The Ecosystem Concept

Gaston E Small, Department of Biology, University of St. Thomas, Saint Paul, MN, United States
Jane Shevtsov, Life Sciences Core Education, University of California Los Angeles, Los Angeles, CA, United States
r 2024 Elsevier Inc. All rights reserved.

This is a update to Eugene P. Odum, Ecosystem, Concept of, Editor(s): Simon A Levin, Encyclopedia of Biodiversity (Second Edition), Academic
Press, 2001, Pages 59–63, ISBN 9780123847201, https://doi.org/10.1016/B978-0-12-384719-5.00040-X.

Introduction 326
History of the Ecosystem Concept 327
The Stock-and-Flow Conceptualization of Ecosystems 328
Structural Conceptualization of Ecosystems 329
Ecosystem Engineers 330
Critical Transitions 330
Applications of the Ecosystem Concept to the Human-Dominated Environment, and Implications for the Future of Human
Life on (and Beyond) Earth 332
Conclusions 333
References 333
Further Reading 334

Abstract
The ecosystem concept encompasses relationships among different species while also including non-living components. The term
“ecosystem” was first used by Tansley, in 1935, but the concept was described in the late 19th Century. One major usage of the term,
advanced by E.P. Odum, has been the stock-and-flow conceptualization, which explicitly analyzes the movement of material or energy
through various ecosystem compartments. A structural conceptualization of the term has also become common, with the term
“ecosystem” referring to a local instance of a biome. The ecosystem concept was originally developed for natural systems but can also
be applied to human-dominated environments.

Key Points
• The ecosystem concept encompasses relationships among different species while also including non-living components
that are sources of material and energy.
• The term “ecosystem” was first used by the British plant ecologist A.G. Tansley in 1935, but the underlying concept was
described in the late 19th Century.
• One major usage of the term, advanced by E.P. Odum, has been the stock-and-flow conceptualization, which explicitly
analyzes the movement of material or energy through various ecosystem compartments.
• A structural conceptualization of the term has also become common, with the term “ecosystem” referring to a local
instance of a biome.
• The ecosystem concept was originally developed for natural systems but can also be applied to human-dominated
environments, yielding insights relevant to sustainability.

Introduction

The ecosystem concept incorporates the biotic and abiotic components that interact to allow life to exist. The concept encompasses
the ecological community (consisting of populations of different species, and the interactions among those organisms) and it also
includes non-living components that act as reservoirs of material and energy. Soil, water, air, and sunlight are essential compo-
nents of many ecosystems. Thus, the ecosystem is the first level along the biological hierarchy that has all of the ingredients
required for life. Whereas the study of populations focuses on numbers of organisms, and the metrics of community ecology
include numbers of different species, ecosystem ecology typically focuses on metrics such as biomass, energy, and mass of carbon
and nutrients.
In contrast to landscape ecology, which explicitly focuses on spatial heterogeneity, the ecosystem concept is generally not
spatially explicit, as it implicitly assumes homogeneity within the defined area. There is also no inherent size to an ecosystem.

326 Encyclopedia of Biodiversity 3rd edition, Volume 6 doi:10.1016/B978-0-12-822562-2.00341-8

 The Ecosystem Concept 327

Ecosystems can be thousands of square kilometers (e.g., the Greater Yellowstone Ecosystem) or as small as several milliliters of
water contained in a pitcher plant. Ecosystems can be naturally bounded (e.g., an island, a lake, a watershed) or the concept can be
applied to arbitrary spatial units (e.g.,1000 ha of forest, or 10 km3 of ocean). The biosphere (or ecosphere) is the part of Earth
where life exists and may be considered the worldwide sum of all ecosystems, or the largest ecosystem.
Over the past century, the ecosystem concept has developed and expanded through theoretical and empirical advances. In this
entry, we will review the history of the ecosystem concept and explore two different conceptualizations of the term: a stock-and-
flow ecosystem concept focusing on storage and movement of material and energy, and a structural conceptualization of the term
focusing on dominance of a particular (typically plant) species. Finally, we will explore the application of the ecosystem concept to
the highly engineered systems that support the vast majority of humans on Earth today, and we will examine the value of this
concept for informing sustainability decisions on Earth and for potentially supporting human life beyond Earth in the future.

