Professional Documents
Culture Documents
For Man 1995
For Man 1995
3 pp 133-142 (1995)
SPB Academic Publishing bv, Amsterdam
Keywords: aggregate-with-outliers, grain size, indispensable pattern, land mosaic, landscape change,
landscape ecology, metapopulation dynamics, mosaic sequence, patch-corridor-matrix, patch
shape, principle, regional ecology
Abstract
A dozen general principles of landscape and regional ecology are delineated to stimulate their evaluation,
refinement, and usage. Brief background material and a few references provide entr6es into the subjects. The
principles are presented in four groups: landscapes and regions; patches and corridors; mosaics; and applica-
tions. Most appear useful in solving a wide range of environmental and societal land-use issues.
complex objects render experimentation difficult, study or environmental issue, one makes the terms
e.g., in geology, astronomy, and physics, great ad- operational, usually by stating assumptions and as-
vances are made by linking observable phenomena signing scales for quantitative measurement. In ad-
to existing first principles. dition, mathematical descriptions of the principles
This approach should play an important role in can be developed, which may be useful in identify-
understanding landscapes and regions. Thus mass ing hypotheses to be tested.
flow principles, gravity and rheotaxis, central place The dozen principles are organized into four
theory, counter current principle, and form and categories: landscapes and regions; patches and
function principles all contribute to the ecology of corridors; mosaics; and applications. The world-
land mosaics. Some of the preceding are actually wide literature and evidence for the principles is
second- or intermediate-level principles based on analyzed in depth by Forman (1995). In the follow-
background fine-scale theory. Models, such as frac- ing list a brief elaboration and a few references for
tal, percolation, network, and graph theory, are each principle provide some perspective and an
simplifications of complex systems to enhance un- entr6e into the literature.
derstanding. They may or may not be empirically
supported and robust, but they often provide high-
ly useful insight that may lead to principles and ap- Landscapes and regions
plications.
The attributes or tests of a principle represent a 1. Landscape and region
time-honored and endless discussion. Six attributes
seem especially relevant to me. A general principle: A mix o f local ecosystem or land use types is re-
(1) integrates diverse areas of knowledge; (2) ad- peated over the land forming a landscape, which is
dresses significant questions; (3) has broad applica- the basic element in a region at the next broader
bility, though exceptions usually exist; (4) has scale composed of a non-repetitive, high-contrast,
predictive ability; (5) is founded in theory, which in coarse-grained pattern of landscapes.
turn has considerable supporting evidence; and (6) The landscape and region are two scales, both at
has some direct supporting evidence. All of the the 'human scale'. They are land mosaics. Mosaics
general principles presented satisfy all or most of are evident at all scales from submicroscopic to the
these tests. planet and universe. All mosaics are composed of
Some of the principles are familiar to all in the spatial elements. Those at the landscape scale are
field, while others result from newer syntheses. commonly called landscape elements, and those at
None of the general principles presented is a 'law', the regional scale are landscapes (Forman and
such as for conservation of matter, where no excep- Godron 1986; Forman 1995). Just as full under-
tions are expected. However, I suggest that each standing of a liver or a town requires information
principle correctly holds for the bulk of the cases in- from both broader and finer scales, understanding
volved, e.g., more than 95% of the time. Although a landscape requires information on the broader
parts of all the principles stated are falsifiable, this region and on the finer-scale local ecosystems.
