Pteridophyta O.P.

Sharma
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Series on Diversity of Microbes and Cryptogams
 O P Sharma, with his 36 research articles published in national and
international journals, 31 books written for university students, and 40 years
of teaching experience, is an able researcher, established Indian author, and
an experienced teacher. His areas of research include pollen morphology,
angiosperm’s anatomy and mycology, with special focus on Indian Cyperaceae
(with particular interest on Cyperus). Over a dozen of Dr Sharma’s books have
been published through internationally known publishers. He has also revised
Economic Botany, an internationally renowned text by the late Professor Albert
F Hill (Harvard University, USA), a publication of McGraw-Hill, New York.
Encouraging reviews of his books have been published in reputed scientific
journals such as Current Science, and his books on Practical Botany have received appreciations from
some eminent botanists including J D Dodge (England), T Christensen (Denmark), J M Herr (USA)
and C R Metcalfe (England).
Immediately after completing his postgraduation (MSc) in Botany with first division from CCS
University, Meerut, and thereafter obtaining a PhD from the same university, Dr Sharma started his
teaching career in 1967 as a faculty member of the Botany Department, Meerut College, Meerut,
and retired from active services as a Reader from the same department in 2007. Besides attending
several national and international workshops, symposia and conferences during the four decades of his
teaching career, Dr Sharma is still enjoying his post-retirement innings as an active author.
Recently, Dr O P Sharma revised some of his widely circulated books published through
McGraw-Hill Education, which include Plant Taxonomy (released first in 1993 and reprinted 19
times), Textbook of Algae (released first in 1986 and reprinted 20 times) now titled Algae, and Textbook
of Fungi (released first in 1989 and reprinted 17 times) now titled Fungi and Allied Microbes.
Series on Diversity of Microbes and Cryptogams

O P Sharma
Reader (Retired)
Department of Botany
Meerut College, Meerut

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Series on Diversity of Microbes and Cryptogams: PTERIDOPHYTA

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 DEDICATION
“Gurus” have a bigger place in life than parents; that’s what my parents told me,
and I am very proud to announce that one of my gurus has been
Professor S K Roy.
He taught me the A, B, C.... of Pteridophyta and many more things for a better life.
I humbly dedicate this book to Professor S K Roy.
O P Sharma
 Contents

1. Introduction .................................................................................................................... 1.1-1.22

2. Development of Pteridology in India ................................................................................. 2.1-2.3
3. Threatened Pteridophytes of India: Handle Them With Care ........................................... 3.1-3.6
4. Economic Importance of Pteridophytes ............................................................................ 4.1-4.5
5. Classification of Pteridophytes......................................................................................... 5.1-5.10
6. Psilophytopsida ............................................................................................................... 6.1-6.14
7. Psilotopsida ..................................................................................................................... 7.1-7.19
8. Lycopsida: Protolepidodendrales...................................................................................... 8.1-8.6
9. Lycopsida: Lycopodiales ................................................................................................. 9.1-9.21
10. Lycopsida: Lepidodendrales ........................................................................................ 10.1-10.6
11. Lycopsida: Isoetales ................................................................................................... 11.1-11.12
12. Lycopsida: Selaginellales ........................................................................................... 12.1-12.28
13. Sphenopsida(= Equisetopsida) ................................................................................... 13.1-13.32
14. Pteropsida ................................................................................................................. 14.1-14.10
15. Primofilices: Cladoxylales and Coenopteridales ........................................................... 15.1-15.5
16. Eusporangiatae: Marattiales...................................................................................... 16.1-16.11
17. Eusporangiatae: Ophioglossales................................................................................. 17.1-17.17
18. Osmundidae: Osmundales ......................................................................................... 18.1-18.13
19. Leptosporangiatae: Filicales (Gleicheniaceae, Schizaeaceae
and Hymenophyllaceae) ............................................................................................... 19.1-19.8
20. Filicales (Cyatheaceae, Matoniaceae, Aspleniaceae and Aspidiaceae) ........................... 20.1-20.6
21. Filicales(Dennstaedtiaceae) ........................................................................................ 21.1-21.13
22. Filicales: Adiantaceae, Dryopteridaceae and Polypodiaceae ....................................... 22.1-22.14
23. Leptosporangiatae: Marsileales ................................................................................. 23.1-23.29
24. Leptosporangiatae: Salviniales .................................................................................... 24.1-24.6
25. Gametophyte in Pteridophytes ................................................................................... 25.1-25.11
26. Apospory, Apogamy and Parthenogenesis ................................................................... 26.1-26.5
27. Origin and Evolution of Pteridophytes......................................................................... 27.1-27.6
28. Telome Theory and Enation Theory ............................................................................ 28.1-28.8
29. Cytogenetics: Polyploidy, Chromosome Number and Organelle Genome..................... 29.1-29.7
30. Stelar System ............................................................................................................... 30.1-30.6
31. Heterospory and Seed Habit ........................................................................................ 31.1-31.7
32. Morphogenesis: Spore, Prothallus, Sexuality and Sporophyte ...................................... 32.1-32.7
33. Nuclear DNA Amounts in Pteridophytes ...................................................................... 33.1-33.5

Answers to Questions ......................................................................................................... Q.1-Q.6

Recommended Readings ..................................................................................................... R.1-R.8
Index................................................................................................................................. I.1-I.12
 PREFACE

My main objective for writing this book has been the complete non-availability of any Indian text on
Pteridophyta that discusses almost all topics of the syllabi of Indian universities in the light of modern
developments of the last three to four decades. Even the best available and globally accepted text of an
Indian author on Pteridophyta was revised last in 1977. After that, a book appeared in 2002, but this too
discusses only selected topics of pteridology by about a dozen Indian authors, is more research-oriented
and least student-friendly.
Some recently published books of foreign authors restrict themselves to a detailed treatment of
the distribution, structure, reproduction and evolution of selected pteridophytic taxa, and a few more
present a very elementary treatment of pteridophytes as a whole, together with discussion of some
selected general topics. These too do not fulfill the need of Indian students.
A book on Pteridophyta was, therefore, planned to fulfill this lacuna of a suitable modern text
for undergraduate and postgraduate students of Indian universities. It was even more considered an
essentiality because Pteridophyta (Vascular Cryptogams) constitutes a compulsory course of study
for UG and PG students of all Indian universities teaching Botany as a subject. Thus, my goal was to
prepare a textbook that would adequately and interestingly display to both students and teachers the
latest available state of our knowledge of pteridophytes. I must say it was indeed a formidable and
challenging task!
The book Pteridophyta contains 33 chapters, of which 18 chapters (7 to 24) discuss all recognised
classes, orders and selected living and fossil genera of this group, and the remaining 15 chapters
(1 to 6 and 25 to 33) discuss the general aspects. Life-history details of almost all major pteridophytes
living today in the world are also discussed duly in the book.
Considerable palaeobotanical information has been included in order to provide a historical
background for the more extended discussions of the surviving genera of pteridophyta. Particular
attention has been given in the form of chapterwise treatment of fossil pteridophytes belonging to
Psilophytopsida, Protolepidodendrales, Lepidodendrales, Cladoxylales and Coenopteridales. Fossils
of orders Hyeniales, Sphenophyllales and Calamitales of Sphenopsida have also been given due
importance in the text.
Some major highlights of the book are as follows:
• Complete coverage of all important topics in Pteridophytes
– Development of Pteridology in India
– Gametophyte in Pteridophytes
xviii ®
– Apospory, Apogamy and Parthenogenesis
– Origin and Evolution of Pteridophytes
– Telome Theory and Enation Theory
– Stelar System
– Heterospory and Seed Habit
• New and modern topics
– Threatened Pteridophytes of India: Handle Them with Care
– Nuclear DNA Amounts in Pteridophytes
• Four application-based chapters
– Economic Importance of Pteridophytes
– Classification of Pteridophytes
– Cytogenetics: Polyploidy, Chromosome Number and Organelle Genome
– Morphogenesis: Spore, Prothallus, Sexuality and Sporophyte
• Examination preparation tools to help students
– 623 chapter-end questions with answers of many of them at the end of the book
– 17 tables of comparison spread in the entire text
– 14 illustrated and pictorial life cycles of major living genera of Pteridophytes
– Detailed life histories of more than 30 genera of Pteridophytes
– Due coverage of almost all fossil groups of Pteridophytes
• Up-to-date coverage as per details given in
– Classification of Pteridophytes proposed by Smith et al. (2006)
– Books and research articles published up to 2010 and mentioned under Recommended
Readings at the end of the book
• Rich pedagogy
– 320 well-labelled line diagrams to make the text clear to the reader
– Dozens of website addresses, related to pteridophytes, spread in the entire text
The book caters to all needs of readers, particularly undergraduate and postgraduate students of
Botany and Agriculture of all universities the world over, in general, and of India in particular. It should
also be equally helpful for students preparing for all competitive examinations, especially AIPMT,
CPMT, IFS, IAS, PCS, NET, SLET, etc.
I wish to acknowledge my special thanks to all those individuals who have offered me advice and
suggestions in the preparation of the manuscript of this book. In particular, I am greatly obliged to my
dear student Professor Mayank Uday Charaya and his team of research students of CCS University,
Meerut, who took pains in searching the latest information for this book on the Internet. For critically
reviewing the entire manuscript, prior to its publication, I am particularly grateful to the following
reviewers:
• J K Mishra, of Sri Jai Narain PG College, Lucknow, Uttar Pradesh
• Beulah Vijayakumar, Stella Maris College, Chennai
• Jyoti Pratap Handique, Gauhati University, Guwahati, Assam
 ® xix

Several of the suggestions made by them and incorporated in the final manuscript have really
upgraded the present book.
I am also happy to pay tributes to my wife, Dr (Mrs) Kanti Devi Sharma, PhD, for her unfailing
patience and assistance with the proofreading. Although deeply unwilling, I had no option but to ignore
my dearest grandchildren (Kuhu, Karan and Krish) during the long period of preparation of this book.
I express my fullest love for them at this stage, and honestly wish they become worthy citizens of this
great country. Last, but not the least, I express my gratitude to the publishing team of McGraw-Hill
Education who made this book see the light of day.
Comments and suggestions for improvement of this book may be sent to tmh.sciencemaths
feedback@gmail.com or directly to me.

O P Sharma
09837566555
0121-2401400
 CHAPTER 1
Introduction

1.1 PTERIDOPHYTES OR VASCULAR CRYPTOGAMS: WHAT ARE THEY?

Pteridophytes are a group of higher cryptogams possessing a well-developed vascular system, hence
also called vascular cryptogams. In the traditional system of classification of plants, the other groups
of cryptogams are algae, fungi and bryophyta, which together represent lower cryptogams.
The name Pteridophyta was originally assigned to this assemblage of plants by Haeckel (1866)
because of the presence of pinnate or feather-like leaves (pteron meaning feather and phyton meaning
plant).
Engler (1886) treated Pteridophyta as a division of the subkingdom Embryophyta, which has
also been named Telomobionta by Takhtajan (1964) and Embryobionta by Cronquist et al. (1968).
According to Parihar (1977), ‘Pteridophytes are the most primitive living and fossil vascular plants’.
In the plant kingdom, pteridophytes occupy a position in between bryophytes and gymnosperms, and
therefore they have some similarities with bryophytes on the one hand (e.g. presence of sterile jacket
around the antheridium and archegonium; water or moisture requirement for fertilization; presence
of alternation of generations; formation of spores, etc.) and with gymnosperms (e.g. sporophytic
plant body; differentiation of sporophytes into roots, stem and leaves; presence of vascular tissue for
conduction; independent nature of the sporophyte, etc.) on the other hand.
In the so-called ‘modern system of classification’, as mentioned by Fuller and Tippo (1954),
pteridophytes are included in the phylum Tracheophyta, which is divisible into four subphyla, viz.,
Psilopsida, Lycopsida, Sphenopsida and Pteropsida. The subphylum Pteropsida, however, includes
two classes, i.e. Gymnospermae and Angiospermae, along with a third class, named Filicinae, which
includes the ferns. The pteridophytes of the older system are, therefore, restricted only up to the
subphyla Psilopsida, Lycopsida, Sphenopsida and a class Filicinae of the subphylum Pteropsida in the
modern classification of the plant kingdom.
1.2 ®
1.2 DEFINITIONS OF PTERIDOPHYTA

In the Longman Illustrated Dictionary of Botany, Andrew Sugden (1984) defined pteridophyte
as “a member of the division Pteridophyta, which includes ferns, clubmosses and horsetails. In
pteridophytes, the sporophyte is the main vegetative stage of the life cycle. The gametophyte is small,
and is independent of the sporophyte, as in bryophytes”. Large tree-like pteridophytes were common
during the Carboniferous period (i.e. 345–280 million years ago), and were fossilized to form coal.
In the Chambers Biology Dictionary, M B Walker (1989) defined Pteridophyta as “a division of
the plant kingdom containing all the vascular plants which do not bear seeds, i.e. ferns, clubmosses,
horsetails, etc. There is an alternation of generations of, typically, a smaller, more or less thalloid,
independent gametophyte and larger, longer-lived sporophyte usually with roots, stems and leaves”.
In The Penguin Dictionary of Botany, E Tootill (1984) defined Pteridophyta as “a division containing
all the nonseed-bearing vascular plants. Its members show a heteromorphic alternation of generations
and the gametophyte is often nutritionally independent of the sporophyte. The sporophyte differs from
that of seed-bearing plants in lacking vessels in the xylem”.
Recently, Masahiro Kato (2005), a Japanese pteridologist, defined pteridophytes as “free-sporing
vascular land plants that evolutionarily link bryophytes and seed plants”. They “have a long (420
million years) evolutionary history” and possess a total of about 12000 species, which may be primitive
or advanced.

