grasslands, systems analysis and...

Grasslands, systems analysis and man

www.cambridge.org© in this web service Cambridge University Press

Cambridge University Press978-1-107-40416-8 - International Biological Programme 19: Grasslands, Systems Analysis and ManEdited by A. I. Breymeyer and G.M. Van DyneFrontmatterMore information

http://www.cambridge.org/9781107404168

http://www.cambridge.org


THE INTERNATIONAL BIOLOGICAL PROGRAMME

The International Biological Programme was established by the International Council of Scientific Unions in 1964 as a counterpart of the International Geophysical Year. The subject of the IBP was defined as 'The Biological Basis of Productivity and Human Welfare', and the reason for its establishment was recognition that the rapidly increasing human population called for a better understanding of the environment as a basis for the rational management of natural resources. This could be achieved only on the basis of scientific knowledge, which in many fields of biology and in many parts of the world was felt to be inadequate. At the same time it was recognized that human activities were creating rapid and comprehensive changes in the environment. Thus, in terms of human welfare, the reason for the IBP lay in its promotion of basic knowledge relevant to the needs of man.

The IBP provided the first occasion on which biologists throughout the world were challenged to work together for a common cause. It involved an integrated and concerted examination of a wide range of problems. The Programme was co-ordinated through a series of seven sections representing the major subject areas of research. Four of these sections were concerned with the study of biological productivity on land, in freshwater, and in the seas, together with the processes of photosynthesis and nitrogen fixation. Three sections were concerned with adaptability of human populations, conservation of ecosystems and the use of biological resources.

After a decade of work, the Programme terminated in June 1974 and this series of volumes brings together, in the form of syntheses, the results of national and international activities.






INTERNATIONAL BIOLOGICAL PROGRAMME 19

Grasslands, systems analysis and man

EDITED BY

A. I. B R E Y M EYE R Associate Professor of Ecology, Institute of Geography, Polish Academy of Sciences

AND

G.M.VAN DYNE Professor of Biology, College of Forestry and Natural Resources, Colorado State University

CAMBRIDGE UNIVERSITY PRESS CAMBRIDGE LONDON NEW YORK NEW ROCHELLE MELBOURNE SYDNEY






CAMBRIDGE UNIVERSITY PRESS

Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, Sao Paulo, Delhi, Tokyo, Mexico City

Cambridge University Press The Edinburgh Building, Cambridge CB2 8RU, UK

Published in the United States of America by Cambridge University Press, New York

www.cambridge.org Information on this title: www.cambridge.org/978n07404168

© Cambridge University Press 1980

This publication is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press.

First published 1980

A catalogue record for this publication is available from the British Library

Library of Congress Cataloguing in Publication Data

Main entry under title:

Grasslands, systems analysis and man.

(International Biological Programme; 19) Includes bibliographies and index. 1. Grassland ecology. 2. Grassland ecology - Mathematical models. 3. International Biological Programme. I. Breymeyer, A. I., 1932-. II. Van Dyne, George M., 1932-. III. Series.

QHW·5·P7G74 574.5'264 77-28249

ISBN 0 521 21872 I

ISBN 978-0-521-21872-6 Hardback ISBN 978-1-I07-40416-8 Paperback

Cambridge University Press has no responsibility for the persistence or accuracy ofURLs for external or third-party internet websites referred to in this publication, and does not guarantee that any content on such websites is, or will remain, accurate or appropriate.






Contents

List of contributors page xiii

Foreword xvii J. B. Cragg

Contents of IBP 18 xxii

Introduction A. I. Breymeyer

Part I. Processes and productivity

Abiotic subsystem 11 W. T. Hinds & G. M. Van Dyne

2 Autotrophic subsystem 59 J. S. Singh, M. J. Tl'lica. P. G. Risser. R. E. Redmann & J. K. Marshall

3 Small herbivore subsystem 201 L. Andrzejewska &: G.Gyllenberg

4 Large herbivore subsystem 269 G. M. Van Dyne, N. R. Brockington, Z. Szocs, J. Duek & C. A. Ribic

5 Invertebrate predator subsystem 539 A. Kajak

6 Vertebrate predator subsystem 591 L. D. Harris & G. B. Bowman

7 Decomposer subsystem 609 D. C. Coleman, A. Sasson, A. I. Breymeyer, M. C. Dash, Y. Dommergues, H. W. Hunt. E. A. Paul, R. Schaefer, B. Ulehlova & R. I. Zlotin

