Table Of ContentStream
Ecology
EcoIogy
Structure and
function of
running waters
J. David Alian
School of Natural Resources and
Environment, University of Michigan,
USA
lanl
SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.
First edition 1995
© 1995 J. David Allan
Originally published by Chapman and Hall in 1995
Softcover reprint of the hardcover lst edition 1995
Typeset in 10V2/12~pt Sabon by WitweU Ltd, Southport
ISBN 978-0-412-35530-1 ISBN 978-94-011-0729-7 (eBook)
DOI 10.1007/978-94-011-0729-7
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Contents
Preface
Xl
1 Channels and flow 1
Hydrology 1
Characteristics of river channels 12
Summary 20
2 Streamwater chemistry 23
Dissolved gases 24
Major dissolved components of river water 25
The bicarbonate buffer system, alkalinity and hardness 35
Influence of chemical factors on the biota 36
Summary 42
3 Physical factors of importance to the biota 45
Current 45
Substrate 59
Temperature 69
Oxygen 78
Summary 81
4 Autotrophs 83
Periphyton 84
Macrophytes 100
Phytoplankton 103
Summary 107
5 Heterotrophic energy sources 109
Decomposition of coarse particulate organic matter 109
Fine particulate organic matter 118
Dissolved organic matter 121
Summary 129
6 Trophic relationships 131
The microbial loop 132
Invertebrate consumers 134
Feeding ecology of riverine fishes 150
vii
Contents
Lotic food webs 159
Summary 159
7 Predation and its consequences 163
Choice and vulnerability 163
Predator control of prey distribution and abundances 176
Summary 185
8 Herbivory 187
Periphyton-grazer interactions 187
Herbivory on macrophytes 200
Grazing on lotic phytoplankton 201
Summary 202
9 Competitive interactions 205
Distributional patterns and resource partitioning 206
Experimental studies of competition 213
Summary 218
10 Drift 221
Composition and periodicity 221
Drift and downstream displacement 224
Functional basis of drift 229
Summary 237
11 Lotic communities 239
Local and regional diversity 239
Community structure 248
Summary 256
12 Organic matter in lotic ecosystems 259
Dynamics of dissolved and particulate organic matter 261
Fate of dissolved and particulate organic carbon 265
Qrganic matter budgets 270
The river continuum concept 276
Summary 281
13 Nutrient dynamics 283
Basic principles of nutrient cycling 285
Nutrient concentrations in running waters 288
Transport and transformation of nutrients 293
Summary 303
14 Modification of running waters by humankind 305
Brief history of river modification 305
Dams and impoundments 308
Transformation of the land 320
vii
Alien species 330
Climate change 335
Imperilment of the biota 338
Recovery and restoration of running waters 339
Summary 341
References 343
Index 379
Ix
Preface
When my general interest in aquatic ecology of river systems that brings a unifying frame
began to focus increasingly on streams, imme work to this area of study. The inputs and trans
diately after completing my PhD, it was my good formation of energy and materials are important
fortune that H.B.N. Hynes classic, The Ecology in all river systems, regional species richness and
of Running Waters, had just appeared. Many of local species interactions influence the structure
the current generation of stream ecologists bene of all riverine communities, and the interaction
fitted from that masterful survey of the field. of physical and biological forces is important to
During the 1970s and 1980s, ecological studies virtually every question that is asked. It appears
of streams and rivers made many advances. to be the processes acting in running waters that
Major new paradigms emerged, such as the river are general, and the settings that often are
continuum concept and nutrient spiralling. Com unique.
munity ecologists made impressive advances in As is true for every area of ecology during the
documenting the occurrence of predation, com closing years of the twentieth century, the study
petition and other species interactions. The of streams and rivers cannot be addressed exclu
importance of physical processes in rivers drew sive of the role of human activities, nor can we
increasing attention, particularly the areas of ignore the urgency of the need for conservation.
hydrology and geomorphology, and the inter This is a two-way street. Ecologists who study
relationships between physical and biological streams without considering how past or present
factors became better understood. While each human modifications of the stream or its valley
advance in knowledge has opened new ques might have contributed to their observations,
tions, the progress made in the past 20 years has do so at the risk of incomplete understand
been considerable, and has formed the basis for ing. Conservation efforts that lack an adequate
continuing and future work. In this volume I scientific basis are less likely to succeed. One
attempt to summarize these developments for trend that seems safe to forecast in stream eco
the current generation of students. logy is toward a greater emphasis on understand
The diversity of running water environments ing human impacts. Fortunately, signs of this
is enormously broad. When one considers tor trend are already apparent.
