Landscape Ecology

Author(s): Dean L. Urban, Robert V. O'Neill and Herman H. Shugart, Jr.

Reviewed work(s):
Source: BioScience, Vol. 37, No. 2 (Feb., 1987), pp. 119-127
Published by: University of California Press on behalf of the American Institute of Biological Sciences
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 Landscape
Ecology
A hierarchicalperspectivecan help scientists
understandspatialpatterns
Dean L. Urban, Robert V. O'Neill, and Herman H. Shugart, Jr.
A landscapeis a mo-
terrestrial
saic of heterogeneous land
forms, vegetation types, and
land uses. The study of landscape-its
spatial patterns and how they devel-
op-is presently emerging as a new
discipline in the field of ecology (For-
man 1981, 1983, Forman and Go-
dron 1981, 1986, Naveh and Lieber-
man 1984, Noss 1983, Risser et al.
1984). Landscape ecology is motivat-
ed by a need to understand the devel-
opment and dynamics of pattern in
ecological phenomena (Clark et al.
1978, Levin 1976a,b, 1978, Whitta-
ker and Levin 1977, Wiens 1976), the
role of disturbance in ecosystems
(Mooney and Godron 1983, Pickett
and White 1985, Sousa 1984, White
1979), and characteristic spatial and
temporal scales of ecological events
(Allen and Starr 1982, O'Neill et al.
1986).
Pattern, generated by processes at
various scales, is the hallmark of a
landscape. In this paper we outline an
approach to landscape study that em-
ploys a hierarchical paradigm of pat-
tern and behavior. Although our em-
Dean L. Urbanis a researchassociateand
HermanH. Shugart,Jr. is the Corcoran
Professor of EnvironmentalSciences in
the Department of EnvironmentalSci-
ences, Universityof Virginia,Charlottes-
ville, VA 22903. Robert V. O'Neill is a
senior researchecologist in the Environ-
mentalSciencesDivision, Oak RidgeNa-
tionalLaboratory,Oak Ridge,TN 37831.
They study the developmentand dynam-
ics of scaled pattern in ecosystems. ?
1987 American Institute of Biological
Sciences.
February 1987
A landscape is a
mosaic of patches,
the components
of pattern
phasis is on forested landscapes,we
can generalizea theory of landscape
ecology.
Landscape pattern and process
We will first focus on the wide range
of phenomenain a naturalterrestrial
landscapeby consideringthe appar-
ent complexityof landscapedynamics
and illustrating how a hierarchical
paradigm lends itself to simplifying
suchcomplexity.Ourperspectivealso
affordsinsightsinto the management
of man-dominatedlandscapes.
Developmentof landscapepattern.A
landscapeis a mosaic of patches, the
componentsof pattern.The agentsof
pattern formation on natural land-
scapes can be categorizedas distur-
bances, biotic processes (especially
the demographicprocesses of birth,
death, and dispersal), and environ-
mental constraints (Levin 1978).
Each of these agents can be consid-
ered across a spectrumof spatial and
temporalscales. For example, distur-
bances that affect terrestrial land-
scapes vary in spatial extent, recur-
rence interval, and intensity (Pickett landscapes would appear futile.
and White 1985, Sousa 1984, White
1979). Disturbancesrange from the Organization of landscape pattern.
localized effects of an individual But importantly, the complexity of
death to the large-scale effects of
wildfires, drought, and epidemic dis-
ease. Biotic, or regenerative, processes
also vary in scale from the regrowth
of an individual to the reorganization
