Evo Edu Outreach (2008) 1:16–24

DOI 10.1007/s12052-007-0016-5

ORIGINAL SCIENTIFIC ARTICLES

The Hierarchical Structure of Ecosystems:

Connections to Evolution
William Miller III

Published online: 20 November 2007

# Springer Science + Business Media, LLC 2007

Abstract Ecologic systems, which are involved mainly in Introduction

the processing of energy and materials, are actually nested
one inside another—they are simultaneously parts and This essay is about the organization of ecologic systems of
wholes. This fundamental hierarchical organization is easy all kinds—from individual organisms and their functional
to detect in nature but has been undervalued by ecologists parts up to regional ecosystems and on to the entire
as a source of new insights about the structure and biosphere. The approach will involve characterization of
development of ecosystems and as a means of understand- the basic properties of such systems. I will also explore two
ing the crucial connections between ecologic processes and rather serious misconceptions in evolutionary biology,
large-scale evolutionary patterns. These ecologic systems related to hierarchical organization. One of these is the
include individual organisms bundled into local popula- notion that everything that is interesting or significant about
tions, populations as functional components of local ecology can be explained in terms of (or reduced to)
communities or ecosystems, local systems making up the processes associated with individual organisms, or at most,
working parts of larger regional ecosystems, and so on, populations of such organisms. The other misconception is
right up to the entire biosphere. Systems at any level of embodied in the phrase “evolution takes place on an
organization can be described and interpreted based on ecologic stage.” Both of these views have not only resulted
aspects of scale (size, duration, and “membership” in more in a general underestimation of the complexity of the living
inclusive entities), integration (all the vital connections both world, but also have led to grossly inaccurate paleoecologic
at a particular focal level and across levels of hierarchical reconstructions, have made it seem that neoecology was
organization), spatiotemporal continuity (the “life history” running out of steam, and – I think most seriously – have
of each system), and boundaries (either membranes, skins, limited the growth and expansion of evolutionary theory.
or some other kind of border criterion). Considering hierar- Serious discussion of these issues began two decades
chical organization as a general feature of ecologic systems ago with a burst of activity in hierarchy theory applied to
could reinvigorate theoretical ecology, provide a realistic biology. A series of fascinating publications in the 1980s
scaling framework for paleoecologic studies, and – most summarized the characteristics of hierarchical systems,
importantly – forge new and productive connections explored the ontologic status of entities or systems at
between ecology and evolutionary theory. various levels within both genealogic (evolutionary) and
economic (ecologic) hierarchies, and began to focus on the
Keywords Hierarchy theory . Neoecology . Paleoecology . connections between the behavior of ecologic systems of
Ecologic theory . Organization and development . varied scale and evolutionary processes and patterns
Evolution . Scientific realism (Eldredge and Salthe 1984; Salthe 1985; Eldredge 1985;
Allen and Starr 1982). Later publications elaborated on
these same themes (O’Neill et al. 1986; Eldredge 1989,
W. Miller III (*)
1990; Allen and Hoekstra 1992; Ahl and Allen 1996). Any
Geology Department, Humboldt State University,
1 Harpst Street, Arcata, CA 95521, USA serious student would want to examine these contributions
e-mail: [email protected] firsthand to get an idea of the nature of the debate about the
Evo Edu Outreach (2008) 1:16–24 17

