Table Of ContentArne Pommerening · Pavel Grabarnik
Individual-
based Methods
in Forest
Ecology and
Management
Individual-based Methods in Forest Ecology
and Management
Arne Pommerening Pavel Grabarnik
(cid:129)
Individual-based Methods
in Forest Ecology
and Management
123
Arne Pommerening PavelGrabarnik
Department ofForest Ecology Institute of Physicochemical andBiological
andManagement Problems in SoilScience
SwedishUniversityofAgriculturalSciences Russian Academy of Sciences
Umeå,Sweden Pushchino, Russia
ISBN978-3-030-24527-6 ISBN978-3-030-24528-3 (eBook)
https://doi.org/10.1007/978-3-030-24528-3
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Sic ergo quaeramus tamquam inventuri,
et sic inveniamus tamquam quaesituri.
Let us therefore search as one who expects to
find, and find as one who is determined to
search further.
Augustine of Hippo, De Trinitate
Foreword
Forests are vast assemblages of trees, understory plants, large and small animals,
andcomplexbioticsoils.Thecomposition,structure,anddynamicsofforestsresult
from interacting ecological processes across gradients of space and time. The vast
amount of information embodied in a forest is almost incomprehensible.
The early development offorest science and forest management was based on
only a very small fraction of the total information residing in forests. Trees were
aggregated into stands, and stands were represented by collective traits such as
percent species composition and tree sizes. Rates of change in stocking (trees/ha)
andstemvolumeormasswerecalculatedfrominformationonstructuralchangesof
standsbetweensamplingperiods.Theuseofinformationtocharacterizestandswas
sometimes expanded by characterizing variances around mean values, and by
characterizing distributions among tree size classes. This level of information
supported more than a century of insights into forests and how they change from
oneperiodtoanother,andacrosssites.Differencesingrowthratesacrosssiteswere
classified into site productivity classes, based on information such as heights of
dominanttreesatagivenage.Familiarsynthesesofinformationincludedsiteindex
curves(treeheightinrelationtoage),andtablesofstandvolumesandyields.Inthe
second half of the last century, computerization offorest science and management
greatly expanded the insights that could be gleaned from information present in
forests.
Curiously, all of the processes that lead to the amalgamated stand-level char-
acteristicsofforestsdonothappenatthelevelofstands.Standgrowthoccursatthe
scale of individual trees, and a stand-level representation of forest growth misses
many of the information present in tree-level processes. For example, changes in
stand-level growth as forests develop over decades result from time trends in the
growth ofindividual trees.Arising(or declining) rate ofstand-levelgrowth might
result from a synchronous trend among all trees, or from the net outcome of trees
showing opposing growth trends. Some of this information was used in classic
representation of dominant and suppressed trees, and silvicultural approaches used
this information to compare stocking regimes that periodically removed small or
large trees.
vii
viii Foreword
A major leap in the use of information from trees and forests developed late in
thetwentiethcenturyasmethodsofsamplingandhandlingdataexpandedbyorders
of magnitude. The quantitative leap in the accounting for growth of thousands of
individual trees was joined by spatial information on tree locations. This combi-
nationofdataonmanyindividualsatdefinedlocationsprovidedalensforseeinga
new dimension of forest information: relationships between individual trees. The
growth rate of a given tree could be evaluated not only in terms of environmental
factors (such as the supply of soil water and nutrients) but also in relation to
competition for resources with neighboring trees. Dominance could be considered
in a dynamic sense, accounting for different rates of resource utilization among
trees, the efficiencies of using resources to support growth, and how the“story” of
each tree’s change over time depended on interactions with the stories of neigh-
boring trees.
The examination of individual-tree dynamics in a spatially explicit context
enabledscientistsinecologyandforestrytoexploreandtestideastoexplainforest
structureanddynamics.Advancesinsciencebegantoexplaintheprocessesbehind
the stand-level patterns that had long been the focus of ecology and forest man-
agement. But could analyses of growth of individual trees, and neighborhoods of
trees,beharnesseddirectlytoimproveforestanalysis,planning,andoperations?A
decade ago, the answer to this question was a nebulous “we hope so, someday”.
