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Pteridology in the New Millennium

Pteridology in theNew Millennium

NBRI Golden Jubilee Volume

Edited by

Subhash ChandraNational Botanical Research Institute,

CSIR, New Delhi, India, Pteridology Laboratory, NBRI, Lucknow, India

and

Mrittunjai SrivastavaNational Botanical Research Institute,

CSIR, New Delhi, India, Pteridology Laboratory, NBRI, Lucknow, India

SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.

A C.I.P. Catalogue record for this book is available from the Library of Congress.

ISBN 978-90-481-6222-2 ISBN 978-94-017-2811-9 (eBook)

DOI 10.1007/978-94-017-2811-9

Printed on acid-free paper

All Rights Reserved© 2003

No part of this work may be reproduced, stored in a retrieval system, or transmittedin any form or by any means, electronic, mechanical, photocopying, microfilming, recording

or otherwise, without written permission from the Publisher, with the exceptionof any material supplied specifically for the purpose of being entered

and executed on a computer system, for exclusive use by the purchaser of the work.

Cover Photo: Pteris wallichiana Ag. - A threatened plant of Kumaun

Springer Science+Business Media Dordrecht Originally published by Kluwer Academic Publishers in 2003

NBRI GOLDEN JUBILEE VOLUME

This volume is a collection of research papers on

pteridology published in honour of Professor B. K. Nayar,

founder of the Pteridology Laboratory of the National

Botanical Research Institute, Lucknow, India, on the eve of

the golden jubilee year, 2002, of the NBRI, a National

Laboratory under C.S.I.R., New Delhi, India.

v

EDITORIAL ADVISORY BOARD

Professor S. S. Bir

Manouli House, 33-Y,

The Mall, Patiala-147 001 (India)

Professor S. C. Verma

5452/1, Cat-II, Modern Complex, Manimajra

Chandigarh-160 101 (India)

Email: [email protected]

Professor U. Sen

Botany Department, Kalyani University

Kalyani, West Bengal (India)

Professor G. K. Srivastava

Botany Department, University of Allahabad

Allahabad-211 002 (India)

E. mail: [email protected]

Professor P. G. Windisch

Universidade do Vale do Rio dos Sinos (UNISINOS)

Centro de Ciências da Saúde.

Laboratório de Botânica Av. Unisinos 950, São

Leopoldo, RS. 93.022-00 (Brazil)


Professor B. C. Tan

Cryptogamic Laboratory

Department of Biological Sciences

National University of Singapore, 10 Kent Ridge Crescent

119260 (Singapore)


Dr. X. C. Zhang

Institute of Botany, The Herbaria P. E.

The Chinese Academy Of Sciences

20 Nanxineun, Beijing-100093 (China)


vi

PROF. BALA KRISHNAN NAYAR

15 September, 1927�

vii

CONTENTS

Chapter Page No.

List of Contributors xiii

Foreword xxi

Editors’ Note xxiii

Preface I xxv

Preface II xxvii

Preface III xxix

Preface IV xxxi

GENERAL PAPERS

1 Professor Bala Krishnan Nayar: A biographical sketch

Subhash Chandra

1-10

2 The Pteridology Laboratory, N.B.R.I., Lucknow (India)

Mrittunjai Srivastava

11-13

SYSTEMATICS

3 Taxonomy and biogeography of Indian pteridophytes

Gopinath Panigrahi

15-24

4 Phylogenetics, biogeograpgy and classification of the

woodwardioid ferns (Blechnaceae)

Raymond Cranfill and Masahiro Kato

25-48

5 Review of the family Woodsiaceae (Diels) Herter of

Eurasia

Alexandr Ivanovich Shmakov

49-64

6 An Introduction to the pteridophytes of Hainan Island,

South China

Xian-Chun Zhang

65-71

7 Notes on new or interesting Asplenium species from

Western Asia, including comments on Ching & Wu

(1985), and Fraser-Jenkins (1992) Reliquiae

Reichsteinianae 1

Ronald Louis Leo Viane and Tadeus Reichstein

73-105

8 Diversity and distribution patterns of Aspleniaceae in

Brazil

Lana da Silva Sylvestre and Paulo Günter Windisch

107-120

9 Two new species of Cyathea from North-Eastern India

Tanoy Bandyopadhyay, Tuhinsri Sen, and Uday Sen

121-132

10 Distributional study of the genus Cyrtomium C. Presl

(Dryopteridaceae)

Jin Mei-Lu and Xiao Cheng

133-141

ix

11 Taxonomic studies on the family Sinopteridaceae and

the genus Ctenitopsis (Tectariaceae) from Hainan

Island, China

Gang-Min Zhang, Shi-Yong Dong, Xian-Chun Zhang, and

Shu-Gang Lu

143-151

12 Valuable and noteworthy Chinese fern specimens

preserved at the herbarium of Singapore Botanic

Gardens (SING)

Su Gong Wu and Benito Ching Tan

153-164

13 Pteridophytic flora of Northern part of Bihar (India)

adjacent to Indo-Nepal border

Alka Kumari and Ranjit Bahadur Srivastava

165-175

MORPHOLOGY, ANATOMY AND CYTOLOGY

14 Studies on the Chinese Selaginellaceae-V: Observations

on the forliar epidermis of Selaginella Beauv. from

Hainan Island in China

Ping Yang and Xiang-Chung Zhang

177-184

15 Morphological observations on the co-occurring

geophytes Isoetes duriei Bory (Lycophyta, Isoetaceae)

and Romulea columnae Sebast. et Mauri (Anthophyta,

Iridaceae)

Angelo Troìa, Elisabetta Oddo, and Enrico Bellini

185-189

16 Cytomorphological studies on some Indian population

of Isoëtes coromandelina L. f.