reality of hierarchical entities and the utility of a hierarchi- energy zones called riffles. Collection of aquatic organisms
cal approach to ecologic and evolutionary theory. In my from these two different fluvial settings would reveal that
view, the most reliable and useful of these guides to the two different habitats harbor several different species,
hierarchy theory are those that accept hierarchical organi- although some kinds of organisms would probably live in
zation as a real feature of living systems in general, not both places. Within any of these pools or riffles, one would
merely an empirical (and potentially disposable) position, notice immediately that some species are exceedingly
heuristic device, or convenient taxonomic system. As I show abundant and easy to collect, while others might be rare
below, biologic hierarchies are obviously real, but their and would require a bit of effort to find. With respect to one
importance has not been much appreciated. of the numerically dominant species, we could collect
enough specimens to reveal growth or developmental
stages of that particular kind of organism, or we could
Hierarchical Properties of Ecologic Systems spend time observing the behavior of living individuals
belonging to that species. So far, we would have described
Accepting a hierarchical perspective involves neither the or delineated, in the order described, local ecosystems of
abandonment of orthodox biologic orientations nor much of the stream, populations within each of these local ecosys-
a stretch of a person’s imagination. Consider a small stream tems (known in the hierarchy literature as avatars), and
together with its surrounding countryside (Figs. 1 and 2). individual organisms.
Keep in mind that the properties of the systems I am about Now, look at the same picture “in the other direction.”
to describe have to do primarily with “making a living” – The local ecosystems within the channel of the stream
the processing of energy and materials – or the economy of certainly have more properties in common than they do
nature as Charles Darwin put it (Darwin 1859). with local ecosystems of the adjacent floodplain. Such
At any particular time, the channel of the stream might closely connected local systems are termed biotope systems,
consist of quiet-water pools separated by rocky, high- the most obvious linkages in this case involving down-
Fig. 1 A hypothetical stream and
surrounding countryside. Local
ecosystems coincide with: A up-
land plateau; B talus-covered
slope and beach; C small, recent
landslide; D riffle zones in the
stream channel; E quiet water
pools; F sandy point bar; G oak–
hickory forest; H pine forest; and
I pond/freshwater marsh on the
floodplain. Diagram at bottom
shows the hierarchical position
of any of these local ecosystems
18 Evo Edu Outreach (2008) 1:16–24

Salthe 1984; Salthe 1985, 1993; Eldredge 1985). These

authors approached metasystems like the hypothetical
channel–flood plain example described above as real things
in the world, with each level of organization featuring
unique properties (Fig. 3). In doing this, they have provided
one of the clearest windows on hierarchy theory as it
applies to organization of ecologic systems. The following
is based on their work, and on my more recent papers
(Miller 1990, 1991, 1996).

Scale Probably the most obvious property of hierarchical

entities of all kinds has to do with their relative sizes and
related process rates. Large, inclusive ecologic systems
cover larger areas and usually “work slower” and last
longer than the smaller, embedded systems. Although it
Fig. 2 The same ecologic systems depicted in Fig. 1, extended may seem unnecessary to say so, any consideration of scale
through time
must also take into account membership: sometimes
component systems have exactly the same geographic
stream drift of organisms during regular flow stages and extent as the enclosing system (some avatars have the same
especially during intervals of high discharge (floods). Such distribution pattern as the encompassing local ecosystem),
closely connected local systems also share energy and but their status as components or working parts places them
materials, such as organisms normally residing in upstream at a lower hierarchical level. In addition, component systems
locations ending up as prey for downstream predators, or are characterized by faster rate constants compared to
simply as downstream transport of organic particles that enclosing systems (an avatar of salamanders may wax and
could be exploited by detritus feeders. The bars and wane in terms of abundance of individuals over the course of
beaches might also be included in this biotope if their decades, but each individual salamander completes its life
resident organisms have similar close connections to the cycle in just 2 years and physiologic processes within
channel systems. But what about local ecosystems of the salamander bodies take place on a scale of minutes to hours).
adjacent floodplain? Could they be construed as connected
in some way to the channel ecosystems, thereby forming a Integration Ecologists have devoted much effort to the
much larger regional ecosystem? Ponds and other low-lying study of interactions, from idealized pair-wise linkages
areas would receive colonists from the riffle and pool (such as competitive interaction between two different
ecosystems during major floods; organic material that avatars in the same local ecosystem) to complex networks
accumulated in such settings might be flushed into the
channel during the same events. Riparian forests could
contribute to the habitat structure of channels as trunks and
crowns of trees that have fallen into the stream; floodplain
animals could depend on fluvial invertebrates as a food
resource.
It is not hard at all to picture ecologic organization at
varied scales in this way and to see that ecologic systems
are simultaneously parts and wholes—which is what they
would have to be to qualify as real hierarchical entities
(Salthe 1985). It can also be shown that all of these systems –
big and small – interact with each other, have boundaries of
some sort, and go through something akin to “life cycles.”