rential mountain brooks, large rivers of low I cannot pretend to do justice to the entire
lands, and great rivers whose basins occupy range of ecological topics pertaining to streams
subcontinents, it is apparent how location and rivers. The diversity of running water
specific environmental factors contribute to the environments, the richness of taxa from micro
sense of uniqueness and diversity of running bes to higher vertebrates, and the range of topic
waters. At the same time, however, our improved areas all are too great. The reader will surely
understanding of ecological, hydrological and detect biases towards studies based in small
geomorphological processes provides insight streams, at middle or higher latitudes, and
into the functional and structural characteristics reported in the English-language literature,
xl
Preface
although I have done my best to resist these drawings for figures. The stream group at the
tendencies. I greatly appreciate the construc University of Maryland was a continual source
tive suggestions of many colleagues who were of ideas during my tenure there. I began writing
generous in their advice on draft chapters of the this book while a sabbatical visitor at the
book. They have done much to clarify and cor University of Lund, and benefitted greatly from
rect what I have written, but the responsibility my interactions with Bjorn Mamlqvist, Per
for choice of topics as well as any errors or omis Sjostrom, Christian Otto and other Swedish
sions is mine. Scott Cooper, Bobbi Peckarsky colleagues. To all of these individuals I extend
and Scott Wissinger used an early draft of the my deepest appreciation. I also wish to thank
entire manuscript in their stream ecology courses those people who helped with many of the details
and provided valuable feedback from their of manuscript preparation. Tara Burke and
students as well as their own reactions. Indiv Cindy Smith helped to compile and prepare
iduals whose efforts improved individual chap figures and Mary Lammert and Robin Abell
ters include Fred Benfield, Art Benke, Art were most careful and efficient editorial
Brown, Scott Cooper, Alex Flecker, Nancy assistants. I thank my editors at Chapman &
Griffith, David Hart, Chuck Hawkins, Bob Hall, Bob Carling and Clem Earle, for their
Hughes, Steve Kohler, Gary Lamberti, Rex combination of patience and prodding.
Lowe, Rich Merritt, Diane McKnight, Judy Lastly, I wish to express my deepest apprecia
Meyer, Bobbi Peckarsky, Pete Ode, Walt tion to my extremely supportive and tolerant
Osterkamp, M.L. Ostrofsky, Margaret Palmer, family, Susan, Jennie and Brian. Their encour
LeRoy Poff, Karen Prestegaard, Ike Schlosser, agement has been a constant throughout my
Len Smock, Al Steinman, Scott Wissinger and career, and I especially appreciate their help and
Jack Webster. I thank Ken Cummins, AIen understanding during the time this book was
Flecker, Jim Gore and AI Steinman for providing being written.
xl
Chapter one
Channels and
flow
Our first impressions, when we gaze upon a river, 1.1 Hydrology
are of the strength of the current, the dimensions
of the channel, and perhaps the boulders in the
1.1.1 The water cycle
streambed or the shape of the banks. Mentally we
catalog this flowing water system as a stream or Until the sixteenth century, oceans were thought
a river, in flood or in repose, and perhaps take to be the source of rivers and springs via under
note of bends, pools and meanders. Very likely ground seepage. Palissy and others suggested
we have encountered other streams with similar that storage of rain-water was the real source,
features, and some that are very different. We based on several lines of reasoning. It was
might also note whether this waterway is suitable suggested that springs would not dry up in
for various activities, such as the passage of summer if oceans were the source, since the
boats, recreational uses and whether it poses a oceans do not decrease noticeably. Springs
hazard to humans. With a moment's further should be more common at low elevations
reflection it is apparent that these same channel if they derive from oceanic water. However
and flow characteristics likely influence the func springs often dry up in summer, they are more
tioning of running water ecosystems, and the common on mountain slopes, and finally,
biota found therein. springs are fresh. In 1674, measurements by
Perrault showed that precipitation into the Seine
Physical processes acting on river systems basin was six times greater than discharge
traditionally have been the domain of hydro (Morisawa, 1968). This finding changed the
logists and geomorphologists. It is increasingly focus of interest from studying whether rainfall is
apparent, however, that our understanding of sufficient to provision rivers, to studying where
the ecology of rivers will benefit from an appre the rest of the rainfall goes.
ciation of flow variability, channel structure The hydrologic cycle describes the continuous
and other physical aspects of river ecosystems. cycling of water from atmosphere to earth and
Running waters are enormously diverse. They oceans and back again (Figure 1.1). Con
range from small streams to great rivers, ceptually this cycle can be viewed as a series of
and occur under widely differing conditions of storage places and transfer processes, although
climate, vegetation, topography and geology. water in rivers is both a storage place, however
In order to make sense of biological findings temporary, and a transfer between land and
from such disparate settings, it is important to sea. The hydrologic cycle is powered by solar
have a framework that reflects the physical energy. This drives evaporation and evapotran
dimensions of the study system. spiration, transferring water from the surface of
1