Gopal Krishna Srivastava, Meena Rai, and Mrittunjai

Srivastava

191-203

17 Studies on the morpho-taxonomy of Indian vittarioid

ferns

Jnan Bikas Bhandari and Radhanath Mukhopadhyay

205-215

18 Morphological and cytological variations of Japanese

Dryopteris varia group (Dryopteridaceae)

Su-Juan Lin, Masahiro Kato, and Kunio Iwatsuki

217-231

ECOLOGY AND FLORISTICS

19 Environmentally related demography: field studies on

Isoëtes lacustris L. (Lycophyta, Isoëtaceae) in Europe

Margrit Vöge

233-260

20 The vertical distributions of live and dead fern spores in

the soil of a semi-natural woodland in Southeast

Scotland and their implications for spore movement in

the formation of soil spore banks

Luciano Maurico Esteves and Adrian Francis Dyer

261-282

x

21 Species richness of pteridophytes in natural versus

man-made lowland forest in Malaysia and Singapore

Farida Binti Yusuf, Benito Ching Tan, and Ian Mark

Turner

283-298

22 The taxonomy and ecology of the pteridophytes of Mt.

Iraya and vicinity, Batan Island, Batanes Province,

Northern Philippines

Julie Fenete Barcelona

299-325

23 Some aspects of the fern flora (Filicopsida) of the

Netherlands

Piet Bremer

327-340

24 Demographic studies of homosporous fern populations

in South Siberia

Irina Ivanovna Gureyeva

341-364

25 Eco-morphological characteristics of Pyrrosia petiolosa

(Christ et Baroni) Ching (Polypodiaceae) in the

primorye territory, in far-Eastern Russia

Nina Mihailovna Derzhavina

365-374

26 Herbivory on three tropical fern species of a Mexican

cloud forest

Klaus Mehltreter and Javier Tolome

375-381

27 Animal interaction with pteridophytes with emphasis

on Indian records

Subir Bera, Biplab Patra, and Narayan Ghorai

383-395

PHYTOREMEDIATION / ECONOMIC UTILITY

28 Tolerance of heavy metals in vascular plants: Arsenic

hyperaccumulation by Chinese brake fern (Pteris

vittata L.)

Bhaskar Rao Bondada and Lena Qiying Ma

397-420

29 Economically viable pteridophytes of India

Har Bhajan Singh

421-446

REPRODUCTIVE BIOLOGY/TISSUE CULTURE

30 Gametophyte morphology and breeding systems in

ferns

Donald Ray Farrar

447-454

31 Some aspects of reproductive biology of the

gametophyte generation of homosporous ferns

Satish Chander Verma

455-484

32 Mating systems of Cyatheaceae native to Taiwan

Wen-Liang Chiou, Yao-Moan Huang and Pei-Hsuan

Lee

485-489

xi

33 Plantlet production of the Philippine Giant Staghorn

Fern [Platycerium grande (Fee) C. Presl] through spore

culture

Cecilia Beltran Amoroso and Victor Bucad Amoroso

491-495

34 Micropropagation and conservation of rare and

endangered ferns of the Southern Western Ghats

through in vitro culture

Visuvasam Soosai Manickam, Sambantham

Vallinayagam, and Marimuthu Johnson

497-504

PALEOBOTANY

35 Heterosporous water ferns from the late Cretaceous to

Neogene periods of India

Bharati Nandi and Saibal Chattopadhyay

505-520

xii

LIST OF CONTRIBUTORS

Cecilia Beltran Amoroso (Chapter 33)

Co-authors: Victor Bucad Amoroso

Department of Biology, College of Arts and Sciences

Central Mindanao University, 8710 Musuan, Bukidnon, Philippines

E. mail: [email protected]; [email protected]

Victor Bucad Amoroso (Chapter 33)

Co-authors: Cecilia Beltran Amoroso

Department of Biology, College of Arts and Sciences

Central Mindanao University, 8710 Musuan

Bukidnon, Philippines


Tanoy Bandyopadhyay (Chapter 9)

Co-authors: Tuhinsri Sen and Uday Sen

Department of Botany, Kalyani University, Kalyani – 741 235, India

Julie Fenete Barcelona (Chapter 22)

Philippine National Herbarium (PNH), Botany Division

National Museum of the Philippines, Manila, C.P.O. 2659, Philippines


Enrico Bellini (Chapter 15)

Co-authors: Angelo Troìa and Elisabetta Oddo

Dipartimento di Scienze Botaniche dell’Università, via Archirafi 38

Palermo, Italy

Subir Bera (Chapter 27)

Co-authors: Biplab Patra and Narayan Ghorai

Department of Botany, University of Calcutta, Calcutta – 700 019, India

E-mail: [email protected]

Jnan Bikas Bhandari (Chapter 17)

Co-author: Radhanath Mukhopadhyay

Pteridology laboratory, Department of Botany

The University of Burdwan, Burdwan – 713104, India

Bhaskar Rao Bondada (Chapter 28)