Some Formalities: The Properties of Hierarchical Entities

Niles Eldredge and Stan Salthe did more than anyone else
to outline the fundamental properties of biologic hierarchies
and the characteristics of hierarchical entities (Eldredge and Fig. 3 Fundamental properties of hierarchical entities of all kinds
Evo Edu Outreach (2008) 1:16–24 19

that attempt to portray all of the components and connections ecosystems would be wiped out by this catastrophe—their
in a community or local ecosystem. This “internal wiring” “lives” would end as the component avatars underwent local
gives local ecosystems the appearance of being things extinction or individual organisms migrated to more favor-
separate from other things at the same scale and at the same able habitats. Perhaps some species from the pools and pond
time (Salthe 1985). Ecologic interactions involve the transfer environments would flourish in the new regime, together
of energy and materials—the fundamental currencies of with recruits from other areas that prefer shallow lacustrine
ecologic systems of all types and scales. settings. In short, the previous regional ecosystem would be
It is important to realize that interaction, or networking, replaced by a new one having different ecologic properties
not only takes place between entities but also across (in terms of species composition, relative abundances, and
hierarchical levels. For example, an avatar of freshwater organization of food webs). The event producing ecosystem
snails in our hypothetical example would not only derive its replacement certainly has a random quality to it, but the
essential ecologic properties from interactions with the organization and function of the new system would not look
other co-occurring avatars in a stream pool, but also from all that random.
the interaction of its internal working parts (the behaviors This new regional system might persist for centuries,
of individual snails) and from its position in the food web until the landslide deposits are washed away in a major
of the local environment (an organizational property of the flooding event. In the meantime, the lentic system would
local ecosystem). Hierarchy theorists have formalized this have experienced disturbances and recoveries at varied
complex network of interactions involving entities that are scales, caused by droughts that lowered water level,
simultaneously parts and wholes as the “triadic system of intervals of low O2, elevated turbidity, or events involving
causation” (Salthe 1985; Miller 1990): the avatar of snails short-lived invasion of predators—all of which would have
is the focal level of interest in this case, the component caused ecologic properties to fluctuate but would not have
organisms present initiating conditions (relating to possi- resulted in regional system collapse. In this view, the
bilities), and the encompassing local ecosystem presents regional system really has a life span (between establish-
boundary conditions (relating to constraints). In other ment of the lake and its eventual disappearance) and a
words, understanding the organization and development of unique life history (related to primary organization of the
avatars must include an account of one-on-one interaction component local ecosystems and to disturbances and
with other avatars, plus interaction between the avatar level recoveries affecting those component systems). When
and the level of individuals on the one hand and interaction ecologists speak of disturbance regimes, succession, and
of the avatar with the encompassing system on the other. replacement, or when they show that a community has a
Stable networks should produce stable ecologic systems. certain geographic extent on a base map, they are owning
For example, a long-lasting metapopulation structure of up to the fact – whether they realize it or not – that things
freshwater snails in a large fluvial system probably supports like local and regional ecosystems actually have spatiotem-
regional ecosystem stability over long intervals of time. poral continuity.