is a minor consideration compared with providing The boundary between landscapes is easily deter-
enhanced understanding, which often includes mined by recording the landscape elements present
many additional important and pioneering ad- along transects or in randomly or regularly dis-
vances. Furthermore, though the core of landscape tributed plots. For example, a line of plots across an
and regional ecology is science, the field explicitly agricultural landscape might contain cornfields,
embraces and integrates other slices of knowledge. beanfields, hedgerows, farmsteads, farm roads,
The principles stated below are conceptual and woodlots, paved tarmac roads, stream corridors,
often include relative terms, such as large or small, and some rarer ecosystems or land uses. Each plot
fine- or coarse-grain, and distinct or gradual contains most of these types, although the percen-
boundary. In applying a principle to a particular tages of individual elements may vary considerably
135
species, provide core habitat and escape cover for tive to flows in the landscape, i.e., the orientation
most large-home-range vertebrates, and permit angle, is a key to several ecological phenomena
near-natural disturbance regimes. (Forman and Godron 1986; Turner 1987; Skidmore
Large natural-vegetation patches serve many 1987; Brandle et al. 1988; Gutzwiller and Anderson
major ecological roles and provide many benefits in 1992). These include wind and water flows, which
a landscape, including those highlighted above sculpt patch shapes, produce distinct areas of tur-
(Forman 1995). Consequently a landscape without bulence, and cause soil erosion. The orientation of
large patches is eviscerated, picked to the bone. A wooded patches has also been linked to the proba-
landscape with only large patches of natural vegeta- bility of their use by migrating birds (Gutzwiller
tion misses few values. On the other hand, small and Anderson 1987, 1992).
natural-vegetation patches serve as stepping stones
for species dispersal or recolonization, protect
scattered rare species or small habitats, provide 5. Interactions among ecosystems
heterogeneity in the matrix, and habitat for an oc-
casional small-patch-restricted species. In effect, All ecosystems in a landscape are interrelated, with
small patches provide different benefits than large movement or flow rate of objects dropping sharply
patches, and should be thought of as a supplement with distance, but more gradually for species inter-
to, but not a replacement for, large patches. actions between ecosystems of the same type.
We may hypothesize that an optimum landscape A first 'law' of geography states that everything
has large patches of natural vegetation, supple- is interrelated, but near objects are more related
mented with small patches scattered throughout the than distant objects. Examples of interactions
matrix. Alternatively, most of the small-patch among nearby ecosystems or land uses are numer-
functions can be provided by small corridors in the ous and familiar (Swingland and Greenwood 1983;
matrix. Saunders et al. 1987; Senft et al. 1987; Shaver et al.
1991; Saunders and Hobbs 1991; Noss 1993).
From ecosystem science we learn that energy and
4. Patch shape mineral nutrients flow from one object to another
within, or between, ecosystems. From behavioral
To accomplish several key functions, an ecologi- science, because certain habitats are more suitable
cally optimum patch shape usually has a large core than others for a species, many locomotion-driven
with some curvilinear boundaries and narrow movements are directional, toward patches of the
lobes, and depends on orientation angle relative to same type. Combining these principles with the
surrounding flows. geography law provides this spatial-flow pr!neiple
A compact or rounded form is effective in con- (Forman 1987, 1995), useful for example, in es-
serving internal resources, by minimizing the ex- timating which ecosystems of the mosaic to focus
posed perimeter to outside effects (Harris and on in planning and management.
Kangas 1979). But patches affect, and are affected
by, manifold ecological processes in a landscape
(Forman 1995). Interactions with adjacent eco- 6. Metapopulation dynamics
systems, e.g., for multihabitat species or to escape
from predators, are enhanced by curvilinear For subpopulations on separate patches, the local
boundaries. Interactions with more distant por- extinction rate decreases with greater habitat quali-
tions of the landscape are enhanced with narrow ty or patch size, and recolonization increases with
lobes, e.g., to increase recolonization rate follow- corridors, stepping stones, a suitable matrix habi-
ing local extinction in the patch, or for species to tat, or short inter-patch distance.
disperse to other patches. A metapopulation is a population consisting of
The orientation of the long axis of a patch rela- spatially-separate subpopulations that are con-
137
nected by the dispersal of individuals (Wilson 1975; aries separating spatial elements are locations where
Fritz 1979; Merriam 1988; Opdam 1991; Hanski objects usually slow down (or in some cases acceler-
1991). Metapopulation dynamics is of special ate) (Swingland and Greenwood 1983; Forman and
importance, because subpopulations may drop to Godron 1986; Brandle et al. 1988; Forman and
zero (local extinctions), especially in small isolated Moore 1992). Hence, boundary-crossing frequen-
patches. If each subpopulation dropped to zero, cy, i.e., the number of boundaries per unit length
this would mean extinction of the whole metapopu- of route, appears to be a useful measure of re-
lation. However, because individuals sometimes sistance.