Co-author: Lena Qiying Ma

Soil and Water Science Department, University of Florida

Gainesville, FL32611-0290, USA


xiii

Piet Bremer (Chapter 23)

Provincial Board of Overijssel, Luttenbergstraat 2

8011 EE Zwolle, The Netherlands


Subhash Chandra (Chapter 1)

Pteridology Laboratory

National Botanical Research Institute, Lucknow – 226 001, India


Saibal Chattopadhyay (Chapter 35)

Co-author: Bharati Nandi

Department of Botany, University of Kalyani

Kalyani – 741 235, India

Present corresponding address:

District Land & Land Reforms Office, Jiban Pal’s Garden

PO and Dist: Hooghly – 712123, India


Xiao Cheng (Chapter 10)

Co-author: Jin Mei-Lu

Kunming Institute of Botany

Academia Sinica, Kunming, Yunnan 650204, P. R. China


Wen-Liang Chiou (Chapter 32)

Co-authors: Yao-Moan Huang and Pei-Hsuan Lee

Division of Forest Biology, Taiwan Forestry Research Institute

53 Nan-Hai Rd., Taipei 100, Taiwan


Raymond Cranfill (Chapter 4)

Co-author: Masahiro Kato

University Herbarium, 1001 Valley Life Sciences Building,

University of California, Berkeley, CA 94720-2465, USA


Nina Mihailovna Derzhavina (Chapter 25)

Faculty of Botany, Orel State University, Bld. 95

Komsomolskaya, Orel, Russia

E. mаil: tb-d@mаil.ru

Shi-Yong Dong (Chapter 11)

Co-authors: Gang-Min Zhang, Xian-Chun Zhang and Shu-Gang Lu

Institute of Botany, the Chinese Academy of Sciences

Beijing 100093, P.R. China

xiv

Adrian Francis Dyer (Chapter 20)

Co-author: Luciano Maurico Esteves

Royal Botanic Garden Edinburgh, 20A Inverleith Row

Edinburgh EH3 5LR, Scotland, UK


Luciano Mauricio Esteves (Chapter 20)

Co-author: Adrian Francis Dyer

Instituto de Botanica de Sao Paulo, Caixa Postal 4005

CEP 01061-970, Sao Paulo, SP, Brazil


Donald Ray Farrar (Chapter 30)

Department of Botany, Iowa State University, Ames, Iowa 50011, USA


Narayan Ghorai (Chapter 27)

Co-authors: Subir Bera and Biplab Patra

Post Graduate Department of Zoology, Presidency College, Kolkata – 700 073, India

Irina Ivanovna Gureyeva (Chapter 24)

Krylov Herbarium, Tomsk State University,

Prospekt Lenina, 36, Tomsk 634050, Russia


Yao-Moan Huang (Chapter 32)

Co-authors: Wen-Liang Chiou and Pei-Hsuan Lee



Kunio Iwatsuki (Chapter 18)

Co-authors: Su-Juan Lin and Masahiro Kato

The University of the Air, 2-11 Wakaba, Mihama-ku, Chiba 261-8586, Japan


Marimuthu Johnson (Chapter 34)

Co-authors: Visuvasam Soosai Manickam and Sanbantham Vallinayagam,

Centre for Biodiversity and Biotechnology

St. Xavier's College (Autonomous), Palayamkottai – 627 002, India

Masahiro Kato (Chapter 4 & 18)

Co-author: Raymond Cranfill (Chapter 4)

Co-authors: Su-Juan Linand Kunio Iwatsuki (Chapter 18)

Department of Biological Sciences, Graduate School of Science

University of Tokyo, Hongo 7-3-1, Tokyo 113-0033, Japan


xv

Alka Kumari (Chapter 13)

Co-author: Ranjit Bahadur Srivastava

Botany Department, M. S. College, Motihari, Bihar, India


Pei-Hsuan Lee (Chapter 32)

Co-authors: Wen-Liang Chiou and Yao-Moan Huang



Su-Juan Lin (Chapter 18)

Co-authors: Masahiro Kato and Kunio Iwatsuki

Department of Biological Science and Technology, School of Life Science

Nanjing University. 22 Hankou-lu, Nanjing, Jiangsu 210093, P.R. China


C/O Department of Biological Science, Graduate School of Science

University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan


Shu-Gang Lu (Chapter 11)

Co-author: Gang-Min Zhang, Shi-Yong Dong, Xian-Chun Zhang

Department of Environmental Science, Yunnan University

Kunming 650091, P. R. China

Lena Qiying Ma (Chapter 28)

Co-author: Bhaskar Rao Bondada

Soil and Water Science Department, University of Florida

Gainesville, FL32611-0290, USA


Visuvasam Soosai Manickam (Chapter 34)

Co-authors: Sambantham Vallinayagam and Marimuthu Johnson

Centre for Biodiversity and Biotechnology, St. Xavier's College (Autonomous)

Palayamkottai – 627 002, India


Klaus Mehltreter (Chapter 26)

Co-author: Javier Tolome

Departamento Ecología Vegetal, Instituto de Ecología, A.C.

