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Cover photographs:

(1) Sand dunes in Death Valley, California (Photo by Ruth and Louis Kirk).
(2) Forest and agriculture by Richmond, Virginia (USDA Soil Conservation
Service). (3) Bull reindeer on western Alaska tundra (D. R. Klein).

Copyright © 1986, by John Wiley & Sons, Inc.

All rights reserved. Published simultaneously in Canada.

Reproduction or translation of any part of

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or further information should be addressed to
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Library of Congress Catakil}iflg in fublication Data:

Forman, Richard T T.
Landscape ecology.

Includes index.
1. Ecology. 2. Landscape protection.
3. Hun1an ecology. I. Godron, Michael. II. Title.
QH541.F67 1986 712 85-12306
ISBN 0-471-87037-4

Printed in the United States of America

10 9 8 7 6 5 4 3 2
 Contents

Part I.
Overview
1. Landscape and Principles 3
PERCEPTIONS OF THE LANDSCAPE 4
Breadth of Concepts. The Artist's Landscape. The Concept in the
Social and Natural Sciences.
A LANDSCAPE FROM AN ECOLOGICAL PERSPECTIVE 8
What Makes a Landscape? A Usable Scientific Concept. Landscape
Elements and Tesserae. Related Concepts.
CONCEPT AREAS AND PRINCIPLES OF LANDSCAPE ECOLOGY 17
Diversity Among Landscapes. Concept Areas. Emerging
Genera'I' Principles.
THE EMERGENCE OF LANDSCAPE ECOLOGY:
MAJOR LITERATURE 28
Geography and Ecology. Related Disciplines. International
Perspectives Today.
SUMMARY 31
QUESTIONS 31

2. Ecological Concepts in Brief 33

THE PHYSICAt ENVIRONMENT 35
Climate. Soil. Fire.
AQUATIC ENVIRONMENTS 44
Hydrologic Cycle. Lotic and Lentic Environments.
Marine Environments.
POPULATIONS AND ｔｈｅｾｒ REGULATION 49
Populations and Growth. Competition. Predation.
Population Regulation.

