Levels of Biological Organization: An Organism-Centered Approach

Author(s): James A. MacMahon, Donald L. Phillips, James V. Robinson, David J. Schimpf

Source: BioScience, Vol. 28, No. 11 (Nov., 1978), pp. 700-704
Published by: University of California Press on behalf of the American Institute of Biological Sciences
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 Levels of
An Organism-Centered
Approach
James A. MacMahon, Donald L. Phillips,
James V. Robinson, and David J. Schimpf
The concept of "Levels of Organiza-
tion" in the biological sciences has had a
checkered history. The general nature of
this concept is one of increasing com-
plexity from molecule through cell, orga-
nism, and ecosystem, though details
vary among authors. Most contemporary
biologists accept the concept, the useful-
ness of which has been stated by Orians
(1973, p. 3):
Since biological investigations range
from determiningthe structureof some
importantmolecule to measuring the
transfer of solar energy through com-
plex systems of interactingorganisms,
some conception of the level of organi-
zation at which a problemis relevantis
essential. At any given level the kinds
of questions that can be asked about
every problem must also be under-
stood.
Early treatises concerned systems at
the level of the organism or below. At
the same time, basic concepts of hier-
archies above the organism level were
being formulated by ecologists, though
in a comparatively diffuse manner. Rowe
(1961) provides an extensive review of
the history of "levels-of-organization"
concept in the ecological context. In ad-
dition, he alludes to the importance of
the individualistic approach (a la Glea-
son 1927). Increased interest in levels of
organization may have been due to sche-
matic presentations in textbooks (e.g.,
The orderof authorshipis strictlyalphabetical.This
paperis a contributionfromthe Departmentof Biol-
ogy and the Ecology Center, UMC 53, Utah State
University, Logan, UT 84322. Currentaddress for
Phillipsis Departmentof Biology, EmoryUniversi-
ty, Atlanta,GA 30322,and for Robinsonis Depart-
mentof Biology, Universityof Texas, Arlington,TX
76010. ? 1978AmericanInstituteof BiologicalSci-
ences. All rightsreserved.
700
Biological Organization:
Odum 1959, Weisz 1959). Since then,
schemata of various forms have ap-
peared in many texts, sometimes depict-
ed in more complexity (e.g., Weisz
1966).
Guttman (1976) doubted the veracity
and usefulness of the levels-of-organiza-
tion concept in biology, pointing out that
the composition and interactions of each
level in the hierarchy are not determined
merely by systems of the level immedi-
ately below; systems of all levels inter-
act, and some levels may be absent. Al-
though Guttman's objections may make
the concept less simplistic, they do not
violate the principles of hierarchy theory
(Bossort et al. 1977). Hence, they do not
preclude the existence of a hierarchical
structure in the biological world worthy
of study.
We believe that an organizational con-
cept of this type has heuristic value, pro-
viding insights to both students and re-
searchers. we here
Consequently,
propose an alternative schema which,
because of our biases, elaborates on the
supraorganismal levels. Our concept em-
phasizes the organism as the pivotal unit,
not just another rung on the ladder, and
changes the emphasis from "levels of or-
ganization" to "biological relationships
of the organism." Our nontraditional use
of some biological terms might initially
cause conceptual dyspepsia, but we be-
lieve consistancy in our definitions will
offset any novelty of the usages we pro-
pose. We will discuss briefly each branch
of our schema (Fig. 1) and allude to some
of its implications, especially those in
ecology.
THE SCHEMA
Our organism-centered diagram (Fig.
1) should be read from the organism out-
ward along each of the four lines of rela-
tionship types. Some categories might be
skipped; e.g., for unicellular organisms,
tissue-level relationships are not mean-
ingful nor is the deme concept appropri-
ate for an asexual form. Our categories
are not exhaustive, but merely the most
commonly used; others, such as sub-
family or organelle, can be used.