A. P. 63, Xalapa 91000, Veracruz, Mexico


Jin Mei-Lu (Chapter 10)

Co-author: Xiao Cheng

Kunming Institute of Botany

Academia Sinica, Kunming, Yunnan 650204, P. R. China

xvi

Radhanath Mukhopadhyay (Chapter 17)

Co-author: Jnan Bikas Bhandari

Pteridology laboratory, Department of Botany

The University of Burdwan, Burdwan – 713104, India


Bharati Nandi (Chapter 35)

Co-author: Saibal Chattopadhyay

Department of Botany, University of Kalyani



Elisabetta Oddo (Chapter 15)

Co-authors: Angelo Troìa and Enrico Bellini


Palermo, Italy

Gopinath Panigrahi (Chapter 3)

Botanical Survey of India

A13/3, Kalindi Housing Estate, Kolkata – 700 089, India


Biplab Patra (Chapter 27)

Co-authors: Subir Bera and Narayan Ghorai

Department of Botany, Anandamohan College

Kolkata – 700 009, India

Meena Rai (Chapter 16)

Co-authors: Gopal Krishna Srivastava and Mrittunjai Srivastava

C. M. P. Degree College

Allhabad – 211 002, India


Tadeus Reichstein (Chapter 7)

Co-author: Ronald Louis Leo Viane

Institute for Org. Chemistry, St. Johanns-Ring 19

CH-4056 Basle, Switzerland

Tuhinsri Sen (Chapter 9)

Co-authors: Tanoy Bandyopadhyay and Uday Sen

Department of Botany, Kalyani University, Kalyani – 741 235, India

Uday Sen (Chapter 9)

Co-authors: Tanoy Bandyopadhyay and Tuhinsri Sen

Department of Botany, Kalyani University


xvii

Alexandr Ivanovich Shmakov (Chapter 5)

South-Siberian Botanical Garden

Altai State University

Barnaul 656099, Russia


Har Bhajan Singh (Chapter 29)

Raw Materials Herbarium & Museum

National Institute of Science Communication

Dr. K. S. Krishnan Marg, New Delhi – 110 012, India


Gopal Krishna Srivastava (Chapter 16)

Co-authors: Meena Rai and Mrittunjai Srivastav

Botany Department, University of Allahabad

Allhabad – 211 002, India

Mrittunjai Srivastava (Chapter 2 & 16)

Co-authors: Gopal Krishna Srivastava and Meena Rai (Chapter 16)

Pteridology Laboratory, National Botanical Research Institute

Lucknow – 226 001, India


Ranjit Bahadur Srivastava (Chapter 13)

Co-author: Alka Kumari

Botany Department, M. S. College, Motihari, Bihar, India


Lana da Silva Sylvestre (Chapter 8)

Co-author: Paulo Günter Windisch

Universidade Federal Rural do Rio de Janeiro (UFRRJ)

Departamento de Botânica. Seropédica, RJ. 23.871-970, Brazil


Benito Ching Tan (Chapter 12 & 21)

Co-author: Su-Gong Wu (Chapter 12)

Co-authors: Farida binti Yusuf and Ian Mark Turner (Chapter 21)

Department of Biological Sciences

National University of Singapore

119260 Singapore


Javier Tolome (Chapter 26)

Co-author: Klaus Mehltreter

Departamento Ecología Vegetal, Instituto de Ecología, A.C.

A. P. 63, Xalapa 91000, Veracruz, Mexico

xviii

Angelo Troìa (Chapter 15)

Co-authors: Elisabetta Oddo, and Enrico Bellini


Palermo, Italy


Ian Mark Turner (Chapter 21)

Co-authors: Farida binti Yusuf and Benito Ching Tan

National Parks Board,

Singapore Botanic Gardens, 1 Cluny Road

259569 Singapore

Sambantham Vallinayagam (Chapter 34)

Co-authors: Visuvasam Soosai Manickam and Marimuthu Johnson

Centre for Biodiversity and Biotechnology, St.

Xavier's College (Autonomous)

Palayamkottai – 627 002, India

Satish Chander Verma (Chapter 31)

Department of Botany, Punjab University, Chandigarh, India


5452/1, CAT-2, Modern Complex, Manimajra

Chandigarh – 160 101, India

E mail: [email protected]

Ronald Louis Leo Viane (Chapter 7)

Co-author: Tadeus Reichstein

Ghent University, Dept. of Biology, K.L. Ledeganckstraat 35

B-9000 Ghent, Belgium


Margrit Vöge (Chapter 19)

22117 Hamburg, Pergamentweg 44b, Germany


Paulo Günter Windisch (Chapter 8)

Co-author: Lana da Silva Sylvestre

Universidade do Vale do Rio dos Sinos (UNISINOS)

Centro de Ciências da Saúde. Laboratório de Botânica

Av. Unisinos 950, São Leopoldo, RS. 93.022-000, Brazil


Su-Gong Wu (Chapter 12)

Co-author: and Benito Ching Tan

Kunming Institute of Botany, The Chinese Academy of Sciences

Kunming 650091, P. R. China


xix

Ping Yang (Chapter 14)

Co-author: Xian-Chun Zhang

College of Life Science, Henan Normal University, P.R. China

Farida Binti Yusuf (Chapter 21)

Co-authors: Benito Ching Tan and Ian Mark Turner

Cryptogam Laboratory, Department of Biological Sciences

The National University of Singapore 119260, Singapore

Xian-Chun Zhang (Chapter 6, 11 & 14)

Co-authors: Gang-Min Zhang, Shi-Yong Dong, and Shu-Gang Lu (Chapter 11)