move between subpopulations, two results occur. In several landscape areas studied in The Nether-
First, the local extinction rate (the number of spe- lands, resistance to species movement increases as
cies disappearing from a patch per unit time) is the percent of area deforested increases (Harms and
lowered. Second, when local extinction does take Opdam 1990; Knaapen et al. 1992). In built areas
place, recolonization of individuals may reestablish (more or less continuous lots containing buildings)
a new subpopulation at the site. Consequently, with butterfly movement is most inhibited, and move-
extinctions followed by colonizations the metapop- ment of forest birds least inhibited. However, in
ulation as a whole persists. The local (subpopula- rural areas bird movement is most inhibited, and
tion) scale may be highly unstable, while the broad large mammals least affected. Highway and river/
(metapopulation) scale exhibits more stability. canal corridors are serious barriers to bird move-
Extinction rate and colonization rate must be ment, and least inhibitory to large mammals.
related both to patch attributes and the surround- Certain landscape elements are more suitable,
ing pattern (Opdam 1991). Extinction rate tends to and others less suitable, to movements and flows
be high in small patches and in low-quality patches. (Forman 1995). In addition, corridors can act to
Extinction rate also increases with higher environ- channel or enhance flow, or act as barriers or filters
mental variability (den Boer 1981; Karr 1982), such inhibiting spread. Finally, highly-heterogeneous
that a population widely fluctuating in size is more areas have a high probability of containing unsuit-
prone to extinction. Recolonization is enhanced by able elements, thus requiring a high boundary-
spatial patterns such as corridors, networks, a row crossing frequency and/or a convoluted route.
of stepping stones, and a cluster of small patches. Therefore, we expect heterogeneous fine-grained
Some evidence exists that recolonization correlates areas to have a high resistance.
with the area of woodland surrounding a patch (van
Noorden 1986), and with the average distance to
other patches occupied by the species (Verboom et 8. Grain size
al. 1991; Opdam 1991).
A coarse-grained landscape containing fine-grained
areas is optimum to provide for large-patch eco-
Mosaics logical benefits, multihabitat species including
humans, and a breadth of environmental resources
7. Landscape resistance and conditions.
The grain size of a landscape mosaic is measured
The arrangement o f spatial elements, especially as the average diameter or area of all patches
barriers, conduits, and highly-heterogeneous areas, present (Forman and Godron 1986; Norton and
determines the resistance to flow or movement of Lord 1990; Wiens 1990; Angelstam 1992; Wiens et
species, energy, material, and disturbance over a al. 1993; Forman 1995). A coarse-grained land-
landscape. scape with only large patches may provide large
Landscape resistance is described as the effect of natural-vegetation patches for aquifer protection
spatial pattern impeding the rate of flow of objects, and specialist interior species, large built areas for
such as species and materials. For example, bound- industrial specialization, and so forth. Coarse grain
138
is moderately monotonous, that is, although land- Perforation is the process of making holes in an
scape diversity is high (being in a farmland is very object such as a habitat or land type (e,g., dispersed
different than in a city), site diversity is low (moving houses or fires in a forest). Dissection is the carving
from one site or point to the adjacent point in up or subdividing of an area using equal-width lines
almost all cases involves no change in land use). Ex- (e.g., by roads or powerlines). Fragmentation is the
cept near boundaries, movement is costly (consider- breaking of an object into pieces (that are often
able distance required) for multihabitat species, widely and unevenly separated). Shrinkage is the
i.e., those that use two or more habitats. decrease in size of objects, and attrition is their dis-
In contrast, a fine-grained landscape has pre- appearance.
dominantly generalist species, since specialists re- These five spatial processes overlap through the
quiring a large patch of one land use cannot sur- period of land transformation. They also are usu-
vive. A fine-grained landscape is monotonous ally ordered in their importance, with perforation
(every portion is about the same), although site and dissection both peaking in relative importance
diversity is high (each adjacent point is a different at the outset. Fragmentation and shrinkage pre-
land use). Species that survive need only move short dominate in the middle phases, and attrition peaks
distances. near the end.
