from energetics to...
TRANSCRIPT
From Energetics to Ecosystems:The Dynamics and Structure of Ecological Systems
THE PETER YODZIS FUNDAMENTALECOLOGY SERIES
VOLUME 1
Series Editor
K. S. McCANN
From Energetics to Ecosystems:The Dynamics and Structure
of Ecological Systems
Edited by
N. ROONEYUniversity of Guelph, Canada
K. S. McCANNUniversity of Guelph, Canada
and
D. L. G. NOAKESOregon State University, Corvallis, Oregon, USA
A C.I.P. Catalogue record for this book is available from the Library of Congress.
ISBN-10 1-4020-5336-3 (HB)ISBN-13 978-1-4020-5336-8 (HB)ISBN-10 1-4020-5337-1 (e-book)ISBN-13 978-1-4020-5337-5 (e-book)
Published by Springer,P.O. Box 17, 3300 AA Dordrecht, The Netherlands.
www.springer.com
Printed on acid-free paper
Dedication page by Susan Yodzis� 2007 For Chapter 12 by H. Caswell
All Rights Reserved�2007 SpringerNo 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 theexception of any material supplied specifically for the purpose of being entered andexecuted on a computer system, for exclusive use by the purchaser of the work.
THANK YOU PETER FOR LEADING THE WAY . . .
PETER YODZIS
AT THE ARBORETUM, UNIVERSITY OF GUELPH, CANADA
The water of spring
Has nothing to do
But flow over these rocks.
24.4.77
Designed by Susan Yodzis
TABLE OF CONTENTS
List of Contributors ..................................................................... ixPreface ....................................................................................... xiii
SECTION I
1. A Process-Oriented Approach to the MultispeciesFunctional Response.............................................................. 1Mariano Koen-Alonso
2. Homage to Yodzis and Innes 1992: Scaling upFeeding-Based Population Dynamics to ComplexEcological Networks .............................................................. 37Richard J. Williams, Ulrich Brose and Neo D. Martinez
3. Food Webs, Body Size and the Curse of the LatinBinomial ............................................................................... 53Dave Raffaelli
4. An Energetic Framework for Trophic Control.......................... 65Adrian M.H. DeBruyn, Kevin S. McCann, John C. Moore,Donald R. Strong
SECTION II
5. Experimental Studies of Food Webs: Causes andConsequences of Trophic Interactions...................................... 87Peter Morin
6. Interplay Between Scale, Resolution, Life History andFood Web Properties ............................................................. 101Kirk O. Winemiller
7. Heteroclinic Cycles in the Rain Forest: Insights fromComplex Dynamics................................................................ 127John Vandermeer
8. Emergence in Ecological Systems ............................................ 157James A. Drake, Michael Fuller, Craig R. Zimmermanand Javier G.P. Gamarra
9. Dynamic Signatures of Real and Model Ecosystems.................. 185Peter Yodzis and Kevin McCann
vii
SECTION III
10. Evolutionary Branching of Single Traits................................... 191Junling Ma, Lee Worden, Simon A. Levin
11. Feedback Effects Between the Food Chain and InducedDefense Strategies.................................................................. 213Donald L. DeAngelis, Matthijs Vos, Wolf M. Mooijand Peter A. Abrams
12. Evolutionary Demography: The Invasion Exponent and theEffective Population Density in Nonlinear Matrix Models ......... 237Hal Caswell
13. Of Experimentalists, Empiricists, and Theoreticians .................. 257Neil Rooney
Index ........................................................................................... 261
viii Table of Contents
LIST OF CONTRIBUTORS
Mariano Koen-Alonso
Northwest Atlantic Fisheries Centre
Fisheries and Oceans Canada
80 East White Hills Road
PO Box 5667, St. John’s
Newfoundland, A1C 5X1
Canada
Richard J. Williams
Pacific Ecoinformatics and Computational
Ecology Lab
PO Box 10106, Berkeley
CA 94709
USA
Ulrich Brose
Pacific Ecoinformatics and Computational
Ecology Lab
PO Box 10106, Berkeley
CA 94709
USA
and
Darmstadt Technical University
Department of Biology
Schnittspahnstr
10, 64287 Darmstadt
Germany
Neo D. Martinez
Pacific Ecoinformatics and Computational
Ecology Lab
PO Box 10106, Berkeley
CA 94709
USA
Dave Raffaelli
Environment Department
University of York