Co-author: Ping Yang (Chapter 14)


Beijing 100093, P. R. China


Gang-Min Zhang (Chapter 11)

Co-authors: Shi-Yong Dong, Xian-Chun Zhang and Shu-Gang Lu


Beijing 100093, P.R. China


xx

FOREWORD

The National Botanical Research Institute came into being as the 13th among a

chain of National Laboratories established during April, 1953 under the Council of

Scientific and Industrial Research by the Govt. of India for advanced research in

fields of specialisation which have a direct bearing on socio-economic, industrial

and scientific advancement of the nation. Christened initially as National Botanic

Gardens, the nucleus around which the institution took shape under its founder-

Director Late Prof. K. N. Kaul, was a large herbarium of Indian flora and a century-

old botanical garden spread over 35 ha of land on the banks of River Gomti in the

heart of Lucknow city.

It’s a matter of great pleasure and profound satisfaction to me that a Golden

Jubilee volume entitled, “Pteridology in the New Millennium” is being published

and released during the Golden Jubilee year of NBRI in the honour of Professor

B. K. Nayar who laid the foundation of the Pteridology Laboratory of the NBRI,

which is now a well equipped laboratory for the study of Indian pteridophytes.

Professor Nayar is a holistic Botanist as evident through his contributions and

publications in almost all the areas of study of Pteridophyta. The contribution of

Professor Nayar towards the development of modern Pteridology and the role of

NBRI in it is indeed great and very important. His publications will be valuable for

the younger generation of scientists in the field as well as for the more mature

research workers and teachers.

The editors deserve to be commended on their initiative in bringing out NBRI

Golden Jubilee volume. This publication will go a long way in helping the scientific

community of the world.

P. Pushpangadan

Director, NBRI, Lucknow (India)

July 15, 2002

xxi

EDITORS’ NOTE

On the eve of the Golden Jubilee year of National Botanical Research Institute,

Lucknow, we have much pleasure in presenting this commemoration volume in the

honour of Prof. B.K. Nayar, Ex-Head of Botany Department, Calicut University,

Kerala who is the founder of Pteridology Division of NBRI, Lucknow. It was due to

the untiring efforts and dedication of Prof. Nayar during his tenure as head of

Pteridology Division of NBRI, that it has become a center for research in

pteridology in the world and established its lofty traditions. It goes to the credit of

Prof. Nayar that those who were trained by him and who succeeded him kept up the

traditions and even enhanced them so that today the NBRI is well recognized as the

foremost center of pteridological research in this country.

Besides being a good scientist, Prof. Nayar is a great teacher, a social worker

and a good humanist. A sincere and painstaking guide of the research scholars who

had the privilege of working under his guidance and one who would go to any

extent to help them, it is but natural that he won their respect, affection and

confidence in abundance. Prof. Nayar is a thorough gentleman and one who is ever

ready to help, and full of affection goodwill and sympathy for his colleagues.

Prof. Nayar has devoted an entire lifetime to research work in pteridology and

has made many significant contributions in diverse lines of investigations on

pteridophytes during his long research career beginning with the time when he

worked as a Lecturer in Botany at Gauhati University (1951-1955). His research

activity reached its peak at NBRI, Lucknow where he was Head of the Pteridology

Division (1955-1970). And it continued at University of Calicut where he was

Professor and Head of the Department of Botany (1970-1987) and finally Emeritus

Professor (1987-1993). Prof. Nayar established two thriving Schools of Pteridology,

one at NBRI and the other at Calicut University, and both continue to contribute

actively to the different lines of research initiated by him while proliferating into

several others.

This book commemorates Prof. B.K. Nayar’s contribution to the advancement

of Pteridology and is presented as our humble homage in his honour. Many

prestigious learned bodies and organizations have earlier honoured Prof. Nayar,

including the Council of Scientific and Industrial Research (India) and he is a

Fellow of Indian Academy of Sciences, Linnaean Society of London, Indian Fern

Society and Palynological Society of India. In comparison, this commemoration

volume is only a trivial tribute to him. The list of well-known contributors to this

volume is indicative of the respect he commands from fellow pteridologists from all

over the world, and the affection from his colleagues and academic associates in the

country. The wide spectrum of the contributions is a rich tribute to his great repute

in the field of pteridology.

We would like to express here our deep regards for this great scientist, who is

our trusted friend and well-wisher and congratulate him for the international stature

he has earned by his rich contributions to the vast expanding knowledge of

pteridological science in the world.

xxiii

The senior editor (SC) has been associated with Prof. B. K. Nayar ever since his

joining the Pteridology Laboratory of NBRI in 1963 and is deeply indebted to him

for motivating and inducting him to pteridological research; it is Prof. Nayar’s

continuous encouragement, sincere advice, help and co-operation that earned for the

senior editor his present standing in pteridology which gave him the courage to

undertake the present task and make this Golden Jubilee Volume possible.

We thankfully acknowledge the co-operation extended to us by the eminent

pteridologists who have contributed different chapters to this book, each one of

which has greatly enhanced its worth. We wish to extend our humble and warmest

thanks to the Director, NBRI, Lucknow, without whose continued encouragement

and blessing, this work would not have been possible. We are deeply indebted to

the members of the Editorial Advisory Committee and the learned reviewers for

their valuable suggestions, kind co-operation and encouragement during the

preparation of this volume. Special sincere thanks are also due to Dr. Xian-Chun

Zhang (Beijing, China) and Dr. S. N. Srivastava (Allahabad, India) for all the help,

co-operation and encouragement they have extended to us. We wish to record here

our sincere appreciation of the ungrudging help rendered by our colleagues Dr. P.