Physiological-Anatomical
Relationships
The various relationships from the or-
ganism through the subatomic particle
are by far the most often discussed in the
context of "levels of biological organiza-
tion." One reason this line has had scien-
tific appeal is that all levels are com-
posed of aggregations of units of lower
levels in the hierarchy. For example,
molecules are composed of atoms and in
turn subatomic particles; cells are com-
posed of subcellular structures, mole-
cules, atoms, and subatomic particles.
The physiological-anatomical line of
relationships is particularly reasonable
because most levels are clearly bounded
in space and time, i.e., contained within
a living organism. Important exceptions
occur when some of an organism's mole-
cules or other constituents are used out-
side of the individual, as is the case with,
e.g., plant allelochemics, bacterial exo-
enzymes, insect pheromones, or fish
electric fields.
This line represents a hierarchy of gen-
erally increasing structural complexity
culminating in the organism. Each level
of complexity appears to have some
emergent properties, but structure and
function are not determined merely by
these emergent properties.
BioScience Vol. 28 No. 11
 Although structure and function are
usually thought of as intimately associat-
ed, there is no clear functional hierarchy
represented in the physiological-anatom-
ical line. In fact, this line, as much as any
of our relationship lines, shows that a
given level may coact with other levels
above or below it in the series. All such
coactions determine function. For ex-
ample, a tissue's functions are based on
molecules affecting it, the state of its
constituent cells and extracellular ma-
trix, and the milieu of the organ and or-
gan system of which it is a part.
We could subdivide the connections
between any two of our levels if a specif-
ic area of interest required this closer
scrutiny; e.g., there are important in-
creasing structural complexities going
from a nucleotide to a nucleotide pair to
a triplet, etc., in a DNA molecule. The
possibility of such refinements does not
negate the existence of the more general
pattern, which we choose to emphasize
for simplicity.
Phylogenetic Relationships
Historically, this is probably the oldest
and least modified series of relationships
of the organism, at least in its structure.
Conceptually, however, the nature of
each of these relationships is highly con-
troversial. Recently, new kingdom ar-
rays have been proposed (Whittaker
1969, Woese and Fox 1977), while the
species concept is periodically reas-
sessed, even to the point of calling the
organism the "species" (the hyper-
modern species concept of Ghiselin
1974, Platnick 1977). Moreover, the very
active controversies concerning cladis-
tics, phenetics, and evolutionary system-
atics continue (Mayr 1974). Despite
these philosophical paroxysms, the orga-
nism through genetic descent from com-
mon ancestors remains variously related
to other organisms.
This continuum of relatedness is arbi-
trarily partitioned by a hierarchy of
terms (Fig. 1). In the ideal case, the ter-
minology will reflect levels of phyloge-
netic relationship. The problems in the
phylogenetic line are generally those of
choosing appropriate characters to es-
tablish levels. The characters should re-
flect degree of difference. Certainly there
is no current consistency in application
of terms. (Are two passeriform bird fami-
lies as distinct from each other as two
araneomorph spider families?)
We do not enter any of these con-
troversies. We merely suggest that none
of the fluidity of conceptualization in the
November 1978
KINGDOM COMMUNITY BIOSPHERE
DIVISION
or PHYLUM
CLASS
ORDER POPULATION
2 FAMILY 3 4
GENUS
I
SPECIES DEME ECOSYSTEM
ORGANISM
ORGANSYSTEM BI"
OLOGICAL RELATIONSHIPS
of the
I ORGANISM
ORGAN Physiological-Anatomical
I 2 Phylogenetic
TISSUE
1 I Coevolutionary
CELL 4. Matter-Energy Exchange
SUBCELLULAR
STRUCTURE
I
MOLECULE
I
ATOM
SUBATOMIC
PARTICLE
SUBATOMIC
PARTICLE
Fig. 1. Biological relationships of the organism.