v






Contents

Part II. Systems synthesis

8 Nutrient cycling F. E. Clark, C. V. Cole & R. A. Bowman

9 Comparative studies of ecosystem function N. I. Bazilevich & A. A. Titlyanova

10 Total-system simulation models G. S. Innis, I. Noy-Meir, M. Godron & G. M. Van Dyne

11 Trophic structure and relationships A. I. Breymeyer

Part III. System utilization

12 Management impacts on structure and function of sown grasslands

K. J. Hutchinson & K. L. King

13 Simulation of intensively managed grazing systems N. G. Seligman & G. W. Arnold

14 Reflections and projections G. M. Van Dyne

Index

VI

page 659

713

759

799

823

853

881

923






Table des matieres

Liste des collaborateurs

Avant-propos J. B. Cragg

Contenu du PBI 18

Introduction A. 1. Breymeyel'

liere partie. Processus et productivite

Sous-systeme abiologique W. T. Hinds & G. M. Van Dyne

2 Sous-systeme autotrophe J. S. Singh, M. J. Trlica. P. G. Risser. R. E. Redmann & J. K. M arsha/l

3 Sous-systeme des petits herbivores L. Andrzejewska & G. Gy/lenberg

4 Sous-systeme des grands herbivores G. M. Van Dyne, N. R. Brockington. Z. Szocs. J. Duek & c. A. Ribie

5 Sous-systeme des Tnvertebres predateurs A. Kajak

6 Sous-systeme des Vertebres predateurs L. D. Harris & G. B. Bowman

page xiii

xvii

xxii

11

59

201

269

539

591

VB






Table des matieres

7 Sous-systeme des decomposeurs 609 D. C. Coleman, A. Sasson, A. I. Breymeyer, M. C. Dash, Y. Dommergues, H. W. Hunt, E. A. Paul" R. Schaefer, B. Ulehlova & R.I. Zlotin

IIieme partie. Synthese des systemes

8 Cycle des substances nutritives 659 F. E. Clark, C. V. Cole & R. A. Bowman

9 Etudes comparatives sur Ie fonctionnement de l'ecosysteme 713 N. 1. Bazilevich & A. A. Titlyanova

10 Modelisation simulatoire du 'systeme-total' 759 G. S. Innis, I. Noy-Meir, M. Godron & G. M. Van Dyne

11 Structure et relations trophiques 799 A. I. Breymeyer

IIIieme partie. Utilisation des systemes

12 Impacts de l'exploitation sur la structure et Ie fonctionnement des prairies artificielles 823


13 Simulation de systemes de paturage intensivement exploites 853 N. G. Seligman & G. W. Arnold

14 Reflexions et remarques prospectives 881 G. M. Van Dyne

hldex 923

VIU






Co,n;ep)I(aHHe

CnucoK. a8mopo8

IIpeOuc/lo8ue J. B. Cragg

CO,D;ep)J(aHHe MEII 18

BBe,D;eHHe

A. I. Breymeyer

l{aCYL I. ITpo-.eccLi H npo,D;YKTHBHOCTh

1 A6HOTHQecKaJI nO,D;CHCTeMa

W. T. Hindv & G. M. Van Dyne

2 ABTOTpo4>HaJi nO,D;CHCTeMa

J. S. Singh, M. J. Trlica, P. G. Risser, R. E. Redmann & J. K. Marshall

3 IIO,D;CHCTeMa MeJIKHX TpaoOJl)lHbIX

L. Andrzejewska & G. Gyllenberg

4 IIO)lCHCTeMa KPYIIHbIX TpaB05l,lUlblX

G. M. Van Dyne, N. R. Brockington, Z. Szocs, J. Duek & C. A. Ribic

5 ITO,D;CHCTeMa XHIIUlbIX 6ecno3BOHOQHbIX

A. Kajak

6 IIO,D;CHCTeMa XHIIUlbIX n03BOHOtfHbIX

L. D. Harris & G. B. Bowman

CmpaHUl/O xiii

xvii

xxii

1

11

59

201

269

539

591

ix






CoiJepJlCaHUe

7 TIo.n;CHCTeMa pe.n;y~eHToB 609 D. C. Coleman, A. Sasson, A. I. Breymeyer, M. C. Dash, Y. Dommergues, H. W. Hunt, E. A. Paul, R. Schaefer, B. Ulehlova & R. I. Zlotin