Heslington York, YO10 5DD
United Kingdom
Adrian M.H. deBruyn
School of Resource and Environmental
Management
Simon Fraser University
Burnaby, BC
V5A 1S6
Canada
Kevin S. McCann
Department of Zoology
University of Guelph
Guelph, ON
N1G 2W1
Canada
John C. Moore
Natural Resource Ecology Laboratory
Colorado State University
Fort Collins
CO 80523-1499
USA
Donald R. Strong
Department of Environmental
Sciences and Policy
University of California
Davis, CA 95616
ix
Peter Morin
Department of Ecology, Evolution, and
Natural Resources
Rutgers University
14 College Farm Road
New Brunswick, NJ 08901
USA
Kirk O. Winemiller
Section of Ecology and Evolutionary
Biology
Department of Wildlife and Fisheries
Sciences
Texas, A&M University
College Station, TX 77843-2258
USA
John Vandermeer
Department of Ecology and Evolutionary
Biology
University of Michigan
Ann Arbor, MI 48109
USA
James A. Drake
Complex Systems Group
Department of Ecology and Evolutionary
Biology
University of Tennessee
Knoxville, TN 37996
Michael M. Fuller
The Institute for Environmental
Modeling
Department of Ecology and Evolutionary
Biology
University of Tennessee
Knoxville, TN 37996
USA
Craig R. Zimmermann
Department of Biological Sciences
Texas Tech University
Lubbock, TX 79409
USA
Javier G.P. Gamarra
Department of Renewable Resources
University of Alberta
Edmonton, AB
Canada T6G 2H1
Peter Yodzis
Department of Integrative Biology
University of Guelph
Guelph, ON
Canada
Kevin McCann
Department of Integrative Biology
University of Guelph
Guelph, ON
Canada
Junling Ma
Department of Mathematics and Statistics
University of Victoria
Victoria, BC
Canada V8W 3P4
Lee Worden
Department of Environmental Science and
Policy
University of California
Davis, CA 95616
USA
Simon A. Levin
Department of Ecology and Evolutionary
Biology
Princeton University
Princeton, NJ 08544
USA
Donald L. DeAngelis
US Geological Survey and Department of
Biology
University of Miami
Coral Gables, FL 33124
USA
x List of Contributors
Matthijs Vos
Netherlands Institute of Ecology (NIOO-
KNAW)
Department of Food Web Studies
Nieuwersluis
The Netherlands
and
Department of Multitrophic Interactions
Heteren
The Netherlands
Present address:
Netherlands Institute of Ecology
(NIOO-KNAW),
Department of Ecosystem Studies
P.O. Box 140, 4400 AC Yerseke
The Netherlands
Wolf M. Mooij
Netherlands Institute of Ecology
(NIOO-KNAW)
Department of Food Web Studies
Centre for Limnology,
3631 AC Nieuwersluis
The Netherlands
Peter A. Abrams
Department of Zoology
University of Toronto
Toronto, ON M5S 3G5
Canada
Hal Caswell
Biology Department
MS-34 Woods Hole Oceanographic
Institution
Woods Hole, MA 02543
USA
Neil Rooney
Department of Integrative Biology
University of Guelph
Guelph, Ontario
Canada, N1G 2W1
List of Contributors xi
PREFACE
A BRIEF INTRODUCTION: WEDNESDAYS WITH PETER
‘‘If the theory is a true piece of deductive logic—and not just a guess—then theory in advance of
data has the power to help us collect and use data efficiently. The reverse of the coin? Data are
treacherous when offered in support of explanations free of theory. Do not trust such explanations,
no matter how much data seems to support them.’’
Mike Rosenzweig
Not long ago John Vandermeer visited me at McGill. Knowing the severity of
Peter Yodzis’ neurological condition—amyotrophic lateral sclerosis (ALS) or
the Lou Gehrig’s disease—he suggested we organize an academic symposium in
Peter’s honor at the University of Guelph. In a sense John’s suggestion was
similar to the story of Mohammed and the mountain. Peter could not go to a
conference but we could bring a conference to him. I agreed, in principle, but
was paralyzed by such a seemingly monumental organizational task. I did little.
Sometime later I serendipitously related this story to Neil Rooney and David
Noakes. Both listened carefully, agreed wholeheartedly with John, and soon
after, had planned a colloquium and a book that honored Peter and his
significant scientific career. I would like to emphasize that although it was my
wish to see the realization of this colloquium and book, they were undoubtedly
the result of John’s inspiration, and David and Neil’s resourcefulness. Such was
the way the colloquium and book series in Fundamental Ecology started.