B. Khare, Dr. R. D. Tripathi, Mr. Sandip Behra, Mr. Manoj Kumar Srivastava, Mr.

Amitosh Verma and Mr. Shyam Babu during the preparation of this volume. Busy

scientists as they are, Prof. P. G. Windisch (Brazil), Prof. H. P. Nooteboom (The

Netherlands), Prof. Su Gong Wu (China) and Prof. S. S. Bir (India) has found time

at our request to write Prefaces to this book, and Dr. P. Pushpangadan the

Foreword, and it is a pleasure for us to record here our gratitude to each of them.

We are extremely thankful to Ms. Zuzana Bernhart Ing., Publishing Editor of

Kluwer Academic Publishers, The Netherlands for all the help she has rendered in

the publication of this volume in its present form and to Ms. Amber Tanghe-Neely

(Dordrecht, The Netherlands) for her editorial comments and suggestions for

improvement, which proved invaluable to us. We are also indebted to Mr. Dilip

Kumar Chakraborty (Lucknow, India) for the preparation of camera-ready

manuscripts for the press.

In editing this volume we have made an attempt to highlight the current

significance some of the emerging fields of pteridological research. We sincerely

hope that pteridologists will welcome this volume as a worthy treatise on

pteridology in the new millennium.

Subhash Chandra

Mrittunjai Srivastava

Pteridology Division, NBRI, Lucknow

xxiv

PREFACE I

NBRI, National Botanical Research Institute (originally known as National Botanic

gardens till October 1978) located at Lucknow in Uttar Pradesh was established in

April 1953. After fifty years of its existence, the NBRI is celebrating its Golden

Jubilee with activities spanning over one year (2002-2003).

Amongst the centers of active research in Pteridology in India the NBRI group

played a significant role. The credit for initiating this goes to the broad vision of

Prof. K.N. Kaul, the founder director of the institute who created the Division of

Cryptogamic Botany and appointed Prof. B. K. Nayar (previously at Gauhati

University, Assam) as scientist in charge, in 1955. Soon, a well-equipped laboratory

was established for the study of Indian Pteridophytic flora especially covering the

aspects of their taxonomy, nomenclature, anatomy, morphology, palynology, etc.

Cultivation of ferns and fern allies received special attention; with the result that

today NBRI has the best sanctuary of Pteridophytic flora of India in the country.

The credit goes to the leadership of Prof. Nayar (who was at NBRI till 1970).

Devoted work of Prof. Nayar and subsequent scientists in charge of the section as

Dr. Surjit Kaur and Dr. Subhash Chandra together with their colleagues resulted in

high academic output from the Pteridology Laboratory. This has been possible only

due to the farsighted and encouraging approach of the various Directors of NBRI

who headed the institute from time to time after Prof. Kaul. I will not detail out the

research activities of especially Prof. Nayar, the person who headed the section, and

Pteridologists at NBRI during 50 years of institute’s existence and provided

leadership, since separate write-ups on Pteridology Division of NBRI and Bio-data

and Research Contributions of Prof. Nayar are given elsewhere in this book.

I join the scientists of the institute, especially the present Director, in felicitating

Prof. Nayar for setting fair traditions of devoted and high quality research, which

brought laurels to the institute. Here, scientists are presently involved in multifarious

activities in researches in plant sciences with attainments of high quality. NBRI has

emerged as a jewel amongst the plant based research institutes in the country.

The present volume, which contains papers covering wide spectrum of various

aspects of Pteridology from prominent scientists of the world is intended to be Prof.

B. K. Nayar Commemoration Volume in recognition of his highly meritorious

research in Pteridology. He made valuable additions to the existing knowledge

about this interesting group of plants. I have also been engaged in India in the study

of various aspects of ferns and fern allies for over 5 decades now. This is an added

reason that I hold Prof. Nayar in high esteem for his scholastic contributions on

ferns. Also, I have always valued his friendship.

I heartily felicitate Prof. Nayar on his 75th

birth anniversary and it is my good

fortune to be associated with this auspicious occasion in the form of this write-up.

S.S. Bir

Punjabi University, Patiala, India

August 2, 2002

xxv

PREFACE II

My first contact with Prof. Bala Krishnan Nayar occurred in an indirect way, back

in 1971. As an apprentice to the Brazilian pteridologist Alexander Brade, one of the

tasks assigned to me was to study the then recently published paper by Prof. Nayar

(in Taxon), on a new classification of leptosporangiate ferns. I started reading it to

Prof. Brade, whose eyesight was quite impaired due to age. As I went on reading, a

series of "oh, yes", affirmative shakes of the head and his final emphatic statement

"the man is quite right!!!" got my attention to the author of that particular

publication, a landmark in taxonomic history. Since then, publications by Prof.

Nayar and his students were of primary interest and respect for me. My discussions

with late Prof. Karl Kramer, on the significance of Prof. Nayar’s work in the

upsurge of pteridology in India and its wide reaching influence the world over,

always elicited a very positive response.