History of the Ecosystem Concept

The term “ecosystem” was first used by British Ecologist Tansley (1935), who argued that, while biologists’ attention may focus on
organisms, these organisms cannot be understood without consideration of the complex of physical factors forming their
environment. The concept of the ecosystem pre-dates the use of the term, however. In 1877, German biologist Karl Möbius
described the community of organisms in an oyster reef as a “biocoenosis”, alluding to the tightly coupled nature of this level of
organization. A decade later, the American biologist S.A. Forbes provided the classic example of “the lake as a microcosm”,
describing the interplay between physical and biological factors. In 1942, the publication of Raymond Lindeman’s “The Trophic-
Dynamic Aspect of Ecology” described for the first time the coupled movement of both energy and nutrients through multiple
trophic pathways within a lake (notably, a relatively closed ecosystem), with the microbially-dominated “ooze” playing a central
role. The Soviet geochemist Vernadsky (1929) developed the concept of the biosphere, and fellow Soviet ecologist Sukachev
(1959) expanded on the terminology of Möbius in describing the concept of “geobiocoenosis,” further integrating biogeochemical
cycling into the ecosystem concept.
The use of the ecosystem concept as a central organizing framework in Odum’s popular Fundamentals of Ecology textbook
(originally published in 1953) helped spur rapid growth in empirical and theoretical ecosystem research. Yet, following this
growth, ecologists have not had a common understanding of the concept (Golley, 1993). One usage has been what we refer to
hereafter as the “stock-and-flow conceptualization,” focusing on the storage and flows of elements or energy among biotic and
abiotic compartments. Another, more general usage has been what we refer to as the “structural ecosystem concept”, referring to a
local instance of a biome. Tansley, in his 1935 paper that introduced the term, foreshadowed each of these different usages. He
wrote that “the more fundamental conception is, as it seems to me, the whole system (in the sense of physics), including not only
the organism-complex, but also the whole complex of physical factors forming what we call the environment of the biome.” But
Tansley also described the term “ecosystem” as a category equal in rank to the biome and acknowledged that the term could be
used in a general sense, similar to the term “community”.
Shortly after the end of World War II and the development of nuclear weapons, the study of energy and nutrient flow in
ecosystems gained both new tools (radioactive tracers) and new support from the US government. For example, influential studies
of energy flow in coral reef ecosystems were carried out at Enewetak Atoll, a US nuclear test site in the Marshall Islands (Odum and
Odum, 1955). Other such studies, done with and without radioactive tracer methods, followed (e.g., Odum, 1957; Teal, 1962;
Tilly, 1968). They aimed at mathematically modeling energy and nutrient flows in comprehensive ways, taking all organisms and
relevant abiotic components, into account. There were also attempts to formulate general principles of ecosystem development
and energy flow (e.g., Odum, 1969; Odum and Pinkerton, 1955; Patten and Odum, 1981). This line of research culminated in the
International Biological Program of 1964–1974.
This research program was not perceived as successful at the time. It collected large amounts of data, but computational
modeling was still difficult, with programs and data still often entered on punched cards and simple simulations taking hours or
days to run. Model predictions were sometimes thrown off by parameters that no one had thought to measure or that had not
been measured with enough precision (Golley, 1996; Hagen, 1992). With hindsight, we might say that these early ecosystem
ecologists attempted too much, too soon. Younger ecologists increasingly turned to population and community studies and the
ecosystem approach, while never disappearing, went out of fashion.
In this context, there emerged critiques and defenses of ecosystem research and the ecosystem concept itself that were essentially
philosophical, written by both practicing ecologists and philosophers of science. For example, in a prominent critique of the ecosystem
concept, O’Neil (2001) described and criticized the traditional view of ecosystems as being (mostly) spatially closed and homogenous,
treating species as substitutable, neglecting natural selection and human influence. He also considered the concept of stability to be
ambiguous, although much of the ambiguity he pointed out appears to stem from the implicit assumption that a stable system should be
able to recover from any disturbance. As a solution, he proposed a highly species- and population-centric approach to ecosystems related
to what we call the structural conception, but this conceptualization appeared to omit energy and nutrient flows.
A similar critique offered by the philosopher Mark Sagoff (2003) attracted a response from deLaplante and Odenbaugh (2006).
They pointed out that “ecosystem” can refer to: (1) an object of scientific study; (2) theories of the nature of such objects; or (3) a
general methodology for doing science. deLaplante and Odenbaugh identified three commonly cited false dichotomies: holism vs.
reductionism, Clementsian vs. Gleasonian community structures, and the combinatorial assumption that the first in each pair
328 The Ecosystem Concept