Spatiotemporal Continuity Genealogic entities have distinct Boundaries Because local ecosystems may have gradational
beginnings, unique histories, and eventual terminations. For borders coinciding with gradual changes in physical and
example, species have a kind of birth (speciation); a life chemical factors of the environment (called ecoclines),
history involving geographic deployment and fate of instead of always having sharp borders defined by steep
demes, ecologic involvements, and phylogenetic potential; environmental gradients (ecotones), some ecologists have
and an eventual death (species-lineage extinction). However, decided that local multispecies assemblies cannot be con-
can we say that ecologic systems have the same kind of strued as entities that are anything like an individual
spatiotemporal extent (Fig. 2)? Many ecologists would organism. However, ecologic systems should be expected
simply preempt the discussion here by saying “no”— to have different kinds of boundaries, depending on the type
claiming that most are happenstance, fluctuating assemblies and scale of the system, compared to more familiar
of organisms thrown together by random processes. I think individual organisms with their skins and membranes. In
there is more to it than this. Picture again the stream and some cases, abrupt change in the features of the enclosing
flood plain example (Fig. 1). Imagine a large landslide environment reveals (or coincides with) the borders of local
derived from a rocky cliff bordering one side of the stream and regional ecosystems. It might be more appropriate,
and covering part of the valley floor. An event such as this however, to “map” such entities using a kind of circuit
could dam the channel and produce a shallow lake, causing diagram, showing which subsystems are included and which
inundation of the flood plain and conversion of the lotic are excluded from the dynamic structure of a system of
(flowing water) environment of the channel to a lentic interactors. It would not matter if the positions or intensities
(standing water) environment. Many of the established local of boundaries changed during system development. They
20 Evo Edu Outreach (2008) 1:16–24

certainly do with organisms (think about the life cycle of a hierarchical perspective, but without seriously considering
colonial animal, such as a coral, involving growth, morphol- the ontologic reality of the nested systems they had
ogic change, and expansion on the seafloor). Moreover, we encountered—and therefore never suspected that important
begin to lose concern for the “boundary problem” when we processes and patterns unique to different levels of organi-
look at satellite images of North America or Europe, which zation might have been overlooked or that crucial connec-
reveal the extent of different types of forests and grasslands tions to evolutionary theory might have been missed.
controlled largely by climate, geology, and evolutionary A consideration of larger, more inclusive units of
history of the plant groups—large, fairly long-lived systems organization is clearly needed in some cases. For example,
apparently having geographic borders or transition zones how can we talk about the global carbon cycle and climate
coinciding with entity boundaries (some of which are change only in terms of individuals and populations? How
produced by human activity). can we determine the processes that result in the stable
In the hypothetical channel–flood plain regional system, species-lineages of punctuated equilibria without consider-
a stroll through the woods to the edge of the steam would ing the properties of the enclosing regional or provincial
require either a change of route or else a splash in the water ecosystems, which sometimes also appear to be stable over
(detection of an abrupt system boundary we could easily millions of years? What happens when climatic or geologic
plot on a base map). Noticing the more subtle differences in processes lead to reorganization or replacement of these
species composition and relative abundance within the regional systems, producing waves of abandonment, inva-
forested parts of the flood plain could reveal similar, if less sion, extinction, and especially speciation events? What are
dramatic, transitions corresponding to much more gradual the major ecologic controls of species richness and
changes in soil type, moisture, topography and disturbance development of clades over even longer time spans and
regime. In all these cases, we are confirming the boundary over larger geographic areas? Reductionism does not serve
criterion. us very well in these cases.
A related problem has affected the development of
paleoecology. Beginning in the 1960s, many marine
paleoecologists interpreted vertical sequences of fossil
Some Consequences of Ignoring Hierarchical assemblages as examples of ecologic succession (initiation
Organization or reestablishment and maturation of local ecosystems
controlled mostly by internal processes)—no matter how
Attempting to practice ecology admitting only individuals thick the associated stratigraphic succession, what type of
and populations as units of description and interpretation paleoenvironmental context, or how drastic the changes in
has turned out to be a rather serious form of scientific the composition of the fossil assemblages. This obviously
tunnel vision. For example, many community ecologists was not right. It is now accepted that most of these
have always doubted the existence of assembly rules or purported cases of succession in the fossil record were
emergent properties of various kinds of multispecies really replacement sequences consisting of the remains of
aggregates. Behavior of individual organisms and popula- different local or regional ecosystems, paced by environ-
tion-level processes explained everything. (It is noteworthy mental fluctuations often caused by large-scale climatic and
that the systems ecologists never followed this route; but it geologic processes (Miller 1986). Notwithstanding this
is curious that they never really explored the ontologic fundamental scaling correction, many paleoecologists con-
properties of the large dynamic systems they analyzed.) In tinue to refer to almost any sample of fossils – of almost
the late twentieth century, as ecology became preoccupied any size – as a “community.” Is it any wonder that much of
with mathematical modeling, some practitioners began to the rich literature of paleoecology has been ignored by
think their discipline had progressed (theoretically, concep- neoecologists?
tually) about as far as it could go. Reinvigoration of
ecology came as workers “scaled up” their studies and
started to think seriously about the consequences of historic Rewards for Getting it Right
legacy and contingency. New approaches included work on
metapopulation networks, “metacommunities” and regional It is an odd fact that ecology and evolutionary biology
trophic webs, large-scale distribution patterns of species, developed as essentially separate fields during most of the
and the long-term development of landscapes, often twentieth century. Other than the notion of adaptation (a
pursued in the cause of conservation ecology (Likens term used in many ways, both as verb and noun—process
1989; Delcourt and Delcourt 1991; Gilpin and Hanski and product), evolutionary theorists had little of substance
1991; Brown 1995; Gaston and Blackburn 2000; Polis et al. to say about the connections between the economic (ecologic)
2004; Holyoak et al. 2005). Ecologists were embracing a and genealogic (evolutionary) realms of life. The most
Evo Edu Outreach (2008) 1:16–24 21