1.3 VALIDITY OF THE CLASSIFICATION OF PTERIDOPHYTA

Jeffrey’s (1902) work created some controversy about the validity of classification of Pteridophyta.
According to him all vascular plants can be divided into two different types called stocks. Of these
two ‘stocks’, the Lycopods and Horsetails have microphyllous leaves and lack any leaf gaps, and
thus represent one ‘stock’, i.e. Lycopsida. The second ‘stock’, i.e. Pteropsida is being represented
by ferns, gymnosperms and angiosperms. Scott (1923) separated the horsetails as Sphenopsida and
Psilophytales as Psilopsida. And thus four separate ‘stocks’ or ‘divisions’ (Psilopsida, Lycopsida,
Sphenopsida and Pteropsida) were formed. Eames (1936) opined that all these four ‘stocks’ should be
treated collectively as subdivisions of a single division Tracheophyta but Smith (1955) has been of the
opinion that all these four should be elevated to the rank of divisions. However, Smith has also stated
that ‘the ferns seem to be sufficiently distinct from the seed plants and thus be treated in a separate
division’. He mentioned further that in view of the recommendations of the International Code of
Botanical Nomenclature (1950) each division and subdivision should be ended by the words ‘phyta’
and ‘opsida’ respectively. He has therefore suggested that pteridophyta should include four divisions,
viz., Psilophyta, Lepidophyta, Calamophyta and Pterophyta, of which the last division (i.e. Pterophyta)
includes only ferns.

1.4 GENERAL CHARACTERISTICS: AT A GLANCE

1. Pteridophytes are photosynthetic vascular plants in which the female sex organ is an
archegonium.
 ® 1.3

2. Plants show clear alternation of generations.

3. The sporophytic generation is dominant and conspicuous, but dependent on gametophytic
generation early in life.
4. The gametophyte is a free-living generation. It is however, highly reduced and called a
prothallus.
5. The gametophyte bears antheridia and archegonia on same or different prothalli.
6. The prothallus is made of a thin plate of cells, and usually lacks a cuticle.
7. The sporophytic generation consists of true roots, stems and leaves, all with vascular tissue.
8. Leaves possess cuticle and stomata, except in some fossil members and primitive modern
forms.
9. The ovum remains in the archegonium.
10. The spermatozoids swim up to the ovum by chemotaxis to effect fertilization.
11. Members are both homosporous or heterosporous.
12. In heterosporous forms, large megaspores develop into female prothalli, bearing archegonia;
and small microspores develop into male prothalli, bearing antheridia.
13. Vascular tissue is usually confined to sporophytes.
14. Vascular tissue is usually arranged in various kinds of steles.
15. The xylem usually consists of only tracheids.
16. Vessels are absent or very rare.
17. Leaves are microphyllous or megaphyllous.
18. In living pteridophytes, sporangia are usually borne on leaves or on stems associated with them
or on modified leaf-like structures associated with them called sporophylls.
19. Several fossil forms of the Devonian period had no leaves. Sporangia developed on their stems
in these forms.

1.5 LIVING AND FOSSIL PTERIDOPHYTES

Pteridophyta is represented by most primitive living vascular plants, such as Selaginella, Lycopodium,
Equisetum, etc., and many fossil vascular plants such as Rhynia, Hornea, Zosterophyllum,
Asteroxylon, etc.
Reimers’s (1954) classification, also followed in the present book, suggests that fossil pteridophytes
are represented by all the members of the division Psilophytopsida; Protolepidodendrales and
Lepidodendrales of the division Lycopsida; Hyeniales, Sphenophyllales and Calamitales of the division
Sphenopsida; and Cladoxylales and Coenopteridales of the division Pteropsida. On the other hand, living
pteridophytes are represented by all the members of the division Psilotopsida, Lycopodiales, Isoetales
and Selaginellales of the division Lycopsida; Equisetales of the division Sphenopsida; and Marattiales,
Ophioglossales, Osmundales, Filicales, Marsileales and Salviniales of the division Pteropsida.
Pteridophyta was a prominent group of the plant kingdom in the Carboniferous era when tree-like
forms were very common in forests. The living representatives form a small group, which attain their
greatest development in the tropical regions.
1.4 ®
1.6 OCCURRENCE OF PTERIDOPHYTES

Most of living members are terrestrial and prefer to grow in cool and shady places while some are
xerophytic, e.g. Selaginella rupestris. Many pteridophytes are found in aquatic conditions, e.g.
Marsilea, Salvinia, Azolla, etc.
Four particular types of habitats that pteridophytes are found in are (i) moist shady forests, (ii)
crevices in rock faces specially when sheltered from the full sun, (iii) acid wetlands including swamps
and bogs, and (iv) tropical trees bearing many epiphytic species.
In general, it may be said that ferns live in a wide variety of habitats, from remote mountain
elevations, to dry rock faces, to bodies of water or in open fields.

1.7 PLANT BODY

The plant body shows great variation in form, size and structure in different members (Fig. 1.1). It is
sporophytic and characterised by possession of roots, stem and leaves. Except a few woody tree ferns,
all living members are herbaceous. Rhizophore, an intermediate stage between root and stem, is present
in Selaginella. A young sporophyte is partially or completely dependent on the gametophyte for some
time, but at maturity it becomes independent.
The roots are generally adventitious. The primary roots are ephemeral. The stem, leaves and roots
develop with the help of a single two-sided or three-sided (Figs. 1.2 and 1.3 A and B) apical cell.
Bhambie’s (1957, 1960, 1970 and 1972) work on the development, structure and organisation of the
root and shoot apices of some Indian pteridophytes is worth mentioning. Bhambie and Puri (1963) have
worked on the shoot apex organisation of Lycopodiales, while Bhambie and Rao (1973) have worked
on the root apex organisation of Actinopteris radiata, Lygodium flexuosum, Microsorium alternifolium,
Nephrolepis exaltata, Osmunda regalis, Angiopteris erecta, Ophioglossum costatum, O. nudicaula,
Isoetes coromandelina and I. panchananii. Bir and Randhawa (1985) studied the apical meristems of
seven ferns and stated that, “always a single apical cell occupies the tip of each growing organ’, and
‘shoot apical cell invariably has three cutting faces in all ferns’.
The leaves may be simple, small and sessile as in Selaginella, Lycopodium, Equisetum, etc., or very
large and petiolate as in Filicinae. On the basis of the size of the leaves (smaller-sized microphylls
and large-sized megaphylls), pteridophytes may be grouped in two categories, i.e. microphyllous
and megaphyllous. Besides their smaller size, microphylls can be differentiated from megaphylls by
their extremely simple vascular system. Large-sized megaphyll of ferns branches many times and have
branching veins. On the other hand, the small-sized microphylls ‘rarely branch and possess either a
limited vascular supply or none at all’ (Sporne, 1975).
New leaves typically expand by unrolling of a tight spiral called a crozier or fiddlehead. This
uncurling of the leaf is termed circinate vernation.
The leaf trace, if present, is single and generally unbranched but sometimes it shows dichotomous
branching. In megaphylls, the leaf traces are usually associated with leaf gaps. Stomata are present
on various aerial parts of the plants including stem and leaves. Mehra and Soni (1983) studied the
stomatal patterns in 358 species belonging to 117 genera of pteridophytes, and designated four main
types, i.e. Psilophytaceous (Pattern I), Equisetaceous (Pattern II), Marattiaceous (Pattern III) and
 ® 1.5

(B)
(C)
(A) (D)

(E) (F)
(G)
1.6 ®
Filicinean (Pattern IV), According to them, the Psilophytaceous pattern was the basic pattern for all
pteridophytes.

Apex

Apical cell
Apical cell
Segment
Leaf apical cell Apical region Scale
Leaf

Subapical
region
Procambium
Leaf trace Leaf gap Maturation region
(A) (B)

The sporangia-bearing leaves are called sporophylls. In heterosporous forms, the sporophylls
are dfferentiated into microsporophyll or megasporophyll on the basis of presence or absence of
microsporangium or megasporangium. Some recent authors divide leaves of pteridophytes into the
following three types:
1. Trophophyll A leaf that does not produce spores, instead only produces sugars by
photosynthesis.
 ® 1.7

2. Sporophyll A leaf that produces spores.

3. Brophophyll A leaf that produces abnormally large amount of spores.

1.8 TROPHOPOD

Trophopod, a food-storage organ formed by the enlarged and modified leaf base, has been reported
in Asplenium platyneuron, Onoclea, Maheuccia and Dryopteris fragrans by Wagner and Johnson
(1983).

1.9 CAROTENOIDS

Czeczuga (1985) reported 27 carotenoids in 66 representatives of Pteridophyta. According to him,

the ‘carotenoids characteristic to clubmoss and horsetail species are b-carotene, b-cryptoxanthin,
lutein epoxide and zeaxanthin; and fern species are b-cryptoxanthin, lutein epoxide, violaxanthin and
rhodoxanthin’.

1.10 VASCULAR SYSTEM

A well-developed vascular system is present in all pteridophytes. Only because of the presence of
vascular structures pteridophytes are called vascular cryptogams. The vascular system consists of
xylem and phloem. As mentioned earlier, the leaf traces supplying microphylls are simple and cause
little disturbance to the stele. However, leaf traces supplying megaphylls are usually associated with
leaf gaps.
Cambium is generally absent. Sporne (1975) has, however, mentioned that “some pteridophytes
possess a vascular cambium from which secondary xylem and secondary phloem are formed. Cambial
cells possess the power of cell division.” .... “While relatively uncommon in living pteridophytes, a
vascular cambium was widely present in coal-age times, when many members of the group grew to the
dimensions of trees.”
A stele without leaf gaps is called a protostele. The stele in members like Lycopodium and Selaginella
is a simple protostele, but in Marsilea it is a siphonostele. In the aerial axis of Equisetum, the stele
is ectophloic siphonostele, whereas in Marsilea rhizome, it is amphiphloic siphonostele. However, in
genera like Pteris, Aspidium, Polypodium, Dryopteris, etc., the stele is of dictyostelic type.
The stelar system has been described in detail in Chapter 23.

1.11 SPORANGIA

Plants reproduce by the spores, which are formed in sporangia. The sporangia develop either on the
ventral surface or in the axils of the leaves. However, in Psilophytales (e.g. Rhynia, Horneophyton,
etc.) the sporangia were cauline.
The pteridophyta includes both homosporous (i.e. all the spores of one type, e.g. Equisetum) and
heterosporous (having two different types of spores, i.e. microspores and macrospores, e.g. Selaginella)
1.8 ®
forms. In homosporous members, exosporic gametophytes are present. In such gametophytes, spore
germination and gametophyte development are controlled by external conditions.
In heterosporous members, endosporic gametophytes are present. In such gametophytes, spore
germination and gametophyte development are independent of external conditions. These processes
take place in darkness in such gametophytes. Heterospory is the unique evolutionary feature of
pteridophytes.
In different genera, the sporangia are either present in a restricted area or throughout the vegetative
portion. In Equisetum and Selaginella they are present in the form of compact structures called strobili
or cones.
In genera, such as Azolla, Marsilea and Salvinia, the sporangia are present in specialised bodies
called sporocarps.
According to Bower (1935), each sporangium contains either 64 or less spores in leptosporangiate
ferns. If the number is less, it may be 24 or 48. On the other hand, the ‘greatest spore output occurs in
the microsporangia of Isoetes, where it is estimated that from 150,000 to 1,000,000 spores may develop
in a single microsporangium’(Foster and Gifford, 1959).

1.12 DEVELOPMENT OF SPORANGIA

Goebel (1881) classified sporangia into two types, i.e. eusporangiate and leptosporangiate, mainly
on the basis of their development.
In the eusporangiate type (Fig. 1.4), the sporangium originates from a group of superfical cells that
divide periclinally into outer primary wall cells and inner primary sporogenous cells. The primary-wall
cells develop into the several-layered wall of the sporangium, the innermost of which is the tapetum.
The sporogenous cells divide meiotically and give rise to haploid spores. Such a sporangium is called
a eusporangium. The wall of the mature eusporangium is single-layered and all its other layers
degenerate at maturity, e.g. Lycopodium, Selaginella, Equisetum, etc. But in Psilotum and Tmesipteris,
the mature sporangium retains its multilayered wall.