of species assemblages. Environmen-
tal constraints include microclimatic
and fine-scale soil conditions govern-
ing seed germination, and also sub-
continental climatic regimes that de-
lineate biomes, such as the Eastern
Deciduous Forest.
The agents of pattern formation are
interwoven in landscape develop-
ment. This interaction allows some
sites to be especially prone to, or
sheltered from, disturbances. For ex-
ample, topographic position interacts
with fire frequency; dry ridges burn
more frequently than moist (mesic)
coves. Regenerative processes are in-
fluenced by site quality, and also vary
with the age and life-history attri-
butes of the regenerating individuals
(Odum 1969, Shugart and Hett
1973). Moreover, both disturbances
and regeneration may be constrained
by the existing spatial pattern (Curtis
1956, Forman 1981, Watt 1947). Fi-
nally, new patches are continually
superimposed on existing patches
(Reiners and Lang 1979). The emer-
gent scenario is a mosaic of patches of
various size, of various origins, in
various stages of regeneration, ap-
proaching microenvironmental equi-
libria at various rates. Such complex-
ity would seem overwhelming at first
and any attempt to fully understand
119
landscape pattern is organized in a
special way: the component events
and patches occur at characteristic
scalesthat are positivelycorrelatedin
time and space. Disturbances,for ex-
ample, occur over a wide range of
space and time scales, but those af-
fectinga particularlandscapetypical-
ly can be divided into events occur-
ring on characteristic scales. For
example, Romme and Knight (1982)
were able to break the fire regimein
Yellowstone Park into two compo-
nents:frequent,small fires (everyfew
decades, affecting areas of less than
100 ha); and larger,less-frequentfires
(everyfew centuries,affectingareasof
severalhundredha). Similarly,Hein-
selman (1973) documented a two-
scale fire regimein the BoundaryWa-
ters Canoe Area of Minnesota. The
concept of disturbance is itself implic-
itly scaled; if disturbances are events
that kill tress prematurely, then "dis-
turbance" is confined to a relatively
narrow window of time correspond-
ing to the lifespan of trees (Figure la).
Events outside this domain, while
they may affect trees, are not consid-
ered disturbances. Small-scale distur-
bances tend to occur more frequently
while larger events tend to be less
frequent.
Demographic processes are also
scaled. Individuals have characteristic
sizes and lifespans that dictate corre-
spondingly scaled patterns (Watt
1947). Species dispersal distances and
rates define the scale of larger and
longer-term patterns (Figure lb). At
still larger scales and over longer
spans of time, genetic fluxes define
the domainsof adaptationand evolu-
tionaryprocesses.
Environmental constraints are
scaled by the manner in which we
witness them. Atmospheric condi-
tions can be observedat virtuallyany
scale, but "microclimate,""weath-
er," and "climate"connotephenome-
na witnessed over increasinglylarger
areasand longertimespans.Likewise,
we could also measuresoil processes
at virtuallyany scale, but we would
not measurethe chemicalweathering
of mineralsover the entire Canadian
Shield (we would use a small, repre-
sentativesurface),nor would we ob-
servethe evolutionof landformson a
single hillslope (where we would