1:1aCTh II. CHRTel CHCTeM

8 060pOT mlTaTeJIbHbIX Bem;eCTB 659 F. E. Clark, C. V. Cole & R. A. Bowman

9 CpaBHHTeJIbHbIe HCCJIe.n;OBaHHR <PYHK~OHHpoBaHHR 3KOCHCTeM 713 N. I. Bazilevich & A. A. Titlyanova

10 06m;eCHCTeMHbIe HMHTa~HOHHbIe Mo.n;eJIH 759 G. S. Innis, I. Noy-Meir, M. Godron & G. M. Van Dyne

11 TPO<PHqecKaR CTpYKTypa H B3aHMOOTHoweHHSI 799 A. I. Breymeyer

'Iacn. III. HCDo~loBaHHe CHCTeM ...

12 X03mCTBeHHOe B03.n;eHCTBHe Ha crpYKTypy If <PYHK~lfoHHpoBaHHe CeRHhIX rpaBRHbIX CHCTeM


13 Mo.n;eJIHpOBaHHe HHTeHCHBHO HCnOJIb3yeMbIx naCT6Hm;HbIx

CHCTeM

N. G. Seligman & G. W. Arnold

14 06cY)I(.n;eHHe H nepCneKTHBhI

G. M. Van Dyne

x

823

853

881

923






Sumario

Lista de eolaboradores

Pr610go J. B. Cragg

Sumario de IBP 18

Introducci6n A. I. Breymeyer

1 a parte. Procesos y productividad

EI subsistema abi6tico W. T. Hinds & G. M. Van Dyne

2 EI subsistema aut6trofo J. S. Singh, M. J. Trliea, P. G. Risser, R. E. Redmann & J. K. Marshall

3 EI subsistema de los pequeiios herbivoros L. Andrzejewska & G. Gyllenberg

4 EI subsistema de los grandes herbivoros G. M. Van Dyne, N. R. Brockington, Z. Szoes, J. Duek & c. A. Ribie

5 EI subsistema de los invertebrados predadores A. Kajak

6 EI subsistema de los vertebrados predadores L. D. Harris & G. B. Bowman

pagina xiii

xvii

xxii

1

11

59

201

269

539

591

xi






Sumario

7 El subsistema de los descomponedores 609 D. C. Coleman, A. Sasson, A. I. Breymeyer, M. C. Dash, Y. Dommergues, H. W. Hunt, E. A. Paul, R. Schaefer, B. Ulehlova & R. I. Zlotin

IIa parte. Siotesis del sistema

8 Ciclo de nutrientes 659 F. E. Clark, C. V. Cole & R. A. Bowman

9 Estudios comparativos de la funci6n del ecosistema 713 N. I. Bazilevich & A. A. Titlyanova

10 Modelos de simulaci6n del sistema total 759 G. S. Innis, I. Noy-Meir, M. Godron & G. M. Van Dyne

11 Estructura y relaciones tr6ficas 799

IlIa parte. Empleo del sistema

12 Impacto de la acci6n del hombre sobre la estructura y funci6n de las praderas sembradas 823


13 Simulaci6n de sistemas de pasto intensivos 853 N. G. Seligman & G. W. Arnold

14 Reflexiones y proyecciones 881 G. M. Van Dyne

Indice 923

xu






Contributors

Andrzejewska, L. Institute of Ecology, Polish Academy of Sciences, PO Lomianki 05 150, Dziekanow Lesny, Poland.

Arnold, G. W. Division of Land Resources Management, CSIRO, Private Bag, PO Wembley, WA 6014, Australia.

Bazilevich, N. I. Dokuchaev Soil Institute, Pyzhevski 7, Moscow-I7, USSR.

Bowman, G. B. School of Forest Resources and Conservation, Institute of Food and Agricultural Sciences, University of Florida, Gainsville, FL 32611, USA.

Bowman, R. A. USDA-ARS Soil Fertility and Management Research, W106 Plant Science, Colorado State University, Fort Collins, CO 80523, USA.

Breymeyer, A. I. Institute of Geography and Spatial Organization, Polish Academy of Sciences, Krakowskie Przedmiescie 30, 00 927 Warsaw, Poland.

