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Ecological Systems in Hierarchical Perspecitive: Breaks in Community Structure and Other

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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Ecology, 70(1), 1989. pp. 36-47


?c 1989 by the Ecological Society of America

ECOLOGICAL SYSTEMS IN HIERARCHICAL PERSPECTIVE: BREAKS IN


COMMUNITY STRUCTURE AND OTHER CONSEQUENCES'
JERZY KOLASA2
Institute of Ecosystem Studies, The New York Botanical Garden, Box AB,
Millbrook, New York 12545 USA

Abstract. Explanation of the patterns of species abundances is important because it


may help in understanding mechanisms structuring communities. A general conceptual
model is proposed and examined as an alternative to previous propositions, which focused
too narrowly on specific mechanisms. According to this model, viewing the structure of
an environment as a nested hierarchy of habitat units provides a general mechanism
sufficient to account for empirically established regularities in species abundances. Various
biotic and abiotic factors can be considered as specific mechanisms sorting biological
components into respective levels and compartments of the hierarchical structure of the
environment. Because "sizes" of these compartments vary in a predictable way as a function
of their position in the hierarchy, so should the abundances. The model may provide a
conceptual framework that allows evaluation of the relative contributions of competition,
predation, and other biological interactions. The operational and highly simplified version
of the model uses spatial or temporal measures of ecological ranges of species to facilitate
testing of the general model. The operational model makes three qualitative and quantitative predictions: (1) although the species display a continuous gradation of properties,
the nested hierarchy of habitat units should lead to clustering of species at distinct levels;
(2) generalists should be relatively more successful than specialists; (3) relative abundances
of species should be predictable from their position in the hierarchical structure. An analysis
of eight communities, including flatworms, aquatic insects, foraminiferans, rodents, and
birds, supports these predictions. The clusters, or "breaks" in community structure, appear
to be a new and possibly general property. Moreover, quantitative predictions of relative
abundances for these communities are in surprisingly good agreement with the actual
abundances. A hierarchical structure of the environment appears promising in accounting
for some poorly explained community-level phenomena, such as correlation between the
species range and abundance, and differences between abundance structures of communities
in rigorous and less severe environments. The model is compatible with commonly observed as well as irregular patterns of distribution of species abundances, with high local
abundance of some species, with differentiation of extinction probabilities, and with scale
dependence of ecological phenomena.
Keyi words: abundance structure, community structure; habitat; heterogeneity; hierarchy; scales.
INTRODUCTION

Recently, hierarchy theory has made inroads into


ecology (MacMahon et al. 1978, Sugihara 1980, Allen
and Starr 1982, Allen et al. 1984, Rudd et al. 1984,
Maurer 1985). The major focus has been on development of a conceptual framework and mathematical
methods, e.g., fractals, to aid in identification of and
differentiation between phenomena occurring at various temporal and spatial scales. For example, Allen
and Wyleto (1983) demonstrated that the interpretation of fire disturbance in prairie communities depends
on the spatial extent and frequency of fire. Morse et
al. (1985) examined the relationship of leaf structure,
scale, and the abundance of leaf animals. O'Neill et al.
I Manuscriptreceived 28 October 1986; revised 22 May
1988; accepted24 May 1988.
2
Presentaddress:Departmentof Biology, McMasterUniversity, Hamilton, Ontario,CanadaL8S 4K1.

(1986) have synthesized much of the recent hierarchy


and scale related work. Their particular focus has been
on ecosystems and on the implication of observation
scale for research methodology. The progress achieved
by these authors permits concentration on further consequences of the hierarchical nature of ecological systems.
The purpose of this paper is to explore the possibility
for direct applications of the concept of hierarchy to
problems traditionally assigned to community ecology.
This paper is based on the assumption that, although
patterns of community structure are produced by a
variety of interactions, such patterns can be considered
separately from the detailed knowledge about these
interactions, and the patterns themselves can be analyzed profitably. The proposed perspective has the potential for helping in the formulation of testable hypotheses about the underlying environmental structure
and mechanisms responsible for observed patterns of