9 9
I I I I I I I I I I
8 a- 8

7 7

a6 I6
0
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-.5 . 5
0O
03 4 <
(4
-J -I 4
O
0 3 0.3

2 -2 2

1 - 1

U
I I I I I I I I I 0
0 2 4 6 8 10 12 0 2 4 6 8 10 12
SPATIAL SCALE (log m2) SPATIAL SCALE (log m2)

9 i I I
I I I I I I I 9

8 - 8

7 7

0) 6
2 GLACIAL 6
CYCLES
w
Lu 5 PEDOGENESIS,
-J
4 CLIMATE FLUX
0 4 -i 4
o4
(1) TOPOGRAPHY
3
- _ L 3
IL
1
2 22

1 MICRO- 1
ENVIRONS
C u
0 2 4 6 8 10 12 0 2 4 6 8 10 12
SPATIAL SCALE (log m2) SPATIAL SCALE (log m2)

Figure1. (a) Disturbanceregimes, (b) forest processes,(c) environmentalconstraints,and (d) vegetationpatterns,viewed in the
context of space-timedomains.Modifiedfrom Delcourtet al. (1983).

120 BioScience Vol. 37 No. 2

study erosional processes).The phe- Hierarchical structure. Components
nomenawe study tend to be positive- of a hierarchical system are organized
ly correlatedin time and space (Figure into levels according to functional
Ic).
Vegetationpatternscan be resolved
on differentscales (Figureld). Whit-
taker's (1953) notion of climax pat-
tern was a two-level description of
vegetationpatternthat reconciledthe
broad-scale idea of a single stable
communityof plants dictated by cli-
mate (Clements1916) with fine-scale
observationsof individualplant com-
munities (Gleason 1939). Moreover,
the large-scalegeneralizationis only
loosely coupled to the fine-scalede-
tails of a phenomenon. The succes-
sional schemesdescribedby Clements
may apply in general to large areas,
but they do not predictwith certainty
what one will actuallyfind on a par-
ticularsite at a giventime. This loose-
ly coupled, multileveledorganization
of landscapesrequiresa new concep-
tual model. We suggest a paradigm
that comes fromhierarchytheory(Al-
len and Starr1982, Allen et al. 1984,
O'Neill et al. 1986, Pattee 1973,
Whyte et al. 1969).
The paradigm
Hierarchytheory is concerned with
systems that have a certain type of
organizedcomplexity. Hierarchically
organizedsystems can be divided,or
decomposed,into discretefunctional
components operating at different LEVEL3
scales (Simon 1962). As applied to
landscape ecology, the hierarchical
paradigmprovides guidelinesfor de-
finingthe functionalcomponentsof a
system,and definesways components
at differentscales are related to one
another (e.g., lower-levelunits inter-
act to generatehigher-levelbehaviors
and higher-levelunits controlthose at
lower levels). This paradigmcan aid
the design of studies in landscape LEVEL 2
ecology and the prediction of how
external factors will alter an
ecosystem.
Natural phenomena often are not
perfectly decomposable: spatial
boundariesmay be difficultto define
preciselyand componentsmay inter-
act. Yet many complex, naturalphe-
nomenaare nearlydecomposable(Al- LEVEL1
len and Starr 1982, O'Neill et al.
1986, Simon 1973) and thus can be
conceptualized usefully as hierarchi-
cal systems.
February 1987
scramble competition ensues, until
another tree grows large enough to
become dominant. When this tree
scale (Figure 2). Events at a given dies, the cycle repeats. The spatial
level have a characteristic natural fre- unit of gap dynamics is equal to the
quency and, typically, a correspond- area affected by the death of a cano-
ing spatial scale. In general, low-level py-dominant tree; its natural frequen-
events are comparatively small and cy reflects the lifespan of the domi-
fast; higher-level behaviors are larger nant species (Shugart 1984, Shugart
and slower. More strictly, compo- et al. 1981).
nents of a hierarchical system may be Trees within a forest gap interact
ordered into levels according to a much more among themselves, by vir-
number of criteria (Allen et al. 1984). tue of their shared regime of available
Higher levels may be larger than, resources (especially light), than they
slower than, constrain (control), or do with trees beyond the gap. These
contain lower levels. In many of the interactions define the boundaries of
hierarchies we will consider, all crite- a gap. By extension, a larger forest
ria will apply. The rules structuring area can be decomposed into a mosa-
hierarchies can be conveniently illus- ic of gap-sized patches, in which each
trated through an extended example. gap undergoes its own dynamics. But
Let us develop a hierarchy to study the gaps are neither identical nor
the species-composition dynamics of completely independent. The gaps
a deciduous forest system in the east- comprising a mesic cove share similar
ern United States. The forest gap has species under similar growing condi-
long been recognized as a functional tions, and they exchange seeds and
unit in forest systems (Watt 1925, nutrients more often within the cove
1947; see also Bormann and Likens than with gaps on a nearby ridge.
1979, Shugart 1984). When a large Again, these similarities allow us to
tree dies, it creates a gap where subor- delineate an area of characteristic size
dinate trees may thrive under a re- within which gaps interact at a char-
gime of greater resources now avail- acteristic frequency, and allow us to
able to them. A transitional stage of define stands as higher-level compo-
---- -- C-
0-0
C-
C
..'< /\ ?
. . . --- o -
o
Figure2. A generalizedhierarchicalsystem.Thick arrowsindicatestronginteractions;
brokenarrows,weak interactions.
121
 FOUR LEVELS OF A FOREST HIERARCHY relationto a specifiedphenomenonof
interest. Different phenomena may
LEVEL BOUNDARYDEFINITION SCALE call for differences in the hierarchies
