member, micro- and mega- spores, on germination, produce male and female
gametophytes, respectively (i.e., heterothallic or dioecious). Here, the sex determination takes place
much earlier, at the stage of sporogenesis
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Gametophyte:
 The spores germinate to form haploid gametophytes or prothalli. The gametophytes of pteridophytes are
small and inconspicuous as compared to the sporophytes.
 The germination of spores in the homosporous forms are of three types.
a) Bipolar (e.g., Lycopodium, Equisetum)
b) Tripolar (e.g., Hymenophyllum)
c) Amorphous and irregular (e.g., Angiopteris).
 The gametophytes are of two types. In homosporous forms, the development of gametophyte is
exosporic in which the prothallus develops outside the spore wall (e.g.-, Psilotum, Lycopodium,
Ophioglossum). Therefore, it is vulnerable to outside environment.
 Gametophytes that develop from heterospores are endosporic in which the development of prothallus
is confined within the spore wall (e.g., Selaginella, Isoetes and Marsilea). The development of endosporic
gametophyte is independent of external environment.
Sex Organs: The gametophytes or prothalli (singular – prothallus) bear the sex organs, the male
antheridia (singular antheridium) and female archegonia (singular archegonium).
Antheridium:
 The antheridium is a sessile or shortly stalked globular structures surrounded by a well-defined jacket
inside containing androcytes or antherozoid mother cells.
 Each androcyte gives rise to a single motile antherozoid.
 The number of antherozoids per antheridium varies from 4 (e.g., Isoetes) to few thousands (e.g.,
Ophioglossum).
 The antherozoids are unicellular, uninucleated and spirally coiled bearing apical flagella (e.g.,
Lycopodium, Selaginella) or are multiflagellate (e.g., Psilotum, Isoetes, Equisetum and ferns).
Archegonium:
 The archegonium is a flask-shaped structure consisting of a basal, swollen venter and a short neck, the
venter is embedded in the prothallus while the neck is projected.
 The venter encloses an egg and a ventral canal cell. Neck is made up of vertical rows of neck cells with
neck canal cells inside.
 At maturity the neck canal cells disintegrate to form a passage for the antherozoids to reach the egg.
Fertilisation:
 The disintegration of neck canal cells also produces a mucilagenous substance which contains
organic compounds like malic and fumaric acid. These substances act as sperm attractant.
 Water is essential for fertilisation and sperms swim over a thin film of water and attracted
chemotactically towards the archegonium.
 The antherozoid and egg — of haploid chromosome number — fuse to form a diploid zygote, which is
the mother cell of sporophytic generation.
The Embryo:
 The zygote undergoes repeated divisions to form embryo. Further development of embryo results into a
well-developed sporophyte differentiated into roots, stem and leaves. The first division of the zygote
determines the polarity of the sporophyte.
 According to the polarity, the embryo may be categorised into two types, Exoscopic embryo and
Endoscopic embryo. In excoscopic embryo, the shoot-forming apical cell is directed towards the neck of
the archegonium i.e., directed outward (e.g., Psitotum, Equisetum, Ophioglossum).
 In endoscopic embryo the shoot-forrping apical pole is directed downward i.e., towards the base of the
archegonium (e.g., Lycopodium, Selaginella, Isoetes).
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Classification of Pteridophytes
(1) Psilopsida:
(i) has 8 species.
(ii) Order-psilotales
Genus- psilotum spp, tmesipteris spp.
(iii) Roots are absent.
(iv) The organization of the plant body of the members is very simple. It is differentiated into a
subterranean (underground) rhizome and an erect aerial portion.
(v) Rhizome bears tufts of unicellular Rhizoids.
(vi) Aerial portion is sparingly or profusely branched. The branching is usually of dichotomous type.
(vii) Aerial axis may be leafless or sometimes may bear scaly appendages (e.g., Psilotum) or large foliage
leaves (e.g., Tmesipteris)
(viii) The vascular tissue is of primitive type i.e., simple, cylindrical protostele with annular or spiral
racheids.
(ix) The reproductive organs are in the form of sac like sporangia.
(x) Sporangia are borne at the apex of the aerial shoots. They are either solitary (e.g., Rhynia) or in groups
and terminal in position. There was nothing like that of sporophyll.
(xi) Sporangia always bearing the same type of spores i.e., they are homosporous.
(xii) The gametophyte is known only in Psilotum and Tmesipteris.
(xiii) The gametophyte is cylindrical or branched, subterranean and colourless.
(xiv) Sex organs are partially embedded in the prothallus.
(xv) Antherozoids are multiciliate in Psilotales.