Spore
Primary Sporangium Tapetum Sporangium tetrads
Sporophyll
wall cells wall wall
Meiosis

Sporangial Primary
initials sporogenous Sporocytes
cells
(A) (B) (C) (D) (E) (F)

In the leptosporangiate type (Fig. 1.5), the sporangium develops only from a single superficial cell.
It divides transversely into an outer and an inner cell. The entire sporangium develops from the outer
 ® 1.9

cell. The inner cell is responsible for the formation of the stalk of the sporangium. The outer cell divides
by three successive divisions, and thus a tetrahedral apical cell is formed. The cells are cut off from
this apical cell on its three lateral faces. Thus formed upper segments form the wall of the sporangium
while the lower segments contribute to the three-rowed sporangial stalk. Finally, a periclinal division
takes place in the apical cell and thus form the outer jacket cell and the inner primary sporogenous cell.
The outer jacket cell divides by anticlinal divisions and forms the upper half of the sporangium wall
that is single-layered. The primary sporogenous cell divides into tapetal initial and sporogenous tissue.
The tapetal initial forms the two-layered tapetum (Fig. 1.5 E–F), while the sporogenous tissue divides
meiotically to give rise to haploid spores.

Primary
Tapetal
sporogenous initial
Apical cell Jacket cell cell
Sporangial
Sporangial initial Outer cell Inner cell Young
wall
stalk
cell

(A) (B) (C) (D) (E)

Primary
Annulus
sporogenous
cells
Spore tetrads
Tapetum
Sporangial wall Meiosis
Stalk
Stalk

(F) (G) (H)

A summary of the eusporangiate and leptosporangiate type of sporangial development is mentioned

below:

(1) Eusporangiate Type:

Outer primary Single-layered
wall cells sporangial wall
and tapetum

Group of superficial
initial cells

Inner primary Haploid

sporogenous spores (x)
cells (2x) Meiosis
1.10 ®
(2) Leptosporangiate Type:
Divides by three
Outer cell successive
divisions

Single superficial
initial cell

Inner cell (either it remains

inactive or divides to form
the stalk of the sporangium)

Tetrahedral Outer Divides anticlinally into

apical cell jacket single-layered upper half
cell of the wall of sporangium
Tapetal Two-layered
Inner initials tapetum
primary
sporogenous Sporogenous Haploid
First develops cell Meiosis
tissue (2x) spores (x)
into lower part of
capsule wall and
upper part of stalk

According to Williams (1928), the stalk development of the sporangia in Osmundaceae neither fits
in the eusporangiate type nor in the leptosporangiate type. Foster and Gifford (1959) have mentioned
that in Osmundaceae “the form of the intial cell from which the sporogenous cell, tapetum and wall
originate is variable” (Fig. 1.6 A–C). The stalk is sometimes truncated at the base, like eusporangium
(Fig. 1.6A) while at times it is pointed at the base, like leptosporangium (Fig. 1.6 B). Further, in
such sporangia, it cannot be traced easily whether the complete sporangium has developed from a
single cell.

1.13 SORUS

The sporangia in higher ferns are present in the form of well-organised groups called sori (singular
sorus). A sorus in which all the sporangia appear, grow and mature at the same time is called a simple
sorus (Fig. 1.7 A).
A sorus in which the centre is occupied by the oldest sporangium and the successive younger
sporangia are present towards the base is called a gradate sorus (Fig. 1.7 B).
But sometimes, sporangia of different ages are present in a sorus, without any definite arrangement.
Such a sorus is called a mixed sorus (Fig. 1.7 C).

1.14 SPORE AND ITS GERMINATION

Spores are haploid and form after reduction division in the sporogenous cells of the sporangium.
The form of the spore is variable. Often various types of wall configurations are present in mature
fern spores. The spore form and wall ornamentations sometimes serve as points of taxonomic
differentiation. The spore germinates into a multicellular gametophytic body called the prothallus.
 ® 1.11

Sporangial wall

Sporogenous cells

Primary Tapetum
sporogenous cells

(A) (B) (C)

Oldest sporangium Receptacle

Youngest
sporangium
Sporangia

Germination of spore is a two-phase process, i.e. spore distension and spore extension. In the
former process, spore swelling takes place; while in the latter process, the formation of germ tube and
germ rhizoid take place. Spore germination in pteridophytes has been studied by Mohr (1963), Sugai
(1971) and Whittier (1973).
Nayar and Kaur (1968) have classified spore germination of homosporous pteridophytes into
three categories, i.e. bipolar as in Lycopodium (Fig. 1.8 A–D), Equisetum (Fig. 1.8 E–G),Osmunda,
Anemia, Cyathea, Gleichenia, etc.; tripolar as in Trichomanes (Fig. 1.8 H and I); Hymenophyllum
and Macodium, etc. and; amorphous as in Angiopteris (Fig. 1.8 J and K). In the amorphous type of
germination, irregular cell divisions take place, and thus the ultimate result is growth in the irregular
direction.

1.15 PROTHALLUS

The spore germinates into a prothallus. Generally, the prothalli are green, simple, somewhat branched
and aerial structures. But in some genera, such as Lycopodium, they are subterranean, well-branched,
tuberous, colourless and saprophytic structures. Two sex organs, i.e. antheridia and archegonia,
develop on the prothallus. Generally, the prothalli of homosporous pteridophytes are monoecious and
protandrous. But in genera such as Equisetum they are protogynous.
In both homosporous and heterosporous pteridophytes, generally the archegonium remains
embedded in the prothallus. On the other hand the antheridium does not have a consistent structure.
1.12 ®

(A) (B) (C) (D)

Bipolar

(E) (F) (G)

Tripolar
(H) (I)

Amorphous

(J) (K)

In some members the antheridium is embedded, e.g. Lycopodium, Selaginella, Isoetes, Equisetum,
Ophioglossaceae, Marattiacease and heterosporous ferns; while in others they are emergent or projected
out, e.g. Psilotum, Tmesipteris and leptosporangiate ferns.
The antheridium is always surrounded by a well-defined jacket. In projected antheridia, the jacket is
always one-cell thick. Only one opercular cell is present. The antherozoids are unicellular, uninucleate
and biciliate structures in Lycopodium, Selaginella, etc., but they are multiciliate in Psilotales (Psilotum
and Tmesipteris), Isoetes, Equisetum and ferns. The number of antherozoids varies from four (Isoetes)
to few thousands (Ophioglossum).
The archegonium consists of a projecting neck and a lower embedded portion, the venter. The
neck region encloses neck canal cells, and the venter region encloses ventral canal cells and an egg.
Many neck canal cells are present in Lycopodium; while in leptosporangiate ferns only one binucleate
neck canal cell is present.

1.16 FERTILISATION AND ZYGOTE FORMATION

Fertilisation takes place with the help of water and results into the formation of a diploid zygote.
Antherozoids are attracted chemotactically towards the archegonium. A chemical substance comes
out from the open archegonium, most likely from the egg. This substance, which functions as a sperm
attracter, contains a large number of organic and inorganic compounds, especially malic acid and
fumaric acid.
The zygote is the first cell of the sporophyte and it is diploid. It develops into a well-developed
sporophyte bearing roots, stem and leaves.
Yamanouchi (1908) and Masuyama (1972) have made a detailed study of fertilisation in
pteridophytes.
 ® 1.13

1.17 LIFE CYCLE AND ALTERNATION OF GENERATIONS

The life cycle of a typical pteridophyte may be summarised as under:

A diploid sporophytic phase produces haploid spores by meiosis. The haploid spore grows by mitosis
into a gametophyte, which is typically a photosynthetic prothallus. The gametophyte produces gametes
(sperms and eggs) by mitosis. A mobile flagellate sperm fertilises an egg that remains attached to the
prothallus. The fertilised egg is now a diploid zygote and grows by mitosis into a sporophytic plant.
Pteridophytes show alternation of generations. In homosporous pteridophytes (Fig. 1.9 A), the
flagellated antherozoids are liberated from the antheridia. The antherozoids remain swimming in water.
The egg cell in the archegonium is non-motile. The fusion between the antherozoid (X) and egg cell
(X) results into the formation of a diploid zygote (2X). The zygote develops into a sporophytic plant
body, which is also diploid. At the time of spore formation in the sporophyte, the reduction of nuclear
contents takes place by meiosis, and thus haploid spores are formed. The spore germinates into a
gametophyte that again bears sex organs (Fig. 1.9 A).

X
Gametophyte

Egg X X Antherozoid

Spores

2X 2X

Zygote Sporophyte Meiosis

(A)

X
Gametophyte

X
X Gametophyte
X
Egg
Antherozoid
Meiosis

Microspores
2X
2X
Zygote Megaspores

Sporophyte Meiosis
(B)
1.14 ®
On the other hand, the general life cycle of a heterosporous pteridophyte (Fig. 1.9 B) shows the
formation of two types of spores, i.e. microspores and megaspores. These spores are formed after
meiosis in sporogenous cells present in the sporangia on the diploid (2X) sporophytic plant body.
The microspores and megaspores are thus haploid structures. The microspore develops into the male
gametophyte (X) that bears the antheridia having male gametes or antherozoids. The megaspore
develops into the female gametophyte (X) that bears the archegonium having a single female gamete
(X) or egg. During the process of fertilisation, fusion takes place between male and female gametes and
the result is the diploid zygote (2X) that directly germinates into a sporophytic plant body (2X).

1.18 RESEMBLANCES OF PTERIDOPHYTES WITH BRYOPHYTES

1. Members of both the groups show heteromorphic alternation of generations.

2. Antheridia and archegonia are present in both.
3. Sex organs are jacketed and multicellular in both.
4. Motile male gamete and nonmotile female gamete are present in both.
5. Absence of asexual spores formed by the ordinary division of mitosis is shown by both the
groups.
6. Members of both bryophyta and pteridophyta show oogamous type of sexual reproduction.
7. In all the bryophytes and some pteridophytes (Rhynia, Psilotum), rhizoids are present instead
of roots.
8. All the bryophytes as well as a large number of pteridophytes possess only one type of spores
showing homosporous condition.
9. Fertilisation in both bryophytes and pteridophytes takes place in the presence of water.

1.19 DIFFERENCES BETWEEN PTERIDOPHYTES AND BRYOPHYTES

1. Plant body in pteridophytes is sporophytic, whereas it is gametophytic in bryophytes.

2. Plant body in pteridophytes is differentiated into roots, stem and leaves, whereas it is thallus-
like in bryophytes.
3. Vascular tissues are absent in bryophytes while present in pteridophytes.
4. Roots are absent in bryophytes while present in pteridophytes.
5. Bryophytes, in general, are much smaller in size than pteridophytes.
6. Vegetative reproduction is more prevalent in bryophytes than pteridophytes.
7. Bryophytes are strictly homosporous whereas many pteridophytes are heterosporous.

1.20 RESEMBLANCES OF PTERIDOPHYTES WITH GYMNOSPERMS

1. Plant body in both the groups is sporophytic, and the sporophyte is differentiated into roots,
stem and leaves.
2. A regular alternation of gametophytic and sporophytic generations is seen in both.
 ® 1.15

3. In both the groups young leaves show circinate vernation.

4. Vascular elements are present in both groups.
5. Vessels are absent in a majorty of pteridophytes as well as gymnosperms (except Gnetales).
However, Bierhorst (1971) has mentioned that true vessels occur in vegetative parts of
Pteridium, Marsilea, Equisetum and Selaginella. Parkash and Bhambie (1980) also reported
true vessels in Helminthostachys.
6. Phloem lacks companion cells in both.
7. The gametophytic generation, in both the groups, is dependent upon the sporophytic
generation.
8. There is permanent retention of megaspores within the megasporangium in both groups.
9. ‘Ferns with seeds’ (Pteridosperms) was the common name given to the Cycadofilicales because
of their resemblances with ferns.

1.21 DIFFERENCES BETWEEN PTERIDOPHYTES AND GYMNOSPERMS

1. Tap roots are present in gymnosperms while roots are adventitious in pteridophytes.
2. Pteridophytes are hygrophytes, i.e. grow in shady humid places, while gymnosperms show
xerophytic characters.
3. Cambium is absent in pteridophytes while present in gymnosperms.
4. Neck canal cells and ventral canal cells are present in the archegonia of pteridophytes, while
they are generally absent in gymnosperms.
5. The microspores and megaspores develop independently after being shed from their sporangia
in pteridophytes, whereas in gymnosperms the microspores are shed only for a short time and
the megaspore is permanently retained within the megasporangium.
6. Due to the permanent retention in ovule, the megaspores in gymnosperms form the seeds,
while no seed formation is observed in pteridophytes.
7. Pollen-tube formation is observed in gymnosperms, while it is absent in pteridophytes.