Now the answer is an emphatic “yes”.
Theforestsofourfuturewillnotrepeatthepatternsofforestsinthepast.Human
interactionswithforestsarenowpervasivearoundtheglobe.Forestcompositionis
managed directly by fostering some species over others, and indirectly by modi-
fying factors such as populations of tree herbivores. The success of species may
now depend on invasions from across the globe, including pathogens that reshape
interactions among tree species across landscapes and regions. The changing cli-
mate of the twenty-first century will have ecological effects of a magnitude that
rivals moving southward by a hundred kilometers or more. The impacts of winds,
fires, diseases, and herbivores will have novel impacts under future climates.
A retrospective, stand-level view of how forests grew in the past would lack vital
information about factors shaping future dynamics of trees and forests. Our ability
to envision and explain the forests of our future depends on a foundation of
dynamics at the level of trees (and neighborhoods). This book develops the con-
cepts, information, and tools for building such a foundation.
Flagstaff, Arizona Dan Binkley
May 2019
Preface
Individual-based forest ecology and management, where thefocus ison individual
plants in the context of their populations, draws on a large body of different
quantitative methods that constantly grows and is not well documented in a com-
prehensive way. This book is intended to serve as a text and reference for under-
graduate and postgraduate courses in forest ecology and management. It combines
and links the synthesis of important theories and concepts of these two academic
fields with new quantitative principles which—although they form a natural and
logical unit—so far have not been published together in a single volume.
This book has many boundary encounters in stock. It is about interdisciplinary
research situated between ecology, forest science, statistics and computing and
facilitates knowledge exchange between different disciplines and research com-
munities.Thetextattemptstoadvanceforestscienceinthefieldsofforestecology
and management through quantitative, individual-based methods and at the same
timeprovidesincentivestoadvancemathematicalstatisticsthroughaconsideration
of applications in forest ecology and management.
Inatimewherefirmboundariesbetweenacademicfieldsarerapidlyfading,itis
animportantfeatureofthebooktochallengeandre-interprettraditionalconceptsof
forest ecology and management through the findings in tree interaction and forest
structure research that we and others have worked on during the last 25 years.
Therefore,thisbookpresentsanewvisionofinterdisciplinary,quantitativeresearch
and deliberately abandoned the structure of traditional textbooks in forest science
and ecology. The text is also instrumental in streamlining the vast amount of
localized theories and methods by presenting them in a new, more concise and
coherent system.
The book is designed as an English language textbook for a wide range of
countries without any particular geographic reference. This textbook stresses the
theoretic principles of forest ecology and management and generic quantitative
methods which can easily be adapted to different local situations. It is ideal for
classes of mixed nationalities with English as the medium of teaching and can
therefore be used almost anywhere in the world. Such classes of mixed language
ix
x Preface
anddifferentacademicbackgroundsareontheincreaseparticularlyaspartofM.Sc.
and Ph.D. courses as well as summer schools.
The book should also appeal as a reference to researchers and is useful as a
numerical handbook for practitioners and consultant ecologists or foresters. An
effort has been made to prepare the text to be comprehensible also for
non-specialists willing to do some limited additional reading.
Chapter1introducesindividual-basedforestecologyandmanagement,explains
the vision of this book, and thus sets a common frame.
Chapter2synthesizesimportantecologicaltheoriesthatformthefoundationand
the frame of individual-based forest ecology and management.
Chapter 3 reviews important generic concepts and techniques of ecology-based
silviculture that are pre-requisites of individual-based forest management.
Chapter4isakeypartofthebookexplainingspatialmethodsoftreeinteraction
researchrequired inindividual-based forest ecologyand management. The chapter
particularlyfocusesonmethodsofpointprocessstatisticsfortheanalysisofspatial
plant patterns.
Chapter5servesasanintroductiontospatialmodellingasanaturalcontinuation
ofpointprocessstatisticsoutlinedinChap.4.Weintroduce here methodsofpoint
process modeling, spatial reconstruction, and individual-based models.
Chapter 6 presents tree and forest growth analysis and modelling with a par-
ticular focus on relative growth rates. This is an important pre-requisite of
individual-based research.