phylogenetic relationships will alter the tionary relationships. Coevolution is the
fact that the individual organism has a subset of evolution resulting from selec-
phylogeny and is variously related to tive forces produced by interactions with
other organisms with some level of ge- other organisms. Traditionally, the term
netic difference involved. Regardless of coevolution has referred to the result of
what one calls the levels (ours are tradi- interspecific interactions, and most com-
tional) or how one determines member- monly reciprocal effects have been im-
ship in the level, the organism is related plied. Since we see no qualitative dif-
to others phylogenetically. ference in the nature of the interactions
Parenthetically, we note the lack of a between different species or within the
universally accepted formal term for the same species, we suggest that coevolu-
ultimate taxon, i.e., one containing sev- tionary interactions can occur between
eral kingdoms. Since the erection of such organisms, demes, or populations of the
a level implies a decision as to a common same species and may affect only one of
origin for two or more kingdoms, we the interactants.
gladly leave this problem for others. The combinations of positive, nega-
tive, and neutral effects on expected re-
productive success are embodied in the
Coevolutionary Relationships
familiar concepts of competition, mu-
Organisms are associated with other tualism, commensalism, etc. (Burkhold-
organisms in ways that affect each oth- er 1952). These are specific kinds of coe-
er's expected reproductive success; we volutionary interactions. Though usually
will term all such interactions coevolu- depicted for pairs of organisms or popu-
701
lations, these interactions may be in-
direct, i.e., mediated through other orga-
nisms, demes, or populations. Although
the interactions occur between orga-
nisms directly, it is often useful to treat
statistically aggregated groups of orga-
nisms as though the groups interact
directly. Organisms and multiorganism
aggregations known as demes, popu-
lations, and communities constitute
coevolutionary units.
When organisms interbreed with a
high frequency, they may essentially
meet the criteria for panmixis and thus
constitute a deme. Demes are not neces-
sarily genetically isolated from each oth-
er. At a lower frequency of genetic ex-
change (interdemic), we term the
aggregate of interacting conspecific
demes apopulation. Populations can, of
course, be comprised of coevolutionarily
interacting conspecific hermaphrodites
or nonsexual organisms. Groups of inter-
acting populations, among which no
gene exchange is taking place but whose
demography or gene pools are affected
by the interaction, are termed commu-
nities. In the usual case, these are gener-
ally not conspecific populations. By this
definition, the simplest community
would be comprised of two genetically
isolated, but coevolutionarily inter-
acting, non-conspecific populations. De-
pending on the exact criteria one uses to
define species, one can conceive of the
situation where two populations of the
same species are essentially reproduc-
tively isolated, yet co-occur, e.g., in-
cipient sympatric speciation (White
1978). These populations might affect
each other's reproductive success and
are, thus, acting qualitatively no dif-
ferently from the interspecific inter-
actions we use to define community. De-
spite this, we opt to designate the
community as a multispecies entity.
The expanded sense of our definition
of community implies that if every orga-
nism in the world is linked with every
other organism through coevolutionary
interactions, no matter how remote, they
all belong to a large, but scientifically un-
interesting, community. As we detail be-
low, it is coevolutionary interactions
more proximate than those implied
above that are of interest and measurable
by scientists, but the possibility of
a "world community" cannot be dis-
missed a priori. The existence of a world
community is probably untestable and of
little meaning in the time frame that we
observe organisms.
Our definition of community differs
from those extant. For example, Odum
702
(1971) stated: "... community in the
ecological sense (sometimes designated
'biotic community') includes all of the
populations occupying a given area."
Conceptually, we do not use a spatial
boundary, but rather a coevolutionary-
effect boundary. Although it is a truism
that a community occupies a finite space,
the only a priori spatial boundaries for
the communities we study are those of
the biosphere itself. A recent definition
of community (Connell and Slatyer 1977)
implies some of what we propose here:
"We define a community as the set of or-
ganisms that occur together and that sig-
nificantly affect each other's distribution
and abundance. It is the interactions that
make a community a worthy unit of
study" (see also Curtis 1959, Menge and
Sutherland 1976).
This emphasis on interaction repre-
sents a common, though often unspoken,
assumption in definitions of community
and population. In our definition, inter-
specific coevolutionary interaction is a
requirement for membership in a com-
munity (intraspecific interaction for pop-
ulation). Any a priori spatial bounda-
ries, other than those of the biosphere,
are strictly arbitrary. Organisms might
co-occur in space and time yet not be
connected by coevolutionary inter-
actions; they, then, do not belong to the
same community (or population) as we
define it.