1.22 FOSSIL STUDY

1.22.1 What is a Fossil?

The word ‘fossil’ is derived from the Latin verb fodere, which means ‘to dig’. Fossils may be precisely
defined as ‘the remains of ancient plants and animals preserved in rocks or different layers of the earth’.
Sporne (1975) defined a fossil as “anything which gives evidence that an organism once lived”. The
branch of science in which we study the plants of past geological ages through the investigation of
fossils is called palaeobotany.

1.22.2 How are Fossils Formed?

Normally, the death of a plant occurs due to decomposition by bacteria, fungi or other microorganisms,
or because of the eating up by insects or other animals. But sometimes it happens that the whole
1.16 ®
organism is buried under earth, either alive or soon after its death. In such cases the decomposition
proceeds under such conditions that check or retard the activity of decaying and allows such changes
to occur which ultimately result in the preservation of the parts of the plant body. Such preserved parts
represent fossils.

1.22.3 Types of Fossils

Some common types of fossils include the following:
1. Petrifactions These are formed because of the slow infiltration of tissues by some minerals
like calcium, silica or magnesium. In such fossils, the organic matter is replaced by mineral
matter. In petrifactions, the tissues are so well-preserved by mineral substances that almost all
the details of even cell walls are visible under the microscope.
2. Compressions or Mummifications These are the fossils in which the parts of the organism are
flattened by the vertical pressure of the overlying rocks. Therefore, a lens-shaped fossil would
be formed if the original organism was spherical.
In the presence of some anaerobic conditions, e.g. in marine mud, plant tissues slowly turn
to coal in which no structures can be recognised except spores or cuticles of leaves. These
portions of the plant that can be separated from each other by mud or sand during deposition
give rise to fossils called compressions or mummifications. Outlines of epidermal cells, hair,
stomata, etc, can be studied by these fossils.
3. Impressions In these fossils the material of the plant disorganises but the ‘impression’
remains. These are formed because of the burial of the parts of the plant in the soil, which later
on harden into rock. Only external features of the plant are preserved in such fossils.
4. Casts or Molds These are formed when an organism, submerged in water containing lime,
remains covered by a crust of mineral matter. The decay of such incrusted organisms leaves a
cast or mold. Casts actually exhibit nothing of the original tissues of the organisms. They are,
however, valuable in showing their shape.
Petrifactions, impressions, compressions and cast or mold can also be summarised as under:
Petrifactions are minerally impregnated plant parts, showing internal details of the cell. There is no
decay of materials in these fossils, and the infiltrating substance forms crystals and hardens in the cell
lumen resulting in the formation of a hard matrix. Silica, iron hydroxide and carbonates of calcium and
magnesium are common petrifying materials.
In compressions, parts of the plant get covered with sediment that makes the object flattened due to
weight. No internal cellular details are preserved in these fossils.
An impression is the negative of compression. On being split open, these fossils show a negative
imprint.
 ® 1.17

Plant remains decompose after compression and ultimately a space is left behind. This space may
get filled with another sediment, and in such a condition the space is called a mold. Simultaneously,
the sediment, which acquires the form of the object in a mold, is called cast.
Differing from petrifactions, no cellular details are preserved in compression, impression, cast and
mold.

1.22.4 Geological Time Scale

The geological history of the earth has been separated into five geological eras by geologists. Each
of the eras is divided into periods, and each period into smaller subperiods called epochs. Geologists
believe that the oldest rocks of the earth’s crust are about 2 billion (2003 million) years old. Cenozoic
is the recent era, of which our present time is also a part. Archeozoic is the oldest of the geological eras.
The different eras, along with their approximately estimated duration in years, are as follows:
Era Duration in years
1. Cenozoic 60,000,000
2. Mesozoic 125,000,000
3. Paleozoic 368,000,000
4. Proterozoic 900,000,000
5. Archeozoic 550,000,000+
2003,000,000 years
A method of preparing a geological time scale is based on radioactivity of isotopes* of certain
elements, e.g. uranium. Uranium (238U) disintegrates into lead (206Pb) in 4.5 billion years. If some lead
has been formed in a sample of uranium ore, then the ratio of 206Pb/238U related to half-life** of 238U is
the measure of this time.
Another method used for dating recent strata is the 14C method. In this method, the death of an
organism is estimated by the ratio of 14C in a fossil sample to the amount of 14C in a living tissue.

1.22.5 Fossil History of Pteridophytes

Pteridophytes, an ancient group of plants, contain a long fossil history as evidenced from Table 1.1.
Although the question as to when did pteridophytes first appear on the earth has been a debatable
subject but the characters like (i) presence of cutinised spores, (ii) a waxy cuticle with stomata, and (iii)
a vascular system containing lignified xylem element, suggest that these plants were first recognised
in the Silurian period of the Middle Paleozoic age (about 381 million years ago) and flourished well
between the Devonian (about 354 million years back) and Upper Carboniferous (about 271 million
years ago) periods of the Paleozoic era. Psilopsida, Lycopsida, Sphenopsida and many ferns flourished
well during the Devonian period and developed during the Lower Carboniferous (about 309 million

*
Isotopes: Any of the two or more forms of the same element having the same atomic number and nearly the same chemical
properties but with different atomic weights.
**
Half-life: The length of time in which one half of the radioactive atoms present in a substance will decay.
 1.18
S. No. Era with duration Major divisions Period and number of Epoch Important events in Dominant organisms

®
in years years from the present plant life of the era
time

1. CENOZOIC Quaternary 2,000,000 years Recent Dominance of Age of herbs

(60,000,000 years) herbaceous plants and man
Pleistocene Speciation of
herbaceous plants
Tertiary Late Tertiary Pliocene Spreading of Angiosperms, birds
herbaceous dicots
Miocene Restriction in plant Mammals
distribution; Reduction
in forests; Rise of
herbaceous angiosperms

Early Tertiary Oligocene Dispersal of woody

(60,000,000 years angiosperms
from the present)
Eocene Development of flowering
plants; Extensive forests
Paleocene Modernisation of
angiosperm families

2. MESOZOIC
(125,000,000 years) Late
Mesozoic Upper Cretaceous Dominance of
angiosperms;
Dwindling of
gymnosperms
Middle Cretaceous Rapid development Higher gymnosperms
of angiosperms; Many and reptiles
genera are present
even today
Lower Cretaceous Rise and development
of cycads, conifers
and angiosperms
Contd.
 Early Jurassic Origin of angiosperms;
Mesozoic Dominance of Cycadales
and Coniferales; Definite
evidence of diatoms
Triassic Conifers and Ginkgoales
(185,000,000 years increased; Disappearance
from the present) of seed ferns; Rise of
cycads

3. PALEOZOIC
(368,000,000 years) Late Permian Rise of conifers; Lycopods, seed
Paleozoic (223,000,000 years Extinction of coal ferns and
from the present) swamp flora amphibians

Upper Dominance of ferns,

Carboniferous Calamitales, Lepidoden-
(271,000,000 drales, Cordaitales
years from (gymnosperms); Extensive
the present) coal formations
Lower Carboniferous Development of Lycopsida,
(309,000,000 years Calamitales, seed ferns;
from the present) Early coal deposits
Middle Paleozoic Devonian Flourishing Psilopsida, Fishes and early
(354,000,000 years Lycopsida, Sphenopsida, land plants
from the present) ferns, Bryophytes, some
algae
Silurian (381,000,000 First-known land plants; Algae, higher
years from the present) Dominance of algae; invertebrates
Cooksonia, the oldest
known vascular plant
Early Paleozoic Ordovician (448,000,000 Marine red and green
years from the present) algae; Perhaps the time
of rise of land plants
Cambrian (553,000,000 Evidence of algal origin
®

years from the present)

Contd.
1.19
4. PROTEROZOIC 1,500,000,000 years Algae and bacteria Marine invertebrates

1.20
(900,000,000 years) from the present

5. ARCHEOZOIC
(550,000,000 years +)
2,000,000,000 years
from the present
Probably very simple,
unicellular organisms.
Unicellular life (?) ®
Origin of prokaryotic cell?
Origin of the earth.
 ® 1.21

years ago) period. There was a complete dominance of ferns, Calamitales and Lepidodendrales during
the Upper Carboniferous (about 271 million years ago) period. Because of this the Late Paleozoic era
can be easily regarded as the age of pteridophytes.

1.23 HOW TO EXAMINE A FOSSIL?

Fossils can be examined by preparing a peel by the following method:

First of all, a polished surface of the specimen is prepared, which is then treated for some time with
dilute hydrochloric acid. In case of a siliceous matrix, hydrofluoric acid is used in place of hydrochloric
acid. Treatment by the acid removes the mineral matrix and due to this, cell walls of the tissue get
slightly elevated over the rock surface. The used acid is removed soon and the surface of the specimen
is flooded with acetone. A thin sheet of cellulose acetate is now laid on the specimen surface. On being
dried, this sheet, which contains a thin preparation of fossil, is peeled off. The peel is mounted on a
clean glass slide and studied under microscope.

TEST YOUR UNDERSTANDING

1. Give a suitable definition of pteridophyta.

2. What are vascular cryptogams?
3. In the traditional system of plant classification, what are included under Cryptogams?
4. The so-called Lower Cryptogams include algae, fungi and ________.
5. The so-called higher cryptogams include ____________.
6. In the plant kingdom, pteridophytes occupy a position in between bryophytes and _______.
7. Write any six general characteristics of pteridophytes.
8. Two categories in which pteridophytes may be grouped on the basis of size of leaves, are ____
and __________.
9. Define the following mentioning one sentence for each:
(a) Circinate vernation (b) Trophopod
(c) Brophophyll (d) Trophophyll
10. In pteridophytes, is the cambium generally present or absent?
11. A stele without leaf gaps is called __________.
12. Differentiate between the following, mentioning only one sentence about each:
(a) Homosporous and heterosporous
(b) Strobilus and sporocarp
(c) Sporangium and sorus
(d) Eusporangiate and leptosporangiate
(e) Exosporic gametophyte and endosporic gametophyte
13. Give a detailed illustrated account of development of eusporangiate and leptosporangiate type
of sporangia.
1.22 ®
14. In Isoetes, a single microsporangium may contain as many as 1,50,000 to __________
spores.
15. How can you diagrammatically represent the following?
(a) Simple sorus (b) Gradate sorus (c) Mixed sorus
16. Define the following terms:
(a) Tapetum (b) Protandrous
(c) Prothallus (d) Protogynous
17. Describe the patterns of spore germination in homosporous ferns.
18. Describe in brief the life cycle of a typical pteridophyte with reference to alternation of
generations.
19. Write at least three points of differentiation between:
(a) Bryophytes and pteridophytes
(b) Pteridophytes and gymnosperms
20. Define briefly:
(a) Petrifactions (b) Compressions (c) Impressions
21. With reference to geological time scale, each geological era is divided into _______
22. The Late Paleozoic era is regarded as the ‘Age of Pteridophytes’. Comment.
 CHAPTER 2
Development of
Pteridology in
India

Even a brief mention of all researches and contributions related to the development of pteridology in
India is neither possible to provide in a book of this nature, nor is it required for its prospective readers.
Therefore, only some selected developmental aspects are enlisted here in this chapter.

2.1 MAJOR DEVELOPMENTS PRIOR TO 1900 AD

1. Linnaeus (1753) described many Indian pteridophytes in his Species Plantarum.

2. ‘Genera et Species Filicum’ by Swartz (1806) contains a detailed discussion of several Indian
ferns.
3. Hooker and Greville (1827–1831) and Hooker (1846–1864) described a number of Himalayan
ferns.
4. A detailed treatment of Indian ferns was published by Beddome (1863) in his two well-known
books entitled Ferns of Southern India and Handbook of Ferns of British India.
5. Several ferns were described by Clarke (1880) in his book Ferns of Northern India.
6. C W Hope (1899–1902) published a detailed account of Indian ferns in his Ferns of North-
Western India.