we use to study them. While the hier-
LANDSCAPE PHYSIOGRAPHIC PROVINCES; 10000s ha archy, gap-stand-watershed-land-
CHANGES IN LAND USE OR scape, might be appropriate for a
DISTURBANCE REGIME study of either species composition or
nutrient cycling in forests, these levels
WATERSHED LOCAL DRAINAGE BASINS; 100s-1000s ha need not be relevant for other pur-
TOPOGRAPHIC DIVIDES poses. For a landscape dominated by
recurrent fires, hierarchical levels cor-
STAND TOPOGRAPHIC POSITIONS; 1s-10s ha
responding to the scales of individual
DISTURBANCES PATCHES fires and to the larger fire-mosaic
GAP LARGE TREE'S INFLUENCE 0.01-0.1 ha might be a more appropriate concep-
tualization. For a study of forest
birds, an obvious focal scale would be
Figure3. A forestedlandscapeas a hierarchyof gaps, stands,and watersheds. territories. The higher levels, for ex-
ample, the forest stand, may or may
not be suitable to studies of fires and
nents of a forest hierarchy. of a higher-level component (Koestler birds. Thus, the species-composition
Moving upscale once again, we 1967). hierarchy for a forest system is not the
might define watershedsat the next Landscapes have a special kind of only one possible.
higherlevel, because stands within a vertical structure: they are nested spa-
watershed share a similar resource tially. Each level of the hierarchy con- Mechanistic explanation. The behav-
base and interactmore among them- tains the levels below it. This property ior of a forest gap through time de-
selves than they do with stands in of containment provides for a number pends on the individual trees in the
other watersheds. At a still higher of special features (Allen and Hoek- gap. Their growth rates, lifespans,
level, we might define landscapesas stra 1984). Levels in a nested hierar- shade tolerance, response to moisture
units of similar, interacting water- chy may coincide when defined by a and nutrient levels, and initial sizes
sheds.At the landscapescale, bound- variety of ordering criteria or mea- collectively determine which trees
aries might be coincidentwith large- sured attributes. Forest gaps might be come into dominance and which trees
scale physiographic features (e.g., recognizable in terms of species com- are suppressed. Gaps, in turn, interact
mountainranges)that governweath- position, biomass, ambient sunlight to generate stand dynamics. Whether
er patterns and limit frequenciesof (insolation), or other attributes. They sugar maple (Acer saccharum) will
species movement. Of course, such might also be recognizable in terms of become more important than white
landscapes interact as well, giving the limited basic resources that con- oak (Quercus alba) in a particular
definition to still higher levels, for strain tree growth. Thus, the hierar- stand may depend on the interaction
example, regional forest provinces, chy is also ordered on the criterion of of nearby gaps. If oak-dominated
such as a spruce-fir forest. constraint. gaps tend to become maple-dominat-
We have constructed a four-level Indeed, it seems that constraint and ed gaps as a result of seed rain from
hierarchy to represent a forest (Figure interaction can be mutually reinforc- nearby maples, stand composition
3). At each level, similar and interact- ing as ordering criteria: patches delin- will shift toward maple. This general
ing components become the function- eated by a spatially distributed con- rule carries upscale: stands, then wa-
al aggregates at the next higher level. straint (e.g., topographic moisture) tersheds, then landscapes interact to
This is a rate-structured hierarchy, may interact to generate higher-level generate successively higher-level
because components of one aggregate aggregates. Forest stands defined on behaviors.
interact more frequently and inten- topographic moisture may be joined We can go a long way toward un-
sively among themselves than with by seed dispersal to generate an inter- derstanding a complex phenomenon
components of other aggregates. This acting landscape mosaic. As a further when we explain its behavior in terms
rule defines the horizontal structure consequence, note that because land- of interactions among its parts. But
(within levels) as well as the vertical scapes contain stands, higher levels of understanding a hierarchical phe-
structure (between levels) of a hierar- this hierarchy are larger than lower nomenon requires more than mecha-
chical system. Interactions among levels; also, because stands interact nisms. Understanding requires that
components at one level generate the more within themselves than among the mechanisms be considered in
behaviors of a component at the next themselves, higher levels are slower context.
higher level. A gap has its own inter- than lower levels. This special rela-
nal dynamics, but it also contributes tionship among ordering criteria in a Constraint and higher-level context.
to the behavior of a stand. In turn, a landscape hierarchy makes its struc- Specific conditions within the gap in-
stand's behaviors are not only its ture very robust. The hierarchical lev- fluence the processes, such as tree
own, but also are a part of watershed els are often evident as patches in growth and longevity, that generate
function. Each patch, at any level, is nature. gap dynamics. The pattern of avail-
at once an integral whole and a part Each hierarchy is constructed in able light in the gap, for example,