Some variables such as air and water quality tend These spatial processes all increase habitat loss
to be low in a fine-grained landscape, because pol- and isolation. However, average patch size de-
lution sources may be widely distributed through- creases in the first four processes, and typically in-
out the land. Indeed, the overall resource base is creases upon attrition, because small patches are
truncated and narrower, due to the absence of most likely to disappear. Connectivity across an
specialized resources in large patches. area in continuous corridors or matrix typically
A medium-grained landscape misses the large- decreases with dissection and fragmentation. The
patch benefits and offers no other advantages. In total boundary length between original and new
short, all of the preceding benefits, and few short- land types increases in the first three processes, and
comings, are provided by a coarse-grained land- decreases with shrinkage and attrition. In short,
scape that contains some fine-grained areas. each spatial process has a highly distinctive effect
on spatial pattern, and consequently on ecological
processes, in a changing landscape.
9. Landscape change
initial-land-type at the outset, and expands outward themselves. In the mid-transformation phase, the
on opposite sides. (3) Nucleus: spread from a single thickening jaws covered with scattered bits hold a
nucleus within the landscape proceeds radially, and few large separated chunks. In the late transforma-
leaves a shrinking ring of the initial-land-type. tion phase the huge jaws covered with bits hold a
(4) Nuclei: growth from a few nuclei produces new- single large chunk. At the end the bits disappear.
land-type areas expanding radially toward one an- This may represent an 'ecologically optimum' land
other. (5) Dispersed: widely dispersing new patches transformation.
rapidly eliminates large patches of the initial-land-
type, produces a temporary network of the initial-
land-type, and prevents the emergence of large Applications
patches of the new-land-type until near the end.
Mosaic sequences can be analyzed for many 11. Aggregate-with-outliers
changing spatial attributes, such as patch size and
boundary length (Richter 1984; Wiens et al. 1986; Land containing humans is best arranged ecologi-
O'Neill et al. 1988; Turner 1989; Odum and Turner cally by aggregating land uses, yet maintaining
1990; Forman 1995). The five sequences can then be
small patches and corridors o f nature throughout
compared based on a wide range of ecological
developed areas, as well as outliers of human activi-
characteristics known to correlate with the spatial
ty spatially arranged along major boundaries.
attributes (Forman and Godron 1986; Ambuel and
Seven, mainly landscape-ecological attributes are
Temple 1983; Franklin and Forman 1987; Brandle
incorporated into or solved by this spatial principle
et al. 1988; Noss 1993). Here we assume the initial-
or model (Forman 1995; Forman and Collinge
land-type is more ecologically suitable than the new
1995): (a) large patches of natural vegetation;
type. Then the mosaic sequences are compared to
(b) grain size; (c) risk spreading; (d) genetic diver-
determine which retains the ecologically best ar-
sity; (e) boundary zone; (f) small patches of natural
rangement of initial-land-type for the longest
vegetation; and (g) corridors. An example will illus-
period.
trate the principle.