It was at this point that David, Neil, and I began to meet every Wednesday
with Peter. Although the intended goal of the meetings was to organize the
colloquium, the thread of our conversations often strayed. Ever the passionate
scientist, Peter also brought to the table a variety of interests, from the Balti-
more Orioles to Haiku poetry, although food seemed to top his list. As such, our
discussions were seemingly light in appearance. This was not quite true though.
These conversations, although light in some aspects, were underlined with the
utmost seriousness—Peter’s was dying and he was dying rapidly. On several
occasions he would let us know this directly and without fear. An extremely
modest but confident man, he wanted to use this last vehicle to make some final
statements to his scientific colleagues and friends. Even the light topics of our
Wednesday meetings, in hindsight, seem to be interlaced with threads of his
personal philosophy on life and science. These same philosophical threads also
permeated my graduate experience with Peter. This book is both a Festschrift
xiii
for Peter and an attempt to bring some of his unwritten scientific views to life. In
what follows, I briefly walk through some of the topics that he quietly and
modestly recapitulated to me and others, time and time again.
MATHEMATICS AND ECOLOGY
Peter was a fearless man. He was never afraid of standing alone and perhaps,
one could argue, sometimes he chose to stand alone. As he frequently put it,
‘‘sometimes one must fight the good fight.’’ He never used this phrase to elevate
himself, but rather to point out his undying belief that a person ultimately must
stand up for what he or she believes. One of those good fights to him was the
formal and rigorous use of mathematics and logic in science.
While studying general relativity theory in physics he saw the power of
simplifying or abstracting complex systems. He firmly believed that ecology
must develop and hone these same analytical and quantitative skills. Not
surprisingly, this belief struck fear into the hearts of almost all undergraduate
biology students at the University of Guelph. On another level, however, there
were the few who truly held him in highest esteem precisely because he
demanded so much of them. He was feared, on average, because he made
them learn theoretical ecology, which meant that biology students must be at
least modestly able to work with some of the basic mathematical tools of
algebra, calculus, and dynamical systems. He was fearless because this does
not win you any popularity contests. That being said, I have met more than one
student who, although being terrified by his course material, spoke volumes of
what Peter actually did for them as ecologists or biologists in the long run.
Peter was disturbed by his belief that ecology was frequently misled simply by
the lack of clarity in thought. As a graduate student of Peter’s, I quickly realized
how thorough he was, as every problem he undertook was subjected to an
almost infinite number of perspectives. Calculation after calculation was neces-
sary as rigor was of utmost importance to him. I also think Peter believed that
slowly immersing oneself in a problem had the interesting tendency to produce
delightful insights. This tendency, he felt, was lost in the push to create volumes
of papers as opposed to fewer more sound papers. As a result of his firm beliefs,
working with Peter was often challenging. He frequently pushed you to explore
further, even when you felt certain that your question had been answered. As we
came to know more about a particular problem, he would begin to raise the
nastiness of the explorative tasks to new heights, as though even greater insights
into a problem demanded a nonlinear increase in workload. As challenging
as this process was, and is, I find myself doing similar things to my own
students now.
Although he held many beliefs firmly, Peter was not resistant to change
as a scientist or a person. This, I imagine, was also part of the ‘‘good fight.’’
Like everyone, certain things never pushed him to change his views but when
something did potentially modify his view of the world he responded, not
xiv Preface
surprisingly, by weeks on weeks of rigorous investigation. Such intense bouts
were occasionally followed by a change in viewpoint, a very well-informed
change of viewpoint indeed. It was likely this ability to change that enabled
him to abruptly abandon theoretical physics for theoretical ecology while he was
in Bern, Switzerland. Peter, tweaked by an environmental essay written by Barry
Commoner in the New Yorker, became engrossed by the problems of the envir-
onment. He felt that mathematical theory and his rigorous set of tools might be
able to play a role in ecological development. He left physics to change the world,
and soon found himself working in the lab of an empirical ecologist (Professor
Hans Burla) who helped educate Peter on ecological theory at that time.