(2) Lycopsida:
(i)Has 963 species
(ii) it has the following orders;
1) Lycopodiales (186 spp)
a) Genus; Lycopodium.
2) Selaginellalis (700 spp)
a) Genus; selaginella.
3) Isotales (77 spp)
a) Genus; Isoetes spp
(iii) Its history indicates that these Pteridophytes developed during the Devonian period of the Palaeozoic
era.
(iv) The leaves are small (microphyllous), simple with a single mid vein.
(v) They are usually spirally arranged, sometimes in opposite fashion and or even in whorls.
(vi) In some cases the leaves are ligulate (e.g., Selaginella, Isoetes). The ligule is present at the base of each
leaf.
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(vii) The vascular tissue may be either in the form of plectostele, siphonostele or sometimes even polystele.
(viii) Leaf gaps are absent.
(ix) Sporangia are quite large in size and develop on the adaxial surface of the leaves (sporophylls).
(x) Sporophylls are loosely arranged and form strobilus.
(xi) Some members are homosporous (e.g., Lycopodium) while others are heterosporous (e.g., Selaginella).
(xii) Antherozoids are biflagellate or multiflagellate.
(xiii) Gametophytes which are in the form of prothalli are formed by the germination of spores.
(xiv) Heterosporous forms have endoscopic gametophytes while in homosporous forms the gametophyte is
exoscopic.

(3) Sphenopsida:
(i) Has 25 species
(ii) Includesthe following order;
1) Order; Equisetales.
a) Genus; Equisetum
(iii) These Pteridophytes evolved during the Carboniferous period of the Palaeozoic era.
(iv) The stem in majority of the forms is long, jointed or articulated and is ribbed i.e., having ridges and
grooves.
(v) Stem is divisible into nodes and internodes and is developed as upright aerial branches from the
underground creeping rhizome.
(vi) Leaves are thin, small, scaly brown and are arranged in transverse whorls on the nodes of the aerial
branches.
(vii) Branches also develop in whorls from the axil of the scaly leaves.
(viii) As the foliage leaves are reduced to scales, the process of photosynthesis is taken up by the stem and
hence it becomes green.
(ix) The stem has a solid protostele (e.g., Sphenophyllum) or medullated protostele (e.g., Equisetum).
(x) Sporangia are developed at the apex of the fertile branches in whorls forming compact cone.
(xi) Living members are homosporous but some fossil forms are heterosporous (e.g., Catamites).
(xii) Spores germinate to give rise to gametophytes (prothalli) which may be monoecious or dioecious.
(xiii) Antherozoids are large and multiflagellate.
(xiv) Embryo is without suspensor.

(4) Pteropsida:
(i) This class includes the plants which are commonly known as ‘ferns’, it has > 10,000 spp.
1) Sub-class- Eusporangiate
a) Order- Ophioglossales (70 spp)
i) Genus- Opioglossuum.
b) Order- osmundales (19 spp)
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i) Genus- osmunda.
c) Order- Marratiales (>200 spp)
i) Genus- Marratia.
2) Sub-class- Leptosporangiate.
a) Order- Filicales (7800 spp)
i) Genus- Oryopteris spp
b) Order- Marsileales (67 spp)
i) Genus- Marsiles
c) Order- Saviniales (16 spp)
i) Genus- Salvinia.
(ii) These Pteridophytes were originated during the Devonian period.
(iii) They occur in all types of habitats. Majority of the ferns are terrestrial and prefer to grow in moist and
shady places. Some are aquatic (e.g., Azolla, Salvinia, Marsilea), xerophytic (e.g., Adiantum emarginatum),
epiphytic (e.g., Asplenium nidus), halophytic (e.g., Acrostichum aureum) or climbing (e.g., Stenochlaena).
(iv) Some members are very small while some members are tall tree like (e.g., Angiopteris).
(v) Majority of the members (except some tree ferns e.g., Angiopteris) have short and stout rhizome. The
rhizome may be creeping, upright or growing above the soil.
(vi) Leaves are large, may be simple (e.g., Ophioglossum) or compound (majority of the ferns for example,
Pteridium, Marsilea, Adiantum etc.) and described as fronds.
(vii) Young fronds are circinately coiled.
(viii) Leaves are exstipulate (e.g., Filicales) while stipulate in some other groups.
(ix) The vascular cylinder varies from a protosete to a complicated type of siphonostele.
(x) Vegetative reproduction takes place by fragmentation (e.g., Adiantum, Pteridium), stem tubers e.g.,
Marsilea), adventitious buds (e.g., Asplenium bulbiferum) or by apogamy (e.g., Marsilea).
(xi) Sporangia arise from placenta (a swollen cushion of cells) in groups (sori).
(xii) Sori develop on the margins or abaxial surface of the leaves (sporophylls) or leaflets.
(xiii) Sori are protected by true (e.g., Marsilea) or false indusia (e.g., Adiantum, Pteris).
(xiv) The sporangial development may be leptosporangiate (e.g., Osmunda) or eusporangiate type e.g.,
Ophioglossum).
(xv) The sporangia in most cases have a distinct annulus and stomium.
(xvi) Members may be homosporous (e.g., Pteris, Adiantum etc.) or heterosporous (e.g., Marsilea,
Regnellidium, Azolla, Salvinia etc.)
(xvii) Spores on germination form autotrophic prothalli (gametophyte).
(xviii) Antheridia and archegonia are partially or completely embedded in the gametophyte.