implies the second. They point out that, as with key concepts in other sciences, it may not be necessary to precisely define
“ecosystem” or be able to put unambiguous boundaries around one in order to do productive research on ecosystems. Also, self-
organization and complexity are important features of ecosystems and their study can provide insight into ecology.
In the 75th Tansley review in the journal New Phytologist, Currie (2011) reached conclusions similar to those of de Laplante and
Odenbaugh; namely, that although we usually cannot draw precise boundaries around ecosystems, the concept still makes sense.
Ecosystem processes, which are characterized by biotic-abiotic coupling, occur at a range of scales. Self-organization is important
and humans are a part of many ecosystems.
The utility of the ecosystem concept in various applications is the focus of the remainder of this article.

The Stock-and-Flow Conceptualization of Ecosystems

One conceptualization of the ecosystem focuses on flows of energy and/or material into and out of the system (however defined),
and among compartments (or stocks) within the system. This conceptualization builds upon laws of thermodynamics and
conservation of mass.
After defining the boundaries of the ecosystem (which may be natural or arbitrary), one or multiple currencies of interest are
selected (e.g., energy, carbon, nitrogen, phosphorus, water, etc.). Compartments are delineated such that all of the currency of interest
within the system boundary is accounted for. The appropriate resolution (the number of compartments and identity of those
compartments) depends on the purpose of the model. For example, a model of a lake ecosystem with a purpose of understanding the
effects of harvesting on dynamics of different fish species may represent all phytoplankton in a single compartment while repre-
senting different fish species as individual compartments, whereas a lake model with a purpose of understanding how warmer spring
temperatures affect algal blooms may need to explicitly represent various algal taxa as individual compartments, but may require less
resolution for higher trophic levels. The benefits of increasing model resolution may tend to peak at intermediate complexity and
decline with additional complexity (Jørgensen, 1988; Costanza and Sklar, 1985), underscoring the trade-off for the modeler between
knowing much about little or knowing little about much (Jørgensen and Bendricchio, 2001). Once compartments (also known as
stocks, or state variables) are defined, flows of the currency of interest are then identified and measured or simulated (Fig. 1).

Fig. 1 An ecosystem model illustrating flows of materials (thick lines) and energy (thin lines) within an ecosystem. Autotrophs are represented as
the compartment labeled “A”; heterotrophs are represented as compartments labeled “H”. Soil (“S”) plays an important role in the recycling of
nutrients within the ecosystem. Reprinted from Odum, E. P. (2001). Concept of ecosystem. Encyclopedia of biodiversity, vol. 2, 305–310.
 The Ecosystem Concept 329

Fig. 2 A conceptualization of an ecosystem illustrating the exchange of energy and materials with the external environment. Flows of energy and materials
within the ecosystem are not depicted in this diagram. Reprinted from Odum, E. P. (2001). Concept of ecosystem. Encyclopedia of biodiversity, vol. 2,
305–310.