122 BioScience Vol. 37 No. 2

 provides a context that constrains in- at the next lower level, and its signifi- derstanding the event of interest. For
dividual tree growth, usually limiting cance in the context of higher-level example, in describing gap dynamics,
replacement of a dominant tree to constraints(O'Neill et al. 1986). we know that safe sites for germina-
shade-tolerant species. This knowl- tion are important to seedling estab-
edge may not, however, allow us to Hierarchyas an analytictool. Simul- lishment. However, we need not con-
predict with certainty which shade- taneouslyconsideringa complex eco- sider these details at the level of the
tolerant species will come into domi- logical system on many scales is an gap, because within gaps safe sites
nance: it might be sugar maple, or intimidating prospect that is made occur with predictable frequency. We
beech (Fagus grandifolia), or bass- simplerby the mannerin which pat- also know that the specific forest ex-
wood (Tilia americana), or some oth- ternstranslateacrosshierarchicallev- ists because the post-Pleistocene cli-
er equally tolerant species. The great- els. Hierarchyisolates the phenome- mate favors trees, but this knowledge
er the functional redundancy among non of interest. In a forest, a tree has little bearing on the event at hand.
trees, the less predictable is case-spe- integrates fine-scale variation in its To recap, a hierarchical perspective
cific behavior. The constraint in effect physical environment, experiencing would emphasize three strategic con-
judges all trees by a single standard. only a blend. For example, a tree's cerns in an analysis of landscape pat-
Some other constraints on tree growthringis an integrationof grow- tern: (1) to detect pattern and define
growth are soil moisture levels, nutri- ing conditionsduringthe year, and it its spatial and temporal scale, which
ent availability, and disturbance re- is an instant cross-section of any long- is to define functional patches at a
gimes such as fires or wind. All these er-term trends. In addition, a tree specific level; (2) to infer which fac-
constraints affect the trees within the experiences large patterns as a rela- tors generate the pattern, whether
gap, favoring some over others. The tively constant context. Each tree they be demographic processes, envi-
constraints sort individually since within a gap is not subject to the full ronmental constraints, disturbances,
trees differ individually with respect range of soil patterns in a watershed, or a combination of these; and (3) to
to such characteristics as fire toler- but only to the small sample of such relate this pattern to adjacent levels.
ance, drought tolerance, and ability patterns that constitute the local We have emphasized that the notions
to withstand wind. condition. of mechanistic interaction and con-
The factors producing these con- A component at a given level of a straining context explicitly involve
straints are themselves patterned on hierarchy experiences as variable only multiple levels of reference. In the
some spatial scale. Topographic pat- those patterns that are similarly next section we pursue a variation on
tern governs soil factors, insolation, scaled in rate, as well as in size. By this theme.
and moisture and defines the charac- comparison lower-level dynamics are
teristic spatial scale of these con- so fast that they are experienced as
straints. Each of these constraints average values; higher-level dynamics Consequences of landscape
provides a context for the behaviors are too slow to be experienced as
pattern
of the lower levels of the hierarchy. variable. Thus, the complexity of We have seen that the apparent com-
Nested hierarchical organization dynamics and spatial patterns at sev- plexity of landscapes can be partially
further specifies the context for low- eral scales is resolved into a few varia- resolved by decomposing them into a
er-level behaviors. Returning to the bles and a set of constants, defined hierarchical framework. It is when
example of tree species replacement relative to the reference level. one considers landscape phenomena
in gap dynamics, we might not be A powerful consequence of a hier- at different levels that the conse-
able to specify precisely which tree archical paradigm is that it allows one quences of pattern at characteristic
will come into dominance, but we can to focus on an event at a particular scales emerge. These consequences
predict that it will be a species that is scale, while recognizing that there are are far reaching; reviews by Levin
well-represented in the local seed pool other scales relevant to that event. (1976a) and Wiens (1976) illustrate
and that has not been excluded by the When describing an event, its charac- the scope of the subject. One funda-
other constraints acting on that par- teristic scale dictates an appropriate mental consequence of hierarchical
ticular site. Collectively, the various sampling scale and frequency. The structure is that events causing pat-
constraints provide a context that al- scale of an event defines the observa- tern on one scale can be incorporated
lows us to make sense of what we tional level through which the rest of into higher-level behavior (O'Neill et
observe in the forest at a given time, a the hierarchy is accessed. The next al. 1986). Through this incorpo-
contextual explanation. In general, lower level provides the components ration, effectively nonequilibrium
the more constraints we consider at of the event and its mechanistic expla- dynamics or spatial heterogeneity at
one level that are relevant to one nation. Higher levels provide the con- one scale can be translated to equilib-
criterion, the greater our predictive text that gives the event greater signif- rium or constancy at a higher level.
power. icance. These higher-level factors can There are two aspects of incorpo-
Thus, to understand a complex, be treated as constants when viewed ration that are especially pertinent to
hierarchically organized system we from the reference level, though they landscape ecology. The first concern
must consider multiple levels. The may be quite variable at larger scales. is whether the biological mechanisms
reference level is the scale on which Levels of the hierarchy further re- necessary to a pattern disturbance at
the phenomenon is witnessed as an moved from the reference level are a given scale are available in the sys-
interesting event. Once specified, the not of immediate concern, in that tem; that is, do the mechanisms exist?
event has its mechanistic explanation they contribute very little toward un- Obviously a logical consequence of