XIII
 57 STRIP CORRIDORS 142
EVOLUTIONARY ECOLOGY Corridors Lower Than Surroundings. Corridors Higher
Variation and Selection. Speciation. Than Surroundings.
60
ECOLOGICAL COMMUNITIES STREAM CORRIDORS 146
Spatial Structure of a Community. Ecological Niche and Species SUMMARY 153
Diversity. Succession. Island Biogeography. Biogeography in
QUESTIONS 155
Geologic Time.
68
ENERGY AND MAnER IN ECOSYSTEMS
Energy. Plant and Animal Production. Food Chains and Webs.
Mineral Nutrient Cycling. Ecosystem Models. 5. Matrix and Network 157
77 DISTINGUISHING A MATRIX 159
SUMMARY
78 Three Matrix Criteria. How the Matrix Criteria Influence Change: The
QUESTIONS Desertification Example.
POROSITY AND BOUNDARY SHAPE 168
The Concept and Its Implications. Porosity Patterns in Different
Part 1/.
Landscapes. Boundary Shape.
Landscape Structure
NETWORKS 178
Chapter 3. Patches 83 Intersections in the Network. Reticulate Pattern. Mesh Size. Factors
83 Determining Network Structure.
PATCH ORIGINS AND CHANGE
Basic Concepts. Disturbance Patches. Remnant Patches. MATRIX HETEROGENEITY 184
Environmental Resource Patches. Introduced Patches. SUMMARY 186
Additional Concepts.
98 QUESTIONS 186
PATCH SIZE
APPENDIX: MEASURES OF PATCH CHARACTERISTICS
Effects on Energy and Nutrients. Effects on Species.
IN A MATRIX 188
106
PATCH SHAPE
Significance in Ecology. Edge Effect. Isodiametric and Elongated
Patches. Rings. Peninsulas. 6. Overall Structure 191
117
PATCH NUMBER AND CONFIGURATION MICROHETEROGENEITY AND MACROHETEROGENEITY 194
119
SUMMARY Analysis of a Microheterogeneous Landscape. Analysis of a
120 Macroheterogeneous Landscape.
QUESTIONS
" CONFIGURATION OF PATCHES, CORRIDOR, AND MATRIX 204
Chapter 4. Corridors 121 A Few Distinctive Configurations. Determining
122 Distinctive Configurations.
CORRIDORS AND THEIR ORIGINS
125 CONTRAST IN THE LANDSCAPE 211
CORRIDOR STRUCTURE
The Vulture's View. The Hiker's View. Microenvironment Low-Contrast Structure. High-Contrast Structure.
Within a Corridor. GRAIN SIZE OF THE LANDSCAPE 216
131
LINE CORRIDORS ADDITIONAL STRUCTURAL CONSIDERATIONS 218
Hedgerows as Line Corridors. Animal Communities of Hedgerows.
SUMMARY 220
Other Types of Line Corridors.
CONTENTS xv
xiv CONTENTS
 220 9. Flows Between Adjacent Landscape Elements 313
QUESTIONS
MECHANISMS UNDERLYING LINKAGES 315
APPENDIX: A METHOD FOR MEASURING 222 Vectors. Forces.
LANDSCAPE HETEROGENEITY
AIRFLOW AND LOCOMOTION 320
, Wind Patterns. Sound. Gases, Aerosols, and Particulate
Matter. Locomotion.
SOIL FLOWS 335
Part 11/.
Landscape Oynamics General Concepts. Erosion and Surface Flow. Subsurface Flow.
INTERACTION BETWEEN LAND AND STREAM 342
7. Natural Processes in Landscape Development 229 The Stream Corridor Role. Corridor Filtration of Mineral Nutrients.
230
GEOMORPHOLOGY HEDGEROW INTERACTIONS WITH ADJACENT
Landforms and Their Effects. River Systems. Bedrock, Climate, and LANDSCAPE ELEMENTS 347
Stasic Processes. Geomorphic Processes Under Different Climates. How Hedgerows Affect Fields. How Fields, Woods, and Homes
241 Affect Hedgerows.
ESTABLISHMENT OF LIFE FORMS
Plant Establishment. Animal Establishment. Speciation in Landscapes. SUMMARY 354
Increase in Vegetation Stature. QUESTIONS 354
259
SOIL DEVELOPMENT
Law of Zonality. Toward a Soil Classification Based on Origin and 10. Animal and Plant Movement Across a Landscape 357
Development. The Role of Decomposers.
265 PATIERNS OF MOVEMENT 357
NATURAL DISTURBANCE Continuous Movement. Saltatory Movement.
268
SUMMARY MOVEMENT OF ANIMALS 361
269
QUESTIONS Home Range, Dispersal', and Migration. Barriers and Conduits
270 Perceived by Different Species. Patterns Based on the
APPENDIX: GEOLOGICAL TIME
Radiotracking Studies.
MOVEMENT OF PLANTS 381
8. The Human Role in Landscape Development 273
274 The Cheatgrass Invasion of the American West. Australian Eucalypt
MODIFICATION OF NATURAL RHYTHMS versus California Pine. Forest Species in Hedgerows.
Disturbance and Rhythms. Daily Rhythms. Seasonal Rhythms.
SOME SPECIES MOVEMENTS IN AGRICULTURE
Rhythms of Several Years or Centuries.
281 AND PEST CONTROL 392
METHODS OR TOOLS USED IN LANDSCAPE MODIFICATION
SUMMARY 394
Natural Resource Extraction and Alteration. Introduction of
Agricultural Methods. Decision Catalysts.
QUESTIONS 394
286
A LANDSCAPE MODIFICATION GRADIENT
Natural Landscapes. Managed Landscapes. Cultivated Landscapes. 11 ｾ Landscape Functioning 397
Suburban Landscapes. Urban Landscapes. The Megalopolis. CORRIDORS AND FLOWS 397
310 The Functioning of Conduits, Barriers, and Breaks.
SUMMARY
311 Hedgerow Function.
QUESTIONS