The choice of coevolutionary-effect
boundaries depends on the scientific
questions addressed by the investigator.
The vagility, longevity, and other life
history characteristics of the organisms
of interest to the researcher will combine
with the strength of interaction he or she
deems significant to determine the space-
time dimensions studied. Conspicuous
organisms of low vagility that have short
generation times (relative to the period
of observation) and are involved in inter-
actions easily observed are those most
favorable for studies of coevolutionary
relationships.
Arbitrary truncation of a coevolution-
ary unit in space may be acceptable if the
investigator adequately accounts for the
interactions crossing the boundaries. In
many studies, the community is parti-
tioned taxonomically, and one phyloge-
netic group (taxocene) is studied. Such
studies are also acceptable, as long as
the investigator is mindful that the inter-
actions with organisms outside that tax-
ocene may be coevolutionarily impor-
tant. The resultant increased tractability
may be worth the reduced coevolution-
ary resolution.
Matter-EnergyExchange
Relationships
Our greatest departure from tradition-
al thinking comes in the removal of com-
munity and population from in-line, sub-
ecosystem levels. The reasons for this
include the fact that the term interaction
has had at least two different meanings
for ecologists. One meaning is in the
sense of effects of one organism or popu-
lation on the expected reproductive suc-
cess of another. In the preceding section,
we suggested that such interactions char-
acterize coevolutionary systems. The
second meaning suggests that environ-
mental interactions consist of matter or
energy flow across the surface of an or-
ganism, e.g., ". . . in the functional
sense, an interaction represents the
transfer of energy or mass between an
organism and its environment" (Spomer
1973).
Simply stated, an ecosystem is a set of
organisms and inanimate entities con-
nected by exchanges of matter or ener-
gy. The entities exchanging and the
items being exchanged together consti-
tute the ecosystem components. In gen-
eral, the strongest interactions occur
among spatially proximate ecosystem
components.
Atmospheric and oceanic circulation
tie all the earth's organisms into a com-
mon ecosystem known as the biosphere.
The biosphere is an open system, e.g.,
radiation impinges on the earth from out-
er space and in turn reradiates away
from the earth. All lesser ecosystems
are, therefore, also open systems; spatial
boundaries for them are strictly arbi-
trary. By our definition, ecosystems con-
taining a single organism are conceiv-
able. This is contrary to the traditional
view (Odum 1971, Whittaker 1975) that
an ecosystem consists of a community
plus its environment (which is true in
some cases).
The biosphere is not merely the largest
terrestrial ecosystem. It differs qualita-
tively from lesser ecosystems in that
biotic flux across its boundaries is negli-
gible, though abiotic flux is not. The bio-
sphere also differs from any "world com-
munity" in that the latter, by definition,
is a closed system.
As it is for the community, the choice
of ecosystem boundaries, in practice, is
investigator-dependent. Scientific gener-
alization depends on the recurrence of
similar units in space or time. Ecosystem
units are frequently chosen by spatial
discontinuities in such features as physi-
ognomy, physiography, taxonomic com-
BioScience Vol. 28 No. 11
position or any environmental factor.
Temporal discontinuities are manifest in
such regular phenomena as diurnality,
lunar cycles, seasonality, etc. Tempo-
rally irregular events may be important
to some ecosystems, e.g., those follow-
ing a pulse-reserve paradigm (Noy-Meir
1973), but such events are difficult to use
for bounding ecosystems.
IMPLICATIONS
We suggest that our schema has some
conceptual advantages over those pre-
viously proposed and that our organism-
centered approach is a step closer to de-
picting realistically the mode of opera-
tion of organisms in nature.
