Professional Documents
Culture Documents
Consequences
Author(s): Jerzy Kolasa
Source: Ecology, Vol. 70, No. 1 (Feb., 1989), pp. 36-47
Published by: Ecological Society of America
Stable URL: http://www.jstor.org/stable/1938410
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37
CONSEQUENCES OF HIERARCHY
February 1989
General assumptions
The conceptual model developed here entails general
assumptions on (1) the nature of the habitat, (2) properties of species, and (3) the relation between the habitat and the species. The habitat is considered to be
hierarchically heterogeneous (see next paragraph for
explanation). Species in a collection are assumed to
display many degrees of specialization to the habitat.
The species abundances are assumed to reflect the relative size as well as the degree of fragmentation of the
habitat used by species.
Nature of the habitat. -Any habitat unit, whether a
decaying log, a forest stand, a lake, a mountain, a
mountain range, or a system of tributaries, is composed
of subunits, and these subunits are composed of even
smaller subunits. The units, as well as their subunits,
A BROAD RANGESPECIES
INTERMEDIATE
00
RANGE
NARROW
SPECIES
VARIABLEI
FIG. 1. Schematicdepiction of a habitat where homogeneity or heterogeneitydepend on the resolutionwith which
speciessee theirmicrohabitats.The top level may be occupied
by a single(if unitsareconsideredhomogeneous[Smith 1972])
generalistthat does not respondnumericallyto heterogeneity
at lower levels. The next lower level may be used by a few (4
in this example)species specializedat this particularscale of
resolution,and the lowest level may be used by 16 species.
Under the saturatedcondition, all the 21 species share the
same physical space, although not all second-level species
overlap all third-levelspecies.
can be described by a set of variables in the multidimensional ecological space analogous to that of the
niche (cf. Hutchinson 1957). If two variables and two
hierarchical decompositions are used for simplicity, it
is possible to visualize the structure of habitat as a
hierarchy of subdivisions (Fig. 1). In this hierarchy
habitat fragments of increasingly smaller size appear
as a function of increasing resolution.
Operationally, these fragments may be either narrow
ranges of the most important variable(s) (but see
McNaughton and Wolf 1970) or integrated sets thereof
(e.g., microhabitats, larger patches, or whole ecosystems). Although the fragments of ecological space are
construed as multidimensional sets of variables, they
are likely to have a spatial expression and to appear
as identifiable patches. In fact, Dueser and Shugart
(1978), who studied the relationship between the habitat structure and distribution of forest rodents, have
translated a raw, multivariable description of rodent
habitat into spatial patches by using discriminant analysis. At present, the spatial patches will be treated as
a tentative approximation to the hierarchy of environment.
A question remains whether subdivisions of habitat
can be identified accurately. In my view we have sufficient statistical tools that can be applied to measurements at various spatial and temporal scales to find
out which variables form integrated, and thus potentially relevant, sets. From such analyses one can construct a model of the habitat and test it against perceptions of species in the taxonomical assemblage of
interest.
Properties of species. -Species differ in the degree of
specialization and thus in their resource requirements.
A sufficiently large collection of species is likely to
38
JERZY KOLASA
territories, nest materials, etc.) and the size of the habitat unit used by a species. This relation can be further
generalized by saying that the habitat type which a
species requires is a resource itself, and that number
and/or area of suitable patches of that habitat may be
a measure of quantity of the resource (Whittaker 1965).
The idealized form of the model is not to be taken
literally. It is meant to emphasize a possible relationship between species and their environment, and the
implications of that relationship. The relationship itself
appears much more important than the question of
abundance patterns that has led to it.