Chapter 7 is dedicated to the analysis of individual human decision making in
forests and how the selection of individual trees impacts on forest structure. The
chapter outlines theory and concepts of disturbance analysis and the analysis of
human tree selection behaviour.
The appendices provide additional background material on forest ecology and
management and computational pre-requisites that may not be known to
non-specialists.
A book on quantitative forest ecology and management is only complete if
explored in practical computer exercises. For this purpose, we integrated quanti-
tative concepts and computations. In our own experience, using computer code in
classesgreatlyenhancestheunderstandingandlearningofquantitativeconcepts,as
students acquire both the concepts and programming through their own experi-
mentation at the same time. In this book, we selected the R language but other
programming languages would have been suitable as well. Thus, R accompanies
thereaderthroughoutthebookasavirtuallaboratorywithawealthofpre-designed
packages and possibilities stimulating self-learning. Finally, the included R code
allowsthereadertoreproducepresentedexamplesandtoexpandormodifythemto
beusefulforher/hisowndataanalyses.Inordertogetthemostoutofthisbook,it
ishighlyrecommendedtoexploretheconceptsintroducedinthetextbyusingtheR
software as a virtual laboratory. The code listings are provided directly following
mathematicalequationssothatthecodeoffersanotherpointofaccessinadditionto
the sign language of mathematics.
Preface xi
Good research does not unfold without inspiration and intergenerational
exchangeofideas.Afterall,everynewgenerationofresearchersnaturallystandson
the shoulders of the previous generation as Stephen Hawking put it, based on an
expression by Isaac Newton and earlier metaphors. In this spirit, the first author
gratefully wishes to acknowledge his academic mentors Klaus von Gadow
(Göttingen),HannsH.Höfle(Göttingen), HansPretzsch (München), Jean-Philippe
Schütz (Zürich), Hubert Sterba (Wien), Dietrich Stoyan (Freiberg), and Günter
Wenk(Dresden),whoexcited himwith different aspects ofindividual-basedforest
ecologyandmanagementatthebeginningofhisresearch career andprovidedhim
withathoroughscientificeducation.Forthemanylivelyandinspiringdiscussions,
we cannot thank them enough. Their enthusiasm and continuous ready support
throughout the years, even from the comfort of their retirement, have been a great
assetandaprivilege.Thisworkowesalottothemanyfruitfuldiscussionswehad.
The mutually beneficial knowledge transfer from statistics to ecology and forest
science and vice versa has had a significant positive impact on the lives of both
authors and with this book we wish to pass on this wonderful experience.
Many good ideas, teaching materials, and computer codes were also developed
during (guest) professorships in Austria, China, Denmark, Estonia, Finland,
Germany, Ireland, Poland, Spain, Sweden, Switzerland, the UK, and in the US.
Discussionswithstudentsandcolleaguesinthesecountrieshavehadaconsiderable
influence on shaping the text. It is to this international origin that the book owes
much of its spirit.
Finally and most importantly, we wish to extend our gratitude to a number of
colleagues, friends, and family members who provided helpful suggestions and
review comments for various parts of this text including Uta Berger, Harold
Burkhart,CésarPérez-Cruzado,RoqueRodríguez-Soalleiro,HongxiangWangand
Zhonghua Zhao. Christopher Guest kindly reviewed Appendix A, and Francis
GwynJonesandJensHaufecontributedfeedbacktoAppendixB.Naturally,forall
remaining errors or shortcomings in this work, we alone assume responsibility.
Discussions with Dan Binkleyduringhis extendedWallenberg visit toUmeåhave
inspired many reflections on individual-based forest ecology and management and
consequently he kindly contributed the foreword to this book.
Umeå, Sweden Arne Pommerening
Pushchino, Russia Pavel Grabarnik
May 2019
Acknowledgements A. P. acknowledges the support of the Swedish University of Agricultural
Sciences (SLU) and particularly of the SLU Department of Forest Ecology and Management at
Umeåwhilepreparingthebookmanuscript.ThecontributionofP.G.wasfundedbytheRussian
ScienceFoundation(projectno.18-14-00362).