Brockington, N. R. The Grasslands Research Institute, Hurley, Maidenhead, Berkshire S16 5LR, UK.

Clark, F. E. Agricultural Research Service, US Department of Agriculture, Fort Collins, CO 80521, USA.

Cole, V. C. Agricultural Research Service, US Department of Agriculture, Fort Collins, CO 80521, USA.

Coleman, D. C. Natural Resource E~ology Laboratory and Zoology! Entomology Dept, Colorado State University, Fort Collins, CO 80523, USA.

Dash, M. C. Postgraduate Department of Biological Science, Sambalpur University, Burla, Sambalpur, Orissa, India.

Dommergues, Y. ORSTOM, PO Box 1386, Dakar, Senegal.

Duek, J. J. Institute of Applied Statistics and Computing, University of the Andes, Merida, Venezuela.

Xlll






Contributors Godron, M. P.

Gyllenberg, G.

Harris, L. D.

Hinds, W. T.

Hunt, W. H.

Hutchinson, K. J.

Innis, G. S.

Kajak, A.

King, K. L.

Marshall, J. K.

Noy-Meir, I.

Paul, E. A.

Redmann, R. E.

Ribic, C. A.

Risser, P. G.

Sasson, A.

xiv

CNRS Louis Emberger Centre for Phytosociological and Ecological Studies, PO Box 5051, Route de Mende, 34033 MontpeIIier, France.

Zoological Institute, Helsinki University, N. Jarnvagagatan 13, Helsinki, Finland.

School of Forest Resources and Conservation, Institute of Food and Agricultural Sciences, University of Florida, Gainesville, FL 32611, USA.

Battelle, Pacific Northwest Laboratories, PO Box 999, Richland, Washington 99352, USA.

Biology Department, West Virginia University, Morgantown, West Virginia 26506, USA.

CSIRO Pastoral Research Laboratory, Armidale, NSW 2350, Australia.

Department of Wildlife Sciences, Utah State University, Logan, Utah 84321, USA.

Institute of Ecology, Polish Academy of Sciences, PO Lomianki 05-150, Dziekanow Lesny, Poland.

CSIRO Pastoral Research Laboratory, Armidale, NSW 2350, Australia.

CSIRO, Western Australian Labs, Division of Land Resource Management, Private Bag, PO Wembley 6014, Western Australia.

Botany Department, Hebrew University of Jerusalem, Jerusalem, Israel.

Saskatchewan Institute of Pedology, University of Saskatchewan, Saskatoon, Saskatchewan, Canada S7NOWO.

Department of Plant Ecology, University of Saskatchewan, Saskatoon, Saskatchewan, Canada S7N OWO.

Department of Ecology and Behavioral Biology, University of Minnesota, St Paul, MN 55108, USA.

Botany Department, University of Oklahoma, Norman, Oklahoma 73069, USA.

Division of Biology, UNESCO, Place de Fontenoy, Paris 75007, France and Dept of Biology, University of Mohammed V, Rabat, Morocco.






Schaefer, R.

Seligman, N. G.

Singh, J. S.

Szocs, Z.

Titlyanova, A. A.

Trlica, M. J.

Ulehlova, B.

Van Dyne, G. M.

Zlotin, R. I.

Contributors Plant Ecology Laboratory, Faculty of Sciences, University of Paris-XI, 91045 Orsay, France.

Agricultural Research Organization, The Volcani Center, PO Box 6, Bet Dagan 50200, Israel.

Department of Botany, Kumaun University, Nainital-263002, India.

Research Institute for Botany of the Hungarian Academy of Sciences, H-2163 Vacratot, Hungary.

Institute of Soil Science and Agrochemistry, USSR Academy of Sciences (Siberian Branch), Sovetskaya 18, Novosibirsk 99, USSR.

Department of Range Science, College of Forestry and Natural Resources, Colorado State University, Fort Collins, CO 80523, USA.

Institute of Botany, CASU, Stara 18, Brno, Czechoslovakia.

Department of Range Science, College of Forestry and Natural Resources, Colorado State University, Fort Collins, CO 80523, USA.