In an ideal situation, the abundance structure of the
community should thus be a strict reflection of the
structure of the environment. Other aspects of the community structure (e.g., phenology, reproductive strategies, functional roles) may also be related to the hierarchy of environment in a similar way. As stated
earlier, the model ignores specific mechanisms. Although the role of deterministic vs. stochastic factors
has been found to vary among species and communities in determining abundances of species, as much
as have the opinions on this matter (Wiens 1984), these
differences do not affect the main propositions of the
model. It is irrelevant that some species may be sorted
to their fragments of the ecological space by stochastic
factors, while others are assorted by deterministic factors, or by a mixture of both. Such distinctions may
depend on the scale chosen for description (S. A. Levin,
personal communication, 1987). The net outcome produced by these factors is an association of individual
species with respective habitat units at appropriate
levels of resolution.
Operational and conceptual problems. -Related to
the model is the question of discreteness, multidimensionality, and identifiability of habitat units. In principle, whether micro- or macrohabitats are considered,
the subunits of the ecological space may be viewed as
quite discrete from the species' perspective. Identification of these units by means independent of the species
distribution will surely face some difficulties similar to
those encountered in the measurement of the niche (cf.
Colwell and Futuyma 1971). However, some important aspects of the habitat can be measured and used
to construct models of hierarchical structure without
reference to species. Mosaics of patches at various scales
are one such aspect.
Ecological space is often considered multidimensional when nongeometrical dimensions are present
(e.g., Hutchinson 1957, May 1976, Harvey and Lawton
1986). Although this may be a necessary consideration
at a detailed level of description, such multidimensionality can be ignored at a coarse resolution because
most variables have a spatial dimension and therefore
can be mapped into a two- (or, in aquatic habitats more
suitably into three-) dimensional space (Cohen 1978).
A similar argument may be applied to successional
continua in general (cf. Whittaker 1972), providing a
February 1989
CONSEQUENCES OF HIERARCHY
39
(1)
Rx
Ry one obtains
Rx_
NA- -
J\RX
(2a)
(2\
R2)Y
(2b)
The qualitative and quantitative predictions are confronted with a turbellarian community whose species
composition and microhabitat preferences have been
described (Kolasa 1983). The results (Fig. 2) show that
the more specialized the species, the less the total abundance per habitat (Pearson correlation, r = 0.61, P <
.001). One measure of specialization used in this case
was the number of microhabitats in which the species
JERZY KOLASA
40
31
0
C,
0
0
C')
29
19
18
17 LZ
U016
13
W 12 .
7IiLIIZ
EACH
6
5=
4
IS A SPECIES
=
W 32
_~
3
2?
1
34
RECTANGLE
?-- - - - - - - - - - - - -
191
151
()
FIG. 2. Turbellariancommunity abundancestructurein
a stream(Fosso Contesora,Tuscany, Italy). Thirty-fourmicrohabitatswere analyzedalong the stream from the source
springto the mouth, coveringa wide gradientof concurring iB
physical and biotic variables. Fifty-seven species of Microturbellariawere identified. Species are arrangedfrom those
living in the largestnumberof habitats(31 at top), to those
living in singlehabitats,bottom. The heightof the rectangles
correspondsto the total abundanceof all the speciesidentified Ir
at a given level of resolution. Data from Kolasa (1983; see
also Appendix).
II
1
LEVEL I
.......
131
6-
2 -
...
0...
LEVEL 2
LEVEL3
*.
41
CONSEQUENCES OF HIERARCHY
February 1989
r2 =82.9
nomenon. In fact, I have found similar breaks in rodent, bird, foraminiferan, water bug, chironomid, and
additional turbellarian communities studied at different scales and in different settings (Figs. 3, 4, Table 2).
If a three-level organization is assumed for the community shown in Fig. 4, one can make predictions on
the basis of the model as to the expected abundance
of species and species clusters which form these levels,
and compare the predicted value with the actual abundances. Results of such a comparison (Fig. 5) indicate
that the behavior of the model closely matches the
behavior of the data, and the prediction is good (stan-
80_
o
z
n =28
~~~/
o
z
60
40
/
0
>~~~~~~~
X~~~~~~
W/
m 20
/70/
0/ /0
/?