However the ecosystem is defined, external inputs of energy and materials, dissipation of energy heat (entropy), and export of
materials occur to varying extents (Fig. 2). Energy inputs are typically derived from solar radiation, but interesting exceptions
include deep sea hydrothermal vents (where food webs are supported by microbial metabolism of hydrogen sulfide) and cave
ecosystems which are supported by inputs of terrestrial organic matter (ultimately derived from sunlight). The degree of internal
cycling relative to external exchange depends in part on the conceptual boundaries of the ecosystem and also on ecological
dynamics inherent to the system. Odum (1969) formalized the role of succession in ecosystem development, describing general
predictions of 24 different ecosystem properties in developmental and mature successional stages. Mature ecosystems, Odum
argued, are characterized by increased complexity, higher biomass relative to production, and higher internal cycling. Subsequent
empirical studies have generally supported Odum’s characterization (Corman et al., 2019).
The stock-and-flow conceptualization of ecosystems is perhaps best illustrated empirically by studies that systematically
quantified food web structure and energy flows through a coral reef on Enewetok Atoll (Odum and Odum, 1955); Silver Springs in
central Florida (Odum, 1957), a salt-marsh ecosystem in coastal Georgia (Teal, 1962), and the Chesapeake Bay (Baird and
Ulanowicz, 1989). The use of isotope tracers has been used to elucidate the movement of nutrients along a stream and within
stream food webs (Newbold et al., 1983) led to the formalization of the concept of “nutrient spiraling”, describing the combi-
nation of downstream hydrologic transport coupled with biological uptake and recycling.
This stock-and-flow conceptualization of ecosystems also underlies (at least implicitly) ecosystem manipulation experiments,
such as clearcutting a watershed to examine effects on nitrate export (Likens et al., 1970), a long-term phosphorus fertilization of a
lake to investigate effects on phytoplankton communities and nitrogen dynamics (Schindler et al., 2008), and excluding terrestrial
leaf litter from a headwater stream to examine the effects on the invertebrate food web (Wallace et al., 1997). More recently, the
development of ecological stoichiometry (Sterner and Elser, 2003) leverages mass balance constraints inherent in considering
multiple currencies (typically an element that limits production, and another element in excess) to make predictions about
retention and recycling of nutrients within ecosystems.
The stock-and-flow conceptualization of ecosystems has also been foundational in theoretical advances in systems ecology
(Ulanowicz, 2004). Ecosystem budgets (whether determined empirically or estimated from simulations) can be written as
matrices, where each entry gives the flow rate from one compartment to another. These budget models can then be iterated to
obtain long-term behavior. This iteration allows us to use tools from linear algebra to study currency flows within the ecosystem.
Such studies have led to insights such as showing that energy or material cycling allows flows to return to compartments multiple
times and become evenly distributed in the network, and that these indirect effects a higher degree of mutualism within networks
than is apparent from direct interactions alone (Fath and Patten, 1999).

Structural Conceptualization of Ecosystems

Although the textbook definition of ecosystem ecology is the stock-and-flow concept, the term is also used more generally. One of the most
common is speaking of a “forest ecosystem” or a “coral reef ecosystem.” Here, the word “ecosystem” refers to a local instance of a biome.
This conception of ecosystems is species-centric but differs from community ecology in focusing on foundational species – for
example, the trees in a forest or the corals in a reef – without much attention to species identity. In this approach, the replacement
330 The Ecosystem Concept

of a foundational species by other functionally similar species (e.g., the replacement of the American chestnut, Castanea dentata, by
other deciduous trees in Appalachian forests) does not change the identity of the ecosystem per se. By contrast, a lake that shifts
from a clear water, macrophyte-dominated state to a turbid, phytoplankton-dominated state has become a structurally different
ecosystem. Thus, we will refer to this as the structural concept of ecosystems.
Despite its difference from the stock-and-flow ecosystem concept, the structural conceptualization is also a genuine ecosystem
concept because it often refers to internally driven dynamics and responses to environmental change that are more than the sum of
species’ responses. Such applications include the concepts of ecosystem engineers and regime shifts or critical transitions in
ecosystems.

