Table Of ContentBryophyte Ecology
Bryophyte Ecology
Edited by A.J .E. Smith
LONDON NEW YORK
CHAPMAN AND HALL
First published 1982 by Chapman and Hall Ltd
11 New Fetter Lane, London, EC4P 4EE
Published in the USA by Chapman and Hall
in association with Methuen, Inc.
733 Third Avenue, New York, NY 10017
© 1982 Chapman and Hall
Softcover reprint ~fthe hardcover 1st edition 1982
Photoset by Enset Ltd
Midsomer Norton, Bath, Avon
All rights reserved. No part of
this book may be reprinted, or reproduced
or utilized in any form or by any electronic, mechanical
or other means, now known or hereafter
or in any information storage or retrieval
system, without permission in writing
from the publisher.
British Library Cataloguing in Publication Data
Bryophyte ecology.
1. Bryophytes
I. Smith, A.l.E.
588 QK533.7
ISBN-13: 978-94-009-5893-7 e-ISBN-J3: 978-94-009-5891-3
DOl: 10.1007/978-94-009-5891-3
Contents
Contributors Vll
Preface 1X
1 Quantitative Approaches in Bryophyte Ecology 1
l.W. Bates
2 Life-forms of Bryophytes 45
K. Magdefrau
59
3 Tropical Porest Bryophytes
T. Pocs
4 Desert Bryophytes 105
G.A.M. Scott
5 Bryophyte Vegetation in Polar Regions 123
R.E. Longton
6 Alpine Communities 167
Patricia Geissler
7 Epiphytes and Epiliths 191
A.l.E. Smith
8 The Ecology of Sphagnum 229
R.S. Clymo and P.M. Hayward
9 Bryophytes and Invertebrates 291
Uri Gerson
10 Physiological Ecology: Water Relations, Light and 333
Temperature Responses, Carbon Balance
M.C.P. Proctor
vi Contents
11 Mineral Nutrition 383
D.H. Brown
12 Responses of Bryophytes to Air Pollution 445
Dhriva N. Rao
13 Quaternary Bryophyte Palaeo-ecology 473
H.J.B. Birks
Author Index 491
Subject Index 505
Contributors
J.W. Bates, Department of Pure and Applied Biology, Imperial
College Field Station, Silwood Park, Ascot, Berks., SL5 7PY, Great
Britain.
H.J.B. Birks, Botany School, Downing Street, Cambridge, CB2
3EA, Great Britain.
D.H. Brown, Department of Botany, University of Bristol,
Woodland Road, Bristol, BS81UG, Great Britain.
R.S. Clymo, Department of Botany and Biochemistry, Westfield
College, Kidderpore Avenue, London, NW3 7ST, Great Britain.
Patricia Geissler, Conservatoire et Jardin Botaniques, Case Postale
60, CH-1292 Chambesy/Geneve, Switzerland.
Uri Gerson, Faculty of Agriculture, Hebrew University of
Jerusalem, Rehovot, Israel.
P.M. Hayward, Department of Botany and Biochemistry, Westfield
College, Kidderpore Avenue, London NW3 7ST, Great Britain.
R.E. Longton, Department of Botany, Plant Science Laboratories,
The University of Reading, Whiteknights, Reading, RG6 2AS,
Great Britain.
K. Magdefrau, 8024 Deisenhoffen bei Munchen, Waldstrasse 11,
Germany.
T. Pocs, Research Institute for Botany of the Hungarian Academy of
Sciences, 2163 Vacratot, Hungary.
M.C.F. Proctor, Department of Biological Sciences, University of
Exeter, Hatherly Laboratories, Prince of Wales Road, Exeter,
EX4 4PS, Great Britain.
D.N. Rao, Department of Botany, Banaras Hindu University,
Varanasi-221005, India.
viii Contributors
G .A.M. Scott, Botany Department, Monash University, Clayton,
Victoria, Australia.
A.J .E. Smith, School of Plant Biology, University College of North
Wales, Memorial Buildings, Bangor, LL57 2UW, Great Britain.
Preface
There has been an increasing interest in bryophyte ecology over the
past 100 or so years, initially of a phytosociological nature but,
additionally, in recent years, of an experimental nature as well. Early
studies of bryophyte communities have led to detailed investigations
into the relationships between the plants and their environment.
Ecological papers, the large number of which is evidenced by the
length of the bibliographies in the subsequent chapters, have
appeared in numerous journals. Yet, apart from review chapters, by
H. Gams and P.W. Richards in Manual of Bryology, edited b:; H.
Verdoorn in 1932 and chapters in E.V. Watson's Structure and Life of
Bryophytes, Prem Puri's Bryophytes - A Broad Perspective and
D.H.S. Richardson's The Biology of Mosses, published in 1972,1973
and 1981 respectively, no general accounts of bryophyte ecology
have been published.
Although the Bryophyta is a relatively small division of plants,
with between 14000 and 21000 species the interest that they have
aroused is out of all proportion to the size either of the plants or of the
division. It is evident, however, that despite their relative insigni
ficance they play an important ecological role, especially in extreme
environments and, in the case of bryophytes in tropical cloud forests
and of Sphagnum, may even be a dominant factor in the ecology of
the area concerned. Analysis of bryophyte communities presents
particular problems; aspects of their physiological ecology, especially
drought resistance and nutrition, are topics of special interest, and
the recent upsurge of interest in atmospheric pollution has revealed a
new aspect of the relevance of the group.
In attempting to edit a book on bryophyte ecology one is limited by
space and the availability of appropriate and willing authors. Hence it
has not proved possible to cover all aspects of the subject but it is
x Preface
hoped that a sufficient diversity of topics has been dealt with to
provide an adequate overview. Some of the chapters are general
reviews, others are more detailed accounts of the authors' own
research interests. In either case it is evident that there remains a
great deal of research to be done on all aspects of bryophyte ecology
and it is hoped that this volume, as well as providing an account of
what is known, will stimulate further study.
Chapter 1
Quantitative Approaches
in Bryophyte Ecology
1.W.BATES
1.1 INTRODUCTION
Studies of bryophyte vegetation are normally undertaken to answer one or
more of the following questions:
(1) Which species occur and in what quantities and spatial arrangement?
(2) How does community structure vary with variation in the intensities of
environmental factors?
(3) How do different species manage to co-exist in the community?
(4) How does the community maintain itself through time given the avail
able resources?
These four topics, community structure, community vanatIOn, species
strategies and community function, form a series which poses increasing
technical difficulties to the would-be investigator. The difficulties are
increased by certain peculiarities of bryophytes themselves; their small
size, the often fragmentary nature of their communities, the tendency of
many species to grow on highly irregular surfaces, their frequent presence
as relatively low biomass components in communities dominated by other
types of vegetation and their lack of economic importance leading to
restricted financial support for research. It is, therefore, perhaps not
surprising that, although simple descriptive accounts of particular bryo
phyte communities are reasonably common in the literature, relatively few
quantitative studies of correlation between vegetation and environmental
factors have been undertaken. More sophisticated studies of functional
and strategic aspects of bryophyte communities are scarce.
The history of bryophyte ecology, in common with other branches of
ecology, shows an increasing tendency towards the use of an objective,
quantitative approach. This must be regarded a~ a healthy sign and results
1
Description:There has been an increasing interest in bryophyte ecology over the past 100 or so years, initially of a phytosociological nature but, additionally, in recent years, of an experimental nature as well. Early studies of bryophyte communities have led to detailed investigations into the relationships b