37

CONSEQUENCES OF HIERARCHY

February 1989

abundance of species. To explore this potential I focus


on the question of species abundance patterns.
Large taxonomic collections normally contain a few
very abundant species and a long list of rare species
(Preston 1962, Williams 1964). Many attempts have
been made to explain this pattern (Patrick et al. 1954,
MacArthur 1957, Preston 1962, Whittaker 1965, Pielou 1975, Sugihara 1980) but controversies continue
to abound (e.g., Routledge 1980, Ugland and Gray
1982, Connor et al. 1983, Brown 1984, May 1986).
Kolasa and Biesiadka (1984) and Kolasa and Strayer
(1988) have proposed that the habitat may be conceived of as hierarchical in structure, i.e., that each unit
of environment may be composed of a number of subunits, and that this structure may be responsible for
the observed patterns of species abundances. Here, this
conceptual model is further developed.
Some ideas incorporated in the proposed model have
been postulated earlier. Colwell and Futuyma (1971)
indicated advantages of measuring niche parameters
by taking into account the species' point of view.
McNaughton and Wolf (1970) supported the idea of
using the ecological range as a measure of the niche
width but cautioned against inconsistencies of using
orderings based on a single gradient. MacArthur and
Levins (1964) have discussed significance of recognizing grain of the habitat. Finally, Dueser and Shugart
(1978) and Dueser and Porter (1986) proposed, and
provided evidence, that species segregate into microhabitats, and that their relative abundance is positively
correlated with the range of microhabitats used. The
latter authors have found competition to be the dominating sorting force.
In this paper I explore the advantages of combining
attributes of the environment with community phenomena. The specific purpose is to create a number of
testable statements, but, more important, to demonstrate how such statements can be generated and to
show the results to which they can lead.
OUTLINE OF THE CONCEPT

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

reveal a continuum of degrees of specialization (e.g.,


Whittaker 1965, Diamond 1986).
The meaning of "specialization" needs to be clarified
at this point. Specialization, in the context of the model, stands for the real range of environment used by a
species, regardless of ecological mechanisms that determine this range. The terms specialist and generalist
are used in the same, or similar, sense that MacArthur
(1968), McNaughton and Wolf (1970), and Dueser and
Shugart (1978) applied them. More complex measures
of the specialization have been proposed that involve
both abundance and range of species (Lane et al. 1975).
For the purpose of this model it is sufficient that a
species operate in a narrow range of values of temporal
and spatial variables. The reasons for that limited use
may involve either morphological or physiological specializations and/or restrictions imposed by the habitat
(predators, competitors, unfavorable physical conditions, resource availability). Specialization is thus considered in this model to be a relational property between a species and its environment and does not
necessarily imply higher efficiency of specialists over
generalists. Indeed, once a species is restricted to use
only a smaller part of the ecological space, its only
evolutionary recourse is make the best of it: specialization would then be an unintended consequence of
reduced opportunities. Such an interpretation of specialization might be consistent with a view that there
is an advantage to specialization, as well as with the
claim of the concept presented here that specialists are
at a disadvantage.
Relation between species and habitat. -The concept
of specialization outlined above can now be combined
with the hierarchical structure of the habitat. The model assumes that the specialists use small subdivisions
of the hierarchical structure of habitat and the generalists use larger fragments of the same structure. For
the unit of environment shown earlier (Fig. 1) we can
imagine that there will be a species using the whole
unit, and a number of other species that will occupy
only fragments of the ecological space. The more specialized the species, the smaller the fragments to which
it responds. The generalists may be said to operate at
a coarser grain of environment than do the specialists.
The difference is of scale. In the simplest terms, one
can conceive of a model where one species is permitted
per fragment of habitat distinguished at any particular
level of resolution. This is because at this level the
habitat appears homogeneous and, consequently, the
competitive exclusion principle might apply (e.g., Murray 1986). Note also that with the exception of the top
generalist, all other species have to cope with some
degree of fragmentation of their habitat.
The important assumption of the model is the link
between the size of the habitat fragment and the abundance of species. The model assumes a strong, although
not necessarily straightforward, association between the
overall resource availability (shelter, food, breeding