Ecosystem Engineers
Although the fact that organisms affect their environment had been recognized and specific cases have been studied since the
beginning of ecology, the term “ecosystem engineer” was not formally coined until 1994. Jones et al. defined ecosystem engineers
as organisms that affect the availability of a resource (such as light, water, space or nutrients) to other organisms, either by their
own bodies or by modifying the abiotic environment in some way. Notably, just serving as a resource does not make an organism
an ecosystem engineer; rather, it must either alter the availability of an abiotic resource or modify a consumer-resource interaction
(Hastings et al., 2007; Jones et al., 1994).
Beavers are the classic example of an ecosystem engineer. By felling trees and blocking water flow, beavers fundamentally
modify their abiotic environment; they create ponds and wetlands, changing the flows of water and sediment, which has a range of
effects throughout the whole ecosystem. The presence of beavers even reduces the susceptibility of forests to fire (Fairfax and
Whittle, 2020).
Marine environments can also contain ecosystem engineers. Jones et al. (1994) listed marine meiofauna and zooplankton as
ecosystem engineers, in part because of their production of fecal pellets that enhance nutrient transport. On a larger scale, Roman
et al. (2014) and (Willis, 2014) described whales as ecosystem engineers for the same reason. The “whale pump” brings N and Fe
from the deep ocean to the photic zone, where they can be used by phytoplankton. In addition, whales can stir ocean waters
sufficiently to impact primary productivity by increasing the availability of nutrients (Roman et al., 2014).
Other ecosystem engineers included in the original list of Jones et al. (1994) are snails and bagworm caterpillars that eat
endolithic lichens and the rocks they grow in, increasing nutrient cycling rates. By that logic, it makes sense to include mycorrhizal
fungi, particularly those that directly break down rock, as ecosystem engineers. Mycorrhizae are symbiotic associations between
fungi and plants, in which the plant provides C to the fungus and the fungus helps the plant obtain nutrients, as well as enhancing
stress resistance, drought tolerance, and various other possible effects. Some mycorrhizal fungi, specifically ectomycorrhizae, can
break down minerals, liberating the nutrients found in them and making them available to plants (van Schöll et al., 2008). Since
these fungi modulate the availability of a resource, they are also ecosystem engineers.

Critical Transitions
The critical transition concept originated from the application of nonlinear dynamics to population ecology, the classic example
being a model of spruce budworm outbreaks (Holling, 1977). Later, similar math was applied to models of lakes that could be
either plant- or algae-dominated, depending on nutrient input (Scheffer et al., 2001). The key result of this research was that regime
changes could not be reversed by undoing the increase in nutrient input that led to them. The new regime represented a stable state
and the system could only be driven back to the old one by significant reductions in nutrient input, possibly coupled with
perturbations to the ecosystem itself (e.g., algae removal).
The key concepts here are positive feedback loops and bistability. Plants in a plant-dominated lake take up nutrients, which
keeps algae populations low, which promotes plant growth by keeping water clear. On the other hand, the dense mat of algae in
an algae-dominated lake prevents light from reaching plants, and the absence of benthic plants in turn results in more nutrients
available for algae, promoting algal growth. Thus, the system has two (sometimes more) stable states maintained by these
feedback loops and will tend to stay at whichever state it currently occupies. However, when a change does occur, it can be sudden
and unexpected (Fig. 3).
Another example of critical transitions in ecosystems that is of great concern today is the Amazon rainforest (Fig. 4). Much of
the Amazon lies in a climatic region that could support either forest or savanna. However, the large amount of evapotranspiration
caused by large expanses of forest maintains the ecosystem in a forest state (Scheffer et al., 2001; Shukla et al., 1990; Staal et al.,
2020; Da Silviera Lobo Sternberg, 2001). Essentially, the forest creates its own rain via evapotranspiration and the formation of
cloud condensation nuclei (Pöschl et al., 2010). Modeling studies show that deforestation and climate change can reduce pre-
cipitation enough to change the forest to a savanna state over a short time, a very real threat. (Staal et al., 2020; Da Silviera Lobo
Sternberg, 2001).
 The Ecosystem Concept 331

Fig. 3 Illustration of how small perturbations may lead to drastic changes in ecosystem state. Reprinted with permission from Scheffer, M.,
Carpenter, S., Foley, J. A., Folke, C. and Walker, B. (2001). Catastrophic shifts in ecosystems. Nature 413, 591–596, courtesy Springer Nature BV.

Fig. 4 The high rates of evapotranspiration from Amazon rainforest produces the rainfall that maintains this forest. Extensive forest clearing may
reduce regional rainfall, driving a shift from forest to savannah in a drier climate. (Image: Jorge.kike.medina, CC BY 3.0 ohttps://creativecommons.
org/licenses/by/3.04, via Wikimedia Commons).
332 The Ecosystem Concept

Applications of the Ecosystem Concept to the Human-Dominated Environment, and Implications for the Future of
Human Life on (and Beyond) Earth