Institute of Geography, USSR Academy of Sciences, Moscow W-17, Staromonetny 29, USSR.

xv






Foreword

The first official mention of the Grasslands Biome studies in IBP is in a report of a meeting of the Productivity of Terrestrial Ecosystems section - IBP(PT) - held in Warsaw 30 August to 6 September 1966. The main purpose of the technical sessions at Warsaw was to consider' ... the principles and methodology of secondary productivity of terrestrial ecosystems'. Among the working groups which met during that meeting was one devoted to the study of grassland ecosystems. As IBP News No.7 (1966) records: 'The working team concerned with grassland ecosystems discussed the project of a fully coordinated research programme. Five specialists from different countries were concerned with this during the meeting, and it was decided to elaborate the project subsequently by mail.'

The grasslands studies became Theme 1 of IBP(PT). Later, when the distinction was sharpened between major studies on ecosystems and investigations on special topics such as certain groups of consumers, the grasslands projects became collectively known as the Grasslands Biome Studies. By 1972, when the majority ofIBP field studies were terminated, some thirty nations had shared in the work of the grasslands group, and several hundreds of scientists had been involved along with a much greater number of support staff.

The general aims of the grasslands theme were summarized in IBP News No. 13 (1969) as follows: 'Grasslands are one of the most important of terrestrial ecosystem types. Large areas occupy the interior of the principal continents, and provide, when managed for crop or meat production, a major source of man's food. The object of the IBP programme is to learn more about the organic production and overall energy flow of the world's grassland areas. Wherever feasible, concurrent studies with those on natural grasslands will be made on managed areas (pastures and croplands) and successional grasslands in the same region. This will make possible the provision of ecologically sound recommendations for the future utilization of the world's grassland areas.'

The considerable task of drawing up detailed plans and later of ensuring a high degree of coordination among the various projects was performed by a small Working Group. It met for the first time in 1967 under the chairmanship of Dr R. G. Weigert and was attended by scientists from six countries. It was held, appropriately, at Saskatoon, Canada, where, under the direction of Dr R. T. Coupland. a Canadian IBP grasslands team was already at work.

xvii






Foreword Over the period 1967-74 the Biome Coordinating Committee, with various changes in chairmanship, met on fifteen occasions.

In 1972 the Scientific Committee of IBP (SCIBP) appointed Chief Editors who, in association with coordinators or associate editors, were to be responsible for the preparation and completion of the synthesis volumes.

IBP brought together scientists from countries with different types of government; with very different attitudes towards the place of science in their culture; and with very different levels of scientific development. In addition, the acceptance of English as the language of communication within IBP placed an added burden on many scientists throughout the world. These strains were compounded because those of us with ready access to tape recorders, typewriters, secretaries, copying machines, a superabundance of paper and large, if not unlimited, postage accounts, produced a mass of voluminous reports and correspondence that must have added many hours of labour to scientists beyond the English-speaking world. Now, having studied once again many of the reports received during the course of the grasslands studies, I want to place on record the appreciation of the English-speaking contingent to those many workers who had to struggle with a language which lacks the logic of a formalized grammar. Not least among their problems was that of appreciating those subtleties of meaning which, although not clearly defined in a mother tongue, are, through custom and practice, understood by those born to it.

Whilst the basic approaches to scientific discovery may appear universal, the degree of universality tends to be exaggerated and this is certainly the case in a diverse and very often 'diffuse subject such as ecology. Ecology was, and to a considerable extent remains, a highly personalized science. In The Evolution of IBP (IBP Synthesis Series 1, 1975), I quoted a statement of Margalef: 'Ecosystems reflect the physical environment in which they have developed, and ecologists reflect the properties of the ecosystems in which they have grown and matured. All schools of ecology are strongly influenced by a genius loci that goes back to the local landscape.'

Nowhere else in IBP(PT) was the 'ecologist-Iocale'link more in evidence than in the Grasslands Biome Studies. On many occasions the energy expended in arguments on aims, methods and significance of results appeared to be approaching an explosion level but compensating control systems, usually in the form of good humour, succeeded in keeping the 'reactors' below the critical point of disintegration. It is not the first time in the history of biology that heated discussions have paved the way for scientific clarification. IBP productivity studies in particular have been fraught with arguments arising from differences in philosophy, definitions, and the complexity of the organic systems being studied. It is worth recalling a comment by Professors K. Petrusewicz and A. Macfadyen in their preface to Productivity of terrestrial animals (TBP Handbook 13, 1970): 'It has not always been easy to agree on

xviii






Foreword what to say or how to say it. Our discussions have been many and prolonged and the final outcome has usually been based, we believe, on understanding rather than on compromise.' It can certainly be said that discussions within the grasslands theme were many and prolonged.