February 1987 123

 patchinesswould be the evolution of over the landsscape, although not at changed the level of resolution in the
mechanisms to deal with it. Both specific local sites. Cherry wins the data, and accessed a hierarchical sys-
plants and animals have evolved di- war, thoughit loses every battle. Thus tem at two different levels (Allen et al.
versetacticsto utilizepatchilydistrib- the dynamics of regional patchiness 1984).
uted resources.One strategyfor deal- are incorporatled at a higher level. Incorporation can be passive; a dis-
ing with patchiness is to employ That species;assemblages in nature turbance is incorporated simply by
superiorcompetitiveabilityto persist can be catego*rized into types (con- increasing the scale of reference. In
within a given patch, maybe even stant assembl; ages) at a variety of the example of cherry abundance, the
exerting some control over the envi- scales (e.g., Braun 1950, Holdridge cherry does not affect the regional
ronment.An alternativestrategyis to 1967, von Humboldt 1807) suggests disturbance regime. In other cases,
concede competitive advantage by that many natlural patterns are incor- adaptive mechanisms may evolve
playingthe role of fugitiveon a larger poratedby eco)logical systems. In par- such that the disturbance regime is
scale. ticular,a disturbance regime that can actually modified. Many fire-adapted
The cherry (Prunus spp.), which be incorporate :d is not disturbing at systems evolve such that the member
requiresopen sites for regeneration,is all (Allenand .Starr1982). At a higher species produce volatile substances
a fugitive in both time and space level, disturba.nce frequency can be and serotinous seeds, which require
(Auclair and Cottam 1971, Marks viewedas a coinstraint that governs an the heat of a fire to germinate. These
1974). Cherryseedsmay be dispersed equilibrium slpecies assemblage. An species both exert some control over
great distances by birds, and at the analysisof prairie plant species com- fire frequency and capitalize on fires
site of deposition, the seeds may re- position in response to fire illustrates for episodic regeneration (Mutch
main viable for several decades. Thus, this two-level approach. Allen and 1970). This incorporation with evolu-
cherry can take advantage of an ap- Wyleto (1983:)used the "time since tionary integration has repercussions
propriate regeneration site elsewhere, fire" variable to emphasize the suc- that we will discuss later.
or it can await the recurrence of a cessional dyno amics of plant species A second aspect of incorporation
disturbance that brings an appropri- composition f(ollowing a burn. Using pertinent to landscape ecology con-
ate site to it. At the level of gap the "fire frequency" as a variable, cerns whether incorporation can be
dynamics, cherry is a loser: it is even- they demonstr;ated that different equi- realized within a particular bounded
tually replaced by longer-lived, shade- librium species assemblages were system. That is, given a geographical-
tolerant species. But the cherry main- maintained uinder different fire re- ly defined region (e.g., a park) and the
tains a nearly constant abundance gimes. In effec:t, data transformation perturbation affecting it, is the region
of sufficient scale to incorporate the
disturbance?Shugart and West (1981)
8 have addressed this idea, using a for-
I l l r est simulation model. In their simula-
V>\
\?i6s tions, individual trees suffered sto-
7 - chastic, age-related mortality, and
G>r/
each model plot (0.08-ha gap) was
(-J
c?
E 6 (\0/S> \ _ independent of other plots. They
" found that 50 model plots were neces-
Q.).x,
0
-1
/<;GS \
'/~O ~ sary to stabilize the statistical vari-
- 5
- ance in biomass associated with gap
4
LU
dynamics, that is, to incorporate gap
III dynamics using biomass as a standard.
s4 Shugart and West then compared
LU
O several bounded landscapes to the
z scale of their disturbance regimes.
43
m
cc
They defined a quasi-equilibrating
landscape as one in which the area
n 2 ratio of bounded landscape to distur-
bance regime was at least 50:1. Small-
er landscapes were called effectively
1 4- nonequilibrating(Figure4).
HED PARKS A bounded landscape that is large
0 I I I enough to incorporate the factors that
7~8 9 10 disturb its component patches has a
0 1 2 3 4 5 6
constant frequency distribution of
FOCAL AREA [LOG(m2)]
patches of all types at all times, and is
Figure 4. Classification of landscapes relative to the spatial scale of their disturbance considered to be a strictly equilibrat-
regimes,accordingto a 50:1 ratio as calculatedfor forestbiiomassdynamics(Shugart ing landscape. A smaller landscape
and West 1981). A forest stand may incorporate single treefalls but not larger that is unable to incorporate a distur-
windthrowsor fires;a smallwatershedcan incorporatetreef;allsbut perhapsnot larger bance has a transient frequency distri-
windthrows,and not wildfire;a largerparkmay incorporateall of these disturbances. bution of patch types, which changes