It is a delicate task to present a foreword to this assemblage of contributions

reflecting the state of Pteridology at the beginning of this new millennium,

presented to honour Prof. Nayar. The chapters are related to the diverse fields of his

interest, which owe much to him, through the more than 200 research papers he

published in addition to several monographs on Indian Ferns, as well as books on

nomenclature and fern floras. Among Prof. Nayar's major contributions to

pteridology are his study of the spores of ca. 3500 taxa of pteridophytes, and study

and analysis of the morphology and development of the gametophytic generation of

more than 2000 taxa, each of which constitutes by itself a Herculean task well

carried out. Bringing all this information into a general evolutive framework, and

reinterpreting diverse morphologic and phylogenetic concepts as he has done

requires far more than perseverance and hard work. It calls for a brilliant and

dedicated mind, and an enormous appreciation and love for the ferns. His

dedication to pteridology gave us new perspectives on the phylogeny of ferns, and a

wealth of information on their richness, structure, biology and importance to

mankind. His in-depth knowledge of the structure of vascular plants, associated

with his interest in evolution, taxonomy, floristics, palynology, spore germination,

gametophyte morphology and development, brought a holistic approach to his

discussions and interpretations. His well-substantiated treatise on evolution of

primitive vascular plants and their cormophytic plant body has a far-reaching

impact on this challenging subject.

Born in 1927, he began his career (in 1950) as a lecturer at Gauhati University

(Assam), where the wealth and diversity of the fern flora kindled in him a passion

for pteridology, which was to last for ever. His establishing and leading the

Pteridology Division (1955-1970) at the National Botanical Research Institute

brought a landmark to Science in India and advancement of pteridology in the

world. In his research endeavors at NBRI he was very ably assisted by Dr. Prakash

Chandra, Dr. Surjit Kaur, and Dr. Subhash Chandra among others, and each of them

made significant contributions following his footsteps. The devotion and hard work

of Prof. Nayar and his followers made the Pteridology Division the largest and most

xxvii

productive research center in the world devoted to ferns and allied plants, with over

400 research papers, 4 books and 25 monographs produced so far. In 1970 Prof.

Nayar joined the University of Calicut as the first Professor and Head of the Botany

Department, bringing his scholarship, excellence in research and organizational

experience to this new Institution and to the associated Botanical Garden, which he

got established. He retired in 1987, but continued for another five years guiding

students and research projects in the University. Many students got inspired by him

to follow the paths his pioneering steps marked in pteridological research.

Even with its rich contents, the present book is only a very small homage to

Prof. Nayar's extensive work. His studies on the spores and gametophytes, inspired

extensive studies on gametophyte morphology and breeding systems as well as

spore banks in Europe, the production of Platycerium grande plantlets through

spore culture, researches on reproductive biology and mating systems. His

contributions to floristics and taxonomy are reflected in such publications as

Sinopteridaceae and Tectariaceae of Hainan, Valuable and Noteworthy Chinese

Ferns, Aspleniaceae in Brazil and Western Asia, and A Synopsis of Woodsiaceae of

Eurasia, and studies on the Indian vittarioid ferns as well as of the Dryopteris varia

and the Dryopteris sinofibrillosa complexes. His conservationist concerns are

reflected in the chapters on the Pteridophyte Preservation Areas in the Philippines,

and The Concept of Conservation Through Micropropagation. His interest in

ecological studies makes itself strongly felt through the contributions on the

application of ferns in phytoremediation of contaminated sites, establishment and

ecology of ferns in the Netherlands, demographic studies in South Siberia, animal

interactions and herbivory, and the impact of the alteration of tropical rain forests in

the fern flora. Prof. Nayar's pioneer landmark in modern fern phylogeny has greatly

influenced the new classification of the Woodwardioid ferns for the twenty first

century.

The wide range of topics and geographic origin of the contributed chapters

included in this commemoration volume bear witness to the far-reaching impact of

Prof. Nayar's research activities, as well as to the achievements of the Pteridology

Division at National Botanical Research Institute for which science and

pteridologists all over the world will always be grateful.

Prof. Paulo G. Windisch

Universidade do Vale do Rio dos Sinos - UNISINOS

Brazil

May 2, 2002

xxviii

PREFACE III

The contribution of Prof. B. K. Nayar in the advancement of modern pteridology and

in establishing a very productive school of research for the subject in NBRI as well as

at Calicut University is very great and important indeed. Throughout the world he is

well known as an accomplished and holistic botanist through his research contributions

and wealth of publications. Due to his untiring efforts and deep dedication, the

Pteridology Division of NBRI, which he founded and got established, earned a coveted

reputation as a center of excellence in research. Even after he left NBRI in 1970 to join

as Professor & Head of the Botany Department of Calicut University, the strong

traditions which he established and the pteridologists he trained kept high the

excellence and even enhanced it so that the Pteridology Laboratory continued to be

well known as an active center of pteridological research in India. In addition to his

superb organizational capabilities, as evidenced by the excellence of the two schools of

pteridology which he got established at NBRI and at Calicut University, Prof. Nayar is

an eminent scientist of international stature and admired and respected by

contemporary botanists the world over for his rich and lasting contributions to the vast

expanding knowledge of pteridological science in the world. As a teacher he is well

respected and loved by his students, and it goes to his credit that many among them

have in later years established themselves as eminent pteridologists following his

footsteps. This Golden Jubilee Volume, published in his honour, is in itself a testimony

to the love and respect he commands from his students and fellow scientists.