The USA and Australian IBP grasslands teams were composed largely of scientists whose background and interests centred on domestic animal production. The European, Japanese and Canadian teams on the other hand were less concerned with 'large' consumers and devoted considerable attention to the invertebrate and small vertebrate components of grassland ecosystems. In the early stages ofIBP many European ecologists viewed with some scepticism the wholehearted devotion of their North American confreres to the 'systems approach'. They pointed to many unresolved taxonomic problems; to the paucity of knowledge on the feeding habits of key organisms; to the fragmentary information available on the seasonal changes in the contribution of known species to different trophic levels; and, above all, to the need for a thoroughgoing analysis of production within species or groups of organisms as distinct from obtaining knowledge on yields or standing crops.

I believe that it is important to recall that the operations phase of IBP(PT) reached its maximum in 1971, the year when ecologists should have been celebrating the centenary of the birth of A. G. Tansley who in 1935 added the term ecosystem to the ecologist's vocabulary. Until the advent of the computer and the creative imagination of a new generation of ecologists with the ability to use mathematical techniques to the full, the ecosystem as conceived by Tansley remained a mental construct, valuable and certainly genuine, unlike the Philosopher's Stone, but seemingly beyond total analysis.

Thirty years after Tansley's publication, IBP provided the opportunity for a thorough examination of the interaction of organisms and environment which together, to use Tansley's phrase, formed one physical system. First, the Canadian IBP(PT) Committee, and later the USA National Committee, decided on a systems approach in their programmes, using large multidisciplinary teams. In both cases grasslands were selected as the first ecosystems to be studied intensively, using the still-embryonic ideas of systems ecology.

At the time that IBP(PT) embarked upon a systems approach, it should be remembered that the systems which were to be studied were largely unknown. In engineering and economics, where systems dynamics had become a way of life, the systems could be simulated with a high degree of precision and the relationship between individual components and particular outputs could be explored. The systems ecologist, at the beginning of IBP, was faced with a very different situation. Knowledge of the component parts and processes was fragmentary even in the most thoroughly explored ecosystems. Thus, the systems ecologist was forced to look at his systems in a manner very different

XIX






Foreword

from that followed by his colleagues in aeronautics or in business management.

Natural and cultivated grasslands provide man with the major part of his food supply. They cover at least 23 % of the land surface of the globe. Whilst some parts are becoming desert, elsewhere forests are being felled and, on balance, the grasslands biome is on the increase. Because of its extensive nature and diverse vegetational form, the grasslands biome has straddled three biomes in IBP - grasslands, tundra and arid lands. Scientists from all three biomes met at various workshops to discuss common problems, especially those arising from the utilization of the biomes for grazing. In 1969, largely as a result of the successful application of the systems approach in the IBP Grasslands Biome Studies, it was proposed that grasslands specialists from IBP and FAO should meet to explore how far the knowledge gained from IBP studies could be applied and developed further for use on a world scale. The discussions took place and other international organizations showed interest, particularly UNESCO. A proposal for an International Grazing Lands Programme was approved by SCIBP on behalf of the IBP(PT) biome investigators and it received the support of FAO and UNESCO, but it failed to obtain adequate financial support. Nevertheless the exploratory studies made by the tundra, arid lands and grassland biomes were not completely discarded. The MAB Project No.3 on the Impact of human activities and land use practices on grazing lands: savanna, grassland (from temperate to arid areas), tundra owes much to these earlier discussions initiated by IBP.

The two volumes produced on the IBP grassland investigations can be regarded as independent approaches to the problems of studying grassland ecosystems. One volume, titled Grassland ecosystems of the world (ed. R. T. Coupland), concentrates on the structure, development and utilization of the world's grasslands, largely by producing extensive word models of the types of grassland which exist. The volume ends with a summary of the major components of the world's grassland ecosystems. The other volume, Grasslands, systems analysis and man (ed. A. Breymeyer & G. M. Van Dyne), is a synthesis of the massive amount of data collected during IBP grasslands studies. Its main purpose is to emphasize the dynamic aspects of the grassland ecosystem by giving emphasis to those processes related to productivity. It provides models and sub-models which permit the assessment of the effects of changes or stresses within a grassland ecosystem.