2.2 DEVELOPMENT OF PTERIDOLOGY IN THE 20TH CENTURY

1. The credit of publishing the first Indian research paper on Equisetum goes to Shiv Ram Kashyap
(1914).
2. Soon, Birbal Sahni began studies on Nephrolepis at Lucknow University around 1915. In
1925, Professor Sahni studied Psilotales, in general, and Tmesipteris, in particular.
2.2 ®
3. Chowdhary (1937) and Mahabale (1937, 1948) made significant contributions on
Lycopodium.
4. Ninan (1950) studied chromosome number in Psilotum and also made valuable contributions
on Lycopodium in 1958.
5. S Bhambie (1957, 1960, 1970, 1972) along with C G P Rao and V Puri published a series
of papers on the development, structure and organisation of root and shoot apices of many
Indian pteridophytes and made worthy contributions on Isoetes, Lygodium, Lycopodiales,
Nephrolepis, Osmunda, Angiopteris and Ophioglossum.
6. Panigrahi (1958), and Panigrahi and Chowdhary (1962) made extensive studies on the genus
Selaginella.
7. T S Mahabale (1933–1972) of Pune Vidyapeeth made extensive collections of Indian
pteridophytes and studied gametophytes of several ferns.
8. P N Mehra and D S Loyal (1956, 1959) studied effects of colchicine on the prothalli of
ferns and cytological details of Marsilea. Mehra also studied the impact of cytology on the
phylogeny of ferns. Morphological, anatomical, taxonomical and palynological studies of
several pteridophytes was made during 1936 and 1976 by Mehra and co-workers, such as
D S Loyal (1956, 1958, 1959, 1965), S S Bir (1957, 1958, 1960), S C Verma (1957, 1960) and
H K Palta (1971).
9. A monographic account of Marsilea was published by K M Gupta (1962). The morphological
nature of sporocarp of Marsilea was studied in detail by V Puri and M L Garg (1953).
10. Genera of Ophioglossaceae, including Botrychium, have been investigated by L N Rao
(1962).
11. K R Surange (1966) gave an excellent account of fossil pteridophytes of India.
12. During 1962 and 1984, D D Pant and co-workers (Bharti Mehra,1964; P F Kidwai, 1968;
P K Khare, 1974; D D Mishra, 1976; L Mishra, 1976; D D Nautyal, 1984; D R Mishra, 1984,
etc.) made valuable contributions on Indian pteridophytes, such as Lycopodium, Isoetes, etc.
13. S S Bir made valuable contributions on the taxonomy and cytogenetics of Indian pteridophytes
during 1954–1987 along with several of his co-workers.
14. B K Nayar also made significant contributions on Indian pteridology during 1954 and
1975 along with several co-workers including S Kaur (1969, 1971), P Chandra (1963), N
Bajpai (1964), S Chandra (1965) and others. Nayar developed a strong centre of research in
pteridology at NBRI (National Botanical Research Institute), Lucknow. He also worked at
Gauhati University and Calicut University. Dr Nayar concentrated his researches mainly on
the morphology of gametophytes, palynology, taxonomy and anatomy of Indian pteridophytes.
A culture laboratory was also established by Nayar at NBRI. Nayar also focussed his researches
on evolutionary trends of gametophytic generation of pteridophytes. He also worked on the
ecology, spread and eradication of Salvinia molesta, a freshwater pteridophytic weed.
15. G Panigrahi conducted researches on taxonomy and phylogeny of pteridophytes during
1962–2002.
16. H K Cheema (1994) focussed on morphogenetic studies on pteridophytes in India.
17. B D Sharma (1994) presented a detailed account of pteridophytes in his article ‘Pteridophyta:
Morphology, Anatomy and Reproductive Biology’.
18. Subhash Chandra (1996) of National Botanical Research Institute, Lucknow, published
excellent accounts of various aspects of Indian pteridophytes, including (i) stelar morphology,
 ® 2.3

(ii) taxonomy, and (iii) nomenclature of Indian ferns. Articles of reputed pteridologists were
edited and published by Dr Subhash Chandra (2002) in the book Pteridology of the New
Millennium.

2.3 DEVELOPMENT OF PTERIDOLOGY IN THE 21ST CENTURY

1. In the year 2002, excellent articles were published by A Kumari and R B Srivastava on
‘Pteridophytes of North Bihar’, S C Verma on Reproductive Biology of Gametophytic
Generation of Homosporous Ferns’, Shukla and Srivastava on ‘Isoetes’, and P B Khare on
‘Pteris vittata’ and ‘Threatened Pteridophytic Taxa’ of the country.
2. Pravin Chandra Trivedi (2002) edited ‘Advances in Pteridology’, which discusses several
major aspects of pteridophytes in India including (i) systematics and classification of ferns
by S M Vasudeva, (ii) gametophytic reproductive biology of homosporous ferns by S C
Verma, (iii) researches on Marsilea by F D’Souza, (iv) researches on Isoetes by Dilip Gena,
(v) biodiversity of vascular cryptogams by P C Pande and H C Pande, (vi) morphoanatomical
studies of Selaginella by R Mukhopadhyay, (vii) Ophioglossum in Rajasthan by B L Yadav
and M K Tripathi, (viii) ethnomedicinal uses of pteridophytes by P C Trivedi, and (x) Cyathea
by N Punetha and J N Pant.
3. In 2008, a detailed article on the ‘Threatened Pteridophytes of India’ was published by S
Chandra, C R Fraser-Jenkins, A Kumari and A Srivastava in Taiwania (2008). They have
enlisted 414 species of pteridophytes as ‘threatened’, which include 219 species ‘at risk’, 82
species as ‘near threatened’ and 113 species as ‘rare’.
4. Recently, in 2009, S D Rout, T Panda and N Mishra published their ‘ethnomedicinal studies’
of 23 Indian species belonging to 21 families of pteridophytes, with particular emphasis
on Adiantum, Asplenium, Lygodium and Pteris. Their research has been published in the
International Journal of Medicine and Medical Sciences.
5. G Perumal (2010) studied ethnomedicinal uses of some pteridophytes available in some parts
of Tamil Nadu, and their research has been published in Ethnobotanical Leaflets. Perumal has
specifically studied medicinal importance of some species of Adiantum, Angiopteris, Drynaria,
Dryopteris, Marsilea, Pteridium and Selaginella.

TEST YOUR UNDERSTANDING

1. Give at least ten major steps of the development of pteridology in India in chronological
order.
2. Describe in brief the major events of the development of pteridology in our country.
3. Name the Indian botanist who was the first to publish a research paper in pteridology.
4. The first authentic books on Indian ferns were published by Beddome (1863). Name them.
5. Give an account of pteridological development in India in the 20th century.
6. Name the place where the following pteridologists worked in India.
(a) S Bhambie (b) T S Mahabale (c) D D Pant
(d) B K Nayar (e) Subhash Chandra
 CHAPTER 3

Threatened
Pteridophytes
of India: Handle
Them with Care

3.1 WHAT ARE THREATENED PTERIDOPHYTES?

Threatened species* of any group include any indigenous species of taxonomic significance, which
are said to be either ‘at risk’ or ‘near threatened’ or ‘rare’. According to Chandra et al. (2008), 414
species of pteridophytes fall under the threatened status out of a total number of 950–1000 species of
pteridophytes reported from political India. Thus, the threatened pteridophytes constitute 41–43% of
the total number of pteridophytes in the country. The 414 species of threatened pteridophytes include
219 at risk, 82 near-threatened and 113 rare (Chandra et al., 2008). Out of the 219 species included ‘at
risk’, 160 are ‘critically endangered’.
To protect our environment, we should handle these species of threatened pteridophytes with utmost
care. With increased utilisation of land and natural resources in the recent past, it is feared that several
of these threatened taxa may soon become yet rarer, more vulnerable and endangered, and in some
cases may finally become extinct (i.e. species which no longer exist).

3.2 HOW TO FIND DETAILS OF EXTINCTION OF SPECIES?

The major and widely accepted international authority, which provides details of the international
concern on the loss or extinction of species is the International Union for the Conservation of Nature
and Natural Resources (IUCN). Its original Red Data Book was first published in 1966 and now revised

*
For details, Consult Chandra, S. et al. (2008), Taiwania 53(2): 170–209.
3.2 ®
annually under the name IUCN Red List. It became electronic since 2000, and its details can be found
on the following websites.
(i) www.iucn.org.
(ii) http://www.iucnredlist.org.
Major details of the threatened plants of India and of Indian Plant Red Data Book can be found in
Jain and Sastry (1980), Jain (1984) and Jain and Rao (1984).
The International Association of Pteridologists (IAP), using criteria established by IUCN, developed
a list of threatened ferns of the world. Their World Conservation Monitoring Centre, Cambridge
(England) listed 1650 threatened species of pteridophytes worldwide according to Jermy (1990).
Sharpe (2001), however, accepted only 770 threatened ferns and fern-allies worldwide.

3.3 MAJOR CATEGORIES OF THREATENED PTERIDOPHYTES

On the basis of the details mentioned in 2001 Red List of IUCN (revised in 2006), Chandra et al.,
(2008) mentioned the following nine categories of threatened plants:
1. Extinct (EX)—with no reasonable doubt after exhaustive surveys
2. Extinct in the Wild (EW)—only known in cultivation after exhaustive surveys
3. Critically Endangered (CR)—extremely high risk of extinction in the wild
4. Endangered (EN)—very high risk of extinction in the wild
5. Vulnerable (VU)—high risk of extinction in the wild
6. Near Threatened (NT)—likely to be threatened in the near future
7. Least Concern (LC)—widespread and abundant
8. Data Deficient (DD)—available information is inadequate to assess risk
9. Not Evaluated (NE)—not evaluated against the criteria
Details of these categories are also available on the website: http://www.iucnredlist.org/.
Except that of some developed countries, all these above-mentioned nine categories are not
appropriate for pteridophytes mainly because of lack of required information. In most countries, the
highly sophisticated and detailed system of local recorders feeding data into a Central database does
not exist. Therefore, in majority of the countries, the threatened status is given only to indigenous
species of taxonomic significance, which fall into one of the three categories of threat:
1. At Risk
2. Near-Threatened
3. Rare

3.3.1 At Risk
The species included under the ‘At Risk’ category should show any one of the following criteria:
1. The decline in species is considered to be rapid and their number and habitat have been reduced
drastically to a critical level.
2. A considerable fall in the reproductive capacity of the species is seen. This results in the
decline in the number and size of the population of this species.
 ® 3.3

3. The species is available only in three or even lesser known localities.

Chandra et al. (2008) included 219 species of pteridophytes under the ‘At Risk’ category in
India.

3.3.2 Near-Threatened
The species included under this category are those which are declining because of:
1. Over-exploitation,
2. Extensive habitat destruction, and
3. Major environmental disturbances.
The species included under the ‘Near-Threatened’ category are likely to be in danger if timely
modification to their habitat does not stop. Chandra et al. (2008) included 82 species of pteridophytes
under the ‘Near-Threatened’ category in India.

3.3.3 Rare
Species are called ‘Rare’ when they (i) occur in widely separated small sub-populations, which helps in
reducing or restricting interbreeding, or (ii) are thinly scattered over a more extensive range. According
to Chandra et al. (2008) there are at present 113 species of pteridophytes in India which fall under the
category of ‘Rare’.

HUMAN ACTIVITIES RESULTING INTO THREATENED

3.4 SPECIES OF PTERIDOPHYTES IN INDIA

A very large number of pteridophytes in India are threatened due to several human activities. Some
such activities are listed below:
1. Environmental degradation of preferred habitat for pteridophytic species by activities such as
(i) uncontrolled tree-felling for fuel, timber, etc., (ii) grazing, (iii) crop plantation, (iv) building
construction, and (v) draining of wetlands for agriculture, etc.
2. Environmental pollution of (i) roadsides, (ii) lakes, and (iii) banks of canals, rivers, etc.
3. Unregulated commercial collection of pteridophytic species, such as tree-fern trunks for
growing orchids, etc.
4. Regular unrequired collection of pteridophytic species, known to be rare, by students, research
scientists, botanists, etc.
5. Encroachment upon huge areas of natural vegetation by businesspersons and several other
organisations for personal gains.
All these activities are continuously resulting in the (i) overall decrease in forest cover, (ii) climate
change, such as decrease in rainfall, increase in temperature, floods, draughts, etc., and finally (iii)
global warming.
3.4 ®
3.5 INDIAN THREATENED PTERIDOPHYTES IN THE ÔAT-RISKÕ CATEGORY