Ecology, Vol. 70, No. 1

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

way around the complexity and continuous character


of ecological variables.
Predictions
The model implies a number of consequences. These
consequences can be tested in several ways. The model
suggests that: (1) the number of specialist species should
be higher than that of the generalists; (2) the specialists
should, on average, be less abundant than the generalists; (3) the density of specialists should be less than
that of the generalists; (4) the ecological range and
abundance should be positively correlated; (5) there
should be groups of species clustered by similarities in
their ecological range and abundance; and (6) specialists should be more vulnerable to disturbance.
The first two consequences do not require special
discussion because they are a direct result of the structure of habitat (Fig. 1). However, other consequences
are not always obvious. One of the counterintuitive
consequences of the model is a prediction (item 3) of
the relative disadvantages of being a specialist. Consider the generalist in Fig. 1. According to the general
model, its abundance will be proportional to the continuous area of the top habitat unit defined by the two
variables. The situation is different for the specialists
of the two lower levels. Although their abundances are
also proportional to the area of squares used, the expected abundance is different for the following reason.
Their habitats appear as single "patches" in the generalist's habitat. In a habitat unit even larger than one
shown in Fig. 1, these habitats would multiply but
would remain separated from each other by other, unusable patches. The more specialized the species, the
greater the geometrical and ecological distance (e.g.,
barriers of "hostile" habitats) between suitable units
of environment. In a sense, resources available to a
specialist can be viewed as being diluted in the patchwork of other habitat units. Seagle and Shugart (1985),
who modeled effects of the mosaic of habitat patches
on the species richness and species-area relationships,
have found that at least two factors may contribute to
extinction of a species in a habitat island: area and
disturbance-related habitat patch dynamics. It is not
unreasonable to assume that a species faces higher energy and population costs when it uses patchily distributed habitats, as specialists in the model do, than
if the habitat is continuous, as in the case of the generalist (see also MacArthur 1968, Fahrig and Merriam
1985). Theoretically, there may be several components
to these costs. Energy costs of getting from one patch
to another are likely to increase. The mortality due to
these costs as well as to predation and exposure to
unfavorable physical conditions is likely to increase as
well. And finally, resource utilization may become less
efficient if some of these scattered resources are not
found.
From now on, the sum of these costs will be assumed
to reduce the abundance in proportion to the degree

39

that the habitat is fragmented at the specialist level.


The ratio of the specialist's ecological range to the generalist's range is adopted as the measure of this fragmentation and called the dilution factor (D). For example, if the generalist in Fig. 1 had a range of 1, then
each specialist at the lowest level would have 1/16 of
that range. The expected abundance of the specialists
would thus be reduced 16 times if all other factors were
the same.
More generally, if the resource of species Y (Ay) is
diluted relative to that of species X by some factor
(Dy), then the energy and population costs to Yincrease
as a function of this factor. The relationship between
abundance, decrease of the size of habitat units, and
dilution of resources due to habitat fragmentation at
higher levels of resolution can be generally expressed
as follows:
Nx =ftAD,).

(1)

The relative amount of resource available to Y(Ay) is


R. Substituting

for A and D in Eq. 1, where D

Rx

Ry one obtains
Rx_

NA- -

J\RX

(2a)

or, if one prefers Ni,


NY=f

(2\
R2)Y

(2b)

where R is the ecological range of species X or Y and


N is abundance. Eq. 2b will be used to predict species
abundances used in Fig. 5. Specifically, Ry was obtained from the Appendix, column Ecological Range,
and R, was set equal to the Total Range.
Tests of the model. -Because information required
to construct a hierarchical model of the habitat structure is rarely available, a different solution to testing
the model is offered. Ecological ranges of species are
assumed to be a reflection of habitat units relevant for
the species in question, and as such, these ranges are
substituted for both spatial and temporal habitat units.
Such a substitution is later used in the validation of
the model.
The other consequences of the model (items 4-6) are
explained by examining real community examples.
EXAMPLES FROM REAL COMMUNITIES

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

Ecology, Vol. 70, No. 1

JERZY KOLASA

40

the realization that species do group according to their


ecological range, and that such a grouping has an impact on the average density of species, offers a potential
for new questions on comparative ecology of species.
The correlation between specialization and abundance
is supported by other data sets (Table 1). The statistical
evidence for the pattern of breaks comes from the analysis of clustering along the environmental range (runs
test: Table 2) and from analysis of departures from
normality of the distribution of species abundances
(Kolmogorov-Smirnov test: Table 2). Departures from
normality can be considered as an indication of clustering along this axis, for generally the distribution of
abundances in collections of species tends to normality
(Preston 1962, Williams 1964). There are no mechanisms for consistent generation of such departures in
other models that address patterns of species abundances. Nevertheless, the clustering of species at certain
levels of habitat resolution may be a common phe-

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).

occurred (see legend to Fig. 2 for more details). Another


revealing feature of this community is the presence of
discontinuities between 9 and 12 habitats (practically
between 9 and 16 habitats if abundance is also considered) and between 19 and 29 habitats, indicating a
sharp division between levels of resolution. In addition
to the discontinuities in the distribution of species over
the environmental range, abrupt changes in the abundance of species may be the second strong criterion to
identify the breaks. The discovery of these breaks is
important, for it lends strong support to the hierarchical model. This is the only model that implies clustering of species by range and abundance. Such clusters
are thought to reflect abrupt changes in the grain of
patches brought about by shifts in resolution. Finally,

II
1

LEVEL I

.......