124 BioScience Vol. 37 No. 2

in response to each disturbance event.
These are called nonequilibrating.
The implication is that the relative
abundance of each patch type can be
predicted readily at the higher level, if RESCALE PATCH DYNAMICS
the lower-level dynamics can be in-
corporated. In a national park that is
large enough to incorporate wildfire,
a constant and predictable proportion
of fire-successional vegetation (per- RESCALE BOUNDED REGIONS REDEFINE FROM EQILIBRATING
haps critical habitat for some species)
is maintained. Conversely, a park that
is too small to incorporate a natural INTRODUCE NOVEL PATCHES RENDER ADAPTIVE MECHANISMS
wildfire regime does not lend itself to
straightforward predictions of the rel-
ative abundance of patch types.
The phenomenon of incorporation
is a natural extension of pattern at
characteristic scales (O'Neill et al. HOMOGENIZE PATTERNS
1986). This perspective is especially
illuminating when applied to man-
dominated landscapes.
Figure 5. Summaryof the effects of anthropogenicrescalingof natural landscape
patternsand processes.
Man-dominated landscapes
There is a tendency to view man-
dominated landscapes as being differ-
ent from natural landscapes. Excel-
lent discussions of human impacts on
landscapes are available (Burgess and
Sharpe 1981, Forman 1981, Forman
and Godron 1981 and 1986, Mooney
and Godron 1983). We focus here on
man's influence on the characteristic
scales of landscape phenomena.
Effects of anthropogenic scaling. A
primary influence of man is to rescale
patterns in time and space (Figure 5).
Human control of forest fires illus-
trates several ramifications of this re-
scaling. Fire suppression retards the
natural frequency of burns in systems
that have incorporated fire. When
wildfires do occur as a result of fuel
buildup, they may escape to burn
over a larger area and at a greater
intensity than they would otherwise.
A thick-barked tree that could survive
a low-intensity fire might succumb to
a hotter fire. A species with seeds that
require episodic fires in order to re-
generate might decrease in regional
abundance because fire suppression
removes opportunities for germina-
tion. In each case, an incorporating
mechanism has been short-circuited.
Under the constraint of periodic
burns, the role of a fugitive, like the
cherry, might have a winning strategy
at the landscape scale. With fire sup-
pression, the constraining rules
February 1987
ANTHROPOGENIC EFFECTS ON LANDSCAPES
HUMANACTIVITY CONSEQUENCES
RENDER ADAPTIVE MECHANISMS
LESS EFFECTIVE
CHANGE CONSTRAINING RULES
ALTER PATCH INTERACTIONS
TO NONEQUILIBRATING STATE
AND DYNAMICS LESS EFFECTIVE
REDUCE POTENTIAL FOR SPECIES
TO EVOLVE ADAPTIVE
MECHANISMS
REDUCE TREE SPECIES DIVERSITY
THROUGH LAND USE REDUCE HABITAT DIVERSITY
FOR FOREST WILDLIFE
change and that role may no longer bations that might differ in spatial or
be advantageous. Finally, small burns temporal scale from natural regimes.
in a fire mosaic might depend on an For example, the spatial scale and
immediate seed rain from adjacent dynamics of human land use may be
unburned forest in order to regener- different from any natural forest
ate. Rescaling the burns to cover larg- process. Many changes in land use
er areas might decrease the influx of cover large areas but are frequent or
seeds, slowing the regenerative chronic, contrary to the natural rule
dynamics of the mosaic. Though of large/slow or small/fast. Man-dom-
these effects are not independent, the inated landscapes may change ac-
specific actions of rescaling are (1) to cording to such nonecological factors
render natural incorporating mecha- as price of commodities or transfers
nisms less effective; (2) to change the of land ownership. One would expect
set of constraints (including distur- that such anthropogenic regimes
bance frequencies) governing lower- would disrupt the natural system,
level biotic processes; and (3) to leaving only behaviorally plastic spe-
change the degree of interaction cies. This is consistent with the fre-
among patches, thus altering behav- quent association of generalists and
iors that influence higher levels. weeds with man-dominated regions.
Man also rescales natural regions Man's activities at some scales may
by establishing new boundaries. Pipe- homogenize a forest stand's fine-scale
lines, drainage canals, and roads all patterns that result from gap dynam-
set new bounds if they are effective ics. Chronic use of woodlots for graz-
barriers to patch interactions, espe- ing or as a fuelwood source can oblit-
cially species dispersal. This is critical erate natural patterns in regeneration,
when the scale is redefined relative to so that the entire woodlot assumes a