Chandra et al. (2008) listed the following 219 pteridophytic species in the ‘At-Risk’ category:
1. Huperzia cancellata 2. H. carinata 3. H. quasipolytrichoides 4. H. nummulariifolia
5. Heterogonium pinnatum 6. H. vernicosa 7. Lycopodium annotinum 8. L. complanatum
9. L. dendroideum 10. Selaginella aitkisonii 11. S. cataractarum 12. S. megaphylla
13. S. ornithopodioides 14. S. pulvinata 15. S. waltii 16. Equisetum palustre17. Botrychium virginianum
18. Helminthostachys zeylanica 19. Ophioglossum gramineum 20. O.lusitanicum 21. O. pendulum
22. Christensenia aesculifolia 23. Marattia fraxinea 24. Osmunda cinnamomea 25. O. javanica
26. O. angustifolia 27. Plagiogyria glauca 28. Lygodium circinnatum 29. L. polystachyum
30. Schizaea dichotoma 31. S. digitata 32. Dicranopteris curranii 33. D. subpectinaty
34. Dipteris wallichii 35. Arthromeris notholaenoides 36. A. wardii 37. Drymotaenium miyoshianum
38. Drynaria bonii 39. D. parishii 40. Goniophlebium persicifolium 41. Lemmaphyllum micraphyllum
42. Leptochilus minor 43. L. subhemionitideus 44. L. thwaitesianus 45. Loxogramme grammitoides
46. Microgramma lycopodioides 47. Microsorum chinense 48. Neocheiropteris ensata 49. Phymatoperis
connexa 50. P. nigrovenia 51. P. tibetana 52. Phymatosorus longissimus 53. Platycerium wallichii
54. Polypodiodes dielsiana 55. P. manmeiense 56. P. wattii 57. Pyrrosia boothii 58. P. drakeana
59. P. laevis 60. P. longipilosa 61. P. rasamalae 62. P. subfurfuracea 63. Selliguea chrysotricha
64. S. majoensis 65. S. subparsa 66. S. tricuspis 67. Thylacopteris papillosa 68. Ctenopterella blechnoides
69. Oreogrammitis attenuata 70. O. austroindica 71. O. pilifera 72. Prosaptia alata 73. P. contigua
74. P. khasyana 75. P. obliquata 76. Scleroglossum sulcatum 77. Tomophyllum perplexum
78. Hymenophyllum acanthoides 79. H. barabatum 80. Trichomanes agasthianum 81. T. apiifolium
82. T. exiguum 83. T. graude 84. T. griffithii 85. T. henzaianum 86. T. javanicum 87. T. maximum
88. T. mindorense 89. T. motleyi 90. T. parvifolium 91. T. sublimbatum 92. Cibotium barometz
93. Cyathea albosetacea 94. C. andersoii 95. C. brunoniana 96. C. chinensis 97. C. contaminans
98. C. crinita 99. Dennstaedtia wilfordii 100. Microlepia calvescens 101. M. majuscula 102. Lindsaea
commixta 103. L. malabarica 104. L. vensusta 105. Taenitis blechnoides 106 Tapeindium pinnatum
107. Acrostichum speciosum 108. Aleuritopteris argentea 109. A. thwaitesii 110. Cheilanthes nitidula
111. Negripteris scioana 112. Notholanea delavayi 113. N. lanuginosa 114. N. dipinnata 115. Pellaea
calomelanos 116. P. longifolia 117. Pteris amoena 118. P. barbigera 119. P. geminata 120. P. griffithii
121. P. griffithii senju 122. P. mertensioides 123. P. pleuricaudata 124. P. tripartita 125. P. venulosa
126. Syngramma alismifolia 127. Adiantum soboliferum 128. A. stenochlawys 129. Asplenium affine
130. A. anogvammoides 131. A. batueuse 132. A. delavayi 133. A. exiguum 134. A. hymenophylloides
135. A. khasianum 136. A. pellucidum 137. A. scalare 138. A. serricula 139. A. sublase rpitiifolium
140. A. athunbergii 141. Thelypteris beddomei 142. T. confluens 143. T. cuspidata 144. T. didymoch
laeneides 145. T. kurzii 146. T. latebrosa 147. T. menisciicarpa 148. T. namburensis 149. T. paludosa
150. T. palustris 151. T. opulente 152. T. srilankensis 153. Athyrium atratum 154. A. cumingianum
155. A. niponicum 156. A. repens 157. A. roseum 158. Deparia macdonellii 159. Diplazium
crenatoserratum 160. D. cordifolium 161. D. griffithii 162. D. heterophlebium 163. D.
pinfaense 164. D. pinnatifidopinnatum 165. D. tomentosum 166. D. virescens 167. Diplazium
sp. 168. Gymnocarpium oyamense 169. Matteuccia orientalis 170. Woodsia cycloloba 171.
W. glabella 172. Acrorumohra diffracta 173. Ctenitis mannii 174. Cyrtomium fortunei
175. C. micropterum 176. Dryopsis ferruginea 177. D. angusttifrous 178. Dryopteris
assamensis 179. D. austroindica 180. D. assamensis 181. D. austroindica 182. D. basisora
 ® 3.5

183. D. costalisora 184. D. deparioides 185. D. dickinsii 186. D. hasseltii 187. D. himachalensis 188.
D. nobilis 189. D. camusiae 190. D. varia 191. Heterogonium pinnatum 192. Lastreopsis tenera 193.
Polystichum anomalum 194. P.duthiei 195. P. glaciale 196. P. grandifrous 197. P. manickamianum
198. P. polydon 199. P. subinerme 200. P. wattii 201. P. sp. indet 202. Pteridrys cnemidaria
203. P. syrmatica 204. P. zeylanica 205. Tectaria kehdingiana 206. T. subconfluens
207. T. zeilanica 208. Arthropteris palisotii 209. Oleandra undulata 210. Bolbitis nodiflora
211. B. situata 212. Elaphoglossum nilgiriosm 213. E. stigmatolepis 214. Lomagramma sumatrana
215. Davallia pectinata 216. D. heterophylla 217. Blechnum finlaysonianum 218. B. melanopus
219. B. melanocaulou.

INDIAN THREATENED PTERIDOPHYTES IN THE

3.6 ÔNEAR-THREATENEDÕ CATEGORY

Chandra et al. (2008) listed the following 82 species of pteridophytes in the ‘Near-Threatened’
category:
1. Lycopodium wightianum 2. Psilotum nudum 3. P. complanatum 4. Osmunda huegeliana
5. Plagiogyria adnata 6. Lygodium longifolium 7. Belvisia henryi 8. Goniophlebium mengtzeense
9. Lepisorus oligolepidus 10. L. sordidus 11. Leptochilus superficialis 12. Microsorum fortunei
13. M. zippelii 14. Pleopeitis macrocarpa 15. Polypodioides niponicum 16. P. subamoena
17. Pyrrosia ceylanica 18. Hymenophyllum denticulatum 19. H. edentulum 20. H. khasianum
21. H. levingei 22. H. simonsianum 23. Trichomanes christii 24. T. humile 25. T. latemarginate
26. T. nitidulum 27. Cyathea khasyana 28. C. nilgirensis 29. Microlepia trichocarpa
30. Lindsaea parasitica 31. L. repens 32. L. walkerae 33. Aleuritopteris duthiei 34. A. wollenweberi
35. Anogramma leptophylla 36. Pellaea boivinii 37. P. faleata 38. Pteris cadieri
39. P. subindivisa 40. P. tricolor 41. Adiantum poiretii 42. Antrophyrum henryi 43. Asplenium caudatum
44. A. grevillei 45. A. magnificum 46. A. nidus 47. A. rivulare 48. A. simonsianum 49. A. Tenerum
50. Thelypteris chandrae 51. T. elwesii 52. T. heterocarpa 53. T. hirsutipes 54. T. immersa 55. T. parishii
56. T. sbelata 57. T. thwaitesii 58. Athyrium nakanoi 59. A. kumaonicum 60. Diplazium beddomei
61. D. cognatum 62. D. crinitum 63. D. leptophyllum 64. D. muricatum 65. D. travancoricum
66. Woodsia alpina 67. W. lanosa 68. W. hancockii 69. W. rosthorniana 70. Arachniodes superba
71. Dryopsis scabrosa 72. Dryopteris khullarii 73. D. sledgei 74. D. yoroii 75. Tectaria chattagramica
76. T. dubia 77. T. ingeus 78. T. melanocaulos 79. T. sifolia 80. Elaphoglossum beddomei
81. Lomagramma sorbifolia 82. Brainea insignis.

3.7 INDIAN THREATENED PTERIDOPHYTES IN THE ÔRAREÕ CATEGORY

Chandra et al. (2008) listed the following 113 species of pteridophytes in the ‘Rare’ category.
1. Huperzia ceylanica 2. H. nilagirica 3. H. selago 4. H. phlegmaria 5. Lycopodium veitchii
6. L. casuarinoides 7. Selaginella adunca 8. S. miniatospora 9. S. picta 10. S. tenuifolia
11. Botrychium daucifolium 12. B. lunaria 13. B. ternatum 14. Ophioglossum nudicaule
15. O. parvifolium 16. O. polyphyllum 17. Osmunda regalis 18. Arthomeris tenvicauda
19. Leptochilus pothifolius 20. L. pedunculatus 21. Neocheiropteris ovata 22. Phymatopteris
crenatopinnata 23. P. erythrocarpa 24. Polypodiodes hendersonii 25. Pyrrosia nummulariifolia
3.6 ®
26. P. stenophylla 27. Selliguea rhynchophylla 28. Hymenophyllum gardneri 29. Trichomanes
bilabiatum 30. T. bimarginatum 31. T. bipunctatum 32. T. intramarginale 33. T. obscurum
34. T. saxifragoides 35. Cyathea holtumeana 36. C. nicobarica 37. Microlepia hallbergii 38. Lindsaea
bouillodii 39. L. gueriniana 40. L. javanensis 41. L. malayensis 42. L. oblanceolata 43. L. obtusa
44. L. teuera 45. L. tetragona 46. Anogramma reichsteinii 47. Cerosora microphylla 48.Cheilanthes
persica 49. Doryopteris lindens 50. Onychium tenuifrons 51. Pteris grevilleana 52. P. heteromorpha
53. P. multiaurita 54. P. quadriaurita 55. P. roseolilacina 56. P. vittata 57. Vittaria microlepis
58. Adiantum flabellulatum 59. A. wattii 60. Antrophyum callifolium 61. A. parvulum 62. Vittaria
ensiformis 63. V. zosterifolia 64. Asplenium aitchisonii 65. A. apogamum 66. A. auritum 67. A. nesii
68. A. pekinense 69. A. prolongatum 70. A. rockii 71. A. viride 72. Thelypteris griffithii 73. T. himalaica
74. T. hirtisora 75. T. phegopteris 76. T. rectangularis 77. T. siamensis 78. T. subpubenceus
79. Athyrium davidii 80. A. gymnogrammoides 81. A. puncticaule 82. A. praetereuissum
83. A. rubricaule 84. Diplazium bellum 85. D. sibricum 86. D. squamigerum 87. D. sylvaticum 88. Ctenitis
subglandulosa 89. Dryopteris filix-mas 90. D. odontoloma 91. D. sikkimensis 92. D. subtriangularis
93. Pleocnemia submembranacea 94. Polystichum acutidens 95. P. luctuosum 96. P. mannii
97. P. woodsioides 98. Tectaria fissa 99. T. griffithi 100. T. simouisii 101. Nephrolepis radicans
102. Oleandra musaefolia 103. Bolbitis presliara 104. B. semicordata 105. Elaphoglossum angulatum
106. Araiostegia hymenophylloides 107. Araiostegiella hookeri 108. A. perdurams 109. Davallia
assamica 110. D. denticulata 111. D. divaricata 112. D. repens 113. D. solida.

3.8 STATUS OF ISOETACEAE AND MARSILEACEAE

Many species of Isoetaceae and Marsileaceae have been described from India as if endemic. But
according to Chandra et al. (2008), “Their status as species is doubtful and perhaps most of them
appear to represent no more than infraspecific variation”.... “Further study is required into separation
of species, and their correct nomenclature and synonymy”. These workers have, however, listed 26
species of Isoetes and 15 species of Marsilea in their detailed and commendable contribution of 414
threatened pteridophytes of India, for all of which we should be cautious while using them for our
classroom/laboratory/research or other requirements.

TEST YOUR UNDERSTANDING

1. What are included under ‘threatened species’ of any group of plants?

2. Describe in brief various aspects of threatened pteridophytes of India.
3. How many species of pteridophytes did Chandra et al. (2008) describe under the status of
threatened pteridophytes of India?
4. Out of 219 species of pteridophytes included in the ‘At Risk’ category, how many are critically
endangered?
5. Write a brief note on Red Data Book of IUCN.
6. Give an account of major categories of threatened pteridophytes.
7. How will you distinguish between the following categories of threatened pteridophytes?
(a) At Risk (b) Near-Threatened (c) Rare
8. Name any ten Indian threatened pteridophytes which fall in the ‘At-Risk’ category.
 CHAPTER 4

Economic
Importance of
Pteridophytes

Pteridophytes are not as economically important as seed plants but still have considerable importance.
Some are used as food plants and cooked as vegetables (e.g. Pteridium aquilinum), others are well-
known biological fertilisers (e.g. Azolla) and a few are noxious weeds (e.g. Salvinia molesta). The
medicinal value of several pteridophytes is widely accepted. Ferns are known for their ornamental
value throughout the world.
About 170 species of pteridophytes have been found to be used as food, flavour, dyes, medicines,
biofertilisers, oils, fibres and biogas production (Manickam and Irudayaraj, 1992). Recently, Perumal
(2010) stated “about the medicinal value of pteridophytes against bacteria, fungi, viruses, cancer,
rheumatism, diabetes, inflammation, fertility, diuretics, pesticides, hepatoprotectives and sedatives.
Besides sugar, starch, proteins and amino acids, ferns contain a variety of alkaloids, glycosides,
flavonoids, terpenoids, sterols, phenols, etc., as potential components used in various industries”.

