Landscape Ecology
Part 1 - Introduction: what is landscape ecology, really?
Terms/people:
landscape ecology
Turner 1989
Wiens 1992
interdisciplinary
Hobbs 1997
Space: The Final Frontier
Space is the final frontier in ecology (as in Star Trek). Most of ecology’s history has been
played out in a nonspatial arena: predators perfectly mixed with prey, ecosystems were
conceived as homogeneous pools of carbon or nutrients, and so on. This is not to say that there
weren't a few odd souls who worked on spatial problems, but they were definitely in the
minority. And yet, space really is inherent in ecology:
 Ecology is defined as the interaction of organisms and their environment, and their
environment is profoundly spatial (because everything has to happen somewhere).
 Ecology is also defined in terms of interactions, and one thing that seems to be true of
interactions in general is that their strength varies with distance: this is true of planetary
gravitation, and it's true of competition for resources between individual plants. And
distance implies spatial location.
The study of spatial patterns is the primary focus of a relatively young field called landscape
ecology. "Landscape ecology explicitly addresses the importance of spatial configuration for
ecological processes" (Turner et al. 2001).
How landscape ecology has variously been defined:
- "the effect of pattern on process" (Turner 1989)
- "focuses on (1) the spatial relationships among landscape elements, or ecosystems, (2) the
flows of energy, mineral nutrients, and species among the elements, and (3) the ecological
dynamics of the landscape mosaic through time" (Forman 1983)
- "deals with the effects of the spatial configuration of mosaics on a wide variety of ecological
phenomena" (Wiens et al. 1993)
- the interrelationship among "pattern, process, and design" (Joan Nassauer's keynote address, 9
July 2007, World Congress of Landscape Ecology)
Does landscape ecology pertain only to large spatial areas?
Landscape ecology focuses on six major areas of inquiry:
-to identify and understand how various processes generate and maintain landscape pattern
-to quantify landscape pattern and determine the scale(s) at which spatial pattern emerges
-to explore the effect that spatial pattern has on biotic and abiotic processes
-to study the interactions and exchanges that occur across spatially heterogeneous landscapes
-to understand the role of human land-use activities on landscape structure and function
-to develop sound ecological principles for the management of landscape heterogeneity
Some general questions that are addressed within the scope of landscape ecology are:
What biotic and abiotic processes are responsible for the formation and dynamics of landscape
patterns?
At what scale does spatial structure emerge on landscapes?
How does spatial pattern affect ecological processes, such as the movement of organisms (or the
flow of water, materials or nutrients) across landscapes?
How does landscape structure affect the spread of disturbances, such as fire or disease or
invasive species?
How can a landscape ecological perspective contribute to better resource or land management?
What can landscape ecology contribute to the conservation of biodiversity?
Click here for a PDF of a list of important topics in landscape ecology. (You'll need Adobe
Reader from www.adobe.com to read the PDF file.)
What is landscape ecology, really?
John Wiens (a key player in landscape ecology and former president of the International
Association for Landscape Ecology) performed a meta-analysis of papers published in the
journal Landscape Ecology over its first 5 years (Wiens 1992). His findings showed:
-Most studies are conducted at a large scale: therefore, landscapes are usually seen as being large
entities;
-Most studies are descriptive or conceptual, indicative of a young discipline;
-Methodologically, it is logistically difficult to do experiments with landscapes (because of
replication/size issues);
-There is a considerable investment in models, owing to the daunting logistics of field studies;
-As study objects, there is a preoccupation with vegetation pattern and land use pattern;
-In particular, landscape ecologists tend to recognize humans as being an important part of the
system (as creators of certain spatial patterns).
Richard Hobbs (another past president of the International Association for Landscape Ecology)
later (1997) summarized the second 5 years of the journal. His findings:
-A decrease in purely descriptive work (i.e., with no quantification) and much more statistical
analysis of patterns (landscape ecology discovers spatial statistics), but still no experiments.