Prof. Nayar’s work has received international acclaim. He has made extensive

contributions to diverse fields of investigations on the pteridophytes. “THE

PHYLLORIZE THEORY”, which he developed and elaborated based on his studies on

the gametophytic as well as sporophytic generations of ferns, could as well prove to be a

turning point in our understanding of evolution of all vascular plants. The evidences he

provides from morphology of extant pteridophytes, especially their gametophytes and

ontogeny of vascular system, to support his hypothesis is quite convincing, and in

contrast to earlier treatises on the subject, totally obviates the need for conjectures of

hypothetical life forms. His theory visualized the successive steps in evolution of

vascular plants from thalloid non-vascular ancestors, in place of the Axial Theory and

the Stelar Theory, which dominated taxonomy and morphology. If universally accepted

this must prove to be a milestone in pteridological research. Special mention may be

made here also of his comprehensive review on the gametophyte of homosporous ferns,

a new phylogenetic classification of ferns, his books on nomenclature of Indian ferns

and on the fern flora of Malabar.

In this context it is very apt that this Golden Jubilee Volume entitled “Pteridology

in the new Millennium”, is published to honour Prof. Nayar and the extensive

contributions he has made to pteridology.

Dr. Hans P. Nooteboom

National Herbarium of the Netherlands, Leiden

May 27, 2002

xxix

PREFACE IV

Professor B. K. Nayar is one of the most eminent pioneers in research studies on

Pteridophyta. I came to know him and his extensive contributions quite late in life,

because it was only after the “Cultural Revolution” of China that I started my study

(in the 1970’s) of the Tibetan Pteridophyta to compile the Pteridophyta part in the

Flora of Xizangica (Tibet) under the direction of Prof. Ching Renchang. Prof.

Ching, the doyen of Chinese pteridology, then told me that in any study of

pteridophytes of Tibet, the work of Indian Pteridologists, especially of Prof. B. K.

Nayar and Prof. S. S. Bir, is very important and must be referred to without fail.

Soon I realized that the book by B. K. Nayar and S. Kaur, “Companion to R. H.

Beddome’s Handbook to the Ferns of British India”, was an unavoidable reference

for my work, as also the many monographs published by them on Indian fern

genera. Later, the elucidation of fern phylogeny contained in a large number of

publications of Prof. Nayar, particularly in the “New phylogenetic classification of

ferns” (in Taxon) and the many supplementary papers, which followed, proved to

be of invaluable help. Without the extensive contributions of contemporary Indian

researchers like Prof. Nayar on the taxonomy and nomenclature of Indian

pteridophytes, the Pteridophyta part of the Flora of Xizangica (Tibet) would not

have been possible.

By then I began to collect and read the vast wealth of papers written by Indian

Pteridologists. It was difficult to get them in China and there was little contact

between the scholars of the two countries, but eminent Indian pteridologists like

Prof. Nayar had close contact with Prof. Ching and this helped. Prof. Nayar’s

publications constituted the bulk of my collection, even though his contributions

covered very many aspects of pteridology other than systematic taxonomy. Perhaps

the most important work of his is the study on the gametophytes of ferns and the

morphology of spores, the taxonomic and phylogenetic implications of which he

amply demonstrated. The review published by Prof. Nayar and Dr. S. Kaur on

“Gametophytes of Homosporous Ferns” is a classical work, which for the first time

unambiguously brought out the significance of gametophyte morphology in studies

on taxonomy and evolution. Similarly, the PHYLLORHIZE THEORY put forward

by Prof. Nayar, based on his extensive study of gametophytes as well as

sporophytes including ontogeny of their vascular system, may prove to be the most

important contribution to pteridology during the last century. If Prof. Nayar’s

hypothesis of origin and evolution of the characteristic plant body of vascular plants

proves to be true, it could revolutionize the study of phylogenetic taxonomy, not

only of the pteridophytes but also of the flowering plants. Let us look forward to his

students continuing the study initiated by him and revolutionize the science of

pteridology in the new millennium. It is my regret that I have had no chance to meet

him and study from him. However, I am lucky to have met his contemporary, Prof.

S. S. Bir, with whom I had long discussions on Indian contributions to pteridology

and who have helped me to understand the extensive Indian work on the subject.

xxxi

Just as India, China is a vast country with a very rich pteridophyte flora, which

is the richest in East Asia and perhaps in the world, and both have, in the last

millennium, produced such stalwarts in pteridology like Prof. B. K. Nayar and Prof.

R. C. Ching. According to “Flora of China” published towards the end of last

century, there are 215 genera and 2535 species of pteridophytes in China. But many

problems of species concepts, nomenclature and phylogeny still persist. The

situation is perhaps better in India, through the effort of eminent pteridologists like

Prof. Nayar, and the prospects seem brighter because of his equally eminent

students like Dr. S. Chandra and Dr. P. V. Madhusoodanan who continue the work.

China and India have much in common in diversity of natural environment and of

floristic wealth especially of pteridophytes. In co-operation, the botanists of the two

countries could make significant and substantial contributions to Pteridology in the

New Millennium on the basis of the work of pioneers like Prof. Nayar and Prof.

Ching who have carved for themselves pre-eminent positions among pteridologists

of the world.

Su Gong Wu

Kunming Institute of Botany, Kunming,

Yunnan, 650204, China

June 5, 2002

xxxii

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