131

6-

2 -

...
0...

LEVEL 2

LEVEL3

*.

FIG.3. A possible hierarchyof habitat specializationsof


anotherassemblage.Speciesare arrangedverticallyaccording
to the numberof habitatsthey occur in. Horizontalarrangement and connectionsare arbitrary.Levels (-- -) correspond
to the averagenumberof habitatsper species in each of the
three clusters.The levels may also be determinedby abunThe insertedsequenceof rectanglesat
danceaverages(.).
upperleft is for comparisonwith Fig. 2. Data were collected
in 1984 from an artificialbottom formed by 45 15 x 15 x
5 cm Nitex netting boxes filled with nine sizes of substrate
grainsand colonized throughperiods varyingfrom 3 to 60 d
(J. Kolasa, R. Short, and E. Stanley, personal observation;

presentedat ASLO meeting, Minneapolis, 1985).

41

CONSEQUENCES OF HIERARCHY

February 1989

from the model deserves a working hypothesis as to


the impact of the variables not included in the model.
0
*l''.
Such a hypothesis can then be clearly and quantitatively formulated within the framework provided by
*
/
\0
.0'!
the model.
Three other consequences result from this particular
model. First is the possibility of evaluating effects of
0
*
disturbance at various levels. If the lower level subunits
of the habitat are incompletely filled by specialists,
relative to the unit used by the generalist, the relative
abundance will significantly shift in favor of the higher
WATER
MAYFLIES
BIRDS
levels, i.e., the generalistss. Such a departure may often
BUGS
be indicative of a recent elimination of the specialists.
Because departures may be different at different levels,
one might be able to evaluate a degree to which various
categories of species were affected, and speculate on
the nature of the disturbance. A synthesis of this proposition with the Seagle and Shugart's Markov model
of species extinctions under various disturbance re*
0
gimes seems to be an intriguing possibility. The two
approaches appear to be highly complementary.
A second consequence is the possibility of making
1
qualitative and quantitative predictions about changes
of the community structure resulting from certain types
of stress. If, for example, the environmental range of
.,*
*
X
./
00
the system shown in Fig. 1 shrank by 50%, then 6-8
0
00
*
*
species would be lost among specialists (38-50%), 02 among the intermediate-range species (0-50%), and
RODENTS
FORAMINIFERANS
none among the generalists. The expected relative
FIG. 4. Distribution of species along ecological ranges abundances should also change in a predictable way.
(vertical axis; scales differ). Horizontal positions and lines
connectingspecies are arbitrary,showing possibilitiesrather
than actualrelations.Informationon the measureof the eco/
100
logical range used for a particulartaxon is provided in the
/
0
Y = - 2.34 + 1. 08x
Appendix.

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

Some theoretical consequences

/
0

>~~~~~~~
X~~~~~~

W/

m 20

/70/
0/ /0

/?
0

aL

On the basis of the model one could predict the


abundance for a species of any rank in a community
from the structure of habitat and the number of species
alone. Because of the good predictions for three-species
communities (Kolasa and Strayer 1988) and multispecies communities (this paper), any major departure

a~~0/
/o o
/

<:_

0/o

/
:D

dard error = 2.0353).


DISCUSSION

oL
20

I I

I
40

PREDICTED

I
60

I-

I
80

I
100

ABUNDANCE

FIG. 5. Predictedabundancesof clusters of species ob-

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

Ecology, Vol. 70, No. 1

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

Third, the model helps to define ecological units as


a function of species resolution at each level, with the
basic unit corresponding to the range of the broadest
species in the community, and subunits of different
rank defined by species of higher resolution (at lower
levels); i.e., units will differ depending on the level of
resolution. Such a treatment may help to reduce arbitrariness of selecting operational ecological units. For
different purposes, of course, there may be other approaches more appropriate for identifying ecological
units.
Compatibility with other observations
Finally, what is the relation of this model to empirical facts associated with community abundance structures? The model is not only consistent with a number
of well-documented facts associated with the species
abundance patterns, but also it does suggest deeper or
alternative explanations. I list the consequences and
predictions implied by the model, which are not new,
but which have not been derived from a single conceptual device before:
1) Species of broad ecological range will be much
more abundant than species of narrow ecological range
(Macan 1976, Brown 1984, Bock 1987 and new data
in Table 1);
2) Distribution of species abundances in small communities (comprising one unit as shown in Fig. 1) will
approach the geometric series, and tend to log-normal
distribution in large species collections (Whittaker 1965,
May 1975);
3) The probability of extinction dramatically increases with the species specialization. Notably, it has
been found (Jablonsky 1986) that the probability of a
lade undergoing a mass extinction in geological time
is correlated negatively with the geographical range it
occupied, but not with the number of species in the
lade;
4) In view of the relative disadvantage of specialists
postulated by this model, a good defensive strategy
would be to present a foraging predator with a diversified behavior, forcing it to act as a specialist. In fact,
oaks allocate tannins to some leaves only, increasing
energy costs and predation risks for the gypsy moth