Fig. 4 Species-lineage extinc-

tions and speciation events dur-
ing a turnover pulse; invasions
and abandonments would also
occur. The varied patterns in-
clude: A extinction of an abun-
dant, ecologically dominant
species; B extinction of a mod-
erately abundant species; C rare
species originating from an
abundant ancestor; D an abun-
dant species that undergoes a
reduction in abundance, but
subsequently recovers; E for-
merly abundant species reduced
to rarity; F a rare species that
persists through the turnover
pulse; G a rare species that
becomes abundant and ecologi-
cally dominant in the subsequent
regime; H an abundant species
derived from a rare ancestral
species; I a rare species that
vanishes early in the turnover
pulse; and J a rare species yield-
ing many descendant species
(based on Miller 2005, Fig. 1)

important property of life was the reproduction of fertile large, inclusive regional ecosystems. At this higher level of
offspring, as well as the perfection and spread of such organization, we must look for novel processes that
reproductive systems. Ecology was seen as mostly the maintain composition and organization over times spans
backdrop to these most essential processes. Although a in the order of hundreds-of-thousands to millions of years
reunion of evolutionary biology and neoecology has been (Miller 1996)—opening nothing less than a new frontier for
heralded in recent articles and books (reviewed in Johnson ecologic theory.
and Stinchcombe 2003), the most dramatic breakthroughs A related development has to do with regional extinc-
seem to be coming from reevaluation of the fossil record tions and waves of more or less concurrent speciation
from a hierarchical perspective. events that mark the intervals of reorganization/replacement
One of the surprises in the recent paleoecologic literature
was a series of papers and an entire journal issue (Ivany and
Schopf 1996) evaluating a pattern of apparent ecosystem
stability lasting, in some cases, for millions of years,
referred to as “coordinated stasis.” It appears that large
marine ecologic systems can be exceedingly durable,
reminiscent of (and probably related to) the stable species
lineages of punctuated equilibria. This pattern of paleoecol-
ogic stasis was detected in the “deep” fossil record, but is
not in accordance with patterns of repeated reassortment of
terrestrial species documented by ecologists over the last
few tens-of-thousands of years, paced by late Pleistocene–
Holocene climate changes. The difference in perspective
between neo- and paleoecology needs to be worked out, Fig. 5 The sloshing bucket model of evolution. A represents the
both here and with respect to other observations and frequency of disturbances of different intensities and geographic scope
through time, N; B represents the accumulation of adaptive speciation
conceptual issues. What we seem to be seeing in the fossil during the Phanerozoic, S. Note that, in this view, most speciation is
record, however, is a pattern of stability not associated with associated with intermediate levels of disturbance, not with “back-
communities or local ecosystems but, rather, involving ground” processes or with global mass extinctions (Miller 2004, Fig. 1)
22 Evo Edu Outreach (2008) 1:16–24