The implication is that LE is maturing as a discipline, although it still has a ways to go. In
particular, Hobbs (1997) admonishes that landscape ecology has thus far offered precious little of
practical utility. He calls for a renewed emphasis on applying the concepts and theory of
landscape ecology to real-world applications.
Landscape ecology is not simply a synonym for spatial ecology, or for large-scale ecology, or for
applied landscape planning. Instead, LE combines all of these. LE is an interdisciplinary field
whose focus is the effects of pattern on process. To quote Risser et al. (1984), "Landscape
ecology is not a distinct discipline or simply a branch of ecology, but rather is the synthetic
intersection of many related disciplines that focus on the spatial-temporal pattern of the
landscape."
What we will cover in this course:
We will cover both the theoretical and empirical foundations of LE. We will examine how we
study landscapes, and how these studies impact other disciplines like conservation biology. The
sequence we will follow is: 1) roots/history of LE, 2) how to define and detect patterns, 3)
importance of scale, 4) techniques we use to analyze patterns, 5) responses to patterns by
individuals, populations, and communities, and 6) applications of LE.
1) Roots and history of LE
2) Detecting and characterizing landscape patterns
-Defining what a "patch" is, and devising aggregate descriptions of collections of patches (their
sizes, diversity, and such);
-Examining landscape fragmentation (and its converse, connectedness);
-How does pattern develop: there are three agents of pattern formation (physical template of
environmental constraints -- soils, topography, climate; biotic processes -- establishment and
growth, dispersal, and mortality; and disturbance regimes -- fires, floods, storms, and human land
use (which we will contrast to natural disturbances);
-Landscape dynamics, or how landscape pattern changes through time.
3) Importance of scale
4) Techniques we use to analyze spatial pattern
5) Implications of landscape heterogeneity and pattern
-Individuals (habitat selection);
-Populations and metapopulations (including source-sink relations; habitat connectedness and
population persistence);
-Communities, and questions about diversity.
6) practical applications of the principles from landscape ecology
-conservation biology and natural resource management
-landscape architecture
-ameliorating effects from urbanization
References:
Forman, R.T.T. 1995. Land Mosaics: The Ecology of Landscapes and Regions. Cambridge
University Press, Cambridge, UK.
Forman, R.T.T., and M. Godron. 1986. Landscape Ecology. John Wiley & Sons, New York, NY.
Naveh, Z., and A. Lieberman. 1994. Landscape Ecology: Theory and Applications. SpringerVerlag, New York, NY.
Pickett, S.T.A., and M.L. Cadenasso. 1995. Landscape ecology: spatial heterogeneity in
ecological systems. Science 269:331-334.
Risser, P.G., J.R. Karr, and R.T.T. Forman. 1984. Landscape Ecology: Directions and
Approaches. Special Publication Number 2, Illinois Natural History Survey, Champaign, IL.
Turner, M.G. 1989. Landscape ecology: the effect of pattern on process. Annu. Rev. Ecol. Syst.
20:171-197.
Turner, M.G., R.H. Gardner, and R.V. O’Neill. 2001. Landscape Ecology in Theory and
Practice. Springer-Verlag, New York, NY.
Urban, D.L., R.V. O’Neill, and H.H. Shugart. 1987. Landscape ecology. BioScience 37:119127.
Wiens, J.A., N.C. Stenseth, and B. Van Horne. 1993. Ecological mechanisms and landscape
ecology. Oikos 66:369-380.
Part 2 - History of landscape ecology
Terms/people:
Carl Troll
Izaak Zonneveld
Richard Forman
Frank Golley
John Wiens
Risser et al. (Allenton Park workshop)
European vs. North American paradigms
Landscape ecology as a discipline has two evolutionary lines, which might be caricatured as the
"European School" (which is also represented in the United States and elsewhere), and the
"American School" (which is also common in Australia, Canada, and elsewhere). Click here for
a figure.
The European school:
Some key European (or European-style) landscape ecologists:
Carl Troll:
Izaak Zonneveld:
Richard Forman (an American):
Many others!