Based on the ecological characteristics correlated
Start with a coarse-grained landscape with only
with the spatial attributes, the 'edge' mosaic se-
quence is considered ecologically the best of the five large patches or areas of the major land uses
transformation sequences. It has no perforation, present, e.g., natural-vegetation, agricultural, and
dissection, or fragmentation. It is best for the large- built areas. Scatter small patches (and corridors) of
patch attributes, and good for connectivity. Yet, natural vegetation over the agricultural and built
shortcomings of the 'edge' sequence include no areas to provide bits of heterogeneous nature over
'risk spreading', a progressive narrowing of the these developed areas, to protect dispersed rare spe-
remnant initial-type until it is only a strip, and an cies and small habitats, and to provide stepping
extensive area of new-land-type without small stones for species movement. Add major corridors
patches and corridors. connecting the large natural-vegetation patches to
A theoretical mosaic sequence (labelled with a facilitate movement of patch-interior species. Add
'jaws and chunks' metaphor) overcomes these small patches of agriculture near the boundaries be-
shortcomings (Forman 1995; Forman and Collinge tween natural-vegetation and built areas. Also with
1995). Draw an isolated square landscape of initial- increasing distance from the large agricultural
land-type, where L-shaped or wide-open 'jaws' of areas, make the small farm-patches further apart
new-land-type will progressively move from upper from one another. Then distribute small built
to left to lower right. Early in land transformation patches (towns, villages, houses) in exactly the same
the jaws appear to grip a huge 'chunk' of initial- manner; near boundaries between natural-vege-
land-type, and 'bits' of initial-land-type (small tation and agricultural areas, and increasingly iso-
patches and corridors) are scattered over the jaws lated with distance from large built areas. This dis-
140
tribution of small patches of all types enhances the of several individual patterns. The second is a fine-
development of genetic diversity, and provides risk scale pattern, i.e., the irregular, curvy, mosaic-like,
spreading against severe disturbance. The bound- aggregated, and complex nature of nature. Its eco-
ary zones between large patches, especially around logical benefits are surprisingly little documented,
junctions (convergency points) of three land-use but probably cannot be otherwise replicated.
types, are fine-grained areas within a coarse-grained
landscape. These fine-grained corridor and junc-
tion areas are effective for multihabitat species. Conclusion
The aggregate-with-outliers model for land plan-
ning thus has numerous ecological benefits The objective of this article is to identify general
(McHarg 1969; Forman and Godron 1986; Franklin principles largely emerging from a decade of work
and Forman 1987; Turner 1989; Hansen et al. in landscape ecology to understand the ecology of
1992). In addition, a range of direct benefits to land mosaics or large spatially-heterogeneous
humans is suggested by this spatial approach. These areas. The objective is also to stimulate evaluation,
include an exceptionally wide range of locations for discussion, and additional work to further solidify
urban dwellers to hermits, fine-scale areas where the conceptual foundations.
jobs, homes and shops are close together, efficient Presenting a dozen principles is somewhat ar-
transportation connecting built areas, large patches bitrary; some number between ten and twenty
for resource harvest or extraction, and high visual seems appropriate today. A higher number means
diversity. The principle appears applicable in any adding principles whose significance or applica-
landscape, from dry to forest, and from agriculture bility is narrower or more limited in scope. Alterna-
to suburb. tively it means adding ones with less supporting evi-
dence. Other scholars in the field would articulate
a different list, and the preceding list will change in
12. Indispensable patterns the years ahead as new questions are posed and ad-
ditional evidence accumulates.
Top-priority patterns for protection, with no Finally, these principles should be applicable for
known substitute for their ecological benefits, are a any environmental or societal land-use objective. In
few large naturaL vegetation patches, wide vegetat- a probabilistic or uncertain world, principles should
ed corridors protecting water courses, connectivity be applied intelligently, not blindly, to solving our
for movement o f key species among large patches, land use problems. They are useful in growing
and small patches and corridors providing heter- wood, protecting species, locating houses, pro-
ogeneous bits o f nature throughout developed tecting soil, enhancing game, protecting water re-
areas. sources, providing recreation, locating roads, and
The background and references for most of these creating sustainable environments. Each objective
have been introduced in the preceding pages. The is accomplished more effectively, and for a longer
indispensables should be essential foundations in time frame, using a healthy dose of landscape and
any land plan, since they accomplish major ecologi- regional ecology principles.
cal or human objectives, and no other practical
mechanism is known to accomplish them (Forman
1995; Forman and Collinge 1995). Other patterns Acknowledgements
appear to simply be efficient or optimal solutions to
difficult land planning problems. As evidence ac- I very much appreciate Michael W. Binford for his
cumulates, two additional patterns may be consid- perceptive contributions to this article, as well as
ered indispensables in the future. One is the aggre- Bruce T. Milne and Thomas R. Crow for signifi-
gate-with-outliers pattern, which is an integration cant reviews of the manuscript.
141
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