During this time, he found out that other physicists like Robert May and Don
DeAngelis had also switched to ecology and had begun extending the theories of
Alfred Lotka, Vito Volterra, and Robert MacArthur. Although the ecological
culture seemed poised for change at this time (e.g., Robert MacArthur’s com-
petition theories were forefront in ecology), the academic culture of zoology and
botany departments seemed to lag behind. In many departments, a league of
natural historians had gathered and, although they collectively represented an
enormous wealth of biological knowledge, they had little, if any, training in
mathematics. The stage was set for departments to be deeply suspicious of these
new theoreticians. Peter’s story fit this scenario rather cleanly.
I want to emphasize that the University of Guelph showed tremendous vision
to even consider Peter for a position. A good friend and colleague of Peter,
David Lavigne, related to me the story of Peter’s hiring at the University of
Guelph. I imagine that many universities have no such stories simply because
most departments resisted change. This is evidenced by the fact that Peter was
one of a mere handful of theoreticians in Canada for many years. This is not to
say that Peter’s hiring was completely embraced by all members of the depart-
ment, however. In David Lavigne’s words:
One day, one of my colleagues on the search committee came to my office with a letter of reference
received in support of Peter’s application. By this time, if I remember correctly, some of us had
identified him as the best candidate, but others weren’t so sure. Regardless, my colleague handed the
letter to me and said, ‘‘What do you make of this?’’ I read the letter and the sentence I particularly
remember went something like this:
‘‘Peter Yodzis is the best young theoretical ecologist on the market today . . . if you can get him, grab
him immediately, before someone else does.’’
I remember thinking, ‘‘Great letter,’’ and I said so to my colleague.
His reply went something like this: ‘‘Well, yes, it is a very positive letter . . . but who is Robert May?
Have you ever heard of him?’’
This story points out that the hiring of a new breed of ecologist, a mathematical
ecologist, was an uphill battle. It took such brave foot soldiers as Peter to pave
the way for change in ecology, and an open-minded department to provide him
with that opportunity. This is nothing small; ecology has changed dramatically
because of the intellectual curiosity of a fearless few.
Preface xv
The merging of different cultures is rarely seamless and, as ecologists know so
well, trade-offs exist. Like many mathematical theoreticians, Peter’s daily work
habits were at variance with the civil service attitudes that prevailed during the
early part of his career. He tended to get in after lunch and stay late into the
night, his room shining like a beacon across the night campus. When eyebrows
were raised about his peculiar schedule, Peter used to say, somewhat tongue-in-
cheek, ‘‘I’m a theoretical ecologist; I’m always working!’’
More to the point, the sudden prominence of mathematics in ecology also
threatened to divide ecologists into two solitudes. Such division is common in
most scientific disciplines (e.g., see the famous physicist Richard Feynman’s
remarks on those who know mathematics and those who do not, in his lecture,
The Relation of Mathematics to Physics). Mathematical theory may be espe-
cially divisive in a developing science where the cultures have had little time to
develop an ability to communicate. Encouragingly, when Peter first started
teaching a graduate course at the University of Guelph, three or four faculty
actually sat in on the lectures, collected the handouts, and did the assignments.
This was the beginning of Peter’s final ‘‘good fight,’’ to demystify theoretical
ecology for empiricists and so begin a discourse in a way that accelerated
ecological understanding.
OF GEDANKEN EXPERIMENTS AND THEORETICAL ECOLOGY
‘‘In mathematics the art of asking questions is more valuable than solving problems’’
Georg Cantor
If there is one thing I remember most about my meetings with Peter, it is his love
of the Gedanken or thought experiment, which can be explained as the distilla-
tion of a complex problem into a simpler form. Einstein, and physicists in
general, may be the most renowned for the Gedanken experiment. Many
physical laws result from setting up complex problems in an answerable way
(e.g., ‘‘given a world with no gravity then . . . ’’). This simplification, if done well,
brings clarity in understanding, new questions, and ultimately new hypotheses
that speak to an otherwise murky situation. I think, to Peter, the Gedanken
experiment was a thing to be exalted, and it exemplified the things he loved
about the power and beauty of theory.
In a simple sense, the Gedanken experiment has two parts: (1) the simplifi-
cation process itself and (2) the new intuition it brings. The first part—simpli-
fication with controlled loss of information—is ultimately the task of every
scientist investigating problems in a complex world. I recall Peter telling me that
Robert May was a champion of posing problems in a way that made them
answerable and yet shed light on a situation. I would argue that if you study
both Robert May and Peter Yodzis’ work, you will see that a lion’s share of
their brilliance is in the setting up of the problem. In essence, they created useful
mathematical thought experiments. In this light of posing answerable questions,
xvi Preface