Summary of the European tradition:
-origin in geography
-stimulated by the development of aerial photography
-emphasis on human-dominated (esp. ag) landscapes, cultural importance of landscapes
-integrated into current discipline of land-use planning

The American school:
Some key American landscape ecologists:
Frank Golley:
Oak Ridge Nat’l Lab ("Oak Ridge Gang"): personnel included:
-Robert V. O’Neill
-Robert H. Gardner
-Monica G. Turner
-also: Kim With, Bill Hargrove, Virginia Dale, Tony King
John Wiens:
Many others!
Summary of the North American tradition:
-some researchers (e.g. Forman, Golley) use an essentially European style
-most other adopt a style that emphasizes quantitative methods (incl. spatial stats, GIS,
remote sensing), ignores or de-emphasizes social and cultural roles of landscapes and human
aesthetics
I will be taking a North American approach in this course.
There are, of course, many other well-regarded landscape ecologists around the world, including
(in alphabetical order): Jack Ahern, Lenore Fahrig, Almo Farina, Eric Gustafson, Richard Hobbs,
Louis Iverson, Jianguo (“Jack”) Liu, Bruce Milne, David Mladenoff, Joan Nassauer, Zev Naveh,
Paul Opdam, Jianguo (“Jingle”) Wu, and many others.
Key events in the development of landscape ecology (particularly North American
landscape ecology):
- 1972: first scientific society for landscape ecology was formed, in The Netherlands
- 1981: emergence of landscape ecology in North America; BioScience article on the field by
Forman and Godron
- 1982: International Association for Landscape Ecology (IALE) was established at the 6th
International Symposium on Landscape Ecological Research in Piestany, Czechoslovakia
- 1983: The Allerton Park (Illinois) Workshop on Landscape Ecology, organized by Risser et al.,
with 25 participants
- 1984: publication of one of the first texts, Landscape Ecology: Theory and Application, by
Naveh and Lieberman
- 1984: 1st Contributed Papers session on Landscape Ecology and Biogeography at the annual
meeting of the Ecological Society of America (national recognition)
- 1986: 1st North American meeting for landscape ecology was held at the Univ. of Georgia,
hosted by Monica Turner and Frank Golley, with 100 participants
- 1986: book Landscape Ecology (Forman and Godron) published
- 1986: US Section of IALE (IALE-US) was established; 1st President – D.M. Sharpe
- 1987: The first and foremost journal of the field, Landscape Ecology, was established with
Frank Golley as the first Editor-in-Chief
Landscape ecology is a relatively young field, despite a long-standing human recognition of the
importance of pattern on process. Its recent emergence can be attributed to 3 factors (Turner et
al. 2001):
1) increasing environmental issues and land-management problems (most often
associated with an ever-burgeoning human population);
2) recognition of importance of scale elsewhere in ecology and biology;
3) technological advances for collecting and analyzing spatial data.
Epilogue
This two-pronged historical trunk has given landscape ecology a somewhat schizophrenic
personality. However, it has also given landscape ecology its primary strength: landscape
ecology is an interdisciplinary/multidisciplinary field that draws upon expertise from
geographers, ecologists, land-use planners, landscape architects, and other specialists.
Landscape ecology is not a narrowly-focused discipline. Instead, it is a broadly encompassing
one that is thus well-suited to address the complicated issues faced by humans in a world that is
being rapidly transformed.
References:
Forman, R.T.T. 1995. Land Mosaics: The Ecology of Landscapes and Regions. Cambridge
University Press, Cambridge, UK.
Forman, R.T.T., and M. Godron. 1986. Landscape Ecology. John Wiley & Sons, New York,
NY.
Naveh, Z., and A. Lieberman. 1994. Landscape Ecology: Theory and Applications. SpringerVerlag, New York, NY.
Risser, P.G., J.R. Karr, and R.T.T. Forman. 1984. Landscape Ecology: Directions and
Approaches. Special Publication Number 2, Illinois Natural History Survey, Champaign, IL.
Turner, M. G. 2005. Landscape ecology in North America: past, present and future. Ecology
86:1967-1974.