CONTENTS xvii
xvi CONTENTS
 404 ASCENDING TYPOLOGY 484
FLOWS AND THE MATRIX
The Landscape Attributes. Multivariate and Direct Methods for
Matrix Connectivity. Landscape Resistance. Narrows. Porosity and
Typology Construction.
Interaction Among Patches. Influence Fields. Peninsular
Interdigitation. Spatial Orientation Relative to Flows. Distance. TOWARD A PHYLOGENETIC TYPOLOGY 490
415
NETWORKS SUMMARY 491
....
Nodes and Corridors. Network Connectivity. Loops and Circuitry. QUESTIONS 492
Gravity Model. Spatial Diffusion Processes. Optimization.
425
SUMMARY ｾｉ 14. Landscape Management 495
426 WHERE HUMANS GATHER 496
QUESTIONS
PRODUCTION IN LANDSCAPES 498
12. Landscape Change 427
428 PLANNING AND MANAGEMENT OF MAJOR LANDSCAPE TYPES 499
STABILITY I The Natural Landscape. Landscapes With Forestry. The Agricultural
Variation Curves. Stability and Instability. An Example of Stability Landscape. The Built Landscape.
Followed by Degradation.
435 LANDSCAPE QUALITY 509
METASTABILITY A Direct Method of Estimating Landscape Quality. Survey
Models. Species Coexistence Patterns. Questionnaires. Perception of Landscape Quality by Animals.
440 Protection of Landscape Quality.
PATIERNS OF OVERALL LANDSCAPE CHANGE
The Transition Matrix. The Shifting Mosaic. MODELLING AND LANDSCAPE MANAGEMENT 515
446 Models from Maps. Model Construction. Tensor Models in
LANDSCAPE DYNAMICS
Management. Model Sensitivity, Risks, and Timing.
Active Forces. Levels of Force. Stabilizing Properties.
456 SOME BROADER PERSPECTIVES 526
LINKAGES AMONG LANDSCAPES
458 SUMMARY 528
SUMMARY
458 QUESTIONS 530
QUESTIONS
References 533

Glossary 589
Part IV.
Heterogeneity and Management Index 603

13. Heterogeneity and Typology 463

463
LANDSCAPE HETEROGENEITY
Thermodynamic Laws. Mechanisms Causing Heterogeneity.
ANIMAL RESPONSE TO HETEROGENEITY 470
GUIDELINES FOR LANDSCAPE TYPOLOGY 473
THE INHERENT HIERARCHY IN NATURE 477
Five Levels in a Descending Hierarchy. A Parallel Hierarchy. Map
Overlay Approach.