0
aL
a~~0/
/o o
/
<:_
0/o
/
:D
oL
20
I I
I
40
PREDICTED
I
60
I-
I
80
I
100
ABUNDANCE
servedin eightcommunities(treatedas the independentvariable) regressedon observedabundancesof these clusters(dependent variable). Sources and transformations of data,
together with assignments of species to clusters, are summarized in the Appendix. Broken lines show the standard
deviation about the mean regressionline.
42
JERZY KOLASA
1. Correlation (r) between ecological specialization
and density of species.
TABLE
Community
Species
Source of data
Turbellaria
Rodents
Birds of prey
Foraminifera
Water bugs
Chironomidae
Turbellaria
Ephemeroptera
57
12
23
14
17
29
17
15
0.611
0.696
0.574
0.848
0.786
0.749
0.920
0.744
<.00 1
<.01
<.002
.001
.001
.001
.001
.01
Kolasa 1983
Brown 1984
Brown 1984
Corliss 1985
Macan 1976
Wappinger's Creek
Wappinger's Creek
Wappinger's Creek
(Feeny 1970). Analogously, an insect generalist, Romalea microptera, shows a remarkable idiosyncrasy of
repellent secretions both among individuals and over
time (Jones et al. 1986).
5) Larger species, which usually have broader ecological ranges, should, according to the model, command more resources. Yet the well-established fact that
larger animals are less abundant (e.g., Peters 1983)
might indicate otherwise. Consideration of the energy
use might be a better measure of ecological efficiency
of larger animals. In fact, Brown and Maurer's (1986)
study suggests that larger species utilize relatively more
energy per population than smaller species in the same
community consuming similar resources. This last
finding removes the apparent discrepancy between the
range and abundance of larger animals.
Difficulties
The rationale for using the spatial substitute of multidimensional ecological space is that most variables
have a spatial expression. However, the operational
model used for calculating relative predictions postulates the presence of a single species at the top level,
which is in disagreement with the data set used as the
example (Fig. 2). For greater precision, models capable
of handling dimensions that cannot be reduced to spatial or temporal dimension must be developed. Such
dimensions may, for example, include trophic differences: of the two species at the top level, one (Stenostomum unicolor, living in 31 microhabitats) is believed to be predatory, while the other (S. leucops; 29
microhabitats) is omnivorous. Thus, these species are
separated along a trophic axis. This means that, at least
for the top level, explanation (and prediction) will require another dimension that could not have been
quantified using the spatial substitute applied in these
computations.
2. Statistical analysis of breaks in the distribution of
species (runs test [RT]) and of species abundances (Kolmogorov-Smirnov test [KS]) along a measure of ecological
range, where a cell is a discrete value of that measure (e.g.,
five sites). Data on ecological ranges of species are shown
in the Appendix.
TABLE
Community*
No.
species
No.
cells
with 0
species
No.
runs
p
RT
KS
17
12
Rodents
103
.01
<.01
14
9
8
<.01
Foraminifera
NS
17
12
.01
<.01
Turbellaria
31
17
9
9
<.01
Water bugs
NS
17
29
.01
<.01
Chironomids
26
59
<.01
23
19
.001
Birdst
57
17
10
<.01
Turbellaria
.001
17
<.01
15
30
NS
Ephemeroptera
* Sources the same as in Table 1.
t Geographical ranges of bird species, originally provided
in square miles, were transformed for the runs test into a new
quantity in which one unit equals 130000 ? 65 000 km2
(50 000 ? 25 000 mi2).
February 1989
CONSEQUENCES OF HIERARCHY
Another difficulty lies in the use of direct proportionality to quantify the effect of habitat fragmentation
on populations. It is not a highly realistic assumption.
Good results obtained by using this linear function,
however satisfying, cannot be mechanistically explained at this stage.