The ecosystem concept was developed through the study of pristine natural systems, but over the past century, human population
has quadrupled and human impacts are now evident throughout the biosphere. The ecosystem concept has increasing relevance
for understanding how humans interact with (and are intrinsically connected to) our environment, therefore informing decisions
about sustainability.
The advent of agriculture was transformational for humans, as humans learned to engineer ecosystems through domestication of
plants and animals so that more primary production (plant growth) and secondary production (animal growth) supported humans.
Today, humans appropriate approximately 38% of global terrestrial net primary production (Running, 2012), and the biomass of
humans and livestock far exceeds the biomass of wild mammals on Earth (Bar-On et al., 2018). Pre-industrial agriculture (and some
organic farms today) featured high diversity of crops and integration of nutrient recycling by grazers, resulting in farms that
functioned somewhat similarly to natural ecosystems. Modern industrial agriculture, by contrast, is characterized by low biodiversity
(typically monoculture), managed to maximize productivity (crop yield) through high inputs of fertilizers and pesticides. Odum
(1969) pointed out the difference between these industrial ecosystems and natural ecosystems which support most life on Earth, and
typically have high biodiversity, high biomass relative to productivity, and high rates of internal nutrient cycling.
Cities can also be viewed through the lens of ecosystem ecology. Cities are inherently open systems, relying on inputs of energy,
food, and raw materials from the hinterlands, and exporting waste products. For this reason, (Odum, 1989) referred to cities as
“parasites on the landscape”. This idea was further developed through the ecological footprint concept, with Wackernagel and Rees
(1998) estimating that more than 100 ha of ecologically productive land may be required to support every 1 ha of urban land.
Increasing recycling of nutrients and production of food within cities (Fig. 5) is one approach towards making cities more
sustainable. Even individual households have been conceptualized as ecosystems, as Fissore et al. (2011) quantified fluxes of
carbon, nitrogen, and phosphorus into and out of homes along an urban to rural gradient around Saint Paul, Minnesota. The
relevance of the ecosystem concept to human-designed systems is illustrated by the development of the field of industrial ecology,
which has a goal of modeling industrial systems after natural systems to turn waste products into resources.
While all ecosystems on Earth are characterized by the exchange of materials with the outside environment to some degree, the
design of long-term remote space habitats necessitates the recycling of all elements required for life. The Biosphere 2 experiment in
southern Arizona featured 8 humans sealed in a 12.5 ha materially closed ecosystem from 1991 to 1993. This experiment encountered
many challenges including pest management and producing adequate amounts of food, regulating atmospheric chemistry and water
nutrient levels, and very high extinction rates, leading ecologists to conclude that we do not yet know how to engineer systems that
replicate the life-support systems inherent in natural ecosystems (Cohen and Tilman, 1996; Silverstone and Nelson, 1996).

Fig. 5 This urban farm in Saint Paul, Minnesota, uses compost from municipal organics waste to recycle nutrients back into the human food system,
providing an example of cycling in urban ecosystems where flows are typically dominated by inputs and outputs. Food and materials are transported
into and out of the city via highways, railroads, and barges. The Mississippi River is a source of drinking water, and it receives treated wastewater. The
haze in the background of this photograph is smoke from wildfires in central Canada, representing another input from outside the ecosystem.
 The Ecosystem Concept 333

Conclusions

Early ecologists including Möbius, Forbes, and Tansley, recognized that organisms cannot be understood outside of the context of
the other species and abiotic environment with which they interact. Over the past century, the ecosystem concept has developed
through empirical and theoretical advances. The concept gained wide exposure through the textbook and other writings of E.P.
Odum, but different usages of the term have led to some ambiguity. The stock-and-flow conceptualization is useful in analyses of
the internal dynamics of an ecosystem, whereas the structural conceptualization is useful as a characterization of habitat type.
Importantly, both of these versions of the ecosystem concept allude to the importance of internal dynamics, and to a whole that is
greater than the sum of its parts. While the ecosystem concept was initially developed to describe natural systems, the concept has
demonstrated utility in human dominated systems, as seen in the development of the field of industrial ecology and through the
design of space habitats.

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Further Reading
Forbes, S. A. (1926) The lake as a microcosm. Bulletin Illinois Natural History Survey 15, 537–550.
Lindeman, R. L. (1942) The trophic-dynamic aspect of ecology. Ecology 23, 399–417.
Möbius, K. (1877) Wiegundt Die Auster und die Austernwirtschaft. Berlin: Hempel und Parey.
Odum, E. P. (1953) Fundamentals of ecology. W.B. Saunders Company.