the scale at which disturbances can be high degree of similarity. Such an
incorporated. In such cases, an equili- effect can be indirectly imposed by
brating system may be rescaled to a natural edge effects in very small
nonequilibrating state. Forest frag- woodlots. In small woodlots, in-
ments in the eastern United States creased insolation and convection can
illustrate this effect. Some fragments alter the physical environment to such
are large enough to incorporate dis- an extent that natural gap-phase re-
turbances; most are not (Pickett and placement mediated by shade and
Thompson 1978). moisture is not expressed. Very small
Man also introduces novel pertur- woodlots do not develop an interior
125
of mesic species; they remain essen- thoughtfully cutting or planting just a as a conceptualand analyticmodel to
tially all edge (Levenson 1981). This few trees. be exercised as long as it is useful.
has an obvious effect on local and A third mode of prescriptive scaling That hierarchytheoryis conceptually
regional forest diversity (Noss 1983) proceeds from our discussion of appropriate to landscape ecology
but may have further consequences. incorporation. We suggest that an in- should be apparentfrom our discus-
The regional abundance of many for- herently nonequilibrium landscape sion. We have statedvery little that is
est birds and small mammals may that cannot incorporate its internal actually novel: we have merely re-
depend on a continual availability of dynamics can be equilibrated by re- phrasedfamiliarnotions in terms of
specific forest microhabitats (Seagle scaling its internal dynamics to effect patterns at characteristicscales. In-
et al. 1984, Whitcomb et al. 1981). smaller patches. Thus, prescribed deed, severalearlyclassics(e.g., Tans-
Man's homogenizing effect may thus burning of small patches could rein- ley 1935, Watt 1947, Whittaker
contribute to a regional decline in state fire into a park that could other- 1953) and a host of more recent
forest microhabitat specialists. wise not incorporate wildfire. In efforts (Allen and Starr 1982) em-
Man's various effects on landscape rangelands, controlled rotation of braceideas that are implicitlyhierar-
pattern are neither exclusive nor inde- grazing pressure in small paddocks chical. Hierarchy theory takes this
pendent but are typically interactive mimicks but rescales the natural re- conceptuallygood fit and pushes it
and cumulative. A forest fragment gime of far-ranging herbivores, effec- deductivelytoward new insightsinto
has an imposed size. It may have its tively creating rangeland microcosms. complex, natural phenomena. As a
component events rescaled and its It seems fruitful to attempt to general- nascent interdisciplinary endeavor,
internal patterns altered. It may be ize these familiar examples to other landscapeecology can benefitfrom a
operating under a new or rescaled set systems in managed landscapes. hierarchytheory as a unifying con-
of higher-level constraints. Each of ceptualand analyticframework.
these factors contributes to the con- Experimental landscapes. Man-domi-
founded and confounding behavior of nated landscapes can provide natural
woodlots (Burgess and Sharpe 1981, experiments from which we can learn
Acknowledgments
Curtis 1956, 1959). a great deal about ecological scaling This researchwas sponsored by the
in natural systems. Human land-use National Science Foundation's Eco-
Prescriptive scaling in land manage- patterns may be more variable than system Study Program,under Inter-
ment. A knowledge of the characteris- many natural environmental patterns, agency Agreement No. BSR-
tic spatial scale and natural frequency because human land use reflects not 8103181-A02 with the US
of patch dynamics on a landscape only natural constraints (Bowen and Department Energy, under Con-
of
lends itself to prescriptive applica- Burgess 1981) but also the financial tractNo. DE-AC05-840R21400with
tions in natural resource manage- resources and personal whims of pri- MartinMariettaEnergySystems,Inc.
ment. In general, resource manage- vate landowners. Thus, these land- PublicationNo. 2807, Environmental
ment should be scaled to mimic scapes often provide a spectrum of SciencesDivision, ORNL. We appre-
natural patch dynamics, so as to take anthropogenic patches of various ciate the helpfulcommentsof T. E H.
advantage of preselected adaptive sizes within the same area (e.g., Am- Allen, B. Milne, D. Miller, P. Del-
mechanisms in the local species pool. buel and Temple 1983, Burgess and court, D. DeAngelis,and A. King.
Foresters use prescriptive scaling Sharpe 1981, Forman et al. 1976).
when they mimic natural distur- Such landscapes can provide the nec- References cited
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