(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

other models appear

to be conceptually related to the model presented in


this paper. Hanski's (1982) core and satellite species
model is the only proposition that indicates qualitative
differences among broad-range species and narrowrange species. Hanski has provided a corrected version
of Levins' (1969) model in which the probability of
local extinction of a species is a decreasing function of
distribution in a heterogeneous mosaic of habitats. One
consequence of his model is a bimodal distribution of
species along the site (=ecological range?) axis. In the
context of the habitat-based model presented here, the
core and satellite species model might be treated as
one of the specific mechanisms contributing to the population costs due to the habitat fragmentation. It is
important to point out, however, that the core and
satellite model does not predict polymodality, or multiple clusters, in the community structure. This last
prediction requires a substantially different conceptualization of the relationship between species and their
habitat. In any case, the habitat-based model appears
to be more general. This can be illustrated by its treatment of r-selected and K-selected species. The generalists in this model, which roughly correspond to the
core species of the core and satellite species model, can
recruit from either category (see also Diamond 1975),
while according to Hanski's model "core species are
certainly not r-selected" (Hanski 1982).
The sequential breakage model (Sugihara 1980) also
involves the concept of hierarchy. Kolasa and Strayer
(1988) have analyzed differences between the Sugihara
and the habitat-based model. The habitat-based model
offers a unique way of conceptualizing the structure of
the environment as a function of species resolution.
The habitat-based model also rests on the assumption
of strong difference in species specialization. It implies
a limited number of vertical subdivisions and multiple
horizontal subdivisions, i.e., many different fragments
of ecological space available at the same level of resolution, except for the coarsest one. As a result it predicts effects of habitat fragmentation and formation of
species clusters. Such clusters group species that use
different resources and, presumably, have different ecological requirements (cf. Harvey and Lawton 1986).
None of these features is implied in the sequential
breakage model.
CONCLUSIONS

The hierarchical model can be considered as a higher


level theory that reads and interprets summary outcomes of many simultaneous processes (competition,
predation, habitat preference, abiotic controlling fac-

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

This is a contribution to the program of the Institute of


Ecosystem Studies of The New York Botanical Garden. Financial support was provided by the Mary Flagler Cary Charitable Trust. I thank Dr. S. A. Levin, R. McIntosh, S. T. A.
Pickett, R. Patrick, and the following colleagues at IES for
discussions and critical comments on drafts of the manuscript:
S. Compton, A. Cundiff, S. Findlay, L. Hedin, G. E. Likens,
G. Parker, J. Pijanowska, M. Shachak, D. Strayer, and C.
Wilczynski. I also wish to thank anonymous reviewers for
valuable and insightful comments.
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February 1989

45

CONSEQUENCES OF HIERARCHY
APPENDIX
Summary of data used in Figs. 2-5.

Taxon

Source of data

Measure of ecological range

. Abundance (no. inds./


unit range)
Eco- Speciunitrange)
es
Species logical
number range cluster Expected* Observed

Rodents

Brown 1984

number of sites occupied (total range available: 115 sites)

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

number of microhabitats occupied (total


range available: 15 microhabitats)

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

home range estimated from figure (total


available range assumed: 14.5 x 106 km2
[9 x 106 square miles])

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

number of microhabitats occupied (total


available range: 43 microhabitats)

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

Ecology, Vol. 70, No. 1

JERZY KOLASA

46
Continued.

Source of data

Measure of ecological range

. Abundance (no. inds./


unit range)
Eco- Speciunitrange)
es
Species logical
number range cluster Expected* Observed

Turbellaria

Wappinger

data as mayflies above

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

number of years of occurrence (total range:


21 yr)

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

data as mayflies above

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

Measure of ecological range


number of microhabitats occupied (total
available range: 34 microhabitats)

.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.