4.1 FOOD PLANTS

Some pteridophytes are widely used as a source of food and fodder. These include fiddleheads of
bracken fern (Pteridium aquilinum), ostrich fern (Matteuccia struthiopteris), and cinnamon fern
(Osmunda cinnamomea). Diplazium esculentum (vegetable fern) is also used by some tropical people
as food. Marsilea is used as a substitute for clover to feed animals. In several parts of south-east Asia,
the young circinately coiled leaf tips of ferns are eaten as table delicacies. In Canada and adjacent USA,
the croziers of Matteuccia struthiopteris are served as common spring vegetables. They are also canned
and frozen for later use.
Pteridium aquilinum fiddleheads, however, should be used with utmost care. Their cooked vegetable
is believed to be responsible for the high rate of stomach cancer in Japan. It is also one of the world’s
most common agricultural weeds, especially in the British highlands, and often poisons horses and
cattle.
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Title: Pompei e le sue rovine, Vol. 3 (of 3)

Author: Pier Ambrogio Curti

Release date: December 5, 2023 [eBook #72323]

Language: Italian

Original publication: Milano: Sanvito, 1872

Credits: Barbara Magni and the Online Distributed Proofreading

Team at http://www.pgdp.net (This file was produced
from images made available by The Internet Archive)

*** START OF THE PROJECT GUTENBERG EBOOK POMPEI E

LE SUE ROVINE, VOL. 3 (OF 3) ***
 POMPEI
E LE SUE ROVINE
VOL. III
 POMPEI
E LE
SUE ROVINE
PER L’AVVOCATO
PIER AMBROGIO CURTI
GIÀ DEPUTATO AL PARLAMENTO NAZIONALE
DIRETTORE DELLA SOCIETÀ ITALIANA DI ARCHEOLOGIA
E DI BELLE LETTERE DI MILANO

VOLUME TERZO
 1874
MILANO — F. SANVITO, EDITORE.
NAPOLI — DETKEN E ROCHOLL.
 Proprietà letteraria.
Legge 25 Giugno 1865. Tip. Guglielmini.
INDICE
 CAPITOLO XIX.
Il quartiere de’ soldati e il Pagus Augustus
Felix.

Quartiere de’ soldati, o Ludo gladiatorio? — Pagus

Augustus Felix — Ordinamenti militari di Roma —
Inclinazioni agricole — Qualità militari — Valore personale
— Formazione della milizia — La leva — Refrattari —
Cause d’esenzione — Leva tumultuaria — Cavalleria —
Giuramento — Gli evocati e i conquisitori — Fanteria:
Veliti, Astati, Principi, Triarii — Centurie, manipoli, coorti,
legioni — Denominazione delle legioni — Ordini della
cavalleria: torme, decurie — Duci: propri e comuni —
Centurioni — Uragi, Succenturiones, Accenti,
Tergoductores, Decani — Signiferi — Primopilo — Tribuni
— Decurioni nella cavalleria — Prefetti dei Confederati —
Legati — Imperatore — Armi — Raccolta d’armi antiche
nel Museo Nazionale di Napoli — Catalogo del comm.
Fiorelli — Cenno storico — Armi trovate negli scavi
d’Ercolano e Pompei — Armi dei Veliti, degli Astati, dei
Principi, dei Triarii, della cavalleria — Maestri delle armi —
Esercizj: passo, palaria, lotta, nuoto, salto, marce —
Fardelli e loro peso — Bucellatum — Cavalleria numidica
— Accampamenti — Castra stativa — Forma del campo
— Principia — Banderuole — Insegne — Aquilifer —
Insegna del Manipolo — Bandiera delle Centurie —
Vessillo della Cavalleria — Guardie del campo —
Excubiæ e Vigiliæ — Tessera di consegna — Sentinelle —
 Procubitores — Istrumenti militari: buccina, tuba, lituus,
cornu, timpanum — Tibicen, liticen, timpanotriba —
Stipendj militari — I Feciali, gli Auguri, gli Aruspici e i
pullarii — Sacrifici e preghiere — Dello schierarsi in
battaglia — Sistema di fortificazioni — Macchine
guerresche: Poliorcetiæ: terrapieno, torre mobile,
testuggine, ariete, balista, tollenone, altalena, elepoli,
terebra, galleria, vigna — Arringhe — La vittoria, inni e
sacrificj — Premj: asta pura, monili, braccialetti, catene —
Corone: civica, murale, castrense o vallare, navale o
rostrale, ossidionale, trionfale, ovale — Altre distinzioni —
Spoglia opima — Preda bellica — Il trionfo — Veste
palmata — Trionfo della veste palmata — In Campidoglio
— Banchetto pubblico — Trionfo navale — Ovazione —
Onori del trionfatore — Pene militari: decimazione,
vigesimazione e centesimazione, fustinarium, taglio della
mano, crocifissione, fustigazione leggiera, multa, censio
hastaria — Pene minori — Congedo.

Nel capitolo di quest’opera I Fori, dicendo del Foro nundinario o

venale, toccai dell’opinione manifestata da molti che anzi essere un
simile foro, quel luogo fosse invece un quartiere di soldati, e venni
osservando come da essi si avesse per avventura a scambiare la
parte per il tutto, riconoscendo io con altri come in tal foro si
ritrovasse un quartiere sia di soldati, sia di gladiatori, come forse
meglio sembrasse al Padre Garrucci.
Bréton, malgrado le dimostrazioni fatte da quest’ultimo scrittore, e
malgrado che sia tratto a riconoscere ch’egli abbia nella maniera la
più positiva stabilito in una dissertazione inserta nel numero
tredicesimo del Bollettino Archeologico napolitano del gennajo 1855,
che si trattasse in questo luogo di un di que’ Ludi gladiatorii, di cui
parla Giusto Lipsio [1], non sa risolversi ancora a non ritenerlo per un
quartiere di soldati.
«Io non so in verità, scrive egli, perchè siasi cercato di sostituire
questa denominazione di foro nundinario a quella di Quartiere de’
Soldati, che venne a siffatto luogo fin dall’origine assegnato. È
evidente che una città dell’importanza di Pompei, una città fortificata,
dovesse avere una guarnigione e, per alloggiarla, una caserma.
Perchè dunque cercare questa caserma altrove e non nel
monumento il più piano e così conforme alla sua destinazione? Le
porte strette e poco numerose sarebbero state incommodissime e
perfino dannose per un mercato dove una folla numerosa si sarebbe
pigiata in disordine, mentre che esse potevano perfettamente
bastare a soldati che marciavano per due e regolarmente. Una
bardatura di cavallo da sella, armi, cimieri, memidi o calzari di
bronzo vennero qui trovati, che a dir vero sono più proprii di
gladiatori, che non di soldati; ma che conchiuderne per ciò? Che
gladiatori di passaggio a Pompei presero alloggio nella caserma, ciò
che è più naturale che prender alloggio nel mercato. Nelle camere
non si rinvennero letti: ma si sa che i soldati giacevano per lo più
sulla paglia. Il solo grande appartamento che esiste dovette essere
destinato al capo della guarnigione. Una sola cucina sarebbe stata
insufficiente se il nutrimento di tutti non fosse stato ammanito in
comune, e noi vediamo che quella del quartiere de’ soldati era
evidentemente destinata alla preparazione degli alimenti d’una gran
quantità di persone. Finalmente, quale sarebbe stata in un mercato
la destinazione di una prigione così severa come quella che qui vi fu
riconosciuta? S’arroge che in nessun luogo della città si rinvenne
una così grande quantità di scheletri, non essendosene contati meno
di sessantatrè, ripartiti principalmente nelle camere del primo piano.
Non è egli quindi supponibile che taluni motivi di disciplina abbiano
ritenuto i soldati al loro posto troppo lungo tempo per permettere a
tutti di sottrarsi alla morte?» [2]
Questi sono gli speciosi argomenti di Bréton, che potrebbero ben
anco essere conformi a verità, se quelli addotti dal Garrucci non
fossero stati buoni del pari e convincenti, per conchiudere che
dovesse essere invece un ludo gladiatorio. Io non presumo di
mettere innanzi un perentorio giudizio; solo permettendomi di
ricordare che ho già esternato l’avviso mio che inclina all’ipotesi del
Garrucci. Se non che al mio presente argomento ciò che preme di
stabilire si è che alla vita romana in Pompei non potesse mancare
quanto aveva tratto alla vita militare, e quindi dovevan esistere e una
caserma, se forse non ve n’erano anche di più, e posti e stazioni;
che infatti alla Porta di Ercolano si trovò morta, l’alabarda in pugno,
la sentinella, che fida alla sua consegna, anzichè mancarvi, e
cercare come tutti gli altri cittadini lo scampo nella fuga, erasi
lasciata soffocare dall’aere graveolenta e seppellire sotto le ceneri e
i lapilli.
Ma se qui non erano alloggiati i soldati, se questa non era la
caserma, ma un ludo gladiatorio, o locali attinenti solo al foro venale,
e dove trovar dovevansi soldati, posto che Pompei, come non è
contraddetto, fosse città importante, e di una importanza ben anco
militare, avesse mura, saracinesche, opere di fortificazione, e se
anzi ben due volte vi furono dedotte colonie militari, l’una volta al
tempo e per gli ordini di Silla e l’altra per quelli di Augusto?
Potrebbesi rispondere a siffatta domanda con quei dati storici che
Bréton medesimo prepose all’opera sua: «Silla ordonna que Pompei
fût reduite en colonie militaire sous le double nom de Colonia
Veneria, Cornelia, emprunté aux noms du dictateur et de la divinité
protectrice de la ville. Il y envoya des troupes sous le
commandement de son neveu Publius Sylla: mais les Pompéiens,
regardant ces colons comme des étrangers, leur refusèrent les droits
de cité...»
E più sotto:
«Quoiqu’il en soit, les colons furent forcés d’habiter hors de la ville
dans un faubourg, qui, lorsque plus tard, Auguste eut envoyé une
nouvelle colonie de vétérans, prit le nom de Pagus Augusto-
Felix [3].»
Da queste nozioni di storia pompeiana, che sono conformi a quelle
che ho pur io date nei capitoli del primo volume di quest’opera,
inferisco: a che dunque cercar in città caserme e stazioni militari, se i
soldati dovevano rimanere fuori della città? Vero è che quanto è
scoperto del Pagus Augustus-Felix non ha rivelato quartiere di sorta,
ma solo quella parte che l’attraversa ed è la Via delle Tombe e che
percorreremo nell’ultimo capitolo di quest’opera; ma rammentiamoci
altresì che ancor molto rimane a trarre in luce e che gli scavi ulteriori
ponno co’ loro risultamenti diradare ogni dubbio e risolvere la
quistione.
Dopo ciò, dinnanzi al fatto delle sentinelle summentovate e dopo le
diverse guerre e fazioni guerresche narrate in quest’opera nei
capitoli della storia, a soddisfare agli intenti dell’opera ed a chiudere
di essa quanto ha riferimento alla vita publica romana, riprodotta in
Pompei, entrerò a dire degli ordinamenti militari e di quanto ha tratto
all’armamento; ben francando la spesa il conoscere siccome
fossero, perocchè non di poco avessero a contribuire a quei trionfali
successi ch’ebbero sempre le armi romane. Gli scavi di Ercolano e
di Pompei portarono discreto contributo all’archeologia per farci
conoscere armi ed attrezzi militari e guerreschi, ed io di questi più
innanzi ne tratterò il meglio che mi sarà dato.
Ho già provato, trattando del commercio de’ Romani, che lungi
costoro dall’essere, come si crede erroneamente dall’universale, un
popolo soldato per istinto, lo fosse invece costretto dalla necessità, e
conquistando l’universo non lo facesse che per proteggere la sua
indipendenza o per difendersi, che non pugnò insomma che
vagheggiando le dolcezze della pace, alla quale, appena il poteva, si
abbandonava. Orazio compendia le aspirazioni de’ Romani quando
esclama:

O rus quando ego te aspiciam [4]?