Our selection of the term coevolution-
ary relationships (rather than evolution-
ary relationships) should not be read as a
denial of the importance of abiotic forces
in the evolutionary process. Rather, we
chose our terminology to underscore the
fact that evolutionary change in popu-
lations occurs in a community context,
not in a biotic vacuum. The term coevo-
lution has traditionally been applied to
adaptations, often complex, resulting
from reciprocal biotic interactions. We
see these as merely the extreme and ob-
vious outcomes of the processes we label
as coevolutionary.
Knowledge of the coevolutionary lega-
cy from previous generations is impor-
tant to understanding the organization of
extant demes, populations, and commu-
nities. The Founder Principle (Mayr
1942) in population biology and the role
of historical factors in ecology are ex-
amples of such legacies. Our concept of
coevolutionary relationships does not re-
quire that the constituent units affect
each other reciprocally, only that each
unit affects or is affected by another. An
extreme interpretation might suggest
that, since previous generations are co-
evolutionarily related to current ones via
one-way interactions, then dead orga-
nisms belong with living ones in coevolu-
tionary units. Although extant coevolu-
tionary units cannot be perfectly
understood without considering their an-
cestry, most studies emphasize inter-
actions among living organisms.
Potentially the most controversial por-
tion of our schema, in view of contempo-
rary definitions, comes with our use of
the terms community and ecosystem.'
The reason we feel compelled to break
with tradition (though one based on im-
precise definition and usage) is that we
believe that much disagreement by ecol-
ogists regarding the nature and definition
November 1978
of communities and ecosystems is, in
fact, a semantic problem confounded by
arbitrarily delimiting entities in nature
that are weakly differentiated at best.
Thus, if one chooses to study a defined
plot of ground, say a hectare of desert, to
elucidate the "structure and function" of
a desert ecosystem or community, one
has set an arbitrary boundary for all
processes of interest. The organisms af-
fecting the matter and energy flow of the
hectare may differ from those organisms
that affect the expected reproductive
success of the component organisms.
Organisms common to both groups may
still differ in the relative importance of
their ecosystem vs. community effects.
For example, pollinators affect the re-
productive success of plants out of pro-
portion to their impact on the plants'
matter or energy exchange. Hence, one
should be cognizant that an understand-
ing of ecosystem structure and function
need not always provide insight into
community properties; the inverse is al-
so true. When organisms occupy proxi-
mate positions in the space-time contin-
uum, the probability of their being
coevolutionarily related or connected by
the flow of matter and energy increases.
This is especially true in the latter case
because at the very least the decompos-
ing remains of dead organisms become a
diffuse ecosystem resource.
In a practical vein, contemporary use
of phrases like "bird community of
Green Canyon" are acceptable as long
as one realizes that community in that
sense is really a partial and truncated
community; i.e., the coevolutionary in-
teractions probably involve more than
just birds and more than just the orga-
nisms of Green Canyon itself. For con-
venience, one can truncate the commu-
nity in space or time or partition it
taxonomically with the proper series of
modifying words.
The term for all of the organisms of
an ecosystem by previous definitions has
usually been community. Since we do
not view a community strictly as a subset
of an ecosystem, another term must be
used. The traditional terms biota and as-
sociation might seem to obtain but these
have slightly different connotations. To
avoid coining a new word, we will simply
refer to the organisms as the biotic com-
ponents of the ecosystem.
Deductive theory of the organization
of communities, populations, or demes
(e.g., Cody and Diamond 1975, May
1976) should apply to the coevolutionary
relationships line. Testing of such theory
is usually attempted in arbitrarily delim-
ited space-time units. Such units usually
include truncated demes, populations, or
communities and not the integral coevo-
lutionary units on which the theory is
based. These truncated units may not
have the same properties as the integral
units. Furthermore, the space-time unit
may include organisms that are not inter-
acting coevolutionarily with the other or-
ganisms; the biotic diversity of such ag-
gregations may exceed that predicted by
theories based on coevolutionary inter-
action. This fundamental difference be-
tween the nature of effect and dimen-
sional boundaries makes more difficult
the task of understanding ecological
patterns.
Two interpretations of the nature of an
organism's community are possible. One
alternative defines a community as an ag-
gregation of populations connected by
statistically detectable effects on their
size or genetic composition. An orga-
nism belongs to such a community by
virtue of membership in one of those
connected populations.