Alternative propositions. -Two
43
tors, density-dependent factors). Integrated, these processes can be regarded as a complex mechanism that
sorts species into various microhabitats and levels. The
process of sorting species into various habitat units has
a behavior and properties that can be informative on
their own. If the model is correct, the regularities I
describe would be obtained no matter which factors,
or combination of factors, force species into using fragments of ecological space. At the same time, this theory
might provide a skeleton to be fleshed out with specific
mechanisms. The skeleton provides the context for the
mechanisms, which otherwise might be difficult to relate. The assumptions of (1) equality of ecological efficiency of all species; (2) linear effects of fragmentation
of resources on specialists; (3) negligible metabolic and
trophic differences; etc., are obviously untrue. Nevertheless, the model founded on them leads to quantitative predictions that can be used as a standard. The
departures of the reality from this standard are of considerable ecological interest. Such departures can be
evaluated with the help of this model; the Hardy-Weinberg law with its unrealistic assumptions is similarly
used in population genetics (Rosenberg 1985). The hierarchical approach can probably incorporate other
specific approaches, whenever the incorporation is appropriate or useful. For any microhabitat and its occupants, density-dependent (e.g., competition, predation) mechanisms (May 1976) may be invoked to
explain the distribution of species, or the organismcentered and density-independent interpretations could
be applied as advocated by some other authors
(MacMahon et al. 1981, Andrewartha and Birch 1984).
For the sake of simplicity and data availability, the
implications of this approach are illustrated through
analysis of the community species and abundance
structures only.
ACKNOWLEDGMENTS
44
JERZY KOLASA
February 1989
45
CONSEQUENCES OF HIERARCHY
APPENDIX
Summary of data used in Figs. 2-5.
Taxon
Source of data
Rodents
Brown 1984
1
2
3
4
5
6
7
8
9
10
11
12
1
2
9
10
12
21
51
52
62
65
81
89
3
3
3
3
3
3
2
2
2
2
1
1
0.004
0.014
0.283
0.349
0.503
1.541
9.086
9.446
13.428
14.759
22.919
27.670
6
11.5
1
8
8.5
1
8
33
26
8
18.8
46
Foraminifera
Corliss 1985
13
14
15
16
17
18
19
20
21
22
23
24
25
26
1
1
1
2
2
2
3
3
3
4
6
7
11
11
3
3
3
3
3
3
3
3
3
3
2
2
1
1
0.260
0.260
1.260
1.039
1.039
1.039
2.338
2.338
2.338
4.156
9.351
12.727
31.429
31.429
1
1
1
1
2
3.5
1.3
4
5.6
2
9.2
12.1
9.3
9.5
Birds
Brown 1984:
Fig. 7
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
25
25
25
25
50
50
50
50
50
50
300
350
1200
1250
1500
1600
1650
1800
2450
2600
2700
2750
2000
3300
3700
4
4
4
4
4
4
4
4
4
4
4
4
3
3
3
3
3
3
2
2
2
2
2
1
1
0.001
0.001
0.001
0.001
0.003
0.003
0.003
0.003
0.003
0.003
0.129
0.165
1.945
2.110
3.039
3.458
3.667
4.376
8.107
9.130
9.846
10.214
10.589
14.708
18.490
0.003
0.003
0.013
0.018
0.003
0.003
0.003
0.003
0.013
0.013
0.005
0.018
0.053
0.016
0.009
0.100
0.006
0.003
0.028
0.003
0.163
0.005
0.166
0.253
0.041
Mayflies
Kolasa et al.;
data depicted schematically in Fig. 4
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
2
2
3
3
4
6
7
8
14
15
18
20
26
29
40
4
4
4
4
4
4
4
4
3
3
3
3
2
2
1
0.090
0.090
0.202
0.202
0.360
0.808
1.100
1.437
4.402
5.053
7.276
8.983
15.181
18.886
35.93 1
1
1
1
1.3
1.3
1.7
2.6
2.1
2.3
3.7
16.9
6.0
4.2
3.8
25.2
APPENDIX
Taxon
JERZY KOLASA
46
Continued.