I Romani in fatti ebbero a preservarsi dai Sabini, dagli Etruschi, dai

Latini, dai Sanniti, in tutti i quali erano elementi di grandissima
resistenza; onde Properzio, che tutto abbracciava il sentimento
dell’antichità, era nel vero giudicando l’Italia in quel verso che già
m’avvenne di dover riferire:
Armis apta magis tellus quam commoda noxæ [5],

più propria, cioè, alle armi, che non alla aggressione e distruzione.
Se dunque al soldato romano si può rimproverare un sol vizio,
l’avarizia, perchè l’orgoglio è più spesso nel soldato una virtù; di
ricambio ebbe l’onor militare come noi l’intendiamo pure oggidì, il
rispetto al giuramento, la devozione al suo capo, il gusto della
disciplina. I Romani vinsero il mondo con la tattica, la disciplina, la
forza d’insieme, la costanza e il sentimento d’essere Romani. Gli altri
popoli usavano di armi straordinarie e di macchine, il popolo romano
della spada. Anche contro i Germani, individualmente sì bravi e sì
forti, era colla pugna corpo a corpo e colla spada che i Romani
avevano la vittoria; onde Germanico così poteva dire alle sue truppe:
Non enim immensa barbarorum scuta, enormes hastas inter truncos
arborum et enata humo virgulta, perinde haberi quam pila et gladios,
et hærentia corporis tegmina. Densarent ictus, ora mucronibus
quærerent [6]. Potrebbesi e vizio e virtù che ho mentovati,
comprovare coi fatti alla mano; ma la storia di Roma è troppo notoria
per avere d’uopo di ricorrere a ciò.
Piuttosto m’occuperò qui ad informare il lettore della formazione di
questa famosa milizia conquistatrice dell’universo.
E prima devesi portar l’attenzione sulla scelta, dilectus, che era il
raccogliere e l’iscrivere i soldati in codici o matricole. Tale scelta
veniva fatta tra cittadini e socj o confederati; — rado avvenne che si
ricorresse ai poveri ed agli schiavi, — e venivano poi ascritti o a’ fanti
o a’ cavalieri. Nella milizia navale si accoglievano anche le persone
più abbiette e i libertini.
Il principio della milizia era al diciassettesimo anno, la fine al
cinquantesimo. Chi per altro serviva di continuo, terminava i suoi
obblighi a’ trentasette anni; gli altri, dove non avessero compiuto il
lor servizio al quarantesimo sesto anno, non n’erano liberati e si
potevano costringere finchè non avessero compiuti i cinquant’anni.
Altro requisito della milizia era il censo, solo volendovisi i ricchi, gli
onesti e coloro i quali, avendo beni tutti proprii, in certo modo
presentassero solidarietà d’interessi colla cosa publica.
La leva, o coscrizione, delle truppe, testimonio Dionigio
d’Alicarnasso [7], si faceva ogni anno, designandosi all’uopo due
consoli, che alla loro volta, congiuntamente al popolo, creavano
ventiquattro tribuni per capi di quattro legioni.
La cernita si faceva, previa publicazione dell’editto a mezzo del
banditore, dai tribuni in Campidoglio, estraendo a sorte dalle tribù e
classi, alla presenza dei consoli assisi nelle sedie curuli. I refrattarj
che si sottraevano alla milizia, i consoli comandavano venissero
ricercati e tradotti in carcere, talvolta puniti di verghe, venduti i loro
beni e qualche volta benanco multati dell’estremo supplizio o della
morte civile, venduti cioè pubblicamente schiavi o notati d’infamia.
Tre giuste cause sottrar potevano al servizio: la prima era la
dispensa, vacatio, per l’età, se già raggiunto il cinquantesimo anno;
per onore, se fosse taluno nella magistratura o nel sacerdozio; per
beneficio se il Senato e il Popolo consentivano: la seconda causa
dicevasi emeritum, ed era per chi aveva compiuti venti stipendj: la
terza era vizio o malattia, come i mancini, i gracili, chi mancasse di
pollici o altre dita, gli inetti a reggere scudo o gladio.
Nella leva così detta tumultuaria, od anche subitaria [8], che seguiva
nell’imminenza di qualche pericolo, non si osservavano grandi
formalità, esentandosi soltanto quelli ch’erano gravemente infermi od
inabili affatto.
Per ciò che riguardava la cavalleria, spettava ai censori il
determinare chi vi dovesse appartenere. Duplice poi era il corpo da’
cavalieri, l’uno costituivasi di quelli che ottenevano dal publico il
cavallo e il suo mantenimento, ed erano i soli che una volta
dicevansi cavalieri, equites; l’altro di coloro che non l’ottenevano.
Costoro potevano allora servire tra i pedoni, pedites. Riguardavasi
molto a’ costumi per concedersi il cavallo, e però spettavane la
decisione al censore. Dopo, tal facoltà si arrogarono i principi.
Finita la coscrizione, i tribuni congregavano i militi delle rispettive
legioni e lor facevano prestare giuramento. Ignorasi però se
giurassero uno per uno, o se insieme. Consisteva la formula nel
giurare: sarebbero per seguire i consoli a qualunque guerra fossero
essi per chiamarli, non mai tentar cosa contraria al popolo, non
disertar mai le bandiere, raccogliersi al cenno de’ consoli, nè partir
mai senza l’ordine loro.
Eranvi poi gli evocati, che formavano spesso la forza degli eserciti,
quasi assunti dietro preghiera o domanda, ed erano per lo più
veterani, esperti e prudenti della milizia, che comunque avessero
assolti i loro servizj, li riassumevano tuttavia in grazia de’ consoli o
de’ capitani. Gli evocati venivano dispensati da certe opere faticose,
come del vallo e degli accampamenti e tenuti in maggior onore,
spesso considerati quasi centurioni.
I conquisitori erano coloro che si mandavano nelle campagne ad
ingaggiare la gioventù per la milizia od a scoprire i refrattarj che vi si
tenevan nascosti ed a persuaderli di costituirsi.
Toccato fin qui della cernita, veggiamo dell’ordine della milizia.
Vario era esso sia ne’ militi che ne’ duci: Giusto Lipsio lo considera e
distingue, rispetto ai primi, in generi e in parti.
Generi dei pedoni erano i Veliti, gli Astati, i Principi e i Triarj. Veliti
coloro che per poca età e ricchezza venivano assegnati a questo
infimo genere, e quasi inermi venivano esposti di fronte al nemico;
astati perchè dapprima combattevano colle aste, dopo poi, serbando
sempre lo stesso nome, combattevano coi pili, specie di giavellotti, e
coi gladii; principi, perchè nello schierarsi dell’esercito venivano nel
terzo ordine. Parrebbe tuttavia dal loro nome dovessero trovarsi
invece nella prima.
Parti della fanteria erano poi queste, nelle quali i generi si dividevano
dai tribuni e dai centurioni i militi pedoni, eccettuati i veliti: la
Centuria, che si componeva di sessanta militi ed era assegnata ad
un centurione; il Manipolo, che si costituiva di due centurie; la
Coorte, composta di tre manipoli ed aveva astati, principi o triarii, ed
anche per consueto i veliti. Scipione Africano istituì anche la Coorte
Pretoria, nella quale s’ascrivevano i volontarii e gli amici e che mai si
dipartiva dal Pretore, ad imitazione della Coorte Regia presso i
Macedoni; finalmente la Legione, perchè comprendeva tutti gli altri
ordini. Romolo l’istituì di tremila uomini; cacciati i Re, crebbe a
quattromila; a cinquemila montò nella guerra contro Annibale ed a
seimila la portò Scipione quando passò in Africa.
Essendo molte le legioni, — perchè se prima sotto i consoli furono
quattro, nella seconda guerra Punica ascesero a venticinque, nella
guerra civile fra Cesare e Pompeo se ne contarono quaranta e
nell’assedio di Modena cinquanta — ebbero diversa denominazione:
il più spesso si distinsero col numero progressivo come prima,
secunda, tertia; talvolta col nome del fondatore, come Augusta,
Claudiana; alcune dal nome degli Dei, di Marte, di Minerva, di
Apollo; altre dalle provincie trionfate, come Italica, Gallica, Cirenaica;
ed altre finalmente da qualche onorifica qualità, come la Vittrice, la
Fulminante, la Valente, la Ferrea, la Pudica, la Fedele [9].
Gli ordini della cavalleria, erano le torme e le decurie. Dividevansi in
dieci corpi. Ogni legione aveva dieci torme tricenarie, ossia tremila
cavalieri. Le ale, erano così chiamate a motivo della loro posizione
nella battaglia; onde dicevasi ala destra ed ala sinistra,
componevansi di soci e di confederati; le torme o compagnie
suddividevansi in tre decurie o brigate di dieci uomini.
Tito Livio ne fa sapere, come nel principio della seconda guerra
Punica, i Romani, veduta l’inferiorità della loro cavalleria rimpetto a
quella de’ Cartaginesi, usassero dei veliti come arcieri e frombolieri
per appiccar zuffa avanti le linee e spazzar la via all’esercito [10].
Questa era, per usare del linguaggio militare, la bassa forza: essa
per altro si completava coi suonatori di militari strumenti, con operai
armajuoli e costruttori di macchine guerresche, tormenta bellica et
impedimenta, e conduttori di bagagli, pel trasporto de’ quali non si
faceva uso, come di presente, di carriaggi, ma di bestie da soma,
perchè di minor impaccio e di servizio più pronto.
Ora dei duci.
Questi pure erano di due generi: proprii, quelli che erano preposti ad
una o a qualche parte dell’esercito, come i centurioni, e i tribuni;
comuni, coloro che erano preposti a tutti, come i legati e il
comandante in capo, imperator.
I centurioni venivan, d’ordine o consenso dei consoli, eletti dai tribuni
fra quelli della loro classe: sovente però si toglievano anche da
classi superiori, ma per segnalati meriti, massime per militari, distinti.
L’elezione dei centurioni era duplice. Nella prima se ne eleggevano
trenta, ed altrettanti nella seconda. Il primo eletto denominavasi
Primopilo ed era nel suo diritto di intervenire nei consigli militari, in
un coi tribuni e coi legati. Talvolta accadde, più per qualche
occasione e volontà del duce, che per diritto o costume, che tutti i
centurioni venissero ammessi nel consiglio. Insegna poi dei
centurioni era un baston di vite, di cui si valevano a punizione de’
soldati.
I centurioni già eletti si eleggono pure alla loro volta gli uffiziali, uragi,
o più veramente chiamati optiones. Quando venivano creati dai
tribuni, dicevansi accensi; ma quando la loro nomina fu devoluta ai
centurioni, ebbero quel nome di optiones ed anche di
sottocenturioni, succenturiones. Tergoductores erano que’
sott’uffiziali che compivano le funzioni che or sarebbero de’ sargenti;
decani quelli che or si direbbero caporali.
In mezzo a’ centurioni si trasceglievano due, prestanti per vigoria
d’animo e di corpo, per essere signiferi, o portatori del vessillo;
perocchè quantunque un solo fosse il vessillo, due tuttavia erano i
vessillarii o signiferi, acciò l’uno succedesse all’altro in caso di fatica,
essendo i vessilli pesanti, od anche all’evenienza di malattia.
Il Primopilo, primopilus ed anche primipilus, era il capo di tutti i
centurioni, come il prefetto e principe delle legioni. Egli aveva
autorità anche sul collega suo Primopilo sinistro e la tutela
dell’aquila, che era lo stendardo principale della legione [11], tanto
così che si dicesse aquila come sinonimo di primopilato.
Come i centurioni eran preposti ai manipoli, così i tribuni della milizia
erano a tutta la legione che ne aveva sei. Li istituì Romolo, li
mantennero i Re ed i Consoli; ma il popolo se ne avocò il diritto e il
suffragio, poi esclusa ancora la facoltà nel popolo, e questi volendola
rivendicare, restò convenuto che parte ne avesse il popolo e un
numero eguale i consoli; i primi detti anche comitiati, avuti in maggior
onore; gli altri detti Rutili o Rufuli. I tribuni erano eziandio di due
sorta, cavalieri e plebei. Spettava ai tribuni render giustizia e
conoscere delle cause capitali, dare il segnale alle guardie e
sentinelle, curare le veglie, vigilare le munizioni, provvedere agli
esercizi tutti. Portavano l’anello d’oro, gli altri militi non potendolo
portar che di ferro.
La cavalleria dividendosi in dieci turme, si pigliavano tre cavalieri per
ciascuna turma, e così si avevano trenta duci. Erano insomma tante
turme nella cavalleria quante erano nella fanteria le coorti; tante le
decurie quanti i manipoli: e per conseguenza altrettanti i duci; con
questo solo divario che nella turma ve ne era un solo, mentre nel
manipolo ve n’erano due. Nella turma erano tre le decurie e l’uffiziale
che comandava la decuria appellavasi decurione. Ciascuna turma
aveva un solo vessillo. Spettava ai duci delle turme la nomina degli
uragi od optiones; come nella fanteria.
I soci o confederati, in luogo dei tribuni, che solo spettavano ai militi
cittadini, avevano i prefetti e venivano costituiti dai consoli, pari nel
resto nei diritti e nella podestà ai tribuni. I Legati erano applicati agli
imperatori o comandanti in capo, non tanto per comandare, quanto
per giovar di consiglio, ed era il Senato che li destinava come pratici
della milizia presso del capo. Era grande dignità codesta, perocchè
col potere dell’imperatore avesse altresì diritto alla venerazione
dovuta ai sacerdoti. Cicerone li chiamava numi di pace e di guerra,
curatori, interpreti, autori di bellici consigli, ministri del provinciale
interesse. Il loro potere per altro era subordinato a quello del
comandante. Incerto era il numero loro, talvolta destinato uno per
legione, sempre come sembrava conveniente al Senato. Quelli
ch’eran preposti a tutto l’esercito, dicevansi consolari: pretorii quelli
assegnati alle legioni.