CONTENTS xix
xviii CONTENTS
 logically. With a low level of biomass present (relative to the same habitats
elsewhere in the landscape), the system has little resistance to change
but may recover rapidly from disturbance, as in the case of a cultivated
field. With considerable biomass present, as in a forest, the system is
usually rather resistant to disturbance but recovers slowly from it. Note
that biomass (sometimes considered an index of the "information" in a
system) is not only a photosynthetic surface but also includes a huge
number of organic and inorganic chemicals involved in protection, growth,
reproduction, and much more.
Stability of the landscape mosaic may increase in three distinct
ways, toward (a) physical system stability (characterized by the
absence of biomass), (b) rapid recovery from disturbance (low
biomass present), or (c) high resistance to disturbance (usually high
biomass present).
We label the seven statements as emerging general principles or theory
of landscape ecology. They are supported by considerable direct and
indirect evidence, rather than being conclusively proven. They deal with
major issues of the field, have predictive ability within broad limits, and
are applicable to any landscape.
The emerging principles are briefly introduced here to allow the reader
to see the breadth of the field. Much of the rationale for them emerges
in the following chapters. It is instructive to try to think of examples that
may be inconsistent with a statement and to consider whether a given
example is an exception or the norm for landscapes in general. It is also
instructive to articulate additional general statements (especially appli-
cable to landscapes), either equivalent in breadth to the emerging prin-
ciples above or more narrowly focused in a hierarchy of generalization.
Such is the process of developing human knowledge.
THE EMERGENCE OF LANDSCAPE ECOLOGY: MAJOR LITERATURE
Geography and Ecology
Since humans have always lived in landscapes, the roots of landscape
ecology could be traced to the writings of scholars in every period of
history. In their own way, for example, Herodotus (484-420 B.C.), von
Humboldt (1769-1859 A.D.), Darwin 0809-1882), and countless others
have contributed to the foundations of landscape ecology. However, the
outlines of a distinct discipline or field of study were provided by a series
of pioneering geographers and biogeographers, primarily during the 1960s,
who interpreted the landscape as being composed of landscape elements
(or biotopes or ecotopes). The term "landscape ecology" is credited to
the German geographer, C. Troll (1950, 1968).
28 OVERVIEW
E. Neef (1963,1967), J. Schmithusen (1964,1967; Bobek and Schmi-
thusen, 1967), and G. Haase (1964), all in Germany, provided early
insights into the ecological structure of landscapes, with Neef going into
particular detail'. The Americans C. Sauer (1925, ] 963) and J. Jackson
(Meinig, 1979b) analyzed landscape structure, particularly as influenced
by human culture. G. A. Hills (1960) in Canada, H. Carol (1956) in
Switzerland, V. Vinogradov (1966) and V. Sochava (1967) in the Soviet
Union, and I. Zonneveld (1979) in tne Netherlands provided additional
important insights into landscape structure from studies in diverse regions.
R. Park (1968) and K. Lynch (1960) particularly elucidated structure and
change in the urban landscape. Concepts of landscape function or dy-
namics received less attention. C. Troll discussed dynamics, but C. Van
Leeuwen (1966, 1973) in the Netherlands provided an original perspec-
tive by linking temporal variation to spatial heterogeneity in landscapes
(the relation theory).
Landscape ecologists quickly recognized the relevance of work in
several sister disciplines. From ecology, the development of the ecosys-
tem concept (e.g., Tansley, 1935; Odum, 1959, 1971; Woodwell and
Whittaker, 1968; Bormann and Likens, 1967), in the United States and
elsewhere, provided an essential holistic view. Sukachev in the Soviet
Union particularly developed the similar biogeocenosis concept (Su-
kachev and Dylis, 1964; Johnson and French, 1981). In England, C. Elton
(1958) focused attention on the invasions of animals and plants. R.
MacArthur and E. Wilson (1967), in the United States, catalyzed interest
in island biogeography and changed its focus to species diversity, rates
of species flow, and island characteristics. Traditional animal and plant
geography (e.g., Schimper, 1903, 1935; Darlington, 1957; Dansereau,
1957; Eyre, 1977; Ozenda, 1964; Walter, 1964, 1968; Udvardy, 1970)
and vegetation methodology (Braun-Blanquet, 1932, 1964; Greig-Smith,
1964; Godron et aI., 1968; Kershaw, 1975; Mueller-Dombois and El-
lenberg, 1974) are related to landscape ecology and have provided some
useful contributions.
A specific series of integrated ecological investigations on hedges or
hedgerows in Britain and France Wollard et al., 1974; Les Bocages, 1976)
turn out to be exceptiona'lly instructive for landscape ecology. These
studies not only elucidate aspects of landscape structure, function, and
change but also consider animals, plants, soil, meteorology, people, land
use, and more. Therefore, we will periodically refer to hedgerow studies
as examples in this volume.
Related Disciplines
The field of regional studies (e.g., Rey, 1960; Dickinson, 1970; Cliff and
Ord, 1,973; Isard, 1975) has provided many useful perspectives on land-
LANDSCAPE AND PRINCIPLES 29
 Preobrazhensky,  1984;  Risser  et  al.,  1984;  Brandt  and  Agger,  1984). 
scapes within regions (Miller, 1978a). Urbanization theory (Christaller,
Such  signs  indicate  that  this  new  field,  as  it  is  weeded  and  cultivated, 