All who have contributed to these volumes are aware of gaps in their attempt to produce a synthesis. The end result is a distillation of numerous points of view and the volumes reflect differences of emphasis, of approach and, above all, disagreements - may it be remembered that disagreements in science are a way of exploring weaknesses and, when accepted in a constructive manner, are the lifeblood of scientific development.

It is always easy to say 'If only it had been possible to .. .'. All of the major

xx






Foreword biome studies were inadequately funded during the synthesis phase. This limited the opportunities to bring editors, chapter authors and contributors together for discussion. Furthermore, some contributors were unable to give enough time to the synthesis operation because they had to become involved full-time in other occupations. Nevertheless, in spite of these and other difficulties the grassland biome studies have advanced our knowledge of grassland ecosystems and they are a major contribution towards the formulation of a theory of ecosystems. These are major achievements. Taken together, they should ensure that the management of the world's grasslands can be placed on a more rational basis than has been possible in the past. The results are especially timely because the world may be facing changes in weather patterns which could well reduce food yields below today's levels. Thus, IBP Theme 1 has made a contribution towards the fulfilment of the overall aim of IBP to investigate The biological basis of productivity and human welfare.

J. B. Cragg Killam Memorial Professor, Faculty of Environmental Design University of Calgary, Canada

xxi






Contents of IBP 18*

List of contributors

Foreword J. B. Cragg

Contents of IBP 19

Part I. Introduction

Background R. T. Coupland

2 The nature of grassland R. T. Coupland

3 Problems in studying grassland ecosystems R. T. Coupland

Part II. Natural temperate grasslands

4 Introduction N. R. French

5 Producers P. L. Sims & R. T. Coupland

6 Consumers M. I. Dyer

7 Micro-organisms E. A. Paul, F. E. Clark & V. O. Biederbeck

8 Systems synthesis R. T. Coupland & G. M. Van Dyne

9 Use and management R. T. Coupland

page xvii

XXI

xxvii

3

23

31

41

49

73

87

97

101

* Grassland ecosystems of the world. ed. R. T. Coupland. Cambridge University Press. 1979.

xxii






Contents of IBP 18

Part III. Semi-natural temperate meadows and pastures Subeditor: M. Rychnovska

10 Introduction 115 E. Balatova-Tu/ackova

11 Primary producers in meadows 127 M. Numata

12 Consumers in meadows and pastures 139 Meadows 139

H. Haas Pastures 147

G. A. E. Ricou

13 Micro-organisms in meadows 155 B. Ulehlova

14 Ecosystem synthesis of meadows 165 Energy flow 165

M. Rychnovska Nutrient cycling 170

A. A. Titlyanova & N. I. Bazilevich

15 Use and management of meadows 181 B. Speidel

Part IV. Tropical grasslands Subeditor: R. Misra

16 Introduction 189 K. C. Misra

17 Primary production 197 J. S. Singh & M. C. Joshi

18 Consumers 219 M. C. Dash

19 Micro-organisms 227 R. S. Dwivedi

20 Ecosystem synthesis 231 J. S. Singh, K. P. Singh & P. S. Yadava

21 Use and management 241 R. K. Gupta & R. S. Ambasht

xxiii






Contents of IBP 18

Part V. Arable grasslands Subeditor: W. M. Willoughby

22 Introduction 247 W. M. Willoughby & P. J. Vickery

23 Producers 253 P. J. Vickery

24 Consumers K. J. Hutchinson & K. L. King 259

25 Micro-organisms 267 R. L. Davidson

26 Nutrient cycling 277 A.R.Till

27 Use, management and conservation 287 W. M. Willoughby & R. L. Davidson

Part VI. Croplands Subeditor: L. Ryszkowski

28 J ntroduction 301 L. Ryszkowski

29 Producers 305 Z. Wojcik

30 Consumers 309 L. Ryszkowski

31 Micro-organisms 319 J. Golebiowska

32 Ecosystem synthesis 327 L. Ryszkowski

Part VII. Conclusion

33 Conclusion 335 R. T. Coupland

References 356

Index 389

xxiv






grasslands, systems analysis and...

Documents