Source of data
Turbellaria
Wappinger
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
1
1
1
1
2
3
3
3
4
5
5
6
13
15
19
32
34
3
3
3
3
3
3
3
3
3
3
3
3
2
2
2
1
1
0.033
0.033
0.033
0.033
0.130
0.293
0.293
0.293
0.521
0.814
0.814
1.172
5.501
7.324
11.751
33.333
37.630
1
1
2
5
1
1
1
2
2.3
1
3.4
2.2
5.8
3.6
5.8
9.7
11.6
Water
bugs
Macan 1976
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
1
1
1
2
2
3
4
6
6
7
7
9
10
17
18
20
21
3
3
3
3
3
3
3
2
2
2
2
2
2
1
1
1
1
0.054
0.054
0.054
0.217
0.217
0.489
0.869
1.956
1.956
2.661
2.661
4.400
5.431
15.698
17.599
21.727
23.594
1
1
2
5
1
1
1
2
2.3
1
3.4
2.2
5.8
3.6
5.8
9.7
11.6
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
1
1
1
1
2
2
3
3
3
3
4
4
5
6
7
7
12
13
19
25
25
31
33
33
35
35
40
42
43
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
3
3
3
2
2
2
2
2
1
1
1
Chironomids
0.008
0.008
0.008
0.008
0.031
0.031
0.069
0.069
0.069
0.069
0.123
0.123
0.193
0.278
0.3789
0.3789
1.111
1.304
2.784
4.821
4.821
7.412
8.400
8.400
9.440
9.440
12.341
13.606
14.262
1
1
1
1
1
1.5
1.3
2
1
1.7
1
1.5
1.8
1.2
1
2.9
2.6
1.3
2.4
3.6
2.1
3.5
16
4.2
10.6
13.7
6.9
34.7
24.6
February 1989
APPENDIX
Taxon
Turbelleria
CONSEQUENCES OF HIERARCHY
47
Continued.
Source of data
Kolasa 1983
.Abundance
(no. inds./
.coSpciunit range)
Eco- Speciunitrange)
es
Species logical
number range cluster Expected* Observed
130
1
131
1
132
1
133
1
134
1
1
135
1
136
1
137
138
1
1
139
1
140
141
1
142
1
143
1
144
1
2
145
2
146
2
147
2
148
2
149
150
2
151
2
2
152
2
153
3
154
3
155
3
156
3
157
3
158
3
159
160
4
4
161
4
162
163
4
164
5
165
5
166
5
167
6
168
6
6
169
7
170
7
171
172
8
173
8
174
9
175
9
176
9
177
9
12
178
179
13
180
16
181
16
17
182
183
18
184
19
185
29
186
31
* Abundance, expressed here in relative terms, was predicted from Eq. 2b: N, = f(Ry2/Rx2),
specialist species Y and R is the ecological range of species X or Y. Rx = the Total Range given
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
3
3
3
3
3
3
3
3
3
3
3
3
3
2
2
2
2
2
1
1
0.022
1
0.022
1
0.022
1
0.022
1
0.022
1
0.022
1
0.022
1
0.022
1
0.022
3
0.022
3
0.022
5
0.022
5
0.022
5
0.022
5
0.022
5
1
0.089
1
0.089
1
0.089
1
0.089
0.089
1.5
2
0.089
0.089
2.5
3
0.089
0.089
8
0.200
1.7
0.200
2.3
0.200
2.7
0.200
4.7
0.200
7.3
0.200
11.7
0.355
3.3
0.355
3.8
4
0.355
0.355
7.8
0.555
1.6
3
0.555
0.555
12.2
0.800
1.2
0.800
1.5
0.800
1.8
1.088
5.7
1.088
80.1
1.421
2.3
1.421
13.5
1.799
3.1
1.799
9
1.799
14.6
1.799
14.7
3.198
6.3
3.753
2.7
5.685
19.3
5.685
22.1
6.418
24.1
7.195
3.4
8.017
7.5
18.676
122.4
21.341
35.3
where N, = abundance of
in column 3.