The second interpretation is more
strictly organism-centered. Consider the
case of a bird. In this view, the bird's
community is composed of those other
organisms that affect its reproductive
success. In this organism-centered ex-
treme, other coevolutionary units af-
fected by the bird are not members of its
community (unless they also happen to
affect the bird); however, the bird is a
member of their communities.
Both views of the community are use-
ful. The organism-centered extreme is
the approach preferred in attempts to un-
derstand coevolution of phenotypic
characteristics, e.g.. life-history traits
(Steams 1977). Coevolutionary units not
affecting a bird's population are not im-
portant for its evolution. Excluding them
from the coevolutionarily defined com-
munity simplifies the task of understand-
ing population evolution in a community
context. The connected populations or
statistical view is the paradigm of choice
in attempts to understand community
evolution (Cody and Diamond 1975). In
this case, the focus is on the presence or
absence of species in the community and
not on the effects of the community on
the individual, with consequent popu-
lation evolution.
The emphases herein are that the orga-
nism is the central unit of biological rela-
tionships and participates in all four rela-
tionship lines simultaneously. Any line
of relationships can influence any other
through their common component level,
the organism. The same organisms that
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exchange matter or energy may affect
each other's expected reproductive suc-
cess and be phylogenetically related all
at the same time. In our organism-cen-
tered view, the environment affects all
four lines by acting directly on the
organism.
Furthermore, the organism is the fun-
damental unit of natural selection. Other
units may be significant under specific
situations, but evidence clearly indicates
that selection operates predominantly at
the organismal level (Lewontin 1970, see
also Dawkins 1976). This does not mean
that organisms can genetically contribute
to future generations only through per-
sonal fecundity; kin selection (Maynard-
Smith 1964) may also be important.
In general, the array of relationships
depicted in Fig. 1 has an implicit time di-
mension. If one starts with the physio-
logical-anatomical line and proceeds in a
counter-clockwise direction, the rela-
tionship types are ordered by the in-
creasing relative time intervals over
which important detectable effects oc-
cur. For many organisms, effect may be
felt instantaneously or over a very short
period (physiological-anatomical); on a
day-to-day, year-to-year basis (matter-
energy); on a generation-to-generation
704
basis (coevolutionary); or over many
generations (phylogenetic). In fact, these
are relative time intervals for any given
organism; the absolute time intervals of
importance are specific to the organism
in question. Thus, an event recurring at a
given temporal frequency may be dealt
with physiologically by longer-lived or-
ganisms but may produce marked popu-
lation fluctuations in shorter-lived orga-
nisms (Slobodkin and Rapoport 1974).
In summary, we propose the following
system of definitions for some ecological
aggregations of organisms:
* Deme-a group of coevolutionarily in-
teracting organisms that interbreed in a
manner approximating panmixis.
* Population-a
group of coevolutiona-
rily interacting demes or non-outcrossing
organisms, which due to a common de-
scent are genetically similar enough to be
considered conspecific, yet do not inter-
breed sufficiently to be a panmictic unit.
* Community-a group of coevolutiona-
rily interacting populations connected by
effects of one population on the demog-
raphy or genetic constitution of the
other.
* Ecosystem-the plexus
composed
both of abiotic entities and at least one
organism, which are united by the ex-
change of matter and energy.
* Biosphere-the ecosystem which in-
cludes all of the organisms on Earth dur-
ing a defined time interval.
ACKNOWLEDGMENTS
We thank our colleagues D. Andersen,
M. Caldwell, K. Dixon, J. Haefner, I.
Palmblad, and J. Warner for reviewing
an early draft of the manuscript. In addi-
tion, B. Guttman, R. McIntosh, G.
Orians, and R. Whittaker critically read
a revised manuscript and offered numer-
ous useful comments. L. Finchum typed
the various drafts. R. Bayn executed the
figure. Part of this paper was made pos-
sible by NSF Grant DEB 75-13996 to
MacMahon.
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