1938; Rapoport, 1982a) and transportation theory, studying the move-
will  be  used  to  solve  many theoretical  and  practical  problems of tomor-
ment of  people,  have  also  proven  valuable,  especially  in  understanding 
row. 
corridors and  networks (Taaffe and Gauthier,  1973; Lowe and  Moryadas, 
1975;  Haggett  et  aI.,  1977).  Planning  and  landscape  architecture  have 
provided valuable information on landscape structure, function, and change,  SUMMARY
as  well  as  underlining the  roles  of aesthetics  and  of  human  perceptions 
of  the  landscape  (Eckbo,  1975;  Kiemstedt,  1975;  Zube  et  aI.,  1975;  Common  threads  from  several  disciplines  help  provide  a  rigorous  sci-
Fabos,  1979;  Steinitz,  1979;  Haber,  1980;  Ruzicka  and  Miklos,  1981,  entific  concept  of the  landscape.  It  may  be  usefully  defined  as  a  heter-
1984).  McHarg's treatise  (1969),  on designing with nature,  focused  peo- ogeneous land area composed of a cluster of interacting ecosystems which 
ple's  attention  on  landscape  modification  within  ecological  constraints.  is  repeated  in  similar  form  throughout  its  kilometers­wide  extent.  Geo-
The land evaluation techniques of Zonneveld (1979)  in The  Netherlands,  morphic processes,  colonization of organisms,  and  disturbance mold the 
Christian and Stewart (1968) in Australia,  and Long (1974) in France have  structure  of  landscapes.  Landscape  ecology,  therefore,  focuses  on  (a) 
given  us  valuable perspectives on scale  and landscape structure.  Finally,  structure,  the  spatial  patterns  of landscape  elements  and ecological  ob-
pioneering individuals in wildlife biology, forestry,  and  pest management  jects  (such  as  animals,  biomass,  and  mineral  nutrients);  (b)  function,  the 
have  provided  excellent  case  studies  for  landscape  ecology.  flows of objects between  landscape elements;  and  (c)  change,  alterations 
in  the  mosaic  through  time. 
Views  of  four  contrasting  landscapes  show  a  common  fundamental 
International Perspectives Today
structure of patches,  corridors,  and  a matrix; their configurations indicate 
A broad schematic look at the present status of landscape ecology suggests  the  wide variety of landscapes on earth.  From  an  array of concept areas, 
the  following  in  different regions.  In  North  America,  landscape ecology  seven  general  principles  of  landscape  ecology  have  emerged.  These  in-
has  just  been  born  and  seems  to have an  initial  major focus  on ecology.  volve  structure  and  functi,on,  biotic  diversity,  species  flow,  nutrient  re-
But several  related  disciplines are  involved,  and  the  primary focus  is  on  distribution,  energy  flow,  landscape  change,  and  landscape  stability. 
developing basic concepts (Burgess and Sharpe,  1981; Forman and  God- The  combination of modern ecological  principles with a geographical 
ron,  1981,  1984;  Romme  and  Knight,  1982;  Phipps,  1984;  Merriam,  foundation  developed  since  the  1960s  has  provided  a  synergism  for  the 
1984;  Risser  et aI.,  1984).  present  burst  of  activity  in  landscape  ecology.  Important  contributions 
In  the  Soviet  Union,  under  the  umbrella of biology and  especially  of  from  related  disciplines have helped establish  the  importance of an  eco-
geography,  landscape ecology has a strong economic and social planning  logical  understanding,  not  only  of  ecosystems,  but  also  of  the  hetero-
component.  Emphasis is  on the complex of land development, increasing  geneous  landscapes  in  which  they  coexist. 
productivity,  and  recognition of environmental  impacts (Vinogradov,  1980; 
Gerasimov  et  aI.,  1979;  Johnson  and  French,  1981;  Preobrazhensky, 
1984).
QUESTIONS
In  Europe and  the Middle East,  a rich  mixture of approaches­includ-
ing  nature  conservation,  planning,  landscape  architecture,  wildlife  bi- 1. What  characteristics  are  repeated  in  similar form  across  a landscape? 
ology, land evaluation, geography, environmental science, forestry,  land- 2.   How would you  have defined  the word  landscape before reading this  chapter? 
scape history, and concept development­promises important synergisms. 2 How  does  your  definition  differ from  that  of the  dictionary?  Of geography? 
Interest  in  landscape ecology  is  present  in  many other countries,  as  evi- Of ecology?  Why are  the  concepts  so  diverse? 
denced  by  participation  in  international  meetings  in  recent  years,  and  is  3.   How would  you  differentiate a landscape  from  an  ecosystem?  A  region?  A 
watershed  or  basin?  A landscape element?  A tessera? 
growing  rapidly  (Tjallingii  and  de  Veer,  1982;  Ruzicka  et  aI.,  1982; 
4. Compare  the  agricultural,  boreal  forest,  tropical  rainforest,  and  Mediterra-
'See,  for example,  Van  Leeuwen,  1973, 1982; Rackham,  1975,  1976, 1980; Haase,  1975,  nean  landscapes. Which had the greatest contrast? Diversity of corridor types? 
1984;  Peterken,  1977;  Ruzicka  and  Miklos,  1981,  1984;  Ruzicka  et  aI.,  1982; Schreiber,  Prevalence  of linear features?  Evidence  of recent  natural  disturbances? 
1977;  Naveh,  1982a;  Naveh  and  Lieberman,  1984;  Godron  and  Forman,  1983;  Vink,  5.   In  what  ways  are  the  structure,  function,  and  change of a landscape  similar 
1980; ten  Haute de  Lange,  1984;  Agger  and  Brandt,  1984;  Antrop,  1984;  Baudry,  1984;  to  those  of the  human  body?  Explain  any  fundamental  differences. 
Opdam,  1984;  Harms  et  aI.,  1984. 