of these same regional ecosystems (Figs. 4, 5, and 6). However, the regional turnovers are both hundreds of times
Elisabeth Vrba (1980, 1985, 2005) was the first to draw more frequent than mass extinction and represent the
attention to these events, referring to them as turnover thresholds where invasions/abandonments and extinctions/
pulses. Eldredge (1999, 2003) has built on this theme with speciation events start to happen. In his thinking, nothing
his sloshing bucket model of evolution. In his broad view of much happens in terms of adaptive evolution until the
evolutionary theory, the larger the environmental jolt, the regional systems get into trouble—a connection between
bigger the evolutionary reaction, an inherently hierarchical ecologic and evolutionary systems I call macroevolutionary
approach. Small disturbances produce little in the way of consonance (Miller 2004). In other words, most of the
phenotypic evolution, and global-scale mass extinctions are species-level evolution during the Phanerozoic Eon (i.e.,
very rare—albeit associated with some of the grandest during the time when the fossil record has been rich enough
evolutionary transformations in the history of life on Earth. to record these patterns) probably occurred at the interme-

Fig. 6 The ecologic structure of

turnover pulses. a Reduction of
an abundant, widespread species
(a metapopulation consisting of
sources and sink populations, 1–
5) to a few isolated remnants (6–
8)—a situation likely to promote
a bout of allopatric speciation (if
the population remnants are not
too small). b Simultaneous col-
lapse of numerous local ecosys-
tems (1) that were connected in
various ways (2) to produce a
regional network, resulting in a
few disconnected remnants (3).
c Turnover in terms of changes
in relative abundance of the
resident organisms (a–j), with
some rare species becoming
abundant (1), prominent species
reduced in abundance (2), and
reduction or complete elimina-
tion of other species (3)
Evo Edu Outreach (2008) 1:16–24 23

diate scale of the sloshing bucket (Fig. 5)—not during the systems and experience at least some of their basic
longer “background” intervals or in the recovery phases properties. We know what the possibilities are: the question
following mass extinctions. This is an extremely interesting becomes, how can we use this more inclusive view of life
idea that needs more empirical work, but it would never to expand and improve ecologic and evolutionary theory, to
have been conceived in the first place without the kind of make instruction in evolutionary biology more comprehen-
hierarchical perspective I have described. I think these sive and realistic, and to forge new and mutually enriching
developments clearly demonstrate that evolution does not connections to related disciplines?
simply take place on an ecologic stage; evolutionary and
ecologic processes are interconnected and interwoven at Acknowledgements This essay is based on a presentation in the
Hierarchy Symposium, held as part of the Genova Festival di Scienza,
varied scales, often take place concurrently and in coordina- November 2006. I thank Niles Eldredge and Telmo Pievani for their
tion, and simultaneously propel and constrain one another in invitation to participate, and the organizers for their generous support.
ways we have barely imagined when it comes to turnovers, Eldredge and Bud Rollins provided insightful reviews.
extinctions, and appearance of new species (Miller 2004).

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