Economic Importance of Pteridophytes:

i. Pteridophytes Used as Food:
The young leaf tips of ferns, the circinate ptyxis or the chroziers are used as vegetable. Leaves of Marsilea
are used as vegetable.
The rhizome of many ferns such as Pteris, rich in starch, is used as food.
The corm (modified stem) of Isoetes is used as food by pigs, ducks and other animals.
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ii. Pteridophytes Used as Fodder:

Dry fronds of many ferns form the livestock for catties. The quadrifid lamina of Marsilea resembles a clover
(Trifolium) has been used as fodder for animals as a substitute for clover.
iii. Pteridophytes Used as Medicine:
The spores of Lycopodium have been widely used in pharmacy as protective dusting powder for tender skin
and also as water-repellants. The foliages of Lycopodium are used as tincture, powder, ointment and cream
as a stomachic and diuretic.
The foliage decoction is used in homeopathy to treat diarrhoea, bladder irritability, eczema, rheumatism,
constipation and inflammation of liver.
Equisetum is rich in silicic acid and silicates. Potassium, aluminium and manganese, along with fifteen
types of flavonoid compounds, have been reported from Equisetum. The flavonoids and saponins are
assumed to cause the diuretic effect. The silicon is believed to exert connective tissue-strengthening and
anti-arthritic action.
Several ferns have been used as herbal medicine.
The decoction of Asplenium is used for cough and a good hair wash. The expectorant of Polypodium is used
as a mild laxative, while the tonic is used for dyspepsia, loss of appetite and hepatic problem.
The root decoction of Osmunda regalis is used for treatment of jaundice. The ointment made from its root is
used for application to wound. The extraction of Osmanda vulgaris, commonly known as ‘Green oil charity’,
is used as remedy for wounds.
The chemically active principal ‘Marsiline’ isolated from Marsilea is found to be very effective against
sedative and anti-convulsant principal.
The rhizome and frond bases of Dryopteris have been used to determine the origin and pathways of
dispersed pathogenic insects like corn ear- worm. The preparation of Ophioglossum vulgatum as ‘Green oil
charity’ is also used as remedy for wounds.
iv. Pteridophytes Used as Horticultural Plants:
Many species of pteridophytes are cultivated for their aesthetic value.
Some epiphytic species of Lycopodium (e.g., L. phlegmaria, L. lucidulum) are aesthetically more valued and
can be grown on hanging baskets.
Several species of Selaginella are used as a ground cover in an undisturbed area because of their decent
foliage and colour.
Several ferns such as Angiopteris, Asplenium, Marattia, Microsorium, Nephrolepis, Phymatodes, etc., have
aesthetic values for their beautiful habit, graceful shape of the leaves, and beautiful soral arrangement.
Thus, these characteristics make them horticulturally important plants.
v. Pteridophytes Used as Biofertiliser:
Azolla is a free-floating water fern which can multiply very quickly through vegetative propagation. There
are hundreds of moss-like leaves harbouring live colonies of dinitrogen fixer Cyanobacterium — Anabaena
azollae.
The relationship between the alga and Azolla is symbiotic where the alga provides nitrogen to the plant.
Thus, Azolla in full bloom in the waterlogged rice fields may serve as a green manure. Rice farmers of our
country are using Azolla as biofertiliser for the better production of their crops.
vi. Pteridophytes Used as Indicator Plants:
Like angiosperms, pteridophytes are being used as indicator plants.
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Equisetum accumulates minerals, especially gold, in their stem. The rate of accumulation even reaches up
to 4.5 ounce per ton. Equisetum may be referred to as gold indicator plants which help in searching a region
for gold ore deposits. Similarly, Asplenium adulterinum is an indicator of nickel and Actinopteris australis
is a cobalt indicator plant. Thus, these plants are found to be valuable in prospecting for new ore deposits.
vii. Pteridophytes Used for Various Purposes:
The stem of Equisetum was used for polishing wood in ancient times and to clean utensils.
The roots and stems of Osmunda are used to make beds for growing orchids. Water boiled with Lycopodium
clavatum is used for dyeing the woollen clothes which becomes blue when dipped in a bath of Brazil wood.
The powder of Lycopodium is highly inflammable and is used in pyrotechny and for artificial lighting. Thus,
Lycopodium powder finds its wide use in demonstration of artificial lighting on the stage, because it
disperses easily in the air and only a small quantity is needed to produce an explosion.