LANDSCAPE  AND  PRINCIPLES  31

30 OVERVIEW 
 2 
6.   Describe two of the emerging general  principles,  explaining how each  could 
be  useful  in  landscape  management. 
7.   If you  were  a  wildlife  manager  responsible  for  both  protecting  rare  species 
and  maximizing wildlife for hunting,  how would you  use  the biotic diversity 
principle? 
8.   If you  were  a forester  faced  with  managing  a  whole  forested  landscape  for 
both  maximum wood  production and  soil  conservation,  how would you  use 
Ecological Concepts in Brief
the  nutrient  redistribution  principle? 
9.   If you were a planner or landscape architect designing a suburban landscape, 
how  would  you  use  the  energy  flow  principle?  Nature.
10.   In  geography,  the  scale  of a map  is  always given  in  the  legend  as  a ratio  of  A  vast, tightly interwoven fabric of activity.
map  distance  to  actual  distance.  What would  be  the  approximate  scale  of  Paul B. Sears, 1962
a  map  of  your  immediate  neighborhood?  For  a  map  of the  continent?  For  The Biology of the Living Landscape
the  landscape  in  which  you  live?  Briefly  describe  your  landscape. 
11.   The landscape change principle concerns changing structure, while the land- Isaac Newton, the noted physicist and  discoverer of gravitation, observed 
scape  stability  principle  concerns  the  types  of  stability.  What  insight  might  that  he  could  see  further  than  his  predecessors  because  he  was  able  to 
be gained  by  integrating the  two  principles together?  stand  on  the  shoulders  of  giants.  So,  too,  we  can  develop  landscape 
12.  What are  the  major areas  of thought contributing to  landscape ecology,  and  ecology  only  by  standing  on  earlier  foundations.  Several  foundations 
how have  they  meshed  to  produce  the  current  state  of the  field? 
underpin  landscape  ecology.  Ecology, the  scientific  study  of  the  rela-
tionships  between  organisms  and  their environments,  is  one.  If ecology 
is  part of you r  repertoi re,  feel  free  to  move  directly to  Chapter  3.  If  not, 
the  present  chapter,  laying out  most of the  main  ideas of the  science of 
ecology,  should  provide a  useful  background  to  landscape  ecology. 
Early  Greek  and  Roman  scholars  discussed  how  plants  and  animals 
were related  to the environment (Figure  2.1 l.  But today's ecological  prin-
ciples mainly have their roots  in the work of seventeenth­ and eighteenth-
century  natural  historians,  agriculturalists,  geographers,  and  human  de-
mographers.  During the  nineteenth­century,  plant geographers,  aquatic 
biologists,  and  zoologists  developed  principles  that  gave  birth  to  the 
discipline. The early twentieth century saw a quickening and broadening 
of  the  ecological  sphere  to  include  the  work  of  animal  behaviorists, 
evolutionists,  statisticians,  animal  geographers,  and  plant  community 
ecologists  (phytosociologistsl.  As  in  any  field,  scholars  of  many  more 
disciplines  have  contributed  and  continue to  contribute significantly. 
Two  broad  perspectives  permeate  all  branches  of  ecology.  The  first 
concerns  feedback  systems.  In  a  feedback loop,  or  system,  one  com-
ponent  affects  a  second  component that  in  turn  affects  the  first  compo-
nent;  such  loops  underlie  most  ecological  processes.  Feedback  loops 
may be  positive or negative.  When  one component stimulates a second, 
but  the  second  inhibits the  first,  we  call  it  a  negative feedback. In  this 
case,  both components may oscillate within a relatively predictable range 
through  time,  thus  operating  as  a  regulating  or  control  mechanism  to 
produce a relatively stable equilibrium. Certain  predator and  prey cycles 
are familiar examples of a negative feedback system.  Other examples are 

33 
32  OVERVIEW